reviewer acknowledgement koedoe 2011; 53(2)http://www.koedoe.co.za koedoe african protected area conservation and science recognises the value and importance of the peer reviewer in the overall publication process – not only in shaping the individual manuscript, but also in shaping the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely, productive peer review process. we would like to take this opportunity to thank all reviewers who participated in shaping this volume of koedoe: we appreciate the time taken to perform your review successfully. page 1 of 1 in an effort to facilitate the selection of appropriate peer reviewers for koedoe, we ask that you take a moment to update your electronic portfolio on www.koedoe.co.za for our files, allowing us better access to your areas of interest and expertise, in order to match reviewers with submitted manuscripts. if you would like to become a reviewer, please visit the journal website and register as a reviewer. to access your details on the website, you will need to follow these steps: 1. log into the online journal at http://www. koedoe.co.za 2. in your ‘user home’ [http://www.koedoe. co.za/index.php/koedoe/ user] select ‘edit my profile’ under the heading ‘my account’ and insert all relevant details, bio statement and reviewing interest. 3. it is good practice as reviewer to update your personal details regularly to ensure contact with you throughout your professional term as reviewer to koedoe. please do not hesitate to contact me if you require assistance in performing this task. margo martens submissions@koedoe.co.za tel: +27 (0)86 1000 381 fax: +27 (0)86 5004 974 koedoe african protected area conservation and science abraham nkhata alan anderson anna treydte belinda reyers bruce brockett charles breen dave richardson david edge ernita van wyk fred kruger gregory kiker jesse kalwij jon rodriguez kevin murray kevin rogers louis liebenberg marius claassen mark robertson mathieu rouget nick abel nick rivers-moore nyambe nyambe oonsie biggs patrick o’farrell pierre bonnet richard fynn richard stirzaker robert scholes steve mccool tim lynam tony swemmer wayne twine william bond filelist convert a pdf file! koedoe 19: 179-180 (l976) a case of abomasal impaction in a captive giraffe (giraffa camelopardalis) d. v. gradwell s tate veterinarian department of veterinary services skukuza jj50 introduction a male giraffe calf was brought into the experimental pens at skukuza after being found on its own in poor condition in the kruger national park, republic of south africa. the mother of the calf could not be traced. the giraffe was fed on full cream powdered milk and given dry lucerne, used to feed all other herbivores in the bomas. the lucerne was placed on a wooden board on top of a normal bovine feeding rack, approximately two metres above ground. the giraffe was also part of an experiment to determine the susceptibility of a number of species to heartwater and had been injected with positive heartwater blood 18 days before it died. the enclosure had an earth floor but the giraffe was penned at night in a stall with bedding on a concrete floor, as a protection against predators. shade consisted of a number of knob thorn (acacia nigrescens) trees in the enclosure. the leaves of these trees were soon consumed to the maximum reach of the giraffe. it was noticed that the animal then started grazing on grass which had sprouted after very good rains . the giraffe would lower its head to graze, after spreading and bending the forelegs in the same procedure as is normally followed for drinking. the tongue was used for grazing as when browsing. the grass was collected into the mouth and pulled upward by a jerk of the head. due to the damp soil the entire root systems of tussocks were uprooted. leaves, stems, roots and soil adherring to the roots were masticated and swallowed. no treatment was given to the giraffe when noticed to be ill. this coincided with the expected reaction to the artificial infection with heartwater. symptoms of anorexia and general apathy were at first presumed to be a reaction to the disease. no fever developed, and the animal died during the night after symptoms were first noticed. discussion post-mortem examination revealed slight ascites but no other typical heartwater lesions were observed. brain smears stained with giemsa revealed no organisms of cowdria ruminantium in endothelial cells. the 179 abomasal contents were extremely dry and consisted of a number of roughly circular five cm diameter bolus-like conglomerates of lucerne, gravel and sand. one such bolus was situated at the pylorus and completely blocked the lumen. this particular bolus did not break up after dropping onto a concrete slab from a height of approximately one metre. the rumen, reticulum and omasum were well filled with ingesta but the remainder of the in testinal canal, except for normal pellets in the final '15 cm of rectum, was empty. a diagnosis of abomasal impaction was made. conclusion cases of a similar nature are not unknown and grazing is fairly common in captive giraffe with insufficient green feed. in the case of giraffe in captivity or in quarantine camps before relocation, an adequate supply of suitable green foliage should be supplied . if this is not available, care should be taken to prevent fresh, highly palatable grass from sprouting in such pens which will produce an abnormal habit of grazing in the giraffe and a real danger of severe losses. surgical intervention, however, could be attempted after diagnosis but a knowledge of this danger will make preventive measures more effective. acknowledgements thanks are due to the director of veterinary services, department of agricultural technical services for permission to publish this article and the director of nature conservation, national parks board of trustees for criticising the manuscript. 180 page 1 page 2 book information book title: environmental risk management in south africa book cover author: mike mentis isbn: 9780620507523 publisher: publisher.co.za; 2010, r228.00* *book price at time of review book review information reviewer title: review of environmental risk management in south africa reviewer: harry biggs1 affiliation: 1sanparks, skukuza, south africa email: biggs@sanparks.org postal address: private bag x402, skukuza 1350, south africa how to cite this article: biggs, h., 2011, ‘review of environmental risk management in south africa’, koedoe 53(1), art. #1065, 1 page. doi:10.4102/koedoe.v53i1.1065 copyright notice: ©2011. the authors. licensee: aosis openjournals. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) review of environmental risk management in south africa congratulations to mike mentis on producing such a concise, clear and powerful guide and critique! i would recommend this book to any environmental manager or consultant who is seeking something beyond the largely once-off and routinised processes in use in south africa for integrated environmental management (iem) especially if they have an existing or potential interest in a more adaptive, risk-based approach. i would also hope that open-minded officials responsible for the legislation he criticises, would give the arguments put forward in the book a fair hearing. for active practitioners, the book is useful because it uses ‘risk’ as the central notion to improve on what was probably all along intended by the word ‘impact’. this is convincingly justified in the course of the book. many practitioners will have heard of, or even used some components of risk-based techniques. however, i believe that the steps and techniques that he clearly lays out, provide the necessary bridge from a more conventional eia to a much improved eia with a central emphasis on adaptation and risk analysis. these approaches can also be used in wider environmental contexts. in fact i wondered whether we are not eventually getting to the point that so-called ‘ecosystem management’ will at last, in actual practice, be seen in a stronger continuum with ‘environmental management’, the latter usually understood in the sense of dealing with consequences of ‘developments’. i always found it curious that ‘ecosystem management’ textbooks (and now slowly, ecosystem management practice, at least in some settings) shows far more uptake of the adaptive ethos. regarding the latter, he makes the point over and over again (rising to a crescendo in the last chapter) that the law’s prescriptions, although not excluding this, do very little to encourage a learning approach through iterations – the vast majority of eias and environmental management plans produce close to static blueprints. he likens this, amongst other analogies, to a general heading out to war with a full and irrevocable plan written out of his units’ movements, intentions and assumed outcomes for the next several years! a semi-independent chapter on decision-making deals with several complementary ideas which add further value to the whole risk theme, but which, again, can be used in many other settings too. these useful principles are not restricted to iem, though most of his examples are. regarding the author’s critique of the existing de facto practice of iem in south africa as an outflow of the law, the author builds a solid interconnected argument (which i won’t spoil for the reader) that a serious re-think is necessary and overdue. this includes the paranoia of omission – the underlying fear of not covering every possible risk in an eia. if the risks are sensibly prioritised by analysis (one subject of the book) and action restricted, at least initially, to those that have the best chance of delivering, then achieving clearly set environmental goals actually becomes possible. what really locks us into the ‘once-off blueprint mode’ is the fact that the environmental departments of government, who in good but misguided faith want to control everything centrally, cannot even deal with the ‘once-offs’, let alone the updates which are actually essential to make the system work properly. he suggests several possible changes to the way the country deals with this, including placing the onus on developers to actually buy into and defend their reputations based on the real underlying fundamentals of the constitution (the right to an environment not harmful to health, free from pollution and degradation) rather than seeing that they as quickly or easily as possible pass the ‘tick-box’ test of the current eia requirements. the ultimate changes he suggests need to be radical, but he suggests these could be phased in for workability. except in a few very small ways, the 2010 regulation changes (which appeared just after his book came out) under the south african national environmental management act, do not move in the directions he proposes. so all those with an interest in robust debate on this topic – this book is also for you! there are a few minor glitches (some semi-estranged footnotes and some typos/miscalculations in a key decision-tree graphic) but nothing serious enough to do much more than keep the reader on her or his feet. many messages are repeated over and over again, and the final chapter reads a bit like a sermon. but in my opinion he is close to spot on, and i hope we see practitioners using more of this approach, and ultimately that we see a revamp in south africa (he does not discuss whether other countries are any better – i fear not – and in which case it is indeed another chance for south africa to show some leadership). readers who don’t agree with him will grow by at least facing the suggestions and challenges articulated in this highly readable book with its clear examples. my suggestion is to either consider adopting this line of thinking, or to produce something better, but not to let it leave one cold. reviewer acknowledgement http://www.koedoe.co.za open access koedoe recognises and acknowledges the value and importance of reviewers in the publication process – not only in shaping the individual manuscript, but also in upholding the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely, productive peer review process. we would like to take this opportunity to thank all reviewers who participated in shaping this volume of koedoe: we appreciate the time taken to perform your review. page 1 of 1 in an effort to facilitate the selection of appropriate peer reviewers for koedoe, we ask that you take a moment to update your electronic portfolio at www.koedoe.co.za for our records. this will allow us better access to your areas of interest and expertise, in order to match reviewers with submitted manuscripts. if you would like to become a reviewer, please visit the journal website and register as a reviewer. to access your details on the website, you will need to follow these steps: 1. log into the online journal at http://www. koedoe.co.za 2. in your ‘user home’ [http://www.koedoe. co.za/index.php/koedoe/ user] select ‘edit my profile’ under the heading ‘my account’ and insert all relevant details, bio statement and reviewing interest. 3. it is good practice as a reviewer to update your personal details regularly, to ensure contact with you throughout your professional term as reviewer to koedoe. please do not hesitate to contact me if you require assistance in performing this task. margo martens submissions@koedoe.co.za tel: +27 (0)21 975 2602 fax: +27 (0)21 975 4635 koedoe african protected area conservation and science adrian craig armin seydack bill carter brian van wilgen charlie m. shackleton candice janse van rensburg carl anhaeusser corli coetsee cornelis van der waal daniele colombaroli dave balfour dave druce dave thompson david le maitre duan biggs emma a. van garderen frances siebert georgette lagendijk graeme shannon gretel van rooyen herbert prins hugo bezuidenhout ian whyte izak smit james pryke jay barton joop schaminée joris cromsigt kate meares lauren coad laurence watson leif peterson leon lotz leslie brown lindsey gilson louise swemmer lubomír tichý lynne isbell maartin strauss mago maila markus gusset marna herbst matthys dippenaar melodie mcgeoch mervyn lötter michael panagos moses cho neil crouch neville pillay niels dreber peter scogings pieter bester piotr migon rachel mostert rina grant rob slotow samson chimphango sharon thompson simon power stefan foord stefan grab theo mostert tony rebelo tony swemmer ute schmiedel vaughan spaull vhalinavho khavhagali wendy annecke 172 filelist convert a pdf file! koedoe 1 9: 169-17 0 (1976) the black-necked grebe podiceps nigricollis a new bird record for the kruger national park a. c. van bruggen department oj systematic zoology and evolutionary biology c/o ri;ksmuseum van natuurlijke historie leiden the netherlands the avifauna of the kruger national park (knp) was described in detail by kemp (1974). however, species new to the list are bound to turn up occasionally. joubert and english (1973), who discovered the occurrence of the crimson-breasted shrike laniarius atroccoccineus burchell, write: "many of these species include migrants and vagrants entering the park during rather abnormal times such as cyclones or periods of heavy rainfall." after having listed the scaly-feathered finch sporopipes squamifrons (a. smith), for the kruger national park list some 15 years ago (van bruggen 1960), this note deals with an additional new record i.e. the black-necked grebe podiceps nigricollis brehm. on 1975.04.16 a specimen in winter plumage was observed by the author, for some time desporting itself on kumana dam alongside the main road h 1-3 between tshokwane and satara. notwithstanding rain and drizzle the bird was clearly recognizable and identification was confirmed by observation through binoculars 10 x 50. mclachlan and liversidge (1957) show this grebe to be a vagrant occurring as far east as the eastern transvaal, so that the knp record naturally fits into this pattern of distribution. usually this bird is to be found on the coast and only occasionally does it range far inland. this also applies to east africa. in 1974 we observed a specimen in winter plumage in kenya on the dam at keekorok lodge in the masai mara game reserve on 29 and 31 january; this is much further inland in a decidedly arid area. the knp record may be correlated with the unusually wet summer of 1974/7 5 in southern africa. in this part of africa the black-necked grebe is both a resident breeder and a migrant from the north. odd specimens are probably likely to turn up at irregular intervals in the eastern transvaal. the knp obviously has seemingly suitable stretches of open water to temporarily accommodate the black-necked grebe. references joubert, s. c. j. and m. english. 1973. a new bird record for the kruger national park. koedoe 16: 199-200. 169 kem p, a. c. 1974. the distribution and status of the birds of the kruger national park. koedoemonograph 2:1-352. mclachlan, g. r. and r. liversidge. 1957. roberts birds of south africa. 2nd ed. cape town: trustees of the s.a. bird book fund. van bruggen, a. c. 1960. notes and observations on birds in the transvaal, southern rhodesia, and portuguese east africa. ostrich 31 :30-31. 170 page 1 page 2 article information author: robert j. scholes1 affiliation: 1global change and sustainability research institute, university of the witwatersrand, south africa correspondence to: robert scholes email: bob.scholes@wits.ac.za postal address: private bag x3, wits 2050, south africa how to cite this article: scholes, r.j., 2015, ‘bigpicture ecology for a small planet’, koedoe 57(1), art. #1328, 4 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1328 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. big-picture ecology for a small planet in this commentary... open access • abstract • the multiple scales of ecosystem change and response • large national parks and their surrounds are a useful platform for research • datasets of great value • contributions to global and local knowledge    • the global carbon balance    • climate change    • biodiversity loss    • land cover change • implications of big-scale process for management, policy and research priorities • acknowledgements    • competing interests • references abstract top ↑ for a number of years, the extensive ecosystems of southern africa have been a testing ground for ideas and techniques useful for studying and managing large-scale complex systems everywhere, and in particular for tackling issues of global change. the first contribution has been through making consistent, long-term, large-scale observations on climate, vegetation and animal dynamics and disturbances. these have been crucial in developing and testing hypotheses regarding how the earth system works at large space and timescales. the observational techniques have evolved dramatically over time: from notes kept by individuals, to systematic measurement programmes by organisations, to continuous and sophisticated measurements made by automated systems such as satellites and flux towers. the second contribution has been experimental, developing the notion that ecosystems can be the subject of deliberate experimental manipulation. sometimes this has taken the form of large-scale treatments, such as fire trials or herbivore exclusion plots. more frequently, it has made use of the ‘experiment’ of the protected area in contrast to its surrounds, or has exploited the information in natural or human-induced gradients. ecosystem experimentation has required rethinking the fundamentals of experimental design: what is the experimental unit? what is the meaning of a control? what constitutes replication? the third contribution has been theoretical. how does the functioning of warm, dry, species-rich ecosystems differ from the cool, moist, species-poor ecosystem examples that dominate the literature? what are the roles of disturbance and competition is maintaining ecosystem diversity, and top-down versus bottom-up control in maintaining ecosystem structure? the fourth contribution concerns the management of large-scale complex systems in the face of limited knowledge. how can the gap between science and policy be narrowed? what advantages and challenges does participatory co-management offer? how do you implement adaptive management? the multiple scales of ecosystem change and response top ↑ ecosystems and social systems are hardly ever static: they change over time, sometimes slowly and sometimes abruptly, and in response to both internal dynamics and external drivers. these changes play out over the surface of the earth in the form of observable ‘patterns’, at a range of scales, from that of individual organisms, to patches showing similar attributes, to whole ecosystems, and to the entire biosphere. there is a broad tendency (which is by no means a fixed rule) that slow processes are associated with large spatial scales, and fast processes with small scales (vance & doel 2010). for useful information to be gained, it is necessary to match the scales of observation, in both time and space, with the scales of what is being observed (englund & cooper 2003). furthermore, it is plausible that interventions in ecosystem processes are more effective if they are matched to the scale of the phenomenon being addressed. it is therefore an area of concern that the majority of the data collected relating to the state and functioning of the natural and human world is on small scales and over short periods of time (carpenter 1996; englund & cooper 2003; levin 1992). this is partly because information at large scales and over long periods was not part of our human experience until relatively recently: we observed locally, and usually for no longer than a human lifespan (and often much less). written records in archives, travel, telecommunications and space exploration have changed all that. at the same time, human dominance of the world has led to a range of phenomena on regional and global scale – such as climate change, biodiversity loss and ecosystem degradation – which require investigation and intervention on scales far beyond the local and immediate. there has been a trend in science towards finer and finer resolution, more detail, increased specialisation and narrower disciplines. the scientific approach of reductionism – looking for the causes of things by isolating them and delving down into their underlying elements – has been enormously successful in many fields; however, it has been unhelpful in solving issues which are systemic in nature. system problems emerge from interactions and context, and require a wider rather than a narrower view; a view which is often associated with larger scales in space and time. fortunately, theoretical and technological advances the past few decades have made it increasingly feasible to observe, experiment with and manage systems on the scales at which they actually operate (schindler 1998). there is a growing realisation that the phenomena relevant to environmental management operate on a range of scales, rather than on a single scale. even within a class of issues (‘desertification’, for instance) several nested scales are often relevant. thus there is seldom a single ‘best’ scale for all purposes. the trend is towards multi-scale observation and analysis, and in particular towards trying to understand how phenomena propagate across scales and interact between scales (scholes et al. 2013). in classical science, still reflected in science education and attitudes towards data ownership, the researcher made his or her own observations. this is not technically feasible on large scales and over long times; we depend on the accumulation of data from many individuals, the custodianship of institutions and complex and expensive equipment, such as satellites, which are designed and operated by specialists. as a result, the activity of making accurate, repeated observations for use by other scientists is a legitimate aspect of science in its own right, which must be funded and recognised as such. large national parks and their surrounds are a useful platform for research top ↑ the act which governs south african protected areas (national environment management: protected areas act [act no. 57 of 2003]) states that the purpose of national parks, amongst other things, is to ‘provide spiritual, scientific, educational, recreational and tourism opportunities which are environmentally compatible’. therefore, conducting scientific activities within parks should not be at the whim of individual park managers: it is part of their mandate, provided it is not in conflict with the broad objective of environmental protection. nor should such research be selected solely on the basis of its immediate usefulness for park management. the scale, relative absence of human impact, institutional stability and infrastructure of national parks make them uniquely appropriate for certain types of research. research activities, properly designed and communicated, seldom conflict materially with biodiversity management and tourism activities: generally, visitors are interested in the research and find that it adds value to their experience. visits by scientists for purposes of research are a significant source of income to the park system. datasets of great value top ↑ national parks have accumulated data of value to both themselves and the wider community. kruger national park (knp) is exemplary here, because of its history and size. in particular, the long-term records pertaining to weather, vegetation composition and state, the populations of mammals and birds and areas affected by fire are near-unique. the philosophical shift from keeping such data for internal use only to making it openly and freely available which occurred in knp over the past two decades has resulted in a massive increase in scientific publications relating to the park. it is recognised that collecting and servicing such datasets involves a significant cost to the park system. experience in many organisations has shown that attempting to recover this cost directly from users is counter-productive: data becomes unaffordable to the users and the expense of invoicing exceeds the income stream. a more sensible approach economically (i.e. considering the needs of society as a whole, rather than the finances of the accounting unit alone) is to treat the cost of data collection as an operating expense to be centrally borne (perhaps supported by specific external grants) and then make the information freely available in order to maximise its use. use of such data by third parties carries obligations: the data source must be correctly acknowledged; publications arising from its use must be lodged with the data collectors for reporting purposes; and data collected in or on the park must in turn become publically and freely available. contributions to global and local knowledge top ↑ there are many domains in which research in large protected areas in south africa has contributed to global scientific understanding; the following are a few examples amongst many. the global carbon balance climate change can be thought of as a symptom of disruption to the global carbon metabolism. the emission of fossil fuel–derived carbon dioxide has perturbed a global carbon cycle previously in near-equilibrium, resulting in a rise in atmospheric concentrations and global mean temperature. about half of the emitted carbon dioxide is removed from the atmosphere by marine and terrestrial ecosystems, approximately equally. the uptake by terrestrial ecosystems is globally distributed, and because of their large extent and relatively high productivity, a substantial fraction is probably being stored in savannas such as those covering large parts of africa. the flux towers operating in the knp since 2000 help to illuminate this issue. they show that this savanna landscape is alternately a source and sink for carbon, depending on the climate of particular years (archibald et al. 2009; kutsch et al. 2008; williams, hanan & scholes 2009). at the decadal timescale, the two landscapes monitored in the knp are most likely small carbon sources rather than sinks. climate change temperature and rainfall records began at skukuza in 1912, and have continued daily since then, almost uninterrupted (kruger, makamo & shongwe 2002). records at many other camps and ranger stations are similarly long and good. there are very few records of this duration in africa, and they are invaluable in proving the upward trend in global mean (and local) temperature over the past century. the work on greenhouse gas emissions from savanna fires in knp has been seminal (swap et al. 2003).the studies relating natural climate variability to plant and animal responses (archibald & scholes 2007; eckhardt, wilgen & biggs 2000; wessels et al. 2006) as well as the experimental manipulations of fire (biggs et al. 2003) and water supply (february et al. 2013) have provided insights into the possible ecological impacts of future climate change. biodiversity loss national parks exist to protect biodiversity. especially when large and well managed, they act as reference points for the potential state of biodiversity. even so, they are not immune from loss of species themselves – or the gain of species through invasions or migration (chirima, owen-smith & erasmus 2012; thrash 1998). because of the research focus on biodiversity dynamics in national parks, they are often key sources of information for tracking changes and for unscrambling global effects, such as climate change, from local effects, such as changes in habitat and the effects of increases and decreases in key populations (trollope et al. 1998; whyte 2004). land cover change the contrast in land use across national park boundaries is often clearly apparent as a difference in land cover. this sets up an ‘experiment’ by which the consequences for climate, ecosystem services and biodiversity can be explored. the effect of differing fire regimes within the knp, and between it and neighbouring lands, has been especially illuminating (van wilgen 2009; van wilgen et al. 2004). implications of big-scale process for management, policy and research priorities top ↑ much of the day-to-day management in protected areas relates to symptomatic issues, immediate concerns and small scales. this bias affects the information needs as perceived by conservation managers; for example, they routinely prioritise issues relating to population excesses of ‘problem species’, population viability in rare species, localised degradation and specific instances of conflict with neighbours. the slower, larger-scale processes, which are more likely to threaten the ability to satisfy the mandate of protected area management, and which may be the root cause of many of the crises which dominate management attention, often go unrecorded and unrecognised until too late – land use changes in adjacent areas, climate and atmosphere changes, and economic and political shifts at regional to global scale are some examples. the knp has been a leader in the practical implementation of adaptive management, the style suggested as most appropriate for ecosystem management in the presence of limited knowledge (van wilgen & biggs 2011). acknowledgements top ↑ competing interests the author declares that he has no financial or personal relationships which may have inappropriately influenced him in writing this article. references top ↑ archibald, s., kirton, a., van der merwe, m., scholes, r.j., williams, c.a. & hanan n. 2009, ‘drivers of interannual variability in net ecosystem exchange in a semi-arid savanna ecosystem, south africa’, biogeosciences 6, 261–266. http://dx.doi.org/10.5194/bg-6-251-2009 archibald, s. & scholes, r.j., 2007, ‘leaf green-up in a semi-arid african savanna – separating tree and grass responses to environmental cues’, journal of vegetation science 18, 583–594. biggs, r., biggs, h.c., dunne, t.t., govender, n. & potgieter a.l.f., 2003, ‘experimental burn plot trial in the kruger national park: history, experimental design and suggestions for data analysis’, koedoe 46(1), 1–15. http://dx.doi.org/10.4102/koedoe.v46i1.35 carpenter, s.r., 1996, ‘microcosm experiments have limited relevance for community and ecosystem ecology’, ecology, 77(3), 677–680. http://dx.doi.org/10.2307/2265490 chirima, g., owen-smith, n. & erasmus, b., 2012, ‘changing distributions of larger ungulates in the kruger national park from ecological aerial survey data’, koedoe 54 art. #1009, 11 pages, http://dx.doi.org/10.4102/ eckhardt, h.c., wilgen, b.w. & biggs, h.c., 2000, ‘trends in woody vegetation cover in the kruger national park, south africa, between 1940 and 1998’, african journal of ecology 38(2), 108–115. http://dx.doi.org/10.1046/j.1365-2028.2000.00217.x englund, g. & cooper, s.d., 2003, ‘scale effects and extrapolation in ecological experiments’, advances in ecological research, 33, 161–213. http://dx.doi.org/10.1016/s0065-2504(03)33011-9 february, e.c., higgins, s.i., bond, w.j. & swemmer, l., 2013, ‘influence of competition and rainfall manipulation on the growth responses of savanna trees and grasses’, ecology 94(5), 1155–1164. http://dx.doi.org/10.1890/12-0540.1 kruger, a.c., makamo, l.b. & shongwe, s., 2002, ‘an analysis of skukuza climate data’, koedoe 45(1), 1–7. http://dx.doi.org/10.4102/koedoe.v45i1.16 kutsch, w.l., hanan, n., scholes, r.j., mchugh, i., kubheka, w., eckhardt, h. & williams, c., 2008, ‘response of carbon fluxes to water relations in a savanna ecosystem in south africa’, biogeosciences 5, 1797–1808. http://dx.doi.org/10.5194/bg-5-1797-2008 levin, s.a., 1992, ‘the problem of pattern and scale in ecology’, ecology 73, 1943–1967. http://dx.doi.org/10.2307/1941447 schindler, d.w., 1998, ‘whole-ecosystem experiments: replication versus realism – the need for ecosystem-scale experiments’, ecosystems 1(4), 323–334. http://dx.doi.org/10.1007/s100219900026 scholes, r.j., reyers, b., biggs, r., spierenburg, m.j. & duriappah, a., 2013, ‘multi-scale and cross-scale assessments of social–ecological systems and their ecosystem services’, current opinion in environmental sustainability 5, 1–10. http://dx.doi.org/10.1016/j.cosust.2013.01.004 swap, r.j., annegarn, h.j., suttles, j.t., king, m.d., platnick, s., privette, j.l. & scholes, r.j., 2003, ‘africa burning: a thematic analysis of the southern african regional science initiative (safari 2000)’, journal of geophysical research: atmospheres 108(d13), 8465. http://dx.doi.org/10.1029/2003jd003747 thrash, i., 1998, ‘impact of water provision on herbaceous vegetation in kruger national park, south africa’, journal of arid environments 38(3), 437–450. http://dx.doi.org/10.1006/jare.1997.0318 trollope, w.s.w., trollope, l.a., biggs, h.c., pienaar, d. & potgieter, a.l.f., 1998, ‘long-term changes in the woody vegetation of the kruger national park, with special reference to the effects of elephants and fire’, koedoe 41(2), 103–112. http://dx.doi.org/10.4102/koedoe.v41i2.255 vance, t. & doel, r., 2010, ‘graphical methods and cold war scientific practice: the stommel diagram's intriguing journey from the physical to the biological environmental sciences’, historical studies in the natural sciences 40(1), 1–47. http://dx.doi.org/10.1525/hsns.2010.40.1.1 van wilgen, b.w., 2009, ‘the evolution of fire management practices in savanna protected areas in south africa’, south african journal of science 105(9–10), 343–349. van wilgen, b.w. & biggs, h.c., 2011, ‘a critical assessment of adaptive ecosystem management in a large savanna protected area in south africa’, biological conservation 144(4), 1179–1187. http://dx.doi.org/10.1016/j.biocon.2010.05.006 van wilgen, b.w., govender, n., biggs, h.c., ntsala, d. & funda, x.n., 2004, ‘response of savanna fire regimes to changing fire-management policies in a large african national park’, conservation biology 18(6), 1533–1540. http://dx.doi.org/10.1111/j.1523-1739.2004.00362.x wessels, k.j., prince, s.d., zambatis, n., macfadyen, s., frost, p.e. & van zyl, d., 2006, ‘relationship between herbaceous biomass and 1-km2 advanced very high resolution radiometer (avhrr) ndvi in kruger national park, south africa’, international journal of remote sensing 27(5), 951–973. http://dx.doi.org/10.1080/01431160500169098 whyte, i.j., 2004, ‘ecological basis of the new elephant management policy for kruger national park and expected outcomes’, pachyderm 36, 99–108. williams, c.j, hanan, n. & scholes, r.j., 2009, ‘complexity in water and carbon dioxide fluxes following rain pulses in an african savanna’, oecologia 161, 469–480. http://dx.doi.org/10.1007/s00442-009-1405-y book information title: the limits to scarcity: contesting the politics of allocation book cover: book cover the limits to scarcity: contesting the politics of allocation editor: lyla mehta isbn: 978-84407-542-3 publisher: earthscan: london and washington dc; 2010, r400* *book price at time of review review title: scarcity in the time of over-consumption reviewer: wendy annecke1affiliation: 1cape research centre, south african national parks, south africa postal address: po box 216, steenberg 7947, cape town, south africa how to cite this article: annecke, w., 2012, ‘scarcity in the time of over-consumption’, koedoe 54(1), art. #1089, 2 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1089 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. scarcity in the time of over-consumption in this book review... open access resource use and the sustainability of resources constitute a significant concern for many scientists and field staff in south african national parks (sanparks). according to its mission ‘to develop, manage and promote a system of national parks that represents biodiversity and heritage assets by applying best practice, environmental justice, benefit sharing and sustainable use’, sanparks is committed to sharing the benefits of the areas it protects with the wider society. this wider society, in immediate terms, primarily incorporates tourists and the communities living on park borders. tourists require access to resources such as roads, water, energy, food and waste facilities, whereas people living on the boundaries of the parks require access to grazing, medicinal and food plants, wood fuel and bush-meat. consequently, some of the biodiversity resources that sanparks was established to protect, are under severe pressure from ‘the wider society’ in general. for example many medicinal plants are threatened by over harvesting (both legal and illegal), whilst other resources, such as water, are threatened by pollution, changes in land use and increased demand for these. this book will be of interest to conservationists who are concerned with the ethics and sustainability of resource use, as well as the status of biodiversity health in the parks. the limits to scarcity does not provide answers to the difficult issues arising with regard to resource use, but it does provide a different way of thinking about scarcity and why this occurs. in doing so it raises questions about who the primary resource users are, and encourages a shift in perspective about the rights of all users. the limits to scarcity is compiled from of a series of papers delivered at a conference on ‘scarcity and the politics of allocation’ held at the institute of development studies, at the university of sussex, in june 2005. the fourteen chapters are divided into three parts with each part preceded by an introduction and commentary by the editor, leyla mehta. the guidance she provides is useful for novice readers in the field of scarcity or for readers with limited time. each chapter explores the notion of scarcity from a different angle and includes how the notion of scarcity is constructed and whose purposes it serves. the three chapters in part i discuss why scarcity matters, part ii explores the different perspectives of scarcity within economics and part iii investigates the politics of scarcity in case studies of water, energy and food supplies. authors include well known nicholas xenos, fred luks and ben fine. the latter is better known to most south africans for his and zavareh rustomjee’s seminal work on the minerals-energy-complex. the limits to scarcity makes an important contribution to resource use debates, particularly in the light of the rampant consumer culture which is seen by most people as the only way out of the on-going global economic woes. chapter by chapter the book unpacks the dynamics of how resources are allocated in various societies: who has access to what resources, and under what circumstances. the authors are careful to acknowledge the ‘biophysical realities of falling groundwater levels, melting icecaps and declining soil fertility’ (mehta 6) whilst they dissect the politics of need and those involved in the access and allocation of resources. the book has chapters of interest to economists (such as that by fine), engineers (such as that by lankford) and to philosophers and non-economists (xenos). for those interested in social justice and equitable societies as well as those involved in development and the poverty industry, there is plenty of significant content over which to ponder. what is most disturbing about this book is the evidence demonstrating how all-pervasive neo-malthusian economics has become, even in ‘development thinking’. in the first chapter, mehta provides a historical overview about the legacy of and approaches to normalizing the ‘scarcity postulate’. this overview sets the tone for the rest of the book by demonstrating that scarcity is not necessarily a natural or universal concept. she offers the chapters that follow as evidence for her argument as well as her challenge to the inertia evident in much mainstream thinking and economics. the ensuing chapters provide ample evidence that there is enough water, energy and food for everyone in the world (fao 2000 in hilyard 2010:152). one has only to visit the landfill sites and the dump pickers industry in south africa to observe the magnitude of waste generated by society, but, as raynor (p. xviii) argues, scarcity has become a rationale for the inequitable allocation of resources: it constitutes a gate-keeping mechanism. fine (p. 82) follows this up by asking uncomfortable questions about the manner in which a society chooses to rank one person’s welfare over another; their criteria for selecting one group of users who are able to access resources easily and use them intensely, whilst another group may have only minimal access to the same resources. i would suggest that feminists have become adept at conducting analyses that evaluate access to, control over and benefits from resources, and that feminist literature could contribute to this debate and perhaps shift its dynamics. in dipak gyawali and ajaya dixit’s chapter 13, scarcity is highlighted as a modern concept driven by technological choices to meet specific socio-political needs (p. 234). the authors quote helena norberg-hodge who worked in laddakh when there was no word for ‘poor’ in that region, but on returning some ten years later, they found her former translator pleading poverty and that his country desired a western notion of development. he and his countrymen (and women) had fallen victim to the notion of wealth that emanates from material goods. there are chapters on soil fertility in africa, agriculture and hunger, and two chapters for those involved in the water sector and the shared responses to its scarcity. the editor, mehta, has herself worked extensively on issues around water scarcity and is well known in this field. bruce lankford argues, in chapter 11, that water scarcity should not only be understood as ‘volumetric imbalance to be dealt with by saving, storing and delivering more water’ but suggests a framework for supply, demand and shared responses. he highlights how the natural and social scarcity of water can be exacerbated by people and institutions that are not designed to react to the rapidly occurring variations in water scarcity, in time and space, and provides a table comparing approaches for managing the dynamic supply of water in sub-catchment areas. from this he draws a number of conclusions, that provide persuasive arguments for the adoption of his composite and complex framework. in chapter 12, jasveen jairath re-examines the meaning and explanations of water scarcity and critiques what he calls the ‘conceptual weakness and politics’ of integrated water resource management. he highlights the importance of historical conditions, suggests the kind of questions that should be asked when bringing the weak and the strong together at a table, and points out that the democratic governance of water is seldom achieved. whilst the concept of scarcity itself is thoroughly unpacked, the book is weak on recommendations about how mainstream and pervasive thinking might be changed from valuing over-consumption to valuing sufficiency. xenos argues that historically the time for appealing to the morals and ethics of a society appears to be over. such appeals did not, for example, win the friends and support that president carter had hoped for in 1979. likewise, in south africa, appealing to the adherence to ex-president mandela’s values has worn thin. the question is what could provide the impetus for change? whilst none of the authors define the term ‘enough’, some point to individuals and communities that have described it for themselves: the amish and po chi-i are mentioned in this regard. i particularly like the words of po chi-i, the retired mandarin who wrote poetry, from about 850 ad, (in thompson 2010): what i shall need are a very few things. a single rug to warm me through the winter; one meal to last me the whole day. it does not matter that my house is rather small; one cannot sleep in more than one room! it does not matter that i have not many horses; one cannot ride two horses at once! (p. 131) there are typing errors throughout the book including the sub-title of a book in the science in society series: participation and exalusion in nuclear decision-making. mehta herself thanks the science and society programme – this name is an unlikely change from the series name and there are various other errors that should have been noticed and corrected. nonetheless, these issues should not detract from a provocative and worthwhile book. abstract introduction methods results discussion acknowledgements references about the author(s) david a. youldon african lion & environmental research trust, livingstone, zambia school of biomedical sciences, edinburgh university, united kingdom jackie abell african lion & environmental research trust, livingstone, zambia faculty of health and life sciences, coventry university, united kingdom joanne s. briffitt african lion & environmental research trust, livingstone, zambia lackson chama department of zoology and aquatic sciences, copperbelt university, zambia michaela d. channings african lion & environmental research trust, livingstone, zambia anastasia kilundo department of national parks and wildlife, mosi-oa-tunya area management unit, zambia christine k. larsen african lion & environmental research trust, livingstone, zambia dabwiso sakala african lion & environmental research trust, livingstone, zambia bruce a. schulte department of biology, western kentucky university, united states citation youldon, d.a., abell, j., briffitt, j.s., chama, l., channings, m.d., kilundo, a. et al., 2017, ‘patch-occupancy survey of elephant (loxodonta africana) surrounding livingstone, zambia’, koedoe 59(1), a1372. https://doi.org/10.4102/koedoe.v59i1.1372 short communication patch-occupancy survey of elephant (loxodonta africana) surrounding livingstone, zambia david a. youldon, jackie abell, joanne s. briffitt, lackson chama, michaela d. channings, anastasia kilundo, christine k. larsen, dabwiso sakala, bruce a. schulte received: 27 nov. 2015; accepted: 31 oct. 2016; published: 30 jan. 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract wild elephants represent the biggest human–wildlife conflict issue in livingstone, zambia. however, little is known about their movements. this survey investigated elephants’ habitat use outside a core protected and fenced zone that forms part of mosi-oa-tunya national park, zambia. using ‘patch-occupancy’ methodology, indications of elephant presence (feeding behaviour, dung and tracks) were surveyed. the survey aimed to assist proposed future monitoring exercises by defining the geographical extent that should be considered to improve accuracy in species abundance estimates. results were supplemented using collected indications of elephant presence from prior monitoring exercises, and during this survey. elephant presence was confirmed up to 8 km from the boundary of the protected core habitat, focussed in: (1) an unfenced zone of the national park, (2) along a road leading from the national park to the dambwa forest to the north and (3) along two rivers located to the west (sinde river) and east (maramba river) of the core area. detection probability of elephant presence was high using these methods, and we recommend regular sampling to determine changes in habitat use by elephants, as humans continue to modify land-use patterns. conservation implications: identification of elephant ranging behaviour up to 8 km outside of the mosi-oa-tunya national park in southern zambia will assist in managing human–elephant conflict in the area, as well as in assessing this seasonal population’s abundance. introduction in fragmented land-use mosaics, the home ranges of african elephants (loxodonta africana) cover a combination of protected and unprotected areas (douglas-hamilton, krink & vollrath 2005; graham et al. 2007; karidozo & osborn 2015). the ranging by elephants in human-dominated landscapes inevitably leads to interaction, and therefore conflict, with communities. most notably, conflict occurs with farmers as a result of crop-raiding, but also includes destroying water supplies, demolishing grain stores and houses, and sometimes injuring and killing people and livestock (dublin & hoare 2004; graham et al. 2007; sitati et al. 2003). wild elephants represent the biggest human–wildlife conflict issue in livingstone, zambia. however, little is known about their movements. understanding elephants’ use of land outside of protected areas is considered important to the future conservation and management of african elephant populations (graham et al. 2007; hoare 2000; okello et al. 2015). the mosi-oa-tunya national park (mot np) on the outskirts of livingstone in southern zambia supports a seasonal population of elephants, peaking in the months of may to october (african lion & environmental research trust [alert] 2012). a fenced section of the mot np, known as the ‘old zoological park’ (ozp), appears to be the core habitat used by elephants in the area. however, at least some elephants are also known to utilise the protected dambwa local forest no. 22, to the north of the mot np, as well as entering surrounding unprotected, human-dominated habitats (alert 2012), causing significant conflict with communities (chishika 2010). no robust survey has been performed to date to assess habitat use by elephants outside the mot np. further, the scope of long-term population monitoring in the region is difficult to define without knowledge of the species’ ranging behaviour. elephants are known to move out of the ozp only after dark (zambia department of national parks and wildlife, pers. comm., n.d.), whilst much of the land outside of the ozp is heavily forested. as such, direct observation of elephants from the ground or air, to assess ranging behaviour, in this area is problematic. ‘patch-occupancy’ or ‘presence–absence’ methodology is well established and provides an effective technique to monitor elephants’ ranging behaviour over large geographic areas (buij et al. 2007; mackenzie et al. 2002, 2003). this survey therefore aims to use the patch-occupancy method for elephants outside of the core area of the ozp to assist planning of long-term monitoring efforts in the region. aerial surveys conducted between september and december 2008 recorded an elephant population for mot np of 31 individuals (simukonda 2009). however, monitoring of elephants both within the ozp and in the surrounding areas between 2010 and 2012 identified 409 individuals that were either resident or transient at some point during the survey period (alert 2012), suggesting a more robust survey of the elephant population is needed. at present, elephants’ ranging behaviour outside of the ozp is unconfirmed. additionally, whether the ozp forms the core habitat for all elephants in the region, is unknown. for more accurate estimates of species abundance and density in the region, it is important to include areas outside the ozp in a robust survey design. however, the necessary geographic extent of any survey for improved survey accuracy, whilst also remaining cost-efficient, can currently only be postulated. results from this survey aim to address knowledge gaps in the ranging behaviour of elephants in livingstone to assist future population and conflict management in this area. methods located in the southern province of zambia, outside the town of livingstone, is the mot np that lies between 17° 48.897’–17° 58.300’ s and 25° 45.040’–25° 53.490’ e. it covers an area of approximately 6600 ha, with a fenced zone in the western section of approximately 2990 ha, known as the ‘old zoological park’ (ozp). the dambwa local forest no. 22 lies between 17° 39.962’–17° 49.300’ s and 25° 46.122’–25° 52.798’ e. it covers an area of 13 746 ha and lies to the north of, and shares a 4-km border with the ozp. these protected areas are surrounded by livingstone town, several villages, small scale and commercial farms, and communal lands. livingstone itself lies to the northeast of the mot np and has an estimated population of ca. 135 000 residents. the population surrounding the dambwa forest was estimated to be over 1680 people in 2005, residing in 11 villages. the principal livelihood system is subsistence agriculture with the main crops being maize, cassava, groundnuts, beans, sorghum, pumpkins and sweet potatoes. some livestock are also kept, including cattle, goats, pigs and chickens (ensvol consult 2007). the survey was undertaken during august to october 2015, coinciding with maximum seasonal elephant presence within the region (alert 2012). concentric distance bands of 4-km width were defined emanating from the ozp boundary (figure 1). a grid of 2 km × 2 km blocks (buij et al. 2007) was overlain on the distance bands. each block was assigned to a distance band based on within which band most of the block’s area fell. thirty nine blocks to be surveyed were randomly selected based on 20% block sampling intensity within each distance band, giving five sample blocks out of 24 in the 4-km distance band, eight sample blocks out of 40 in the 8-km distance band, 12 blocks out of 60 blocks in the 12-km distance band and 14 blocks out of 72 in the 16-km distance band. a survey start point was randomly selected within each block (buij et al. 2007) (figure 1). if the selected survey start point was inaccessible, it was displaced at 0° from the original location as far as necessary, for transects operated from it to be undertaken (buij et al. 2007). figure 1: map of the livingstone area showing the boundary (red line) of the ozp (green shaded area), the distance bands at 4-km intervals from the boundary of the ozp (black lines) and the 2 km × 2 km grid (with block colours indicating into which distance and the block falls). sampled blocks are highlighted (green outline), with the location of survey start points in sampled blocks shown (black cross). thirty eight out of the 39 sample blocks were surveyed (total area covered on 152 transects was 18.85 ha, with four replicates of each transect). one block within the 4-km distance band was not surveyed as it was located on livingstone airport runway and could not be displaced to an accessible area using the method described. three further blocks (two in the 4-km distance band and one in the 8 km distance band) were displaced at 0° from the original location as far as necessary for transects operated from them to be undertaken. at each survey start point, 155-m straight line transects were conducted in each cardinal direction, starting 6 m from the centre of the survey start point to avoid transect overlap. at the end of each transect line, a second parallel transect was undertaken 6 m to the right of the original and back towards the centre (buij et al. 2007). presence or absence was recorded as zero or one within 2 m either side of the transect line based on observation of signs of elephant presence, being either dung, tracks or debris from elephants feeding within the transect area. the types of indication observed were also noted. detectability creates variation in patch-occupancy surveys as not all signs of elephant presence may be observed (mackenzie et al. 2002, 2003). to overcome this, each of the four transect lines was independently surveyed by four different trained observers with no communication between them. surveys were also conducted during dry season, which reduces the negative association between detection and vegetation density (buij et al. 2007). inter-observer reliability was assessed using the results from all transects. the calculation was made using the formula a/(a+d), where a is the number of agreements and d is the number of disagreements (caro et al. 1979). detection probability was assessed from the findings of all observers from all transects with elephant presence. the calculation was made for all indications combined, and for each indication independently, using the formula p = n/tr, where n is the total number of agreements of elephant presence, t is the number of transects with elephant presence and r is the number of independent replications of those transects. to supplement data collected from transects, additional data sources were utilised to assess habitat occupancy: opportunistic sightings of elephant presence (animals, dung, tracks or feeding) observed between 2011 and 2012 during a previous monitoring programme (alert 2012); opportunistic sightings of elephant presence obtained whilst travelling to and from survey start points; and opportunistic questioning of community members in the region. whilst travelling to and from survey start points, community members encountered were asked to confirm if they were currently located at their normal residence. if a positive response was given, they were asked whether they had observed any presence of elephants within 1 km of their homestead during the current year (2015), and the same question over the prior years (2010–2014). results elephant presence was confirmed at four (10.5%) survey start points on 13 (8.6%) transects (figure 2). of these 13 transects, debris from feeding behaviour was observed on all 13, dung on 12 and tracks on 9. elephant presence as a result of transect surveys was therefore confirmed within the 4and 8-km distance bands. elephant absence, or non-detection, was confirmed beyond the 8-km distance band. opportunistically observed presence of elephants was determined in 2010 (n = 9), in 2011 (n = 13) and in 2012 (n = 3). opportunistically observed presence of elephant spoor was evident for 2011 (n = 64), for 2012 (n = 14) and for 2015 (n = 18). community advised presence of elephants was established for the period 2010–2014 (n = 2) and for 2015 (n = 6), and community advised absence was acquired for the period 2010–2015 (n = 14) (see figure 2 for all determinations). figure 2: map of the livingstone area with indications of elephant presence or absence. inter-observer reliability was assessed to be very high at 0.99, with agreement by all four observers on 150 out of 152 transects. detection probability for any indication of elephant presence was p = 0.96 (n = 52). detection probability varied, however, for each indication of presence: feeding p = 0.89 (n = 52), dung p = 0.73 (n = 48) and tracks p = 0.75 (n = 36). discussion detection probability for any indication of elephant presence was very high, suggesting that using multiple indicators of elephant presence, along with multiple replicates of each transect, is a robust method for patch-occupancy surveys of elephants. inter-observer reliability was also high, suggesting that the chosen indicators (feeding behaviour, dung and tracks) can be reliably assessed as present or absent by trained observers. the survey methodology undertaken by buij et al. (2007) focussed only on dung as an indicator of elephant presence. detection probability for dung in this survey was assessed as being the lowest of the three indicators. large tracts of burnt areas due to late dry season fires throughout the region may have negatively impacted the detectability of elephant presence indicators in affected survey areas. with all sources of elephant presence considered, elephant occupancy is concentrated within 8 km of the boundary of the ozp. further, presence is focussed around three apparent corridors: (1) around the sinde river to the west of the ozp, (2) the dambwa road, that runs from the ozp along the west of the dambwa forest, and (3) around the maramba river to the east of the ozp. significant elephant presence was also observed in the unfenced section of the mot np to the east of the ozp. seven community members advised of elephant presence outside of the 8-km distance band, with all but one stating that elephant presence was from one occasion only. one informant, located 11 km due west of the ozp, reported that in 2015 a breeding herd was present in the area for 4 days and that four bulls were resident in the area. the authors recommend that any planned intensive elephant monitoring in the region should focus on the ozp and the area up to 8-km distance from it, where presence is greatest, to maximise accuracy and cost-efficiency in determining the abundance of elephants. additional resources should be employed, if possible, beyond this limit, and also along the three identified corridors to assess the level of usage of these areas, and the extent of each corridor, including ascertaining whether they are migration corridors to other protected areas or local movement feeding corridors. the authors also recommend that a patch-occupancy survey is repeated at regular intervals to assess changes in elephant habitat usage in the region over time, and in response to changing land-use patterns and human–elephant conflict mitigation measures that may be introduced. acknowledgements the authors would like to thank the zambia department of national parks and wildlife and forestry department and the dambwa trust for facilitating the survey. thanks also go to wildlife encounter (zambia) for enabling the survey through the provision of vehicles, drivers and research assistants. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions d.a.y. was the project leader. d.a.y., j.a., l.c., a.k., c.k.l. and b.a.s. were responsible for project design. d.a.y., j.s.b., m.d.c., a.k. and d.s. undertook most of the data collection. c.k.l. undertook g.i.s. mapping. d.a.y., j.a, l.c., a.k. and b.a.s. wrote the manuscript. references african lion & environmental research trust (alert), 2012, elephant monitoring in the mosi-oa-tunya national park and livingstone area, unpublished report to the zambia wildlife authority. buij, r., mcshea, w.j., campbell, p., lee, m.e., dallmeier, f., guimondou, s. et al., 2007, ‘patch-occupancy models indicate human activity as major determinant of forest elephant loxodonta cyclotis seasonal distribution in an industrial corridor in gabon’, biological conservation 135, 189–201. http://dx.doi.org/10.1016/j.biocon.2006.10.028 caro, t.m., roper, r., young, m. & dank, g.r., 1979, ‘inter-observer reliability’, behaviour 69(3), 303–315. http://dx.doi.org/10.1163/156853979x00520 chishika, j., 2010, ‘current status of hec in zambia, zambian wildlife authority perspective’, in a. songhurst (ed.), working towards coordinated regional approaches in human-elephant conflict management: proceedings of a workshop for the kwando-kavango region organised by conservation international, pp. 46–47, conservation international, gabarone, botswana. douglas-hamilton, i., krink, t. & vollrath, f., 2005, ‘movements and corridors of african elephants in relations to protected areas’, naturwissenschaften 92, 158–163. http://dx.doi.org/10.1007/s00114-004-0606-9 dublin, h.t. & hoare, r.e., 2004, ‘searching for solutions: the evolution of an integrated approach to understanding and mitigating human–elephant conflict in africa’, human dimensions of wildlife 9(4), 271–278. http://dx.doi.org/10.1080/10871200490505701 ensvol consult, 2007, environmental impact assessment: dambwa forest joint management area lion rehabilitation project. chapter 3: description of existing environment, envsol consult, lusaka, zambia. graham, m.d., douglas-hamilton, i., adams, w.m. & lee, p.c., 2007, ‘the movement of african elephants in a human-dominated land-use mosaic’, animal conservation 12, 445–455. http://dx.doi.org/10.1111/j.1469-1795.2009.00272.x hoare, r., 2000, ‘african elephants and humans in conflict: the outlook for co-existence’, oryx 34, 34–38. http://dx.doi.org/10.1017/s0030605300030878 karidozo, m. & osborn, f.v., 2015, ‘community based conflict mitigation trials: results of field tests of chilli as an elephant deterrent’, journal of biodiversity and endangered species 3, 144. http://dx.doi.org/10.1111/10.4172/2332-2543.1000144 mackenzie, d.i., nichols, j.d., hines, j.e., knutson, m.g. & franklin, a.b., 2003, ‘estimating site occupancy, colonization and local extinction when a species is detected imperfectly’, ecology 84, 2200–2207. http://dx.doi.org/10.1890/02-3090 mackenzie, d.i., nichols, j.d., lachman, g.b., droege, s., royle, j.a. & langtimm, c.a., 2002, ‘estimating site occupancy rates when detection probabilities are less than one’, ecology 83, 2248–2255. http://dx.doi.org/10.1890/0012-9658(2002)083[2248:esorwd]2.0.co;2 okello, m.m., njumbi, s.j., kiringe, j.w. & isiiche, j., 2015, ‘habitat use and preference by the african elephant outside of the protected area, and management implications in the amboseli landscape, kenya’, international journal of biodiversity and conservation 7(3), 211–226. http://dx.doi.org/10.5897/ijbc2014.0795 simukonda, c., 2009, the elephant survey (2008) – a country report, zambia wildlife authority, lusaka. sitati, n.w., walpole, m.j., smith, r.j. & leader-williams, n., 2003, ‘predicting spatial aspects of human–elephant conflict’, journal of applied ecology 40, 667–677. http://dx.doi.org/10.1046/j.1365-2664.2003.00828.x article information authors: olaf l.f. weyl1,2 bruce r. ellender1,2 ryan j. wasserman1,2 darragh j. woodford1,2 affiliations: 1south african institute for aquatic biodiversity, south africa 2centre for invasion biology, south african institute for aquatic biodiversity, south africa correspondence to: olaf weyl email: o.weyl@saiab.ac.za postal address: private bag 1015, grahamstown 6140, south africa dates: received: 03 dec. 2014 accepted: 23 mar. 2015 published: 07 july 2015 how to cite this article: weyl, o.l.f., ellender, b.r., wasserman, r.j. & woodford, d.j., 2015, ‘unintended consequences of using alien fish for human benefit in protected areas’, koedoe 57(1), art. #1264, 5 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1264 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. unintended consequences of using alien fish for human benefit in protected areas in this essay... open access • abstract • introduction • conflicting stakeholder agendas in the limietberg nature reserve, western cape • angler access and illegal alien fish introductions in the goukamma nature reserve, western cape • creating dependencies may compromise long-term conservation objectives • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ there is increasing pressure on conservation agencies to allow access to natural resources within protected areas for human benefit. alien fishes are often seen as a convenient resource because their harvest does not conflict with conservation goals. however, allowing such access may have unintended consequences for managers. this opinion essay is intended to provide some insights into how promoting access to alien fish resources can add to the complexity of conservation interventions, may facilitate additional fish introductions and create dependencies on alien fish that could compromise potential eradication efforts. conservation implications: management plans for the utilisation of alien fishes by external stakeholders must include clear exit strategies so that the ability to eradicate when necessary or feasible is not compromised. introduction top ↑ by mitigating anthropogenic impacts, protected areas play an important role in the conservation of freshwater biodiversity (russell 2011). in south africa, there is increasing pressure for protected areas to sustain themselves economically and provide tangible benefits to bordering communities (e.g. van wilgen et al. 2013). for this reason, managers are often faced with the task of developing innovative avenues for increasing revenue and for developing livelihood opportunities for local communities. in south africa, freshwater fishes are often used to provide such opportunities and there are examples of recreational, subsistence and even commercial fishing in protected areas (e.g. ellender et al. 2010; mccafferty et al. 2012; van wilgen et al. 2013). public access to protected areas for angling has a long history, and until the 1980s the production and stocking of fishes to provide angling opportunities was part of the mandate of many provincial conservation authorities (see mccafferty et al. 2012). the tradition of catch and release recreational angling in protected areas is illustrated by the fierce opposition to suggested catch and kill regulations for alien fishes proposed in draft versions of the recently promulgated national environmental management: biodiversity act (10/2004): alien and invasive species regulations, 2014 (republic of south africa 2014a) and alien and invasive species list, 2014 (republic of south africa 2014b). as a result of this opposition, rainbow trout oncorhynchus mykiss and brown trout salmo trutta, fishes listed among the world's 100 worst invasive species (lowe et al. 2000), were excluded from the alien and invasive species list and catch and release angling for other alien and invasive fish species in formal protected areas was specifically included (republic of south africa 2014b). at first glance, alien fish appear to be a convenient resource for public use. after all, alien fishes are generally acknowledged as a major threat to biodiversity both inside and outside of protected areas because they impact on aquatic ecosystems by hybridising with native species, altering habitats, introducing parasites and disease, and by competing with and preying on native biota (ellender & weyl 2014). while their removal from protected areas is thus desirable (russell 2011), there are, however, several examples where the use of alien fishes in protected areas has resulted in unforeseen and unintended consequences for management. in this opinion essay, we present examples of how promoting access to alien fish resources can add to the complexity of conservation interventions, facilitate additional fish introductions and create dependencies on alien fish that may compromise potential eradication efforts. conflicting stakeholder agendas in the limietberg nature reserve, western cape top ↑ river fisheries for alien brown trout and rainbow trout in the western cape were managed by the cape department of nature conservation (cdnc) until this role was transferred to local angling organisations in the 1980s (skelton & davies 1986). a formal agreement in 1991 gave the cape piscatorial society (cps) control of trout fishing for five years in several streams in the limietberg nature reserve, which the cps is still managing even though the formal agreement is now outdated. the krom river, which flows through the limietberg nature reserve (figure 1a), is of particular interest at present. the current trout fishery in this river is managed for long-term sustainability by the cps, who promotes the utilisation of rainbow trout within the reserve by anglers. the provincial conservation agency, capenature, benefits through conservation fees generated when anglers book a reach of river to fish through the cps. the relationship between the conservation authority (capenature and its various precursors) and the cps has historically been positive and co-operative when there was historical legislative protection for trout in the cdnc ordinance (mccafferty et al. 2012). this protection was removed in the mid-1980s and the current management of the fishery, which includes limiting the number of anglers and enforcing a strict catch and release policy, ensures the long-term presence of rainbow trout (figure 1b) within the reserve, which would appear to be in conflict with the conservation objectives of protected areas. figure 1: (a) map of south africa indicating the location of examples of unintended consequences of promoting the use of alien fish. in the limietberg nature reserve, (b) rainbow trout onchorhynchus mykiss threaten (c) giant redfin minnows pseudobarbus skeltoni and management interventions are complicated by management agreements between the angling organisations and the conservation agency. in the goukamma nature reserve, the introduction of (d) common carp cyprinus carpio into groenvlei was most likely facilitated by the access for angling for introduced fishes such as (e) largemouth bass micropterus salmoides. (f) one of 450 subsistence anglers whose livelihood depends on access to lake gariep via the oviston nature reserve to harvest common carp. developing similar dependencies elsewhere may negate the ability of reserve managers to eradicate alien fishes. a significant conservation threat recently emerged with the discovery of a new species of redfin minnow within the krom river. the newly described giant redfin pseudobarbus skeltoni (figure 1c) has yet to be given an official conservation status, but since it is currently known only from the krom river and one other stream in the breede river catchment, evaluation using iucn red list criteria is likely to result in a ‘critically endangered’ status (chakona & swartz 2013). this discovery places the management of rainbow trout within the krom river in a new light, as the species is known to deplete or replace populations of redfin minnows (shelton, samways & day 2015). the presence of rainbow trout within the limietberg nature reserve directly threatens a native fish species (chakona & swartz 2014), and is therefore in conflict with the biodiversity conservation mandate of capenature. however, the cps is considered an important stakeholder for capenature and the existence of historical formal management agreements now adds an additional level of complexity to managing the krom river rainbow trout population. logically, capenature should take steps to reduce the trout population in this protected area to benefit the giant redfin. capenature has already successfully embarked on alien fish control projects to better conserve threatened fish species, notably the eradication of alien invasive smallmouth bass micropterus dolomieu from the rondegat river (weyl et al. 2014). however, project proposals by capenature to control trout in rivers have met with substantial opposition from trout anglers (ellender et al. 2014; marr, impson & tweddle 2012; weyl et al. 2014). as a result, capenature is unable to take decisive conservation action until they have amended the outdated management agreement with the cps to exclude the krom river as a trout water. these negotiations are now underway and it is hoped that a new management agreement with the cps will be concluded in 2015 (d. impson [capenature] pers. comm., 11 february 2015). angler access and illegal alien fish introductions in the goukamma nature reserve, western cape top ↑ illegal introductions of alien fishes for angling are a major problem faced by conservation authorities worldwide (johnson, arlinghaus & martinez 2009). in south africa, one such example is the introduction of common carp cyprinus carpio (figure 1d) into groenvlei, a natural lake in the goukamma nature reserve (figure 1a). historically, groenvlei contained only indigenous estuarine roundherring gilchristella aestuaria and cape silverside atherina breviceps (phair et al. 2015). as these fishes are too small (< 12 cm) to be of interest to anglers, largemouth bass micropterus salmoides (figure 1e) were legally introduced into the lake to develop a fishery in 1934 (de moor & bruton 1988). subsequently, alien mosquitofish gambusia affinis were introduced for mosquito control, bluegill lepomis macrochirus as a forage species for largemouth bass and mozambique tilapia oreochromis mossambicus for angling and to reduce algal growth (coetzee 1980; de moor & bruton 1988). largemouth bass are a popular target species for recreational anglers while bluegill and mozambique tilapia are now harvested by subsistence anglers (weyl, unpublished data). common carp were first reported from this lake in the 1990s (olds et al. 2011). the movement and stocking of common carp has been prohibited in the western cape province since 1947 (mccafferty et al. 2012) and since groenvlei lacks an inflowing river – thus depending entirely on rainfall, spring flow and seepage for inflow (parsons 2009) – their presence in the system is a direct result of an illegal introduction. as common carp are a popular angling species in south africa (ellender et al. 2014) we contend that the introduction was driven by the intention to develop an additional angling opportunity in the lake. recent surveys indicate that the two indigenous fish have co-existed with largemouth bass and other alien fishes for 80 years (phair et al. 2015; weyl, unpublished data). the main conservation concern regarding the recent common carp introduction therefore revolves around the potential impact of this fish on water quality – a conservation priority because groenvlei lacks an outlet and water exits the system mainly by evaporation (parsons 2009). common carp have been shown to affect water quality because their bottom grubbing during feeding suspends sediments thus increasing nutrient availability and turbidity (lougheed, crosbie & chow-fraser 2009). carp anglers also often pre-bait areas for carp fishing, a practice that under certain conditions has been shown to contribute substantially to anthropogenic eutrophication (arlinghaus & mehner 2002). for this reason, goukamma management is using gill nets to try to control carp numbers and discourages carp anglers from pre-baiting (keith spencer [capenature] pers. comm., 17 february 2015). we contend that in the case of groenvlei, the introduction of bass and subsequent angler access to this fishery facilitated the introduction of an additional invader which is likely to impact on water quality both directly (through feeding) and indirectly (through ground baiting), resulting in additional management costs. creating dependencies may compromise long-term conservation objectives top ↑ it is generally accepted that in the south african context the eradication of alien fishes is only possible in small, relatively isolated environments such as headwater streams (weyl et al. 2014). even then, this is an expensive exercise. the removal of alien smallmouth bass from a 4 km stretch of the lower rondegat river in the western cape, for example, cost zar3.3 million ($290 000) (impson, van wilgen & weyl 2013). this situation may be changing, as a result of rapidly evolving technologies for controlling alien fishes over larger spatial scales (britton, gozlan & copp 2011). researchers in australia, for example, are investigating the use of the cyprinid herpes virus-3 (mccoll, cooke & sunarto 2014) and genetically modified carp to produce male-only offspring (thresher et al. 2014) as potential agents to control common carp. this presents somewhat of a conundrum for conservation authorities in south africa, where poverty eradication, job creation and economic empowerment are national policy objectives, and where alien fishes provide considerable opportunities for food security and economic empowerment, either directly through commercial harvest or indirectly by the provision of equipment and services to recreational anglers (ellender et al. 2014). on lake gariep, for example, some 450 subsistence anglers (figure 1f) access the oviston (eastern cape) and gariep (free state) nature reserves (figure 1a) to harvest common carp, which is an important source of household food security and income (ellender et al. 2010). this dependency on the fish resource has already resulted in conflicts between fishers and reserve managers regarding access to the lakeshore (leon barkhuisen [department of economic development, tourism and environmental affairs, free state] pers. comm., 17 february 2015) and may result in an inability to eradicate common carp even if it becomes feasible to do so. conclusion top ↑ in south africa, aquatic ecosystems are under pressure from a variety of anthropogenic impacts, including catchment degradation, habitat modification, dewatering, pollution and the introduction of alien fishes (weyl et al. 2014). protected areas often offer the last refugia for native biota as many threats are significantly lower here than in surrounding areas. as a result the eradication of alien fishes should be prioritised as this is often the most significant threat in protected areas (russell 2011). while eradication may not currently be feasible because of the scale of areas affected, managers of protected areas need to be careful when promoting utilisation of alien fishes because this may increase costs of management and compromise long-term conservation objectives. in the limietberg nature reserve, for example, the ultimate removal of trout now requires considerable investment in stakeholder consultations, an essential component in management and insurance against possible reintroductions (ellender et al. 2014). in groenvlei, it was access to the lake for anglers that most likely led to the illegal introduction of common carp. this situation is not limited to south africa. in the pyrenees, for example, fishing has been banned in lakes protected by national parks as this is the only reliable management policy to avoid new introductions of trout (miró & ventura 2013). finally, managers of protected areas should take care when developing subsistence and commercial fisheries, as the concomitant social dependencies on alien fish could lead to an inability to eradicate them if it becomes feasible to do so. with regard to aquatic ecosystems, we believe that the needs of biodiversity conservation should generally trump social and economic considerations within protected areas. while this article is not intended to call for a change in the current management practice, we believe that stakeholder agreements regarding the utilisation of alien fishes must include clear exit strategies so that the potential eradication of alien fishes from protected areas is not compromised. acknowledgements top ↑ we would like to thank the national research foundation of south africa (nrf, uid: 77444, 86147, 88092) for its ongoing support. dean impson (capenature), keith spencer (capenature), leon barkhuisen (department of economic development, tourism and environmental affairs) and two anonymous reviewers are thanked for their valuable comments which helped shape this manuscript. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions o.l.f.w. (south african institute for aquatic biodiversity) was the project leader and lead author, b.r.e. (south african institute for aquatic biodiversity), r.j.w. (south african institute for aquatic biodiversity) and d.j.w. (south african institute for aquatic biodiversity) contributed to the study conceptualisation, writing and editing the manuscript. references top ↑ arlinghaus, r. & mehner, t., 2002, ‘socio-economic characterisation of specialised common carp (cyprinus carpio l.) anglers in germany, and implications for inland fisheries management and eutrophication control’, fisheries research 6, 19–33. http://dx.doi.org/10.1016/s0165-7836(02)00243-6 britton, j.r., gozlan, r.e. & copp, g.h., 2011, ‘managing non-native fish in the environment’, fish and fisheries 12, 256–274. http://dx.doi.org/10.1111/j.1467-2979.2010.00390.x chakona, a. & swartz, e.r., 2013, ‘a new redfin species, pseudobarbus skeltoni (cyprinidae, teleostei), from the cape floristic region, south africa’, zootaxa 3686, 565–577. http://dx.doi.org/10.11646/zootaxa.3686.5.5 coetzee, d.j., 1980, ‘zooplankton and environmental conditions in groenvlei, southern cape, during 1976’, journal of the limnological society of southern africa 6, 5–11. http://dx.doi.org/10.1080/03779688.1980.9633200 de moor, i. & bruton, m., 1988, atlas of alien and translocated indigenous aquatic animals in southern africa, south african national scientific programmes report no 144, csir, pretoria. ellender, b.r. & weyl, o.l.f., 2014, ‘a review of current knowledge, risk and ecological impacts associated with non-native freshwater fish introductions in south africa’, aquatic invasions 9, 117–132. http://dx.doi.org/10.3391/ai.2014.9.2.01 ellender, b.r., weyl, o.l.f., winker, h., stelzhammer, h. & traas, g.r.l., 2010, ‘estimating angling effort and participation in a multi-user, inland fishery in south africa’, fisheries management and ecology 17, 19–27. http://dx.doi.org/10.1111/j.1365-2400.2009.00708.x ellender, b.r., woodford, d.j., weyl, o.l.f. & cowx, i.g., 2014, ‘managing conflicts arising from fisheries enhancements based on non-native fishes in southern africa’, journal of fish biology 85, 1890–1906. http://dx.doi.org/10.1111/jfb.12512 impson, n.d., van wilgen, b.w. & weyl, o.l.f., 2013, ‘coordinated approaches to rehabilitating a river ecosystem invaded by alien plants and fish’, south african journal of science 109(11/12), art. #a0041, 4 pages. http://dx.doi.org/10.1590/sajs.2013/a0041 johnson, b.m., arlinghaus, r. & martinez, p.j., 2009, ‘are we doing all we can to stem the tide of illegal fish stocking?’, fisheries 34, 389–394. http://dx.doi.org/10.1577/1548-8446-34.8.389 lougheed, v.l., crosbie, b. & chow-fraser, p., 1998, ‘predictions on the effect of common carp (cyprinus carpio) exclusion on water quality, zooplankton, and submergent macrophytes in a great lakes wetland’, canadian journal of fisheries and aquatic sciences 55, 1189–1197. http://dx.doi.org/10.1139/f97-315 lowe, s., browne, m., boudjelas, s. & de poorter, m., 2000, ‘100 of the world's worst invasive alien species: a selection from the global invasive species database’, invasive species specialist group of the world conservation union (iucn), auckland. marr, s.m., impson, n.d. & tweddle, d., 2012, ‘an assessment of a proposal to eradicate non-native fish from priority rivers in the cape floristic region, south africa’, african journal of aquatic science 37, 131–142. http://dx.doi.org/10.2989/16085914.2012.666654 mccafferty, j.r., ellender, b.r., weyl, o.l.f. & britz, p.j., 2012, ‘the use of water resources for inland fisheries in south africa’, water sa 38, 327–344. http://dx.doi.org/10.4314/wsa.v38i2.18 mccoll, k.a., cooke, b.d. & sunarto, a., 2014, ‘viral biocontrol of invasive vertebrates: lessons from the past applied to cyprinid herpesvirus-3 and carp (cyprinus carpio) control in australia’, biological control 72, 109–117. http://dx.doi.org/10.1016/j.biocontrol.2014.02.014 miró, a. & ventura, m., 2013, ‘historical use, fishing management and lake characteristics explain the presence of non-native trout in pyrenean lakes: implications for conservation’, biological conservation 168, 17–24. http://dx.doi.org/10.1016/j.biocon.2013.07.016 olds, a.a., smith, m.k.s., weyl, o.l.f. & russell, i.a., 2011, ‘invasive alien freshwater fishes in the wilderness lakes system, a wetland of international importance in the western cape province, south africa’, african zoology 46, 179–184. http://dx.doi.org/10.3377/004.046.0109 parsons, r., 2009, ‘is groenvlei really fed by groundwater discharged from the table mountain group (tmg) aquifer?’, water sa 35, 657–662. http://dx.doi.org/10.4314/wsa.v35i5.49425 phair, n., barendse, j., smith, m.k.s. & von der heyden, s., 2015, ‘molecular analyses confirm genetically distinct populations of two indigenous estuarine fish species in an isolated coastal lake: implications for the management of introduced ichthyofauna’, conservation genetics, 05 february. http://dx.doi.org/10.1007/s10592-015-0701-9 republic of south africa, 2014a, national environmental management: biodiversity act (10/2004): alien and invasive species regulations, government printer, pretoria. republic of south africa, 2014b, national environmental management: biodiversity act (10/2004): alien and invasive species list, government printer, pretoria. russell, i.a. 2011, ‘conservation status and distribution of freshwater fishes in south african national parks’, african zoology 46, 117–132. http://dx.doi.org/10.3377/004.046.0103 shelton, j.m., samways, m.j. & day, j.a., 2015, ‘predatory impact of non-native rainbow trout on endemic fish populations in headwater streams in the cape floristic region of south africa’, biological invasions 17, 365–379. http://dx.doi.org/10.1007/s10530-014-0735-9 skelton, p.h. & davies, m.t.t., 1986, ‘trout in south africa’, ichthos special edition 1, 1–20. thresher, r., van de kamp, j., campbell, g., grewe, p., canning, m., barney, m. et al., 2014, sex-ratio-biasing constructs for the control of invasive lower vertebrates’, nature biotechnology 32(5), 424–427. http://dx.doi.org/10.1038/nbt.2903 van wilgen, n.j., dopolo, m., symonds, a., vermeulen, w., bester, e., smith, k. et al., 2013, ‘an inventory of natural resources harvested from national parks in south africa’, koedoe 55(1), art. #1096, 5 pages. http://dx.doi.org/10.4102/koedoe.v55i1.1096 weyl, o.l.f., finlayson, b., impson, n.d., woodford, d.j. & steinkjer, j., 2014, ‘threatened endemic fishes in south africa's cape floristic region: a new beginning for the rondegat river’, fisheries 39, 270–279. http://dx.doi.org/10.1080/03632415.2014.914924 filelist convert a pdf file! koedoe 19: 67-74 (1976) populasiesamestelling van die bontebok damaliscus dorcas dorcas in die bontebok nasionale park gedurende januarie 1974. g. de graaff posb11.\ 787, pretoria 0001 p. t. van der walt privaatsak x404, skukuw jj 50 l.j. van zyl posb11.\ 149, swellendam 6740 samevalling populasiegegewens van die bontebok damaliscus dorcaj dorcas in die bontebok nasionale park word verskaf. die spesie se sosiale struktuur sentreer basies rondom verskeie teeltroppe en 'n ramtrop. voortplantingsgegewens vanaf 1960--1973 word bespreek. abstract population data for the bontebok damaliscus dorcas dorcas in the bontebok national park are presented . the social structure of the species centres around several breeding herds and a ram herd. information pertaining to reproduction for the period 1960--1973 is discussed. inleiding die bontebok damaliscus dorcas dorcas is een van die seldsaamste soogdierspesies in afrika en waarskynlik die seldsaamste in suiderafrika. verder is dit een van die opvallendste en kleurvolste bokke wat ongetwyfeld bygedra het tot die afname in getalle wat sedert die jaar 1800 plaasgevind het. gepaardgaande hiermee, hey sy natuurlike verspreidingsgebied ook gekrimp. vollediger gegewens oor hierdie gebeure kan nageslaan word in bigalke (1955), barnard en van del' walt (1961) van der merwe (1968), de graaff (1972), david (1973) en van rensburg (1975). bespreking van die taksonomie en beskrywings van die anatomie en uiterlike voorkoms van die bontebok is te vinde in ansell (1968), harper (1945) en david (1973). die bontebok se voortbestaan is tans egter grootliks verseker, onder andere, in die bestaande bontebok nasionale park. dit volg ondermeer 67 dat daar sedert 1960 (toe die park tot stand gekom hetl 'n groter insig verkry is in die behoeftes van die populasie in die park ten einde die spesies lewenskragtig te hou. met verloop van jare is die aandag toegespits op die sosiale struktuur van die populasie en die gepaardgaande voortplantingspotensiaal. fisiografie en klimaat van die bontebok nasionale park die bontebok nasionale park is ongeveer 7 km suid van swellendam gelee (20°30' 0; 34°03's) en is ongeveer 2 786 ha groot. die kortste afstand na die suidwaarts-gelee indiese oseaan is ongeveer 50 km, terwyl die oos-wes langebergreeks ongeveer 9 km noord van die park gelee is. die hoogte bo seespieel wissel tussen 61 m en 200 meter. die standhoudende breerivier vorm die suidwestelike grens van die park. volgens theron ( 1967) bestaan sowat 90% van die parkoppervlakte uit alluviale en gruisterrasse terwyl dagsome van gesteentes van die serie witteberg slegs langs die breerivier voorkom. die park is gelee in die winterreenvalstreek van die republiek van suid-afrika en toon normaalweg 'n piekreenval periode in die lente. vema' 1961-1973 word 'n gemiddelde jaarlikse reenval van 526mm aangctckcn. die temperatuur-uiterstes wissel tussen 2°c tot 42°c. die heersende winde is suidoos in die somer en noordw(;c in die winter. plantegroei die algemene plantegroeigeaardheid in die oostelike, suidelike en noordelike gedeeltes van die park is ' n oorwegende onsmaaklike, onbenutbare struikgemeenskap met 'n onbeduidende graskomponent. teen die hange van enkele valleie wat voorkom, is elytropappus rhinocerotis en cliflortia ruscifolia dominant terwyl die ander gruisterrasse gcdomineer word deur leucadendron salignum, pelargonium ovale, berkheya armata en serruria fucifolia. grasse wat hier verspreid voorkom is themeda triandra, cymbopogon marginatus en eragrostis capensis (grobler en marais 1967). i n die suidwestelike asook 'n klein gedeelte van die oostelike flank van die park, word 'n oorwegende grasstand aangetref in 'n sanderige laagland wat swak dreineer. kenmerkende grasse alhier is merxmuellera stricta, stipagroslis z.eyheri en eragroslis curvula. die park besit 'n merkwaardige rykdom aan plantsoorte. tot op datum, is daar 460 spesies gedentifiseer waarvan 61 geofiete is. verder behoort 52 tot die familie graminae en 21 tot die familie cyperaceae. min gegewens is beskikbaar oor die weidingsvoorkeure van die bontebok. wat wel bekend is, is dat die dier 'n strawwe benutter is van die kort-tot medium-hoogte grassoorte soos cynodon dactylon, themeda triandra, digitaria eriantha en die eragrostis-soorte. van die 52 bekende grasse in die park, kan ii as smaaklik vir die bontebok beskou word, 18 halfsmaaklik en 23 as onsmaaklik. slegs ongeveer 5% van die struike in die park word deur die bontebok benut. 68 m ateriaal en m etode gegewens gebruik in hierdie artikel is verkry uit diverse literatuurverwysings, uit inligting ter insae in leers van die nasionale parkeraad, uit persoonlike waarnemings van die plaaslike natuurbewaarder (l.j. van zyl) en is grotendeels opgestel ten tye van 'n besoek aan die park gedurende 1974.01.06-14. resultate en bespreking a. struktuur aangesien die bontebok primer 'n kuddedier is vind tropvorming plaas en behels dit ook 'n bepaalde sosiale struktuur. teeltroppe dit is gevestigde eenhede wat lewensbelangrik is vir die bontebok in die park. dit bestaan gewoonlik uit 'n aantal ooie en lammers (gewoonlik jonger as 12 maande) wat in 'n afgebakende gebied loop, tesame met 'n territoriale ram wat die ooie en lammers deurgaans probeer bymekaar hou enlander ramme uit sy afgebakende gebied hou. die ooie en lammers is egter nie streng gebiedsgebonde nie. ramtrop die ramtrop funksioneer basies as 'n enkele eenheid in die park. die trap beweeg op 'n losse basis oor die hele park en is nie gebiedsgebonde nie. konfrontasies tussen volwasse ramme van hierdie trop en 'n teeltropram wat sy gebied wil verdedig, kom dikwels voor. daar is geen rangorde of dominasie-hierargie in die ramtrop nie. uitgewerkte ramme kom daarin voor asook jong ramme wat vandaar uitbreek om eventueel suksesvol territoriaal te word. aanvulling vind gewoonlik plaas vanuit die teeltroppe deurdat jaaroue lammers geleidelik die teeltrappe verlaat, veral wanneer die teelooie weer begin lam. alleenloperramme breek ook van die ramtrop weg en kom verspreid oor die park voor. dit is moontlik dat van hulle besig is met die afbakening van territoria. 'n skematiese voorstelling van die bontebokpopulasie in die park gedurende januarie 1974 word in fig. 1 weergegee asook die populasiesamestelling in tabel 1. b. voortplanting die bontebok is 'n seisoenale teler. die paartyd strek vanaf januarie tot begin april en die draagtyd wissel van 230 tot 254 dae. david (1973) meld dat die ooie gedurende die tweede jaar in oestrus kom, d.w.s. ongeveer op 27 maande ouderdom. die jaarlikse verloop van geboorte gedurende 1964-1972 word weer69 o 1 inpiwiov i ramme ~ ocye .20 0 lammers m oveii. a5 i jaai{ /5 ~vj(iig.b.4its t'.lr"o,.."e ~ ~ on~rii"ific~'" ~flt.aiiif"'i(o~e ~ ~l()sldi'fi;1( 1t,aiiiimii ~ te~ltif.opfi>e fig. i. individuele tropsamestelling van die bontebok populasie in die bontebok nasionale park (1974.01.12). tabell populasiesamestelling van die bontebok in die bontebok nasionale park (1974.01.12) lammers lammers lammers ramme ooie (onder (ouer as (ouer as 1 jaar) 1 jaar) 2 jaar) territoriale ramme met ooie en lammers 18 53 37 6 territoriale ramme met ooie sonder lammers 9 16 alleenloperramme 13 ramtrop 104 28 76 totaal 144 69 37 34 76 70 gegee in tabel 2 waaruit dit blyk dat die jaarlikse geboorteperiode 'n redelike konstante aanvangstyd het aan die begin van september, waarna die geboortetempo progressief toeneem en gewoonlik eers na twee maande (november/desember) 'n toppunt bereik. hierdie toppunt word dan geredelikerwys gehandhaaf vir ongeveer 3 tot 4 maande. die gemiddelde jaarlikse getalaanwas sedert 1960 is ongeveer 54 persent. die jaarlikse getaltoename in bontebokramme, -ooie en -lammers (asook die vrektes) vanaf 1960-1973 word weergegee in fig. 2, terwyl die moontlike verwantskap tussen geslagsverhouding (9 :0') en lampersentasie weerspieel word in fig. 3. / 1'\ 1-"'" _ r~ares i d ii'hl i#( 9 vaovt/1( v l laiiimm$ / ....'" 1/ / v /' 1\ / v v 1/ 1f~~ 1/ i~ /' v :'" / 1"'1-2 60 61 6. '5 " 61 " " lj ! " :rarie~ fig. 2. lewenshistogram van die bontebok in die bontebok nasionale park 1960-1973. 9 :d' l~m % / : 1.,0 100 1: i, q ,. /:/, b '0 / : ',7 >. 1: 1, 6 60 1 : 1,5 <0 /:'," '0 i 1,3 30 i : '}1 '0 f : ~ i 10 i : /jo 0 i i " i i i i ;ll: , , , , " , -"\ , , , ';..-..... ~ : geslagsv£~ho(/ding , , , i " i )( , '", , 60 61 62 i., bit 6s 6' 67 6' h 70 71 72 73 fig. 3. geslagsverhouding (9 :0') en lampersentasie van die bontebokke in die bontebok nasionale park 1960-1973 . 71 ....., i'..:) 1964/ 65 27 aug. m idde! sept. 5 eind sept. 9 middcl okt. 19 eind okl. 29 eind nov. 29':' m iddcl des. 30 eind des. 31 ,', = 2 vrektes 196 5/66 8 sept. eind sept. 12 eind okt. 25 eind nov. 25':' eind des. 26 eindjan. 23 ':' = 4 vrektes 1966/ 67 eind aug. eind sept. eind okt. eind nov. eind des. eindjan. tabcl2 j aarlikse vetloop van bontebok-geboottes 1964-1972 1967/68 1968/69 1969/70 1970/71 1971/72 eind aug. :j 15 aug. 1 1 sept. 1 10 aug . 1 19 aug. 3 13 eind sept. ? 22 aug. 2 middel sept. 2 eind sept. 11 :' eind okt. 38 eind sept. 25 eind sept. 4 middel okt. 17 ? eind nov. 40 eind okt. 44 mid.okt. 24 eind okt. 20 27 eind des. ? find nov. 46 eind okt. 30 middel nov. 33 30 9jan. 45 eind des. 49 eind feb. 31 eind nov. 33':' 20 feb. 47 eindjan. 51 middel feb . 35 eind feb. 53 eind feb. 37 mid. mrt. 39 ceen mid. april 41 waarneillings ':' = 5 vrektes tabel3 lewenstabel van die bontebokkudde in die bontebok nasionale park 196{}--1973 totaal geslagslamjaar 9 aanwas vrektes in verpersenpark houding tasie 0 9 9:0 1960 39 22 11 2 2 68 1 : 1,8 50% 1961 43 25 17 4 80 1 : 1,7 68% 1962 50 30 21 6 3 92 1 : 1,6 70% 1963 59 33 32 5 4 121 1 : 1,5 82% +6':' 1964 74 47 31 9 145 1 : 1,4 58% +3':' +6':' 1965 83 62 35 12 10 158 1 : 1,3 56% 1966 91 67 30 3 5 180 1 : 1,3 44% 1967 103 77 47 13 6 208 1 : 1,3 61% 1968 120 88 55 4 4 255 1 : 1,3 62,5% 1969 151 104 53 9 5 294 1 : 1,45 51% 1970 172 122 36 45 22 263 1 : 1,40 29% 1971 145 118 39 12 10 280 i : 1,2 33% 1972 153 105 55 13 15 285 1 : 1,5 52% ( ?) (?) (?) 1973 164 84 34 5 6 263 1 : 2,0 41% -4':":' -4':":' geskenk deur mnr. bowker ::::;: geskenk aan belgie vit be ide figure blyk dit duidelik dat daar 'n oonnaat manlike diere tot stand begin kom het wat miskien 'n nadelige uitwerking het op die teeltroppe: die territoriale ram moet sy gebied en ooie beskerm teen besoekers energie wat hy andersins aan die ooie kon bestee (om daardeur , n hoer ooi-besetting te verkry) word nou aan gebiedsbeskerming gewy en wat indirek 'n rol kan speel in verlaagde lammeroeste. 73 vanaf 1960-1963 het die lampersentasie van die populasie toegeneem tot 'n rekord 83 persent. daarna het 'n afname gevolg en die dalende tendens is op 'n oneweredige wyse voortgesit blykbaar onafhanklik van die geslagsverhoudingstendense. dit kan daarop dui dat daar 'n proteientekort in die dieet van die ooie teenwoordig is deels as gevolg van die lae grasbedekking. 'n verdere moontlike oorsaak is die teeltroppe sonder lammers (33% van die totale aantal diere) wat voorkom. die 9 ramme·wat die 16 ooie van hierdie troppe beheer is waarskynlik onvrugbaar. ten slotte kan die aandag gevestig word op 'n vooriopige lewenstabel van die bontebokpopulasie in die park (tabel 3), gebaseer op gegewens ingewin tussen 1960 en 1973. op hierdie stadium is daar nog te min feitlike gegewens oor die populasie (bv. geen groeireekse is beskikbaar vir ouderdomsbepalings nie) en daarom is die lewenstabel 'n voorlopige poging. dit gee egter 'n aanduiding van die aanwas, mortaliteit, volwasse geslagsverhouding, lampersentasie en die groot totaal van die bontebokke wat per jaar in die park teenwoordig was vanaf 1960 tot 1973. wat op hierdie stadium dus nog ontbreek is gegewens insake oorlewingskurwes, ouderdomspiramides en die rol van omgewingsweerstand versus die reproduktiewe potensiaal van die populasie. verwysings ansell, w. f. h.1971. order artiodactyla. in meester,]. en h. w. setzer (reds.) the mammals oj africa an identification manual. washington d.c.: smithsonian institution press. barnard, p.]. en k. van der walt. 1961. translocation of the bontebok from bredasdorp to swellendam. koedoe 4: 105-109. bigalke, r. 1955. the bontebok with special reference to its history and preservation. fauna flora pretoria 6 :95-115. david,]. h. m. 1973. the behaviour of the bontebok damaliscus dorcas riorcaj (pallas 1766) with special reference to territorial behaviour. z. tierpsychol. 33:38-107. de graaff, g. 1972. die bontebok-trots van die suid-westelike distrikte. custos 1(4):3,5,19,41. grobler, p.]. en]. marais 1967. die plantegroei van die nasionale bontebokpark, swellendam. koedoe 10: 132-146. harper, f. 1945. extinct and vanishing mammals oj the world. am. comm. int. wildlife protn. special pub!. no. 12 :850 pp. theron,]. m. 1967. geologie van die bontebokpark, distrik swellendam. koedoe 10: 147-148. van der merwe, n.]. 1968. die bontebok. koedoe 11 :161-168. van rensburg, a. p.]. 1975. die geskiedenis van die nasionale bontebokpark, swellendam. koedoe 18: 165-190. 74 page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 abstract introduction rwandan culture history and forest the context of buhanga sacred forest material and methods results discussion conclusion acknowledgements references appendix 1 about the author(s) runyambo irakiza africa rice center (africarice), east and southern africa, weed sciences, dar es salaam, tanzania association pour la conservation de la nature au rwanda (acnr), kigali, rwanda minani vedaste institute of scientific and technological research, national herbarium of rwanda, huye, rwanda bizuru elias college of science and technology, university of rwanda, rwanda brigitte nyirambangutse college of science and technology, university of rwanda, rwanda nsengimana joram serge association pour la conservation de la nature au rwanda (acnr), kigali, rwanda ndimukaga marc association pour la promotion des etudes d’impacts environnementaux au rwanda (apeier), kigali, rwanda citation irakiza, r., vedaste, m., elias, b., nyirambangutse, b., serge, n.j. & marc, n., 2016, ‘assessment of traditional ecological knowledge and beliefs in the utilisation of important plant species: the case of buhanga sacred forest, rwanda’, koedoe 58(1), a1348. http://dx.doi.org/10.4102/koedoe.v58i1.1348 original research assessment of traditional ecological knowledge and beliefs in the utilisation of important plant species: the case of buhanga sacred forest, rwanda runyambo irakiza, minani vedaste, bizuru elias, brigitte nyirambangutse, nsengimana joram serge, ndimukaga marc received: 21 aug. 2015; accepted: 05 feb. 2016; published: 22 july 2016 copyright: © 2016. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract traditional ecological knowledge is an integrated part of the african people and indeed the rwandese for cultural purpose. buhanga sacred forest is a relict forest of tremendous ecological importance to rwandan society located in musanze district. the aim of this study was to assess the traditional ecological knowledge and belief in the utilisation of some important plant species for the conservation of buhanga sacred forest. ecological information about ethnomedicinal and traditional practices were collected following structured questionnaire through interview involving eight traditional healers and three focus group discussions. data were collected from the natural habitats, home gardens, farmlands and roadsides of buhanga sacred forest. a total of 45 botanical taxa belonging to 28 families were reported to be used by the local community. species such as brillantaisia cicatricosa and senna septemtrionalis were the popular species cited by traditional healers to treat human and animal diseases and ailments, respectively. the results of the study indicated that because of the cultural norms and values associated with the sacred forest, this has led to non-exploitation. the study presents key sites and plant species in which their use and belief can lead to their conservation. however, not only is it imperative to conserve traditional local knowledge for biocultural conservation motives but there is also need to train traditional healers on how to domesticate indigenous species as conservation measure because some species have become susceptible to extinction. conservation implications: highlighting indigenous species investigated in this research will provide a powerful tool for ensuring biodiversity conservation through community participation in a country of high population density in africa. some plant species that provided satisfactory local health traditions among communities surrounding buhanga can contribute as good material for further research in rwanda. introduction traditional ecological knowledge is defined as a cumulative body of knowledge, practice and belief. it is a mutual relationship between living beings (including humans) and its environment which evolves by adaptive processes and are handed down through generations by cultural transmission (berkes 1999). such knowledge has contributed to conservation of biodiversity (gadgil, berkes & folke 1993), rare species (colding 1998) and protected areas (johannes 1998) as well as to sustainable resource use (berkes 1999) from one area to another. in africa, there are important elements to take into account regarding indigenous or traditional spirituality towards the nature. the value of traditional knowledge of natural resources is widely recognised among indigenous people because it was accumulated and transferred orally across generations. according to charnley, paige and jones (2007) and berkes, colding and folke (2000), the integral success of the traditional ecological knowledge into biodiversity conservation depends on active participation of the knowledge holders, such as indigenous communities and institutions. to avoid loss of biodiversity, traditional rules play an important role when developing conservation strategies for rare species by limiting illegal activities such as selective trees felling (ormsby 2013). some sacred species constitute excellent totems. totems are often plants or animals that are a stand-in of its owner and their presence dissuade from illegal access to sacred areas. furthermore, these sacred species contribute significantly in conservation by acting as umbrella species to the overall ecosystem (ndimukaga 2009). sacred place functions under the principle of faith and fear of god and other supernatural forces. this principle requires honesty, respect of ancestors and observation of moral values (kamga-kamdem 2008). rwandan culture history and forest in many sub-saharan african societies, the traditional ecological knowledge among indigenous people is synonymous with spirituality, which contributed to sustain the sacred forests. rwanda, one of the smallest countries in the world with only 26, 338 km2, has a rich forest heritage. the value of rwandan culture is based on oral history, ecology and nature, geographical sites and by igitaramo. igitaramo constitutes a gathering of families, communities and friends with songs and dances. during pre-colonial time, rwandan society was inherently linked to forest, which hides great lessons concerning rites, ancestral customs, taboos, arts, crafts, music, dance and many more. such integral links led to the emergence of traditional medicine (karangwa 1997). the rwandese are aware of the existence of these traditional healers who provide home-based healthcare. locally, they are known as abavuzi gakondo or abapfumu. the former denotes traditional healers involved in the use of plants and the latter denotes those who are involved especially in the ritual component of traditional medicine (rwangabo 1993). but it has been said that sometimes abapfumu use leaves, roots or barks of plants to protect their clients in exchange for money (adenkule 2007). as a matter of fact, traditional ecological knowledge is still very much alive in rwanda because it contributes significantly to solve health problems. in rwanda, cutting of certain tree species is seen as a taboo as well as killing of indigenous animal such as primates, elephants, leopards, birds, etc. according to kimenyi (1989), this conservation emphasised the symbiotic and interdependence relationship between people, plants and animals. it is for these reasons that cultural values are an essential component of every society and they act as checks and balances in the management of natural resources (verschuuren et al. 2010). the context of buhanga sacred forest buhanga sacred forest is a relict forest of tremendous ecological importance to rwandan’s society for cultural purposes. it lies in volcanic belt and has one of the highest population density in rwanda and africa, with up to 500–700 people per km2 (nisr 2012; plumptre, masozera & vedder 2001). buhanga relict forest contains sacred water-spring, marshland, medicinal plants, native trees and even a last refuge for arboreal, migratory birds. for example, african pitta pitta angolensis is a migrant bird that breeds in central tanzania, malawi, southeast democratic republic of congo, eastern zambia, zimbabwe and possibly northern south africa that was sighted in buhanga sacred forest (birdlife international africa partnership e-bulletin 2008). unfortunately, reports indicate that this relict forest is facing many threats as a result of human interference on the environment in various ways, such as harvesting of firewood and agriculture (minitere & cgis-nur 2007; rema 2009). in fact, high pressure of disturbance of natural reserves in rwanda was noticed until the resettlement of refugees after the genocide during the year 1997–1998 where people both settled and farmed within many natural reserves (plumptre et al. 2001). it should be mentioned that in most cases natural resources are still viewed as limitless in the mind of the resource-poor local communities in rwanda as a source of income. however, it should be seen furthermore that without any practical actions for sustainable utilisation of so-called ‘limitless resources’ and usage of traditional ecological knowledge, there is a risk of genetic loss and lack of traditional information from generation with time. to address the issue of loss of this critical cultural forest in rwanda, the goal of this study is to document the traditional ecological knowledge and belief in the utilisation of some important plant species of buhanga sacred forest. hence, there is a need for such information and documentation with the help of local communities to develop comprehensive and useful strategies for public awareness but also to formulate recommendations for monitoring natural resources for better future (figure 1). figure 1: view of buhanga sacred forest. material and methods geo-ethnographical overview of the study area buhanga sacred forest is an archaic forest located in northern province, district of musanze, nkotsi sector and bikara cell, at the right side of the road that leads towards vunga business centre. the site is situated at approximately 114 km from kigali, the capital city of rwanda and covers an area of 33.252 ha, of which 16.0011 ha is still intact and located between 1°34’00” and 1°34’30” south latitude and 29°34’30” and 29°37’30” east longitude at an altitude ranging between 1623 m a.s.l. and 1657 m a.s.l. geo-climatically the area falls within the tropical rainforest of the volcano massif where climate is controlled by various factors such as altitude, latitude and vegetation. the climate is cool and wet and temperatures fluctuate around 20 °c during the day. most of the lands surrounding buhanga sacred forest are used for agriculture subsistence. near subsistence agriculture, coffee, tea and pyrethrum plantations are prominent land uses as cash crop generating local employment. projects funds from government and international non-governmental organisations have been used for a range of activities from environment protection (tree planting, soil erosion control and beekeeping) to limit access in the sacred forest (figure 2). figure 2: map of the study area. ethnobotanical survey our study was carried out during the period of 26–31 may, 2008. a reconnaissance survey was conducted in abadahemuka cooperative at nkotsi-bikara, musanze district. in rwanda, most of the traditional healers are grouped into cooperatives under the supervision of the ministry of health in close collaboration with the institute of scientific and technological research. the traditional healers provided the information about traditional healing practices for both humans and animals, while the elders provided information regarding rituals and beliefs surrounding the forest. the information regarding the use of medicinal plants was collected following structured questionnaire through interviews involving eight traditional healers (five men and three women) aged between 41 and 80 years. these traditional healers were recognised by the local government authorities and selected based on their reputation on herbal medicinal uses. plant species mentioned by traditional healers were visited inside the relict forest and home gardens to verify the reliability of data obtained during interviews (alexiades 1996; gerique 2006). the recorded field data include the following: collection number, plant local name, scientific name, habit, plant parts used, mode of preparation, habitat and locality (martin 1995). the information regarding the traditional practices and utilisation of buhanga sacred forest were captured through three focus discussion groups. each discussion group comprised 15 persons (10 men and 5 women) who were natives of the nkotsi-bikara village aged between 50 and 80 years; inclusion was based on a good knowledge of buhanga sacred forest history. during the discussion, each member of the group took about 5–8 min to talk about the importance of cultural knowledge and use of buhanga sacred forest. species information provided either during the interview or group discussion were recorded. when a species was physically observed in home garden and not in the forest and vice versa, the information regarding that species was collected. species recognised only through vernacular names without physical identification either in home garden or in the forest was considered unreliable and rejected. species botanical identification was carried out in the national herbarium of rwanda using voucher specimens, identification keys, field guides and the flora of rwanda (troupin 1966, 1978, 1982, 1983, 1985, 1988; and fischer & dorothee 2008) (see picture showing different sites of study area in appendix 1). data analysis validation of plant names, family and plant authority were carried out using the royal botany garden and missouri botanic garden plant names database (http://www.theplantlist.org). collected data were analysed using descriptive statistics. the determination of relative frequency of citation of reported medicinal plant species was done using the following formula: rfc (%) = (fc/n) × 100             [eqn 1] where, fc is the number of informants mentioning the use of medicinal plant species and n is the total number of informants participating in the survey. results plant diversity this study recorded 45 plant species that were used by local communities in the vicinity of buhanga sacred forest. of the 45 species, 38 plant species were categorised in 34 genera and 19 families to have medicinal value. in terms of species composition of medicinal plants, the family of asteraceae, acanthaceae and fabaceae had three species each followed by the family of chenopodiaceae, rutaceae, rhamnaceae, ranunculaceae, polygonaceae, solanaceae, menispermaceae, verbenaceae, urticaceae and cucurbitaceae with two species each. the remaining 14 families were represented by a single species each (see figure 3). regarding life form, 21 species (47%) were herbs, 8 species (18%) were trees, 10 species (22%) were climbers, 4 species (8%) were shrubs and 2 species (4%) were grasses (see figure 4). figure 3: distribution of 45 identified plant species among botanical families. figure 4: life form proportion of plant species. plant species role in ecological, social and cultural welfare results of medicinal plants collected from different habitats show that most of the plant species, about 29 species (62.22%) were harvested from natural vegetation followed by 9 species (20.00%) under cultivation in home gardens, 6 species (13.33%) collected in farming plots and 2 species (4.44%) found near the roadsides. the goods and uses of the vegetation in our study area showed that most of the plant species are used for medicinal purposes (38 plant species [74.51%]) followed by raw materials (8 species [15.69%]), social symbol (3 species [5.88%]) and 2 species (3.92%) for spiritual purpose (see figure 5 and table 1). figure 5: (a) origin of medicinal plants and (b) social welfare of plant species. table 1: plant species uses by local communities as raw material. interviewees from the three group discussions recounted stories of transgressors who had taken resources from a sacred forest and misfortunes fell on them (see table 2). for example it was said that villagers would not dare to enter the buhanga sacred forest for fear of angering some spirits in control of the forest. they believe that these spirits were able to curse them by sending a resident giant snake protective of the surrounding marshland called igishanga cya gihanga. moreover, buhanga sacred forest is a habitat for many rare species of snakes which upon sight a curse follows. calamities such as drought were also associated by such omens. table 2: name of sites and species of spiritual importance among local people. medicinal flora of the 38 plant species reported to have medicinal uses, 29 species distributed in 19 families were reported to be used against human health problems, six species belonging to six families were used against cattle diseases and three species belonging to three families were used against both human and cattle diseases. the most striking diseases recorded in 17 human health problems were the poison vomiting treated by six plant species followed by mental diseases and kilondatumbo (symptoms of kilondatumbo: painful sensation in the digestive system with presence of ulcers in the ileum and rectum) treated with five specific plant species (table 3). stomach ache was treated by three plant species while other plant species were used to treat one or two diseases (see table 3). in regard to ethnoveterinary medicine, the most remarkable diseases were tick-borne diseases (east coast fever, babesiosis and anaplasmosis) treated by four species followed by sexual dysfunction treated by two species. all other diseases were treated by one species each (see table 4). table 3: medicinal plants used against human health problems. table 4: plants used in veterinary medicine. regarding of some medicinal plants popular than others, brillantaisia cicatricosa was the most popular plant cited by 5 traditional healers for it medicinal value (relative frequency of citation [rfc] = 62.5%) followed by three plant species senna septemtrionalis, thalictrum rhynchocarpum and solanum terminale mentioned by 4 traditional healers (rfc = 50%). the species zanthoxylum sp., rumex abyssinicus, prunus africana, ranunculus multifidus, tragia brevipes, urtica massaica and ajuga alba were mentioned by 3 informants (rfc = 37.5%) while chenopodium opulifolium, cyathula cylindrica, solanum aculeastrum and solanecio mannii were mentioned by only 2 traditional healers (rfc = 25%) (see tables 3 & 4). concerning plant parts used frequently for medicinal preparation, the leaves were the most used parts followed by both leaves and tree barks in human health problem representing 81% and 6% respectively. regarding the ethnoveterinary medicine the leaves were also the most plant part used followed by both roots and tree barks and flowers representing 78% and 11% each. the majority of remedies were prepared in the form of juice from freshly collected plant parts. the juice was prepared by pounding or crushing a plant part in a wooden or stone motor and pesters. water was the most liquid substance used to dilute the prepared juice. the remedies were taken orally, accounting 84% of medicinal plants use, followed by external application applied typically on skin representing 8% and 5% on ear application except remedies from the species eucalyptus maidenii giving anally as smoke to treat haemorrhoids and fresh leaves from t. brevipes rubbing on painful join to treat rheumatism accounting both 3% (see figure 6). figure 6: proportion of plant parts used for human and cattle health problems: (a) proportion of plant parts used for human health problems; (b) proportion of plant parts used for cattle health problems. discussion taking consideration the size of buhanga sacred forest, there is a high percentage of plants being naturally harvested in the forest by local communities. the fact that more than a half of remedies were prepared from herbs in natural habitat than cultivated or home gardens indicates that the natural area of buhanga sacred forest is very important for local communities to satisfy their home-based healthcare. our finding is similar to the finding of edwards (2004) reporting that 2/3 of medicinal plants used worldwide are harvested from the natural environment. during interview, traditional healers reported that more medicinal plants were harvested previously than now in buhanga sacred forest and some species formerly forest-habiting species are now rarely encountered in the area. this is the case of the species p. africana that became extinct in the natural forest while c. cylindrica still existed occasionally due to overharvesting in the past. studies involving rwanda on medicinal plants reveal decline of these resources (stewart 2003). our research findings revealed that the poison vomiting, mental diseases and kilondatumbo in human health problems were the most cited diseases in our study area because remedies against these three predominant health problems were mostly prepared. in a situation where several drugs in one area are required, this indicates a prepotency of a particular disease (dawit & ahadu 1993). the potential of medicinal plants recorded in this study were also confirmed by other research conducted elsewhere in rwanda (dessouter 1991; kamagaju et al. 2013; kayonga & habiyaremye 1987; mbarubukeye & niang 1996; rwangabo 1993). as per the ethnoveterinary medicine represented mostly by tick-borne diseases, suggests that these diseases were very important in the area. according to byavu et al. (2000), mbarubukeye (1991) and van puyvelde et al. (1985), the east coast fever (theileriosis) played a vital role in cattle mortality in rwanda, particularly in exotic pure breed crossing with the ankole breed while anaplasmosis and babesiosis represented an enormous cost in term of time and money. also study conducted by nshimiyimana and mutandwa (2010) indicates that damages attributable to ticks borne diseases in rwanda were considered as the first economic damage of bovine production transmitting sucking blood parasites diseases such as theileriosis, anaplasmosis and babesiosis. with more than 70% of leaves use as ingredient in making of medicinal concoction indicates that leaves are more required to solve the health problem in our study area. in reference to dawit and ahadu (1993) and poffenberger et al. (1992), the normal harvesting leaves estimated of up to 50% does not significantly affect the growth of plant species or does not cause a great danger to existence of individual plant when compared to the collection of underground part, stem or whole plant from both ecological points of view and survival of species. the extinction of p. africana as reported by the healers is due to overutilisation of both its leaves and barks as ingredients and was replaced by agricultural activities. the same were reported by sayer, harcourt and collins (1992) stewart (2003) and minitere (2003). in general there was no really threat of overharvesting of plant in buhanga sacred forest as it used to be in the past because the remedies are prepared and used locally, and therefore harvested in small quantities. traditional healers reported that they do not store remedies for a prolonged period of time because when the need comes, they go out to collect plants, prepare the remedies and use it for treatment. however the importance of buhanga were known a long time ago, this sacred forest has acquired since 2005 a status of protected area as a part of the volcanoes national park from the former rwanda office of tourism and national parks (ortpn), now rwanda development board (rdb) (rema 2009). in the same context, culture and spiritual beliefs plays an important aspect in the conservation of buhanga sacred forest. recently, water springs from buhanga are still used as source of clean water for local people in the area and caves and some tree species (dracaena steudneri, erythrina abyssinica and ficus thonningii) are important for their ritual and ceremonies. conclusion this study has revealed the importance of traditional ecological knowledge in buhanga sacred forest plays a vital role in our heritage. the high plant diversity used in traditional medicine to treat different ailments was localised mostly in the natural habitat and does not really present threat of overharvesting because remedies are prepared on a need-to-need basis. also this study has shown how cultural values and beliefs are crucial tools for conservation and should therefore be incorporated in all management plans of buhanga sacred forest. in light of this research finding, we recommended that traditional healers should be trained on how to domesticate natural plant species as some of them has becoming rare or extinct in the surrounding area due to past anthropogenic activities. among others 15 species were documented and reported in our study area as priority species for preservation of stored grains for their medicinal importance uses. these include: b. cicatricose, s. septemtrionalis, t. rhynchocarpum, s. terminale, zanthoxylum sp., r. abyssinicus, p. africana, r. multifidus, t. brevipes, u. massaica, a. alba, c. opulifolium, c. cylindrica, s. aculeastrum and s. mannii. acknowledgements this study was funded by the regional non-governmental organisation (ngo) of albertine rift (arcos) in partnership with association pour la conservation de la nature au rwanda (acnr). we thank the national herbarium of rwanda for their support in species identification and other relevant botanical information, the local government authorities’ of musanze district and traditional healers, members of abadahemuka cooperative. special thanks go to ir. gloriose barebwayire from projet d’appui du développement de l’élevage bovin laitier (padebl/minagri) for excellent technical support in cattle diseases identification, mrs usambyimbabazi madeleine, a student at the national university of rwanda (nur)-botany department for her kind field assistance and mr felix m. waweru (africarice center dar es salaam) for editing and proof-reading. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions n.j.s. was the project leader, m.v., b.e. and b.n. made conceptual contribution, n.m. was involved in data collection and r.i. was responsible for data collection and analysis, writing up the article and project design. references adenkule, j.o., 2007, culture and customs of rwanda, greenhood press, london. alexiades, m.n., 1996, collecting ethnobotanical data: an introduction to basic concepts and techniques, institute of economic botany, the new york botanical garden, new york, viewed 20 october 2013, from http://www.academia.edu/446138/alexiades_m._1996._collecting_ethnobotanical_data association pour la conservation de la nature au rwanda, 2008, biodiversity survey of buhanga relict forest. report, acnr, kigali, rwanda. berkes, f., 1999, in f. berkes (ed.), ‘sacred ecology: traditional ecological knowledge and resource management’, in chapter i: context of traditional ecological knowledge (part i, concept), p. 209, taylor and francis, philadelphia, pa. berkes, f., colding, j. & folke, c., 2000, ‘rediscovery of traditional ecological knowledge as adaptive management’, ecological applications 10(5), 1251–1262, viewed 12 january 2015, from http://www.fws.gov/nativeamerican/pdf/tek-berkes-2000.pdf birdlife international africa partnership e-bulletin, 2008, ‘an african pitta pitta angolensis pays a visit to buhanga relict forest, rwanda’ (16th e-bulletin june 2008), viewed 24 november 2015, from http://www.birdlife.org/sites/default/files/attachments/16th%20ebulletin%20june%202008_final.pdf byavu, n., henrard, c., dubois, m. & malaisse, f., 2000, ‘phytothérapie traditionnelle des bovins dans les élevages de la plaine de la ruzizi’, biotechnology, agronomy, society and environment 4(3), 135–156, viewed 20 december 2012, from http://www.researchgate.net/publication/26392451_phytothrapie_traditionnelle_des_bovins_dans_les_levages_de_la_plaine_de_la_ruzizi charnley, s., paige, a.f. & jones, t.e., 2007, ‘integrating traditional and local ecological knowledge into forest biodiversity conservation in the pacific northwest’, forest ecology and management 246, 14–28. http://dx.doi.org/10.1016/j.foreco.2007.03.047 colding, j., 1998, ‘analysis of hunting options by the use of general food taboos’, ecological modelling 110, 5–17. http://dx.doi.org/10.1016/s0304-3800(98)00038-6 dawit, a. & ahadu, a., 1993, medicinal plants and enigmatic health practices of northern ethiopia, birhanna selam printing enterprise, addis ababa, ethiopia. dessouter, s., 1991, pharmacopée humaine et vétérinaire du rwanda. annales sciences economiques 22, institut national de recherche scientifique, butare, rwanda, musée royal de l’afrique centrale tervuren. edwards, r., 2004, ‘no remedy in sight for herbal ransacks’, new science 181, 10–11. fischer, e. & killmann, d., 2008, illustrated field guide to the plants of nyungwe national park, rwanda. koblenz geographical colloquia, series biogeographical monographs 1: 1–772, germany. gadgil, m., berkes, f. & folke, c., 1993, ‘indigenous knowledge for biodiversity conservation’, ambio 22, 151–156. gerique, a., 2006, an introduction to ethnoecology and ethnobotany theory and methods. integrative assessment and planning methods for sustainable agroforestry in humid and semiarid regions, department of geography, university of giessen, senckenbergstr, germany, viewed 28 october 2013, from https://www.researchgate.net/publication/239591352_an_introduction_to_ethnoecology_and_ethnobotany_theory_and_methods_-_integrative_assessment_and_planning_methods_for_sustainable_agroforestry_in_humid_and_semiarid_regions iraguha, g., 2004, database of rwanda and gps point of buhanga relict forest, in association pour la conservation de la nature au rwanda, 2008, biodiversity survey of buhanga relict forest, report, acnr, kigali, rwanda. johannes, r.e., 1998, ‘the case for data-less marine resource management: examples from tropical nearshore fisheries’, trends in ecology and evolution 13, 243–246. http://dx.doi.org/10.1016/s0169-5347(98)01384-6 kamagaju, l., bizuru, e., minani, v., morandini, r., stévigny, c., ghanen, g. et al., 2013, ‘an ethnobotanical survey of medicinal plants used in rwanda for voluntary depigmentation’, journal of ethnopharmacology 150, 708–717, viewed 26 november 2015, from http://www.sciencedirect.com/science/article/pii/s0378874113006788 kamga-kamdem s.l., 2010, ‘ancestral beliefs and conservation: the case of sacred sites in bandjoun, west cameroon’, in b. verschuuren, r. wild, j.a. mcneely and g. oviedo (eds.), sacred natural sites. conserving nature & culture, pp. 120–128, earthscan ltd. london and washington dc. karangwa, c., 1997, validating traditional knowledge: rwanda. institute of scientific and technological research (irst), general information, butare, rwanda, viewed 28 december 2012, from http://tcdc2.undp.org/gssdacademy/sie/docs/vol10/v10_s3_validknowledge.pdf kayonga, a. & habiyaremye, f.x., 1987, médecine traditionnelle et plantes médicinales rwandaises, contribution aux études ethnobotaniques de la flore rwandaise, préfecture de gisenyi, univ. nat. rwanda centre universitaire de recherche sur la pharmacopée et la médecine traditionnelle, curphametra, inédit, p. 121. kimenyi, a., 1989, kinyarwanda and kirundi names: a semiotic analysis of bantu onomastics, the edwin mellen press, new york. martin, g.j., 1995, ethnobotany: people and plants conservation manuals, chapman and hall, london. mbarubukeye, s., 1991, ‘immunization against east coast fever in rwanda: ticks and tick-borne diseases control’, proceedings of a joint oau, fao and ilrad workshop held in kampala, uganda, the international laboratory for research on animal diseases, kenya, september 12–14, 1991, viewed 02 december 2012, from http://pdf.usaid.gov/pdf_docs/pnabs630.pdf mbarubukeye, s. & niang, a.i., 1996, les usages particuliers des plantes ligneuses agroforestières autochtones du rwanda; utilisation en médecine traditionnelle humaine et vétérinaire, en alimentation humaine, en apiculture et en artisanat, rubilizi. minitere, 2003, national strategy and action plan for the conservation of biodiversity in rwanda, ministry of lands resettlement and environment, republic of rwanda, kigali, p. 44, viewed 23 december 2013, from http://www.vub.ac.be/klimostoolkit/sites/default/files/documents/bd_policy_rwanda.pdf minitere & cgis-nur, 2007, final report on the mapping of rwandese forests, vol. 1. the ministry of lands, environment, forests, water and natural resources (minitere) and the geographic information systems & remote sensing research and training center of the national university of rwanda (cgis – nur). ndimukaga, m., 2009, non timber wetland products and their sustainable use: case of rugezi wetland, rwanda, association pour la conservation de la nature au rwanda and alcoa foundation, kigali, rwanda, report, p. 15. nisr, 2012, population and housing census-november 2012, national institute of statistics of rwanda (nisr)-republic of rwanda, kigali, viewed 14 december 2012, from https://www.statistics.gov.rw/system/files/user_uploads/files/books/2012_census_final_draft.pdf nshimiyimana, j. & mutandwa, e., 2010, ‘seasonal dynamics and distribution of ticks in rwanda: implications for tick control strategy in rwanda’, international journal of animal and veterinary advances 2(1), 21–25, viewed 13 august 2015, from http://maxwellsci.com/print/ijava/v2-21-25.pdf ormsby, a., 2013, ‘analysis of local attitudes toward the sacred groves of meghalaya and karnataka, india’, conservation and society 11, 187–197, viewed 19 november 2015, from http://www.conservationandsociety.org/temp/conservatsoc1121874854658_132906.pdf plumptre, a.j., masozera, m. & vedder, a., 2001, the impact of civil war on the conservation of protected areas in rwanda. washington, dc: biodiversity support program (bsp). poffenberger, m., mcgean, b., ravindranath, n.h. & gadgil, m., 1992, ‘field methods manual, volume ii. community forest economy and use pattern: participatory rural appraisal (pra) methods in south gujarat, india-society for promotion of wasteland development’, in j.a. sayer, c.s. harcourt & n.m. collins (eds.), the conservation atlas of tropical forests: africa, pp. 288, macmillan, basingstoke. rema, 2009, rwanda state of environment and outlook: chap ii. population, health and human settlements, rwanda environment management authority (rema), kigali, viewed 11 november 2015, from http://rema.gov.rw/soe/chap2.php rwangabo, p.c., 1993, la médicine traditionnelle au rwanda, edition karthala, acct, paris. sayer, a.j., harcourt, c.s. & collins, n.m., 1992, ‘the conservation atlas of tropical forests: africa’, in j.a. sayer, c.s. harcourt & n.m collins (eds.), part ii: country studies, p. 104, macmillan publishers ltd., london, united kingdom, viewed 21 december 2014, from https://archive.org/details/conservationatla92sa stewart, k.m., 2003, ‘the african cherry (prunus africana): can lessons be learned from an over-exploited medicinal tree?’, journal of ethnopharmacology 89, 3–13, viewed 11 november 2012, from http://www.sciencedirect.com/science/article/pii/s037887410300299x troupin, g., 1966, etude phytocénotique du parc national de l’akagera et du rwanda oriental, institut national de recherche scientifique (inrs), butare. troupin, g., 1978–1988, flore du rwanda: vols 1-4. musée royal de l’afrique centrale. tervuren and institut national de recherche scientifique, butare. van puyvelde, l., geysen, d., ayobangira, f.x., hakizamungu, e., nshimiyimana, a. & kalisa, a., 1985, ‘screening of medicinal plants of rwanda for acaricidal activity’, journal of ethnopharmacology 13, 209–215. http://dx.doi.org/10.1016/0378-8741(85)90008-x verschuuren, b., wild, r., mcneely, j. & gonzalo, o., 2010, natural sites conserving nature and culture, international union for conservation of nature (iucn), earthscan press, pp. 20–32, london and washington dc. appendix 1 figure 1-a1: (a) photo showing the village surrounding the study area, (b) one of the group discussion with local indigenous, (c) and some important sites and species of buhanga sacred forest including the mysterious water-spring, (d) cave (e) and dracaena steudneri (igihondohondo). figure 2-a1: remarkable places in rwanda. issn 0075-6458 39 koedoe 49/2 (2006) the current status of freshwater molluscs in the kruger national park c.t. wolmarans and k.n. de kock wolmarans, c.t. and k.n. de kock. 2006. the current status of freshwater molluscs in the kruger national park. koedoe 49(2): 39–44. pretoria. issn 0075-6458. surveys regarding the freshwater mollusc population of the kruger national park commenced before 1966 and was repeated in 1995, 2001 and recently in 2006. during this period the park was exposed to several droughts as well as floods, all of which could have had some influence on the various habitats in the park. industrial developments and urbanisation near some of the major rivers outside the park, but which flow through the park, could also have a detrimental effect on the species diversity of the freshwater molluscs. the recent survey in 45 habitats included several dams and rivers. the survey was done to determine the current status of the mollusc species diversity. by comparing it with previous surveys, an attempt was made to establish the overall health of freshwater mollusc habitats. from the results it is clear that there was a drastic decrease in the mollusc species diversity in five of the habitats. although there was a slight change in the species composition in some of the major rivers there was no difference in the number of species collected in these habitats. tarebia granifera, an alien invader mollusc, was found in several rivers and dams. gyraulus costulatus, melanoides victoriae and eupera ferruginea, which could not be found during the survey of 2001, were collected during the current survey. key words: freshwater molluscs, species diversity, rivers, dams, alien invaders. c.t. wolmarans and k.n. de kock, school of environmental sciences and development (zoology), north-west university, potchefstroom campus, p/bag x6001, potchefstroom, 2520 republic of south africa. issn 0075-6458 39 koedoe 49/2 (2006) introduction the largest diversity of freshwater molluscs is usually present in the tropical and subtropical areas of the world (de kock et al. 2002). in south africa, this is especially true for mpumalanga, limpopo and kwazulunatal (de kock & wolmarans 1998). the kruger national park is situated within the first two provinces mentioned. various species of freshwater snails act as intermediate hosts for trematode parasites (de kock & wolmarans 1998) while they are also known for their ability to act as indicators for water health (schoonbee 1963). the poor ability of these organisms to migrate, makes them vulnerable to changes in their habitats, such as global warming, pollution or prolonged periods of drought. although the influence of some of these factors like global warming, is difficult to determine over the short to medium term, the influence of other factors like environmental contamination, may be more evident over a shorter period of time. the degree of these influences on molluscs may also be influenced by a stream, river, small pond or dam. although the kruger national park is recognised worldwide for its conservation efforts, it is not isolated from the factors mentioned above. a number of rivers in the park, including the letaba, olifants, crocodile, sand and sabie, meander through adjacent, highly populated and industrialised areas and is it possible that the changes in the water quality occurring in these areas have an effect on the freshwater biodiversity in the park. molluscs present in isolated habitats may, however, escape such changes. the aim of this investigation was to determine the current status of freshwater molluscs in the park and to compare this with a survey done in 2001. wolmarans.indd 39 2006/10/15 10:59:49 pm koedoe 49/2 (2006) 40 issn 0075-6458 methods and materials this study concentrated mainly on water bodies that had been surveyed for molluscs during the 2001 collection. a few habitats, namely the mtombeni spruit, rabelais dam (does not exist any more), lower sabie rest camp, tsende drift, skukuza fish pond and jones’s dam were not surveyed. habitats surveyed for the first time included the mpanamana dam, timbavati river below the piet grobler dam wall, sabie river below the lower sabie dam wall, crocodile river, mbyamiti river (weir), and the letaba river below the engelhard dam wall. a total of 45 habitats were surveyed during 2006. all of these habitat types were screened for molluscs by means of aluminium mollusc scoops. although a quantitative survey of molluscs was not intended, the same time (man hours) was spent at each of these habitats as in previous surveys in order to recover as many species as possible. the molluscs collected at each habitat were largely full-grown specimens and were transferred to 50 ml glass screw-capped bottles and fixed in 70 % ethanol. each bottle was provided with a label containing information on date of collection, locality, and collection number. the specimens were identified by means of a stereo microscope and were then added to the national freshwater snail collection (nfsc) located at the north-west university, potchefstroom campus. results the results of this investigation are presented in table 1. from this it is evident that 11 of the habitats, namely the crocodile river, mpanamana dam, nhlanganzwane dam, ngotso dam, letaba river below the engelhard dam, olifants river, lower sabie dam, the sabie river below the lower sabie dam, the mbyamiti river (weir), luvuvhu river and leeupan were not surveyed in 2001. this was mainly due to a lack of suitable habitats at that time and the fact that some of the habitats were situated in concession areas. of the 45 habitats surveyed in 2006, no molluscs were found in 12 of the habitats, i.e., the mpanamana, nhlanganzwane and ngotso dams (not surveyed previously), the nsemani dam, shisa spruit, vervoer dam, manzimahle dam, sunset dam, sable dam, silolweni dam, leeupan and the luvuvhu river. comparing the 34 habitats surveyed in both 2001 and 2006, it is evident that only the piet grobler dam had no molluscs in 2001. contrary to this, no molluscs were found in seven of these habitats in 2006, namely the shisa spruit, the nsemani, vervoer, manzimahle, sunset, sable and silolweni dams, four species, not found in 2001 were collected in 2006. tarebia granifera (an alien invader snail) was found in 12 habitats including several rivers and dams (table 1). according to the records of the nfsc this species was not previously found in the kruger national park. the other three species, namely gyraulus costulatus, eupera ferruginea and melanoides victoriae were found in numerous habitats during the first extensive survey done in 1964. eupera ferruginea was also found during the 1995 survey. lanistes ovum and pisidium pirothi, collected in 2001, were not found during the recent survey. in comparison with the 2001 survey, there was a drastic decrease in the species diversity evident in five of the habitats surveyed in 2006. these include the matjulu and shisa spruits, the engelhard dam, the nwanetsi water hole and the gudzani and sirheni dams. whether the presence of the invader t. granifera played a role in the decrease in three of these habitats, namely the engelhard dam, nwanetsi water hole and the gudzani dam is unknown as it was not found in the remaining three habitats. a drastic increase in species diversity was not experienced in any of the habitats surveyed in 2006. the major rivers (timbavati, letaba, and sabie), showed no significant difference in the mollusc species diversity between the two surveys. discussion direct comparisons of the species diversity recorded during the respective surveys could lead to scientifically unfounded conclusions. aspects that should be taken into account, include amongst others, the number of collections made in a habitat, the exact collection site, the time of the year when the collection was made, as well as the rainfall patterns prior to each survey (de kock & wolmarans 1998). in spite of this, it still seems possible to outline specific trends. although there was no obvious differences in the number of wolmarans.indd 40 2006/10/15 10:59:49 pm issn 0075-6458 41 koedoe 49/2 (2006) table 1 molluscs found in water bodies of the kruger national park surveyed in 2001 and 2006 rainfall monitoring station collection site grid reference species collected in 2006 species collected in 2001 berg-en-dal matjulu spruit 25º21’s; 31º31’e bulinus globosus bulinus globosus biomphalaria pfeifferi bulinus forskalii lymnaea columella biomphalaria pfeifferi lymnaea natalensis lymnaea columella aplexa marmorata burnupia caffra melanoides tuberculata mlambane spruit 25º21's; 31º31'e bulinus globosus bulinus forskalii biomphalaria pfeifferi aplexa marmorata crocodile bridge gezantombi dam 25º20's; 31º52'e bulinus forskalii bulinus forskalii cleopatra ferruginea lymnaea columella aplexa marmorata burnupia caffra crocodile river 25º21's; 31º54'e tarebia granifera (not surveyed) mpanamana dam 25º17's; 32º0'e no molluscs found (not surveyed) nhlanganzwane dam 25º14's; 31º57'e no molluscs found (not surveyed) houtboschrand timbavati river 24º31's; 31º38'e bulinus globosus bulinus africanus lymnaea natalensis lymnaea natalensis gyraulus costulatus burnupia caffra ngotso dam 24º11's; 31º43'e no molluscs found (not surveyed) letaba letaba river 23º45's; 31º22'e lymnaea collumella bulinus forskalii physa acuta physa acuta aplexa marmorata melanoides tuberculata melanoides tuberculata engelhard dam 23º50's; 31º28'e physa acuta bulinus globosus aplexa marmorata bulinus tropicus tarebia granifera bulinus forskalii biomphalaria pfeifferi physa acuta melanoides tuberculata corbicula fluminalis pisidium pirothi letaba river below the 23º31's; 31º39'e lymnaea columella (not surveyed) engelhard dam wall physa acuta melanoides tuberculata melanoides victoriae tarebia granifera corbicula fluminalis lower sabie olifants river 24º31's; 31º41'e physa acuta (not surveyed) mpondo dam 25º12's; 31º43'e bulinus forskalii bulinus forskalii tarebia granifera lymnaea natalensis ceratophallus natalensis lymnaea collumella bulinus forskalii spathopsis petersi wolmarans.indd 41 2006/10/15 10:59:50 pm koedoe 49/2 (2006) 42 issn 0075-6458 lower-sabie dam 25º07's; 31º56'e tarebia granifera ( not surveyed ) sabie river 25º07's; 31º55'e bulinus forskalii physa acuta burnupia cf. caffra apexa marmorata sabie river below 25º07's; 31º55'e tarebia granifera ( not surveyed ) lower sabie dam wall corbicula fluminalis eupera ferruginea mooiplaas pioniers dam 23º31's; 31º23'e bulinus globosus bulinus globosus biomphalaria pfeifferi bulinus forskalii lymnaea natalensis biomphalaria pfeifferi grootvlei dam 23º20's; 31º30'e bulinus tropicus spathopsis sp nwanetsi nwanetsi water hole 24º28's; 31º58'e biomphalaria pfeifferi bulinus globosus tarebia granifera bulinus tropicus bulinus forskalii biomphalaria pfeifferi ceratophallus natalensis burnupia caffra spathopsis wahlbergi spathopsis petersi corbicula fluminalis melanoides tuberculata oxyloma striata gudzani dam 24º22's; 31º55'e bulinus globosus bulinus globosus biomphalaria pfeifferi bulinus tropicus lymnaea natalensis biomphalaria pfeifferi melanoides tuberculata lymnaea natalensis tarebia granifera ceratophallus natalensis melanoides tuberculata spathopsis petersi corbicula fluminalis pretoriuskop mbyamiti river 25º13's; 31º34'e bulinus globosus bulinus globosus biomphalaria pfeifferi lymnaea columella aplexa marmorata mbyamiti river (weir) 25º15's; 31º37'e bulinus globosus ( not surveyed ) biomphalaria pfeifferi tarebia granifera mestel dam 25º06's; 31º12'e bulinus globosus bulinus globosus melanoides tuberculata lymnaea columella tarebia granifera aplexa marmorata shitlehave dam 25º08's; 31º19'e bulinus forskalii biomphalaria pfeifferi menanoides tuberculata punda maria klopperfontein dam 22º37's; 31º10'e aplexa marmorata bulinus tropicus bulinus forskalii satara nkaya pan 24º31's; 31º45'e bulinus forskalii bulinus forskalii piet grobler dam 24º31's; 31º38'e bulinus globosus no molluscs found tarebia granifera timbavati river below piet grobler dam wall 24º30's; 31º38'e tarebia granifera bulinus forskalii nsemani dam no molluscs found bulinus forskalii burnupia caffra table 1 (continued) wolmarans.indd 42 2006/10/15 10:59:50 pm issn 0075-6458 43 koedoe 49/2 (2006) shingwedzi krapkuil dam 23º14's; 31º18'e bulinus tropicus burnupia caffra sirheni dam 22º56's; 31º14'e bulinus globosus bulinus globosus biomphalaria pfeifferi biomphalaria pfeifferi lymnaea natalensis lymnaea columella lanistes ovum shisa spruit 22º57's; 31º15'e no molluscs found cleopatra ferruginea melanoides tuberculata spathopsis petersi corbicula fluminalis kanniedood dam 23º08's; 31º27'e spathopsis wahlbergi bulinus tropicus spathopsis petersi bulinus forskalii lymnaea natalensis skukuza olifantdrinkgat 24º53's; 31º44'e bulinus forskalii bulinus forskalii vervoer dam 25º07's; 31º29'e no molluscs found lymnaea columella aplexa marmorata manzimahle dam 24º53's; 31º43'e no molluscs found bulinus forskalii sunset dam 25º06's; 31º54'e no molluscs found aplexa marmorata phalaborwa sable dam 23º56's; 31º14'e no molluscs found bulinus tropicus bulinus forskalii orpen dam 24º47's; 31º53'e aplexa marmorata lymnaea columella ceratophallus natalensis mazithi dam 24º43's; 31º50'e bulinus forskalii bulinus forskalii kumana dam 24º37's; 31º47'e bulinus forskalii bulinus forskalii nwaswitsontso river 24º37's; 31º47'e bulinus forskalii bulinus forskalii silolweni dam 24º29's; 31º50'e no molluscs found bulinus forskalii burnupia caffra leeupan 24º50's; 31º48'e no molluscs found (not surveyed) pafuri luvuvhu river 22º26's; 31º13'e no molluscs found (not surveyed) species found in the larger rivers (timbavati, letaba and sabie rivers) when comparing the 2001 and 2006 surveys, slight changes in the species diversity were evident (table 1). this may be an indication that any changes in the water quality of these rivers before entering the park, still did not have a noticeable effect on the mollusc species diversity in these habitats. the presence of physa acuta (also known as the sewage snail, appleton 2003) the second-most successful alien invader mollusc species in south africa (de kock et al. 1989) in the olifants river, may be an indication of the abnormal conditions in this river. the unfavourable ph of the water due to old mine activities, the pollution contributed by various industries in the witbank area, and poor farming practises where overgrazing causes top soil to wash into the river, all create a situation where p. acuta may be table 1 (continued) wolmarans.indd 43 2006/10/15 10:59:50 pm koedoe 49/2 (2006) 44 issn 0075-6458 one of the few species able to survive. the occurrence of a number of floods in this river since 1995 should have enhanced the overall water quality but did not contribute much to the increase in species diversity of molluscs recovered. the fact that there was a decrease in species diversity of molluscs in a number of habitats and that no snails were found in several of the dams also surveyed in 2001 (table 1) could either be ascribed to a decrease in the overall number of molluscs in these dams due to a lack of suitable habitats or to environmental influences like specific rainfall patterns directly before the survey. as these dams are not connected to rivers arising outside the park, it seems highly improbable that industrial pollution could play a significant role in this phenomenon. the presence of t. granifera in a number of habitats in which it was not previously found, could be reason for great concern especially if its presence would have a detrimental effect on the survival of other freshwater fauna. this is an aspect that should, in our opinion, be investigated in great depth. although the increase in species diversity found in 2001 as compared to the survey of 1995 was ascribed to the high rainfall average between 1995 and 2001 (de kock et al. 2002) the recent survey was done shortly after a period of high rainfall. it is, however, important to keep in mind that the actual period between the last significant rainfall and the time of the survey may play a role in the species diversity and it is therefore premature to conclude that there was a decrease in the general species diversity as far as the freshwater molluscs are concerned. therefore more surveys should be done to clarify the reasons for the decrease in species diversity recently encountered. conclusions comparisons between the respective habitats surveyed in 2001 and 2006 revealed slight differences in the mollusc species diversity of the letaba, timbavati and sabie rivers but no differences in the number of species collected. although there was a decrease in the mollusc species diversity in five of the habitats during the 2006 survey and no molluscs were found in another 12 habitats as compared to only one during the 2001 survey, it would seem premature to conclude that there was an overall decrease in the species diversity. the occurrence of t. granifera in 12 of the habitats is a matter of concern especially because of its ability to invade a variety of habitats in a relatively short period, a situation that should be monitored very closely in the future. acknowledgements the authors wish to thank the following persons and institutions for their assistance: the national parks board for permission to conduct the research, as well as for their financial support; mr hendrik sithole who was instrumental in assisting us with the necessary arrangements for the survey; the research focus area, environmental science and sustainable management of the potchefstroom campus of the north-west university for financial support. references appleton, c.c. 2003. alien and invasive fresh water gastropoda in south africa. african journal of aquatic science 28: 69–81. de kock, k.n., p.h. joubert & s.j. pretorius. 1989. geographical distribution and habitat preferences of the invader freshwater snail species lymnaea columella (mollusca: gastropoda) in south africa. onderstepoort journal of veterinary research 56: 271–275. de kock, k.n. & c.t. wolmarans. 1998. a re-evaluation of the occurrence of freshwater molluscs in the kruger national park. koedoe 41(1): 1–8. de kock, k.n., c.t. wolmarans & l.h. du preez. 2002. freshwater mollusc diversity in the kruger national park: a comparison between a period of prolonged drought and a period of exceptionally high rainfall. koedoe 45(2): 1–11. schoonbee, h.j. 1963. pollution studies in the umgeni river, part 3: a study of the biological and chemical effects of effluent discharges in the pinetown-new germany industrial complex. c.s.i.r. spec. rep., w25, 1–34. wolmarans.indd 44 2006/10/15 10:59:50 pm filelist convert a pdf file! koedoe 19: 181-184 (1976) lewensloop van 'n rooihartbeesbevolking alcelaphus buselaphus caama in die bontebok nasionale park p t van der walt ajdeling n atuurbewaring privaatsak x404 skukuw 1350 g de graaff nasionale parkeraad posbus 787 pretoria 0001 lj van zyl bontebok nasionale park posbus 149 swellendam 6740 abstract the life history of a re-introduced red hartebeest alcelaphus buselaphus caama population in the bontebok national park, near swellendam, is discussed and illustrated. inleiding die rooihartbees alcelaphus buselaphus caama kan beskou word as een van suidelike afrika se mees aanpasbare boksoorte. historiese verspreidingsrekords, soos weergegee deur du plessis (1969), dui daarop dat die bok vroeer voorgekom het oor die hele kaapprovinsie, oranje-vrystaat, suidwes-afrika, botswana en die westelike dele van natal en transvaal. wyd uiteenlopende plantegroei en klimaatstreke kom in hierdie dele voor wat wissel vanaf 'n droe sanderige woestyn tot sub-humiede grasveld en boomryke gebiede. suid van die oranjerivier word daar vandag slegs enkele geisoleerde populasies aangetref in bewaarde gebiede. tot onlangs was die bontebok nasionale park (bnp), digby swellendam, een van hierdie gebiede. hierdie verslag dien as 'n dokumentering van die lewensloop van 'n rooihartbeestrop vanaf hul hervestiging in die bn p (april 1960) tot die verwydering daarvan uit die bnp (1975.05.28). die redes vir die inbring van diersoorte in nasionale parke asook 'n beskrywing van die geskiedkundige agtergrond van die bnp, word in besonderhede deur bateman (1961), penzhorn (1971) en van rensburg (1975). 181 resultate en bespreking 'n bykans volledige beeld van die bevolkingsamestelling van die rooihartbeesindiebnpoordiejare 1961-1975,wordinfig.l uitgebeeld. ...j < ? geslagsverhouding by benadering • yr~ktes 16 ~manlik 14 • vroulik o kalwers 12 19 !z 8 < < 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 jare fig. i. lewenshistogram van die rooihartbeesbevolking in die bontebok nasionale park vanaf 1961-1975. gelyktydig met die hervestiging van bontebokke damaliscus dorcas dorcas in die bestaande bn p gedurende april 1960, het die rooihartbees sy beslag in die park gekry toe ses kalwers vanuit die kalahari-gemsbok nasionale park losgelaat is. die aanvangs-geslagverhouding was 20'0':1<:(. . vanuit fig. 1 is dit duidelik dat 'n algemene vermeerdering onder die vroulike diere oor die jare ingetree het. dit het egter nie volgens 'n spesifieke patroon geskied nie wat daarop dui dat die hartbeestrop deurentyd heelwat steurnisse ondervind het wat inderdaad die geval was. die jaarlikse tropontleidings in fig. 1 toon tot 'n groot mate aan dat die frekwensie en intensiteit van vrektes op sy hoogste is wanneer daar 'n geslagsverhouding van 1: 1 heers . dominansiegevegte onder die bulle kom dan dikwels voor wat lei tot vrektes veral onder die jonger diere. feitlik al die vrektes wat aangetoon word, kan hieraan toegeskryf word. die trop het vanaf 1962-1975 'n gemiddelde kalfpersentasie van 40% gehandhaaf. dit stem min of meer ooreen met van die ander boksoorte wat in die bnp aangetref word, soos die eland taurotragus oryx (41%) en die bontebok (54%). ten spyte hiervan kon die rooihartbees nie in dieselfde mater as die eland, bontebok en springbok antidorcas marsupialis oor ' n periode van 15 jaar in 'n lewensvatbare populasie in die bnp ontwikkel nie. 182 faktore wat moontlik hiertoe aanleiding kon gee is 'n beperkte lewensruimte (oppervlakte van bnp is 2786 hal, 'n te klein aantal van die diere wat aanvanklik hervestig is, en die heersende toestand van die plantegroei in die park. die huidige oorwegende onsmaaklike struikgemeenskap in die bnp, het volgens getuienis (barrow 1801; lichtenstein 1811) vroeer bestaan as 'n grasveld. die retererende graskomponent in die bnp as gevolg van boerderyaktiwiteite in die verlede, kan moontlik tot gevolg gehad het dat die bnp tans meer as 'n marginale verspreidingsgebied beskou kan word vir die rooihartbees. dit volg daaruit dat die dier primer 'n grasbenutter is. die vroeere seisoenale benutting deur rooihartbeeste van bepaalde gebiede in die ruensveld en omgewing waarin die bnp gelee is, het enige voedingstekorte wat mag ontstaan uitgeskakel. in die bnp was die diere beperk tot slegs 'n baie klein gedeelte van hul vroeere verspreidingsgebied. dit tesame met die reeds genoemde nie-wenslike plantontwikkelingstendense in die park, het waarskynlik aanleiding gegee tot 'n proteientekort by die diere wat weer sekondere simptome soos 'n lae reproduksie en 'n afname in massa tot gevolg gehad het. hierdie diere was in grootte en massa heelwat kleiner as die rooihartbeeste in die kalahari-gemsbok nasionale park, wat nog na willekeur rondbeweeg. al die bogenoemde beweegredes het gedurende maart 1975 daartoe gelei dat die raad van kuratore vir nasionale parke besluit het om die rooihartbeestrop (40', 49, 2 kalwers) uit die bnp te verwyder. as gevolg van beperkte natuurlike hulpbronne in die bnp, sal prioriteit voortaan verleen word aan die bewaring van bepaalde diersoorte, soos die bontebok, vaalribbok pelea capreolus en verskeie kleinere antiloopsoorte soos veral die kaapse grysbok raphicerus melanotis. optimum bestaanstoestande sal sover moontlik vir hulle geskep word, veral met behulp van beproefde veldbestuurspraktyke. verwysings barrow, j. 1801 . an account of travels into the interior of southern africa in the years 1797 and 1798. 1. london: t. cadell jun. 8 w. davies. bateman, j. a. 1961. the mammals occurring in the bredasdorp and swellendam districts, c. p., since european settlement. koedoe 4: 78-100. du plessis, s. f. 1969. the past and present geographical distribution of the perissodactyla and artiodactyla in southern africa. m.sc. thesis, univ. of pretoria. lichtenstein, h. 1811. reisen in sudlichen afrika in der jahren 1803, 1804, 1805 und 1806. i. berlin: c. g. liideritz. penzhorn, b. l. 1971. a summary of the re-introduction of ungulates into south african national parks (to 31 december 1970). koedoe 14: 145-159. van rensburg. a. p. j. 1975. die geskiedenis van die nasionale bontebokpark, swellendam. koedoe 18: 165-190. 183 page 1 page 2 page 3 article information authors: george j. chirima1,2,3 norman owen-smith1 barend f.n. erasmus1 affiliations: 1school of animal, plant and environmental sciences, university of the witwatersrand, south africa2geoinformatics department, agricultural research council, south africa 3south african environmental observation network, pretoria, south africa correspondence to: george chirima postal address: private bag x79, 600 belvedere street, pretoria 0001, south africa dates: received: 08 june 2010 accepted: 13 apr. 2012 published: 24 july 2012 how to cite this article: chirima, g.j., owen-smith, n. & erasmus, b.f.n., 2012, ‘changing distributions of larger ungulates in the kruger national park from ecological aerial survey data’, koedoe 54(1), art. #1009, 11 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1009 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. changing distributions of larger ungulates in the kruger national park from ecological aerial survey data in this original research... open access • abstract • introduction • methods    • study area    • data source    • data analysis       • distribution range estimation       • regional distribution range change       • habitat associations       • comparison with earlier maps • results    • impala    • buffalo    • zebra    • wildebeest    • kudu    • giraffe    • waterbuck    • warthog    • sable antelope    • tsessebe    • eland    • roan antelope • trustworthiness • discussion • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ documenting current species distribution patterns and their association with habitat types is important as a basis for assessing future range shifts in response to climate change or other influences. we used the adaptive local convex hull (a-locoh) method to map distribution ranges of 12 ungulate species within the kruger national park (knp) based on locations recorded during aerial surveys (1980–1993). we used log-linear models to identify changes in regional distribution patterns and chi-square tests to determine shifts in habitat occupation over this period. we compared observed patterns with earlier, more subjectively derived distribution maps for these species. zebra, wildebeest and giraffe distributions shifted towards the far northern section of the knp, whilst buffalo and kudu showed proportional declines in the north. sable antelope distribution contracted most in the north, whilst tsessebe, eland and roan antelope distributions showed no shifts. warthog and waterbuck contracted in the central and northern regions, respectively. the distribution of impala did not change. compared with earlier distributions, impala, zebra, buffalo, warthog and waterbuck had become less strongly concentrated along rivers. wildebeest, zebra, sable antelope and tsessebe had become less prevalent in localities west of the central region. concerning habitat occupation, the majority of grazers showed a concentration on basaltic substrates, whilst sable antelope favoured mopane-dominated woodland and sour bushveld on granite. buffalo showed no strong preference for any habitats and waterbuck were concentrated along rivers. although widespread, impala were absent from sections of mopane shrubveld and sandveld. kudu and giraffe were widespread through most habitats, but with a lesser prevalence in northern mopane-dominated habitats. documented distribution shifts appeared to be related to the completion of the western boundary fence and widened provision of surface water within the park. conservation implications: the objectively recorded distribution patterns provide a foundation for assessing future changes in distribution that may take place in response to climatic shifts or other influences. introduction top ↑ populations of certain large ungulates have declined within the kruger national park (knp) as well as elsewhere in africa (craigie et al. 2010; ogutu & owen-smith 2003). the challenge is to establish the causal influences underlying these population changes and, in particular, to distinguish extrinsic drivers such as climatic shifts from intrinsic factors such as fencing, water distribution and fire pattern, which can be managed more directly. changing distribution patterns provide potentially helpful clues to the nature of mechanisms negatively operating on a population within a conservation area such as the knp (gaston 1990, 2003; lawton 1993). climatic influences should be reflected by distributional shifts along gradients in temperature or rainfall and underlain by consequent changes in habitat suitability. more local causes would be indicated by disparate changes in different regions of the species distribution.the foundational requirement is for prevailing distribution ranges to be established as sufficiently rigorous for future changes to be identified with confidence. pienaar (1963) mapped the distribution patterns of all of the larger mammal species within the knp from rangers’ observations over the preceding five years, road strip counts (1956–1961) and aerial counts covering the central section in 1960 and 1962. whilst historically useful, these maps are obviously subjective and vague in the time period that they represent. between 1977 and 1995, annual ecological aerial surveys were conducted recording the locations of all animals seen, which comprehensively covered almost the entire knp from 1980 to 1993. count totals for the larger ungulate species, partitioned amongst eight census compartments, were summarised in annual reports (e.g. viljoen 1993). however, distribution patterns revealed at a finer scale by the actual animal locations recorded have not been synthesised. hence, our starting aim was to document the recent distribution patterns of the larger ungulate species for comparison with the earlier patterns depicted by pienaar (1963) and as a basis for further monitoring. subsequent to the time period covered by pienaar (1963), various changes potentially influencing the distribution of the larger herbivore species have occurred. the southern and western borders of the knp became completely fenced in 1961, blocking the movement of animals between the park and adjoining private nature reserves, followed by the completion of a fence along the eastern border with mozambique in 1976 (joubert 2007a, 2007b). between 1965 and 1975, the provision of artificial water sources in the form of earth dams, weirs on seasonal streams and drinking troughs supplied from boreholes was greatly expanded (grant et al. 2002). after 1977, very little of the park area remained more than 5 km from the nearest perennial water source (redfern et al. 2003). fire policy has also changed in various ways, potentially influencing vegetation features and hence habitat conditions for large herbivores (van wilgen et al. 2004). the elephant population grew from a little over 1000 animals in 1962 to a total of over 7000 by 1968, after which annual removals curtailed further increase until this culling was suspended in 1995 (whyte et al. 1999). persistently low rainfall conditions prevailed in the late 1960s and then from 1982 through 1995, including exceptionally severe droughts in 1982–1983 and 1991–1992 (owen-smith & ogutu 2003). this rainfall variation was associated with substantial changes in the abundance of certain ungulate species, including marked declines in populations of some of the less common antelope species (mills, biggs & whyte 1995; ogutu & owen-smith 2003; owen-smith, mason & ogutu 2005; owen-smith & mills 2006). with these changes in mind, our specific objectives were: • to establish shifts in distribution accompanying the population changes of larger ungulates over the period spanned by the annual aerial surveys, in particular comparing the period prior to 1986 with that thereafter, when declines of less common species took place in association with persistently low rainfall conditions. • to compare recent distribution patterns of these ungulates with those around 1960, as mapped by pienaar (1963). • to relate distribution patterns to the habitat types preferred or avoided by these species. • to assess how changes in surface water availability might have affected regional concentrations. methods top ↑ study area gertenbach (1983) and venter, scholes and eckhardt (2003) provided detailed descriptions of climate, vegetation and geology within the nearly 20 000 km2 area of the knp. briefly, mean annual rainfall declines from about 750 mm in the south-west to around 400 mm in parts of the north, falling mostly during the summer wet season between october and march. in the southern half, the vegetation is predominantly knob thorn (acacia nigrescens) and marula (sclerocarya birrea) savannah on basalt substrates in the east, and combretum spp. savannah on granitic substrates in the west. mopane (colophospermum mopane) dominates the woody vegetation in the northern half on both substrates. data source the ecological aerial surveys were conducted between may and august when visibility conditions are best (viljoen 1993). hence, our distributional analysis represents dry season conditions only. four observers, besides the pilot and recorder, counted all animals seen using transects spaced 800 m apart covering successive blocks. from 1980 through 1986, animal locations were mapped within a 2 km × 2 km grid (joubert 1983). after 1986, a palmtop computer coupled with a gps unit recorded the coordinates of animals seen (viljoen & retief 1994). hence, the positional inaccuracy could be up to 2 km prior to 1987 but within 0.8 km from 1987 onwards (viljoen & retief 1994). the area north of punda maria was not consistently covered and hence was excluded from our analysis. the hilly region in the extreme south-west was also less reliably covered. the species considered were, in order of abundance, impala (aepyceros melampus), plains zebra (equus quagga), african buffalo (syncerus caffer), blue wildebeest (connochaetes taurinus), greater kudu (tragelaphus strepsiceros), giraffe (giraffa camelopardalis), common waterbuck (kobus ellipsiprymnus), warthog (phacochoerus africanus), sable antelope (hippotragus niger), tsessebe (damaliscus lunatus), eland (taurotragus oryx) and roan antelope (hippotragus equinus). data analysis distribution range estimation we used the adaptive local convex hull (a-locoh) method, developed for home range analysis (getz & wilmers 2004; getz et al. 2007), to assess the distribution ranges. this method is more sensitive to gaps in occurrence, and less influenced by outliers, than parametric kernel methods. we generated point shape files of the geographical locations of animal herds in arc map 10.0, (environmental systems research institute 2010) and applied the a-locoh spatial analyst tool (getz et al. 2007). local minimum convex hulls were constructed from a variable number (k) of neighbouring points to a location or root point. initially, we fixed the value of k at 3 and the value of the distance (a) from the root point at 1 km. we then plotted the area of the estimated distribution range versus increasing values of a to find the point where the area began to level off (i.e. the minimum spurious hole covering [mshc]) value of a (getz et al. 2007). with a fixed at this value, we varied k to find its mshc value. thereafter, we used these joint mshc values of a and k to construct the final distribution ranges. the union of hulls moving up from the smallest were used to construct isopleths (getz et al. 2007). the 0.99 isopleth was used to define range limits for the most common species, because the 0.95 isopleth excluded a substantial number of animals. for remaining species, ranges limits were mapped using the 0.95 isopleth. the 0.75 isopleth was used consistently to demarcate core regions where most of the population was concentrated. we excluded records of single animals generally representing solitary males, which were potentially found outside the distribution range of breeding herds. for each species, location records were aggregated for the 7-year periods 1980–1986 (including the 1982–1983 drought but before population declines by rarer species were initiated) and 1987–1993 (covering the time when population declines by less common species were under way). for impala, the total number of location records exceeded computing capacity and so were processed in separate batches representing the northern and southern halves of the knp, later merged for display purposes. regional distribution range change to investigate regional and temporal disparities in distribution patterns, we divided the knp into four sections, separated by major rivers (figure 1). the relative presence of herds consisting of two or more individuals in 5 km × 5 km cells in these sections was assessed separately for the pre-1987 and post-1986 periods. cells that contained multiple records were treated as a single ’presence‘. we used log-linear models because both predictor and response variables were categorical, judging goodness-of-fit by the likelihood ratio statistic (l2) (agresti 1996). to establish whether the distribution of a species had changed significantly between the two periods, we dropped the three-way or two-way interaction terms from models incorporating region and period as factors, checking whether the omission brought about a significant reduction in model fit. we examined z-scores for reduced models to establish which interactions contributed the most to the lack of fit when their effects were removed and to establish whether the effect was positive or negative (christensen 1997; knoke & burke 1980). figure 1: distribution of major habitat types in the kruger national park. habitat associations gertenbach (1983) described 35 landscape types representing 15 major ecological units defined by vegetation structure and composition. we reduced his 15 ecological units to nine distinct habitat types (see online appendix) and grouped four minor habitats into the category ’other‘ (figure 1). to assess relative habitat preferences, we compared the proportion of animals of each ungulate species mapped within each habitat type with the proportional availability of these habitats. availability was estimated as the proportion of total area of the park covered. habitat preference or avoidance was assessed separately for the pre-1987 and post-1986 periods. we interpreted a habitat type as being preferred if the proportional occupation exceeded twice the relative availability and avoided if this proportion was less than 0.5 of the relative availability. for species that exhibited distributional changes, we also assessed how this shift was reflected in habitat occupation. changes in the proportion of occupied 5 km × 5 km cells per habitat type were compared between the pre-1987 versus the post-1986 periods, supported by chi-square tests. comparison with earlier maps because the distribution patterns presented by pienaar (1963) were somewhat subjective, they cannot be compared rigorously with those documented during the annual aerial surveys. hence, we merely draw attention to the notable differences that seem apparent. results top ↑ impala the overall distribution of impala expanded slightly between the pre-1987 and post-1986 periods (l2 = 9.171, df = 3, p = 0.027), without any significant change in regional proportions (figure 2a). impala occupied all habitats, showing a notable preference only for delagoa thorn thicket (table 1). however, occupation of sour bushveld by impala did increase significantly, by roughly 50%, after 1986 (χ2 = 12.136, df = 1, p = 0.001). figure 2: changes in the overall proportion of 5 km × 5 km cells (n = 804) occupied between the two periods (thick black lines) and in the proportional distribution of occupied cells among the four sections of the kruger national park (stacked bars), where (a) impala, (b) buffalo, (c) zebra, (d) wildebeest, (e) giraffe, (f) kudu, (g) warthog, (h) waterbuck, (i) sable antelope, (j) eland, (k) tsessebe and (l) roan antelope. table 1: proportion of animal sightings recorded in 0.99 probability isopleths for common species within each habitat type, expressed as a percentage relative to the proportional extent of these habitats in the kruger national park. in addition, although impala showed no obvious change in distribution pattern between the two periods, they remained absent from some regions of the knp. gaps were evident in sections of the east and in the extreme south-west (figure 3). impala were more common in the southern half of the knp than in the north, noting that distribution patterns must be assessed independently between the two halves of the knp. concentration areas were associated with perennial and seasonal rivers on the western granitic region of the northern part, but more widespread away from rivers in the wetter southern part. the distribution of impala mapped by pienaar (1963) showed a tighter concentration along rivers, particularly in the northern half of the knp, than was apparent after 1980. figure 3: distribution range of impala comparing the periods (a) 1980–1986 and (b) 1987–1993. coancentration zones enclosing 75% of the population (dark shade) are distinguished from the 95% or 99% range limits (grey shade). buffalo buffalo were likewise distributed through most of the knp, with local concentrations near rivers during the dry season. however, their occurrence in the south-western region appeared somewhat more patchy prior to 1987 than subsequently (figure 4). the buffalo distribution mapped by pienaar (1963) showed a strong concentration along rivers in the dry season and an expansion over most of the park in the wet season. relatively few buffalo were found in the south-west region prior to 1963. figure 4: distribution range of buffalo comparing the periods (a) 1980–1986 and (b) 1987–1993. concentration zones enclosing 75% of the population (dark shade) are distinguished from the 95% or 99% range limits (grey shade). the change in buffalo distribution between the two periods differed between sections (l2 = 13.276, df = 3, p = 0.004). buffalo became distributed more widely in the north (z = 2.879, p = 0.004) and south (z = 6.490, p = 0.001) at the expense of the central and far north sections (figure 2b). buffalo occupied all habitat types and showed a marked increase in their presence in sour bushveld after 1986 (χ2 = 9.733, df = 1, p = 0.002). zebra zebra occurred throughout the knp, except for gaps south-west of the sabie river and near the crocodile river (figure 5). pienaar’s (1963) map shows a basically similar distribution, except for his demarcation of the south-eastern region of the central section as wet season range. figure 5: distribution range of zebra comparing the periods (a) 1980–1986 and (b) 1987–1993. concentration zones enclosing 75% of the population (dark shade) are distinguished from the 95% or 99% range limits (grey shade). dropping two-way interactions from models incorporating period and region as factors brought about a significant reduction to model fit (l2 = 96.103, df = 10, p = 0.001), suggesting that zebra distribution changed between the two periods, with most range expansion occurring in the far north (z = 2.429, p = 0.027) (figure 2c). zebra occupied all habitats, but tended to avoid thorn thickets near rivers and sour bushveld, although their presence in the latter habit increased slightly, but significantly, after 1986 (table 1; χ2 = 5.595, df = 1, p = 0.032). wildebeest wildebeest were concentrated in the central section and distributed more patchily in the south as well as north of the olifants river (figure 6). pienaar’s (1963) map shows additional dry season concentrations along the western border of the central and south sections. the eastern region of the central section was shown formerly as wet season range. a pocket in the far north-west was no longer present after 1980. figure 6: distribution range of wildebeest comparing the periods (a) 1980–1986 and (b) 1987–1993. concentration zones enclosing 75% of the population (dark shade) are distinguished from the 95% or 99% range limits (grey shade). although the proportion of wildebeest in broader regions of the knp appeared to have changed little, dropping two-way interactions incorporating period and region as factors brought about a significant reduction of model fit (l2 = 92.835, df = 10, p = 0.001), indicating the distribution changed between the regions after 1986, with most of the range expansion occurring in far north (z = 5.352, p = 0.001) (figure 2d). wildebeest strongly favoured knob thorn – marula savannah on basalt and tended to avoid mopane woodland, thorn thickets near rivers, sour bushveld and sandveld (table 1). their presence in bushwillow woodland increased marginally after 1986 (χ2 = 5.760, df = 1, p = 0.016). kudu kudu were distributed throughout the knp, but with a greater concentration in the southern half than in the northern sections (figure 7). pienaar’s (1963) map shows a blanket distribution of kudu throughout the knp. figure 7: distribution range of kudu comparing the periods (a) 1980–1986 and (b) 1987–1993. concentration zones enclosing 75% of the population (dark shade) are distinguished from the 95% or 99% range limits (grey shade). although the overall occurrence of kudu remained high after 1986, their relative presence in the northern half of knp contracted (figure 2e; l2 = 23.407, df = 3, p = 0.001). kudu positively selected mountain bushveld and avoided sandveld (table 1). their presence in mopane shrubveld decreased significantly after 1986 (χ2 = 6.572, df = 1, p = 0.017), but it increased in sour bushveld (χ2 = 7.419, df = 1, p = 0.012). giraffe giraffe occurred throughout the southern half of knp, but were more sparsely distributed in the northern half (figure 8). pienaar’s (1963) map shows that they had formerly been absent from the mopane zone north of the letaba river, except for a pocket in the north-eastern basaltic plains. figure 8: distribution range of giraffe comparing the periods (a) 1980–1986 and (b) 1987–1993. concentration zones enclosing 75% of the population (dark shade) are distinguished from the 95% or 99% range limits (grey shade). giraffe continued expanding their presence in the northern half of the knp during the survey period (l2 = 9.454, df = 3, p = 0.024), particularly in the far north (z = 6.907, p = 0.001) (figure 2f). nevertheless, in general they avoided mopane-dominated vegetation as well as the sandveld, but occurred fairly evenly through other habitat types (table 1). waterbuck waterbuck showed a ribbon distribution concentrated along perennial and seasonal rivers (figure 9). pienaar’s (1963) map shows a restricted distribution of waterbuck along rivers very similar to that documented during the subsequent surveys. figure 9: distribution range of waterbuck comparing the periods (a) 1980– 1986 and (b) 1987–1993. concentration zones enclosing 75% of the population (dark shade) are distinguished from the 95% or 99% range limits (grey shade). although the overall proportion of occupied 5 km × 5 km cells in the knp appeared to have increased much (l2 = 18.954, df = 10, p = 0.041), the actual regional distribution did not change significantly (figure 2g; l2 = 2.595, df = 3, p = 0.476) after 1986. waterbuck favoured mountain bushveld, whilst avoiding bushwillow woodland, thorn thickets and sandveld (table 2). table 2: proportion of animal sightings recorded in 0.95 probability isopleths for less common species within each habitat type, expressed as a percentage relative to the proportional extent of these habitats in the kruger national park. warthog warthog occurred over most of the knp prior to 1987, but after 1986 had disappeared from most of the basaltic region of the north-east (figure 10). warthog appeared to be restricted more narrowly to the vicinity of rivers in the northern half of the knp around 1960 than was evident after 1980 (pienaar 1963). figure 10: distribution range of warthog comparing the periods (a) 1980–1986 and (b) 1987–1993. concentration zones enclosing 75% of the population (dark shade) are distinguished from the 95% or 99% range limits (grey shade). warthog showed a slight shrinkage in their distribution, which seemed to be concentrated more in the southern half of the knp (figure 2h; l2 = 5.257, df = 3, p = 0.154). warthog did not appear to favour any habitat type, whilst avoiding sour bushveld and sandveld. after 1986, warthog became less commonly present in mopane-shrubveld (χ2 = 33.893, df = 1, p = 0.001) and mountain bushveld (χ2 = 6.812, df = 1, p = 0.014). sable antelope sable antelope showed a patchy distribution in the southern half of the knp, but occurred more continuously in the northern part (figure 11). they had disappeared from sections of the north-east after 1986, following their general population decline. sable formerly had a wider distribution in the south-west and in the western region of the central section of the knp than was recorded after 1980 (pienaar 1963). figure 11: distribution range of sable antelope comparing the periods (a) 1980– 1986 and (b) 1987–1993. concentration zones enclosing 75% of the population (dark shade) are distinguished from the 95% or 99% range limits (grey shade). overall, the proportion of cells occupied by sable contracted after 1986 (l2 = 131.640, df = 10, p = 0.001), despite no significant change in their regional distribution (l2 = 3.523, df = 3, p = 0.318) (figure 2i). although most sable occurred in mopane woodland, they showed a relative preference for sour bushveld and sandveld, whilst avoiding knob thorn – marula savannah, mountain bushveld and thorn thickets (table 2). after 1986, they had declined most substantially in their presence in mopane-shrubveld (χ2 = 8.812, df = 1, p = 0.012). tsessebe tsessebe were found mostly in the north-east region of the knp, apart from an isolated pocket in the south-east (figure 12). around 1960, tsessebe had occurred throughout the far north as well as along the western border of the central section (pienaar 1963). figure 12: distribution range of tsessebe comparing the periods (a) 1980–1986 and (b) 1987–1993. concentration zones enclosing 75% of the population (dark shade) are distinguished from the 95% or 99% range limits (grey shade). the distribution range of tsessebe remained unchanged despite their population decline after 1986, both absolutely and relatively (figure 2j; l2 = 1.026, df = 3, p = 0.795). tsessebe showed a restriction almost entirely to mopane-shrubveld (table 2). eland eland were present through much of the northern half of the knp, with a concentration in the far north (figure 13). their distribution appears to have been somewhat more extensive in the northern half around 1960 than recorded more recently (pienaar 1963). eland remained restricted almost entirely to mopane woodland and shrubveld (table 2). figure 13: distribution range of eland comparing the periods (a) 1980–1986 and (b) 1987–1993. concentration zones enclosing 75% of the population (dark shade) are distinguished from the 95% or 99% range limits (grey shade). roan antelope roan antelope were present patchily in the north-eastern region of the knp and the isolated pockets that had occurred in the north-west and south-west had disappeared after 1986 (figure 14). prior to 1963, roan were widely distributed through most of the far north and occurred also in a pocket in the north-west of the central section (pienaar 1963). their presence in the south-west appears to have been somewhat wider than the isolated herd recorded there after 1980. roan occupied mostly the mopane shrubveld (table 2). figure 14: distribution range of roan antelope comparing the periods (a) 1980– 1986 and (b) 1987–1993. concentration zones enclosing 75% of the population (dark shade) are distinguished from the 95% or 99% range limits (grey shade). trustworthiness top ↑ this study comprises our joint original research, apart from our dependence on the aerial survey data made available to us by national parks scientists. aerial surveys are inevitably undercounts and the extent of the bias can be affected by prevailing conditions. however, from the experience of park scientists, which is also supported by differential ground counts, we do not think undercounting will bias the current interpretation. furthermore, surveys were conducted in well-defined census blocks using similar techniques and methods over the 14-year study period, a factor which improves the reliability of the data and validity of the conclusions reached. discussion top ↑ kudu, zebra, buffalo and impala were the species distributed most widely throughout the knp. although wildebeest and warthog also occurred widely in the southern half, they showed a more restricted distribution in the northern half of the park, a pattern exhibited more extremely by giraffe. waterbuck were concentrated mainly along rivers, but also in mountain bushveld along the eastern border. less common antelope species, such as sable antelope, tsessebe, eland and roan antelope, showed patchy distributions concentrated mostly in the northern half of the park. wildebeest, giraffe, zebra and warthog favoured the knob thorn – marula savannah on basalt substrates, whilst sable antelope, tsessebe, eland and roan antelope favoured mopane-dominated vegetation in the north and, in the case of sable antelope, also the sour bushveld and sandveld that was negatively selected by other herbivore species. relative declines in the proportion of the population found in the more arid northern half of the knp could be a consequence of progressive habitat degradation following the extreme drought conditions experienced in 1982–1983 and 1992–1993 and persistently low rainfall from 1987 through 1994. the greatly widened surface water distribution in this region of the knp may have contributed by attracting increased presence of zebra, which interacted with the rarer grazers either by pre-empting resources or by attracting greater numbers of lions under these stressful conditions (owen-smith & mills 2006). the increased presence of buffalo herds in the south-west could have contributed to the decline in sable numbers in this region (owen-smith et al. 2012). the maps presented by pienaar (1963) suggest that the distributions of impala, buffalo and warthog were concentrated more strongly near rivers during the dry season prior to 1963 than was evident over 1980–1993, particularly in the northern half of the knp. their widened occupation of areas away from rivers was perhaps a consequence of the widened provision of artificial water sources during the 1970s. around 1960, wildebeest, tsessebe and sable antelope had apparently been more prevalent in the west of the central section of the park than more recently. their decreased presence in this region is most likely a consequence of the western boundary fence completed in 1961, blocking dry season movements towards the private nature reserves to the west. both wildebeest and zebra showed dry season concentrations over 1980–1993 in parts of the eastern central section that were indicated as wet season range by pienaar (1963). zebra and giraffe widened their presence in the far north compared with their occurrence there prior to 1963, whilst buffalo extended their distribution towards the south-west region of the south section. for zebra, the shift towards the north appeared to be a direct consequence of the widened availability of surface water there (harrington et al. 1999). sable antelope, roan antelope and tsessebe were formerly more widely distributed through the northern part of the knp than was recorded after 1980. whether this was caused by increasing aridity in this region or the increased presence of zebra, and hence a shared predator, cannot be established from the distribution change alone, although other evidence implicates increased predation as a contributory factor (owen-smith et al. 2012). since 1995, many artificial waterpoints have been closed (smit, grant & whyte 2007) and fences along parts of both the western and eastern boundaries have been removed. vegetation changes are evident towards an opening of tree canopy cover in knob thorn – marula parkland, but with a greater density in shrub cover evident on granitic substrates (eckhardt, van wilgen & biggs 2000; trollope et al. 1998). rainfall within the park has recently shown wider annual variation than during the earlier historical record (ogutu & owen-smith 2003). elephants and white rhinos continue to increase towards abundance levels not manifested in the knp region within the past 150 years, expanding their substantial impacts on vegetation structure. the consequences of increased atmospheric carbon dioxide levels for temperature regimes, rainfall and the competitive balance between c4 plants (primarily tropical grasses) and c3 plants (particularly woody trees and shrubs) are likely to become additional influences on vegetation trends and water flow in rivers. hence, further changes in the populations and distributional ranges of large herbivores within the knp are to be expected subsequent to the period covered by our data base. our objectively derived maps provide a reference for future changes in distribution patterns to be identified. conclusion top ↑ changes in the distribution patterns of several ungulate species have occurred since 1963 and seem to be continuing. we have identified the closure of the western boundary and expanded availability of perennial surface water sources as likely contributory influences recently, but further changes in distribution in response to the effects of climate change on vegetation are to be anticipated. further monitoring of species presence that is both spatially comprehensive and sufficiently explicit locally will be needed to document the continuing effects of both extrinsic and intrinsic drivers on distribution ranges. acknowledgements top ↑ we express special thanks to sandra macfadyen for making available to us the spatial data for the knp and for her quick responses to our queries. j.g. chirima’s study was supported by a bursary awarded by the south african national research foundation. competing interests the authors declare that they have no financial or personal relationship(s) which may have inappropriately influenced them in writing this paper. authors’ contributions g.j.c. (university of the witwatersrand) performed all the data analyses and wrote the manuscript. n.o-s. (university of the witwatersrand) supervised g.j.c.’s (university of the witwatersrand) phd study and contributed to in-depth discussions through sharing his knowledge of animal and plant ecology pertaining to the knp to improve the manuscript. b.f.n.e. (university of the witwatersrand) co-supervised the phd study and provided valuable comments to improve the manuscript. for publication, the manuscript was prepared when g.j.c. (university of the witwatersrand) was working for the south african environmental observation network and later the agricultural research council. references top ↑ agresti, a., 1996, an introduction to categorical data analysis, john wiley and sons, new york.christensen, r., 1997, log-linear models and logistic regression, springer-verlag inc., new york. craigie, i.d., baillie, j.e.m., balmford, a., carbone, c., collen, b., green, r.e. et al., 2010, ’large mammal population declines in africa’s protected areas’, biological conservation 143, 2221–2228. http://dx.doi.org/10.1016/j.biocon.2010.06.007 eckhardt, h.c., van wilgen, b.w. & biggs, h.c., 2000, ‘trends in woody vegetation cover in kruger national park, south africa, between 1940 and 1998’, african journal of ecology 38, 108–115. http://dx.doi.org/10.1046/j.1365-2028.2000.00217.x environmental systems research institute, 2010, arcmap version 10.0, computer software, esri, redlands. gaston, k.j., 1990, ‘patterns in the geographic ranges of species’, biological reviews 65, 105–29. http://dx.doi.org/10.1111/j.1469-185x.1990.tb01185.x gaston, k.j., 2003, the structure and dynamics of geographic ranges, oxford university press, oxford. gertenbach, w.p.d., 1983, ‘landscapes of the kruger national park’, koedoe 26(1), 9–12. getz, w.m. & wilmers, c.c., 2004, ‘a local nearest-neighbor convex-hull construction of home ranges and utilization distributions’, ecography 27, 498–505. http://dx.doi.org/10.1111/j.0906-7590.2004.03835.x getz, w.m., fortmann-roe, s., cross, p.c., lyons, a.j., ryan, s.j. & wilmers, c.c., 2007, ‘locoh: nonparametric kernel methods for constructing home ranges and utilization distributions’, plos one 2, 1–11. http://dx.doi.org/10.1371/journal.pone.0000207, pmid:17299587 grant, c.c., davidson, t., funston, p.j. & pienaar, d.j., 2002, ‘challenges faced in the conservation of rare antelope: a case study on the northern basalt plains of the kruger national park’, koedoe 45(2), 45–62. harrington, r., owen-smith, n., viljoen, p.c., biggs, h.c., mason, d.r. & funston, p., 1999, ‘establishing the causes of roan antelope decline in the kruger national park, south africa’, biological conservation 90, 69–78. http://dx.doi.org/10.1016/s0006-3207(98)00120-7 joubert, s.c.j., 1983, ‘a monitoring programme for an extensive national park’, in r.n. owen-smith (ed.), management of large mammals in african conservation areas, pp. 201–212, haum, pretoria. joubert, s.c.j., 2007a, the kruger national park – a history, vol. 1, high branching, johannesburg. joubert, s.c.j., 2007b, the kruger national park – a history, vol. 2, high branching, johannesburg. knoke, d. & burke, p.j., 1980, log-linear models, sage publications inc., newberry park.  lawton, j.h., 1993, ‘range, population abundance and conservation’, trends in ecology and evolution 8, 409–413. http://dx.doi.org/10.1016/0169-5347(93)90043-o mills, m.g.l., biggs, h.c. & whyte, i.j., 1995, ‘the relationship between rainfall, lion predation and population trends in african herbivores’, wildlife research 22, 75–88. http://dx.doi.org/10.1071/wr9950075 ogutu, j.o. & owen-smith, n., 2003, ‘enso, rainfall and temperature influences on extreme population declines among african savanna ungulates’, ecology letters 6, 412–419. http://dx.doi.org/10.1046/j.1461-0248.2003.00447.x owen-smith, n. & ogutu, j., 2003, ‘rainfall influences on ungulate population dynamics’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 310–331, island press, washington, dc. owen-smith, n., mason, d.r. & ogutu, j.o, 2005, ‘correlates of survival rates for ten african ungulate populations: density, rainfall and predation’, journal of animal ecology 74, 774–788. http://dx.doi.org/10.1111/j.1365-2656.2005.00974.x owen-smith, n. & mills, g.l.m., 2006, ‘manifold interactive influences on the population dynamics of a multispecies ungulate assemblage’, ecological monographs 76, 73–92. http://dx.doi.org/10.1890/04-1101 owen-smith, n., chirima, g.j., macandza, v. & le roux, l., 2012, ‘shrinking sable antelope numbers in kruger national park: what is suppressing population recovery?’, animal conservation 15, 195–204. http://dx.doi:10/1111/j.1469-1795.2011.00504.x pienaar, u. de v., 1963, ‘the large mammals of the kruger national park – their distribution and present-day status’, koedoe 6, 1–37. quinn, g.p. & keough, m.j., 2002, experimental design and data analysis for biologists, cambridge university press, cambridge. redfern, j.v., grant, r., biggs, h. & getz, w., 2003, ‘surface-water constraints on herbivores foraging in the kruger national park, south africa’, ecology 84, 2092–2107. http://dx.doi.org/10.1890/01-0625 smit, i.p.j., grant, c.c. & whyte, i.j., 2007, ‘landscape-scale sexual segregation in the dry season distribution and resource utilization of elephants in kruger national park, south africa’, diversity and distributions 13, 225–236. http://dx.doi.org/10.1111/j.1472-4642.2007.00318.x trollope, w.s.w., trollope, l.a., biggs, h.c., pienaar, d. & potgieter, a.l.f., 1998, ‘long-term changes in the woody vegetation of the kruger national park, with special reference to the effects of elephants and fire’, koedoe 41, 103–112. van wilgen, b.w., govender, n., biggs, h.c., ntsala, d. & funda, x.n., 2004, ‘response of savanna fire regimes to changing fire management policies in a large african national park’, conservation biology 18, 1533–1540. http://dx.doi.org/10.1111/j.1523-1739.2004.00362.x venter, f.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation pattern’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 82–129, island press, washington, dc. viljoen, p.c., 1993, ‘ecological aerial surveys in the kruger national park: 1992’, scientific report 2/93, national parks board, skukuza. viljoen, p.c. & retief, p.f., 1994, ‘the use of global positioning system for real-time data collected during ecological aerial surveys in the kruger national park, south africa’, koedoe 37(2), 149–157. whyte, i.j., biggs, h.c., gaylard, a. & braack, l.e.o., 1999, ‘a new policy for the management of the kruger national park’s elephant population’, koedoe 42(1), 111–133. filelist convert a pdf file! koedoe 19: 171-174 (1976) additions to the check-list of birds of the mountain zebra national park b. l. penzhorn and p. j. l. bronkhorst mountain zebra national park private bag x66 cradock 5880 abstract twenty-two additional bird species have been recorded in the park, increasing the check-list total to 170 species. in his report on the bird-life in the mountain zebra national park, skead (1965) reported 148 bird species in the park. he predicted that vagrants could increase the total to 190-200 species. since the publication of skead's check-list, several bird-watchers have visited the park and compiled lists of the birds seen. the authors have also kept a look-out for species not previously reported. twenty-two additional species have been recorded, increasing the number of species recorded in the park to 170. the 22 additional species are listed below. the numbers denote those used by mclachlan and liversidge (1957) in roberts birds oj south africa and the nomenclature follows the check list oj the birds oj south africa (s.a.o.s. list committee 1969). 47 white-breasted cormorant phalacrocoraxlucidus witborsduiker (lichtenstein) white-breasted cormorants are recorded regularly at the large dam near the museum. 50 reed cormorant rietduiker phalacrocorax africanus (gmelin) a few vagrants have been recorded occasionally at the large dam near the museum. 61 cattle egret bosluisvoel ardeola ibis (linnaeus) cattle egrets are recorded regularly, although not resident in the park. skead (1965) predicted that vagrants from the lucerne lands in the great fish river valley would visit the park. 79 black stork ciconia nigra (linnaeus) groot swartsprinkaanvoel black storks have been recorded infrequently during the summer months. 171 85 spoonbill lepelaar platalea alba scopoli spoonbills are observed occasionally at dams in the park. on one occasion six were seen together at the large dam near the museum. 95 black duck swarteend anas sparsa eyton black duck are seen on the wilgerboom river, usually where there is some cover, but a pair has been observed on a few occasions in an open valley at the upper reaches of the river. a black duck was flushed off a nest containing seven eggs on 1975.09.19. the nest was situated on a ledge on a cliff-face, about six metres above the level of the river. 114 lanner falcon falco biarmicustemminck edelvalk the presence of this species, predicted by skead (1965), can now be confirmed . it appears to be rare in the park. 122 greater kestrel groot rooivalk falco rupicoloides a. smith members of the eastern cape wild bird society observed six greater kestrels near the look-out point in the park in march 1969 (brown 1969a). 134 tawny eagle kouvotl aquila rapax (temminck) a single bird was seen on the rooiplaat plateau on three consecutive days during december 1973 (b.l.p.) . this species was regarded by skead (i 967) as rather a vagrant of irregular appearance anywhere throughout the eastern cape. 158 little sparrowhawk klein sperwer accipiter minullus (daudin) a single bird was observed once in the dense riverine scrub in march 1973 (b .l.p.). in the eastern cape this species usually occurs in the forests of the coastal and sub-coastal belt (skead 1967). 210 moorhen waterhoender gallinula chloropus (linnaeus) a single moorhen was seen on a few consecutive days in september 1975 below the museum at the drift in the wilgerboom river. 245 blacksmith plover bontkiewiet vanellus armatus (burchell) blacksmith plovers have been recorded infrequently, mainly at the small dam near the nature conservator's house. 172 258 common sandpiper gewone ruiter tringa hypoleucos linnaeus single birds have been seen on sandbanks along the wilgerboom river. 367 cape eagle-owl kaapse ooruil bubo capensis a. smith an injured owl found in the park was identified as a cape eagleowl by p. j. l. b. 368 spotted eagle-owl gevlekte ooruil bubo ajricanus (temminck) these owls are occasionally seen at dusk and at night. 373 fiery-necked nightjar suid-afrikaanse naguil caprimulgus pectoralis cuvier on moonlit nights, these night jars are infrequently heard calling. recorded at berghof, babylons toren, the museum and in fonteinkloof. 397 malachite kingfisher kuifkopvisvanger alcedo cristata pallas recorded once at a small dam in march 1971 and along the wilgerboom river in september 1975. 485 grey-backed finch-lark eremopterix verticalis (a. smith) grys kaffertjie recorded infrequently on the rooiplaat plateau. 517 fork-tailed drongo mikstertbyvanger dicrurus adsimilis (bechstein) recorded regularly in the dense acacia karroo scrub in the vahey near the nature conservator's house and in the tall exotic trees at the house itself. 784 house sparrow engelse mossie passer domesticus (linnaeus) house sparrows were first recorded in the park in june 1968 by penzhorn and morris (1968). they must have arrived after 1964, when skead (1965) concluded his survey of the bird-life in the park. house sparrows are resident at babylons toren, berghof, weltevrede and the museum. 808 red bishop-bird rooi kaffervink euplectes orix linnaeus red bishop-birds have been recorded by p.j. l. b. 173 859 yellow-eye canary geeloogsysie serinus mozambicus (m tiller) brown (1969b) recorded a small flock of these canaries in the park in march 1969. the following eight species have also been reported, but require confirmatiqn: 146 black-breasted snake-eagle circaetus pectoralis a. smith 160 african goshawk accipiter tachiro (daudin) 371 european night jar caprimulgus europaeus linnaeus 395 giant kingfisher ceryle maxima (pallas) 458 rufous-naped lark mirafra africana a. smith 519 golden oriole oriolus oriolus (linnaeus) 540 rock-jumper chaetops frenatus (temminck) 649 tawny-flanked prinia prinia subjlava (gmelin). acknowledgements thanks are due to mr c. j. skead for reading and commenting on the manuscript. references brown, d. m . v. 1969a. new distributional data: 2. greater kestrel. ostrich 40: 131. brown, d. m. v. 1969b. new distributional data: 2. yellow-eye canary. ostrich 40: 132. mclachlan, g. r. and r. liversidge. 1957. roberts birds oj south africa. 2nd ed. cape town: trustees of the s.a. bird book fund. penzhorn, b. l. and a. k. morris. 1968. new distributional data: i. house sparrow. ostrich 39:272. s.a.o.s. list committee. 1969. check list oj the birds oj south africa. cape town: south african ornithological society. skead, c. j. 1965. report on the bird-life in the mountain zebra national park, cradock, c.p. 1962-1964. koedoe 8:1-40. skead, c. j. 1967. ecology of birds in the eastern cape province. ostrich suppl. 7: 1-103. 174 page 1 page 2 page 3 page 4 filelist convert a pdf file! supplement to koedoe. 1977: 143-151. the status of wildlife conservation in botswana w von richter f a 0 wildlife ecologist department of wildlife, national parks and tourism pobox 131 gaberone republic of botswana the sparse human population and the general lack of surface water over most parts of the republic of botswana, which has hampered rapid expansion of agricultural activities into the less suitable areas in the past, have contributed to the fact that botswana still supports a varied and rich wildlife population. the long history of hunting by the local populae makes them understand and appreciate the concept of wildlife conservation and utilization and has assisted in general to implement a policy for rational conservation and utilization. the next decade will be decisive whether this laudable state of affairs will continue or whether the wildlife resource will be depleted and finally restricted only to formal conservation areas as it has happened in many other countries on the african continent. the government is fully aware of the significance of wildlife conservation and utilization and the necessity to integrate it into overall landuse planning. the environment botswana is a flat, landlocked country of approximately 569800 km2 , situated in the middle of the dry southern african plateau at an average altitude of 1 000 m. the majority of the country belongs to the south west arid zone . approximately two-thirds of the country is covered by a deep layer of kalahari sands of geologically recent wind blown origin. surface water is restricted to the limpopo watershed in the east, the kwando-linyanti-chobe system, which forms part of the zambezi system, in the north and the okavango river and delta in the north west. at one time both the latter have been largely infested by tsetse fly glossina morsitans, which inhibited settlement by man and his cattle; the okavango delta is still largely infested by the fly. the makgadikgadi pans complex formed in ge?logical times a huge lake, but is now a salina only filled to a depth of a few centimeters during times of exceptional high local rainfall. the kalahari covers two-thirds of the country, and is interspersed with numerous pans, round or elleptical depressions, and ancient river beds which emptied in wetter periods into 143 national parks and game reserves in botswana national parks game reserves proposed consolidations and new reserves roads 144 (1) existing national parks and game reserves 1 chobe national park (10 800 km2) est. 1961, elevated to national park 1968. 2 mikaelelo game reserve (360 km2) est. 1974. 3 moremi wildlife reserve (3 880 km2) est. 1965, enlarged in 1976. 4 nxai pan national park (2 100 km2) est. 1971. 5 makgadikgadi pans game reserve (3 900 km2) est. 1970. 6 central kalahari game reserve (52 800 km2) est. 1957. 7 gemsbok national park (24 800 km2) est. 1932, enlarged and upgraded. 8 mabuasehube game reserve (1 800 km2) est. 1971 to national park in 1971. 9 khutse game reserve (2 500 km2) est. 1971. 10 gaborone dam national park (5 km2) est. 1975. 15 maun educational game park (5 km2) est. 1969. (2) proposed new conservation areas 11 okavango river front. 12 tsodilo hills, aha mountains, kwihabe hills. 13 waterfowl sanctuary lake ngami. 14 kwebe hills. 16 tamafupajari pans complex. 17 pataletsabe hills and mannyelanong hills. 18 mannyelanong hill game reserve (pending gazettement). 19 shashe elephant reserve. 20 lepokola hills. 21 tswapong and mabeleapodi hills. 22 shoshong hills. 23 mokwane hills. 24 notwane limpopo area. the makgadikgadi. the soil and vegetation of these pans and river beds are of a different type than the surrounding sandveld and are of vital importance to humans, wildlife and cattle for their survival in this harsh environment. rainfall is varied and erratic, confined mainly to the summer (november to april), declining from an average of 650 mm in the north east to under 200 mm in the south west. shrub and tree savanna vegetation types are dominant over most of botswana. human population is low (approximately 1 person / km2); approximately 80% of the population is, however, confined to th e areas of better soils and more reliable rainfall in the east, and the fringes of the okavango delta, leaving large tracts of the country virtually uninhabited, save the few groups of nomadic bushman. most people have a history of pastoralism, although the ownership of the national cattle hercl is very skewed , i.e. a minority owns the majority of cattle. subsistence arable agriculture is very small and mostly confined to the more suitable soils. wildlife utilization has played and still plays a significant role in the life of the people of botswana. in times of catastrophic events such as rinderpest, foot and mouth disease or drought, the rural population has turned to wildlife as a means for support. in the rural areas wildlife still is a significant factor for the provision of protein and cash, although there is a noticeable trend away from purely subsistence hunting to commercial exploitation of the resource. 145 policy of wildlife conservation the policy for wildlife conservation in botswana has to match government's four objectives for development of the country: rapid economic growth, social justice, economic independence and sustained production. rural development and the raising of the standards ofliving in the rural areas is and integral part of this policy. policies for the conservation and utilization of the wildlife resource follows therefore from the above, and they can be summarised as such: (i) the establishment of national parks and game reserves to protect and conserve a cross section of all species and habitats occurring in botswana; (ii) complete protection of rare and endangered species; (iii) the implementation of rational and realistic conservation measures outside established conservation areas as a condition for optimum u tiliza tion; (iv) to provide adequate and relatively cheap hunting for the rural population who still depend to a significant degree on wildlife for their day-to-day living; (v) to optimise the financial returns from non-resident recreational hunting; (vi) to encourage non-hunting photographic tourism to reduce the dependence on non-resident hunting as the major revenue earner; and (vii) to educate all strata of society of the value of wildlife conservation, and that its utilization should benefit all. the status of conservation in botswana currently approximately 17% of botswana enjoys national park and game reserve status, comprising four national parks and six major game reserves (fig. 1). the national parks and game reserves system covers adequately all major habitat types in botswana, although none of these constitute ecological entitles. it is the intention of the department to consolidate some of the existing national parks and game reserves to make for better and more meaningful conservation and administration. in addition a number of new sites have been identified (fig. 1) for inclusion in the national parks and game reserves system, primarily on the grounds of their scientific, cultural or scenic values. the dichotomy of conserved areas in botswana is the result of historical and political reasons, i.e. national parks may only be established on stateland or any other land donated or bequeathed for this purpose to the president through an act of parliament. the national parks act of 1967 provides total protection to the entire environment of a park. game reserves are established under the fauna conservation proclamation of 1961 by the president on advice of cabinet after 146 consultation with the local authorities. in a strict legal sense game reserves provide only protection for vertebrate animals as no mention of any other environmental features are made; in practice, however, they also provide protection for the entire ecosystem . in 1976 it was decided that the national parks act should apply to all game reserves after their elevation to national parks status, regardless whether they are situated on state or tribal land . in this way more permanency in tenure and an uniform approach to management and development of all conserved areas will be achieved . as a number of new sites will not qualify for national park status, the introduction of a new category of reserved areas is contemplated. nature reserves will be designated those areas which warrant formal conservation status for various reasons, although they do not qualify for national parks status. certain activities will be permitted as long as they are compatible with the purpose for which the reserve was established. recognising the potential conflict between conserving an ecosystem and its development for tourism, all national parks and game reserves are being zoned in the following categories: wilderness area, low, medium and high density tourist areas; each of the four categories will be managed differently, according to their classification. the fauna conservation proclamation of 1961 makes provision for the gazetting of rare and endangered species as conserved animals which may not be hunted except in the case of defence of life and property. table i lists all currently conserved animals. the white rhinoceros is the table 1 conserved animals in botswana antbear aardwolf black-footed cat brown hyena cheetah civet giraffe hippopotamus honey badger klipspringer mountain rheedbuck night-ape oribi otter pangolin puku rhinoceros (black and white) roan antelope serval vaal rhebok yellow-spotted dassie waterbuck 147 all cranes all eagles all egrets fishing owl all flamingoes hammerkop all herons all jacanas kgori bustard all pelicans secretary bird spoonbill stanley bustard all storks all vultures only species which became exterminated during this centuary in botswana and a re-introduction programme is well under way to re-establish the species. wildlife utilization in botswana has two aspects, hunting by residents and non-residents and photographic tourism. consumptive utilization of wildlife in botswana can be separated in three categories: (i) traditional or subsistence huntong carried out by the local population and accounting for approximately 85% of the legal offtake (± 15 000 animals in 1975). the financial return to local or central government from this type of hunting is small, but the significance has to be seen in the socio-economic context of a rural population; (ii) superimposed on traditional hunting is non-resident and resident recreational hunting, which accounts for approximately 83% of all direct revenue accrued from wildlife utilization; and (iii) an industry utilising the trophies, hides and skins of wild animals is well established in botswana and benefits from the hunting activities. the most recent development is the emergence of non-hunting photographic tourism centred around the national parks and game reserves, primarily in northern botswana. currently 31 tourist operators conduct tours in botswana, ranging from the more luxury type to the "do-ityourself' arrangements. for the purpose of recreational hunting, botswana is subdivided into 40 controlled hunting areas (cha) of which presently 13 are leased as concession areas to four safari companies . the remainder of the cha's are open to booking by residents as non-residents are only permitted to hunt through a safari company. annual quotas are being set by the department of wildlife, national parks and tourism in consultation with the local authorities for each controlled hunting area. residents and non-residents pay c.h.a. booking fee (rio and r50/ week/hunter respectively) while tribesmen or stateland residents hunting in their areas are exempted from paying booking fee. after purchasing a general game licence (r50 for 4 animals for non-resident and r3 for 7 animals for a resident) the hunter may obtain supplementary licences which vary between ri to rio for a resident and r5 to r200 for a non-resident. protected species (lion, leopard, sable and eland) command the same fee both for resident and non-resident (r500, r300, r250, rl20 respectively). hunting regulations for tribesmen and stateland residents are complicated and the numbers of animals made available on a licence and fees vary considerably from area to area, due to historical and political reasons, introducing a measure of discrimination between citizens of botswana, depending where they reside. recognising this fact, the department is presently drafting regulations to introduce a unified 148 hunting system, differentiating basically only between residents/citizens and non-residents. recreational hunting, conducted through safari companies, has proved to be the most lucrative and easily to administer form of wildlife utilization in botswana, and it is the intention of the department to continue this aspect of hunting, modifying it in such a way as to optimise the returns of the country and make the best possible use of the wildlife resource. recognising the need to diversify the utilization of wildlife, encouragement is given to the development of non-hunting photographic tourism. the greatest asset botswana can offer to the wildlife viewing tourist is the unspoilt wilderness of her national parks and game reserves, and botswana therefore encourages high price/ low volume tourism. no permanent and costly tourism infrastructure is developed in the conserved areas, apart from an essential road network. development of the more luxury facilities is, however, encouraged through private enterprise outside the national parks and game reserves. wildlife utilization contributed r427 144 in direct revenue to local and central government in 1974, while the annual economic contribution of the wildlife based industry amounted to r3 ,7 million after deduction of all overheads and costs. a detailed knowledge of the ecology and biology of the habitats and species is a prerequisite for rational conservation and utilization of the wildlife resource, and since its inception the department has encouraged wildlife research. owing to the lack of finance and trained manpower in botswana, most of the research has been financed and carried out by outside research workers and donors. the department has initiated most of the projects, and co-ordinates and supervises them, in order to obtain the necessary information for the proper conservation and utilization of th e wildlife resource. the research projects range from ecological surveys of important wildlife areas, the monitoring of seasonal distribution and abundance of major wildlife species, to more detailed studies of individual species for management and conservation purposes. wildlife forms an important asset to botswana and its integration in the overall development is essential if wildlife is to survive in m ea ningful population in the country. the recently introduced tribal land grazing policy calls for a major input into landuse planning on a regional and national scale. agreement has been reached for the establishing and gazetting of wildlife management areas in which wildlife utilization will take preference over any form of landuse, and detrimental and competing landuses may be prohibited. it is anticipated that some of the areas will eventually be given over to other forms of landuse, but pending detailed assessment and evaluation for their ultimate use, their productivity is conserved. the involvement of the department in the landuse planning process is assured through the various regional and central landuse planning committees. 149 the conservation of wildlife in botswana for posterity will only succeed in the long run if the human population effected by the conservation measures accepts those as valid and reasonable. the education of all strata of the population to the value of wildlife conservation, in terms of financial as well as ethical and aesthetic benefits, is therefore of a paramount importance. the department has, with the assistance of outside donors, embarked on an education programme, aiming at the various groups of the population, and using all available media, to bring greater awareness of the importance of conservation to the people. the department is hampered in the effective execution of its duties and responsibilities by the lack of appropriate funds and trained manpower. the current worldwide financial crisis does not help to alleviate the situation in the near future and it is therefore necessary to continue as best as possible with the available resources. recognising the need for more appropriate training for the field staff to increase their efficiency and morale, it is hoped to establish in the near future a training centre in botswana, while medium level staff will continue to receive training at the college for african wildlife management in tanzania. future prospects for conservation in botsw;ma botswana is fortunately not as much plagued by overpopulation and scarcety of land as many other african countries. development during the last decade has been gradual and the ecological impact on the environment has not been too severe as a result. government is aware of the fragility of most of botswana's environment and the risks involved in large scale modification to it. various legislation has been passed in the last few years relating to the conservation and wise use of the natural renewable resources. the tribal land grazing policy attempts to arrest the uncontrolled spread oflow management cattle raising at the expense of the land and introduce forms of mangement assuring sustained production, raising at the same time the standard of living of the rural population. wildlife has been recognised as a significant factor in the economy of the country, and the necessity of its integration into the overall land use planning process is adcepted. botswana has at the present stage of the development, still most options open to make rational land use decisions, insuring that wildlife will not be restricted only to conservation areas, but continue to roam freely in many parts of botswana in the future. it is not unrealistic to expect that botswana may be one of the few countries where wildlife conservation and utilization will complement rather than compete with other, more conventional forms of land use. 150 conclusion during the last 10 years botswana has developed a viable national parks and reserve system, covering 17% of the co"jlntry and giving conservation to a cross section of all major habitats. a hunting system has been developed where the rural population enjoys cheap and generous licences and still permitting a lucrative non-resident hunting industry based on safari companies. non-hunting photographic tourism is emerging as an additional form of wildlife utilization, attracted by the wilderness the visitors can experience. there is growing recognition of wildlife as an alternative and complementary form of landuse, and the regional and national landuse planning exercise attempts to integrate wildlife . considering the limited resources available to botswana for the conservation of wildlife and its rational utilization, the results of these efforts are remarkabje. botswana has reached a stage of her development, where still many options remain open for continuous and increased efforts to conserve wildlife for future generations, as an integral part of the country's devel?pment process. lsi page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 article information authors: frederik j. venter1 navashni govender2 affiliations: 1conservation management, kruger national park, south africa 2scientific services, kruger national park, south africa correspondence to: navashni govender postal address: private bag x402, skukuza 1350, south africa dates: received: 10 dec. 2010 accepted: 21 nov. 2011 published: 14 may 2012 how to cite this article: venter, f.j. & govender, n., 2012, ‘a geomorphic and soil description of the long-term fire experiment in the kruger national park, south africa’, koedoe 54(1), art. #1037, 10 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1037 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. a geomorphic and soil description of the long-term fire experiment in the kruger national park, south africa in this short communication... open access • abstract • introduction    • study site    • experimental burn plots • methods    • assumptions • results    • pretoriuskop    • skukuza    • satara    • mopane • discussion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ in 1954, the experimental burning programme into fire research was initiated in the kruger national park (knp), south africa. it is viewed as one of the last remaining long-term landscape fire experiments in africa. throughout the more than five decades of fire treatments in the experiment, numerous surveys (expanding various spatial and temporal scales), research projects (covering biotic and abiotic components) and analyses have been conducted with the aim to assess the impacts of different fire regimes on the savannah biome. the design of the experiment intended to test the effect of season and frequency of burning on vegetation within four major landscapes in the knp. however, these effects have been partly obscured by factors not fully taken into account by the experimental design, namely, herbivory, artificial water provision and soil variation. soil variation between replicates in the same landscape, as well as within individual replicates, has raised the issue of the representivity of the trial. this paper provided a description and ranking of the experimental burning trial according to the geomorphic and soil characteristics of each plot in comparison to the surrounding landscape. conservation implications: the knp burn plots are one of the largest and longest-running fire experiments on fire ecology in african savannahs. however, studies need to consider the underlying geomorphic and soil template when designing experiments and interpreting results. this work describes the representivity of the plots across, and within, treatments. introduction top ↑ fire has long been recognised as an essential determinant of the structure and function of african savannahs (higgins et al. 2007; sankaran, ratanam & hanan 2008; scholes & walker 1993; trollope 1982) and is often used as a management tool within conservation agencies. the application of appropriate fire regimes is usually informed and adapted by the findings of ongoing research, which is undertaken on the experimental application of selected fire regimes on fixed areas (see e.g., andersen, cook & williams 2003; knapp et al. 1998), with treatments often repeatedly applied for many decades. the changing attitudes towards fire management within south african conservation areas – from the orthodox equilibrium theory (fixed rotational fire regimes) to one that embraces a dynamic savannah system based on non-equilibrium theory (patch mosaic fires over space and time) (bond & archibald 2003; mentis & bailey 1990; van wilgen et al. 2000; van wilgen, biggs & potgieter 1998) – has questioned the validity of results from historical long-term fixed fire experiments in influencing fire management policies (van wilgen et al. 2004; van wilgen, govender & biggs 2007). a recent critical review of the long-term fire experiment in bringing about changes in the fire management policy at the kruger national park (knp) found that it had little direct influence. however, the experiment has had numerous other unforeseen benefits, particularly its support and use in ongoing key research projects undertaken on the experiment to improve our understanding of fire as a key driver of savannah dynamics (van wilgen et al. 2007). the spatio-temporal scale and the underlying template of various fire experiments often brings into question the representivity of the results from these plot-based experiments when scaling up to landscape processes. the knp long-term fire experiment (biggs et al. 2003) is often criticised regarding the difficulty in separating the effects of the experimental fire treatments from other factors that might be causing the observed differences. the experimental burn plot (ebp) trial was designed with the initial aim of testing the effects of different seasons and frequencies of burning on the vegetation of the knp. these effects have been partly obscured by factors not fully taken into account by the experimental design, most importantly, herbivory, artificial water provision and soil variation (biggs et al. 2003). the response of the vegetation and animal population to the template presented by the geology (which is reflected by the soils) and the changes caused by the ecosystem drivers such as rainfall, hebivory and fire, have led to a complex patch mosaic template within the park (venter, scholes & eckhart 2003). to quantitatively assess the effect that the varying fire regimes have on the dominant vegetation communities, it was envisioned that vegetation surveys would be conducted annually, whilst animal life would be observed periodically (nel 1953). a comprehensive baseline vegetation (woody and herbaceous) survey was conducted prior to establishment of all treatments in 1954. vegetation re-surveys were undertaken in the late 1990s and early 2000s (van wilgen et al. 2007). during the 50-year lifespan of the experiment, various researchers conducted surveys (which were predominantly of vegetation) on the experiment (govender, potgieter & biggs 2003; van wilgen et al. 2007). webber (1979) was the first to investigate the soil characteristics on the skukuza plots; thereafter, other work looking at soil characteristics was undertaken by mills and fey (2004a, 2004b), whilst the majority of research pertaining to soils on the experiment focused on the nutrient content, cycling and variation in nutrient levels with different fire regimes (coetsee 2007; feig 2004; otter 1992; shackleton & scholes 2000; van wilgen et al. 2007). analysis, understanding and incorporation of the results from the experiment of the effects of fire on savannah systems have only been undertaken in the past decade. the van wilgen et al. (2007) paper outlines, in detail, the contribution that the experiment had to effect change in management actions and policy in the park. key findings on the biota studied on the experiment include the following (van wilgen et al. 2007): • woody tree and shrub density are unresponsive to fire regimes and fire is not critical for the maintenance of woody plant richness and composition. exclusion of fire promotes an increase in woody biomass, with the most marked effects on woody vegetation in the extreme treatments (annual burning, burning in summer or wet season, or long periods of fire exclusion). these effects were also greater in areas of higher rainfall. • manipulation of fire regimes is not critical for the maintenance of herbaceous plant species diversity. herbaceous composition changed little with fires in the dormant season, but changes were significant in the wet, growing season and with fire exclusion. impacts caused by fire were most marked in wetter areas and increased grazing pressure on the plots after the treatments influenced the grass diversity at drier sites. • there were noticeable effects of fire on small mammal communities, with unburnt sites supporting the most species and highest densities of small mammals. • bird species richness and composition did not respond to the different fire regimes. • no significant effect on ant species composition and diversity between the fire treatments but significant differences in ant assemblage composition were found between the extreme fire treatments (burnt vs unburnt plots). • regardless of fire treatment, nitrogen loss is replenished soon after the fire. frequent, annual burning increases soil crusting on the plots. mycorrhizal colonisation increases with root branching and fine root development decreases with decreasing fire frequency, allowing for optimal acquisition of resources under different fire frequencies. • fire intensity varies as a result of changes in fuel moisture content rather than post-fire age, particularly because of the seasonal differences in fuel moisture content that override those of fuel load. since the inception of the experiment in the 1950s, our understanding of the role that fire plays within savannah ecosystems continues to improve. therefore, current and future fire management policies implemented in the knp will continue to be supported by the best available science that promotes appropriate adaptive fire management. owing to the lack of baseline soil surveys or soil characterisation of the ebps, this paper provides a soil and geomorphic description of the experiment in order to determine which replicate or individual plots within replicates are not representative of the land types in which they are set. it is well known that differences in soil types cause significant differences in vegetation in the knp (venter 1990) and knowledge of the variation and extent of different types of soil and geomorphology on the ebps is therefore necessary, in order to make future studies on the plots more relevant and representative of the surrounding landscape. study site the knp was proclaimed in 1926, and covers approximately 1 948 528 ha within the low-lying savannahs of north-eastern south africa. the vegetation is characterised by savannahs within 37 landscapes (gertenbach 1983), with the dominant trees being knobthorn (acacia nigrescens), marula (sclerocarya birrea), leadwood (combretum imberbe), red-bush willow (combretum appiculatum), silver cluster leaf (terminalia sericea) and mopane (colophospermum mopane). mean annual rainfall varies from 350 mm in the north to over 700 mm in the south. geologically, the knp is underlain by granites in the west, whilst the eastern sector is underlain predominantly by basalt (venter 1990). the flora of the park comprises 1983 species, including more than 400 tree and shrub species and more than 220 grass species. experimental burn plots fire research formally began in the knp in 1954 with the establishment of one of the few long-term fire ecology research experiments in africa (van der schijff 1958). the experiment consisted of the application of fires at varying return intervals and seasons, as well as protection from fire, on a series of plots approximately 7 ha in area within specific vegetation types represented by four of the major ecological regions in the park (pretoriuskop: sourveld vegetation; skukuza: combretum vegetation; satara: knobthorn and marula vegetation; mopane: mopane vegetation)(figure 1). the treatments were replicated four times in each of these ecological regions. within each ecological region, four different replicates or strings were scattered within 20 km of each other. a double firebreak road surrounds the individual replicates (with the area between the firebreak roads burnt annually in autumn) to ensure protection from wild fires. figure 1: location of the experimental burn plots within the land types as described by venter (1990), in the kruger national park. the treatments originally included annual (b1) winter fires in august and biennial (b2) and triennial (b3) fires in august (winter), october (after first spring rains), december (early summer), february (late summer) and april (autumn). in 1976, further treatments to examine the effects of fires every four (b4) and six (b6) years in october were added to selected landscapes (satara and mopane). full details and history of the experimental design and application of treatments are available from biggs et al. (2003). description of the areas in which the knp’s fire experiment was replicated within is outlined in table 1. the associated fire treatments and plots numbers for each replicate is given in online appendix 1a–d. table 1: salient features of the four major vegetation types in which the kruger national park’s fire experiment was replicated. methods top ↑ a series of aerial photographs (digital, colour, shutter speed: 1/800, scale: 1:50 000, altitude: 914.4 m, aircraft speed: 185.2 km/h) taken in april 2000 and field surveys were used to map the land units with their associated soil and vegetation patterns on each of the plots. sections that are atypical of the specified norm were delineated and assigned a score according to the system described below (table 2). however, the survey was not conducted to identify, in detail, all soil types or to give detailed descriptions of vegetation within the plots. reference made to land types in this report is according to the definitions and descriptions of venter (1990). table 2: the allocated representivity scores and description of each score assigned to plots and replicates. soil differences usually cut through individual plots so that part of the plot may score 5, whilst another part may have a score of 1. the plot as a whole was scored depending on the dominant soil types found on it and how representative it is of the environmental parameters within each particular replicate was assessed visually. assumptions the following assumptions were made with regard to this survey:• when the ebps were laid out in the 1950s, the initiators attempted to locate the different replicates on the crests in order to avoid too much soil variation. this assumption is based on the layout of the replicates in both granitic and basaltic areas, where minor portions of mid-slope and foot-slope areas are represented in some of the plots. • the presence or absence of roads influenced the layout of the ebps as they were only laid out alongside existing roads. this negatively influenced the attempt to restrict the ebps to upland areas, so that certain portions of plots occur in foot-slope areas. this scoring system was used as a means to investigate the following questions: • how representative of the broad ecological region or landscape is each group of four replicates and series? • how comparable is each of the four replicates? this is to test if one or more of the replicates in a series are outliers, why it is regarded as outliers and whether it is enough of an outlier to warrant complete exclusion from analyses under certain circumstances. • how comparable is each the individual plots in one replicate? are there particular outliers based on any geological, geomorphologic or soil criteria? results top ↑ pretoriuskop the pretoriuskop replicates of the ebps are all fairly well representative of the pretoriuskop land type (venter 1990). the fayi replicate generally has more sandy and yellow soils with patches of avalon–clovelly and shallow hutton–glenrosa–kroonstad forms (soil classification working group 1991) dominant with many t. sericea trees (figure 2a). the downstream part of some of the plots on the shabeni replicate are influenced by seepage areas and duplex soils, namely deep clovelly form (soil classification working group 1991) (figure 2b). this replicate seems to have less tall grass species than the other pretoriuskop plots. the numbi replicate features deep, red, sandy loam soils with minor areas of duplex soils (usually on the foot slope of plots), shallow soils (steep areas) or clay soils associated with intrusions of dolerite or gabbro. a clayey area caused by the presence of some gabbro may occur, for example on plot 4 (fire treatments – online appendix 1a–d) of the numbi replicate (figure 2c). the kambeni replicate has been deeply incised by a stream (plot 11) but the influence of this is limited and is characterised by very shallow soils (escourt form) (soil classification working group 1991), caused by the steep slope in this area (figure 2d). table 3a and 3b illustrates the overall representivity scores for each replicate and individual plot within pretoriuskop. figure 2: description of the soils types on the pretoriuskop replicates (a) fayi (b) shabeni (c) numbi and (d) kambeni. for the complete list of soils names and abbreviations see online appendix 2. table 3a: the overall representivity scores for each replicate within pretoriuskop. table 3b: the overall representivity scores for each plot within the four replicates in pretoriuskop. skukuza huge granite rock outcrops and boulders occur on some of the biyamiti plots. the soils within these plots are usually hydromorphic and therefore slightly different from the rest of the plots. the south-western plots (plots 9–12) feature more clayey soils as a result of the presence of dolerite dykes in this area. these plots are generally quite different from the other plots in this replicate, as well as the series, as they are dominated by estcourt–swartlands–kroonstad and hutton–clovelly–fernwood forms (soil classification working group 1991) (figure 3a). the skukuza replicate of the ebps is characterised by quite large areas of sodic and duplex soils that occur on the foot slopes of parts of plots 6 and 7. these sodic soils have relatively thin a horizons (< 15 cm), which makes these areas completely different from the other areas in the plots. serious erosion within firebreaks is already evident where the firebreaks cross into sodic soils. unfortunately, the control or no burn plot (plot 7) (see online appendix 1a–d for treatments) at this replicate is one of those most seriously affected by the sodic soils, which covers more than half of the no burn treatment (figure 3b). the n’washitshaka replicate is characterised by reddish soils (glenrosa–hutton forms), but plot 6 is almost completely associated with hydromorphic or duplex soils (kroonstad–estcourt forms) (soil classification working group 1991) (figure 3c). the napi replicate also has some duplex soils (fernwood–kroonstad and hutton–clovelly–fernwood forms), as well as granite boulders or rocky outcrops (figure 3d). table 4a and 4b illustrates the overall representivity scores for each replicate and individual plot within skukuza. figure 3: description of the soils types on the skukuza replicates (a) biyamiti (b) skukuza (c) n’waswitshaka and (d) napi. for the complete list of soil names and abbreviations see online appendix 2. table 4a: the overall representivity scores for each replicate within skukuza. table 4b: the overall representivity scores for each plot within the four replicates in skukuza. satara it is assumed that these replicates were located to represent the s. birrea–a. nigrescens savannah of the central basalt plains. the soils and general vegetation of this zone are described as the satara land type by venter (1990). the satara replicate occurs on a very flat part of the basalt plains. the geology is sabie river basalt formation with dolerite dykes and the soils are mainly moderately deep shortlands–swartland soil (soil classification working group 1991), that are red structured clays. because of the flatness of this area, a number of shallow pans and associated black vertic soils occur. some thin, elongated, rocky outcrops associated with dolerite dykes also occur (figure 4a). the lindanda replicate (figure 4b) is representative of soils and vegetation of three different land types, namely satara, mavumbye and nwanetsi (venter 1990), as it is characterised by soils associated with both the sabie river basalt formation and the letaba basalt formation. it has been influenced by colluviation from the lebombo hills. this colluvium has influenced soils in the north-eastern corner of the replicate (portions of plots 5 and 6 and plot 7). the occurrence of more clayey soils along the south-western part of the replicate (plots 8, 9, 10, 11, 12 and 14) is ascribed to the occurrence of picrite basalts of the letaba basalt formation. this area is characterised by very clayey soils of the bonheim and arcadia forms. stunted a. nigrescens dominates this area and no s. birrea occurs. a drainage line through plot 2 is the cause of alluvial soils (bonheim and swartland) (soil classification working group 1991) and typical associated dense vegetation. the influence of the valley bottom and foot-slope extends about 50 m on either side of the drainage line. the area to the west of the drainage line, covered by plot 1 and part of plot 2 corresponds to a transition between the red, medium clayey soils associated with sabie river basalt and the black very clayey soils associated with letaba basalt. the woody vegetation on parts of plots 5 and 6 and all of plot 7 indicates that sodic soil conditions occur, for example sterkspruit, with acacia welwitschii as the dominant tree. some pans also occur in this area. these plots are therefore not completely representative of the lindanda replicate or the satara series. the area covered by significant portions of plot 2 and plots 3, 4, 5, 6 and 13 of the lindanda replicate is characterised by soils of the swartland and shortlands forms and features woody plants such as s. birrea, a. nigrescens, acacia tortilis, et cetera; it is comparable with the rest of the series. the western part of the replicate is associated with letaba basalt. the marheya replicate is considered to be representative of the satara land type, but a drainage line through plot 9 also influences plots 8 and 10 with darker, calcareous soils representative of very shallow mispah–glenrosa–maya forms (soil classification working group 1991) (figure 4c). the nwanetsi replicate is regarded as very representative of the satara land type, as most of the area is flat and characterised by red, structured, clay soils, which are characterised as moderately deep shortlands–swartland forms (soil classification working group 1991) (figure 4d). table 5a and 5b illustrates the overall representivity scores for each replicate and individual plot within satara. figure 4: description of the soils types on the satara replicates (a) satara (b) lindanda (c) marheya and (d) nwanetsi. for the complete list of soil names and abbreviations see online appendix 2. table 5a: the overall representivity scores for each replicate within satara. table 5b: the overall representivity scores for each plot within the four replicates in satara. mopane three of the four replicates are situated in the mopane shrub vegetation, with relatively shallow, calcareous, dark, clay soils associated with the picrite lavas of the letaba basalt formation. the dzombo replicate is regarded as the best replicate as it is very homogeneous with respect to soils and slope and is characterised by soils from deep milkwood and shallow bonheim forms (soil classification working group 1991) (figure 5a). two of the replicates – mooiplaas (figure 5b) and tsende (figure 5c) – have parts that represent slightly incised, sloping land where soils are shallower and the mopane has a tendency to grow taller. sometimes, drainage lines or depressions represent deeper and vertic clay soils, for example arcadia (soil classification working group 1991). on the shallow or rocky areas, different vegetation may be found, namely trees or shrubs such as terminalia prunioides, c. apiculatum and acacia exuvialis, as well as grasses such as heteropogon contortus and aristida spp. the nshawo replicate is located in an area where three types of parent material occur and where the soils and vegetation are therefore fairly heterogeneous over the replicate as a whole. the nshawo replicate is located on the interface of the letaba basalt and sabie river basalt formations, as well as on the edge of the shawu vlei area where very deep alluvial soils occur. soils associated with the letaba basalt formation are dark, calcareous clays, often shallow as on the western side of the replicate, whereas those associated with the sabie river basalt formation are reddish and usually not calcareous. on the reddish soils, trees or shrubs such as combretum imberbe and albizia harveyi may be dominant. on the shallow calcareous clays, mopane may dominate and on the alluvial clay deposits thickets of mopane occur (figure 5d). table 6a and 6b illustrates the overall representivity scores for each replicate and individual plot within mopane. figure 5: description of the soils types on the mopane replicates (a) dzombo (b) mooiplaas (c) tsende and (d) nshawo. for the complete list of soil names and abbreviations see online appendix 2. table 6a: the overall representivity scores for each replicate within mopane. table 6b: the overall representivity scores for each plot within the four replicates in mopane. discussion top ↑ present long-term studies, such as those in the serengeti (stronach & mcnaughton 1989), the knp (van wilgen et al. 2007), and yellowstone national park (romme et al. 2011) usually begin as short-term academic projects and are not envisioned to continue for decades. however, they progress over time as new events lead to further research, with the combined data and results leading to insights that were unintended and support additional questions and projects for further understanding (sinclair et al. 2007). the few long-term ecological experiments in south africa are crucial to understanding and interpreting the complex ecosystem behaviour involving slow and rapid changes at multiple ecosystem states which only became apparent over a period of several decades (thirgood et al. 2007). as previously outlined, the original objective of the experiment was to determine the effect of fire frequency and season on four of the major vegetation types in the knp (biggs et al. 2003) and therefore were spaced across the landscape to maximise the representivity of the landscape, particularly with reference to the underlying soils (van der schijff 1958). the results from this geomorphic and soil description of the individual plots and replicates in the experiment has indicated that not all plots within replicates and not all replicates within the landscape can be considered homogeneous treatments. it is for this reason that researchers undertaking their projects within the ebps, can, for statistical analyses, exclude any outlier replicates (based on the soil and geomorphic description) from their sampling protocol. however, it must also be noted that it is the outlier replicate (least representative of the surrounding landscape) that encompasses the variation and heterogeneity that is inherent within knp ecosystems and therefore it should not be completely ignored. finally, the knp’s long-term fire experiment has been instrumental in highlighting that ecosystems are not static and management should not aim to maintain the status quo; rather, it should allow natural change to take place within an adaptive management framework. this understanding could not have been achieved without the array of research and projects that have been undertaken on the ebps over the past five decades (van wilgen et al. 2007) acknowledgements top ↑ we thank the south african national parks for access to data and sandra macfadyen for the preparations of the figures. the role of the fire team that maintained the fire experiment (especially andre potgieter) is also gratefully acknowledged, as are the valuable inputs provided by harry biggs. comments from the two reviewers are also appreciated. competing interests the authors declare that they have no financial or personal relationship(s) which may have inappropriately influenced them in writing this article. authors’ contributions f.j.v. (kruger national park) undertook the geomorphic and soil survey on the experiment. he also developed the scoring system for plot and replicate representivity. the initial project report was prepared by f.j.v. (kruger national park), whilst n.g. (kruger national park) wrote the manuscript for publication. references top ↑ andersen, a.n., cook, g.d. & williams, r.j., 2003, fire in tropical savannas: the kapalga experiment, springer verlag, new york.biggs, r., biggs, h.c., dunne, t.t., govender, n. & potgieter, a.l.f., 2003, ‘experimental burn plot trial in the kruger national park: history, experimental design and suggestions for data analysis’, koedoe 46(1), 1–15. bond, w.j. & archibald, s., 2003, ‘confronting complexity: fire policy choices in south african savanna parks’, international journal of wildland fire 12, 381–389. http://dx.doi.org/10.1071/wf03024 coetsee, c., 2007, ‘a study of the interactions between fire, vegetation and nitrogen dynamics in the kruger national park’, phd thesis, department of botany, university of cape town. higgins, s.i., bond, w.j., february, e.c., bronn, a., euston-brown, d.i.w., enslin, b. et al., 2007, ‘effects of four decades of fire manipulation on woody vegetation structure in savanna’, ecology 88, 1119–1125. http://dx.doi.org/10.1890/06-1664, pmid:17536398 feig, g., 2004, ‘the effect of fire regime on plant carbon allocation and associated soil microbial community composition’, msc thesis, science faculty, university of the witwatersrand. gertenbach, w.p.d., 1983, ‘landscapes of the kruger national park’, koedoe 26, 1–121. govender, n., potgieter, a.l.f. & biggs, h.c., 2003, ‘scientific value of research conducted on the experimental burn plots in the kruger national park’, international rangelands conference 2003 proceedings, durban, south africa, 26 july – 01 august. knapp, a.k., briggs, j.m., hartnett, d.c. & collins, s.l., 1998, grassland dynamics: long-term ecological research in tallgrass prairie, oxford university press, new york. mentis, m.t. & bailey, a.w., 1990, ‘changing perceptions of fire management in savanna parks’, journal of the grassland society of southern africa 7, 81–85. http://dx.doi.org/10.1080/02566702.1990.9648211 mills, a.j. & fey, m.v., 2004a, ‘a simple laboratory infiltration method for measuring the tendency of soils to crust’, soil use and management 20, 8–12. http://dx.doi.org/10.1079/sum2003213 mills, a.j. & fey, m.v., 2004b, ‘frequent fires intensify soil crusting: physicochemical feedback in the pedoderm of long-term burn experiments in south africa’, geoderma 121, 45–64. http://dx.doi.http://dx.doi.org/10.1016/j.geoderma.2003.10.004 nel, t.g., 1953, verslag oor veldbrand [report on wildfire], south african national parks, skukuza. otter, l.b., 1992, ‘soil carbon fractionation of sand and clay soils under different burning regimes’, bsc honours thesis, science faculty, university of the witwatersrand. romme, w.h., boyce, m.s., gresswell, r.e., merrill, e.h., minshall, g.w., whitlock, c. et al., 2011, ‘twenty years after the 1998 yellowstone fires: lessons about disturbance and ecosystems’, ecosystems 91, 1–20. sankaran, m., ratanam, j. & hanan, n.p., 2008, ‘woody cover in african savannas: the role of resources, fire and herbivory’, global ecology and biogeography 17, 236–245. http://dx.doi.org/10.1111/j.1466-8238.2007.00360.x scholes, r.j. & walker, b.h., 1993, an african savanna: synthesis of the nylsvley study, cambridge university press, cambridge. http://dx.doi.org/10.1017/cbo9780511565472 shackleton, c.m. & scholes, r.j., 2000, ‘impact of fire frequency on woody community structure and soil nutrients in the kruger national park’, koedoe 43, 75–81. sinclair, a.r.e., mduma, s.a.r., hopcraft, j.c.g., fryxell, j.m., hilborn, r. & thirgood, s.j., 2007, ‘long-term ecosystem dynamics in the serengeti: lessons for conservation’, conservation biology 21, 580–590. http://dx.doi.org/10.1111/j.1523-1739.2007.00699.x, pmid:17531037 soil classification working group, 1991, ‘soil classification – a taxonomic system for south africa’, memoirs on the agricultural natural resources of south africa 15, department of agricultural development, pretoria. stronach, n.r.h. & mcnaughton, s.j., 1989, ‘grassland fire dynamics in the serengeti ecosystem and the potential method of retrospectively estimating fire energy’, journal of applied ecology 26, 1025–1033. http://dx.doi.org/10.2307/2403709 thirgood, s.j., mduma, s.a.r., keyyu, j.d. & laurenson, m.k., 2007, ‘value of long-term research: conservation science and practice in tanzania’, conservation biology 21, 576–579. http://dx.doi.org/10.1111/j.1523-1739.2007.00707.x, pmid:17531036 trollope, w.s.w., 1982, ‘ecological effects of fire in south african savannas’, in b.j. huntley & b.h. walker (eds.), ecology of tropical savanna, pp. 292–306, springer-verlag, berlin. van der schijf, h.p., 1958, inleidende verslag oor veldbrandnavorsing in die nasionale krugerwildtuin [preliminary report on wildfire research in the kruger national park]. koedoe 1, 60–92. van wilgen, b.w., biggs, h.c. & potgieter, a.l.f., 1998, ‘fire management and research in the kruger national park, with suggestions on the detection of thresholds of potential concern’, koedoe 41, 69–87. van wilgen, b.w., biggs, h.c., o’reagan, h. & mare, n., 2000, ‘a fire history of the savanna ecosystems in the kruger national park, south africa, between 1941 and 1996’, south african journal of science 96, 167–178. van wilgen, b.w., govender, n., biggs, h.c., ntsala, d. & funda, x.n., 2004, ‘response of savanna fire regimes to changing fire management policies in a large african national park’, conservation biology 18, 1533–1540. http://dx.doi.org/10.1111/j.1523-1739.2004.00362.x van wilgen, b.w., govender, n. & biggs, h.c., 2007, ‘the contribution of fire research to fire management: a critical review of a long-term experiment in the kruger national park, south africa’, international journal of wildland fire 16, 519–530. http://dx.doi.org/10.1071/wf06115 venter, f.j., 1990, ‘a classification of land for management planning in the kruger national park’, phd thesis, department of geography, university of south africa. venter, f.j., scholes, r.j. & eckhart, h., 2003, ‘the abiotic template and its associated vegetation patterns’, in j. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83–129, island press, new york. webber, n.w., 1979, ‘the effects of fire on soil/plant ecological relationships in the southern part of the kruger national park: a study on soil geography’, msc thesis, department of geography, university of south africa. article information author: alfred maroyi1 affiliation: 1department of biodiversity, university of limpopo, south africa correspondence to: alfred maroyi postal address: private bag x1106, sovenga 0727, south africa dates: received: 27 apr. 2011 accepted: 18 may 2012 published: 09 oct. 2012 how to cite this article: maroyi, a., 2012, ‘the casual, naturalised and invasive alien flora of zimbabwe based on herbarium and literature records’, koedoe 54(1), art. #1054, 6 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1054 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/ koedoe.v54i1.1054-1 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. the casual, naturalised and invasive alien flora of zimbabwe based on herbarium and literature records in this checklist... open access • abstract • introduction • research method and design    • compiling the list    • species data • results • discussion • conclusion • acknowledgements    • competing interests • references abstract top ↑ zimbabwe’s casual, naturalised and invasive alien plant species were analysed with regard to their habit, origin, mode or purpose of introduction and their invasion status in the country. this alien flora of 391 taxa belonged to 239 genera and 73 families, corresponding to 6.6% of the total flora of zimbabwe. of these, 153 (39.1%) plant species were casual aliens, 154 (39.4%) were naturalised and 84 (21.5%) were invasive species. most invasions in terms of numbers of alien species were in the central and eastern parts of the country. asteraceae (53 species), poaceae (48 species) and fabaceae sensu lato (49 species) families were prominent in all the floristic regions of the country. annual and perennial herbaceous species formed the majority of life forms of the casual, naturalised and invasive alien flora of zimbabwe. genera with the highest number of alien species were ipomoea with nine species, acacia and euphorbia with eight species each, chenopodium and senna with seven species each, eucalyptus with six species, oenothera, physalis and solanum with five species each. more than 49.6% of the alien plants in zimbabwe originated primarily from south, central and north america, followed by europe (24.6%), asia (23.8%), africa (10.5%) and australasia (5.9%). conservation implications: this research provides baseline information and historical invasion patterns of casual, naturalised and invasive alien flora in zimbabwe. this inventory is a crucial starting point in trying to understand and initiate the management of biological invasions. this is also important for monitoring new introductions and management of existing alien plants in zimbabwe. introduction top ↑ alien plant species invasions are causing major conservation problems in many regions of the world and are viewed as an important component of human caused global change (vilà, meggaro & weber 1999; vitousek et al. 1997). the spread of alien species is often associated with human activities which include disturbance and fragmentation of the natural environment, urbanisation, agriculture and the use of alien plants for landscaping and erosion control (vilà et al. 1999). the transport of organisms by humans since earliest times and now through increased levels of trade and tourism has led to the widespread breaching of natural biogeographic barriers of alien species (hulme 2009). according to vilà et al. (1999), there is a need to document naturalised flora before investigating the ecological relationship between invader and invaded habitats. documentation of alien species and analysing patterns of floristic status, biological attributes and geographical distribution has proved to be a useful approach in understanding alien species richness in various regions (e.g. khuroo et al. 2007; pyšek, sádlo & mandák 2003; pyšek & richardson 2006; rozefelds et al. 1999; stadler et al. 2000; weber, sun & li 2008).basic information on naturalised plant species, plant invasions and their impacts is lacking in zimbabwe. although occasional attention has been paid to alien plants and weeds in zimbabwe (e.g. biegel 1977, 1980; drummond 1975, 1984; mapaura & timberlake 2004; maroyi 2006; mullin 1996, 2000; timberlake, fagg & barnes 1999; wild 1955), comprehensive studies on plant invasions are still lacking. the compilation of a catalogue of casual, naturalised and invasive species, hereafter referred to as alien flora, can serve as an initial step towards further detailed studies on the impact of alien plant species on the indigenous flora and fauna. therefore, the primary objective of this investigation was to provide an overview of the alien species present, and their invasion status and geographical extent in zimbabwe, based on a literature survey and herbarium records. research method and design top ↑ compiling the list a database of casual, naturalised and invasive alien flora occurring in zimbabwe was compiled based on a literature survey and herbarium records. all alien plant species ever recorded in zimbabwe as escapes from cultivation or naturalised at least once in the wild were included in the database. species introduced or cultivated without any evidence of having escaped were not considered. alien species were divided into three categories: casual, naturalised and invasive aliens. according to pyšek et al. (2004), casual aliens reproduce occasionally outside cultivation, do not form self-sustaining populations and rely on repeated introductions for their persistence. naturalised species are defined as aliens that reproduce consistently without direct human intervention, and invasive aliens as naturalised species producing offspring in large numbers and at considerable distances from the parent plants with the potential to spread over a large area. this definition of invasive alien species used in this study is different from the convention on biological diversity (cbd) conference of parties’ definition of an invasive alien species, where an alien is defined as a species outside its indigenous geographic range, whose introduction and spread threatens biodiversity (unep 2002). the national herbarium, harare, zimbabwe (srgh), was visited on numerous occasions between 1996 and 2010. information on herbarium labels, including locality and year of collection of the plant species, was examined and recorded. species data for each taxon, information on its life history traits, such as habit, origin, year when plant species was first collected, mode or purpose of introduction and its invasion status in zimbabwe were extracted from literature and herbarium records. economic uses were included and whether the introduction was accidental or intentional. a taxon was allocated to more than one use category if different uses applied to it. a wide range of published literature and websites was consulted (see online appendix). the correct identities and names of all the taxa listed were checked and, where names have changed, the currently accepted name was applied. the species nomenclature and classification generally follow glen (2002). infra-specific taxa up to the variety level have been recognised. author citation abbreviations follow those given in brummitt and powell (1992). each taxon’s distribution in zimbabwe was indicated by letters showing the floristic divisions used in flora zambesiaca (see pope & pope 1998): northern region (n), western region (w), central region (c), eastern region (e) and southern region (s). results top ↑ the present study revealed that the casual, naturalised and invasive alien flora of zimbabwe is comprised of 391 taxa belonging to 239 genera and 73 families, representing 6.6% of the total zimbabwean flora. although this checklist is not exhaustive, it provides good baseline data for future work on casual, naturalised and invasive flora of zimbabwe. of these, 153 (39.1%) plant species were casual aliens, 154 (39.4%) were naturalised aliens, and 84 (24.5%) were invasive aliens. pteridophytes were represented by five plant species (1.3%): adiantum raddianum c. presl. (pteridaceae), azolla filiculoides lam. (azollaceae), microlepia strigosa (thunb.) c. presl (dennstaedtiaceae), microsorum scandens (g. forst.) tindale (polypodiaceae) and salvinia molesta d.s. mitch. (salviniaceae). all five species were introduced to zimbabwe as ornamentals (online appendix). gymnosperms were represented by cupressaceae and pinaceae families composed of callistris calcarata f. muell., cupressus lusitanica mill., cupressus torulosa d. don, pinus elliottii engelm., pinus patula schltdl. and cham. and pinus roxburghii sarg.; all introduced to zimbabwe at the turn of the 19th and 20th centuries, mainly for timber and as ornamentals (online appendix). dicotyledons constituted the majority of alien flora, with 323 species (82.6%), and monocotyledons contributed 58 species (14.8%). a large number of casual, naturalised and invasive alien plant species in zimbabwe (289, 73.9%) are from 16 families (table 1). plant families with the highest number of casual, naturalised and invasive alien plant species were: asteraceae (53 species), fabaceae sensu lato (49 species), poaceae (48 species) and solanaceae (21 species). the genera with the highest number of casual, naturalised and invasive alien plant species were ipomoea with nine species, acacia and euphorbia with eight species each, chenopodium and senna with seven species each, eucalyptus with six species, and oenothera, physalis and solanum with five species each.the earliest recorded dates of the alien plants in this study are based on herbarium collections (online appendix). the oldest collections date back to the late 1800s, with nicotiana tabacum l. collected in 1870, populus x canescens (aiton) sm collected in 1874, c. torulosa d.don collected in 1895, eucalyptus camaldulensis dehnh. collected in 1897 and c. lusitanica mill. collected in 1900. most of the species were first collected between 1941 and 1960 (online appendix). intensive collection of casual, naturalised and invasive alien plant species occurred between 1921 and 1980 (figure 1). nineteen twenty-one to 1940 was a period of relatively slow accumulation of casual, naturalised and invasive alien plant species records, with a total of 72 species (18.4%) collected during this period (figure 1, online appendix). most casual, naturalised and invasive alien plant species (153 in total, 39.1%) were collected between 1941 and 1960 (figure 1). the accumulation of casual, naturalised and invasive alien plant species slowed and levelled off between 1961 and 1980, resulting in the collection of 83 species (21.2%) (figure 1). table 1: families with the largest number of casual, naturalised and invasive alien species (with more than 5 taxa) in zimbabwe. figure 1: cumulative number of species against date of earliest herbarium collection for 391 casual, naturalised and invasive alien plant species in zimbabwe. the region with the highest number of casual, naturalised and invasive alien plant species was central (296, 75.7%), followed by east (279, 71.4%), west (158, 40.4%) and northern (156, 39.9%) zimbabwe, whilst southern zimbabwe (122, 31.2%) had the lowest number (figure 2). most of the species are annual and perennial herbs (figure 3). more than 49.6% originated primarily from south, central and north america (figure 4, online appendix), followed by europe (24.6%), asia (23.8%), africa (10.5%) and australasia (5.9%) (figure 4). those plant species that originated from europe, asia and africa are mostly casual aliens (figure 4). figure 2: the distribution of the 391 casual, naturalised and invasive alien plant species in zimbabwe. the numbers in the figure indicate the number of alien species in the five floristic regions of the country. figure 3: the growth forms of 391 casual, naturalised and invasive alien plant species recorded in zimbabwe. figure 4: regions of origin of alien plant flora in zimbabwe. the alien flora arrived in zimbabwe through both intentional and unintentional introductions. about 261 species (66.8%) appeared to have been intentionally introduced for various purposes (table 2). amongst these were 149 species (38.1%) introduced as ornamentals, 51 species (13%) as food plants, 31 species (7.9%) for fodder, 18 species (4.6%) for timber, six species (1.5%) for fibre, three species each (0.8%) for green manure and medicinal use, and one species (0.3%) as a crop plant (table 2). in addition, 150 species (38.4%) appear to have arrived in the country unintentionally (table 2). table 2: possible reasons for the introductions of the alien flora into zimbabwe. species may have been allocated to more than one category. discussion top ↑ the species list includes a total of 391 casual, naturalised and invasive alien plant species and infraspecific taxa is higher than 328 species listed by maroyi (2006). the current figure is higher because it utilised herbarium records, an online database (http:// www.zimbabweflora.co.zw/ [hyde & wursten 2011]) and published literature (e.g. volumes of flora zambesiaca and kirkia), unlike findings of maroyi (2006), which utilised mainly herbarium records. this figure is relatively low when compared with the 5930 indigenous zimbabwean taxa recorded by mapaura and timberlake (2004). this could be a result of under-collection, as plant collectors tend to collect mainly indigenous species, ignoring alien plants (pyšek et al. 2008). the rather small alien flora in zimbabwe confirms the findings of foxcroft et al. (2010), le floc’h, le houerou and mathez (1990) and vitousek et al. (1997) that african countries generally have few naturalised species compared to the other regions of the world. however, research by mcgeoch et al. (2010), and nuñez and pauchard (2010) showed that the number of documented invasive alien species is underestimated due to lack of information and research. the taxonomic composition of the alien flora of zimbabwe is not random and families with the largest number of species belong to the largest families worldwide, for example asteraceae, poaceae and fabaceae. a number of taxa belonging to these families and those of amaranthaceae, caryophyllaceae, chenopodiaceae and solanaceae have been recorded as weeds of agroecosystems in zimbabwe (drummond 1984; wild 1955). heywood (1989) examined the patterns and extent of invasions by agricultural weeds and concluded that most weeds come from the largest plant families like the asteraceae and poaceae. thus, it is also not surprising that annuals and herbaceous perennials are the prevalent life forms amongst the alien flora recorded in this study, as it has been shown in other studies that the herbaceous habit is most frequent amongst the invasive species (daehler 1998; kolar & lodge 2001). the early phase of exotic plant species introduction involved early settlers who arrived and settled in major towns like harare and gweru (zimbabwe central), bulawayo (zimbabwe west) and mutare (zimbabwe east). historically, colonists from europe brought food plants, ornamentals, structural timber species, et cetera, with them during their migration (maroyi 2006; mullin 1996). according to maroyi (2006), small-scale plantings of alien plant species in zimbabwe started with european settlement in the 1890s and early 20th century. this was a period when acclimatisation societies and botanical gardens flourished in the british colonies and economic plants were transferred from one continent to another (maroyi 2006). over 80% of the alien flora in zimbabwe originated from america, europe and asia. alien plants were introduced into zimbabwe from these regions probably through the european colonisers. the relatively lower number of alien species in the southern part of zimbabwe is due to the harshness of the region’s climate. southern zimbabwe is hot and dry, characterised by low and erratic rainfall. the majority of alien plant species recorded in zimbabwe have been introduced intentionally for economic purposes, although a sizeable proportion of species arrived in the country by accident. several species of eucalyptus were introduced from australia for structural timber, furniture, poles, charcoal and medicinal oils (mullin 1996). about 38.1% of the casual, naturalised and invasive alien plant species in zimbabwe were introduced into the country as ornamentals. similar studies in other countries have shown that most invasive plants are ornamentals (reichard & white 2001). previous research has shown that whilst many herbaceous ornamentals have been accidentally introduced, woody ornamentals have been almost entirely introduced intentionally, mainly for landscaping (reichard & white 2001). the accidental introduction of alien plant species into zimbabwe is also due both to the use of contaminated seeds in agricultural systems and as a result of human transport systems. nowadays, intensive trade and the constant flow of people have resulted in a large plant transfer, crossing frontiers and geographical boundaries (del monte & aguado 2003). an understanding of the establishment and distributional changes of alien flora over time is important for making informed decisions in managing existing introductions and predicting future invasions. although the zimbabwean alien flora has been noted for some time (wild 1955), it is only in the last four decades that a profound interest has developed, owing to the important economic impacts of alien weed species on agricultural crops (drummond 1975, 1984). the majority of the species (63.2%) considered in this investigation were already casuals, naturalised or invasives in zimbabwe by 1955, when an initial survey of exotic weeds was carried out (wild 1955). about 30 alien species were recognised as weeds by wild (1955). drummond (1975) reported 45 naturalised woody plant species. drummond (1984) systematically recorded 62 alien species that grew as weeds in croplands, causing important economic damage in productive lands. the period between 1921 and 1980 was characterised by increased study of the zimbabwean flora, which boosted awareness and recording of new invaders. equally intriguing is the fact that the following ten notable introduced species classified as invasive aliens in this study were not documented until the 1960s: a. filiculoides (azollaceae), bidens bipinnata (asteraceae), browallia americana (solanaceae), coreopsis lanceolata (asteraceae), erigeron karvinskianus (asteraceae), jacaranda mimosifolia (bignoniaceae), myriophyllum aquaticum (haloragaceae), persicaria lapathifolia (polygonaceae), rubus niveus (rosaceae) and solanum mauritianum (solanaceae). the number of areas an alien weed species occupies increases with the number of years since the first record of naturalisation. biological invasions are made up of several stages, which include introduction, naturalisation and spread. previous research by binggeli (2001) has shown that there are time-lags between the introductions of exotic plants, their establishment and subsequent spread. it has since been established that only a small proportion of introduced species are capable of spreading and becoming weeds (kowarik 1995). for example acacia mearnsii (fabaceae-mimosoideae), cajanus cajan (fabaceae-papilionoideae), solanum betaceum (solanaceae), mangifera indica (anacardiaceae), nicotiana glauca (solanaceae), passiflora edulis (passifloraceae), p. patula (pinaceae), prunus serotina var. salicifolia (rosaceae) and syzygium jambos (myrtaceae) were recorded as naturalised only in eastern zimbabwe by drummond (1975), but this investigation showed that they are also naturalised in other regions of the country. information on naturalisation or rate of spread of alien plants increases with more collecting and monitoring efforts. a better understanding of their establishment and distributional changes is important for the development of appropriate alien plant species management plans. conclusion top ↑ the catalogue of casual, naturalised and invasive alien plant species presented in this study is a critical starting point in trying to understand and initiate the management of biological invasions; and it will serve as a foundation for future research on plant invasions in zimbabwe. however, the documentation of casual, naturalised and invasive plant species in zimbabwe is far from sufficient. it is here recommended that extra attention be paid at studying the ecological and socio-economic impacts of introduced plant species on the native communities in zimbabwe. a better understanding of their establishment, spread, social and economic impacts, and distributional changes over time is vital for making informed decisions on new introductions and managing existing casual, naturalised and invasive alien plant species. since complete eradication of introduced species is unachievable (moles et al. 2012), there is need to have a historical invasion patterns of alien species and their long-term effects on native communities. this field of study is currently under researched in zimbabwe and there are few examples where casual, naturalised and invasive alien plant species have been fully evaluated. the result is that, where investigations are done at all, research on such alien plants are often short-term, highly localised, sporadic, largely ad hoc and restricted to field studies without underpinning agricultural, scientific or economic studies. therefore, elucidation of alien plant naturalisation patterns in zimbabwe will prove crucial in the scientific management of casual, naturalised and invasive alien flora in the country. acknowledgements top ↑ the head of the national herbarium and botanic garden, harare, zimbabwe (srgh) and staff are thanked for granting access to their facilities, helpful advice and plant identifications. i am also grateful to four anonymous reviewers for constructive comments and sandra macfadyen for figure 2. competing interests the author declares that he has no financial or personal relationship(s) which may have inappropriately influenced him in writing this paper. references top ↑ biegel, h.m., 1977, checklist of ornamental plants used in rhodesian parks and gardens, rhodesia agricultural journal research report no. 3, department of research & specialist services, harare.biegel, h.m., 1980, common and botanical names of garden plants in zimbabwe, zimbabwe agricultural journal research report no. 4, department of research & specialist services, harare. binggeli, p., 2001, ‘time-lags between introduction, establishment and rapid spread of introduced environmental weeds’, paper presented at the iii international weed science congress, international weed science society, corvallis, 6–11 june 2000, viewed n.d., from http://members.multimania.co.uk/woodyplantecology/docs/timelag.rtf brummit, r.k. & powell, c.e., 1992, authors of plant names, royal botanic gardens, kew. daehler, c.c., 1998, ‘the taxonomic distribution of invasive angiosperm plants: ecological insights and comparison to agricultural weeds’, biological conservation 84, 167–180. http://dx.doi.org/10.1016/s0006-3207(97)00096-7 del monte, j.p. & aguado, p.l., 2003, ‘survey of the non-native plant species in the spanish iberia in the period 1975-2002’, flora mediterranea 13, 241–259. drummond, r.b., 1975, ‘a list of trees, shrubs and woody climbers indigenous or naturalized in rhodesia’, kirkia 10(1), 229–286. drummond, r.b., 1984, arable weeds of zimbabwe: a guide to the recognition of the more important weeds of crops, agricultural research trust of zimbabwe, harare. foxcroft, l.c., richardson, d.m., rejmánek, m. & pyšek, p., 2010, ‘alien plant invasions in tropical and sub-tropical savannas: patterns, processes and prospects’, biological invasions 12, 3913–3933. http://dx.doi.org/10.1007/s10530-010-9823-7 glen, h.f., 2002, cultivated plants of southern africa, jacana press, national botanical institute, pretoria. heywood, v.h., 1989, ‘patterns, extents, and modes of invasions by terrestrial plants’, in j.a. drake, h.a. mooney, f. dicastri, r.h. groves, f.j. kruger, m. rejmánek & m. williamson (eds.), biological invasions: a global perspective, pp. 31–60, wiley, new york. hulme, p.e., 2009, ‘trade, transport and trouble: managing invasive species pathways in an era of globalization’, journal of applied ecology 46, 10–18. http://dx.doi.org/10.1111/j.1365-2664.2008.01600.x hyde, m.a. & wursten, b., 2011, flora of zimbabwe, viewed 05 january 2012, from http://www.zimbabweflora.co.zw khuroo, a.a., rashid, i., reshi, z., dar, g.h. & wafai, b.a., 2007, ‘the alien flora of kashmir himalaya’, biological invasions 9, 269–292. http://dx.doi.org/10.1007/s10530-006-9032-6 kolar, c.s. & lodge, d.m., 2001, ‘progress in invasion biology: predicting invaders’, trends in ecology & evolution 16, 199–204. http://dx.doi.org/10.1016/s0169-5347(01)02101-2 kowarik, i., 1995, ‘time lags in biological invasions with regard to the success and failure of alien species’, in p. pyšek, k. prach, m. rejmánek & m. wade (eds.), plant invasions, general aspects and special problems, pp. 15–38, spb academic publishers, amsterdam. le floc’h, e., le houerou, h.n. & mathez, j., 1990, ‘history and patterns of plant invasion in northern africa’, in f. d’castri, a.j. hansen & m. debussche (eds.), biological invasions in europe and the mediterranean basin, pp. 105–133, kluwer academic publishers, dordrecht. mapaura, a. & timberlake, j. (eds.), 2004, a checklist of zimbabwean vascular plants, southern african botanical diversity network report no. 33, sabonet, pretoria. maroyi, a., 2006, ‘preliminary checklist of introduced and naturalized plants in zimbabwe’, kirkia 18(2), 177–247. mcgeoch, m.a., butchart, s.h.m., spear, d., marais, e., kleynhans, e.j., symes, a. et al., 2010, ‘global indicators of biological invasion: species numbers, biodiversity impact and policy responses’, diversity and distributions 16, 95–108. http://dx.doi.org/10.1111/j.1472-4642.2009.00633.x moles, a.t., flores-moreno, h., bonser, s.p., warton, d.i., helm, a., warman, l. et al., 2012, ‘invasions: the trail behind, the path ahead, and a test of a disturbing idea’, journal of ecology 100, 116–127. http://dx.doi.org/10.1111/j.1365-2745.2011.01915.x mullin, l.j., 1996, eucalyptus in zimbabwe, kirkia 16(2), 95–107. mullin, l.j., 2000, conifers in zimbabwe, kirkia 17(2), 199–217. nuñez, m.a. & pauchard, a., 2010, ‘biological invasions in developing and developed countries: does one model fit all?’, biological invasions 12, 707–714. http://dx.doi.org/10.1007/s10530-009-9517-1 pope, g.v. & pope, d.g., 1998, collecting localities in the flora zambesiaca area, flora zambesiaca management committee, royal botanic gardens, kew. pyšek, p., richardson, d.m., pergl, j., jarošík, v., sixtová, z. & weber, e., 2008, ‘geographical and taxonomic biases in invasion ecology’, trends in ecology and evolution 23, 237–244. http://dx.doi.org/10.1016/j.tree.2008.02.002 pyšek, p. & richardson, d.m., 2006, ‘the biogeography of naturalization in alien plants’, journal of biogeography 33, 2040–2050. http://dx.doi.org/10.1111/j.1365-2699.2006.01578.x pyšek, p., richardson, d.m., rejmanek, m., webster, g.l., williamson, m. & kirschner, j., 2004, ‘alien plants in checklists and flora: towards better communication between taxonomists and ecologists’, taxon 53, 131–143. http://dx.doi.org/10.2307/4135498 pyšek, p., sádlo, j. & mandák, b., 2003, ‘alien flora of the czech republic, its composition, structure and history’, in l.e. child, j.h. brock, g. brundu, k. prach, p. pyšek, p.m. wade & m. williamson (eds.), plant invasions: ecological threats and management solutions, pp. 113–130, backhuys, leiden. reichard, s.h. & white, p., 2001, ‘horticulture as a pathway of invasive plant introductions in the united states’, bioscience 51, 103–113. http://dx.doi.org/10.1641/0006-3568%282001%29051%5b0103:haapoi%5d2.0.co;2 rozefelds, a.c.f., cave, l., morris, d.i. & buchanan, a.m., 1999, ‘the weed invasion in tasmania since 1970’, australian journal of botany 47, 23–48. http://dx.doi.org/10.1071/bt97054 stadler, j., trefflich, a., klotz, s. & brandl, r., 2000, ‘exotic plant species invade diversity hot-spots: alien flora of north-western kenya’, ecography 23, 169–176. http://dx.doi.org/10.1111/j.1600-0587.2000.tb00272.x timberlake, j.r., fagg, c. & barnes, r., 1999, field guide to the acacias of zimbabwe, cbc publishing, harare. unep (united nations environmental programme), 2002, alien species that threaten ecosystems, habitats or species, viewed 23 december 2011, from http://www.cbd.int vilà, m., meggaro, y. & weber, e., 1999, ‘preliminary analysis of the naturalized flora of northern africa’, orsis 14, 9–20. vitousek, p.m., d'antonio, c.m., loope, l.l., rejmánek, m. & westbrooks, r., 1997, ‘introduced species: a significant component of human-caused global change’, new zealand journal of ecology 21, 1–16. weber, e., sun, s.g. & li, b., 2008, ‘invasive alien plants in china: diversity and ecological insights’, biological invasions 10, 1411–1429. http://dx.doi.org/10.1007/s10530-008-9216-3 wild, h., 1955, common rhodesian weeds, salisbury, government printer. filelist convert a pdf file! supplement to koedoe. 1977: 255-259. suid-afrika se nasionale parke sisteem wat van die toekoms? sy edele h schoeman* minister van landbou republiek van suid-afrika abstract the significance of the 50th anniversary of the national parks board of trustees is highlighted and homage is paid to all those who have contributed to the development of the national parks system in the republic of south africa. the government never lightly refuses requests on sustained pressures for new national parks and establishment of a new national park near beaufort west in the karoo, is announced. mnr die voorsitter, uitgelese gaste, dames en here. u sal my toelaat om reg by die begin verskoning te maak dat sy edele minister h schoeman nie aanwesig kan wees nie weens omstandighede buite sy beheer. u sal my ook toelaat om sommer nou aan die voorsitter 'n woordjie toe te voeg van 'n belofte wat hy gemaak het en dit is toe hy ons uitgenooi het na hierdie funksie, dat ek hier kan kom en op my gemak kom sit en die tydjie in die wildtuin net kan geniet. hy het die belofte egter vergeet, ek is seker daarvan. maar, weens omstandighede buite die beheer van sy edele die minister, moet ek nou vandag hier optree soos wat dit in engels gestel word "i am deputising, true to the real meaning of what a deputy minister means". u weet minister schoeman het van tyd tot tyd gese 'n adjunk-minister is 50% van niks en ek dink hy is nie baie ver uit nie! mnr die voorsitter, u sal my dit vergun dat ek vir u gelukwens met u persoonlike verjaardag. ons weet dat dr knobel 'n besondere vindingryke persoonlikheid is. ek wil hom nie blameer en ek glo nie iemand hier in die gehoor sal hom blameer, dat hy dit so bewerkstellig het dat die nasionale parkeraad se verjaardag saamval met sy eie nic. maar ek wil dit darem net as 'n opmerking laat dat ek nie 'n enkele amptenaar ken, maak nie saak van watter status, wat so vindingryk is om te sorg dat soveel uitgelese gaste sy verjaardag bywoon nie! ek wil hom gelukwens daarmee en daarmee gaan dit ook na mev knobel. ons weet dat sy hom beslis in alles bystaan en beslis veel kleur gee aan dit wat hy doen. ons hoop dat hulle hierdie jaar wat voorle, baie sal geniet en dat u ook vee i *referaat voorgedra deur sy ed ele j j malan, adjunk-minister van landbou. 255 sukses sal he met u werk hierso as hoofdirekteur van die nasionale parkeraad. waar ek namens minister schoeman hier optree wi! ek dan ook hoop dat u sover 'n besondere vrugbare simposium gehad het en as ek geluister het na wat dr. martiny hier as opsomming aan ons voorgehou het, is ek jammer dat ek nie die hele tyd hier kon gewees het nie. dit wil my voorkom onder andere, en dan is ek heeltemal objektief, dat iemand uit ons departement landbou-tegniese dienste minstens gesorg het dat die boer se standpunt hier gestel is en dat u u voete op die grond hou. dit is vir my baie interessant toe dr. martiny gepraat het, en waar hy dit so mooi gestel het dat daar selfs van owerheidswee meer gedoen moet word, kon ek egter nie help as om daaraan te dink dat dit darem eienaardig is dat mense nie die essensie van dinge snap nie. wanneer 'n demokratiese land soos ons s'n aan mense eise stel, verwagtinge na vore bring, dan se hulle by implikasie, ons wil dit he, en ons is gewillig om daarvoor te betaal. dit is ook waar ek 'n probleem het met die regering. u weet ek het drie besware teen die regering t.w.: as jy hulle nader dan het hulle probleme in die eerste instansie met finansies, in die tweede instansie het hulle probleme met verkryging van fondse en in die derde instansie het hulle probleme met geld! dit is vir my 'n eer om vandag hier namens my minister op te tree by hierdie besondere geleentheid. dit is vir my dan ook aangenaam om sy toespraak hier aan u voor te lees. die bereiking van so 'n belangrike mylpaal as die 50ste verjaardag van die totstandkoming van die liggaam soos die raad van kuratore vir nasionale parke is ongetwyfeld 'n trotse gebeurtenis. dit getuig van bestendigheid en dat die liggaam tot groot hoogte die taak wat aan hom opgedra is suksesvol uitgevoer het. geen liggaam wat nie redelikerwys voldoen aan die eise wat aan hom gestel word, sal 50 jaar lank kan bestaan en gedy nie. op 'n dag soos hierdie is dit paslik om 'n terugblik te werp in die verlede. by so 'n terugblik is dit baie duidelik dat daar veral rede is tot groot dankbaarheid, maar ook om trots te kan wees. soos ook elders vandag by 'n ander geleentheid staan ons in groot ootmoed en erkentlikheid voor die herinnering van die historiese figuur paul kruger. aan hom kom die hulde toe vir sy grootmoedigheid en gebruiklike versiendheid om in die pionierstyd van die ontwikkeling van ons land en volk reeds 'n gebied af te sonder wat die natuur in sy ongeskonde staat bewaar en sou beskerm. hy staan natuurlik nie aileen in die opsig nie, maar aan die voorpunt van 'n lang ry mense, wat groot bydraes gel ewer het tot die totstandkoming en ontwikkeling van hierdie en ons ander nege nasionale parke. daar was lede van die regering van ons land wat van tyd tot tyd groot entoesiaste van natuurbewaring was en wat deur regeringsoptrede die ontwikkeling van die stelsel van ons nasionale parke bevorder het. ek verswyg liewer hulle name omdat dit seker is dat wanneer ek hulle probeer noem die name van sommige van hulle onregverdiglik weggelaat sal word. 256 mr chairman, we cherish the memory of many private individuals who have already passed away but who during their lifetimes were enthusiastic nature lovers and contributed to the development of our national parks by the grant of land, money and other gifts. it is gratifying to know that also the present generation has not been lacking in displaying the true participation in our company here today and who have also made many material and substantial contributions both in money and in respect of services rendered towards the further development of our national parks. it is only because i realise that they would not like to advertise the fact that they are serving the country also in this respect, that i refrain from mentioning their names. we remember the many persons who have served as members of the national parks board of trustees. some of them with distinction. it is a fact that the board has created an image of itself which is looked upon by many people with such high respect that they regard an appointment to the board as the highest achievement possible. dit is interessant om net daaraan te dink hulle se dit word oorvertel in die wandelgange van die parlement " ... that the definition of a national park is a place where people are sent when they are tired and need to relax, but appointment to the board is for those who have spent their lives there!". ons neem vandag afskeid van een van die dienende lede van die parkeraad. hy is mnr tiny faure. hy het met onderskeiding en met groot toewyding as lid van die parkeraad gedien vir nie minder as 26 jaar nie. oom tiny sou seker nog graag voortgegaan het as lid van die parkeraad as dit nie was dat hy noodwendig sy aktiwiteite wil begin inkort nie. ek se dankie aan hom vir sy lange jare van uitmuntende diens as lid van die raad en wens hom die allerbeste toe op die pad vorentoe. ons is jammer om te hoor dat mev. faure nie so wel is nie en ons wil graag dit wat ons van oom tiny se, dat hy dit aan haar sal oordra. ons is dankbaar ook teenoor haar vir wat sy gedoen het en ons wens is dat sy geheel-en-al sal hers tel. ons dink ook aan die aandeel van die amptenare van die raad oor die jare. die van hulle wat vandag nog in die diens van die parkeraad is, diegene wat die parke geskep het, waarop ons so trots is en wat aan ons volk en aan ons mense asook mense van oor die hele wereld so 'n unieke vreugde en genot verskaf. ons wil ook met groot waardering verwys na die verskillende staatsdepartemente en ander owerheidsinstansies wat oar die jare steeds uiters simpatiek opgetree het wanneer hulle met sake van die nasionale parkeraad te doen gekry het. hierdie welwillendheid het in 'n aansienlike mate die totstandkoming en ontwikkeling van ons nasionale parke moontlik gemaak. mr chairman, we regard what has been achieved in the field of nature conservation in our country with justifiable pride. nature conservation in what may be termed national parks, is rightly the function of the national parks board of trustees. national parks to my mind are 257 necessary to protect and nourish the continued existence of objects of nature for the benefit of the nation, where such conservation bears a national character and calls for action on a national scale. enthusiasts from the ranks of the board and its officials and other nature lovers are continually pressing for the creation of further national parks, or for the extension of their areas, or for improvements of the facilities in existing national parks. i wish to assure them that the government never lightly refuses these requests in this regard and i am sure the chairman will bear me out on that. hoe graag ookal die regering hoogsprioriteit sa! wil verleen aan aansoeke om ondersteuning van die nasionale parkeraad is hy nogtans soms verplig om sulke aansoeke af te wys. dit gebeur in elk geval nie dikwels nie. daarmee moet egter rekening gehou word dat aansprake op die land se bates, veral in die vorm van grond en geld geweldig is, en dat dit al hoe moeiliker word om selfs die verdienstelikste voorstelle te bevredig. dit is vir my as minister van landbou ook 'n voorreg om te kan getuig dat in die landbou, van die grootste gedeelte van die oppervlakte van ons land wat landboukundig aangewend word, die belangrikheid van bewaring van die gawes van die natuur in die vorm van landbouhulpbronne oor die jare heen al hoe meer besef word. vandag is juis hierdie aspek een van die belangrikste onderdele van die boerdery. ek maak die stelling dat geweldig baie reeds gedoen word om die gawes wat die natuur ons vaderland bied, te beskerm en te bewaar. ook is daar groot vordering gemaak met die inskerping van 'n nasionale bewustheid van die waarde van ons natuurlike hulpbronne by die publiek. terwyl ons erken dat baie meer nog gedoen moet word , hoef ons land nie skaam te wees vir wat reeds bereik is en wat tans onderneem word nie. moontlik moet ondersoek ingestel word of daar met inagneming van beskikbare fondse nie uitbreidings aan die volgende aksies bewerkstellig kan word nie . ek noem hulle: (a) ek dink dat dit noodsaaklik is om te kyk tot watter mate die bergkwagga nasionale park, die bontebok nasionale park en die tsitsikamabos en -seekus nasionale park vergroot kan word. dat dit nodig is, is seker. of dit gedoen kan word, dit moet die toekoms bepaal. (b) die stigting van nuwe nasionale parke om meeste van die volgende belangrike gebiede te bewaar: die rigtersveld; sommige voeleilande en robeilande aan ons kuste; die kosibaai-kompleks; deel van die sederberge; deel van die drakensberg eskarpement; 'n namakwaland blommepark; die bobbejaanskloof; 'n grot nasionale park en dan natuurlik 'n karoo nasionale park. dit is dan met groot vreugde dat ek kan aankondig dat die kabinet in beginsel goedkeuring verleen het aan die stigting van 'n karoo nasio258 nale park by beaufortwes. die park word moontlik gemaak deur 'n wonderlike skenking van grond op die dorpsmeent deur die munisipaliteit van beaufort-wes. die munisipaliteit word vandag hier verteenwoordig en ons se baie dankie vir die besondere bydrae deur 'n enkele munisipaliteit vir 'n skenking wat 'n groot nasionale bate gaan word. tot hierdie grond word bygevoeg grond wat bygekoop word deur gelde voorsien deur die suid-afrikaanse natuurstigting wat bykans rloo 000 beloop asook 'n bedrag uit die staatskis van rioo 000. die suidafrikaanse natuurstigting wat hier verteenwoordig word, moet spesiale vermelding kry omdat hulle dit reggekry het om gelde van die algemene publiek en veral skoolkinders vir die doe! in te same!. ek dink dit was 'n besonderse poging. ons wil die stigting en deur hulle aan almal wat bygedra het tot die karoo parkefonds baie dankie se. ons vertrou dat die wonderlike aksie wat hulle begin het, voortgesit sal word en dit die algemene publiek meer en meer sal aanspoor om grond vir nasionale parke te verkry, sodat die raad nie net na die staat hoef te kyk vir die verkryging van fondse nie. die gronde bly 'n volksbesit en dit gee my dus groot vreugde om op die 50ste bestaansjaar van die nasionale parkeraad die stigting van die lode nasionale park te mag aankondig. ek sluit af met die gedagte, mnr die voorsitter, dat die nasionale parkeraad veral kan trots wees op wat hy tot stand gebring het. die hele land kan trots wees op wat tot stand gebring is en kan spog met dit wat die raad opgebou het. ons is dankbaar daarvoor en ons vertrou dat die raad op die pad vorentoe net sovee! sukses sal he as op die pad wat reeds agter ie. mag die ideale wat vir die toekoms gekoester word, almal bewaarheid word. baie dankie. 259 page 1 page 2 page 3 page 4 page 5 a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe reviewer acknowledgement 88 koedoe african protected area conservation and science koedoe african protected area conservation and science recognises the value and importance of the peer reviewer in the overall publication process – not only in shaping the individual manuscript, but also in shaping the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely, productive peer review process. we would like to take this opportunity to thank all reviewers who participated in shaping this volume of koedoe: alex sliwa angela gaylard anna spenceley berndt j. van rensburg campbell murn chris sandbrook colleen t. downs craig t. symes danny govender dave i. thompson edward witkowski frauke dogley graeme shannon graham alexander kevin p. kirkman leslie r. brown m. ernest daemane marietjie landman we appreciate the time taken to perform your review successfully. i i in an effort to facilitate the selection of appropriate peer rievewers for koedoe, we ask that you take a moment to update your electronic portfolio on www.koedoe.co.za for our files, allowing us better access to your areas of interest and expertise, in order to match reviewers with submitted manuscripts. if you would like to become a reviewer, please visit the journal website and register as a reviewer. to access your details on the website, you will need to follow these steps: 1. log into the online journal at http://www. koedoe.co.za 2. in your ‘user home’ [http://www.koedoe. co.za/index.php/koedoe/ user] select ‘edit my profile’ under the heading ‘my account’ and insert all relevant details, bio statement and reviewing interest. 3. it is good practice as reviewer to update your personal details regularly to ensure contact with you throughout your professional term as reviewer to koedoe. please do not hesitate to contact me if you require assistance in performing this task. rochelle flint submissions@koedoe.co.za tel: +27 (0)21 914 5100 fax: +27 (0)21 914 5105 mark alderweireldt michelle d. henley mike rutherford nick zambatis norbert hahn petri viljoen phyllis lee pieter bester priscilla burgoyne richard h. taylor rudy jocque sarel s. cilliers stefan siebert sue j. milton theo mostert timm m. hoffman willem j. myburgh willie j.l. coetzee filelist convert a pdf file! supplement to koedoe. 1977: 230-242. marine conservation in the republic of south africa with special reference to marine parks and reserves g a robinson park warden: tsitsikama forest and coastal national parks p a storms river 6308 introduction conservation in the republic of south africa (rsa) is dominated by big game and terrestrial reserves. protecting these, have been in existence for many years. recently man has turned his attention more and more to the sea and due to adverse effects of pollution, over-exploitation and habitat destruction the marine environment is threatened. however, conservationists have learnt lessons about both terrestrial and marine ecosystem-destruction and are now demanding with increasing persistence the establishment of marine parks and reserves. it is therefore not surprising that the iucn's marine conservation programme, is to be given special emphasis during the two-year period, 1977 and 1978. with the establishment of a ministerial committee to investigate marine parks and reserves south africa has not lagged behind. the committee's objectives are to promote marine conservation by the probable establishment of a system of marine parks and reserves. the rsa has an exceptionally long and beautiful coastline (2 961 km). its west coast is lapped by the cold benguela current which originates in the antarctic and supports one of the most productive fisheries in the world. on the east coast the shores are bathed by a current which arises in the indian ocean, the warm agulhas current, and gives a tropical flavour to the marine life along this coast. the intermediate region or "south coast" lies between these two spheres of influences and has representatives of both warm water and cold water communities living side by side (fig. 1). a wide spectrum of marine fauna and flora is therefore, encountered along the shores of south africa. marine conservation in the past "conservation" means the wise use of resources. in a sense, man has conserved (used) the biological resources of the sea since prehistoric 230 ~ l>:> atlantic ocean fig. 1. existing marine parks and reserves in south africa ~ benguela current (cold) ''''',,-'" ;; j) -~:t-' ;'~;"~'il ~, \ \ co~'ow ,,( • s\ 0 " & / i . so,th coa" · : <? ~_~ c;::----~ ~;:~ ;/ //~ l;;/ {j~ (1 l / ~ indian ocean times. the normandic aborigines of the "south coast", the strandlopers, lived predominantly off the coast for thousands of years (goodwin 1946). at first he collected mainly shellfish, as seen from the numerous strandloper middens found along the coast, but he later built enclosures in the tidal zone in which he trapped fish. the enclosure is a stone wall built to such a height that the high tides will cover the enclosed area allowing fish to swim into it. as the tide recede, the fish were trapped and caught by hand. these aborigines probably conserved the marine resource more intelligently than our modern "civilized" society is apparently doing he lived in harmony with his environment. the value of the marine resources has in more recent times been recognised and the need for conservation fully appreciated. biological and environmental research has been conducted by the government in order to rationally exploit the fishery resources for many years. the regulatory agency in the earlier days was known as the fisheries and marine biological survey and was directed by dr j f gilchrist. later this conservation agency became the division of sea fisheries and has recently become the sea fisheries branch. this branch is responsible for regulating and conserving the exploitation of all marine biological resources with maximum sustainable yield as objective. the south african fishing industry is dominated by the pelagic west-coast fishery and the trawl fishery, but has also a valuable rock lobster fishery. it is, therefore, understandable that most of the research effort has been directed towards the species supporting these fisheries. regulatory measures have been imposed on by the branch to conserve the resources. these include the mesh-size of nets, gear to be used or not used, size limits, bag limits , closed seasons, quotas and sanctuaries. these sanctuaries were proclaimed to protect species such as rock lobsters , oysters and sea-horses. the sea fisheries branch employs a large staff of inspectors at a considerable expense to enforce thes e regulations. this entails the inspection of catches, the hounding of rock lobster poachers and patrolling (both sea and land) of the sanctuaries. marine conservation at present in the past marine conservation went through two phases . the first, when man lived in harmony with the sea, he used the marine resources and they were conserved. but there was very little " intelligent" exploitation done. the second phase of marine conservation was the rational exploitation on a maximum sustainable yield basis. here man used his intelligence to devise in some cases sophisticated, mathematical models whereby regulating his harvest. marine conservation implies the wise use of the marine resources and by this it does not only imply use in terms of human consumption. wise use must also incorporate aesthetic and recreation values. the estab232 lishment of marine parks and reserves is then the third phase of marine conserva tion. participants at the first world conference on national parks in seattle washington, in 1962, after discussing underwater preservation, passed resolution no. 15 that stated: "whereas it is recognized that the oceans and their teeming life are subject to the same dangers of human interference and destruction as the land, that the sea and land are ecologically independent and indivisible, that population pressures will cause man to turn increasingly to the sea, and especially to the underwater scene, for recreation and spiritual refreshment, and that the preservation of unspoiled marine habitat is urgently needed for ethical and esthetic reasons, for the protection of rare species, for the replenishment of stocks and valuable food species, and for the provision of undisturbed area for scientific research. the first world conference on national parks invites the governments of all those countries having marine frontiers, and other appropriate agencies, to examine as a matter of urgency the possibility of creating marine parks or reserves to defend underwater areas of special significance from all forms of human interference and further recommends the extension of existing national parks and equivalent reserves with shorelines, into the water to the ten fathom depth or the territorial limit or some other appropriate off-shore boundary." this recommendation gave considerable momentum to the third phase of marine conservation and the establishment of several parks and reserves in various parts of the world resulted. during more recen t years meetings such as the second world conference on national parks in 1972, the international conference on marine parks and reserves in tokyo 1975 and the 12th general assembly of the iucn, held in zaire 1975, have greatly emphasized the need for marine conservation and manne reserves. three out of 12 general resolutions passed at kinshasa, zaire in september 1975 by the general assembly related to marine conservation read as follows: resolution number 2 calls on governments to establish marine parks and reserves of sufficient area to protect the vulnerable features of vulnerable marine habitats; and recommends to governments with terrestrial parks or reserves adjoining coastal areas of value for nature conservation that they extend the boundaries of those parks to include a marine component. 233 resolution number 3 recommends that areas as large as possible of sea-grass, coral reefs and mangroves be set aside as reserves for research and the protection oflife. urges the coastal states with such habitats within their jurisdiction to evolve a policy to maintain their continued health and stability, and urges the research and monitoring programmes in such areas be strengthened or initiated to establish guidelines for their management. resolution number 4 requests governments to take urgent steps to protect and manage key areas, large enough to form viable ecological limits, containing mangroves, salt marshes and related wet-lands together with adjacent estuarine waters as reserves for the preservation of these ecosystems and the species for which they provide habitat. from the above it is transparently clear that internationally there is a growing concern regarding the state of the marine environment. there is a restless feeling amongst conservationists time is running out. threats to marine resources why the haste to establish marine parks and reserves? the reason is that many marine species and habitats are today threatened . this is due to man's irrational approach to the conservation of these natural resources. we are all, by and large, fond of the coast but the sea or ocean appeals to but a few of us. of these few, fewer still appreciate it as an environment. the ocean is valued asa resource to be exploited and misused. but let us look at the causes that threaten our marine environment. foremost in most marine situations is pollution. from the earliest times, man has used the sea as a dum.ping place for his refuse. it was generally accepted to be a "bottomless pit". when our waste was still mostly organic matter, this dumping was of little consequence but now, with inorganic chemicals, pesticides, and radio-active wastes, they are not easily disposed of. there are some that become extremely concentrated in filter-feeding animals such as bivalves. oysters are known to store contaminants in their tissues at concentrations 70 000 times greater than in their surrounding water. although these animals may themselves be in a healthy condition they can be lethal to animals feeding on them. when one speaks of pollution, sewage and oil spillage is usually thought of. pollution is not always unsightly, sticky or smelly. nuclear power stations for instance pump heated effluent into the sea which can and do disrupt the marine environment. secondly, over-exploitation of these resources by fishermen can cause serious problems. i t is hardly necessary to repeat the tragic history of the exploitation of the whales (chapman 197 i) to illustrate the illogical 234 .,,"'---' i i i i i i i i i \ \ \ \ \ \ \ , , , ... ••••••• 0 •• oaoe.ooco _-------_-------~8 ... .... .......... ' ................... ... ~.... -.~---( -----------------------------------_.----235 / ,/ i i i i i o • o o • o e • · • o q • • o o '" '" n '" "' ., :" > 00 m • · '" • !" n m c.) r·· .. 0 ("'-j ('0 i.':-~ r:: (0 ") .c': o +j <'l u ro fa ,j.: r: 0) .. :; ('j w way in which man has over-killed one species after another (fig. 2). fortunately species are only exploited until it becomes uneconomical to harvest them, they are then left alone. u nfortuna tely they have to then fend for themselves in what probably is an unstable environment. thirdly, development of coastal towns, roads, industries and resorts have devastating abilities of not only shrinking the available natural coastal environment, but also to change the surrounding areas. lastly, there is the relentless destruction caused by uncontrolled tourist activities. these include spear-fishing, shell-collecting, baitcollecting and collection of living specimens. these activities which together, even if they do not exterminate or endanger entire species, may yet cause serious ecological disruptions. marine turtles are endangered through over-harvest of adults and unlimited collection of eggs and the loss or reduction of suitable nesting areas by the activities of man (hughes 1976). purposes of marine parks and reserves an area managed along sound ecological principles is highly significant for man's ultimate understanding of the dynamics of a particular ecosystem. therefore, an area protected solely for the reason of preserving it in its pristine condition, is relevant to the study of the natural processes, and to the protection of endangered species and habitats from man's parturbations. r eserves have, however, four main purposes, depending on th eir use (pollard 1976): (i) the conservational purpose of marine reserves would imply a sanctuary for the preservation of a commercially exploited species or the protection of a habitat; (ii) the scientific purpose of marine reserves would be to protect an area in its pristine condition as a reference area or to monitor a disturbed area or manipulate it in order to gain an understanding of the ecosystem; (iii) the educational purpose of a marine reserve would be the interpretation of the ecology of the area to both students and the general public to instill a general appreciation for the marine conservation; (iv) the recreational purpose of marine reserves, if managed along sound ecological lines, would not include the entire spectrum of man's enjoyment possibilities . it would only include his more passive recreational activities such as snorkelling, underwater photography, animal watching (birds, whales, dolphins, fish and invertrebrates), or aesthetic values. existing marine parks and reserves the rsa can only boast with three marine reserves in the sense that they are protected for aesthetic, educational and recreational purposes 236 (fig. 1) . only one of these is spacious enough to have been proclaimed a marine national park. the tsitsikama coastal national park is a 67 km stretch of shoreline, mainly rocky, but has a few sandy beaches and numerous riverlets running into the sea. proclaimed in 1964, this unspoiled rugged coast is administered by the national parks board of trustees (robinson 1969). it is situated along the south coast and therefore has a wide spectrum of marine life covering warm tropical and subtropical indo-pacific and cold atlantic and sub-antarctic species. the park extends 0,8 km out into the sea and everything is protected including flotsam and jetsam, dead trees, pebbles and rocks . controlled angling is, however, allowed at specific well demarcated places. the objectives for this park are: (i) the conservation of a south coast marine environment in its pristine state; (ii) research on an ecosystem orientated basis; and (iii) marine ecology interpretation and education. the park is zoned and only half of the coastline is accessible to the public for recreational purposes in the form of hiking, birdwatching, botanising and snorkelling. the h f verwoerd reserve is a small sanctuary close to cape town. although only two kilometres in length, it represents the west coast limit of the south coast , supporting a fine example of a kelp-bed community with huge macrophytes, rock lobsters and perlemoen. the sanctuary is administered by a local authority in co-operation with the sea fisheries branch (heydorn and hughes 1969). all marine life, both flora and fauna is protected with the exception that controlled angling is allowed. the sardinia bay marine r eserve, situated on the eastern limit of the south coast near port elizabeth. the sanctuary protects only the marine fauna and does not allow fishing of any kind along its fiv e kilometre shoreline (grindley 1974). it is important in that it supplements the tsitsikama national park in having good representation of the more tropical and sub-tropical south coast species. the point to stress here is that only the south coast is represented in the above three reserves. apart from the above "true" reserves the sea fisheries branch has created an entire system of sanctuaries and protected areas in order to manage the commercially important species. there are exceptions to this as in the case of the knysna sea-horse hippocampus capensis. this system is rather involved and table 1 is hopefully sufficient to indicate what this protection entails. 237 table 1 existing marine reserves and protected areas i. national marine parks tsitsikama coastal national park 2. nature marine reserves i sardinia bay reserve ii h.f. verwoerd reserve (bet ty ' s bay) iii buffalo bay and ro b berg reserves iv knysna lagoon 3. guano islands (government permits required for admission.) 4. commercial fish reserves i si. helena bay, saldanha bay, table bay and luderlitz bay ii pal grave point to kunene river (s.w.a.) iii algoa bay (hougham park) protected organisms all marine life (flora and fauna) controlled angling is permi tted in a few specified areas. all marine fish (" fish" includes all marine fauna) marine life excluding angling fish bait organisms seahorses, pipefish and seaweeds all marine life and seabirds, although seals & guano are exploi ted. rock lobsters pelagic fish oysters 5. areas closed for specific fishing gear i all commercial gear perle moen (a belone) ii all ncts all fish iii all nets iv all nets v ncts except purse seines vi purse seine nets vii trawl nets (bottom/mid water) viii gill nets (set/drift/staked) sharks yellowtail all fish mull e t and pelagic fish west coast rock lobsters all fish number of areas 2 40 (including islands off s.w.a. coast) 4 1 5 2 3 9 combined coastline nautical miles 36 5 2 12 6 40 47 220 0,2 20 36 200 130 ii 115 163 645 6. remarks about sixty species or specific groups of organisms are protected partially or entirely. completely protected organisms include seafans, dolphins, and six different whale species. (information mainly from de villiers, pers. comm.) 238 proposed marine parks and reserves marine national parks as previously stated the rsa has three distinct coasts, only one of which is represented by the tsitsikama coastal national park. it is therefore desirable that a spacious stretch of coastline be set aside to represent each of the other two coasts. a namaqualand marine national park, if proclaimed, would represent the cold benguela current system of the west coast. it should be situated between the orange river mouth in the north and the olifants river in the south. hopefully the entire coastline could be proclaimed as such. where private property is adjacent to this coast, the terrestrial boundary will obviously be the high water mark. the coast is dominated by dune and sandveld but representatives of rocky shore and kelp-bed areas should also be included . a tongaland marine national park has been proposed as a marine reserve by many biologists (heydorn and hughes 1969, grindley 1975) and every effort should be made to establish it as such. it is probably the most beautiful coast along south africa. besides preserving the eastcoast in its pristine condition, the nesting area of turtles could be protected (hughes 1976). a de hoop marine national park would represent a unique example of the cape shores (including its weathered sandstone shores, fish species as well as relevant cape macchia. as such a likewise addition would imply an excellent contribution to the spectrum of national parks). in addition to these, five islands should also be given national status. a dassen island national park which would represent an excellent example of a west-coast island system with both a rich avi-fauna as well as a rich interand subtidal fauna. a dyer island national park would represent a south-coast island environment with a unique variety of birds visiting the island. a seal island national park, a small rocky out-crop with a healthy cape fur-seal arctocephalus pusillus population situated in mossel bay. a bird island national park (algoa bay) a 25 ha island in algoa bay supporting a population of some 50 000 gannets sula capensis. the marion national park (antarctic). if proclaimed it would protect an unique environment. ecosystem research would be fundamental to its management. 239 n ~ .... 0 fig. 3. proposed marine parks and reserves in south africa \ r :;f1 910 1 namaqualand marine national park 2 olifants ri ver estuary 3 verlorenvlei estuary 4 langebaan lagoon and saldanha bay tslands 5 dassen island national park 6 bokbaai coast 7 robben island 8 cape point coast 9 dyer island national park 10 quoin point coast 11 de hoop marine national park 12 seal island national park 13 goukamma coast 14 knysna western head 15 noetsie coast 16 st croix island 17 bird island national park 18 trafalgar impanjati coast 19 umdoni park 20 peace cottage 21 jex estate 22 umlalazi inyoni coast 23 leven point coast 24 tongaland marine national park 11 ills .-.. 16)7 )" 20 " / i 18 4; ). ") 000000 11'1 .., 2 :!! ~ km provincial marine parks and reserves the following list are proposed marine reserves and if established will afford endangered marine species and habitats better protection. (this list endorses the list by grindley 1975). (a) west coast (see fig. 3) olifants river estuary, verlorenvlei estuary, langebaan lagoon and saldanha bay islands, bokbaai coast, robben island and cape point coast (b) south coast quoin point coast, goukamma coast, knysna western head, noetsie coast (eastern knysna head to robberg) and st. croix island (c) east coast (the ciskei and transkei coasts have been omitted) trafalgar-impanjate coast, umdoni park, peace cottage, jex estate, umlalazi-inyone coast and leven point coast. conclusion if the rsa is to protect its marine environment from the adverse effects of pollution, over-exploitation and habitat destruction, drastic action is needed. the establishment of marine parks and reserves can no longer wait. it is urgent that they be proclaimed immediately. south africa has been a leader in terrestrial nature conservation since the establishment of the kruger national park. let us prove that we are not only sympathetic towards marine nature conservation but also develop a rational and practical system for the protection of the marine environment for future generations. 241 references chapman, d g 1971. estimation of population parameters of antarctic baleen whales. quantitative science paper no . 22. university of washington. mineograph. goodwin, a] h 1946. prehistoric fishing methods in south africa . antiquity 20: 134-141. grindley, ] r 1974. the establishment of the sardinia bay marine reserve. e. cape naturalist 54: 2-4 grindley, ] r 1975. the need for the establishment of marine reserves in south africa. proceedings if the council for the habitat symposium on coastal conservation. durban. april. heydorn, a e f, and g r hughes. 1969. urgent need for more marine reserves. afr. wildl. 23 (3):270-278 hughes, g r 1976. sea turtles in south east africa . proceedings if a symposium on endangered wildlife in southern africa . university of pretoria (81-87). mineograph. pollard, d a 1976. international conference on marine parks and reserves, tokyo, may 1975. australian marine science bulletin 52: 7-11. robinson, g a 1969. the tsitsikama coastal national park, south africa. bioi. cons. 2 (i): 72-73. 242 page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 article information author: susan snyman1,2 affiliations: 1environmental economics policy research unit, university of cape town, south africa2wilderness safaris, johannesburg, south africa correspondence to: susan snyman postal address: po box 5219, rivonia 2128, south africa dates: received: 01 feb. 2013 accepted: 30 nov. 2013 published: 24 june 2014 how to cite this article: snyman, s., 2014, ‘assessment of the main factors impacting community members’ attitudes towards tourism and protected areas in six southern african countries’, koedoe 56(2), art. #1139, 12 pages. http://dx.doi.org/10.4102/ koedoe.v56i2.1139 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. assessment of the main factors impacting community members’ attitudes towards tourism and protected areas in six southern african countries in this original research... open access • abstract • introduction    • conceptual framework and literature review • research method and design • results    • reasons given for the importance of conservation    • probit models for tourism and protected area attitudes • ethical considerations • trustworthiness    • reliability    • validity • discussion • conclusion • acknowledgements    • competing interests • references abstract top ↑ in southern africa, many early conservation efforts from the late 1800s and early 1900s either displaced local communities or restricted their access to natural resources. this naturally affected community attitudes towards protected areas and efforts were later made to rectify growing tensions. in the last few decades of the 20th century, these efforts led to conservation and ecotourism models that increasingly included communities in the decision-making and benefit-sharing process in order to garner their support. although the results of these policies were mixed, it is clear that the future success of conservation and, consequently, ecotourism in many areas will depend on the attitudes and behaviour of communities living in or adjacent to protected areas. managing and understanding community expectations and attitudes under varying socio-economic circumstances will lead to more efficient, equitable and sustainable community-based conservation and ecotourism models. this study was based on 1400 community interview schedules conducted in botswana, malawi, namibia, south africa, zambia and zimbabwe, allowing for an accurate comparison of attitudes across countries, protected areas and communities. the results highlighted important demographic and socio-economic factors to consider in terms of understanding the attitudes of those living in and around protected areas. suggestions were put forward for managing community relationships and garnering long-term support for protected areas and ecotourism.conservation implications: it was observed that, in general, community members living in or adjacent to conservation areas in southern africa have an understanding and appreciation of the importance of conservation. formal education was found to positively impact attitudes and human–wildlife conflict negatively impacted attitudes, highlighting important policy focus areas. introduction top ↑ it is clear that sustainable african conservation has to include local populations (hoon 2004; musumali, larsen & kaltenborn 2007). many have argued that if communities receive benefits from protected areas (pas) and ecotourism they will tend to hold positive attitudes towards conservation as a land use and to care for the natural resources in their area (currie 2001; emptaz-collomb 2009; hulme & murphree 2001; mcneely, in groom & harris 2008; wang & pfister 2008; waylen et al. 2009). ecotourism allows local communities to receive benefits from conservation, both directly, through wages and salaries, and indirectly, as suppliers of goods and services. numerous authors have argued that community support is critical to the long-term success of ecotourism operations and their associated pas (alexander 2000; allendorf et al. 2006; chandralal 2010; gillingham & lee 2003; makindi 2010; nepal 2002; newmark et al. 1994; sifuna 2010). an analysis of community attitudes towards pas across southern african would highlight the determinants of community support. such information may encourage governments and ecotourism operators to focus on these factors in policy decisions. although pas and ecotourism are interlinked, community attitudes towards them may be sensitive to the direct costs and benefits of ecotourism, particularly employment and human–wildlife conflict. managing community expectations of ecotourism requires an understanding of the factors driving their attitudes. education and awareness-raising programmes also benefit from an understanding of community attitudes (allendorf et al. 2006; chidakel 2011; sifuna 2010; simelane, kerley & knight 2006). as emerton (1999) pointed out, benefit distribution is a necessary, but not sufficient, condition for communities to engage in wildlife conservation (snyman 2012a). the understanding of local residents’ attitudes, by the managers of pas and eco-lodges, will naturally affect their interactions with the community and it is hoped that this understanding will allow more focused planning in pas and associated ecotourism operations. the specific objectives of the present study were to, (1) enrich the tourism and pa literature with an analysis of factors impacting rural community members’ attitudes to tourism and pas across six southern african countries and (2) provide useful policy options and practices for pa and ecotourism managers. from these objectives three research questions emerged: • what demographic and socio-economic factors drive rural community members’ attitudes towards tourism and pas? • how do rural community members’ attitudes differ across six southern african countries? • what policy implications emerge from an understanding of factors impacting community members’ attitudes? conceptual framework and literature review following allport and mcdougall and munro (both in wang & pfister 2008:85) we shall treat ‘attitudes’ as ‘respondents’ particular feelings and perceptions towards the stated questions relating to tourism and/or conservation’. wang and pfister (2008) found that examinations of attitudes towards tourism showed that these are influenced by people’s values and personality and are therefore slow to change. they are also influenced by factors invisible to outsiders (allendorf et al., in waylen et al. 2009) making them difficult to observe or understand.in addressing individual community members’ attitudes, social exchange theory assumes that potential beneficial outcomes will create positive attitudes towards tourism (andereck et al. 2005; teye, sönmez & sirakaya 2002). the theory postulates that individuals perceiving net benefits from an exchange are likely to view it positively and those perceiving net costs are likely to view it negatively (snyman 2012a). this approach is supported by numerous studies which showed that residents dependent on tourism for support, or who perceive it offering net personal benefits, tend to view its impacts more positively than others (brunt & courtney and child, in groom & harris 2008; various sources, in andereck et al. 2005:1061; shibia 2010; walpole & goodwin 2001; wang & pfister 2008). the fact that people with lower household incomes are often less supportive of pas and tourism is largely explained by needs theories. such theories argue that an individual’s basic needs are attended to first before higher needs such as supporting community, conservation or tourism initiatives (doyal & gough and maslow, in emptaz-collomb 2009). emptaz-collomb (2009) stresses that it would therefore be prudent for companies or individuals engaged in such initiatives to assist in improving the lives of local people through improving access to and the quality of education, health, transport and communication. in the long run, this would create a more supportive environment for tourism and conservation projects and ensure their sustainability. government, non-governmental organisations (ngos) and the private sector can all play a role in this through infrastructural and other development projects. the importance of understanding attitudes lies in their theoretical connection with behaviours (manfredo et al., in lepp & holland 2006). the links between attitudes and behaviours are, however, not automatic (scanlon & kull 2009). even though nunkoo and ramkissoon (2011) found that residents in their mauritian study engaged in behaviours congruent with their attitudes, this will not always be the case. although past research has shown that many communities hold positive attitudes towards conservation (alexander 2000; currie 2001; de boer & baquete 1998; mehta & heinen 2001; sekhar 2003; weladji, moe & vedeld 2003) and ecotourism development (chandralal 2010; lepp 2007; mehta & kellert 1998), there are a number of different factors affecting attitudes. some studies (gadd 2005; lepp & holland 2006; rodriguez 2008) have also found that communities, for varying reasons, may hold negative attitudes towards conservation. some studies have also not found a correlation between economic benefits from tourism and positive attitudes to conservation (stem et al. 2003; walpole & goodwin 2001) or suggested that economic benefits alone were insufficient to encourage conservation (stem et al. 2003; stronza & pégas 2008). stem et al. (2003) and stronza and gordillo (2008:450) found that non-economic benefits, such as new skills, broader experiences in managing projects and people, ideas exchange, expanded circles of contacts, empowerment and support for community efforts, could also influence attitudes towards pas and tourism. these non-economic benefits are, however, often more difficult to measure and assess. creating and maintaining positive attitudes towards pas is especially important when other mechanisms for changing behaviour, such as regulation, are inappropriate or ineffective (waylen et al. 2009). over the years, regulation has often failed to ensure conservation success and the resultant need for a more appropriate and effective means of ensuring conservation has arisen. in summary, studying community members’ attitudes towards tourism and pas is important for a number of reasons (snyman 2012a): • it can disclose whether or not negative attitudes exist towards a pa and/or tourism operation, which, in some cases, may help to explain behaviour (lepp & holland, in anthony 2007). • it can inform policymakers and managers about which factors influence attitudes and this can assist with prioritising avenues for action (anthony 2007; browne-nuñez & jonker 2008). • it can reveal opportunities for improving relationships and outreach programmes with communities living adjacent to pas (anthony 2007). • it can assist in developing appropriate benefit-sharing and cost-minimising programmes for communities based on their attitudes and experiences. • it can highlight areas important for education and training programmes. overall, there is widespread consensus that community attitudes matter and these attitudes may vary over time and be influenced by various factors (e.g. changing incomes, land management arrangements, etc.) (andereck & vogt 2000; anthony 2007; gillingham & lee 1999). factors such as the level of education, recent experiences and degree of politicisation may also play a role. in an attempt to capture the relevant issues, this article looks at the impact of a number of socio-economic variables on community attitudes across different countries. despite the many attitude studies that have been conducted, they are not always comparable as they have been conducted under different circumstances and with different measurement tools. this study was however conducted consistently over six study countries allowing for a more accurate comparison of attitudes across countries, pas, tourism operations and communities. research method and design top ↑ in this study (adapted from snyman 2012a, 2013) extensive socio-economic interviews were conducted in over 30 rural communities, covering more than 16 different ethnic groups and an average of approximately 25% of households in each study village (ranging from 10% to 84%). the selection of study sites was dictated by the presence of a community–ecotourism relationship or partnership, or because the community lived in or adjacent to the conservation or pa where the ecotourism operation was located, or a combination of these (in this study all ecotourism operations were owned or managed by wilderness safaris, see http://www.wilderness-safaris.com). conservation areas need not be government owned, but can include areas owned and/or managed by communities, private individuals, ngos or companies (makindi 2010). the common feature of the areas covered in this study is that all have been specifically set aside for conservation. at one extreme, this involved the total relocation of local people formerly living in the area (e.g. the makuleke community in south africa); at the other extreme are conservation areas in which people still live and have access to the natural resources (e.g. the namibian conservancies). few of the study sites had accurate or recent maps of households, dictating other means of sample size selection. sample sizes varied with the size of the community being surveyed. an attempt was made to interview at least 10% of households in all adjacent communities. logistically, however, this was not always possible. if the survey area was small, then the majority of the households in the area were interviewed. if the survey area was large, then for logistical reasons, a subset of villages and households was selected. household numbers were obtained from the latest census figures and/or from the respective headman, chief or community trust. households were selected randomly, either by walking through the village interviewing every second household (or the next household where someone was available) or interviewing a household member in a public area, for example, a local shop or meeting area. in total, 1400 community interviews were conducted in rural villages either within or adjacent to pas (see table 1). attempt was made to interview the household head, but if they were not available, then either the spouse, or next oldest person in the household was interviewed. table 1: details of communities surveyed in each country. as the concept of community is central to the analysis in this article the term needs to be clarified. for the purposes of this article, we follow borrini’s (as cited in borrini-feyerabend, kothari & oviedo 2004:9) description of it as ‘a human group sharing a territory and involved in different but related aspects of livelihoods – such as managing natural resources, producing knowledge and culture, and developing technologies and practices’, whilst a local community is a group who interact regularly or who influence one another’s daily lives. in this study, such local communities may be mobile, permanently settled or semi-nomadic such as the himba people of north-west namibia. the communities are found living either within or adjacent to the pa, or in some cases, having left the pa, are now living further afield (e.g. the makuleke community which is located two hours’ drive from pafuri camp, but is still impacted by it). all communities described in this article are either directly or indirectly affected by the conservation and ecotourism strategies in their area, whilst their activities, in turn, impact nearby pas and ecotourism operations. the camps associated with the study communities were all ecotourism camps in terms of their location and the activities offered. defining ecotourism and its relationship to conservation is also contextually important at this stage. in summarising the literature, de witt, van der merwe and saayman (2011:1139) suggest that the key principles of ecotourism are that it should foster a genuine interest in nature, contribute to conservation, respect and conserve local culture, make non-consumptive use of natural resources, yield benefits to the local community and create tourist awareness of conservation and local community issues. based on this definition, ecotourism in this article is taken to include activities which are nature-based and culture-based, sustainable, promote conservation and provide benefits to local people in the area. it is therefore not simply tourism that is based on the sale of access to an interesting natural area, but tourism that also provides benefits to local communities (snyman 2013). the interviews were conducted by both male and female interviewers and local translators were used in circumstances where the respondent could not speak or understand english. the interview schedules contained questions relating to demographics, social welfare and living standards, education, employment patterns, income and expenses, health and safety and attitudes toward tourism and conservation. this article focuses on the sections relating to attitudes. each interview was conducted verbally, with the interviewer completing the questionnaire during the interview. each interview took approximately 20 min – 45 min depending on the respondent’s educational level and whether or not translation was required. the interview schedule consisted of a structured set of questions, with the majority being close-ended and some having the option for further explanation. communities associated with a single ecotourism enterprise, wilderness safaris (ws), were surveyed in this study because it was the only ecotourism company that had parallel ecotourism operations, operating according to a standard policy framework, over the six anglophone countries in the region. the use of a single company made for ease of comparison because the head office imposes a consistent management style over its different camps in southern africa. the company itself wished to quantify the impact of its ecotourism operations on rural communities and gave the author access to its camps and staff and to the communities with whom they engage. the process followed does, however, mean that some caveats attach to this research: • although the camps and communities were diverse (with varying land management systems, ethnic groups and tourism camp price ranges), as only one ecotourism operator was included in the analysis, there could be limitations to the generalisability of the research. • the author was employed by ws to conduct a study on the impact of the company’s activities on rural communities. this study was, however, performed as an independent researcher looking to discover the realities of ecotourism and community development and was in no way influenced by the company. • local residents would have associated interviewers with ws because of the use of ws vehicles in some areas and through the introduction process. this may have biased responses to questions about ws. it is however impossible to predict the direction of the bias a priori; some respondents may have been strategically negative in order to ensure changes or positive in order to win favour with the private sector operator in the area (allendorf et al. 2006). the results showed both positive and negative responses in all areas and many respondents were clearly comfortable expressing negative responses. • the presence of the researcher during the administration of the interview schedule may have influenced some respondents and their answers to attitudinal questions regarding tourism and conservation. the bulk of the interview schedule was, however, socio-economic in nature and this should not have proven problematic. there remained the risk of strategic bias. when it was felt that this was occurring, the author re-iterated to the respondent that the interview was anonymous and honest answers were required. all data collected were analysed using spss version 12 (2004) and stata version 10.2 (2008), as well as a combination of descriptive statistics, t-tests and probit models. results top ↑ table 2 summarises the respondents’ demographic details in order to provide context to the other results. these results are both aggregated across the southern african region and, in some cases, disaggregated by country to show specific country nuances. table 3 shows the results for the tourism and conservation attitudes by location. overall, the zambian community respondents held the most positive attitudes to tourism and conservation. this is most likely because there were few employment opportunities in the area other than ecotourism or with the zambian wildlife authority. table 2: summary of respondents' demographic characteristics. table 3: attitudes towards tourism and conservation by sampled communities. the importance of tourism and conservation in these remote, rural areas is highlighted by the percentage of respondents (45%) who had family members working in tourism and/or conservation. this result does, however, need to be interpreted with caution in terms of total employment impact. there is a risk of double-counting, as many village respondents are related and a number of separate households may have been referring to the same employed family member. in all, 87% of community respondents felt that conservation was important, despite 83% of them having problems with wild animals. reasons given for the importance of conservation respondents who felt that conservation was important were asked for their reasons. a number of different rationales emerged (see table 4). many respondents said that it was important for tourism. other reasons included: for their children and/or the future and to be able to use the natural resources in the future for food, firewood, et cetera. some respondents said because the ‘trees bring rain and/or prevent wind’. some respondents said that conservation was important for tourism, but not for people’s crops or livestock and others said that it was important to conserve trees and plants, but not animals because they were dangerous. a number of respondents gave more than one reason and others said that they knew it was important (often because they had been told at school), but did not know why. in some areas, respondents said it was important because the government said so. table 4: main reasons given for the importance of conservation by location. probit models for tourism and protected area attitudes probit models were run for each of the attitude questions. the demographic variables in the probit were chosen as a result of a descriptive statistical analysis and an extensive literature review of factors found to impact community attitudes. table 5 presents the probit results, reporting marginal effects for each attitude question. table 5: probit results reporting marginal effects for each attitude question. although factors affected attitudes differently, there was a trend observed in terms of the impact of education on attitudes, with respondents reporting higher levels of formal education being more positive. this was statistically significant in four of the questions. those who were more educated also had fewer problems with wild animals.statistically, demographic variables had varying impacts on attitudes. age was however, surprisingly, positively related, suggesting that older respondents were more positive about tourism and pas than younger respondents. with respect to age of the household head however, this relationship was largely negative, with respondents with older household heads being less positive about tourism and pas. areas with higher population densities were shown to be less positive about tourism and pas. distance had mixed impacts on attitudes, as did the number of years that the tourism camp had been operating. respondents with more household income sources tended to be more positive about tourism having a positive change in their village, as well as about conservation being important. a consistent issue raised in all study communities was that of wild animals interfering with households’ livelihoods and, in some cases, personal safety. one female respondent in malawi told of a night she had been terrified inside her house with her children, whilst a bull elephant tried to push the house over to reach her maize stored inside. this highlights the real threat wildlife can pose to rural households and their livelihoods. table 6 shows that across all communities surveyed, 83% of respondents had problems with wild animals at home. table 6: percentage of respondents who had problems with wild animals by location. elephants were the most frequently mentioned animals (55%) that caused problems, followed by lions (28%). other animals mentioned included cheetah, hippo, leopard, hyaena, baboon and jackal. in general, human–wildlife conflict (hwc) resulted in less positive attitudes towards ecotourism in the study areas. respondents who had family employed in tourism or conservation were generally observed to be more positive towards tourism and pas and had fewer problems with wild animals. having family employed in tourism or conservation positively impacted four attitude questions and could be because unemployment and poverty levels are high; having a family member employed in tourism or conservation therefore has a significant impact on a household and therefore on attitudes. the country variable was significant in the ‘reducing poverty’ question only, indicating that, in general, communities across southern africa did not differ significantly in their attitudes towards tourism and pas. ethical considerations top ↑ the interview schedule was approved by the university of cape town’s ethics committee. all respondents were told the reason for the study and signed a written consent form prior to the interviewer beginning the interview. respondents were told that the interview schedules were confidential and their participation in answering all questions was voluntary. this resulted in some questions not being answered. non-response to questions did not cluster on particular questions, as no particular question had a greater non-response rate than any other question. trustworthiness top ↑ although every effort was made to ensure the interviews and data analysis were conducted correctly, certain assumptions and limitations of the research necessarily arise, which should be factored into the data analysis and interpretation. reliability cross-cultural research can have limitations in terms of respondents reacting or answering differently because of the presence of an expatriate working in a developing country that was previously colonised (bruyere, beh & lelengula 2009). it was hoped that the use of local translators (who received informal training from the author) would help minimise this limitation. the author and other interviewers (where not local) also made efforts to learn some of the local languages and customs, to greet respondents in their own language and to help them feel relaxed. de boer and baquete (1998) warned that formal questionnaires become a drawback when people are unwilling to express negative opinions or attitudes to a third party, in particular where interviewees are reluctant to confess to illegal exploitation practices, such as snaring or collecting plants in a restricted area. questionnaires are, however, a cost-effective method of research. this needs to be kept in mind when analysing the data collected on opinions and attitudes to tourism and conservation in the present study. that negative attitudes were expressed by some respondents suggests that they did not feel constrained. in order to minimise bias and inaccurate information, respondents were also informed at the beginning of the interview that it was anonymous, was part of a research study, and that their responses would be aggregated and impossible to identify in the larger study. interviews run a number of other risks including the researcher leading the respondent, variation in the delivery of the survey between interviewers, respondent anticipation or desire to please the researcher and discrepancies between what people report and what they actually do or feel (de boer & baquete 1998; gadd 2005). despite this, surveying attitudes and quantifying them is necessary if one is to compare attitudes towards conservation and ecotourism in different regions or within the same region over time (gadd 2005). every effort was made to keep the interviews uniform and to ask questions in such a manner as to reduce bias or at least keep it consistent. in order to render any existing bias relatively constant, the author conducted over 1000 of the interviews herself. eight other interviewers assisted across the six countries. validity the sample size was not standardised in all study countries because of logistical constraints in some areas where communities were large. these differences in the percentage of the community interviewed could result in some issues relating to external validity. it was however felt that all sample sizes were sufficient in the areas surveyed and no new information was found after a certain percentage (approximately 100 households) of the community had been interviewed. the zambian sample is included in the analysis because, despite being small, it was felt to be relevant and representative of the area where the interviews were conducted. inferences drawn from the zambian results should however be viewed with caution. discussion top ↑ the present study analysed the impact of various demographic and socio-economic variables on attitudes of community members in six southern african countries and, as in past studies, (allendorf et al. 2006; simelane et al. 2006) found mixed results relating to various factors. individual’s attitudes, although generally long term, can change over time and also vary between and within different communities and countries. there are a number of factors that cannot be controlled or manipulated, so it is important that policy is aimed at those factors that can, to some extent, be influenced – for example, education, land ownership, et cetera. in general, community members in this study felt that tourism creates employment and can help reduce poverty. if tourism-related jobs were to disappear in an area (for example in the zambian study area where tourism is one of few employers), it is unlikely that people would feel as positive about tourism and pas in the future. this highlights the importance individuals attach to tourism in terms of employment, household income, et cetera. it is critical that this is managed in an appropriate manner to ensure that expectations are met and that there is not large-scale disappointment on the part of communities. this is highlighted by the fact that in communities where tourism had been operational for longer, respondents were less positive about conservation than those areas where tourism was new. allendorf (2007) highlighted that although there has been little attention given to the importance of non-economic benefits that residents may value in developing countries, studies indicate that people do value pas for non-economic reasons, such as ecosystem services, for the benefit of future generations and the conservation of wildlife. this was supported by the findings in the present study. although many respondents felt that conservation was important because of the income to be derived from it through employment and tourism, many felt that it was important for their children and future generations, as well as for the wood, thatch and food it provides. as found in the present study, various other studies have found differing results relating to the impact of demographic variables on attitudes towards tourism and conservation (akyeampong 2011; allendorf et al. 2006; baral & heinen 2007; currie 2001; de boer & baquete 1998; gadd 2005; gillingham & lee 1999; kideghesho, røskaft & kaltenborn 2007; larson 2010; mbaiwa & stronza 2011; mehta & heinen 2001; sarker & røskaft 2010; sekhar 2003; shibia 2010; stem et al. 2003; tessema et al. 2007; teye et al. 2002; weladji et al. 2003). it appears, therefore, that it is difficult to use demographic variables to predict attitudes. there are, however, some areas of commonality in terms of demographics that can be used as potential predictors. other studies (anthony 2007; currie 2001) have largely found that younger respondents hold more positive attitudes towards tourism and pas. this may be because many younger people have more education than older people, are less reliant on natural resources and therefore less affected by a lack of access to them and/or have alternative livelihoods that reduce the risk that they face. age was significant only in the ‘conservation important’ attitude, with a positive coefficient, indicating that as age increases, there was an increase in the predicted probability of having a positive attitude. this contradicts past research relating to the impact of age. the number of children was only significant in the ‘problem animal’ attitude question, with a positive relationship. this could be because respondents with more children perceived more problems with wild animals, as any conflict would negatively affect them as they had more mouths to feed. communities living in or adjacent to pas frequently incur direct costs associated with living next to wildlife, such as damage to crops, loss of livestock and, occasionally, loss of human life. other costs associated with wildlife include the additional direct cost of guarding crops and livestock by paying someone to look after them or the opportunity cost of protecting them by giving up one’s time which could have been put to a more productive use. this often results in a disruption to children’s schooling as they are kept out of school to guard household fields (pers. obs. author, december 2009). logic implies that those who are negatively impacted by wildlife are likely to have less positive attitudes towards conservation and consequently ecotourism. this is detrimental to the long-term success of pas and ecotourism and therefore requires attempts to mitigate hwc in rural areas. in countries where there was a high percentage of respondents who had problems with wild animals combined with less positive attitudes (e.g. malawi), it is even more important that the surrounding communities receive benefits from ecotourism and conservation, or measures to mitigate hwc. hill (2004) argues that people who believe that they do not have control over a conflict situation are often more likely to inflate their perceptions of risk and dublin and hoare (2004:274) emphasise that it is often the potential for suffering large losses, especially at harvest time, that is a major factor influencing rural communities’ attitudes, rather than actual losses. this could explain the high incidence of hwc expressed in the surveys in the present study. the close-knit community life of many rural african villages could also result in an exaggeration of hwc based on the perceptions of residents, rather than the actual number of incidences (romañach, lindsey & woodroffe 2007; woodroffe, thirgood & rabinowitz 2005). in general, respondents who said they had problems with wild animals were found to have less positive attitudes towards tourism and conservation. walpole and thouless (2005:130) highlight that tourism will only improve tolerance towards wildlife where the benefits of tourism actually reach those bearing the costs of living with wildlife and where the local communities can understand and act upon the linkages between tourism benefits and wildlife conservation. those bearing the costs are frequently not the same people receiving the benefits and this needs to be taken into consideration for long-term sustainability (walpole & thouless 2005). based on past studies (groom & harris 2008; mehta & kellert 1998), it was assumed that those with higher monthly household incomes would have more positive attitudes towards tourism and pas. this assumption is based on the fact the wealthier households have the ‘luxury’ of being able to either enjoy the tourism, pas and/or are not as greatly affected by the negative impacts of conservation, for example hwc, loss of access to natural resources, et cetera. income had mixed effects on attitudes in this study and suggests that household wealth is not always a driver of positive attitudes. of importance to policymakers is that groom and harris (2008:250) found in their kenyan study that although financial incentives from wildlife can improve community attitudes towards wildlife and conservation, the actual distribution of benefits is more important in shaping attitudes. this is supported by the results from the torra conservancy in namibia, where respondents’ attitudes were still (i.e. in 2009) influenced by a dividend payout in 2003 (see snyman 2012a). the lack of consistency of the overall importance of household income as a determinant of attitudes in the probit models suggests that income, as an incentive, is not the only incentive that matters to these communities. past studies have found that increased education impacted positively on attitudes towards tourism and pas (see chidakel 2011; larson 2010; shibia 2010; tessema et al. 2007; teye et al. 2002). teye et al. (2002), in their ghanan study, surmised that a more positive attitude towards tourism by more educated people could be explained by the fact that much has been written in english regarding the benefits of tourism in both print and electronic media, therefore educated people would be more familiar or aware of the potential benefits than those with less education. overall in the present study, formal education was found to have a statistically significant positive impact on individual’s attitudes to tourism and pas, highlighting the importance of formal education in terms of garnering support from rural communities. the benefits of formal education are numerous in terms of long-term poverty reduction, as well as biodiversity conservation (see snyman 2012c). one would surmise that if a respondent had a family member, or in the case of the community respondents, was themselves, employed in tourism and/or conservation, then they would have more positive attitudes towards tourism and pas. this is premised on the fact that they, or their family members, would be receiving direct, tangible benefits from tourism and/or pas and would therefore be more positive towards it. this premise was supported by the results in this study. in fact, having a family member employed in tourism or conservation was one of the most significant variables in a number of the analyses. this illustrates that it is not necessarily required that someone has to be receiving direct benefits themselves to be aware of the benefits that can result from tourism and pas. conclusion top ↑ it is frequently the vulnerability of poorer households and the risks they face that leads to less positive attitudes towards tourism and pas. as discussed, the costs that communities have to bear are often high. if there are no concomitant benefits associated with these costs, then it is unsurprising that households would hold negative attitudes towards pas and the associated tourism operations in the area.not only are there numerous direct factors affecting people’s attitudes to tourism and pas, they may also be influenced by costs and benefits that accrue to others, including those in different households (emerton, in sandbrook & adams 2012). this would explain why some households, who are not directly benefiting from, or negatively affected by, tourism or pas, still hold certain attitudes towards it. a factor that was shown to affect attitudes to conservation and that can be managed in order to ensure future positive attitudes was the length of operation of the ecotourism camp in the area. in general, the longer the camp had been open, the more negative respondents were about conservation; although, they interestingly tended to be more positive about tourism. this does not bode well for the long-term success of pas and needs to be addressed. the main reason for this is likely because of unmet expectations, unfulfilled promises and high levels of hwc. the management of community expectations and benefit distribution before an ecotourism camp starts operating, as well as during the operational phase, including realistic goal setting, is important to the long-term success of ecotourism in pas. it will help to reduce negative attitudes that can result from unmet expectations and the consequent dissatisfaction of communities living adjacent to pas. formal employment of any kind was shown to have a largely positive impact on attitudes as it allows a diversification of household livelihoods and reduces the risk they face. the important point to note is that there are few formal employment opportunities in these remote rural areas, other than in ecotourism. the results of this article highlight the diverse array of factors affecting people’s attitudes towards ecotourism and pas. monetary benefits from ecotourism alone will not serve to improve local people’s attitudes towards ecotourism and pas, as there are a number of factors shaping attitudes. these include receipt of tangible, as well as intangible benefits, demographic factors, local economic situation, past beliefs, cultural beliefs, land ownership systems, population density and the diversity of livelihood strategies available to households in the area. the more livelihood strategies people have available to them, the less dependent they are exclusively on natural resources for survival and livelihoods. it is premised that people will then value conservation and pas more. dependence on ecotourism as the sole livelihood can also be risky and vulnerable to external shocks. ideally, ecotourism therefore needs to be part of a diverse livelihood portfolio. conservation, for most rural africans, is an investment for present and future value, with the main goal being the maintenance or enhancement of their livelihoods (hulme & murphree 2001). ecotourism can therefore assist communities in earning much-needed income, as well as assist them in conserving their natural resources in order to maintain or enhance other livelihoods. such a scenario would however only apply in areas where communities have access to the natural resources in the area where the ecotourism camp operates (snyman 2013). communities who are excluded from pas will require further incentives to conserve natural resources, as they will not be benefitting directly from their conservation. global benefits to be derived from ecosystem services are also important and communities can play a role in this through watershed protection, preventing deforestation, et cetera. the importance of education in terms of obtaining employment, as well as more positive attitudes is important in terms of the sustainability of tourism and conservation as land uses. increasing access to formal education, improving education infrastructure, as well as implementing awareness and education campaigns in communities can serve to increase people’s knowledge and awareness of pas and tourism (snyman 2013). this, in turn, may result in a greater willingness to accept the costs of living with wildlife and more positive attitudes towards pas and tourism in the area. positive attitudes do not however necessarily suggest that behaviours will also promote conservation and tourism. poor rural households face many economic and time constraints that can prevent them from supporting conservation (snyman 2013). parry and campbell (in emptaz-collomb 2009:101) suggest that improving the living conditions and social welfare of rural people is therefore an important part of any conservation strategy. ecotourism can play an important role here, through donations towards community development projects, tourism-related infrastructure developments, directly through wages and salaries and indirectly as suppliers of goods and services. in summary, some implications for management drawn from this research include: • in areas where government owns the land (in this study, national parks) and there is no contractual community involvement (e.g. malawi, zambia and zimbabwe), there have to be benefits, both tangible and intangible, received by the community, as well as a mitigation of the negative impacts associated with conservation (i.e. hwc). outreach programmes, introduced by the private sector tourism operator, in communities abutting the park could include educational programmes as well as social welfare projects. such programmes would serve to link pas and tourism directly to benefits (snyman 2012b). • government, ngos or the private sector need to raise awareness relating to ecotourism, conservation and pas. ecotourism operators can play an important role in this through environmental talks and conservation and tourism awareness-raising days in communities, as well as offering environmental lessons and game drives to community school children, as many have never been inside the pa adjacent to their homes (snyman 2012b). • there must be a clear, structured process of setting and managing expectations prior to an ecotourism operator starting in an area, as well as through the operational phases. • overall, it is not only important to maximise benefits to communities, there needs to be a concomitant process of minimising costs, as often there are more who will bear the costs than there are those who will benefit from the conservation and ecotourism in the area (snyman 2012a). • alternative livelihoods (such as ecotourism employment) may assist in steering households away from absolute dependence on the consumptive use of natural resources for survival, which could, in turn, promote biodiversity conservation and long-term sustainable use, as well as positive attitudes (snyman 2012b). • formal education is important and has been shown to influence attitudes. improved educational infrastructure and improved access to education (e.g. scholarship programmes) should therefore be a priority in rural areas. managing community expectations, through an understanding of community members’ attitudes under varying socio-economic conditions, will lead to more efficient, equitable and sustainable community-based conservation and tourism models. also important for long-term sustainability is that, over and above this understanding, one needs to know what factors directly affect behaviours. future research should focus on monitoring behaviours related to pas, for example collection of natural resources, engagements with pa and ecotourism staff, et cetera. positive community behavioural changes towards pas and natural resources will ensure their long-term sustainability. a focus on formal education, improving social welfare and increasing the local linkages from ecotourism operations will go a long way in improving local community attitudes towards tourism and pas. acknowledgements top ↑ competing interests as stated above, the author was employed by ecotourism group, wilderness safaris, to conduct a study on the impact of the company’s activities on rural communities. however, the study itself was performed by the author as part of her phd research and in her capacity as an independent researcher looking to discover the realities of ecotourism and community development. it was in no way influenced by the company. references top ↑ akyeampong, o.a., 2011, ‘pro-poor tourism: resident’s expectations, experiences and perceptions in the kakum national park area of ghana’, journal of sustainable tourism 19(20), 197–213. http://dx.doi.org/10.1080/09669582.2010.509508alexander, s.e., 2000, ‘resident attitudes toward conservation and black howler monkeys in belize: the community baboon sanctuary’, environmental conservation 27(4), 341–350. allendorf, t.d., 2007, ‘resident’s attitudes toward three protected areas in nepal’, biodiversity conservation 16, 2087–2102. http://dx.doi.org/10.1007/s10531-006-9092-z allendorf, t.d., swe, k.k., oo, t., htut, y., aung, m., aung, m. et al., 2006, ‘community attitudes towards three protected areas in upper myanmar (burma)’, environmental conservation 33(4), 344–352. http://dx.doi.org/10.1017/s0376892906003389 andereck, k.l. & vogt, c.a., 2000, ‘the relationship between resident’s attitudes toward tourism and tourism development options’, journal of travel research 39, 27–36. http://dx.doi.org/10.1177/004728750003900104 andereck, k.l., valentine, k.m., knopf, r.c. & vogt, c.a., 2005, ‘residents’ perceptions of community tourism impacts’, annals of tourism research 32(4), 1056–1076. http://dx.doi.org/10.1016/j.annals.2005.03.001 anthony, b., 2007, ‘the dual nature of parks: attitudes of neighbouring communities to kruger national park, south africa’, environmental conservation 34(3), 236–245. http://dx.doi.org/10.1017/s0376892907004018 baral, n. & heinen, j.t., 2007, ‘resources use, conservation attitudes, management intervention and park–people relations in the western terai landscape of nepal’, environmental conservation 34, 64–72. http://dx.doi.org/10.1017/s0376892907003670 borrini-feyerabend, g., kothari, a. & oviedo, g., 2004, indigenous and local communities and protected areas: towards equity and enhanced conservation, iucn, gland, viewed 23 january 2012, from http://plateauperspectives.org/pubs/indigineous%20peoples.pdf browne-nuñez, c. & jonker, s.a., 2008, ‘attitudes toward wildlife and conservation across africa: a review of survey research’, human dimensions of wildlife 13, 47–70. http://dx.doi.org/10.1080/10871200701812936 bruyere, b.l., beh, a.w. & lelengula, g., 2009, ‘differences in perceptions of communications, tourism benefits, and management issues in a protected area of rural kenya’, environmental management 43, 49–59. http://dx.doi.org/10.1007/s00267-008-9190-7 chandralal, k.p.l., 2010, ‘impacts of tourism and community attitudes towards tourism: a case study in sri lanka’, south asian journal of tourism and heritage 3(2), 41–49. chidakel, a., 2011, conservation attitudes and community-based natural resource management in an understocked game management area of zambia, fiu electronic theses and dissertations, paper 450, viewed 10 august 2011, from http://digitalcommons.fiu.edu/etd/450 currie, j.a., 2001, ‘towards sustainable conservation: community perceptions of game reserves in the maputoland region of south africa: comparative case studies of phinda and mkhuze game reserves’, master of sustainable environmental management thesis, middlesex university. de boer, w.f. & baquete, d.s., 1998, ‘natural resource use, crop damage and attitudes of rural people in the vicinity of the maputo elephant reserve, mozambique’, environmental conservation 25(3), 208–218. http://dx.doi.org/10.1017/s0376892998000265 de witt, l., van der merwe, p. & saayman, m., 2011, ‘an ecotourism model for south african national parks’, proceedings of the international conference on tourism and management studies, algarve, portugal, 26–29 october, vol. ii, viewed 02 february 2012, from http://tmstudies.net/index.php/ectms/article/view/311 dublin, h.t. & hoare, r.e., 2004, ‘searching for solutions: the evolution of an integrated approach to understanding and mitigating human-elephant conflict in africa’, human dimensions of wildlife 9, 271–278. http://dx.doi.org/10.1080/10871200490505701 emerton, l., 1999, ‘the nature of benefits and the benefits of nature: why wildlife conservation has not economically benefitted communities in africa’, community conservation research in africa: principles and comparative practice 9, institute for development policy and management, university of manchester. emptaz-collomb, j-g.j., 2009, ‘linking tourism, human wellbeing and conservation in the caprivi strip (namibia)’, phd thesis, university of florida, viewed 15 february 2012, from http://etd.fcla.edu/uf/ufe0041031/emptazcollomb_j.pdf gadd, m.e., 2005, ‘conservation outside of parks: attitudes of local people in laikipia, kenya’, environmental conservation 32(1), 50–63. http://dx.doi.org/10.1017/s0376892905001918 gillingham, s. & lee, p.c., 1999, ‘the impact of wildlife-related benefits on the conservation attitudes of local people around the selous game reserve, tanzania’, environmental conservation 26(3), 218–228. http://dx.doi.org/10.1017/s0376892999000302 gillingham, s. & lee, p.c., 2003, ‘people and protected areas: a study of local perceptions of wildlife crop-damage conflict in an area bordering the selous game reserve, tanzania’, oryx 37(3), 316–325. http://dx.doi.org/10.1017/s0030605303000577 groom, r. & harris, s., 2008, ‘conservation on community lands: the importance of equitable revenue sharing’, environmental conservation 35(3), 242–251. http://dx.doi.org/10.1017/s037689290800489x hill, c.m., 2004, ‘farmers’ perspectives of conflict at the wildlife–agriculture boundary: some lessons learned from african subsistence farmers’, human dimensions of wildlife 9(4), 279–286. http://dx.doi.org/10.1080/10871200490505710 hoon, p.n., 2004, ‘impersonal markets and personal communities? wildlife, conservation and development in botswana’, journal of international wildlife law & policy 7(3), 143–160. http://dx.doi.org/10.1080/13880290490883223 hulme, d. & murphree, m. (eds.), 2001, african wildlife and livelihoods: the promise and performance of community conservation, pp. 106–130, james currey ltd, woodbridge. kideghesho, j.r., røskaft, e. & kaltenborn, b.p., 2007, ‘factors influencing conservation attitudes of local people in western serengeti, tanzania’, biodiversity conservation 16, 2213–2230. http://dx.doi.org/10.1007/s10531-006-9132-8 larson, k., 2010, ‘attitudes concerning conservation in two different wildlife areas in kenya’, swedish university of agricultural services, department of animal and environment and health ethology and animal welfare programme, student report 320, viewed 03 may 2013, from http://stud.epsilon.slu.se/1584/ lepp, a., 2007, ‘resident’s attitudes toward tourism in bigodi village, uganda’, tourism management 28, 876–885. http://dx.doi.org/10.1016/j.tourman.2006.03.004 lepp, a. & holland, s., 2006, ‘a comparison of attitudes toward state-led conservation and community-based conservation in bigodi village, uganda’, society and natural resources 19, 609–623. http://dx.doi.org/10.1080/08941920600742377 makindi, s.m., 2010, ‘communities’ perceptions and assessment of biodiversity conservation strategies: the case of protected areas in kenya’, phd thesis, school of environmental sciences, university of kwazulu-natal. mbaiwa, j.e. & stronza, a.l., 2011, ‘changes in resident attitudes towards tourism development and conservation in the okavango delta, botswana’, journal of environmental management 92(8), 1950–1959. http://dx.doi.org/10.1016/j.jenvman.2011.03.009, pmid:21482448 mcgehee, n.g. & andereck, k.l., 2004, ‘factors predicting rural residents’ support of tourism’, journal of travel research 43, 131–140. http://dx.doi.org/10.1177/0047287504268234 mehta, j.n. & heinen, j.t., 2001, ‘does community-based conservation shape favourable attitudes among locals? an empirical study from nepal’, environmental management 28(2), 165–177. http://dx.doi.org/10.1007/s002670010215 mehta, j.n. & kellert, s.r., 1998, ‘local attitudes toward community-based conservation policy and programmes in nepal: a case study in the makalu-barun conservation area’, environmental conservation 25(4), 320–333. http://dx.doi.org/10.1017/s037689299800040x musumali, m.m., larsen, t.s. & kaltenborn, b.p., 2007, ‘an impasse in community based natural resource management implementation: the case of zambia and botswana’, oryx 41(3), 306–313. http://dx.doi.org/10.1017/s0030605307000518 nepal, s.k., 2002, ‘linking parks and people: nepal’s experience in resolving conflicts in parks and protected areas’, international journal of sustainable development and world ecology 9(1), 75–90. http://dx.doi.org/10.1080/13504500209470104 newmark, w.d., manyanza, d.n., gamassa, d-g.m. & sariko, h.i., 1994, ‘the conflict between wildlife and local people living adjacent to the protected areas in tanzania: human density as a predictor’, conservation biology 8(1), 249–255. http://dx.doi.org/10.1046/j.1523-1739.1994.08010249.x nunkoo, r. & ramkissoon, h., 2011, ‘developing a community support model for tourism’, annals of tourism research 38(3), 964–988. http://dx.doi.org/10.1016/j.annals.2011.01.017 rodriguez, s.l., 2008, ‘perceptions and attitudes of a maasai community regarding wildlife-damage compensation, conservation and the predators that prey on their livestock’, human dimensions of wildlife 13(3), 205–206. http://dx.doi.org/10.1080/10871200801886137 romañach, s.s., lindsey, p. & woodroffe, r., 2007, ‘determinants of attitudes towards predators in central kenya and suggestions for increasing tolerance in livestock dominated landscapes’, oryx 41(2), 185–195. http://dx.doi.org/10.1017/s0030605307001779 sandbrook, c. & adams, w.m., 2012, ‘accessing the impenetrable: the nature and distribution of tourism benefits at a ugandan national park’, society and natural resources 25(9), 915–932. http://dx.doi.org/10.1080/08941920.2011.644394 sarker, a.h.m.r. & røskaft, e., 2010, ‘human attitudes towards conservation of asian elephants (elephas maximus) in bangladesh’, international journal of biodiversity and conservation 2(10), 316–327, viewed 02 april 2013, from http://www.academicjournals.org/article/article1380538795_sarker%20and%20røskaft.pdf scanlon, l.j. & kull, c.a., 2009, ‘untangling the links between wildlife benefits and community-based conservation at torra conservancy, namibia’, development southern africa 26(1), 75–93. http://dx.doi.org/10.1080/03768350802640107 sekhar, n.u., 2003, ‘local people’s attitudes towards conservation and wildlife tourism around sariska tiger reserve, india’, journal of environmental management 69, 339–347. http://dx.doi.org/10.1016/j.jenvman.2003.09.002, pmid:14680895 shibia, m.g., 2010, ‘determinants of attitudes and perceptions on resource use and management of marsabit national reserve, kenya’, journal of human ecology 30(1), 55–62. sifuna, n., 2010, ‘wildlife damage and its impact on public attitudes toward conservation: a comparative study of kenya and botswana, with particular reference to kenya’s laikipia region and botswana’s okavango delta region’, journal of asian and african studies 45, 274–296. http://dx.doi.org/10.1177/0021909610364776 simelane, t.s., kerley, g.i.h. & knight, m.h., 2006, ‘reflections on the relationships between communities and conservation areas of south africa: the case of five south african national parks’, koedoe 49(2), 85–102. http://dx.doi.org/10.4102/koedoe.v49i2.121 snyman, s., 2012a, ‘ecotourism joint ventures between the private sector and communities: an updated analysis of the torra conservancy and damaraland camp, namibia’, tourism management perspectives 4, 127–135 http://dx.doi.org/10.1016/j.tmp.2012.07.004 snyman, s., 2012b, ‘the impact of land management systems on community attitudes towards tourism and conservation in six southern african countries’, parks 18(2), 20–31. snyman, s., 2012c, ‘the role of ecotourism employment in poverty reduction and community perceptions of conservation and tourism in southern africa’, journal of sustainable tourism 20(3), 395–416. http://dx.doi.org/10.1080/09669582.2012.657202 snyman, s., 2013, ‘high-end ecotourism and rural communities in southern africa: a socio-economic analysis’, phd dissertation, school of economics, university of cape town. spss version 12, 2004, computer software, ibm corportation, new york. stata version 10.2, 2008, computer software, statacorp, college station. stem, c.j., lassorie, j.p., lee, d.r., deshler, d.d. & schelhas, j.w., 2003, ‘community participation in ecotourism benefits: the link to conservation practices and perspectives’, society and natural resources 16, 387–413. http://dx.doi.org/10.1080/08941920309177 stronza, a. & gordillo, j., 2008, ‘community views of ecotourism’, annals of tourism research 35(2), 448–468. http://dx.doi.org/10.1016/j.annals.2008.01.002 stronza, a. & pêgas, f., 2008, ‘ecotourism and conservation: two cases from brazil and peru’, human dimensions of wildlife 13, 263–279. http://dx.doi.org/10.1080/10871200802187097 tessema, m.e., ashenafi, z.t., lilieholm, r.j. & leader-williams, n., 2007, ‘community attitudes toward wildlife conservation in ethiopia’, proceedings of the 2007 george wright society conference, assessing public attitudes and experiences, st paul, minnesota, 16–20 april, viewed 03 may 2013, from http://www.georgewright.org/0755tessema.pdf teye, v., sönmez, s.f. & sirakaya, e., 2002, ‘resident attitudes toward tourism development’, annals of tourism research 29(3), 668–688. http://dx.doi.org/10.1016/s0160-7383(01)00074-3 walpole, m.j. & goodwin, h.j., 2001, ‘local attitudes towards conservation and tourism around komodo national park, indonesia’, environmental conservation 28(2), 160–166. http://dx.doi.org/10.1017/s0376892901000169 walpole, m.j. & thouless, c.r., 2005, ‘increasing the value of wildlife through non-consumptive use? deconstructing the myths of ecotourism and community-based tourism in the tropics’, in r. woodroffe, s. thirgood & a. rabinowitz (eds.), people and wildlife: conflict or coexistence?, pp. 122–139, the zoological society of london and cambridge university press, cambridge. wang, y. & pfister, r.e., 2008, ‘resident’s attitudes toward tourism and perceived personal benefits in a rural community’, journal of travel research 47(1), 84–93. http://dx.doi.org/10.1177/0047287507312402 waylen, k.a., mcgowan, p.j.k., pawi study group & milner-gulland, e.j., 2009, ‘ecotourism positively affects awareness and attitudes but not conservation behaviours: a case study at grand riviere, trinidad’, oryx 43(3), 343–351. http://dx.doi.org/10.1017/s0030605309000064 weladji, r.b., moe, s.r. & vedeld, p., 2003, ‘stakeholder attitudes toward wildlife policy and the bénoué wildlife conservation area, north cameroon’, environmental conservation 30(4), 334–343. http://dx.doi.org/10.1017/s0376892903000353 woodroffe, r., thirgood, s. & rabinowitz, a., 2005, ‘the future of co-existence: resolving human–wildlife conflicts in a changing world’, in r. woodroffe, s. thirgood & a. rabinowitz (eds.), people and wildlife: conflict or coexistence?, pp. 388–405, the zoological society of london and cambridge university press, cambridge. article information authors: graham pasternak1,2 leslie r. brown2 stefan kienzle2,3 andrea fuller4 louise barrett1,2 s. peter henzi1,2 affiliations: 1department of psychology, university of lethbridge, canada2applied behavioural ecology and ecosystems research unit, university of south africa, south africa 3department of geography, university of lethbridge, canada 4school of physiology, university of the witwatersrand, south africa correspondence to: peter henzi postal address: department of psychology, university of lethbridge, lethbridge, alberta t1k 3m4, canada dates: received: 25 feb. 2012 accepted: 20 aug. 2012 published: 20 feb. 2013 how to cite this article: pasternak, g., brown, l.r., kienzle, s., fuller, a., barrett, l. & henzi, s.p., 2013, ‘population ecology of vervet monkeys in a high latitude, semi-arid riparian woodland’, koedoe 55(1), art. #1078, 9 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1078 copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. population ecology of vervet monkeys in a high latitude, semi-arid riparian woodland in this original research... open access • abstract • introduction • research method and design    • study area    • vegetation surveys    • group counts    • activity, diet and ranging    • body mass    • statistical analysis • results    • vegetation structure    • population structure    • inter-birth intervals    • day journey length    • home range size and population density    • activity    • diet    • body mass • ethical considerations • discussion • conclusion • acknowledgements    • competing interests    • authors' contributions • references • appendix abstract top ↑ narrow riparian woodlands along non-perennial streams have made it possible for vervet monkeys to penetrate the semi-arid karoo ecosystem of south africa, whilst artificial water points have more recently allowed these populations to colonize much more marginal habitat away from natural water sources. in order to better understand the sequelae of life in these narrow, linear woodlands for historically ‘natural’ populations and to test the prediction that they are ecologically stressed, we determined the size of troops in relation to their reliance on natural and artificial water sources and collected detailed data from two river-centred troops on activity, diet and ranging behaviour over an annual cycle. in comparison to other populations, our data indicate that river-centred troops in the karoo were distinctive primarily both for their large group sizes and, consequently, their large adult cohorts, and in the extent of home range overlap in what is regarded as a territorial species. whilst large group size carried the corollary of increased day journey length and longer estimated interbirth intervals, there was little other indication of the effects of ecological stress on factors such as body weight and foraging effort. we argue that this was a consequence of the high density of acacia karroo, which accounted for a third of annual foraging effort in what was a relatively depauperate floristic habitat. we ascribed the large group size and home range overlap to constraints on group fission.conservation implications: the distribution of group sizes, sampled appropriately across habitats within a conservation area, will be of more relevance to management than average values, which may be nothing more than a statistical artefact, especially when troop sizes are bimodally distributed. introduction top ↑ models that derive the limits of group size from the constraining effects of environmental variables on activity schedules have been very successful in predicting the distribution of social primates (dunbar 1996, korstjens, verhoeckx & dunbar 2006). such modelling has value, not only for recreating a species’ evolutionary biogeography, but also for simulating the future consequences of climate change. nevertheless, by necessity, these models operate at a relatively coarse spatial scale and are not therefore able to discern the local presence or absence of species. a very good example of this is the inability of a recent activity budget model to detect the presence of vervet monkeys (cercopithecus aethiops) in the semi-arid karoo biome of south africa (figure 5 in willems & hill 2009), whereas they have had a historically documented presence since at least the 18th century (skead 1987) and are, in fact, now regionally widespread.although the provision of artificial point water sources for livestock has increased their distribution, local vervet monkey populations in the karoo have been confined historically to narrow strips of acacia karroo woodland along non-perennial rivers and streams in otherwise inhospitable open country. in this regard, then, the failure of the activity budget model to predict their presence is a consequence only of the fact that these riparian woodlands are, in effect, anomalous features in the larger landscape. nevertheless, they are very important components of the local ecosystem, both as refugia and corridors (puth & wilson 2001), especially in the context of increasing temperatures and decreasing rainfall in this part of south africa (hoffman et al. 2009). consequently, the general research question we address here is directed at the local viability of these vervet populations and the consequences of life for an obligate social mammal restricted to these narrow riparian strips, where migration pathways are severely constrained (see also isbell, cheney & seyfarth 2002) and the effects of drought are exacerbated by both high summer temperatures and very low winter ones. taken in conjunction with the modelled expectation that vervet monkeys should not be found in the karoo, we test the obvious prediction that existing natural populations should exhibit clear signs of ecological stress and be susceptible to extirpation. we present data on the demographic and ecological characteristics of one such population, not only to describe its general characteristics, but also to provide a qualitative test of the prediction by comparison with available data from vervet populations elsewhere. research method and design top ↑ study area the data come from an ongoing study of a vervet monkey population in the samara private game reserve, eastern cape province, south africa (32°22’s, 24°52’e, figure 1). the reserve comprises 27 000 ha of mountains and nama-karoo grassland transected by the milk river and its tributaries. our study site is located in the north of the reserve where the river, which flows only intermittently, has not been dammed and where the monkeys have no access to artificial water sources. such point sources of water have been established away from the river and generally have vervet groups associated with them. the area receives a declining average of 330 mm rain per annum (wet season, october–march; dry season, april–september) and experiences a mean maximum temperature of 27 °c and a mean minimum temperature of 10 °c. the coldest month is july when snow falls on the surrounding mountains (mean minimum, 4 °c), whilst december and january are the hottest (mean maximum, 34 °c). in addition to a variety of ungulates, the reserve has an established predator guild. animals either observed to prey on the study population, or to elicit alarm calling, include cheetah (acinonyx jubatus), caracal (caracal caracal), black-backed jackal (canis mesomelas), martial eagle (polemaetus bellicosus), verreaux’s eagle (aquila verreauxii) and cape eagle owl (bubo capensis). whilst there are no large constrictors in the region, venomous snakes were also a confirmed source of mortality. figure 1: the location of the general study area within south africa and in relation to the town of graaff-reinet, where the rectangle indicates the study site enlarged in the inset. vegetation surveys we stratified the study site into physiognomic–physiographic units using a 1:12 000 aerial photograph and located 35 sample plots of 400 m2 on a randomly stratified basis within the various identified units, where the number of plots was determined by each unit’s area. we conducted braun-blanquet vegetation and habitat surveys in each of the sample plots during march 2010 to define plant communities by recording all plant species in each of the plots and estimating the percentage cover for the tree, shrub, and herbaceous layers using the modified 9-degree braun-blanquet sampling scale (kent & coker 2008). this method is used to define plant communities by grouping sets of vegetation samples on the basis of their floristic attributes (barbour, burk & pitts 1987). we recorded all plant species present in each of the plots and estimated the percentage cover for the tree, shrub, and herbaceous layers using the braun-blanquet cover abundance scale. we first analysed the floristic data using the multivariate classification programme, two-way indicator species analysis (twinspan) (tichý et al. 2007) to obtain a statistically derived approximation of the main plant communities. we then refined this classification by applying braun-blanquet procedures (brown & bredenkamp 1994). group counts two or more observers simultaneously counted vervet monkey groups during 2009 and 2010, either from vehicles or on foot at distances of 10 m – 100 m. all groups were counted at least twice and their locations specified using a global positioning system (gps) recorder. most groups were located early in the mornings at their sleeping sites and counted as they departed. this allowed us to obtain repeated, reliable counts of smaller troops in open country away from the river. counts of groups with home ranges centred on the river were made as they crossed roads or dry river-beds. apart from the two study troops, for which exact troop size records have been kept, counts of such river-centred groups are best regarded as conservative estimates of troop size. activity, diet and ranging data on the diurnal activity schedules and diet of adult female monkeys were obtained over ten months, from february to november, during 2010. they came from two habituated groups (rbm, rst) using scan samples (altmann 1974) taken over a 10 min period every 30 min during all-day follows (n2010 = 36 898 records). our data indicate no systematic bias in the number of records across the day and we derive percentages from overall frequencies. all animals could be followed at distances of one to five metres. during each scan period, the activity of all visible animals was assigned to one of four mutually exclusive states – foraging, moving, socialising or resting – and recorded using an electronic data logger. when animals were foraging, we also recorded both the food species and the part eaten. at the end of each scan period, we took a gps reading from the troop’s estimated centre. these readings, together with additional data from january and december 2010, were used to derive both day journey lengths and, subsequently, home range areas for each of the two troops. after importing all relevant gps data, we derived day journey length and travel velocities using both the et geowizards tool (tchoukanski 2010) and arcgis. we applied the home range tools (hrt) for arcgis (rodgers et al. 2007) to determine the minimum convex polygon (mcp) estimates of both the home range size (99% mcp) and each troop’s core area (50% mcp), using an adaptive kernel with a bandwidth of 25 m. home range overlaps were calculated from gps readings taken of the location of all other troops when these were observed in areas used by the two study troops, where accuracy was ensured by taking the readings from the observed locations once the other troops had moved away.in addition to these behavioural data, records have been kept of all births, deaths, immigration and emigration since november 2008. body mass we obtained body mass data from adults of both sexes that were anaesthetised for the surgical implantation of temperature loggers (university of the witwatersrand animal ethics screening committee clearance number 2010/41/04). statistical analysis all tests were conducted with the jmp 9 statistical package (sas institute 2007), with alpha set at 0.05. results top ↑ vegetation structure the area has relatively low species richness, with 124 different plant species identified within the home ranges of the two study vervet troops. we identified three plant communities that could be grouped into two major structural units: (1) open dwarf shrubland on the slightly higher lying and drier areas, which continues into (2) dense a. karroo woodland along the seasonal river (figure 2). figure 2: cross-sectional vegetational profile of the study area, indicating the three main communities. the largest plant community – lycium oxycarpum–a. karroo woodland – occurs along the river and floodplain and is dominated by the trees, a. karroo and l. oxycarpum, with the tree rhus lancea and the tall shrub rhus longispina also prominent. large single individuals (> 6 m) of the pepper bark tree (schinus molle) are scattered along the river bank. the woody species range in height from 1 m to 5 m and form dense canopies that cover up to 80% of the area. the herbaceous layer is dominated by the palatable grass, panicum maximum, and the pioneer grass, cynodon incompletus. the two communities that comprise dwarf shrubland – pentzia globosa–grewia robusta shrubland to the west and p. globosa–c. incompletus shrubland to the east – extend on the higher-lying areas adjacent to the acacia woodland and are both relatively open areas dominated by the dwarf shrub, p. globosa, and the tall shrub, r. longispina. whereas the woodland is still in very good condition, both shrubland communities are degraded – a consequence of grazing in the p. globosa–g. robusta shrubland and cultivation in the p. globosa–c. incompletus shrubland prior to 1998. population structure we obtained repeated, reliable counts from 29 troops. this gave us a mean troop size of 26.62 (± 18.11 s.d.) for the population. troops were distinguished, however, by whether their ranges were centred on the river or whether their source of water was artificial and maintained by humans. river troops (n = 15) had a significantly larger mean size (40.13 ± 15.53 s.d.) than troops relying on human-provided water (n = 14, mean = 12.14 ± 3.23 s.d., analysis of variance [anova]: f1.27 = 43.57, p < 0.0001, figure 3). the sizes of the two study troops, occupying adjacent home ranges, were nrbm ≈ 48 and nrst ≈ 72 respectively. the modal adult sex ratio (male [m] or female [f]) was 0.67 for rbm (nmales = 10, nfemales = 15) and 0.43 for rst (nmales = 10, nfemales = 23). figure 3: mean troop size +/1 s.d. and distribution for non-river and rivercentred troops. inter-birth intervals we recorded 65 births over 36 months (nrbm = 30; nrst = 35), with an overall birth rate of 0.57 infants/female/year (birth raterbm = 0.67; birth raterst = 0.5), allowing us to estimate an inter-birth interval of 21 months (rbm = 17.9 months; rst = 24 months). day journey length we analysed 295 entire day journeys (nrbm = 138, nrst = 157). the mean distance travelled by rbm was 2806.3 m (range: 1013 m – 5229 m) whilst that for rst was 2353.6 m (range: 912 m – 4320 m). the data come from a year of low rainfall when both study troops frequently visited a distant water hole (n = 100) during periods when there was no water available in their core areas (mcdougall et al. 2010). as this increased the mean day journey by more than 500 m, we therefore ran a full-factorial anova with troop identity, season wet or dry and water hole visits (yes or no) as factors and day journey length as the dependent variable. the whole model was significant (f7.287 = 11.22, adj. r2 = 0.195, p < 0.001) and there were significant main effects for troop identity (f1.1 = 15.1, p < 0.001), season (f1.1 = 4.5, p < 0.05, mean distancedry = 2760.3 m, mean distancewet = 2387.8 m) and for water hole visits (f1.1 = 32.75, p < 0.0001, mean distanceyes = 2972.3 m, mean distanceno = 2431.97). the only significant interaction was for season water hole visit (f1.1 = 4.4, p < 0.05), with visits to the water hole increasing day journey length in the wet season. home range size and population density we used the day journeys to estimate the annual home range sizes for the two study troops (figure 4). in the habituated group rbm’s 99% mcp covered an area of 176.1 ha of which the four discrete core areas constituted 10.45 ha. they shared 23% of their home range with five other troops. the habituated group rst used 63.7 ha over the same period, with a single core area of 7.42 ha. they shared 86% of their home range with 4 other troops. the larger home range of the smaller group is likely to be due to their having no other troops on their western boundary. on the assumption that overlap areas were shared equally by the study troops and their neighbours (cf. whiten, byrne & henzi 1987), the data from rbm generate a population density of 30.79 animals/km2, whilst the value for rst is 202.24 animals/km2. combining the data for rbm and rst and taking account of the 40.7 ha overlap between their two home ranges, gave a composite population density estimate of 54.68 animals/km2. we estimated the defendability of these home ranges (a) in relation to the average day journey length (d), using mitani and rodman’s (1979) defendability index (d), where d = d/d’ and d’ = 4a/∏0.5, where d ≥ 1, home ranges are assumed to be defendable. the values for samara were computed as drbm = 1.05; drst = 1.44. figure 4: home ranges of the study troops, indicating core areas and overlap. activity the relative allocation of time by the females of the two troops to the four activity categories is provided in table 1. table 1: the annual percentage of adult scan samples allocated to each of four activities. diet the animals foraged on 26 different plant species as well as fungi and a number of different insects, of which grasshoppers and termites were eaten most frequently (appendix 1). the five most frequent items in the diet accounted for 68.7% of annual foraging effort. acacia karroo products alone accounted for 33.8% of all foraging records and were used consistently across the 10 months (figure 5). figure 5: contributions of the five most commonly used food sources to monthly foraging effort. body mass adult males weighed 5.93 kg +/0.42 s.d. (n = 9) and females weighed 3.3 kg. +/0.3 s.d. (n = 13). these are compared to values from other populations in figure 6. figure 6: comparison of body mass across different vervet monkey populations. where available, standard deviations are indicated. ethical considerations top ↑ the behavioural data collection was conducted under the terms of reference of animal welfare protocol 0702 (university of lethbridge) and clearance for the body mass measurements was obtained from by the university of the witwatersrand animal ethics screening committee (clearance number 2010/41/04). discussion top ↑ karoo vervet troops whose ranges centre on local rivers are significantly larger than those found away from rivers. colonisation of areas away from these rivers is made possible by the presence of artificial water points that therefore provide a population sink for large river troops, but carries the corollary that the animals are confronted by habitat that is both naturally less productive and currently degraded. this would explain their small troop sizes and suggests a vulnerability to downturns in local environmental conditions. indeed, at least three non-river troops that were monitored in 2009 as part of the census were no longer locatable by the end of 2010, over which period the local drought had worsened. whilst the presence of water means that their disappearance was likely to have been associated with a decline in food availability, it is not known whether they suffered extirpation or had returned to the river and fused with troops there, as observed by isbell, cheney and seyfarth (1991). the absence of artificial water points along the milk river and the relative lack of degradation of the riparian habitat itself suggests that river-centred troops constitute the ‘natural’ regional population and can therefore be used to characterise the response of the species to these narrow habitable corridors that run through an otherwise inhospitable landscape. perhaps the most surprising finding in this regard, given the inability of the activity budget models to predict their presence in the region, is that they are not a marginal population but an apparently flourishing one, with most parameter values used to determine population viability well within the range of those reported from other study sites (table 2; figure 6). there are two values, however, that might, at first glance, be assumed to indicate ecological stress. the first is the longer interbirth interval in comparison to those reported from the long-term study at amboseli and the second is the longer day journeys undertaken by our two study groups. although data from more years will be valuable, both may reasonably be explained in terms of increased group size: the associated intragroup competition leads to lower reproductive rates, whilst larger groups need to cover more ground in order to forage (borries et al. 2008; clutton-brock & harvey 1977). the latter is corroborated by data from 12 troops (barrett 2009; willems 2009, this study) that indicate a positive correlation between troop size and day journey length (rs = 0.78, n = 12, p < 0.001). table 2: comparative data for different vervet monkey populations arranged by latitude. what therefore needs to be explained are the distinctively large size of river-centred groups and – given that vervet monkeys are territorial (struhsaker 1967) – the marked extent of home range overlap. in the absence of indications of gross ecological stress, the large troops living at high densities are clearly being sustained by the consistent availability of a. karroo products, as is true, not only for the historically high density population of vervets at amboseli in kenya, where acacia xanthophloea and acacia tortilis played much the same role (struhsaker 1967, lee & hauser 1998), but also for vervets living at lower density in the northern south africa (barrett 2004, 2009). at the same time, the inclusion of both insects and succulents in the diet (pasternak 2011) buffered the animals during those periods when free-standing water was absent (mcdougall et al. 2010). whilst one might expect some broad positive relationship between group size and population density (van schaik 1983), three things suggest that this alone will not provide an adequate explanation for population structure at samara. the first is that there is no indication of a correlation between density and troop size for the vervet populations in table 2 (rs = -0.09, n = 11, p = 0.77). the second is that mean troop size was very much smaller at a number of other sites where density was comparably high (table 2). the third is that the time budget model for vervet troop size derived by willems and hill (2009) predicts a maximum ecologically tolerable troop size of 46 animals at samara (henzi et al. in prep.). whereas the maximum troop size exceeded observed sizes for almost all their populations (figure 6 in willem & hill 2009), both our study troops exceeded model predictions. one possibility, of course, is that predation risk is responsible (van schaik 1983) but, again, for example, the amboseli population had troops of significantly smaller size whilst being as much, if not more, at risk as the samara river-centred troops (pasternak 2011, struhsaker 1967).to our minds, the most viable explanation is that successful troop fission, which would offer the benefits of shorter day ranges and inter-birth intervals, requires the maintenance of access to the riparian woodland and, given that this habitat is probably saturated, this is likely to managed only rarely, fission away from the river is likely only to occur when conditions are good and when, historically, temporary water sources are available for colonisation. interestingly, in this regard, following unusually high rainfall and improvements in resource availability during 2011, our two study troops have shown increased dispersion, with small subgroups foraging and, occasionally, sleeping independently within the home range (unpublished data). whilst it might be argued that subgroups of this kind could simply carve out territories within the larger home range and establish themselves in this way at any time, the high d values mean that any putative subgroup will intersect with other troop members frequently, making it difficult to sustain coherence in membership, especially since social coherence is reduced in these large groups in any case (henzi et al. in prep.). given this, the unusually high levels of home range overlap appear to be a natural corollary of what is, in effect, a super-saturated riparian population, sustained by high yields from a few food sources. under such conditions, we have found that troops do not invariably fight when they meet. if we are correct, we should record fission from these troops in phases of improved ecological conditions when the areas around artificial water points can support small groups, at least temporarily. in summary, as at amboseli prior to the population crash (isbell et al. 1991), which in many regards resembles samara most closely, the fact that acacia provides a consistent source of food products and is abundant, allows the vervet population to reach high densities in an otherwise inhospitable environment. unlike amboseli and other localities, the narrow riparian distribution of acacia and the sharp transition to marginal habitat, makes it very difficult for troops to divide into smaller units. these large river-centred troops, in addition to any implications they carry for conservation in the region, offer an excellent opportunity to study the social dynamics of larger male and female cohorts under natural conditions and in the absence of any obvious ecological stressors. conclusion top ↑ the vervet population in this region of the semi-arid karoo experiences little sign of ecological stress, and our estimates of population viability are in line with those from areas comprising more suitable vervet habitat (in terms of resource and water availability). troops living in the riparian zone were significantly larger than those living away from the river, and were sustained by the constant availability of a. karroo. we suggest that these larger group sizes occur because group fission is inhibited by the poor quality of habitat away from the river. this gives rise to a super-saturated riparian population, in which territorial behaviour is reduced. such groups offer an excellent opportunity to test hypotheses regarding the social dynamics and life history of vervet monkeys. our results also emphasise that effective conservation and management practices in the karoo should be predicated on an assessment of the distribution of group sizes, and not simply the population mean, as the bimodal distribution of large riparian groups and smaller non-riparian groups renders the latter a statistical artefact, rather than a measure that possesses biological relevance. acknowledgements top ↑ this project has been funded by grants from nserc canada, nrf south africa and unisa. we are most grateful to mark and sarah tompkins for logistical assistance and permission to work at samara. we thank ria boner, natalie freeman, and tricia rubi for help with data collection and richard and kitty viljoen for many acts of kindness. two anonymous referees provided advice that improved the manuscript substantially. competing interests the authors declare that they have no financial or personal relationship(s) which may have inappropriately influenced them in writing this article. authors’ contributions g.p. (university of lethbridge) collected and, with s.p.h. (university of lethbridge) and l.b. (university of lethbridge), analysed the behavioural data. l.r.b. (university of south africa) undertook the vegetation survey, a.f. (university of the witwatersrand) collected body mass data, s.p.h., g.p. and l.b. did the population survey. s.k. (university of south africa) and g.p. undertook the spatial analyses. all authors participated in the writing of the manuscript. references top ↑ altmann, j., 1974, ‘observational study of behaviour: sampling methods’, behaviour 49, 227–267.baldellou, m., 1992, ‘implications of the multimale troop structure in vervet monkeys (cercopithecus aethiops pygerythrus)’, phd thesis, department of psychology, university of natal. barbour, m.g., burk, j.h. & pitts, w.d., 1987, terrestrial plant ecology, benjamin cummings, menlo park. barrett, a.s., 2004, ‘foraging ecology of the vervet monkey (chlorocebus aethiops) in mixed lowveld bushveld and sour lowveld bushveld of the blydeberg conservancy, northern province, south africa’, mtech dissertation, department of environmental sciences, university of south africa. barrett, a.s., 2009, ‘spatial and temporal patterns in resource dispersion and the structure of range use and coexistence in a social omnivore (chlorocebus aethiops)’, phd thesis, department of environmental sciences, university of south africa. bolter, d.r. & zihlman, a.l., 2003, ‘morphometric analysis of growth and development in wild-collected vervet monkeys (cercopithecus aethiops), with implications for growth patterns in old world monkeys, apes and humans’, journal of zoology 260, 99–110. http://dx.doi.org/10.1017/s0952836903003522 borries, c., larney, c., lub, a., ossib, k. & koenig, a., 2008, ‘costs of group size, lower developmental and reproductive rates in larger groups of leaf monkeys’, behavioral ecology 19, 1186–1191. http://dx.doi.org/10.1093/beheco/arn088 brown, l.r. & bredenkamp, g.j., 1994, ‘the phytosociology of the southern section of the borakalalo nature reserve, south africa’, koedoe 37, 59–72. http://dx.doi.org/10.4102/koedoe.v37i2.337 clutton-brock, t.h. & harvey, p.h., 1977, ‘primate ecology and social organization’, journal of zoology 183, 1–39. http://dx.doi.org/10.1111/j.1469-7998.1977.tb04171.x dunbar, r.i.m., 1996, ‘determinants of group size in primates: a general model’, proceedings of the british academy 88, 33–57. enstam, k.l. & isbell, l., 2007, ‘the guenons (genus cercopithecus) and their allies: behavioral ecology of polyspecific associations’, in c.j. campbell, a. fuentes, k.c. mckinnon, m. panger & s.k. bearder (eds.), primates in perspective, pp. 252–274, oxford university press, new york. hall, k.r.l. & gartlan, j.s., 1965, ‘ecology and behaviour of the vervet monkey, cercopithecus aethiops, lolui island, lake victoria’, proceedings of the zoological society london 145, 37–56. http://dx.doi.org/10.1111/j.1469-7998.1965.tb01999.x harrison, m.j.s., 1983, ‘territorial behaviour in the green monkey, cercopithecus sabaeus: seasonal defense of local food supplies’, behavioral ecology and sociobiology 12, 85–94. http://dx.doi.org/10.1007/bf00296937 harrison, m.j.s., 1985, ‘time budget of the green monkey, cercopithecus sabaeus: some optimal strategies’, international journal of primatology 6, 351–376. http://dx.doi.org/10.1007/bf02736383 henzi, s.p., barrett, l., forshaw, n., boner, r. & lusseau, d., in prep., ‘scalar social dynamics in female vervet monkey cohorts’. hoffman, m.t., carrick, p.j., gillson, l. & west, a.g., 2009, ‘drought, climate change and vegetation response in the succulent karoo, south africa’, south african journal of science 105, 54–60. http://dx.doi.org/10.1590/s0038-23532009000100021 isbell, l.a., cheney, d.l. & seyfarth, r.m., 1991, ‘group fusions and minimum group sizes in vervet monkeys’, american journal of primatology 25, 57–65. http://dx.doi.org/10.1002/ajp.1350250106 isbell, l.a., cheney, d.l. & seyfarth, r.m., 2002, ‘why vervet monkeys (cercopithecus aethiops) live in multimale groups’, in m.e. glenn & m. cords (eds.), the guenons: diversity and adaptation in african monkeys, pp. 173–187, kluwer academic/plenum publishers, new york. kaghira, j.m., ngotho, m., thuita, j.k., maina, n.w. & hau, j., 2007, ‘hematological changes in vervet monkeys (chlorocebus aethiops) during eight months’ adaptation to captivity’, american journal of primatology 69, 1053–1063. pmid:17294427 kent, m. & coker, p., 2008, vegetation description and analysis: a practical approach, belhaven press, london. korstjens, a.h., verhoeckx, i.l. & dunbar, r.i.m., 2006, ‘time as a constraint on group size in spider monkeys’, behavioral ecology and sociobiology 60, 683–694. http://dx.doi.org/10.1007/s00265-006-0212-2 lee, p.c. & hauser, m.d., 1998, ‘long-term consequences of changes in territory quality on feeding and reproductive strategies of vervet monkeys’, journal of animal ecology 67, 347–358. http://dx.doi.org/10.1046/j.1365-2656.1998.00200.x mcdougall, p., forshaw, n., barrett, l. & henzi, s.p., 2010, ‘leaving home: responses to water depletion by vervet monkeys’, journal of arid environments 74, 924–927. http://dx.doi.org/10.1016/j.jaridenv.2010.04.003 mitani, j.c. & rodman, p.s., 1979, ‘territoriality: the relation of ranging pattern and home range size to defendability, with an analysis of territoriality among primate species’, behavioral ecology and sociobiology 5, 241–251. http://dx.doi.org/10.1007/bf00293673 nakagawa, n., 1999, ‘differential habitat utilization by patas monkeys (erythrocebus patas) and tantalus monkeys (cercopithecus aethiops tantalus) living sympatrically in northern cameroon’, american journal of primatology 49, 243–264. http://dx.doi.org/10.1002/(sici)1098-2345(199911)49:3<243::aid-ajp3>3.0.co;2-4 pasternak, g., 2011, ‘environmental effects on group structure and vigilance in vervet monkeys’, msc thesis, dept. of psychology, university of lethbridge. pruetz, j.d.e., 2009, the socioecology of adult female patas monkeys and vervets in kenya, pearson prentice hall, upper saddle river, nj. puth, l.m. & wilson, k.a., 2001, ‘boundaries and corridors as a continuum of ecological flow control: lessons from rivers and streams’, conservation biology 15, 21–30. rodgers, a.r., carr, a.p., beyer, h.l., smith, l. & kie, j.g., 2007, home range tools for arcgis, viewed on 13 june 2011, from http://blue.lakeheadu.ca/hre/ van schaik, c.p., 1983, ‘why are diurnal primates living in groups?’, behaviour 87, 120–144. http://dx.doi.org/10.1163/156853983x00147 skead, c.j., 1987, historical mammal incidence in the cape province, volume 2 – the eastern half of the cape province, including the ciskei, transkei and east griqualand, dept. of nature and environmental conservation of the provincial administration of the cape of good hope, cape town. skinner, j.d. & cimimba, c.t., 2005, the mammals of the southern african subregion, university of pretoria press, pretoria. struhsaker, t.t., 1967, ‘ecology of vervet monkeys (cercopithecus aethiops) in the masai-amboseli game reserve, kenya’, ecology 48, 891–904. http://dx.doi.org/10.2307/1934531 tichý, l., rolocek, d., zeleny, c. & chytry, m., 2007, new stopping rules for twinspan, european vegetation survey, 16th workshop, march 22–27, 2007. turner, t.r., anapol, f. & jolly, c.j., 1997, ‘growth, development and sexual dimorphism in vervet monkeys (cercopithecus aethiops) at four sites in kenya’, american journal of physical anthropology 103, 19–35. http://dx.doi.org/10.1002/(sici)1096-8644(199705)103:1<19::aid-ajpa3>3.0.co;2-8 tchoukanski, i., 2010, et geo wizards. version 10.0 for arcgis, viewed on 13 june 2011, from http://www.ian-ko.com whiten, a., byrne, r.w. & henzi, s.p., 1987, ‘the behavioural ecology of mountain baboons’, international journal of primatology 8, 367–388. http://dx.doi.org/10.1007/bf02737389 whitten, p.l., 1983, ‘diet and dominance among female vervet monkeys (cercopithecus aethiops)’, american journal of primatology 5, 139–159. http://dx.doi.org/10.1002/ajp.1350050205 willems, e.p., 2009, ‘from space to species: integrating remotely sensed information on primary productivity into investigations and systems models of vervet monkey (cercopithecus aethiops) socio-ecology’, phd thesis, department of anthropology, durham university. willems, e.p. & hill, r.a., 2009, ‘a critical assessment of two species distribution models: a case study of the vervet monkey (cercopithecus aethiops)’, journal of biogeography 36, 2300–2312. http://dx.doi.org/10.1111/j.1365-2699.2009.02166.x wolfheim, j.h., 1983, primates of the world: distribution, abundance and conservation, university of washington press, seattle and london. appendix top ↑ appendix 1: food species and plant parts consumed by vervet monkeys at samara game reserve. article information authors: elizabeth h. bouchard1 lawrence e. little2 cassandra m.l. miller3 susan m. rundell4 elana m. vlodaver5 kristine maciejewski6,7 affiliations: 1department of environmental science, wheaton college, united states 2department of biology, washington university, united states 3department of biology, grinnell college, united states 4department of ecology and evolutionary biology, yale university, united states 5department of biology, bowdoin college, united states 6organisation for tropical studies, cape town, south africa 7department of biological sciences, university of cape town, south africa correspondence to: kristine maciejewski email: krismacski@gmail.com postal address: 407 rusdon park, college road, rondebosch 7701, south africa dates: received: 25 may 2015 accepted: 17 aug. 2015 published: 08 oct. 2015 how to cite this article: bouchard, e.h., little, l.e., miller, c.m.l., rundell, s.m., vlodaver, e.m. & maciejewski, k., 2015, ‘undeclared baggage: do tourists act as vectors for seed dispersal in fynbos protected areas?’, koedoe 57(1), art. #1323, 9 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1323 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. undeclared baggage: do tourists act as vectors for seed dispersal in fynbos protected areas? in this original research... open access • abstract • introduction • research method and design    • study area    • interview survey    • plant seed collection and categorisation    • alien plant survey    • data analysis • results    • seed survey and collection    • vegetation survey • discussion • conclusion • acknowledgments    • competing interests    • authors’ contributions • references • appendix 1 abstract top ↑ encroachment by alien species is the second greatest threat to biodiversity worldwide. as south africa's cape floristic region has a botanical endemism of nearly 70%, conservation efforts are a high priority. estimates suggest that alien species cost the country over r6.5 billion per year. despite significant research on alien species dispersal, the role of tourists as seed dispersers requires further exploration. to investigate the potential role tourists play in introducing alien seeds into protected areas, long-bristle brushes were used to scrape seeds off the shoes of hikers, dog walkers and cyclists, as well as the wheels of mountain bikes and dogs themselves, upon entering the silvermine nature reserve section of the table mountain national park in the western cape province, south africa. in addition, a vegetation survey was conducted. this comprised 18 transects at various distances from the recreational paths in the park, and used a prioritisation ranking system that identified the alien species of greatest concern. it was concluded that the greatest number of alien plant species could be found along dog paths, in comparison to the hiking trails and cycling trails. this corresponded to the findings that dog walkers had the highest incidence of seeds on their shoes, suggesting that tourists were possibly dispersing seeds from their gardens. alien species significantly covered more of the vegetation transects closer to the trails than they did in transects further into the matrix. because more alien species were present in areas susceptible to human disturbance, the data suggest that tourists can act as vectors for alien seed dispersal. these findings emphasise the need for active tourism management in line with the south african national parks biodiversity monitoring programme in order to prevent the introduction and spread of alien species into south africa's protected areas. conservation implications: tourism is the main source of revenue for south african national parks, and one of the organisation's principal goals is to create a tourism management policy conducive to conservation. this research explores the potential role that tourists may play in the introduction of non-native species into a protected area, thereby providing novel information that could assist managers in the sustainable management of protected areas. introduction top ↑ south africa's cape floristic region is a world-renowned hotspot for botanical biodiversity, home to some 9000 plant species, 70% of which are endemic (forest et al. 2007). however, alien species – the second greatest threat to biodiversity worldwide (richardson & van wilgen 2004) – jeopardise the long-term survival of this region's staggering diversity. an alien species can be defined as a non-native species, the introduction of which has the potential to cause health-related, economic or environmental damage (swearingen et al. 2010). in south africa alone, nearly 200 plant taxa are legally recognised as ‘alien’ (wilson et al. 2013). these species degrade ecosystems through the excessive consumption of resources, the alteration of fire regimes, and the disruption of soil stability and composition (richardson & van wilgen 2004). alien species cost south africa an average of r6.5 billion per year, a sum set to grow by an order of magnitude in the near future (wilson et al. 2013). the management of alien species is, therefore, vital to the biological and economic well-being of the country (wilson et al. 2013). given that south africa is home to 3 of the world's 25 biodiversity hotspots, managing alien species presents a particularly daunting task due to the already established vulnerability of these areas (myers et al. 2000). south africa has one of the worst problems with alien flora of any country in the world, and managers have never successfully eradicated an alien plant species nationwide (richardson & van wilgen 2004; wilson et al. 2013). one of the challenges of containment and eradication is that, while managers may successfully clear above-ground individuals from a plot, the alien seed bank can persist in the soil for many decades, encouraging reinvasion (wilson et al. 2011). accordingly, a more effective management strategy is the prevention of seed dispersal (wilson et al. 2011). the introduction of any new alien species into a protected area is against the mandate of south african national parks (sanparks), even when the potential for a particular species to become invasive is unknown (foxcroft 2009). as such, one of the key functions of the sanparks biodiversity management programme for invasive alien species is the monitoring of potential introduction pathways (mcgeoch et al. 2011). research on such pathways has expanded from such vectors such as animal transport and wind to include the role of tourists (pickering et al. 2011). tourists are often cited as playing a role in the dispersal of seeds through activities such as hiking and mountain biking. however, there is only a small body of evidence to support these claims. to date, only a single study on seed dispersal by tourists has been conducted in the entire continent of africa (pickering & mount 2010). circumstantial evidence suggests that alien species are found most commonly near paths in protected areas, and a correlation exists that links an increased visitor presence to a greater number of alien species in protected areas in southern africa (usher 1988). this emerging field has no standardised techniques for collecting seed from clothing, vehicles, animals, or equipment used by tourists, limiting the utility of comparisons between data sets (pickering & mount 2010). despite the dearth in research on the subject, it is likely that tourists do have a profound impact on the dispersal of alien seeds. hikers, for instance, have been shown to carry seeds on their clothing for an average of 13 km, and much further if their travels include a car, aeroplane, train or boat (ansong & pickering 2014a). protected areas are usually associated with minimal artificial disturbance, reducing the impact of certain other pathways, which highlights the importance of managing this particular pathway of introduction (pickering & mount 2010; usher et al. 1988). tourism and tourism-related activities comprise upwards of 80% of sanparks’ annual revenue, distinguishing it as one of the most successful protected area networks in the world (biggs et al. 2014). nonetheless, there are ecological risks associated with an increase in visitor numbers, which highlights the need for environmentally responsible tourism (cini & saayman 2014). one of the goals of sanparks’ strategic adaptive management process is to find a balance between the goals of increased ecotourism and the needs of conservation (biggs et al. 2014). achieving this goal will require greater investigation into the role that tourists play in ecosystem processes, and integrating the results of such studies into tourism management (biggs et al. 2014). a study on the role that tourists play in the dispersal of alien plants presents an opportunity to inform and improve tourism management practices in a way that protects the native flora of south africa's iconic national parks. the aim of this study is to investigate the role that three categories of tourists – hikers, dog walkers and cyclists – play in alien seed dispersal in a fynbos protected area. by sampling the shoes of visitors, dogs and bicycles for seeds, the role that visitors play in introducing seeds into a protected area is assessed, as well as the types of tourists that act as vectors for seed dispersal. at the same time, vegetation transects at different distances from trails used by these tourists illustrate the distribution of established alien species within the protected area. since roads have been shown to act as propagules for alien species in protected areas globally (pauchard & alaback 2004), a hypothesis is put forward that greater tourist disturbance closer to trails leads to a greater percentage of cover of alien species in these areas as opposed to areas further into the matrix. research method and design top ↑ study area the cape peninsula has a mediterranean climate, with an average winter rainfall of approximately 2000 mm per year (trinder-smith 2006). furthermore, the region is characterised by strong gulf winds, blowing primarily south-east to north-west. the table mountain national park covers 57 km2 and offers over 650 km of hiking trails (clarke, mackensie & merry 2013). the silvermine nature reserve section of the table mountain national park, in cape town, western cape province, south africa (18°40's, 34°07’e), is known for its diverse fynbos vegetation (figure 1). the soil is very sandy and nutrient poor, with limited phosphorus and nitrogen availability (trinder-smith 2006). this section of the park provides various trails for visitors to use, including paths for dog-walking, cycling and hiking. of the three trails considered in this study, dogs were restricted to the dog-walking trail, as were cyclists to the cycling trail. hikers, however, could walk on all three trails. figure 1: location study area within the silvermine section of the table mountain national park, western cape province, south africa. the yellow, blue and red rings represent the vegetation transects conducted along the dog-walking, hiking and cycling trails respectively. the stretches of the dog-walking and cycling paths investigated in this study were wider than the hiking path. furthermore, the hiking path utilised in this study was one of the most commonly trafficked trails in the park, which also serves as the start to many other hiking routes. interview survey data were collected for 3 days in early february 2015 – the end of the dry season – during peak visiting hours (between 08:00 and 11:00). whilst it was partly cloudy during the first 2 days of data collection, it rained heavily on the third day, resulting in a decrease in the number of participants and overall park traffic on that day. survey stations were set up at the trailhead of three different paths in silvermine: a hiking path (18°40's, 34°08’e), a cycling path (18°41's, 34°08’e) and a dog-walking path (18°40’ s, 34°08’ e). tourists entering the park were classified into three categories: hikers, dog walkers and cyclists. each participant was asked a series of questions about their activities (see appendix 1). the data were further assessed to determine tourists’ role in seed dispersal, and the introduction and potential source of alien vegetation into protected areas. plant seed collection and categorisation at the three survey stations, prior to entering the park, each participant stepped into a 40 cm × 30 cm plastic container. here, the soles of shoes were swept, using a long-bristle brush to collect any seeds the individual may have been carrying. the debris was collected into individual plastic bags for later classification. this same procedure was followed to brush off the entirety of the front tyre of each bicycle that was surveyed. the front legs and flank of each dog surveyed was brushed using a dog brush and all the debris collected. each debris sample was then examined under an olympus ch light microscope (model lsk [20w], olympus corporation of america, new hyde park, ny, usa). all of the seeds in these debris samples were counted and categorised by morph. an attempt was made to germinate the seeds in order to identify the species from which each morph had come. however, this was unsuccessful, preventing seed identification beyond the original morph categorisation. alien plant survey in order to examine the occurrence of alien plants in protected areas, eighteen 10-m transects were set up along recreational trails in silvermine. alien species were defined as those that are non-native to south africa. to compare the potential for different human activities at dispersing alien seeds, three paths were selected: a dog-walking path, a hiking path and a cycling path. six transects were set up in two groups on each trail. using a measuring tape, the first set of 10-m transects was set up 5 m from the trailhead; the second group of transects, 100 m from the trailhead (figure 2). to evaluate the relationship between the percentage of cover of plant species, and distance from the trail, three vegetation transects were conducted: one along the edge of the trail; one 1 m away; and a third, 10 m away from the trail (figure 2). figure 2: experimental design of vegetation transect survey conducted in the silvermine section of the table mountain national park in order to investigate the distribution of invasive plants. three 1-m2 quadrats, each represented by a grey square, were sampled along a 10-m line transect. this technique was repeated at 5 m (a, b and c) and 100 m (a, b and c) from the trailhead. at both of these locations, transects were established at the edge of the trail (1), 1 m into the matrix (2) and 10 m into the matrix (3). at each vegetation transect point, a 1 m2 quadrat was used to examine the vegetation cover every 5 m along each line transect, for a total of three plots. the percentage of cover for all plant species within the quadrats was estimated by identifying the known species to at least genus level and collecting samples that were later identified at a laboratory. to approximate the relative abundances of each plant, the estimated proportion of the plot covered by each particular species was done visually. to assess the immediacy and severity of the risks posed by the identified alien species, the prioritisation system was used to score each species and to rank them accordingly (robertson et al. 2003). a prioritisation score was calculated for each species based on an assessment of the impact the species poses in a range of categories. these included long-distance dispersal mechanism, invasive elsewhere, density, biodiversity, impact on water resources, and poison status. each category score was first standardised by dividing it by the potential maximum (robertson et al. 2003). these category scores were summed up and divided by the number of categories, resulting in a weighting score. species were then ranked according to the product of their total prioritisation score and total confidence interval. a score of 5 represented a species that posed the greatest threat, whilst a score of 0 demarcated no threat. data analysis to determine the proportion of seeds found per vector, and to account for sampling effort, the number of seeds collected per vector was divided by the sample size of each vector. a permanova analysis was conducted to analyse the difference between percentage of cover of alien plant species between trail types, distance from the trailhead, and distance into the matrix (primer software version 6.1.5, clarke & gorley 2006). results top ↑ seed survey and collection in order to examine their potential as vectors for seed dispersal, a survey and debris collection was conducted among 68 participants entering silvermine nature reserve. of these 68 participants, 10 were cyclists, 12 were dog walkers (with 19 dogs in total) and 46 were hikers. all of the cyclists and dog walkers lived in the cape town area. of the 46 hikers, 16 were either non-local south africans, living in johannesburg or port elizabeth, or international visitors from scotland, england or germany. fifty of the sampled visitors were not carrying any seeds that could be detected. from the remaining 18 participants, a total of 41 seeds and 17 unique seed morphs were isolated (table 1). it was not possible to identify the seeds to species level, and it therefore remains unknown if these were native or non-native seeds. table 1: the total number and morph of seeds collected from tourists (cyclists, dog walkers and hikers) entering the silvermine section of the table mountain national park, and their potential source of origin. of the participants who were carrying seeds, 16 were from the cape town area, one was visiting from england (9 seeds) and one from germany (1 seed). the english hiker carrying 9 seeds had last worn his shoes in london, and the german hiker had last worn his shoes at lion's head, a different part of the table mountain national park. no seeds were found on bicycle tyres, but seeds of two distinct morphs were found on a single cyclist's shoes (table 1). when sampling effort and population sample size were taken into account, it was discovered that a larger proportion (75%) (9 out of 12 that were sampled) of dog walkers carried seeds than any other vector, comprising 42% of seed morphs (table 1; figure 3). hikers were shown to be the second highest seed disperser type. of the hikers, 56% (27 out of 46 that were sampled) carried seeds, representing 19.6% of the total seed morphs. only 20% of the cyclists (2 out of 10 that were sampled) carried seeds on their shoes, whilst 15% of the dogs (3 out of 19 that were sampled) carried seeds (figure 3). figure 3: the proportion of seeds collected from each vector, calculated from the number of seeds divided by vector sample size, to account for sampling effort and population sample size. the black line represents the number of different seed morphs found per vector as a proportion of total seed morphs found. vegetation survey a total of 54 plots in 18 line transects along the three selected trails in silvermine were surveyed, and a total of seven invasive plant species were documented (table 2). the most frequently occurring invasive species were briza maxima (greater quaking grass, found in 17 quadrats), lagurus ovatus (hare's tail, found in 7 quadrats) and stenotaphrum secundatum (coastal buffalo grass, found in 6 quadrats) (table 2). the dog path had the greatest number of invasive plant species (n = 6), whilst the mountain bike path had the fewest invasive plant species (n = 2). furthermore, the dog-walking path had the highest average percentage of cover of invasive species (13.8%), whilst the hiking path had 9.6% and the cycling path had only 3.0% average percentage of cover. table 2: the percentage of cover of plant species identified to family or genus level, averaged from three 1-metre2 quadrats used in the vegetation transects along the various trails in the silvermine section of the table mountain national park. the permanova analysis demonstrated a significant increase in the percentage of cover of alien plant species in the plots located at the trail edge compared to further into the vegetation matrix (pseudo-f = 3.392, p < 0.005, n = 18; figure 4). there was also a significantly higher percentage of cover of alien plants along the dog-walking path than the cycling path (pseudo-t = 2.592, p < 0.05, n = 12). there was no significant difference, however, between plots taken 5 m and 100 m along the path for all trails. figure 4: the average proportional cover of alien plant species along three different paths in the silvermine section of the table mountain national park, taken at 0 m, 1 m and 10 m into the matrix and measured at 5 m and 100 m from the trailhead. the highest ranking species with regards to the threatening status of alien species identified in this study obtained a score of 2.74 (giant cane, arundo donax), whilst the lowest ranking species (l. ovatus) scored 1.56 (table 3). species that had high prioritisation and confidence scores, such as a. donax, were of most concern (confidence score = 11.16; table 3). the high ranking of this species was mainly due to its severe impact on biodiversity (giessow et al. 2011; guthrie 2007) and its negative economic impact. this was followed closely by s. secundatum (prioritisation score = 2.54, confidence score = 9.479) and b. maxima (prioritisation score = 2.22, confidence score = 9.527) (table 3). table 3: the prioritisation scores of alien plants species surveyed on the trails in the silvermine section of the table mountain national park. the scores were calculated using the alien species prioritisation system. discussion top ↑ this study demonstrates that park visitors, in the form of dog walkers, hikers and cyclists, act as seed dispersers in protected areas, corroborating existing research. one study found that humans working in a series of meadows carried seeds from this location on their clothing for an average distance of 13 km (auffret & cousins 2013). furthermore, it has been demonstrated that seeds can remain on hikers’ shoes for distances as great as 5 km (wichmann et al. 2009). the seeds of 16 taxa were found in a single study specifically investigating dispersal via shoes (clifford 1956). the vehicles considered in studies as a means of seed dispersal, however, have been exclusively motorised, and the only animals investigated have been donkeys and horses (ansong & pickering 2014a; pickering & mount 2010). the finding that the areas closest to human activity (i.e. near the edge of the trails) are also the areas that are most susceptible to invasion from non-native plants suggests a path-user-mediated mechanism for alien seed dispersal. this evidence is strengthened by a correlation between the relative importance of different potential vectors of seed dispersal and the percentage of cover of alien species on different paths. for instance, dog walkers had the highest incidence of seeds on their shoes; correspondingly, the dog-walking path had the highest percentage of cover of alien plants. likewise, cyclists carried virtually no seeds, and the cycling trail had the lowest percentage of cover of alien species. the difference in tendency for different categories of tourists for carrying seeds into the park, therefore, suggests a possible explanation for the differences in the percentage of cover between paths. whilst the sample sizes used in this study were too small to establish an unambiguous relationship between seed incidence on trail users and the percentage of cover of alien plants along these trails, it is an observation worthy of further investigation. to confirm such a relationship, future studies need to identify the species of collected seeds, particularly since the attempts in this study to propagate and identify seeds were unsuccessful. even though the non-native species identified along the sides of the trails are all classified as ‘alien’, they all play different roles in invading or threatening the environment. in this study, the alien cane species a. donax represented the largest alien risk compared to the other species, which was also found by robertson et al. (2003) where this species received a total prioritisation score of 3.08. arundo donax, a tall perennial cane native to eastern asia, has been introduced into many areas for erosion control (mariani et al. 2010). however, it has a tendency to outcompete native plant species, reduce wildlife habitat and modify river hydrology (giessow et al. 2011). invasion by a. donax alters biomass, thereby increasing the fuel load and creating a potential fire hazard (guthrie 2007). even though this species only covered a small area in the hiking trail 100 m away from the trailhead, a monitoring programme is needed to prevent further invasion. this species may easily spread to other areas since reproduction occurs almost entirely from rhizomes and stem fragments (boose & holt 1999). significantly, a number of the invasive species that were found along the paths’ edge are used in southern africa for decorative purposes, including b. maxima and l. ovatus (van oudtshoorn 2002). furthermore, pauchard and alaback (2004) found that the land-use type surrounding protected areas influences the numbers and types of plants that are able to invade protected areas. the highly developed landscape surrounding silvermine is likely rich in decorative plants, considering the large anthropocentric role in determining the ecological makeup of urban areas (aronson et al. 2014). together, these findings suggest that tourists are possibly dispersing seeds from their gardens, a source of alien species now recognised as one of the most challenging to combat (mack 2003; mack & lonsdale 2001). the spread of ornamental grasses such as b. maximus and l. ovatus has been attracting greater attention in south africa (milton 2004), and researchers have begun to investigate the threat posed to south african protected areas by ornamental plant species (foxcroft, richardson & wilson 2008). it seems likely that this same mechanism is operating in silvermine. priorities for future studies should include more comprehensive seed collection techniques. through the course of this study, it was observed that more seeds appeared to become caught on the socks of researchers rather than on the shoes. this observation suggests that clothing other than shoes could be important mechanisms for dispersal. the socks of participants were not a priority in this study because socks are typically washed between each use. it was therefore less likely that more seeds could be brought into silvermine on socks than on shoes, which was the primary focus of this study. socks may play a more important role in dispersing seeds within protected areas than in introducing seeds into those protected areas. in addition, car tyres represent a potential vector for invasive seeds that remains unexamined in the context of south african protected areas. future studies should take this into account when considering the method of seed collection – it is suggested that the clothes and vehicles of participants are sampled in addition to shoes. to limit the spread of non-native plant species into pristine ecosystems, protected area managers must take an active role in monitoring tourists in accordance with the sanparks biodiversity management programme for invasive alien species (mcgeoch et al. 2011). tourism is the main source of revenue for south african protected areas, and a major goal for sanparks is to encourage ecotourism that does not compromise the agency's commitment to conservation (biggs et al. 2014). some research on the means of minimising tourist-mediated seed dispersal has already been conducted in other countries. a study by ansong and pickering (2014b) found that the vast majority of tourists entering an australian protected area were aware of the potential capacity for unintentionally dispersing invasive plant seeds. over half of the individuals surveyed by ansong and pickering (2014b) supported greater funding toward non-native species removal, and a vast majority of respondents either supported or felt neutral towards an initiative requiring tourists to remove seeds from their clothing and equipment before entering the park (ansong & pickering 2014b). these results indicate that an educational campaign about seed dispersal by tourists could have widespread public support (ansong & pickering 2014b). primarily, these efforts should focus on the correct form of disposal for seeds attached to clothing, shoes, dogs or recreational equipment, since many tourists interviewed said they threw seeds away indiscriminately (ansong & pickering 2014b). information posters at trail heads about the dangers of alien species and the correct method for the removal of seeds could be one easy method of raising awareness and decreasing the incidence of unintentional dispersal. shoe-sweeping stations in trailhead parking lots could similarly provide a cost-effective and convenient method for tourists to remove seeds from their shoes. the implementation of such measures could help to prevent the further spread of non-native plants species into silvermine nature reserve, and elsewhere in south africa's national parks. conclusion top ↑ this research adds to the small body of existing evidence concerning the spread of invasive seeds by tourists. tourist surveys and vegetation transects conducted in the silvermine nature reserve section of the table mountain national park demonstrate a clear link between tourist disturbance – measured as distance from the path into the matrix – and percentage of cover of invasive plants. at the same time, the path used by dog walkers, who were found to have the highest load of seeds, have the highest overall percentage of cover of invasive species. these results demonstrate that tourists can act as vectors for invasive alien species dispersal, highlighting the need for active tourism management in line with sanparks’ biodiversity monitoring programme. acknowledgments top ↑ the authors would like to thank dr laurence kruger for his assistance in plant identification. they would also like to thank the organisation for tropical studies and sanparks for their roles in facilitating this project. competing interests the authors declare that they have no financial or personal relationships, which may have inappropriately influenced them in writing this article. authors’ contributions k.m. (organisation for tropical studies) was the project leader and responsible for project design. e.h.b. (wheaton college), l.e.l. (washington university), c.m.l.m. (grinnell college), s.m.r. (yale university) and e.m.v. (bowdoin college) were responsible or conducting the fieldwork and the analyses of data. references top ↑ ansong, m. & pickering, c., 2014a, ‘weed seeds on clothing: a global review’, journal of environmental management 144, 203–211. pmid: 24956465, http://dx.doi.org/10.1016/j.jenvman.2014.05.026 ansong, m. & pickering, c., 2014b, ‘weed hygiene: what do we do with seeds we find on our clothing?’, nineteenth australasian weeds conference, pp. 42–45. aronson, m.f., la sorte, f.a., nilon, c.h., katti, m., goddard, m.a., lepczyk, c.a. et al., 2014, ‘a global analysis of the impacts of urbanization on bird and plant diversity reveals key anthropocentric drivers’, proceedings of the royal society b: biological sciences 281(1780), 20133330. pmid: 24523278, http://dx.doi.org/10.1098/rspb.2013.3330 auffret, a.g. & cousins, s.a.o., 2013, ‘humans as long-distance dispersers of rural plant communities’, plos one 8(5), e62763. pmid: 23658770, http://dx.doi.org/10.1371/journal.pone.0062763 biggs, d., swemmer, l., phillips, g., stevens, j., freitag, s. & grant, r., 2014, ‘the development of a tourism research framework by south african national parks to inform management’, koedoe 56(2), art. #1164, 9 pages. http://dx.doi.org/10.4102/koedoe.v56i2.1164 boose, a.b. & holt, j.s., 1999, ‘environmental effects on asexual reproduction in arundo donax’, weed research 39, 117–127. http://dx.doi.org/10.1046/j.1365-3180.1999.00129.x cini, f. & saayman, m., 2014, ‘which age group spends the most in a national park?’, koedoe 56(2), art. #1158, 8 pages. http://dx.doi.org/10.4102/koedoe.v56i2.1158 clarke, k.r. & gorley, r.n., 2006, primer version 6.1.5 user manual/tutorial, plymouth, primer-e ltd, uk. clarke, h., mackensie, b. & merry, c., 2013, common wild flowers of table mountain & silvermine, struik nature, cape town. clifford, h.t., 1956, ‘seed dispersal on footwear’, proceedings of the botanical society of the british isles 2, 129–131. forest, f., grenyer, r., rouget, m., davies, t.j., cowling, r.m., faith, d.p. et al., 2007, ‘preserving the evolutionary potential of floras in biodiversity hotspots’, nature 445, 757–760. pmid: 17301791, http://dx.doi.org/10.1038/nature05587 foxcroft, l.c., 2009, ‘developing thresholds for potential concern for invasive alien species: hypotheses and concepts’, koedoe 50(1), art. #157, 6 pages. http://dx.doi.org/10.4102/koedoe.v51i1.157 foxcroft, l.c., richardson, d.m. & wilson, j.r., 2008, ‘ornamental plants as invasive aliens: problems and solutions in kruger national park, south africa’, environmental management 41(1), 32–51. pmid: 17943344, http://dx.doi.org/10.1007/s00267-007-9027-9 giessow, j., casanova, j., leclerc, r., macarthur, r., fleming, g. & giessow, j., (else) 2011, arundo donax (giant reed): distribution and impact report, agreement no. 06-374-559-0, state water resources control board, california plant council (cal-ipc). guthrie, g., 2007, ‘impacts of the invasive reed arundo donax on biodiversity at the community-ecosystem level’, msc thesis, biodiversity and conservation biology department, university of cape town, cape town. mack, r.n., 2003, ‘global plant dispersal, naturalization, and invasion: pathways, modes, and circumstances’, in j. carlton (ed.), invasive species: vectors and management solutions, pp. 3–30, island press, washington, dc. mack, r.n. & lonsdale, w.m., 2001, ‘humans as global plant dispersers: getting more than we bargained for: current introductions of species for aesthetic purposes present the largest single challenge for predicting which plant immigrants will become future pests’, bioscience 51(2), 95–102. http://dx.doi.org/10.1641/0006-3568(2001)051[0095:hagpdg]2.0.co;2 mariani, c., cabrini, r., danin, a., piffanelli, p., fricano, a., gomarasca, s. et al., 2010, ‘origin, diffusion and reproduction of the giant reed (arundo donax l.): a promising weedy energy crop’, annals of applied biology 157, 191–202. http://dx.doi.org/10.1111/j.1744-7348.2010.00419.x mcgeoch, m.a., dopolo, m., novellie, p., hendriks, h., freitag, s., ferreira, s. et al., 2011, ‘a strategic framework for biodiversity monitoring in south african national parks’, koedoe 53(2), art. #991, 10 pages. http://dx.doi.org/10.4102/koedoe.v53i2.991 milton, s.j., 2004, ‘grasses as invasive alien plants in south africa’, south african journal of science 100(1/2), 69–75. https://www.dwa.gov.za/wfw/docs/milton,2004.pdf myers, n., mittermeier, r.a., mittermeier, c.g., da fonseca, g.a.b. & kent, j., 2000, ‘biodiversity hotspots for conservation priorities’, nature 403, 853–858. pmid: 10706275, http://dx.doi.org/10.1038/35002501 pauchard, a. & alaback, p.b., 2004, ‘influence of elevation, land use, and landscape context on patterns of alien plant invasions along roadsides in protected areas of south-central chile’, conservation biology 18, 238–248. http://dx.doi.org/10.1111/j.1523-1739.2004.00300.x pickering, c. & mount, a., 2010, ‘do tourists disperse weed seed? a global review of unintentional human-mediated terrestrial seed dispersal on clothing, vehicles and horses’, journal of sustainable tourism 18(2), 239–256. http://dx.doi.org/10.1080/09669580903406613 pickering, c., mount, a., wichmann, m.c. & bullock, j.m., 2011, ‘estimating human-mediated dispersal of seeds within an australian protected area’, biological invasions 13(8), 1869–1880. http://dx.doi.org/10.1007/s10530-011-0006-y richardson, d.m. & van wilgen, b.w., 2004, ‘invasive alien plants in south africa: how well do we understand the ecological impacts?’, south african journal of science 100(1/2), 45–52. http://dx.doi.org/10.1007/s10021-012-9605-4 robertson, m.p., villet, m.h., fairbanks, d.h.k., henderson, l., higgins, s.i., hoffmann, j.h. et al., 2003, ‘a proposed prioritization system for the management of invasive alien plants in south africa’, south african journal of science 99, 37–43. swearingen, j., reshetiloff, k., slattery, b. & zwicker, s., 2010, plant invaders of mid-atlantic natural areas, national park service and u.s. fish & wildlife service, washington, dc. trinder-smith, t., 2006, wild flowers of the table mountain national park: south african wild flower guide 12, botanical society of south africa, cape town. usher, m.b., 1988, ‘biological invasions of nature reserves: a search for generalizations’, biological conservation 44, 119–135. http://dx.doi.org/10.1016/0006-3207(88)90007-9 usher, m.b., kruger, f.j., macdonald, i.a.w., loope, l.l. & brockie, r.e., 1988, ‘the ecology of biological invasions into nature reserves: an introduction’, biological conservation 44, 1–8. http://dx.doi.org/10.1016/0006-3207(88)90002-x van oudshoorn, f., 2002, guide to grasses of southern africa, briza publications, pretoria. wichmann, m.c., alexander, m.j., soons, m.b., galsworthy, s., dunne, l., gould, r. et al., 2009, ‘human-mediated dispersal of seeds over long distances’, proceedings of the royal society b: biological sciences 276, 523–532. pmid: 18826932, http://dx.doi.org/10.1098/rspb.2008.1131 wilson, j.r.u., gairifo, c., gibson, m.r., arianoutsou, m., bakar, b.b., baret, s. et al., 2011, ‘risk assessment, eradication, and biological control: global efforts to limit australian acacia invasions’, diversity distribution 17(5), 1030–1046. http://dx.doi.org/10.1111/j.1472-4642.2011.00815.x wilson, j.r.u., ivey, p., manyama, p., nänni, i., 2013, ‘a new national unit for invasive species detection, assessment and eradication planning’, south african journal of science 109(5/6), art. #0111, 13 pages. http://dx.doi.org/10.1590/sajs.2013/20120111 appendix 1 top ↑ interview questionnaire presented to participants at the entrance of visitor trails in the silvermine section of the table mountain national park where are you from? what are you doing? cycling walking/hiking/dog-walking where did you last: wear your shoes? ride your bike? walk your dog? article information authors: edward s. riddell1 ahmed khan1 benjamin mauck1 simphiwe ngcobo1 jonathan pasi1 andrew pickles1 jennifer pickles1 zinhle sithole1 simon a. lorentz1 navashni govender2 affiliations: 1school of agriculture, earth and environmental science, university of kwazulu-natal, pietermaritzburg campus, south africa 2scientific services, south african national parks, kruger national park, south africa correspondence to: edward riddell postal address: private bag x01, scottsville 3209, south africa dates: received: 23 dec. 2011 accepted: 08 may 2012 published: 27 july 2012 how to cite this article: riddell, e.s., khan, a., mauck, b., ngcobo, s., pasi, j., pickles, a. et al., 2012, ‘preliminary assessment of the impact of long-term fire treatments on in situ soil hydrology in the kruger national park’, koedoe 54(1), art. #1070, 7 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1070 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. preliminary assessment of the impact of long-term fire treatments on in situ soil hydrology in the kruger national park in this original research... open access • abstract • introduction • research method and design • results • trustworthiness    • reliability    • validity • discussion • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ there has been significant attention focused on the impacts of fire frequency and season of burn on ecological processes in the kruger national park (knp). whilst there has been some examination of these fire effects on soil properties, the explicit linkages of these effects to the hydrology of soils in burnt areas has remained a gap in our understanding. during august 2010, a field scoping campaign was undertaken to assess the impacts, if any, of long-term fire treatments on the hydrology of soils on the experimental burn plots (ebps) in the knp. using various hydrometric and soil physical characterisation instruments soil, hydraulic conductivity and soil strength variations were determined across the extreme fire treatment on the ebps, the annual august (high fire frequency) plots and the control (no burn) plots, on both the granite and basalt geologies of pretoriuskop and satara, respectively. it was found that there were soil hydrological and structural differences to fire treatments on the basalt burn plots, but that these were not as clear on the granite burn plots. in particular, hot, frequent fires appeared to reduce the variation in soil hydraulic conductivity on the annual burn plots on the basalts and led to reduced cohesive soil strength at the surface.conservation implications: the knp burn plots are one of the longest running and well studied fire experiments on african savannahs. however, the impacts of fire management on hydrological processes in these water-limited ecosystems remains a gap in our understanding and needs to be considered within the context of climate and land-use changes in the savannah biome. introduction top ↑ savannahs are tropical grasslands with scattered trees; they occupy about 20% of the land surface of the earth and 40% of africa (scholes & hall 1996). these ecosystems are dynamic in their structure, composition and function. fire is an important driver in savannah ecosystems, where it acts as both a generalist herbivore (bond & keeley 2005) and as a facilitator of the coexistence of trees and grasses (higgins, bond & trollope 2000). the spatio-temporal availability of water is also considered a key driver of savannah heterogeneity (walter 1971). studies of catchment hydrological processes response to natural and induced fires have received the greatest attention in various land-systems within australia (e.g. brown 1972; townsend & douglas 2000). studies elsewhere include mountain catchments under exotic forestry (scott 1997) and fynbos (lindley, bosch & van wyk 1988) in south africa; however, few studies have examined explicitly pyro-hydrological interactions and the role of various burning regimes within african savannahs (scholes & walker 1993).fire has long been used in the management of savannahs and this management has been informed and adapted by the findings of ongoing research on experimental sites of selected fire regimes on fixed areas (andersen, cook & williams 2003; knapp et al. 1998). the long-term fire experiment in the kruger national park (knp) was established in 1954, with the initial objectives being to study the effects of fire frequency and season on the vegetation of the knp under indigenous grazing pressure (van der schijff 1958). the experimental burn plots (ebps) were laid out in four of the 37 main vegetation landscapes of the knp described by gertenbach (1983): the pretoriuskop sourveld vegetation, skukuza combretrum vegetation, satara knobthorn and marula vegetation and mopani mopane vegetation. whilst there has been some examination of fire effects on in situ soil properties, the explicit linkages of these to the hydrological parameters of soils in burnt areas has remained a knowledge gap (pierson et al. 2011). to date, a variety of soil hydrological responses have been described; for instance, frequent burning reduced the rate of infiltration of rainwater into soil as noted in the drakensberg (bijker et al. 2001). this is caused by crusting of the soil, thereby yielding lower electrical conductivity as a result of the removal of ash via increased surface water runoff and increasing soil erosion and as shown particularly for the granite ebps in the knp (mills & fey 2004). webber (1979) showed that the unburnt control ebps near skukuza retained greater moisture levels well into the dry season and, where burning occurred on duplex soils, this made them susceptible to erosion. in light of these previous findings, the objectives of this study therefore were to determine the impacts, if any, of long-term high fire frequency and intensity treatments versus fire exclusion on in situ soil hydrological characteristics in the knp. moreover, this included an assessment of the effects of potential fire impacts on soil hydrology in the two distinct geologies of the knp: the granites and basalts. research method and design top ↑ the august 2010 field campaign focused on describing various in situ soil hydraulic and strength characteristics on these plots on the two extreme fire treatments, within an ebp string, on the two dominant geologies in the knp, namely the granites at pretoriuskop (lowveld sour bushveld, sandy soils with mean annual precipitation [map] of 737 mm) and basalts at satara (acacia nigrescens savannah, clay soils with map of 537 mm) (biggs et al. 2003). an ebp string is located generally along the crests of the hillslope and consists of twelve to fourteen 7 ha plots bounded by fire breaks to prevent natural and accidental fires from entering them. the fire treatments include winter august (dry season, hot), october (first spring rains), december (early summer wet season), february (late summer) and april (autumn) with annual, biennial and triennial treatments.because one string at both sites was deemed unrepresentative of the broader landscapes (venter & govender 2012), only three of the four strings at these sites were assessed, as shown in figure 1. a semi-systematic approach was taken in order to dismiss the inherent variability associated with the different soil morphologies linked to subtle changes in topography. predetermined areas were selected according to diagnostic soil form characteristics, identified by a soil scientist (le roux pers. comm., 06 march 2010), which resulted in attention to sites characteristic of the shortlands and huttons soil forms (soil classification working group 1991) which were identified as being dominant on the basalt and granite plots, respectively. the reader is referred to venter and govender (2012) for detailed descriptions of the soil morphologies on the ebps. hydrological measurements and soil characterisations were then taken randomly within these areas. a variety of techniques were used to determine the soils’ hydraulic conductivity within 30 cm of the soil surface. these included: • double-ring infiltrometers, to determine saturated soil conductivities (ksat) at the soil surface, according to the method of lorentz et al. (2003). • guelph permeameters, to determine ksat within the subsurface at 5 cm, 15 cm and 30 cm depths (eijkelkamp agrisearch equipment 2008). • tension disc infiltrometers, to determine unsaturated soil conductivities (kunsat) at the soil surface, according to the method of lorentz et al. (2003). two replicates of each of the above techniques were made at each ebp treatment per landscape and results are discussed simply based on visual interrogation of the data. sampling sites were selected in a semi-random manner by first identifying areas a minimum of 10 m from large trees and 15 m from the fire breaks and then randomly placing instruments on bare ground. the unconfined cohesive soil strength on the ebps was assessed using the following equipment: • a dynamic cone penetrometer (herrick & jones 2002), consisting of a 1 cm diameter rod, conical extremity of 90° and a 2 kg weight dropped from a height of 100 cm • a pocket penetrometer (eijkelkamp agrisearch equipment 2009) • a pocket vane tester (eijkelkamp agrisearch equipment 2011). these assessments were performed within a 5 m radius of the hydrology measurements and typically 10 readings were taken at each of the ebps randomly within the immediate vicinity of the soil hydrology measurements. the resulting data was then subject to non-parametric statistical analysis using the mann-whitney u-test for unmatched samples using the statext v. 1.4 software (2011). figure 1: the kruger national park experimental burn plot strings assessed in this study. results top ↑ the ksat values determined at the soil surface of the granite ebps clearly illustrate higher variation across all the plots and between fire treatments (figure 2). unfortunately, no measurements were taken on the numbi control plot; however, it was noted that, for both replicates, the shabeni control plots had significantly greater ksat values than the annual burn plots. at kambeni, the annual and control plots had similar ksat values, although in both replicates the values were slightly lower for the control. the ksat values for the basalt ebps had generally lower variation than the granite ebps (figure 3). with the exception of marheya replicate 1 (0.015 cm/s-1 beyond the axis range), there appeared to be no distinct difference in the ksat between annual or control treatments on the basalt ebps.the results of ksat tests at depths between 5 cm and 30 cm (as depicted in figure 2 and figure 3) revealed that soils on the granite ebps have, on average, a greater hydraulic conductivity on the annual frequent burn plots than the control plots; this was most noticeable at kambeni and shabeni. there was also a switch in the trend of ksat where annual burn plots had lower ksat at the soil surface than the control plots, a trend which reversed at depth. interestingly, it appeared that there was less variation in ksat at 5 cm within the control treatments on the granite ebps. the ksat distribution of the basalt ebps, meanwhile, was an order of magnitude lower than the granite ebps; however, there was no distinct difference in ksat between annual and control treatments in the soil sub-surface. figure 4 shows the kunsat values of the granite ebps soil surface, where, on average, the annual burn plots tended to have greater kunsat than the controls. also, there was a distinctly greater kunsat on the annual burn plots at the lowest soil moisture tensions close to saturation (0.5 cm). the opposite trend of the annual ebps having a lower kunsat on average than the controls was demonstrated on the basalt ebps (figure 5). the outliers to the above trend on the basalt ebps were the marheya string and, with the exception of the satara annual plot replicate, the controls had greater variation in kunsat than the annual burn plots. drop cone penetrometer tests for granite ebps revealed that the first approximate 150 j was the threshold for maximum depth penetration across annual and control plots; thereafter, there was a steady increase in depth at each energy increment (figure 6). the greatest variations in soil strength properties were noted for control plots compared to annual burn plots on the soil of the granite ebps. whilst with the basalt ebps (figure 7), it was the annual plots which again had distinctly less variation in soil strength compared to the control plots. an interesting aspect depicted in figure 7 was the slightly lower threshold in energy over depth, which was reached at about 130 j before further energy resulted in less depth penetration. table 1 presents the statistical results of the raw drop cone penetrometer data at two energy levels (2 kg × 2 kg weight drops, or 34 j/cm-2 and 10 kg × 2 kg weight drops, or 307 j/cm-2). there was no significant difference in the cohesive soil strength between control and annual burn plots on the granite ebps; however, the basalt ebps showed a significantly greater mean depth of penetration at both energy levels on the annual burn plots. the pocket penetrometer data supported this with no significant difference on the granite ebps, but there was a statistical difference on the basalt ebps. no significant difference was observed within the soil shear strength between annual and control treatments on both geological settings. trustworthiness top ↑ the data presented in these results are based on human measurements and thus aspects pertaining to the trustworthiness of the collected data need to be considered in the following contexts. figure 2: field saturated hydraulic conductivity versus depth on the granite experimental burn plots in the kruger national park. figure 3: field saturated hydraulic conductivity versus depth on the basalt experimental burn plots in the kruger national park. reliability repeatability of acquiring the same results of both the kunsat and ksat measurements is achieved from standard in situ hydrological measurements. however, replication of the results would require the placing of the instruments at precisely the same locations, otherwise inherent heterogeneities in the soil matrix would yield different results. a potential observer bias arises when determining whether kunsat and ksat is reached at a steady-state infiltration rate. to ensure that this bias inherently did not affect the collected data, we followed the convention of 10 equal time intervals per volume of water infiltrated interval (for further details consult lorentz et al. 2003).the same principle applies for soil strength characteristics determined via the drop cone penetrometer method, where consistency of measurements is achieved by the dropping of a known weight from a standard height on the penetrometer device. however, reliability of the data may be questioned should the penetrometer not be held vertically above the soil. care was taken during this study to ensure that each measurement was taken with a vertical deployment of the penetrometer device. similarly, this principle also applies to the pocket penetrometer and pocket vane tester. however, we did observe a potential effect of pressure applied to the soil surface and angle of contact by these devices, meaning that each operator has the potential to influence the outcome of the measurement. to minimise this risk, the same operator conducted these measurements during this study. validity 1 the validity of the reported measurements was believed by the authors to be appropriate to achieving the objectives of this scoping exercise within these specific study areas. nevertheless, significantly greater replication of these measurements should be performed in future to quantify the variability of results from significant soil surface spatial heterogeneities on the ebps and to answer comprehensively those causal mechanisms postulated in the following discussion. table 1: mann-whitney u-test results for unconfined cohesive soil strength parameters in the kruger national park. figure 4: unsaturated hydraulic conductivity at the surface of the granite experimental burn plots in the kruger national park. figure 5: unsaturated hydraulic conductivity at the surface of the basalt experimental burn plots in the kruger national park. discussion top ↑ these measurements reveal interesting differences across the soils associated with the two geologies of granite and basalt in relation to long-term fire and no fire treatments. in the absence of greater spatial and temporal replication, the interpretation at this stage is speculative, but sets the scene for future data collection and analysis. nevertheless, key aspects and questions from these initial interpretations and assessments should be explored further. as expected, and as a result of their finer texture soils, the basalt ebps have a ksat distribution of an order of magnitude lower than the granite ebps. meanwhile, there is generally greater kunsat at the surface on the granite ebps annual burn plots versus the control plots, than compared to the similarity observed within the basalt annual burn and control plots. furthermore, the ksat at the surface is distinctly higher than at depth on the annual burn plots of the granite ebps. these effects seen on the granite ebps could be attributed to the loss of organic residues at the soil surface caused by long-term fire treatments, which would explain these higher ksat and kunsat values of the annual plots. as demonstrated by fynn, haynes and o’connor (2003), organic carbon was significantly reduced within the first 2 cm of the soil profile under repeated grassland dry annual and biennial winter fire treatments at ukalinga in kwazulu-natal. similar results were described by mills and fey (2004) on the granite ebps at numbi, but also the napi plots in the skukuza ebp strings. recently, soil carbon was also shown to be lower on dry annual burn plots than the controls in the mopani ebps on basalt (holdo, mack & arnold 2011). whilst the n’wanetsi string, also on basalt and following 30 years of fire treatment, showed a quarter lower organic carbon content in the control than on the annual, biennial and triennial burn plots (jones et al. 1990). the observed switch in ksat trends between the surface and deeper layers, as noted on the granite ebps, is also a point of interest. the differences noted at depths below the surface, such as ksat and the soil strength, could simply be a result of inherent site differences such as soil morphological characteristics associated with different slope angles and position of measurements in the landscape; this is highly likely in the undulating landscape of pretoriuskop. this points to the need for greater replication and spatial coverage in future studies of this type to isolate for such heterogeneous influences in the analysis. however, it is feasible that the lower ksat on the annual plots on the granite ebps soil surface may result from changes in the soils sorptivity resulting from ash deposition, the effects of which can be markedly variable depending on the texture of the soil (see e.g. moody, kinner & úbeda 2009). this also points to the need in future to examine explicitly soil sorptivity characteristics resulting from fire. however, the basalt ebps show a smaller variation in ksat and kunsat on the annual treatments than the control plots, which may result from the development of a relatively homogenous soil layer structure on the more frequently burnt plots. this could be caused by a loss of organic matter and downward illuviation of finer material and ash deposits. the basis for this theory arises from two aspects. firstly, the influence and importance of organic matter content on soil sheer strength is further demonstrated by ekwue (1990), who showed that soil shear strength increased as a result of grass derived organic matter content. therefore, the annually burnt basalt plots of the knp have reduced soil strength as a result of a loss of organic carbon caused by frequent grassland vegetation fires at the site. this will make the soils on burnt areas within basalt both susceptible to erosion and have reduced soil moisture retention properties. the above results of deeper soil penetration per unit energy on the annual basalt plots described in this communication would support this hypothesis. secondly, debano (2000) discusses the role of fires in facilitating the development of a water repellent layer in the soil subsurface. the frequent annual burning on the basalt ebps could lead to the formation of a water repellent layer below the surface, rather than as a crust at the surface as suggested by mills and fey (2004). the small variation in ksat at 5 cm on the basalt annual burn plots compared to the control plots is supportive of this theory, but warrants more detailed assessment, replication and analysis. seemingly, the same effects could also have occurred on the coarser soils of the granite ebps. whilst these may not be apparent within the limited analysis described here, the intricacies of different fire severities associated with soils of different textures on changes in aggregate stability warrants further determination (for a review see mataix-solera et al. 2011). moreover, both granite and basalt ebps showed high compaction beyond 5 cm soil depth in the penetrometer analysis and a concomitant similarity in ksat between annual burn and control treatments beyond 5 cm. this suggests that fire effects on soil hydrology may be most significant within the first 5 cm of soil and should also be investigated further by analysis using parameters such as soil bulk density and rooting density, especially to complement further penetrometer data. results from this preliminary assessment of the impact of long-term fire treatments on in situ soil hydrology indicate that frequent hot fires lead to structural homogenisation of the soil surface particularly on basalt soils. interpretation of this dataset requires cognisance of other recent findings on the effects of fire frequency and season on the knp landscape. the reduction in woody vegetation cover as a result of frequent burning (smit et al. 2010) and increase in small tree dominance under frequent burning (higgins et al. 2007) are bound to have an effect on soil structure, as suggested by the scale dependent controls on biogeochemical processes resulting from tree canopy size (holdo et al. 2011). results will need to be discussed in this light. whilst most interpretations and explanations have focused on the vegetation (both woody and grass) feedbacks on soil hydrological properties through a fire driven system, changes in herbivore utilisation and density (e.g. burns et al. in review) and resultant trampling and nutrient deposition as a result of fire treatments also need to be considered. figure 6: drop cone penetrometer data from the granite experimental burn plots in the kruger national park. figure 7: drop cone penetrometer data from the basalt experimental burn plots in the kruger national park. conclusion top ↑ this study indicated interesting geologically specific feedbacks between fire and soil hydrology. these need to be more thoroughly determined in order to quantify the impacts of soil crusting and water repellency resulting from fire on plant available water in the savannah landscape. the ebp experiment provides a valuable opportunity to explore the role of fire management policies on soil hydrological properties at point and plot scales, with the added benefit that the experiment allows for the inter-comparison between geological settings. the results presented here have suggested that the effects of fire management on soil hydrology become more apparent on the finer texture basalt soils than the coarser granites. however, future studies of this nature should be based on a more comprehensive, replicated and multifactorial analysis. these should focus on quantifying the variation of pyro-hydrological interaction both between geologies, but also the nuances that may be associated with fire frequency and intensity of burn within geologies, as well as an explicit assessment of the influence of slope and landscape position, amongst other aspects. acknowledgements top ↑ the authors acknowledge the contributions from the following people: dr pieter le roux (university of the free state) for assisting with site selection based on soil diagnostics, renson thethe and thomas rikombe (both south african national parks) for protection whilst in the field and johannes hachmann (vrije universiteit, the netherlands) for providing additional assistance during the field campaign. the authors are also grateful for the useful comments provided by two anonymous referees. competing interests the authors declare that they have no financial or personal relationship(s) which may have inappropriately influenced them in writing this paper. authors’ contributions e.s.r. (university of kwazulu-natal) was responsible for the methodological design, data collection and analyses used in this study. he was also responsible for analysing the results and compiling the manuscript. a.k. (university of kwazulu-natal), b.m. (university of kwazulu-natal), s.n. (university of kwazulu-natal), j.p. (university of kwazulu-natal), a.p. (university of kwazulu-natal), j.p. (university of kwazulu-natal) and z.s. (university of kwazulu-natal) were responsible for data collection and analysis. s.a.l. (university of kwazulu-natal) contributed to the methodological design and was also responsible for data collection. n.g. (south african national parks) provided the team with background information on the ebps, logistical support during the campaign and made valuable contributing comments to the manuscript. references top ↑ andersen, a.n., cook, g.d. & williams, r.j., 2003, fire in tropical savannas: the kapalga experiment, springer verlag, new york.biggs, r., biggs, h.c., dunne, t.t., govender, n. & potgieter, a.l.f., 2003, ‘experimental burn plot trial in the kruger national park: history, experimental design and suggestions for data analysis’, koedoe 46(1), 1–15. bijker, h.j., sumner, p.d., meiklejohn, k.i. & bredenkamp, g.j., 2001, ‘documenting the effects of veld burning on soil and vegetation characteristics in giant’s castle game reserve, kwazulu-natal drakensberg’, south african geographical journal 83, 28–33. http://dx.doi.org/10.1080/03736245.2001.9713716 bond, w.j. & keeley, j.e., 2005, ‘fire as a global herbivore: the ecology and evolution of flammable ecosystems’, trends in ecology and evolution 20(7), 387–394. http://dx.doi.org/10.1016/j.tree.2005.04.025 brown, j.a.h., 1972, ‘hydrologic effects of a bushfire in a catchment in south-eastern new south wales’, journal of hydrology 15, 77–96. http://dx.doi.org/10.1016/0022-1694(72)90077-7 burns, c.e., zinn, a.d., govender, n., buis, g.m. & smith, m.d., in review, ‘the distribution of large herbivores in a fire-managed savanna grassland: assessing the role of bottom-up processes’, oikos. debano, l.f., 2000, ‘the role of fire and soil heating on water repellency in wildland environments: a review’, journal of hydrology 231, 195–206. http://dx.doi.org/10.1016/s0022-1694(00)00194-3 eijkelkamp agrisearch equipment, 2008, operating instructions: geulph permeameter, giesberg, the netherlands, viewed 18 december 2011, from http://pkd.eijkelkamp.com/portals/2/eijkelkamp/files/manuals/m1-0907e%20guelph%20permea.pdf eijkelkamp agrisearch equipment, 2009, operating instructions: pocket penetrometer, giesberg, the netherlands, viewed 18 december 2011, from http://pkd.eijkelkamp.com/portals/2/eijkelkamp/files/manuals/m1-0603e%20pocket%20penetromet.pdf eijkelkamp agrisearch equipment, 2011, operating instructions: pocket vane tester, giesberg, the netherlands, viewed 18 december 2011, from http://pkd.eijkelkamp.com/portals/2/eijkelkamp/files/manuals/m1-1410e%20pocket%20vane%20test.pdf ekwue, e.i., 1990, ‘organic-matter effects on soil strength properties’, soil and tillage research 16(3), 289–297. http://dx.doi.org/10.1016/0167-1987(90)90102-j fynn, r.w.s., haynes, r.j. & o’connor, t.g., 2003, ‘burning causes long-term changes in soil organic matter content of a south african grassland’, soil biology and biochemistry 35(5), 677–687. http://dx.doi.org/10.1016/s0038-0717(03)00054-3 gertenbach, w.p.d., 1983, ‘landscapes of the kruger national park’, koedoe 26, 9–121. herrick, j.e. & jones, t.l., 2002, ‘a dynamic cone penetrometer for measuring soil penetration resistance’, soil science society of america journal 66, 1320–1324. http://dx.doi.org/10.2136/sssaj2002.1320 higgins, s.i., bond, w.j. & trollope, w.s.w., 2000, ‘fire, resprouting and variability: a recipe for grass–tree coexistence in savanna’, journal of ecology 88, 213–229. http://dx.doi.org/10.1046/j.1365-2745.2000.00435.x higgins, s.i., bond, w.j., february, e.c., bronn, a., euston-brown, d.i.w., enslin, b. et al., 2007, ‘effects of four decades of fire manipulation on woody vegetation structure in savanna’, ecology 88(5), 1119–1125. http://dx.doi.org/10.1890/06-1664, pmid:17536398 holdo, r.m., mack, m.c. & arnold, s.g., 2011, ‘tree canopies explain fire effects on soil nitrogen, phosphurus and carbon in a savanna ecosystem’, journal of vegetation science 23(2), 352–360. http://dx.doi.org/10.1111/j.1654-1103.2011.01357.x jones, c.l., smithers, n.l., scholes, m.c. & scholes, r.j., 1990, ‘the effect of fire frequency on the organic components of a basaltic soil in the kruger national park’, south african journal of plant soil 7(4), 236–238. knapp, a.k., briggs, j.m., hartnett, d.c. & collins, s., 1998, grassland dynamics: long-term ecological research in tallgrass prairie, oxford university press, new york. lindley, a.j., bosch, j.m. & van wyk, d.b., 1988, ‘changes in water yield after fire in fynbos catchments’, water sa 14(1), 7–12. lorentz, s., thornton-dibb, s., pretorius, j.j. & goba, p., 2003, hydrological systems modelling research programme: hydrological processes. phase ii quantification of hillslope, riparian and wetland processes, report to the water research commission on the project a field study of two and three dimensional processes, wrc report k5/1061, water research commission, pretoria. mataix-solera, j., cerda, a., arcenegui, v., jordán, a. & zavala, l.m., 2011, ‘earth-science reviews fire effects on soil aggregation: a review’, earth science reviews 109(1–2), 44–60. http://dx.doi.org/10.1016/j.earscirev.2011.08.002 mills, a.j. & fey, m.v., 2004, ‘frequent fires intensify soil crusting: physicochemical feedback in the pedoderm of long-term burn experiments in south africa’, geoderma 121, 45–64. http://dx.doi.org/10.1016/j.geoderma.2003.10.004 moody, j.a., kinner, d.a. & úbeda, x., 2009, ‘linking hydraulic properties of fire-affected soils to infiltration and water repellency’, journal of hydrology 379(3–4), 291–303. http://dx.doi.org/10.1016/j.jhydrol.2009.10.015 pierson, f.b., williams, c.j., hardegree, s.p., weltz, m.a., stone, j.j. & clark, p.e., 2011, ‘fire, plant invasions, and erosion events on western rangelands’, ecology 64(5), 439–449. scholes, r.j. & hall, d.o., 1996, ‘the carbon budget of tropical savannas, woodlands and grasslands’, in a.i. breymeyer, d.o. hall, j.m. melillo & g.i. agren (eds.), global change: effects on coniferous forests and grasslands, pp. 69–100, wiley, chichester. scholes, r.j. & walker, b.h., 1993, an african savanna – synthesis of the nylsvley study, cambridge university press, cambridge. http://dx.doi.org/10.1017/cbo9780511565472 scott, d.h., 1997, ‘the contrasting effects of wildfire and clearfelling on the hydrology of a small catchment’, hydrological processes 11, 543–555. http://dx.doi.org/10.1002/(sici)1099-1085(199705)11:6<543::aid-hyp474>3.0.co;2-j smit, i.p.j., asner, g.p., govender, n., kennedy-bowdoin, t., knapp, d.e. & jacobson, j., 2010, ‘effects of fire on woody vegetation structure in african savanna’, ecological applications 20(7), 1865–1875. http://dx.doi.org/10.1890/09-0929.1 soil classification working group, 1991, soil classification: a taxonomic system for south africa. memoirs on the agricultural natural resources of south africa no. 15. soil and irrigation research institute, department of agricultural development, pretoria. statext version 1.4, 2011, computer software, viewed 06 december 2011, from http://www.statext.com townsend, s. & douglas, m., 2000, ‘the effect of three fire regimes on stream water quality, water yield and export coefficients in a tropical savanna (northern australia)’, journal of hydrology 229, 118–137. http://dx.doi.org/10.1016/s0022-1694(00)00165-7 van der schijff, h.p., 1958, inleidende verslag oor veldbrandnavorsing in die nasionale krugerwildtuin [preliminary report on wildfire research in the kruger national park]. koedoe 1, 60–93. venter, f.j. & govender, n., 2012, ‘a geomorphic and soil description of the long-term fire experiment in the kruger national park, south africa’, koedoe 54(1), art. #1037, 10 pages. http://dx.doi.org/10.4102/koedoe.v54i1.1037 walter, h., 1971, ecology of tropical and subtropical vegetation, 1st edn., oliver & boyd, edinburgh. webber, c.n., 1979, ‘the effects of fire on soil–plant ecological relationships in the southern part of the kruger national park: a study in soil geography’, msc thesis, dept. of geography, university of pretoria. article information author: adrian j.f.k. craig1 charlene bissett1,2 mark d. galpin2 bryan olver2,3 pat e. hulley1 affiliations: 1department of zoology and entomology, rhodes university, south africa 2kwandwe private game reserve, grahamstown, south africa 3phinda private game reserve, hluhluwe, south africa correspondence to: adrian craig email: a.craig@ru.ac.za postal address: department of zoology and entomology, rhodes university, grahamstown 6140, south africa dates: received: 22 jul. 2010 accepted: 03 dec. 2010 published: 02 mar. 2011 how to cite this article: craig, a.j.f.k., bissett, c., galpin, m.d., olver, b. & hulley, p.t., 2011, ‘the avifauna of kwandwe private game reserve, eastern cape, south africa’, koedoe 53(1), art. #1015, 5 pages. doi:10.4102/koedoe.v53i1.1015 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) the avifauna of kwandwe private game reserve, eastern cape, south africa in this original research... open access • abstract • introduction    • study area • methods • results and discussion    • past records    • current records    • single records    • species indicative of range expansion    • seasonal visitors    • re-introductions    • conservation significance • conclusion • acknowledgements • references abstract (back to top) a protected area since 1999, kwandwe private game reserve incorporates several former farms, for which past records of bird occurrences are available. no bird species appear to have been lost from the area. between 2001 and 2005, a group of observers conducted systematic bird surveys in most months, which allowed the status (resident, migrant or irregular visitor) of most bird species to be determined. at least three species have established breeding populations in the reserve over the past 10 years. of 302 species reliably recorded to date, 182 (60.3%) appear to be resident, 46 (15.2%) are seasonal migrants and 74 (24.5%) are vagrant visitors. eight vulnerable and eight near-threatened bird species have been recorded; blue crane, kori bustard and african crowned eagle have bred in the reserve. together with other state-owned and private protected areas in this region, this reserve holds a significant portion of the inland bird species recorded from the eastern cape. conservation implications: the varied thicket vegetation types of the great fish river valley support a considerable diversity of bird species. if these habitats are managed for biodiversity conservation, they can support a large component of the terrestrial avifauna of the eastern cape region. introduction (back to top) over the past 20 years there has been a significant shift in land usage in the eastern cape. several commercial farms have been converted to game ranches either for hunting or for ecotourism (palmer 2008), leading to the formation of extensive private reserves along with the expansion of state-owned conservation areas such as the addo elephant national park. this has particularly affected the vegetation type originally described as valley bushveld by acocks (1975), and now treated as a distinct biome, albany thicket, by mucina and rutherford (2006). in this paper, we present some baseline data to establish how this management change has influenced the avifauna in one protected area in the eastern cape. kwandwe private game reserve (pgr) was established in 1999 and the first lodges opened for guests in 2001. there are bird records from different sources for three distinct periods: museum specimens (1916–1960), bird club observations (1980–1999), and surveys by reserve staff and associates (2001–2005). the present study investigated four questions. 1. what bird species occurred in the area prior to the establishment of the reserve? 2. have any species been lost since then and, if so, can we explain their disappearance? 3. have new species become established and, if so, can we account for their arrival? 4. do periodic vehicle-based surveys reflect the current species diversity? figure 1:vegetation map of kwandwe private game reserve (from bissett 2007). study area kwandwe pgr (33°09’s, 26°37’e ) lies in the great fish river valley, approximately 35 km north of grahamstown in the eastern cape, south africa. the farms purchased for the reserve had previously been used primarily for stock farming, with irrigated lands bordering on the great fish river and ostrich pens on some sections. in 2005 additional land was purchased on the northern bank of the great fish river, increasing the area of the reserve to some 20 000 ha. the perennial great fish river flows through the reserve for about 25 km and all watercourses drain towards the river (figure 1).the main vegetation types represented in the reserve are great fish noorsveld and great fish thicket, within the albany thicket biome. the kwandwe pgr preserves a significant area of great fish noorsveld (mucina & rutherford 2006). at a finer scale, bissett (2007) described 11 vegetation types in the reserve, of which five cover more than 1000 ha. these form a mosaic of thicket and bush-clump types, with acacia karoo, cussonia spicata, euclea undulata, euphorbia spp., maytenus spp., pappea capensis, portulacaria afra, searsia spp. and schotia afra as the dominant larger plant species. there are also old cultivated lands, now covered by grasses, and two large dams. recent additions to the property include some valleys with dry forest, including species typical of both valley thicket and afro-montane forest (mucina & rutherford 2006), on the north bank of the great fish river, where large trees such as harpephyllum caffrum occur. the altitude ranges from 170 m a.s.l in the great fish river valley to 600 m a.s.l. on the ridges, and mean annual rainfall for the past 7 years was 394 mm and 358 mm at brandeston and krantzdrift, respectively (bissett 2007). table 1: summary of the bird groups recorded on kwandwe private game reserve since october 2001. methods (back to top) past records examined in this study included the published literature on birds of the eastern cape, the bird collection at the albany museum, records of outings by the diaz cross bird club, and an unpublished list of birds on the farm brandeston, compiled by c.a. whittington-jones. during 2001 the staff at kwandwe pgr began a systematic survey conducted on a specific day each month for the ’birds in reserves project’ (birp), coordinated by the (then) avian demography unit at the university of cape town. birp records exist for five months in 2001, for every month in 2002, for eight months in both 2003 and 2004, and for all months in 2005. these records allowed for an assessment of the birds that are likely to be resident as opposed to vagrants. we have also included more recent records (up to october 2009). during the birp surveys we followed a standard route, with surveying starting at 06:00 in summer and at 07:00 in winter. the route was designed to traverse all the different vegetation types in the reserve and to pass close by the major water bodies. the total distance was about 35 km, which was covered in 8–10 hours. we also included other sightings by rangers during the observation period; these reports typically included only large birds such as bustards and cranes, or birds of prey. between three and eight observers participated in the surveys, depending on staff commitments on the particular day. normally an open game-viewing vehicle was used; if another vehicle was used, there was always one observer standing on the back. all observers were experienced bird watchers equipped with binoculars and all species confirmed by call were included. the bird names used follow those in roberts birds of southern africa (eds. hockey, dean & ryan 2005). only standard english names are used in the text, but both english and scientific names are given in online appendix 1. results and discussion (back to top) past records specimens from the albany museum included 68 study skins from the present reserve area (collected on the farms brandeston and the fort, which both had river frontage). all but three specimens were collected between 1920 and 1927: a long-billed crombec collected on brandeston in 1916, a dark-capped bulbul from the fort collected in 1939, and a crowned hornbill from brandeston collected in 1960. the collections were made by v.e. watson on the fort and a.t. rivett-carnac on brandeston. hewitt (1931) cited rivett-carnac as one of his bird informants and mentioned a single sighting of a white-fronted bee-eater from the great fish river valley, but gave no precise locality. despite this being a biased sample in which larger birds such as waterbirds, gamebirds and raptors are poorly represented, it does provide insight into the avifauna history of the area. of the 56 species, 55 have been recorded on the reserve in the past 5 years. the only exception is the red-headed finch, which was categorised as ‘locally nomadic’ by dean (1997) and makes occasional incursions into this region according to recent records from brandeston. seven sets of eggs collected at heatherton towers (now the visitor reception centre on kwandwe pgr) in the 1930s, also formed part of the past records. the eggs represent species that currently breed on kwandwe pgr. they are listed together with the study skins in the category ’museum’ in online appendix 1. c.j. skead visited all the farms that have now been incorporated into kwandwe pgr during his research on eastern cape birds, but no specific records are mentioned in his reviews (skead 1965, 1967). the diaz cross bird club visited the farms brandeston, krantzdrift and the fort (now incorporated into kwandwe pgr) on four occasions between 1980 and 1998, during which 117 bird species were recorded within the area of the present reserve. three of these species (glossy ibis, southern pochard and cape canary) have not yet been recorded since the establishment of the reserve. club members also submitted sightings of interest to the club newsletter and we have extracted from the ‘diaz diary’ all such reports (listed under ’diaz’ in online appendix 1). one species, the european bee-eater, was reported in march 1988 but has not been recorded from the area since. there are unconfirmed records for denham’s bustard (formerly often confused with ludwig’s bustard – see herholdt 1988), sand martin (likely erroroneous identification for the brown-throated martin, formerly the african sand martin), yellow wagtail, and thick-billed weaver (the latter both from an unreliable observer). from july 1997 to june 1999, c.a. whittington-jones made regular visits to brandeston during his study of red-billed queleas at ostrich feedlots. he recorded 136 bird species on the farm, of which only the red-headed finch (sighted in may 1998 and may 1999) has not subsequently been recorded on the reserve. since the removal of the ostrich feedlots, the red-billed quelea has become an occasional vagrant in the area. table 2: bird species recorded at kwandwe private game reserve, listed in the current red data book for south africa (ed. barnes 2000). current records to date, 302 bird species have been recorded reliably on kwandwe pgr (online appendix 1). of this total, 274 species were recorded during the 45 birp surveys included in this overview, representing more than 90% of the species recorded up to 2006. the avifauna was subdivided into broad habitat classes within which the birds could be categorised as regulars, vagrants or migrants based on the monthly distribution of records (table 1).compared with three other eastern cape conservation areas where thicket is the predominant vegetation type, kwandwe pgr has the most diverse avifauna (302 species), of which 11% have been recorded only on this reserve. some 194 species have been recorded from the original addo elephant national park area of 12 000 ha (urquhart & klages 1997), excluding the recently added colchester, zuurberg and the greater addo sections towards the coast. these include habitat types such as southern coastal forest, suurberg quartzite and suurberg shale fynbos (mucina & rutherford 2006), which are absent from the other reserves. the checklist for the great fish river reserve complex records 245 species from the area west of the great fish river, which spans the former andries vosloo kudu reserve and the sam knott nature reserve (23 000 ha combined). shamwari game reserve currently covers 22 000 ha, with 257 species having been recorded. although addo elephant national park is thus the smallest of these conservation areas and no river runs through the reserve, all four reserves lie below the inland escarpment and at a similar distance from the coast. the great fish river reserve is almost adjacent to kwandwe pgr and shamwari game reserve is less than 20 km from addo elephant national park in direct line of flight. these reserves form part of a belt of conservation areas between the coast and the more arid interior and include a significant proportion of the south african avifauna (combined total = 334 species), despite none being mentioned in the first national survey of important bird areas (ed. barnes 1998). single records several species have been recorded only once. however, in all cases there are other records for single sightings in adjoining regions of the eastern cape and in some instances the timing of vagrant sightings suggests that the same individual birds could have been observed. this group includes the african pygmy goose, ground woodpecker, african black oystercatcher (sighted after a period of onshore gales along the coast), caspian tern, tawny eagle, bateleur, baillon’s crake, greater flamingo, abdim’s stork, dusky lark, chat flycatcher, chorister robin-chat, capped wheatear, white-browed sparrow-weaver and long-tailed paradise whydah. species indicative of range expansion the goliath heron, a rarity in the eastern cape in the 1980s, is now well established along coastal rivers, down the east coast (craig 2006) and on inland dams, perhaps spreading from major impoundments along the orange river.kurrichane buttonquails were not reported from the eastern cape during the bird atlas survey (harrison et al. 1997), although there are historic records (skead 1967). several recent records at different times of the year have led to suggestions of a range expansion into the eastern cape (eds. hockey et al. 2005). birds on kwandwe pgr were most often flushed from areas of thicket–grass patch habitat mosaic. in 2001 an adult bird was seen with a single, very small chick, indicating that breeding had occurred. the white-fronted bee-eater was previously regarded as a vagrant in the eastern cape (harrison et al. 1997; skead 1967). a pair was found breeding on the sundays river at colchester in 1998 (martin 1999) and since then birds have been seen at this site annually. found nesting on kwandwe pgr in january 2002, the birds were initially thought to be summer-breeding visitors, but in recent years they have been sighted in most months and may now be considered resident. african palm swifts were not recorded south of kwazulu-natal by skead (1967) nor quickelberge (1989), who reviewed data for the transkei region. nevertheless, the first records for the eastern cape date back to the 1980s in steytlerville (ward et al. 1984). this species is now regular along the coast where suitable palms are available and has also been recorded in grahamstown. a small colony has been present on kwandwe pgr at the brandeston homestead since 2005. the double-banded courser was not found in the grahamstown region during the bird atlas surveys (harrison et al. 1997), which supports vernon’s (1982) observation that coursers are rarely sighted in the eastern cape today. however, this courser has been sighted in one section of the kwandwe reserve quite regularly, in association with other birds typical of the arid interior (e.g. the rufous-eared warbler). breeding had been recorded in 2001 and 2002. the violet-backed starling is an intra-african migrant, which has been recorded breeding in the eastern cape to the east of the great fish river. in recent summers there have been an increasing number of reports from this southern extremity of its range in south africa. the scaly-feathered finch was first reported in 2006 and is now apparently a breeding resident, but its distribution is very localised within the reserve. to date there has been a single sight record of the white-browed sparrow-weaver, but nesting birds were found on a farm south-west of kwandwe pgr recently. this species has become established on farms in the queenstown district in the past 20 years and is currently expanding its range there (k. webster, pers. comm., may 2010). it colonised the mountain zebra national park from the late 1960s onwards (craig, hulley & parker 2005; skead 1966). passing largely unnoticed, the southern grey-headed sparrow underwent a major range expansion during the last century, colonising the albany district during the 1960s (craig, every & summers-smith 1987) and reaching the south-western cape by the 1990s (harrison et al. 1997). seasonal visitors barratt’s warbler, a bird from the mountain forests of the eastern cape often moves to lower altitudes in winter (vernon 1989). this is reflected by higher winter records on the coast and occasional sightings en route (craig 1986). the warbler was present in thick cover close to a game-viewing hide for several weeks in june 2003; it can easily be overlooked in areas traversed only by vehicles. a winter record of chorister robin-chat may also represent an individual moving from montane forest towards the coast.the dusky sunbird is evidently a winter visitor to the great fish river valley, with several sightings on the reserve where it often feeds on the honey thorn (lycium oxycarpum). there were also ringing records from brandeston and resolution farms in the fort brown area prior to the establishment of the reserve and later captures at aloes on kwandwe pgr and hounslow, a farm on the western border of the reserve. re-introductions along with other reserves, kwandwe pgr has been stocked with fauna which formerly occurred in the area (skead 1987) and, more controversially, with species such as giraffe (giraffa camelopardalis), which can be considered alien to the eastern cape (castley, boshoff & kerley 2001). by contrast, only two bird species have been considered for re-introduction.the red-billed oxpecker, which had been locally extinct, was re-introduced to the eastern cape from the kruger national park in 1990, with birds released in addo elephant national park, the great fish river reserve and on a private game farm in the highlands area (craig & weaver 1990). birds have subsequently also been introduced to shamwari game reserve in 1998 and there was a second release in the great fish river reserve in 2003. breeding has been reported from addo elephant national park, the great fish river reserve and shamwari game reserve, with regular sightings from properties adjoining these sites. the birds are often seen on giraffe, one of their favourite hosts throughout southern africa. currently oxpeckers are recorded in kwandwe pgr most months and over the past 2 years there have been sightings of dark-billed juvenile birds, despite no direct evidence of breeding within the reserve. the southern ground hornbill may be introduced to kwandwe in the future if the pilot project at shamwari game reserve proves successful, although this region clearly represents the edge of the natural range of this species (hulley & craig 2007). conservation significance the avifauna of kwandwe pgr includes 16 species of conservation concern, all of which are potential breeding species in the area (table 2). the cape vulture has suffered a major decline throughout the eastern cape, with many former breeding sites now abandoned (boshoff, piper & michael 2009). only occasional vagrants are now reported in this part of the province. similarly, the tawny eagle was formerly widespread in the eastern cape and until 1981 a regular breeding species. it is suspected to have suffered heavily from indiscriminate use of poisoned baits (ed. barnes 2000). the sighting in january 2005 was the first in the grahamstown district for some 15 years (craig 2005). bateleurs have seldom been seen south of the orange river over the past 50 years and a record of an adult bird at kwandwe pgr in june 2009 was a major surprise. african crowned eagles nest on the reserve and martial eagles still nest in the district. blue cranes nest on the reserve annually, while kori bustards have nested here occasionally. these ground-nesting species are, however, vulnerable to predators, particularly the black-backed jackal (canis mesomelas). cranes and bustards frequently collide with power lines (jenkins, smallie & diamond 2010) and it is thus laudable that all power cables on this reserve are now underground. the great fish river valley appears to form a conduit both for southward penetration of species typical of the arid interior, such as the red-billed firefinch, and for inland movements of species generally restricted to the coastal belt, such as the yellow weaver (recorded in the great fish river reserve, but not yet on kwandwe pgr). the thickets also provide acceptable habitat for some forest species such as the narina trogon and the dark-backed weaver. conservation of the thicket biome is currently considered a priority, since this habitat has been extensively modified by agriculture and is regarded as important in carbon sequestration. thus, the mosaic of privately managed and state-owned reserves in this region of the eastern cape should be seen as a significant contributor to conserving south african biodiversity. conclusion (back to top) although we lack a comprehensive checklist for the region prior to the establishment of the reserve, there is no clear evidence that formerly resident bird species have been lost, while at least three species (white-fronted bee-eater, african palm swift and scaly-feathered finch) have established breeding populations since the area had become fully protected. regular vehicle-based surveys by experienced observers seem to record a high proportion of the bird species that occur within the reserve. such monitoring should form a part of future management plans, in the interest of both conserving biodiversity and highlighting additional attractions for visiting tourists. acknowledgements (back to top) we are most grateful to the management of kwandwe pgr, in particular angus sholto-douglas, for supporting this project and placing vehicles at our disposal. our thanks also go to all the rangers, trainees, and other volunteers who have taken part in the surveys. brad fike (great fish river reserve) and antony collett (shamwari game reserve) kindly provided copies of unpublished checklists of the birds of these reserves, and craig whittington-jones offered us all his dated records for brandeston. a.c. and p.h. received research funding from rhodes university. c.b. received funding for research on carnivores from kwandwe pgr during the time of the study. references (back to top) acocks, j.p.h., 1975, ‘veld types of south africa’, 2nd edn., memoirs of the botanical survey of south africa 40, 1−128. barnes, k.n. (ed.), 1998, the important bird areas of southern africa, birdlife south africa, johannesburg. barnes, k.n. (ed.), 2000, the eskom red data book of birds of south africa, lesotho and swaziland, birdlife south africa, johannesburg. bissett, c., 2007, ‘the feeding and spatial ecologies of the large carnivore guild on kwandwe private game reserve’, phd thesis, department of zoology and entomology, rhodes university. boshoff, a., piper, s. & michael, m., 2009, ‘on the distribution and breeding status of the cape griffon gyps coprotheres in the eastern cape province, south africa’, ostrich 80(2), 85−92. doi:10.2989/ostrich.2009.80.2.4.831 castley, j.g., boshoff, a.f., kerley, g.i.h., 2001, ‘compromising south africa’s natural biodiversity: inappropriate herbivore introductions’, south african journal of science 97, 344−348. craig, a., 1986, ‘forest birds on rhodes university campus’, diaz diary 155, 4−5. craig, a., 2005, ‘will the tawny eagle return to the eastern cape?’, diaz diary 33(1), 11−12. craig, a., 2006, ‘goliath herons on the eastern cape coast’, bee-eater 57(2), 36−37. craig, a., every, b. & summers-smith, d., 1987, ‘the spread of the southern greyheaded sparrow in the cape province’, annals of the cape provincial museums (natural history) 16(9), 191−200. craig, a.j.f.k., hulley, p.e. & parker, d., 2005, ‘a re-assessment of the avifauna of the mountain zebra national park’, koedoe 48, 95−113. craig, a. & weaver, a., 1990, ‘the relocation of the redbilled oxpecker’, bee-eater 41(4), 58−61. dean, w.r.j., 1997, ‘the distribution and biology of nomadic birds in the karoo, south africa’, journal of biogeography 24, 769−779. doi:10.1046/j.1365-2699.1997.00163.x harrison, j.a., allan, d.g., underhill, l.g., herremans, m., tree, a.j., parker, v., et al., 1997, the atlas of southern african birds, birdlife south africa, johannesburg. herholdt, j.j., 1988, ‘the distribution of stanley’s and ludwig’s bustards in southern africa: a review’, ostrich 59, 8−13. doi:10.1080/00306525.1988.9633917 hewitt, j., 1931, a guide to the vertebrate fauna of the eastern cape province. part 1. mammals and birds, albany museum, grahamstown. hockey, p.a.r., dean, w.r.j. & ryan, p.g. (eds.), 2005, roberts birds of southern africa, 7th edn., john voelcker bird book fund, cape town. hulley, p.e.h. & craig, a.j.f.k., 2007, ‘the status of the southern ground-hornbill in the grahamstown region, eastern cape’, ostrich 78, 89−92. doi:10.2989/ostrich.2007.78.1.13.57 jenkins, a.r., smallie, j.j. & diamond, m., 2010, ‘avian collisions with power lines: a global review of causes and mitigation with a south african perspective’, bird conservation international 20(3), 152−278. doi:10.1017/s0959270910000122 martin, p., 1999, ‘first breeding record of white-fronted bee-eaters’, bee-eater 50(2), 13. mucina, l. & rutherford, m., 2006, ‘vegetation of south africa, lesotho and swaziland’, strelitizia 19, 1−807. palmer, a., 2008, ‘a study of land-use trends and land tenure changes in the makana local municipality (albany registration district) for the period 1994 to 2008’, unpublished report to ardri, university of fort hare, east london. quickelberge, c.d., 1989, the birds of the transkei, durban natural history museum, durban. skead, c.j., 1965, ‘birds of the albany (grahamstown) district’, south african avifauna series 30, 1−46. skead, c.j., 1966, ‘white-browed sparrow-weaver, plocepasser mahali a. smith, in the eastern and north-eastern cape’, ostrich 37, 128−129. skead, c.j., 1967, ‘ecology of birds in the eastern cape province’, ostrich suppl. 7, 1−103. skead, c.j., 1987, historical mammal incidence in the cape province. vol. 2. the eastern half of the cape province, including the ciskei, transkei and east griqualand, chief directorate, nature and environmental conservation, cape town. urquhart, c. & klages, n., 1997, addo, more than just elephants, bluecliff publishing, port elizabeth. vernon, c.j., 1982, ‘coursers in the eastern cape’, bee-eater 33(3), 31−34. vernon, c.j., 1989, ‘observations on the forest birds around east london’, ostrich 14, suppl., 14, 75−84. ward, d., martin, r., martin, j. & martin, e., 1984, ‘palm swifts in the cape province’, bokmakierie 36, 107. article information authors: angela gaylard1 sam ferreira2 affiliations: 1savanna research unit, conservation services, south african national parks, garden route regional office, knysna, south africa 2savanna research unit, conservation services, south african national parks, skukuza, south africa correspondence to: angela gaylard email: angela.gaylard@sanparks.org postal address: po box 3542, knysna 6570, south africa dates: received: 24 may 2010 accepted: 17 mar. 2011 published: 13 may 2011 how to cite this article: gaylard, a. & ferreira, s., 2011, ‘advances and challenges in the implementation of strategic adaptive management beyond the kruger national park – making linkages between science and biodiversity management’, koedoe 53(2), art. #1005, 8 pages. doi:10.4102/koedoe.v53i2.1005 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) advances and challenges in the implementation of strategic adaptive management beyond the kruger national park – making linkages between science and biodiversity management in this essay... open access • abstract • introduction • bridging the science–management gap    • linking scientists and managers by creating a forum for developing a shared rationale    • revisiting the objectives hierarchy to ensure that it is linked to management concerns    • linking the objectives hierarchy with tpcs, monitoring and research    • case studies: unpacking concerns to identify the linkages between the objectives hierarchy and tpcs • evolution of the tpc concept    • use of the tpc concept by other organisations • challenges, uncertainties and the way forward    • challenges involved in adopting the new nonequilibrium paradigm    • institutional constraints to implementing sam    • making sam tangible to managers yet rigorous enough for scientists – the search for a ‘requisite simplicity’ • conclusion • acknowledgments • references abstract (back to top) south african national parks (sanparks) makes use of strategic adaptive management (sam) to achieve its primary mandate of biodiversity conservation. this involves an iterative adaptive planning, management and review cycle to ensure appropriate alignment of stakeholder values with conservation objectives, to address the uncertainty inherent in complex social–economic–ecological systems and to learn explicitly whilst doing so. adaptive management is recognised as the most logical framework for continuous improvement in natural resource management; nevertheless, several challenges in its implementation remain. this paper outlined these challenges and the various modifications to sanparks’ adaptive planning and management process that have emerged during its development. we demonstrated how the establishment of a regular science–management forum provides opportunities for social co-learning amongst resource managers and scientists of a particular park, whilst providing other positive spin-offs that mature the sam process across the organisation. we discussed the use of particular conceptual constructs that clarify the link between monitoring, management requirements and operational endpoints, providing the context within which thresholds of potential concern (tpcs) should be set, prioritised and measured. the evolution of the tpc concept was also discussed in the context of its use by other organisations, whilst recognising its current limitations within sanparks. finally, we discussed remaining implementation challenges and uncertainties, and suggested a way forward for sam. conservation implications: this paper outlined practical methods of implementing sam in conservation areas, beyond what has already been learnt within, and documented for, the kruger national park. it also highlighted several implementation challenges that prove useful to other conservation agencies planning to adopt this approach to managing complex ecosystems. introduction (back to top) south african national parks (sanparks) makes use of an adaptive planning process to ensure that the principles of strategic adaptive management (sam) and the managing of its parks conform to legislation as well as to its own mandates, values and organisational structure (rogers & sherwill 2008). the planning process is adaptive because it cycles sequentially, but often iteratively, through its major development phases. sanparks’ adaptive planning process originated along with the development of sam in the kruger national park (knp) in the late 1990s (biggs 2003). attempts to implement the approach in the other national parks have met with various extents of success since the mid-2000s (see biggs et al. 2011; ferreira et al. 2010). these attempts followed the mckinsey report (mckinsey & company 2002), which highlighted the lack of a formal biodiversity policy or management plans for most of the parks under sanparks’ management authority (rogers & sherwill 2008). such policies are now a requirement under new national legislation for protected areas and the conservation of biodiversity. implementing this legislative requirement carries significant challenges for a variety of reasons. included in these are the uncertainty, expectations and preferences inherent in using adaptive management (cundill & fabricius 2009), and, in particular, thresholds of potential concern (tpcs), when managing complex social–economic–ecological systems (biggs et al. 2011; jacobson et al. 2006; rogers, roux & biggs 2000; venter et al. 2008). disjuncts between the understanding of scientists and conservation practitioners (specifically rangers and park managers, henceforth referred to as managers) in terms of how systems may function often leads to inertia in management decisions (duncan & wintle 2008; rogers 2003) and hence a great deal of frustration for conservation agencies trying to deliver their mandate. in this paper, we address this challenge and illustrate how regular science–management interactions can generate positive spinoffs that mature sam across the organisation. this includes the explicit use of particular conceptual constructs and supplementary tools that enable the development of joint mental models between scientists and managers. we will also demonstrate how the latter, in turn, has clarified the links between monitoring, management requirements and operational endpoints, and thus provide the context within which tpcs should be set (see biggs et al. 2011), prioritised and measured. finally, we reflect on some of the implementation challenges faced by sanparks and consider the areas that still require development to take sam forward. bridging the science–management gap (back to top) linking scientists and managers by creating a forum for developing a shared rationale implementing sam in any south african national park under sanparks’ management authority begins with an adaptive planning process involving facilitated stakeholder meetings, in which a vision and mission, as well as high level objectives are jointly agreed upon (rogers & sherwill 2008). thereafter, this broad values-based vision is translated into a fully fledged objectives hierarchy that documents the sequential reasoning of these values into science-based system outcomes (rogers & sherwill 2008). decision-making towards achieving these outcomes is based on the setting and subsequent monitoring of the tpcs that define the acceptable limits of change in ecosystem or biodiversity composition, structure and function (biggs 2003). finally, it is essential that there is reflection at each step, in order to ensure learning (cundill & fabricius 2009). whilst sanparks’ application of sam originated in the knp, several modifications have emerged as a consequence of its implementation in other national parks. an early and far-reaching example of such a modification was the establishment of a science–management forum (smf) for each park. smfs are regular meetings of groups of sanparks scientists and managers, with the primary aim being to provide a platform for meaningful interaction regarding conservation issues. the adaptive planning process requires technical input once the high level objectives have been compiled along with stakeholders. the smfs therefore present an opportunity for scientists and managers to provide this technical input, working their way through the various stages of their park’s adaptive management cycle after the preliminary stakeholder meetings. consequently, smfs also provide opportunities for essential discourse amongst scientists and managers about the practical implications of sanparks’ recent paradigm shift towards a complex systems approach to conservation (rogers 2003). nevertheless, this paradigm shift has been only a recent occurrence and many managers are still reluctant to relinquish the agriculturally based conservation practices of their training and accept the uncertainty inherent in managing the complex ecosystems expounded by scientists (rogers et al. 2000). yet managers have a wealth of valuable ‘on the ground’ knowledge and practical experience of feasible management actions. a critical function of smfs is therefore to establish regular interactions between scientists and managers, in which they can share their perspectives and develop a shared rationale that can be used to meaningfully proceed with the adaptive management process. it is during this process with the smfs that several additions or modifications have developed that have proved very constructive to the adaptive planning and management process. we outline these below and summarise their benefits for sam. revisiting the objectives hierarchy to ensure that it is linked to management concerns once the high level objectives have been set with the involvement of stakeholders, the technical details for unpacking each objective to create an objectives hierarchy is left to the conservation practitioners, that is park managers and scientists. lower levels of management are not always required to be present at the stakeholder meetings where the vision, mission, vital attributes and high level objectives are derived. however, it is essential that all members of the smf have had an opportunity to articulate their values, because these components are the direct translations of stakeholder values into the top tiers of the objectives hierarchy. agreement at this level of the adaptive planning process prevents conflicts lower down the objectives hierarchy at the levels where these managers have to operate. if the managers have not been involved in the stakeholder meetings, it is particularly important that they are provided with an opportunity to list all of their management concerns. this contributes significantly towards buy-in by these managers, whilst also ensuring that the scientists become aware of the particular suite of issues faced by these managers on the ground. similar concerns are then grouped into what have become known as ‘concern themes’ and cross-checked against the objectives hierarchy. apart from ensuring that no important issues are omitted from the objectives hierarchy, this exercise is particularly useful for scientists and managers to gauge the alignment of their own concerns (which, to a large extent, represent their values) against one another, and against the values of the park’s stakeholders. once the concern themes have been listed, the managers are asked to prioritise them according to the urgency with which they feel each concern needs to be addressed. concern themes most often receiving high priority involve the adverse effects of herbivory, predation and degradation such as erosion and alien plant infestation. importantly, these concerns are articulated in terms of ecosystem processes, in line with sanparks’ paradigm shift away from its previous species focus. linking the objectives hierarchy with tpcs, monitoring and research following a complex systems approach requires the recognition that ecosystems need to fluctuate in order to maintain their resilience (jacobson et al. 2006; pickett, kolasa & jones 2007). sam makes use of upper and lower tpcs to quantify how much to allow a system to fluctuate before managers should become concerned that it is about to change into an undesirable state (biggs 2003). once concern themes have been identified and prioritised, the next step is to identify tpcs for each. several factors challenged the derivation of these endpoints at the non-knp smfs. firstly, there were fewer (internal and external) scientists present with knowledge of the particular ecological system in smaller parks. secondly, fewer staff members present were familiar with the process of adaptive planning and management. thirdly, there was less familiarity with the concepts of flux and resilience and, lastly, the managers present largely still focused on managing species rather than ecosystem processes. consequently, what may have been intuitively obvious to those well-versed in the decade or so of tpc development in the knp (i.e. the linkage between lower level objectives and operational endpoints and tpcs) was often conspicuously absent in the smfs held in other parks. in the absence of such a linkage, the indicators, endpoints, and hence tpcs, are either difficult to identify or become disjointed from the mechanisms underlying the ecological processes involved. a key modification of the sam process has developed through its implementation beyond the knp to overcome this disjunct between higher and lower level planning. this modification involves unpacking the particular concerns related to each of the lower level objectives, identifying the ecological processes underlying these concerns and then exploring the mechanism and its drivers and modulators jointly with scientists and managers during the smfs. this is achieved explicitly by making use of a framework to extract a joint science–management understanding of these mechanisms (see lindenmayer & likens 2009). the use of a framework cooperatively exposes the mental models of scientists and managers, capitalising on the diverse experience and knowledge of both groups. in addition, in the true spirit of adaptive management, the framework acts as a knowledge management system, to which improvements to conservationists’ existing understanding can be added over time, ensuring explicit learning (gaylard, cadenasso & pickett in press). a multitude of frameworks exist for compiling conceptual models of ecological processes (cundill & fabricius 2009). the ultimate premise of sanparks’ objectives hierarchies is the maintenance of a ‘desired state’ (or set of conditions allowing a certain amount of flux), which meant that our need for a framework resonated strongly with the framework developed by pickett, cadenasso and benning (2003). their framework explores the creation of, and response to, heterogeneity, with its basic building blocks representing a change from one (desirable) condition to another (undesirable condition; see figure 1). figure 1:pickett et al.’s (2003) framework for understanding the generation of and responses to heterogeneity, used during the process of unpacking concerns in science–management forum meetings. this naturally poses the question of what brings about the change (i.e. the agent or driver), from which point any other factors that members of the smf understand or hypothesise to influence the state change (through their action on the driver, i.e. modulators) can be incorporated into the conceptual framework. once the conceptual model is complete, at least according to the joint understanding of the smf, the drivers and modulators are easily identifiable from the resultant diagram of the mechanism. the drivers (ideally) or modulators are the practical translation of the management objective into a measurable tpc because, through the use of the framework, they provide the link between the objective and the required tpc. the members of the smf quantify the tpc using their own knowledge and expertise, and within the context of the values articulated through the objectives hierarchy. scientists (within and external to sanparks) are then consulted to comment on and/or fine-tune these quantifications. it is important that the facilitator incorporates the aspect of scale into the discussion of the mechanism in question, because focusing on just one scale might obscure important controlling processes at other scales (cundhill & fabricius 2009). moreover, identification of the scale at which, ultimately, the tpc’s indicator will be measured is that at which the mechanism operates. in addition, in order to quantify the upper and lower levels of the tpc between which the particular component of the system will be allowed to flux, the smf assesses the risk to the system should the tpc be exceeded. a very useful construct for balancing ‘risk tolerant’ versus ‘risk averse’ notions is use of the concept of ‘reversibility’ (i.e. how reversible is the process?). processes that are irreversible or would take much longer to recover tend to have risk averse tpcs (i.e. they are not allowed to flux very widely), whereas tpcs for processes that can easily be reversed (e.g. supplementing game after predation has reduced prey biomass) may be allowed to have wider limits of tolerance. risk assessments have also been used by sanparks to prioritise objectives (see box 1) and, more recently, to evaluate the trade-offs between various management options to achieve a desired objective. box 1: comparative risk assessment as an effective nuanced means of prioritising objectives. if there is consensus that there are no practical issues with the monitoring, the measurement of the quantified tpc is incorporated into the monitoring programme, which is developed in parallel to this process. this ensures that all monitoring is linked explicitly to tpcs at each level of the objectives hierarchy (see ferreira et al. 2011). the only exception to this is what sanparks scientists refer to as ‘background’ data, which is data used to interpret models of how the system works. these data can be of two distinct types, (1) factors that we have no control over or cannot do anything about (e.g. climate) and (2) factors that we do have control over but, in themselves, are not necessarily drivers (e.g. park-wide animal numbers when they are not high enough to act as modulators). monitoring data should be constantly checked against the tpcs and tabled at smfs if they are exceeded, or predicted to be exceeded. a tpc maintenance system (biggs 2003) then guides the smf through the process of acting on the exceeded tpc. unpacking a concern in this way is a key modification of the sam process, providing four critical benefits aside from the common understanding to which we have already referred. firstly, it provides clear guidance in terms of what to manage in order to address the park’s objectives. it thus has direct management implications and provides transparent and mutually agreed upon directions for action. in fact, it is sensible to set such ‘decision’ tpcs (see martin et al. 2009) based on the modulators of the mechanisms involved. secondly, this will remove some of the unpredictability and, most importantly, the subjective setting of a tpc solely based on outcome measures such as number of species (see biggs et al. 2011, for the prediction value judgements that need to be made). thirdly, the unpacking of a concern provides clear directions about what to measure, as well as the justification for this action. unpacking of concerns inadvertently imposes prioritisation, because, often, the unpacking exercise identifies perceived concerns for which sometimes conservationists simply cannot do anything about at the park level. climate change is a good example of this – although monitoring may confirm changing climate, there is little that park managers themselves can do to reverse the trend and hence this concern is ranked lower. an additional fundamental spinoff of using a framework in this way is that the process forces managers and scientists to expose their various mental models to one another. the joint mental model that emerges therefore recognises the managers’ instinctual knowledge, as well as the scientists’ empirical or theoretical knowledge, combining them to create a rigorous, yet practical means of dealing with real management concerns. moreover, the compilation of such conceptual models exposes knowledge gaps, information which can then be used to direct research and monitoring. case studies: unpacking concerns to identify the linkages between the objectives hierarchy and tpcs although to date, the process of unpacking concerns has been confined largely to biodiversity issues, table 1 illustrates attempts to apply it within the areas of sanparks’ other core businesses: stakeholders and tourism. as such, we provide three examples of the outcomes of unpacking a concern using the process discussed in the previous section (table 1). table 1:three case studies of different types of concerns unpacked in three different national parks in south africa. the first example relates to a typical concern regarding a biodiversity objective, the key mandate of sanparks. the concern is over the loss of particular plant species in the addo elephant national park as a result of elephant impacts. in this example, elephants are the driver of loss of plant species, but unpacking the mechanism reveals that water provision homogenises the elephants’ use of space, causing the animals to use landscapes more intensely. as such, individual plants are subjected to more regular foraging, with less time to recover before being visited by elephants again. the extent or rate of this homogenisation is modulated by elephant density, becoming more extensive or rapid with higher elephant densities, resulting in a high risk to the ecosystem because recovery is slow. hence, the measurement indicator is identified as the elephants’ use of space, and how this is related to their density. this case study illustrates how unpacking the concern facilitates the definition of how the particular problem is created. in this case, dealing with the problem requires a change in water distribution long before managers target reductions in abundances of mega-herbivores. the second example that we unpacked (table 1) relates to how stakeholders affected sanparks’ achievement of its suite of strategic objectives. the concern was over the influence of a bad relationship between augrabies national park and its stakeholders, on the park’s ability to carry out certain activities nearby such as anti-poaching. unpacking this concern revealed various sources of stakeholder misconceptions about sanparks and, hence, ways of establishing a better relationship with them. explicit measures of the park forum and other community meetings were then identified to track whether and/or how stakeholder perceptions were changing through pro-active engagement activities with stakeholders. this case illustrates how the unpacking of a mechanism at a social level is analogous to unpacking a biodiversity concern. the mechanism, as well as how to address the concern and measure it, is defined in much the same way as with the previous example. the final example addresses the third pillar of sanparks’ strategic objectives: providing benefits to people, in this case through the experience of tourism (table 1). this example addresses a tourism concern about lack of game observations by visitors to marakele national park. unpacking the mechanism revealed that viewing game was dependent on whether game and tourists were at the same place at the same time, which was constrained by the existing road network and the timing of game drives. monitoring should therefore be focused on tourists’ spatial use of this infrastructure over time, related to large mammal space use. unpacking this concern revealed that the tacit assumption of increasing game densities to improve a tourist product was not valid, and addressing the concern required a more nuanced approach involving spatial and temporal considerations. this example illustrates that the process does allow for incorporating uncertainty, because, in this instance, the risk component is not clearly understood. whilst the three examples represent very different kinds of concerns, the unpacking approach suggested here serves to illustrate how it may greatly enable the generation of common understanding and outcomes within the mandate of a conservation agency. evolution of the tpc concept (back to top) perhaps the most significant area of tpc development within sanparks has therefore been the use of the framework described above. this framework has allowed for the formation of important linkages between the objectives hierarchy and appropriate tpcs by unpacking the mechanisms and modulators that create particular management concerns. practical experiences in the smfs have demonstrated how this can also reduce conflict between members of the smf who hold different viewpoints, thereby easing decision-making and improving our understanding of the system. there are several examples of rigorously determined biophysical tpcs for the knp, particularly regarding those for fire (van wilgen et al. 2003). however, beyond the knp there has been a greater focus on ensuring that links are made with objectives, to ensure that appropriate tpcs are identified for a particular objective. having established these links, there is now a need to focus on the quantification of these tpcs for other parks. yet there is also room for developing the conceptual basis of tpcs further (duncan & wintle 2008). for example, setting directionality as a prerequisite of a tpc is the central tenet of revisiting tpcs in the sanparks context (ferreira et al. 2011). directionality can generalise the applicability of tpcs across parks, because well protected parks can have tpcs for which the rate of change in a measure centres on zero, whilst that for degraded parks centres away from zero. directionality also generalises across ecological, social, economical and financial spheres (biggs et al. 2011), a property that bodes well for the social learning required in adaptive management (cundhill & fabricius 2009). in addition, because tpcs are based on varying degrees of uncertainty, resource managers new to sam are still uncomfortable with their use (duncan & wintle 2008). it is therefore critical that the development process moves towards making tpcs more tangible to resource managers to avoid an implementation crisis (rogers 2003). we suggest that the degree of uncertainty associated with any particular tpc be classified along a continuum from empirical-based, through experience-based, to hypothetical situations. this will allow scientists to prioritise the tpcs that require more hard data, whilst managers develop confidence in the use of tpcs. use of the tpc concept by other organisations whilst sanparks has been concerned with managing ecosystems, using tpcs to measure unacceptable change, the south african biodiversity institute (sanbi) has been striving to maintain representative vegetation types by meeting national targets. these operational approaches apparently contradict one another – from the sanparks perspective the state of the system can vary whilst still retaining its essential structure and function, whilst sanbi’s outlook was that at least a fixed amount of each representative unit needed to be ‘conserved’, although at a coarser national scale. maintenance of representative vegetation types carries ecological non-realities and short-sightedness on the part of advisory ecologists. however, ongoing interactions between sanparks and sanbi have resulted in a growing realisation that ‘thresholds’ and ‘targets’ are similar constructs. whilst sam ‘thresholds’ delineate the boundaries of the desired state when within them, ‘targets’ define a trajectory towards a desired state or condition from the outside. this emerging understanding has resulted in sanbi proposing the use of national biodiversity thresholds in place of the national biodiversity targets, representing decision end-points based on unacceptable, but justifiable, system change. challenges, uncertainties and the way forward (back to top) in addition to demonstrating the advances that sanparks has made in its implementation of sam, it is also useful to reflect on the challenges and uncertainties that constrain its development (see jacobson et al. 2006). in doing so, we focus on three challenge areas that have emerged during the smfs. challenges involved in adopting the new non-equilibrium paradigm possibly the most pervasive challenge to the implementation of adaptive management outside of the knp is the institutional difficulty of shifting to the new ecological paradigm. managers and scientists alike are faced with the daunting prospect of letting go of decades-old approaches to conservation management that involved intensive, largely command-and-control management (rogers et al. 2000) of problem species based largely on agricultural principles such as carrying capacity. moreover, this approach was underpinned by an equilibrium-based understanding of ecosystem dynamics. adopting the non-equilibrium paradigm involves ‘unlearning’ all of these principles and learning a whole suite of new and unfamiliar terminology. whilst the use of stocking rates provided a simple formula with which to make decisions about wildlife management historically, the use of tpcs to inform management decisions has proven intangible to many managers beyond the knp to date. and because tpcs were introduced to managers in the context of how to deal with flux, the non-equilibrium paradigm and adaptive management are still often used synonymously, rather than it being understood that adaptive management is merely the tool with which to address the complexity and uncertainty of non-equilibrium systems. critics of the non-equilibrium paradigm are therefore quick to dismiss the use of tpcs to inform management decisions. finally, adoption of the non-equilibrium paradigm is constrained by pervasive perceptions amongst scientists and managers that small park size places limitations on ecosystem processes such that they cannot possibly operate in a similar manner to larger parks. sanparks now addresses this possible limitation by attempting to mimic such lost processes. institutional constraints to implementing sam rogers (2003) has provided a comprehensive review of the major institutional constraints facing the implementation of sam and we therefore do not attempt to repeat these here. rather, we outline some additional constraints that have emerged during the implementation of sanparks’ own adaptive planning and management process. one of the key limitations to the successful use of sam that has emerged during its development is reflected by the fact that conflicting objectives still manage to creep into parks’ objectives hierarchies, placing serious limitations on achieving their mandates. we suggest that this is as a direct result of lack of buy-in to the prioritisation of its high level objectives on a departmental level during the adaptive planning process. this is particularly the case with non-biophysical departments. consequently, departmental-scale management priorities are not aligned with corporate strategies or values at a park level. in situations where a park must sustain itself with limited or no state subsidies, the upshot is that the objective associated with the best (often short-term) financial outcome most often receives the higher priority, despite the risks this creates for failing to address the remaining objective. the vision and mission for each park reflect its stakeholders’ values, which, in turn, provide the rationale for the park’s operating principles. consequently, it should be possible to trace each and every management action back to stakeholder values through the park’s objectives hierarchy. apart from making future management decisions more defensible, this traceability of a shared value system also makes management decisions less prone to conflict amongst departments within a particular park. without park-wide departmental involvement in the adaptive planning process, the successful implementation of sam is severely compromised. feedback about what has been learnt through monitoring, research and management intervention is an essential component of sam that allows conservation organisations to improve the outcomes of their biodiversity conservation mandate incrementally (see cundhill & fabricius 2009; duncan & wintle 2008). although various performance evaluation and outcome assessment initiatives exist, sanparks still faces the challenge of adequately and timeously providing the necessary feedback in a format that ensures it will be useful and used. incorporated into this feedback should be a mechanism that tracks the progress made with sam explicitly at each stage of the adaptive management cycle, as well as across various scales within the organisation. such a tracking system represents the institutional memory that is essential for adaptive learning. finally, a remaining challenge is to outline the possible management actions that can be undertaken once a tpc is reached and to do so pro-actively or strategically, rather than once the tpc has already been exceeded. such strategic planning and weighing up of options is one requirement of sam that has, as yet, not been explicitly incorporated into the adaptive planning and management process. consequently, it has not yet fully matured in sanparks, and is one which, with adequate attention, should make the use of tpcs much more tangible to managers. making sam tangible to managers yet rigorous enough for scientists – the search for a ‘requisite simplicity’ managers of complex systems complain that science delivers information that is not useful for implementation, often because it is cluttered with complicated detail. stirzaker et al. (2010) suggest that identifying a ‘requisite simplicity’ (holling 2001) may provide the tangibility that managers seek from science. a requisite simplicity attempts to discard some detail, whilst retaining conceptual clarity and scientific rigor, and therefore helps conservationists move to a new position where they can more usefully benefit from new knowledge (detail). whilst the key elements of the requisite simplicity must be measureable, it is not falsifiable in the same sense as an experimental hypothesis (stirzaker et al. 2010). the requisite simplicity does represent a conceptualisation of the problem and conservationists’ measurements can show whether events are unfolding in a way that is consistent with this conceptualisation. it provides a basis for conservationists’ actions, although they expect that, as context and values change and knowledge is gained, it will be superseded (stirzaker et al. 2010). our mechanism framework fulfils all of the requirements of this definition. we therefore suggest that, in addition to providing the critical links between management objectives and actions, this recent modification of the adaptive planning process also provides the sought after requisite simplicity. conclusion (back to top) this paper has outlined the development of sam and its implementation beyond the knp. we have done this primarily by discussing biophysical issues, but have also illustrated how we incorporate social factors into the case studies that we presented. we have also highlighted the areas that still require substantial development before the use of sam within sanparks can be considered to be matured. we believe that these developments and deficiencies can provide useful lessons for other conservation agencies using adaptive management to achieve their conservation mandates. establishing a forum for regular discourse between scientists and managers has been a key supplementation to the sam process developed in the knp. through these smfs the use of a framework that links the objectives hierarchy with operational endpoints has been developed. this ensures the identification of appropriate indicators for each tpc, whilst developing a joint mental model of the system between scientists and managers. tpc quantification beyond the knp now requires attention and developing the concept of tpcs to incorporate such components as directionality and rate of change could benefit the sam process throughout sanparks and internationally. herein lies the importance of establishing and maintaining the smfs, as these forums are structured to address all of the limitations identified above. acknowledgments (back to top) we gratefully acknowledge mr cillie malan and dr harry biggs for their contribution to this paper in the form of the evaluation of comparative risk assessment outlined in box 1. references (back to top) biggs, h.c., 2003, ‘integration of science: successes, challenges, and the future’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience. ecology and management of savanna heterogeneity, pp. 469−487, island press, washington dc. biggs, h.c., ferreira, s.m., freitag-ronaldson, s. & grant-biggs, r., 2011, ‘taking stock after a decade: does the ‘thresholds of potential concern’ concept need a socio-ecological revamp?’, koedoe 53(2), art. #1002, 9 pages. doi:10.4102/koedoe.v53i2.1002 cundill, g. & fabricius, c., 2009, ‘monitoring in adaptive co-management: toward a learning based approach’, journal of environmental management 90(11), 3205−3211. doi:10.1016/j.jenvman.2009.05.012, pmid: 19520488 duncan, d. & wintle, b.a., 2008, ‘ towards adaptive management of native vegetation in regional landscapes’ in c. pettit et al. (eds.), landscape analysis and visualisation. spatial models for natural resource management and planning, springer-verlag gmbh, berlin. ferreira, s.m., deacon, a., sithole, h., bezuidenhout, h., deamane, e. & herbst, m., 2011, ‘from numbers to ecosystems and biodiversity – a mechanism approach to monitoring’, koedoe 53(2), art. #998, 12 pages. doi:10.4102/koedoe.v53i2.998 gaylard, a., cadenasso, m. & pickett, s.t.a., in press, ‘a conceptual framework for managing species as ecosystem drivers: spatial and temporal heterogeneity of surface water and elephant impacts in kruger national park’, bioscience. holling, c.s., 2001, ‘understanding the complexity of economic, ecological and social systems’, ecosystems 4, 390−405. doi:10.1007/s10021-001-0101-5 jacobson, s.k., morris, j.k., sanders, j.s., wiley, e.n., brooks, m., bennetts, r.e., et al., 2006, ‘understanding barriers to implementation of an adaptive land management program’, conservation biology 20(5), 1516−1527. doi:10.1111/j.1523-1739.2006.00476.x, pmid: 17002769 lindenmayer, d.b. & likens, g.e., 2009, ‘adaptive monitoring: a new paradigm for long-term research and monitoring’, trends in ecology and evolution 24(9), 482−486. doi:10.1016/j.tree.2009.03.005 martin, j., runge, m.c., nichols, j.d., lubow, b.c. & kendall, w.l., 2009, ‘structured decision making as a conceptual framework to identify thresholds for conservation and management’, ecological applications 19(5), 1079−1090. doi:10.1890/08-0255.1, pmid: 19688917 mckinsey & company, 2002, sanparks and mckinsey final meeting report, following strategic review, sanparks, skukuza. pickett, s.t.a., cadenasso, m.l. & benning, t.l., 2003, ‘biotic and abiotic variability as key determinants of savanna heterogeneity at multiple spatio-temporal scales’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience. ecology and management of savanna heterogeneity, pp. 22−40, island press, new york. pickett, s.t.a., kolasa, j. & jones, c.g., 2007, the nature of theory and the theory of nature, 2nd edn., academic press inc., san diego. rogers, k.h., 2003, ‘adopting a heterogeneity paradigm: implications for biodiversity management in protected areas’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience. ecology and management of savanna heterogeneity, pp. 41−58, island press, new york. rogers, k.h., roux, d. & biggs, h., 2000, ‘challenges for catchment management agencies: lessons from bureaucracies, business and resource management’, water sa 26(4), 505–511. rogers, k.h. & sherwell, t., 2008, a framework for developing and implementing management plans for south african national parks, viewed n.d., from http://www.sanparks.org/parks/kruger/conservation/scientific/key_issues/plans/adaptive/default.php stirzaker, r., biggs, h., roux, d. & cilliers, p., 2010, ‘requisite simplicities to help negotiate complex problems’, ambio: a journal of the human environment 39(8), 600−607. van wilgen, b.w., trollope, w.s.w., biggs, h.c., potgieter, a.l.f. & brockett, b.h., 2003, ‘fire as a driver of ecosystem variability’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience. ecology and management of savanna heterogeneity, pp. 149−170, island press, new york. venter, f., naiman, r.j., biggs, h.c. & pienaar, d.j., 2008, ‘the evolution of conservation management philosophy: science, environmental change and social adjustments in kruger national park’, ecosystems 11, 173−192. doi:10.1007/s10021-007-9116-x article information authors: mark roberts1 kevin mearns2 victoria edwards3 affiliations: 1school of environmental design and management, university of portsmouth, united kingdom2department of environmental sciences, university of south africa, south africa 3school of real estate and planning, henley business school, university of reading, united kingdom correspondence to: kevin mearns postal address: private bag x6, florida 1710, south africa dates: received: 27 mar. 2013 accepted: 06 nov. 2013 published: 24 june 2014 how to cite this article: roberts, m., mearns, k. & edwards, v., 2014, ‘evaluating the effectiveness of guided versus non-guided interpretation in the kruger national park, south africa’, koedoe 56(2), art. #1160, 8 pages. http://dx.doi.org/10.4102/ koedoe.v56i2.1160 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. evaluating the effectiveness of guided versus non-guided interpretation in the kruger national park, south africa in this original research... open access • abstract • introduction    • literature review       • visitor satisfaction       • knowledge gain       • attitude change       • behavioural change    • evaluation of interpretive programmes • research method and design • results    • level of visitor satisfaction    • level of knowledge gain    • degree of attitude change    • intent to modify behaviour • ethical considerations • trustworthiness • discussion    • visitor preference of interpretation type • conclusion • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ in the face of growing pressure placed on the natural environment, the study on which this article is based considered the effectiveness of interpretive provision in mitigating the harmful effects of tourism on the environment. the aim of this research was to determine whether guided or non-guided interpretation is most effective in reaching the stated goals of interpretation. the four key goals of interpretation, namely visitor satisfaction, knowledge gain, attitude change and modification of behaviour intent, were used in the assessment of the relative effectiveness of guided and non-guided interpretation in the kruger national park, south africa. through comparing responses to questionnaires from post-visit samples and observing both guided and non-guided interpretation, the research found that guided interpretation was only marginally more effective in reaching the four key goals of interpretation than the non-guided interpretive media. guided interpretation was found to be more effective in terms of visitor satisfaction, whilst guided and non-guided interpretation had only marginal differences in effectiveness in relation to knowledge gain, attitude change and intent to modify behaviour.conservation implications: the necessity of implementing an appropriate interpretation programme within protected areas cannot be overemphasised. the interpretation programme should be designed to include elements of both guided and non-guided interpretation in order to achieve a predetermined goal. the effectiveness of the programme should be evaluated periodically and amended where appropriate. introduction top ↑ the worldwide increase in tourism to natural areas has led to the notion that we are ‘loving national parks to death’ and this is becoming a major concern for national park managers (vaske, donnelly & whittaker 2000). the inherent fragile and unique nature of national parks and conservation areas suggests that they will be more severely affected by human activity (weaver 2000). one of the most pressing problems of many national parks is how to cope with the growing number of visitors (world tourism organisation [wto] 1992). an important measure of both the success and the sustainability of tourism in national parks and protected areas is the management of visitor impacts to ensure the long-term protection of the natural and cultural resources as well as continued visitor enjoyment and use (marion & farrell 1998). allowing visitor use makes impacts unavoidable; these impacts need to be managed. visitor management in national parks seeks to redress this situation through the protection of natural and cultural resources and the provision of tourist activities and experiences. without effective visitor management, tourism can lead to adverse impacts on the natural, cultural and heritage environments to the extent that they may also negatively affect visitor satisfaction (shackley 1998). effective interpretation can make a substantial contribution to improving the sustainability of tourism and, as a result, reduce the associated impacts from tourism (moscardo 1996, 1998). interpretation has already been recognised as no longer being a frill or a ‘luxury’ but an essential management function for every park or recreation area (herbst 1979). consequently, national park managers agree on the necessity to introduce interpretation programmes. the questions that arise for these managers within a constrained financial paradigm relate to the type of interpretation that should be implemented, as well as the effectiveness of the chosen type of interpretation. this research examined visitor perceptions in order to determine whether guided or non-guided interpretation is most effective in reaching predetermined goals of interpretation. literature review tourism is increasingly being seen as an important means through which the natural resources of national parks and conservation areas can generate revenues to ensure their long-term economic sustainability (weaver 2000). tourism growth to natural areas is now reaching levels where the impacts may be just as detrimental as other industrial sectors. concern for the environmental consequences of tourism has grown since the 1990s (buckley & pannell 1990; hjalager 1996; wto 1995). interpretation has been recognised as an important solution to reduce these impacts whilst also educating the visitor to achieve the goals of sustainable tourism development (ballantyne et al. 2007). sustainable living and conservation have become hotly debated topics over recent years, with many commentators seeing social learning as a high priority on the environmental agenda. through the concept of citizenship, it is thought that teaching in formal or informal settings might encourage and enable all members of society to contribute to the ‘common good’. with the high visitor numbers recorded to nature reserves and national parks, it has been acknowledged that interpretation could aid development of environmental citizenship and, as a result, it is seen as an essential component of national park management.although there are numerous definitions of interpretation, they all centre on the idea of translating information from the scientist, the historian and the manager to the visitor or layperson (ed. mcarthur 1998; ward & wilkinson 2006). the most widely accepted definition of interpretation is that of tilden (1977): interpretation is an educational activity which aims to reveal meaning and relationships through the use of original objects, by first-hand experience, and by illustrative media, rather than simply to communicate factual information. (p. 8) as tilden’s definition suggests, interpretation is an approach to communicating which stresses the transfer of ideas and relationships rather than just facts and figures. this separates interpretation from conventional education. since tilden first published his work in 1977, many people and organisations have adopted their own interpretation or definition of ‘interpretation’. the overarching theme of these definitions indicates that there is consensus that interpretation is a means of communicating visitor relationships with the environment, not just imparting scientific facts. environmental interpretation has been in existence for a long time. in 1919, the us national park service started to develop guided activities for visitors (aldridge 1973). at the same time in south africa, guides for visitors to national parks were being produced. the recent development of the interpretive philosophy and techniques has not just been related to natural areas, but has moved to include all aspects of heritage of a geographical area which are worth conserving for future generations. interpretation is increasingly believed to play a role in influencing visitor beliefs, attitudes, knowledge and behaviours (hughes & morrison-saunders 2005; kohl 2004; kuo 2002; moscardo 1998). ham, housego and weiler (2005) suggest that when interpretation is carried out in protected areas, it is expected that it will encourage appropriate use, support responsible management and foster long-term conservation goals. ward and wilkinson (2006) divide interpretation into two types, namely personal and non-personal. wearing et al. (2007) concur with this notion and divide interpretation into two categories, (1) guided (e.g. guided walks) or (2) non-guided (e.g. boards) on the basis of delivery technique. guided interpretive programmes are developed to utilise direct contact between the public and an interpreter or guide. non-guided interpretive programmes are developed so that visitors do not have physical interaction or discourse with an interpreter or guide, but rather connect through different objects, media and resources. the investigation on which this article is based compared these two types of interpretation and their effectiveness. although there is no consensus on what successful or effective interpretation is, a number of authors have argued that interpretation should do one or more of the following, (1) enhance visitor experiences, (2) protect resources at sites, (3) protect visitors, (4) increase public support for an agency and its management policies, (5) add to or broaden visitors’ perspectives about a place or idea and (6) enhance their knowledge and foster positive attitudes and behaviours with respect to the natural and cultural environment (beck & cable 1998; ham 1992; ham & krumpe 1995; ham et al. 2005; knudson, cable & beck 1995; lewis 1980; moscardo, ballantyne & hughes 2006; regnier, gross & zimmerman 1994; sharpe 1982; ward & wilkinson 2006). as a result, the goals of interpretation are, (1) to satisfy visitors, (2) to instil knowledge gain, (3) to achieve attitude change and, consequently, (4) to achieve behavioural change (hughes & morrison-saunders 2005; kohl 2004; kuo 2002; moscardo 1998). for the greatest achievement of all four goals, managers need to know how to deliver interpretation effectively. lubbe (2003) argues that in south africa there is a dire need for good interpretation, so that tourists ‘will keep coming back’. whilst what is communicated to tourists is important, the manner in which it is communicated is equally important (lubbe 2003). the aim of this research was to compare the effectiveness of guided and non-guided interpretation in achieving the four stated goals of interpretation in the kruger national park (knp) and to draw conclusions for a wider audience. the research analysed visitor perceptions after guided or non-guided interpretation experiences in relation to the stated goals of interpretation. uzzell (1998) contends that the evaluation of interpretation is recommended, but is rarely conducted. it is essential for managers to know if their chosen management tool is effective, thus preventing managers from spending scarce resources on tools that do not work (brown, mccool & manfredo 1987; gunderson et al. 2000). the most commonly evaluated outcomes of interpretation in research studies have been increased knowledge and attitude change, as they are generally easier to measure. results of previous studies evaluating interpretation have left uncertainty over the effectiveness of interpretation – for this reason further research is necessary. weiler and smith (2009) state that there is an urgent need to develop strategies for assessing the effectiveness of interpretation programmes. this study attempted to aid in the development of a method for the assessment of the effectiveness of interpretation programmes in relation to the four goals of interpretation. visitor satisfaction making recreational experiences more enjoyable is almost always a goal of interpretation (knudson et al. 1995). pearce and moscardo (1998) found that interpretation at the skyrail rainforest cableway near cairns, australia was positively linked to enhanced visitor satisfaction. in turn, ham and weiler (2000) found that five interpretive services in the panama canal protected areas contributed to overall tourist satisfaction. interpretation must be enjoyable in order to hold visitors’ attention (ham 1992; sharpe 1982). although entertainment and satisfaction are not the only goals of interpretation they are important indicators of successful interpretation. knowledge gain interpretation can reveal a world many may not have seen before (ham 1992; ed. larsen 2003; pastorelli 2003; regnier et al. 1994; ward & wilkinson 2006). lee and balchin (1995) suggest that in psychological terms the aim of interpretation is to achieve learning and, by helping visitors to better understand the idea of coexisting with the environment, their awareness of their place in the total environment may be enhanced (sharpe 1982; tisdell & wilson 2001). phillips (1989) believes that the chief measure of interpretive effectiveness is whether information has been conveyed to the visitor. orams (1996) concurs with this statement and further suggests that through increasing visitors’ knowledge or understanding, interpretation could potentially prompt more environmentally responsible behaviour. attitude change if interpretation succeeds in increasing visitors’ understanding of the places they visit, it is hoped that it will lead to the respect for an area (bramwell & lane 1993). interpretation aims to broaden awareness and concern and to assist people in building relationships with, or understanding interrelationships between, what they are observing at the site and in their lives (ed. larsen 2003; lewis 1980). iozzi (1989) argues that interpretation must include activities that are specifically designed to change the attitudes of the visitor. behavioural change if interpretation is to be an effective technique in managing tourist–nature interaction and achieving sustainable development, it should do more than simply increase knowledge and understanding; it should prompt behaviour change (moscardo 1996; orams 1997). at a basic level, interpretation should seek to manage people’s behaviour through the encouragement of their own personal development and self-realisation so that the way they behave is respectful of the potential impact their actions may have (krippendorf 1987; orams 1997). it is proposed that through interpretation, people will be encouraged to act in more appropriate ways (ballantyne et al. 2007; o’riordan, shadrake & wood 1989; stewart, hayward & devlin 1998). higham and carr (2002) suggest that interpretation programmes may foster behavioural change relating to domestic lifestyle, which may contribute to the long-term benefit of the environment. evaluation of interpretive programmes there have been a number of studies since the 1970s that have evaluated the influence of interpretation, with the large majority of these focusing on knowledge or attitude change amongst visitors. some authors (mcgehee & santos 2005; russell & hodson 2002; ryan, hughes & hirgwin 2000) have highlighted the importance of evoking emotion as a means to encourage positive environmental behaviour amongst visitors. munro, morrison-saunders and hughes (2008) suggest that this may be difficult to achieve through text-based signs compared with interaction through a guide. other authors (mallick & driessen 2003; orams 1996; townsend 2003) have linked programme success with providing visitors with an opportunity to act upon newly formed attitudes and intentions. there is evidence that guided interpretation enhances the quality of the visitor experiences. forestry tasmania (1994), hughes (1991), moscardo (1998) and schänzel and mcintosh (2000) linked success of interpretive programmes with guided communication. the evaluation of interpretive programmes is difficult for two main reasons. firstly, there can be many ensuing experiences as a result of interpretation, such as inspiration and enjoyment, which cannot be measured by standard methodologies (beckmann 1991). secondly, interpretation takes place in recreational areas where many other aspects influence both visitors and interpretation: these influences can be difficult to control (mcdonough 1986). in a literature study reviewing 21 articles that evaluated the outcomes of environmental interpretation programmes, munro et al. (2008) found that about half of the studies had used post-experience sampling to evaluate interpretive influences on visitors. the munro et al. (2008) study illustrates the spectrum of evaluation methods used in the field and that any attempt to apply a single evaluative process in environmental interpretation would probably favour certain methodologies. moscardo (1998) and beaumont (2001) propose that research efforts to measure the effectiveness of interpretation can be put into two categories. the first is research that measures visitors’ perceived satisfaction, knowledge gain, attitude and behavioural intentions (e.g. asking visitors to reflect on what they think they learned as a result of their experience). the second category includes research that aims to measure actual outcomes. knowing how visitors feel when they come away from an experience is important as it provides a measure of the quality of the experience and visitors’ satisfaction with their experience. this article follows the first approach. there have been mixed results from studies that have evaluated the effectiveness of interpretation on influencing visitors’ enjoyment, knowledge gain, attitude change and behavioural intentions. some studies have found that interpretation has increased visitor enjoyment and knowledge, modified attitudes and increased behavioural intentions, whilst other studies have found that interpretation has had no effect. munro et al. (2008) found that out of the 21 case studies reviewed, 19 considered the interpretive programme that was evaluated to be successful or at least partly successful. guided experiences are generally accepted as the most effective form of interpretation, provided that it is well planned and implemented (armstrong & weiler 2002; beaumont 2001; brody, tomkiewicz & graves 2002; ham 1992; hughes & morrison-saunders 2005; luck 2003; tubb 2003). however, an extensive review of the literature revealed that no direct comparison has been made between the relative effectiveness of guided and non-guided interpretation. this is supported by weiler and smith (2009), which confirms that there is a general lack of research on different interpretation techniques. the study that informed this article made a direct comparison between guided and non-guided interpretation in the knp. research method and design top ↑ moscardo (1998) and beaumont (2001) separate interpretation evaluation into two categories, (1) visitor perceptions and (2) actual outcomes. the research reported in this article followed the first methodological approach, making visitor perceptions the focus of the study in evaluating the effectiveness of interpretive programmes. the research used a post-visit survey questionnaire to identify visitors’ perceived outcomes of interpretation, assessed against the four main goals of interpretation, in order to evaluate the effectiveness of the two types of interpretation (30 guided and 30 non-guided questionnaires). observations were also used to verify data collected in the survey responses and to add depth of understanding to the survey data (kuo 2002; novey & hall 2007; tubb 2003). the knp was proclaimed in 1926; since then tourism has developed significantly and, in 2008, the park already attracted in excess of one million visitors annually (van der merwe & saayman 2008). the knp is located in the north-east of south africa, bordering the limpopo national park of mozambique. this research was conducted in 2010 in the letaba rest camp in the knp. letaba is situated on the bend of the letaba river, midway between the southern and northern boundaries of the knp. the guided interpretation evaluation was conducted after night drives that were undertaken in the company of trained field guides. night tours last for two hours, during which the guides accompany up to 24 people whilst offering commentary on geographical sites and wildlife sightings. the non-guided interpretation was undertaken after visitors exited the elephant hall museum in letaba, which covers elephant evolution, biology, behaviour, ecology and research. the elephant hall also showcases the ivory of eight of the knp’s greatest tuskers, including six of the ‘magnificent seven’. this non-guided experience consists of several interpretive styles, which include both interactive and standard displays. the visitor survey was designed in order to determine visitor satisfaction, perceived knowledge gain, attitude change and the visitor’s intent to change their behaviour. socio-demographic questions were also included in order to obtain a profile of the visitors participating in the study. the questionnaire was designed to allow for comparison between guided and non-guided interpretation. this meant that the questionnaire was designed to be non-specific to site and interpretive media (munro et al. 2008). the questionnaire used a combination of open-ended and closed-ended questions, as well as likert scale questions. in the likert scale questions respondents were shown a statement and were asked to indicate their level of agreement or disagreement with that statement with a ranking score of 1–5 (1 being ‘strongly disagree’, showing lower effectiveness levels; 2, 3 and 4 representing a continuum, until 5 being ‘strongly agree’, showing higher levels of effectiveness). these responses were used to determine the effectiveness of the different interpretation techniques in terms of the four goals of interpretation, namely visitor satisfaction, knowledge, attitude and behavioural intent. the survey was conducted on site at exit points of guided and non-guided interpretive settings. data were collected from a self-administered questionnaire. the survey scope was explained to each respondent. the researchers also explained that the survey was anonymous and confidential. the questionnaire was then handed to any visitor over the age of eighteen. visitors were asked to reflect on the experience they had just received and complete the questionnaire on site. the researchers were present to clarify any possible issues or concerns. observations were conducted as a secondary tool of research. as the actions and behaviour of people are a central aspect in this study, a natural and obvious technique is to watch what they do, record this information and then describe, analyse and interpret what has been observed. the researchers participated in the tours and the interpretive experience whilst observing visitors’ actions, behaviours and interpretation participation, as well as assessing both guided and non-guided interpretation programmes. observations were undertaken as discretely as possible to limit the intrusion on the guides and the visitors. digital photographs were taken as a memory aid. novey and hall (2007) explains that by immersing themselves in the research setting and remaining anonymous, researchers can observe dimensions of visitors’ behaviour, interactions and actions. observations added a depth of understanding to data collected through the visitor survey. results top ↑ the discussion of the results follows the four goals of interpretation, namely, (1) visitor satisfaction, (2) knowledge gain, (3) attitude change and (4) intent to modify behaviour. level of visitor satisfaction guided interpretation in the knp was found to be statistically more effective than the non-guided interpretive media at raising levels of satisfaction (p = 6 × 10-5, p < 0.05). the overall average likert-scale responses by the visitors in term of the series of statements in relation to visitor satisfaction was 4.09 for non-guided interpretation and 4.49 for guided interpretation, indicating that the visitor satisfaction levels were higher for the guided interpretation than the non-guided interpretation. amongst the reasons given by respondents for their heightened satisfaction levels were, (1) the excellent knowledge of the guide, (2) the guide’s personality, (3) seeing animals and (4) the ability of the guide to answer questions when asked. the guide was also considered to be very humorous and this seemed to hold the visitors’ attention, drawing a good response from them. the visitors seemed to be very relaxed during the drive, joking and laughing with each other, which could reflect the guide’s ability to act as a successful host. however, although guided interpretation was found to be more enjoyable, some reasons for dissatisfaction were given. for example, visitors noted that they were given the incorrect time for the tour departure and that the guide was sometimes difficult to understand because of a heavy accent and being softly spoken. reasons for satisfaction with the non-guided interpretation were given as information, elephant skulls and skeleton displays and the interpretive displays. for the non-guided interpretation, respondents indicated that they had enjoyed the interactive displays that were present at the elephant hall. this suggests that a large proportion of respondents enjoyed learning through interactive means. interactive displays showing the heights of elephants compared to humans were very popular with visitors. visitors were observed measuring themselves against elephant and human heights at various ages. reasons for dissatisfaction in relation to non-guided interpretation by respondents indicated that there were not enough interactive displays and that many of the displays were old. the findings in relation to visitor satisfaction in this research concur with the findings of moscardo (1998) and schänzel and mcintosh (2000) that visitor satisfaction is higher in guided interpretation programmes. level of knowledge gain the non-guided interpretive media at the knp received a higher mean likert-scale score for respondents’ perceived knowledge gain (3.68), compared to 3.63 for guided interpretation; however, the difference was not found to be statistically significant (p = 0.03, p > 0.05). when asked to mention two things they had learnt as a result of the interpretation, over 80% of the guided tour respondents were able to give one or more specific details they had learnt compared to only 30% of those respondents who had attended the non-guided interpretation. this shows that those who had attended the guided interpretation in the knp were able to recall facts passed on by the interpreter better than those who participated in the non-guided interpretation. the guide of the nightly game drives was able to hold the visitors’ attention with anecdotes, enthusiasm and knowledge. in contrast, the researchers noted that, on average, visitors spent between 20 min and 25 min in the elephant hall (non-guided interpretation). the researchers believe that it is not possible to read all the information on display in the elephant hall within that time scale. a time estimate for reading all the interpretive material in the elephant hall will be around 3 h. the non-guided interpretation was unable to hold the visitors’ attention for long periods of time, which has an impact on the knowledge gained by the visitor. section signs within the elephant hall tell the visitor what they are about to read in the forthcoming section. this interpretation draws the visitor’s eye in an effective manner. however, the majority of the interpretation on show in the elephant hall is not of recent design and has a great deal of information packed onto the displays, which means that there is a large amount for visitors to read and take in. some of the displays are difficult to follow, as there is no obvious route to follow and display cases offer no natural flow to visitors. the small blocks of writing within the cases appeared to be unappealing to the visitors as observing people’s behaviour revealed few people stopped to read about the contents of the display cases. the researchers also noted that the explanatory cards contained a great deal of writing. large amounts of writing may affect the visitors’ desire to read further, resulting in the possibility that the viewers ignore the signage completely. however, there seemed to be a greater perceived knowledge gain from non-guided interpretation amongst the respondents. degree of attitude change guided interpretation was found to be more effective at provoking attitude change amongst respondents, with an average likert-scale score of 4.31, compared to 4.29 for non-guided interpretation. however, the difference was not found to be statistically significant (p = 0.47, p > 0.05). in terms of attitude change, guided and non-guided interpretation delivered similar results. intent to modify behaviour guided interpretation received a marginally higher mean score for respondents’ intent to change behaviour (3.78) than that found for respondents of the non-guided interpretation (3.73). however, the difference was not found to be statistically significant (p = 0.48, p > 0.05). both groups of respondents had high levels of education and showed prior environmental commitment through membership of environmental organisations and completion of courses and training, which may have affected their behavioural intent. at the start of the nightly game drives in the knp, the guide gives a safety and behaviour awareness talk before the game viewing vehicle leaves the camp. the guide explains that visitors should remain in the vehicle at all times, refrain from allowing any body parts to protrude from the vehicle and, for the safety of all visitors, they are advised to remain seated whilst the vehicle is in motion. the researcher noticed that on several occasions the third message was not adhered to as many guests chose to stand whilst the vehicle was in motion, which suggests that the introductory safety message was not effective in that respect. according to hu and wall (2012), tour guides play a critical role in striving towards sustainable tourism in national parks and protected areas. the intent to modify behaviour was very similar in guided and non-guided interpretation. ethical considerations top ↑ ethical approval for undertaking the research was obtained through the university of portsmouth, school of environmental design and management ethics committee. the research was also conducted under the consent and approval of the south african national parks board social sciences research department coordinator.all the participants in the study took part voluntarily, after they were informed of the objectives of the study and the completion of an informed consent agreement. all participants were entitled to withdraw from the study at any point. the completed questionnaires were also completed anonymously and confidentially. the original completed questionnaires were scanned and digitally stored with password protection, after which the original questionnaires were destroyed. the password protection was also maintained through the data encoding and analysis phases of the research. trustworthiness top ↑ to ensure validity and reliability of responses with the survey technique, the questionnaire was pre-tested on five visitors exiting the elephant hall at letaba camp in the knp. after the collection of the five pretested questionnaires, the results were briefly analysed in order to determine if the required research instrument would deliver the expected results. this was found to be the case. during the entire period during which the questionnaires were being completed, the researchers were available should additional clarity be sought by the participants. discussion top ↑ using the above results, the average scores for all four goals of interpretation are listed in table 1. the combined interpretation scores for the four goals of interpretation indicated that there is only a marginal difference in the effectiveness between guided (4.05) and non-guided (3.95) interpretation, the only really notable difference being in terms of visitor satisfaction, which is much higher in guided interpretation. table 1: effectiveness scores measured on a 5-point likert scale against the four goals of interpretation. in order to further explore the effectiveness of the different interpretation methods, visitors were asked to indicate their preferences in terms of interpretation type. these results are discussed below. visitor preference of interpretation type firstly, 83% and 87% of the respondents indicated a preference for guided interpretation to non-guided interpretation, respectively (table 2). the visitors who completed both the guided and the non-guided interpretation indicated that their preferred type of interpretation was guided interpretation. secondly, 39% of the respondents indicated that they believe guided interpretation to be more educational, whilst 22% of the respondents preferring guided interpretation attributed their preference to enjoying the personal aspect of having a guide (table 3). a respondent from the non-guided elephant hall interpretation attributed their preference of guided interpretation to it being ‘more informative’ and to their view that guided interpretation requires ‘less effort in receiving information’. table 2: preferred interpretation type. table 3: reasons for the preference of guided interpretation. visitors preferring non-guided interpretation stated that freedom (50%) and reduced costs (18%) were the most important reasons for respondents to prefer non-guided interpretation. conclusion top ↑ increased tourism and the growing pressure on tourist sites and the environment have called for national parks and conservation areas to recognise the need for appropriate visitor management and interpretation. the aim of the research reported in this article was to utilise visitor perceptions to determine whether guided or non-guided interpretation is most effective in reaching the goals of interpretation. it was found that only marginal differences in the effectiveness between guided and non-guided interpretation exist. although visitor satisfaction levels were higher in guided interpretation, the other three goals of interpretation, namely knowledge gain, attitude change and intent to change behaviour were only marginally different. the research has added depth of understanding to the general agreement amongst many authors that guided or personal forms of interpretation are the most effective, provided they are well planned and implemented (armstrong & weiler 2002; beaumont 2001; brody et al. 2002; ham 1992; hughes & morrison-saunders 2005; luck 2003; tubb 2003). this study provides valuable evidence that the effectiveness of interpretation can be measured through ascertaining visitor perceptions in relation to the four goals of interpretation. interpretive planners should redesign both guided and non-guided interpretation programmes within the kruger national park to be more effective in terms of all four goals of interpretation. interpretation should be seen as being vital to the conservation and preservation of natural areas as it aims to change visitors’ attitudes and behaviours towards creating more environmentally responsible tourists that will ensure the long-term viability and sustainability of natural areas. acknowledgements top ↑ competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions m.r. (university of portsmouth) conducted the research as part of his masters dissertation in ecotourism and co-authored the article. k.m. (university of south africa) supervised the academic research and co-authored the article, whilst v.e. (university of reading) was the supervisor of the academic research. references top ↑ aldridge, d., 1973, ‘upgrading park interpretation and communication with the public’, in iucn (ed.), second world conference on national parks, yellowstone and grand teton, usa, 18–27 september 1972.armstrong, e. & weiler, b., 2002, ‘getting the message across: an analysis of messages delivered by tour operators in protected areas’, journal of ecotourism 1(2&3), 104–121. http://dx.doi.org/10.1080/14724040208668118 ballantyne, r., packer, j., hughes, k. & dierking, l., 2007, ‘conservation learning in wildlife tourism settings: lessons from research in zoos and aquariums’, environmental education research 13(3), 363–383. http://dx.doi.org/10.1080/13504620701430604 beaumont, n., 2001, ‘ecotourism and the conservation ethic: recruiting the uninitiated or preaching to the converted?’, journal of sustainable tourism 9(4), 317–341. http://dx.doi.org/10.1080/09669580108667405 beck, l. & cable, t., 1998, interpretation for the 21st century, sagemore publishing, champaign. beckmann, e.a., 1991, ‘environmental interpretation for education and management in australian national parks and other protected areas’, phd thesis, department of ecosystem management, university of new england. bramwell, b. & lane, b., 1993, ‘interpretation and sustainable tourism: the potential and pitfalls’, journal of sustainable tourism 1(2), 71–80. brody, m., tomkiewicz, w. & graves, j., 2002, ‘park visitors’ understandings, values and beliefs related to their experience at midway geyser basin, yellowstone national park, usa’, international journal of science education 24(11), 1119–1141. http://dx.doi.org/10.1080/09500690210134820 brown, p.j., mccool, s.f. & manfredo, m.j., 1987, ‘evolving concepts and tools for recreation user management in wilderness: a state-of-knowledge review’, in r.c. lucas (ed.), proceedings of national wilderness research conference: issues, state-of-knowledge, future directions, us department of agriculture, forest service, intermountain research station, ogden. buckley, r. & pannell, j., 1990, ‘environmental impacts of tourism and recreation in national parks and conservation reserves’, journal of tourism studies 1(1), 24–32. forestry tasmania, 1994, guided nature-based tourism in tasmania’s forests: trends, constraints and implications, forestry tasmania, hobart. gunderson, k., barns, c.v., hendricks, w.w. & mcavoy, l.h., 2000, wilderness education: an updated review of the literature and new directions for research and practice, us department of agriculture, forest service, rocky mountain research station, ogden. ham, s., 1992, environmental interpretation: a practical guide for people with big ideas and small budgets, north american press, golden. ham, s. & krumpe, e., 1995, ‘how can site-based interpretation be used to change behaviors and contribute to biodiversity and ecosystem conservation in central america?’, paper presented at academy for educational development (greencom project), technical advisory group meeting, 16 october. ham, s. & weiler, b., 2000, profile of tourists in the panama canal watershed protected areas: final research report, academy for educational development and greencom project, washington, dc. ham, s., housego, a. & weiler, b., 2005, tasmanian thematic interpretation planning manual, tourism tasmania, hobart. herbst, r.l., 1979, ‘introduction’, trends 16(1), 2. higham, j. & carr, a., 2002, ‘ecotourism visitor experiences in aotearoa/new zealand: challenging the environmental values of visitors in pursuit of pro-environmental behaviour’, journal of sustainable tourism 10(4), 277–294. http://dx.doi.org/10.1080/09669580208667168 hjalager, a., 1996, ‘tourism and the environment: the innovation connection’, journal of sustainable tourism 4(4), 201–218. http://dx.doi.org/10.1080/09669589608667268 hu, w. & wall, g., 2012, ‘interpretation guiding and sustainable development: a framework’, tourism management perspectives 4, 80–85. http://dx.doi.org/10.1016/j.tmp.2012.05.003 hughes, k., 1991, ‘tourist satisfaction: a guided cultural tour in north queensland’, australian psychologist 26(3), 161–171. http://dx.doi.org/10.1080/00050069108257243 hughes, m. & morrison-saunders, a., 2005, ‘international activity participation, and environmental attitudes of visitors to penguin island: western australia’, society of natural resources 18, 611–624. http://dx.doi.org/10.1080/08941920590959587 iozzi, l.a., 1989, ‘what research says to the educator – part one: environmental education and the effective domain’, journal of environmental education 20(3), 3–9. http://dx.doi.org/10.1080/00958964.1989.9942782 knudson, d., cable, t. & beck, l., 1995, evaluation: interpretation of cultural and natural resources, venture publishing, state college. pmcid:pmc40522 kohl, j., 2004, ‘putting environmental interpretation to work for conservation in a park setting: conceptualizing principal conservation strategies’, applied environmental education and communication 4, 1–19. krippendorf, j., 1987, the holiday makers, heinemann, london. kuo, i.l., 2002, ‘the effectiveness of environmental interpretation at resource-sensitive tourism destinations’, international journal of tourism research 4, 87–101. http://dx.doi.org/10.1002/jtr.362 larsen, d. (ed.), 2003, meaningful interpretation – how to connect hearts and minds to places, objects and other resources, u.s. national park service, fort washington. lee, t. & balchin, n., 1995, ‘learning and attitude change at british nuclear fuel’s sellafield visitors centre’, journal of environmental psychology 15, 283–298. http://dx.doi.org/10.1006/jevp.1995.0025 lewis, w., 1980, interpreting for park visitors, eastern acorn press, philadelphia. lubbe, b.a., 2003, tourism management in south africa, pearson education, cape town. luck, m., 2003, ‘education on marine mamma tours as agent for conservation – but do tourists want to be educated?’, ocean & coastal management 46(9), 943–956. http://dx.doi.org/10.1016/s0964-5691(03)00071-1 mallick, s.a. & driessen, m.m., 2003, ‘feeding of wildlife: how effective are the “keep wildlife wild” signs in tasmania’s national parks?’, ecological management and restoration 4(3), 199–204. http://dx.doi.org/10.1046/j.1442-8903.2003.00157.x marion, j.l. & farrell, t.a., 1998, ‘managing ecotourism visitation in protected areas’, in k. lindberg, m.e. wood & d. engeldrum (eds.), ecotourism: a guide for planners and managers, vol. 2, pp. 155–181, the ecotourism society, washington, dc. mcarthur, s. (ed.), 1998, introducing the undercapitalised world of interpretation, vol. 2, the ecotourism society, washington, dc. pmid:10451319 mcdonough, m.h., 1986, ‘evaluation: the interpreter’s dilemma’, in g.e. machlis (ed.), interpretive views: opinions on evaluating interpretation in the national parks service, pp. 99–105, national parks conservation association, washington, dc. pmcid:pmc239163 mcgehee, n.g. & santos, c.a., 2005, ‘social change, discourse and volunteer tourism’, annals of tourism research 32(3), 760–779. http://dx.doi.org/10.1016/j.annals. 2004.12.002 moscardo, g., 1996, ‘mindful visitors – heritage and tourism’, annals of tourism research 23(2), 376–397. http://dx.doi.org/10.1016/0160-7383(95)00068-2 moscardo, g., 1998, ‘interpretation and sustainable tourism: functions, examples and principles’, journal of tourism studies 9(1), 2–13. moscardo, g., ballantyne, r. & hughes, k., 2006, interpretive signs and exhibits – principles in practice, fulcrum books, golden. pmcid:pmc1600016 munro, j.k., morrison-saunders, a. & hughes, m., 2008, ‘environmental interpretation evaluation in natural areas’, journal of ecotourism 7(1), 1–14. http://dx.doi.org/10.2167/joe137.0 novey, l.t. & hall, t.e., 2007, ‘the effect of audio tours on learning and social interaction: an evaluation at carlsbad caverns national park’, science education 91(2), 201–221. http://dx.doi.org/10.1002/sce.20184 orams, m.b., 1996, ‘using interpretation to manage nature-based tourism’, journal of sustainable tourism 4(2), 81–94. http://dx.doi.org/10.1080/09669589608667260 orams, m.b., 1997, ‘the effectiveness of environmental education: can we turn tourists into ”greenies”?’, progress in tourism and hospitality research 3(4), 295–306. http://dx.doi.org/10.1002/(sici)1099-1603(199712)3:4<295::aid-pth85>3.0.co;2-d o’riordan, t., shadrake, a. & wood, c., 1989, ‘interpretation, participation and national park planning’, in d. uzzell (ed.), heritage interpretation – the natural and built environment, pp. 179–189, belhaven, london. pmcid:pmc1385425 pastorelli, j., 2003, enriching the experience: an interpretive approach to tour guiding, hospitality press, frenchs forest. pearce, p. & moscardo, g., 1998, ‘the role of interpretation in influencing visitor satisfaction: a rainforest case study’, in w. faulkner, c. tidswell, & d. weaver, (eds.), progress in tourism and hospitality research, part 1, proceedings of the 8th australian tourism and hospitality research conference, gold coast, 11–14 february, pp. 309–319, bureau of tourism research, canberra. phillips, a., 1989, ‘interpreting the countryside and the natural environment’, in d. uzzell (ed.) heritage interpretation – the natural and built environment, pp. 121–131, belhaven, london. regnier, k., gross, m. & zimmerman, r., 1994, techniques for programs and presentations, 3rd edn., uw-sp foundation press, stevens point. pmcid:pmc395234 russell, c.l. & hodson, d., 2002, ‘whale watching as critical science education?’, canadian journal of science, mathematics and technology education 2(4), 485–504. http://dx.doi.org/10.1080/14926150209556537 ryan, c., hughes, k.c. & hirgwin, s., 2000, ‘the gaze, spectacle and ecotourism’, annals of tourism research 27(1), 148–163. http://dx.doi.org/10.1016/s0160-7383(99)00061-4 schänzel, h.a. & mcintosh, a.j., 2000, ‘an insight into the personal and emotive context of wildlife viewing at the penguin place, otago peninsula, new zealand’, journal of sustainable tourism 8(1), 36–52. http://dx.doi.org/10.1080/09669580008667348 shackley, m., 1998, visitor management – case studies from world heritage sites, butterworth-heinemann, oxford. pmcid:pmc1364099 sharpe, g. (ed.), 1982, an overview of interpretation, 2nd edn., john wiley & sons, new york. stewart, e.j., hayward, b.m. & devlin, p.j., 1998, ‘the ”place” of interpretation: a new approach to the evolution of interpretation’, tourism management 19(3), 257–266. http://dx.doi.org/10.1016/s0261-5177(98)00015-6 tilden, j., 1977, interpreting our heritage: principles and practices for visitors in parks, museums and historic places, 3rd edn., university of north carolina press, chapel hill. pmid:832384 tisdell, c. & wilson, c., 2001, ‘wildlife-based tourism and increased support for nature conservation financially and otherwise: evidence from sea turtle ecotourism at mon repos’, tourism economics 7(3), 233–249. http://dx.doi.org/10.5367/000000001101297847 townsend, c., 2003, ‘marine ecotourism through education: a case study of divers in the british virgin islands’, in b. garrod & j.c. wilson (eds.), marine ecotourism: issues and experiences, pp. 138–154, channel view publications, clevedon. tubb, k., 2003, ‘an evaluation of the effectiveness of interpretation within dartmoor national park in researching the goals of sustainable tourism development’, journal of sustainable tourism 11(6), 476–498. http://dx.doi.org/10.1080/09669580308667217 uzzell, d., 1998, ‘strategic considerations and practical approaches to the evaluation of heritage interpretation’, in d. uzzell & r. ballantyne (eds.), contemporary issues in heritage and environmental interpretation: problems and prospects, pp. 185–202, stationery office, london. van der merwe, p. & saayman, m., 2008, ‘travel motivations of tourists visiting the kruger national park’, koedoe 50(1), 154–159. http://dx.doi.org/10.4102/koedoe.v50i1.140 vaske, j.j., donnelly, m.p. & whittaker, d., 2000, ‘tourism, national parks and impact management’, in r.w. butler & s.w. boyd (eds.), tourism and national parks, pp. 203–222, wiley, chichester. ward, c. & wilkinson, a., 2006, personal interpretation – a field guide for success, fulcrum books, golden. wearing, s., archer, d., moscardo, g. & schweinsberg, s., 2007, best practice interpretation research for sustainable tourism: framework for a new research agenda, sustainable tourism crc, queensland. weaver, d., 2000, ‘tourism and national parks in ecologically vulnerable areas’, in r.w. butler & s.w. boyd (eds.), tourism and national parks, pp. 107–124, wiley, chichester. pmid:11011800 weiler, b. & smith, l., 2009, ‘does interpretation lead to greater outcomes? an assessment of the impacts of multiple layers of interpretation in a zoo context’, journal of sustainable tourism 17(1), 91–105. http://dx.doi.org/10.1080/09669580802359319 world tourism organisation, 1992, guidelines: development of national parks and protected areas for tourism, wto, madrid. world tourism organisation, 1995, what tourism managers need to know: a practical guide to the development and use of indicators of sustainable tourism, wto, madrid. abstract introduction research method and design results discussion conclusion acknowledgements references appendix 1: methods and materials used for soil analyses appendix 2: braun–blanquet scale for vegetation cover appendix 3: ant species lists for various data sets appendix 4: tables illustrating sampling completeness calculated for the rehabilitated and control sites appendix 5: list of indicator values about the author(s) samantha-leigh jamison department of plant science, university of pretoria, south africa mark robertson centre for invasion biology, department or zoology & entomology, university of pretoria, south africa ian engelbrecht department of zoology and entomology, university of pretoria, south africa peter hawkes department of zoology and entomology, university of pretoria, south africa afribugs cc, pretoria, south africa citation jamison, s-l., robertson, m., engelbrecht, i. & hawkes, p., 2016, ‘an assessment of rehabilitation success in an african grassland using ants as bioindicators’, koedoe 58(1), a1383. http://dx.doi.org/10.4102/koedoe.v58i1.1383 original research an assessment of rehabilitation success in an african grassland using ants as bioindicators samantha-leigh jamison, mark robertson, ian engelbrecht, peter hawkes received: 02 feb. 2016; accepted: 23 june 2016; published: 29 sept. 2016 copyright: © 2016. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract many studies that evaluate rehabilitation make use of invertebrate bioindicators. invertebrates, especially ants, make useful indicators as they are sensitive to environmental change. we compared ant assemblages in rehabilitated and control sites in the rietvlei nature reserve, a protected area important for grassland conservation in south africa. pitfall traps were used to sample ant assemblages at six control sites and six rehabilitated sites. in addition, environmental and vegetation surveys were conducted at each site. we found that the ant assemblages differed significantly between the control and rehabilitated sites, although there was considerable overlap; the control sites supported a greater species density and higher abundance of ants than the rehabilitated sites. in total, 36 ant species were collected (control sites: 34 species; rehabilitated sites: 26 species). the environmental survey revealed that percentages of bare ground and coarse sand, as well as soil ph, differed significantly between the control and rehabilitated sites. the control and rehabilitated sites also supported significantly different plant assemblages. three indicator ant species were identified for the control sites: crematogaster rectinota, crematogaster amita and monomorium fastidium. no indicator species were identified for the rehabilitated sites. these results suggest that recovery from the previous agricultural use of the area is still incomplete and highlights the lack of research examining the success of rehabilitation in the grassland biome. conservation implications: the present study illustrates the need for further research on rehabilitation techniques utilised in the grassland biome. this is of value as the remainder of south african grasslands are considered critically endangered. introduction rehabilitation and the need for bioindicators increasing human impacts on the environment have resulted in widespread degradation of ecosystems (hobbs & norton 1996). as a result, there is a global drive to rehabilitate terrestrial and freshwater environments (lake 2001). typically, rehabilitation projects monitor aspects such as vegetation structure, species diversity and richness as well as nutrient cycling (ruiz-jaén & aide 2005; van aarde & smit 1997). a number of rehabilitation studies make use of biological indicators. these indicators provide a means to monitor and assess the state of an ecosystem over time (cairns, mccormick & niederlehner 1993; morellet et al. 2007). however, in order to be practical, these bioindicators need to meet certain criteria. they need to be sensitive enough to provide an early warning in the event of a disturbance, they should have a wide spatial distribution and they must provide a continuous assessment for several disturbance factors (noss 1990). such indicators can aid us in understanding the rehabilitation of previously disturbed land and prove valuable for future management projects. ants as bioindicators invertebrates are potentially valuable indicators for monitoring the success of rehabilitation (andersen & sparling 1997). in particular, ants (hymenoptera: formicidae) have received much attention as ecological bioindicators. they are widespread, abundant, have a manageable species richness and perform many keystone functions and thus have the potential to be used for the assessment of rehabilitation (fagan et al. 2010; van hamburg et al. 2004). they are important ecosystem engineers that are responsible for turning soil, seed dispersal, forming mutualistic relationships and are among the top predators of other invertebrates (hölldobler & wilson 1990; lach, parr & abott 2010). as they are colonial, their response to disturbance may be very different from other invertebrates that are more susceptible to disturbance events (lach et al. 2010). the removal of individuals, resulting from mortality caused by disturbance, may not lead to the eradication of the colony from the habitat. thus, the response of ants to disturbance differs from that of other terrestrial plants and animals that may become locally extinct after a disturbance event (andersen 2000; lach et al. 2010). in response to disturbance, such as agriculture or mining activities, ant communities may show changes in species composition and interspecific interactions, as well as loss of taxonomic diversity and variation in the provision of ecosystem services (lach et al. 2010). ants have been used as bioindicators in several projects involving ecosystem management, land rehabilitation and monitoring the degree of land degradation, as well as investigating the conservation status of various zones of land (majer 1983). they have been successfully used as bioindicators of mine site rehabilitation in northern australia (andersen, hoffman & somes 2003) and on rehabilitated ash dams associated with coal-fired power stations around the highveld of south africa (van hamburg et al. 2004). they have also been used to indicate pesticide contamination in cotton-growing areas in australia (weir 1978). ants thus provide an important indication of the biotic and abiotic state of an ecosystem and ultimately reveal the impacts that various environmental changes can have on an ecosystem (hodkinson & jackons 2005). the importance of grasslands grasslands are regions of global importance and provide a series of important ecosystem services. these include the purification of water, flood attenuation and nutrient cycling, as well as carbon sequestration and storage (south african national biodiversity institute [sanbi] 2013). the south african grassland biome covers nearly 30% of the country’s land surface and is home to a large majority of the country’s population and several endangered and endemic species (neke & du plessis 2004; sanbi 2013). increasing urbanisation and development has resulted in a substantial amount of land degradation in the south african grassland biome, resulting in its classification as critically endangered (neke & du plessis 2004). despite this, grasslands throughout the country have received little conservation protection (neke & du plessis 2004). despite the necessity to rehabilitate grasslands following disturbances such as mining or agriculture, there appears to be limited knowledge on grassland rehabilitation (zaloumis & bond 2010). as rehabilitation may take many years, there is a need to quantify the extent to which ecosystems (such as grasslands) have been rehabilitated (legg & nagy 2006). bioindicators such as ants can fulfil this purpose. however, to date surprisingly little research has been done in south africa on ant diversity in grasslands and how ant communities resemble each other following a disturbance. this study investigated the success of the rehabilitation measures applied in an old agricultural land area on rietvlei nature reserve. we compared ant assemblages in areas where rehabilitation measures were applied and in nearby untransformed, natural grassland areas as an indication of the effectiveness of rehabilitation. we also aimed to identify whether any indicator species were present in the rehabilitating and untransformed (control) sites of the reserve. research method and design study area the study took place on the southern region of rietvlei nature reserve (between 25°50’s, 28°15’e and 25°56’s, 28°19’e), a protected area important for grassland conservation in south africa. the reserve covers an area of approximately 3870 ha. it is situated in close proximity to urban centres. the climate of the area is characterised by warm, wet summers that are followed by cold, dry winters, when frost is prevalent. summer temperatures may reach highs of 34 °c, which are contrasted by the low winter temperatures that can fall to –2 °c (marais 2004). the area has a mean annual rainfall of approximately 724 mm (1970–1999; marais 2004). the plant composition of the reserve is typical of the grassland biome; the vegetation type of the area is described as rand highveld grassland (mucina & rutherford 2006). around 23 years ago, the reserve extended its southern boundary and incorporated land that was previously used for agriculture. rehabilitation measures were employed on the land in an attempt to improve species diversity and ecological function. the rehabilitated areas were ploughed and sown with a mix of indigenous grass seeds (r. marais 2014, pers. comm., rietvlei nature reserve, pretoria). procedure ant sampling sampling was carried out at 12 sites, 6 located in the rehabilitated area and 6 in untransformed (control) areas of the reserve. all sites were at a minimum distance of 200 m from old land boundaries to avoid edge effects and 300 m from other sampling sites to ensure independent sampling. at each site, 10 plastic pitfall traps (55 mm in diameter and 70 mm deep) were arranged in two rows of five, positioned 10 m apart. the study was designed in such a manner so as to avoid or minimise the possibility of pseudoreplication that may have been introduced by having a single larger array with many traps and treating each array (rather than each trap) as a replicate. the traps were placed in the soil with the rim flush with the soil surface and partially filled with a mixture of water and propylene glycol. this non-toxic solution ensures the successful preservation of the captured specimens. a rain cover (20-cm plastic lid with four wire legs) was placed above each trap to avoid flooding. the traps were left in place for 3 days from 08 to 11 april 2014. the ant specimens found in each trap were removed and identified to species level where possible and their abundances recorded. ants were identified to genus using bolton (1994) and then to species where possible, using the most recent available keys to the genera concerned: for agraulomyrmex, prins (1983); anoplolepis, prins (1982); crematogaster, arnold (1920); lepisiota, arnold (1920); leptogenys, bolton (1975); meranoplus, bolton (1981); monomorium, bolton (1987); technomyrmex, bolton (2007) and tetramorium, bolton (1980). for some genera, such as camponotus, carebara, dorylus, pheidole, plagiolepis and solenopsis, no keys are presently available for the afrotropical fauna; for these genera, as well as for species in other genera that could not be identified using the available keys, specimens were identified by comparison with reference material in the afribugs collection (afrc). where no formal name could be assigned, a morphospecies code matched to voucher material in the afrc collection was applied. these codes are globally unique and images of most are available on antweb (http://www.antweb.org) to allow cross-referencing between studies. all identifications were performed by s.l. jamison under the guidance of p.g. hawkes. a specimen of each of the species identified was mounted and labelled to produce a reference collection, which is housed at the department of zoology & entomology, university of pretoria. environmental factors estimates of the percentage bare ground, rock, grass, shrub cover and height of the tallest plant were recorded in five 1-m2 quadrats at each site. these quadrats were randomly selected from the immediate area around each of the sites. the quadrats were placed within a 10-m perimeter around the pitfall traps in order to establish the best estimate of environmental factors within each of the respective sites. the braun–blanquet cover-abundance scale was used to quantify the cover of all plant species in three 1-m2 quadrats at each site. a disk pasture meter was used to obtain measures of above-ground vegetation biomass. a total of 20 disk pasture meter height measurements were recorded at random at each site. we refer to these height measurements as a biomass index from here onwards. a soil auger was used to collect a soil sample from the first 20 cm of soil at each of the sites. soil was randomly sampled within a 10-m perimeter around the pitfall traps. three soil samples were randomly taken from each of the 12 sites. the three soil samples collected at each site were then combined and used as one sample to get an indication of the average soil conditions at each site. the percentage coarse sand, sand, silt and clay content of the soil as well as the soil ph and soil organic matter were determined using standard methods (appendix 1). all soil analyses were conducted at the university of pretoria in the soil sciences department. analysis species diversity indices for the ant data were generated using primer 5.2.0 software package (clarke & warwick 2001). the indices included the pielou (pielou 1969) and shannon–weiner diversity indices (shannon 2001). these diversity indices were compared between the rehabilitated and control sites with welch two-sample t-tests (welch 1947). furthermore, welch two-sample t-tests were used to test for significant differences between the number of ant species and the number of individuals found between rehabilitated and control sites. species richness in the two sites was compared using rarefaction curves with 95% confidence intervals derived from unconditional variance estimates (colwell et al. 2012). richness estimators were generated using estimate-s (v 9.1.0; colwell 2013). we used three non-parametric species richness estimators and one parametric estimator in our analysis to correct for sampling bias. the non-parametric estimators were the incidence-based coverage estimator (ice), the chao 2 abundance-based richness estimator (chao 2) and the second-order jackknife richness estimator (jack 2). we used the michaelis–menton mean (mmmean) as the parametric estimator. we chose this suite of estimators as estimator performance varies depending on a range of factors, and no single estimator is universally accepted as superior to others (smith & jones 2005). sampling was considered adequate if the sample-based rarefaction curves and the curves of the species richness estimators converged closely at the highest observed values (cardoso 2009; cardoso et al. 2009; longino, coddington & colwell 2002; magurran 2004). observed richness as a percentage of estimated richness was used as a measure of inventory completeness (jiménez-valverde et al. 2010). the mean sampling completeness was calculated for the control and rehabilitated sites by averaging the completeness calculated for each of the different species richness estimators. species richness was compared between the rehabilitated and control sites by plotting rarefaction curves with their 95% confidence intervals. if the intervals overlapped, the differences between the treatments were taken to be not significant (colwell et al. 2012). an analysis of similarity (anosim) and non-metric multidimensional scaling (nmds) were performed to compare the ant assemblages in the rehabilitated and control areas. the similarity matrix was calculated using a bray–curtis similarity measure and a fourth-root transformation in primer 5.2.0 (clarke & warwick 2001). the fourth-root transformation reduces the contribution of very abundant species (french & major 2001). anosim generates a global r statistic that provides an indication of average dissimilarity between the assemblages being compared. values closer to 1 indicate distinct differences, while values closer to 0 indicate high levels of similarity in species composition. to assess the contribution of different species to the differences between the rehabilitated and control sites, the similarity percentages for species contributions were applied to the data (simper, primer 5.2.0). a fourth-root transformation was again used to reduce the influence of dominant species. characteristic ant species (indicator species) were identified for each of the sites with the use of the indicator values method (dufrêne & legendre 1997) in the package labdsv (roberts 2014), run in r 3.1.1 (r development core team 2011). an analysis of the relative indicator values (indval) was performed to determine the specificity (uniqueness to specific sites) and fidelity (rate of recurrence within each site) of each species for a particular site and treatment (e.g. rehabilitating or control). species that were found to have a value greater than 70%, that is, species that occur predominantly in a particular habitat and occur frequently in that habitat were considered as reliable indicator species for the different sampling units. rank abundance curves were also generated and allowed comparison of species composition between control and rehabilitated sites. the values for percentage of bare ground, rock, grass, shrub and height of tallest plant were compared between control and rehabilitated sites using welch’s two-sample t-tests, which were also used to compare the percentages of coarse sand, sand, silt and clay, as well as soil ph and soil organic matter. anosim and nmds were performed to compare the plant assemblages in the rehabilitated and control sites. the similarity matrix was calculated using a bray–curtis similarity measure and a fourth-root transformation in primer 5.2.0 (clarke & warwick 2001). the braun–blanquet scale was rescaled to range from 1 to 7 for the purpose of the anosim and nmds (table 1-a2). results ants in total, 3206 individuals representing 36 species were collected during the study (table 1-a3). the control sites had a significantly higher species density (t = 6.618, p < 0.001) and abundance (t = 4.674, p < 0.001) than the rehabilitated sites (figure 1). additionally, the control sites had a significantly higher pielou evenness index (t = 3.125, p < 0.05) and shannon–weiner index (t = 6.789, p < 0.001) than the rehabilitated sites (figure 1). in total, 34 ant species were collected from the control sites, whereas only 26 species were collected from the rehabilitated sites. of the 36 species collected in this study, 24 species (66.7%) were present in both the control and rehabilitated sites. in total, 10 species (27.8%) were unique to the control sites, whereas only 2 species (5.6%) were unique to the rehabilitated sites (table 1-a3). figure 1: box plots of ant diversity in the control and rehabilitated sites on rietvlei nature reserve, showing (a) the number of species, (b) the number of individuals, (c) the pielou evenness (j’) and (d) the shannon–weiner index (h’). the estimated species richness (sest) for the ant samples collected from the control and rehabilitated sites did not converge closely with the richness estimators (figure 1-a1). the mean sampling completeness for the rehabilitated sites was found to be 65.70%, whereas the average sampling completeness for the control sites was 70.31% (from table 1-a4 to table 5-a4). however, as it is unlikely that we will ever reach an asymptotic endpoint, particularly for ants and other invertebrates (gotelli et al. 2014), this level of sampling completeness is likely sufficient to draw conclusions about grassland rehabilitation. the confidence intervals on the rarefaction curves for the rehabilitated and control sites overlapped, indicating that there were no significant differences in species richness (figure 2). figure 2: rarefaction curves with 95% confidence intervals comparing species richness in the control and rehabilitated (rehab) sites. species richness should be compared for equal numbers of species in each treatment. anosim indicated that ant assemblages differed significantly between the treatments (global r = 0.507, p = 0.002). the nmds plot revealed that there was some separation between the control and rehabilitated sites, although some overlap is still evident (figure 3). the replicates representing the control sites (c) are clustered together at the top of the plot indicating smaller assemblage differences between these replicates than for the rehabilitated sites. the replicates representing four of the rehabilitated sites (1r, 3r, 5r and 6r) are clustered together in the lower right quadrant of the plot. they are clustered less tightly than the control sites at the top of the plot. this indicates fairly large assemblage differences between the replicates within the rehabilitated sites compared with those of the control sites. in particular, sites 1r and 6r are plotted in close proximity to the control sites indicating that the ant assemblages at these rehabilitated sites are similar to those of the control sites. sites 2r and 4r are isolated from the clusters on either side of the plot indicating that the ant assemblages found at these sites are different from those found at the other sites of both treatment types. it is interesting that 2c is located near the rehabilitated sites within the nmds plot. the potential impact of human-mediated activities could have influenced the habitat at this site. however, we have no information regarding the use of the area before the establishment of the reserve and cannot give a fully informed explanation. figure 3: non-metric multidimensional scaling plot indicating the similarity of ant assemblages among replicates of control and rehabilitated sites. the assemblage differences are further supported by the rank abundance curves observed for the different treatments (figure 4). these rank abundance curves show only species that were represented by more than five individuals. the three most abundant species found in the control sites were monomorium albopilosum emery (641 individuals), meranoplus peringueyi emery (209 individuals) and tetramorium bothae forel (193 individuals). in comparison, the three most abundant species found in the rehabilitated sites included m. albopilosum (411 individuals), tetramorium setuliferum emery (278 individuals) and pheidole megacephala fabricius (177 individuals). although many of the highly abundant species were present in both the rehabilitated and control sites, the control sites had more unique species than the rehabilitated sites (control: 10 unique species, rehabilitated: 2 unique species; table 1-a3). the criteria needed to identify indicator species (indval > 70%) were fulfilled for three species present in the control sites (table 1-a5). these species include crematogaster rectinota forel (indval = 0.932, p = 0.004), crematogaster amita forel (indval = 1, p = 0.003) and monomorium fastidium bolton (indval = 0.808, p = 0.031). no indicator species were identified for the rehabilitated sites. the results from the rank abundance curves are further supported by the simper results, which found c. amita to have the greatest contribution to the separation of the control and rehabilitated sites (6.31% of total abundance; 51 individuals in total for the control sites), followed by m. peringueyi (5.67%; 209 individuals in control sites) and lastly m. fastidium (5.53%; 167 individuals across control and rehabilitated sites). although m. albopilosum had a total of 1052 individuals for both the control and rehabilitated sites, the overall contribution of this species to the separation of sites was only 4.27%. the contribution of the other species was greater because of the difference in average abundance of the species between the sites. the mean abundance of m. albopilosum was 106.83 in the control sites and 68.50 in the rehabilitated sites. figure 4: rank abundance plot for the rehabilitated and control grassland sites on rietvlei nature reserve. environmental variables the mean percentage of bare ground was significantly higher in the rehabilitated than the control sites (t = -3.193, p = 0.003; figure 1-a2). there were no significant differences in the percentage cover of grass, rock, height of tallest plant and biomass index between the rehabilitated and control sites. the ph of the soil (t = -2.754, p = 0.021) and the percentage coarse sand (t = 2.275, p = 0.0495) differed significantly between the rehabilitated and control sites (figure 1-a3). however, the t-tests revealed that there was no significant difference in soil organic matter and percentage of sand, silt and clay content between the rehabilitated and control sites. vegetation assemblages anosim indicated that the plant assemblages differed significantly between the treatments (global r = 0.712, p = 0.001). the global r statistic indicates fairly large assemblage differences between the control and rehabilitated sites. the nmds plot illustrates that there is a clear distinction between the control and rehabilitated sites (figure 1-a4). there is a larger amount of variation in the plant communities between the various control sites than there is between the rehabilitated sites. it can be seen that the plant community of the control and rehabilitated sites is still very different, 23 years after rehabilitation. discussion this study found that the control sites on the rietvlei nature reserve had significantly higher ant species density and abundance than the rehabilitated sites. in total, 34 ant species were collected from the control sites, whereas only 26 species were collected from the rehabilitated sites. moreover, the control sites had a significantly higher pielou evenness index and shannon–weiner diversity index than the rehabilitated sites. however, species richness was not significantly different between control and rehabilitated sites. there was a significant difference in the ant assemblages between the rehabilitated and control sites, although some convergence was evident. influence of dominant ant species of the 36 ant species collected in this study, 66.7% were present in both the control and rehabilitated sites (i.e. 24 species shared). however, a large portion of the difference between ant assemblages of the rehabilitated and control sites was because of the higher abundance of individuals present in the control sites (control: 1954 individuals, rehabilitated: 1252 individuals). the difference between ant assemblages may thus be because of a combination of the abundance of individuals and the composition of species present in the control and rehabilitated sites. in particular, two ant species (t. setuliferum and p. megacephala) had higher abundances in the rehabilitated sites than in the control sites, though neither species was identified as an indicator. pheidole megacephala and several species of tetramorium and monomorium are said to be characteristic of disturbed areas (andersen 2003; samways, caldwell & osborn 1996). furthermore, dominant ant species are known to affect the ant assemblages present in a region as well as influence species coexistence (majer, delabie & smith 1994; samways et al. 1996). samways et al. (1996) attributed the lower abundance of species recorded in the disturbed sites in their study to the presence of p. megacephala. consequently, the high abundance of an aggressive species such as p. megacephala may explain the lower abundance of other ant species recorded in the rehabilitated sites in this study. environmental variables many of the environmental variables investigated in this study did not differ between the control and rehabilitated sites. the sites were similar in terms of total grass cover, other cover, rock cover, height of tallest plant and several soil properties (e.g. percentage sand, silt and clay content as well as soil organic matter). as a result, it could be said that the old agricultural land on the reserve has started to recover. despite this, the percentage bare ground, soil ph and the percentage coarse sand differed significantly between the rehabilitated and control sites. this is important as such factors could influence the ant assemblages present in the sites. for instance, soil type is known to have an effect on the ant species present in a specific area (andersen 1993; koen & breytenbach 1988; lindsey & skinner 2001). unfortunately, the exact habitat requirements of ants are not well known (lindsey & skinner 2001). however, the little research that has been done on this topic has revealed that ant assemblages are influenced by a number of habitat variables; soil moisture, soil type, the geology of an area, plant structural complexity and leaf litter cover (andersen 1993; koen & breytenbach 1988; lindsey & skinner 2001). many rehabilitation studies that have compared ant assemblages on rehabilitated and reference sites have established that the vegetation between the two compared treatments is very different. for example, van hamburg et al. (2004) noted that the vegetation composition on rehabilitated ash dams and nearby natural grasslands was distinct. consequently, these differences in plant composition result in different ant species assemblages as well as differences in ant species diversity (van hamburg et al. 2004). also, a study by zaloumis and bond (2010) found differences between rehabilitated and control sites in a south african grassland; rehabilitated sites supported a much smaller community of resprouting forb species than the control sites. this is important as forb species make up much of the diversity of south african grasslands. the poor ability of forb species to reestablish after disturbance is attributed to their poor dispersal ability and lack of propagating sources (dickson & busby 2008; kardol et al. 2008; zaloumis & bond 2010). several environmental factors such as soil properties can alter the direction of succession as well as the plant species present (zaloumis & bond 2010). also, the addition of nutrients such as nitrogen from fertilisers can influence the spread and species richness of weeds and alien plants, which has negative consequences for the native flora of the region (zaloumis & bond 2010). thus, a thorough understanding of the region’s environmental variables and their impact on plant species is of importance, as ants are dependent on vegetation structure (andersen 1995). the failure of successful rehabilitation of the plant community in grasslands could consequentially affect the ant assemblages present within the area. this is important as the present study found a significant difference between the plant assemblages of the control and rehabilitated sites on rietvlei nature reserve. this difference in plant assemblages could possibly explain the difference in ant assemblages on the reserve because of factors such as substrate and vegetation have direct effects on the colonisation ability of different ant species (van hamburg et al. 2004). as a result, the differences in plant assemblages and specific environmental factors may explain the variation between ant assemblages of control and rehabilitated sites. similarly, as ants are considered ecosystem engineers and often cause changes to soil, they too could be a contributing factor affecting the plant species composition within the area. as a result, ants could be driving much of the rehabilitation in the reserve old lands; however, more research is required before a conclusive decision can be made. indicator species three indicator species were identified, all of which were characteristic of the control sites. the indicator species were c. rectinota, c. amita and m. fastidium. no indicator species were identified for the rehabilitated sites. specialist ant predators are said to be highly sensitive to disturbance and thus are rarely recorded in disturbed habitats (andersen et al. 2003; hoffmann & andersen 2003). included in the list of specialist predators (andersen 2000) is the genus leptogenys, in the tribe ponerini. only one species of this genus, leptogenys intermedia emery, was recorded during this study. however, the fact that this species was recorded in both the rehabilitated and control sites suggests that the rehabilitated sites have started acquiring the vegetation structure, compostition and arthropod fauna that is necessary to support the specialised diet of these predators (andersen 2000). this is noteworthy as many authors have found specialist predators to be among the last species to colonise rehabilitating areas (majer & beeston 1996; hoffman & andersen 2003). this is interesting as not much is known regarding the potential effects of these species on their environment and thus requires further research. conclusion our results show that the ant assemblages on rehabilitated areas of the reserve are significantly different from those of surrounding undisturbed areas, although some convergence is evident. these differences are likely to be explained by a combination of factors, including the presence of certain dominant ant species, differences in plant species composition and differences in environmental factors (including percentage bare ground, percentage coarse sand and soil ph). a conclusive statement regarding the success of rehabilitation efforts on the reserve is not possible given the lack of background information on recovery rates and compositional changes, but our study shows that rehabilitation of grasslands would benefit from a greater understanding of ant diversity and the factors that are responsible for driving this diversity. further studies are needed to investigate the functional roles of particular ant species, especially those that are considered to be good indicator species. acknowledgements we are grateful to everyone who assisted with this project; chantal ferreira and jonathan fisher are thanked for their assistance with the ant identifications. our gratitude is also extended to the rietvlei nature reserve and its staff members, especially riaan marais, who made this study possible. furthermore, wesley daniels, chantal ferreira, michael potgieter, andré van tonder, cavin shivambu and paul akkermans are gratefully acknowledged for their assistance in the field. our thanks are extended to the soil sciences department of the university of pretoria who allowed us to use their facilities when conducting the soil analyses. we would also like to thank the anonymous reviewers whose comments helped to improve the manuscript. we are grateful for funding received from the national research foundation. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions s-l.j., m.r., i.e. and p.h. conceived and designed the study. s-l.j., m.r. and i.e. performed fieldwork. s-l.j. and p.h. performed ant identifications. s-l.j., i.e. and m.r. performed statistical analyses; p.h. contributed identification tools. s-l.j., m.r., i.e. and p.h. wrote and edited the manuscript. references andersen, a., 1993, ‘ants as indicators of restoration success at a uranium mine in tropical australia’, restoration ecology 1(3), 156–167. http://dx.doi.org/10.1111/j.1526-100x.1993.tb00022.x andersen, a., 1995, ‘a classification of australian ant communities based on functional groups which parallel plant life-forms in relation to stress and disturbance’, journal of biogeography 22(1), 15–29. http://dx.doi.org/10.2307/2846070 andersen, a., 2000, ‘a global ecology of rainforest ants: functional groups in relation to environmental stress and disturbance’, in d. agosti, j.d. majer, l.e. alonso & t.r. schultz (eds.), ants: standard methods for measuring and monitoring biodiversity, pp. 25–34, smithsonian institution press, washington, dc. andersen, a., 2003, ‘ant biodiversity in arid australia: productivity, species richness and community organisation’, records of the south australian museum monograph series 7, 79–92. andersen, a., hoffmann, b. & somes, j., 2003, ‘ants as indicators of minesite restoration: community recovery at one of eight rehabilitation sites in central queensland’, ecological management and restoration 4(1), 12–19. http://dx.doi.org/10.1046/j.1442-8903.4.s.2.x andersen, a. & sparling, g., 1997, ‘ants as indicators of restoration success: relationship with soil microbial biomass in the australian seasonal tropics’, restoration ecology 5(2), 109–114. http://dx.doi.org/10.1046/j.1526-100x.1997.09713.x arnold, g., 1920, ‘a monograph of the formicidae of south africa, myrmicinae’, annuals of the south african museum 14(5), 403–578. bolton, b., 1975, ‘a revision of the ant genus leptogenys roger (hymenoptera: formicidae) in the ethiopian region with a review of the malagasy species’, bulletin of the british museum (natural history) entomology 31(7), 235–305. http://dx.doi.org/10.5962/bhl.part.29487 bolton, b., 1980, ‘the ant tribe tetramoriini (hymenoptera: formicidae). the genus tetramorium mayr in the ethiopian zoogeographical region’, bulletin of the british museum (natural history) entomology 40(3), 193–384. bolton, b., 1981, ‘a revision of the ant genera meranoplus f. smith, dicroaspis emery and calyptomyrmex emery (hymenoptera: formicidae) in the ethiopian zoogeographical region’, bulletin of the british museum (natural history) entomology 42(2), 43–81. bolton, b., 1987, ‘a review of the solenopsis genus-group and revision of afrotropical monomorium mayr (hymenoptera: formicidae)’, bulletin of the british museum (natural history) entomology 54(3), 263–452. bolton, b., 1994, identification guide to the ant genera of the world, harvard university press, cambridge. bolton, b., 2007, ‘taxonomy of the dolichoderine ant genus technomyrmex mayr (hymenoptera: formicidae) based on the worker caste’, contributions of the american entomological institute 35(1), 1–150. cairns, j.r.j., mccormick, p.v. & niederlehner, b.r., 1993, ‘a proposed framework for developing indicators of ecosystem health’, hydrobiologia 263(1), 1–44. http://dx.doi.org/10.1007/bf00006084 cardoso, p., 2009, ‘standardization and optimization of arthropod inventories — the case of iberian spiders’, biodiversity and conservation 18(14), 3949–3962. http://dx.doi.org/10.1007/s10531-009-9690-7 cardoso, p., crespo, l.c., carvalho, r., rufino, a.c. & henriques, s.s., 2009, ‘ad-hoc vs. standardized and optimized arthropod diversity sampling’, diversity 1(1), 36–51. http://dx.doi.org/10.3390/d1010036 clarke, k.r. & warwick, r.m., 2001, change in marine communities: an approach to statistical analysis and interpretation, 2nd edn., primer-e, plymouth. colwell, r.k., 2013, ‘estimates: statistical estimation of species richness and shared species from samples’, version 9.1.0, viewed may 2014, from http://www.purl.oclc.org/estimates colwell, r.k., chao, a., gotelli. n.j., lin, s.y., mao, c.x., chazdon, r.l. et al., 2012, ‘models and estimators linking individual-based and sample-based rarefaction, extrapolation, and comparison of assemblages’, journal of plant ecology 5(1), 3–21. http://dx.doi.org/10.1093/jpe/rtr044 dickson, t.l. & busby, w.h., 2008, ‘forb species establishment increases with decreased grass seeding density and with increased forb seeding density in a northeast kansas, u.s.a., experimental prairie restoration’, restoration ecology 17(5), 597–605. http://dx.doi.org/10.1111/j.1526-100x.2008.00427.x dufrêne, m. & legendre, p., 1997, ‘species assemblages and indicator species: the need for a flexible asymmetrical approach’, ecological monographs 67(3), 345–366. http://dx.doi.org/10.2307/2963459 fagan, k.c., pywell, r.f., bullock, j.m. & marrs, r.h., 2010, ‘are ants useful indicators of restoration success in temperate grasslands?’, restoration ecology 18(3), 373–379. http://dx.doi.org/10.1111/j.1526-100x.2008.00452.x french, k. & major, r.e., 2001, ‘effect of an exotic acacia (fabaceae) on ant assemblages in south african fynbos’, austral ecology 26(4), 303–310. http://dx.doi.org/10.1046/j.1442-9993.2001.01115.x gotelli, n.j., ellison, j.a., dunn, r.r. & sanders, n.j., 2014, ‘counting ants (hymenoptera: formicidae): biodiversity sampling and statistical analysis for myrmecologists’, myrmecological news 15, 13–19. hobbs, r.j. & norton, d.a., 1996, ‘towards a conceptual framework for restoration ecology’, restoration ecology 4(2), 93–110. http://dx.doi.org/10.1111/j.1526-100x.1996.tb00112.x hodkinson, i.d. & jackson, j.k., 2005 ‘terrestrial and aquatic invertebrates as bioindicators of environmental monitoring with particular reference to mountain ecosystems’, environmental management 35(5), 649–666. http://dx.doi.org/10.1007/s00267-004-0211-x hoffmann, b.d. & andersen, a.n., 2003, ‘responses of ants to disturbance in australia, with particular reference to functional groups’, austral ecology 28(4), 444–464. http://dx.doi.org/10.1046/j.1442-9993.2003.01301.x hölldobler, b. & wilson, e.o., 1990, the ants, springer-verlag, cambridge. howard, s., 1998, ‘verbal protocol analysis’, in b. henderson-sellers, a. simons & h. younessi (eds.), the open process specification, pp. 272–274, addison wesley, sydney. jiménez-valverde, a., baselga, a., melic, a. & txasko, n., 2010, ‘climate and regional beta-diversity gradients in spiders: dispersal capacity has nothing to say?’, insect conservation and diversity 3(1), 51–60. http://dx.doi.org/10.1111/j.1752-4598.2009.00067.x kardol, p., wal, a.v.d., bezemer, t.m., boer, w.d., duyts, h., holtkamp, r. et al., 2008, ‘restoration of species-rich grasslands on ex-arable land: seed addition outweighs soil fertility reduction’, biological conservation 141(9), 2208–2217. http://dx.doi.org/10.1016/j.biocon.2008.06.011 koen, j.h. & breytenbach, w., 1988, ‘ant species richness of fynbos and forest ecosystems in the southern cape’, south african journal of zoology 23(3), 184–188. http://dx.doi.org/10.1080/02541858.1988.11448100 lach, l., parr, c.l. & abott, k.l., 2010, ant ecology, oxford university press, oxford. lake, p.s., 2001, ‘on the maturing of restoration: linking ecological research and restoration’, ecological management and restoration 2(2), 110–115. http://dx.doi.org/10.1046/j.1442-8903.2001.00074.x legg, c.j. & nagy, l., 2006, ‘why most conservation monitoring is, but need not be, a waste of time’, journal of environmental management 78(2), 194–199. http://dx.doi.org/10.1016/j.jenvman.2005.04.016 lindsey, p.a. & skinner, j.d., 2001, ‘ant composition and activity patterns as determined by pitfall trapping and other methods in three habitats in the semi-arid karoo’, journal of arid environments 48(4), 551–568. http://dx.doi.org/10.1006/jare.2000.0764 longino, j.t., coddington, j. & colwell, r.k., 2002, ‘the ant fauna of a tropical rain forest: estimating species richness three different ways’, ecology 83(3), 689–702. http://dx.doi.org/10.1890/0012-9658(2002)083[0689:tafoat]2.0.co;2 magurran, a.e., 2004, measuring biological diversity, blackwell, oxford. majer, j.d., 1983, ‘ants: bio-indicators of minesite rehabilitation, land-use, and land conservation’, environmental management 7(4), 375–383. http://dx.doi.org/10.1007/bf01866920 majer, j.d. & beeston, g., 1996, ‘the biodiversity integrity index: an illustration using ants in western australia’, conservation biology 10(1), 65–73. http://dx.doi.org/10.1046/j.1523-1739.1996.10010065.x majer, j.d., delabie, j.h.c. & smith, r.b., 1994, ‘arboreal ant community patterns in brazilian cocoa farms’, biotropica 26(1), 73–83. http://dx.doi.org/10.2307/2389112 marais, r., 2004, ‘a plant ecological study of the rietvlei nature reserve, gauteng province’, magister agriculturae thesis, department of animal, wildlife and grassland sciences, university of the free state. morellet, n., gailllard, j.m., hewison, a.j., ballon, p., boscardin, y., duncan, p. et al., 2007, ‘indicators of ecological change: new tools for managing populations of large herbivores’, journal of applied ecology 44(3), 634–643. http://dx.doi.org/10.1111/j.1365-2664.2007.01307.x mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, south african national biodiversity institute, pretoria. neke, k.s. & du plessis, m.a., 2004, ‘the threat of transformation: quantifying the vulnerability of grasslands in south africa’, conservation biology 18(2), 466–477. noss, r.f., 1990, ‘indicators for monitoring biodiversity: a hierarchical approach’, conservation biology 4(4), 355–364. http://dx.doi.org/10.1111/j.1523-1739.1990.tb00309.x pielou, e.c., 1969, an introduction to mathematical ecology, wiley, new york. prins, a.j., 1982, ‘review of anoplolepis with reference to male genitalia, and notes on acropyga (hymenoptera, formicidae)’, annuals of the south african museum 89(3), 215–247. prins, a.j., 1983, ‘a new ant genus from southern africa (hymenoptera, formicidae)’, annuals of the south african museum 94(1), 1–11. r development core team, 2011, r: a language and environment for statistical computing, r foundation for statistical computing, vienna, viewed june 2014, from http://www.r-project.org rawson-tetley, r., animal shelter manager, computer software, viewed 11 january 2005, from http://sheltermanager.sourceforge.net roberts, d.w., 2014, ‘labdsv: ordination and multivariate analysis for ecology’ r package version 1.6-1’, viewed june 2014, from http://cran.r-project.org/package=labdsv ruiz-jaén, m.c. & aide, t.m., 2005, ‘vegetation structure, species diversity, and ecosystem processes as measures of restoration success’, forest ecology and management 218(1), 159–173. samways, m.j., caldwell, p.m. & osborn, r., 1996, ‘ground-living invertebrate assemblages in native, planted and invasive vegetation in south africa’, agriculture, ecosystems and environment 59(1), 19–32. south african national biodiversity institute (sanbi), 2013, grasslands ecosystem guidelines: landscape interpretation for planners and managers, compiled by cadman, m., de villiers, c., lechmere-oertel, r. & mcculloch, d. south african national biodiversity institute, pretoria. shannon, c.e., 2001, ‘a mathematical theory of communication’, acm sigmobile mobile computing and communications review 5(1), 3–55. http://dx.doi.org/10.1145/584091.584093 smith, k.l. & jones, m.l., 2005, ‘watershed-level sampling effort requirements for determining riverine fish species composition’, canadian journal of fisheries and aquatic sciences 62(7), 1580–1588. http://dx.doi.org/10.1139/f05-098 van aarde, r. & smit, a., 1997, the ecology of rehabilitating coastal dunes at richards bay, university of pretoria, pretoria. van hamburg, h., andersen, a.n., meyer, w.j. & robertson, h.g., 2004, ‘ant community development on rehabilitated ash dams in the south african highveld’, restoration ecology 12(4), 552–558. http://dx.doi.org/10.1111/j.1061-2971.2004.00421.x weir, j.s., 1978, the ant iridomyrmex as a biological indicator of pesticide contamination, report for n.s.w. state pollution control commission, new south wales, australia. welch, b.l., 1947, ‘the generalization of “student’s” problem when several different population variances are involved’, biometrika 34(1/2), 28–35. http://dx.doi.org/10.2307/2332510 zaloumis, n.p. & bond, w.j., 2010, ‘grassland restoration after afforestation: no direction home?’, austral ecology 36(4), 357–366. http://dx.doi.org/10.1111/j.1442-9993.2010.02158.x appendix 1: methods and materials used for soil analyses soil samples were taken from each of the sites with the aid of a soil auger. the auger was twisted into the soil until the chamber filled with soil. the soil was placed into labelled plastic bags. three soil samples were taken randomly from the area around each site. the three samples taken at each site were combined and used as one sample. the soil ph, soil organic matter, sand, silt and clay contents were determined for the 12 sites surveyed. the ph of the soil was determined with the use of a ph meter (accurate to 0.05 ph units). initially, twelve 50-cm3 beakers were labelled and weighted on a balance (accurate to 0.1 g) and their weights recorded. a 10-g sample of each of the 12 soil samples was placed into the respectively labelled beakers. an automatic dispenser was used to dispense 25 cm3 of de-ionised water into each of the beakers. the beakers were shaken for approximately 5 s and allowed to stand for 60 min. each beaker was then shaken before the electrodes of the ph meter were placed into the soil sample. the results obtained are reported as ph (h2o). the sand, silt and clay contents of the soil were determined with the use of the particle size distribution method. twelve beakers (250 cm3 capacity) were weighed and their weights recorded. each beaker received 50 g of soil from a respective site. to each beaker of soil, 10 cm3 of calgon dispersing solution (a mixture that contains sodium hexametaphosphate [napo4] and sodium carbonate [na2co3] and de-ionised water) was added. the contents of a beaker were placed into a dispersion cup and filled to approximately 150 cm3 with de-ionised water. the sample was then mixed for 5 min with the use of an electric mixer (10 000 rpm – 13 000 rpm). the sand fraction of the soil was removed by washing the dispersed sample on a 0.053-mm-size sieve. this sieve would ensure that only silt and clay could pass through the funnel into a 1-dm3 cylinder. the washing process was continued until the percolate was clear. the sieve was then removed from the cylinder, and the sand was transferred into a labelled beaker. this process was repeated for all 12 soil samples. the beakers containing the separated sand were dried in a drying oven at 105 °c to constant mass. the samples were then weighed and the masses of the sand (excluding the weight of the beaker) were recorded. the mass of the sand fraction was given as a. following this, the cylinders containing the silt and clay were filled to the 1-dm3 mark. each of the cylinders was stirred with a hand stirrer for a period of 30 s. a ‘blank’ was prepared by adding 10 cm3 calgon to a 1-dm3 cylinder of water. the cylinders were left undisturbed for a total period of 6 hours 30 ± 5 min. once this time had elapsed, a standard hydrometer (with bouyoucos scale in grams per litre, ranging from –5 to 60) was slowly inserted into the suspension of each cylinder and the recording (c) was taken. the hydrometer was placed into each suspension carefully in order not to mix the solution. at this time, a temperature reading was taken. the particle sizes were calculated as follows: a = mass of sand fraction (g) b = hydrometer reading of blank c = hydrometer reading of sample m = mass (g) of soil used sand fraction: clay fraction: to calculate the clay reading, the blank hydrometer reading was subtracted from the sample reading (c − b) and the percentage clay under the appropriate temperature was determined using a particle size distribution table (where e is the reading obtained from the table). silt fraction: the soil organic matter was determined by weighing 12 labelled beakers (50 cm3) and recording the masses of each. an approximate mass of between 10 g and 20 g of each soil sample was weighed and placed into the relevant beakers. the beakers were placed in a drying oven at 105 °c and left overnight to ensure that any moisture in the samples was removed. when removed from the drying oven, the samples were weighed and transferred to a furnace (550 °c) and left for a period of 16 hours. the beakers were then weighed and the mass of the soil organic matter determined. this percentage was then subtracted from the previously determined sand fraction to produce a new adjusted sand fraction. figure 1-a1: sample-based rarefaction curves indicating the number of species (srar), ice mean (incidence-based coverage estimator), chao 2 mean (abundance-based richness estimator), jack 2 mean (second-order jackknife richness estimator) and the michaelis–menton mean (mmmeans) richness estimators of ants in (a) control sites and (b) rehabilitated sites of the rietvlei nature reserve. appendix 2: braun–blanquet scale for vegetation cover table 1-a2: plant morphospecies abundance quantified with the use of the braun–blanquet abundance scale and the altered scale that was used during the analysis of data. figure 1-a2: box plots indicating the various vegetation indices recorded in 1 m × 1 m quadrats, (a) percentage bare ground, (b) percentage grass cover, (c) percentage of other cover (forbs, shrubs and herbs), (d) percentage rock cover, (e) height (cm) of the tallest plant in each of the quadrats for each of the sites and (f) height of the biomass recorded from the disk pasture meter. appendix 3: ant species lists for various data sets specimens were identified to species level where possible. specimens that could not be identified to species level were identified to genus and assigned to numbered morphospecies. table 1-a3: the number of individuals per species, listed per subfamily, which were collected in total for rehabilitated and control sites on the rietvlei nature reserve, south africa. figure 1-a3: box plots indicating the properties of the soil collected from the different treatments: (a) percentage soil organic matter, (b) percentage sand, (c) percentage silt, (d) percentage clay, (e) ph and (f) percentage coarse sand. appendix 4: tables illustrating sampling completeness calculated for the rehabilitated and control sites table 1-a4: richness estimator values as obtained from estimate-s (v 9.1.0; colwell 2013). table 2-a4: richness estimator values as obtained from estimate-s (v 9.1.0; colwell 2013). table 3-a4: the reliability of each inventory was calculated for the control sites. table 4-a4: the reliability of each inventory was calculated for the rehabilitated sites. table 5-a4: overall sampling completeness. figure 1-a4: non-metric multidimensional scaling plot indicating the similarity of plant assemblages among replicates of control and rehabilitated sites. appendix 5: list of indicator values table 1-a5: indicator values generated in rstudio. species fulfilling the criteria (indval > 70%, p < 0.05) are identified as indicator species. article information authors: graham r.h. grieve1 colleen t. downs1 affiliations: 1school of life sciences, university of kwazulu-natal, south africa correspondence to: colleen downs email: downs@ukzn.ac.za postal address: private bag x01, pietermaritzburg 3209, south africa dates: received: 30 june 2014 accepted: 07 jan. 2015 published: 16 sept. 2015 how to cite this article: grieve, g.r.h. & downs, c.t., 2015, ‘a checklist of the plants of the forests and grasslands in the weza district, southern kwazulu-natal and a review of their status in the red data list’, koedoe 57(1), art. #1237, 7 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1237 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/koedoe.v57i1.1237-1. copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. a checklist of the plants of the forests and grasslands in the weza district, southern kwazulu-natal and a review of their status in the red data list in this checklist... open access • abstract • introduction • methods    • study area    • data sources    • species list arrangement • results and discussion • conservation status and management • acknowledgements    • competing interests    • authors’ contributions • references • footnote abstract top ↑ eastern mistbelt forests are naturally fragmented forests with grassland which occur from the eastern cape to kwazulu-natal, south africa. these were heavily logged by colonial settlers and continue to be harvested despite being protected. consequently we documented a checklist of the plants of the forests and grasslands in the weza district (3029da weza), southern kwazulu-natal, including ngeli forest and nearby indigenous forest patches to highlight their biodiversity status and need for conservation. we also reviewed their status in the red data list. of the 1554 records included in this summary of plant species for the weza district, there were 6 lichens (0.4%), 46 bryophytes (3.0%), 58 pteridophytes (3.7%), 6 gymnosperms (0.4%) and the remaining 1424 species angiosperms (92.5%). of the angiosperms, 27.3% were monocotyledons and 72.7% were dicotyledons. the most species-rich family was asteraceae (239 species) followed by fabaceae (115 species), liliaceae (used for purposes of comparison against older studies – 89 species), orchidaceae (89 species), iridaceae (59 species), poaceae (58 species), asclepidaceae (again used for purposes of comparison against older studies – 57 species), scrophulariaceae (42 species), euphorbiaceae (32 species), lamiaceae (32 species) and rubiaceae (27 species). these 10 families each comprised more than 2% of the species in the list. together they contributed 55% of the angiosperm species and 34.1% of the angiosperm genera. the biodiversity and conservation value of the study area are conserved pockets of eastern mistbelt forest, drakensberg foothill moist grassland and mistbelt grassland. more than 4% of the species are under some degree of threat, as was evidenced by the number of species regarded as endangered (5), vulnerable (18), near threatened (10), critically rare (1), rare (20) or declining (11) amongst the 1554 species covered in the list. conservation implications: in terms of taxa under some degree of threat, number of endemic and near endemic species the biodiversity and conservation value of the ngele (3029da weza) area is reinforced. this necessitates that the area is appropriately protected from invasive alien species, and uncontrolled and illegal use by the neighbouring communities. introduction top ↑ globally africa currently has one of the highest levels of deforestation, which affects both habitat quality and species diversity (bodart et al. 2013; fao 2010; lawes, macfarlane & eeley 2004; wade et al. 2003). in south africa the forest biome is the smallest of the eight biomes (mucina & geldenhuys 2006). eastern mistbelt forests, which form part of the southern mistbelt forest group, are naturally fragmented forests with grassland which occur from the eastern cape to kwazulu-natal (kzn) (mucina & geldenhuys 2006). these forests are dominated by afrocarpus/podocarpus species (yellowwoods) (adie, rushworth & lawes 2013; moll 1972). drakensberg foothill moist grassland and midlands mistbelt grassland comprise the largest part of the untransformed grassland habitat in the study area and are home to a wide variety of plants (mucina & geldenhuys 2006). the first forest legislation in kzn was passed in 1853, a forest service was started in 1891, and a functional forestry department in 1902 (fourcade 1889; king 1941). however, most of the forests had already been exploited by european settlers, including colonial woodcutters (cawe & mckenzie 1989; cooper 1985; king 1941). fourcade (circa 1889) was seconded to the natal government to prepare a report on the condition of indigenous forests on crown and trust lands (fourcade 1889). his report subsequently led to the decision in 1891 to protect the crown forests (fourcade 1889; storrar 1990). remnants of sawpits can still be seen in ngeli forest and others, and although the forests appear to have recovered substantially since protection was afforded (pers. obs.), regeneration is slow and the canopies are generally irregular and areas reduced (geldenhuys 2007; lawes et al. 2004; moll 1972). following the colonial logging exploitation in the mid-1800s and intermittently to the 1940s, these forests in kzn have been allowed to recover by natural succession processes (adie et al. 2013). however, as most of these forests are in close proximity to communal lands, subsistence wood harvesting and use of the forest and grasslands to graze cattle has continued (adie et al. 2013; lawes, griffiths & boudreau 2007; robertson & lawes 2005; wirminghaus et al. 1999; pers. obs.). despite being protected, generally conservation of these forests and their associated grasslands are difficult (adie et al. 2013; cawe & mckenzie 1989; downs 2005; wirminghaus et al. 1999). it is clearly difficult to determine whether the present plant species diversity in ngeli forest and nearby forest patches and associated grasslands is similar to what existed before the exploitation that began in the mid to late 19th century, although some of the specimens collected and included in this species list date from that period. therefore we documented a checklist of the plants of the forests and grasslands in the weza district (3029da weza), southern kzn, including ngeli forest and nearby indigenous forest patches to highlight the biodiversity importance of this area and need for conservation. the list could serve as an important conservation management tool in future. we also reviewed the plants’ status in the red data list. methods top ↑ study area the study area was the 3029da quarter-degree square described as 3029da weza (figure 1; table 1). this includes the village of weza in the southern interior of kzn, south africa, close to the border with the eastern cape, situated between harding and kokstad. it is a centre for timber growing and has a sawmill to process the pine pinus sp. logs extracted from the plantations in the surrounding hills. the weza state forests, which include pine plantations as well as indigenous forest patches and grassland (firebreaks and the tops of the mountain range), lie to the south-east of the ngeli mountain range. the range runs from a peak at 30°34.3's, 29°35.5’e on the eastern cape border, past the dominant peak of ngeli mountain at 2267.7 m and slowly reduces in height towards the north-east. there is a high point to the north of the n2 highway on which fire lookout towers are situated at 30°31.2's, 29°41.9’e. the study area lies generally to the south-east of the axis joining these extremes. figure 1: map of the study area. source: authors’ own creation table 1: areas of different habitat or land use in the study area. the ngeli forest and nearby forest patches and associated grasslands lie between an altitude of 950 m adjacent to weza and the altitude of the higher peaks at about 2200 m. the ngeli mountain range is regarded as an outlier of the drakensberg alpine centre (carbutt & edwards 2006). one of the larger forest patches straddles the n2 highway roughly midway between harding and kokstad, but the bulk of the study area is to the south-west of the n2 highway and south-east of the ngeli range. the umtamvuna river rises in the ngeli range and flows in a south-easterly direction to reach the sea just south of port edward. apart from a short section in the ngeli range, the umtamvuna forms the border between the eastern cape and kzn. the forests in this area fall under the foz 3 southern mistbelt forest category, whilst the grasslands are categorised as gs 9 midlands mistbelt grassland and gs 10 drakensberg foothill moist grassland (mucina & rutherford 2006). the average rainfall in this area is 991 mm/a, although there appears to have been periods of below-average (664 mm/a) and above-average (1476 mm/a) rainfall during the period covered by the data (whyle, unpublished data). if the rainfall is considered as the precipitation during the rainy season from the beginning of july until the end of june in the following year, there are similar peaks and troughs but there are also much greater swings in total rainfall, from a low of 580 mm to a high of 1624 mm (whyle, unpublished data). the main parts of the exposed rocks of the ngeli range are composed of dolerite rocks and associated soil types (wirminghaus et al. 2001). norman (2012) notes that the dolerite occurs in sills and dykes, and these are mixed with un-deformed sedimentary strata from the lower karoo age (permian). the sedimentary horizons and sills are almost horizontal. data sources the plant species list for the study area is based on data mainly from voucher specimens collected by a wide range of botanists, although there are several species, mostly naturalised invasive alien species, for which only ‘observation’ data are currently available. in addition, voucher specimens were collected from 2009 to 2014 whilst conducting monthly research in the area (hart, grieve & downs 2013) and specimens were identified with the aid of pooley (1998, 2003) and boon (2010). the voucher specimens included in this species list are housed at a number of herbaria around the country, namely kbg, nbg, nh, nu, pce and pre. an attempt was made to obtain an extract of data from the schweikert herbarium at pretoria university (as many records of a. abbott and a.e. van wyk are known to be housed in the schweikert herbarium), but they advised they were in the process of migrating their database records from precis to brahms and were unable to assist at the time the request was made. although the collecting of species on which this study is based took place over an extended period (1933–2014), there may still be species which have been overlooked. the initial stimulus for this work was provided by t. abbott, who made a plant data list of ngeli species available (abbott 2010). the south african national biodiversity institute (sanbi) was asked to make similar lists available for the collections at pre and nh, and use was made of the sibis (sanbi's integrated biodiversity information system) website (http://sibis.sanbi.org/) for further data. information was received in discrete subsets which had to be consolidated and duplication removed. where necessary, additional information on collector id, voucher number and location of voucher were also derived from the sibis site. species list arrangement the arrangement of the species list and species nomenclature follows the red list of south african plants (raimondo et al. 2009). this was used as it was felt important to determine the red list status of as many of the species in the 3029da quarter-degree square as possible to facilitate and reinforce conservation measures for the area. however, this publication only covers vascular plants. in the case of the bryophytes, use was made of ‘a checklist of south african plants’ (germishuizen et al. 2006), and for the few lichens amongst the data, google searches had to suffice; in neither of these cases was there any information on conservation status of these species. use was made of the lists of endemic and near endemic species for the drakensberg alpine centre (carbutt & edwards 2006) to show which species in the ngeli area are endemic and near endemic. this information was included, where applicable, for the angiosperms in the species list (online appendix 1). naturalised exotic species (weeds) were included and appropriately indicated. the web-based red data list (rdl) (raimondo et al. 2009) was used to determine the status of each of the vascular plant species recorded for the study area. percentages of each rdl category falling within the order breakdown were determined. results and discussion top ↑ of the 1554 records included in this summary of plant species for the weza district, there were 6 lichens (0.4%), 46 bryophytes (3.0%), 58 pteridophytes (3.7%), 6 gymnosperms (0.4%), and 1438 angiosperms (92.5%) (table 2). the number of plant species was highest in the grasslands, and most of these were herbaceous species (table 3). of the angiosperms, 27.3% were monocotyledons and 72.7% were dicotyledons. the ratio of monocotyledons to dicotyledons for ngeli was 1:2.7. this was higher than reported for mahwaqa mountain (1:2.1) (meter et al. 2002). table 2: analysis of plant orders, families, genera and species found in the ngeli study area. table 3: habitat preference of species listed in online appendix 1 (a) and summary of vascular plants listed in online appendix 1 (b). recently, the flora of the great winterberg-amatholes area (7382 km2 in extent) was described (clark et al. 2014). whilst this region has similar numbers of families for the pteridophytes, gymnosperms, monocotyledons and dicotyledons, the numbers of genera and species are higher, possibly because of the larger area studied compared with the current study area. the number of endemics recorded in this study (43) exceeds the number of endemics in the clark et al. (2014) study (35) in a much larger area. only plant families with more than 1% of the total number of species for the ngeli quarter-degree square were included (table 4). to make this comparable with a similar list (meter et al. 2002), the family structure of asclepidaceae before consolidation into apocynaceae was used (table 4). the 24 families contributed 71.0% of the species and 57.2% of the genera of this list. table 4: synopsis of families which contribute 1% or more of total number of species for the ngeli study area together with number of genera in each family. as is the case with mahwaqa mountain, kzn (meter et al. 2002), the most species-rich family for 3029da weza was asteraceae, with 239 species (table 4, online appendix 1). there were 14 endemics (5.9%) and 67 near endemics (28.0%) amongst the listed asteraceae. amongst the asteraceae, the following species fell into ‘endangered’ categories in the rdl: berkheya pannosa (rare, endemic); felicia wrightii (critically rare, endemic); gnaphalium griquense (rare, near endemic); helichrysum tenax var. palidum (rare, near endemic); phymaspermum villosum (rare); and senecio poseidonis (threatened, near endemic) (table 5). asteraceae also included 11 (4.6%) naturalised species. table 5: consolidated summary of the red data list status of vascular plants for 3029da weza. the second most species-rich family for 3029da weza was fabaceae, with 115 species (table 4, online appendix 1), in contrast to poaceae in meter et al. (2002) (table 6). however, this was consistent with the ranking of fabaceae as the second most important species contributor to the kzn flora (hilliard & burtt 1987). there was one endemic (0.9%) and 11 near endemic species (9.6%) amongst the listed fabaceae. amongst the fabaceae, the following species fell into ‘endangered’ categories in the rdl (table 5): crotalaria dura subsp. dura (near threatened) and psoralea abbottii (vulnerable). table 6: nine of the largest families and their contributions to the flora of the ngeli study area, compared against those for mahwaqa mountain, for the southern drakensberg, and for kwazulu-natal as a whole. using the old family structure liliaceae for comparative purposes, this family is the third most species-rich family, with 109 species (table 4). there were 5 endemic (4.6%) and 14 near endemic species (12.8%) amongst the listed liliaceae. amongst the liliaceae, the following species fell into ‘endangered’ categories in the rdl (table 5): aloe kniphofioides (vulnerable), aloe linearifolia (near threatened), sandersonia aurantiaca (declining), bowiea volubilis (vulnerable), eucomis bicolor (near threatened), eucomis comosa (declining) and merwilla plumbea (near threatened). the orchidaceae was the fourth most species-rich family in 3029da weza, with 89 species (table 4, online appendix 1). there were three endemics and 25 near endemics amongst the listed orchidaceae. four orchid species collected were listed as rare. these were: disa oreophila subsp. erecta (rare, endemic); disa tysonii (rare, near endemic); satyrium microrhynchum (rare, near endemic) and schizochilus flexuosus (rare, near endemic). the iridaceae was the fifth most species-rich family in 3029da weza, with 59 species (table 4, online appendix 1) three of these were endemic (5.1%) and 21 were near endemic (35.6%) (table 5). one of these species was listed as rare, one as vulnerable, one as declining and one as endangered; they were: dierama ambiguum (endangered, near endemic) (table 5); dierama tysonii (vulnerable, near endemic); gladiolus oppositiflorus subsp. salmoneus (declining, near endemic) and hesperantha ingeliensis (rare, near endemic). the poaceae ranked sixth, with 58 species in 3029da weza (table 4, online appendix 1). a comparison of the relative richness of poaceae at ngeli against that reported for mahwaqa mountain, the southern drakensberg (meter et al. 2002) and kzn as a whole suggest that poaceae might have been under-collected at ngeli (table 6). indeed, once a base species list was available, it proved to be relatively easy to collect previously uncollected grass species in the study area; there are undoubtedly further grass species awaiting collection there. this historical under-collecting of poaceae could explain the difference in monocotyledon to dicotyledon ratio between the present study and that of meter et al. (2002) (table 6). one species was endemic (1.7%) and three were near endemic (5.2%) (table 3). poaceae included 5 naturalised species (table 4, online appendix 1).there were no ‘endangered’ species in this list (table 5, online appendix 1). the asclepiadaceae (57 species) (table 4, online appendix 1), again used for purposes of comparison against older studies, was the seventh most numerous family in 3029da weza. three of these species were endemic (5.3%) and 17 near endemic (29.8%). five species were listed as rare and three as vulnerable. these were: asclepias xysmaloboides (rare, endemic); aspidonepsis cognate (rare, near endemic); brachystelma molaventi (vulnerable); brachystelma tenellum (vulnerable); emplectanthus gerrardii (rare); schizoglossum elingue subsp. purpureum (rare, endemic); schizoglossum montanum (rare, endemic); schizoglossum singulare (vulnerable).1 the scrophulariaceae had 42 species in 3029da weza (table 4, online appendix 1). of these, 4 were endemic (9.5%) and 11 were near endemic (26.1%) (table 3). none of these scrophulariaceae were listed as endangered in the rdl (table 5). the euphorbiaceae in 3029da weza had 32 species (table 4, online appendix 1). amongst these were two near endemic species (table 5), one near threatened species (alchornia hirtella forma glabrata) and one vulnerable species (euphorbia flanaganii). the lamiaceae had 32 species in 3029da weza (table 4, online appendix 1). amongst these were four near endemic species (table 5). none of these lamiaceae were in ‘endangered’ categories in the rdl, but there were two naturalised species (table 5). the 10 families above all comprised more than 2% of the species in the list (online appendix 1). together they contributed 53.5% of the species and 41.9% of the genera. comparative data for this study, for mahwaqa mountain, for the southern drakensberg (meter et al. 2002) and for kzn as a whole, are shown in table 6. as mentioned before, the poaceae in 3029da weza appeared to be comparatively under-represented, despite additional collecting. fabaceae, in contrast, appeared to be over-represented when compared with mahwaqa and the southern drakensberg (meter et al. 2002), although as a proportion of all recorded species, their data were more in line with the data for kzn as a whole. these apparent anomalies might justify a review of the poaceae (some work already done) and fabaceae to determine whether these relative proportions are real or as a result of collecting bias. the latter is unlikely given the significant number of collectors involved here. conservation status and management top ↑ the biodiversity and conservation value of the study area lie in the conserved pockets of eastern mistbelt forest and drakensberg foothills moist grassland and eastern mistbelt grassland. this was evidenced by the number of species regarded as endangered (5), vulnerable (18), near threatened (10), critically rare (1), rare (20) or declining (11) amongst the 1554 species covered in the list, that is, more than 4% of the species are under some degree of threat. this, together with the fact that, amongst the angiosperms, there are 44 (3.1%) endemic species (11 monocotyledons and 33 dicotyledons) and 256 (17.8%) near endemic species (79 monocotyledons and 177 dicotyledons), reinforces the biodiversity and conservation value of the 3029da weza area. furthermore, this area supports important fauna (adie et al. 2013; c. potgieter [university of kwazulu-natal], pers. comm.). this necessitates that the responsible authorities ensure that the area is appropriately protected from invasive alien species, and uncontrolled and illegal use by the neighbouring communities (adie et al. 2013). an interesting question to be investigated is whether all the species collected in the period leading up to 1891, when protection was afforded this and other kzn forests (mccracken 2004), are still present. acknowledgements top ↑ w. rozani, o. zikhali and b. dzidza from the department of agriculture, forestry and fisheries (daff) are thanked for permission to undertake the research work in the ngeli forests. the daff forestry security guards s. cwele, d. masuku and r. madwe are thanked for their considerable assistance in the field during data collection. t. abbott kindly supplied a significant portion of the data on which this paper is based, and assisted with tree identifications at the outset of the project. he also kindly facilitated the first author's involvement with the nicholson/abbott herbarium at the umtumvuna nature reserve which led to the preparation of the most recent specimens and accession of these into that collection. b. bytebier provided valuable guidance on the structuring of the species list. y. ehlers-smith kindly generated the maps used. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions both g.r.h.g. (university of kwazulu-natal) and c.t.d. (university of kwazulu-natal) conceived the project and wrote the manuscript. in addition, c.t.d. sourced funding and g.r.h.g. collected and analysed the data. references top ↑ abbott, a.t.d., 2010, checklist of ngele mountain and environs weza state forest 3029da collectors. unpublished report, nicholson/abbott herbarium, port edward. adie, h., rushworth, i. & lawes, m.j., 2013, ‘pervasive, long-lasting impact of historical logging on composition, diversity and above ground carbon stocks in afrotemperate forest’, forest ecology and management 310, 887–895. http://dx.doi.org/10.1016/j.foreco.2013.09.037 bodart, c., brink, a.b., donnay, f., lupi, a., mayaux, p. & achard, f., 2013, ‘continental estimates of forest cover and forest cover changes in the dry ecosystems of africa between 1990 and 2000’, journal of biogeography 40, 1036–1047. http://dx.doi.org/10.1111/jbi.12084 boon, r., 2010, pooley’s trees of eastern south africa, flora & fauna publications trust, durban. carbutt, c. & edwards, t.e., 2006, ‘the endemic and near endemic angiosperms of the drakensberg alpine centre’, south african journal of botany 72, 105–132. http://dx.doi.org/10.1016/j.sajb.2005.06.001 cawe, s.g. & mckenzie, b., 1989, ‘the afromontane forests of transkei, southern africa ii. a floristic classification’, south african journal of botany 55, 31–39. clark, v.r., dold, a.p., mcmaster, c., mcgregor, g., bredencamp, c. & barker, n.p., 2014, ‘rich sister, poor cousin: plant diversity and endemism in the great winterberg-amatholes (great escarpment, eastern cape, south africa)’, south african journal of botany 92, 159–174. http://dx.doi.org/10.1016/j.sajb.2014.01.008 cooper, k.h., 1985, the conservation status of indigenous forests in transvaal, natal and o.f.s., south africa, wildlife society of southern africa, durban. downs, c.t., 2005, ‘abundance of the endangered cape parrot, poicephalus robustus, in south africa: implications for its survival’, african zoology 40, 15–24. food and agriculture organization of the united nations (fao), 2010, global forest resources assessment, forestry paper 163, fao, rome. fourcade, h.g., 1889, report on the natal forests, watson, pietermaritzburg. http://dx.doi.org/10.5962/bhl.title.23002 geldenhuys, c.j., 2007, ‘growth, ingrowth and mortality patterns over stands and species in the forest growth sites at weza and ngome indigenous forests, kwazulu-natal: report on first remeasurement, 2003/2004’, project no. 2008-450, department of water affairs and forestry, pretoria. germishuizen, g., meyer, n.l., steenkamp, y. & keith, m., 2006, ‘a checklist of south african plants’, south african national biodiversity network report no. 41, sabonet, pretoria. hart, l., grieve, g. & downs, c.t. 2013, ‘fruiting phenology and implications of fruit availability in the fragmented ngeli forest complex, kwazulu-natal, south africa’, south african journal of botany 88, 296–305. http://dx.doi.org/10.1016/j.sajb.2013.08.003 hilliard, o.m. & burtt, b.l., 1987, the botany of the southern natal drakensberg, annals series, vol 15, national botanical gardens, cape town. king, n.l., 1941, ‘the exploitation of the indigenous forests of south africa’, journal of the south african forestry association 6, 26–48. http://dx.doi.org/10.1080/03759873.1941.9631098 lawes, m.j., 1990, ’the distribution of the samango monkey (cercopithecus mitis erythrarchus peters, 1852 and cercopithecus mitis labiatus i. geoffroy, 1843) and forest history in southern africa’, journal of biogeography 17, 669–680. http://dx.doi.org/10.2307/2845148 lawes, m.j., griffiths, m.e. & boudreau, s., 2007, ‘colonial logging and recent subsistence harvesting affect the composition and physiognomy of a podocarp dominated afrotemperate forest’, forest ecology and management 248, 48–60. http://dx.doi.org/10.1016/j.foreco.2007.04.012 lawes, m.j., macfarlane, d.m. & eeley, h.a.c., 2004, ‘forest landscape pattern in the kwazulu-natal midlands, south africa: 50 years of change or stasis?’, austral ecology 29, 613–623. http://dx.doi.org/10.1111/j.1442-9993.2004.01396.x mccracken, d.p., 2004, ‘dependence, destruction and development: a history of indigenous timber use in south africa’, in m.j. lawes, h.a.c. eeley, c.m. shackleton & b.g.s. geach (eds.), indigenous forests and woodlands in south africa, pp. 77–308, university of kwazulu-natal press, pietermaritzburg. meter, e.b., edwards, t.j., rennie, m.a. & granger, j.e., 2002, ‘a checklist of the plants of mahwaqa mountain, kwazulu-natal’, bothalia 32, 101–115. moll, e.j., 1972, ‘the current status of mistbelt mixed podocarpus forest in natal’, bothalia 10, 595–598. mucina, l. & geldenhuys, c.j., 2006, ‘afrotemperate, subtropical and azonal forests’, in l. mucina & m.c. rutherford (eds.), the vegetation of south africa, strelitzia 19, pp. 586–655, south african national biodiversity institute, pretoria. mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, strelitzia 19, south african national biodiversity institute, pretoria. norman, p.g., 2012, ‘the geology of the ngele area: a brief geological description’, unpublished report, southport geology and natural history society, uvongo. pooley, e.a., 1998, field guide to wild flowers kwazulu-natal and the eastern region, natal flora publications trust, durban. pooley, e.a., 2003, mountain flowers: a field guide to the flora of the drakensberg and lesotho, flora publications trust, durban. raimondo, d., von staden, l., foden, w., victor, j.e., helme, n.a., turner, r.c. et al., 2009, red list of south african plants, strelitzia 25, south african national biodiversity institute, pretoria. robertson, j. & lawes, m.j., 2005, ‘user perceptions of conservation and participatory management of igxalingenwa forest, south africa’, environmental conservation 32, 64–75. http://dx.doi.org/10.1017/s0376892905001979 storrar, c., 1990, the four faces of fourcade: the biography of a remarkable scientist, maskew miller longman, cape town. wade, t.g., riitters, k.h., wickham, j.d. & jones, k.b., 2003, ‘distribution and causes of global forest fragmentation’, conservation ecology 7(2), 7. wirminghaus, j.o., downs, c.t., symes, c.t. & perrin, m.r., 1999, ‘conservation of the cape parrot in southern africa’, south african journal of wildlife research 29, 118–129. wirminghaus, j.o., downs, c.t., symes, c.t. & perrin, m.r., 2001, ‘fruiting in two afromontane forests in kwazulu-natal, south africa: the habitat type of the endangered cape parrot poicephalus robustus’, south african journal of botany 67, 329–336. footnote top ↑ 1. in a subsequent collecting trip with two other postgraduate researchers, collections were made of several asclepiadaceae believed to be undescribed species. additional work done by these other researchers may lead to the publication of new species descriptions. filelist convert a pdf file! supplement to koedoe. 1977: 190-196. conservation education on the academic level in southern africa j du p bothma eugene marais chair of wildlife management university of pretoria pretoria 0002 introduction conservation in some form, albeit dormant at times, has probably been with man for many centuries. yet wildlife conservation as a science is a relatively new concept, which basically originated in the united states of america (usa). that country also led the world in developing conservation education. this lead was followed by most progressive countries, although the nature of conservation and its related educational processes has been adopted to the attitudes and needs of individual countries. the oldest und ergraduate course in wildlife conservation (through management) started in the usa in 1924, and this was followed five years later by two more courses. by 1948 th ere were 23 colleges offering wildlife management training in the usa (trippensee 1948) and this increased to 97 in 1970 (evenden 1971) . the first textbook on wildlife conservation, and one still widely used, appeared in 1933 when leopold published his book: "game management" (trippensee 1948). the expanding interest in academic education in conservation has recently become the focus for careful re-evaluation in the usa and also in the r epublic of south africa (rsa). however, before discussiong conservation education, on e must first consider what is conservation. conservation cons ervation and management of wildlife have become almost synonymous since leopold (1933) expounded on his principle of conservation through wis e use. different authors give varying views on what they consider to be the essence of conservation. clarke (1973) said that cons ervation is religion because the religious impulse is directed primarily to the conservation and promotion of life. like wildlife conservation, this leads to the riddance of anything considered as hostile and the enhancement of what is thought to be favourable to life. warren and goldsmith (1974) see conservation as the meeting point of cultural, 190 social and economic systems on the one hand and natural systems at the other and say that conservation actions aim at the planned , harmonious interlocking of these two sets of systems and requires a knowledge of both. darwin already showed man's place in the panorama of creation, but it remained for leopold to define man as a geological and ecological force in nature and to hold that man has an inescapable responsibility for the land. lately the importance of the landscape has been added to man's list of conservation concern (hickey 1974) . wildlife the conservation actions that we are dealing with in this paper are those associated with wildlife. originally wildlife was written as two words, but since the early 1930's it is regarded as one word . to the ecologist wildlife simply means all living things (hickey 1974). leopold (1933) stated the objective of wildlife conservation to be: to retain for the average citizen the opportunity to see, admire and enjoy, and the challenge to understand the varied forms of wildlife indigenous to his state. universities and conservation training in southern africa conservation education on the academic level in southern africa is found at many institutions. the extent and emphasis of this training depends on the aims and interest of the specific university, and are often those of one or more lecturer. i t is not the aim of this paper to discuss these various courses or even to attempt to evaluate them. what will be tried, however, is to examine why and how we are training students in wildlife conservation as seen against the background of what happens to these people once they leave the universities. the major universities involved in wildlife conservation education in southern africa are: natal, pretoria, rhodesia, stellenbosch and to a lesser extent now also the witwatersrand . a broader course in environmental conservation is presented at cape town. the graduates produced by these institutions find employment in the whole conservation spectrum in southern africa, with the universities of natal, pretoria and rhodesia the main suppliers. this crop of conservation-trained biologists has led to the question of whether we are not overstocking a limited market with qualified graduates who may not all find the employment they were specifically trained for and with the resultant frustration of some graduates. this problem, however, is not limited to sou thern africa and has recen tly been discussed in depth in the u ni ted states of america. the results of their evaluation of wildlife student training versus available employment can also be applied to southern africa. before doing so, however, let us briefly look at some aspects of conservation curricula. 191 conservation curricula conservation training is so broad that the setting up of anyone detailed curriculum is as impossible as it is undesirable. however, a wellrounded programme can be viewed as an equilateral triangle of environment, biology and land-use (trippensee 1948). the basic approach is to develop students qualified to tackle management and research problems on an inter-agency and interdisciplinary basis. the renewable resources of the world have been so thoroughly altered by man that conservation without management is seldom possible (warren and goldsmith 1974). the modern conservation student must therefore, be trained in environmental analysis and control, the characteristics and needs of wildlife and in management techniques (trippensee 1948). this takes time to acquire and it is usually built on a cornerstone of biological sciences (larson 1971). the most important point to remember, however, is that the natural environment, although infinitely complex and varied, is a dynamic, organic whole and therefore cannot be properly investigated in isolation (owen 1975). furthermore, whatever his training, the benefit of well-planned education is that the student may be successful in a wide spectrum of employment situations (scott 1975). employment opportunities and graduate training a student entering a university for conservation training is from three to ten years away from terminal graduation depending on the nature of his research and training. this makes predictions of the employment opportunities that he will face upon completion of his studies impossible, and emphasises that training in any field of science must be more than a route to eventual employment. education, especially at the graduate level, should aim at more than preparing a student for a narrow professional job; it should aim at producing a better citizen (scott 1975). universities must provide students with a realistic appraisal of the current employment facilities in their chosen field of study. having done this, no education institution should prohibit or dissuade a suitably qualified student from participating in any study course provided that that institution has the necessary room and manpower to ensure quality instruction to all individuals (sanderson 1975). this approach eliminates arbitrary enrollment quotas (hale 1975) and supports the concept of supply and demand in which competition for the available employment positions should benefit the best qualified graduates and in turn enhances the stature of the conservation profession (peterle 1975). in accepting a student for graduate tr~ining we must resist the temptation to accept doubtful students simply to increase class sizes, to meet research obligations or to initiate research that we are urged to undertake. acceptance of mediocre students is unfair to student and the 192 community. this approach should result in enrollment limitations unrelated to employment opportunities (scott 1975). the wildlife profession is the mainstream of the environmental movement and a change in some wildlife aspect often signals a " dis ease" of the environment . thus wildlife graduates are also well qualified to work in many interdisciplinary environmental studies (scott 1975). there is also no need for the student qualified in conservation to limit himself to that field of employment. we need knowledgeable citizens in all spheres of life. for example, one suitably train ed and motivated high school teacher can probably do more long-term good for conservation than a host of professional wildlife biologists . a wildlife graduate does not necessarily lose the value of his training ifhe is employed outside the direct conservation sphere (sanderson 1975). training of students must also be advanced to keep pace with new d evelopments in their chosen subject. this cannot be achieved by preventing acceptable candidates from study simply because they are not assured of immediate employment in their major field of study (scott 1975). correct use of available graduates another serious question affecting the training of conservation (wildlife) graduates is: are we using our graduates correctly, even when they secure professional wildlife jobs? i believe not. in most conservation bodies in southern africa the bulk of the research staff hav e either completed four years (b.sc. honours) or five to six years (m.sc.) of study. these men are then required to do in depth research which in turn is used to make long-term management proposals. these actions, i believe, should only be entrusted to men with doctoral training . while exceptions do occur, the average student entering employment with a b.sc (honours) degree is at best qualified to act as a conservation field officer, reserve manager or technician. we are making too many conservation and management decisions based on information obtained through research which has barely scratched the surface. one of the most serious problems in this connection is that while most conservation graduates receive interdisciplinary training (the basis of all ecology), they are upon formal employment often required to work in isolation or with the minimum of interand intra-agency co-operation. this results in a waste of time and manpower. scott (1975) states that if we have learned anything at all from the environmental crisis it is that problems must be resolved through interdisciplinary and interagency co-operation . the conservation graduate of today receives such training but usually can only apply it on a limited scale in his own work and research once he is professionally employed. this matter calls for a serious re-evaluation by conservation agencies on how they are using their available manpower. 193 in southern africa we should strive for increased training as the basis of all levels of conservation employment. field staff such as law enforcement officers, reserve managers and game rangers are increasingly required to collect wildlife management and research information and must be truly professional in their training. this can only be done by setting a b.sc. degree in wildlife conservation as a minimum requirement for their training (sigler 1975). the law enforcement or nature conservation officer of today is a broadly-based resource manager. his actions must be based on the correct applicable biological and political principles. to convince the layman of the need for conservation the field officer must have the knowledge to do so and the ability to explain his conservation actions. field and research staff also too often clash with each other, to the detriment of conservation. according to sigler (1975) this is partly the result of administrative restrictions involving educational requirements, working hours, social status and salary. it is suggested that field staff be given equal status and opportunities at education and career-building. this is the basis of any conservation action. a field officer who understands the actions of research staff can better evaluate and defend them. the law enforcement officer or game ranger must therefore, have a professional status and attitude of his own. he often not only creates the public image of conservation but he usually is the public image of any conservation body. this requires basic training as a biologist and advanced training in several fields of his own (sigler 1975), a love for outdoor work and a real interest in wildlife (trippensee 1948). conservation action without reliable factual information usually leads to the deterioration of the situation (dasmann 1972). basic conservation research is the field of the master's candidate, and in depth reserach requires training at the doctoral level (trippensee 1948). when conservation bodies do not have the staff to carry out research on this basis they should seriously consider using suitable graduates on ad hoc projects involving temporary (project) employment ties only. research by a master's degree candidate usually only produces a man who has a good understanding of the research techniques (not principles) in his field of study and practice in writing a finished report (trippensee 1948). the woman in conservation more and more women are becoming interested in conservation employment. in 1971 in the usa 13% of all students at the top 20 wildlife schools were women (rongstad and hickey 1972). these people can all be used successfully, especially as public relations experts. women school teachers with conservation training can also be of inestimiable benefit to the whole conservation movement. 194 conservation at the national level successful students in conservation enter a variety of employments at the national level. let us briefly consider this environment into which they venture and examine their role in it. government and semi-government conservation action may often be considered slow and difficult to initiate, but the powers of legislation, taxation and access to funds make it ultimately the most effective (warren and goldsmtih 1974) and one which employs most conservation graduates. the important prerequisite for any conservation action or body is a clear statement of demands. it is my belief that we in the rsa need a continuous reassessment of our conservation progress and our use of men trained in conservation matters. this can only be achieved effectively if we have a well-planned comprehensive national conservation action plan to serve as our reference guide. in setting up this plan it will be well to reassess our whole conservation movement and to redefine the tasks of the various conservation bodies in south africa. we cannot live in isolation. to pretend that the problems of other nations, and also other conservation bodies, can be ignored, is like telling a fellow passenger that his end of the boat is sinking (dasmann 1972). one of our most immediately urgent conservation tasks in the rsa is to obtain all the land needed for conservation and then to use it properly. in doing so we must remember that cheap budgets often only lead to cheap results! once we have secured this land we can start worrying about who shall manage it and how it is to be done. conclusion it should be clear that there is scope for reassessment of our entire attitude to conservation education, its products and their employment environment. the student with a bachelor's degree does not automatically have a certificate to professional employment and in depth research should only be done by senior students. field officers constitute the largest single career group in conservation, should become better qualified and should become more involved with research and management programmes. a broad approach to environmental conservation has become a necessity. this can only be achieved by a national conservation action plan administered on a national level. the guidance and impetus for the implementation of such a national conservation plan should come from top government executive level. one-sided approaches to conservation usually end in failure. nobody can escape some share of responsibility; nobody can avoid the world in which he lives. failure to act can only result in the need to live with the consequences of other people's action (dasmann 1972). 195 references clarke, c h d 1973. conservation revisited. wildl. soc. bull. 1(2): 106-108. dasmann , r f 1972 environmental conservation. 3rd ed. new york: john wiley and sons. evenden , f g 1971. wildlife student enrollment up 25,4% over '68. wildl. soc. ne ws 132: 1-3. hale,j b 1975. comments on scott' s views on the training of wildlife students. wildl. soc. bull. 3(2): 62-63. hickey, j j 1974. some historical phases in wildlife conservation. wildl. soc. bull. 2(4): 164-170. larson , j s 1971. the baccalaureate dilemma. wildl. soc. n ews 132 :4. leopold, a 1933. gam e manag ement. new york: charles scribner's sons. owen, 0 s 1975. natural resource conservation. 2nd ed . new york: macmillan . peterle, t j 1975. comments on scott's views on the training of wildlife stud ents. wildl. soc. bull. 3(2): 64-65. rongstad, 0 j and j j hickey. 1972. survey of qualifications of graduate students enrolled in wildlife programs fall of 1971. wildl. soc. ne ws 143: 45. sanderson, g c 1975. comments on scott 's views on th e training of wildlife students. wildl. soc. bull. 3(2): 65-67 . scott, t a 1975. the training role of co-operative units . wildl. soc. bull. 3(2): 59-62. sigler, w f 1975. recommended: b.s . degree for state wildlife law-enforcement officers . wildl . sol. bull. 3(4) : 173-175. trippensee, r e 1948 . wildlife management . vol. 1. new york: mcgraw-hill. warren, a and f b goldsmith eds . 1974. conservation in practice . london: john wiley and sons. 196 page 1 page 2 page 3 page 4 page 5 page 6 page 7 article information authors: martinette kruger1 melville saayman1 affiliation: 1tourism research in economic environs and society, north-west university, potchefstroom campus, south africa correspondence to: martinette kruger postal address: private bag x6001, potchefstroom 2520, south africa dates: received: 21 sept. 2012 accepted: 03 dec. 2013 published: 24 june 2014 how to cite this article: kruger, m. & saayman, m., 2014, ‘the determinants of visitor length of stay at the kruger national park’, koedoe 56(2), art. #1114, 11 pages. http://dx.doi.org/10.4102/ koedoe.v56i2.1114 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. the determinants of visitor length of stay at the kruger national park in this original research... open access • abstract • introduction    • literature review • research method and design    • the questionnaire    • sampling method and survey    • statistical analysis • results    • profile of respondents    • factor analysis    • poisson and negative binomial regression analyses       • northern region       • southern region • trustworthiness • discussion    • limitations    • recommendations • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ this study analysed the determinants of tourists’ length of stay at one of south africa’s oldest and largest national parks, the kruger national park. it took the different regions of this park into account and analysed visitors to the northern and southern regions separately to distinguish the different determinants of length of stay. the results showed clear differences between the determinants of length of stay for the two regions, indicating that for a destination with the size and scope of the kruger park, a regional approach should be followed to improve management and encourage visitors to stay longer.conservation implications: the northern and southern regions of the kruger national park differ significantly in terms of ecosystems, rainfall, climate and wildlife. from a tourism perspective, these regions should be managed separately taking the distinct differences of the two regions into consideration. different variables influence visitors’ length of stay in these two regions. conservation practitioners can use the results of this study to manage visitors to these areas. introduction top ↑ this study looked at what determines the length of time tourists stay at one of south africa’s oldest and largest national parks, the kruger national park. menezes, moniz and vieira (2008:205) state that ‘the overall impact of tourists in a given economy hinges on length of stay’. length of stay is of fundamental importance for tourist destinations, because the tourist product adapts itself to the amount of time the tourist has available at the destination and promotional campaigns must also be adjusted to the tourist’s decisions in this regard (martinez-garcia & raya 2008:1064). thrane and farstad (2009:2) and barros, butler and correia (2010:13) agree that length of stay is of major importance to any tourism destination, because longer stays are positively associated with the total earnings from tourist activities and with higher bed-occupancy rates. barros and machado (2010) posit that: tourist destinations should aim to attract tourists who wish to stay longer, particularly during off-peak seasons, since tourists who visit only for short periods tend to stay centrally and visit only the major tourist attractions. (p. 2) longer-stay tourists, on the other hand, visit a wider range of attractions, explore more peripheral regions and generate more diverse economic, social and environmental impacts. alegre and pou (2006) point out that: length of stay at a holiday destination, a basic characteristic of the holiday, has important repercussions for the tourist destination since it affects occupancy rates and final income derived from tourists (in other words, the economic impact and value of tourists). (p. 1352) identifying the factors that make tourists stay longer is therefore extremely beneficial to any destination seeking to increase visitor spending (peypoch et al. 2011:1). given that the topic is so important, there were several strong motives for researching length of stay and for choosing the kruger national park as a case study. firstly, alegre and pou (2006) claim that most studies on tourism demand fail to pay attention to length of stay, at least at a micro-economic level, where the researcher can control for individual heterogeneous behaviour. in addition, even though most tourism economists and managers agree that length of stay has a direct impact on spending (see cannon & ford 2002; jang, bai & o’leary 2004; kozak, gokovali & bahar 2008; mules 1998; seaton & palmer 1997; spotts & mahoney 1991; taylor, fletcher & clabaugh 1993, amongst others), little attention has been paid to this important topic (decrop & snelders 2004; gokovali, bahar & kozak 2007:737). secondly, the topic of tourists visiting a south african nature-based destination has not attracted more research thus far, despite the essential role these destinations play in tourism and conservation (holden 2008:180; south african national parks [sanparks] 2009:31). thirdly, the huge numbers of visitors to the kruger park make it south africa’s most profitable national park (sanparks 2009:19). as an all-inclusive holiday destination that provides a unique nature and leisure experience, the kruger park is a magnet for tourists, both domestic and international. it is a major export earner and constitutes an important part of the south african tourism industry (cook, yale & marqua 2010:212; saayman & saayman 2008; uysal, mcdonald & martin 1994:18). the park attracts over one million visitors per annum and is one of the top five international tourist destinations in the country (sanparks 2009; van der merwe & saayman 2008:154). currently, 80% of sanparks revenue comes from kruger park accommodation and admission fees (mabunda & wilson 2009:118). the findings of this study, a contribution to the still small literature on the topic, will be valuable in helping to sustain this growth and ensure long-term sustainability and competitiveness. finally, the size of the kruger park also makes it an ideal subject for this research, as it allows for comparison, within one park, of different tourist motives for length of stay. this is one of the largest game reserves in the world, covering approximately 20 000 km2, and extending 350 km from north to south and 60 km from east to west; an area larger than, for example, israel, wales or the netherlands. the kruger park includes two climatic transitional zones, namely the tropical and subtropical north and the temperate south; each with a clear rainfall gradient (lower rainfall in the south compared to the north) (mabunda, pienaar & verhoff 2003:16). from an ecological point of view, it is divided into four regions, namely the far northern and northern (shingwedzi, pafuri and punda maria), central (mopani, satara, orpen and letaba) and southern (berg-en-dal, crocodile bridge, pretoriuskop, biyamiti, malelane and lower sabie) (see figure 1), each with its diverse ecosystem, vegetation and wildlife. from a tourism perspective, it is divided into northern and southern regions and tourists are attracted to these regions for different reasons, depending on what they want to see and do. notable differences between the two regions include the number of rest camps and accommodation available (e.g. the southern region has more rest camps), accessibility (e.g. certain parts of the northern region are only accessible by 4×4s and there are fewer entry gates to this region), as well as wildlife and vegetation found (e.g. owing to a higher vegetative biomass to game ratio, the central and southern regions have more game and are known for sightings of the big five, whilst the northern region is more known for its large hippo, elephant and wild dog populations) (sa venues 2013; smit, grant & whyte 2007:227; tinker & tinker 2011:4). figure 1: map of the kruger national park. tourists’ length of stay in the kruger park, and consequently the determinants thereof, will therefore depend on their specific choices. taking this into consideration, this study analysed the northern and southern tourists separately so as to distinguish between the different determinants. the results of the analysis made it possible to propose strategies to increase the amount of time tourists spend in the kruger park. literature review how long to spend on a visit is one of the main considerations in peoples’ decision-making processes when making holiday plans, so destination managers need to know how to intervene in this process so as to encourage them to stay longer and thus spend more (alegre & pou 2006; peypoch et al. 2011). tourists weigh up the benefits of different holiday destinations, assessing the cost of each and the length of stay they can afford, financially and time-wise (alegre & pou 2006:1343). the chosen length of stay may reflect the attractiveness of a destination, but several other factors may come into play, as gokovali et al. (2007) indicate: the amount of free time the tourist has available, the availability of flexible package tour deals, the level of prices, the number of people in the party, and familiarity with the destination, to name but a few. (p. 737) the length of a holiday, like the decision to take a trip or the choice of a particular destination, may therefore be explained by a combination of internal and external factors, as shown in figure 2. internal factors may be personal and family characteristics, as well as economic factors such as the tourist’s level of income (alegre & pou 2006; machado 2010). external variables may include destination-specific (in this case park-specific) attributes such as destination image, type of destination, activities and amenities on offer or the price of the holiday (saayman & saayman 1997; machado 2010; kruger et al. 2012). figure 2: factors influencing visitors’ length of stay at a destination (national park). once the combination of factors that influence length of stay have been determined, marketing strategies can be planned that will encourage longer stays at a destination so that economic benefits can be maximised (barros & machado 2010:2; gokovali et al. 2007:737; menezes et al. 2008:206). menezes et al. (2008) state that: uncovering the determinants of length of stay is critical to the design of marketing policies that promote longer stays, associated with higher occupancy rates and revenue streams: an enterprise ever more valuable given the increasingly pervasive pattern of shorter length of stays. (p. 207) length of stay can also be an indicator of the profile of tourists visiting a particular destination and their propensity to spend whilst on holiday; information regarding their length of stay can consequently be used to increase tourist spending (davies 2002; gokovali et al. 2007; legohérel 1998; mok & iverson 2000). taking the length of stay into account can help in the effective management and planning of tourism destinations (barros & machado 2010; ritchie & crouch 2003). visitors who stay longer experience more and the destination earns more as they are more likely to become aware of and use facilities and services at the location where they are staying and in surrounding regions (gokovali et al. 2007; kozak 2001). saarinen (2005) points out that identifying the determinants of length of stay is important for research on sustainable tourism because: they are useful in forecasting tourists’ on-site time and, concomitantly, the stress on local resources caused by tourism activity; an important issue for analysing carrying capacity, especially at nature-based tourism destinations such as the kruger park. (p. 35) however, despite the numerous benefits associated with identifying the determinants of length of stay, studies of this variable in tourism are rare (barros & machado 2010:3), especially within a nature-based context. those available are mainly descriptive – mostly of popular ‘sea-and-sun’ destinations – and they analyse the way different tourist profiles or types of trip affect the length of stay (see alegre & pou 2006; barros & machado 2010; barros et al. 2010; gokovali et al. 2007; menezes et al. 2008; martinez-garcia & raya 2008; oppermann 1995, 1997; seaton & palmer 1997; sung et al. 2001). with regard to socio-demographic determinants, collectively these studies found that older, male and married tourists tend to stay longer at a destination than tourists with a higher level of education and a high-income profession (alegre & pou 2006:1349–1352; alegre, mateo & pou 2011; barros & machado 2010:10–13; barros et al. 2010:18–20; gokovali et al. 2007:742–744; martinez-garcia & raya 2008:1070–1073; menezes et al. 2008:214–218; peypoch et al. 2011). some specific findings were that visitors aged 25–34 years had a lower probability of long stays than those aged 35–44 years (gokovali et al. 2007:742–744) and that as the size of the party and daily expenditure increased, the probability of staying longer decreased (alegre & pou 2006:1349–1352). based on this, the following hypothesis was formulated: • hypothesis 1 (socio-demographic characteristics): the length of stay is determined by individual socio-demographic characteristics such as home language, marital status, level of education, annual gross income and province of origin. with regard to behavioural determinants, studies show that tourists who travel shorter distances stay a shorter time than those who travel longer distances. travel motives (leisure, visiting friends or relatives, business and others), repeat visits and an increased number of tourist attractions visited all increased the expected length of stay (menezes & moniz 2006:10–12; menezes et al. 2008:214–218). gokovali et al. (2007:742–744) found that tourists who choose an ‘all-inclusive’ type of holiday tend to have a lower probability of a long stay than those who choose a ‘half-board’ type of accommodation and that as the number of previous visits to the destination increases, the probability of a long stay increases; however, as the daily expenditure increases, the probability of a long stay decreases. when it comes to a destination’s image, machado (2010) reveals that tourists visiting destinations with a better reputation tend to stay longer. other determinants, such as nationality, type of flight, environmental initiatives, repeat visits, word-of-mouth recommendations, as well as quality and hospitality also tend to encourage longer stays (alegre & pou 2006: 1349–1352, 2011; barros & machado 2010:10–13; gokovali et al. 2007:742–744; menezes & moniz 2006:10–12; menezes et al. 2008:214–218; peypoch et al. 2011). based on this, the second hypothesis was formulated: • hypothesis 2 (behavioural characteristics): the length of stay is determined by individual behavioural characteristics such as mode of transport and travel motives. the findings of the abovementioned studies show collectively that determinants of length of stay are destination specific, but that tourists’ socio-demographic characteristics and economic factors are also significant determinants. furthermore, it is important to note that tourists cannot be considered homogeneous in terms of the effects of different factors on how long they decide to stay. given the size of the kruger park and the variety of its offerings, the determinants could differ for tourists visiting the northern and southern regions. this led to the third hypothesis: • hypothesis 3 (destination characteristics): the length of stay is determined by the destination characteristics (northern and southern region of the park – for example, seeing certain types of animals found in each region) experienced and valued by the tourist. research method and design top ↑ the questionnaire the questionnaire used during the two surveys (north and south) consisted of three sections: section a captured the respondents’ demographic details, including language, gender, age, race, and marital status, country of residence, province, highest qualification and occupation, whilst section b captured economic information such as size of travel group, number of people paid for, the type of accommodation used, the number of nights spent at the park, the number of visits over the last 3 years and spending dynamics. the spending questions were detailed by spending category, including entrance and conservation fee, accommodation, food and beverages, shopping, recreation and transport. section c captured the respondents’ reasons for visiting the park, where 12 items were listed and respondents were asked to rate the importance of each item on a five-point likert scale of importance: 1 = not at all important, 2 = slightly important, 3 = important, 4 = very important and 5 = extremely important (cf. kruger, saayman & manners 2012). this section also captured respondents’ perception of the value of the big five (i.e. the five most difficult african animals to hunt on foot: lion, african elephant, cape buffalo, leopard and rhinoceros). for the purpose of this research, the information from all three sections was included in the analyses. sampling method and survey the survey sample consisted of respondents to this questionnaire, which was distributed to visitors staying in the kruger park from 24 june 2011 to 02 july 2011 (northern region, winter survey) and from 27 december 2011 to 04 january 2012 (southern region, summer survey). the rest camps surveyed for the northern region were olifants, letaba, mopani and punda maria and for the southern region satara, skukuza, lower sabie and berg-en-dal. fieldworkers distributed questionnaires in the evenings and collected the questionnaires later during the same evening or early the next morning. a convenience sampling method was followed and questionnaires were therefore distributed to available overnight visitors (camping and chalet). only one questionnaire per group was allowed and only adults (18 years and above) were allowed to complete the questionnaire. the latest available sanparks statistics for 2009 were used to calculate the proportion of responses that would constitute a representative sample. according to these statistics, 384 249 (n) tourists stayed for at least one night in the park in 2009 (stevens 2010). furthermore, du plessis, saayman and erasmus (2010) state that the average travelling group of tourists to the park was 3.4 persons. the total population (n) was divided by 3.4 and this resulted in 112 132 tourist groups (n). the minimum number of completed questionnaires sufficient for this study would be 399 (israel 2009). of the total of 853 questionnaires collected in the two surveys, 410 had been fully completed, 175 from the northern region and 235 from the southern region. only fully completed questionnaires were used in the analysis to ensure that the important socio-economic and behavioural variables were all included. statistical analysis microsoft excel was used to capture the data and ibm spss statistics 20 and r 2.14.1 to analyse it. the analysis was conducted in two stages. firstly, data were captured in microsoft excel and analysed using spss (2013). factor analyses were conducted to determine respondents’ motives for travelling to the northern and southern regions of the kruger national park. to determine whether a factor analysis could have been conducted on the motivational data variables, the barlett’s test of spehricity and kaiser-meyer-olkin (kmo) measure of sampling adequacy was performed on both the expectations, as well as the experience scales. according to barlett (1954), a factor analysis is appropriate when the result of the barlett test for sphericity indicates significance, in other words p < 0.05. in agreement, kaiser (1970, 1974) explains that the minimum value of the kmo (the measure of sampling adequacy) for a factor analysis to be performed should be 0.6. in order to determine the smallest number of factors from the respective data variables, the pattern matrix of the principle component factoring extraction technique was applied. the kaiser normalisation (eigenvalues above 1.0 or more) guided the decision on the amount of facto rs retained. to name the factors, according to dancey and reidy (2004:431), a decision should be made on how strong the factor loadings must be to be included in a factor, but this tends to be arbitrary and varies amongst authors. all items with a factor loading above 0.4 were considered as contributing to a factor and all with loadings lower than 0.4 as not correlating significantly with this factor (steyn 2000). in addition, any item that cross-loaded onto two factors, with factor loadings greater than 0.4, was categorised in the factor where interpretability was best. the internal consistency of each factor was also calculated by means of the reliability coefficient (cronbach’s alpha). only reliability coefficients above 0.6 were considered as acceptable for the study because a reliability coefficient below 0.6 indicates that the scale has poor reliability and unsatisfactory internal consistency (malhotra 2007:285; zikmund et al. 2010:306). additionally the inter-item correlations were also calculated as another reliability measure, which, as recommended by briggs and cheek (1986), should be between 0.2 and 0.4. secondly, the dependent (predicted) variable was understood to be length of stay, defined as the number of days at the park; accordingly, it is a non-negative count variable. a poisson regression was applied to identify the determinants of length of stay for both the northern and southern region. the link function for a poisson regression model is the log link function and the model (with a log link) used in this research is as follows: [eqn 1]the mean satisfies the exponential relationship: [eqn 2] where: a one-unit increase in x has a multiplicative impact of eβ on μ. the mean of y at x + 1 equals the mean of y at x multiplied by eβ. if β = 0, then eβ = e0 = 1 and the multiplicative factor is 1. then, the mean of y does not change as x changes. if β > 0, then eβ > 1, and the mean of y increases as x increases. if β < 0, the mean decreases as x increases. (agresti 2007:75) a poisson regression model was also used to find the determinants of length of stay for the southern region; however, the over-dispersion parameter was 2.45. a negative binomial regression was used because this model takes this over-dispersion into account. according to agresti (2007:81), the negative binomial is another distribution that is concentrated on the non-negative integers. unlike the poisson, it has an additional parameter such that the variance can exceed the mean. the negative binomial distribution has: [eqn 3] the index d, which is non-negative, is called a dispersion parameter. the negative binomial distribution occurs as a type of mixture of poisson distributions. greater heterogeneity in the poisson means results in a larger value of d. as d → 0, var(y) → μ and the negative binomial distribution converges to the poisson distribution. the further d falls above 0, the greater the over-dispersion relative to poisson variability (agresti 2007:81). results top ↑ this section presents the profile of respondents as well as the results of the factor analysis (travel motives), the poisson regression analysis (northern region) and the negative binomial regression analysis (southern region). profile of respondents table 1 shows the profile of the respondents in each of the two regions surveyed in 2011–2012. there are notable similarities between the northern and southern region tourists, especially in terms of home language, average age, marital status, level of education, mode of transport, length of stay and being holders of a wild card (a loyalty card that can be used at all 22 national parks in south africa and offers the benefits of discounted entrance fees and a contribution to conservation). the main differences between the two profiles are in terms of province of origin, average spending and financial responsibility, with the northern region attracting more visitors from the western cape, who are financially responsible for more people during their trip and, consequently, have a higher average spending than tourists who visit the southern region. table 1: comparison of profiles of respondents staying in the northern and southern regions of the kruger national park, 2011–2012. factor analysis the pattern matrix of the principal component factor analyses using an oblimin rotation with kaiser normalisation identified four motivational factors each for the northern and the southern regions. these were labelled according to similar component items (table 2). these factors accounted for 58% and 64% of the total variance for north and south, respectively. the average of all items contributing to a specific factor revealed factor scores that interpret the factor to the original five-point likert scale. all factors indicated very good convergent validity with cronbach alphas above 0.6 and inter-item correlations of between 0.31 and 0.63. in the case of the northern region, ‘to photograph animals and plants’ was not included in the factor analysis, because it did not have a loading greater than 0.4. the kaiser-meyer-olkin measure of sampling adequacy of, respectively, 0.70 and 0.75 indicated that patterns of correlation were relatively compact and yielded distinct and reliable factors (field 2005:640). in both cases, bartlett’s test of sphericity also reached statistical significance (p < 0.001), supporting the factorability of the correlation matrix (pallant 2007:197). as table 2 shows, the following reasons for visiting the northern region of the park were identified: ‘escape’ (factor 1), ‘exploration’ (factor 2), ‘education’ (factor 3) and ‘value and amenities’ (factor 4). with a mean value of 4.391, ‘education’ was found to be the most important motive for travelling to the northern region, followed by ‘exploration’ (3.509) and ‘escape’ (3.293). ‘value and amenities’ (2.860) was rated the least important motive. for the southern region, the motives and the grouping of the items and their importance were different. ‘escape’ (factor 1), ‘group togetherness and exploration’ (factor 2), ‘photography and spiritual fulfilment’ (factor 3) and ‘value and amenities’ (factor 4) were identified as the reasons for travelling to this part of the park. ‘photography and spiritual fulfilment’ (4.027) was rated as the most important motive, followed by ‘value and amenities’ (3.506), ‘escape’ (3.420) and ‘group togetherness and exploration’ (3.177). it is clear from these results that the tourists in this sample visited the two areas of the park for somewhat different reasons and that the characteristics of the two areas influence these motives. table 2: results of the factor analyses – motives of visitors to the kruger national park. poisson and negative binomial regression analyses as table 3 shows, the dummy variables (socio-demographic and behavioural variables, as well as the travel motives indicated in table 2) for both the northern and southern regions were coded 1 and 0 to be included in the poisson regression and the negative binomial regression analyses. table 3: relationship between variables and respondents’ length of stay. northern region a poisson regression model was performed with all the independent variables indicated in table 3, after which a stepwise selection procedure was carried out to decrease the number of variables. the null hypothesis was that the poisson regression model provided an adequate fit to the data. the residual deviance is a measure of how much the data deviates from the model, so the larger the residual deviance is, the worse the fit. the residual deviance here was 176.55, to be compared with a chi-square distribution with 166 degrees of freedom. the p-value was 0.274, which was large enough that the null hypothesis did not have to be rejected. the model therefore appeared to be adequate. as shown in table 4, gauteng as a province of origin (p = 0.023), total spending (p = 0.002), decision to visit made long in advance (p = 0.042), being a wild card holder (p = 0.004), lion and leopard as ‘must-see’ big five animals (respectively p = 0.010 and p = 0.034) and donating money for conservation (p = 0.010) had a statistical significant influence on length of stay. with regard to socio-demographic determinants, based on the signs of the coefficients, a person coming from gauteng would have a predicted stay at the park 0.866 times shorter than that of the person not coming from this province. therefore, coming from gauteng decreased the length of stay when controlling for the other variables. the reason for this could be that, for the tourists travelling from this province, the park is close and they can therefore travel to the park more often and, consequently, do not have to stay long in the park. tourists with a higher average spending per trip tended to have a longer length of stay. these tourists would have a predicted stay at the park 1.000 times longer than those with a lower spending per trip. these results validated hypothesis 1 that stated that the length of stay is determined by individual socio-demographic characteristics. table 4: results of the stepwise poisson regression analysis. with regard to behavioural determinants, tourists who made their decision to travel to the park well in advance would have a predicted stay 1.191 times longer than that of the person who made a spontaneous decision. in addition, tourists who had a wild card would have a predicted stay at the park 1.711 times longer than that of a person who did not have a wild card. having a loyalty card therefore increased the length of stay when controlling for other variables, validating hypothesis 2 that individual behavioural characteristics influence length of stay. with regard to the popularity of the big five, tourists who most wanted to see lion and leopard would have predicted stays 1.314 and 1.240 times longer, respectively, than those who did not want to see these animals. this could be because the park is synonymous with the lion, its image being visible in the park’s logo and used extensively in marketing and branding. leopards are rarely sighted – this is the big five animal that most often eludes visitors. this makes them sought-after and visitors perhaps stay longer to increase their chances of seeing one. another reason may be that june and july (the winter months) are the prime game-viewing time in the park. visitors who donated money for conservation causes tended to stay fewer days at the park and would have a predicted stay 0.832 shorter than that of a person who did not support these causes. therefore, making the decision to travel well in advance and wanting to see lion and leopard increased the length of stay at the park when controlling for other variables, whilst donating money for conservation decreased the length of stay. regarding the big five as important to extremely important (p = 0.114) was not statistically significant. this determinant did, however, still have an important effect on length of stay. tourists who considered seeing the big five to be very important would have a predicted stay 1.103 times longer than those for whom this was less important. these results supported hypothesis 3 that destination characteristics also influenced length of stay. southern region here the residual deviance was 218.89, to be compared with a chi-square distribution with 222 degrees of freedom. the p-value was 0.546, which was large enough that the null hypothesis did not have to be rejected. the model therefore appeared to be adequate. as table 5 shows, afrikaans as home language (p = 0.036), number of people paying for (p = 0.004), decision to visit made long in advance (p = 0.006), sedan as mode of transport (p = 0.004), total spending (p = 0.000), being a wild card holder (p = 0.000) and being motivated by ‘escape’ (p = 0.043) had a statistically significant influence on length of stay. with regard to socio-demographic determinants, afrikaans-speaking tourists and those who used a sedan to travel to the park would have a predicted stay respectively 0.840 and 0.755 times shorter than that of either english-speaking or foreign language tourists and tourists who used other modes of transport to travel to the park, validating hypothesis 1. with regard to behavioural determinants, travelling with a larger group had a negative influence on length of stay and visitors who travelled in larger groups to the park would have a predicted stay 0.932 shorter than that of tourists who travelled in smaller groups. here, the result was similar to that obtained for the northern region: making the decision to travel to the park well in advance had a positive influence on length of stay and these tourists would have a predicted stay 1.275 longer than that of the tourists who made a spontaneous decision. tourists motivated more by the motive ‘escape’ also tended to stay longer at the park (1.088 times longer). in addition, those with a higher total average spending and who were wild card holders tended to stay longer at the park (respectively 1.000 and 2.048 times longer). these results confirmed hypothesis 2. the other variables did not have a statistically significant influence on length of stay; however, tourists who were local residents from south africa tended to stay 1.276 times longer than tourists travelling from outside the borders of the country. like the tourists who travelled with a sedan to the park, those who used a 4×4 to travel tended to have a shorter length of stay (0.872 times shorter than that of visitors who used other modes of transport). contradicting the results obtained for the northern region, wanting to see a lion or leopard had a negative influence on length of stay (respectively 0.877 and 0.892 shorter than the stay of tourists who did not particularly want to see them), rejecting hypothesis 3 that destination characteristics influenced length of stay. trustworthiness top ↑ the following steps, as proposed by field (2003) as well as zikmund et al. (2010) were followed to design and validate the questionnaire as well as the results:• content validity: an in-depth literature analysis was performed to identify the relevant motivational factors for travelling to the kruger national park. the questionnaire was based on van der merwe and saayman (2008), saayman and saayman (2009) and kruger and saayman (2010). • face validity: statistical consultation services advised on the formulation of the statements as well as the measuring scales used. the park manager also gave their opinion on the included items and whether it captured the essence of the study. • construct validity: factor analyses were performed on the motivational factors for travelling to the northern and southern region of the park in order to determine the degree to which the statements measures what it claims, or purports, to be measuring (please see the ‘statistical analysis’ and ‘factor analysis’ sections for more detail). • reliability: to test the reliability of the identified factors, reliability coefficients (cronbach’s alpha) and inter-item correlations were calculated (please also see the ‘statistical analysis’ and ‘factor analysis’ sections for more detail). discussion top ↑ this article identified the determinants of length of stay for tourists at a nature-based destination in south africa, namely the kruger national park. the size, nature and characteristics of this park made it possible for the first time to differentiate between determinants of length of stay for different regions, namely the northern and southern regions of the kruger park. the results revealed clear differences between the determinants of length of stay for the two regions. the general conclusion was that tourists travelling to the northern region with a higher total average spending, who made their decision to visit the kruger park long in advance, who were wild card holders and who were keen to see lion and leopard whilst on holiday were likely to stay longer in this region of the park. a different set of determinants were found to affect the length of stay in the southern region: here, afrikaans-speaking tourists who travelled in smaller groups, with a higher total average spending, who made their decision to travel to the park long in advance, who were wild card holders and who were motivated by the need to escape, were likely to stay longer. this study corroborates the finding by alegre and pou (2006) that travelling in larger groups has an inverse influence on length of stay: the larger the group, the shorter the stay. in addition, the results support the finding by gokovali et al. (2007) that tourists who spend more stay longer. however, it contradicts the findings by menezes and moniz (2006) and menezes et al. (2008) that tourists who travel shorter distances stay longer than those who travel further. the results supported all three of the study’s hypotheses. firstly, for both the northern and southern regions of the park, tourists’ individual socio-demographic characteristics (total spending, province of origin, group size and home language) influenced the length of stay, validating hypothesis 1. the socio-demographic determinants were, however, different for the two regions. secondly, behavioural variables (decision-making, mode of transport, being loyalty members, desire to see the big five and not donating to conservation causes) explained the length of stay, validating hypothesis 2. here again the determinants for the two regions were different; however, behavioural characteristics seemed to have a stronger influence on length of stay in both regions than socio-demographic characteristics. thirdly, the length of stay was also determined by the characteristics of the two regions (the two destinations), such as the different likelihood of seeing the big five, validating hypothesis 3. it was clear from the results that the length of stay was specific to each of the two tourism destinations, supporting previous research by alegre and pou (2006), alegre, mateo and pou (2011), barros and machado (2010), barros et al. (2010), gokovali et al. (2007), martinez-garcia and raya (2008), menezes and moniz (2006), menezes et al. (2008) and peypoch et al. (2011). the determinants of length of stay are also more internal rather than external factors (see figure 2). however, the study makes a further contribution: on the basis of its findings for the two regions, it cautions marketers to approach the park from a regional perspective (i.e. making a distinction between two different parts of the same national park) rather as a whole. this might also apply to other destinations in the world that consist of regions offering different products. limitations one apparent limitation was that the survey was conducted separately during the winter (northern region) and summer (southern region). the type of visitor differs during these months and future research should perhaps consider conducting a survey in both regions at the same time either during the winter or summer in order to accurately compare results. recommendations the results of this study provide a clear, focused view of the reasons for tourists’ length of stay at the nature-based destination analysed. this method furthermore takes into account that the characteristics of the different regions themselves may influence length of stay. from a methodological point of view, a regional approach is therefore suitable for analysing a destination with multifaceted characteristics, such as the kruger national park, where tourists also resemble homogeneous tendencies. as it enables a concise comparison of the effect on length of stay of the same variables in different regions, it should prove valuable to destination marketers, planners and managers, as well as academics. findings produced by this method will enable them to differentiate between and compare different regions and plan their strategies accordingly to encourage visitors to stay longer.it is, of course, not possible to control all the determinants influencing length of stay in the kruger park. however, it is possible to concentrate on the important ones and to highlight them in advertising and promoting the different regions of the park. on the basis of the findings of this study, a diversified strategy is recommended. if the park is to increase tourists’ length of stay in the northern region, its marketing campaigns should focus on the big five, especially lion and leopard. more information about these animals, their behaviour and preferred terrain should be made available to tourists at the different rest camps and in the game drives and guided walks. this might help to attract longer-staying visitors. to increase length of stay in the southern region, marketing campaigns should focus on people’s desire to escape from the routine of their daily lives and emphasise that the park is the ideal relaxing and breakaway holiday destination. the benefits of the current loyalty card (the wild card) should also be marketed more intensively, because this will encourage tourists to stay longer irrespective of which of the two regions they visit. conclusion top ↑ in summary, because a combination of socio-demographic and behavioural determinants influences length of stay, and length of stay may differ for different regions of a destination, at a nature-based destination like the kruger park a regional analysis is advisable and a diversified marketing strategy will be more likely to increase the average length of stay than an undifferentiated mass strategy. as more research is needed to confirm these findings, it is recommended that a similar method be applied to other nature-based destinations, as well as ‘sea-and-sun’ destinations. acknowledgements top ↑ competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions m.k. (north-west university) was the lead author on the article, conceptualised the idea, conducted the survey, analysed the data together with the statistical consultation services at the north-west university and did the final technical editing. m.s. (north-west university) focused on the findings and implications, as well as general flow of the research. references top ↑ agresti, a., 2007, an introduction to categorical data analysis, 2nd edn., john wiley and sons, gainesville. http://dx.doi.org/10.1002/0470114754 alegre, j. & pou, l., 2006, ‘the length of stay in the demand for tourism’, tourism management 27(1), 1343–1355. http://dx.doi.org/10.1016/j.tourman.2005.06.012 alegre, j., mateo, s. & pou, l., 2011, ‘a latent class approach to tourists’ length of stay’, tourism management 32(3), 555–563. barros, c.p. & machado, l.p., 2010, ‘the length of stay in tourism’, annals of tourism research 1(1), 1–15. barros, c.p., butler, r. & correia, a., 2010, ‘the length of stay of golf tourism: a survival analysis’, tourism management 31(1), 13–21. http://dx.doi.org/10.1016/j.tourman.2009.02.010 bartlett, m.s., 1954, ‘a note on the multiplying factors for various chi-square approximations’, journal of the royal statistical society 16(series b), 296–298. briggs, s.r. & cheek, j.m., 1986, ‘the role of factor analysis in the development and evaluation of personality scales’, journal of personality 54(1), 106–149. cannon, t.f. & ford, j., 2002, ‘relationship of demographic and trip characteristics to visitor spending: an analysis of sports travel visitors over time’, tourism economics 8(3), 263–271. http://dx.doi.org/10.5367/000000002101298106 clark, l.a. & watson, d., 1995, ‘constructing validity: basic issues in objective scale development’, psychological assessment 7(3), 309–319. http://dx.doi.org/10.1037/1040-3590.7.3.309 cook, r.a., yale, l.j. & marqua, j.j., 2010, tourism: the business of travel, 4th edn., pearson education, upper saddle river. dancey, c.p. & reidy, j., 2004, statistics without maths for psychology: using spss for windows, 3rd edn., prentice hall, new york. davies, l.e., 2002, ‘consumers’ expenditure on sport in the uk: increased spending or underestimation?’, managing leisure 7(2), 83–102. http://dx.doi.org/10.1080/13606710210137237 decrop, a. & snelders, d., 2004, ‘planning the summer vacation: an adaptable process’, annals of tourism research 31(4), 1008–1030. http://dx.doi.org/10.1016/j.annals.2004.03.004 downward, p. & lumsdon, l., 2004, ‘tourism transport and visitor spending: a study in the new york moors national park, uk’, journal of travel research 42(4), 415–420. http://dx.doi.org/10.1177/0047287504263038 du plessis, l., saayman, m. & erasmus, l.j.j., 2010, ‘a marketing and management analysis of overnight and day visitors to the kruger national park december 2009/10’, unpublished report for the institute for tourism and leisure studies, north-west university, potchefstroom. field, a., 2005, discovering statistics using spss, sage, thousand oaks. gokovali, u., bahar, o. & kozak, m., 2007, ‘determinants of length of stay: a practical use of survival analysis’, tourism management 28(1), 736–746. http://dx.doi.org/10.1016/j.tourman.2006.05.004v holden, a., 2008, environment and tourism, 2nd edn., routledge, london. israel, g.d., 2009, determining sample size, viewed 12 july 2010, from http://www.edis.ifas.ufl.edu/pdffiles/pd/pd00600.pdf jang, s., bai, b., hong, g. & o’leary, j.t, 2004, ‘understanding travel expenditure patterns: a study of japanese pleasure travelers to the united states by income level’, tourism management 25(3), 331–341. http://dx.doi.org/10.1016/s0261-5177(03)00141-9 kaiser, h., 1970, ‘a second generation little jiffy’, psychometrika 35(4), 401–415. kaiser, h., 1974, ‘an index of factorial simplicity’, psychometrika 39(1), 31–36. kozak, m., 2001, ‘comparative assessment of visitors’ satisfaction with destinations across two nationalities’, tourism management 22(4), 391–401. http://dx.doi.org/10.1016/s0261-5177(00)00064-9 kozak, m., gokovali, u. & bahar, o., 2008, ‘estimating the determinants of tourist spending: a comparison of four models’, tourism analysis 13(2), 143–155. kruger, m., saayman, m. & saayman, a., 2010, ‘expenditure-based segmentation of visitors to tsitsikamma national park’, acta commercii 10(1), 137–149. kruger, m. & saayman, m., 2010, ‘travel motivation of tourists to kruger and tsitsikamma national parks: a comparative study’, south african journal of wildlife research 40(1), 93–102. kruger, m., saayman, m. & manners, b., 2012, ‘determinants of visitor expenditure at the tsitsikamma national park’, journal of economics and financial science 5(1), 11–30. krugerpark.com, 2011, full map of the kruger national park, viewed 07 august 2013, from http://www.krugerpark.com/maps/full-map-of-the-kruger-national-park/ legohérel, p., 1998, ‘toward a market segmentation of the tourism trade: expenditure levels and consumer behaviour instability’, journal of travel and tourism marketing 7(3), 19–39. http://dx.doi.org/10.1300/j073v07n03_02 leones, j., colby, b. & crandall, k., 1998, ‘tracking expenditures of the elusive nature tourists to south-eastern arizona’, journal of travel research 36, 56–64. http://dx.doi.org/10.1177/004728759803600306 mabunda, d.m., pienaar, d.j. & verhoff, j., 2003, ‘the kruger national park: a century of management and research’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 3–21, island press, washington, dc. mabunda, d.m. & wilson, d., 2009, ‘commercialisation of national parks: south africa’s kruger national park as an example’, in j. saarinen, f. becker, h. manwa & d. wilson (eds.), sustainable tourism in southern africa, pp. 116–133, channel view publications, bristol. machado, l.p., 2010, ‘does destination image influence the length of stay in a tourism destination’, tourism economics 16(2), 443–456. http://dx.doi.org/10.5367/000000010791305554 malhotra, n.k., 2007, marketing research: an applied orientation, 5th edn., prentice hall, upper saddle river. martinez-garcia, e. & raya, j.m., 2008, ‘length of stay for low-cost tourism’, tourism management 29(1), 1064–1075. http://dx.doi.org/10.1016/j.tourman.2008.02.011 mehmetoglu, m., 2007, ‘nature-based tourists: the relationship between their trip expenditures and activities’, journal of sustainable tourism 15(2), 200–215. http://dx.doi.org/10.2167/jost642.0 menezes, a.g. & moniz, a., 2006, ‘determinants of length of stay: a parametric survival analysis’, working paper no. 19/2006, centro de estudos de economia aplica do atlântico, viewed 25 march 2014, from https://repositorio.uac.pt/handle/10400.3/1152 menezes, a.g., moniz, a. & viera, j.c., 2008, ‘the determinants of length of stay of tourists in the azores’, tourism economics 14, 205–222. mok, c. & iverson, t.j., 2000, ‘expenditure-based segmentation: taiwanese tourists to guam’, journal of travel research 16(1), 1–4. mules, t., 1998, ‘decomposition of australian tourist expenditure’, tourism management 19(30), 267–271. http://dx.doi.org/10.1016/s0261-5177(98)00014-4 oppermann, m., 1995, ‘travel life cycle’, annals of tourism research 22(3), 535–552. http://dx.doi.org/10.1016/0160-7383(95)00004-p oppermann, m., 1997, ‘first-time and repeat visitors to new zealand’, tourism management 18(1), 177–181. http://dx.doi.org/10.1016/s0261-5177(96)00119-7 pallant, j., 2007, spss survival manual: a step-by-step guide to data analysis using spss version 15, mcgraw-hill, new york. peypoch, n., randriamboarison, r., rasoamananjara, f. & solonandrasana, b., 2011, ‘the length of stay of tourists in madagascar’, tourism management 1(1), 1–6. ritchie, j.r.b. & crouch, g.i., 2003, the competitive destination, cabi publishing, cambridge. saarinen, j., 2005, ‘tourism in the northern wildernesses: wilderness discourses and the development of nature-based tourism in northern finland’, in c.m. hall & s. boyd (eds.), nature-based tourism in peripheral areas: development or disaster?, pp. 36–49, channel view publications, clevedon. saayman, m. & saayman, a., 2008, ‘estimating the economic contribution of visitor spending in the kruger national park to the regional economy’, journal of sustainable tourism 14(1), 67–81. http://dx.doi.org/10.1080/09669580608668592 saayman, a. & saayman, m., 1997, ‘die ekonomiese impak van toerisme op die suid-afrikaanse ekonomie’, south african journal of economic management sciences 21, 162–174. saayman, m. & saayman, a., 2009, ‘why travel motivation and socio-demographics matter in managing a national park’, koedoe 51(1), 9 pages. sa venues, 2013, kruger national park, viewed 07 august 2013, from http://www.sa-venues.com/game-reserves/mpl_kruger.htm seaton, a.v. & palmer, c., 1997, ‘understanding vfr tourism behaviour: the first five years of the united kingdom tourism survey’, tourism management 1(6), 345–355. http://dx.doi.org/10.1016/s0261-5177(97)00033-2 smit, i.p.j., grant, c.c. & whyte, i.j., 2007, ‘landscape-scale sexual segregation in the dry season distribution and resource utilisation of elephant in kruger national park, south africa’, diversity and distributions 13, 225–236. http://dx.doi.org/10.1111/j.1472-4642.2007.00318.x south african national parks, 2009, annual report 2008–2009, viewed 15 february 2010, from http://www.sanparks.org/about/annual/2009.pdf spotts, d.m. & mahoney, e.m., 1991, ‘segmenting visitors in a destination region based on the volume of their expenditure’, journal of travel research 29(4), 24–31. http://dx.doi.org/10.1177/004728759102900405 spss version 20, 2013, computer software, ibm, chicago. stevens, j., 2010, ‘number of visitors to sanparks’, 12 april, joeps@sanparks.org steyn, h.s., 2000, ‘practical significance of the difference in means’, south african journal of industrial psychology 26(3), 1–3. sung, h.h., morrison, a.m., hong, g.s. & o’leary, j.t., 2001, ‘the effects of household and trip characteristics on trip types: a consumer behavioural approach for segmenting the us domestic leisure travel market’, journal of hospitality and tourism research 25(1), 46–68. http://dx.doi.org/10.1177/109634800102500105 taylor, d.t., fletcher, r.r. & clabaugh, t., 1993, ‘a comparison of characteristics, regional expenditures and economic impact of visitors to historical sites with other recreational visitors’, journal of travel research 32(1), 30–35. http://dx.doi.org/10.1177/004728759303200105 thrane, c. & farstad, e., 2009, ‘domestic tourism expenditures: the non-linear effects of length of stay and travel party size’, tourism management 1(1), 1–7. tinker, a. & tinker, l., 2011, ‘kruger national park: guide and map’, andy tinker photography, witrivier, mpumalanga. van der merwe, p. & saayman, m., 2008, ‘travel motivations of tourists visiting kruger national park’, koedoe 50(1), 154–159. van der merwe, p., saayman, m. & krugell, w., 2007, ‘the determinants of spending by biltong hunters’, south african journal of economics and management science 10(2), 184–194. uysal, y., mcdonald, c.d. & martin, b.s., 1994, ‘australian visitors to us national parks and natural areas’, international journal of contemporary hospitality management 6(3), 18–24. http://dx.doi.org/10.1108/09596119410059209 zikmund, w.g., babin, b.j., carr, j.c. & griffin, m., 2010, business research methods, 8th edn., south western cencage learning, mason. article information authors: mahlomola e. daemane1 sarel s. cilliers2 hugo bezuidenhout1 affiliations: 1conservation services, south african national parks, south africa 2school of environmental sciences and development, north-west university, south africa correspondence to: mahlomola daemane postal address: po box 110040, kimberley 8306, south africa dates: received: 11 aug. 2010 accepted: 16 sept. 2011 published: 18 jan. 2012 how to cite this article: daemane, m.e., cilliers, s.s. & bezuidenhout, h., 2012, ‘classification and description of the vegetation in the spitskop area in the proposed highveld national park, north west province, south africa’, koedoe 54(1), art. #1020, 7 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1020 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. classification and description of the vegetation in the spitskop area in the proposed highveld national park, north west province, south africa in this original research... open access • abstract • introduction    • study area • methods    • stratification of sample sites    • sampling procedure    • data processing    • description of plant communities • results    • vegetation classification    • detailed description       • 1. acacia caffra – seriphium plumosum woodland       • 1.1. acacia robusta – melinis repens woodland       • 1.2. mundulea sericea subsp. sericea – eragrostis racemosa shrubland       • 1.3. acacia karroo – asparagus suaveolens shrubland       • 2. setaria sphacelata var. torta – eragrostis curvula grassland       • 3. cymbopogon pospischilii – elionurus muticus grassland    • ordination • discussion • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ the objective of the proposed highveld national park (hnp) is to conserve a considerable area of the poorly conserved rocky highveld grassland and dry sandy highveld grassveld of the western grassland biome in south africa. the park has not yet been proclaimed, but is currently under the management of the north west parks and tourism board. the main aim of this study was to classify and describe the vegetation in the spitskop area in the hnp. the areas affected by soil degradation were on the midslopes, footslopes, valley bottomland and the floodplains around the spitskop hill. the concentrated grazing around the spitskop area was also influenced by the existing dam in the floodplains. floristic and soil degradation data were collected and used to classify and describe the plant communities of the spitskop area. vegetation sampling was performed by means of the braun-blanquet method and a total of twenty plots were sampled. a numerical classification technique (twinspan) was applied to the floristic data to derive a first approximation of the main plant communities. further refinement was achieved by braun-blanquet procedures. the final results of the classification procedure were presented in the form of a phytosociological table, with three major communities and three subcommunities being described. canonical correspondence analysis was used to determine the direct correlation between plant communities and soil degradation types. soil compaction and sheet erosion were found to be the most significant variables determining plant community composition. rill and gully erosion were shown to be of lesser significance in explaining the variation in plant communities. conservation implications: grasslands are amongst the most threatened biomes in south africa, yet less than 1.3% are currently being conserved. the hnp has significant value for biodiversity conservation and the protection of this area will contribute to the preservation of the highly threatened highveld vegetation types. introduction top ↑ the proposal for the development of the highveld national park (hnp) dates back to1984, when the then department of agriculture decided to inspect all the agricultural campsites surrounding potchefstroom (north west province, south africa) with the aim of renting (daemane 2007). the investigation showed that the conservation value of the hnp grassland was far higher than its agricultural value and the thought of establishing a conservation area emerged. however, poorly conserved rocky highveld grassveld and dry sandy highveld grassveld (bredenkamp & van rooyen 1996a, 1996b) in the hnp are currently threatened by increasing urbanisation (daemane, cilliers & bezuidenhout 2010). on the urban fringe of potchefstroom, urban sprawl has not only transformed large natural areas but also fragmented existing plant communities extensively(cilliers, müller & drewes 2004). practices such as harvesting of trees for firewood, uncontrolled fires and overgrazing are some of the challenges facing the management of the hnp (daemane et al. 2010). a previous study on the phytosociology of the hnp described and subsequently classified the vegetation into nine plant communities (daemane et al. 2010). however, vegetation sampling for the spitskop area in the hnp was undertaken separately as some of the areas were characterised by soil degradation with very sparse vegetation cover. in that study soil degradation was not evident in the upper topographical positions such as the crest and the upper slopes of the spitskop hill (daemane et al. 2010).the sheet, gully and rill erosions were associated with the midslopes and footslopes of the spitskop area. studies on soil degradation found rills and gullies are rarely evident on gradients greater than 10° and typically occur on unconsolidated colluvial or alluvial materials (cobban & weaver 1993; liggitt 1988; snyman, van rensburg & opperman 1986). previous studies on soil degradation found soil erosion to be highly depended on land use, slope and climatic factors (garland 1987;liggitt 1988). brady (1993) found the influence of the slope to be strongly interrelated with factors such as soil type. in the spitskop area, the vegetation communities in the valley bottomland and footslopes were close to the spitskop dam and were influenced by overgrazing and trampling (daemane 2007). the main aim of the present study was therefore to classify and describe the vegetation in the soil-degraded spitskop area. an understanding of the plant communities and their associated habitats is of fundamental importance for conservation of vegetation and soil and compiling sound management and conservation strategies (e.g. rehabilitation and monitoring). study area the study focused on the spitskop area in the hnp, west of potchefstroom (26°42’45”s–26°43’21”s and 26°59’45”e–27°00’16”e; figure 1). the areas affected by soil degradation were on the midslopes, footslopes, valley bottomlands and the floodplains of the spitskop hill. the entire park occupies approximately 10 200 ha and lies between the n12 motorway between potchefstroom and klerksdorp in the south, and eleazer road in the north. the relatively heterogeneous geology in the hnp is represented by the witwatersrand and ventersdorp supergroups and the transvaal sequence, with isolated occurrences of archaean granites and karoo sequence sediments (daemane et al. 2010). the geology in the spitskop area comprises mostly rocky quartzite outcrops. soils are heterogeneous and range from sandy to clayey owing to great variation in the parent rock material (land type survey staff 1984). soil forms consist mainly of glenrosa, hutton and mispah (daemane 2007). mean annual rainfall in the area is approximately 600 mm per year, but can be as high as 900 mm in exceptional seasons (institute for soil, climate and water 2003). the area experiences great seasonal and daily variation in temperature: the summers are very hot (daily mean maximum temperatures may exceed 32 °c in january), whilst winters are mild to cold (mean minimum monthly temperatures are approximately 12 °c) (iscw 2003). the vegetation of the spitskop area is dominated by vangueria infausta subsp. infausta – loudetia simplex shrubland on the crest. diheteropogon amplectens – trychopogon spicatus grassland is associated with the upper midslopes (daemane et al. 2010). acacia caffra – setaria sphacelata var. sphacelata woodland is associated with the lower midslopes, but also extend to the footslopes and bottomlands (daemane et al. 2010). bredenkamp and van rooyen (1996a, 1996b) classified the hnp area as rocky highveld grassland and dry sandy highveld grassland. the bankenveld vegetation type (veld type 61), which acocks (1953) described as ‘false grassveld’, is dominant in the spitskop area. the climax veld type is open a. caffra woodland. figure 1: vegetation map of the plant communities of the highveld national park, potchefstroom, south africa, with the spitskop area highlighted in red. methods top ↑ stratification of sample sites vegetation sampling involved 20 relevés of 900 m2each, randomly placed within the stratified units. stratification was performed based on 1:50 000 aerial orthophotographs showing relatively homogeneous physiographic and physiognomic units. further stratification was carried out by recognising the terrain types associated with topographical positions. sampling procedure plant species were identified in each sampling plot and the cover abundance (barkman, doing & segal 1964) of each species was estimated visually. unknown specimens were identified in the kimberley south african national parks herbarium (ksan). plant nomenclature follows plants of southern africa (south african national biodiversity institute 2009). average height and canopy cover of tree, shrub and herbaceous strata were estimated and the data were used in the description of the plant communities. features of the habitat such as geology, soil forms (macvicar, loxton & lambrechts 1991), and rockiness of the soil surface were recorded for each relevé. classification of erosion type, such as sheet, rill and gully erosion, follows torrion (2002). data processing the twinspan classification (hill 1979), which is regarded as a very successful approach by many phytosociologists (bredenkamp & bezuidenhout 1995; cilliers 1998; mucina & van der maarel 1989), was used as a first approximation to analyse the floristic data. the twinspan classification was then refined further by the application of braun-blanquet procedures by means of the bbpc-program (bezuidenhout, biggs & bredenkamp 1996). canonical correspondence analysis, using the canoco software version 2.1 (ter braak 1986), was employed to determine the direct correlation between plant communities and soil degradation variables. the stepwise ordination approach followed by mucina and van tongeren (1989) was adopted to cope with the heterogeneity of the data set. description of plant communities descriptive terms such as ‘differential species’ and ‘dominant species’ were used following kent and coker (1992) and jennings et al. (2009). ‘differential species’refers to species of medium to low constancy that tend to occur together in a series of quadrats and can thus be used to characterise groups. ‘dominant species’ refers to species with the highest percentage of cover, usually in the uppermost dominant layer. results top ↑ vegetation classification daemane et al. (2010) previously described nine plant communities (figure 1) for the hnp. a further three major plant communities and three subcommunities occurring in the spitskop area are described in this study (figure 2). in the descriptions of the different plant communities, all species groups refer to the phytosociological table (online appendix 1). therefore no repeated reference will be made to online appendix 1 in the descriptions of the various plant communities. the vegetation comparisons, where applicable, were made between the plant communities of the spitskop area and those previously described in the hnp (daemane et al. 2010) and in the grassland of the former western transvaal (bezuidenhout & bredenkamp 1991; bredenkamp, joubert & bezuidenhout 1989;cilliers, van wyk & bredenkamp 1999). the hierarchical classification of the spitskop communities is as follows: 1. acacia caffra – seriphium plumosum woodland 1.1. acacia robusta – melinis repens woodland 1.2. mundulea sericea subsp. sericea – eragrostis racemosa shrubland acacia karroo – asparagus suaveolens shrubland setaria sphacelata var. torta – eragrostis curvula grassland cymbopogon pospischilii – elionurus muticus grassland figure 2: vegetation map showing degraded and non-degraded plant communities of the spitskop area in the highveld national park, potchefstroom, south africa. detailed description 1. acacia caffra – seriphium plumosum woodland the acacia caffra – seriphium plumosum woodland occurred in the rocky midslope of the quartzite outcrops and was associated with sheet, gully and rill erosion. the glenrosa soil form was dominant in this habitat. differential species of this community, indicated by species group a, included acacia caffra, seriphium plumosum, gomphocarpus fruticosus subsp. fruticosus, aristida bipartita and convolvulus sagittatus. the dominant species in this community were diospyros lycioides subsp. lycioides (species group d); mundulea sericea subsp. sericea (species group f); eragrostis racemosa, urochloa panicoides, searsia pyroides var. pyroides and euclea undulata (species group h); acacia karroo, setaria sphacelata var. sphacelata and ziziphus zeyheriana (species group l), and melinis repens var. repens, aloe greatheadii var. davyana, cynodondactylon, themeda triandra, hyparrhenia hirta and asparagus suaveolens (species group m). on average, 18 species were recorded per sample plot. the average height of the tree stratum was 1.6 m with a canopy cover of 10.5%. the average height of the shrub stratum was 0.5 m and the canopy cover was 5.7%. the average height of the herbaceous layer was 0.2 m, with a canopy cover of 3.1%. a closely related community in the park, acacia caffra – setaria sphacelata var. sphacelata woodland (daemane et al. 2010), had an average of 40 species per plot and herbaceous and tree cover of 61.7% and 11.2%, respectively. species such as s. plumosum, a. greatheadii var. davyana and a. suaveolens were the most common and abundant species in both these plant communities. acocks (1988) attributed the increase of these species to conditions of severe degradation due to overgrazing. uncontrolled fires, gathering of trees for firewood and accompanying disturbances, such as trampling and soil compaction by livestock, were found to be the major threats to these plant communities (cilliers et al. 1999; daemane et al. 2010). the acacia caffra – seriphium plumosum woodland community was divided into the three sub-communities described below. 1.1. acacia robusta – melinis repens woodland this plant community occurred in the rocky midslope of the quartzite outcrops and was associated with sheet, gully and rill erosion. sheet erosion was severe, covering up to 92% of the soil surface. glenrosa soil form was dominant in this habitat. differential species of this community, indicated by species group b, includedthe grass panicum maximum and the tree acacia robusta subsp. robusta. the dominant shrubs in this community were acacia caffra and seriphium plumosum (species group a); eragrostis racemosa and searsia pyroides var. pyroides (species group h); and diospyros lycioides subsp. lycioides (species group d). on average, 16 species were recorded per sample plot. the average height of the tree stratum was 1.3 m and the canopy cover was 5.2%. the average height of the shrub stratum was 0.5 m andthe canopy cover was 1.8%. the average height of the herbaceous stratum was 0.2 m, with a low canopy cover of 1.0%. 1.2. mundulea sericea subsp. sericea – eragrostis racemosa shrubland the mundulea sericea subsp. sericea – eragrostis racemosa shrubland occurred in the rocky midslope of the quartzite outcrops and was associated with sheet, gully and rill erosion. sheet erosion was severe, covering 83% of this plant community. glenrosa was the dominant soil form. the differential species in this community was found in species group c and consisted of only one species, indigofera sp.; species group b was absent from this subcommunity. dominant species in this communityincluded acacia caffra, seriphium plumosum and gomphocarpus fruticosus subsp. fruticosus (species group a); diospyros lycioides subsp. lycioides and clematis brachiata (species group d); mundulea sericea subsp. sericea (species group f); eragrostis racemosa (species group h); setaria sphacelata var. sphacelata (species group l), andmelinis repens var. repens, aloe greatheadii var. davyana, cynodon dactylon and themeda triandra (species group m). on average, 17 species were recorded per sampleplot. the average height of the tree stratum was 1.5 m, with a canopy cover of 9.3%. the average height of the shrub stratum was 0.5 m and the canopy cover was 5.3%. the average height of the herbaceous layer was 0.1 m and the canopy cover was 2.3%. 1.3. acacia karroo – asparagus suaveolens shrubland the acacia karroo – asparagus suaveolens shrubland occurred in the footslopes and was associated with sheet, rill and gully erosion. sheet erosion covered 71% of the surface area. hutton and glenrosa were the dominant soil forms. the differential species of this community, indicated by species group e, included senecio sp., atriplex sp. and lantana rugosa. the dominant species were acacia caffra, seriphium plumosum, aristida bipartita and convolvulus sagittatus (species group a); mundulea sericea subsp. sericea (species group f); urochloa panicoides, searsia pyroides var. pyroides and eragrostis racemosa (species group h); acacia karroo, setaria sphacelata var. sphacelata and ziziphus zeyheriana (species group l), and melinis repens var. repens, aloe greatheadii var. davyana, cynodon dactylon, themeda triandra, hyparrhenia hirta and asparagus suaveolens (species group m). on average, 21 species were recorded per sample plot. this plant community was affected by sheet, rill and gully erosion. sheet erosion covered 71% of the surface area. the average height of the tree stratum was 2.1 m and the canopy cover was 17%. the average height of the shrub stratum was 0.6 m, with a canopy cover of 10%. the average height of the herbaceous stratum was 0.2 m and the canopy cover was 6.0%. a closely related plant community, acacia karroo – ziziphus zeyheriana woodland, was described by daemane et al. (2010). bredenkamp et al. (1989) described a closely related community, the a. karroo woodland, also occurring on severely degraded land but with a higher tree cover (30%). species such as z. zeyheriana, a. greatheadii var. davyana and a. suaveolens were common in all these plant communities and seem to be increasing under conditions of severe overgrazing, as noted by bredenkamp et al. (1989). 2. setaria sphacelata var. torta – eragrostis curvula grassland this plant community occurred in the footslopes and was associated with sheet, rill and gully erosion. sheet erosion covers an average of 76% of the soil surface. hutton and mispah were the dominant soil forms. the differential species in this community were in species group g and included eragrostis curvula, digitaria eriantha, setaria sphacelata var. torta, grewia flava, asparagus laricinus and solanum lichtensteinii. dominant species in this community were eragrostis racemosa and searsia pyroides var. pyroides (species group h); chrysocoma ciliata and eragrostis lehmanniana var. lehmanniana (species group k); acacia karroo (species group l), and melinis repens var. repens, aloe great headii var. davyana, cynodon dactylon, themeda triandra, hyparrhenia hirta and asparagus suaveolens (species group m). on average, 23 species were recorded per sample plot. the average height of the tree stratum was 1.2 m and the canopy cover was 11.2%. the average height of the shrub stratum was 0.3 m and the canopy cover was 6.8%. the average height of the herbaceous layer was 0.2 m, with a canopy cover of 5.4%. 3. cymbopogon pospischilii – elionurus muticus grassland the cymbopogon pospischilii – elionurus muticus grassland occurred in the valley bottomland and floodplains. this plant community was characterised by bare patches of compacted soils resulting from overgrazing and trampling by livestock. hutton and mispah were the dominant soil forms. the differential species for this community were in species group i and included elionurus muticus, eustachys paspaloides, aristida congesta subsp. congesta, sida alba, vernonia oligocephala and gomphrena celosioides. dominant species in this community included cymbopogon pospischilii (species group j); setaria sphacelata var. sphacelata, acacia karroo and ziziphus zeyheriana (species group l), and melinis repens var. repens, aloe greatheadii var. davyana and cynodon dactylon (species group m). on average, 17 species were recorded per sample plot. the average height of the tree stratum was 1.0 m, with a canopy cover of 0.5%. the average height of the shrub stratum was 0.6 m and the canopy cover was 1.0%. the average height of the herbaceous layerwas 0.4 m and the canopy cover was 2.9%. a closely related plant community, cymbopogon pospischilii – themeda triandra grassland, was described by daemane et al. (2010), with an average of 31 species per sample plot. the herbaceous canopy cover for the cymbopogon pospischilii – themeda triandra grassland was 77% compared to 0.5% in the cymbopogon pospischilii – elionurus muticus grassland in the spitskop area. another closely related plant community, heteropogon contortus – themeda triandra – elionurus muticus grassland, with an average cover of 71%, 24 species per sample plot and absolute dominance of themeda triandra, was described by bredenkamp et al. (1989). this type of vegetation was previously used as pasture and often overgrazed (bredenkamp et al. 1989). ordination the environmental gradients and the relative importance and intercorrelation of the environmental variables are shown by the arrows by the canonical correspondence analysis (figure 3). on axis 1, compacted soil was the most important variable determining the species composition. the eigenvalue for the firstordination axis of the plant communities/environment biplot was 0.28, representing 34.9% of the total variance. on axis 2, sheet erosion was the most important variable determining species composition. the eigenvalue for the second axis was 0.23, representing 63.5% of the total variance. thus, the two axes account for 98.4% of the variance in the plant communities. rill and gully erosion were of lesser significance in explaining the species composition. the two different associationsbetween vegetation and soil compaction or sheet erosion emerged clearly in two separate parts of the diagram. the seriphium plumosum – acacia caffra woodland (including three subcommunities) and the setaria sphacelata var. torta – eragrostis curvula grassland were associated with sheet erosion. the cymbopogon pospischilii – elionurus muticus grassland was associated with compacted soils. figure 3: canonical correspondence analysis showing the correlation between plant communities and soil erosion gradients in the spitskop area in the proposed highveld national park. discussion top ↑ many of the plant communities previously described in the grassland of the north west province (bredenkamp et al. 1989; cilliers et al. 1999; daemane et al. 2010)showed some floristic affinities with the plant communities described in the spitskop area. according to guardia and ninot (1992), the vegetation of the degraded areas retains a strong floristic similarity to the better preserved vegetation patches in close proximity. in the spitskop area, the vegetation was characterised by relatively low plant cover and lower species richness compared to the non-degraded plant communities described by daemane et al. (2010) and other plant communities previously described in the area (bredenkamp et al. 1989; cilliers et al. 1999; daemane et al. 2010). the general trends concerning the regressive dynamics of plant communities caused by soil degradation include decrease of vegetation cover and a reduction in species richness (guerrero-campo & montserrat-marti 2000; eds. morgan & rickson 1995). the results from the spitskop area showed that woody species such as acacia caffra, searsia pyroides var. pyroides and acacia karroo have decreased in their cover and abundance compared to the similar, less degraded plant community of the hnp (daemane et al. 2010). other woody species, such assearsia leptodictya and ehretia rigida found in the acacia caffra – setaria sphacelata var. sphacelata community of the hnp (daemane et al. 2010) have been lost in the degraded communities of the spitskop area. grasses such as themeda triandra, digitaria eriantha, elionurus muticus, heteropogon contortus and cymbopogon pospischilii have also declined in cover in the spitskop area. the grass cynodon dactylon has been totally lost in the areas affected by sheet erosion and where the soil has been compacted by livestock. overall, the ordination results indicated that the species composition was affected largely by sheet erosion and soil compaction by livestock. the spitskop area is also close to the dam and concentrated grazing was one of the factors that contributed to the formation of sheet erosion and soil compaction. according to valentin, poesen and yong (2005) water infiltration is low in areas affected by sheet erosion and compacted soil, resulting in low vegetation cover and species richness. cattletracks were observed to cause the formation of rill and gully erosion where a concentration of water runoff occurred (daemane 2007). rill and gully erosion were mostly associated with the midlopes and the footslopes of the spitskop hill. however, gully and rill erosion were found to be of lesser significance in explaining species composition in the spitskop area. in general, the destruction of the protective vegetation cover by overgrazing, trampling and wood harvesting contributed to soil and vegetation degradation in the spitskop area (bredenkamp et al. 1989; cilliers et al. 1999; daemane 2007). the combination of these factors seemed to have enhanced the aridity of the area becauseof the reduction in vegetation cover. the rate of soil erosion is also highly dependent on land use, slope and climatic factors (garland 1987). observations in the spitskop area showed that the slope of the spitskop hill also contributed to the accelerated erosion. although experimental trials were not undertaken to validate the effect of slope on erosion, rills and gullies were associated with the midslopes and footslopes of the spitskop hill. according to valentin et al. (2005), water runoff from steep slopes reach lower slopes with intense velocity, resulting in soil erosion. in areas where sheet erosion and compacted soil were pronounced, the upper layers of the soil profile had already been lost, without any organic matter left. only a few species, such as seriphium plumosum, ziziphus zeyheriana, mundulea sericea subsp. sericea and aloe great headii var. davyana, were abundant and their covers seemed to increase under conditions of severe sheet erosion and soil compaction bylivestock. conclusion top ↑ the results obtained in this study highlighted the ecological relationship between vegetation and soil degradation. the proposed strategies for soil conservation are based on covering the soil for protection from natural and human-induced disturbances. rehabilitation measures such as mechanical soil cultivation, brush packing and stone packing proved effective in areas elsewhere affected by sheet erosion and soils compacted by livestock (coetzee 2005; van der merwe & kellner 1999;wight & white 1974). the gullies and rills can also be addressed by stone packing and the use of gabions can be applied in severe conditions (coetzee 2005). the dam also needs to be rehabilitated to reduce the concentrated grazing around the spitskop area. development of a fire management plan for the park to minimise the impact of fire on vegetation structure and cover is suggested. a resource use policy also needs to be developed to address the issues such as firewood collection within the park. the ecological interpretation derived in this study will therefore be a valuable tool for development of a rehabilitation plan and the general conservation management of this grassland area. acknowledgements top ↑ the following individuals and institutions are sincerely thanked: south african national parks for financing this study; north-west university for supporting the research; dr. stephen holness and ms. phozisa mamfengu for assisting with maps; mr. abbey legari for assisting with fieldwork and the hnp founding partners for allowing us to undertake this study. competing interests the authors declare that they have no financial or personal relationship(s) which may have inappropriately influenced them in writing this paper. authors’ contributions m.e.d. (south african national parks) was responsible for vegetation sampling and writing of the manuscript, s.s.c. (north-west university) assisted with the ordination analysis and h.b. (south african national parks) assisted with the twinspan data analysis and classification of plant communities. references top ↑ acocks, j.p.h., 1953, ‘veld types of south africa’, memoirs of the botanical survey of south africa 28, 1–192. acocks, j.p.h., 1988, ‘veld types of south africa’, 3rd edn, memoirs of the botanical survey of south africa 57, 1–146. barkman, j.j., doing, h. & segal, s., 1964, ‘kritische bemerkungen und vorschläge zur quantitativen vegetationsanalyse [critical remarks and suggestions for the quantitative analysis of vegetation]’, acta botanica neerlandica 13, 394–419. bezuidenhout, h. & bredenkamp, g.j., 1991, ‘the vegetation of the bc land type in the western transvaal grassland, south africa’, phytocoenologia 19(4), 497–518. bezuidenhout, h., biggs, h.c. & bredenkamp, g.j., 1996, ‘a process supported by the utility bbpc for analysis of braun-blanquet data on personal computer’, koedoe 39, 107–112. brady, h.m., 1993, ‘an investigation into the nature of gully erosion at golden gate highlands national park’, msc thesis, dept. of geography, university of natal. bredenkamp, g.j. & bezuidenhout, h., 1995, ‘a proposed procedure for the analysis of large data sets in the classification of south african grasslands’, koedoe 38, 33–39. bredenkamp, g.j., joubert, a.f. & bezuidenhout, h., 1989, ‘a reconnaissance survey of the vegetation of the plains in the potchefstroom–fochville–parys areas’, south african journal of botany 55(2), 199−206. bredenkamp, g.j. & van rooyen, n., 1996a, ‘rocky highveld grassland’, in a.b. low & a.g. rebelo (eds.), vegetation of south africa, lesotho and swaziland, p. 39, department of environmental affairs and tourism, pretoria. bredenkamp, g.j. & van rooyen, n., 1996b, ‘dry sandy highveld grassland’, in a.b. low & a.g. rebelo (eds.), vegetation of south africa, lesotho and swaziland, p. 41, department of environmental affairs and tourism, pretoria. cilliers, s.s., 1998, ‘a phytosociological study of urban open spaces in potchefstroom, north west province, south africa’, phd thesis, school of environmental sciences and development, potchefstroom university for christian higher education. cilliers, s.s., müller, n. & drewes, e., 2004, ‘overview on urban nature conservation: situation in the western-grassland biome of south africa’, urban forestry & urban greening 3, 49–62. cilliers, s.s., van wyk, e. & bredenkamp, g.j., 1999, ‘urban nature conservation: vegetation of natural areas in the potchefstroom municipal area, north west province, south africa’, koedoe 42(1), 1–30. cobban, d.a. & weaver, a.v.b., 1993, ‘a preliminary investigation of the gully features in the tsolwane game reserve, ciskei, south africa’, south african geographical journal 75, 14–21. coetzee, k., 2005, caring for natural rangelands, university of kwazulu-natal press, scottsville. daemane, m.e., 2007, ‘an ecological study of the plant communities and degraded areas of the highveld national park, north west province, south africa’, msc thesis, school of environmental sciences and development, north-west university. daemane, m.e., cilliers, s.s. & bezuidenhout, h., 2010, ‘an ecological study of the plant communities in the proposed highveld national park, in the peri-urban area of potchefstroom, south africa’, koedoe 52(1), art #708, 8 pages. http://dx.doi.org/10.4102/koedoe.v52i1.708 garland, g.g., 1987, ‘rates of soil loss from mountain footpaths: an experimental study in the drakensberg mountains, south africa’, applied geography 7, 41–54. http://dx.doi.org/10.1016/0143-6228(87)90006-3 guardia, r. & ninot, j.m., 1992, ‘on the vegetation–topography relationship in the badlands of the upper llobregat basin (southeastern pyrenees)’, studia geobotanica 12, 83–103. guerrero-campo, j. & montserrat-marti, g., 2000, ‘effects of soil erosion on the floristic composition of plant communities on marl in northeast spain’, journal of vegetation science 11, 329–336. http://dx.doi.org/10.2307/3236625 hill, m.o., 1979, twinspan – a fortran program for arranging multivariate data in an ordered two-way table by classification of individuals and attributes, cornell university, ithaca, ny. institute for soil, climate and water, 2003, comprehensive weather report: brakspruit, potchefstroom, agricultural research council, pretoria. iscw. see institute for soil, climate and water. jennings, m.d., faber-langendoen, d., loucks, o.l., peet, r.k. & roberts, d., 2009, ‘standards for associations and alliances of the u.s. national vegetation classification’, ecological monographs 79, 173–199. kent, m. & coker, p., 1992, vegetation description and analysis: a practical approach, john wiley & sons, chichester. land type survey staff, 1984, ‘land types of the maps 2626 west rand and 2726 kroonstad’, memoirs of agricultural natural resources of south africa 4, 1–441. liggitt, b., 1988, ‘an investigation into soil erosion in the mfolozi catchment’, final report, kwazulu bureau of natural resources, university of natal. macvicar, c.n., loxton, r.f. & lambrechts, j.j.n., 1991, ‘soil classification: a taxonomic system for south africa’, memoirs on the agricultural natural resources of south africa 15, 1–257. morgan, r.p.c. & rickson, r.j. (eds.), 1995, slope stabilization and erosion control: a bioengineering approach, e & fn spon, london. mucina , l. & van der maarel, e., 1989, ‘twenty years of numerical syntaxonomy’, vegetatio 81, 1–5. mucina, l. & van tongeren, o.f.r., 1989, ‘a coenocline of the high-ranked syntaxa of ruderal vegetation’, vegetatio 81, 117–125. snyman, h.a., van rensburg, w.l.j. & opperman, d.p.j., 1986, ‘application of a soil loss equation on natural veld of the central orange free state’, journal of the grassland society of southern africa 3, 4–9. soil classification working group, 1991, ‘soil classification: a taxonomic system for south africa’, memoirs on the agricultural natural resources of south africa 15, 1–262. south african national biodiversity institute, 2009, plants of southern africa: an online checklist, viewed 26 august 2011, from http://posa.sanbi.org. ter braak, c.j.f., 1986, ‘canonical correspondence analysis: a new eigenvector technique for multivariate direct gradient analysis’, ecology 65, 1167–1179. torrion, j.a., 2002, ‘land degradation detection, mapping and monitoring in the lake naivasha basin, kenya’, msc thesis, international institute for geo-information science and earth observation, enschede. valentin, c., poesen, j. & yong, li., 2005, ‘gully erosion: impacts, factors and control’, catena 63,132–153. van der merwe, j.p.a. & kellner, k., 1999, ‘soil disturbance and increase in species diversity during rehabilitation of degraded arid rangelands’, journal of arid environments 41, 323–333. wight, j.r. & white, l.m., 1974, ‘interseeding and pitting on a sandy range site in eastern montana’, journal of range management 27, 206–210. article information authors: corli coetsee1 benjamin j. wigley1 affiliations: 1school of natural resource management, nelson mandela metropolitan university, george campus, south africa correspondence to: corli coetsee postal address: private bag x6531, george 6530, south africa dates: received: 26 nov. 2012 accepted: 22 apr. 2013 published: 15 july 2013 how to cite this article: coetsee, c. & wigley, b.j., 2013, ‘virgilia divaricata may facilitate forest expansion in the afrotemperate forests of the southern cape, south africa’, koedoe 55(1), art. #1128, 8 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1128 copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. virgilia divaricata may facilitate forest expansion in the afrotemperate forests of the southern cape, south africa in this original research... open access • abstract • introduction • methods    • study site    • sample collection and analyses    • statistical analyses • results • discussion • conclusion • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ virgilia divaricata is a fast-growing nitrogen-fixing tree species often found on the margins of forest in the southern cape of south africa and is particularly abundant after fire. however, v. divaricata may invade fynbos even in the absence of fire and it has been described as a forest precursor. we investigated whether v. divaricata enriches soil fertility after its invasion into fynbos areas adjacent to forests. we measured soil organic carbon and soil nutrients at four sites. at each site, three vegetation types (forest, v. divaricata and fynbos) were examined on the same soil type and at the same elevation. our results showed that, on average, soils taken from v. divaricata stands had higher nitrogen and phosphorus values than the adjacent fynbos soils, with either lower or similar values to the adjacent forest soils. higher soil fertility under v. divaricata, together with their shading effect, may create conditions favourable for shade-loving forest species dependent on an efficient nutrient cycle in the topsoil layers, and less favourable for shade-hating fynbos species, which are generally adapted to low soil fertility. we suggest that the restoration of the nutrient cycle found in association with forest may be accelerated under v. divaricata compared with other forest precursor species, which has important consequences for the use of v. divaricata in ecosystem restoration.conservation implications: alien plantations in the outeniqua mountains are being phased out and the areas are being incorporated into the garden route national park. fynbos areas are increasingly being invaded by forest and thicket species owing to fire suppression in lower-lying areas. an improved understanding of the fynbos–forest boundary dynamics will aid in efficient management and restoration of these ecosystems. introduction top ↑ in south africa, natural forest covers about half a million hectares, which equates to only 0.5% of the land surface. apart from being the smallest biome, forest is also the most fragmented and most vulnerable biome within the country (van der merwe 2011). the largest forest complex is the afrotemperate knysna forest, which covers 65 000 ha of the southern cape. heath-like vegetation that is adapted to frequent fire, known locally as fynbos, forms a matrix with these forests (manders, richardson & masson 1992). fynbos is evergreen and sclerophyllous. geldenhuys (1994) has shown that although environmental factors such as rainfall and substrate determine the potential limits of forest distribution, actual forest distribution is determined by the interaction between prevailing winds during dry periods, terrain physiography and the effect of this interaction on fire pattern. in our study area, forests are usually found in the topographic shadow areas of warm and desiccating berg winds, which are common in autumn and winter (geldenhuys 1994). however, extensive plantation forestry, which has protected fynbos against fire, has led to forest expansion in the southern cape in recent times (c. geldenhuys, pers. comm., january 2011). in the prolonged absence of fire, forest species start to invade the surrounding fynbos and the dominant vegetation may switch to closed-canopy forest (kruger 1984; masson & moll 1987). forest trees face certain inhibitory factors when expanding into fynbos. these include the lack of perches for birds, which inhibits seed dispersal, the inability of seedlings to survive summer droughts, lack of protection from wind and radiation and low levels of soil nutrients (campbell & killick 1985; manders 1990; manders & richardson 1992; white 1978). manders and richardson (1992) propose a model of forest development in fynbos where the mechanism of succession involves facilitation. forest precursor species act as ‘nurse plants’ by increasing the attractiveness of the area to avian dispersers, ameliorating summer water deficits owing to a mulching effect of litter, protecting forest trees from radiation and wind, and decreasing competition for water owing to decreased cover beneath these plants. the concept of pioneer plants acting as nurse plants and ‘nucleation’ sites for subsequent woody vegetation development has been reported in many ecosystems (e.g. barnes & archer 1999; yarranton & morrison 1974). such modification of the environment by an organism has been referred to as ‘niche construction’ or ‘ecosystem engineering’ (crain & bertness 2006; laland, odling-smee & feldman 1999; odling-smee, laland & feldman 1996). examples include changing the chemical nature of soils, the pattern of nutrient cycling, the temperature, humidity, fertility, acidity and salinity of soils, and the patterns of shade in the habitat (laland et al. 1999). virgilia divaricata adamson and the closely related virgilia oroboides (p.j. bergius) salter (considered as one species by some authors and locally known as keurboom) are commonly observed growing on forest margins in the southern cape (phillips 1926). virgilia spp. seeds can be viable for at least 230 years, enabling the plant to recover or regenerate rapidly after fire (geldenhuys 1994). this fast-growing tree can establish without shade. seedlings and saplings of forest canopy species can usually be found 5–10 years after the development of virgilia spp. stands and become established after the natural suppression, mortality and consequent thinning of the pioneer stand (geldenhuys 1994). phillips (1926) has shown that the growth of seedlings and saplings of forest species co-dominant with virgilia spp. is considerably more rapid under virgilia spp. stands than in normal climax forest. the mechanism of this enhanced growth was not investigated; however, phillips (1926) suggests that it may be related to light conditions under virgilia spp. being more conducive to sapling growth than the low light conditions beneath forest. we suggest that an alternative mechanism for facilitation of forest sapling growth may be that v. divaricata increases soil fertility, as both virgilia spp. are members of fabaceae, which have root nodules containing nitrogen-fixing bacteria (palgrave 1981). nitrogen-fixing alien acacias, which regularly invade fynbos, have been shown to increase soil nutrient mineral content (musil 1993; stock, wienand & baker 1995; witkowski 1991a; yelenik, stock & richardson 2004). however, little prior research has considered the effects of v. divaricata on soil fertility and the current study therefore aimed to investigate whether the establishment of v. divaricata in fynbos patches adjacent to forests enhances soil fertility. we hypothesised that soils underneath the legume v. divaricata will become nitrogen enriched as a result of the addition of nitrogen-rich litter and that the higher nitrogen levels will be perpetuated by forest development. we also hypothesised that levels of other nutrients, including carbon, will increase as forest establishment progresses. methods top ↑ study site the study sites were situated in the garden route national park in the western cape: two on the northern edges of the groeneweide forest, 5 km east of george (groenkop 1: 33.939436° s 22.560760° e; groenkop 2: 33.943334° s 22.562730° e) and two in the tsitsikamma bloukrans forest area near nature’s valley (plateau: 33.565690° s 23.45856° e; covie: 33.943334° s 22.562730° e). the average rainfall varies between 500 mm and 1200 mm, with peaks in autumn and early summer and a minimum in december. temperatures are mild, with mean minima and maxima of 7 °c and 19 °c, respectively, in june and 15 °c and 26 °c, respectively, in january (bond 1981). soils are derived from quartzitic sandstone of the table mountain group. the topsoil in the groenkop sites is between 40 cm and 50 cm deep and overlays poorly drained subsoil with indications of a fluctuating water level, more so at groenkop 1 than groenkop 2. the two tsitsikamma sites were situated on old dunes. (see geldenhuys [1982, 1991] for a thorough environmental description of the area.) the forests studied fall within the southern afrotemperate forest vegetation type (mucina & rutherford 2006). the surrounding vegetation is mostly fynbos, with south outeniqua sandstone fynbos bordering the groeneweide forest and tsitsikamma sandstone fynbos in the tsitsikamma sites (mucina & rutherford 2006). the study sites at groeneweide (groenkop 1 and 2) burnt in 1996 and well-established adult v. divaricata trees now form mostly consocies stands, approximately 10 m – 15 m high at the ecotone between forest and fynbos. the fynbos at covie is dominated by young leucadendron saplings, indicative of a recent fire, with a narrow band of v. divaricata between forest and fynbos areas. the fynbos at plateau is mostly lacking in proteaceae, probably because the area now conserved as a nature reserve has been used for (potato) farming until quite recently. the v. divaricata bordering the forest at plateau established after a fire in 2005. finding sites where the fynbos was in pristine condition was a typical problem during site selection. in the past, fynbos adjacent to forest was either farmed or planted with exotic plantations. currently, fire suppression, especially in the lower-lying fynbos patches, has led to encroachment by thicket species, forest precursors or invasive alien trees. sample collection and analyses at each of the four sites, soils were collected from within the forest, adjacent fynbos vegetation and v. divaricata stands. care was taken to select sites where forest, fynbos and v. divaricata occurred at the same elevation and on the same soil type. at each site, the three treatments (forest, fynbos and v. divaricata) were close to one another. adult stands of v. divaricata formed narrow strips (~20 m – 80 m) of consocies with an understorey of sparsely distributed forest saplings on the margin between forest and fynbos. we tried to avoid areas where pteridium aquilinum (bracken) occurred as we were uncertain how the strong allelopathic substances exuded by bracken would affect soils (chapman & reiss 1992); however, bracken occurred in quite dense patches in some areas adjacent to v. divaricata, especially at plateau. soil was collected on transects spanning across the v. divaricata stands and on transects running parallel to either side of the v. divaricata stands in forest and fynbos. soil samples were collected about 20 m – 30 m apart, with a soil auger at three depths: 0 cm – 10 cm, 10 cm – 20 cm and 20 cm – 50 cm (4 sites × 3 vegetation types × 3 depths × 5 repetitions = 180 samples). after collection, soils were sieved through a 2-mm sieve to remove organic material and roots, and air-dried for several days. all samples were analysed for organic carbon, nitrogen, phosphorus, calcium, magnesium, sodium and potassium at the institute for plant production, stellenbosch, south africa. carbon content was determined by a rapid dichromate oxidation method using the walkley–black procedure (walkley 1947). samples were not pre-treated for carbonates as the ph was well below 7.4, which suggested that carbonates were not present in significant amounts in our samples (nelson & sommers 1996). repeatedly run samples analysed for carbon had a standard deviation of 0.03% and coefficient of variation (cv) of 0.07. extractable phosphorus, calcium, magnesium, sodium and potassium were extracted with 1% citric acid and analysed by using a thermo icp icap 6000 series spectrometer (thermofisher scientific, surrey, uk). total soil nitrogen was analysed by total combustion at bemlab (pty) ltd., somerset west, south africa, using a leco nitrogen analyser fp 528 (leco corporation, st. joseph, usa). to determine bulk density, a known volume was removed from each soil depth using a core sampler. known volumes were dried, weighed and sieved to remove all roots and stones, and the weight and volume of rocks were measured. the bulk density of each sample was calculated by dividing the dry mass of soil by the volume of soil, with the mass of the rocks subtracted in each case. bulk density was used to convert concentration data to content data (i.e. aerial data). we underestimated bulk density for samples with high root biomass by not subtracting root volume from soil volume. as forests typically have high root biomass, forest bulk densities were consistently underestimated. we therefore considered both the concentration and content data in our analyses. at the two tsitsikamma sites, we dried and weighed the roots before the samples were sent for analysis. litter was also collected from beneath the canopy of the three treatments (forest, fynbos and v. divaricata) and both root and litter samples were analysed for nitrogen and carbon using a leco truspec cn analyser (leco corporation, st. joseph, usa). statistical analyses all statistical analyses in this study were performed using r (r development core team 2011). principle component analyses (pca) were used to explore the relationships between the three vegetation types and various elemental soil variables. the broken-stick distribution (pca signifance criterion in r) was used as the most reliable criterion to test which of the pca axes contributed significantly to the ordination. pca axes with larger percentages of accumulated variance than the broken-stick variance were considered significant (kindt & coe 2005; legendre & legendre 1998). a three-way analysis of variance (anova) was fitted to explore the effects of site, vegetation type, depth and the respective interactions on soil factors. where necessary, variables were transformed to attain approximate normality and homogeneity of variance. post hoc tukey tests were used to test for differences between levels of each factor in the anovas. results top ↑ pc1 of the pca explained 59% of the variance in soils (eigenvalue 4.19; x-axis in figure 1), with high loads for nitrogen (-0.39), carbon (-0.39), potassium (-0.41), magnesium, sodium and phosphorus (all -0.38). the three vegetation types were arranged along a gradient of soil fertility, according to which forests were found on the most fertile soils, v. divaricata was associated with intermediate soil nutrients, and fynbos was found on low-nutrient soils. pc2 (y-axis in figure 1) explained a further 16% of the variance and separated sites with high nitrogen (+0.42) and carbon (+0.23) from those with high calcium (-0.71) and magnesium (-0.44); however, this axis was not deemed significant. this axis reflected site differences to a great extent. covie soils, for example, had very high levels of calcium and magnesium. figure 1: principle component analysis based on measures of soil fertility (soil nutrients). anova results showed that the interactive effect of site and vegetation type on nitrogen concentration was statistically significant but not the interaction amongst the three factors or any of the other two-way interactions (table 1). although depth had less of an effect on nitrogen content, it was an important determinant of soil concentrations (table 1). when nitrogen concentration was converted to nitrogen content, both the interactions between site and vegetation type and site and depth became important in determining nitrogen (table 1). table 1: results of three-way analysis of variance, showing the effect of site, block (vegetation type) and depth on various soil variables. across sites, forest soils had the highest nitrogen concentration (0.44% ± 0.05% and 34.2 g.m–2 ± 3.18 g.m–2), followed by v. divaricata soils (0.29% ± 0.02% and 25.4 g.m–2 ± 1.55 g.m–2) and fynbos soils (0.18% ± 0.01% and 19.2 g.m–2 ± 0.86 g.m–2). for individual sites, groenkop 1 and groenkop 2 reflected the general pattern for both nitrogen concentration (figure 1a) and nitrogen content (table 2), with the highest values associated with forest and the lowest values associated with fynbos. at covie, forest and v. divaricata soils had similar nitrogen concentrations (0.24% ± 0.02% and 0.26% ± 0.04%, respectively), but these were higher than for fynbos (0.14% ± 0.01%; figure 1a). nitrogen content was similar for forest and v. divaricata, but v. divaricata had higher nitrogen levels than fynbos (table 2). at plateau, all three vegetation types had similar nitrogen concentrations (figure 2) and nitrogen content (table 2). as expected, the shallowest soils generally had the highest nitrogen concentration (0.44% ± 0.56%), followed by the 10-cm – 20-cm depths (0.19% ± 0.01%) and then the deeper soils (0.27% ± 0.19%). table 2: the effect of interactions between site and vegetation type on mean nitrogen and phosphorus content in the southern cape (g.m–2 ± se). figure 2: the effect of an interaction between site and vegetation type on soil nitrogen in the southern cape. significant interactions affecting phosphorus concentration are shown in table 1. when phosphorus concentrations were converted to phosphorus content, the interaction between vegetation type and depth, in addition to the other factors and interactions, was also statistically significant (table 1). although the three-way interaction was significant, its importance was relatively small compared to site × vegetation type and vegetation type × depth (table 1). however, to confirm that the three-way interaction was significant, we compared the interaction between site and vegetation type for each depth (figure 3). at 0 cm – 10 cm and 10 cm – 20 cm, covie forest and v. divaricata soils had higher phosphorus concentrations than fynbos; groenkop 1 had higher phosphorus concentrations associated with forest than fynbos (figure 3). no differences were found for phosphorus concentration between vegetation types at plateau. at 40 cm – 50 cm, fynbos still had a lower phosphorus concentration than forest and v. divaricata at covie, but phosphorus concentration for fynbos was lower than for both forest and v. divaricata at groenkop 1 and plateau (figure 3). when phosphorus concentrations were converted to content data, no differences were found between vegetation types at groenkop 2. virgilia divaricata had higher phosphorus content than fynbos at covie and groenkop 1, whilst fynbos soils had higher phosphorus content than the other two vegetation types at plateau (table 2). figure 3: soil phosphorus (mg.kg–1) for different vegetation types at four sites at different depths: (a) 0 cm – 10 cm, (b) 10 cm – 20 cm and (c) 40 cm – 50 cm. we measured soil carbon-to-nitrogen ratios (c:n) at all four sites, as well as c:n for litter and root material at covie and plateau. site had a very pronounced influence on soil c:n and all interactions with site were significant (table 1). apart from the interactions with site, the interaction between vegetation type and depth was also statistically significant. the three-way interaction was not significant. when comparing individual sites, soils from groenkop 1 and groenkop 2 showed no differences in c:n between the three vegetation types (figure 4). at covie, soils beneath forest and v. divaricata showed the highest c:n, both of which were higher than for soils beneath fynbos (figure 4). at plateau, forest and v. divaricata had similar values, and v. divaricata and fynbos had similar values, but forest had higher soil c:n than fynbos. the site × vegetation type interaction was statistically significant for root c:n (p = 0.002). at covie, v. divaricata soils had lower root c:n than fynbos soils, but no differences were found at plateau (figure 4). unfortunately, we did not have enough repetitions to compare c:n for litter statistically and can therefore comment only on the general trend. litter c:n showed the same pattern as root c:n, with v. divaricata samples having the lowest litter c:n (22.8 ± 1.75), followed by forest (35.7 ± 3.16) and fynbos samples (51.8 ± 6.41). figure 4: carbon-to-nitrogen ratios for different ecosystem components: (a) soil and (b) root. discussion top ↑ in this study, we compared differences in soil fertility between stands dominated by forest, v. divaricata and fynbos vegetation, respectively, to establish the effect of v. divaricata on soil fertility. firstly, we used pca to explore relationships amongst soil variables and found that the different vegetation types separated in ordinational space, with forest, v. divaricata and fynbos on a gradient of high to low soil nutrients and carbon. we also compared in more detail the differences in nitrogen and phosphorus amongst the three vegetation types. although the results were site depended, both nitrogen and phosphorus concentrations appeared to be low in fynbos soils, with higher values found in forest and v. divaricata soils. it was only at plateau where phosphorus concentration was higher in fynbos soils compared with the other vegetation types. the lower nutrient status of fynbos soils is not unexpected, as previous work has shown that fynbos soils are nutrient poor, with low concentrations and availability of nitrogen and phosphorus (richards, stock & cowling 1997; stock & lewis 1986; stock, wienand & baker 1995; witkowski & mitchell 1987). fynbos vegetation has several adaptations to low nutrient availability, including cluster (e.g. proteoid, capillaroid and dauciform) roots and mycorrhizal associations that confer efficient nutrient uptake (allsopp & stock 1993; lamont 1982) and high nutrient-use efficiency (stock & lewis 1984; van daalen 1981). the higher phosphorus concentration and content of fynbos soils at plateau was an unexpected result. however, the fynbos area at this site was farmed until recently and we cannot rule out that phosphorus-rich fertilisers were used at the site. as phosphorus is bound quite strongly by soils, it is not easily leached from soils. this could explain the higher phosphorus values found for the top 20 cm, despite the lower phosphorus values in deeper soils compared with the other vegetation types.the establishment of forest on fynbos soils is not an uncommon phenomenon and forest invasion into fynbos during prolonged fire-free periods has been reported for many sites (geldenhuys 2004; kruger 1984; manders 1990). although fynbos soils are generally low in nitrogen (stock & lewis 1986) and phosphorous (witkowski & mitchell 1987), van daalen (1984) has shown that in the southern cape forest can establish on most soils. fynbos and forest soils seldom appeared to differ and therefore van daalen (1984) described any existing differences as an artefact of the vegetation. van daalen (1981) rejected the hypothesis that the poorer nutrient status of fynbos soils precludes the colonisation of fynbos by forest species. bond (2010) supported this when he showed that most fynbos soils have enough micronutrients to build forest when deeper soil layers are included. however, van daalen (1981) found no regeneration of forest in fynbos at some sites in our study area and attributed this to the dependence of forest species on the ‘closed’ nutrient cycling of the forest ecosystem, where nutrients are tightly and efficiently cycled within the root zone. increasing research findings point to vegetation feedbacks having significant effects on soil fertility to explain establishment of trees on nutrient-poor fynbos soils (vitousek 1990; vitousek & walker 1989; wedin & tilman 1990). for example, invasive australian acacia species invade fynbos over vast areas in south africa, with profound changes in soil as a result (higgins et al. 1999; van wilgen et al. 2001). these alien nitrogen-fixing acacia species regenerate rapidly after fire (milton & hall 1981) and the subsequent large inputs of high-nitrogen leaf litter elevate total soil nitrogen and organic matter (stock et al. 1995; witkowski 1991a; yelenik et al. 2004). alien acacias have been found to have higher leaf nitrogen concentrations than indigenous vegetation and no differences in resorption were found between fynbos species and alien acacias during a study in another fynbos area, which meant that more nitrogen was returned to soil in the acacia litter (witkowski 1991b). a combination of high tolerance to low shade conditions and fast modification of the nutrient cycle thus results in successful invasion of alien acacias into fynbos. in contrast with alien acacias, forest trees typical of afrotemperate forests do not tolerate regeneration in sunny micro-climates well and are, as a rule, quite sclerophyllous (van daalen 1981). this leads to low decomposition rates (witkowski 1991b). van daalen (1984) suggests that for forest to re-establish in a fynbos habitat or after disturbance, forest precursor species that can tolerate low nutrient levels and summer water deficits establish first. decomposition and nutrient return were found to be higher in a study comparing strandveld (sclerophyllous vegetation with a higher tree component on the west coast of south africa) with fynbos (witkowski 1991b). although nitrogen-fixing trees increase soil nitrogen over time owing to nitrogen-enriched litter being returned to the soil, soil nutrients may also be enhanced in other ways. trees may increase soil nutrients by: • drawing nutrients from deep soil layers and areas beyond the canopy and depositing them in the form of root exude, litter and/or canopy leaching (belsky 1994; jobbágy & jackson 2004; kellman 1979; scholes & hall 1996) • acting as an atmospheric dust trap and increasing nutrients from canopy leaching (bernhard-reversat 1988; escudero et al. 1985) • attracting mammals and roosting birds by providing shade and refuge, who then add nutrients by means of their droppings (belsky 1994; giorgiadis 1989). when forest precursor trees establish in the current study area, litter builds up over time and eventually a dense mat of decomposing root material and organic material is formed (geldenhuys 2011; geldenhuys & theron 1994). shallow feeder roots depend on the nutrients released from decomposing material and take up these nutrients before they are leached; in this way nutrients are increased and cycled quite tightly within the rooting zone. small additions of nitrogen to nutrient-poor fynbos soils resulted in significant changes in the growth-form composition of the vegetation and led to increases in graminoid forms of fynbos elsewhere (witkowski 1991b). some fynbos species show decreased growth rates with increased levels of nitrogen or phosphorus in the soil (witkowski 1989). some restios have been shown to decrease allocation to reproduction when treated with nitrogenand phosphorus-rich fertilisers (witkowski 1990). apart from decreasing the competitiveness of fynbos vegetation and restoring the nutrient cycle, forest precursor species provide suitable shade, with accompanying higher dispersal opportunities and ameliorated soil water conditions. once these conditions re-establish, forest trees re-establish (geldenhuys 2004). if nitrogen-fixing v. divaricata enriches soils in a similar manner to the invasive acacias, then the nutrient cycle should be re-established much faster than under sclerophyllous forest precursors. invasive acacias show faster decomposition rates and nitrogen return compared with the local forest precursor tree pterocelastrus tricuspidatus as a result of nitrogen-rich litter (witkowski 1991b). similarly, we found that v. divaricata litter (and roots) tends to have higher nitrogen concentrations than either forest or fynbos litter (with accompanying low c:n). in some ways v. divaricata has a very similar ecology to that of australian acacias in that it recruits rapidly after fire and has the potential to fix nitrogen. however, some major differences between invasive acacias and v. divaricata are that v. divaricata is not as persistent and long lived as the acacias. this allows other forest species to establish over time, as virgilia spp. stands usually die off naturally after approximately 20 years owing to caterpillars of the leto venus moth ringbarking trees (c. geldenhuys, pers. comm., january 2011). virgilia divaricata stands are typically also less dense than invasive acacia stands, thereby allowing gaps for indigenous shade-tolerant forest species to establish. conclusion top ↑ we compared soil fertility between three co-occurring vegetation types: v. divaricata-dominated stands with forest and fynbos on either side. our findings suggested that v. divaricata enhances soil fertility. however, owing to the nature of our work, the link between v. divaricata and higher soil nutrients is only tentative and further experimental work should be undertaken to confirm that v. divaricata enhances soil nutrients. nonetheless, owing to its ability to grow fast and establish without shade, we suggest that v. divaricata can be gainfully used to restore the ‘closed’ nutrient cycle found beneath forest and in this way aid in efficient forest regeneration. in contrast, maintaining ‘healthy’ fynbos (i.e. with a competitive advantage over trees) requires regular fires, both to maintain the ideal micro-climate for fynbos vegetation and to disrupt the nutrient cycle associated with tree-dominated vegetation. acknowledgements top ↑ the authors want to thank the following people and organisations: btech students oiva akudhenga, iikela sioni, muzuma uakendisa, don de swart, andrew jackson and roxane moulet for performing fieldwork, sanparks for allowing us to work in the garden route national park, the haupt family for allowing us to work at plateau and mike cameron for comments on an earlier draft of the manuscript. competing interests the authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article. authors’ contributions c.c. (nelson mandela metropolitan university) and b.j.w. (nelson mandela metropolitan university) designed the research questions and organised fieldwork. c.c. wrote the text and b.j.w. assisted in creating artwork. references top ↑ allsopp, n. & stock, w.d., 1993, ‘mycorrhizal status of plants growing in the cape floristic region, south africa’, bothalia 23, 91–104.barnes, p.w. & archer, s., 1999, ‘tree-shrub interactions in a subtropical savanna parkland: competition or facilitation?’, journal of vegetation science 10, 523–536. http://dx.doi.org/10.2307/3237187 belsky, a.j., 1994, ‘influences of trees on savanna productivity: tests of shade, nutrients, and tree-grass competition’, ecology 75, 922–934. http://dx.doi.org/10.2307/1939416 bernhard-reversat, f., 1988, ‘soil nitrogen mineralization under a eucalyptus plantation and a natural acacia forest in senegal’, forest ecological management 23, 233–244. http://dx.doi.org/10.1016/0378-1127(88)90054-0 bond, w.j., 1981, ‘vegetation gradients in the southern cape mountains’, msc thesis, university of cape town. bond, w.j., 2010, ‘do nutrient-poor soils inhibit development of forests? a nutrient stock analysis’, plant and soil 334, 47–60. http://dx.doi.org/10.1007/s11104-010-0440-0 campbell, b.m. & killick, d.j.b., 1985, a classification of the mountain vegetation of the fynbos biome, memoirs of the botanical survey of south africa, vol. 50, botanical research institute, department of agriculture, south africa. chapman, j.l. & reiss, m.j., 1992, ecology principles and applications, cambridge university press, cambridge, uk. crain, c.m. & bertness, m.d., 2006, ‘ecosystem engineering across environmental gradients: implications for conservation and management’, bioscience 56, 211–218. http://dx.doi.org/10.1641/0006-3568(2006)056[0211:eeaegi]2.0.co;2 escudero, a., garcia, b., gomex, j.m. & luis, e., 1985, ‘the nutrient cycling in quercus rotundifolia and q. pyrenaica ecosystems (‘dehesas’) of spain’, acta oecologica. oecological plantarum 6, 73–86. geldenhuys, c.j., 1982, ‘the management of the southern cape forests’, paper presented at the eighth regular meeting, sarccus standing committee for forestry, louis trichardt, 09–13th october 1981. geldenhuys, c.j., 1991, ‘distribution, size and ownership of forests in the southern cape’, south african forestry journal 158, 51–68. http://dx.doi.org/10.1080/00382167.1991.9630375 geldenhuys, c.j., 1994, ‘bergwind fires and location pattern of forest patches in the southern cape landscape, south africa’, journal of biogeography 21, 49–62. http://dx.doi.org/10.2307/2845603 geldenhuys, c.j., 2004, ‘concepts and processes to control invader plants in and around natural evergreen forest in south africa’, weed technology 18, 1386–1391. http://dx.doi.org/10.1614/0890-037x(2004)018[1386:captci]2.0.co;2 geldenhuys, c.j., 2011, ‘how to live with trees and keep them healthy’, south african forestry magazine, june, pp. 36–37. geldenhuys, c.j. & theron, j.m., 1994, ‘litterfall and nutrient cycling in mixed evergreen forest near george, south africa’, report for-dea 746, division of forest science and technology, csir, pretoria. giorgiadis, n.j., 1989, ‘microhabitat variation in an african savanna: effect of woody cover and herbivores in kenya’, journal of tropical ecology 5, 93–108. http://dx.doi.org/10.1017/s0266467400003254 higgins, s.i., richardson, d.m., cowling, r.m. & trinder-smith, t.h., 1999, ‘predicting the landscape-scale distribution of alien plants and their threat to plant diversity’, conservation biology 13, 303–313. http://dx.doi.org/10.1046/j.1523-1739.1999.013002303.x jobbágy, e.g. & jackson, r.b., 2004, ‘the uplift of soil nutrients by plants: biogeochemical consequences across scales’, ecology 85, 2380–2389. http://dx.doi.org/10.1890/03-0245 kellman, m., 1979, ‘soil enrichment of neotropical savanna trees’, journal of ecology 67, 565–577. http://dx.doi.org/10.2307/2259112 kindt, r. & coe, r., 2005, tree diversity analysis: a manual and software for common statistical methods for ecological and biodiversity studies, world agroforestry centre (icraf), nairobi. kruger, f.j., 1984, ‘effects of fire on vegetation structure and dynamics’, in p. de v. booysen & n.m. tainton (eds.), ecological effects of fire in south african ecosystems, pp. 219–244, springer verlag, berlin. http://dx.doi.org/10.1007/978-3-642-69805-7_10 laland, k.n., odling-smee, f.j. & feldman, m.w., 1999, ‘evolutionary consequences of niche construction and their implications for ecology’, proceedings of the national academy of sciences of the united states of america 96, 10242–10247. http://dx.doi.org/10.1073/pnas.96.18.10242 lamont, b., 1982, ‘mechanisms for enhancing nutrient uptake in plants, with particular reference to mediterranean south africa and western australia’, the botanical review 48, 597–689. http://dx.doi.org/10.1007/bf02860714 legendre, p. & legendre, l., 1998, numerical ecology, 2nd edn., elsevier, amsterdam. manders, p.t., 1990, ‘fire and other variables as determinants of forest/fynbos boundaries in the cape province’, journal of vegetation science 1, 483–490. http://dx.doi.org/10.2307/3235782 manders, p.t. & richardson, d.m., 1992, ‘colonization of cape fynbos communities by forest species’, forest ecology and management 48, 277–293. http://dx.doi.org/10.1016/0378-1127(92)90150-8 manders, p.t., richardson, d.m. & masson, p.h., 1992, ‘is fynbos a stage in succession to forest? analysis of the perceived ecological distinction between two communities’, in b.w. van wilgen, d.m. richardson, f.j. kruger & h.j. van hensbergen (eds.), fire in south african mountain fynbos: ecosystem, community and species response at swartboskloof, pp. 81–107, springer verlag, berlin. [ecological studies vol. 93]. http://dx.doi.org/10.1007/978-3-642-76174-4_5 masson, p.h. & moll, e.j., 1987, ‘the factors affecting forest colonization of fynbos in the absence of recurrent fire at orange kloof, cape province, south africa’, south african journal of forestry 143, 5–10. http://dx.doi.org/10.1080/00382167.1987.9630294 milton, s.j. & hall, a.v., 1981, ‘reproductive biology of australian acacias in the south-western cape province, south africa’, transactions of the royal society of south africa 44, 465–487. http://dx.doi.org/10.1080/00359198109520589 mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, south african national biodiversity institute, pretoria. [strelitzia no. 19]. musil, c.f., 1993, ‘effect of invasive australian acacias on the regeneration, growth and nutrient chemistry of south african lowland fynbos’, journal of applied ecology 30, 361–372. http://dx.doi.org/10.2307/2404637 nelson, d.w. & sommers, l.e., 1996, ‘total carbon, organic carbon, and organic matter’, in d.l. sparks, a.l. page, p.a. helmke, r.h. loeppert, p.n. soluanpour, m.a. tabatabai, c.t. johnston & m.e. sumner (eds.), methods of soil analysis part 3: chemical methods, pp. 961–1010, soil science society of america, inc. and american society of agronomy, inc., madison, wisconsin, usa. odling-smee, j., laland, k.n. & feldman, m.w., 1996, ‘niche construction’, the american naturalist 147, 641–648. http://dx.doi.org/10.1086/285870 palgrave, k.c., 1981, trees of southern africa, struik, cape town. phillips, j.f.v., 1926, ‘virgilia divaricata capensis lam. (‘keurboom’): a contribution to its ecology and sylviculture’, south african journal of science 23, 435–454. r development core team, 2011, r: a language and environment for statistical computing, r foundation for statistical computing, vienna. richards, m.b., stock, w.d. & cowling, r.m., 1997, ‘soil nutrient dynamics and community boundaries in the fynbos vegetation of south africa’, plant ecology 130, 143–153. http://dx.doi.org/10.1023/a:1009742225383 scholes, r.j. & hall, d.o., 1996, ‘the carbon budget of tropical savannas, woodlands, and grasslands’, in a. breymeyer, d. hall, j. melillo & g. agren (eds.), modelling terrestrial ecosystems, pp. 69–100, wiley, chichester. stock, w.d. & lewis, o.a.m., 1984, ‘uptake and assimilation of nitrate ammonium by an evergreen fynbos shrub species protea repens l. (proteaceae)’, new phytologist 97, 261–268. http://dx.doi.org/10.1111/j.1469-8137.1984.tb04130.x stock, w.d. & lewis, o.a.m., 1986, ‘soil nitrogen and the role of fire as a mineralizing agent in a south african coastal fynbos ecosystem’, journal of ecology 74, 317–328. http://dx.doi.org/10.2307/2260257 stock, w.d., wienand, k.t. & baker, a.c., 1995, ‘impacts of invading n2-fixing acacia species on patterns of nutrient cycling in two cape ecosystems: evidence from soil incubation studies and 15n natural abundances’, oecologia 101, 375–382. http://dx.doi.org/10.1007/bf00328825 van daalen, j.c., 1981, ‘the dynamics of the indigenous forest-fynbos ecotone in the southern cape’, south african forestry journal 119, 14–23. http://dx.doi.org/10.1080/00382167.1981.9630218 van daalen, j.c., 1984, ‘distinguishing features of forest species on nutrient-poor soils of the southern cape’, bothalia 1/2, 229–239. van der merwe, i., 2011, green heritage: trees, forests and woodlands of south africa, department of agriculture, forestry and fisheries, pretoria. van wilgen, b.w., richardson, d.m., le maitre, d.c., marais, c. & magadlela, d., 2001, ‘the economic consequences of alien plant invasions: examples of impacts and approaches to sustainable management in south africa’, environment, development and sustainability 3, 145–168. http://dx.doi.org/10.1023/a:1011668417953 walkley, a., 1947, ‘a critical examination of a rapid method for determining organic carbon in soils: effect of variations in digestion conditions and of inorganic soil constituents’, soil science 63, 251–263. http://dx.doi.org/10.1097/00010694-194704000-00001 vitousek, p.m., 1990, ‘biological invasions and ecosystem processes: towards an integration of population biology and ecosystem studies’, oikos 57, 7–13. http://dx.doi.org/10.2307/3565731 vitousek, p.m. & walker, l.w., 1989, ‘biological invasions by myrica faya in hawai’i: plant demography, nitrogen fixation, ecosystem effects’, ecological monographs 59, 247–265. http://dx.doi.org/10.2307/1942601 wedin, d.a. & tilman, d., 1990, ‘species effects on nitrogen cycling: a test with perennial grasses’, oecologia 84, 433–441. white, f., 1978, ‘the afromontane region’, in m.j.a. werger (ed.), biogeography and ecology of southern africa, pp. 463–513, junk, the hague. http://dx.doi.org/10.1007/978-94-009-9951-0_11 witkowski, e.t.f., 1989, ‘effects of nutrients on the distribution of dry mass, nitrogen and phosphorus in seedlings of protea repens (l.) l. (proteaceae)’, new phytologist 112, 481–487. http://dx.doi.org/10.1111/j.1469-8137.1989.tb00341.x witkowski, e.t.f., 1990, ‘effect of nutrient additions on the allocations of dry mass, phosphorus and nitrogen in the cape fynbos plant, thamnochortus punctatus pill. (restionaceae)’, south african journal of ecology 1, 38–44. witkowski, e.t.f., 1991a, ‘effects of invasive alien acacias on nutrient cycling in the coastal lowlands of the cape fynbos’, journal of applied ecology 28, 1–15. http://dx.doi.org/10.2307/2404109 witkowski, e.t.f., 1991b, ‘growth and competition between seedlings of protea repens (l.) l. and alien invasive, acacia saligna (labill.) wendl., in relation to nutrient availability’, functional ecology 4, 101–110. http://dx.doi.org/10.2307/2389560 witkowski, e.t.f. & mitchell, d.t., 1987, ‘variations in soil phosphorus in the fynbos biome, south africa’, journal of ecology 75, 1159–1171. http://dx.doi.org/10.2307/2260320 yarranton, g.a. & morrison, r.g., 1974, ‘spatial dynamics of a primary succession: nucleation’, journal of ecology 62, 417–428. http://dx.doi.org/10.2307/2258988 yelenik, s.g., stock, w.d. & richardson, d.m., 2004, ‘ecosystem level impacts of invasive acacia saligna in the south african fynbos’, restoration ecology 12, 44–51. http://dx.doi.org/10.1111/j.1061-2971.2004.00289.x filelist convert a pdf file! koedoe 19: jj3-144 (976) a survey of the mammals occurring in the golden gate highlands national park i. l. rautenbach transvaal museum p.o. box 413 pretoria 0001 abstract this paper reports on a survey of the mammals of the golden gate highlands national park, republic of south africa. fifty-seven species are mentioned, the majority recorded through material or sight records. those species which may occur in the park, as deducted from their overall distribution ranges, or from other indirect observations such as spoor or droppings, are considered as well. habitat preferences are mentioned wherever possible, and the conservation status and relocation histories of the antelope species are quoted. introduction the main object of this study is to provide a checklist of mammals for the golden gate highlands national park, republic of south africa, but stems from an interest in the taxonomy and zoogeography of the mammals of southern africa. these interests can only be successfully pursued through intensive and continuous collecting. when the late dr. n.]. van der merwe, and later his successor dr. g. de graaff, prompted myself and my predecessor mr c. g. coetzee to produce such a checklist for this park, we obligingly agreed without hesitation. this park in the northeastern orange free state on the border of lesotho, was proclaimed in 1963, primarily for its scenic beauty. six farms were dispossessed, comprising a total area of 4 792 ha of mountain sourveld (van rensburg 1968). the park constitutes the upper catchment area of the little caledon river, and varies in altitude from 1 800 m at the riverbed to 3 000 m at the peak of rhebokkop. the watershed between the orange-and vaal river systems is at the eastern boundary of the park. the national parks board of trustees, in agreeance with its conservation policy, subsequently reintroduced the game animals that long ago disappeared from the area as a result of human settlement and agricultural interests (liebenberg 1964; penzhorn 1971) . for obvious reasons the board could not reintroduce some of the 133 former residents such as lion, wild dog, hippopotamus and elephant. primarily as a result of pressing conservation problems in other national parks, very little research has thus far been conducted on the fauna and flora of the golden gate highlands national park. roberts (1969) pioneered research within the park by conducting a botanical survey. material and methods in aid of this survey, the area has been visited three times, i.e. for a ten-day period during july 1965, as well as two additional ten-day periods during may and november 1969. the species discussed in the text include not only those whose presence have been confirmed through sight or material records, but also those whose occurrence are suspected through circumstantial evidence such as general species range, or spoor and faeces encountered in the park. this latter group comprises predominantly mammals in the medium-size range which migrate or wander mostly unnoticed over great distances. specimens of only those species still presenting taxonomic problems were collected, being without exception in the smallermammal species range. specimens were procured through the normal mammal-collecting techniques, which need no elaboration here as it has been extensively described in the literature. trapping was done predominantly in those area offering the best habitat. the animals collected were prepared as standard study-material and have been incorporated in the permanent collection of the transvaal museum. results the following species have been noted to occur, or may occur in the park: erinaceusfrontalis smith, 1831 hedgehog krimpvarkie although as yet not recorded from the park, it almost certainly occurs here, since suitable habitat is available, and since the park falls within the known range of the species. it has been recorded from the nearby reitz and senekal districts (lynch 1975). myosorex varius smuts, 1832 forest shrew bos skeerbek a large series of 17 males and 21 females have been collected, mostly on the grassy slopes of bakenkop. a few specimens were also procured in dense grass on the riverbanks near wilgenhof. 134 crocidura jlavescens i. geoffroy, 1827 red musk shrew rooi skeerbek although this species has never been recorded from the o.f.s., it is almost certain to occur in the park, as it falls within the species range and affords suitable habitat. faeces, presumably of this species, was found in old rock walls, but attempts at trapping animals were unsuccessful. crocidura cyanea duvernoy, 1838 reddish-grey musk shrew rooigrys skeerbek a single specimen was procured in dense grass along a stream. amblysomus hottentotus a. smith, 1829 hottentot golden mole hotnotkruipmol four females were collected during november 1969 under willow trees on the banks of the small caledon river. at this locality this species coexists with the rodent mole (cryptomys hottentotus lesson, 1826) in the same area, albeit in separate tunnel systems. on the surface the tunnel systems of these two unrelated fossorial animals cannot be distinguished. the substrate was of a black clayish type . two of the females were reproductively active. chlorotalpa sclateri broom, 1907 sclater's golden mole sclaterse kruipmol no material has as yet been collected within the park, but three specimens in the transvaal museum collection have been collected from the n'earby clocolan district (2827dc). the characteristic shallow subsurface tracks of this species have been found in the low-lying areas of the park. pipistrellus kuhli n atterer, 1817 kuhl's pipistrelle kuhlse vlermuis recorded by lynch (j 975) from areas adjacent to the park. the species is as a consequence almost certain to occur within the boundaries of the park. eptesicus capensis a. smith, 1829 cape serotine kaapse dakvlermuis lynch (op . cit.) recorded this species within 25 km east of golden gate, and it is therefore not unlikely that it will in time be recorded from within the park. papio ursinus kerr, 1792 chacma baboon kaapse bobbejaan a fairly large troop has regularly been observed in the park on the mountain slopes and crests. 135 lepus saxatilis f. cuvier, 1823 scrub hare kolhaas a single specimen was collected during november 1969. hares of the genus lepus are not at all common in golden gate. lynch (1975) unfortunately did not verify the identification of this specimen while extracting transvaal museum records for his paper, and consequently incorrectly cited it under lepus capensis linnaeus, 1758. lepus capensis linnaeus, 1758 cape hare vlakhaas it is not impossible that this species is to be found at golden gate, but no material exists from the near vicinity of the park to substantiate such a speculation. lynch (1975) claims the presence of this species in the park and vicinity, but apart from a misidentified specimen recorded, the remainder of his records are non-material. he did not state whether the latter are from roadkills or sightings. it is of course impossible to distinguish between l. capensis and l. saxatilis in the wild state. pronolagus rupestris a. smith, 1834 smith's red hare smith se rooihaas two specimens, a male and a female, were collected in the hills behind glen reenen camp during november 1969. the female was lactating. judging from the number of toilet-sites found, so characteristic of the genus, this species is relatively abundant in the park. the measurement parameters quoted by petter ( 197 i) in his identification key are not clear, and as a consequence the key and taxonomic approach of ellerman, morrison-scott and hayman (1953) is followed here. cryptomys hottentotus lesson, 1826 common mole rat hotnot grysmol three specimens were collected in the park to date. this species occurs in the low-lying areas as well as on the higher platos and hillslopes, indicating a wider habitat tolerance than the golden moles. hystrix africaeaustralis peters, 1852 cape porcupine ystervark although this species has never been observed during collecting trips, its presence has been confirmed by several quills found in the veld, particularly at the bases of rockfaces on the lower hillslopes. pedetes capensis forster, 1778 springhare springhaas springhares have never been observed during night-hunting operations, but they almost certainly occur here. lynch (i 975) reports sight records from the near vicinity of the park . 136 graphiurus murinus desmarest, 1822 forest dormouse boswaaierstertmuis although the vernacular name of this insectivorous rodent denotes its preferred habitat, it is also to be found in rocky places, as for instance the golden gate material. all three specimens were trapped amongst rock debris at the foot of the sentinel. the only female in the series was pregnant upon collection during november; four foetuses, one in the left and three in the right uterus horn. aethomys namaquensis smith, 1834 namaqua rock rat namakwalandse klipmuis this is an abundant inhabitant of the rocky mountain slopes of golden gate. aethomys chrysophilus de winton, 1897 african bush rat afrikaanse bosrot not recorded through this project. lynch (i 97 5) claims its presence in the near vicinity of this park, which entails a 100 km westwards extension of the known range as defined by davis (1974). however, the streambanks lined with ouhout leucosidea sericea as found in the park, do not appear to offer suitable habitat when compared with the bushveld conditions selected over the rest of the species range. leggada minutoides a. smith, 1834 dwarf mouse dwergmuis a series of four was collected in the long grass near the brandwag camp. mus musculus linnaeus, 1758 house mouse huismuis a single specimen was procured in the non-white compound. praomys natalensis smith, 1834 multimammate mouse vaalveldmuis a common rodent in the denser grass oflow-lying areas. rattus rattus linnaeus, 1758 house rat huisrot several specimens were trapped in the living quarters of the bantu staff of the park . rhabdomys pumilio sparrman, 1784 striped mouse streepmuis a common rodent in the park, especially in dense tall grass. according to coetzee (in litt.), it was also found to construct its grassnests in old rockwalls up to almost a meter above ground level, which is a deviation from the normal behaviour. 137 dendromus melanotis smith, 1834 grass climbing mouse grassklimmuis a single specimen was collected in long grass along the road near the brandwag camp. dendromus mystacalis heuglin, 1863 lesser climbing mouse kleiner klimmuis four specimens were collected, all from dense tall grass either along the road , the river or the base of a hill at wilgenhof. one specimen had the remains of insects in its stomach. this is a new record for the orange free state as the result of a westwards range extention along the 29° latitude . malacothrix typica smith, 1834 large-eared mouse grootoormuis recorded by lynch (1975) in the bethlehem/kestell districts, near the park . mystromys albicaudatus smith, 1834 white-tailed rat witstertrot as yet not recorded from the park itself. it is essentially a grasslandplains species, but may in time prove to be a resident of golden gate as it has been recorded from the nearby districts (davis 1974; lynch 1975). tatera brantsi a. smith, 1834 highveld gerbil hoeveldse nagmuis several colonies were found in sandy patches oflow-iying areas. otomys irroratus brants, 1827 vlei rat vleirot a common rodent of the grassy areas near water. somewhat atypically, the species was also recorded on the lower hillslopes, especially where the ground is soggy at the bases of rocks. coetzee collected two specimens with pure-white hindquarters in the park during a museum collecting trip in 1965. according to him (in lilt.) this is the only instance of partial albinism known in this species south of zaire. note: without examining the specimen, lynch (1975) accepts as authentic the identity of an otomys specimen from golden gate, housed in the tm collection, but misidentified as o. unisulcatus. the unfortunate result is that he plots the occurrence of o. unisulcatus at golden gate, some 200 km out of range (davis 1974) in completely atypical habitat. i examined the specimen in question, and found it answers to the description of o. irroratus. consequently lynch's (1975) record of o. unisulcatus from golden gate is rejected here. 138 otomys sloggetti thomas, 1902 rock karoo rat/ice rat klip karroorot/ysrot this species certainly occurs in the park, but unfortunately no substantiating material could as yet be collected. the typical otomys faeces have, however, been found higher up on the mountain slopes in typical o. sloggetti habitat. vulpes chama a. smith, 1833 silver fox silwervos not recorded during our visits, but as this is a widely distributed species in the orange free state, individuals will undoubtedly occur in the park from time to time. canis mesomelas schreber, 177 5 black-backed jackal rooijakkals an individual was encountered during night-collecting operations. aonyx capensis schinz, 1821 cape clawless otter groototter the spoor and characteristic droppings of this species was found along the river as well as at the big dam in the game camp. lutra maculicollis lichtenstein, 1835 spotted-necked otter klein-otter the presence of this species in the park could as yet not be confirmed. since it has been reported from the nearby districts (lynch 1975), a resident population, or at least migrants through the park, is very likely. jctonyx striatus perry, 1810 striped polecat stinkmuishond individuals have been observed in the park, but as yet none could be collected. a roadkill is reported by coetzee (in litt.) on the main road near the park entrance. genetla trigrina tigrina schreber, 1776 large-spotted genet grootkolmuskejaatkat two specimens were live-trapped in the shrub along watercourses. after deliberation with c. g. coetzee and j. pringle (pers. comm.) these specimens were assigned to this subspecies, in difference to lynch (1975) who recognizes only g. genetta linnaeus, 1758 in this area and the rest of the orange free state. consequently this constitutes a new record for this province . cynictis penicillata g. cuvier, 1829 yellow mongoose geelmeerkat no specimens of this species were collected, but a colony to the east of the game camp has often been observed. 139 herpe5tes pulverulentes wagner, 1839 cape grey mongoose kleingrysmuishond certainly the most common carnivore in the park, and several were livetrapped of which two were prepared as specimens. they seem to prefer the wooded banks of the various streams flowing through the park. a female collected during november was lactating. ichneumia albicauda g. cuvier, 1829 white-tailed mongoose witstertmuishond lynch ( 1975) reports this species from areas adjacent to golden gate. as the park offers suitable habitat this animal may in time be shown to occur here as well. alilax paludinosus g. cuvier, 1777 marsh mongoose kommetjiesgatmuishond the spoor and faeces of this animal was found, but it was not collected or seen during our visits. protele5 cri5tatus spaarman, 1783 aardwolf maanhaarjakkals this species was never encountered, but spoor seen in the park was probably that of this animal. the fact that the aardwolf is widely distributed in the orange free state renders it very likely to be present in the park. felis caracal schreber, 1776 caracal rooikat spoor were found on several occasions, while a former ranger has seen a caracal in the park. felis lib,yea forster, 1780 cape wild cat vaalboskat several of these animals were seen during night collecting trips, but unfortunately none were collected, procavia capensis pallas, 1766 cape dassie klipdas three specimens were collected, of which one was heavily infected with lungworms. dassies are particularly abundant in golden gate. equus burehelli gray, 1824 burchell's zebra bontkwagga the burchell's zebra were reintroduced from the kruger national park (penzhorn 1971). the population has settled down well and is expanding (national parks board of trustees annual report 49: 197417 5). 140 phacochoerus aethiopicus pallas, i 766 warthog vlakvark reintroduced from the hluhluwe game reserve (penzhorn 197 i), but the animals have apparently not adapted very well. syncerus calfer sparrman, 1779 cape buffalo kaapse buffel reintroduced from the addo elephant national park (penzhorn 197 i) . taurotragus oryx pallas, i 766 eland reintroduced from the kalahari gemsbok national park, addo elephant national park, and the willem pretorius game reserve (o.f.s.) (penzhorn 1971 ). the population is slowly expanding (national parks board of trustees annual report 49). redunca arundinum boddaert, 1785 reedbuck rietbok reintroduced from northam (tv!.) (penzhorn 197 i), but these animals are adapting poorly to their new environment. rrduncafulvorufula at:zelius, 1815 mountain reedbuck rooiribbok reintroduced from luckhoff (o.f.s.) (penzhorn 197 i). population numbers have not increased significantly since 1971 (national parks board of trustees annual reports 46-48). alcelaphus buselaphus caama g. cuvier, 1804 cape hartebeest rooihartbees reintroduced from the addo elephant national park, kalahari gemsbok national park, willem pretorius game reserve (o.f.s.), rooipoort estates (kimberley, c.p.), and setlagodi (c.p.) (penzhorn 197 i). only three have survived since 1971 and they are not reproducing (national parks board of trustees annual reports 46-49). damaliscus dorcas phillipsi harper, 1939 blesbok reintroduced from the mountain zebra national park, willem pretorius game reserve (o.f.s.), van riebeeck nature reserve (pretoria, tv!.), kestell (o.f.s.), petrus steyn (o.f.s.), pietersburg (tv!.), and nduli national reserve (umtata, transkei) (penzhorn 197 i). the population has settled down very well and is rapidly expanding (national parks board of trustees annual reports 46-49). 141 connochaetes gnou zimmermann, 1780 black wildebeest swartwildebees reintroduced from the willem pretorius game reserve (o.f.s.), odendaalsrus (o.f.s.), and makwassie (tvl.) (penzhorn 1971). the population is viable and expanding rapidly (national parks board of trustees annual reports 46-49). antidorcas marsupialis zimmermann, 1780 springbok reintroduced from the bontebok national park, kalahari gemsbok national park, mountain zebra national park, and cookhouse (c.p.) penzhorn 197 i). the population is rapidly expanding (national parks board of trustees annual reports 46-49) . the animals were reintroduced from the ranges of two subspecies, i.e. a. m. marsupia/is from the republic and a. m. hofmeyri thomas, 1926 from the kalahari gemsbok national park. the present golden gate population therefore could elucidate the hitherto unsettled state of subspeciation in this species. ourebia ourebi zimmermann, 1783 oribi oorbietjie reintroduced from grey town (natal) (penzhorn 1971). births have been recorded (national parks board of trustees annual report 48) and the population seems to be thriving. raphicerus campestris thunberg, 1811 steenbok the present day population is endemic (penzhorn 197 i), but its status is unknown. peiea capreo/us forster, 1790 grey rhebuck vaalribbok the existing population is also endemic (penzhorn 1971), and appears to be stable although exact numbers are not known. acknowledgements i wish to express my gratitude to the national parks board of trustees for permission to collect material from the golden gate highlands national park, as well as for accommodation provided . my sincere thanks are due to mr a. van zyl, the ranger of the park, for his assistance and advice. the late dr nico van der merwe initiated the project, and later dr g. de graaff encouraged the completion of the work, and rendered advice and assistance whenever needed . this project was carried out and the results are published with the permission of the director of the transvaal museum. 142 references davis, d. h. s. 1974. the distribution of some small southern african mammals (mammalia:insectivora, rodentia). ann. transvaal mus. 29(9):135-184. ellerman,j. r., t. c. s. morrison-scott and r. w. hayman, 1953. southern african mammals 1758 to 1951: a reclassification. trustees of the british museum. liebenberg, l. c. c. 1964. die grotere soogdiere wat vroeer dae voorgekom het in die omgewing van die golden gate hooglandpark. koedoe 7 :99-104. lynch, c. d. 1975. the distribution of mammals in the orange free state, south africa. navorsinge van die nasionale museum 3(6): 109139. penzhorn, b. l. 1971. a summary of the re-introduction of ungulates into south african national parks (to 31 december 1970). koedoe 14: 145-159. petter, f. 1971. order lagomorpha. in: meester, j. and h. w. setzer (eds.) 1971. the mammals of africa: an identification manual . washington, d.c . : smithsonian institution press. roberts, b. r. 1969. the vegetation of golden gate highlands national park. koedoe 12:15-28. van rensburg, a. p. j. 1968. golden gate die geskiedenis van twee plase wat 'n nasionale park geword het. koedoe 11 :83-138. 143 page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 133.pdf page 1 van staden woud.qxd a floristic analysis of forest and thicket vegetation of the marakele national park p.j. van staden & g.j. bredenkamp van staden, p.j. and g.j. bredenkamp. 2006. a floristic analysis of forest and thicket vegetation of the marakele national park. koedoe 49(1): 15–32. pretoria. issn 00756458. one of the major plant communities identified in the marakele national park was forest. it became clear that this major forest community contained various forest and thicket communities. relevés compiled in the forest were classified by twinspan and braun-blanquet procedures identified six communities that are hierarchically classified. the forests dominated by podocarpus latifolius and widdringtonia nodiflora represent afromontane forests, whereas the buxus macowanii-dominated dry forests and olea europaea subsp. africana represent northern highveld forests. a further group of communities represent thickets on termitaria with floristic affinities to both savanna and forest. the floristic composition and relationships of the forest and thicket communities are discussed. key words: afromontane forest, afrotemperate forest, classification, national park, termitaria, thicket, twinspan. pj van staden, centre for wildlife management, university of pretoria, pretoria, 0001 republic of south afirca; gj bredenkamp , department of botany, university of pretoria, pretoria, 0001 republic of south afirca (george.bredenkamp@.up.ac.za). issn 0075-6458 15 koedoe 49/1 (2006) introduction in an overview of the vegetation of marakele national park, van staden & bredenkamp (2005) recognised five major plant communities, of which one represents forests. these forests are mostly restricted to moist, sheltered valleys, and represent typical marakele afromontane forest (von maltitz 2003). although du preez et al. (1991) classified this type of forest as typical afromontane forests, the marakele forests exist under much drier conditions than in the drakensberg (von maltitz 2003). these forests occur in specific niches in deep valleys, protected gorges and ravines along the eastern and western slopes of the drakensberg mountain range. indeed, some of the marakele forests are very dry and represent a very rare and restricted plant community classified by von maltitz (2003) under northern highveld forests. van staden (2002) indicated that various plant communities occurred within these forests, though they are not yet described. this paper aims to describe the forest plant communities of the marakele national park, to provide an inventory of the biodiversity of the plant community at the plant species level. study area the study area was described in detail by van staden (2002) and van staden & bredenkamp (2005), and are therefore only briefly mentioned here. the study area covers 290.51 km² in the southwestern part of the limpopo province between 27°30'e– 27°45'e and 24°15's–24°30's. marakele national park is situated mainly in the waterberg moist mountain bushveld (low & rebelo 1996), in the savanna biome (rutherford & westfall 1994). the vegetation of the study area includes acocks’ van staden woud.qxd 2006/04/17 08:38 pm page 15 (1988) sour bushveld (veld type 20), mixed bushveld (veld type 18), sourish mixed bushveld (veld type 19) and northeastern mountain sourveld (veld type 8). the underlying parent rock of the biggest part of the study area consists of sandstone of the kransberg subgroup, sandriviersberg formation. sandstone of the matlabas subgroup, aasvoëlkop formation, occurs in the southwestern and southern parts; shale and mudstone of the matlabas subgroup aasvoëlkop formation, groothoek mudstone member; a conglomerate outcrop of the matlabas subgroup, aasvoëlkop formation are found in the western part. these rock formations resulted in the complex mountain topography of the waterberg. within this complex topography, the forests are restricted to the relatively moist kloofs, mostly with small rivers. the soils that have developed on the parent materials range from shallow sandy to deep sandy soils on sandstone and clayey soils on diabase and mudstone. the rainfall varies from 556 mm to 630 mm per annum and occurs mainly during the summer months. this is considerably lower than the rainfall limits described for forests in summer rainfall area by rutherford & westfall (1994). the study area experiences warm, wet summers with temperatures of up to 32 ºc and cool dry winters with frost in the low-lying areas. methods analysis in the overview of the vegetation of the marakele national park, van staden (2002) used 130 sample plots, located in a stratified random manner in physiographically and physiognomically homogeneous units, delineated on 1:50 000 scale aerial photographs (bredenkamp & theron 1978; westfall 1981; gertenbach 1987). the number of sample plots for each delineated physiographic-physiognomic unit was determined according to the size of each delineated unit. termitaria were not included in the placing of the sample plots. although they could be recognised on the aerial photographs, they were too small to delineate. additional sample plots were subjectively placed within the termitaria vegetation. for the purpose of this study, a standard sample plot size of 10 m x 20 m was fixed and used throughout the study area. this plot size is considered adequate for surveys in savanna vegetation by coetzee (1975), coetzee et al. (1976), westfall (1981), van rooyen (1983) and gertenbach (1987). at each sample plot a list is compiled of all the species that occur. a cover-abundance value was given to each species according to the cover-abundance scale, used by braun-blanquet and given by mueller-dombois & ellenberg (1974) and werger (1974), and modified by barkman et al. (1964): 5 any number of plants, with cover >75 % of the sample plot. 4 any number of plants, with cover > 50–75 % of the sample plot. 3 any number of plants, with cover > 25–50 % of the sample plot. 2a covering between 5–12 % of the sample plot area independent of abundance (indicated as a in the tables). 2b -covering between 13–25 % of the sample plot area independent of abundance (indicated as b in the tables). 1 numerous, with cover of 5 % or less. + individuals with cover of < 1 %. r usually a single individual with a cover of < 1 %. although van staden (2002) recorded various habitat variables, only the following habitat classes are used in this study (see table 1): the altitude of each sample plot was recorded using an altimeter and is given in meters. the slope of the terrain of each sample plot was measured in degrees, using an optical clinometer. the following classification of slope units (westfall 1981), were used in this study: symbol description class l level 0.00° 3.49° g gentle 3.50° 17.62° m moderate 17.63°36.39° s steep 36.40° the aspect of the terrain where each sample plot is situated was determined using a compass. aspect is given in the eight compass directions, namely: n north s south ne northeast sw southwest e east w west se southeast nw northwest koedoe 49/1 (2006) 16 issn 0075-6458 van staden woud.qxd 2006/04/17 08:38 pm page 16 issn 0075-6458 17 koedoe 49/1 (2006) the surface rock cover in each sample plot was estimated as a percentage of stones (> 20 mm diameter), boulders and rocky outcrops. the following five classes were used, based on its potential influence on mechanical use (ploughing) (van der meulen 1979; westfall 1981): symbol class description o < 1 % no limitation on mechanical utilisation l 1 4 % low limitation on mechanical utilisation m 5 34 % moderate limitation on mechanical utilisation h 35 84 % high limitation on mechanical utilisation v 85 100 % no mechanical utilisation possible land type was read from the land type maps. synthesis the full dataset was captured on the mainframe computer of the university of pretoria, in the bbnew software package. it was then exported to be used in the software package bbpc (bezuidenhout et al. 1996). after a twinspan classification (hill 1979), the output of the resulting classification was imported into a spreadsheet, for refinement by braun-blanquet procedures (behr & bredenkamp 1988). the final classification of the relevés was then interpreted for identification of major communities for the entire park. the relevés representing the forests and thickets were then extracted, subjected to twinspan (hill 1979) and exported to be used in the software package bbpc (bezuidenhout et al. 1996) for refinement by braun-blanquet procedures. the results are summarised in a phytosociological table (table 1). results the phytosociological classification of forests and thickets in the study area resulted in the identification of five forest communities with one of the communities having two sub-communities. olea europaea subsp. africana-diospyros whyteana major community van staden & bredenkamp (2005) identified the forest and thicket vegetation that occur on the marakele national park as the olea europaea subsp. africana-diospyros whyteana major community. obvious variation in plant species composition and dominance was observed within this major community. the floristic composition of the olea europaea subsp. africana-diospyros whyteana major community is given in table 1. van staden & bredenkamp (2005) described this major community as having a large number of diagnostic species, all being trees or shrubs, which distinguish this forest vegetation from any other plant community occurring in the park. these diagnostic species included pappea capensis, mimusops zeyheri, olea europaea subsp. africana, cussonia paniculata, diospyros whyteana, euphorbia ingens, rhus leptodictya, cryptolepis transvaalensis, podocarpus latifolius, maytenus undata, zanthoxylum capense, myrsine africana, canthium gilfillanii, ficus sur, calpurnia aurea, olea capensis and grewia occidentalis. although different plant communities can be recognised within these forests and bush clumps, certain species, e.g., olea europaea subsp. africana, ficus sur, cryptolepis transvaalensis, cussonia paniculata and euphorbia ingens were prominent throughout the range of this major community. the forest communities occur in the kloofs, or as thickets in bush clumps on south and east-facing slopes and also as dense thickets on termitaria on the plains. the kloofs are the most sheltered of the geomorphology classes found in the study area, with water in the watercourses draining down the kloofs. these kloofs are also protected against the veld fires that occur frequently in the adjacent savanna or grassland vegetation. geldenhuys (1994) described a similar situation for forests in the western cape. coetzee (1975) described a related community from the rustenburg nature reserve as hypoestes verticillaris-mimusops zeyheri forests, and westfall (1981) described a similar community from the farm groothoek in the waterberg as kloof forest communities on moderately deep soils in moist, sheltered van staden woud.qxd 2006/04/17 08:38 pm page 17 koedoe 49/1 (2006) 18 issn 0075-6458 ta bl e 1 p hy to so ci ol og ic al t ab le o f t he fo re st s an d bu sh c lu m ps o n m ar ak el e n at io na l p ar k van staden woud.qxd 2006/04/17 08:38 pm page 18 issn 0075-6458 19 koedoe 49/1 (2006) ta bl e 1 (c on tin ue d) van staden woud.qxd 2006/04/17 08:38 pm page 19 koedoe 49/1 (2006) 20 issn 0075-6458 ta bl e 1 (c on tin ue d) van staden woud.qxd 2006/04/17 08:38 pm page 20 issn 0075-6458 21 koedoe 49/1 (2006) ta bl e 1 (c on tin ue d) van staden woud.qxd 2006/04/17 08:38 pm page 21 koedoe 49/1 (2006) 22 issn 0075-6458 ta bl e 1 (c on tin ue d) van staden woud.qxd 2006/04/17 08:38 pm page 22 issn 0075-6458 23 koedoe 49/1 (2006) habitats. du preez et al. (1991) classified this type of forest as typical afromontane forests, where these forests occur in specific niches in deep valleys, protected gorges and ravines along the eastern and western slopes of the drakensberg mountain range. coetzee et al. (1976) described termitaria bush clump communities from the nylsvley nature reserve. these have diagnostic species similar to the bush clump communities on termitaria in the marekele national park. in the hierarchical phytosociological classification, the following plant communities are classified under the olea europaea subsp. africana-diospyros whyteana major community (table 1): 1 olea capensis-cheilanthes viridis forest 1.1 podocarpus latifolius-curtisia dentata forest 1.1.1 widdringtonia nodiflorapodocarpus latifolius moist short forest. 1.1.2 podocarpus latifolius-rothmannia capensis moist tall forest. 1.2 buxus macowanii-kirkia wilmsii dry low forest. 2 pappea capensis-ziziphus mucronata thickets and bush clumps 2.1 rhus leptodictya-mimusops zeyheri termitarium thickets. 2.1.1 rhus leptodictya-carissa bispinosa sub-community 2.1.2 rhus leptodictya-berchemia zeyheri sub-community 2.2 olea europaea subsp. africanacalpurnea aurea tall closed woodland. a dendrogram showing the habitat relationships of the various plant communities is shown in fig. 1. 1. olea capensis-cheilanthes viridis forest this community represents all the forests that occur in moist kloofs or sheltered valleys in the marakele national park. based on species composition, this vegetation can be regarded as afromontane forest (white 1978). according to the classification of fig. 1. a dendrogram showing the habitat relationships of the plant communities classified under the olea europaeasubsp. africana-diospyros whyteana major community. olea europaeadiospyros whyteana major community 1. dense forest pristine ib land type shallow lithosols > 50 % rock cover 2. dense thicket degraded mostly ad land type mostly yellow apedal soils 0-30 % rock cover 1.1 moist & cool perennial/seasonal streams southern & eastern slopes 1.1.1 steep ravines 1.1.2 sandstone slopes 2.1.1 large bush clumps 25 m diameter 2.1.2 small bush clumps 11 m diameter 2.1 flat plains ad land type deep yellow apedal soils termitaria 0 % rock cover 1.2 dry & warm seasonal stream northern & western slopes 2.2 southern slopes ad & ib land types shallow rocky soils 30 % rock cover van staden woud.qxd 2006/04/17 08:38 pm page 23 koedoe 49/1 (2006) 24 issn 0075-6458 south african forests by von maltitz (2003), the forests of the waterberg area represent marakele afromontane forests and northern highveld forests, both these types being classified under the northern afrotemperate group of forests. the olea capensischeilanthes viridis forest represents a northwestern island of afromontane (white 1978) or afrotemperate (von maltitz 2003) forests. within the study area this community is characterised by species group e (table 1), with the trees olea capensis and canthium gilfillanii and the ferns cheilanthus viridis and pellaea calomelanos as the diagnostic species. olea capensis, in particular, can be regarded as a typical forest species, though the other three species may occur widespread on rocky areas of the waterberg and bankenveld (bredenkamp & brown 2003; van staden & bredenkamp 2005). these forests can be subdivided into podocarpus latifolius-curtisia dentata forest, restricted to moist, sheltered valleys, and representing typical marakele afromontane forest (von maltitz 2003), and buxus macowanii-kirkia wilmsii dry low forest, a very specialized dry forest, representing a very rare and restricted plant community classified by von maltitz (2003) under northern highveld forests. 1.1 podocarpus latifolius-curtisia dentata forest these forests represent the typical moist short to tall forests of the waterberg area, tall forest is found at 1300 m–1730 m a.s.l. along perennial or seasonal streams and moist kloofs. this forest community is a representative of north-eastern mountain sourveld (acocks 1988), afromontane forest (white 1978) and typical marakele afromontane forest (mucina & van staden 2003), classified under the northern afrotemperate forest group (von maltitz 2003). this is a local forest type found only on the waterberg, marakele national park (van staden 2002) and some kloofs of magaliesberg (coetzee 1975). these northern afrotemperate forests are, according to von maltitz (2003), restricted to mountains and low ridges (strydpoortberg, waterberg, pilanesberg, witwatersrand, magaliesberg, suikerbosrand, sekhkukhuneland) interrupting the relatively flat northern highveld grassland or bushveld. the forests are found in kloofs on the northern and eastern flanks of the drakensberg. the diagnostic species of podocarpus latifolius-curtisia dentata forest are listed under species group c, with the trees podocarpus latifolius, curtisia dentata, rothmannia capensis, faurea saligna, rhus dentata, myrsine africana and ochna holstii, the liana secamone alpinii and the sedge cyperus albostriatus as diagnostic species. the tall-growing podocarpus latifolius is mostly the dominant species, locally reaching a cover of up to 70 %. 1.1.1 widdringtonia nodiflora-podocarpus latifolius short forest typical: relevé 130 habitat the widdringtonia nodiflora-podocarpus latifolius short forest is found at high altitudes, 1500 m–1730 m a.s.l. along perennial streams and deep, sheltered, moist kloofs, on gentle to moderate (6–22°) east and southfacing sandstone slopes within the ib land type. the soils are of the mispah form derived from sandstone of the sandriviersberg formation, and the soil is relatively shallow (< 500 mm) (land type survey staff 1988). although this community is not totally protected from fire, the woody vegetation and structure are seldom damaged due to the poorly developed grass layer and the high percentage rock cover that provides shelter against fire. large rocks cover more than 76 % of the soil surface. during the rainy season and for some time thereafter, water constantly seeps from the sandstone, creating an ideal moist habitat where this community is typically found. van staden woud.qxd 2006/04/17 08:38 pm page 24 issn 0075-6458 25 koedoe 49/1 (2006) floristics and structure the widdringtonia nodiflora-podocarpus latifolius short forest is differentiated by the following diagnostic plant species (species group a, table 1): the woody widdringtonia nodiflora, pittosporum viridiflorum, ilex mitis, halleria lucida, syzygium cordatum and cliffortia linearifolia; the shrubby clutia pulchella and phylica paniculata; the ferns blechnum giganteum, pteridium aquilinum, asplenium splendens and cheilanthus hirta; and the forb gerbera piloselloides. this community is represented by five relevés and an average of 19 species was recorded per sample plot. the tree layer is on average 8.4 m tall with an average canopy cover of 79 % (table 2). widdringtonia nodiflora is totally dominant, whereas podocarpus latifolius may locally be very prominent. apart from the diagnostic woody species, other prominent trees are curtisia dentata, canthium gilfillanii (species group c), and olea capensis (species group e). the latter species has a wider distribution within various forest communities in the park (table 1). the shrub layer is on average 1.4 m tall, with an average canopy cover of 22 % (table 2). prominent shrubs occurring in this community are myrsine africana and ochna holstii (species group c) (table 1). the woody liana secamone alpinii (species group c) is frequently found in the tree and shrub strata. the herbaceous layer has an average height of 0.5 m and an average canopy cover of 11.0 % (table 2). it is dominated by the diagnostic ferns blechnum giganteum, pteridium aquilinum and asplenium splendens (species group a) and the sedge cyperus albostriatus (species group c) as ground cover in the more open areas. the tree fern, cyathea dregei, which was not recorded in any sample plots, also occurs as isolated individuals in the forests in the study area. general the widdringtonia nodiflora-podocarpus latifolius short forest has many characteristic species in common with the ilex mitispittosporum viridiflorum forest community described by coetzee (1975) and the kloof forest communities described by westfall (1981). this community occupies the most mesic habitats of sheltered parts of kloofs on the eastern side of the waterberg mountain. the dominant woody species widdringtonia nodiflora and podocarpus latifolius are, however, absent from the forest communities described by coetzee (1975). communities 1.1.1 and 1.1.2 are floristically related through the mutual presence of the species of species group b, emphasised especially by podocarpus latifolius, curtisia dentata, rothmannia capensis and myrsine africana. these forests are also remotely related to buxus macowanii-kirkia wilmsii low forest (community 3), through species group e (table 1). 1.1.2 podocarpus latifolius-rothmannia capensis tall forest. typical: relevé 109 habitat the podocarpus latifolius-rothmannia capensis tall forest is found at 1300 m– 1660 m a.s.l. along perennial or seasonal streams and moist kloofs on sandstone slopes (4–30°), facing southeast, north, northeast and east. it is represented by five relevés and an average of 26 species was recorded per sample plot. this forest community is a representative of acocks’ (1988) north-eastern mountain sourveld, with the structure as a tall forest (edwards 1983) and it all occurs in the ib land type (land type survey staff 1988). during the rainy season and for some time thereafter water constantly seeps from the sandstone, creating an ideal moist habitat, characteristic for this community. the soils are of the mispah form derived from sandstone of the sandriviersberg formation, and the average soil depth is relatively shallow (< 500 mm) (land type survey staff 1988). large rocks cover more than 70 % of the soil surface. van staden woud.qxd 2006/04/17 08:38 pm page 25 koedoe 49/1 (2006) 26 issn 0075-6458 floristics and structure the podocarpus latifolius-rothmannia capensis tall forest is differentiated by the diagnostic following plant species (species group b, table 1): the woody combretum moggii, strychnos usambarensis, pterocelastrus echinatus, celtis africana, tetradenia brevispica, solanum giganteum, dovyalis zeyheri, englerophytum magalismontanum, pavetta gardeniifolia, diospyros lycioides and nuxia congesta; the herbaceous asparagus setosum; and the grass oplismenus hirtellus. the tree layer is on average 12 m tall with an average canopy cover of 79 %. other prominent trees are pittosporum viridiflorum (species group a), podocarpus latifolius and curtisia dentata (species group c), cussonia paniculata, ficus thonningii, olea europaea subsp. africana, euphorbia ingens, ficus sur (species group k), mimusops zeyheri (species group j), and olea capensis (species group e). the tree fern cyathea dregei, which was not recorded in the sample plots, also occurs in this community. the shrub layer is on average 1.7 m tall with an average canopy cover of 49 %. apart from the diagnostic species, other prominent shrubs occurring in this community are myrsine africana (species group c), canthium gilfillanii (species group e), and diospyros whyteana and grewia occidentalis (species group k). the woody liana secamone alpinii and cryptolepis transvaalensis are frequently found in the tree and shrub strata. the herbaceous layer has an average height of 0.5 m and an average canopy cover of 22 %. it is dominated by cyperus albostriatus and oplismenus hirtellus. general this community is closely related to the celtis africana-osyris lanceolata kloof forest described by westfall (1981). these two communities have many distinctive habitat features and many, distinctive species in common. the podocarpus latifolius-rothmannia capensis tall forest is floristically related to the widdringtonia nodiflorapodocarpus latifolius short forest, both being classified under the podocarpus latifolius-curtisia dentata forest. 1.2 buxus macowanii-kirkia wilmsii low forest typical: relevé 64 habitat the buxus macowanii-kirkia wilmsii low forest is found at 1110 m–1250 m a.s.l. the buxus macowanii-kirkia wilmsii low forest is restricted to hot, dry kloofs of north facing, east and northwest-facing slopes (2– 29°) of the study area (fig. 1) this low forest occupies a relatively drier and warmer habitat than the podocarpus latifolius-curtisia dentata forest. the structure of this forest community is a low forest (edwards 1983) and it occurs in the ib land type (land type survey staff 1988). the soils are of the mispah form derived from sandstone of the sandriviersberg formation, and the soil is relatively shallow (< 500 mm) (land type survey staff 1988). rocks cover more than 50 % of the soil surface. floristics and structure the buxus macowanii-kirkia wilmsii low forest is represented by four relevés and an average of 23 species was recorded per sample plot. it is differentiated by the following woody plant species (species group d): buxus macowanii, kirkia wilmsii, apodytes dimidiata, ochna pretoriensis, croton gratissimus, euphorbia turicalli, acacia ataxacantha and heteromorpha trifoliata. the tree layer is on average 7.8 m tall, with an average canopy cover of 35 %. other prominent trees are mimusops zeyheri, berchemia zeyheri, dombeya rotundifolia (species group j) and olea europaea subsp. africana, cussonia paniculata and euphorbia ingens (species group k). the shrub layer is on average 2.3 m tall, with an average canopy cover of 84 %. prominent shrubs occurring in this community are van staden woud.qxd 2006/04/17 08:38 pm page 26 issn 0075-6458 27 koedoe 49/1 (2006) ochna pretoriensis, croton gratissimus, euphorbia tirucallii and acacia ataxacantha (species group d), diospyros whyteana, and and heteropyxis natalensis (species group k). the herbaceous layer has an average height of 0.3 m and an average canopy cover of 7 %. the herbaceous layer is poorly represented and is dominated by the xerophytic ferns cheilanthes viridis and pellaea calomelanos (species group e). general westfall (1981) described an erythrina lysistemon-celtis africana kloof forest with buxus macowanii as a conspicuous woody species with more than 5 % mean canopy cover and occurring in more than 50 % of the relevés. this community is similar to the community found in the study area. these shrubs are often gregarious, forming pure stands (table 1). the distribution of the tree/shrub, buxus macowanii, is restricted to warm valleys and coastal dunes in the eastern cape, with the limpopo province population as an unexpected outlier (coatespalgrave 1983). this could indicate a former afromontane forest biogeographic connection via the waterberg and stydpoort mountains to the northeastern escarpment and also to the blouberg and western soutpansberg. the buxus macowanii-kirkia wilmsii dry low forest, a very specialised dry forest, representing a very rare and restricted plant community classified by von maltitz (2003) under northern highveld forests. according to von maltitz (2003), these latter forests are restricted to suikerbosrand (bredenkamp & theron 1978, 1980), witwatersrand (behr & bredenkamp 1988; ellery et al. 2001), magaliesberg (coetzee 1974, 1975), waterberg (westfall 1981; westfall et al. 1985; van staden 2002), sekhukhuneland (siebert 2001), and some other ridges occurring throughout the western central bushveld region (van vuuren 1961; van vuuren & van der schijff 1970; van der meulen 1979). these forest patches are imbedded within an ecotone between sourveld grasslands and subtropical savannas of the central bushveld. 2. pappea capensis-ziziphus mucronata thickets and bush clumps dombeya rotundifolia subsp. rotundifolia, berchemia zeyheri and ziziphus mucronata (species group j) show strong floristic affinities between these thickets and the buxus macowanii-kirkia wilmsii dry low forest. 2.1 rhus leptodictya-mimusops zeyheri termitarium thickets typical: relevé 18 habitat the rhus leptodictya-mimusops zeyheri termitarium thickets are found from 1160 m– 1300 m a.s.l. this widespread community occupies an even drier habitat than that of the buxus macowanii-kirkia wilmsii low forest, though the termitaria are moister than the surrounding savanna (van der meulen 1979). they are therefore classified with the forests. the bush clumps on termitaria are restricted to the ad land type on level and gentle slopes (land type survey staff 1988) (fig. 1). before proclamation of marakele national park, this community was subjected to heavy grazing by cattle and the field layer varies in height and cover depending on soil salinity and the amount of grazing (coetzee et al. 1981). the vegetation developed on huge termite mounds to form this specific plant community. the size of these termitaria determines the type of vegetation that occurs. the trees are usually very tall because of the depth and aeration and the fine texture and higher nutrient status of the soil (coetzee et al. 1976). this community is a representative of acocks’ (1988) sour bushveld, with the structure as a short closed woodland (edwards 1983) floristics the rhus leptodictya-mimusops zeyheri termitarium thickets are differentiated by the following diagnostic plant species (species group g, table 1): the woody rhus leptovan staden woud.qxd 2006/04/17 08:38 pm page 27 koedoe 49/1 (2006) 28 issn 0075-6458 dictya, euclea crispa, grewia flavescens, flueggia virosa, ficus thonningii and pavetta zeyheri; the liana asparagus transvaalensis; the grass eragrostis curvula; the herbaceous schistostephium heptalobum; and the weedy annual tagetes minuta. 2.1.1 rhus leptodictya-carissa bispinosa variation typical: relevé 18 habitat the rhus leptodictya-carissa bispinosa variation is found at 1160 m–1300 m a.s.l. this variation occurs on level ground or/and gentle slopes (1-160º). the soils are yellow, apedal and well drained and relatively deep (> 500 mm), belonging to one of the following forms: inanda, kranskop, magwa, hutton, griffin and/or clovelly (land type survey staff 1988) (fig. 1). the average diameter of the rhus leptodictya-carissa bispinosa variation bush clumps is 25 m. it is represented by seven relevés and an average of 35 species was recorded per sample plot (table 1). floristics and structure the rhus leptodictya-carissa bispinosa variation is differentiated by the following diagnostic plant species (species group f, table 1): the woody carissa bispinosa, gymnosporia buxifolia, vangueria infausta, clerodendrum glabrum, adenia glauca, dichrostachys cinerea, gardenia volkensii, psiadia punctulata, aloe marlothii, acacia karroo, maerua angolensis and pterocarpus rotundifolius; the forbs pavonia burchelli, sida dregei, kalanchoe rotundifolia, schkuhria pinnata, lippia javanica, sida cordiifolia, anthospermum hispidulum, ipomoea magnusiana, hemizygia pretoriae, solanum panduraeforme, scadoxus puniceus; and the grasses panicum maximum, aristida congesta, eragrostis rigidior, eragrostis lehmanniana and cynodon dactylon. many of these diagnostic species indicate previous over-utilisation and trampling. the tree layer is on average 7 m tall with an average canopy cover of 79 %. other prominent trees are pappea capensis and schotia brachypetala (species group i), olea capensis (species group e), mimusops zeyheri, dombeya rotundifolia subsp. rotundifolia, berchemia zeyheri and ziziphus mucronata (species group j), and olea europaea subsp. africana, ficus sur, cussonia paniculata and grewia occidentalis (species group k). the shrub layer is on average 1.4 m tall, with an average canopy cover of 46 %. prominent shrubs are grewia flavescens, flueggia virosa (species group k) and calpurnia aurea (species group h). the herbaceous layer has an average height of 0.4 m and an average canopy cover of 20 %. it is dominated by the grass panicum maximum and eragrostis rigidior, the forbs pavonia burchelli, schkuhria pinnata and lippia javanica (species group f). general coetzee et al. (1976) described similar communities from flat bottomlands with termitaria at nylsvley nature reserve, namely the pappea capensis-acacia tortilis variation which shows strong affinities with the rhus leptodictya-carissa bispinosa variation, with the most common plant species being carissa bispinosa and pappea capensis. 2.1.2 rhus leptodictya-berchemia zeyheri variation typical: relevé 16 habitat the rhus leptodictya-berchemia zeyheri variation is found at 1250 m–1360 m a.s.l. the soils are the same as for the rhus leptodictya-carissa bispinosa variation, but here the bush clumps are much smaller, with an average diameter of 11 m, whereas the average diameter of the bush clumps rhus leptodictya-carissa bispinosa variation is 25 m. floristics and structure this variation is represented by four relevés and an average of 35 species was recorded per sample plot. although it seems that no van staden woud.qxd 2006/04/17 08:38 pm page 28 issn 0075-6458 29 koedoe 49/1 (2006) diagnostic species occur in this variation, it can be distinguished from the rhus leptodictya-carissa bispinosa variation by the absence of species in the species group f and the presence of species in species group h (table 1), with maytenus undata, calpurnea aurea and calodendron capense being differential species. the tree layer is on average 6.5 m tall with an average canopy cover of 66 %. the shrub layer is on average 1.6 m tall with an average canopy cover of 46 %. the herbaceous layer has an average height of 0.4 m and an average canopy cover of 10 %. 2.2 olea europaea subsp. africana-calpurnea aurea tall closed woodland typical releve: relevé 97 habitat the olea europaea subsp. africanacalpurnea aurea tall closed woodland is found from 1180–1500 m a.s.l. in deep kloofs on southwest-facing and south-facing sandstone slopes (2–250º), usually along seasonal streams (table 1, fig. 1). this bush clump community is a representative of acocks’ (1988) sour bushveld, with the structure as a tall closed woodland (edwards 1983). it occurs on the ad land type and the ib land type (fig. 2.7) (land type survey staff 1988). the soils are of the mispah form derived from sandstone of the sandriviersberg formation (land type survey staff 1988) and large rocks cover more than 31 % of the soil surface (fig. 1). floristics and structure the olea europaea subsp. africanacalpurnea aurea tall closed woodland is characterised by the presence of species group h and absence of species group g. no diagnostic species group was identified, though the species of species group h are differential, it also indicates a floristic relationship with the rhus leptodictyaberchemia zeyheri variation (community 2.1.2). the differential species are the woody maytenus undata, calpurnia aurea, calodendron capense, gymnosporia polyacantha, vepris lanceolata and tricalysia lanceolata, the herbaceous asparagus virgatus and the grass setaria lindenbergiana. the tree layer is on average 10.5 m tall with an average canopy cover of 56 %. other prominent trees are celtis africana (species group b); rhus leptodictya, euclea crispa, ficus thonningii (species group g); pappea capensis, ziziphus mucronata, zanthoxylum capense, schotia brachypetala, euclea natalensis and acacia caffra (species group i); mimusops zeyheri and berchemia zeyheri (species group j); and olea europaea subsp. africana, ficus sur and euphorbia ingens (species group k). the shrub layer is on average 1.8 m tall with an average canopy cover of 41 %. cryptolepis transvaalensis is prominent. the herbaceous layer has an average height of 0.4 m and an average canopy cover of 21 %. the herbaceous layer is dominated by the grass setaria lindenbergiana and the forb hypoestes forskaolii (species group k). general the olea europaea subsp. africana-calpurnia aurea tall closed woodland has floristically strong affinities with the rhus leptodictyamimusops zeheri termitarium thickets (species group i) and especially the rhus leptodictya-berchemia zeyheri variation, indicated by species group h. the habitat difference between these two bush clump communities is that the olea europaea subsp. africana-calpurnia aurea tall closed woodland occurs in deep, sheltered kloofs on rocky substrates without termitaria, while the rhus leptodictya-berchemia zeyheri variation occurs on termitaria on the plains without rocks. discussion according to white (1978) the majority of the tree species in the afromontane forests are very widespread. amongst them are podocarpus latifolius, ilex mitis and halleria lucida that occur in the study area. these van staden woud.qxd 2006/04/17 08:38 pm page 29 koedoe 49/1 (2006) 30 issn 0075-6458 species could almost be used to define the afromontane region as a whole. not one species occurs throughout, but most species of the assemblage are represented on virtually every ‘island’ of afromontane forest. according to von maltitz (2003) the northern highveld forest type is a relict forest type in a progressed stage of ‘erosion’ from the original afromontane composition. they are considered remnants of the magaliesburg extension (white 1978)—an afromontane (forest) intrusion linking the forests of northern escarpment to those found in sheltered sites of the central bushveld and northern highveld regions. these forests are afrotemperate (afromontane) in character, however holocene climate changes have caused their confinement to fire-sheltered habitats and a concomitant decline in forest elements, but an increase in woodland elements (von maltitz 2003). this fragmentary afromontane type forest forms transitions to surrounding savanna and to the riverine forests with combretum erythrophyllum (van rooyen 1983; von maltitz 2003). there is also floristic similarity between the kloof forest communities and thickets found on termitaria. termitaria are known to increase water infiltration and percolation, as well as nutrient availability, compared to the surrounding savanna woodlands (van der meulen 1979). therefore, termitaria do support dense thickets with savanna species, however some forest species are also present. low diversity of the woody element places this type close to drakensberg montane forests and northern kwazulu-natal mistbelt forests (von maltitz 2003). the distribution of the shrub buxus macowanii is of biogeographic interest. this species is restricted to coastal forests of the eastern cape and forms an outlying population in the waterberg region (coates palgrave 1983). acknowledgements south african national parks is thanked for the opportunity to use the offical research project for a msc degree. references acocks, j.p.h. 1988. veld types of south africa, 3rd edn. memoirs of the botanical survey of south africa 57: 1–146. barkman, j.j., j. moravec & s. rauchert. 1964. kritische bemerkungen und vorschläge zur quantitativen vegetationanalyse. acta botanica neerlandia 13: 394–449. behr, c.m. & g.j. bredenkamp. 1988. a phytosociological classification of the witwatersrand national botanical garden. south african journal of botany 54: 525–533. bezuidenhout, h., h.c.biggs & g.j. bredenkamp. 1996. a process supported by the utility bbpc for analysing braun-blanquet data on a personal computer. koedoe 39(1): 107–112. bredenkamp, g.j. & l.r. brown. 2003. a reappraisal of acocks’ bankenveld: origin and diversity of vegetation types. south african journal of botany 69(1): 7–26. bredenkamp, g.j. & g.k. theron. 1978. a synecological account of the suikerbosrand nature reserve. 1. the phytosociology of the witwatersrand geological system. bothalia 12: 513–529. bredenkamp, g.j. & g.k. theron. 1980. a synecological account of the suikerbosrand nature reserve, part ii: the phytosociology of the ventersdorp geological system. bothalia 13(1&2): 199–216. coates palgrave, k. 1983. trees of southern africa. 2nd ed. cape town: struik. coetzee, b.j. 1974. a phytosociological classification of the vegetation of the jack scott nature reserve. bothalia 11: 329 – 347. coetzee, b.j. 1975. a phytosociological classification of the rustenburg nature reserve. bothalia 11: 561–580. coetzee, b.j., f. van der meulen, s. zwanziger, p. gonsalves & p.j.weisser. 1976. a phytosociological classification of the nylsvley nature reserve. bothalia 12: 137–160. coetzee, b.j., p. van wyk, w.p.d. gertenbach, a. hall-martin & s.c.j. joubert. 1981. 'n plantekologiese verkenning van die waterberggebied in die noord-transvaalse bosveld. koedoe 24:1–23. du preez, p.j., g.j.bredenkamp & h.j.t. venter. 1991. the syntaxonomy and synecology of the forests in the eastern orange free state, south africa. i. the podocarpetalia latifolii. south african journal of botany 57: 198–206. van staden woud.qxd 2006/04/17 08:38 pm page 30 edwards, d. 1983. a broad-scale structural classification of vegetation for practical purposes. bothalia 14 (3 & 4): 705–712. ellery, w.n., k. balkwill & k. ellery, r.a. reddy. 2001. conservation of the vegetation on the melville ridge, johannesburg. south african journal of botany 67: 261–273. geldenhuys, c.j. 1994. bergwind patches and the location of forest patches in the southern cape landscape south africa. journal of biogeography 21: 49–62. gertenbach, w.p.d. 1987. 'n ekologiese studie van die suidelikste mopanieveld in die nasionale krugerwildtuin. d.sc proefskrif, universiteit van pretoria, pretoria. hill, m.o. 1979. twinspan – a fortran program for arranging multivariate data in an ordered two way table by classification of individuals and attributes. ithaca, new york: cornell university. land type survey staff. 1988. land types of the maps 2426 thabazimbi, 2428 nylstroom. memoirs of the agricultural natural resoures of south africa no.10: 1–431. mucina, l. & p.j. van staden. 2003. marakele afromontane forests. pp. 99–104. in: von maltitz, g. (project leader). classification system for south african indigenous forests: an objective classification for the department of water affairs and forestry. pretoria: council for scientific and industrial research. (environmentek report env-p-c 2003-017). mueller-dombois, d. & h. ellenberg. 1974. aims and methods of vegetation ecology. new york: wiley. rutherford, m.c. & r.h. westfall. 1994. biomes of southern africa: an objective categorization. memoirs of the botanical survey of south africa 63: 1–94. siebert, s.j. 2001. vegetation of the ultramafic soils of the sekhukhuneland centre of endemism. ph.d. thesis, universty of pretoria, pretoria. van der meulen, f. 1979. plant sociology of the western transvaal bushveld. south africa: syntaxonomic and synecological study. vaduz: cramer. (dissertationes botanica 49.) van rooyen, n. 1983. die plantegroei van die roodeplaatdam-natuurreservaat ii. die plantgemeenskappe. south african journal of botany 2: 115–125. van staden, p. j. 2002. an ecological study of the plant communities of the marakele national park. m.sc. thesis, university of pretoria, pretoria. van staden p.j. & g.j. bredenkamp. 2005. major plant communities of the marakele national park. koedoe 48(2): 59–70. van vuuren, d.r.j. 1961. 'n ekologiese studie van 'n noordelike en suidelike kloof van die magaliesberge. m.sc. thesis, universty of pretoria, pretoria. van vuuren, d.r.j. & h.p. van der schijff. 1970. 'n vergelykende ekologiese studie van die plantegroei van 'n noordelike en suidelike kloof van die magaliesberg. tydskrif vir natuurwetenskappe 10: 16–75. von maltitz, g. (project leader). 2003. classification system for south african indigenous forests: an objective classification for the department of water affairs and forestry. pretoria: council for scientific and industrial research. (environmentek report env-p-c 2003-017). werger, m.j.a. 1974. on concepts and techniques applied in the zürich-montpellier method of vegetation survey. bothalia 11(3): 309–323. westfall, r.h. 1981. the plant ecology of the farm groothoek, thabazimbi district. m.sc. thesis, university of pretoria, pretoria. westfall, r.h., n. van rooyen & g.k. theron. 1985. the plant ecology of the farm groothoek, thabazimbi district. 2. classification. bothalia 15: 655–688. white, f. 1978. the afromontane region. pp. 465–513. in: werger, m.j.a. (ed.). biogeography and ecology of southern africa. the hague: junk. issn 0075-6458 31 koedoe 49/1 (2006) van staden woud.qxd 2006/04/17 08:38 pm page 31 filelist convert a pdf file! koedoe 19: 175-176 (1976) two new bird records for the tsitsikama national parks g . a. robinson tsitsikama coastal national park p. o. stormsrivier 6)08 a comprehensive check-list of the birds that possibly could occur in the vicinity of the parks was published by skead and liversidge (1967). they refer to 210 bird species but only record 205; three species were duplicated in part 1 (forest habitats) and part 2 (sea habitats). it would appear that two species were omitted from part 1. however, one of the species recently recorded should be included in part 1 while the other belongs to part 2. on 1973.07.03 a lesser gallinule porphyrio alieni, (roberts number 209) was observed outside the boathouse at storms river mouth. this bird was captured , fed to a healthy state and eventually released at nature's valley. prior to the sighting of this bird the weather had been foul with strong south-westerly winds and rough seas and rain. the bird was in a poor state of health weak and undernourished . as this species selects marshy reeded areas to hive in and no such area exists in the immediate vicinity of storms river mouth, it was surmised that the bird had been blown in from one of these habitats west of storms river. nature's valley, nearly 40 km to the western boundary seemed to be a likely area that would attract gallinules. identification was made by comparing the dimensions of the wing, tarsus, culmen, shield width and tail length to those of the purple, american and lesser gallinules as presented by mclachlan and liversidge (1970). these are listed in table 1. more recently on 1975.07.15 another lesser gallinule was collected at storms river mouth. this bird was found shortly after the same adverse weather conditions had prevailed as the first, i.e. stormy southwesterly winds accompanied by heavy rain. the bird was a juvenile and in a weak and hungry condition, and it was therefore assumed that the bird probably could not have flown. because the storms river was in flood during the period of both sightings both birds could have come down the river which is marshy and has reeds upstream. on the 1973 .07. 10 a sea-bird in poor condition, which later died, was found on the beach at nature's valley. professor j. m. winterbottom (pen . comm.), identified the bird as a blue petrel halobaena caerlea (roberts number 20). its measurements are given in table 1. these two additional records brings the total of birds species for the parks to 207. 175 table 1 comparative measurements of gallinules and blue petrels gallinules blue petrel measurements storms river purple' american lesser .... nature's blue peu-e!· specimen purple' valley specimen wing 153mm 225-248 mm 162-181 mm 144-160mm 217mm 205-221 mm tarsus 44mm 70--92 mm 58--jj2 mm 45-56 mm 32mm 39-33 mm cu lmen 37 mm 56--71 mm 38-48 mm 35-40 mm 25mm 25-28 mm shield width ilmm 18-24 mm tail length 65mm 82-101 mm 64-mm 66--73 mm 104mm 77-91 mm r.: measurements from roberts references mclachlan, g. r. and r. liversidge. 1970. roberts birds of south africa. cape town: trustees of the s.a. bird book fund. skead, c.j. and r. liversidge. 1967. birds of the tsitsikama forest and coastal national park, 1966. koedoe 10:43-62. 176 page 1 page 2 abstract introduction the evolving association between people, non-native trees and sense of place viewshed and sense of place in a protected area context synthesis of insights acknowledgements references appendix 1 about the author(s) jaco barendse scientific services, south african national parks, south africa sustainability research unit, nelson mandela metropolitan university, south africa dirk roux scientific services, south african national parks, south africa sustainability research unit, nelson mandela metropolitan university, south africa waldo erfmann biodiversity and social projects, south african national parks, south africa johan baard scientific services, south african national parks, south africa tineke kraaij school of natural resource management, nelson mandela metropolitan university, south africa cara nieuwoudt sharples environmental services, cape town, south africa citation barendse, j., roux, d., erfmann, w., baard, j., kraaij, t. & nieuwoudt, c., 2016, ‘viewshed and sense of place as conservation features: a case study and research agenda for south africa’s national parks’, koedoe 58(1), a1357. http://dx.doi.org/10.4102/koedoe.v58i1.1357 original research viewshed and sense of place as conservation features: a case study and research agenda for south africa’s national parks jaco barendse, dirk roux, waldo erfmann, johan baard, tineke kraaij, cara nieuwoudt received: 04 sep. 2015; accepted: 29 mar. 2016; published: 05 aug. 2016 copyright: © 2016. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract sense of place (sop) refers to the meanings and values that people attach to places. the concept can be used to frame how people engage or form a connection with the natural environment. at a sensory level, sop is influenced by people’s visual experiences, which in turn can be linked to the concept of viewsheds. viewsheds can be transformed, either abruptly (e.g. by infrastructure development such as wind turbines) or more gradually (e.g. by non-native trees invading a landscape). in this study, we focus on the garden route national park to explore the potential importance of viewsheds as a conservation feature, specifically in the context of non-native (especially invasive) tree species. using mixed information sources, we explore the potential role of invasive trees on experiences of visitors to this protected area and speculate on how viewsheds may shape sop associations and how such associations may inform protected area management. our investigation shows that people’s experiences regarding natural and modified viewsheds are varied and intricate. both sop and viewsheds have the potential to inform conservation action, and these concepts should form an integral part of objective hierarchies and management plans for national parks. however, while legislation and park management plans make provision for the use of these concepts, associated research in south africa is virtually non-existent. we conclude by proposing a conceptual model and research agenda to promote the use of viewsheds and sop in the management of national parks in south africa. conservation implications: viewshed and sense of place can be used as boundary concepts to (1) facilitate interdisciplinary research between social and natural scientists, (2) help understand the connectedness and feedbacks between people and nature and (3) promote communication between science, management and stakeholders regarding desired conditions of landscapes in and around parks. introduction a growing number of scientists believe that earth has transitioned into a new geological epoch – the anthropocene – characterised by a single, dominant species (homo sapiens) that is affecting the planet’s life support system at an unprecedented scale, including changes to landscapes and ecosystems, biological distributions, climate and atmospheric chemistry (steffen, crutzen & mcneill 2007; zalasiewicz et al. 2010). the proclamation and effective management of representative networks of protected areas (pas) is seen as critical to buffering society against adverse changes to the biosphere attributable to this human-dominated era (watson et al. 2014). in this context, expected benefits from pas include conserving biodiversity, safeguarding ecosystems and the services they provide, mitigating climate change and promoting social–ecological resilience, with associated economic and social benefits, at regional scales. there is increasing recognition of the importance of less tangible or quantifiable benefits that people derive from nature and pas. such benefits are, for example, referred to as ‘nature’s gifts’ in the intergovernmental platform on biodiversity and ecosystem services conceptual framework (díaz et al. 2015). these benefits are often grouped under the collective of ‘cultural ecosystem services’, including experiential, spiritual, educational and recreational interactions with nature that contribute to human well-being (millennium ecosystem assessment 2005). importantly, an appreciation of such benefits is often dependent on a pro-environmental identity or ability to engage or form a connection with the natural environment (hinds & sparks 2008), for example, through ‘meaningful nature experiences’ (zylstra et al. 2014). how people connect with nature is invariably a function of their value systems (see chan, satterfield & goldstein 2012; raymond et al. 2009), which are context-specific and evolve dynamically over time. for example, while one group may value a particular landscape for its tangible materials (such as harvestable fruits and medicinal plants), another may value the same landscape for intangible benefits (such as relaxation and therapy) derived from its tranquil and scenic features. for the purpose of this study, we will focus on intangible benefits that people derive through experiential interactions with nature. in this context, connection between humans and nature relate strongly to aesthetics (plieninger et al. 2015) and therefore link closely with concepts such as ‘viewsheds’ and ‘landscapes’. perceptions of ‘beautiful scenery’ may be a predictor for environmental connection. likewise, the sensitivity of ‘the public’ to scenery or ‘how the landscape looks’ is an important driver of support for conservation actions. this is borne out by spatial metrics studies (palmer 2004) and evidence that positive nature experiences (such as hikes) may predispose people to financially support conservation efforts (zaradic, pergams & kareiva 2009). ironically, although the creation of pas has often led to the separation of humans and nature (west, igoe & brockington 2006), these remnants of wilder, more natural or intact ecosystems and landscapes potentially offer greater opportunity to experience connection with nature and benefits such as psychological rejuvenation (see ulrich et al. 1991). higher ecological integrity should result in higher ‘visual landscape quality’, and together with the notion of ‘sense of place’ (sop) and the related concept of ‘place attachment’ (the environmental psychologist’s equivalent of the geographer’s sop) (farnum, hall & kruger 2005) is subsumed in the landscape-quality construct (daniel 2001). although sometimes regarded as vague (shamai 1991) and elusive (williams & stewart 1998), the concept of sop has been applied widely to describe the relationship between people and physical environment. it is generally used for framing the meanings and values that people attach to places (larson, de freitas & hicks 2013; williams & stewart 1998) and may incorporate experiences of dependence, attachment, identity and satisfaction (jorgensen & stedman 2001; stedman 2003). such meanings and values are typically rich and varied (williams & stewart 1998), commonly based on a mix of cultural histories and natural features in a landscape (larson et al. 2013), and develop as a result of biological, individual and sociocultural processes that take place while interacting with the physical environment (hausmann et al. 2016). many feel that sop resides primarily in human experiences, interpretations and value endowment, rather than being intrinsic to the physical setting itself – ‘space becomes place when we endow it with value’ (tuan 1977; but see stedman 2003). at a sensory level, what people do (e.g. fish from the bank of a river), feel (e.g. grass under their feet or warmth of the sun), hear (e.g. the sound of birds or the wind in the trees) and see (e.g. a seascape or forest) will contribute to their experiences in relation to a place. such experiences are likely to change over time (e.g. different seasons) and space (e.g. vantage points) and to be mediated by memory of previous such experiences. while the natural sciences have found ways to measure, for example, changes in soundscapes (pijanowski et al. 2011) and viewsheds (camp, sinton & knight 1997), neither of these concepts has been incorporated into the predominantly social construct of sop. apart from the social variable, there may be many different ways in which sop can be altered or lost, through changes in physical appearance within a landscape, and thus its aesthetics. let us consider viewsheds in ‘natural’ or ‘wilderness’ areas, defined in the us wilderness act (public law 88–577, 16 u.s. c. 1131–1136, 88th congress, second session, september 3, 1964) as: … undeveloped … land retaining its primeval character and influence, without permanent improvements or human habitation, which is protected and managed so as to preserve its natural conditions and which … generally appears to have been affected primarily by the forces of nature, with the imprint of man’s work substantially unnoticeable; and … has outstanding opportunities for solitude or a primitive and unconfined type of recreation. (n.p.) it stands to reason that such viewsheds (and associated place-value) may be transformed, or even destroyed, by evidence of human presence or activities. viewshed transformation can intuitively be linked to structural developments, for example, housing on a lake shoreline (stedman 2003), presence of roads (selva et al. 2011) or power infrastructure such as overhead pylons and wind turbines (gee 2010). however, there are other, less explicit, human-mediated changes. one such ‘slow’ transformer of viewsheds may be non-native plants that have become invasive, especially large, woody, tree species (figure 1) – widely referred to as weeds, invaders or invasive alien plants (iaps). while we acknowledge the variety of terms and connotations associated with describing iaps (e.g. richardson et al. 2000; schlaepfer, sax & olden 2011), we choose to use the more neutral term ‘non-native’ except when explicitly referring to declared invasive alien species or ‘weeds’ as defined by the conservation of agricultural resources act (cara, no. 43 of 1983, and amendments). the impacts of iaps on biodiversity and ecosystems, and resultant ecosystem service loss, such as water quality (e.g. chamier et al. 2012; van wilgen et al. 2008), are widely known and are often the primary determinants for allocation of funding and human resources to manage invasions (marais, van wilgen & stevens 2004). less well understood is the impact of iaps on cultural ecosystem services, such as mediated through sop experiences (but see le maitre et al. 2011), and this aspect is generally not considered when prioritising areas for iap clearing and restoration. figure 1: non-native trees and invasive alien plants are examples of ‘slow transformers’ (compared to other human developments) of landscapes and viewsheds in and around the garden route national park (also see figure 3), as shown by photographs taken in different years near the iconic ‘kaaimansgat’ – where the kaaimans and swart rivers enter the indian ocean and located within the western buffer/viewshed protection zone of the grnp. the view to the north from the dolphin point lookout in 1929 (a) and 2015 (b); the view to the south from next to the n2/kaaimans pass, pre-1910 (c), 1927 (d) and 2015 (e) – note cellular phone tower ‘disguised’ as pine tree shown by arrow and insert; and the ‘map of africa’ viewpoint, pre-1910 (f) and in 2015 (g). in this study, we use an interdisciplinary narrative approach, including the use of various pieces of ‘evidence’ and ‘exhibits’, to explore the potential importance of viewsheds as conservation features, specifically in the context of non-native (especially invasive) tree species. we restrict our attention to the western cape, the province regarded as the epicentre for development of awareness about iaps, mainly because of regional public reverence for the native vegetation of the globally recognised cape floristic region (bennett 2014). furthermore, we focus on the garden route national park (grnp) along the southern cape coast (garden route), a region widely known for its scenic beauty and an example of a pa embedded or integrated within a greater social–ecological landscape. using mixed information sources, including media, unpublished studies, scientific literature and management documents, we explore the potential role of invasive plants (especially trees) on experiences of visitors to this pa and speculate on how viewsheds may shape sop associations and the interplay between such associations and pa management. to this end, we present a conceptual model relating the concepts of viewshed and sop to the high-level objectives in a management plan for a national park in south africa. lastly, we propose a research agenda to inform the future incorporation of viewsheds and sop in park management decisions. the evolving association between people, non-native trees and sense of place when considering terms such as ‘naturalness’, or whether non-native trees have a place in natural viewsheds, it is important to acknowledge that associated perceptions develop over time. societal perceptions influence, and are influenced by, the reigning utilitarian values (e.g. forestry or dune stabilisation) and socio-economic circumstances of the period (baard & kraaij 2014). therefore, sentiments about a specific ‘immigrant species’ may change as a society does, or according to whether it is associated with a negative impact or trait, such as being invasive or harmful (coates 2007). perceptions may further depend on how informed individuals are, their level of knowledge of biodiversity, or on more personal values of aesthetics, or notions of scenic beauty (dhami & deng 2010; garcía-llorente et al. 2008) (also see box 1 and figure 2). a case in point is the black wattle (acacia mearnsii), arguably one of the most problematic iap species in coastal areas of the western cape, originally introduced to support the tanning industry (carruthers et al. 2011). especially for a botanist or invasion biologist today, it may be unsettling to travel through the garden route where black wattle, together with other acacia and pinus species (see photographs in cowling et al. 2009), dominate the viewsheds of coastal, riparian, mountain and production landscapes (henderson 1998). that the wattle is now considered unwanted in this region was not always the case, as shown by a letter written to the george and knysna herald on 13 september 1893, titled ‘wattle growing at knysna’: last sunday the writer took a stroll ‘over the hills’, but by no means so ‘far away’ to the nursery at concordia, and would recommend to those of his fellow townsmen who are blessed with the aptitude for enjoying natures beauties to lose no time in hieing tither ere [hastening there before] the many varieties of acacia lose their magnificent bloom. the wattles are just now simply one blaze of bright yellow, and are really a sight worth beholding. (n.p.) figure 2: four scenes, three from around the garden route national park: views from (a) the ‘seven passes road’ near the woodville ‘big tree’ looking towards the upper touw river catchment; and (b) donaghy’s hill (42011’47.04”s, 145056’01.97”e) in the franklin-gordon wild rivers national park, tasmania; (c) the area north of karatara in the garden route national park, looking towards the mountains traversed by the outeniqua hiking trail; (d) the non-invasive australian red flowering ‘gum’ trees corymbia ficifolia in full bloom at bergplaas forestry station is a common and much-loved sight in the garden route; along with related alien eucalypts (rejmánek & richardson 2011; van staden 2015) many consider these part of the region’s natural-cultural heritage. box 1: appreciating natural viewsheds and sense of place (see figure 2). the writer then goes on to suggest that establishing wattle plantations in the area would not only provide economic benefits on otherwise ‘idle ground’ or ‘sour waste lands’, but that anyone who has seen them in full bloom (as described) would be supportive of his suggestions, presumably on aesthetic grounds. indeed, wattle growing did become an important economic mainstay in the region well into the 1960s and was strongly supported by the local authorities, as shown by a piece titled ‘the municipal wattle plantations’ in 03 june 1914 edition of the herald: the public of george will learn with pleasure that at the last meeting of the municipal council it was unanimously resolved to prepare another 6 morgen of land for the planting of wattle. this is a move in the right direction and if steadily persevered in, this city will find itself within a few years the owner of a very valuable asset … (n.p.) the above example shows that the lower value attributed to local landscapes and vegetation and the higher value to the non-native species played an important role in promoting their deliberate spread. contrast this with the present-day situation, where a. mearnsii is now one of the top invaders in the garden route (baard & kraaij 2014) of especially riparian habitats (holmes et al. 2005). country-wide, it has cost an estimated r62 million from 1997 to 2006 to control invasions by acacia spp. alone (marais & wannenburgh 2008) through national initiatives such as working for water. the ‘duality’ in perspective on whether invasive species are ‘good’ or ‘bad’ has a strong cultural dimension (tassin & kull 2015) and ties closely to the leading discourse at any given time (bennett 2014; carruthers et al. 2011), but it can also vary between different industry sectors, for example, forestry and conservation. in king’s 1951 paper ‘tree planting in south africa’, he stated: there is a small section of the population who wage a wordy warfare against the planting of exotic trees. this element with fanatical zeal, presents only one side of the picture. let us look at the other side. criticism is often levelled against wattles on the cape flats and pines on the mountains of the cape peninsula. in order to put the matter in proper perspective we must go back to the time when much of the cape flats was a barren waste of drift sands and the only trees on the mountains were contained in small patches … [on the cape flats there are] … only shrubs called blombos (metalasia) and waxberry (myrica) both of which are much less valuable than the australian wattles … [that] yield excellent firewood. can anyone be so foolish as to imagine that without wattles a population starved for firewood would not have stripped the mountains of woody vegetation? the claim can safely be made that indirectly the wattles have saved the mountain flora from extermination. (p. 13) in a similar vein, king (1951) stated the following about pines: despite their high intrinsic value, pines have been described as weeds, mainly on the grounds that they tend to spread. this is only true of pinus pinaster, which can readily be kept in check. the allegation that pine plantations are ousting indigenous vegetation is not entirely true, but, even if it were, it cannot be taken seriously, because the plantations occupy less than 3,000 of the 120,000 acres in the [cape] peninsula. (p. 14) while these remarks may appear naïve given (expert) knowledge about the current causes, extent and impacts of pine tree invasions (mcconnachie et al. 2015), they do represent views that persist to the present, even in scientific circles. for example, a recent ‘pine plantation vs. fynbos’ debate (de ronde 2012) publicly pitched ecologists (stellenbosch university 2012) against forestry scientists (du toit 2012), showing that some perceptions remain entrenched (de wit, crookes & van wilgen 2001). the article that sparked much of the discussion (van wilgen & richardson 2012) did recognise that exotic conifers may offer aesthetical and recreational benefits to people, especially in peri-urban areas. such opposing perceptions about non-native species are also pervasive among members of the public, often reflecting incomplete understanding about ecological processes or biodiversity conservation, as shown by 62% of park visitors to the addo elephant national park who did not consider the potential presence of introduced fauna to be in conflict with conservation objectives (boshoff et al. 2008). being uninformed can lead to the confusion of unrelated issues, for example, the felling of plantations perceived as synonymous with deforestation of natural forests, or where ‘saving’ a tree – any tree – is seen as positive (van wilgen 2012). perceptions are highly context-specific, for example, species that have been naturalised for a long time are not necessarily perceived as ‘alien’ even by traditional communities (shackleton et al. 2007); or, despite knowing a species’ alien status, a high utility value may reduce the support for its outright eradication (de neergaard et al. 2005). this may further vary according to socio-economic variables, where people of higher economic status or better education may rate non-consumptive values of indigenous plants (e.g. aesthetics) higher than poorer or less educated people (le maitre et al. 1997). the resultant conflict (between informed and uninformed parties) over non-native species (dickie et al. 2014) may reduce the potential support for clearing iaps in pas, in particular those located close to urban areas, or ‘embedded’ in cities (van wilgen 2012). this does not mean that the potential contribution of pas in preserving natural viewsheds or landscapes on the basis of aesthetics has escaped recognition. possibly the earliest local proponent of this cause was wicht (1943) who – using the cluster pine pinus pinaster (=maritima) as an example – suggested (almost prophetically, see cowling et al. 2009; kraaij, cowling & van wilgen 2011) that exotic plants would, over time, dominate everywhere except in nature reserves and that ‘to botanists and all other lovers of nature the thought that such a change is likely to come is very distressing’ (p. 34). he further suggested that ‘species that are spreading into natural vegetation … [are] undesirable from an aesthetic or scientific view’ (p. 43) and (quoting the forest and veld conservation act of 1941) that nature reserves should be set aside for the ‘preservation of natural scenery, forests, flora or fauna thereon’ (p. 45) (wicht 1943). the above raises three important questions, which we discuss in the ensuing sections, using the grnp and surrounds as example: is the preservation of natural viewsheds and associated sop included or provided for in current national legislation relating to pas? do park management plans recognise the potential impact of iaps on viewshed and sop as conservation features? do visitors to pas value ‘natural’ viewsheds and perceive the presence of iaps as a threat to sop in such areas? viewshed and sense of place in a protected area context national legislation the national environmental management: protected areas act (57/2003) (nem:pa – norms and standards for the management of pas in south africa) stipulates (chapter 1, p. 3) under guiding principle (a) that pas should ‘protect ecologically viable areas representative of south africa’s biological diversity and its natural landscapes and seascapes’; and under (k) ‘contribute to human, social, cultural, spiritual … development’. in chapter 2 (p. 4), one of the indicators of relative importance is that an area ‘protects a representative sample or iconic feature of south africa’s land/seascapes’. while provision is made for a zoning plan that determines different activities and conservation objectives within a pa (p. 15), there is no mention of sop or viewsheds, and the section referring to visitor’s experiences mainly considers visitor facilities (p. 24). in an amendment to nem:pa (strategy on buffer zones for national parks, notice 106 in government gazette no. 35020, 08 february 2012) in the context of managing activities in buffer zones surrounding national parks, a viewshed is defined as ‘an area of land, or other environmental element that is visible to the human eye from a fixed vantage point’. in chapter 3 (p. 12), two of the stated objectives of the policy are to (1) ‘protect, enhance and restore the unique and memorable character – the sop – that underpins the image of the national parks and their approaches’; and (2) ‘protect and enhance the wilderness experience of park users’. to this end, the policy calls for ‘viewshed protection’ in buffer zone areas that are visible from the park, especially ‘from wilderness areas, or … night lights which negatively effect [sic] the ambience of the park’ (p. 13). forestry is specifically listed as a development that ‘may have a negative impact or effect on a national park’ (p. 13) and the control of iaps in buffer zones (p. 19) is mentioned in the context of local community involvement and ecosystem rehabilitation; however, the policy falls short of making the link between the potential impact of iaps or non-native trees (other than in plantations) on viewsheds, or sop. another area of governance that holds implications for activities in park buffer zones and takes into consideration viewshed and sop is land-use planning and environmental management. at a national level, the national environmental management act and the environmental conservation act, which guide environmental impact assessments, include visual impact considerations as a component. further, the national heritage resources act (act no. 25 of 1999) provides protection for listed or proclaimed heritage resources and sites, such as urban conservation areas, nature reserves, and recognised scenic routes. western cape provincial legislation requires the preparation of a spatial development framework (sdf) and an integrated development plan for each sub-region or municipality. these documents aim to guide land-use to be compatible with cultural and scenic landscapes and could include reference to open-space and scenic resources, together with management guidelines for the area covered by these plans, for example, the knysna sdf (http://www.knysna.gov.za/wp-content/uploads/2012/12/knysna-sdf-nov-2008-core.pdf). under these management guidelines, protection is provided not only to existing natural assets such as existing and proposed pas but also to transformed, so-called ‘productive green areas’, that include existing agricultural and commercial forestry areas, which: have historically been, and should remain important sources of productive economic activity in the municipal area, as well as being contributors to the sense of place. (n.p.) this role of sop and visual amenity of non-native vegetation thus finds legal application (which may be in conflict with the application of nem:pa) in urban expansion developments. in a recent case at the coastal town of plettenberg bay on the garden route, a part of a residential development was not granted environmental authorisation in order to retain a stand of mature, non-native eucalyptus trees alongside an indigenous forest on the same property. park management plans a somewhat more detailed consideration of viewshed is found at the park management plan level. for example, the garden route national park management plan (grnpmp; sanparks 2012) – and in fact all other park management plans – make generic provision for viewshed protection areas and defines a viewshed protection zone as ‘an area where any developments should be screened to prevent excessive impact on the aesthetic appeal of the park’. the grnpmp recognises five different zones (table 1, figure 2) that stipulate limits of acceptable change in terms of aesthetics and recreational activities, including consideration of facilities and infrastructure development and visitor numbers. although terms such as ‘wild appearance and character’, ‘natural appearance’ and ‘wilderness characteristic’ are used to distinguish between the different types of zones, there is no specific mention of iaps and their potential influence on the aesthetic appeal in these zones, or visitor’s experiences. we are unaware of any initiatives in the national parks through which viewsheds or the sop experiences of visitors in relation to viewsheds are being monitored. this might be a function of nem:pa providing only limited support for monitoring and reporting against progress in implementing plans, with legal obligations to monitor largely confined to the impact of revenue-generating activities. table 1: zonation and limitations of acceptable change to aesthetics and recreational activities. public perceptions towards invasive alien plants in protected areas information on the perception of park visitors regarding the presence of iaps in viewsheds, or more generally on sop in south african pas, appears to be non-existent. here, we present results from an exploratory and an opportunistic survey. the first was conducted during 2013 in the knysna section of the grnp at three sites: spitskop viewpoint and the fisantehoek and sinclair huts on the outeniqua hiking trail (figure 3). the second opportunistic survey was conducted along the course of the 5-day tsitsikamma hiking trail (in january 2014), which crosses mountain fynbos and indigenous forest areas managed by south african national parks (tsitsikamma section of the grnp) and pine plantations (cape pine). figure 3: the location of the garden route national park wilderness and knysna sections, main geographical features and places, and the viewshed protection zones defined in table 1, relative density of alien invasive plants within the grnp and selected points of interest mentioned in the text or illustrated in other figures. in the first survey, a composite panoramic photograph was taken of the available viewshed at each of the three sites, and each photo was delineated into numbered sections based on the visible extent of iap coverage (see examples in figure 4a), but excluding barren areas such as roads. for each of the identified numbered sections, this was equated to actual iap densities (see table 1-a1) derived from aerial vegetation cover and transformation surveys (vlok, euston-brown & wolf 2008) and further classified according to the height class of the predominant vegetation (≤ 2 m or > 2 m) and state of transformation by iaps, relative to the desired natural state (on a five-point scale, from fully transformed to completely natural). overnight hikers and day visitors were approached between 08 may and 07 july 2013 and shown a photo of the viewshed (with numbered sections) at the given site. they were then asked to score each section on how it influenced their viewshed experience, where 1 = a reduced experience, 2 = no effect and 3 = an enhanced experience. they were asked to motivate each score: every time iaps were mentioned, it was recorded against the specific section. the survey was concluded with the question, ‘why did you specifically choose to visit this site’? if the answer included words or phrases alluding to the aesthetic appeal of the site (e.g. ‘naturalness’, ‘wildness’, ‘prettiness’), it was recorded (see box 2 and figure 5). figure 4: examples of photographs shown to park visitors to determine viewshed preferences and assess awareness about invasive alien plants: views at (a) fisantehoek hut and (b) sinclair hut showing 4 and 10 delineated sections, respectively, to illustrate the methodology used; and composite panoramic photo of the view at (c) spitskop viewpoint (11 sections, not shown). figure 5: influence on visitor’s experience attributable to (a) vegetation height and (b) state of transformation of the vegetation in perceivable viewshed sections on photographs, combined for three sites. also indicated is the number of times invasive alien plants were mentioned per vegetation height class or transformation state. the number of respondents who specifically mentioned (c) invasive alien plants per viewshed section or (d) aesthetics per survey site. box 2: results from visitor surveys in the knysna section of the garden route national park. during the second opportunistic survey, hikers encountered were asked three questions. this was after hiking for at least 2 days through a landscape with a high prevalence of non-native trees, including dense stands of pines (see figure 6) and wattles in the fynbos sections (see box 3). figure 6: invasive pine trees growing along the tsitsikamma hiking trail may, or may not, impact on the viewshed and sense of place experiences of hikers (box 3). box 3: responses of nine hikers on the tsitsikamma hiking trail. synthesis of insights our varied assemblage of evidence suggests that sop, although provided for in legislative spheres and by national park and environmental management, remains a poorly developed concept in south africa. where and when considered, it rarely relates to a holistic appreciation of viewsheds, landscapes or biodiversity, including non-native trees. specifically for the grnp, the extant zonation scheme (table 1) recognises changes to aesthetics due to human activities, albeit at a very coarse scale, but it does not consider non-native trees or iaps. the same applies to the demarcated viewshed protection zone (figure 3) where, ironically, high aip densities may occur in areas zoned as ‘remote’ or along the outeniqua hiking trial (figure 2c), particularly in fynbos vegetation. in buffer zones, our examples show that over the past century, non-native trees have become established features of iconic view sites (figure 1) and main tourist routes (figure 2b). this is indicative that wattles and pines continue to form part of a publicly acceptable viewshed, firstly, due to the historic role of these non-native trees in the development of the region, and secondly, due to a lack of a collective mental model of what a representative garden route viewshed should look like. thus, while le maitre et al. (2011) argue that invasive australian acacias can negatively affect both tourist experience and sop by reducing ‘landscape diversity’ and degrading recreational areas, our exploratory surveys suggest that this relationship is not so straightforward. even where park visitors were aware of non-natives, such knowledge did not necessarily translate into negative experiences (as suggested by results from our survey – figure 5a), and it seems that mature (> 2 m) non-native trees may even contribute to positive scenic or aesthetic experiences. likewise, while some hikers displayed fairly good understanding of invasive pines and alluded to negative experiences, others were apparently oblivious or tended towards absolute biocentricity – simply enjoying any trees (box 3). therefore, while scientists may be able to distinguish between ecosystem service benefits offered by initial introductions of non-natives, and ecosystem service losses due to subsequent invasions, this may not be intuitive for non-scientists, especially when such species also provide tangible benefits (e.g. de neergaard et al. 2005). this may hinder public support or understanding of clearing and restoration efforts. the same applies at park management level, where managers may be aware of the negative impact of iaps on biodiversity and provision of ecosystem services, but may not value the preservation of aesthetics (e.g. in california – funk et al. 2013) or consider the potential impact on tourism (forsyth et al. 2012) as a reason to combat iaps. this is similar for attitudes towards non-native ornamental species that are found in rest camps in kruger national park (foxcroft, richardson & wilson 2008), where it required education and increased awareness of staff to gain support for the removal of such plants. similarly, tourists in pilanesberg who were aware of the invasive prickly pear (opuntia stricta) indicated a willingness to contribute financially to its control (nikodinoska et al. 2014). in an ‘open-access’ scenario such as the grnp where visitors do not necessarily have to pass through a gate to experience the park, the demarcation between pas, surrounding buffer zones and the rest of the landscape is less easily observable. given that non-native trees are entrenched as part of the garden route’s cultural heritage, it will require a particularly pragmatic (and creative) approach to use feedbacks from visitor sop experiences to inform park management. the role of shifting baselines in sense of place given how attachment to place and sop develop over time and is closely related to individual baselines, it is worth considering how the concepts of naturalness or reference states are formed. retaining historical character of a region may serve as motivation for aip eradication but over time people become accustomed to non-natives (schlaepfer et al. 2011). there may be differences between experiences of local and foreign visitors, making ‘novel’ ecosystems more acceptable to some due to a lack of historic perspective, or personal familiarity with similar viewsheds elsewhere. internationally, there is some indication that the way in which people appreciate natural areas is evolving. a recent case study in the river piedra floodplain (spain), has shown that over the past 50 years, there has been a positive shift towards appreciation of social and cultural ecosystem services, including aesthetics, inspiration and sop (see figure 5 in felipe-lucia, comín & escalera-reyes 2015). similarly, land managers of the nature conservancy in the usa consider impacts of non-natives on aesthetics at least equally important to degradation of other (provision) ecosystem services (kuebbing & simberloff 2015). while restoration and aip clearing can have positive effects on ecosystem service provision and tourism in certain biomes or vegetation types (e.g. fynbos – currie, milton & steenkamp 2009), the ‘human element’ also needs due consideration. an example from finnish national parks showed that the natural characteristics such as scenery and biotype diversity are significant determinants of park visitation, along with factors such as recreational opportunities, for example, availability of trails (neuvonen et al. 2010). while it is easy to understand the danger of an ‘absolute ecocentric’ attitude (sharp et al. 2011), which would oppose the clearing of any tree at all costs, a single-minded, negative focus on non-native species without considering other factors may be counter-productive. in the case of the garden route, the contribution of non-native trees and plantation forestry to the historic heritage (e.g. ‘flower gums’, figure 2d) deserves consideration. there are legal mechanisms to guide some of these trade-off situations, such as the ‘champion tree’ project by the department of agriculture, forestry and fisheries (in terms of section 12 of the national forests act of 1998) which recognises individual specimens or clumps of non-native trees of historic value or exceptional size. also, the south african national heritage resources act (act no. 25 of 1999) mandates protection of ‘heritage objects’ including natural landscapes of cultural significance; even non-native trees associated with such landscapes, for example, historic arboretums. this emphasises the need for a deliberate process of consultation informed by history, public participation and science to agree on acceptable viewsheds, both native and non-native, that contribute to sop. conceptual framework for incorporating viewsheds and sense of place in the management plans of national parks in south africa, the nem:pa makes it obligatory for authorities responsible for pas to develop a management plan for each such area, to submit this plan for the approval of the minister responsible for the environment and to manage the pa in accordance with the approved plan. in the case of sanparks, an adaptive planning approach is followed to formulate a hierarchy of objectives that serve as a basis for developing management plans for the various national parks (see biggs & rogers 2003; foxcroft & mcgeoch 2011). adaptive planning takes place in consultation with relevant organs of state, local communities and other affected parties. in figure 7 we propose a conceptual framework for linking the concepts of viewshed and sop to the typical high-level management objectives of a national park. according to our framework, viewsheds (and other sensory features such as soundscapes) can be described in both biophysical and social or cultural terms. sop experiences can be viewed as an emergent property of the interaction between people and the environment. such experiences are mediated by factors such as community history, identity and value systems and can be facilitated or hindered by park management actions. regarding the latter, there is an obvious interplay between management action and visitor experiences: while conservation initiatives have the potential to build on existing, or to create new, sop associations (larson et al. 2013), being aware of sop experiences can be an effective driver of conservation actions (ardoin 2014). figure 7: schematic showing how viewshed and sense of place relate to a typical set of high-level objectives that serve as a basis for developing park management plans in south african national parks. viewshed and sense of place are cross-cutting concepts likely to be influenced via multiple objectives and in turn impact on the responsible tourism objective. several arrows in figure 7 indicate bi-directional influence. such two-way feedbacks should be important considerations in designing monitoring and management interventions for incorporating viewsheds and sop into park management plans. concluding thoughts and proposing a research agenda in this article, we have explored the potential impact of iaps on viewsheds, within a broader context of sop as experienced by visitors to grnp. the novelty of our study also results in limitations, in that insights derived from focussing on iaps, the grnp and park visitors are not necessarily applicable to other landscape transformers, pas or sectors of society. however, our study also highlights some pertinent points with generic relevance to pas in south africa. firstly, the mixed information sources considered in this study suggest that viewshed and sop are important conservation features from both conceptual and legal perspectives. as such, these concepts need to inform conservation action and therefore should be incorporated into park management plans. secondly, viewshed and sop should be considered through both natural and social lenses to facilitate discussions of the ‘desired future conditions’ of landscapes under conservation from both ecological and social perspectives (williams & stewart 1998). to this end, viewshed and sop can potentially serve as ‘boundary concepts’ to promote interdisciplinary learning between social and natural scientists as well as communication between science, management and stakeholders (chapin iii & knapp 2015). thirdly, the links and feedbacks between conservation features such as viewshed and sop, disturbances such as non-native and invasive plants and various park management objectives are multiple and intricate. these relationships may straddle human history (shifting baselines) and thus environmental and social contexts. place-specific monitoring will be required to meaningfully incorporate these concepts into park management practice. fourthly, the current lack of formal research in south africa on viewsheds and sop, especially relating to national parks and their buffer areas, represents a considerable void in our understanding of the relationship between park management and visitor experiences. some studies on visitors’ motivations to visit south african pas have identified activities such as photography (e.g. saayman, saayman & ferreira 2009), implying recognition of scenic values. however, there is a need to explicitly evaluate the role and value of sop and natural viewsheds in pas as well as the potential implications that may result from various threats. threats are likely to be regionand context-specific (e.g. iaps in grnp, hydraulic fracturing activities and infrastructure in the karoo parks and wind farms in coastal areas), further emphasising the breadth of research opportunities. how do the relationships depicted in figure 7 play out in the real-world setting of a specific park? we conclude by proposing research questions that could serve as a basis from which to develop a more comprehensive research programme for improved appreciation of viewsheds and sop as conservation constructs. firstly, we consider questions related to viewsheds: considering a broad definition of biodiversity, encompassing genetic, species and ecosystem (including habitat) diversity, could viewshed serve as a surrogate feature (similar to an umbrella species) for conservation? should south africa be concerned with conserving a representative sample of natural viewsheds (e.g. per bioregion or biome) and to what degree can or should national parks contribute to such a purpose? what are ‘representative’ or ‘iconic’ viewsheds for specific pas in terms of historic naturalness and biogeography? how should specific sites for representative and iconic viewsheds be identified? should such viewsheds be restored where they no longer exist, and which methods should be used to reconstruct acceptable baselines (e.g. soliciting park visitors to submit historic photographs from of chosen sites)? what are the main threats to, and modifiers of, natural or cultural viewsheds and how do these affect sop experiences of visitors? what is the role of buffer zones in viewshed conservation? should thresholds of potential concern (tpcs – biggs et al. 2011) be developed for viewsheds and how could such tpcs inform monitoring (e.g. through fixed-point photography) for compliance with set objectives? questions related to sop are: how should sop experiences (based on feedback from stakeholders and visitors) be considered in the design, establishment and management of pas? can we characterise sop experiences for each national park and surrounding areas? how do activities, such as guided hikes, animal tracking (e.g. cheetah tracking in mountain zebra national park), trail running and mountain biking (in the grnp) influence sop experiences of participating and other visitors to these pas? how do sop experiences differ across age groups, cultures and nationalities of visitors, as well as local versus non-local residents, or day versus overnight visitors? are the dynamics of sop experiences different in open-access pas to those in fenced-off pas with distinct boundaries? what is the relationship between individual and collective experiences in developing attachment to place? implementing a research agenda as suggested here could significantly contribute to people-centred conservation while at the same time promoting south african national parks’ vision of ‘connecting to society’ (http://www.sanparks.co.za/about/connecting_to_society/). acknowledgements j.b. was supported by a postdoctoral research fellowship from nelson mandela metropolitan university research capacity building and sanparks. we would like to thank lynne thompson at the george research library for providing historic photographs. we are grateful for helpful colleague reviews from wessel vermeulen, lizette van der vyver, marna herbst, wendy annecke, liandi slabbert and peter bradshaw. we appreciate the constructive comments provided by three anonymous reviewers. the relevant components of this study obtained ethics clearance from nelson mandela metropolitan university (reference h2013-sci-nrm-008). competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions j.b. (south african national parks and nelson mandela metropolitan university) and d.r. developed the project concept, j.b. (south african national parks and nelson mandela metropolitan university), d.r. and w.e. were responsible for experimental design, data collection and analyses. t.k. and c.n. made conceptual contributions and literature suggestions. j.b. (south african national parks) drew the map in figure 3. j.b. (south african national parks and nelson mandela metropolitan university) and d.r. wrote the manuscript with contributions by w.e., t.k., j.b. (south african national parks) and c.n. references ardoin, n., 2014, ‘exploring sense of place and environmental behavior at an ecoregional scale in three sites’, human ecology 42, 425–441. http://dx.doi.org/10.1007/s10745-014-9652-x baard, j.a. & kraaij, t., 2014, ‘alien flora of the garden route national park, south africa’, south african journal of botany 94, 51–63. http://dx.doi.org/10.1016/j.sajb.2014.05.010 bennett, b.m., 2014, ‘model invasions and the development of national concerns over invasive introduced trees: insights from south african history’, biological invasions 16, 499–512. http://dx.doi.org/10.1007/s10530-013-0601-1 biggs, h., ferreira, s., freitag-ronaldson, s. & grant-biggs, r., 2011, ‘taking stock after a decade: does the ‘thresholds of potential concern’ concept need a socio-ecological revamp?’, koedoe 53(2), 9. http://dx.doi.org/10.4102/koedoe.v53i2.1002 biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j. du toit, k. rogers & h. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59–80, island press, washington, dc. boshoff, a.f., landman, m., kerley, g.i.h. & bradfield, m., 2008, ‘visitors’ views on alien animal species in national parks: a case study from south africa’, south african journal of science 104, 326–328. camp, r.j., sinton, d.t. & knight, r.l., 1997, ‘viewsheds: a complementary management approach to buffer zones’, wildlife society bulletin 25, 612–615. carruthers, j., robin, l., hattingh, j.p., kull, c.a., rangan, h. & van wilgen, b.w., 2011, ‘a native at home and abroad: the history, politics, ethics and aesthetics of acacias’, diversity and distributions 17, 810–821. http://dx.doi.org/10.1111/j.1472-4642.2011.00779.x chamier, j., schachtschneider, k., le maitre, d., ashton, p. & van wilgen, b., 2012, ‘impacts of invasive alien plants on water quality, with particular emphasis on south africa’, water sa 38, 345–356. http://dx.doi.org/10.4314/wsa.v38i2.19 chan, k.m., satterfield, t. & goldstein, j., 2012, ‘rethinking ecosystem services to better address and navigate cultural values’, ecological economics 74, 8–18. http://dx.doi.org/10.1016/j.ecolecon.2011.11.011 chapin, iii, f.s. & knapp, c.n., 2015, ‘sense of place: a process for identifying and negotiating potentially contested visions of sustainability’, environmental science & policy 53, 38–46. http://dx.doi.org/10.1016/j.envsci.2015.04.012 coates, p., 2007, american perceptions of immigrant and invasive species: strangers on the land, university of california press, berkeley, ca. cowling, r., van wilgen, b., kraaij, t. & britton, j., 2009, ‘how no-man’s-land is now everyone’s problem. the renowned cape flora is everywhere in retreat as runaway pine invasions transform the outeniqua and tsitsikamma mountains’, veld & flora 95(3), 147–149. currie, b., milton, s.j. & steenkamp, j.c., 2009, ‘cost-benefit analysis of alien vegetation clearing for water yield and tourism in a mountain catchment in the western cape of south africa’, ecological economics 68, 2574–2579. http://dx.doi.org/10.1016/j.ecolecon.2009.04.007 daniel, t.c., 2001, ‘whither scenic beauty? visual landscape quality assessment in the 21st century’, landscape and urban planning 54, 267–281. http://dx.doi.org/10.1016/s0169-2046(01)00141-4 de neergaard, a., saarnak, c., hill, t., khanyile, m., berzosa, a.m. & birch-thomsen, t., 2005, ‘australian wattle species in the drakensberg region of south africa – an invasive alien or a natural resource?’, agricultural systems 85, 216–233. http://dx.doi.org/10.1016/j.agsy.2005.06.009 de ronde, n., 2012, pine vs fynbos, viewed n.d., from http://saforestryonline.co.za/articles/environment/pine_vs_fynbos/ de wit, m.p., crookes, d.j. & van wilgen, b.w., 2001, ‘conflicts of interest in environmental management: estimating the costs and benefits of a tree invasion’, biological invasions 3, 167–178. http://dx.doi.org/10.1023/a:1014563702261 dhami, i. & deng, j., 2010, ‘modeling the scenic beauty of the highland scenic highway, west virginia’, in clifton e. watts, jr. & cherie leblanc fisher (eds.), proceedings of the 2009 northeastern recreation research symposium, pp. 66, gen. tech. rep. nrs, u.s. department of agriculture, forest service, northern research station. díaz, s., demissew, s., carabias, j., joly, c., lonsdale, m., ash, n. et al., 2015, ‘the ipbes conceptual framework — connecting nature and people’, current opinion in environmental sustainability 14, 1–16. http://dx.doi.org/10.1016/j.cosust.2014.11.002 dickie, i.a., bennett, b.m., burrows, l.e., nuñez, m.a., peltzer, d.a., porté, a. et al., 2014, ‘conflicting values: ecosystem services and invasive tree management’, biological invasions 16, 705–719. http://dx.doi.org/10.1007/s10530-013-0609-6 du toit, b., 2012, “no” to biased and pseudo scientific reporting, says forest scientist, viewed n.d., from http://blogs.sun.ac.za/news/files/2012/05/forestry-in-fynbos-reply-bdt.pdf farnum, j., hall, t. & kruger, l.e., 2005, sense of place in natural resource recreation and tourism: an evaluation and assessment of research findings, department of agriculture, forest service, pacific northwest research station, portland, or. felipe-lucia, m., comín, f. & escalera-reyes, j., 2015, ‘a framework for the social valuation of ecosystem services’, ambio 44, 308–318. http://dx.doi.org/10.1007/s13280-014-0555-2 foxcroft, l. & mcgeoch, m., 2011, ‘implementing invasive species management in an adaptive management framework’, koedoe 35(2), art. #1006, 11 pages. http://dx.doi.org/10.4102/koedoe.v53i2.1006 foxcroft, l., richardson, d. & wilson, j.u., 2008, ‘ornamental plants as invasive aliens: problems and solutions in kruger national park, south africa’, environmental management 41, 32–51. http://dx.doi.org/10.1007/s00267-007-9027-9 forsyth, g.g., le maitre, d.c., o’farrell, p.j. & van wilgen, b.w., 2012, ‘the prioritisation of invasive alien plant control projects using a multi-criteria decision model informed by stakeholder input and spatial data’, journal of environmental management 103, 51–57. funk, j.l., matzek, v., bernhardt, m. & johnson, d., 2013, ‘broadening the case for invasive species management to include impacts on ecosystem services’, bioscience 64, 58–63. http://dx.doi.org/10.1093/biosci/bit004 garcía-llorente, m., martín-lópez, b., gonzález, j.a., alcorlo, p. & montes, c., 2008, ‘social perceptions of the impacts and benefits of invasive alien species: implications for management’, biological conservation 141, 2969–2983. http://dx.doi.org/10.1016/j.biocon.2008.09.003 gee, k., 2010, ‘offshore wind power development as affected by seascape values on the german north sea coast’, land use policy 27, 185–194. http://dx.doi.org/10.1016/j.landusepol.2009.05.003 hausmann, a., slotow, r., burns, j.k. & di mini, e., 2016, ‘the ecosystemn service of sense of place: benefits for human well-being and biodiversity conservation, environmental conservation’ 43(2), 117–127. http://dx.doi.org/10.1017/s0376892915000314 henderson, l., 1998, ‘invasive alien woody plants of the southern and southwestern cape region, south africa’, bothalia 28(1), 91–112. http://dx.doi.org/10.4102/abc.v28i1.624 hinds, j. & sparks, p., 2008, ‘engaging with the natural environment: the role of affective connection and identity’, journal of environmental psychology 28, 109–120. http://dx.doi.org/10.1016/j.jenvp.2007.11.001 holmes, p., richardson, d., esler, k., witkowski, e. & fourie, s., 2005, ‘a decision-making framework for restoring riparian zones degraded by invasive alien plants in south africa: review article’, south african journal of science 101, 553–564. jorgensen, b.s. & stedman, r.c., 2001, ‘sense of place as an attitude: lakeshore owners attitudes toward their properties’, journal of environmental psychology 21, 233–248. http://dx.doi.org/10.1006/jevp.2001.0226 king, n.l., 1951, ‘tree-planting in south africa’, journal of the south african forestry association 21(1), 1–102. http://dx.doi.org/10.1080/03759873.1951.9631082 kraaij, t., cowling, r.m. & van wilgen, b.w., 2011, ‘past approaches and future challenges to the management of fire and invasive alien plants in the new garden route national park’, south african journal of science 107(9/10), art. #633, 11 pages. http://dx.doi.org/10.4102/sajs.v107i9/10.633 kuebbing, s.e. & simberloff, d., 2015, ‘missing the bandwagon: nonnative species impacts still concern managers’, neobiota 25, 73–86. http://dx.doi.org/10.1007/s40362-014-0021-3 larson, s., de freitas, d.m. & hicks, c.c., 2013, ‘sense of place as a determinant of people’s attitudes towards the environment: implications for natural resources management and planning in the great barrier reef, australia’, journal of environmental management 117, 226–234. http://dx.doi.org/10.1016/j.jenvman.2012.11.035 le maitre, d., gelderblom, c., maphasa, l., sarel, y., van den belt, m. & manuel, t., 1997, ‘communicating the value of fynbos: results of a survey of stakeholders’, ecological economics 22, 105–121. http://dx.doi.org/10.1016/s0921-8009(97)00572-7 le maitre, d.c., gaertner, m., marchante, e., ens, e.-j., holmes, p.m., pauchard, a. et al., 2011, ‘impacts of invasive australian acacias: implications for management and restoration’, diversity and distributions 17, 1015–1029. http://dx.doi.org/10.1111/j.1472-4642.2011.00816.x marais, c., van wilgen, b.w. & stevens, d., 2004, ‘the clearing of invasive alien plants in south africa: a preliminary assessment of costs and progress’, south african journal of science 100, 97–103. marais, c. & wannenburgh, a.m., 2008, ‘restoration of water resources (natural capital) through the clearing of invasive alien plants from riparian areas in south africa — costs and water benefits’, south african journal of botany 74, 526–537. http://dx.doi.org/10.1016/j.sajb.2008.01.175 mcconnachie, m.m., van wilgen, b.w., richardson, d.m., ferraro, p.j. & forsyth, a.t., 2015, ‘estimating the effect of plantations on pine invasions in protected areas: a case study from south africa’, journal of applied ecology 52, 110–118. http://dx.doi.org/10.1111/1365-2664.12366 millennium ecosystem assessment (mea), 2005, ecosystems and human well-being: synthesis, island press, washington, dc. neuvonen, m., pouta, e., puustinen, j. & sievänen, t., 2010, ‘visits to national parks: effects of park characteristics and spatial demand’, journal for nature conservation 18, 224–229. http://dx.doi.org/10.1016/j.jnc.2009.10.003 nikodinoska, n., foxcroft, l.c., rouget, m., paletto, a. & notaro, s., 2014, ‘tourists’ perceptions and willingness to pay for the control of opuntia stricta invasion in protected areas: a case study from south africa’, koedoe 56(1), art. #1214, 8 pages. http://dx.doi.org/10.4102/koedoe.v56i1.1214 palmer, j.f., 2004, ‘using spatial metrics to predict scenic perception in a changing landscape: dennis, massachusetts’, landscape and urban planning 69, 201–218. http://dx.doi.org/10.1016/j.landurbplan.2003.08.010 pijanowski, b.c., villanueva-rivera, l.j., dumyahn, s.l., farina, a., krause, b.l., napoletano, b.m. et al., 2011, ‘soundscape ecology: the science of sound in the landscape’, bioscience 61, 203–216. http://dx.doi.org/10.1525/bio.2011.61.3.6 plieninger, t., bieling, c., fagerholm, n., byg, a., hartel, t., hurley, p. et al., 2015, ‘the role of cultural ecosystem services in landscape management and planning’, current opinion in environmental sustainability 14, 28–33. http://dx.doi.org/10.1016/j.cosust.2015.02.006 raymond, c.m., bryan, b.a., macdonald, d.h., cast, a., strathearn, s., grandgirard, a. et al., 2009, ‘mapping community values for natural capital and ecosystem services’, ecological economics 68(5), 1301–1315. http://dx.doi.org/10.1016/j.ecolecon.2008.12.006 rejmánek, m. & richardson, d.m., 2011, ‘eucalypts’, in d. simberloff & m. rejmánek (eds.), encyclopedia of biological invasions, pp. 203– 209, university of california, berkeley, ca. richardson, d.m., pyšek, p., rejmánek, m., barbour, m.g., panetta, f.d. & west, c.j., 2000, ‘naturalization and invasion of alien plants: concepts and definitions’, diversity and distributions 6, 93–107. http://dx.doi.org/10.1046/j.1472-4642.2000.00083.x saayman, m., saayman, a. & ferreira, m., 2009, ‘the socio-economic impact of the karoo national park’, koedoe 51(1), art. #158, 10 pages. sanparks, 2012, garden route national park: park management plan, south african national parks, pretoria. viewed n.d., from https://www.sanparks.org/assets/docs/conservation/park_man/grnp-approved-plan.pdf schlaepfer, m.a., sax, d.f. & olden, j.d., 2011, ‘the potential conservation value of non-native species’, conservation biology 25, 428–437. http://dx.doi.org/10.1111/j.1523-1739.2010.01646.x selva, n., kreft, s., kati, v., schluck, m., jonsson, b.-g., mihok, b. et al., 2011, ‘roadless and low-traffic areas as conservation targets in europe’, environmental management 48, 865–877. http://dx.doi.org/10.1007/s00267-011-9751-z shackleton, c.m., mcgarry, d., fourie, s., gambiza, j., shackleton, s.e. & fabricius, c., 2007, ‘assessing the effects of invasive alien species on rural livelihoods: case examples and a framework from south africa’, human ecology 35(1), 113–127. shamai, s., 1991, ‘sense of place: an empirical measurement’, geoforum 22, 347–358. http://dx.doi.org/10.1016/0016-7185(91)90017-k sharp, r.l., larson, l.r. & green, g.t., 2011, ‘factors influencing public preferences for invasive alien species management’, biological conservation 144(8), 2097–2104. stedman, r.c., 2003, ‘is it really just a social construction?: the contribution of the physical environment to sense of place’, society & natural resources 16, 671–685. http://dx.doi.org/10.1080/08941920309189 steffen, w., crutzen, p.j. & mcneill, j.r., 2007, ‘the anthropocene: are humans now overwhelming the great forces of nature’, ambio 36, 614–621. http://dx.doi.org/10.1579/0044-7447(2007)36[614:taahno]2.0.co;2 stellenbosch university, 2012, ‘no’ to forestry in fynbos areas, says invasive species specialists, viewed n.d., from http://blogs.sun.ac.za/news/2012/05/17/no-to-forestry-in-fynbos-areas-says-invasive-species-specialists/ tassin, j. & kull, c.a., 2015, ‘facing the broader dimensions of biological invasions’, land use policy 42, 165–169. http://dx.doi.org/10.1016/j.landusepol.2014.07.014 tuan, y.-f., 1977, space and place: the perspective of experience, university of minnesota press, minneapolis. ulrich, r.s., simonds, r.f., lositio, b.d., fiorito, e., miles, m.a. & zelson, m., 1991, ‘stress recovery during exposure to natural and urban environments’, journal of environmental psychology 11, 201–230. http://dx.doi.org/10.1016/s0272-4944(05)80184-7 van staden, f., 2015, ‘a sticky question: do gum trees really have to go?’, south magazine 15, viewed n.d., from http://southmagazine.co.za/archives/2015/a-sticky-question/ van wilgen, b.w., 2012, ‘evidence, perceptions, and trade-offs associated with invasive alien plant control in the table mountain national park, south africa’, ecology and society 17(2), 23. http://dx.doi.org/10.5751/es-04590-170223 van wilgen, b.w., reyers, b., le maitre, d.c., richardson, d.m. & schonegevel, l., 2008, ‘a biome-scale assessment of the impact of invasive alien plants on ecosystem services in south africa’, journal of environmental management 89, 336–349. http://dx.doi.org/10.1016/j.jenvman.2007.06.015 van wilgen, b.w. & richardson, d.m., 2012, ‘three centuries of managing introduced conifers in south africa: benefits, impacts, changing perceptions and conflict resolution’, journal of environmental management 106, 56–68. http://dx.doi.org/10.1016/j.jenvman.2012.03.052 vlok, j.h.j., euston-brown, d.i.w. & wolf, t., 2008, a vegetation map for the garden route initiative, unpublished 1:50 000 maps and report supported by cape fsp task team. watson, j.e.m., dudley, n., segan, d.b. & hockings, m., 2014, ‘the performance and potential of protected areas’, nature 515, 67–73. http://dx.doi.org/10.1038/nature13947 west, p., igoe, j. & brockington, d., 2006, ‘parks and people: the social impacts of protected areas’, annual review of anthropology 35, 251–277. http://dx.doi.org/10.1146/annurev.anthro.35.081705.123308 wicht, c.l., 1943, ‘preservation of the vegetation of the south western cape’, transactions of the royal society of south africa 30, 7–18. http://dx.doi.org/10.1080/00359194309519824 williams, d.r. & stewart, s.i., 1998, ‘sense of place: an elusive concept that is finding a home in ecosystem management’, journal of forestry 96, 18–23. zalasiewicz, j., williams, m., steffen, w. & crutzen, p., 2010, ‘the new world of the anthropocene’, environmental science & technology 44, 2228–2231. http://dx.doi.org/10.1021/es903118j zaradic, p.a., pergams, o.r.w. & kareiva, p., 2009, ‘the impact of nature experience on willingness to support conservation’, plos one 4, e7367. http://dx.doi.org/10.1371/journal.pone.0007367 zylstra, m., knight, a., esler, k. & le grange, l.l., 2014, ‘connectedness as a core conservation concern: an interdisciplinary review of theory and a call for practice’, springer science reviews 2, 119–143. http://dx.doi.org/10.1007/s40362-014-0021-3 appendix 1 table 1-a1: the density classes (percentage ground cover) and description used in the analysis of results of viewshed zone preference by visitors as surveyed at spitskop viewpoint, sinclair and fisantehoek huts. article information authors: emma f. gray1,2 william j. bond2 affiliations: 1south african environmental observation network fynbos node, cape town, south africa2department of botany, university of cape town, south africa correspondence to: emma gray postal address: private bag x7, claremont 7735, africa dates: received: 05 aug. 2012 accepted: 29 apr. 2013 published: 02 aug. 2013 how to cite this article: gray, e.f. & bond, w.j., 2013, ‘will woody plant encroachment impact the visitor experience and economy of conservation areas?’, koedoe 55(1), art. #1106, 9 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1106 copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. will woody plant encroachment impact the visitor experience and economy of conservation areas? in this original research... open access • abstract • introduction • methods    • study area    • visibility of animals in different habitats    • visitor surveys    • vegetation change and desirability    • visitor demography and reasons for visiting    • effect of encroachment on visitor satisfaction • results    • visibility of animals in different habitats    • visitor perceptions    • demographic profiles of visitors    • reasons for visiting hluhluwe–imfolozi park    • visitor response to changing vegetation • discussion • conclusion • acknowledgements    • competing interests    • authors’ contributions • references • footnote abstract top ↑ woody plant encroachment into savannas is a globally prevalent phenomenon and impacts ecosystem goods and services such as biodiversity, carbon storage, nutrient cycling, grazing and hydrology. the direct ecological and economic consequences for rangelands have been fairly well studied, but, to our knowledge, the economic impact on conservation efforts has not been investigated. african savannas are important as conservation areas because they support large numbers of the world’s remaining megafauna. this study used visitor surveys and long-term mammal distribution data to investigate how an increase in tree density might affect the visibility of animals in a conservation area, which could reduce the satisfaction of visitors to the area. we found that apparent herd sizes and density of animals were much reduced in woody areas, suggesting that visibility is negatively impacted. visitor surveys determined that a large fraction (almost half) of potential future visitors to the park may be lost if animals became more difficult to see and that the majority of these would be the higher-spending visitors. responses differed depending on the origin of visitors, with international visitors being more interested in seeing animals, whilst local visitors were more content with just being away from the city. the results suggest that woody plant encroachment may have significant impacts on visitor numbers to savanna conservation areas, whilst animal numbers and densities may also be significantly impacted.conservation implications: the results pointed to potentially significant economic consequences for conservation efforts as visitors become less satisfied with their experience. perceptions of visitors are important for management decisions as park fees contribute significantly to conservation efforts. this could ultimately result in a reduced capacity for african conservation areas to conserve their biodiversity effectively. the results suggest that management may need to re-evaluate their approach to controlling woody plant encroachment. introduction top ↑ savannas and grasslands are economically important as rangelands and agricultural lands (costanza et al. 1997; sankaran, ratnam & hanan 2004). in africa in particular, savannas have an intrinsic sentimental value because they are home to most of the world’s last remaining megaherbivores (animals > 1000 kg). this translates into an economic importance because they attract visitors from all over the world, which contributes significantly to the gross domestic product of many african countries (akama & kieti 2003; akama, maingi & camargo 2011; di minin et al. 2012). the experiences of these visitors are an important source of information to guide park managers and planners in appropriate measures to achieve both visitor satisfaction and conservation goals (boshoff et al. 2007; obua & harding 1996).woody plant encroachment has been extensive in both farmed and conserved savannas all over the world over the last century (archer, schimel & holland 1995; asner et al. 2003; britz & ward 2007; fensham, fairfax & archer 2005; skarpe 1990; van auken 2000; wigley, bond & hoffman 2010). the drivers are strongly debated, but recent literature recognises the importance of both local drivers, such as changes in land use (fire and herbivore stocking rates), and global drivers, such as increases in atmospheric carbon dioxide concentrations (bond 2008; bond & midgley 2012; bowman, murphy & banfai 2011; wigley et al. 2010). besides environmental repercussions such as loss of biodiversity (bond & parr 2010; parr, gray & bond 2012; ratajczak, nippert & collins 2012) and altered ecosystem goods and services such as water supply and carbon sequestration (coetsee et al. 2013), woody thickening also has economic consequences on commercial rangelands. stocking rates are likely to drop as trees increasingly restrict animal access and forage is reduced as grass cover declines. the consequence is a reduction in the overall carrying capacity of the land (moleele et al. 2002; roques, o’connor & watkinson 2001). in namibia, cattle numbers have been reduced to 36% of what they were in 1959, and woody plant encroachment is estimated to cause a loss of more than n$700 million per annum (de klerk 2004). what is often overlooked is that in savannas under conservation there are likely to be direct consequences for wild animals, which may translate into unforeseeable indirect economic consequences for conservation areas. savannas support a diversity of habitats for mammals and birds (sirami et al. 2009). different animals show diverse preferences for certain habitats owing to direct (sinclair, mduma & brashares 2003) and indirect (brown 1988, 1992, 1999; brown, laundré & gurung 1999; laundré, hernández & altendorf 2001; ripple & beschta 2004) effects of predation and the availability of grazing. this makes for a diversity of wildlife viewing experiences for visitors to conservation areas. this spatial heterogeneity is crucial for maintaining diverse species assemblages and may help to buffer against changes in resource availability due to a changing climate (wang et al. 2006) by compensating for temporal variability in resource availability (fryxell et al. 2005). woody plant encroachment will reduce spatial heterogeneity in vegetation structure. this change in habitat structure is likely to affect animal distributions, herd sizes and densities, with the direction of change depending on the size and feeding habits of the species concerned (du toit & owen-smith 1989). however, whether actual animal numbers are reduced may be of little importance to visitors. an increase in tree cover will reduce visibility and, as more woody plants encroach, it is likely that animals will become more difficult to see (marshall, lovett & white 2008). an inability to see animals may lead to a reduced wildlife experience, which could lead to a decrease in visitor numbers in conservation areas. regular visitors to a conservation area are likely to notice reduced animal visibility due to woody plant encroachment, which could translate to a reduction in their desire to return to the park. here we report how woody plant density affects wildlife viewing in a savanna park and how visitors are likely to respond to woody plant encroachment. woody plant encroachment is widespread in african savannas and is well documented in our study area, which was far more open in the early and mid-20th century but has become heavily encroached in the last few decades (watson & macdonald 1983; wigley et al. 2010). we addressed the following questions: • does denser vegetation result in fewer visible animals and therefore poorer game viewing opportunites? • do visitors go to parks primarily to see wildlife? • if visitors’ ability to view wildlife is reduced, will they be less likely to visit parks and, if so, could this lead to reduced visitor numbers in parks and subsequent loss of revenue? methods top ↑ study area this study was conducted in the hluhluwe–imfolozi park (hip) complex, south africa (28°00’–28°26’s; 31°43’–32°09’e), which consists of the hluhluwe game reserve (225 km²) in the north, the imfolozi game reserve (447 km²) in the south and a corridor (227 km²) joining the two (whateley & porter 1983). hip, which is managed by ezemvelo kwazulu-natal (kzn) wildlife, is a major contributor to the tourism industry of kzn (aylward & lutz 2003).hip contains the full complement of large mammal herbivores characteristic of the region (including black and white rhinoceros) and is the oldest conservation area in south africa. the park is characterised by high habitat heterogeneity, with vegetation ranging from open grassland and savanna to closed woodland and forest (whateley & porter 1983). the vegetation in the park has thickened significantly over the last century (watson 1995; watson & macdonald 1983) and a study by wigley et al. (2010) found an increase in woody cover from 14% in 1937 to 58% in 2004 in a significant portion of the northern hluhluwe section of the park. visitors to hip go there predominantly to see wildlife (di minin et al. 2012) and are an important source of income for the park, surrounding communities and smaller game reserves (aylward & lutz 2003). the annual economic contribution of hip to ezemvelo kzn wildlife is disproportionately large for its area and expenditures (table 1) and therefore any loss in revenue that hip suffers is likely to have repercussions for ezemvelo kzn wildlife parks in general. reduced revenue generation from tourism may ultimately influence operational costs and conservation goals of the parks authority. table 1: the relative contribution of hluhluwe–imfolozi park to annual income of ezemvelo kwazulu-natal wildlife. visibility of animals in different habitats to determine the impact of vegetation on animal visibility we used census data collected by the earthwatch institution in 2008, 2010 and 2012. during censuses, observation teams walked a total of 26 fixed-line transects of between 3.9 km and 10.4 km (cromsigt et al. 2009). teams of two observers walked transects just after sunrise over a three-month period in the dry season (from july to october). all sightings of herbivore species larger than a hare within 500 m of both sides of a transect were recorded. the species and abundance, the gps coordinates of the observer and the distance and bearing of the animal(s) from the observer were recorded (cromsigt et al. 2009). the vegetation in each transect was classified into five different categories according to increasing woodiness: grassland, open savanna, closed savanna, thicket (dense, impenetrable woody vegetation) and forest. we used these data to make inferences about whether animal visibility is significantly reduced in habitats with dense tree cover. besides analysing the visibility of all species in combination, we also looked at nyala (tragelaphus angasi), kudu (tragelaphus strepsiceros) and impala (aepyceros melampus) in isolation. nyala and kudu are predominantly browsers and so would be expected to be found in forested areas. impala are predominantly grazers and so would be expected to be found mainly in savanna areas. we hypothesised that regardless of feeding preferences, larger herd sizes and higher animal densities would be recorded in open grassland and savanna areas because animals would more likely have been counted in these habitats, as visibility is not obstructed by trees (marshall et al. 2008). we used the statistical software package r to build a generalised linear model to predict the response variable density of animals per area (km2). we used habitat as an explanatory variable to determine if it explained a significant amount of deviance in the model. we included transect length as an offset to account for different sizes of each habitat. this produced a response variable of animal density. in this step, we were interested in the apparent rather than the actual density of animals in each habitat, so there was no need to standardise our estimates using correction factors. visitor surveys we used structured questionnaires to survey 220 visitors to the park in july 2009 and may 2010. we approached visitors in accommodation camps, at the entrances to the park and in recreational areas throughout the park. the park has two main accommodation areas, with hilltop being a luxury camp in the hluhluwe section of the park and mpila a budget camp in the imfolozi section of the park. all visitor groups present were approached, with one representative per group completing the questionnaire. the questionnaire included both open-ended and multiple-choice questions (see online appendix 1). vegetation change and desirability we asked repeat visitors to describe any change in vegetation they may have noticed since their first visit, without explaining the woody plant encroachment phenomenon to them. visitors were also shown photographs depicting three different vegetation types (open savanna, heterogeneous landscapes with savanna and forest elements, and encroached thicket) and asked which vegetation they most enjoyed driving through. the responses to these questions were used to establish the main reasons for visiting the park and which aspects of the park and its wildlife visitors were most attracted to. visitors were also asked to identify, from the same pictures, in which vegetation they had seen the most wildlife. visitor demography and reasons for visiting at the onset of the survey we established where visitors were from, where they were staying, whether they had visited the park before and, if so, how often. visitors were asked why they had chosen to visit hip and to indicate which animals they most wanted to see in the park. this allowed us to determine whether responses differed depending on visitor origin and accommodation choice using contingency tables and chi-square tests. effect of encroachment on visitor satisfaction we then explained the woody plant encroachment phenomenon. visitors were asked to record, on a scale between 1 and 5, how strongly they agreed with a series of statements (5 = strongly agree; 1 = strongly disagree; also see online appendix 1) to establish whether they would return to visit the park despite a reduction in the visibility of wildlife. the results were subsequently related to the demographic profiles of visitors using contingency tables and chi-square tests. results top ↑ visibility of animals in different habitats results from a generalised linear model of the form:density = ß0 + ß1 × habitat + ε [eqn 1] suggested that mean density was significantly higher in grasslands and open habitats than in closed habitats (see results of model summarised in table 2). forests had the lowest observed animal densities (0.38 animals/km2). some 30% of the deviance in the model was explained by habitat. the open habitats had much larger animal groups, with a maximum of 300 individuals in one group in grassland, and a maximum of just 50 in forest (figure 1a). this pattern was repeated regardless of feeding ecology when animals were grouped according to species (figure 1b–d). these results suggest that an increase in woody plants would likely cause a reduction in the apparent density of animals. table 2: results from a generalised linear model investigating how much of the variation in the density of animals is explained by differences in habitat. figure 1: boxplots showing medians and quartiles of the number of animals recorded at each sighting in each habitat type: (a) all species, (b) kudu, (c) impala and (d) nyala. visitor perceptions before woody plant encroachment was explained to visitors, they were asked how long they had been visiting the park and whether they had noticed a change in vegetation since they had first visited. of the 64.7% of returning visitors (n = 66) who answered that they had noticed a change, 80% (n = 53) described a vegetation change that could be identified as woody plant encroachment. figure 2 shows the percentage of visitors who had observed vegetation change as a function of how long ago their first visit was. time since first visit increased the likelihood of having noticed vegetation change (χ2 = 22.30, df = 6, p = 0.001). of the 20 visitors who had been visiting the park for more than 30 years, 80% (n = 16) had noticed woody encroachment. a further 10% had noticed some change in the vegetation but did not describe it. this was in contrast to responses from visitors who had been visiting for less than 10 years (n = 48), with only 31.2% having noticed woody plant encroachment and 12.5% having noticed some change in vegetation. figure 2: percentage of return visitors who noticed vegetation change over time. when visitors were asked to identify, using photographs, the habitat they thought they had seen most animals in, 63% chose photographs showing open savannas or landscapes with both forest and savanna elements. only 37% of visitors indicated that they had seen more animals in closed thicket and forest landscapes. we used the same photographs to ask visitors which scenery they enjoyed most when driving through the park. according to the responses, 48% of visitors preferred driving through landscapes with forest and savanna elements (i.e. heterogeneous landscapes), 27% preferred open savanna landscapes and 25% preferred closed habitats. demographic profiles of visitors the demographic profiles of visitors are shown in table 3 and table 4. although the majority of visitors to the park were local travellers (i.e. from kzn or elsewhere in or close to south africa1) a large proportion (45.5%) of visitors were international guests. most guests were day visitors (58.1%). of the overnight visitors (n = 92), 65.2% stayed at the hilltop camp and the majority of these (60%) were international visitors. a chi-square test confirmed that local overnight visitors were less likely to stay at hilltop, whilst international visitors were more likely to stay there than would be expected by chance (χ2 = 13.04, df = 4, p = 0.011). table 3: demographic profile and accommodation choice of visitors. table 4: contingency table showing the number of return and first-time visitors to the park and their respective accommodation choices. of the surveyed visitors, 47.3% had been to the park before. more than half of the overnight visitors (n = 48) were return visitors. most of the day visitors (56.1%, n = 72) had not visited the park before. likelihood of staying in park accommodation was not different for first-time or regular visitors (χ2 = 3.62, df = 2, p = 0.16) (table 4). reasons for visiting hluhluwe–imfolozi park this was an open-ended question, so answers were combined into categories. the most commonly cited reason for choosing to visit hip was that it was close or convenient (figure 3). for international visitors, the most commonly cited reason for visiting was that it was recommended by someone or because it formed part of a tour. tranquillity or being in nature was most commonly cited by local visitors as a reason for choosing the park. the park’s heritage and its well-known rhino conservation efforts appeared to be more important to international than local visitors. a chi-square test confirmed that the origin of visitors (i.e. local vs international) influenced their reasons for choosing to visit hip (χ2 = 26.8, df = 10, p = 0.003). surprisingly, only 17.7% of visitors cited seeing animals in general, and the ‘big five’ (lion, leopard, elephant, white rhino and buffalo) in particular, as the most important reason for visiting the park. however, in response to the statement: ‘the main reason i am visiting this park is to see animals,’ 58.6% strongly agreed, whilst only 5.58% strongly disagreed. visitor demography strongly influenced how they answered this question (χ2 = 38.9, df = 8, p < 0.0001): 46.0% of visitors who strongly agreed with the statement were international guests, whilst 83.3% of respondents who strongly disagreed with the statement were local visitors, specifically from kzn. visitors from elsewhere in africa were more likely to have a neutral view, although the majority of them still strongly agreed with the statement. figure 3: reasons cited for visiting the park for (a) all visitors combined, (b) visitors from kwazulu-natal (local), (c) visitors from the rest of south africa and africa (excluding kwazulu-natal province) and (d) international visitors. the ‘big five’ tended to be the most desirable animals (figure 4) for visitors to see whilst in the park, although giraffe were more popular than buffalo. animal preferences were affected by visitor origin (χ2 = 51.90, df = 26, p = 0.002), with giraffe and zebra being more popular sightings amongst international visitors, and buffalo and birds being more popular sightings amongst local visitors. whether visitors had been to the park before or not also affected their desire to see certain animals (χ2 = 78.7, df = 52, p = 0.0098): the desire to see leopard, impala, nyala and cheetah increased if people had been to the park before, but the desire to see zebra decreased. figure 4: animals that visitors most wanted to see, as a percentage of all visitors. visitor response to changing vegetation although the vast majority of visitors (87.4%) indicated that they would like to return to the park given the opportunity, 42.4% of visitors agreed or strongly agreed with the statement that they would not return to the park if it became more difficult to see animals (table 5). accommodation choice and whether they had been to the park before affected their response to this statement (χ2 = 36.86, df = 22, p = 0.025). day visitors or visitors staying at hilltop were less likely to return if game became more difficult to spot than those staying at mpila. first-time visitors were also less likely to return than regular visitors if animals became more difficult to see. table 5: contingency table showing the proportion of return visitors, their accommodation choice, and response to the statement ‘i would not visit the park again if game became more difficult to spot’. a chi-square test found that visitors whose responses indicated that they were unlikely to return if encroachment made wildlife more difficult to see were likely to agree with the statement that encroachment should be controlled (χ2 = 50.18, df = 16, p < 0.0001). day visitors were also more likely to agree with controlling encroachment than overnight guests staying in park accommodation (χ2 = 19.46, df = 8, p = 0.01). discussion top ↑ woody plant encroachment will impact on many savanna ecosystem goods and services such as fodder availability in rangelands (de klerk 2004), biodiversity (parr et al. 2012), hydrology and nutrient cycling (archer, boutton & mcmurtry 2004). it is also likely to have regional-scale feedbacks to the earth–atmosphere system because of changes in albedo and gas exchange (asner et al. 2004; beerling & osborne 2006). this study set out to determine whether woody plant encroachment would have additional economic impacts due to a reduction in game viewing capacity in hip, where evidence of extensive woody plant encroachment has been seen over the last century (wigley et al. 2010). specifically, we set out to investigate (1) whether woody encroachment would reduce visibility of animals, (2) whether guests visited the park predominantly to see animals and (3) whether reduced visibility of animals could therefore result in reduced visitor numbers and, consequently, reduced park revenue. our study showed clear evidence that wildlife visibility is compromised by woody plant encroachment (figure 1), regardless of the species of animal. in addition, most visitors indicated that they had seen more animals in open and heterogeneous habitats. visitors also found heterogeneous landscapes with typical savanna characteristics, such as open plains and single large trees, more aesthetically pleasing than encroached areas. besides the aesthetics, heterogeneous vegetation is ecologically important in increasing the carrying capacity of populations, because it provides a diversity of resources (fryxell et al. 2005; wang et al. 2006). in a previous study undertaken in hip, cromsigt, prins and olff (2009) suggested that habitat heterogeneity might facilitate herbivore diversity in savanna ecosystems. whilst the conclusion that denser vegetation will result in reduced visibility of animals may seem an obvious one, it is likely to have numerous less obvious knock-on effects, one of which is the potential reduction in visitor numbers to game reserves. our results established that whilst wildlife viewing may not be the main reason for visitors choosing hip (figure 3), the majority of visitors were indeed intent on seeing wildlife. this was, however, affected by visitor origin, with international visitors being more interested in seeing animals than local visitors. di minin et al. (2012) found a similar trend when investigating the relative importance of the ‘big five’ in hip, with more wealthy and less experienced international visitors most interested in seeing these animals. whether visitors will stop visiting the park because of encroachment was not clear from our results, but the responses to our questions have led us to make certain inferences. people who visit the park because it is close or convenient (figure 3) are unlikely to be much affected by a change in woody vegetation as the location of the park and its proximity to other areas will not change. those who visit the park because of its heritage and rhino conservation efforts, as well as those who have come to escape city life and experience the tranquillity of nature, are also unlikely to stop visiting the park as a results of encroachment. for international visitors, however, recommendations and choice of tour operator were important reasons for visiting hip. mmopelwa, kgathi and molefhe (2007) found that in botswana international visitors were more willing to pay higher rates for wildlife tourism, a finding mirrored by our observations that international tourists often stayed in the more expensive accommodation. tour operators are unlikely to take tourists to a game reserve where wildlife viewing success is poor, and the same may be expected from guide book recommendations. this suggests that the park could lose a significant fraction of their international visitors (and therefore highest spenders) because of woody plant encroachment. although the majority of visitors indicated they had enjoyed their experience and would like to return to the park, a large portion of visitors confirmed that reduced visibility of animals would impact their decision to the return to the park (with nuances depending on whether they were regular visitors and choice of accommodation). based on visitor responses we estimated that hilltop could lose at least half of its potential future visitors and about 40% of its regular visitors if animals became more difficult to see. if we consider return visitors as definite future visitors, the park would lose about 40% of its definite future visitors and about one third of these would be visitors to hilltop (i.e. the visitors who are likely to spend the most money whilst in the park). in addition, with an estimated return rate of 47% for first-time visitors, hilltop could lose nearly half of potential future visitors. this combines to a possible total loss of about 40% of potential future visitors to the park if animals became more difficult to see. these results suggest that park managers should consider suitable encroachment control measures if they are to ensure return visits. on the whole, visitors did not object to the suggestion of controlling encroachment, so park managers are not likely to face public outcry if such measures were put in place. conclusion top ↑ our study has shown strong evidence that woody plant encroachment influences visitor experience in hip and suggests that visitor numbers might diminish in heavily encroached areas. woody plant encroachment is not merely a debate on conservation objectives for park management, but could negatively influence the economy of the park. our results suggest that the park may lose a significant portion of visitors if woody encroachment continues unchecked. however, studies have shown that visitor demand for large wildlife viewing is unlikely to disappear and the overall number of wildlife tourists in africa is unlikely to drop (boshoff et al. 2007; goodwin & leader-williams 2000; lindsey et al. 2007; mmopelwa et al. 2007; obua & harding 1996). rather, we may see a shift in visitor numbers to different parks as visitors choose parks where they are more likely to have their desired experience. to ensure success of their parks in the future, managers need to make decisions based on conservation principles, but with an understanding that conservation will be facilitated by the revenue generated by visitors (boshoff et al. 2007; kiss 2004; lindsey et al. 2007; mmopelwa et al. 2007). as hip contributes significantly to the revenue of ezemvelo kzn wildlife as a whole (aylward & lutz 2003), a decrease in revenue may affect the organisation significantly. in addition, ezemvelo kzn wildlife employs many members of the local community (aylward & lutz 2003) and many livelihoods could therefore be affected by a drop in visitor numbers, as well as the many smaller businesses and private game reserves and local ecotourism ventures that benefit from being in close proximity to such a large tourist attraction (bookbinder et al. 1998; kiss 2004). our conclusions on the impact of habitat change on park visitors are not restricted to hip, as woody encroachment is a widespread phenomenon in many savannas in africa. it will be prudent for managers to consider strategic management of the phenomenon using fire and other management tools. acknowledgements top ↑ this study was made possible through funding from the university of cape town woolworths fellowship and the german academic exchange programme (daad). thanks to the earthwatch institute for mammal census data, geoffrey clinning for providing data, and amy rose wilkinson for assistance in the field. thanks also to field and administrative staff at hluhluwe–imfolozi park for allowing us to undertake the study and providing us with assistance at every stage. competing interests the authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article. authors’ contributions w.j.b. (university of cape town) was the project leader. e.f.g. (university of cape town) and w.j.b. were both responsible for project design. e.f.g. undertook all fieldwork and analyses and wrote the manuscript. w.j.b. made conceptual contributions to the manuscript. references top ↑ akama, j.s. & kieti, d.m., 2003, ‘measuring tourist satisfaction with kenya’s wildlife safari: a case study of tsavo west national park’, tourism management 24(1), 73–81. http://dx.doi.org/10.1016/s0261-5177(02)00044-4 akama, j.s., maingi, s. & camargo, b., 2011, ‘wildlife conservation, safari tourism and the role of tourism certification in kenya: a postcolonial critique’, tourism recreation research 36, 1–11. archer, s., schimel, d.s. & holland, e.a., 1995, ‘mechanisms of shrubland expansion: land use, climate or co2’, climatic change 29, 91–99. http://dx.doi.org/10.1007/bf01091640 archer, s., boutton, t. & mcmurtry, c., 2004, ‘carbon and nitrogen accumulation in a savanna landscape: field and modeling perspectives’, in m. shiomi, h. kawahata, h. koizumi, a. tsuda & y. awaya (eds.), global environmental change in the ocean and on land, pp. 359–373, terra scientific, tokyo. asner, g.p., archer, s., hughes, r.f., ansley, r.j. & wessman, c.a., 2003, ‘net changes in regional woody vegetation cover and carbon storage in texas drylands, 1937–1999’, global change biology 9(3), 316–335. http://dx.doi.org/10.1046/j.1365-2486.2003.00594.x asner, g.p., elmore, a.j., olander, l.p., martin, r.e. & harris, t., 2004, ‘grazing systems, ecosystem responses, and global change’, annual review of environment and resources 29, 261–299. http://dx.doi.org/10.1146/annurev.energy.29.062403.102142 aylward, b.a. & lutz, e. (eds.), 2003, nature tourism, conservation, and development in kwazulu-natal, south africa, world bank publications, washington. beerling, d.j. & osborne, c.p., 2006, ‘the origin of the savanna biome’, global change biology 12(11), 2023–2031. http://dx.doi.org/10.1111/j.1365-2486.2006.01239.x bond, w.j., 2008, ‘what limits trees in c4 grassland and savannas?’, annual review of ecology, evolution and systematics 39, 641–659. http://dx.doi.org/10.1146/annurev.ecolsys.39.110707.173411 bond, w.j. & midgley, g.f., 2012, ‘co2 and the uneasy interactions of trees and savanna grasses’, philosophical transactions of the royal society biological sciences 367, 601–612. http://dx.doi.org/10.1098/rstb.2011.0182, pmid:22232770 bond, w.j. & parr, c.l., 2010, ‘beyond the forest edge: ecology, diversity and conservation of the grassy biomes’, biological conservation 143(10), 2395–2404. http://dx.doi.org/10.1016/j.biocon.2009.12.012 bookbinder, m.p., dinerstein, e., rijal, a., cauley, h. & rajouria, a., 1998, ‘ecotourism’s support of biodiversity conservation’, conservation biology 12(6), 1399–1404. http://dx.doi.org/10.1046/j.1523-1739.1998.97229.x boshoff, a.f., landman, m., kerley, g.i.h. & bradfield, m., 2007, ‘profiles, views and observations of visitors to the addo elephant national park, eastern cape, south africa’, south african journal of wildlife research 37(2), 189–196. http://dx.doi.org/10.3957/0379-4369-37.2.189 bowman, d.m.j.s., murphy, b.p. & banfai, d.s., 2011, ‘has global environmental change caused monsoon rainforests to expand in the australian monsoon tropics?’, landscape ecology 25, 1247–1260. http://dx.doi.org/10.1007/s10980-010-9496-8 britz, m.l. & ward, d., 2007, ‘dynamics of woody vegetation in a semi-arid savanna, with a focus on bush encroachment’, african journal of range and forage science 24(3), 131–140. http://dx.doi.org/10.2989/ajrfs.2007.24.3.3.296 brown, j.s., 1988, ‘patch use as an indicator of habitat preference, predation risk, and competition’, behavioral ecology and sociobiology 22(1), 37–47. http://dx.doi.org/10.1007/bf00395696 brown, j.s., 1992, ‘patch use under predation risk: i. models and predictions’, annales zoologici fennici 29, 301–309. brown, j.s., 1999, ‘vigilance, patch use and habitat selection: foraging under predation risk’, evolutionary ecology research 1(1), 49–71. brown, j.s., laundré, j.w. & gurung, m., 1999, ‘the ecology of fear: optimal foraging, game theory, and trophic interactions’, journal of mammalogy 80(2), 385–399. http://dx.doi.org/10.2307/1383287 coetsee, c., gray, e.f., wakeling, j., wigley, b.j. & bond, w.j., 2013, ‘low gains in ecosystem carbon with woody plant encroachment in a south african savanna’, journal of tropical ecology 29(1), 49–60. http://dx.doi.org/10.1017/s0266467412000697 costanza, r., d’arge, r., de groot, r., farber, s., grasso, m., hannon, b. et al., 1997, ‘the value of the world’s ecosystem services and natural capital’, nature 387(6630), 253–260. http://dx.doi.org/10.1038/387253a0 cromsigt, j.p., van rensburg, s.j., etienne, r.s. & olff, h., 2009 ‘monitoring large herbivore diversity at different scales: comparing direct and indirect methods’, biodiversity and conservation 18(5), 1219–1231. http://dx.doi.org/10.1007/s10531-008-9506-1 cromsigt, j.p., prins, h.t.h. & olff, h., 2009, ‘habitat heterogeneity as a driver of ungulate diversity and distribution patterns: interaction of body mass and digestive strategy’, diversity and distributions 15, 513–522. http://dx.doi.org/10.1111/j.1472-4642.2008.00554.x de klerk, j.n., 2004, bush encroachment in namibia: report on phase 1 of the bush encroachment research, monitoring, and management project, windhoek, namibia, ministry of environment and tourism, directorate of environmental affairs, windhoek. di minin, e., fraser, i., slotow, r. & macmillan, d.c., 2012, ‘understanding heterogeneous preference of tourists for big game species: implications for conservation and management’, animal conservation 16(3), 249–258. http://dx.doi.org/10.1111/j.1469-1795.2012.00595.x du toit, j.t. & owen-smith, n., 1989, ‘body size, population metabolism and habitat specialization among large african herbivores’, american naturalist 133, 736–740. fensham, r.j., fairfax, r.j. & archer, s., 2005, ‘rainfall, land use and woody vegetation cover change in semi-arid australian savanna’, journal of ecology 93(3), 596–606. http://dx.doi.org/10.1111/j.1365-2745.2005.00998.x fryxell, j.m., wilmshurst, j.f., sinclair, a.r.e., haydon, d.t., holt, r.d. & abrams, p.a., 2005, ‘landscape scale, heterogeneity, and the viability of serengeti grazers’, ecology letters 8, 328–335. http://dx.doi.org/10.1111/j.1461-0248.2005.00727.x goodwin, h.j. & leader-williams, n., 2000, ‘tourism and protected areas – distorting conservation priorities towards charismatic megafauna’, in a. entwhistle & n. dunstone (eds.), priorities for the conservation of mammalian diversity: has the panda had its day?, pp. 257–275, cambridge university press, cambridge. kiss, a., 2004, ‘is community-based ecotourism a good use of biodiversity conservation funds?’, trends in ecology and evolution 19(5), 232–237. http://dx.doi.org/10.1016/j.tree.2004.03.010, pmid:16701261 laundré, j.w., hernández, l. & altendorf, k.b., 2001, ‘wolves, elk, and bison: reestablishing the “landscape of fear” in yellowstone national park, usa’, canadian journal of zoology 79(8), 1401–1409. http://dx.doi.org/10.1139/z01-094 lindsey, p.a., alexander, r., mills, m.g.l., ramanach, s. & woodroffe, r., 2007, ‘wildlife viewing preferences of visitors to protected areas in south africa: implications for the role of ecotourism in conservation’, journal of ecotourism 6(1), 19–33. http://dx.doi.org/10.2167/joe133.0 marshall, a.r., lovett, j.c. & white, p.c.l., 2008, ‘selection of line-transect methods for estimating the density of group-living animals: lessons from the primates’, american journal of primatology 70, 452–462. http://dx.doi.org/10.1002/ajp.20516, pmid:18240143 mmopelwa, g., kgathi, d.l. & molefhe, l., 2007, ‘tourists’ perceptions and their willingness to pay for park fees: a case study of self-drive tourists and clients for mobile tour operators in moremi game reserve, botswana’, tourism management 28(4), 1044–1056. http://dx.doi.org/10.1016/j.tourman.2006.08.014 moleele, n.m., ringrose, s., matheson, w. & van der post, c., 2002, ‘more woody plants? the status of bush encroachment in botswana’s grazing areas’, journal of environmental management 64(1), 3–11. http://dx.doi.org/10.1006/jema.2001.0486, pmid:11876072 obua, j. & harding, d.m., 1996, ‘visitor characteristics and attitudes towards kibale national park, uganda’, tourism management 17(7), 495–505. http://dx.doi.org/10.1016/s0261-5177(96)00067-2 parr, c.l., gray, e.f. & bond, w.j., 2012, ‘cascading biodiversity and functional consequences of a global change-induced biome switch’, diversity and distributions 18(5), 493–503. http://dx.doi.org/10.1111/j.1472-4642.2012.00882.x ratajczak, z., nippert, j.b. & collins, s.l., 2012, ‘woody encroachment decreases diversity across north american grasslands and savannas’, ecology 93(4), 697–703. http://dx.doi.org/10.1890/11-1199.1, pmid:22690619 ripple, w.j. & beschta, r.l., 2004, ‘wolves and the ecology of fear: can predation risk structure ecosystems?’, bioscience 54(8), 755–766. http://dx.doi.org/10.1641/0006-3568(2004)054%5b0755:wateof%5d2.0.co;2 roques, k.g., o’connor, t.g. & watkinson, a.r., 2001, ‘dynamics of shrub encroachment in an african savanna: relative influences of fire, herbivory, rainfall and density dependence’, journal of applied ecology 38(2), 268–280. http://dx.doi.org/10.1046/j.1365-2664.2001.00567.x sankaran, m., ratnam, j. & hanan, n.p., 2004, ‘tree–grass coexistence in savannas revisited – insights from an examination of assumptions and mechanisms invoked in existing models’, ecology letters 7, 480–490. http://dx.doi.org/10.1111/j.1461-0248.2004.00596.x sinclair, a.r.e., mduma, s. & brashares, j.s., 2003, ‘patterns of predation in a diverse predator–prey system’, nature 425(6955), 288–290. http://dx.doi.org/10.1038/nature01934, pmid:13679915 sirami, c., seymour, c., midgley, g. & barnard, p., 2009, ‘the impact of shrub encroachment on savanna bird diversity from local to regional scale’, diversity and distributions 15(6), 948–957. http://dx.doi.org/10.1111/j.1472-4642.2009.00612.x skarpe, c., 1990, ‘structure of the woody vegetation in disturbed and undisturbed arid savanna, botswana’, plant ecology 87(1), 11–18. http://dx.doi.org/10.1007/bf00045650 van auken, o.w., 2000, ‘shrub invasions of north american semiarid grasslands’, annual review of ecology and systematics 31, 197–215. http://dx.doi.org/10.1146/annurev.ecolsys.31.1.197 wang, g., hobbs, n.t., boone, r.b., illius, a.w., gordon, i.j., gross, j.e. et al., 2006, ‘spatial and temporal variability modify density dependence in populations of large herbivores’, ecology 87(1), 95–102. http://dx.doi.org/10.1890/05-0355, pmid:16634300 watson, h.k. & macdonald, i.a.w., 1983, ‘vegetation changes in the hluhluwe–umfolozi game reserve complex from 1937 to 1975’, bothalia 14(2), 265–269. watson, h.k., 1995, ‘management implications of vegetation changes in hluhluwe-imfolozi park’, south african geographical journal 77(2), 77–83. http://dx.doi.org/10.1080/03736245.1995.9713595 whateley, a. & porter, r.n., 1983, ‘the woody vegetation communities of the hluhluwe-corridor-umfolozi game reserve complex’, bothalia 14(3), 745–758. wigley, b.j., bond, w.j. & hoffman, m.t., 2010, ‘thicket expansion in a south african savanna under divergent land use: local vs. global drivers?’, global change biology 16(3), 964–976. http://dx.doi.org/10.1111/j.1365-2486.2009.02030.x footnote top ↑ 1. very few african visitors were from outside south africa and therefore all african and south african visitors were grouped together. hudson.qxd new records of 45 bird species in the desert margins area of the northwest province, south africa a. hudson & h. bouwman hudson, a. & h. bouwman. 2006. new records of 45 bird species in the desert margins area of the north-west province, south africa. koedoe 49(1): 91–98. pretoria. issn 0075-6458. new records and possible range extensions are reported on 45 bird species (ca 25% of the 167 species recorded during surveys) in the bophirima district of the north-west province, south africa. the findings were compared with data in the atlas of southern african birds. the main reasons for these new records may be ascribed to the low number of visits during the atlas project, higher precipitation during the time of our observations versus that of the atlas project, and possibly an increased suitability of the area for some bird species due to human habitation. these new records also provide additional information that may be useful in conservation planning, especially in arid areas. heuningvlei pan in particular, should be considered for additional conservation measures. key words: arid areas, molopo nature reserve, heuningvlei pan, wetland. adrian hudson (drkah@puk.ac.za ) & henk bouwman, school for environmental sciences and development (zoology), north-west university (potchefstroom campus), private bag x6001, potchefstroom, south africa. issn 0075-6458 91 koedoe 49/1 (2006) introduction birds, to a greater extent than other animals, have the ability to move away from adverse conditions and towards areas with more favourable conditions (dean 2004). their ability to fly reduces the significance of distance from one area to the other, and negates boundaries that would keep other species (such as larger game) restricted to a certain area (o' halloran et al. 2002). the two basic strategies desert animals use for survival are to be sedentary and using behavioural or physiological adaptations, or to move seasonally or opportunistically to more suitable areas (dean 2004). one form of migration found in birds, including desert birds, is nomadism (dean 2004). nomadism occurs when birds move from area to area in order to utilise spatially and temporally patchy resources. nomadic avifauna is characterised by the dramatic increase of numbers of birds in suitable areas or decrease in unsuitable areas. the topic of a great deal of modern-day literature is the dynamic environment in which we as humans and other organisms on earth find ourselves (parody et al. 2001). stochastic, natural and long-term alterations of the environment can take on various forms, namely, changing climatic conditions, changing geological structure and concomitant changes in vegetation structure. the environmental changes can also be of an anthropogenic nature. humans change the environment in a number of ways, for instance: changing watercourses, draining wetlands, destruction of natural habitats for various reasons, urbanisation or practices that promote desertification. these can all lead to areas becoming unsuitable for bird species that previously occurred in that area (parody et al. 2001). other anthropogenic influences may even increase an area’s suitability for birds. thousands of small man-made wetlands are being constructed due to watering points in the more arid areas of southern africa to provide water for livestock. large areas of grain crops are providing food hudson.qxd 2006/04/17 10:55 pm page 91 sources for granivorous birds. tree plantations are providing suitable habitats for some tree dwelling birds in areas where they did not previously exist, and urbanisation is providing additional habitats for a number of species in areas with an otherwise flat topography (hockey 2003). the changing environment is causing bird species to move away from habitats that have become unsuitable, expanding their ranges into areas that have become more suitable. in the 1950s, the range of black-chested prinia prinia flavicans had a southern boundary that extended as far as the southeastern free state. by the 1990s, this had shifted as far as the southern cape coast. similarly, the african goshawk accipiter tachiro occurred only as far as the eastern cape in the 1950s, but has expanded its range as far as the cape west coast (hockey 2003). because of these changing ranges it is necessary for ornithologists to regularly update data as to where species occur. here we present some significant new records and possible range extensions of birds observed, while on research and training surveys in and around the molopo nature reserve and heuningvlei pan in the bophirima district of the north-west province (south africa). normally new bird records for many areas are to be expected, but the remoteness of the area in question, the increased attention given to desert margin areas in a global context, as well as the number of new records obtained, warrants closer investigation and reporting. the surprising number of new records reported here may also have conservation implications. study area the study area (fig. 1) included four sites in the bophirima district of the north-west province in south africa, ranging from morokweng (26°27's; 23°29'e) northwestwards to the botswana border. the vegetation type of the area is kalahari plains thorn bushveld (kalahari thornveld), characterised by a fairly well-developed tree stratum, a moderately-developed shrub stratum and a varying amount of grass cover (low & koedoe 49/1 (2006) 92 issn 0075-6458 fig. 1. location of grids for new records of bird species, in the bopherima district of the north-west province, south africa. hudson.qxd 2006/04/17 10:55 pm page 92 rebelo 1996). the vegetation types of the study sites are identical, but vegetation structure varies due to overgrazing, management practices and/or the use of fire as a management tool in some areas. one site includes a natural saltpan (heuningvlei; 2623ac) surrounded by heavily overgrazed kalahari thornveld. the northwest corner of the pan is edged by a rocky outcrop and fed by a number of permanent freshwater springs. these springs cause that section of the pan to exist as a small but perennial marsh-type wetland with brackish water. other ephemeral pans—most of them highly impacted by overgrazing and urbanisation—are also located throughout the region. the molopo nature reserve (2522dd) is one of the very few places that can be considered as natural. materials and methods the atlas of southern african birds (harrison et al. 1997a, 1997b) was used as baseline of existing records. the new records reported here are the result of more than 12 surveys, conducted in the area over a four-year period (2003–2006), for various activities in support of the desert margins programme in south africa. intensive bird ringing was also done on some of these surveys. none of these surveys were specifically undertaken to gather new records. binoculars, a telescope, mistnetting equipment, and a global positioning system (gps) were used. new records were obtained with the following methods: a) transects measuring either 500 m or 1 km each were surveyed at specific sites over a twentyfour month period. these transects were randomly selected and walked at a constant pace, or were selected based on the condition of the vegetation before and after a fire. bird species were observed along these transects up to a distance of about 100 m from the transect centre line. more detailed findings of these transects will be reported elsewhere. b) mistnetting was also done at the study sites in order to trap and identify less conspicuous birds (approximately 3000 birds were ringed). mistnetting was, however, hampered by the openness of the vegetation, heat and strong winds. mistnetting was carried out strictly according to its code of conduct. c) observations of opportunity during transit. results in total, 167 bird species were identified between july 2002 and february 2006. of the 167 species observed in the study area, 45 were new records for quarter degree grids when compared with the sa bird atlas (harrison et al. 1997a, 1997b). the species are named and arranged, according to evolutionary relationships, as per hockey et al. (2005). brief descriptions of these sightings and conditions follows, and is summarised in table 1. white-faced duck dendrocygna viduata a group of 13-17 was recorded during each of the surveys at heuningvlei pan. this flock appeared to be resident at the pan. no breeding was observed. egyptian goose alopochen aegyptiaca this bird is common at water bodies throughout the bophirima district. however, according to harrison et al. (1997a) this is a new record for grid block 2623ac (heuningvlei pan). no breeding was observed. cape teal anas capensis eight birds were recorded at heuningvlei pan during march and december (2003), but were absent from the site during the winter months. yellow-billed duck anas undulata a small flock of seven birds was recorded during each of the surveys at heuningvlei pan. this group also appeared to be resident. no breeding was observed. cape shoveller anas smithii one bird was recorded at heuningvlei pan in march 2003. no subsequent sightings were recorded. red-billed teal anas erythroryncha one bird was recorded at heuningvlei pan in march 2003. no subsequent sightings were recorded. hottentot teal anas hottentota two birds were recorded at heuningvlei issn 0075-6458 93 koedoe 49/1 (2006) hudson.qxd 2006/04/17 10:55 pm page 93 koedoe 49/1 (2006) 94 issn 0075-6458 pan during march and december (2003), but were absent from the site during the winter months greater honeyguide indicator indicator one bird of this species was ringed on the molopo nature reserve and another seen on driefontein communal farm during the march 2003 surveys. brown-hooded kingfisher halcyon albiventris a single bird was sighted during a survey on molopo nature reserve in july 2004. eurasian bee-eater merops apiaster european bee-eaters were recorded once at both heuningvlei pan and molopo nature reserve. speckled pigeon columba guinea a large number of these birds were recorded at both heuningvlei pan and driefontein communal farm (2623ad). african crake crex egregia a single individual was recorded at heuningvlei pan during the march survey in 2003, but not on any of the subsequent surveys. baillons’s crake porzana pusilla a single individual was seen at heuningvlei pan in february 2006. common moorhen gallinula chloropus a single individual was recorded at heuningvlei pan during the march survey in 2003, but not on any of the subsequent surveys. red-knobbed coot fulica cristata a large population of between 40-50 individuals appear to be resident at heuningvlei pan. these individuals were recorded during all the surveys done at the site and showed little or no variation in numbers. chicks were observed. greenshank tringa nebularia a single bird was seen at heuningvlei pan during the march survey in 2003, but not on any of the subsequent surveys. common sandpiper tringa hypoleucos two birds were recorded at heuningvlei pan during march and december (2003), but were absent from the site during the winter months curlew sandpiper calidris ferrugine one or possibly more of this species were seen on two occasions at heuningvlei pan. ruff philomachus pugnax a single bird was recorded at heuningvlei pan, during the march survey in 2003, but not on any of the subsequent surveys. african jacana actophilornis africana a single bird was sighted during each of the six surveys done at heuningvlei pan. pied avocet recurvirostra avosetta a resident group of eight birds seemed to be resident at heuningvlei pan. no breeding was observed. burchell’s courser cursorius rufus a total of five birds were spotted at different times at lafras commercial farm (2522dd) and heuningvlei pan. gymnogene polyboroides typus one individual was recorded at molopo nature reserve in august 2003. no subsequent sightings were recorded jackal buzzard buteo rufofuscus a single individual is resident at heuningvlei pan. peregrine falcon falco peregrinus one individual was recorded at molopo nature reserve in november 2003. no subsequent sightings were recorded little grebe tachybaptus ruficolli a single bird was recorded at heuningvlei pan during the 2003 winter surveys. hudson.qxd 2006/04/17 10:55 pm page 94 black-necked grebe podiceps nigricolis two birds were observed at heuningvlei pan during all of the surveys, indicating residency. black heron egretta ardesiaca single birds were recorded during two of the visits to heuningvlei pan in 2003. grey heron ardea cinerea one grey heron was recorded during the first survey heuningvlei pan. during a later visit a group of five grey herons was recorded. these birds do not appear to be resident. great egret casmerodius albus single birds were recorded during two of the visits to heuningvlei pan in 2003. cattle egret bubulcus ibis cattle egrets are abundant throughout the study area, but according to harrison et al. (1997a) specimens recorded in grid blocks 2623ac and 2623ad are new records. little bittern ixobrychus minutes a single bird was observed during the autumn 2003 visit to heuningvlei pan. it is probable that this individual was a vagrant. greater flamingo phoenicopterus rubber four birds were recorded during each of the december surveys (2003 and 2004) at heuningvlei pan. these birds were not recorded during any of the other surveys. glossy ibis plegadis falcinellus a single bird was recorded during a visit to heuningvlei pan during march 2003. hadeda ibis bostrychia hagedash two birds were recorded at the molopo nature reserve. it is quite reasonable to assume that these birds were just passing through the area. however the range of this species has increased greatly over the last two decades (hockey 2003). african spoonbill platalea alba one bird was recorded at heuningvlei pan during each of three of the surveys. white stork ciconia ciconia two birds were recorded during a survey at heuningvlei pan during 2004. there have been no further sightings of these birds at this location. eurasian golden oriole oriolus oriolus this species was recorded on one occasion during the summer survey on the molopo nature reserve. greater striped swallow hirundo cucullata this species was recorded on one occasion at heuningvlei pan. arrow-marked babbler turdoides jardineii four individuals were sighted on one of the transects at driefontein communal farm in march 2003. these birds were not sighted again during any of the other surveys done in the area. dusky lark pinarocorys nigricans one bird was identified at heuningvlei pan during the march 2003 survey, with an additional sighting recorded during december 2003. short-toed rock-thrush monticola brevipes this species was recorded on two occasions at heuningvlei pan. cape robin-chat cosypha caffra one individual was caught in a mistnet and banded at heuningvlei pan in march 2003. the same individual was identified on two subsequent occasions during october 2003 and december 2003. white-winged widowbird euplectus albonotatus two individuals were recorded at driefontein communal farm (2623ad) during the march 2003 survey. this species was not recorded again during the rest of the study. issn 0075-6458 95 koedoe 49/1 (2006) hudson.qxd 2006/04/17 10:55 pm page 95 koedoe 49/1 (2006) 96 issn 0075-6458 table 1 new records of bird species in the bophirima district of the north-west province, south africa common name species previous new record no. of closest record sightings white-faced duck dendrocygna viduata 2722aa 2623ac 6 egyptian goose alopochen aegyptiaca 2622dd 2623ac 6 cape teal anas capensis 2723ad 2623ac 2 yellow-billed duck anas undulate 2622dc 2623ac 6 cape shoveller anas smithii 2723ad 2623ac 1 redbilled teal anas erythroryncha 2623ba 2623ac 1 hottentot teal anas hottentota 2722dd 2623ac 2 greater honeyguide indicator indicator 2524ca 2623ac 1 brown-hooded kingfisher halcyon albiventris 2623dc 2522dd & 2623ac 1 eurasian bee-eater merops apiaster 2523cb 2522dd & 2623ac 1 speckled pigeon columba guinea 2623ad 2522dd 6 african crake crex egregia 2624bc 2623ac 1 baillon’s crake porzana pusilla 2523dc 2623ac 1 common moorhen gallinula chloropus 2723aa 2623ac 1 red-knobbed coot fulica cristata 2623bb & 2523ab 2623ac 6 greenshank tringa nebularia 2523dc 2623ac 3 common sandpiper tringa hypoleucos 2523db 2623ac 2 curlew sandpiper calidris ferruginea 2523bd & 2723ca 2623ac 2 ruff philomachus pugnax 2623bb 2623ac 2 african jacana actophilornis africana 2523bc 2623ac 6 pied avocet recurvirostra avosetta 2623bb 2623ac 6 burchell’s courser cursorius rufus 2622ab 2522dd & 2623ac 5 gymnogene polyboroides typus 2622ab 2522dd 1 jackal buzzard buteo rufofuscus 2624cb 2623ac 6 peregrine falcon falco peregrinus 2723cc 2522dd 1 little grebe tachybaptus ruficolli 2623bc 2623ac 1 black-necked grebe podiceps nigricolis 2523db 2623ac 6 black heron (egret) egretta ardesiaca 2524cc 2623ac 2 grey heron ardea cinerea 2722bb 2623ac 1 great egret casmerodius albus 2723ad & 2624ca 2623ac 2 cattle egret bubulcus ibis 2622dd 2623ac & 2623ad 6 little bittern ixobrychus minutus 2622dd 2623ac 1 greater flamingo phoenicopterus ruber 2723ad 2623ac 1 glossy ibis plegadis falcinellus 2722bb 2623ac 1 hadeda ibis bostrychia hagedash 2622ab & 2522ab 2522dd & 2623ac 2 african spoonbill platalea alba 2523ac & 2722bb 2623ac 3 white stork ciconia ciconia 2722dc 2623ac 1 eurasian golden oriole oriolus oriolus 2622ab 2522dd 1 greater striped swallow hirundo cucullata 2622db 2522dd & 2623ac 1 arrow-marked babbler turdoides jardineii 2523db 2623ad 1 dusky lark pinarocorys nigricans 2623cc 2623ac 1 short-toed rock-thrush monticola brevipes 2623aa 2623ad 1 cape robin-chat cosypha caffra 2723ad 2623ac 2 white-winged widowbird euplectus albonotatus 2624ad 2623ad 1 yellow-throated petronia petronia superciliaris 2523cb 2522dd 1 hudson.qxd 2006/04/17 10:55 pm page 96 issn 0075-6458 97 koedoe 49/1 (2006) yellow-throated petronia petronia superciliaris these birds are relatively common in small groups on the molopo nature reserve. discussion the data collected showed that 45 of the 167 species observed in the study area were present outside of their previously recorded ranges (table 1). this constitutes ca 25 % of the total number of species identified in the study area since the beginning of 2003. of these, 26 species were water birds or birds closely associated with water. in this regard the herons, egrets and ducks were conspicuous. these were mostly found at heuningvlei pan (the only permanent natural water body within at least 100 km), but were also observed on ephemeral pans, when these had water. some of these pans are closely associated with urban areas. ganyesa completely encloses one such pan, where red-knobbed coot, ruff and curlew sandpipers were observed (these sites have established records). at morokweng, where the town partially encompasses a larger ephemeral pan, the same birds, larger waders such as greenshank and common sandpiper, as well as egyptian goose were also present in low numbers. heuningvlei also has an associated urban community. when the pan is dry (except for the small permanently inundated corner), people use the pan as a shortcut for motor vehicles and donkey carts (but not intensively), closely skirting the permanently inundated area. the birds seemed to have adapted to this, but some disturbance is evident, and the community seems to be made up of transient birds visiting in summer, rather then permanent residents. greater flamingos, african spoonbills and the larger herons were also regularly observed at heuningvlei, but with little pattern indicating a transient use of the pan. ducks, especially the white-faced and yellow-billed ducks were always present at heuningvlei, but no breeding or juveniles were observed. a recent visit to heuningvlei pan (2005) has shown a new threat developing. the dolomitic springs that were previously protected by fencing are now in disrepair, and cattle are grazing within these sensitive areas. three new records of raptors were made. a jackal buzzard was regularly observed at heuningvlei (very likely the same individual), while a gymnogene and peregrine falcon was sighted on the molopo nature reserve. this area is well known for its raptors, and the molopo raptor conservancy is closely associated with the reserve. speckled rock pigeons were only found associated with human habitation, and mostly at ganyesa and morokweng. their numbers seem to be increasing with every visit, and this may therefore constitute a range extension. for terrestrial birds, most new records were obtained from the molopo nature reserve. the brown-hooded kingfisher is a notable new record for this reserve. whether these records represent range extensions or just new records of birds in these areas are not clear. some may be range expansions occurring due to structural changes in the environment caused by management practices. an increase in the amount of surface water available to birds on communal farms, commercial farms and the nature reserve may make the area more habitable for species, such as granivorous birds, that need daily access to water (hockey 2003). the atlas of southern african birds (harrison et al. 1997a, 1997b) (which was used as a guide to assess which birds were new records) was compiled from data obtained during a period with below average rainfall (harrison et al. 1997a, 1997b). this may account for many of the new records for terrestrial birds. this however, does not explain the large number of new records at heuningvlei pan, because this area has a permanent water supply. one factor might be the number of cards completed for these areas, during the southern african bird atlas project. the number of cards completed for hudson.qxd 2006/04/17 10:55 pm page 97 the 2522dd grid square, in which the molopo nature reserve occurs, was 28. this is not a very high number when compared with the 311 cards that were completed for potchefstroom for the same project. only five cards were completed for heuningvlei pan, indicating very few visits. conclusion the new records presented here are probably due to a combination of lack of previous surveys, higher rainfall during this study when compared with the period when data for the atlas of southern african birds was collected, and possibly changes in the availability of resources for birds due to human influences. it would appear that the low reporting intensity for this area during the southern african bird atlas project may be the largest single factor influencing the number of new records. these new records, combined with those from previous efforts, indicate that existing and additional conservation efforts in arid areas should receive more attention. the large number of birds and species recorded at heuningvlei (27 new records for waterassociated birds) indicates that the current protection measures should be assessed to enhance the conservation of this apparently important wetland site in this arid region. acknowledgements we thank the desert margins programme (klaus kelner), national research foundation and the north-west university (potchefstroom campus) for support. we are also indebted to lieb and belinda venter, leone hudson, nikki smith, rina and riaan booysen, ester van der westhuizen and retha hoffman for their eyes and ears. we thank steven gore, previous park warden of the molopo nature reserve, and the north-west province parks and tourism board for access and permission. cecile combrink is thanked for language editing and rené navarro for the data from the southern african bird atlas project. references dean, w.r.j. 2004. nomadic desert birds. berlin: springer-verlag. harrison, j.a., d.g. allan, l.g. underhill, m. herremans, a.j. tree, v. parker & c.j. brown (eds.). 1997a. the atlas of southern african birds. volume 1: non-passerines. johannesburg: birdlife south africa. harrison, j.a., d.g. allan, l.g. underhill, m. herremans, a.j. tree, v. parker & c.j. brown (eds.). 1997b. the atlas of southern african birds. volume 2: passerines. johannesburg: birdlife south africa, hockey, p. 2003. land invasions. africa birds and birding 8(5): 35-41 low, a.b. & t.c. rebelo. 1998. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. o’ halloran, j., p.m. walsh, p.s. giller & t.c. kelly. 2002. forestry and bird diversity in ireland: a management and planning guide. oikos 79: 247-258 parody, j.m., f.j. cuthbert & e.h.decker. 2001. the effect of 50 years of landscape change on species richness and community composition. global ecology & biogeography 10: 305-313. koedoe 49/1 (2006) 98 issn 0075-6458 hudson.qxd 2006/04/17 10:55 pm page 98 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice zietsman.qxd dune vegetation and coastal thicket plant communities in threatened limestone fynbos of andrew’s field and tsaba-tsaba nature reserve, struisbaai, western cape m.m. zietsman and g.j. bredenkamp zietsman, m.m. and g.j. bredenkamp. 2006. dune vegetation and coastal thicket plant communities in threatened limestone fynbos of andrew’s field and tsaba-tsaba nature reserve, struisbaai, western cape. koedoe 49(1): 33–47. pretoria. issn 00756458. the coastal thicket and dune vegetation of andrew’s field and tsaba-tsaba nature reserve was classified using braun-blanquet procedures and twinspan. the vegetation was sampled using 74 randomly stratified sample plots. the floristic composition, cover-abundance of each species, and several environmental variables were recorded in each sample plot. six plant communities were identified, namely, rhus glauca euclea racemosa low to tall closed thicket community; chrysanthemoides monilifera solanum africanum low closed dune shrub community; chrysanthemoides monilifera ehrharta villosa var. maxima low to high closed dune shrub community; ehrharta villosa var. maxima low to short closed dune grassland community; ammophila arenaria low to short closed dune grassland community; and arcthotheca populifolia thinopyrum distichum low to short open beach community. these were subdivided into eight subcommunities and four variants. all communities, sub-communities and variants were described and ecologically interpreted. the distribution of the communities, sub-communities and variants can mainly be ascribed to differences in landform, rockiness of the soil surface the degree of protection / exposure of the vegetation to the dominating winds of the area. key words: coastal thicket, conservation area, endemic plants, limestone fynbos, phytosociology, plant communities, twinspan, western cape m.m. zietsman and g.j. bredenkamp, african vegetation and plant diversity research centre, department of botany, university of pretoria, pretoria, 0002 republic of south africa. issn 0075-6458 33 koedoe 49/1 (2006) introduction a sound knowledge of the ecology of the area is an essential prerequisite for the establishment of efficient wildlife management programmes and compilation of conservation policies for any area (edwards 1972). vegetation science has been applied in the fields of nature conservation for years, but recent developments relate to the application of plant ecological knowledge to environmental management (scheepers 1983). bredenkamp & brown (2001) emphasise the use of plant communities as a reliable basis for any ecological planning and management. to obtain knowledge of the ecology of the study area, a study of the vegetation of the area should be made at the plant community level of organisation (bredenkamp & brown 2001). studying the vegetation of an area allows the identification, description and classification of plant communities. on the basis of the plant communities as vegetation units, management units can be delineated. vegetation studies also allow the identification of ecologically sensitive areas, bush encroached areas or areas infested with alien plants, degraded areas, habitats of rare or endangered plant species, and habitats of specific animals. the andrew’s field and tsaba-tsaba reserve is situated in the bredasdorp/riversdale centre of endemism (cowling 1992). this centre of endemism refers to a welldefined group of plants, confined to the limezietsman.qxd 2006/04/17 08:48 pm page 33 stone of the bredasdorp formation and associated colluvial deposits (heydenrych 1994). according to hilton-taylor & le roux (1989) limestone fynbos is one of the most threatened vegetation types in the cape floristic region. factors threatening limestone fynbos vegetation include alien plants (considered the biggest problem), land clearing, resort development, inappropriate fire management and over-harvesting of flowers (heydenrych 1994). thicket encroachment into the adjacent limestone fynbos and renosterveld is therefore a serious threat to the species-rich fynbos with many endemic and red data species. the species richness of coastal thicket in the reserve is very low compared to those of the adjacent limestone fynbos and renosterveld communities (zietsman & bredenkamp in press) and the true thicket species like rhus glauca and euclea racemosa are neither rare nor endemic. regarding the importance of the conservation of limestone fynbos in areas where the thicket communities occur, the effective management of the area becomes all the more important, and fully justifies a detailed vegetation study of the coastal thicket. study area the reserve is situated in the bredasdorp district, western cape, between struisbaai north in the south and de mond state forest in the north. it is bordered in the west by the bredasdorp/struisbaai road, and in the south by the coastline. the study area is situated close to the coastal town of struisbaai. the andrew’s field (129 ha) and tsaba-tsaba nature reserve is approximately 979 ha in extent. approximately one third of the study area consists of the coastal vegetation, which is dealt with in this report. the remainder of the study area consists of inland plains and hills, described by zietsman & bredenkamp (in press). the dominant substrate in the study area is the bredasdorp group (malan et al. 1994). two formations of the bredasdorp group namely strandveld and waenhuiskrans formations are found in the study area (malan et al. 1994). beach and terrace deposits, not formally named as a formation, but forming part of the bredasdorp group, are found in the form of roll-stones on the coastal beach (malan et al. 1994). previous changes in sea level resulted in the presence of roll-stones of table mountain sandstone. the strandveld formation is restricted to the coast. it consists of white to light-grey dune sands with a high percentage shell fragments. partial cementing of sands is a result of the high calcium carbonate content in the sand. the lithology of this formation is described as white dune sand, strand sand with finely divided shell and alluvial stones (malan et al. 1994). the waenhuiskrans formation forms outcrops adjacent to the current coastline. the waenhuiskrans stratiotype is 12.4 m deep, is locally overlain with 1 m thick calcretes and consists of medium-grained cross-layered calcarenite with well-rounded quartz and a few glauconite grains. large-scale aeolic cross-layers are characteristic of the unit. the lithology of this formation is described as partially calcified dune sand (malan et al. 1994). the study area is situated on the agulhas plain, a coastal lowland. the majority of the area falls below the 10 m contour (jefferey 1996). coastal dunes and dune plains are typical in this area, with the highest point on a ridge 31 m above sea level. the soil can be described as shallow sandy soil, overlying limestone, or shallow to deep sandy soil, overlying clay, silt and gravel. four different soil forms have been distinguished in the area (macvicar 1991): coega: orthic a on hard bank carbonate horizon; family marydale: lime containing a-horizon. this soil form is found on the limestone hills and shallow-soil limestone plain. immerpan: melanic a on hard bank carbonate horizon; family kalkpan: lime containing a-horizon. this soil form is found on the proteoid dominated limestone plain koedoe 49/1 (2006) 34 issn 0075-6458 zietsman.qxd 2006/04/17 08:48 pm page 34 where the soil is deep. brandvlei: orthic a on soft carbonate horizon; family kolke: signs of wetness in carbonate horizon. the marsh area is characterised by this soil form. namib: orthic a on regic sand; family beachwood: containing lime within 1500 mm from soil surface. this soil form is found on the dune plain and the deep-sand plains. according to tinley (1985) two distinct sand characteristics occur in dune fields. bare dune sands, yellow in colour, absorbs all rain and consequently there is no surface runoff. grey sands, stained by humus and covered with woody vegetation, have a water repellent layer near the surface, beneath the litter, causing a massive surface runoff when heavy rains follow a dry period (tinley 1985). the main factors affecting the climate are the contrasting sea surface temperatures of the two major ocean currents and the inshore circulation (tinley 1985). according to the geographical division of heydorn & tinley (1980), the reserve is situated on the south coast, which is a transitional zone between tropical and temperate waters. the study area is situated in the transitional zone between the winter-rainfall region in the west and the non-seasonal rainfall region in the east (mustart et al. 1997). the south coast has a warm temperate climate with allseasons and bimodal equinoctial rainfall (strydom 1992). the average annual precipitation of the area is 444 mm. precipitation is mostly rain, but it also occurs in the form of fog. the maximum precipitation is in june, and the minimum during february and december. according to heydorn & tinley (1980) the study area is situated in a low rainfall region, within an arid belt protruding into the interior from the coast. the low rainfall appears to be the result of cold inshore waters that inhibit shoreline rains. a walter climatic diagram was compiled (fig. 1) from data obtained from the agulhas weather station. the average annual maximum and minimum temperatures for the area are 6 ºc and 13.3 ºc respectively. the maximum temperature for the area was 36.1 ºc, obtained in february. the minimum temperature for the area is 3.9 ºc, obtained in june (zietsman & bredenkamp in press). the wind along the south coast is bidirectional. southeasterly winds alternate with issn 0075-6458 35 koedoe 49/1 (2006) fig. 1. a walter climatic diagram for andrew’s field and tsaba-tsaba nature reserve. a = altitude, b = mean annual temperature, c = mean annual rainfall, d = mean daily minimum (coldest month), e = mean daily maximum (hottest month) zietsman.qxd 2006/04/17 08:48 pm page 35 northwesterly and southwesterly winds (heydorn & tinley 1980). methods relevés were compiled in 74 stratified sample plots, placed in relatively homogeneous areas, representative of particular plant communities that occur in a mosaic distribution pattern. a plot size of 10 m² was used, and considered as large enough to ensure that all species of regular occurrence in the stand are present in the sample plot. sample plots were placed in such a way as to ensure that each plot adequately represents the structure of the particular vegetation (werger 1974). a list of plant species found in each sample plot was compiled. the cover-abundance of each species in the sample plot was assessed, using the braun-blanquet cover-abundance scale (werger 1974). cover of the height classes of the different strata (edwards 1983) was estimated and the vegetation was structurally classified according to the edwards (1983) structural classification system. the following habitat characteristics were recorded in each sample plot: altitude, topographical position, exposure to wind (seaward or towards the land), slope angle, slope direction, geology, soil, percentage rock cover and biotic influence. six main different habitat types are found in the study area, namely dune plain, renosterveld plain, limestone plain, limestone hills, dunes and beach. these were recorded in each sample plot. to obtain a first approximation of the plant communities of the area, relevés were classified using twinspan (hill 1979), and these results were then refined by application of the classical braunblanquet methodology (behr & bredenkamp 1988). results classification the results obtained from the classification are presented in a phytosociological table (table 1). in naming the plant communities, diagnostic and/or dominant species were used in combination with the edwards (1983) structural classification. 1. rhus glauca euclea racemosa low to tall closed thicket community 1.1 rhus lucida euclea racemosa low to short closed thicket sub-community 1.2 pterocelastrus tricuspidatus euclea racemosa short to tall closed thicket sub-community 1.2.1 olea exasperata euclea racemosa short to high closed thicket variant 1.2.2 carissa bispinosa euclea racemosa short to tall closed thicket variant 1.3 acmadenia obtusata euclea racemosa short to tall closed thicket subcommunity 1.4 helichrysum dasyanthum euclea racemosa low to short closed thicket sub-community 1.5 acacia cyclops euclea racemosa high to tall closed thicket sub-community 1.6 thamnocortus insignis euclea racemosa high to tall closed thicket subcommunity 1.6.1 elytropappus rhinocerotus euclea racemosa short to tall closed thicket variant 1.6.2 leucadendron coniferum euclea racemosa high to tall closed thicket variant 2. chrysanthemoides monilifera solanum africanum low closed dune shrub community 3. chrysanthemoides monilifera ehrharta villosa var. maxima low to high closed dune shrub community 3.1 chrysanthemoides monilifera rhus crenata short to high dune shrub subcommunity 3.2 chrysanthemoides monilifera morella cordifolia low to short closed shrub sub-community 4. ehrharta villosa var. maxima low to short closed dune grassland community 5. ammophila arenaria low to short closed dune grassland community 6. arcthotheca populifolia thinopyrum distichum low to short open beach community the cover and height of various strata of the main plant communities are given in table 2. the hierarchical classification and associated environmental interpretation of the plant communities is given in fig. 2. the community numbers in fig. 2 correspond with the plant community numbers used in the descriptions in the text. 1. rhus glauca euclea racemosa low to tall closed thicket community the community is found close to the sea on the dune plains, limestone plains, renosterkoedoe 49/1 (2006) 36 issn 0075-6458 zietsman.qxd 2006/04/17 08:48 pm page 36 issn 0075-6458 37 koedoe 49/1 (2006) dune plains limestone plains rhenoster plains dune slopes slopes and peaks of limestone hills flat dune plain sandy shallow limestone plain dune plains deep to flat limestone plains sandy footslopes, slopes and shoulders of limestone hills sheltered, steep north-facing slopes of back-dune deep limestone plains peaks, foot-slopes of limestone hills flat limestone plains peaks and shoulders of limesone hills rhenosterveld plain limestone hills foot-slopes dune slopes sheltered from south-eastern wind dune slopes parallel to south-eastern wind dune plain deep sandy slopes and shoulders of limestone hills limestone plains acacia cyclops encroachment in community 1 rhenosterveld plains limestone hills, footslopes very steep dry, warm dune slopes coastal dunes and dune troughs south-facing slopes, sparsely vegetated dunes northern steep dry mid-slopes of dunes beach and gravel plain frontal dune slopes 1.1 1.2.1 1.2.2 1.3 1.4 1.5 1.6.1 1.6.2 2 3.1 3.2 4 5 6 fig. 2. hierarchical classification and associated environmental characteristics of the coastal thicket, dunes and strand of andrew’s field and tsaba-tsaba nature reserve. zietsman.qxd 2006/04/17 08:48 pm page 37 ta bl e 1 p hy to so ci ol og ic al ta bl e of c oa st al th ic ke t, du ne s an d st ra nd o f a nd re w 's f ie ld a nd t sa ba -t sa ba n at ur e r es er ve zietsman.qxd 2006/04/17 08:48 pm page 38 ta bl e 1 (c on tin ue rd ) zietsman.qxd 2006/04/17 08:48 pm page 39 ta bl e 1 (c on tin ue rd ) zietsman.qxd 2006/04/17 08:48 pm page 40 issn 0075-6458 41 koedoe 49/1 (2006) veld plains, northern slope of back-dunes, and on the slopes and peaks of limestone hills. these areas have a low cover of small irregular white-grey limestone pebbles. the rhus glauca euclea racemosa short to tall closed thicket community consists of scattered, dense groups of broad-leaved shrubs. these groups have high shrubs near the centre and lower shrubs near the periphery. the structure of the vegetation is given in table 2. the shrub layer is most prominent, while restioid and grass layers cover 10 % or less. forbs and sedges have a very low cover. the shrubs euclea racemosa, rhus glauca and rhus laevigata (species group a, table 1) are diagnostic and dominant in this community. chrysanthemoides monilifera and metalasia muricata (species group q) are also prominent in the community. in the absence of fire, these thickets tend to encroach into adjacent threatened limestone fynbos and rhenosterveld communities which have several endemic and red data species (cowling & richardson 1995; heydenrych 1994; zietsman & bredenkamp in press). six sub-communities, two of which have two variants, were identified: 1.1 rhus lucida euclea racemosa low to short closed thicket sub-community this sub-community is found on the flat to undulating dune plain, namib soil form and limestone plain, brandvlei and coega soil forms (fig. 2). the vegetation forms scattered groups of low to short broad-leaved shrubs. rhus lucida (species group b, table 1) is the only diagnostic species for this sub-community. the shrubs euclea racemosa, rhus glauca and rhus laevigata var. laevigata for. coangoa (species group a, table 1) are the dominants in this sub-community. the shrubby helichrysum dasyanthum (species group g) and passerina paleacea (species group i) are locally prominent in the subcommunity. an average of only four species per relevé was recorded in this sub-community. 1.2 pterocelastrus tricuspidatus euclea racemosa short to tall closed thicket sub-community the sub-community is found on the namib soil form on the dune plain and on the immerpan and coega soil forms on sandy foot-slopes and shoulders of limestone hills (fig. 2). table 2 percentage cover (c ) (%) and height (h)(m) of the strata of the main plant communities community 1 2 3 4 5 6 stratum c h c h c h c h c h c h tall shrub 14 2.5 6 2.5 6 2.5 high shrub 20 1.5 5 1.5 10 1.5 10 1 short shrub 28 0.75 5 0.75 29 0.75 low shrub 15 0.4 51 0.15 16 0.3 forb 3 0.3 1 0.3 2 0.3 10 0.5 short restoid 4 0.75 0.3 0.2 low restoid 7.5 0.4 sedge 1 0.15 0.5 0.15 high grass 0.5 1.5 3.5 1.5 short grass 0.5 0.5 0.6 0.75 2 0.75 25 0.75 44 0.7 5 0.5 low grass 10 0.3 2 0.3 15 0.3 2 0.3 zietsman.qxd 2006/04/17 08:48 pm page 41 koedoe 49/1 (2006) 42 issn 0075-6458 the vegetation forms scattered groups of short to tall broad-leaved shrubs, with some lower shrubs at the periphery. species group c (table 1) is diagnostic for this sub-community, with the shrubby trees robsonodendron maritimum and pterocelastrus tricuspidatus dominant diagnostic species. the shrubs euclea racemosa, rhus glauca and rhus laevigata var. laevigata for. coangoa (species group a), are also very prominent in the sub-community. 1.2.1 olea exasperata euclea racemosa short to high closed thicket variant the variant occurs on the slopes and peaks of the limestone hills, and on the sandy, deep-soil limestone plains (fig. 2). species group d (table 1) is diagnostic, with olea exasperata and osyris compressa prominent diagnostic species. the diagnostic species of the community and sub-community, namely the shrubs euclea racemosa, rhus glauca and rhus laevigata var. laevigata for. coangoa (species group a), robsonodendron maritimum and pterocelastrus tricuspidatus (species group c) are dominant in the variant. an average of 12 species per relevé was recorded for this variant. 1.2.2. carissa bispinosa short to tall closed thicket variant this variant is restricted to the flat limestone plains and the shoulders and peaks of limestone hills (fig. 2). this soil is of the immerpan and coega forms. species group e (table 1) is diagnostic with the shrubs carissa bispinosa and sideroxylon inerme the most prominent diagnostic species. the shrubs euclea racemosa, rhus glauca (species group a), robsonodendron maritimum and pterocelastrus tricuspidatus (species group c) are dominant whereas the alien shrub acacia cyclops (species group h) is also locally prominent. an average of 13 species per relevé was recorded for this variant. 1.3 acmadenia obtusata euclea racemosa short to tall closed thicket sub-community the sub-community is restricted to a small patch close to the sea, in the central part of the reserve, on a steep (26° to 45°), sheltered northern slope of a back-dune. the soil is of the namib form and no rocks are found here. only a single relevé represents this sub-community. the vegetation forms scattered groups of short to tall closed broad-leaved shrubs, with some lower shrubs at the periphery. species group f (table 1) is diagnostic, with the shrub acmadenia obtusata, the forbs arctopus echinatus and zaluzianskya villosa and the grass festuca scabra, as diagnostic species. the shrubs euclea racemosa (species group a), otholobium bracteolatum (species group g), rhus crenata (species group n) and passerina rigida (species group p) are dominant. eighteen species were recoded in the single releve. 1.4 helichrysum dasyanthum euclea racemosa low to short closed thicket sub-community this sub-community is widespread on the dune plain, namib soil form, the sandy shallow-soil limestone plain and sandy slopes of limestone hills, immerpan and coega soil forms (fig. 2). the vegetation forms scattered groups of low to short broad-leaved shrubs, with some high shrubs at the centre. otholobium bracteolatum and helichrysum dasyanthum (species group g, table 1) are the diagnostic species for this sub-community. the shrubs euclea racemosa, and rhus laevigata var. laevigata for. coangoa (species group a) are dominant, while rhus glauca (species group a), pterocelastrus tricuspidatus (species group c), passerina paleacea (species group i), chrysanthezietsman.qxd 2006/04/17 08:48 pm page 42 issn 0075-6458 43 koedoe 49/1 (2006) moides monilifera (species group q) are also locally prominent. the sub-community contains an average of only six species per relevé. 1.5 acacia cyclops euclea racemosa high to tall closed thicket sub-community locally the alien acacia cyclops encroached into the rhus glauca euclea racemosa short to tall closed thicket community. acacia cyclops, the shrubby tree protea obtusifolia, and the shrub passerina galpinii are the diagostic species for this sub-community (species group h, table 1). acacia cyclops and the shrubs euclea racemosa, rhus glauca and rhus laevigata var. laevigata for. coangoa (species group a), and metalasia muricata (species group q) are dominant in the sub-community. the sub-community contains an average of seven species per relevé. 1.6 thamnochortus insignis euclea racemosa high totall closed thicket sub-community the sub-community is found on the renosterveld plain, brandvlei soil form and on sandy foot-slopes of limestone hills, namib soil form (fig. 2). the vegetation forms scattered groups of high to tall broad-leaved shrubs, with some low and short shrubs at the periphery. low forbs and some high, low and short restioids are also present in the sub-community. species group i (table 1) is diagnostic for this sub-community with the restioid thamnochortus insignis and the shrubs helichrysum patulum, passerina paleacea, metalasia densa the most prominent diagnostic species. the shrubs euclea racemosa, rhus glauca (species group a), chrysanthemoides monilifera and metalasia muricata (species group q), are also prominent in the subcommunity. two variants are recognised within this subcommunity: 1.6.1 elytropappus rhinocerotus short to tall closed thicket variant the variant is found at an altitude of 3 m on the renosterveld plain, brandvlei soil form (fig. 2). the vegetation scattered groups of short to tall broad-leaved shrubs, with some lower shrubs at the periphery, as well as some high restioids. the diagnostic species are listed in species group k (table 1), which includes the shrubs elytropappus rhinocerotis, leucadendron linifolium, oedera uniflora, lycium afrum, rhus laevigata var. villosa, asparagus lignosus, euryops hebecarpus, and the forbs limonium scabrum var. scabrum, hermannia althaeifolia and pelargonium myrrhifolium. the shrubs euclea racemosa and rhus glauca (species group a) and the restioid thamnochortus insignis (species group i) are dominant whereas the shrubs helichrysum patulum, passerina paleacea (species group i), chrysanthemoides monilifer and metalasia muricata (species group q) are locally prominent. the variant has an average of 22 species per relevé. 1.6.2 leucadendron coniferum high to tall closed thicket variant the variant is found on the sandy foot-slopes of limestone hills, namib soil form (fig. 2). the vegetation forms scattered groups of high to tall closed broad-leaved shrubs, with some low to short shrubs at the periphery, as well as some high and low restioids. a large number of species given in species group k (table 1) are diagnostic for this variant, the most prominent being the shrubs eriocephalus kingesii, leucadendron coniferum, protea susannae, maytenus procumbens, agathosma serpyllacea, agathosma collina, phylica stipularis, trigogyne repens and tephrosia capensis and the restioid chondopetalum microcarpum. the shrubs euclea racemosa and rhus glauca (species group a), and the restioid thamnochortus zietsman.qxd 2006/04/17 08:48 pm page 43 koedoe 49/1 (2006) 44 issn 0075-6458 insignis (species group i) are dominant, while the shrubs olea exasperata (species group d), cassine peragua (species group i), solanum africanum (species group l), chrysanthemoides monilifera (species group q) are locally prominent. the variant is rich in species and has an average of 43 species per relevé. 2. chrysanthemoides monilifera solanum africanum low closed shrub dune community the sparsely vegetated community is found in the shifting sand dune area adjacent to the sea. it occurs in a mosaic distribution pattern with communities 3, 4 and 5. the community is situated at an altitude of 1–20 m, on steep gradients. this vegetation is restricted to the drier northerly mid-slopes of coastal dunes, facing inland (fig. 2). the soil is of the namib form and no rocks are present. the vegetation consists of low prostrate shrubs, with a few short, high and tall shrubs also present. the low shrub stratum is prominent with an average cover of 51 % (table 2). the low shrub solanum africanum (species group l, table 1) is not only diagnostic but also dominant in the community. the shrub chrysanthemoides monilifera (species group q) is also locally prominent. the community contains an average of only four species per relevé. 3. chrysanthemoides monilifera ehrharta villosa var. maxima low to high closed dune shrub community the community is found in the sparsely vegetated shifting dune area adjacent to the sea, in a mosaic distribution pattern with communities 2, 4 and 5. it is also present in a mosaic distribution pattern with communities 2 and 4 in the more densely vegetated sand dune area at the seashore and directly behind to the shifting sand dune area (fig. 2). no rocks are present in this habitat and the soil is of the namib form. the vegetation consists of short and low shrubs, with some short, high and tall shrubs as well as some low forbs, and low and short grasses present (table 2). the shrub stoebe cinerea and the grass ehrharta villosa var. maxima (species group m, table 1), are diagnostic for the community and chrysanthemoides monilifera (species group q) is the dominant species. the shrubs solanum africanum (species group l), stoebe cinerea (species group m), morella cordifolia (species group o), passerina rigida (species group p) and metalasia muricata (species group q) are locally prominent species. the community has an average of ten species per relevé. 3.1 chrysanthemoides monilifera rhus crenata short to high dune shrub sub-community the sub-community is found on north-facing coastal dune slopes with steep gradients of 26°–45°, sheltered from the south-eastern wind carrying salt spray. no rocks are found here (fig. 2). the vegetation consists of short to high shrubs, with a few tall shrubs. low forbs, low restios, low sedges, and low, short and high grasses have low cover values. the shrub rhus crenata (species group n, table 1) is the only diagnostic species. passerina rigida (species group p), chrysanthemoides monilifera and metalasia muricata (species group q) are dominant whereas the shrubs solanum africanum (species group l), stoebe cinerea (species group m), morella cordifolia (species group o), and the grass ehrharta villosa var. maxima (species group m) are also locally prominent in the sub-community. the sub-community contains an average of 12 species per relevé. 3.2 chrysanthemoides monilifera morella cordifolia low to short closed dune shrub sub-community the steep (16–>45°) southerly-facing midslopes dunes where this sub-community occurs are larger, more exposed areas than zietsman.qxd 2006/04/17 08:48 pm page 44 issn 0075-6458 45 koedoe 49/1 (2006) those where sub-community 3.1 occurs. the vegetation is exposed to sea mist (fig. 2). the leaf structure and orientation of the most dominant plant species, morella cordifolia, enhances the ability of the plant to catch moisture from sea fog. the vegetation consists of low to short shrubs, with some high and tall shrubs, as well as some low forbs, low restios, low sedges, and low, short and high grass. the shrubs morella cordifolia, lessertia frutescens (species group o, table 1), are dominant diagnostic species, and chrysanthemoides monilifera (species group q is also dominant. the shrubs stoebe cinerea (species group m), passerina rigida (species group p) and metalasia muricata (species group q), and the grass ehrharta villosa var. maxima (species group m) are locally prominent. the sub-community contains an average of 11 species per relevé. 4. ehrharta villosa var. maxima low to short closed dune grassland community the community is found on steep, exposed southern slopes of sparsely vegetated stable sand dunes, or the more mobile shifting sand dunes adjacent to the sea. (fig. 2). no rocks are present in the namib soil form. the community is found in a mosaic distribution pattern with communities 2, 3 and 5 and also in a mosaic distribution pattern with community 6 in the coastal beach area, adjacent to the sea. the vegetation is low to short dune grassland, with a few high grasses. (table 2). the grass ehrharta villosa var. maxima (species group m, table ) is the sole diagnostic and dominant species in the community. no other species occur here. 5. ammophila arenaria low to short closed dune grassland community the community occurs on the steep, dry and warm north-facing slopes of sparsely vegetated shifting sand dunes, in a mosaic distribution pattern with communities 2, 3 and 4 (fig. 2). the vegetation is low to short dune grassland. the shrub stratum has an average height of 1 m and an average canopy cover of 10 % and the grass stratum has an average height of 0.7 m and an average canopy cover of 44 % (table 2). the exotic grass ammophila arenaria (species group r, table 1) is the diagnostic and dominant species in the community. other species present are lessertia frutescens and senecio arenarius. the community contains an average of only three species per relevé. 6. arctotheca populifolia thinopyrum distichum short to low open beach community the community is situated on the beach and south-facing slopes of low fore dunes, directly adjacent to the sea, and also on gravel plains behind the low fore dunes (fig. 2). there are no rocks on the beach and dunes and 30–50 % rock cover on the gravel plain. the rocks on the gravel plain are mostly sandstone and quartzite roll-stones, deposited by the sea while the lea level was higher. the vegetation consists of short to low open beach grassveld with low forbs (table 2). the forb arctotheca populifolia and the grass thinopyrum distichum (species group s, table 2) are diagnostic and are the only species found in this community. the community contains an average of only two species per relevé. discussion the use of twinspan and the application of braun-blanquet procedures for refinement, were successful to classify the plant communities of this vegetation type. six plant communities, with eight sub-communities and four variants, were identified. the plant communities could all be related to specific environmental conditions and are therefore zietsman.qxd 2006/04/17 08:48 pm page 45 koedoe 49/1 (2006) 46 issn 0075-6458 floristically and ecologically distinguishable and interpretable. the thicket community (community 1) forms part of the dune thicket vegetation type of the thicket biome (low & rebelo 1998), whereas the other communities form part of the fynbos biome (low & rebelo). the thicket was mostly found in a mosaic with the vegetation of the inland plains and hills (zietsman & bredenkamp in press). the thicket community does not only differ from the fynbos communities concerning species composition and structure, but also in conservation importance. according to low & rebelo (1998) 14.49 % of dune thicket is currently being conserved, as opposed to the 1.42 % of south and southwest coast renosterveld and the 13.84 % of limestone fynbos (low & rebelo 1998). according to cowling & richardson (1995), if fire could be excluded from fynbos for a century or two, many of the landscapes would become densely populated with just a few species of thicket shrubs and trees. inland thicket development is therefore a feature of vegetation that has not been subjected to fire for some time. the species richness of this thicket is very low compared to those of fynbos and renosterveld (zietsman & bredenkamp in press) and the true thicket species like rhus glauca and euclea racemosa, are neither rare nor endemic. the limestone communities and the renosterveld community, within which the thicket patches are found, have high conservation priority. renosterveld has a very high conservation priority due to the small portion left, mainly because of agricultural land clearing (low & rebelo 1998; zietsman & bredenkamp in press). limestone fynbos is one of the most threatened vegetation types in the cape floristic region (hilton-taylor & le roux 1989). there are 110 plant species endemic to limestone outcrops. limestone fynbos also contains many species that are classified as rare (heydenrych 1994). thicket forming should be seen as a threat to the natural fynbos and renosterveld vegetation, and should not be encouraged above the conservation of the other fynbos and renosterveld vegetation. the presence of sub-community 1.5 (acacia cyclops euclea racemosa high totall closed thicket sub-community) is a reason for concern. acacia cyclops occurs scattered in various communities, fairly well distributed throughout andrew’s field and tsabatsaba nature reserve, but the larger stands found locally represent sub-community 1.5. all areas containing acacia cyclops, should be managed by implementing regular prescribed burning, and alien-invasive plant control, ideally by means of the cut-and-burn method (cowling 1992), but the current woodcutting method (promoting job provision), can also be followed. the conservation of limestone and renosterveld vegetation should be treated as a high priority. fire should be kept out of the dune area, because, according to tinley (1985) fire might cause an increase in dune instability. fire should also be kept out of areas that are heavily infested with the cape dune molerat. according to tinley (1985) fire would lead to an undesired increase in burrowing by cape dune molerats to obtain sufficient food (roots). therefore, as extensive dune molerat burrowing is already a problem in some areas in the reserve, fire should be kept out of those areas. other areas should be burned using a 15-year block-burn cycle to avoid the possible substitution of limestone and renosterveld vegetation by dense thickets. a single red data species (solanum africanum) is found quite abundantly in the dune area (communities 2 and 3). the presence of limestone endemic species in the dune area can be explained by the fact that both the dune sands and the limestone soil are alkaline. the shifting sand dune area should be protected from disturbances, like trampling and vehicle tracks. protection from disturbances should also provide sufficient protection to the red data and limestone endemic species. community 9 is dominated by ammophila arenaria, an exotic grass which is an european weed, commonly found on coastal dunes (goldblatt & manning 2000). although community 6 contains no rare or endemic species, the vegetation unit should zietsman.qxd 2006/04/17 08:48 pm page 46 issn 0075-6458 47 koedoe 49/1 (2006) be protected due to its uniqueness, and due to the fact that this gravel-plain is the breeding site of the rare damara tern sterna balaenarum (jeffery 1996). andrew’s field and tsaba-tsaba nature reserve is an area of great conservation significance. this area comprises an important part of the natural heritage found at the southern tip of africa, and should be protected and conserved for future generations. the resulting classification provides a useful tool, not only for the management of the plant communities of andrew’s field and tsaba-tsaba nature reserve, but also for similar vegetation areas, found in the surrounding region. acknowledgements mrs. f. siebert is thanked for her assistance during data processing. references behr, c.m. & g.j. bredenkamp. 1988. a phytosociological classification of thevegetation of the witwatersrand national botanic garden. south african journal of botany 54(6): 525–533. bredenkamp, g.j. & l.r. brown. 2001. vegetation – a reliable ecological basis for environmental planning. urban greenfile nov-dec 2001: 38–39. cowling, r.m. (ed.). 1992. the ecology of fynbos: nutrients, fire and diversity. cape town: oxford university press. cowling, r.m. & d. richardson. 1995. fynbos: south africa’s unique floral kingdom. cape town: fernwood press. edwards, d. 1972. botanical survey and agriculture. proceedings of the grassland society of southern africa 7: 15–17. edwards, d. 1983. a broad-scale structural classification of vegetation for practical purposes. bothalia 14: 705–712. goldblatt, p. & j. manning. 2000. cape plants – a conspectus of the cape flora of south africa. pretoria: national botanical institute of south africa. heydenrych, b.j. 1994. limestone fynbos – unique but threatened. an investigation into the conservation status of limestone fynbos in the southern cape. kirstenbosch: botanical society of south africa. (flora conservation committee report no. 94/1.) hill, m.o. 1979. twinspan: a fortran program for arranging multivariate data in an ordered two-way table by classification of the individuals and attributes. new york: cornell university. hilton-taylor, c. & a. le roux. 1989. conservation status of the fynbos and karoo biomes. pp. 202–233. in: huntley, b.j. (ed.). biotic diversity in southern africa: concepts and conservation. cape town: oxford university press. heydorn, a.e.f. & k.l. tinley. 1980. estuaries of the cape, part 1: synopsis of the cape coast natural features, dynamics & utilization. stellenbosch: council for scientific & industrial research, national research institute for oceanology. (council for scientific & industrial research research report; no. 380.) jeffery, d. 1996. an abbreviation of the botanical sensitivity analysis report for portion 7 and remainder portion 8 of the farm zoetendalsvlei no. 280 of july 1995. environmental consulting and facilitation services, klapmuts. low, a.b. & a.g. rebelo. 1998. vegetation of south africa, lesotho and swaziland: a companion to the vegetation map of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. macvicar, c.n. 1991. grondklassifikasie ‘n taksonomiese sisteem vir suid-africa. pretoria: departement van landbou-ontwikkeling, malan, j.a., j.h.a. viljoen, h.p. siegfried & h. de v. wickens. 1994. die geologie van die gebied riversdale. pretoria: staatsdrukker. mustard, p., r. cowling & j. albertyn. 1997. southern overberg: south african wildflower guide 8. cape town: national book printers. scheepers, j.c. 1983. vegetation studies in south africa. bothalia 14: 683–690. strydom, s. 1992. a classification of southern cape coastal dunes and the associated vegetation. msc: thesis, university of port elizabeth, port elizabeth. tinley, k.l. 1985. coastal dunes of south africa. pretoria: council for scientific and industrial research, national scientific programmes unit. (south african national scientific programmes report; no. 109.) werger, m.j.a. 1974. on concepts and techniques applied in the zürich-montpellier method of vegetation survey. bothalia 11: 309–323. zietsman, m.m. & g.j. bredenkamp (in press). threatened limestone fynbos plant communities of andrew’s field and tsaba-tsaba nature reserve. bothalia. zietsman.qxd 2006/04/17 08:48 pm page 47 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <feff004f007000740069006f006e007300200070006f0075007200200063007200e900650072002000640065007300200064006f00630075006d0065006e00740073002000500044004600200064006f007400e900730020006400270075006e00650020007200e90073006f006c007500740069006f006e002000e9006c0065007600e9006500200070006f0075007200200075006e00650020007100750061006c0069007400e90020006400270069006d007000720065007300730069006f006e00200070007200e9007000720065007300730065002e0020005500740069006c006900730065007a0020004100630072006f0062006100740020006f00750020005200650061006400650072002c002000760065007200730069006f006e00200035002e00300020006f007500200075006c007400e9007200690065007500720065002c00200070006f007500720020006c006500730020006f00750076007200690072002e0020004c00270069006e0063006f00720070006f0072006100740069006f006e002000640065007300200070006f006c0069006300650073002000650073007400200072006500710075006900730065002e> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice article information authors: jan a. venter1 werner conradie2,3 affiliations: 1school of natural resource management, nelson mandela metropolitan university, george campus, south africa 2port elizabeth museum (bayworld), port elizabeth, south africa 3south african institute for aquatic biodiversity, grahamstown, south africa correspondence to: jan venter email: jan.venter@nmmu.ac.za postal address: private bag x6531, george 6530, south africa dates: received: 24 nov. 2014 accepted: 27 may 2015 published: 17 nov. 2015 how to cite this article: venter, j.a. & conradie, w., 2015, ‘a checklist of the reptiles and amphibians found in protected areas along the south african wild coast, with notes on conservation implications’, koedoe 57(1), art. #1247, 25 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1247 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. a checklist of the reptiles and amphibians found in protected areas along the south african wild coast, with notes on conservation implications in this checklist... open access • abstract • introduction • study areas    • cape morgan nature reserve    • manubi state forest    • dwesa nature reserve    • hluleka nature reserve    • silaka nature reserve    • mkambati nature reserve • material and methods • results and discussion • conclusion • acknowledgements    • competing interests    • authors’ contributions • references • appendix 1    • species accounts    • amphibia    • arthroleptidae    • arthroleptis wahlbergii smith, 1849 (bush squeaker)    • brevicipitidae    • breviceps cf. bagginsi minter, 2003 (bilbo's rain frog)    • hyperoliidae    • afrixalus spinifrons cope, 1862 (natal leaf-folding frog)    • hyperolius poweri loveridge, 1938    • pyxicephalidae    • cacosternum boettgeri boulenger, 1882 (boettger's caco)    • natalobatrachus bonebergi hewitt & methuen, 1912 (kloof frog)    • reptilia    • crocodylidae    • crocodylus niloticus laurenti, 1768 (nile crocodile)    • gekkonidae    • afroedura pondolia hewitt, 1925 (pondo flat gecko)    • lygodactylus capensis smith, 1848 (common dwarf gecko)    • lacertidae    • tropidosaura montana natalensis fitzsimons, 1947 (common mountain lizard)    • gerrhosauridae    • gerrhosaurus flavigularis wiegmann, 1828 (yellow-throated plated lizard)    • tetradactylus cf. fitzsimonsi (fitzsimons’ long-tailed seps)    • scincidae    • acontias plumbeus bianconii, 1849 (giant legless skin)    • trachylepis homalocephala wiegmann, 1828 (red-sided skink)    • chamaeleonidae    • bradypodion caffer boettger, 1889 (pondo dwarf chameleon)    • bradypodion kentanicum hewitt, 1935 (kentani dwarf chameleon)    • leptotyphlopidae    • leptotyphlops sylvicolus broadley & wallach, 1997 (forest thread snake)    • pythonidae    • python natalensis smith, 1840 (southern african python)    • lamprophiidae    • amblyodipsas concolor smith, 1849 (kwazulu-natal purple-glossed snake)    • macrelaps microlepidotus günther, 1863 (kwazulu-natal black snake)    • psammophis brevirostris peters, 1881 (short-snouted grass snake)    • elapidae    • dendroaspis polylepis günther, 1864 (black mamba)    • colubridae    • thelotornis capensis capensis smith, 1849 (south-eastern savannah vine-snake) • references for appendix 1 • appendix 2    • list of all amphibian material from the port elizabeth museum examined for the study    • amphibians    • reptiles • appendix 3    • additional records used in compiling the final checklist of amphibians and reptiles of the wild coast reserves • amphibians    • reptiles • appendix 4 abstract top ↑ we surveyed six protected areas along the wild coast of the eastern cape, south africa, to determine general herpetofaunal diversity as well as the representation of species of special conservation concern. visual encounter survey methods and standard y-shape trap arrays were used to conduct surveys from 2011 to 2013. a total of 59 species (22 amphibians and 37 reptiles) were recorded. a number of previously unknown populations of threatened species and one potential novel species were discovered in these protected areas, and the known ranges of several other species were extended. a total of 243 quarter-degree grid-cell unit records were documented, of which 90 (23% amphibians and 50% reptiles) represented the first records for the area. amphibian and reptile diversity increased along the coast and a number of species of conservation concern were well represented in current protected areas. our study provides a comprehensive amphibian and reptile checklist for an under-sampled region and highlights the importance of baseline data for improving conservation management. conservation implications: small protected areas play an important role in conserving a number of threatened herpetofaunal species along the wild coast. the region is currently under significant and increasing pressure from anthropogenic-induced environmental degradation, which affects biodiversity and subsequently the local inhabitants. the information presented here represents an important baseline for future conservation management. introduction top ↑ amphibians and reptiles are declining globally owing to habitat loss and degradation, pressure from alien invasive species, environmental pollution, disease, unsustainable use and global climate change (böhm et al. 2013). protected areas are often useful tools for the protection of herpetofauna and contribute to the conservation of many species (ochoa-ochoa et al. 2009; tuberville et al. 2005). management actions within protected areas may, however, have a significant impact on herpetofaunal ecology in a number of ways (measey, armstrong & hanekom 2009; russell, lear & guynn jr. 1999). it is therefore critical that protected area managers consider herpetofaunal diversity, abundance and distributions within a local and regional context in conservation management. southern africa, the geographical region south of the cunene and zambezi rivers, has the richest reptile diversity in africa (branch 2006). within this area, south africa (including lesotho and swaziland) has the highest reptile diversity (384 indigenous species) (branch 2014). the wild coast, in the former transkei (‘transkei’ refers to a former homeland in pre-democratic south africa), is regarded as having very low reptile diversity compared with southern and north-eastern areas of south africa (branch 2014). although the pondoland region of the wild coast (eastern coastal region of the eastern cape, south africa) is regarded as a centre of high plant diversity and endemism (van wyk & smith 2001), the terrestrial faunal species of the entire wild coast region (coastal transkei and pondoland combined), in many cases, remain poorly represented compared with other areas in southern africa (bates et al. 2014; dippenaar, hamer & haddad 2011; minter et al. 2004; mokhatla et al. 2012; perera, ratnayake-perera & proches 2011; skead 2007). most herpetological studies from this area date back to the early 1900s (fitzsimons 1930; power 1935), with only a few recent ad hoc surveys added (branch & haagner 1999; burger 1996; haagner 1994). this could possibly be attributed to the remoteness of the area and unfavourable road conditions, as well as perceived safety risks, which made the logistics of field sampling challenging and, in turn, may have deterred herpetologists from working in the area. some authors go as far as to describe the phenomenon as the ‘transkei faunal distribution gap’; that is a real gap in species distributions caused by environmental features (botts, erasmus & alexander 2011; perera et al. 2011; skead 2007; c. vernon pers. comm., 13 november 2013). this perceived distribution gap is observed for many reptile species, particularly chelonians (bates et al. 2014; branch, benn & lombard 1995). there are several examples of currently known reptile species distributions (e.g. for trachylepis capensis, bitis arietans and lamprophis aurora) extending all the way to the kei river along the east coast, with a gap in distribution through the coastal transkei–pondoland region, and afterwards extending further north along the kwazulu-natal coastline (bates et al. 2014; minter et al. 2004). the lack of herpetofaunal records along the wild coast is a biodiversity conservation concern. it is not clear whether the current conservation initiatives in the wild coast region, which forms part of the larger maputaland–pondoland–albany global biodiversity hotspot (mittermeier et al. 2004), are sufficient for protecting herpetofauna. this relates specifically to species of special conservation concern, as well as to providing a sufficient north-east to south-west corridor for species dispersal. in addition, the resulting lack of knowledge increases the risk of negative impacts on amphibian and reptile species through poor management and development. the objectives of this study were (1) to provide an annotated list of amphibians and reptiles occurring in protected areas along the wild coast and (2) to assess representation of species of special conservation concern (threatened and endemic species) in relevant protected areas. study areas top ↑ we surveyed a total of six protected areas along the wild coast in the eastern cape province, south africa, from 2011 to 2013 (figure 1). the area that constitutes the ‘wild coast’ stretches from north-east of east london to the kwazulu-natal border and includes the former transkei and the pondoland regions. protected areas comprise (from south to north) the cape morgan nature reserve, manubi state forest, dwesa nature reserve, hluleka nature reserve, silaka nature reserve and mkambati nature reserve. figure 1: the location of the six protected areas along the wild coast, eastern cape, south africa, where reptile and amphibian surveys were conducted. cape morgan nature reserve east london coast nature reserves comprises a number of small nature reserves (nrs) and state forests (sfs) situated primarily along the coastline between the great kei river in the north and the great fish river in the south (figure 1). the survey was conducted only in the cape morgan (214 ha) and double-mouth (199 ha) sections of this reserve. vegetation types consist of southern coastal forest (low forests with a well-developed low tree and shrub component) (mucina & geldenhuys 2006) and albany coastal belt grassland (hoare et al. 2006). the survey was conducted from 26 march to 02 april 2012 (venter & conradie 2012a). manubi state forest the manubi sf (with an adjacent proposed expansion area) is located in the south-western part of the wild coast, south-east of the town butterworth (figure 1). manubi sf and the adjacent area contain small patches of subtropical dune thicket, but are dominated by scarp forest and transkei coastal belt vegetation types (mucina & geldenhuys 2006; mucina et al. 2006). the survey was conducted from 18 to 27 february 2013 (venter et al. 2013). dwesa nature reserve the dwesa nr (3623 ha) is located along the western side of the mbashe river estuary (figure 1). the reserve occupies a narrow coastal strip of approximately 9 km in length and 1 km – 2 km in width. dwesa nr comprises roughly 80% indigenous coastal scarp forest and 20% transkei coastal belt grassland (mucina & geldenhuys 2006; mucina et al. 2006). dune forest and scrub forest are also found close to the coast and are classified as sensitive vegetation types (mucina et al. 2006). the survey was conducted from 05 to 15 february 2012 (venter & conradie 2012b). hluleka nature reserve the hluleka nr (577 ha) is situated nearly in the centre of the wild coast (figure 1), south-west of port st johns along the coast. scarp forest and coastal forest cover approximately 70% of the reserve, whereas the remainder of the vegetation is made up of thicket and transkei coastal belt grassland (mucina & geldenhuys 2006; mucina et al. 2006). the survey was conducted from 20 to 28 october 2011 (venter & conradie 2012c). silaka nature reserve silaka nr is a small (400 ha) protected area situated south-west of port st johns (figure 1). it is made up mainly of a forested valley located between second beach and sugar loaf rock. vegetation types include transkei coastal belt grassland (mucina et al. 2006) and scarp forest (mucina & geldenhuys 2006). the survey was conducted from 29 september to 06 october 2012 (venter, conradie & greeff 2012). mkambati nature reserve mkambati nr (7736 ha) is situated along the north-eastern pondoland coast (figure 1). the vegetation types are dominated by fire-prone pondoland ugu sandstone coastal sourveld with patches of scarp and swamp forest in fire refuge areas (mucina & geldenhuys 2006; mucina et al. 2006). this reserve also contains a large population of wild ungulates (venter et al. 2014). the survey was conducted from 07 to 13 february 2011 (venter & conradie 2011). material and methods top ↑ we used a combination of visual encounter survey methods and standard y-shape trap arrays. each trap array consisted of three drift fences of 10 m long and 50 cm high, positioned in a y-shape, with four pitfall traps at the ends and middle, and two one-way funnels per fence – see appendix 4. surveys of the six wild coast protected areas were conducted from 2011 to 2013. diurnal searches were conducted by actively searching specific microhabitats, including beneath rocks and decaying logs. nocturnal surveys were also carried out with the use of headlamps or flashlights, and advertisement calls were recorded in the field using either an olympus vn-3500pc voice recorder or nagra ares-ml recorder with an external sony f-v4t microphone. all specimens other than voucher specimens were captured by hand or net (e.g. tadpoles), photographed and released back to the original capture site. voucher specimens were humanely euthanised by injecting reptiles with and submerging amphibians in a tricaine methanesulfonate (ms222) solution (conroy et al. 2009), after which they were fixed in formalin for 48 hours and transferred to 50% isopropanol for long-term storage in the herpetological collection of the port elizabeth museum (pem). specimens were identified using field guides and publications (branch 1998; channing 2001; channing & baptista 2013; channing et al. 2013; du preez & curruthers 2009; marais 2004; petzold et al. 2014). nomenclature was mainly based on frost (2014) for amphibians and uetz (2014) for reptiles, or updated taxonomy in recent journal publications (channing & baptista 2013; petzold et al. 2014). conservation status is based on the latest iucn assessments, which use updated evaluations for amphibians (measey 2011) and reptiles (bates et al. 2014). all fieldwork was supervised by the first author, who was a staff member of the eastern cape parks and tourism agency at the time, and formed part of the agency's operational activities as the appointed management authority of the protected areas stipulated by the eastern cape parks and tourism agency act (no. 2 of 2010). collection and handling of specimens was according to ethical standards of bayworld/port elizabeth museum (bayworld ethical clearance 2013_001). rating of the conservation importance of each reserve was based on the sum of the categorised scoring of the current iucn-listed species (critically endangered = 4; endangered = 3; vulnerable = 2; near threatened = 1; least concern = 0), level of endemism based on the proposed maputaland–pondoland–albany regional list of endemic to near-endemic species compiled by perera et al. (2011) (endemic = 2; near endemic = 1), and the total number of species found per reserve. additional records were sourced from the southern african reptile conservation assessment (sarca) (bates et al. 2014), south african frog atlas project (safap) (minter et al. 2004) and historical literature records pertaining to the study area (appendix 3). records with appropriate supporting evidence (identifiable photographs or voucher specimens) were included in the final species checklist, for example burger (1996) and the grobler photograph (see figure 4c). figure 2: a comparison of the number of (a) amphibian and (b) reptile species and families and number of iucn red listed species, as well as the conservation importance scores recorded in the six protected areas during the study. figure 3: amphibian species of special or conservation concern recorded in protected areas along the wild coast, including (a) arthroleptis wahlbergii (hluleka nr), (b) leptopelis natalensis (silaka nr), (c) breviceps cf. bagginsi (silaka nr), (d) afrixalus spinifrons (hluleka nr), (e) hyperolius poweri (mkambati nr) and (f) natalobatrachus bonebergi (manubi sf). figure 4: reptile species of special or conservation concern recorded in protected areas along the wild coast, including (a) bradypodion caffer (mkambati nr), (b) tetradactylus cf. fitzsimonsi (hluleka nr), (c) python natalensis (mkambati nr), (d) amblyodipsas concolor (hluleka nr), (e) macrelaps microlepidotus (manubi sf) and (f) crocodylus niloticus (dwesa nr). results and discussion top ↑ a total of 22 amphibian species, representing 8 frog families and 15 genera (table 1), and 37 reptile species (20 snakes, 16 lizards, 1 terrapin), representing 17 families and 33 genera (table 2), were recorded from the six protected areas surveyed during this study (appendix 2). this represents a near-complete list of species expected to occur along the wild coast (27 amphibian species were recorded by minter et al. [2004] and 50 reptile species were recorded by bates et al. [2014]). although our survey excluded marine species, four species of marine turtle (dermochelys coriacea, chelonia mydas, eretmochelys imbricata and caretta caretta) and one sea snakes species (hydrophis platurus) are expected to occur along the coastline (bates et al. 2014). a total of three threatened amphibian and five threatened reptile species are represented in these protected areas. a total of 243 quarter-degree-unit (qdu) records were recorded, of which 37% (23% amphibians and 50% reptiles) represented first records for the respective units (table 3), with southerly range extensions for 11 species. these include arthroleptis wahlbergii, breviceps cf. bagginsi, hyperolius poweri, leptopelis natalensis, natalobatrachus bonebergi, amblyodipsas concolor, leptotyphlops sylvicolus, psammophis brevirostris, python natalensis, thelotornis capensis capensis and tropidosaura montana natalensis (see tables 1, 2 and appendix 1). the most southerly distribution in africa was also recorded for the following genera: amblyodipsas, arthroleptis, leptopelis, natalobatrachus and thelotornis (appendix 1). furthermore, the distribution of bradypodion caffer was extended northwards. range extensions exceeding 100 km were also recorded for a. concolor and p. brevirostris (appendix 1). the distribution range of one species, t. montana natalensis, was extended to the eastern cape herpitofaunal list. a potential new species, which is related to tetradactylus fitzsimonsi from the port elizabeth area, was discovered in hluleka nr (appendix 1). hluleka, silaka and mkambati nrs had the greatest numbers of amphibian species, whereas dwesa, hluleka and mkambati nrs had the greatest numbers of reptile species (table 3 and figure 2). with regard to amphibians, silaka nr has a higher conservation importance, which can be attributed to the presence of three threatened, five endemic and three near-endemic species. both dwesa and hluleka nrs are important for reptile conservation, as they are home to at least two threatened, three endemic and two near-endemic species each. regarding overall conservation importance, hluleka nr scored the highest, followed by, in descending order, dwesa nr, mkambati nr, silaka nr, manubi sf and cape morgan nr (table 3 and figure 2). species of special conservation concern or noteworthy records are illustrated in figures 3 and 4. table 1: amphibian checklist for six conservation areas surveyed along the wild coast, eastern cape, south africa. table 2: reptile checklist for six conservation areas surveyed along the wild coast, eastern cape, south africa. table 3: new quarter-degree-unit records for south africa and conservation importance score of six conservation areas studied along the wild coast, eastern cape, south africa. conclusion top ↑ we provide valuable data on amphibian and reptilian species presence and distribution for a fairly under-sampled region in south africa. our study further highlights the importance of baseline survey work to inform conservation management planning for protected areas and species of special conservation concern. the discovery of several healthy and thriving populations of threatened species, such as b. cf. bagginsi, n. bonebergi, afrixalus spinifrons, macrelaps microlepidotus and bradypodion spp., is very encouraging. it further indicates that even small protected areas, as is the case with silaka nr, hluleka nr and manubi sf, play an important role in conserving a number of threatened species, considering the development pressure many of these species face in kwazulu-natal (minter et al. 2004). future research should focus on investigating the impact of current land-use activities (communal subsistence farming) on species conservation along the wild coast compared with that of commercial agriculture, forestry and high-density development in areas such as south-western kwazulu-natal. this is especially important because of the significant and increasing pressure placed on the wild coast region owing to slash-and-burn agriculture, illegal activities such as sand mining, road construction and logging, as well as water pollution from neglected or non-functional municipal sewage treatment works (de villiers & costello 2013). significant portions of land north-east of mkambati nr (xolobeni area) are currently being considered for mining, which, together with the construction of the proposed national (n2) toll road, will result in additional habitat loss and fragmentation for several important herpetofaunal species. it is therefore critical that the current protected areas in the region remain secure and that opportunities are sought to increase the size of the protected areas in the region to form corridors for dispersal. in addition, it is crucial that local, provincial and national government agencies take note of the potential negative consequences of increased land degradation in the region, the effect on biodiversity and the subsequent impact on local communities. our study provides the first checklist for amphibians and reptiles from the wild coast protected areas, but gaps remain owing to factors related to spatial and temporal under-sampling. the results reported here represent only one sampling trip per protected area, conducted during either early or late summer, which may have led to species being missed owing to non-breeding or inactivity. it is important that further surveys and monitoring are conducted to increase sampling effort and to capture seasonal differences in species assemblages and densities. monitoring at regular intervals, in order to measure the effect of land use or climate change on the populations of threatened species especially, is recommended. our surveys focused on the coastal areas of the wild coast, with the inland area of the transkei remaining under-sampled. a rich source of undiscovered herpetofaunal diversity is likely to still be uncovered. acknowledgements top ↑ the eastern cape parks and tourism agency (ecpta) funded the study. neil brown and numerous students from the parks and people program of penn state university provided valuable field assistance. juan greeff (ecpta) provided technical assistance. gillian watson (port elizabeth museum, bayworld) allowed the examination of specimens in her care. lemmy mashini (ditsong national museum of natural history) provided data of tm specimens. andrew turner (capenature) provided additional records. rené navarro (avian demography unit, university of cape town) provided safap and sarca data. krystal tolley (south african national biodiversity institute), savel daniels (stellenbosch university), div de villiers (eastern cape department of economic development and environmental affairs) and marius burger provided valuable inputs on certain species account sections. christa conradie provided proofreading services. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions j.a.v. (nelson mandela metropolitan university) was the project leader. he was responsible for experimental design, conducted fieldwork and performed data analysis. w.c. (port elizabeth museum) assisted in experimental design, conducted fieldwork, handled all preservation and lodging of specimens and performed data analysis. references top ↑ bates, m.f., branch, w.r., bauer, a.m., burger, m., marais, j., alexander, g.j. et al., 2014, atlas and red list of the reptiles of south africa, lesotho and swaziland, south african national biodiversity institute, pretoria (suricata series; no. 1). böhm, m., collen, b., baillie, j.e.m., bowles, p., chanson, j., cox, n. et al., 2013, ‘the conservation status of the world's reptiles’, biological conservation 157, 372–385. http://dx.doi.org/10.1016/j.biocon.2012.07.015 botts, e., erasmus, b.n. & alexander, g., 2011, ‘geographic sampling bias in the south african frog atlas project: implications for conservation planning’, biodiversity and conservation 20(1), 119–139. http://dx.doi.org/10.1007/s10531-010-9950-6 branch, w.r., 1998, field guide to the snakes and other reptiles of southern africa, struik, cape town. branch, w.r., 2006, ‘priorities for systematic studies on southern african reptiles’, in w.r. branch, k.a. tolley, m. cunningham, a.m. bauer, g. alexander, j.a. harrison et al. (eds.), a plan for phylogenetic studies of southern african reptiles: proceedings of a workshop held at kirstenbosch, february 2006, pp. 1–15, south african national biodiversity institute, pretoria (sanbi biodiversity series; no 5). branch, w.r., 2014, ‘conservation status, diversity, endemisim, hotspots and threats’, in m.f. bates, w.r. branch, a.m. bauer, m. burger, j. marais, g.j. alexander et al. (eds.), atlas and red data list of the reptiles of south africa, lesotho and swaziland, pp. 22–50, south african national biodiversity institute, pretoria (suricata series; no. 1). branch, w.r., benn, g.a. & lombard, a.t., 1995, ‘the tortoises (testudinidae) and terrapins (pelomedusidae) of southern africa: their diversity, distribution and conservation’, south african journal of zoology 30, 91–102. branch, w.r. & haagner, g.v., 1999, ‘geographical distribution: psammophis brevirostris and acontias sp.: the value of road kills’, african herp news 30, 37–38. burger, m., 1996, ‘report on a herp[e]tofaunal survey conducted in the former transkei region’, unpublished report, ministry of economic affairs, environment & tourism, eastern cape nature conservation. channing, a., 2001, amphibians of central and southern africa, cornell university press, new york. channing, a. & baptista, n., 2013, ‘amietia angolensis and a. fuscigula (anura: pyxicephalidae) in southern africa: a cold case reheated’, zootaxa 3640, 501–520. http://dx.doi.org/10.11646/zootaxa.3640.4.1 channing, a., hillers, a., lotters, m.o., schick, s., conradie, w., rodder, d. et al., 2013, ‘taxonomy of the super-cryptic hyperolius nasutus group of long reed frogs of africa (anura: hyperoliidae), with descriptions of six new species’, zootaxa 3620, 301–350. http://dx.doi.org/10.11646/zootaxa.3620.3.1 conroy, c.j., papenfuss, t., parker, j. & hahn, n.e., 2009, ‘use of tricaine methanesulfonate (ms222) for euthanasia of reptiles’, journal of the american association for laboratory animal science 48, 28–32. de villiers, d. & costello, j., 2013, mkambati and the wild coast, 2nd edn., div de villiers and john costello, port st johns. dippenaar, a.s., hamer, m. & haddad, c.r., 2011, ‘spiders (arachnida: araneae) of the vegetation layer of the mkambati nature reserve, eastern cape, south africa’, koedoe 53(1), 1–10. du preez, l.h. & curruthers, v.c., 2009, a complete guide to the frogs of southern africa, struik nature, cape town. fitzsimons, v.f.m., 1930, ‘descriptions of new south african reptilia and batrachia, with distribution records of allied species in the transvaal museum collection’, annals of the transvaal museum 14, 20–48. frost, d.r., 2014, amphibian species of the world: an online reference, version 6.0, online database, viewed 24 june 2014, from http://research.amnh.org/herpetology/amphibia/index.html, american museum of natural history, new york. haagner, g.v., 1994, ‘geographical distribution: amblyodipsas concolor’, african herp news 21, 26. hoare, d.b., mucina, l., rutherford, m.c., vlok, j., euston-brown, d.i.w., palmer, a.r. et al., 2006, ‘albany thicket biome’, in l. mucina & m.c. rutherford (eds.), the vegetation of south africa, lesotho and swaziland, south african national biodiversity institute, pretoria. marais, j., 2004, a complete guide to the snakes of southern africa, struik, cape town. measey, g.j., 2011, ensuring a future for south africa’s frogs: a strategy for conservation research, south african national biodiversity institute, petoria (sanbi biodiversity series; no. 19). measey, g.j., armstrong, a.j. & hanekom, c., 2009, ‘subterranean herpetofauna show a decline after 34 years in ndumu game reserve, south africa’, oryx 43(2), 284–287. http://dx.doi.org/10.1017/s0030605307002311 minter, l.r., burger, m., harrison, j.a., braack, h.h., bishop, p.j. & kloepfer, d., 2004, atlas and red data book of the frogs of south africa, lesotho and swaziland, smithsonian institute, cape town. mittermeier, r.a., robles-gil, p., hoffman, m., pilgrim, j.d., brooks, t., mittermeier, c. et al., 2004, revisited: earth’s biologically richest and most endangered ecoregions, cemex, mexico city. mokhatla, m.m., measey, g.j., chimimba, c.t. & van rensburg, b.j., 2012, ‘a biogeographical assessment of anthropogenic threats to areas where different frog breeding groups occur in south africa: implications for anuran conservation’, diversity and distributions 18(5), 470–480. http://dx.doi.org/10.1111/j.1472-4642.2011.00870.x mucina, l. & geldenhuys, c.j., 2006, ‘afrotemperate, subtropical and azonal forests’, in l. mucina & m.c. rutherford (eds.), the vegetation of south africa, lesotho and swaziland, pp. 584–614, south african national biodiversity institute, pretoria. mucina, l., scott-shaw, c.r., rutherford, m.c., camp, k.g.t., matthews, w.s., powrie, l.w. et al., 2006, ‘indian ocean coastal belt’, in l. mucina & m.c. rutherford (eds.), the vegetation of south africa, lesotho and swaziland, pp. 568–583, south african national biodiversity institute, pretoria. ochoa-ochoa, l., urbina-cardona, j.n., vázquez, l.-b., flores-villela, o. & bezaury-creel, j., 2009, ‘the effects of governmental protected areas and social initiatives for land protection on the conservation of mexican amphibians’, plos one 4(9), e6878, 9 pages. perera, s.j., ratnayake-perera, d. & proches, s., 2011, ‘vertebrate distributions indicate a greater maputaland-pondoland-albany region of endemism’, south african journal of science 107(7/8), 1–15. http://dx.doi.org/10.4102/sajs.v107i7/8.462 petzold, a., vargas-ramirez, m., kehlmaier, c., vamberger, m., branch, w.r., du preez, l. et al., 2014, ‘a revision of african helmeted terrapins (testudines: pelomedusidae: pelomedusa), with descriptions of six new species’, zootaxa 3795, 523–548. http://dx.doi.org/10.11646/zootaxa.3795.5.2 power, j.h., 1935, ‘contribution to the herpetology of pondoland’, proceedings of the zoological society of london 1935, 333–346. http://dx.doi.org/10.1111/j.1469-7998.1935.tb06253.x russell, k.r., lear, d.h.v. & guynn jr, d.c., 1999, ‘prescribed fire effects on herpetofauna: review and management implications’, wildlife society bulletin 27(2), 374–384. skead, c.j., 2007, historical incidence of the larger mammals in the broader eastern cape, 2nd edn., nelson mandela metropolitan university, port elizabeth. tuberville, t.d., willson, j.d., dorcas, m.e. & gibbons, j.w., 2005, ‘herpetofaunal species richness of southeastern national parks’, southeastern naturalist 4(3), 537–569. http://dx.doi.org/10.1656/1528-7092(2005)004[0537:hsrosn]2.0.co;2 uetz, p., 2014, the reptile database, viewed 23 june 2014, from http://www.reptile-database.org/ van wyk, a.e. & smith, g.f., 2001, regions of floristic endemism in southern africa, umdaus press, hatfield. venter, j.a. & conradie, w., 2011, ‘the herpetological fauna of mkambati nature reserve: a report on the herp[e]tological survey conducted in february 2011’, unpublished report, eastern cape parks and tourism agency, east london. venter, j.a. & conradie, w., 2012a, ‘the amphibians and reptiles of cape morgan and double-mouth nature reserves’, unpublished report, eastern cape parks and tourism agency, east london. venter, j.a. & conradie, w., 2012b, ‘the amphibians and reptiles of dwesa nature reserve: a report on a rapid biodiversity survey conducted in february 2012’, unpublished report, eastern cape parks and tourism agency, east london. venter, j.a. & conradie, w., 2012c, ‘the amphibians and reptiles of hluleka nature reserve: a report on a rapid biodiversity survey conducted in october 2011 ’, unpublished report, eastern cape parks and tourism agency, east london. venter, j.a., conradie, w. & greeff, j., 2012, ‘the amphibians and reptiles of silaka nature reserve: a report on a rapid biodiversity survey conducted in october 2012’, unpublished report, eastern cape parks and tourism agency, east london. venter, j.a., conradie, w., pfeiffer, m. & greeff, j., 2013, ‘the reptiles, frogs, birds, mammals and dragonflies of manubi forest and expansion areas’, unpublished report, eastern cape parks and tourism agency, east london. venter, j.a., prins, h.h.t., balfour, d.a. & slotow, r., 2014, ‘reconstructing grazer assemblages for protected area restoration’, plos one 9(3), e90900, 10. appendix 1 top ↑ species accounts this section provides selective species accounts (excluding common and widespread species, marine turtles and sea snakes) to highlight special records for the region. species account information was sourced from the individual species account sections in minter et al. (2004) and bates et al. (2014), and updated with our new records unless stated otherwise. families are organised according to frost (2014) for amphibia and bates et al. (2014) for reptilia. species of special conservation concern or noteworthy records are illustrated in figures 3 and 4 in the main text. all additional pem holdings collected and used in this study are listed in appendix 2 and additional records derived from other sources are listed in appendix 3. records labelled as pem a refer to amphibian records from the port elizabeth museum, those labelled pem t refer to tadpole records from this collection and those labelled pem r refer to reptilian records from the collection. amphibia arthroleptidae arthroleptis wahlbergii smith, 1849 (bush squeaker) this species is endemic to the coastal forest patches of the east coast of south africa, from the border of mozambique to just south of port st johns. because of this restriction, the species is under encroachment pressure owing to agricultural and urban development. the hluleka nr (3129cd) specimens represent the most southern distribution of the species, as well as for the genus in africa. two additional inland records from mqanduli (pem a7368, pem a7369) are included in the pem collection. fitzsimons (1930) considered the population around port st johns as a new species, namely arthroleptis wageri, citing a broader head, shorter hind limbs and larger general size. this arrangement was also followed by laurent (1941). however, poynton (1964) synonymised this species with the more common a. wahlbergii. genetic studies currently underway on this species show substantial genetic differentiation for this eastern cape population, pointing to the possibility that a. wageri might be a valid species (k. tolley pers. comm., 14 july 2014). the species was recorded only from hluleka and silaka nrs during this study. brevicipitidae breviceps cf. bagginsi minter, 2003 (bilbo's rain frog) a series of specimens collected from silaka nr (3129cb, –da) during this study and one additional pem specimen (pem a11122) previously collected 15 km west of mkambati nr (3129bb, new qdu record) have temporarily been assigned to breviceps bagginsi, based on the call and morphology. in silaka nr, specimens were captured in traps associated with grassland habitat (see ‘study area’). these new records potentially expand the known distribution of this species southwards to the port st johns area. burger (1996) reported an unidentified breviceps species from mkambati nr (3129bd), recorded as breviceps mossambicus by minter (2004b) and later identified as b. bagginsi (measey 2011). the specific specimen is not available in any museum collection and thus could not be verified. this species belongs to the b. mossambicus complex and further studies are needed in the northern coastal eastern cape to determine whether the applicable records of b. mossambicus in minter (2004b) can be assigned to b. bagginsi. minter (2004b) mistakenly reported b. mossambicus as occurring as far south as mkambati nr, but the provided qdu actually represents port st johns. b. bagginsi is a recently described species known only from four kwazulu-natal qdu records (minter 2004a) and is believed to be endemic to kwazulu-natal. thus, the new distribution records for the eastern cape and the presence in a protected area will contribute to protecting the species. the species is currently rated as vulnerable (measey 2011). the species was recorded only from silaka nr during this study. hyperoliidae afrixalus spinifrons cope, 1862 (natal leaf-folding frog) an endemic south african species distributed from st. lucia village in the north to cintsa bay and kei road in the south. pickersgill (1996) assigned the eastern cape populations to a. spinifrons spinifrons, but tarrant (2012) showed that, according to molecular description, eastern cape populations should be referred to afrixalus spinifrons intermedius. the species is currently rated as near threatened (measey 2011). whilst documenting and photographing freshly laid eggs in situ at silaka nr, an adult ran up the stem of the leaf to the clutch and assumed a threatening posture. this behaviour can possibly be attributed to a form of parental care or protection, which has never been reported for the species. the species was found in both silaka and hluleka nrs during the study. hyperolius poweri loveridge, 1938 channing et al. (2013a) assigned the kwazulu-natal and eastern cape populations of the hyperolius nasutus complex (for which mkambati nr is the most southern distribution) to h. poweri based on genetics and calls. because of the recent taxonomical change, the species’ threatened status has not been assessed. however, owing to its patchy distribution along the east coast of south africa and loss of habitat due to agricultural and urban development, the species may in the future be considered threatened and treated as a separate conservation unit. the species was found in mkambati nr during the study, which is the only known occurrence of the species in the eastern cape. pyxicephalidae cacosternum boettgeri boulenger, 1882 (boettger's caco) recent revision of the genus by channing et al. (2013b) indicated the presence of cryptic diversity and led to the description of four new species (cacosternum aggestum, cacosternum australis, cacosternum nanogularum and cacosternum rhythmum), with conradie (2014) describing another species within the c. boettgeri complex (cacosternum thorini). in both studies, no material from coastal eastern cape or kwazulu-natal was used. populations from these areas may hold further cryptic diversity. burger (1996) reported this species to be present in mkambati nr based on the call only, and the safap database lists a voucher specimen collected from dwesa nr in late 1991. the specimen is unaccounted for in any of the south african museums; thus, its identification cannot be verified. owing to its possible distribution, we tentatively include this species in our records. however, no new material has been collected from any of the reserves surveyed during the current study. further surveys in optimal conditions are needed to confirm the presence of this species from coastal eastern cape. natalobatrachus bonebergi hewitt & methuen, 1912 (kloof frog) the species is currently known from only nine localities in kwazulu-natal and the eastern cape (measey 2011). historically, it has been recorded from 18 qdus (du preez 2004), of which three are in the eastern cape. during the frog atlas period (1996–2002), only two of the historical sites were confirmed and one new qdu was added (dwesa nr, 3228bb), which represented its most southern distribution (du preez 2004). we report three new qdu records (3228bd, 3129cb and 3228bc), which now represent the most southern distribution, and confirm the two historical qdu sites (3129da and 3129cd) previously recorded. this increases the existing distribution to seven qdu sites within the eastern cape, including 3228bb and the historical record (3229cc). the habitat of n. bonebergi is currently being heavily affected by agricultural clearing (especially for sugar cane) and urbanisation, particularly in kwazulu-natal (measey 2011). as a result, populations of this species are becoming severely fragmented and are thus regarded as endangered (measey 2011). reptilia crocodylidae crocodylus niloticus laurenti, 1768 (nile crocodile) crocodiles are restricted to the northern and eastern parts of south africa. they were re-introduced to dwesa nr in 1979 (pooley 1980), which has been criticised as ‘ill-advised’ by marais (2014a). this criticism may be unwarranted because good evidence exists that the species historically (during the 19th and early 20th centuries) occurred even further south than the current location in dwesa nr, with the last record of a crocodile killed in the former transkei recorded in the mendwana river, dwesa forest in 1903 (feely 2010). evidence also exists that the re-introduced dwesa nr population did successfully breed (feely 2010; venter & conradie 2012), but the survival rate of young crocodiles is still to be determined. only two adults were recorded in the latest game census (peinke & gibisela 2013). this species is rated as vulnerable (bates et al. 2014). gekkonidae afroedura pondolia hewitt, 1925 (pondo flat gecko) this species is endemic to the eastern cape and kwazulu-natal coast and associated with scattered inland records. it reaches its southern distribution limit around the coastal dwesa nr (3228bd) and inland butterworth (3228ad) area. it is present in mkambati, silaka (tm 76520), hluleka and dwesa nrs. lygodactylus capensis smith, 1848 (common dwarf gecko) this is a common and widespread diurnal gecko species, which is not native to the eastern cape (branch 2014). populations at silaka and cape morgan nrs were most likely accidentally introduced by tourists visiting the reserves. all specimens collected or observed were limited to buildings and other human-made structures. lacertidae tropidosaura montana natalensis fitzsimons, 1947 (common mountain lizard) this subspecies is restricted to southern kwazulu-natal, where it is found in both coastal and montane grassland. a single record from mkambati nr was recorded during this study, its remains being regurgitated by a lycophidion capense. this is the first record for this species in the eastern cape. gerrhosauridae gerrhosaurus flavigularis wiegmann, 1828 (yellow-throated plated lizard) this species occurs mostly in north-eastern parts of south africa, with an apparently isolated southern population. in a recent revision of the entire genus, three well-structured clades were formed (i.e. eastern south africa, northern south africa and east africa) and current research may split these populations (bates et al. 2013). if this is the case, the southern population will keep the name g. flavigularis, which will make the silaka nr material the most northern record. the species was observed at three of the six reserves surveyed. tetradactylus cf. fitzsimonsi (fitzsimons’ long-tailed seps) tetradactylus fitzsimonsi is restricted to only four qdus in the port elizabeth area and one near george. the new records from hluleka nr were found 600 km to the north-east of the t. fitzsimonsi distribution range. identification of these specimens is based on fitzsimons (1943), with the main identifying characteristic being the lack of front limbs. current genetic and morphometric work indicates that the hluleka specimens might be a novel species related to t. fitzsimonsi (k. tolley pers. comm., 14 july 2014). more than a century of herpetological studies in the eastern cape have failed to find any additional t. fitzimonsi records from the coastal eastern cape or the former transkei. furthermore, overlap of the distribution for t. fitzsimonsi and the congener, tetradactylus africanus, is not clear. fitzsimons (1943) gives the distribution of t. africanus as pondoland and northwards (i.e. northern eastern cape), but no recent records have been collected from this area (bates 2014a, 2014b). the ditsong national museum of natural history (tm 69067) houses a specimen of t. africanus from msikaba river, near mkambati nr (bates 2014c). additional surveys in coastal grassland along the eastern cape coastline will improve information on this species’ distribution. for the purpose of this species inventory we consider the species as t. fitzsimonsi. scincidae acontias plumbeus bianconii, 1849 (giant legless skin) this species represents the world's largest species of legless skink, with a maximum size recorded at 490 mm, and is restricted to well-forested areas (branch 1998). an isolated eastern cape population around east london shows a number of subtle morphological differences from northern populations (bourquin & lambiris 1996), and possibly represents a new species (branch 2002). genetic studies are currently underway to determine the relationship of this population to northern populations (daniels, in preparation). its sister species, acontias poecilus, is known from only five specimens in south-eastern kwazulu-natal (bourquin & lambiris 1996) and only a single record from the eastern cape (branch & haagner 1999), but may be present in mkambati nr (bauer 2014). dwesa nr has a selection of preserved specimens on display for educational purposes, which includes a preserved a. plumbeus from the reserve (m. burger pers. comm., 23 april 2014). this forms the basis of the present record in sarca. we confirmed this distribution during the recent survey of dwesa nr and reported a new distribution record for the manubi sf area. trachylepis homalocephala wiegmann, 1828 (red-sided skink) this species is found mostly in the eastern cape and western cape, with scattered populations in kwazulu-natal, lesotho, the free state, mpumalanga and limpopo. in the recent reptile atlas (bates et al. 2014) there is a noticeable gap in the distribution from east london to the closest kwazulu-natal populations. this gap has been described as a biodiversity-poor gap due to negative environmental conditions (see main text). we recorded this species from all the reserves, indicating a sampling gap rather than a natural distribution gap. more surveys need to be conducted outside the reserves to determine the full extent of this species’ distribution. chamaeleonidae bradypodion caffer boettger, 1889 (pondo dwarf chameleon) dwarf chameleons from mkambati nr have been assigned to bradypodion melanocephalum (raw 2001; tolley & burger 2007), given their morphological similarity to the species. they were therefore regarded as the population with the most southern distribution limit of the species (tolley 2014a). in contrast, raw (2001) regarded the mkambati population as a new species. recent molecular work on this population shows a close relationship to b. caffer, which may represent a grassland ecomorph (k. tolley pers. comm., 14 july 2014). b. caffer is a forest-adapted species, whereas the mkambati population was found only in natural grassland. although morphological similarity might be expected within a species (i.e. amongst populations), structure is a known driver of morphological variation in dwarf chameleons habitat (da silva & tolley 2013; hopkins & tolley 2011; tolley 2014a). the grassland population of mkambati most likely represents an ecomorph of b. caffer and is another example of ecological adaptation due to differences in habitat structure in this group. the forest and grassland ecomorphs should be considered as separate management units with regard to conservation and management. despite a close genetic relationship, the two ecomorphs are distinct with regard to the morphological adaptations that allow them to persist in the different environments. additional molecular and morphological work is required to better place the mkambati population. b. caffer was also recorded from the forests at silaka and hluleka nrs (k. tolley pers. comm., 14 july 2014) and has been classified as endangered (tolley 2014b). bradypodion kentanicum hewitt, 1935 (kentani dwarf chameleon) this species is restricted to coastal scarp forest from kentani in the south to coffee bay in the north (tolley & burger 2007). it was represented in dwesa nr and manubi sf and has been assessed as vulnerable (tolley 2014c). leptotyphlopidae leptotyphlops sylvicolus broadley & wallach, 1997 (forest thread snake) the species is restricted to coastal moist forest (except for the matatiele area, where they were found in grassland). eastern cape specimens differ morphologically from the main type and genetic differences indicate that a number of cryptic species exist. this is specifically the case with l. sylvicolus (adalsteinsson et al. 2009). further investigations of the isolated eastern cape specimens are needed and the populations may warrant special conservation concern. the species is currently regarded as data deficient (alexander 2014a). it was collected from only two of the reserves surveyed and both represented new records for this species. at mkambati nr, specimens were collected under rocks in gardens, but the surrounding vegetation type is grassland. in hluleka nr, specimens were collected amongst the chalets, which are surrounded by coastal forest. pythonidae python natalensis smith, 1840 (southern african python) southern african pythons once extended to port alfred (eastern cape), but the last known specimen from this area was killed in bathurst in 1927 (broadley 1990; c. vernon pers. comm. 13 november 2013). there are unconfirmed historical records of pythons killed in the manubi or dwesa forest region (records seem to be unclear which forest) in 1845 (c. vernon pers comm., 13 november 2013). in the early 1980s, specimens were re-introduced in the andries vosloo nr outside grahamstown, but no follow-up work has been done on this population (alexander 2014b; c. vernon pers. comm., 13 november 2013). anecdotal records exist for mkambati nr (burger 1996; de villiers & costello 2013), as well as a single photographic record by ecpta staff (see figure 4c). the occurrence of p. natalensis in mkambati nr is noteworthy, given that almost nothing has been recorded regarding python distribution in the former transkei region and this is the only recent record for the eastern cape. lamprophiidae amblyodipsas concolor smith, 1849 (kwazulu-natal purple-glossed snake) this is a fossorial species restricted to moist forested areas in the eastern and northern parts of southern africa, feeding mostly on other snakes and lizards. this cryptic species can easily be confused with sympatric amblyodipsas polylepis, macrelaps microlepidotus and atractaspis bibronii, which occur further north. they differ from the latter two as they do not have preocular scales, but do have paired subcaudals and divided anal scales (broadley 1990; marais 2004). they are also differentiated from a. polylepis by having lower mid-body scale counts (17 vs 19) (broadley 1990; marais 2004). previously the only record for the eastern cape was based on a specimen from mtumbane, port st johns area (haagner 1994: tm 69016). we extend the species’ distribution 120 km southwards to manubi sf and add two additional new records south of port st johns (hluleka and dwesa nrs). no records of either a. polylepis or a. bibronii exist for the eastern cape and more surveys are needed in the most north-eastern parts of the eastern cape to confirm the presence of this species. macrelaps microlepidotus günther, 1863 (kwazulu-natal black snake) macrelaps microlepidotus is a semi-fossorial snake that inhabits moist forest and feeds on frogs, lizards and other snakes (marais 2004). its distribution is restricted mainly to the coastal eastern parts of south africa, with an inland expansion into the amatole region (eastern cape) and the escourt area (kwazulu-natal). it has previously been known from only seven qdus in the eastern cape. recently, the most inland eastern cape record was confirmed from the katberg area and two new qdus (one based on the collection from dwesa nr during this study) were added in the eastern cape (conradie, venter & nicolau 2012). we provide another new locality (manubi sf), further indicating continuous distribution along the east coast of the transkei. owing to habitat destruction this species is rated as near threatened (bates et al. 2014). psammophis brevirostris peters, 1881 (short-snouted grass snake) this species occurs mostly in the northern and eastern parts of south africa, inhabiting grassland and savannah. it has previously been found only in three localities in the eastern cape (bates et al. 2014; branch & haagner 1999), with its southern distribution limit around mpande beach (3129cd: pem r13694). we extend the distribution 100 km south to manubi sf and add two additional new records for the eastern cape (mkambati and hluleka nrs). a recently donated specimen to the pem collection expands the distribution of this species even further south to the kei river mouth (pem 20166). elapidae dendroaspis polylepis günther, 1864 (black mamba) only two records exist for this species in the eastern cape, namely from port st johns (tm 21455) and mkambati nr. the latter is based only on observational data (burger 1996; de villiers & costello 2013). it is likely that this species will occur more widely in the north-eastern parts of the eastern cape. we found none during this study. colubridae thelotornis capensis capensis smith, 1849 (south-eastern savannah vine-snake) t. capensis capensis is an arboreal species that occurs in coastal thicket and savannah. the only record for the eastern cape is an anecdotal sighting by d. de villiers (pers. comm., 23 april 2014). the specimen was sighted and photographed (photo could not be found) near the msikaba river vulture colony in mkambati nr (d. de villiers pers. comm., 23 april 2014). this forms the basis of the sarca record for the eastern cape (marais 2014b) and has been listed in de villiers and costello (2013). the record needs to be confirmed and voucher specimens need to be collected. references for appendix 1 top ↑ adalsteinsson, s.a., branch, w.r., trape, s., vitt, l.j & hedges, s.b., 2009, ‘molecular phylogeny, classification, and biogeography of snakes of the family leptotyphlopidae (squamata, scolecophidia)’, zootaxa 2244, 1–50. alexander, g.j., 2014a, ‘leptotyphlops sylvicolus broadley & wallach, 1997’, in m.f. bates, w.r. branch, a.m. bauer, m. burger, j. marais, g.j. alexander et al. (eds.), atlas and red data list of the reptiles of south africa, lesotho and swaziland, pp. 322–323, south african national biodiversity institute, pretoria (suricata series; no. 1). alexander, g.j., 2014b, ‘python natalensis a. smith, 1840’, in m.f. bates, w.r. branch, a.m. bauer, m. burger, j. marais, g.j. alexander et al. (eds.), atlas and red data list of the reptiles of south africa, lesotho and swaziland, p. 328, south african national biodiversity institute, pretoria (suricata series; no. 1). bates, m.f., 2014a, ‘tetradactylus africanus gray, 1838’, in m.f. bates, w.r. branch, a.m. bauer, m. burger, j. marais, g.j. alexander et al. (eds.), atlas and red data list of the reptiles of south africa, lesotho and swaziland, p. 231, south african national biodiversity institute, pretoria (suricata series; no. 1). bates, m.f., 2014b, ‘tetradactylus fitzsimonsi hewitt 1915’, in m.f. bates, w.r. branch, a.m. bauer, m. burger, j. marais, g.j. alexander et al. (eds.), atlas and red data list of the reptiles of south africa, lesotho and swaziland, p. 234, south african national biodiversity institute, pretoria (suricata series; no. 1). bates, m.f., 2014c, ‘geographical distributions: tetradactylus africanus (gray, 1838)’, african herp news 61(2), 35–36. bates, m.f., branch, w.r., bauer, a.m., burger, m., marais, j., alexander, g.j. et al., 2014, atlas and red data list of the reptiles of south africa, lesotho and swaziland, south african national biodiversity institute, pretoria. bates, m.f., tolley, k., edwards, s., davids, z., da silva, j. & branch, w.r., 2013, ‘a molecular phylogeny of the african plated lizards, genus gerrhosaurus wiegmann, 1828 (squamata: gerrhosauridae), with the description of two new genera’, zootaxa 3750(5), 465–493. http://dx.doi.org/10.11646/zootaxa.3750.5.3 bauer, a.m., 2014, ‘acontias poecilus bourquin & lambiris’, in m.f. bates, w.r. branch, a.m. bauer, m. burger, j. marais, g.j. alexander et al. (eds.), atlas and red data list of the reptiles of south africa, lesotho and swaziland, pp. 249–250, south african national biodiversity institute, pretoria (suricata series; no. 1). bourquin, o. & lambiris, a.j.l., 1996, ‘a new species of acontias [c]uvier (sauria: scincidae) from southeastern kwazulu-natal, south africa’, annals of the transvaal museum 36(17), 223–227. branch, w.r., 1998, field guide to the snakes and other reptiles of southern africa, struik, cape town. branch, w.r., 2002, ‘terrestrial fauna’, in ‘n2 wild coast toll road between durban and east london’, environmental impact assessment report, cca environmental, cape town. branch, w.r., 2014, ‘lygodactylus capensis capensis a. smith 1849’, in m.f. bates, w.r. branch, a.m. bauer, m. burger, j. marais, g.j. alexander et al. (eds.), atlas and red data list of the reptiles of south africa, lesotho and swaziland, p. 118, south african national biodiversity institute, pretoria(suricata series; no. 1). branch, w.r. & haagner, g.v., 1999, ‘geographical distribution: psammophis brevirostris and acontias sp.: the value of road kills’, african herp news 30, 37–38. broadley, d.g., 1990, fitzsimons’ snakes of southern africa, jonathan ball & ad. donker, parklands. burger, m., 1996, ‘report on a herp[e]tofaunal survey conducted in the former transkei region’, unpublished report, ministry of economic affairs, environment & tourism, eastern cape nature conservation. channing, a., hillers, a., lotters, m.o., schick, s., conradie, w., rodder, d. et al., 2013a, ‘taxonomy of the super-cryptic hyperolius nasutus group of long reed frogs of africa (anura: hyperoliidae), with descriptions of six new species’, zootaxa 3620, 301–350. http://dx.doi.org/10.11646/zootaxa.3620.3.1 channing, a., schmitz, a., burger, m. & kielgast, j., 2013b, ‘a molecular phylogeny of african dainty frogs, with the description of four new species (anura: pyxicephalidae: cacosternum)’, zootaxa 3701, 301–350. http://dx.doi.org/10.11646/zootaxa.3701.5.2 conradie, w., 2014, ‘the king of the dwarves: a new cryptic species of dainty frog (anura: pyxicephalidae: cacosternum) from the eastern great escarpment of south africa’, zootaxa 3785(3), 438–452. http://dx.doi.org/10.11646/zootaxa.3785.3.6 conradie, w., venter, j.a. & nicolau, j.r., 2012, ‘geographical distribution: macrelaps microlepidotus günther, 1860’, african herp news 58, 19–21. da silva, j.m. & tolley, k.a., 2013, ‘ecomorphological variation and sexual dimorphism in a recent radiation of dwarf chameleons (bradypodion)’, biological journal of the linnean society 109(1), 113–130. http://dx.doi.org/10.1111/bij.12045 de villiers, d. & costello, j., 2013, mkambati and the wild coast, 2nd edn., div de villiers and john costello, port st johns. du preez, l.h., 2004, ‘natalobatrachus bonbergi hewitt and methuen, 1913’, in l.r. minter, m. burger, j.a. harrison, h.h. braack, p.j. bishop & d. kloepfer (eds.), atlas and red data book of the frogs of south africa, lesotho and swaziland, pp. 246–247, smithsonian institute, washington, dc (si/mab series; no. 9). feely, j.m., 2010, ‘on the southeastern range limits of the nile crocodile: a review of its past and present occurrences in the eastern cape and western cape, south africa’, south african journal of wildlife research 40(2), 169–175. http://dx.doi.org/10.3957/056.040.0209 fitzsimons, v.f.m., 1930, ‘descriptions of new south african reptilia and batrachia, with distribution records of allied species in the transvaal museum collection’, annals of the transvaal museum 14, 20–48. fitzsimons, v.f.m., 1943, ‘the lizards of south africa’, transvaal museum memoirs 1, 1–528. frost, d.r., 2014, amphibian species of the world: an online reference, version 6.0, online database, viewed 24 june 2014, from http://research.amnh.org/herpetology/amphibia/index.html, american museum of natural history, new york. haagner, g.v., 1994, ‘geographical distribution: amblyodipsas concolor’, african herp news 21, 26. hopkins, k.p. & tolley, k.a., 2011, ‘morphological variation in the cape dwarf chameleon (bradypodion pumilum) as a consequence of spatially explicit habitat structure differences’, biological journal of the linnean society 102(4), 878–888. laurent, r.f., 1941, ‘contribution à l’ostéologie et à la systématique des ranides africains, première note [contribution to the osteology and the systematics of african ranidae, primary notes]’, revue de zoologie et de botanique africaines 34, 74–96. marais, j., 2004, a complete guide to the snakes of southern africa, struik, cape town. marais, j., 2014a, ‘crocodylus niloticus laurenti, 1768, nile crocodile’, in m.f. bates, w.r. branch, a.m. bauer, m. burger, j. marais, g.j. alexander et al. (eds.), atlas and red list of of the reptiles of south africa, lesotho and swaziland, pp. 87–88, .south african national biodiversity institute, pretoria. marais, j., 2014b, ‘thelotornis capensis capensis a. smith 1849’, in m.f. bates, w.r. branch, a.m. bauer, m. burger, j. marais, g.j. alexander et al. (eds.), atlas and red data list of the reptiles of south africa, lesotho and swaziland, p. 423, south african national biodiversity institute, pretoria (suricata series; no. 1). measey, g.j., 2011, ensuring a future for south africa’s frogs: a strategy for conservation research, south african national biodiversity institute, petoria (sanbi biodiversity series; no. 19). minter, l.r., 2004a, ‘breviceps bagginsi minter, 2003’, in l.r. minter, m. burger, j.a. harrison, h.h. braack, p.j. bishop & d. kloepfer (eds.), atlas and red data book of the frogs of south africa, lesotho and swaziland, pp. 174–175, smithsonian institute, washington, dc (si/mab series; no. 9). minter, l.r., 2004b. ‘breviceps mossambicus peters, 1854’, in l.r. minter, m. burger, j.a. harrison, h.h. braack, p.j. bishop & d. kloepfer (eds.), atlas and red data book of the frogs of south africa, lesotho and swaziland, pp. 184–186, smithsonian institute, washington, dc (si/mab series; no. 9). minter, l.r., burger, m., harrison, j.a., braack, h.h., bishop, p.j. & kloepfer, d., 2004, atlas and red data book of the frogs of south africa, lesotho and swaziland, smithsonian institute, washington, dc (si/mab series; no. 9). peinke, d. & gibisela, z., 2013, ‘game census report 2013’, unpublished report, eastern cape parks and tourism agency, east london. pickersgill, m., 1996, ‘a new subspecies of afrixalus (anura: hyperoliidae) from kwazulu-natal, south africa, with comments on its superspecies affinities’, durban museum novitates 21, 49–59. pooley, a.c., 1980, ‘crocodile research in maputaland’, in m.n. bruton & k.h. cooper (eds.), studies on the ecology of maputaland, pp. 293–299, wildlife society of southern africa, grahamstown. poynton, j.c., 1964, ‘the amphibia of southern africa: a faunal study’, annals of the natal museum 17, 1–334. raw, l.r.g., 2001, ‘revision of some dwarf chameleons (sauria: chamaleonidae: bradypodion) from eastern south africa’, msc dissertation, school of botany and zoology, university of natal. tarrant, j., 2012, ‘conservation assessment of the threatened amphibians on kwazulu-natal, and a national assessment of the chytrid infection in south african threatened species’, phd thesis, school for biological sciences, north-west university. tolley, k., 2014a, ‘bradypodion melanocephalum (gray, 1865 “1864”)’, in m.f. bates, w.r. branch, a.m. bauer, m. burger, j. marais, g.j. alexander et al. (eds.), atlas and red data list of the reptiles of south africa, lesotho and swaziland, pp. 291–292, south african national biodiversity institute, pretoria (suricata series; no. 1). tolley, k.a., 2014b, ‘bradypodion caffer boettger, 1889’, in m.f. bates, w.r. branch, a.m. bauer, m. burger, j. marais, g.j. alexander et al. (eds.), atlas and red data list of the reptiles of south africa, lesotho and swaziland, p. 288, south african national biodiversity institute, pretoria (suricata series; no. 1). tolley, k.a., 2014c, ‘bradypodion ketanicum hewitt 1935’, in m.f. bates, w.r. branch, a.m. bauer, m. burger, j. marais, g.j. alexander et al. (eds.), atlas and red data list of the reptiles of south africa, lesotho and swaziland, pp. 290–291, south african national biodiversity institute, pretoria (suricata series; no. 1). tolley, k. & burger, m., 2007, chameleons of southern africa, struik, cape town. venter, j.a. & conradie, w., 2012, ‘the amphibians and reptiles of dwesa nature reserve: a report on a rapid biodiversity survey conducted in february 2012’, unpublished report, eastern cape parks and tourism agency, east london. appendix 2 top ↑ list of all amphibian material from the port elizabeth museum examined for the study amphibians arthroleptis wahlbergii: silaka nr (pem a10548–10554), hluleka nr (pem a9967, 10001). leptopelis natalensis: mkambati nr (pem a9565; pem t434), silaka nr (pem a10588–10591), hluleka nr (pem a9979, 9980; pem t497, 505), dwesa nr (pem t535, 541, 542), manubi (pem t640). breviceps cf. bagginsi: silaka nr (pem a10565–10569). breviceps verrucosus: silaka nr (pem a10570). amietophrynus gutturalis: mkambati nr (pem a6908, 9553–9556), silaka nr (pem a10563, 10564; pem t592), hluleka nr (pem a2141, 2142, 6942, 9970, 9971, 9978), dwesa nr (pem a10115, 10130; pem t536, 545), cape morgan nr (pem a10164, 10172). amietophrynus rangeri: mkambati nr (pem a9557–9559), hluleka nr (pem a6936), dwesa nr (pem a3460, 6751, 10126, 10155), manubi sf (pem a10723, 10727, 10728), cape morgan nr (pem a10165, 10175, 10177, 10178). afrixalus spinifrons: silaka nr (pem a10555–10560; pem t593, 632), hluleka nr (pem a9984, 9986). hyperolius marmoratus verrucosus: mkambati nr (pem a6903, 6909, 6910, 9560–9563; pem t428–431, 669), silaka nr (pem a10577–10580), hluleka nr (pem a2147–2151, 6924–6927, 9963, 9964, 9982, 9983, 9990, 9991), dwesa nr (pem a6753, 10116, 10120, 10124, 10125, 10136, 10141; pem t539, 543), manubi sf (pem a10730, 10734, 10735; pem t636, 637), cape morgan nr (pem a10184, 10185; pem t549). hyperolius poweri: mkambati nr (pem a6911–6914, 9545, 9546, 11106, 11107; pem t426, 427). hyperolius pusillus: silaka nr (pem a10581, 10582), hluleka nr (pem a6928–6930, 9985, 9987–9989), dwesa nr (pem a10122, 10135, 10140, 10142, 10148, 10153; pem t538), manubi sf (pem t638). hyperolius semidiscus: silaka nr (pem a10583–10586), hluleka nr (pem a6939–6941, 9995–9997), dwesa nr (pem a10111–11014, 10117–10119, 10121, 10137). kassina senegalensis: mkambati nr (pem a9564; pem t432, 433, 665, 667), silaka nr (pem a10587), hluleka nr (pem a9962, 9969, 9973, 9974, 9977), dwesa nr (pem t537, 546), manubi sf (pem t639), cape morgan nr (pem a10170, 10174, 10186, 10187). phrynobatrachus natalensis: mkambati nr (pem a6917, 9566; pem t440, 441, 669), silaka nr (pem a10595–10597; pem t594), hluleka nr (pem a9959–9961, 9965, 9968; pem t506, 507), dwesa nr (pem a8194–8196, 10127, 10129, 10145, 10151; pem t540), manubi sf (pem a10733), cape morgan nr (pem a10166–10168). xenopus laevis: mkambati nr (pem a9573, 9574; pem t737), silaka nr (pem t599), hluleka nr (pem a9975, 9976), dwesa nr (pem 10138), manubi sf (pem a10731, 10736), cape morgan nr (pem a10179). ptychadena oxyrhynchus: mkambati nr (pem a9567; pem t443–446), silaka nr (pem a10598–10601), hluleka nr (pem a2143, 9981, 9994), dwesa nr (pem a10139; pem t544), cape morgan nr (pem a10182). ptychadena porosissima: mkambati nr (pem a9568–9570; pem t442). amietia quecketti: mkambati nr (pem a7049–7053, 9547–9552; pem t438, 439), silaka nr (pem a10561, 10562), hluleka nr (pem a2144–2146, 7062, 9966, 9998; pem t509), dwesa nr (pem a3953–3955, 10108, 10123, 10128, 10132–10134, 10147; pem t588), manubi sf (pem a10729, 10732; pem t633), cape morgan nr (pem a10180, 10181, 10183). cacosternum nanum: silaka nr (pem a10571–10576; pem t595, 596), hluleka nr (pem a9999, 10000; pem t495, 496, 499, 503, 508), dwesa nr (pem a10143, 10144, 10146, 10149, 10150, 10152, 10154), manubi sf (pem t634, 635), cape morgan nr (pem a10109, 10110, 10163). natalobatrachus bonebergi: silaka nr (pem a10592–10594; pem t597), hluleka nr (pem a9992, 9993; pem t502, 504, 510), manubi sf (pem a10724, 10725; pem t641). strongylopus fasciatus: mkambati nr (pem a6918), silaka nr (pem a10602; pem t598), dwesa nr (pem a4091, 10131), manubi sf (pem a10726), cape morgan nr (pem a10169, 10171, 10173, 10176, 10188). strongylopus grayii: mkambati nr (pem a6922). tomopterna natalensis: mkambati nr (pem a6915, 6916, 9571, 9572; pem t435, 436), hluleka nr (pem a9972, 10002). reptiles crocodylus niloticus: dwesa nr (ecpta). pelomedusa galeata: mkambati nr (pem r19172), silaka nr (ecpta), hluleka nr (ecpta), dwesa nr (pem r19787, 19790), manubi sf (pem r20297). afroedura pondolia: mkambati nr (pem r9711, 12031, 19152), hluleka nr (pem r13377–13379, 19427, 19435, 19436), dwesa nr (pem r13327, 13328, 13448, 13842, 13857, 19783, 19799, 19803). hemidactylus mabouia: mkambati nr (pem r9710, 13367, 19155, 19156), silaka nr (pem r20140–20144), hluleka nr (pem r13375, 13376, 19426, 19440), dwesa nr (pem r19798, 19804), manubi sf (pem r20290), cape morgan nr (pem r19863, 19864, 19873). pachydactylus maculatus: silaka nr (pem r20145, 20146), manubi sf (pem r20291), cape morgan nr (pem a19870). agama atra: manubi st (pem r13008, 20296), cape morgan nr (pem r19872). material: mkambati nr (pem r13370, 19153, 19154, 20686), silaka nr (pem r20133), manubi st (pem r20282, 20283), cape morgan nr (pem r19874, 19877). bradypodion caffer: mkambati nr (pem r20685, 20687–20689). bradypodion ketanicum: dwesa nr (pem r5503, 5505, 17101, 17102, 19789, 19800), manubi sf (pem r16616, 16711). acontias plumbeus: dwesa nr (pem r19811), manubi sf (pem r20279). trachylepis homalocephala: mkambati nr (pem r19157), silaka nr (pem r20151), hluleka nr (pem r19433), dwesa nr (pem r19807), manubi sf (pem r20299, 20300), cape morgan nr (pem r19868, 19871). trachylepis varia: mkambati nr (pem r13368, 19158–19161), silaka nr (pem r20149, 20150), hluleka nr (pem r19428, 19434, 19452), dwesa nr (pem r13779, 19788, 19801), manubi sf (pem r20301, 20302), cape morgan nr (pem r19875, 19876). tropidosaura montana natalensis: mkambati nr (pem r19173). chamaesaura anguina anguina: hluleka nr (pem r19443), dwesa nr (pem r19802). gerrhosaurus flavigularis: silaka nr (pem r20138, 20139), dwesa nr (pem r19797, 19809, 19810), cape morgan nr (pem r19869). tetradactylus cf. fitzsimonsi: hluleka nr (pem r19424, 19425, 19453, 19456). varanus niloticus: mkambati nr (pem r20864), silaka nr (ecpta), hluleka nr (pem r19451), dwesa nr (pem r13329, 19808), cape morgan nr (ecpta). leptotyphlops sylvicolus: mkambati nr (pem r18079, 19164, 19165), hluleka nr (pem r19439). afrotyphlops bibronii: silaka nr (pem r20130–20132), hluleka nr (pem r19429, 19430, 19438), dwesa nr (pem r13693, 13836, 19805), manubi sf (pem r20280, 20281). crotaphopeltis hotamboeia: mkambati nr (pem r19163), hluleka nr (pem r19444), dwesa nr (pem r19796), manubi sf (pem r20287) dasypeltis inornata: hluleka nr (pem r19445), dwesa nr (pem r13692). dispholidus typus typus: hluleka nr (pem r19447, 19448), dwesa nr (pem r13725, 19785). philothamnus hoplogaster: hluleka nr (pem r13690). philothamnus natalensis occidentalis: mkambati nr (pem r19167, 19168), silaka nr (pem r20147, 20148), cape morgan nr (pem a19878). python natalensis: mkambati nr (ecpta). amblyodipsas concolor: hluleka nr (pem r19437), dwesa nr (pem r19795), manubi sf (pem r20284, 20285). boaedon capensis: mkambati nr (pem r13369), silaka nr (pem r20135), hluleka nr (pem r19454, 19455). duberria lutrix lutrix: silaka nr (pem r20137), hluleka nr (pem r19431, 19432), dwesa nr (pem r13724), manubi sf (pem r20288, 20289), cape morgan nr (pem r19866). lycodonomorphus rufulus: dwesa nr (pem r19792, 19793), manubi sf (pem r20292, 20295). lycophidion capense: mkambati nr (pem r19166), dwesa nr (pem r19794). macrelaps microlepidotus: dwesa nr (pem r19786, 19791), manubi sf (pem r20294, 20295). psammophis brevirostris: mkambati nr (pem r19169–19171), hluleka nr (pem r19446), manubi sf (pem r20298). bitis arietans: silaka nr (pem r20134). causus rhombeatus: mkambati nr (pem r19162), silaka nr (pem r20136), hluleka nr (pem r19449, 19450), dwesa nr (pem r19806), manubi sf (pem r20286), cape morgan nr (pem r13714, 19865, 19867). appendix 3 top ↑ additional records used in compiling the final checklist of amphibians and reptiles of the wild coast reserves tm, ditsong national museum of natural history (formerly transvaal museum); cas, california academy of science; dmr, durban natural science museum. amphibians top ↑ arthroleptis wahlbergii: hluleka nr (safap 116500, 141009), silaka nr (safap 14100). leptopelis natalensis: dwesa nr (safap 116240), hluleka nr (safap 141006), mkambati nr (safap 116162, 116226), silaka nr (safap 116243; cas 11606, 211607). breviceps bagginsi: mkambati nr (safap 116148, 116742, 117408), silaka nr (safap 116147). breviceps verrucosus: mkambati nr (safap 116743, 117409), silaka nr (safap 116146). amietophrynus gutturalis: hluleka nr (safap 141004), mkambati nr (safap 116154, 116161, 116223, 116744), silaka nr (safap 116247, 140996). amietophrynus rangeri: cape morgan nr (safap 115819, 115821), dwesa nr (safap 117285). hyperolius marmoratus verrucosus: cape morgan nr (safap 115817, 115822), dwesa nr (safap 117289, 116238), hluleka nr (cas 21162, cas 211611), manubi sf (safap 115772), mkambati nr (safap 116157, 116164, 116224, 116745), silaka nr (safap 116244, 140998). hyperolius poweri: mkambati nr (safap 116158, 116746). hyperolius semidiscus: cape morgan nr (safap 115823), hluleka nr (cas 211609, 211610), mkambati nr (safap 116165), silaka nr (safap 115618, 116245). kassina senegalensis: cape morgan nr (safap 115825), manubi sf (safap 115774), mkambati nr (safap 116155). phrynobatrachus natalensis: cape morgan nr (safap 115818), mkambati nr (safap 116160, 116163, 116227), silaka nr (cas 211604, 211605,). xenopus laevis: dwesa nr (safap 116219, 117460). ptychadena oxyrhynchus: mkambati nr (safap 116156, 116220, 117415), silaka nr (safap 116242). ptychadena porosissima: mkambati nr (safap 116221, 117416). amietia quecketti: cape morgan nr (safap 115740), dwesa nr (safap 127053), hluleka nr (safap 141005), mkambati nr (safap 116748, 117417, 117418, 117458, 117459), silaka nr (safap 115616, 140999). cacosternum boettgeri: mkambati nr (safap 11652). cacosternum nanum: cape morgan nr (safap 115820, 115824), hluleka nr (safap 117439), manubi sf (safap 115773), mkambati nr (safap 116149, 116152, 116166). natalobatrachus bonebergi: silaka nr (safap 116246), dwesa nr (safap 116239). strongylopus fasciatus: dwesa nr (safap 127052), hluleka nr (safap 141007), mkambati nr (safap 116153, 116222). tomopterna natalensis: mkambati nr (safap 116150, 116159, 116217, 116225). reptiles afroedura pondolia: silaka nr (tm 76520), dwesa nr (tm 69607–69621). hemidactylus mabouia: manubi sf (tm 69605, 69606). agama atra: mkambati nr (dmr1124). bradypodion caffer: silaka nr (sarca 100520, 100521). bradypodion ketanicum: dwesa nr (sarca 100721). acontias plumbeus: dwesa nr (sarca 129704). dasypeltis inornata: hluleka nr (sarca 1576). dispholidus typus: dwesa nr (sarca 4373). lycophidion capense: mkambati nr (dmr1107). appendix 4 top ↑ table 1-a4: a summary of the locations, habitats and trapping dates where drift fence with pitfall or funnel combinations were set in the six protected areas along the wild coast, eastern cape, south africa. figure 1-a4: (a–ai) habitat type and trap layout of the respective trap sites surveyed during this study. figure 1-a4 (continues…): (a–ai) habitat type and trap layout of the respective trap sites surveyed during this study. figure 1-a4 (continues…): (a–ai) habitat type and trap layout of the respective trap sites surveyed during this study. figure 1-a4 (continues…): (a–ai) habitat type and trap layout of the respective trap sites surveyed during this study. figure 1-a4 (continues…): (a–ai) habitat type and trap layout of the respective trap sites surveyed during this study. article information authors: casparus j. crous1 james s. pryke1 michael j. samways1 affiliations: 1department of conservation ecology and entomology, stellenbosch university, south africa correspondence to: casper crous email: cjcrous@gmail.com postal address: private bag x1, matieland 7602, south africa dates: received: 09 jan. 2015 accepted: 08 july 2015 published: 31 aug. 2015 how to cite this article: crous, c.j., pryke, j.s. & samways, m.j., 2015, ‘conserving a geographically isolated charaxes butterfly in response to habitat fragmentation and invasive alien plants’, koedoe 57(1), art. #1297, 9 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1297 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. conserving a geographically isolated charaxes butterfly in response to habitat fragmentation and invasive alien plants in this original research... open access • abstract • introduction • research method and design    • study area    • design    • procedure       • behavioural observations       • landscape variables    • analyses • results • discussion    • the presence of charaxes xiphares occidentalis across this fragmented landscape    • tree community settling patterns of charaxes xiphares occidentalis in a heterogeneous landscape • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ in south africa, much of the forest biome is vulnerable to human-induced disturbance. the forest-dwelling butterfly charaxes xiphares occidentalis is naturally confined to a small forest region in the south-western cape, south africa. most of the remaining habitat of this species is within a fragmented agricultural matrix. furthermore, this geographical area is also heavily invaded by alien plants, especially acacia mearnsii. we investigated how c. x. occidentalis behaviourally responds to different habitat conditions in the landscape. we were particularly interested in touring, patrolling and settling behaviour as a conservation proxy for preference of a certain habitat configuration in this agricultural matrix. remnant forest patches in the agricultural matrix showed fewer behavioural incidents than in a reference protected area. moreover, dense stands of a. mearnsii negatively influenced the incidence and settling pattern of this butterfly across the landscape, with fewer tree settlings associated with more heavily invaded forest patches. this settling pattern was predominantly seen in female butterflies. we also identified specific trees that were settled upon for longer periods by c. x. occidentalis. distance to a neighbouring patch and patch size influenced behavioural incidences, suggesting that further patch degradation and isolation could be detrimental to this butterfly. conservation implications: we highlight the importance of clearing invasive tree species from vulnerable forest ecosystems and identify key tree species to consider in habitat conservation and rehabilitation programmes for this butterfly. we also suggest retaining as much intact natural forest as possible. this information should be integrated in local biodiversity management plans. introduction top ↑ in south africa, the forest biome is the smallest of all biomes (< 1% surface cover) and is highly sensitive to environmental disturbances (eeley, lawes & piper 1999). as many people continue to rely on native forests for their livelihoods, this biome is one of the most vulnerable and threatened vegetation types in the country (shackleton et al. 2007). in these forests is a charismatic butterfly species: the forest king emperor (nymphalidae: charaxinae: charaxes xiphares), a large and fast-flying butterfly. moreover, this species has nine subspecies, all of which are geographically distinct (woodhall 2005). unfortunately, fruit-feeding butterflies are often highly sensitive to forest disturbance (fermon et al. 2000; uehara-prado, brown & freitas 2007). little information is available on forest-dwelling butterfly ecology in the threatened south african forest biome. the south-western cape region of south africa has many small natural forest patches that today are mainly within a cultivated agricultural landscape. many of these forest patches have been reduced in size to provide pasture and fodder for cows. as a result, few of the forest patches remain intact, especially with regard to their original size. within this region lies an important protected area, namely the grootvadersbosch nature reserve. here, the subspecies c. x. occidentalis pringle is endemic. this is the most southern subspecies of c. xiphares (woodhall 2005). this protected area and many of the fragmented forest patches are situated in mainly a dairy farming matrix. these farms are part of the grootvadersbosch conservancy, although there is no conservation plan (other than the formally protected area) to protect the habitat of this highly localised butterfly population. many of the remnant forest patches in the grootvadersbosch conservancy are along rivers and streams, which bring them into contact with invasive alien plant species, another threat to the biodiversity in this landscape (hood & naiman 2000). south african river systems are particularly threatened, with less than a third of the main rivers currently ecologically intact (nel et al. 2007). these riverbanks are heavily invaded by the alien tree acacia mearnsii de wild (fabaceae). acacia mearnsii trees often form homogenous stands within catchment areas, which severely alter stream hydraulic characteristics across the landscape, and ultimately negatively affect plant community structure and biodiversity in the catchment area (dye & jarmain 2004; enright 2000; le maitre et al. 2002, 2011). biotic homogenisation is strongly associated with anthropogenic mass extinction events (mckinney & lockwood 1999). production landscapes are particularly prone to these events, as biodiversity reductions associated with such landscapes often increase the invasion potential of alien species (fargione & tilman 2005). therefore, managing forest-dwelling butterflies effectively in a forest–agricultural matrix will require a deeper understanding of the role that natural environmental heterogeneity plays in butterfly dispersal patterns (hamer et al. 2003). for example, preserving heterogeneity in native tree canopy structure and prohibiting invasive tree species from forming dense stands within native forests are important for conserving fruit-feeding butterflies (hamer et al. 2003). indeed, many butterflies are sensitive to changes in habitat conditions due to alien plant invasions (edge 2005; florens et al. 2010; hanula & horn 2011; lawrence & samways 2002; lu & samways 2002; but see graves & shapiro 2003). as invasive acacia trees in south africa severely degrade the local plant community structure (le maitre et al. 2011), which could include reducing the presence or density of the butterfly's larval and adult feeding plants, the effect of a. mearnsii on the dispersal of c. x. occidentalis in this increasingly fragmented landscape requires understanding. the relevance of this information is underscored by there being no evidence that the larvae or adults of this charaxes subspecies utilise fabaceae plants as a habitat resource. as there is currently no research on the response of c. x. occidentalis to disturbed southern afrotemperate forest, it is essential to identify the landscape variables that can contribute to this butterfly's conservation in this production landscape. behavioural studies can be particularly useful to identify these preferences (new 1991; shreeve 1984; thomas 1984). behavioural studies can reveal specific habitat requirements needed for a species or community, and subsequently how to optimally conserve their habitat or biotope (crous, samways & pryke 2014; sutherland 1998; turlure et al. 2011). understanding how different resource conditions influence butterfly behaviour is crucial for understanding their distribution patterns across space and time (shreeve & dennis 2011). in this study we use behavioural observations to determine the response of c. x. occidentalis to remnant patches of southern afrotemperate forest embedded in a dairy farming matrix. furthermore, we examine the effect that a. mearnsii invasion has on aspects of the butterfly's behaviour. our key questions related to effective conservation are therefore two tiered. at one level, we aim to determine (1) the activity (number of behavioural incidences) of adults of c. x. occidentalis in remaining forest fragments with different invasion levels, (2) the activity of adults of this species in the protected area vis-à-vis in the forest fragments and (3) which environmental variables may help explain the different behavioural activities across the landscape. at the second level, we aim to establish (1) whether there is a consistent native tree community upon which the adult butterfly population shows a positive settling response (with settling as a conservation proxy for preference) and (2) which tree species are especially important to this butterfly within the fragmented forest fragments. research method and design top ↑ study area we recorded the behaviour of c. x. occidentalis at 11 sites in the grootvadersbosch conservancy and the grootvadersbosch nature reserve (the latter abbreviated here to pa for ‘protected area’ to distinguish this reference site from the forest fragments in the conservancy), western cape, south africa (figure 1). the 2.5-km2 pa is considered the second-largest patch of southern afrotemperate forest, preceded only by the considerably larger knysna–tsitsikamma forest (narrow coastal strip of 250 km), about 100 km east of this pa. the langeberg mountain range runs north of both the closest town and the pa. the geology of the langeberg mountain range is mostly table mountain sandstone and forms part of the greater cape fold belt. patches of southern afrotemperate forest occupy the wetter southern slopes of this mountain range. this forest vegetation type occurs at elevations between 10 m a.s.l. near its coastal distribution range and 1060 m a.s.l. at the highest measured peak (mucina & rutherford 2006). all sites were situated on the south-facing lower slopes of this mountain range. the elevation range of the study sites was 115 m a.s.l. – 324 m a.s.l., with an average elevation of 190 m a.s.l. the grootvadersbosch region lies in the transitional zone between the winter and all-year rainfall regions, with up to 1050 mm of mean annual precipitation recorded for the pa. the existing remnant afrotemperate forest patches within the dairy farming matrix of the conservancy are mostly restricted to rivers or streams. figure 1: location of study sites within the grootvadersbosch conservancy, near the town of suurbraak in the western cape province, south africa. design we selected 11 sites along four river systems that flow through the catchment area (figure 1). the duivenhoks river, which is a small stream that also flows through the pa (not indicated on the map), flows towards the south-east. one site (pa1) was along this river. we focus on forest along these rivers because of the remnant patches here still being intact. as so little is known about the presence of the butterfly across this fragmented landscape, we also focused, as a theoretical starting point, on forest patches where at least scutia myrtina, the larval food plant of c. x. occidentalis, was present (woodhall 2005). of the 11 selected sites, two were along streams in the 250-ha patch of continuous forest that constitutes the pa (sites were 2398 m apart). six sites were remnant forest patches in the conservancy of the agricultural matrix (natural fragments; nf), which we classified as patches that showed little or no indication of disturbance, either morphologically (riverbank structure) or biologically (alien plant species). these sites were between 319 m and 9182 m apart. lastly, we selected three remnant forest patches invaded by a. mearnsii, also in the conservancy agricultural matrix and interspersed with the natural fragments (invaded fragments; if). the riverbanks here were degraded morphologically. specifically, an invaded site was where > 75% of the trees were a. mearnsii (homogenous stands). the latter sites were between 1031 m and 2829 m apart. the variable number of studied sites in each of the mentioned categories was a factor of the availability of similar sites across this small region (a discontinuous forest–agriculture matrix; see uehara-prado et al. 2007). for example, the ifs were not consistent across the area owing to some land users actively weeding invasive alien plants whereas others do not. this also means that the sizes of the studied remnant forest fragments differed. however, to standardise the sampling in this case, we surveyed and compared a fixed forest fragment area where at least the butterfly's larval food plant was present. procedure behavioural observations behaviour of both sexes of the butterfly was observed at each of the 11 sites by the same two observers in january 2014. the study period was during the peak flying season of this butterfly. observation took place between 10:00 and 14:00 on warm or hot days with < 10% cloud cover (average temperature: 30.4 °c ± 2.6 sd). eight hours of total observation time were spent per site. behaviour recording started as soon as a c. x. occidentalis individual entered a circle of a 10-m radius and ended when the butterfly left the circle. a 10-m radius was used for each observer to observe at least a 20-m strip of forest trees in the remaining habitat. as proxies for the response of this butterfly to its immediate environment, only the following behavioural observations were recorded: patrolling – the individual flies low over the area at low speed, as if investigating or searching touring – the individual flies high over the area and at high speed, without showing any searching or investigating behaviour settling – the individual is settled on an object; settling may integrate other behaviours, such as resting, basking, feeding or ovipositing. the amount of time an individual exercised a particular behaviour was also recorded. landscape variables we chose four landscape variables for correlation with the behavioural observations: tree species richness in the sampled belt size of the patch of native forest in which the sampling belts occurred distance of the fragments to the pa distance of the fragments to the nearest large protected patch on a land-user property. the latter variable refers to a group of isolated forest patches on a farm, extending more inland or up the mountain as opposed to patches mainly being riparian, and ultimately present owing to ongoing tree protection efforts by the land user (non-nature reserve or informal protection; in this case an approximately 20-ha patch). for this variable, the pa could also be considered a protected patch on a large land-user property in this landscape. prior to butterfly sampling, tree species richness of each fragment (identified to at least genus level) was estimated by two trained observers using a 50-m × 10-m belt (50 m parallel to the river-facing forest edge and < 10 m into the forest). at nf5, however, the forest did not always extend past 10 m owing to uneven fragmentation (ranging between 8 m and 15 m). we used the 10-m cut-off perpendicular to the stream to complement the 10-m radius of the butterfly sampling circle. this variable was chosen to determine whether decreasing tree richness in the fragments has an effect on the butterfly activity in the area. tree identifications were in random directions (non-linear searches) within each sampled belt, and it took up to an hour to identify all species. the distance of each forest patch to the pa and the two largest and actively conserved forest patches was calculated from images from google earth (google, inc., mountain view, usa). analyses statistical analysis software (statistica version 12, statsoft, tulsa, usa) was used for initial statistical analyses. to observe the number of behavioural incidences recorded between the pa and the nfs and ifs (non-nature reserve sites), a bar graph was constructed showing standard deviations. to compare the mean number of behavioural incidences recorded in each behavioural category across these non-nature reserve sites, we used one-way analyses of variance. a shapiro–wilk test showed that values in each category were normally distributed and their variances were homogenous. however, quantile–quantile plots suggested that a non-parametric analysis would be more appropriate for the touring category. non-parametric mann–whitney u tests were performed to establish whether there were sex-specific responses in each behavioural class across the nfs and ifs. this test was performed as male settling, female settling and female touring behaviour values were not normally distributed. a spearman non-parametric rank correlation matrix was constructed to determine the possible relationship between the total number of behavioural incidences observed for each behavioural trait and the four measured environmental variables. this test was chosen because some of the environmental variables were not normally distributed. in addition to behavioural incidence patterns, we also determined which community of trees c. x. occidentalis spent the most time settling upon, and whether this was consistent across sites, using a non-metric multidimensional scaling (nmds) ordination in primer 6 (primer-e, lutton, uk). nmds is a non-linear, distance-based technique that ranks different samples or groups in ordination space according to their similarity/dissimilarity (ter braak 1995). for this particular nmds, we used time (seconds) as a measure of the ‘abundance’ of the time individual butterflies spent on different trees within a site. we excluded s. myrtina (the known larval food plant) from this analysis. this ensured that we focused on trees used by c. x. occidentalis for reasons other than ovipositing. a bray–curtis similarity matrix (which takes into account abundance) was used and data were fourth-root transformed. nmds therefore indicated the butterfly's tree community settling pattern (tcsp) across the study area. furthermore, to determine which tree species strongly structured differential tree community settling (i.e. were settled upon for longer periods), we vectored (correlated) those tree species with pearson correlations (r) > 0.8 onto the nmds axes in primer 6. we specifically chose the strongest possible correlation to ensure that we identify the tree species that most strongly influences the variation in settling behaviour of this butterfly amongst the studied sites. to test for possible spatial autocorrelation between tree use activity patterns and distance between sites (i.e whether closer sites are more alike in tree activity use), we used the relate function in primer 6. this is a mantel-type test that correlates two similarity matrices with each other. to obtain the between-site distance matrix, we used the geographic distance matrix generator v.1.2.3 (ersts n.d.). for this relate test, we used spearman's rho with 9999 permutations. results top ↑ the highest number of behavioural incidences recorded was in the pa (figure 2). in total, and for each behavioural category, there was a general, stepwise decrease in the number of behavioural incidences recorded when scaling from the pa to the ifs (figure 2). the more intact forest patches also had the highest number of individuals entering a sampling circle, of which most were female (table 1). specifically, the more intact fragments had generally more behavioural incidences than the invaded fragments (f = 5.18, p = 0.057). this trend was mainly due to the significantly higher settling behaviour in the nfs than in the ifs (f = 7.98, p = 0.026; table 2). notably, this overall higher settling behaviour in the nfs was specifically a function of female settling behaviour (z = 2.21, p = 0.027; table 3). neither touring behaviour nor patrolling behaviour differed significantly between natural and invaded patches (table 2). figure 2: means (±sd) of total observed butterfly behavioural incidences, as well as the incidences in each behavioural classification, in the grootvadersbosch nature reserve forest (n = 2), the natural forest fragments (n = 6) and the forest fragments invaded by acacia mearnsii (n = 3). table 1: the average number of charaxes xiphares occidentalis individuals that entered a sampling radius (as a proxy for abundance) from which behavioural data were recorded, as well as their sexual delineation. table 2: results of one-way analyses of variance and a mann–whitney u test to compare the number of incidences in all and within each studied butterfly behavioural classification across natural and invaded forest patches in a dairy farming matrix. table 3: results of mann–whitney u tests of sex-specific behavioural incidences within each studied behavioural classification in natural and invaded forest patches in a dairy farming matrix. settling incidences (as a conservation proxy for resource preference in a site) and patrolling incidences were both positively correlated with tree species richness (rs = 0.90 and rs = 0.75, respectively) and patch size (rs = 0.92 and rs = 0.75, respectively). however, settling and patrolling incidences were negatively correlated with distance to a large protected forest patch at a land user in the landscape (rs = −0.71 and rs = −0.67, respectively) (n = 11; table 4). in turn, touring behaviour was significantly correlated only with patrolling (n = 11; rs = 0.62; table 4). patch size was positively correlated with tree species richness (rs = 0.86) and the amount of time an individual butterfly settled in the patch (rs = 0.62; table 4). table 4: a rank correlation matrix of spearman rank correlations (rs) to compare the relationships between measured environmental variables and behavioural traits across all sites (n = 11). there was no spatial autocorrelation between the tcsp of c. x. occidentalis and the distances between sites (rs = −0.054, p = 0.596). this suggests that in situ patch conditions in this particular landscape are more important for determining longer settling behaviour than the proximity of sites to one another. furthermore, there was a strong tendency for different tree selection by individual butterflies under certain site conditions (nmds, 2d stress 0.05; figure 3). in particular, the two pa sites grouped together, with olea capensis and canthium inerme being the tree species that were most strongly correlated with driving tcsp. notably, two nfs and an if also had a more similar adult tcsp. here, the settling time of c. x. occidentalis on buddleja saligna structured this site grouping (figure 3). across all sites, the most consistent settling behaviour was on s. myrtina (table 5), the butterfly's larval food plant that was present in all studied sites. this confirms the importance of this species for conservation. there was only one incident of a male individual settling on an a. mearnsii tree (for 2 min) during the entire sampling period (table 5). figure 3: non-metric multidimensional scaling ordination depicting the observed tree community settling activity of charaxes xiphares occidentalis across all study sites. the indicated tree species had the strongest influence (r > 0.8) in structuring the observed dissimilarity in tree community settling pattern across the study area. table 5: average settling time (seconds) of charaxes xiphares occidentalis individuals on trees at 11 sites in the grootvadersbosch conservancy. discussion top ↑ the presence of charaxes xiphares occidentalis across this fragmented landscape it was encouraging to see that this butterfly species was recorded in all the remnant patches here, despite invasion by a. mearnsii. however, it was clear that the number of individuals entering a sampling radius and the total recorded behavioural incidences were fewer in fragments invaded by a. mearnsii. this was especially evident when directly comparing the invaded sites with the intact and formally protected sites, where > 3 times the number of behavioural incidences were observed. there were also more behavioural incidences in the pa sites than in the natural remnants. so, although c. x. occidentalis was present at all sites, it was more active at the more physically and biologically intact sites. this variance in activity patterns can be attributed to the tree settling incidences, recorded here as an indication of resource preference in their immediate habitat, and which were significantly higher in the nfs than the ifs. this result supports findings from further north in the country, where dense stands of a. mearnsii negatively affected the behaviour of butterflies, with individuals mostly touring over invaded areas (lawrence & samways 2002). these results are also consistent with a study on lupine-invaded areas in finland, where higher alien plant cover resulted in more fly-over (touring) behaviour amongst butterfly individuals, resulting in fewer settling and feeding events (valtonen, jantunen & saarinen 2006). however, the overall higher settling response in nfs observed in this study was primarily a female-driven pattern. thus, the fragmented nature of the remnant patches, aggravated by the specific presence of dense stands of a. mearnsii trees, is particularly negative towards habitat suitability for female individuals of c. x. occidentalis. the mechanism as to how and why a. mearnsii trees negatively affected the behaviour of female c. x. occidentalis is related to the strong positive correlation between native tree species richness, size of the remnant patch and activity incidences. it is likely that more female c. x. occidentalis individuals patrol the larger intact forest patches with higher native tree species richness and subsequently settle on a tree. in south african riparian areas, a. mearnsii often forms dense stands. this essentially almost homogenises riverbanks, with the consequence of decreased biodiversity (dye & jarmain 2004; enright 2000; le maitre et al. 2002). as the presence of the host plant, as well as its density, will affect female butterfly behaviour in the landscape (rutowski 1991), the current (and expanding) dense stands of a. mearnsii trees in this catchment will inevitably curb host plant presence and density. furthermore, alien plants may reduce the presence of certain specialist canopy-dwelling butterfly species (ghazoul 2004). homogenisation of gallery forest canopy structure in the remaining natural patches may therefore also be a major contributor to c. x. occidentalis settling less often in such environments, as heterogeneity in the tree canopy cover is often critical for maintaining butterfly species in an area (francesconi et al. 2013; koh & sodhi 2004). essentially, certain forest fragments, and especially those heavily invaded by a. mearnsii trees, seem to create matrix resistance (ricketts 2001) regarding the settling (resource use) activity of female c. x. occidentalis. as there was only one incidence of a male individual settling on an a. mearnsii tree, it means, theoretically, that the resource base of suitable forest habitat for this butterfly is declining in this agricultural landscape. we therefore support the view on the conservation importance of understanding the role of biotope structure and its changes for shaping butterfly movement across space and time (dover & settele 2009). notably, distance of sites to the extensive and intact pa (grootvadersbosch nature reserve) was not correlated with higher behavioural incidences in c. x. occidentalis. however, it was remarkable how important the proximity of a forest remnant to a larger and land-user-protected forest patch was to the number of settling and patrolling incidences. the closer a large area of protected forest at a land user was to a site (considering that the pa also acted as such a patch), the more active individuals of c. x. occidentalis were. however, baguette and van dyck (2007) caution against extrapolating from simple euclidean distance for describing species behaviour patterns. still, there are some important precautionary conservation management implications for this butterfly in this unique agricultural matrix: it seems that heavily invaded fragments further away from a large intact and managed patch in the landscape (more isolated), negatively affect the behaviour of c. x. occidentalis. this finding should be taken into account for the conservation management priority of such spatially distributed forest remnants. tree community settling patterns of charaxes xiphares occidentalis in a heterogeneous landscape to determine whether this butterfly has a specific tcsp, we also recorded the amount of time that each individual spent settling on a tree. in a sense, settling functions as a conservation or management proxy for preference to a resource in an individual's immediate environment, as butterfly settling behaviour is strongly associated with a variety of life history strategies such as feeding, ovipositing, basking, resting and mate location (dennis 2004, 2010; dennis, shreeve & van dyck 2003, 2006). within the limits of this summer-time observation, there was a clear dissimilarity in tcsp between sites, with the two pa sites showing similar tree use activity, as did a group of nf and if patches. in the latter group, it was the settling time of individuals on b. saligna that most strongly affected dissimilarities in the tcsp. buddleja saligna is an endemic southern african tree species (2 m – 7 m tall) and a forest margin species when associated with forest (coates palgrave 2005). charaxes xiphares occidentalis feeds on the stem sap of this tree (figure 4). such branch-sap feeding behaviour has also been observed for two charaxes species native to the north-eastern parts of south africa (forrester 2011). indeed, charaxes butterflies are considered frugivorous, feeding on the sap of rotting fruit, but have also been shown to feed on a variety of surface-oozing liquids, such as from tree branches (krenn, zulka & gatschnegg 2001; molleman et al. 2005), or decaying animal matter and faeces (molleman & midgley 2009). therefore, b. saligna interaction seems to be an important habitat resource for c. x. occidentalis in disturbed forest fragments, especially in remnants that are now essentially isolated forest edges. figure 4: a male charaxes xiphares occidentalis individual feeding on the stem sap of a non-forest inhabiting buddleja saligna tree. although there is no information on the particular charaxes–buddleja interaction in our study, this finding is consistent with buddleja species being important as an adult butterfly food source (andersson 2003; bruner et al. 2006; corbet 2000). for example, two swedish nymphalidae species showed positive antennal behaviour towards floral scent compounds of buddleja davidii, which suggests that these compounds signal food availability (andersson 2003). as this plant species is not indigenous to sweden, the response perhaps emphasises the theoretical base of local adaptations by some butterflies to varying resource conditions in space and time (schlaepfer et al. 2005). in comparison, a study comparing fruit-feeding butterflies between fragmented and continuous brazilian forests highlighted how brazilian charaxinae species are likely able to adapt to forest fragments as they are opportunist feeders of edge-associated plant species (uehara-prado et al. 2007). the relevance of buddleja spp. as a known adult butterfly food source – being edge inhabitants – and the tcsp investigated here underscore that this plant must be considered in conservation planning of remnant forest fragments. the tcsp of c. x. occidentalis in southern afrotemperate forest in this dairy farming matrix suggests that at least two specific tree species are of importance as a habitat resource of this butterfly when active in forest fragments: s. myrtina as a larval food plant, which was the only species consistently settled upon across all sites (table 5), and b. saligna, as an edge-specialist adult food plant. however, this butterfly did settle on a larger variety of indigenous trees in the pas, most notably on the trees c. inerme and o. capensis, emphasising the butterfly's current need to have both intact forest and edges for two different resource behaviours in this fragmented landscape. of course, there might be other species that could also be of significance but were not observed during this limited summer period; thus the best strategy would be simply to observe the precautionary principle and maintain as much intact forest as possible (nyafwono et al. 2014a). conclusion top ↑ native forests in south africa are sensitive to environmental change due to high commodity production activity and, given their isolation, require conservation efforts that are focused on advanced matrix management strategies (eeley et al. 1999). whilst we can now confirm that the remnant patches of southern afrotemperate forest across this conservancy are still suitable environments for the naturally restricted c. x. occidentalis, the settling patterns of this butterfly vary greatly, especially for the female individuals. this is likely because of certain habitat conditions to which they show a decrease in behavioural activity. first, dense stands of a. mearnsii may limit niche dynamics (see holt 2009), as there was no clear evidence in this study that the butterfly uses a. mearnsii trees in this catchment as a resource. this emphasises the importance of the national initiative to clear this invasive tree species from the river systems in the country (gerber et al. 2008; hanula & horn 2011; le maitre et al. 2011). second, there seem to be specific native tree species that are more often visited by adults of c. x. occidentalis, which also vary across this fragmented forest landscape. this is a positive response of c. x. occidentalis to different forest habitat structure. however, more information is needed regarding the specific behaviour exercised on these trees (basking or resting, as opposed to merely settling), as this would help determine the specific niche a tree provides to this butterfly. nonetheless, this initial identification of utilised trees now allows better focus on the conservation management and rehabilitation processes for the forest patches in which c. x. occidentalis dwells, which should involve habitat rehabilitation through establishing native plants as well as removing invasive alien plants (galatowitsch & richardson 2005; nyafwono et al. 2014b). to achieve the current set of milestones for sustainable farming, for example the convention on biological diversity's aichi 2020 biodiversity targets, a resource-based butterfly conservation strategy based on behavioural ecology has great potential in the farm-scale conservation planning process globally (crous et al. 2014; dennis 2010; dennis et al. 2003, 2006; shreeve & dennis 2011). acknowledgements top ↑ this research was undertaken under a western cape nature conservation board (capenature) research permit (no. 0056-aa008-00023). we thank j. groenewald for field assistance, as well as the following land users who permitted access to their estates: h. barry, p. groenewald and k. moody. the department of science and technology and national research foundation of south africa funded this project. we also thank three anonymous reviewers for their constructive comments. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions c.j.c. (stellenbosch university) and m.j.s. (stellenbosch university) conceived the study. c.j.c. was responsible for the sampling and j.s.p. (stellenbosch university) contributed to the analysis. all three authors were involved in interpretation of the results, and all contributed to writing the manuscript. references top ↑ andersson, s., 2003, ‘antennal responses to floral scents in the butterflies inachis io, aglais urticae (nymphalidae), and gonepteryx rhamni (pieridae)’, chemoecology 13, 3–20. http://dx.doi.org/10.1007/s000490300001 baguette, m. & van dyck, h., 2007, ‘landscape connectivity and animal behavior: functional grain as a key determinant for dispersal’, landscape ecology 22, 1117–1129. http://dx.doi.org/10.1007/s10980-007-9108-4 bruner, l.l., eakes, d.j., keever, g.j., baier, j.w., whitman, c.s., knight, p.r. et al., 2006, ‘butterfly feeding preferences for buddleja selections in the landscape’, journal of environmental horticulture 24, 39–44. coates palgrave, m., 2005, keith coates palgrave: trees of southern africa, struik, cape town. corbet, s.a., 2000, ‘butterfly nectaring flowers: butterfly morphology and flower form’, entomologia experimentalis et applicata 96, 289–298. http://dx.doi.org/10.1046/j.1570-7458.2000.00708.x crous, c.j., samways, m.j. & pryke, j.s., 2014, ‘rockiness determines meso-scale conservation of butterflies in afro-montane grassland’, journal of insect conservation 18, 77–83. http://dx.doi.org/10.1007/s10841-014-9616-x dennis, r.l.h., 2004, ‘butterfly habitats, broad-scale biotope affiliations, and structural exploitation of vegetation at finer scales: the matrix revisited’, ecological entomology 29, 744–752. http://dx.doi.org/10.1111/j.0307-6946.2004.00646.x dennis, r.l.h., 2010, a resource-based habitat view for conservation: butterflies in the british landscape, wiley-blackwell, chichester. dennis, r.l.h., shreeve, t.g. & van dyck, h., 2003, ‘towards a functional resource-based concept for habitat: a butterfly biology viewpoint’, oikos 102, 417–426. http://dx.doi.org/10.1034/j.1600-0579.2003.12492.x dennis, r.l.h., shreeve, t.g. & van dyck, h., 2006, ‘habitats and resources: the need for a resource-based definition to conserve butterflies’, biodiversity and conservation 15, 1943–1966. http://dx.doi.org/10.1007/s10531-005-4314-3 dover, j. & settele, j., 2009, ‘the influences of landscape structure on butterfly distribution and movement: a review’, journal of insect conservation 13, 3–27. http://dx.doi.org/10.1007/s10841-008-9135-8 dye, p. & jarmain, c., 2004, ‘water use by black wattle (acacia mearnsii): implications for the link between removal of invading trees and catchment streamflow response’, south african journal of science 100, 40–44. edge, d.a., 2005, ‘butterfly conservation in the southern cape’, metamorphosis 16, 28–46. eeley, h.a.c., lawes, m.j. & piper, s.e., 1999, ‘the influence of climate change on the distribution of indigenous forest in kwazulu-natal, south africa’, journal of biogeography 26, 595–617. http://dx.doi.org/10.1046/j.1365-2699.1999.00307.x enright, w.d., 2000, ‘the effect of terrestrial invasive alien plants on water scarcity in south africa’, physics and chemistry of the earth, part b: hydrology, oceans and atmosphere 25, 237–242. http://dx.doi.org/10.1016/s1464-1909(00)00010-1 ersts, p.j., n.d., geographic distance matrix generator (v 1.2.3), computer software, viewed 16 april 2014, from http://biodiversityinformatics.amnh.org/open_source/gdmg (american museum of natural history, center for biodiversity and conservation). fargione, j.e. & tilman, d., 2005, ‘diversity decreases invasion via both sampling and complementarity effects’, ecology letters 8, 604–611. http://dx.doi.org/10.1111/j.1461-0248.2005.00753.x fermon, h., waltert, m., larsen, t.b., dall’asta, u. & mühlenberg, m., 2000, ‘effects of forest management on diversity and abundance of fruit-feeding nymphalid butterflies in south-eastern côte d’ivoire’, journal of insect conservation 4, 173–189. http://dx.doi.org/10.1023/a:1009638808635 florens, f.b.v., mauremootoo, j.r., fowler, s.v., winder, l. & baider, c., 2010, ‘recovery of indigenous butterfly community following control of invasive alien plants in a tropical island's wet forests’, biodiversity and conservation 19, 3835–3848. http://dx.doi.org/10.1007/s10531-010-9930-x forrester, w.s., 2011, ‘fruit-feeding butterfly assemblages at dlinza and entumeni nature reserves, kwazulu-natal: a quantitative biodiversity study’, msc dissertation, school of environmental science, north-west university. francesconi, w., nair, p.k.r., levey, d.j., daniels, j. & cullen, l., 2013, ‘butterfly distribution in fragmented landscapes containing agroforestry practices in southeastern brazil’, agroforestry systems 87, 1321–1338. http://dx.doi.org/10.1007/s10457-013-9640-y galatowitsch, s. & richardson, d.m., 2005, ‘riparian scrub recovery after clearing of invasive alien trees in headwater streams of the western cape, south africa’, biological conservation 122, 509–521. http://dx.doi.org/10.1016/j.biocon.2004.09.008 gerber, e., krebs, c., murrell, c., moretti, m., rocklin, r. & schaffner, u., 2008, ‘exotic invasive knotweeds (fallopia spp.) negatively affect native plant and invertebrate assemblages in european riparian habitats’, biological conservation 141, 646–654. http://dx.doi.org/10.1016/j.biocon.2007.12.009 ghazoul, j., 2004, ‘alien abduction: disruption of native plant-pollinator interactions by invasive species’, biotropica 36, 156–164. http://dx.doi.org/10.1111/j.1744-7429.2004.tb00308.x graves, s.d. & shapiro, a.m., 2003, ‘exotics as host plants of the california butterfly fauna’, biological conservation 110, 413–433. http://dx.doi.org/10.1016/s0006-3207(02)00233-1 hamer, k.c., hill, j.k., benedick, s., mustaffa, n., sherratt, t.n., maryati, m. et al., 2003, ‘ecology of butterflies in natural and selectively logged forests of northern borneo: the importance of habitat heterogeneity’, journal of applied ecology 40, 150–162. http://dx.doi.org/10.1046/j.1365-2664.2003.00783.x hanula, j.l. & horn, s., 2011, ‘removing an exotic shrub from riparian forests increases butterfly abundance and diversity’, forest ecology and management 262, 674–680. http://dx.doi.org/10.1016/j.foreco.2011.04.040 holt, r.d., 2009, ‘bringing the hutchinsonian niche into the 21st century: ecological and evolutionary perspectives’, proceedings of the national academy of sciences usa 106, 19659–19665. pmid: 19903876, http://dx.doi.org/10.1073/pnas.0905137106 hood, w.g. & naiman, r.j., 2000, ‘vulnerability of riparian zones to invasion by exotic vascular plants’, plant ecology 148, 105–114. http://dx.doi.org/10.1023/a:1009800327334 koh, l.p. & sodhi, n.s., 2004, ‘importance of reserves, fragments, and parks for butterfly conservation in a tropical urban landscape’, ecological applications 14, 1695–1708. http://dx.doi.org/10.1890/03-5269 krenn, h.w., zulka, k.p. & gatschnegg, t., 2001, ‘proboscis morphology and food preferences in nymphalid butterflies (lepidoptera: nymphalidae)’, journal of zoology 254, 17–26. http://dx.doi.org/10.1017/s0952836901000528 lawrence, j.m. & samways, m.j., 2002, ‘influence of hilltop vegetation type on an african butterfly assemblage and its conservation’, biodiversity and conservation 11, 1163–1171. http://dx.doi.org/10.1023/a:1016017114473 le maitre, d.c., gaertner, m., marchante, e., ens, e.-j., holmes, p.m., pauchard, a. et al., 2011, ‘impacts of invasive australian acacias: implications for management and restoration’, diversity and distributions 17, 1015–1029. http://dx.doi.org/10.1111/j.1472-4642.2011.00816.x le maitre, d.c., van wilgen, b.w., gelderblom, c.m., bailey, c., chapman, r.a. & nel, j.a., 2002, ‘invasive alien trees and water resources in south africa: case studies of the costs and benefits of management’, forest ecology and management 160, 143–159. http://dx.doi.org/10.1016/s0378-1127(01)00474-1 lu, s.-s. & samways, m.j., 2002, ‘conservation management recommendations for the karkloof blue butterfly, orachrysops ariadne (lepidoptera: lycaenidae)’, african entomology 10, 149–159. mckinney, m.l. & lockwood, j.l., 1999, ‘biotic homogenization: a few winners replacing many losers in the next mass extinction’, trends in ecology and evolution 14, 450–453. http://dx.doi.org/10.1016/s0169-5347(99)01679-1 molleman, f., krenn, h.w., van alphen, m.e., brakefield, p.m., devries, p.j. & zwaan, b.j., 2005, ‘food intake of fruit-feeding butterflies: evidence for adaptive variation in proboscis morphology’, biological journal of the linnean society 86, 333–343. http://dx.doi.org/10.1111/j.1095-8312.2005.00533.x molleman, f. & midgley, j.j., 2009, ‘δ15n analyses of butterfly wings and bodies suggest minimal nitrogen absorption in carrion and dung puddling butterflies (lepidoptera: nymphalidae)’, journal of research on the lepidoptera 41, 14–16. mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, south african national biodiversity institute, pretoria. nel, j.l., roux, j.l., maree, g., kleynhans, c.j., moolman, j., reyers, b. et al., 2007, ‘rivers in peril inside and outside protected areas: a systematic approach to conservation assessment of river ecosystems’, diversity and distributions 13, 341–352. http://dx.doi.org/10.1111/j.1472-4642.2007.00308.x new, t.r., 1991, butterfly conservation, oxford university press, melbourne. nyafwono, m., valtonen, a., nyeko, p. & roininen, h., 2014a, ‘butterfly community composition across a successional gradient in a human-disturbed afro-tropical rain forest’, biotropica 46, 210–218. http://dx.doi.org/10.1111/btp.12085 nyafwono, m., valtonen, a., nyeko, p. & roininen, h., 2014b, ‘fruit-feeding butterfly communities as indicators of forest restoration in an afro-tropical rainforest’, biological conservation 174, 75–83. http://dx.doi.org/10.1016/j.biocon.2014.03.022 ricketts, t.h., 2001, ‘the matrix matters: effective isolation in fragmented landscapes’, american naturalist 158, 87–99. rutowski, r.l., 1991, ‘the evolution of mate-locating behavior in butterflies’, american naturalist 138, 1121–1139. schlaepfer, m.a., sherman, p.w., blossey, b. & runge, m.c., 2005, ‘introduced species as evolutionary traps’, ecology letters 8, 241–246. http://dx.doi.org/10.1111/j.1461-0248.2005.00730.x shackleton, c.m., shackleton, s.e., buiten, e. & bird, n., 2007, ‘the importance of dry woodlands and forests in rural livelihoods and poverty alleviation in south africa’, forest policy and economics 9, 558–577. http://dx.doi.org/10.1016/j.forpol.2006.03.004 shreeve, t.g., 1984, ‘habitat selection, mate location, and microclimatic constraints on the activity of the speckled wood butterfly pararge aegeria’, oikos 42, 371–377. shreeve, t.g. & dennis, r.l.h., 2011, ‘landscape scale conservation: resources, behaviour, the matrix and opportunity’, journal of insect conservation 15, 179–188. http://dx.doi.org/10.1007/s10841-010-9336-9 sutherland, w.j., 1998, ‘the importance of behavioural studies in conservation biology’, animal behaviour 56, 801–809. http://dx.doi.org/10.1006/anbe.1998.0896 ter braak, c.j.f., 1995, ‘ordination’, in r.h.g. jongman, c.j.f. ter braak & o.f.r. van tongeren (eds.), data analysis in community and landscape ecology, pp. 91–173, cambridge university press, cambridge. thomas, j.a., 1984, ‘the behaviour and habitat requirements of maculinea nausithous (the dusky blue butterfly) and m. teleius (the scarce large blue) in france’, biological conservation 28, 325–347. http://dx.doi.org/10.1016/0006-3207(84)90040-5 turlure, c., baguette, m., stevens, v.m. & maes, d., 2011, ‘speciesand sex-specific adjustments of movement behavior to landscape heterogeneity in butterflies’, behavioral ecology 22, 967–975. http://dx.doi.org/10.1093/beheco/arr077 uehara-prado, m., brown, k.s. & freitas, a.v.l., 2007, ‘species richness, composition and abundance of fruit-feeding butterflies in the brazilian atlantic forest: comparison between a fragmented and a continuous landscape’, global ecology and biogeography 16, 43–54. http://dx.doi.org/10.1111/j.1466-8238.2006.00267.x valtonen, a., jantunen, j. & saarinen, k., 2006, ‘flora and lepidoptera fauna adversely affected by invasive lupinus polyphyllus along road verges’, biological conservation 133, 389–396. http://dx.doi.org/10.1016/j.biocon.2006.06.015 woodhall, s., 2005, field guide to butterflies of south africa, struik nature, cape town. article information authors: melodie a. mcgeoch1 hendrik sithole2 michael j. samways3 john p. simaika3 james s. pryke3 mike picker4 charmaine uys4 adrian j. armstrong5 ansie s. dippenaar-schoeman6,7 ian a. engelbrecht8 michelle hamer10 affiliations: 1cape research centre, south african national parks, cape town, south africa 2savanna and arid parks, south african national parks, kimberley, south africa 3department of conservation ecology and entomology, stellenbosch university, south africa 4zoology department, university of cape town, south africa 5ezemvelo kzn wildlife, pietermaritzburg, south africa 6arc-plant protection research institute, pretoria, south africa 7department of zoology and entomology, university of pretoria, south africa 8gauteng department of agriculture, conservation and environment, johannesburg, south africa 9centre for invasion biology, department of botany and zoology, stellenbosch university, south africa 10biosystematics division, south african national biodiversity institute, south africa correspondence to: melodie mcgeoch email: melodiem@sanparks.org postal address: po box 216, steenberg 7947, south africa dates: received: 05 may 2010 accepted: 27 jan. 2011 published: 13 may 2011 how to cite this article: mcgeoch, m.a., sithole, h., samways, m.j., simaika, j.p., pryke, j.s., picker, m., et al., 2011, ‘conservation and monitoring of invertebrates in terrestrial protected areas’, koedoe 53(2), art. #1000, 13 pages. doi:10.4102/koedoe.v53i2.1000 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) conservation and monitoring of invertebrates in terrestrial protected areas in this essay... open access • abstract • introduction • why monitor invertebrates in protected areas? • tackling the knowledge challenge    • advances    • species inventories for protected areas • use of invertebrates in conservation planning • threats to invertebrate diversity in protected areas    • invasive invertebrates • what to monitor? selection of invertebrate taxa and habitats    • odonata    • formicidae    • selection of habitats for monitoring • conclusion    • recommendations • acknowledgements • references abstract (back to top) invertebrates constitute a substantial proportion of terrestrial and freshwater biodiversity and are critical to ecosystem function. however, their inclusion in biodiversity monitoring and conservation planning and management has lagged behind better-known, more widely appreciated taxa. significant progress in invertebrate surveys, systematics and bioindication, both globally and locally, means that their use in biodiversity monitoring and conservation is becoming increasingly feasible. here we outline challenges and solutions to the integration of invertebrates into biodiversity management objectives and monitoring in protected areas in south africa. we show that such integration is relevant and possible, and assess the relative suitability of seven key taxa in this context. finally, we outline a series of recommendations for mainstreaming invertebrates in conservation planning, surveys and monitoring in and around protected areas. conservation implications: invertebrates constitute a substantial and functionally significant component of terrestrial biodiversity and are valuable indicators of environmental condition. although consideration of invertebrates has historically been neglected in conservation planning and management, substantial progress with surveys, systematics and bioindication means that it is now both feasible and advisable to incorporate them into protected area monitoring activities. introduction (back to top) monitoring biodiversity in protected areas (pas) forms an integral component of assessing their performance and providing the necessary information for effective management. invertebrates constitute a significant proportion of terrestrial and freshwater biodiversity (mound & gaston 1993), serve a series of critical ecosystem functions (losey & vaughn 2006), and as a consequence must necessarily be considered in pa monitoring systems (vane-wright 1993; woodroffe & ginsberg 1998). however, monitoring with invertebrates is associated with a series of regularly cited and well-recognised challenges (dobson 2005; lovell et al. 2010). these include their enormous richness and diversity of habits and habitats, inadequate systematic and biological knowledge for many groups, and the associated shortage of expertise and capacity (mound & gaston 1993). nonetheless, the field of insect bioindication and monitoring has developed substantially over the past two to three decades. a range of solutions, as well as several highly successful case studies of the use and incorporation of invertebrates in monitoring, now exist. as a consequence, the advantages of their inclusion have started to outweigh perceived and real disadvantages. monitoring has a long history in entomology with long-term data series being used, for example, to develop forecasting systems in pest management (conrad, fox & woiwod 2007). the rothamsted insect survey (uk) has, for example, been monitoring the relative abundance of insects since the 1960s (woiwod & harrington 1994). indeed, butterfly monitoring in the uk is amongst the most comprehensive current monitoring programmes using insects. here, four schemes provide complementary information on the distribution, abundance and diversity of butterfly assemblages. these schemes include mapping and multiscale atlases to monitor changes in species distributions, red data books on species conservation status, transects that provide information on population trends and a system of occasional surveys of selected species (thomas 2005). together these data are extremely valuable for butterfly conservation and they provide substantial insight into annual and long-term trends in the diversity, abundance and distribution of the species concerned (conrad et al. 2007). significant advances have also been made in south africa, and elsewhere, over the last three decades (mcgeoch 2002), both with regard to improvements in invertebrate systematics (speight & castella 2001; telford & littlewood 2008) and development of invertebrate bioindicators and their use in monitoring (mcgeoch 2007; samways, mcgeoch & new 2010). in this article, we use a selection of taxa and case studies (which is by no means comprehensive) to demonstrate the value and feasibility of both monitoring invertebrates in pas and including them in broader conservation assessments and biodiversity monitoring programmes. we discuss the objectives, opportunities and constraints specific to invertebrate monitoring in pas, with a particular focus on south africa. we identify suitable taxa for monitoring based on a range of monitoring objectives and experiences, and make recommendations for the implementation of an invertebrate monitoring programme for pas in south africa. why monitor invertebrates in protected areas? (back to top) one of the key reasons for conserving and monitoring invertebrates in their own right (i.e. as the subject of monitoring), particularly in pas, is to ensure the adequate protection of rare and threatened invertebrate species and communities (samways 1993a). nonetheless, invertebrates are also useful, appropriate and often highly effective and informative indicators of other elements of biodiversity, ecosystem function and restoration, system health and associated threats, including invasive alien species (mcgeoch 2007). as discussed later, invertebrates provide sensitive, appropriate and logistically feasible target taxa for monitoring a wide cross section of pa management objectives (table 1). in november 1995, south africa ratified the convention on biological diversity (cbd) under which signatories are obligated to develop a strategic plan for the conservation and sustainable use of biodiversity. meeting the goals of the convention requires intensive national effort including (1) discovery, description and inventory of species, (2) analysis and synthesis of information into predictive classification systems, and (3) organisation of this information into an efficiently retrievable form that best meets the needs of science, conservation and society. invertebrates necessarily form a significant component of this task as they not only perform critical roles in ecosystems (via pollination, decomposition, soil property maintenance, predation, parasitism, and herbivory; see for example, lawrence (1953), scholtz and holm (1985) and scholes and walker (1993)) but also contribute disproportionately to biodiversity and outnumber plant and vertebrate richness several fold (grimaldi & engel 2005; samways 2005). although lesser known than in the case of vertebrates and plants, several south african invertebrate taxa also display high levels of endemism. many invertebrates have narrow ranges; for example, in kwazulu-natal alone seven species of butterfly, about 80 species of snail, and more than 200 species of millipede are found exclusively in the province (pringle, henning & ball 1994; hamer 1998; herbert & kilburn 2004). these species require conservation action to ensure their survival because of their small distribution ranges and increasing rates of land transformation (thompson 2008). another example is the very high concentration of taxa with a southern gondwanan ancestry that are restricted to the cape fold mountains. this very rich south african gondwanan fauna (800 species, 200 families) has its primary hotspot on the cape peninsula (a substantial proportion of which constitutes table mountain national park [tmnp] in the western cape [day 2005]). high levels of endemism (111 cape peninsula endemics, according to picker and samways [1996]) are concentrated in refugial habitats such as mountain streams, southern afrotemperate forest and caves. the cape floristic region, in general, has a distinctive invertebrate fauna, with high levels of endemism in gall-forming insects (wright & samways 1998), leafhoppers (stiller 2002), dung beetles (davis 2002), scorpions (prendini 2005), millipedes (hamer & slotow 2002), velvet worms, (onychophora) (daniels et al. 2009; hamer, samways & ruhberg 1997), heelwalkers (mantophasmatodea) (damgaard et al. 2003) and lycaenid butterflies (cottrell 1985). other biomes are probably less well studied in terms of their invertebrate fauna and levels of endemicity, but the more northern afrotemperate forests, and even grassland and savanna have high levels of narrow endemism for some taxa that have been investigated in this respect (e.g. hamer, slotow & lovell 2006; hamer & slotow 2009). pas play a vital role in the conservation of such narrow range endemics, and these species are generally considered ‘species of special concern’ in pa management objectives (table 1). for example, in the tmnp several invertebrates require special attention, such as colophon westwoodi (coleoptera: lucanidae) and trimenia malagrida malagrida (lepidoptera: lycaenidae). peripatopsis alba is an endangered cave-dwelling velvet worm confined to table mountain (hamer et al. 1997). these taxa have been found only at very few localities and these sites thus require special conservation management. table 1: examples of biodiversity objectives in protected areas (see mcgeoch, m.a., dopolo, m., novellie, p., hendriks, h., freitag, s., ferreira, s., et al., 2011, ‘a strategic framework for biodiversity monitoring in sanparks’, koedoe 53(2), art. #991, 10 pages. doi:10.4102/koedoe.v53i2.991) and relevant associated invertebrate monitoring. tackling the knowledge challenge (back to top) the conservation and monitoring of invertebrates, including those in pas in south africa, are well-known to present unique taxonomic challenges given their overwhelming diversity (vane-wright 1993; grimaldi & engel 2005; scholtz & chown 1995). with the comparative exception of butterflies, dragonflies, trapdoor and baboon spiders (dippenaar-schoeman 2002; henning & henning 1989; eds. henning, terblanche & ball 2009; samways 2006), the conservation status of most invertebrate groups in south africa is poorly known (other comparatively well-known groups include terrestrial molluscs, one genus of millipede, and onychophora) (scholtz & chown 1995). for example, a systematic conservation planning project, initiated by the gauteng department of agriculture, conservation and environment (dacel) in 2000, aimed to include broad representation of threatened species from a range of taxonomic groups (dacel 2003). however, inclusion of the invertebrates proved a particular challenge owing to the lack of available information for identifying species of conservation concern. as a result, an expert consultation approach was followed to identify candidate species for inclusion. through this process, insect and arachnid species were identified and included in the first version of the conservation plan (dacel 2003). surveys were then conducted for several of these species to obtain additional distribution data. however, as a result of these surveys several species were removed from the expert list when they were found to be far more widespread than originally thought. this example well illustrates the challenges involved in identifying invertebrate species of conservation concern and deciding which need to be monitored. expert opinion in itself may be insufficient and available distribution data are often inadequate. therefore, whilst surveying and quantifying invertebrate biodiversity are valuable in their own right (lovell et al. 2010), a focus on better-known groups is likely to be more effective for the purposes of conservation planning and monitoring in pas. at present, management decisions are often based on information on the distribution of vertebrates or plants, with the assumption that the latter will also benefit invertebrates. however, whilst there often is a broad relationship between plant and insect diversity at large spatial scales (e.g. procheş et al. 2009), this approach is insufficient to guarantee the conservation of rare invertebrate species at the finer spatial scales at which conservation action usually occurs (e.g. lovell et al. 2007; panzer & schwartz 1998). advances despite the rather dismal reflection of the status of invertebrate knowledge as described above, knowledge of selected invertebrate taxa in south africa is increasing exponentially. monitoring efforts in some instances are also supported by adequate keys, distribution information and baseline conservation status assessments. for example, the south african butterfly conservation atlas (sabca) project, launched in 2007 to determine the distribution and conservation status of butterflies in south africa, lesotho and swaziland (http://sabca.adu.org.za/), will be published in 2011. it will incorporate field data (2007–2010) and museum and private collection records (eds. henning et al. 2009). the lepidopterists’ society of southern africa has been instrumental in these activities and their achievements provide an excellent example of the role of citizen scientists along with semiprofessional and professional entomologists in promoting invertebrate conservation and improving the knowledge base with regard to particular taxa. as a charismatic group of taxa, butterflies have long received the attention of entomologists, collectors and conservationists across the world. similar approaches and activities for other invertebrate groups are nonetheless possible. for example, south africa has a rich arachnid fauna with about 5000 known species, representing 6% of global arachnid diversity. although they constitute a diverse and ecologically important group of invertebrates in south africa, they have traditionally been comparatively poorly sampled in some areas. however, in 1997 the south african national survey of arachnida (sansa) was initiated by the biosystematics unit (arc-plant protection research institute). sansa aims to inventory the arachnofauna of the country and forms part of the threatened species programme of the south african national biodiversity institute (sanbi) to determine the distribution and conservation status of spiders and scorpions. one of the focus areas of sansa is to survey pas to obtain species-specific information, compile inventories and to determine which species in south africa receive some protection. more than 86 surveys in pas are currently underway, of which 25 have already resulted in published annotated checklists that provide information on abundance, behaviour and the distribution of arachnid species. several forms of participation have involved pas, including sansa surveys, surveys by pa managers and rangers, student research projects (seven msc projects completed), by-catch data from other research projects and records submitted by the public. this has dramatically increased the knowledge of the spider fauna of pas and with material made available to taxonomists has resulted in the description of several new species (e.g. wesolowska & haddad 2009). during this survey a record 430 species were recorded, which represent more than 20% of the south african spiders and include several new and undescribed species (haddad, dippenaar-schoeman & wesolowska 2006; wesolowska & haddad 2009). long-term surveys in pas also increase the number of species sampled. for example, during the survey of the kruger national park (knp) 152 species were reported in 2003, increasing the number currently listed for the park to 305 (dippenaar-schoeman & leroy 2003, see also dippenaar-schoeman 1988, 2006). pas have proven to be particularly valuable sites to sansa, both from the perspective of encountering pristine habitat and high diversity, as well as for the safety of survey teams (dippenaar-schoeman et al. 1999, 2005). species inventories for protected areas species inventories are critical for effective pa management and are generally considered to be important by pa managers, specialist taxon scientists, citizen scientists and the layperson alike. such inventories form the basis of identifying the presence of species of special conservation concern in pas (table 1), are valuable in regional conservation planning processes, and are the first step in any monitoring programme across a range of monitoring objectives (rohr, kim & mahan 2007). species inventories, when repeated over time, may also be used as a tool to monitor the loss or gain of species in pas (droege, cyr & larivée 1998). because of the importance of inventories in species conservation assessment and pa management, engelbrecht (2010) investigated to what extent and how invertebrate species inventories are currently applied in pa management in south africa. pa managers and ecologists were asked, using a questionnaire-based survey, to comment on whether (1) a documented management plan existed for the pa in question, (2) any invertebrate species inventories existed for the pa, and if so, (3) whether those inventories had been used to guide the development of any of the management goals or actions for the pa. although the overwhelming majority considered inventories desirable, very few of the responses received indicated that invertebrates were specifically considered in management goals or actions. those that did included the brenton blue butterfly reserve (western cape) and the suikerbosrand nature reserve (gauteng), which is home to the heidelberg copper (engelbrecht 2010). in other cases, the principal value of inventories was to tourists and scientists visiting the pa. the outcome of this study illustrates that although insect conservation and monitoring have not yet become mainstream in pa planning and management processes, the potential value of inventories is (1) widely acknowledged and (2) would be more suited to this purpose where accompanied by information on richness patterns, distributions, and rare and indicator species (engelbrecht 2010). use of invertebrates in conservation planning (back to top) invertebrates have been successfully used in systematic conservation planning in south africa. for example, in kwazulu-natal, a range of spatial products that include invertebrates (e.g. databases of distribution records, geographical information system layers and predicted distribution maps) have been developed to support the assessment of land use change applications by the integrated environmental management (iem) team of ezemvelo kzn wildlife (see also an example by edge (2005) on butterflies in the southern cape). these products include the ezemvelo kzn wildlife biodiversity database, which, as of july 2009, contains distribution records for 3649 invertebrate species and subspecies, both within and outside of pas. these records are used in species distribution modelling and to indicate which species occur, or are likely to occur, on land subject to applications for land use change. to date, only red listed species and those endemic to the province have been included in this modelling. the first spatial product incorporating modelled invertebrate distributions included 98 invertebrate species and subspecies (armstrong 2002; goodman 2000). as shown in table 2, despite this and other available and related tools in kwazulu-natal, invertebrates have been explicitly included in only a small proportion of land use change applications under the national environmental management act (nema) (act 107 of 1998), where the proposed land use change may have significant impacts on species of special concern. various mitigation measures were proposed by ezemvelo kzn wildlife as befitting the range of invertebrate taxa concerned. the recommendations ranged from no mitigation specific to invertebrates, through restriction of the proposed land use to certain portions of the land (e.g. to already transformed portions of the land), to set-asides for invertebrate conservation. in one case, refusal of the permit was recommended on invertebrate conservation grounds. some of these recommendations were incorporated into the record of decision issued by the provincial department of agriculture and environmental affairs (now the department of agriculture, environmental affairs and rural development), whilst others were not. one of the insights gained from the aforementioned process is the need to strengthen the profile of invertebrates in conservation planning, both within and outside of pas, so that charismatic umbrella vertebrate taxa are not relied upon to effect the conservation of invertebrates and their habitats. the ezemvelo kzn wildlife iem team now communicates to all relevant parties when umbrella species are used for invertebrate conservation purposes. the need to conserve invertebrates is not readily accepted or appreciated by some permitting authorities, applicants and environmental consultants, and some lack expertise on invertebrates and therefore ignore them. profiling of invertebrates and training of consultants are required to successfully integrate invertebrate conservation into processes legislated by nema and its subsidiary acts. this will allow invertebrates to be considered explicitly and conserved in areas subject to change in land use. table 2: the number of land use change applications in kwazulu-natal with species concerns, and the number of these with invertebrate concerns as ascertained by ezemvelo kzn wildlife, the mitigations proposed, and the final decisions by the competent authority. threats to invertebrate diversity in protected areas (back to top) invertebrates are susceptible to the same key threats as other biodiversity elements, namely habitat loss and degradation, invasive alien species and climate change (mcgeoch 2002; new 1995; scholtz & chown 1993). a recent assessment of invertebrate diversity in the tmnp provides a good example of threats to invertebrates (pryke & samways 2008, 2010). as is the case elsewhere in the fynbos biome, most of the upper elevational areas of the tmnp are well protected, whereas the low-lying areas are of significant concern. for example, the importance of conserving lowlands is well illustrated by noordhoek wetlands (a new addition to tmnp), with two new water beetle species recently identified from the area (closest known localities were zaire and mozambique), and a third species new to science (pryke & samways 2009a). lower-lying areas that are covered by plantations tend to have lower invertebrate species richness, and for some species lower abundance, compared to natural forests and fynbos (pryke & samways 2009b). invertebrate assemblages of indigenous natural forests are likely to be more difficult to restore than those of fynbos and thus the protection of the remaining natural forests in the tmnp is a conservation priority (pryke & samways 2009b). monitoring invertebrate responses to fire (e.g. frequency and intensity) is also important in pas within biomes where fire is a key habitat management tool, such as savanna and fynbos (parr et al. 2002). fire is a particular conservation risk in mediterranean-type biomes and, although it is an essential element in the maintenance of biodiversity, fire regimens (frequency, intensity and timing) have been significantly altered by increasing human population densities, resulting in too frequent, too infrequent and other ecologically undesirable fire properties (forsyth & van wilgen 2008). a study by pryke (2008) showed that the invertebrates in the tmnp demonstrated differential short-term resilience to fire, especially with regard to species richness and abundance. some components of the invertebrate assemblage were remarkably resilient to fire (particularly ants), whilst others were far more conservative (pollinators and detritivores). invertebrates were also found to recolonise high-altitude grasslands in the drakensberg within a short space of time after burning (uys, hamer & slotow 2006). however, although not well studied, fire in afrotemperate forest may have a negative impact on some less mobile taxa such as molluscs (uys, hamer & slotow 2010). in terms of fire-related monitoring objectives in pas, invertebrates are thus an important element of monitoring the status of and trends in habitat persistence, degradation and rehabilitation (table 1). invasive invertebrates monitoring programmes for invertebrate conservation in pas should also consider invasive invertebrates (table 1). established invaders may thrive and come to dominate local communities, often causing a loss of biodiversity and impacting on ecosystem functioning (vilà et al. 2010). whilst a few terrestrial alien invasive invertebrates are well known in south africa (e.g. the argentine ant (linepithema humile) and the european wasp (vespula germanica)), the majority remain inconspicuous in local faunas owing to a lack of thorough taxonomic surveys. in the absence of precise taxonomic identification by experts, the number of alien species is often greatly underestimated. alien invertebrates in south african pas span most of the higher-level taxa and include earthworms, slugs, snails, millipedes, woodlice, spiders, mites and insects such as earwigs, wasps and ants (macdonald et al. 2003; raharinjanahary 2007). whilst some of these alien species have not become invasive, many invaders have the potential to impact native ecosystems, either directly through predation or indirectly by disrupting mutualisms or through interference competition (kenis et al. 2009). for example, many alien earthworms are highly invasive and often dominate faunas even in relatively untransformed habitats (e.g. horn, plisko & hamer 2007). most biological invasions arise through human-mediated extra-range dispersal pathways (wilson et al. 2009) and can also include range extensions of native species, such as the cape honeybee (apis mellifera capensis) (braack & kryger 2003; valéry et al. 2008). monitoring or research programmes that include accurate taxonomic identifications are the most likely way of detecting new alien species and structured monitoring programmes for target species allow for early detection and potentially rapid control. those involved in invertebrate monitoring should not only be familiar with known invasives, but be on the lookout (early detection) for species that are problematic in other countries, but have not yet been recorded in south africa. examples include the yellow crazy ant (anoplolepis gracilipes) and red imported fire ant (solenopsis invicta) (lowe et al. 2000). what to monitor? selection of invertebrate taxa and habitats (back to top) the debate on which taxa are best suited for inclusion in monitoring programmes is long running and closely related to the debate on which taxa are good bioindicators of environmental change, or of broader biodiversity per se (kremen et al. 1993; mcgeoch 1998). more than 32 criteria have been identified for guiding the choice of taxa suitable for bioindication and monitoring (mcgeoch 1998). the relative importance of these particular criteria is determined principally by the monitoring objective. biodiversity monitoring in pas encompasses several broad objectives and invertebrates are potentially both relevant and useful for most of these (table 1). however, all taxa have a range of advantages and disadvantages as monitoring tools and these should be evaluated to assess their suitability in a particular monitoring context (samways 1993b). decisions on what to monitor must also necessarily consider the focal taxa and habitats along with logistic feasibility of the proposed exercise. different invertebrate taxa are obviously more, or less, appropriate for monitoring in particular environments and are differentially sensitive to particular stressors. table 3 provides an assessment of the value of selected higher-level taxa to monitoring based on a range of key generic criteria. the outcome of this assessment provides guidance for selecting which higher taxa to focus on a priori, before proceeding with more specific indicator testing as would be required to achieve more specific monitoring objectives (see mcgeoch [1998] and mcgeoch, van rensburg & botes [2002] for general approach). we selected 16 objectives and criteria that are most relevant in a pa context and assessed a range of higher taxa against these. rather than assessing all higher taxa, a subgroup of seven, which were considered to be those broadly most feasible in a south african context (based on the collected specialist opinion and experience of the authors), was selected a priori. other taxa may well be relevant and suitable for more narrowly defined purposes. the taxa were scored for each criterion (see explanation in table 3) and ranked according to a descending score (left to right, table 3). odonata, formicidae, lepidoptera (butterflies) and scarabaeidae were considered best suited for monitoring in pas. interestingly, coleoptera, hymenoptera and lepidoptera (along with molluscs and arachnids) currently receive most research attention in south african national parks (figure 1). for example, dung beetles, whilst potentially useful indicators as a result of their specialised distribution patterns, require fine-scale spatial data to realise this potential (davis 2002). given the rapid increase in knowledge and expertise on south africa’s arachnid fauna, particularly in pas, the score for this taxon is likely to increase in the foreseeable future. we consequently use the odonata and formicidae (hymenoptera) as two of the highest-scoring taxa in our assessment to illustrate aspects of the suitability and application of invertebrates for monitoring in pas. table 3: an assessment of the suitability of taxonomic groups to be used for monitoring in protected areas, according to a range of objectives (see table 1) and general suitability criteria. figure 1: percentage of research projects (n = 63) on invertebrates (a) per taxon and (b) per park (including nontaxon specific projects) currently registered with south african national parks (as at november 2009). odonata several freshwater monitoring schemes involving macroinvertebrates are well developed with a long history, for example the river invertebrate prediction classification system used in the uk to monitor the pollution status of water courses (wright et al. 2000) and the south african scoring system (sass) (revenga et al. 2005). good taxonomic and biological information, along with knowledge of species conservation status and responses to habitat quality are amongst the key suitability criteria in invertebrate monitoring (table 3). the odonata in south africa perhaps best exemplify this with existing red data information and comprehensive field guides that make working with the group broadly accessible to biologists, the public and citizen scientists (samways 2006, 2008). south africa’s freshwater systems are under intense pressure (nel et al. 2007) and the odonata are sensitive indicators of the quality of these systems. this includes their response to pollution and invasive alien species impacts (chovanec 2000), as well as their recovery after alien plant removal in rivers in pas in south africa (samways & sharratt 2010). in addition to the well-known and widely used sass system (dallas & day 1993; revenga et al. 2005), the dragonfly biotic index (dbi) provides a measure of ecological integrity for freshwater systems. the dbi is a weighted index (see table 4) based on the quantitative assessment of three subindices of species distribution, threat status and sensitivity to disturbance (simaika & samways 2008, 2009a). the total dbi of a water body (stream, river or pool) reflects the total odonate assemblage, thus allowing for water bodies to be compared and restoration success to be monitored (simaika & samways 2008). every south african odonate species has been assigned a score (samways 2008). the dbi has been tested and applied in biomonitoring (simaika & samways 2009a), 2011 to measure habitat recovery (simaika & samways 2008) and select sites for conservation (simaika & samways 2009b). previous work has shown a strong correlation between adult dragonfly scores and macroinvertebrate scores (simaika & samways 2011; smith, samways & taylor 2007). an advantage of the dbi over conventional macroinvertebrate indices is that it operates at the species level and is therefore highly sensitive to habitat condition. it, therefore, has good potential for environmental assessment and monitoring freshwater biodiversity and quality, alongside sass (simaika & samways 2011). the low field effort required to obtain a dbi score for a site also makes this a low-cost and readily applied method. odonata are therefore particularly useful for monitoring freshwater quality and landscape physiognomy around riverine and other aquatic habitats in south africa’s pas (samways 1993b). table 4: the subindices of the dragonfly biotic index (dbi) range from 0 to 3. the dbi is based on three subindices relating to geographical distribution, level of threat, and sensitivity to habitat change, with particular reference to invasive alien riparian trees. the dbi is the sum of the scores for the three subindices, and ranges from 0 to 9. a common, widespread, not-threatened and highly tolerant (of disturbance) species would score 0 (0 + 0 + 0), whilst a highly range-restricted, threatened and sensitive species would score 9 (3 + 3 + 3). formicidae other important criteria to consider when selecting a taxon for monitoring include the taxonomic richness of the group, their ubiquitousness, the diversity of habitats that they occupy, their importance in ecosystem functioning and the range of functional roles that they perform in the environment (table 3). ant assemblages have been widely tested in the assessment of environmental disturbance and ecosystem condition and their abundances and distributions are strongly correlated with temperature and vegetation communities (andersen & majer 2004; botes et al. 2006b; kaspari & majer 2000; underwood & fisher 2006). however, equally important is the cost, effort and capacity associated with sampling, identifying and analysing the taxon and, for example, the feasibility of using parataxonomists for such species-rich groups (new 1998; underwood & fisher 2006). ant monitoring in south africa has gained significant momentum over the last decade. for example, the iimbovane project (http://academic.sun.ac.za/iimbovane) is a spatially extensive and temporarily replicated ant monitoring programme (currently focussed in the western cape), where equivalent information on invertebrates is rare (driver et al. 2005; koch et al. 2000). it thus delivers baseline data of species distributions in different vegetation types in two biodiversity hotspots (the fynbos biome and the succulent karoo biome), against which the impact of future environmental change can be assessed (braschler et al. 2010). the project uses sites that are disturbed in a variety of ways and control sites that are typically located within pas. spatially and temporally replicated surveys are used to detect trends and assess natural variation, including variation due to seasonality, climatic events or fire events. however, such surveys face many challenges, including securing expertise and funding for long periods. continuity was built into the iimbovane project by combining monitoring with a long-term outreach programme (teaching biodiversity to high school learners participating in the monitoring; see braschler [2009]) and by involving technical staff on long-term contracts. including both pristine and disturbed sites means that the data may be used for a variety of purposes. it is inevitable that much conservation will have to be done outside pas, including transformed landscapes, which makes monitoring disturbed sites necessary. control sites in pas are particularly valuable and serve to separate various fine-scale and large-scale disturbance effects. furthermore, where restoration is the goal for conservation, baseline data are needed on what is natural to an area and this information is often best obtained from control sites with natural vegetation such as those found inside pas. faced with adopting and implementing an invertebrate monitoring programme, the knp has also assessed the feasibility, resource requirements and cost of ant monitoring in the park. ants were selected as one of three possible groups (the others were dung beetles and butterflies) because of the presence of taxonomic expertise in the park, a history of active collaboration with external researchers on this taxon, good baseline data on ant assemblages in the knp being available and the ubiquitousness and relevance of the taxon in the area (parr & chown 2001; parr et al. 2004). to assess the costs associated with ant monitoring, ants were sampled from 34 plots (2000 m2 each) using 20 pitfall traps per plot. traps were left open in the field for 5 days. pitfall trapping was chosen because it is widely used for ant sampling and is considered efficient in savanna ecosystems (parr & chown 2001; underwood & fisher 2006). sampling was done from 2003 to 2005 and again in 2009. the optimum workforce needed for the field work of a single 2000 m2 area was found to be four people, which is similarly labour intensive to other monitoring projects in sanparks (figure 2). the total time spent sampling and identifying ants on a single plot was 460 min, including 80 min spent on setting and removing traps, 80 min for screening an average of 300 individual ants, and 300 min for identifying them to species level under the microscope. although comparatively inexpensive (see below), ant sampling was found to be more time consuming than other monitoring projects in the knp (figure 2). this may be mitigated by constantly using the same trained assistants and applying simplified assessments such as those developed for australia (andersen et al. 2002). the resources required for postcollection sorting, identification, curation and analyses are often not carefully considered and these are all critical for the collection and long-term use and storage of monitoring information. considered per appropriate sample area, ant monitoring was also found to be one of the least expensive monitoring projects in the knp (figure 2). ants have been demonstrated to provide particularly valuable management-based information across a range of regions of the world, in particular for invasive species monitoring, detecting trends in functionally important species and those of specific conservation concern and assessing long-term changes in ecosystems and the consequences of management action (underwood & fisher 2006). this series of factors makes the formicidae particularly suitable for inclusion in pa monitoring systems in south africa. figure 2:estimated total resource costs per unit sampling area of the ant monitoring project in the kruger national park compared with six other monitoring projects in sanparks (different area for each project, but the minimum relevant unit area is used in each case). selection of habitats for monitoring an alternative to selecting particular taxa is the selection of particular habitats as the focus for invertebrate monitoring (dennis, shreeve & sheppard 2007; see samways [1993a] for a conceptual framework). the rationale for this approach may be either to conduct monitoring across a range of representative habitat types or to focus on particularly threatened or sensitive (priority) habitat types known to support unique assemblages of invertebrates and/or other taxa (new 1995). systems and habitats worth monitoring will in many cases vary across pas, as a result of biome-specific ecosystem characteristics. component species and communities in such habitats may then be given weightings based on rarity, threat, endemism and phylogenetic uniqueness (balmford, jayasuriya & green 1996; bonn, rodrigues & gaston 2002; davis 2002; kier et al. 2009). the advantage of a habitat-based approach is that a clear association of habitat and critical elements of biodiversity can serve to prioritise a system worth investing in for long-term monitoring. the habitat can then be monitored for threats such as invasive species, climate change and water quality change. initial monitoring will form a baseline against which future changes in species composition can be compared and indicator taxa may be used to monitor responses to environmental change (botes et al. 2006a,b; davis 2002; mcgeoch et al. 2002; rösch, chown & mcgeoch 2001; uys et al. 2010). an example of habitat-focused monitoring is the streams in the tmnp. because of the evolutionary significance and high levels of endemism, both streams and their associated gondwanan biota could be prioritised for monitoring. further reasons for doing so include the fact that gondwanan taxa tend to be stenothermal (cold-adapted) (mckie, cranston & pearson 2004) and thus are most at risk from elevated temperatures and the synergistic influence of reduced precipitation predicted by models of global warming for the western cape (de wit & stankiewicz 2006). baseline data for future monitoring in this habitat would include a thermal and biotic profile of a selected stream, encompassing both spatial (longitudinal stream profile) and seasonal variability. against this baseline, future monitoring would then be conducted to detect the influences of climate change on both water temperature and invertebrate biota. because water temperature is also influenced by flow, monitoring would simultaneously provide a baseline against which effects of the possible abstraction of the table mountain aquifer on the stream and its biota can be assessed. a year’s worth of baseline data have already been collected for this project at window gorge stream (ketley 2009) and the project has been transferred to sanparks for future monitoring. regular, ongoing monitoring is essential in this system if trends in change drivers (especially changing flow rates, water temperatures and biotic responses) are to be captured. both temporal and small-scale turnover, as well as habitat association, are generally poorly understood in invertebrate communities (lovell et al. 2010) and would need to be carefully considered in establishing and maintaining a habitat-based or community-based monitoring programme. for example, turnover of epigaeic invertebrates in afrotemperate forest in the drakensberg ranged from complete to 50%, even in forests within the same valley (hamer & slotow 2009; uys, hamer & slotow 2009). similarly, temporal turnover between seasons and years may be high for epigaeic invertebrates and flying insects, particularly (lovell et al. 2010). conclusion (back to top) we have shown that there are already several examples in south africa of invertebrates (1) being successfully inventoried, (2) having their conservation status assessed, (3) being used in conservation planning, and (4) being used as bioindicators in monitoring programmes. furthermore, a significant component of such activity has taken place in pas. generally such programmes focus on taxa that are well studied and readily identifiable and which are responsive to the variable under consideration or appropriate to the pa management objective. nonetheless, for invertebrate information to be relevant to pa management, it is best linked explicitly to one or more management objectives for the pa in question (walker & salt 2006). importantly, sufficient information and support are now available for particular taxa to be included practically in monitoring in pas, and in many instances significant levels of endemism render invertebrates important for monitoring in their own right. indeed, several current initiatives are set to improve this situation substantially. good data on species richness, species distributions and population trends remain essential if invertebrates are to be considered in conservation management decisions and if effectiveness of these decisions for their protection is to be assessed. currently such data are available for only a few key taxa. the south african biosystematics initiative of the national research foundation currently provides some funding for invertebrate survey and monitoring research, if projects are well justified and placed in an appropriate conceptual framework. significant opportunities thus exist to develop collaborations between pa agencies needing data and researchers skilled to provide these data. collaboration between pa and external researchers on pa invertebrate research can produce valuable baseline data, taxonomic knowledge and inventories, which then enable pas to (1) integrate invertebrates into pa monitoring programmes in a feasible manner and (2) continue key invertebrate monitoring in the longer term. recommendations the following points serve as general recommendations: • although invertebrates are relevant and important across a range of pa management objectives, they should be explicitly and clearly linked to these objectives. • monitoring activities that involve invertebrates should focus on taxonomically well-known groups that are supported by, for example, good keys, available expertise and published evidence of their value in the specific context. the success of initiatives such as sansa, iimbovane and sabca illustrates the feasibility and value of invertebrate surveys and monitoring in pas. • an important point to communicate to conservation planners is that vegetation or other surrogates do not adequately capture invertebrate diversity and that setting targets without considering at least some invertebrates is unlikely to capture invertebrate diversity (e.g. hamer & slotow 2009). • in a south african pa context, dragonflies, ants, butterflies and dung beetles (and increasingly also spiders) are appropriate for monitoring a broad range of management objectives. • invertebrate species of special concern represent a particular knowledge gap for pas and here research is required to identify localised endemics and threatened species along with surveys targeting potentially important areas and taxa. for threatened species, some idea of habitat requirements and basic biology is required for adequate management, as well as approaches to monitoring threatened invertebrate species. • similarly, surveys of invasive alien invertebrate species are required along with research on their impacts and approaches to prevent further introductions. • profiling of invertebrates and training specialist invertebrate consultants are required to successfully integrate invertebrate conservation into processes legislated by nema and its subsidiary acts. this will allow invertebrates to be explicitly considered and conserved in areas subject to change in land use. • collaborative relationships between entomologists, particularly systematists, and pa agencies should be fostered to maximise both research opportunities and opportunities to improve our knowledge of the invertebrate fauna of pas. citizen scientists continue to make an extremely valuable contribution to invertebrate surveys and monitoring in south africa and their potential role in pa monitoring is significant. acknowledgements (back to top) this manuscript arose from a workshop on the use of invertebrates in monitoring in pas, followed by a symposium on this theme at the 16th congress of the entomological society of southern africa (stellenbosch, july 2009). the authors thank the pioneers of mainstreaming invertebrate conservation in kwazulu-natal, the iem team at ezemvelo kzn wildlife, sharon louw, john craigie, peter goodman and colleagues, and rachel ndlovu for assistance with obtaining information from iem case studies. cost estimates for fish, lion, elephant and woody vegetation monitoring projects in sanparks were kindly provided by sanparks staff sam ferreira, hugo bezuidenhout and ian russell. we also thank two anonymous reviewers for their helpful comments. the national research foundation, andrew w. mellon foundation and dst–nrf centre for invasion biology are acknowledged for financial support. references (back to top) andersen, a.n., hoffmann, b.d., müller, w.j. & griffiths, a.d., 2002, ‘using ants as bioindicators in land management: simplifying assessment of ant community responses’, journal of applied ecology 39, 8−17. doi:10.1046/j.1365-2664.2002.00704.x andersen, a.n. & majer, j.d., 2004, ‘ants show the way down under: invertebrates as bioindicators in land management’, frontiers in ecology & environment 2, 291−298. doi:10.1890/1540-9295(2004)002[0292:astwdu]2.0.co;2 armstrong, a.j., 2002, ‘insects and the determination of priority areas for biodiversity conservation in kwazulu-natal province, south africa’, african entomology 10, 11−27. armstrong, a.j. & louw, s., 2010, ‘kzn biodiversity status assessment report: millar’s tiger moth, callioratis millari’, ezemvelo kzn wildlife report, pietermaritzburg. balmford, a., jayasuriya, a.h.m. & green, m.j.b., 1996, ‘using higher-taxon richness as a surrogate for species richness: ii local adaptations’, proceedings of the royal society of london b 263, 1571−1575. doi:10.1098/rspb.1996.0230, doi:10.1098/rspb.1996.0186 bonn, a., rodrigues, a.s.l. & gaston, k.j., 2002, ‘threatened and endemic species: are they good indicators of patterns of biodiversity on a national scale?’, ecology letters 5, 733−741. doi:10.1046/j.1461-0248.2002.00376.x botes, a., mcgeoch, m.a. & van rensburg b.j., 2006a, ‘elephantand human-induced changes to dung beetle (coleoptera: scarabaeidae) assemblages in the maputaland centre of endemism’, biological conservation 130, 573−583. doi:10.1016/j.biocon.2006.01.020 botes, a., mcgeoch, m.a., robertson, h.g., van niekerk, a., davids, h.p. & chown, s.l., 2006b, ‘ants, altitude and change in the northern cape floristic region’, journal of biogeography 33, 71−90. doi:10.1111/j.1365-2699.2005.01336.x braack, l. & kryger, p., 2003, ‘insects and savannah heterogeneity’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience, pp. 263–275, island press, london. braschler, b., 2009, ‘successfully implementing a citizen-scientist approach to insect monitoring in a resource-poor country’, bioscience 59, 103−104. doi:10.1525/bio.2009.59.2.2 braschler, b., mahood, k., karenyi, n., gaston, k.j. & chown, s.l., 2010, ‘realizing a synergy between research and education: how participation in ant monitoring helps raise biodiversity awareness in a resource-poor country’, journal of insect conservation 14, 19−30. doi:10.1007/s10841-009-9221-6 chovanec, a., 2000, ‘dragonflies (insecta: odonata) as indicators of the ecological integrity of aquatic systems – a new assessment approach’, verhandlungen der internationalen vereiningung für theoretische und angewandte limnologie 27, 887−890. chown, s.l., scholtz, c.h., klok, c.j., joubert, f.j. & coles, k.s., 1995, ‘ecophysiology, range contraction and survival of a geographically restricted african dung beetle (coleoptera, scarabaeidae)’, functional ecology 9, 30−39. doi:10.2307/2390087 conrad, k.f., fox, r. & woiwod, i.p., 2007, ‘monitoring biodiversity: long-term changes in insect abundance’, in a.j. stewart, t.r. new & o.t. lewis (eds.), insect conservation biology, pp. 203–225, cabi, wallingford. doi:10.1079/9781845932541.020 cottrell, c.b., 1985, ‘the absence of coevolutionary associations with capensis floral element plants in the larval/plant relationships of southwestern cape butterflies’, transvaal museum monographs 4, 115−124. dacel, 2003, a conservation plan for gauteng, gauteng department of agriculture, conservation, environment and land affairs, johannesburg. dallas, h.f., 2007, ‘the influence of biotope availability on macroinvertebrate assemblages in south african rivers: implications for aquatic bioassessment’, freshwater biology 52, 370−380.doi:10.1111/j.1365-2427.2006.01684.x dallas, h.f., 2008, ‘water temperature and riverine ecosystems: an overview of knowledge and approaches for assessing biotic responses, with special reference to south africa’, water sa 34, 393−404. dallas, h.f. & day, j.a., 1993, the effect of water quality variables on riverine ecosystems: a review, technical report series no. tt 61/93, south africa, water research commission, pretoria. damgaard, j., klass, k-d., picker, m.d. & buder, g., 2008, ‘phylogeny of the heelwalkers (insecta: mantophasmatodea) based on mtdna sequences, with evidence for additional taxa in south africa’, molecular phylogenetics and evolution 47, 443−462. doi:10.1016/j.ympev.2008.01.026, pmid:18396416 daniels, s.r., picker, m.d., cowling, r.m. & hamer, m., 2009, ‘unravelling evolutionary lineages among south african velvet worms (onychophora: peripatopsis) – evidence for cryptic species complexes’, biological journal of the linnean society 97, 200−216. doi:10.1111/j.1095-8312.2009.01205.x davis, a.l.v., 2002, ‘dung beetle diversity in south africa: influential factors, conservation status, data inadequacies and survey design’, african entomology 10, 53−65. day, b., 2005, ‘the distribution of the palaeorelictual invertebrate fauna of south africa’, report for table mountain fund (project number za 5061), wwf, cape town. de kock, a.e. & giliomee, j.h., 1989, ‘a survey of the argentine ant, iridomyrmex humilis (mayr), (hymenoptera: formicidae) in south african fynbos’, journal of the entomological society of southern africa 52, 157−164. de kock, k.n. &. wolmarans, c.t., 2008, ‘invasive alien freshwater snail species in the kruger national park, south africa’, koedoe 50, 49−53. de wit, m. & stankiewicz, j., 2006, ‘changes in surface water supply across africa with predicted climate change’, science 311, 1917−1921. doi:10.1126/science.1119929, pmid:16513946 dennis, r.l.h., shreeve, t.g. & sheppard, d.a., 2007, ‘species conservation and landscape management: a habitat perspective’, in a.j. stewart, t.r. new & o.t. lewis (eds.), insect conservation biology, pp. 92–126, cabi, wallingford. dippenaar-schoeman, a.s., 1988, ‘annotated check list of the spiders (araneae) of the mountain zebra national park’, koedoe 31, 151−160. dippenaar-schoeman, a.s., 2002, baboon and trapdoor spiders of southern africa: an identification manual, plant protection research institute handbook 13, agricultural research council, pretoria. dippenaar-schoeman, a.s., 2006, ‘new records of 43 spider species from the mountain zebra national park, south africa (arachnida: araneae)’, koedoe 49, 23−28. dippenaar-schoeman, a.s. & leroy, a., 2003, ‘a check list of the spiders of the kruger national park, south africa (arachnida: araneae)’, koedoe 46, 91−100. dippenaar-schoeman, a.s., leroy, a., de jager, m. & van den berg, a., 1999, ‘spider diversity of the karoo national park, south africa (arachnida: araneae)’, koedoe 42, 31−42. dippenaar-schoeman, a.s., van der walt, a.e., le roux, e. & van den berg, a., 2005, ‘the spiders of the swartberg nature reserve in south africa (arachnida: araneae)’, koedoe 48, 77−86. dobson, a., 2005, ‘monitoring global rates of biodiversity change: challenges that arise in meeting the convention on biological diversity (cbd) 2010 goals’, philosophical transactions of the royal society london b 360, 229−241. doi:10.1098/rstb.2004.1603, pmid:15814342 , pmcid:1569458 driver, a., maze, k., rouget, m., lombard, a.t., nel, j., turpie, j.k., et al., 2005, national spatial biodiversity assessment 2004: priorities for biodiversity conservation in south africa, strelitzia 17, south african national biodiversity institute, pretoria. droege, s., cyr, a. & larivée, j., 1998, ‘checklists: an under-used tool for the inventory and monitoring of plants and animals’, conservation biology 12, 1134−1138. doi:10.1046/j.1523-1739.1998.96402.x edge, d.a., 2005, ‘butterfly conservation in the southern cape’, metamorphosis 16, 28−46. engelbrecht, i.a., 2010, ‘invertebrate species inventories in protected area management: are they useful?’, african entomology 18, 235−245. doi:10.4001/003.018.0203 forsyth, g. & van wilgen, b., 2008, ‘the recent fire history of table mountain national park and implications for fire management’, koedoe 50, 3−9. goodman, p.s., 2000, ‘determining the conservation value of land in kwazulu-natal’, final report, kwazulu-natal nature conservation service, pietermaritzburg. grimaldi, d. & engel, m.s., 2005, evolution of the insects, cambridge university press, cambridge. haddad, c.r., dippenaar-schoeman, a.s. & wesołowska, w., 2006, ‘a checklist of the non-acarine arachnids (chelicerata: arachnida) of the ndumo game reserve, maputoland, south africa’, koedoe 49, 1−22. hamer, m.l., 1998, ‘checklist of southern african millipedes’, annals of the natal museum 39, 11−82. hamer, m.l. & slotow, r.h., 2002, ‘conservation application of existing data for south african millipedes (diplopoda)’, african entomology 10, 29−42. hamer, m. & slotow, r., 2009, ‘a comparison and conservation assessment of the high-altitude grassland and forest-millipede (diplopoda) fauna of the south african drakensberg’, soil organisms 81, 701−717. hamer, m.l., samways, m.j. & ruhberg, h., 1997, ‘a review of the onychophora of south africa, with discussion of their conservation’, annals of the natal museum 38, 283−312. hamer, m.l., slotow, r.h. & lovell, s., 2006, ‘the south african savanna millipede (diplopoda) fauna: taxonomic diversity, endemism and spatial and temporal effects on conservation assessments’, norwegian journal of entomology 53, 321−324. hanski, i. & pöyry, j., 2007, ‘insect populations in fragmented habitats’, in a.j. stewart, t.r. new & o.t. lewis (eds.), insect conservation biology, pp. 175–202, cabi, wallingford. doi:10.1079/9781845932541.0175 henning, s.f. & henning, g.a., 1989, ‘south african red data book: butterflies’, south african national scientific program report no. 158, foundation for research development, pretoria. henning, g.a., terblanche, r.f. & ball, j.b. (eds.), 2009, south african red data book: butterflies, south african national biodiversity institute, pretoria. herbert, d. & kilburn, d., 2004, field guide to the land snails and slugs of eastern south africa, natal museum, pietermaritzburg. horn, j.l., plisko, j.d. & hamer, m.l., 2007, ‘the leaf-litter earthworm fauna (annelida: oligochaeta) of forests in limpopo province, south africa: diversity, communities and conservation’, african zoology 42, 172−179. doi:10.3377/1562-7020(2007)42[172:tlefao]2.0.co;2 kaspari, m. & majer, j.d., 2000, ‘using ants to monitor environmental change’, in d. agosti, j.d. majer, l.e. alonso & t.r. schultz (eds.), ants: standard methods for measuring and monitoring biodiversity, pp. 89–98, smithsonian institution press, washington dc. kenis, m., auger-rozenberg, m.-a., roques, a., timms, l., péré, c., cock, m.j.w., et al., 2009, ‘ecological effects of invasive alien insects’, biological invasions 11, 21−45. doi:10.1007/s10530-008-9318-y ketley, z.a., 2009, ‘stream invertebrates and water temperature: evaluating thermal tolerances in the cape floristic region (south africa) – implications of climate change’, msc thesis, dept. of zoology, university of cape town. kier, g., kreft, h., lee, t.m., jetz, w., ibisch, p.i., nowicki, c., et al., 2009, ‘a global assessment of endemism and species richness across island and mainland regions’, proceedings of the national academy of sciences of the united states of america 106, 9322−9327. doi:10.1073/pnas.0810306106,pmid: 19470638 , pmid: 2685248 koch, s.o., chown, s.l., davis, a.l.v., endrödy-younga, s. & van jaarsveld, a.s., 2000, ‘conservation strategies for poorly surveyed taxa: a dung beetle (coleoptera, scarabaeidae) case study from southern africa’, journal of insect conservation 4, 45−56. doi:10.1023/a:1009634318926 kremen, c., colwell, r.k., erwin, t.l., murphy, d.d., noss, r.f. & sanjayan, m.a., 1993, ‘terrestrial arthropod assemblages: their use in conservation planning’, conservation biology 7, 796−808. doi:10.1046/j.1523-1739.1993.740796.x lawrence, r.f., 1953, the biology of the cryptic fauna of forests. with special reference to the indigenous forests of south africa, a.a. balkema, cape town. losey, j.e. & vaughan, m., 2006, ‘the economic value of ecological services provided by insects’, bioscience 56, 311−323. doi:10.1641/0006-3568(2006)56[311:tevoes]2.0.co;2 lovell, s., hamer, m., slotow, r. & herbert, d., 2007, ‘assessment of congruency across invertebrate taxa and taxonomic levels to identify potential surrogates’, biological conservation 139, 113−125. doi:10.1016/j.biocon.2007.06.008 lovell, s., hamer, m., slotow, r. & herbert, d., 2010, ‘assessment of sampling approaches for a multi-taxa invertebrate survey in a south african savanna-mosaic ecosystem’, austral ecology 35, 357–370. doi:10.1111/j.1442-9993.2009.02052.x lowe, s., browne, m., boudjela, s. & de poorter, m., 2000, ‘100 of the world’s worst invasive alien species. a selection from the global invasive species database’, the invasive species specialist group (issg), iucn, gland. lyons, c., picker, m.d. & carrick, p., 2008, ‘evaluating restoration success of alluvial diamond-mined sites using invertebrate community indicators’, mine closure proceedings 3, 821−830. macdonald, i.a.w., reaser, j.k., bright, c., neville, l.e., howard, g.w., murphy, s.j., et al. (eds.), 2003, ‘invasive alien species in southern africa: national reports & directory of resources’, global invasive species programme, cape town. mcgeoch, m.a., 1998, ‘the selection, testing and application of terrestrial insects as bioindicators’, biological reviews 73, 181−201. doi:10.1017/s000632319700515x, doi:10.1111/j.1469-185x.1997.tb00029.x mcgeoch, m.a., 2002, ‘insect conservation in south africa: an overview’, african entomology 10, 1−10. mcgeoch, m.a., 2007, ‘insects and bioindication: theory and practice’, in a.j. stewart, t.r. new & o.t. lewis (eds.), insect conservation biology, pp. 144−174, cabi, wallingford. doi:110.1079/9781845932541.0144 mcgeoch, m.a., van rensburg, b.j. & botes, a., 2002, ‘the verification and application of bioindicators: a case study of dung beetles in a savanna ecosystem’, journal of applied ecology 39, 661−672. doi:10.1046/j.1365-2664.2002.00743.x mcgeoch, m.a., le roux, p.c., hugo, a.e. & chown, s.l., 2006, ‘species and community responses to short-term climate manipulation: microarthropods in the sub-antarctic’, austral ecology 31, 719−731. doi:10.1111/j.1442-9993.2006.01614.x mcgeoch, m.a., dopolo, m., novellie, p., hendriks, h., freitag, s., ferreira, s., et al., 2011, ‘a strategic framework for biodiversity monitoring in sanparks’, koedoe 53(2), art. #991, 10 pages. doi:10.4102/koedoe.v53i2.991 mckie, b.g., cranston, p.s. & pearson, r.g., 2004, ‘gondwanan mesotherms and cosmopolitan eurytherms: effects of temperature on the development and survival of australian chironomidae (diptera) from tropical and temperate populations’, marine & freshwater research 55, 759−768. doi:10.1071/mf04023 mound, l.a. & gaston, k.j., 1993, ‘conservation and systematics the agony and the ecstasy’, in k.j. gaston, t.r. new & m.j. samways (eds.), perspectives on insect conservation, pp. 185–196, intercept, andover. nel, j.l., roux, d.j., maree, g., kleynhans, c.j., moolman, j., reyers, b., et al., 2007, ‘a systematic conservation assessment of the ecosystem status and protection levels of main rivers in south africa’, diversity and distributions 13, 341−352. doi:10.1111/j.1472-4642.2007.00308.x new, t.r., 1995, an introduction to invertebrate conservation biology, oxford university press, oxford. new, t.r., 1998, invertebrate surveys for conservation, oxford university press, oxford. odendaal, l.j., haupt, t.m. & griffiths, c.h., 2008, ‘the alien invasive land snail theba pisana in the west coast national park: is there cause for concern?’, koedoe 50, 93−98. panzer, r. & schwartz, m.w., 1998, ‘effectiveness of a vegetation-based approach to insect conservation’, conservation biology 12, 693−702. doi:10.1046/j.1523-1739.1998.97051.x parr, c.l. & chown, s.l., 2001, ‘inventory and bioindicator sampling: testing pitfall and winkler methods with ants in a south african savanna’, journal of insect conservation 5, 27−36. doi:10.1023/a:1011311418962 parr, c.l., bond, w.j. & robertson, h.g., 2002, ‘a preliminary study of the effect of fire on ants (formicidae) in a south african savanna’, african entomology 10, 101−111. parr, c.l., robertson, h.g., biggs, h.c. & chown, s.l., 2004, ‘response of african savanna ants to long-term fire regimes’, journal of applied ecology 41, 630−642. doi:10.1111/j.0021-8901.2004.00920.x picker, m.d. & samways, m.j., 1996, ‘faunal diversity and endemicity of the cape peninsula, south africa a first assessment’, biodiversity and conservation 5, 591−606. doi:10.1007/bf00137611 prendini, l., 2005, ‘scorpion diversity and distribution in southern africa: pattern and process’, in b.a. huber, b.j. sinclair & k.-h. lampe (eds.), african biodiversity: molecules, organisms, ecosystems, pp. 25−28, springer, amsterdam. pringle, e.l.l., henning, g.a., & ball, j.b., 1994, pennington’s butterflies of southern africa, 2nd edn., struik winchester, cape town. procheş, s., forest, f., veldtman, r., chown, s.l., cowling, r.m., johnson, s.d., et al., 2009, ‘dissecting the plant–insect diversity relationship in the cape’, molecular phylogenetics and evolution 51(1), 94−99. pryke, j.s., 2008, ‘conservation of invertebrate fauna of the cape peninsula’, phd thesis, dept. of conservation ecology and entomology, stellenbosch university. pryke, j.s. & samways, m.j., 2008, ‘conservation of invertebrate biodiversity on a mountain in a global biodiversity hotspot, cape floral region’, biodiversity and conservation 17, 3027−3043. doi:10.1007/s10531-008-9414-4 pryke, j.s. & samways, m.j., 2009a, ‘conservation of the insect assemblages of the cape peninsula biodiversity hotspot’, journal of insect conservation 13, 627−641. doi:10.1007/s10841-009-9213-6 pryke, j.s. & samways, m.j., 2009b, ‘recovery of invertebrate diversity in a rehabilitated city landscape mosaic in the heart of a biodiversity hotspot’, landscape and urban planning 93, 54−62. doi:10.1016/j.landurbplan.2009.06.003 pryke, j.s. & samways, m.j., 2010, ‘significant variables for the conservation of mountain invertebrates’, journal of insect conservation 14, 247−256. doi:10.1007/s10841-009-9253-y raharinjanahary, d., 2007, ‘impact of argentine ants (linepithema humile mayr) on saproxylic invertebrates in afromontane forest and pine plantation of the cape peninsula (south africa)’, msc thesis, dept. of zoology, university of cape town. revenga, c., campbell, i., abell, r., de villiers, p. & bryer, m., 2005, ‘prospects for monitoring freshwater ecosystems towards the 2010 targets’, philosophical transactions of the royal society of london b 360, 397−413. doi:10.1098/rstb.2004.1595, pmid:15814353 , pmid: 1569454 rohr, j.r., kim, k. & mahan, c., 2007, ‘developing a monitoring program for invertebrates: guidelines and a case study’, conservation biology 21, 422−433. doi:10.1111/j.1523-1739.2006.00578.x, pmid: 17391192 rösch, m., chown, s.l. & mcgeoch, m.a., 2001, ‘testing a bioindicator assemblage: gall-inhabiting moths and urbanization’, african entomology 9, 85−94. samways, m.j., 1993a, ‘a spatial and process sub-regional framework for insect and biodiversity conservation research and management’, in k.j. gaston, t.r. new & m.j. samways (eds.), perspectives on insect conservation, pp. 1−28, intercept, andover. samways, m.j., 1993b, ‘dragonflies (odonata) in taxic overlays and biodiversity conservation’, in k.j. gaston, t.r. new & m.j. samways (eds.), perspectives on insect conservation, pp. 111−124, intercept, andover. samways, m.j., 2005, insect diversity conservation, cambridge university press, cambridge. doi:10.1017/cbo9780511614163 samways, m.j., 2006, ‘national red list of south african odonata’, odonatologica 35, 341−368. samways, m.j., 2008, dragonflies and damselflies of south africa, pensoft, sophia. samways, m.j. & sharratt, n.j., 2010, ‘recovery of endemic dragonflies after removal of invasive alien trees’, conservation biology 24, 267−277. doi:10.1111/j.1523-1739.2009.01427.x ,pmid: 20121847 samways, m.j., mcgeoch, m.a. & new, t.r., 2010, insect conservation: a handbook of approaches and methods, oxford university press, oxford. scholes, r.j. & walker, b.h., 1993, an african savanna: synthesis of the nylsvley study, cambridge university press, cambridge. doi:10.1017/cbo978051156547 scholtz, c.h. & chown, s.l., 1993, ‘insect conservation and extensive agriculture: the savanna of southern africa’, in k.j. gaston, t.r. new & m.j. samways (eds.), perspectives on insect conservation, pp. 75−96, intercept, andover. scholtz, c.h. & chown, s.l., 1995, ‘insects in southern africa: how many species are there?’, south african journal of science 91, 124−126. scholtz, c.h. & holm, e., 1985, insects of southern africa, butterworths, durban. sharratt, n.j., picker, m.d. & samways, m.j., 2000, ‘the invertebrate fauna of the sandstone caves of the cape peninsula (south africa): patterns of endemism and conservation priorities’, biodiversity and conservation 9, 107−143. doi:10.1023/a:1008968518058 simaika, j.p. & samways, m.j., 2008, ‘valuing dragonflies as service providers’, in a. cordoba-aguilar (ed.), dragonflies: model organisms for ecological and evolutionary research, pp. 109−123, oxford university press, oxford. simaika, j.p. & samways, m.j., 2009a, ‘an easy-to-use index of ecological integrity for prioritizing streams for conservation action’, biodiversity and conservation 18, 1171−1185. doi:10.1007/s10531-008-9484-3 simaika, j.p. & samways, m.j., 2009b, ‘reserve selection using red listed taxa in three global biodiversity hotspots: dragonflies in south africa’, biological conservation 142, 638−651. doi:10.1016/j.biocon.2008.11.012 simaika, j.p. & samways, m.j., 2011, ‘comparative assessment of indices of freshwater habitat conditions using different invertebrate taxon sets’, ecological indicators 11, 370−378. doi:10.1016/j.ecolind.2010.06.005 sithole, h., smit, i.p.j. & parr, c.l., 2010, ‘preliminary investigations into a potential ant invader in kruger national park, south africa’, african journal of ecology 48, 736−743. smith, j., samways, m.j. & taylor, s., 2007, ‘assessing riparian quality using two complementary sets of bioindicators’, biodiversity and conservation 16, 2695−2713. doi:10.1007/s10531-006-9081-2 speight, m.c.d. & castella, e., 2001, ‘an approach to interpretation of lists of insects using digitised biological information about the species’, journal of insect conservation 5, 131−139. doi:10.1023/a:1011399800825 stiller, m., 2002, ‘leafhopper (hemiptera: cicadellidae) diversity in the fynbos biome of south africa’, denisia 4, 379−400. telford, m.j. & littlewood, d.t.j., 2008, ‘the evolution of the animals: introduction to a linnean tercentenary celebration’, philosophical transactions of the royal society of london b 363, 1421−1424. doi:10.1098/rstb.2007.2231, pmid:18192193 , pmcid:2394567 thomas, j.a., 2005, ‘monitoring change in the abundance and distribution of insects using butterflies and other indicator groups’, philosophical transactions of the royal society of london b 360, 339−357. doi:10.1098/rstb.2004.1585, pmid: 15814349 , pmid: 1569450 thompson, m.w., 2008, kzn province land-cover mapping (from spot 2 / 4 satellite imagery 2005–2006), data users report and meta data (version 2.0), geoterraimage, pretoria. underwood, e.c. & fisher, b.l., 2006, ‘the role of ants in conservation monitoring: if, when, and how’, biological conservation 132, 166−182. doi:10.1016/j.biocon.2006.03.022 uys, c., hamer, m. & slotow, r., 2006, ‘the effect of burn area on invertebrate recolonisation in grasslands in the drakensberg, south africa’, african zoology 41, 51−65. doi:10.3377/1562-7020(2006)41[51:eobaoi]2.0.co;2 uys, c., hamer, m. & slotow, r., 2009, ‘turnover in invertebrate species composition over different spatial scales in afrotemperate forest in the drakensberg, south africa’, african journal of ecology 47, 341−351. doi:10.1111/j.1365-2028.2008.00968.x uys, c., hamer, m. & slotow, r., 2010, ‘step process for selecting and testing surrogates and indicators of afrotemperate forest invertebrate diversity’, plos one 5(2), e9100. doi:10.1371/journal.pone.0009100, pmid:20161757, pmcid:2817749 valéry, l., fritz, h., lefeuvre, j.-c. & simberloff, d., 2008, ‘in search of a real definition of the biological invasion phenomenon itself’, biological invasions 10, 1345−1351. doi:10.1007/s10530-007-9209-7 vane-wright, r.i., 1993, ‘systematics and the conservation of biodiversity: global, national and local perspectives’, in k.j. gaston, t.r. new & m.j. samways (eds.), perspectives on insect conservation, pp. 197−212, intercept, andover. veldtman, r., mcgeoch, m.a. & scholtz, c.h., 2002, ‘variability in cocoon size in southern african wild silk moths: implications for sustainable harvesting’, african entomology 10, 127−136. vilà, m., basnou, c. & pyšek, p., josefsson, m., genovesi, p., gollasch, s., et al., 2010, ‘how well do we understand the impacts of alien species on ecosystem services? a pan-european, cross-taxa assessment’, frontiers in ecology and the environment 8, 135−144. doi:10.1890/080083 walker, b. & salt, d., 2006, resilience thinking: sustaining ecosystems and people in a changing world, island press, washington dc. warren, m.s., bourn, n., brereton, t., fox, r., middlebrook, i. & parsons, m.s., 2007, ‘what have red lists done for us? the values and limitations of protected species listing for invertebrates’, in a.j. stewart, t.r. new & o.t. lewis (eds.), insect conservation biology, pp. 76−91, cabi, wallingford. doi:10.1079/9781845932541.0076 wesołowska, w. & haddad, c.r., 2009, ‘jumping spiders (araneae: salticidae) of the ndumo game reserve, maputaland, south africa’, african invertebrates 50, 13−103. wilson, r.j., davies, z.g. & thomas, c.d., 2007, ‘insects and climate change: processes, patterns and implications for climate change’, in a.j. stewart, t.r. new & o.t. lewis ot (eds.), insect conservation biology, pp. 245−279, cabi, wallingford. wilson, j.r.u., dormontt, e.e., prentis, p.j., lowe, a.j. & richardson, d.m., 2009, ‘something in the way you move: dispersal pathways affect invasion success’, trends in ecology and evolution 24(3), 136−144. doi:10.1016/j.tree.2008.10.007 woiwod, i.p. & harrington, r., 1994, ‘flying in the face of change: the rothamsted insect survey’, in r. leigh & a. johnston (eds.), long-term experiments in agricultural and ecological sciences, pp. 321−342, cab international, wallingford. woodroffe, r. & ginsberg, j.r., 1998, ‘edge effects and the extinction of populations inside protected areas’, science 280, 2126−2128. doi:10.1126/science.280.5372.2126, pmid:9641920 wright, m.g. & samways, m.j., 1998, ‘insect species richness tracking plant species richness in a diverse flora: gall-insects in the cape floristic region, south africa’, oecologia 115, 427−433. doi:10.1007/s004420050537 wright, j.f., sutcliffe, d.w. & furse, m.t., 2000, assessing the biological quality of fresh waters, the freshwater biological association, ambleside. filelist convert a pdf file! koedoe 19: 75-82 (i976) the re-introduction of springbok antidorcas marsupialis into south african national parksa documentation g. de graaff and b. l. penzhorn national parks board of trustees p.o . box 787 pretoria 0001 abstract the introduction and establishment of springbok populations in four south african national parks are discussed. springbok have £ailed to establish themselves in the addo elephant national park but are thriving in the mountain zebra, golden gate highlands and bontebok national parks, although the latter park is extralimital to their original range. introduction the policy of the national parks board of trustees is to reintroduce wildlife species in newly acquired national parks only if the particular species in question occurred in an area formerly and naturally. the extent to which this approach has been implemented for the springbok (antidorcas marsupialis) over the years will be discussed below. presently there are 10 national parks administered by the board and springbok are encountered in six. the national parks where they do not occur allude to kruger, the tsitsikama parks and groenkloof. the reintroductions and subsequent histories of the various springbok populations introduced into mountain zebra, addo, bontebok and golden gate will be sketched and the situation pertaining to augrabies will be referred to. the large, resident springbok population (c. 20000 individuals) occurring in the kalahari gemsbok is excluded in the' ensuing paragraphs. re-introduction histories not only make interesting reading but are of practical importance as well provided the facts and data are carefully and meticulously recorded. the published accounts pertaining to species handled by the national parks board are few in number and for detailed information the reader is referred to barnard and van der walt (i 96 1), van niekerk and pienaar (1962), pienaar and van niekerk (1963) and penzhorn (i 971). 75 material and methods the data presented in this paper have been compiled from records kept in the national parks board head office in pretoria. it included perusal of annual reports, minutes of board meetings, quarterly reports, monthly returns by rangers as well as diaries kept by the latter. this information was also amplified by personal experience and observation during handling and culling operations. both the addo elephant national park (near port elizabeth) and the bontebok national park (near bredasdorp) were established in 1931. proclamation of the mountain zebra national park (near cradock) occurred ill 1937. during 1960 the original bontebok national park was superseded by the present park (near swellendam) and followed by the realisation of the golden gate highlands national park (near bethlehem) and the augrabies falls national park (near upington) in 1963 and 1966 respectively. chronologically, springbok were reintroduced to mountain zebra (1941), addo (1956), bontebok (1960) and golden gate (1964) . 1200 1000 800 en ~ 600 ;;; z « ::; 400 a: w ld ::; ~ 200 1940 1945 1950 1955 year 196 a 1965 1970 1975 fig. 1. populatioll histol), of springbok antidorcas marsupialis in the mountain zebra national park, rsa. 76 • results mountain zebra national park. (mznp) this park falls well within the historical range of the springbok and during 1941 eight animals were released into this park having been donated b y e. bowker esq. of bedford (not grahamstown as stated by penzhorn 197 i). the introduction was done as an experiment and their numbers increased to 43 in 1946 . the subsequent increase in the population is shown in fig. 1. by 1954 their numbers had reached 300+ and the first control of their numbers took place. a total of 106 individuals were captured and sold to farmers cw r8,00 each for restocking on their farms. the remaining 250 animals consisted of a 2 « « : 1 0 ratio . eleven animals were transported to the addo elephant national park in 1956 and another 11 in 1958 . 100 80 (i) ~ 80 ~ z < u.. 040 a: w iii ~ ;:) z 20 1956 10 65 70 75 year fig. 2. po pul a ti o n hi story o f springbok antidorc(lj mars upialis m the add o elephant nati o na l park , rsa . 77 towards the end of 1961 they reached a peak with 450 individuals and some 205 were caught and made available to interested farmers. twenty animals were translocated to the bontebok national park. during 1962 the overall population strength was further diminished by removal of 155 animals leaving a remnant population of c. 90 animals . annual increases were also harvested and by 1964 only 70 animals remained. in 1964 the park was substantially enlarged by the acquisition of adjoining land. within eight years (1974) the springbok population increased to an all-time peak of c. 1267 animals, which prompted the initiation of control procedures. during 1972 and 1973 a total of 823 individuals were removed, mainly by shooting (penzhorn 1974), and by 1975 the population had been reduced to 345 animals. addo elephant national park (aenp) eleven springbok were translocated to this park from the flourishing population in the mznp in 1956. these were later augmented by further springbok (16) from the mznp (1958) and the nduli nature reserve, umtata, transkei (penzhorn 1971). the springbok increased steadily to ul ....i « :i z « 200 150 o 100 ii: w iii ::ii :;) z 110 1900 1965 1970 1975 year fig. 3. population history of springbok antidorcas marsupialis in the bontebok national park, rsa. 78 over 100 individuals by 1962 (fig. 2) but by 1964 more than 60% of the population succumbed, mainly after contracting heartwater. since then there has been a steady decrease and only a few individuals remained in 1974 (9 9) which was further reduced to a single female in 1975. bontebok national park (bnp) the springbok in the bnp were also introduced from the mznp. a total of 20 animals was transported, seven of which died shortly afterwards. the population had increased to 246 by 1974 (fig. 3). during 1966, animals from this population were transferred to the golden gate highlands national park. c/) -i ~ :e z 200 150 ~ 100 u. o c: w co :e :::> z 50 1964 66 68 year 70 72 74 fig. 4. population history 01 springbok antidorcas marsupialis in the golden gale highland national park, rsa . golden gate highlands national park (gghnp) this population also hails from mznp originally, when 10 animals were introduced in 1964. these numbers were augmented by a mixed bag from mznp and bnp respectively in 1966/67 and in 1968/69 the 79 population stood at 86 animals. subsequently, they have increased to just over 200 individuals by the end of 1974. augrabies falls national park (afnp) although this park falls within the past and present range of distribution of springbok, this species has not been re-introduced as yet. there are small resident herds on adjoining farms and these animals occasionally stray into the park. discussion it is clear that although we do have a fair amount of information available amassed over the years, there are glaring gaps in the data, pertaining especially to dynamics of the different populations. valuable opportunities have been missed for constructive studies pertaining to the process of adaptation which occurs in populations placed in new habitats. there is no detailed information available on the social organization of herds within the populations, or, for that matter on ecology and ethology in the broad sense of the words. this emphasizes once again the importance of meticulous documentation and the sustained upkeep thereof. to what extent has the re-introduction of springbok been successful? there seems to be no uncertainty of the positive results obtained in the mzn p. since 1941, some 1750 animals have been removed from the population, while the populations in three other parks also originated from this bedford stock. this response is and was to be expected, for the mznp falls right within the typical springbok habitat and therefor in the historical range of distribution of the species. although springbok occurred in the vicinity of the aenp in historic times (skead 1958), the park itself does not represent ideal springbok habitat. the dense addo bush is unsuited to the plains-inhabitating springbok and attempts at artifically creating habitat for certain antelope by clearing the natural vegetation have fortunately been abandoned. sclater (1900) stated that springbok did not occur south of the swartberg. bateman (i 960, who made a survey of the mammals indigenous to the swellendam district, stated that springbok are not generally considered to have inhabited this region. in view of the stated policy of the national parks board, it is therefore surprising that springbok were introduced to the bnp shortly after the park was established at swellendam in 1960. according to du plessis (969), the bnp falls outside the original distribution of springbok, while ansell (1971) termed this introduction " .. . very probably, outside the natural range of the species." although the springbok population in the bnp is well-established and flourishing, its entire removal is recommended . in contrast, the gghnp falls well within the past and present range of springbok distribution and all accounts point to a well-adapted population . the finite size of the park, however, has to be kept in mind and in 80 all probability their numbers will have to be reduced considerably in due course. although earlier documentation of springbok introductions was incomplete, this is partially levelled-up by recent work. in a pilot study of the parasitology and pathology of springbok in mznp, young, zumpt, basson, erasmus, boyazoglu and boomker (1973) reported 13 species of parasitic metozoa and two protozoa. pathological effects were described where applicable. serological tests for virus diseases were negative but biochemical analyses of liver samples suggest deficiencies of mn, co, mg and especially cu in the diet. a serious by-product of the introduction of springbok to the bnp is the accompanying infestation of bontebok damaliscus dorcas dorcas with parasites associated with springbok. the lungworm dictyocaulus magna was not recorded from bontebok before they were transferred to swellendam (barnard et al. 1961). during 1960, 20 springbok were also introduced and d. magna, " ... a well known parasite of the springbok ... " (verster, imes and smit 1975) was undoubtedly introduced at the same time. in this case the introduction of springbok into the bnp, beyond their original range was not only ill-conceived but proved to be detrimental to the bontebok, the raison d'etre of this park. references ansell, w. f. h. 1971. order artiodactyla. in: ]. meester and h. w. setzer (eds). the mammals of africa an identification manual. washington, d.c.: smithsonian institution press. barnard, p.]. and k. van der walt. 1961. translocation of the bontebok (damaliscus pygargus) from bredasdorp to swellendam. koedoe 4: 105-109. bateman, j. a. 1961. the mammals occuring in the bredasdorp and swellendam districts, c.p., since european settlement. koedoe 4: 78-100. du plessis, s. f. 1969. the past and present geographical distribution of the perissodactyla and artiodactyla of southern africa . m.sc. thesis, university of pretoria (unpublished). penzhorn, b. l. 1971. a summary of the re-introduction of ungulates into south african national parks (to 31 december 1970). koedoe 14: 145-159. penzhorn, b. l. 1974. sex and age composition and dimensions of the springbok (antidorcas marsupialis) population in the. mountain zebra national park.}. sth . afr. wildl. mgmt ass. 4 :63-65. pienaar, u. de v. and]. w. van niekerk. 1963. the capture and translocation of three species of wild ungulates in the eastern transvaal with special reference to r05-2807/b-5f (roche) as a tranquillizer in game animals. koedoe 6 :83-90. 81 sclater, w. l. 1900. the mammals oj south africa. london: r. h. porter. skead, c. j. 1958. mammals of the uitenhage and cradock districts, c.p. in recent times. koedoe 1: 19-59. van niekerk, j. w. and u. de v. pienaar. 1962. adaptations of the immobilizing technique to the capture, marking and translocation of game animals in the kruger national park. koedoe 5: 137-143. verster, a., g. d. imes andj. p.j. smit. 1975. helminths recovered from the bontebok, damaliscus dorcas dorcas (pallas, 1766) . onderstepoortj. vet . res . 42 :29-32. young, e., f. zumpt, p. a. basson, b. erasmus, p. a. boyazoglu and j. boomker. 1973. notes on the parasitology, pathology and bio-physiology of springbok in the mountain zebra national park. koedoe 16 : 195-198. 82 page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 filelist convert a pdf file! koedoe 19: 185-188(1976) lewensloop van 'n elandbevolking taurotragus oryx in die bontebok nasionale park g de graaff n asionale p arkeraad posbus 787 pretoria 0001 p t van der walt ajdeling natuurbewaring privaatsak x404 skukuw 1j50 l j van zyl bontebok nasionale park posbus 149 swellendam 6740 abstract a description of the life history of an eland population taurotragus oryx in the bontebok national park, near swellendam is given. the reasons which led to the eventual removal of the population from the park are discussed. lnleiding vanuit historiese gegewens (du plessis 1969) is dit duidelik dat die eland taurotragus oryx in die meeste dele van die suidelike kaapprovinsie voorgekom het. daar is egter geen aanduidings hoe volop die diere was nie en teen die einde van die agtiende eeu was die spesies reeds skaars in die suidkaap. hierdie proses van getalle-afname het later daartoe gelei dat elande hier beperk was tot beskermde gebiede. een so 'n gebied was die bontebok nasionale park (bnp), naby swellendam, waar elande gedurende 1960 hervestig is vanuit die kalahari-gemsbok nasionale park (kgnp) (penzhorn 1971) om die wildbevolking van die bnp verteenwoordigend te laat wees van wat daar vroeer in die geweste voorgekom het. die verblyf van die diere in die bnp is op 1975.05.25 kortgeknip as gevolg van redes wat in hierdie verslag omlyn word. 185 ta u cl i lt'wt'!) ~tabd van die elande in die bontebok .'jasiuoale park ,\f wdikediat' ji rouilkt'dlrr( aldj .lllar r l j('i/' ll/i('i a/11<11f1i r"{<l111 i'rr1<"i{ww' i"o!um,\\f ,\/ ft'r}ang a"a l!,jfrl vo/w,h5f ivl eer;arig ka lwfr.l 1960 4 ela ndc ' in palk herv('s!i g vd llu it k,lbha ri g('lll~b()k na ~ i otlal( ' palk 1961 ku c'i c' dr'i)i;lig icjti :! lka lw('! ~r"djc)1'(' 100 jfl63 s \'('!' sc ingc h rin g va nuit di(' kal a hari gelll sho k '\ja sio nalt' p;n k 2 kahil's g-e b(h -{' mei en j lilli{' " 100 1964 j \'ol wd s.<;(' koei \"r('k j j()llg: bul in gt'n:g illet '1\ li\ll'r bul g('d()od 2 k;tll i('~ gd)()r<' 4(11 7\1j 13 2r 19fi s .3 ka llie~ gl'hore iii 37 1966 i mlwa,,, ,,,,i imd} ,>il"" he('n i \'olwas\t' b tl l breck 1l1t't1g lwell i n)h,'a ~,l' koci vr e k 3 b ui('s gt-' boh: augustu s en oktober "" :-1 1 ih 2) 16 37 ..... 196 7 1 volw;tsse bul vrek 00 01 4 blfie's g-ebon:' 61 -1 if) 57 1968 3 kalfi t's )l;t'bon' 22 33 1969 l ouhu[v r ek (fi 9 .07_201 4 ka ltit'~ g ebolt' julie en augustus 8': -1 i " i i vnsuip in ill hret'd t'ri vierl 1(1) " so 1970 3 kalfics geboh' 1211\ 16 , oil koci vrek 2s 1971 2 kalfics gebore " 2(1j 12 1(-1) [(-i ) 2s 16 , ind, 1 yt'rse vrek 1971 t-.-1ci!i kaltil'~ gd)()rt , i bul \'t'!-k(l ()p de s(,tllbcr 3 kalfi es \te k 1 bul \tek. oucienjolll dc st' lil ber 19 71 12(3) 5(· 3 ) 1:2(-j) 23 41 1 bul geskict vi,post mort e lll b a ic o uoi ermei 1972 i koc i " rek. o uderdo m 19 73 pn pll las ie sanl('std li n g 13 29 40 in r"\' o\'cmb cr 1973 1974 i bul en :2 koeie vcrkoop 1 bul c n i koci g(' ski('t vir p. hl 101-2) 13(-.)) 25 23 2 volwasst' ko eic v rc k 1975 popuiasic saole~tejling op 197.'>.05.25 toe al die dand e uit p a rk vci-wyder i$ ._---~ .---------resultate en bespreking met die totstandkoming van die nuwe bnp te swellendam in 1960 (van rensburg 1975), het die gebied 1 435 ha beslaan en is vier elande (twee jong bulle en twee jong verse) daar hervestig. in 1962 is die eerste kalfles gebore (tabel 1). in 1963 is die getalle verder versterk deur die inbring van vyf versies, ook vanuit die kgnp. in 1964 is die bnp vergroot tot 2786 ha en met die verloop van jare het die trop aangegroei tot 'n maksimum van 29 individue. die wel-en-wee van die elande in die bnp word in tabel 1 weergegee waarin die populasie-struktuur vanaf 1960-1975 verstrek word tesame met gegewens oor reproduksie en mortaliteit. die elande het deurgaans 'n lae kalfpersentasie van 41% gehandhaaf terwyl hul algemene bouvorm ook veel te wense oorgelaat het. dit spruit skynbaar vanuit 'n voedingstekort wat aanleiding gee tot sekondere simptome soos 'n lae reproduksie en afname in massa. hierdie tekort vererger hoe ouer die dier word want dit was deurentyd opvallend dat die .iong elande se kondisie goed is. dit dui daarop dat die probleem gesentreer het rondom 'n proteientekort wat veroorsaak word deur 'n verlaagde voedselinname hoe ouer die dier word. dit is bekend (louw, steenkamp en steenkamp 1968) dat die ruproteieninhoud van smaaklike bossies en struike oor die algemeen hoog is. in die bnp is die plantegroeisamestelling egter van so 'n aard dat daar weinig smaaklike struike en bossies voorkom (de graaff, van der wal' en van zyl 1976) maar heelwat smaaklike grasse. dit bring mee jat die clande tog noodwendigerwys gaan peusel aan hierdie bossies tep. einde te pro beer voldoen aan sy proteienbehoeftes. hierdie genoemde bossies en struike besit deurgaans 'n hoe eterekstrak (louw et al. 1968). dit, tesame met die teenwoordigheid va~: n hoc persentasie harde grasse, lei tot 'n relatief vinnige slytasie van die elande sc tande. die geslyte gebit van die dier gee weer aanleiding tot 'n verlaagde inname van smaaklike grasse wat redelik volop in die bnp voorkom. aangesien 'n grasblaar primer uit ruvesel bestaan, gee hierdie verlaagde inname aanleiding tot 'n verlaagde energie omdat ruvesel die belangrikste bron van energie vir enige herkouer uitmaak. om 'n aanduiding te gee van die rol wat siektes en/of parasiete in die elandpopulasie gespeel het, is die volgende bevindings van 'n veeartsenykundige ondersoek op 'n ou bul gedurende 1972 van waarde: (i) die dier was baie maer, moontlik as gevolg van 'n voedingstekort. geen vetreserwes behalwe om die hart nie. (ii) behalwe vir 'n paar volwasse paramphistome (peervormige slakke) was die dier vry van inwendige parasiete. (iii) eiertellings ten opsigte van wurms in die mis was negatief. (iv) uitwendige parasiete het hoofsaaklik uit die volgende bosluise bestaan: rhipicephalus evertsi evertsi hyalomma turanicum (v) die bloedwaardes was soos volg: 187 n mg/l 00 illi g/loo ml po, mg ca na k total e plasma hb" % prote'ien hie' ,~ 1,3 2, 2 10,0 326 20,3 7,4 11,0 32 die waardes ie redelik goed behalwe in die fosfate en ca, asook die hb-waarde wat moontlik 'n bietjie laag is. ':' haemoglobin ':' ':' haemokrit (vi ) die lewer het die volgende waardes getoon: d.p.m. droee basis cli fe mg k 107 345 618 696 die waardes blyk normaal te wees. (vii) die rumen, sowel as die abomasum, het lang stukke veselrige plantegroei bevat wat te wyte is aan die afgeslyte gebit van die dier. di e velerlei reeds genoemde gebreke wat die elandbevolking in die bnp oor die jare geopenbaar het en ten spyte van voorkomende maatreels, 500s die verskateng van mineraallekke, om hul voerinname te stimuleer, het die raad van kuratore vir nasionale parke gedurende m2art 1975 tot die oortuiging gebring dat die bestaande habitattoestande ill die bnp nie bevordelik is vir die welvaart van die eland nie. in hierdie verband het oorwegings ook gegeld soos uitgestippel is in van der walt, de graaflen van zyl (976). verwysings de graaff, g., p. t. van der walt en l. j. van zyl, 1976 . populasie-samestelling van die bontebok damaliscus dorcas dorcas in die bontebok nasionale park, gedurendejanuarie 1974. koedoe 19: du plessis, s. f. 1969. the past and present geographical distribution of the perissodactyla and artiodactyla in southern africa. m .sc. thesis, univ. of pretoria. louw , g. n., c. w. p. steenkamp en e. l. steenkamp. 1968. chemiese sames telling van die vernaamste plantspesies in die noorsveld. regn. med. no. 77 pretoria: staatsdrukker. penzhorn, b. l. 1971. a summary of the re-introduction of ungulates into south african national parks (to 31 december 1970) . koedoe 14: 145-159. van der walt, p. t., g. de graaff en l. j. van zyl. 1976. lewensloop van 'n rooihartbeesbevolking alcelaphus buselaphus caama in die bontebok nasionale park. koedoe 19: 181-184. van rensburg, a. p. j. 1975. die geskiedenis van die nasionale bontebokpark, swellendam. koedoe 18: 165-190. 188 page 1 page 2 page 3 page 4 abstract introduction material and methods discussion conclusion acknowledgements references appendix 1 footnotes about the author(s) sam m. ferreira scientific services, south african national parks, skukuza, south africa charlene bissett scientific services, south african national parks, kimberley, south africa carly r. cowell scientific services, south african national parks, cape town, south africa angela gaylard scientific services, south african national parks, knysna, south africa cathy greaver scientific services, south african national parks, skukuza, south africa jessica hayes scientific services, south african national parks, knysna, south africa markus hofmeyr veterinary wildlife services, south african national parks, skukuza, south africa lizette moolman-van der vyver scientific services, south african national parks, knysna, south africa david zimmermann veterinary wildlife services, south african national parks, port elizabeth, south africa citation ferreira, s.m., bissett, c., cowell, c.r., gaylard, a., greaver, c., hayes, j. et al., 2017, ‘the status of rhinoceroses in south african national parks’, koedoe 59(1), a1392. https://doi.org/10.4102/koedoe.v59i1.1392 original research the status of rhinoceroses in south african national parks sam m. ferreira, charlene bissett, carly r. cowell, angela gaylard, cathy greaver, jessica hayes, markus hofmeyr, lizette moolman-van der vyver, david zimmermann received: 19 mar. 2016; accepted: 29 aug. 2017; published: 31 oct. 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract african rhinoceroses (rhinos) experienced a poaching onslaught since 2008 with the epicentre in south africa where most of the world’s rhinos occur. south african national parks, under the management of south african national parks (sanparks), are custodian to 49% of south africa’s white and 31% of the country’s black rhinos. we collated information on rhino population sizes in seven national parks from 2011 to 2015. we include and report on rhino surveys in kruger national park during 2014 and 2015. south-western black rhinos increased over the study period, which allows sanparks to achieve its contribution to south africa’s 2020 target of 260 individuals. south-central black rhinos declined over the study period because of poaching in the kruger national park, making it difficult for sanparks to realise a 9% increase per annum for its expected contribution to the south african target of 2800 individuals. for southern white rhinos, sanparks requires 5% annual growth for its contribution to the south african target of 20 400 individuals. to continue to evaluate the achievement of these targets, sanparks needs annual population estimates relying on total counts, mark-recapture techniques and block-based sample counts to track trends in rhino populations. sanparks’ primary challenge in achieving its contribution to south africa’s rhino conservation targets is associated with curbing poaching in kruger national park. conservation implications: the status and trends of rhino species in sanparks highlight key challenges associated with achieving the national targets of south africa. conservation managers will need to improve the protection of southern white rhino, while the department of environmental affairs need to be made aware of the challenges specifically associated with not achieving targets for south-central black rhino. outcomes for south-western black rhino have already realised and the good conservation efforts should continue. introduction rhinoceroses (rhinos) are under threat worldwide. of the african rhinos, the southern white rhino ceratotherium simum simum is the most abundant (amin et al. 2006a), with its total numbers exceeding that of a minimum viable population (reed et al. 2003). in contrast, the african black rhino sub-species experienced severe declines with the eastern black rhino diceros bicornis michaeli and south-western black rhino diceros bicornis bicornis listed as critically endangered and the south-central black rhino diceros bicornis minor listed as vulnerable (amin et al. 2006a). the northern white rhino ceretatherium simum cottoni is functionally extinct (groves, fernando & robovský 2010) and the western black rhino diceros bicornis longipes (emslie 2011) recently went extinct (travers, waterland & stroud 2011). asian rhinos (greater one-horn rhino rhinoceros unicornis; javan rhino rhinoceros sondaicus and sumatran rhino dicerorhinus sumatrensis) all survive only in small numbers, with the sumatran and javan rhinos each with less than 100 in the wild. there were 3557 greater one-horn rhino by the end of 2015 (emslie et al. 2016). by 2014, south africa was home to approximately 90% of the global population of southern white rhinos, and 36% of the world’s black rhinos, consisting of south-western, south-central and a small extra-limital population of eastern black rhinos (emslie et al. 2016). by then, the south african national parks board (sanparks) was custodian to roughly half of the white rhinos (49%) and 31% of the black rhinos within south africa (emslie et al. 2016). south african national parks are thus home to a substantial proportion of the world’s rhino and play a key role in ensuring their survival. rhino conservation efforts, including anti-poaching, range from maximising species protection to restoring ecosystem processes. for instance, african rhino conservation plans typically seek to maximise population growth (e.g. amin et al. 2006b; knight, balfour & emslie 2013; milliken, emslie & talukdar 2009; mills et al. 2006), induce meta-population dynamics (knight et al. 2013) and maintain genetic integrity (emslie & brooks 1999; karstens et al. 2011). although the most serious threat to rhinos in africa is poaching (emslie 2013), intensified anti-poaching alone may not curb the poaching onslaught (emslie 2013; ferreira, pfab & knight 2014; ferreira & okita-ouma 2012; kagande & musarurwa 2014) and broader socio-economic approaches are required (haas & ferreira 2016). hence some plans incorporate social and economic objectives (knight et al. 2015). several regional, national and international rhino conservation strategies and targets thus underpins sanparks’ rhino management. the black rhino biodiversity management plan (knight et al. 2013) aims for a south african black rhino population growth rate of 5% per annum, with 2800 south-central and 260 south-western black rhinos by the end of 2020. the white rhino biodiversity management plan (knight et al. 2015) also aims for a white rhino population growth rate of 5% per annum, with 20 400 southern white rhinos by the end of 2020 in south africa. in addition to these rhino biodiversity management plans (bmps), south africa’s cabinet adopted an integrated four-pronged approach to curb rhino poaching during 2014 (department of environmental affairs 2014). these are: (1) compulsory interventions to protect rhinos by implementing widespread and intensive anti-poaching programmes as well as creating particular zones of management using technology and intelligence, (2) game-changing interventions, targeted simultaneously at disrupting organised crime and creating opportunities for more equitable benefit-sharing of ecosystem services with all south africans, (3) long-term sustainability interventions to explore the development of a legal and sustainable rhino trade system and (4) biological management interventions that focus on strategic removals from areas of high poaching risk to create rhino strongholds elsewhere. biological management historically served as the core of south africa’s rhino conservation approach and successes (knight et al. 2015). as an identified implementing agency of the rhino bmps, sanparks should contribute 22% (538–676) and 65% (169) of the individuals to the south-central and south-western black rhino targets for 2020, respectively, based on population estimates during 2014. for the southern white rhino, sanparks needs to contribute 49% (9854–10 232 individuals) of the 2020 targets. as rhino conservation targets are set around rhino population sizes and growth rates, it is important to survey populations frequently to detect statistical changes. estimates and trends allow for evaluating progress towards the targets, and assessing the effectiveness of current management interventions. conservation authorities, however, face several challenges when counting and detecting trends in rhinos and other large animal population sizes. these relate to the feasibility of techniques at different population sizes and densities (table 1), and sources of bias or error (e.g. caughley 1974) linked to different methods and habitats. optimal survey designs and intervals are thus context-specific, influencing the precision and accuracy of counts (e.g. ferreira & van aarde 2009). table 1: survey techniques, situation suitability and supporting literature. although authorities recognise the effect of rhino poaching on southern white rhinos in kruger national park (ferreira et al. 2015), there was no previous published attempt to quantify sanparks’ overall progress towards the national rhino conservation targets across the sanparks estate. moreover, sanparks have not yet assessed whether the techniques employed to count rhinos are optimal at current population sizes or densities. this study therefore aims to collate all sanparks’ data on rhino population estimates for the period 2011–2015, across the range of methodologies employed to (1) evaluate the population status of southern white rhino, south-western and south-central black rhino populations in sanparks, (2) assess whether the techniques employed to count rhinos are optimal within park-specific contexts and (3) recommend optimal survey designs for the three sub-species in various national parks. for south-central black and southern white rhinos in kruger national park we also derive estimates of recruitment rates (i.e. proportion of rhinos less than 1 year old reflect rhinos born and that survived the first year after birth) between two surveys and compare these to estimates of poaching rates. the results will allow us to evaluate sanparks’ progress in achieving south africa’s rhino conservation targets and help inform management interventions. material and methods study populations, data collection and collation the study populations include all rhinos in sanparks’ protected areas from 2011 to 2015 (appendix 1). seven national parks contain rhinos in a single management unit. addo elephant national park has rhinos in three separate sections, but for analytical purposes we consider these combined. rhinos experience different features and management histories in the various national parks (appendix 1). we collated data from all rhino census surveys that took place in these parks during our study period. south-western black rhinos occur in four national parks: addo elephant, mountain zebra, karoo and mokala national parks. in the kuzuko section of addo elephant national park, field ranger patrols and game guides record daily sightings of individually recognisable rhinos, through ear notch patterns (greaver, ferreira & slotow 2014; ngene et al. 2011). the nyathi section has low visibility in the mesic thicket vegetation and requires ground or camera-trap observations of individually marked rhinos. however, the presence of several un-notched rhinos that are not individually recognisable, challenges the maintenance of an up-to-date rhino register. regular patrols, however, allow rangers to identify individuals by sex and age occurring in a specific area. rangers monitor rhinos in the colchester-main camp section through individual observation from the ground or using aerial surveys. the populations of black rhino in karoo, mountain zebra and mokala national parks are small enough to maintain a register of all individuals through observations of individually notched rhinos. we collated data on south-western black rhino from sanparks records (sanparks unpublished data1) for addo elephant, mountain zebra, karoo and mokala national park annually from 2011 to 2015. south-central black rhinos occur in two national parks: marakele and kruger national parks. we extracted data for 2015 for marakele from ferreira and greaver (2016). this study made use of a mark-recapture approach to estimate population size. for kruger national park, we collated south-central black rhino data from ferreira, greaver and knight (2011) as well as ferreira et al. (2015). these publications provided estimates with 95% confidence intervals for 2009 and 2013, respectively. to obtain estimates for 2011 and 2012, we used monte-carlo simulation techniques (fishman 1995) to randomly draw from the probability distributions of the estimates collated for 2009 and 2013, and calculated an annual exponential growth rate (caughley 1977). our study also made use of surveys targeting the southern parts of kruger national park during september 2014 and 2015 using block count techniques (ferreira et al. 2011, 2015). the technique focused on surveying 470 and 487 randomly placed blocks 3 km × 3 km in size during 2014 and 2015, respectively. surveyors systematically completed transects comprising a 200 m observation strip on each side of a helicopter within each block with flights 45 m above ground at 65 knots. the survey team comprised a pilot, a data recorder and two observers. using the block count data with an estimator (jolly 1969) during 2014 and 2015 allowed landscape-specific estimates and overall estimates for kruger national park after accounting for the following biases: availability bias came from relationships between vegetation cover and rhino visibility (ferreira et al. 2015), and observer bias came from estimates made during a previous survey (ferreira et al. 2011). detectability bias was minimal as the size of observation strips was narrower than it was in previous studies when detectability was noted as negligible (kruger, reilly & whyte 2008). using the observed age distribution and sex ratios, we could derive recruitment rates for kruger national park between the surveys of 2013 and 2014, as well as 2014 and 2015. this allowed us to compare the number of rhinos recruited through birth and first year survival in the total black rhino population with the number of rhinos poached between the two surveys. we thus collated poaching data in kruger national park for those periods (sanparks unpublished records2). southern white rhinos occur in four national parks – kruger, marakele, mapungubwe and mokala national parks. we collated all southern white rhino observation data for these parks from 2011 to 2015.3 annual and bi-annual helicopter surveys flying at a height of 50 m at a speed of 50–60 knots used 200 m wide transects to systematically cover a park. this provided total counts of southern white rhinos at mokala national park and marakele national parks, respectively. southern white rhinos in kruger national park also occur mostly south of the olifants river. we collated estimates for 2011, 2012 and 2013 (ferreira et al. 2012, 2015). during september 2014 and september 2015 we made use of the same survey and analytical approaches as used for south-central black rhinos in kruger national park. rangers have a full record of every individual in mapungubwe national park based on regular observations. analysis of trends in population estimates and growth rates we calculated sub-species-specific exponential growth rates for each park using an exponential model (caughley 1977). for parks and species that included sample-based estimates with associated 95% confidence intervals in any particular year we made use of monte carlo simulations (fishman 1995) to define exponential population growth rates and their associated 95% confidence intervals. for each year when a sample-based estimate was available, we extracted a value of the population size from the probability distribution defined by the estimate and its 95% confidence interval of that specific population estimate, and calculated exponential growth. we repeated this process 100 000 times and calculated the median, as well as 2.5% and 97.5% percentiles as a definition of the 95% confidence interval. if the confidence intervals of exponential growth rates excluded zero, we concluded that a population is changing. this allowed us to identify the national parks that play an important role in sanparks’ ability to achieve contributions to national rhino objectives. in order to evaluate sanparks’ requirements to meet national targets by 2020, we calculated the annual rhino population growth rate required for each sub-species to reach the stipulated targets, given the population status during 2015 (see the ‘results’ section). these evaluations allowed us to assess whether authorities require revised conservation interventions. determining optimal survey requirements surveying populations living at low densities present similar statistical challenges as surveying populations with few individuals. we generated a two-way matrix that describes the population sizes and densities of all rhino species in sanparks (table 2). for ease of classification we defined population density categories on exponential scales. this allowed us to place a population in the smallest category based on size or density and identify the most suitable techniques given the category (table 3). table 2: summary of rhino populations within south african national parks in categories of population sizes (number of rhinos) and densities (n.km−2). table 3: a summary of techniques associated with evaluating various aspects of african rhino conservation plans. note that suitable techniques can be informed by the total size of a rhino population and/or the density of rhinos in an area of interest. as the survey techniques employed to estimate south-central black and southern white rhinos had estimates of confidence intervals, and thus precision (i.e. the likely spread of estimates given the uncertainties introduced by biases such as availability, observer and detectability biases, see caughley 1974 and thompson 1992), these values allowed us to define optimal monitoring intervals directed at detecting required population growth. we used the required growth rates to achieve targets along with recorded confidence intervals to define optimal survey requirements using power analyses to detect trends (gerrodette 1987). when designing optimal survey requirements, authorities need to trade-off the magnitude of change to detect, survey intervals, the number of surveys required and the total change by the time a trend is detected. this results in authorities deciding how many surveys and intervals are needed to obtain reliable data for detecting trends. results contribution of various subpopulations to rhino population size targets addo elephant national park makes the largest contribution towards south-western black rhino populations, with the combined numbers in kuzuko, nyathi and colchester-main camp sections comprising a minimum of 120 individuals (table 4). over half of the south-central black rhinos within sanparks occur in kruger national park (table 4). between the 2013 and 2014 surveys, the number of south-central black rhinos born that survived the first year in kruger national park was similar to the number of rhinos poached (table 5). more south-central black rhinos were poached between the 2014 and 2015 surveys than what were born and survived the first year. table 4: population estimates of various subspecies of rhinos within south african national parks. we provide 95% confidence intervals for estimates that made use of formal statistical techniques. table 5: a summary of first year recruitment (the number of rhinos born and surviving the first year of life) and poached individuals recorded for south central black rhinos and southern white rhinos in kruger national park between the 2013, 2014 and 2015 surveys. the block count of southern white rhinos in kruger national park produced an estimate of 8821 (95% ci: 8335–9307) south of the olifants river during 2015. this figure increased to a total of 8875 (95% ci: 8365–9337) after including field ranger observations north of the olifants river (table 5). elsewhere, an additional 252 individuals occur in mapungubwe, mokala and marakele national parks combined. more southern white rhinos were born and survived the first year than what were poached between the 2013 and 2014 surveys (table 5). between the 2014 and 2015 surveys, the number of rhinos born and surviving the first year were similar to that poached. this result confirms that, despite significant numbers of southern white rhino poached, kruger national park remains the most important contributor towards the conservation of wild free-ranging individuals of this species within its natural range in south africa. contemporary trends and requirements to meet conservation targets south-western black rhinos increased at a rate of 0.17 (95% ci: 0.10–0.25) per annum from 2011 to 2015, based on annual exponential growth calculated using observed total population estimates. during 2015, sanparks was already contributing 166 individuals of the 169 that sanparks must contribute to the 260 south-central black rhino required by 2020. if sanparks maintain trends noted from 2011 to 2015, 388 (95% ci: 186–810) south-western black rhinos will reside in south african national parks by 2020. the sanparks target is achievable assuming that the current constraints on sanparks south-western black rhinos do not change substantially. sanparks’ south-central black rhinos declined overall at an annual exponential rate of −0.11 (95% ci: −0.03 to −0.18). in kruger national park from 2011 to 2015 there was an exponential decline of −0.15 (95% ci: −0.05 to −0.25), largely as a result of rhino poaching, which overrode the exponential annual rate of 0.48 (95% ci: 0.26–0.69), because of management introductions, at marakele national park. if sanparks is to contribute 538–676 individuals to meet the south african target of 2800 south-central black rhinos by 2020, this will require sanparks’ south-central black rhinos to reach an annual exponential growth rate of 0.09 (95% ci: 0.02–0.18) from 2016 to 2020. southern white rhino populations occurring in the parks outside of kruger national park increased at a rate of 0.14 (95% ci: 0.10–0.19), partly because of introductions from kruger. however, these populations contributed only 3% to the total number of southern white rhinos occurring in south african national parks (table 4). within kruger national park, southern white rhinos may be declining at an exponential rate of −0.05 (95% ci: 0.01 to −0.09) from 2011 to 2015, because of rhino poaching. as a result, sanparks’ southern white rhinos exhibited at best a fluctuating population overall. if sanparks is to contribute the required 9854–10 232 individuals to the 20 400 target of southern white rhinos in south africa, their populations will need to increase at an annual exponential rate of 0.03 (95% ci: 0.01–0.05) from 2016 to 2020. this is unlikely if the current rates of poaching are sustained. optimal survey intervals and requirements survey methods for south-western black rhinos (table 4) do not allow for estimates of confidence intervals and hence do not allow definition of optimal surveys through trade-offs (gerrodette 1987). consequently, south-western black rhino numbers require estimation on an annual basis making use of total counts as before and mark-recapture techniques in future using ear-notching (greaver et al. 2014; ngene et al. 2011) and or genetic marking (brook et al. 2012). in contrast, estimates for sanparks’ south-central black rhinos made it possible to generate coefficients of variance (8% during the study period) with which optimal survey intervals could be calculated. two surveys between 1 and 4 years apart are needed to detect the required annual increase from 2016 to 2020. at this survey interval, there is a trade-off with the total amount of change detected at the time of the second survey’s completion. if declining trends continue as at present, a total reduction of 11% south-central black rhinos will be detected during two surveys 1 year apart. in contrast, a total reduction of 44.2% individuals will be detected with surveys every 4 years (e.g. 2016–2020). optimal survey intervals for south-central black rhinos should also be annual. techniques should focus on block counts and mark-recapture estimates in kruger national park and mark-recapture approaches elsewhere. estimates of southern white rhinos living in south african national parks had a coefficient of variance of 5% at the end of 2015. to detect an annual decline of −0.05 as well as the required annual exponential increase of 0.05 from 2016 to 2020, sanparks can estimate southern white rhinos twice with surveys 1–4 years apart. the total change after 1 year, if potential declining trends realise (i.e. 5% decline in numbers), carries less risk than the total change after 4 years (i.e. 22.6% decline in numbers). southern white rhinos thus also require annual estimates. techniques should focus on block counts in kruger national park and total counts elsewhere. discussion sanparks play a key role in the conservation of three sub-species of rhino within free-ranging conditions in south africa. for south-central black rhinos and southern white rhinos, however, contemporary trends predict that sanparks will not be able to meet contributions to south africa’s rhino population targets if observed annual population growth rates remain the same. encouraging is the significant increases noted for south-western black rhinos living in addo elephant, mountain zebra, karoo and mokala national parks. the observed annual growth exceeds the physiological capability of south-western black rhinos – having their first calf at 5–11 years of age, and at best, a calf every 2 years thereafter (hitchins & anderson 1983). the high observed annual growth can result from small population effects on accuracy and precision of population estimates (gerrodette 1987), accentuating estimated vital rates (akςakaya 2002). high population growth can also result from conservation husbandry seeking to maximise population growth, including skewing sex ratios (holand et al. 2003) and moving rhinos to maximise the social requirements of individuals (reid et al. 2007). over the past 5 years, sanparks moved three south-western black rhinos between four national parks, placed five under custodianship and introduced an additional seven individuals from elsewhere into national parks. the overall growth comes from having a female sex skew resulting from the initial introductions and high female calving rate in the initial years after introduction. the trends noted for the numbers of south-central black rhinos in marakele and kruger national parks significantly contrasts numbers noted for south-western black rhinos. overall, south-central black rhinos occurring in south african national parks declined from 2011 to 2015. even so, the numbers of south-central black rhinos in marakele national park increased significantly over a 5-year period. similar factors influence this trend as for the south-western black rhino population. importantly the introduction of a robust mark-recapture estimate during 2015 (ferreira & greaver 2016) compared to the records on individual rhinos used for the previous year’s may also contribute. even so, the 2015 survey highlights that marakele national park performs well (ferreira & greaver 2016). the trends in numbers of south-central black rhinos in kruger national park dominate in south african national parks. at kruger, south-central black rhino numbers declined significantly over a 5-year period. southern white rhinos occurring in south african national parks followed similar trends to south-central black rhinos where combined numbers, in the three small populations, in mokala, marakele and mapungubwe national parks increased from 2011 to 2015. in kruger national park confidence intervals of estimates from 2011 to 2015 overlapped, but point estimates suggest 1% increase to a potential 9% decline. kruger national park is thus a key park that influence sanparks’ ability to contribute to south africa’s rhino conservation objectives by 2020. many of these patterns across south african national parks result from disruptive effects of poaching on rhinos, particularly in kruger national park (ferreira et al. 2015). from 2011 to 2015, poachers killed one rhino in mapungubwe, five in marakele and a staggering 2936 in kruger national park (sanparks unpublished data4). a key question is: what would have been the rhino losses if sanparks had not implemented an integrated approach through the rhino protection in the smaller parks, anti-poaching in kruger national park and biological management across south african national parks? nonetheless, curbing rhino poaching in kruger national park remains the highest priority for sanparks’ contribution to south africa’s rhino initiatives. the status of rhinos in south african national parks at the end of 2015 provides guidelines for requirements to achieve the 2020 targets. that in itself is challenging. contemporary trends in south-western black rhinos suggest the 2020 contribution will easily be met by sanparks if conditions prevail. sustaining the contemporary trends is unlikely as annual growth rates recorded are higher than what optimal survival and fecundity schedules predicts for closed populations (rachlow & berger 1998). sanparks can enhance population performances through conservation husbandry in addo elephant, mountain zebra, karoo and mokala national parks that seeks to skew sex ratios in favour of adult cows and introducing more young adult cows. removals of existing rhinos to skew sex ratios should focus on sub-adult males. even so, biological management interventions require more land – removed sub-adult males need localities to contribute to creating new populations; however, south-western black rhinos notoriously have density-dependent social constraints that impose on interventions such as translocations (sanparks unpublished data5). several other south african national parks, such as tankwa karoo, namaqua, richtersveld, kalahari, augrabies and cambedoo national parks that may have suitable habitat within the historical distribution of south-western black rhinos (skead 1980) provide additional options, but currently do not have security measures in place or do not have adequate fencing. a further challenge is to source young adult cows. by 2014, namibia was the stronghold for south-western black rhinos (afrsg unpublished data6). the rise in poaching incidents in namibia (afrsg unpublished data7) reduces the likelihood of sourcing south-western black rhinos from there. an encouraging prospect is that sanparks’ conservation husbandry approach from 2011 to 2015 resulted in an inflated increase in south-western black rhino numbers within south african national parks. south-central black rhinos require a 9% increase in numbers per annum to reach the sanparks contribution to south africa’s 2020 target. this exceeds the maximum growth predicted by best survival, the ages at which cows have their first calves (hitchins & anderson 1983) and how often thereafter (ferreira et al. 2011; hitchins & anderson 1983). in addition, sanparks only have two parks where south-central black rhinos occur within their natural distribution. conservation husbandry and expanding marakele national park (ferreira & greaver 2016) should continue, but these practices were not enough to compensate for the losses experienced in kruger national park from 2011 to 2015. the poaching onslaught in kruger national park is the most significant threat to sanparks’ ability to achieve its south-central black rhino target by 2020. the 5% annual increase required for southern white rhino populations is biologically realistic. in fact, the age at which a cow has her first calf, along with birth intervals and survival rates (owen-smith 1988) predict >5% potential annual growth rates. for instance, southern white rhinos realised 9% annual growth in kruger national park prior to the poaching onslaught that began in 2008 (ferreira & okita-ouma 2012). sanparks should continue to implement rhino protection in small parks, anti-poaching in kruger national park and biological management across all of its national parks. implementation has borne some results – in the 18 months before the end of 2015, only one rhino was confirmed poached in the small parks (sanparks unpublished data8). the number of confirmed rhinos poached in kruger national park was one less during 2015 (826) compared to 2014 (827) (sanparks unpublished data11). this contrasts the pattern from 2008 to 2014 when there was a 32% annual increase in the number of rhinos poached (sanparks unpublished data11). sanparks anticipate that the full implementation of zone-, technologyand intelligence-driven anti-poaching in kruger national park will provide improved control of poaching within the park. if complimented with game-changing interventions, including disrupting organised crime (haas & ferreira 2015) and empowering people (lunstrum 2013) outside the park, authorities can curb rhino poaching. several ongoing national and international collaborations, as well as new initiatives could aid in ensuring the long-term persistence of rhinos in kruger national park. the results of our analysis suggest three key directives for conserving south-central black rhinos: (1) the existing conservation targets require revision of the 5% annual increase requirements in the black rhino biodiversity management plan (knight et al. 2013). given the effect of poaching (ferreira et al. 2015), the targeted numbers of sanparks’ contribution will require higher annual growth rates, (2) taking control and curbing the effect of poaching on south-central black rhinos in kruger national park requires urgent interventions including a prioritised security response using interdiction patrol and hot pursuit tactics (haas & ferreira 2017; park et al. 2016) informed by predictions of spatial distribution of rhinos (rachlow, kie & berger 1999) extracted from intensive biological monitoring programmes and (3) we recommend that land for use by south-central black rhinos be increased. the northern part of kruger national park provides an ideal opportunity for expansion. although south-central black rhinos mostly occur in southern kruger at present, rangers noted several cases of south-central black rhinos previously using landscapes in northern kruger (ferreira et al. 2011). expansion into the area can only be considered if authorities adequately address the internal and external factors driving rhino poaching in the park. conclusion the trends of two of the three sub-species of rhinos in national parks are of key concern. poaching, specifically in kruger national park, has disrupted the ability of sanparks to achieve its contribution to south africa’s rhino conservation targets for 2020. achieving targets is feasible for south-western black rhinos, less so for southern white rhinos, and most challenging for south-central black rhinos. this requires sanparks to continue to implement compulsory anti-poaching and innovative biological management interventions in kruger national park. for monitoring, it is critical that sanparks undertakes annual assessments of rhino population status in order to detect trends in population growth. for south-western black rhinos, registration studies that make use of a variety of individual recognition techniques including ear notching and genetic markers obtained through direct, indirect and technology-based observations provide for sanparks needs. south-central black rhinos require mark-recapture and aerial sample-based approaches, while southern white rhinos need aerial total counts and sample-based approaches. ensuring funding to undertake the surveys, as well as capacity to perform the analysis, needs emphasis. the lessons learnt from this study are similarly applicable to surveys of large mammalian herbivores, particularly those undergoing either rapid increases or declines. acknowledgements we are grateful for the assistance provided during the kruger national park surveys. we would like to thank observers ben wigley, scott ronaldson, steven khosa, marius renke, etienne le roux, adolf manganyi, corli wigley, pauli viljoen, isaac sibiya, izak smit and elana mol. we also thank pilots john bassi, grant knight, charles thompson, jaco mol and brad grafton. we appreciate the support from sanparks senior management for census requirements during resource restricted times. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions s.m.f. was responsible for study conceptualisation, analyses, report writing and editing. c.b., a.g. and c.g. were responsible for data collection and contributed to study design, report writing and editing. c.r.c., j.h., m.h., l.m-v.d.v. and d.z. contributed to report writing and editing. references akςakaya, h.r., 2002, ‘estimating the variance of survival rates and fecundities’, animal conservation 5, 333–336. https://doi.org/10.1017/s1367943002004092 amin, r., okita-ouma, b., adcock, r., emslie, r., mulama, m. & pearce-kelly, p., 2006b, ‘an integrated management strategy for the conservation of eastern black rhinoceros, diceros bicornis michaeli, in kenya’, international zoo yearbook 40, 118–129. https://doi.org/10.1111/j.1748-1090.2006.00118.x amin, r., thomas, k., emslie, r.h., foose, t.j. & van strien, n., 2006a, ‘an overview of the conservation status of and threats to rhinoceros species in the wild’, international zoo yearbook 40, 96–117. https://doi.org/10.1111/j.1748-1090.2006.00096.x brook, s.m., van coeverden de groot, p., scott, c., boag, p., long, b., ley, r.e. et al., 2012, ‘integrated and novel survey methods for rhinoceros populations confirm the extinction of rhinoceros sondaicus annamiticus from vietnam’, biological conservation 155, 59–67. https://doi.org/10.1016/j.biocon.2012.06.008 buckland, s.t., anderson, d.r., burnham, k.p., laake, j.l., borchers, d.l. & thomas, l., 2004, advanced distance sampling, oxford university press, new york. caughley, g., 1974, ‘bias in aerial survey’, the journal of wildlife management 38(4), 921–933. https://doi.org/10.2307/3800067 caughley, g., 1977, analysis of vertebrate populations, john wiley & sons, new york. department of environmental affairs, 2014, minister edna molewa leads implementation of integrated strategic management of rhinoceros in south africa, viewed 23 february 2017, from https://www.environment.gov.za/mediarelease/molewa_integratedstrategicmanagement_rhinoceros emslie, r., 2011, diceros bicornis ssp. longipes, iucn 2011, iucn red list of threatened species, version 2011.2. emslie, r.h., 2013, african rhinoceroses – latest trends in rhino numbers and poaching, an update to doc 54-2-annexe 2 from the iucn species survival commission’s (iucn/ssc), african rhino specialist group to the cites secretariat pursuant to resolution conf. 9.14, (rev. cop15) cites cop16 inf doc 51. emslie, r. & brooks, m., 1999, african rhino. status survey and conservation action plan, iucn/ssc african rhino specialist group, iucn, switzerland. ix + 92 pp. emslie, r.h., milliken, t., talukdar, b., ellis, s., adcock, k. & knight, m.h., 2016, african and asian rhinoceroses – status, conservation and trade, cop17 doc. 68, annex 5, viewed 23 february 2017, from https://cites.org/sites/default/files/eng/cop/17/workingdocs/e-cop17-68-a5.pdf ferreira, s.m., botha, j.m. & emmett, m.c., 2012, ‘anthropogenic influences on conservation values of white rhinoceros’, plos one 7(9), e45989. https://doi.org/10.1371/journal.pone.0045989 ferreira, s.m. & funston, p.j., 2010, ‘estimating lion population variables: prey and disease effects in kruger national park, south africa’, wildlife research 37(3), 194–206. https://doi.org/10.1071/wr09030 ferreira, s.m. & greaver, c.c., 2016, ‘re-introduction success of black rhino in marakele national park’, african journal of wildlife research 46(2), 135–138. https://doi.org/10.3957/056.046.0135 ferreira, s.m., greaver, c.c. & knight, m.h., 2011, ‘assessing the population performance of the black rhinoceros in kruger national park’, south african journal of wildlife research 41(2), 192–204. https://doi.org/10.3957/056.041.0206 ferreira, s.m., greaver, c., knight, g.a., knight, m.h., smit, i.p.j. & pienaar, d., 2015, ‘disruption of rhino demography by poachers may lead to population declines in kruger national park, south africa’, plos one 10(6): e0127783. https://doi.org/10.1371/journal.pone.0127783 ferreira, s.m. & okita-ouma, b., 2012, ‘a proposed framework for short-, medium-and long-term responses by range and consumer states to curb poaching for african rhino horn’, pachyderm 51, 52–59. ferreira, s.m., pfab, m. & knight, m., 2014, ‘management strategies to curb rhino poaching: alternative options using a cost-benefit approach’, south african journal of science 110(5–6), 1–8. https://doi.org/10.1590/sajs.2014/20120055 ferreira, s.m. & van aarde, r.j., 2009, ‘aerial survey intensity as a determinant of estimating population sizes and trends for large herbivores’, south african journal of wildlife research 39, 181–191. https://doi.org/10.3957/056.039.0205 fishman, g.s., 1995, monte carlo: concepts, algorithms, and applications, springer, new york. gerrodette, t., 1987, ‘a power analysis for detecting trends’, ecology 68(5), 1364–1372. https://doi.org/10.2307/1939220 greaver, c.c., ferreira, s.m. & slotow, r., 2014, ‘density dependent regulation of the critically endangered black rhinoceros population in ithala game reserve, south africa’, austral ecology, 39, 437–447. https://doi.org/10.1111/aec.12101 groves, c.p., fernando, p. & robovský, j., 2010, ‘the sixth rhino: a taxonomic re-assessment of the critically endangered northern white rhinoceros’, plos one 5(4), e9703. https://doi.org/10.1371/journal.pone.0009703 haas, t. & ferreira, s.m., 2015, ‘federated databases and actionable intelligence: using social network analysis to disrupt transnational wildlife trafficking criminal networks’, security informatics 4, 1. https://doi.org/10.1186/s13388-015-0018-8 haas, t.c. & ferreira, s.m., 2016, ‘combating rhino horn trafficking: the need to disrupt criminal networks’, plos one, 11(11), p.e0167040. https://doi.org/10.1371/journal.pone.0167040 haas, t.c. & ferreira, s.m., 2017, ‘optimal patrol routes: interdicting and pursuing rhino poachers’, police practice and research, https://doi.org/10.1080/15614263.2017.1295243 hitchins, p.m. & anderson, j.l., 1983, ‘reproduction, population characteristics and management of the black rhinoceros diceros bicomis minor in the hluhluwe/comdor/umfolozi game reserve complex’, south african journal of wildlife research 13(3), 78–85. holand, ø., røed, k.h., mysterud, a., kumpula, j., nieminen, m. & smith, m.e., 2003, ‘the effect of sex ratio and male age structure on reindeer calving’, the journal of wildlife management 67, 25–33. https://doi.org/10.2307/3803058 jolly, g.m., 1969, ‘sampling methods for aerial censuses of wildlife populations’ east african agricultural and forestry journal 34, 46–49. kagande, s.m. & musarurwa, l.k., 2014, ‘conserving the african rhinoceros’, biodiversity and conservation 23(2), 497–502. https://doi.org/10.1007/s10531-013-0613-2 karsten, m., van vuuren, b.j., goodman, p. & barnaud, a., 2011, ‘the history and management of black rhino in kwazulu-natal: a population genetic approach to assess the past and guide the future’, animal conservation 14(4), 363–370. https://doi.org/10.1111/j.1469-1795.2011.00443.x knight, m.h., balfour, d. & emslie, r.h., 2013, biodiversity management plan for the black rhinoceros (dicero,s bicornis) in south africa 2011–2020, government gazette south africa, 36096, pp. 5–76. knight, m.h., emslie, r.h., smart, r. & balfour, d., 2015, biodiversity management plan for the white rhinoceros (ceratotherium simum) in south africa 2015-2020, department of environmental affairs, pretoria, south africa. kruger, j.m., reilly, b.k. & whyte, i.j., 2008, ‘application of distance sampling to estimate population densities of large herbivores in kruger national park’, wildlife research 35(4), 371–376. https://doi.org/10.1071/wr07084 lunstrum, e., 2013, ‘green militarization: anti-poaching efforts and the spatial contours of kruger national park’, annals of the association of american geographers, https://doi.org/10.1080/00045608.2014.912545 milliken, t., emslie, r.h. & talukdar, b., 2009, african and asian rhinoceroses–status, conservation and trade, a report from the iucn species survival commission (iucn/ssc) african and asian rhino specialist groups and traffic to the cites secretariat pursuant to resolution conf. vol. 9.14 (rev. cop15), viewed n.d., from https://cites.org/sites/default/files/eng/cop/17/workingdocs/e-cop17-68-a5.pdf mills, a., morkel, p., amiyo, a., runyoro, v., borner, m. & thirgood, s., 2006, ‘managing small populations in practice: black rhino diceros bicornis michaeli in the ngorongoro crater, tanzania’, oryx 40(3), 319–323. https://doi.org/10.1017/s0030605306000901 moss, c.j., 2001, ‘the demography of an african elephant (loxodonta africana) population in amboseli, kenya’, journal of zoology 255(2), 145–156. https://doi.org/10.1017/s0952836901001212 ngene, s., bitok, e., mukeka, j., okita-ouma, b., gakuya, f., omondi, p. et al., 2011, ‘census and ear-notching of black rhinoceros (diceros bicornis michaeli) in tsavo east national park, kenya’, pachyderm, 49, 61–69. olivier, p.i., ferreira, s.m. & van aarde, r.j., 2009, ‘dung survey bias and elephant population estimates in southern mozambique’, african journal of ecology 47(2), 202–213. https://doi.org/10.1111/j.1365-2028.2008.00983.x owen-smith, r.n., 1988, megaherbivores: the influence of very large body size on ecology, cambridge university press, cambridge. park, n., serra, e., snitch, t. & subrahmanian, v.s., 2016, ‘ape: a data-driven, behavioral model based anti-poaching engine’, (to appear in) ieee transactions on computational social systems, 2(2). patton, f., campbell, p. & parfet, e., 2007, ‘establishing a monitoring system for black rhinos in the solio game reserve, central kenya’, pachyderm (43), 87–95. picard, n., chagneau, p., mortier, f. & bar-hen, a., 2009, ‘finding confidence limits on population growth rates: bootstrap and analytic methods’, mathematical biosciences 219, 23–31. https://doi.org/10.1016/j.mbs.2009.02.002 rachlow, j.l. & berger, j., 1998, ‘reproduction and population density: trade-offs for the conservation of rhinos in situ’, animal conservation 1, 101–106. https://doi.org/10.1111/j.1469-1795.1998.tb00017.x rachlow, j.l., kie, j.g. & berger, j., 1999, ‘territoriality and spatial patterns of white rhinoceros in matobo national park, zimbabwe’, african journal of ecology, 37, 295–304. https://doi.org/10.1046/j.1365-2028.1999.00175.x redfern, j.v., viljoen, p.c., kruger, j.m. & getz, w.m., 2002, ‘biases in estimating population size from an aerial census: a case study in the kruger national park, south africa’, south african journal of science 98(9 & 10), 455–461. reed, d.h., o’grady, j.j., brook, b.w., ballou, j.d. & frankham, r., 2003, ‘estimates of minimum viable population sizes for vertebrates and factors influencing those estimates’, biological conservation, 113(1), 23–34. https://doi.org/10.1016/s0006-3207(02)00346-4 reid, c., slotow, r., howison, o. & balfour, d., 2007, ‘habitat changes reduce the carrying capacity of hluhluwe-umfolozi park, south africa, for critically endangered black rhinoceros diceros bicornis’, oryx 41, 247–254. https://doi.org/10.1017/s0030605307001780 seber, g.a.f., 1982, the estimation of animal abundance, backburn press, caldwell. skead, c.j., 1980, historical mammal incidence in the cape province. volume 1, the western and northern cape, the department of nature and environmental conservation of the provincial administration of the cape of good hope, cape town, pp. 277–311. thompson, s.k., 1992, sampling, john wiley & sons, new york. travers, w., waterland, s. & stroud, a., 2011, ‘extinction of western black rhinoceros’, cites afrique 4, 1–3. van aarde, r., whyte, i. & pimm, s., 1999, ‘culling and the dynamics of the kruger national park african elephant population’, animal conservation 2(4), 287–294. https://doi.org/10.1111/j.1469-1795.1999.tb00075.x appendix 1 table 1-a1: features of south african national parks where rhinos occur, along with a brief summary of rhino history in each park. table 1-a1(continues…): features of south african national parks where rhinos occur, along with a brief summary of rhino history in each park. footnotes 1. data available from angela gaylard, angela.gaylard@sanparks.org and charlene bissett charlene.bissett@sanparks.org 2. data available from ken maggs, ken.maggs@sanparks.org 3. data available from cathy greaver, cathy.greaver@sanparks.org and charlene bissett, charlene.bissett@sanparks.org 4. data available from ken maggs, ken.maggs@sanparks.org 5. dave zimmermann, sanparks, dave.zimmermann@sanparks.org 6. african rhino specialist group, mike knight, mike.knight@sanparks.org 7. african rhino specialist group, mike knight, mike.knight@sanparks.org 8. ken maggs, sanparks, ken.maggs@sanparks.org article information authors: marietjie landman1 angela gaylard2 thando mendela2 graham i.h. kerley1 affiliations: 1department of zoology, nelson mandela metropolitan university, south africa 2south african national parks scientific services, garden route regional office, knysna, south africa correspondence to: marietjie landman email: marietjie. landman@nmmu.ac.za postal address: po box 77000, port elizabeth 6031, south africa dates: received: 28 mar. 2014 accepted: 31 aug. 2014 published: 17 nov. 2014 how to cite this article: landman, m., gaylard, a., mendela, t. & kerley, g.i.h., 2014, ‘impact of elephant on two woody trees, boscia oleoides and pappea capensis, in an arid thicket-nama karoo mosaic, greater addo elephant national park’, koedoe 56(1), art. #1231, 3 pages. http://dx.doi.org/10.4102/koedoe.v56i1.1231 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. impact of elephant on two woody trees, boscia oleoides and pappea capensis, in an arid thicket-nama karoo mosaic, greater addo elephant national park in this short communication... open access • abstract • introduction • methods    • study area    • field survey • results • discussion    • recommendations for monitoring • acknowledgments    • competing interests    • authors’ contributions • references abstract top ↑ despite extensive evidence of the influences of elephant on woody trees in savannah habitats, effects on trees in the succulent thickets of the eastern cape are relatively poorly described. our study investigates the role and intensity of elephant impacts on pappea capensis and the relatively rare boscia oleoides in an arid thicket-nama karoo mosaic habitat of the greater addo elephant national park. we show that roughly 19% of the b. oleoides and nearly half of the p. capensis individuals recorded showed signs of elephant impact. elephant often toppled our study trees, and where these individuals were uprooted, mortalities occurred: b. oleoides ∼ 44% of the impacted trees (4 individuals); p. capensis ∼ 22% of the impacted trees (29 individuals). conservation implications: whilst this study is restricted by limited spatial and temporal replication, p. capensis mortalities caused by elephant occurred at a rate exceeding that of other processes. our results provide insight into the severity of the measured changes and the need to reduce the impacts. however, it would be critically important to establish the specific driver of elephant–tree interactions before any management intervention is implemented. introduction top ↑ understanding the distribution of elephant effects amongst elements of biological diversity is important for the conservation management of both elephant and biodiversity (landman et al. in press). in particular, such an understanding provides key insights into the scale of monitoring needed to avoid mismatch at the management scale (lindenmayer & likens 2009). in savannah habitats, the effects of elephant on woody trees are well described; elephant (and fire) are considered to be the primary drivers of tree mortalities, changing habitat composition and structure (kerley et al. 2008; o'connor, goodman & clegg 2007). in the succulent thickets of the eastern cape, however, impacts on trees are relatively poorly described (notable exceptions include effects on euclea undulata and schotia afra [landman et al. in press; stuart-hill 1992]). this is probably because research has focused almost exclusively on coastal thicket types (see the most recent review by kerley & landman 2006), with a relatively poorly developed tree component. for these thicket types, elephant appear to have a disproportionate impact on rare and endemic low-growing succulents and geophytes, potentially causing local extinctions (lombard et al. 2001). however, shifts in the relative importance of tree species (between thicket types) may also mean shifting impacts and increased vulnerability of trees. the kuzuko section of the greater addo elephant national park comprises a mosaic of arid thicket and nama karoo types, with a relatively well-developed woody tree component. following an absence of nearly 100 years (boshoff et al. 2002), elephant were re-introduced into the park during 2005 and have since had substantial impacts on this component of the flora: pappea capensis and the relatively rare boscia oleoides appear to be particularly vulnerable, suffering extensive toppling effects (figure 1). this raises significant concerns regarding their midto long-term longevity, particularly as both species are exceptionally long-lived (some experts estimate up to ∼ 600 years) and form a key structural element of the landscape (lechmere-oertel, kerley & cowling 2005). this article describes the role and intensity of elephant effects on these woody trees in the study area. figure 1: example of a pappea capensis tree toppled by elephant in the kuzuko section, greater addo elephant national park. methods top ↑ study area we conducted the study in the kuzuko section (covering ∼ 110 km2) of the greater addo elephant national park, located north of the zuurberg mountains in the eastern cape, south africa. sixteen elephant were re-introduced into the area in 2005, with the population increasing to twenty-one individuals during 2011. the region is semi-arid, with 200 mm–300 mm rainfall annually, often experiencing extended droughts. more than half of the study area comprises a series of low, undulating hills dominated by a mosaic of arid thicket and nama karoo types, whilst the steeper slopes of the zuurberg mostly give way to arid thicket with a varying abundance of the tree-succulent portulacaria afra (boshoff et al. 2002). field survey during january 2011 we located 17 transects across the arid thicket-nama karoo habitats of the study area. transects were 10 m wide and 1 km long. along each transect we recorded all b. oleoides and p. capensis individuals, distinguishing between individuals that (1) showed no impacts (no impact), (2) were impacted by elephant (elephant; easily differentiated given the severity of the impacts) or (3) were affected by other or unknown factors (other/unknown). for all the affected trees (i.e. 2 and 3 above), we also determined the intensity of the impact according to three categories: (1) broken branches only, (2) toppled, but alive and (3) dead. results top ↑ in total, we recorded 47 b. oleoides (0–10 indiv.transect−1) and 291 p. capensis (6–26 indiv.transect−1) trees (all mature) along transects. roughly 19% of the b. oleoides and nearly half of the p. capensis individuals recorded showed signs of elephant impact (figure 2). whilst impacts in the other/unknown category appeared extensive, exceeding that caused by elephant in the case of b. oleoides, these were generally less severe (i.e. broken branches accounted for most of the impacts; figure 3). importantly, the impacts in this category never resulted in tree mortalities. figure 2: relative impact of elephant on woody trees in the kuzuko section. numbers in brackets indicate the number of individuals of each species recorded. figure 3: intensity of elephant impact on woody trees, distinguishing between individuals with broken branches, toppled trees and those that suffered mortalities. numbers in brackets indicate the number of individuals of each species recorded. elephant often toppled our study trees (figure 3), and where these individual trees were uprooted, mortalities occurred: b. oleoides ∼ 44% of the impacted trees (4 individuals); p. capensis ∼ 22% of the impacted trees (29 individuals). it is not known which of the toppled trees that were alive at the time of sampling later died. because of their size, elephant are the primary drivers of tree toppling in our study area. thus, it is very likely that the remaining toppled and dead individuals recorded (see the other/unknown category in figure 2 and figure 3) reflect background mortalities; in other words, mortalities related to tree demographics or other abiotic factors (e.g. lightning). for p. capensis, mortalities caused by elephant were nearly five times that caused by other processes (i.e. from 6 to 29 individuals). our small b. oleoides sample population precluded any robust conclusions in this regard. discussion top ↑ our results show the extensive impacts of elephant, particularly on p. capensis, in our study area. interestingly, and despite numerous descriptions of the influences of elephant on savannah trees (e.g. kerley et al. 2008; o'connor, goodman & clegg 2007) and the wide geographic distribution of our study species, we provide the first record of elephant effects on these species. thus, our study contributes towards understanding the distribution of elephant effects between plant species in succulent thickets and elsewhere. however, our study is restricted by limited spatial and temporal replication. that is, transects were located in a single habitat, on relatively even terrain and with abundant water provisioning, which probably elevated the observed intensity of the impact (e.g. kerley et al. 2008; landman et al. in press). the steeper slopes of the zuurberg mountains may be inaccessible to elephant, providing a refuge habitat. furthermore, we only measured the impacts once and consequently could not determine rates of toppling or mortality. nevertheless, for p. capensis at least, mortalities caused by elephant occurred at a rate exceeding that of other processes. it is important to recognise that whilst elephant were historically abundant on the coastal lowlands and along the river valleys of the eastern cape, the inland interfluves – such as kuzuko – were only utilised ephemerally (boshoff et al. 2002). this suggests that the woody trees in the region probably escaped intensive utilisation, both spatially and temporally. also, most thicket plants showed limited recruitment from seeds (lechmere-oertel, kerley & cowling 2005), which could mean that the vulnerable, slow-growing and long-lived mature trees may not be readily replaced following these impacts. this provides insight into the severity of the measured changes and the need to reduce the impacts. management interventions that may be implemented to reduce the impacts should be based on robust cause-and-effect relationships between elephant and the impacts (landman, kerley & schoeman 2008). without this, conservation management interventions may fail in their objectives. as an example, the culling of elephant in the kruger national park did not prevent the loss of tall trees (owen-smith et al. 2006). interestingly, tree toppling in our study area was often not associated with other foraging impacts (e.g. extensive branch breaking; m. landman pers. obs., n.d.). thus, reducing elephant numbers per se may be a costly intervention if trees are toppled during the strength training of mature bulls (midgley, balfour & kerley 2005). thus, it would be critically important to establish the specific driver of elephant–tree interactions in kuzuko before any management interventions are implemented. recommendations for monitoring our study only provides a first approximation of the impact of elephant on two key species in the study area. it is important to expand this work to be representative of the habitat and topographic heterogeneity of the area, including information on the age structure and levels of recruitment of trees. furthermore, there currently exists no information on the consequences of the impacts for other components of biodiversity. from figure 1 it is clear that the toppling effects cause disturbance to soil processes and probably ecological functioning. in addition, these effects may (at least initially) facilitate access to forage for co-occurring browsing herbivores. clearly, these knock-on effects require further research. acknowledgments top ↑ south african national parks provided financial support to conduct the study. izak smit provided comments that improved the manuscript. competing interests the authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article. authors’ contributions m.l. (nelson mandela metropolitan university) and g.i.h.k. (nelson mandela metropolitan university) analysed the data and wrote the article. a.g. (garden route regional office) and t.m. (garden route regional office) designed the experiment and conducted the field survey. references top ↑ boshoff, a.f., kerley, g.i.h., cowling, r.m. & wilson, s.l., 2002, ‘the potential distributions, and estimated spatial requirements and population sizes, of the medium to large-sized mammals in the planning domain of the greater addo elephant national park project’, koedoe 45, 85–116. http://dx.doi.org/10.4102/koedoe.v45i2.19 kerley, g.i.h. & landman, m., 2006, ‘the impacts of elephants on biodiversity in the eastern cape subtropical thickets’, south african journal of science 102, 395–402. kerley, g.i.h., landman, m., kruger, l., owen-smith, n., balfour, d., de boer, w.f. et al., 2008, ‘effects of elephant on ecosystems and biodiversity’, in r.j. scholes & k.g. mennell (eds.), elephant management: a scientific assessment for south africa, pp. 146–204, wits university press, johannesburg. landman, m., kerley, g.i.h. & schoeman, d.s., 2008, ‘relevance of elephant herbivory as a threat to important plants in the addo elephant national park, south africa’, journal of zoology 274, 51–58. landman, m., schoeman, d.s., hall-martin, a.j. & kerley, g.i.h., in press, ‘long-term monitoring reveals differing impacts of elephant on elements of a canopy shrub community’, ecological applications. lechmere-oertel, r.g., kerley, g.i.h. & cowling, r.m., 2005, ‘patterns and implications of transformation in semi-arid succulent thicket, south africa’, journal of arid environments 62, 459–474. http://dx.doi.org/10.1016/j.jaridenv.2004.11.016 lindenmayer, d.b. & likens, g.e., 2009, ‘adaptive monitoring: a new paradigm for long-term research and monitoring’, trends in ecology and evolution 24, 482–486. http://dx.doi.org/10.1016/j.tree.2009.03.005 lombard, a.t., johnson, c.f., cowling, r.m. & pressey, r.l., 2001, ‘protecting plants from elephants: botanical reserve scenarios within the addo elephant national park, south africa’, biological conservation 102, 191–203. http://dx.doi.org/10.1016/s0006-3207(01)00056-8 midgley, j.j., balfour, d. & kerley, g.i.h., 2005, ‘why do elephants damage savanna trees?’, south african journal of science 101, 213–215. o'connor, t.g., goodman, p.s. & clegg, b., 2007, ‘a functional hypothesis of the threat of local extirpation of woody plant species by elephant in africa’, biological conservation 136, 329–345. http://dx.doi.org/10.1016/j.biocon.2006.12.014 owen-smith, n., kerley, g.i.h., page, g., slotow, r. & van aarde, r.j., 2006, ‘a scientific perspective on the management of elephants in the kruger national park and elsewhere’, south african journal of science 102, 389–394. stuart-hill, g.c., 1992, ‘effects of elephants and goats on the kaffrarian succulent thicket of the eastern cape, south africa’, journal of applied ecology 29, 699–710. http://dx.doi.org/10.2307/2404479 filelist convert a pdf file! apologies and acknowledgements ladies and gentlemen: we have reached the end of our symposium proceedings and allow me to have the final word. mr e t matenge , director of national parks of the republic of botswana, as well as mr a b bridgens, secretary-general ofsarccus (southern african regional commission for the conservation and utilization of the soil) have tended their apologies for absence. i would like to thank the cabinet (in particular the hon. min. h schoeman) for the announcement of the institution of the 10th national park. this gratitude not only comes from the officials of the national parks board, but also from the people of the entire rsa, be they white , brown or black. i also thank my board for permission to go ahead with the arrangements for this symposium. the person who had to bite the bit was dr g de graaff and with the aid of his colleagues, everything ran smoothly. these efforts are greatly appreciated. furthermore, i would like to thank the international conference organisers. their loudspeakers and translation system never let us down. we had a large group of donours, financially and otherwise. these include the rembrandt organisation, the rennies organisation, coca-cola international, the united tobacco company, mr and mrs dingler, winter minerals (nelspruit) and the total petroleum company. to these organizations and individuals our thanks are due. without becoming sentimental, it is to be emphasized that no symposium can be held without people willing to participate. i would like to thank every speaker, especially dr f vollmar, who came all the way from switzerland to open this symposium. the reaction of the audience was wonderful. in particular i would also like to thank his royal highness, prince m. dlamini (swaziland) as well as the many leaders representing the developing territories, for their esteemed presence. my thanks are also due to the south african broadcasting corporation. we hope that whatever was summarized in such an efficient way by dr martiny, that it is clear that this symposium emphasized the importance of education and carried a message directed at the youth of southern africa. r. knobel symposium chairman 260 page 1 article information authors: armin h. seydack1 cornelia c. grant2 izak p. smit2 wessel j. vermeulen1 johan baard1 nick zambatis2 affiliations: 1scientific services, garden route national park, south african national parks, south africa 2scientific services, kruger national park, south african national parks, south africa correspondence to: armin seydack postal address: po box 3542, knysna 6570, south africa dates: received: 08 mar. 2011 accepted: 05 oct. 2011 published: 08 feb. 2012 how to cite this article: seydack, a.h., grant, c.c., smit, i.p., vermeulen, w.j., baard, j. & zambatis, n., 2012, ‘climate and vegetation in a semi-arid savanna: development of a climate–vegetation response model linking plant metabolic performance to climate and the effects on forage availability for large herbivores’, koedoe 54(1), art. #1046, 12 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1046 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. climate and vegetation in a semi-arid savanna: development of a climate–vegetation response model linking plant metabolic performance to climate and the effects on forage availability for large herbivores in this original research... open access • abstract • introduction    • study area • materials and methods    • development of an explanatory framework    • definition of indicators of forage quantity and quality • results    • climate–vegetation response model       • indicators of forage availability       • herbivore species guilds of key nutritional requirements       • plant–climate response processes • discussion    • model postulates and predictions       • key forage resources       • herbivore species guilds of nutritional requirements       • spatial distribution of key forage resources       • climate-driven temporal trends in forage availability       • spatiotemporal interaction in forage availability • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ a framework to establish the expected effects of climate on forage quantity and quality in a local savanna system was developed to interpret large herbivore population performance patterns in the kruger national park. we developed a climate–vegetation response model based on interpretation and synthesis of existing knowledge (literature review) and supported by investigation and analyses of local patterns of climate effects on forage plant performance and chemical composition.developing the climate–vegetation response model involved three main components, namely (1) defining indicators of forage availability to herbivores (nitrogen productivity, nitrogen quality, carbon-nutrient quality), (2) identifying herbivore species guilds of similar nutritional requirements with respect to these indicators [bulk feeders with tolerance to fibrous herbage (buffalo, waterbuck), bulk feeders with preference for high nitrogen quality forage (short grass preference grazers: blue wildebeest and zebra) and selective feeders where dietary items of relatively high carbon-nutrient quality represented key forage resources (selective grazers: sable antelope, roan antelope, tsessebe, eland)] and (3) developing a process model where the expected effects of plant metabolic responses to climate on key forage resources were made explicit. according to the climate–vegetation response model both shorter-term transient temperature acclimation pulses and longer-term shifts in plant metabolic functionality settings were predicted to have occurred in response to temperature trends over the past century. these temperature acclimation responses were expected to have resulted in transient pulses of increased forage availability (increased nitrogenand carbon-nutrient quality), as well as the progressive long-term decline of the carbon-nutrient quality of forage. conservation implications: the climate–vegetation response model represents a research framework for further studies contributing towards the enhanced understanding of landscape-scale functioning of savanna systems with reference to the interplay between climate, vegetation and herbivore population dynamics. gains in such understanding can support sound conservation management. introduction top ↑ herbivore population trends in the kruger national park (knp) over the past century were traditionally explained with reference to decimation by hunting and epizootics (joubert 2007a; pienaar 1963), effects of rainfall (dunham, robertson & grant 2004; owen-smith & ogutu 2003; whyte & joubert 1988) and predation (harrington et al. 1999; joubert 2007a, 2007b; owen-smith, mason & ogutu 2005; owen-smith & mills 2006; owen-smith & mills 2008). however, the long-term and spatially widespread nature of population trends suggested that over-arching landscape-scale influences may also be involved to an as yet unknown extent. a greater role of climate effects as a long-term landscape-scale factor was therefore implicated. accordingly, we initiated a study to explore the extent to which climate–vegetation responses could plausibly explain diverse and divergent spatiotemporal patterns in population performance of eight large herbivore species, namely african buffalo (syncerus caffer), blue wildebeest (connochaetes taurinus), plains zebra (equus burchelli), waterbuck (kobus ellipsiprymnus), sable antelope (hippotragus niger), roan antelope (hippotragus equinus), tsessebe (damaliscus lunatus lunatus) and eland (taurotragus oryx). to interpret large herbivore population performance patterns over space and time in relation to climate it was necessary to develop a framework in which the expected effects of climate on forage quantity and quality were established. towards this purpose we developed a climate–vegetation response model. this model was developed through interpretation and synthesis of existing knowledge (literature review) and supported by analyses of local patterns of climate effects on forage plant performance and chemical composition. forage availability to herbivores was taken to be a function of plant productivity and the nutritional quality of the resultant plant material (fryxell 1991; holdo, holt & fryxell 2009), based on concentrations of protein (nitrogen), non-structural carbohydrates (tnc) and minerals in plant tissues. key resources for herbivores thus relate to nitrogen productivity, nitrogen quality and carbon-nutrient quality. a central notion of the climate–vegetation response model is that levels of these three forage resource indicators are affected differentially by plant metabolic processes in response to the nature of edaphoclimatic resources (water, temperature and nutrients) available to plants. to the extent that different herbivore species have divergent nutritional requirements in respect of these key forage resources, we expected that their population performances would be affected differentially by conditions of the edaphoclimatic environment (spatial scale) and climate (temporal scale). the climate–vegetation response model developed here accordingly consists of three main components: • definition of indicators of forage availability to herbivores (nitrogen productivity, nitrogen quality and carbon-nutrient quality) • identification of herbivore species guilds of similar nutritional requirements in respect of these indicators • development of a climate–plant metabolic response process model where the expected effects of plant metabolic responses to climate on key forage resources are made explicit. with the climate–vegetation response model as explanatory framework, large herbivore population performance patterns in both space and time in the knp were investigated and interpreted (seydack et al. 2012). in a broader sense, however, our climate–vegetation response model is intended to represent a research framework for further studies contributing towards the enhanced understanding of landscape-scale functioning of savanna systems in respect of the interplay between climate, vegetation and herbivore population dynamics. study area the knp is situated in north-eastern south africa and represents a large (approximately 2 000 000 ha) semi-arid savanna system. the area lies between the drakensberg escarpment in the west and the mozambique coastal plain in the east. covering an altitudinal range of between 200 m a.s.l. and 700 m a.s.l., the knp falls within two disparate climate zones as defined by the south african weather service (venter, scholes & eckhardt 2003). the area north of the olifants river is in the northern arid bushveld zone, receiving 300 mm – 500 mm rain per year, whilst the southern part of the park falls into the lowveld bushveld zone, with an average annual rainfall of 500 mm – 700 mm per year. years of pronounced below-average rainfall were 1963–1964, 1973, 1983–1984, 1987, 1991–1992 and 1998 (figure 1). growing season duration (v. schulze 1997) is on average about 1.5 months shorter in the north than in the south (96 days compared with 142 days, respectively). figure 1: mean annual rainfall in the kruger national park (1954–2006). temperature records from stations within or near the knp followed trends congruent with those recorded globally (figures 2 and 3). three phases of temperature increases were discernable. the first reached peak values between 1937 and 1947, followed by a second phase of increasing temperature towards the period 1958–1960 and a third, still ongoing, warming phase which commenced between 1975 and 1978. locally differentiated temperature trends are shown in figure 3. regional temperature lapse rates for monthly means of daily maximum and minimum temperatures are –0.777 °c×100 m-1 and –0.465 °c×100 m-1, respectively (schulze 1995). as indicated by higher summer-to-winter heat unit differentials (determined according to schulze 1997), northern areas of the park experience more pronounced temperature seasonality. figure 2: southern hemisphere surface air temperature anomalies between 1850 and 2009. figure 3: mean annual temperatures at musina (525 m a.s.l), approximately 100 km north-west of the kruger national park (south african weather service). the park is longitudinally divided into granitic substrates forming relatively nutrient-poor sandy soils in the west and more nutrient-rich, basalt-derived clay soils in the east (venter et al. 2003). the savanna vegetation on nutrient-poor substrates tends to be dominated by trees of combretaceae (e.g. combretum and terminalia species) and caesalpineaceae, whereas those on more nutrient-rich clay soils are dominated by trees of mimosaceae (especially acacia species) (gertenbach 1983; venter et al. 2003). the vegetation of the northern knp is characteristically dominated by mopane (colophospermum mopane) and this species is well represented in broad-leaved bushveld vegetation types on granites and broad-leaved shrubveld associated with basalts (gertenbach 1983; venter et al. 2003). the herbaceous layer of the knp is dominated by c4 grass species (kennedy, biggs & zambatis 2003) and the more nutrient-rich savanna vegetation types on clay soils carry dense stands of nutritious, high-bulk grasses (venter et al. 2003). materials and methods top ↑ development of an explanatory framework exploration of spatiotemporal patterns in population performance of eight large herbivore species revealed three population performance response groups (i.e. groups of large herbivore species that shared broad-scale habitat preferences and similar population performance trends over time) and group-specific correlations with climate variables (seydack et al. 2012). we developed an explanatory framework – the climate–vegetation response model – which permits the interpretation of these correlative patterns in relation to climate effects. as part of the climate–vegetation response model we (1) defined indicators of forage availability (quantity and quality), (2) established forage selection requirements characteristic of and unique to the three herbivore population performance response groups (nutritional guilds), and (3) developed a plant-climate response process submodel. the resulting climate–vegetation response model defined the expected effects of climate on indices of forage availability, thereby providing an explanatory framework for the interpretation of spatiotemporal patterns in herbivore population performance in relation to climate. the model was developed largely based on the interpretation and synthesis of relevant literature. some results of local research contributed towards the definition of key forage resource indices. definition of indicators of forage quantity and quality leaf and stem material of the grass species panicum maximum, panicum coloratum and themeda triandra were collected in and beyond the roan enclosure (nwashitshumbe, northern knp) during september 2007, november 2007, august 2008 and february 2009. the apical 20 cm of grass stems were collected as samples of stem material. the samples originated from a study to explore the effects of fire frequency on forage quality, where sampling was undertaken in four treatment blocks (three inside and one outside of the enclosure) defined by the time since the last burn. each treatment block contained three sampling localities (replications) to represent a total of 12 samples per sampling event. the collected grass material was dried at 55 °c overnight and mill ground through a 1-mm sieve to form homogeneous powdered samples. the concentrations of ash minerals and tnc were determined, as well as in vitro organic matter digestibility (omd) of the powdered sample material. these analyses were performed by the analytical services department of the agricultural research council (arc) in irene. ashing was performed in a furnace at 600 °c. tnc were analysed as reducing sugars after complete enzymatic hydrolysis to monosaccharides. the method entailed gelatinisation of all starch in the sample by autoclaving, followed by enzymatic hydrolysis of the starch to glucose and determination of glucose content by spectrometric measurement. duplicate results were within 3% of the mean for each pair. in vitro omd was determined following an adaptation of the two-phase technique described by tilley and terry (1963), with some modification introduced by engels and van der merwe (1967). it involved a 24-hour-long fermentation by rumen micro-organisms in a buffer solution followed by a 48-hour-long pepsin digestion after acidification. the reduction in the organic matter content was ascribed to digestion of the sample. for the determination of d13c and d15n, samples were combusted in an automated thermo1112 elemental analyser (carlo erba, milan). the resultant co2 and n2 gases were then introduced to a thermo delta stable light isotope xp mass spectrometer via a continuous flow-through inlet system (thermo conflo iii). 13c/12c and 15n/14n ratios were expressed in the delta (d) notation in parts per thousand (o/oo) relative to international standards (pee dee belemnite standard for carbon; n2 air for nitrogen). standard deviations of repeated measurements were less than 0.1o/oo for d13c (stable light isotope unit, department of archaeology, university of cape town). grass sample collections during november 2007 and february 2009 represented spring and summer sampling events, respectively, and plant growth under conditions of concurrent high levels of temperature and water availability (tw conditions). dry-season grass and stem material collected during september 2007 had been subject to summer greening in november–december 2006 (tw metabolic cueing at sub-mature stages; mean monthly maximum and minimum temperatures at 33.9 °c and 21.1 °c, respectively) and water-stressed summer growth conditions (tw growth realisation at mature stages). dry-season material collected in august 2008 represented conditions following spring greening in september–october 2007 (tw metabolic cueing at sub-mature stages; mean monthly maximum and minimum temperatures at 30.6 °c and 17.7 °c, respectively) and summer growth subject to high rainfall conditions (tw growth realisation at mature stages). the main objective of this part of the study was to determine how these varying growth conditions affected the chemical composition and digestibility of the grass tissues and the implications for key indices of forage quality defined in the context of this study. results top ↑ climate–vegetation response model indicators of forage availability nitrogen productivity: nitrogen productivity is defined as the quantitative availability of forage items for herbivores of a given nitrogen quality (plant protein content) as a result of biomass production. grass productivity is known to be strongly promoted by water availability [positively correlated with rainfall (zambatis 2004)] and temperature, especially in the case of tropical grass species (long 1999; luo 2007; wan et al. 2005), which have high temperature optima for growth (dwyer et al. 2007; ’t mannetje 1982; tudsri, matsuoka & kobashi 2002; wilson & ford 1971). warming-induced increases in biomass production of c4 grasses have been demonstrated under experimental conditions (an et al. 2005; bijoor et al. 2008). accordingly, nitrogen productivity of c4 grasses and subtropical woody vegetation is expected to increase over time as ambient temperature increases (figure 2). however, at progressively higher nitrogen productivity, an inverse relationship to nitrogen quality due to nitrogen dilution is generally encountered (an et al. 2005; fryxell 1991; fryxell et al. 2005; mutanga et al. 2004; wilson 1982).nitrogen quality: high nitrogen quality of forage implies high nitrogen concentrations of plant tissues; that is, plant protein content not unduly diluted by structural or non-structural carbon and of relatively high digestibility. an index of nitrogen quality is defined here as the product of nitrogen concentration and omd of the plant tissues involved. wet-season nitrogen quality tended to be higher early in the wet season (spring) compared with later in the wet season (summer) (figure 4c). nitrogen content in summer leaf material (sul) was diluted by structural carbon relative to that of spring (spl) leaf material (analysis of variance for carbon content, followed by a post hoc bonferroni test showed sul > spl; p < 0.00001) and carbon-nutrient levels (figure 4d: sul > spl). this is congruent with nitrogen dilution through growth under conditions of relatively high temperature and water availability (tw conditions), as typically experienced by tropical grasses (wilson & ford 1971). figure 4: monthly rainfall at vlakteplaas (northern basalt plains, kruger national park) for two consecutive years (a) 2007/2008 and (b) 2006/2007, and forage quality indices of selected grass tissues (c) index of nitrogen forage quality and (d) index of carbon-nutrient forage quality. dry-season leaf (wl) and stem (ws) material were sampled to represent two sets of conditions (figure 4): summer greening and water-stressed summer growth (wl-su/ws-su) and spring greening and summer growth with high rainfall (wl-sp/ws-sp), respectively. nitrogen quality of dry-season wl and ws material (figure 4c) was enhanced following water-stressed summer growth (wl-su/ws-su: figure 4b; tw conditions indicated by low d15n values) compared with such material following summer growth in higher rainfall conditions (wl-sp/ws-sp: figure 4a; tw conditions indicated by high d15n values; n-quality: leaf material: f = 113.36, n = 72, p < 0.000001; stem material: f = 50.77, n= 72, p < 0.000001). omd of dry-season grass tissue (both leaf and stem material) following water-stressed summer growth (tw conditions; figure 4b) was significantly higher than of dry-season material following summer growth under tw conditions (figure 4a; omd: anova: bonferroni post hoc test: wl-su > wl-sp; p < 0.0001; ws-su > ws-sp:p < 0.00001). omd of dry-season grass leaf material was positively related to nitrogen content and negatively correlated with d15n values (multiple regression of omd on n and d15n: ß of n = +0.758; ß of d15n = –0.348, r2 = 0.42, n = 72). according to the results of codron et al. (2005) seasonal shifts in plant d15n corresponded positively with seasonal summer rainfall, supporting our interpretation that d15n values index summer growth activity under tw conditions. these relationships are consistent with general findings that metabolic activity under water stress promotes digestibility (wilson 1982) and that high growth activity under tw conditions results in reduced forage quality. metabolic activity under conditions of high temperature and water availability (tw conditions) is expected to negatively affect dry matter digestibility as it promotes more rapid growth and stem development of, particularly, tropical grasses (’t mannetje 1975; wilson 1982; wilson, taylor & dolby 1976). high growth temperatures also hasten the maturation of individual tissues and the digestibility of plant tissue is usually higher when grown at low temperatures, such as in spring or winter (’t mannetje 1984; wilson 1982).carbon-nutrient quality: carbon-nutrient quality of forage items is defined here as the product of tnc and ash mineral concentrations (figure 4d). dry-season tnc content of grass leaf material following spring greening was significantly higher than that of leaf material following summer greening (dry-season tnc content: wl-sp > wl-su; f = 25.02, p < 0.00001). a similar pattern was found for ash (mineral) contents (anova, followed by post hoc bonferroni test: wl-sp > wl-su and ws-sp > ws-su, p < 0.00001). these results are consistent with a metabolic response mode of low-temperature growth during spring (tw metabolic cueing at sub-mature stages) resulting in storage allocation of tnc (estiarte & peñuelas 1999; herms & mattson 1992; wilson 1975) and of minerals (balasko & smith 1971). in accordance with these findings, high carbon-nutrient quality was typically encountered in dry-season leaf material (figure 4d) and was higher in dry-season leaf and stem tissue in grass cohorts that had been subject to spring greening (figure 4d; anova followed by a post hoc bonferroni test: wl-sp > wl-su, p < 0.00001; ws-sp > ws-su, p < 0.00001). carbon-nutrient quality forage items are thus characterised by relatively high concentrations of tnc and ash minerals resulting from substantial metabolic allocation to storage under circumstances promoted by constrained growth (particularly owing to low temperatures, but also moisture stress: tw conditions) but sustained photosynthesis. carbon-nutrient quality of dry-season plant leaf and stem material of t. triandra (andropogoneae) was generally higher than for the two species of paniceae (figure 4d; anova and post hoc bonferroni tests: p < 0.01). this reflected the predisposition of andropogonoid species, geared to cope with more pronounced seasonality (osborne 2008), towards greater metabolic scope for storage investment. the typical accumulation of tannin-like substances in the foliage of andropogoneae indicates a carbon excess metabolism with pronounced allocation to storage (baas 1989; herms & mattson 1992). such physiological characteristics are considered adaptive to ensure sustained metabolic performance under seasonally variable conditions of resource availabilities (temperature and water: tw/tw/tw conditions). all species of andropogoneae function according to the nadp-me biochemical subtype and these are implicated to operate at lower temperature optima than species of the other subtypes, including, inter alia, species of paniceae (ehleringer, cerling & helliker 1997). grass species of this biochemical subtype, and particularly of andropogoneae, also exhibited the smallest responses to changes in environmental conditions (buchmann et al. 1996). these findings are suggestive of the capacity of andropogoneae species for sustained metabolic performance at variable and lower resource levels (dry-season growth), underpinning their metabolic scope for storage of metabolites (tnc and minerals for enhanced carbon-nutrient quality). herbivore species guilds of key nutritional requirements in this section we present evidence suggesting that herbivore species within particular population performance response groups share similar broad-scale forage selection requirements. buffalo and waterbuck are considered roughage feeders able to digest fibrous food relatively efficiently (hofmann 1973, 1989). despite the relatively unselective bulk grazing habit of buffalo, they adjust their feeding patterns in relation to seasonally changing forage quantity and quality (macandza, owen-smith & cross 2004). however, for both these species food quantity seems to override exacting quality requirements. this is in contrast to blue wildebeest and zebra, which have clear preferences for short-grass areas on nutrient-rich soils (grange & duncan 2006; traill 2004). both species select grass patches of high nitrogen quality for feeding, although zebra, as a hindgut fermenter, can tolerate more fibrous and longer-grass feeding conditions (bodenstein, meissner & van hoven 2000; smuts 1972).apart from species-specific, often seasonally changing, feeding preferences (dunham et al. 2004; heitkönig & owen-smith 1998; joubert & bronkhorst 1977; magome et al. 2008; watson & owen-smith 2000), all four selectively feeding species (i.e. tsessebe, eland, sable and roan antelope) are able to adapt to a wide range of grass heights and often include medium to tall grasses in their diets (joubert 1976; skinner & chimimba 2005; traill 2004). according to a study by parrini (2006) sable antelope do not avoid stemmy grass tufts and include tall stemmy species generally avoided by short-grass grazers. sable antelope were found to be particularly tolerant of stems in habitats with intermediate to high nutrient levels. heitkönig and owen-smith (1998) found a similar pattern in roan antelope, which tolerated stems when leaf quality was high. the preferred feeding sites of tsessebe typically have soils with a high clay content (dörgeloh 2006). topi (damaliscus lunatus jimela), with foraging habits believed to be similar to those of tsessebe, selected for the tallest plants and species with relatively long leaves when feeding on grass with more than 20% green leaves (duncan 1975 op. cit. parrini 2006). these patterns of ingesting stem material together with leaf matter and the relatively high importance of nutrient contents of these are commensurate with the maintenance of relatively high dietary tnc and mineral nutrient levels resulting in carbon-nutrient ratios as required for sustained carbon-nutrient quality. numerous studies have revealed the prominence of andropogonoid grass species in the preferred habitat and forage species assemblages of sable antelope (le roux 2010; magome 1991; parrini 2006), roan antelope (joubert 1976; van lavieren & esser 1979), tsessebe (gureja & owen-smith 2002) and eland (watson & owen-smith 2000). as indicated in the previous section, andropogonoid grass species would tend towards a sustained metabolic performance type metabolism and thus have a propensity towards carbon and nutrient storage, particularly when subject to seasonally low temperature conditions. under such metabolically favourable conditions, these species are expected to have high carbon-nutrient quality; that is, a combination of favourable levels of tnc, nitrogen and minerals. all these features of forage selectivity, including tolerance of stemminess, prominence of andropogonoid grass species, adaptability to a wide range of grass heights and, in the case of browsing, feeding on freshly fallen tnc-rich leaves [see skinner & chimimba (2005) for reference to eland] indicate that these species can be classed as selective feeders of plant parts of relatively high carbon-nutrient quality. in summary, the eight species studied here are grouped into three guilds of nutritional requirements: bulk feeders with tolerance to fibrous herbage (buffalo and waterbuck), bulk feeders with preference for high nitrogen quality (short-grass grazers: blue wildebeest and zebra) and selective feeders for which dietary items of relatively high carbon-nutrient quality are of particular importance (selective grazers: sable antelope, roan antelope, tsessebe and eland). plant–climate response processes plant metabolic performance trade-off relationships: owing to resource limitations plants are subject to trade-offs between growth, maintenance, storage, reproduction and defence (herms & mattson 1992). consequently, there are, inter alia, strong inverse relationships between the functional allocation to growth and non-growth processes, notably carbon storage (herms & mattson 1992). accordingly, there is an allocation trade-off between quantity of plant tissue produced through growth (structural plant content) and quality, represented by the proportion of non-structural plant content (metabolically active components and storage products). thus, abundance of resources such as water, nutrients and temperature enhance productivity, but increasing productivity then progressively reduces plant tissue quality for herbivores through nutrient dilution (an et al. 2005; dwyer et al. 2007; ’t mannetje 1982, 1984; tudsri et al. 2002; van soest, mertens & deinum 1978; wilson & ford 1971; wilson 1982), reduction in digestibility (’t mannetje 1975, 1984; van soest et al. 1978; wilson & ford 1973; wilson & ‘t mannetje 1978) and reduced allocation to storage (low tnc content) (estiarte & peñuelas 1999; herms & mattson 1992; van soest et al. 1978).the climate–vegetation response model was developed to integrate existing knowledge and to form part of an explanatory template for spatiotemporal patterns of climate–vegetation response processes controlling forage quantity and quality. at the core of this model is the metabolic performance trade-off model, from which predictions can be made regarding divergent plant responses to site and climate-linked resource constraints and their impact on parameters of forage availability to herbivores. the principal trade-off in the context of this model is between maximised peak performance, closely tracking and responsive to conditions of resource surplus (rmp; resource responsive metabolic performance mode), and sustained performance when subject to conditions of varying resource levels or deficits (smp; sustained metabolic performance mode). the capacity for maximum metabolic performance under conditions of resource surplus is at the cost of the capacity for sustained performance under conditions of resource deficits or fluctuations. in line with this reasoning, we expect a propensity towards the rmp mode under conditions of high and less fluctuating levels of temperature and water availability (tw conditions: combination of relatively high and sustained levels of water and temperature during the growing season), whereas under conditions of fluctuating resource levels and pronounced deficits (regarding temperature and water; tw or tw conditions), the smp mode would be expected to predominate. high peak-level performance characteristics of the rmp mode require high turnover metabolism (capacity for high instantaneous rates of photosynthesis; amax). this is generally associated with high foliar nitrogen levels (lambers, chapin & pons 1998) resulting in increased nitrogen/carbon ratios of plant tissues. in contrast, sustained performance capacity of the smp mode requires resource use efficiency and is associated with the propensity towards somatic accumulation [size, storage, longevity: relatively high carbon/nitrogen ratios of plant tissues; herms and mattson (1992)]. furthermore, stress-tolerant plants (smp mode) tend to allocate an especially high proportion of available energy to maintenance respiration required to tolerate adverse environmental conditions (taylor 1989). plants in a relative smp mode are accordingly expected to be even more sensitive to resource conditions inducing elevated maintenance respiration, such as increased mean mimimum temperature (reflecting nocturnal temperature conditions) (tmin). resource constraints on growth are generally more intense than their inhibitory effect on photosynthesis. as a consequence of carbon acquisition exceeding growth requirements, the products of photosynthesis accumulate, primarily as carbohydrate storage products (baas 1989; herms & mattson 1992; lambers & poorter 1992). when tracking resource surplus conditions favourable for both photosynthesis and growth in rmp-mode plants, the resource-level response relationships for growth and photosynthesis tend to be similar (wilson & ford 1973). in smp-mode plants, stress tolerance capacity is associated with photosynthesis being relatively less sensitive to resource levels than growth. accordingly, there is greater scope for source (photosynthesis) to sink (growth) imbalance in smp-mode plants, favouring allocation to secondary metabolism and storage as carbon assimilation levels exceed those which are required for growth. plants in relative smp mode are thus comparatively stress tolerant in order to cope with fluctuations in resource levels. as such their metabolism is capable of functioning at relatively low temperature and water availability levels/optima (broad metabolic amplitude; smp-tw functional efficiency). such metabolic capacity is, however, associated with relatively high storage-to-growth allocation priority and comparatively high maintenance respiration costs (i.e. temperature sensitivity). in contrast, plants in relative rmp mode are geared to the maximum use of resource surplus conditions, functioning at relatively high temperature and water resource level optima (narrow metabolic amplitude) and high growth-to-storage priority (figure 5). figure 5: sustained metabolic plant performance (smp) to resource-responsive metabolic performance (rmp) mode adjustments to resource conditions, also representing the smp to rmp temperature acclimation trajectory endpoints (climate–vegetation response process model). plant metabolic performance mode settings: the relative expression of smp to rmp mode functionality/acclimation is a function of the combined availability levels of temperature and water (tw resource conditions). the higher the concurrent and uninterrupted levels of temperature and water available to plants, the more pronounced the intraspecific metabolic adjustment towards higher rmp mode functionality settings. the relative expression of smp or rmp depends on the spatiotemporal nature of resource availability for plants (water and temperature resource levels and variability or seasonality), shaping forage quantity and quality as a consequence of the relative prevalence of smp-tw/tw to rmp-tw functionality (figure 5). resource conditions in the northern knp are characterised by greater moisture stress levels and higher degrees of resource level fluctuations regarding both moisture availability and temperature compared to the southern knp (increasing north to south rainfall gradient: knp-north = 483 mm/year; knp-central = 535 mm/year; knp-south = 632 mm/year). growth season duration (v. schulze 1997) is, on average, about 1.5 months shorter in the northern knp. according to the metabolic performance trade-off model, we therefore expect the smp mode to predominate (intraspecifically) in northern knp areas and the rmp mode to be more prevalent in southern areas of the park. conditions prevailing in the central knp are expected to promote an intermediate position in this regard. conditions for plant functionality are also expected to change along an altitudinal gradient [temperature lapse rates for monthly means of daily maximum and minimum temperatures are –0.777 °c×100 m–1 and –0.465 °c×100 m–1, respectively (schulze 1995)]. a pattern of lower temperatures and increased rainfall at higher altitudes represents tw conditions, with tw conditions prevailing at lower altitudes. combining the north–south water availability and altitudinal temperature gradients, plant functionality settings were identified and the associated expected implications for forage quality indicated in accordance with the metabolic performance trade-off model (table 1). table 1: plant functionality settings according to prevailing environmental conditions in various parts of the kruger national park and their expected consequences for forage quality (climate–vegetation response model). as a general notion, nitrogen quality is usually considered to be higher in plants on nutrient-rich geological substrates, resulting in higher palatability for herbivores (skidmore et al. 2010; venter et al. 2003). this pattern may, however, be modified at various spatial scales by the availability of water (mutanga et al. 2004) and factors resulting in nitrogen dilution (accentuated smp or rmp functional expression as in northern and southern knp, respectively). at a spatial scale we thus generally expect the prevalence of relatively high nitrogen quality forage on nutrient-rich geological substrates in the central knp (table 1). temperature acclimation and forage quality: ambient temperature has been increasing over the past century (figure 2). in particular, tmin, which represents nocturnal temperature conditions, has increased more steeply than the mean periodic maximum temperature (tmax), which represents diurnal temperature conditions (houghton et al. 1996). increasing tmin is expected to stimulate nocturnal maintenance respiration (wan et al. 2009), promoting temperature acclimation in order to restore the original steady-state respiration-to-photosynthesis ratio (dewar, medlyn & mcmurtrie 1999). as temperature increases, leaf metabolic pools adjust until the original steady-state respiration-to-photosynthesis ratio is re-established. leaf labile carbon, starch and protein pools decrease in response to increased temperature, eventually reaching a new, lower steady state, consistent with the general pattern of plant carbohydrate and soluble protein content being negatively correlated with growth temperature (dewar et al. 1999; wan et al. 2009). these acclimation responses represent intraspecific smp to more rmp functionality shifts (smp-to-rmp temperature acclimation). this has consequences for forage quality, such as reduced carbohydrate and mineral storage levels (progressively lowered carbon-nutrient quality) and reduced nitrogen quality due to nitrogen dilution and lowered digestibility resulting from high rmp mode-based growth rates (figure 5). in the climate–vegetation response model the expression of relative rmp to smp functionality results from, inter alia, the acclimation to changing temperature conditions. in response to increasing tmax (daytime temperatures), the productivity of tropical and subtropical forage plants (especially c4 grasses) is increased (long 1999; luo 2007; wan et al. 2005), resulting in greater quantities of plant protein in the landscape (increased nitrogen productivity). tmax was found to be positively correlated with tmin (r = 0.52, n = 30, p < 0.004) and negatively correlated with the amounts of summer rainfall (r = 0.54, n = 30, p < 0.003). phases of relatively elevated tmax therefore indicate conditions of increased solar radiation (reduced cloud cover) and reduced water availability for plant growth (reduced rainfall). the associated increased photosynthesis-to-growth ratio is conducive to enhanced forage quality (increased storage allocation; reduced nitrogen dilution). during the initial phases of increasing temperature, tmax is relatively high compared with tmin and relatively high diurnal temperature ranges are thus encountered, supporting a pulse of enhanced smp productivity (increased tmax/tmin ratio, increased tmax – tmin differential, and increased ratio of photosynthesis to respiration). this forms the basis of the first of four defined temperature acclimation phase states (smp-p, phase of increased smp productivity) constituting the temperature acclimation sequence resulting from (pulses of) increasing atmospheric temperature (figure 6). however, as the tmax/tmin differential (diurnal temperature range; dtr) declines as nocturnal warming (tmin) increases faster than daytime temperatures, smp-to-rmp mode acclimation occurs (decreased tmax/tmin ratio, decreased tmax – tmin differential, and increased ratio of respiration to photosynthesis) and sequential temperature acclimation phase states come into effect (figure 6). the initially enhanced smp mode productivity declines and rmp-mode growth increases [referred to as over-compensation by wan et al. (2009)]. the associated shift from smp-tw/tw to rmp-tw functionality has physiological consequences which compromise forage quality (figure 5), notably owing to a shift from storage-based to relative growth-dominated metabolism. this results in a reduction of stored metabolites (tnc, minerals and protein) relative to the accumulation of structural carbon through accelerated growth (reduced carbon-nutrient quality; figure 5). furthermore, rmp-tw mode functionality implies metabolic growth activity at higher water and temperature optima. growth realisation thus takes place under conditions of concurrently high temperature and water availability (tw conditions; at high water and temperature optima), constraining plant productivity under conditions when growth-curbed physiological activity (based on smp-tw functional efficiency) associated with the production of high-quality forage (early morning hours, early in season, dry season growth: smp-tw functionality) could have occurred. dry-season rainfall supports smp-tw functionality, thus promoting the production of forage items of high carbon-nutrient and nitrogen quality (figure 5). however, with progressive smp-to-rmp acclimation smp-tw efficiency is expected to decline as rmp-tw functionality favours growth under conditions of higher temperature and water availability. smp-to-rmp temperature acclimation (shifts from smp-tw/tw/tw to rmp-tw functionality) is therefore expected to result in forage with lower nitrogen quality (figure 5: nitrogen dilution and lowered dry-season forage digestibility) and notably lower carbon-nutrient quality (figure 5: low metabolic storage allocation priority). figure 6: sustained metabolic performance (smp) to resource-responsive metabolic performance (rmp) mode temperature acclimation sequence involving four phase states: an initial smp productivity pulse (smp-t/smp-t → smp-p) followed by temperature acclimation (smp-t/smp-p → smp-t → rmp-t) along the smp-rmp acclimation trajectory (figure 5) of intraspecific plant metabolic responses to temperature variables. spatiotemporal interactions in temperature acclimation and functionality settings: following from the climate–vegetation response patterns postulated under the climate–vegetation response model we expect pulses of increased forage availability induced by pseudo-cyclic temperature increases and progressive long-term shifts towards increased rmp functionality settings.• pulses of increased forage availability due to increased productivity of quality forage induced by pseudo-cyclic pulses of temperature increases (figure 2 and 3: increasing towards 1940; increasing towards 1960; increasing after 1975): in response to transient rising tmax, pulses of smp productivity (smp-p temperature acclimation phase state) occur when the temperature optima for plant functionality are still relatively low (tw metabolic activity). however, as acclimation to tmin takes effect, increased rmp-tw metabolic activity of plants results in reductions in forage quality and thus forage availability (figure 6). • progressive long-term shifts towards relatively advanced rmp functionality settings in response to cumulative acclimation to tmin and declining diurnal temperature ranges: trends of progressively deteriorating carbon nutrient quality and increasing nitrogen productivity-to-nitrogen quality ratios (figure 5) over the past century were accordingly anticipated (figure 2). owing to higher and more predictable levels of water availability to plants, such long-term shifts towards higher rmp functionality settings would have taken effect earlier (see figures 2 and 3, notably in response to prolonged temperature increases between approximately 1930 and 1940–1945) and to more advanced rmp settings in the southern knp (table 1). furthermore, as plants are already functioning at higher rmp settings in the southern knp, smp–rmp–smp temperature response pulses (figure 6) are expected to be attenuated relative to those in the northern knp, where pronounced associated pulses of forage availability are expected to be encountered. discussion top ↑ model postulates and predictions the postulates and predictions inferred from the climate–vegetation response model represent an explanatory framework geared to permit, inter alia, the interpretation of herbivore population performance patterns in space and over time. based on the postulates of this model inferences can be made as to how the metabolic responses of plants to their dynamic edaphoclimatic environment shape forage availability for herbivores by affecting the levels of key forage resources (relating to nitrogen productivity, nitrogen quality and carbon-nutrient quality). key forage resources the climate-vegetation response model differentiates three key indicators affecting forage availability to herbivores: nitrogen productivity, nitrogen quality and carbon-nutrient quality. nitrogen productivity refers to the quantitative availability of forage items for herbivores of a given nitrogen quality (plant protein content) as a result of biomass production. high nitrogen quality of forage implies high nitrogen concentrations of plant tissues; that is, plant protein content not unduly diluted by structural or non-structural carbon and of relatively high digestibility. carbon nutrient quality forage items are characterised by relatively high concentrations of non-structural carbohydrates and ash minerals resulting from metabolic allocation to storage under circumstances promoted by constrained growth (low temperatures, moisture stress: tw conditions), but sustained photosynthesis. levels of these key forage resource indicators in plants (referring primarily to c4 grasses) are determined by plant metabolic responses to the spatiotemporally dynamic availability of resources (temperature, water and nutrients; figure 5). herbivore species guilds of nutritional requirements as inferred from the nature of the diets of eight large savanna herbivore species, three herbivore species guilds of nutritional requirements were identified: bulk feeders with tolerance to fibrous herbage (buffalo and waterbuck), bulk feeders with preference for high nitrogen quality forage (short-grass grazers: blue wildebeest and zebra) and selective feeders for which dietary items of relatively high carbon-nutrient quality represent key forage resources (selective grazers: sable antelope, roan antelope, tsessebe and eland). spatial distribution of key forage resources plant metabolic adjustments to the north–south water availability and altitudinal temperature gradients are expected to determine the spatial distribution of smp to rmp functionality settings across the knp (table 1). the general pattern involved higher smp functional expression in the drier, northern areas of the park and at higher altitudes, and advanced rmp functionality settings towards the south. as the smp mode involves relatively growth-curbed, storage-based metabolism, the availability of forage of enhanced carbon-nutrient quality is accordingly expected to be higher in the northern knp and at higher altitudes. metabolic performance responses of intermediate smp–rmp modes favour lower levels of nitrogen dilution and are associated with forage of relatively high nitrogen quality, especially of plants on nutrient-rich geological substrates (table 1: central knp). climate-driven temporal trends in forage availability according to the climate–vegetation response model, both shorter-term transient temperature acclimation pulses (figures 2 and 3: increasing temperature towards 1940; increasing towards 1960; increasing after 1975 ) and longer-term shifts towards advanced rmp functionality settings are predicted to have occurred in response to temperature trends over the past century (figure 2). these temperature acclimation responses are expected to have resulted in transient pulses of increased forage availability (increased nitrogen and carbon-nutrient quality; figure 6), as well as the progressive long-term decline of carbon-nutrient quality of forage and an increasing nitrogen productivity/nitrogen quality ratio associated with advanced rmp functionality settings (figure 5). spatiotemporal interaction in forage availability accentuated temperature acclimation responses (figure 6) are predicted for circumstances where low and unpredictable water availability favour the prevalence of smp mode settings; that is, in northern relative to southern knp. more pronounced associated effects on nitrogen quality and carbon-nutrient quality are thus expected towards the northern areas of the park, with attenuation or absence of such responses where relatively high rmp functionality settings already predominate towards the southern knp. conclusion top ↑ the climate–vegetation response model as outlined here was applied as an explanatory framework with respect to herbivore population performance patterns in the knp. such application resulted in the internally consistent interpretation of spatiotemporal patterns of large herbivore population performance in relation to climate (seydack et al. 2012). acknowledgements top ↑ we acknowledge the effort of rangers, other field staff and scientists in collecting diverse data sets (e.g. animal population censuses and grassland monitoring) over many decades, without which this study would not have been possible. mss judith kruger, sandra macfadyen and sharon thomson of scientific services, knp, are acknowledged for providing data used in this study. the south african weather service provided temperature and rainfall data. this study represents an output of the specialist scientist research programme in systems ecology conducted by members of the conservation services division (knysna and skukuza) of south african national parks. a.h.s, in his capacity as honorary research associate at the botany department, university of cape town, acknowledges general support received from this institution. competing interests the authors declare that they have no financial or personal relationship(s) which may have inappropriately influenced them in writing this paper. authors’ contributions a.h.s. (garden route national park) was responsible for conceptualising and developing the climate–vegetation response model. c.c.g. (kruger national park), i.p.s. (kruger national park) and n.z. (kruger national park) contributed to in-depth discussions through sharing their knowledge of animal and plant ecology pertaining to the knp. w.j.v. (garden route national park) acted as field project leader of the research into climate effects on forage quality and j.b. (garden route national park) was involved in data processing. references top ↑ an, y., wan, s., zhou, z., subedar, a.a., wallace, l.l. & luo, y., 2005, ‘plant nitrogen concentration, use efficiency, and contents in a tallgrass prairie ecosystem under experimental warming’, global change biology 11, 1733–1744. http://dx.doi.org/10.1111/j.1365-2486.2005.01030.xbaas, w.j., 1989, ‘secondary plant compounds, their ecological significance and consequences for the carbon budget. introduction to the carbon/nutrient cycle theory’, in h. lambers, m.l. cambridge, h. konings & t.l. pons (eds.), causes and consequences of variation in growth rate and productivity of higher plants, pp. 313–340, s p b academic publishing, the hague. balasko, j.a. & smith, d., 1971, ‘influence of temperature and nitrogen fertilization on the growth and composition of switchgrass (panicum virgatum l.) and timothy (phleum pratense l.) at anthesis’, agronomy jounal 63, 853–857. http://dx.doi.org/10.2134/agronj1971.00021962006300060009x bijoor, n.s., czimczik, c.i., pataki, d.e. & billings, s.a., 2008, ‘effects of temperature and fertilization on nitrogen cycling and community composition of an urban lawn’, global change biology 14, 2119–2131. http://dx.doi.org/10.1111/j.1365-2486.2008.01617.x bodenstein, v., meissner, h.h. & van hoven, w., 2000, ‘food selection by burchell’s zebra and blue wildebeest in the timbavati area of the northern province lowveld’, south african journal of wildlife research 30(2), 63–72. buchmann, n., brooks, j.r., rapp, r.d. & ehleringer, j.r., 1996, ‘carbon isotope composition of c4 grasses is influenced by light and water’, plant, cell and environment 19, 392–402. http://dx.doi.org/10.1111/j.1365-3040.1996.tb00331.x codron, j., codron, d., lee-thorp, j.a., sponheimer, m., bond, w.j., de ruiter, d. et al., 2005, ‘taxonomic, anatomical, and spatio-temporal variations in the stable carbon and nitrogen isotopic compositions of plants from an african savanna’, journal of archaeological science 32, 1757–1772. http://dx.doi.org/10.1016/j.jas.2005.06.006 dewar, r.c., medlyn, b.e. & mcmurtrie, r.e., 1999, ‘acclimation of the respiration/photosynthesis ratio to temperature: insights from a model’, global change biology 5, 615–622. http://dx.doi.org/10.1046/j.1365-2486.1999.00253.x dörgeloh, w.g., 2006, ‘habitat suitability for tsessebe damaliscus lunatus lunatus’, african journal of ecology 44, 329–336. http://dx.doi.org/10.1111/j.1365-2028.2006.00654.x dunham, k.m., robertson, e.f. & grant, c.c., 2004, ‘rainfall and the decline of a rare antelope, the tsessebe (damaliscus lunatus lunatus), in kruger national park, south africa’, biological conservation 117, 83–94. http://dx.doi.org/10.1016/s0006-3207(03)00267-2 dwyer, s.a., ghannoum, o., nicotra, a. & von caemmerer, s., 2007, ‘high temperature acclimation of c4 photosynthesis is linked to changes in photosynthetic biochemistry’, plant, cell and environment 30, 53–66. http://dx.doi.org/10.1111/j.1365-3040.2006.01605.x, pmid:17177876 ehleringer, j.r., cerling, t.e. & helliker, b.r., 1997, ‘c4 photosynthesis, atmospheric co2, and climate’, oecologia 112, 285–299. http://dx.doi.org/10.1007/s004420050311 engels, e.a.n. & van der merwe, f.j., 1967, ‘application of an in vitro technique to south african forages with special reference to the effect of certain factors on the results’, south african journal of agricultural science 10, 983. estiarte, m. & peñuelas, j., 1999, ‘excess carbon: the relationship with phenotypical plasticity in storage and defense functions of plants’, orsis 14, 159–203. fryxell, j.m., 1991, ‘forage quality and aggregation by large herbivores’, american naturalist 138(2), 478–498. http://dx.doi.org/10.1086/285227 fryxell, j.m., wilmshurst, j.f., sinclair, a.r.e., haydon, d.t., holt, r.d. & abrams, p.a., 2005, ‘landscape scale, heterogeneity, and the viability of serengeti grazers’, ecology letters 8, 328–335. http://dx.doi.org/10.1111/j.1461-0248.2005.00727.x gertenbach, w., 1983, ‘landscapes of the kruger national park’, koedoe 26, 9–121. grange, s. & duncan, p., 2006, ‘bottom-up and top-down processes in african ungulate communities: resources and predation acting on the relative abundance of zebra and grazing bovids’, ecography 29, 899–907. http://dx.doi.org/10.1111/j.2006.0906-7590.04684.x gureja, n. & owen-smith, n., 2002, ‘comparative use of burnt grassland by rare antelope species in a lowveld game ranch, south africa’, south african journal of wildlife research 32(1), 31–38. harrington, r., owen-smith, n., viljoen, p.c., mason, d.r. & funston, p.j., 1999, ‘establishing the causes of the roan antelope decline in the kruger national park, south africa’, biological conservation 90, 69–78. http://dx.doi.org/10.1016/s0006-3207(98)00120-7 heitkönig, i.m.a. & owen-smith, n., 1998, ‘seasonal selection of soil types and grass swards by roan antelope in a south african savanna’, african journal of ecology 36, 57–70. http://dx.doi.org/10.1046/j.1365-2028.1998.114-89114.x herms, d.a. & mattson, w.j., 1992, ‘the dilemma of plants: to grow or defend’, quarterly review of biology 67(3), 283–335. http://dx.doi.org/10.1086/417659 hofmann, r.r., 1973, the ruminant stomach: stomach structure and feeding habits of east african game ruminants, vol. ii, east african literature bureau, nairobi. hofmann, r.r., 1989, ‘evolutionary steps of ecophysiological adaptation and diversification of ruminants: a comparative view of their digestive systems’, oecologia 78, 443–457. http://dx.doi.org/10.1007/bf00378733 holdo, r.m., holt, r.d. & fryxell, j.m., 2009, ‘opposing rainfall and plant nutritional gradients best explain the wildebeest migration in the serengeti’, american naturalist 173(4), 431–445. http://dx.doi.org/10.1086/597229, pmid:19243258 houghton, j.t., meira filho, l.g., callander, b.a., harris, n., kattenberg, a. & maskell, k. (eds.), 1996, climate change 1995: the science of climate change, cambridge university press, cambridge. joubert, s.c.j., 1976, ‘the population ecology of the roan antelope (hippotragus equinus equinus) (desmarest, 1804) in the kruger national park’, dsc thesis, faculty of science, university of pretoria. joubert, s.c.j. & bronkhorst, p.h.l., 1977, ‘some aspects of the history and population ecology of the tsessebe damaliscus lunatus lunatus in the kruger national park’, koedoe 20, 125–145. joubert, s., 2007a, the kruger national park: a history, vol. i, high branching, johannesburg. joubert, s., 2007b, the kruger national park: a history, vol. ii., high branching, johannesburg. kennedy, a.d., biggs, h. & zambatis, n., 2003, ‘relationship between grass species richness and ecosystem stability in kruger national park, south africa’, african journal of ecology 41, 131–140. http://dx.doi.org/10.1046/j.1365-2028.2003.00391.x lambers, h. & poorter, h., 1992, ‘inherent variation in growth rate between higher plants: a search for physiological causes and ecological consequences’, advances in ecological research 23, 187–261. http://dx.doi.org/10.1016/s0065-2504(08)60148-8 lambers, h., chapin iii, f.s. & pons, t.l., 1998, plant physiological ecology, springer verlag, berlin. le roux, e., 2010, ‘habitat and forage dependency of sable antelope (hippotragus niger) in the pretoriuskop region of the kruger national park’, msc thesis, faculty of science, university of the witwatersrand. long, s.p., 1999, ‘environmental responses’, in r.f. sage & r.k. monson (eds.), c4 plant biology, pp. 215–249, academic press, san diego. http://dx.doi.org/10.1016/b978-012614440-6/50008-2 luo, y., 2007, ‘terrestrial carbon-cycle feedback to climate warming’, annual review of ecology, evolution and systematics 38, 683–712. http://dx.doi.org/10.1146/annurev.ecolsys.38.091206.095808 macandza, v.a., owen-smith, n. & cross, p.c., 2004, ‘forage selection by african buffalo in the late dry season in two landscapes’, south african journal of wildlife research 34(2), 113–121. magome, d.t., 1991, ‘habitat selection and feeding ecology of the sable antelope, hippotragus niger niger (harris 1838), in pilanesberg national park, bophuthatswana’, msc thesis, faculty of science, university of the witwatersrand. magome, h., cain, j.w.i., owen-smith, n. & henley, s.r., 2008, ‘forage selection of sable antelope in pilanesberg game reserve, south africa’, south african journal of wildlife research 38(1), 35–41. http://dx.doi.org/10.3957/0379-4369-38.1.35 ‘t mannetje, l., 1975, ‘effect of daylength and temperature on introduced legumes and grasses for the tropics and subtropics of coastal australia. 2. n-concentration, estimated digestibility and leafiness’, australian journal of experimental agriculture and animal husbandry 15, 256–263. http://dx.doi.org/10.1071/ea9750256 ‘t mannetje, l., 1982, ‘problems of animal production from tropical pastures’, in j.b. hacker (ed.), nutritional limits to animal production from pastures, pp. 67–85, commonwealth agricultural bureaux, farnham royal. ‘t mannetje, l, 1984, ‘nutritive value of tropical and subtropical pastures, with special reference to protein and energy deficiency in relation to animal production’, in f.m.c. gilchrist & r.i. mackie (eds.), herbivore nutrition in the subtropics and tropics, pp. 56–66, the science press, craighall. mutanga, o., prins, h.h.t., skidmore, a.k., van wieren, s., huizing, h., grant, r. et al., 2004, ’explaining grass-nutrient patterns in a savanna rangeland of southern africa’, journal of biogeography 31, 819–829. http://dx.doi.org/10.1111/j.1365-2699.2004.01072.x osborne, c.p., 2008, ‘atmosphere, ecology and evolution: what drove the miocene expansion of c4 grasslands?’, journal of ecology 96, 35–45. pmid:18784799, pmcid:2517376 owen-smith, n. & ogutu, j.o., 2003, ‘rainfall influences on ungulate population dynamics’, in j.t. du toit, h.c. biggs & k.h. rogers (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 310–331, island press, washington dc. owen-smith, n., mason, d.r. & ogutu, j.o., 2005, ‘correlates of survival rates of ten african ungulate populations: density, rainfall and predation’, journal of animal ecology 74, 774–778. http://dx.doi.org/10.1111/j.1365-2656.2005.00974.x owen-smith, n. & mills, m.g.l., 2006, ‘manifold interactive influences on the population dynamics of a multispecies ungulate assemblage’, ecological monographs 76, 73–92. http://dx.doi.org/10.1890/04-1101 owen-smith, n. & mills, m.g.l., 2008, ‘shifting prey selection generates contrasting herbivore dynamics within a large-mammal predator-prey web’, ecology 89(4), 1120–1133. http://dx.doi.org/10.1890/07-0970.1, pmid:18481536 parrini, f., 2006, ‘nutritional and social ecology of the sable antelope in a magaliesberg nature reserve’, phd thesis, school of animal, plant and environmental sciences, university of the witwatersrand. pienaar, u de v., 1963, ‘the large mammals of the kruger national park – their distribution and present-day status’, koedoe 6, 1–37. schulze, r.e., 1995, hydrology and agrohydrology: a text to accompany the acru 3.00 agro-hydrological modeling system, department of agricultural engineering, university of natal, pietermartizburg. schulze, r.e., 1997, south african atlas of agrohydrology and –climatology, department of agricultural engineering, university of natal, pietermaritzburg. seydack, a.h., grant, c.c., smit, i.p., vermeulen, w.j., baard, j. & zambatis, n., 2012, ‘large herbivore population performance and climate in a south african semi-arid savanna’, koedoe 54(1), art. #1047, 20 pages. http://dx.doi.org/10.4102/koedoe.v54i1.1047 skidmore, a.k., ferwerda, j.g., mutanga, o., van wieren, s.e., peel, m., grant, r.c. et al., 2010, ‘forage quality of savannas: simultaneously mapping foliar protein and polyphenols for trees and grass using hyperspectral imagery’, remote sensing of environment 114, 64–72. http://dx.doi.org/10.1016/j.rse.2009.08.010 skinner, j.d. & chimimba, c.t., 2005, the mammals of the southern african subregion, 3rd edn., cambridge university press, cambridge. smuts, g.l., 1972, ‘seasonal movements, migration and age determination of burchell’s zebra (equus burchelli antiquorum, h. smith, 1841) in the kruger national park’, msc thesis, faculty of science, university of pretoria. taylor, g.j., 1989, ‘maximum potential growth rate and allocation of respiratory energy to stress tolerance in plants’, plant physiology and biochemistry 27, 605–611. tilley, j.m.a. & terry, r.a., 1963, ‘a two-stage technique for the in vitro digestion of forage crops’, journal of the british grassland society 18, 104–111. http://dx.doi.org/10.1111/j.1365-2494.1963.tb00335.x traill, l.w., 2004, ‘seasonal utilization of habitat by large grazing herbivores in semi-arid zimbabwe’, south african journal of wildlife research 34(1), 13–24. tudsri, s.h., matsuoka, h. & kobashi, k., 2002, ‘effect of temperature on seedling growth characteristics of panicum maximum’, tropical grasslands 36, 165–171. van lavieren, l.p. & esser, j.d., 1979, ‘numbers, distribution and habitat preference of large mammals in bouba ndjida national park, cameroon’, african journal of ecology 17, 141–153. van soest, p.h., mertens, d.r. & deinum, b., 1978, ‘preharvest factors influencing quality of conserved forage’, journal of animal science 47, 712–720. venter, f.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation patterns’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83–129, island press, washington dc. wan, s., hui, d.f., wallace, l.l. & luo, y., 2005, ‘direct and indirect effects of experimental warming on ecosystem carbon processes in a tallgrass prairie’, global biogeochemical cycles 19, gb2014. http://dx.doi.org/10.1029/2004gb002315 wan, s., xia, j., liu, w. & niu, s., 2009, ‘photosynthetic overcompensation under nocturnal warming enhances grassland carbon sequestration’, ecology 90(10), 2700–2710. http://dx.doi.org/10.1890/08-2026.1, pmid:19886480 watson, l.h. & owen-smith, n., 2000, ‘diet composition and habitat selection of eland in semi-arid shrubland’, african journal of ecology 38, 130–137. http://dx.doi.org/10.1046/j.1365-2028.2000.00229.x whyte, i.j. & joubert, s.c.j., 1988, ‘blue wildebeest population trends in the kruger national park and the effect of fencing’, south african journal of wildlife research 18(3), 78–87. wilson, j.r., 1975, ‘influence of temperature and nitrogen on growth, photosynthesis and accumulation of non-structural carbohydrate in a tropical grass, panicum maximum var. trichoglume’, netherlands journal of agricultural science 23, 48–61. wilson, j.r., 1982, ‘environmental and nutritional factors affecting herbage quality’, in j.b. hacker (ed.), nutritional limits to animal production from pastures, pp. 111–131, commonwealth agricultural bureaux, farnham royal. wilson, j.r. & ford, c.w., 1971, ‘temperature influences on the growth, digestibility, and carbohydrate composition of two tropical grasses, panicum maximum var. trichoglume and setaria sphacelata, and two cultivars of the temperate grass lolium perenne’, australian journal of agricultural research 22, 563–571. http://dx.doi.org/10.1071/ar9710563 wilson, j.r. & ford, w., 1973, ‘temperature influences on the in vitro digestibility and soluble carbohydrate accumulation of tropical and temperate grasses’, australian journal of agricultural research 24, 187–198. http://dx.doi.org/10.1071/ar9730187 wilson, j.r., taylor, a.o. & dolby, g.r., 1976, ‘temperature and atmospheric humidity effects on cell wall content and dry matter digestibility of some tropical and temperate grasses’, new zealand journal of agricultural research 19, 41–46. wilson, j.r. & ‘t mannetje, l., 1978, ‘senescence, digestibility and carbohydrate content of buffel grass and green panic leaves in swards’, australian journal of agricultural research 29, 503–516. http://dx.doi.org/10.1071/ar9780503 zambatis, n., 2004, ‘summary report of trends in herbaceous veld condition in the ranger sections of the kruger national park during the period 1989 to 2004’, scientific services, kruger national park, skukuza. article information authors: adrian j. armstrong1 sharon l. louw2 affiliations: 1biodiversity research & assessment division, ezemvelo kwazulu-natal wildlife, south africa2ecological advice, ezemvelo kwazulu-natal wildlife, south africa correspondence to: adrian armstrong postal address: po box 13053, cascades 3202, south africa dates: received: 05 mar. 2013 accepted: 17 aug. 2013 published: 10 oct. 2013 how to cite this article: armstrong, a.j. & louw, s.l., 2013, ‘monitoring of the eggs of the karkloof blue butterfly, orachrysops ariadne, for its conservation management’, koedoe 55(1), art. #1150, 11 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1150 copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. monitoring of the eggs of the karkloof blue butterfly, orachrysops ariadne, for its conservation management in this original research... open access • abstract • introduction • research methods and design    • monitoring area at the orachrysops ariadne colony near nkandla    • development of a monitoring method for the three orachrysops ariadne colony sites near howick       • mapping the host plant distribution boundaries at the three colony sites near howick       • pilot study at the start orachrysops ariadne colony       • camponotus natalensis distribution in relation to the distribution of indigofera woodii var. laxa    • implementation of the 'mature host plants index method' at the start    • statistical analyses and geographic information systems • results    • monitoring at the nkandla colony       • browsing, fire and host plants       • egg numbers    • pilot study to determine a monitoring method for the three colony sites near howick       • host-plant distribution boundaries       • comparison of the plot and the 'mature host plants sample' monitoring methods       • mapping the host-ant distribution at the stirling colony breeding area       • implementation of the 'mature host plants index' method at the start • discussion    • monitoring and habitat management at the nkandla colony    • monitoring and habitat management at the three colony sites near howick    • future priorities • conclusion • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ the endangered orachrysops ariadne (butler 1898) (karkloof blue butterfly) is endemic to the endangered moist midlands grassland in kwazulu-natal, south africa, and is extant at four sites. the results from the monitoring of the eggs laid by o. ariadne in a grassland area that is frequently burnt by poor rural people to ensure that palatable grass is available to their livestock, suggested the implementation of management interventions (fencing and firebreak burning) to prevent the local extinction of the butterfly. the number of eggs at the monitoring site declined dramatically between 2002 and 2003 and fluctuated after the management interventions were initiated properly in 2008, but had nearly reached the target number of 250 by 2013. an index count method for the monitoring of o. ariadne eggs at the other three known colonies, where plant invasion rather than uncontrolled burning is a major threat, was developed and shown to be efficient with regard to time relative to the number of eggs sampled. the host ant camponotus natalensis (f. smith 1858) (natal sugar ant) was found to be present in all the host-plant patches at one colony site, indicating that all host-plant patches are likely to be breeding areas for the butterfly. invasive plant control at and appropriate burning of the habitat of o. ariadne should assist in ensuring the survival of these colonies.conservation implications: adaptive monitoring and management of threatened endemic invertebrates and their habitats may be crucial for their continued survival. the development of efficient methods for the monitoring of such species is required where resources are limited, as threats to the species may cause sudden and irreversible declines in population size. introduction top ↑ endangered species should not lose any more of their habitat if they are to survive in the wild in the long term (pfab, victor & armstrong 2011). monitoring of populations of threatened lepidopteran species that have obligatory relationships with host plants or ants will almost always be required for their effective conservation, especially where land transformation or habitat degradation is proceeding apace (jewitt 2012; murphy & weiss 1988). adaptive monitoring that enables adaptive management interventions to be initiated as and when appropriate, and that informs managers of the nature of the threats to the species and potential counters to those threats, would be very useful (campbell et al. 2002; greenwood & robinson 2006; lindenmayer & likens 2009; lindenmayer et al. 2011). monitoring can be costly and therefore should be initiated only if clear and tangible benefits for the conservation of the species will result (field et al. 2007).the threats to a threatened species and knowledge of the biology of the species should guide the development of a monitoring programme and implementation of management actions for that species (campbell et al. 2002), although certain threats may be detected only once a monitoring programme is underway. monitoring of lepidoteran species needs to be long term, as responses of threatened butterflies to management interventions may not be immediate and may depend on other interacting factors such as weather conditions (murphy & weiss 1988; smee et al. 2011; thomas 2005). long-term monitoring of threatened butterfly species is well established on other continents (e.g. swengel et al. 2011; thomas 2005), but is only in its infancy in africa (for south africa, see edge et al. 2013; mcgeoch et al. 2011). the endangered orachrysops ariadne (butler 1898) (karkloof blue butterfly) is known from only four colonies, all of which are in the highly transformed and endangered midlands mistbelt grassland in kwazulu-natal, south africa (figure 1; armstrong et al. 1998; armstrong & edge 2013; henning, terblanche & ball 2009; lu & samways 2001, 2002a, 2002b; mucina & rutherford 2006). none of the colonies has protected area status yet. the butterfly has an obligate host relationship with indigofera woodii bolus var. laxa bolus (lax variety of wood’s indigo plant) and camponotus natalensis (f. smith 1858) (natal sugar ant), laying eggs only on the host plant and apparently only where the host ant is present (lu & samways 2001). (see pierce et al. [2002] for further discussion of ant-dependent oviposition, but cf. edge, cilliers & terblanche [2008] for the related orachrysops niobe [trimen 1862] [brenton blue butterfly]). the larvae feed only on the host plant and the adults feed on a limited number of nectar-plant species that flower during their flight period (lu & samways 2001). any conservation action implemented for the butterfly must therefore conserve the butterfly’s host-plant, host-ant and nectar-plant relationships and other required habitat conditions for it to be successful (swengel et al. 2011; tudor et al. 2004). the work by lu and samways (2001, 2002a, 2002b) elucidated some of the life history and behavioural ecology of the butterfly and enabled the formulation of conservation management recommendations. a conceptual ecological model of required and threatening interactions during the life cycle of o. ariadne is presented in figure 2. figure 1: location of the four known colonies of orachrysops ariadne in relation to the original extent of the moist midlands mistbelt vegetation type and two urban areas. figure 2: ecological model of required and threatening interactions during the life cycle of orachrysops ariadne. at least one known colony of o. ariadne, near balgowan, became extinct last century and from verbal reports and the literature it seems probable that other colonies have also been extirpated through land transformation or habitat degradation (armstrong 2010; henning et al. 2009; pennington 1946; pringle, henning & ball 1994; samways & lu 2007). mixed-species alien plant infestation was observed at the extinct colony site by the first author. potential threats and other factors relating to the four extant colonies are presented in table 1. all current landowners or land occupiers are very supportive of the monitoring of the colonies and management of the colony sites for the conservation of o. ariadne. table 1: characteristics of, and factors relating to, the known orachrysops ariadne colonies and colony sites. monitoring of the o. ariadne colonies would be an important component of the management programme for this species, because it enables the current threats to the colonies to be determined, the response of the populations to management actions to be ascertained and timely corrective action to be initiated and maintained (campbell et al. 2002; greenwood & robinson 2006). monitoring of the eggs is a simple and efficient way of determining the trend in a population of o. ariadne, as well as ascertaining the factors that are causing a decline in the population (lu & samways 2001). the eggs are clearly visible on the host plant as they are deposited near the ends of growth tips of the mature host plant, which are not covered by other vegetation, and the eggs whiten with age from their initial green colour (lu & samways 2001; pers. obs.). monitoring of non-adult life cycle stages can be, and has been, used to detect population size changes in some butterfly species (e.g. hula et al. 2004; murphy & weiss 1988; smee et al. 2011). monitoring methods that are inexpensive and easy to implement by the managers or inhabitants of the land on which the colonies occur would be very useful (mcgeoch et al. 2011).a conceptual model for the management and monitoring of o. ariadne is presented in figure 3. the management objectives for o. ariadne will be achieved through various means, including adaptive management of the fire regime, fire control, livestock exclusion, invasive plant control, and through restoration of habitat for and re-introductions of o. ariadne. the key performance indicator for success of the monitoring and management programme is a sustained increase in the number of eggs laid (the primary response variable) until a set target number is reached, and thereafter the maintenance of egg numbers near this target indefinitely (figure 3). some variables that may have an influence on the value of the primary response variable are presented in table 2. these variables may be measured during monitoring. figure 3: conceptual model for the management and monitoring of orachrysops ariadne. table 2: variables that may have an influence on the primary response variable (number of orachrysops ariadne eggs laid). only a short monitoring period for o. ariadne is consistently available each year as the species has a short flight period and embryonic development period (lu & samways 2001). few personnel will be available to undertake the monitoring. these constraints would have to be taken into account when developing the monitoring programme. the objectives of the present study were to:• determine whether management of threats to the nkandla o. ariadne colony that caused its rapid decline, as ascertained through the long-term monitoring of egg numbers at the colony site, would change the trajectory of egg numbers towards a target of at least the number of eggs recorded during the first monitoring occasion • develop and test a cost-effective monitoring index method for the other o. ariadne colonies to ensure that major declines in population size can be detected in time to prevent possible extirpation of the colonies. research methods and design top ↑ monitoring area at the orachrysops ariadne colony near nkandla a small valley, with an intermittent stream, was chosen as the monitoring area (0.24 ha) for the nkandla colony, the breeding areas of which are discrete patches distributed in nearly 4 ha of grassland. this monitoring site was chosen because of its accessibility and the large number of host plants present and because the number of eggs present on the host plants could be counted during a day by a team of at least six staff. the latter requirement refers to the amount of time and number of personnel that are available to do the monitoring. host plants growing in the several other small valleys inhabited by the butterfly were not included in the monitoring owing to resource constraints and to avoid excessive disturbance of the breeding habitat (samways, mcgeoch & new 2010). the boundary of the monitoring area was taken to be the edge of the valley (i.e. the top of the small ridge or watershed on both sides) and where the host plant distribution ended below the cliff at the top and above the road bank at the bottom.the adult o. ariadne are on the wing from early march to late april, with male butterflies eclosing before the female insects. the maturation period for the developing larvae ranges from about 18–30 days in the field (lu & samways 2001). monitoring was carried out 1 month after the last eggs were likely to have been laid in order to minimise the number of eggs that might not have hatched yet and also the number of eggshells that might be dislodged by frost or other factors. the monitoring commenced in 2002. all host plants considered large enough to be suitable for oviposition were searched for eggs; that is, single-stemmed seedlings were excluded. the precise number of i. woodii var. laxa surveyed in 2002 was not recorded as this was not deemed necessary then. avoidance of double counting was implemented through awareness, communication between the monitors and the vigilance of the recorder. the number of monitors varied over the years, from 3 in 2003 to 19 in 2009 (median and mode: 10 monitors; mean ± 1 sd = 9.5 ± 4.25 monitors; n = 12). there was no significant correlation between egg numbers and number of monitors (r = -0.172; n = 12). eggs were classified visually according to the system of lu and samways (2001), using the naked eye or a magnifying glass. an egg from which a larva has successfully hatched had a round, clean-edged exit hole. we classified eggs that showed no signs of damage or parasitism as unhatched, being either infertile or still unhatched (sometimes the head capsule of the unhatched larva could be distinguished). eggs that had a minute round hole on the surface in addition to the micropyle were classified as parasitised and combined with the damaged class, and all other eggs were classified as damaged (e.g. those that had ragged openings or were misshapen). i. woodii var. laxa were considered overgrown if covered by surrounding vegetation without the protrusion of at least two i. woodii var. laxa shoots on which the adult female butterflies could lay. i. woodii var. laxa were classified as senescing if yellowing leaves, leaf fall and withered stems were evident, and as browsed by mammals where nipped shoots (shoots with obvious obliquely truncated ends) were evident. plant heights were estimated to the nearest 10 cm and the presence of seed pods was recorded. the management interventions at the nkandla monitoring site included the burning of a firebreak around the site in 2006 and most years thereafter, and the fencing of the site in 2008. alien plant control occurred once (2006) during the monitoring period. since the approach taken is not experimental, the success of management interventions can only be inferred from the data until further factors that could potentially have caused the observed changes have been measured and a long-term dataset is obtained. development of a monitoring method for the three orachrysops ariadne colony sites near howick the monitoring of adult o. ariadne using a mark–recapture method was shown not to be feasible (samways & lu 2007) because of the cryptic habits of the female butterflies. a total count of eggs laid at the three more southern colonies is impractical because their breeding areas are much greater than at the monitoring area of the nkandla colony, a more limited number of monitors will be available, and the daily period of daylight suitable for monitoring eggs is less. therefore, a subsample of the total egg number at a colony would have to be monitored. the boundary of the host plant distribution at each colony would have to be mapped and the distribution of the host ant be determined in relation to that of the host plant to prevent potential bias in where sampling takes place. both hosts are essential to the completion of the butterfly’s life cycle and the mobility of early instar o. ariadne larvae is likely to be limited. as a result, the eggs may be patchily distributed. these aspects of the study were investigated using the following methods. mapping the host plant distribution boundaries at the three colony sites near howick the boundary of the i. woodii var. laxa distribution was determined at each of the three colonies near howick by searching for the host plants at the edge of suitable habitat and recording the positions of the edge host plants using a gps receiver. the boundary was mapped by connecting the outer host plant positions that were at most 10 m apart. single-stemmed seedlings were again ignored, as there was no certainty as to which would survive to maturity. the potential area of the o. ariadne breeding area could then be calculated. at stirling, i. woodii var. laxa were surveyed by walking transects and recording the positions of host plants encountered between april 2007 and january 2008. two to four observers walked approximately 2 m – 3 m abreast along transects, recording i. woodii var. laxa locations on and to about a metre on either side of the transects. these transects included straight transects through the habitat and zigzag transects along the apparent boundaries of the host plant patches to delimit the boundaries more accurately. the aim was not to record every single i. woodii var. laxa, but rather to define the limits of the various patches of host plants on the site. ‘outlier’ host plants were those more than 10 m away from the next nearest host plants, including groups of up to three host plants that were more than 10 m away from the next nearest host plant. from visual observations, the distribution of the i. woodii var. laxa appeared to be closely correlated to the distribution of the miscanthus capensis (nees) andersson (daba grass) on the site. therefore areas of this grass were targeted for searching during the survey. other areas were also searched, and from the data it became apparent that the distributions of i. woodii var. laxa and m. capensis were closely related, although the host plant was not as widely distributed as the grass species. pilot study at the start orachrysops ariadne colony both a plot method and an index method were compared for efficiency. the plot method might permit an estimate of the total number of eggs laid at the colony as the areas of both the plot and of the host plant distribution were known. it would also give an indication of whether a full egg count at the site was feasible given the constraints.a plot of 20 m × 20 m was placed roughly in the middle of the start study area (approximately 1090 m.a.s.l.), where the setup was facilitated by the reasonably even slope. the plot was divided into four smaller quadrants (10 m × 10 m each), identified as quadrants a (south-east quadrant), b (south-west quadrant), c (north-west quadrant) and d (north-east quadrant). this enabled accurate searching for host plants and detection of variation in host plant densities and egg densities over the whole plot. each quadrant was systematically searched for i. woodii var. laxa and o. ariadne eggs by three people walking roughly parallel along the contours of the hillside and recording the number of host plants and eggs. only plants rooted inside the plot were counted. the following features of the i. woodii var. laxa were recorded: presence of seed pods, signs of browsing by mammals and signs of senescence. whether the surrounding vegetation completely covered the host plant was also recorded, as were the number of flowering indigenous encroacher phymaspermum acerosum (dc.) källersjö (currys post plant) and the number of alien invader rubus cuneifolius pursh (american bramble) plants. the start and finish times were recorded for each quadrant. since 0.038 ha of the colony’s breeding area fell within the plot, the densities of host plants and eggs in this area were used to estimate the number of host plants and eggs in the whole breeding area of the colony, respectively. the densities of the invader or increaser plant species were calculated from the numbers of each species recorded in the entire 0.04-ha plot. an index monitoring method might be very efficient in that specific host plants could be targeted to increase the number of eggs recorded per unit effort. when developing an index monitoring method it will have to be considered that: • the number of host plants suitable for oviposition will vary from year to year according to the post-burn age of the vegetation • the density of host plants will vary over the colony area • the spatial distribution of the host ant is not likely to change in response to fire as the ants live underground and therefore should not be affected directly by fire • as a result of these and perhaps other factors, the distribution of o. ariadne eggs is likely to change somewhat each year (figure 4). figure 4: distribution of host plants with eggs at the nkandla monitoring site, as recorded during monitoring in 2002, 2008 and 2012, to illustrate changes in the distribution of eggs. therefore, the index monitoring method will have to include some randomness in its spatial component and the number of host plants that can be searched for eggs in a day will be one of the necessary considerations. during the pilot phase of the study, two observers walked through the colony’s breeding area and searched 50 large (mature) host plants suitable for oviposition as an indexing method, as they were encountered randomly in various parts of the site outside the plot. the data recorded were similar to those during the plot count (excluding the number of encroacher and invader plants), but also included a visual estimate (to the nearest 10 cm) of the relative height of each i. woodii var. laxa, as assessed against an observer’s height. camponotus natalensis distribution in relation to the distribution of indigofera woodii var. laxa the distribution of the host ant, c. natalensis, in relation to the host plant was investigated at stirling farm. sampling for the host ant was initiated in february 2008 and was completed by mid-march 2008. weather conditions were variable during the survey period and nights were frequently misty and wet. sampling was abandoned in rain. a grid of points 20 m apart was laid out over the site encompassing the area where i. woodii var. laxa grew. each point was used as a baiting station and the actual coordinates on the ground were captured using a hand-held gps receiver. each baiting station was a perspex square of 10 cm × 10 cm placed on cleared ground of approximately 30 cm in diameter. bait was applied in the late afternoon and left for 90 min before surveying. the bait consisted of a honey-and-brown sugar mix, diluted with water to give a watery consistency, and a small amount of bovril added to provide protein. the baiting stations were checked for ant activity after dark. each baiting site (including the baiting station, the bare ground surrounding the station and the peripheral vegetation, up to the top of the grass layer) was searched very carefully for about 1 min and the c. natalensis were counted. implementation of the ‘mature host plants index method’ at the start the ‘mature host plants index method’ was refined by determining the number of mature host plants that should be sampled at the start to ensure an appropriate egg sample size. an appropriate egg sample size was estimated using the formula (watson 2001):n = {(p[1–p])/(a2/z2)+p[1-p]/n}/r [eqn 1] where n = sample size required, n = estimated number of eggs in the population, p = estimated variance in the population, a = the precision desired (± 5%), z is the confidence level (95%, i.e. 1.96), and r is the estimated response rate (100%, i.e. 1.0). an estimate of the number of reproductively mature host plants that should be searched to obtain the required sample size of eggs was then calculated from the data collected during the pilot study at the start in 2006. two observers searched for eggs on this number of mature host plants (i.e. those with pods) at the start on 16 and 17 may 2012. they used a hand-held gps receiver to guide them along predetermined transects spaced 20 m apart and running north–south through the colony’s breeding area. these transects were numbered consecutively from west to east. the order of transects to be walked until the calculated number of mature host plants had been searched for eggs was determined via a random number generator. mature host plants observed within 2 m of the centre line of each transect walked were searched for eggs. statistical analyses and geographic information systems all statistical analyses were performed using statistica 10 (statsoft, inc., tulsa, oklahoma). spatial data building was performed using cartalinx (clark labs, clark university, worcester ma). the idrisi 15 andes version (clark labs, clark university, worcester ma, usa) and arcview 3.2 (esri, inc., redlands ca) geographic information systems were used for geographical coverage analyses. results top ↑ monitoring at the nkandla colony browsing, fire and host plants the relatively extensive browsing of the host plants was curtailed by the erection of a fence in 2008, after which no or little browsing was recorded (figure 5). uncontrolled fires burnt through the monitoring site each year between 2002 and 2005 (figure 5). the establishment of the firebreak enabled the fires and their effects on the host plants to be controlled most years thereafter. exceptions included the burning of the site by an uncontrolled fire in july 2007 after the egg monitoring took place that year, and in 2011, again after the egg count. during the latter fire, about half the site was burnt in august as part of the site management programme, followed later by an unauthorised fire that burnt the rest of the site. figure 5: status of the host plants indigofera woodii var. laxa at the nkandla monitoring site as recorded during monitoring. a decline in the number of i. woodii var. laxa was recorded between 2004 and 2006, when fewer than half the number of host plants surveyed in 2003 were visible owing to the uncontrolled fires (figure 5). a recovery in the numbers of i. woodii var. laxa surveyed occurred between 2007 and 2010, when the number exceeded the number in 2003, owing to the presence of the firebreak. another decline in visible host plant numbers was recorded in 2011 due to the fires that year, after which the number of visible host plants increased. the percentage of host plants with pods was low from 2006 to 2008, but increased markedly from 2009 to 2011 owing to the protective effect of the firebreak, which allowed the host plants and their re-growth to mature. the rejuvenating effect of fire is seen in the decline in the percentage of host plants showing signs of senescence between 2004 and 2007, and the subsequent increase from 2010 to 2011 in the absence of fire. signs of senescence declined to near zero and the percentage of host plants with pods declined in 2012 after the site was burned in 2011.the mean height of the host plants varied according to the frequency of uncontrolled fires, decreasing after each burn and generally increasing over the years without fire, as seen between 2008 and 2011 (figure 5). in 2005, after a series of annual burns and with extensive browsing, many of the host plants were overgrown by surrounding vegetation (figure 5). the percentage of host plants overgrown by surrounding vegetation gradually increased from 2009 to 2011, after control over fire and browsing was established. this percentage dropped practically to zero in 2012 after the entire monitoring site was burned in 2011. the number of i. woodii var. laxa with o. ariadne eggs declined overall by 93% between 2002 and 2005, fluctuated somewhat thereafter until 2010, and then increased until 2013 (table 3). table 3: number and fate of orachrysops ariadne eggs, number of indigofera woodii var. laxa host plants with eggs, number of host plants surveyed, and standardisation of the number of eggs at the nkandla monitoring site. egg numbers the results of the o. ariadne egg monitoring at the nkandla colony during the study period are presented in table 3. the number of eggs recorded decreased by 77% between 2002 and 2003, and by 95% between 2002 and 2005. after the management interventions were effectively implemented in 2008, the recorded number of eggs fluctuated and then increased year on year from 2011 to 2013, when the number of eggs recorded was nearly at target. no significant correlation was found between the number of eggs and the number of host plants between 2003 and 2013 (r = 0.167, n = 11, p > 0.05). the management of threats appears to have contributed to reversing the population trajectory of both o. ariadne and i. woodii var. laxa at the nkandla monitoring site. pilot study to determine a monitoring method for the three colony sites near howick host-plant distribution boundaries the host-plant distribution at the start was 1.35 ha in extent, consisting of two patches divided by a narrow strip of riparian vegetation. this was the estimated extent of the o. ariadne breeding area here. the area of the host-plant patch at wahroonga farm was calculated to be 8.15 ha. the boundaries of the host-plant patches (1.57 ha in total area) and the location of outlier i. woodii var. laxa at stirling farm are indicated in figure 6. host plants were not uniformly distributed over the site, but were concentrated on south-east-facing slopes. figure 6: distribution of patches of the host plant indigofera woodii var. laxa in relation to the presence and absence of the host ant camponotus natalensis on the stirling farm study site. comparison of the plot and the ‘mature host plants sample’ monitoring methods the 50 mature host plants sample method was much more efficient than the plot method (table 4). in the sample, a significantly higher proportion of host plants had pods (c2 = 70.49; df = 1; p < 0.0001) and eggs (c2 = 124.80; df = 1; p < 0.0001), and a significantly lower proportion were covered by other plants (c2 = 186.97; df = 1; p < 0.0001) compared with the plot (the other comparisons of proportions were not significant). even in the plot, which had host plants of a variety of ages, the majority of the host plants selected for oviposition by female o. ariadne were reproductively mature. the density of host plants and eggs in each plot quadrant varied between 31 and 106 and 0 and 7 per 100 m2, respectively. from the density of eggs and host plants recorded in the plot, an estimated 22% (71 out of 320) of the eggs at the colony were in the egg sample from the 50 mature host plants. a total of 38 p. acerosum plants, one maesa lanceolata forssk. (false assegai) shrub and one r. cuneifolius were recorded from the entire plot. from these figures, a total of 8384 i. woodii var. laxa, 320 eggs (107 hatched), 1283 p. acerosum plants, 34 m. lanceolata and 34 r. cuneifolius plants were estimated for the breeding area of the o. ariadne colony. table 4: comparison of results from the two monitoring methods employed at the start in 2006. mapping the host-ant distribution at the stirling colony breeding area the distribution of c. natalensis in relation to the i. woodii var. laxa distribution at stirling farm is shown in figure 6. the host ant was present in all the host-plant patches, but absent from some of the areas where host plants were absent. there were significantly more c. natalensis occurrences in the host-plant patches than expected and no absences of the host ant there (χ2 = 21.88; df = 1; p < 0.01). c. natalensis was recorded near many of the outlier host plants. from these results, only the distribution of the host plant needs to be taken into account in the monitoring method. implementation of the ‘mature host plants index’ method at the start using equation (1) and the estimates from the pilot study, the appropriate egg sample size was estimated at 88 eggs, with n estimated at 320 eggs and p estimated at 0.4 (approximately 40% of the 50 mature host plants sampled when testing the index method did not have o. ariadne eggs, suggesting that the population could increase by a maximum of 40%). searching of the 50 mature host plants during the pilot study returned 71 eggs. therefore, to have recorded an adequate sample of 88 eggs using the index method during the pilot study, the estimated number of mature host plants that would have had to be searched was 88/71*50 = 61.97, (i.e. 62) mature host plants.following from this calculation, 62 reproductively mature host plants were searched for eggs at the start colony in may 2012. most of the 31 eggs observed had hatched; 29% did not hatch because they were damaged (6) or assumed parasitised (3). three of the host plants with eggs showed signs of being browsed. the total area covered by the 547-m transect length walked was 2188 m2, representing 16.22% of the total colony breeding area. discussion top ↑ monitoring and habitat management at the nkandla colony the nkandla o. ariadne population was relatively large before and at the start of the monitoring programme (lu & samways 2001; this study). this population is a priority for conservation as it is separated from the other three known colonies by at least 120 km (figure 1). the precarious situation of the monitored population of o. ariadne at the nkandla site between 2003 and 2008 was the result of a lack of appropriate management of the colony area. the management guidelines for this colony included the necessity for maintaining a fire return frequency of 2–4 years (lu & samways 2002b) and for domestic livestock to be excluded from the breeding areas. the acceptance of, and involvement with, the management programme at nkandla by local residents was a very important factor in eventually achieving appropriate management of the site and one that should assist in eventually achieving the target number of eggs and the overall success of the project (louw & armstrong 2010). such cooperative management should become a feature of future invertebrate conservation programmes in africa to increase the probability that the management goals will be achieved.the large decline in the number of eggs at the monitored site between 2002 and 2003 was apparently partly caused by the browsing of the i. woodii var. laxa by domestic stock and perhaps also indigenous antelope. the eggs are usually laid within 3 cm of the tips of the buds of the i. woodii var. laxa and therefore the eggs are very vulnerable to accidental ingestion when the shoots are eaten by mammalian browsers (lu & samways 2002a). fencing of the monitoring site was finally achieved in 2008 (louw & armstrong 2010), which prevents cattle and goats from accessing the host plants at the site. cattle are known to have detrimental impacts on threatened butterfly host-plant abundance and habitat structure elsewhere (e.g. schtickzelle, turlure & baguette 2007). the decline in the number of visible i. woodii var. laxa through the agency of fire between the 2003 and 2004 monitoring events indicated the need for careful management of the habitat through fire. the firebreak around the nkandla monitoring site has protected the monitoring site from uncontrolled fires since 2008, allowing appropriate fire management of the site. fire is the best means of defoliation on the site as the vegetation and invertebrates such as o. ariadne are adapted to fire and many are dormant or underground during the fire season. the host plants are sensitive to fire before they are 2 years old (lu & samways 2001), but resprout as adults and also reproduce via seed. the vigour and health of the host plant community and the habitat as a whole are maintained by fire, including by enabling the recruitment of i. woodii var. laxa to offset the number that senesce. very few mature i. woodii var. laxa were visible in the monitoring area in 2005 owing to the prior uncontrolled fires (one in august 2004 and two in 2003). most of the remaining host plants were young and many were covered by other vegetation. female o. ariadne generally lay their eggs on the larger, mature i. woodii var. laxa and not on host plants covered by other vegetation (this study; lu & samways 2002a). browsing of i. woodii var. laxa also continued. as a result, few eggs were recorded in 2005, 2006 and 2007. the exclusion of fire in 2008, 2009 and 2010 caused an increase in the percentage of host plants showing signs of senescence from 0.6% in 2007 to 59.8% in 2011. this indicates the need for rejuvenation of the host plant population by fire. the controlled fire in 2011 (and apparently also the unauthorised burn that year) achieved this purpose, and in 2013 the egg numbers (235) had nearly reached the target of 250, the number recorded in 2002. the trend in egg numbers is positive and a potential extirpation of the species at the site has been avoided. although inappropriate burning of habitat affects some threatened butterfly species or subspecies (powell, busby & kindscher 2007; swengel et al. 2011), other threatened butterfly species require regular but not very frequent (and sometimes small-scale) burning of their habitats at the appropriate time of year and day to maintain habitat heterogeneity and the availability of required resources without killing vulnerable stages of the life cycles of the butterflies (see, e.g. new et al. 2010; pickens & root 2009). the management interventions appear to have contributed to reversing the population trajectory of both o. ariadne and i. woodii var. laxa and need to be extended to the adjacent grassland areas, including the south-facing seasonal stream valleys where o. ariadne is known to occur. the areas that would require fencing are relatively small and the loss of browse for domestic stock would not be extensive. the total activity area for an o. ariadne colony is larger than the area in which the host plant is distributed, as the adult nectar plants may be more widely distributed than the host plant (dennis, shreeve & van dyck 2006; samways & lu 2007). the monitoring of nectar resources for butterflies of conservation concern, such as o. ariadne, is important because such butterflies can be flower-feeding specialists that have short flight periods, low mobility and biotope specialisation (lu & samways 2001; samways & lu 2007; schultz & dlugosch 1999; tudor et al. 2004). monitoring of nectar plants should be added to the present monitoring programme. monitoring and habitat management at the three colony sites near howick the results obtained using the index method at the start suggested that sampling a certain number of mature i. woodii var. laxa for eggs is an efficient monitoring method with regard to time and resources and appears to be suitable for monitoring the status of other o. ariadne populations. furthermore, it avoids the greater disturbance of the vegetation around the host plants associated with the plot counts. congruence in the distribution of the host plant and the host ant, as determined at the stirling colony site, indicates that transects can be placed either systematically or randomly over the breeding area. assuming similar mean densities of host plants at all colony sites, and considering the ratio between the size of the breeding area at the start and the breeding areas at the two other colonies near howick, we recommend that at least 80 and 411 mature host plants are sampled for eggs on each monitoring occasion at stirling and wahroonga, respectively. at least four monitors would be required at wahroonga to ensure that the sampling is finished within 2 days. the index method will have to be put into practice for some years and tested to see whether it is sensitive enough to detect negative unidirectional trends in population size rapidly enough to enable effective management interventions to be implemented to reverse the declines (field et al. 2007). an increase in the robustness of estimates of the number of mature host plants to sample at each of the three colonies near howick is desirable in the future. to this end, sampling a sufficient number of randomly placed plots or transects at each site would allow measures of variability within and between populations to be calculated. the results of the monitoring at the nkandla colony suggest that monitoring should eventually be carried out at least every other year so that any major decline in population size is noticed quickly enough to effect a reversal.the percentage of i. woodii var. laxa in the plot (24.2%) and in the index method sample (16%) at the start that had signs of being browsed suggests that monitoring the eggs around the time of hatching of the last eggs is most appropriate as monitoring later may result in greater egg loss and monitoring earlier may under-estimate the number of eggs that hatched. browsing by antelope could lead to an underestimation of the number of eggs laid and the host plants, being legumes, are suitable browse in the relatively dry and cold winter of in the mistbelt region of kwazulu-natal. lu and samways (2001) found that all the eggs monitored at the start colony between 1997 and 1999 had hatched by 24 may. monitoring of eggs at the colonies near howick should therefore be carried out around this time of year. rotational burning of about a third of the grassland at each site each year may be useful for promoting seedling growth and survival in the burnt area and in rejuvenating the foliage of mature host plants that resprout after fire. how fire might affect the c. natalensis is unknown, but the butterfly’s habitat should be burnt during the day as the host ant is nocturnal and is underground during the day (lu & samways 2002a). burning should also be performed only after the butterfly eggs have hatched and when the larvae are underground in ant nests. the main threat to the habitat of o. ariadne at the three colonies near howick appears to be encroachment by indigenous plants and invasion by alien plants into the breeding areas. p. acerosum, m. lanceolata and r. cuneifolius should be monitored as these species might out-compete the host plants and other plants for resources such as light, water, soil nutrients and space. the number of p. acerosum at the start colony was estimated at several thousand, and although only one r. cuneifolius was recorded in the plot, many were found along the edges of the watercourses and stream. edge et al. (2008) found that the encroacher pteridium aquilinum (l.) kuhn (bracken fern) regrows rapidly after fire and preoccupies space from the slower-growing indigofera erecta thunb. (erect indigo plant), the host plant of o. niobe, resulting in a low recruitment success for the host plant. a high biomass of the relatively large p. acerosum could lead to intense fires, which could kill the younger i. woodii var. laxa. a programme to actively control these species is in place, but the efficacy of the programme in reducing the number of invasive plants needs to be monitored. future priorities apart from monitoring the o. ariadne colonies, a priority is to continue the search for other colonies of these species and potential re-introduction sites, guided by the predicted distribution of the species (armstrong 2010). if possible, grassland linkages of adequate width and quality (pryke & samways 2001, 2003) between colonies will need to be negotiated with landowners on the assumption that o. ariadne can disperse between colonies. although individuals observed away from the known colony areas may lend support to the latter assumption (pennington 1946), the assumption needs to be tested through genetic studies. the results of samways and lu (2007) indicate that male o. ariadne tend to remain in the colony area but that they are powerful fliers. the genetic distance between the nkandla population and the populations near howick needs to be established so that the sources of any re-introduced butterflies can be wisely chosen. the overall vision is to have thriving o. ariadne populations in various parts of the probable former distribution range of the species one day. conclusion top ↑ this study indicated that adaptive monitoring and management of threatened and endemic invertebrate species, with the cooperation of the owners, occupiers or managers of the land on which the species occur, may be crucial for the successful conservation of such species. examples such as that of the erikssonia edgei gardiner & terblanche 2010 (waterberg copper butterfly) exist, where the local extinction of the species may have been prevented with the implementation of landowner-assisted adaptive monitoring and management (henning et al. 2009). as land transformation continues in south africa (e.g. driver et al. 2012; jewitt 2012) and elsewhere in africa, such inclusive conservation efforts will be required for more invertebrate species. the need for more species-specific management protocols to ensure the long-term persistence of threatened habitat-specialist butterflies has been highlighted elsewhere (new et al. 2010; swengel et al. 2011). in order to determine the most appropriate habitat management protocols for threatened south african grassland-specialist butterfly species, studies of the life history and ecology of each (e.g. see lu & samways [2001, 2002a, 2002b] for o. ariadne and thomas [2005] for grassland species elsewhere), should be prioritised. the long-term conservation of these species will be more assured if adaptive management of their habitats in response to the information obtained through monitoring is effectively implemented, given the other requirements that may pertain for each species, such as the adequate spatial arrangement of sufficiently large colonies or populations within dispersal distance. acknowledgements top ↑ we gratefully thank the midlands meander association, and especially nikki brighton, for their continued sponsorship and encouragement of our research into the conservation of the karkloof blue butterfly, and especially for the bursaries that the association awarded to keenan steers and nonku mkhize to assist them with their university studies and the fieldwork reported on here. we thank hylton adie for sampling ants and for his report, dino santoro for permission to work on stirling farm, rose kunhardt for permission to work on wahroonga farm, and john craigie, sihle madonsela, callum beattie, george pizzey, liandra bertolli and caitlin thorpe for assistance in the field. all the monitors are thanked for their participation in the study. douglas macfarlane and the managers of the start property from sappi are thanked for inviting and allowing us to undertake the work there. several anonymous reviewers are thanked for their comments on previous versions of this manuscript. competing interests the authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article. authors’ contributions a.j.a. (ezemvelo kwazulu-natal wildlife) was the project leader, designed the study, carried out the fieldwork together with s.l.l. (ezemvelo kwazulu-natal wildlife), analysed the data and wrote the manuscript. s.l.l. organised and carried out the fieldwork at the nkandla monitoring site and helped capture the data, liaised with the amachube traditional authority, provided suggestions for the methodology, organised and guided the management of the nkandla monitoring site and commented on drafts of the article. references top ↑ armstrong, a.j., benn, g., bowland, a.e., goodman, p.s., johnson, d.n., maddock, a.h. et al., 1998, ‘plantation forestry in south africa and its impact on biodiversity’, south african forestry journal 182, 59–65. http://dx.doi.org/10.1080/10295925.1998.9631191armstrong, a.j., 2010, ‘box 9.1 the use of cartographic modelling to map potential habitat for threatened endemic species’, in m.j. samways, m.a. mcgeoch & t.r. new (eds.), insect conservation: a handbook of approaches and methods, pp. 290–292, oxford university press, oxford. pmid:19967358 armstrong, a.j. & edge, d.a., 2013, ‘orachrysops ariadne (butler, 1898a)’, in s. mecenero, j.b. ball, d.a. edge, m.l. hamer, g.a. henning, m. kruger et al. (eds.), conservation assessment of butterflies of south africa, lesotho and swaziland: red list and atlas, p. 542, saftronics (pty) ltd., johannesburg & animal demography unit, cape town. campbell, s.p., clark, j.a., crampton, l.h., guerry, a.d., hatch, l.t., hosseini, p.r. et al., 2002, ‘an assessment of monitoring efforts in endangered species recovery plans’, ecological applications 12, 674–681. http://dx.doi.org/10.1890/1051-0761(2002)012[0674:aaomei]2.0.co;2 dennis, r.l.h., shreeve, t.g. & van dyck, h., 2006, ‘habitats and resources: the need for a resource-based definition to conserve butterflies’, biodiversity and conservation 15, 1943–1966. http://dx.doi.org/10.1007/s10531-005-4314-3 driver, a., sink, k.j., nel, j.n., holness, s., van niekerk, l., daniels, f. et al., 2012, ‘national biodiversity assessment 2011: an assessment of south africa’s biodiversity and ecosystems’, synthesis report, south african national biodiversity institute and department of environmental affairs, pretoria. edge, d.a., cilliers, s.s. & terblanche, r.f., 2008, ‘vegetation associated with the occurrence of the brenton blue butterfly’, south african journal of science 104, 505–510. http://dx.doi.org/10.1590/s0038-23532008000600028 edge, d.a., woodhall, s.e., ball, j.b., henning, g.a., armstrong, a.j. & mecenero, s., 2013, ‘future priorities for butterfly conservation and research’, in s. mecenero, j.b. ball, d.a. edge, m.l. hamer, g.a. henning, m. kruger et al. (eds.), conservation assessment of butterflies of south africa, lesotho and swaziland: red list and atlas, pp. 35–40, saftronics (pty) ltd., johannesburg & animal demography unit, cape town. field, s.a., o’connor, p.j., tyre, a.j. & possingham, h.p., 2007, ‘making monitoring meaningful’, austral ecology 32, 485–491. http://dx.doi.org/10.1111/j.1442-9993.2007.01715.x greenwood, j.j.d. & robinson, r.a., 2006, ‘principles of sampling’, in w.j. sutherland (ed.), ecological census techniques, pp. 11–86, cambridge university press, cambridge. http://dx.doi.org/10.1017/cbo9780511790508.003 hamer, m., 2005, ‘report on the monitoring of karkloof blue butterfly populations in 2005’, inland invertebrate initiative, university of kwazulu-natal, pietermaritzburg. henning, g.a., terblanche, r.f. & ball, j.b. (eds.), 2009, south african red data book: butterflies, sanbi biodiversity series 13, south african national biodiversity institute, pretoria. hula, v., konvicka, m., pavlicko, a. & fric, z., 2004, ‘marsh fritillary (euphydryas arinia) in the czech republic: monitoring, metapopulation structure, and conservation of an endangered butterfly’, entomologica fennica 15, 231–241. jewitt, d., 2012, ‘land cover change in kwazulu-natal’, environment 10, 12–13. lindenmayer, d.b. & likens, g.e., 2009, ‘adaptive monitoring: a new paradigm for long-term research and monitoring’, trends in ecology and evolution 24, 482–486. http://dx.doi.org/10.1016/j.tree.2009.03.005, pmid:19409648 lindenmayer, d.b., likens, g.e., haywood, a. & miezis, l., 2011, ‘adaptive monitoring in the real world: proof of concept’, trends in ecology and evolution 26, 641–646. http://dx.doi.org/10.1016/j.tree.2011.08.002, pmid:21880394 louw, s. & armstrong, a.j., 2010, ‘box 11.5 community involvement in the conservation of threatened endemic invertebrate species’, in m.j. samways, m.a. mcgeoch & t.r. new (eds.), insect conservation: a handbook of approaches and methods, pp. 368–370, oxford university press, oxford. lu, s-s. & samways, m.j., 2001, ‘life history of the threatened karkloof blue butterfly, orachrysops ariadne (lepidoptera: lycaenidae)’, african entomology 9, 137–151. lu, s-s. & samways, m.j., 2002a, ‘behavioural ecology of the karkloof blue butterfly orachrysops ariadne (lepidoptera: lycaenidae) relevant to its conservation’, african entomology 10, 137–147. lu, s-s. & samways, m.j., 2002b, ‘conservation management recommendations for the karkloof blue butterfly, orachrysops ariadne (lepidoptera: lycaenidae)’, african entomology 10, 149–159. mcgeoch, m.a., sithole, h., samways, m.j., simaika, j.p., pryke, j.s., picker, m. et al., 2011, ‘conservation and monitoring of invertebrates in terrestrial protected areas’, koedoe 53, art. #1000, 13 pages. http://dx.doi.org/10.4102/koedoe.v53i2.1000 mucina, l. & rutherford, m.c. (eds.), 2006, the vegetation of south africa, lesotho and swaziland, strelitzia 19, south african national biodiversity institute, pretoria. murphy, d.d. & weiss, s.b., 1988, ‘a long-term monitoring plan for a threatened butterfly’, conservation biology 2, 367–374. http://dx.doi.org/10.1111/j.1523-1739.1988.tb00201.x new, t.r., yen, a.l., sands, d.p.a., greenslade, p., neville, p.j., york, a. et al., 2010, ‘planned fires and invertebrate conservation in south east australia’, journal of insect conservation 14, 567–574. http://dx.doi.org/10.1007/s10841-010-9284-4 pennington, k.m., 1946, ‘notes on some rare south african butterflies’, journal of the entomological society of south africa 9, 20–27. pfab, m.f., victor, j.e. & armstrong, a.j., 2011, ‘application of the iucn red listing system to setting species targets for conservation planning purposes’, biodiversity and conservation 20, 1001–1012. http://dx.doi.org/10.1007/s10531-011-0009-0 pickens, b.a. & root, k.v., 2009, ‘behavior as a tool for assessing a managed landscape: a case study of the karner blue butterfly’, landscape ecology 24, 243–251. http://dx.doi.org/10.1007/s10980-008-9302-z pierce, n.e., braby, m.f., heath, a., lohman, d.j., mathew, j., rand, d.b. et al., 2002, ‘the ecology and evolution of ant association in the lycaenidae (lepidoptera)’, annual review of entomology 47, 733–771. http://dx.doi.org/10.1146/annurev.ento.47.091201.145257, pmid:11729090 powell, a.f.l.a., busby, w.h. & kindscher, k., 2007, ‘status of the regal fritillary (speyeria idalia) and effects of fire management on its abundance in northeastern kansas, usa’, journal of insect conservation 11, 299–308. http://dx.doi.org/10.1007/s10841-006-9045-6 pringle, e.l.l., henning, g.a. & ball, j.b. (eds.), 1994, pennington’s butterflies of southern africa, struik, cape town. pryke, s.r. & samways, m.j., 2001, ‘width of grassland linkages for the conservation of butterflies in south african afforested areas’, biological conservation 101, 85–96. http://dx.doi.org/10.1016/s0006-3207(01)00042-8 pryke, s.r. & samways, m.j., 2003, ‘quality of remnant indigenous grassland linkages for adult butterflies (lepidoptera) in an afforested african landscape’, biodiversity and conservation 12, 1985–2004. http://dx.doi.org/10.1023/a:1024103527611 samways, m.j. & lu, s-s., 2007, ‘key traits in a threatened butterfly and its common sibling: implications for conservation’, biodiversity and conservation 16, 4095–4107. http://dx.doi.org/10.1007/s10531-007-9209-z samways, m.j., mcgeoch, m.a. & new, t.r. (eds.), 2010, insect conservation: a handbook of approaches and methods, techniques in ecology & conservation series, oxford university press, oxford. pmcid:pmc2847917 schtickzelle, n., turlure, c. & baguette, m., 2007, ‘grazing management impacts on the viability of the threatened bog fritillary butterfly proclossiana eunomia’, biological conservation 136, 651–660. http://dx.doi.org/10.1016/j.biocon.2007.01.012 schultz, c.b. & dlugosch, k.m., 1999, ‘nectar and hostplant scarcity limit populations of an endangered oregon butterfly’, oecologia 119, 231–238. http://dx.doi.org/10.1007/s004420050781 smee, m., smyth, w., tunmore, m., ffrench-constant, r. & hodgson, d., 2011, ‘butterflies on the brink: habitat requirements for declining populations of the marsh fritillary (euphydryas aurinia) in sw england’, journal of insect conservation 15, 153–163. http://dx.doi.org/10.1007/s10841-010-9334-y spatial data, 2013, biodiversity spatial planning and information division, ezemvelo kwazulu-natal wildlife, south africa. swengel, s.r., schlicht, d., olsen, f. & swengel, a.b., 2011, ‘declines of prairie butterflies in the midwestern usa’, journal of insect conservation 15, 327–339. http://dx.doi.org/10.1007/s10841-010-9323-1 thomas, j.a., 2005, ‘monitoring change in the abundance and distribution of insects using butterflies and other indicator groups’, philosophical transactions of the royal society b 360, 339–357. http://dx.doi.org/10.1098/rstb.2004.1585, pmid:15814349, pmcid:pmc1569450 tudor, o., dennis, r.l.h., greatorex-davis, j.n. & sparks, t.h., 2004, ‘flower preferences of woodland butterflies in the uk: nectaring specialists are species of conservation concern’, biological conservation 119, 397–403. http://dx.doi.org/10.1016/j.biocon.2004.01.002 watson, j., 2001, ‘how to determine a sample size: tipsheet #60’, penn state cooperative extension, university park, pa, viewed 14 may 2012, from http://www.extension.psu.edu/evaluation/pdf/ts60.pdf grobler.qxd natural woodland vegetation and plant species richness of the urban open spaces in gauteng, south africa c.h. grobler, g.j. bredenkamp and l.r. brown grobler, c.h., g.j. bredenkamp and l.r. brown. 2002. natural woodland vegetation and plant species richness of the urban open spaces in gauteng, south africa. koedoe 45(1): 19–34. pretoria. issn 0075-6458. it is estimated that approximately 60 % of the world’s population will be living in urban areas by 2025. in gauteng, the most densely populated province in south africa, the natural open spaces are continually under threat from development. vegetation is the most physical representation of the environment on which all animals are ultimately dependent. in order to evaluate an areas potential for development or conservation it is necessary to make a thorough inventory of the plant communities and their associated habitats. a survey of the natural woodlands was undertaken as part of a project describing the vegetation of the natural open spaces within the gauteng region. relevés were compiled in 73 stratified random sample plots in selected open spaces within the study area. a twinspan classification, refined by braun-blanquet procedures, indicated six woodland communities that can be grouped into two major woodland communities. a classification and description of these communities as well as their species richness are presented. the results indicate that there are still patches of well-preserved natural vegetation within the study area and contribute to the limited knowledge that presently exists for the vegetation of the area. key words: braun blanquet analysis, classification, plant communities, urban open space, twinspan. c.h. grobler and g.j. bredenkamp, department of botany, university of pretoria, pretoria, 0002 south africa; l.r. brown , applied natural sciences, technikon sa, private bag x6, florida, 1710 south africa. issn 0075-6458 19 koedoe 45/1 (2002) introduction although the smallest of the nine provinces of south africa, covering an area of only 16 191 km², gauteng has a population of approximately 7.3 million people making it the most densely populated province in south africa (gauteng 2000). it is therefore understandable that urbanisation and its associated impacts are the biggest threats to the natural areas that still persist in the gauteng urban areas. kowarik (1990) confirms this by stating that human impact has been recognised as one of the most important influences on the composition of vegetation in urban environments. these impacts often include the loss of complete habitats due to the construction of residential, industrial or other developments. natural areas adjacent to urban areas are equally affected by humanrelated activities such as trampling, footpaths and exotic plant invasions, including stray products from gardens. management practices such as mowing of natural grasslands and changing the natural veld fire frequency also have an effect on these natural ecosystems. in the past, little attention was paid to the natural environment when making decisions on development in gauteng. by overlooking the natural environment in urban development planning, wasteful exploitation of the natural resources occurred (bredenkamp & brown 1998a, 1998b). in european countries, landscape ecological mapping and evaluation grobler.qxd 2005/12/09 11:29 page 19 have become an essential part of the planning process (spellerberg 1992). such planning assists in restricting disturbance in areas with sensitive habitats or species and prevent undue fragmentation of wildlife habitat (helliwell 1973). until recently, vegetation surveys in urban areas in south africa were unknown. after a pioneer survey in the durban municipal area (roberts 1993), surveys were undertaken in the urban areas of potchefstroom (cilliers & bredenkamp 1998, 1999a, 1999b, 1999c, 2000; cilliers et. al. 1998), klerksdorp (van wyk et al. 1997) in the north-west province, and bloemfontein (dingaan 1999) in the free state. the natural areas of the gauteng province presently support a large number of plant species. limited vegetation studies, mostly unpublished, have been done on small areas in the urban gauteng (behr & bredenkamp 1988; ellery 1992, 1994; bredenkamp 1991, 1992, 1997a, 1997b; bredenkamp & brown 1998a, 1998b). surveys of similar natural woodland vegetation in the gauteng province outside urban areas, include those of bredenkamp & theron (1978, 1980) and coetzee et.al. (1993a, 1993b, 1994, 1995). the present study reports on a wider assessment of the vegetation of the urban areas of gauteng, attempting to include areas of high species richness in urban areas and to consolidate the studies previously done in the study area. the purpose of this study was therefore to identify the plant communities of different types of habitat present in the urban areas of gauteng. this information will help to motivate conservation actions, ensuring that nature conservation strategies are incorporated into land-use planning initiatives within the urban environment. study area the study area is located between 27º40'e– 28º25'e and 25º40's–26º20's approximately in the center of the gauteng province (fig. 1) and includes pretoria, midrand, johannesburg and parts of the west rand. altitudes in gauteng vary from 1081 m to 1899 m above sea level, with a mean altitude of 1512 m (gauteng 2000). mean monthly temperature in the study area is 16.8 ºc with a mean maximum of 22.6 ºc and a mean minimum of 10.8 ºc. the mean winter temperature in the study area is 13.8 ºc and mean summer temperature, 25.6 ºc (weather bureau 2000). mean annual rainfall in gauteng is 670 mm per year (gauteng 1997). the geology of the area includes the rock types dolomite, chert, quartzite, granite, diabase, shale and andesitic lava. the most important land type units in this study area are ab, bb, ba that are mainly associated with with flat or slightly undulating landscapes and the ib land type which is often associated with ridge areas. the main soil forms found in the study area are mispah and glenrosa. (land type survey staff 1985, 1987a, 1987b). acocks (1988) described the vegetation of the study area as “false” grassland of the central variation of bankenveld (veld type 61b). patches of woodland vegetation are found at sheltered sites on hillslopes and rocky outcrops within this veld type. bredenkamp & van rooyen (1996a) described this vegetation as the rocky highveld grassland vegetation type, which covers the largest part of the study area. a small patch of moist cool highveld grassland (bredenkamp & van rooyen 1996b) occurs in the southeastern corner of the study area. the area, therefore, predominantly represents the grassland biome (rutherford & westfall 1986). the open and closed woodland patches in the grassland biome resemble the vegetation of the waterberg moist mountain bushveld (bredenkamp & van rooyen 1996b) also described by acocks (1988) as sour bushveld (20). the woodland layer of the area is further represented by the mixed bushveld vegetation type (bredenkamp & van rooyen 1996a) representing the savanna biome. currently, only 3.05 % of mixed bushveld and 8.55 % of the waterberg moist mountain bushveld is conserved (bredenkamp & van rooyen 1996a, 1996c). koedoe 45/1 (2002) 20 issn 0075-6458 grobler.qxd 2005/12/09 11:29 page 20 methods the open spaces within the study area were identified and delineated using 1:50 000 aerial photographs and were stratified into relatively homogeneous areas within which sample plots were randomly allocated. a total of 73 sample plots were randomly located within the woodland vegetation to ensure that all variations were considered and sampled. plot sizes of 200 m² were used in accordance with bredenkamp & theron (1978). total floristic composition was noted for every sample plot. additionally, the average height and percentage cover of the tree, shrub and herbaceous layers were estimated and recorded. the percentage cover of bare soil was also estimated and recorded. the coverage of the tree layer was based on crown cover where the canopies are vertically projected onto the ground (werger 1974) the same technique was used for the shrub and herbaceous layers. the dominant tree, shrub, and herbaceous species were noted and the cover/abundance for each species was estimated according to the braun blanquet coverabundance scale (mueller-dombois & ellenberg 1974). environmental data included slope inclination measured in degrees, topography including hills, slopes and crests, aspect (north, south, west, east, and aspects in between) and disturbance factors such as exotic vegetation, trampling and erosion. information on geology, land type, soil forms and clay contents were obtained from land type survey staff (1985, 1987). soil classification is according to macvicar (1991). the computer programme turboveg (hennekens 1996a) and the twinspan classification algorithm (hill 1979) were used for capture, processing and presentation of phytosociological data. further refinement was achieved with braun blanquet procedures by using megatab (hennekens 1996b), a visual editor for phytosociological tables. names and authors of taxa are in accordance with arnold & de wet (1993). for the purposes of this study, aloe greatheadii var. davyana and aloe transvaalensis were lumped as one species and referred to as aloe greatheadii var. davyana. the classification of species growth forms were done according to van wyk & malan (1998). issn 0075-6458 21 koedoe 45/1 (2002) fig. 1. location of the study area within the gauteng province. grobler.qxd 2005/12/09 11:29 page 21 results a mixture of tree, shrub and herbaceous species characterises the woodland areas of the entire study area. general species present in the woodlands are listed under species group k (table 1). the vegetation of all the woodland communities is characterised by the presence of the small trees diospyros lycioides, canthium gilfillanii, zanthoxylum capensis, rhus leptodictya, maytenus heterophylla and rothmannia capensis together with the exotics lantana camara and solanum mauritianum. the small shrub asparagus suaveolens, the forbs commelina africana, helichrysum rugulosum, the succulent aloe greatheadii var. davyana, the slender twining herb pentarrhinum insipidum, the exotic weeds tagetes minuta, bidens pilosa, the noxious parasite cuscuta campestris and the fern pellaea calomelanos are often encountered. common grasses include melinis repens, melinis nerviglumis, themeda triandra, setaria sphacelata, cymbopogon validus, panicum maximum, hyparrhenia hirta, eragrostis curvula, brachiaria serrata, heteropogon contortus and digitaria eriantha. these species are listed in species group k (table 1) and will therefore not be repeated in the description of the various plant communities. hills and ridges in the study area are often covered by a mosaic of open woodland on the warmer and drier north-facing slopes, closed woodland in more sheltered sites below rocky cliffs, and grassland patches on more exposed, high altitude areas. closed woodland communities are also found along rivers and streams. cooler south-facing slopes may have protea caffra woodland communities. two major plant communities were identified namely the acacia caffra-rhus pyroides open to closed major woodland community, mostly found on cooler and more moist, south facing, or exposed habitats and the combretum molle-setaria lindenbergiana open to closed major woodland community, mostly associated with warmer and drier north-facing slopes. the classification obtained from the twinspan and refined by braun-blanquet procedures resulted in the following six plant communities that can be grouped into two major communities being identified in the woodland areas surveyed. 1. acacia caffra – rhus pyroides open to closed major woodland community 1.1 hypoestes forskaolii – acacia caffra closed woodland community 1.2 acacia caffra – setaria lindenbergiana closed woodland community 1.3 acacia caffra – themeda triandra open to closed woodland community 1.3.1 acacia caffra – themeda triandra – asparagus laricinus open to closed woodland sub-community 1.3.2 acacia caffra – themeda triandra – hyparrhenia hirta open to closed woodland sub-community 2. combretum molle–setaria lindenbergiana open to closed major woodland community 2.1 combretum molle – burkea africana open to closed woodland community 2.2 combretum molle – englerophytum magalismontanum open woodland community discussion floristically the acacia caffra-rhus pyroides open to closed woodland shows strong floristic relationships with the sub-humid cool temperate mountain bushveld of coetzee et al. (1995), and also exhibits an affinity to the afro-montane vegetation of the drakensberg. the plant species composition of the combretum molle-setaria lindenbergiana open to closed woodland relates to the sub-humid warm temperate bushveld as described by coetzee et al. (1993b), and shows an affinity to the bushveld vegetation koedoe 45/1 (2002) 22 issn 0075-6458 grobler.qxd 2005/12/09 11:29 page 22 of the savanna biome (rutherford & westfall 1986). bredenkamp & brown (1998a, 1998b) also described vegetation to the west and north of johannesburg indicating floristic resemblance to this vegetation unit. the vegetation of the ruimsig entomological reserve in roodepoort, also shows a floristical resemblance to the woodland species identified in this study (deutschländer & bredenkamp 1999). the common species associated with the rhus pyroides forest and canthium gilfillanii-aristida transvaalensis-cymbopogon marginatus savanna communities identified by bredenkamp & theron (1978), and the euclea crispa-rhoicissus tridentata bush and savanna communities as well as the trachypogon spicatus-themeda triandra grassland communities bredenkamp & theron (1980) compare well with the common species identified in the woodland communities for this study. vegetation description 1. acacia caffra – rhus pyroides open to closed major woodland community habitat this plant community is found scattered throughout the study area, among granite boulders, on rocky outcrops and hills. slopes varies from gradual to very steep. a total of 54 % of the sample plots were on the bb, and ib land types. this plant community occurs on rocky areas with shallow soils and is associated with the glenrosa , mispah and wasbank soil forms, with a clay content of 10–30 %. vegetation species group a (table 1) characterises this community. the diagnostic species are the trees acacia caffra, celtis africana, rhus pyroides, ehretia rigida, dombeya rotundifolia, ziziphus mucronata, cussonia paniculata, heteromorpha trifoliata and euclea crispa, the geophytes ledebouria revoluta and scadoxus puniceus, the exotic perennial weed achyranthes aspera, and the annual weed conyza albida. the species that dominate this plant community are the trees rhus leptodictya, celtis africana, acacia caffra, canthium gilfillanii and zanthoxylum capense, the shrub diospyros lycioides and small shrub asparagus suaveolens, the succulent aloe greatheadii var. davyana, the common weed tagetes minuta, the asteraceous forb helichrysum rugulosum, and the grasses themeda triandra, hyparrhenia hirta and setaria sphacelata. the high occurrence of the weed tagetes minuta as well as the other exotic weedy species verbena bonariensis, conyza albida, lantana camara, zinnia peruviana and the succulent opuntia ficus-indica in the herbaceous layer together with melia azedarach in the shrub and tree layer, indicates that this plant community is susceptible to disturbance and generally not in a good condition in the study area. an average of 37 species per 200 m² was recorded in this plant community. in general, the species composition of this community compares well with that of the acacia caffra – euclea crispa woodland within the sub-humid cool temperate mountain bushveld described by coetzee et al. (1994, 1995), found on north-facing slopes, crests and upper south-facing steep slopes of the rocky outcrops mainly within the ib land type in the pretoria area. this community also shows floristic resemblance with the acacia caffra – setaria spacelata closed woodland described by coetzee et al. (1995), the acacia caffra – setaria sphacelata closed woodland in the roodeplaatdam nature reserve (van rooyen 1984), the vegetation on the bronberg in pretoria (bredenkamp 1997), the olea europaea – cymbopogon excavatus savanna and olea europeae schistotephium heptalobium bush clumps vegetation of the rietfontein nature reserve (bredenkamp 1991), the acacia robusta woodland (ellery 1994) and the acacia caffra vegetation of the issn 0075-6458 23 koedoe 45/1 (2002) grobler.qxd 2005/12/09 11:29 page 23 koedoe 45/1 (2002) 24 issn 0075-6458 ta bl e 1 a ph yt os oc io lo gi ca l t ab le o f t he w oo dl an d ve ge ta tio n of th e ur ba n ar ea s of g au te ng grobler.qxd 2005/12/09 11:29 page 24 issn 0075-6458 25 koedoe 45/1 (2002) ta bl e 1 (c on tin ue d) grobler.qxd 2005/12/09 11:29 page 25 koedoe 45/1 (2002) 26 issn 0075-6458 ta bl e 1 (c on tin ue d) grobler.qxd 2005/12/09 11:30 page 26 issn 0075-6458 27 koedoe 45/1 (2002) ta bl e 1 (c on tin ue d) grobler.qxd 2005/12/09 11:30 page 27 fort klapperkop area in pretoria (bredenkamp 1997b). the kloof communities: rhus pyroides forests, rhus pyroides-buddleja salviifolia-celtis africana forest and dombeya rotundifolia-ziziphus mucronata savanna identified by bredenkamp & theron (1978) also show resemblance to this community. bredenkamp & brown (1998a, 1998b) described similar vegetation to the west and north of johannesburg. three communities were recognised under this major woodland community: 1.1 hypoestes forskaolii – acacia caffra closed woodland community habitat this woodland community is found on gradual to moderately steep slopes and a variety of aspects of hills, ridges and between granite boulders as well as along rivers in lower lying areas in the pretoria, midrand, sandton and randburg areas. a 10-15 % clay content was observed in the soils associated with this plant community. this plant community was predominantly found in the bb land type. vegetation this community is characterised by the presence of the trees combretum erythrophyllum and rhamnus prinoides (along rivers), acalypha glabrata, pavetta gardeniifolia var. gardeniifolia and buddleja saligna, the shrub asparagus sp. and the shrublet aerva leucura, the herb hypoestes forskaolii, the erect cyperus esculentus the shrubby perennial herb hibiscus calyphyllus, the exotic climber araujia sericifera and the grass setaria megaphylla (species group b table 1) prominent species in this plant community include the trees rhus pyroides, celtis africana, acacia caffra, euclea crispa, canthium gilfillanii, rhus leptodictya, zanthoxylum capense, the shrub diospyros lycioides, the small shrub asparagus suaveolens, and the weeds achyranthes aspera and tagetes minuta. the average number of species recorded in this plant community per 200 m² is 32. showing resemblance to this community is the combretum erythrophyllum celtis africana major community described by behr & bredenkamp (1988) from the witwatersrand botanic garden in roodepoort and the celtis africana – ziziphus mucronata community (bredenkamp 1997b) from the fort klapperkop area in pretoria and the lonehill nature reserve (bredenkamp 1992). bredenkamp (1992) conducted a more detailed assessment on the small lonehill nature reserve and described more communities. bredenkamp & brown (1998a, 1998b) described vegetation to the west and north of johannesburg indicating resemblance to this community. 1.2 acacia caffra – setaria lindenbergiana closed woodland community habitat this closed woodland community has a limited distribution in the study area. it was observed on steep to very steep, north and south-facing slopes of hills and ridges throughout the study area except in the midrand-fourways area. it is generally associated with the land type ib. vegetation the species group c (table 1) is diagnostic for this plant community. the diagnostic species are the grass setaria lindenbergiana and the climber shrub rhoicissus tridentata. the most prominent species in this plant community include the trees acacia caffra, celtis africana and euclea crispa, the shrub diospyros lycioides and the bulbous geophyte ledebouria revoluta, the noxious weed tagetes minuta and the succulent aloe greatheadii var. davyana. disturbance to this plant community was recorded at most of the sites, with specific reference to the low herbaceous cover under the trees. an average of 31 species per 200 m² was recorded for this plant community. koedoe 45/1 (2002) 28 issn 0075-6458 grobler.qxd 2005/12/09 11:30 page 28 this community is related to the setaria lindenbergiana – ehretia rigida bush clumps (coetzee et al. 1995), the setaria lindenbergiana – acacia caffra woodland described by coetzee (1975) from the rustenburg nature reserve, and the olea europaea – cymbopogon excavatus savanna and olea europaea schistotephium heptalobium bush clumps vegetation of the rietfontein nature reserve (bredenkamp 1991). 1.3 acacia caffra – themeda triandra open to closed woodland community habitat this open to closed woodland sub-community was found distributed throughout the study area on a variety of aspects, on gradual to moderately steep slopes of hills and ridges. it was recorded predominantly on land type ib, characterised by mispah and glenrosa soil forms with a clay content of 10–20 %. vegetation the diagnostic species for the community are the small tree rhus zeyheri, the grasses eragrostis chloromelas, aristida congesta subsp. barbicollis, the geophyte hypoxis rigidula, the forbs nidorella hottentotica, graderia subintegra, pearsonia sessilifolia subsp. sessilifolia, justicia anagalloides, crabbea angustifolia and scabiosa columbaria, the small shrub lantana rugosa and the exotic shrub solanum elaeagnifolium (species group d – table 1). the woody layer is dominated by trees rhus pyroides, canthium gilfillanii, rhus leptodictya, and the shrub diospyros lycioides, together with the shrublet asparagus suaveolens. the succulent aloe greatheadii var. davyana, the asteraceous helichrysum rugulosum, the weed tagetes minuta together with the grasses themeda triandra, hyparrhenia hirta, brachiaria serrata and setaria sphacelata are also prominent. the average number of species recorded in this plant community per 200 m² is 41. in parts of the plant community stands of protea caffra (species group f) and the scarcer protea roupelliae (species group f) are prominent, indicating an affinity with the drakensberg highveld sourveld (sourveld) vegetation. the olea europaea – cymbopogon excavatus savanna and olea europeae schistotephium heptalobium bush clumps vegetation of the rietfontein nature reserve (bredenkamp 1991) also show resemblance to this community. two sub-communities are recognised: 1.3.1 acacia caffra – themeda triandra – asparagus laricinus closed woodland sub-community habitat this sub-community is found on gradual north and south-facing slopes predominantly in the pretoria area, but also in clayville, fourways and alberton. this plant community is associated with closed woodland in association with rocky outcrops. this plant community was recorded from the ib1b land types. vegetation species that dominate the plant community are the trees acacia caffra, celtis africana and ehretia rigida, the shrublet asparagus suaveolens, the grasses themeda triandra, setaria sphacelata, the succulent aloe greatheadii var. davyana and the asteraceous helichrysum rugulosum. rhus pyroides is locally dominant and panicum maximum is sometimes prominent. species group e (table 1) is diagnostic for this community. diagnostic tree species include acacia karroo, the shrub asparagus laricinus that forms impenetrable thickets, the slender shrub sida dregei, the erect shrublets hibiscus lunarifolius and teucrium trifidum, the herbaceous shrublet pavonia burchellii, the twining forb clematis brachiata, the forb conyza podocephala and the exotic weeds oxalis corniculata and conyza albida. issn 0075-6458 29 koedoe 45/1 (2002) grobler.qxd 2005/12/09 11:30 page 29 the average number of species recorded in this plant community per 200 m² is 36. the acacia karroo – lippia javanica closed woodland (coetzee et al. 1995) shows resemblance to this community. 1.3.2 acacia caffra – themeda triandra – hyparrhenia hirta open to closed woodland sub-community habitat this sub-community is found throughout the study area on a variety of aspects and on gradual to steep slopes of ridges and hills and rocky outcrops. this plant community is prominent on the higher altitude slopes and crests. vegetation species group f (table 1) is diagnostic for this vegetation. the diagnostic species are the trees kiggelaria africana, protea caffra, protea roupelliae and rhus pyroides, the shrublets athrixia elata, pentanisia angustifolia, rhus discolor, and shrub lippia javanica, the forbs chaetacanthus costatus, vernonia galpinii and rhynchosia totta, and the grasses tristachya leucothrix, eragrostis racemosa, cynodon dactylon, elionurus muticus and alloteropsis semialata subsp. eckloniana. the plant community is dominated by the tree species rhus leptodictya and canthium gilfillanii the shrub diospyros lycioides, the succulent aloe greatheadii var. davyana, the asteraceous helichrysum rugulosum and the grasses setaria sphacelata, melinis repens, melinis nerviglumis, themeda triandra, hyparrhenia hirta and brachiaria serrata. an average of 44 species per 200 m² was recorded in this plant community. this community resembles the protea caffra – athrixia elata open woodland (coetzee 1995) from the pretoria area. the common species of the tracypogon spicatus-themeda triandra grassland of bredenkamp & thereon (1980) compare well with this community. bredenkamp & brown (1998a, 1998b) described vegetation to the west and north of johannesburg indicating resemblance to this vegetation unit. 2. combretum molle–setaria lindenbergiana open to closed major woodland community habitat this plant community is mostly found on warmer and drier north-facing aspect on gradual to very steep slopes in kloofs, between granite boulders and on rocky outcrops, drainage lines, at the foot of quartzite ridges and hills and on rocky outcrops in the pretoria and johannesburg area. this plant community is mainly found in the ib and ba land types. vegetation the diagnostic species of this plant community are the trees combretum molle, vangueria infausta, ochna pulchra, combretum zeyheri, nuxia congesta, lannea discolor and englerophytum magalismontanum. other species include the shrub ochna pretoriensis, the shrublet indigofera melanadenia, and the succulents kalanchoe paniculata, crassula swaziensis and euphorbia schinzii, forbs such as the tufted coleochloa setifera, commelina erecta, oldenlandia herbacea var. herbacea and grasses such as setaria lindenbergiana and loudetia simplex (species group g; table 1). the species that dominate this plant community are the succulent aloe greatheadii var. davyana, the weed tagetes minuta, the herb commelina africana and the scrubby xerophyta retinervis, the fern pellaea calomelanos, and the grasses cymbopogon validus, melinis repens, melinis nerviglumis and diheteropogon amplectens. an average of 38 species per 200 m² was recorded in this plant community. invasion of exotic vegetation was often recorded in this plant community. invasive exotic species include lantana camara, cereus peruviana and jacaranda mimosifolia. koedoe 45/1 (2002) 30 issn 0075-6458 grobler.qxd 2005/12/09 11:30 page 30 in general, this community compares well with the sub-humid warm temperate mountain bushveld found on north-facing slopes, crests, and upper south-facing steep slopes of the rocky outcrops mainly within the ib land type (coetzee et al. 1995). also resembling this community are the burkea africana–bequartiodendron magalismontanum open woodland (coetzee 1994), setaria lindenbergiana-combretum molle community (bezuidenhout 1993), sporobolus pectinatus–chaetacanthus setiger community (coetzee 1993), canthium gilfillanii– combertum molle savanna community from the witwatersrand national botanic garden (behr & bredenkamp 1988), nuxia congesta-combretum molle savanna (bezuidenhout et al. 1988) vegetation on the bronberg in pretoria (bredenkamp 1997a), the cryptolepis oblongifolia–englerophytum, burkea africana–combretum molle and brachylaena rotundata–burkea africana communities from the fort klapperkop area (bredenkamp 1997b). two sub-communities were recognised: 2.1 combretum molle – burkea africana open to closed woodland community habitat this sub-community is predominantly found on shallow to moderately steep northern to north-eastern slopes of ridges and hills in the pretoria area. it was mainly recorded in 1b land type. vegetation species group h (table 1) is diagnostic for this sub-community. diagnostic species include the trees burkea africana and strychnos pungens and the dwarf shrubs rhynchosia nitens and helichrysum kraussii and geoxylic suffrutescent dichapetalum cymosum. the most prominent species in this plant community are the trees combretum molle and vangueria infuasta, the shrubs xerophyta retinervis, the succulent aloe greatheadii var. davyana, the fern pellaea calomelanos and the grass melinis nerviglumis. the average number of species recorded in this plant community is 40 per 200 m². 2.2 combretum molle – englerophytum magalismontanum open woodland community habitat this sub-community was found at high altitudes on northern aspects with gradual to very steep (55º and 50º at bush hill in randburg and on northcliff hill in northcliff) slopes, on top of ridges and hills, and in drainage lines on ridges in the pretoria and johannesburg areas. the plant community was recorded specifically in land types ba and ib. the soil form is mainly mispah with a clay content of 10–20 % also indicating shallow soils on rock. vegetation this sub-community is characterised by species group i (table 1). characteristic species associated with this plant community are the trees pittosporum viridiflorum, croton gratissimus, osyris lanceolata and pouzolzia mixta and the shrubs ancylobotrys capensis and rhus magalismontana and the sbrublet waltheria indica. other species include forbs like the succulent crassula setulosa, vernonia sutherlandii, cyperus sp. and sutera caerulea and the grasses aristida transvaalensis and enneapogon scoparius. species that dominate the plant community are the trees combretum molle, vangueria infausta, canthium gilfillanii and diospyros lycioides the grasses cymbopogon validus, melinis repens and the forb commelina africana. the average number of species recorded in this plant community is 36 per 200m². floristically the bequartiodendron magalismontanum shrub land of ellery (1994) also compares well with this community. issn 0075-6458 31 koedoe 45/1 (2002) grobler.qxd 2005/12/09 11:30 page 31 discussion the results of this study indicate that patches of well-preserved vegetation do occur in urban gauteng. the sites chosen were relatively undisturbed and many could be important to conserve. some sites are presently utilised for agricultural purposes. in spite of this, weedy species, eg., tagetes minuta and bidens pilosa occur on a large number of plots indicating disturbances of the herbaceous layer. species group j (table 1) indicates a floristic relationship between the acacia caffra–themeda triandra–hyparrhenia hirta open to closed woodland and the combretum molle–setaria lindenbergiana open to closed woodland. both these communities occur on slopes of fairly high altitude. the classification procedures followed, resulted in clearly defined vegetation units that can be related to the environmental factors observed. this classification contributes to the limited knowledge that presently exists for the vegetation of urban open spaces in gauteng. the results of this study can be used as a baseline study and future studies can be measured against it. the species richness determined per 200 m² per vegetation type could also be used to determine environmental and ecological degradation and actions could be taken to manage these changes. the results of this study could further be used to determine how the surrounding landuses have influenced the vegetation of urban open spaces. the results could assist in guiding development in future open spaces in a constantly expanding urban environment. acknowledgements the research was funded by the gauteng department of agriculture, conservation and environment. references acocks, j.p.h. 1988. veld types of south africa, 3rd ed. memoirs of the botanical survey of south africa 57: 1–146. arnold, t.h. & b.c. de wet. 1993. plants of southern africa. names and distribution. memoirs of the botanical survey of south africa 62: 1–825. behr, c.m. & g.j. bredenkamp. 1988. a phytosociological classification of the vegetation of the witwatersrand national botanical garden. south african journal of botany 54: 525–533. bezuidenhout, h. 1993. syntaxonomy and synecology of western transvaal grasslands, south africa. ph.d. thesis. university of pretoria, pretoria. bezuidenhout, h., g.j. bredenkamp & j.h. elsenbroek. 1988. die plantegroei van die alkaligraniet en aangrensende kwartsiet in die vredefortkoepel noordwes van parys. suid afrikaanse tydskrif vir natuurwetenskap en tegnologie 7: 4–9. bredenkamp, g.j. 1991. die plantegroei van die rietfontein natuureservaat. unpublished report, sandton municipality. bredenkamp, g.j. 1992. the vegetation of the lonehill nature reserve, sandton. unpublished report, sandton municipality. bredenkamp, g.j. 1997a. a vegetation assessment of a proposed residential development on the willows 340 jr portion 234. pretoria: van riet & louw landscape architects. bredenkamp, g.j. 1997b. a vegetation assessment of the fort klapperkop area. pretoria: van riet & louw landscape architects. bredenkamp, g.j.& l.r. brown. 1998a. a vegetation assessment of open spaces of the western metropolitan local council area. report for the western metropolitan local council. bredenkamp, g.j. & l.r. brown. 1998b. a vegetation assessment of the northern areas of the northern metropolitan local council. report for the northern metropolitan local council. bredenkamp, g.j. & g.k. theron. 1978. a sinecological account of the suikerbosrand nature reserve. i: the phytosiciology of the witwatersrant geological system. bothalia 12: 513–529. bredenkamp, g.j. & g.k. theron. 1980. a synecological account of the suikerbosrand nature reserve 11: the phytosociology of the ventersdorp geological system. bothalia 13(192): 199–216. bredenkamp, g.j. & n. van rooyen. 1996a. mixed bushveld. p. 26. in: low, a.b. & a.g. rebelo. (eds). vegetation of south africa, lesotho and swaziland. pretoria: dept. of environmental affairs and tourism. bredenkamp, g.j. & n. van rooyen. 1996b. moist cool highveld grassland. p. 43. in: low, a.b. & koedoe 45/1 (2002) 32 issn 0075-6458 grobler.qxd 2005/12/09 11:30 page 32 a.g. rebelo. (eds). vegetation of south africa, lesotho and swaziland. pretoria: dept. of environmental affairs and tourism. bredenkamp, g.j. & n. van rooyen. 1996c. waterberg moist mountain bushveld. p. 22. in: low, a.b. & a.g. rebelo. (eds). vegetation of south africa, lesotho and swaziland. pretoria: dept. of environmental affairs and tourism. cilliers, s.s. & g.j. bredenkamp. 1998. vegetation of railway reserves in the potchefstroom municipal area, north west province, south africa. south african journal of botany. 64(5): 271–280. cilliers, s.s. & g.j. bredenkamp. 1999a. urban nature conservation: vegetation of natural areas in the potchefstroom municipal area, north west province, south africa. koedoe. 42(1): 1–30. cilliers, s.s. & g.j. bredenkamp. 1999b. analysis of the spontaneous vegetation of intensely managed urban open spaces in the potchefstroom municipal area, north west province, south africa. south african journal of botany 65(1): 59–68. cilliers, s.s. & g.j. bredenkamp. 1999c. ruderal and degraded natural vegetation on vacant lots in the potchefstroom municipal area, north west province, south africa. south african journal of botany 65(2) 163–173. cilliers, s.s. & g.j. bredenkamp. 2000. vegetation on road verges on an urbanisation gradient in potchefstroom, south africa. landscape and urban planning 46: 217–239. cilliers, s.s., l.l. schoeman & g.j. bredenkamp. 1998. wetland plant communities in the potchefstroom municipal area, north west, south africa. bothalia 28(2): 213–229. coetzee, b.j. 1975. a phytosociological classification of the rustenberg nature reserve. bothalia 11: 561–580. coetzee, j.p., g.j. bredenkamp & n. van rooyen. 1993a. the plant communities of the sub-humid cool temperate mountain bushveld within the pretoria-wtbank-heidelberg area, south africa. south african journal of botany 61(3): 114–122. coetzee, j.p., g.j. bredenkamp & n. van rooyen. 1993b. the sub-humid warm temperate mountain bushveld plant communities of the pretoriawitbank-heidelberg area. south african journal of botany. 59(6): 623–632. coetzee, j.p., g.j. bredenkamp, n. van rooyen & g.k. theron. 1994. an overview of the physical environment and plant communities of the ba and ib land types of the pretoria-witbank-heidelberg area. south african journal of botany 60: 49–60. coetzee, j.p., g.j. bredenkamp & n. van rooyen. 1995. the phytosociology of the grasslands of the ba and ib land types in the pretoria-witbank-heidelberg area. south african journal of botany 61(3): 123–133. deutschländer, m.s. & g.j. bredenkamp. 1999. importance of vegetation analysis in the conservation management of the endangered butterfly aloeides dentatis dentatis (swierstra) (lepidoptera, lycaenidae). koedoe 42(2): 1–11. dingaan, 1999. the phytosociology of the natural open spaces in bloemfontein, free state. m.sc. thesis. university of the orange free state, bloemfontein. ellery, w.n. 1992. the vegetation ecology and conservation value of the western extension of melville koppie nature reserve, and proposals for its management. johannesburg: botanical society of south africa. ellery, w.n. 1994. the vegetation ecology of melville koppies nature reserve and louw geldenhuys view site: proposals for their management. johannesburg: botanical society of south africa. gauteng. 1997. state of the environment in gauteng: a preliminary report. gauteng department of agriculture, conservation and environment. unpublished report, csir environmentek. gauteng. 2000. spatial development framework. gauteng provincial government: development planning and local government. unpublished report, aps planafrica in association with labat – anderson africa, johannesburg. helliwell. d.r. 1973. priorities and values in nature conservation. journal of environmental management 1: 85–127 hennekens, s.m. 1996a. turboveg: software package for input, processing and presentation of phytosociological data. university of lancaster: ibn-dlo. hennekens, s.m. 1996b. megatab: a visual editor for phytosociological tables. ulft: giesen. hill, m.o. 1979. twinspan – a fortran program for arranging multivariate data in an ordered twoway table by classification of the individuals and attributes. new york: cornell university, ithaca. kowarik, i. 1990. some responses to flora on vegetation to urbanisation in central europe. pp. 45–74. in: sukopp, h. & s. hejny (eds.). urban ecology. the hague: spb academic publishing. land type survey staff. 1985. land types of the maps 2628 east rand. 2630 mbabane. memoirs on the agricultural natural resources of south africa 4: 1–261. land type survey staff. 1987a. land types of the maps 2526 rustenburg, 2528 pretoria. memoirs on the agricultural natural resources of south africa 8: 1–391. land type survey staff. 1987b. land types of the maps 2626 wes-rand, 2726 kroonstad. memoirs on the agricultural natural resources of south africa 4: 1–342. issn 0075-6458 33 koedoe 45/1 (2002) grobler.qxd 2005/12/09 11:30 page 33 macvicar, c.n. 1991. soil classification – a taxonomic system for south africa. pretoria: department of agricultural development. mueller-dombois, d. & h. ellenberg. 1974. aims and methods of vegetation ecology. new york: wiley. roberts, d.c. 1993. the vegetation of the municipal durban, natal. florist classification. bothalia 23(2): 271–326. spellerberg, i. 1992. evaluation and assessment for conservation. london: chapman & hall. van rooyen, n. 1984. ‘n fenologiese studie van die plantegroei van die roodeplaatdam-natuurreservaat. d.sc. –thesis. universiteit van pretoria. van wyk, e., s.s. cilliers & g.j. bredenkamp. 1997. ranteveldplantegroei van klerksdorp, noord-wes provinsie, suid-afrika. suidafrikaanse tydskrif vir natuurwetenskap en tegnologie 16(2): 74–85. van wyk, b. & s. malan. 1998. field guide to the wild flowers of the highveld. cape town: struik. rutherford, m.c. & r.h. westfall. 1986. the biomes of southern africa – an objective categorisation. memoirs of the botanical survey of south africa 51: 1–98. weather bureau. 2000. prints of the climate statistics of the johannesburg leeukop, pretoria forum wb, johannesburg joubert park and krugersdorp kroningspark weather stations. pretoria: department of environment affairs and tourism. werger, m.j.a. 1974. on concepts and techniques applied in the zurich-montpellier method of vegetation survey. bothalia 11: 309–323. koedoe 45/1 (2002) 34 issn 0075-6458 grobler.qxd 2005/12/09 11:30 page 34 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <feff00560065007200770065006e00640065006e0020005300690065002000640069006500730065002000450069006e007300740065006c006c0075006e00670065006e0020007a0075006d002000450072007300740065006c006c0065006e00200076006f006e0020005000440046002d0044006f006b0075006d0065006e00740065006e0020006d00690074002000650069006e006500720020006800f60068006500720065006e002000420069006c0064006100750066006c00f600730075006e0067002c00200075006d002000650069006e00650020007100750061006c00690074006100740069007600200068006f006300680077006500720074006900670065002000410075007300670061006200650020006600fc0072002000640069006500200044007200750063006b0076006f0072007300740075006600650020007a0075002000650072007a00690065006c0065006e002e00200044006900650020005000440046002d0044006f006b0075006d0065006e007400650020006b00f6006e006e0065006e0020006d006900740020004100630072006f0062006100740020006f0064006500720020006d00690074002000640065006d002000520065006100640065007200200035002e003000200075006e00640020006800f600680065007200200067006500f600660066006e00650074002000770065007200640065006e002e00200042006500690020006400690065007300650072002000450069006e007300740065006c006c0075006e00670020006900730074002000650069006e00650020005300630068007200690066007400650069006e00620065007400740075006e00670020006500720066006f0072006400650072006c006900630068002e> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice abstract introduction research method and design results and discussion conclusion acknowledgements references about the author(s) edward s. riddell centre for water resources research, university of kwazulu-natal, south africa conservation management, south african national parks, kruger national park, south africa werner kilian etosha ecological institute, okaukeujo, etosha national park, namibia wilferd versfeld etosha ecological institute, okaukeujo, etosha national park, namibia martin kosoana etosha ecological institute, okaukeujo, etosha national park, namibia citation riddell, e.s., kilian, w., versfeld, w. & kosoana, m., 2016, ‘groundwater stable isotope profile of the etosha national park, namibia’, koedoe 58(1), a1329. http://dx.doi.org/10.4102/koedoe.v58i1.1329 short communication groundwater stable isotope profile of the etosha national park, namibia edward s. riddell, werner kilian, wilferd versfeld, martin kosoana received: 21 july 2015; accepted: 23 may 2016; published: 29 july 2016 copyright: © 2016. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the etosha national park (enp) is a large protected area in northern namibia. while the enp has received a lot of research attention in terms of terrestrial ecosystem process understanding in recent decades, aquatic and hydrological research has to date been limited to a descriptive form. this study provides a baseline hydrological data set of the spatial representation of oand h-isotope ratios in the groundwater at a park scale, with a focus on three water point types utilised by game, namely natural artesian and contact springs as well as artificial boreholes. the data are used to infer broad-scale hydrological process from groundwater recharge mechanisms dominated by direct rainfall recharge in the west of the enp to evaporative controls on surface water recharge pathways in the east of the enp close to fishers pan. the findings are used to recommend further targeted research and monitoring to aid management of water resources in the enp. conservation implications: the terrestrial ecosystem, particularly large game, are tightly coupled to the distribution of available surface water in the enp, notably contact and artesian springs. within the enp there is a perceived desiccation of these springs. this study provides a baseline upon which more comprehensive studies should be undertaken to differentiate natural from anthropogenic causes for this phenomenon. introduction the requisite holistic understanding of natural and social environments within and around protected areas includes determining the complexities and challenges of managing water in their terrestrial environment (gonzález et al. 2013). these challenges include increasing water demands and often altered catchment processes arising from increasingly dense and urbanising populations on the peripheries of small and large protected areas, as seen frequently in southern africa. the etosha national park (enp) in north-central namibia is a case in point. the enp encompasses a biologically diverse semi-arid woodland savanna ecosystem, whose core is occupied by a large depression, the etosha pan. this is a superpan formed during the pleistocene by the coalescing of several smaller pans following the tectonic separation of the owambo basin from the kunene basin and consequent deflection of the kunene river towards the coast during the late pliocene (hipondoka 2005; miller, pickford & senut 2010). the etosha pan is also a ramsar wetland of international importance. in the definition of rodríguez-iturbe and porporato (2004), the enp would be classified as a water-controlled ecosystem, not only because of its water scarcity but also because of its intermittent and sometimes erratic availability. it is for this reason and the fact that the spatio-temporal distribution of biota in enp is tightly coupled to its water distribution (auer 1997) that baseline hydrological studies are required to be undertaken within it. within this context, a snapshot (conducted over the period of 1 week) survey similar to that developed by grayson et al. (1997) was undertaken for the enp during the dry winter months to provide a baseline stable isotope hydrology profile of the enp. research method and design the hydrochemical sampling of the water points in enp was conducted between 05 and 11 may 2013 and took the form of both a west-east and a south-north transect (figure 1). three types of water points were considered: groundwater boreholes, artesian springs and contact springs. twenty-five millilitre veterinary polystyrene vials, which were rinsed three times with sample water before collection and filled to the brim to prevent evaporative fractionation, were used in the following manner: groundwater borehole: by collection from borehole taps (ensuring that the pump was running continuously to the water trough in order that an unevaporated sample was collected from the aquifer). artesian springs: a sink-bottle sampler (500 ml plastic drinks bottle with weight tied to the bottle opening and three pin holes introduced at the base, so that bottle sinks into the water column) tied to a rope and thrown as close to the artesian spring eye as possible. contact spring: free flowing water in the contact spring. figure 1: location of groundwater sampling points following an approximate west-east (dolomietpunt to kameldoring) and south-north (eindpal to mbari) transect in the etosha national park, may 2013. field samples were stored in a cooler bag and then stored in a fridge before transport to the laboratory. a second sample was taken in the field to determine temperature (°c) ± 0.5 °c, electrical conductivity (ms/cm) ± 1% full scale and ph ± 0.01 ph using a calibrated eutech instruments pcstestr 35 pocket tester. samples were then sent to the university of kwazulu-natal to measure the stable isotope compositions of oxygen-18 (o18) and deuterium (h2) using a los gatos research (lgr) dt-100 liquid water isotope laser analyser and analysed six times to provide a high-value precision using in-house standards of indian ocean water (δ2h 4.75‰ ± 0.41‰; δ18o 0.06‰ ± 0.30‰) and spring water (avian bottled water δ2h -61.43‰ ± 0.56‰; δ18o -4.05‰ ± 0.29‰) and a 50:50 mixture of these. the standards were prepared against international atomic energy agency (iaea) standards lgr2, vsmow2 and ia-ro53 during which standard deviations of δ2h were less than 2‰ and δ18o less than 0.3‰. at each point, a differential gps reading was also taken with trimble geoexplorer 3® to determine accurate elevations above the mean sea level (± 10 cm). results and discussion field data may be found in table 1 and are discussed here in summary form to give an overall picture of the isotope composition of groundwater in the enp during the short time period of 1 week. this data set may thus be used to augment the spatially representative but temporally disparate data set presented by auer (1997). table 1: water hydrochemical parameters recorded in the etosha national park during may 2013. as seen in figure 2, in all cases the water sampled during may 2013 in the enp show signs of having been evaporated. because all samples plot below the global meteoric water line (gmwl), this gives a generalised local evaporation line (lel) from these groundwater samples. the lel with a slope of 5.1 is in similar agreement with the 5.2 lel slope developed in lindenmaier and christelis (2012). the windhoek meteoric water line is plotted for reference, because this is the closest available long-term data set for the region. while this has a slope of 7.13, gaj et al. (2016) produced a local meteoric water line for the cuvelai-etosha basin to the north with a slope of 7.3. the plotting of all groundwater stable isotope data below the gmwl would be expected given the high potential evapotranspiration (2600 mm – 2800 mm per annum) for the region and the convective nature of rainfall in northern namibia, whose origins are in the zaire air boundary of the inter-tropical convergence zone (see engert 1997). overall, there is a clustering of borehole, contact spring and artesian spring water (central), but several contact and artesian spring waters show evaporative loss (plotting to the right and below the gmwl). these data are plotted in figures 3 and 4 with spatial and altitudinal distribution and as a function of the deuterium excess, or d-excess (δ2h -8* δ18o). where the lower the numeric value, the influence of evaporation becomes more likely as δ18o increases, as result of the different equilibrium fractionation between the water molecules of 2h1h16o as related to 1h1h18o (dansgaard 1964). figure 2: stable isotope results (δ18o/δ2h) for samples of different water point types in the etosha national park, may 2013 (windhoek meteorological water line developed from international atomic energy agency global network of isotopes in precipitation database 1961–2001). figure 3: variation of altitude, δ2h and δ18o (above) and δ2h excess (below) along an approximate east-west transect for different water point types in the etosha national park, may 2013, with altitude reference. figure 4: variation of altitude, δ2h and δ18o (above) and δ2h excess (below), along an approximate north-south transect for different water point types in the etosha national park, may 2013, with altitude reference. figure 3 shows that most of the sampled water points in enp plot between -5‰ and 5‰ for d-excess while several artesian water points (homob, ngobib, chudop, klein namutoni, koinahas and groot okevi) have d-excess lower than -5‰. in general terms then, it can be speculated that the artesian water points suggest more variability in their water origins. the fact that this variability is pronounced in the east of enp points to the role of the oshivelo artesian and phreatic aquifers as being the primary groundwater pathways in this region as described by christelis and struckmeier (2011). meanwhile, mushara and kameldoring plot in a very similar range (< -6‰ for δ18o and < -50‰ for δ2h) as groundwater samples reported from 2008 in the north-east corner of the etosha pan by lindenmaier and christelis (2012). the boreholes in the west of enp all show relatively low delta values compared with the entire range of samples. it was noted by department of water affairs and forestry (2006) that the borehole at dolomietpunt showed a relatively large and rapid response to significant rains during the 2005 rainy season and that soils in the west are typically sandy (beugler-bell & buch 1997). the boreholes that align in an approximate south-north orientation to the west of the etosha pan (figure 4) also show low delta values, similar to those further west (as in figure 3). hence, these depleted groundwaters in the west of enp may then be indicative of direct recharge from large rain events through hydraulically conductive kalahari sands in an otherwise low rainfall region (300 mm pa – 350 mm pa). this will cause a minor evaporation of the recharging water body as it moves rapidly below the root zone of the evapotranspiring vegetated surface. in general, the ec readings (figure 5) reveal the same trends described by auer (1997) in that the western region of enp has low ec values. in this case, < 250 ms/m, which is in line with auer’s (1997) alkaline earth-hydrogen carbonate water, which is low in ions in the extreme west of enp, which up to toespad is < 300 ms/m. this freshwater chemistry is similar to auer’s (1997) data in the south of enp at ombika and also gruneveld and panpoint. we found high ec at > 500 ms/m at eindpal and narawandu, and this is similar to that found by auer (1997); however, they showed that ec was greater in the case of the former, whereas we found the opposite. meanwhile, we found mbari to have a low ec, which appeared to have not been sampled by auer (1997). in the east of enp, water sampled in this study at both contact and artesian springs along the shore of the pan generally had ec > 300 ms/m. this is generally in agreement with auer’s (1997) data, although there are some spatial inconsistencies between the two data sets, at specific points. although not revealed in our sampling, in the auer (1997) data there is an increase in ec closer to the shore of the pan. figure 5: spatial distribution of ec (ms/cm) for different water point types in the etosha national park, may 2013. (possible instrument error as uncalibrated spare multi-parameter meter had to be used for all the south-north transect points, 10 may 2013.) the spatial distribution of temperature (°c) in enp is shown in figure 6. typically, there is a general decrease in groundwater temperature at the boreholes west of deiwelsvuur, which augments the theory of more direct recharge through the kalahari sands and local dolomite formations by incoming rainfall as one moves further west. in the east of enp, around the shores of the pan, temperature is more variable and indicative therefore of the various source pathways of water from the oshivelo aquifer system emerging at artesian discharge or contact points. of interest are the contact springs halali and sueda, which both have cool temperatures (19.4 °c and 20.3 °c, respectively) and very high ec (10.58 ms/m and 9.72 ms/m) indicative perhaps of deep water sources traversing through weathered limestone in that region. in the far east of the pan, around fisher’s pan, the temperature is also reasonably low (23 °c – 24 °c) while the waters show some evaporative loss as suggested by the very low d-excess values. this suggests direct shallow recharge by water flowing into fisher’s pan from the omuramba-owambo river but where it is subject to significant evaporation in and around the pan before replenishing groundwater in the area. figure 6: spatial distribution of temperature for different water point types in the etosha national park, may 2013. the distinction in water point types that arises from the temperature data is reaffirmed in figure 7, which displays a ternary plot of d-excess, ec and temperature, which along with k-means analysis (table 2) depict three distinct clusters to the water samples. cluster 1 represents the majority of the water samples and includes those from all three water source types throughout the enp. it is also important to note within cluster 1 the close clustering of borehole water samples, suggesting very similar hydrochemistry, at least under this limited analysis. meanwhile, cluster 2 represents only artesian boreholes. all are at the far east of the enp with the exception of homob. cluster 3 consists of only two contact springs on the southern shores of the etosha pan at sueda and halali. figure 7: ternary plot with normalised data (percentiles) for ec (ms/cm), temperature (°c), and δ2h excess for different water point types in the etosha national park, may 2013, inclusive of k-means centroids (north-south transect samples omitted from the analysis because of unreliable multi-parameter probe readings). table 2: k-means centroids for groundwater ec (ms/cm), temperature (°c), and δ2h excess recorded in the etosha national park during may 2013. based on this snapshot analysis of groundwater in the enp, a preliminary picture is inferred from the data to broadly distinguish two hydrogeological areas: the west of the enp is likely to have groundwater that is recharged relatively directly from incoming rainfall through the sandy substrate in that landscape and from the dolomite ridge that runs along the south-western boundary of the park; meanwhile, the east is much more complex, as revealed by the k-means analysis, which clusters artesian and contact springs into three water facies. the network of artesian and contact springs that are likely to be connected to the oshivelo aquifer arise from the tsumeb-grootfontein dolomite complex. the data here support the conceptual model of a stratified oshivelo aquifer system as described by christelis and struckmeier (2011). recommendations for management there are a number of questions and recommendations raised by this limited study that could be used to inform future water availability and management strategies in the enp. first and more locally would be to determine the hydrological processes maintaining the okondeka freshwater spring, which makes it geographically distinct from other springs in the region, because it is the only active contact spring on the western shores of the etosha pan. the possibility that the dune fields to the west supply local recent event rainfall freshwater could be explored, as this would warrant that spring system a unique ecosystem type within the enp. the second broader, pertinent issue is to develop a groundwater hydrodynamic and hydrochemical monitoring programme for the enp and its groundwater catchments. the purpose of this will be to further characterise the hydrogeological processes in different water regions of the park, for example, western region and eastern region. for instance, feedback from managers in the enp suggests that there has been a steady desiccation of the springs along the south-west of the pan. it will need to be determined whether this is as a result of inter-glacial climate cycling or something more serious, such as groundwater over-abstraction elsewhere in the catchment and/or enhanced precipitation interception and transpiration from bush thickening in the karst aquifer of the region (e.g. namibia agriculture union 2010). conclusion the results of this snapshot hydrochemical and stable isotope characterisation of groundwater in the enp have revealed a close clustering of borehole water properties towards the west of the park and the d-excess values there suggest direct recharge by rainfall through the kalahari sands. the west of the park clearly has a more complicated groundwater flow pathway and the variable hydrochemistry reported here for artesian and contacts springs along the southern shore of the etosha pan support the conceptualised flow paths of other studies. the results of this baseline study could be used in future long-term hydrological change monitoring of the etosha ecosystem, especially with respect to understanding the causes for apparent desiccation of contact springs along the southern shores of the pan. acknowledgements dr jaco nel of groundwater consulting services is acknowledged for his advice in preparation for the field study and lending of some field equipment. mr cobus pretorius is acknowledged for analysing the water samples for stable isotopes at soil & water laboratory of the centre for water resources research at the university of kwazulu-natal. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions e.s.r. led the research design and was responsible for data analysis and compiling the manuscript. w.k., w.v. and m.k. assisted with field data collection and site interpretation. references auer, c., 1997, ‘chemical quality of water at waterholes in the etosha national park’, madoqua 20(1), 121–128. beugler-bell, h. & buch, m.w., 1997, ‘soils and soil erosion in the etosha national park, northern namibia’, madoqua 20(1), 91–104. christelis, g. & struckmeier, w., 2011, groundwater in namibia: an explanation to the hydrogeological map, ministry of agriculture, water and rural development, windhoek, namibia, p. 132. dansgaard, w., 1964, ‘stable isotopes in precipitation’, tellus 16, 436–468. http://dx.doi.org/10.1111/j.2153-3490.1964.tb00181.x department of water affairs and forestry, 2006, cuvelai-etosha groundwater investigation, version 1.1. bittner water consult cc, windhoek, namibia, p. 72. engert, s., 1997, ‘spatial variability and temporal periodicity of rainfall in the etosha national park and surrounding areas in northern namibia’, madoqua 20(1), 115–120. gaj, m., beyer, m., koeniger, p., wanke, h., hamutoko, j. & himmelsbach, t., 2016, ‘in situ unsaturated zone water stable isotope (2h and 18o) measurements in semi-arid environments: a soil water balance’, hydrology and earth system science 20, 715–731. http://dx.doi.org/10.5194/hess-20-715-2016 gonzález, h.f., porras, j.l.c., guitérez, i.b. & lamoreaux, j.w., 2013, management of water in protected areas, springer-verlag, berlin heidelberg, p. 351. grayson, r.b., gippel, c.j., finlayson, b.l. & hart, b.t., 1997, ‘catchment-wide impacts on water quality: the use of “snapshot” sampling during stable low flow’, journal of hydrology 199(1–2), 121–134. http://dx.doi.org/10.1016/s0022-1694(96)03275-1 hipondoka, m.h.t., 2005, ‘the development and evolution of etosha pan, namibia’, phd thesis, universität würzburg, p. 152. lindenmaier, f. & christelis, g., 2012, groundwater for the north of namibia: summary report of activities of phase i, exploration of ohangwena ii aquifer and preliminary isotope study, vol. 1a, ministry of agriculture, water and forestry, windhoek, namibia, p. 69. miller, r, mcg., pickford, m. & senut, b., 2010, ‘the geology, palaeontology and evolution of the etosha pan, namibia: implications for terminal kalahari deposition’, south african journal of geology 113, 307–334. http://dx.doi.org/10.2113/gssajg.113.3.307 namibia agriculture union, 2010, the effect of bush encroachment on groundwater resources in namibia: a desk top study, final report, colin christian & associates, environmental consultants, windhoek, namibia, p. 126. rodríguez-iturbe, i. & porporato, a., 2004, ecohydrology of water-controlled ecosystems: soil moisture and plant dynamics, cambridge university press, cambridge. filelist convert a pdf file! supplement to koedoe. 1977: 171-180. the status of nature conservation in the developing territories of south and south west africa m j de beer chief professional officer (nature conservation) department of bantu administration and development pobox 384 pretoria 0001 comments on the status of nature conservation in an area require an evaluation of progress. apart from ad hoc nature conservation actions taken in the past by the agricultural and administrative staff of the department of bantu administration and development (hereafter called the department) and of the governments of the various developing territories, or homelands, the more formal and specialised nature conservation action by these bodies started almost exactly three years ago. at that time the t ranskei already had a nature conservation section for about a year. three years ago, under the guidance of a senior agriculturalist, the department made its first appointment of a professional conservationist in pretoria, as well as of a number of nature conservation officers. these last-named officials were immediately seconded to some of the developing territories to fill vacancies in newly established nature conservation sections. since these "sections" frequently consisted and still consist of only one or two posts, it has not yet been possible to fill all these posts because of a scarcity of suitable applicants, and since the nature conservation section in the department itself has subsequently only grown from one to two officials, the question arises whether this is at all the correct time for stocktaking and evaluation . to see the matter in perspective it is necessary also to consider the diversity and distribution of the vast areas involved, scattered as they are from the northernmost boundaries of south west africa down to the eastern cape and covering a surface of more than 45 million hectares. t able i provides a more complete picture. weighing available facilities and personnel against the scope of the task, there can be no doubt that insufficient time has passed for the achievement of much more than preliminary objectives. on the other hand, a stocktaking at this stage will provide the general public with a clearer picture of the exten t of the task, the nature of the obstacles and of such progress as has been made. understandably, progress at this stage will be mainly in the fields of planning and organisation. 17! table 1 territories where nature conservation is controlled by the homelands and the department of bantu administration and development territories where the department*2 is sti ll territories responsible for nature conservation r es ponsible for nature conservation south africa *' south west africa sou th africa *3 south west africa territory area (ha) t erritory area (ha) territory area (ha) t erritory area (ha) transkei 4 168000 kavango 4 170050 swazi 372000 kaokoland 5525 129 ciskei 533000 owambo 5607200 ndebele 75000 damaraland 4779021 bophuthatswana 3826000 eastern caprivi i 153387 bushmanland i 837000 lebowa 2268000 hereroland 5899680 venda 650000 other 863350 gazankulu 675000 kwazulu 3 174000 qwaqwa 48000 total 15342 000 10930637 447000 18 904 180 i *' a sma ll portion of the land class ified here still falls und er t he jurisdiction of the departm ent. *2 th e departm ent of bantu administration and d evelo pm ent. *3 in these territories provincia l nature conservation legis lation is sti ll appli cab le and law enforcem ent rests with the provincial authorities. c'i r-the total area involved (i.e. about 45 million ha) has already been mentioned. the transkei will be discussed in a separate paper on this symposium. if its area is subtracted we are still left with about 41 million ha, which should be comparable with and probably exceeds that under the jurisdiction of many conservation organisations in southern africa. there are other factors to consider. these territories are variable, ranging from mountainland to plain, from rain forest to desert. according to acocks (1975) the territories falling within the republic of south africa (rsa) include no fewer than 34 of its 70 main veld types. endemicity of plant and animal life occurs , and scenic beauty unsurpassed anyhere in southern africa. with the transfer of nature conservation to homeland authorities at the required level of self-government, this enormous task is being broken down into more manageable portions, but this also necessitates a certain amount of organisational and functional replication. viewed against the background of political evolution towards self-government, however, this is to be welcomed and the sooner all the peoples concerned make a start with their own conservation programmes, the better. this is being done with the assistance of the department. legal background until recently the nature conservation organisations of the provinces and of south west africa provided the necessary legislation and general executive functions for all the territories under discussion. the complexity ofa situation where large tracts of populated land were either under state control or undergoing political evolution at varying rates and where cultural diversity required different approaches to legislation, lawenforcement and executive conservation action in general, necessitated a more specialised approach. although occasional regulatory functions such as the issue of permits for hunting or plant collection had been undertaken by the department or local authorities within the homelands, the central government decided to establish the legal framework for more efficient action. today this is to be found in the bantu homelands constitution act (act no. 21 of 1971) which provides for the transfer of wildlife management and the conservation function to the developing territories inside the republic of south africa. for south west africa it is to be found in the development of self-government for native nations in south west africa act (act no. 54 of 1968). the responsibility for nature conservation has already been assumed by the transkei, ciskei, kwazulu, lebowa, gazankulu, venda, bophuthatswana, qwaqwa, owambo, kavango and the eastern caprivi. all of these, with the exception of the eastern caprivi, have already adopted the necessary legislation. admittedly this legislation has in most instances been based largely on the ordinances of the provinces and may eventually have to be refined and adjusted more specifically to condi173 tions within each territory. at present, however, it provides a suitable basis for the management functions required at this stage. until the eastern caprivi adopts its own nature conservation legislation the south west african ordinance, number 5 of 1927, remains in force there. all the above-mentioned homelands are responsible for law-enforcement and the implementation of legislation within their territories. in terms of act no. 54 of 1968 the department has also prepared regulations on nature conservation to replace the relevant south west african ordinance in respect of kaokoland, damaraland, hereroland and bushmanland . all actions in terms of these regulations, which are now being promulgated , are the department's responsibility. the various provincial ordinances still apply to all land of the south african bantu trust within the rsa which has not yet been allocated to a specific homeland. law enforcement there is still the duty of the relevant provincial nature conservation authority. the same holds true for territori es not yet at the required level of self-government such as those of the swazi and south ern ndebele. the only exception here is the manyeleti gam e reserve, which was established on trust land in terms of the bantu administration act (act no. 38 of 1929) and the bantu trust and land act (act no. 18 of 1936) and which is being maintained by the department. these acts of course allow the future establishment of similar conservation areas on trust land. organisational development the first specialised nature conservation staff needs in the department itself arose with the establishment of the manyeleti game reserve in 1967. since its inception this reserve has been manned nearly exclusively by black people. with the exception of the manager and one assistant the present staff of approximately 60 and including the clerks, a public relations officer and game rangers, are all black people. all are departmental officials. the manager and his assistant are responsible for supervision and the training of staff, apart from supervisory assistance on other trust land situated nearby. control over manyeleti still rests with the department. during june 1972 a start was made with the appointment of nature conservation staff in the homelands. the first appointments were made in the transkei and other territories followed in 1973. because of a shortage of suitably qualified blacks, the department requested and obtained the approval of homeland governments to appoint and delegate for homeland service a number of white officials. they were to initiate the new conservation organisations and hold their positions until these are taken over by black people with the appropriate training and experience. since homeland organisations necessarily had to grow from small beginnings, they were attached to the only existing government departments involved in the utilisation of biological resources, namely 174 forestry and agriculture. at present nature conservation staff in all cases are attached to the departments of agriculture. through the kind co-operation of the kwazulu government service training facilities for black nature conservation officers from all developing territories inside the rsa now exist at the cwaka agricultural college. the first students obtained their two-year post standard 8 diplomas at the end of 1975 and have already assumed duty in their respective homelands. as they gain in experience and prove their mettle they can be expected to gradually fill the more senior positions. until that happens and for as long as they are needed, the department will assist by appointing white nature conservationists for homeland service. unfortunately our advertisements for nature conservationists have been drawing a poor response , at least from suitably qualified and experienced people. although nature conservation sections in the developing territories have to parallel all functions normally undertaken by a division or department of nature conservation, it has already been mentioned that in many instances these sections started with only one post. fortunately, nature conservation received sympathetic treatment from the various government service commissions and quite a number of new posts have subsequently been created. at present the available posts for all our developing territories are as follows (the t ranskei is not included, although one senior nature conservation officer of the department still does duty there): table 2 nature conservation posts in developing territories nature cons ervation professional officers (college diploma rangers (limited staff (unior extensive experience) formal education versity qualiin-service territory fications) senior ranks ordinary training) ciskei 0 i 4 4 kwazulu i 3 5 14 gazankulu 0 1 4 0 venda 0 i 4 17 bophuthatswana 0 3 21 48 lebowa 0 3 4 25 qwaqwa 0 0 0 0 eastern caprivi 2 i 3 14 kavango 0 i 2 19 owambo i i 2 i total 4 15 49 142 175 at least three more territories are expected to have professional posts in the near future. of the four professional posts in homelands, only one (in the eastern caprivi) is filled at the moment. of the 15 posts for nature conservation officers of senior or higher rank , only 10 are filled at the time of writing. this shortage of staff is the greatest single obstacle in the way of progress. the department's own nature conservation section started with the appointment of the first professional conservationist in pretoria in 1973. at present this section consists offour posts , of which two are filled. the function of this departmen tal section is to take direct charge of conservation management on land of the south african bantu trust not ye t under the jurisdiction of a specific homeland and on the land of the swazi and southern ndebele, as well as the areas known as kaokoland, damaraland, bushmanland and hereroland in south west africa. because of their distance from pretoria steps have been taken to establish a nature conservation section for thes e territories in south west africa. at present this south west african section consists of one senior and three ordinary nature conservation posts to which the first appointments are now being made . use is also being made of a professional post created for agriculture. this section will be assisted by the nature conservation staff in pretoria who also carry out the centralised control and administration and give advice and aid to all hom elands undertaking nature conservation themselves. this assistance could cover any conceivable aspect of conservation management on the combined 26 million hectares. because of the extent of this task the present two officials understandably are con centrating on th e more basic aspects such as the improvement of staff structures in the homelands, assistance with draft legislation, assistance with work programmes and assistance with broad planning, in addition to becoming better acquainted with their area of operations. conservation action much of our progress has been made possible by the generous assistance received from other conservation organisations in the rsa and south west africa . not a single request for advice or help has yet been turned down. this is sincerely appreciated. secondly, the magnificent work done by the totally inadequate homelands' staff deserves the highest praise and is mainly responsible for what has been achieved. the task of training rangers has rested fully on the senior conservation staff in the homelands. a number of short courses have been held and in-service training was done. manyeleti game reserve was made available to officials and rangers from developing territories to allow them to obtain practical experience. the progress here is considered to be satisfactory, with the exception of those territories which have had no senior staff to do the necessary training. 176 an important prerequisite for progress is thorough planning. it is indeed fortunate that nature conservation and tourism form part of the planning projects now being undertaken on a national scale in the ciskei, kwazulu, lebowa, gazankulu, venda, bophuthatswana, qwaqwa and owambo. population pressure and political and other factors sometimes prevent wise land-use, but such national plans will at least provide guidelines to follow up with more specific and detailed conservation plans . only sound planning will accord nature conservation and tourism their rightful places. only through sound planning can tourism dependent on natural areas be prevented from eroding its underlying resource. detailed management and conservation plans of wildlife resources for every homeland deserve priority recommendation to his cabinet by every minister of agriculture and forestry. a master plan for nature conservation and tourism for kaokoland and damaraland has been commissioned by the department and is now nearing completion. this will be followed by a detailed plan for conservation management and tourism developm ent which will be completed during 1977. fair progress can also be reported in respect of law-enforcement, the use of mass media and meetings for extension purposes, and the identification of potential conservation areas. the situation varies from territory to territory, but in general the approach has been to consider the less sophisticated rural population as more in need of education than law-enforcement. this view is supported by the fact that when wild animal and plant life are destroyed, it mostly takes place in an effort to obtain necessary food or to meet demands from outside the homeland, rather than as the result of vandalism, recreational activities or the pursuit of luxuries. some use has been made of radio bantu, while schools and colleges have been lectured and the meetings of headmen attended. it is of the greatest importance , however, that the concrete advantages of wildlife conservation should be practically demonstrated for the benefit of the general public as well as the decision-makers in homeland policy and land-use. for this more time is needed, although a start has been made. bophuthatswana's revenue from this source during the previous financial year was ri2 000, mostly from hunting and fishing permits, confiscated articles and fines. since a start has now been made with the sale of live game and the amounts paid for permits can with justification be made more realistic , this source of income should grow substantially in future. the department itself is making good progress with projects which should eventually provide workopportunities and a high income from tourism. i t is hoped that the national conservation plans which i referred to previously will identify the major conservation areas. in addition, the nature conservation staff of the homelands and of the department have all proceeded with the identification of areas that for fairly obvious reasons should be devoted to conservation and tourism. in some cases 177 cabinet approval has already been obtained and negotiations for compensatory land and the actual establishment of reserves have reached and advanced stage. examples (my list is not complete) include a major portion of about 32000 ha of the pilanesberg complex in bophuthatswana where only the lack of co-operation of one tribe is preventing the conservation of the whole complex; the klipvoordam area, also in bophuthatswana, where a game reserve of approximately 10 000 ha is being established as part of a complex which will also include the dam, a recreation resort and a fish production unit; the blouberg complex of 4900 ha in lebowa which it is hoped will eventually include a scenic area of high botanical value adjoining a large game reserve. in kavango an already game-proof fenced area of about 40 000 ha in the mangetti, and another of about 700000 ha in the khaudum, are presently under consideration as nature reserves. negotiations to safeguard an estimated 40 000 ha of the andonivlakte in owambo have also been initiated. this last-named area adjoins the etosha national park. the department itself has also made satisfactory progress. the establishment of conservation areas (eloff 1975) covering an estimated 34000 km 2 for the peoples of kaokoland and damaraland was approved in principle on 7 may 1975. this has subsequently been announced by the prime minister. another 3 423 ha has also been added to the manyeleti game reserve this year bringing it to a total area of 20 629 hectare. in addition to the above-mentioned, some other possible conservation areas were identified and brought to the notice of the various planning groups. as work progresses, increasing attention will be paid to the extent to which different veld types and ecosystems are being safeguarded. at present the department does not have the staff and facilities to undertake research. until now available knowledge was sufficient to allow for considerable progress. as research becomes imperative, the homelands and the department can be expected to approach research institutions to undertake specific projects. some work on fresh water fish and limnology has already been done by a departmental official at lake liambezi in the eastern caprivi. lebowa has the benefit of research done by the zoology department of the university of the north on dams in that territory, while a botanist from the same university is doing plant surveys in manyeleti. periodic investigations have also been undertaken by the wildlife society of southern africa and private individuals in a number of territories. it is hoped that the black universities will play an increasingly important role as the need for applied research in the field of wildlife management grows. problems and obstacles many of the obstacles we encounter are not exclusive to the nature conservation movement, but also frustrate other efforts towards home178 land development. examples are population size in relation to available resources and traditional forms of land-use and occupation. people do not move easily from a place they consider to be an ancestral abode and a repository for holy objects. such a place above all other places is considered to be theirs on the basis of a complexity of values relating to ownership, tradition and religion. if conservationists demand evacuation of such a piece ofland in the interests of a very rare species of plant or animal, conflict is likely. extension work, financial assistance, agricultural planning, rotational grazing and stock reduction might alleviate the problem. it should be borne in mind, however, that traditionally livestock numbers are more important than livestock quality. stock reduction therefore, does not come easily either. should one decide to move the people, alternative land will first have to be found. this is frequently not possible. if such land is available, one tries to obtain the co-operation of the people concerned. if they agree, the problem is solved. if not, the obvious alternative appears to be force. a government consists of politicians. unpopularity with your own people means political suicide in any language. i see no solution to such a dilemma other than to convince local decision-makers of the advantages to be had, either on the alternative land, or from the specific conservation project. this takes time. such conflicts do not arise very often, although they do occur. it remains a fact, however, that within most of the developing territories land for conservation is scarce. forestry, agriculture and other forms of land-use with an apparently more immediate or short term survival value or potential for profit take priority. the very sympathetic treatment conservation proposals are accorded at cabinet level in the homelands is highly encouraging. it is also very fortunate that the pres en t steps of the sou th african governmen t towards better consolidation of the developing territories sometimes make available land which can either be used as compensation for other land that has to be specially conserved, or which itself merits conservation or tourism projects. conservationists from the homelands consider as a major problem the fact that decision-makers at all levels of homeland development are insufficiently informed about the nature, motives and advantages of nature conservation. they believe that specialised, full-time extension workers are required. such appointments have not yet been made. although their youth and ethnic grouping may sometimes be a drawback, the newly trained black conservationists should find it much easier to overcome communication problems arising from differences in cultural background and language. what is needed here is not the usual promotional or informative type of work done by nature conservation organisations in the rsa, but carefully planned and programmed activities that will change attitudes. in one or two developing territories military operations in the interests of national security represent occasional practical hindrances. in south 179 west africa the political future is hard to predict and therefore dictates the highest priority to our efforts there. until now homeland conservationists have made only modest financial demands and received no grounds for complaint. inevitably these demands will have to increase. this happens at a time when the general financial climate is not exactly favourable. we can only hope that these conservationists will be provided with the means to prove the substantial contribution which nature conservation, inland fisheries and tourism can make towards homeland development. conclusion i t is sometimes said that the extent to which a nation practises conservation indicates its level of civilisation. although we should have preferred more time before having to report on progress, i think enough has been achieved to indicate the direction which has been taken and the support the conservation movement is receiving from the governments of all the developing territories. may their actions in this respect, now and in future, bear witness on their behalf. references acocks , j p h 1975. veld types of south africa. mem. bot. surv. s. afr. 40 . eloff, f c 1975. meesterplan vir die bewaring, bestuur en benutting van natuurreservate in damaraland en kaokoland. interimverslae nrs . i en 2. (ongepu bliseer) . 180 page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 49 53 original research abstract an account is given of all invasive alien freshwater snail species samples found in the kruger national park currently on record in the national freshwater snail collection (nfsc) database. this report mainly focuses on samples collected during surveys of selected water bodies in the kruger national park (knp) during 1964, 1995, 2001 and 2006. the progress made by four alien invasive freshwater snail species, lymnaea columella, physa acuta, aplexa marmorata and tarebia granifera, in colonising water bodies since first being recorded in the knp is addressed. the results clearly show that all four species are in the process of becoming more widespread than they were when first recorded. however, t. granifera is the only one of these species of which exceptionally dense populations were encountered in several water bodies. all indications are that this species could already have had a negative impact on the species diversity of molluscs in these water bodies, and it is recommended that the situation be closely monitored. keywords: invader snails, lymnaea columella, physa acuta, aplexa marmorata, tarebia granifera invasive alien freshwater snail species in the kruger national park, south africa kenné n. de kock cornelius t. wolmarans school of environmental sciences and development (zoology) north-west university potchefstroom south africa correspondence to: kenné de kock e-mail: kenne.dekock@nwu.ac.za postal address: school of environmental sciences and development, north-west university, private bag x6001, potchefstroom, 2520, south africa in an assessment of the ecological impact and economic consequences of the introduction and dispersal of invasive alien organisms into southern african aquatic ecosystems, ashton et al. (1986) concluded that available information pertaining to molluscs was limited. in his revision of the available knowledge on the biology of invasive gastropods and the impact they may have on systems they have colonised in south africa, appleton (2003) lists four species as being invasive or as becoming so. two of these, lymnaea columella (say 1817) and physa acuta (draparnaud 1805), were introduced in the early 1940s (brown 1994, appleton & brackenbury 1998), while aplexa marmorata (guilding 1828) was collected for the first time in this country in 1986 (appleton et al. 1989). the most recently introduced species, tarebia granifera (lamarck 1816), was reported for the first time in africa by appleton and nadasan (2002) after it was discovered in 1999 in a reservoir supplying water to a paper mill in kwazulu-natal. in a recent survey of the mollusc fauna of the knp, we collected all four of these invasive alien species from several water bodies. although more than 25 different mollusc species from the knp are on record in the nfsc database, no other alien mollusc species are on record. this computerised database was compiled from more than 18 000 samples, the majority of which were collected during surveys conducted by staff of government and local health authorities. samples were then dispatched to the former snail research unit at the potchefstroom university for identification. however, at least 25% of these samples were collected by snail research unit staff during research projects aimed at studying the taxonomy, geographical distribution and ecology of the indigenous freshwater mollusc fauna. the database, along with more than 35 000 vials of preserved species samples, is located at the potchefstroom campus of the north-west university. the aim of the present contribution is to evaluate the progress of the four alien species on record in colonising water bodies in the knp by comparing the results of our recent survey with data in the nfsc database, dating back to 1959. methods and materials during our recent survey (wolmarans & de kock 2006) we concentrated mainly on water bodies that were screened for molluscs in 1995 (de kock & wolmarans 1998) and 2001 (de kock et al. 2002). molluscs were collected by means of aluminium scoops, preserved in alcohol, identified by the first author and subsequently incorporated into the nfsc. sites where the four alien invasive snail species were recovered, records of previous collections extracted from the database of the nfsc, as well as sites positive for other mollusc species but negative for the four alien species, were plotted on a map. information on the type of water body, the grid reference and the date of collection for each locality where freshwater mollusc species were recovered during our 1995, 2001 and 2006 surveys are presented in table 1. results of the four alien invader freshwater snail species under discussion, only three samples of l. columella from the knp were recorded prior to our 1995 survey. although samples of this species were recovered from only a single locality during our extensive survey in 1995 (de kock & wolmarans 1998), the results of our 2001 (de kock et al. 2002) and our 2006 (wolmarans & de kock 2006) surveys added eight and two new localities to its distribution respectively (see table i). the collection sites of all samples of l. columella currently on record in the nfsc database for the knp are depicted in fig. 1. the single sample of p. acuta recovered from the sabie river in 1995 (de kock & wolmarans 1998) was the first record of this species from the knp reported in print (see table i). two new localities were added as a result of the survey in 2001 (de kock et al. 2002) and another two as a result of our 2006 survey (wolmarans & de kock 2006) (see table i). the collection sites of all samples of p. acuta currently on record in the nfsc database for the knp are depicted in fig. 1. 49 2008 original research de kock & wolmarans vol. 50 no. 1 pp. 49 53koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za although the results of the survey in 1995 (de kock & wolmarans 1998) yielded only one sample of a. marmorata, it was also the fi rst recorded instance of the species in the knp. respectively, seven and three new localities were added to the list as a result of our 2001 (de kock et al. 2002) and 2006 (wolmarans & de kock 2006) surveys (see table i). the collection sites of all a. marmorata samples currently on record in the nfsc database for the knp are depicted in fig. 1. the recently introduced alien invader species, t. granifera, (appleton & nadasan 2002; appleton 2003), was collected from 12 water bodies during our 2006 survey (wolmarans & de kock 2006) (see table i). these collection sites are depicted in fig. 1. discussion lymnaea columella was not mentioned in a monograph by connolly (1939) after an in-depth revision of the extensive mollusc material collected by him and many other researchers. it is therefore reasonable to conclude that this species was extremely rare in south africa before 1944 (van eeden & brown 1966; brown 1967; 1978; 1980). according to brown (1994), the earliest record of l. columella dates back to 1942 when it was found near somerset west. since then, 2 302 samples of this species have been recorded in the nfsc database. these collection sites are distributed over 353 loci (�∕16 square degrees), ranking l. columella the third-most-widespread freshwater snail species in south africa. it is only surpassed by the two indigenous species, bulinus tropicus (krauss 1848) (939 loci) and s n ew 0 60 120 km berg-en dal crocodilecrocodilecrocodile bridge skukuza lower lower sabie pretoriuskop tshokwanetshokwane nwanetsi nwanetsi satara mooiplaas shingwedzi shingwedzi shingwedzi punda maria pafuripafuri vlakteplaas phalaborwa letaba kingfisherspruitkingfisherspruitkingfisherspruit figure 1 distribution of the alien invader freshwater snail species lymnaea columella (●), physa acuta (∆), aplexa marmorata (▲), tarebia granifera (○) and rainfall monitoring stations (♦). lymnaea natalensis (krauss 1848) (616 loci). in 1964, during the fi rst concentrated effort to systematically survey the knp for freshwater molluscs, l. columella was collected in two localities (oberholzer & van eeden 1967). however, it was identifi ed fi ve years earlier in a collection from the crocodile river in 1959. an extensive survey of water bodies in the knp in 1995 (de kock & wolmarans 1998) yielded only one new collection site. the fact that only three new collection sites were recorded for the 35 years up to 1995 suggests that l. columella has been less successful in invading water bodies in the knp than elsewhere in south africa (de kock et al. 1989). however, 10 new localities were recorded during our surveys in 2001 and 2006 (see table�i) and it would seem that this species has now become well established, especially in the southern areas of the knp. with regard to p. acuta, the species was reported by van bruggen (1964) from a single locality in pretoria, thereby establishing the fi rst published record of the species in south africa, although the oldest record in the nfsc database dates back to the umsindusi river, pietermaritzburg, in 1954. since then it has dispersed and colonised many water bodies in several south african river systems (hamilton-attwell et al. 1970, de kock et al. 1989). it is therefore rather surprising that the fi rst record of this species in the knp was only established more than fi ve decades after its introduction into this country, during our 1995 survey (de kock & wolmarans 1998). in view of the fact that only four new collection sites were added as a result of our extensive 2001 (de kock et al. 2002) and 2006 (wolmarans & de kock 2006) surveys, it seems justifi able to regard p. acuta as a less aggressive invader than l. columella. this view is supported by the fact that, without exceptions, only a small number of p. acuta specimens were collected in any of the water bodies in which it was present. this is surprising in view of the fact that population densities as high as ±3 000 m2 for this species were reported by brackenbury and appleton (1993), as well as a much higher reproductive rate than for the rselected bulinus tropicus (brackenbury & appleton 1991), which is generally considered to be the most widespread freshwater snail species in south africa (de kock et al. 2002). perhaps the fact that p. acuta commonly occurs in areas where sewerage pollution is obvious and that it seems to be closely associated with human activities (brown 1994) might explain its relatively slow progress in colonising the relatively unpolluted freshwater habitats in the knp. appleton (2003) also mentions the fact that p. acuta has acquired the nickname of ‘the sewage snail’ and is often the only mollusc living in water with high levels of faecal pollution. the fi rst report of the presence of the more recently introduced invader species, a. marmorata, in south africa was by appleton et al. (1989) from specimens collected near durban, kwazulunatal in 1983. these authors also reported two additional records for this species: one from kwazulu-natal and the other from tzaneen, in limpopo province, collected in 1986 and 1988 respectively. according to appleton (2003), a. marmorata is now widespread in the durban area. the fi rst sample of this species from the knp was collected by us in 1995, just over a decade after it was fi rst discovered in this country. this is much sooner than the fi ve decades that elapsed before the presence of p. acuta was fi rst recorded in the knp after its introduction in the early 1940s. in view of the fact that both species were initially recorded in localities in kwazulu-natal, it seems justifi able to assume that a. marmorata might be a more aggressive invader of relatively undisturbed habitats. this is supported by the results in fig. 1, which not only depicts the higher number of collection sites but also the wider distribution of a. marmorata in the knp. as mentioned earlier, the most recently introduced invader species of the four under discussion, tarebia granifera, was reported for the fi rst time in africa by appleton and nadasan (2002). according to these authors, it was discovered in 1999 in 50 invasive alien snail species original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 49 53 rainfall monitoring station water body grid reference 1995 2001 2006 mollusc species found invader mollusc species found mollusc species found invader mollusc species found mollusc species found invader mollusc species found berg en dal matjulu spruit 25˚ 21’s;31˚31’e none none yes lymnaea columella aplexa marmorata yes lymnaea columella mlambane spruit 25˚ 21’s;31˚31’e not surveyed yes aplexa marmorata yes none crocodile bridge crocodile river 25˚21’s;31˚53’e not surveyed not surveyed yes tarebia granifera gezantombi dam 25˚ 20’s;31˚52’e none yes lymnaea columella aplexa marmorata yes none lower sabie lower sabie dam 25˚ 07’s;31˚56’e not surveyed not surveyed yes tarebia granifera mpondo dam 25˚ 12’s;31˚43’e not surveyed yes lymnaea columella yes tarebia granifera sabie river below lower-sabie dam wall 25˚ 07’s;31˚56’e not surveyed yes tarebia granifera sabie river 25˚ 07’s;31˚55’e yes lymnaea columella physa acuta aplexa marmorata yes physa acuta aplexa marmorata yes none sunset dam 25˚ 06’s;31˚54’e not surveyed yes aplexa marmorata none pretoriuskop mbyamiti river 25˚ 13’s;31˚34’e yes none yes lymnaea columella aplexa marmorata yes none mbyamiti river weir 25˚ 13’s;31˚37’e yes none yes none yes tarebia granifera mestel dam 25˚ 06’s;31˚12’e not surveyed yes lymnaea columella aplexa marmorata yes tarebia granifera shitlhave dam 25˚ 08’s;31˚19’e none yes none yes none skukuza manzimahle dam 24˚ 35’s;31˚43’e none yes none none olifantdrinkgat 24˚ 53’s;31˚44’e none yes none yes none vervoer dam 25˚ 07’s;31˚29’e none yes lymnaea columella aplexa marmorata none tshokwane kumana dam 24˚ 37’s;31˚47’e none yes none yes none mazithi dam 24˚ 43’s;31˚50’e none yes none yes none nwaswitsontso river 24˚ 37’s;31˚47’e yes none yes none yes none orpen dam 24˚ 47’s;31˚53’e yes none yes lymnaea columella yes aplexa marmorata silolweni dam 24˚ 29’s;31˚50’e none yes none none nwanetsi gudzani dam 24˚ 22’s;31˚55’e yes none yes none yes tarebia granifera nwanetsi water hole 24˚ 28’s;31˚58’e yes none yes none yes tarebia granifera satara nkaya pan 24˚ 31’s;31˚45’e none yes none yes none nsemani dam 24˚ 23’s;31˚43’e none yes none none houtboschrand ngotso dam 24˚ 10’s;31˚43’e not surveyed none none piet grobler dam 24˚ 13’s;31˚38’e not surveyed yes none yes tarebia granifera timbavati river 24˚ 16’s;31˚38’e none yes none yes none timbavati river below piet grobler dam wall 24˚ 12’s;31˚38’e not surveyed yes none yes tarebia granifera phalaborwa sable dam 23˚ 56’s;31˚14’e not surveyed yes none none letaba engelhard dam 23˚ 50’s;31˚38’e no suitable habitat yes physa acuta yes physa acuta aplexa marmorata tarebia granifera letaba river 23˚45’s;31˚22’e none none yes physa acuta yes physa acuta aplexa marmorata letaba river below engelhard dam wall 23˚ 50’s;31˚39’e no suitable habitat yes none yes lymnaea columella physa acuta tarebia granifera olifants river 24˚ 31’s;31˚41’e no suitable habitat no suitable habitat yes physa acuta mooiplaas grootvlei dam 23˚ 20’s;31˚30’e not surveyed yes none yes none pioniers dam 23˚ 31’s;31˚23’e yes none yes none yes none shingwedzi kanniedood dam 23˚ 08’s;31˚27’e not surveyed yes none yes none krapkuil dam 23˚ 14’s;31˚18’e none yes none yes none shisa spruit 22˚ 57’s;31˚15’e not surveyed yes none none sirheni dam 22˚ 56’s;31˚14’e not surveyed yes lymnaea columella yes none punda maria klopperfontein dam 22˚ 37’s;31˚10’e none yes none yes aplexa marmorata pafuri luvuvhu river 22˚ 26’s;31˚13’e not surveyed no suitable habitat none none table 1 positive and negative records of four alien invader freshwater mollusc species collected in water bodies in the kruger national park during surveys conducted in 1995, 2001 and 2006 51 original research de kock & wolmarans vol. 50 no. 1 pp. 49 53koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a reservoir supplying water to a paper mill at mandeni, 95 km north-east of durban. subsequently, the collection of another 16 samples of t. granifera – 14 from kwazulu-natal and two from the crocodile river at malelane in the knp – were reported by appleton (2005). no specimens of t. granifera were found during either of our 1995 and 2001 surveys of selected water bodies in the knp; however, during our 2006 survey, 12 localities were recorded (see table i). from the fact that the first record of this species in south africa was only collected in 1999 and the fact that no specimens were found during our 1995 and 2001 surveys, it seems reasonable to assume that t. granifera did not reach the knp earlier than 1999. the results in fig. 1 support reports in the literature that this species readily becomes invasive after its introduction into a new country. it took a. marmorata and l. columella 10 and 47 years respectively to invade more than 10 water bodies in the knp, while t. granifera accomplished this in only about five years. according to the data in the nfsc database, only relatively small numbers of specimens of the l. columella, p. acuta and a. marmorata could be collected in each locality in the knp. in contrast to this, extremely high numbers of t. granifera were found at five of the 12 collection sites during our 2006 survey. appleton (2003) also reports very high numbers of this species from two habitats in kwazulu-natal. in a discussion of the ecological impacts of invasive snails, appleton (2003) raises the point that high densities of alien snails must surely reduce the food available to indigenous grazing invertebrates and that in high-density situations, physical interaction between invasive snails and indigenous invertebrates could also be important. the negative effects of increased physical contact between snails in high density situations on reproduction and growth have been reported by several authors in the past (dewitt 1954, chernin & michelson 1957, berrie 1968). in puerto rico and cuba, invasion by t. granifera was also associated with the disappearance of an indigenous snail species (pointier 1999). in accordance with these observations, a marked decrease in species diversity was found at five of the 12 sites where t. granifera was collected in 2006, as compared to the results of our survey in 2001. from fig. 1 it is clear that three of the four invader species have become well established in the southern part of the knp. the success of l. columella as invader is partly ascribed to its amphibious habits (van eeden & brown 1966, appleton 2003), wide temperature tolerance (brown 1994) and its relatively high intrinsic rate of natural increase (de kock et al.1989). likewise, tarebia granifera is r-selected and also parthenogenetic (appleton 2003); these attributes contribute to its success as invader. while the introduction into south africa of l. columella, p. acuta and t. granifera was probably via the aquarium trade, it seems possible that a. marmorata has been present in south-eastern africa for at least 160 years (appleton 2003). the airborne dispersal of flightless small animals is an established fact and is known to be facilitated mainly by dispersal agents such as wind, insects and birds (rees 1965). according to rees (1965), it is well documented that birds play a more important role in the dispersal of molluscs than either wind or insects. to substantiate this statement, he lists a number of cases reported in the literature that incriminate bird species (ducks, plovers, herons) as dispersal agents for a variety of gastropod and bivalve species. passive transport by water currents in lotic habitats can obviously play a role in the dispersal of aquatic fauna, and it has been found that p. acuta is also capable of actively migrating upstream against the current (appleton & branch 1989). immigration from refugia after heavy floods also plays an important role in the recolonisation of a habitat, as reported by brackenbury and appleton (1993) for p. acuta in the umsindusi river, kwazulu-natal. egg clutches and snails can also be successfully transported over relatively short distances on the hooves of animals, which could have contributed to the gradual colonising of proximate suitable habitats after a first introduction to the kruger national park. however, birds most probably contributed largely towards the spreading of the alien invader snail species – a view supported by the discontinuous distribution pattern plotted for l. columella, p. acuta and a. marmorata in fig. 1 and also by the presence of t. granifera in isolated localities such as the mestel dam. conclusions from the results of this investigation it is clear that all the invasive alien freshwater snail species currently present in the kruger national park are in the process of becoming more widespread. however, the rate at which t. granifera seems to be spreading is cause for concern, and we fully support the view of appleton (2005) that this is the species that should be monitored. acknowledgements we are indebted to the south african national parks for permission to conduct our research and for partially sponsoring the surveys; to prof l. van rensburg, director: research focus area, environmental science and sustainable management of the potchefstroom campus of the north-west university for financial support and to prof l.h. du preez of the school of environmental sciences and development of the potchefstroom campus of the north-west university for assistance in compiling the map. references appleton, c.c. 2003. alien and invasive fresh water gastropoda in south africa. african journal of aquatic science, 28: 69– 81. appleton, c.c. 2005. invasive freshwater invertebrates in south africa. plenary address to the congress of the zoological society of southern africa, grahamstown, july, 2005. appleton, c.c. & brackenbury, t.d. 1998. introduced freshwater gastropods in africa with special reference to physa acuta. in: madsen, h., appleton, c.c. & chimbari, m. (eds). proceedings of a workshop on medical malacology in africa, september 1997. harare: danish bilharziasis laboratory, pp. 22–26. appleton, c.c., brackenbury, t.d. & tonin, a.f.g. 1989. physa mosambiquensis (clessin, 1886) rediscovered? south african journal of zoology, 24: 340–344. appleton, c.c. & branch, g.m. 1989. upstream migration by the invasive snail physa acuta in cape town, south africa. south african journal of science, 85: 189–190. appleton, c.c. & nadasan, d.s. 2002. first report of tarebia granifera (lamarck, 1816) (gastropoda: thiaridae) from africa. journal of molluscan studies, 68: 399–402. ashton, p.j., appleton, c.c. & jackson, p.b.n. 1986. ecological impacts and economic consequences of alien invasive organisms in southern african ecosystems. in: macdonald, i.a.w., kruger, f.j. & ferrar, a.a. (eds). the ecology and management of biological invasions in southern africa. cape town: oxford university press, pp. 247– 257. berrie, a.d. 1968. prolonged inhibition of growth in a natural population of the freshwater snail biomphalaria sudanica tanganyicensis (smith) in uganda. annals of tropical medicine and parasitology, 62: 45–51. brackenbury, t.d. & c.c. appleton. 1991. effect of controlled temperatures on gametogenesis in the gastropods physa acuta (physidae) and bulinus tropicus (planorbidae). journal of molluscan studies, 57: 461–469. brackenbury, t.d. & appleton, c.c. 1993. recolonization of the umsindusi river, natal, south africa, by the invasive gastropod, physa acuta (basommatophora, physidae). journal of medical & applied malacology, 5: 39–44. brown, d.s. 1967. a review of the freshwater mollusca of natal and their distribution. annals of the natal museum, 18: 477– 494. 52 invasive alien snail species original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 49 53 brown, d.s. 1978. freshwater molluscs. in: werger, m.j.a. (ed.) biogeography and ecology of southern africa 2. the hague: junk, pp. 1155-1180. brown, d.s. 1980. freshwater snails of africa and their medical importance. london: taylor & francis ltd. brown, d.s. 1994. freshwater snails of africa and their medical importance, 2nd edition. london: taylor & francis. chernin, e. & michelson, e.h. 1957. studies on the biological control of schistosome-bearing snails. iv: further observations on the effects of crowding on the growth and fecundity in australorbis glabratus. american journal of hygiene, 65: 71–80. connolly, m. 1939. a monographic survey of south african non-marine mollusca. annals of the south african museum, 33: 1–660. de kock, k.n., joubert, p.h. & pretorius, s.j. 1989. geographical distribution and habitat preferences of the invader freshwater snail species lymnaea columella (mollusca: gastropoda) in south africa. onderstepoort journal of veterinary research, 56: 271–275. de kock, k.n. & wolmarans, c.t. 1998. a re-evaluation of the occurrence of freshwater molluscs in the kruger national park. koedoe, 41: 1–8. de kock, k.n., wolmarans, c.t., bornman, m. & maree, d.c. 2002. verspreiding en habitats van bulinus tropicus, tussengasheerslak van die peervormige bot, callicophoron microbothrium, in suid-afrika. suid-afrikaanse tydskrif vir natuurwetenskap en tegnologie, 21: 114–120. de kock, k.n., wolmarans, c.t. & du preez, l.h. 2002. freshwater mollusc diversity in the kruger national park: a comparison between a period of prolonged drought and a period of exceptionally high rainfall. koedoe, 45: 1–11. dewitt, r.m. 1954. reproductive capacity in a pulmonate snail (physa gyrina say). american naturalist, 88: 159–164. hamilton-attwell, v.l., de kock, k.n. & van eeden, j.a. 1970. the occurrence and distribution of physa acuta draparnaud in the republic of south africa. wetenskaplike bydraes van die potchefstroomse universiteit vir christelike hoër onderwys, reeks b: natuurwetenskappe, 26: 1–11. oberholzer, g. & van eeden, j.a. 1967. the freshwater molluscs of the kruger national park. koedoe, 10: 1–42. pointier j.p. 1999. invading freshwater gastropods; some conflicting aspects for public health. malacologia, 41: 403– 411. rees, w.j. 1965. the aerial dispersal of mollusca. proceedings of the malacological society of london, 36: 269–273. van bruggen, a.c. 1964. the distribution of introduced mollusc species in southern africa. beaufortia, 11: 161–169. van eeden, j.a. & brown, d.s.. 1966. colonization of fresh waters in the republic of south africa by lymnaea columella say (mollusca: gastropoda). nature, 210: 1172–1173. wolmarans, c.t. & de kock, k.n. 2006. the current status of freshwater molluscs in the kruger national park. koedoe, 49: 39–44. 53 abstract introduction materials and methods results discussion conclusion acknowledgements references about the author(s) kerry-ann van der walt department of ichthyology and fisheries science, rhodes university, south africa ernst r. swartz south african institute for aquatic biodiversity, grahamstown, south africa darragh woodford school of animal, plant and environmental sciences, university of the witwatersrand, south africa olaf weyl south african institute for aquatic biodiversity, grahamstown, south africa south african centre for invasion biology, grahamstown, south africa citation van der walt, k-a., swartz, e.r., woodford, d. & weyl, o., 2017, ‘using genetics to prioritise headwater stream fish populations of the marico barb, enteromius motebensis steindachner 1894, for conservation action’, koedoe 59(1), a1375. https://doi.org/10.4102/koedoe.v59i1.1375 review article using genetics to prioritise headwater stream fish populations of the marico barb, enteromius motebensis steindachner 1894, for conservation action kerry-ann van der walt, ernst r. swartz, darragh woodford, olaf weyl received: 07 dec. 2015; accepted: 31 oct. 2016; published: 03 feb. 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract south africa has a relatively large number of threatened freshwater fish species and limited resources to implement conservation programs. enteromius motebensis was regionally prioritised for action because of its conservation status and flagship status in a nationally important aquatic ecosystem. genetic diversity of e. motebensis in headwater refugia of the groot marico river catchment was assessed to determine if genetic diversity is important for conservation planning for this species. the results of the genetic analysis indicate that some prioritisation was possible, with two populations showing evidence of recent isolation. conservation implications: we recommend that at least three populations be prioritised for conservation action to ensure maintenance of most of the remaining genetic diversity of the species. introduction headwater stream ecosystems are often seen as areas of freshwater conservation priority (abell, allan & lehner 2007). the geographical isolation of headwater streams gives rise to high levels of endemism (abell et al. 2007) and often genetically distinct species (gomi, sidle & richardson 2002). globally and locally, headwater ecosystems are under increasing threat from human disturbance (aparicio et al. 2000; collen et al. 2014; ellender, weyl & swartz 2011; fausch et al. 2009; tweddle et al. 2009; vörösmarty et al. 2010). these threats include, but are not limited to, habitat alteration and water extraction for irrigation purposes and inter-basin transfers, which mediate the introduction of extra-limital and non-native invasive species. such environmental stressors can permanently alter these systems, driving the extinction of the biota that inhabit them (fausch et al. 2009; vörösmarty et al. 2010). non-native invasive species pose one of the greatest challenges to conservation in freshwater ecosystems (dudgeon et al. 2006; lowe et al. 2000) because they can have a serious impact on native organisms (moyle & light 1996; rahel 2002; whittier & kincaid 1999). in south africa, for example, the largemouth bass micropterus salmoides (lacepede 1802) was first introduced in 1928 for recreational angling in impoundments (hargrove et al. 2015). largemouth bass have subsequently invaded some headwater streams where their predation has altered invertebrate communities (weyl et al. 2010) and resulted in fragmentation of native fishes such as the marico barb enteromius motebensis (steindachner 1894) in the groot marico river catchment, limpopo river system (kimberg et al. 2014). the groot marico river catchment in the north west province (figure 1) is part of the limpopo river system and contains several perennial headwater tributaries, which contain unique landscape features and are of significant conservation importance (nel et al. 2011; skelton & ribbink 1994; smith-adao et al. 2006). these unique landscape features include groundwater-fed aquifers (dolomitic eyes) that arise from fractures at the contact zones between underlying dolomite intrusions of igneous rock, which provide permanent flow to headwater tributaries (wellington 1995). in an otherwise arid region, these features support a high number of endemic invertebrate taxa (grobler, roux & koni 2007; nel et al. 2011). the high level of endemism in conjunction with its free-flowing nature has resulted in the marico river and its catchment being listed as a national freshwater ecosystem priority area (nfepa) (nel et al. 2011). figure 1: map illustrating the nine sites where enteromius motebensis specimens were collected for genetic analysis, across the upper groot marico river catchment, north west province. the marico barb is near-endemic to the groot marico river catchment in the north west province, south africa, and is considered to be of particular conservation importance because of being red-listed as vulnerable (engelbrecht & bills 2007). it is a small (80 mm standard length) cyprinid fish characterised by numerous conical tubercles on the snout, forehead and lower jaw and has two pairs of barbels (skelton 2001). its reproductive biology suggests that it has an opportunistic life history with high fecundity, an extended breeding season (spring to summer) and early maturity at 50% of their maximum length (kindler, wagennar & weyl 2015). taxonomically, e. motebensis belongs to the chubbyhead barb group (engelbrecht 1996). according to skelton (2001), the other members of this group are enteromius anoplus weber, 1897 (chubbyhead barb), enteromius gurneyi günther, 1868 (redtail barb), enteromius amatolicus skelton, 1990 (amatola barb) and enteromius breviceps trewavas, 1936 (shorthead barb). previous studies using an allozyme analysis suggest that members of the chubbyhead barb group are typically restricted to upper catchments of rivers and are often isolated in headwaters of rivers, which results in extensive genetic structuring among populations (engelbrecht 1996; engelbrecht & van der bank 1994, 1996). in the upper reaches of the groot marico river catchment, e. motebensis is limited to the headwater streams of the study area, with a small area of occupancy, which is less than 2000 km2 (engelbrecht 1996). this limited distribution may be caused by natural barriers, weirs, dams, water abstraction, habitat alterations and predation by m. salmoides (see smith-adao et al. 2006). in addition, the fragmentation of this population may further be a result of dispersal ability, habitat preference, isolation by distance and physiological adaptations (chakona, swartz & chakona 2015), all of which result in reduced dispersal success and limited gene flow, which may result in the development of novel species or facilitate localised extinctions (narum 2006; segelbacher et al. 2010). therefore, an understanding of the genetic diversity and structuring of e. motebensis is an important first step towards developing a conservation strategy for this species (kimberg et al. 2014). the aims of the study were thus: (1) to assess the genetic structure and diversity of e. motebensis in the groot marico catchment across tributary locations to determine whether e. motebensis comprises one population or genetically distinct populations separated by natural barriers, (2) determine whether genetic isolation has occurred, (3) use these data to develop theories on how the ecology of e. motebensis, natural landscape and climatic processes may have influenced the genetic structuring of e. motebensis populations and (4) identify which populations of e. motebensis may require a conservation plan in the groot marico catchment. materials and methods study area and sample collection this study was conducted in the groot marico river catchment in the north west province (figure 1) part of the limpopo river system. samples were collected from the kaaloog se loop (henceforth abbreviated to kaaloog), draaifonteinspruit, bokkraal, ribbokfontein and vanstraatensvlei tributaries as well as from the groot marico main stem (figure 1). there are five known potential in-stream barriers (roux 2010) that may restrict upstream migration (hughes, schmidt & finn 2009). four of these barriers are natural in the form of waterfalls and are found in the draaifonteinspruit, bokkraal and polkadraaispruit tributaries and in the upper reaches of the groot marico main stem. the last barrier is an artificial weir on the lower reaches of the draaifonteinspruit tributary (figure 1). sixty-eight e. motebensis individuals were collected from nine locations (figure 1, table 1) in march and november 2012 using a samus backpack electrofisher (90-hz pulsed-dc current) and seine netting. at each locality, up to 10 individuals of e. motebensis were euthanised with clove oil (0.2 ml/l). the rest of the captured fish were returned alive. small pieces of muscle tissue from each specimen were then removed using a dissection blade and placed into 1.5-ml vials containing 98% ethanol. the remaining specimens served as voucher specimens and were fixed in 10% formalin and later transferred to 70% ethanol. voucher specimens were accessioned into the national collection of the south african institute for aquatic biodiversity (saiab) in grahamstown, south africa. the tissue vials were then stored in a freezer at -20 °c for dna extraction, and once all laboratory work was completed, the tissue vials were transferred to saiab’s biomaterial bank for long-term storage in ultra-deep freezers at -70 °c. table 1: localities and sample sizes (n) analysed for the mitochondrial nucleotide diversity 2 gene of enteromius motebensis. dna extraction, amplification and sequencing total genomic dna was isolated from muscle tissue using the nucleic acid and protein purification extraction kit, nucleospin tissue (machery-nagel, germany). a 947-base pair fragment of the mitochondrial nd2 gene was amplified with the primers nd2-f and nd2-r (kocher et al. 1995). the polymerase chain reactions (pcrs) were performed with a veriti® 96-well thermal cycler (applied biosystems, north america). each reaction mixture (25 µl) contained 1x reaction buffer, 2.5 mm mgcl2, 0.8 mm of each dntp (kapa biosystems [pty] ltd, cape town), 0.2 mm of each primer, 1 u super-therm taq polymerase (southern cross biotechnology, cape town), and 100 ng – 200 ng of template dna. the pcr profile for amplification for the mitochondrial nd2 gene was 95 °c for 3 min, followed by 35 cycles of (1) 95 °c for 45 s, (2) 50 °c for 45 s and (3) 72 °c for 1 min, and then a final extension at 72 °c for 5 min. the nd2 gene region of 68 individuals was sequenced with the forward primer only, as no ambiguities or sequencing errors were detected. automated sequencing was performed at macrogen inc. (south korea) using an abi 3730xl dna analyser (applied biosystems). statistical analysis sequences were cleaned and trimmed manually to equal lengths using the program seqman 10.2.1 (dnastar, madison, wi, usa) and were aligned using the program clustalx (larkin et al. 2007). nucleotide sequences of the protein-coding nd2 gene were translated to amino acid sequences to check for stop codons and errors using the program dnasp 5.10 (librado & rozas 2009). all sequences were submitted to genbank (accession numbers: kt070326–kt070393). unique haplotypes were identified using dnasp 5.10 (librado & rozas 2009). haplotype diversity (h) and nucleotide diversity (π) were calculated for each location and the overall locations using arlequin ver. 2.000 (schneider, roessli & excoffier 2000) (table 1). parsimony networks were constructed using the program tcs 1.2.1 (clement, posada & crandall 2000).the best fitting model of molecular evolution for the nd2 data was estimated using akaike’s information criterion (burnham & anderson 2002) as implemented in jmodeltest 2 (darriba et al. 2012) and pairwise φst values were calculated in arlequin ver. 2.000 (schneider et al. 2000) to determine the degree of genetic differentiation among locations. model-corrected distances between unique haplotypes were calculated using paup* (swofford 2002). results for the mitochondrial nd2 gene, trn+g (tamura & nei 1993) was the substitution model that best fit the data, with a gamma value of 0.128. analysis of 68 e. motebensis individuals from nine locations resulted in 24 haplotypes defined by 38 variable sites. the overall nucleotide diversity for all tributary locations was low (π = 0.007), with the vanstraatensvlei, lower draaifonteinspruit and below bokkraal waterfall tributary locations having the lowest nucleotide diversity (π = 0.001) (table 1). overall haplotype diversity was high (hd = 0.883), and haplotype diversities at each of the locations ranged from 0.439 (bokkraal tributary) to 1.00 (upper kaaloog tributary) (table 1). the parsimony network revealed that the draaifonteinspruit tributary did not share any of its haplotypes (1, 2 and 3) with other tributary locations (figure 2). there was also minor differentiation among the remaining tributary locations that could indicate disrupted gene flow patterns (figure 2). for example, the vanstraatensvlei tributary did not share any of its haplotypes (18, 20 and 24) with other tributary locations. the kaaloog, bokkraal, ribbokfontein tributaries and the groot marico main stem shared at least a proportion of the remaining haplotypes (figure 2). sequence divergence in the draaifonteinspruit tributary (0% < d < 0.3%) and the vanstraatensvlei tributary (0% < d < 0.2%) was relatively low compared to the range of genetic divergence (0% < d < 2.43%) among the remaining 18 haplotypes (table 2). figure 2: parsimony network with 95% plausible set of mitochondrial nd2 allele connections (numbered circles) constructed with the program tcs 1.2.1 (clement et al. 2000) for enteromius motebensis (see table 1 for location key codes). the size of the circles indicate the relative frequency of haplotypes (total n = 68) (see table 1). table 2: ranges of genetic divergences (%) (above the diagonal), φst values (below the diagonal) for pairwise comparisons for enteromius motebensis from nine populations in the upper groot marico river based on nucleotide diversity two sequences. significant high values were observed in all analyses of genetic structure because of the lack of sharing of haplotypes between draaifonteinspruit and the other tributary locations. however, because of the small sample size, the full extent of diversity at each location may not have been captured. high φst values (> 0.600) were found between the upper kaaloog and other tributary localities (table 2). a similar pattern (φst > 0.600) was found between the middle vanstraatensvlei and six other localities (table 2). low φst values (< 0.200) among some of the locations suggest relatively recent (e.g. before bass introduction) or potentially ongoing gene flow (table 2). discussion enteromius motebensis is a key component of the unique headwater stream biota that characterises the groot marico nfepa, being near-endemic and restricted to the near-pristine headwaters of this vulnerable catchment (engelbrecht & bills 2007; nel et al. 2011). it is directly threatened by invasive largemouth bass m. salmoides (kimberg et al. 2014) and is likely to require conservation action to avoid a decline in its conservation status from vulnerable to endangered. to aid such future efforts, molecular analyses demonstrated the presence of significant genetic structuring among the sampled populations of e. motebensis and consequently identified which populations require conservation prioritisation within the groot marico catchment. the current study demonstrates that e. motebensis locations in the upper groot marico river catchment show relatively recent isolation of at least one location, the draaifonteinspruit tributary, and significant genetic structuring and diversity among some of the remaining locations (vanstraatensvlei tributary). this genetic structuring may be a result of a number of factors such as in-stream barriers, habitat preferences, tributary flow regimes or isolation by distance. furthermore, results indicated that the overall haplotype diversity across locations of e. motebensis was high and haplotype diversities at each location for e. motebensis was relatively high. in the upper groot marico river catchment, the lack of gene flow from the draaifonteinspruit tributary to the remaining populations is potentially the result of a natural waterfall barrier below the confluences of the kaaloog and ribbokfontein tributaries (figure 1). this barrier may interrupt upstream dispersal of e. motebensis from draaifonteinspruit. however, the apparent lack or limited downstream migration from upstream areas to the draaifonteinspruit and vanstraatensvlei tributaries cannot be explained by natural barriers because e. motebensis should be able to move downstream over such obstacles. in addition, the waterfall is only approximately 1 m high. therefore, it is possible that this barrier may be breached during periods of flooding, which would allow upstream migration. while the artificial weir on the draaifonteinspruit can prevent present-day upstream migration of e. motebensis or invasion by m. salmoides, the genetic results are likely a result of more historic processes (vicariance, habitat fragmentation, and drainage alterations etc., hughes et al. 2009). therefore, the high level of genetic structuring between these areas is surprising. enteromius motebensis inhabits shallow slow-flowing pools in small streams (engelbrecht & bills 2007), making it a rhithronic pool specialist typically associated with headwaters of mountain streams. therefore, this habitat preference may limit downstream migration of e. motebensis, a supposition that is supported by kimberg et al. (2014), who found that e. motebensis was most abundant in the upper reaches of the catchment and seldom penetrated into the lower stream reaches where tributary confluences are located. moreover, ellender, woodford and weyl (2015) demonstrated that the eastern cape redfin pseudobarbus afer (peters 1864) resisted displacement during a high-magnitude flood in a headwater stream in the eastern cape. therefore, the possibility that e. motebensis may display similar resistance to downstream movement cannot be excluded as a potential mechanism for restricted gene flow. the vanstraatensvlei and draaifonteinspruit tributaries have a lower flow volume and lower gradient compared to other tributaries in the groot marico river catchment (roux 2010), which may have contributed to disruption of gene flow. the draaifonteinspruit, in particular, has a weak seasonal base flow because of lower dependence on dolomitic groundwater, resulting in naturally intermittent flow near its confluence with the marico river (roux 2010). thus, naturally intermittent connectivity between draaifonteinspruit and the other tributary locations may also have contributed to isolation. it is also possible that isolation by distance played a role in isolating the draaifonteinspruit and vanstraatensvlei populations, and a further investigation to formally test this hypothesis should be conducted. isolation by distance occurs when organisms have limited dispersal ability, which increases genetic differentiation between sites with increasing geographic distance between them (wright 1943). slatkin (1993) further suggested that in highly connected populations, isolation by distance will occur if dispersal distance of an individual is less than the range of the species. therefore, it may be possible that e. motebensis individuals have small home ranges or very strong preference for tributary habitats and are able to maintain their position in the tributaries during flooding periods. conclusion with limited resources, the maintenance of the genetic structuring and high genetic diversity of e. motebensis can be achieved by prioritising a subset of populations. at least three populations from the genetic evidence are necessary to represent most of the detected genetic diversity of the species. however, in order to validate this finding, more than one genetic marker needs to be analysed (nuclear markers and microsatellites), sample sizes of e. motebensis need to be increased and specific adaptations of e. motebensis need to be studied. from the results of this study, it can be seen that the draaifonteinspruit and vanstraatensvlei tributaries have to be secured for e. motebensis because they have unique haplotypes that do not occur elsewhere. this was also evident in populations of the stargazer mountain catfish amphilius uranoscopus (pfeffer 1889) that also occurs in the groot marico catchment (ka van der walt, unpublished data). therefore, among the remaining populations, the kaaloog tributary has the highest genetic diversity of all the e. motebensis populations and will contribute most towards maintaining the detected genetic diversity. micropterus salmoides is present in the dolomitic eye of the kaaloog tributary, which prevents this area from being an effective refuge area for e. motebensis. therefore, removal of these non-native invasive fish, together with local stakeholder education to prevent their re-introduction is the single highest priority conservation action. furthermore, non-native invasive fish management and reduction of other threats to the groot marico river catchment will directly and indirectly help to improve the conservation status of e. motebensis in future, should more resources become available. acknowledgements we thank h. roux and p. kimberg for their support during this project. we also thank j. du plessis (riverstill guest farm) for accommodation, and the many landowners of the groot marico who allowed access to their streams. we thank t. msezane (university of pretoria), a. basson and d. kindler (golder associates) for assistance in the field. we thank a. chakona and g. gouws (south african institute for aquatic biodiversity) for assistance with the genetic analysis. all fieldwork was performed under south african institute for aquatic biodiversity animal ethics guidelines (saiab 2012/03), and research permits were provided by the north west sub-directorate of biodiversity management and conservation (permit no. 042 nw-ii). funding for this research was provided by the water research commission (project k5/2039), with logistical support from the south african institute for aquatic biodiversity, the national research foundation of south africa (uid: 77444), golder associates ltd and department of rural, environment and agricultural development (north west provincial government). competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions this article is based on the master’s thesis of k-a.v.d.w. k-a.v.d.w. was responsible for sample collection, sample processing, laboratory analysis, genetic analysis, interpretation of data and writing the article. e.r.s. played a co-supervisor role. assisted conceptualisation and planning of the research, assisted with genetic analysis, interpretation of data and writing the article. d.w. assisted with sample collection, construction of map, interpretation of data and writing the article. o.w. conceptualised the overall research programme, sourced funding, planned field research, assisted with sample collection, interpretation of data and writing the article. references abell, r., allan, j. & lehner, b., 2007, ‘unlocking the potential of protected areas for freshwaters’, biological conservation 134, 48–63. https://dx.doi.org/10.1016/j.biocon.2006.08.017 aparicio, e., vargas, m.j., olmo, j.m. & de sostoa, a., 2000, ‘decline of native freshwater fishes in a mediterranean watershed on the iberian peninsula: a quantitative assessment’, environmental biology of fishes 59, 11–19. https://dx.doi.org/10.1023/a:1007618517557 burnham, k.p. & anderson, d.r., 2002, model selection and multi-model inferences: a practical information-theoretic approach, 2nd edn., springer, new york. chakona, g., swartz, e.r. & chakona, a., 2015, ‘historical abiotic events or human-aided dispersal: inferring the evolutionary history of a newly discovered galaxiid fish’, ecology and evolution 5, 1369–1380. https://dx.doi.org/10.1002/ece3.1409 clement, m., posada, d. & crandall, k.a., 2000, ‘tcs: a computer program to estimate gene genealogies’, molecular ecology 9, 1657–1659. https://dx.doi.org/10.1046/j.1365-294x.2000.01020.x collen, b., whitton, f., dyer, e., baillie, j.e.m., cumberlidge, n., darwall, w.r.t. et al., 2014, ‘global patterns of freshwater species diversity, threat and endemism’, global ecology and biogeography 23, 40–51. https://dx.doi.org/10.1111/geb.12096 darriba, d., taboada, g.l., doallo, r. & posasa, d., 2012, ‘jmodeltest 2: more models, new heuristics and parallel computing’, nature methods 9, 772. https://dx.doi.org/10.1038/nmeth.2109 dudgeon, d., arthington, a.h., gessner, m.o., kawabata, z., knowler, d.j., levȇque, c. et al., 2006, ‘freshwater biodiversity: importance, threats, status and conservation challenges’, biological reviews 81, 163–182. https://dx.doi.org/10.1017/s1464793105006950 ellender, b.r., weyl, o.l.f. & swartz, e.r., 2011, ‘invasion of headwater stream by non-native fishes in the swartkops river system, south africa’, african zoology 46, 39–46. https://dx.doi.org/10.3377/004.046.0116 ellender, b.r., woodford, d.j. & weyl, o.l.f., 2015, ‘the invisibility of small headwater streams by an emerging invader, clarius gariepinus’, biological invasions 17, 57–61. https://dx.doi.org/10.1007/s10530-014-0744-8 engelbrecht, j. & bills, r., 2007, barbus motebensis in iucn 2013, iucn red list of threatened species, version 2013.2, viewed 05 june 2016, from http://www.iucnredlist.org/ engelbrecht, j.s., 1996, ‘phylogenetic relationships between morphologically similar barbus species, with reference to their taxonomy, distribution and conservation’, phd thesis, rand afrikaans university, johannesburg, south africa. engelbrecht, j.s. & van der bank, f.h., 1994, ‘isozyme and allozyme differences in four shortfin barb (barbus brevipinnis jubb 1966) populations with reference to undescribed species from the transvaal’, biochemical systematics and ecology 22, 269–276. https://dx.doi.org/10.1016/0305-1978(94)90100-7 engelbrecht, j.s. & van der bank, f.h., 1996, ‘genetic relationships between seven species within the chubbyhead and goldie barb groups of minnows (pisces, cyprinidae)’, journal of african zoology 110, 381–396. fausch, k.d., rieman, b.e., dunham, j.b., young, m.k. & peterson, d.p., 2009, ‘invasion versus isolation: trade-offs in managing native salmonids with barriers to upstream movement’, conservation biology 23(4), 859–870. http://dx.doi.org/10.1111/j.1523-1739.2008.01159.x gomi, t., sidle, r.c. & richardson, j.s., 2002, ‘understanding processes and downstream linkages of headwater systems’, bioscience 52(10), 905–916. https://dx.doi.org/10.1641/0006-3568(2002)052[0905:upadlo]2.0.co;2 grobler, d.f., roux, h. & koni, s., 2007, final habitat integrity assessment report of selected rivers in north west province, dace report number: ace 199/06, department of agriculture conservation and environment, mafikeng, south africa. hargrove, j.s., weyl, o.l.f., allen, m.s. & deacon, n.r., 2015, ‘using tournament angler data to rapidly assess the invasion status of alien sport fishes (micropterus spp.) in southern africa’, plos one 10(6):e0130056. http://dx.doi.org/10.137/journal.pone.0130056 hughes, j.m., schmidt, d.j. & finn, d.s., 2009, ‘genes in streams: using dna to understand the movement of freshwater fauna and their riverine habitat’, bioscience 59, 573–583. https://dx.doi.org/10.1525/bio.2009.59.7.8 kimberg, p.k., woodford, d.j., roux, h. & weyl, o.l.f., 2014, ‘species-specific impacts of introduced largemouth bass (micropterus salmoides) in the groot marico freshwater ecosystem priority area’, african journal of aquatic science 39, 451–458. https://dx.doi.org/10.2989/16085914.2014.976169 kindler, d., wagennar, i., & weyl, o.l.f., 2015, ‘an assessment of the reproductive biology of the marico barb barbus motebensis steindachner 1894 from the upper groot marico catchment’, african journal of aquatic science 40, 425–431. https://dx.doi.org/10.2989/16085914.2015.1106400 kocher, t.d., conroy, j.a., mckaye, k.r., stauffer, j.r. & lockwood, s.f., 1995, ‘evolution of nadh dehydrogenase subunit 2 in east african cichlid fish’, molecular phylogenetics and evolution 4, 420–432. https://dx.doi.org/10.1006/mpev.1995.1039 larkin, m.a., blackshields, g., brown, n.p., chenna, r., mcgettigan, p.a., mcwilliam, h. et al., 2007, ‘clustal w and clustal x version 2.0’, bioinfomatics 23, 2947–2948. https://dx.doi.org/10.1093/bioinformatics/btm404 librado, p. & rozas, j., 2009, ‘dnasp v5: software for comprehensive analysis of dna polymorphism data’, bioinformatics 25, 1451–1452. https://dx.doi.org/10.1093/bioinformatics/btp187 lowe, s., browne, m., boudjelas, s. & pe poorter, m., 2000, 100 of the world’s worst invasive alien species: a selection from the global invasive species database, published by the invasive species specialist group (issg), a specialist group of the species survival commission (ssc) of the world conservation union (iucn), viewed n.d., from http://www.gisp.org/publications/brochures/100worst.pdf moyle, p.b. & light, t., 1996, ‘fish invasions in california: do abiotic factors determine success?’, ecology 77(6), 1666–1670. https://dx.doi.org/10.2307/2265770 narum, s.r., 2006, ‘beyond bonferroni: less conservative analyses for conservation genetics’, conservation genetics 7, 783–787. https://dx.doi.org/10.1007/s10592-005-9056-y nel, j.l., driver, a., strydom, w.f., maherry, a., petersen, c., hill, l. et al., 2011, atlas of freshwater ecosystem priority areas in south africa: maps to support sustainable development of water resources, water research commission, wrc report no. tt 500/11, water research commission, pretoria, south africa. rahel, f., 2002, ‘homogenization of freshwater faunas’, annual review of ecology and systematics 33(1), 291–315. https://dx.doi.org/10.1146/annurev.ecolsys.33.010802.150429 roux, h., 2010, ‘a comparison between regional and multivariate classification of biomonitoring sites based on aquatic macroinvertebrate assemblages in the marico river catchment’, phd thesis, university of johannesburg, gauteng. schneider, s., roessli, d. & excoffier, l., 2000, arlequin ver.2.000: a software for population genetic analysis, genetics and biometry laboratory, university of geneva, switzerland. segelbacher, g., cushman, s.a., epperson, b.k., fortin, m-j., francois, f., hardy, o.j. et al. 2010, ‘applications of landscape genetics in conservation biology: concepts and challenges’, conservation genetics 11, 375. https://dx.doi.org/10/1007/s10592.009/0044.5 skelton, e.e. & ribbink, t.-j.v., 1994, the conservation of dolomitic ecosystems in the western transvaal, south africa, j.l.b. smith institute for ichthyology, grahamstown. skelton, p.h., 2001, a complete guide to freshwater fishes of southern africa, southern book publishers (pty) ltd, jlb smith institute of ichthyology, grahamstown, south africa. slatkin, m., 1993, ‘isolation by distance in equilibrium and non-equilibrium populations’, evolution 47, 264–279. https://dx.doi.org/10.2307/2410134 smith-adao, l.b., nel, j.l., roux, d.j., schonegevel, l., hardwick, d., maree, g. et al., 2006, a systematic conservation plan for the freshwater biodiversity of the crocodile (west) and marico water management area, department of water affairs and forestry, csir report no csir/nre/eco/2006/0133/c, csir natural resources and the environment, pretoria, south africa. swofford, d.l., 2002, paup* phylogenetic analysis using parsimony (*and other methods), version 4, sinauer associates, sunderland, ma. tamura, k. & nei, m., 1993, ‘estimation of the number of nucleotide substitutions in the control region of mitochondrial dna in humans and chimpanzees’, molecular biology and evolution 10, 512–526. tweddle, d., bills, r., swartz, e.r., coetzer, w., da costa, l., engelbrecht, j. et al., 2009, ‘the status and distribution of freshwater fishes’, in w.r.t. darwall, k.g. smith, d. tweddle & p.h. skelton (eds.), the status and distribution of freshwater biodiversity in southern africa 3, pp. 21–37, iucn red list of threatened species regional assessment, saiab, grahamstown, south africa. vörösmarty, c.j., mcintyre, p.b., gessner, m.o., dudgeon, d., prusevich, a., green, p. et al., 2010, ‘rivers in crisis: global water insecurity for humans and biodiversity’, nature 467(7315), 555–561. https://dx.doi.org/10.1038/nature09440 wellington, j.h., 1995, southern africa a geographical study, volume 1: physical geography, cambridge university press, london. weyl, p.s.r., de moor, f.c., hill, m.p. & weyl, o.l.f., 2010, ‘the effect of largemouth bass, micropterus salmoides on aquatic macro-invertebrate communities in the wit river, eastern cape, south africa’, african journal of aquatic science 35(3), 273–281. https://dx.doi.org/10.2989/16085914.2010.540776 whittier, t.r. & kincaid, t.m., 1999, ‘introduced fish in the north-eastern usa lakes: regional extent, dominance, and effect in native species richness’, transactions of the american fisheries society 128, 769–783. https://dx.doi.org/10.1577/1548-8659(1999)128<0769:ifinul>2.0.co;2 wright, s., 1943, ‘isolation by distance’, genetics 114, 114–138. filelist convert a pdf file! koedoe 19: j-15 (j976) an outbreak of dermatophilosis in sable hippotragus niger and roan hippotragus eq.uinus in the kruger national park " v. de vas divijion ?fnature conservation pn'vate bag x404 skukuz.a jj50 c. d . imes h lt. col. , usaf, vc, zoonotic pathology divi5ion armed forces institute of pathology walhinglon d.c. , 20j06 ab5t racl a severe skin disease diagnosed as dennatophilosis took on alarming proportions in roan antelope hippotragus equinus and sa ble antelope hippotragw niger herds, which were kept in a 256 ha enclosure in the kruger national park, republi c of south africa. only calves were effected. numerous dmnatophilul organisms were identified in skin biopsies taken from affected areas and examined histologically. complete recovery fo llowed a single treatment of intramuscular injections of a co mbination of a long-acting penicillin and stre ptomycin and the topical application of a preparation which included copper sulphate and benzene hexachloride. factors which probably contributed to the outbreak included prolonged wetness, flies , a high density of susceptible animals and playful butting behaviour of the calves. this is the first report of the occurrence of dennatophilosis as a disease entity in wildlife species in southern africa. 6 the views expressfd he rei71 are those oj the aulhotj a7ld are not to be cqmtrued aj official or as reflecting the ljiews oj the u. s . air foret or the department of dejenu . u tnlljwwl')' ajsignmwt secti071 rif pathology, velen'nary resea rch /mlilu le, onderstepoorl oj 10, republic oj south africa. introduction the distribution of the disease entity known as dermatophilosis (c utaneous streptothricosis, senkobo disease, mycotic dennatilis l among domestic animals such as caule, sheep and horses is world-wide (plowright 1956; austwick 1969; searcy and hulland 1968). the disease was initially reponed in south africa during 1928 from sheep (bekker 19281. since then it has been found that the disease is widespread in south africa among cattle and sheep (steyn 1931; schulz 1956), particularly along the eastern escarpment with its high summer rainfall and greater humidity (rossiter 1969). this area lies adj"acent to south afi·ica '5 greatest wildlife concentration and largest wild ife sanctuaries. yet so far as we aware the disease has not hitheno been described in southern afri can wildlife species. the position is very much the same elsewhere. austwick ( 1969) commented on this phenomenon by pointing oul that the actinomycete dermatophilw congo/emu causes one of the commonest skin diseases of domestic animals and yet few cases have been reponed in wild animals. vague reports from tropical africa indicate that the girailc giraffa came/opardalis, thomson's gazelle ga u lle thormorn· (mackenzie and simpson 1964 ) and a zebra equus sp. (green 1960) have been found to be mildly infected. in temperate regions the di sease has been reponed from the white-tailed deer odocoilew virginia nw (dean, gordon, severinghaus, kroll and reilly 1961; kistner, sholts and green 1970), a cottontail rabbit sylvilagus jloridanw (shotts and ki stn er 1970 ), young chamois ru.picapra rupicapra (nicolet, klinger and fey 1967 ), polar bears rhalarctos maritimw (smith and cordes 1972 ), owl monkeys aotw trivirgatw (king, fraser, garcia, wolf and williamson 1971), a fox vulpes vulpes (austwick 1969 ) and sea lions otaria bryonia (frese and weber 1971) . according to stewart (1972 ) kusel'tan reported infection in hares, hedgehogs and gerbils. the disease is zoonotic and was experimentally produced in man by memery and thiery (according to stewart 1972). natural infection occurred in four men who skinned and handled an infected deer carcass (dean et al. 1961) and one of us (g.d.i.) had a focal, self limiting skin lesion on a finger from careless handling of infc:ctious material. rubel (1972 ) strongly implicated d. congolemu as the cause of pitted keratolysis, a skin condition affecting the fool soles of people in zaire. history the roan antelope hippotragw equ.inw is comparatively rare througho ut its range in africa. this is especially true for the kruger national park (k.n.p ') where there is a small population of 250 to 300 animals causing considerable concern for their chances of survival uouben 1970). it was, therefore, decided to launch an intensive investigation into the bio-ecology of this rare species. to provide realistic study opportunities and simultaneously build up a population from which 2 groups could be released periodically to augment the natura l populati o n, it was decided to erect an invio late enclosure of 256 ha in the most favo urable roan hab itat in the no rthern distric t o f the k.n .p. this encl os ure is hereafter referred to as the camp. a nucleus herd of 10 ro an were released in the camp without mishap in 1967 (pienaar 1968 ; j o ub ert 1970). after promising res ults were achieved initiall y, 13 sab le antelope hippolraguj niger, a lso a rare species in south africa, were placed in th e sa me enclosure during au gust 1969 and may 1970. the respective nucleus herds did exceedingly well initiall y. the roan antelope herd, however , suffered a severe setback in 1970 when 15 out of a to tal o f 27 animals died from anthrax (unpublished data, de vos ). in spite of morta li ty of unkno wn aetio logy which th e yo ung calves of both species suffered sporad icall y, the numbers aga in increased to 48 for sable and 26 for roa n a ntelo pe at the beginning of march 1974 . shortly th ereafter, however, two deaths among young sable antelope ca lves occurred but du e to interfere nce by scavenger birds no worth while specimens for diagnosis were available. it was decided to th o ro ughl y examine rema ining anima ls in the camp in an attempt to detemline the ca uses o f the unexplained deaths. materials and method.s operational the enclosure was entered and the roan and sable herds approached by means of a vehicle. preliminary observations o f the animals o n hoof were made from th e ve hicle. a pair of 10 x 50 binoculars aided in the close'· exa mination o f individuals. affected animals were subseq uentl y caug ht by th e drug immo bilization technique. a pa lmer cap-c hur gaspo~cred rifle was used to pro pel two ml capacity darts or projectill' synnges. to lessen the disturban ce factor the recumbent a nimals were saken by vehi cle to a properl y eq uipped po int of operations o n th e periphery of the camp for closer examination, sampling and treaonent. skin bio psies were taken fro m the worst affected areas of each individua l a nd preserved in milio nig' s 10% buffered fo nnalin . in addition deep skin scrapings were collected in a 50% glycerine/a lco ho l solution . these specimens were processed and examined at a later date. the formalin-fixed tissues were prepared in a routine manner for light microsco py and stain ed with haematoxylin and eosin (he), giemsas and gram's stains and the periodic acid schiff (pas ) reaction. trea ted animals were marked by hairclipping, every individual having its own distinct and easily recog nizable identificatio n mark ·for fulure reference. after resuscitat io n th e animals were re leased in the immediate vicinity of the herd . during 15 to 20 april 1974 12 affected anima ls were ca ught a nd trea ted in this manner. in order to assess the co urse o f the disease without chemotherapeutic s interference, the leas t infected individual was omitted from the initial treatment , but was treated on the second visit to the camp io days afterwards. at the same time one of the previo usly treated indi viduals was caught and re -examined to determine the reaction to therapy. to det ennine whether infection occurred in the immediate surround ings o f the camp, the area was surveyed intensively b y means of binocula rs. the comparative abundance o f ticks inside and o utside the camp was subjectively assessed by merely walking through the veld in the res pe<:tive areas. the amount of ticks attached to the person was taken as an indi cation of abu ndance. climate data were obtained from the meteorological station at the shingwedzi rest camp, situated 40 k.m to the so uth of the enclosure. drugs the immobi lizing drugs and antidote used were : elorphim hydrochloride, m99 1 was used as the basic immobilizing age n t (pienaar 1974 ). azaperom, r19 29' was used as a neuroleptic together with the m99 in the dan combinatio ns (pienaar 1974 ). nalorphine hydrobrol."lide, uthidrone' was used for its ' properties to antago nize th e narco ti c effects of mo rphine or a mo rphine derivative such as m99 (pienaar 1974 ). for th era py o f th e skin co ndition , associated debility and stre ss, a se ri es of drugs and combinatio ns were used . the faculties for which th e drug was choosen, are mentio ned briefly in each case. thioctic acid {tioctan)4 was administered for liver invigoration, det o xification and meta bolic improvement. procaim penicillin g and benetluzmim penidlfin (comprope n )5 was adminis tered in co mbination with dihydro streptomycin (d istrep)6 for its kno wn sy nergisti c action against d . congolemis (roberts 1967; smith and cordes 1972). selenium-vitamin e (bo-self was administered fo r it 's beneficial effect on capture myo pahty as cle imed by basso n and hofmeyr ( 1973 ). hydroxycobalamin ( vit . b12b) (neo cytamen)1 with its prolonged blood level was used for its known beneficial effects in debilitated cases. benum i rrd/ill and som /) d . 1) (l1w(i! pharlflauulica. ' burrollghj ivrllcomt and company. ffuji:mwfl i'hamwuutical co. ud. jglaxo. alittlbuty.l s.a. ( pi} ) lid . 6a. s. rllifr/(lji\'i /jr/ . is.a. ()'llil11midt /..id. 'g/axo·a llrnbrttyj s. ii . ( ply) ltd. 4 hexachloride, s. h.c. diluted in water to a concentration o f 0,0396 of th e gamma isomer was used as a wash for its know n paraciticidal effects and iu claimed beneficial effects o n th e co urse of d ermatophilosis (plowright 1956 ). a dermatological preparation, applied topically was made up of: "healing oi l"9, co ntaining 33,396 vegetable o il, 58,996 o leo resins and 0,596 gennicidal agent as the basis with copper julphale b.p. added to make 5% of th e fina l solution. copper sulpha te was used for its useful antiseptic and potent fungicida l properties. it has been found beneficial in th e trea tment o f dermatophilosis in sheep (moule 1948 ; rossiter 19 56; roberts 1957). finally b. h .c. was added to two concentrations of about 0,025% of the gamma iso mer. 9c(){)p a and nephews sa. ( /)1.1' ) uri . environmental geographica ll y the camp is situated in the northern region of the k. n. p. and occ upi es a 1,6 km area to the north and south of the 22<'47' latitude sou th a nd 3 1° 15' longitude east lines respectively. as is typica l for the rest of th e region the area is rather flat , with an a ltitude of about 390 m above sea level. the area co nsists mainly of a heavy '" '" ... '" • , . , " . • , • no • .. .. m jjasonoj' .. a .. jj .. sonoj ...... "yoiogo li71/h (1' -'.0' porlod) .... .... m. 1 ... ' ....... fi g . 1. h islogram of the monthl y rainfall dislribulion for shingwedzi rest c,ljnp, kruger national park , during the period of and directly prt·, ced ing the outbrea k of dermalophilosis . yearly rainfall figures are includ ed li)r comparat ive purposes. 5 black basaltic soil type and the savannah vegetation type is dominated by th e tree and shrub [ann of the mopane (colopho5permum mopane), hence the designation of this vegetation type as "rnopane scrub and tree sava nnah " (pie naar 1963) . other important tree and shrub species inelude combrelum imberbe, sc/erocarya caffta, lqnchocqrpus capajjo, dalbergia melanoxylon and grewia spp . the grass stratum is mainly co mprised o f themeda lriandra, panicum coloratum, schmidtia pappophoroidej, cmchrw ciljaris, heteropogon contortu.s, digitaria spp. and arislida spp. giving an overall " tall grass" appearance. being at its peak stage of growth and as a result of exceptionally good rains (fig. i), the green grass cover was es pecia l\ y lush during the lime of the disease outbreak. for the greater pan of th e camp th e grass exceeded the sho ulder height of the yo unger , or affected animals. as depicted b y fig. 1, exceptionally heavy rains were experienced during the season preceding the disease outbreak. precipitation in th e form of heavy dew further intensified and prolonged exposure to moisture. the biting ny lipoptena paradoxa was panic ularly abundant and when wo rkin g with the calves these flies would swarm from them and settle o n the handlers. a subjective assessment of tick infestatio n indicated that there were greater numbers o n the vegetation inside than there was o u tside the cam p . initial examination familiar to the occasional proximity of a vehicle within the camp, it was poss ible to a pproach the animals with some caution to within 20 metres. at this distance and with th e aid o f 10 x 50 binoculars it was clear tha t some debilitating skin disease existed which affected the total calf crop, at the time consisting of seven sable and six roan ca lves. the ages ranged from about 2-4 months. they were variably infected , so me being obvio usl y in the early and others in more advanced stages of the di sease. debilitation clearly paralleled the progressive stages of infection . the more severel y infected individuals showed a droopiness of the head and ea rs and frequently shoo k the ir head s as if the lesions were irritant. they often lagged b ehind the rest of the herd . mild lachrymation was show n by the most seve rely infected individuals. binocular examination o f animals outside the camp for infection was also ca rried out, the immediate surrounding area being covered more intensively. no affected animals were detected. immobiliuzlion on 1974 .04.16 catching o peratio ns commenced and by 1974.04 . 19 12 of the 13 affected ca lves were ca ught for close examination, sampling of the sk in lesio ns and treatment. the animals were of uniform size and age a nd a d osage rate of 2 mg m99 and 40 mg azaperone was used throughout. animals went down in three to nine minutes after intramuscalar d eposition of the drugs by cap-chur dart. this was, however, 6 only execuled with the utmost caution and patience. with too much activity around them, o r when hard pressed, the calves often resorted to hiding or laying up behaviour for long periods or even the rest of the day. effective co unteraclion of the immobilizing drugs was achieved with the administration of 100 mg lethidrone (60 mg intravenous and 40 mg intramuscu lar). the calves invariab ly got up within two minut es and joined the herd shortly afterwards. no fatalities occurred. fig. 2. fig. 3. derrnatophilosis. sable ca lf. patchy lesions around eyes and base orcai'. encrus ted hair in lesions over the withers. no te cap ture dart in ri ght hip. dermatophilosis, sab le calf. patchy lesions on cars and heavy enc r ustatiun of mane down to with er s. 7 grojj appearanu the sk.in condition was an ex udative dermatitis characterized by the [onnation o f crusts. lesions ranged in size from small nodu le·li kc fom13tion s to large patches (fig. 2) and in the more advanced stages who le regions of sk.in were covered with heavy encrustations of dry exuda te admixed with hair (fig. g ). in the more severe cases a pumlent serous exudate exuded through cracks in the crust. in the early stages (he crusts were tenacious and on removal the skin beneath would appear moist and hyperaemic with minor haemorrhages in some cases. in the later stages the crusts co uld be removed more easily and in so me instances were o nl y held in place by penetrating hairs. the variety of lesions. or vario us stages of infectivity, made it possible to follow the seq uence of development of the les ions. in the earliest stage • 6 fig . 4. derma tophi los i s. fig. 5. dennatophilosis. • low magnificati on showing layered exudate and acanthosis of epidermis. h ex 30. concent rat ion of organism s around hai r shaft in ('xudall" gicto sa sta in x 500. fig. 6. d . collgoltj js. branchin g and horizo ntal div isio ns. giem sa slain x i zoo . fig. 7. d . ( ollgoltj ij. ho rizontal and transvcrse divisions fanning coccoi d spo res . giemsa stain x izoo . 8 on ly the rorehead and immediate vicini£}, or the eyes were atlected. lesio ns then appeared o n the dorsal aspects o r the head. neck. thorax and lumbar regions. from there it appeared to spread down the sides or the body and on the icgs. the most severe case showed heavy coalesced encrusta ti o ns o r the dorsal aspects or the body, the lateral rib cage behind th e elbow, the sternal aspect and lower parts or th e legs. in this case the hair or the rest o r the body were unnaturally erecl and on palpation revea led ro ughenin g or the skin surface due to numero us small (peasize) isolated wart-li ke elevations. nu mero us immature srages or boophilw ticks were round o n th e animals, some even bei ng imbedded in the lesion encrustations. a rew ambl)'omma and rhipicephalw ticks were also round, mostly co nfined to the anal region. hi jtopathologjcalfinding5 and dtjinitive diag1loju pathological changes were primarily in the epid~rmis and co nsi sted or acamhos is and piling or cornified epithel ium and exudate on the epi dermal surraces in a la yered rash io n (fig. 4. 5, 6 and 7). this layered material contained neutro phils, which in some foci were conce ntra ted in a pustu la r-like fonnation . within the exudate orga nisms were easily see n in h e sta ined sections. they were most plentiful around, but not invading hair sharts (fig. 5) and in foci containing large numb ers of inflamatol)' cells. organisms grew in a mycel ia -like ra shion (fig. 6) and the growth fonnalion was divided both horizontally and longit udinall y form ing cocco id bodies up to 6 or 8 organisms across in the widest portions (fig. 7). they stained well with giemsa's and gram's stains but pas reaction was negative. morphologically and histologicall y they were identified as d. congolmjij. the underlying dermis was infiltrated by neutrophils a nd small mononuclear cells and there was some fibroblastic proliferation. therapy ten days afler treatment (vide supra) a binocular re-examination revea led drastic and a clearcut improvement of previously affected indi viduals. this was further borne o ut by catching and closely examining one of the ca lves which was judged to be severely infected prio r to treatmen t 12 days earli er. scab fonnation had stop ped and the sk in was dl)' and pliable. the overall impression which was ga ined was that th e a nimal was on its wa y to full recovel)'. the disease in the untreated control calf had progressed fro m a limited inrect ion o n the forehead and around the eyes to a severe stage with crustformation o n the head. ears, back, sides, stern um and legs. treatment was instituted at t.his time. as witnessed a month later al animals showed a co mpletc clinical recovel)'. 9 dijcwjion and conciujiom the gross skin lesio ns o f dennatophilosis are essentially th f: same in a ll animal species (sm ith , j o nes and hunt 1972) and those seen in the roa n and sable calves of this report co nfonn with th ose which have been described. the constant finding of dt rmatophiltu organisms in skin lesions therefore warranted a diagnosis of dermatophilosis. the fungallike appearance o f d . congolensij has led to names for the di sease such as mycolic dermatitis and cutaneo us streptothricosis. cordo n ( 1964 ) d isproved its relations hip with fungi and regarded the above na mes as mislead ing impl ying rreaufl ent should be antifungal in stead of antiba cteria l. roberts ( 1967 ) suggested " dennatoph ilus infecti o ns" be used for t he disease and stewart ( 19 72) slates the term "denna top hilosis" is eq ua ll y exp li cab le. jubb and kennedy (1970) expressed the opinion that the older en trenched names, le. cutaneous streptothricos is in ca ttle, mycotoc dennatitis in sheep and strawberry footrol in sheep be retained b ut recent reports of the disease have favo ured derma lophi losis. alth ough th e epizootiologic pattern of dennatop hilosis is not completely und ers too d , some fa ctors contributing to dis se mination of this di sease are kno wn . organisms o f the genus dtrma/.()philuj undergo fragmentation of the mycelia as a result o f multidimensio nal division and give rise to motile zoospores (gordon and edwards 1963; cordon 1964 ; ro berts 196 7). the reselvoir o f d. congoleruij in nat ure is, however, un known. attempts to isolate it from soil have been unsuccessful (roberts 1963a ; kaplan and j o hnsto n 1966 ). roberts (1967) ma inlains tha t chro ni ca ll y affected indi viduals are probab ly the o rga nism's chief mea ns of surv iva l with in a nock or herd during donna m periods. there is a lso evidence that the mi cro-orga nism co ncerned leads a sa prophytic life on th e skin and that it is no t until the equilibrium be[wee n th e paras ite and host is disturbed by extra neous facto rs that it becomes a true pathogen (cord on 1964 ). the mi cro-orga nis m which gave rise to th e present outbreak must therefore have originated from a biological carr ier within th e camp. there has been a rather co nsistem association between wet co nd itions o r hum idity a nd infec ti o n (chod nik 1956; plowright 1956; scarnell 1961; ie ri che 1968 ). in add itio n richard and pier ( 1966) fo und that fli es, co ntaminated with d. congoiemil, infected rabbits more readily il' their skin was first wetted . intense o r prolonged wetting of the skin appa rentl y results in emulsification and di sr uption of th e se baceo us film (roberts 1963b ; robe rts and gra ham 1966) as well as maceration o f the stratum corneum , whi ch then becomes more penneable to irri tant substa nces (suskind and ishihara 1965) and probably more susceptible to di sturbance by mechanical agents (roberts 1967). as depicted by fi gure i exceptionally heavy rains were ex perienced during the seaso n preceding the disease oulbreak. wet co nd itions were furt her augmented by a high humidity with regular ni ghtly precipita ti o ns in the form of heavy dew which nol o nl y dampened individua ls 10 dose to the ground but also thoroughly moistened the t.all grass cove r. this provided almost contin uo us moist cond itions for the young animals and must be considered partially respo nsible for the dramatic differential morbidity in which the ca lves were exclusively affected. however, in australia young animals appear more susceptible to infection than older sheep (nesb it and bannaryne 1955) and in kansas, u.s.a. calves accounted for 33 of 39 cases reponed in callie (kelly, huston, imes and weide 1964 ), roberts (l963b ) maintains that sheep lambs are bo rn with an incomplete sebaceous film that does not become fully protective for a pe riod varying from a few days to five or six weeks. this should not be a faclor in the present outbreak as th e youngest roan and sable calves were at least two months o ld . the lesions of the infection are usually distributed in either of two quite different patterns over the face , ears and body sugge st in g trans fer by insects , or on the li ps and legs suggesting transfer via the pasture (roberts 196 7). in cattle in africa the disease is closely associa ted with tick infestation, and the lesions develop main ly in those ventra l areas of the body most heavily infes ted with ticks (plowright 1956; macadam 1964b ). ticks of the genera amblyomma, boophilus, haemaphysalis, hyalomma, ixodes and rhipicephalus have been incriminated as mechanical vec tors o f thi s dis eas e entity (plowright 1956; vandemaele 1961; macadam 1964a ). the transmission of zoospores by flies has also been shown to occur (roberts 1963a; macadam 1964a ; richard and pier 1966 ; robens and graham 1966). in experiments with rabbit s both musca domeslica and stomoxys ca!citrarn were implicated {richard and pier 1 966 j. in this case three ixodidae parasites were recovered. since boophilus which was the most abundam, is a one-host tick, its potential as a vector wou ld be limited. the three-host ticks, amblyomma and rhipicephalus were not present in large enough numb ers as to be significant in th e dis seminat ion of the disease. the distribution of the initial les ions, being dorsally over the muzzle, head and neck also indicates the importance o f a flying vector rather than ticks. by virtue of their abundance th e biting flies lipoptena paradoxa must be considered important vectors and one of the most significant epizootiological factors of this o utbreak . a wellkn own epizootiological co ncept is that di sease transmission is usually in direct proportion to the density of susceptible individual s. the 26 roan antelope inside the camp provided a density figure of one anima l per 0,1 km 2 as opposed to one an imal per 14 ,5 km 2 for comparable habitat outside. the position for the sable antelope is even more dramatic, there bing o ne animal per 0,05 km 2 within the camp and very few sedentary animals in that particular habitat outside. even th e most favourable sab le amelope habitat in the k.n. p. does not exceed more tha n one anima l per 3,6 km 2 (s. c. j j o ubert, pm . comm . ). this means that both species are lumped together in a restricted space (2,56 km 2) in densit ies which far exceed those for their free-ra nging com peers in the k.n.p. and must therefore. under the specific circumstances, be reii garded as abnonnally high densities. density and probably an associated higher a rthropo d incidence are some factors at variance between the two populations in the same habitat inside and outside the camp. freeroaming individuals (part icul arl y calves) are also in a bener position to seek. o ut areas of shott grass during periods of heavy precipitation, and thus· escape from conditions which are favourable for the development of this di sease. the failure o f finding infection outside might therefore be regarded as significant in this context. the importance o f density as an epizootiological factor is further stressed by findings to th e eltect that the u-ansmi ssio n of dennatophilosi.s ca n occur by contact between sheep, especially if th ey are wet (ie ri che 1968). the behavio ur pauern of young roan and sable calves also afford amp le op portuni ty o f contact between members of a herd. on this aspectjouben (970) wrote : "certainly one of the maj or characteristics in th e compositio n o f the roan herd is the dose relationship betwee n the young animals and especially those of the same age group. young roan genera ll y associated together in a closely knit unit o r nursery in the close proximity of one o r m o re of the adult cows, or at times even seek the co mpan y of the herd bull" . " q.uite frequently even the very young calves interlude the wild running by co nfronting one another o n their knees and after a few seconds of playful butting with their foreheads and pushing they again jump up and co mmence running around. these playfights may be repeated several times between different members during each 'play-session '." this butting behaviour sho uld provide ideal opport unity for transm ission o f zoospores to take place. this theory is funher corroborated b y th e fact that the earliest lesio ns were seen on the forehead and around the eyes. disease manifesta tio ns in roa n and sable calves were very similar to th at which is frequently see n in domesticated stock. without chemotherapeutic interference dea th wo uld undoubtedly have occurred in a hi gh percentage of a ffected an ima ls and probably was the ca use of death of two calves. an occurrence which initiated the investiga tio n. the efficacy o f therapy, as instituted. was further proven by a complete cure after a single treatment. ac4nowltdgemtnts the assistance rend ered during capturing and treatment operations by mr l. e. van rooye n, senior game ranger. kruger national park, and mr p. j. l. bro nkhorst, warden, mountain zebra national park. is gra tefully acknow ledged. 12 references austwick. p. k. c. 1969. mycotic infections. i n mc diarmid, a. (ed ), 1969 . diseases in freeliving wild animals. symp. z.ool. soc. land. 24 :249-271. the zoo logical society of london, london: academic press. 8asson, p. a. and]. m. hofmeyr. 197 3. mortalities associated with wildlife capture operations. in young e. (ed. ), 1973 . the capture and care of wild anima ls. 15 1-160. cape town: human and rous seau. bekker, ] . g . 1928. undescribed skin diseases of sheep in south africa . } i. s . aft. vet . ttlld. all" 1:5 1-57 . chodnik, k. s. 1956 . mycotic dermatitis of ca ttl e in british west afri ca .} . romp. path . tiilr . 66: 179-186. dean , d. ). , m . a. gordon, c. w. seyeringhaus, e. t . kroll and j. r. rei lly, 1961. streptothricosis: a new zoonotic disease. new yorllj. m ed. 61 : 128$--1287. frese, k. and a. weber. 1971. dermatitis in sea li ons (diana bryonia) caused by dmnatophilus congoknsis. ber/. munch . titrdrt1.)l w schr. 84: 50-5 4. go rdo n, m . a. 1964 . the genus dmnatophilw . } . bad . 88 :509--522. go rdo n, m. a. and m . r . edwards. 196.!1 . micromo rphology o f dt rmatophilul congolmju .} . bact. 86: 1101-111 5. green, h. f. 1960. streptothricosis in zebra and m o nkeys and demodec tic mange in e land in ken ya. vet . rec. 72 : 1098 . joubert, s. c. j 19 70. a study of the social behaviour of the roan ante lope. hippotragus equinus equinus (desmarest, 1804 ) in the kruger national park. m .sc. thes is, faculty of science, university of preto ria. j ubb , k. v. f. and p. c . kennedy. 1970. patholotj of domestic ani mals.2. 2nd ed . new york; academic press. kaplan , w. and w. j. johnston . 1966. equine d ermatophilosis (cutaneous streptothricosis ) in georgia. j. am. vet . mtd. au. 149 : 116211 71. kelly, d . c., k. h uston, g. d . imes and k. d. weide. 1964 . cutaneous streptothricosis in kansas cattie. vet . med . 59:7~78, 175-178. kin g, n. w., c. e. d . fraser, f. g. garcia, l . a. wolf and m. e. williamson. 1971. cutaneous streptothricosis (dermatophiliasis) in owl monkeys. lab. anim. sa. 21 :67-74 . kistner , t . p ., e. b. sholts and e. w . greene. 1970. naturally occurring cutaneous streptothricos is in a white tailed d eer (odocoi lew virginianuj). ) . am. vet . mtd. ajs. 151 :633-635. le riche, p. d . 1968. the transmission of dermatophilosis (myco tic . dermatitis ) in sheep. a wl. vtt . } . 44 :64-67 . macadam, i . 1964a . observations on the effects of flies and humidity on the natural lesions of strep tothricosis. vtl. rtc. 76 : 194-198. " macadam , i. 1964b . observations on the effects of ecloparasites and humidity on natura l lesions of streptothricosis . vel. rec. 76:354. macke nzie , p. z. and r. m. simpson. 1964 . the african veterinary handbooa. 4th ed. nairobi : sir isaac pinnan and sons ltd. mo ule, g . r. 194 8. lump y wool of sheep. oj. agric . } . 67 :272-273 . nicolet, j., k. kli n gler and h. fey. 1967. dermatophilw congolen sis, agem de la streptoth ichose du chamois. patholgia microbiol. 30 :831-837 . nisbet, d. 1. and c. c. bannatyne. 1955 . a d e rmatitis of sheep associated with an o rganism of the genus actinomyces. vel. rtc. 6 7: 71 3-715 . pj en aar, u. de v. 1963. t he large mammals of the kruger national pa rk the ir d is tri bution and present-day status. koedot 6 : 1-37 . pi en aar , u. de v. 1968. the use of immobilizing drugs in conservatio n procedures fo r roa n antel o pe. acta 1.001. palh . amve rp . 46 :39-51. pienaar, u. de v. 1974 . the drug immobilization of antel ope species . in young , e. (ed .)' the capture and care oj wild animals . pretoria: human and rousseau. plowright, w. 1956. cutaneous streptothricosis of cattle : i. introdu cti o n and e pizoo ti o lo gical features in nigeria . vel. rec. 68 :350-355 . ri c hard, m. s. a nd a. c. pier. 1966. transmission o f dtrmatophiiw congoleru is b y slomoxys calcil rans and mwca domeslica . am. j . vet. res. 270419-4 23. roberts , d. s. 1957. an ecolo gica l study of th e mycotic dermatilis organi sm . a ils i. vel. }. 33 :23 3--236. roberts , d. s. 1963a . the release and survival of dermatophilw derma lon omus zoospores. aw l. }. agric. res. j4 :386-399. roberts , d. s. 1963b. the influence of carbon di oxide on the growth a nd sporu lation of dermalophilw dermat otwmw . aw l. }. agric. res. 14 : 41 2-423 . roberts , d. s. 196 7. derma toph il m infectio n . vel. bull. 37 :521. roberts , d. s. and n. p. h . graham . 1966 . co ntrol ofovine cutaneous actinom ycos is. awl. vel. } . 42 :74--78 a nd 392-394 . rossiter , l. w. 1956. experiments in the treatment of lumpy woo l. ji. s. afro vel. med. au. 27 : 179-182 . rubel, l. r. 1972 . pitted kerato lysis and dermal ophilw congo/emu . arch . da m. 105 :584--5 86. scarnell, j. 1961. clinical observations on dermatitis of the horse cau sed by "dermatophi lus" species. vet. rec . 73 :795--797. schulz, k. c. a. 1956. mycotic dermatitis {senkobo skindiseasel of callie in the uni o n ofsoulh afri ca . bull. ephoot. dis. aft. 3 :244--261. searcy, g . p. and t. j hulland . 1968 . de rmatophilw dennatitis (streptothri cos is) in ontario. i. clinical observations. can . vet. }. 9 :715 and 16-2 1. shotis , e. b. and t. p. kistner. 1970. naturally occ urring cuta neous stre ptothricosis in a conontail rabbit. } . am. vel. as5 . 15 7: 667-670. 14 smith, c. f. and d. o. cordes. 1972. dermatitis caused by dermatophiluj congolensis inrection in polar bears (thalarcto$ maritimujj br. vet . l. 128,366-371. smith, h . a., t. c.jones and r. d. hunt. 1972. veterinary pathology 4th ed. philadelphia: lea & febiger. steyn, d. g. 1931. investigations into the cause and transmission or lumpy wool affecting merino sheep and its treatment. rep. vel. res. un. s. afr. 17:205-213. stewart, g. h. 1972. dermatophilosis : a skin disease or anima ls and man . vel. rec. 91 :537-544 and 555-561. susk ind, r. r. and m. ishihara. 1965. the effects or wetting or cutaneous vulnerability. arclu. envit. hltlt . ii :529-537. vande rma ele, f. p. 1961. enqucte sur la streptothericose cutanee en afrique. bull. epiwot. dis. aft. 9:251-259. 15 page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 page 14 page 15 filelist convert a pdf file! koedoe 19: 177-178 (1976) additions to the check-list of birds of the addo elephant national park b. l. penzhorn and p. f. van straaten addo elephant national park private bag x6027 port elizabeth 6000 in his original check-list of the birds of the addo elephant national park, liversidge (1965) recorded 120 species. in a subsequent publication six additional species were reported from the park (penzhorn and morris 1969). a further seven species are reported here, increasing the species total for the park to 133. the seven additional species recorded are listed below. the numbers denote those used by mclachlan and liversidge (1957) in roberts birds oj south africa and the nomenclature follows the check list oj the birds oj south africa (s.a.o.s. list committee 1969). 47 white-breasted cormorant phalacrocorax lucidus witborsduiker (lichtenstein) white-breasted cormorants were first reported from caesar's dam by lockhart (1968a) in july 1967. they have subsequently been recorded regularly at the dam. 52 african darter slanghalsvot.l anhinga tufa (lacepede et daudin) three darters were seen at caesar's dam by lockhart (1968b) in july 1967. 133 black eagle witkruisarend aquila verreauxi lesson a single bird was seen soaring above the elephant enclosure in june 1972. this was probably a vagrant from the suurberg range north of the park where black eagles are resident (skead 1967). 343 red-chested cuckoo cuculus solitarius stephens piet-my-vrou red-chested cuckoos have infreqeuntly been heard calling during the early summer months. 352 didric cuckoo diedrikkie chrysococcyx caprius (boddaert) didric cuckoos have frequently been heard calling during the summer months. 177 356 burchell's coucal centro pus superciliosus hemprich vleiloerie and ehrenberg these coucals have been seen among the reeds and rushes at caesar's dam and heard calling at the small dam east of the nature conservator's house. 373 fiery-necked nightjar suid-afrikaanse naguil caprimulgus pectoralis cuvier the calls of these night jars have infrequently been heard on moonlit nights. liversidge (1965) predicted that his total of 120 species of birds would be increased, "particularly in respect to birds of the habitats in and around caesar's dam". of the 13 species subsequently reported, six were recorded at the dam. references liversidge, r. 1965. the birds of the addo national park. koedoe 8: 41-67. lockhart, p. s. 1968a. new distributional data: 1. white-breasted cormorant. ostrich 39 :271. lockhart, p. s. 1968b. new distributional data: 1. african darter. ostrich 39: 2 71. mclachlan, g. r. and r. liversidge. 1957. roberts birds oj south africa. 2nd ed. cape town: trustees of the s.a. bird book fund. penzhorn, b. l. and a. k. morris. 1969.a supplementary checklist of the birds recorded in the addo elephant national park. koedoe 12: 106-10 7. s.a.o.s. list committee. 1969. check list oj the birds oj south africa. cape town: south african ornithological society. skead, c. j. 1967. ecology of birds in the eastern cape province. ostrich suppl. 7: 1-103. 178 page 1 page 2 article information authors: kerry slater1 konrad muller2 affiliations: 1department of environmental sciences, university of south africa, south africa2shamwari game reserve, eastern cape, south africa correspondence to: kerry slater postal address: private bag x6, florida 1710, south africa dates: received: 05 feb. 2013 accepted: 18 sept. 2013 published: 28 feb. 2014 how to cite this article: slater, k. & muller, k., 2014, ‘the diet of brown hyaenas (hyaena brunnea) in shamwari game reserve, eastern cape, south africa’, koedoe 56(1), art. #1143, 5 pages. http://dx.doi.org/10.4102/ koedoe.v56i1.1143 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. the diet of brown hyaenas (hyaena brunnea) in shamwari game reserve, eastern cape, south africa in this short communication... open access • abstract • introduction • research method and design    • study area    • procedure • results • discussion • conclusion • acknowledgements    • competing interests    • authors’ contributions • references • appendix 1 abstract top ↑ brown hyaenas (hyaena brunnea) were introduced to shamwari game reserve in the eastern cape province during 2002, but their feeding ecology is poorly understood. feeding observations of brown hyaena by field guides and the collection of 31 scats from the study area took place over an 11 month period. standard techniques were used to analyse the scats and identify prey items present. ten dietary categories were identified from the scats, with a mean of 3.2 dietary categories per scat. large mammal remains were found in 30 of the 31 scats, with kudu being the most abundant (61.0% of scats). overall the two methods indicated at least 14 mammal species being fed on by the brown hyaena. the presence of mainly large mammal remains and invertebrates (in 38.7% of all scats), together with the feeding observations of mainly large mammals by field guides, suggests that brown hyaena in shamwari are mainly scavengers and that sufficient carrion is available, thereby reducing the need for them to hunt. a 52.0% occurrence of plant matter was found in the scats, suggesting that plant material is an important component of their diet. further studies are underway to investigate the feeding ecology of brown hyaena in shamwari and surrounding areas. conservation implications: comprehensive scat analysis over a number of years, monitoring of individual movement patterns and population numbers of brown hyaena in and around conservation areas will be beneficial in quantifying resource use of this species. introduction top ↑ investigating the diet of free-ranging animals provides information of resources available to and utilised by these animals, but this information is not always available to wildlife area managers because of constraints on time-consuming field work. the diet of a carnivore is a reflection of its feeding type (specialist or opportunistic), the availability of its potential prey and its morphological, behavioural and physiological adaptations (kok & nel 2004). previous studies on brown hyaenas (hyaena brunnea) have shown it to have a diverse diet and, although primarily a scavenger of mammals, it also forages for insects, reptiles, birds and their eggs, and plants (burgener & gusset 2003; kuhn, wiesel & skinner 2008; maude 2005; mills & mills 1978; owens & owens 1978; skinner 1976; skinner & chimimba 2005; skinner & van aarde 1981; van der merwe et al. 2009). despite being generalist feeders, they appear to lean towards specific prey items depending on their habitat. for example, in the southern kalahari, 58 different food types were identified in faecal samples (mills & mills 1978), but along the namib desert coast, the bulk of the brown hyaena’s diet is cape fur seals (arctocephalus pusillus) (goss 1986; kuhn et al. 2008; siegfried 1984; skinner & van aarde 1981). brown hyaenas are inefficient predators and food obtained by hunting is rare (mills & mills 1978). in the southern kalahari, only 4.7% of hunting attempts were successful and those vertebrates hunted successfully made up only 4.2% of the total vertebrate diet (mills 1978). vertebrates hunted were also of small sizes and included birds such as korhaan (eupodotis sp.), small mammals such as springbok lamb (antidorcus marsupilis), springhare (pedetes capensis), striped polecat (ictonyx striatus) and bat-eared fox (otocyon megalotis) (mills 1990). along the namib coast, between 2.9% (goss 1986) and 9.6% (wiesel 2010) of the cape fur seal pups eaten were killed by the brown hyaenas themselves. in areas where brown hyaena ranges overlap with agricultural lands (skinner 1976; skinner & van aarde 1981) or human settlements (kuhn et al. 2008), remains of domestic dogs, cats, cattle, goats, sheep, horses and donkeys (maude 2005) have been found in their faecal scats. brown hyaenas are sparsely distributed over the arid and semi-arid regions of southern africa (mills & hofer 1998). owing to the shy behaviour of this animal, only a handful of papers report on their diet and are mainly restricted to arid areas. there is therefore generally a lack of this information in most of their distribution range. during the early 1990s, the eastern cape province transformed large areas of agriculture to more economically viable game farming, ecotourism and conservation areas (hayward et al. 2007). this change in land use led to numerous wildlife species being reintroduced to former domestic livestock farms. these reintroductions included large predator species that had historically occurred in the eastern cape region, with the main aim of these reintroductions being to restore ecological integrity, conserve threatened species and maximise ecotourism (hayward et al. 2007). although translocation of large carnivores is common, there is often very little or no post-release monitoring (hofmeyer & van dyk 1998). research method and design top ↑ study area this article describes a preliminary study on the diet of brown hyaenas at shamwari game reserve in south africa’s eastern cape province (33°25′40′′s, 26°05′32′′e). shamwari is approximately 25 000 ha, traversing four geological formations (i.e. bokkeveld series shale, witteberg quartzite, karoo sandstone and sundays river formations) (o’brien 2004). vegetation types in shamwari include: kowie thicket, bhisho thornveld, suurberg quartzite fynbos and suurberg shale fynbos (mucina & rutherford 2006). shamwari is situated in the spring-dominant rainfall region of the province and receives about 550 mm of rainfall per annum (low & rebelo 1996; stone, weaver & west 1998). permanent water is available in the semi-perrenial bushman’s river that flows through the reserve for 27.6 km and there are a number of small earth dams. carnivores that have been reintroduced to shamwari include cheetah, leopard, lion, serval, african wild dog and brown hyaena. historical records indicate that brown hyaenas had not been seen in the eastern cape region since the 1960s (hayward et al. 2007). during 2002, two adult male and four adult female brown hyaenas wild caught from northern cape province farms were reintroduced to the reserve and, by 2004, an estimated population of 15 hyaenas could be found in shamwari. during 2007, seven adults and sub-adults were relocated from shamwari to other eastern cape reserves. despite this apparent success of reintroducing brown hyaenas into shamwari, no detailed study on their diet has been conducted. procedure data on feeding observations of brown hyaenas documented by field guides whilst out on game drives with guests were collected between april 2008 and june 2009 (figure 1). these recordings included the date, species being fed on, size and any other related information. thirty-one fresh faecal scats were also collected from 11 latrines throughout the study area between april 2008 and may 2009 (figure 2). dried scats were placed into a nylon netting envelope and rinsed with warm water to loosen the hair and other objects trapped inside. the scats were dissected manually and all hair samples were removed. a representative sample was selected for analysis by inspecting all the hairs visually from each faecal scat and selecting at least one hair for each size, thickness, shape, colour and length (maude 2005). this increased the chances of identifying all mammal species represented by hair in each faecal sample. a scale imprint was made of each hair sample by placing the hair onto a glass slide that was covered with liquid gelatine. once the gelatine had dried, the hair was removed from the slide leaving the imprint of the hair’s scale on the slide. each hair sample was placed into the front section of a pasteur pipette. wax was melted and sucked up into the pipette and then dipped into cold water to solidify the wax. a cross-section of the hair was made by cutting the front end of the pipette into small slices with a surgical blade. the size and shape of the medulla and cortex of each cross-section, together with the scale imprints, were used to identify mammal species represented in each sample from reference material (keogh 1983; maude 2005; perrin & campbell 1980). samples from mammal species occurring in the reserve were also used to make up a reference key (see appendix 1). remains of insects, reptiles, fruit, bone, hair, feathers and any other objects were identified by manually sorting through the faecal scats.dietary diversity (h) was calculated from the scats using the brillouin index and the cumulative diversity (hk) was plotted against the number of scats to determine if the brown hyaenas diet had been adequately sampled (glen & dickman 2006). the percentage occurrence of each prey item or species in the total number of scats was calculated as an indication of how frequently the hyaenas fed on each dietary component (loveridge & macdonald 2003). this was calculated by the number of scats containing a particular food item / total number of scats × 100. the relative percentage occurrence (number of times a specific food item was encountered / total number of occurrences of all food items × 100) indicated the importance of each food item to the overall diet (loveridge & macdonald 2003). a 95% confidence interval for both the percentage occurrence and the relative percentage occurrence were generated from 1000 bootstrap simulations (andheria, karanth & kumar 2007). figure 1: locations of brown hyaena feeding observations by field guides in shamwari game reserve, eastern cape province during the study period, from april 2008 to june 2009. figure 2: locations of brown hyaena latrines used for faecal scat collection in shamwari game reserve, eastern cape province during the study period, from april 2008 to june 2009. results top ↑ eighty-one feeding observations of brown hyaenas were made by field guides during the study period (table 1). fifty-seven (70.00%) of the feeding observations were of brown hyaenas feeding on large mammals, of which 62.00% were of ruminantia or perrisodactyla. during the observations these larger items were always carried away from the carcass to be eaten elsewhere. on one occasion, a female brown hyaena was observed carrying a large piece of a gemsbok for over 3 km during 30 min of observation. twelve observations (14.80%) of brown hyena feeding on two white rhino carcasses that had been killed in a territorial dispute were made during the study period. a giraffe that had died of natural causes was observed being fed on by brown hyaenas eight times (9.88%). of the total observations, 11.00% (five blesbok, one kudu and three springbok) were kills that brown hyaenas had stolen from cheetah. brown hyaenas were also observed to kill and eat one leopard tortoise (geochelone pardalis) and three angulate tortoises (chersina angulata). the use of a cumulative brillouin index (hk) indicated that the scat sample size of 31 scats was sufficient for brown hyaenas (figure 3). we found a mean of 3.2 dietary categories per hyaena scat. large mammal remains (> 10 kg) were found in 30 of the 31 scats and were clearly the most important dietary component for brown hyaenas (table 2). five species of large mammals were identified from the scats, with kudu being the most frequent, representing 61.0% occurrence in the diet. blesbok and bushbuck contributed 58.1% and 41.9% to the brown hyaena’s diet, respectively. plains zebra contributed 29.0% and warthog 12.0% to diet. overall, at least 14 mammal species were found to be included in the hyaena’s diet, as well as a variety insects, reptiles, birds and plant material. table 1: feeding observations of brown hyaenas in shamwari game reserve, eastern cape province, south africa. table 2: diet of brown hyaenas in shamwari game reserve, eastern cape province, south africa, as determined by faecal scats, expressed as the percentage occurrence and relative percentage occurrence. figure 3: species accumulation curve based on the cumulative brillouin index for brown hyaena scat samples in shamwari game reserve, eastern cape province, south africa. discussion top ↑ brown hyaenas usually only hunt small mammals (owens & owens 1978) and therefore the high number of large mammals being observed to be eaten and the hair found in the faecal scats was more than likely indicative of these prey items having been scavenged. this study confirms the findings of other such studies where remains in brown hyaena scats were mainly from members of the order ruminantia (burgener & gusset 2003; van der merwe et al. 2009). the low frequency of small species in this study may suggest that sufficient carrion is available due to the presence of other predators in the same area as the hyaenas, thereby reducing the need for them to hunt. although the occurrence of plant material found in the scats (52%) is not as high as found in brown hyaenas living in arid areas (mills & mills 1978), plant material is still an important component of the diet for brown hyaenas in shamwari. invertebrates that were consumed were more likely ingested during feeding off carcasses. conclusion top ↑ this article represents the first documented study of the dietary habits of brown hyaenas in shamwari game reserve in the eastern cape. further studies are currently underway to explore the feeding ecology and the influence of apex predators on the scavenging behaviour of the brown hyaenas in shamwari game reserve and surrounding areas. acknowledgements top ↑ we thank shamwari game reserve for permission to conduct the project. nadia muller, dr murray stokoe and the shamwari rangers department assisted with logistics of the project. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions k.m. (shamwari game reserve) was responsible for data collection and the initial draft of the article. k.s. (university of south africa) was the project supervisor, analysed the data and wrote the final article. references top ↑ andheria, a.p., karanth, k.u. & kumar, n.s., 2007, ‘diet and prey profiles of three sympatric large carnivores in bandipur tiger reserve, india’, journal of zoology, london 273, 169–175. http://dx.doi.org/10.1111/j.1469-7998.2007.00310.x burgener, n. & gusset, m., 2003, ‘the feeding habits of brown hyaenas (hyaena brunnea) on a game ranch in limpopo province, south africa’, african zoology 38, 181–184. glen, a.s. & dickman, c.r., 2006, ‘diet of the spotted tailed quoll (dasyurus maculatus) in eastern australia: effects of season, sex and size’, journal of zoology, london 269, 241–248. goss, r.a., 1986, ‘the influence of food source on the behavioural ecology of brown hyena in the namib desert’, msc thesis, department of zoology and entomology, university of pretoria. hayward, m.w., adendorff, j., o’brien, j., sholto–douglas, a., bissett, c., moolman, l.c. et al., 2007, ‘the reintroduction of large carnivores to the eastern cape, south africa: an assessment’, oryx 41, 205–214. http://dx.doi.org/10.1017/s0030605307001767 hofmeyer, m. & van dyk, g., 1998, ‘cheetah introductions to two north west parks: case studies from pilanesburg national park and madikwe game reserve’, in b.l. penzhorn (ed.), cheetahs as game ranch animals, pp. 60–71, wildlife group of the south african veterinary association, onderstepoort. keogh, h.j., 1983, ‘a photographic reference system of the microstructure of the hair of southern african bovids’, south african journal of wildlife research 13, 89–132. kok, o.b. & nel, j.a.j., 2004, ‘convergence and divergence in prey of sympatric canids and felids: opportunism or phylogenetic constraint?’, biological journal of the linnaean society 83, 527–538. http://dx.doi.org/10.1111/j.1095-8312.2004.00409.x kuhn, b.f., wiesel, i. & skinner, j.d., 2008, ‘diet of brown hyaenas (parahyaena brunnea) on the namibian coast’, transactions of the royal society south africa 63(2), 150–159. http://dx.doi.org/10.1080/00359190809519219 loveridge, a.j. & macdonald, d.w., 2003, ‘niche separation in sympatric jackals (canis mesomelas and canis adustus)’, journal of zoology, london 259, 143–153. http://dx.doi.org/10.1017/s0952836902003114 low, a.b. & rebelo, a.g., 1996, vegetation of south africa, lesotho, and swaziland, department of environmental affairs and tourism, pretoria. maude, g., 2005, ‘the comparative ecology of the brown hyena (hyaena brunnea) in the makgadikgadi national park and a neighbouring community cattle area in botswana’, msc thesis, department of zoology and entomology, university of pretoria. mills, m.g.l., 1978, ‘foraging behaviour of the brown hyaena (hyaena brunnea thunberg, 1820) in the southern kalahari’, zeitschrift fur tierpsychologie 48, 113–141. http://dx.doi.org/10.1111/j.1439-0310.1978.tb00252.x mills, m.g.l., 1990, kalahari hyenas: the comparative behavioral ecology of two species, chapman & hall, london. mills, m.g.l. & hofer, h., 1998, hyaenas – status survey and conservation action plan, iucn/ssc hyaena specialist group, gland. mills, m.g.l. & mills, e.j., 1978, ‘the diet of the brown hyaena (hyaena brunnea) in the southern kalahari’, koedoe 21, 125–149. http://dx.doi.org/10.4102/koedoe.v21i1.968 mucina, l. & rutherford, m.c. (eds.), 2006, ‘the vegetation of south africa, lesotho and swaziland’, strelitzia 19, south african national biodiversity institute, pretoria. o’brien, j.w., 2004, ‘vegetation management units of shamwari game reserve’, msc thesis, terrestrial ecology research unit, university of port elizabeth. owens, m.j. & owens, d.d., 1978, ‘feeding ecology and its influence on social organization in brown hyaena (hyaena brunnea, thunberg) of the central kalahari desert’, east africa wildlife journal 16, 113–135. perrin, m.r. & campbell, b.s., 1980, ‘key to the mammals of the andries vosloo kudu reserve (eastern cape) based on their hair morphology, for use in predator scat analysis’, south african journal of wildlife research 10, 1–4. siegfried, w.r., 1984, ‘an analysis of faecal pellets of the brown hyaena on the namib coast’, south african journal of zoology 19, 61. skinner, j.d., 1976, ‘ecology of the brown hyaena hyaena brunnea in the transvaal with a distribution map for southern africa’, south african journal of science 72, 262–269. skinner, j.d. & chimimba, c.t., 2005, the mammals of the southern african subregion, 3rd edn., cambridge university press, cambridge. http://dx.doi.org/10.1017/cbo9781107340992 skinner, j.d. & van aarde, r.j., 1981, ‘the distribution and ecology of the brown hyaena hyaena brunnea and spotted hyaena crocuta crocuta in the central namib desert’, madoqua 12, 231–239. stone, a.w., weaver, a.b. & west, w.o., 1998, ‘climate and weather’, in r.a. lubke & i. demoor (eds.), field guide to the eastern and southern cape coasts, pp. 41–49, university of cape town press, cape town. van der merwe, i., tambling, c.j., thorn, m., scott, d.m., yarnell, r.w., green, m. et al., 2009, ‘an assessment of diet overlap of two mesocarnivores in the north west province, south africa’, african zoology 44(2), 288–291. wiesel, i., 2010, ‘killing of cape fur seal (arctocephalus pusillus pusillus) pups by brown hyenas (parahyaena brunnea) at mainland breeding colonies along the coastal namib desert’, acta ethologica 13, 93–100. appendix 1 top ↑ appendix 1: animal species other than brown hyaena that occur on shamwari game reserve. watson.qxd check list of the mammals of tussen-die-riviere provincial nature reserve, free state province, south africa j.p. watson watson, j.p. 2006. check list of the mammals of tussen-die-riviere provincial nature reserve, free state province, south africa. koedoe 49(1): 111–117. pretoria. issn 0075-6458. tussen-die-riviere nature reserve falls within the eastern mixed nama-karoo biome. the surface area is approximately 23 000 ha and it is located in a summer rainfall region. sixty six different mammal species were recorded for the reserve over the past 13 years which constitutes 65 % of the indigenous mammals recorded for the free state province. the highest number of species for the rodents and insectivores were recorded in riverine habitat but the highest abundance were recorded in rocky habitat and was constituted by only one species, aethomys namaquensis. key words: check list, distribution, mammals, conservation area, south africa. j.p. watson, scientific support services, free state department of tourism, environmental and economic affairs, private bag x20801, bloemfontein, 9300 republic of south africa. (watsonj @dteea.fs.gov.za). issn 0075-6458 111 koedoe 49/1 (2006) introduction species lists of occurrence or inventories of species on nature reserves, national parks or specific conservation areas have been published extensively in the past (rautenbach 1976; nel et al. 1980; pienaar et al. 1980; rautenbach 1982; lynch 1983; lynch 1989; lynch & watson 1990; lynch 1994; avenant 1997; avenant 2000a). this refined the knowledge of the distribution of mammal species in south africa. this information formed part of the reference base when the status of species and distribution delineations were determined for the biodiversity of south africa. very few provincial reserves in the free state (fs) have published mammal inventories (rautenbach 1976; avenant 1997; avenant 2000b; avenant & watson 2002; wandrag et al. 2002). this is one of the reasons why the presence of some species in the fs were omitted or not properly documented during the compilation of the recent red data book of the mammals of south africa (friedman & daly 2004). lynch (1983) conducted a survey of the mammals in the free state but never sampled any of the provincial or national conservation areas during this survey. all the effort was focused on private farms, although sampling did occur adjacent to some provincial reserves. the intention of this paper is to publish data that has been collected over the past 13 years at tussen-die-riviere nature reserve (tdr) in the free state and hopefully it will contribute to the distribution of information on mammals in central south africa. study area tussen-die-riviere nature reserve is situated in the southern free state province of south africa, between the caledon and orange rivers, with the western boundary at the confluence of the two rivers (26º03'–26º 20'e, 30º25'–30º35's). the reserve was established in 1967 and proclaimed as a nature reserve in 1972 (van aarde 1984; vrahimis 1991). altitude varies between 1150–1480 m above sea level. the absolute maximum and minimum temperatures recorded are 38.3 ºc and -9.3 ºc respectively, with a mean annual precipitation of 441 mm (vrahimis 1991). tussen-die-riviere is about 23 000 ha in size, consisting of dolomite koppies with domiwatson.qxd 2006/04/17 11:02 pm page 111 nant tree species being olea, rhus, celtis and maytenus. the grass plains consist of themeda, aristida, cymbopogon, eragrostis, chrysocoma, lesertia, osteospermum, salsola and nestlera species. the two major rivers, the orange and caledon, that runs through the reserve are fringed by species such as diospyros, phragmitis, salix babalonica, acacia karoo, lycium and celtis. according to acocks (1988), it falls within the false upper karoo (vegetation type 300), but low & rebelo (1996) relegates it to eastern mixed nama-karoo (vegetation type 52). originally tdr was utilised exclusively as a hunting venue but was also managed as a conservation area where a number of artiodactyla and perissodactyla were introduced into the reserve over the years. hunting formed part of the game management actions of the reserve. recently the function of tdr changed from primarily being a hunting venue to a conservation area and the management plan was adjusted accordingly. materials and methods recording of mammal occurrence data has been conducted since 1992. small mammal surveys were carried out on various occasions utilising sherman type live traps, snap traps (museum specials) and catching specimens by hand. a general bait, consisting of peanut butter, rolled oats, sunflower oil and beef extract (bovril) was used to bait traps. traps were checked and re-baited at sunrise and sunset and the specimens collected were either released after identification or in cases where animals were sampled the specimens were deposited in the mammal collection of the national museum in bloemfontein. the trap success of rodents and insectivores caught in traps was calculated following the method described by rowe-rowe & meester (1982). the following habitat types were sampled: rocky ridges, riverine vegetation, grass/ shrub land, and old buildings. in the case of the latter, traps were set outside the buildings next to walls in vegetation. macro mist nets were used to collect bats at night. likely sites were investigated during the day and bats were collected by hand or shot with .22 dust shot. sight records of various species were taken during game drives, night drives with a spotlight, and during culling operations. other signs like quills, tracks, scats and burrows were also used to determine the presence of some species on the reserve. sightings recorded by the reserve personnel were incorporated in the species lists. the nomenclature followed in this manuscript is according to bronner et al. (2003). results and discussion according to hilton-taylor & le roux (1989) the nama-karoo and succulentkaroo are two of the most inadequately protected biomes in south africa. the namakaroo, in which tdr is located, is particularly inadequately protected and of concern (gelderblom & bronner 1995). tussen-dieriviere has grassland elements present which places it in the transitional zone between the nama-karoo and grassland biomes. gelderblom et al. (1995) are of the opinion that the southern free state is an unprotected hotspot with high species richness, endemic and red data species and is a conservation priority area for carnivores. augmentation of the conservation area network in this region will protect the maximum number of species within a minimum land surface (gelderblom et al. 1995). apart from the alien invasive fallow deer cervus dama (nature conservation ordinance no. 8 of 1969) which entered the reserve by dispersal from neighbouring farms, only one species of the 66 recorded species for tdr, viz., impala aepyceros melampus, occurs extralimitly in tdr (plug & badenhorst 2001). this species was originally introduced for hunting purposes. the species composition (table 1) on tdr represents 65 % of the 101 indigenous mammals for the free state (lynch 1983; watson 1990a; watson 1990b; watson 1998; avenant & watson 2002; ferreira & avenant 2003). the chacma baboon papio hamadryas ursinus is shown as being introduced in table 1, although this species did occur originally in the region. baboons that were used in a behavioural study by a student were released koedoe 49/1 (2006) 112 issn 0075-6458 watson.qxd 2006/04/17 11:02 pm page 112 issn 0075-6458 113 koedoe 49/1 (2006) into the resident troop at tdr in the early 1970s after completion of the project. the study animals did not originate from this region but were animals captured at loskop dam nature reserve in mpumalanga. there is evidence that two genetically different groups of baboons occur in south africa, a northern and southern group (friedman & daly 2004; bloomer pers. comm.). the result is that the troop on tdr might have been contaminated with animals from the northern group. fallow deer, an exotic or alien invasive species (nature conservation ordinance no 8 of 1969), occasionally enter tdr from neighbouring farms. the tdr personnel cull these animals whenever this species is encountered. re-entry from adjacent farms does, however, occur periodically and it is a continuous process to remove these animals, underlining the importance of regulating the importation of alien species into any area. ferreira & avenant (2003) recorded otomys saundersiae or saunders’ vlei rat, dendromus melanotis the grey climbing mouse, tatera leucogaster the bushveld gerbil, mystromys albicaudatus the white tailed rat and crocidura cyanea the reddish-grey musk shrew at the reserve. four other species, crocidura fuscomurina the tiny musk shrew (dd), desmodillus auricularis the short tailed gerbil, ichneumia albicauda the whitetailed mongoose and lutra maculicollis the spotted necked otter (nt) were recorded by lynch (1983) in close proximity to reserve but these species have not to date been recorded on the reserve. of these four species, one is near threatened (nt) and one is data deficient (dd), resulting in two of the four species listed in a red data category as defined by friedman & daly (2004). it is important to establish whether these species occur on the reserve, and management practices should be of such a nature as to be able to accommodate these species. most of the above-mentioned four species are not common, occur in low numbers or are very secretive and difficult to observe or capture. rodentia the rocky outcrop habitat offered the lowest variety of species. only one species, aethomys namaquensis, was trapped although sightings of the insectivores elephantulus myurus, were made while working in the veld. none were successfully trapped. even with the lowest species variety this habitat had the highest trap success (table 2). aethomys namaquensis is a common and abundant species in rocky habitat in the free state. high trap success percentages are usually experienced when trapping in rocky habitat. old buildings gave the second highest trap success with only two species present, viz., mastomys (sensu lato) and graphiurus murinus. the mastomys (sensu lato) was trapped in only two habitats; the other being riverine vegetation. this species is a generalist and its presence is also indicative of habitat disturbance. graphiurus murinus was only captured next to an old building in the spes bona area of the reserve (table 2). these animals are, to a large extent, arboreal and old buildings have shelter and food plants, such as diospyros lyciodes, growing along the walls. traps were set for this species where eaten fruits were detected at the old buildings. factors such as shelter, food and signs of their presence enhanced the trappabilty of this species at the old buildings although they do mostly occur in rocky habitats, or in big trees with holes in the trunks and branches (skinner & smithers 1990). grass and shrub plains had the lowest trap success. only two species were recorded, i.e., aethomys namaquensis and saccostomys campestris (table 2). the former is usually confined to rocky areas and the inclusion of the species in this habitat is due to the fact that traplines were set too close to rocky areas and an ecotone effect manifested in this species being trapped in grass/ shrub plains. the pouched mouse, saccostomys campestris, is catholic, occupying a variety of vegetation and climate types and is widely distributed (skinner & smithers 1990). lynch (1983) shows this species to occur only in the watson.qxd 2006/04/17 11:02 pm page 113 koedoe 49/1 (2006) 114 issn 0075-6458 table 1 mammals recorded in tussen-die-riviere 1 = re-introduced, 2 = listed red data species (friedman & daly 2004). 3 = exotic. dd = data deficient; nt = near threatened; vu = vulnerable. scientific and common names are according to bronner et al. (2003) scientific name common name status observed signs/ tdr in surveys personnel insectivora 2 suncus varilla lesser dwarf shrew dd x 2 atelerix frontalis south african hedgehog nt x 2 crocidura cyanea reddish-grey musk shrew dd chiroptera rhinolophus clivosus geoffroy’s horseshoe bat x neuromicia capensis cape serotine bat x tadarida aegyptiaca egyptian free-tailed bat x primates 1 papio hamadryas ursinus chacma baboon x cercopithecus pygerythrus vervet monkey x carnivora vulpes chama cape fox x canis mesomelas black-backed jackal x aonyx capensis african clawless otter x ictonyx striatus striped polecat x 2 poecilogale albinucha striped weasel dd x genetta genetta small-spotted genet x suricata suricatta suricate x cynictis penicillata yellow mongoose x galerella pulverulenta cape grey mongoose x atilax paludinosus marsh mongoose x proteles cristatus aardwolf x 2 parahyaena brunnea brown hyaena nt x caracal caracal caracal x felis silvestris african wild cat x felis nigripes black footed cat x otocyon megalotis bat-eared fox x panthera pardus leopard x perissodactyla 1 ceratotherium simum white rhinoceros x 2,1 diceros bicornis minor black rhinoceros vu x 1 equus quagga plains zebra x hyracoidea procavia capensis rock hyrax x tubulidentata orycteropus afer aardvark x artiodactyla 1 phacochoerus africanus common warthog x 1 connochaetus gnou black wildebeest x 1 alcelaphus buselaphus red hartebeest x 1 damaliscus pygargus phillipsi blesbok x sylvicapra grimmia common duiker x antidorcas marsupialis springbok x rhaphicerus campestris steenbok x 1 aepyceros melampus impala x 1 oryx gazella gemsbok x 1 syncerus caffer african buffalo x 1 tragelaphus strepsiceros greater kudu x watson.qxd 2006/04/17 11:02 pm page 114 extreme southern free state province and is of the opinion that they are absent from the rest of the province. this species has however been recorded at sandveld nature reserve in the western free state (avenant & watson 2002). next to the drainage lines where riverine vegetation was present, the highest number of species (5) were recorded. the two most common species were generalists, rhabdomys pumilio and mastomys (sensu lato). these two species usually have wide habitat requirements and are catholic. the high species variety is probably due to the greater number of micro habitats that are available, more diverse food range and the denser vegetation cover in these areas. otomys irroratus, a specialist and vegetarian, was only collected in the riverine areas as this species tends to keep to areas with firstly dense vegetation cover and secondly, areas associated with higher moisture which also promotes plant production (table 2). conclusion as mentioned above, tussen-die-riviere nature reserve has 66 recorded mammal species present. a number of these species are endemic to south africa and some are listed in red data book categories (friedissn 0075-6458 115 koedoe 49/1 (2006) table 1 (continued) scientific name common name status observed signs/ tdr in surveys personnel 1 tragelaphus oryx eland x redunca fulvorufula mountain reedbuck x 1 redunca arundinum southern (common) reedbuck x 3 cervus dama fallow deer x rodentia xerus inauris south african ground squirrel x pedetes capensis springhare x hystrix africaeaustralis cape porcupine x cryptomys hottentotus african mole-rat x otomys irroratus vlei rat x otomys saundersiae saunders’ vlei rat 2 mystromys albicaudatus white-tailed rat en saccostomus campestris pouched mouse x malacothrix typica large eared mouse x dendromus melanotis gey climbing mouse rhabdomys pumilio four-stripe grass mouse x mus minutoides (sensu lato) pygmy mouse x mastomys (sensu lato) multimammate mouse x aethomys namaquensis namaqua rock mouse x grahiurus murinus woodland dormouse x tatera brantsii highveld gerbil x tatera leucogaster bushveld gerbil lagomorpha lepus saxatilis scrub hare x lepus capensis cape hare x pronolagus rupestris smith’s red rock rabbit x macroscelidea elephantulus myurus rock elephant shrew x species in bold are records from ferreira & avenant (2003). watson.qxd 2006/04/17 11:02 pm page 115 man & daly 2004). tussen-die-riviere nature reserve falls within the transitional zone between the nama-karoo and grassland biomes and is an important area for the conservation of south african mammals. the reserve supports a high species richness, including some endemics and is pointed out as a priority area for carnivores (gelderblom et al. 1995). furthermore, the reserve forms an integral part of the conservation network in this region. this also supports the recommendation by gelderblom et al. (1995) that maximum conservation effort with minimum land surface can be achieved in this region. this should be investigated by the various adjoining provinces and an effort must be made to increase the surface area of the conservation areas in this biome. the formation of a trans-provincial park, linking all the nature reserves and having a jointly-managed management plan for the whole area will also enhance the biodiversity conservation effort in this biome. acknowledgements gratitude is extended to the free state department of tourism, environmental and economic affairs for the time and funds that were available to collect the data and publish the results. further gratitude is extended to all the reserve personnel at tussen-dieriviere who contributed with observations. the recommendations of dr naas rautenbach, savvas vrahimis and dr. pierre nel on the manuscript are appreciated and incorporated in the document. references acocks, j.p.h. 1988. veld types of south africa, 3rd edn. memoirs of the botanical survey of south africa 57: 1-146. avenant, n.l. 1997. mammals recorded in qwaqwa national park (1994-1995). koedoe 40(1): 31-40. avenant, n.l. 2000a. terrestrial small-mammal diversity in koranaberg conservancy, free state, south afrika. navorsinge van die nasionale museum, bloemfontein 16(4): 69-82. avenant, n.l. 2000b. small mammal community characteristics as indicators of ecological disturbance in the willem pretorius nature reserve, free state, south afrika. south african journal of wildlife research 30(1): 26-33. avenant, n.l. & j.p. watson. 2002. mammals of sandveld nature reserve, free state province, south africa. navorsinge van die nasionale museum, bloemfontein 18(1): 1-12. bronner, g.n., m. hoffmann, p.j. taylor, c.t. chimimba, p.b. best, c.a. mathee & t.j. robinson. 2003. a revised systematic checklist of the extant mammals of the southern african subregion. durban museum novitates 28: 56 106. ferreira, s.m. & n.l. avenant. 2003. modelling the effects of trap-spacing on small mammal community descriptors in grasslands at tussendie-riviere nature reserve, free state province, koedoe 49/1 (2006) 116 issn 0075-6458 table 2 species variety and trap success for the various habitats sampled in tussen-die-riviere nature reserve, free state province, south africa habitat nights trap number of individuals (percentage trap-success) total no. individuals aethomys saccostomus rhabdomys mastomys otomys mus graphiurus and namaquensis campestris pumilio sensu lato irroratus minitoides murinus trap-success rocky outcrop 120 5 (4.2%) 5 (4.2%) grass/ shrub plains 280 2 (0.7%) 4 (1.4%) 6 (2.1%) riverine 640 4 (0.6%) 7 (1.1%) 7 (1.1%) 1 (0.2%) 1 (0.2%) 20 (3.2%) old shed 140 3 (2.1%) 2 (1.4%) 5 (3.5%) total 1180 7 (0.6%) 8 (0.7%) 7 (0.6%) 10 (0.9%) 1 (0.08%) 1 (0.08%) 2 (0.2%) 36 (3.1%) watson.qxd 2006/04/17 11:02 pm page 116 south africa. navorsinge van die nasionale museum, bloemfontein 19(2): 21-30. friedman, y. & b. daly (eds.). 2004. red data book of the mammals of southern africa: a conservation assessment. cbsg soutern africa, conservation breeding specialist group (ssc/iucn), endangered wildlife trust. south africa. gelderblom, c.m. & g.n. bronner. 1995. patterns of distribution and current protection status of the endemic mammals in south africa. south african journal of zoology 30(3): 127-135. gelderblom, c.m., g.n. bronner, a.t. lombard & p.j. taylor. 1995. patterns of distribution and current protection status of the carnivora, chiroptera and insectivora. south african journal of zoology 30(3): 103-114. hilton-taylor, c. & a. le roux. 1989. conservation status of the fynbos and karoo biomes. pp. 202-223. in: huntley, b.j. (ed.). biotic diversity in southern africa. cape town: oxford university press. low, a.b. & a.g. rebelo. 1996. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. lynch, c.d. 1983. mammals of the orange free state. memoirs van die nasionale museum, bloemfontein 18: 1-218. lynch, c.d. 1989. the mammals of the north-eastern cape province. memoirs van die nasionale museum, bloemfontein 25: 1-116. lynch, c.d. 1994. the mammals of lesotho. navorsinge van die nasionale museum, bloemfontein 10(4): 177-241. lynch, c.d. & j.p. watson. 1990. mammals of sehlabathebe national park, lesotho. navorsinge van die nasionale museum, bloemfontein 6(12): 523-554. nel, j.a.j., i.l. rautenbach & j. breytenbach. 1980. mammals of the kammanasie mountains, southern cape. south african journal of zoology 15: 225-261. pienaar, u. de v., i.l. rautenbach, & g. de graaff. 1980. the small mammals of the kruger national park (a checklist and atlas). pretoria: national parks board of trustees. plug, i. & s. badenhorst. 2001. the distribution of macromammals in southern africa over the past 30 000 years. transvaal museum monograph 12: 1-234. rautenbach, i.l. 1976. a survey of the mammals occurring in the golden gate highlands national park. koedoe 19: 133-144. rautenbach, i.l. 1982. mammals of the transvaal. ecoplan monograph no. 1: 1-211. rowe-rowe, d.t. & j. meester. 1982. habitat preferences and abundance relations of small mammals in the natal drakensberg. south african journal of zoology 17: 202-209. skinner, j.d. & r.h.n. smithers. 1990. the mammals of the southern african subregion. pretoria: university of pretoria, south africa. south africa. 1969. nature conservation ordinance no. 8 of 1969. pretoria: government printer. van aarde, r.j. 1984. reproduction in the porcupine hystrix africae-australis peters. d.sc. thesis, univ. pretoria, pretoria. vrahimis, s. 1991. activity patterns of black wildebeest (connochaetes gnou zimmerman, 1780) in the orange free state. m.sc. thesis, university of the orange free state. wandrag, g.f., j.p. watson & n.b. collins. 2002. rodent and insectivore species diversity of seekoeivlei provincial nature reserve, free state, south africa. south african journal of wildlife research 32(2): 137-143. watson, j.p. 1990a. westward range extention of temminck’s hairy bat in south africa and lesotho. south african journal of wildlife research 20(3): 119-121. watson, j.p. 1990.b. new distribution records of laephotis in southern africa and lesotho. navorsinge van die nasionale museum, bloemfontein 7(4): 61-70. watson j.p. 1998. new distributional records for three microchiropteran bats (vespertillionidae, rhinolophidae) from the free state province, south africa. south african journal of wildlife research 28(4):127-131. issn 0075-6458 117 koedoe 49/1 (2006) watson.qxd 2006/04/17 11:02 pm page 117 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice article information author: ishbel cullen1 affiliation: 1fenner school of environment and society, australian national university, australia correspondence to: ishbel cullen email: u5006844@anu.edu.au postal address: 54 bonython st, downer, act, australia how to cite this article: cullen, i., 2015, ‘world parks congress 2014: parks, people and planet’, koedoe 57(1), art. #1338, 3 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1338 note: ishbel cullen is an honours student at the fenner school of environment and society, australian national university. she was a volunteer at the world parks congress in sydney. copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. world parks congress 2014: parks, people and planet in this commentary... open access • introduction • parks • people • planet • conclusion • acknowledgements    • competing interests introduction top ↑ the 6th international union for conservation of nature (iucn) world parks congress took place in november 2014 in sydney, australia (figure 1 and 2). this once-in-a-decade event is the flagship global forum on protected areas and the conservation of nature. for seven days, sydney olympic park was teeming with over 6000 park rangers, conservation scientists, politicians, government officials, business people and concerned citizens from 170 countries. they came together to share their experience and ideas about the challenges and opportunities facing the global protected area network. figure 1: pavilions established at the international union for conservation of nature world parks congress 2014. source: photograph taken by ishbel cullen figure 2: leaders of pacific island nations make a commitment to expand marine protected areas at the international union for conservation of nature world parks congress 2014. source: photograph taken by ishbel cullen the breadth of issues addressed at the congress was impressive. each day was structured around eight themes: reaching conservation goals; responding to climate change; improving health and well-being; supporting human life; reconciling development challenges; enhancing diversity and quality of governance; respecting indigenous and traditional knowledge and culture; and inspiring a new generation. these different themes involved discussions that spanned all scales of activity, from removal techniques for individual invasive species to updates on international legal frameworks. given this overwhelming amount of content, it was not possible for one individual to engage in all the workshops, dialogues and plenaries the conference had on offer. this article is a reflection of the sessions that were attended by the author and how they were relevant to the catchphrase of the congress: ‘parks, people, planet: inspiring solutions’. parks top ↑ preserving natural ecosystems in protected areas or parks is the main international strategy to conserve biodiversity. sustained efforts to increase the coverage of legally recognised protected areas have resulted in significant gains in recent years. the united nations environment programme protected planet report 2014, which was released at the congress, outlined that, since 2010, 6.1 million km2 of new protected areas have been created. this is close to the size of the entire australian continent and means that we are on track to meet target 11 of the convention on biological diversity aichi biodiversity targets, which aims for 17% of terrestrial and inland water areas to be managed for conservation by 2020. though encouraging, this increase in protected area coverage is yet to halt global biodiversity loss. species decline and rates of extinction continue to accelerate. according to the convention on biological diversity's global biodiversity outlook 4, published in 2014, this may in part be due to time lags in environmental responses to increased conservation activity. however, this disconnect is rooted in broader pressures including habitat destruction, pollution, climate change, over-exploitation and invasion by exotic species. much more needs to be done to reduce and counteract these pressures inside and outside formal protected areas if global biodiversity loss is to be arrested. a major focus of the congress was identifying new and adaptive methods to enhance conservation outcomes in light of these pressures. one of the discussions focused on linking natural assets with the ‘real economy’ as a way to prevent further degradation and generate the funds required to scale up conservation efforts. achim steiner, executive director of the united nations environment programme, stressed the importance of conservationists becoming economically literate by using the concept of ‘natural capital’ and putting a price on the value of ecosystem services. members of the finance and business community spoke about trends in this direction, through continued development of markets for environmental goods such as carbon and biodiversity. the move towards integrating natural capital could help address the fundamental problem of environmental externalities in the current economic system and generate funds for conservation through payment for ecosystem services. representatives from the world wide fund for nature and credit suisse presented their 2014 report on conservation finance at the congress, showing that investment in biodiversity conservation needs to be around $200 – $300 billion globally per year to adequately protect the world's most important ecosystems. while this figure is 20–30 times the current rate of investment, it represents only 1% of the total private sector's annual investment on a global scale. the speakers argued for increased involvement by the private sector in order to complement existing conservation finance from public and philanthropic channels. this will require creating opportunities for conservation activities that can generate returns for investors as well as protect biodiversity. people top ↑ the contribution of protected areas to human well-being was widely acknowledged at the congress, with many sessions relating to the physical, psychological and cultural benefits of parks and how these interactions can be used to realise positive outcomes for conservation as well as human populations. the opportunities for indigenous peoples to achieve economic, social and cultural benefits through the management of protected areas was another area of focus, with a strong representation of indigenous communities at the congress. there are many relevant examples from australia, where there are 51 declared indigenous protected areas covering over 36 million hectares. for instance, managing fire regimes in the savannah country of the northern territory has resulted in carbon credits that the traditional owners can sell to generate revenue. monitoring wildlife and removing marine debris has provided opportunities for employment for many indigenous peoples in the sea country of north queensland. these examples demonstrate the potential synergies and co-benefits of biodiversity conservation management. a recent iucn report, urban protected areas: profiles and best practice guidelines (2014), was presented by lead author dr ted trzyna. he explained that urban protected areas are accessible to a large number of visitors from diverse backgrounds and therefore play an important political role in building urban constituencies that value conservation. by researching urban protected areas around the world, dr trzyna and his team have collected a number of important observations and lessons. not surprisingly, urban areas face the most pressure from development, pollution and high levels of use. however, there are also many positive reports. in hong kong, a remarkable 40% of land is in protected parklands. in taipei, yangmingshan national park has an extraordinary volunteering culture, with over 700 000 hours of volunteer work dedicated per year. in san francisco, the parks conservancy, a nonprofit organisation, collaborates with the golden gate national parks to deliver extensive education programmes, with a focus on climate change education. instead of talking about climate change politics or predictions, they are highlighting changes currently occurring in the park to make climate change real and accessible. whilst there are important opportunities for mutually beneficial outcomes for both human populations and biodiversity conservation, in many contexts exploitation and degradation remain dominant. as with most discussions on natural resource management, at the congress notions of balancing resource use and resource preservation were ubiquitous. a pragmatic approach to achieving conservation outcomes often involves a compromise. however, ‘balancing’ terminology is frequently used to give the impression of measured consideration, but in reality it is used to justify poor environmental outcomes. with many politicians and representatives from extractive industries present at the congress, it was important to question how decisions were balanced, by whom and according to what assumptions. this was clearly demonstrated in a session discussing biodiversity offset policies (figure 3). in some contexts biodiversity offsets do represent an opportunity to balance human needs and achieve positive outcomes for biodiversity. for example, where the status quo would see continued degradation and loss of habitat, offsetting provides opportunities for net biodiversity benefits. however, biodiversity offsets can be misused, resulting in a net loss of protected habitat. for example, one speaker identified the potential for biodiversity offsets to resource existing commitments to protected areas, which runs counter to the principle of additionality. figure 3: congress delegates discuss biodiversity offsets and their application in different countries. source: photograph taken by ishbel cullen the emphasis at the congress on integrating human needs and biodiversity conservation was progressive, and accurately represented current realities. however, some cases lacked caution and neglected the importance of ‘no-go’ areas exclusively dedicated to the conservation of nature. planet top ↑ while most direct threats to biodiversity occur at a local scale, the ultimate affliction on nature is now occurring on a global scale, with continued global warming threatening climate stability. this overwhelming problem, with its varied and pervasive impacts, will likely define the coming century for humanity as well as the managers of biodiversity conservation. at the congress there was a climate change stream; however, given its universal effects, it needed to be integrated into all areas of discussion. this separation reflects the genuine challenge of reconciling the temporal and spatial scales of climate change in planning and management today. the challenge for all conservationists is how to focus on immediate and local conservation goals whilst considering, and acting upon, the bigger-picture issues. whilst maintaining this perspective is difficult, it is essential, given the devastating effect unabated climate change will have on biodiversity, but also the increased strategic importance of protected areas in a warming world. many of the presenters who addressed the issue of climate change spoke about the role protected areas will play in adaptation through increasing resilience to environmental extremes. for example, representatives of the iucn from the pacific islands explained the potential for ecosystem-based adaptation to sea level rise by planting mangroves to buffer coastlines against king tides. the immense opportunities for integrating biodiversity outcomes with greenhouse gas emission mitigation activities were also discussed, especially in relation to the united nations reducing emissions from deforestation and forest degradation projects. however, presenters from participating projects expressed frustration at the lack of demand for carbon credits globally. it is hoped that the shift to a carbon-constrained world will increase demand in carbon markets, with co-benefits for biodiversity conservation. conclusion top ↑ the iucn had a difficult task in organising the 2014 world parks congress. it had to be mainstream and inclusive, creating an open forum where politicians and business leaders were included in conversations with dedicated conservationists. it had to confront, in an open and honest manner, the many challenges and the scale of action required. the congress also needed to celebrate significant achievements and progress whilst acknowledging the current biodiversity crisis and our uncertain future. these different tensions were reflected in the formal conclusion of the congress, ‘the promise of sydney’ (see http://worldparkscongress.org/about/promise_of_sydney.html), which was collaboratively written by congress participants. this representation of the congress outcomes included a vision for the future, a set of innovative approaches and solutions to solve challenges facing protected areas, and commitments made at the congress by countries and organisations. cynics might say that these outcomes amount to hand waving and rhetoric. we all know that talk is cheap, but human interaction and the sharing of ideas with other individuals is motivating and helps create the global networks required to address global problems. acknowledgements top ↑ competing interests the author declares that she has no financial or personal relationships which may have inappropriately influenced her in writing this article. issn 0075-6458 103 koedoe 49/2 (2006)issn 0075-6458 103 koedoe 49/2 (2006) the quality and effectiveness of strategic environmental assessment (sea) as a decision-aiding tool for national park expansion—the greater addo elephant national park case study f. retief retief, f. 2006. the quality and effectiveness of strategic environmental assessment (sea) as a decision-aiding tool for national park expansion—the greater addo elephant national park case study. koedoe 49(2): 103–122. pretoria. issn 0075-6458. internationally south africa is regarded as a leading developing country in the application of sea. research also indicates that the conservation sector within south africa has been particularly prominent in exploring its application. however, very little empirical research has been conducted to learn from practice and to establish the added value of sea to strategic decision making. to address this gap in knowledge the paper explores the input quality and output effectiveness of the greater addo elephant national park sea. the results show that the sea achieved average to good input quality and output effectiveness performance, respectively. a number of lessons were learned for the application of sea to future park expansion initiatives. these relate to the timing of the sea, addressing social impacts, setting of boundaries as well as dealing with scale. the research results concluded that sea made valuable contributions to decision making and could be considered an important decision-aiding tool for the conservation sector. to take the debate forward and to benchmark best practice it is proposed that further comparative performance evaluation research of multiple sea case studies within the conservation sector be conducted. key words: strategic environmental assessment, quality and effectiveness, performance evaluation, park expansion, conservation sector f. retief, senior lecturer, school of environmental sciences and development, north west university (potchefstroom campus), private bag x6001, potchefstroom, 2520, south africa, (ggffpr@puk.ac.za). introduction south africa, with its exceptional natural beauty and cultural diversity is sometimes fittingly described as a world in one country and the rainbow nation. although the country is world renowned for its achievements in the field of conservation and biodiversity management, it is also known for its turbulent political past marred by inequality and social injustice. it is a context where sustainability cannot be considered lightly because it deals with pressing short term survival issues as well as longer term concerns with quality of life. strategic environmental assessment (sea) was introduced as a tool that could assist with integrating the concept of sustainability into strategic level decision making (csir 1996; deat 2000; rossouw et al. 2000; deat 2004b). the introduction of sea has been on the back of extensive international refinement and wide adoption within developed as well as developing country contexts (dalal-clayton & sadler 2005; jones et al. 2005; schmidt et al. 2005). in terms of developing countries, south africa has received international acclaim for its homegrown approach to sea (therivel & partidario 2000; dalal-clayton & sadler 2005). the extent of sea practice within the country is also considered exceptional, with 50 seas conducted across nine different sectors between 1996 and 2003 (retief 2005). the three best-represented sectors were planning, conservation and water management. however, very little research has been conducted retief.indd 103 2006/10/15 10:57:39 pm koedoe 49/2 (2006) 104 issn 0075-6458 on the quality and effectiveness of sea within these different sectors. from a conservation perspective, south africa is classified by the world conservation monitoring centre as the third most biologically diverse country in the world (world conservation monitoring centre 1992, 2002). however, this status also inevitably complicates the trade-offs that need to be made between the country’s pressing development needs and its wealth in biodiversity. wynberg (2002) tracked the progress in biodiversity conservation in south africa from the rio earth summit in 1992 to the johannesburg world summit on sustainable development (wssd) in 2002 and suggests that a paradigm shift has taken place where biodiversity shifted from the realms of pure conservation synonymous with ‘saving the rhino’, to encompassing politics, culture and economy. this shift has come about through the democratisation process in south africa, but also changes in international thinking on resource management. subsequently, sustainable development is understood as a development approach rather than a biodiversity conservation strategy. since the wssd in 2002, south africa, in an effort to address the lack of a national strategic framework has promulgated the national environmental management biodiversity act (nemba) 10 of 2004, published a national biodiversity strategy and action plan as well as a national spatial biodiversity assessment (deat 2004a). moreover, in line with international thinking (kolhoff & slootweg 2005; treweek et al. 2005), specific biodiversity-focused approaches to planning, such as the bioregional approach, have emerged more strongly (nbi 2002). an example is the cape action plan for the environment (cape) that explicitly incorporated sea elements (lochner et al. 2003). however, due to continued institutional and legal fragmentation, as well as until recently the lack of a national sustainability strategy, little progress has been made to incorporate biodiversity issues into mainstream decision making (rossouw & wiseman 2004; glazewski 2005). as wynberg (2002) states: “…, biodiversity considerations at the strategic planning level remain marginalised and narrow, and sorely neglected within existing plans. … strategic environmental assessment allows issues of biodiversity to be more strongly integrated into the formulation of policies, plans and programmes.” thus, although the exact role and potential added value of sea to conservation and biodiversity decision making is still unclear, and has not entered formal debate, sea already seems to have started to make important contributions (brownlie et al. 2005). this paper explores the quality and effectiveness of sea as a tool to inform strategic decision making related to conservation. the aim is to highlight the added-value of sea but also to gain experience in tailoring sea as a tool to assist initiatives involving the establishment of new and extension of existing conservation areas within south africa. the complexities associated with sea performance evaluation required a case study research approach. for this reason, a specific high-profile sea case study was identified, namely the greater addo elephant national park (gaenp) sea. case study—greater addo elephant national park the addo elephant national park (aenp) had its humble beginnings in 1931 when it was proclaimed to protect the last dwindling population of eastern cape elephants. since then, the park has been expanded from its original 2 200 ha to over 120 000 ha in 2004, and potentially over 400 000 ha in future (fig. 1). but the aenp was not the only conservation area established in the region. several other provincial reserves and private reserves were established, as well as three conservancies and two heritage sites. the integration of these different conservation areas was desirable, since from a biodiversity management perspective, ‘single large’ is better than ‘several small’. in 1997, kerley & boshoff (1997) completed an award winning proposal that set out the framework retief.indd 104 2006/10/15 10:57:39 pm issn 0075-6458 105 koedoe 49/2 (2006) for a greater aenp that would integrate and extend these existing conservation areas. the challenge for the park was accepted by south african national parks (sanparks) and set the scene for one of the most ambitious, and potentially significant, contributions to biodiversity conservation in africa. the new enlarged park would be the third largest national park in south africa and cover six of the seven south african biomes. the implementation of such an ambitious endeavour would, however, require extensive funding, which could not be accessed fully nationally. due to the potential international contribution the project could make to conservation it was feasible to approach international funding agencies (the overall gaenp expansion initiative was referred to as a ‘project’ which might cause some confusion since sea normally relates to policy, plan and programme (ppp) levels of decision fig. 1. extent of the gaenp planning domain (ces 2002) making). ultimately the global environment facility (gef) provided a grant, administered through the world bank (wb), to conduct a series of specialist studies to form the basis for a full project proposal for further funding. the wb has a number of safeguard policies, which had to be adhered to if further funding was to be secured. in 2002, a sea was commissioned as a requirement in accordance with wb safeguard policy requirements and aimed to consolidate all the specialist information in a manner that would facilitate sound strategic planning and decision making. the lead agent of the sea was sanparks who appointed coastal and environmental services (ces) as the consultancy to carry out the sea in line with the 2000 south african sea guidelines (deat 2000; ces 2002). the entire area specifically considered as part of the sea for the gaenp is referred to as the ‘planning domain’. it includes a retief.indd 105 2006/10/15 10:57:39 pm koedoe 49/2 (2006) 106 issn 0075-6458 terrestrial area of approximately 340 000 ha (around which a 5 km buffer is included) and a marine area of approximately 100 000 ha inclusive of the st croix and bird island groups (fig. 1). the criteria used to determine the boundaries of the planning domain related to the potential of the park to achieve certain conservation and development goals in line with south africa’s development policies and strategies on the environment (kerley & boshoff 1997; ces 2002). these goals relate to the significance of the biodiversity assets as well as the need to facilitate compatible development, such as tourism to address social and economic needs typical of the underdeveloped rural areas of the eastern cape, south africa’s poorest province. examples of the biophysical assets for the area covered by the park include: terrestrial environment (csir 2002)— the terrestrial environment covers six biomes of which the thicket biome is best represented, covering 69 % of the planning domain. the others are forest (10 %), nama karoo (7 %), fynbos (5 %), grassland (5 %) and savanna (4 %). the alexandria forest is unique and endemic to the eastern cape while fynbos is renowned for its diversity of floral species. the most conspicuous faunal species are the addo elephant (loxondonta africana) and the black rhinoceros (diceros bicornis bicornis and d. b. michaeli). other less visible but extremely significant species include the endemic dune grasshopper (arctylos hirsutus) and hairy-footed gerbil (gerbillurus paeba exilis). freshwater environment (csir 2001b)— the planning domain includes nine different types of water bodies having a complement of biotopes, namely: perennial and seasonal rivers; episodic rivers; ephemeral rivers; permanent and semipermanent vleis; seasonal vleis; springs and seeps; episodic endorheic pans; forest swamps and marches. a number of threatened faunal species rely on these bodies for survival such as the redfin minnow (barbus afer). marine environment (csir 2001a)— the proposed marine protected area (mpa) is of national and international significance. due to its location, the fish fauna comprises species found on both the east and west coasts of south africa, many of which are endemic. it is also known for its population of great white shark (carcharadon carcharias) and 9 species of whale, dolphin and seal are relatively common. the two island groups accommodate a number of bird species of conservation significance such as the african penguin (spheniscus demersus) and roseate tern (sterna dougallii), south africa’s most threatened marine bird. finally the alexandria coastal dune field is regarded as the largest and least degraded coastal dune field in south africa. in light of these bio-physical features the area also presents significant social and development challenges, such as: land ownership and acquisition (connor 2002; wren & msutu 2002)—sensitivity surrounding the issue of land ownership presented a particular challenge for the expansion of the park. the majority of farm owners and farm workers have a long-term occupancy in the region, which made the land ownership issue even more contentious. the process used for the acquisition of land was criticised and the possibility of expropriation created negative perceptions. the management scenarios for the park had to consider innovative solutions to accommodate conservation on private land such as contractual agreement initiatives. it also led to the formulation of a land acquisition policy. employment (antrobus 2002; connor 2002)—the gaenp project had to show that it would contribute to economic development and employment within the planning domain. analysis of existing employment patterns related to the different land uses was required to make such estimates. it concluded that conservation combined with tourism should be able to create more employment than existing agricultural land uses, except for dairy farming. also, ways retief.indd 106 2006/10/15 10:57:39 pm issn 0075-6458 107 koedoe 49/2 (2006) of retraining and redeployment of existing labourers had to be achieved within the requirements of the wb safeguard policies prohibiting any form of involuntary resettlement. methods due to the substantial difficulties experienced with sea performance evaluation (sadler 2004; partidario & arts 2005), and specifically in dealing with effectiveness (therivel & minas 2002; cashmore et al. 2004; owens et al. 2004) no international methodological blue print exists. the performance evaluation methodology for the research was based on the quality and effectiveness sea review protocol designed by retief (2005). it was based on national as well as international sea literature, which provided the basis for the development of a conceptual framework and the identification of existing performance criteria for sea quality and effectiveness. in line with international literature, the protocol distinguishes between input quality at the level of application and output effectiveness at the level of implementation (sadler 1996; lawrence 1997; thissen 2000a). furthermore, the protocol distinguishes between three sea input quality components, namely: process, methodology and documentation. in terms of output effectiveness, two components were identified namely direct and indirect outputs. direct outputs are understood to relate to the objectives of sea, which include aspects such as changes to decisions, improvement in the environmental quality and changes to the contents of plans or programmes. indirect outputs are more difficult to measure and include outputs such as changes in attitudes towards the environment, improved awareness, changes in institutional arrangements and departmental traditions. to provide some form of measurement, indicators were developed. the use of the term indicators was preferred to criteria because indicators suggest that they are indicative, whereas criteria imply precision not always achievable due to the complex nature of sea (todd 2001; retief 2005). the methodology for the design of key performance areas (kpas) and key performance indicators (kpis) was based on the agreed notion that overarching context specific sea principles and objectives should form the basis for the application of sea (marsden 1998; rossouw et al. 2000; thissen 2000b; verheem & tonk 2000). examples of such sea principles have been designed for sea internationally (iaia 2002), and also for south africa specifically (deat 2000). ultimately, 14 kpas and 48 kpis were identified for quality review of sea. it was based on the understanding that sea involves a context-specific, sustainabilityled, participative, pro-active and efficient process. this requires different sea methodologies during screening, situation analysis, scoping, environmental assessment and monitoring, and review phases. the results of these need to be documented and communicated to decision makers and interested and affected parties (iaps). in terms of effectiveness, four kpas and nine kpis were developed. table 1 provides a summary of the kpas and kpis for quality and effectiveness and their relation to sea principles and objectives. to deal with the qualitative and subjective nature of the conformance measurement only three scales were used namely, conformance, partial conformance and non-conformance. the level of conformance was determined through in-depth analysis of the sea documentation as well as interviews with selected key role players. the basic assumption is that conformance to more indicators implies better quality and/or effectiveness performance (for a more detailed description of the protocol see retief (2005)). however, the literature warns against just adding up different variable scores as with a quantitative survey approach (miles & huberman 1994; yin 2003). this would destroy the local web of causality and result only in a smoothed down set of generalisations. as reflected in the following sections the research ultimately aimed to tell a story rather than produce a quantitative performance score. results and discussion table 2 provides a summary of the overall review results. the input quality results show that the sea failed to conform to 13 indicators, partly conformed to 20 and conformed to 15 of the 48 indicators. the larger number of partly conformed and conformed indicators suggests an average to good quality performance. on the other hand, the output effectiveness performance shows conformance or partial conformance to all except one of the indicators (the status of two of the indicators could not be established). this also suggests an average to good effectiveness performance. although it is not possible at this stage to identify specific linkages between input quality and output effectiveness it can be concluded that the input quality does seem to be reflected in the output effectiveness with what could be considered average to good performance on both sides. retief.indd 107 2006/10/15 10:57:39 pm koedoe 49/2 (2006) 108 issn 0075-6458 sea principles “basic building blocks for the context specific perspectives on sea in south africa” (adapted from deat 2000) sea objectives “indication of what needs to be achieved” (adapted from deat 2000) kpas “theme related to principles” kpis “questions that provide an indication if sea objectives were achieved” key process principles key process objectives process kpas process kpis there is not one sea process to be used in all contexts. this requires a sea process to be flexible and adaptable, in order to integrate with the decision making context. • to integrate the sea with the decision making context. • to avoid the duplication of processes. context specific kpi 1.1: was the sea fully integrated with the plan or programme formulation process, from conceptualisation to implementation? kpi 1.2: did the sea make provision for tiering with project eia? kpi 1.3: did the sea formulate actor and process configurations? development must be socially, environmentally and economically sustainable. sea provides a practical means of integrating the concept of sustainability into plan and programme formulation. • to integrate the concept of sustainability into plan and programme level decision making. • to facilitate the development of local definitions and understandings of sustainability. sustainabilityled kpi 2.1: did the sea documentation provide a definition for sustainability, which is consistent with the way sustainability is understood in the local context? kpi 2.2: was sustainability included as a specific objective of the sea? kpi 2.3: was an attempt made, as part of the sea, to measure sustainability by means of parameters, objectives, criteria or indicators? kpi 2.4: did the sea give equal consideration to the biophysical, social and economic aspects? public participation forms an integral part of sea because sea puts people and their needs at the forefront of its concern. • to inform and involve iaps throughout the sea process. • to incorporate public inputs and concerns into decision making processes. • to facilitate information sharing. participative kpi 3.1: was a formal public participation process followed, which informed and involved the iaps throughout the sea process? kpi 3.2: were the iaps satisfied with the public participation process? kpi 3.3: did the sea explicitly address public inputs and concerns? kpi 3.4: were all key state departments and other governing bodies consulted during the sea? table 1 summary of linkages between sea principles, objectives, kpas and kpis developed for south africa (retief 2005) retief.indd 108 2006/10/15 10:57:39 pm issn 0075-6458 109 koedoe 49/2 (2006) sea provides a means of influencing decision making throughout its life cycle, from conceptualisation to implementation in an incremental and iterative way while facilitating the concepts of pre-caution and continuous improvement. • to ensure that the sea is implemented early enough to influence decision making. • to facilitate continual improvement. proactive kpi 4.1: did the sea ensure availability of the assessment results early enough to influence the decision making process? kpi 4.2: was commitment confirmed to ensure that the results of the sea be considered in future decisionmaking? the benefits of implementing sea exceed the costs. sea adds value to existing decision making by focusing on key strategic environmental issues. • to provide sufficient, reliable and usable information. • to optimise the use of time and resources. • to focus decision making on the key environmental issues. efficient kpi 5.1: did the sea provide sufficient information for decision-making according to the relevant decisionmakers? kpi 5.2: were sufficient resources and time allocated to conduct the sea according to the relevant sea role players? kpi 5.3: did the sea focus on key significant strategic issues? key methodology principles key methodology objectives method ology kpas methodology kpis sea has to justify why it needs to be applied and what it aims to achieve. • to justify the need for the sea. • to clearly define project objectives of the sea. screening kpi 6.1: was a formal screening method or criteria applied? kpi 6.2: was the need for the sea clearly defined? kpi 6.3: was the purpose and/or objectives of the sea clearly defined that could serve as reference for effectiveness review? sea determines the opportunities and constraints that the environment places on development. • to provide sufficient information on environmental attributes to identify opportunities and constraints. situation analysis kpi 7.1: was a resource inventory prepared which describes the social, economic and biophysical aspects in the area at the appropriate scale and level of detail? kpi 7.2: was the state of the environment (including economic, social and bio-physical) determined against set objectives, criteria or indicators? kpi 7.3: were environmental opportunities and constraints identified by means of a justified methodology? table 1 (continued) retief.indd 109 2006/10/15 10:57:39 pm koedoe 49/2 (2006) 110 issn 0075-6458 sea identifies the most significant key strategic environmental issues. • to ensure that key strategic environmental issues are identified. scoping kpi 8.1: was a formal scoping method applied? kpi 8.2: did scoping assist in defining the scope and extent of the sea? kpi 8.3: did the scoping method(s) focus the sea on key significant strategic issues? kpi 8.4: were public inputs considered during scoping? sea determines the implications of strategic decisions on the environment. • to ensure that environmental implications of strategic decisions are considered. environmental assessment kpi 9.1: was an assessment conducted against a sustainability framework (it may include sustainability parameters / objectives / criteria and indicators)? kpi 9.2: were different scenarios and/or alternatives considered to identify the best option? kpi 9.3: were the assessment techniques appropriate in terms of the context, available resources as well as data quality and availability? kpi 9.4: were cumulative effects considered? sea aims for continuous improvement, which relies on monitoring and review mechanisms. • to ensure that the sea is reviewed and the implementation of proposals monitored. monitoring and review kpi 10.1: did the sea propose a plan for monitoring? kpi 10.2: has the sea been independently reviewed? kpi 10.3: has environmental monitoring been conducted? key documentation principles key documentation objectives documentation kpas documentation kpis information should be documented in a sound and justifiable manner. • to provide sound and justifiable information, which allows for verification of results. • to contribute to existing environmental data and information. description of context kpi 11.1: were the purpose and objectives of the sea described in the documentation? kpi 11.2: was the decision making contexts and linkages with other decision making processes described? kpi 11.3: was a description provided of the sea process followed? kpi 11.4: were those involved in consultation and participation indicated? table 1 (continued) retief.indd 110 2006/10/15 10:57:40 pm issn 0075-6458 111 koedoe 49/2 (2006) description of the state of the environment kpi 12.1: was a description provided of the current state of the environment (either as a separate volume or integrated with the description of the baseline environment)? kpi 12.2: was the state of the environment described against clear thresholds and/or limits of acceptable change in a way that highlights relative significance? description of assessment methodology and results kpi 13.1: were the different methods applied in the sea described (relating to for instance screening, scoping and environmental assessment)? kpi 13.2: was a description of key significant strategic environmental issues given? kpi 13.3: were different scenarios and/or alternatives described? kpi 13.4: were the recommendations and/or terms of approval described? kpi 13.5: was a summary provided of difficulties encountered and subsequent uncertainties in results? sea should be documented in a manner that ensures effective communication of results in order to optimise the possibility of it influencing decision making. • to communicate the results of the sea to decision makers. • to communicate the results of the sea to iaps. communication of results kpi 14.1: were the contents clearly explained, justified and logically arranged in sections or chapters? kpi 14.2: were the specialist reports well referenced and integrated in a way that promotes a self-contained document? kpi 14.3: was a non-technical summary provided of the main results and conclusions? kpi 14.4: were the inputs received from iaps incorporated in the report? key direct outputs principles key direct outputs objectives direct outputs kpas direct outputs kpis sea influences the contents of plans and programmes. • to influence the contents of plans and programmes. policies, plans and programmes kpi 15.1: were any plans or programmes amended based on the proposals of the sea? kpi 15.2: did the sea facilitate the incorporation of sustainability objectives into relevant plans or programmes? table 1 (continued) retief.indd 111 2006/10/15 10:57:40 pm koedoe 49/2 (2006) 112 issn 0075-6458 sea facilitates the achievement of sustainability objectives • to achieve the sea project objectives. • to achieve the sea sustainability / environmental objectives. sea objectives kpi 16.1: were the sea project objectives achieved (as described in the tor)? kpi 16.2: were the sustainability / environmental objectives achieved (as might be described in relation to the vision)? sea influences decision making • to influence decision making. decision making kpi 17.1: were decisions changed or amended based on the outcomes and proposals of the sea? kpi 17.2: was the sea implemented as a decision-support guideline for future development proposals? kpi 17.3: did the sea inform/guide subsequent project level decision making (such as eia or water licensing)? sea improves environmental quality • to improve environmental quality environmental quality / sustainability kpi 18.1: were changes to the environment observed since the completion of the sea process, which could be attributed to the influence of the sea? kpi 18.2: did the sea accurately identify the key significant strategic environmental issues? interpretation of overall results: poor: failure to conform to large majority of indicators. average to poor: failure to conform, and/or partial conformance, to the majority of indicators. average: partial conformance to the majority of indicators, or an even spread in performance. average to good: partial, and/or conformance, to the majority of indicators. good: conformance to large majority of indicators. table 1 (continued) input quality components sea process component the review results show that the sea failed to conform to four, partly conformed to seven, and conformed to five of the 16 indicators. this section describes the sea process quality with specific reference to kpis 1.1, 2.4, 3.1, 4.1, 4.2 and 5.2 (table 2). tracing the origins of the sea through documentation and interviews revealed that considerable work on the conceptualisation of the project had already been completed by the time the sea was initiated (kerley & boshoff 1997) (kpi 4.1). importantly, this included a public participation process where, during a public workshop on 23 february 1999, consensus was reached and a retief.indd 112 2006/10/15 10:57:40 pm issn 0075-6458 113 koedoe 49/2 (2006) table 2 overall performance evaluation results: sea for the gaenp kpas kpi results in pu t q ua lit y pr oc es s context specific 1 1.1 1.2 1.3 sustainability led 2 2.1 2.2 2.3 2.4 participative 3 3.1 3.2 3.3 3.4 pro-active 4 41 4.2 efficient 5 5.1 5.2 5.3 m et ho do lo gy screening 6 6.1 6.2 6.3 situation analysis 7 7.1 7.2 7.3 scoping 8 8.1 8.2 8.3 8.4 environmental assessment 9 9.1 9.2 9.3 9.4 monitoring and review 10 10.1 10.2 10.3 d oc um en ta tio n description of context 11 11.1 11.2 11.3 11.4 description of state of the environment 12 12.1 12.2 description of assessment methodology and results 13 13.1 13.2 13.3 13.4 13.5 communication of results 14 14.1 14.2 14.3 14.4 o ut pu t e ff ec tiv en es s d ir ec t o ut pu ts plans and programmes 15 15.1 15.2 sea objectives 16 16.1 16.2 decision making 17 17.1 17.2 17.3 environmental quality 18 18.1 18.2 non-conformance partial conformance conformance status could not be established indicators with a particular relevance to this case study retief.indd 113 2006/10/15 10:57:40 pm koedoe 49/2 (2006) 114 issn 0075-6458 unanimous declaration passed in support of the gaenp initiative (teru 1999). thereby a real sense of optimism and passion was generated amongst sanparks to realise the vision and objectives formulated for the gaenp initiative. based on this initial project conceptualisation phase, sanparks realised that the project would need to comply with eia legislation because of the change in land use from agriculture to conservation and also expansion of existing services infrastructure such as roads. but, due to the geographic scale and extent of the project, deat considered eia to be unsuitable and advised sanparks that a sea be conducted instead (knight 2004 pers. comm.). the consultant remarked that at that stage “sea was seen as the panacea for all environmental problems” and there did not seem to be clarity on sea, “anything not called eia was considered sea” (avis 2004 pers. comm.). outcomes of the interviews suggest that the sea did not include a specific screening phase and was initiated largely based on the limitations of eia to deal with the particular scale rather than a clear understanding of sea and what it could deliver. in view of the timing of the sea, it is not surprising that one of the major shortcomings was its lack of integration with the conceptualisation phase (kpi 1.1), which resulted in the sea being unable to influence the original gaenp vision and primary objectives. the terms of reference (tor), prepared mainly by sanparks, also did not task the sea to revisit the vision and primary objectives (ces 2002), which the consultant highlighted as a key weakness of the sea (avis 2004 pers. comm.). additionally the public did not understand what the purpose of the sea would be if it did not have the mandate to question or influence the primary project objectives (wren & msutu 2002). this jeopardised the sea and risked that it “… be reduced to a validation exercise for a pre-supposed vision and objectives” (wren 2004 pers. comm.) from a sustainability perspective one got the impression that there were so-called green and brown lobbies (kpi 2.4). the green lobby seemed to be represented by the proponent and those responsible for conservation planning and the brown lobby by those responsible for the social and economic specialist studies. it was unanimously expressed by those involved in the sea (including sanparks) that during the early stages of the sea “the conservation planners did not fully appreciate the social aspects and their constraints to the expansion of the park” (knight 2004 pers. comm.) the irony is that, due to its very nature, the gaenp initiative (which involves the expansion of a conservation area and rehabilitation of farm land) would not present any negative biophysical impacts, and the main contribution of the sea turned out to be towards the social aspects. the perception from iaps was that sanparks did not anticipate, nor were they sensitive to, the social implications of the park expansion. the biggest social issue was the re-employment of retrenched farm workers (connor 2002; wren & msutu 2002). the title deeds of many farms stipulated a vacancy clause, which requires the farm to be vacant on transfer from the farmer to sanparks. this implied eviction of the workforce. in light of the long-term residency (spanning generations) many farm workers would not adapt easily to any other way of life. low levels of education also limit their future employment prospects. it was argued by land owners that sanparks should at least be partly responsible for the livelihoods of these rural communities, especially where expropriation was involved (connor 2002). to underline the negative perceptions comments were made by some iaps that “coega [controversial industrial development in the area] would never have gotten away with the same approach to social issues that the addo project did” and that, “if it were not for the world bank safeguard policies on resettlement the social issues would never have been addressed” (anonymous pers. comm.) retief.indd 114 2006/10/15 10:57:40 pm issn 0075-6458 115 koedoe 49/2 (2006) due to the (largely unanticipated) significant social issues related to the expansion of the park, the public participation process turned out to be a vitally important contribution to the sea process (kpi 3.1). probably the biggest problem for the legitimacy and credibility of the participation process was the continuation of land purchase by sanparks while the sea process was being conducted (wren 2004 pers. comm.). the consultants objected to this (avis 2004 pers. comm.). it was only reasonable and understandable that certain sectors of the public interpreted this as confirmation that the sea was only a window dressing exercise and that the project was going ahead despite their inputs. in certain areas people withdrew and distanced themselves totally from the process because of this (wren & msutu 2002). however, sanparks argued that they have been purchasing land on a continuous basis since the establishment of the park in the 1930s. also, the rural areas presented a particular challenge for the participation process where the high profile of the gaenp initiative generated various rumours among the farming community that caused uncertainty and distrust, which included amongst others, expropriation threats. many farmers have been farming the land for generations and were unwilling to become involved. in this regard farm workers proved even more difficult to liase with since they were isolated on farms and were not organised into representative bodies such as trade unions (wren & msutu 2002). sanparks did employ an effective and well managed project management regime and was very specific in terms of the products they wanted (knight 2004 pers. comm.). they were clearly passionate about the establishment of the park, which probably manifested in their approach to their project management function as a contribution towards realising the gaenp vision. it also ensured efficiency in terms of time and resources (kpi 5.2). however, the lead sea consultant (who was also responsible for the social specialist studies) indicated that communication with the conservation planners was inadequate and left some room for improvement (griffith 2004 pers. comm.). although the sea did make provision for a review period of 28 days for iaps, environmental authorities (national and provincial) had not commented on the sea by the time of the review in july 2004 (avis 2004 pers. comm.). the consultant specifically expressed the need to tidy up the review process (avis 2004 pers. comm.). moreover, sanparks never formally adopted the sea and confirmed that it only serves as an informal information document (knight 2004 pers. comm.) (kpi 4.2). also, the sea was never given closure through formal commitment from sanparks to implement the proposals. two other factors worried the consultants, namely the fact that the sea was conducted outside a formal regulatory framework as well as the status of sanparks as a statutory body of the national department of environmental affairs and tourism (deat), effectively positioning them as referee and player. the latter prompted the consultant to write a formal letter on 1 august 2002 to deat asking “… could you please explain how the decision making process would work. in this instance national deat is both referee and player, and iaps have been concerned that all decisions would fall in favour of sanparks” and “could you therefore map out for us the regulatory framework within which decisions regarding the establishment of the gaenp will be made.” no formal reply was received (avis 2004 pers. comm.). not surprising, some key role players expressed their doubts on whether sanparks would actually implement the proposals of the sea. however, from the following sections it is clear that sanparks, in the end, did indeed use and implement some of the sea proposals. sea methodology component the methodology review results show that the sea failed to conform to five indicators, partly conformed to eight, and conformed to four of the 17 indicators. this section presents the quality performance of the methodretief.indd 115 2006/10/15 10:57:40 pm koedoe 49/2 (2006) 116 issn 0075-6458 ology with specific reference to kpis 6.1, 6.2, 6.3, 7.1, 7.2, 7.3, 8.1, 8.3, 8.4, 9.1 and 10.1. the methodology of the sea was primarily designed to deliver the terms of reference (tor). a formal screening method was not applied (kpi 6.1) and sanparks, in consultation with deat, drafted the tor. the first and most significant aspect of the tor was that it did not include a requirement to specifically assess anything (kpi 6.3). rather it specified that the sea should essentially integrate the specialist information in order to identify opportunities and constraints and describe the socio-economic and biophysical implications of establishing the gaenp, and finally, make proposals for the way forward (ces 2002). this was exactly what the sea did. so it could almost be considered more of a feasibility study on the desirability of the park expansion than an assessment. as part of the situational analysis, a total of 17 specialist studies were conducted, covering biophysical, social, economic and institutional aspects (kpi 7.2). it was explained that the sea followed a nested approach where the biophysical and social assessments focussed on the planning domain and the legal, economic and institutional assessments focussed more broadly, looking at interactions within the region. each specialist study had its own tor and applied its own methodology, which ranged from detailed gis analysis (for the conservation planning) to simple desktop studies and descriptions (for the economic and institutional studies). the budgets for each study also varied greatly, with the majority of funds allocated towards conservation planning. all this resulted in different levels of detail and formats, which made integration of specialist studies exceedingly difficult (kpi 7.1). for example, the conservation plan (c-plan) classified detailed land parcels according to irreplaceability from where priority expansion areas were identified (csir 2001b). the social data relied on a representative sample, consisting of 16 farms spread throughout the planning domain (connor 2002). sanparks confirmed that the aim to link the social and biophysical data at the same geographic scale and level of detail, by means of gis, was never achieved (gordon 2004 pers. comm.; knight 2004). this illustrates that the broadbrush social analysis could not purposefully influence the specific land purchases envisaged by the c-plan. the boundary of the study area was also a key aspect influencing methodology. the boundaries of the gaenp initiative were determined based on certain conservation and development goals even before the sea was initiated. the sea study area was subsequently referred to as a planning domain wherein, crucially, the boundaries were not fixed, primarily because sanparks argued that the future extent of the park would be continuously influenced by new information and the development of relationships with private landowners (ces 2002). the flexible nature of the boundaries caused various problems for data gathering and public participation respectively (kpi 7.1). this was because the planning domain was largely determined based on biodiversity significance and subsequently did not correspond to administrative boundaries. since most existing economic and social data were linked to municipal or district boundaries it was at times difficult to match data with the boundaries of the planning domain (geach 2002; timmermans & sisitka 2002). the public participation process suffered because iaps could not gauge the influence of the exact extent of the park on them. it also created feelings of mistrust and uncertainty (wren & msutu 2002). it is understandable that for private landowners it was imperative for the park to indicate exact boundaries so that they could assess the potential impact on their livelihoods. as part of the situation analysis, a key deliverable of the sea was the identification of opportunities and constraints for the gaenp proposal (kpi 7.3). afterwards sanparks described the opportunities and constraints approach as amorphous and not very rigorous (knight 2004 pers. comm.). on the other hand the consultant considered it a valuable and beneficial approach but pointed out one specific limitation. what was an opportunity retief.indd 116 2006/10/15 10:57:40 pm issn 0075-6458 117 koedoe 49/2 (2006) for one alternative (such as conservation) was a constraint for the other (such as agriculture) and vice versa, which implied that the interpretation differed depending on the proponent (avis 2004 pers. comm.). for instance, the lower agricultural potential of the zuurberg area provides an opportunity for conservation and a threat to agriculture. similarly, the high agricultural potential of the alexandria area presents an opportunity for agriculture and a threat to conservation. the latter was even more significant in light of the fact that only one land-use alternative, namely conservation, was considered and other alternatives such as improving the status of agriculture were not. the consultant highlighted the failure to pursue any alternatives (for the area) other than conservation, as a significant limitation of the sea (ces 2002). however, it was argued by sanparks that other alternatives were not forthcoming due to the fact that they were not considered viable and that conservation was the only alternative worth pursuing (ces 2002; world bank 2002). there was also no proponent for other alternatives or funds to investigate their viability. although only one overriding land-use alternative was put forward, various development scenarios (in terms of tourism products and game farming) as well as management arrangements (in terms of contractual parks) were considered as part of the specialist studies (davies 2002; timmermans & sisitka 2002). these were referred to as incremental alternatives (ces 2002). although no specific scoping phase could be distinguished, issues were scoped based on public participation and specialist studies (ces 2002; wren & msutu 2002) (kpi 8.1). however, these issues were not prioritised or weighted in terms of relevant significance (kpi 8.3). since the sea ended up with a relatively small number of key issues, prioritisation was probably not that critical, although it would have been especially helpful in the implementation of the sea proposals. however, the consultant indicated that an attempt was made to prioritise through a scoring exercise during a specialist workshop but that it was abandoned due to time constraints (griffith 2004 pers. comm.). the contribution that public participation made to the identification of key issues for the area can hardly be overstated (kpi 8.4). it is significant to note that the majority of strategic issues raised through the specialist studies also seemed to have been captured during the participation process, which might be an indication that public participation could serve as a valuable substitute where specialist inputs are not available or affordable. the fundamental methodological limitation of the sea, which negated the possibility of actually conducting an assessment, was that it failed to set objectives, targets and indicators (which could have formed part of a sustainability framework) and nobody seemed to know how to address this limitation (kpi 9.1). this was especially true for the social aspects, but less so for the biophysical because conservation targets were set for the terrestrial environment (csir 2002). however, since overall targets and indicators were not set it was difficult to determine the sustainability and viability of, for instance, the conservation targets against social targets. the lack of targets and indicators also meant that no monitoring arrangements were put in place (kpi 10.1). so, sanparks highlighted that there was a definite need for an implementation phase to bridge the gap between planning and implementation (gordon 2004 pers. comm.). for instance, there was no direct interaction between the lead sea consultant and the park management, which suggests that the integration of the sea with actual park management was limited and that the new integrated environmental management system (iems) for the park was designed separately from the sea. sea documentation component the sea documentation conformed to six indicators, partly conformed to five, and did not conform to four of the 15 indicators. this section presents the performance of the sea with specific reference to kpis 11.2, 11.3, 12.2, 13.1, 13.3, 14.1, 14.3 and 14.4. retief.indd 117 2006/10/15 10:57:40 pm koedoe 49/2 (2006) 118 issn 0075-6458 the sea documentation consisted of 17 specialist study reports, the results of which were integrated into a single sea report (ces 2002), that described opportunities and constraints and made proposals for the way forward. an executive summary was also included (kpi 14.3). not surprising, the areas of partial and non-conformance could be related back to limitations also reflected in the process and methodology components. for instance, the sea process did include a formal participation process, which was well recorded and thus those involved in consultation and participation were clearly described and inputs received from the public were incorporated in the report (kpi 4.4). also the sea did not consider any other fundamental land-use alternative than conservation, thus no alternatives were described in the documentation (kpi 13.3). the review results show that the context of the sea was not well described, apart from those involved with consultation and participation (kpi 11.2). the reader could not identify linkages between the sea and other decision making processes. in addition, more importantly, the actual sea process was not described (in relation to the gaenp development process) (kpi 11.3). the lack of objectives, targets and indicators and clear methodologies for screening, scoping and assessment resulted in the lack of descriptions of the state of the environment (kpi 12.2) and assessment methodologies (kpi 13.1). however, the sea did communicate its results well. the documents were logically arranged, well referenced and the inputs received from the public were clearly incorporated (kpi 14.1). this might also be one of the advantages of having a formal public participation process which ensures that information is presented in a user-friendly format. the availability of the sea documentation on the internet was also a definite advantage of the sea. it can be concluded that the documentation can only provide what the sea process and methodology were designed to deliver. output effectiveness components the review results suggest an average to good effectiveness performance and show that the sea did deliver on the tor and partly influenced policy and decision making. the following sections discuss the direct and indirect outputs of the sea. direct outputs component the sea recommendations highlighted gaps in information and policy as well as shortcomings in participation and communication, which needed to be addressed as part of the way forward. during interviews with sanparks it was confirmed that the majority of these information gaps have been addressed since the completion of the sea, and that the shortcomings in consultation and participation were also addressed through the restructuring of the addo planning forum (knight 2004 pers. comm.). in terms of policy, the sea directly influenced the decision by sanparks to draft a resettlement policy as well as a land acquisition policy (kpi 15.1). most significantly, the sea influenced decisions relating to land acquisition (kpi 17.1). the best example was the decision by sanparks to abandon the prospect of purchasing land in the coastal dairy farming region, especially around alexandria (avis 2004 pers. comm.; knight 2004). this region is ideally suited to dairy farming and serves as one of the most productive dairy regions in the country. the sea public participation process pointed out that there were serious public objections against expansion of the park in this area (wren & msutu 2002). unfortunately, isolated cases were even found where farmers cleared indigenous forest to influence the biodiversity value of their land in an effort to make it unattractive for expropriation. the decision by sanparks to exclude this area from their short to mediumterm land acquisitions, took a lot of heat off the project (knight 2004 pers. comm.). but, the sea also provided information to sanparks on the biodiversity, social and economic aspects in the area. it showed retief.indd 118 2006/10/15 10:57:40 pm issn 0075-6458 119 koedoe 49/2 (2006) that land prices in some areas were high and rehabilitation would be time consuming and expensive, giving it a low priority from a conservation planning perspective. also, and probably more importantly, the financial scenarios indicated that the income generated through tourism and game farming (r103/ ha) could not compete with that generated through dairy farming (r177/ha) (antrobus 2002). the sea also highlighted the potential impact the park may have on the kommadagga, glenconner and kleinpoort areas which fall within some of the highest mohair producing areas in the world. although land purchase proposals in this area have not been abandoned by sanparks, they decided to investigate in more detail the potential impact of the park on mohair production. the question whether the sea influenced environmental quality was difficult to gauge since no monitoring (social or biophysical) had been conducted. however, the perception was that more land under conservation, due to the gaenp initiative, implies improved environmental quality from a conservation perspective (knight 2004 pers. comm.) (kpi 18.1). however, sanparks confirmed that, in retrospect, failure to address the social issues could have jeopardised the entire project (gordon 2004 pers. comm.). what remained unresolved was the need to formulate targets and indicators for the social and biophysical environments to gauge environmental quality from a sustainability, as well as conservation, perspective. the use of environmental reporting was being considered by sanparks and might form part of their management system in future (gordon 2004 pers. comm.; knight 2004). indirect outputs component the most significant indirect output of the sea was a raised awareness by sanparks towards social issues related to conservation. sanparks admitted that they were surprised about the social aspects and initially did not fully anticipate them (knight 2004 pers. comm.). this state of affairs might also reflect a general tendency amongst conservationists to underestimate social issues in decision making. the fact that sanparks has since improved their data and policy on the social environment as well as communication channels with communities is testimony that they are serious about the social aspect and intend to integrate them fully with their iems. according to sanparks the sea also raised awareness of the gaenp initiative with local authorities and ways of integrating the sea with local integrated development plans and spatial development frameworks were being investigated at the time (gordon 2004 pers. comm.) (kpi 15.2). however, this remained a difficult task since local authorities are under severe resource constraints. moreover, nobody seems to know whose responsibility it was to ensure the integration of the gaenp initiative with local authority decision making. finally, it was also confirmed that eia consultants have contacted sanparks to obtain the sea information to assist in their eias in the area (gordon 2004 pers. comm.). although this cannot be considered a formal tiering arrangement, it at least indicates awareness of the sea and that it was being considered as a source of background information (kpi 17.3). to address the gaps in information relating to tourism, sanparks initiated a study on the impact of the gaenp on provincial economy. with regard to the lack of information on farm occupancy, a detailed farm survey has also been conducted and a farm register established (gordon 2004 pers. comm.). finally, various information gaps were indicated in terms of the marine environment. initially, sanparks hoped to use the same management model for the terrestrial and marine components of the park but through the sea realised that this would not be feasible. subsequently a study to address information gaps on the marine environment has also been initiated (knight 2004 pers. comm.). conclusion the sea for the gaenp proved particularly challenging due to the extent and diversity of the planning domain, the national and international significance of the initiative, retief.indd 119 2006/10/15 10:57:41 pm koedoe 49/2 (2006) 120 issn 0075-6458 diverse interests in the region, and the fact that ultimately it involved the contentious issue of land ownership. however, notwithstanding some limitations it integrated well with strategic decision making and involved a comprehensive public participation process. ultimately, the research results showed that the sea achieved an average to good effectiveness performance where it influenced key controversial decisions on park expansion. the main lessons learned for application of sea to future park expansion initiatives are: firstly, to be effective, sea needs to be initiated as early as possible. because of the lack of integration with early conceptualisation the gaenp sea was limited, by its very design, to purposefully influence important strategic decisions concerning the vision and strategic objectives. moreover, its restrictive mandate (as reflected in the tor) and lack of legal framework further hampered implementation. it is evident that sea should ideally be conducted within a clearly defined regulatory and institutional framework to ensure that outcomes are considered and incorporated into decision making. secondly, from an environmental assessment point of view, the main impacts to consider relate to the social domain. as illustrated by the case study, the expansion of a conservation area mainly complements and improves the bio-physical environment, but could imply potentially significant social impacts. thus, although understanding the bio-physical environment is critically important from a park management and planning perspective, more prominence towards dealing with social issues are required from an environmental impact assessment perspective. a balance needs to be struck between conservation targets and socio-economic development priorities. thirdly, setting and communicating spatial boundaries is a key consideration for the expansion of conservation areas. the case study has shown that it is very difficult to engage with iaps if park expansion boundaries are not clarified up front and well communicated to those affected. vague boundaries create mistrust within the communities and hampers ultimate buy-in. rather, conservation targets and boundaries need to be communicated through a consultative process exploring alternative scenarios. finally, dealing with scale is important from a data gathering and ultimately data integration perspective. it is extremely difficult to consider trade-offs between bio-physical and social impacts if the scale and level of detail of the data differs. thus special consideration in the design of the sea methodology needs to be given towards aligning the different specialist studies. this could be addressed through the formulation of the tors and assigning comparable time and resources to different specialist inputs. based on the research results it can thus be concluded that sea is a useful tool to assists strategic decision making for park expansion. to take the debate forward and further explore the potential of sea as a decision support tool it is proposed that comparative case study research be conducted for a number of case studies. existing research and anecdotal evidence suggests that more than a dozen seas have been conducted in the conservation sector. such multiple case study research would allow for the identification of communalities and patterns between cases, leading towards increased generalisation and the potential to benchmark best practice. acknowledgements the contribution of all the various interviewees are gratefully acknowledged, with special reference to dr michael knight and jill gordon from sanparks, dr ted avis and michelle griffith from ‘coastal and environmental services’ as well as sandy wren from ‘sandy and mazizi consulting’. valuable inputs were also received from dr carys jones, university of manchester and dr stephen jay, sheffield hallam university. the author would like to thank the external reviewers for their insightful comments which added value to the revised version. retief.indd 120 2006/10/15 10:57:41 pm issn 0075-6458 121 koedoe 49/2 (2006) finally, appreciation to the commonwealth scholarship commission for their financial support without which, the research would not have been possible. references antrobus, g. 2002. strategic environmental assessment of the greater addo elephant national park—specialist report no.10—income from livestock farming. grahamstown: rhodes university. brownlie, s., c, de villiers, a. driver, n. job, a. von hase & k. maze. 2005. systematic conservation planning in the cape floristic region and succulent karoo, south africa: enabling sound spatial planning and improved environmental assessment. journal of environmental assessment, policy and management 7(2): 201–228 cashmore, m., r. gwilliam, r. morgan, d. cobb, d. & a. bond. 2004. the interminable issue of effectiveness: substantive purposes, outcomes and research challenges in the advancement of environmental impact assessment theory. impact assessment and project appraisal 22(4): 295–310 coastal and environmental services (ces). 2002. strategic environmental assessment of the greater addo elephant national park. grahamstown: coastal and environmental services. connor, t. 2002. strategic environmental assessment of the greater addo elephant national park—specialist report no. 6—social assessment. port elizabeth: south african national parks. council for scientific and industrial research (csir). 1996. strategic environmental assessment (sea)—a primer. stellenbosch: council for scientific and industrial research. council for scientific and industrial research (csir). 2001a. final report on the proposed marine protected area for the greater addo elephant national park aquatic conservation planning. stellenbosch: council for scientific and industrial research. council for scientific and industrial research (csir). 2001b. preliminary survey and desktop approach to conservation planning of freshwater ecosystems in the greater addo elephant national park. stellenbosch: council for scientific and industrial research. council for scientific and industrial research (csir). 2002. final report: conservation planning framework for the greater addo elephant national park. stellenbosch: council for scientific and industrial research. dalal-clayton, b. and sadler, b. 2005. strategic environmental assessment: a sourcebook and reference guide to international experience. london: earthscan. davies, r. 2002. strategic environmental assessment of the greater addo elephant national park—specialist report no. 11—ecotourism scenarios. port elizabeth: south african national parks. department of environmental affairs and tourism (deat). 2000. strategic environmental assessment (sea) in south africa: guideline document. pretoria: department of environmental affairs and tourism. department of environmental affairs and tourism (deat). 2004a. national biodiversity strategy and action plan. pretoria: department of environmental affairs and tourism. department of environmental affairs and tourism (deat). 2004b. strategic environmental assessment, integrated environmental management, information series 10. pretoria: department of environmental affairs and tourism. geach, b. 2002. strategic environmental assessment of the greater addo elephant park—specialist report no.9—economic activities in the planning domain. johannesburg: south african national parks. glazewski, j. 2005. environmental law in south africa. cape town: butterworths. international association for impact assessment (iaia). 2002. strategic environmental assessment performance criteria. iaia special publication series no.1, international association for impact assessment: fargo. jones, c., m. baker, j. carter, s. jay, m. short & c. wood. 2005. strategic environmental assessment and land use planning: an international evaluation. london: earthscan. kerley, g. & a. boshoff. 1997. a proposal for a greater addo national park—a regional and national conservation and development opportunity. terrestrial ecology research unit, university of port elizabeth. port elizabeth. kolhoff, a. & r. slootweg. 2005. biodiversity in sea for spatial plans—experiences from the netherlands. journal of environmental assessment, policy and management 7(2): 267–286 lawrence, d. 1997. quality and effectiveness of environmental impact assessments: lessons and insights from ten assessments in canada. project appraisal 12(4): 219–232 lochner, p., a. weaver, c. gelderblom, p. raewyn, t. sandwith & s. fowkes. 2003. aligning the diverse: the development of a biodiversity conservation strategy for the cape floristic region. biological conservation 112: 29–43 marsden, s. 1998. importance of context in measuring the effectiveness of strategic environmenretief.indd 121 2006/10/15 10:57:41 pm koedoe 49/2 (2006) 122 issn 0075-6458 tal assessment. impact assessment and project appraisal 16(4): 255–266 miles, m. & a. huberman. 1994. an expanded source book: qualitative data analysis. london: sage. national botanical institute (nbi). 2002. mainstreaming biodiversity at the bioregional scale. parallel workshop event at the world summit on sustainable development, august 2002. national botanical institute: johannesburg. owens, s., t. rayner & o. bina. 2004. new agendas for appraisal: reflections on theory, practice, and research. environment and planning 36: 1943–1959 partidario, m. & j. arts. 2005. exploring the concept of sea follow-up. impact assessment and project appraisal 23(3): 246–257 retief, f. 2005 quality and effectiveness of strategic environmental assessment (sea) in south africa. ph.d. thesis, school of environment and development. university of manchester, manchester. rossouw, n., m. audouin, p. lochner, s. heather-clark, & k. wiseman. 2000. development of strategic environmental assessment in south africa. impact assessment and project appraisal 18(3): 217–223 rossouw, n. & k. wiseman. 2004. learning from the implementation of environmental public policy instruments after the first ten years of democracy in south africa. impact assessment and project appraisal 22(2): 131–140 sadler, b. 1996. international study of the effectiveness of environmental assessment—final report. environmental assessment in a changing world: evaluating practice to improve performance. ottawa: international association for impact assessment and the canadian environmental assessment agency. sadler, b. 2004. on evaluating the success of eia and sea. pp. 248–285. in: morrison-saunders, m. & j. arts (eds.). assessing impact: handbook of eia and sea follow-up. london: earthscan. schmidt, m., e. joao & e. albrecht. 2005. implementing strategic environmental assessment. new york: springer. terrestrial and ecology research unit (teru). 1999. report—greater addo national park— stakeholder workshop, 23 february 1999. terrestrial and ecology research unit, university of port elizabeth. port elizabeth. therivel, r. & p. minas. 2002. ensuring effective sustainability appraisal. impact assessment and project appraisal 20(2): 81–91 therivel, r. & m. partidario. 2000. the future of sea. pp. 271–280. in: partidario, m. & r. clark (eds). perspectives on strategic environmental assessment. boca raton: crc press. thissen, w. 2000a. criteria for the evaluation of sea. pp. 113–127. in: partidario, m. & r. clark (eds). perspectives on strategic environmental assessment. boca raton: crc press. thissen, w. 2000b. strategic environmental assessment at a crossroads. impact assessment and project appraisal 18(3): 174–176 timmermans, h. & l. sisitka. 2002. strategic environmental assessment of the greater addo elephant national park—specialist report no. 14—institutional review. grahamstown: institute of social and economic research, rhodes university. todd, s. 2001. measuring effectiveness of environmental dispute settlement efforts. environmental impact assessment review 21: 97–110 treweek, j., r. therivel, s. thompson & m. slater. 2005. principles for the use of strategic environmental assessment as a tool for promoting the conservation and sustainable use of biodiversity. journal of environmental assessment, policy and management 7(2): 173–199 verheem, r. & j. tonk. 2000. strategic environmental assessment: one concept multiple forms. impact assessment and project appraisal 18: 177–182 world bank. 2002. project brief—funding for the establishment of the greater addo elephant national park. johannesburg: world bank. world conservation monitoring centre. 1992. global biodiversity: status of the earth’s living resources. london: chapman and hall. world conservation monitoring centre. 2002. world atlas of biodiversity—earth’s living resources in the 21st century. california: university of california press. wren, s. & m. msutu. 2002. strategic environmental assessment of the greater addo elephant national park—specialist study no 5—public participation. port elizabeth: sandi & mazizi consulting. wynberg, r. 2002. a decade of biodiversity conservation and use in south africa: tracking progress from the rio earth summit to the johannesburg world summit on sustainable development. south african journal of science 98: 233–243 yin, r. 2003. case study research: design and methods–applied social research methods series. volume 5. london: sage. retief.indd 122 2006/10/15 10:57:41 pm issn 0075-6458 67 koedoe 49/2 (2006)issn 0075-6458 67 koedoe 49/2 (2006) introduction one corollary of the preservation of natural habitats is an increased probability of conflict between conservation objectives and other land-use practises in the vicinity. while the exact nature and extent of this conflict will obviously be locally determined (hill 2000), it stems ultimately from the fact that conservation areas are designed to serve as reservoirs for both flora and fauna. in the south african context, primates are regarded as particularly problematic residents of protected areas, as conventional fences do not readily enclose them, their eclectic diets allow them to benefit from a range of agricultural endeavours, and they are behaviourally opportunistic (von dem bussche & van der zee 1985; cowlishaw & dunbar 2000). where their depredations are not tolerated, attempted extirpation is the most common population structure and habitat use of baboons (papio hamadryas ursinus) in the blyde canyon nature reserve a.j. marais, l.r. brown, l. barrett and s.p. henzi marais, a.j., l.r. brown, l. barrett and s.p. henzi. 2006. population structure and habitat use of baboons (papio hamadryas ursinus) in the blyde canyon nature reserve. koedoe 49(2): 67–76. pretoria. issn 0075-6458. baboons are highly intelligent and ecologically flexible animals with attributes that allow them to exploit diverse habitats. as a result of their dietary flexibility they often exploit human habitats, causing damage to crops and forest plantations as well as to human dwellings. in the south african context this has led to baboons being regarded as problem animals and attempted extirpation is the most common approach to the damage they cause. this perception of and attitude toward baboons gives many conservationists cause for concern since all southern african cercopithecine primates are cites listed and it has not been proven that this strategy is the best long-term solution. as part of a research programme focusing on the damage done by chacma baboons in pine plantations along the drakensberg escarpment in mpumalanga, a single troop in the blyde canyon nature reserve was studied to describe their patterns of habitat use. vegetation and habitat surveys were conducted within the home range of the troop. the troop was habituated and each member’s activity, location and food items utilised were recorded over a 12 month period. the results of this study indicate that baboons utilised plant communities based on food production and availability rather than size in hectares. the results also indicate that the group size, foraging and food search strategies of this troop resembles that of the drakensberg troops previously studied. the study troop employs two different forage modes of engagement depending on where they choose to forage while they avoid utilising an easily accessible pine plantation. due to the troop’s long inter-birth intervals it is likely that the current forestry practice of extirpation may have a negative influence on baboon population viability in these areas. key words: chacma baboons, mpumalanga, home range, population structure, habitat use. a.j. marais and l.r. brown, applied behavioural ecology and ecosystem research unit, university of south africa; private bag x6, florida, 1710 republic of south africa; l. barrett and s.p. henzi, applied behavioural ecology and ecosystem research unit, university of south africa, (present address: department of psychology, university of central lancashire). marais.indd 67 2006/10/15 10:54:42 pm koedoe 49/2 (2006) 68 issn 0075-6458 approach to the damage they cause. this is problematic for three reasons. first, all southern african cercopithecine primates are cites listed (appendix ii). second, there is as yet little evidence, in the absence of complete and permanent removal, that hunting and trapping are the most cost-effective strategies in the medium to long term. third, there is little understanding of its effect on the refugial populations themselves. it is possible that control measures compensate for the reduction in predator numbers often associated with conservation islands (cowlishaw & dunbar 2000), thereby allowing surrounding areas to serve as demographic sinks. it is, however, at least equally likely that such populations are not sustainable and, where this is so, the integrity of the refugium is threatened. any effective policy for both conservation and problem animal control must address these issues and doing so clearly requires an enumeration of the extent of the problems. our own research focuses on the damage done by chacma baboons in pine (pinus spp.) plantations along the drakensberg escarpment in mpumalanga province, south africa. advantages of rainfall and climate in this area have led to very large commercial plantings, while the escarpment itself and, especially, nature reserves in the area harbour baboon populations that generally have easy access to plantations. baboons strip the bark of pine trees, thereby disrupting growth and preventing trees from being harvested (bigalke & van hensbergen 1990). they have the potential to make large inroads into the profitability of plantations, which in turn has consequences for local conservation policies. estimating costs accurately, and determining viable solutions, however, which is our overall objective, must begin with an understanding of the natural structure and ecology of the baboon populations involved. this is necessitated both by the need to predict the effects of control measures as well as the observation that not all trees are damaged and not all baboon troops that enter plantations 23 os bourke ’s luck section blyde canyon nature reserve mpumalanga province botswana republic of south africa namibia fig. 1. a map of the study site’s location within south africa and mpumalanga province (shaded). the dotted line indicates the approximate location of the drakensberg escarpment. marais.indd 68 2006/10/15 10:54:42 pm issn 0075-6458 69 koedoe 49/2 (2006) are destructive in this way. our aims in this paper, accordingly, are to outline the structure and density of the baboon population in the blyde canyon nature reserve (bcnr) and to describe the patterns of habitat use of one representative, habituated troop. the reserve has commercial plantations on its boundary and the troop was chosen because it had, potentially, easy access to these plantations. study area blyde canyon nature reserve is a 280 km² provincial reserve in mpumalanga province running along the escarpment that divides the south african highveld from the lowveld (fig. 1). the vegetation is primarily a mixture of mountain sourveld (acocks veld type 8) and mixed bushveld (acocks veld type 18) (acocks 1988) and as north-eastern mountain grassland (type 43) by bredenkamp et al. (1996) which reflects the rugged topography and altitudinal gradient (600 m– 2000 m asl). it is distinctive in containing populations of all five non-human primates that occur in southern africa as well as their most important predators (python, crowned eagle and, especially important for baboons, the leopard). the climate along the escarpment is mild. the mean temperature of the hottest months (january; february) is 21.3 ºc, while that of the coldest month (june) is 11.2 ºc (fig. 2). absolute temperatures vary between a maximum of 34.1 ºc (february) to -2 ºc (june). the months of may, june, july and august are normally the coldest months with early morning temperatures ranging between -2 ºc to 2 ºc but increasing in midday to between 26.8 ºc and 28 ºc. during the warmer months (december, january, february, march) the temperatures vary between 8.6 ºc and 14 ºc in the morning and 30.2 ºc and 34.1 ºc in the middle of the day. rainfall is the primary driving force influencing the productivity of vegetation and therefore food available to animals. mean annual rainfall for the study area is 848 mm, with most precipitation occurring in the austral spring and summer (marais 2005). we used r ai nf al l ( m m ) te m pe ra tu re ºc month 200 160 120 80 40 0 jul aug sep oct nov dec jan feb mar apr may jun 35 30 25 20 15 10 5 0 these data to divide the year into a wet (october–april) and dry season (may–september) for analysis. mild mist would normally only be encountered during the raining season, but is restricted to the canyon sections only. methods vegetation description and mapping. the vegetation of the home range of the troop was stratified into physiognomicphysiographic units using 1:20 000 stereo aerial photographs. to ensure that all variations in the vegetation were considered and sampled, a total of 61 sample plots (200 m²) were located on a randomly stratified basis within the different units (bredenkamp 1982; bezuidenhout 1993). vegetation and habitat surveys were conducted in each of the sample plots by recording all plant species present. the cover of the tree, shrub and herbaceous layers was estimated using the braun–blanquet cover abundance scale (mueller-dombois & ellenberg 1974). population survey the baboon population of the bcnr was surveyed during bi-monthly walks along pre-determined routes. when troops were seen, they were counted and their location marked on a 1:50 000 topographical map. once the troops had been identified in this way, repeated sightings enabled confirmation of size and the estimation of home range size by the fitting of minimum convex polygons fig. 2. climate of the study area. marais.indd 69 2006/10/15 10:54:42 pm koedoe 49/2 (2006) 70 issn 0075-6458 to the sightings of each troop using commercial gis software (arcinfo). each troop was allocated to one of four visually identifiable habitat types (grassland, wetland/grassland, protea grassland and mixed bushveld) within the broad acocks categories. study troop habituation the study troop at bourke’s luck (n = 18) was chosen because it was close to average size (see below) and because its home range extended at least to the reserve boundary. the troop was habituated by following it on foot until it could be observed at a distance of 30 m. during habituation, data were collected on diet. once it was habituated we recorded activity and location (as well as food items) over a 12-month period from june 2001 to may 2002. data collection activity budgets were determined through 15-minute scan samples of all visible group members. the age and sex of each animal was recorded together with its state (moving, resting, foraging, socialising). data were collected using a hand-held data logger (handspring) and commercial software (pendragon forms). during all-day follows, the location of the troop was recorded using a gps logger whenever the estimated centre of mass shifted by more than 20 m. these data were downloaded into a commercial gis package and linked to a fine-grained habitat map of the home range. this allowed us to identify the habitat type occupied by the troop at each point sample. home range area was estimated by fitting a minimum convex polygon to the points. day range and habitat use information comes from 25 complete day ranges. data analysis data were downloaded from the data logger into a commercial spreadsheet package and exported to spss for analysis. we used the shapiro-wilk test to determine the normality of the data and found no significant devia0 10 20 30 40 1.0 2.0 2.5 3.0 group size d en si ty (in d/ km ²) 1.5 fig. 3. the relationship between group size and population density. the best-fit regression line (density = 0.042size + 0.756) is provided together with its 95 % confidence limits (dashed lines). the study troop is indicated by an open circle. marais.indd 70 2006/10/15 10:54:42 pm issn 0075-6458 71 koedoe 49/2 (2006) tions from the normal distribution. all tests were two-tailed with alpha set at 0.05. the floristic data was analysed according to braunblanquet procedures using turboveg (hennekens 1996). by applying the two-way indicator species analysis (twinspan) (hill 1979) to the floristic data, a first approximation of the main plant communities was derived. further refinement of the classification was achieved by braun-blanquet procedures (bredenkamp et al. 1989; kooij et al. 1990; bezuidenhout 1993; eckhardt 1993; brown & bredenkamp 1994). results population structure and density. twentyone troops were counted, ranging in size from 9–37 animals. the mean troop size was 18.3 animals (± 6.8 sd). troop size did not differ with recorded habitat type (f3,17 = 1.35; ns). one small troop, using a tourist residential zone, had a home range area that was significantly smaller than expected for its size (centred leverage value > 0.5) and was excluded from the following analyses. the mean estimated home range area was 10.2 km² (± 2.3 sd). there was a significant positive correlation between troop size and estimated home range area (r = 0.81; n = 20; p < 0.01). density was calculated as home range area/ group size. the mean estimated population density was 1.8 baboons/km² (± 0.4 sd). this was also positively correlated with group size (r = 0.76; n = 20, p < 0.01) (fig. 3). day-journey length. the average annual day-journey length for the bourke’s luck troop during the study period was 3.37 km (± 0.73 sd; n = 25). the mean daily dry season distance was 3.78 km (± 0.74 sd; n = 12), while the mean wet season distance was 2.99km (±0.48 sd; n = 13). this difference in seasonal day journey length was significant (t23 = 3.16; p < 0.01). day and home range size. the baboons utilised, on average, 34.47 ha per observation day (± 20.31 sd), resulting in a recorded annual home range size of 10.35 km² (fig. 4 – home range and day ranges). the mean day range area for the wet season was 31.44 ha (n = 13 ± 10.56 sd) while that for the dry season was 52.3 ha (n = 12 ± 23.0). this seasonal difference was significant (t23=2.95; p < 0.01). there was, overall, a positive correlation between day journey length and day journey area (r = 0.69; n = 25; p < 0.01). home range overlap. two other baboon troops (n1=18; n2=25) overlapped with the bourke’s luck troop. the percentage overlap for both troops combined was relatively small at 7.5 %. habitat use. topographically, the troop’s home range varied from gentle to relatively moderate slopes of 2–6° on the higher lying areas to valley bottoms where slopes were as steep as 60°. the habitat covered by the study troop’s home range comprises six major plant communities, some with recognisable sub-communities or variants, generating a total of 12 distinctive floristic zones (fig. 4) (marais 2005). the first point to note from fig. 4 is that despite the presence of a large pine plantation the troop chose not to enter it; in fact, they went right up to it on at least two occasions and then turned away. given this, we calculated the proportion of the annual home range contributed by each of the remaining plant communities (fig. 4) and used the electivity index (ei) (krebs 1989) to determine whether the study troop occupied plant communities only in relation to their availability or whether some were preferred and others avoided. the ei describes habitat as a score between -1 and +1, where 0 indicates that a community is used in accordance with its availability, while plus and minus signs signal that communities are favoured or avoided respectively. the data show, overall, that the baboons had distinct preferences for certain plant communities and avoided others (fig. 5). in particular, there was a very high preference for the englerophytum magalismontanumacacia ataxacantha woodland (community 6.1), which is the smallest plant community present at bourke’s luck, covering an area of only 9.3 ha (1.8 % of the home range). the baboons also showed a preference for marais.indd 71 2006/10/15 10:54:42 pm koedoe 49/2 (2006) 72 issn 0075-6458 n fi g. 4 . t he h om e ra ng e of th e st ud y tr oo p su pe ri m po se d on th e id en tif ie d ve ge ta tio n co m m un iti es ( a) . t he d ay r an ge s us ed to d er iv e th e es tim at e of h om e ra ng e ar e di vi de d in to th os e fr om e ith er th e dr y (b ) o r t he ra in y (c ) s ea so ns . t he b ar in di ca te s 1k m . n um be rs in f ig . 4 a re fe r t o th e fo llo w in g pl an t c om m un iti es a nd s ub -c om m uni tie s, a s id en tif ie d by m ar ai s (2 00 5) : ( 1) . h yp er th el ia d is so lu ta -h et er op og on c on to rt us g ra ss la nd ; ( 2) . d ih et er op og on a m pl ec te ns -l ou de tia s im pl ex g ra ss la nd ; ( 2. 1) . d ih et er op og on a m pl ec te ns -m on oc ym bi um c er es iif or m e g ra ss la nd ; ( 2. 1. 1) . h em iz yg ia tr an sv aa le ns is -t he m ed a tr ia nd ra g ra ss la nd ; ( 2. 1. 2) . d ih et er op og on a m pl ec te ns p ea rs on ia s es si lif ol ia s hr ub la nd ; (2 .2 ). h el ic hr ys um w ilm si ip an ic um n at al en se g ra ss la nd ; (2 .3 ). d ih et er op og on a m pl ec te ns -b ra ch ia ri a se rr at a g ra ss la nd ; (2 .3 .1 ). e ra gr os tis g um m ifl ua -l ou de tia s im pl ex g ra ss la nd ; (2 .3 .2 ). (2 .4 ). f ad og ia t et ra qu et ra -e uc le a lin ea ri s sh ru bl an d; s en ec io g la be rr im us -p ea rs on ia s es si lif ol ia s hr ubl an d; ( 3) . e ng le ro ph yt um m ag al is m on ta nu m -h el ic hr ys um k ra us si i s hr ub la nd ; ( 4) . p te ro ca rp us a ng ol en si se ng le ro ph yt um m ag al is m on ta nu m w oo dl an d; ( 5) . f au re a sa lig na c ym bo po go n va lli du s w oo dl an d; ( 6) . c om br et um k ra us ii a ca ci a at ax ac an th a w oo dl an d; ( 6. 1) . e ng le ro ph yt um m ag al is m on ta nu m a ca ci a at ax ac an th a w oo dl an d; ( 6. 2) . c om br et um k ra us si i a ca ci a at ax ac an th a w oo dl an d; ( 7) . p in us p at ul a w oo dl an d. marais.indd 72 2006/10/15 10:54:43 pm issn 0075-6458 73 koedoe 49/2 (2006) combretum kraussii-acacia ataxacantha woodland (community 6.2), which covers a relatively large area (55 ha—10.6 % of home range). together, these findings suggest that although the baboons are most attracted to the acacia woodland they prefer a specific sub-community within the woodland to the other sub-community and not acacia woodland in general. the faurea saligna–cymbopogon validus woodland (community 5) comprising 18.8 ha (3.6 % of home range) was slightly less preferred than the other woodland communities, but still a preferred community and utilised throughout the year. the next most preferred habitats consisted of grassland and shrubland (communities 2.3 and 2.4) (42.5 ha—8.1 % of home range and 14.6 ha—2.8 % of home range respectively). the least-preferred communities were the hyperthelia dissoluta–heteropogon contortus grassland (community 1) and particularly, the helichrysum wilmsii–panicum natalense grassland (community 2.2). the baboons mostly preferred only 140.2 ha (26.9 %) of their home range, showing the highest preference for the smallest community namely the englerophytum magalismontanum-acacia ataxacantha woodland (community 6.1) which relates to food production and availability rather than plant community size. qualitatively, the only observed seasonal effect was for community 3 (englerophytum magalismontanum-helichrysum kraussii shrubland), which was avoided during the wet season and sought out during the dry (fig 5b). -0.8 -0.6 -0.4 -0.2 0.0 0.2 0.4 0.6 0.8 1 2.1 2.2 2.3 2.4 3 4 5 6.1 6.2 plant community e le ct iv ity in de x -0.8 -0.6 -0.4 -0.2 0.0 0.2 0.4 0.6 0.8 e le ct iv ity in de x 1 2.1 2.2 2.3 2.4 3 4 5 6.1 6.2 plant community a b -0.8 -0.6 -0.4 -0.2 0.0 0.2 0.4 0.6 0.8 1 2.1 2.2 2.3 2.4 3 4 5 6.1 6.2 -0.8 -0.6 -0.4 -0.2 0.0 0.2 0.4 0.6 0.8 1 2.1 2.2 2.3 2.4 3 4 5 6.1 6.2 plant community b fig. 5. the overall utilisation of plant communities in relation to their availability (a), expressed as an electivity index, together with seasonal preferences (b) for both the dry (open bars) and wet seasons (black bars). plant community numbers are the same as those used in fig. 4. marais.indd 73 2006/10/15 10:54:43 pm koedoe 49/2 (2006) 74 issn 0075-6458 activity budgets. the allocation of time to each of the four behavioural states was determined as a percentage of the total activity budget. overall, the baboons spent 62.3 % of their time foraging, 24.3 % moving and 10.2 % resting. the remainder was allocated to social interaction. no seasonal differences were noted for foraging (wet: 62.5 %; dry 62.0 %), moving (wet: 23.1 %; dry: 26.3 %) or resting (wet: 10.6 %; dry: 9.6 %). discussion comparison with data from other chacma study sites (stoltz & saayman 1970; davidge 1978; watson 1985; gaynor 1995; henzi & lycett 1995; hill 1999) reveals that in their low densities, small mean group size, high investment in foraging and food search strategies that result in relatively short day ranges, the bcnr baboons bear the closest resemblance to the drakensberg mountain population (population density = 2 ind/km²; mean group size = 21.4; percentage time feeding = 68 %; mean day range = 4.6 km) (henzi & lycett 1995; henzi et al. 1997). together, these two populations are outliers in a subspecies that is, itself, characterised by small group sizes and high foraging demands (henzi et al. 1999). the similarity is not surprising since both populations are subject to the direct and indirect effects of the elevation and relief of the drakensberg escarpment, which result in high thermal demand and low food availability, at least in winter (henzi et al. 1992). unlike the natal drakensberg population, though, the bcnr baboons forage in an environment that, by virtue of its lower latitude, is characterised by a much greater variability of habitat types within the home range (henzi et al. 1992). our results indicate that this corresponds to a greater variability in utilisation with a particular, but not surprising, preference for woodland, as this is likely to yield a better return for foraging effort (hill & dunbar 2002). while we will consider this further elsewhere (brown et al. 2005), there are two issues that can be pursued now. the first is that the baboons clearly employ two different modes of engagement with their home range that is conditional on where they choose to forage on any one day. successful foraging in grassland requires careful searching for cryptic food items and results in uniform, slow travel speeds and small day ranges 10 20 30 40 50 60 70 fitted value (ha) -3 -2 -1 0 1 2 3 s td .r es id ua l 10 20 30 40 50 60 70 -3 -2 -1 0 1 2 fig. 6. the relationship between fitted values for area utilised (ha) and the standardised residuals for day journey length. marais.indd 74 2006/10/15 10:54:43 pm issn 0075-6458 75 koedoe 49/2 (2006) (henzi et al. 1992; henzi et al. 1997), whereas utilisation of woodland generally involves greater variation in travel speed and distance travelled as animals move to a foraging area where they can subsequently obtain necessary resources in a small, circumscribed zone (gaynor 1995). the general correlation we found between day-journey length and dayjourney area masks an additional relationship between the absolute size of the standardised residual and day-journey area that supports this (fig. 6) (rho = 0.69; n = 25; p < 0.01). figure 6 indicates that longer day journeys, a characteristic of the dry season, were associated with the exploitation of either very small or very large areas. in our reading, and given the absence of seasonal effects in activity patterns, this suggests that the baboons were both moving and foraging consistently but slowly in small areas of grassland or moving rapidly over some distance (hence a large, positive residual) and then increasing their intake rate rapidly within a circumscribed area of relatively high food availability. the point is that the actual foraging zone of the troop—once travel is factored out—is always likely to be relatively small. sample size precludes us investigating this but it remains a topic for further, more detailed analysis that considers exploitation in relation to the mix of habitat types encountered. the second issue is that, despite the damage that they cause elsewhere, the study troop chose to avoid an easily accessible pine plantation. this avoidance, especially given the positive correlation between group size and number of group members/km² that suggests some local crowding, is telling and confirms strum’s point (1994) that raiding by baboons is not inevitable, nor is its occurrence predictable from gross local features. one or both of two possible reasons suggest themselves: either the baboons have available to them outside the plantation all the keystone resources they need or they target only forestry compartments of particular age/ size classes. determining how these alternatives might interact requires comparative data and we have now expanded the study to include troops that do make use of plantations. results will be reported elsewhere. lastly, one implication for conservation policy stems from the similarity of the bcnr baboon profile to that of the drakensberg population. long-term data from the drakensberg reveals that recruitment to the baboon population is low, with female reproductive rate constrained by climate (henzi & lycett 1995; lycett et al. 1998). if, as seems likely, baboons of the mpumalanga escarpment have comparably long inter-birth intervals, then the current forestry management practise of extirpation may reduce the long-term viability of baboon populations in protected areas. acknowledgments we thank the nrf, global forest products and komatiland forests for funding, and mpumalanga parks board for permission to conduct the study. references acocks, j.p.h. 1988. veld types of south africa. memoirs of the botanical survey of south africa 57: 1–146. bezuidenhout, h. 1993. syntaxonomy and synecology of the western transvaal grasslands. ph.d. thesis, university of pretoria, pretoria. bredenkamp, g.j. 1982. ‘n plantekologiese studie van die manyeleti-wildtuin. d.sc. thesis. university of pretoria, pretoria. bredenkamp g.j., e. granger & n. van rooyen. 1996. north-eastern mountain grassland. p. 46. in: low, a.b. & a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. bredenkamp, g.j., a.f., joubert & h. bezuidenhout. 1989. a reconnaissance survey of the vegetation of the plains in the potchefstroomfochville-parys area. south african journal of botany 55: 199–206. bigalke, r.c. & h.j. van hensbergen. 1990. baboon damage in plantation forestry in south africa. south african forestry journal 152: 26–33. brown, l.r. & g.j. bredenkamp. 1994. the phytosociology of southern section of the borakalalo nature reserve, south africa. koedoe 37(2): 59–72. brown, l.r., h. marais, s.p. henzi & l. barrett. 2005. vegetation classification as the basis for baboon management in the bourke’s luck marais.indd 75 2006/10/15 10:54:43 pm koedoe 49/2 (2006) 76 issn 0075-6458 section of the blyde canyon nature reserve, mpumalanga. koedoe 48(2): 71–92. cowlishaw, g. & r.i.m. dunbar. 2000. primate conservation biology. cambridge: cambridge university press. davidge, c. 1978. activity patterns of chacma baboons (papio ursinus) at cape point. zooogica africana 13: 143–155. eckhart, h.c. 1993. a synecological study of the vegetation of the north-eastern orange free state. m.sc. thesis. university of pretoria, pretoria. gaynor, d. 1995. foraging and feeding behaviour of chacma baboons in a woodland habitat. ph.d. thesis, university of natal. hennekens, s.m. 1996. megatab: a visual editor for phytosociological tables. ulft: giesen. henzi, s.p. & j.e. lycett. 1995. population structure, dynamics and demography of mountain baboons. american journal of primatology 35: 155–163. henzi, s.p., r.w. byrne & a. whiten. 1992. patterns of movement by baboons in the drakensberg mountains: primary responses to the environment. international journal of primatology 13: 601–629. henzi, s.p., j.e. lycett, a. weingrill, r. byrne & a. whiten. 1997. the effect of troop size on travel and foraging in mountain baboons. south african journal of science 93: 333–335. henzi, s.p., a. weingrill & l. barrett. 1999. male behaviour and the evolutionary ecology of chacma baboons. south african journal of science 95: 240–242. hill, c.m. 2000. conflict of interest between people and baboons: crop raiding in uganda. international journal of primatology 21: 299–315. hill, m.o. 1979. twinspan: a fortran programme for arranging multivariate data in an ordered two-way table classification of individuals and attributes. new york: cornell university. hill, r.a. 1999. ecological and demographic determinants of time budgets in baboons: implications for cross-populational models of baboon sociobiology. phd thesis, university of liverpool. hill, r.a. & r.i.m. dunbar, 2002. climatic determinants of diet and foraging behaviour in baboons. evolutionary anthropology 16: 579–593. kooij, m.s., g.j. bredenkamp & g.k. theron. 1990. classification of the vegetation of the b land type in the north-western orange free state. south african journal of botany 56: 309–318. krebs, c.j. 1989. ecological methodology. new york: harper and row. lycett, j.e., s.p. henzi & l. barrett. 1998. maternal investment in mountain baboons and the hypothesis of reduced care. behavioral ecology and sociobiology 42:49–56. marais a.j. 2005. resource utilisation of the chacma baboon in different vegetation types in northeastern mountain sour veld, blyde canyon nature reserve. m.tech dissertation, university of south africa. mueller-dombois, d. & h. ellenburg. 1974. aims and methods of vegetation ecology. new york: wiley. stolz, l.p. & g.s. saayman. 1970. ecology and behaviour of baboons in the northern transvaal. ann. transvaal museum 26, 99–143. strum, s.c. 1994. prospects for management of primate pests. terre et vie 49: 295–306. von dem bussche, g.h. & d. van der zee. 1985. damage by samango monkeys to pine trees in the northern transvaal. south african forage journal 133: 43–48. watson, l.h. 1985. the feeding ecology of chacma baboons and vervet monkeys in the hluhluweumfolozi reserve. m.sc. thesis, university of natal. marais.indd 76 2006/10/15 10:54:44 pm filelist convert a pdf file! koedoe 19: 83-88 (i9 76) growth and development of the impala aepyceros melampus n. fairall department oj nature conservation p. o. box 659 cape town 8000 h. h. braack division oj nature conservation private bag x 404 skukuz.a jj50 introduction growth and development of the prenatal impala has been described by fairall (1969) and roettcher, hoffman and kayanja (1970) while weights and measurements of the mature animal have been reported by sachs (1967) and talbot and mcculloch (1965). child (1964) has investigated growth in this species. this paper serves as a baseline for comparison of future changes in morphology of the species, as the animals were all obtained at the beginning of the culling programme. material and methods impala became available in the kruger national park as a result of culling operations and for a separate study project on the reproductive physiology of this animal (fairall 1972). mass and measurements of 300 animals were analysed and this information was used to quantify growth and size for this species from the park. all animals were shot and transported to a central point where they were weighed. some measurements were taken in the field, the majority were, however, taken when the animals were weighed. mass was determined with the aid of a spring balance. standard measurements taken were: length of vertebral column from the atlas joint to the sacrococcygeal joint; heart girth measured directly behind the shoulder; and shoulder height taken from the tip of the hoof to the highest point of the shoulder. measurements were all taken along the body curves. the lambing season of the impala in this area is very clearly defined (fairall 1968, 1972). ages of immature animals were, therefore, deduced from the months in which they were shot. females can be accurately 83 aged up to two years and males up to three years by using horn size and shape as additional ageing criteria. in attempting to derive a method to determine mass from body measurements, regressions of each separate measurement and multiple regressions of all combinations were tested using standard methods. results mean mass and measurements are shown in table i and growth is illustrated graphically in fig. 1. the standard deviation of mass should be noted, as this shows a deviation of between 6 to 11 percent. the breaks in the growth curves at about eight months are also interesting and can be compared with the curves presented by child (1964). simple linear regression using heart girth as independent variable, was found to give as accurate a representation of the relation between mass and measurement as any combination; it was, therefore, used. the growth curves in fig. 1 show that it would be unrealistic to fit one linear equation over the whole curve. separate regressions for ages under and above 12 months were therefore calculated . there is very little difference between the sexes of the immature animals and the linear function for both is adequately represented by mass = 0,8744 heart girth 31,1218. this has a coefficient of determination of 0,86. mature growth difhts between the sexes and two linear functions describe the relationship: for males the line is defined by mass = 0,9746 heart girth 30,7913 and for females by mass = 0,5180 heart girth + 0,2926; the coefficient being 0,71 and 0,93 respectively. table 2 is an attempt to use this information in a practical manner. for any measured heart girth the equivalent mass may be determined direc tl y. 50 0male .. female 40 ----------------------. .. e ~ ! 30 10 6 8 10 12 18 24 36 age in months fig. i. growth curves of kruger national park impala . 84 00 '" age 6 months 8 months 10 months 12 months 18 months 24 months 36 months mature old mass ± 25,00 2,19 22,47 3,65 25,93 2,99 32,59 1,07 35,24 4,33 36,72 1,76 40,58 2,21 49,22 1,02 47,80 2,19 table 1 mean mass and measurements of kruger national park impala with their standard deviations male impala female impala length ± shoulder ± heart ± mass ± length ± shoulder ± heart ± height girth height girth 77,69 2,18 74,65 1,90 64,64 2,97 24,32 1,63 7s,1 1,57 73,03 1,51 63,33 1,88 79,48 8,72 74,46 3,28 65,42 9,70 23,00 2,31 75,78 4,34 72,63 3,12 63,33 3,29 si,02 1,72 75,28 3,21 64,15 2,95 23,03 3,63 80,73 4,39 75,4s 3,22 62,36 3,12 s7,78 1,16 80,50 0,74 71,26 0,91 27,34 1,01 s5,16 1,57 7s,42 0,93 67,93 1,41 91,04 3,03 83,16 1,96 72,41 2,18 31,s9 1,95 88,45 2,75 80,24 1,55 69,43 1,36 90,00 2,06 83,22 1,12 73,00 1,21 34,s2 1,33 91,00 2,s4 ilo,43 1,93 70,43 1,25 89,39 8,68 85,772 1,52 76,05 1,52 96,81 0,74 88,51 0,62 82,15 0,71 3s,30 1,79 91,12 5,57 1l3,69 0,51l 75,43 2,24 96,99 2,06 88,44 0,83 81,03 1,49 4 u~3 :1 , 44 99,57 4,97 1l4,29 1,57 77,57 2,87 table 2 m ,w ejtimates from heart girth measurements for different classes of impala immature mature females mature males heart heart heart girth mass girth mass girth mass 55 16,9 65 34,0 70 37,4 56 17,8 66 34,5 7 i 38,4 57 18,6 67 35,0 72 39,4 58 19,5 68 35,5 73 40,4 59 20,4 69 36,0 74 41,3 60 21,3 70 36,6 75 42,3 61 22, i 71 37, i 76 43,3 62 23,0 72 37,6 77 44,3 63 23,9 73 38, i 78 45 ,2 64 24,7 74 38,6 79 46,2 65 25,6 75 39, i 80 47,2 66 26,5 76 39,7 81 48,2 67 27,4 77 40,2 82 49,1 68 28,2 78 40,7 83 50, i 69 29 , i 79 41,2 84 51 , i 70 29,9 80 41,7 85 52,1 81 42,3 86 53 ,0 82 42,8 87 54,0 83 43,3 88 55 ,0 84 43,8 89 56,0 85 44,3 90 57,0 discussion measurements quoted by sachs (1967) show that impala from the serengeti region in tanzania are heavier and larger in all body measurements. talbot and mcculloch (1965) confirm this fact for east african animals. they derive a regression of mass = 3,09 heart girth -149 when using mass in pounds and measurements in centimeters. when our data is treated in this way we derive an equation mass = 3,01 heart girth -140,8 for all age classes and both sexes combined. these equations are amazingly similar considering the difference in size of the two races of impala. mcculloch and talbot (1965) studied the relationship between mass and body measurements of various african ungulates. after extensive statistical tests using various linear and non-linear models they confiml that the simple relationship between heart girth and mass is as accurate as any other. the growth curves follow the normal pattern for large ungulates, the break at 8 months most likely being caused by the fact that the animals are weaned at about this time which coincides with a low nutritional value of the vegetation. 86 the information in table 2 can be used to construct a mass tape for the impala. in an attempt to assess the accuracy of this information deviations from the mean and deviations from the estimated value were determined. these indicate an accuracy of 4 per cent for the mean and 6 per cent for the estimate. therefore, while individual estimates could deviate up to 10 per cent, the estimates of any sample should be as accurately determined as with a balance. as mass determination by means of a tape is much easier and more practical in the field, it is felt that in most studies data from more animals could be obtained in this way with an eventual accuracy equal to that obtained by the conventional method. the estimates were applied to data from wankie national park in rhodesia and from the serengeti supplied by dr. jarman and worked well on our original data. this, taken together with the close relationship between our regression and that of talbot and mcculloch (1965) seems to indicate that this information may have a universal applicability. acknowledgements this study was done while the senior author was employed by the national parks board of trustees and the facilities provided are gratefully acknowledged. thanks are due to persons who helped with the collection of data, especially district ranger thys mostert. references child, g. 1964. growth and ageing criteria of impala aepyceros melampus. amoldia 27: 128-135. fairall, n. 1968. the reproductive seasons of large mammals in the kruger national park. zool. afr. 3: 189-2lo. fairall, n. 1969. prenatal development of the impala aepycer05 melampus. koedoe 12:97-lo3 . f ai rall, n. 1972. die voortplantingsfisiologie van die rooibok aepyceros melampus. d.se. thesis, university of pretoria, pretoria. mcculloch, j. s. g. and l. m. talbot. 1965. comparison of weight estimation methods for wild animals and domestic livestock.}. appl. ecol. 2, 59-69. roettcher, d., r. r. hoffmann and f. i. b. kayanja. 1970. ergebnisse der prae und postnatalen alterbestimmung beim ostafrikanishen impala. z. saugetierk. 35 :289-305. talbot, l. m. and j. s. g. mcculloch. 1965. weight estimates for east african mammals from body measurements. j .. wildl. mgmt. 29 :84-89. sachs, r. 1967. liveweights and body measurements of serengeti game animals. e. afr. wildl.}. 5 :24-36. 87 page 1 page 2 page 3 page 4 page 5 article information authors: anna s. dippenaar-schoeman1,2 michelle hamer3,4 charles r. haddad5 affiliations: 1arc-plant protection research institute, queenswood, south africa 2department of entomology and zoology, university of pretoria, south africa 3south african biosystematics institute, pretoria, south africa 4school of biological and conservation science, university of kwazulu-natal, south africa 5department of zoology and entomology, university of the free state, south africa correspondence to: charles haddad email: haddadcr@ufs.ac.za postal address: p.o. box 339, bloemfontein 9300, south africa dates: received: 27 may 2011 accepted: 07 sept. 2011 published: 11 nov. 2011 how to cite this article: dippenaar-schoeman, a.s., hamer, m. & haddad, c.r., 2011, ‘spiders (arachnida: araneae) of the vegetation layer of the mkambati nature reserve, eastern cape, south africa’, koedoe 53(1), art. #1058, 10 pages. http://dx.doi.org/10.4102/koedoe.v53i1.1058 copyright notice: © 2011. the authors. licensee: aosis openjournals. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) spiders (arachnida: araneae) of the vegetation layer of the mkambati nature reserve, eastern cape, south africa in this checklist... open access • abstract • introduction • materials and methods    • study area and period    • sampling methods    • grassland sampling    • forest sampling • results    • species inventory and faunal composition    • species prevalence, rarity, endemism and habitats • discussion • conclusion • acknowledgements    • author’s contributions • references • appendix abstract (back to top) the pondoland region of the eastern cape province, south africa is very poorly studied with regard to invertebrate diversity, particularly in the case of arachnids. accordingly, and in view of proposed infrastructural and mining developments in this ecologically sensitive area of high plant endemism, baseline data are provided on spiders (araneae) of the vegetation layer (i.e. excluding the ground-dwelling fauna) of the mkambati nature reserve (mnr). spiders were collected at 26 sites (six forest and 20 grassland sites) in the mnr over an eight-day period, using sweep sampling and active searching of flowers in grassland and tree beating in forests, as part of a broader biodiversity survey. additional specimens were collected with malaise and pan traps. a total of 1275 specimens were sampled, representing 132 species (6.6% of the total number recorded in south africa) in 103 genera and 29 families. theridiidae and araneidae were the most diverse spider families in the reserve, represented by 22 species each (16.7% of the total), followed by thomisidae with 19 species (14.4%) and salticidae with 18 species (13.6%). grassland and forest had distinct spider faunas, with only 24.2% of species being recorded from both biomes. the average number of species sampled per site in grassland and forest was 26 species for both habitats, although values for the two biomes are not directly comparable because different sampling methods were used. all 132 species are new records for the reserve, of which 20 were new records for the eastern cape and at least eight spider species may be new to science. the spider diversity captured despite temporal and methodological limits indicates that many additional species are likely to occur in the reserve. conservation implications: if the mnr is not adequately conserved at least five new species, which may be confined to the area, would be at high risk of extinction and 15 other species endemic to the pondoland and kwazulu-natal region would have their risk of extinction increased. introduction (back to top) the conservation of biodiversity is critical to the well-being of humans owing to its provision of ecosystem services (millennium ecosystem assessment 2005). in addition, biodiversity, especially where unique and rich, can also contribute to economic development through ecotourism (kepe 2001). however, to identify key areas for conservation and ecotourism and to monitor the effects of protection, management and/or threats, biodiversity inventories are essential and are considered by conservationists as being good investments (balmford & gaston 1999; rohr, mahan & kim 2007; ward & larivière 2004). invertebrates have become recognised as an important component of biodiversity. they are important in all ecosystems with regard to species richness (an estimated 95% of all species are invertebrates), abundance and biomass, and they play vital roles in ecosystem functioning (see, for example, black, shepard & allen 2001; janzen 1987; luck, daily & ehrlich 2003). spiders (araneae) are one of the largest orders of terrestrial invertebrates, with more than 40 000 described species (platnick 2010), and are predacious in all of their life stages. they therefore play an important role in terrestrial food webs as arthropod population regulators, whilst also providing food for other predators (gruner 2004; schmitz 2003). the south african national survey of arachnida was initiated in 1997 to document the diversity and distribution of arachnids in the country (dippenaar-schoeman & craemer 2000). surveys have been carried out across many of the provinces, biomes, agro-ecosystems and protected areas of south africa (dippenaar et al. 2008; dippenaar-schoeman 2006; dippenaar-schoeman & leroy 2003; dippenaar-schoeman et al. 1999, 2005; dippenaar-schoeman, van den berg & prendini 2009; foord et al. 2008; haddad & dippenaar-schoeman 2009; haddad, dippenaar-schoeman & wesolowska 2006), thereby generating the data that are critical for a conservation assessment of species and for understanding habitats and vegetation types of conservation concern for arachnids. whilst the pondoland region of the eastern cape, south africa is recognised as a centre of high plant diversity and endemism (van wyk & smith 2001), the spider fauna of the region is still poorly known and no structured surveys or sampling using standard methods and a consistent number of samples across sites (i.e. quantified surveys) have previously been carried out in the region. the environmental impact assessment report for a proposed national toll road through the region acknowledged the poor state of knowledge regarding invertebrates of the area (branch 2002). quantified surveys are critical for comparisons of areas and for providing future baseline data for monitoring (rohr et al. 2007). the mkambati nature reserve (mnr) was selected for the survey of invertebrates in 2008 because of possible tourism development in the reserve (kepe 2001), the proposed construction of a toll road adjacent to the reserve (branch 2002; farrington & davies 2004), the threat of controversial open cast dune mining in the region and the lack of invertebrate data to support planning and decision-making. the survey targeted a range of invertebrates, including spiders. the objectives of this particular paper are (1) to provide an annotated species list of spiders sampled from the vegetation layer at the mnr, (2) to provide an analysis of the spider faunal composition with regard to taxonomic breakdown and guilds, and (3) to assess the fauna with regard to species abundance or rarity, levels of endemism and the habitats used by species. spider guilds can be broadly divided into ground dwellers, plant dwellers and web dwellers. sampling in the mnr focused mainly on two of these guilds, namely plant dwellers and web dwellers. it should be noted that this survey is still incomplete because few ground dwellers, usually sampled with pitfall traps and litter sampling methods, were collected. community and detailed habitat analyses will be presented as part of a broader invertebrate publication (hamer et al. in prep.). materials and methods (back to top) study area and period the mnr (31°15’s, 29°56’e) is situated in north-eastern pondoland, 30 km south of port edward in the eastern cape, south africa (figure 1). this coastal reserve is 7720 ha in size, and comprises mainly open grasslands with patches of coastal, dune and swamp forest (figure 2). the climate is subtropical, with summer rainfall (annual average = 1200 mm). figure 1: location and extent of the mkambati nature reserve, south africa and its main river systems. figure 2: habitats sampled at the mkambati nature reserve: a) site 1 (grassland), b) site 11 (forest), c) site 19 (grassland), d) site 12 (forest), e) site 15 (forest), f) site 10 (grassland), g) site 25 (grassland) and h) site 26 (forest). we surveyed the mnr between 24 january and 3 february 2008 and sampled a total of 26 sites, which included six forest (coastal forest, swamp forest and dune forest) and 20 grassland sites. the sample sites covered localities from the coast at 16 m.a.s.l., those near wetlands, on rocky hills or slopes, and inland sites at 311 m.a.s.l. (figure 2; table 1). sampling was undertaken primarily by seven earthwatch institute volunteers, who had no previous biodiversity survey experience, and seven experienced technicians or scientists under the supervision of the second author. the volunteers attended an introductory presentation on the project and sampling methods and were then trained on site. they received close guidance from the experienced project staff during the actual sampling. previous studies have shown that volunteers are capable of producing similar diversity results when compared with experienced researchers (lovell et al. 2009) and their contribution to the current biodiversity survey was thus considered adequate to get a representation of the spider fauna of the area. considering the size of the survey team, the sampling effort was 112 ‘person-days’. table 1: details of sites and sampling methods used during the survey at mkambati nature reserve (january–february 2008). sampling methods details of sampling methods used at each site are provided in table 1. grassland sampling grassland sampling included setting one malaise trap and five each of yellow, blue and white pan traps, which were collected after five days, at each site (approximately 1 ha). six sweep samples, each comprising 40 sweeps (approximately 1 m apart) with a 50-cm diameter net, were taken from selected sites. active searching for spiders on 40 flowers was carried out at 10 of the grassland sites. this sampling involved examination of the upper and lower surfaces of each flower and the capture of any spiders observed using a vial. there was no time limit set for examination of flowers. forest sampling ten trees in each forest site were beaten to sample selected invertebrates, including spiders. for this method each tree was struck 10 times with a beating stick and a white tray (80 cm x 80 cm) was held underneath to collect specimens that dropped from the tree. an aspirator was used to capture small specimens. specimens were preserved in 80% ethanol and later sorted in the laboratory according to sample and morphospecies. the first and third authors subsequently identified the specimens to species level, where possible. voucher specimens are housed at the national collection of arachnida of the arc–plant protection research institute in pretoria, south africa. results (back to top) species inventory and faunal composition a total of 1275 specimens were sampled, representing 132 species (6.6% of the total recorded in south africa) in 103 genera (95 identified) and 29 families (appendix and table 2). theridiidae and araneidae were the most diverse spider families sampled in the reserve, represented by 22 species each (16.7% of the total number of species), followed by thomisidae (19 species; 14.4%) and salticidae (18 species; 13.6%). neoscona blondeli (simon, 1885) (araneidae) represented 13.0% of the spider specimens sampled, followed by heliophanus proszynskii wesołowska, 2003 (salticidae) at 7.8% (table 2). of the sampled species, 38 (28.8%) could not be identified beyond genus level and a further eight species (6.1%) could not be assigned to a genus. an equal number (66) of web-dwelling and wandering species were sampled (appendix). table 2: spider families sampled at the mkambati nature reserve and diversity within each family, in order of decreasing species richness. the highest number of individuals (n = 635, 49.8%) was sampled using sweep-netting, followed by tree beating, with which 342 individuals (26.8%) were sampled. pantraps sampled 206 individuals (16.2%) and other methods provided 92 specimens (7.2%). species prevalence, rarity, endemism and habitats of the total number of species sampled, 35 (26.5%) occurred only in forest and 65 (49.2%) were confined to grassland. only 32 species (24.2%) occurred in both habitat types. at one site 45 species (34.1%) were recorded, whilst 40 species (30.3%) were represented by only a single individual (singletons). no species was sampled at all the sites and only seven species (5%) were recorded from more than 50% of the sampled sites (both forest and grassland). the most regularly collected species across sites were n. blondeli and copa flavoplumosa simon, 1885 (corinnidae), which were sampled at 20 sites, theridion sp. 1 (theridiidae) and clubiona sp. 3 (clubionidae), which were collected from 17 sites, and h. proszynskii, which was collected at 16 sites. these were also the most abundant species. all 132 species are new records for the reserve and eight species (6.1% of the total) may be new to science. these species belong to the genera chresiona simon, 1903 (amaurobiidae), araniella chamberlin & ivie, 1942, cyrtophora simon, 1864, hypsosinga ausserer, 1871 and poltys c.l. koch, 1843 (araneidae), typhistes simon, 1894 (linyphiidae), cheiramiona lotz & dippenaar-schoeman, 1999 (miturgidae) and tibitanus simon, 1907 (philodromidae). five of these species may be confined to the reserve or the pondoland region as they have not been collected previously in surveys of other areas. a further 15 species are known only from the eastern cape and one other province, and another 11 species are south african endemics (appendix). the number of species recorded at a grassland site, where the full suite of sampling methods was used, ranged between 15 and 43, with an average number of 26 species per site. the range for forests was 17–45 species, also with an average of 26 species per forest. site 17, a flat, open inland area with frequently burnt grass, had the highest measured richness for grassland. site f26, a large forest in an inland gorge (known as the superbowl), had the highest measured forest richness (table 1). discussion (back to top) it should be noted that this survey produced a preliminary species list and that additional methods, sites and survey timing will produce many more species. other spider surveys have shown that species numbers increased with increased sampling effort (e.g. cardoso et al. 2008, 2009; muelelwa et al. 2010). however, given the large efforts and costs required to sample, sort and identify invertebrates such as spiders, completing the inventory is unlikely to occur in the short term. given the urgency for biodiversity data, especially with regard to invertebrates, even incomplete data sets have relevance. the identification of at least eight possibly new species over eight days of sampling illustrates just how poorly known the fauna of pondoland is. the 30% of singletons sampled at the mnr is close to the average of 32% calculated by coddington et al. (2009) from 71 studies. the authors suggested that very high percentages of singletons indicate undersampling, but they also recognised that undersampling is virtually inevitable in most tropical regions when dealing with arthropods. many other published studies on spider diversity included specimen numbers similar to the total sampled at the mnr, ranging between 75 and 9375 individuals (see coddington et al. 2009). this suggests that the data collected in the current study are not unacceptably incomplete. almost half the species recorded could not be identified beyond genus or even family level, which highlights a common problem with invertebrate surveys. in a survey of the polokwane nature reserve by dippenaar et al. (2008), a similar proportion (35%) of unidentifiable spider species was found. expertise for identification is usually limited to selected families and global experts may need to be consulted, which creates delays in obtaining data; in many cases even global expertise does not exist. the large number of unidentifiable specimens is also an indication of the incomplete knowledge of the fauna and the figures include the eight species recognised as new. it is possible that additional new species may be found amongst the material not identified to species level. some of the species could not be identified because only immature specimens were collected, which is another common problem associated with invertebrate surveys. comparisons of recorded species richness across spider surveys of other biomes and areas are difficult because in some of the similar spider surveys sampling was not quantified and sampling effort may be unequal. however, haddad and dippenaar-schoeman (2009) provided a range of between 76 and 431 species for south african conserved areas for which published surveys exist. none of the reserves they provided data for were in the forest or grassland biome. for the polokwane nature reserve, 13 821 specimens sampled over a year yielded 275 species (or a sample intensity of 50 specimens per species) (dippenaar et al. 2008). in a heterogeneous area in india, hore and uniyal (2008) sampled 3666 individuals over a year, representing 160 species (a sample intensity of 23 specimens per species). at the mnr the sample intensity was approximately nine specimens per species. in spider surveys the ratio of individuals to species is initially low and increases as an increasing number of samples are taken and fewer new morphospecies are sampled (e.g. cardoso et al. 2008). this indicates that the mnr survey (which provided a low specimens-to-species ratio) is only in the initial phase of discovery of biodiversity and supports the suggestion that much more sampling is needed to approach a complete inventory (high ratio value). coddington et al. (2009) calculated sample intensities of less than 10 for some individual sampling methods, where they estimated the survey completeness to be around 60%. the low sample intensity at the mnr could also indicate higher diversity than for areas where large numbers of specimens are required to record additional species. forests have been shown in many studies to have high species richness across many taxa, including spiders. this is often attributed to the higher structural complexity in forests (hore & uniyal 2008). however, in india, hore and uniyal (2008) found that grassland also had high species richness and a diverse assemblage of spiders relative to forests. a similar pattern was evident at the mnr, with grassland sites showing high species richness. this would appear not to conform to the general trend of diversity being associated with obvious habitat heterogeneity (yen 2009). of course, the sampling methods used in the two biomes may not be directly comparable with regard to effort, but this aspect of spider diversity should be investigated further. hore and uniyal (2008) found that annual burning of grassland did not seem to affect spider diversity negatively. this was also seen at the mnr, where two of the frequently burnt grassland sites were amongst the 10 sites of highest species density. this could be explained by higher herbivore density in newly resprouted grass and thus higher prey density, but the diversity of spider species rather than simple abundance is interesting. however, this study did not specifically investigate the effects of burning on spider diversity and conclusions cannot be drawn for the small number of sites sampled. the site with the highest species density was the largest forest site (f26, the ‘superbowl forest’). this might be explained by higher sampling effort exercised than for the other forests (tree beating was performed in 20 rather than 10 trees), but even so, this forest does house at least one new species that may be a narrow endemic, seven species that are near endemics for the eastern cape, and 26 species that were not identified to species level. additional sampling will be required to validate this preliminary assessment of the sites, but the superbowl forest appears to be a high priority for the spider fauna. the survey suggests that additional sampling of the spider fauna would be productive, especially in view of the threats to the region. other aspects that require investigation are the comparison of the grassland and forest habitats with regard to their invertebrate fauna using comparative sampling methods and efforts, and further investigation of fire impacts in the grassland areas. conclusion (back to top) baseline data on spiders are provided for the first time for a conserved area in pondoland, eastern cape. a total of 132 species, including at least eight new species, were recorded from the mnr. grassland and forest sites had a similar average number of species and each had a distinct fauna, indicating that both biomes require conservation measures. a single site yielded 34% of the total number of species recorded, whilst only seven species (5%) were sampled from more than 50% of the sites. although the survey was not comprehensive, preliminary data suggest a high turnover of species even at the small scale at which the survey was carried out, which means that a large area is likely to be required to conserve the spider fauna. this finding should be considered in setting conservation targets for pondoland grassland and forest. in addition, any loss of area in the mnr may result in the loss of species in the reserve because most areas have unique species. of the eight new species recorded, five may be confined to the mnr or the pondoland area. this means that these species would be at high risk of extinction without the conservation area, and 15 other species endemic to the pondoland and kwazulu-natal region would have their risk of extinction increased if the mnr is not adequately conserved. the results from this study provide a platform for much needed future spider research in pondoland, eastern cape, and the species data can be used in future conservation and environmental impact assessments. acknowledgements (back to top) we thank the earthwatch institute for sponsoring the project and for field assistance by the volunteers. eastern cape nature conservation, through dr dave balfour and mr jan venter, also contributed to funding for the project and provided logistical support during fieldwork. funding was also provided by the national research foundation through a grant to m.h. various field and laboratory assistants (jessica cochrane, lynne stone and patsy birkett, in particular) and collaborator rob slotow of the university of kwazulu-natal are thanked for their contributions to the project. the university of kwazulu-natal is thanked for providing access to a vehicle and laboratory facilities. two anonymous reviewers are thanked for useful suggestions to improve the manuscript. author’s contributions a.s.d. identified most of the material and presented statistical data. m.h. was the project leader, supervised the fieldwork and sorting of material, drafted the manuscript and took habitat photographs. c.h. identified corinnidae and salticidae, designed the map, and prepared the manuscript for submission. a.s.d., m.h. and c.h. all contributed to writing the manuscript. references (back to top) balmford, a. & gaston, k.j., 1999, ‘why biodiversity surveys are good value’, nature 398, 204–205. http://dx.doi.org/10.1038/18339black, s.h., shepard, m. & allen, m.m., 2001, ‘endangered invertebrates: the case for greater attention to invertebrate conservation’, endangered species update 18, 42–50. branch, w.r., 2002, ‘terrestrial fauna’, in n2 wild coast toll road between durban and east london, environmental impact assessment report, cca environmental (pty) ltd, cape town. cardoso, p., crespo, l.c., carvalho, r., rufino, a.c. & henriques, s.s., 2009, ‘ad-hoc vs. standardized and optimized arthropod diversity sampling’, diversity 1, 36–51. http://dx.doi.org/10.3390/d1010036 cardoso, p., scharff, n., gaspar, c., henriques, s.s., carvalho, r., castro, p.h. et al., 2008, ‘rapid biodiversity assessment of spiders (araneae) using semi-quantitative sampling: a case study in a mediterranean forest’, insect conservation and diversity 1, 71–84. coddington, j.a., agnarsson, i., miller, j.a., kuntner, m. & hormiga, g., 2009, ‘undersampling bias: the null hypothesis for singleton species in tropical arthropod surveys’, journal of animal ecology 78, 573–584. http://dx.doi.org/10.1111/j.1365-2656.2009.01525.x, pmid:19245379 dippenaar, s.m., modiba, m.a., khoza, t.t. & dippenaar-schoeman, a.s., 2008, ‘a checklist of the spiders (arachnida, araneae) of the polokwane nature reserve, limpopo province, south africa’, koedoe 50, 10–17. dippenaar-schoeman, a.s., 2006, ‘new records of 43 spider species from the mountain zebra national park, south africa (arachnida: araneae)’, koedoe 49, 23–28. dippenaar-schoeman, a.s. & craemer, c., 2000, ‘the south african national survey of arachnida’, plant protection news 56, 11–12. dippenaar-schoeman, a.s. & leroy, a., 2003, ‘a check list of the spiders of the kruger national park, south africa (arachnida: araneae)’, koedoe 46, 91–100. dippenaar-schoeman, a.s., leroy, a., de jager, m. & van den berg, a., 1999, ‘a check list of the spider fauna of the karoo national park, south africa (arachnida: araneae)’, koedoe 42, 31–42. dippenaar-schoeman, a.s., van den berg, a. & prendini, l., 2009, ‘spiders and scorpions (arachnida: araneae, scorpiones) of the nylsvley nature reserve, south africa’, koedoe 51, 1–9. http://dx.doi.org/10.4102/koedoe.v51i1.161 dippenaar-schoeman, a.s., van der walt, a.e., de jager, m., le roux, e. & van den berg, a., 2005, ‘the spiders of the swartberg nature reserve in south africa (arachnida: araneae)’, koedoe 48, 77–86. farrington, k. & davies, k., 2004, ‘‘the gentle art of letting the other fellow have your own way’: viewpoints on a media narrative used to promote the proposed n2 toll road’, southern african journal of environmental education 21, 166–173. foord, s.h., mafadza, m.m., dippenaar-schoeman, a.s. & van rensburg, b.j., 2008, ‘micro-scale heterogeneity of spiders (arachnida: araneae) in the soutpansberg, south africa: a comparative survey and inventory in representative habitats’, african zoology 43, 156–174. http://dx.doi.org/10.3377/1562-7020-43.2.156 gruner, d.s., 2004, ‘attenuation of top-down and bottom-up forces in a complex terrestrial community’, ecology 85, 3010–3022. http://dx.doi.org/10.1890/04-0020 haddad, c.r. & dippenaar-schoeman, a.s., 2009, ‘a checklist of the non-acarine arachnids (chelicerata: arachnida) of the de hoop nature reserve, western cape province, south africa’, koedoe 51, 1–9. http://dx.doi.org/10.4102/koedoe.v51i1.149 haddad, c.r., dippenaar-schoeman, a.s. & wesolowska, w., 2006, ‘a checklist of the non-acarine arachnids (chelicerata: arachnida) of the ndumo game reserve, maputaland, south africa’, koedoe 49, 1–22. hore, u. & uniyal, v.p., 2008, ‘diversity and composition of spider assemblages in five vegetation types of the terai conservation area, india’, journal of arachnology 36, 251–258. http://dx.doi.org/10.1636/ct07-53.1 janzen, d.h., 1987, ‘insect diversity of a costa rican dry forest: why keep it, and how?’, biological journal of the linnean society 30, 343–356. http://dx.doi.org/10.1111/j.1095-8312.1987.tb00307.x kepe, t., 2001, ‘tourism, protected areas and development in south africa: views of visitors to mkambati nature reserve’, south african journal of wildlife research 31, 155–159. lovell, s., hamer, m., slotow, r. & herbert, d., 2009, ‘an assessment of the use of volunteers for terrestrial invertebrate biodiversity surveys’, biodiversity and conservation 18, 3295–3307. http://dx.doi.org/10.1007/s10531-009-9642-2 luck, g.w., daily, g.c. & ehrlich, p.r., 2003, ‘population diversity and ecosystem services’, trends in ecology and evolution 18, 331–336. http://dx.doi.org/10.1016/s0169-5347(03)00100-9 millennium ecosystem assessment, 2005, ecosystems and human well-being: biodiversity synthesis, world resources institute, washington dc. muelelwa, m.i., foord, s.h., dippenaar-schoeman, a.s. & stam, e.m., 2010, ‘towards a standardized and optimized protocol for rapid assessments: spider species richness and assemblage composition in two savanna vegetation types’, african zoology 45(2), 273–290. http://dx.doi.org/10.3377/004.045.0206 platnick, n.i., 2010, the world spider catalog, version 10.5, viewed 08 june 2010, from http://research.amnh.org/entomology/spiders/catalog/index.html (american museum of natural history). rohr, j.r., mahan, c. & kim, k., 2007, ‘developing a monitoring program for invertebrates: guidelines and a case study’, conservation biology 21, 422–433. http://dx.doi.org/10.1111/j.1523-1739.2006.00578.x, pmid:17391192 schmitz, o.j., 2003, ‘top predator control of plant biodiversity and productivity in an old-field ecosystem’, ecology letters 6, 156–163. http://dx.doi.org/10.1046/j.1461-0248.2003.00412.x van wyk, a.e. & smith, g., 2001, regions of floristic endemism in southern africa, umdaus press, pretoria. ward, d.f. & larivière, m.c., 2004, ‘terrestrial invertebrate surveys and rapid biodiversity assessment in new zealand: lessons from australia’, new zealand journal of ecology 28, 151–159. yen, j.d.l., 2009, ‘spider family richness and habitat complexity’, victorian naturalist 126, 18–22. appendix (back to top) appendix: list of spider species sampled at the mkambati nature reserve (january–february 2008). filelist convert a pdf file! koedoe 19: 189-191 (1976) lewensloop van 'n kaapse buffelbevolking syncerus caffer in die bontebok nasionale park p. t. van der walt ajdeling natuurbewaring privaatsak x404 skukuw ljso l. j. van zyl bontebok nasionale park posbus 149 swellendam 6740 g de graaff nasionale parkeraad posbus 787 pretoria 0001 abstract the course of an experiment to re-introduce the cape buffalo syncerus caifer in the bontebok national park, near swellendam, is described and illustrated. inleiding die historiese verspreiding van die kaapse buffel syncerus caifer in suidelike afrika soos weergegee deur du plessis (1969), dui daarop dat die dier voorkeur gee aan grasveldgebiede waarbinne bosse en borne volop voorkom. klimaatstoestande tree skynbaar nie beperkend op in die buffel se verspreiding nie aangesien hy dwarsoor die kontinent vanaf die dorre westelike klimaatsones tot by die subtropiese oostelike kusstroke voorgekom het. langs die suidoos kaapse kusgebied word daar vandag slegs 'n betekenisvolle buffelpopulasie, ongeveer 250, aangetref in die addo-olifant nasionale park (aonp) naby port elizabeth. in 'n poging om die voortbestaan van die buffel in hierdie dele verder te stabiliseer (liebenberg 1963) het die nasionale parkeraad gedurende 1963 ses buffels hervestig in die bontebok nasionale park (bnp), naby swellendam, vanuit die aonp. vele faktore het egter daartoe gelei dat die populasie oor 'n 12 jaar periode nie juis kon toeneem in getalle nie. 189 jaar periode nie juis kon toeneem in getalle nie. hierdie verslag skets kortliks die lewensgang van die buffelbevolking in die bnp tot met die verwydering daarvan uit die bnp gedurende september 1974. rejuitate en bespreking die verloop van 'n eksperiment om 'n diersoort met 'n temperamentele geaardheid te hervestig in 'n bewaringsgebied van slegs 2786 ha in omvang en wat gelee is midde in 'n intensief bewerkte graanstreek, word saamgevat in fig. i . ...j ;! z oc[ oc[ • vrektes 18 ~ manlik 16 ii vroulik 14 o kalwers +3 getal i ngevoer vanuit addo. 12 10 a 6 .. 2 63 64 65 66 67 68 71 72 jare fig. i. lewenshistogram van die buftelbevolking in die bontebok nasionale park vanaf 1963-1974. daar was beide 'n bewaringsen 'n bestuursmotief agter die plasing van die bufteis in die bnp. liebenberg (1963) meld dat die diere se vreetgewoontes hopelik sou bydra om die vermeerdering van onsmaaklike grasse, so os die verskillende pentaschistis spp., stop te sit. sodoende is gehoop om die gebruik van brand uit te skakel. du plessis (j 969) meld dat die bnp op een na die mees suidelike gebied is waar builds voorgekom het soos blyk vanuit bekende verspreidingsrekords. reeds kort na die plasing van ses jong buffels in die bnp in 1963, het 'n mate van onrustigheid onder die diere na yore getree. dit dwing liebenberg (1963) om aan te beveel dat die tropgrootte aangevul moet word na 10 jong diere 30'0'en 799, om sodoende 'n "rustige teeltrop" te verseker. slegs een bul en twee koeie is daarop toegevoeg. gedurende 1965 is die bevolking aangevul met 'n verdere vyf individue. 190 eers vroeg in 1966 het die eerste nakomelinge hulle verskyning gemaak, met 'n eenmalige 100% kalfpersentasie. sedertdien het 'n relatief lae kalfpersentasie oor die jare voorgekom, naamlik 19%. dit was skynbaar nie te wyte aan 'n proteientekort in die diere se dieet nie aangesien hul fisiese kondisie deurgaans daarop gedui het dat omgewingsfaktore soos die heersende klimaat en plantegroei bevorderlik is vir die trop se welsyn. die geslagsverhoudings was ook altyd taamlik gunstig (fig. 1), alhoewel dit lyk of die effense getalle-oorwig van die bulle steurings in die paringsaktiwiteite kon teweeggebring het. die groot aantal vrektes wat daar oor die jare onder die buffels voorgekom het, kan grotendeels gewyt word aan moontlike gebrek aan lebensraum vir die trop waartydens van die kalwers deur die ouer diere gedood is. verder het opportunitiese verliese voorgekom tydens hul sporadiese uitbreke uit die bnp sedert 1967. die kwaai daling in hul getalle gedurende 1971 is tydens een van die laasgenoemde uitbreke aangeteken, aangesien daar tydens die proses van terugplasing van die diere in die park heelwat verliese voorgekom het. sedert 1967 het die buffels feitlik jaarliks daarin geslaag om buite die bnp te kom. dit het hulle vermag om bloot deur die omheining te breek in hul haas om by die aanloklike naburige koringlande te kom. hulle het verder tydens die tye wat die breerivier in vloed was, oor die versperring geswem wat deel van die bnp se grens vorm in die suidwestelike hoek. die eksperiment om die kaapse buffel te laat bly voortbestaan in die suid-kaap is dus teen groot koste voortgesit vir 12 jaar. op 1974.06.28 besluit die nasionale parkeraad dat hierdie bewaringspoging vrugteloos was omdat die sukses daarvan onderhewig was aan eksterne faktore soos veral parkgrootte, standaard van omheining en omringende boerederyaktiwiteite . die aard van die buffel was dus beperkend op sy voortbestaan alhier. verwysing du plessis, s. f. 1969. the past and present geographical distribution of the perissodoctyla and artiodactyla in southern africa. m.sc. thesis, univ. of pretoria. liebenberg, l. c. c. 1963. a memorandum on the place of the buffalo in the bontebok park. unpublished. national parks board meeting, june. 191 page 1 page 2 page 3 article information authors: stoffel p. bester1,2 ronell r. klopper3,4 hester m. steyn1 hugo bezuidenhout5,6 affiliations: 1national herbarium, south african national biodiversity institute, south africa2school of environmental sciences and development, north-west university, south africa 3biosystematics research & biodiversity collections division, south african national biodiversity institute, south africa 4department of plant science, university of pretoria, south africa 5scientific services, south african national parks, south africa 6applied behavioural ecology and ecosystem research unit, university of south africa, south africa correspondence to: hester steyn postal address: private bag x101, pretoria 0001, south africa dates: received: 13 oct. 2011 accepted: 07 aug. 2012 published: 12 nov. 2012 how to cite this article: bester, s.p., klopper, r.r., steyn, h.m. & bezuidenhout, h., 2012, ‘new plant records for tankwa karoo national park, south africa’, koedoe 54(1), art. #1066, 9 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1066 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. new plant records for tankwa karoo national park, south africa in this checklist... open access • abstract • introduction • research method and design • ethical considerations • results and discussion • conclusion • acknowledgements    • competing interests    • authors' contributions • references • appendix 1: detailed information and relevant specimens for new plant records from tankwa karoo national park • apocynaceae    • quaqua acutiloba (n.e.br.) bruyns    • quaqua parviflora (masson) bruyns subsp. gracilis (c.a.lückh.) bruyns    • stapelia arenosa c.a.lückh.    • stapelia surrecta n.e.br.    • tridentea parvipuncta (n.e.br.) l.c.leach subsp. truncata (c.a.lückh.) bruyns    • tromotriche thudichumii (pillans) l.c.leach • asphodelaceae: alooideae    • aloe microstigma salm-dyck    • aloe falcata baker    • aloe variegata l. • hyacinthaceae    • lachenalia ameliae w.f.barker • iridaceae    • moraea tanquana goldblatt & j.c.manning • scrophulariaceae    • nemesia suaveolens k.e.steiner • pteridophyta: sinopteridaceae    • cheilanthes parviloba (sw.) sw.    • cheilanthes induta kunze    • cheilanthes namaquensis (baker) schelpe & n.c.anthony abstract top ↑ the tankwa karoo national park has been enlarged from 27 064 ha to 143 600 ha. this whole area is severely under-collected for plants in general and therefore it was an obvious target for the south african national parks (sanparks) programme, a component of the pretoria national herbarium (pre) plant collecting programme. this programme not only aims to survey national parks that have been poorly surveyed, but also inadequately known taxa, unique habitats, remote and inaccessible areas and plant species flowering at irregular times, especially after events such as fire or unusual timing of, or high, rainfall. general collecting in the tankwa karoo national park has already led to the description of two new taxa, from two families. it furthermore resulted in new distribution records for the park and for the northern cape province. these are reported on here.conservation implications: although the tankwa karoo national park falls within the succulent karoo biome (a biodiversity hotspot of international importance), information on its plant diversity is insufficient because it is an under-collected area. results of this study will guide conservation and supply occurrence and distribution data required to compile management plans for the park. introduction top ↑ the tankwa karoo national park (tknp) is a relatively new addition to south africa’s treasure of national parks and was only declared a national park in 1986 (rubin 1998). the original park consisted of eight farms amounting to 27 064 ha, which comprised mainly long gentle slopes to low-lying plains and isolated hills (rubin 1998). by 2011 the park was extended to 143 600 ha.the south africa national parks (sanparks) plant collection programme is a facet of the larger national herbarium (pre) plant collecting programme, where priority is given to collecting in national parks that contain under-collected areas (less than 200 species known within a quarter degree squared [qds] grid according to the south african national biodiversity institute [sanbi]’s plant database pre computerised information system [precis]). this programme officially started in 2005 with collecting efforts in the tankwa karoo and namaqua national parks. the work within these parks not only targets the under-collected areas, but also inadequately known taxa, unique habitats, remote and inaccessible areas and plant species flowering at irregular times, especially after events such as fire or unusually high rainfall. the aim of such surveys in under-collected areas, such as the tknp, is to enhance our knowledge and general understanding of these conservation areas in terms of their floristic component. general collecting in these areas will enable us to compile comprehensive species checklists for the areas that will, in turn, inform decisions regarding the conservation and monitoring of these conservation areas and the plants which they contain. research method and design top ↑ tankwa karoo national park is situated in the southern section of the succulent karoo biome (mucina & rutherford 2006) and stretches from the plains in the south-west to the escarpment of the roggeveld mountains in the east (figure 1). for details on the vegetation of the tankwa karoo and the tknp in particular the reader is referred to rubin (1998 [for major plant communities]), van der merwe, van rooyen and van rooyen (2008a [fynbos related vegetation]) and van der merwe, van rooyen and van rooyen (2008b [succulent karoo vegetation]). figure 1: map of tankwa karoo national park showing the original boundaries of the park at its inception in 1986, the expanded park as in 2005, and its current size (2011). all the qds grids in the park are regarded as under-collected, and therefore the whole park was targeted for general collecting. this entails collecting herbarium specimens of all plants that are flowering or fruiting at the various collecting sites that were chosen. collecting sites were selected to represent the variety of habitat types, for example hill slopes, plains and so forth, found within the specific qds. collecting trips targeted different areas of the park and were carried out at various times of the year (july–october), to reflect the floristic component of the park as best as possible.a total of seven collecting trips were conducted within the park in 2005–2010 with collecting efforts concentrated in the six qdss that comprised the park in 2005. the collecting was carried out by 10 collectors (all staff of the national herbarium in pretoria), including the first three authors. most trips included two or three collectors at a time, with one large trip where six collectors were involved. information accompanying each specimen includes the standard information recorded on herbarium collecting labels, such as locality, habitat, substrate, lithology, exposure, aspect and slope. global positioning system (gps) readings were taken, but these mostly represent a few general readings for a collecting site as a whole, and are not necessarily specific for each individual plant collected. fertile herbarium specimens were collected and identified mainly at the national herbarium in pretoria (pre). cuttings were taken from sterile stapeliads and these were grown at the pretoria national botanical garden’s nursery until flowering. flowers were then added to the herbarium sheets with sterile stems. specimens are housed at pre and duplicates, when available, were distributed to the kimberley south african national parks herbarium (ksan), the compton herbarium in cape town (nbg), and mcgregor museum herbarium in kimberley (kmg). herbarium acronyms follow holmgren, holmgren and barnett (1990). specimen information was incorporated into the main database, the pretoria national herbarium (pre) computerised information system (precis), to allow analysis of distribution data. new specimen records were compared to existing records in precis. where a first record for a taxon in tknp was indicated, the existing checklist for the area (rubin 1998) was consulted to verify the novelty of the record. all relevant specimens acting as vouchers of the new records reported on here, are cited under each taxon in appendix 1. the citation of these voucher specimens are listed in the following format: province.– grid latitude and longitude (grid name): locality description where the specimen was collected (–qds indicator), date, collector & collector’s number (acronym of herbaria where the specimen and its duplicates are housed) ethical considerations top ↑ ethical consideration has been given to the collection of plant specimens by not collecting indiscriminately and taking care not to damage plants. results and discussion top ↑ during recent field trips targeting the tankwa karoo region and especially the tknp, approximately 1500 specimens were collected by staff from the national herbarium (pre). this substantially increased our knowledge of the floristics of the region: at the start of the programme the lowest number of species per qds was 16 (3219ba); this number was raised to 108 (3219ba) by 2011 (table 1). table 1: number of species per quarter degree square grid cell according to the pretoria national herbarium computerised information system data. many significant collections were made as part of this programme: new distribution records were added to the pre collection, for example bulbine triebneri dinter (first record for the northern cape, klopper et al. 2008); first records for the park were collected, for example dioscorea elephantipes (l'hér.) engl. (collected on the gannaga pass); a new species (moraea tanquana goldblatt & j.c.manning) was described from leeuberg (goldblatt & manning 2009) and a new monotypic genus in the apiaceae (scaraboides magee & b.-e. van wyk) was described from elandsberg (magee et al. 2009).further significant range extensions and new park and provincial records from within the park are provided (table 2) with additional information on each species (appendix 1). table 2: family, taxon and status of new plant distribution records from tankwa karoo national park. in the checklist of species for the tknp (rubin 1998) only two stapeliads were listed, namely hoodia gordonii (masson) sweet ex decne. and a stapelia species. stapeliads, in general, are poorly represented in herbaria because it is usually difficult to locate these plants. many are obscured and hidden amongst nurturing plants that provide shade and shelter or they may grow in inaccessible habitats. furthermore, because of their collector’s value, many plants are collected and kept in living collections without making representative herbarium specimens. the correct preservation of herbarium specimens of stapeliads is also perceived to be difficult. cuttings collected in the park were grown until they flowered which made positive identification possible – the list was extended to include six more taxa (table 2).no aloes (asphodelaceae) or ferns (pteridophyta) were represented on the previous checklist (rubin 1998) and general collecting led to three taxa being added from each of these plant groups (table 2). although records of aloe microstigma salm-dyck from the calvinia area has been known since the 1920s, this species has not been collected in the tknp until 2006 when a few individuals of this aloe were found growing on the gannaga pass. apart from moraea tanquana (vulnerable) and nemesia suaveolens k.e.steiner (rare), the only other taxon with a red list status of conservation concern is lachenalia ameliae w.f.barker (near threatened) (l. von staden pers. comm., february 2010). following the new records and range extensions resulting from this study, the extent of occurrence could be re-determined and the statuses of these species were revised. all other taxa reported on here are listed in the latest red list (raimondo et al. 2009) as least concern, and these new records have no influence on their status. figure 2: images of selected species reported on: (a) quaqua acutiloba, (b) stapelia surrecta, (c) tridentea parvipuncta subsp. truncata, (d) tromotriche thudichumii, (e) aloe microstigma growing on the gannaga pass, (f) lachenalia ameliae growing on gravel plain, (g) moraea tanquana and (h) nemesia suaveolens. conclusion top ↑ the general collecting carried out thus far in the tknp has increased our knowledge of the flora of a severely under-collected and unique habitat significantly. knowledge of the full distribution of taxa, especially taxa with limited distribution ranges, is imperative in helping us fully understand these plant species; for instance, without this information the assessment of red data list status of such taxa cannot be carried out accurately. data generated by the sanparks programme in tknp will be used for better informed decisions regarding the conservation of this unique area and also will assist sanparks with management of these plant species. the data are essential in conservation and monitoring actions within the park and will be taken up in the park management plan and conservation development framework. furthermore, it is useful in terms of ecotourism to show visitors which species occur in the park and to highlight the importance of the park in the conservation of these plants. the occurrence of rare and threatened plants in the tknp will also focus the attention of researchers working on these groups on the park and will thus stimulate further research in the area. acknowledgements top ↑ we would like to thank our colleagues for helpful comments and suggestions on this manuscript. cape nature and northern cape nature conservation provided collecting permits, and lize von staden assessed the threat status of species. we also acknowledge with appreciation the two anonymous referees for suggesting improvements to the manuscript. furthermore, we would like to thank sanparks for permission to work in the arid parks with a special mention of conrad strauss, letsie coetzee and the staff of tankwa karoo national park. competing interests the authors declare that they have no financial or personal relationship(s) which may have inappropriately influenced them in writing this paper. authors' contributions s.p.b. (south african national biodiversity institute), r.r.k. (south african national biodiversity institute) and h.m.s. (south african national biodiversity institute) were equally responsible for data collection and analyses used in this study, as well as compiling the manuscript. h.b. (south african national parks) provided the team with logistical support for field work and made valuable contributing comments and improvements to the manuscript. references top ↑ brown, n.e., 1909, ‘stapelia surrecta’, in w.t. thiselton-dyer (eds.), flora capensis, vol. 4(1), pp. 970–971, reeve, london.bruyns, p.v., 2005, stapeliads of southern africa and madagascar, vol. ii, umdaus press, hatfield. crouch, n.r., klopper, r.r., burrows, j.e. & burrows, s.m., 2011, ferns of southern africa: a comprehensive guide, struik, cape town. goldblatt, p. & manning, j.m., 2009, ‘new species of moraea (iridaceae: iridoideae), with range extensions and miscellaneous notes for southern african species’, bothalia 39, 1–10. holmgren, p.k., holmgren, n.h. & barnett, l.c., 1990, index herbariorum, part 1: the herbaria of the world, 8th edn., new york botanical garden, new york. klopper, r.r., klopper, a.w., baijnath, h. & smith, g.f., 2008, ‘bulbine triebneri, an earlier name for bulbine alba, as well as additional and new localities in eastern and northern cape, south africa’, bothalia 38, 67–69. klopper, r.r. & smith, g.f., 2007, ‘the genus aloe (asphodelaceae: alooideae) in namaqualand, south africa’, haseltonia 13, 38–51. http://dx.doi.org/10.2985/1070-0048(2007)13[38:tgaaai]2.0.co;2 klopper, r.r. & smith, g.f., 2010, ‘the genus aloe l. (asphodelaceae: alooideae) in the eastern cape province of south africa’, haseltonia 16, 16–53. http://dx.doi.org/10.2985/1070-0048-16.1.16 leach, l.c., 1985, a revision of stapelia l. (asclepiadaceae), excelsa taxonomic series no. 3, aloe books, johannesburg. magee, a.r., van wyk, b.-e., tilney, p.m. & downie, s.r., 2009, ‘generic delimitations and relationships of the cape genera capnophyllum, dasispermum, and sonderina, the north african genera krubera and stoibrax, and a new monotypic genus of the subfamily apioideae (apiaceae)’, systematic botany 34(3), 580–594. http://dx.doi.org/10.1600/036364409789271218 mucina, l. & rutherford, m.c. (eds.), 2006, the vegetation of south africa, lesotho and swaziland, strelitzia 19, south african national biodiversity institute, pretoria. raimondo, d., von staden, l., foden, w., victor, j.e., helme, n.a., turner, r.c. et al. (eds.), 2009, red list of south african plants, strelitzia 25, south african national biodiversity institute, pretoria. rubin, f., 1998, ‘the physical environment and major plant communities of the tankwa karoo national park’, koedoe 41, 61–94. steiner, k., 2009, ‘two new species of nemesia (schrophulariaceae) from arid areas of the northern cape, south africa’, bothalia 39, 67–72. van der merwe, h., van rooyen, m.w. & van rooyen, n., 2008a, ‘vegetation of the hantam-tanqua-roggeveld subregion, south africa. part 1: fynbos biome related vegetation’, koedoe 50(1), 61–71. van der merwe, h., van rooyen, m.w. & van rooyen, n., 2008b, ‘vegetation of the hantam-tanqua-roggeveld subregion, south africa. part 2: succulent karoo biome related vegetation’, koedoe 50(1), 160–183. appendix 1: detailed information and relevant specimens for new plant records from tankwa karoo national park apocynaceae top ↑ quaqua acutiloba (n.e.br.) bruyns this species was discovered by robert templeman in 1898 and described by n.e. brown in 1909 (bruyns 2005). quaqua acutiloba has a relatively wide distribution range, occurring from south of laingsburg on the edge of the western escarpment northwards to the vicinity of aus in namibia (figure a-1). in the tknp this species is associated with rocky and gravelly plains and foot slopes of hills, but also grows on the lower plateaux (for instance the elandsberg plateau). these plants are usually associated with various species of ruschia which, in most cases, act as nurturing plants to this species (bruyns 2005). the stems can become ± 2 cm in diameter (usually 4-angled) and up to 300 mm long. the plants form clumps of up to 0.3 m in diameter. the flowers are about 1 cm in diameter and are extremely variable in the expression of colour that ranges from plain yellow-green to dark maroon with various degrees of maroon-spotted on yellow-green background forms. the flowers are open, rotate, sometimes with a shallow basal cup (figure 2a).relevant specimens: northern cape.–3219 (wuppertal): tknp. plato above elandsberg (–bb), 07 august 2007, s.p. bester 7792 (pre); on the road between potkleiberg and pramberg. south-western foot slopes and flats at potkleiberg. farm: folmoesfontein (–bd), 05 august 2007, s.p. bester 7731 (pre), s.p. bester 7732 (pre). quaqua parviflora (masson) bruyns subsp. gracilis (c.a.lückh.) bruyns the material from which lückhoff described this taxon was collected between calvinia and pakhuis pass (bruyns 2005). compared to the previous species the flowers in quaqua parviflora subsp. gracilis are much smaller and the stems generally much thinner. the flowers are characteristic with long narrow segments that are constricted more or less in the middle, ending in a thickened apical part. the apical section is usually homogenous in colour in contrast to the rest of the corolla that are maroon-spotted on a cream background. based mainly on this apical swelling and somewhat longer segments of the corolla, it is possible to distinguish it from the typical subspecies. this taxon has a much more restricted distribution than the typical species, occurring from the ceres karoo north-westwards to the west of garies (figure a-1). plants are usually confined to the lower foot slopes of mountainous areas along the escarpment in stony and rocky soils. figure a-1: distribution of quaqua acutiloba, quaqua parviflora subsp. gracilis and tridentea parvipuncta subsp. truncata. relevant specimens: northern cape.–3220 (sutherland): tknp. langkloof gorge (–aa), 07 august 2006, s.p. bester 7139 (pre).in the tknp quaqua and stapelia can be distinguished from each other in the sterile state based on the hairiness of the stems. in quaqua the stems are always glabrous compared to stapelia where it is velvety pubescent (there are, however, some exceptions in stapelia that are also glabrous, but these do not occur in tknp). stapelia arenosa c.a.lückh. joseph archer was the first to collect this species which came from the northern cederberg. it was later described by c.a. lückhoff (bruyns 2005). in the distribution range of this species it is encountered only sporadically. it has been recorded from the kamiesberg southwards to the karoo poort in the ceres karoo (leach 1985) (no specimens in the database). in august 2007 it was collected in the elandsberg area (figure a-2). the plants form few-flowered inflorescences with short pedicels causing the flowers to be held quite close to the stems. with repeated formation of flowers in inflorescences the stem may become irregularly thickened. as the stems develop, flowers may occur all along the stem, from its base to the tips. the stems are diffuse and form lax clumps. figure a-2: distribution of stapelia arenosa, stapelia surrecta and tromotriche thudichumii. relevant specimens: northern cape.–3219 (wuppertal): tknp. southern slopes of elandsberg, (–bb), 06 august 2007, s.p. bester 7749a (ksan, pre); plato above elandsberg, (–bb), 07 august 2007, s.p. bester 7795b (kmg, ksan, nbg, pre). stapelia surrecta n.e.br. this species was first collected by rudolph marloth and later described by n.e. brown in flora capensis (1909). stapelia surrecta (figure 2b) is one of few species in the genus that carries flowers with long pedicels concentrated at the tips of the stems (another example is some forms of stapelia kwebensis). this species has a restricted range and is only known from the ceres karoo. it occurs from karoo poort in the south to bloukrans pass in the north (figure a-2). in the tknp plants were collected in rocky and sandy soils in the langkloof, but according to bruyns (2005) it is restricted to shale and tillite. this species can only be distinguished from stapelia arenosa when flowering because the vegetative stems are indistinguishable.relevant specimens: northern cape.–3220 (sutherland): tknp. langkloof gorge (–aa), 07 august 2006, s.p. bester 7161a (ksan, pre); 7163a (pre). tridentea parvipuncta (n.e.br.) l.c.leach subsp. truncata (c.a.lückh.) bruyns the subspecies was originally discovered by james lückhoff in 1931 and described by c.a. lückhoff in 1937 (bruyns 2005). it is distinguished from the typical subspecies in that the outer corona lobes are truncate and emarginate and not deeply bifid with two lobes diverging as in the typical subspecies (figure 2c). tridentea parvipuncta subsp. truncata are found from the western section of karoo poort to botterkloof (figure a-1). it is separated from the typical subspecies which is found south of merweville westwards.relevant specimens: northern cape.–3219 (wuppertal): tknp. on the road between potkleiberg and pramberg. south-western foot slopes and flats at potkleiberg. farm: folmoesfontein (–bd), 07 november 2007, s.p. bester 7730a (kmg, ksan, pre). tromotriche thudichumii (pillans) l.c.leach although this taxon was already discovered in 1937 and re-collected in 1940 by marthinus malherbe, it was only described some 20 years later by n.s. pillans in 1959 (bruyns 2005). despite a manuscript name that was given to it in the bolus herbarium (stapelia malherbei), it was eventually named after jacques thudichum (curator of the karoo botanical garden at the time), who found a number of these plants in cultivation at the garden which he could not identify. the plants grow in rocky and gravelly soils on the flats of the tanqua karoo and roggeveld escarpment (bruyns 2005), mainly in the ceres karoo (north of karoo poort) and calvinia districts (figure a-2). characteristic of this species is the corolla that is so strongly reflexed that the flowers usually seem to be clasping the stem (figure 2d).relevant specimens: northern cape.–3219 (wuppertal): tknp. southern slopes of elandsberg (–bb), 06 august 2007, s.p. bester 7749 (kmg, ksan, pre); plato above elandsberg, (–bb), 07 august 2007, s.p. bester 7795a (kmg, ksan, pre); on the road between potkleiberg and pramberg. the south-western foot slopes and flats at potkleiberg. farm: folmoesfontein (–bd), 05 august 2007, s.p. bester 7730 (pre). asphodelaceae: alooideae top ↑ aloe microstigma salm-dyck aloe microstigma occurs fairly widespread in the western parts of the eastern cape, the central areas of the western cape and just into the northern cape, with a disjunction to south-western namibia (klopper & smith 2010). although records from the calvinia area has been known since the 1920s, this species has not been collected in the tknp until 2006 when a few individuals of this aloe were found growing on steep slopes in tankwa escarpment shrubland (mucina & rutherford 2006) on the gannaga pass. this population represents the yellow flowering form of a. microstigma, previously known as aloe brunthallerii a.berger ex cammerl. and is one of the northernmost records of this colour form (figure 2e and figure a-3). figure a-3: distribution of aloe microstigma and lachenalia ameliae. relevant specimens: northern cape.–3220 (sutherland): tknp, top of gannaga pass, farm kleinfontein 1027, near the viewpoint, (–aa), 05 august 2006, r.r. klopper 321 (ksan, pre); 10 july 2010, r.r. klopper & a.w. klopper 336 & 337 (pre). aloe falcata baker aloe falcata is known to occur from the richtersveld south to the klawer area and inland to around calvinia (klopper & smith 2007). it was first observed on a rocky ridge in the tanqua karoo vegetation type (mucina & rutherford 2006) during a general collecting trip in tknp in august 2006, but was not in flower and no material was collected. the significance of this population was only realised afterwards: it is the most south-eastern record known for this aloe to date. subsequently, during a specialised collecting trip focussing on aloes in 2010, a herbarium record from this population was collected (figure a-4). figure a-4: distribution of aloe falcata. relevant specimens: northern cape.–3220 (sutherland): tknp, farm roodewerf 1091, houtkraalkloof, (–aa), 10 july 2010, r.r. klopper & a.w. klopper 339 (pre). aloe variegata l. this very widespread and small aloe, with its cryptically mottled and camouflaged leaves (klopper & smith 2007, 2010), is very inconspicuous when not in flower. it is thus not surprising that it has not been collected in the tknp before 2006, even though it occurs at several localities within the park, all within the tanqua karoo vegetation type of the rainshadow valley karoo bioregion (mucina & rutherford 2006) (figure a-5). figure a-5: distribution of aloe variegata and cheilanthes namaquensis. relevant specimens: northern cape.–3219 (wuppertal): tknp, hill east of leeuberg, (–bb), 04 august 2006, h.m. steyn 888 (ksan, pre); grasberg south 1103, (–bc), 16 august 2008, h.m. steyn 1457 (pre); foot of slope west of potklysberg, (–bd), 05 august 2007, a.c. mudau 254 (pre). 3220 (sutherland): tknp, border of farms manus zyn dam and roodewerf 1019, remhoogte, (–aa), 08 august 2006, r.r. klopper 361 (ksan, pre); maansedam, on the plain surrounding the house, south of the house, (–aa), 06 august 2006, h.m. steyn 941 (ksan, pre). hyacinthaceae top ↑ lachenalia ameliae w.f.barker lachenalia ameliae was thought to be endemic to the western cape, occurring mainly in the montagu and touws river areas. the northernmost specimen was collected in 1921 at gansfontein in the ceres karoo (marloth 10479, pre). during august 2006, 2008 and 2011 four populations were found in three adjacent grids in the north-western parts of tknp. these populations comprise several plants growing on shale-derived soils on gravel plains in the tanqua karoo vegetation type (mucina & rutherford 2006), at altitudes between 352 m and 407 m. this represents a northward extension of the known distribution range by ± 50 km and a new record for the northern cape (figure 2f and figure a-3). because of the new records, the extent of occurrence changed from 4000 km² to 17 696 km², but this is still within the near threatened (nt) range and the status remains unchanged (l. von staden pers. comm., february 2012).relevant specimens: northern cape.–3219 (wuppertal): tknp. sandy wash west of leeuberg, (–bb), 03 august 2006, h.m. steyn 863b (pre); tknp, between varsfontein and prambergfontein, (–bb), 17 august 2008, h.m. steyn 1459 (ksan, pre); tknp, grasberg south 1103, (–bc), 16 august 2008, h.m. steyn 1452 (ksan, pre); tknp, track between luiperdskop and varsfontein, (–bd), 15 august 2011, h.m. steyn 1864 (pre). iridaceae top ↑ moraea tanquana goldblatt & j.c.manning as a result of general collecting, a previously unknown, blue flowering moraea species was collected in tknp in august 2006 (figure 2g). it was subsequently described as m. tanquana (goldblatt & manning 2009). when described, this species was only known from the type locality, but during follow-up field trips to different parts of the park, three more populations were found. in august 2008, a population was found on the lower north-eastern slopes of leeuberg (no specimens were collected), as well as on a plateau to the southeast of grasberg. in august 2009, yet another population on the eastern side of the same plateau was discovered. all three populations were found in tanqua karoo vegetation (mucina & rutherford 2006) on dolerite-derived stony or rocky soil. the populations were estimated to consist of between 80 and 120 plants each and the maximum range extension is between 13 km and 15 km (figure a-6). a preliminary threatened status of vu d1 (vulnerable) was assigned to this species based on the localised distribution and relatively small population sizes (l. von staden pers. comm., february 2010). figure a-6: distribution of moraea tanquana and nemesia suaveolens. relevant specimens: type.–northern cape, 3219 (wuppertal): tknp, small koppie east of leeuberg, (–bb), 04 august 2006, h.m. steyn 872 (nbg, holo.; ksan, pre, iso.).northern cape.–3219 (wuppertal): grasberg south 1103, (–bc), 16 august 2008, h.m. steyn 1431 (pre); hill west of track between varsfontein and luiperdskop, (–bd), 06 august 2009, h.m. steyn 1524 (pre). scrophulariaceae top ↑ nemesia suaveolens k.e.steiner a number of nemesia specimens with yellow and purple or pink flowers were collected (figure 2h). this species was usually found on gravelly sand in the tanqua karoo vegetation type (mucina & rutherford 2006). at the time, these specimens could not be matched to any nemesia species at pre and they were only recently identified as n. suaveolens k.e.steiner. this is an annual species known from a very limited area of the central tanqua karoo. the flowers have a straight spur and a pleasant spicy fragrance (steiner 2009). with this new information the national threatened status of this species was determined as rare, because it has a restricted range (eoo 478 km2) and is potentially vulnerable to grazing and population fluctuations in response to rainfall. subpopulations, however, are large and a large proportion of the population is protected in the tknp (l. von staden pers. comm., february 2010).the known distribution of this species is shown in figure a-6. the original localities are included in this map, not only to highlight the new records, but to rectify the original map (steiner 2009:70). in the original map all the collections were plotted in 3219 (–bc), although specimens were listed from (–ba), (–bc), (–bd), (–da) and 3220 (–ac) (steiner 2009). relevant specimens: northern cape.–3219 (wuppertal): tknp, western end of park. dry riverbed, (–ba), 02 august 2007, s.p. bester 7687 (pre); tknp, between varsfontein and the gate at the ceres-calvinia road, (–ba), h.m. steyn 1434 (pre); calvinia district, tknp, gravel plains west of leeuberg, (–bb), 03 august 2006, b. sachse 31b (pre); calvinia district, tknp, biesjesfontein (–bb), 15 august 2011, s.p. bester 10713 (pre); tknp, grasberg south 1103, (–bc), h.m. steyn 1442 (pre); calvinia district, tknp, luiperdskop farm se of the koppie (–bd), 15 august 2011, s.p. bester 10712 (pre). western cape.–3220 (sutherland): along the road between paulshoek and oudebaaskraal (–ac), 05 august 2009, m. koekemoer 3752 (pre); ceres district, tknp, between roodewerf and oudebaaskraal, platfontein farm (–ac), 13 august 2011, s.p. bester 10706 (pre). pteridophyta: sinopteridaceae top ↑ cheilanthes parviloba (sw.) sw. this fern is widespread from the western cape, through the eastern cape to the south of kwazulu-natal, the free state, lesotho, north west and limpopo, with a disjunction to central namibia, and also occurs in southern zimbabwe (crouch et al. 2011). it was first collected in the tknp in 2006 in the tankwa escarpment shrubland (mucina & rutherford 2006) near the top of gannaga pass. it had not been collected in the northern cape previously; thus this collection represents a range extension of ± 150 km from the nearest known record in the western cape (figure a-7). figure a-7: distribution of cheilanthes parviloba. relevant specimens: northern cape.–3220 (sutherland): tknp, farm kleinfontein 1027. plateau at top of gannaga pass, at the top edge of the pass, (–aa), 05 august 2006, r.r. klopper 319 (ksan, pre). cheilanthes induta kunze cheilanthes induta is a fairly distinctive species, characterised by long, reddish hairs on the stipe, rachis and abaxial lamina surface, as well as its unusual convoluted stipe (crouch et al. 2011). it occurs widespread in the western cape and adjacent parts of the northern cape, as far north as springbok, and the eastern cape, extending into the isolated mountains of the great karoo (crouch et al. 2011). it was first collected in the tknp in 2007 in the roggeveld shale renosterveld (mucina & rutherford 2006) (figure a-8). figure a-8: distribution of cheilanthes induta. relevant specimens: northern cape.–3220 (sutherland): tknp, klipfontein farm, (–aa), 14 september 2007, m.s. mothogoane 736 (pre). cheilanthes namaquensis (baker) schelpe & n.c.anthony cheilanthes namaquensis is a rare fern that occurs from southern namibia, through namaqualand in the northern cape, into the western cape (crouch et al. 2011). it is easily confused with cheilanthes deltoidea kunze subsp. deltoidea that has a very similar, although slightly wider, distribution range. cheilanthes namaquensis can be distinguished by its narrower fronds with basal pinnae that are not basiscopically developed and its rachis that is only winged in the apical portion of the frond and not winged throughout, as in the broadly triangular fronds of c. deltoidea with its basiscopically developed basal pinnae (crouch et al. 2011). this fern was first collected in tknp in 2006 in tankwa escarpment shrubland (mucina & rutherford 2006) (figure a-5).relevant specimens: northern cape.–3220 (sutherland): tknp, farm langekloof 60. langkloof, ± 1.5 km – 2 km north of malansgat river, (–aa), 07 august 2006, r.r. klopper 353 (ksan, pre). filelist convert a pdf file! koedoe 19: 145-152(1976) notes on mammals encountered in the tsitsikama national parks g. a. robinson tsitsikama coastal national park p.o . stormsrivier 6308 abstract data and observations on mammals collected since the proclamation of the tsitsikama national parks in 1964, are presented. although the swedish naturalist j. f. victorin (1970) and other early collectors and explorers have referred to elephants, hippos, buffaloes and many other big game species occurring in the area between knysna and humansdorp, only data and observations collected subsequent to the establishment of the tsitsikama national parks in 1964 will be presented. de graaff and nel (1970) referred to the smaller mammals occurring in the parks while de g raaff (1974) listed the occurrence of rodents. the material observations were collected as a routine duty of field staff in the parks but in a few cases traps were set by visiting professionals. the numbers referred to below the notes of some of the species are the museum accession numbers of the tsitsikama coastal national park housed at storms river mouth. nomenclature follows that of the smithsonian institute (meester and setzer 1971) as far as possible, otherwise roberts (1951) is accepted. order primates family cercopithecidae papio ursinus ursinus (kerr, 1792) chacma baboon widespread and abundant, troops at bloukrans pass, storms river mouth, paul sauer bridge and at nature's valley. the troop at storms river mouth occasionally ventures down to the intertidal zone where they have been observed eating intertidal invertebrates. they are caught in adjacent areas to the park by the department of forestry where they cause considerable damage to the pine (pinus spp.) plantations. cercopithecus pygerythrus pygerythrus (f. cuvier, 1821) vervet frequently seen at groot river near nature's valley and at the de plaat forest near paul sauer bridge although they are found throughout the area in the indigenous forests. the troops usually number between 15-25 individuals. 145 order carnivora family mustelidae mellivora capensis capensis (schreber, 1776) honey badger only one has been observed near lottering river in 1969. a skeleton was, however, found during 1967 while the otter trail was being constructed and it is likely that others exist in this area. tnp ibi 1967 110 poecilogale albinucha albinucha (gray, 1864) snake mongoose a vel)' rare find, collected at the de plaat forest. the distribution of this species seems uncertain and this record probably extends its known distribution. (skin and skull collected). tnp/b/1974/28 aonyx capensis capensis (schinz, 1821) clawless otter found all along the park's coast usually in the vicinity of estuaries. often seen in the late afternoons during the winter months when the sea is rough. most commonly seen at storms river mouth and at groot river estuary. i ctonyx striatus striatus (perry, 1810) cape polecat one specimen collected after it had been run over on the national road near witelsbos. not usually seen due to its nocturnal habits. only the skin was preserved. tnp/b/197512 family viverridae genetta genetta (thunberg, 1811) small spotted genet commonly seen at storms river mouth and since the prohibition of dogs in the park, has become quite tame. three have been trapped and tagged for further studies. herpestes pulverulentus pulverulentus (wagner, 1839) widespread in the parks and regularly seen. family felidae panthera pardus melanotica (gunther, 1885) cape grey mongoose leopard records of sightings at paul sauer bridge, storms river mouth, robbehoek and bloukrans pass. the populations must consist of at least eleven individuals as a mother with three kittens were reported being seen in the bloukrans pass. the specimen on display in the open-air museum (tsitsikama coastal national park), was shot in the langkloof. tnp/b/1968/14 146 felis caracal caracal (schreber, 1776) caracal have been observed on several occasions by rangers and the public. two kittens were caught at stonns river in 1966 by local labourers and reared by the author. order hyracoidea family procaviidae procavia capensis capensis (pallas, 1766) cape dassie extremely abundant in rocky outcrops and since the prohibition of dogs in the park have become very tame at stonns river mouth. the breeding season usually starts during the latter part of november and continues into december. order cetacea sub-order odontoceti family physeteridae kogia breviceps (blainville, 1838) pygmy sperm whale one specimen collected along the coast during 1969 by the port elizabeth museum. it was a male 2,6 m in length. kogia simus (owen, 1866) dwarf spenn whale this identification is doubtful. on saturday 1974.05.31 the author was infonned of a stranded dolphin at mooibaai. it was noted that it was still alive and the beast was hurriedly given a few violent pumps to clear its lungs. the dolphin was dragged back into the water. after swimming around in the bay for a short while it headed for deeper water and was not seen again. j. a. fourie and t. dearlove confinned the tentative identification. this could, however, also have been a k. breviceps. family delphinidae delphinus delphis linnaeus, 1758 common dolphin occurring in enormous schools frequently as large as several thousand individuals. these schools are usually found in deeper water. stenella euphrosyne (gray, 1846) euphrosyne dolphin another pelagic dolphin found in huge schools and not readily distinguishable from the abovementioned species. saayman, bower and taylor (1972) refers to it as stenella caeruleoalbus. 147 tutsiops aduncus (ehrenberg, 1833) bottle-nose dolphin occurring in smaller schools totalling several hundred individuals. seen frequently throughout the year close to the coast. best (1971) has suggested using "gadamn" as this species' common name but, "bottlenose" is more commonly used. solalia lenliginosa (owen, 1866) humpback dolphin seen throughout the year close inshore in small groups. saayrnan el al. (1972) describes this species as sousa sp. best (1971) has suggested that the common name "speckled dolphin" be accepted for this species . family ziphiidae mesoplodon heclori (gray, 1871) hector's beaked whale during march 1967 two specimens were collected at lottering river mouth. their skulls are housed in the port elizabeth museum (1511/15 and 1511/16) . mesoplodon densitostris (blainville, 1817) blainville's beaked whale the most recent stranding (1975.06.26) found at the western end of the caravan park at storms river mouth. it was a female weighing 184,4 kg and had a total length of 2,61 m. all the skeletal material is available. tnp/bi7 5/3 sub-order mysticeti family balaenidae e ubalaena glacialis (m liller, 1 7 76) southern right whale these whales spend their summers closer to the antarctic and are only seen during the winter months migrating to and from the warmer regions. they select protected bays in which to calve and storms river bay has been regularly visited for this purpose. usually seen alone but often in pairs and occasionally a pair with a calf. best (1971) also suggests "back right whale". order pinnipedia family otariidae subfamily arctocephalinae arctocephalus pusillus (schreber, 1776) south african fur seal often seen out at sea but occasionally enters river mouths and has 148 been observed by the author catching fish in the storms river estuary. one or two mortalities occur annually along the tsitsikama coastal national park. family phocidae subfamily monachinae lobodon carcinophagus (hombron andjacquinot, 1842) craheater seal collected at nature's valley beach on 1974.04.15. a rare seal for the south coast and only a few records have been published (nel 1971). the skin is on display at skukuza. tnp/bi1974/1. order chiroptera sub-order megachiroptera family pteropodidae rousettus aegyptiacus (e. geoffroy, 1810) egyptian fruit bat found in large cave in storms river estuary. colony probably consisting of more than 3 000 individuals. other caves visited along the coast and at groot river showed no signs of this bat. reported by b. whitcher to occur in the bloukrans river area (pers. comm.). sub-order microchiroptera family rhinolophidae tnp/b/1972/2. rhinolophus capensis lichtenstein, 1823 cape horseshoe bat found in most of the caves situated along the coast. tnp/b/1 974/26 family vespertilionidae subfamily miniopterinae m iniopterus schreibersi (kuhl,1819) schreiber's long-fingered bat commonly found in the caves caused by faults in the table mountain sandstone layers. tnp/b/1974/22 order artiodactyla family suidae potamochoerus porcus (linnaeus, 1758) bush pig occurs throughout the coastal and forest area. regularly seen at night and judging from its diggings must be fairly abundant. tnp/b/1 974/10 149 family bovidae subfamily bovinae tribe tragelaphini tragelaphus scriptus pallas, 1 766 bushbuck never seen in groups, usually alone, but on occasions a pair have been seen consisting of either a ram and a doe or of a doe and a lamb. tnp/b11967111 taurotragus oryx (pallas, 1766) eland it was.re-introduced by the department of forestry to the lottering forest reserve during 1973 and has subsequently been seen in the immediate vicinity of both the coastal as well as the forest national parks . subfamily cephalophinae c ephalophus monticola (thunberg, 1789) blue duiker recently more observations have been recorded. a nucleus herd of four (two does and two rams) was initiated during 1974. these were presented by the director of the queens park zoo, east london. unfortunately the two rams died shortly after arrival. post-mortems revealed white muscle as the cause of death. subfamily antilopinae tribe neotragini raphicerus melanotis (thunberg, 1811) grysbok a common antelope in the parks usually seen at night. prefers the macchia to the dense forest. order rodentia sub-order hystricomorpha family hystricidae hystrix africae-australis peters, 1852 cape porcupine seems to be quite common in the coastal national park and quills are often found on the otter trail between storms river and groot river. they are not frequently seen, due to their nocturnal habits. family muridae praomys (m astomys) natalensis multimammate mouse collected above caravan park on loerie trail, otter trail and in the restcamp at storms river mouth. seems to be very common. tnp/b/1974/14 150 rhabdomys pumilio (sparrman, 1784) cape striped mouse probably the most common mammal in these parks. found all over and collected at both storms river and groot river. tnp/b/1974118 family cricetidae atomys irroratus irroratus brants, 1827 vlei otomys it is a very common rodent with numerous runs all over the parks. sub-order bathyergomorpha family bathyergidae georychus capensis capensis (pallas, 1 7 78) fairly common at storms river mouth. order chrysochloridea family chrysochloridae amblysomus iris corriae thomas schwann, 1905 tnp/b/1968/9 cape mole-rat tnp /b/ 1969/6 zulu golden mole common in the parks where the soil allows them to burrow. order insectivora family crociduridae myosorex cafer (sundevall, 1847) tnp/bi1974/30 natal forest shrew from the trapped specimens it would appear that these animals are very numerous. tnp/b/1974/16 crocidura jlavescens jlavescens (i. geoffroy, 1827) red musk shrew this shrew is common along the fringes of the coastal forest. collected at storms river mouth. tnp/b11974115 acknowledgements thanks are due to the national parks board of trustees for permission to publish these notes, and to colleagues who helped with the collection and identification of specimens. 151 references best, p. 1971. order cetacea. in: meester j. and h. w. setzer (eds): the mammals of africa: an identification manual. washington, d.c.: smithsonian institution press . de craaff, c. 1974. notes on the occurrence of rodents in south african national parks. koedoe 17: 173-183. de craaff, c. andj. a.j. nel. 1970. notes on the smaller mammals of the eastern cape national parks. koedoe 13:147-149 . nel, j. a. j. 1971. order pinnipedia. in: meester, j. and h. w. setzer (eds): the mammals of africa: an identification manual. washington d.c.: smithsonian institution press. roberts, a. 1951. the mammals of south africa. johannesburg: "the mammals of south africa" book fund. saayman, c. s., d. bower and c. k. taylor. 1972. observations on inshore and pelagic dolphins on the south eastern cape coast of south africa. koedoe 15:1-24. victorin, j. f. 1970. the travels in the cape the years 1853-1855. english translation. cape town: c. struik. 152 page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 filelist convert a pdf file! word of welcome mr chairman, dr. vollmar, ladies and gentlemen! it is indeed a pleasure on behalf of the national parks board of trustees to welcome all the delegates to the symposium which coincides with the 50th anniversary of the national parks board. i think it human after 50 years to think back and to give a summary of the years past. when we talk about nature conservation, it is natural to make a summary of what has been done not only in the republic of south africa (rsa) but in th e whole of southern africa. therefore, knowledgeable persons have been approached to evaluate critically th e position of nature conservation in southern africa . in some instances , certain countries could not be represented due to political upheaval. on behalf of the national parks board of trustees i extend a very hearty word of welcome to each and everyone and to those who have come from afar to take this opportunity of not only noting what has been done in the republic of south africa , but also look with a critical eye on what they think should be done. i would like to welcome representatives of the national parks board of trustees and similarly the welcome is extended to representatives of lesotho, rhodesia, swaziland , bophutatswana, the ciskei, gazankulu, kwa-zulu, lebowa, quaqua, the swazi territorial authority, the republic of the transkei, the venda, the provincial administrations of the rsa, the universities of pretoria and stellenbosch, the south african nature foundation, satour, the national zoological gardens, the national botanical gardens, different government d epartments and last, but not least, th e wildlife society of southern africa. you will permit me to mention one name i.e. dr fritz vollmar and his good lady. dr vollmar is the director general of the world wildlife fund (wwf) employed in finding the necessary funds for the establishment of new national parks all over the world . we are very pleased to have them here. i sincerely hope, ladies and gentlemen, that you will leave our boundaries (after completion of our proposed programme) with optimism and enthusiasm as far as nature conservation is concern ed. we appreciate your presence at this, our 50th anniversary. we shall try our best to do what is still to be done (albeit a long-time programme) and i would emphasise the fact that we are to learn considerably from this symposium on your account the mere fact ofypur presence here being testimonial of our common idealism. the hon s g j van niekerk chairman national parks board qf trustees v page 1 van der merwe.qxd game farms as sustainable ecotourist attractions p. van der merwe and m. saayman van der merwe, p. and m. saayman. 2005. game farms as sustainable ecotourist attractions. koedoe 48(2): 1–10. pretoria. issn 0075-6458. the main goal of this research was to determine aspects that will contribute to the development of sustainable ecotourism on game farms. in order to achieve this, empirical research was done by means of a survey sample consisting of all the active members of the south african game farmers organisation. there are a total of 1244 members, of whom 50 % (n = 622) were randomly sampled for the research. in the main findings of the research, aspects were identified that are developed according to the criterion for sustainable ecotourism. these aspects include natural/conservation/environmental considerations as well as cultural aspects, such as learning local languages. aspects that are neglected include working closely with the local community to develop new products; developing partnerships and joint ventures in which the community has a significant stake; fostering the development of community-based tourism products by providing marketing and mentoring support; and considering using local entrepreneurs in the development of community initiatives. key words: game farm tourism, ecotourism, sustainable development, tourism p van der merwe (ontpvdm@puk.ac.za) and m saayman, north-west university: potchefstroom campus, school of entrepreneurship, marketing and tourism management: institute for tourism and leisure studies, private bag x 6001, potchefstroom, 2520 republic of south africa. issn 0075-6458 1 koedoe 48/2 (2005) introduction for international as well as local tourists, scenic beauty and wildlife remain the major tourism attractions that south africa has to offer (gcis 1998; gcis 1999). ecotourism is responsible for 20 % of the world’s total tourism expenditure. according to the world tourism organisation (wto), the fastest growing of all tourism sectors is any form of tourism in an area of unspoilt nature (van zyl 1999). in southern africa, there is also an increasing awareness of the importance of wildlife, as well as the socio-economic advantages of well-conserved wildlife. this shows that ecotourism is an important product for south africa and a drawcard for international as well as local tourists. van der merwe (2004), points out the important role of game farms in ecotourism and conservation. a game farm is defined as land that is adequately fenced, with a variety of game species that can be used for hunting, photographic opportunities, environmental education, meat production, live game sales and which provides infrastructure and suprastructures for ecotourists. it includes both consumptive and non-consumptive utilisation of wildlife (van der merwe & saayman 2004). eloff (2000) and fox & du plessis (2000) have found that 80 % of nature conservation in south africa is taking place on privately owned land, such as game farms, and according to bothma (2002) there was a 2.5 % increase in land utilised for game farming from 1998 to 1999. the latter translates into an increase of 300 000 ha per year. flack (2002b) reports that it is estimated that the conversion rate from cattle to game farming was nearly 500 000 ha in 2002, which is 200 000 ha more than the 1998 to 1999 increase. south africa has in fact experienced a sharp increase in the supply of game over the past decade because of the growth in game farms and private reserves as well as the fact that one hardly found any game on farms in the previous century (eloff 2002, 2003, 2004). in 2000, it was estimated that there were approximately 7000 privately-owned game farms in south africa, with a total surface area of 16 million ha (ebedes 2002), of which 5061 were exempted game farms, with a surface area of 10 364 154 ha (erasmus 2000). research conducted in 1993 indicated that the surface area of exempted game farms constituted 8.5 % of the total agricultural land in south africa, which increased to 12.5 % in 2000, with a total of 1.7 m head of game (eloff 2002; erasmus 2000). statistics for 2002 showed that 13.3 % of the agricultural land was used for game farming, while national and provincial game reserves only covered 6 % or 6.1 m ha of land in south africa (flack 2002a; van der walt 2002). the wildlife sector in south africa differs considerably from the rest of the world in the sense that game farm tourism is practised with indigenous species that are privately owned. in addition, it seems that game farm tourism in south africa is better controlled and regulated (fox & du plessis 2000). sustainable tourism is based on three pillars namely, economic efficiency, environmental conservation and social equity (coccossis et al. 1996). furthermore, coccossis et al. (1996:9) indicated that from a sustainability point of view it is important that money generated from consumptive use (for example hunting) of wildlife is ploughed back into ecotourism, local communities and nature conservation. due to a lack of research in this regard it is not known whether this is taking place in south africa. mcintosh et al. (1995) also argue that, to be competitive, it is important for game-farm owners to develop competitive products that are sustainable in the long term. van der merwe & saayman (2001) also indicate that in order to be competitive game farms should concentrate on ecotourism, breeding game and rare/endangered game species, processed game products, and hunting (fig. 1). this will also enhance their level of economic sustainability. van der merwe (2004) identified a variety of activities that may be applied by game-farm owners (fig. 1). according to van der merwe & saayman (2004), the type or combination of activities on a game farm is determined by a number of aspects, for example the size and location of the farm, climate, infrastructure and number of game. these authors have found that more activities may result in more income, but that the environment also largely determines which activities can be provided (van der merwe & saayman 2001). eloff (2001), eloff (1999), du plessis (2000), hamman et al. (2003), saayman & van der merwe (2005) and schack (2004) identified a number of problems that game farms are currently experiencing and that will impact on sustainability. these problems include: over-dependence on game; game farms that are too small to allow for greater variety of activities; cross-breeding or hybridisation; freak animals; rand / dollar exchange rate; fire-arms control act; slow implementation of national qualifications framework; long delays for checking in at international arrivals; low level of domestic awareness of game farm ecotourism; little diversification; unregulated relocation of animals; lack of tourism knowledge; game farm owners in general stick to traditional services, for example hunting and breeding of game; lack of game farmers working together to create a better product for tourists; lack of involvement and disinterest among game farm owners. game farm owners are not part of regional or national organisations. as a result they are unaware of the latest developments in the industry. briassoulis (2002) argues that sustainable tourism development revolves around the central issue of how to manage the natural, built, and socio-cultural resources of host koedoe 48/2 (2005) 2 issn 0075-6458 communities in order to meet the fundamental criteria of promoting their economic wellbeing, preserving their natural and socio-cultural capital, achieving intra and inter-generational equity in the distribution of costs and benefits, securing their self-sufficiency, and satisfying the needs of tourists. sustainable tourism development may be seen as tourism development which is responsibly planned and managed. it is the exact opposite of tourism that has been developed for short-term gains (myburgh & saayman 1999). according to goodwin (2002), there are three primary requirements for sustainable tourism. firstly, it must meet the needs of the host community in terms of standards of living (short and long term); secondly, it must satisfy the demands of increasing tourist numbers and continue to attract them in order to achieve an improved standard of living; and thirdly, it must safeguard the environment for this purpose. thus, to achieve true sustainable ecotourism development, one must find a delicate balance between conflicting economic efficiency, environmental conservation and social equity objectives. the resulting economic growth must be distributed equably, and the environmental impacts of these actions must be minimised. the ‘three e’s’ of sustainable tourism may be best explained by a triangle that represents policy. each vertex portrays an imaginary situation in which policy is exclusively dominated by one goal, economic efficiency, social equity or environmental conservation (fig. 2). the first analysis of sustainable tourism may be identified as the policy area near the vertex corresponding to economic efficiency. the second analysis, ecologically sustainable tourism, may be identified near the vertex corresponding to environmental conservation. the third is really a combination of economic efficiency and environmental conservation policies and may be identified somewhere along the side connecting these two vertices of the triangle. the last analysis seems to fit better in the centre of the triangle. there is no one, ideal tourism strategy. the above-mentioned approaches reflect difissn 0075-6458 3 koedoe 48/2 (2005) fig. 1. four pillars of game farm tourism. game farm tourism hunting ecotourism processed game products breed rare/endangered species breed game species trophy hunting biltong hunting bird hunting green safaris bow hunting fishing trips bird watching 4 x 4 trails adventure activities hot-air balloon safaris horse trails mountain bike trails educational walks helicopter game views rhino & elephant walks canoe safaris photographic safaris water-based activities hiking trails sell venison overseas sell venison locally different meat products breeding game and rare/endangered game ferent priorities of which each has its own merits and could be appropriate for different cases and settings, whether mature or emerging destinations, in growth or decline, natural areas or developed resorts (coccossis et al. 1996). if game-farm owners stick to the above-mentioned aspects, they will contribute to achieving the goals of sustainable tourism development, which are explained by inskeep (1991) as follows: to develop a greater awareness and understanding of the significant contributions that game farm tourism can make to the environment, the people and the economy; to promote equity in development; to improve the quality of life of the host community; to provide a high quality of experience for the visitor; to maintain the quality of the environment on which the foregoing goals depend. the outcome of these factors will indicate which aspects are neglected and will impact on the sustainability of game farms in south africa. methods based on the literature review (hobson & essex 2001; mohasi 1999; harris & leiper 1995; myburgh & saayman 2002; milne 1998; yunis 2001; coetzee 2004) a questionnaire was developed with a total of seventy-three items covering sustainability. the questionnaire made use of two different likert scales. the first was a five-point scale and was used to determine the importance of the different activities on game farms, where 1 = not at all important and 5 = extremely important. the second likert scale was a three-point scale and was used to test the importance of the different sustainable development aspects, where 1 = not important; and 3 = very important. the empirical research was conducted from june 2003 until march 2004. the survey sample included all the active members of the south african game farmers organisation, namely: northern game farmers organisation, with 645 members (gauteng, north west, limpopo, and mpumalanga); free state game organisation, with 250 members; northern cape game organisation, with 109 members; natal game ranchers association, with 120 members; western cape game management association, with 20 members; eastern cape game management association, with 100 members. koedoe 48/2 (2005) 4 issn 0075-6458 fig. 2. analysis of sustainable tourism (adapted from coccossis et al. 1996). economic efficiency economic sustainability of tourism sustainable tourism development ecologically sustainable tourism environmental conservationsocial equity sustainability tourism as part of sustainable development in a pilot study, ten questionnaires were administered so as to identify possible constraints and problems. modifications were then made to develop the final questionnaire. the membership added up to a total of 1244, of which 50 % (n = 622) were randomly sampled for the research. a questionnaire was sent to each of the selected respondents via e-mail or fax. members were also visited in person during various workshops that were held by the south african game farmers organisation. a total of 97 questionnaires were received from the selected 622 members. the low response rate (15 % response) was ascribed to the fact that game farm owners were unwilling to complete the questionnaire. to increase the return rate of questionnaires, many follow-up phone calls were made and e-mails were sent to the different game farmers associations and respondents, to request them to complete and return the questionnaires. however, this also had little success. the statistical analysis involved statistical processes such as cronbach alpha (factor analysis) and effect sizes for the relationship in contingency tables (phi coefficient) (steyn 2000). the cronbach alpha statistical process is of major importance for this research. the programme that was used is for statistical analysis was sas institute inc. (sas system for windows release 9.1 ts level 1m0). results firstly, the activities of game farms will be discussed, and secondly the application of the aspects of sustainable development will be addressed. importance of activities respondents had to rate the variety of activities that are currently applied on game farms (table 3). the following activities were rated as very important to extremely important by respondents: breeding game species (69 %); offering trophy hunting (60 %); breeding rare/endangered species (47 %); hunting game for biltong (45 %); offering ‘green’ safaris (39 %). traditional activities, for example biltong hunting or trophy hunting, remain the most issn 0075-6458 5 koedoe 48/2 (2005) table 3 activities according to scale of importance 5 extremely important 4 very important 3 important 2 less important 1 not important at all activities 1 2 3 4 5 breeding rare/ endangered species 21 20 12 20 27 breeding game species 2 8 21 29 40 offering trophy hunting 14 10 16 24 36 hunting game for biltong 22 18 15 23 22 offering bird hunting 38 24 19 8 11 offering green safaris 29 12 20 18 21 offering bow hunting 26 18 17 22 17 offering fishing trips 61 11 10 13 5 selling venison overseas 44 16 12 8 20 selling venison local market 35 13 22 17 13 providing bird watching 13 17 25 22 23 providing hiking trails 16 20 26 20 18 offering photographic safaris 20 18 31 10 21 providing 4 x 4 trails 45 20 16 11 8 providing adventure activities 33 22 24 10 11 providing hot-air balloon safaris 58 26 10 3 3 providing horse trails 41 13 21 12 13 providing mountain bike trails 44 20 21 7 8 providing educational walks 23 15 30 19 13 other helicopter game views rhino and elephant walks canoe safaris game drives with ranger important. this supports the notion that game farm owners do not utilise all four pillars, namely breeding game; hunting (trophy and/or biltong); game sales; and selling game products. this also implies that activities such as walking safaris, horse riding, educational talks, mountain bike trails and hiking do not form part of their product base. a reason for not implementing the four pillars could be that game farms are too small koedoe 48/2 (2005) 6 issn 0075-6458 table 4 factor analysis factor 1 conservation and resources factor loading b7dd invest profit in habitat 0.80 b7aa encourage visitors to respect nature 0.79 b7gg personnel informed 0.77 environmental impact b6cc switch to less polluting fuels. 0.76 b7ee use environmental friendly 0.74 chemicals b4gg proud service among participants 0.73 b6aa measure water quantity and 0.73 energy consumed b7ff work with conservation authorities 0.72 b4hh ensure respect for and dignity of 0.72 local people b6bb measure electricity usage/ saving 0.68 b5gg smallest possible ecological impact 0.68 b4ii ensure that tourism does not 0.60 undermine local resources b7bb no purchase that exploits wildlife 0.56 b7cc enterprise and guests assist in 0.55 conservation b4cc use local guides 0.49 b6jj assist conservation through 0.41 sustainable trails factor 2 local culture / architectural factor loading b5dd avoid noise or light pollution 0.85 b3dd visits by local school 0.82 b5ee buildings with natural ventilation 0.72 b3mm visitor safety 0.70 b3aa respect nature, local culture 0.70 b3ll hospitality 0.69 b3hh local flavour 0.68 b3jj tourists learning a few words 0.67 b4aa commercialisation and 0.62 over-exploitation. b711 keep local character 0.61 b5ff teach visitors about 0.55 environmentally friendly conduct b3ff showcase local cultural artefacts 0.53 b4bb respect local culture 0.52 b3ee scholarships 0.49 factor 3 involvement: tourists/ f a c t o r host community loading b2kkcc cooperation relationship: 0.84 businesses sector b2kkdd cooperation relationship: 0.82 local government b2kkee cooperation relationship: 0.79 government b4ee visitors to interact with 0.73 local culture b2kkbb cooperation relationship: 0.72 tourists b4dd interaction between local 0.63 community and tourists factor 4 environmentally friendly factor products loading b6hh minimise packaging 0 . 8 4 b6ee reduce consumption of 0.80 natural resources b6gg recycling waste produced 0.79 b6ii reduce "food miles" 0.78 b6dd energy used from renewable 0.74 sources b6kk environmentally friendly transport 0.72 issn 0075-6458 7 koedoe 48/2 (2005) table 4 (continued) factor 3 (continued) involvement: f a c t o r tourists/host community loading b2kkaa cooperation relationship: 0.60 local community b2jj monitor progress in 0.59 achieving objectives b4kk educate tourists about 0.58 local culture b1bb create revenue from 0.56 cultural heritage b3kk complementary product 0.55 opportunities b4ff provide visitors with 0.52 reliable information b1aa extend the season 0.48 factor 4 (continued) environmentally friendly factor products loading b4jj negative cultural impacts 0.64 monitored b6ff monitor sewage 0 . 6 1 b5aa plan new developments 0.49 factor 5: local community factor loading b2ee community-based tourism 0.81 b2ii be transparent community 0.72 benefits b2hh use local entrepreneurs 0 . 6 9 b2bb partnerships with community 0.69 b2ff visitors to spend more 0.65 money local economy. b2aa local community products. 0 . 6 4 b2cc identify projects 0.62 benefit the poor b2dd development of the local 0.61 community b5mm conserving fresh water 0.58 resources b2gg including shebeens, 0.57 museums and arts/craft b5ll avoid environmentally 0.53 dangerous substances b3ii cultural history 0.49 local community b3gg maintain the authenticity 0.45 and cultural values factor 6: environmental issues factor loading b5ii minimise environmental impact 0.72 b5bb environmentally friendly products 0.72 b5jj environmentally friendly 0.71 guidelines for construction firms b5hh planting local indigenous species 0.65 b5cc local materials 0 . 6 4 b3bb tourism catalyst for 0.63 human development b5kk fines contractors 0 . 5 9 b7hh no tourists into ecologically 0.59 sensitive areas b3cc respect for social/ 0.41 cultural/religious rights to host all four pillars. game-farm owners are possibly also uninformed in terms of ecotourism as well as the needs of tourists in general. research by van der merwe & saayman (2001) showed that the majority of game farm owners are not trained and educated in the field of tourism. these results support the above findings and confirm that most game farms (56 %) were converted from cattle or crop farms. factor analysis (chronbach alpha) a factor analysis was conducted for the results regarding sustainable ecotourism development. six factors were identified (table 4), and 75.5 % of the total variance in the data was explained by the six factors. all factor loadings above 0.4 were obtained from the data (table 4), which is significant. — factor one: conservation and resources. game farms constituted approximately 12.5 % of the total agricultural land in south africa in 2000. this increased to 13.3 % in 2002, with a total of 1.7 m head of game, while national and provincial game reserves only covered 6 % or 6.1 million ha of land in south africa (eloff 2002; erasmus 2000; flack 2002a; van der walt 2002). hence, game farm owners are clearly very aware of their role in conservation. — factor two: local culture and architecture, which include aspects such as visits by local schools, respect for nature and local culture, a local flavour, retaining the local character and learning a few words of the local language. researchers such as hobson & essex (2001); mohasi (1999); harris & leiper (1995); myburgh & saayman (2002); milne (1998) and yunis (2001) emphasise these aspects as part of sustainable tourism. — factor three: the relationship between tourists and the host community. relevant aspects here are cooperation with local government and national government, interaction between visitors and local culture, cooperation with the local community and the education of tourists regarding the local culkoedoe 48/2 (2005) 8 issn 0075-6458 table 5 standard deviation of factors factor n mean standard coefficient min. max. deviation of variation 1 75 2.44 0.45 18% 1 3 2 77 2.33 0.46 19% 1 3 3 82 2.20 0.44 20% 1 3 4 74 2.26 0.53 23% 1 3 5 81 2.11 0.51 24% 1 3 6 78 2.37 0.39 16% 1 3 ture (hobson & essex 2001; pigram & wahab 1997). — factor four: the use of environmentally friendly products. it was indicated that it is important to minimise packaging that can harm nature, reduce the consumption of natural resources, recycle waste, reduce food miles and monitor sewage and energy from non-renewable resources. — factor five: the local community. the following aspects had a high loading factor: community-based tourism, using local entrepreneurs, partnerships with the local community, visitors spending more money in the local community, buying local community products and visiting shebeens and local museums. — factor six: environmental issues, such as minimising environmental impacts, environmentally friendly products, local indigenous species and not taking tourists into ecologically sensitive areas. this goes hand in hand with conservation and hence it is rated as important by game farmers. table 5 shows the mean and standard deviation, indicating that all factors are considered important to very important, considering the scale that was used in the questionnaire (1 = not important; 2 = important; 3 = very important). this is also evident in columns six and seven of table 5, where the minimum was 1 and the maximum 3. factors that respondents consider very important (column 3 of table 5) are factors one (conservation and resources; mean 2.44), two (local culture/architecture; mean 2.33) and six (environmental issues; mean 2.37). factor five (local community) was not deemed as important as the other factors (mean 2.11). conclusion the primary objective of this research was to determine aspects that will contribute to the sustainable ecotourism development of game farms. these aspects were identified as natural/conservational/environmental considerations as well as cultural aspects, such as learning local languages. aspects that are neglected were: working closely with the local community to develop new products; developing partnerships and joint ventures in which the community has a significant stake; fostering the development of communitybased tourism products by providing marketing and mentoring support; and using local entrepreneurs in developing community initiatives. failure to address aspects like these could result in negative community relationships and even poaching, due to unemployment and poverty. the majority of game farms (56 %) were converted from stock/cattle farms to game farms and have been managed as game farms for 1–10 years (64 %). the high conversion rate has resulted in a situation where the majority of game-farm owners were not trained and educated in tourism. this is in line with other research on tourism-related issues. it also supports the notion that game farmers do not have the required skills to manage tourism as part of game farms. the contribution of this research is captured below. firstly, the statistical analysis of the results gauged the most important aspects of sustainable development as rated by game farmers: (a) promote and ensure respect for and dignity of people in the development, marketing and promotion of tourism; (b) promote a sound, proud service ethic among all participants in the tourism sector; and (c) avoid damage to the environmental quality of the enterprise’s neighbourhood by noise or light pollution. secondly, aspects that need attention and are neglected by game farm owners were also identified. the most important of these are the development of partnerships and joint ventures with the local community; encouraging tour operators to be more innovative in their itineraries by, for example, including shebeens, local museums and arts and craft shops in their tour itineraries; and identifying projects that can support entrepreneurial activities. lastly, this research was among the first of its kind in south africa and has laid the foundation for further research to be accommodated in this field of study. references bothma, j. du p. 2002. some economics of wildlife ranching. pp. 23-29. in: penzhorn, b.l., (ed.). game ranch planning and management. symposium held at onderstepoort, pretoria on 1-2 nov. 2002. pretoria: wildlife group of south african veterinary association. briassoulis, h. 2002. sustainable tourism and the question of the commons. annals of tourism research 29(4):1065-1085. oct. coccossis, h., j.a. edwards & g.k. priestly. 1996. sustainable tourism: european experiences. guilford: biddles. coetzee, w.j.l. 2004. an integrated sustainable tourism development strategy for the vredefort dome as a world heritage site. ph.d thesis, north-west university, potchefstroom. du plessis, p.c. 2000. game farmers not concerned enough. sa game & hunt 6(7): 9. jul. ebedes, h. 2002. preface. pp. 7-9. in: ebedes, h., b. reilly, w. van hoven & b. penzhorn (eds.). sustainable utilization-conservation in practice. proceedings of the 5th international wildlife ranching symposium held at pretoria on 20-23 march 2001. pretoria: south african game ranchers association. eloff, t. 1999. professionalisme in die wildbedryf. klk nuus 29(1): 9. jan. eloff, t. 2000. die omvang van die wildbedryf in suid-afrika. pu vir cho. potchefstroom ongepubliseerd. eloff, t. 2002. the economic realities of the game industry in south africa. pp. 78-86. in: ebedes, issn 0075-6458 9 koedoe 48/2 (2005) h., b. reilly, w. van hoven & b. penzhorn (eds.). sustainable utilization-conservation in practice. proceedings of the 5th international wildlife ranching symposium held at pretoria on 20-23 march 2001. pretoria: south african game ranchers association. eloff, t. 2003. gemiddelde wildveilingpryse 2002. wild & jag 9(2): 21. feb. eloff, t. 2004. game sales 2003 in south africa. africa indaba 2(2): 5. mrt. erasmus, s.d. 2000. die ekonomiese lewensvatbaarheid van ‘n wildvleisbewerkingsaanleg in suid-afrika. m.com. verhandeling, pu vir cho, potchefstroom. flack, p.h. 2002a. the conservation revolution. game & hunt 8(10): 29-33. oct. flack, p.h. 2002b. exotic game catching up with texas? magnum: 76-80. oct. fox, t. & p. du plessis. 2000. hunting in southern africa. africa indigo 3(3): 40-47. gcis (government communication and information system). 1998. south africa official yearbook. cape town: rustica. gcis (government communication and information system). 1999. south africa official yearbook. cape town: rustica. 576 p. goodwin, h. 2002. contribution of ecotourism to sustainable development in africa. wto seminar on planning, development and management of ecotourism in africa. maputo, mozambique held on 5-6 march 2001. hamman, k., s. vrahimis & h. blom. 2003. can current trends in the game industry be reconciled with nature conservation? african indaba 1(5): 3-5. sep. harris, r. & n. leiper. 1995. sustainable tourism: an australian perspective. chatswood: butterworth. hobson, k. & s. essex. 2001. sustainable tourism: a view from accommodation businesses. service industry journal 21(4): 133-146. oct. inskeep, e. 1991. tourism planning: an integrated and sustainable development approach. new york: van nostrand reinhold. mcintosh, r.w., c.r. goeldner & j.r. ritchie. 1995. tourism, principles, practices and philosophies. new york: wiley. 551 p. milne, s.s. 1998. tourism and sustainable development: exploring the global-local nexus. pp. 3549. in: hall, c.m. & a.a. lew (eds.). sustainable tourism. harlow: prentice hall. mohasi, l. 1999. adventure travel and sustainable tourism development in lesotho: the case of pony trekking. m.sc. dissertation, university of natal, pietermaritzburg. myburgh, e. & m. saayman. 1999. ecotourism in action: practical guidelines and principles. first edition. potchefstroom: leisure consultants and publications. myburgh, e. & m. saayman. 2002. ecotourism in action: practical guidelines and principles. second edition. potchefstroom: leisure consultants and publications. pigram, j.j. & s. wahab. 1997. the challenge of sustainable tourism growth. pp. 3-13. in: wahab, s. & j.j. pigram (eds.). tourism, development and growth. london: routledge. saayman, m. & p. van der merwe. 2005. gun laws and its impact on the sustainibility of the hunting industry. wildlife and hunting symposium: facing challenges, 6 july, 2005. lephalale: dbsa. schack, w. 2004. wildboerdery van wêreldgehalte: algemene filosofie, potensiaal, bedreigings, doelwitte. wild & jag 10(4): 22-25. apr. steyn, h.s. (jr.) 2000. practical significance of the difference in means. journal of industrial psychology 26(3):1-3. van der merwe, p. & m. saayman. 2001. managing game farms from a tourism perspective. potchefstroom: ons drukkers. van der merwe, p. 2004. game farms as sustainable ecotourism attractions. ph.d thesis, north-west university, potchefstroom. van der merwe, p. & m. saayman. 2004. managing game farms from a tourism perspective. potchefstroom: leisure c publications. van der walt, j. 2002. proliferation of game ranches. game & hunt 8(10): 7. oct. van zyl, j. 1999. kies ‘n wenplaas met ‘n sakeplan. finansies & tegniek 51(41): 30. nov. yunis, e. 2001. conditions for sustainable ecotourism development and management. [web:] http://www.world-tourism.org/sustainable/ iye/regional_activities/mozambique/spee [date of access: 21 feb. 2004]. koedoe 48/2 (2005) 10 issn 0075-6458 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! supplement to koedoe. 1977: 197-202. interpretation and education in nature conservation j t geddes page director natal parks game and fish preservation board p a box 662 pietermaritzburg 3200 it is pleasant to have an opportunity to deal with the subject of interpretation in nature conservation at a time when so many diversified and often ingenious efforts over the years are becoming formalized into systems with specially trained personnel supplied with specific and reasonabl e budgets to enable them to really get on with the job. preparations for the writing of this paper have led to the decision that education in nature conservation, although unquestionably running as a thread throughout most of our interpretative efforts is truly a very different approach . from earliest 'pupil' nature studies of old faithfuls like the grasshopper and the mealie to the most sophisticated university courses involving most intricate ecological studies of environmental interests , and th e most valuable educational programmes of volunte er societies, education must, in the main, yield pride of place in this contribution to interpretation as practiced by the professionals privileged to control visiting in proclaimed parks, game and nature reserves in the republic of south africa (rsa) today. indulgence is, however, sought for references to overseas examples and illustrations which in fact so often form the basis of our own efforts. objectives if education is the "hard sell" approach for nature conservation, interpretation should be the "soft sell" part of our approach to its primary objective of increasing visitor satisfaction and enjoyment. management objectives can, however, also be facilitated and a high degree of public understanding of the policies of the nature conservation authority can also be obtained. ecological objectives of interpretation include telling the whole natural story by interrelating all plant and animal communities with other natural features of the environment. the identification of the 'superlatives', like the "biggest", "oldest" and "smallest" all adds to visitor enjoyment and often simplifies the task of protecting the "rarest" of species for which we are responsible. 197 tilden's principles tilden's six principles of interpretation are restated as follows: (i) relate interpretative features to something within the experience of the visitor; (ii) interpretation must cover more than information; why is it there?, what are its significance and implications?, and what is not known about it?; (iii) interpretation is an art; using all the senses to build concepts and to comm unica te; (iv) interpretation must be presented as a whole; interrelate features and stories into a conceptual entity; (v) interpretation is provocation; arousing curiosity by describing significance and implications; and (vi) interpretation should be variously directed to specific audiences; children or adults for example. available media media most suited for interpretational use include: (i) personal contact through information desk welcoming staff, guided tours and personal talks; (ii) simulated personal services like recorded audio and visual messagerepeaters; (iii) written material like signs, labels, published brochures, checklists and guides; (iv) self-guided nature trails along 'loop' routes where the visitor increases his enjoyment by 'teaching' himselffrom specially written and illustrated pamphlets; (v) indoor and outdoor displays telling brief interesting facts illustrated by pictures, dendrograms, models, etc; and (vi) visitor centres which can combine most of these features concentrated upon one point. they must be easy of access; include reception desk, auditorium, exhibit area, rest rooms, staff rooms. they usually cost a lot of money, but the value of such investments is incalcuable. visitor centres should ideally be constructed on good quality natural sites en route between entrance and accommodation. they should preferably be sited separated from other buildings and designed to blend into the environment concerned. naturally it has to be accepted that the provision of a full interpretative service can seldom just 'happen'. small starts are more usually the beginnings which can be built up to the ideal of a visitor centre for every major reserve combined with educational programmes provided by the nature conservation authority, by schools, universities, zoos, and voluntary conservation societies. 198 considerations factors which are well worth consideration before de ciding upon interpretational programmes include: (i) where visitors come from and in what proportions; (ii) the average numbers in visitor parties; (iii) their ages and edu cational levels; and (iv) the average length of stays and numbers of 'repeater' visitors. well d esigned interpretation programmes could embrace all or some of th e following steps, although the priorities are not ne cessarily inflexibl e: (i) an inventory of what the reserve has to offer; (ii) the erection of elementary directional and interpretational signs; (iii) th e selection of interesting natural features to form the basis of a nature trail and prepara tion of self-guiding pamphlets for such trails; (iv) roadside stops, shelters and artwork; (v) the appointment of (perhaps seasonal) a ranger(s) (naturalist) and preparation of evening talks; preparation of slides and arranging use of films etc. initially senior students and junior university lecturers offer good holiday-staff material; (vi) the remodelling of an existing building as a visitor ce ntre; th e preparation of models, displays , audio and video a ids plus furth er nature trails; (vii) th e appointm ent of one or more full tim e naturalists and perhaps, other part-time naturalists the preparation of radio and tel evision programmes and the expansion of publications and their distribution; and (viii) the construction of a properly designed visitor centre/s and stepping up of whole programmes and so on. a future as wonderful as those concerned choose to make it! surely non e can doubt the importance of concentration upon this field so vital to beneficial conservation-experiences for th e " piper who plays the tune" the taxpayer. progress let us deal first with the provincial nature conservation authorities and the national parks board of trustees collectively as "the professionals" and secondly with some details of the invaluable work of volunteer societies, " schools " and other operators of educational safaries or trails. the national and provinc!al authorities pride of place in this general field belongs perhaps to the cape province. the inland fisheries department, forerunner of the present nature conservation authority in that province, started in the 'forties' and 'fifties' with publications designed to inform the public on nature 199 conservation subjects and most of us are familiar with their delightful and informative booklets on protected cape flora; "the culture of freshwater fish in south africa"; "the keeping of fish in ponds and aquaria"; handbooks on waterfowl and wildlife management and the propagation of wild flowers. dr d hey deserves special mention for his pioneering work in producing most of these publications, for posters, stickers, radio programmes and regular press features; also for his early personal nature conservation films and for the wonderful educational efforts of the museums of that province. more generally, we find that all the provinces and the national parks board of trustees offer informative annual reports, scientific papers (often in journals rapidly becoming well known like koedoe, madoqua and lammergeyer) as well as brochures, maps and other publications. in addition, growing information and extension services exist, staffed specifically by professional, administrative and technical personnel, but assisted by virtually all the experienced officers in the employ of these authorities who are expected to be willing and able to provide lectures, slide and film illustrated talks to interested clubs, societies and to the public generally. in the orange free state lecturing and nature trails arranged in close collaboration with the education department have proved a great success. the young are considered an especially fertile field in which to sow and cultivate the nature conservation messages. nature trails, accompanied by experienced interpretative officers (naturalists), who are able to put across the nature conservation story to visitors are popular features in increasing numbers of reserves and national parks. individual feature films have been made in the past but television features have also been produced and others are planned as this most powerful medium must be used to best advantage in future. excellent interpretative and aquarium exhibits exist in the head office of the national parks board of trustees, as well as in the head office of the division of nature conservation in pretoria. progress with the building and furnishing of visitor centres at blyde canyon by transvaal and at giant's castle and royal natal by the natal parks board deserve special mention, whilst the stevenson hamilton library in the kruger national park and other visitor centres in national parks throughout the rsa have long made important contributions in this important field. many of the provincial departments, divisions and the natal parks board concentrate upon agricultural and other specialised shows as excellent media for contact with the farming community. prominence has been given to game capture techniques and to the desirability of restocking depleted areas with species which previously occurred there. the previously mentioned educational services of the cape's museums are duplicated by all the national and provincial museums, to a greater or lesser degree, and form an integral part of the cultural lives of many towns and cities in south africa. the efforts of the national 200 zoological gardens in pretoria, however, also deserve special mention. lectures given by trained personnel are aimed at school children, students and adult audiences, and a few statistics indicate the scope and importance of this service. financial years: 1973/ 74 attendance 602 financial years: 1975/ 76 lectures given 68 190 attendance i 161 lectures given 100 795 since the 1969/70 financial year, the service has expanded 300% with 80% of all visitors now visiting both the zoological gardens and the new aquarium and only 20% visiting only the one or the other. programmes are designed particularly: (i) to emphasise that certain species are endangered thereby bringing to the attention of the individual visitor the importance of nature conservation; (ii) to create an interest in the creatures displayed and the countries whence they originated, and (iii) to stimulate visitors to contribute financially or personally to conservation efforts which are presented in the light of their economic, scientific, aesthetic, recreational and prestige values. whereas it is true that many non-indigenous species loom large in zoological garden exhibits, our national organisation has made great strides in using man's fascination for most wild creatures to highlight the importance of nature conservation in our own country. the other organisations (a) it must be clearly repeated that it has proved beyond the ability of the author to cover the basic and extremely valuable contributions of all education departments (both provincial and national), the universities and institutes (often linked to those universities) to the fields of interpretation and nature conservation education in the rsa and it is to be hoped that other may be able to catalogue those efforts elsewhere in due course. (b) of the wildlife and hunting-orientated societies, the efforts of the wildlife society of south africa through its various branches and, in particular, in association with other organisations, deserve particular attention. the society has the largest membership of any comparable group in the rsa and through its meetings, lectures, film shows and particular educational programmes it reaches a huge public. further planning within the society includes a multi-racial educational project for scholars and teacher training throughout southern africa. practical involvement will, it is hoped, become an 201 integral part of schooling, from primary to matric standards, all designed to promote a positive attitude among young south africans, with field excursions serving as an educational and recreational outlet offering opportunities for aesthetic experiences of the wonders of nature under the guidance of trained staff. it is not the wish of the society to usurp the functions of any education authority, but rather to compliment, encourage and assist them to reach the young and to provide the tools, in many cases, so that the job may better be done. in the latter regard, it is to be hoped that this programme will greatly increase the audio-visual aids available for educating the young. (c) "joint venture" is singled out as a particular programme prepared and executed in natal by the combined efforts of the wildlife society, the wilderness leadership school and the natal hunters association. launched in 1974,joint venture was aimed at creating a conservation awareness among children between 11 and 16 years of age. many have contributed to the success of this scheme which offers courses in the u mgeni valley game ranch, the karkloof environmental education park, the lnanda park and at the midmar public resort. (d) the wilderness leadership school and veld & vlei are now world famous for their efforts to assist in producing better leaders for the future by exposing the young (and perhaps some of the not so young) to experiences of a personal 'testing' type either specifically in wildnerness areas or generally through contact with natural environments and with the effects of the forces of nature. conclusion it must be expected in a presentation of this kind that the efforts of some organisations or individuals will have been missed and others, perhaps, over emphasised as a result of the personal experiences and knowledge of the author. let none of these aspects detract from the most important fact which, it is hoped, should emerge from this paper and that is how fortunate we are in the rsa that so many are concious of the need for nature conservation. never before have so many actively sought knowledge and experience in this field; never before have so many been ready, willing and able to offer time and effort to supply the demand. once again it is reiterated that we have (all of us) good cause to give thanks for a hopeful and ever-improving prognosis for the future of nature conservation in south africa. it could be considered irresponsible not to confirm the need for all to increase our efforts, especially in the fields of nature conservation, interpretation and education, but let us also take time off from all our important and continuing battles to improve our lot, to consider for a moment just how lucky we are for what we already have to enjoy. 202 page 1 page 2 page 3 page 4 page 5 page 6 a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe original research vol. 50 no. 1 pp. 3 9 2008 the recent fire history of the table mountain national park and implications for fire management abstract this paper provides an assessment of fire regimes in the table mountain national park over the past four decades. we compiled a gis database of all fires between 1970 and 2007 and analysed the fire regime in terms of the frequency, season and size of fires and the relationship between fire occurrence and fire weather. most fires (90.5% of area burnt) occurred in summer and autumn, the ecologically acceptable season for fires. however, mean fire return intervals declined by 18.1 years, from 31.6 to 13.5 years, between the first and last decades of the record respectively. the area subjected to short (≤ six years) intervals between fires covered > 16% of the park in the last two decades of the record, compared to ~ 4% in the first two decades. a relatively small number of large fires dominated in terms of area burnt. of the 373 fires recorded, 40 fires > 300 ha burnt 75% of the area, while 216 fires < 25 ha burnt 3.4% of the area. fires occurred under a wide range of weather conditions, but large fires were restricted to periods of high fire danger. prescribed burning was a relatively unimportant cause of fires, and most (> 85%) of the area burnt in wildfires. areas subjected to short fire return intervals should be considered for management interventions. these could include the re-establishment of extirpated fire-sensitive species, the clearing of invasive alien plants and increased precautions for the prevention or rapid suppression of future accidental fires. keywords: cape floral kingdom, fire weather, fynbos, ignitions, invasive alien plants the table mountain national park is a rugged 265 km2 area surrounded by the city of cape town, south africa. it obtained world heritage site status in line with its global importance as a hotspot of biodiversity for higher plants and invertebrates (cowling, macdonald & simmons 1996). the dominant vegetation of the area is fynbos, a mediterranean-climate shrubland that is both fire prone and fire adapted (van wilgen, bond & richardson 1992, van wilgen, richardson, kruger & van hensbergen 1992). the park is home to 2 285 plant species, of which 90 are endemic. the park was proclaimed in 1998, prior to which it was managed by 14 separate public bodies; as a result, the earlier conservation management of the area was uncoordinated and fragmented (van wilgen 1996). prior to proclamation as a national park, the area’s unique biodiversity faced numerous threats, including inappropriate fire regimes arising from a lack of clear fire management policies (richardson et al. 1996). one of the major aims of consolidating land parcels into a single conservation area was to promote a coordinated approach to fire management (van wilgen 1996). the current fire management policy of the park is flexible (forsyth et al. 2000). it recognises the need for periodic fire as an important ecological process and that prescribed fire will be needed from time to time. it also recognises that some wildfires are inevitable and that these should be taken into account by adjusting prescribed burning schedules as fire patterns develop. fire management in the park is complicated by an extensive urban-wildland boundary. this means that houses and other property are at risk from wildfires and that fire suppression for nonecological reasons will be needed from time to time. it also means that prescribed fires need to be planned and conducted carefully to avoid the risk of damage. widespread alien plant invasions bring added requirements in terms of fire management; these plants (dominated by trees and shrubs) are spread by fire and require careful treatment prior to burning (richardson et al. 1994). good conservation practice includes both the use of prescribed burning and allowing wildfires to burn in areas where they will do no ecological harm. however, the use of these approaches in close proximity to densely populated areas invites criticism, and managers need good information on past fires to support their actions. in addition, densely settled areas bring increased risks of unwanted ignitions. managers need to be able to identify areas that would be at risk if such fires occurred, so that these areas could be afforded additional protection. future fire management actions within the park will be based on the assessment of fire records. such assessments should identify areas where biodiversity is at risk (either from excessively frequent burning or from protracted fire-free intervals) as well as areas that should be scheduled for burning. the thresholds for fire frequency and season can be determined by plant species’ sensitivity to elements of the fire regime. significant shifts in any of these elements may eliminate some species (van wilgen & forsyth 1992). if these shifts manifest themselves over large areas, management action would be required to counter these trends. to support these assessments, fires that occur in the park are recorded in a geographic information system. this paper provides the first comprehensive assessment of fire regimes in the park over the past four decades. the work aimed to determine whether the fire regimes were within acceptable limits in terms of their impact on biodiversity and if not, how, where and why they deviated from these limits. it also makes recommendations for the improvement of fire management in the park. methods study area the table mountain national park (centred at 34º00’ s, 18º 25’ e) was proclaimed in may 1998 and covers 26 553 ha. the park includes most of the public undeveloped land bordered by the greg g. forsyth brian w. van wilgen centre for invasion biology csir natural resources and the environment south africa correspondence to: brian w. van wilgen e-mail: bvwilgen@csir.co.za postal address: centre for invasion biology, csir natural resources and the environment, p.o. box 320, stellenbosch, 7599, south africa. 3 2008 original research forsyth & van wilgen koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 3 9 city of cape town and its southern suburbs. the park and the city are located on the cape peninsula, a 50 x 10 km mountainous area with about 100 km of coastline. elevations range from sea level to 1 086 m. mean annual rainfall ranges from 1 700 mm in the north to 400 mm in the south, and rain falls predominantly in the winter months (june to august). sandstones and shales of the cape supergroup dominate the geology, and soils are generally sandy and low in nutrients. the predominant vegetation is fynbos, a fire-prone shrubland covering 22 906 ha of the park. renosterveld (a low, grassy shrubland) occurs on shale areas and covers 828 ha of the park; this vegetation type is also fire prone. finally, well-developed afromontane forests covering ~1 000 ha occur in sheltered areas; these tend to be fire free (van wilgen et al. 1990), although forest margins are scorched by fire from time to time. further details of the area are provided by cowling et al. (1996). the fire regime of the area is characterised by dry-season (summer) fires at intervals of between 10 and 20 years (bond 1997) compilation of a fire database fires in the area that now constitutes the table mountain national park have been recorded by earlier land management agencies for many years. typical fire records included a mapped boundary of each fire, along with the dates on which the fire occurred. most records were transferred to the national park following proclamation of the park. we assembled these records and captured the boundaries of all fires > 1 ha in a geographic information system. records were checked for completeness, and a concerted effort was made to locate missing records. newspaper reports, aerial photographs, satellite images and references to unmapped fires were all used to ensure that a comprehensive database was compiled. the earliest fire records were from the 1950s, but reasonably complete records were only available from 1970 onwards. for this reason, we restricted further analyses to the period between 1970 and may 2007. fire return periods the mean fire return period was calculated as rp = y/(b/a), where rp is the return period in years, b is the area of all fires recorded over y years and a is the area over which fires were recorded. we calculated this statistic for the area as a whole as well as for the three dominant vegetation types by overlaying fire records on a vegetation map, using a geographic information system. return periods were also calculated by decade to establish whether any trends in fire frequency were evident. the above method delivers a single estimate of the mean fire return period and does not provide insight into individual fire return intervals (the time between individual fires in the same area). very short fire return intervals can eliminate obligate reseeding shrubs if their juvenile periods are longer than the fire return interval (noble & slatyer 1980, van wilgen & forsyth 1992). in order to assess fire return intervals, we delimited areas of unique fire history by overlaying fire records for the park (de klerk et al. 2007). each polygon of unique fire history was characterised by a number of fire return intervals (except for polygons where no fires were recorded or where only one fire was recorded). we examined the fire record for the occurrence of short fire return intervals (≤ six years) for two separate periods. intervals of ≤ six years were chosen as this approximates the age at which the slower-growing obligate reseeding plants mature. overlapping time periods (1970–1990 and 1985–2007) were used rather than successive periods (1970– 1990 and 1991–2007) to ensure that short-interval fires that occurred between 1985 and 1990 would be identified. fire season we calculated the total area burnt in different seasons for each year on the data record. the seasons were summer (november– february inclusive); autumn (march–april inclusive); winter (may–august inclusive); and spring (september and october). we grouped the annual estimates of the occurrence of fires in different seasons into the periods 1970–1988 and 1989–2007, for comparative purposes. cause of fires fire records included, where available, the cause of the fire. the main causes of fires included fires of human origin (divided into prescribed burns and a range of other causes including arson, escaped barbecue fires, flares, power lines and so on), fires caused by lightning or falling rocks and fires of unknown origin. fire size distribution and fire weather fire size may be important for a number of ecological reasons (for example in terms of the relative proportion of ‘edge effects’ or the distances that must be covered by species reliant on dispersal for recolonising burnt sites). we also wished to examine the relationship between fire size and fire weather. the size of individual fires was examined with a view to determining the proportion of area burnt in fires of different sizes. we also examined the fire record in terms of fire size distribution for the periods 1970–1988 and 1989–2007 to establish whether or not these proportions were changing over time. we obtained daily weather data from a recording weather station at cape town international airport (33º58’s, 18º36’e) for the period 1970 to 2007 and used the mcarthur forest fire danger index (noble et al. 1980) to calculate daily indices of fire danger. the data used included daily maximum temperature and minimum relative humidity, wind speed, the time since the last rain fell and moisture deficit to estimate a fire danger index (fdi). fdi values of 0–5 are considered low, 5–12 moderate, 13– 24 high, 25–50 very high and > 50 extreme. results fire database a total of 373 fires > 1 ha, covering 45 757 ha, were recorded between 1970 and 2007. while some fires could have been overlooked, it is not possible to estimate how large this potential error would be. in addition, not every fire record was complete. not all fire records were accompanied by a full date (day, month and year), and fires covering 19.7% of the total area burnt were associated with a particular year only. for these fires, the season of burning was not known, and it was not possible to associate them with weather conditions on the day of the fire. the causes of fires were also incompletely recorded, and 244 of the 373 fires were of unknown origin. these fires were assumed to be unplanned fires, as the origin of prescribed fires would have been known. figure 1 mean fire return periods per decade in the table mountain national park and in three component vegetation types. note the difference in scale for forest vegetation. 4 fire history of table mountain original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 3 9 fire return periods the mean fire return period for the whole area between 1970 and 2007 was 22 years. mean fire return periods declined in each decade of the fire record (fig. 1). in fynbos vegetation, mean fire return periods declined by 18.1 years from 31.6 years in the 1970s to 13.5 years between 2000 and 2007. equivalent estimates for renosterveld and forest are 37.4 to 17.7 years and 144.6 to 45.6 years respectively (fig. 1). the incidence of short-interval (≤ six yrs) fires also increased significantly from the first half of the data record to the second half (table 1). the extent of these frequently burnt areas was four times greater between 1985 and 2007 than it was between 1970 and 1990 in fynbos vegetation (fig. 2). similarly, frequently burnt areas were five times more extensive in renosterveld. fire season most of the area (90.5%) for which fire season was recorded burnt in summer or autumn (fig. 3). the total area burnt during the first half of the data record was less than in the second half, and the proportion of fires burnt in summer and autumn increased from 86 to 92% between these two periods. a shift away from the practice of prescribed burning in winter and spring (when weather conditions were safer) to autumn could have contributed to this difference. the shift came about with the recognition that prescribed burns in winter and spring would have detrimental effects on the vegetation (van wilgen, bond & richardson 1992). fires in summer and autumn are regarded as optimal for the health of fynbos ecosystems (van wilgen, bond & richardson 1992), while the occurrence of a smaller proportion of fires in other seasons is probably acceptable (van wilgen et al. 1994). from this, we conclude that the historic seasonal distribution of fires within the park should not be a cause for concern. cause of fires the total area burnt in 373 fires on record was 45 757 ha. there were 45 prescribed fires on record, and these burnt 6 661 ha; a further 3 164 ha was burnt in 16 prescribed fires that escaped (and burnt a larger area than intended). prescribed burning thus accounted for only 9 825 ha (21% of the total area burnt), some of which was unintentional; thus most (> 80%) of the area burnt in wildfires. fires of unknown origin (244 fires) burnt 29 118 ha or 63% of the area burnt in all fires. six fires were attributed to natural causes (lightning, five fires and falling rocks, one fire), and these burnt 540 ha or 1% of the area. cba 37.5 18.8 12.5 9.4 < 7.5 fire re turn period (ye ars) n 1985 20071970 1990 n n 5 0 5 kilo meters 5 0 5 kilo meters 5 0 5 kilo meters vegetation type area covered by vegetation type (ha) proportion (%) subjected to short-interval fire 1970–1990 1985–2007 fynbos 22 906 4.6 17.3 renosterveld 828 5.6 29.2 forest 1 009 0.8 0.6 table 1 areas of three vegetation types subjected to at least one short-interval (≤ six yrs) fire during two time periods in the table mountain national park figure 2 fire return periods in the table mountain national park. a = mean fire return periods based on all fires between 1970 and 2007. note that unshaded areas within the park boundary did not burn (i.e. mean fire return period was > 37.5 years), while unshaded areas outside of the park boundary were not included in the analysis. b = the spatial occurrence of short (≤ six yrs) fire return periods between 1970 and 1990. c = the spatial occurrence of short (≤ six yrs) fire return periods between 1985 and 2007. 5 original research forsyth & van wilgen koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 3 9 fire size large numbers of small fires occurred in the table mountain national park, but they burnt a relatively small area; on the other hand, a relatively small number of large fires burnt most of the area (fig. 4). the two biggest fires on record (3 363 and 3 204 ha respectively) both started on the same day (16 january 2000). of the 373 fires on record, only 8 were over 1 000 ha, and the 40 fires > 300 ha burnt 75% of the area. on the other hand, 216 fires were < 25 ha but burnt only 3.4% of the area. the distribution of fire sizes did not differ for the two periods compared, suggesting that fire size distribution is stable (fig. 4). fire weather the analysis of fire weather data and associated fire size revealed that fires in the park can take place under a relatively wide range of weather conditions, including conditions of low fire danger (fig. 5). however, larger fires (> 1 000 ha) only occurred when the fire danger index was in or above the high category, and the two largest fires on record occurred when the index was in the very high category (fig. 5). unfortunately, the complete dates of many of the large fires were not recorded, and these fires are not depicted in fig. 5 (9 out of the 16 largest fires on record, which ranged from 723 to 3 363 ha, did not have full dates). the point should also be made that the table mountain national park experiences relatively mild fire weather conditions when compared to fynbos areas further inland as a result of its proximity to the sea, which exercises a moderating effect on the weather (van wilgen 1984). discussion history of prescribed burning in the park prescribed burning as a land management practice was introduced to most fynbos conservation areas in 1970 (bands 1977). although prescribed burning was accepted as a conservation practice by some land management agencies on the cape peninsula in the 1970s and 1980s, the execution of prescribed fires proved problematic due to public resistance to the practice (van wilgen 1996). by 1996, only 3 out of 10 public land owners, managing just over 30% of the undeveloped land on the peninsula, had adopted the use of prescribed burning (van wilgen 1996). the proclamation of the park in 1998 was followed in 2000 by the formal acceptance of a unified fire management plan, recognising the need for occasional prescribed burning (forsyth et al. 2000). the largest fires on record occurred shortly after this in january 2001. these fires destroyed 14 houses and necessitated the temporary evacuation of many residents, both reducing the immediate need for further burning and hardening public attitudes towards fire. as a result, almost no large prescribed burns were subsequently conducted despite the revised policy. in 2007, a prescribed burn was initiated in the south of the park in an area that had not been burnt for some time; this fire escaped and burnt a larger area than had been planned, resulting in further criticism of the practice. thus, despite overwhelming scientific support for prescribed burning to rejuvenate fire-adapted vegetation and as a vital component of invasive alien plant control, broader public support is not universal. ongoing public awareness programmes will be needed to counter some of this criticism if prescribed burning is to take its place as a useful conservation management practice. increases in short-interval fires short-interval fires in recent years were fairly widespread across the park (fig. 2). the increase in short-interval fires identified in this study is almost certainly attributable to increased ignition sources resulting from increases in human population. the relaxation of rigid influx controls under the apartheid government in the 1980s, which accelerated economic growth over the past decade, and increasing immigration into urban areas have all contributed to the rapid growth of the human population in the area. increases in human population have been shown to be correlated with increases in fire frequency in californian chaparral ecosystems (keeley et al. 1999, keeley & fotheringham 2001), and it appears that similar trends may now be occurring in the table mountain national park. ecological effects of fire in fynbos managers charged with conserving biodiversity in firedependent ecosystems need to assess whether fire regimes remain within acceptable limits. the elements of the fire 0 5 0 0 1 0 0 0 1 5 0 0 2 0 0 0 2 5 0 0 3 0 0 0 a u t u m n s p rin g s u m m e r w in t e r ar ea bu rnt (h a) s e a s o n figure 3 mean annual area burnt in different seasons in all vegetation types in the table mountain national park in two time periods (unshaded bars = 1970–1988; shaded bars = 1989–2007). bars are standard deviations from the mean. figure 4 the relationship between fire size and area burnt in the table mountain national park between 1970 and 2007 (upper figure). the proportion of the area burnt in fires of different sizes did not change over time (lower figure). size classes are small (0–25 ha); moderate (25–100 ha); large (100–500 ha); and very large (> 500 ha). unshaded and shaded columns are for the periods 1970–1988 and 1989–2007 respectively. bars are one standard deviation from the mean. mcarthur forest fdi 0 5 0 0 1 0 0 0 1 5 0 0 2 0 0 0 2 5 0 0 3 0 0 0 3 5 0 0 0 1 0 2 0 3 0 4 0 5 0 fdi a re a bu rn t ( ha ) . figure 5 the fire weather conditions (as measured by the mcarthur forest fire danger index, fdi) associated with fires of different sizes in the table mountain national park between 1970 and 2007. 0 . 0 2 0 . 0 4 0 . 0 6 0 . 0 8 0 . 0 1 0 0 . 0 pr op or tio n of a re a bu rn t ( % ) n u m b e r o f fire s 2 0 0 2 5 0 3 0 0 3 5 0 4 0 01 5 01 0 05 0 0 1 0 2 0 3 0 4 0 5 0 6 0 7 0 s m a ll m e d iu m l a rg e v e ry la rg e f ire s iz e c la s s pr op or tio n of a re a bu rn t ( % ) 6 fire history of table mountain original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 3 9 regime normally assessed with respect to fynbos relate to fire frequency and fire season (van wilgen & scott 2001). in terms of fire season, managers of fynbos ecosystems should seek to keep the majority (> 80–90%) of fires in summer or autumn to avoid negative impacts (van wilgen, bond & richardson 1992). as most (> 90%) fires occur at this time of the year, there is no indication that this aspect should be a cause for concern in the table mountain national park. long fire-free intervals (> 35 years) may also be a cause for concern as they lead to senescence (bond 1980, van wilgen 1982). over the past 37 years, only 12.7 % of the peninsula has remained fire free and about a fifth of this was in fire-independent forest or coastal vegetation. in this regard, therefore, a lack of fire does not present a serious cause for concern. nonetheless, areas that remain fire free for long periods should be identified and monitored to ensure that important fire-dependent species are not affected. on the other hand, the significant increase in fire frequency and the accompanying increase in areas subjected to short-interval fires are causes for concern. most species in fynbos vegetation are resilient to a wide range of fire return intervals. for example, van wilgen and forsyth (1992) established the regeneration strategies of 210 co-occurring fynbos species; of these, most were able to resprout after fire, and only 29 species were classified as obligate seeders (species that have their growth cycle terminated prematurely by fire and are unable to sprout). large, serotinous shrubs with relatively long juvenile periods are an important component among obligate seeders. these species, typically in the family proteaceae, are killed by fire and rely on canopy-stored seed for regeneration (bond, vlok & viviers 1984). while only a small proportion of the total number of species fall into this category (van wilgen & forsyth 1992), they can be the dominant component (in terms of cover and biomass) of the vegetation. short intervals between fires (less than the juvenile periods) can eliminate these species from the vegetation and cause dramatic structural changes (van wilgen 1982). as a result, they are usually the species that are used to determine acceptable fire return intervals (van wilgen, bond & richardson 1992). for example, kruger and lamb (1979) proposed that the minimum interval between prescribed fires in fynbos should be equivalent to the time needed for at least 50% of the individuals in a population of the slowest-maturing of the obligate reseeding species to have flowered and developed seed for at least three successive seasons. application of this rule normally suggests a minimum period of between 10–12 years between fires, and this is currently not being met in over 10% of the park (table 1). there have been several detailed studies on the responses of fynbos to fire in the table mountain national park (bond, le roux & erntzen 1990, cowling & gxaba 1990, moll & gubb 1981, thuiller et al. 2007, yeaton & bond 1991). of the 40 extant species of proteaceae in the park, at least 9 (in the genera protea, leucadendron, leucospermum and mimites) are killed by fire (protea atlas 2007) and rely on seed for regeneration. the juvenile periods of these species are not accurately known but are presumably similar to species of proteaceae elsewhere (i.e. three to six years) (van wilgen & forsyth 1992). comparison of distribution data (protea atlas 2007) with the extent of shortinterval fires (fig. 2) shows that at least five of these (including one endemic) are at risk from frequent burning. in addition, the 34 species of rare or threatened erica (including 29 endemics) (golding 2002, richardson et al. 1996) that occur in the park are likely to be adversely affected by frequent burning, although detailed data on their distribution and ecology are not available. finally, the shrub staavia dodii (also endemic) is known to be at risk (moll & gubb 1981), although its range is currently outside of the areas affected by short-interval fires. there is evidence that variation in fire regimes is necessary to maintain plant diversity in the landscape (thuiller et al. 2007). variation in the intervals between fires, in fire season or in fire intensity will induce variation in the density of overstorey shrubs (for example leucadendrom laureolum), and this variation is in turn associated with the maintenance of diversity in understorey species (cowling & gxaba 1990, thuiller et al. 2007). thuiller et al. (2007) concluded that recurrent fires will safeguard plant populations against extinction by ensuring stable co-existence over time, despite localised extirpation by individual fires. this finding was based on two surveys of plant composition on fixed sites 30 years apart. these sites were, however, in the southern part of the park that had not been subjected to repeated frequent burning (fig. 2). it is well known that repeated frequent burning can eliminate important overstorey shrubs in fynbos (van wilgen 1981, 1982). while we accept that variation in fire regimes is acceptable and even necessary, there are limits beyond which elements of the vegetation may well suffer. ecological effects of fire in non-fynbos vegetation fire frequencies and the occurrence of short-interval fires have also increased in renosterveld vegetation over the past few decades. this is not expected to have serious consequences for species diversity, but data are lacking to test this hypothesis. while the fire regimes in renosterveld vegetation have been poorly studied (van wilgen 1987), fire return intervals are assumed to be shorter than those in fynbos due to the dominance of species with short maturation times (boucher 1983). at present there is no evidence that increases in fire frequency in renosterveld are a cause for concern. the apparent increase in fire frequency (fig. 1) in forests is probably not a cause for concern. most forest vegetation has remained fire free (table 1), but where fires do occur, they appear to be getting more frequent (although not as frequent as in fynbos). the vegetation mapped as forest by cowling et al. (1996) included areas of scrub thicket with a potential to develop into forest. these authors recognised that prolonged (‘centuries-long’) fire-free periods would be needed to allow forests to develop on these sites. afromontane forests persist in fire refugia in fynbos vegetation, and while their expansion into surrounding areas would be possible under current climates, this is limited by recurrent fires (manders & richardson 1992). the apparent increase in fire frequency in forest vegetation is probably a result of fires in scrub thicket as well as the fact that areas recorded as being burnt would contain forest patches that in fact did not burn. at present there is no evidence that fires are causing forest patches to shrink. comparison to other areas the fire management situation in the table mountain national park bears strong similarities to some other areas in fireprone mediterranean climates. for example, the santa monica national park (los angeles, california) and the blue mountain national park (sydney, australia) are both important fire-prone conservation areas within an urban setting. as with the table mountain situation, both have a substantial urban-wildland interface, and both are subjected to periodic wildfires that cause substantial damage to property (bradstock et al. 1998, keeley & fotheringham 2006). in the los angeles situation, fire frequency has been shown to be increasing over the past century, and this is linked to increasing human populations, providing increasing sources of ignition (keeley & fotheringham 2001). this problem now appears to be manifesting itself in the table mountain case. in both cases, increasing fire frequency will impact on the survival and post-fire recovery of dominant shrubs, and in both cases invasive alien plants are able to take the place of dominant indigenous shrubs, with negative consequences for the conservation of biodiversity and ecosystem services. in the californian case, the alien species are grasses, which promote more frequent fire (keeley, keeley & fotheringham 2005). in the table mountain case, they are largely alien trees and shrubs, which increase fire intensity and erosion after fire (van wilgen & scott 2001). despite the similarities, the scale of the fire problem is orders of magnitude larger in los angeles 7 original research forsyth & van wilgen koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 3 9 bond, w.j., le roux, d. & erntzen, r. 1990. fire intensity and regeneration of myrmecochorous proteaceae. south african journal of botany 56: 326–331. bond, w.j., vlok, j. & viviers, m. 1984. variation in seedling recruitment of cape proteaceae after fire. journal of ecology 72: 209–221. boucher, c. 1983. floristic and structural features of the coastal foreland vegetation south of the berg river, western cape province, south africa. bothalia 14: 669–674. bradstock, r.a., gill, a.m., kenny, b.j. & j. scott, j. 1998. bushfire risk at the urban interface estimated from historical weather records: consequences for the use of prescribed fire in the sydney region of south-eastern australia. journal of environmental management 52: 259–271. cowling, r.m. & gxaba, t. 1990. effects of a fynbos overstorey shrub on understorey community structure: implications for the maintenance of community-wide species richness. south african journal of ecology 1: 1–7. cowling, r.m., macdonald, i.a.w. & simmons, m.t. 1996. the cape peninsula, south africa: physiographical, biological and historical background to an extraordinary hot-spot of biodiversity. biodiversity and conservation 5: 527–550. de klerk, h., schutte-vlok, a., vlok, j., shaw, k., palmer, g., martens, c., viljoen, p., marshall, t., van ross, g., forsyth, a.t., wessels, n., geldenhuys, d., wolfaardt, a. & kirkwood, d. 2007. ecological fire monitoring manual. stellenbosch: cape nature scientific services. forsyth, g.g., van wilgen, b.w., ruddock, g., nel, j.l., le maitre, d.c., smith, c., cullinan, c., & chapman, r.a. 2000. a fire management plan for the cape peninsula national park. report no. env-s-c 2000-111, csir, stellenbosch. golding, j. 2002. southern african plant red data lists. national botanical institute, pretoria. keeley, j.e. & fotheringham, c.j. 2001. historic fire regime in southern california shrublands. conservation biology 15: 1536–1548. keeley, j.e. & fotheringham, c.j. 2006. wildfire management on a human-dominated landscape: california chaparral wildfires. in: wuerthner, g. (ed.). the wildfire reader: a century of failed forest policy. pp 116–131. washington: island press. keeley, j.e., fotheringham, c.j. & morais, m. 1999. re-examining fire suppression impacts on brushland fire regimes. science 284: 1829–1832. keeley, j.e., keeley, m.b. & fotheringham, c.j. 2005. alien plant dynamics following fire in mediterranean-climate california shrublands. ecological applications 15: 2109–2125. kruger, f.j. & lamb, a.j. 1979. conservation of the kogelberg state forest. preliminary assessment of the effects of management from 1967 to 1978. report 79-02, jonkershoek forestry research centre, stellenbosch. manders, p.t. & richardson, d.m. 1992. is fynbos a stage in succession to forest? analysis of the perceived ecological distinction between two communities. in: van wilgen, b.w., richardson, d.m., kruger, f.j. & van hensbergen, h.j. (eds.). fire in south african mountain fynbos: species, community and ecosystem response in swartboskloof. pp 81–107. heidelberg: springer verlag. moll, e.j. & gubb, a.a. 1981. aspects of the ecology of staavia dodii in the south western cape of south africa. in: h. synge. (ed.). the biological aspects of rare plant conservation. pp 331–342. new york: john wiley & sons. noble, i.r. & slatyer, r.o. 1980. the use of vital attributes to predict successional changes in plant communities subject to recurrent disturbances. vegetatio 43: 5–21. noble, i.r., gill, a.m & bary, g.a.v. 1980. mcarthur’s fire danger meters expressed as equations. austral ecology 5: 201–203. protea atlas. 2007. (http://protea.worldonline.co.za/idm_ cp.htm). maintained by rebelo a., south african national biodiversity institute, kirstenbosch, south africa. (accessed 26 august 2007). richardson, d.m., van wilgen, b.w., higgins, s.i., trinder-smith, t.h., cowling, r.m. & mckelly, d.h. 1996. current and or sydney when compared to the table mountain area. for example, the 2003 fires in los angeles burnt 364 000 ha and destroyed 3 361 homes (keeley & fotheringham 2006). the fires in december 2001 in sydney burnt 250 000 ha and destroyed 109 homes (shakesby et al. 2003). in contrast, the fires in january 2000 in the table mountain national park (the worst by far on record) only burnt 7 000 ha and destroyed 14 homes (van wilgen & scott 2001). these differences could be attributed to higher fuel loads and more severe fire weather in australia and california than in the table mountain area. fire and conservation in urban-dominated landscapes as a result of the establishment of a comprehensive spatial database on fires, managers in the table mountain national park should now be able to make better and more informed decisions about fire management. the key will be to maintain and regularly update the database and to analyse it periodically. the existence of relevant analyses will enable managers to identify areas of concern (such as areas that have been subjected to repeated, short-interval fires) and to take appropriate action. it will also provide them with a defensible basis for making decisions about the ignition of prescribed burns or the tolerance of wildfires. both of these will usually elicit comment and possibly criticism from interested and affected parties. it is also important that managers ensure that fire records are complete so that the best possible information can be used to support management decisions. we found, for example, that several fire records were incomplete in that the dates, durations and causes were not always consistently recorded, and this will weaken any subsequent analysis. appropriate responses to human-caused increases in fire frequency would be to increase both vigilance and public awareness in areas close to human habitation, to improve the chances of preventing further ignitions and/or the early identification and rapid containment of fires should they occur in such priority areas. secondly, in areas where short-interval fires have been found to occur, it would be prudent to conduct surveys of plant regeneration as soon as possible following fire and to consider, for example, the reintroduction of important plant species should they be found to have been eliminated from the area. finally, the situation with regard to the regeneration and spread of invasive alien plants in areas subjected to higherthan-normal fire frequencies should be carefully managed. it is often the case that invasive alien species are superior competitors in the face of fire regimes that differ from those under which the indigenous biota evolved, and this represents a significant ongoing threat to the conservation of the park’s unique flora. acknowledgements this work was jointly funded by south african national parks, cape nature and the south african council for scientific and industrial research. we thank south african national parks for access to fire records and lucille schonegevel and patrick o’farrell for assistance with data capture and processing. references bands, d.p. 1977. prescribed burning in the cape fynbos. in: mooney, h.a. & conrad, c.e. (eds.). proceedings of the symposium on the environmental consequences of fire and fuel management in mediterranean ecosystems. pp 245–256. usda forest service, general technical report wo-3. bond, w.j. 1980. fire and senescent fynbos in the swartberg, southern cape. south african forestry journal 114: 68–74. bond, w.j. 1997. fire. in: cowling, r.m., richardson, d.m. & pierce, s.m. (eds.). the vegetation of southern africa. pp 421– 446. cambridge: cambridge university press. 8 fire history of table mountain original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 3 9 future threats to plant biodiversity on the cape peninsula, south africa. biodiversity and conservation 5: 607–647. richardson, d.m., van wilgen, b.w., le maitre, d.c., higgins, k.b., & forsyth, g.g. 1994. a computer-based system for fire management in the mountains of the cape province, south africa. international journal of wildland fire 4: 17–32. shakesby, r.a., chafer, c.j., doerr, s.h, blake, w.h., wallbrink, p., humphreys, g.s. & harrington, b.a. 2003. fire severity, water repellency characteristics and hydrogeomorphological changes following the christmas 2001 sydney forest fires. australian geographer 34: 147–175. thuiller, w., slingsby, j.a., privett, s.d.j. & cowling, r.m. 2007. stochastic species turnover and stable coexistence in a species-rich, fire-prone plant community. plos one 2(9): doi:10.1371/journal.pone.0000938. van wilgen b.w., bond, w.j. & richardson, d.m. 1992. ecosystem management. in: cowling, r.m. (ed.). the ecology of fynbos: nutrients, fire and diversity. pp 345–371. cape town: oxford university press. van wilgen, b.w. & forsyth, g.g. 1992. regeneration strategies in fynbos plants and their influence on the stability of community boundaries after fire. in: van wilgen, b.w., richardson, d.m., kruger, f.j. & van hensbergen, h.j. (eds.). fire in south african mountain fynbos: species, community and ecosystem response in swartboskloof. pp 54–80. heidelberg: springer verlag. van wilgen, b.w. & scott, d.f. 2001. managing fires on the cape peninsula: dealing with the inevitable. journal of mediterranean ecology 2: 197–208. van wilgen, b.w. 1981. some effects of fire frequency on fynbos plant community composition and structure at jonkershoek, stellenbosch. south african forestry journal 118: 42–55. van wilgen, b.w. 1984. fire climates in the southern and western cape province and their potential use in fire control and management. south african journal of science 80: 358–362. van wilgen, b.w. 1996. management of the natural ecosystems of the cape peninsula: current status and future prospects. biodiversity and conservation 5: 671–684. van wilgen, b.w. 1982. some effects of post fire age on the above ground biomass of fynbos (macchia) vegetation in south africa. journal of ecology 70: 217–225. van wilgen, b.w. 1987. fire regimes in the fynbos biome. in: cowling, r.m., le maitre, d.c., mckenzie, b., prys-jones, r.p. & van wilgen, b.w. (eds.). disturbance and the dynamics of fynbos biome communities. pp 6–14. south african national scientific programmes report no. 135. pretoria: csir. van wilgen, b.w., higgins, k.b. & bellstedt, d.u. 1990. the role of vegetation structure and fuel chemistry in excluding fire from forest patches in the fire-prone fynbos shrublands of south africa. journal of ecology 78: 210–222. van wilgen, b.w., richardson, d.m. & seydack, a. 1994. managing fynbos for biodiversity: constraints and options in a fire-prone environment. south african journal of science 90: 322–329. van wilgen, b.w., richardson, d.m., kruger, f.j. & van hensbergen, h.j. 1992. fire in south african mountain fynbos: species, community and ecosystem response in swartboskloof. heidelberg: springer verlag. yeaton, r.i. & bond, w.j. 1991. competition between two shrub species: dispersal differences and fire promote coexistence. american naturalist 138: 328–341. 9 vol. 50 no. 1 pp. 10 17koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research a checklist of the spiders (arachnida, araneae) of the polokwane nature reserve, limpopo province, south africa abstract as part of the south african national survey of arachnida (sansa), spiders were collected from all the field layers in the polokwane nature reserve (limpopo province, south africa) over a period of a year (2005–2006) using four collecting methods. six habitat types were sampled: acacia tortillis open savanna; a. rehmanniana woodland, false grassland, riverine and sweet thorn thicket, granite outcrop; and aloe marlothii thicket. a total of 13 821 spiders were collected (using sweep netting, tree beating, active searching and pitfall trapping) represented by 39 families, 156 determined genera and 275 species. the most diverse families are the thomisidae (42 spp.), araneidae (39 spp.) and salticidae (29 spp.). a total of 84 spp. (30.5%) were web builders and 191 spp. (69.5%) wanderers. in the polokwane nature reserve, 13.75% of south african species are presently protected. keywords: arachnida, araneae, diversity, habitats, conservation in the early 1990s, south africa was recognised, in terrestrial terms, as a biologically very rich country and even identified as the world’s ‘hottest hotspot’ (myers 1990). after the united nations conference on environment and development in 1992, south africa ratified the convention on biological diversity in 1995 with the main goal to develop a strategic plan for conservation and sustainable use of biodiversity. one of the national efforts identified as necessary to accomplish the goal was to discover, describe and to make an inventory of the species of south africa. as a result of this, the south african national survey of arachnida (sansa) was established in 1997 with the aim to develop an inventory of the arachnofauna of south africa (dippenaar-schoeman & craemer 2000). arachnids are well represented in south africa, with nine of the twelve arachnid orders occurring here. however, even though they constitute an abundant and successful group, they are still poorly sampled with little existing knowledge about their diversity and distribution within certain ecoregions. south africa has a very rich spider fauna represented by 69 families, 469 genera and about 2 000 species that occur in all of the ecoregions of south africa (dippenaar-schoeman 2002a). unfortunately, the lack of taxonomic expertise makes it impossible to identify some spider families to species and even generic levels, which could lead to under-appreciation and estimation of the actual species pool and undermines meaningful conservation. as part of sansa, this study addressed specifically the objective to compile species lists within the savanna biome, starting with checklists of the conserved areas where conservation strategies are already in place. the savanna biome is one of the world’s major biomes and covers about one third of south africa (low & rebelo 1996). it is especially well developed in the kalahari, parts of limpopo and kwazulu-natal, mpumalanga and the eastern cape. savanna is characterised by a grassy ground layer and a distinct upper layer of woody plants that can be identified as shrubveld, woodland or bushveld. the delimiting factor is complex for the savanna biome and includes mainly rainfall (235 to 1 000 mm per year) and/or frost (0–120 days/year). reports on the spiders of the savanna biome in south africa are restricted to that of roodeplaat dam nature reserve (dippenaar-schoeman et al. 1989), makalali game reserve (whitmore et al. 2001, 2002), western soutpansberg (foord et al. 2002), kruger national park (dippenaar-schoeman & leroy 2003), sovenga hill (modiba et al. 2005) and ndumo game reserve (haddad et al. 2006). this is the second survey of arachnids from the polokwane region, limpopo province. the study was undertaken in the polokwane nature reserve (pnr) which lies in the savanna biome, and is situated in the central bushveld bioregion, specifically the polokwane plateau bushveld vegetation unit (mucina et al. 2005, mucina & rutherford 2006). a significant portion of the reserve includes a unique and threatened habitat type known as the pietersburg plateau false grassland. the reserve was given conservation status and was proclaimed as a municipal reserve after officially being opened in 1971. currently the reserve conserves one of the largest pristine examples of this habitat type, along with its associated plant and animal species. as part of the msc studies of the third and fourth authors, spiders were sampled from six habitat types over a period of a year using four collecting methods, sampling all the field layers. this paper is the first in a series of papers on the reserve and provides only an annotated checklist for the spiders of the reserve. detailed information about the abundance and species richness of spiders collected from different habitat types, during different seasons and using different sampling methods will be published in succeeding papers. susan m. dippenaar 1department of biodiversity school of molecular & life sciences university of limpopo south africa ansie s. dippenaar-schoeman arc-plant protection research institute south africa mokgadi a. modiba1 thembile t. khoza1 correspondence to: susan m. dippenaar e-mail: susand@ul.ac.za postal address: private bag x1106, sovenga 0727, republic of south africa 10 2008 spiders: arachinda, araneae original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 10 17 study area the polokwane nature reserve (23º58’s 29º28’e) covers an area of approximately 3 200 ha and lies on an elevated plateau with an altitude ranging between 1 200 and 1 500 m above sea level. the climate is moderate with mean summer day-time high of 28 ºc and a mean winter night-time low of 5 ºc. the average annual rainfall for the area is 478 mm. the polokwane plateau bushveld vegetation unit is characterised by open themeda grassland with scattered acacia trees and bush clumps. the dominant trees are acacia tortillis (forssk.) hayne and a. rehmanniana schinz. other interesting plant communities and habitats in the reserve include riverine and sweet thorn thickets, granite outcrops, quartzite pebble slopes, saline patches and a mountain aloe, aloe marlothii a. berger thicket situated on a shale outcrop (grosel pers. comm.). methods six different habitat types were selected subjectively to represent most of the habitat diversity in the reserve and included: 1. acacia tortillis open savanna, 2. a. rehmanniana woodland, 3. false grassland, 4. riverine and sweet thorn thicket, 5. a northern slope granite outcrop, and 6. aloe marlothii thicket on shale outcrop. four collecting methods were used, namely grass sweep netting, tree beating, active searching and pitfall trapping. leaf litter sifting was attempted once on the a. rehmanniana woodland site but was discontinued because of insufficient leaf litter. sampling was conducted once a month in each of the six habitat types for a duration of 12 months (march 2005 to february 2006). pitfall trapping: containers with a 10 cm diameter were planted with the upper rim level with the ground surface and covered by a funnel leading to a smaller container filled with 70% etoh to immobilise and preserve caught specimens. ten traps, 10 m apart, were planted per habitat site, consisting of a central trap with three arms in three different directions, each arm consisting of three traps. the traps were checked every day during the week of sampling per site. the pitfall traps were open for five consecutive days, resulting in 60 trap days per site and therefore 360 trap days for all sampling sites during the sampling period of one year. this resulted in a total of 3 600 pitfall trap samples taken during the study. sweep netting: a sweep net with a diameter of 31 cm was used to sample through grass and herbs. sweeping was conducted once a month for two hours at each site, while walking from the central pitfall trap throughout the sampling site. collected material was gathered into marked plastic bags and sorted in the laboratory. tree beating: trees and shrubs were beaten with a 0.21 kg baton to dislodge specimens which were caught on a sheet and collected in plastic bags. bigger trees received up to 20 beatings while smaller trees and shrubs received fewer beatings. this method was applied once a month for two hours at each site. collected material was sorted in the laboratory. active searching was undertaken once a month for two hours at each site. the area covered was from the central pitfall trap working outwards and covered the whole sampling site. specimens were searched and collected from all field layers. collected spiders were fixed and preserved in 70% etoh and identified up to family level using the african spider family keys (dippenaar-schoeman & jocqué 1997) while genus and species determinations were done by the second author. voucher specimens were deposited in the national collection of arachnida (nca) at the plant protection research institute in pretoria, an institute of the agricultural research council. two main guilds can be distinguished among spiders, namely wanderers and web builders. the wanderers can be further divided into plant wanderers (pw) and ground wanderers (gw). plant wanderers were separated and placed in a guild based on the vegetation type that they were most commonly found on, namely plant wanderers found on foliage (pwf), plant wanderers found on grass (pwg) and plant wanderers found on bark (pwb) while ground wanderers may be free living or burrow dwelling (bd). the web-building spiders can also be subdivided into different guilds based on the structure of the webs that they build: sheet-web builders (swb), spaceweb builders (spwb), orb-web builders (owb), funnel-web builders (fwb), retreat-web builders (rwb), tube-web builders (twb) and gumfoot-web builders (gwb). results and discussion a total of 13 821 spiders were caught during the sampling period, representing 39 families, 156 determined genera and 275 species (see table 1). the 39 families represent 56.5% of the currently recognised families for south africa (dippenaarschoeman & jocqué 1997). the representation and numbers caught (irrespective of other factors such as length of sampling period, size of sampling area etc.) compare favourably with table 1 spider families collected in the polokwane nature reserve indicating the number of species and percentage of the total number of specimens collected families genera species specimens % agelenidae 2 2 11 0.08 amaurobiidae 1 1 1 0.01 ammoxenidae 1 1 61 0.44 araneidae 23 39 916 6.63 barychelidae 1 1 3 0.02 caponiidae 1 1 16 0.12 clubionidae 1 2 17 0.12 corinnidae 4 4 7 0.05 ctenidae 1 1 3 0.02 cyrtaucheniidae 1 1 6 0.04 dictynidae 1 1 5 0.04 eresidae 3 3 2 547 18.43 gnaphosidae 13 25 499 3.61 hersiliidae 1 1 11 0.08 idiopidae 3 3 7 0.05 linyphiidae 4 4 5 0.04 lycosidae 8 21 784 5.67 mimetidae 1 1 4 0.03 miturgidae 1 3 289 2.09 nemesiidae 1 1 2 0.01 nephilidae 1 1 31 0.22 oonopidae 1 1 1 0.01 oxyopidae 3 16 3 415 24.71 palpimanidae 2 3 53 0.38 philodromidae 7 12 621 4.49 pholcidae 1 1 4 0.03 pisauridae 5 6 232 1.68 prodidomidae 1 1 9 0.07 salticidae 18 29 1 056 7.64 scytodidae 1 1 2 0.01 segestriidae 1 1 4 0.03 selenopidae 2 2 9 0.07 sparassidae 3 5 25 0.18 tetragnathidae 1 2 4 0.03 theraphosidae 3 3 10 0.07 theridiidae 9 22 778 5.63 thomisidae 17 42 2 212 16.00 uloboridae 2 4 42 0.30 zodariidae 6 7 119 0.86 total 156 275 13 821 100 (*value excluding undetermined genera) 11 original research dippenaar, dippenaar-schoeman, modiba, khoza koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 10 17 table 2 checklist of the spiders of the polokwane nature reserve, limpopo province, south africa (pw: plant wanderer; gw: ground wanderer; bd: burrow dweller; swb: sheet-web builder; spwb: space-web builder; owb: orb-web builder; fwb: funnel-web builder; rwb: retreat-web builder; gwb: gumfoot-web builder; twb: tube-web builder) family species guild field type agelenidae agelena sp. (immature) fwb on soil olorunia sp. (immature) fwb on soil amaurobiidae pseudauximus sp. (immature) rwb debri ammoxenidae ammoxenus amphalodes dippenaar & meyer, 1980 gw soil araneidae acanthepeira sp. * owb vegetation araneidae sp. (genus undetermined) owb vegetation araneus apricus (karsch, 1884) owb vegetation araneus coccinella pocock, 1898 owb vegetation araneus nigroquadratus lawrence, 1937 owb vegetation araniella sp. 1* owb vegetation araniella sp. 2* owb vegetation araniella sp. 3* owb vegetation argiope aurocincta pocock, 1898 owb vegetation argiope australis (walckenaer, 1805) owb vegetation argiope lobata (pallas, 1772) owb vegetation argiope trifasciata (forskal, 1775) owb vegetation argiope sp. (immature) owb vegetation caerostris sexcuspidata (fabricius, 1793) owb vegetation chorizopes sp. * owb vegetation cyclosa insulana (costa, 1834) owb vegetation cyphalanothus larvatus (simon, 1881) owb vegetation cyrtophora citricola (forskal, 1775) owb vegetation gasteracantha sanguinolenta c.l. koch, 1884 owb vegetation gea infuscata tullgren, 1910 owb vegetation hypsosinga lithyphantoides caporiacco, 1947 owb vegetation hypsosinga sp. 2 owb vegetation larinia natalensis (grasshoff, 1971) owb vegetation lipocrea longissima (simon, 1881) owb vegetation mahembea hewitti (lessert, 1930) owb vegetation nemoscolus elongatus lawrence, 1947 owb vegetation nemoscolus sp. 2 owb vegetation neoscona blondeli (simon, 1886) owb vegetation neoscona moreli (vinson, 1863) owb vegetation neoscona penicillipes (karsch, 1879) owb vegetation neoscona quincasea roberts, 1983 owb vegetation neoscona subfusca (c.l. koch, 1837) owb vegetation paraplectana sp. 1 owb vegetation pararaneus cyrtoscapus (pocock, 1898) owb vegetation prasonica sp. 1* owb vegetation pycnacantha tribulus (fabricius, 1781) owb vegetation singa lawrencei (lessert, 1930) owb vegetation singa sp. 2 owb vegetation singa sp. 3 owb vegetation caponiidae caponia chelifera lessert, 1936 gw soil clubionidae clubiona abbajensis strand, 1906 pw vegetation clubiona sp. (immature) pw vegetation corinnidae casteineira sp. 1 gw soil cetonana simoni (lawrence, 1942) gw soil copa flavoplumosa simon, 1885 gw soil graptartia mutillica haddad, 2004 gw soil ctenidae ctenus sp. 1 gw soil cyrtaucheniidae ancylotrypa brevipalpis (hewitt, 1916) bd burrow soil dictynidae archaeodictyna sp. 1 rwb vegetation eresidae dresserus colsoni tucker, 1920 rwb soil debri gandanameno fumosus (c.l. koch, 1837) rwb tree stegodyphus dumicola pocock, 1898 rwb tree * possibly new 12 spiders: arachinda, araneae original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 10 17 family species guild field type gnaphosidae aneplasa sp. 1 asemesthes ceresicola tucker, 1923 gw soil asemesthes decoratus purcell, 1908 gw soil camillina aestus tucker, 1923 gw soil camillina maun platnick & murphy, 1987 gw soil camillina procurva (purcell, 1908) gw soil drassodes bechuanicus tucker, 1923 gw soil drassodes solitarius purcell, 1907 gw soil drassodes splendens tucker, 1923 gw soil drassodes stationis tucker, 1923 gw soil echemus sp. 1 gw soil poecilochoa sp. 1 gw soil scotophaeus marleyi tucker, 1923 gw soil setaphis anchoralis purcell, 1908 gw soil setaphis arcus tucker, 1923 gw soil setaphis subtilis (simon, 1897) gw soil trachyzelotes jaxartensis (kroneberg, 1875) gw soil trephopoda hanoveria tucker, 1923 gw soil upognampa parvipalpa tucker, 1923 gw soil xerophaeus appendiculatus purcell, 1907 gw soil xerophaeus bicavus tucker, 1923 gw soil zelotes reduncus (purcell, 1907) gw soil zelotes tuckeri roewer, 1951 gw soil zelotes ungulus tucker, 1923 gw soil zelotes sp. 4 gw soil hersiliidae tyrotama soutpansbergensis foord & dippenaar-schoeman, 2005 rwb soil idiopidae ctenolophus fernoulheti hewitt, 1913 bd burrow soil idiops monticola (hewitt, 1916) bd burrow soil idiops sp. 1 bd burrow soil linyphiidae pelecopsis sp. 1 swb vegetation meioneta sp. 1 swb vegetation mecynidis sp. 1 swb vegetation microlinyphia sterilis (pavesi, 1883) swb vegetation lycosidae evippomma squamulatum (simon, 1898) gw soil geolycosa sp. 1 gw soil lycosa sp. 1 gw soil lycosa sp. 2 gw soil hogna sp. 1 gw soil hogna sp. 2 gw soil lycosidae sp. 1 (genus undetermined) gw soil lycosidae sp. 2 (genus undetermined) gw soil lycosidae sp. 3 (genus undetermined) gw soil lycosidae sp. 4 (genus undetermined) gw soil lycosidae sp. 5 (genus undetermined) gw soil lycosidae sp. 6 (genus undetermined) gw soil lycosidae sp. 7 (genus undetermined) gw soil pardosa leipoldti purcell, 1903 gw soil pardosa sp. 2 gw soil pardosa sp. 3 gw soil pardosa sp. 4 gw soil proevippa wanlessi (russell-smith, 1981) gw soil proevippa sp. 2 gw soil trabea purcelli roewer, 1951 gw soil zenonina albocaudata lawrence, 1952 gw soil mimetidae ero sp. 1* pw vegetation miturgidae cheiracanthium africanum lessert, 1921 pw vegetation cheiracanthium furculatum karsch, 1879 pw vegetation cheiracanthium vansoni lawrence, 1936 pw vegetation nemesiidae hermacha mazoena hewitt, 1915 bd burrow soil table 2 continued . . . 13 original research dippenaar, dippenaar-schoeman, modiba, khoza koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 10 17 family species guild field type nephilidae nephila senegalensis (walckenaer, 1842) owb vegetation oonopidae gamasomorpha humicola lawrence, 1947 gw soil oxyopidae hamataliwa fronticornis (lessert, 1927) pw grass hamataliwa kulczynskii (lessert, 1915) pw grass hamataliwa strandi caporriacco, 1939 pw grass oxyopes affinis lessert, 1915 pw grass oxyopes bedoti lessert, 1915 pw grass oxyopes bothai lessert, 1915 pw grass oxyopes hoggi lessert, 1915 pw grass oxyopes jacksoni lessert, 1915 pw grass oxyopes pallidecoloratus strand, 1906 pw grass oxyopes russoi caporiacco, 1940 pw grass oxyopes schenkeli lessert, 1927 pw grass oxyopes tuberculatus lessert, 1915 pw grass oxyopes sp. 10 pw grass oxyopes sp. 11 pw grass oxyopes sp. 12 pw grass peucetia viridis (blackwall, 1858) pw grass palpimanidae diaphorocellus biplagiatus simon, 1893 gw soil palpimanus armatus pocock, 1898 gw soil palpimanus transvaalicus simon, 1893 gw soil philodromidae ebo sp. 1* pw grass gephyrota sp. 1* pw grass hirriusa variegata (simon, 1895) pw grass philodromus browningi lawrence, 1952 pw grass philodromus grosi lessert, 1943 pw grass philodromus guineensis millot, 1942 pw grass suemus punctatus lawrence, 1938 pw grass thanatus dorsilineatus jézéquel, 1964 pw grass thanatus sp. 2 pw grass tibellus gerhardi van den berg & dippenaar-schoeman, 1994 pw grass tibellus hollidayi lawrence, 1952 pw grass tibellus minor lessert, 1919 pw grass pholcidae smeringopus atomarius simon, 1910 spwb soil pisauridae afropisaura sp. 1 pw vegetation euprosthenops australis simon, 1898 fwb vegetation euprosthenopsis vuattouxi blandin, 1977 fwb vegetation maypacius bilineatus (pavesi, 1895) pw vegetation maypacius stuhlmanni (bösenberg & lenz, 1895) pw vegetation rothus purpurissatus simon, 1898 pw vegetation prodidomidae theuma parva purcell, 1907 gw soil salticidae aelurillus sp. 1* gw soil baryphas ahenus simon, 1902 pw vegetation brancus bevisi lessert, 1925 pw vegetation cosmophasis sp. 1 gw soil dendryphantes sp. 1 pw vegetation dendryphantes sp. 2 pw vegetation euophrys sp. 1 pw vegetation heliophanus debilis simon, 1901 pw vegetation heliophanus demonstrativus wesolowska, 1986 pw vegetation heliophanus insperatus wesolowska, 1986 pw vegetation heliophanus transvaalicus simon, 1901 pw vegetation hyllus treleaveni peckham & peckham, 1902 pw vegetation hyllus sp. pw vegetation hyllus sp. 3 pw vegetation hyllus sp. 4 pw vegetation hyllus sp. 5 pw vegetation langelurillus sp. 1 gw soil table 2 continued . . . 14 spiders: arachinda, araneae original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 10 17 family species guild field type mogrus sp. 1 pw vegetation natta horizontalis karsch, 1879 gw soil pellenes sp. 1 pw vegetation phlegra sp. 1 gw soil pseudicius sp. 1 pw vegetation rhene machadoi berland & millot, 1941 pw vegetation stenaelurillus nigricaudus simon, 1886 gw soil stenaelurillus sp. 1 gw soil stenaelurillus sp. 2 gw soil stenaelurillus sp. 3 gw soil thyene inflata (gerstäcker, 1873) pw vegetation thyenula aurantiaca (simon, 1902) pw vegetation scytodidae scytodes sp. 1 gw soil segestriidae ariadna sp. 1 twb soil selenopidae anyphops sp. 1 pw tree selenops sp. 1 pw tree sparassidae olios sp. 1 pw vegetation olios sp. 2 pw vegetation olios sp. 3 pw vegetation palystes superciliosus l. koch, 1875 pw vegetation pseudomicrommata longipes (bösenberg & lenz, 1895) pw vegetation tetragnathidae leucauge decorata (blackwall, 1864) owb vegetation leucauge festiva (blackwall, 1866) owb vegetation theraphosidae augacephalus junodi (simon, 1904) bd burrow soil brachionopus pretoriae purcell, 1904 bd burrow soil harpactira sp. 1 bd burrow soil harpactirella sp. 1 bd burrow soil theridiidae argyrodes convivans lawrence, 1937 gwb webs other spiders argyrodes zonatus (walckenaer, 1842) gwb webs other spiders dipoena sp. 1 gwb vegetation enoplognatha molesta o.p.-cambridge, 1904 gwb vegetation latrodectus geometricus c.l.koch, 1841 gwb vegetation latrodectus renivulvatus dahl, 1902 gwb vegetation phoroncidia eburnea (simon, 1895) gwb vegetation steatoda sp. 1 gwb soil theridiidae sp. 1 (genus undetermined) gwb vegetation theridiidae sp. 2 (genus undetermined) gwb vegetation theridiidae sp. 3 (genus undetermined) gwb vegetation theridiidae sp. 4 (genus undetermined) gwb vegetation theridiidae sp. 5 (genus undetermined) gwb vegetation theridion purcelli o.p.-cambridge, 1904 gwb vegetation theridion sp. 2 gwb vegetation theridion sp. 3 gwb vegetation theridion sp. 4 gwb vegetation theridion sp. 5 gwb vegetation theridion sp. 6 gwb vegetation theridion sp. 7 gwb vegetation theridion sp. 8 gwb vegetation tidarren sp. 1 gwb vegetation thomisidae avelis hystriculus simon, 1895 pw vegetation camaricus nigrotesselatus simon, 1895 pw vegetation heriaeus crassispinus lawrence, 1942 pw grass heriaeus transvaalicus simon, 1895 pw grass hewittia gracilis lessert, 1928 pw grass misumenops rubrodecoratus millot, 1942 pw grass monaeses austrinus simon, 1910 pw tree monaeses fuscus dippenaar-schoeman, 1984 pw tree monaeses gibbus dippenaar-schoeman 1984 pw tree monaeses paradoxus (lucas, 1846) pw tree table 2 continued . . . 15 original research dippenaar, dippenaar-schoeman, modiba, khoza koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 10 17 family species guild field type monaeses pustulosus pavesi, 1895 pw tree monaeses quadrituberculatus lawrence, 1927 pw tree oxytate argenteooculata (simon, 1886) pw tree ozyptila sp. 1 pw soil pactactes trimaculatus simon, 1895 pw soil pherecydes tuberculatus o.p.-cambridge, 1883 pw tree pherecydes sp. 2 pw tree runcinia aethiops (simon, 1901) pw grass runcinia affinis simon, 1897 pw grass runcinia erythrina jézéquel, 1964 pw grass runcinia flavida (simon, 1881) pw grass stiphropus affinis lessert, 1923 pw soil synema decens (karsch, 1878) pw vegetation synema diana (audouin, 1826) pw vegetation synema imitator (pavesi, 1883) pw vegetation synema nigrotibiale lessert, 1919 pw vegetation thomisops sulcatus simon, 1895 pw vegetation thomisus blandus karsch, 1880 pw grass thomisus citrinellus simon, 1875 pw grass thomisus congoensis comellini, 1957 pw grass thomisus granulatus karsch, 1880 pw grass thomisus kalaharinus lawrence, 1936 pw grass thomisus scrupeus (simon, 1886) pw grass thomisus stenningi pocock, 1900 pw grass tmarus africanus lessert, 1919 pw grass tmarus cameliformis millot, 1942 pw grass tmarus cancellatus thorell, 1899 pw grass tmarus comellini garcia-neto, 1989 pw grass tmarus foliatus lessert, 1928 pw grass tmarus longicaudatus millot, 1941 pw grass tmarus sp. 7 pw grass xysticus fagei lessert, 1919 gw soil uloboridae miagrammopes longicaudus o.p.-cambridge, 1882 owb vegetation uloborus plumipes lucas, 1846 owb vegetation uloborus sp. 2 owb vegetation uloborus sp. 3 owb vegetation zodariidae capheris decorata simon, 1904 gw soil chariobas cylindraceus simon, 1893 gw soil cydrela sp. 1 gw soil diores auricula tucker, 1920 gw soil diores sp. 2 gw soil psammoduon sp. 1 gw soil ranops sp. 1 gw soil table 2 continued . . . other surveys undertaken in the savanna biome. in the makalali private game reserve, 4 832 spiders were caught over a year represented by 38 families and 268 species (whitmore et al. 2001); at lajuma in the western soutpansberg 127 species were caught that represented 46 families (foord et al. 2002); on sovenga hill in the polokwane district 793 spiders were caught representing 29 families and 76 species (modiba et al. 2005). in the kruger national park, 152 species from 40 families were recorded (dippenaar-schoeman & leroy 2003). the survey of the ndumo game reserve reported the highest spider diversity from any protected area in south africa so far with 46 families and 431 species (haddad et al. 2006). the most diverse families collected at the pnr were the thomisidae (42 spp.), representing 15.3% of the total number of species sampled, followed by the araneidae (39 spp.; 14.2%) and the salticidae (29 spp.; 10.5%) (see table 1). sixteen families were represented by a single species only. at roodeplaat dam nature reserve and lajuma, the thomisidae were also the most diverse with 25 and 15 species respectively, while at ndumu nature reserve, the salticidae were the most diverse with 82 species. the total spider diversity (275 spp.) represents 13.75% of the species of south african spiders (2 000 spp.), presently protected in the polokwane nature reserve. the vast majority of the spiders (191 spp.) collected were wanderers (69.5%), with web builders (84 spp.) contributing 30.5% (see table 2). this follows the reported pattern for species of the savanna biome. at ndumo game reserve, 75.9% were wanderers, with web builders contributing 24.1% of the species (haddad et al. 2006). at makalali private game reserve, the most abundant spiders collected were also wanderers (59%), with web builders contributing 41% (whitmore et al. 2002), while on sovenga hill, 64.9% of species were wanderers while the web builders only contributed 35.1% (modiba et al. 2005). similar results were found in the western soutpansberg with 64% of collected species being wanderers and 36% web builders (foord et al. 2002). most of the mygalomorphae spiders are protected due to the pet trade and it is important that species are protected in reserves. according to dippenaar-schoeman (2002b), there are 14 species of mygalomorphae occurring in the limpopo 16 spiders: arachinda, araneae original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 10 17 of facilities and equipment; ms b.p. jordaan (department of biodiversity) for spending a lot of time assisting and organising to accommodate our needs in order to successfully complete the study; ms a. van den berg (arc plant protection research institute) for her assistance with the family identification of specimens; and mr c. haddad (university of the free state) for the identification of the corinnidae specimens. references dippenaar-schoeman, a.s. 2002a. status of south african arachnida fauna. proceedings of the symposium on the status of south african species organised by the endangered wildlife trust (ewt) of south africa. rosebank. 4 7 september 2002. dippenaar-schoeman, a.s. 2002b. baboon and trapdoor spiders of southern africa: an identification manual. plant protection research institute handbook no. 13. pretoria, agricultural research council. dippenaar-schoeman, a.s. & craemer, c. 2000. the south african national survey of arachnida (sansa). plant protection news 56: 11–12. dippenaar-schoeman, a.s. & jocqué, r. 1997. african spiders, an identification manual. plant protection research institute handbook no. 9. pretoria, agricultural research council. dippenaar-schoeman, a.s. & leroy, a. 2003. a check list of the spiders of the kruger national park, south africa (arachnida: araneae). koedoe 46: 91–100. dippenaar-schoeman, a.s., van den berg, a. & van den berg, a.m. 1989. species composition and relative seasonal abundance of spiders from the field and tree layers of the roodeplaat dam nature reserve. koedoe 32: 25–38. foord, s.h., dippenaar-schoeman, a.s. & van der merwe, m. 2002. a checklist of the spider fauna of the western soutpansberg, south africa (arachnida: araneae). koedoe 45: 35–43. haddad, c.r., dippenaar-schoeman, a.s. & wesolowska, w. 2006. a checklist of the non-acarine arachnids (chelicerate: arachnida) of the ndumo game reserve, maputaland, south africa. koedoe 49: 1–22. low, a.b. & rebelo, a.g. (eds). 1996. vegetation of south africa, lesotho and swaziland. pretoria, department of environmental affairs and tourism. myers, n. 1990. the biodiversity challenge: expanded hot-spot analysis. the environmentalist 10: 243–255. modiba, m.a., dippenaar, s.m. & dippenaar-schoeman, a.s. 2005. a checklist of spiders from sovenga hill, an inselberg in the savanna biome, limpopo province, south africa (arachnida: araneae). koedoe 48: 109–115. mucina, l. & rutherford, m.c. (eds). 2006. the vegetation of south africa, lesotho and swaziland. strelitzia 19. pretoria, south african biodiversity institute. mucina, l., rutherford, m.c. & powrie, l.w. 2005. vegetation map of south africa, lesotho and swaziland. south africa national biodiversity institute (sanbi), kirstenbosch research centre. whitmore, c., slotow, r., crouch, t.e. & dippenaar-schoeman, a.s. 2001. checklist of spiders (araneae) from a savanna ecosystem, northern province, south africa: including a new family record. durban museum novitates 26: 10–19. whitmore, c., slotow, r., crouch, t.e. & dippenaar-schoeman, a.s. 2002. diversity of spiders (araneae) in a savanna reserve, northern province, south africa. journal of arachnology 30: 344–356. family species cyrtaucheniidae ancylotrypa brevipalpis (hewitt, 1916) idiopidae idiops sp. ctenolophus fernoulheti hewitt, 1913 segregara monticola (hewitt, 1916) nemesiidae hermacha mazoena hewitt, 1915 theraphosidae augacephalus junodi (simon, 1904) brachionopus pretoriae purcell, 1904 harpactira sp. harpactirella sp. table 3 checklist of the mygalomorphs of the polokwane nature reserve, limpopo province, south africa province. only one of the previously recorded species (i.e. augacephalus junodi (simon, 1904)) was encountered during the study period. however, eight additional mygalomorph species (see table 3) have been collected: ancylotrypa brevipalpis (hewitt, 1916), brachionopus pretoriae purcell, 1904, ctenolophus fenoulheti hewitt, 1913, segregara monticola (hewitt, 1916), hermacha mazoena hewitt, 1915, idiops sp., harpactira sp. and harpactirella sp. all of the above-mentioned species are burrow dwelling. these species are all new geographical records for the province. conclusion considering the duration of the current study (one year) and the large number of specimens caught (13 821), it may be expected that the study provides a good representation of families occurring in the polokwane nature reserve and that an extension of the study will probably increase the number of genera and species, but will probably not increase the number of families significantly. all the reported families and species are new geographical records for the polokwane nature reserve. the large number of species (97) that could not be identified emphasises the taxonomic impediment of invertebrates in south africa. during this study, nine possible new species were collected, and several genera were collected in south africa for the first time. additionally, the study provided new information on the distribution of all species concerned and provides material that can be used in future taxonomic work. this highlights the important contribution that the sansa initiative makes to a better knowledge of the diversity of arachnids in south africa, especially in conserved areas. acknowledgements we thank the nrf (thuthuka programme) and the university of limpopo (research development and administration) for financial aid that supported this work; the polokwane municipality for allowing the study on their property; the rotary club (polokwane) for allowing two of the authors to stay at the education centre for the duration of field work and for supplying additional accommodation together with wood worx; die drift instant lawn for sponsorship of a bicycle trailer and saloojee cycle works for discount on bicycle repairs. additionally we thank the university of limpopo for provision 17 article information authors: michelle j. tedder1 kevin p. kirkman1 craig d. morris1 winston s.w. trollope2 mpaphi c. bonyongo3 affiliations: 1school of life sciences, university of kwazulu-natal, south africa2working on fire international, nelspruit, south africa 3okavango research institute, university of botswana, botswana correspondence to: michelle tedder postal address: private bag x01, pietermaritzburg 3209, south africa dates: received: 15 july 2012 accepted: 11 june 2013 published: 31 july 2013 how to cite this article: tedder, m.j., kirkman, k.p., morris, c.d., trollope, w.s.w. & bonyongo, m.c., 2013, ‘classification and mapping of the composition and structure of dry woodland and savanna in the eastern okavango delta’, koedoe 55(1), art. #1100, 8 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1100 copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. classification and mapping of the composition and structure of dry woodland and savanna in the eastern okavango delta in this original research... open access • abstract • introduction • research method and design    • setting    • sampling procedure    • data analysis • results    • tall mopane woodland (group b)    • tall mixed mopane woodland (group c)    • short mixed mopane woodland (group d)    • short mixed broadleaf woodland (group e)    • open mixed acacia woodland (group f)    • closed acacia–combretum woodland (group g)    • closed mixed acacia veld (group h)    • closed acacia erioloba savanna (group i)    • tall mixed broadleaf woodland (group j)    • open acacia erioloba savanna (group k)    • palm thornveld (group l) • discussion • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ the dry woodland and savanna regions of the okavango delta form a transition zone between the okavango swamps and the kalahari desert and have been largely overlooked in terms of vegetation classification and mapping. this study focused on the species composition and height structure of this vegetation, with the aim of identifying vegetation classes and providing a vegetation map accompanied by quantitative data. two hundred and fifty-six plots (50 m × 50 m) were sampled and species cover abundance, total cover and structural composition were recorded. the plots were classified using agglomerative, hierarchical cluster analysis using group means and bray-curtis similarity and groups described using indicator species analysis. in total, 23 woody species and 28 grass species were recorded. acacia erioloba and colophospermum mopane were the most common woody species, whilst urochloa mossambicensis, panicum maximum, dactyloctenium gigantium and eragrostis lehmanniana were the most widespread grasses. eleven vegetation types were identified, with the most widespread being short mixed mopane woodland, tall mopane woodland and tall mixed mopane woodland, covering 288.73 km2 (28%), 209.14 km2 (20%) and 173.30 km2 (17%) of the area, respectively. despite their extensive area, these three vegetation types were the least species-rich, whilst palm thornveld, short mixed broadleaf woodland and open mixed acacia woodland were the most taxonomically variable. by contrast, closed mixed acacia woodland and closed acacia–combretum woodland had the most limited distribution, accounting for less than 1% of the mapped area each. conservation implications: the dry woodland and savanna vegetation of the okavango delta comprises a much wider suite of plant communities than the acacia-dominated and mopane-dominated classifications often used. this classification provided a more detailed understanding of this vegetation and essential background information for monitoring, management and research. introduction top ↑ de cáceres and wiser (2011:387) state that, ‘vegetation classification is a useful tool for basic and applied research as well as for environmental management.’ the major goal when conducting such an exercise is to simplify a complex continuous pattern of change in vegetation composition and structure across the landscape in order to understand how species are distributed and why, as well as to predict how species, communities and landscapes will respond to environmental changes (symstad 2002). as a result of the varied applications of vegetation classification, the properties utilised in their construction are equally varied (de cáceres & wiser 2011; symstad 2002). the more commonly used properties, listed by symstad (2002) are: structure, taxonomy, resource use, response to change and role in ecosystem function. vegetation classification becomes an even more powerful research and management tool when combined with some form of mapping indicating the distribution and spatial coverage of the groupings produced.current vegetation mapping practices are strongly influenced by the availability of remote-sensing tools (bredenkamp et al. 1998). over reliance on these remote sensing tools may, however, result in map inaccuracies because poor resolution of the available predictor variable maps and low levels of correlation between predictor variables and vegetation units may affect map accuracy (van etten 1998). but remote sensing can be an extremely powerful tool if used in conjunction with appropriate ground truthing (bredenkamp et al. 1998). one of the more efficient methods for the construction of multipurpose vegetation maps is a floristic-based classification, supported by multivariate analysis and structural data (dias, elias & nunes 2004). this allows the classification to be used to investigate both long-term change indicated by the floristic composition and short-term change indicated by the structural composition (lux & bemmerlein-lux 1998). vegetation maps and descriptions can also be used for a range of purposes such as: landscape planning, development and implementation of management plans, forestry, conservation, monitoring and many varied forms of research (bredenkamp et al. 1998; dias et al. 2004). although a vegetation map covering the whole of ngamiland (110 000 km2), the north-western region of botswana, was produced in 2002 (jellema, ringrose & matheson 2002), it has limited applicability to research within the okavango delta as the delta itself only comprises 26% of the mapped area (ramberg et al. 2006). several other maps focused on the peripheral dryland regions of the okavango delta (mccarthy, gumbricht & mccarthy 2005; ringrose, van der post & matheson 2003; rűbbel & child 1976; tinley 1966), referring to woody cover at a finer scale than the broad division of acacia-dominated woodland and tall and short c. mopane-dominated woodlands (ellery & ellery 1997). the objectives of this study were to, firstly, taxonomically and structurally classify the vegetation of the dry woodland and savanna regions in the eastern okavango delta. secondly, they aimed to provide a broad description of those vegetation types in terms of their, (1) woody species composition, (2) mean canopy cover, (3) indicator woody species, (4) diversity, (5) height structure of the woody species layer (based on the relative cover of pre-defined tree height classes) and (6) composition of the most prominent grasses. thirdly, was to produce a vegetation map that provides background data for the ongoing research in the eastern okavango delta. research method and design top ↑ setting the study was conducted in the southern okavango delta, botswana, in the controlled hunting areas ng33 and ng34 and a portion of moremi game reserve (ng28) (figure 1). figure 1: location of the study area (ng33, ng34 and moremi game reserve) in the okavango delta, north-western botswana. these three sections of the study area cover an area of 60.55 km2, 851.68 km2 and 579.59 km2, respectively. these dry woodland regions in the south-eastern okavango delta function as a transition zone between the okavango swamps and the kalahari desert, resulting in a region of high species richness and endemism (mazizi resources pty ltd 2009). the selected area is predominantly dry woodland and savanna and provides good representations of both acacia-dominated savanna and mopane-dominated woodland. the area experiences a dry winter season from april to september, with minimum temperatures dropping to 15 °c, and a warm summer from october to march, with maximum temperatures rising to 40 °c (mazizi resources pty ltd 2009). the mean annual rainfall is 560 mm, with the majority falling between november and march (mazizi resources pty ltd 2009). the area provides habitats for a wide range of mammal, reptile and bird species, many of which are rare and protected (mazizi resources pty ltd 2009). sampling procedure the composition and structure of the dryland woody-dominated vegetation in ng33 and ng34 was sampled between february and may 2009 using 256 plots, each 50 m × 50 m in size. this plot size is considered to be suitable for mapping vegetation on a landscape scale as it minimises standard error over sampling cost (xiao et al. 2004). sample sites were systematically located along existing tracks, with the minimum distance between any two sites being 1 km, to cover the study area as extensively as possible and to facilitate access.at each sample plot, all woody species were recorded and each species was allocated an absolute cover abundance value (goodall 1980). this was achieved by estimating the percentage of the whole site cover occupied by each species separately. in addition, the total woody cover was visually estimated and gps coordinates were recorded for each plot. the structural composition of the woody layer was sampled by visually estimating the absolute cover of each of the following height classes, < 0.50 m, 0.50 m – 0.99 m, 1.00 m – 1.99 m, 2.00 m – 2.99 m and > 3.00 m. absolute cover (aerial) of each grass species in the understory was also estimated and later relativised so that total plot cover equalled 100% to describe the grass species composition. identification was performed to species level and nomenclature was according to coates-palgrave (2002), roodt (1998) and van oudtshoorn (1999) and names were confirmed as current by the south african national biodiversity institute (accessed through the sibis portal, http://sibis.sanbi.org on 05 april 2013). data analysis the woody species absolute cover data were subjected to an agglomerative, hierarchical cluster analysis via the software program primer (clarke & gorley 2006), using the group mean clustering method and bray-curtis similarity to create groups. a similarity profile test (simprof), also conducted in primer (clarke & gorley 2006), was used to identify sub-groups for which species composition was not statistically distinguishable (p ≥ 0.05, n = 10 000 permutations) and thus could be retained as a single branch of the dendrogram. the woody layer within each vegetation type was described in terms of species composition, based on mean cover abundance of each species in that vegetation group, and indicator species. in addition, mean woody canopy cover and species richness were calculated for each vegetation type. vegetation classes with a mean woody canopy cover of greater than 50% were considered to have a closed canopy and those with a mean woody canopy cover of less than 50% were described as open canopy. indicator species for each vegetation type were identified using indicator species analysis, conducted in pc-ord (mccune & mefford 1997) using 9999 permutations in a monte carlo test for indicator values (mccune & mefford 1997). diversity within the vegetation types was described using mean hill’s diversity indices and mean pielou’s evenness co-efficient (begon, townsend & harper 2005) using the following formulae: hill’s n1 = eh’ [eqn 1] where h’ = -σpi.ln(pi) and pi = the proportion of species i in the sample. hill’s n2 = 1/λ [eqn 2] where λ = σni(ni – 1)/n(n-1), ni = number of individuals of species i in the sample and n = total number of individuals in the sample. pielou’s evenness co-efficient = h’/lns [eqn 3] where s = total number of species in the sample. hill’s diversity numbers n1 and n2 describe the diversity of abundant and very abundant species respectively, whilst pielou’s evenness co-efficient ranges from 0 to 1 with low values indicating dominance by a few species (begon et al. 2005). differences in height structure between the vegetation types were examined using an analysis of similarity (anosim) in primer based on bray-curtis similarity and tested with 9999 permutations (clarke & gorley 2006). the p-values were controlled for false discovery rate using the benjamani and hochberg procedure (verhoeven, simonsen & mcintyre 2005). a similarity percentages routine (simper), also conducted in primer (clarke & gorley 2006) was then used to determine the contribution of each height class to the difference in bray-curtis similarity between vegetation types. vegetation was considered to be tall if more than 35% of the woody layer fell into the height class > 3 m, whilst it was considered to be short if less than 30% of the woody component fell into this height class. the vegetation types were named according to their dominant species, height structure and level of woody cover. to provide a complete overview of the vegetation types, the mean abundance for each grass species occurring in each vegetation type was calculated. the vegetation types were then mapped using arcmap 10 (esri 2010) by overlaying the categorised sample sites on aerial photographs obtained from the okavango research institute. results top ↑ twenty-three woody species belonging to 16 genera and 28 grass species belonging to 19 genera were identified in the study area. the cluster analysis and simprof test produced 18 groups, which were significantly different from one another (p < 0.05). these groups were examined and those with a mean similarity of greater than 70% and comparable woody cover and species richness were combined. this resulted in 12 final groups, of which one (with two sites) was subsequently excluded from the woody classification because it had less than 5% woody cover (group a) (figure 2). the vegetation types were then named according to the dominant species, and then further differentiated according to their dominant height classes (tall or short) and percentage canopy cover (open or closed). if there were numerous co-dominant species present, then the term mixed was used. the 11 vegetation types derived from this classification are as follows (figure 2): figure 2: dendrogram of an agglomerative cluster analysis (based on braycurtis similarity measure) of canopy cover composition of woody species in the eastern okavango delta, botswana. • group b: tall mopane woodland. • group c: tall mixed mopane woodland. • group d: short mixed mopane woodland. • group e: short mixed broadleaf woodland. • group f: open mixed acacia veld. • group g: closed acacia–combretum woodland. • group h: closed mixed acacia veld. • group i: closed acacia erioloba savanna. • group j: tall mixed broadleaf woodland. • group k: open acacia erioloba savanna. • group l: palm thornveld. tall mopane woodland (group b) tall mopane woodland almost solely comprised the indicator species c. mopane (p < 0.001) (table 1) and was the densest of all 11 woody vegetation types, with a mean woody cover of 92.0% (± 0.9%) (table 2). the majority of this cover was taller than 3 m (table 4). tall mopane woodland covered an area of 209 km2 (table 2), occurring mainly in the northern and eastern regions of ng34 and continuing north into moremi game reserve (figure 3). it had a low mean species richness (1.90 ± 0.14) (table 2), with a small proportion of the woody cover comprising acacia, combretum and grewia species growing in the understory (table 1). the grass layer was fairly diverse, comprising 18 species, but was dominated by u. mossambicensis (25.0%) and eragrostis rigidior (19.0%) (table 3). table 1: mean estimated relative cover (%) of woody species for the vegetation types of ng33 and ng34 in the eastern okavango delta. table 2: number of sites, hill’s diversity indices, mean (± standard error) species richness, maximum species richness and mean (± standard error) woody cover (%) for the vegetation types of ng33 and ng34 in the eastern okavango delta. table 3: mean estimated relative cover (%) of grass species in the vegetation types of ng33 and ng34 in the eastern okavango delta. table 4: mean (± standard error) relative cover (%) of height classes and mean within group similarity calculated using analysis of similarities of the vegetation types in ng33 and ng34. figure 3: distribution of vegetation types in ng33 and 34 in the eastern okavango delta, botswana. tall mixed mopane woodland (group c) tall mixed mopane woodland was dominated by the indicator species c. mopane (p < 0.001) (table 1), with some a. erioloba occurring in more open patches. tall mixed mopane woodland had a mean woody cover, predominantly taller than 3 m (table 4), of 75.0% (± 3.4%) (table 2) and a mean species richness of 3.30 (± 0.32) (table 2). other species contributing to the understory cover were acacia hebeclada, acacia luederitzii, boscia albitrunca, combretum species and terminalia sericea (table 1). the grass layer comprised 14 species with u. mossambicensis (21.0%) and p. maximum (19.0%) dominating (table 3). tall mixed mopane woodland, covering 173 km2, occurred mainly in the central and eastern regions of ng34 along the edges of the tall mopane woodland (figure 3) (table 2). short mixed mopane woodland (group d) short mixed mopane woodland had the lowest cover of the three mopane-dominated vegetation types, with a mean woody cover of 66.0% (± 3.1%) (table 2). this cover was spread evenly between the three tallest height classes (table 4). colophospermum mopane comprised approximately one-third of the woody cover, with the remainder a mix of acacia species, b. albitrunca, grewia species, philenoptera nelsii and t. sericea (table 1), resulting in a mean species richness of 5.20 (± 0.32) (table 2). the grass layer predominantly comprised e. lehmanniana (17.0%), d. gigantium (16.0%), p. maximum (14.0%), pogonarthria squarrosa (13.0%) and u. mossambicensis (12.0%), with occasional occurrences of eight other species (table 3). short mixed mopane woodland, the most widespread vegetation type covering 288 km2, occurred in large patches in ng34 south of the boundary of ng33 and east of the gomoti river and as small patches on the boundaries of tall mopane woodland and tall mixed mopane woodland in the central and northern sections of ng34 (figure 3). short mixed broadleaf woodland (group e) short mixed broadleaf woodland covered an area of 43 km2 and was characterised by the indicator species p. nelsii (p = 0.034), t. sericea (p = 0.003) and dichrostachys cinerea (p = 0.030) (table 1). short mixed broadleaf woodland had a mean woody cover of 66.0% (± 3.2%) (table 2), which was generally shorter than 3 m in height (table 4) and contained small proportions of acacia species, b. albitrunca and c. mopane (table 1), resulting in a mean species richness of 5.0 (± 0.3) (table 2). eragrostis lehmanniana (18.0%), stipagrostis hirtigluma (15.0%) and p. maximum (14.0%) dominated the grass layer, with limited cover of 11 other species (table 3). short mixed broadleaf woodland occurred mainly between the southern tip of ng33 and the southern buffalo fence, with some patches occurring east of the boundary of ng33 and along the cutline between ng33 and moremi game reserve (figure 3). open mixed acacia woodland (group f) open mixed acacia woodland was characterised by the presence of widely scattered individuals of the indicator species a. hebeclada (p = 0.001) and a. luederitzii (p = 0.009) growing with a. erioloba. other species that were present but with low canopy cover include boscia mosambicensis, c. mopane, combretum imberbe, croton megalobotrys, diospyros lycioides and p. nelsii (table 1). open mixed acacia woodland had a mean canopy cover of 41.0% (± 4.9%) (table 2), the majority of which was taller than 2 m (table 4) and a mean species richness of 5.20 (± 0.57) (table 2). the grass layer comprised 14 species with u. mossambicensis (24.0%) and e. lehmanniana (17.0%) being dominant (table 3). open mixed acacia woodland covered an area of 14 km2 (table 2) and occurred mainly along the northern boundary of ng33 and ng34, in small patches in the central and eastern regions of ng34 and east of the gomoti river in moremi game reserve, north of the ng34 boundary (figure 3). closed acacia–combretum woodland (group g) closed acacia–combretum woodland had a mean woody cover of 67.0% (± 14.5%) (table 2), a mean species richness of 5.70 (± 0.33) (table 2) and was dominated by a. erioloba and the indicator species c. imberbe (p = 0.001) (table 1). acacia hebeclada, b. mosambicensis, c. mopane, c. megalobotrys, gymnosporia senegalensis, hyphene petersiana and searsia tenuinervis contributed a small proportion to the canopy cover (table 1). the majority of the woody cover was between 1 m and 3 m tall (table 4). the grass layer comprised 11 species, with u. mossambicensis (32.0%) being dominant and e. lehmanniana (15.0%) and p. maximum (15.0%) being sub-dominant (table 3). closed acacia–combretum woodland covered an area of approximately 1 km2 and occurred in small patches in central and western ng34 (figure 3). closed mixed acacia veld (group h) closed mixed acacia veld had a mean canopy cover of 73.0% (± 22.5%) and was characterised by the presence of the indicator species acacia nigrescens (p < 0.001), a. hebeclada (p = 0.001) and b. albitrunca (p = 0.013) growing with a. erioloba (table 1). the woody cover was mostly between 1 m and 3 m tall and had a mean species richness of 7.0 (± 1.0) (table 2). the grass layer was dominated by e. lehmanniana (25.0%) and u. mossambicensis (25.0%), with d. giganteum (15.0%) and p. maximum (15.0%) being sub-dominant and four other species occurring occasionally (table 3). closed mixed acacia woodland occurred in small patches in south-western ng34 close to the gomoti river (figure 3), totalling an area of 7 km2 (table 2). closed acacia erioloba savanna (group i) closed a. erioloba savanna was dominated by tall a. erioloba trees (table 1), the indicator species for this structural vegetation type (p = 0.009), with low cover of a. hebeclada, c. mopane, c. imberbe, c. megalobotrys, p. nelsii and ziziphus mucronata in the understory (table 1). closed a. erioloba savanna had a mean woody cover of 50.0% (± 1.8%) (table 2), with half the woody cover taller than 3 m (table 4), and a mean species richness of 4.20 (± 0.24) (table 2). the grass layer comprised 14 species, with d. giganteum (17.0%) and digitaria eriantha (17.0%) dominating a. erioloba savanna covered an area of 73 km2 (table 2) and occurred in large patches in central ng33, in ng34 south and east of the ng33 boundary and in smaller patches between the c. mopane-dominated belts in eastern ng34 (figure 3). tall mixed broadleaf woodland (group j) tall mixed broadleaf woodland is characterised by a combination of tall a. erioloba with an understory of c. megalobotrys, the indicator species for this vegetation type (p < 0.001) (table 1). a small proportion of the woody cover also comprised c. mopane, c. imberbe, g. senegalensis, p. nelsii, s. tenuinervis and z. mucronata (table 1), resulting in a mean species richness of 5.90 (± 0.64) (table 2). the mean woody cover in tall mixed broadleaf woodland was 66.0% (± 4.5%) (table 2), with 70.0% of this being taller than 2 m (table 4). the grass layer comprised 15 species, but was dominated by p. maximum (22.0%), d. giganteum (19.0%) and u. mossambicensis (19.0%) (table 3). the 30 km2 of tall mixed broadleaf woodland (table 2) occurred in scattered patches in ng34, mainly east of the ng33 boundary and in central and western ng33 along the moremi game reserve boundary (figure 3). open acacia erioloba savanna (group k) open a. erioloba savanna was characterised by an open canopy (23.0% ± 12.7%) (table 2), with a scattered distribution of tall a. erioloba (table 1) as an indicator species (p = 0.009). other species present included c. mopane, p. nelsii, t. sericea, a. hebeclada, b. albitrunca, c. imberbe, h. petersiana and z. mucronata (table 1), resulting in a mean species richness of 4.40 (± 1.98) (table 2). the woody cover was predominantly taller than 1 m, with a fairly even spread between the three tallest height classes (table 4). the grass was dominated by d. giganteum (20.0%) and u. mossambicensis (17.0%), with 14 other species occurring occasionally (table 3). open a. erioloba savanna occurred in two main patches: in ng34 south of the boundary of ng33 and moremi game reserve and in central ng34 between the c. mopane-dominated belt to the north and the closed a. erioloba savanna to the south (figure 3), totalling an area of 122 km2 (table 2). palm thornveld (group l) palm thornveld had an open canopy (36.0% ± 4.3%) (table 2) and was characterised by a sparse cover of z. mucronata and the indicator species h. petersiana (p = 0.017), with some occurrences of a. erioloba, a. hebeclada, a. nigrescens, c. megalobotrys, d. lycioides, p. nelsii and c. imberbe (table 1), resulting in a mean species richness of 4.40 (± 0.34) (table 2). the majority of the woody cover was between 2 m and 3 m tall. the grass layer comprised 15 species, with cynodon dactylon (27.0%) dominating (table 3). the 60 km2 area (table 2) of palm thornveld was limited to the old floodplain regions south of ng33 and east of the gomoti river (figure 3). discussion top ↑ overall, the structure of the woody vegetation in the eastern okavango delta was short (< 3 m) according the broad-scale classification guidelines stated by edwards (1983). the two most widespread woody species occurring in varying densities in every vegetation type were a. erioloba and c. mopane, whilst the most widespread grass species were u. mossambicensis, p. maximum, d. gigantium and e. lehmanniana. the three c. mopane-dominated vegetation types accounted for approximately half the woody cover in ng33 and ng34. the structural differences observed between the tall and short c. mopane vegetation may be caused by several factors. nutrient poor, sandy soils (roodt 1998) and the associated decrease in available moisture can result in stunted c. mopane trees (february et al. 2007), whilst disturbance factors such as fire, wood harvesting and extensive utilisation by elephants result in multi-stemmed coppice growth (lombard 2003; van voorthuizen 1976) and may cause the development of short mixed mopane woodland in areas where soil conditions allow for tall mopane woodland. by contrast, tall stands of c. mopane trees tend to occur on nutrient-rich alluvial soils (roodt 1998) and, although slow-growing initially, they can reach heights of up to 30 m (venter & venter 2002). there is great variation in the number of woody vegetation classes previously described by the four vegetation studies conducted in the dryland regions of the okavango delta between 1966 and 2005. tinley (1966) describes eight vegetation classes predominantly based on species composition but including structure in some descriptions, whilst rübbel and child (1976) described five, distinguished mainly by flooding frequency and dominant woody species. ringrose et al. (2003) described twelve ecological units in total, of which seven are woody dominated, whilst mccarthy et al. (2005) described only four. both ringrose et al. (2003) and mccarthy et al. (2005) base their descriptions on plant species composition. of these four studies, the one which is most closely correlated to the eleven vegetation types described here is the oldest classification, conducted by tinley (1966) in moremi game reserve. seven of the eight vegetation types described by tinley (1966) can be matched to groups produced by this classification, whilst the same can be said for only five of the seven groups described by ringrose et al. (2003), who mapped vegetation in the xudum region of the south-western okavango delta. ringrose et al. (2003) do, however, provide a slightly more detailed description of the species composition within the c. mopane-dominated vegetation types, describing three distinct groups, whilst tinley (1966) describes only two forms of c. mopane woodland focusing on structural composition. the absence of any broadleaf-dominated communities in the ecoregions described by ringrose et al. (2003) and the limited description given by tinley (1966) emphasises the level of variation observed in the dry woodland and savanna vegetation in different regions of the okavango delta and also the importance of using detailed field-based vegetation studies in conjunction with remote-sensing techniques. conclusion top ↑ this study highlights the broad suite of vegetation communities present in the dry woodland and savanna vegetation of the eastern okavango delta. the vegetation classification and map produced by this study not only provides valuable geo-referenced field data to augment the delta-wide maps produced in 2002 (ramberg et al. 2006), but also important background data for both the habitat-based monitoring projects to be conducted in ng33 and ng34 and the carnivore and herbivore research being conducted in the area. acknowledgements top ↑ we would like to thank mr r.g. fuhr for his generous financial support for the duration of this project and the staff and fellow researchers of dubatana camp and the okavango research institute for their assistance and support in the field. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions all field work was conducted by m.j.t. (university of kwazulu-natal) as part of a phd project. k.p.k. (university of kwazulu-natal) was the main supervisor and c.d.m. (university of kwazulu-natal), w.s.w.t. (working on fire international) and m.c.b. (university of botswana) were co-supervisors. m.j.t., k.p.k. and w.s.w.t. were responsible for project design. m.c.b. assisted in acquiring the necessary permits and permissions for fieldwork. m.j.t. and c.d.m. conducted data analysis and m.j.t. wrote the manuscript which was commented on and edited by all four co-authors. references top ↑ begon, m., townsend, c. & harper, j., 2005, ecology: from individuals to ecosystems, blackwell scientific publications, oxford.bredenkamp, g., chytery, m., fischer, h., neuhauslova, z. & van der maarel, e., 1998, ‘vegetation mapping: theory, methods and case studies: introduction’, applied vegetation science 1, 161–164. http://dx.doi.org/10.1111/j.1654-109x.1998.tb00397.x clarke, k. & gorley, r., 2006, primer, primer-e ltd, ivybridge. coates-palgrave, m., 2002, keith coates-palgrave trees of southern africa, 3rd edn., struik publishers, cape town. de cáceres, m. & wiser, s., 2011, ‘towards consistency in vegetation classification’, journal of vegetation science 23, 387–393. http://dx.doi.org/10.1111/j.1654-1103.2011.01354.x dias, e., elias, r. & nunes, v., 2004, ‘vegetation mapping and nature conservation: a case study in terceira island (azores)’, biodiversity and conservation 13, 1519–1539. http://dx.doi.org/10.1023/b:bioc.0000021326.50170.66 edwards, d., 1983, ‘a broad-scale structural classification of vegetation for practical purposes’, bothalia 14, 705–712. ellery, k. & ellery, f., 1997, ‘plants of the okavango delta’, tsaro publishers, durban. pmid:15093481 esri, 2010, arcmap version 10, esri, redlands. february, e., higgins, s., newton, r. & west, a, 2007, ‘tree distribution on a steep environmental gradient in an arid savanna’, journal of biogeography 34, 270–278. http://dx.doi.org/10.1111/j.1365-2699.2006.01583.x goodall, d.w., 1980, ‘numerical methods of classification’, classification of plant communities 5(1), 247–286. jellema, a., ringrose, s. & matheson, w., 2002, ‘northern botswana vegetation mapping project’, harry oppenheimer okavango research centre, university of botswana, maun. lombard, p., 2003, ‘the long-term effects of fire frequency and season on the colophospermum mopane shrubveld of the kruger national park’, magister technologiae thesis, faculty of the built environment, port elizabeth technikon. lux, a. & bemmerlein-lux, f., 1998, ‘two vegetation maps of the same island: floristic units versus structural units’, applied vegetation science 1, 201–210. http://dx.doi.org/10.2307/1478949 mazizi resources pty ltd, 2009, ‘management plan for ng33 and ng34’, mazizi resources pty ltd, maun. mccarthy, j., gumbricht, t. & mccarthy, t., 2005, ‘ecoregion classification in the okavango delta, botswana from multitemporal remote sensing’, international journal of remote sensing 26, 4339–4357. http://dx.doi.org/10.1080/01431160500113583 mccune, b. & mefford, m., 1997, pc-ord multivariate analysis of ecological data version 6, mjm software design, gleneden beach. ramberg, l., hancock, p., lindholm, m., meyer, t., ringrose, s., sliva, j. et al., 2006, ‘species diversity of the okavango delta, botswana’, aquatic sciences 68, 310–337. http://dx.doi.org/10.1007/s00027-006-0857-y ringrose, s., van der post, c. & matheson, w., 2003, ‘mapping ecological conditions in the okavango delta, botswana using fine and coarse resolution systems including simulated spot vegetation imagery’, international journal of remote sensing 24, 1029–1052. http://dx.doi.org/10.1080/01431160210155046 roodt, v., 1998, the shell field guide series: part i – trees and shrubs of the okavango delta: medicinal uses and nutritional value, shell oil botswana (pty) ltd, gaberone. rűbbel, h. & child, g., 1976, ‘notes on the ecology of the moremi wildlife reserve’, department of wildlife, national parks and tourism, botswana. symstad, a., 2002, ‘an overview of ecological plant classification systems’, in r. ambasht & n. ambasht (eds.), modern trends in terrestrial ecology, pp. 13–50, kluwer academic, new york. http://dx.doi.org/10.1007/978-1-4615-0223-4_2 tinley, k.l., 1966, ‘an ecological reconnaissance of the moremi wildlife reserve, northern okavango swamps, botswana’, the okavango wildlife society, johannesburg. van etten, e., 1998, ‘mapping vegetation in an arid, mountainous region of western australia’, applied vegetation science 1, 189–200. http://dx.doi.org/10.2307/1478948 van oudtshoorn, f., 1999, guide to grasses of southern africa, briza, pretoria. van voorthuizen, e., 1976, ‘the mopane tree’, botswana notes and records 8, 223–230. venter, f. & venter, j., 2002, making the most of indigenous trees, 2nd edn., briza publications, pretoria. verhoeven, k.j.f., simonsen, k.l. & mcintyre, l.m., 2005, ‘implementing false discovery rate control: increasing your power’, oikos 108, 643–647. http://dx.doi.org/10.1111/j.0030-1299.2005.13727.x xiao, x., gertner, g., wang, g. & anderson, a.b., 2004, ‘optimal sampling scheme for estimation landscape mapping of vegetation cover’, landscape ecology 20, 375–387. http://dx.doi.org/10.1007/s10980-004-3161-z article information authors: ben j. strohbach1 willem j. jankowitz2 affiliations: 1national botanical research institute, windhoek, namibia2school of natural resources and tourism, polytechnic of namibia, namibia correspondence to: ben strohbach postal address: private bag 13184, windhoek, namibia dates: received: 14 dec. 2010 accepted: 16 sept. 2011 published: 29 mar. 2012 how to cite this article: strohbach, b.j. & jankowitz, w.j., 2012, ‘phytosociology of the farm haribes in the nama-karoo biome of southern namibia’, koedoe 54(1), art. #1038, 13 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1038 note: nomenclature follows craven (1999) and subsequent name changes used by the national herbarium of namibia (wind). copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. phytosociology of the farm haribes in the nama-karoo biome of southern namibia in this original research... open access • abstract • introduction    • study area • research method • results    • description of the plant associations       • 1. lycio cinereum – salsoletum ass. nov.       • 1.1 the zygophylo simplicis – salsoletosum subass. nov.       • 1.2 the lycio cinereum – salsoletosum subass. nov.       • 2. the eragrostis rotifer – leptochloa fusca community       • 3. salsolo – tetragonietum schenckii ass. nov.       • 4. the cullen obtusifolia – leptochloa fusca community       • 5. anthephoro pubescentis – ziziphodetum mucronatae ass. nov.       • 5.1 ehretrio albae – ziziphodetosum mucronatae subass. nov.       • 5.2 heteropogo contorti – ziziphodetosum mucronatae subass. nov.       • 6. zygophylo pubescentis – leucosphaeretum bainesii ass. nov.       • 7. monsonio umbellatae – boscietum foetidae ass. nov.       • 8. acacio senegal – catophractetum alexandri ass. nov.       • 8.1 the schmidtio kalahariensis – boscietosum foetidae subass. nov.       • 8.2 the acacio senegal – catophractetosum alexandri subass. nov.    • vegetation mapping • discussion • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ limited historic vegetation data (prior to the 1980s) are available for namibia. finding such historic data at haribes prompted a follow-up survey of the vegetation. we present a classification of the recent data in this paper as a first step towards comparing the two data sets. six new associations (three with two subassociations each) are formally described. the landscape at haribes is dominated by a pan with surrounding hummock dunes. the pan supports the lycio cinereum – salsoletum, whilst on the hummock dunes the salsolo – tetragonietum schenckii can be found. the surrounding plains and escarpment can be divided into three landforms: the torras with the monsonio umbellatae – boscietum foetidae, the ranteveld with the acacio senegal – catophractetum alexandri (and two subassociations) and calcrete ridges with the zygophylo pubescentis – leucosphaeretum bainesii. dry river beds on the farm support two subassociations of anthephoro pubescentis – ziziphodetum mucronatae. the area covered by each dominant landform has been calculated after being mapped. the composition and diversity of the associations are briefly compared to other known vegetation descriptions within the nama-karoo. since november 2011, haribes has been used as a resettlement farm. this may result in the overutilisation of the limited grazing resources, to the extent that the present, fairly dense acacio senegal – catophractetum alexandri of the ranteveld is feared to become degraded to resemble the monsonio umbellatae – boscietum foetidae of the torras.conservation implications: this paper describes six plant associations of the nama-karoo biome in arid southern namibia. the information presented forms a baseline description, which can be used for future monitoring of the vegetation under altered land use. introduction top ↑ vegetation in namibia has been described only on a few attempts in the past (burke & strohbach 2000; strohbach & jürgens 2010). to date, the only concise vegetation map of namibia was produced in the 1970s, based on field observations rather than actual surveys (giess 1971, 1998), and depicts broad categories at biome rather than landscape level. later attempts to map the vegetation of namibia had to rely on partial, often incomplete, regional updates used in conjunction with the giess vegetation map to fill in gaps (mendelsohn et al. 2002). grazing capacity maps dating from the 1970s (departement landbou tegniese dienste 1979) are used for land-use planning, including the ongoing land reform programme of the government of namibia. however, these maps are regarded as outdated and in urgent need of revision (espach 2006; espach, lubbe & ganzin 2006; lubbe 2005). current efforts to upgrade the grazing capacity estimation with the aid of satellite images require extensive ground surveys, including surveys of the structure and composition of vegetation (espach et al. 2006; espach, lubbe & ganzin 2010). with this in mind, but also as basis for a biodiversity inventory, the vegetation survey of namibia project was initiated (strohbach 2001).prof. o.h. volk was one of only a few vegetation scientists active in namibia before the 1980s. during 1956, he conducted extensive vegetation surveys on selected farms in the former south west africa. unfortunately, only one of these data sets had been analysed and formally published (volk & leippert 1971); the others that could be retrieved were all in the form of raw data sets with brief comments on the vegetation, generally held in farm archives. nonetheless, these data sets have proved to be of immense value in reconstructing vegetation changes over the past five decades of land use, especially with regard to the effects of land use and/or global climate change. one of these data sets, surveyed in april 1956, was from the farm haribes. this study was initiated to provide data on the current vegetation, for later comparison with volk’s historic data. the aim of this paper, however, is to classify and describe the present vegetation of the farm haribes as an attempt towards filling the immense gap in vegetation data in namibia (strohbach & jürgens 2010). in a subsequent paper, we intend to compare the present vegetation with that documented in 1956. study area haribes, in the mariental district of the hardap region in southern namibia (figure 1), belonged to the neue haribes company until 2010. in 1956 the farm covered 77 500 ha, but has been reduced to its present 50 100 ha through a process of selling off blocks of land. the present survey was conducted across blocks 019/1, 019/rem, 020, 672 and 673, which total 55 487 ha. initially, haribes was used for extensive commercial karakul sheep farming. the most important land-use change took place in 1992, when all the sheep and goats were sold and replaced by cattle as the only domesticated commercial species. as a result of the selective utilisation of the grass layer, woody species, especially catophractes alexandri, rhigozum trichotomum and acacia nebrownii, increased in abundance, resulting in a phenomenon commonly referred to as ’bush encroachment‘ (de klerk 2004). a wide variety of game, often in large numbers, are found on the farm. most numerous are springbok and oryx. there are also many bird species, including many ostriches. the farm was sold in may 2010 to the government of namibia as part of its land reform and resettlement scheme (cloete 2010a; pretorius 2010), with the first 13 families receiving their plots in november 2010 (cloete 2010b; kleinhans 2010). figure 1: location of the farm haribes in the hardap region of southern namibia. also indicated on this map are the long-term average annual rainfall isohyets (in grey), with their values (in mm) indicated to the right. haribes is situated approximately 30 km west of mariental and southwest of the hardap dam nature reserve within the central plateau 4-8 agro-ecological zone (de pauw et al. 1998). this zone is described as a strongly dissected plain on karoo rocks. the average rainfall is about 200 mm per year, with a coefficient of variation between 50% and 60% (botha 1996). temperatures rise to above 36 °c in summer, with minimum temperatures of between 2 °c and 4 °c in winter (mendelsohn et al. 2002).the geomorphology can be described as a plateau broken up by many dry washes and some bigger ephemeral rivers that are responsible for the drainage of the area [a dissected plain according to the terminology of the digital soils and terrain database (soter) (fao 1995; oldeman & van engelen 1993]. the riet river drains into a big pan surrounding the farmhouse complex. in and around the pan several well-developed dune hummocks have formed over time. the farm lies between 1120 m and 1320 m above sea level. the geological strata belong to the fish river subgroup of the nama sediments. the eastern part of the farm is situated on the gross aub formation dominated by red shale and sandstone, whilst the larger, western part of the farm forms part of the older nababis formation with red sandstone and shales (geological survey 1980). the soils in the region (at a scale of 1:1 000 000) are classified as eutric leptosols, which means that the soils are fertile but generally very shallow (icc, mawrd & aeci 2000; mendelsohn et al. 2002). the farm falls within the nama-karoo biome (irish 1994) and its vegetation is classified as dwarf shrub savanna (giess 1971, 1998). this can broadly be described as open, short-to-low shrublands dominated by c. alexandri, r. trichotomum, boscia foetida and several dwarf shrubs. the grass layer, which includes species such as stipagrostis uniplumis, eragrostis nindensis, enneapogon cenchroides and schmidtia kalahariensis, is relatively sparse. trees and larger shrubs, including species such as a. nebrownii, acacia karroo, euclea pseudebenus and ziziphus mucronata, are largely confined to dry washes and rivers. research method top ↑ a total of 177 plots were sampled between march and may 2005. below average rain (c. 150 mm, d. poggenpoel, pers. comm., may 2005) had fallen that season, with the first substantial rains falling only in march. to compare the present composition with that documented by volk in 1956, we made exhaustive attempts to relocate his original sampling sites according to route descriptions, odometer distance readings (in miles) and old farm maps. although this information greatly eased the task of relocation, the exact position of those plots could not be determined. additional vegetation types were sampled using a stratified sampling approach to achieve a good average for the main vegetation types. for the purpose of stratification, a satellite map of the farm (based on landsat 7 etm scene path 177 row 077, 26 march 2001) was used as base map. relatively homogenous areas were delineated as derived from segmentations calculated by definiens 5 (definiens 2006). occasional azonal vegetation types (especially small vleis and related features) were also sampled. surveying followed the braun-blanquet concept as applied during the vegetation survey of namibia project (strohbach 2001). for each 20 m × 50 m plot, all species were recorded, as well as their typical growth form and average height. abundance was estimated according to the modified braun-blanquet scale (mueller-dombois & ellenberg 1974). in addition, the position of the plot and environmental parameters such as altitude, aspect, slope, topography, stone cover, erosion and disturbances were noted. owing to time constraints, no soil samples were taken. instead, available soil data (icc et al. 2000; mendelsohn et al. 2002) were interpolated for the various vegetation associations described. unknown plant species were collected and identified at the national herbarium of namibia. the relevé data were captured in the turboveg database (hennekens & schaminée 2001). the data were classified with juice (tichý 2002), using the modified twinspan classification procedure (roleček et al. 2009). the pseudospecies cut level was set to presence/absence, and total inertia was selected as heterogeneity measure. the classification process concluded after 14 divisions. at this stage, two communities (hummock dunes and the ranteveld) were already subdivided owing to high internal heterogeneity and had to be recombined manually (figure 2). the classification result was ordered as a table using the phi coefficient fidelity measure (chytrý et al. 2002) as sorting criterion. figure 2: dendrogram based on the modified twinspan classification of the vegetation types in the study area. the classification results were confirmed with an ordination using detrended correspondence analysis (dca) (hill & gauch 1980), prepared with pc-ord 5 (mccune, grace & urban 2002). two iterations were performed, with the first excluding only two wetland relevés (as outliers) whilst the second also excluded the pans and associated hummock dunes. although dca has been criticised in recent years for ’detrending‘ the arch effect, this very effect proved useful in illustrating the relationships between the various vegetation associations, as reciprocal averaging (ra) depicted only a severe arch effect. the dca ordination diagram was also used to depict habitat relations, which could often only be overlaid as categorical data. mapping was performed using a landsat 7 etm satellite image (path 177, row 077; 26 march 2001). in addition, a digital terrain model derived from shuttle radar topography mission (srtm) data (available from http://srtm.csi.cgiar.org/) (jarvis et al. 2008) was used to elucidate the topography. the satellite image was clipped to an area slightly larger than the study area before further processing. this clip was imported into the definiens software package (definiens 2006) and segmented into relatively homogenous areas. the segments were classified using the classified sample sites as ground truth data. in a final step, the resulting shape file was clipped according to the farm boundaries and areas for each landscape calculated using the idrisi andes software package (eastman 2006). results top ↑ a total of 218 species, representing 42 families, were observed on the 177 relevés. poaceae was the most abundant familiy, represented by the most species (54), followed by the fabaceae (27 species), asteraceae (16 species) and the aizoaceae (13 species). the classification of these relevés revealed eight communities, of which three were divided into subcommunities. the phytosociological table is presented in an online appendix 1, whereas online appendix 2 is a synoptic table. figure 2 depicts a dendrogram of the classification result. the ordination diagrams of the dca are shown in figure 3a (all data except for two outlier relevés) and figure 3b (also excluding the relevés representing pans and associated hummock dunes). in figure 3c, the lithology (i.e. the rock types which gave rise to the soils) has been imposed onto the dca ordination diagram of figure 3b to illustrate the habitat relationships between vegetation communities. figure 3: detrended correspondence analysis ordination diagrams compiled from (a) all data, excluding two outlying wetland relevés and (b) data excluding wetland, hummock dune and pan relevés; (c) indicates the lithology of each plot, with a biplot indicating the stoniness of the substrate. the pan and hummock dune communities form a well-defined continuum to the right of the ordination diagram in figure 3a, indicating most likely a combined salinity and soil moisture gradient from the pan centre (community 1.2; upper right corner) to the surrounding hummock dunes (community 3; lower right corner). in contrast, the remaining communities (plains, ranteveld and rivers) are compressed on the left of the main axis. splitting out the relevés representing the pan and hummock dune communities resulted in a continuum spread between the stony ranteveld, plain and torra communities as well as the rivers (figure 3b). the limited habitat data available indicate that this is a gradient of increased stoniness towards the left of the main gradient (figure 3c), representing the ranteveld. together with the lithology (i.e. the rock type from which the substrate is derived), the stoniness forms the main ecological gradient in this environment. description of the plant associations we formally describe six of the observed communities as associations, three with subassociations, following the international code of phytosociological nomenclature (weber, moravec & theurillat 2000). the two wetland communities, however, are described only informally, as the number of samples taken was too small for proper description. 1. lycio cinereum – salsoletum ass. nov the typicum for this association is relevé 2514, sampled at 24°35’36”s, 17°32’58”e on 09 march 2005 (figure 4b). the association is represented by 21 relevés and characterised by the presence of salsola spp. dwarf shrubs, which also dominate the vegetation. the structure is a short, open shrubland (sensu edwards 1983), with an average shrub cover of 6%. total vegetation cover barely reaches 10%. in total, 58 species have been observed on these pans, with an average species density of 12 per 1000 m2.the lycio cinereum – salsoletum is typical of the large haribes pan on the farm. the soils originate from weathered sandstones and some calcretes consisting of fine material which is very prone to capping and surface sealing (figure 4c). the cation levels are probably elevated to form a halophitic environment, but not sufficiently so to render the pan a salt pan (m. coetzee, pers.comm., march 2010). two subassociations can be distinguished, as described below. 1.1 the zygophylo simplicis – salsoletosum subass. nov. the typicum for this subassociation is relevé 2518, sampled at 24°37’53”s, 17°32’31”e on 09 march 2005 (figure 4a). only five relevés represent this subassociation. the occurrence of tetragonia schenkii, trianthema sp. and salsola spp. is characteristic and s. uniplumis var. uniplumis, s. kalahariensis, boerhavia repens and a. nebrownii also occur constantly.along the outer parts of the pan and on the floodplains to the pan the vegetation is better developed, forming a tall, moderately closed shrubland (figure 5). few perennial herbaceous species occur here; this layer is dominated by ephemerals. the species density is on average 14 species per 1000 m2 and in total 39 species have been observed here. 1.2 the lycio cinereum – salsoletosum subass. nov. this subassociation forms the typicum for the lycio cinereum – salsoletum. it is represented by 16 relevés and characterised by near-monospecific stands of salsola shrubs. other species that occur here are mostly of ephemeral nature. the shrubs are on average only 0.5 m – 1 m high and reach a cover of only about 6%, thus forming a short, open shrubland (figure 5). on average, only five species occur per 1000 m2, with a total of 31 species being observed on these pans. 2. the eragrostis rotifer – leptochloa fusca community this community was found once in relevé 2599 from a sample on a deeper vlei, which is regularly inundated. this is indicated by the presence of the geophytic hydrophyte aponogeton cf. desertorum and eragrostis rotifer, also generally associated with water-logged soils (figure 4d). these uncommon vleis form in relatively small depressions, mostly on top of the plateau area in pans that range from 20 m to 50 m in diameter. the pans are surrounded by sandstone stones varying in size between 5 cm and 80 cm. the reddish colour indicates that the soil is most probably relatively high in nutrient content (m. coetzee, pers. comm., march 2010). 3. salsolo – tetragonietum schenckii ass. nov. the typicum for this association is relevé 2501, sampled at 24°37’00”s, 17°31’01”e on 08 march 2005 (figure 4e). eleven relevés were classified within this association. the association is characterised by the presence of the endemic shrub t. schenckii. constant species include s. uniplumis var. uniplumis, stipagrostis ciliata, salsola spp. and r. trichotomum. the association is dominated by t. schenckii. the typical structure is a tall, moderately closed shrubland (figure 5), with the shrubs reaching a height of between 1 m and 1.5 m. on average 11 species are found per 1000 m2, with a total diversity of 56 species.the salsolo – tetragonietum schenckii is found on hummock dunes probably originating from a deflation process during the formation of the haribes pan, similar to that described for the koës pan and other depressions in the kalahari (bullard & nash 2000; lancaster 1986; shaw 1988; shaw & thomas 1997). the dunes vary in length, generally between 30 m and 50 m. the slopes of the dunes are often quite steep. the occurence of salsola spp. on these dunes indicate a high base status, which, in turn, supports the hypothesis that the dunes are the result of the deflation process of the pan. 4. the cullen obtusifolia – leptochloa fusca community this community was identified in three relevés within the larger haribes pan ecosystem. the presence of leptochloa fusca and indigofera alternans indicates water-logging at these vleis, whilst the absence of e. rotifer and aponogeton sp. and the presence of the flat-growing therophytes cullen obtusifolia and platycarpha carlinoides indicate a drier environment than occurs in the deep vleis of the e. rotifer – leptochloa fusca community. typically for these vleis, a. nebrownii is dominant along the fringes (figure 4f). these pans are small (50 m – 80 m in diameter) and very shallow. the high clay content of the soil allows the water to remain in the pan for approximately 2–3 weeks. in most cases, these pans are surrounded by reddish gravel, but no larger stones. there are very few of these small pans that differ greatly from the big pans on the farm. some are relatively small depressions, mostly on top of the plateau area, but there are also small pans below the plateau, especially near the old road along the tar road between mariental and maltahöhe. these depressions allow water to accumulate, resulting in microhabitats with a wetland character. 5. anthephoro pubescentis – ziziphodetum mucronatae ass. nov. the typicum for this association is relevé 2649, sampled at 24°38’17”s, 17°28’12”e on 16 may 2005 (figure 4h). a total of 30 relevés have been classified within this association. it is characterised by the diagnostic species z. mucronata, ocimum americanum var. americanum, aristida meridionalis, grewia flava, anthephora pubescens, monechma divaricatum and melinis repens. these are constantly associated with the species s. uniplumis var. uniplumis, s. kalahariensis, lycium cinereum, e. cenchroides, gisekia africana, a. nebrownii, eragrostis porosa, r. trichotomum and b. foetida. the structure is a low open woodland (sensu edwards 1983), dominated by trees between 4 m and 6 m high that attain a cover of about 2%. shrubs attain a cover of between 5% and 10%, at a height of about 1 m to 2 m. a total of 130 species have been observed in this association.the anthephoro pubescentis – ziziphodetum mucronatae represents the riverine woodlands (sensu giess 1998). there are two types of drainage system on the farm, with dry washes, which contain water only for very short periods during rain storms, being most common. geomorphologically they can be classified as very young river systems. in many cases the slope is relatively steep and the run-off very fast. they occur mostly on the plateau or at the edge of the plateau. towards the centre of the farm, more mature rivers, like the riet, holz and lewer, are responsible for drainage. there is a big dam near the farmhouse, where the water of these rivers is captured. these two different habitats support two different vegetation subassociations (5.1 and 5.2). although the vegetation of the riverbed and the riverbank differ, they were sampled as one for the purposes of this study and are thus included in the same plant association. 5.1 ehretrio albae – ziziphodetosum mucronatae subass. nov. the typicum of this subassociation is relevé 2574, sampled at 24°32’34”s, 17°32’45”e on 11 may 2005 (figure 4g). a total of 17 relevés belong to this subassociation. there are no diagnostic species for this association, but the following are constantly present: s. uniplumis var. uniplumis, s. kalahariensis, l. cinereum, a. nebrownii, r. trichotomum, limeum sulcatum, g. africana, b. foetida, e. cenchroides, g. flava, e. porosa and b. repens. the structure is a low, open woodland (figure 5), with the trees only 4 m – 5 m tall and shrubs reaching height of 1 m – 2 m. in this subassociation, 99 species have been recorded; on average 21 species occur per 1000 m2.the ehretrio albae – ziziphodetosum mucronatae occurs along the dry washes. their slopes vary from steep to almost level (15°–20° and < 3°, respectively). erosion is a common occurrence on the steeper slopes. most of these dry washes start on top of the plateau and generally run eastward or southward to join the riet, holz and lewer rivers. they are all typically ephemeral and are active only during and shortly after thunderstorms. varied soil character is observed: although most soils are very shallow, occasionally with big rocks, gravel and small stones also represent a common soil cover. at the base of the plateau some of the washes have deeper soils, which are also higher in nutrient content. 5.2 heteropogo contorti – ziziphodetosum mucronatae subass. nov. this subassociation, with 13 relevés, is the typicum of the anthephoro pubescentis – ziziphodetum mucronatae (figure 4h). it is characterised by the presence of the diagnostic species heteropogon contortus, cleome hirta, kyphocarpa angustifolia and senecio sp. in addition to the diagnostic species defining the anthephoro pubescentis – ziziphodetum mucronatae. these are also constantly accompanied by tephrosia dregeana, otoptera burchellii, g. flava, tribulus terrestris, mollugo cerviana, kohautia caespitosa subsp. brachyloba and cenchrus ciliaris (in addition to the constant species listed for the association). owing to the higher moisture content in the soils, the trees grow up to 7 m tall, thus forming a short, open woodland with a tree cover of 2% (figure 5). species density is on average 28 species per 1000 m2, whilst a total of 109 species have been observed in this subassociation.the heteropogo contorti – ziziphodetosum mucronatae occurs along the more mature rivers (riet, holz and lewer). their slopes are normally less than 3° and water remains standing in river pools for long periods. the soils in the riverbeds are normally rough gravels, which can be very stony with big rocks. the nutrient level in these gravels is low, but water content is usually relatively high. the soils of the river banks are sandy loams with a relatively high nutrient content, whilst the moisture content varies depending on the slope of the river. 6. zygophylo pubescentis – leucosphaeretum bainesii ass. nov. the typicum for this association is relevé 2526, sampled at 24°32’34”s, 17°37’08”e on 10 march 2005 (figure 4i). the association comprises 14 relevés and is characterised by the diagnostic species leucosphaera bainesii, commiphora glandulosa, zygophyllum pubescens, stipagrostis obtusa and eriocephalus pubescens. these are constantly associated with b. foetida, s. uniplumis var. uniplumis, r. trichotomum, c. alexandri, e. cenchroides, o. burchellii, enneapogon desvauxii, dicoma capensis and tribulus sp. the structure is a short, moderately closed shrubland (figure 5), with shrubs standing approximately 0.5 m – 2 m tall and a canopy cover of about 10%, reaching up to 20%. here, 65 species have been observed, with an average of 19 species per 1000 m2. zygophylo pubescentis – leucosphaeretum bainesii is associated with shallow calcrete soils, either along the northern haribes pan edge, as part of the flood plain system of larger rivers or on dolerite ridges. the soils are shallow (20 cm – 30 cm deep), whitish in colour and have a high clay content. the soils are expected to be alkaline with a ph of more than 7 (m. coetzee, pers. comm., march 2010). stone cover in the form of calcrete gravel and pebbles is often found mixed with medium to large dolerite blocks and can reach up to 80%. steep terraces, with slopes of between 5° and 10°, are formed along the pan edge. 7. monsonio umbellatae – boscietum foetidae ass. nov. the typicum for this association is relevé 2632, sampled at 24°37’10”s, 17°37’03”e on 15 may 2005 (figure 4j). this association was found on 17 plots in the study area. the occurrence of the low spreading forb monsonia umbellata (‘rabas’) is characteristic of this association. in addition, the constant presence of s. uniplumis var. uniplumis, r. trichotomum, c. alexandri, b. foetida, s. kalahariensis, tribulus cristatus, ptychtolobium biflorum subsp. biflorum, g. africana, e. desvauxii, d. capensis and t. dregeana is typical for this association. the vegetation structure can be defined as a tall, open shrubland (figure 5). the shrubs are between 0.5 m and 2 m high and represent less than 5% of the ground cover. density is on average 17 species per 1000 m2, with a total of 67 species being observed in this association.the monsonio umbellatae – boscietum foetidae typically occurs on the torras on the farm. torras are defined as flat plains, with little or no internal relief, covered with reddish brown gravel on the surface (often more than 80%). the soils are loamy, with a crust, and with very low water infiltration capacity. this low water infiltration capacity results in patchy, near-banded vegetation, with run-off from the open patches feeding the vegetated patches (run-on). similar vegetation patterns have been described as brousse tigree in the sahel (boaler & hodge 1962; bromley et al. 1997; couteron et al. 2000; thiéry, d’herbès & valentin 1995) and banded mulga in arid australia (dunkerley 1997a, 1997b). 8. acacio senegal – catophractetum alexandri ass. nov. the typicum for this association is relevé 2606, sampled at 24°32’31”s, 17°28’59”e on 13 may 2005 (figure 4l). the association comprises 80 relevés. it is characterised by eragrostis nindensis as characteristic species, with the following constant species: s. uniplumis var. uniplumis, c. alexandri, b. foetida, l. cinereum, e. cenchroides, s. kalahariensis, triraphis ramosissima, t. dregeana and aristida adscensionis. the structure is a typical tall, open shrubland, with shrubs of 1 m – 2 m in height, reaching a cover of about 5%. the total cover varies between 20% and 50%. species density is on average 19 species per 1000 m2, whilst a total of 131 species have been recorded for this association. two subassociations have been identified within this association. 8.1 the schmidtio kalahariensis – boscietosum foetidae subass. nov. the typicum of this subassociation is relevé 2546, sampled at 24°41’17”s, 17°34’54”on 14 march 2005 e (figure 4k). this subassociation was represented by 28 relevés and is characterised by the following constant species: s. uniplumis var. uniplumis, b. foetida, l. cinereum, r. trichotomum, c. alexandri, s. kalahariensis, e. cenchroides, tribulus sp., d. capensis, a. nebrownii, t. dregeana and aptosimum albomarginatum. no characteristic species were identified. the structure is a tall, open shrubland (figure 5), with a shrub cover of less than 5%. shrub height ranges between 0.5 m and 1.8 m. the species density is on average 18 species per 1000 m2, whilst a total of 87 species have been found in this subassociation.the schmidtio kalahariensis – boscietosum foetidae occurs on flat plains and footslopes as well as on ranteveld proper and forms a transition between the torras and the ranteveld. these plains are underlain by sandstones of the nababis formation. fragments of these form the stone cover on the soil surface, ranging from gravel to medium-sized stones and covering up to 80% of the soil surface. the low vegetative cover indicates a tendency to degrade to a similar vegetation as the monsonio umbellatae – boscietum foetidae on the present torras (figure 4j). 8.2 the acacio senegal – catophractetosum alexandri subass. nov. this subassociation forms the typicum of the acacio senegal – catophractetum alexandri (figure 4l). a total of 52 relevés have been classified into this subassociation. it is characterised by e. nindensis as diagnostic species, with s. uniplumis var. uniplumis, c. alexandri, b. foetida, l. cinereum, e. cenchroides, t. ramosissima, g. africana, s. kalahariensis, a. adscensionis, t. dregeana and eragrostis porosa as constant species. this subassociation forms a tall, open shrubland or occasionally a low bushveld (figure 5), with shrubs of 0.5 m – 3 m tall. occasional trees stand 4 m – 5 m tall. on average, 19 species have been found per 1000 m2, whilst the total species diversity is 105.the acacio senegal – catophractetosum alexandri forms the ranteveld and occurs mainly in areas on top of the plateau. the plateau is dominated by very stony outcrops and small escarps (rante) of nababis sandstone. the soils are leptosols, red in colour and very shallow, of moderately coarse to medium texture, with many coarse fragments and well drained. the surface layer is typically 10 cm – 20 cm thick. these soils have a moderate hydraulic conductivity and present a moderate infiltration rate with a low water holding capacity (icc et al. 2000). the landscape is relatively flat on top of the plateau, but forms occasional low hills and an escarpment area to the west of the farm, with steep-sloped ravines in between. figure 4: typical vegetation at different sampling sites on the farm haribes. figure 5: box-and-whisker plots of the various structural components for each of the described associations or subassociations. vegetation mapping it was not possible to map all vegetation types, for several reasons. firstly, many units are just too small to map with the 30-m ground resolution of landsat 7 etm satellite images. this holds true for the two vlei communities, e. rotifer – l. fusca and c. obtusifolia – l. fusca, as well as the zygophylo pubescentis – leucosphaeretum bainesii, which occurs on ridges with very limited spatial extent.differentiating between the monsonio umbellatae – boscietum foetidae and the schmidtio kalahariensis – boscietosum foetidae proved extremely difficult owing to their similar structures and especially similar low vegetative cover. for this reason, a landscape map was produced (figure 6) to differentiate between plains and torras, and ranteveld. table 1 lists the mapped landscapes, their spatial extent and their included vegetation associations or subassociations. table 1: landscapes and associated vegetation associations or subassociations on the farm haribes. discussion top ↑ four distinct higher syntaxonomic groupings can be recognised, namely the saline pans and associated hummock dunes (as represented by the lycio cinereum – salsoletum and the salsolo – tetragonietum schenkii), the wetlands (as represented by the two vlei communities), the river systems (as represented by the anthephoro pubescentis – ziziphodetum mucronatae) and the surrounding plateaus (as represented by the zygophylo pubescentis – leucosphaeretum bainesii, the monsonio umbellatae – boscietum foetidae and the acacio senegal – catophractetum alexandri). unfortunately the data set is too limited to define and describe such higher syntaxonomic levels at this stage. volk and leippert (1971) refer to the class rhigozetea as representing the larger dwarf shrub savanna of southern namibia; this encompasses all those syntaxa containing r. trichotomum (i.e. specifically the latter grouping). giess (1998) superficially described the dwarf shrub savanna, as follows: characteristic for the greatest part is rhigozum trichotomum (…). parkinsonia africana, acacia nebrownii, boscia foetida subsp. foetida, b[oscia] abitrunca var. albitrunca and catophractes alexandri as well as karoo bushes such as pentzia spp., eriocephalus spp., and others are typical for this vegetation type. (giess 1998) this description is illustrated with three plates. yet the acacio senegal – catophractetum alexandri has a too tall structure (2 m – 3 m) to be regarded as true dwarf shrub savanna. within the broader nama-karoo (of which giess’ dwarf shrub savanna forms the northern extent), the acacio senegal – catophractetum alexandri can structurally be compared to the acacia mellifera open shrubland of the blouputs karroid thornveld as described for the augrabies falls national park (bezuidenhout 1996; mucina & rutherford 2006). however, the shrub component here is dominated by a. mellifera subsp. detinens, a species rarely found in the acacio senegal – catophractetum alexandri. in this respect, the acacio senegal – catophractetum alexandri is to be regarded as a xeric outlier of the highland savanna (sensu giess 1998). the rocky habitat with shallow soils, and often steep slopes, is similar in both vegetation types. the salsolo – tetragonietum schenckii resembles the saline desert and dwarf shrub savanna fringe (sensu giess 1998) described for the etosha pan. here the structure of the hummock dune vegetation can specifically be compared to the okondeka duneveld (le roux 1980). however, the composition varies widely, with t. schenkii not occurring in northern namibia. for the lycio cinereum – salsoletum no comparable vegetation type is known. it is likely that these, in a future synopsis of syntaxa, will form a class on their own. the very close relationship between the monsonio umbellatae – boscietum foetidae and the two subassociations of the acacio senegal – catophractetum alexandri is well illustrated in the ordination diagram in figure 3b. this leads to the assumption that these three units form degradation stages of one another: acacio senegal – catophractetosum alexandri degrading to schmidtio kalahariensis – boscietosum foetidae, and in very extreme cases (possibly also owing to soil degradation processes such as erosion and capping) to monsonio umbellatae – boscietum foetidae. this hypothesis needs to be tested and described in a paper in which dynamic trends are considered. with reference to biodiversity patterns, species density is comparable to other descriptions within the nama-karoo biome in southern namibia. the nico observatories (nico north and nico south) of the biota southern africa project are the nearest to haribes and are also situated on relatively stony soil, albeit of origin from the dwyka formation (geological survey 1980). the nico observatories display an average of 21 and 20 species per 1000 m2, respectively (jürgens et al. 2010), compared with the 17–19 species per 1000 m2 for the zygophylo pubescentis – leucosphaeretum bainesii, monsonio umbellatae – boscietum foetidae and acacio senegal – catophractetum alexandri. the slightly lower values observed during the current study can be related to the survey being conducted during a single, comparatively dry year, whilst the nico observatories had been resurveyed over a period of seven years (but unfortunately not in 2005). however, the overall number of species observed at haribes (218) is higher than the total number of species observed at both nico north (166) and nico south (204). this can, in turn, be attributed to the fact that the area sampled at haribes is far bigger and more diverse than the 1 km2 sample areas of the biota observatories. comparing the zygophylo pubescentis – leucosphaeretum bainesii to the narias and duruchaus observatories, which also have shallow soils on calcretes (thus a comparable edaphic habitat), a far lower species density was observed (19 vs 36–37 per 1000 m2). the climatic environment at the narais and duruchaus observatories is, however, far more moist, with a mean annual rainfall of et al. 2010). conclusion top ↑ haribes is unlikely ever to become a conservation area. on the contrary, the recent change in ownership also introduces an entirely different land use, with the farm being targeted for resettlement. some 13 families have already been resettled on small parcels of the farm, each ranging between 1700 ha and 7500 ha (cloete 2010b; kleinhans 2010). these units, meant to support the livelihoods of these families (werner & odendaal 2010), have been calculated according to grazing capacity maps from the 1970s (department landbou tegniese dienste 1979). however, the general indications are that the grazing capacity has drastically reduced throughout the entire country over the past decades (espach et al. 2010). this is due to a multitude of factors generally associated with land degradation [e.g. bush encroachment (de klerk 2004)]. it is thus unlikely that these small farming units will be sustainable and severe degradation of grazing resources may result (werner & odendaal 2010).within the nama-karoo biome in namibia, livestock farming depends mainly on browsing of dwarf shrub species (espach et al. 2006), with grazing playing a secondary role. this does not hold true for haribes. a comparison of the shrub, dwarf shrub, grass and total vegetation cover of the various associations (figure 5) shows that farming on haribes is more dependent on grazing than browsing. the pans and surrounding dunes are mostly covered by shrubs dominated by salsola spp. and t. schenckii, which are both known to be poisonous (mannheimer, marais & schubert 2008). this makes roughly 13% of the farm unsuitable for grazing. the zygophylo pubescentis – leucosphaeretum bainsii on the calcrete ridges provide some dwarf shrub browse, but is very limited in extent. the monsonio umbellatae – boscietum foetidae (torras) and the acacio senegal – catophractetum alexandri (ranteveld) provide the main grazing resources on the farm. however, as pointed out in the discussion, these seem to form a degradation gradient from each other; thus, extensive overgrazing will lead to desert-like torra plains. figure 6: map of the major landscapes on the farm haribes. global climate change has been identified as a reality in recent years. temperature increases coupled with less precipitation, in more erratic events, are predicted for southern africa (parry et al. 2007). based on these forecasts, it is predicted that large parts of southern namibia will become unsuitable for livestock farming, as it will turn into a virtual desert (midgley et al. 2005). similar scenarios have been developed for the kalahari basin, with forecasts of remobilisation of sand dunes as far north as southern angola (thomas, knight & wiggs 2005). in addition, a human-induced drought, as described for other nama-karoo regions (booysen & rowswell 1983), threatens. this is aggravated by the fact that farming on haribes depends largely on grazing, meaning that protection of the perennial grass cover is essential for sustainable use of the farm in the long term. perennial grasses have been shown to be able to withstand prolonged droughts (i.e. periods of below-normal precipitation), but are unable to withstand heavy grazing together with arid conditions (o’connor 1991, 1994). it is thus likely that overgrazing in the ranteveld may lead to a composition (and vegetation cover) similar to that of the present torras. when planning the redistribution of the farm haribes, it should be kept in mind that only 75% of the farm (the ranteveld) provides adequate grazing and that even this grazing needs to be utilised judiciously to prevent severe degradation to desert-like conditions. acknowledgements top ↑ we are grateful to mr and mrs poggenpoel, previous managers at the farm haribes, for making volk’s data of 1956 available to us and for their kind support during fieldwork. the assistance of staff members at the national herbarium of namibia (wind) during specimen identification is gratefully acknowledged. many thanks also go to mrs marina coetzee of the school of natural resources and tourism, polytechnic of namibia for providing information on soil properties. this work was financially supported by the german federal ministry of education and research under their biolog programme, promotion number 01 lc 0024a, as part of the biota southern africa project. competing interests the authors declare that they have no financial or personal relationship(s) which may have inappropriately influenced them in writing this paper. authors’ contributions b.j.s. (national botanical research institute) initiated the project. w.j.j. (school of natural resources and tourism) was responsible for the vegetation surveys and the habitat and structural descriptions of the associations, whilst b.j.s. (national botanical research institute) was responsible for data analysis, diagnostic descriptions of the associations, and mapping. references top ↑ bezuidenhout, h., 1996, ‘the major vegetation communities of the augrabies falls national park, northern cape. 1. the southern section’, koedoe 39, 7–24.boaler, s.b. & hodge, c.a.h., 1962, ‘vegetation stripes in somaliland’, journal of ecology 50, 465–474. http://dx.doi.org/10.2307/2257457 booysen, j. & rowswell, d.i., 1983, ‘the drought problem in the karoo areas’, proceedings of the grassland society of southern africa 18, 40–45. botha, l., 1996, rainfall as an indicator of the agroclimate of namibia, m.dip.tech. thesis, faculty of engineering, technikon pretoria. bromley, j., brouwer, j., barker, a.p., gaze, s.r. & valentin, c., 1997, ‘the role of surface water redistribution in an area of patterned vegetation in a semi-arid environment, south-west niger’, journal of hydrology 198, 1–29. http://dx.doi.org/10.1016/s0022-1694(96)03322-7 bullard, j.e. & nash, d.j., 2000, ‘valley-marginal sand dunes in the south-west kalahari: their nature, classification and possible origins’, journal of arid environments 45, 369–383. http://dx.doi.org/10.1006/jare.2000.0646 burke, a. & strohbach, b.j., 2000, ‘vegetation studies in namibia’, dinteria 26, 1–24. chytrý, m., tichý, l., holt, j. & bottadukat, z., 2002, ‘determination of diagnostic species with statistic fidelity measures’, journal of vegetation science 13, 79–90. http://dx.doi.org/10.1111/j.1654-1103.2002.tb02025.x cloete, l., 2010a, ‘government buys resettlement farm for n$10m’, the namibian, 06 august, p. 5. cloete, l., 2010b, ‘landless families become proud owners of land’, the namibian, 22 november, p. 7. couteron, p., mahamane, a., ouedraogo, p. & seghieri, j., 2000, ‘differences between banded thickets (tiger bush) at two sites in west africa’, journal of vegetation science 11, 321–328. http://dx.doi.org/10.2307/3236624 craven, p. (ed.), 1999, a checklist of namibian plant species, southern african botanical diversity network, windhoek (southern african botanical diversity network report, no 7). de klerk, j.n., 2004, ‘bush encroachment in namibia’, report on phase 1 of the bush encroachment research, monitoring and management project, ministry of environment and tourism, windhoek. de pauw, e., coetzee, m.e., calitz, a.j., beukes, h. & vits, c., 1998, ‘production of an agro-ecological zones map of namibia (first approximation). part ii: results’, agricola 10, 33–43. definiens version 5, 2006, computer software, definiens ag, munich. departement landbou tegniese dienste, 1979, die afbakening van redelike homogene boerderygebiede van die noordelike en sentrale substreke van s.w.a. met die heersende knelpunte en beoogde ontwikkleingsprogramme vir die verskillende bedryfstakke [the delimitation of relative homogenous farming areas in the northern and central subregions of swa with the present constraints and intended development programmes for the different enterprises], departement landbou tegniese dienste, windhoek. dunkerley, d.l., 1997a, ‘banded vegetation: survival under drought and grazing pressure based on a simple cellular automaton model’, journal of arid environments 35, 419–428. http://dx.doi.org/10.1006/jare.1995.0158 dunkerley, d.l., 1997b, ‘banded vegetation: development under uniform rainfall from a simple cellular automaton model’, plant ecology 129, 103–111. http://dx.doi.org/10.1023/a:1009725732740 eastman, j.r., 2006, idrisi. the andes edition, computer software, clark university, worcester, massachusetts. edwards, d., 1983, ‘a broad-scale structural classification of vegetation for practical purposes’, bothalia 14, 705–712. espach, c., 2006, ‘rangeland productivity modelling: developing and customising methodologies for land cover mapping in namibia’, agricola 16, 20–27. espach, c., lubbe, l.g. & ganzin, n., 2006, ‘determining grazing capacity in namibia: approaches and methodologies’, agricola 16, 28–39. espach, c., lubbe, l.g. & ganzin, n., 2010, ‘determining grazing capacity in namibia with the aid of remote sensing’, agricola 20, 41–61. fao. see food and agriculture organization. food and agriculture organization, 1995, global and national soils and terrain digital databases (soter), food and agriculture organization, water development division, rome (world soil resources reports, no. 74 rev. 1). geological survey, 1980, south west africa/namibia geological map 1:1000000. giess, w., 1971, ‘a preliminary vegetation map of south west africa’, dinteria 4, 1–114. giess, w., 1998, ‘a preliminary vegetation map of namibia’, dinteria 4, 1–112. hennekens, s.m. & schaminée, j.h.j., 2001, ‘turboveg, a comprehensive data base management system for vegetation data’, journal of vegetation science 12, 589–591. http://dx.doi.org/10.2307/3237010 hill, m.o. & gauch, h.g., 1980, ‘detrended correspondence analysis: an improved ordination technique’, vegetatio 42, 47–58. http://dx.doi.org/10.1007/bf00048870 icc, mawrd & aeci, 2000, ‘project to support the agro-ecological zoning programme (aez) in namibia’, report, institut cartogràfic de catalunya (icc), namibian ministry of agriculture, water and rural development (mawrd) and spanish agency for international co-operation (aeci), windhoek. irish, j., 1994, ‘the biomes of namibia, as determined by objective categorisation’, navorsinge van die nasionale museum bloemfontein 10, 549–592. jarvis, a., reuter, h.i., nelson, a. & guevara, e., 2008, hole-filled seamless srtm data v4, viewed 03 september 2010 from http://srtm.csi.cgiar.org/ jürgens, n., haarmeyer, d.h., luther-mosebach, j., dengler, j., finckh, m. & schmiedel, u. (eds.), 2010, patterns at local scale: the biota observatories, biodiversity in southern africa, klaus hess, göttingen & windhoek. kleinhans, t., 2010, ‘staat gee suide-grond – met voorwaardes [government gives land in south – with conditions]’, republikein, 25 november, p. 5. lancaster, n., 1986, ‘pans in the southwestern kalahari: a preliminary report’, palaeoecology of africa and the surrounding islands 17, 59–67. le roux, c.j.g., 1980, ‘vegetation classification and related studies in the etosha national park’, dsc thesis, dept. of plant production, university of pretoria. lubbe, l.g., 2005, ‘towards an updated carrying capacity map for namibia: a review of the methodologies currently used to determine carrying capacity in namibia’, agricola 15, 33–39. mannheimer, c., marais, a. & schubert, s., 2008, toxic plants of veterinary importance in namibia, 2nd ed., ministry of agriculture, water and forestry, windhoek. (addendum to agricola 2007) mccune, b., grace, j.b. & urban, d.l., 2002, analysis of ecological communities, mjm software, gleneden beach, oregon. mendelsohn, j., jarvis, a., roberts, c. & robertson, t., 2002, atlas of namibia, david phillips, cape town, pp. 55-100. midgley, g., hughes, g., thuiller, w., drew, g. & foden, w., 2005, ‘assessment of potential climate change impacts on namibia’s floristic diversity, ecosystem structure and function’, report, climate change research group, south african national biodiversity institute for the namibian national biodiversity programme, directorate of environmental affairs, cape town. ministry of agriculture, water and rural development, 2001, namibian agricultural resources information system (naris), ministry of agriculture, water and rural development, windhoek. mucina, l. & rutherford, m.c. (eds.), 2006, the vegetation of south africa, lesotho and swaziland, south african national biodiversity institute, pretoria (strelitzia, no. 19). mueller-dombois, d. & ellenberg, h., 1974, aims and methods of vegetation ecology, john wiley & sons, new york. o’connor, t.g., 1991, ‘influence of rainfall and grazing on the compositional change of the herbaceous layer of a sandveld savanna’, journal of the grassland society of southern africa 8, 103–109. o’connor, t.g., 1994, ‘composition and population responses of an african savanna grassland to rainfall and grazing’, journal of applied ecology 31, 155–171. http://dx.doi.org/10.2307/2404608 oldeman, l.r. & van engelen, v.w.p., 1993, ‘a world soils and terrain digital database (soter): an improved assessment of land resources’, geoderma 60, 309–325. http://dx.doi.org/10.1016/0016-7061(93)90033-h parry, m.l., canziani, o.f., palutikof, j.p., van der linden, p.j. & hanson, c.e. (eds.), 2007, contribution of working group ii to the fourth assessment report of the intergovernmental panel on climate change, cambridge university press, cambridge. pretorius, m., 2010, ‘50 100 hektaar in suide vir hervestiging gekoop [50 100 hectares bought in south for resettlement]’, republikein, 10 augustus, p. 7. roleček, j., tichý, l., zelený, d. & chytrý, m., 2009, ‘modified twinspan classification in which the hierarchy respects cluster heterogeneity’, journal of vegetation science 20, 596–602. http://dx.doi.org/10.1111/j.1654-1103.2009.01062.x shaw, p.a., 1988, ‘lakes and pans’, in b.p. moon & g.f. dardis (eds.), the geomorphology of southern africa, pp. 120–144, southern book publishers, johannesburg. shaw, p.a. & thomas, d.s.g., 1997, ‘pans, playas and salt lakes’, in d.s.g. thomas (ed.), arid zone geomorphology, pp. 293–317, john wiley & sons, chichester. strohbach, b.j., 2001, ‘vegetation survey of namibia’, journal of the namibia scientific society 49, 93–124. strohbach, b.j. & jürgens, n., 2010, ‘towards a user-friendly vegetation map of namibia: ground truthing approach to vegetation mapping’, in u. schmiedel & n. jürgens (eds.), biodiversity in southern africa: patterns and processes at regional scale, pp. 46–56, klaus hess publishers, göttingen & windhoek. thiéry, j.m., d’herbès, j.m. & valentin, c., 1995, ‘a model simulating the genesis of banded vegetation patterns in niger’, journal of ecology 83, 497–507. http://dx.doi.org/10.2307/2261602 thomas, d.s.g., knight, m. & wiggs, g.f.s., 2005, ‘remobilization of southern african desert dune systems by twenty-first century global warming’, nature 435, 1218–1221. http://dx.doi.org/10.1038/nature03717, pmid:15988522 tichý, l., 2002, ‘juice, software for vegetation classification’, journal of vegetation science 13, 451–453. http://dx.doi.org/10.1111/j.1654-1103.2002.tb02069.x volk, o.h. & leippert, h., 1971, vegetationsverhältnisse im windhoeker bergland, südwestafrika [vegetation relationships in the windhoek mountainveld, south-west africa], journal der s.w.a. wissenschaftliche gesellschaft xxv, 5–44. weber, h.e., moravec, j. & theurillat, j.p., 2000, ‘international code of phytosociological nomenclature, 3rd edn.’, journal of vegetation science 11, 739–768. http://dx.doi.org/10.2307/3236580 werner, w. & odendaal, w., 2010, livelihoods after land reform. namibia country study, land, environment and development project, legal assistance centre, windhoek. article information authors: milingoni p. tshisikhawe1 magaretha w. van rooyen2 jerome y. gaugris2 affiliations: 1department of botany, university of venda, south africa 2department of plant science, university of pretoria, south africa correspondence to: milingoni tshisikhawe postal address: private bag x5050, thohoyandou 0950, south africa dates: received: 01 feb. 2012 accepted: 05 july 2013 published: 04 sept. 2013 how to cite this article: tshisikhawe, m.p., van rooyen, m.w. & gaugris, j.y., 2013, ‘is the present brackenridgea nature reserve large enough to ensure the survival of brackenridgea zanguebarica oliv.?’, koedoe 55(1), art. #1072, 5 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1072 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/ koedoe.v55i1.1072-1 copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. is the present brackenridgea nature reserve large enough to ensure the survival of brackenridgea zanguebarica oliv.? in this short communication... open access • abstract • introduction • methods    • step 1    • step 2    • step 3    • step 4    • step 5    • step 6    • step 7    • step 8    • step 9    • step 10    • step 11    • step 12 • results    • minimum viable population size    • identification of populations with similar disturbance    • potential habitat    • potential habitat surveyed    • density of mature individuals    • minimum target area    • proportion of land lost    • proportion irreversibly damaged in 50 years    • compensation    • assessment of catastrophes    • combination of targets    • evaluation of current strategies • discussion • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ the brackenridgea nature reserve is a 110 ha protected area established by the provincial limpopo department of economic development, environment and tourism as a way of protecting the population of brackenridgea zanguebarica, a species classified as critically endangered in south africa. in the whole of south africa, the species is found in only one small area around thengwe–mafukani in venda. it is threatened with extirpation due to high demand for its medicinal bark. this study investigated the adequacy of the reserve to conserve the species using a method established in 2001 by burgman et al. this method involves 12 steps to quantify the risk of the decline or possible extinction of the species and takes current human activities, disturbances and the viability of the population into consideration for setting a conservation target. from the results, it was clear that more area is needed for the current population to survive beyond 50 years. assuming the status quo, it will require 410 ha to maintain the population, whereas a 50% reduction in human-related activities, such as cultivation, harvesting and livestock grazing, will lower the required potential habitat to 203 ha and a conservation option, which allows for bark harvesting, will require 179 ha.conservation implications: the results of this study will have conservation implication on management of viable species population within a nature reserve. it will require managers to take into consideration the reserve size in relation to potential habitats for the development of species under their management. introduction top ↑ the brackenridgea nature reserve (bnr), or better known as the mutavhatsindi nature reserve, is a protected area established in 1987 by the provincial limpopo department of economic development, environment and tourism in a proactive attempt to protect the population of brackenridgea zanguebarica oliv. (ochnaceae), commonly known as yellow peeling plane (english) or mutavhatsindi (venda). in south africa, the species is confined to a small area around thengwe–mafukani in venda, but is common in zimbabwe and mozambique northwards to tanzania. it is threatened with extirpation due to its high demand as a magical and medicinal plant species (netshiungani & van wyk 1980; todd et al. 2004) and is currently classified as critically endangered in south africa (williams & raimondo 2008).the population at mafukani is facing a serious threat due to the uncontrolled bark harvesting of b. zanguebarica. the questions therefore arise as to whether the current brackenridgea nature reserve is adequate to ensure the survival of the species and, if not, in view of the current threats, what would be an adequately sized area to ensure the survival of the species? population viability analysis (pva) is regarded as one of the cornerstones of conservation science and has been used to estimate the minimum viable population for threatened taxa (beissinger & mccullough 2002; menges 2000; pfab & witkowski 2000). it provides a framework to determine how stochastic events and processes affect the extinction probability of a species. pva can inform whether the size of a reserve is large enough to conserve a particular species, but data needed for a realistic pva may take many years to gather (menges 2000). furthermore, estimation of extinction risks for a large number of species requires an immense database (burgman et al. 2001; cabeza & van teeffelen 2009) that is seldom available in developing countries. consequently, conservationists are faced with a problem because they seldom have the time or budget for detailed, long-term population viability analysis and habitat modelling needed for setting ecologically acceptable targets for the size of conservation areas. in response to the general deficiency in time and data, burgman et al. (2001) developed a method for setting conservation targets for plant species when a limited amount of relevant information is available. these authors suggest that by using their method, a reserve system, adequate to conserve a viable population of a species, can be designed. the present study aims to apply the methodology of burgman et al. (2001) to assess whether the size of the bnr is currently large enough to conserve a viable population of b. zanguebarica. several scenarios were run to investigate different levels of human-induced impact to derive the most promising and realistic target area to conserve the species. methods top ↑ the study was undertaken in the surrounds of the bnr in the vhembe district municipality of the limpopo province (figure 1). the reserve is currently 110 ha in size. the vegetation in and around the reserve is vhavenda miombo (mucina & rutherford 2006), which is limited to a small area in the upper reaches of the mbodi river valley. several species, amongst which brachystegia spiciformis and b. zanguebarica, find their southernmost distribution within this isolated miombo vegetation unit (mucina & rutherford 2006). overall, the vegetation type is heavily degraded by overgrazing, wood-collecting, agriculture and alien invasion (mucina & rutherford 2006). figure 1: grid map of the mafukani–thengwe region showing the brackenridgea nature reserve, south africa. the mafukani population of b. zanguebarica covers an area of approximately 2500 ha (todd et al. 2004). within the reserve it is a dominant species of the b. zanguebarica – digitaria sanguinalis open scrub vegetation, with emergent trees of up to 10 m high (todd et al. 2004). the method developed by burgman et al. (2001) and modified by gaugris and van rooyen (2010) was used to set conservation goals for b. zanguebarica. this method accounts for deterministic and stochastic processes that lead to a decline in a population. the method should ideally be conducted by a panel of experts or people with local knowledge, who guide the decision process and assess the various risks considered. a brief summary of the 12-step method is provided below to guide the reader. step 1 the first step was to get a value for f, the minimum viable population size likely to persist demographic and environmental influences. this was defined by burgman et al. (2001) as the population size that faces a 0.1% probability of falling below 50 adults in the next 50 years, assuming no detrimental human effects. the f-value was obtained by applying the empirical method proposed by gaugris and van rooyen (2010) for practitioners. this was achieved by fitting the exponential function [y = aeb(x)] to the graph derived from the f-value (y) against life expectancy of known species (x) (gaugris & van rooyen 2010). a life expectancy of 150 years was assumed for b. zanguebarica.the adjusted f-value was based on available knowledge regarding the species and environmental factors against the list of 25 ecological factors (online appendix table 1), with each factor having two alternative states: one related to the species resilience and the other one to the species vulnerability (burgman et al. 2001; gaugris & van rooyen 2010). the percentage adjustment needed to the f-value was derived by calculating the ecological factor score for the species. an all positive score of 25 was assumed to need zero adjustment and an all negative -25 score was assumed to need a 100% adjustment (equal to 2f) (gaugris & van rooyen 2010). step 2 in step 2, areas experiencing similar sources and intensity of disturbance were identified. for an easy assessment of the area through expert judgement, an area of 6000 ha was mapped into 24 cells of 250 ha each (figure 1). the disturbance in each of the 24 cells was classified into one of three classes, (1) insignificant, (2) light (human activity disturbances associated with light grazing and resource harvesting) or (3) heavy (disturbances associated with building, cultivation and overgrazing). step 3 the potential b. zanguebarica habitat per cell was evaluated using knowledge from reconnaissance and fieldwork surveys. step 4 this step indicated the potential habitat that was surveyed (ha). the area surveyed in detail to establish a plant density consisted of 110 ha in the bnr. step 5 density of adults trees per ha (d) was established from 16 plots, of 500 m2 each, sampled inside the bnr. although the method allows different density values for the different disturbance regions, it is preferable to use a single density value based on the most undisturbed habitat (burgman et al. 2001), that is the bnr in this instance. individuals with a stem diameter of ≥ 100 mm were considered as mature as this is the threshold size where plants commence flowering. step 6 the preliminary minimum target area (target area a0) required for conservation was calculated as:target area a0 = adjusted f / d (in ha). [eqn 1] step 7 in this step, the proportion of land that remains in 50 years, after yearly disturbance, was estimated (s) in each cell. all activities causing disturbances that may reduce the potential habitat of b. zanguebarica species were assessed. it was assumed that these small-scale disturbances (e.g. fire) are reversible and that the species will be able to recover within the 50-year period. the reduction in potential habitat was used to calculate an adjusted target area (a1) as:a1 = a0 / s (in ha). [eqn 2] step 8 the area expected to be irreversibly damaged in the next 50 years (ci) was evaluated per cell. the remaining area (1 – ci) was used to refine the adjusted target area (a2) as:a2 = a1 / (1 – ci) (in ha). [eqn 3] step 9 compensation for expected density-reducing human related activities was achieved through adjustment of the target area per block and was expressed as ri, the estimated proportion of remaining habitat (burgman et al. 2001). the following four human related activities were considered: cultivation, grazing, fencing material and bark harvesting for medicinal material. for each of these activities, a proportion habitat remaining was calculated and the product of these proportions was used for further refinement of the target area (a3) as:a3 = a2 / ri (in ha). [eqn 4] four scenarios were assessed in order to determine which scenario could provide the best acceptable management option for b. zanguebarica. the four scenarios were as follows: scenario 1 looked at the current status of the species management, scenario 2 was when grazing was removed from the management system, scenario 3 investigated the effect of reducing all the human-related activities by half, whereas scenario 4 looked at the management system in which all the human-related activities, except bark harvesting, had been removed from the management system. step 10 identification of catastrophic events such as landslides, earthquakes and volcanic eruptions (burgman et al. 2001), that could affect the species’ potential habitat was not performed because such events are unexpected in the area. step 11 combining target areas across different regions was also not conducted, because the method was applied on the only population of b. zanguebarica that exists within south africa. step 12 the final step entailed an evaluation of the adequacy of current strategies and the setting out of objectives accounting for spatial and species constraints (burgman et al. 2001). the ratio of available to required habitat was calculated for each of the cells. results top ↑ the study results are described in order, following the abovementioned steps outlined in burgman et al. (2001). minimum viable population size based on a life expectancy of 150 years for b. zanguebarica, an f-value of 1071 was derived by applying the empirical method of gaugris and van rooyen (2010). the ecological factor score of 6 (online appendix table 1) necessitated an adjustment of 38% for an adjusted f of 1478. identification of populations with similar disturbance only three cells showed an insignificant disturbance level, whereas nine cells showed a light disturbance level and 12 showed a heavy disturbance level (online appendix table 2a). potential habitat six cells exhibited quality habitat for b. zanguebarica and small proportions of suitable habitat were found in another five cells (online appendix table 2a). potential habitat surveyed the potential habitat surveyed for a density estimate amounted to 110 ha in the bnr. density of mature individuals the mean density of mature b. zanguebarica individuals across all transects within the reserve was 61 individuals per ha. minimum target area the minimum target area or raw area for b. zanguebarica (a0 = adjusted f/density), assuming no threats, was 24 ha. proportion of land lost over the projected 50-year period, the calculation for proportion of land lost would need to include all small-scale activities that affect the population of b. zanguebarica. for the bnr, this increased the adjusted target area (a1 = a0 / s) to 43 ha (online appendix table 2a). proportion irreversibly damaged in 50 years the refinement of the adjusted target area [a2 = a1 / (1 – ci)] through irreversible damage in the next 50 years (ci) brought about a further increase in the target area to 112 ha. it was found that the main cause of these irreversible losses of habitat in the region was human settlement expansion. compensation four scenarios of expected density-reducing human-related activities (cultivation, grazing, fencing material and bark harvesting for medicinal material) were assessed. scenario 1 assessed the current status of the area and revealed that 410 ha will be needed in order to maintain a viable population of b. zanguebarica (online appendix tables 2a and 2b, cells with suitable habitat). scenario 2 was to prohibit livestock grazing from the area. it was found that removing grazing from the area can reduce the amount of land needed to conserve the species from 410 ha to 298 ha (online appendix table 2c, cells with suitable habitat). under scenario 3, the local communities were allowed to carry on with their activities, but at a reduced rate. by reducing all four human activities (cultivation, grazing, building and harvesting) by half, an area of 203 ha would be needed to conserve b. zanguebarica (online appendix table 2d, cells with suitable habitat). finally, scenario 4 assessed a scenario of removing cultivation, grazing, harvesting for fencing material, but retaining bark harvesting for medicinal purposes and found that this approach would require an area of 179 ha for a viable population of b. zanguebarica (online appendix table 2e, cells with suitable habitat). assessment of catastrophes as stated above, catastrophic events were not considered as the area has no historical records of any such catastrophes and none are anticipated for the foreseen 50-year period of evaluation. combination of targets combining target areas was not conducted in this study because there are no other populations of b. zanguebarica within south africa. evaluation of current strategies from the ratio of available-to-required habitat calculated in step 12, it was evident that cells c2, c3, d2 and e2 were the most promising for the conservation of the species. the bnr can therefore be expanded to the east along the ridgeline to obtain the required 179 ha for conservation under scenario 4. the reserve could also be expanded to the south, but would then have to cross the road (figure 1). discussion top ↑ when evaluating rare taxa, it is important to understand the distribution, biology and threats in order to devise efficient strategies for their protection (lozano & schwartz 2005; rodgers et al. 2010; wessels, freitag & van jaarsveld 1999). understanding the dynamics of the resource base is important to develop a sound management system for resource harvesting (ghimire, mckey & aumeeruddy-thomas 2005; obiri, lawes & mukolwe 2002). although the burgman et al. (2001) method relies heavily on expert opinion, it nevertheless provided a framework to systematically quantify the risk of decline or possible extinction of b. zanguebarica, taking into consideration current human activities, disturbances and the viability of the population. the method illustrated the effect of different land-use options on the size of the conservation target. from this study, it is evident that the current reserve is inadequate to ensure a viable population of the species and that additional land will have to be set aside. the size of the additional land will depend on the land use. a conservation land use (scenario 4) will require a total of 179 ha. if the human-related activities are reduced by 50% (scenario 3), then a total of 203 ha will be needed; however, if only grazing is prohibited (scenario 2) then a total of 298 ha will be needed. if the human-related activities are allowed to continue unabated the total area needed to conserve the species increases to 410 ha. amongst the scenarios assessed, scenario 4 corresponds to the current conservation option within the bnr where harvesting is occurring, but no cultivation, grazing or other harvesting activities are allowed. selecting this option would necessitate another 69 ha to be incorporated into the reserve. an area adjoining the reserve (d2) should be selected to avoid the need for corridors between conservation areas. by selecting an area where no cultivation is practiced this would imply that 69 ha of grazing land will have to be sacrificed for the enlargement of the reserve. the use of b. zanguebarica for fences will have to be prohibited entirely, but the controlled harvesting for medicinal purposes can continue because of the high demand for the bark. furthermore, the enlargement of the reserve would be supported by the fact that the vhembe region, in which the bnr is situated, is a unesco-declared biosphere reserve. should increasing the size of the reserve not be a viable option, then human-related activities will need to be reduced in the area surrounding the reserve. however, the challenge will arise in the monitoring of the level of utilisation, which is to be reduced by half from the current level in scenario 3. prohibiting only grazing (scenario 2) might be a more realistic option, because this can be more easily monitored than bark harvesting. however, in this case the size of the additional area will have to be almost doubled. excluding herbivores from an area has the further advantage that it reduces trampling and promotes seedling establishment and the amount of dead plant material that acts as protective mulch (angassa & oba 2010). although the creation of a protected area facilitates the conservation of medicinal plant species by restricting access and extractive use (mcgeoch, gordon & schmitt 2008), it has also been found that whenever the economic value of a natural resource carries more weight than the cultural value, traditional management of such a resource can fail to guarantee its sustainability (saidi & tshipala-ramatshimbila 2006). strict control over the volume of bark harvested will have to be exercised and measures will have to be put in place to prevent illegal bark harvesting. conclusion top ↑ the study revealed that the current reserve size is not sufficient for conservation of a viable population of b. zanguebarica. the study identified potential areas, which can be utilised for the expansion of the reserve. it also identified the size of the area to be set aside under different management options. the most promising option appears to enlarge the reserve by adding at least 70 ha of adjacent land. the conservation of a viable b. zanguebarica population can only be achieved through a reduction of human activities that have a negative impact on the population dynamics of the species and by strict control over the volume of bark harvested. acknowledgements top ↑ the authors gratefully acknowledge the assistance of mr magwede khathutshelo and mr siaga moses with data collection. staff members at the brackenridgea nature reserve, especially mr rasivhaga, mr maluta, and mr manenshe are thanked for their unconditional support. dr nethengwe from the department of geography at the university of venda is also acknowledged for helping out with geographic information system during the production of the area map. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions m.p.t. (university of venda) was the project leader who carried out the experiments, analysis and preparation of the manuscript. m.w.v.r. (university of pretoria) designed the experiments, analysed the data and proofread the manuscript. j.y.g. (university of pretoria) provided most of the calculations during data analysis and also helped in proofreading of the manuscript. references top ↑ angassa, a. & oba, g., 2010, ‘effects of grazing pressure, age of enclosures and seasonality on bush cover dynamics and vegetation composition in southern ethiopia’, journal of arid environments 74, 111–120. http://dx.doi.org/10.1016/j.jaridenv.2009.07.015 beissinger, s.r. & mccullough, d.r., 2002, population viability analysis, the university of chicago press, chicago. burgman, m.a., possingham, h.p., lynch, j.j., keith, d.a., mccarthy, m.a., hopper, s.d. et al., 2001, ‘a method for setting the size of plant conservation target areas’, conservation biology 15, 603–616. http://dx.doi.org/10.1046/j.1523-1739.2001.015003603.x cabeza, m. & van teeffelen, a.j.a., 2009, ‘strategies of reserve selection’, encyclopedia of life sciences, john wiley & sons, chichester. http://dx.doi.org/10.1002/9780470015902.a0021224 gaugris, j.y. & van rooyen, m.w., 2010, ‘evaluating the adequacy of reserves in the tembe–tshanini complex: a case study in maputaland, south africa’, oryx 44, 399–410. http://dx.doi.org/10.1017/s0030605310000438 ghimire, s.k., mckey, d. & aumeeruddy-thomas, y., 2005, ‘conservation of himalayan medicinal plants: harvesting patterns and ecology of two threatened species, nardostachys grandiflora dc and neopicrorhiza scrophulariiflora (pennell) hong’, biological conservation 124, 463–475. http://dx.doi.org/10.1016/j.biocon.2005.02.005 lozano, f.d. & schwartz, m.w., 2005, ‘patterns of rarity and taxonomic group size in plants’, biological conservation 126, 146–154. mcgeoch, l., gordon, i. & schmitt, j., 2008, ‘impacts of land use, anthropogenic disturbance, and harvesting on an african medicinal liana’, biological conservation 141, 2218–2229. http://dx.doi.org/10.1016/j.biocon.2008.06.021 menges, e.s., 2000, ‘population viability analysis in plants: challenges and opportunities’, trends in ecology and evolution 15, 51–56. http://dx.doi.org/10.1016/s0169-5347(99)01763-2 mucina, l. & rutherford, m.c. (eds.), 2006, ‘the vegetation of south africa, lesotho and swaziland’, strelitzia 19, south african national biodiversity institute, pretoria. netshiungani, e.n. & van wyk, a.e., 1980, ‘mutavhatsindi – mysterious plant from venda’, veld and flora september, 87–90. obiri, j., lawes, m. & mukolwe, m., 2002, ‘the dynamics and sustainable use of high value tree species of the coastal pondoland forests of the eastern cape province, south africa’, forest ecology and management 166, 131–148. http://dx.doi.org/10.1016/s0378-1127(01)00665-x pfab, m.f. & witkowski, e.t.f., 2000, ‘a simple population viability analysis of the critically endangered euphorbia clivicola r.a. dyer under four management scenarios’, biological conservation 96, 263–270. http://dx.doi.org/10.1016/s0006-3207(00)00088-4 rodgers, h.m., glew, l., honzak, m. & hudson, m.d., 2010, ‘prioritizing key biodiversity areas in madagascar by including data on human pressure and ecosystem services’, landscape and urban planning 96, 48–56. http://dx.doi.org/10.1016/j.landurbplan.2010.02.002 saidi, t.a. & tshipala-ramatshimbila, t.v., 2006, ‘ecology and management of a remnant of brachystegia spiciformis (miombo) woodland in north eastern soutpansberg, limpopo province’, south african geographical journal 88, 205–212. http://dx.doi.org/10.1080/03736245.2006.9713862 todd, c.b., khorommbi, k., van der waal, b.c. & weisser, p.j., 2004, ‘conservation of woodland biodiversity: a complementary traditional approach and western approach towards protecting brackenridgea zanguebarica’, in m.j. lawes, h.a.c. eeley, c.m. shackleton & b.g.s. geach (eds.), indigenous forests and woodlands in south africa – policy, people and practice, pp. 737–750, university of kwazulu-natal press, durban. wessels, k.j., freitag, s. & van jaarsveld, a.s., 1999, ‘the use of land facets as biodiversity surrogates during reserve selection at a local scale’, biological conservation 89, 21–38. http://dx.doi.org/10.1016/s0006-3207(98)00133-5 williams, v.l. & raimondo, d., 2008, brackenridgea zanguebarica oliv. national assessment: red list of south african plants version 2011.1, viewed 16 april 2012, from http://redlist.sanbi.org/species.php?species=2924-2 article information authors: simon d. capon1 alison j. leslie1 bruce clegg2 affiliations: 1department of conservation ecology and entomology, stellenbosch university, south africa 2research department, malilangwe wildlife reserve, zimbabwe correspondence to: alison leslie postal address: private bag x1, matieland 7602, south africa dates: received: 20 apr. 2012 accepted: 17 feb. 2013 published: 05 apr. 2013 how to cite this article: capon, s.d., leslie, a.j. & clegg, b., 2013, ‘the use of population viability analysis to identify possible factors contributing to the decline of a rare ungulate population in south-eastern zimbabwe’, koedoe 55(1), art. #1081, 9 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1081 copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. the use of population viability analysis to identify possible factors contributing to the decline of a rare ungulate population in south-eastern zimbabwe in this original research... open access • abstract • introduction • methods    • study site    • population data collection    • age class mortality rates    • the population viability analysis model       • scenario settings       • reproduction system, age of first reproduction and longevity       • reproduction rate       • catastrophic events       • initial population       • carrying capacity       • supplementation and harvest    • model scenarios    • sensitivity analysis • results    • demographic parameter estimation, population size and reproductive rate    • replicating the historical population trend    • future population viability    • population sensitivity • ethical considerations • trustworthiness • discussion    • management implications • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ populations that are vulnerable to decline are of particular concern to wildlife managers and uncovering the mechanisms responsible for downward trends is a crucial step towards developing future viable populations. the aims of this study were to better understand the mechanisms behind the historic decline of the sable antelope, hippotragus niger, population at the malilangwe wildlife reserve (mwr), to assess its future viability and to use this analysis to determine key areas of breakdown in population growth and link these to potential limiting factors. vortex, a population viability model was used to assess the future viability of the sable antelope population and a sensitivity analysis was applied to identify the key areas of breakdown in growth. the sable population is currently viable, but remains highly vulnerable to changes in adult female survival, a factor which had the greatest influence on overall population fitness. lion predation, impacting on the adult segment of the population, appeared to be the main factor responsible for the historic decline at the mwr. conservation implications: sable generally occur at low densities in the lowveld region of zimbabwe and, as such, populations are vulnerable to increases in mortality rates. the role of lions in driving the decline at the mwr suggests a need to control their numbers and develop prey refuges through improved management of artificial water. introduction top ↑ crisis management is a common element of conservation biology (begon, harper & townsend 1996) because small or declining populations often require urgent action to prevent extinctions. small or declining populations are particularly vulnerable to extirpation because they are governed by a high level of insecurity resulting from demographic and environmental stochasticity and spatial uncertainty (begon et al. 1996). grant and van der walt (2000) explain that at low densities populations are particularly at risk of extinction and any mortality in the population can be disastrous. furthermore, it has been shown that at low densities stochastic events can wipe out small and declining populations altogether (ginsberg, mace & albon 1995).past studies have focused on identifying factors that influence population abundance, but less effort has focused on determining how age class survival rates may affect population dynamics (gaillard, festa-bianchet & yoccoz 1998). studies of population dynamics can often give insights into the possible causes of decline by identifying areas of breakdown in recruitment. they can also provide a better understanding of population trends and provide vital rates crucial to accurate viability analyses. population viability analysis (pva) is a useful tool for predicting future population sizes and assessing viability under a range of management scenarios (brook et al. 2000). pva makes use of real life-history data of a species or population to train the model and project the future population utilising stochastic variation in demography (boyce 1992; brook et al. 2000). owing to their long generation time, large mammals were often excluded from population ecology studies because of the length of time taken to reveal interesting results (gaillard et al. 1998). gaillard et al. (1998) go on to suggest, however, that large mammals make ideal subjects for population studies because they can readily be distinguished into discrete age classes. this allows for accurate estimation of age-specific vital rates, an important component of any pva. the low density of occurrence and strict seasonality of reproduction in sable antelope, hippotragus niger, makes them ideal subjects for population studies and pva may be a very useful tool in assessing the viability of isolated populations, such as that on the malilangwe wildlife reserve (mwr). the long-term growth rate of a population can be used as a direct measure of fitness and, as such, a sensitivity analysis assessing the sensitivity of the long-term growth rate to changes in vital rates can provide important clues as to the areas most susceptible to breakdown in the growth of a population. the sable population at the mwr in the south-eastern lowveld of zimbabwe declined from over 200 animals in 1994 to only 62 by 2005. this was followed by a period of relative recovery where the population increased to 77 by 2010, but the population still remains highly vulnerable. this declining trend (1994–2005) is not unique to the mwr and is similar to that reported for sable in the kruger national park (knp), south africa. grant and van der walt (2000) note that the sable population in the knp declined from 2240 in 1986 to only 505 by 1999. a number of hypotheses have been put forward to explain the decline, including: competition with other herbivores (harrington et al. 1999), increased predation (grant & van der walt 2000; owen-smith, mason & ogutu 2005), rainfall variation (owen-smith et al. 2005), disease (nijhof et al. 2005; oosthuizen et al. 2008) and habitat loss (grant & van der walt 2000). an understanding of the population dynamics of sable on the mwr may give insights into the factors responsible for the decline and could, in turn, help to formulate a mitigating management strategy. the main objectives of this study were, (1) to gain an understanding of the dynamics of the sable population at the mwr and assess its future viability, (2) to identify the possible factors leading to the decline of the species and (3) to discuss the implications thereof. we used a pva to broadly assess the viability of the sable population under different scenarios and further applied the pva to a sensitivity analysis of the overall fitness of the sable population. methods top ↑ study site the mwr (39 378 ha) is located in the south-eastern lowveld of zimbabwe. the reserve is bordered by the gonarezhou national park to the south, the hippo valley game section to the west and by communal land to the north and east (figure 1). the area has three distinct seasons: a hot, wet season from november to march, a cool, dry season from may to august and a hot, dry season from september to october. the long-term average annual rainfall for the area is 562.22 mm ± 193.81 mm, measured at malilangwe headquarters from 1951 to 2009 (figure 2). between march 2009 and january 2011, the study period, the mwr received 785.40 mm of rainfall and the average monthly maximum and minimum temperatures were 30.50 °c ± 3.20 °c and 17.60 °c ± 3.88 °c, respectively. the vegetation is dominated by a variety of open colophospermum mopane woodland, with the malilangwe sandstone range being characterised mostly by brachystegia woodlands. the property has been classified into 38 vegetation types based on the species composition of both the woody and herbaceous layers, the structure of the woody component and the ecological factors that influence the plant communities (clegg 2010). population data collection data on the population structure of sable antelope at the mwr were collected between march 2009 and january 2011. to facilitate this, an adult female from four of the sable herds at the mwr was immobilised by a licensed professional using a combination of 6.0 mg m99 and 1.0 mg medetomidine and fitted with a gps–vhf collar (qauntum 5000, weight: 700 g; telemetry solutions, concord, california). once the collar had been securely fitted, the anaesthesia was reversed using 18.0 mg m5050 and 2.5 mg antisedan and, in all cases, the collared animals moved off unharmed, rejoining their respective herds within 1–4 h. a fifth herd, the only other herd on the mwr, was monitored opportunistically over the study period, allowing us to ensure all sable individuals, except for itinerant males, were accounted for. although not fitted with a collar, this herd was encountered on average every 2–3 weeks and from these encounters a clear demographic structure of the herd was gathered. the vhf signal emitted by the collars was used to locate each herd every 3 days. on locating a herd, a total count was made, followed, where possible, by an age-sex specific count. individuals were assigned to either male adult (3+ years), female adult (3+ years), sub-adult male (2–3 years), sub-adult female (2–3 years), yearling (1–2 years) and juvenile (0–1 year) age classes, according to grobler (1980a), who suggested the use of a combination of horn characteristics and physical appearance for classification. by combining data from the counts of each herd, changes in the number of animals in each age class were recorded over the study period. in most cases, all four collared herds were seen in 1 day and the fifth herd was encountered at least once every fortnight. for all the female age-classes and the male juvenile, yearling and adult age-classes, if an individual disappeared and did not return before the next breeding cycle, that individual was considered to be dead. consistent monitoring of sub-adult males was more difficult as they do not remain with the herds until adulthood; accurate observations were therefore not possible. it was not considered necessary to recognise each individual sable because the entire population was being monitored over the study period. during the wet season, sable herds often split up into smaller units, but in nearly all cases reliable counts were achieved at least twice per month. this provided a near-continuous data set for the entire study period, covering two calving seasons. these demographic data were then used to estimate vital rates and develop a baseline for the pva. figure 1: map showing the location of malilangwe wildlife reserve. figure 2: the historical trend in the sable population on the malilangwe wildlife reserve between 2000 and 2010. evident are two distinct trends in the population from 2000 to 2005 and from 2005 to 2010. age class mortality rates age-specific mortality rates for sable were calculated for each breeding cycle of the study period using population data gathered during the fieldwork phase of the study. the start and end of the breeding cycle was taken as april and january, respectively, because february–march is the calving season for sable at the mwr. this meant that by april a new crop of calves would emerge pushing existing animals into the next age-class. data derived from the life tables were used to calculate age-class mortality rates using the formula developed from methods described in caughley (1966):qс = [{nχ nχ+1} / nχ × 100] [eqn 1] where qс is the mortality rate of age class c, nχ is the number of individuals in age class c at time χ and nχ+1 is the number of individuals still alive in age class c at time χ+1. this method of deriving mortality rates was employed for all but the juvenile age-class. the mortality rate for the juvenile age-class was calculated by dividing the number of calves alive at the end of the breeding cycle by 0.96 times the number of adult females in the herd at the beginning of the cycle and subtracting this value from one. this figure was then multiplied by 100 to convert the mortality rate into a percentage. the value of 0.96 was derived from observations made from sable antelope at the mwr during the 2009 and 2010 breeding seasons, where 96% of adult females fell pregnant and were assumed to give birth to a healthy calf. pregnancy was estimated somewhat subjectively following grobler (1980b), who used the shape of the stomach and presence of a bulging udder to determine pregnancy. in all but the adult age-class, no distinction in mortality rate was made between male and female sable. at the younger stages, it was not considered necessary as both males and females stay within the herd and in theory face the same risk of mortality. determining a mortality rate for sub-adult males was more difficult because they often leave the herd making it impossible to monitor them regularly. the same mortality estimates developed for sub-adult female sable were therefore used for sub-adult males, possibly a rather conservative estimate for the part of sub-adult males. the population viability analysis model for the analysis and simulation of the pva, vortex version 9.99b was used (lacy, borbat & pollack 2009). the model was developed to stochastically simulate the risk of extinction faced by a species. the vortex model takes into account a description of the species reproduction system and reproductive rates, a specified age structure, age-specific mortality rates, catastrophic events, demographic and environmental stochasticity, density dependence and options for the harvest and augmentation of the population.assessing the viability of the sable population at the mwr using vortex involved a stepwise process. the first stage entailed calibrating the model and replicating the historical trend of the population. an analysis of the historical population trend revealed two distinct periods. the first was the period of population decline from 2000 to 2005 and the second, the period of population recovery from 2005 to 2010. accordingly, the calibration of the population model was divided into these two distinct periods. the second stage of the process involved projecting the population forward 100 years using data ratified during the calibration of the model for the periods 2000–2005 and 2005–2010. baseline settings used in the model are described in the following subsections. scenario settings although five separate sable herds exist on the mwr, the reserve is essentially a closed system and therefore the mwr sable were described as a single population. during the calibration process, the pva was simulated 500 times over two 5-year periods, from 2000 to 2005 and from 2005 to 2010. following calibration, the model was again simulated 500 times for a period of 100 years in each of the subsequent scenarios. reproduction system, age of first reproduction and longevity age of first reproduction in the vortex model was set at 3 and 6 years old for female and male sable, respectively. based on personal observations at the mwr and discussions with other experts, under natural conditions of predation, disease and environmental variability, the maximum age of reproduction in sable was set at 12 years of age. reproduction rate in the study, it was assumed from field observations, that 96% of the adult females in the population would fall pregnant each year and each would produce a single calf. from observations made over two calving seasons at the mwr, the calf sex ratio at birth was estimated at 1:1. a breeding population of 29% of males was used and this was determined by dividing the number of dominant, territorial bulls seen to be mating with females by the number of adult bulls in the population. catastrophic events catastrophic events by their very nature are unpredictable, usually with devastating consequences, making managing for a catastrophe almost impossible. the purpose of this study was not to create a generic population model for sable but to investigate specifically what drove the decline in sable at the mwr using a population viability tool. as no catastrophic events were encountered over the period under review, they were excluded from the model. initial population data on the size of the sable population (figure 2) in 2000 and 2005 on the mwr used in calibrating the model were taken from the annual aerial census data conducted by helicopter between 2000 and 2010 (goodman 2009). as the age-specific population structure was unknown in both 2000 and 2005, the model was run using a stable age distribution in both scenarios to test the baseline data and calibrate the model. once calibrated, the model was simulated using the age specific population structure (table 1) determined at the beginning of the study period in march 2009. carrying capacity a value of 200 was used in the pva as a measure of carrying capacity (k) on the mwr and was taken from historical population estimates made by booth (1980). it was not considered essential to use an entirely precise figure for k, as the purpose of this study was to assess the risk of extinction of the current population and to assess the trend of the population under a number of different scenarios, rather than being concerned with actual figures for final population size after 100 years. supplementation and harvest the release of a further 30 sable took place in 2011, the first supplementation of the population since 1994. the population model was run using current population estimates, as well as testing the effect of boosting the population with a further 30 individuals (10 adult females, 5 sub-adult males, 5 sub-adult females, 5 juvenile males and 5 juvenile females). no harvest of the sable population was included in the pva.in the vortex model, standard deviation in vital rates or carrying capacity as a result of environmental variance was stated as zero in all cases. although it was recognised that environmental variation may well cause year to year variation in survival, no data were available on the actual effect of this variation. it was therefore decided not to include this effect, but rather to work with a relatively simplified version of the model. table 2 shows a summary of those baseline data input for each scenario of the vortex model. model scenarios in scenario 1, the current conditions using vital rates recorded over the study period were simulated and these were tested against the trend observed from 2005 to 2010. historical data from scout patrol reports on sable carcasses were then used to estimate a factoring for the adjustment of current mortality estimates for the second scenario. five times as many sable carcasses were found by scouts on patrol during the period 2000 to 2005 than during 2005 to 2010, despite a relatively even patrol effort between the periods (two-sample t(5) = -1.39, p = 0.22). of the carcasses discovered, 88% were from the adult and sub-adult age classes, hence mortality rates for only the adult and sub-adult age classes were increased by a factor of five. it was impossible to estimate a factoring for the juvenile age class as juvenile carcasses are notoriously difficult to find, often being completely eaten leaving no trace (owen-smith & mills 2008). the new vital rates developed from the factoring process were then tested in the model against the observed trend from 2000 to 2005. in scenarios 3a and 4a, the population was projected forward by 100 years using the vital rates derived for scenarios 1 and 2, respectively. additionally, the effect of augmenting the sable population with 30 individuals was simulated in scenarios 3b and 4b. table 1: demographic structure of the sable population on the malilangwe wildlife reserve in march 2009. table 2: summary of baseline data used in the vortex model under various scenarios. sensitivity analysis in the sensitivity analysis, the vortex model was run varying survival of each age class in turn, whilst keeping all others at their average value over the study period (2009–2011). the percentage change in survival rate for each age class was plotted against the vortex output value of mean long-term population growth rate (r). this allowed the determination of the sensitivity of the long-term population growth rate, and hence overall fitness, to changes in the survival rates of each age class. results top ↑ demographic parameter estimation, population size and reproductive rate demographic parameters were calculated from the field data collected over the entire study period (table 2). in general, mortality rates (mr) were considered low for all but the juvenile age class (mr = 57%). adult female sable had the lowest mortality rate of all the age classes (mr = 4%), followed by the yearling age class (mr = 6%). adult male mortality rate was more than double that of adult females at 9% and this was reflected in the sable population structure at the mwr. from 2000 to 2005, the sable population declined from 110 individuals to only 62 (figure 2). the population then began to stabilise and had grown to 77 individuals by 2010. during the 2009–2010 breeding season, 21 out of 22 adult females were pregnant and due to give birth in march 2010, but by april 2010 there were only nine surviving calves. this suggests that calves were either still-born or died during the hiding phase after birth. notably though, three calves, all from the same herd, were recorded alive soon after birth and seemed healthy, but by may 2010 this herd had no calves accompanying them, indicating that many more calves may have been born healthy but perished during the calf hiding phase. replicating the historical population trend results from the overall training and testing of the model proved highly comparable with that of the observed population trend (figure 3). during the 5-year period, 2000–2005, the modelled population declined from a total of 110 individuals to only 61 by 2005. this was similar to observed trends which showed a decline from the same initial population to only 62 individuals over the same period. vortex predicted a growth rate of -0.1209 (s.d. = 0.0777) and a probability of extinction within the 5 years of zero.the modelled population projected from 2005 to 2010 tracked the observed trend closely (figure 4), increasing from an initial population of 62 individuals to 79 by 2010. similarly, the observed population increased from 62 to 77 over the same period, falling well within the 95% confidence limits set by the modelled population. the modelled and observed populations increased at average rates of 4.80% ± 5.02% and 4.90% ± 12.6% per year, respectively. future population viability using the vital rates estimated for the period 2000–2005, the modelled population is predicted to decline sharply at an overall rate of 16.7% per annum (figure 5), with extinction predicted after just 18 years. augmenting the population with the further 30 sable will not curb this trend and the sable population is predicted to remain extant on the mwr for only the next 22 years. the addition of the 30 sable to the population slows the decline to 16.1% per annum, but there remains no chance of recovery for the population under these circumstances.using the vital rates estimated for the period 2005–2010, the sable population is predicted to increase steadily at a rate of 4.9% per year, quickly stabilising at k (figure 5). in this case, the population reaches k after only 30 years and the model predicts no chance of extinction within 100 years. under these conditions, supplementation of the population with a further 30 sable causes the population to increase again at a rate of 4.9% per annum, stabilising around k after just 20 years. the model forecasts no chance of extinction within the 100 year period. figure 3: modelled versus observed trends in the sable population on the malilangwe wildlife reserve from 2000 to 2005. figure 4: modelled versus observed trends in the sable population on the malilangwe wildlife reserve from 2005 to 2010. population sensitivity the sensitivity of the population growth rate to changes in survival of the different age classes is shown in figure 6. proportional changes to adult female survival had the greatest effect on the average long-term population growth rate. as little as a 10% reduction in survival of adult females led to a negative growth rate (r = -0.007) in the population. by comparison, a reduction of 25% in the survival rate of the other age classes still resulted in positive growth in the overall population. though marginally so, sub-adult survival had the next greatest effect on the overall population growth rate, with a 25% reduction in survival reducing population growth to 0.006 per year. adult male survival had the least impact, if any, on the overall growth rate of the population. ethical considerations top ↑ all work was carried out under the guidance of the malilangwe trust and its code of conduct. in all cases, licensed professionals and veterinarians were used to perform immobilisations and any risks to the animals were kept to an absolute minimum (see ‘acknowledgements’). trustworthiness top ↑ all data were collected using standard scientific procedures and, as such, the results are considered to be entirely objective and accurate. unfortunately, the nature of ecological work means that some level of inconsistency is achieved, in that the results may not be directly repeatable if the study were performed over another time period because of the nature of variation in natural systems. however, this does not detract from the findings of the research in any way whatsoever. discussion top ↑ there have been heated debates in the literature regarding the value and accuracy of pva models for predicting future population trends (coulson et al. 2001; ellner et al. 2002; taylor 1995). in spite of this, pva has been employed widely as a tool for assessing the vulnerability of threatened populations and for evaluating possible options for their recovery (brook et al. 2000; cancino, rodriguez-estrella & miller 2010; gusset et al. 2009; kelly & durant 2000). the usefulness of a pva model, however, depends entirely on the quality and accuracy of the baseline data (coulson et al. 2001). the probability of extinction of a population is influenced by both uncertainties in the natural environment, as well as uncertainty in the estimation of parameters used in the model (taylor 1995). although environmental variation plays a role in varying the predictions of the model, uncertainty in the estimation of input parameters had a much greater effect on the overall output (taylor 1995). mcloughlin and owen-smith (2003) state that models that ignore predation pressure and take into account only demographic and environmental stochasticity may greatly underestimate extinction risk. in this study, the data used were collected intensively over a 2-year period to derive vital rates for the population. these rates were tested by comparing the modelled trend to the observed trend in the population over the past 5 years. the observed population fluctuated above and below the modelled population, which showed a steady increase from 2005 to 2010, a result of taking an average of 500 iterations. the short duration of the study meant that the effect of environmental stochasticity was essentially excluded from the model. however, testing of the output against historical data lent strong support to the model parameters despite the limited period of observation. figure 5: mean sable population trend on the malilangwe wildlife reserve over 100 years, modelled using estimates of vital rates taken from 2000 to 2005 and 2005 to 2010 (3a and 4a). also shown are the effects of augmenting the population with 30 sable (3b and 4b). figure 6: sensitivity of the population growth rate to changes in survival rate of different age classes of sable on the malilangwe wildlife reserve. replicating the declining trend of the sable population from 2000 to 2005 proved more challenging as it was clear that, under current conditions, the sable population on the mwr was predicted to increase. historical data on the number of sable carcasses found by scouts on patrol clearly indicated a period of considerably higher mortality in sable from 2000 to 2005 (table 3), particularly amongst the older age classes. this period coincides with a period of markedly higher lion numbers at the mwr (figure 7). of the total number of sable carcasses found from 2000 to 2005, 70% were attributed to lion predation. lions show a clear preference for medium-bodied to large-bodied prey (hayward & kerley 2005) and this was evident in the fact that 88% of sable killed by lions between 2000 and 2010 fell into the adult and sub-adult age classes. this fact further justified the factoring of the vital rates for the 2000–2005 period. the output from the model followed the observed trend well, again lending support for the estimates of the vital rates for the period 2000–2005. the rate of decline over this period was -0.1209 (s.d. = 0.0777), markedly higher than that reported by nicholls et al. (1996) for the knp between 1983 and 1992. under the current conditions, the sable population on the mwr is predicted to remain viable, steadily increasing to carrying capacity over the next 30 years. however, the sensitivity analysis shows that even a small change in adult female survival (10%) could have disastrous effects. even more alarming is that a 10% reduction in survival of adult females equates to a total loss of only four adult female sable per year. this means that the sable population on the mwr, although currently stable, remains highly vulnerable to changes in predation pressure, environmental and demographic stochasticity and management interventions. similarly, sable antelope in the knp are at serious risk of extinction within the next 100 years (nicholls et al. 1996) and owen-smith et al. (2005) attribute the decline of the species there to increased predation by lions. on the mwr, evidence points to a similar pattern where increased lion numbers were primarily driving the decline of the species until 2005 (figure 7). in 2005, emigration of two lion prides effectively halved the lion population (figure 7), following an outbreak of anthrax that devastated prey populations in 2004 (clegg et al. 2007). since then, the relatively high rate of juvenile mortality has prevented the sable population from growing at its full potential. figure 7: estimates of lion population on the malilangwe wildlife reserve from 2000 to 2010, taken from call ups, spoor counts and known pride numbers. table 3: causes of death of sable carcasses discovered by the malilangwe wildlife reserve scout force on their daily patrols between 2000 and 2010. the long-term growth rate of a population can be used as a direct measure of the fitness of that population (ranta, lundberg & kaitala 2006) and, as such, changes in the demographic components within the population can have direct impacts on its overall viability. sensitivity analyses can determine which vital rates have the greatest impact on overall population growth and thus fitness. examining the results from the sensitivity analysis allows one not only to determine which demographic parameters have the greatest impact on overall population growth, but to assess the effect of variability or uncertainty in these parameters. the finding that a change in adult female survival has the greatest effect on overall population growth rate is not unusual and has been documented for other taxa (gaillard et al. 2000; saether & bakke 2000). the extent of this sensitivity in relation to that of other age classes indicates that the historic decline of the species on the mwr must have been caused by, inter alia, an increase in mortality in the adult female age class. this observation again lends support to the theory that increased predation, particularly by lions, was responsible for the decline of the species. in general, gaillard et al. (1998) state that adult survival amongst large ungulates is always less variable than juvenile mortality and attribute this to the fact that recruitment is often the foremost target of limiting factors, both density-dependent and independent. highly variable juvenile mortality has been linked to population declines (gaillard et al. 1998) and owen-smith and mason (2005) state that variations in juvenile survival could be a key factor responsible for fluctuations in population numbers. on the mwr, it seems unlikely that juvenile mortality alone could be responsible for the decline of the species from 2000 to 2005, based on the fact that a reduction in juvenile survival of as much as 25% still led to an overall positive growth rate in the population. it is possible to suggest, however, that variable juvenile mortality potentially causes short-term fluctuations in the population growth rate and increased mortality targeting the adult segment of the population is the driving factor in long-term sharp declines. management implications using baseline data designed to simulate conditions observed from 2000 to 2005, the pva model predicted a local extinction of the sable population on the mwr within the next 20 years. the augmentation of the population under these circumstances does little to curb the trend and management would need to look at ways of reducing the rate of mortality in the older age classes. this is similarly the case with roan antelope (hippotragus equinus) in the knp, where predation risk remains high for adult roan and augmentation of the free-range population is seen as being a wasteful option at this stage (mcloughlin & owen-smith 2003). in the long-term, manipulation of water points on the mwr to reduce the density of other herbivores and thus predators within the range of the sable herds may be a feasible option. however, managers within the knp have attempted this with limited success (mcloughlin & owen-smith 2003). mcloughlin and owen-smith (2003) suggest this could be because of only a partial emigration by predators from these areas. these authors warn that reducing the density of other herbivores in these areas may leave predators more reliant on the low density ungulates, with devastating consequences. a further complication arises because water points are an important resource for many other species, as well as being a focus of ecotourists on the mwr. therefore, with the limited number of alternatives, some control of predators, particularly lions, in combination with a review of artificial water management practices may be necessary to reduce the predation risk faced by the sable population. the management implications of controlling predator numbers have been poorly studied, however, so investigations into the methods and effects of predator control are required. conclusion top ↑ the sable population on the mwr is vulnerable to declines. adult female mortality showed the greatest sensitivity and indicated that the decline of sable on the mwr was driven by, inter alia, an increase in mortality in the adult female segment of the population. the results suggest that predation was the main driver of the historic decline on the mwr and future declines could be prevented through reducing the risk of predation on the sable population. these findings underpin the importance of effective management of predator populations in small-enclosed reserves, ideally achieved through better management of artificial water distribution. we recommend that future research should focus on the management of artificial water and its implications for predator numbers and the development of predator refuges. acknowledgements top ↑ this study was carried out as a collaboration between the department of conservation ecology and entomology at the university of stellenbosch and the malilangwe wildlife research department. the study was conducted under the ethical guidance of the malilangwe trust and its board of trustees and licensed professionals were used where necessary in all cases. we would like to thank the malilangwe trust for providing us with the resources and funding needed to make this study possible. several people are thanked for assisting with fieldwork, in particular: c. wenham, b. fivaz, p. chivambu, k. mahuhle, p. chitimela and g. coombes. competing interests the authors declare they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions s.d.c. (stellenbosch university) was the primary field researcher and msc student on whose dissertation this article is based. a.j.l. (stellenbosch university) was the project leader and student’s supervisor, whilst b.c. (malilangwe wildlife reserve) was the on-site ecologist an d co-supervisor. all three authors were responsible for the experimental and research design and production of the final manuscript. references top ↑ begon, m., harper, j.l. & townsend, c.r., 1996, ecology: individuals, populations and communities, blackwell, oxford. http://dx.doi.org/10.1002/9781444313765booth, v.r., 1980, ‘a study on lichtenstein’s hartebeest, alcelaphus lichtensteini (peters 1852) and its habitat in south-eastern rhodesia’, msc thesis, university of pretoria. boyce, m.s., 1992, ‘population viability analysis’, annual review of ecology and systematics 23, 481–506. http://dx.doi.org/10.1146/annurev.es.23.110192.002405 brook, b.w., o’grady, j.j., chapman, a.p., burgman, m.a., akçakaya, h.r. & frankham, r., 2000, ‘predictive accuracy of population viability analysis in conservation biology’, nature 404, 385–387. http://dx.doi.org/10.1038/35006050, pmid:10746724 cancino, j., rodriguez-estrella, r. & miller, p., 2010, ‘using a population viability analysis for management recommendations of the endangered endemic peninsular pronghorn’, acta zoologica mexicana (n.s.) 26, 173–189. caughley, g., 1966, ‘mortality patterns in mammals’, ecology 47, 906–917. http://dx.doi.org/10.2307/1935638 clegg, b.w., 2010, ‘habitat and diet selection by the african elephant at the landscape level: a functional integration of multi-scale foraging processes’, phd thesis, university of the witwatersrand. clegg, b.w., 2011, ‘lion population estimates for malilangwe wildlife reserve between 2000 and 2010’, unpublished internal report for the malilangwe trust, zimbabwe. clegg, s.b., turnbull, p.c.b., foggin, c.m. & lindeque, p.m., 2007, ‘massive outbreak of anthrax in wildlife in the malilangwe wildlife reserve, zimbabwe’, veterinary record 160, 113–118. http://dx.doi.org/10.1136/vr.160.4.113, pmid:17259452 coulson, t., mace, g.m., hudson, e. & possingham, h.p., 2001, ‘the use and abuse of population viability analysis’, trends in ecology and evolution 16, 219–221. http://dx.doi.org/10.1016/s0169-5347(01)02137-1 ellner, s.p., fieberg, j., ludwig, d. & wilcox, c., 2002, ‘precision of population viability analysis’, conservation biology 16, 258–261. http://dx.doi.org/10.1046/j.1523-1739.2002.00553.x gaillard, j.m., festa-bianchet, m. & yoccoz, n.g., 1998, ‘population dynamics of large herbivores: variable recruitment with constant adult survival’, trends in ecology and evolution 13, 58–63. http://dx.doi.org/10.1016/s0169-5347(97)01237-8 gaillard, j.m., festa-bianchet, m., yoccoz, n.g., loison, a. & toigo, c., 2000, ‘temporal variation in fitness components and population dynamics of large herbivores’, annual review of ecology and systematics 31, 367–393. http://dx.doi.org/10.1146/annurev.ecolsys.31.1.367 ginsberg, j.r., mace, g.m. & albon, s., 1995, ‘local extinction in a small and declining population: wild dogs in the serengeti’, proceedings: biological sciences 262, 221–228. http://dx.doi.org/10.1098/rspb.1995.0199, pmid:8524914 goodman, p.s., 2009, ‘large mammal population estimates for malilangwe estate’, unpublished report for the malilangwe trust, zimbabwe. grant, c.c. & van der walt, j.l., 2000, ‘towards an adaptive management approach for the conservation of rare antelope in the kruger national park: outcome of a workshop held in may 2000’, koedoe 43, 103–112. http://dx.doi.org/10.4102/koedoe.v43i2.202 grobler, j.h., 1980a, ‘body growth and age determination of the sable hippotragus niger niger (harris, 1838)’, koedoe 23, 131–156. http://dx.doi.org/10.4102/koedoe.v23i1.641 grobler, j.h., 1980b, ‘breeding biology and aspects of social behaviour of sable hippotragus niger niger (harris, 1838) in the rhodes matopos national park, zimbabwe’, south african journal of wildlife research 10, 149–153. gusset, m., jakoby, o., muller, m.s., somers, m.j., slotow, r. & grimm, v., 2009, ‘dogs on the catwalk: modelling re-introduction and translocation of endangered wild dogs in south africa’, biological conservation 142, 2774–2781. http://dx.doi.org/10.1016/j.biocon.2009.07.007 harrington, r., owen-smith, n., viljoen, p.c., biggs, h.c., mason, d.r. & funston, p., 1999, ‘establishing the causes of the roan antelope decline in kruger national park, south africa’, biological conservation 90, 69–78. http://dx.doi.org/10.1016/s0006-3207(98)00120-7 hayward, m.w. & kerley, g.i.h., 2005, ‘prey preferences of the lion (panthera leo)’, journal of zoology, london 267, 309–322. http://dx.doi.org/10.1017/s0952836905007508 kelly, m.j. & durant, s.m., 2000, ‘viability of the serengeti cheetah population’, conservation biology 14, 786–797. http://dx.doi.org/10.1046/j.1523-1739.2000.98329.x lacy, r.c., borbat, m. & pollak, j.p., vortex: a stochastic simulation of the extinction process, computer software, version 9.99b, viewed march 2009, from http://www.vortex9.org/vortex.html mcloughlin, c.a. & owen-smith, n., 2003, ‘viability of a diminishing roan antelope population: predation is the threat’, animal conservation 6, 231–236. http://dx.doi.org/10.1017/s1367943003003287 nicholls, a.o., viljoen, a.j., knight, m.h. & van jaarsveld, a.s., 1996, ‘evaluating population persistence of censused and unmanaged herbivore populations from the kruger national park, south africa’, biological conservation 76, 57–67. http://dx.doi.org/10.1016/0006-3207(95)00094-1 nijhof, a.m., pillay, v., steyl, j., prozesky, l., stoltsz, w.h., lawrence, j.a. et al., 2005, ‘molecular characterisation of theilleria species associated with mortality in four species of african antelope’, journal of clinical microbiology 43, 5907–5911. http://dx.doi.org/10.1128/jcm.43.12.5907-5911.2005, pmid:16333074 pmcid:1317158 oosthuizen, m.c., zweygarth, e., collins, n.e., troskie, m. & penzhorn, b.l., 2008, ‘identification of a novel babesia sp. from a sable antelope (hippotragus niger harris, 1838)’, journal of clinical microbiology 46, 2247–2251. http://dx.doi.org/10.1128/jcm.00167-08, pmid:18508943 pmcid:2446884 owen-smith, n. & mason, d.r., 2005, ‘comparative changes in adult vs juvenile survival affecting population trends of african ungulates’, journal of animal ecology 74, 762–773. http://dx.doi.org/10.1111/j.1365-2656.2005.00973.x owen-smith, n. & mills, g., 2008, ‘predator-prey size relationships in an african large-mammal food web’, journal of animal ecology 77, 173–183. http://dx.doi.org/10.1111/j.1365-2656.2007.01314.x, pmid:18177336 owen-smith, n., mason, d.r. & ogutu, j.o., 2005, ‘correlates of survival rates for 10 african ungulate populations: density, rainfall and predation’, journal of animal ecology 74, 774–788. http://dx.doi.org/10.1111/j.1365-2656.2005.00974.x ranta, e., lundberg, p. & kaitala, v., 2006, ecology of populations, cambridge university press, new york. pmcid:1634913 saether, b.e. & bakke, e.o., 2000, ‘avian life history variation and contribution of demographic traits to the population growth rate’, ecology 81, 642–653. taylor, b.l., 1995, ‘the reliability of using population viability analysis for risk classification of species’, conservation biology 9, 551–558. http://dx.doi.org/10.1046/j.1523-1739.1995.09030551.x article information authors: hendri c. coetzee1 werner nell2 elize s. van eeden3 engela p. de crom4 affiliations: 1institutional community engagement office and unit for environmental sciences and management, north-west university, potchefstroom campus, south africa 2optentia research programme, north-west university, vaal campus, south africa 3research group for integrative multidisciplinary eco health and well-being studies, north-west university, vaal campus, south africa 4department of nature conservation, tshwane university of technology, south africa correspondence to: hendri coetzee email: hendri.coetzee@nwu.ac.za postal address: private bag x1290, potchefstroom 2520, south africa dates: received: 06 aug. 2014 accepted: 04 feb. 2015 published: 12 june 2015 how to cite this article: coetzee, h.c., nell, w., van eeden, e.s. & de crom, e.p., 2015, ‘artisanal fisheries in the ndumo area of the lower phongolo river floodplain, south africa’, koedoe 57(1), art. #1248, 6 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1248 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. artisanal fisheries in the ndumo area of the lower phongolo river floodplain, south africa in this original research... open access • abstract • introduction • research method and design    • study area    • design and participants    • procedure and data collection    • data analysis and interpretation • results and discussion    • consumers of fish    • local fishermen and fisherwomen    • fish traders • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ the aim of this study was to investigate the status of artisanal fisheries in the lower phongolo river floodplain in northern kwazulu-natal, south africa. a cross-sectional quantitative survey design was used that included the development of a questionnaire and a systematic survey among the five villages bordering the ndumo game reserve. data were collected over a 5-day period by a group of 16 fieldworkers and analysed using the statistical package for the social sciences, version 21. the results revealed that fish was the third most consumed protein in the area, that people consumed fish on average twice a week, that at least six fish species (and one recently introduced crayfish species) were consumed regularly, and that most fish were obtained from local vendors, who in turn bought it from local fishermen and -women. fishing activities also appeared to occur predominantly along the river system and targeted mainly red-breasted tilapia (tilapia rendalli) and mozambique tilapia (oreochromis mossambicus) and, to a slightly lesser extent, african sharptooth catfish (clarias gariepinus) and brown squeaker (synodontis zambezensis). conservation implications: given that ndumo residents predominantly catch fish by means of non-commercial methods, that they do so for reasons of personal consumption and subsistence, and that they mostly target rivers and dams rather than the ecologically sensitive pans in the region, it would seem likely that fishing in the region might be sustainable for the moment. however, it is recommended that studies on the local fish populations and their reproductive rates be conducted so that the actual impact on local fish populations can be determined more accurately. this study serves to provide the necessary baseline data on fish utilisation in the region, which would enable the impact of artisanal fishing on fish reserves in the ndumo region to be determined once population studies have been completed. introduction top ↑ artisanal or small-scale fisheries that support people's livelihoods are a worldwide phenomenon. they are characterised by the use of traditional fishing techniques, low-level technologies (e.g. handlines, rods, traditional boats, etc.) and normally take place close to the shore. the fish that are caught are intended mostly for local markets. artisanal fisheries tend to have a much smaller impact on local fish populations compared to large-scale commercial fishing practices and thus tend to be more sustainable (béné, macfadyen & allison 2007). throughout central and southern africa, inland water bodies are important sources for artisanal fisheries (andrew, rouhani & seti 2000; turpie et al. 1999). these multi-species fisheries are generally highly productive and a major contributor to the livelihoods of many local communities (baker 2008; neiland & béné 2006), especially in the large lake systems in central and eastern africa (heck, béné & reyes-gaskin 2007). in south africa, however, the relative scarcity of wetlands means that those who operate artisanal fisheries, especially in floodplains, are not experienced in generating high levels of fishery productivity, with the exception of some of the larger systems such as the phongolo floodplain (acreman & hollis 1996; merron & weldrick 1995; ross & ross 2012). the ecological value of the phongolo river floodplain ecosystem, and particularly the varying natural and anthropogenic impacts on the system, have been recognised decades ago (breen et al. 1973; heeg & breen 1982). recently, a number of additional studies were conducted on human activity in this region and the various ways in which such activities affect the ecosystem of the phongolo floodplain (chhotray 2010; ellender et al. 2010; lankford et al. 2010; pongola river ecosystem services for poverty alleviation report 2009). ross and ross (2012) found that fish was the main protein source in the floodplain area. however, whilst there seemed to be consensus about the importance of fishing in the region, far less is known about aspects such as the types of species most commonly consumed and the nature and extent of local trade in fish. given the ecological sensitivity of the phongolo river floodplain, especially around the ndumo game reserve, as well as the possible pressures that artisanal fisheries might exert on the system, this study set out to investigate artisanal fisheries in the region by focusing on fishing, fish consumption and the commercial utilisation of fish in the ndumo area. research method and design top ↑ study area the ndumo area is situated in the north-eastern corner of south africa's kwazulu-natal (kzn) province, south of swaziland and southern mozambique. the area is comprised of a small town (with limited facilities and infrastructure) and a number of rural villages that border the southern and western parts of the ndumo game reserve. ndumo is a provincial game reserve in south africa that is managed by ezemvelo kzn wildlife. the whole area falls under the jozini local municipality, which in turn forms part of the larger umkhanyakude district municipality in northern maputaland (see figure 1). figure 1: location of the ndumo area, kwazulu-natal, south africa. to the north of the ndumo area, the usutu river separates south africa from mozambique. in the east, the pongola river runs through the ndumo area from north to south and into the larger phongolo river and its floodplain. the area lies within a summer rainfall area and has a subtropical climate with an average annual rainfall of between 670 mm and 1000 mm, with temperatures ranging from 13 °c to 40 °c (morgenthal et al. 2006). design and participants a cross-sectional quantitative survey design (sarantakos 2005) was adopted as a basis for this study. in total, 521 participants participated in the survey. their average age was 37.28 years (s.d. = 13.12) and participants had been living in the ndumo area for a mean of 28.18 years (s.d. = 16.24). table 1 depicts additional demographic characteristics of the sample. table 1: social and biographical characteristics of participants. procedure and data collection entry into ndumo (i.e. various villages) was negotiated on behalf of the research team by an experienced research assistant. the research assistant also used the opportunity to gather basic data that was used as a basis for the development of a questionnaire. further data was collected by means of a structured questionnaire (creswell 2007) during a second visit to the area. the questionnaire consisted of 40 questions that were divided into 4 sections: demographic characteristics; questions relating to the consumption of fish and fish products; questions relating to local fishing habits; and, finally, questions relating to local trading in fish. a group of 16 fieldworkers were sourced from local villages and trained to recruit participants, to obtain informed consent, and to administer the questionnaire by means of structured face-to-face interviews. all data was collected over a 5-day period in august 2013, with fieldworkers receiving a set of questionnaires in the morning, which they administered during the course of the day. questionnaires were collected at the end of each day and scrutinised for completeness. during the entire process of data collection, care was taken to do so in as ethical a manner as possible. full, informed and signed consent was obtained from all participants. participants were advised that their responses would be treated confidentially and that they had the right to withdraw from the research at any time without penalty. data analysis and interpretation data was analysed using the statistical package for the social sciences, version 21 (spss 21). basic descriptive statistics (means and measures of variance) were calculated for all items in the questionnaire. wherever relevant, findings were compared across demographic groups using chi-square tests, with cross tabulation and independent t-tests to verify the statistical significance of any differences that emerged, while bivariate correlations were used to examine relationships between variables (field 2005). results and discussion top ↑ consumers of fish as reflected in table 2, fish was found to be the third most frequently consumed protein per week (after chicken and beef) among all participants. this contradicts earlier findings (ross & ross 2012) that fish was the main protein source in the floodplain area. among those who consumed fish, which included 88.9% of the participant group, fish was consumed on average twice a week. on those days that fish was consumed, participants did so 1.4 times per day on average (s.d. = 0.59). table 2: mean types of protein sources consumed by villagers in the ndumo area. additionally, participants were asked to specify which species of fish they ate and how frequently they did so. table 3 provides a summary of these findings. table 3: percentage of participants who consumed the species as listed, and mean number of fish or crayfish consumed per week. overall, it was found that all species of fish listed in table 3 were consumed in fairly high numbers by a substantial portion of the participant group, with the exception of the recently introduced redclaw crayfish (cherax quadricarinatus) (see du preez & smit 2013). reasons for the low levels of utilisation of this (non-native invasive) species are still unclear and should be examined more closely in future studies. the two tilapia species that occur in the region were the most preferred and frequently consumed species of fish, with red-breasted tilapia (tilapia rendalli) being much more popular than mozambique tilapia (oreochromis mossambicus). however, while the african sharptooth catfish (clarias gariepinus) was consumed by only 54.8% of the participants, it was the third most frequently consumed fish of all the species. bivariate correlations were used to examine the relationships between age, the number of years that participants resided in the ndumo area and their consumption of fish, both generally and in relation to specific species. age was uncorrelated to general fish consumption, but it was found that the longer participants lived in the ndumo area, the more likely they were to eat red-breasted tilapia (t. rendalli) (r = 0.12, p = 0.02), brown squeaker (synodontis zambezensis) (r = 0.15, p = 0.02), mozambique tilapia (o. mossambicus) (r = 0.13, p = 0.02) and, in particular, tigerfish (hydrocynus vittatus) (r = 0.24, p = 0.00), suggesting that those living in this region develop an acquired taste for these species. asked how they obtained the fish they consumed, 46.1% of participants (54.4% of females and 37.3% of males) indicated that they bought the fish from a vendor in the area; 37.8% (similar for both genders) obtained the fish from family or friends, whereas 16.1% (25.5% of males and 7.2% of females) caught the fish themselves. a chi-square test with cross tabulation confirmed that the association between gender and where participants obtained the fish was significant [χ²(2, n = 457) = 31.18, p = 0.00] and of moderate strength (ɸ = 0.26). local fishermen and fisherwomen participants indicated that they commonly fished in an average of 4.32 (s.d. = 5.07) different locations. however, the majority of participants (70.48%) only fished in one to three locations. among these sites, 81.02% of fish was caught in rivers and streams, 13.60% in pans and 1.9% in other (unspecified) locations, suggesting that fishing was mostly focused around the river system and that pans were not highly exploited for fishing. the main reason for this seems to be that the majority of the pans in the ndumo region falls within the confines of the ndumo game reserve and, as such, are protected. participants indicated that the bulk of fish caught was intended to be sold (54.73%) and that 44.28% of fish caught was for personal consumption. the most commonly employed fishing techniques were handlines (26.77%), traditional baskets (25.81%) and drag nets (22.26%), followed by gill nets (17.10%) and, to a much lesser extent, fishing rods (8.19%). thus, non-commercial techniques were used and no evidence of large-scale fishing was uncovered. this indicates that local communities in the area fish primarily for personal subsistence use rather than for large-scale commercial reasons. the species of fish that participants usually caught and the average number of fish per species is summarised in table 4. table 4: summary of type of species caught, and number of each species caught in an average week. red-breasted tilapia and, to a lesser degree, mozambique tilapia emerged as the species that were most frequently caught in the region. whilst the vast majority of participants indicated that they actively sought to catch african sharptooth catfish, a much lower number of these fish was caught. however, as can be seen from the large standard deviations associated with the number of fish caught, great variability exists in this regard, with some fishermen and -women catching 1 or 2 fish per week and 1 individual reporting catching as many as 200 tilapia species per week. these differences could possibly be attributed to the fact that whereas most fishermen and -women fished for personal consumption and as such only needed a relatively small number of fish, a few residents did so for small-scale commercial purposes and therefore made far larger catches. no statistically significant gender differences were found in relation to the types or numbers of fish species caught. in order to determine whether any changes or trends pertaining to fish populations had been observed by local fishermen and -women, participants were asked whether they had noticed any changes in the number of fish over the past 10 years. responses were measured on a five-point scale ranging from ‘decreased a lot’ (1) to ‘increased a lot’ (5). a score of 3 indicated that there was no observed change. participants’ mean score was 2.91 (s.d. = 1.22), suggesting that they observed very little change. the same was true for the average size of the fish, which was viewed as decreasing only fractionally (m = 2.89; s.d. = 1.25). fish traders lastly, data were also collected from participants who sold fish, but did not catch it themselves. the average trader indicated that he or she had been selling fish in the area for 7.50 years (s.d. = 5.87) and that they obtained 85% of the fish from local fishermen and -women and 15% from other undisclosed suppliers. traders indicated that they mostly sold their fish in nearby towns (40.7%) (mainly ndumo, but occasionally jozini, approximately 75 km farther south) and in their own villages (30.7%). to a lesser extent, fish was also sold in other villages where they did not reside (15.9%) or ‘next to the road’ (referring to areas outside towns or villages) (11.1%). this suggests that the local fishing industry, though limited in opportunities, contributes directly and meaningfully to the economy of the ndumo area, in which income-generating opportunities are very scarce (ogilvie pers. comm. 2014). table 4 indicates the number of fish that were sold by traders during the course of a typical week. it also reflects the mean and modal prices (in south african rand [zar]) that is charged for an average-sized fish of a given species. the findings reported in table 5 suggest that the two species of tilapia and african sharptooth catfish were the two species most frequently sold and also the most expensive of all the fish species that were exploited in the area. however, when the modal prices for all the fish species were considered, it would appear that fish of all species were most commonly sold for zar 10 (with the exception of african sharptooth catfish, which sold for a modal amount of zar 20; and the redclaw crayfish, which sold for a modal amount of zar 15). as such, these species likely play an important role in the context of the local economy within the ndumo and the greater phongolo river floodplain region. table 5: number of fish sold during a typical week, and mean and modal prices per average-sized specimen for each species (2013 data). none of these species are regarded as threatened or endangered, with the exception of the tigerfish, which is a protected species in south africa. however, even though residents tend to increase their consumption of this species the longer they reside in the ndumo area, this species is consumed fairly infrequently and does not appear to be specifically targeted as much as being incidentally caught. assessing the impact of human activities on fish populations in the ndumo area is hampered by the fact that very little research has been conducted to determine the size and reproductive rate of fish populations in this ecologically sensitive region (smit pers. comm. 2014). as such, it is hoped that the findings of the present study would provide the necessary baseline data on fish consumption in the region so that the ecological impact on local fish populations can be assessed once future studies on fish populations and reproductive rates in the area have been completed. conclusion top ↑ the results of this study suggest that artisanal fishery activities in the ndumo area of kzn play an important role in supporting the livelihood of members of this community, both by serving as an important form of sustenance and by constituting an important economic resource. however, given the ecological sensitivity of the floodplain, it is also necessary to assess the impact that these interactions might have on the fish populations in the region, for example, by determining the size and reproductive rate of the most important species and populations. to fully understand human habits and needs in the floodplain, the value of extended socio-economic research in the future in this region should also be seriously considered. finally, continuous monitoring of artisanal fishery activities is important to assess and ensure the sustainability of these interactions in the region. acknowledgements top ↑ the authors would like to thank the water research commission (wrc) of south africa (project k5-2185 n.j. smit, pi) for the financial support of this study. views expressed are those of the authors and not those of the wrc. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions e.v.e. (north-west university) was the project leader. all four authors were involved in the conceptualization of the research. h.c.c. (north-west university) and w.n. (north-west university) were responsible for all data collection and analysis. all authors provided inputs and helped in the writing of the article. references top ↑ acreman, m.c. & hollis, g.e., 1996, water management and wetlands in sub-saharan africa, international union for the conservation of nature (iucn), gland, switzerland. andrew, t.g., rouhani, q.a. & seti, s.j., 2000, ‘can small-scale fisheries contribute to poverty alleviation in traditionally non-fishing communities in south africa?’, african journal of aquatic science 25(1), 49–55. http://dx.doi.org/10.2989/160859100780177938 baker, c.f., 2008, ‘seasonal floodplain wetlands as fish habitat in oregon and washington’, phd thesis, department of fisheries science, oregon state university, corvallis. béné, c., macfadyen, g. & allison, e.h., 2007, ‘increasing the contribution of small-scale fisheries to poverty alleviation and food security’, p. 125, food and agriculture organization of the united nations (fao) fisheries technical paper no. 481, rome. breen, c.m., furness, h.d., heeg, j. & kok, j. (eds.), 1973, ‘bathymetric studies on the pongola river floodplain’, journal of the limnological society of southern africa 4(2), 95–100. http://dx.doi.org/10.1080/03779688.1978.9633158 chhotray, v., 2010, ‘the impacts of the ecosystem service and environmental governance on human well-being in the pongola region, south africa’, working paper, uea, norwich and institute for natural resources, pietermaritzburg. https://ueaeprints.uea.ac.uk/37820/ creswell, j.w., 2007, qualitative inquiry and research design: choosing among five approaches, sage publications, thousand oaks, california. du preez, l.h. & smit, n.j., 2013, ‘double blow: alien crayfish infected with invasive temnocephalan in south african waters’, south african journal of science 109(9/10), 1–4. http://dx.doi.org/10.1590/sajs.2013/20130109 ellender, b.r., weyl, o.l.f., winkler, h., stelzhammer, h. & traas, g.r.l., 2010, ‘estimating angling effort and participation in a multi-user, inland fishery in south africa’, fisheries management and ecology 17, 19–27. http://dx.doi.org/10.1111/j.1365-2400.2009.00708.x field, a., 2013, discovering statistics using ibm spss statistics, 4th edn., sage publications, london. heck, s., béné, c. & reyes-gaskin, r., 2007, ‘investing in african fisheries: building links to the millennium development goals’, fish and fisheries 8(3), 211–226. http://dx.doi.org/10.1111/j.1467-2679.2007.00251.x heeg, j. & breen, c.m., 1982, ‘man and the pongola floodplain’, report no. 56 of the committee for inland water ecosystems, national programme for environmental sciences, council for scientific and industrial research (csir), south africa. https://ueaeprints.uea.ac.uk/33628/ lankford, b., pringle, c., dickens, c., chhotray, v., mander, m., goulden, m. et al., 2010, ‘the impacts of ecosystem services and environmental governance on human well-being in the pongola region, south africa’, working paper, university of east anglia and institute of natural resources, norwich, uk and pietermaritzburg, south africa. mavundza, e.j., maharaj, r., finnie, j.f., kabera, g. & van staden, j., 2011, ‘an ethnobotanical survey of mosquito repellent plants in umkhanyakude district, kwazulu-natal province, south africa’, journal of ethnopharmacology 137(3), 1516–1520. http://dx.doi.org/10.1016/j.jep.2011.08.040 merron, g.s. & weldrick, s.k., 1995, fisheries management of the phongola river floodplain, investigational report no. 51, jlb smith institute of ichthyology, grahamstown, south africa. morgenthal, t.l., kellner, k., van rensburg, l., newby, t.s. & van der merwe, j.p.a., 2006, ‘vegetation and habitat types of the umkhanyakude node’, south african journal of botany 72, 1–10. http://dx.doi.org/10.1016/j.sajb.2005.03.003 neiland, a.e. & béné, c., 2006, tropical river fisheries valuation: a global synthesis and critical review, international water management institute, colombo, sri lanka. pongola river ecosystem services for poverty alleviation (prespa), 2009, ‘livelihoods and poverty linkages to the ecosystem services provided by the pongola floodplain’, project brief no. 3. https://www.uea.ac.uk/polopoly_fs/1.140758!livelihoods%20project%20brief%20prespa%20dec%2009..pdf ross, m. & ross, t., 2012, umkhanyakude district municipality, status quo report: rivers & wetlands, environmental management framework report, johannesburg. sarantakos, s., 2005, social research, 3rd edn., palgrave macmillan. turpie, j., smith, b., emerton, l. & barnes, j., 1999, economic value of the zambezi basin wetlands, zambezi basin wetlands conservation and resource utilization project, international union for the conservation of nature (iucn), regional office for southern africa, harare. abstract introduction research method and design results and discussion general discussion acknowledgements references about the author(s) alexis a. olds capenature, scientific services, south africa department of ichthyology and fisheries science, rhodes university, south africa center for invasion biology, south african institute for aquatic biodiversity, south africa nicola c. james south african institute for aquatic biodiversity, grahamstown, south africa m. kyle s. smith south african national parks, rondevlei scientific services, south africa olaf l.f. weyl department of ichthyology and fisheries science, rhodes university, south africa center for invasion biology, south african institute for aquatic biodiversity, south africa south african institute for aquatic biodiversity, grahamstown, south africa citation olds, a.a., james, n.c., smith, m.k.s. & weyl, o.l.f., 2016, ‘fish communities of the wilderness lakes system in the southern cape, south africa’, koedoe 58(1), a1364. http://dx.doi.org/10.4102/koedoe.v58i1.1364 original research fish communities of the wilderness lakes system in the southern cape, south africa alexis a. olds, nicola c. james, m. kyle s. smith, olaf l.f. weyl received: 12 oct. 2015; accepted: 22 may 2016; published: 31 aug. 2016 copyright: © 2016. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the wilderness lakes system, a temporarily open and closed estuary with three associated lakes situated in the southern cape region of south africa, was sampled using a range of sampling gears to assess the fish community. a total of 25 species were sampled throughout the system, with the highest diversity in the touw estuary (23 species) and the lowest in langvlei (11 species). estuary-associated marine species (13 species) dominated species richness with smaller proportions of estuarine resident (7 species), freshwater (3 species) and catadromous species (2 species). estuarine resident species dominated the catch numerically. the size–class distribution of euryhaline marine species indicated that upon entering the touw estuary as juveniles, the fish move up the system towards rondevlei where they appear to remain. three freshwater species were recorded in the system, all of which are alien to the wilderness lakes system. decreasing salinity in the upper lakes appears to be a driving factor in the distribution and increasing abundance of the freshwater fishes. sampling followed a drought, with the system experiencing substantially increased levels of mouth closure compared to a similar study conducted in the 1980s. the timing of mouth opening and the degree of connectivity between the lakes influence the nursery function of the system as a whole. management actions need to focus on improving ecological functioning of this system, in particular how mouth opening is managed, to facilitate nursery function and limit the establishment of invasive species. conservation implications: key management actions are required to improve fish recruitment potential into and within the system. these include maintenance of adequate marine inflow through adherence to artificial mouth breaching protocols and improving connectivity between the lakes through sediment removal from localised deposition points within the connecting channels. introduction the wilderness lakes system (figure 1) comprise a series of three coastal lakes and a temporarily open and closed estuary connected by shallow channels. it is located on the warm-temperate south coast of south africa between the towns of george (16 km) and knysna (40 km). the national conservation importance of the wilderness lakes system (hereafter referred to as wilderness lakes) has been assessed in several studies (maree, whitfield & quinn 2003; turpie et al. 2002). the wilderness lakes is an important component of the garden route national park vision, namely ‘an integrated protected area that effectively conserves a functionally linked mosaic of diverse terrestrial, freshwater, estuarine and marine ecosystems, landscapes, and cultural heritage, representative of the garden route, that contributes to the well-being of present and future generations’ (sanparks 2012:28). figure 1: map of the wilderness lakes system. major threats to the system include alien invasive species, development pressure, water resource management and climate change (sanparks 2012). to be effective, management interventions will need to be guided by a sound understanding of the physical, biological and social components of the system. estuaries constitute important nursery areas for a number of estuary-associated fish species (potter & hyndes 1999; whitfield 1994), and as such fish assemblages have been used as indicators of estuary ecosystem health (e.g. harrison & whitfield 2004; whitfield & elliott 2002). an understanding of the relative abundances and species composition of fish communities in estuaries is therefore important for guiding management actions as they reflect the state of the estuary. although fish communities in the wilderness lakes have been previously studied (hall 1985; hall, whitfield & allanson 1987; olds et al. 2011; russell 1996, 1999), the last assessment of the fish community throughout the system using multiple gear types was undertaken in the 1980s by hall (1985) and hall et al. (1987). as a result, an assessment of the present state of the fish community allows for spatial comparisons across estuaries and temporal assessments of change within an estuary. this paper describes a basic assessment of the abundance and distribution of fish species throughout the wilderness lakes in 2010–2011 and uses the relative abundance and species composition data to highlight management needs for this important estuarine lakes system. study area the wilderness lakes is comprised of three coastal lakes: the first two, eilandvlei and langvlei, are barrier lagoons and the third, rondevlei, is a deflation basin that has been artificially connected by a channel to langvlei (hall et al. 1987). the lakes provide a flood plain for the touw river and estuary (figure 1). the depth of the touw estuary varies from 1.5 m to 3 m (hall et al. 1987; russell 2003). rondevlei is the smallest lake with a surface area of 1.43 km2 and a maximum depth of 6 m. langvlei is the shallowest lake with a maximum depth of 4 m, but has the largest surface area of 2.16 km2. eilandvlei has a maximum depth of 6.5 m and is the deepest in the system, with a surface area of 1.5 km2 (hall et al. 1987). under drought conditions, the water levels in the system drop significantly, resulting in the separation of rondevlei, langvlei and the connecting channel. this scenario was observed between march and may 2010 during the study period. the touw estuary is a temporarily open and closed estuary that is closed off from the sea for varying periods by a sandbar at the mouth. under pristine conditions, the estuary opened approximately 40% of the time. opening of the estuary mouth has been managed by sanparks since the 1970s to prevent flooding of private property adjacent to the estuary (russell 2013). when water levels in the estuary reach between 2.1 m and 2.4 m above mean sea level, the mouth is opened manually by sanparks. declining freshwater inputs and artificial breaching have resulted in reduced opening of the estuary, with the estuary now only open approximately 28% of the time (russell 2013). during this study, the estuary mouth was only open in july 2010 (4 days) and between december 2010 and january 2011 (28 days) (figure 2b). figure 2: mouth state of the touw estuary between (a) april 1982 and april 1983 (hall 1985) and (b) between april 2010 and april 2011. the wilderness lakes often exhibits a reverse salinity gradient, with the salinity increasing higher up the system towards rondevlei (allanson & whitfield 1983). tidal penetration into the lakes is minor during open phases, with higher salinities in rondevlei and langvlei caused by low or no freshwater input into these lakes (russell 2013). a reverse salinity gradient was present throughout the study period, with the highest salinities (mean ± s.d.: 10.1 ppt ± 0.2 ppt) recorded in langvlei and the lowest in the touw estuary (mean ± s.d.: 4 ppt ± 2.3 ppt). surface water temperature varied little across the system, but varied seasonally with relatively warm summer (mean ± s.d.: 26.0 °c ± 0.9 °c) and mild winter (mean ± s.d.: 13.6 °c ± 0.6 °c) temperatures. the wilderness lakes are bordered by a margin of emergent aquatic plants predominantly made up of phragmites australis (common reed) and, to a lesser extent, juncus kraussii (dune slack rush), schoenoplectus scirpoides (biesiegoed) and typha capensis (bulrush) (russell 2003). submerged aquatic macrophytes occur in each of the lakes and are mostly made up of stands of potamogeton pectinatus (fennel pondweed), charophyta and filamentous algae (cladophora spp.). zostera capensis (eelgrass) occurs sporadically in the touw estuary (allanson & whitfield 1983; howard-williams & liptrot 1980; weisser & howard-williams 1982). during drought conditions, the density of aquatic vegetation decreases, which is assumed to be the result of increased salinity levels. research method and design the system was sampled using a seasonally and geographically stratified sampling design between april 2010 and january 2011. the year was divided into four seasonal sampling periods, namely, autumn (april 2010), winter (july 2010), spring (october 2010) and summer (january 2011). three different net types were used to sample as wide of a range of habitats, species and size classes as possible. each season, 12 double-ended fyke nets (8 m guiding net, first-ring diameter of 55 cm and 10 mm mesh size at the cod end) were set randomly in each of the lakes (a total of 48 fyke nets per lake) whilst six fyke nets were randomly set within the middle and upper reaches of the touw estuary (a total of 24 fyke nets). fyke nets are a passive gear type and were set in water at a depth of approximately 1 m – 1.5 m. many of these areas were on the verge between sandy substrates and beds of p. pectinatus. all fyke nets were fitted with an ‘otter guard’ comprising plastic mesh with openings no larger than 10 cm × 10 cm to prevent cape clawless otters, aonyx capensis, from entering. although the use of otter guards influenced the maximum size of fish that could be sampled, their use was considered critical to avoid bycatch. fyke nets were set between 16:00 and 18:00 and lifted the next morning in sequence between 06:00 and 08:00 with an average soak time of 16 h. a set of three multi-meshed gill nets each measuring 35 m × 2.75 m with stretch meshes of 35 mm, 45 mm, 57 mm, 73 mm, 93 mm, 118 mm and 150 mm (5 m per mesh size) were set for two consecutive nights per season in different positions in each of the lakes. two nets were set once per season in the upper and lower reaches of the touw estuary. gill nets were deployed at sunset and had an average soak time of 2 h. several mesh sizes were used to eliminate the size and species selectivity, which could have caused potential problems when investigating population and size frequency analyses (prchalová et al. 2009). a large seine net (30 m long × 2 m deep with a bag, 12 mm stretched mesh), laid from a boat approximately 30 m offshore, was used to target fish from the littoral and pelagic zones. initially, three pulls at three sites were completed in each of the lakes and the touw estuary per season. because of progressively increasing water levels and filamentous epiphytic algae (enteromorpha spp.) growth, the number of pulls had to be reduced. fish caught were identified to species level, counted and measured to the nearest millimetre fork length (fl) or total length (tl) depending on species, and released alive where possible. sub-sampling was used in cases where the catch comprised large numbers of small fish, from which total abundances were estimated. where appropriate, the length frequency distribution (in 10 mm size classes) was calculated for the most abundant species within the system. results and discussion species composition during this study, a total of 25 species were caught (tables 1, 2 and 3). a total of 23 species were recorded in the touw estuary and eilandvlei, 11 in langvlei and 15 in rondevlei. of the species caught, euryhaline marine species were dominant (15 species), followed by estuarine resident species (7 species) and freshwater species (3 species). estuarine resident species, particularly estuarine roundherring (gilchristella aestuaria) and cape silverside (atherina breviceps), were the most abundant species recorded (with more than 10 000 individuals caught). seine net catches in all water bodies were dominated numerically by estuarine roundherring (g. aestuaria) and cape silverside (a. breviceps), which together comprised between 66% (rondevlei) and 99% (touw estuary) of the catch (table 1). the freshwater species mozambique tilapia (oreochromis mossambicus) was also numerically abundant in the system, with more than 5000 individuals sampled. mozambique tilapia comprised between 63% of the catch (touw estuary) and 99.9% (langvlei and rondevlei) of the fyke net catch (table 2). gill nets, which sampled larger individuals of marine and freshwater species, were dominated numerically by euryhaline marine species, particularly southern mullet (liza richardsonii), which comprised between 10% (langvlei) and 40% (eilandvlei) of the catch, and oval moony (monodactylus falciformis), which comprised between 3% (eilandvlei) and 48% (langvlei) of the catch. table 1: relative abundance (% abundance) of fish species caught in seine nets in the wilderness lakes system between april 2010 and january 2011. table 2: relative abundance (% abundance) of fish species caught in fyke nets in the wilderness lakes system between april 2010 and january 2011. table 3: relative abundance (% abundance) of fish species caught in gill nets in the wilderness lakes system between april 2010 and january 2011. euryhaline marine species oval moony was present throughout the system and was common in langvlei and rondevlei, where it comprised, respectively, 48% and 27% of the gill net catch (table 3). although this species commonly uses the middle and upper reaches of estuaries as nursery areas, adults (> 120 mm standard length [sl]) are generally found in the near-shore marine environment (whitfield 1998). individuals captured during this survey ranged in size from 13 mm fl to 228 mm fl and showed a bimodal length frequency distribution. the majority of small individuals in the 90 mm fl – 100 mm fl size class were caught in the touw estuary. larger specimens represented by the 190 mm fl – 200 mm fl size class were caught in langvlei and rondevlei (figure 3). the presence of individuals > 120 mm fl in the upper lakes suggests that they were trapped in the system. figure 3: length frequency of (a) monodactylus falciformis, (b) liza richardsonnii, (c) myxus capensis, (d) rhabdosargus holubi, (e) lithognathus lithognathus and (f) pomadasys commersonnii captured in the wilderness lakes system between april 2010 and may 2011. southern mullet was present in all water bodies and common (comprised more than 20% of the catch) in gill net catches in eilandvlei and rondevlei (table 3). common as juveniles and adults in estuaries in the eastern and western cape, southern mullet is normally more abundant in the adjacent near-shore marine environment (bennett 1989). individuals captured during this survey were all large, mature individuals ranging in size between 200 mm fl and 483 mm fl, with most individuals in the 340 mm fl – 350 mm fl size class (figure 3). the smallest individuals were recorded in eilandvlei. maturing at 225 mm fl, with spawning taking place in the near-shore marine environment (de villiers 1987; van der horst & erasmus 1981), they are usually more abundant in the lower and middle reaches of southern cape estuaries (whitfield & kok 1992). the abundance of adults, particularly in langvlei and rondevlei, during this study indicates possible entrapment in the upper lakes. although it is found as juveniles and adults in both estuaries and rivers, freshwater mullet (myxus capensis) is often referred to as a catadromous species, with spawning occurring in the near-shore marine environment (bok 1979). freshwater mullet was present throughout the system, and increased in abundance in rondevlei where it comprised 31% of the gill net catch (table 3). individuals showed a bimodal length frequency distribution; the first size class comprised individuals with a mode of 300 mm fl – 310 mm fl and the second size class comprised individuals with a mode of 380 mm fl – 390 mm fl. the larger individuals were found in rondevlei (figure 3). cape stumpnose (rhabdosargus holubi), considered to be entirely dependent on estuarine habitats for the first year of life (whitfield 1994), was present throughout the system and more common in rondevlei where it comprised 5% of the gill net catch (table 3). cape stumpnose is often the most abundant marine species recorded in both temporarily open and closed and permanently open estuaries in the warm-temperate region of south africa (harrison 2005; vorwerk et al. 2001). individuals captured during this survey ranged in size from 120 mm fl to 420 mm fl with a polymodal length frequency distribution; the first size class comprised individuals with a mode of 140 mm fl – 160 mm fl, caught mainly in the touw estuary. the second size class comprised individuals with a mode of 230 mm fl – 240 mm fl, mainly from eilandvlei, and the third size class comprised individuals with a mode of 340 mm fl – 350 mm fl from rondevlei (figure 3). cape stumpnose normally migrate back to sea after reaching 120 mm sl. some individuals, however, remain trapped in closed estuaries, where they reach sizes greater than 200 mm sl (james, cowley & whitfield 2007a). large, predatory species such as leervis (lichia amia) and springer (elops machnata) were present in the estuary, eilandvlei and rondevlei, with one large leervis caught in rondevlei (> 900 mm fl). spotted grunter (pomadasys commersonnii) and white steenbras (lithognathus lithognathus), two important fishery species, showed a similar trend, with a greater proportion of smaller individuals sampled in the touw estuary and eilandvlei and only a few very large individuals caught in rondevlei (table 3 and figure 3). these species are all dependent on estuaries as nursery areas, but usually return to sea as sub-adults (whitfield 1998). estuarine resident species seven estuarine resident species were caught, of which estuarine roundherring and cape silverside were common or abundant in seine net catches in all water bodies. the other estuarine resident species, including prison goby (caffrogobius gilchristi) (touw estuary and eilandvlei) and speckled sandgoby (psammogobius knysnaensis) (touw estuary, eilandvlei and rondevlei), were not caught in all water bodies. cape silverside captured during this survey were between 16 mm fl and 100 mm fl. the majority (> 85%) caught in eilandvlei and rondevlei were mature individuals (< 40 mm, ratte 1989). in contrast, only 53% of the individuals caught in the touw estuary and 59% of the individuals caught in langvlei were mature (figure 4). estuarine roundherring captured during this survey were between 20 mm fl and 89 mm fl, with the largest individuals caught in rondevlei (figure 4). estuarine roundherring is estimated to mature at a size of 28 mm sl (talbot 1982). figure 4: length frequency of (a) gilchristella aestuaria and (b) atherina breviceps captured in the wilderness lakes system between april 2010 and may 2011. freshwater species three alien freshwater species, mozambique tilapia, western mosquitofish (gambusia affinis) and common carp (cyprinus carpio), were sampled. mozambique tilapia was abundant throughout the system (dominated fyke net catches in all sampling areas). although endemic to southern africa, mozambique tilapia does not occur naturally south of the bushmans estuary in the eastern cape, but the species has been introduced extensively into farm dams as an angling fish in systems further south along the coastline, from which it has escaped. it is most abundant in temporarily open and closed estuaries during the closed phase (whitfield & blaber 1979), possibly because of a combination of increased suitable nesting habitats during the closed phase and reduced recruitment success during open phases (ellender et al. 2008). although large individuals (> 200 mm fl) were caught throughout the system, the majority were small (< 180 mm fl) individuals (figure 5). figure 5: length frequency of (a) oreochromis mossambicus and (b) cyprinus carpio captured in the wilderness lakes system between april 2010 and may 2011. the first formal record of the invasive common carp within the wilderness lakes was from langvlei in 2009 (olds et al. 2011), with fish measuring between 150 mm and 300 mm. larger specimens (500 mm fl – 800 mm fl) were sampled in 2010 in each of the lakes, whilst in 2011 three juveniles were sampled in the touw estuary (figure 5). together the data show a progression in size classes from 2009 to 2011 and confirms a successful spawning event having occurred in the wilderness lakes. western mosquitofish were sampled in the 30 m seine net within the touw estuary during this study and previously in the touw and duiwe rivers by russell (1999). this small species is most likely underrepresented in seine net samples because it is generally associated with marginal vegetation in the shallow littoral zone (sloterdijk et al. 2015). sloterdijk et al. (2015) sampled the wilderness lakes using scoup nets concurrently with this study and showed this species to be widespread and abundant throughout the system. general discussion the wilderness lakes is one of a few estuarine lakes systems in southern africa, with only 3% of southern african systems comprising estuarine lakes (whitfield 1998). in common with other estuarine lakes in the warm-temperate region (such as swartvlei, bot and klein), the fish fauna of the wilderness lakes is largely made up of euryhaline marine species, with smaller proportions of estuarine species and freshwater species (bennett et al. 1985; scott, harrison & macnae 1952; whitfield & kok 1992). although the overall fish community is predominantly made up of marine species, estuarine resident fishes were numerically dominant in the seine nets, with estuarine roundherring and cape silverside comprising the highest number of fish caught overall. the numerical dominance by cape silverside and estuarine roundherring in the system was also noted by hall (1985), james and harrison (2008) and olds et al. (2011). within warm-temperate temporarily open and closed estuaries, estuarine resident species comprise well over 50% of the catch (by numbers) (james et al. 2007b). although only a few species are able to complete their entire life cycle in estuaries (day, blaber & wallace 1981), short-lived estuarine resident species are well adapted to the estuarine environment and can dominate the fish communities of estuaries numerically (potter et al. 1990). cape silverside and estuarine roundherring comprised a larger percentage of the catch (by numbers) in temporarily open and closed estuaries on the southeast coast than in permanently open estuaries and also had a smaller mean size in temporarily open and closed estuaries than in permanently open estuaries (vorwerk et al. 2001, 2003). this was attributed to a higher survival rate of smaller individuals in closed estuaries. the presence of a range of size classes of these species in the wilderness lakes may be attributed to the predominantly closed mouth and retention of size classes within the system. the touw estuary and eilandvlei had the highest species diversity (23 species), which was predominantly made up of euryhaline marine species, whilst langvlei had the lowest species diversity (11 species). the low species diversity higher up the system was also noted by hall (1985), with langvlei and rondevlei having low fish diversity. hall (1985) sampled the estuary seasonally between april 1982 and april 1983 using similar gear. hall (1985) and hall et al. (1987) found that the depth of the interconnecting channels in wilderness lakes has a major influence on fish community structure in the lakes, with the marked decline in marine species richness between eilandvlei and langvlei attributed to the tenuous connection between these two lakes (figure 6). in the first half of this study period, langvlei and rondevlei were also completely isolated from each other owing to a drought. in contrast, marine species dominate the lake compartments of the subtropical kosi bay system owing to a consistent connection between the lakes (blaber 1978). hall (1985) also recorded nine marine species in the estuary that were not found in the lake components of the system. these were mainly predatory species, such as dusky kob, leervis and white seacatfish (galeichthys feliceps). hall (1985) attributed this to the preference of these species for the deep, turbid waters of estuaries and their unwillingness to negotiate the shallow waters of the connecting serpentine channel. interestingly, during this study, only one species, white seacatfish, was recorded in the touw estuary and not in the lakes. white steenbras, leervis and spotted grunter were most abundant in eilandvlei and in the touw estuary, with low numbers of very large fish sampled from rondevlei. the decreasing abundance but increase in size of estuary-associated marine species up the system was also reported by hall (1985). the smaller fish found in eilandvlei suggests that this lake serves as the system’s initial nursery ground for the marine-spawned species (hall 1985). once these fish reach rondevlei, it appears that they remain trapped there, which accounts for the presence of very large euryhaline marine fishes in rondevlei. langvlei had the lowest number of euryhaline marine species and it appears that these species either stay in eilandvlei or move through to rondevlei. with few euryhaline marine species being sampled by both hall (1985) and us (in this study), it may be suggested that langvlei acts as an intermediary or transition zone between the fresher eilandvlei and more saline rondevlei. numerous authors have suggested that the timing, frequency and duration of mouth opening events in temporarily open and closed estuaries affect species composition, particularly of marine species (beckley 1984; griffiths 1998; kok & whitfield 1986; wallace & van der elst 1975; young et al. 1997). in the warm-temperate east kleinemonde estuary, the timing of mouth opening had a significant effect on species composition (james, whitfield & cowley 2008). spring opening allowed for the regular recruitment of marine species into the estuary, many of which start entering cape estuaries in late winter (august), with peak recruitment occurring during spring (september–november) (whitfield 1998). the duration and frequency of mouth opening within a particular season appeared to have little impact on species composition in the east kleinemonde estuary (james et al. 2008). during hall’s (1985) study, the estuary was open for 86% of the time, particularly during peak recruitment periods (late winter and spring) (figure 2a), and mullet species in the touw estuary and eilandvlei were dominated by 0+ juveniles (> 50 mm sl), indicating good recruitment had occurred. in addition, hall (1985) recorded 32 species from the system. four of these species, blacktail (diplodus sargus capensis), strepie (sarpa salpa) (sampled in the touw estuary), cape stumpnose (eilandvlei) and super klipfish (clinus superciliosus) (rondevlei), were not sampled during this study. in contrast, the estuary only opened for 32 days (8% of the time) during this study between april 2010 and april 2011, and this was in summer and autumn (figure 2b). although the overall fish community composition was similar between the two periods, with catches dominated numerically by estuarine resident species, mullet and mozambique tilapia, there were no 0+ juvenile southern mullet (< 180 mm, de villiers 1987), cape stumpnose (< 100 mm, blaber 1974), freshwater mullet (< 100 mm, bok 1984), spotted grunter (< 150 mm, wallace 1975) or white steenbras (< 120 mm, bennett 1989) recorded. this indicates poor recruitment into the system during this period (figure 6). russell (1996) sampled the fish communities of both the wilderness and swartvlei lake systems and found that breaching of the estuaries during optimum fish recruitment periods resulted in an increase in the abundance of 0+ juveniles in both systems. the duration of mouth opening had little impact on successful recruitment. figure 6: comparison of the fish communities between a predominantly open phase (1982–1983) and a predominantly closed phase (2010–2011). the widespread distribution and abundance of one extra-limital and two alien invasive freshwater fishes is potentially concerning. although mozambique tilapia and western mosquitofish have been in the system for some time, the more recent introduction and spread of common carp in particular could have negative consequences. because of its benthic foraging behaviour, the common carp resuspends particulate matter, which increases turbidity leading to decreased light penetration and submerged macrophyte abundance (parkos, santucci & wahl 2003). in addition, the foraging behaviour can increase the available nitrogen, ammonia and phosphorus available in the water column (chumchal, nowlin & drenner 2005), potentially promoting algal blooms and further environmental degradation (khan, wilson & khan 2003; weber & brown 2009). a long-term water quality dataset for the wilderness lakes (russell 2013) indicates a significant decrease in salinity in the upper lakes; if this continues it may improve conditions for these three species at the expense of endemic euryhaline marine and estuarine resident species. future research work should shift towards understanding the ecological role and impacts of these species on the system. conservation implications improving or maintaining the ecological functioning of the wilderness lakes requires management actions to halt observed directional changes in water quality trends and improve connectivity between lakes. in particular, as in other systems, the structure of the marine fish community is determined by the timing of the mouth opening. in addition, the community is also influenced by the degree of flow and the ease of movement between lakes. the directional change in water chemistry from an estuarine to a more lacustrine system noted by russell (2013) needs to be halted. recognition of the national and international importance of the system dictates that the artificial mouth opening needs to take into consideration biodiversity and ecosystem processes and should only occur at the required height of 2.1 m above mean sea level. premature breaching due to human pressure (flooding risk) will only continue to exacerbate the trends in water quality and overall ecosystem functioning. to ensure adequate water flow and fish recruitment throughout the system, restrictions (sediment deposition) within the connecting channels need to be assessed and, where possible, removed. in addition, the dense macrophyte stands occurring within these channels not only restrict water movement but also could limit fish movement. both submerged and emergent aquatic plants within the channels should be regularly cut. importantly, the cut plants should be removed from the channel to prevent possible anoxic conditions during decomposition. monitoring of fish recruitment within and throughout the system should continue to assess efficacy of management actions, whilst research should focus on the impacts of the invasive fish species on endemic species and broader ecosystem functioning. acknowledgements the national research foundation of south africa and the dst-nrf centre of excellence in invasion biology are thanked for financial support. this research was conducted under daff/dea permit res2010/61 and was a sanparks registered research project. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions a.a.o. was responsible for data collection as part of her msc project, writing the research methods and design section and editing the manuscript. n.c.j. was responsible for data analysis and calculations, writing and editing the manuscript. m.k.s.s. contributed to study conceptualisation, study design, data capture and manuscript editing. o.l.f.w. was the project leader and contributed to study conceptualisation, funding, study design, data collection and editing the manuscript. references allanson, b.r. & whitfield, a.k., 1983, the limnology of the touw river floodplain, south african national scientific programmes report no. 79, council for scientific and industrial research, pretoria. beckley, l.e., 1984, ‘the ichthyofauna of the sundays estuary, south africa, with particular reference to the juvenile marine component’, estuaries 7, 248–258. http://dx.doi.org/10.2307/1352145 bennett, b.a., 1989, ‘a comparison of the fish communities in nearby permanently open, seasonally open and normally closed estuaries in the south-western cape, south africa’, south african journal of marine science 8, 43–55. bennett, b.a., hamman, k.c.d., branch, g.m. & thorne, s., 1985, ‘changes in the fish fauna of the bot river estuary in relation to opening and closure of the estuary mouth’, transactions of the royal society of south africa 45, 449–464. http://dx.doi.org/10.1080/00359198509519503 blaber, s.j.m., 1974, ‘the population structure and growth of juvenile rhabdosargus holubi (steindachner) (teleostei: sparidae) in a closed estuary’, journal of fish biology 6, 455–460. http://dx.doi.org/10.1111/j.1095-8649.1974.tb04561.x blaber, s.j.m., 1978, ‘fishes of the kosi system’, lammergeyer 24, 28–41. bok, a.h., 1979, ‘the distribution and ecology of two mullet species in some fresh water rivers in the eastern cape, south africa’, journal of the limnological society of south africa 5, 97–102. http://dx.doi.org/10.1080/03779688.1979.9633193 bok, a.h., 1984, ‘freshwater mullet in the eastern cape – a strong case for fish ladders’, the naturalist 28, 31–35. chumchal, m.m., nowlin, w.h. & drenner, r.w., 2005, ‘biomass-dependent effects of common carp on water quality in shallow ponds’, hydrobiologia 545, 271–277. http://dx.doi.org/10.1007/s10750-005-3319-y day, j.h., blaber, s.j.m. & wallace, j.h., 1981, ‘estuarine fishes’, in j.h. day (ed.), estuarine ecology with particular reference to southern africa, pp. 197–221, a.a. balkema, cape town. de villiers, g., 1987, ‘harvesting harders liza richardsoni in the benguela upwelling region’, south african journal of marine science 5, 851–862. http://dx.doi.org/10.2989/025776187784522540 ellender, b.r., weyl, o.l.f., shanyengange, m.k. & cowley, p.d., 2008, ‘juvenile population dynamics of oreochromis mossambicus in an intermittently open estuary at the limit of its natural distribution’, african zoology 43(2), 277–282. http://dx.doi.org/10.3377/1562-7020-43.2.277 griffiths, s.p., 1998, ‘diversity and distribution of fishes in an intermittently open coastal lagoon at shellharbour, new south wales’, wetlands (australia) 18, 13–24. hall, c.m., 1985, ‘some aspects of the ecological structure of a segmented barrier lagoon system with particular reference to the distribution of fishes’, msc thesis, department of ichthyology & fisheries science, rhodes university. hall, c.m., whitfield, a.k. & allanson, b.r., 1987, ‘recruitment, diversity and the influences of constrictions on the distribution of fishes in the wilderness lakes system, south africa’, south african journal of zoology 22, 163–168. harrison, t.d., 2005, ‘ichthyofauna of south african estuaries in relation to the zoogeography of the region’, smithiana bulletin 6, 1–27. harrison, t.d. & whitfield, a.k., 2004, ‘a multi-metric fish index to assess the environmental condition of estuaries’, journal of fish biology 65, 683–710. http://dx.doi.org/10.1111/j.0022-1112.2004.00477.x howard-williams, c. & liptrot, m.r.m., 1980, ‘submerged macrophyte communities in a brackish south african estuarine-lake system’, aquatic biology 9, 101–116. james, n.c., cowley, p.d. & whitfield, a.k., 2007a, ‘abundance, recruitment and residency of two sparids in an intermittently open estuary in south africa’, african journal of marine science 29, 527–538. http://dx.doi.org/10.2989/ajms.2007.29.3.18.348 james, n.c., cowley, p.d., whitfield, a.k. & lamberth, s.j., 2007b, ‘fish communities in temporarily open/closed estuaries from the warmand cool-temperate regions of south africa: a review’, reviews in fish biology and fisheries 17, 565–580. james, n.c. & harrison, t.d., 2008, ‘a preliminary survey of the estuaries on the south coast of south africa, cape st blaize, mossel bay robberg peninsula, plettenberg bay, with particular reference to the fish fauna’, transactions of the royal society of south africa 63, 111–127. http://dx.doi.org/10.1080/00359190809519216 james, n.c., whitfield, a.k. & cowley, p.d., 2008, ‘long-term stability of the fish assemblages in a warm-temperate south african estuary’, estuarine, coastal and shelf science 76, 723–738. http://dx.doi.org/10.1016/j.ecss.2007.07.036 khan, t.a., wilson, m.e. & khan, m.t., 2003, ‘evidence for invasive carp mediated trophic cascade in shallow lakes of western victoria, australia’, hydrobiologia 506, 465–472. kok, h.m. & whitfield, a.k., 1986, ‘the influence of open and closed mouth phases on the marine fish fauna of the swartvlei estuary’, south african journal of zoology 21, 309–315. http://dx.doi.org/10.1080/02541858.1986.11448004 maree, r.c., whitfield, a.k. & quinn, n.w., 2003, prioritisation of south african estuaries based on their potential importance to estuarine-associated fish species, wrc report no. tt 2003/03, water research commission, pretoria. olds, a.a., smith, m.k.s., weyl, o.l.f. & russell, i.a., 2011, ‘invasive alien freshwater fishes in the wilderness lakes system, a wetland of international importance in the western cape province, south africa’, african zoology 46, 179–184. parkos, j., santucci, v.c. & wahl, d., 2003, ‘effects of common carp (cyprinus carpio) on multiple torphic levels in shallow mewocosms’, canadian journal of fisheries and aquatic sciences 60, 182–192. http://dx.doi.org/10.1139/f03-011 potter, i.c., beckley, l.e., whitfield, a.k. & lenanton, r.c.j., 1990, ‘comparisons between the roles played by estuaries in the life cycles of fishes in temperate western australia and southern africa’, environmental biology of fishes 28, 143–178. http://dx.doi.org/10.1007/bf00751033 potter, i.c. & hyndes, g.a., 1999, ‘characteristics of the ichthyofaunas of the southwestern australian estuaries, including comparisons with holarctic estuaries and estuaries elsewhere in temperate australia: a review’, australian journal of ecology 24, 395–421. http://dx.doi.org/10.1046/j.1442-9993.1999.00980.x prchalová, m., kubečka, j., říha, m., mrkvička, t., vašek, m., jůza, t., et al., 2009, ‘size selectivity of standardized multimesh gillnets in sampling coarse european species’, fisheries research 96, 51–57. http://dx.doi.org/10.1016/j.fishres.2008.09.017 ratte, t.w., 1989, ‘population structure, production, growth, reproduction and the ecology of atherina breviceps valenciennes, 1935 (pisces: atherinidae) and gilchristella aestuaria (gilchrist, 1914) (pisces clupeidae), from two southern cape coastal lakes’, phd thesis, zoology department, university of port elizabeth. russell, i.a., 1996, ‘fish abundance in the wilderness and swartvlei lake systems: changes relative to environmental factors’, south african journal of zoology 31, 1–9. http://dx.doi.org/10.1080/02541858.1996.11448389 russell, i.a., 1999, ‘freshwater fish of the wilderness national park’, koedoe 42, 73–78. http://dx.doi.org/10.4102/koedoe.v42i1.223 russell, i.a., 2003, ‘changes in the distribution of emergent aquatic plants in a brackish south african estuarine-lake system’, african journal of aquatic science 28, 103–122. http://dx.doi.org/10.2989/16085910309503776 russell, i.a., 2013, ‘spatio-temporal variability of surface water quality parameters in a south african estuarine lake system’, african journal of aquatic science 38, 53–56. http://dx.doi.org/10.2989/16085914.2012.742006 sanparks, 2012, garden route national park – management plan, viewed 07 october 2015, from http://www.sanparks.org/docs/parks_grnp/planning_products/management_plan.pdf scott, k.m.f., harrison, a.d. & macnae, w., 1952, ‘the ecology of south african estuaries. part 2: the klein river estuary, hermanus, cape’, transactions of the royal society of south africa 33, 282–331. sloterdijk, h., james, n.c., smith, m.k.s., ekau, w. & weyl, o.l.f., 2015, ‘population dynamics and biology of an invasive population of mosquitofish gambusia affinis in a temperate estuarine lake system’, african zoology 50(1), 31–40. talbot, m.m.j.f., 1982, ‘aspects of the ecology and biology of gilchristella aestuarius (g & t) (pisces: clupeidae) in the swartkops estuary’, msc thesis, zoology department, university of port elizabeth. turpie, j.k., adams, j.b., joubert, a., harrison, t.d., colloty, b.m., maree, r.c., et al., 2002, ‘assessment of the conservation priority status of south african estuaries for use in management and water allocation’, water sa 28, 191–206. http://dx.doi.org/10.4314/wsa.v28i2.4885 van der horst, g. & erasmus, t., 1981, ‘spawning time and spawning grounds of mullet with special reference to liza dumerili (steindachner, 1869)’, south african journal of science 77, 73–78. vorwerk, p.d., whitfield, a.k., cowley, p.d. & paterson, a.w., 2001, ‘a survey of selected eastern cape estuaries with particular reference to the ichthyofauna’, ichthyological bulletin of the j.l.b. smith institute of ichthyology 72, 1–52. vorwerk, p.d., whitfield, a.k., cowley, p.d. & paterson, a.w., 2003, ‘the influence of selected environmental variables on fish assemblage structure in a range of southeast african estuaries’, environmental biology of fishes 66, 237–247. http://dx.doi.org/10.1023/a:1023922521835 wallace, j.h., 1975, ‘the estuarine fishes of the east coast of south africa. part 1. species composition and length distribution in the estuarine and marine environments. part 2. seasonal abundance and migrations’, investigational report of the oceanographic research institute 40, 1–72. wallace, j.h. & van der elst, r.p., 1975, ‘the estuarine fishes of the east coast of south africa. part 4. occurrence of juveniles in estuaries. part 5. ecology, estuarine dependence and status’, investigational report of the oceanographic research institute 42, 1–63. weber, m.j. & brown, m.l., 2009, ‘effects of common carp on aquatic ecosystems 80 years after “carp as a dominant”: ecological insights for fisheries management’, reviews in fisheries science 17, 524–537. http://dx.doi.org/10.1080/10641260903189243 weisser, p.j. & howard-williams, c., 1982, ‘the vegetation of the wilderness lakes system and the macrophyte encroachment problem’, bontebok 2, 19–40. whitfield, a.k., 1994, ‘an estuary-association classification for the fishes of southern africa’, south african journal of science 90, 411–417. whitfield, a.k., 1998, biology and ecology of fishes in southern african estuaries, ichthyological monographs of the j.l.b. smith institute of ichthyology, no 2, grahamstown. whitfield, a.k. & blaber, s.j.m., 1979, ‘the distribution of the freshwater cichlid sarotherodon mossambicus in estuarine systems’, environmental biology of fishes 4, 77–81. http://dx.doi.org/10.1007/bf00005931 whitfield, a.k. & elliott, m., 2002, ‘fishes as indicators of environmental and ecological changes within estuaries: a review of progress and some suggestions for the future’, journal of fish biology 61(suppl a), 229–250. http://dx.doi.org/10.1111/j.1095-8649.2002.tb01773.x whitfield, a.k. & kok, h.m. 1992. ‘recruitment of juvenile marine fishes into permanently open and seasonally open estuarine systems on the southern coast of south africa’, ichthyological bulletin, 57, 1–15. young, g.c., potter, i.c., hyndes, g.a. & de lestang, s., 1997, ‘the ichthyofauna of an intermittently open estuary: implications of bar breaching and low salinities on faunal composition’, estuarine, coastal and shelf science 45, 53–68. http://dx.doi.org/10.1006/ecss.1996.0165 article information authors: llewellyn c. foxcroft1,2 melodie mcgeoch2,3 affiliations: 1south african national parks, conservation services, skukuza, south africa 2centre for invasion biology, stellenbosch university, south africa 3south african national parks, cape research centre, cape town, south africa correspondence to: llewellyn foxcroft email: llewellyn.foxcroft@sanparks.org postal address: private bag x402, skukuza 1350, south africa dates: received: 31 may 2010 accepted: 14 jan. 2011 published: 13 may 2011 how to cite this article: foxcroft, l.c. & mcgeoch, m., 2011, ‘implementing invasive species management in an adaptive management framework’, koedoe 53(2), art. #1006, 11 pages. doi:10.4102/koedoe.53i2.1006 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) implementing invasive species management in an adaptive management framework in this essay... open access • abstract • introduction    • need for an invasive alien species policy and management framework    • need for flexibility and rapid response for management of invasive alien species • constructing the foundations: developments to date • a framework for advancing adaptive management of invasive alien species    • sanparks vision and mission statements (1)    • park management plan (2)    • management forum (3)    • monitoring (4)       • long-term trends in ecosystem change       • operational management effectiveness       • project auditing    • indicators and thresholds (5)    • invasive alien species management programmes (6)    • multiple feedback processes (7)    • conclusion    • acknowledgements • references • supplementum abstract (back to top) adaptive management theory has attracted substantial interest in recent years, in natural resource management in general and also for invasive alien species management. however, whilst many theoretical and conceptual advances have been made, documented cases of practical applications are rare. coupling invasive species management components with adaptive feedback processes is not without challenges, requiring a substantial change in the thinking and practice of all those involved. drawing on a decade of experience in south african national parks, we suggest an approach to implementing adaptive management for controlling invasive alien species. whilst efforts have been made to advance components of the overall management strategy, the absence of a framework for decision making and feedback mechanisms, inflexibility in the system and shortcomings in the governance structure are all identified as barriers to learning and knowledge integration for the purposes of effective invasive alien species management. the framework provided here, encompassing documents, committees and processes, is aimed at addressing these shortcomings. conservation implication: adaptive management theory offers a robust tool for managing inherently complex systems. its practical application, however, requires distilling the theory into useable functions. we offer a framework to advance implementation of strategic adaptive management for the control of invasive alien species using experiences gained from south african national parks. introduction (back to top) ecosystems are dynamic, changing in both space and time and over multiple scales (pickett, cadenasso & benning 2003). these fluctuations are both necessary and desirable. adaptive management (am) has emerged as a preferred method of management for natural resources in the face of such change and inherent uncertainty (biggs & rogers 2003; gregory, ohlson & arvai 2006; jacobson et al. 2006; salafsky, margoluis & redford 2001; scholes & kruger 2011). accordingly, south african national parks (sanparks) has adopted the principles of am to guide decision making (rogers 2003). am generally includes the following steps: (1) developing a conceptual understanding of the social-ecological system, (2) developing objectives and management plans, (3) implementing management, (4) monitoring trends and system changes, (5) analysing data, and (6) purposefully modifying management in response to outcomes where necessary (e.g. salafsky et al. 2001). strategic adaptive management (sam), as applied in sanparks, encompasses the general am process, but differs by focussing more explicitly on (1) developing a mutual understanding and appreciation by stakeholders of the way ecosystem goods and services are valued, (2) building an understanding of the social, technological, economic, environmental and political context and (3) establishing an adaptive assessment process for defining spatially and temporally explicit monitoring programmes that assess the response of ecosystems to management actions (biggs & rogers 2003; roux & foxcroft 2011). the focus on these aspects is an attempt to overcome some of the barriers (communication, attitudinal and conceptual) to am implementation (jacobson et al. 2006). a further contribution that sanparks has made to am theory has been the development of thresholds of potential concern (tpcs) (roux & foxcroft 2011). based on best available science and updated as new understanding emerges, tpcs allow for some measure of system flux whilst highlighting points in a trend that reflect unacceptable environmental consequences and therefore require management interventions. as shown in figure 1, a three-dimensional space represents possible changes in the system attribute of interest over time and space. the sphere represents the outer boundaries of the desirable system, which are represented by tpcs. any directional change in the system within the boundaries (or tpcs) is considered acceptable – even desirable – ecosystem flux (blue to green arrow). should the system be outside or on a trajectory leading out of the desired set of conditions (blue or green to orange arrow) – a tpc thus having been breached – an assessment of the cause of the system change and possible management interventions are called for. for alien species, tpcs are breached when new alien species are introduced or species already in the system spread (red arrow). these thresholds are low because once highly invasive species have established, they are likely to spread over a wide area, with long-lasting impacts on the system attribute of interest (e.g. plant diversity in riparian areas). the use of tpcs creates an opportunity to implement management strategies as landscape scale experiments, from which the outcomes of the management interventions must necessarily be assessed. tpcs also represent another difference between general am approaches and sam as applied in sanparks, in that tpcs aim to pre-empt impending unacceptable system change and thus allow for proactive management options (scholes & kruger 2011). long-term monitoring aims to detect these trends and changes in various ecological features (buckley et al. 2008) and is therefore an essential component in the implementation of am (rogers 2003). the outcome of monitoring and analysis is measured against predefined indicators and associated tpcs (biggs & rogers 2003). this assessment provides feedback on the outcomes of management actions and, in the process, provides both a formal and an informal means of learning and knowledge diffusion (biggs & rogers 2003). most ecosystems in south africa have been invaded by a large number of taxonomic groups (richardson et al. 2000), resulting in a wide range of negative impacts on biodiversity and social-economic systems (pyšek & richardson 2010). the most well-documented, ecologically harmful taxa are plants, many of which are woody plants (richardson, rouget & rejmánek 2004). although conservation areas are protected from some threats (e.g. large-scale habitat transformation or over-exploitation of resources), other threats are more difficult to exclude (kareiva et al. 2007; millennium ecosystem assessment 2005; woodroffe & ginsburg 1998). globally, invasive alien species are a substantial and escalating threat to the integrity of many ecosystems within protected areas (mea 2005). this can be attributed, in part, to the fact that protected areas often exist as islands in a sea of diverse, human-dominated land uses (pickett & thompson 1978), where disturbances and propagule sources are strong drivers of biological invasions (hobbs & huenneke 1992). for example, the kruger national park (knp), with an area of approximately 20 000 km2, has been invaded by about 370 alien plants; 27 000 plant records having been collected (foxcroft et al. 2009). knp also has a history of alien species management policies based either on national legislation (initially the conservation of agricultural resources act; act 43 of 1983, and later amendments in 2001) or on the internal needs of the park (e.g. a policy on the use and management of alien ornamental plants as described by foxcroft [2001] and foxcroft and freitag-ronaldson [2007]). more recently, the development of the national environmental management: biodiversity act (act 10 of 2004) and the national environmental management: protected areas act (act 57 of 2003) has determined the legislative context within which all management and monitoring must take place, including those for invasive alien species. this, together with the experience gained in sanparks (and knp) over the last decade in applying sam to invasive alien species (bestbier, jacoby & rogers 2000; foxcroft & richardson 2003; foxcroft & downey 2008; foxcroft 2009), provides a useful platform for proposing future management options. however, three key shortcomings are considered to exist in the current approaches to management of invasive alien species in sanparks. these are forms of institutional barriers to the implementation of am (jacobson et al. 2006) and include (1) the lack of a broad framework for decision making, (2) the inflexibility in existing management systems and administration and (3) shortcomings in the governance structure for invasive alien species management. figure 1: the threshold of potential concern may be represented as a threedimensional space, showing possible changes in the system attribute of interest over time and space. figure 2: an overview of the adaptive management process for an overall invasive alien species management framework. each number in the figure is related to the specific point in the ensuing discussion. the monitoring programme (number 4) is further expanded on in figure 3. need for an invasive alien species policy and management framework although frameworks for implementing sam for alien species management have been proposed (bestbier et al. 2000; freitag-ronaldson & foxcroft 2003), emphasis has been largely on developing specific tpcs to date. since 1999, 37 cases of breached thresholds have been documented (supplementary material (table 2) in foxcroft [2009]). in some cases the lack of clarity of departmental responsibility for responding to the breached tpc led to confusion, delaying management action. thus, although the thresholds were designed to trigger appropriate interventions (mainly rapid eradication or containment strategies as described by foxcroft [2009]), the lack of an overall policy and management framework within which they could operate impeded a coordinated response. moreover, although the tpcs include monitoring for new incursions of alien species into parks, almost all attention has been on control, with little focus on prevention. although this is a global trend (keller & lodge 2010), a substantial increase in focus on prevention is essential if parks are to manage invasive alien species successfully in the long term. in a large natural system such as the knp, where many alien species have been introduced (foxcroft et al. 2003), not all species can or necessarily need to be managed. some alien species will also experience fluctuations in their population density and spatial distribution over time, although the most invasive and potentially damaging species will tend to become, and remain, highly abundant and widespread (freitag-ronaldson & foxcroft 2003:414). it is these ‘transformer’ species that alter the landscape over a wide area, often in both structure and function (pyšek et al. 2004), that are of most concern and thus the focus of management attention. without an overarching framework and coordinated effort to identify where and when tpcs are required (e.g. tu 2009), and to respond when they are breached, efforts will remain largely opportunistic and potentially ineffective (hall & fleishman 2009). another aspect that requires further consideration is adaptive governance, which is the process by which social systems are organised or reorganise themselves (folke et al. 2005). governance provides the structures and processes needed in which collective decisions are taken. adaptive governance networks join teams that contain different experiences to develop a common understanding of management and policies. the governance networks function across multiple levels within organisations, between organisations and between individuals (folke et al. 2005). these networks provide for enhanced flexibility (a requisite for rapid response to invasions), but still operate within the organisational hierarchy, holding specific agencies, management levels or individuals accountable for inaction or mismanagement. in discussing biosecurity risks of invasive alien species in australia, cook et al. (2010:1308) discuss governance structures for interagency cooperation to improve information transfer for surveillance, response, diagnostics, risk analysis and border inspections. these functions are carried out by a number of organisations in a number of different ways. similarly, we suggest that greater focus on the governance structure of invasive alien species management is required for improved utility of the framework we present. an organisational policy for alien and invasive species management will, for example, contribute significantly to the governance of management of invasive alien species in sanparks. need for flexibility and rapid response for management of invasive alien species globally, management of invasive alien species has developed in a number of forms, including risk assessment, vector and pathway control, early detection and rapid response (pyšek & richardson 2010). although eradication is touted as the main aim of control, containment and maintenance of target populations at low levels are often the only feasible option (rejmánek & pitcairn 2002), particularly with transformer species. as part of a containment strategy, best management practices include setting fixed goals, delineating fixed areas, listing priority species for control, applying available resources to these areas, and follow-up operations, all of which are commonly planned on an annual time scale (eds. wittenberg & cock 2001). these are all an essential part of the overall planning process, but the associated rigidity does not always allow for the surprises that are common features in dynamic ecological systems. examples of such unforeseen events include large, infrequent disturbances, such as floods and fires. in response to these disturbance events, invasive species that were previously being maintained at a low abundance or contained within a small area may re-emerge rapidly and over a large area after a long period of apparent stability (e.g. crooks 2005). the events given in these examples are likely to cause the tpcs to be exceeded, requiring immediate assessment and action rather than reliance on a set of fixed medium-term or long-term goals. moreover, such events may necessitate complete reorganisation of control projects, either to take advantage of a particular situation or to mitigate undesirable changes. this process requires reassessing the order of planned control actions, revising priorities and redirecting resources, often at relatively short notice. in large bureaucracies this is a time-consuming process, which is likely to delay or prohibit opportunities for eradication or early containment. consistent long-term management and control of invasive alien species is essential. should follow-up operations cease, all the resources used and advances achieved in clearing a particular area may be lost in a very short time. we suggest here that an additional layer of planned management action, designed for and focussed on enabling rapid response to unexpected changes, should be implemented. successful management of invasive alien species necessarily includes effective rapid response along with consistent clearing and follow-up operations. we also aim here to place these actions, together with those already in practice, in their broader framework and context. whilst management action in sanparks has to date been virtually aimed exclusively at invasive alien plants, the proposal outlined here is generally relevant to other alien taxa. constructing the foundations: developments to date (back to top) sanparks has invested substantially in adopting and further developing am principles and a number of unique constructs have been instituted. notably, these include a hierarchy of objectives (cascading down from the organisational vision to ground level or individual goals) to structure park management priorities and the use of decision thresholds. although a number of management functions are in place, albeit in varying stages of development, they have largely functioned in isolation of one another to date. these components include: • the sanparks vision and individual park mission statements • park management plans, comprising the park-specific objectives • lower level plans (llps), which are specific operational plans for all management concerns in a park; the llps stem from the park management plan objectives and are written, for example, for fire management, invasive alien species management, artificial water provision, rehabilitation and many other aspects of protected area management • management unit clearing plan (mucp) and annual plan of operation (apo); these documents are medium-term to short-term plans used by the sanparks invasive species control unit (iscu) to prepare and guide project implementation and independent contractor management • physical implementation of control programmes for invasive alien species by iscu and/or park management • indicators and tpcs, which, for the purposes of invasive alien species management, have largely been decision thresholds (i.e. mtpcs as in foxcroft [2009]), indicating points at which management decisions and intervention are usually required; these have only been used in a few parks to date • fora where selected invasive alien species management issues are discussed intermittently; however, in most cases this dialogue has taken place informally between park managers or rangers and the iscu project managers, but needs to be complemented by more formal discourse • functional feedback loops, which are inherent in am processes (stafford-smith et al. 2009) and are aimed at enhancing understanding and ensuring continuous improvement in management effectiveness. together, the listed objectives, plans, interactions and processes should complement one another in the functions they perform, and thus form a foundation for consolidating effective management of alien and invasive species. the relative roles of these management functions and the relationships between them should also be clear. however, a number of gaps remain and further components are necessary, such as a coordinated monitoring programme to enhance the optimal implementation of an overall response to invasive alien species management and efficient feedback mechanisms. here we identify these gaps and suggest ways to address them and also clarify the relative roles and relationships between each of the management functions. finally, we suggest how the various components may be integrated into an overall sam framework for invasive alien species management. a framework for advancing adaptive management of invasive alien species (back to top) conceptual frameworks are tools to organise and evaluate the state of a particular system (pickett, kolasa & jones 2007). a framework provides a way of deriving a common understanding of a system or process, whilst also indicating various levels of feedback and learning (tu 2009). the schematic diagrams we use to illustrate the framework that is required for an overall, coordinated approach to managing invasive alien species (figures 2 and 3) are embedded within the overarching sam paradigm in use by sanparks. we suggest specific components in the framework and illustrate explicit pathways between these components. we further provide a detailed description of each aspect of the management cycle, highlighting their key attributes. the number shown in parentheses at each subsection in the ensuing discussion relates to the numbered boxes in figures 2 and 3. although a broader context informs the development of each step (e.g. deriving vision statements and vital attributes at the organisational or park scale [biggs & rogers 2003]), we assume these processes have taken place and frame our discussion in the more focussed context of management of invasive alien species. sanparks vision and mission statements (1) the sanparks vision and mission statements provide the overall direction that guides all decision making. this cascades down to park level, where the vital attributes of the park steer local decision making. for example, the main focus of mapungubwe national park as a world heritage site is based on its rich cultural heritage (south african national parks 2006). the knp is a representative of a relatively intact large savanna ecosystem, with various processes and species that interact over large spaces and long time frames (pickett et al. 2003), whilst table mountain national park supports a significant portion of rare and endemic taxa forming part of the globally renowned cape floristic region (cowling et al. 2003). these attributes guide the management focus of each park. park management plan (2) the park management plan, a legislative requirement in south africa for all protected areas (national environmental management: protected areas act), provides park-specific context and guides actions. the components of the park management plans are hierarchical (e.g. biggs & rogers 2003:63) and are sequentially split into themes, which are then expanded upon in llps. in the context of invasive alien species, the llp presents a risk assessment of species that present the greatest threat or concern, the planned management interventions, financial and resource-related requirements, monitoring, and research needs. therefore the llp essentially forms the roadmap for managing invasive alien plants in a park over a five-year cycle. the llp is, in turn, translated into an mucp (a five-year plan that runs concurrently with the llp) and from there to an apo. the apo provides details of each work area to be contracted out for clearing, specific details of the areas to be cleared (with maps), species lists and information about species abundance, human resources, control methods and other project-specific details. management forum (3) a forum for each park (commonly referred to as a science–management forum), should be the nexus for integrating all components of the invasive alien species management cycle. this forum includes park managers, staff of the iscu, project managers and regional ecologists and also receives input from specialist researchers where needed. the development of the park plans, specifically the llps and apos, should be formulated by this group – a process which has happened very rarely. ideally, these fora should take place at least biannually to reassess progress and revise plans, thereby providing feedback to all. the proposed planning process would contribute to achieving a common understanding of the overall invasive alien species control programme. moreover, experience of all role players and short-term internal feedback loops between park managers, iscu project managers and the regional ecologists, are essential for adapting and improving operational plans to increase their flexibility and effectiveness. monitoring (4) monitoring is the cornerstone on which all forms of am rest (salafsky et al. 2002). this is because am requires implementing management actions as experiments. as a consequence, monitoring is needed to determine how the system responds to management (biggs & rogers 2003). the term ‘monitoring’ is not used consistently, leading to subsequent confusion about applying monitoring methods in a particular situation (see dewey & anderson 2004). we differentiate between the following: 1. monitoring long-term trends in the ecosystem as a response to management interventions 2. assessing operational or management effectiveness 3. project auditing (stem et al. 2005). long-term trends in ecosystem change such monitoring assesses the ecological outcome of management efforts. here we refer to monitoring as the ‘collection and analysis of repeated observations or measurements to evaluate changes in condition and progress toward meeting a management objective’ (elzinga et al. 2001). the main objective of the sanparks monitoring programme for alien and invasive species is to assess the status of and trends in invasion, distribution and management of alien species in sanparks, such that the information contributes to management and policy development at a local as well as global level. the approach forms part of the sanparks biodiversity monitoring system (mcgeoch et al. 2011). in order to achieve this higher objective, three subobjectives are proposed as outlined in the following: • subobjective 1: to determine and monitor the rate of introduction of all alien species into the sanparks estate. the main target of this objective is to prevent the introduction of any new invasive alien species into a park. • subobjective 2: to determine and monitor the distribution of existing naturalised and invasive species within and across parks. the target of this objective is to contain the spread of invasive species that are already present. • subobjective 3: to determine and monitor the increase in the abundance and distribution of naturalised and invasive alien species in parks (with distribution used as a predictor and proxy for abundance, as proposed by he and gaston [2000]). the target of this objective is to maintain the abundance of invasive species below an acceptable threshold. operational management effectiveness monitoring aims to provide rapid feedback on control activities that will allow project managers to assess progress and strategically adapt their plans as necessary. owing to the vast areas being controlled across the sanparks estate, this will most likely have to be a rapid assessment of a few randomly selected sites (see stem et al. 2005). within a management unit, data may be collected on, for example, whether a target species has been cleared, whether plants have been missed, are coppicing (resprouting), and whether there have been any negative impacts on nontarget species. in addition, distribution data should be collected opportunistically for any newly observed species or an increase in the distribution of a listed species. project auditing in this case monitoring aims to assess the implementation of the project and procedures (stem et al. 2005). these include logistic arrangements, contractor implementation and contractual obligations, occupational health and safety compliance, compliance with herbicide use policies and other aspects which directly relate to the smooth implementation of projects. many of these components are legislative requirements and need to be performed according to a fixed schedule. indicators and thresholds (5) tpcs have long been under development and refinement in the knp (biggs & rogers 2003) and more recently also in other parks. foxcroft (2009) provides the theoretical background to the current suite of tpcs for invasive alien species. the use of indicators and their associated tpcs provide endpoints against which monitoring of the response of an ecosystem to management intervention may be assessed. in other words, indicators are dependent variables (e.g. the number of alien species or area of a park invaded by invasive alien species), whereas a tpc is the specific value(s) of that indicator for which well-considered intervention is needed (e.g. the introduction of one new alien species to a park or a 10% increase in the area invaded). the indicators should be assessed according to a 3–5-year time frame and any thresholds which have been breached at this point should be raised at the relevant park forum (figure 3). if a tpc has been breached at any other time, it should also be documented and presented at the appropriate science–management forum. in this way any unacceptable trends that suggest potential deleterious ecological or biodiversity consequences are given specific attention. using tpcs as a reference point for management interventions does not imply that ongoing control operations are stopped and initiated only when a tpc is breached. rather, parallel to ongoing control operations, tpcs are used to highlight the threshold in a trend where undesirable impacts are expected (foxcroft 2009) and where additional control, or reorganising and reprioritising, of projects is required. management actions should, therefore, aim to prevent the system from reaching such thresholds. where a tpc is breached, the background, cause of concern, potential impacts or consequences and proposed management actions should be discussed. although tpcs represent hypotheses that are open to challenge, testing and refinement at any time, the validity of the tpc should not be challenged once a breach of a tpc has been tabled (biggs & rogers 2003). this is to prevent ‘management paralysis’, where the threshold or indicator is continuously questioned and further assessments and research are called for, instead of decisive management action being taken (biggs & rogers 2003). invasive alien species management programmes (6) in sanparks, the iscu is largely responsible for the implementation of invasive alien species management programmes, although park managers, rangers, regional ecologists and scientists are all required to provide input at various stages. the iscu operations are largely funded by the national working for water programme (see van wilgen, le maitre and cowling [1998] and van wilgen, khan and marais [2010] for a discussion on the working for water programme), with sanparks providing management, financial, logistic and other support. multiple feedback processes one of the key features of all forms of am is multiple feedback loops between stages (biggs & rogers 2003; maris & béchet 2010; stafford-smith et al. 2009). continuous evaluation and self-assessment of all stages of the management process provide information on the outcome of previous interventions and allow for ongoing improvement (timko & innes 2009). further, purposeful reflection and assessment provide growing understanding of the desired management goal and progress impediment (venter et al. 2008). purposeful learning and continuous feedback between the various players involved in management of invasive alien species in sanparks have largely been absent to date, despite substantial efforts to enhance communication. we suggest that the schematic frameworks presented here outline a governance structure and provide clear linkages to assist role players in visualising the contribution of all components of invasive alien species management and the feedbacks required amongst them. figure 3: a schematic figure of the monitoring process (as a more detailed subset of figure 2). the aim of the monitoring programme is to detect the change and trends in the number and distribution of alien species in a predefined area. conclusion (back to top) institutionalising changes to an existing management paradigm is not without challenges. the framework we illustrated aims to combine and integrate management functions that are already in place or are being developed. despite a number of barriers (see jacobson et al. 2006) that will require ongoing assessment and reflection, the framework provides one approach for fully implementing am. in future, options should be explored for streamlining and simplifying the array of documents, bodies and/or committees and processes that constitute invasive alien species management (as reflected in the supplementary material, figure 1). the framework for invasive alien species management presented here formally links and establishes the relationships between various existing and some new management functions. implementation of such a framework, supported by a clearer governance structure, will better enable effective alien species management via improved flexibility of management operations and the speed at which management actions take place. it will also allow for the assessment of environmental effectiveness and management costs (hall & fleishman 2009). in the context of managing invasive alien species, sam should assist with integrating project management, park management and monitoring of invasive alien species and include more effective feedback mechanisms. we therefore suggest that the proposed framework provides a simple yet robust way of organising and connecting the various facets of invasive species management in sanparks. acknowledgements (back to top) l.c.f. and m.m. thank the dst–nrf centre of excellence in invasion biology, stellenbosch university for funding and support. this work was supported financially by the national research foundation, with grants to both l.c.f. and m.m. we thank two anonymous reviewers for their valuable insights and comments. references (back to top) bestbier, r.x., jacoby, d.l. & rogers, k.h., 2000, a goal maintenance system for the management of the kruger national park riverine alien vegetation: developing a protocol and a prototype, water research commission, pretoria. biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring, and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59–80, island press, washington dc. buckley, r., robinson, j., carmody, j. & king, n., 2008, ‘monitoring for management of conservation and recreation in australian protected areas’, biodiversity and conservation 17, 3589–3606. doi:10.1007/s10531-008-9448-7 cook, d.c., liu, s., murphy, b. & lonsdale, w.m., 2010, ‘adaptive approaches to biosecurity governance’, risk analysis 3(9), 1303–1314. doi:10.1111/j.1539-6924.2010.01439.x cowling, r.m., pressey, r.l., rouget, m. & lombard, a.t., 2003, ‘a conservation plan for a global biodiversity hotspot: the cape floristic region, south africa’, biological conservation 112(1–2), 191–216. doi:10.1016/s0006-3207(02)00425-1 crooks, j.a., 2005, ‘lag time and exotic species: the ecology and management of biological invasions in slow-motion’, ecoscience 12(3), 316–329. doi:10.2980/i1195-6860-12-3-316.1 dewey, s.a. & anderson, k.a., 2004, ‘distinct roles of surveys, inventories, and monitoring in adaptive weed management’, weed technology 18, 1449–1452. doi:10.1614/0890-037x(2004)018[1449:drosia]2.0.co;2 elzinga, c.l., salzer, d.w., willoughby, j.w. & gibbs, j.p., 2001, monitoring plant and animal populations, blackwell science, london. folke, c., hahn, t., olsson, p. & norberg, j., 2005, ‘adaptive governance of social-ecological systems’, annual review of environment and resources 30, 441–473. doi:10.1146/annurev.energy.30.050504.144511 foxcroft, l.c., 2001, ‘a case study of the human dimensions in invasion and control of alien plants in the personnel villages of kruger national park’, in j.a. mcneely (ed.), the great reshuffling: human dimensions of invasive alien species, pp. 127–134, iucn, gland. foxcroft, l.c., 2009, ’developing thresholds of potential concern for invasive alien species: hypotheses and concepts’, koedoe, 50(1), 6. foxcroft, l.c. & downey, p.o., 2008, ‘protecting biodiversity by managing alien plants in national parks: perspectives from south africa and australia’, in b. tokarska-guzik, j.h. brock, g. brundu, l. child, c.c. daehler & p. pyšek (eds.), plant invasions: human perception, ecological impacts and management, pp. 387–403, backhuys publishers, leiden. foxcroft, l.c. & freitag-ronaldson, s., 2007, ‘seven decades of institutional learning: managing alien plant invasions in the kruger national park, south africa’, oryx 41(2), 160–167. doi:10.1017/s0030605307001871 foxcroft, l.c. & richardson, d.m., 2003, ‘managing alien plant invasions in the kruger national park, south africa’, in l. child, j. brock, g. brundu, k. prach, p. pyšek, p. wade, et al. (eds.), plant invasions: ecological threats and management solutions, pp. 385–403, backhuys publishers, leiden. foxcroft, l.c., henderson, l., nichols, g.r. & martin, b.w., 2003, ‘a revised list of alien plants for the kruger national park’, koedoe 46(2), 21–44. foxcroft, l.c., richardson, d.m., rouget, m. & macfadyen, s., 2009, ‘patterns of alien plant distribution at multiple spatial scales in a large national park: implications for ecology, management and monitoring’, diversity and distributions 15, 367–378. doi:10.1111/j.1472-4642.2008.00544.x freitag-ronaldson, s. & foxcroft, l.c., 2003, ‘anthropogenic influences at the ecosystem level’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 391–421, island press, washington dc. gregory, r., ohlson, d. & arvai, j., 2006, ‘deconstructing adaptive management: criteria for applications to environmental management’, ecological applications, 16(6), 2411–2425. doi:10.1890/1051-0761(2006)016[2411:damcfa]2.0.co;2 hall, j.a. & fleishman, e., 2009, ‘demonstration as a means to translate conservation science into practice’, conservation biology 24, 120–127. doi:10.1111/j.1523-1739.2009.01297.x he, f. & gaston, k.j., 2000, ’occupancy-abundance relationships and sampling scales’, ecography 23, 503–511. doi:10.1034/j.1600-0587.2000.230412.x hobbs, r.j. & huenneke, l.f., 1992, ‘disturbance, diversity, and invasion: implications for conservation’, conservation biology 6(3), 324–337. doi:10.1046/j.1523-1739.1992.06030324.x jacobson, s.k., morris, j.k., sanders, j.s., wiley, e.n., brooks, m., bennetts, r.e., et al., 2006, ‘understanding barriers to implementation of an adaptive land management program’, conservation biology 20, 1516–1527. doi:10.1111/j.1523-1739.2006.00476.x kareiva, p., watts, s., mcdonald, r. & boucher, t., 2007, ‘domesticated nature: shaping landscapes and ecosystems for human welfare’, science 316, 1866–1869. doi:10.1126/science.1140170 keller, r.p. & lodge, d.m., 2010, ‘prevention: designing and implementing national policy and management programs to reduce the risks from invasive species’, in c. perrings, h.a. mooney & m. williamson (eds.), bioinvasions and globalization, ecology, economics, management and policy, pp. 220–234, oxford university press, maris, v. & béchet, a., 2010, ‘from adaptive management to adjustive management: a pragmatic account of biodiversity values’, conservation biology 24(4), 966–973. mcgeoch, m.a., dopolo, m., novellie, p., hendriks, h., freitag–ronaldson, s., ferreira, s., et al., 2011, ‘a strategic framework for biodiversity monitoring in sanparks’, koedoe 53(2), art. #991, 10 pages. doi:10.4102/koedoe.v53i2.991 mea. see millenium ecosystem assessment. millennium ecosystem assessment, 2005, ecosystems and human well-being: current state and trends, island press, washington dc. pickett, s.t.a. & thompson, j.n., 1978, ‘patch dynamics and the design of nature reserves’, biological conservation 13, 27–37. doi:10.1016/0006-3207(78)90016-2 pickett, s.t.a., cadenasso, m.l. & benning, t.l., 2003, ‘biotic and abiotic variability as key determinants of savanna heterogeneity at multiple spatiotemporal scales’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity’, pp. 22–40, island press, washington dc. pickett, s.t.a., kolasa, j. & jones, c.g., 2007, ecological understanding: the nature of theory and the theory of nature, 2nd edn., academic press, san diego. pyšek, p. & richardson, d.m., 2010, ‘invasive species, environmental change and management, and health’, annual review of environment and resources 35, 1–31. pyšek, p., richardson, d.m., rejmánek, m., webster, g.l., williamson, m. & kirschner, j., 2004, ‘alien plants in checklists and floras: towards better communication between taxonomists and ecologists’, taxon 53(1), 131–143. rejmánek, m. & pitcairn, m.j., 2002, ‘when is eradication of exotic pest plants a realistic goal?’, in c.r. veitch & m.n. clout (eds.), turning the tide: the eradication of invasive species, pp. 249–253, iucn, gland. richardson, d.m., bond, w.j., dean, w.r.j., higgins, s.i., midgley, g.f., milton, s.j., et al., 2000, ‘invasive alien organisms and global change: a south african perspective’ in h.a. mooney & h.a. hobbs (eds.), invasive species in a changing world, pp. 303–349, island press, washington dc. richardson, d.m., rouget, m. & rejmánek, m., 2004, ‘using natural experiments in the study of alien tree invasions: opportunities and limitations’, in m.s. gordon & s.m. bartol (eds.), experimental approaches to conservation biology, pp. 180–201, university of california press, berkeley. rogers, k.h., 2003, ‘adopting a heterogeneity paradigm: implications for management of protected savannas’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 41–58, island press, washington dc. roux, d.j. & foxcroft, l.c., 2011, ‘the development and application of strategic adaptive management within south african national parks’, koedoe 53(2), art. #1049, 5 pages. doi:10.4102/koedoe.v53i2.1049 salafsky, n., margoluis, r. & redford, k.h., 2001, adaptive management: a tool for conservation practitioners, biodiversity support programme, washington dc. salafsky, n., margoluis, r., redford, k.h. & robinson, j.g., 2002, ‘improving the practice of conservation: a conceptual framework and research agenda for conservation science’, conservation biology 16(6), 1469–1479. doi:10.1046/j.1523-1739.2002.01232.x scholes, r.j. & kruger, j.m., 2011, ‘triggers for management intervention in uncertain, varying and time-lagged ecosystems’, koedoe 53(2), art. #987, 8 pages. doi:10.4102/koedoe.v53i2.987 south african national parks, 2006, ‘mapungubwe national park: park management plan’, unpublished management plan, south african national parks, pretoria. stafford-smith, d.m., abel, n., walker, b.h. & chapin iii, f.s., 2009, ‘drylands: coping with uncertainty, thresholds and changes in state’, in f.s. chapin iii, g.p. kofinas & d. folke (eds.), principles of ecosystem stewardship: resilience-based natural resource management in a changing world, pp. 171–195, springer, new york. stem, c., margoluis, r., salafsky, n. & brown, m., 2005, ‘monitoring and evaluation in conservation: a review of trends and approaches’, conservation biology 19, 295–309. doi:10.1111/j.1523-1739.2005.00594.x timko, j.a. & innes, j.l., 2009, ‘evaluating ecological integrity in national parks: case studies from canada and south africa’, biological conservation 142, 676–688. doi:10.1016/j.biocon.2008.11.022 tu, m., 2009, ‘assessing and managing invasive species within protected areas’ in j. ervin (ed.), protected area quick guide series, pp. 1–40, the nature conservancy, arlington, va. van wilgen, b.w., khan, a. & marais, c., 2011, ‘changing perspectives on managing biological invasions: insights from south africa and the working for water programme’, in d.m. richardson (ed.), fifty years of invasion ecology: the legacy of charles elton, pp. 373–393, blackwell publishing, oxford. van wilgen, b.w., le maitre, d.c. & cowling, r.m., 1998, ‘ecosystem services, efficiency, sustainability and equity: south africa’s working for water programme’, trends in ecology and evolution 13, 378. doi:10.1016/s0169-5347(98)01434-7 venter, f.j., naiman, r.j., biggs, h.c. & pienaar, d.j., 2008, ‘the evolution of conservation management philosophy: science, environmental change and social adjustments in kruger national park’, ecosystems 11, 173–192. doi:10.1007/s10021-007-9116-x wittenberg, r. & cock, m.j.w. (eds.), 2001, invasive alien species: a toolkit of best prevention and management practices, cab international, wallingford. woodroffe, r. & ginsburg, j.r., 1998, ‘edge effects and the extinction of species inside protected areas’, science 280, 2126–2128. doi:10.1126/science.280.5372.2126 supplementum (back to top) supplementum figure 1: an expanded overview of the proposed framework to implementing the control of invasive alien species. the framework indicates the key components, with the salient points, and their linkages. the numbers cross link to the discussion. we also differentiate between documents, activities and/or processes, actions, and bodies and/or committees. article information authors: clinton carbutt1 peter s. goodman2 affiliations: 1biodiversity research and assessment, scientific services, ezemvelo kzn wildlife, south africa2private consultant, conservation solutions, south africa correspondence to: clinton carbutt postal address: po box 13053, cascades 3202, south africa dates: received: 03 sept. 2012 accepted: 23 july 2013 published: 15 oct. 2013 how to cite this article: carbutt, c. & goodman, p.s., 2013, ‘how objective are protected area management effectiveness assessments? a case study from the isimangaliso wetland park’, koedoe 55(1), art. #1110, 8 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1110 copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. how objective are protected area management effectiveness assessments? a case study from the isimangaliso wetland park in this original research... open access • abstract • introduction • methods    • the study area    • national and provincial assessments       • national assessment and attendees       • provincial task team, participatory assessment and attendees    • comparing the national and provincial assessments within a framework of best practice    • provincial assessment methodology       • normalising the scores       • pressures and threats assessment       • correlation exercises • results    • provincial assessment       • comparison of normalised scores per management unit       • pressures and threats       • correlation exercises: relationship between management effectiveness, resource inputs and pressures and threats    • national assessment    • parity in results between national and provincial assessments? • discussion    • disparity between national and provincial scores    • the need for robust and standardised operating procedures       • ‘the right players’: the careful selection of a broad range of relevant staff       • ‘let them speak’: fostering the peer review dynamic       • ‘do not rush’: allow for sufficient time and a sufficient level of detail    • the added value of a pressure and threats assessment • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ the assessment of protected area management effectiveness was developed out of a genuine desire to improve the way protected areas are managed and reported on, in relation to a formalised set of conservation objectives. for monitoring and reporting purposes, a number of participatory methods of rapidly assessing management effectiveness were developed. most rapid assessment methods rely on scoring a range of protected area-related activities against an objective set of criteria documented in a formal questionnaire. this study evaluated the results of two applications of the same management effectiveness assessment tool applied to the same protected area, namely the isimangaliso wetland park, south africa. the manner in which the assessments were undertaken differed considerably and, not unexpectedly, so did the results, with the national assessment scoring significantly higher than the provincial assessment. therefore, a further aim was to evaluate the operating conditions applied to each assessment, with a view to determining which assessment was more closely aligned with best practice and hence which score was more credible. the application of the tool differed mainly with respect to the level of spatial detail entered into for the evaluation, the depth and breadth of the management hierarchy that was consulted, the time in which the assessment was undertaken and the degree of peer review applied. disparate scores such as those obtained in the assessments documented here are likely to bring the discipline of management effectiveness assessment into disrepute unless an acceptable and standardised set of operating procedures is developed and adopted. recommendations for such a set of ‘indispensable constants’ were made in this article to ensure that management effectiveness assessments remain robust and reputable, thereby ensuring an honest picture of what is happening on the ground.conservation implications: we proposed that standard operating procedures should be in place when protected area management effectiveness assessments are undertaken, in order for the results to be credible. this involves ensuring that the right people participate and that each participant is allowed sufficient time to peer review each other. introduction top ↑ protected areas are amongst the most efficient and cost-effective ways of conserving biodiversity (balmford, leader-williams & green 1995) and therefore form the cornerstone of most conservation strategies (bertzky et al. 2012; hockings 2003). the global protected area estate has increased significantly over the past decade, covering some 12.7% of the world’s terrestrial and inland water areas and 1.6% of the global ocean area (bertzky et al. 2012). this amounts to a global protected area estate of 23 million km2, making protected areas the world’s largest form of planned land use (bertzky et al. 2012; convention on biological diversity [cbd] 2010; dudley et al. 2010; ervin 2007; leverington, hockings & costa 2008). despite this significant area under protection, the current system of protected areas does not adequately conserve a representative sample of the world’s biodiversity at any level in the biodiversity hierarchy (i.e. landscapes, ecosystems, communities, species and genetics) (cbd 2010; mcneely, harrison & dingwall 1994). furthermore, protected areas are facing increasing pressures and threats such as habitat loss, fragmentation, isolation, illegal exploitation, invasive species, lack of capacity, inappropriate policies and incentives, the inequitable distribution of costs and benefits, breaches in security and global climate change (international union for conservation of nature [iucn] 2004; leverington et al. 2008; leverington et al. 2010). this suite of external and internal pressures is affecting the conservation community’s ability to manage its conservation estate effectively, thereby undermining its overall contribution to biodiversity conservation (cbd 2010). management effectiveness is defined by the iucn world commission on protected areas (wcpa) as the assessment of how well a protected area is being managed – primarily the extent to which it is protecting the values and achieving goals and objectives of the protected area (hockings, stolton & dudley 2000; hockings et al. 2006). such assessments have generally considered four areas: protected area design (both individual sites and systems) (leverington et al. 2008; leverington et al. 2010), appropriateness of management systems and processes (leverington et al. 2008; leverington et al. 2010), delivery of objectives (ervin 2003a; hockings 2003; leverington et al. 2008; leverington et al. 2010) and ecological integrity (ervin 2003b; parrish, braun & unnasch 2003). if applied broadly across an entire organisation, protected area management effectiveness assessments can enable policymakers to refine their conservation strategies, re-allocate budget expenditures and develop strategic, system-wide responses to the most pervasive threats and management weaknesses (ervin 2003b). protected area management effectiveness assessments are therefore not performance assessments of an individual; rather, they reflect how well a conservation authority is managing its entire conservation estate at the protected area level (carbutt & goodman 2010). protected area management effectiveness is now a well-developed branch of protected area monitoring and evaluation; a number of tools have been developed since the mid–1990s to help assess the effectiveness of protected area management (hockings 2003). however, a diverse range of circumstances and needs require different methods of assessment (world wildlife fund [wwf] & world bank 2007). therefore the iucn’s wcpa, under the guidance of a management effectiveness task force, has developed a framework for the development and standard-setting of a range of management effectiveness methodologies within a consistent overall approach (hockings et al. 2000; hockings 2003; hockings et al. 2006). currently, some 54 assessment tools have been developed in line with this framework (leverington et al. 2010); the choice of the appropriate tool depends on the information and management needs of the individual or organisation applying the tool. in this respect, ezemvelo kzn wildlife has applied protected area management effectiveness assessment methodologies for organisational performance assessment since 2002 (carbutt & goodman 2010; goodman 2002, 2003a, 2003b). management effectiveness assessments will always be criticised for being ‘soft science’ as they are rapid and qualitative in nature, essentially devoid of independently collected empirical data. the risk of using qualitative assessments is that the scoring system can be subject to one-sided opinions and perspectives in the absence of peer review, thereby introducing subjectivity and bias. however, such assessments remain the primary means for undertaking an appraisal of management effectiveness to rapidly report on progress over time (wwf & world bank 2007). therefore, the onus lies on the assessor to conduct the assessment under strict and consistent operating conditions to ensure that the technique remains robust, objective and reputable. the application of the tool should therefore comply with best practice (see ervin 2007; hockings 2003; hockings et al. 2000; hockings et al. 2006; leverington & hockings 2004; leverington et al. 2008; leverington et al. 2010; williams 2011; wwf & world bank 2007). the national and provincial assessments of the isimangaliso wetland park (iswp), listed in 1999 as south africa’s first world heritage site (whs) (figure 1), afforded the opportunity to compare data parity under two different circumstances. the two primary aims of this study were to, (1) compare the results (outcomes) of the two assessments in light of the manner in which each assessment was conducted given that essentially the same questionnaire was used and (2) recommend the appropriate standard methodology that should be applied to ensure that the results of these qualitative assessments are as robust and objective as possible, thereby reflecting an accurate and credible representation of management effectiveness on the ground. a secondary aim of this study was to determine whether or not a separate pressure and threats assessment is able to add further value to the monitoring, evaluation and mitigation components of the adaptive management framework. figure 1: location of the isimangaliso wetland park in kwazulu-natal, south africa. methods top ↑ the study area the non-marine component of the iswp comprises 11 management units (mus) accounting for a total area of 230 164 ha (figure 1). each mu is linked to an operational budget. the national and provincial assessments did not include the two marine protected areas of the iswp, namely the maputaland and st lucia marine protected areas, which were assessed by an independent consultant (tunley 2009) on behalf of marine and coastal management as part of a separate management effectiveness assessment of south africa’s marine protected areas. the national assessment focused on the iswp as a whole, whereas the ezemvelo kzn wildlife (‘provincial’) assessment was applied at the mu level. in this regard, lake st lucia was included as the ‘unofficial’ 11th mu, even though it does not carry a designated budget and staff establishment. some mus are dominated by terrestrial habitats, whilst others are dominated by aquatic habitats. furthermore, the umkhuze mu included lower umkhuze, whilst the ‘st lucia conservation’ mu referred to the st lucia ‘municipal’ area to the cape vidal gate and included the lower narrows (figure 1). dukuduku and makasa nature reserve did not form part of this assessment. the mean effectiveness score was weighted by the area of each mu. national and provincial assessments the national mandate stems essentially from the cbd’s ambitious ‘programme of work on protected areas’ (dudley et al. 2005). the development of this programme stemmed from the outcomes of the 5th iucn world parks congress held in september 2003 in durban, south africa, which made protected area management effectiveness one of its seven major themes and one of the key messages delivered to the cbd (iucn 2004). in response to this international obligation, the national department of environmental affairs (dea) initiated a nationwide project to assess the management effectiveness of south africa’s whs and national parks. since its listing as a whs, the iswp has been governed by an over-arching wetland park authority appointed by national government, in addition to the long established de facto on-the-ground conservation management agency, ezemvelo kzn wildlife, acting on behalf of provincial government. as the national assessment only involved the wetland park authority, it was essential to undertake an additional assessment involving the conservation management agency on the ground (‘provincial assessment’). the national assessment championed by the dea opted for the second edition of the management effectiveness tracking tool (mett) developed by the wwf and the world bank in 2007 (see britton 2010); this assessment tool was based on the iucn-wcpa evaluation framework (hockings et al. 2000; hockings et al. 2006). the mett is a rapid, site-level, qualitative assessment tool based on an expert scoring approach (hockings et al. 2006) that assesses (depending on which version is used) all six elements of protected area management identified in the wcpa framework (see hockings et al. 2000; hockings 2003), namely establishing the context of existing values and threats, followed by adequate planning and the allocation of adequate resources (inputs) and, as a result of management actions (processes), eventually produces products (outputs) that result in impacts that can be measured against set objectives (outcomes). ezemvelo kzn wildlife therefore also made use of the mett for its assessments, as did capenature in january 2009 (dea 2009), given that all data would feed into the national assessment programme. furthermore, the provincial assessment made use of aspects of the wwf rapid assessment and prioritisation of protected area management (rappam) tool (ervin 2003c), which effectively quantifies the total pressures and threats faced by each protected area under assessment, given that the mett is too limited to allow a detailed evaluation of protected area outcomes, as well as the assessment of pressures and threats (wwf & world bank 2007). the pressures and threats assessment was a separate component of the overall assessment. national assessment and attendees the national assessment undertaken on 16 april 2010 comprised seven participants (officials of the wetland park authority, all of which are a cohort of senior management) and was completed in 2 h. the management staff of ezemvelo kzn wildlife, the de facto on-the-ground management agency, did not partake in the assessment. the iswp was assessed as a whole and not per mu. the first author sat in as an observer. the national assessment comprised only the mett questionnaire. provincial task team, participatory assessment and attendees the application of the protected area management effectiveness (pame) programme across kwazulu-natal (including the iswp) was guided by the ezemvelo kzn wildlife pame task team. protected area members of staff were sensitised to the programme through a series of workshops prior to the assessments. the ezemvelo kzn wildlife version of the mett was endorsed by the national assessment programme of dea, with some minor improvements made by ezemvelo kzn wildlife being carried into the national mett assessment tool. this involved the addition of the ‘protected area outcomes’ category, which had, up to this point, not been included in the early development of the national mett tool. furthermore, the assessment involved three exercises, (1) a cover sheet that captured details of the protected area such as size (area), number of staff, annual operational budget, primary management objectives, protected area values et cetera, (2) the mett questionnaire and (3) the pressures and threats analysis. the 16 ezemvelo kzn wildlife staff members who actively participated in the assessment on 23 july 2010 included the park manager, ecoadvice manager, park ecologist, conservation managers for each mu, the district conservation officer and law enforcement staff. comparing the national and provincial assessments within a framework of best practice in line with the aims of this study, two applications of the same assessment tool to the same protected area afforded an opportunity to compare results and the operating conditions under which each assessment was conducted. this involved objectively comparing the operating conditions of each assessment with best practice, with a view to, (1) understanding which assessment was better aligned with best practice and therefore more credible and (2) making an explicit, general statement of global application regarding a minimum set of standard operating procedures for assessments of this nature in the future. in the quest for objectivity, our basis or framework for adjudicating ‘best practice’ was based on a number of key publications in the field of management effectiveness. although most often expressed only as ‘recommendations’, these publications provide ‘guidelines’ that inform how each assessment should be undertaken if we are to accept the scores as rigorous and reasonable reflections of protected area management effectiveness. benchmark examples include: making use of all six components of the iucn-wcpa management effectiveness evaluation framework (hockings 2003; hockings et al. 2006; leverington & hockings 2004), adequate consultation and participation (hockings et al. 2006), applying the assessment at an appropriate level, scale and frequency (hockings et al. 2006), the need for a credible, non-management assessor (hockings et al. 2006), spending sufficient time on the assessment to reach a considered judgement (ervin 2007; wwf & world bank 2007), appropriate multiple stakeholder involvement (ervin 2007; hockings et al. 2006; williams 2011) and the inclusion of a pressure and threats assessment (ervin 2003c). provincial assessment methodology normalising the scores not all questions (assessed criteria) were applicable to all mus. hence, it was necessary to score these items as ‘non-applicable’ rather than penalise the mu with a score of zero. this meant that the maximum potential score differed across mus; the final score per mu was then divided by the denominator that had been totalled by subtracting the value of the ‘non-applicable’ items and then multiplying by 100 to achieve a normalised percentage effectiveness score. pressures and threats assessment both pressures and threats (as defined in the online appendix) were quantified for each of the 11 mus in the provincial assessment. this entailed a qualitative assessment of up to a maximum of 22 identified pressures and threats. to quantify each relevant pressure and threat, members of staff had to assign a value ranging from 4 (highest) to 1 (lowest) that best reflected the extent, impact and permanence of each identified pressure and threat. the degree of pressure and threat was determined by calculating extent × impact × permanence (online appendix); the maximum degree of pressure or threat for each identified pressure or threat was therefore 64 (i.e. 4 × 4 × 4), which was rated as being ‘severe’, whilst the minimum degree of pressure or threat was 1, which was rated as being ‘mild’ (ervin 2003c). the total degree of pressure and the total degree of threat for each mu was determined by summing all individual pressure and threat scores respectively. the maximum potential score in this regard was 1408 (i.e. 22 pressures or threats × 64), whilst the minimum score was zero (ervin 2003c). it is important to note that any pressures and threats assessment is perception-based (r. uys pers. comm., 05 november 2009); each participant possesses different views and outlooks so it was therefore important that sufficient oversight and consistent interpretation was provided in this regard. correlation exercises correlation exercises were undertaken which compared quantitative data sets such as budget and staff numbers with management effectiveness to identify significant relationships based on the correlation coefficient (r) approaching a value of one. results top ↑ provincial assessment comparison of normalised scores per management unit the provincial assessment realised a mean weighted score of 58% effectiveness (table 1). most mus scored between 51% – 60%, followed by the 61% – 70% category (figure 2). two mus scored between 41% – 50% (figure 2). no mus scored more than 68% and no mus scored below 48% (figure 2). the lowest scoring mu was manzengwenya (coastal forest reserve), whilst the highest scoring mus were eastern shores and umkhuze (figure 2). the effectiveness scores across the mus were contained within a narrow range (< 20%), with no major outlying scores (figure 2). table 1: summary of scores for the national and provincial assessments of the isimangaliso wetland park. figure 2: ranking of the management effectiveness scores per isimangaliso wetland park management unit (provincial assessment) relative to the national assessment of the isimangaliso wetland park. pressures and threats some 22 pressures and/or threats were identified as activities or events that either are, or may in the future, have a detrimental impact on the ecological integrity of the iswp. the ‘top six’ pressures (most of which are also the most significant threats) are climate change, alien plants, dam building, bush encroachment, poaching and protected area isolation, which collectively accounted for 51% of the pressures and threats experienced in the iswp (figure 3). in most instances, the total threat score surpassed the total pressure score, inferring that protected area management anticipated that all pressures will increase into the future if not mitigated (figure 4). figure 3: the full suite of 22 pressures and threats displayed as a mean value across all 11 management units of the isimangaliso wetland park ranked from highest to lowest mean pressure (paired with mean threat). figure 4: the total pressure and threat scores for the 11 management units of the isimangaliso wetland park ranked from highest to lowest total pressure (paired with total threat). correlation exercises: relationship between management effectiveness, resource inputs and pressures and threats the 11 mus of the iswp accounted for 319 permanent staff (at an average of 32 permanent staff members per mu) and a total operational budget (excluding payroll) of c. r4 million (at an average of c. r400 000 per mu). few correlations were statistically significant (i.e. when r > 0.602; table 2). management effectiveness was positively correlated to total budget and negatively correlated to total pressure, whilst budget and the number of permanent staff were positively correlated (table 2). total pressures and total threats were also positively correlated (table 2). false bay has a large staff complement for a relatively small mu, whereas far larger mus such as lake sibaya, st lucia conservation and eastern shores have either a smaller or similar staff establishment when compared with false bay. false bay is also well resourced for its size, whereas ozabeni (in particular) and western shores are poorly funded compared to their respective surface areas. table 2: correlation matrix showing the degree of correlation between management effectiveness and total budget, staff, protected area size, total pressures, and total threats. national assessment the national assessment realised an effectiveness score of 86% (table 1; figure 2). no pressures and threats assessment or correlation exercises were undertaken. parity in results between national and provincial assessments? the outcome in terms of comparing effectiveness scores for the two assessments of the same protected area assessed by the same tool is a difference of 28% effectiveness (table 1; figure 2). the score differences were significantly different across all categories of protected area management (figure 5). it is clearly evident that the two assessments were undertaken under highly contrasting operating conditions, with the provincial assessment far better aligned with best practice than the national assessment (table 3). table 3: the operating conditions applied to the national and provincial assessments. each aspect is adjudicated against published best practice guidelines and recommendations. figure 5: ranking of scores per protected area management category for the isimangaliso wetland park based on the provincial assessment (mean score of 11 management units) and the national assessment (single score). n discussion top ↑ disparity between national and provincial scores in this study, we have demonstrated that two management effectiveness assessments of the same protected area, using the same assessment tool, have achieved two very different results. in detailing the protocols used in each assessment, we have noticed that the only difference relates to how the assessments were undertaken (‘operating conditions’) (table 3). given that management effectiveness assessments are essentially perception-based and qualitative in nature, and therefore subject to interpretive bias and subjectivity, the most objective way to determine whether each assessment was providing a fully defensible and objective picture of what’s happening on the ground is to compare each assessment methodology with best practice. after consultation with the literature, it is evident that the provincial assessment is better aligned with best practice than the national assessment, which makes the provincial assessment more credible and hence a more realistic picture of management effectiveness. the provincial assessment involved a quorum of key, relevant staff engaging in robust peer review. given the diversity of habitats and challenges in the iswp, the assessment of individual mus (not the iswp as a whole) each linked to an operational budget added even further value towards the aim of achieving a credible score. further exercises that collected budget and staff data, together with a pressures and threats assessment, presented a comprehensive and defensible picture of the state of the iswp, both in terms of its management effectiveness, management challenges and its state of biodiversity integrity. the provincial assessment should become the baseline for future assessments of the iswp.from the breakdown of operating conditions, it is clear that the national assessment did not fulfil most of the criteria recognised by best practice as being the essential ingredients required for undertaking assessments of this nature (table 3). therefore the score should be interpreted with caution in light of skewed and incomplete stakeholder representation, the perfunctory assessment style that did not encourage peer review or allow the expression of different opinions and perspectives, the coarse park-wide scale at which it was applied and the brief period of time in which it was conducted. this, in all likelihood, explains the high score and the large disparity with the provincial score, which is problematic. the score of 86% effectiveness for the iswp derived from the national assessment is the highest score across all protected areas assessed under the national programme (britton 2010). the score is therefore misleading, given that the national report (britton 2010) concluded by saying that ‘all whs are soundly managed’. a further concern is that the high score distorts the mean for whs as a whole; a whs site such as the ukhahlamba drakensberg park, which scored 73% effectiveness and therefore falls below the mean, would probably have been above the mean if the iswp national score was a more realistic reflection of management effectiveness. not only does a biased, non peer-reviewed score ‘cloud’ the national average (both for all protected areas and whs separately), it also sends a false message to the cbd and brings the assessment method into disrepute, which is a concern for both the science and practice of protected area management effectiveness. the need for robust and standardised operating procedures in order to obtain an objective, robust score, it is clear from a comparison of these two assessments applied under contrasting operating conditions that a basic set of standard operating procedures (sops) should be agreed to and used in all assessments. these should be viewed as ‘non-negotiable’ and not merely as ‘guidelines’ or ‘recommendations’, as the latter invokes a sense of being negotiable. with these ‘ground rules’ in place, we believe that the results of any assessment will be more credible and robust. in the subsections below, we propose three essential ingredients to be included as sops. ‘the right players’: the careful selection of a broad range of relevant staff the accuracy of the score is dependent on identifying the right people and making sure they are present on the assessment day. it is essential to communicate the need for such staff involvement in timely fashion prior to the assessment to ensure that all participate. the assessment tool comprises a broad range of assessment criteria, with no single individual best placed to answer all of the questions with 100% certainty. it is therefore essential to encourage the participation of a range of relevant staff members, each of whom brings a level of expertise to the assessment table. the failure to do so will result in a ‘false assessment’ and hence a ‘false assessment score’. ‘let them speak’: fostering the peer review dynamic the review dynamic involving multiple parties is essential to achieving a fair score. allowing participants the time and space to debate each question will help eliminate any bias, false perceptions or prejudice inherent in such assessments. we have noticed that field staff members tend to be so closely involved with day-to-day activities that they lose objectivity, or tend to be too negative and score low. senior management come with a more strategic viewpoint and, in the absence of the day-to-day realities, tend to score too high. hence the need to encourage a range of viewpoints and opinions and facilitate dialogue until a consensus score is reached. ‘do not rush’: allow for sufficient time and a sufficient level of detail the facilitator should not feel under any pressure or obligation to rush to the next question; rather, the facilitator should only move on to the next question when all participants have had sufficient time to comment and then agree on the score. the assessments should be applied at a spatial scale that is meaningful and should provide sufficient time to encourage peer review, as well as to accommodate the required detail in terms of spatial scale. the added value of a pressure and threats assessment the suite of major pressures and threats identified for the iswp is comparable with those identified in carbutt and goodman (2010) for kwazulu-natal province as a whole, with the notable exception of dam building identified in this analysis, most likely because the iswp is primarily a hydrologically driven system and the impacts of dam building will have far-reaching consequences. given the significant negative correlation between management effectiveness and the pressures being experienced by the iswp, special attention should be directed towards mitigating these numerous high pressures, particularly given their negative impact on biodiversity target achievement. the alien plants pressure and threat should continue to receive attention and measures should be put in place to educate people on the topical issue of climate change. conclusion top ↑ management effectiveness assessments should not be seen merely as a ‘paper exercise’ to meet reporting obligations. rather, they should be undertaken objectively and with sober judgement and diligence to ensure that the effectiveness score achieved represents a realistic picture of management practices and processes, in the absence of hard quantitative data. as protected area management effectiveness assessments are, by their very nature, open to abuse and one-sided, narrow points of view, it is therefore critical that strict sops and conditions are imposed during the undertaking of such assessments. this will result in a credible and robust rapid assessment methodology that generates objective and defendable data within acceptable limits and standards. furthermore, for protected areas such as the iswp where the complex institutional arrangements involve a de facto conservation management agency and a separate overarching authority, one workshop with full representation of all actors and stakeholders is the most appropriate way to assess the iswp that, in doing so, also ensures good cooperative governance. we have noticed that a clear, empathic and absolute statement on how to best apply the various assessment tools is lacking, because most publications address best practice methodology only in terms of ‘guidelines’ or ‘recommendations’ (see hockings et al. 2000; hockings et al. 2006; leverington et al. 2008; leverington et al. 2010; williams 2011; wwf & world bank 2007). such guidelines should be elevated in practice to non-negotiable sops that provide rigour and credibility to the practice of management effectiveness, including but not limited to, (1) full stakeholder representation of all relevant and knowledgeable actors, (2) an environment that caters for the expression of different perceptions and interpretations, moderated by peer review under the direction of an able chairperson who is not involved in protected area management and (3) the application of the tool at a spatial resolution that is biologically meaningful and with sufficient allocation of time to do so. anything less will bring the application of management effectiveness assessment methodologies into disrepute and open them up to abuse. finally, an additional pressure and threats assessment will add further value to the monitoring, evaluation and mitigation components of the adaptive management framework. acknowledgements top ↑ the ezemvelo kzn wildlife pame working group is thanked for guiding the implementation of the programme in kwazulu-natal. heidi snyman, ezemvelo kzn wildlife cartographer, is thanked for producing the map of the iswp. we also thank the two anonymous reviewers for helpful comments and suggestions that have significantly improved the quality of the publication. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions c.c. (ezemvelo kzn wildlife), author of article and co-ordinator of management effectiveness assessments, collated and analysed all results. p.s.g. (conservation solutions) provided guidance and oversight throughout the assessments and provided the title and abstract. references top ↑ balmford, a., leader-williams, n. & green, j.b., 1995, ‘parks or arks: where to conserve large threatened mammals’, biodiversity and conservation 4, 595–607. http://dx.doi.org/10.1007/bf00222516 bertzky, b., corrigan, c., kemsey, j., kenney, s., ravilious, c., besançon, c. et al., 2012, protected planet report 2012: tracking progress towards global targets for protected areas, iucn and unep-wcmc, cambridge. britton, p., 2010, ‘a report on the application of the mett-sa version 1 (2008) to terrestrial protected areas managed at national and provincial level in south africa’, unpublished report for the department of environmental affairs, pretoria. carbutt, c. & goodman, p.s., 2010, ‘assessing the management effectiveness of state-owned, land-based protected areas in kwazulu-natal’, ezemvelo kzn wildlife unpublished report, pietermaritzburg. convention on biological diversity, 2010, programme of work on protected areas, viewed 13 january 2010, from http://www.cbd.int/protected/pow/ department of environmental affairs, 2009, workshop report on the ‘expert input on protected areas study’, department of environmental affairs, pretoria. dudley, n., mulongoy, k.j., cohen, s., stolton, s., barber, c.v. & gidda, s.b., 2005, towards effective protected area systems: an action guide to implement the convention on biological diversity programme of work on protected areas, technical series 18, pp. 1–95, cbd, montreal. dudley, n., parrish, j.d., redford, k.h. & stolton, s., 2010, ‘the revised iucn protected area management categories: the debate and ways forward’, oryx 44(4), 485–490. http://dx.doi.org/10.1017/s0030605310000566 ervin, j., 2003a, ‘protected area assessments in perspective’, bioscience 53(9), 819–822. http://dx.doi.org/10.1641/0006-3568(2003)053[0819:paaip]2.0.co;2 ervin, j., 2003b, ‘rapid assessment of protected area management effectiveness in four countries’, bioscience 53(9), 833–841. http://dx.doi.org/10.1641/0006-3568(2003)053%5b0833:raopam%5d2.0.co;2 ervin, j., 2003c, wwf: rapid assessment and prioritization of protected area management (rappam) methodology, wwf forests for life programme, gland. ervin, j., 2007, assessing protected area management effectiveness: a quick guide for protected area practitioners, pp. 1–17, the nature conservancy and wwf, arlington. goodman, p.s., 2002, a rapid assessment of terrestrial protected area management effectiveness in kwazulu-natal, south africa, pp. 1–30, ezemvelo kzn wildlife and wwf, pietermaritzburg. pmid:12555798 goodman, p.s., 2003a, south africa: management effectiveness assessment of protected areas in kwazulu-natal using wwf’s rappam methodology, pp. 1–29, wwf international, gland. goodman, p.s., 2003b, ‘assessing management effectiveness and setting priorities in protected areas in kwazulu-natal’, bioscience 53(9), 843–850. http://dx.doi.org/10.1641/0006-3568(2003)053%5b0843:ameasp%5d2.0.co;2 hockings, m., 2003, ‘systems for assessing the effectiveness of management in protected areas’, bioscience 53(9), 823–832. http://dx.doi.org/10.1641/0006-3568(2003)053%5b0823:sfateo%5d2.0.co;2 hockings, m., stolton, s. & dudley, n., 2000, evaluating effectiveness: a framework for assessing the management of protected areas, wcpa best practice protected area guideline series 6, iucn, gland. hockings, m., stolton, s., leverington, f., dudley, n. & courrau, j., 2006, evaluating effectiveness: a framework for assessing management effectiveness of protected areas, wcpa best practice protected area guidelines series 14, 2nd edn., iucn, gland. international union for conservation of nature, 2004, iucn 5th world parks congress, durban, south africa, 08–17 september, viewed 13 january 2010, from a summary within http://cmsdata.iucn.org/downloads/iucn_wpc_2014_discussion_paper_13_5_11.pdf leverington, f. & hockings, m., 2004, ‘evaluating the effectiveness of protected area management: the challenge of change’, in c.v. barber, k.r. miller & m. boness (eds.), securing protected areas in the face of global change: issues and strategies, pp. 169–224, iucn, gland. leverington, f., hockings, m. & costa, k.l., 2008, management effectiveness evaluation in protected areas – a global study. report for the project ‘global study into management effectiveness evaluation of protected areas’, p. 170, the university of queensland, gatton. leverington, f., costa, k.l., courrau, j., pavese, h., nolte, c., marr, m. et al., 2010, management effectiveness evaluation in protected areas – a global study, 2nd edn., pp. 1–101, the university of queensland, brisbane. mcneely, j.a., harrison, j. & dingwall, p., 1994, protecting nature: regional reviews of protected areas, iucn, gland. parrish, j.d., braun, d.p. & unnasch, r.s., 2003, ‘are we conserving what we say we are? measuring ecological integrity within protected areas’, bioscience 53(9), 851–860. http://dx.doi.org/10.1641/0006-3568(2003)053[0851:awcwws]2.0.co;2 tunley, k., 2009, state of management of south africa’s marine protected areas, report series – 2009/marine/001, wwf south africa, cape town. williams, b.k., 2011, ‘adaptive management of natural resources – framework and issues’, journal of environmental management 92, 1346–1353. http://dx.doi.org/10.1016/j.jenvman.2010.10.041, pmid:21075505 world wildlife fund & world bank, 2007, management effectiveness tracking tool: reporting progress at protected area sites, 2nd edn., wwf international, gland. article information authors: louise k. swemmer1 sandra taljaard2 affiliations: 1savanna and arid research unit, south african national parks, phalaborwa, south africa 2people and conservation department, south african national parks, northern cluster, golden gate national park, south africa correspondence to: louise swemmer email: louises@sanparks.org postal address: private bag x402, skukuza, kruger national park, south africa dates: received: 04 aug. 2010 accepted: 13 mar. 2011 published: 13 may 2011 how to cite this article: swemmer, l.k. & taljaard, s., 2011, ‘sanparks, people and adaptive management: understanding a diverse field of practice during changing times’, koedoe 53(2), art. #1017, 7 pages. doi:10.4102/koedoe.v53i2.1017 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) sanparks, people and adaptive management: understanding a diverse field of practice during changing times in this essay... open access • abstract • introduction • the historical evolution of the people and conservation department    • the early days    • the emergence of the social ecology unit    • transformation into ‘people and conservation’    • the emergence and role of social science research • strategic adaptive management – objectives driven management and research    • strategic adaptive management and the people components of parks    • strategic adaptive management and natural resource use    • strategic adaptive management and social and economic monitoring • conclusion • acknowledgements • references abstract (back to top) biodiversity conservation is often measurable and achievable and has been reasonably successful within the boundaries of national parks. however, the concept of parks providing tangible benefits and hence being seen as ‘valuable’ to the majority of the nation has been more difficult to define, measure and, importantly, deliver on. this function has traditionally fallen under what is currently known as the people and conservation department, which has a rich history in south african national parks (sanparks) of change and adaptive learning in terms of defining core functions and associated management strategies, spanning from its original inception as the information services department over 80 years ago. learning from and in some cases, adapting to change, is evident throughout this broad scale national evolution of the department, from an initial focus on information sharing and education in the 1930s, to what we see today. this includes the primary focus areas of cultural resource management and indigenous knowledge, community relations, environmental education, awareness, youth outreach, interpretation and training. at a more local, park scale, there is a current drive to formalise the adaptive management and learning process for the people component of protected areas through the alignment of relevant project, programme and park objectives with those at a corporate or national level. associated with this is an attempt to further align the associated monitoring, evaluation and reporting processes, thereby completing the formal adaptive management loops in order to facilitate and stimulate co-learning within and between relevant responsible departments within the organisation. conservation implications: benefit sharing through biodiversity conservation has been shown to be crucial for the long-term success of protected areas, but the practicalities of implementing this are thwart with challenges. despite this, sanparks is attempting to facilitate and promote benefits through conservation, specifically in the sense of benefits that support livelihoods whilst reducing vulnerability. with this in mind, we acknowledge the importance of the concepts of scale, resilience, complexity and adaptive learning for, and during, this process. introduction (back to top) the primary focus of south african national parks (sanparks) is the conservation of protected areas and their associated biodiversity, for the pride and benefit of the nation (see sanparks 2006). sanparks has a strong history of successful biodiversity conservation, achieved through the identification of well structured management objectives and the implementation of intensive monitoring programmes (sanparks 1997). more recently, the formal implementation of the adaptive management approach by park management and scientists, involving regular monitoring and feedback loops that inform the re-evaluation of management objectives and techniques for biodiversity conservation, has further enhanced the management effectiveness for the biodiversity component of parks at various levels (roux & foxcroft 2011). however, an important component of sanparks’ function is to ensure that the potential benefits of biodiversity conservation are accessible to the broader society. this ensures the delivery of the sanparks mandate, as well as constituency building for conservation. in reality, this ‘people component’ has received somewhat less attention and support in the past, having been previously criticised by some as not making a significant contribution towards the sanparks mission (pollard, shackleton & carruthers 2003). historically, the various stakeholder groups that derived benefits from protected areas were determined primarily by the management era at the time, and this has played a large role in influencing the ways in which the people components of sanparks have been managed over the years. the formal adoption of the strategic adaptive management (sam) (biggs & rogers 2003) approach in sanparks in the early 2000s did not filter through to the people components of sanparks management until very recently and, as a result, there are currently few examples of formal adaptive management being practiced in this field within the organisation. despite the lack of formal strategic adaptive management, the dynamic process illustrated in the longer-term development of the people component of sanparks, then known as the national parks board (npb) or nasionale parkeraad (npr) in afrikaans, over the past 84 years does demonstrate a certain number of lessons learnt to date, some of which have contributed towards a degree of informal adaptation to change. in some cases this adaptation was conscious and in others not, but ultimately it was about attempting to adapt and, probably more accurately, to survive the challenging circumstances and requirements of an ever-changing environment. these adaptations were driven by changes in the national and global approach to conservation, as well as by national and international politics and changes in legislation (pollard et al. 2003; taljaard 2008). this paper aims to outline key events and milestones during the evolution of the people-centred approach to conservation over the last 80 years. we further aim to highlight both the positive and negative influences contributing to the adoption of formal adaptive planning and management tools. the historical evolution of the people and conservation department (back to top) the early days although the people and conservation department (p&c) currently covers a wide range of functions, including cultural resource and indigenous knowledge management, community relationship building, local economic empowerment, environmental education, outreach and awareness, interpretation and training, as well as youth outreach, it began with simpler informational and educational functions from as early as 1952, when the first formal information officer was appointed (taljaard 2008). this official post was initiated as a response to a number of school groups and scientific expeditions that took place in the kruger national park (knp) between 1932 and 1950 (milne 1996; taljaard 2008). as more national parks were established between the 1950s and 1980s, the need for people-interaction on an interpretational and educational level grew rapidly, resulting in the need to employ more staff in the information services department. up to this period, the main focus on the interpretational and educational front was on white south africans, which was in line with the political order of the day. however, in 1981, an explicit, formally documented intent to serve the broader south african public was published (npr 1981a; npr 1981b). this was the first of its kind and marked a watershed of change within the approach of inclusively educating all people on environmental aspects and related issues. this move was clearly linked to the political changes and instability that characterised south africa in the 1980s. in 1987, the first mention was made of a partnership between conservation areas and their neighbours, stressing the importance of ‘conservation education’ (taljaard 2008). in 1989, a report appeared titled ‘ multicultural environmental education’ (npr 1989). this report focused even more directly on the inclusion of neighbouring communities in educational programmes and also noted that the knp accommodated the first ‘non-white’ (standard jargon of that era, now considered an exclusionary term) groups in its bush camps during that year. there were other ‘multicultural’ programmes forthcoming from these efforts and it was reported that these interventions were valuable and successful attempts to encourage the development of interracial relationships between the sanparks and local communities (npr 1989). one example of this was the five non-white schools that attended the national environmental youth symposium (nyes) for the first time in 1987. the nyes was initiated by sanparks in 1981 and hosted annually in the golden gate national park. the main aim of the nyes was to promote environmental awareness and appreciation by the youth and, by 1988, the number of non-white participating schools had grown to sixteen (npr 1989). the emergence of the social ecology unit changes in a new democratic dispensation that picked up momentum in the early 1990s marked the first democratic elections in 1994, and this led to the establishment of the social ecology unit (se) in that same year. the main aim of the se was to improve strained relationships with neighbouring communities, avert threats such as poaching and land grabs and respond to the general trends towards democracy in southern africa (sanparks 2001). the mid-nineties also marked the start of the integrated conservation and development programmes (icdps) in south africa, aimed at integrating environmental protection with poverty reduction (campbell, sayer & walker 2010). land restitution became an important driver of some of the sanparks’ decision-making around land use, the outcomes of which have been mixed and multidimensional thus far (carruthers 2007). during this time, positive discrimination policies in favour of employing black south africans were implemented (fabricius 2004) and the development of the concept of transfrontier conservation areas (tfcas) further supported the ideology of promoting benefits through the broader socioecological systems approach (whande & suich 2009). during this time, the se strived to influence the npb policies to accelerate the shift from the practice of traditional conservation to a more holistic, integrated, natural and cultural heritage management approach (sanparks 2007). the se established and serviced various community structures (such as park forums), with the main aim of facilitating positive relationships with neighbouring communities through transparent communication strategies. the situation that resulted from these changes could almost be described as an ‘over-emphasis’ or ‘over-reaction’ to the ‘social’ aspect of sanparks’ mandate during the intense political turmoil associated with that time and this resulted in a programme that was under an enormous amount of pressure to serve as a focal point of transformation within the organisation. this resulted in unrealistic and often unfair expectations being placed on the se, with many people being under the impression that the unit would be the arm of sanparks that would take on the role of rural development and, hence, address the needs related to regional poverty. in some ways, it was hoped that the establishment of the se would alleviate the pressure felt by the other departments to contribute towards development outside of the park boundaries. as such, the se was almost seen as being solely responsible for alleviating the negative tensions and pressure on resources that were becoming a threat to the survival of national parks in south africa. despite this pressure on the se at a national and parks level, there was very little internal support for what the department was trying to achieve. this was fuelled by an internal resistance to change within sanparks, as well as a broader lack of recognition of the tangible role that the se could and should play in the achievement of the broader conservation mission. furthermore, owing to the historical legacy of biodiversity management dominating the conservation agenda, there was a lack of social-specific scientific support for management of se functions from the research division, which further contributed towards a sense that the se was alienated from the rest of the organisation. this often put the se, as well as the concept of ‘social ecology’ within sanparks in the hot seat. however, it still managed to create a platform for reflection on the social ecology philosophy, which was aimed at achieving biodiversity conservation goals without separating humans and nature (sanparks 2001). creating a balance between meeting human needs and conserving nature was becoming more and more challenging both locally and globally, as a result of human population growth and associated poverty increasing the dependency and pressure on natural resources, all of which made the se mandate exceedingly difficult to manage and achieve. furthermore, the park forums often became centres of conflict on issues of power and access to resources (sanparks 2007). social ecologists were operating on relatively unknown terrain that overlapped with areas that were supposed to be serviced by other government departments. this meant that, to a large extent, the department as a whole was operating independently from the rest of the organisation. as a result of the mounting pressure and emerging challenges, sanparks management marginalised the se towards the late 1990s, arguing that its practitioners were working outside of the scope of sanparks’ core functions (sanparks 2007). transformation into ‘people and conservation’ the se faced much difficulty between the late 1990s and early 2000s, when its value and contribution towards the organisation was in question. however, there was an emphasis on the concepts of ‘people and parks’ and ‘benefits beyond boundaries’ at the world summit on sustainable development (wssd 2000) and the world parks congress (wpc 2003). this message was formalised in the protected areas act of 2003 (act no. 57 of 2003) (department of environmental affairs and tourism 2003), which further highlighted the role sanparks had to play with regard to issues of sustainable economic development. this resulted in the transformation of the se into a new directorate known as the people and conservation department (p&c) in august 2003 (sanparks 2007), which enabled the maturation and growth of the ‘people and parks’ section of sanparks’ mandate to continue unabated. many lessons were learnt from the se era, some of which have, over time, been converted into recommended courses of action and there has been a move towards defining clearer roles and responsibilities within the organisation (sanparks 2001). the intention of establishing the p&c was to support the core business of sanparks in building constituencies for biodiversity conservation through the p&c programme at local, national and international levels. between 2003 and 2008, the main focus areas of the p&c included community relations, environmental education, awareness, interpretation and training, cultural resource management and indigenous knowledge, youth outreach and social science research (sanparks 2007). the emergence and role of social science research the emergence of the p&c also marked the first appearance of a formal post to manage and coordinate social science research within sanparks in 2004. the initial objective of including this area of focus as one of the key functions of the p&c was to create more awareness of it, to identify and encourage more social science research projects and to create a more scientific approach towards the people and parks fraternity within sanparks. however, it became clear that although many more social research projects were being registered formally through the process, and social science research had begun to make a name for itself within the organisation, there was a need to merge social and ecological research in order to address the broader socioecological system through research that was more transdisciplinary in nature. hence the social science research function was transferred to the conservation services department’s scientific services unit in 2008. in that same year, the p&c was merged with the conservation services department in order to further facilitate the collaborative approaches of managing conservation with people. the transfer of social science from the p&c to the conservation services department enabled further growth in this field of research through facilitating the move from what was previously primarily academic driven research at a national level, to research that was driven from a park based management needs perspective. strategic adaptive management – objectives driven management and research (back to top) sanparks has implemented a unique version of adaptive ecosystem management known as sam (biggs & rogers 2003). this was augmented by the acknowledgement that the socioecological systems in which we live and work are complex; in other words, they are unpredictable and constantly changing. management of these systems therefore requires an approach that can be adapted continually where necessary as our understanding and knowledge base grows. the focus of sam is on using a strategy that enables looking forward to promote pro-active as opposed to reactive thinking and decision-making (biggs & rogers 2003). this approach requires clear objectives at both a national and a parks level (in park management plans) in compliance with the protected areas act 2003 (act no. 57 of 2003) (deat 2003). these objectives provide a framework and context for the management programmes; however, there is likely to be poor understanding, learning and agency adaptation if we cannot evaluate the extent of achievement of these objectives and respond to these evaluations effectively. this requires a basic understanding of the system in order to identify the indicators that can be measured, the thresholds that would flag significant changes in these indicators and the drivers that could cause changes in these indicators (water research commission 2008). it also requires the creation of a reporting and feedback process that will inform the learning process through evaluating (amongst others) the effectiveness of management actions and the appropriateness of the objectives, indicators and thresholds (wrc 2008). some of these objectives (particularly those associated with the biodiversity components) have clearly defined indicators and thresholds that enable the evaluation of the degree to which the objectives are being addressed or achieved in each case. clear examples of these objectives and measurables can be found in the river management arena (wrc 2008). strategic adaptive management and the people components of parks although the above process has been well articulated within the biophysical realm of sanparks, numerous challenges exist when it comes to its effective implementation. similarly, implementing sam for the social components of protected areas comes with its own set of challenges and, if it is to be applied effectively, requires an understanding of some specific concepts. we need to acknowledge that protected areas are nested in a mosaic of land uses and it is necessary to appreciate the role that they play in the broader system if we are to understand which drivers support and which drivers threaten their existence or constrain their goals. having a complete understanding of these factors is nearly impossible owing to their complex, dynamic and unpredictable nature. our management strategies and approach to thinking around these issues need to accommodate this incomplete knowledge by being flexible and adaptable to change as our knowledge and understanding of the system grows. protected areas are nested in a system that supports people through a large variety of ecosystem services or benefits that people derive from nature (scholes & biggs 2004). the accessibility of these services to people living within the system has a direct bearing on human livelihoods and well-being (scholes & biggs 2004), with a reduced available supply of services resulting in increased household vulnerability to risks and shock events (parent 2010; scholes & biggs 2004). although sanparks strives to reduce household vulnerability through promoting access to ecosystem services that will have positive effects on livelihoods and well-being, this has only been articulated in a very broad way and not embedded explicitly in much of what the se and, more recently, the p&c has been trying to accomplish. the need for a strategic plan and conceptual framework for benefit sharing and constituency building has long been identified as a priority (pollard 2003) and these specificobjectives have been articulated systematically only recently in the form of a ‘people objectives hierarchy’ across sanparks (figure 1) (sanparks 2010a). figure 1: a diagrammatic representation of the draft high level management objectives for the people component of the south african national parks (sanparks) mandate. broadly speaking, the people objectives of sanparks focus on both beneficiation from parks (through ecotourism, ecosystem services and economic opportunities), cultural heritage and constituency building for conservation (through education, awareness, outreach and building community relationships). gaining a complete understanding of the socioecological system is impossible, but a degree of understanding is necessary in order to identify the main drivers that influence the outcomes of these various components. further understanding is needed to identify appropriate indicators and threshold levels for measuring and monitoring the drivers and this has been a large challenge limiting the implementation of an effectively adaptive, formal learning based management approach for the people components of park management in the past. considerable applied research is still needed to support informed decision making in this regard. strategic adaptive management and natural resource use natural resource use is categorised as a component of ecosystem services under the sanparks beneficiation objectives and sanparks has an extractive natural resource use programme under which a number of examples of formal implementation of the adaptive management approach with regards to resource use can be found (scheepers, swemmer & vermeulen 2011). however, even in these examples, the focus remains on monitoring and evaluating the ecological impacts of resource use, and few go into much detail regarding the social or economic components. ultimately, the primary objective of the sanparks’ extractive resource use programme is the promotion of sustainable beneficiation to predominantly poor neighbouring communities through the improvement of livelihoods and well-being (sanparks 2010b). other objectives for embarking on extractive resource use in sanparks include the effective management of ecosystems (e.g. game sales) and compliance with legislation (sanparks 2008). in this context, ecological as well as social and economic sustainability are important. a holistic, adaptive management framework for the sustainable implementation of resource use initiatives should therefore include a detailed ecologically sustainable harvesting strategy (which includes an ecological monitoring programme), a sound business model that articulates financial and economic components, a social impact assessment and social monitoring strategy and, finally, a cost benefit analysis that would enable the combination of all of these components. one of the requirements for the setting of sustainable harvesting quotas is knowledge of the production rate of the biological material in order to ensure that the off-take rate does not exceed the production rate. however, there is a lack of baseline data when it comes to production and this is partly a result of the dynamic nature of ecosystems. overcoming this requires a strategy that will enable the setting of flexible harvesting quotas, which are informed by a monitoring process that incorporates both temporal and spatial fluxes in production. collecting sufficient data on this before such a strategy can be designed takes much time and energy and the reality is that we may never have enough data that spans the degrees of change in the system, enabling us to make absolute decisions with regards to sustainable harvesting quotas. decisions need to be made in feasible and socially acceptable timeframes; hence, we have opted for the learning-by-doing approach with regards to resource use. as such, we are dependent on detailed, yet flexible monitoring programmes and regular feedback loops to inform and guide the learning process towards the implementation of more effective and adaptable processes that take the precautionary principle into account and are ecologically sustainable, yet flexible through their acknowledgement of the existence and role of ecosystem flux (sanparks 2008). implementing and monitoring the financial, economic and social components of sustainability with regards to resource use poses its own challenges that have not received much attention in sanparks to date. often, resource use harvesting in national parks is not financially sustainable for either the resource harvesters or the park. in these cases the economic (monetary and other nonmonetary livelihoods strategies) as opposed to the financial (strictly monetary) benefits of granting people access to protected areas need to be included in the cost-benefit analysis. an example of this could be the value of the constituency gained through enabling access to parks, which may outweigh the small financial cost of a certain initiative to a park. strategic adaptive management and social and economic monitoring social and economic monitoring is difficult, especially because it may not be easy or possible to valuate or quantify many of the nonmaterial impacts, costs and benefits. furthermore, traditional reporting templates are not able to accommodate for reporting of a qualitative nature, which is what many of the social objectives require. this often results in intangible impacts (both positive and negative) being ignored in cost-benefit analyses. additionally, benefits operate at various levels, with those that operate closer to the source (primary benefits) being clearer and easier to identify, and more quantifiable, than others which operate as ‘knock-on’ or added value benefits (secondary or tertiary benefits). monitoring the social impact is important in order to be aware of and to monitor any social benefits, as well as to mitigate any negative consequences of outreach or beneficiation initiatives. social impact assessment (sia) is a tool used to assess, or estimate in advance, the social consequences that are likely to follow from specific outreach initiatives or projects (lahiri-dutt, nair & dowling 2008). in other words, sia allows for the identification of the likely and realised impacts of a project on people. examples of social impacts could include changes that occur in people’s way of life (how they live, work, play and interact with one another on a daily basis), their culture (shared beliefs, customs and values) and their community (its cohesion, stability, character, services and facilities) (lahiri-dutt et al. 2008). the primary objectives of these in the case of sanparks, would be to ensure that local communities are not adversely affected by initiatives and to facilitate their ability to reap sustainable benefits from development activities (lahiri-dutt et al. 2008). sia can be used in the planning stages of projects, as well as in ongoing monitoring and evaluation processes (lahiri-dutt et al. 2008). the outcome of well planned monitoring programmes for social projects and programmes should include adequate and relevant data at the correct scope and scale that can be evaluated and analysed using both qualitative and quantitative methods in order to determine whether set objectives are being met and where adjustments in management are necessary or appropriate. this highlights the need for clear objectives at a project, programme, park and organisational level. these objectives need to be aligned in a way that park-based implementation is guided ultimately by its respective contribution towards the achievement of the sanparks desired state or mission statement (roux & foxcroft 2011). this, in turn, calls for an aligned and hierarchical monitoring and reporting process from a project level to a national level that facilitates careful evaluation and analysis of data. the current national corporate strategy is articulated in the ‘corporate strategic balanced score card’ (sanparks 2010c), the measurables of which do not necessarily accurately reflect the objectives derived via the articulation of the desired state. one example of this is the corporate strategic objective relating to the growing of constituencies and the provision of access to benefits from the national parks system. the three measurables identified here include, (1) the number of participants in environmental education programmes, (2) the number of internal awareness interventions and (3) the number of sustainable resource use projects. where the objectives are to provide benefits and build constituencies, the true measurables should reflect both a measure of benefits and of what was learnt during the educational programmes; however, the current measurables fail to do this. this does not reflect well at a project level and does not guide the reporting and monitoring process at this level. similarly, whilst the corporate objective related to the facilitation of socioeconomic development is measured as a count of the community-based socioeconomic initiatives implemented, the objective is actually about beneficiation and livelihoods, but, again, the measurables do not reflect this. conclusion (back to top) it is clear that events during the 80-year evolution of the current people component of sanparks management has had a profound influence on the degree to which formal strategic adaptive management has been adopted in the social components of parks. however, despite the fact that biodiversity conservation has a much longer history of this type of management, and is considered to be primarily measurable and achievable, the effective implementation of sam in these more ‘tangible fields’ is also fraught with challenges. the concept of parks providing benefits to people other than direct employment opportunities and recreation is even more difficult to define, measure and, importantly, deliver on, but it is widely accepted that benefit sharing through biodiversity conservation is crucial for the long-term success of protected areas. the effective implementation of the people objectives is also full of challenges. apart from the theoretical difficulties in identifying, quantifying and monitoring both tangible and intangible benefits, contradicting values and belief systems between stakeholder groups partially dictate how benefits are viewed and prioritised by different parties. often, the expectations or demands for benefits far outweigh the reasonable possibilities or sustainable opportunities that arise from protected areas, with the protected areas being viewed naively as a solution to national poverty. within sanparks, there is a need to clearly align the objectives for park-based projects and programmes with national corporate programmes. following this, there is a need for the alignment of associated monitoring and evaluation techniques and reporting protocols at these various levels within the organisation. the formal adoption of sam into the social components of sanparks is becoming more evident and, to date, has been implemented most successfully in the natural resource use arena. however, sanparks is currently attempting to use sam more extensively in other aspects of the people objectives in order to facilitate learning whilst attempting to predict drivers of change that could ultimately impact on the effectiveness of promoting benefits through conservation, specifically in the sense of benefits which support livelihoods whilst reducing vulnerability. acknowledgements (back to top) we would like to acknowledge the sanparks people and conservation department for sharing their experiences, the reviewers for their valuable comments and suggestions and dr harry biggs for his expert input and recommendations on improving the text. references (back to top) biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs, the kruger experience. ecology and management of savanna heterogeneity, pp. 59–80, island press, washington dc. campbell, b.m., sayer, j.a. & walker, b., 2010, ‘navigating trade-offs: working for conservation and development outcomes’, ecology and society 15(2), viewed on 01 august 2010, from http://www.ecologyandsociety.org/vol15/iss2/art16/ carruthers, j., 2007, ‘south africa – a world in one country: land restitution in national parks and protected areas’, conservation and society 5(3), 292–306. department of environmental affairs and tourism, 2003, national environmental management: protected areas act 2003, (act no. 57 of 2003), deat, pretoria. fabricius, c., 2004, ‘the fundamentals of community-based natural resource management’, in c. fabricius, e. koch, h. magome & s. turner (eds.), rights, resources and rural development: community-based natural resource management in southern africa, pp. 3–43, earthscan, london. lahiri-dutt, k., nair, a. & dowling, s., 2008, social impact assessment: a manual for mining projects, resource management in asia pacific programme, australian national university, canberra. milne, i.b., 1996, an investigation into the development, principles and practice of environmental interpretation in south africa: a case study of the national parks board. med thesis, dept. of education, rhodes university. national parks board, 1960, 34th annual report of the national parks board of trustees, 1 april 1959 – 31 march 1960, national parks board, pretoria. nasionale parkeraad, 1981a, beplanning ten opsigte van die raad se inligtingsaksie vir die onmiddellike toekoms [planning in relation to the board’s information action for the immediate future], nasionale parkeraad, pretoria. nasionale parkeraad, 1981b, inligtingsaksie van die nasionale parkeraad [information action of the national parks board], nasionale parkeraad, pretoria. nasionale parkeraad, 1989, multi-kulturele omgewingsopvoeding [multicultural environmental education], nasionale parkeraad, pretoria. parent, g., 2010, ‘kruger feedback’, scientific services project meeting, skukuza, kruger national park, april 22, 2010. pollard, s., shackleton, c. & carruthers, j., 2003, ‘beyond the fence – people and the lowveld landscape’, in j.t. du toit, k.h. rogers & h.c. biggs, the kruger experience. ecology and management of savanna heterogeneity, pp. 59–80, island press, washington dc. roux, d. & foxcroft, l., 2011, ‘the development and application of strategic adaptive management within south african national parks’, koedoe 53(2), art. #1049, 5 pages. doi:10.4102/koedoe.v53i2.1049 scheepers, k., swemmer, l.k. & vermeulen, 2011, ‘applying adaptive management in resource use in sanparks’, koedoe 53(2), art. #999, 14 pages. doi:10.4102/koedoe.v53i2.999 scholes, r.j. & biggs, r., 2004, ‘ecosystem services in south africa: a regional assessment’, in r.j. scholes & r. biggs (eds.), the regional-scale component of the southern african millenium ecosystem assessment, council for scientific and industrial research, pretoria. south african national parks, 1997, ‘a revision of parts of the management plan for the kruger national park volume v11 – an objectives hierarchy for the management of the knp, december 1997’, sanparks, pretoria, unpublished. south african national parks, 2001, a social ecology policy for south african national parks, sanparks, pretoria. south african national parks, 2006, ‘coordinated policy framework governing park management plans, draft 3, work in progress document’, july 2006, sanparks, pretoria, unpublished. south african national parks, 2007, ‘south african national parks week brochure 2007’, sanparks, pretoria, unpublished. south african national parks, 2008, sanparks resource use policy, ref. 17/p – csd/pol/resource use/03-10/vs1, sanparks, pretoria. south african national parks, 2010a, ‘a people objectives hierarchy’, poster presented at the 9th ahead meeting, hazyview, 23–25 february. south african national parks, 2010b, ‘sanparks extractive resource use programme outline and objectives hierarchy’, sanparks, pretoria, unpublished report. south african national parks, 2010c, corporate strategic balanced score card. financial year 2010/2011, sanparks, pretoria, unpublished. taljaard, s., 2008, ‘an investigation into the development of environmental education as a field of practice in south african national parks’, med thesis, department of education, rhodes university. water research commission, 2008, ‘application and testing of a strategic adaptive management system for freshwater protection, associated with implementation of south africa’s national water policy’, project report wrc project k5/1797, deliverable no. 3, wrc, pretoria, unpublished. whande, w. & suich, h., 2009, ‘transfrontier conservation initiatives in south africa: observations from the great limpopo transfrontier conservation area’, in h. suich, b. child & a. spenceley, evolution and innovation in wildlife conservation , pp. 373−393, earthscan, london. world parks congress, 2003, durban accord, world parks congress proceedings, durban, south africa, september 8–27, 2003, viewed 01 august 2010, from http://www.iucn.org/about/work/programmes/pa/pa_event/wcpa_wpc/ world summit on sustainable development, 2000, ‘summary of the 24th special session of the general assembly 26 june – 1 july 2000’, earth negotiations bulletin 10(63), viewed 28 july 2010, from http://www.iisd.ca/download/pdf/enb1063e.pdf issn 0075-6458 59 koedoe 49/2 (2006) introduction the egoli granite grassland is a type of grassland and a mapping unit (gm10) classified under the mesic highveld grassland bioregion according to the new vegetation map of south africa, lesotho and swaziland (mucina et al. 2005). this grassland was mapped as bankenveld (veld type 61) by acocks (1988), and rocky highveld grassland (vegetation type 34) by bredenkamp & van rooyen (1996). acocks (1988) describes the bankenveld as a false grassveld type. the climax vegetation of this veld type should be, according to acocks, an open savanna, that is a bushveld vegetation, but it has been changed to, and maintained as grassveld by regular veld fires. however, bredenkamp & brown (2003) consider bankenveld vegetation as a mosaic of grassland and woodland communities controlled by (micro-)climatic conditions that exist in the topographically heterogeneous landscape in the transition zone between the grassland and savanna biomes. from a conservation planning or urban development viewpoint, the range of structural and floristic variation is considered too large to define bankenveld as a single unit, even on the reconnaissance level of vegetation investigation. bredenkamp & brown (2003) recognised 16 major vegetation types within bankenveld. the egoli granite grassland is considered as mainly belonging to the hyparrhenia hirta anthropogenic grassland (bredenkamp & brown 2003). egoli granite grassland is restricted to the gauteng province of south africa and is located north of the roodepoort / krugersdorp ridge complex and stretches over a distance of approximately 35 km to centurion in the north. this grassland stretches over a distance of approximately 50 km in a westerly direction from the r21 highway in the east. the 28º longitude and 26º latitudiconservation value of the egoli granite grassland, an endemic grassland in gauteng, south africa g.j. bredenkamp, l.r. brown and m.f. pfab bredenkamp, g.j., l.r. brown and m.f. pfab. 2006. conservation value of the egoli granite grassland, an endemic grassland in gauteng, south africa. koedoe 49(2): 59–66. pretoria. issn 0075-6458. gauteng is the most densely populated province in south africa. its remaining natural areas are constantly under threat from urban development and the associated impacts. presently, the natural areas of gauteng support a large diversity of ecosystems. one such ecosystem is the egoli granite grassland, endemic to the province, poorly conserved and therefore highly threatened. this paper describes the original egoli granite grassland and the anthropogenic hyparrhenia hirta dominated grassland that has replaced it in many of the remnant areas. human impacts on this sensitive ecosystem have resulted in an altered species composition, loss of many species, and a change from a species-rich grassland with high conservation value to a species-poor grassland with low conservation value. the conservation of the last remaining relicts of original egoli granite grassland is essential. keywords: egoli granite grassland; conservation value; hyparrhenia hirta, gauteng. g.j. bredenkamp, african vegetation and plant diversity research centre, university of pretoria; l.r. brown, applied behavioural ecology and ecosystem research unit, university of south africa; m.f. pfab, bioregional planning, gauteng department of agriculture, conservation and environment, johannesburg, 2001 south africa. bredenkamp.indd 59 2006/10/15 10:51:01 pm koedoe 49/2 (2006) 60 issn 0075-6458 nal lines cross in the centre of this grassland (fig. 1). due to high demand for developable land in gauteng, and especially the johannesburg / pretoria corridor, the egoli granite grassland is under extreme pressure. large areas are already developed for residential, industrial and commercial purposes. this has led to the destruction of vast tracts of this grassland. at least 61 % of egoli granite grassland has been permanently transformed: 27 % by urban development (fig. 2); 17 % by smallholdings; 12 % by agriculture (cultivated lands); and 5 % by other impacts such as exotic plantations, mining and planted pastures. an estimated additional 17 % of egoli granite grassland is degraded due to overgrazing and the influence of edge effects associated with a highly fragmented landscape (fahrig 2003). other effects, such as trampling, increased pollutants and the infestation of weeds and alien species also contribute to degradation of this grassland. it is therefore highly likely that the national target for conservation of this grassland type, i.e., 25 % of the total extent, will never be realised. egoli granite grassland is extremely poorly conserved, with only 0.02 % (26 ha) of the vegetation type currently protected, including 3 ha in glen austin bird sanctuary, 3 ha in melville koppies nature reserve, 9 ha in ruimsig nature reserve, and 11 ha in the walter sisulu botanical gardens (fig. 2). the current protection status of egoli granite grassland is therefore completely inadequate, with only 0.1 % of the national target actually achieved. in order to meet south africa’s international obligations, it is imperative that the gauteng provincial government put in place measures to improve the conservation status of egoli granite grassland. any viable remnant patch of original egoli granite grassland must therefore be protected from transforming land uses. in order to assess whether a valuable remnant patch will be affected by a proposed development, environmental consultants involved in vegetation impact assessments are required by the gauteng department of agriculture, conservation and environment (gdace) to assess the presence and condition of egoli granite grassland on any site proposed for development. as this grassland was only recently defined with the development of the new vegetation map of south africa, lesotho and swaziland (mucina et al. 2005), very little information on its floristic composition, condition, the 60 0 60 120 km jhb pta fig. 1. locality map of the egoli granite grassland. grey areas indicate urbanisation 10 0 10 20 kilometers n fig. 2. dark gray represents remnant areas of potentially original egoli granite grassland; perennial rivers shown. bredenkamp.indd 60 2006/10/15 10:51:02 pm issn 0075-6458 61 koedoe 49/2 (2006) extent of the grassland, and the management thereof exists, causing some confusion among nature conservationists, environmental consultants and local authorities. the aim of this paper is therefore to provide an ecological overview of this grassland. physical environment this area is generally characterised by rocky undulating plains representing crests, slopes and valley bottoms with shallow, nutrientpoor soils. shallow drainage lines and vleilike wetland areas occur in the valley bottoms. rockiness of the soil surface is a further common characteristic shared by most bankenveld areas. mean monthly temperature in the study area is 16.8 ºc with a mean maximum of 22.6 ºc and a mean minimum of 10.8 ºc. the mean winter temperature is 13.8 ºc and mean summer temperature, 25.6 ºc (weather bureau 2000; grobler et al. 2002). mean annual rainfall in gauteng is 670 mm (gauteng 1997). old granitic and gneissic rocks at least 2 400 million years old (kerfoot 1987) are exposed in the egoli granite grassland area. the halfway house granites (fig. 3) of the johannesburg dome are intensively weathered with deep drainage lines resulting in a gently rolling topography, with shallow, coarse, nutrient-poor, well-drained soils. the granite areas are mostly covered by grassland vegetation though patches of woodland vegetation are found at sheltered sites on hillslopes and rocky outcrops within this veld type (grobler 2000; grobler et al. 2002, 2006). the bb land type predominates over the entire region (land type survey staff 1984, 1985, 1987) (fig. 3). the residual granitic soils are very shallow and poorly drained. the a horizon is often sandy and light in colour with little organic matter and with an increase in clay content with depth. the soils are invariably acid and very rich in silica (kerfoot 1987). vegetation current vegetation this tall grassland occurs over vast areas, usually on shallow, leached soils on the johannesburg granite dome. disturbed grassland or other disturbed areas such as road reserves or old fields, not cultivated for some years, are also usually hyparrhenia species-dominated and low in species richness. although some of these tall grasslands appear to be quite natural, they are mostly associated with an anthropogenic influence from recent or even iron-age times. these grasslands are characterised by the tall-growing dominant grass hyparrhenia hirta and the invader dwarf shrub seriphium plumosum (=stoebe vulgaris), indicating its low successional status or degraded condition. dense hyparrhenia species-dominated grassland mostly has low species richness, with only a few other species able to establish or survive in the shade of the dense sward of tall grass. most of these species are relict pioneers or early seral species. the most prominent species include the grasses cynodon dactylon, eragrostis chloromelas, e. racemosa, e. curvula and aristida congesta. forbs are rarely encountered, though a few individuals of species such as anthospermum hispidulum, pseudognaphalium luteo-album, conyza albida, c. podocephala, crabbea angustifolia, helichrysum nudifolium and h. rugulosum are often present. the woody layer, which has a very low cover, consists of small clumps of indigenous trees and shrubs widely scattered within this grassland. the scanty woody species include the trees rhus pyroides and ziziphus mucronata, together with the scandent shrub ziziphus zeyheriana. declared alien invasive species such as melia azedarach, eucalyptus species, pinus species and planted ornamentals are often present. typical species found in anthropogenic hyparrhenia hirta-dominated egoli granite grassland are indicated below: bredenkamp.indd 61 2006/10/15 10:51:02 pm koedoe 49/2 (2006) 62 issn 0075-6458 trees and shrubs acacia caffra (thunb.) willd. *eucalyptus species gymnosporia buxifolia l. szysyal *melia azedarach l. olea europaea l. ssp. africana (mill.) p.s.green *pinus species rhus lancea l.f. rhus leptodictya diels rhus pyroides burch. ziziphus mucronata willd. ziziphus zeyheriana sond. grasses aristida bipartita (nees) trin. & rupr. aristida canescens henrard aristida congesta roem. & schult. ssp. barbicollis (trin. & rupr.) de winter aristida congesta roem. & schult. ssp. congesta cynodon dactylon (l.) pers. diheteropogon amplectens (nees) clayton elionurus muticus (spreng.) kunth eragrostis chloromelas steud. eragrostis curvula (schrad.) nees eragrostis gummiflua nees eragrostis plana nees eragrostis racemosa (thunb.) steud. eragrostis rigidior pilg. heteropogon contortus (l.) roem. & schult. hyparrhenia filipendula (hochst.) stapf hyparrhenia hirta (l.) stapf melinis repens (willd.) zizka pogonarthria squarrosa (roem. & schult.) pilg. sporobolus africanus (poir.) robyns & tournay trichoneura grandiglumis (nees) ekman forbs acalypha angustata sond. anthospermum hispidulum e.mey. ex sond. asparagus laricinus burch. asparagus suaveolens burch. bidens formosa (bonato) sch.bip. bidens pilosa l. chamaecrista mimosoides (l.) greene cirsium vulgare (savi) ten. commelina africana l. conyza albida spreng. cucumis zeyheri sond. cyperus species elephantorrhiza elephantina (burch.) skeels felicia muricata (thunb.) nees gazania krebsiana less. geigeria burkei harv. gomphocarpus fruticosus (l.) aiton f. helichrysum nudifolium (l.) less. helichrysum rugulosum less. hermannia depressa n.e.br. hibiscus aethiopicus hypoxis rigidula lactuca species ledebouria marginata (baker) jessop nidorella hottentotica dc. pseudognaphalium luteo-album (l.) hilliard & b.l.burtt schkuhria pinnata (lam.) cabrera seriphium plumosum l. tagetes minuta l. verbena bonariensis l. verbena brasiliensis vell. vernonia oligocephala (dc.) sch.bip. ex walp. walafrida densiflora (rolfe) rolfe zinnia peruviana (l.) l. original vegetation the original vegetation is thought to be typical bankenveld (acocks 1988) or rocky highveld grassland (bredenkamp & van rooyen 1996). bredenkamp & brown (1998) found a few relict sites which indicate that the original vegetation on the shallow granitic soils of the johannesburg granite dome could have been a variant of the monocymbium ceressiforme-loudetia simplex grassland (bredenkamp & brown 2003). this vegetation is found in areas where little or no disturbance is evident. the herbaceous layer covers approximately 80 % and is dominated by the grasses loudetia simplex, trachypogon spicatus, schizachyrium sanguineum, monocymbium ceresiiforme, digitaria monodactyla, eragrostis racemosa, andropogon shirensis, brachiaria serrata, alloteropsis semialata, bewsia biflora and themeda triandra. the woody layer consists mainly of a few scattered individuals of the trees rhus pyroides, r. leptodictya, ziziphus mucronata, the dwarf shrubs protea welwitschii, lopholaenia coriifolia, and the geoxylophyte parinari capensis that are locally prominent. the grasses panicum natalense, urelytrum agropyroides, tristachya leucothrix, cymbopogon excavatus and elionurus muticus are also abundant together with the forbs cyanotis speciosa, bulbostylis burchellii, crabbea acaulis, anthospermum hispidulum and senecio venosus. the grass hyparrhenia hirta is often present and may become more prominent at disturbed sites. this grassland is characterised by a high species richness with a patchy dominance of bredenkamp.indd 62 2006/10/15 10:51:03 pm issn 0075-6458 63 koedoe 49/2 (2006) # halfway house granite # black reef # malmani # malmani # hospital hill #government # timeball hill # witwatersrand #muldersdrif # central rand # turffontein # jeppestown # klipriviersberg # klipriviersberg # timeball hill # hekpoort 30 0 30 60 km ab12 fa18 ab2 ab11 bb1 bb2 ab6 ab1 ba9 ba7 ba8 ib4 ba35 ab7 ab7 bb3 ba1 ba36 fa12 ba44 ab4 fa17 ib4 ib41 ib43 ib4 ib7 ib6 ib6 bb3 fb14 n n fig. 3. land types (above) and geology (bottom) within the egoli granite grassland various grass species and a large variety of forbs (louw 1970), representing a climax or close to climax condition. grazing by cattle is often found in this vegetation type, but the dominance of sour grass species often results in a low nutrient status of the grass during winter (kerfoot 1987; bredenkamp & van rooyen 1996). typical species found in the original egoli granite grassland are indicated below: trees and shrubs acacia caffra (thunb.) willd. celtis africana burm.f. elephantorrhiza elephantina (burch.) skeels gymnosporia buxifolia l. szysyal olea europaea l. ssp. africana (mill.) p.s.green rhus lancea l.f. rhus leptodictya diels rhus pyroides burch. ziziphus mucronata willd. ziziphus zeyheriana sond. grasses andropogon schirensis a.rich. aristida adscensionis l. aristida canescens henrard aristida congesta roem. & schult. ssp. congesta brachiaria serrata (thunb.) stapf cymbopogon excavatus (hochst.) stapf ex burtt davy cymbopogon pospischilli (k.schum.) c.e. hubb cynodon dactylon (l.) pers. digitaria brazzae (franch.) stapf diheteropogon amplectens (nees) clayton elionurus muticus (spreng.) kunth eragrostis chloromelas steud. eragrostis curvula (schrad.) nees eragrostis racemosa (thunb.) steud. harpochloa falx (l.f.) kuntze heteropogon contortus (l.) roem. & schult. hyparrhenia hirta (l.) stapf melinis repens (willd.) zizka monocymbium ceresiiforme (nees) stapf panicum natalense hochst. schizachyrium sanguineum (retz.) alston sporobolus pectinatus hack. themeda triandra forssk. trachypogon spicatus (l.f.) kuntze trichoneura grandiglumis (nees) ekman tristachya leucothrix nees tristachya rehmannii hack forbs acalypha angustata sond. aloe transvaalensis kuntze aloe greatheadii schönland anthericum species anthospermum hispidulum e.mey. ex sond. asparagus suaveolens burch. aster bakeranus burtt davy ex c.a.sm. babiana hypogea burch. becium obovatum (e.mey. ex benth.) n.e.br. blepharis subvolubilis c.b.clarke boophane disticha (l.f.) herb. bulbostylis hispidula (vahl) r.w.haines chascanum hederaceum (sond.) moldenke var. hederaceum chaetacanthus burchellii nees chlorophytum fasciculatum (baker) kativu crabbea acaulis n.e.br. crabbea angustifolia nees crassula capitella thunb. ssp. capitella crinum bulbispermum (burm.f.) milne-redh. & schweick. bredenkamp.indd 63 2006/10/15 10:51:03 pm koedoe 49/2 (2006) 64 issn 0075-6458 cucumis zeyheri sond. dicoma anomala sond. eriosema cordatum e.mey. eulophia species felicia muricata (thunb.) nees gladiolus crassifolius baker gnidia capitata l.f. haplocarpha scaposa harv. helichrysum aureonitens sch.bip. helichrysum miconiifolia hilliard helichrysum nudifolium (l.) less. hermannia depressa n.e.br. hypericum aethiopicum (bret.) n.k.b. hypericum lalandii choisy hypoxis hemerocallidea fisch. & mey. hypoxis multiceps buchinger ex baker hypoxis rigidula baker ipomoea obscura (l.) ker gawl. ipomoea ommaneyi rendle justicia anagalloides (nees) t.anderson kohautia amatymbica eckl. & zeyh. ledebouria ovatifolia (baker) jessop ledebouria marginata (baker) jessop lotononis foliosa bolus mariscus congestus (vahl) c.b.clarke neorautanenia ficifolius (benth.) c.a.sm. pentanisia angustifolia (hochst.) hochst. pentanisia prunelloides klotzsch ex eckl. & zeyh.) walp. raphionacme hirsuta (e.mey.) r.a.dyer ex e.phillips scabiosa columbaria l. schistostephium crataegifolium (dc.) fenzl ex harv. senecio venosus harv. tephrosia capensis (jacq.) pers. triumfetta sonderi ficalho & hiern tulbaghia acutiloba harv. turbina oblongata (e.mey. ex choisy) a.meeuse ursinia nana dc. vernonia natalensis sch.bip. ex walp. vernonia oligocephala (dc.) sch.bip. ex walp. origin of the current vegetation the shallow, nutrient-poor soils provide a habitat suited to the climax vegetation as discussed previously. some nutrients are available in the topsoil, supplemented from fallen leaf litter and decomposition, and also from ashes of burned herbaceous layer (if there was a fire). nutrients are quickly utilised by plants and are furthermore also quickly leached from the very coarse sandy soils. nutrient cycling is therefore very rapid. the system is quite stable and fairly predictable without much change caused by normal droughts or grazing. however, if overgrazed or disturbed to such an extent that degradation proceeds beyond a threshold, then recovery is very slow, due to reduced nutrient cycling and decreased nutrient availability and the vegetation may change to another domain of attraction (bosch 1989), different from the original climax vegetation, representing a plagioclimax (fig. 4). a change back to the original domain of attraction is unlikely if not impossible in the short and medium term. due to the granitically derived shallow nutrient poor soils these systems are sensitive and intolerant to frequent impacts such as heavy grazing, ploughing, trampling and general domestic activities. thus degradation occurs easily resulting in a change from the climax (high species richness) vegetation to an anthropogenic hyparrhenia hirta (low species richness) dominated vegetation type. very often hyparrhenia species-dominated grasslands occur on ancient lands in the central variation of the bankenveld (acocks 1988) and in the surroundings of archaeological sites (bredenkamp & brown 2003), where the inhabitants had a mosaic of cultivated lands and grazing of domestic stock. the more recent european settlers also had a profound effect on the natural vegetation as they developed the cities of johannesburg and pretoria following the discovery of gold more than a century ago. it seems that the degraded sites developed into hyparrhenia species-dominated grasslands, which tend to be stable for a very long time. moll (1965) and smits et al. (1999) also indicated that hyparrhenia species-dominated tall grasslands are anthropogenic in origin. importance the substrate of the johannesburg granite dome (halfway house granites, fig. 3) is suitable for development. the area is well situated between johannesburg, the economic powerhouse of south africa, and pretoria the administrative capital of the country. there is an enormous demand for developable land in this area, which has resulted in the loss of large tracts of the original egoli granite grassland. due to general disturbance, ploughing and degradation, an estibredenkamp.indd 64 2006/10/15 10:51:04 pm issn 0075-6458 65 koedoe 49/2 (2006) mated 60 % of the remaining original egoli granite grassland has been transformed to hyparrhenia hirta dominated grassland. only relatively small scattered pockets of the original grassland are still intact and these are considered rare and highly threatened. due to its high species richness and restricted occurrence, this endemic grassland has a high conservation value. furthermore, the bottomland areas and wetlands within the egoli granite grassland provide suitable habitat for various sensitive fauna species such as the grass owl tyto capensis (red listed), marsh sylph metisella meninx (vulnerable), and the giant bullfrog pyxicephalus adspersus (near threatened). pioneer original egoli granite grassland sere sere plagioclimax domain of attraction ii domain of attraction i hyparrhenia dominated grassland threshold = degradation = succession conclusion egoli granite grassland is a poorly conserved, severely transformed, highly fragmented and degraded vegetation type. it is estimated that only 22 % of the original extent of egoli granite grassland remains in its original state. many of these remnant areas are likely to be destroyed in the near future due to previously authorised developments (e.g. cosmo city to the north-west of johannesburg) or illegal activities. it is therefore essential that any viable remnant patch of original egoli granite grassland is conserved. areas associated with transformed grassland, where hyparrhenia hirta is dominant and species richness is low, have a low value with respect to achieving the national conservation target for egoli granite grassland. it should be noted however, that hyparrhenia hirta dominated grassland may be valuable for the conservation of sensitive fauna, e.g. grass owl (tyto capensis) and giant bullfrog (pyxicephalus adspersus). a mixture of various grasses and high forb diversity renders the original egoli granite grassland with a high conservation value and conservation of remnant areas is especially important since it is unlikely that the transformed anthropogenic grassland will return to the original climax vegetation. since egoli granite grassland is endemic to gauteng, its protection is both a provincial and national priority. it also forms part of a global ecoregion in crisis, an area where biodiversity and ecosystem services are at greatest risk and focused on-the-ground conservation action are required (hoekstra et al. 2005). references acocks, j.p.h. 1988. veld types of south africa. 3rd ed. memoirs of the botanical survey of south africa 57: 1–146. barnes, k.n. (ed.) 2000. the escom red data book of birds of south africa, lesotho and swaziland. johannesburg: birdlife south africa. bosch, o.j.h. 1989. degradation of the semi-arid grasslands of southern africa. journal of arid environments 16: 165–175. bredenkamp, g.j & l.r. brown. 1998. a vegetation assessment of the northern areas of the northern metropolitan local council. pretoria: ekotrust cc. unpublished report. bredenkamp, g.j & l.r. brown. 2003. a reappraisal of acocks’ bankenveld: origin and diversity of vegetation types. south african journal of botany 69(1): 7–26. fig. 4. a model indicating degradation of original egoli granite grassland climax and succession towards a hyparrhenia hirta dominated vegetation type. bredenkamp.indd 65 2006/10/15 10:51:04 pm koedoe 49/2 (2006) 66 issn 0075-6458 bredenkamp, g.j. & n. van rooyen. 1996. rocky highveld grassland. p. 39. in: low, a.b. & a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism, fahrig, l. 2003. effects of habitat fragmentation on biodiversity. annua. review of ecology evolution and systematics 34: 487–515. gauteng, 1997. state of the environment in gauteng: a preliminary report. gauteng department of agriculture, conservation and environment. unpublished report, csir environmentek. kerfoot, o. 1987. the geology, physiography and vegetation of the witwatersrand floral region. pp. 1-7. in: lowrey, t.k. & s. wright (eds.). the flora of the witwatersrand volume 1: the monocotyldonae. johannesburg: witwatersrand university press. grobler, c.h. 2000. the vegetation ecology of urban open spaces in gauteng. msc thesis, university of pretoria, pretoria. grobler, c.h., g.j. bredenkamp, l.r. brown. 2002. natural woodland vegetation and plant species richness of the urban open spaces in gauteng, south africa. koedoe 45: 19–34 grobler, c.h., g.j. bredenkamp, l.r. brown. 2006. primary grassland communities of urban open spaces in gauteng, south africa. south african journal of botany 72: 367–377. hoekstra j.m., t.m. boucher, t.h. ricketts & c. roberts. 2005. confronting a biome crisis: global disparities of habitat loss and protection. ecology letters 8: 23–29. land type survey staff. 1984. land types of the maps 2626 wes-rand, 2726 kroonstad. memoirs on the agricultural natural resources of south africa 4: 1–342. land type survey staff. 1985. land types of the maps 2628 east rand. 2630 mbabane. memoirs on the agricultural natural resources of south africa 4: 1–261. land type survey staff. 1987. land types of the maps 2526 rustenburg, 2528 pretoria. memoirs on the agricultural natural resources of south africa 8: 1–391. louw, w.j. 1970. klipveld studies 1: checklist of vegetation. journal of south african botany 36: 199–206 minter, l.r., m. burger, j.a. harrison, h.h. braack, p.j. bishop & d. koepfer (eds.). 2004. atlas and red data book of the frogs of south africa, lesotho and swaziland. washington, dc: smithsonian institution. moll, e.j. 1965. an account of the plant ecology of the upper umgeni catchment. msc thesis, university of natal, pietermaritzburg mucina, l., rutherford, m.c. & l.w. powrie (eds.). 2005. vegetation map of south africa, lesotho & swaziland, 1:1 000 000 scale sheet maps. pretoria: south african national biodiversity institute. smits, n.a.c., g.j. bredenkamp, l. mucina & j.e. granger. 1999. the vegetation of old fields in transkei. south african journal of botany 65: 414–420 weather bureau. 2000. prints of the climate statistics of the johannesburg leeukop, pretoria forum wb, johannesburg joubert park and krugersdorp kroningspark weather stations. pretoria: department of environment affairs and tourism. domain of attraction idomain of attraction i bredenkamp.indd 66 2006/10/15 10:51:04 pm reflections on the relationships between communities and conservation areas of south africa: the case of five south african national parks t.s. simelane, g.i.h. kerley and m.h. knight simelane, t.s., g.i.h. kerley and m.h. knight. 2006. reflections on the relationships between communities and conservation areas of south africa: the case of five south african national parks. koedoe 49(2): 85–102. pretoria. issn 0075-6458. an evaluation of the relationships between communities around addo elephant national park, mountain zebra national park, karoo national park, golden gate highlands national park and vaalbos national park shows that these communities have limited ecological knowledge and understanding of resources occurring within the parks. people within these communities rate relationships with their neighbouring parks as relatively poor. despite this, these communities are keen to support conservation and management of biodiversity through national parks. the study further revealed that two types of communities occur around the national parks of south africa. these are neighbouring and the distant communities. the distant communities are more urban in character than the neighbouring communities. these communities are heterogeneous with people from a variety of cultures. the varying lifestyle, age groups, cultural backgrounds and income levels lead to differences in expectations from the national parks by these communities. this, which is critical in determining the level of appreciation of conservation of biodiversity by communities around conservation areas, requires the attention of the park managers. they need to ensure that when distributing the benefits or opportunities linked to conservation, cultural, income and education differences among communities are considered and used as the basis for development and implementation of community development projects. key words: community based conservation, biodiversity, natural resources t.s. simelane, south african national parks, p.o. box 20419, humewood, 6013 and centre for african conservation ecology, department of zoology, p.o. box 77000, nelson mandela metropolitan university 6013 (saenvironet@ananzi.co.za); g.i.h. kerley, centre for african conservation ecology, department of zoology, p.o. box 77000, nelson mandela metropolitan university 6013 (graham.kerley@nmmu.ac.za); m.h. knight, south african national parks, p.o. box 20419, humewood, 6013 (mknight@nmmu.ac.za). issn 0075-6458 85 koedoe 49/2 (2006) introduction in the past ten years, there has been a major shift in the approach of managing natural resources, land and environment in south africa (kerley et al. 1999; nauta 2001). linked to this had been political demands for redistribution of land, wealth and access to natural resources (anc 1994; cobbett 1987; ramphele 1991; sachs 1990). the call for land redistribution saw an influx of land claim applications from traditional societies, some of which threatened the very existence of well-known conservation areas such as the kruger national park, st lucia wetland park and the kgalagadi transfrontier park. faced with these threats, conservation in south africa has been assumed to be under siege (ledger 1998). thus, conservation agencies like the south african national parks (sanparks) needed to adapt and respond swiftly by providing strategies that would assure the support of national parks by the local communities (hanekom & liebenberg 1994). in dealing with this challenge, sanparks adopted a proactive strategy of conservationbased human development and natural resource management (sanparks 1995, 1997). this included the establishment of various community development projects simelane.indd 85 2006/10/15 10:59:17 pm koedoe 49/2 (2006) 86 issn 0075-6458 in the national parks (sanparks 1995) and the consideration of the possibility of allowing communities around the parks to harvest resources (e.g. fuel wood) (sanparks 1997, 1998). in addition, parks management committees and community fora were formed. these were aimed at increasing dialogue around the management of natural resources and the participation of communities in the management of the parks in general (sanparks 1998). contractual park agreements, such as those introduced in the richtersveld national park (sanparks 1995; krog 2000), were also adopted and used as a strategy to proclaim new national parks, manage land handed over to local communities and when extending national park borders (sanparks 1995). however, sanparks still needed to adhere to, and maintain its main mission and objectives, which are: to conserve south africa’s natural heritage and biodiversity; to manage national parks according to the highest standards; to provide assurance of the appropriate use of the natural resources; and to ensure that national parks represent the diversity of south african plants, animals, landscapes, natural processes and cultural artifacts (sanparks 1995). the recognition of this need is a direct response to the concerns raised about, firstly, the possible negative impacts on biodiversity that might be caused by people gaining access to natural resources within the national parks, and secondly, the possibility of sanparks compromising its focus of promoting the conservation of natural resources and biodiversity (steele 1991; simmonds 1998). in addition to these concerns, sanparks was not sure of the role it should play in community development (e.g. greyling & huntley 1984). it was, in fact, not clear whether or not the parks would be able to meet the expectations of the local communities. this was exacerbated by the lack of welldefined community structures with which sanparks could work, and also the lack of • • • • effective strategies to identify communities for incorporation into established community development projects. these complications are not unique to sanparks (hildgard et al. 1998). many conservation agencies in africa, and elsewhere, face similar challenges when attempting to implement community-based conservation strategies (bell 1982, anon 1989). in general, conservation agencies struggle to embrace the concept of human development through conservation (western & henry 1979; zimbabwe trust 1990; brandon & wells 1992; sibanda 1995), apparently due to the lack of reliable models that can be followed by the conservation agencies (rothley 1999). as a result, most community-based conservation projects initiated by conservation agencies tend to fail (richards 1993; kiss 2004). while heavy reliance of these initiatives on foreign funding and limited participation of communities in their management can be seen as a major cause of their failure in some instances like the zimbabwean campfire project, which drew international admiration, the increase in the number of people who demand benefits and a lack of well defined beneficiaries, is also a common cause for the demise of these initiatives (matzke & nabane 1990; sibanda & omwega 1996). in order to avoid this it is necessary to provide a clear definition of the local communities associated with a specific conservation area and to develop strategies that can enhance the full participation of local communities in the project, as well as to formulate frameworks that will guide the design of the project. with this background, this study evaluates the achievements of sanparks in implementing its community-based conservation strategy, an initiative that was introduced during 1994. this was done by: investigating attitudes towards tourists and the conservation of biodiversity by communities around addo elephant, mountain zebra, golden gate highlands, vaalbos and karoo national parks; • simelane.indd 86 2006/10/15 10:59:17 pm issn 0075-6458 87 koedoe 49/2 (2006) identifying the local communities associated with these parks, who can participate in development projects; highlighting the social, economic and educational profiles of the local communities around the five national parks; determining the level of ecological knowledge among these communities; determining the impact of education and economic status on the understanding and appreciation of conservation by these communities; and by determining if the community development projects initiated by sanparks during the past ten years were effective in developing conservation awareness, and whether or not tangible and intangible benefits were channelled to communities as a result of these projects. the study tested the hypothesis that attitudes towards conservation among local communities around protected areas could be related to ecological knowledge and perceived benefits. such interpretations may differ between age groups and people with different levels of education. the study also tested the assumption that communities around south african national parks desire to extract resources from conservation areas. this assumption may differ with age, between cultural groups and with different income levels. methods a questionnaire survey was conducted among local communities around addo elephant national park (aenp), mountain zebra national park (mznp), golden gate highlands national park (gghnp), karoo national park (krnp) and vaalbos national park (vnp). this aimed to obtain information about community attitudes towards the parks, tourists and resources within the parks. the survey also aimed to determine whether members of the local communities visited their nearby park, and whether or not communities have obtained benefits from the established community development projects. the survey did not attempt to exhaustively cover all socioeconomic issues of the communities, but rather focused on issues identified as being relevant to sanparks. a sample of 400 people, aged between 15 and 60 years, was interviewed in the vicinity of each park, a total of 2000 people. interviews were based on an informal discussion with each interviewee (arber 1996). some interviews took place in household situations where only one person in the household was interviewed. a household is here defined as all people who live together • • • • • and who make joint economic decisions (mohr & fourie 2003). the interviews were conducted by interviewers recruited from each sampled community and were held in appropriate local languages (afrikaans, isixhosa, setswana and sesotho). interviewers undertook a one-day training course. during training, they were acquainted with the format of the questionnaire and were introduced to the acceptable way of approaching interviewees. interviewers conducted trial interviews, using a sample of ten people at each park, before they started the formal interviews. interviewees were classified into six age groups (procter 1996). these were: group i (15–20 yrs), group ii (21–30 yrs), group iii (31–40 yrs), group iv (41–50 yrs), group v (51–60 yrs) and group vi (above 60 yrs). this allowed the assessment of income disparities and differences in education levels among people of different ages, and also the assessment of age-based differences in attitude towards tourists and conservation. for aenp, mznp and krnp, which have more than one cultural group living near the park, representation of all relevant groups was ensured by sampling two settlements around each park. at aenp, the valencia settlement, occupied mainly by coloured people, and nomathamsanqa, dominated by xhosa people, were surveyed. at mznp, michausdal (coloured) and lingelihle (xhosa) were surveyed. at krnp, kwamandlenkosi (xhosa) and newlands (coloured) were surveyed. at vnp, only one settlement was surveyed as only one cultural group existed. at gghnp, two communities—qwaqwa and kgubetswana—of the same cultural group (basotho) were intentionally included in the study, as these two communities had different political histories. under the previous political arrangements, qwaqwa settlement was located within the self-governing state of qwaqwa, while kgubetswana was located within the republic of south africa. it was anticipated that this might influence attitudes of the communities towards the gghnp. due to the large size of some of the settlements (table 1), it was impossible to include all parts of the settlements for each community in the survey. the focus was therefore given to those portions of the settlements that were closest to the national park. this approach was mostly applied to the qwaqwa community. the communities were then defined as either “neighbouring” or “distant”, based on their proximity (see results) to the park, and were assessed as being serviced or less serviced, based on available infrastructure and social services. these definitions accord with the sanparks approach to identifying communities as stakeholders. this approach differs from other similar surveys conducted elsewhere in africa, where communities and conservation areas are more tightly coupled. additional information about the number of people in each settlement and other general information pertaining to community assessment were obtained from the offices of the local municipality, through the social ecologists of each park. simelane.indd 87 2006/10/15 10:59:17 pm koedoe 49/2 (2006) 88 issn 0075-6458 table 1 neighbouring and distant communities of the five national parks (aenp = addo elephant, mznp = mountain zebra, vnp = vaalbos, gghnp = golden gate highlands, krnp = karoo national parks) park settlement race / ethnic group estimated number of people as provided by local municipality distance from the park (km) class type of settlement aenp bergsig xhosa 2000 30 distant less serviced bontmug xhosa 500 30 distant less serviced valenciaa coloured 1500 20 neighbouring serviced enon xhosa 1800 38 distant less serviced nomathamsanqaa xhosa 27 500b 1 neighbouring serviced bersheba xhosa 1500 38 distant less serviced mznp lingelihlea xhosa 35000 27 distant serviced michausdala coloured 9000 27 distant serviced vnp longlandsa batswana & coloured 2375 5 neighbouring less serviced gghnp kgubetswanaa basotho 2500 17 neighbouring serviced qwaqwaa basotho 1000000 35 distant serviced kestell basotho 4500 40 distant serviced krnp kwamandlenkosia xhosa 5157 3 neighbouring serviced rustdene newlandsa essopville newtown hooyvlakte coloured 17341 5 neighbouring serviced a communities in which interviews were conducted. b from geach (1997). statistical analyses a descriptive analysis of age and levels of income and education of interviewees was calculated using sigma stat™. differences in age groups among the parks were compared using a kruskal-wallis anova (zar 1984). differences in income and levels of education between communities of vnp and aenp (parks surrounded by less serviced neighbouring communities) and gghnp, mznp and krnp (parks surrounded by serviced distant communities) were compared using a student’s t-test. the numbers of people within households between parks with distant (gghnp, mznp & krnp) communities and those with neighbouring (vnp and aenp) communities were compared using a kruskal wallis anova. the attitudes of respondents towards tourists were measured according to the likert scaling method, where respondents are asked to respond to an item through answers such as positive, neutral or negative (arber 1996). the prediction that communities around conservation areas have a negative attitude towards conservation was tested using the chi-squared test (zar 1984). differences of visits to the parks by local people as a way of supporting tourism and the management of the parks were compared using the chi-squared test (zar 1984). results descriptive analysis of the results responses to questions varied, with some respondents not answering all questions. this did not appear to create a bias against particular questions, but it did provide a small deviance from the intended target sample of 400 interviewees at each park. the effects of this deviation are reflected in the results. simelane.indd 88 2006/10/15 10:59:17 pm issn 0075-6458 89 koedoe 49/2 (2006) two types of community (neighbouring and distant communities) were identified. neighbouring communities were defined as being closer than 20 km from the park, while distant communities were defined as being further than 20 km from the park (table 1). distant communities are normally associated with the town near where the park is situated. for example, for mznp, the distant community was in cradock (situated 27 km from mznp), while for gghnp, the distant community was in qwaqwa (situated 45 km from gghnp) (table 1). addo elephant national park, krnp and mznp had communities with mixed cultures and were composed of two recognisable language groups, isixhosa and afrikaans. sesotho-speaking people dominated gghnp and setswana and afrikaans-speaking people dominated vnp. communities around aenp, krnp and mznp were notably settled along the cultural lines brought about by the previous group areas act of 1966. such an arrangement was more prevalent in the nomathamsanqa (xhosa) and valencia (coloured) communities of aenp, the lingelihle (xhosa) and michausdal (coloured) communities of mznp, and the kwamandlenkosi (xhosa) and newlands (coloured) communities of krnp. the kgubetswana and qwaqwa communities of gghnp, the nomathamsanqa of aenp, the michausdal and lingelihle of mznp and all communities around krnp (table 1) featured modern amenities such as tarred roads, electricity, tap water and sewage systems. these communities were classified as serviced (table 1), as the presence of these amenities indicated better social conditions and lifestyles. the remaining communities were classified as less serviced (table 1) as they had neither tarred roads nor electricity and appeared more rural in the nature of their services. in all parks, the age distribution was typical of a rapidly growing population (starr & taggart 1987), with young people (i.e. group ii, aged 21–30 yrs) making up the highest proportion of the population. for aenp, the average age of respondents was 39.6 ± 0.9 (mean ± se), vnp (34.7 ± 0.9), mznp (23.8 ± 0.6), krnp (39.7 ± 0.6) and gghnp (30.8 ± 0.7). the age distribution differed significantly between parks (kruskal-wallis anova = 141.5, df = 4, p < 0.001), with gghnp having a larger number of young people (table 2) than all other parks. numbers of people within households differed significantly between the parks with gghnp (mean = 5.0 ± 0.4), mznp (mean = 4.9 ± 0.03) and vnp (mean = 5.7 ± 1.4) having a significantly (kruskalwallis anova, h = 75, df = 4, p < 0.05) table 2 age distribution (within six age groups) of people interviewed around the five national parks (park codes as for table 1). column totals indicate the numbers of respondents who provided their ages for each national park. distribution is expressed as: a) actual numbers (n) recorded in each group, and b) their percentages (%) within the age group group aenp mznp vnp krnp gghnp mean (± sd) across parksn % n % n % n % n % i (15 – 20 yrs) 36 9.0 52 13.6 57 19.3 5 1.3 89 23.8 13.4 ± 7.9 ii (21 – 30 yrs) 122 30.7 153 40.0 106 35.8 109 28.4 139 37.4 34.5 ± 4.3 iii (31 – 40 yrs) 79 19.9 90 23.6 43 14.5 107 27.9 80 21.4 21.5 ± 4.4 iv (41 – 50 yrs) 68 17.4 43 11.3 28 9.5 84 21.9 39 10.4 14.1 ± 4.8 v (51 – 60 yrs) 32 8.0 31 8.0 28 9.5 48 12.5 14 3.5 8.3 ± 2.9 vi (> 60 yrs) 60 15.0 13 3.5 34 11.4 31 8.0 13 3.5 8.3 ± 4.5 total 397 100 382 100 296 100 384 100 374 100 simelane.indd 89 2006/10/15 10:59:17 pm koedoe 49/2 (2006) 90 issn 0075-6458 higher average number of people living in each household (fig. 1) than aenp (mean = 4.4, df = 4, p < 0.05) and krnp (mean = 4.2, df = 4, p < 0.05). the education levels of those people around national parks with distant communities differed significantly (kruskal-wallis, h = 1.35, df = 4, p < 0.05) from those people associated with neighbouring rural communities. mountain zebra national park (mean educational grade = 8), gghnp (mean educational grade = 8) and krnp (mean educational grade = 8), had larger numbers of people with grade fig. 1. mean (± sd) number of people living in households of the communities around the five studied national parks. aenp = addo elephant national park; mznp = mountain zebra national park; vnp = vaalbos national park; krnp = karoo national park; gghnp = golden gate highlands national park). fig. 2. distribution of respondents within ten education levels (standards). the data reflects the status of education within the communities adjacent to the studied national parks. (park codes as for fig. 1). 7 6 5 4 3 2 1 0 m ea n aenp mznp vnp krnp gghnp parks aenp 0 20 40 60 80 100 0 1 2 3 4 5 6 7 8 9 10 education levels nu m be ro fin div idu als gghnp 0 20 40 60 80 100 120 140 160 0 1 2 3 4 5 6 7 8 9 10 education levels n um be ro fi nd iv id ua ls krnp 0 50 100 150 200 250 0 1 2 3 4 5 6 7 8 9 10 education levels nu m be ro fin div idu als mznp 0 50 100 150 200 0 1 2 3 4 5 6 7 8 9 10 education levels n um be ro fi nd iv id ua ls vnp 0 10 20 30 40 50 0 1 2 3 4 5 6 7 8 9 10 education levels n um be ro fi nd iv id ua ls simelane.indd 90 2006/10/15 10:59:18 pm issn 0075-6458 91 koedoe 49/2 (2006) ten education than aenp (mean educational grade = 7) and vnp (mean educational grade = 6) (fig. 2). for all five parks, the highest proportion (47.7 %, n = 648) of people with grade twelve education were aged between 21 and 30 years (group ii). addo elephant national park and vnp had a comparatively larger percentage (23 % and 35 % respectively) of people with no formal education (classified as illiterate) than gghnp, krnp and mznp. this was more common among older people (group v, 51– 60 years and group vi, > 60 years). at mznp and krnp, two age groups (mznp group ii (21–30 years), krnp group iv (41–50 years) had an unusually high proportion of people with no formal education (fig. 3). this contrasts with the other parks where the low education levels were mainly in three groups (groups ii (21–30 yrs), v (51–60 yrs) and vi (> 60 yrs) (fig. 3). aenp (34.0 %), mznp (21.0 %) and vnp (48.0 %) had comparatively more people with a monthly income ranging between r400 to r500 (table 3). this category was classified as a low-income category. income categories, however, differed significantly (kruskalwallis anova, h = 273.8, df = 4, p < 0.001) between the parks, with krnp (14.0 %) fig. 3. percentages of people with no formal education (illiterate) within six age groups, interviewed from the five studied national parks (group i = 15-20 yrs, group ii = 21–30 yrs, group iii = 31-40 yrs, group iv = 41-50 yrs, group v = 51-60 yrs and group vi = > 60 yrs) . (park codes as for fig. 1). aenp 0 10 20 30 40 50 60 i ii iii iv v vi age group p er ce nt ag e krnp 0 10 20 30 40 i ii iii iv v vi age group p er ce nt ag e mznp 0 5 10 15 20 25 30 35 i ii iii iv v vi age group p er ce nt ag e gghnp 0 5 10 15 20 25 30 i ii iii iv v vi age group p er ce nt ag e vnp 0 10 20 30 40 50 i ii iii iv v vi age group p er ce nt ag e simelane.indd 91 2006/10/15 10:59:18 pm koedoe 49/2 (2006) 92 issn 0075-6458 and mznp (11.0 %) having more people in the higher income brackets (r2001–r3000) than aenp (1.0 %), gghnp (7.0 %) and vnp (0.0 %) (table 3). while this was more expressed among parks, the distribution of income within age groups reflected that, for vnp (r = -0.0001), mznp (r = -0.24), gghnp (r = -0.33) and krnp (r = -0.09), income was negatively correlated to age, indicating that young people had higher incomes than older people. for aenp (r = 0.13), it was positively correlated to age, reflecting that older people at this park had higher incomes than young people. the overall comparison of income among the parks reflected that the average monthly incomes of communities near parks with distant communities (i.e. mznp, r1342; gghnp, r1279; and krnp, r2285) were significantly (t = 4.7, df = 1, p < 0.001) higher than those of parks with neighbouring rural communities (i.e. aenp, r650; and vnp, r496) (table 3). with an average of 4.9 ± 2.1 people per household in studied communities (both rural and urban) (fig. 1), it was calculated that people around the national parks we studied survive on an estimated daily income of r3.28 per person. social welfare in the form of state pensions (restricted to groups v and vi) was the main source of income in neighbouring, less serviced communities. private enterprises, such as farms (as in aenp) and diamond mines (as in vnp), provided additional jobs for neighbouring less-serviced communities of aenp and vnp. many of these jobs were casual and seasonal. in all parks, young people (group i) (63 %) in both distant and neighbouring communities were still at school while groups ii (21–30 yrs), iii (31– 40 yrs) and iv (41–50 yrs) of neighbouring communities were either not working (55 %) or had seasonal jobs (32 %) on the farms (as in aenp) or diamond mines (as in vnp). in distant communities, groups ii (21–30 yrs), iii (31–40 yrs) and iv (41–50 yrs) were either not working or held a permanent job, with teaching being a common profession. table 3 income distribution among people in the communities of the five studied national parks. (park codes as for table 1) income category (rands/month) aenp (n = 277) mznp (n = 185) gghnp (n = 145) krnp (n = 250) vnp (n = 141) mean (± sd) across parks % % % % % <100 2.0 0.0 1.0 0.0 0.0 0.6 ± 0.8 100-200 16.0 3.0 8.0 2.0 3.0 6.4 ± 5.2 201-300 12.0 4.0 8.0 2.0 7.0 6.6 ± 3.4 301-400 8.0 2.0 3.0 4.0 14.0 6.2 ± 4.4 401-500 34.0 21.0 15.0 14.0 48.0 26.4 ± 12.9 501-600 3.0 5.0 4.0 1.0 9.0 4.4 ± 2.7 601-700 3.0 4.0 3.0 1.0 8.0 3.8 ± 2.3 701-800 6.0 9.0 8.0 4.0 4.0 6.2 ± 2.0 801-900 1.0 2.0 7.0 1.0 4.0 3.0 ± 2.3 901-1000 9.0 7.0 6.0 5.0 1.0 5.6 ± 2.7 1001-2000 5.0 22.0 20.0 22.0 2.0 14.2 ± 8.8 2001-3000 1.0 11.0 7.0 15.0 0.0 6.8 ± 5.7 3001-4000 0.0 6.0 8.0 14.0 0.0 5.6 ± 5.3 4001-5000 0.0 3.0 1.0 6.0 0.0 2.0 ± 2.3 >5000 0.0 1.0 1.0 8.0 0.0 2.0 ± 3.0 simelane.indd 92 2006/10/15 10:59:18 pm issn 0075-6458 93 koedoe 49/2 (2006) the neighbouring communities of aenp, mznp, gghnp and krnp mainly used electricity as the primary source of energy (table 4). this form of energy was either supplemented with paraffin, gas or fuel wood (table 4). more people (55.5 %) in aenp used electricity as their main source of energy. the longlands community of vnp used either fuel wood or paraffin as their only source of energy (table 4). this indicated an increased level of poverty within this community, as well as the possible demand of fuel wood for energy among this community. an average of 77 % of respondents (n = 1829) were positive about the existence of the parks near their settlements (table 5). eighty percent of respondents in the communities of aenp, krnp, vnp and mznp were positive or satisfied with their national parks (table 5), and assessed their relationship with table 4 percentages of respondents using different forms of energy in the communities sampled around the five parks. (park codes as for table 1) type of energy used aenp mznp gghnp krnp vnp mean (± sd) across parks% % % % % electricity only 55.5 24.4 25.3 36.3 0.0 28.3 ± 18.0 paraffin only 3.8 7.7 19.7 1.5 0.0 6.5 ± 7.1 fuel wood only 6.3 0.0 6.0 8.8 0.0 4.2 ± 3.6 gas only 3.8 0.0 8.5 0.0 0.0 2.5 ± 3.4 paraffin and fuel wood 4.9 0.0 18.0 1.8 100.0 24.9 ± 38.1 paraffin and gas 0.6 0.0 6.5 0.8 0.0 1.6 ± 2.5 electricity and gas 0.5 13.9 6.4 7.5 0.0 5.7 ± 5.1 electricity and paraffin 16.7 5.7 3.3 4.0 0.0 5.9 ± 5.7 electricity and fuel wood 7.9 19.7 2.6 31.5 0.0 12.3 ± 11.7 electricity, paraffin and fuel wood 0.0 28.6 0.0 7.5 0.0 7.2 ± 11.1 table 5 attitude (mean %) of local communities towards tourists and adjacent national parks. attitudes were measured according to the likert scale (arber 1996). (park codes as for table 1) feeling aenp mznp gghnp vnp krnp mean (± sd) across parks attitude towards the park happy 72.0 61.0 73.0 87.0 89.0 76.4 ± 10.4 do not know 16.0 20.0 24.0 9.0 8.0 15.4 ± 6.2 unhappy 12.0 9.0 3.0 4.0 3.0 6.2 ± 3.7 attitude towards tourists happy 83.0 86.0 65.0 81.0 86.0 80.2 ± 7.8 neutral 12.0 9.0 31.0 11.0 13.0 15.2 ± 8.0 unhappy 5.0 5.0 4.0 8.0 13.0 7.0 ± 3.3 assessment of the relationship good 45.0 2.0 57.0 54.0 43.0 40.2 ± 19.8 fair 32.0 17.0 39.0 13.0 42.0 28.6 ± 11.6 poor 23.0 56.0 4.0 33.0 15.0 26.2 ± 17.7 simelane.indd 93 2006/10/15 10:59:18 pm koedoe 49/2 (2006) 94 issn 0075-6458 table 6 opinions of the neighbouring communities regarding the importance of tourists in the five studied national parks and their towns (park codes as for table 1) data expressed as % of respondents) opinion aenp mznp gghnp vnp krnp mean (± sd) across parks parks are economically important to the region 73.0 92.1 65.0 74.1 86.3 78.1 ± 8.9 increase international exposure 12.2 7.9 25.0 24.7 3.6 14.7 ± 8.0 provide information 2.2 0.0 2.5 0.0 4.3 1.8 ± 1.5 create job opportunities 7.9 0.0 0.8 0.0 5.4 2.8 ± 3.0 increase the purchase of local arts and crafts 0.4 0.0 6.7 0.0 0.4 1.5 ± 2.4 do not know 4.3 0.0 0.0 1.2 0.0 1.1 ± 1.5 neighbouring parks as being good (table 5). however, gghnp had comparatively fewer positive responses from respondents (65 % being positive about the park) and mznp had fewer respondents who assessed their relationship as good (table 5). awareness that parks are economically important to local communities was common among all residents around these national parks (table 6). however, communities were not aware of the direct development opportunities that are provided by the presence of the parks near their settlements (table 6). very few respondents indicated that they were aware that parks are the main source of attraction for tourists, or that the park increases the international exposure of their areas and provides job opportunities (table 6). despite the strong support for the parks and awareness of the benefits associated with tourism and conservation, few people were aware of the benefits that they were deriving through development projects initiated by sanparks during the period of the study (table 7). most people (86 %), however, felt that the relationship between the local communities and neighbouring parks could be sustained through job creation (65 %), assistance in skills development (25 %), provision of environmental or conservation education (5 %) and the improvement of communication (5 %). this was interpreted as an indication of the willingness of the 0 10 20 30 40 50 60 70 80 krnp mznp aenp gghnp vnp park % vi sit ed yes no fig. 4. percentages of people who indicated that they visited the nearby parks. (park codes as for fig. 1). communities to cooperate with sanparks’ community conservation initiatives. an average of 42.4 % of respondents (n = 961) indicated that they had visited the nearby park at least once. visits to the parks were positively correlated (r = 0.34, df = 4, p < 0.05) with age, with young people visiting the nearby park either once or twice and older people either never visiting the park or unable to visit due to age. krnp (76 %) and mznp (58 %) had the highest percentages of visits by local residents (fig. 4). visits were for a number of purposes and varied between rural neighbouring communities and distant urban communities (table 8). visits for game viewing, enjoyment, picnics, parties and attending functions were mostly cited by people from urban distant communities (table 8). visits for work simelane.indd 94 2006/10/15 10:59:18 pm issn 0075-6458 95 koedoe 49/2 (2006) table 7 list of benefits realised by communities around the five studied national parks. variations in the number of respondents reflect the deviance created by the willingness of respondents to provide answers. park codes as for table 1. data expressed as % of respondents. (note that some respondents provided more than one answer) park settlement benefits mentioned no. of respondents aenp nomathamsanqa (n = 300) provided building material 18.9 mayibuyendlovu project 15.2 sponsored local sports 6.4 assisted during elections 4.7 do not know 1.7 nothingw 0.6 no response 52.5 total 100 valencia (n = 100) nothing 100 total 100 mznp lingelihle (n = 200) nothing 50 michausdal (n = 200) nothing 50 total 100 gghnp kgubetswana (n = 200) environmental education 43.1 animals (for aesthetic use) 15.5 conservation 6.9 no response 34.5 total 100 qwaqwa (n = 200) do not know 36.4 environmental education 10.8 nothing 7.3 no response 45.5 total 100 vnp longlands (n = 400) jobs during road construction 15.9 environmental education 5.5 do not know 5.9 nothing 1.4 no response 71.3 total 100 krnp kwamandlenkosi (n = 100) environmental education 1.9 skills development programme 2.6 provide job opportunities 21.9 bring tourists closer to us 14.2 do not know 54.2 nothing 5.2 total 100 newlands (n = 300) offers an ideal place to relax 3.3 provide job opportunities 18.6 provide recreation facilities 11.6 environmental education 6.1 education excursion 1.1 conservation awareness 0.8 conservation services 4.2 assist on some school projects 2.5 provide houses for workers 3.0 bring tourists closer to us 1.7 nature conservation clubs 5.5 development opportunities 8.6 nothing 3.9 no response 29.1 total 100 simelane.indd 95 2006/10/15 10:59:18 pm koedoe 49/2 (2006) 96 issn 0075-6458 reasons or to seek employment were mostly cited by people from neighbouring rural communities. parks with distant communities had a significantly higher number of visits (χ2 = 40.6, df = 2, p < 0.01) than those with neighbouring communities (table 8). since their income was comparatively higher than that of neighbouring rural communities, this was interpreted as being due to the availability of extra disposable money for these communities. in addition to the various cited reasons for visiting the parks, activities of the social ecologists contributed towards motivating local people to visit the parks (table 8). this was expressed more in aenp (8.4 %) and krnp (8.3 %), where there were comparatively more people visiting the park to attend meetings or workshops organised by the social ecologist (table 8). although it was expected that past politics and the history of managing parks might have highly influenced communities not to visit their neighbouring park, the results reflected that this was not the case (table 9). very few people indicated that they did not visit the parks either because of a history of being removed or through certain forms of exclusion (table 9). the perception was expressed that national parks were the domain of white people, but this was limited to aenp (table 9). lack of money, transport, awareness and interest were the common reasons for people not visiting the parks. negative attitudes towards the local residents were also cited as an additional reason for people not visiting the parks (table 9), although this was limited to aenp and krnp. in all five parks, the level of knowledge of resources that are available within the parks was significantly low (kruskal-wallis anova, h = 0.34, df = 4, p < 0.05), with interviewees citing not more than five resources as occurring within the parks (aenp (mean ± se = 2.1 ± 0.2), vnp (mean = 1.2 ± 1.01 ), mznp ( mean = 1.3 ± 0.01), gghnp (mean = 2.1 ± 0.4) and krnp (mean = 1.5 ± 0.3)). the majority of resources provided were table 8 reasons provided by respondents for visiting the parks, expressed as the percentage of respondents who visited the park for the specified reason. (park codes as for table 1) reasons for visiting the park aenp (n = 178) mznp (n = 225) gghnp (n = 136) krnp (n = 302) vnp (n = 120) mean (± sd) % across parks game viewing 21.3 19.1 24.3 10.6 18.3 18.7 ± 4.6 educational tour 14.0 4.0 13.2 6.3 0.0 7.5 ± 5.4 attend workshop 8.4 0.9 0.0 8.3 2.5 4.0 ± 3.6 delivery 2.3 0.0 0.0 0.3 0.0 0.5 ± 0.9 was working there 4.5 0.9 2.2 3.3 42.5 10.7 ± 15.9 visit a friend or relative 5.1 7.1 4.4 6.0 5.8 5.7 ± 0.9 looking for job 0.6 10.7 4.9 0.3 15.0 6.3 ± 5.8 picnic / party / function 20.2 16.0 21.8 27.3 12.5 19.6 ± 5.1 to fetch something 1.7 0.0 0.7 0.0 0.0 0.5 ± 0.7 for enjoyment 18.5 41.3 20.6 22.5 1.7 20.9 ± 12.6 with a church 1.7 0.0 0.7 0.3 0.0 0.5 ± 0.6 camping 0.0 0.0 0.0 10.8 0.0 2.2 ± 4.3 hiking 0.0 0.0 0.0 3.3 0.0 0.7 ±1.3 eat out 0.0 0.0 0.0 0.3 0.0 0.1 ± 0.1 the curio shop 0.0 0.0 0.0 0.4 0.0 0.1 ± 0.2 to mix with people 0.0 0.0 2.1 0.0 0.0 0.4 ± 0.8 to relax 0.0 0.0 4.4 0.0 0.0 0.9 ±1.8 simelane.indd 96 2006/10/15 10:59:19 pm issn 0075-6458 97 koedoe 49/2 (2006) animals (23 %). demand on the resources was low, with most interviewees indicating that they do not support the idea of harvesting resources from the parks. this was expressed more by those people in parks with distant urban communities (krnp, gghnp and mznp) than those with neighbouring rural communities (aenp and vnp). discussion the study revealed that, in aenp, mznp, vnp, gghnp and krnp, the communities have limited understanding of the resources that occur within these parks. it also showed that these communities are keen to support conservation of natural resources. their limited understanding of resources is understandable because, in south africa, most conservation areas are surrounded by fences and communities have, for some time, been excluded from either harvesting or accessing resources from the conservation areas. this physical exclusion has thus restricted their understanding of resources, which has in turn limited their understanding of biodiversity within these areas. the study has also revealed that two types of communities occur around the national parks of south africa, namely neighbouring and distant communities. as indicated by the study, the distant communities are more urban in character than the neighbouring communities (table 1). based on their levels of income, lifestyles and age group composition, the expectations of these communities from a national park may differ widely. for example, communities with low incomes, fewer educated people and a high level of unemployment (e.g. vnp and aenp) largely expect sanparks to provide them with immediate solutions to current problems, including creation of job opportunities and access to natural resources. on the other hand, communities with bettereducated people and higher incomes (e.g. krnp and gghnp) expect sanparks to provide them with recreational, pleasure and learning opportunities (table 8). the recognition of these differing expectations, as identified in the study, is critical for the table 9 reasons provided by respondents for not visiting the park, expressed as the percentage of respondents who had not visited the park for the specified reason. (park codes as for table 1) reasons provided for not visiting the park aenp (n = 178) mznp (n = 162) gghnp (n = 240) krnp (n = 96) vnp (n = 400) mean (± sd) % across parks financial problem 27.1 35.9 45.4 4.1 16.4 25.8 ± 14.5 do not have transport 19.8 35.2 3.3 30.2 24.3 22.6 ± 10.9 like to go there but do not have time 5.1 11.1 9.6 10.4 17.9 10.8 ± 4.1 never had an opportunity, now too old to go there 0.0 4.3 0.3 10.4 8.9 4.8 ± 4.3 poor state of health 0.0 0.0 0.0 0.0 10.4 2.1 ± 4.2 not interested 14.7 7.4 20.4 18.7 12.8 14.8 ± 4.6 restricted by apartheid 31.3 1.2 1.3 6.3 0.4 8.1 ± 11.8 not aware of such possibility 1.4 3.7 2.1 5.2 0.7 2.6 ± 1.6 not well informed about the park 0.0 0.0 10.8 12.5 5.0 5.7 ± 5.3 negative attitudes from the staff 3.2 0.0 0.0 1.1 0.0 0.9 ± 1.3 never thought of visiting 0.0 0.0 5.0 0.0 0.0 1.0 ± 2.0 afraid of animals 0.9 1.2 1.3 0.0 3.2 1.3 ± 1.1 poor relationship with the park 1.4 0.0 0.0 1.1 0.0 0.5 ± 0.6 too lazy to go there 0.0 0.0 0.4 0.0 0.0 0.1 ± 0.2 no reason 1.4 0.0 0.1 0.0 0.0 0.3 ± 0.6 simelane.indd 97 2006/10/15 10:59:19 pm koedoe 49/2 (2006) 98 issn 0075-6458 design of community-based projects. such projects need to be aligned with community expectations as guided by income, lifestyle and education level of the people for whom they are intended. the projects must also aim to bring social and economic prosperity to communities that are highly affected by conservation (e.g. rural neighbouring communities). through this study it has emerged that communities around national parks may be heterogeneous with people of different cultures, as is the case with the aenp, mznp and krnp (table 1). in such cases, attention needs to be given to addressing the wide range of expectations of culturally diverse communities (van wyk & rossouw 1992). thus, in order to give fair consideration to community development projects, both during formulation and implementation, sanparks needs to be aware of the differences in the use of natural resources as interpreted by the various cultural groups (brockett 1990; wells & brandon 1992; fourie 1991,1994). social ecologists (as referred to in sanparks) or community liaison officers (as referred to in other conservation agencies) need to establish an understanding of the differences among the communities and their diverse interpretations of the resources that are available in the parks (wells & brandon 1993). of great importance is that, if more than one cultural group exists around the conservation area, conservation agencies need to ensure that they distribute benefits equally among all cultural groups (mcneely 1988; haynes 1998). for example, when establishing development projects aimed at exploiting identified development opportunities, community expectations as motivated by lifestyles should be regarded as a foundation for project development (hales 1989; brown & wyckoff-baird 1995; sibanda & omwega 1996). this may be attained through ensuring that, as far as possible, the projects are established in line with the lifestyles, levels of education and incomes of the communities involved (mcneely 1988; brown & wyckoffbaird 1995). in so doing, care should be taken to address the expectations of all cultural groups. giving more attention to the needs and demands of one cultural group may hamper the chances of establishing the balanced community support that is essential to extend conservation objectives by other groups to communities (byers 1996). this may then thwart the chances of achieving meaningful input from all communities around a conservation area (hackel 1999). the study has indicated that sanparks is battling to attain this balance, and that some communities have not yet realised the benefits to be derived from development projects (table 7). what conservation agencies need to realise is that successful implementation of a community-based conservation strategy lies in the design of the development projects, so that they catch the attention of different cultures and age groups at all levels of the project development and implementation cycle (colchester 1994). such involvement creates a sense of project ownership among the communities (colchester 1994; little 1994). it also ensures that communities realise the benefits of the project at an early stage of development (little 1994; fabricius 1994). this results in the development of a longterm relationship between conservation areas and communities (hanekom & liebenberg 1994; little 1994). communities then start to appreciate the existence of the conservation areas and thus realise the objectives of the existence and purpose of conservation areas (hampicke 1994; engelbrecht & van der walt 1994; fabricius 1994). the study did not clearly show if sanparks was involving communities at all levels of project development but it did indicate that almost a quarter (averaged across all five studied parks) of respondents rated the relationships with the parks as poor. however, the high percentage of people who indicated their willingness to take part in the conservation of natural resources is encouraging. this analysis provides sanparks with an opportunity to implement participatory processes for the support of conservation of biodiversity in national parks. the high level of recognition by the communities of the simelane.indd 98 2006/10/15 10:59:19 pm issn 0075-6458 99 koedoe 49/2 (2006) economic benefits of national parks (table 6) reflects the awareness of the communities towards development opportunities provided by conservation. this allows sanparks to effectively implement strategies of sustainable utilisation of natural resources through options such as ecotourism (geach 1997). this is of particular importance as conservation provides a wide range of sustainable opportunities (cater & lowman 1994; geach 1997; weinberg 2000). these include the creation of sustainable job opportunities and the generation of economic activities while conserving biodiversity (kerley & boshoff 1997). as conservation is starting to gain preference as a form of land use that promotes the sustainable use of natural resources (kerley et al. 1995; ashely 1996), conservation agencies need to ensure that development opportunities that are associated with conservation are realised and utilised by the local communities (kiss 1990; colchester 1994). however, for communities to exploit all opportunities that are provided by conservation of biodiversity, it is necessary that they obtain relevant training, which may include business skills development (fourie 1994; weinberg 2000). although conservation areas are often regarded as associated with poor rural communities (kock 1995), it has here emerged that some communities around national parks are well serviced in terms of infrastructure. as described in this study, modern amenities such as tarred roads, electricity, piped tapwater and sewage systems are found in communities around aenp, mznp and krnp. these amenities provide opportunities for communities to participate in eco-tourism related business initiatives (dbsa 1998). tarred roads promote visits to local settlements by tourists and electricity increases the intensive production of arts and crafts that can be sold to tourists. the participation of communities in such economic activities will not only develop interest in conservation but may also create an awareness of development opportunities that are provided by conservation (wells et al. 1992). it should be noted that the relatively high level of infrastructure available to these communities may explain their relatively low reliance on natural resource use (e.g. firewood). this identifies an urgent need to investigate further the relationship between socio-economic development and reliance on traditional natural resources. in certain cases, this study has identified findings that are not in accord with other studies conducted in africa (bell 1987; beinart 1989; alcorn 1993; alien 1995; mcneely 1997; bolton 1997). this specifically applies to the issue of resource demand and extraction from the parks. for example, the demand for and use of fuel wood has emerged to be relatively low in comparison with similar studies conducted elsewhere in africa. while this serves as a point of departure of this study from other similar studies, it is worthy of note that four of the studied parks (i.e. krnp, vnp, mznp and gghnp) are located in arid, relatively unwooded areas where the use of fuel wood is not as common as in other parts of the country. the positive attitudes towards national parks shown here reflect a change in the attitudes of communities around national parks in south africa (hough 1988; newmark et al. 1993; fourie 1994; boonzaier 1996; weaver 2000; munnik & mhlope 2000; berger & ntiati 2000; hove 2000; jacobson 2000). it remains imperative, however, that conservation areas are seen to be promoting the conservation of nature and biodiversity (cater & lowman 1994). communities around conservation areas need to be made to understand and appreciate that conservation areas exist to conserve nature and biological diversity (alien 1995; alcorn 1995). thus, the development of relationships with communities should be geared at increasing conservation awareness rather than demand for resources within the conservation areas (preston et al. 1991; bookbinder et al. 1998; colchester 1994; bebbington & kopp 1998). references anc (african national congress). 1994. the reconstruction and development programme: a policy framework. cape town: abc printers. simelane.indd 99 2006/10/15 10:59:19 pm koedoe 49/2 (2006) 100 issn 0075-6458 anon. 1989. the politics of conservation in southern africa. journal of south african studies 15 (2): 1–249. arber, s. 1996. the research process. pp. 32–50. in: gilbert, n. (ed.). researching social life. london: sage. alcorn, j.b. 1993. indigenous peoples and conservation. conservation biology 7: 424–426. alcorn, j.b. 1995. ethnobotanical knowledge systems—a resource for meeting rural development goals. pp. 1–12. in: warren, d.m., l.j. slikkerveer & d. brokensha (eds). the cultural dimension of development: indigenous knowledge systems. london: intermediate technology publications. alien, e.k. 1995. people’s participation in the development and management of tiwai-sierra leon’s first game sanctuary. nature et fauna 113: 11–21. ashely, c. 1996. incentives affecting biodiversity conservation and sustainable use: the case of land use options in namibia. windhoek: directorate of environmental affairs. (dea research discussion paper 13: 1–21.) bell, r.h.v. 1982. the problem of traditional use of wildlife resources in malawi. paper presented to counseille internationale du chasse, monaco, june 1982. brandon, k.e. & m. wells. 1992. planning for people and parks: design dilemmas. world development 20 (4): 557–569. brockett, c. 1990. land power and poverty: agrarian transformation and political conflict in central america. boston, usa: unwin hyman. brown, m. & b.wyckoff-baird. 1995. designing integrated conservation and development projects. washington dc: biodiversity support programme (wwf). byers, b.a. 1996. understanding and influencing behaviours in conservation and natural resources management. washington dc: westend press. bolton, m. (ed.). 1997. conservation and the use of wildlife resources. new york: chapman & hall. bookbinder, m.p., e. dinerstein, a. rijal, h. cauley & a. rajouria. 1998. ecotourism’s support of biodiversity conservation. conservation biology 12 (6): 1399–1404. boonzaier, e. 1996. local responses to conservation in richtersveld national park, south africa. biodiversity and conservation 5: 307–314. berger, d. & p. ntiati. 2000. milking the wild herd – maasailand. pp. 75–102. in: weinberg, p. (ed.). once we were hunters. amsterdam: mets & schilt. bebbington, a. & a. kopp. 1998. networking and rural development through sustainable forest management frameworks for pluralistic approaches. unasylva 49: 11–18. bell, r.t. 1987. conservation with a human face: conflict and reconciliation in african land use planning. pp. 79–109. in: anderson, d & d. grove (eds.). conservation in africa: people, policies and practice. cambridge: cambridge university press. beinart, w. 1989. the politics of colonial conservation. journal of south african studies 15: 145–161. colchester, m. 1994. beyond “participation”: indigenous peoples, biological diversity conservation and protected area management. unasylva 186(47): 33–39. cater, e. & g. lowman (eds.). 1994. ecotourism —sustainable option. england: john wiley. cobbett, m. 1987. the land question in south africa: a preliminary assessment. south african journal of economics 55(1): 17–22. dbsa (development bank of southern africa) 1998. infrastructure: a foundation for development. development report, development bank of southern africa, midrand. engelbrecht, w. & p.t. van der walt. 1994. notes on the economic use of the kruger national park. koedoe 36 (2): 113 – 119. fourie, j. 1994. comments on national parks and future relations with neighbouring communities. koedoe 37: 123–136. fourie, j. 1991. the concept of life on the social role of conservation areas. koedoe 32: 157–165. fabricius, c. 1994. towards a decision analysis framework to integrate protected area management and community development. paper presented at saie symposium on land utilization, people & environment, 5/51994. port elizabeth. geach, b.g.s. 1997. the addo elephant national park as a model of sustainable land use through ecotourism. unpubl. msc thesis. university of port elizabeth, port elizabeth, south africa. greyling, t. & b.j. huntley (eds.). 1984. directory of southern african conservation areas. pretoria: council for scientific and industrial research, national scientific programmes unit. (south african national scientific programmes report; no. 98.) hough, j.l. 1988. obstacles to effective management of conflicts between national parks and surrounding communities in developing countries. environmental conservation 15: 129–136. hampicke, u. 1994. ethics and economics of conservation. biological conservation 67: 219–231. hackel, j.d. 1999. community conservation and the future of africa’s wildlife. conservation biology 13: 726 – 734. haynes, j. s. 1998. involving communities in managing protected areas: contrasting initiatives in nepal and britain. parks 8: 54–61. hales, d. 1989. changing concepts of national parks. pp. 139–144. in: western, d. & c. simelane.indd 100 2006/10/15 10:59:19 pm issn 0075-6458 101 koedoe 49/2 (2006) pearl, (eds.). conservation for the 21st century. new york: wildlife conservation international. hanekom, d. & l. liebenberg. 1994. utilization of national parks with special reference to the costs and benefits to communities. bulletin of the grassland society of southern africa 5: 25–36. hildgard, n., d. hedge, p. wolvekamp & s. reddy. 1998. same platform, different train—the politics of participation. unasylva 49:26–34. hove, c. 2000. zimbabwe—people and animals, a tense harmony. pp. 122–146. in: weinberg, p. (ed.). once we were hunters. amsterdam: mets & schilt. jacobsohn, m. 2000. namibia’s kunene region—a new vision unfolds. pp. 105–120. in: weinberg, p. (ed.). once we were hunters. amsterdam: mets & schilt. kerley, g.i.h, a.f. boshoff & m.h. knight. 1999. ecosystem integrity and sustainable land-use in the thicket biome, south africa. ecosystem health 5 (2):104–109. kerley, g.i.h. & a.f. boshoff. 1997. a proposal for a greater addo national park—a regional and national conservation and development opportunity. zoology department, university of port elizabeth. (terrestrial ecology research unit; internal report no. 17.) kiss, a. 1990. living with wildlife: wildlife resource management with local participation in africa. washington, dc: the world bank. (world bank technical paper no 130.) kiss, a. 1994. is community-based ecotourism a good use of biodiversity conservation funds? tree 19: 232–237. kock, r.a. 1995. wildlife utilization: use it or lose it—a kenyan perspective. biodiversity and conservation 4: 241–256. krog, a. 2000. it takes a lot of god to survive here —the richtersveld. pp. 55–92. in: weinberg, p. (ed.). once we were hunters. amsterdam: mets & schilt. little, p. 1994. the link between local participation and improved conservation: review of issues and experiences. pp. 105–117. in: d. western & r. wright (eds.). natural connections: perspectives in community-based conservation. washington dc: island press. ledger, j. 1998. south africa’s protected areas under siege. endangered wildlife 30: 4–9. mcneely, j.a. 1997. conservation and the future: trends and options towards the year 2025. gland, switzerland: iucn. munnik, v & g. mhlope. 2000. uneasy paradise —a journey through maputaland. pp. 32–52. in: weinberg, p. (ed.). once we were hunters. amsterdam: mets & schilt. mcneely, j.a. 1988. economics and biological diversity: developing and using economic incentives to conserve biological resources. gland, switzerland: iucn. mohr, p. & l. fourie. 2003. economics for south african students. pretoria: van schaik. newmark, d.w., n.l. leonard, h.l. sariko & d.m. gamasa. 1993. conservation attitudes of local people living adjacent to five protected areas in tanzania. biological conservation 63: 177–183. procter, m. 1996. measuring attitudes. pp. 116– 134. in: gilbert, n. (ed.). researching social life. london: sage. preston, g.r., r.f. fuggle & e. siegfried. 1991. environmental education in south african nature reserves: promoting desirable resource utilization practices. south african journal of science 87: 544 –547. rothley, k.d. 1999. designing biosphere networks to satisfy multiple conflicting demands. ecological applications 9: 741–750. richards, m. 1996. protected areas, people and incentives in search for sustainable forest conservation in honduras. environmental conservation 23 (3): 207–217. ramphele, m. (ed.). 1991. restoring the land: environment and change in post apartheid south africa. london: parios. sachs, a. 1990. protecting human rights in a new south africa. cape town: oxford university press. sanparks (south african national parks). 1995. national parks board strategic plan, final consolidation of management input. kimberley: south african national parks. sanparks (south african national parks). 1997. draft policy on sustainable use of natural resources in the national parks. pretoria: south african national parks. sanparks (south african national parks). 1998. visions of change—social ecology and south african national parks. pretoria: south african national parks. simmonds, m. 1998. changing conservation aims —who will represent wildlife? oryx 32 (2): 82–83. sibanda, b.m.c. 1995. wildlife conservation in kenya: wildlife or local communities at the crossroads. environmental policy and practice 5(1): 22–24. sibanda, b.m. & a.k. omwenga. 1996. some reflections on conservation, sustainable development and equitable sharing of benefits from wildlife in africa: the case of kenya and zimbabwe. south african journal of wildlife research 26 (4): 175–181. starr, c. & r. taggart. 1987. biology—the unity and diversity of life. belmont, california: wadsworth. simelane.indd 101 2006/10/15 10:59:19 pm koedoe 49/2 (2006) 102 issn 0075-6458 steele, n.a. 1991. natural resource utilization as conservation approach. koedoe 34: 93–95. van wyk, a. & a. rossouw. 1992. a socio-economic survey of nomathamsanqa in the addo district. research report 44. institute for planning research, university of port elizabeth. weaver, t. 2000. kalahari, namibia and botswana— no respite for the san. pp. 12–30. in: weinberg, p. (ed.). once we were hunters. amsterdam: mets & schilt. wells, m.p. & k.e. brandon. 1993. the principles and practice of buffer zones and local participation in biodiversity conservation. ambio 22: 157–162. wells, m.p., k.e. brandon & l. hannah. 1992. people and parks, linking protected area management with local people. washington dc: international bank for reconstruction and development. weinberg, p. (ed.). 2000. once we were hunters. amsterdam: mets & schilt. western, d. & w.r. henry. 1979. economics and conservation in third world national parks. bioscience 29(7): 414–418. zimbabwe trust. 1990. people, wildlife and natural resources: the campfire approach to rural development in zimbabwe. harare, zimbabwe: zimbabwe trust. zar, j. h. 1984. biostatistical analysis. new jersey: prentice hall. simelane.indd 102 2006/10/15 10:59:19 pm article information authors: izak p.j. smit1 edward s. riddell2 carola cullum3 robin petersen1 affiliations: 1scientific services, south african national parks, skukuza, south africa2school of bioresources engineering and environmental hydrology, university of kwazulu-natal, south africa 3centre for water in the environment, university of the witwatersrand, south africa correspondence to: izak smit postal address: private bag x402, skukuza 1350, south africa dates: received: 24 aug. 2012 accepted: 14 jan. 2013 published: 12 apr. 2013 how to cite this article: smit, i.p.j., riddell, e.s., cullum, c. & petersen, r., 2013, ‘kruger national park research supersites: establishing long-term research sites for cross-disciplinary, multiscaled learning’, koedoe 55(1), art. #1107, 7 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1107 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/ koedoe.v55i1.1107-1. copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. kruger national park research supersites: establishing long-term research sites for cross-disciplinary, multiscaled learning in this original research... open access • abstract • introduction    • rationale for research supersites • research method and design    • supersite selection criteria    • supersite selection process    • supersite management • results    • supersite descriptions       • southern granites: stevenson-hamilton supersite       • northern granites: ngwenyeni supersite       • southern basalts: nhlowa supersite       • northern basalts: mooiplaas supersite • discussion    • supersite datasets       • supersite data availability • conclusion • acknowledgments    • competing interests    • authors' contributions • references abstract top ↑ researchers interested in studying the effects of fire or herbivory in the kruger national park (knp) often focus their research activities on the experimental burn plots or herbivore exclosure camps, respectively. these are manipulated sites that apply treatments, for example annual fires or total exclusion of fire and herbivores. however, many projects aim to study or monitor patterns and processes emerging under non-manipulated conditions, typically at sites with contrasting geologies and rainfall. yet, these sites are usually selected in a haphazard and uncoordinated manner for different projects and, as a consequence, it is often not possible to integrate datasets and knowledge. an alternative to the ever-increasing number of unrelated sites scattered across the park are the ‘knp research supersites’ which have been earmarked to geographically focus future research effort, acting as data-rich, long-term sites for monitoring and research. in this paper, we introduced the four recently established knp research supersites, which cover the rainfall gradient and geological contrast of the knp, presenting their rationale, selection criteria and location, along with existing datasets that describe their herbaceous biomass, woody cover, phenology, fire history, levels of herbivory. additional site-specific datasets, which are already available, or which are in preparation, were outlined together with details for assessing these open-source datasets online. conservation implications: the knp research supersites will become increasingly used for research, monitoring and remote-sensing calibration and ground-truthing purposes. scientists are encouraged to gain from, and contribute towards, these sites, which will facilitate long-term data collection, data-sharing and co-learning and, ultimately, lead to a more integrated, multiscaled and multitemporal understanding of savannahs. introduction top ↑ rationale for research supersites the kruger national park (knp) has become an important research location for scientists from across the world studying savannah systems. close to 800 projects have been officially registered in the knp since the 1950s and project numbers have rapidly increased in recent years, with 154 new projects being registered between january 2010 and june 2012 (r. scholtz [south african national parks] pers. comm., 14 august 2012). various reasons are proposed for the popularity of the knp as an important ‘outdoor laboratory’, including that the knp:• has a long history of management being informed by science, which attracts researchers wanting to study and contribute to applied conservation problems. • offers untransformed or largely non-manipulated savannah landscapes, yet with good access and logistical support provided through a dedicated department (scientific services). • has accumulated an excellent database of historical data, including spatially explicit long-term vegetation, fire and herbivore datasets, as well as an ever-expanding base of published research. • supports a well-established and growing research community operating mostly in a collaborative manner, as witnessed though the success of the annual savanna science network meeting, which has been running since 2002 and attracts close to 300 delegates each year. • maintains long-term manipulated and well-studied research sites in the park. long-term, manipulated research sites include the experimental burn plots (ebps), where fire season and frequency have been manipulated since 1954 (biggs et al. 2003; van der schijff 1958). the ebps form one of the oldest and largest uninterrupted fire manipulation experiments in africa and, by june 2012, 21 datasets, 15 theses and 50 peer-reviewed publications, including an overview paper (van wilgen, govender & biggs 2007) have emanated from this experiment (n. govender [sanparks] pers. comm., 20 june 2012). the rare antelope enclosures and herbivore exclosures which manipulate levels of herbivory are additional manipulation sites and are attracting more researchers as the role of herbivores as drivers in african savannah systems is increasingly appreciated (for studies conducted on the knp herbivore enclosures and/or exclosures see e.g. asner & levick 2012; asner et al. 2009; jacobs & naiman 2008; jonsson et al. 2010; levick et al. 2009). there are no equivalent long-term, data-rich sites available for research studies that focus on non-manipulated areas which experience ’natural’ levels of fire, herbivory, et cetera. as a result, studies involving non-manipulated conditions are located haphazardly across the remaining two million hectares of the park. these ad hoc site selections limit the opportunity of linking datasets emanating from different studies over time. the advantages of incentivising a geographic focus for future studies are numerous. previous experience shows that ’data beget data’ – the more data that are generated for a specific location, the more that site is further studied and the more additional data get collected, especially when these datasets are freely available within the scientific community. this results in data-rich sites where the level of understanding of pattern and process can deepen as additional and complimentary datasets are collected. the long-term ecological research (lter) sites are an example of an existing network of data-rich sites in the usa, where baseline data, data sharing and a research-enabling environment act as catalysts for data sharing and collaboration of over 1800 scientists (see http://www.lter.edu). whilst the lter sites are also largely experimental in design, they have revealed that comprehensive understanding of ecological processes can only be achieved by avoiding the temptation to interpret geographically and biologically disjunct datasets (knapp et al. 2012). to this end, the knp research supersites follow a model similar to that proposed for the south african environmental observation network (saeon) for the long-term monitoring of biomes, in which ecological observations are gathered, collaborative research is facilitated and spatially integrated data are archived (van jaarsveld et al. 2007). however, the supersites are applied at a more local scale, with a focus on the dominant rainfall and geomorphologically defined land systems in the knp, and aim to facilitate integrated research rather than monitoring. herein, we provide a description on the supersites and datasets already available for these supersites, with the aim of encouraging researchers wanting to establish new research projects in the knp to choose these sites. although different research projects have diverse requirements for site selection based on the specific aims and scale of a project, there are often multiple potential sites that will fit these criteria. as a result, the final site selection is often largely based on choosing from potential sites that are conveniently located and/or have existing data available. furthermore, the criteria most often used for selecting sites in the knp are simply geology (covering the two main geologies of the park, namely nutrient-rich basalt in the east versus nutrient-poor granite in the west) and rainfall (a gradient of generally increasing rainfall from north to south) (venter, scholes & eckhardt 2003) and researchers are often amenable to working at any specific location within these broad zones. research method and design top ↑ supersite selection criteria a group of south african nation parks (sanparks) scientists and external research collaborators agreed that supersites should be delineated using catchment boundaries, rather than by an ecologically meaningless shape. the spatial and temporal availability of water constrains many ecological processes, ranging from fire to flood to biochemistry to resource availability for animals (jenerette et al. 2012; noy-meir 1973). thus the patterns of geology, climate, morphology, soils and vegetation that largely control water fluxes across and through the landscape form a biophysical template upon which many ecological processes are played out (caylor, d’odorico & rodriguez-iturb 2006). in semi-arid climates, such as that found in the knp, these patterns are largely coupled. for example, catenas of soils and associated vegetation are often found at certain hillslope positions, forming sequences that reflect patterns of water availability within a sub-catchment (milne 1935). furthermore, these catenal sequences are repeated throughout areas with similar climate, geology and morphology (‘physiographic zones’ after cullum & rogers 2011, or ‘land systems’ after venter 1990). using (sub-)catchment boundaries and catenal elements to delineate research and monitoring sites within distinct physiographic zones therefore allows sites to be placed within the ecohydrological contexts that control so many ecological processes. the abovementioned group of scientists also agreed on the following more specific selection criteria; that is, supersites should: • between themselves include contrasts of the two main abiotic variables driving much of the larger scale variation in the park, namely, geology (granite and basalt) and rainfall (higher rainfall in the south versus lower rainfall in the north). • be large enough to contain at least one third-order catchment situated entirely within a single geology, allowing ecological patterns to be studied at the three scales associated with first-order, second-order and third-order catchments. • contain the hillslope vegetation and soil patterns (catenal sequences) that commonly occur in the local land system. • be easily accessible from as many sides as possible by all-weather roads. • be close to research camps and facilities. • be outside of areas demarcated as ‘wilderness’ or ‘remote’ in the knp zoning plan, in order to allow installation of instrumentation. although the sites have been selected to be ‘typical’ of the main geoclimatic settings in the park, care must be taken to account for local heterogeneity when extrapolating data to other locations (see cullum & rogers 2012). at each supersite, at least one third-order catchment has been furnished with hydrological and meteorological instruments for intensive study. however, because a single third-order catchment may be too small or may have sub-optimal access for certain types of studies, the supersites were often delineated to be larger than the catchment (in which case, roads mostly form the boundary). whenever possible, studies should aim to work in the focal sub-catchments, as many patterns and processes are driven by water fluxes associated with catchments and landscape position (see above) and, as such, the sub-catchments form the key focal areas within the supersites. supersite selection process potential sites meeting the above selection criteria were initially identified within a geographic information system environment, overlaying various data layers with high resolution spot 5 satellite imagery as the backdrop. this was followed by a helicopter survey over potential sites in march 2011 and subsequent field visits to allow further aerial and ground-based evaluation of suitability. through this process the most suitable sites emerged and consensus was reached as to which sites were to be tabled for review and approval by the wider scientific and management community involved in the knp. the four proposed sites identified by the process above were discussed with the relevant managers, after which they were also discussed at an open-invitation workshop at the 9th savanna science network meeting, skukuza, kruger national park (march 2011). the workshop allowed a large proportion of the scientific community with research interests and experience in the park to participate in the site review process. the workshop was attended by about 30 scientists, including prominent researchers with a long history of working in the knp. the reaction was overwhelmingly positive regarding the concept of research supersites and, after discussing some potential alternative sites, the originally proposed sites were accepted without objection. in order to maintain the geographic focus of the initiative, additional suggested sites (e.g. sites on gabbros, sites next to perennial rivers, etc.) could be considered at a later stage and even then only with reduced instrumentation and support. supersite management as the supersites were conceived to allow study of non-manipulated conditions, they will be managed in accordance with the rest of the park. the supersites are therefore not available for large-scale manipulative studies (e.g. fertilisation, irrigation, fire experiments, etc.), but instrumentation of the site will be encouraged if and where appropriate. as with all research projects conducted within national parks, projects intended for the supersites will first need to be approved and registered by sanparks. the sanparks committees evaluating the projects will also evaluate which instrumentation is appropriate for inclusion on the sites. results top ↑ supersite descriptions venter (1990) characterised the knp in terms of 11 land systems, defined by similarities in landforms and the hillslope sequences of soils and dominant woody and herbaceous vegetation at a scale of approximately 1:1 000 000. these land systems were further subdivided by venter (1990) into 56 land types, mapped at a scale of approximately 1:250 000. the supersites occur within the four largest land systems (which account for 80% of the total park area), namely, skukuza (southern granites, with relatively high rainfall), phalaborwa (northern granites, with relatively low rainfall), satara (southern basalts, with relatively high rainfall) and letaba (northern basalts, with relatively low rainfall) (figure 1). venter’s (1990) descriptions of the land types within which the supersites fall are summarised in the online appendix tables 1 and 2, together with their hydrogeological characteristics (online appendix table 3). figure 1: the locations of the four supersites within the kruger national park, in relation to the designated land types and land systems. the characteristics of the supersites, including long-term average rainfall, herbaceous biomass and woody cover are described in table 1 (also see online appendix table 4 for annual rainfall history since 1941). a summary of the long-term fire history is provided in table 2 and historic levels of herbivory in figure 2 (also see online appendix table 5 for density of specific herbivore species). figure 3 outlines the average yearly phenological response of the supersites, based on the normalised difference vegetation index time series data, derived using moderate resolution imaging spectroradiometer (modis) imagery collected between 2000 and 2012. aerial and ground photographs of the sites are included in the online appendix figures 1–8 (see also online appendix table 6 for details of historical aerial photography taken of the supersites between 1940 and 2001). the broader park-wide context within which the supersites occur appear as maps in the online appendix figures 9–18 (e.g. depicting supersites in relation to camps and tourist roads, average herbaceous biomass, woody cover, fire return period, dry-season herbivore distribution and elephant distribution). the online appendix figures 19–22 provide the locations (and site identification numbers) for sites on or within 5 km of the supersites for the following datasets, (1) soil and vegetation (herbaceous and woody) data collected by venter (1990) and (2) vegetation condition assessment surveys carried out by the knp rangers since 1989. table 1: average long-term rainfall, herbaceous biomass and woody cover for the kruger national park research supersites. table 2: average long-term (1941–2006) fire characteristics of kruger national park research supersites. figure 2: average herbivore biomass on supersites, based on aerial census data collected between 1987 and 1993. the southern basalt supersite experiences the highest levels of herbivory and both basalt sites support a large density of grazers. figure 3: average phenology of supersites based on the normalised difference vegetation index, derived using moderate resolution imaging spectroradiometer (modis) imagery. time series are based on average index calculated for nine pixels located in the centre of each supersite (averaged on anniversary dates between february 2000 and march 2012). southern granites: stevenson-hamilton supersite the southern granite supersite sits within the renosterkoppies land type, which is a transitional zone between the land types associated with the perennial sabie and crocodile river catchments (figure 4). the landscape is finely dissected, with a high stream density. the woody vegetation of the sandy crests is dominated by combretum apiculatum and combretum zeyheri. there is often a distinct seepline between the crest and the midslope, where terminalia sericea is conspicuous. fine leaved woody species such as acacia nilotica dominate the more clayey midslopes and footslopes. duplex, sodic soils are often found on the footslopes, characterised by the presence of euclea divinorium. concave stream channels generally have gravel beds. figure 4: stevenson-hamilton supersite on southern granites (coordinates of h1-1/s114 intersection: 31.589e; 25.046s). northern granites: ngwenyeni supersite the northern granite supersite falls within the malopeni land type, part of the phalaborwa land system (figure 5). it is an intensely migmatised zone between the gneiss to the south and greenstone to the north. it is characterised by many small headwater tributaries of the letaba river. combretum apiculatum dominates on the shallow soils of the crests, whilst colophospermum mopane is associated with the midslopes. footslopes are dominated by c. mopane and some acacia nigrescens. figure 5: ngwenyeni supersite on northern granites (coordinates of h14/s131 intersection: 31.256e; 23.863s). southern basalts: nhlowa supersite the southern basalt supersite occurs within the satara land type, which is characterised by shallow to moderately deep red olivine-poor clays (figure 6). these nutrient-rich soils support a high density of herbivores, with dense grass cover providing a reliable source of grazing. it is very flat with open tree savannah dominated by sclerocarya birrea and a. nigrescens. stream density is low, with wide grassy valley bottoms in headwater areas. figure 6: nhlowa supersite on southern basalts (coordinates of s137/s28 intersection: 31.955e; 25.213s). northern basalts: mooiplaas supersite the northern basalt supersite occurs within the mooiplaas land type and is characterised by dark olivine-rich soils (figure 7). it is very flat, with a very low stream density, shallow grassy vleis and large alluvial deposits along the major rivers. it is dominated by moderate to dense stands of c. mopane. dichrostachys cinerea is a prominent shrub. figure 7: mooiplaas supersite on northern basalts (coordinates of h1-6/s49 intersection: 31.420e; 23.534s). discussion top ↑ supersite datasets presently, a concerted effort is directed at collecting baseline datasets characterising the supersites. the existing datasets to date are summarised in table 3. table 3: existing baseline datasets available for the kruger national park, including those shortly available through projects registered with south african national parks. supersite data availability the datasets described in table 3 are available (or will become available shortly as they are finalised) from the sanparks data repository (http://www.dataknp.sanparks.org). it is expected that researchers using these datasets will also contribute additional data to the wider research community as they collect or derive new datasets for these sites. new data will be added to the sanparks data repository as they become available. in cases where the researchers want to publish the data before making it publicly available, a lead-time of up to 3 years will be granted. during this period, only an abstract of the data and the contact details of the relevant researcher will be visible on the repository; however, the data will be made available once the data owner provides permission, or at the latest after 3 years from the date of collection, as normally specified by sanparks.titles of all the supersite datasets will appear when searching the data repository using the term ’supersite’ (with additional keywords focusing the search to be more specific). once the list of dataset titles appears during a search, the required data and metadata can be downloaded directly. researchers downloading supersite data from the data repository will be required to acknowledge the data collectors and/or reference the publication associated with the specific dataset, as indicated in the metadata of each dataset. in a few exceptional cases where the data may be sensitive and/or where other restrictions (e.g. copyright) may apply, the users will only see the abstract and contact details of the person from which to request the data. all the data will be provided on an ‘as is’ basis and sanparks and the data collectors will not be responsible for any data errors (however, feedback on any data errors is encouraged). conclusion top ↑ the aim of the recent establishment of four research supersites in the knp and the provision of spatially explicit baseline datasets for these locations is to stimulate a geographic focus for research and monitoring studies which currently happen on sites that are selected in an uncoordinated and haphazard manner. the supersites cover the two main abiotic variables that drive the large-scale variation in the park, namely, the rainfall gradient and the geological contrast, and contain entire lower order catchments. it is believed that these sites will result in a long-term, data-rich environment, which will provide the opportunity for better integration of datasets and disciplines. it is anticipated that this will lead to a deeper understanding of pattern and process on various spatio-temporal scales. the wealth of freely available spatially explicit datasets presently available or in process of finalisation will act as catalysts to attract further research projects to these sites and to stimulate the collection of additional datasets and knowledge generation. acknowledgments top ↑ we want to acknowledge all managers and scientists involved in the discussions and workshops during which the supersites were discussed and finalised. special thanks also to gabriela bucini who co-organised a remote-sensing woody cover mapping workshop together with the primary author in 2009, which was useful in developing the supersite concept. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions i.p.j.s. (sanparks) was the project leader and, together with e.s.r. (university of kwazulu-natal) and c.c. (university of the witwatersrand), was closely involved in the supersite conceptualisation and delineation. i.p.j.s. prepared the manuscript with contributions from e.s.r. and c.c, whilst i.p.j.s. and c.c. created all the maps. r.p. (sanparks) coordinated many of the baseline studies presently occurring on the supersites and also contributed towards the manuscript. references top ↑ asner, g.p., levick, s.r., kennedy-bowdoin, t., knapp, d.e., emerson, r., jacobson, j. et al., 2009, ‘large-scale impacts of herbivores on the structural diversity of african savannas’, proceedings of the national academy of sciences of the united states of america 106(12), 4947–4952. http://dx.doi.org/10.1073/pnas.0810637106, pmid:19258457, pmcid:2650504asner, g.p. & levick, s.r., 2012, ‘landscape-scale effects of herbivores on treefall in african savanna’, ecology letters 15(11), 1211–1217. biggs, r., biggs, h.c., dunne, t.t., govender, n. & potgieter, a.l.f., 2003, ‘experimental burn plot trial in the kruger national park: history, experimental design and suggestions for data analysis’, koedoe 46(1), 1–15. bucini, g., saatchi, s., hannan, n.p., boone, r.b. & smit, i.p.j., 2009, ‘woody cover and heterogeneity in the savannas of the kruger national park, south africa’, in proceedings from the 2009 ieee international geoscience and remote sensing symposium iv, cape town, south africa, july 12–17, 2009, pp. 334–337. bucini, g., hanan, n.p., boone, r.b., smit, i.p.j., saatchi, s., lefsky, m.a. et al., 2010, ‘woody fractional cover in kruger national park, south africa: remote-sensing-based maps and ecological insights’, in m.j. hill & n.p. hanan (eds.), ecosystem function in savannas: measurement and modeling at landscape to global scales, pp. 219–237, crc/taylor and francis, boca raton. caylor, k.k., d’odorico, p. & rodriguez-iturbe, i., 2006, ‘on the ecohydrology of structurally heterogeneous semiarid landscapes’, water resources research 42(7), w07424. http://dx.doi.org/10.1029/2005wr004683 cullum, c. & rogers, k.h., 2011, a framework for the classification of drainage networks in savanna landscapes, wrc report tt498/11, water research commission, pretoria. cullum, c. & rogers, k.h., 2012, using landscape classification within complex, heterogeneous systems, presented at the 10th savanna science networking meeting, skukuza, 05–09 march, viewed 19 july 2012, from http://www.sanparks.org/assets/docs/parks_kruger/conservation/scientific/noticeboard/science_network_meeting_2012/5-15-cullum.pdf du toit, w.h., 1998, geohidrologie van die nasionale krugerwildtuin gebaseer op die evaluering van bestaande boorgatinligting [geohydrology of the kruger national park based on the evaluation of existing borehole information], vol. 1, directorate geohydrology internal report gh 3798, department of water affairs and forestry, pretoria. jacobs, s.m. & naiman, r.j., 2008, ‘large african herbivores decrease herbaceous plant biomass while increasing plant species richness in a semi-arid savanna toposequence’, journal of arid environments 72(6), 891–903. http://dx.doi.org/10.1016/j.jaridenv.2007.11.015 jenerette, g.d., barron-gafford, g.a., guswa, a.j., mcdonnell, j.j. & villegas, j.c., 2012, ‘organization of complexity in water limited ecohydrology’, ecohydrology 5, 184–199. jonsson, m., bell, d., hjalten, j., rooke, t. & scogings, p.f., 2010, ‘do mammalian herbivores influence invertebrate communities via changes in the vegetation? results from a preliminary survey in kruger national park, south africa’, african journal of range and forage science 27(1), 39–44. knapp, k.a., smith, m.d., hobbie, s.e., collins, s.l., fahey, t.j., hansen, g.j.a. et al., 2012, ‘past, present, and future roles of long-term experiments in the lter network’, bioscience 62(4), 377–389. levick, s.r., asner, g.p., kennedy-bowdoin, t. & knapp, d.e., 2009, ‘the relative influence of fire and herbivory on savanna three-dimensional vegetation structure’, biological conservation 142(8), 1693–1700. http://dx.doi.org/10.1016/j.biocon.2009.03.004 milne, g., 1935, ‘some suggested units for classification and mapping, particularly for east african soils’, soil research 4, 183–198. noy-meir, i., 1973, ‘desert ecosystems: environment and producers’, annual review of ecology and systematics 4, 25–51. schulze, r.e., maharaj, m., lynch, s.d., howe, b.j. & melville-thomas, b., 1997, south african atlas of agrohydrology and -climatology, wrc report tt82/96, water research commission, pretoria. smit, i.p.j., 2007, ‘artificial surface-water provision in a semi-arid savanna. a spatio-temporal analysis of herbivore distribution patterns in relation to artificial waterholes under different habitat, rainfall and management scenarios in the kruger national park, south africa’, phd thesis, department of geography, university of cambridge. smit, i.p.j., 2011, ‘resources driving landscape-scale distribution patterns of grazers in an african savanna’, ecography 34(1), 67–74. http://dx.doi.org/10.1111/j.1600-0587.2010.06029.x smit, i.p.j. & ferreira, s.m., 2010, ‘management intervention affects river-bound spatial dynamics of elephants’, biological conservation 143(9), 2172–2181. http://dx.doi.org/10.1016/j.biocon.2010.06.001 smit, i.p.j., grant, c.c. & devereux, b.j., 2007, ‘do artificial waterholes influence the way herbivores use the landscape? herbivore distribution patterns around rivers and artificial surface water sources in a large african savanna park’, biological conservation 136(1), 85–99. http://dx.doi.org/10.1016/j.biocon.2006.11.009 smit, i.p.j., smit, c.f., govender, n., van der linde, m. & macfadyen, s., 2013, ’rainfall, geology and landscape position generate large-scale spatiotemporal fire pattern heterogeneity in an african savanna’, ecography 36(4), 447–459. http://dx.doi.org/10.1111/j.1600-0587.2012.07555.x van der schijff, h.p., 1958, ‘inleidende verslag oor veldbrandnavorsing in die nasionale krugerwildtuin [introductory report on veld burning research in the kruger national park]’, koedoe 1(1), 60–93. van jaarsveld, a.s., pauw, j.c., mundree, s., mecenero, s., coetzee, b.w.t. & alard, g.f., 2007, ‘south african environmental observation network: vision, design and status’, south african journal of science 103, 289–294. van wilgen, b.w., govender, n. & biggs, h.c., 2007, ‘the contribution of fire research to fire management: a critical review of a long-term experiment in the kruger national park, south africa’, international journal of wildland fire 16(5), 519–530. http://dx.doi.org/10.1071/wf06115 venter, f.j., 1990, ‘a classification for land management planning in the kruger national park’, phd thesis, department of geography, university of south africa. venter, f.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation pattern’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83–129, island press, washington, dc. abstract introduction materials and methods results discussion conclusion acknowledgements references about the author(s) zoliswa n. nhleko department of zoology and entomology, rhodes university, south africa dan m. parker department of zoology and entomology, rhodes university, south africa school of biology and environmental sciences, university of mpumalanga, south africa dave j. druce ezemvelo kzn wildlife, kwazulu-natal, south africa school of life sciences, university of kwazulu-natal, south africa citation nhleko, z.n., parker, d.m. & druce, d.j., 2017, ‘the reproductive success of black rhinoceroses in the hluhluwe–imfolozi park, kwazulu-natal, south africa’, koedoe 59(1), a1386. https://doi.org/10.4102/koedoe.v59i1.1386 original research the reproductive success of black rhinoceroses in the hluhluwe–imfolozi park, kwazulu-natal, south africa zoliswa n. nhleko, dan m. parker, dave j. druce received: 17 feb. 2016; accepted: 10 july 2017; published: 15 nov. 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract black rhinoceroses (diceros bicornis) are endangered and the southern-central sub-species (diceros bicornis minor) is considered critically endangered. we assessed the reproductive life-histories of black rhinoceroses in hluhluwe–imfolozi park (hip), kwazulu-natal, south africa, to determine whether this historically important donor sub-population was meeting regional reproductive targets. detailed life-history information for known individuals (n = 79–120) was used to investigate reproductive parameters between 1998 and 2013. mean age at sexual maturity was 12 years, which exceeded a target period of 7 years and 5 months. the mean inter-calving interval was 3 years and 8 months – 8 months longer than the recommended 3 years. the poor population performance of the hip black rhinoceroses could be a result of poor habitat quality, poor animal condition, females losing their first calves, predation of calves or a negative social effect of annual live-harvesting of the population. however, we believe that the estimated ecological carrying capacity of black rhinoceroses at hip (a figure used to ascertain whether the population can be harvested at all) may be incorrect, leading to the poor reproductive performance. we recommend that the accuracy of the ecological carrying capacity estimate be assessed as a matter of urgency and that a moratorium be placed on the live-harvesting of individuals until the estimate has been refined. conservation implications: our results provide key data which can be used to refine black rhinoceros breeding targets in south africa and the region more broadly. introduction globally, the conservation status of the southern-central black rhinoceros (diceros bicornis minor) is critically endangered as a result of a population decrease of approximately 80% over the last three generations (international union for conservation of nature [iucn] 2012). this decrease has come about mainly because of poaching in the 1970’s and 1980’s, and the recent escalation in rhinoceros poaching (iucn 2012). south africa is the most important range state for this sub-species of black rhinoceros and, by the end of 2010, contained 1684 individuals (iucn 2012). the kwazulu-natal province, in the eastern part of south africa, was home to approximately 480 (29%) individuals of this sub-species in 2012 (iucn 2012). in 2011, the south african government produced a biodiversity management plan (bmp) for the black rhinoceros (diceros bicornis) in which two main goals were outlined. with relevance to the southern-central black rhinoceros (hereafter black rhino), these were (1) to achieve a meta-population growth rate of at least 5% per annum and (2) to increase the population to at least 2800 individuals by the end of 2020. these targets were to be achieved through sustainable meta-population growth by annual live-harvesting and translocating 5% of individuals from certain sub-populations to populate additional sites (goodman 2013). the expansion programme was dependent on the intensive monitoring of all sub-populations, including assessments of annual population sizes, population growth rates, fertility rates and mortality rates (goodman 2013). in addition to the national bmp, there is a provincial strategy for the management of black rhino in kwazulu-natal (conway & goodman 2013). the provincial management document describes the overall strategy for the management and recovery of the black rhino populations on state, communal and private land in kwazulu-natal for the period from 2013 to 2018 (conway & goodman 2013). strategic goals are to increase the number of black rhinos in the province to at least 740 individuals by the end of 2022, to achieve an average of 0.25 births per adult female per annum (between 2013 and 2022), and to achieve a growth rate of 5% per annum over any 3-year cycle (conway & goodman 2013). one of the actions required to achieve these goals is to live-harvest populations that are greater than 70% of the estimated ecological carrying capacity, at a minimum rate of 5% up to a maximum of 8% per annum over any 3-year cycle (conway & goodman 2013). hluhluwe–imfolozi park (hip) is located in kwazulu-natal and contains one of the most important black rhino populations in southern africa (brooks & adcock 1997; emslie & brooks 1999). in addition, it has historically served as a donor population for black rhino range expansion programme (brrep) of the world wildlife fund (wwf) (wwf 2012). the hip black rhino population is believed to be above 70% of the estimated ecological carrying capacity and is therefore live-harvested for translocation at a rate of 5% per annum (clinning et al. 2009). for these translocations to be sustainable, an annual population growth rate of at least 5% is required to ensure that black rhinos do not become locally extinct (du toit 2006; hrabar & du toit 2005). however, there has only been one peer-reviewed assessment of the reproductive vital rates of the hip black rhino population and the factors which could be affecting these rates (hitchins & anderson 1983), and this was prior to the establishment of the brrep and the strategy to live harvest 5% of the population annually. population dynamics in herbivorous mammals are governed by the differences between births, deaths, emigration and immigration (fowler 1981). vital rates such as the age at sexual maturity (asm), conception rate, gestation length and inter-calving interval (ici) all influence fecundity, growth potential and turnover or generational time (gaillard, festa-bianchet & yoccoz 1998; gaillard et al. 2000). the minimum age at first conception indicates the potential for the population to increase because the earlier the minimum asm, the longer the lifetime productivity (owen-smith 1988). thus, if females attain sexual maturity and start reproducing at an early age, the growth rate for the population could increase even if no other vital rates change. longer icis reduce the number of calves born each year and therefore result in a reduction in the population growth rate. significantly, long-lived animals (such as black rhinos) can also be heavily influenced by stochastic environmental events such as drought, flood, fire and disease (fowler 1981; gaillard et al. 1998, 2000; greaver, ferreira & slotow 2013; hrabar & du toit 2005). because the hip black rhino population is an important population for both the kwazulu-natal province and the region more broadly, it is important that its vital rates are scrutinised in relation to national and provincial population growth targets. in this study, we assessed the reproductive parameters (i.e. asm, ici, fertility and fecundity rates) of the hip black rhino population. we also suggest potential drivers of the measured reproductive parameters within the hip black rhino population. we predicted that if the hip black rhino population was above 70% of the estimated ecological carrying capacity, then the reproductive parameters of the population would not be negatively affected by annual live-harvesting. materials and methods the hluhluwe and imfolozi game reserves were founded in 1895; are located 280 km north of durban, south africa; and are the oldest formally proclaimed protected areas in africa (brooks 2000; ezemvelo kzn wildlife 2011). the two reserves were amalgamated in 1989 to form the hip which is 895 km² in size (figure 1) (brooks 2000). the park is divided into five management sections, namely, manzimbomvu, nqumeni, masinda, mbhuzane and makhamisa. the park covers a vast and diverse landscape, consisting mainly of steep wooded hills, grass-covered slopes and riverine woodland along the many rivers and streams in the north of the park (brooks 2000) and more undulating; open to bush-encroached savanna woodland in the south. it is situated in the savanna biome, a tropical vegetation type that is characterised by woody plants and grasses (cowling, richardson & pierce 1997). hip is managed by ezemvelo kwazulu-natal wildlife (ezemvelo kzn wildlife 2011), which is a provincial nature conservation and tourism organisation responsible for the conservation of biodiversity and associated activities for the province of kwazulu-natal (ezemvelo kzn wildlife 2011). figure 1: the location of hluhluwe–imfolozi park in south africa. sightings data of individually recognisable animals have been collected at hip from the early 1980’s by field staff, scientists and students working within hip (clinning et al. 2009). for our assessment, we used individual black rhino sightings data from 1998 to 2013. individual black rhinos were identified by unique patterns of notches cut into their ears during marking operations, usually whilst the calves were still with their mothers, and as part of routine black rhino management operations in the park. population estimates were a total of all recognisable animals; animals known by association; and known, clean animals (i.e. individual black rhinos that had not been ear-notched but were recognisable by other physical features, location or by association with notched animals) which had been sighted within the last three calendar years (goodman 2013). the harvest rate (ht) was calculated using the following formula (goodman 2013): where, ht = the harvest rate, ht = the number of animals removed in year t and nt-1 = the population estimate in the preceding year (goodman 2013). population demographics we used population age structure information for the black rhino population at hip from 1998 to the end of 2013. we used the detailed population status reports (i.e. demographics and status), compiled on the 31st of december each year to extract the relevant data. age at sexual maturity age at sexual maturity was determined as the age at which a female gave birth to her first calf. the mean asm for the hip population was determined using a sub-sample of 120 individual female black rhinos which were in the population in 1998 or in subsequent years even though they may have given birth to their first calf prior to 1998. these females had the most complete life-history records (i.e. their birth dates and the birth dates of their first and subsequent calves were known). females which gave birth to their first calf before the end of 2013 were included. inter-calving interval from the same sub-sample of 120 black rhino females, 79 had more than one calf prior to 31 december 2013 and these individuals were used to calculate icis. the mean ici was calculated for each female with more than one calf, and icis for females from the same section (masinda, nqumeni etc.) were then combined to calculate the mean ici for each section. the weighted mean ici for the park was then calculated using the number of females from the five management sections, each of which had more than one calf prior to 31 december 2013. fertility and fecundity rates fertility rate (fert), which is the number of calves born per year as a proportion of mature females (i.e. females aged ≥ 7 years) in the population at the beginning of each year, was calculated as follows (goodman 2013): where, birthst is the number of births in the year under review and ad ft-1 is the number of adult females (i.e. females aged ≥ 7 years) in the preceding year (goodman 2013). the fertility rate at hip was calculated between 1998 and the end of 2013. fecundity rate (fect), which is the number of calves born in a year and still alive at the end of that year as a proportion of adult females (i.e. females ≥ 7 years old) in the population at the beginning of each year, was calculated as follows: where, surviving birthst is the number of calves that survived in the year under review and ad ft-1 is the number of adult females (i.e. females ≥ 7 years old) in the preceding year (goodman 2013). the fecundity rate at hip was calculated from 2010 to 2013 when the relevant information was available. mortality mortality rate was calculated as the number of black rhinos that died per year divided by the number of animals alive at the beginning of that year (sinclair, fryxell & caughley 2006). the mortality rates were calculated from 2009 to 2013, a period when all mortalities were investigated and accurately documented. statistical analysis binomial proportions tests were used to test for any difference in the overall sex ratio of the black rhino calves born on hip between 1998 and 2013 using r (r development core team 2014). data are presented as means ± s.d. (unless stated otherwise), and the level of significance was set at 95% (i.e. p = 0.05). results population demographics adult black rhinos (aged > 7 years) comprised the greatest proportion of the hip population in all years, and there were more females than males (figures 2 and 3). the proportion of adult males in the population has stayed relatively stable (figure 2), whereas the proportion of adult females has declined since 1998 (figure 3). juvenile (birth to 1 year) black rhinos made up the lowest proportion of the overall hip population in all years (figures 2 and 3). the proportion of sub-adult individuals (> 1 year but < 7 years) fluctuated over time but remained relatively stable (± 30% of the population) for the males (figure 2). by contrast, the proportion of female sub-adults increased steadily from ± 24% in 1998 to 33.3% in 2013 (figure 3). figure 2: annual changes in the proportion of male black rhinos in each of the six age classes in hluhluwe–imfolozi park on 31 december, annually from 1998 to 2013. figure 3: annual changes in the proportion of female black rhinos in each of the six age classes in hluhluwe–imfolozi park on 31 december, annually from 1998 to 2013. there were no significant differences in the sex ratios of the black rhino calves born at hip between 1998 and 2013, except in 2002 when significantly more female than male calves were born (χ2 = 4; df = 1; p = 0.045). in all years except 2002, either only one calf of a particular sex was born or the ratio was 1:1. in 2002, the number of female calves born was double that of males. between 1998 and 2003, the hip black rhino population had significantly more adult females than males (table 1). between 2004 and 2013, adult sex ratios were close to parity but remained slightly female biased except for 2005 and 2010 where significantly more females were present (table 1). table 1: the number of adult (> 7 years old) male and female black rhinos and their sex ratio at hluhluwe-imfolozi park between 1998 and 2013. age at sexual maturity the median asm for the entire black rhino population at hip was 11 years (132 months). the range in asm was 4–23 years (mean = 12 years/145 ± 11 months). assuming a gestation length of 15 months (goddard 1967), the mean age of first mating and conception was 130 ± 11 months (10 years). median age of first mating and conception was 117 months (9 years and 9 months). the earliest mating (backdated from birth) recorded was of a female that was 2 years, 8 months (34 months) old and the oldest recorded mating was of a female aged 22 years, 5 months (269 months). inter-calving interval, births per year and births per lifetime of the 120 black rhino females for whom adequate records were available, 79 had more than one calf. this gave a subset of 190 icis (figure 4). the weighted mean ici was 3 years, 8 months (45 ± 6 months), and ici ranged between 39 and 53 months across the five management sections of the park. the median ici was 37 months – the shortest ici was 1 year, 6 months (18 months) and the longest ici was 9 years (108 months). when icis longer than 40 months were removed from the dataset (hitchins & anderson 1983), 109 individual icis remained, and the calculation of the overall ici for the hip became 2 years, 5 months (29 ± 6 months). figure 4: histogram showing the frequency distribution of inter-calf intervals for the 79 black rhino females at hluhluwe–imfolozi park, each of which had more than one calf prior to 31 december 2013. using an ici of 45 months, the births per year per female were 0.27 and when using an ici of 29, births per year increased to 0.41 per female. the oldest female recorded to give birth at hip between 1998 and 2013 was aged 31 years and the mean age at first parturition was 12.08 years (145 ± 11 months). thus, if 31 years is assumed to be the age at which female black rhinos at hip cease reproductive activity, the reproductive lifetime of the population is approximately 19 years. a female black rhino may therefore be expected to have between five and eight calves in her lifetime. fertility and fecundity rates the fertility rate of black rhinos at hip was lowest in 2006 and 2009 (0.17 for each year) and highest in 2000 (0.34). in the 16 years between 1998 and 2013, the fertility rate averaged 0.24 (figure 5). the fecundity of the black rhino population at hip was below 0.25 between 2010 and 2013. the lowest fecundity rate was 0.21 in 2010, whilst the highest rate was 0.24 in 2011 and 2013. the mean fecundity rate over the 4 years (2010–2013) was 0.23. figure 5: the mean fertility rate of the black rhino population at hluhluwe–imfolozi park for 15 years, from 1998 to 2013. the dashed line indicates the targeted fertility rate (25%). mortality the mortality rate of black rhinos at hip was relatively low. the highest mortality rate was recorded in 2010 and 2013, when 0.07% of the population died. the total number of dead rhinos in 2013 was 14, only one calf died this year compared to other years where calf mortalities made up almost half of all mortalities. the remaining 13 deaths were of older individuals. the causes of death included fighting, natural death and poaching. discussion population age structure in large, predator-free systems, the population growth of herbivorous mammals are most sensitive to adult survivorship (gaillard et al. 1998, 2000). however, high temporal variation in juvenile survival typically drives variability in population growth rates (gaillard et al. 1998, 2000) and is thus an important factor in determining fluctuations in the populations of large mammalian herbivores (owen-smith & mason 2005). in expanding mammalian populations, the proportion of juveniles and sub-adult animals will be relatively higher and the mean age of the population will be low (coulson et al. 2004; owen-smith 1988). populations that are close to carrying capacity have a higher proportion of adult animals and a reduced juvenile survival and fecundity (coulson et al. 2004). stable populations are made up of about 60% – 70% reproductively mature adults, whilst increasing populations have 45% – 55% mature adults (owen-smith 1988). for example, an increasing black rhino population in the great fish river nature reserve (gfrnr), eastern cape, south africa, comprised 20% juveniles, 36% sub-adults and 44% adults (fike 2011). by contrast, the thomson’s gazelle (eudorcas thomsonii) population in the serengeti, tanzania, was shown to be in rapid decline when the population was made up of 12% juveniles and 70% adults (borner et al. 1987). owen-smith (1988) demonstrated that an east african black rhino population which consisted of about 60% adults were categorised as stable. our study shows that the current black rhino population at hip consists of 63.2% adults, 25.5% sub-adults and 11.3% juveniles, suggesting that the population is stable (owen-smith 1988). however, the aim is to have a black rhino population which is growing and reproducing at a high rate. in order for this to occur, the proportion of juveniles and sub-adult animals needs to be higher than it currently is and the population should consist of between 45% and 55% mature individuals (coulson et al. 2004; owen-smith 1988). an increase in the proportion of sub-adult females could potentially result in an increase in the fertility rates as more females could breed each year. sex ratio sex ratio is an essential factor influencing the growth rates and population dynamics of many large mammal populations (okita-ouma et al. 2009; weladji & laflamme-mayer 2011). this is because it affects the reproductive potential of a population, which is the relative capacity of a species to reproduce under favourable conditions (okita-ouma et al. 2009; weladji & laflamme-mayer 2011). a population with a female skew, under favourable conditions, is expected to demonstrate high growth rates (okita-ouma et al. 2009). the favourable conditions will allow females to reach the body mass required for reproduction sooner and asm will then be attained earlier (gaillard et al. 2000). rainfall and range conditions at the time of conception have been found to influence conception rates and sex ratios in black rhinos (berkeley & linklater 2010). more female black rhino offspring are born during years with below average rainfall and more males are conceived in the years with above average rainfall (hrabar & du toit 2005; weladji & laflamme-mayer 2011). the extrinsic modification hypothesis suggests that offspring sex can be influenced by the environmental conditions experienced by the mother (weladji & laflamme-mayer 2011). more male calves are born to dams that are gaining body condition during conception, compared to more female calves when dams are losing body condition at conception (cameron & linklater 2007). an assessment and comparison of female body condition at the time of conception and calf sex was not possible during our study. however, research has shown that male foetuses are affected by poor maternal resources and weather experienced by the mother prior to conception and during mid to late gestation (berkeley & linklater 2010; gaillard et al. 2000; weladji & laflamme-mayer 2011). this is because females eating food with low nutrient levels have poor body condition and poor body condition in black rhinos affects the sex of the calf (hrabar & du toit 2005). owen-smith (1988) found that the flexibility in the time of conception in black rhinos could also allow females to selectively abort embryos of the less advantageous sex (males) during unfavourable conditions (owen-smith 1988). previous periods with male-biased sex ratios in hip were associated with conception that occurred during wetter seasons (berkeley & linklater 2010). in umkhuze game reserve, which is ± 30 km north of hip, a male-biased sex ratio was related to lower rainfall during pregnancy and a relatively high population size (weladji & laflamme-mayer 2011). an increase in population size could bring about increased competition for food resources and it would be advantageous to produce male calves that could potentially disperse away from their natal ranges (weladji & laflamme-mayer 2011). however, having more males in a fenced system could also be counter-productive because with limited space, male-male conflict (and hence mortality) is increased. the sex ratio for the black rhino population at the gfrnr in the eastern cape, south africa, was 1.3 females per male (fike 2011). this could be indicative of the lower than average rainfall experienced at gfrnr, eastern cape, south africa, which resulted in more female calves being born (hrabar & du toit 2005). in hip, the overall sex ratio was close to parity but the adult and sub-adult sex ratio was slightly female biased (1.2:1). a slightly female biased sex ratio for adults under favourable conditions should result in high growth rates and thus as increase in the population going forward (okita-ouma et al. 2009). park managers are unable to manipulate the sex ratio of calves, but they could potentially manipulate the adult and sub-adult sex ratio through the removal of males. age at sexual maturity the asm in mammals differs between populations and even within the same species (bronson 1989). the genetic makeup of each individual accounts for the differences in the asm for females in the same population (bronson 1989). environmental factors such as food availability, ambient temperature and, to a lesser extent, humidity can influence reproductive development (bronson 1989). these factors also influence asm by influencing the rate of growth for each individual female (bronson 1989). evidence shows that underweight females and those on nutrient-poor diets may fail to ovulate (hrabar & du toit 2005). in addition, low rainfall reduces the quality and quantity of the food supply, negatively affecting the body condition of black rhino females (hrabar & du toit 2005). a threshold mass must be attained by mammals before they can attain sexual maturity and thus reproduce (gaillard et al. 2000). black rhino populations in poor habitats may attain sexual maturity and thus reproduce much later than black rhino populations in better habitats (gaillard et al. 2000). this is because poor nutrition in black rhino can lead to a delay in attaining the body weight required for reproduction (patton, campbell & parfet 2008). at the gfrnr in the eastern cape, south africa, the asm ranged from 4 years, 10 months to 9 years and 3 months, with the mean asm being 6 years and 8 months (fike 2011). the asm of the black rhino population in ithala game reserve, kwazulu-natal, south africa, was 6 years and 6 months (greaver et al. 2013). a study similar to the current one yielded an asm of between 5 years, 10 months and 6 years, 5 months (hitchins & anderson 1983). our data show that the asm of black rhinos at hip ranged between 4 and 23 years. the mean asm was 12 years. the asm for the current hip population exceeds the 7 years and 5 months quoted in du toit (2006) for poor performing populations and is almost double that of the 1983 hip estimate, and the gfrnr population. a number of potential reasons for this high asm include, poor habitat (gaillard et al. 2000), poor nutrition (patton et al. 2008), changes in predator effects on juvenile black rhinos (plotz & linklater 2009) and potentially negative social effects from annual removals. the black rhino females in the current hip population could be conceiving much earlier but losing their calves before they are born due to poor body condition (garnier, holt & watson 2002; patton et al. 2008). removals may affect the social structure of populations. such imbalance in the social structure may result in shifts in home ranges. productivity may then be affected as individuals would expend more energy creating new home ranges rather than investing in reproductive activities (patton et al. 2008). however, it is important to recognise that the calculation of asm in our study is based on the determination of the first known calf of a female. thus, it is possible that some calves may have died prior to them being recorded, and may therefore have been missed during monitoring, potentially inflating the asm. inter-calving interval the variation in the length of black rhino icis depends on the age of the female and the quality of food within her home range (du toit 2006). under favourable conditions, black rhino females can produce successive calves every 2–3 years (du toit 2006). during unfavourable conditions, the length of the ici can be in excess of 3 years, 6 months (du toit 2006). icis are not restricted to yearly time increments because reproduction in black rhinos is asynchronous (hrabar & du toit 2005). significantly, female megaherbivores can lengthen their ici as an ecological response to food restrictions which occur when populations are close to carrying capacity (owen-smith 1988). female black rhinos that are 28 years or older take longer to regain body condition after weaning calves and tend to have longer intervals between calves (du toit 2006). the icis for other south african black rhino populations range between 19 months and 7 years, 5 months (fike 2011; hall-martin & penzhorn 1977; hitchins & anderson 1983; hrabar & du toit 2005). in our study, the shortest ici was 1 year, 6 months and the longest ici was 8 years, 11 months. the mean ici was 3 years, 9 months and this falls into the category of poor to very poor fecundity (du toit 2006). this poor fecundity could be due to unfavourable conditions in the park (owen-smith 1988). however, similar to the calculation of the asm, any calves that were born but died before they were recorded could have inflated our estimates of ici. nevertheless, freeman et al. (2014) suggested that black rhino icis can be extended in the presence of predators and where intra-specific competition is apparently high. because of the annual live-harvesting of the hip population, intra-specific competition is unlikely to be the major driver of ici extension. by contrast, predation of rhino calves may be an important factor increasing icis at hip (see below and gaillard et al. 2000). births per year and lifetime productivity reproduction in black rhinos in the wild may cease between 30 and 35 years (garnier, bruford & goosens 2001; schenkel & schenkel-hulliger 1969) or at the age of 40 (owen-smith 1988). with a mean ici of 3 years, 8 months, the births per year for black rhino females at hip were 0.3. this rate was less than the births per year of 0.4 found at gfrnr, eastern cape, south africa, (fike 2011) as a result of much longer icis at hip or perhaps predation of calves, which were not detected through monitoring. fertility rate inter-calving interval and the gestation period determine the number of females that give birth each year (owen-smith 1988). ici is affected by both density dependent and independent factors, and these factors also influence fertility rates (hrabar & du toit 2005). at high densities, the ici increases and this causes a decline in the number of females that give birth each year (hrabar & du toit 2005). under favourable conditions, ici is at a minimum and this causes fertility rates to increase (fike 2011; hrabar & du toit 2005). in the gfrnr, eastern cape, south africa, the fertility rate was between 33% and 41% (fike 2011). pilanesberg national park, north west, south africa, had fertility rates of between 16% and 45% (hrabar & du toit 2005). at hip, the highest fertility rates were reached in 2000 when 34% of adult females gave birth. the lowest fertility rates were observed in 2006 and 2009 when only 17% of adult females gave birth. the mean fertility rate for the current population was 24% which is just less than the 25% fertility rate target (goodman 2013). mortality black rhino calf mortality rates within the first year after birth in south africa and namibia range between 8% and 14% (du toit 2006). sub-adult mortalities are between 2% and 4%, less than 2% in young and prime adults and 4% in much older adults (du toit 2006). males have higher mortality rates than females because the main cause of death in males are wounds from fighting, whereas most females die of old age (du toit 2006). mortality at hip was relatively low and deaths were mostly of calves. the highest mortality in the 4 years between 2009 and 2012 was 7% in 2010. most calf mortalities occur within the first month of birth and are linked to maternal care in cases where predation on calves is not high (gaillard et al. 2000). maternal age, size and reproductive experience also have a strong influence on calf survival (gaillard et al. 2000). where predators are present, predation can account for about 50% of calf deaths. maternal experience can lower calf vulnerability to predation (gaillard et al. 2000). however, malnutrition is the main cause of early mortality and may cause females to abandon their young. malnutrition also increases the juvenile’s susceptibility to other sources of mortality such as disease because of a weak immune system (gaillard et al. 2000). calf mortalities made up most of the mortalities at hip between 2009 and 2012. however, in 2013, most of the mortalities involved sub-adults and adults. it is possible that the better monitoring methods employed from 2009 onwards increased the chances of detecting calf births and mortalities than in previous years. it could also be a result of predation on the calves by lions (panthera leo) and hyaenas (crocuta crocuta), or malnutrition. conclusion the black rhino population at hip is an important donor population for the brrep. this means that this population must perform well both genetically and demographically in order to continue contributing to the brrep. the rhino management group (rmg) targets set for black rhino sub-populations in the southern african region include a minimum of 5% growth rate per year and 25% fecundity (goodman 2013). the current black rhino population at hip did not quite attain the 25% fecundity target. however, the difference between the fecundity rate of the hip population and the rmg target is small. therefore, it can tentatively be concluded that the current hip black rhino population is just meeting the minimum performance levels according to rmg standards. however, the delayed asm and longer icis are a concern and need to be addressed to ensure the population reproduces at a higher rate. the poor population performance of the hip black rhinos could be a result of poor habitat quality, poor animal condition, females losing their first calves, predation of calves or a negative social effect of annual live-harvesting of the population. annual rhino removals may be disrupting the social structure of the population, but we do not have any direct evidence to support this contention. it is unlikely that a lack of good-quality habitat is the main cause of the poor observed reproductive rates since reducing overall rhino densities through removals should theoretically reduce competition for food, resulting in improved reproductive rates. however, if the hip black rhino population has (as our data point towards) already exceeded the estimated ecological carrying capacity, then habitat quality for black rhinos may, in fact, be declining in hip as suggested by reid et al. (2007). we agree that the ecological carrying capacity estimate itself is likely incorrect (reid et al. 2007). if this estimate is incorrect, then it would likely explain the poor reproductive performance of the population. we therefore recommend that the accuracy of the ecological carrying capacity estimate be assessed as a matter of urgency to prevent the continued poor performance by the hip black rhino population and that a moratorium be placed on the live-harvesting of individuals until the estimate has been refined. acknowledgements we thank geoff clinning (ezemvelo kzn wildlife) for all his work in constructing the black rhino database for hip, error-checking of historical data and for his assistance in providing data and comments. heleen druce assisted with the capture and error-checking of historical black rhino data. bom ndwandwe, numerous ezemvelo kzn wildlife field and scientific staff and students assisted with the field data collection over the years. we would like to thank the national research foundation and the oppenheimer memorial trust for funding. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions z.n.n. was the project leader. z.n.n., d.m.p. and d.j.d. were responsible for the project design. z.n.n. collected some of the field data and analysed these data with the assistance of d.m.p. and d.j.d. all authors were involved in the writing of the article and the interpretation of the results. references berkeley, e.v. & linklater, w.l., 2010, ‘annual and seasonal rainfall may influence progeny sex ratio in the black rhinoceros’, south african journal of wildlife research 40, 53–57. https://doi.org/10.3957/056.040.0102 borner, m., fitzgibbon, c.d., borner, m., caro t.m., lindsay, w.k., collins, d.a. et al., 1987, ‘the decline of the serengeti thomson’s gazelle population’, oecologia 73, 32–40. https://doi.org/10.1007/bf00376974 bronson, f.h., 1989, mammalian reproductive biology, the university of chicago press, chicago, il. brooks, m. & adcock, k., 1997, conservation plan for the black rhinoceros diceros bicornis in south africa, report, the rhino management group, pietermaritzburg. brooks, s., 2000, ‘re-reading the hluhluwe–umfolozi game reserve: constructions of a natural space’, transformation 44, 63–79. cameron, e.z. & linklater, w.l., 2007, ‘extreme sex ratio variation in relation to change in condition around conception’, biology letters 3, 395–397. https://doi.org/10.1098/rsbl.2007.0089 clinning, g., druce, d., robertson, d., bird, j. & nxele, b., 2009, black rhino in hluhluwe–imfolozi park: historical records, status of current population and monitoring and future management recommendations, report, ezemvelo kzn wildlife, pietermaritzburg. conway, a.j. & goodman, p.s., 2013, strategy for the management of black rhinocerous (diceros bicornis minor) in kwazulu-natal, report, ezemvelo kzn wildlife, pietermaritzburg. coulson, t., guiness, f., pemberton, j. & clutton-brock, t.h., 2004, ‘the demographic consequences of releasing a population of red deer from culling’, ecology 85, 411–422. https://doi.org/10.1890/03-0009 cowling, r.m., richardson, d.m. & pierce, s.m., 1997, vegetation of southern africa, cambridge university press, cape town. du toit, r., 2006, guidelines for implementing sadc rhino conservation strategies, report, sadc regional programme for rhino conservation, pretoria. emslie, r.h. & brooks, m., 1999, african rhino: status survey and conservation action plan, iucn/ssc african rhino specialist group, gland. ezemvelo kzn wildlife, 2011, integrated management plan: hluhluwe–imfolozi park, south africa, report, ezemvelo kzn wildlife, pietermaritzburg. fike, b., 2011, ‘the demography and population dynamics of a re-introduced black rhinoceros population on the great fish river reserve, eastern cape province’, msc thesis, department of zoology & entomology, rhodes university. fowler, c.w., 1981, ‘density dependence as related to life history strategy’, ecolological society of america 62, 602–610. https://doi.org/10.2307/1937727 freeman, e.w., meyer, j.m., bird, j., adendorff, j., schulte, b.a. & santymire, r.m., 2014, ‘impacts of environmental pressures on the reproductive physiology of subpopulations of black rhinoceros (diceros bicornis bicornis) in addo elephant national park, south africa’, conservation physiology 2, cot034. gaillard, j.-m., festa-bianchet, m. & yoccoz, n.g., 1998, ‘population dynamics of large herbivores: variable recruitment with constant adult survival’, trends in ecology and evolution 13, 58–63. https://doi.org/10.1016/s0169-5347(97)01237-8 gaillard, j.-m., festa-bianchet, m., yoccoz, n.g., loison, a. & toigo, c., 2000, ‘temporal variation in fitness components and dynamics of large herbivores’, annual review of ecology systematics 31, 367–393. https://doi.org/10.1146/annurev.ecolsys.31.1.367 garnier, j.n., bruford, m.w. & goosens, b., 2001, ‘mating systems and reproductive skew in the black rhino’, molecular ecology 10, 2031–2041. https://doi.org/10.1046/j.0962-1083.2001.01338.x garnier, j.n., holt, w.v. & watson, p.f., 2002, ‘non-onvasive assessment of oestrus cycles and evaluation of reproductive seasonality in the female wild black rhinoceros (diceros bicornis minor)’, reproduction 123, 877–889. https://doi.org/10.1530/rep.0.1230877 goddard, j., 1967, ‘home range, behaviour and recruitment rates of two black rhinoceros populations’, east african wildlife journal 5, 133–150. https://doi.org/10.1111/j.1365-2028.1967.tb00768.x goodman, p.s., 2013, ezemvelo kzn wildlife monitoring plan: black rhinoceros (diceros bicornis minor), unpublished report, ezemvelo kzn wildlife, pietermaritzburg, south africa. greaver, c., ferreira, s. & slotow, r., 2013, ‘density-dependent regulation of the critically endangered black rhinoceros population in ithala game reserve, south africa’, austral ecology 39, 1–11. hall-martin, a.j. & penzhoen, b.l., 1977, ‘behaviour and recruitment of translocated black rhinoceros diceros bicornis’, koedoe 20, 147–162. https://doi.org/10.4102/koedoe.v20i1.941 hitchins, p.m. & anderson, j.l., 1983, ‘reproduction, population characteristics and management of the black rhinoceros diceros bicornis minor in the hluhluwe/corridor/umfolozi game reserve complex’, south african journal of wildlife research 13, 78–85. hrabar, h. & du toit, j.t., 2005, ‘dynamics of a protected black rhino (diceros bicornis) population: pilanesberg national park, south africa’, animal conservation 8, 259–267. https://doi.org/10.1017/s1367943005002234 international union for conservation of nature (iucn), 2012, iucn red list of threatened species (ver. 2012.2), viewed 17 october 2012, from http://www.iucnredlist.org okita-ouma, b., amin, r., van langevelde, f. & leader-willians, n., 2009, ‘density dependence and population dynamics of black rhinos (diceros bicornis michaeli) in kenya’s rhino sanctuaries’, african journal of ecology 3, 791–799. https://doi.org/10.1111/j.1365-2028.2009.01179.x owen-smith, r.n., 1988, megaherbivores: the influence of very large body size on ecology, cambridge university press, cape town. owen-smith, r.n. & mason, d.r., 2005, ‘comparative changes in adult vs. juvenile survival affecting population trends of african ungulates’, journal of animal ecology 74, 762–773. https://doi.org/10.1111/j.1365-2656.2005.00973.x patton, f., campbell, p. & parfet, e., 2008, ‘biological management of the high density black rhino population in solio game reserve, central kenya’, pachyderm 44, 72–79. plotz, r.d. & linklater, w.l., 2009, ‘black rhinoceros (diceros bicornis) calf succumbs after lion predation attempt: implications for conservation management’, african zoology 44, 283–287. https://doi.org/10.3377/004.044.0216 r development core team, 2014, r: a language and environment for statistical computing, r foundation for statistical computing, vienna. reid, c., slotow, r., howison, o. & balfour, d., 2007, ‘habitat changes reduce the carrying capacity of hluhluwe–umfolozi park, south africa, for critically endangered black rhinoceros diceros bicornis’, oryx 41, 247–254. https://doi.org/10.1017/s0030605307001780 schenkel, r. & schenkel-hulliger, l., 1969, ecology and behaviour of the black rhinoceros (diceros bicornis l.): a field study, paul parey, hamburg. sinclair, a.r.e., fryxell, j.m. & caughley, g., 2006, wildlife ecology, conservation and management, blackwell, oxford. weladji, r.b. & laflamme-mayer, k., 2011, ‘influence of environmental conditions on sex allocation in the black rhinoceros population of mkhuze game reserve, south africa’, african journal of ecology 49, 471–480. https://doi.org/10.1111/j.1365-2028.2011.01280.x wwf, 2012, black rhino range expansion programme, viewed 27 july 2012, from www.wwf.org.za.what_we_do/species/black/rhino/ article information authors: morgan b. pfeiffer1 jan a. venter2,3 colleen t. downs1 affiliations: 1school of life sciences, university of kwazulu-natal, south africa 2department of biodiversity conservation, eastern cape parks and tourism agency, east london, south africa 3centre for wildlife management, university of pretoria, south africa correspondence to: colleen downs email: downs@ukzn.ac.za postal address: private bag x01, scottsville 3209, south africa dates: received: 04 july 2014 accepted: 07 jan. 2015 published: 18 may 2015 how to cite this article: pfeiffer, m.b., venter, j.a. & downs, c.t., 2015, ‘foraging range and habitat use by cape vulture gyps coprotheres from the msikaba colony, eastern cape province, south africa’, koedoe 57(1), art. #1240, 11 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1240 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. foraging range and habitat use by cape vulture gyps coprotheres from the msikaba colony, eastern cape province, south africa in this original research... open access • abstract • introduction • methods    • study area    • cape vulture captures and marking    • data collection    • data analysis    • habitat use and gps tracking • results    • foraging ranges    • habitat use • discussion    • conservation implications • conclusion • acknowledgments    • competing interests    • authors’ contributions • references • appendix 1 abstract top ↑ despite the extent of subsistence farmland in africa, little is known about endangered species that persist within them. the cape vulture (gyps coprotheres) is regionally endangered in southern africa and at least 20% of the population breeds in the subsistence farmland area previously known as the transkei in the eastern cape province of south africa. to understand their movement ecology, adult cape vultures (n = 9) were captured and fitted with global positioning system/global system for mobile transmitters. minimum convex polygons (mcps), and 99% and 50% kernel density estimates (kdes) were calculated for the breeding and non-breeding seasons of the cape vulture. land use maps were constructed for each 99% kde and vulture locations were overlaid. during the non-breeding season, ranges were slightly larger (mean [± se] mcp = 16 887 km2 ± 366 km2) than the breeding season (mcp = 14 707 km2 ± 2155 km2). breeding and non-breeding season mcps overlapped by a total of 92%. kernel density estimates showed seasonal variability. during the breeding season, cape vultures used subsistence farmland, natural woodland and protected areas more than expected. in the non-breeding season, vultures used natural woodland and subsistence farmland more than expected, and protected areas less than expected. in both seasons, human-altered landscapes were used less, except for subsistence farmland. conservation implications: these results highlight the importance of subsistence farmland to the survival of the cape vulture. efforts should be made to minimise potential threats to vultures in the core areas outlined, through outreach programmes and mitigation measures. the conservation buffer of 40 km around cape vulture breeding colonies should be increased to 50 km. introduction top ↑ africa has been inhabited by humans for over 300 000 years (fisher et al. 2013; sheehan & sanderson 2012). within that time, communal grazing of livestock, human-induced fires, depletion of indigenous forests and urbanisation have altered many landscapes (lawes, griffiths & boudreau 2007; sheehan & sanderson 2012; skead 1987; vetter & bond 2012). although heavily human-altered landscapes are often degraded, endangered species can persist in these environments (mckee et al. 2004; phipps et al. 2013b). one opportunistic animal guild that has coexisted with humans for centuries is the vulture (haas & mundy 2013; moleón et al. 2014). vultures perform an important ecosystem service by consuming carcasses. vultures recycle nutrients, reduce the potential for the spread of infectious diseases, and provide a carbon-neutral waste removal service (dupont et al. 2012; prakash et al. 2003; ogada et al. 2012b). in some cultures, vultures are highly revered and, for example, are used to ritually dispose of human corpses (haas & mundy 2013). however, 61% of vulture species worldwide are vulnerable to extinction from a variety of threats (ogada et al. 2012a). understanding how vultures persist in human-altered landscapes will provide information on where and how to focus conservation efforts on a regional and global scale. the cape vulture (gyps coprotheres), a colonial nesting scavenger, is endemic to southern africa. it is listed as ‘vulnerable’ on the red list of threatened species of the international union for conservation of nature and natural resources (iucn) and as ‘endangered’ in the eskom red data book of birds of south africa, lesotho and swaziland (birdlife international 2013). at least 20% of the global population breeds in the former bantustan homeland of the transkei in the eastern cape province of south africa (birdlife international 2013; boshoff, piper & michael 2009; piper 1994). this area was created under segregation laws of the former apartheid government of south africa and is characterised by high human densities and subsistence farmland (kepe 1997; shackleton et al. 1991; statistics south africa 2011). in this area, every resident has access to communal grazing land and livestock numbers are not restricted (vetter & bond 2012). most of the cape vulture breeding colonies in the former transkei are located in formal protected areas or important bird areas (ibas) (birdlife south africa 2013). two of the three protected areas, namely collywobbles vulture colony (iba sa088) and pondoland cape vulture colonies (iba sa126), were designated as ibas specifically to promote the conservation of this threatened species (birdlife international 2014a; birdlife international 2014b). however, foraging vultures are rarely confined to protected areas and are thus exposed to numerous threats elsewhere (bamford et al. 2007; phipps et al. 2013a). for example, cape vultures are illegally killed for the traditional medicine market and are negatively impacted by power line infrastructure in the eastern cape (boshoff et al. 2011; mander et al. 2007). poisoned carcasses, resulting in mass vulture mortalities, appear to be an infrequent occurrence in subsistence farmland areas, but do occur on commercial farms (brown & piper 1988). a possible benefit to vultures in the former transkei is the relatively high livestock mortality rates compared to commercial farming areas, which results in an abundance of carrion (boshoff et al. 2009; vernon 1998). furthermore, the landscape in the former transkei contains numerous suitable cliffs on which cape vultures roost and breed (mundy et al. 1992; piper & ruddle 1986). despite having knowledge of the potential threats and perceived benefits for cape vultures, knowledge of the movement ecology and detailed demographic information of cape vultures in this area is lacking. birdlife south africa, the endangered wildlife trust (ewt) and a number of bird specialists recommend 40 km buffers around cape vulture breeding colonies as conservation priority areas to prevent mortalities from wind turbines and hazardous power infrastructure development (boshoff & minnie 2011; retief et al. 2013). breeding cape vultures of the southern node population are known to forage and move extensively within this range (boshoff & minnie 2011; boshoff, robertson & norton 1984; brown & piper 1988; robertson & boshoff 1986). breeding vultures that forage within 40 km of the colony are better able to relieve their partner of parenting duties so that both can forage on the same day (ruxton & houston 2002). vultures that forage in this manner are thought to have higher breeding success because of a higher food delivery rate to the chick (ruxton & houston 2002). however, telemetry-based cape vulture studies in other regions have indicated that both breeding and non-breeding cape vultures forage considerably farther than 40 km from the breeding colony, which may weaken the conservation goals of the colony buffers (bamford et al. 2007; phipps et al. 2013b). foraging ranges of vultures may be influenced by the surrounding land uses or presence of vulture feeding sites. vulture feeding sites (vulture restaurants) provide an uncontaminated, regular supply of carrion for vultures, which aims to prevent mortalities from food shortages and poisonings (piper, boshoff & scott 1999). most operate on commercial farms in south africa (ewt and ezemvelo kwazulu-natal [kzn] wildlife unpublished data). in the eastern cape, all active cape vulture breeding colonies are in or near subsistence farmland with few vulture feeding sites, but the degree of subsistence farmland use by cape vultures remains unknown. the aim of this study was to document the foraging range and habitat use of adult cape vultures in the former transkei from a colony in the mkambati nature reserve. one intention of the study was to test if 40 km buffers around southern cape vulture breeding colonies are adequate for their intended conservation purposes. the size, shape and habitat use in the overall foraging and core areas were investigated and possible seasonal differences quantified. seasons were separated into either breeding or non-breeding season. adult vultures were expected to conduct fewer foraging trips during the early breeding season and incubation (kendall et al. 2014; spiegel et al. 2013). breeding behaviour may concentrate cape vulture movements to areas that maximise the success of foraging trips. these areas would be ideal to identify for conservation planning. additionally, a small proportion of cape vultures may migrate from the eastern part of the eastern cape to the west in the non-breeding season (boshoff et al. 2009). if this migration occurs, it would be important to isolate any corridors or flight paths. vulture movements may be influenced by the availability of resources across the landscape. therefore, resource selection by the vultures was thought to differ with land use and season (murn & anderson 2008; vernon 1998). the location of the study vulture colony provides a unique opportunity to investigate the vultures’ use of subsistence and commercial farmland, as both land uses are present within 100 km of the colony. methods top ↑ study area the entire former transkei (approximately 27° e – 30° e and 33° s – 30° s) is located in the eastern cape and kwazulu-natal provinces of south africa (boshoff et al. 2009). the indian ocean coastal belt, savanna and grassland are the three major biomes in the study area (mucina et al. 2006). ngongoni grass (aristida junciformis) dominates the savanna and grassland biomes, while the indian ocean coastal belt supports patches of species-rich sour grasslands (mucina et al. 2006). the dominant herbivores in the study area are domestic livestock (either for subsistence or commercial purposes) and wild ungulates in fenced protected areas (boshoff & vernon 1980; shackleton et al. 1991). the msikaba cape vulture colony (31°16ʹ s, 29°59ʹ e; 200 m a.s.l.; figure 1) is located on cliffs formed by the msikaba river in the mkambati nature reserve (boshoff & minnie 2011; piper & ruddle 1986). at least 170 cape vulture pairs breed regularly within the mkambati nature reserve (botha et al. 2012). vulture breeding activity was first documented at msikaba in 1984; however, breeding attempts along the mtentu river, the northern boundary of the mkambati nature reserve, were first documented in the mid-1970s (piper & ruddle 1986). annual rainfall is about 1200 mm and the difference in monthly mean temperature is less than 6 °c along the coast (shackleton et al. 1991). figure 1: location of the former transkei, mkambati nature reserve and the msikaba cape vulture breeding colony in the eastern cape province of south africa. cape vulture captures and marking a 9 m × 6 m × 3 m wooden-framed walk-in cage trap (diekmann et al. 2004) was constructed at the cape vulture feeding site at the mkambati nature reserve. the walls of the cage consisted of wire mesh (100 mm) reinforced with steel cable. translucent shade cloth (50% opaqueness) was attached to the walls to prevent injuries to the vultures. construction and baiting of the trap with ungulate carcasses from the mkambati nature reserve commenced at least 7 months before capture attempts. each vulture captured was fitted with a unique metal south african bird ringing unit (safring) ring and patagial tags on both wings. adults (n = 9) were identified by plumage and eye colour (mundy 1982). avi-track (pietermaritzburg, south africa) global positioning system (gps)/global system for mobile (gsm) transmitters were attached as backpacks (n = 3) and pelvic mounts (n = 3) using teflon® ribbon. cellular tracking technologies (ctt) 1100 gps/gsm transmitters (somerset, pennsylvania, usa) were attached as backpacks (n = 2) and as a pelvic mount (n = 1). the average weights of the avi-track and ctt units were 97 g and 136 g respectively, which is less than 1% of the average weight of an adult cape vulture (piper 2005). data collection the avi-track transmitters were programmed to record the gps location of the vulture, direction of travel and speed at least six times a day in 2 h intervals from 06:00 to 18:00. the ctt transmitters were programmed to record the gps location of the vulture, horizontal dilution of precision, fix quality, direction of travel, speed and altitude every 15 min from sunrise to sunset. for comparison with the avi-track units, a subsample of the ctt data was created by using one data point every 2 h for a total of six gps locations a day. the first and last point of the day (which changed with day length) were used in addition to three points during the day, which were at least 2 h apart. data analysis the vulture transmitter data were entered into arcgis 9.3 (esri, www.esri.com) and projected to the universal transverse mercator (utm) (wgs [world geodetic system] 1984 utm zone 35s). to determine if an asymptote was reached during each season, the minimum convex polygons (mcps) were plotted in relation to the number of gps locations. visually, asymptotes were identified and vultures that reached asymptotes were used for further analyses. although widely criticised, mcp is the most commonly used home range estimator. it entails drawing the smallest polygon that incorporates all of the animal's locations (powell 2000). the mcps (100%) were calculated for each vulture in both breeding and non-breeding seasons and tested for differences. the breeding season data included all fixes from may to october 2013, while the non-breeding season data included fixes from november 2012 to april 2013 (mundy et al. 1992). the mean egg-laying period is may to june, with chicks hatching between july and august. fledglings can be dependent on their parents until october or november, and even into december (mundy et al. 1992; piper 1994). the percentage of mcp overlap was calculated for the two seasons. since mcps generally include areas that are not visited by the vulture, kernel density estimates (kdes) were used to identify high density areas of vultures. in previous studies, 95% kdes were found to produce numerous fragmented areas; hence 99% kdes were used (blundell, maier & debevec 2001; phipps et al. 2013b). fifty percent kdes were used to identify core areas. both 99% and 50% kdes were calculated for the breeding and non-breeding seasons and tested for seasonal differences. all kdes were calculated using bivariate fixed kernels with a reference bandwidth. least-squares cross validation calculations for kdes could not be used because of numerous identical roosting locations. the raster cell size was 1000 m x 1000 m. both mcp and kde contours were produced using the home range tools (hrt) extension for arcgis (rodgers et al. 2007). mann-whitney tests were used to determine differences in foraging range size (mcp, 99% kdes, 50% kdes) and season (breeding vs non-breeding). p-values < 0.05 were read as significant. statistical analyses were conducted in statistica (statsoft 2006). habitat use and gps tracking a land use map was created using the south african national land cover database merged with all the protected areas of south africa and lesotho (south african national botanical institute [sanbi] 2000; iucn and united nations environment programme's world conservation monitoring centre [unep-wcmc] 2014). the 41 original south african national land cover database land use categories were compressed into six land use classes: urban centres, village communities, natural woody vegetation, tree plantations, commercial farmland and subsistence farmland (table a1). as the original map did not illustrate livestock grazing land (only cultivated land), land use classes such as ‘natural grassland’ were separated into commercial or subsistence farmland based on their location to the former political boundaries of the transkei (figure a1). tree plantations and natural woody vegetation were separated because of the level of human transformation in these areas. to account for urban and suburban sprawl, 2 km buffers were placed around the urban and village layers. the polygon layer was converted into a raster with a cell size of 1800 m. the raster assigned one land use value to each cell, based on the cell centre. analysis was limited by the resolution of spatial data available, but was compensated appropriately using buffers and the unbiased method of assigning a land use value based on the cell's centre. the 99% kdes of pooled cape vulture locations from the breeding and non-breeding seasons were clipped to the final categorised land use map, excluding areas that extended into the indian ocean, since vultures do not fly above oceans (pers. obs.). for each 99% kde, areas of all land uses were calculated (km2). the number of vulture gps locations within each land use was also calculated. both procedures were conducted with hawth's analysis tools (beyer 2004). habitat use in proportion to availability, considering each land use separately, was tested using the bonferroni z-statistic in microsoft excel (byers, steinhorst & krausman 1984). in total, 34 cape vultures (including 1 recapture) were captured in 2012 and 2013, during the non-breeding season. the gps locations of birds were highly autocorrelated, with a mean schoener's index value of 0.10 ± 0.07. this index detected that the individual's gps locations were not independent of each other, which may result in underestimating home range estimates (swinhart & slade 1985). to correct this, all data were rescaled to unit variance using home range tools (rodgers et al. 2007). the reference bandwidth for both seasons across all individuals was 0.34 ± 0.02 for kdes. results top ↑ nine transmitters recorded location data for 277 ± 72 days. of the vultures fitted with transmitters, five reached an mcp asymptote during the breeding season and four during the non-breeding season (figure a2). the average number of gps locations for the breeding and non-breeding seasons were 717 ± 122 (n = 5) and 820 ± 123 (n = 4) respectively. the number of fixes required for mcps to become constant varied, but generally, transmitters with fewer than 300 gps locations were found to be insufficient. some vultures were tracked for both seasons, while other transmitters were deployed later in the season, or failed. three transmitters stopped working for unknown reasons (birds were resighted alive) before 300 gps locations were collected; these data were excluded from the analyses. two transmitters only collected data for one season (x023 and x022). of the vultures used for analysis, two (x027 and x023) were confirmed to have successfully raised chicks in 2013. one vulture (x022) was observed at a nesting site arranging nesting material with its partner, but did not breed. foraging ranges general movements of the cape vultures occurred from the breeding colony in the south to the south-western part of the kwazulu-natal province. no vultures travelled south of the mzimvubu river mouth during the tracking period. the pooled breeding season mcp overlapped 92% with the non-breeding season mcp (figure 2). the mean mcp during the breeding season was 14 707 km2 ± 2155 km2 (n = 5, median = 13 282 km2). the mean mcp during the non-breeding season was 16 887 km2 ± 366 km2 (n = 4, median = 16 602 km2). there was no significant difference between individual mcps (mann–whitney test, z = -0.49, p = 0.62). figure 2: the combined minimum convex polygons of adult cape vultures (gyps coprotheres) in the breeding (n = 5) and non-breeding seasons (n = 4) captured at the msikaba cape vulture colony in the eastern cape, south africa. individual 99% kdes were not significantly larger in the non-breeding season (mann–whitney test, z = -0.73, p = 0.46), nor were the 50% kdes (mann–whitney test, z = -1.71, p = 0.09). when cape vulture gps locations were pooled together, mcps and 99% kdes were only slightly larger in the breeding season than the non-breeding season (table 1). pooled 50% kdes were also only slightly larger (908 km2) in the non-breeding season (figure 3). minimum convex polygons and 99% kdes were similar across the breeding and non-breeding seasons, while the number and size of 50% kdes differed slightly. table 1: home range estimates for adult cape vultures (gyps coprotheres) captured at mkambati nature reserve, eastern cape, south africa. figure 3: combined kernel density estimates for adult cape vultures (gyps coprotheres) in the (a) breeding (n = 5) and (b) non-breeding season (n = 4) captured at the msikaba cape vulture colony, eastern cape, south africa. kernel density estimate areas that extended into the indian ocean were removed as vultures cannot forage there. the maximum radius of the 50% kde around the msikaba cape vulture colony was 46 km during the breeding season. the other two core 50% kdes during the breeding season had smaller radii (7 km and 11 km) and were located north of the breeding colony (figure 3). the northernmost 50% kde during the breeding season was mainly created by one bird (x042), which was not recorded at a breeding site at msikaba. in the non-breeding season, there were only two core 50% kdes areas. during the non-breeding season, the maximum radius from the colony to the edge of the 50% kde was 52 km. the 50% kde not located around the breeding colony had a radius of 29 km in the non-breeding season. habitat use when vulture locations were pooled, habitats were not selected in proportion to their availability. habitat selected by vultures differed between the breeding and non-breeding seasons (table 2; figure 4). cape vultures used subsistence farmland and natural woody vegetation more than expected in both the breeding and non-breeding season (table 2). protected areas were used in a greater proportion during the breeding season, while during the non-breeding season protected areas were used less than their availability in the 99% kdes (table 2). in both seasons, commercial farmland, plantations, urban centres and villages were used less than their availability in the 99% kdes of the vultures (table 2). table 2: habitat availability in pooled 99% kernel density estimate based on the reclassified land use map. bonferroni confidence intervals were used to determine cape vulture habitat use in pooled 99% kernel density estimate. figure 4: cape vulture (breeding and non-breeding season) locations overlaid on the land use map used for habitat analysis. discussion top ↑ this study highlights the importance of subsistence farmland, rather than commercial farmland, as foraging habitat for cape vultures from the msikaba colony. although the results presented here are from a small and restructured sample size (n = 9), they illustrate the seasonal foraging and habitat selection patterns of the cape vulture in the southern node population. adult vultures from the msikaba colony exhibited a well-defined foraging range. the tagged vultures did not participate in westerly migratory behaviour as previously reported by boshoff et al. (2009). foraging ranges calculated as mcps were found to overlap considerably in the breeding and non-breeding seasons. other studies have also observed that the distance covered by vultures during the breeding season is similar to the non-breeding season, but foraging trips occurred less frequently in the early breeding season and during incubation (bamford, monadjem & hardy 2007; kendall et al. 2014; spiegel et al. 2013). the 50% core area around the breeding colony was oval shaped and extended towards kwazulu-natal with a radius ranging 17 km – 46 km during the breeding season. during the non-breeding season the 50% kde around the colony increased to a maximum radius of 52 km. the size of the core area around the colony was therefore larger than the proposed 40 km buffer (boshoff & minnie 2011). as the home range represents an area ‘traversed by the individual in its normal activities of food gathering, mating and caring for young’ (burt 1943), a smaller range may be explained by the abundance of food or suitable roosts in the environment. formal protected areas were used more than expected during the breeding season, possibly because breeding sites were located in protected areas, not because there was more carrion available. as adult cape vultures were captured in the mkambati nature reserve and two vultures were confirmed successful breeders, more time was spent at this locale. cape vultures used formal protected areas less during the non-breeding season, while natural woody vegetation and subsistence farmlands were preferred. use of natural woody vegetation by the vultures may have been misinterpreted because of the scale of the habitat classifications, as cliffs were not distinguished in the habitat classifications. vultures did not necessarily use the woody vegetation, but the steep cliffs located above them, as roosting sites. roost sites in the study area were typically located on isolated cliff faces with indigenous forest at the base (boshoff & minnie 2011; pers. obs.). the two 50% kdes created during the non-breeding season were both located in subsistence farmland that contains only one formal (registered with ewt or ezemvelo kzn wildlife) vulture feeding site, which was located in the mkambati nature reserve. other studies have found that african vultures use subsistence farmland less (bamford et al. 2007; bamford et al. 2009). this could be explained by different livestock carcass management (burning or burying) or over-harvesting of forest resources, which prevent tree-nesting vulture species from inhabiting these areas (monadjem & garcelon 2005). in the former transkei, carrion may be more readily available because of inadequate animal husbandry and abundant tick-borne diseases (shackleton et al. 2013). strong cultural traditions may provide another scavenging opportunity for vultures: during traditional ceremonies, local amaxhosa people slaughter and butcher animals, the leftovers of which are discarded for vultures and other scavengers (pfeiffer pers. obs.). commercial farmland areas were used less than expected during both seasons, despite the presence of formal vulture feeding sites. however, the northernmost 50% kde in the breeding season was located near multiple vulture feeding sites. (one feeding site was located inside the northernmost 50% kde.) the number of fixes a day in the current study (six a day including roosting locations) may have been insufficient to identify feeding events, which may have resulted in underestimating the use of vulture feeding sites. furthermore, cape vultures can be grounded at roost sites for long periods of time because of adverse soaring conditions. accordingly, these results may overestimate roosting locations and underestimate feeding events (monsarrat et al. 2013; spiegel et al. 2013). future research should use higher resolution gps data in order to identify feeding events and then calculate habitat use. conservation implications the findings presented here highlight the relatively small foraging ranges of adult cape vultures from the msikaba colony and their extensive use of subsistence farmland. conservation efforts should focus on mitigating threats to vultures in the 50% kdes, which are mainly located in subsistence farmland. three local municipalities (ingquza hill, mbizana and umzimkhulu) were represented in both the breeding and non-breeding season 50% kdes. on-the-ground conservation projects by provincial staff and relevant non-government organisations should be conducted in these areas. as some cape vulture core areas differed between the breeding and non-breeding seasons, other local municipal districts could be targeted based on the time of the year. during the breeding season (may to october), impendle, umngeni and mpofana local municipalities were represented in the 50% kdes. in the non-breeding season (november – april), hibiscus coast, ezinqoleni, umuziwabantu and ubuhlebezwe local municipalities were represented and should be targeted for conservation projects during these months. based on these results, it is recommended that buffers around cape vulture colonies in the southern node population be increased from 40 km to 50 km. for cape vulture roost sites, 40 km buffers appear to be sufficient. in certain areas where this may be in conflict with development, a combination of gps tracking data and risk assessment modelling should be used to construct conservation priority areas (katzner et al. 2012). conclusion top ↑ although vultures are far-ranging foragers that will never be fully secure within protected areas, it is essential to identify and proclaim conservation buffers. tracking of a small sample of adult cape vultures from one colony has successfully identified the main foraging areas of vultures from that colony, and perhaps in the region. these areas can be targeted in focused strategic action plans aimed at avoiding or reducing the mortality of vultures. it will only be with the collaboration of communities, policy makers, conservation organisations and provincial governments that this regionally endangered vulture species will survive. acknowledgments top ↑ the gay langmuir bursary, the rufford grant, the university of kwazulu-natal, and the eastern cape parks and tourism agency are thanked for research funding. thank you to v. mapiya and the staff of the mkambati nature reserve for field assistance. the mazda wildlife fund is thanked for vehicle assistance. j. greeff, b. hoffman, s. kruger, s. mcpherson, m. and k. bowker, c. and k. nelson, d. berens, m. witteveen, d. allan, s. heuner, a. botha and vulpro are thanked for helping with the capture of the vultures. field work was conducted in accordance to the laws of south africa. permits for vulture captures were granted by the department of environmental affairs (tops permit nr. 05052) and approved by the ethics committee of the university of kwazulu-natal. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions m.b.p. (university of kwazulu-natal) was the project leader. m.b.p., j.a.v. (department of biodiversity conservation) and c.t.d. (university of kwazulu-natal) conceived and designed the experiments. m.b.p was responsible for performing the experiments and analysing the data. m.b.p., j.a.v. and c.t.d. wrote the article. references top ↑ bamford, a.j., diekmann, m., monadjem, a. & mendelsohn, j., 2007, ‘ranging behaviour of cape vultures gyps coprotheres from an endangered population in namibia’, bird conservation international 17(4), 331–339. http://dx.doi.org/10.1017/s0959270907000846 bamford, a.j., monadjem, a. & hardy, i.c.w., 2009, ‘nesting habitat preference of the african white-backed vulture gyps africanus and the effects of anthropogenic disturbance’, ibis 151(1), 51–62. http://dx.doi.org/10.1111/j.1474-919x.2008.00878.x beyer, h.l., 2004, hawth's analysis tools for arcgis, viewed n.d., from http://www.spatialecology.com/htools birdlife international, 2013, species factsheet: gyps coprotheres, viewed 01 july 2013, from http://www.birdlife.org birdlife international, 2014a, important bird areas sa088 collywobbles vulture colony, viewed 23 june 2014, from http://www.birdlife.org.za/conservation/iba/iba-directory/184-collywobbles birdlife international, 2014b, important bird areas sa126 pondoland cape vulture colonies, viewed 23 june 2014, from http://www.birdlife.org.za/conservation/policy-and-advocacy/item/381-sa126-pondoland-cape-vulture-colony birdlife south africa, 2013, important birding areas (iba) shapefile january 2013, viewed 15 december 2014, from http://bgis.sanbi.org/iba/project.asp blundell, g.m., maier, j.a.k. & debevec, e.m., 2001, ‘linear home ranges: effects of smoothing, sample size, and autocorrelation on kernel estimates’, ecological monographs 71(3), 469–489. http://dx.doi.org/10.1890/0012-9615(2001)071[0469:lhreos]2.0.co;2 boshoff, a., barkhuysen, a., brown, g. & michael, m., 2009, ‘evidence of partial migratory behaviour by the cape griffon gyps coprotheres’, ostrich 80(3), 129–133. http://dx.doi.org/10.2989/ostrich.2009.80.3.1.964 boshoff, a. & minnie, j., 2011, ‘on the role of the shape and size of foraging area, and colony size, in selecting critical areas for cape griffon gyps coprotheres conservation action’, vulture news 61(1), 4–15. boshoff, a.f., minnie, j.c., tambling, c.j. & michael, m.d., 2011, ‘the impact of power line-related mortality on the cape vulture gyps coprotheres in a part of its range, with an emphasis on electrocution’, bird conservation international 21(3), 311–327. http://dx.doi.org/10.1017/s095927091100013x boshoff, a., piper, s. & michael, m., 2009, ‘on the distribution and breeding status of the cape griffon gyps coprotheres in the eastern cape province, south africa’, ostrich 80(2), 85–92. http://dx.doi.org/10.2989/ostrich.2009.80.2.4.831 boshoff, a.f., robertson, a.s. & norton, p.m., 1984, ‘a radio-tracking study of an adult cape griffon vulture gyps coprotheres in the south-western cape province’, south african journal of wildlife research 14, 73–78. boshoff, a.f. & vernon, c.j., 1980, ‘the past and present distribution and status of the cape vulture in the cape province’, ostrich 51, 230–250. http://dx.doi.org/10.1080/00306525.1980.9633570 botha, a., allan, d., krueger, s., neethling, m., neser, w., pfeiffer, m. et al., 2012, cape vulture task force report 2012: wolter, k., endangered wildlife trust, viewed n.d., from http://www.vulpro.com/publications/cvtf_report_2012_final.pdf brown, c.j. & piper, s.e., 1988, ‘status of cape vultures in the natal drakensberg and their cliff site selection’, ostrich 59(3), 126–136. http://dx.doi.org/10.1080/00306525.1988.9633714 burt, w.h., 1943, ‘territoriality and home range concepts as applied to mammals’, journal of mammalogy 24(3), 346–352. http://dx.doi.org/10.2307/1374834 byers, c.r., steinhorst, r.k. & krausman, p.r., 1984, ‘clarification of a technique for analysis of utilization-availability data’, the journal of wildlife management 48(3), 1050–1053. http://dx.doi.org/10.2307/3801467 diekmann, m., scott, a., scott, m. & diekmann, j., 2004, ‘capture and fitting of satelliteand radio-telemetry equipment onto cape griffon vulture gyps coprotheres, african white-backed vulture gyps africanus and lappet-faced vulture torgos tracheliotus in the waterberg area, namibia in 2004’, vulture news 51(1), 34–45. dupont, h., mihoub, j.b., bobbé, s. & sarrazin, f., 2012, ‘modelling carcass disposal practices: implications for the management of an ecological service provided by vultures’, journal of applied ecology 49(2), 404–411. http://dx.doi.org/10.1111/j.1365-2664.2012.02111.x endangered wildlife trust (ewt) and ezemvelo kwazulu-natal (kzn) wildlife unpublished data, vulture feeding sites. fisher, e.c., albert, r., botha, g., cawthra, h.c., esteban, i., harris, j. et al., 2013, ‘archaeological reconnaissance for middle stone age sites along the pondoland coast, south africa’, paleoanthropology 1(1), 104–137. haas, d. & mundy, p., 2013, ‘vultures revered in time and place’, vulture news 63(1), 4–14. international union for conservation of nature and natural resources (iucn) and united nations environment programme's world conservation monitoring centre (unep-wcmc), 2014, the world database on protected areas (wdpa), cambridge, uk: unep-wcmc, viewed 11 november 2014, from http://www.protectedplanet.net katzner, t.e., brandes, d., miller, t., lanzone, m., maisonneuve, c., tremblay, j.a. et al., 2012, ‘topography drives migratory flight altitude of golden eagles: implications for on-shore wind energy development’, journal of applied ecology 49(5), 1178–1186. http://dx.doi.org/10.1111/j.1365-2664.2012.02185.x kendall, c.j., virani, m.z., hopcraft, j.g.c., bildstein, k.l. & rubenstein, d.i., 2014, ‘african vultures don’t follow migratory herds: scavenger habitat use is not mediated by prey abundance’, plos one 9(1), e83470. http://dx.doi.org/10.1371/journal.pone.0083470 kepe, t., 1997, ‘communities, entitlements and nature reserves: the case of the wild coast, south africa’, ids bulletin 28(4), 47–58. http://dx.doi.org/10.1111/j.1759-5436.1997.mp28004006.x lawes, m.j., griffiths, m.e. & boudreau, s., 2007, ‘colonial logging and recent subsistence harvesting affect the composition and physiognomy of a podocarp dominated afrotemperate forest’, forest ecology and managment 247(1–3), 48–60. http://dx.doi.org/10.1016/j.foreco.2007.04.012 mander, m., diedericks, n., ntuli, l., mavundla, k., williams, v. & mckean, s., 2007, ‘survey of the trade in vultures for the traditional health industry in south africa’, futureworks, everton, viewed n.d., from http://146.141.46.24/craig/raptors/mander%20et%20al%202007%20-%20vulture%20trade%20in%20south%20africa%20-%20report.pdf mckee, j.k., sciulli, p.w., fooce, c.d. & waite, t.a., 2004, ‘forecasting global biodiversity threats associated with human population growth’, biological conservation 115(1), 161–164. http://dx.doi.org/10.1016/s0006-3207(03)00099-5 moleón, m., sánchez-zapata, j.a., margalida, a., carrete, m., owen-smith, n. & donázar, j.a., 2014, ‘humans and scavengers: the evolution of interactions and ecosystem services’, bioscience 64(5), 1–10. http://dx.doi.org/10.1093/biosci/biu034 monadjem, a. & garcelon, d.k., 2005, ‘nesting distribution of vultures in relation to land use in swaziland’, biodiversity & conservation 14(9), 2079–2093. http://dx.doi.org/10.1007/s10531-004-4358-9 monsarrat, s., benhamou, s., sarrazin, f., bessa-gomes, c., bouten, w. & duriez, o., 2013, ‘how predictability of feeding patches affects home range and foraging habitat selection in avian social scavengers?’, plos one 8(1), e53077. http://dx.doi.org/10.1371/journal.pone.0053077 mucina, l., adams, j.b., knevel, i.c., rutherford, m.c., bolton, j.j., van der merwe, j.h. et al., 2006. ‘the vegetation of south africa, lesotho and swaziland’, in mucina, l. & rutherford, m.c. (eds.), strelitzia, 19th ed., pp. 45, 510–511, 579, south african national biodiversity institute, pretoria. mundy, p., butchart, d., ledger, j. & piper, s., 1992, the vultures of africa, acorn books, randburg, pretoria. mundy, p.j., 1982, the comparative biology of southern african vultures, vulture study group, johannesburg. murn, c. & anderson, m.d., 2008, ‘activity patterns of african white-backed vultures gyps africanus in relation to different land-use practices and food availability’, ostrich 79(2), 191–198. http://dx.doi.org/10.2989/ostrich.2008.79.2.9.583 ogada, d.l., keesing, f. & virani, m.z., 2012a, ‘dropping dead: causes and consequences of vulture population declines worldwide’, annals of the new york academy of sciences 1249(1), 57–71. http://dx.doi.org/10.1111/j.1749-6632.2011.06293.x ogada, d.l., torchin, m.e., kinnaird, m.f. & ezenwa, v.o., 2012b, ‘effects of vulture declines on facultative scavengers and potential implications for mammalian disease transmission’, conservation biology 26(3), 1–8. http://dx.doi.org/10.1111/j.1523-1739.2012.01827.x phipps, w.l., willis, s.g., wolter, k. & naidoo, v., 2013a, ‘foraging ranges of immature african white-backed vultures (gyps africanus) and their use of protected areas in southern africa’, plos one 8(1), e52813. http://dx.doi.org/10.1371/journal.pone.0052813 phipps, w.l., wolter, k., michael, m.d., mactavish, l.m. & yarnell, r.w., 2013b, ‘do power lines and protected areas present a catch-22 situation for cape vultures (gyps coprotheres)?’, plos one 8(10), e76794. http://dx.doi.org/10.1371/journal.pone.0076794 piper, s.e., 1994, mathematical demography of the cape vulture, phd thesis, university of cape town, cape town. piper, s.e., 2005, ‘cape vulture’, in hockey, p.a.r., dean, w.r.j. & ryan, p.g. (eds.), roberts birds of southern africa, 7th edn., pp. 489–491, john voelcker bird book fund, cape town. piper, s.e., boshoff, a.f. & scott, h.a., 1999, ‘modelling survival rates in the cape griffon gyps coprotheres, with emphasis on the effects of supplementary feeding’, bird study 46, 230–238. http://dx.doi.org/10.1080/00063659909477249 piper, s.e. & ruddle, p., 1986, ‘an initial evaluation of the cape vulture colonies at mkambati, transkei’, vulture news 15(1), 7–12. powell, r.a., 2000, ‘animal home ranges and territories and home range estimators’, in boitani, l. & fuller, t.k. (eds.), research techniques in animal ecology: controversies and consequences, 2nd edn., pp. 79–80, columbia university press, new york. prakash, v., pain, d.j., cunningham, a.a., donald, p.f., prakash, n., verma, a. et al., 2003, ‘catastrophic collapse of indian white-backed gyps bengalensis and long-billed gyps indicus vulture populations’, biological conservation 109(3), 381–390. http://dx.doi.org/10.1016/s0006-3207(02)00164-7 retief, e.f., diamond, m., anderson, m.d., smit, h.a., jenkins, a., brooks, m. et al., 2013, avian wind farm sensitivity map for south africa: criteria and procedures used. robertson, a.s. & boshoff, a.f., 1986, ‘the feeding ecology of cape vultures gyps coprotheres in a stock-farming area’, biological conservation 35(1), 63–86. http://dx.doi.org/10.1016/0006-3207(86)90027-3 rodgers, a.r., carr, a.p., beyer, h.l., smith, l. & kie, j.g., 2007, home range tools (hrt) for arcgis: version 1.1, ontario ministry of natural resources, centre for northern forestry ecosystem research, thunder bay, ontario, canada. ruxton, g.d. & houston, d.c., 2002, ‘modelling the energy budget of a colonial bird of prey, the ruppell's griffon vulture, and consequences for its breeding ecology’, african journal of ecology 40(3), 260–266. http://dx.doi.org/10.1046/j.1365-2028.2002.00368.x south african national botanical institute (sanbi), 2000, national land cover database, pretoria, south africa. shackleton, c.m., granger, j.e., mckenzie, b. & mentis, m.t., 1991, ‘multivariate-analysis of coastal grasslands at mkambati game reserve, north-eastern pondoland, transkei’, bothalia 21(1), 91–107. shackleton, r., shackleton, c., shackleton, s. & gambiza, j., 2013, ‘deagrarianisation and forest revegetation in a biodiversity hotspot on the wild coast, south africa’, plos one 8(10), 1–12. http://dx.doi.org/10.1371/journal.pone.0076939 sheehan, d.k. & sanderson, f.j., 2012, ‘seeing the bigger picture: how anthropogenic landscape modification in africa affects declining migratory birds and the need for trans-continental research and conservation’, ibis 154(4), 659–662. http://dx.doi.org/10.1111/j.1474-919x.2012.01275.x skead, c.j., 1987, ‘the eastern half of the cape province, including the ciskei, transkei and east griqualand’, in skead, c.j. (eds.), historical mammal incidence in the cape province, pp. 80–82, provincial administration of the cape of good hope, department of nature conservation, cape town. spiegel, o., harel, r., getz, w. & nathan, r., 2013, ‘mixed strategies of griffon vultures’ (gyps fulvus) response to food deprivation lead to a hump-shaped movement pattern’, movement ecology 1(5), 1–12. http://dx.doi.org/10.1186/2051-3933-1-5 statistics south africa, 2011, population per municipality, viewed 05 november 2012, from http://beta2.statssa.gov.za/ statsoft, 2006, statistica (data analysis software system): version 7.1, http://www.statsoft.com swinhart, r. & slade, n., 1985, ‘influence of sampling interval on estimates of home-range size’, the journal of wildlife management 49(4), 1019–1025. http://dx.doi.org/10.2307/3801388 vernon, c., 1998, ‘the cape vulture at collywobbles [sic.]: 1977–1997’, ostrich 70(3–1), 200–202. vetter, s. & bond, w.j., 2012, ‘changing predictors of spatial and temporal variability in stocking rates in a severely degraded communal rangeland’, land degradation & development 23(2), 190–199. http://dx.doi.org/10.1002/ldr.1076 appendix 1 top ↑ figure a1: land use map used for habitat analysis detailing the political boundaries of the former transkei and lesotho. figure a2: incremental area analysis of minimum convex polygons of adult cape vultures (gyps coprotheres) in relation to number of global positioning system location fixes for (a) the non-breeding season and (b) the breeding season, indicating that foraging range asymptotes were reached during each season. table a1: land use map reclassified categories. original land use categories were based on the 2000 south african national land cover database (south african national botanical institute 2000). article information authors: jennifer strickland-munro1 susan moore1 affiliations: 1school of environmental and life sciences, murdoch university, australia correspondence to: jennifer strickland-munro postal address: south st, murdoch wa 6150, australia dates: received: 27 mar. 2013 accepted: 01 nov. 2013 published: 24 june 2014 how to cite this article: strickland-munro, j. & moore, s., 2014, ‘exploring the impacts of protected area tourism on local communities using a resilience approach’, koedoe 56(2), art. #1161, 10 pages. http://dx.doi.org/10.4102/ koedoe.v56i2.1161 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. exploring the impacts of protected area tourism on local communities using a resilience approach in this original research... open access • abstract • introduction    • protected area tourism       • tourism impacts • research method and design • results    • phase 1: system definition    • phase 2: past system change    • phase 3: current system state • ethical considerations    • potential benefits and hazards    • recruitment procedures    • informed consent    • data protection • discussion    • nature conservation    • park access and visitation    • employment • conclusion • acknowledgements    • competing interests    • authors&rsquo; contributions • references • footnotes abstract top ↑ as the protected area mandate expands to include social equity, the impacts of parks and their tourism on neighbouring indigenous and local communities is receiving growing practical and theoretical interest. this article reported on one such study, which explored the impacts of protected area tourism on communities bordering the iconic kruger national park in south africa and purnululu national park in australia. the study drew on interviews with park staff, tourism operators and community members. guided by a conceptual framework grounded in resilience thinking, interactions amongst the parks, tourism and local communities were revealed as complex, contested and multi-scalar. underlying drivers included cultural norms and values based on nature, entrenched poverty, poor western education and economic opportunities associated with tourism. park tourism offered intrinsic opportunities and benefits from nature conservation and associated intangible cultural values. more tangible benefits arose through employment. damage-causing animals and visitation difficulties were negative impacts. interaction with tourists was limited, with a sense of disconnect evident. findings indicated the need for multifaceted, carefully considered policy responses if social equity and benefits for local communities are to be achieved. framing the impacts of protected area tourism through the resilience framework provided a useful way to access local community perceptions whilst retaining awareness of the broader multi-scalar context in which interactions occur. conservation implications: perceptions of separation and lack of education to engage in economic opportunities are major issues. intrinsic appreciation of parks is an important platform for building future opportunities. accrual of future benefits for local communities from park tourism depends on developing diverse economic opportunities, building community capacity and managing expectations and addressing economic disadvantage. introduction top ↑ protected areas and tourism have a longstanding and mutually beneficial relationship (bushell, staiff & eagles 2007; reinius & fredman 2007). such areas offer a unique, desirable setting for the tourism product, whilst tourism offers park managers a revenue source that can assist ongoing conservation efforts. beyond this mutual association, protected area tourism can have considerable benefits for surrounding communities, as well as social costs (e.g. increased competition for resources and facilities). whether the opportunities provided by tourism benefit communities or not is debatable and contingent on a number of interacting factors. increasingly, protected area tourism is being recognised as a complex social-ecological system, with interactions characterised by dynamism, complexity, uncertainty, unpredictability and multi-scalar connections (mccool 2009; strickland-munro, allison & moore 2010) (figure 1). local communities are an integral part of this system, as described by strickland-munro et al. (2010) using a resilience approach. unpredictable and susceptible to external influences beyond local control, a number of barriers can obstruct the delivery of benefits to local communities and confound interactions between protected areas, tourism and local communities. a resilience approach, which acknowledges the importance of understanding interactions between system components, provides a promising alternative to reductionist analyses. a broader, interdisciplinary approach cognisant of multiple interacting variables and perspectives allows exploration of the complex relationships amongst system components (farrell & twining-ward 2005). resilience thinking offers one such framework and is particularly suited to protected area tourism’s inherently ‘messy’ environment of complexity, change and uncertainty (mccool 2009; plummer & fennell 2009). resilience perspectives focus on understanding the dynamics of systems interactions and explicitly acknowledge system uncertainty, complexity and dynamism (resilience alliance 2007a, 2007b; walker et al. 2006). published applications of resilience perspectives to protected area tourism however have been limited to date. this article applies a conceptual framework based on resilience thinking to an exploration of the impacts of tourism associated with protected areas on local communities. the framework was used for local communities adjacent to kruger national park in south africa and purnululu national park in australia. specifically, the study sought to ascertain key socio-economic impacts of protected area tourism on the local communities. local perspectives were privileged, whilst the broader context was accommodated through the framework’s attention to multi-scalar influences and interactions (figure 1). we first outline the role of protected area tourism in south africa and australia, followed by a brief synopsis of the tourism impact literature and the growing demand for park tourism to address social equity and deliver benefits to local communities. the research design and methods follows, with the first part of this section devoted to an overview of the resilience framework guiding this research. the two case study areas are then described as part of the comprehensive description of the case studies as protected area–tourism systems. the article’s conclusion includes implications for park and tourism managers. here we present one of the first published applications of resilience perspectives to protected area tourism and the investigation of such impacts on local communities. the combination of resilience perspectives with ethnographic research provides a synthesised, coherent approach facilitating in-depth insights into local impacts. figure 1: multi-scalar nature of protected area–tourism systems. protected area tourism protected area tourism fills a significant niche in both south africa and australia; this provides the initial rationale for selecting case studies in the two countries. boasting diverse scenery and wildlife (cornelissen 2005; worboys, lockwood & de lacy 2005), the respective protected areas of these countries offer extensive tourism opportunities and experiences. visitation to protected areas in both countries continues to grow, providing valued revenue. in south africa, visitation to protected areas grew by 0.5% from 2009 arrivals, to 4 536 491 arrivals in 2010–2011. associated revenue for this period was r815.26 million (approx. $99.09 million), an increase of 12.1% from the previous year (south african national parks [sanparks] 2011). in australia, nature tourism (based on parks and visits to zoos, aquaria and botanic gardens) contributes an estimated aus$23 billion (approx. $23.85 billion) annually to the economy (tourism research australia 2009). beyond this broad economic impact, protected area tourism is further valued for its touted ability to foster smaller scale socio-economic development and employment (telfer & sharpley 2008). however, these socio-economic benefits often co-exist alongside negative impacts for the communities adjoining protected areas (wall & mathieson 2006). given links between benefit delivery and positive local perceptions of protected areas and tourism (e.g. brockington, duffy & igoe 2008), research into the impacts of park tourism on local communities is of considerable interest. kruger national park and purnululu national park offered promising case studies as they are both iconic, with communities living adjacent to them which have opportunities to engage in tourism but continue to face entrenched economic disadvantage. tourism impacts tourism impacts have been extensively documented (e.g. wall & mathieson 2006). we confine our synopsis to impacts relevant to protected area tourism and local communities. the tangible economic impacts, notably employment opportunities, dominate the literature. employment provides a direct economic link between local communities and park tourism and is recognised as one of tourism’s main development advantages (e.g. simelane, kerley & knight 2006). this development role stems from the generation of both direct (e.g. roles in park or tourism operations) and indirect job opportunities (e.g. derivation of income from related activities such as curio sales), both of which can raise local incomes and living standards (mill & morrison 2006). the need for such economic development via tourism is well recognised in south africa, where poverty is endemic (binns & nel 2002). its necessity is often overlooked in australia, where, in reality, living standards and economic development for many indigenous australians remain significantly below that of non-indigenous australians (altman 2007). tourism can also have negative economic impacts. these include dependency and stifling of economic diversification (telfer & sharpley 2008), as well as inequality in benefits (simpson 2009; trau & bushell 2008). inequity of benefit distribution is heightened by ‘enclave’ tourism, where tourists and local communities are separated by boundaries (e.g. park fences), with community interactions typically limited to transient hawking at park entrances. enclave tourism restricts local access to tourism markets and can also contribute to financial leakage (mill & morrison 2006). novelli and scarth (2007), for example, report that tourists to malawi’s liwonde national park remain within park boundaries, limiting opportunities for interactions with locals. this distancing was linked to a lack of ownership for the park. furthermore, tourism’s financial benefits are often overstated, leading to false expectations (simpson 2008). other negative economic impacts include lost opportunity costs, where tourism dominates resources that could have been more profitably used in other sectors (mill & morrison 2006) such as agriculture. socio-cultural impacts include changes to individual behaviour, value systems, cultural practices and community organisations (andereck et al. 2005). tourism can enhance ethnic identity, increase understanding between cultures, challenge stereotypes and perceptions and revitalise cultural values and traditions (telfer & sharpley 2008), including the use of natural resources. changes to decision-making structures and involvement in park management are further positive impacts (scherl & edwards 2007). enrichment of local skills, local and institutional capacity building, investments in community infrastructures, such as schools and health care, and improved access to information and external environments are further positive impacts. negative socio-cultural impacts include the loss or misuse of cultural resources, perceptions of cultural commoditisation and exploitation, ‘demonstration effects’ where locals adopt tourist behaviours and the marginalisation of locals to jobs of lesser importance (telfer & sharpley 2008). tourism may also cause culture shock, communication difficulties, tension, suspicion and hostility, segregation and perceptions of superiority or inferiority (robinson 1999). in recent years, social equity and the delivery of benefits to local communities have assumed a greater role in protected areas (phillips 2003; worboys et al. 2005). this is certainly true for park managers in the two protected areas of interest to this study, where official attention is directed towards redressing social and economic inequities facing previously disadvantaged communities. south africa’s kruger national park emphasises park–people relations and benefit sharing and seeks to increase the park’s legitimacy amongst neighbouring communities. tourism, and the benefits it can engender, is a key means of achieving these goals (sanparks 2008). similarly, managers of australia’s purnululu national park are guided by an official commitment to ensure indigenous engagement and benefit from park and tourism operations (department of environment and conservation [dec] 2008). however, both parks evidence a dearth of social research, with research on the impacts of park tourism on local communities scarce, despite calls for such from park managers and other interested parties (e.g. sanparks 2008; unesco 2010). the study reported in this article addresses this deficit. research method and design top ↑ research was guided by strickland-munro et al.’s (2010) conceptual framework for investigating the relationship between protected area tourism and local communities and was grounded in resilience thinking (resilience alliance 2007a, 2007b). four interdependent stages of investigation, described in detail in strickland-munro et al. (2010), were modified to comprehensively describe the protected area–tourism system and scale of analysis (figure 2, phase 1), followed by a detailed description of past system changes and current state of the system (figure 2, phases 2 and 3). in all four phases, socio-political aspects, and especially governance, have been emphasised in contrast to much resilience thinking, which has a strong ecological emphasis and often downplays these aspects (miller et al. 2010). phase 1 (figure 2) concerns defining the system of interest – interactions between the national parks, their tourism and associated local communities. investigation prioritised local perspectives, given that the study’s central interest was impacts on local communities. we focused on the local scale, geographically defined ‘community’ and prioritised social over ecological interactions. the views of other stakeholders at local, sub-national and national scales were included, however, to incorporate interests beyond the local (strickland-munro et al. 2010), with such a multi-scalar approach being central to resilience analyses (resilience alliance 2007a, 2007b). phase 2 (figure 2) explores past system change, focusing on system drivers and historical influences affecting current conditions. understanding these drivers and underlying processes influencing system change is necessary to develop appropriate management responses and is central to resilience analyses (resilience alliance 2007a, 2007b). typically, drivers are grouped into categories, including socio-political, demographic, economic and biophysical, and classified as either ‘slow’ or ‘fast’ depending on the rate at which they influence change. slow variables play an important role in controlling the protected area–tourism system (strickland-munro et al. 2010) and influence well-being and governance (kofinas & chapin 2009), whereas fast variables tend to operate at smaller spatio-temporal scales and influence system interactions at a faster rate. although the adaptive cycle heuristic is integral to this phase it was not considered further in this article, given its focus on the dynamics amongst fast and slow variables over time and scale, rather than giving explicit attention to impacts. phase 3 (figure 2) focuses on the state of the current system and explored key issues, such as those relating to values, benefits, conflicts, challenges and governance. phase 4 (figure 2) rests on monitoring changes in how protected areas, tourism and local communities interact. the final phase presented in this article differs from strickland-munro et al. (2010), as a lack of empirical information from the case studies on interactions meant that the threshold development advocated could not be undertaken. for this reason, this final phase was modified to recommend exploring future scenarios and developing indicators, both with the ultimate objective of further understanding interactions and eventually developing thresholds. this article reported on the first three phases, with the aim of providing a thorough analysis of the impacts of protected area tourism on local communities. future scenarios and indicator development, the fourth phase, are not included as they are less central to understanding impacts. for simplicity, a description of the impacts of governance is included in the second phase, although it is evident in all phases. site selection was guided by three criteria, (1) the sites are iconic protected areas, (2) the protected areas have high tourism visitation and (3) they are neighboured by disadvantaged communities, that is, those with high unemployment, low incomes and poor education and health outcomes. information was obtained in this ethnographic study through semi-structured interviews with local community members, park and tourism managers and staff, as well as government and non-government operatives. this wide range of people was selected to obtain the broadest possible worldview of how these local communities are experiencing tourism associated with their adjacent national park. new individuals were interviewed until ‘theoretical saturation’ (bowen 2008) was reached, meaning that the particular interview did not add any new information. in this study, theoretical saturation was reached at 110 interviews (92 at kruger, 18 at purnululu). the interview questions covered perceived benefits, barriers to participation and other factors influencing the engagement of local communities with protected area tourism. these questions were asked to populate the conceptual framework in figure 1 with information. interviews were recorded where possible and transcribed verbatim, with trained local translators used in the south african study. analysis was undertaken using grounded theory, an inductive technique that generates theory from constant comparison of themes and patterns in interview transcripts. the code-based theory development software nvivo (qsr international 2008) facilitated analysis and assisted in the iterative refinement of conceptual codes. these codes, together with information gathered from document review and participant observation, informed the following results. interview excerpts are provided to help describe these codes in the following section, a standard practice in reporting the results of grounded theory research. review of historical and current literature regarding protected areas, tourism and local communities also contributed information. figure 2: conceptual framework for investigating the relationship between protected area tourism and local communities. results top ↑ phase 1: system definition both parks in this protected area–tourism system are of iconic status: kruger is world-renowned and acknowledged as the flagship of south african national parks (sanparks 2011), whilst purnululu is listed a world heritage site for its natural values (unesco 2010). beyond these iconic ties, the parks differ widely. kruger dominates in physical size and scale of tourism, covering approximately 2 million ha and receiving 1.45 million tourists annually (sanparks 2013), in contrast to purnululu’s 208 723 ha and current visitation of 24 602 (dec 2012). operationally, kruger has been in existence since 1926, whereas purnululu’s existence as a national park is much more recent, being formally gazetted in 1987 (department of conservation and land management [calm] 1995). this disparity is reflected in kruger’s well developed tourist infrastructure, including 26 tourist rest camps with varied accommodation options and the broad range of visitor activities (sanparks 2008). purnululu, however, has purposely adopted a minimalist approach to tourism development and offers only two public campsites with basic facilities. several private camps offer a higher standard of accommodation. at the time of research, kruger’s workforce was 2116 permanent and temporary workers (urban-econ 2008), compared to purnululu’s complement of two permanent rangers and two indigenous assistant rangers employed under government welfare schemes.both parks are accessible via road and air links and cater to self-drive visitors and group tours. however, kruger’s accessibility is far superior to purnululu, particularly for road access, with purnululu’s unsealed 52 km entrance track requiring considerable skill and a suitable four-wheel drive vehicle to navigate. additionally, purnululu is affected by limited seasonal accessibility, with its open period restricted to april–december on account of the east kimberley wet season (calm 1995). kruger remains open for business year round. in terms of local communities, the parks offer significant disparity as well as surprising analogies. kruger is surrounded by densely populated ex-bantustan communities, with the park’s southern and western borders1 home to over two million people (department of water affairs and forestry [dwaf] 2008). this research focused on two villages, cork and belfast, located closest to one of kruger’s busiest entrance gates. with a combined population of approximately 15 000 residents, their location close to the park offers opportunities for interactions with passing tourists. purnululu, in contrast, is located in australia’s sparsely populated east kimberley region, with a population density of 0.08 people/km2 (kimberley development commission n.d.). the indigenous community of warmun, with a population around 300 (australian bureau of statistics 2006), is the park’s closest neighbour despite being located 50 km north-west of the park. it was included in this study on account of this geographic proximity, as well as status at the time of research as ‘gateway’ community providing the last service facilities (e.g. petrol station, tourist accommodation and general store) available before entering the park. in addition, many indigenous residents retain strong cultural ties to the purnululu landscape. whilst populations and locations of the parks differ, their neighbouring communities are characterised by a number of similarities that can be linked to entrenched disadvantage: in south africa, the legacies of apartheid (sartorius et al. 2009) and, in australia, legacies of socio-political and economic marginalisation of indigenous people (altman 2009). these legacies are evidenced by limited educational attainment, widespread poverty and unemployment. warmun is characterised by welfare dependency (doohan 2008), with 20.0% of the community receiving government benefits in 2007 (warmun community 2007). in the bushbuckridge local municipality, which includes cork and belfast, unemployment often exceeds 90.0% and 84.0% of the population is classified as ‘indigent’, earning less than r1300 ($169) per household per month (dwaf 2008). educational statistics further illustrate community disadvantage. in warmun, 22.0% of adults have never attended school (taylor 2003). similarly, only 1.7% of people residing in bushbuckridge have received an education beyond high school level (statistics south africa 2007). phase 2: past system change four drivers, three of them slow variables, had a significant influence on how the parks, tourism and local communities interact: poverty, poor (western) education levels and cultural norms and values based on a respect for nature. these slow variables exert a controlling influence over current system interactions. economic opportunity associated with tourist arrivals was a fast variable, and the fourth driver, and potentially able to overwhelm the influence of the slower drivers. each of these drivers influenced park tourism impacts. these drivers are largely beyond the control of local decision-makers, as they represent fundamental, higher-scale processes not easily amenable to change by park managers. the study communities are clearly characterised by disadvantage and underdevelopment. poverty and poor (western) education, for instance, reflect the legacies of past policies and practices of discrimination. their influence is evident in the high incidence of unemployment in both study communities, as well as the entrenched welfare dependence in warmun. in the communities around kruger, there have been grant systems (e.g. for child support), a focus on education and economic development initiatives; however, poverty and disadvantage are still evident. together, these two slow drivers were described by respondents as significant barriers to the ability of local communities to engage with park tourism. the third slow variable, cultural norms and values based on a respect for nature, was particularly evident in the purnululu study. indigenous respondents continually referenced enduring links between traditional culture and the physical landscape, emphasising the fundamental influence of such cultural norms on indigenous worldviews. in the kruger study, the influence of cultural norms on local interactions with park tourism was unexpected yet clearly manifest in community responses. respondents expressed a distinct respect for, and appreciation of, the natural environment and suggested that caring for such was integral to their way of life. governance issues related to power inequities and conflict over resources such as cultural sites, park access and natural resources were also a central concern for respondents in discussing park-tourism-community interactions. importantly, these governance elements were not fundamental determinants of interactions. rather, they were themselves influenced by underlying drivers, such as a lack of the western skills or education necessary for operating in a formal management environment, as identified in the purnululu study. this vulnerability to drivers beyond the control of park managers means that governance is considered a system characteristic rather than a driver. phase 3: current system state three key issues are most pertinent to the impacts of park tourism on local communities: nature conservation and intrinsic values, park access and visitation, and employment. community members were highly appreciative of links between park tourism and the conservation of ‘nature’. a range of intrinsic benefits were derived which appeared to foster positive attitudes towards the parks and tourism. in the kruger study, benefits centred on existence, aesthetic and bequest values. respondents reported personal benefits linked to opportunities to view natural landscapes and iconic wildlife species, as illustrated by the following quotation: ‘the existence of [kruger] itself i think people are benefitting because when they go there, they will see what they haven’t seen outside’ (community member 73, male).in contrast, respondents in the purnululu study emphasised spiritual significance, although bequest values were also noted in relation to the transmission of cultural knowledge: ‘traditional country ... culturally … gives people their identity, their family’s identity, the connections, the stories, the beliefs ... country is very important ... [it] helps to define you as a person and to an extent your lifestyle and our values.’ (community member 8, female) (see strickland-munro & moore 2013:32) this intrinsic appreciation of nature derives from, as well as contributes to, the identified slow driver of underlying cultural norms and values based on respect for nature. the conservation of nature provided further (contested) benefits. these included the use and sale of natural resources such as fuel wood and medicinal plants. respondents associated with kruger linked this benefit to the ever-present demands of poverty and subsistence lifestyles: ‘people see kruger as a resource centre, where they can tap in for anything they want’ (park staff 5, female). community members were resentful of restrictions placed upon resource use by park managers. a further contestation relevant to kruger only was animals ‘escaping’ from the park. this key park tourism resource can cause significant destruction and sometimes deaths in local villages, negative impacts which can decisively influence local views of the park and tourism associated with it. these negative impacts are compounded by complex, multi-stakeholder governance arrangements that restrict locals from dealing with escapee animals themselves: ‘villagers are not allowed to defend their property or commercial interests from the animals’ (tour operator 7, male). in many instances, however, community members appeared unwilling to discuss such animals in negative terms. instead, they highlighted their appreciation of the park and nature. benefits derived from the meat from escaped animals provide another explanation: ‘elephants and the other animals, they are not a problem because we get meat when [kruger] kills them’ (community member 23, male). park access and visitation was a second key issue. typically, community members lacked the physical and financial ability to visit their respective parks. in the kruger study, locals rarely visited the park for pleasure. most repeat visits were attributed to past or current employment or park outreach programmes. in the purnululu study, all community respondents had visited the park previously but described significant difficulties in doing so. poverty was central to these difficulties. transport issues were the key limitation in both case studies, whereas kruger respondents additionally cited the requirement to pay entrance fees.2 community members described these barriers in the following ways: ‘i do like to go to purnululu, but how am i going to get there?’ (community member 5, female). ‘i’ve never been inside kruger. i’ve got no money to go in the gate’ (community member 17, female). in both studies, access difficulties were linked to perceptions of disconnect from the parks and tourism: ‘the main purpose of kruger is to conserve nature for the tourist to come and experience’ (community member 52, female). employment in park tourism posed a third key issue and one directly related to the driver of ‘economic opportunities associated with tourist arrivals’. employment occurred in two ways: direct jobs in park management or tourism operations and indirect employment involving the derivation of income from business or activities based on the parks. direct employment was limited in both studies. in the purnululu study, only one local community member was employed in park operations at the time of research. no community members were employed in tourism operations. in kruger, nine community members were directly employed in varied park and tourism operations. such employment was highly valued by all community members on account of its ability to provide for family dependants and educational needs. however, a lack of skills required to successfully engage in formal park and tourism roles precluded widespread benefit accrual, with benefits typically limited to those community members with the requisite skills and knowledge: ‘it takes a fair bit of experience, knowledge and intent … perhaps there aren’t those people in warmun’ (community member 10, male). indirect employment associated with park tourism was more prominent than direct employment in both studies. at purnululu, one community member was employed at the warmun roadhouse and a further four received income by selling indigenous art. home to a major art movement, a growing number of locals produce art for the warmun art gallery. although intermittent, income derived from sales can be substantial, often reaching tens of thousands of dollars. artists receive 60% of the sale price, with 40% retained by the gallery. art production also provided significant intrinsic cultural benefits relating to maintenance of cultural connections to the country. at kruger, 11 community members were indirectly employed in park tourism through the selling goods at roadside stalls. economic benefits derived from this activity were tempered by variability in tourist patronage: ‘if tourists weren’t driving up and down the road, those curio sellers wouldn’t be there’ (tour operator 4, female). in both studies, respondents described the interactions with tourists as limited. this limited interaction further contributed to a local disconnect from park tourism. ethical considerations top ↑ approval for this research was granted by the murdoch university human research ethics committee, permit no. 2007/153. approval was also granted by the dec’s and conservation’s nature based tourism and recreation research group, as well as the sanparks research board. potential benefits and hazards no risks to interviewees were foreseen. all interviewees were guaranteed anonymity of data collected.  recruitment procedures participation in the research was voluntary and interviewees were able to withdraw from the interview at any stage. feedback was provided to interviewees at the end of the research process. informed consent interviewees were required to sign a consent form indicating their willingness to participate in the research. data protection interview data was securely stored on the password-protected computer of the first author. in accordance with ethical requirements, all data are securely kept for 7 years. discussion top ↑ community benefits are obviously derived from tourism associated with the two case study parks. these exist, however, alongside negative and contested impacts that highlight the complexity of interactions between park tourism and local communities. the findings illustrate the influence of underlying fast and slow drivers on impacts experienced by the communities. the roles of historical contingency and cross-scale influences in determining current conditions are particularly evident. nature conservation community members derived clear intrinsic benefits from the natural park environments. this was expected in the australian context, where links between the physical environment and indigenous traditions are well established (doohan 2008). its presence in the kruger study however was unexpected. the history of exclusion and separation associated with african conservation and south africa’s apartheid regime (garland 2008; kepe 2009) foster assumptions of an inherent disconnect. the results clearly refute this assumption. instead, intrinsic benefits appear to provide local community members with a sense of stewardship for the park and its tourism resources. this appreciation of intrinsic benefits challenges the wider african and broader global literature, which rarely highlights the significance of non-pecuniary benefits (coad et al. 2008; shackleton et al. 2008).use of natural resources and damage-causing animals were more tangible, and highly contested, impacts arising from nature conservation (and hence indirectly related to tourism) in the kruger study. the use of natural resources by local communities appeared influenced by two key slow variables. poverty, in terms of enduring livelihood pressures was central, indicating the ongoing influence of historic legacies. cultural norms associated with respecting nature was a second slow variable influencing natural resource use. locals linked their ability to harvest medicinal plants to intrinsic and tangible benefits associated with the maintenance of cultural traditions and knowledge, although, again, poverty has fostered contestation and resource demands that park managers are unable to meet without causing significant ecological damage (l. swemmer [sanparks] pers. comm., 12 september 2010). impacts associated with damage-causing animals challenge the intrinsic values attributed by community members to kruger, fostering negative views towards the park and its tourism. these negative views accord with other research linking such animals with reduced local support for protected areas (e.g. madden 2004). this issue clearly illustrates the complexity of interactions between kruger national park and local communities and the impact of this issue on benefit perceptions. the socio-political realities that characterise community members in terms of poverty and poor western education levels challenge their ability to dispute existing institutional arrangements governing management of damage-causing animals, to the ongoing detriment of park–people relations. park access and visitation community members were constrained in their ability to visit the parks for pleasure. transport and monetary constraints were prominent barriers to further visitation, as commonly found in other research (e.g. brown 2009; simelane et al. 2006). both barriers reflect the constraints imposed by entrenched poverty on local ability to benefit from park tourism. in kruger, access difficulties precluded opportunities to experience the park as a ‘tourist’, a dichotomy wherein locals view their role as restricted to serving tourist needs rather than holding the right to enjoy kruger as other people do. in purnululu, access difficulties reduced community ability to maintain intrinsic benefits in terms of cultural connections to country. in both studies, access difficulties fostered perceptions of disconnect from the parks, with an associated lesser appreciation of benefits. visitation has been described as a privilege for rich (typically white) tourists (see also novelli & scarth 2007; simelane et al. 2006). for kruger, this disparity is changing, with 29% of the 1 132 564 south african residents visiting this park, as of april 2013, being black (sanpark 2013). employment employment in park tourism presented a central point of benefit and contestation in both case studies. clear benefits were realised and greatly appreciated by community members. such benefits were limited in distribution to a minority of community members, as commonly reported in the wider literature (e.g. simpson 2009; trau & bushell 2008). it appears that whilst tourist arrivals bring potential for economic benefit, the underlying contexts of widespread poverty and poor western education limit the ability of such benefits to permeate communities. these slow drivers restrict the capacity of community members to procure employment or start businesses to tap into tourism benefits. these findings contribute to the growing body of literature contesting views of protected areas and tourism as ‘money spinners’ generating widespread financial benefits for local communities (brockington et al. 2008). pervasive expressions of disconnect from park tourism evident from the interviews provide further support. in contrast, employment in the purnululu study (both direct and indirect) provided intrinsic cultural benefits relating to connections to country. thus again, interactions between park tourism and local communities are seen as contested and complex in terms of benefit provision and influenced by historic and ongoing legacies that fundamentally shape current conditions and benefit accrual. conclusion top ↑ this study has provided in-depth insights into how local communities interact with, and benefit from park tourism in south africa and australia. research was guided by strickland-munro et al.’s (2010) resilience framework for investigating the relationship between park tourism and local communities. findings reveal a complex and contested picture whereby benefit derivation is juxtaposed against negative impacts. benefit perceptions were influenced by four key underlying drivers: poverty, poor (western) education levels, cultural norms and values based on a respect for nature, and economic opportunities associated with tourist arrivals. although significant contextual differences exist, these drivers were common to both case studies and exerted a decisive influence over local perceptions and interactions with park tourism. however, they are multi-scalar in origin and largely beyond the ability of local decision-makers to control. their influence on perceptions of benefit was explored through three key issues identified by respondents: nature conservation, access difficulties and employment in park tourism. an important opportunity for local communities to benefit from park tourism clearly centres on tangible economic benefits from employment, as commonly reported in the literature. in both studies, constraints are evident and reflect the ongoing influence of historical legacies of discrimination and marginalisation, entrained in the underlying slow variables of poverty and poor western education. as such, caution is needed in advocating benefit opportunities, so as not to stimulate unrealistic community expectations. simultaneously, park and tourism managers must recognise the influence of historic legacies on the ability of community members to engage with available opportunities, working with communities over extended periods of time to build local capacity for involvement. evidence of intrinsic socio-cultural benefits derived from opportunities to experience ‘nature’ is an important finding. such benefits were anticipated in the australian context, yet unexpected in the kruger study, where their identification challenges common assumptions regarding non-pecuniary benefits. the often-undervalued presence of cultural norms and values respecting nature potentially offers park and tourism managers a promising and enduring platform for beneficial interactions between park tourism and local communities. the complex nature of the relationships between communities and nature conservation in the parks is illustrated by contestations over benefits and impacts associated with the use of natural resources and damage-causing animals at kruger. a second important finding arising from the research was the feeling of disconnect between community members and the parks and their tourism. this disconnect likely stems from negative impacts associated with park tourism, such as access difficulties and associated views of park tourism as an activity reserved for rich or white tourists rather than locals. cross-scale influences again appear to contribute, in the form of entrenched poverty and poor western education, which, in turn, represent the legacies of policies and practices in south africa and australia. pragmatic policy responses that seek to foster connections between locals and park tourism, for instance alternative methods of facilitating visitation and, in kruger, opportunities to experience the park as a ‘tourist’, provide one means of reducing local disconnect. in closing, a few comments on the conceptual framework are warranted. the framework proved valuable in accessing both local perspectives and multi-scalar influences, allowing the development of nuanced insights into the complexity of park tourism impacts on local communities. using an ethnographic approach provided valuable insights to the local communities; however, its combination with document reviews and interviews with stakeholders at broader scales was essential given the location of drivers beyond the local focal system. applying this novel methodology to two iconic national parks and their communities has revealed the opportunities and challenges in progressing the well-being of these communities and their parks. this resilience approach has importantly exposed the complexity of such systems and through such exposure made an enhanced understanding and well-informed actions possible. acknowledgements top ↑ this research was supported financially by the sustainable tourism cooperative research centre (established and funded by the australian government), the western australian department of environment and conservation (wa dec) and murdoch university. important and essential in-kind support was provided by wa dec and sanparks. the intellectual and logistical contributions by dr stefanie frietag-ronaldson (sanparks) are gratefully acknowledged, as are the contributions of drs helen allison and david palmer, murdoch university for their respective insights to systems thinking and indigenous ways of knowing the world. daryl moncrieff, wa dec also provided important support for and input to this research. the authors gratefully acknowledge the time and cooperation of respondents involved in this research. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions j.s-m. (murdoch university) was the principal investigator in this study and prepared the manuscript. s.m. (murdoch university) contributed to research design and manuscript refinement. references top ↑ altman, j.c., 2007, alleviating poverty in remote indigenous australia: the role of the hybrid economy, centre for aboriginal economic policy research, australian national university, canberra.altman, j.c., 2009, developing a national indigenous policy framework that recognises needs, rights and legacies and delivers results, centre for aboriginal economic policy research, australian national university, canberra. andereck, k.l., valentine, k.m., knopf, r.c. & vogt, c.a., 2005, ‘”residents” perceptions of community tourism impacts’, annals of tourism research 32(4), 1056–1076. australian bureau of statistics, 2006, housing and infrastructure in aboriginal and torres strait islander communities, australia: warmun community profile, commonwealth of australia, canberra. binns, t. & nel, e., 2002, ‘tourism as a local development strategy in south africa’, the geographical journal 168(3), 235–247. bowen, g.a., 2008, ‘naturalistic inquiry and the saturation concept: a research note’, qualitative research 8(1), 137–152. brockington, d., duffy, r. & igoe, j., 2008, nature unbound: conservation, capitalism and the future of protected areas, earthscan, london. brown, s., 2009, protected landscapes and cultural and spiritual values, iucn protected landscapes task force, gland. bushell, r., staiff, r. & eagles, p.f.j., 2007, ‘tourism and protected areas: benefits beyond boundaries’, in r. bushell & p.j.f. eagles (eds.), tourism and protected areas: benefits beyond boundaries, pp. 1–11, cab international, wallingford. department of conservation and land management, 1995, purnululu national park management plan 1995–2005, calm, perth. coad, l., campbell, a., miles, l. & humphries, k., 2008, the costs and benefits of forest protected areas for local livelihoods: a review of the current literature, unep world conservation monitoring centre, cambridge. cornelissen, s., 2005, the global tourism system: governance, development and lessons from south africa, ashgate publishing limited, aldershot. department of environment and conservation, 2008, 2008–2009 annual report, dec, perth. department of environment and conservation, 2012, ‘purnululu national park visitor data’, unpublished data, dec, perth. department of water affairs and forestry, 2008, bushbuckridge case study, dwaf, durban. doohan, k., 2008, making things come good: relations between aborigines and miners at argyle, backroom press, broome. farrell, b.h. & twining-ward, l., 2005, ‘seven steps towards sustainability: tourism in the context of new knowledge’, journal of sustainable tourism 13(2), 109–122. garland, e., 2008, ‘the elephant in the room: confronting the colonial character of wildlife conservation in africa’, african studies review 51(3), 51–73. kepe, t., 2009, ‘shaped by race: why “race” still matters in the challenges facing biodiversity conservation in africa’, local environment 14(9), 871–878. kimberley development commission, n.d., the kimberley, viewed 23 november 2012, from http://www.kdc.wa.gov.au/the-kimberly/ kofinas, g.p. & chapin iii, f.s., 2009, ‘sustaining livelihoods and human well-being during social-ecological change’, in f.s. chapin iii, g.p. kofinas & c. folke (eds.), principles of ecosystem stewardship: resilience-based natural resource management in a changing world, pp. 55–76, springer, new york. madden, f., 2004, ‘creating coexistence between humans and wildlife: global perspectives on local efforts to address human-wildlife conflict’, human dimensions of wildlife 9(4), 247–257. mccool, s.f., 2009, ‘constructing partnerships for protected area tourism planning in an era of change and messiness’, journal of sustainable tourism 17(2), 133–148. mill, r.c. & morrison, a.m., 2006, the tourism system, 5th edn., kendall/hunt publishing company, dubuque. miller, f., osbahr, h., boyd, e., thomalla, f., bharwani, s., ziervogel, g. et al., 2010, ‘resilience and vulnerability: complementary or conflicting concepts?’, ecology and society 15(3), article 11. novelli, m. & scarth, a., 2007, ‘tourism in protected areas: integrating conservation and community development in liwonde national park (malawi)’, tourism and hospitality planning and development 4(1), 47–73. phillips, a., 2003, ‘turning ideas on their head: the new paradigm for protected areas’, the george wright forum 20(2), 8–32. plummer, r. & fennell, d.a., 2009, ‘managing protected areas for sustainable tourism: prospects for adaptive co-management’, journal of sustainable tourism 17(2), 149–168. qsr international, 2008, nvivo 8 [computer software], qsr international, victoria. reinius, s.w. & fredman, p., 2007, ‘protected areas as attractions’, annals of tourism research 34(4), 839–854. resilience alliance, 2007a, ‘assessing and managing resilience in social-ecological systems: a practitioner’s workbook, vol. 1, version 1.0’, viewed 31 july 2007, from http://www.resalliance.org/3871.php resilience alliance, 2007b, ‘assessing resilience in social-ecological systems: a scientist’s workbook’, viewed 31 july 2007, from http://www.resalliance.org/3871.php robinson, m., 1999, ‘cultural conflicts in tourism: inevitability and inequality’, in m. robinson & p. boniface (eds.), tourism and cultural conflicts, pp. 1–32, cabi publishing, wallingford. sartorius, k., sartorius, b., tollman, s., schatz, e., kirsten, j. & collinson, m., 2009, the dynamics of rural poverty in a south african community: a spatial-temporal model, institute of behavioural science, university of colorado, boulder. scherl, l.m. & edwards, s., 2007, ‘tourism, indigenous and local communities and protected areas in developing nations’, in r. bushell and p.f.j. eagles (eds.), tourism and protected areas: benefits beyond boundaries, pp. 71–88, cab international, wallingford. shackleton, c., shackleton, s., gambiza1, j., nel, e., rowntree, k. & urquhart, p., 2008, links between ecosystem services and poverty alleviation: situation analysis for arid and semi-arid lands in southern africa, ecosystem services and poverty reduction research programme, dfid, nerc, esrc, south africa. simelane, t.s., kerley, g.i.h. & knight, m.h., 2006, ‘reflections on the relationships between communities and conservation areas of south africa: the case of five south african national parks’, koedoe 49(2), 85–102. simpson, m.c., 2008, ‘the impacts of tourism initiatives on rural livelihoods and poverty reduction in south africa: mathenjwa and mqobela’, in a. spenceley (ed.), responsible tourism: critical issues for conservation and development, pp. 239–266, earthscan, london. simpson, m.c., 2009, ‘an integrated approach to assess the impacts of tourism on community development and sustainable livelihoods’, community development journal 44(2), 186–208. south african national parks, 2008, kruger national park management plan, sanparks, pretoria. south african national parks, 2011, annual report 2010/11, sanparks, pretoria. south african national parks, 2013, ‘guest access data’, unpublished internal report, sanparks, pretoria. statistics south africa, 2007, community survey 2007, revised edn., statistics south africa, pretoria. strickland-munro, j.k., allison, h.e. & moore, s.a., 2010, ‘using resilience concepts to investigate the impacts of protected area tourism on communities’, annals of tourism research 37(2), 499–519. strickland-munro, j.k. & moore, s.a., 2013, ‘indigenous involvement and benefits from tourism in protected areas: a study of purnululu national park and warmun community, australia’, journal of sustainable tourism 21(1), 26–41. taylor, j., 2003, aboriginal population profiles for development planning in the northern east kimberley, centre for aboriginal economic policy research, australian national university, canberra. telfer, d.j. & sharpley, r., 2008, tourism and development in the developing world, routledge, london. trau, a. & bushell, r., 2008, ‘tourism and indigenous people’, in s.f. mccool & r.n. moisey (eds.), tourism, recreation and sustainability: linking culture and the environment, 2nd edn., pp. 260–282, cabi, wallingford. tourism research australia, 2009, nature tourism fact sheet, viewed 27 september 2012, from http://www.ret.gov.au/tourism/documents/tra/snapshots and factsheets/nature_factsheet_final.pdf unesco, 2010, purnululu national park, viewed 08 february 2010, from http://whc.unesco.org/en/list/1094 urban-econ, 2008, sanparks: economic impact assessment, sanparks, pretoria. walker, b.h., anderies, j.m., kinzig, a.p. & ryan, p., 2006, ‘exploring resilience on social-ecological systems through comparative studies and theory development: introduction to the special issue’, ecology and society 11(1), article 12. wall, g. & mathieson, a., 2006, tourism: changes, impacts and opportunities, pearson prentice hall, new york. warmun community (turkey creek) inc., 2007, community corporate planning and management guide 2007–2010, warmun community (turkey creek) inc., warmun. worboys, g.l., lockwood, m. & de lacy, t., 2005, protected area management: principles and practice, 2nd edn., oxford university press, south melbourne. footnotes top ↑ 1. kruger’s northern and eastern boundaries form international borders and were not included in the scope of this research. 2. purnululu’s management policy exempts local indigenous community members from paying park entrance fees. article information authors: gavin gouws1 nasreen peer2 renzo perissinotto2 affiliations: 1south african institute for aquatic biodiversity, grahamstown, south africa 2department of science and technology, national research foundation research chair for shallow water ecosystems, nelson mandela metropolitan university, south africa correspondence to: gavin gouws email: g.gouws@saiab.ac.za postal address: private bag 1015, grahamstown 6140, south africa dates: received: 29 may 2015 accepted: 27 aug. 2015 published: 17 nov. 2015 how to cite this article: gouws, g., peer, n. & perissinotto, r., 2015, ‘mtdna lineage diversity of a potamonautid freshwater crab in kwazulu-natal, south africa’, koedoe 57(1), art. #1324, 12 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1324 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. mtdna lineage diversity of a potamonautid freshwater crab in kwazulu-natal, south africa in this original research... open access • abstract • introduction • methods • results • discussion • conclusion • acknowledgments    • competing interests    • authors’ contributions • references • appendix 1 abstract top ↑ five species of freshwater crab (genus potamonautes) are known from kwazulu-natal, south africa, whilst a sixth (potamonautes isimangaliso) was recently described from the isimangaliso wetland park. earlier molecular studies of crab diversity in the province were largely limited in geographic scope or employed genetic markers, ill-suited for identifying intraspecific diversity. possible species-level diversity or cryptic taxa may have thus remained undetected. in this study, lineage diversity was examined in a widespread species, potamonautes sidneyi, using mitochondrial sequence data, to determine whether this species harbours cryptic diversity that could be of conservation importance in the province, particularly with respect to the isimangaliso wetland park. the taxonomic status of p. isimangaliso was also assessed. mitochondrial sequence data were generated and analysed to identify unique lineages and to examine their distributions. phylogenetic analyses were used to determine whether these lineages represented known or potentially novel species, using comparative data from southern african potamonautes species. seven independent networks were identified within p. sidneyi and substantial structure was observed amongst sampling localities. phylogenetic analyses revealed two distinct, divergent lineages in p. sidneyi. one was positively assigned to p. sidneyi, whereas the placement of the other suggested a novel species. these results suggested possible species diversity within p. sidneyi, with one lineage occurring in the north-east of the province, around the isimangaliso wetland park. potamonautes isimangaliso was clearly allied to potamonautes lividus, but genetic divergences suggested that p. isimangaliso is a distinct taxon and that p. lividus may represent a species complex. conservation implications: this study confirmed unique freshwater crab diversity, both within kwazulu-natal and associated with the isimangaliso wetland park. introduction top ↑ an intensive, dedicated, large-scale systematic study into the diversity of south africa's freshwater crab fauna was launched during the 1990s (hart, stewart & bickerton 2001; stewart 1997a, 1997b; stewart, coke & cook 1995). through this, seven new species were described (daniels, stewart & burmeister 2001; daniels, stewart & gibbons 1998a; gouws, stewart & coke 2000; gouws, stewart & reavell 2001; stewart & cook 1998; stewart 1997a; stewart et al. 1995) and one re-described (stewart 1997b). at its conclusion, 13 species of potamonautes macleay, 1838, the only genus of freshwater crabs occurring in south africa, were recognised (cumberlidge et al. 2009; daniels, phiri & bayliss 2014; gouws & stewart 2001; hart et al. 2001). more than 50% of these are endemic (cumberlidge et al. 2009). subsequently, ongoing research has identified and described four new species (e.g. daniels et al. 2014; peer et al. 2015; phiri & daniels 2014a) throughout the country. along with the western cape province of south africa, the province of kwazulu-natal (kzn) was a substantial focus of this research programme. previously, only two species were known to occur in kzn: potamonautes sidneyi (rathbun, 1904) and potamonautes depressus (krauss, 1843) (gouws & stewart 2001). potamonautes sidneyi is a widely distributed species, occurring across the eastern part of south africa and extending northwards, at least into zimbabwe (gouws, daniels & stewart 2002). it is generally widespread across kzn, from the foothills of the drakensberg mountains through the lower-lying midlands to the coast (gouws & stewart 2001). potamonautes depressus occurs in the upper, faster-flowing tributaries that drain the drakensberg and its foothills (gouws & stewart 2001). a number of accounts of unexpected species or enigmatic records are known from the province, which likely reflect taxonomic uncertainty and/or the assignment of known (extralimital) species epithets to the then-undescribed species (see gouws & stewart 2001; gouws et al. 2002). with dedicated sampling and systematic investigation using morphometric and genetic approaches, three new species were described from kzn. potamonautes dentatus stewart, coke & cook, 1995 was described from sections of the mngeni and thukela rivers by stewart et al. (1995). gouws et al. (2000) described potamonautes clarus gouws, stewart & coke, 2000 from the headwaters of the thukela river along the drakensberg in the north-western part of the province. this species was delineated from p. depressus, which occurs along the drakensberg to the south (gouws & stewart 2001). daniels et al. (2003) later revealed p. clarus and p. depressus to be a complex of considerable cryptic diversity. potamonautes lividus gouws, stewart & reavell, 2001, a morphologically distinct species associated with swamp forests dominated by barringtonia racemosa and ficus tricopoda, was described from a few localities in the north-eastern region of kzn (gouws et al. 2001). more recently, potamonautes isimangaliso peer & gouws, 2015 was described from the isimangaliso wetland park in the north-eastern part of the province. this ecologically unique, burrowing species, occurring in ephemeral pans, was primarily delineated from the burrowing and morphologically similar p. lividus (peer et al. 2015). although the programme in kzn was underpinned by a substantial sampling effort, there were still notable sampling gaps (particularly in terms of specimens examined genetically), especially in the south and north-east of the province (gouws & stewart 2001). the isimangaliso wetland park, an area of considerable biological, cultural and socio-economic importance, is found in the latter. this park is south africa's first united nations educational, scientific and cultural organization world heritage site and includes three ramsar wetlands of international importance, namely the st lucia estuarine lake, lake sibaya and kosi bay (perissinotto, stretch & taylor 2013). although genetic approaches, specifically allozyme electrophoresis, were used to identify and delineate two of the species from the province (gouws et al. 2000; gouws et al. 2001), these studies were limited in their geographic or taxonomic focus (i.e. in terms of specific species compared to the putatively new species). a wider geographic consideration of genetic diversity and structure amongst and within the species of kzn was attempted, again using allozymes (gouws & stewart 2001). however, it has been consistently recognised that allozyme variation in southern african potamonautid crabs is limited and that these markers may be too conservative to detect finer genetic diversity (daniels 2003; daniels, gibbons & stewart 1999; daniels, gouws & crandall 2006; daniels, stewart & gibbons 1998b; daniels, stewart, ridgway & florence 2001; gouws & stewart 2001; gouws et al. 2001; gouws et al. 2002). aside from studies of cryptic diversity and species delineation within the p. clarus – p. depressus complex (daniels et al. 2003; phiri & daniels 2014b), dna sequence data have generally not been used in studies of kzn crabs and there has been no published dna-based survey of lineage diversity within the most widespread species in the province. the extent of divergence amongst mitochondrial dna sequences generated for p. isimangaliso and published data for p. lividus (daniels et al. 2002) provided initial evidence of the separation of these species (peer et al. 2015). however, the phylogenetic relationships between these species, and between p. isimangaliso and other kzn species, were not considered. the primary aim of this study is to provide a coarse-grained and broad-scale snapshot of mitochondrial dna lineage diversity along the coastal zone in the kzn province for the most widespread species, p. sidneyi, and to consider aspects of its biogeography in the region. highlighting unique diversity within the province would be of interest to provincial conservation (ezemvelo kzn wildlife) and regional management (e.g. the isimangaliso wetlands park authority) agencies. documenting undescribed diversity (novel species or unique genetic lineages) in the area around the isimangaliso wetland park would be noteworthy and add to the conservation value of the park, considering the recent description of p. isimangaliso. this species is presently believed to be endemic to the park (peer et al. 2015) and there is a growing number of cryptic taxa and unique lineages recently documented in this region (see maake et al. 2013). the study also seeks to examine the relationship between p. isimangaliso, the morphologically similar p. lividus, and other species from the province and southern african region to support or refute its specific status. methods top ↑ potamonautes sidneyi and p. isimangaliso were sampled between february 2012 and june 2013 from nine localities across kzn, south africa (figure 1), and were identified using published keys (gouws & stewart 2001; hart et al. 2001; peer et al. 2015). four of these localities – lake sibaya (site 1), mpophomeni stream (site 2), dukandlovu pan (site 3) and hluhluwe (site 4) (figure 1) – were from the north-eastern part of kzn, within or in close proximity to the isimangaliso wetland park. the remaining localities were distributed along a coastal transect extending to the south of the province. figure 1: sampling localities of potamonautes sidneyi (open circles) and potamonautes isimangaliso (filled circle) in kwazulu-natal, south africa: (site 1) lake sibaya, (site 2) mpophomeni stream, (site 3) hluhluwe, (site 4) dukandlovu pan, (site 5) entumeni, (site 6) siyayi, (site 7) mhlanga, (site 8) oribi gorge and (site 9) mtamvuna. total genomic dna was extracted from pereiopod muscle tissue from each specimen using a thermo fisher scientific genejet genomic dna purification kit (massachusetts, usa) following the manufacturer's protocols, except that extracted dna was eluted in a final volume of 100 μl. a fragment of the mitochondrial cytochrome c oxidase subunit i (coi) gene was amplified by a polymerase chain reaction. each 25 μl reaction contained 1 × buffer, 3 mm mgcl2, 0.2 μm of each of folmer et al.'s (1994) primers (lcoi-1490 and hcoi-2198), 0.2 mm of each deoxynucleotide, 0.5 u taq-polymerase (southern cross biotechnology, south africa) and 3 μl template dna, made up to the final volume with ultrapure water. the thermocycling regime for the amplification of this gene fragment included an initial denaturing step of 2 min at 95 °c, followed by 35 cycles of denaturing (95 °c for 30 s), annealing (44 °c for 40 s) and extension (72 °c for 1 min). this was followed by a final extension step of 72 °c for 10 min. a fragment of the 16s ribosomal dna (rdna) was amplified from representative individuals (see below), using the primers (16sar and 16sbr) of palumbi et al. (2007) and the above polymerase chain reaction recipe and thermocycling regime, but with annealing performed at 50 °c. to confirm successful amplification, products were visualised on an ultraviolet transilluminator, following electrophoresis in 1% agarose gels stained with ethidium bromide. amplicons were sent to a commercial sequencing facility (macrogen inc., south korea), where they were purified, sequenced using standard bigdye v3.1 (applied biosystems, austin, texas, usa) terminator chemistry and analysed on an abi 3730xl (applied biosystems) automated sequencer. sequences were checked against their respective chromatograms for ambiguities, misreads and sequencing errors using chromas lite (technelysium pty ltd., south brisbane, australia). alignment and further editing were performed using dnastar® lasergene seqman pro 11 (madison, wisconsin, usa). the final alignments of the coi and 16s data sets were produced using clustalx2 (larkin et al. 2007) and maftt version 6 (katoh & toh 2008), respectively, with an iterative refinement strategy (l-ins-i) (katoh et al. 2005) for the latter. data analyses proceeded in two stages. firstly, to understand diversity within and amongst sampling localities of p. sidneyi and p. isimangaliso, and the diversity of lineages within p. sidneyi, analyses were conducted with the full set of coi sequences generated. to examine genealogical and geographic relationships, a 95%-probability parsimony network was generated using tcs1.21 software (clement, posada & crandall 2000). a midpoint-rooted neighbour-joining tree (saitou & nei 1987), based on uncorrected sequence divergences, was generated in paup*4b10 (phylogenetic analysis using parsimony) software (swofford 2002). uncorrected sequence divergences within and amongst identified lineages were examined using paup. in a second set of analyses, the phylogenetic placement of, and divergences amongst, the above lineages were contextualised and an attempt was made to confirm or assign species identifications to each. representatives of the unique lineages identified were included in a matrix containing published data, sourced from genbank® (table 1), of other southern and eastern african potamonautes species. data from the coi and 16s fragments were analysed as a combined data set and phylogenetic analyses were conducted by unweighted parsimony and bayesian inference. for the parsimony analysis, a heuristic tree search was conducted in paup*4b10, using tree bisection and reconnection branch swapping of a tree obtained by a random stepwise addition of taxa, employing 1000 such iterations. nodal support was evaluated by bootstrapping of the data set (felsenstein 1985), using 1000 pseudoreplicates. table 1: southern and eastern african potamonautes species included in the phylogenetic analysis. liberonautes latidactylus and sudanonautes aubryi were included as outgroup taxa. genbank® accession numbers for the cytochrome c oxidase subunit i and 16s ribosomal dna sequences are included and the data sources provided. prior to the bayesian analysis, the optimal model of nucleotide substitution for each partition (coi and 16s rdna) of the combined data set was identified using jmodeltest 2.1.4 (darriba et al. 2012) with the choice of model determined by the akaike (1974) information criterion. bayesian inference was conducted in mrbayes 3.1.2 (ronquist & huelsenbeck 2003). four simultaneous, independent analyses, each employing three heated and one cold markov chains, were run for 107 generations, sampling every 2000 generations. the chosen models were specified for each partition, with mr bayes estimating the model parameters. default, unlinked priors were used. the standard deviation of split frequencies was observed to monitor convergence of the analyses on a similar part of the posterior distribution. stationarity and effective sample size (ess > 200; such that sampling of the various parameters from the posterior distribution was sufficient) were determined using tracer 1.5 software (rambaut & drummond 2009). a majority rule consensus tree was constructed from the 18 000 trees retained from the four analyses combined, once a 10% burn-in was discarded in each; the frequency of retrieval of particular nodes representing the bayesian posterior probabilities (bpps) for those relationships. liberonautes rubigimanus cumberlidge & sachs, 1989 and sudanonautes floweri (de man, 1901) were included as outgroups in these analyses. previous multi-locus phylogenies (daniels, cumberlidge, pérez-losada, marijnissen & crandall 2006; daniels et al. 2015) revealed liberonautes bott, 1955, and a sudanonautes bott, 1955 and potamonemus clade to be sister taxa to the potamonautes a. milne-edwards, 1887 clade that nests both erimetopus rathbun, 1894 and platythelphusa a. milne-edwards, 1887. representatives of liberonautes and sudanonautes have been used as outgroups in other phylogenetic studies of potamonautes (daniels & bayliss 2012; daniels et al. 2014; phiri & daniels 2013). uncorrected sequence divergences amongst known species and the representatives of the respective lineages were calculated in paup. results top ↑ specimens of p. isimangaliso were sampled only from dukandlovu pan (figure 1, site 4), whilst specimens from all the other localities were positively identified as p. sidneyi. all coi sequences were submitted to genbank® under accession numbers kt275869 to kt275936. the full coi alignment of all 69 specimens was 639 nucleotides in length. upon translation to amino acids, there was no evidence of stop codons in the alignment and it was concluded that the data represented functional proteins and were of mitochondrial origin. one haplotype was found amongst the four p. isimangaliso individuals from dukandlovu pan, including the individual sequenced by peer et al. (2015). nineteen unique haplotypes were identified within p. sidneyi (figure 2). the distribution of haplotypes within this species indicated substantial genetic structuring amongst sampling localities. eight individuals possessed unique haplotypes, while nine haplotypes were found in multiple individuals from the same locality. the remaining two haplotypes were shared amongst sampling localities and only amongst two localities in each instance: amongst the proximate lake sibaya (site 1) and mpophomeni stream (site 2) localities, and amongst the geographically more distant hluhluwe (site 3) and entumeni (site 5) localities, respectively. figure 2: relationships amongst cytochrome c oxidase subunit i mtdna haplotypes found amongst potamonautes sidneyi and potamonautes isimangaliso individuals sampled from kwazulu-natal. the 20 haplotypes observed in this study were placed in 8 independent networks that could not be linked with 95% confidence. five networks (figure 2, networks c, e, f, g and h), all but one consisting of single haplotypes, represented single localities. two larger networks (networks a and d) and one consisting of a single haplotype (network b), found in both lake sibaya (site 1) and mpophomeni stream (site 2), represented multiple localities. haplotypes of network a were found in four localities from the north-eastern to central parts of the province: mpophomeni stream site (site 2), hluhluwe (site 3), entumeni (site 5) and siyayi (site 6). those of network d were found in the localities to the south of kzn: mhlanga (site 7) and oribi gorge (site 8). several localities possessed haplotypes of more than one network: entumeni (site 5), mpophomeni stream (site 2), mhlanga (site 7) and mtamvuna (site 9). uncorrected sequence divergences among haplotypes belonging to different networks ranged from 2.8% to 14.7% (with a mean of 10.7%). the two haplotypes of network e were 0.2% divergent. network a showed the greatest diversity (seven haplotypes), with divergences amongst the constituent haplotypes ranging from 0.2% to 1.3% (with a mean of 0.6%). greater divergences were observed within network d (from 0.6% to 2.5%; mean 1.5%). the diversity and the geographic structure observed were substantiated by the neighbour-joining tree (figure 2). two distinct clusters were retrieved, with a single sample from entumeni (npp47) placed outside of these. the first cluster, corresponding to networks a and b, contained samples identified as p. sidneyi from the north-eastern and central parts of the province: lake sibaya (site 1), mpophomeni stream (site 2), hluhluwe (site 3), entumeni (site 5) and siyaya (site 6). the second cluster contained individuals identified as p. sidneyi, as well as the p. isimangaliso specimens from dukandlovu pan (site 4). these formed separate sub-clusters and corresponded to separate networks (networks d to g for p. sidneyi and network h for p. isimangaliso). the p. sidneyi individuals in this cluster were from the three sites towards the south of the province: mhlanga (site 7), oribi gorge (site 8) and mtamvuna (site 9). for the phylogenetic analyses, a 16s rdna fragment was sequenced for a single representative of each of seven networks (networks a–c, and e–h) and three representatives of network d (representing the three divergent groups of haplotypes in this network). these 16s rdna sequences were lodged under genbank® accession numbers kt275860 to kt275868. this fragment was analysed in combination with the coi data for each of these specimens and data downloaded from genbank®. once sequences were trimmed to equal length, the coi and 16s alignments were 362 and 401 nucleotides in length, respectively, yielding a combined alignment of 763 nucleotides. the parsimony analysis, based on 207 parsimony informative characters (97 from the coi data and 110 from the 16s rdna), yielded a single most parsimonious tree (figure 3a: 1106 steps; consistency index = 0.326; retention index = 0.543; rescaled consistency index = 0.177). figure 3: the phylogenetic placement, based on the parsimony and bayesian analyses of combined cytochrome c oxidase subunit i and 16s ribosomal dna sequence data partitions, of unique kwazulu-natal potamonautes sidneyi lineages and potamonautes isimangaliso, relative to southern and eastern african potamonautes species: (a) the single most parsimonious tree (1106 steps, consistency index = 0.326, retention index = 0.543, rescaled consistency index = 0.177) and (b) the most likely (-lnl = 7841.114) topology obtained across the four bayesian analyses. numbers on the branches indicate bootstrap support (only values > 75% are shown) and bayesian posterior probabilities (only bayesian posterior probabilities ≥ 0.95) for relationships in a and b, respectively. hasegawa, kishino and yano’s (1985) hky model, with unequal base frequencies (a = 0.332, c = 0.206, g = 0.077 and t = 0.385), a transition to transversion rate of 4.826, a proportion of invariable sites (i = 0.487) and a gamma-distribution (α = 0.707) of rate variation was determined to be the most appropriate for the coi partition. the model chosen for the 16s partition included unequal base frequencies (a = 0.394, c = 0.078, g = 0.136 and t = 0.393), mostly independent transition and transversion rates (r[a↔c] = 0.364, r[a↔g] = 7.317, r[a↔t] = r[g↔t] = 1.000, r[c↔g] = 0.675 and r[c↔t] = 2.893), a proportion of invariable sites (i = 0.169) and a gamma-distribution (α = 0.443) of rate variation. the most likely topology (-lnl = 7841.114) obtained through the markov chain monte carlo searches across the four bayesian analyses is presented in figure 3b. both phylogenetic trees showed certain similarities to published phylogenies (daniels & bayliss 2012; daniels, cumberlidge, pérez-losada, marijnissen & crandall 2006; daniels et al. 2014; phiri & daniels 2013) relating to the species represented in the major clades retrieved, but many of the relationships within and among these clades were not consistent. nonetheless, it was possible to resolve, in either of the two analyses, the placement of representatives of the unique lineages with high support. like the neighbour-joining tree of the coi data, the representatives of p. sidneyi were placed in two distinct and well-separated clades in both analyses. the representatives of the lineages from the north-eastern and central parts of the province (npp65 hluhluwe, representing network a; and npp70 lake sibaya, representing network b) formed a clade with the published p. sidneyi sequence with high support (95% bootstrap; 1.00 bpp). npp47 entumeni (network c) was well-supported as a sister-taxon to this group (0.99 bpp) in the bayesian analysis. although this specimen was not associated with this clade in the parsimony analysis, the alternative placement was not supported (figure 3a). regardless, these specimens were all placed in a larger clade containing a number of large-bodied riverine species (sensu daniels et al. 2002; potamonautes barbarai phiri & daniels, 2014, potamonautes barnardi phiri & daniels, 2014, potamonautes bayonianus [brito-capello, 1864], potamonautes granularis daniels, stewart & gibbons, 1998, potamonautes perlatus [h. milne-edwards, 1837], p. sidneyi, potamonautes unispinus stewart & cook, 1998 and potamonautes warreni [calman, 1918]) with strong support (98%; 1.00 bpp). the second group of individuals identified as p. sidneyi (npp13 oribi gorge, npp15 oribi gorge and npp21 mhlanga, representing network d, and the representatives of networks e, f and g) from the south of the province were placed in a clade containing two published p. lividus sequences and the p. isimangaliso (network h) representative, with 1.00 bpp. the p. isimangaliso specimen was consistently the sister-taxon to the sequence of p. lividus from south africa, to the exclusion of the representative from swaziland. sequence divergences amongst all included, recognised species and representatives of the lineages are presented in the appendix, table 1. divergences amongst known species ranged from 2.3% (p. barbarai versus p. granularis) to 21.5% (potamonautes calcaratus [gordon, 1929] versus p. lividus from south africa), with a mean of 13.5%. those specimens of p. sidneyi from the north-eastern and central parts of the province, placed in a clade with p. sidneyi and the large-bodied riverine species, were 1.8% – 4.8% divergent from the published p. sidneyi sequence. these individuals (and p. sidneyi itself) were, in turn, 9.2% – 11.8% divergent from the representatives of the second clade of p. sidneyi individuals from the south of the province. the divergences amongst these two groups were substantially higher than the divergences within (0.4% – 4.8%). this southern clade showed a marginally closer relationship to p. lividus and p. isimangaliso (6.7% – 10.0% divergence) than to the north-eastern p. sidneyi clade. considering the combined coi and 16s rdna data, the p. isimangaliso representative was 7.4% and 7.8% divergent from the swaziland and south african representatives of p. lividus, respectively. discussion top ↑ the analysis of the coi data reveals substantial genetic diversity and structure within p. sidneyi in kzn; 7 divergent, unlinked networks were retrieved (with an additional network representing p. isimangaliso) and only 2 of 19 haplotypes are shared amongst sampling localities. for the most part, the data show a clear isolation and lack of gene flow amongst most sampling localities, even amongst those that are geographically proximate. this is contrary to expectations for a species that is assumed to be highly dispersive (daniels 2003; daniels et al. 1998a, 1998b; daniels et al. 1999; daniels, gouws & crandall 2006; daniels, stewart, ridgway & florence 2001) and follows more closely patterns observed in certain primary, obligate freshwater fishes from south africa, for example galaxias (chakona, swartz & gouws 2013). the patterns observed reflect not only the isolation of individual drainages, but often the isolation of localities within the same system (e.g. amongst hluhluwe and mpophomeni streams, which flow into lake st lucia). in the similarly widespread potamonautes perlatus sensu lato, extensive sharing of haplotypes was observed amongst localities, but these were within the same catchments (daniels, gouws & crandall 2006; phiri & daniels 2014a). very few haplotypes were shared amongst currently isolated systems, much like the present case. the isolation of the individual localities and systems is perhaps facilitated by the hydrology of kzn. in the subtropical to tropical north-east of the province, drainages are generally mature, widely separated and flow over low relief floodplains (skelton 2001). geographic distance may prohibit dispersal amongst systems, despite the area being flood-prone and the presence of ephemeral wetlands (skelton 2001), which should facilitate dispersal. towards the south of the province, in the montane escarpment region, a dense network of shorter, high-gradient, deeply-incised catchments fragment the landscape (skelton 2001). dispersal may be hindered across higher drainage divides. two divergent phylogenetic lineages were found in p. sidneyi, with one of these showing a possible closer affinity to p. lividus and p. isimangaliso. these two lineages were restricted to the north-eastern and central, and southern parts of the province, respectively. the distribution of these lineages and the division amongst them reflects the two aquatic ecoregions mentioned above, as well as the separation of the sibayi and zululand from the mngeni and mzimkhulu aquatic bioregions (see rivers-moore, goodman & nkosi 2007); these regions defined by ecology, species diversity and endemism. previous studies considering the genetic diversity or structure in p. sidneyi in kzn provided conflicting patterns. an allozyme study, delineating p. lividus from p. sidneyi, found low levels of diversity and only shallow structure in the latter (gouws et al. 2001). however, localities from which p. sidneyi was sampled were from the north-eastern region of kzn and may represent only one of the lineages identified in the current study. with slightly wider sampling, a study of morphometric and allozyme variation within and amongst all potamonautes species in kzn provided some evidence of genetic structure and differentiation within p. sidneyi (gouws & stewart 2001). two distinct genetic clusters were identified: one from the central and north-east of the province and the other from the high-lying drakensberg in the extreme west (gouws & stewart 2001). whether these correspond to the two lineages of the present study or represent additional lineage diversity remains to be determined as no p. sidneyi populations from the drakensberg were included in this study. nonetheless, the genetic differentiation of p. sidneyi from the drakensberg was confirmed in a subsequent study (gouws et al. 2002). these populations form a distinct genetic cluster separate to samples from elsewhere in the province; the latter showed closer genetic similarity to p. sidneyi samples from other provinces, that is, mpumalanga, eastern cape and limpopo (gouws et al. 2002). interestingly, the latter study showed genetic differentiation amongst populations of p. sidneyi from different reaches of the same river system, the thukela. this reflects a longitudinal pattern of differentiation involving the phylogenetic separation (see daniels et al. 2002) of high-altitude, small-bodied species (members of the p. depressus – p. clarus complex in this system) in the upper reaches from the robust species of the middle to lower reaches (p. sidneyi) and then possible longitudinal genetic structure within the latter. the co-occurrence of multiple, genetically independent networks at single localities, particularly in the south of the province where drainage is steep and with potential for isolation along the river course, attests to the diversity in the species and may perhaps reflect such longitudinal or altitudinal genetic structure. as the present study focused on sampling in the coastal zone only, this requires further investigation in the thukela river and these southern systems. nonetheless, p. sidneyi clearly shows a complex pattern of lineage diversity and structure within kzn and beyond. this is currently being investigated through a much broader phylogeographic study of this species across its full south african distribution. the taxonomic status of the two p. sidneyi lineages requires consideration. the sequence divergence amongst these lineages (9.2% – 11.8%) was more extensive than was observed amongst other species (p. barbarai, p. barnardi, p. bayonianus, p. granularis, p. perlatus, p. unispinus and p. warreni) in the group of large-bodied, riverine species from south africa, and amongst these species and the reference p. sidneyi individual (2.3% – 8.3%). furthermore, the higher values from individual comparisons amongst the two lineages approached the mean divergence amongst all valid species included in the study. potamonautes sidneyi in kzn shows comparable diversity to p. perlatus (see phiri & daniels 2014a), which, along with p. sidneyi, is one of the most widely distributed south african freshwater crab species (daniels 2003; daniels, gouws & crandall 2006; gouws et al. 2002; phiri & daniels 2014a). however, a deeper divergence was observed between lineages of p. sidneyi in kzn than was observed amongst the lineages of p. perlatus. comparisons amongst representatives of the two p. sidneyi lineages yielded sequence divergences of 6.2% – 9.3% for the 16s data, and 12.2% – 14.3% for coi, which exceeded the maximum values of 6% and 10% for 16s and coi, respectively, in p. perlatus (daniels 2003; daniels, gouws & crandall 2006). two new species (p. barbarai and p. barnardi) were delineated, using genetic evidence alone, amongst the three lineages in p. perlatus sensu lato (phiri & daniels 2014a). the extent of divergence, the phylogenetic placement of lineages identified as p. sidneyi in the present phylogenetic analyses, results from past analyses (gouws & stewart 2001; gouws et al. 2002) and the precedent set by phiri and daniels (2014a) suggests that there may be additional species-level diversity with p. sidneyi, requiring recognition. the identification and description of new species within p. sidneyi is deferred at present, for two reasons. the first is the uncertainty over which lineage or clade represents the ‘true’ p. sidneyi. rathbun (1905), when describing the species, indicated the type locality as being ‘natal’, without providing a more specific locality within the province. the species were described from material collected by sarah abraham in 1871 (rathbun 1905) and archival research may be required to determine the type locality more accurately. secondly, the wider phylogeographic study and a more rigorous consideration of morphological variation across the species’ range would provide greater insight into the taxonomic status of these and any other lineages. while p. isimangaliso shows a clear resemblance to p. lividus (peer et al. 2015), the lack of the distinctive colouration of the latter (gouws & stewart 2001; gouws et al. 2001) aided the identification of p. isimangaliso specimens in the current study. however, recent collections of p. lividus (daniels et al. 2014) suggest that colour may be variable in this species (s.r. daniels, pers. comm.). phylogenetically, the representative p. isimangaliso specimen was associated with p. lividus from south africa (kzn), with the representative from swaziland sister to these (or even placed outside of a clade formed by these and one of the p. sidneyi lineages). the phylogenetic placement of p. isimangaliso suggests that it may be conspecific to p. lividus from kzn. however, the p. isimangaliso specimen was substantially (~ 7.5%) and roughly equally divergent from each of the p. lividus representatives. recently, daniels et al. (2014) reported the occurrence of p. lividus from dweza forest in the eastern cape province. together with the specimens from swaziland (see daniels & bayliss 2012), this represents a significant range extension for the species. however, unpublished sequence data revealed almost no differentiation among p. lividus from kzn and from the eastern cape (daniels et al. 2014). this suggests a genetic cohesion within p. lividus over a range of some ~ 750 km (daniels et al. 2014) and indicates that both p. lividus from swaziland and p. isimangaliso represent unique lineages, both warranting recognition at species level. the description by peer et al. (2015) and support for p. isimangaliso as a valid taxon are significant because gouws et al. (2001) highlighted a number of specimens in museum collections that resembled p. lividus. however, identifications of these specimens could not be confirmed at the time as the seemingly diagnostic colouration (see above) was not preserved. repeated sampling from these localities, which were not typical swamp forests, was unsuccessful. the possibility that these morphotypes may correspond to p. isimangaliso, p. lividus or additional taxa within a possible complex require further investigation. conclusion top ↑ the present study highlights the unique diversity within the north-eastern region of kzn, in and around the isimangaliso wetland park. a unique lineage of p. sidneyi is found here, which is separate to a lineage occurring to the south of the province and possibly an additional lineage occurring along the drakensberg to the west (gouws & stewart 2001). whether this north-eastern lineage is restricted to this area or has a more extensive distribution throughout the range of p. sidneyi remains to be determined. allozyme invariance among populations from this area and from further afield, including other provinces, indicates that this may be the case (gouws et al. 2002). however, the conservative nature of these markers in potamonautid crabs (see introduction) may obscure true lineage diversity. potamonautes isimangaliso is currently endemic to the isimangaliso wetland park (peer et al. 2015), but possibly occurs at other localities in the north-east (gouws et al. 2001), and clearly represents a unique species within the p. lividus – p. isimangaliso complex. these findings suggest a somewhat unique potamonautid fauna within this region, adding to the conservation value of this ecologically-, culturallyand socio-economically important system. the isimangaliso wetland park was proclaimed a united nations educational, scientific and cultural organization world heritage site in 1999 for three outstanding universal values, one of which is biodiversity (perissinotto et al. 2013). the findings of this study confirm that the park truly represents a hotspot of biological diversity. acknowledgments top ↑ this research was financially supported by the national research foundation (nrf) of south africa through the south african research chair initiative (sarchi) (grant no. 84375) of the department of science and technology (dst) and the nrf's support of the first author. we thank the isimangaliso wetland park authority and ezemvelo kzn wildlife (ekznw) for supporting this research. sampling for this research was conducted under permits issued by ekznw (permits op 685/2015). the department of environmental affairs and the department of agriculture, forestry and fisheries granted an integrated scientific investigation permit that allowed the completion of this research. we thank willem coetzer (saiab) for the drafting of the map. all opinions, findings, conclusions or recommendations expressed in this material are those of the authors and the nrf does not accept any liability in this regard. we thank several anonymous referees and editors for input on past versions of this manuscript. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions the study was jointly conceived, conceptualised and designed by n.p. (nelson mandela metropolitan university), r.p. (nelson mandela metropolitan university) and g.g. (south african institute for aquatic biodiversity). fieldwork and sampling were undertaken by n.p. and r.p. data were generated by n.p. and g.g., and analysed by g.g. g.g. took the lead in writing the manuscript, with significant editorial input and direction from r.p. and n.p. the research was supported, financially and logistically, by grants made to r.p. references top ↑ akaike, h., 1974, ‘a new look at the statistical model identification’, institute of electrical and electronics engineers transactions on automatic control 19, 716–723. http://dx.doi.org/10.1109/tac.1974.1100705 chakona, a., swartz, e.r. & gouws, g., 2013, ‘evolutionary drivers of diversification and distribution of a southern temperate stream fish assemblage: testing the role of historical isolation and spatial range expansion’, plos one 8, e70953. pmid: 23951050, http://dx.doi.org/10.1371/journal.pone.0070953 clement, m., posada, d. & crandall, k.a., 2000, ‘tcs: a computer program to estimate gene genealogies’, molecular ecology 9(10), 1657–1659. pmid: 11050560, http://dx.doi.org/10.1046/j.1365-294x.2000.01020.x cumberlidge, n., ng, p.k.l., yeo, d.c.j., magalhães, c., campos, m.r., alvarez, f. et al., 2009, ‘freshwater crabs and the biodiversity crisis: importance, threats, status, and conservation challenges’, biological conservation 142, 1665–1673. http://dx.doi.org/10.1016/j.biocon.2009.02.038 daniels, s.r., 2003, ‘examining the genetic structure among populations of the common cape river crab potamonautes perlatus from river systems in south africa reveals hydrographic boundaries’, journal of crustacean biology 23, 936–950. http://dx.doi.org/10.1651/c-2383 daniels, s.r. & bayliss, j., 2012, ‘neglected refugia of biodiversity: mountainous regions in mozambique and malawi yield two novel freshwater crab species (potamonautidae: potamonautes)’, zoological journal of the linnean society 164, 498–509. http://dx.doi.org/10.1111/j.1096-3642.2011.00773.x daniels, s.r., cumberlidge, n., pérez-losada, m., marijnissen, s.a.e. & crandall, k.a., 2006, ‘evolution of afrotropical freshwater crab lineages obscured by morphological convergence’, molecular phylogenetics and evolution 40, 227–235. http://dx.doi.org/10.1016/j.ympev.2006.02.022 daniels, s.r., gibbons, m.j. & stewart, b.a., 1999, ‘allozyme variation among populations of the freshwater crab, potamonautes perlatus (decapoda: potamonautidae) in the berg river system, western cape’, south african journal of zoology 34, 64–68. daniels, s.r., gouws, g. & crandall, k.a., 2006, ‘phylogeographic patterning in a freshwater crab species (decapoda: potamonautidae: potamonautes) reveals the signature of historical climatic oscillations’, journal of biogeography 33, 1538–1549. http://dx.doi.org/10.1111/j.1365-2699.2006.01537.x daniels, s.r., gouws, g., stewart, b.a. & coke, m., 2003, ‘molecular and morphometric data demonstrate the presence of cryptic lineages among freshwater crabs (decapoda: potamonautidae: potamonautes) from the drakensberg mountains, south africa’, biological journal of the linnean society 78, 129–147. http://dx.doi.org/10.1046/j.1095-8312.2003.00143.x daniels, s.r., phiri, e.e. & bayliss, j., 2014, ‘renewed sampling of inland aquatic habitats in southern africa yields two novel freshwater crab species (decapoda: potamonautidae: potamonautes)’, zoological journal of the linnean society 171, 356–369. http://dx.doi.org/10.1111/zoj.12139 daniels, s.r., phiri, e.e., klaus, s., albrecht, c. & cumberlidge, n., 2015, ‘multilocus phylogeny of the afrotropical freshwater crab fauna reveals historical drainage connectivity and transoceanic dispersal since the eocene’, systematic biology 64, 549–567. pmid: 25649930, http://dx.doi.org/10.1093/sysbio/syv011 daniels, s.r., stewart, b.a. & burmeister, l., 2001, ‘geographic patterns of genetic and morphological divergence amongst populations of a river crab (decapoda, potamonautidae) with the description of a new species from mountain streams in the western cape, south africa’, zoologica scripta 30, 181–197. http://dx.doi.org/10.1046/j.1463-6409.2001.00061.x daniels, s.r., stewart, b.a. & gibbons, m.j., 1998a, ‘potamonautes granularis sp. nov. (brachyura, potamonautidae), a new cryptic species of river crab from the olifants river system, south africa’, crustaceana 71, 885–903. http://dx.doi.org/10.1163/156854098x00905 daniels, s.r., stewart, b.a. & gibbons, m.j., 1998b, ‘genetic and morphometric variation in the potamonautid river crab potamonautes parvispina (decapoda: potamonautidae) from two western cape rivers, south africa’, journal of natural history 32, 1245–1258. http://dx.doi.org/10.1080/00222939800770621 daniels, s.r., stewart, b.a., gouws, g., cunningham, m. & matthee, c.a., 2002, ‘phylogenetic relationships of the southern african freshwater crab fauna (decapoda: potamonautidae: potamonautes) derived from multiple data sets reveal biogeographic patterning’, molecular phylogenetics and evolution 25, 511–523. pmid: 12450755, http://dx.doi.org/10.1016/s1055-7903(02)00281-6 daniels, s.r., stewart, b.a., ridgway, t.m. & florence, w., 2001, ‘carapace dentition patterns, morphometrics and allozyme differentiation amongst two toothed freshwater crab species (potamonautes warreni and p. unispinus) (decapoda: brachyura: potamonautidae) from river systems in south africa’, journal of zoology (london) 255, 389–404. http://dx.doi.org/10.1017/s0952836901001480 darriba, d., taboada, g.l., doallo, r. & posada, d., 2012, ‘jmodeltest 2: more models, new heuristics and parallel computing’, nature methods 9, 772. pmid: 22847109, http://dx.doi.org/10.1038/nmeth.2109 felsenstein, j., 1985, ‘confidence limits on phylogenies: an approach using the bootstrap’, evolution 39, 783–791. http://dx.doi.org/10.2307/2408678 folmer, o., black, m., hoeh, w., lutz, r. & vrijenhoek, r., 1994, ‘dna primers for amplification of mitochondrial cytochrome c oxidase subunit i from diverse metazoan invertebrates’, molecular marine biology and biotechnology 3, 294–299. pmid: 7881515. gouws, g., daniels, s.r. & stewart, b.a., 2002, ‘allozyme electrophoresis demonstrates the presence of a species boundary in freshwater crabs (decapoda: potamonautidae)’, journal of natural history 36, 1199–1222. http://dx.doi.org/10.1080/00222930110045425 gouws, g. & stewart, b.a., 2001, ‘potamonautid river crabs (decapoda, brachyura, potamonautidae) of kwazulu-natal, south africa’, water sa 27, 85–98. http://dx.doi.org/10.4314/wsa.v27i1.5015 gouws, g., stewart, b.a. & coke, m., 2000, ‘evidence for a new species of river crab (decapoda, brachyura, potamonautidae) from the drakensberg, south africa’, journal of crustacean biology 20, 743–758. http://dx.doi.org/10.1651/0278-0372(2000)020[0743:efanso]2.0.co;2 gouws, g., stewart, b.a. & reavell, p.e., 2001, ‘a new species of freshwater crab (decapoda, potamonautidae) from the swamp forests of kwazulu-natal, south africa: biochemical and morphological evidence’, crustaceana 74, 137–160. http://dx.doi.org/10.1163/156854001750096256 hart, r.c., stewart, b.a. & bickerton, i.b., 2001, ‘decapoda (chapter 6)’, in j.a. day, b.a. stewart, i.j. de moor & a.e. louw (eds.), guides to the freshwater invertebrates of southern africa. volume 4: crustacea iii. bathynellacea, amphipoda, isopoda, spelaeogriphacea, tanaidacea and decapoda, pp. 87–123, water research commission, pretoria. hasegawa, m., kishino, h. & yano, t., 1985, ‘dating of the human-ape splitting by a molecular clock of mitochondrial dna’, journal of molecular evolution 22, 160–174. pmid: 3934395. http://dx.doi.org/10.1007/bf02101694 katoh, k., kuma, k., toh, h. & miyata, t., 2005, ‘mafft version 5: improvement in accuracy of multiple sequence alignment’, nucleic acids research 33, 511–518. pmid: 15661851, http://dx.doi.org/10.1093/nar/gki198 katoh, k. & toh, h., 2008, ‘recent developments in the mafft multiple sequence alignment program’, briefings in bioinformatics 9, 286–298. pmid: 18372315, http://dx.doi.org/10.1093/bib/bbn013 larkin, m.a., blackshields, g., brown, n.p., chenna, r., mcgettigan, p.a., mcwilliam, h. et al., 2007, ‘clustal w and clustal x version 2.0’, bioinformatics 23, 2947–2948. pmid: 17846036, http://dx.doi.org/10.1093/bioinformatics/btm404 maake, p.a., mwale, m., dippenaar, s.m. & gon, o., 2013, ‘mitochondrial and nuclear dna reveals a complete lineage sorting of glossogobius callidus (teleostei: gobiidae) in southern africa’, african journal of aquatic science 38, 15–29. http://dx.doi.org/10.2989/16085914.2013.782263 palumbi, s., martin, a., romano, s., mcmillan, w.o., stice, l. & grabowski, g., 2007, the simple fool’s guide to pcr version 2, computer software, department of zoology and kewalo marine laboratory, university of hawai’i, honolulu, hi. peer, n., perissinotto, r., gouws, g. & miranda, n.a.f., 2015, ‘description of a new species of potamonautes macleay, 1838, from the isimangaliso wetland park, south africa’, zookeys 503, 23–43. pmid: 26019669, http://dx.doi.org/10.3897/zookeys.503.9532 perissinotto, r., stretch, d.d. & taylor, r.h. (eds.), 2013, ecology and conservation of estuarine ecosystems: lake st. lucia as a global model, cambridge university press, cambridge. phiri, e.e. & daniels, s.r., 2013, ‘hidden in the highlands: the description and phylogenetic position of a novel endemic freshwater crab species (potamonautidae: potamonautes) from zimbabwe’, invertebrate systematics 27(5), 530–539. http://dx.doi.org/10.1071/is13012 phiri, e.e. & daniels, s.r., 2014a, ‘disentangling the divergence and cladogenesis in the freshwater crab species (potamonautidae: potamonautes perlatus sensu lato) in the cape fold mountains, south africa, with the description of two novel cryptic lineages’, zoological journal of the linnean society 170(2), 310–332. http://dx.doi.org/10.1111/zoj.12103 phiri, e.e. & daniels, s.r., 2014b, ‘a coalescent multilocus species delimitation approach demonstrates widespread cryptic differentiation within two mountain-living freshwater crab lineages’, unpublished data. plachetzki, d.c. & cumberlidge, n., 2001, ‘a preliminary phylogenetic analysis of african freshwater crab (decapoda: potamoidea: potamonautidae) based on mitochondrial dna coi sequences’, unpublished data. rambaut, a. & drummond, a.j., 2009, tracer software version 1.5. http://beast.bio.ed.ac.uk. rathbun, m.j., 1905, ‘les crabes d’eau douce (potamonidae)’, nouvelles archives du muséum d’histoire naturelle (paris) 7, 159–322. rivers-moore, n.a., goodman, p.s. & nkosi, m.r., 2007, ‘an assessment of the freshwater natural capital in kwazulu-natal for conservation planning’, water sa 33, 665–673. ronquist, f. & huelsenbeck, j.p., 2003, ‘mrbayes 3: bayesian phylogenetic inference under mixed models’, bioinformatics 19, 1572–1574. pmid: 12912839, http://dx.doi.org/10.1093/bioinformatics/btg180 saitou, n. & nei, m., 1987, ‘the neighbor-joining method: a new method for reconstructing phylogenetic trees’, molecular biology and evolution 4, 406–425. pmid: 3447015. skelton, p.h., 2001, a complete guide to the freshwater fishes of southern africa, struik publishers, cape town. stewart, b.a., 1997a, ‘biochemical and morphological evidence for a new species of river crab potamonautes parvispina sp. nov. (brachyura. potamonautidae)’, crustaceana 70(6), 737–753. http://dx.doi.org/10.1163/156854097x00168 stewart, b.a., 1997b, ‘morphological and genetic differentiation between populations of river crabs (decapoda: potamonautidae) from the western cape, south africa, with a taxonomic re-examination of gecarcinautes brincki’, zoological journal of the linnean society 119(1), 1–21. http://dx.doi.org/10.1111/j.1096-3642.1997.tb00133.x stewart, b.a., coke, m. & cook, p.a., 1995, ‘potamonautes dentatus, new species, a fresh-water crab (brachyura: potamoidea: potamonautidae) from kwazulu-natal, south africa’, journal of crustacean biology 15(3), 558–568. http://dx.doi.org/10.2307/1548776 stewart, b.a. & cook, p.a., 1998, ‘identification of a new species of river crab (decapoda: brachyura: potamonautidae) from south africa using morphological and genetic data’, journal of crustacean biology 18, 556–571. http://dx.doi.org/10.2307/1549420 swofford, d.l., 2002, paup*, phylogenetic analysis using parsimony (*and other methods) version 4, computer software, sinauer associates, sunderland, ma. appendix 1 top ↑ table 1-a1: sequence divergences among southern african potamonautes species and unique lineages found in kwazulu-natal. divergences are uncorrected sequence divergences, calculated from the combined 16s ribosomal dna and cytochrome c oxidase subunit i mitochondrial dna sequence data. article information authors: colin s. schoeman1 stefan h. foord2 affiliations: 1department of zoology, university of venda, south africa2centre for invasion biology, department of zoology, university of venda, south africa correspondence to: colin schoeman postal address: private bag x5050, thohoyandou 5050, south afica dates: received: 14 oct. 2010 accepted: 16 sept. 2011 published: 23 jan. 2012 how to cite this article: schoeman, c.s. & foord, s.h., 2012, ‘a checklist of epigaeic ants (hymenoptera: formicidae) from the marakele national park, limpopo, south africa’, koedoe 54(1), art. #1030, 7 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1030 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. a checklist of epigaeic ants (hymenoptera: formicidae) from the marakele national park, limpopo, south africa in this checklist... open access • abstract • introduction • methods • results • discussion • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ ant surveys are extensively used to guide conservation decisions and form part of a ‘shopping basket’ of invertebrate taxa proposed for the use in monitoring programmes in south africa’s national parks. however, very few ant inventories exist for these conservation areas. we report on the first quantitative survey of ants in the marakele national park (67 000 ha). ants were sampled in four habitats, covering both the altitudinal range (1000 m a.s.l. – 2000 m a.s.l.) and three vegetation types in the park. a total of 4847 specimens, representing 29 genera and 104 species, were recorded from pitfall traps over a five-day period. myrmicinae was the most abundant and diverse subfamily, representing 82% of all ants sampled, followed by the formicinae subfamily, which represented 18% of the total abundance. the most abundant species were members of the pheidole megacephala group, pheidole sculpturata mayr and members of the monomorium salomonis group. in general, we found that the less complex habitats supported higher ant diversity. the marakele national park contains a quarter of the ant species recorded in south africa and is a potential hotspot for invertebrate conservation.conservation implications: the marakele national park represents an area of high ant – and therefore invertebrate – diversity. ant conservation would require attention to each of the vegetation types to maintain complementarity (beta diversity) of the assemblages as well as consideration to the impact of large herbivores, whose presence positively influence ant richness at a site (alpha diversity). introduction top ↑ the lack of taxonomic invertebrate expertise poses a significant challenge to conservation-related decision making. this statement is particularly relevant to ants (hymenoptera: formicidae): although traditionally considered a ’difficult‘ group (bolton 1984), they are increasingly being used for environmental impact assessments (andersen 1997), monitoring environmental change (kaspari & majer 2000; mcgeoch & sithole 2009) and even as surrogates for insect species richness (uys, hamer & slotow 2010).ant surveys are generally considered to be a very rich source of data for conservation planning and management (kremen et al. 1993; yek et al. 2009). they can also be used to delineate biogeographic zones, areas of high general biodiversity and centres of evolutionary radiation (kremen et al. 1993). ant monitoring systems have been developed in australia to assess restoration success after mining and have since been applied successfully in assessing grazing impacts in rangelands (andersen et al. 2004) and in the succulent karoo (seymour & dean 1999). ant community data have been used predominantly to understand structural diversity in ecosystems, yet may prove to be seminal in studies attempting to answer important questions concerning functional biodiversity (folgarait 1998). ants are used as surrogates of ecosystems per se, because they form a staggeringly diverse and abundant group and constitute a substantial part of animal biomass (hölldobler & wilson 1994). ants have invaded almost all ecosystems across the world; they are important ecosystem engineers (samways 2005) and are significant predators of other terrestrial invertebrates (hölldobler & wilson 1994). there are also different functional groups amongst ants, displaying a wide range of life histories and food preferences (andersen 2000). they perform important ecosystem functions, for example myrmecochory (giliomee 2003; lingyel et al. 2009) and the maintenance of rare lycaenid butterfly populations (pierce et al. 2002). above all, they are important ecosystem engineers. a considerable body of work exists on the ants of africa (arnold 1924; fisher 2004; garcia et al. 2009; koch & vohland 2004; lévieux 1972; marsh 1986; taylor 2005; willis, skinner & robertson 1992). african ant community richness clearly rivals that of other tropical ecosystems worldwide. fisher (2004) collected a total of 310 species across 56 genera on mont doubou in gabon. in recent years, surveys of epigaeic ants in south africa have yielded interesting baseline data that indicate effects of agricultural activity (addison & samways 2000; gaigher 2008), invasions by alien plants and the argentine ant (schoeman & samways 2011), fire regimes (parr, bond & robertson 2002; parr & chown 2003), vegetation types (boonzaaier, mcgeoch & parr 2007) and predicted future climate change (botes et al. 2006; koch & vohland 2004) on ant community assemblages. with such superdiverse assemblages, there are still gaps in our knowledge on the biology, ecology, taxonomy and distribution of ants in africa. mcgeoch et al. (2011) have recently identified five major groups that can serve as surrogates in conservation planning and monitoring, namely araneae, scarabaeidae (coleoptera), lepidoptera, odonata and formicidae (hymenoptera). we report on an invertebrate study in the marakele national park, on the south-western border of the limpopo province. the study aimed to determine ant diversity and generate baseline data that can serve as a reference for future studies in the marakele national park. although there are several type specimens from the waterberg plateau (gardiner & terblanche 2010; szüts & jocqué 2001), this study probably represents only the second over-arching invertebrate survey for this protected area, the first being a study on termites (isoptera) by sileshi et al. (2010). the information from this study will contribute to the knowledge on invertebrates occurring across the network of south african protected areas. methods top ↑ the marakele national park (67 000 ha) is a sanctuary situated in the waterberg region near the town of thabazimbi in the limpopo province, south africa (figure 1). it is a core conservation area in the waterberg biosphere reserve. the park includes the highest point in the limpopo province (2110 m a.s.l.). it receives 500 mm – 750 mm rainfall per year and includes three vegetation types (mucina & rutherford 2006), namely western sandy bushveld, waterberg mountain bushveld and waterberg/magaliesberg summit sourveld (figure 1). the first two vegetation types mentioned are poorly protected in south africa (mucina & rutherford 2006). figure 1: map of the vegetation types in the marakele national park. m1–m4 are the sampling locations (pitfall transects of 400 m) for the sites. ants were sampled as part of the south african national survey of arachnida (sansa). the sansa protocol (dippenaar-schoeman & craemer 2000) includes pitfall trapping as one of its sampling methods and requires that four sites be selected to include vegetation types representative of a survey area as a whole. sites at the marakele national park were located at various altitudes along the western aspect of the mountain, namely at 1074 m (m1), 1182 m (m2), 1587 m (m3) and on the summit at 2044 m (m4), and in each of the vegetation types in the park (figure 2a–d). one site, m3, was ecotonal (figure 1 and figure 2c). a total of 24 pitfall traps (diameter = 6 cm; depth = 10 cm) were placed in different patches at each site in february 2010. the traps were placed 20 m apart, flush with the soil surface, and care was taken not to disturb any existing leaf litter layers or nearby organic material in order to minimise digging-in effects (greenslade 1973). the traps contained 50 ml propylene glycol, which is non-lethal and does not have a scent that can attract or repel ants. the pitfalls were left out for a period of 5 days. pitfall samples from each site were pooled. figure 2: vegetation type at the four sampling sites. (a) m1, western sandy bushveld; (b) m2, waterberg mountain bushveld; (c) m3, ecotone on waterberg–magalies summit sourveld and waterberg mountain bushveld; (d) m4, waterberg–magalies summit sourveld. methods to process ants are described in lattke (2000). sampled ants were washed to remove any rubble, leaves and soil. ants were then placed in 70% alcohol (96% alcohol diluted with glycol) and then sorted according to morphospecies. these were further separated and identified to genus, subgenus, species group or species level according to the classifications by agosti and johnson (2005), bolton (1984), hölldobler and wilson (1994) and taylor (2005). agosti and johnson (2005) and taylor (2005) have a complete list of taxonomic references. a reference collection is housed in the arthropod collection of the department of zoology at the university of venda.species richness estimates and inventory completeness were calculated using chao 1, a non-parametric technique based on the distribution of individuals amongst species (colwell & coddington 1994): where sobs is the observed species, f1 the number of observed species represented by a single individual and f2 the number of observed species represented by more than one individual. with increasing pitfalls added to a sample, the observed species richness usually approaches the estimated species richness, which is depicted graphically. as we did not have replicates for a site, we used the chao1 estimator and calculated inventory completion by comparing the estimated species richness with the observed richness per site. simpson’s diversity index is one of many diversity indices that calculate a measure of diversity for communities based on the number of species represented and also their relative abundances. a diversity index was calculated for each site according to where n is the community size (i.e. σn) and n is each population size. species turnover between sites was calculated from square-root transformed abundance data using the bray–curtis similarity and hierarchically clustered using primer 6 (primer-e, plymouth) (figure 3). figure 3: dendrogram based on group average linking of bray–curtis similarity matrix of ant assemblages at the four sampling sites. results top ↑ a total of 4847 ants were collected. the ants were most abundant at the site of lowest altitude (m1), with 1737 individuals, followed by 1512 individuals collected at m3, 823 individuals at m2, and 775 individuals at m4. a total of 104 species across 29 genera were sampled (tables 1 and 2), of which 18 were singletons that likely represent rare species and are not merely a result of undersampling. observed species richness for all samples approached 98% of the chao 1 species richness estimates. results for separate sites also show that sampling was relatively complete, with observed species richness approaching 89%, 98%, 96% and 97% of the estimated species richness for m1, m2, m3 and m4, respectively. the use of pitfall sampling allowed us to collect strictly epigaeic groups. the ant fauna profile could be completely different with another sampling method. winkler traps may, for example, give different results with a very distinct fauna.diversity differed between sites, as shown by simpson’s diversity index: m3, the ecotonal site, was the most diverse (54 species, n = 1512, d = 3.48) followed by m1 (42 species, n = 1737, d = 2.33), whilst m2 (40 species, n = 823, d = 1.48) and m4 (35 species, n = 775, d = 177) had lower diversity than the other sites. five subfamilies were collected: ponerinae, dolichoderinae, dorylinae, formicinae and myrmicinae. myrmicinae was the most abundant and most diverse subfamily sampled (63 species, n = 4214), comprising 87% of ant abundance and 61% of all species. formicinae was the next most diverse and abundant subfamily (19 species, n = 364), comprising 18% of the total number of ant species sampled (table 1). the least represented subfamilies were dolichoderinae, with six species of tapinoma, and dorylinae, with three species of dorylus. table 1: species richness and abundance of ant subfamilies collected at the marakele national park, limpopo province, south africa. as shown in table 2, the most abundant myrmicinae species were pheidole sp. 1 megacephala group (n = 1137), pheidole sp. 2 capensis group (n = 147), pheidole sculpturata (n = 486), tetramorium quadrispinosum emery (n = 208) and monomorium sp. 4 salomonis group (n = 7780). all these species were common to all the sites. other species that were not so abundant but still occurred across all or most sites were monomorium sp. 3 destructor group, pheidole sp. 2 rugaticeps group, solenopsis sp. 4 and tetramorium sp. 7 similimum group, which is a tramp species that occurred in many different habitats. similarly, t. quadrispinosum co-occurred with the invasive argentine ant (linepithema humile) in studies of plant invasions in the western cape (schoeman & samways 2011). table 2: checklist of ants of the marakele national park, limpopo province, south africa. many species seemed to be localised or restricted to one habitat or altitude, for example calyptomyrmex clavatus, cardiocondyla emeryi, messor denticornis and myrmicaria fusca, as well as some highly localised monomorium, pheidole and tetramorium species. the most abundant formicinae ants were anoplolepis custodiens (n = 35), camponotus sp. 8 rufoglaucus group (n = 125), lepisiota capensis (n = 38) and paratrechina sp. 1 (n = 49). of these, paratrechina sp. 1 was the only species restricted to m3. localised species were less abundant and included camponotus (myrmacrhaphe) sp. 1, camponotus mayri, lepisiota rubrovaria group (both species sampled) and plagiolepis puncta (table 2). all species of the dorylinae and dolichoderinae subfamilies were sampled only in m3, with the exception of tapinoma voeltzkowi forel (found at all sites), tapinoma sp. 2 and dorylus (dorylus) sp. 1. the most abundant ponerine ant was hypoponera similoponera group (n = 102). the remaining ponerine ants were restricted to single sites, with the exception of pachycondyla granosa, which was found in two sites. ponerinae species are generally rarer than other ants, but in the marakele national park the assemblage seems to combine rarity with diversity (12 species, n = 173). it is also worth noting that both the known species of odontomachus occur in the park. some ant species were sampled only in specific vegetation types, indicating that the ant assemblages here differ between the different habitats. nearly half of all sampled species (n = 56; 53%) were restricted to a single site. the percentages of species sampled only in m1, m2, m3, and m4 were 38%, 38%, 33% and 20%, respectively. considerable variation in species assemblage composition was found between sites, with no one site sharing more than 39% of the species. m1 and m2 clustered together with a bray–curtis similarity of 39, whilst m3 and m4 formed a group based on a similarity of 41 (figure 3). m2 and m4 were the least similar (similarity = 25). there was a clear distinction between high (m3 and m4) and low (m2 and m1) elevational ant assemblages according to latitude. discussion top ↑ although sampling lasted only 5 days, it was across a representative altitudinal range and vegetation types occurring in the park. the survey therefore approximates most of the ant genera in the park. inventory completion was satisfactory and suggests that sampling effort was sufficient for the habitats surveyed. approximately 415 ant species are known in south africa (taylor 2005), although there are certainly more. the marakele national park, which covers a land surface of 67 000 ha, hosts approximately a quarter of these species. it is probable that some species found in this study have not previously been recorded in south africa. this observation compares particularly well with the situation in the kruger national park, where 162 species have been recorded over an extended period (parr et al. 2004). the species richness also compares well with that of an altitudinal transect (160 km) in the cederberg mountains in the western cape (botes et al. 2006), in which 86 species were recorded using substantially more intensive sampling (> 50 000 specimens).the survey reported here provides comprehensive baseline data across the altitudinal sweep and vegetation types at the marakele national park. based on the low bray–curtis similarities (i.e. < 40), there is considerable turnover between sites. this highlights the importance of conserving different vegetation types for the maintenance of ant diversity in marakele. the two most diverse sites were m1 and m3. m1 was vegetationally complex, with more trees and more canopy cover than the other (figure 2). m3 was situated on an ecotone between waterberg mountain bushveld and waterberg–magalies summit sourveld (figure 2), which may account for the high diversity at this particular site. this suggests that the physical structure of the vegetation (and hence habitat) may play a role in promoting greater ant species richness (lubertazzi & tschinkel 2003). however, results obtained by lassau and hochuli (2004) showed that ant species richness was negatively associated with ground herb cover, tree canopy cover, soil moisture and leaf litter. during the course of sampling, we also noticed that there was a greater degree of large herbivore trampling in these sites (m1 for instance was close to a rhino latrine). other studies have shown that large herbivore trampling stimulates invertebrate diversity by creating a more heterogeneous habitat (tscharntke & greiler 1995). m2 and m4 yielded a lower species richness than the other sites. the low diversity at m4 may be accounted for by the fact that it is situated at the highest peak in the waterberg. studies have shown that diversity decreases with increasing elevation (mccoy 1990) owing to a decrease in net primary productivity. for the current study, however, there seems to be no clear relationship between elevation and diversity, as our most diverse site was located at 1587 m a.s.l., being the third highest site in the study. with regard to community composition and similarity, m1 and m2, and m3 and m4 clustered together, respectively (figure 3). however, as stated before, the turnover between these sites is considerable. we could not deduce any discernable patterns of higher taxonomic level dissimilarity between the sites, except for m4, which displayed reduced myrmicinae ant species richness, specifically with reference to monomorium species. m3 displayed a greater diversity and abundance of dolichoderinae and dorylinae ants than the other sites, where these subfamilies were poorly represented. so far, though, studies indicate that the physical structure of the habitat can have a distinct effect on composition of the ant communities, affecting niche differentiation and distribution within these habitats (legendre, borcard & peres-neto 2005). in marakele, the presence of large herbivores, combined with vegetational complexity, plays a role in the actual generation of ant species richness within sites, whilst community composition here is largely determined by vegetation type. more realistically, ant community assemblages function as important ecosystem engineers that change a habitat whilst being changed by the habitat in a functionally changing ecosystem. the survey was conducted over a relatively short period using only one sampling technique and therefore our data may not be a true reflection of the total ant species diversity in the park. further studies should include long-term monitoring at a site and management objectives in design of the sampling protocol. surveys of epigaeic ants suggest that abundances and assemblage composition differ between biomes not only as a result of change in vegetation (botes et al. 2006) but also in response to net primary productivity (kaspari, o’donnell & kercher 2000). whether ant community composition differs between vegetation types or units within a biome still needs to be determined; thus far our results suggest that this may be the case. however, we caution that previous studies have shown that differences in invertebrate community assemblage of replicates in the same vegetation type might differ more from each other than assemblages in other vegetation types in the same biome (muelelwa et al. 2010). our assumptions of what exactly constitutes a habitat would therefore need reassessment. conclusion top ↑ our results suggest that the marakele national park, and by extension the waterberg region, is a hotspot for ant diversity in south africa. it may be worth testing whether the groups identified by mcgeoch and sithole (2009) can act as surrogates for one another in any projected future monitoring programmes. to this end, detailed and freely available inventories of ants, butterflies, dragonflies and scarab beetles need to be compiled for all protected areas in south africa. so far, an inventory of spiders has been compiled as part of sansa. such inventories will not only make a significant contribution to existing knowledge of insects in south africa’s protected areas, but will also facilitate movement towards full-scale use of invertebrates in monitoring and other conservation activities. acknowledgements top ↑ we would like to thank the management of the marakele national park for allowing us to conduct this survey, norbert hahn and ovavhu gelebe for help with field work, and sansa and the research and publications committee (university of venda) for financial support. s.h.f. acknowledges support from the dst-nrf centre of excellence for invasion biology. competing interests the authors declare that they have no financial or personal relationship(s) which may have inappropriately influenced them in writing this paper. authors’ contributions c.s.s. (university of venda) was responsible for preparation and identification of specimens and all calculations. s.h.f. (university of venda) was the project leader and made conceptual contributions to the manuscript. references top ↑ addison, p. & samways, m.j., 2000, ‘a survey of ants (hymenoptera: formicidae) that forage in vineyards in the western cape province, south africa’, african entomology 8(2), 251−260.agosti, d. & johnson, n.f. (eds.), 2005, antbase, viewed 13 march 2010, from antbase.org (version 05/2005). andersen, a.n., 1997, ‘using ants as bioindicators: multiscale issues in ant community ecology’, conservation ecology 1(1), 8, viewed 24 november 2011, from http://www.consecol.org/vol1/iss1/art8 andersen, a.n., 2000, ‘a global ecology of rainforest ants: functional groups in relation to environmental stress and disturbance’, in d. agosti, j.d. majer, l.e. alonso & t.r. schultz (eds.), ants: standard methods for measuring and monitoring biodiversity, pp. 25–34, smithsonian institution press, washington dc. andersen, a.n., fisher, a., hoffman, b.d., read, j.l. & richards, r., 2004, ‘use of terrestrial invertebrates for biodiversity monitoring in australian rangelands, with particular reference to ants’, austral ecology 29, 87−92. http://dx.doi.org/10.1111/j.1442-9993.2004.01362.x arnold, g., 1924, ‘a monograph of the formicidae of south africa. part 6. (camponotinae)’, annals of the south african museum 14, 675–766. bolton, b., 1984, identification guide to ant genera of the world, harvard university press, cambridge, ma. boonzaaier, c., mcgeoch, m.a. & parr, c.l., 2007, ‘fine-scale temporal and spatial dynamics of epigaeic ants in fynbos: sampling implications’, african entomology 15(1), 1−11. http://dx.doi.org/10.4001/1021-3589-15.1.1 botes, a., mcgeoch, m.a., robertson, h.g., van niekerk, a., davids, h.p. & chown, s.l., 2006, ‘ants, altitude and change in the northern cape floristic region’, journal of biogeography 33, 71−90. http://dx.doi.org/10.1111/j.1365-2699.2005.01336.x colwell, r.k. & coddington, j.a., 1994, ‘estimating terrestrial biodiversity through extrapolation’, philosophical transactions of the royal society of london b 345, 101−108. http://dx.doi.org/10.1098/rstb.1994.0091, pmid:7972351 dippenaar-schoeman, a.s. & craemer, c., 2000, ‘the south african national survey of arachnida’, plant protection news 56, 11−12. fisher, b.l., 2004, ‘diversity patterns of ants (hymenoptera: formicidae) along an elevational gradient on monts doudou in southwestern gabon’, california academy of sciences memoir 28, 269–286. folgarait, p.j., 1998, ‘ant biodiversity and its relationship to ecosystem functioning: a review’, biodiversity and conservation 7(9), 1221−1244. http://dx.doi.org/10.1023/a:1008891901953 gaigher, r., 2008, ‘the effect of different vineyard management systems on the epigaiec arthropod assemblages in the cape floristic region, south africa’, msc thesis, dept. of conservation ecology and entomology, university of stellenbosch. garcia, f.h., fischer, g., peters, m.k., snelling, r.r. & wägele, j.w., 2009, ‘a preliminary checklist of the ants (hymenoptera: formicidae) of kakamega forest (kenya)’, journal of east african natural history 98(2), 147–165. gardiner, a.j. & terblanche, r.f., 2010, ‘taxonomy, biology, biogeography, evolution and conservation of the genus eriksonia trimen (lepidoptera: lycaenidae)’, african entomology 18(1), 171−191. http://dx.doi.org/10.4001/003.018.0114 giliomee, j.h., 2003, ‘insect diversity in the cape floristic region’, african journal of ecology 41(3), 237−244. http://dx.doi.org/10.1046/j.1365-2028.2003.00442.x greenslade, p.j.m., 1973, ‘sampling ants with pitfall traps: digging-in effects’, insectes sociaux 20(4), 343–353. http://dx.doi.org/10.1007/bf02226087 hölldobler, b. & wilson, e.o., 1994, the ants, springer-verlag, berlin. kaspari, m. & majer, j.d., 2000, ‘using ants to monitor environmental change’, in d. agosti, j.d. majer, l.e. alonso & t.r. schultz (eds.), ants: standard methods for measuring and monitoring biodiversity, pp. 89–98, smithsonian institution press, washington dc. kaspari, m., o’donnell, s. & kercher, j.r., 2000, ‘energy, density, and constraints to species richness: ant assemblages along a productivity gradient’, the american naturalist 155, 280−293. http://dx.doi.org/10.1086/303313, pmid:10686166 koch, f. & vohland, k., 2004, ‘ants along a southern african transect a basis for biodiversity change monitoring (insecta, hymenoptera, formicidae)’, zoosystematics and evolution 80(2), 261–273. kremen, c., colwell, r.k., erwin, t.l., murphy, d.d., noss, r.f. & sanjayan, m.a., 1993, ‘terrestrial arthropod assemblages: their use in conservation planning’, conservation biology 7(4), 796−808. http://dx.doi.org/10.1046/j.1523-1739.1993.740796.x lassau, s.a. & hochuli, d.f, 2004, ‘effects of habitat complexity on ant assemblages: can we generalise across scale?’, ecography 27(2), 157−164. http://dx.doi.org/10.1111/j.0906-7590.2004.03675.x lattke, j.e., 2000, ‘specimen processing: building and curating an ant collection’, in d. agosti, j.d. majer, l.e. alonso & t.r. schultz (eds.), ants: standard methods for measuring and monitoring biodiversity, pp. 155–171, smithsonian institution press, washington dc. legendre, p., borcard, d. & peres-neto, p.r., 2005, ‘analyzing beta diversity: partitioning the spatial variation in community composition data’, ecological monographs 75(4), 435–450. http://dx.doi.org/10.1890/05-0549 lévieux, j., 1972, ‘le rôle des fourmis dans les réseaux trophiques d’une savane préforestière de côte d’ivoire [the role of ants in a trophic network in a preforested savanna in the ivory coast]’, annales de l’université d’abidjan ser. e, 5, 143–240. lingyel, s., gove, a.d., latimer, a.m., majer, j.d. & dunn, r.r., 2009, ‘convergent evolution of seed dispersal by ants, and phylogeny and biogeography in flowering plants: a global review’, perspectives in plant ecology, evolution and systematics 12(1), 43−55. http://dx.doi.org/10.1016/j.ppees.2009.08.001 lubertazzi, d. & tschinkel, d.r., 2003, ‘ant community change across a ground vegetational gradient in north florida’s longleaf pine flatwoods’, journal of insect science 3, 21, viewed 24 november 2011, from http://www.insectscience.org/3.21, pmid:15841237, pmcid:524660 marsh, a.c., 1986, ‘checklist, biological notes and distribution of ants in the central namib desert’, madoqua 14(4), 333–344. mccoy, e.d., 1990, ‘the distribution of insects along elevational gradients’, oikos 58(3), 313–322, viewed 04 may 2011, from http://www.jstor.org/pss/3545222 mcgeoch, m.a., sithole, h., samways, m.j., simaika, j.p., pryke, j.s., picker, m. et al., 2011, ‘conservation and monitoring of invertebrates in terrestrial protected areas’, koedoe 53, art. #1000. http://dx.doi.org/10.4102/koedoe.v53i2.1000 mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, south african national biodiversity institute, pretoria. (strelitzia, no. 19) muelelwa, m.i., foord, s.h., dippenaar-schoeman, a.s. & stam, e.m., 2010, ‘towards a standardized and optimized protocol for rapid biodiversity assessments: spider species richness and assemblage composition in two savanna vegetation types’, african zoology 45(2), 273–290. http://dx.doi.org/10.3377/004.045.0206 parr, c.l. & chown, s.l., 2003, ‘burning issues in conservation: a critique of faunal fire research in southern africa’, austral ecology 28, 384–395. http://dx.doi.org/10.1046/j.1442-9993.2003.01296.x parr, c.l., 2003, ‘ant assemblages in a southern african savanna: local processes and conservation implications’, phd thesis, dept. of entomology, university of pretoria. parr, c.l., bond, w.j. & robertson, h.g., 2002, ‘a preliminary study of the effect of fire on ants (formicidae) in a south african savanna’, african entomology 10, 101–111. parr, c.l., robertson, h.g., biggs, h.c. & chown, s.l., 2004, ‘response of african savanna ants to long-term fire regimes’, journal of applied ecology 41(4), 630–632. http://dx.doi.org/10.1111/j.0021-8901.2004.00920.x pierce, n.e., braby, m.f., heath, a., lohman, d.j., mathew, j., rand, d.b. et al., 2002, ‘the ecology and evolution of ant association in the lycaenidae (lepidoptera)’, annual review of entomology 47, 733−771. http://dx.doi.org/10.1146/annurev.ento.47.091201.145257, pmid:11729090 samways, m.j., 2005, insect diversity conservation, cambridge university press, cambridge. http://dx.doi.org/10.1017/cbo9780511614163 schoeman, c.s. & samways, m.j., 2011, ‘synergisms between the invasive alien argentine ant and alien plants on native ants of the western cape, south africa’, african entomology 19(1), 96–105. seymour, c.l. & dean, w.r.j., 1999, ‘effects of heavy grazing on invertebrate assemblages in the succulent karoo, south africa’, journal of arid environments 43, 267−286. http://dx.doi.org/10.1006/jare.1999.0552 sileshi, g.w., arshad, m.a., konate, s. & nkunika, p.o.y., 2010, ‘termite-induced heterogeneity in african savanna vegetation: mechanisms and patterns’, journal of vegetation science 21(5), 923−937. http://dx.doi.org/10.1111/j.1654-1103.2010.01197.x szüts, t. & jocqué, r., 2001, ‘a revision of the afrotropical spider genus palfuria (araneae: zodariidae)’, journal of arachnology 29, 205−219. http://dx.doi.org/10.1636/0161-8202(2001)029[0205:arotas]2.0.co;2 taylor, b., 2005, the ants of (sub-saharan) africa, viewed 29 june 2010, from http://antbase.org/ants/africa tscharntke, t. & greiler, h.j., 1995, ‘insect communities, grasses and grasslands’, annual review of entomology 40, 535–558. http://dx.doi.org/10.1146/annurev.en.40.010195.002535 uys, c., hamer, m. & slotow, r., 2010, ‘step process for selecting and testing surrogates and indicators of afromontane forest invertebrate diversity’, plos one 5(2), e1900, viewed on 03 october 2010, from http://www.plosone.org/article/info%3adoi%2f10.1371%2fjournal.pone.0009100 willis, c.k., skinner, j.d. & robertson, h.g., 1992, ‘abundance of ants and termites in the false karoo and their importance in the diet of the aardvark orycteropus afer’, african journal of ecology 30(4), 322–334. http://dx.doi.org/10.1111/j.1365-2028.1992.tb00509.x yek, s.h., williams, s.e., burwell, c.j., robson, s.k.a. & crozier, r.h., 2009, ‘ground dwelling ants as surrogates for establishing conservation priorities in the australian wet tropics’, journal of insect science 12, 1−12. http://dx.doi.org/10.1673/031.009.1201, pmid:19613441, pmcid:3011884 article information authors: robin m. cook1 michelle d. henley2,3 francesca parrini1 affiliations: 1school of animal, plant and environmental sciences, university of the witwatersrand, south africa 2applied behavioural ecology and ecosystem research unit, university of south africa, south africa 3elephants alive, hoedspruit, south africa correspondence to: robin cook email: robinmichaelcook@gmail.com postal address: private bag x3, university of the witwatersrand 2050, south africa dates: received: 05 feb. 2015 accepted: 17 aug. 2015 published: 17 nov. 2015 how to cite this article: cook, r.m., henley, m.d. & parrini, f., 2015, ‘elephant movement patterns in relation to human inhabitants in and around the great limpopo transfrontier park’, koedoe 57(1), art. #1298, 7 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1298 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. elephant movement patterns in relation to human inhabitants in and around the great limpopo transfrontier park in this original research... open access • abstract • introduction • research method and design    • study area    • elephants and tracking data    • data analysis       • seasonal locations around villages       • time-of-day differences       • mean linear displacement rates       • statistical analyses • results    • seasonal locations around villages    • time-of-day differences    • mean linear displacement rates • discussion • conclusion and management implications • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ the presence of humans and african elephants (loxodonta africana) in the great limpopo transfrontier park can create situations of potential human–elephant conflict. such conflict will likely be exacerbated as elephant and human populations increase, unless mitigation measures are put in place. in this study we analysed the movement patterns of 13 collared adult african elephants from the northern kruger national park over a period of eight years (2006–2014). we compared the occurrence and displacement rates of elephant bulls and cows around villages in the limpopo national park and northern border of the kruger national park across seasons and at different times of the day. elephants occurred close to villages more often in the dry season than in the wet season, with bulls occurring more frequently around villages than cows. both the bulls and the cows preferred to use areas close to villages from early evening to midnight, with the bulls moving closer to villages than the cows. these results suggest that elephants, especially the bulls, are moving through the studied villages in mozambique and zimbabwe at night and that these movements are most common during the drier months when resources are known to be scarce. conservation implications: elephants from the kruger national park are moving in close proximity to villages within the great limpopo transfrontier park. resettlement of villages within and around the park should therefore be planned away from elephant seasonal routes to minimise conflict between humans and elephants. introduction top ↑ conflict between humans and elephants (loxodonta africana) has been investigated throughout africa, focusing on elephant crop raiding activities (o’connell-rodwell et al. 2000), mapping elephant movement patterns around human settlements (galanti et al. 2006) and investigating possible corridors along which elephants can move safely between reserves (van de perre et al. 2014). however, as human and elephant populations increase, both in and around protected areas, the likelihood of interactions between humans and elephants, as well as the levels of human–elephant conflict (hec), will increase. in southern africa, elephants can move freely throughout the great limpopo transfrontier park (gltp), owing to fences being dropped between south africa's kruger national park (knp), mozambique's limpopo national park (lnp) and zimbabwe's gonarezhou national park (gnp). however, hec is prevalent within this region as humans and elephants share the same landscape and compete for the same resources (dunham et al. 2010; witter 2013). residents have reported that the presence of elephants has placed restrictions on their livelihoods: they are scared of moving between villages and elephants raid their crops (milgroom & spierenburg 2008; witter 2013). in addition, six elephants were shot within the lnp between 2006 and 2008 as a consequence of hec (dunham et al. 2010). hec is further complicated by residents’ unwillingness to be moved to areas outside the gltp (lunstrum 2015; milgroom & spierenburg 2008; witter 2013). global positioning system (gps) technology has been widely used in elephant movement pattern studies since the mid-1990s (douglas-hamilton 1998). this technology allows scientists to track animal positions over time and relate these positions to the animals’ environment (cagnacci et al. 2010). animal movements are influenced by both external factors such as resource availability and internal factors such as reproductive status or fear (nathan et al. 2008). environmental factors such as water distribution (smit, grant & devereux 2007; thomas, holland & minot 2011), vegetation greenness (marshal et al. 2011) and shifts in rainfall patterns (birkett et al. 2012) all drive elephant movements. however, in this study we focus on how the anthropogenic environment within and around the gltp affects elephant movement patterns. the way elephants use the areas surrounding villages will depend on their spatial and temporal knowledge of human activities, such as when crops present alternative food sources, when humans are least active or how humans react to their presence (hoare & du toit 1999). as bulls have different nutrient requirements from cows, preferring bulky diets over quality of vegetation (greyling 2004; shannon et al. 2006), the body size hypothesis predicts sexual segregation in bull and cow movement patterns outside of mating events (stokke & du toit 2000). cows are responsible not only for feeding the young elephants nutrient-rich milk during lactation, but also for regulating the pace of movement to that of the young elephants to ensure their safety (duffy et al. 2011; shannon et al. 2010). we therefore expect differences in movement patterns to occur between the sexes. specifically, the objectives of our study are (1) to analyse the seasonal occurrences of bulls and cows at different distances from villages, (2) to compare the distance at which bulls and cows occur relative to villages over a 24-hour period and (3) to calculate and compare the displacement rates (distance travelled by an elephant between consecutive collar recordings) of bulls and cows at different distances from villages. research method and design top ↑ study area the study area falls within the gltp, an area of 35 000 km² that incorporates the knp, lnp and gnp. we focused on villages within the lnp and bordering the limpopo river of the knp of the two study sites, mozambique and zimbabwe, respectively (figure 1). the gltp is a semi-arid region with a mean annual rainfall of 440 mm in the pafuri land system (venter & gertenbach 1986). the dominant vegetation communities include colophospermum mopane woodland and forest, ficus sycomorus forest, and burkea africana and combretum apiculatum woodland. tree species such as diospyros mespiliformis, combretum imberbe and kigelia africana are widespread along the limpopo and luvuvhu rivers (venter, scholes & eckhardt 2003). figure 1: location of villages and community zones within mozambique's limpopo national park and along the limpopo river of south africa's kruger national park. the gltp is primarily surrounded by small villages along its borders (hughes 2005). each village community has an estimated population size of around 1000 individuals (holden 2001; lunstrum 2015). the study area included villages situated along the zimbabwean border of the knp and villages within the lnp. there are around 27 000 residents within the study area (milgroom & spierenburg 2008). elephants can move freely from the pafuri region in the knp to the vicinity of these villages. villages are situated 5 km – 30 km apart, with each village assigned a community radius of 4 km for crop fields, which comprise maize, beans, tomato, onions, cabbage and carrots, and grazing land for cattle (bos taurus) (a. alexander pers. comm., 17 july 2014; t. chauque pers. comm., 12 march 2015). elephants and tracking data following a near population collapse in the knp region at the turn of the twentieth century, the first elephants were recorded in the knp in 1905, and the first in the pafuri region only after 1945 (whyte 2001). currently, there are approximately 16 700 elephants in the knp (sanparks 2012), which can move freely into the lnp where over 1000 elephants have been recorded (milgroom & spierenburg 2008). in mozambique, where people and elephants compete for resources, there have been reports of crop raiding by elephants (de boer & baquete 1998; de boer, stigter & ntumi 2007; harris et al. 2008) as well as recent reports of intensive illegal killings in mozambique's largest conservation areas (booth & dunham 2014). both illegal killings and human occupancy of their home ranges could affect the elephants’ perception of safety. therefore, to investigate the elephant movements in relation to human occupancy, we used existing data from 13 gps-satellite collars that had been placed on eight bulls and five cows from different breeding herds in the far northern and eastern regions of the knp as part of the elephant monitoring programmes in the region. collaring operations had been carried out by knp wildlife veterinarians using standard operating procedures as approved by the south african national parks animal use and care committee (sanparks 2011). the collars recorded the location of each individual at set eight-hour intervals. elephant tracking data were retrieved from a centralised database using customised software (wall et al. 2014) that employs a data filter to remove erroneous gps fixes based on a maximum rate of travel (see austin, mcmillan & bowen 2003). data were stored in both esri geodatabase (arcgis version 10.1) and comma-separated text file formats. data from 2006 to 2014 were used, with each elephant's collar lasting an average of two to three years (table 1). nine of the collars were replaced during this period (six breeding herd individuals and three solitary bulls) (table 1). the collars recorded the elephant's gps locations, the time of day for each data point, and the linear displacement rate (m/h) at each location. in our analyses, the social units described are bulls and cows. the movement of a bull refers to an adult bull that is either solitary or in a bachelor herd. the movement of an individual cow is assumed to be that of the entire family unit, consisting of adult cows, calves and bulls younger than 15 years. table 1: collar identification codes and the years when the collars were active for the 13 collared elephants used in this study. data analysis distance buffer zones were drawn around villages within the study area. buffer zones included the following areas: anthropogenic infrastructure and fields located within 0 km – 2 km from a village the outer regions of the community zones, 2 km – 4 km from a village, occupied by fields the wilderness zone around the community zone, 4 km – 6 km from a village, which is not used for crop production or grazing. locations within 6 km of a village were analysed for a total of 3524 gps coordinates, of which 1319 were associated with cows and 2205 with bulls. arcmap (arcgis, version 10.1, esri) was used for analysis of the gps locations. seasonal locations around villages for each buffer zone, monthly proportions of bull and cow locations were calculated for the entire study period (2006–2014). mean monthly rainfall data (2006–2013) were used to distinguish between wetter and drier months in the study area in order to compare elephant locations in the early wet season (october–december), late wet season (january–march), early dry season (april–june) and late dry season (july–september). mean monthly rainfall data were acquired from the punda maria and shingwedzi ranger stations. time-of-day differences elephant locations within the three buffer zones were assigned to three time intervals defined by the recording intervals of the elephant's collars. the period from 00:00 to 07:59 was classified as post-midnight to early morning, from 08:00 to 15:59 was classified as late morning to mid-afternoon, and from 16:00 to 23:59 was classified as evening to midnight. we expected that people would likely be the most active in the fields and outside their homes between late morning and mid-afternoon. the proportions of occurrences of bulls and cows within and between the buffer zones were then compared for each period. mean linear displacement rates mean linear displacement rates (referred to as displacement rates from here on) were analysed for each elephant within each buffer zone. displacement rates (m/h) were automatically calculated by the collars as the distance travelled between two consecutive collar recordings (and therefore two locations). displacement rates for bulls and cows were compared between buffer zones. seasonal differences in displacement rates could not be analysed as a result of the small number of locations close to villages during the wet season. statistical analyses all statistical analyses were performed using statistica 8.0 (http://www.statsoft.com). a pearson's chi-squared test was used to test for independence of the mozambican and zimbabwean village rainfall and elephant occurrence data. generalised linear mixed models were applied to both the location and linear displacement rate data. to test the responses of elephant location points as a function of social unit, distance from villages and the time of day, the proportions of location points were used as the dependent variable, with the social unit, buffer zones and times set as fixed factors. the individual elephants were regarded as random factors. to test the responses of linear displacement rates as a function of social unit and distance from villages, the linear displacement rates were used as the dependent variable, with the social unit and buffer zones set as fixed factors. the individual elephants were regarded as random factors. results top ↑ seasonal locations around villages monthly locations were combined for villages along the mozambican and zimbabwean borders, as there was no significant difference between the rainfall data at the two border sites (χ2 = 4.07, df = 2, p > 0.05) and the seasonal elephant patterns (χ2 = 1.02, df = 1, p > 0.05) for the two sites. across the study period, the proportion of elephant locations began to increase during the early dry season in all buffer zones (april–may) as mean monthly rainfall across the study area decreased (figure 2a and 2b). the highest proportion of locations in all three buffer zones occurred during the late dry season (august–october), when the mean monthly rainfall was at its minimum, for both bulls and cows (figure 2a and 2b). the lowest proportion of locations within buffer zones occurred in the late wet season (january–march), when the mean monthly rainfall was at its peak, for both bulls and cows in all buffer zones (figure 2a and 2b). figure 2: monthly location proportions (as measured by global positioning system) of elephant (a) bulls and (b) cows within three buffer zones around studied villages in relation to average monthly rainfall data (2006–2013). time-of-day differences the time recordings of elephant locations in the two study sites were combined as there was no significant difference between recordings at the two sites (three-way interaction term f4,42 = 0.53; p > 0.05). there was a higher proportion of elephant locations in the 0 km – 2 km and 2 km – 4 km buffer zones during the evening–midnight period than in the other periods. a significantly higher proportion of locations was recorded from bulls in the 0 km – 2 km buffer zone during the evening–midnight period than in the other periods (table 2). none of the collared cows used the 0 km – 2 km buffer zone, but rather remained on the outskirts of the community zones around the villages. in the 2 km – 4 km buffer zone, location proportions of both the bulls and the cows were significantly higher in the evening–midnight period than in the two other periods (table 2). there was no significant difference between the time proportions for bulls and cows in this buffer zone. in the 4 km – 6 km buffer zone, there was no significant difference in the location proportions of bulls and cows across the three periods (table 2). table 2: proportion of elephant location points during three periods across the buffer zones surrounding villages in mozambique's limpopo national park and villages bordering the limpopo river of south africa's kruger national park (± standard error). mean linear displacement rates displacement rates of bulls were significantly higher than those of breeding herds around villages (313 m/h for bulls vs 208 m/h for cows) (f1,10 = 5.13; p < 0.05; figure 3). the displacement rate of bulls increased significantly the closer the bulls were to villages, with a mean displacement rate of 500 m/h in the 0 km – 2 km buffer zone, 335 m/h in the 2 km – 4 km buffer zone and 287 m/h in the 4 km – 6 km buffer zone (f2,10 = 6.41; p < 0.05; figure 3). there was significant variation between the displacement rates of the individual bulls in the three buffer zones, and although all of the bulls increased their displacement rates the closer they were to villages, there was variation in the changes of displacement rate when approaching a village (two-way interaction term f10,2186 = 3.69; p < 0.0001). one of the bulls moving around the villages makandazulo a and b decreased its displacement rate when moving from the 4 km – 6 km buffer zone (260 m/h) into the 2 km – 4 km buffer zone (208 m/h). however, this bull's displacement rate again increased when in the 0 km – 2 km buffer zone (329 m/h). furthermore, three of the bulls had displacement rates above 350 m/h (mean = 380 m/h), which was significantly higher than the mean displacement rate for the others (231 m/h) (f1,3 = 41.04; p < 0.005). cows’ displacement rates increased from 205 m/h in the 4 km – 6 km buffer zone to 224 m/h in the 2 km – 4 km buffer zone, albeit not significantly (f1,2 = 13.88; p = 0.07; figure 3). there was no significant variation in the displacement rates of individual cows in the respective buffer zones. figure 3: mean linear displacement rate of bulls and cows across buffer zones surrounding studied villages. error bars indicate standard error. asterisks indicate significantly different displacement rates (p < 0.05). discussion top ↑ we found that the majority of elephant locations in village community zones occurred during the drier months. previous studies have suggested an increase in elephant movements around villages as a result of decreased water availability in protected reserves (kioko, okello & muruthi 2006), an increase in nutrient quality available in villages (o’connell-rodwell et al. 2000) and a decrease in the natural vegetation quality in protected reserves (osborn 2004). in the context of the gltp, no single explanation can be given for the observed higher frequency of elephant locations close to villages without real-time observations during the dry season. most of the villages are located along major river systems, including the rio limpopo (limpopo river), the rio shingwedzi (shingwedzi river) and the rio dos elefantes (olifants river). in contrast to the neighbouring knp, there are no artificial water points in the lnp and elephants appear to concentrate their movements through the lnp along river systems (unpublished data). furthermore, motor and hand-held pumps have been provided to river-bordering villages in the region to enable residents to grow crops throughout the year without having to rely on rainfall (a. alexander, pers. comm., 17 july 2014). elephant locations close to villages may therefore be due to a combination of water being available close to the crops and crops offering accessible and diverse resources (depending on what crop type is being cultivated). during the drier months, when natural resources become scarce, elephants will be more prone to supplement their diets with cultivated crops until the nutritional quality from natural vegetation increases in the wetter months. osborn (2004) found similar behavioural patterns for elephants in northern zimbabwe and witter (2013) mentions elephants utilising not only crops but also fruit grown and stored around villages in the lnp. we found that only the bulls used the 0 km – 2 km buffer zone. bulls have been found to be more likely to move within closer proximity to villages than cows, with hypotheses suggesting that the increased intake of higher quality crops enables the bulls to compete more successfully against other bulls for dominance status and mating opportunities (chiyo et al. 2011; sitati et al. 2003; sukumar & gadgil 1988). although the presence of crops would also be attractive to cows and their smaller family members, the small number of cow locations in the closest community zone could simply be a result of the risk posed by humans coupled with an inability to move to safety as quickly as bulls, who generally have smaller group sizes and whose movements are not constrained to the same level by the slowest members of the group, usually calves (duffy et al. 2011; shannon et al. 2010). despite the presence of smaller-bodied cows, who are often pregnant or lactating and therefore in need to maintain diets of higher quality compared to bulls (stokke & du toit 2000), safety aspects may outweigh the nutrient value of the crops when considering family units. the predation-risk theory predicts that cows with young are more vulnerable to predation than bulls and are therefore less likely to use areas of potential danger (ruckstuhl & neuhaus 2002). the predation-risk theory may explain why elephant breeding herds are less inclined to use areas around villages in comparison to bulls, as the villages may present a potential danger to the cows and their calves. in addition, solitary bulls may be less likely to be detected by humans when crop raiding than a breeding herd with more individuals (sukumar & gadgil 1988). crop raiding may therefore be a more attractive activity for solitary bulls able to avoid human observation. the majority of elephant locations in the 0 km – 2 km and 2 km – 4 km buffer zones occurred in the evening–midnight period, which suggests that the elephants are closest to villages when humans are least active. evening–midnight location trends have been recorded both for elephants moving past villages (douglas-hamilton, krink & vollrath 2005; graham et al. 2009; van de perre et al. 2014) and for actively crop-raiding elephants (chiyo & cochrane 2005; jackson et al. 2008; sukumar & gadgil 1988). the low proportion of locations in the community zones between midnight and early morning may be attributed to elephants moving out of the community zones during this period to rest or sleep. rest periods are associated with less vigilant behaviour, which is easier when not in a potential landscape of fear (wing & buss 1970). however, more information is required based on gps locations recorded at shorter time intervals to test for such fine-scale activity patterns of elephants. the increased displacement rates of bulls moving through the community zones suggest that the displacement rates of elephants increase when moving through areas of potential high risk. this increased displacement rate was particularly evident in the 0 km – 2 km buffer zone, which is occupied by village infrastructure and residents. elephants that move through a community zone with the sole purpose of minimising the time spent in that zone will be likely to have increased displacement rates as their movement patterns are more likely directional (douglas-hamilton et al. 2005; jachowski, slotow & millspaugh 2013). if an elephant is crop raiding, its displacement rate will be lower than that of an elephant moving through in a directional manner, as the distance covered between locations will be minimal. this does not necessarily suggest that the elephant's speed has decreased (graham et al. 2009), as a crop-raiding elephant may be moving at an increased speed amongst crops in a non-directional and tortuous manner to maximise resource intake. therefore, the variation in displacement rates seen between bulls moving through community zones could mean that not all elephants moving through a village carry out the same activity or respond to the presence of a village in the same manner (mccomb et al. 2011). musth is known to influence the movement rates of bulls (rasmussen, wittemyer & douglas-hamilton 2005) and hence individual variability between bulls could also be linked to their reproductive status at the time of crop raiding. although the displacement rate of the cows increased when moving through the community zones, there was no significant change between displacement rates within a buffer zone or the displacement rates of different cows, presumably as cows are equally constrained by their group sizes and can maintain movement at a constant rate when not in the highest danger zone (0 km – 2 km). conclusion and management implications top ↑ elephants from the knp are moving in close proximity to villages within the gltp. this is the first known study focusing on the movement patterns of elephants around villages in the gltp and provides baseline information on how elephants in the gltp use the areas around villages. elephants evidently have to play off the benefits derived from crop raiding against those of being persecuted by humans. bulls and cows were found to handle these treat-versus-threat scenarios differently, with bulls being less risk averse than family units. further understanding of the movements of elephant bulls and breeding herds around villages within this area requires a more in-depth approach. focusing on the human density, crop layout, water availability and previous experience with hec will benefit our understanding of how these factors influence elephant movement patterns. by understanding where, when and in what manner elephants are moving around these villages, conservation authorities can begin to adopt mitigation systems such as beehive fences (king, douglas-hamilton & vollrath 2011) or chili extracts (osborn & rasmussen 1995) in an attempt to minimise hec in the region. in addition, resettlement of villages within and around the gltp should take place away from seasonal elephant routes to prevent conflict between humans and elephants in this region. acknowledgements top ↑ we are grateful for the assistance and information provided by the members of staff from both the lnp and the gnp regarding the villages and their interactions with elephants. the us fish and wildlife services (usfws), wilderness safaris, jerry cohen, madeleine delman and marlene mccay are thanked for financial support (collars, collaring costs and various other financial aspects of the ste–sa transboundary elephant research programme). the knp is thanked for logistical support during collaring operations. craig beech from the peace parks foundation is thanked for supplying gis layers for the lnp. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions r.m.c. (university of the witwatersrand) was responsible for analysis and interpretation of the data. he was also responsible for drafting the article. m.d.h. (university of south africa) was responsible for data acquisition and contributed to data interpretation and article revision. f.p. (university of the witwatersrand) was the project leader. she contributed to the concept and design of the study, supervised data analysis and interpretation, and revised the article. references top ↑ austin, d., mcmillan, j.i. & bowen, w.d., 2003, ‘a three-stage algorithm for filtering erroneous argos satellite locations’, marine mammal science 19(2), 371–383. http://dx.doi.org/10.1111/j.1748-7692.2003.tb01115.x birkett, p.j., vanak, a.t., muggeo, v.m., ferreira, s.m. & slotow, r., 2012, ‘animal perception of seasonal thresholds: changes in elephant movement in relation to rainfall patterns’, plos one 7(6), e38363. pmid: 22761680, http://dx.doi.org/10.1371/journal.pone.0038363 booth, v.r. & dunham, k.m., 2014, ‘elephant poaching in niassa reserve, mozambique: population impact revealed by combined survey trends for live elephants and carcasses’, oryx, 10 pages. http://dx.doi.org/10.1017/s0030605314000568 cagnacci, f., boitani, l., powell, r.a. & boyce, m.s., 2010, ‘animal ecology meets gps-based radiotelemetry: a perfect storm of opportunities and challenges’, proceedings of the royal society b 365(1550), 2157–2162. pmid: 20566493, http://dx.doi.org/10.1098/rstb.2010.0107 chiyo, p.i. & cochrane, e.p., 2005, ‘population structure and behaviour of crop-raiding elephants in kibale national park, uganda’, african journal of ecology 43(3), 233–241. http://dx.doi.org/10.1111/j.1365-2028.2005.00577.x chiyo, p.i., lee, p.c., moss, c.j., archie, e.a., hollister-smith, j.a. & alberts, s.c., 2011, ‘no risk, no gain: effects of crop raiding and genetic diversity on body size in male elephants’, behavioral ecology 22(3), 552–558. http://dx.doi.org/10.1093/beheco/arr016 de boer, w.f. & baquete, d.s., 1998, ‘natural resource use, crop damage and attitudes of rural people in the vicinity of the maputo elephant reserve, mozambique’, environmental conservation 25(3), 208–218. de boer, w.f., stigter, j.d. & ntumi, c.p., 2007, ‘optimising investments from elephant tourist revenues in the maputo elephant reserve, mozambique’, journal for nature conservation 15(4), 225–236. http://dx.doi.org/10.1016/j.jnc.2006.11.002 douglas-hamilton, i., 1998, ‘tracking african elephants with a global positioning system (gps) radio collar’, pachyderm 25, 81–92. douglas-hamilton, i., krink, t. & vollrath, f., 2005, ‘movements and corridors of african elephants in relation to protected areas’, naturwissenschaften 92(4), 158–163. pmid: 15770465, http://dx.doi.org/10.1007/s00114-004-0606-9 duffy, k.j., dai, x., shannon, g., slotow, r. & page, b., 2011, ‘movement patterns of african elephants (loxodonta africana) in different habitat types’, south african journal of wildlife research 41(1), 21–28. http://dx.doi.org/10.3957/056.041.0107 dunham, k.m., ghiurghi, a., cumbi, r. & urbano, f., 2010, ‘human–wildlife conflict in mozambique: a national perspective, with emphasis on wildlife attacks on humans’, oryx 44(2), 185–193. http://dx.doi.org/10.1017/s003060530999086x galanti, v., preatoni, d., martinoli, a., wauters, l.a. & tosi, g., 2006, ‘space and habitat use of the african elephant in the tarangire–manyara ecosystem, tanzania: implications for conservation’, mammalian biology 71(2), 99–114. http://dx.doi.org/10.1016/j.mambio.2005.10.001 graham, m.d., douglas-hamilton, i., adams, w.m. & lee, p.c., 2009, ‘the movement of african elephants in a human-dominated land-use mosaic’, animal conservation 12(5), 445–455. http://dx.doi.org/10.1111/j.1469-1795.2009.00272.x greyling, m.d., 2004, ‘sex and age related distinctions in the feeding ecology of the african elephant, loxodonta africana’, phd thesis, school of animal, plant, & environmental sciences, university of the witwatersrand. harris, g.m., russel, g.j., van aarde, r.i. & pimm, s.l., 2008, ‘rules of habitat use by elephants loxodonta africana in southern africa: insights for regional management’, oryx 42(1), 66–75. http://dx.doi.org/10.1017/s0030605308000483 hoare, r.e. & du toit, j.t., 1999, ‘coexistence between people and elephants in african savannas’, conservation biology 13(3), 633–639. http://dx.doi.org/10.1046/j.1523-1739.1999.98035.x holden, p., 2001, ‘land use planning of coutada 16: part of the gaza-kruger-gonarezhou transfrontier park’, report, peace parks foundation. hughes, d.m., 2005, ‘third nature: making space and time in the great limpopo conservation area’, cultural anthropology 20(2), 157–184. http://dx.doi.org/10.1525/can.2005.20.2.157 jachowski, d.s., slotow, r. & millspaugh, j.j., 2013, ‘corridor use and streaking behaviour by african elephants in relation to physiological state’, biological conservation 167, 276–282. http://dx.doi.org/10.1016/j.biocon.2013.08.005 jackson, t.p., mosojane, s., ferreira, s.m. & van aarde, r.j., 2008, ‘solutions for elephant loxodonta africana crop raiding in northern botswana: moving away from symptomatic approaches’, oryx 42(1), 83–91. http://dx.doi.org/10.1017/s0030605308001117 king, l.e., douglas-hamilton, i. & vollrath, f., 2011, ‘beehive fences as effective deterrents for crop-raiding elephants: field trials in northern kenya’, african journal of ecology 49(4), 431–439. http://dx.doi.org/10.1111/j.1365-2028.2011.01275.x kioko, j., okello, m. & muruthi, p., 2006, ‘elephant numbers and distribution in the tsavo-amboseli ecosystem, south-western kenya’, pachyderm 40, 61–68. lunstrum, e., 2015, ‘green grabs, land grabs and the spatiality of displacement: eviction from mozambique's limpopo national park’, area, 11 pages. http://dx.doi.org/10.1111/area.12121 marshal, j.p., rajah, a., parrini, f., henley, m., henley, s.r. & erasmus, b.f., 2011, ‘scale-dependent selection of greenness by african elephants in the kruger-private reserve transboundary region, south africa’, european journal of wildlife research 57(3), 537–548. http://dx.doi.org/10.1007/s10344-010-0462-1 mccomb, k., moss, c., durant, s.m., baker, l. & sayialel, s., 2001, ‘matriarchs as repositories of social knowledge in african elephants’, science 292(5516), 491–494. pmid: 11313492, http://dx.doi.org/10.1126/science.1057895 mccomb, k., shannon, g., durant, s.m., sayialel, k., slotow, r., poole, j. et al., 2011, ‘leadership in elephants: the adaptive value of age’, proceedings of the royal society b: biological sciences 278(1722), 3270–3276. pmid: 21411454, http://dx.doi.org/10.1098/rspb.2011.0168 milgroom, j. & spierenburg, m., 2008, ‘induced volition: resettlement from limpopo national park, mozambique’, journal of contemporary studies 26(4), 435–448. http://dx.doi.org/10.1080/02589000802482021 nathan, r., getz, w., revilla, e., holyoak, m., kadmon, r., saltz, d. et al., 2008, ‘a movement ecology paradigm for unifying organismal movement research’, proceedings of the national academy of sciences 105(49), 19052–19059. pmid: 19060196, http://dx.doi.org/10.1073/pnas.0800375105 o’connell-rodwell, c.e., rodwell, t., rice, m. & hart, l.a., 2000, ‘living with the modern conservation paradigm: can agricultural communities co-exist with elephants? a five-year case study in east caprivi, namibia’, biological conservation 93(3), 381–391. http://dx.doi.org/10.1016/s0006-3207(99)00108-1 osborn, f.v., 2004, ‘seasonal variation of feeding patterns and food selection by crop-raiding elephants in zimbabwe’, african journal of ecology 42(4), 322–327. http://dx.doi.org/10.1111/j.1365-2028.2004.00531.x osborn, f.v. & rasmussen, l.e.l., 1995, ‘evidence for the effectiveness of an oleo-resin capsicum aerosol as a repellent against wild elephants in zimbabwe’, pachyderm 20, 55–64. rasmussen, h.b., wittemyer, g. & douglas-hamilton, i., 2005, ‘estimating age of immobilized elephants from teeth impressions using dental silicon’, african journal of ecology 43(3), 215–219. http://dx.doi.org/10.1111/j.1365-2028.2005.00571.x ruckstuhl, k.e. & neuhaus, p., 2002, ‘sexual segregation in ungulates: a comparative test of three hypotheses’, biological reviews of the cambridge philosophical society 77(1), 77–96. pmid: 11911375, http://dx.doi.org/10.1017/s1464793101005814 sanparks, 2011, ‘standard operating procedures for the capture, transportation and maintenance in holding facilities of wildlife: elephant’, report, veterinary wildlife services, sanparks. sanparks, 2012, ‘elephant management plan, kruger national park 2013–2022’, report, sanparks. shannon, g., page, b.r., duffy, k.j. & slotow, r., 2006, ‘the role of foraging behaviour in the sexual segregation of the african elephant’, oecologia 150(2), 344–354. pmid: 16927101, http://dx.doi.org/10.1007/s00442-006-0521-1 shannon, g., page, b.r., duffy, k.j. & slotow, r., 2010, ‘the ranging behaviour of a large sexually dimorphic herbivore in response to seasonal and annual environmental variation’, austral ecology 35(7), 731–742. http://dx.doi.org/10.1111/j.1442-9993.2009.02080.x sitati, n.w., walpole, m.j., smith, r.j. & leader-williams, n., 2003, ‘predicting spatial aspects of human–elephant conflict’, journal of applied ecology 40(4), 667–677. http://dx.doi.org/10.1046/j.1365-2664.2003.00828.x smit, i.p.j., grant, c.c. & devereux, b.j., 2007, ‘do artificial waterholes influence the way herbivores use the landscape? herbivore distribution patterns around rivers and artificial surface water sources in a large african savanna park’, biological conservation 136(1), 85–99. http://dx.doi.org/10.1016/j.biocon.2006.11.009 stokke, s. & du toit, j.t., 2000, ‘sex and size related differences in the dry season feeding patterns of elephants in chobe national park, botswana’, ecography 23(1), 70–80. http://dx.doi.org/10.1111/j.1600-0587.2000.tb00262.x sukumar, r. & gadgil, m., 1988, ‘male-female differences in foraging on crops by asian elephants’, animal behaviour 36(4), 1233–1235. http://dx.doi.org/10.1016/s0003-3472(88)80084-8 thomas, b., holland, j.d. & minot, e.o., 2011, ‘seasonal home ranges of elephants (loxodonta africana) and their movements between sabi sand reserve and kruger national park’, african journal of ecology 50(2), 131–139. http://dx.doi.org/10.1111/j.1365-2028.2011.01300.x van de perre, f., adriaensen, f., songorwa, a.n. & leirs, h., 2014, ‘locating elephant corridors between saadani national park and the wami-mbiki wildlife management area, tanzania’, african journal of ecology 52(4), 448–457. http://dx.doi.org/10.1111/aje.12139 venter, f.j. & gertenbach, w.p.d., 1986, ‘a cursory review of the climate and vegetation of the kruger national park’, koedoe 29(1), 139–148. http://dx.doi.org/10.4102/koedoe.v29i1.526 venter, f.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation pattern’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83–129, island press, washington dc. wall, j., wittemyer, g., klinkenberg, b. & douglas-hamilton, i., 2014, ‘novel opportunities for wildlife conservation and research with real-time monitoring’, ecological applications 24(4), 593–601. pmid: 24988762, http://dx.doi.org/10.1890/13-1971.1 whyte, i.j., 2001, ‘conservation management of the kruger national park elephant population’, phd thesis, department of zoology and entomology, university of pretoria. wing, l.d. & buss, i.o., 1970, ‘elephants and forests’, wildlife monographs 19, 3–92. witter, r., 2013, ‘elephant-induced displacement and the power of choice: moral narrative about resettlement in mozambique's limpopo national park’, conservation and society 11(4), 406–419. http://dx.doi.org/10.4103/0972-4923.125756 reviewer acknowledgement http://www.koedoe.co.za open access koedoe recognises and acknowledges the value and importance of reviewers in the publication process – not only in shaping the individual manuscript, but also in upholding the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely, productive peer review process. we would like to take this opportunity to thank all reviewers who participated in shaping this volume of koedoe: page 1 of 1 in an effort to facilitate the selection of appropriate peer reviewers for koedoe, we ask that you take a moment to update your electronic portfolio at www.koedoe.co.za for our records. this will allow us better access to your areas of interest and expertise, in order to match reviewers with submitted manuscripts. if you would like to become a reviewer, please visit the journal website and register as a reviewer. to access your details on the website, you will need to follow these steps: 1. log into the online journal at http://www. koedoe.co.za 2. in your ‘user home’ [http://www.koedoe. co.za/index.php/koedoe/ user] select ‘edit my profile’ under the heading ‘my account’ and insert all relevant details, bio statement and reviewing interest. 3. it is good practice as a reviewer to update your personal details regularly, to ensure contact with you throughout your professional term as reviewer to koedoe. please do not hesitate to contact me if you require assistance in performing this task. margo martens submissions@koedoe.co.za tel: +27 21 975 2602 fax: +27 21 975 4635 koedoe african protected area conservation and science anneli douglas antoinette p. malan brian kuhn bruce h. brockett carla staver cleo gosling clinton carbutt dave balfour david le maitre edwin muchapondwa gabriela bucini george bredenkamp ian a. russell jens oldeland jesse m. kalwij johan van tol john o’brien joop schaminée juergen dengler keenan stears laura k. blamey lucas enrico matthew mcconnachie megan griffiths navashni govender nicola stevens norbert hahn olaf weyl osborn v. ferrel pieter bester richard stirzaker rina c. grant robert buitenwerf robert guldemond sam m. ferreira shaun levick tammy robinson tercia strydom theo h.c. mostert ute schmiedel vaughan spaull yolanda pretorius 94 article information authors: harry biggs1 charles breen2 rob slotow3 stefanie freitag1 marc hockings4 affiliations: 1south african national parks, skukuza, south africa 2centre for environment, agriculture and development, university of kwazulu-natal, south africa 3school of biological and conservation sciences, university of kwazulu-natal, south africa 4school of geography, planning and environmental management, university of queensland, australia correspondence to: harry biggs email: biggs@sanparks.org postal address: private bag x402, skukuza 1350, south africa dates: received: 10 may 2010 accepted: 17 dec. 2010 published: 13 may 2011 how to cite this article: biggs, h., breen, c., slotow, r., freitag, s. & hockings, m., 2011, ‘how assessment and reflection relate to more effective learning in adaptive management’, koedoe 53(2), art. #1001, 13 pages. doi:10.4102/koedoe.v53i2.1001 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) how assessment and reflection relate to more effective learning in adaptive management in this essay... open access • abstract • introduction • methods • initial analysis of how adaptive management has been assessed and reflected upon • moving towards improved learning • assessment, reflection and the learning dualities • discussion of the learning duality approach as applied to the case studies • is the kruger national park good at learning in adaptive ecosystem management? • conclusion • acknowledgements • references abstract (back to top) assessment (an immediate evaluation of significance or performance) and reflection (a lengthy, deep consideration) should be important components of adaptive management leading to learning. in this paper we use a prototype adaptive cycle and feedback framework, which are related to some aspects of learning theory, to examine the extent to which assessment and reflection were applied in a series of studies and initiatives in the kruger national park. in addition to evaluating assessment and reflection, we also considered how the various contributing components of each case were inter-related to provide a holistic view of each initiative. two other studies in the kruger national park, which have examined learning specifically, are also discussed. one of them suggests that in a complex environment, learning necessarily has a dual nature, with each component of seven contrasting pairs of the aspects of learning in partial tension with the other. we use these dualities to further probe assessment, reflection, inter-relatedness and learning in the cases presented. each contrasting aspect of a ‘learning duality’ turns out to emphasise either assessment or reflection, which reinforces the idea that both are needed to facilitate sufficient learning for successful adaptive management. we hope this analysis can act as a springboard for further study, practice and reflection on these important and often underrated components of adaptive management. conservation implications: the better understanding of assessment and reflection as being largely separate but complementary actions will assist adaptive management practitioners to give explicit attention to both, and to relate them better to each other. introduction (back to top) south african national parks (sanparks) formally adopted an adaptive management paradigm (walters 1986) in the form of strategic adaptive management (sam) (roux & foxcroft 2011) in the kruger national park (knp) in the 1990s. the park has become widely known for this initiative, which has now also spread to other protected areas and other natural resource management situations (freitag, biggs & breen, in review). it is therefore fitting to determine the extent to which adaptive management performance in the knp has been assessed and/or reflected upon, and how this has contributed to further learning in a management context that typically embraces spatial and temporal variation and socioecological linkages. this paper examines whether adaptive management, designed for decision making in uncertain environments, is being implemented in a rote, mechanistic way or supported with effective assessment, reflection and learning. evidence-based policy and decision making processes and the theory of learning are used in a wide range of fields (e.g. healthcare, see gray [1997]). it is reasonable and more generic to see the adaptive management processes discussed in this paper as a particular (albeit more unpacked and iterative) form of the construct called the ‘policy cycle’, as exemplified by the well-cited australian policy cycle (bridgman & davis 2004). although the latter has been criticised for simplicity and linearity, broad parallels with several of the sequential steps of adaptive management emerge (roux & foxcroft 2011). it is likely that each has drawn on the other’s underlying thought processes and may continue to do so. from an examination of fundamental references on learning (e.g. kolb 1984; schön 1983) it is clear that such processes are linked to learning; indeed, individuals and organisations adapt (and therefore often survive or prosper) based on re-conceptualisation following reflective observation of past experiences. it is therefore appropriate that this paper frames learning as the central goal of assessment and reflection in adaptive management. after examining assessment and reflection specifically, we draw directly on other learning work (the interviews for which were conducted on knp staff) to examine the relationship between six learning dualities (roux, murray & van wyk 2008) and assessment and reflection, respectively. assessment generally means to ‘evaluate or estimate the nature, quality, ability, extent, or significance of’ whilst reflection is viewed as ‘a calm, lengthy, intent consideration’ (wordnetweb.princeton.edu/perl/webwn). although the words ‘assessment’ and ‘reflection’ are sometimes used interchangeably in the conservation literature, we shall use the definitions as described above (recognising limited overlap), except when citing. in the context of adaptive management, assessment is commonly intuitively thought to lead to reflection. such reflective behaviour should specifically be carried out in the full context of the system under consideration, observing and modifying its own attributes and behaviour, with a view to learn for future action. an organisation that practices such reflective behaviour can be considered an adaptive learning organisation (garvin 1993) in which creating opportunities for reflection through a process of consistent and structured review of past performance can be a critical factor. argouridas and race (2007) have argued for the importance of evidence-based reflection in learning and knowledge management within higher education. however, too narrow a focus on (often limited) evidence may also hide the controlling influence of context. such reflective aspects are widely underemphasised, often as a result of an incentive to progress with implementation and so be seen as ‘doing something’ (allan & curtis 2005) or because the organisational culture does not promote learning and adaptation (fazey & schultz 2009). it is therefore important to consider the prominence given to reflection in the broader learning literature (schön 1983). literature regarding effective management of protected areas describe assessment and evaluation cues in a sequential, iterative and adaptive process at different points in the management life cycle (figure 1), which is itself a version of an adaptive cycle (described as the adaptive review cycle of sam by rogers [2005]). hockings, stolton and dudley (2004) emphasise the need to assess and evaluate continuously during the cycle, rather than only after the outcomes step (figure 1). we specifically avoid the term ‘evaluation’ onwards, except where explicitly used by others (e.g. hockings et al. 2004). we regard ‘evaluation’ to be closer in meaning to ‘assessment’, although it can include the important additional aspect of passing judgement, which suggests some reflection, according to our definition. experience and learning provide improved understanding of the complexity and changing context, which, in turn, allows for refinement or revision of different steps in the cycle. by following all six evaluation cues (the straight solid arrows radiating from the centre) and considering them in relation to one another participants can view the overall process holistically, for example, as applies to park management (hockings et al. 2009). also, it is regarded to be diagnostically useful to ‘step back through the process’ so that each component can be considered with regard to its preceding input. if the actions represented by feedback arrows in figure 1 are ignored, assessment and/or reflection will not occur in appropriate contexts, and hence opportunities for meaningful and holistic learning will be missed. if assessment and reflection are considered to be fundamentally distinct processes, lumping them together may lead to practitioners subsequently acting without one or the other consciously having taken place. this may lead to poorly considered action in a complex, changing setting such as management of a park or natural resources and negate learning opportunities or effectiveness. another characteristic depiction of sam is a series of well-defined nested feedbacks as shown in figure 2a. (‘nested’ means feedback is contained conceptually within another feedback, in a logical sequence.) such nested feedbacks are usually shown most usefully as arising from the postoperationalisation learning opportunity of the cycle (i.e. once things on the ground start happening), but they could be drawn from any step in the cycle (as shown in figure 2b). indeed, the recommended diagnostic approach of sequentially stepping backward, as shown in figure 1, can be considered nested. in the context of the process steps of sam (rogers 2005) there are several opportunities for ‘advance reflection’ or ‘stepping forward’ in the early forward planning components, specifically when predicting consequences of potential management options or the societal acceptability of these, before the specific management action is selected. such nesting for attempted prediction or scenario planning occurs through a process of ‘thought experimentation’ (figure 2b) or even explicit modelling (starfield & bleloch 1991). figure 2 contains a key decision point (‘select options’) and although other decisions are taken at other points, this decision point gives rise to the most direct ‘action on the ground’. an analogous point can be inferred between ‘process’ and ‘outputs’ in figure 1. a combination of figures 1 and 2 hence clarifies opportunities to facilitate structured forms of both assessment and reflection. nevertheless, we need not be bound by only these pathways, as additional unstructured assessment (i.e. without any obvious direct reference to particular heuristics depicted in figures 1 and 2) and particularly unstructured reflection, may add further value. checking that each nested step is being performed constitutes one way of verifying that adaptive management is indeed being practiced, or that it is at least procedurally sound. yet the context in which different situations are assessed or reflected upon varies a great deal and usually changes over time, which makes iterations, including explicit reflection on the changing context itself, essential. furthermore, the subcontexts (e.g. fire management, surface water source management, etc.) that constitute any particular context (e.g. the management of the park as a whole) change differentially and it is the aim of management actions to alter some of these subcontexts. as the context becomes more confined, management influence increases. however, as the context grows and becomes more related to governance, management influence usually decreases (nkhata & breen 2010). the appropriate scale(s) and/or organisational levels at which an assessment or reflection initiative should focus should thus be considered. levels often correspond to the different levels of park objectives. as such, scale of selection may determine whether different contexts or subcontexts are appropriately and fairly assessed and reflected upon and, hence, whether these have been wisely chosen as a basis for thinking about management action. as one proceeds along the cycle in figure 1 (from planning onwards), the scale of reflection usually increases in both space and time. there is a move from specific objectives, as defined through planning and implementation, to more general goals as the process approaches the scale of re-visioning. accordingly, there is a move away from considering only some parts at particular scales towards a more holistic, integrated and systems-based consideration. if processes are institutionally embedded and aligned with the context, within an enabling organisational culture (see growcock, sutherland & stathis 2009; stirzaker, roux & biggs 2011), adaptive management can be improved. on the other hand, organisations typically have several procedures and approaches, many of which are not adaptive in nature (and are even designed not to be), and it can be challenging to dovetail these with adaptive assessment and reflection. in addition to the adaptive processes outlined earlier (and depicted graphically in figures 1 and 2), this paragraph provides a brief history of adaptive management in knp as a backdrop for understanding where assessment and reflection, as described in this paper, fit in. mabunda, pienaar and verhoef (2003) described, before the advent of explicit adaptive management in knp in the 1990s, a series of different management eras, each with a particular style. these eras set favourable precursor conditions (such as regular interaction between scientists and managers and joint responsibility) for explicit adaptive management to be accepted once the opportunity arose (see biggs & rogers 2003, box 1). freitag et al. (in review) describe how adaptive management practice, after expanding from the knp rivers research programme to the general management of the park, spread to influence other aspects of conservation management in south africa during the first decade of this century. examples include drafting guidelines for protected areas and catchment management and planning conservation initiatives, which co-evolved with evaluations of park effectiveness (belokurov et al. 2009) and reporting of the state of biodiversity (knight 2008), both of which had a culture of assessment. figure 1: framework for assessing effectiveness of management of protected areas (adapted from hockings, m., stolton, s. & dudley, n., 2004, ‘management effectiveness: assessing management of protected areas’, journal of environmental policy and planning 6(2), 157-174). figure 2: schematic representation of nested adaptive management feedback loops as (a) described by pollard, s.r. & du toit, d.r., 2007, guidelines for strategic adaptive management. experiences from managing the rivers of the kruger national park. guidelines for management and field staff of protected areas, ecosystems, protected areas and people (epp) project (unep gef no. gf/27-13-03-4679), regional protected areas programme of iucn–world conservation union in asia; and (b) adapted from the same source. methods (back to top) we reviewed as many earlier initiatives that had focused on the operation or performance of adaptive management in the knp as possible. these were mostly in qualitative or narrative form, although some had semiquantitative ‘scores’. some of the initiatives were internal (part of the prescribed procedures in the adaptive management protocols), whilst others had been subjected to previous external review (as one would expect in an independent audit). however, this distinction actually translates to a continuum spanning ‘designed as internal’ to ‘initially mainly external’, because (1) feedbacks are intrinsic to adaptive management and contribute to defining a wider system boundary and (2) implementation of the initiatives often involved the same people who initially advised on the operation of the system, possibly owing to their existing knowledge. these initiatives were subjectively rated for their emphasis on assessment, reflection and inter-relatedness, independent of impression of the study quality (table 1). inter-relatedness pertains to the holistic approach of the initiative and considered how the different levels of adaptive feedbacks are related and considered jointly, given that they can be viewed as nested within one another. we regarded inter-relatedness to enhance assessment and reflection, thereby facilitating more effective learning. we scored the respective columns in table 1 based on our impressions of the extent to which each initiative dealt with assessment, reflection and inter-relatedness. we did not consider the interpretations of how these aspects contributed to the study system, but simply whether the aspects could have been noticed and discussed. cross-correlation results we report in table 2 may therefore partly reflect differential study designs of the various initiatives. however, our interim conclusions of this initial analysis include conclusions based on actual findings of particular studies (as in a literature review), shown as numbered references in table 1. following this initial analysis, which serves as a potential bridge to the issue of learning, we describe two further studies that dealt explicitly with learning in the knp in or close to the context of ecosystem management. one of these studies produced a set of dualities, which we thought could support our own exploration into assessment and reflection. the third leg of the method involved mapping examples of assessment and reflection in adaptive ecosystem management in the knp onto these paired strategy options (roux et al. 2008) or dualities (table 3) and then exploring how assessment and reflection appeared to be related to each other and to learning, respectively (table 4). the dualities are presented as sharply dichotomous (binary), whereas they can, in reality form more of a continuum in some of the cases (e.g. learning cf. unlearning). table 1: studies or initiatives that involved adaptive management in or around the kruger national park, rated for their degree of assessment, reflection and interrelatedness of the particular analysis. table 2: spearman’s rank correlations between ten ordered individual scores for assessment, reflection and inter-relatedness in table 1. table 3: the two facets of each learning duality as aligned with assessment and reflection. each case is related to adaptive processes through pointing to phases in figure 1 (context, planning, inputs, process, outputs, and outcomes) and process linkages in figure 2. (the phrase ‘bottom loops’ refers to loops marked 1–3 and ‘upper loops’ to those marked 4–7 in figure 2a). all themes used here are discussed in detail in one or more of the references given in table 1. table 4: the relationship between assessment and reflection for each paired case in table 3, with comments on overall learning and the relationship to adaptive management. initial analysis of how adaptive management has been assessed and reflected upon (back to top) the knp has explicitly attempted to implement strategic adaptive management, at least in the sphere of ecosystem management, for 15 years, and the methods and intentions have been documented in publications, reports and brochures (e.g. biggs & rogers 2003; iucn freshwater task force and skukuza group [in prep.]; pollard & du toit 2005; sanparks 2008). this has attracted the attention of scholars and managers, with several initiatives or studies aimed at meaningfully looking directly or indirectly at this drive (table 1). the studies and initiatives listed in table 1 provide a useful idea of how well (1) the defined adaptive management processes, and/or (2) actual management outcomes (the ultimate goals of sam) have performed. these are invariably presented in narrative form in the originals, although some do include semiquantitative scores. our general summary after reviewing the findings of these studies follows below. numbers in square brackets refer to the studies listed in table 1 that most clearly support each point: • adaptive planning has progressed and produced a more shared rationale (i.e. more overlap in underlying mental models) amongst most key stakeholder groups, seemingly greater than we can perceive prior to this from documented history (over 100 years). [3, 5, 6, 10] • explicit objectives seem to be well accepted by relevant parties, which results in their commitment. however, prioritisation of objectives remains problematic (comparative risk assessment [box 1, gaylard & ferreira 2011] is currently being tested for prioritisation). [1, 5, 7] • explicit feedbacks now exist at various scales. for example, feedback from monitoring thresholds of potential concern (biggs & rogers 2003) are nested within park-wide state of biodiversity reports. [2, 8, 9] • implementing adaptive processes took time but they are now fairly widespread (but not universally used) in the knp. these processes spread from one theme (rivers) to several others within knp, and then more widely to park planning and river management domains in south africa. [4, 5, 6] • general implementation of adaptive management itself remains variable and somewhat messy, with some distinct ‘pockets of hope’ appearing between barriers that still need to be overcome. these impressions were gleaned from the more recent studies reviewed, which have had the benefit of longer hindsight than is probably the norm. the extent of progress is thought to relate to personal attitudes and institutional issues rather than lack of biophysical understanding. [5, 6, 8] • a better grasp of social-ecological interactions in these systems is still needed, including threshold behaviour. [5, 6, 8] • social capital amongst actors and within networks and enthusiasm for action are critical underlying success factors. [6, 10] table 1 also aimed to examine assessment, reflection and inter-relatedness within each initiative. our generating scores across all the examples provides a subjective and relative idea of the amount of assessment, reflection and inter-relating in each. with ten pairs for comparison (8 d.f.), none of the correlations between columns proved significant (table 2). the relationships are nevertheless informative, with deeper assessment and reflection apparently interfering with rather than potentiating each other; deeper reflection and inter-relation of feedbacks appear positively associated. any possible significant cross-correlations which may arise with the use of greater sample sizes in future, may also reflect differential study design of the various initiatives. this could be considered equally informative, in the sense that the overall system of study and action is in some cases less geared to picking up on or producing, say, actual reflection. moving towards improved learning (back to top) our fundamental premise is that assessment, reflection and their thorough inter-relation create a foundation of shared understanding (amongst all the relevant stakeholders associated with a national park) that directs and shapes deliberations and creates the basis for adaptive learning. how can we then assess whether this learning, in the context of adaptive management, takes place effectively? a learning organisation (garvin 1993) is one skilled at creating, acquiring, and transferring knowledge and is able to modify its behaviour to reflect new knowledge and insights. schön (1983) provides a theoretical understanding of learning and michael (1995) specifically lists criteria to assess learning organisations, illustrated by questions such as whether (1) error is accepted, and fear of error is offset, in participating individuals, (2) group or social learning takes place, with facilitation rather than chairing of meetings, and (3) crises are leveraged as opportunities for learning. these kinds of question are mirrored by broadly similar ones for benchmarking adaptive management (stankey, clark & bormann 2005), determining effective environmental administration (paehlke & torgerson 1990), assessing resilience and sustainability of socioecological systems (walker & salt 2006) and managing businesses sufficiently ‘mindfully’ to sense emergent signals and their meaning in a currently uncertain and complex world (weick & sutcliffe 2001). it would thus appear that these fields are all strongly influenced by the way learning is enabled in individuals and across organisations and by whether or not behaviours have been modified in response to growth in understanding as per the reflection stage of the experiential learning cycle (kolb 1984). a recent consultation delivered to sanparks scientific services staff (narrative lab 2009) deals with factors enabling performance in the sanparks work environment, which staff consider to be complex and dynamic, after having worked through responses to questions of a similar genre to these above. key recommendations, such as more regular use of probes (small, adaptive experiments, mainly in the organisational milieu), to help structure learning emerged. knp staff were interviewed in two studies to probe their learning experiences after the inception of adaptive management. as part of an iucn study, pollard and du toit (2005, 2007) interviewed rangers whose sections included perennial rivers to establish their attitudes towards the adaptive management of rivers. the results showed good, but not universal, uptake of river objectives amongst participants and an understanding that their own inputs guided these objectives as part of a holistic vision (s. pollard, pers. comm.). rangers were aware of, and to an extent also participating in influencing, the changing context (e.g. cultural, legal, climatic and water use context). long-serving participants regarded this as an improvement compared to previous such initiatives. there was fair buy-in for the overall goal to plan and manage actively for river-related objectives and the majority of interviewees appreciated that they had been involved actively. they understood their roles in the process and, moreover, that their behaviour had changed to include more concern for and action on river issues. several mentioned other administrative systems as competing for valuable time when it came to executing the river objectives. the results of the study suggest that rangers participated in a nested way in assessments, were partly involved in reflection, and subsequently learnt in a meaningful way. roux, murray and van wyk (2009) interviewed eight sanparks scientists of knp scientific services to examine the extent to which their department could be considered a learning organisation. interviewees considered 11 learning principles derived from literature and regarded statements reflecting application of nine of these as being ‘largely true’. scores reflecting organisational capability and information and data processing were far lower, suggesting caveats in understanding and/or implementation, perhaps relative to the growth in other learning areas which placed higher demands on these. with reference to the institutional model of anderies, janssen and ostrom (2004), roux et al. (2009:49) found that although the internal learning system (in this specific context referring mainly to within scientific services in the knp) works fairly well, the greater encompassing learning systems (e.g. catchment management agencies) do not necessarily do so. links from the subsystem to these bigger systems may also have been unclear: ‘good knowledge workers [within knp] must step outside of their usual domain to engage issues within the larger learning system and to deal with complexity and response scenarios across a range of organisations and scales’ (roux et al. 2009:49). however, organisational context and ethics sometimes complicate moving beyond the usual domain and functioning effectively, even for those who are willing and keen to do so. this study targeted learning, and although reflection was explicitly probed and found to be taking place to some extent within the organisation, the report did not deal directly with assessment and inter-relatedness as discrete or traceable topics. self-evaluation within the mentioned project suggested limitations of such mechanistic analyses. this led to a more nuanced articulation in which each action was seen as a paradox, of which facets of both components needed simultaneous consideration, similar to the seven paired strategy options described by roux et al. (2008) and which we describe here as learning dualities. they include: • considering both prior knowledge and new evidence • practicing conceptual as well as practical learning • acquiring breadth and depth of knowledge • learning and unlearning • applying single-loop, double-loop and triple-loop learning • learning though patient reflection and rapid responses • focussing on individual and extended group learning. the maturity, purpose and operating realities of an organisation will cause the balance between these dual needs to change over time. without this tension, it is unlikely that adaptive learning can take place effectively and, as such, adaptive management itself should be expected to lead to conscious or subconscious shifts in this balance. assessment, reflection and the learning dualities (back to top) roux et al. (2008) found that these paired dualities, if discussed in depth amongst stakeholders to determine how effective they were at practicing adaptive learning, were extremely useful in eliciting improved understanding of the learning process. using these dualities as a framework against which assessment and reflection (and the relationship between them) in the studies mentioned in table 1 can be compared should help us to gain knowledge of what was learnt, and how this learning relates to adaptive management. we propose that the opposite endpoints of each duality appear to represent an emphasis towards either assessment or reflection (table 3) as defined in this paper. we have intentionally selected examples (from sources in table 1) to illustrate this from a wide range of themes. we use one theme for each duality so that assessment and reflection for the specific duality can be juxtaposed. although assessment is commonly thought of as a precondition for reflection, we try to examine the relationship between the two critically in the narration in table 4. we also comment on how the authors gauged overall learning to have proceeded and the inferred links to adaptive management. this helps cover the full organisational perspective of the cycles represented in figures 1 and 2, including assessment, reflection and integration, and their influence on and from decision making. in this way we hope to illustrate whether, through decision making, learning leads to improved action, and action to further learning, as described by kolb (1984). from tables 3 and 4 it becomes clear that cycles of both assessment and reflection are asserted to have led to learning, albeit at different scales and paces and with varying time frames for decision making and management implementation. effective adaptive management, however, does require a timely response to learning. this has not always been the case in the knp examples, and assessment, reflection, and response times have often been decoupled. this led to reduced effectiveness, which is probably related to either lagged learning or lagged responses to learning. however, not all lags are undesirable; in fact, they can allow for adequate reflection (cilliers 2006). discussion of the learning duality approach as applied to the case studies (back to top) the examples illustrating the learning dualities highlight circumstances which emphasised assessment or reflection, how they appeared to inter-relate, and allude to some obvious gaps (tables 3 and 4). this strongly suggests that it is meaningful to distinguish between assessment and reflection, because they appear to have different roles and can lead to different outcomes and are apparently often decoupled. inter-relatedness of feedbacks appears to augment and/or be the result of deeper reflection, although such evidence is limited at present. in addition, the intuitive impression that assessment precedes reflection seems not to be well supported by these narratives. we believe that in order to span the various needs of the complex challenges in natural resource management, both assessment and reflection are needed, perhaps concurrently. intuitively, we seem to want them to be complementary or even synergistic, although our case material suggests they do not often tend to feature‘naturally’ in the same initiative. the relationship between them is often complex (tables 3 and 4) and it may require explicit effort to create any such synergy or complementarity. it seems that assessment is often linked to acting on the answers that emerge, whilst reflection may generate more depth by asking more questions. the challenges we face may need both action and concurrent contemplation if we are to stay both responsive and resilient in future. is it possible that reflection is meant more to consider our actions and interventions, whilst assessment is meant to trigger more immediate interventions? assessment and reflection can be considered to constitute an overarching duality required for effective learning and, where they lead to decision making, also then for adaptive management. kolb’s (1984) learning cycle refers to observations and reflections following from experiences, with reflections then being distilled into abstract concepts for re-testing in reality. he details different learning styles, of which a rework by honey and mumford (as cited in businessballs [n.d.]), describes activists, reflectors, theorists and pragmatists. we suggest that assessment may be something carried out more by activists and pragmatists, whilst reflection is an activity more related to reflectors and theorists. importantly, these four styles together constitute a full learning cycle of kolb (1984), possibly implying that what we interpret as a duality between assessment and reflection could also (and perhaps better) be expressed as assessment and reflection complementing each other in a dovetailed way. this might also explain why we interpret the preliminary data (table 2) as suggesting a disjunct between assessment and reflection. one likely explanation is that if they are not executed together in the right dovetailed way – that is, if there is an imbalance between them (as there often seems to be in our case studies) – learning is in some sense incomplete. we accept that under the circumstances that unfold in the situations we normally deal with there may be a need for later studies to try to re-align the correct balance between them, and hope that this paper may have contributed in this way. conversely, we suggest that if one can design complementary use of assessment and reflection appropriately from the outset, learning may be more effective. as with the learning dualities, or in the same way that learning styles (kolb 1984) are seen as mutually exclusive at one instant, there is a necessary tension between assessment and reflection (roux et al. 2009). presumably, shifts in emphasis between them over time are, or should be, adaptive. a generalised schematic of the inter-relationship of key elements of assessment and reflection, supported by our analysis, is presented in figure 3. whilst this represents the existing tensions, striving for an improved relationship should also include issues of scaling. with some exceptions, reflection conventionally tends to involve coarse-scale issues and assessment finer-scale issues, as seen in our examples. challenging this ‘convention of scale’ by also reflecting at finer scales and assessing at coarser scales, and interlinking assessment and reflection across these scales, could lead to more useful feedbacks. if better understood and linked, a well-conceptualised and tailored combination of assessment and reflection (i.e. for the particular situation at a particular time) may support enhanced purposeful learning. perhaps we have not made the differences between them sufficiently explicit to be able to exploit the opportunities of this fuller learning. this paper illustrates that they do not have a fixed, linear or unidirectional relationship. even when used independently, but particularly when used together, they appear to underpin learning and, when they result in decision making, also adaptive management. emphasis on one or the other leads to different types of learning (duality 5, table 3) and evoke different adaptive management feedback loops (highlighting loops 1–3 and 5 for assessment, and loops 4, 6 and 7 for reflection; figure 2a). so, there appears to be a need for both to complete the adaptive management cycle, similar to how different learning styles produce a full kolb (1984) learning cycle. our impression is that assessment tends to provide, broadly, the basis for the inputs through output stages, and reflection the basis for the outcome through inputs stages (figure 1). this may be a consequence of managerial and scientific groups in the knp being more involved in the implementation and planning phases, respectively, although older staff recall an era (of the so-called ‘house 31’ meetings and the first adaptively planned management plan in the late 1990s) when managers and scientists appeared to share assessment and reflection more equitably than after the staff restructuring of 2001. our challenge appears to be, for each case, to couple assessment and reflection more productively by ensuring enough of each (there is often too little reflection) at scales appropriate to the case, and ensuring that they complement each other in the most useful way. the outcome stage (figure 1) is a particularly critical step at which considerable reflection is possible. this leads to consequences for implementation, which may, through re-visioning, influence planning in the next iteration. examples where the feedback loop is thought to have been completed, such as rivers and fire management in the knp, share a key bridge that allows for learning from doing. reflection at implementation and output stages, shown in elephant management and tpc examples, is also helpful but more to refine technical aspects of management. in addition, thought experiments (depicted in figure 2b) and modelling can help identify and improve options and enhance the possibility of successfully learning from anticipated doing. in particular, thought experiments that involve both scientific and managerial stakeholders and include elements of both planning and implementation, may assist in coupling assessment and reflection at multiple stages and scales of the cycle. this should lead to more purposeful reflection and learning. decoupling of operational and conceptual issues highlights a known tension in natural resource management. this is illustrated in the threatened species example (tables 3 and 4), where specific knowledge and a lobby for a particular aspect, in this case an individual species, oppose the need to conceptualise ecological frameworks into new management paradigms, which in this case refers to management for overall ecosystem heterogeneity. assessment at the level of managing an individual iconic species was translated into objectives that were not necessarily in line with the overall park vision. no persistent holistic reflection emanated from either the outcomes phase or the emerging changes in context such as climate change. the reflection that did take place, on the other hand, resulted in an abstract conceptualisation of heterogeneity, which was unacceptable or disconnected from on-the-ground-realities in that it appeared to undermine preservation goals for charasmatic species. the examples show that when multidisciplinary, multi-institutional teams come together to reflect in the outcomes stage (in the fire and rivers management programmes and also to some extent in the elephant and alien species invasion programmes) and fully connect the adaptive management cycle though decision making, both learning and adaptive management are deemed more successful. however, these should be accompanied by sufficient assessment. such perceived success is partly lacking in the current relatively inert threatened species programme and the ‘stop-go’ artificial water provision programmes, although impressive individual initiatives may have contributed a great deal (e.g. sanparks 2010). adaptive learning seems to have been deeper and easier when a clear and jointly agreed goal was set, without other conflicting goals, which enabled decision making and implementation. this has not been the case for the threatened species programme in the knp. neither was it the case for the artificial water provisioning programme until fairly recently, when a re-evaluation of context and principles (including acknowledged overriding importance of the heterogeneity paradigm and of climate change) led to consensus for implementation (sanparks 2010). figure 3: general schematic of the existing relationships between learning, adaptive management, context specificity and the duality between assessment and reflection, as evident from analyses in this paper and learning theory in terms of the honey and mumford modification of the kolb learning cycle concept (businessballs n.d.). is the kruger national park good at learning in adaptive ecosystem management? (back to top) whilst it is encouraging to note that many narratives in this paper suggest this to be the case, there appear to be major gaps and opportunities. the evidence presented suggests that assessment-type learning predominates and is inadequately linked to reflection. assessment appears amenable to, and can even be effectively driven by, codification, which enables some stability for decision making. this may be appropriate under conditions of relatively unchanging, or assumed unchanging, contexts, but could result in ‘blind-siding’ and a ‘dulling of the senses’ when confronted with changing and newly emerging contexts. reflection, on the other hand, is often spontaneous or emergent, although, clearly, some structured processes can also be enabling and can lead to more fundamental changes in decision making. the knp may be guilty of often falling back into a narrowly defined version of adaptive management, which may tend to assume a given context. the knp may even have become trapped in a fixed context, which has become inappropriate over time. this is akin to the way in which attitudes around elephant management (eds. scholes & mennel 2008) persisted for decades before a change in thinking was accepted, all the while showing welcome technical improvements and refinements within the particular culling context, which proved unsustainable, as ‘the only answer’. on the other hand, there is generally also an urgent need for effective assessment processes, as depicted on the left side of figure 3. the way forward appears to be, as the dualities suggest, that assessment and reflection should coexist with their complementary function playing out at various scales and with varying strengths. recognition of dynamic complexity, linked to a system that supports policy-oriented decisions, seems crucial. such interlinking requires diverse teams working together on problems holistically. for example, in sanparks this could include, in a stereotypic or conventional interpretation, (1) managers who often tend to focus on finer-scale implementation and assessment issues, (2) scientists who tend to have a broad, theoretical view, focus on planning and outcomes and, intuitively, like to reflect, and (3) ‘bridging’ regional ecologists who have an understanding of both planning and implementation components. it has been stressed before that more shared functionality across these cadres results in better integration. the prescription should therefore also include stretching, within the reasonable capability of people, these stereotypes outside of their immediate operational comfort zones and promoting engagement by at least some individual team members across all aspects of the adaptive management process as illustrated in figure 1. this, in itself, will lead to enhanced achievement of the original dualities or paired options (roux et al. 2008). given the paucity of quantitative empirical evidence (cook, hockings & carter 2009), is it the reality of complex systems that we should expect a mix of fact-based and expert-based evidence and changing ideology to drive management, rather than imagining the trajectory to develop solely on hard factual evidence? how should scientists respond to this? this paper describes that, in reality, several different classes of feedback exist, of which only some are based on hard evidence. this suggests that a mix of these change drivers is inevitable (even if hard evidence plays an increasing role) and that reflection at different scales may help improve decision making and learning amidst this uncertainty, with changing ideology often (but not exclusively) manifesting itself through policy option choice. our case histories showed some examples of assessment patterns being driven by decisions made subsequent to reflection, often after a lag period, which may not always be undesirable (cilliers 2006). these reflections appear, in turn, to have been driven mainly by changing paradigms. the changing paradigm itself is a consequence of primarily, but not entirely, scientists’ reflection and these changed paradigms prompt further management reflection. we have also described how assessment contributes to further reflection, but individually these effects seem to reach triple-loop standing only occasionally (roux et al. 2009). thus, a combination of altered expert opinion (including that of managers) and changing values, in some cases together with the appearance of a certain amount of new evidence, appears to lead to change. all three these drivers may co-evolve or be brought together in a more conscious process. is the overlap of striving for the same goal, although perhaps for different reasons, a key element of practical progress in many cases? a way forward may be to target several key questions: • what influences the balance between assessment and reflection in particular cases? • is assessment more prevalent because it is easier for organisations to implement and/or codify? • what is the trade-off between tedious, detailed assessment and a more practical intervention that is not overly simplistic? • should all systems strive to include and promote personal thoughtfulness and vibrancy, along with other strategies, to prevent becoming rote? • how can we promote more reflection and its meaningful relationship with assessment? these may be addressed in an enlightened way by using one or multiple approaches to evaluate our success at adaptation and learning, as discussed by several others (e.g. michael 1995; narrative lab 2009; paehlke & torgerson 1990; stankey et al. 2005; walker & salt 2006; weick & sutcliffe 2001), and incorporating these into wider fundamental learning theory (kolb 1984). such initiatives could even re-use the case data mentioned in table 1. although they have much in common, each of these approaches also has a unique slant. we propose that sufficient reflection on one or more of these approaches (perhaps emphasising their commonalities) and not only the learning dualities on which this paper has focused, may yield startling and useful consolidations in our particular context of natural resource management. conclusion (back to top) we have shown that both mindful assessment and reflection are necessary for purposeful learning and adaptive management, and that there is much scope for improvement throughout this interacting system. consideration to both learning theory and the correct contextualisation of adaptive management helps to determine the dynamic coupling we observe in natural resource management: complex social-ecological systems are embedded in extensive dynamic contexts. with this paper we have made a start by looking at empirical evidence available after more than a decade of adaptive management in a park, and relating the patterns to meaningful theory. we hope the review can serve as a springboard for more exact work of this kind and hence further enhance learning. acknowledgements (back to top) we thank alice schwarz brunold (office of marketing and communications, university of queensland) for professional assistance with compiling figure 1, eureta rosenberg for insightful discussions after reading an earlier draft of the manuscript, and the two reviewers for their painstaking efforts to help us generate a clear, well-grounded and useful product. references (back to top) allan, c. & curtis, a., 2005, ‘nipped in the bud: why regional scale adaptive management is not blooming’, environmental management 36(3), 414−425. anderies, j.m., janssen, m.a. & ostrom, e., 2004, ‘a framework to analyze the robustness of social-ecological systems from an institutional perspective’, ecology and society 9(1), 18, viewed 28 april 2011, from http://www.ecologyandsociety.org/vol9/iss1/art18/ argouridas, v. & race, p., 2007, ‘enhancing knowledge management in design education through systematic reflection practice’, concurrent engineering: research and applications 15(1), 63−76. belokurov, a., besançon, c., burgess, n.d., dudley, n., hockings, m., leverington, f., et al., 2009, ‘new resources for assessing the effectiveness of management in protected areas’, oryx 43(1), 14. biggs, h.c. & rogers, k.m., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j. du toit, k.m. rogers and h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59−80, island press, covelo. biggs, h. & van wyk, a., 2000, ‘how far are we with our stated goals and how should we modify them along the way?’, presented at annual general meeting of conservation section, kruger national park, skukuza, 9–10 nov 2000. businessballs n.d., kolb learning styles, featuring honey and mumford’s variation, viewed 28 march 2011, from http://www.businessballs.com/kolblearningstyles.htm bridgman, p. & davis, g., 2004, the australian policy handbook, allen & unwin, sydney. cilliers, f.p., 2006, ‘on the importance of a certain slowness. stability, memory and hysteresis in complex systems’, emergence: complexity and organisation 8(3), 106−113. cook, c., hockings, m. & carter, r.w., 2009, ‘conservation in the dark: the information used to support decisions’, frontiers in ecology and the environment 8(4), 181–186. du toit, j., rogers, j.k. & biggs, h.c. (eds.), 2003, the kruger experience: ecology and management of savanna heterogeneity, island press, covelo. duff, j., 2002, ‘an assessment of adaptive management practices in south africa national parks and ezemvelo kwazulu natal wildlife’, msc dissertation, centre of environment and development, faculty of science and agriculture, university of natal. fazey, i. & schultz, l., 2009, ‘adaptive people for adaptive management’ in c. allen & g. stankey (eds.), adaptive environmental management: a practitioner’s guide, pp. 323−338, springer, dortrecht and csiro, collingwood. freitag, s., biggs, h. & breen, c., in review, ‘fifteen years of the spread and maturation of adaptive management in south african national parks: organisational learning in systems perspective’, in w. freimund, s. mccool & c.m. breen (eds.), engaging complexity in protected area management: challenging occam’s razor, university of kwazulu-natal press, pietermaritzburg. gaylard, a. & ferreira, s.m., 2011, ‘modification and maturation of sanpark’s adaptive planning process – making critical linkages between conservation objectives and actions’, koedoe 53(2), art. #1005, 8 pages. doi:10.4102/koedoe.v53i2.1005 garvin, d.a., 1993, ‘building a learning organization’. harvard business review 71(4), 78–91. gray, j.a. muir, 1997, evidence-based health care, churchill livingstone, edinburgh. growcock, a.j., sutherland, e.f. & stathis, p.t., 2009, ‘challenges and experiences in implementing a management effectiveness evaluation program in a protected area system’, australasian journal of environmental management 16, 30−38. hockings, m., cook, c., carter, r.w. & james, r., 2009, ‘accountability, reporting or management improvement? development of a state of the parks assessment system in new south wales’, environmental management 43, 1013–1025. hockings, m., stolton, s. & dudley, n., 2004, ‘management effectiveness: assessing management of protected areas’, journal of environmental policy and planning 6(2), 157−174. iucn freshwater task force and skukuza group, in prep., ‘adaptive management guidelines for effective conservation of freshwater ecosystems in and around protected areas of the world’, australian wetlands and rivers centre, sydney. kingsford, r.t., biggs, h.c. & pollard, s.r., 2011, ‘strategic adaptive management in freshwater protected areas and their rivers’, biological conservation 144, 1194–1203. knight, m.h., 2008, ‘sanparks: state of biodiversity report: comparisons of 2008 & 2007 scores’, internal sanparks document, park planning & development, port elizabeth. kolb, d., 1984, experiential learning, prentice hall, englewood cliffs, nj. mabunda, d., pienaar, d.j. & verhoef, j., 2003, ‘the kruger national park: a century of management and research’, in j. du toit, k.m. rogers and h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 3−21, island press, covelo. michael, d.n., 1995, barriers and bridges to learning in a turbulent human ecology, columbia university press, new york. narrative lab, 2009, ‘investigating appropriate performance measures. south african national parks, conservation services division. final consolidated report to sanparks’, internal report, sanparks, pretoria (available from sanparks archives). nkhata, b.a. & breen, c., 2010, ‘a framework for exploring integrated learning systems for the governance and management of public protected areas’, environmental management 45(2), 403−413. paehlke, r. & torgerson, d., 1990, managing leviathan: environmental politics and the administrative state, broadview press, peterborough. pollard, s.r. & du toit, d.r., 2005, ‘recognizing heterogeneity and variability as key characteristics of savannah systems: the use of strategic adaptive management as an approach to river management within the kruger national park, south africa’, report of unep/gef project no. gf/2713-03-4679, ecosystems, protected areas and people project. iucn, gland, viewed 28 march 2011, from http://www.parksnet.org/files/1/2/65777_documents_document_file_1267.pdf pollard, s.r. & du toit, d.r., 2007, guidelines for strategic adaptive management. experiences from managing the rivers of the kruger national park. guidelines for management and field staff of protected areas, ecosystems, protected areas and people (epp) project (unep gef no. gf/27-13-03-4679), regional protected areas programme of iucn–world conservation union in asia. rebelo, t., freitag-ronaldson, s., cheney, c. & mcgeoch, m., 2011, ‘prioritizing species of special concern in the table mountain national park for monitoring: the challenge of a species-rich, threatened ecosystem’, koedoe 53(2), art. #1019, 14 pages. doi:10.4102/koedoe.v53i2.1019 rogers, k.h., 2005, biodiversity custodianship in sanparks: a protected area management planning framework, south african national parks, pretoria. roux, d.j. & foxcroft, l.c., 2011, ‘the development and application of strategic adaptive management (sam) within south african national parks’, koedoe 53(2), art. #1049, 5 pages.doi:10.4102/koedoe.v53i2.1049 roux, d.j., murray, k. & van wyk, e., 2008, ‘learning to learn for socio-ecological resilience. balancing strategy options in public sector organizations’, in m. burns & a. weaver (eds.), exploring sustainability science: a southern african perspective, pp. 599–625, african sun media, stellenbosch. roux, d.j., murray, k. & van wyk, e., 2009, enabling effective learning in catchment management agencies: research report. report no 1689/1/09, water research commission, pretoria. sanparks, 1999–2011, conservation services management committee, minutes of [regular] meetings, sanparks archive, skukuza. sanparks, 2008, ‘a framework for developing and implementing management plans for south african national parks’, viewed 15 april 2009, from http://www.sanparks.org/parks/kruger/conservation/scientific/key_issues/plans/adaptive/default.php sanparks, 2010, kruger national park manco meeting minutes (march 2010), kruger national park archives, skukuza. scholes, r.j. & mennell, k.g. (eds.), 2008, elephant management. a scientific assessment for south africa, wits university press, johannesburg. schön, d., 1983, the reflective practitioner: how professionals think in action, temple smith, london. stankey, g.h., clark, r.n. & bormann, b.t., 2005, adaptive management of natural resources: theory, concepts, and management institutions, general technical report pnw-gtr-654, us department of agriculture, forest service, pacific northwest research station, washington dc. starfield, a.m. & bleloch, a.l., 1991, ‘building models for conservation and wildlife management’, 2nd ed., burgess international, edina. stirzaker, r., roux, d.j. & biggs, h.c., 2011, ‘learning to bridge the gap between adaptive amangement and organisational culture’, koedoe 53(2), art. #1007, 6 pages. doi:10.4102/koedoe.v53i2.1007 van wilgen, b.w. & biggs, h.c., 2011, ‘a critical assessment of adaptive ecosystem management in a large savanna protected area in south africa’, biological conservation 144, 1179–1187. walker, b. & salt, d., 2006, resilience thinking: sustaining ecosystems and people in a changing world’, island press, washington dc. walters, c., 1986, adaptive management of renewable resources, macmillan, new york. water research commission, n.d., report under project k5/1711, principal investigator: s. pollard, water research commission, pretoria. water research commission, n.d., report under project k5/1797, principal investigator: c. mcloughlin, water research commission, pretoria. weick, k.e. & sutcliffe, k.m., 2001, managing the unexpected: assuring high performance in an age of complexity, jossey-bass, san francisco. article information authors: francesca cini1 melville saayman1 affiliations: 1tourism research in economic environs and society, north-west university, potchefstroom campus, south africa correspondence to: francesca cini postal address: private bag x6001, potchefstroom 2520, south africa dates: received: 26 mar. 2013 accepted: 03 dec. 2013 published: 24 june 2014 how to cite this article: cini, f. & saayman, m., 2014, ‘which age group spends the most in a national park?’, koedoe 56(2), art. #1158, 8 pages. http://dx.doi.org/10.4102/ koedoe.v56i2.1158 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. which age group spends the most in a national park? in this original research... open access • abstract • introduction    • literature review • research method and design    • setting    • design    • procedure       • socio-demographics       • expenditure patterns       • reasons for visiting the park    • analysis • results    • respondents' profile    • reasons for park visit    • expenditure patterns by age    • segment characterisation • ethical considerations • trustworthiness • discussion    • practical implications    • recommendations    • limitations • conclusion • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ age (and its changing structure amongst the wider population) is one of the most relevant aspects required to better understand and forecast the needs, interests and associated consumption behaviours of tourists. this research used age to investigate the expenditure patterns amongst a sample of visitors to the table mountain national park (tmnp), south africa. in march 2010, visitors to the tmnp were found to differ significantly from those at other parks, as they were younger and most of them were foreigners. this study found that younger visitors (18–29 years) were higher spenders when compared to those aged 30–49 years. as parks are generally visited by older people, this study showed the economic importance of the younger market. the research also made clear implications and recommendations for park management as to how to address these findings.conservation implications: conservation is dependent on funding. one of the main sources of income is tourism and tourism related activities. this research can assist marketers and managers to target the right markets in order to be more sustainable. this research also shows the importance of environmental education at an early age in order to grow awareness and to target the right markets. introduction top ↑ the economic importance of natural protected areas is recognised at both national and international levels. since the 2003 world parks congress in durban, south africa, the view of economic development and nature conservation as opposite realities is changing. in this way, a new paradigm for protected areas was fostered: the synergy between conservation and sustainable development was forged. protected areas were seen as providers of benefits beyond boundaries – beyond their boundaries of nation-states (international union for conservation of nature [iucn] 2004b). the relationship between economic development and nature conservation became increasingly important, especially in developing countries, because conservation areas can be used as a tool for poverty alleviation (buckley 1994, 2002a; bushell & mccool 2007; bushell, staiff & eagles 2007; butler & boyd 2000; geldenhuys & saayman 2009; myburgh & saayman 1999; world commission on protected areas 1998; world tourism organization 2002; world wildlife fund 2001). practical, protected areas can provide a number of benefits to the poor. for instance, they can provide some ecosystem services including coastal protection, water purification and carbon sequestration. they also can provide some options for income by providing jobs in the tourism industry. for instance, in countries such as south africa, zimbabwe and zambia, local communities obtain income from sport hunting (child & dalal-clayton 2004; iucn 2004a). in many countries, government funding for parks is reducing in real terms (buckley 2002b; eagles, mccool & haynes 2002; mabunda & wilson 2009; saayman 2009); therefore, it is becoming increasingly important to attract markets that can generate capital that can be used for sustainable development. then, particularly for third-world countries, finance generated and secured from tourism can represent the basis for nature conservation and preservation, as well as for the eradication of economic and social inequalities. in addition, it can provide benefits such as an increase in local jobs, higher local income, higher education level of local people and equal access to quality and affordable basic services (eagles et al. 2002). as promoting the mere increase in the number of visitors to capture economic benefits might be somewhat risky in terms of the negative impacts on the natural environment (see eagles et al. 2002), a possible de-marketing strategy might be developed for targeting only those segments with the stronger propensity to spend. literature review the characteristics of consumers as demographic, behavioural (including the expenditure patterns) and psychographic are widely considered the key elements of a marketing strategy. identifying the factors linked to tourists’ motivations, preferences and behaviours is crucial to foreseeing their potential travel choices (cha, mccleary & uysal 1995; frochot & morrison 2000; kotler 1992). specifically, age (and its changing structure amongst the wider population) is one of the most relevant aspects that is considered to better understand and forecast the needs, interests and associated consumption behaviours of tourists (gibson, attle & yiannakis 1998; johns & gyimóthy 2002; jönsson & devonish 2008; moutinho 1987; oh et. al. 2004; patterson & pegg 2009; seaton 1996). in western countries, the size of the older population (55 years and older) is growing at a faster rate than the younger one and represents new target tourist markets that are healthier, affluent and have a greater desire for novelty and escape than do those of previous generations (patterson & pegg 2009; sherman & schiffman 1991). the use of age as a segmentation variable is supported by several research articles that found age to be one of the primary variables, which explained the differences in people’s travel interests preferences and behaviours. to date, several studies found age as one of the main variables in affecting even travel expenditure patterns as a type of travel behaviours (thrane & farstad 2011). for example, in a study conducted by mok and iverson (2000), age significantly characterised the different spending segments. specifically, heavy spenders were younger than 50 years old. again, jang, ismail and ham (2002) found that association amongst a sample of japanese outbound travellers. specifically, this study revealed that heavy spenders were older, had a larger party size and a longer length of stay. in addition, saayman, saayman and du plessis (2005) found differences between heavy and light spenders according to their age amongst visitors to the world cup cricket matches in south africa. specifically, heavy spenders were older than 35 years old and spent more on accommodation, restaurants and souvenirs, compared to the younger ones. in most studies, profiling the spending segments was conducted using socio-demographics such as age, level of education, annual income, place of residence and trip characteristics such as travel party size, length of stay and type of accommodation. with specific regard to the relationship between travellers’ age and their use of natural areas, for instance, a study conducted by kim, lee and klenosky (2003) amongst visitors to a national park found out that the younger group (less than 50 years) was motivated mainly by the possibility of experiencing adventure and sharing their friendship, whilst the older visitors (50 years and older) by the possibility of appreciating the natural resources and enhancing their state of health. also moscardo and green (1999) found an overall decline in activity participation associated with age, particularly for activities associated with nightlife and entertainment and outdoor, physical, nature-based activities. analysing travellers by age can also have important implications for the management of a tourist destination. types of facilities and access to infrastructures, for example, might be planned on the basis of visitors’ age. strongly associated with people’s income and expenditure patterns (down 2000), age was found as one of the main variables affecting even travel expenditure patterns (jang et al. 2004; saayman et al. 2005). however, a review of the literature revealed a dearth of studies aimed at investigating the expenditure patterns (behavioural characteristics) amongst nature-based tourists, or even of expenditure behaviours by their age. amongst the few, a study conducted by mehmetoglu (2007) investigated the relationship between the daily expenditures and the trip activities of a sample of visitors at two nature-based attractions in northern norway. heavy spenders were more likely to consider nature-based activities when compared to the light spenders who, instead, attached more importance to visiting museums and attractions. whilst motivational differences amongst the spending groups were partially supported by the study, income and age were found to significantly influence their expenditure patterns. specifically, older nature-based tourists (50 years and older) were more likely to be lighter spenders than were younger ones. another study looked at the variables that influence spending patterns and made recommendations to attract high-spending markets to the tsitsikamma national park in south africa (kruger, saayman & saayman 2010). respondents segmented by their real expenditure per person per day showed differences based on their age. two different markets were identified, (1) ‘high-spenders’, aged 35–49 years and older than 50 years, more likely to stay in chalet accommodations and to have a longer stay at the park and (2) ‘low-spenders’, aged 35–49 years old, more likely to stay in camping accommodations and have a shorter stay at the park. however, some other studies of the field did not include age as a segmentation variable (downward & lumsdon 2004) or did not detect any significant relationship between expenditure patterns concerning age (chhabra 2007; spotts & mahoney 1991). since age has been found as one of the most relevant aspects that is considered to better understand and forecast the needs, interests and associated consumption behaviours of tourists, as mentioned above, further research is required on this aspect to provide more precise information for marketers. comparing groups of visitors by age with regard to their expenditure patterns will possibly allow both known and new age lucrative markets to national parks to be explored. this will be useful when proposing strategic plans to grow visitation figures amongst those nature-based segments with the highest propensity to spend. this is particularly important for less developed countries, where a large portion of the world’s biodiversity is concentrated and where the finance generated and secured from nature-based tourism can foster and support the sustainable development of an area. based on this, the purpose of this article is to use age to investigate the expenditure patterns amongst a sample of visitors to the table mountain national park (tmnp), along with other socio-demographics and their reasons for the visit. the latter data are to assist with profiling the age groups and to match the ‘product‘s’ specificities with the public’s specific demands. research method and design top ↑ setting the tmnp is one of 22 national parks in south africa and was officially established in 1988. it stretches from signal hill in cape town to cape point at the southern end of the cape peninsula and covers 25 000 ha. this urban park is famous because of table mountain itself, its historical and cultural heritage and its world heritage status. south african national parks (sanparks), which is the custodian of national parks in south africa, indicated the promotion of the country’s nature-based tourism as one of its core pillars. without compromising sustainability, self-generated revenues from commercial operations are considered necessary to supplement government funding of conservation management. the tmnp is amongst the five top south african parks with the highest number of guests. statistics for 2011 reported 2 344 340 guests for the tmnp, followed by the kruger national park with 1 411 796 guests, the west coast national park with 201 137 guests, the tsitsikamma national park with 180 107 and the addo national park with 138 079 guests. compared to the previous year (2010), total guests-to-parks numbers increased by 3.6%. specifically, the tmnp increased by 5.6%, whilst the kruger national park increased by 1.8% (sanparks 2012). the main reason for selecting the tmnp was that this park attracts a large number of visitors and shows the highest representation of international tourists, as well as a more equal spread of respondents over different age groups, compared to other south african national parks (saayman, kruger & fouche 2009). this can be explained by the facts that tmnp is a scenic park, offering the ‘majestic mountain’ as one of the new seven wonders, together with ocean views, mountains, beaches, forests and a variety of popular recreational activities. design the study sample included day visitors to the park from 27 march to 01 april 2010. six fieldworkers (three groups of two) approached visitors whilst they were queuing at the three popular entry points to the park: the cable car to table mountain, boulders and cape point. it is difficult to have fieldworkers at all entry points as there are also open access areas; therefore, it was decided to focus on these three, as they could be considered the most important access points and tourists generally visit all these areas.those visitors who expressed willingness to take part in the research were first asked a number of screening questions to detect the expenditure patterns from only one member per family or travel group. this information avoided repeat data and biases in the actual expenditure patterns amongst the respondents. in total, 441 questionnaires were collected during the study period, of which 404 were usable after a preliminary screening. according to israel (2009), in a population of > 100 000 (n), 398 respondents (n) are seen as representative and result in a 95% level of confidence with a ± 5% sampling error. thus, 404 usable questionnaires were deemed sufficient for the purposes of this study. procedure the research used a structured self-administered questionnaire, developed on the basis of the questionnaires used by past researchers (oberholzer et al. 2009; saayman, saayman & ferreira 2009). the questionnaire consisted of three main sections, (1) socio-demographic details, (2) expenditure patterns and (3) reasons for the visit. socio-demographics taking into consideration the goal of this study, age was considered pre-eminent compared to the other socio-demographic characteristics gathered here and this factor was detected by an open-ended question: ‘year of birth’. the respondents’ ages were calculated and post-coded in groups on the basis of the strong association found between age and household finance (income and expenditure) (down 2000; mok & iverson 2000). only respondents over 18 years of age were surveyed, as they are more likely to travel by themselves and have money to spend. respondents were categorised into one of the following four age groups: 18–29 years, 30–49 years, 50–64 years and 65–74 years (see table 1). table 1: location of respondents by age group in table mountain national park. the age groups were characterised based on the expenditure patterns per day per person, together with a trip-related characteristic, reasons for the visit and other socio-demographics. two other socio-demographic details were also included in the questionnaire to assist with profiling the age groups: ‘region of residence’ and ‘education’ (‘no school’, ‘matric’, ‘degree’, ‘postgraduate’, ‘professional’ and ‘other’). the region of residence was post-coded as follows: ‘north america’, ‘australasia’, ‘europe’, ‘south africa’ and ‘other countries’. finally, a question about the awareness of being in a national park was included. a dichotomous response format (‘yes’ or ‘no’) was used. expenditure patterns the expenditure patterns section was developed based on past research (kruger et al. 2010; saayman, van der merwe & pienaar 2009). spending behaviours were captured as follows: ‘money spent for categories or services per person’ (‘entrance and conservation fee’, ‘restaurants’, ‘food’, ‘beverages’, ‘transport to the park’, ‘activities’, ‘souvenirs and jewellery’) and ‘number of people paid for’ (including the respondent). on the basis of responses to ‘money spent for categories or services’, the total expenditure was calculated. to compute the total expenditure per person, the total expenditure was divided by the number of people paid for. reasons for visiting the park one measure that was developed ad hoc for this study concerned the reasons for visiting the park and the age groups were also characterised according to this. these reasons were found to be connected to age and can be considered amongst the most important personal variables for a better understanding of the consumers’ profile (see crompton 1979; crompton & mckay 1997; iso-ahola 1999). the scale for this question was developed based on common motivations identified from the tourism literature (crompton 1979; kim, jogaratnam & noh 2006; kruger & saayman in review; richards & wilson 2003), which include, (1) relaxation and getaway, (2) knowledge, (3) family togetherness, (4) natural beauty admiration and (5) activities participation. the scale included 26 items (four to five for each motivation) and participants were asked to indicate the importance of each item using a 5-point likert scale (1 = ‘not at all important’, 2 = ‘less important’, 3 = ‘important’, 4 = ‘very important’, 5 = ‘extremely important’). the relaxation and getaway motivation included items such as, ‘to get away from routine’ and ‘to relax’. the knowledge motivation items included, ‘learn about plants and animals’ and ‘learn more about specific marine life’. the family togetherness motivation included statements such as, ‘family recreation’ and ‘benefit of children’. the natural beauty admiration motivation included items such as, ‘tmnp is world renowned’ and ‘for the scenic beauty and view’. finally, the activities participation included statements such as, ‘mountain biking’ and ‘surfing’. analysis statistical data analysis was performed using spss version 18 (2009). the four age groups were characterised using chi-square, kruskal-wallis and analysis of variance (anova) tests based on expenditure patterns, along with psychographics (reasons for visiting the park) and socio-demographics. results top ↑ respondents’ profile in the tmnp, the largest group of respondents (196, 48.5%) were 30–49 years old. this was followed by those aged 18–29 years (130, 32.2%). fewer in number were respondents aged 65–74 years (49, 12.1%) and those aged 50–64 years (29, 7.2%). during the 5-day study period, the average age of respondents was 41.3 years (s.d. = 15.69).the median of the total expenditure per respondent per day person was r125 and the majority of respondents (71.2%) said that they had paid for one or two people, including themselves. this group was followed by those respondents who paid for three or four people (22.7%). the majority of respondents were well-educated, with 37.6% holding a degree, 22.6% in professional occupations and 18.3% holding a postgraduate qualification. this result, too, is consistent with previous research (e.g. marques, reis & menezes 2010; saayman & slabbert 2004; saayman, kruger & fouche 2009; the international ecotourism society 2006). from this study, it was found that the tmnp is attracting a higher percentage of international tourists compared to local tourists. almost half (46.1%) of the visitors to the tmnp were foreigners from europe. south african tourists made up almost one-third of all respondents (28.9%), whilst north americans represented one-tenth (10.7%). in contrast, previous research at other south african national parks (saayman, kruger & fouche 2009) highlighted a higher percentage of local visitors compared to international ones. reasons for park visit a principal component analysis with oblimin rotation supported a five-factor structure, explaining 71% of the total variance: • ‘activities and adventure sports performance’ • ‘park’s attributes admiration’ • ‘knowledge and inner experience’ • ‘escape’ • ‘social interactions’. the scree plot of eigenvalues suggested that no further factors contained reliable systematic variance. satisfactory reliabilities were observed for the scale scores (0.95, 0.68, 0.84 and 0.79) with the exception of social interactions (0.51). the results are shown in table 2. table 2: principal component analysis on respondents’ motives for visiting table mountain national park. factor scores, calculated as the average of all items contributing to a specific factor, indicated the most important reasons for visiting the park were, ‘escape’ (3.49), followed by ‘park’s attributes admiration’ (3.46). less important reasons were ‘social interactions’ (3.28), ‘knowledge and inner experience’ (3.12) and ‘activities and adventure sports performance’ (2.26). these results are similar to those of studies at other south african national parks (saayman, kruger & fouche 2009), which, in most cases, found the main reason for visiting national parks was the need to escape and relax. expenditure patterns by age the kruskal-wallis test was used for continuous variables that displayed a non-normal distribution. the respondents aged 18–29 years reported the highest expenditure per person and a higher expenditure per person per day when compared to the older group aged 30–49 years. specifically, the median expenditure for the group aged 18–29 years is r160 and the median for the group aged 30–49 years is r120. the former group was above, whilst the latter was slightly below the overall median, which was r125. no significant statistical differences could be established between the groups aged 18–29 years and 30–49 years and the groups aged 50–64 years and 65–74 years, although the median of the expenditures per person was noticeably less than for the groups younger than 50 years. this is because, on the one hand, the older groups (50–64 years and 65–74 years) were fewer in numbers (29 and 49 respondents, respectively) whilst, on the other hand, few of them provided information about their expenditure patterns (10 and 15 respondents, respectively) (see table 3). table 3: expenditures per person by age group for respondents at table mountain national park. most of the travel expenditures were allocated to entrance and conservation fees (48%), whilst smaller percentages characterised restaurants (16%), transport (9%), food (8%) and activities (8%). however, no significant differences were found amongst the age groups in expenditures per person for each of these categories. segment characterisation an anova analysis on the five psychographic characteristics found significant differences for the motivational dimension ‘social interactions’ by age groups (f = 3.18, df = 3/356, p < 0.05). respondents aged 18–29 years or 30–49 years were more likely to be motivated for the visit by the possibility of being with family (or someone special) or with friends, when compared to the group aged 65–74 years. however, these results should be interpreted with scepticism because of the low reliability of the scale (see table 4). table 4: analysis of variance of reasons for the visit to table mountain national park by age group. statistically significant differences were also found for the awareness of being in a national park [χ2 (3=397) = 8.12, p < 0.05]. interestingly, although the majority of visitors amongst all age groups were aware, a higher number of younger respondents aged 18–29 years (21 out of 44) and those aged 30–49 years (20 out of 44) stated they were unaware that they were in a national park, compared to the older groups.statistically significant differences were also found in the region of residence [χ2 (9=353) = 30.1, p < 0.01]. in particular, more respondents aged 30–49 years (45.3% and 31.8%, respectively) and 18–29 years (31.8% and 32.5%, respectively) came from europe and south africa, compared to the older groups. older respondents aged 50–64 years (46.2% and 26.9%, respectively) and 65–74 years (44.2% and 30.2%, respectively) came from europe and north america (see table 5). table 5: region of residence of respondents in table mountain national park by age group. the education level of three groups also differed [χ2 (12=385) = 47.12, p < 0.01]. whilst most of the age groups held a diploma or degree, more respondents aged 65–74 years (32.6%) and 30–49 years (24.7%) were professionals, compared to those aged 18–29 years (12%) and 50–64 years (18.5%). again, more respondents aged 30–49 years (22.1%), followed by those aged 18–29 years (17.6%) and 50–64 years (14.8%) held a postgraduate qualification (see table 6). table 6: education of respondents in table mountain national park by age group. ethical considerations top ↑ respondents participated of their own free will and had the opportunity to decline participation. no information was obtained that could personally identify respondents. trustworthiness top ↑ the authors did everything in their power to ensure that the sample and the survey was conducted in a systematic and scientific manner. results of the profile confirm previous research conducted in national parks. discussion top ↑ unlike previous research, this study specifically targeted visitors to a national park on the basis of their travel expenditure patterns by age range, along with other socio-demographics and their reasons for the visit. the latter assisted with profiling the age groups and to match the ‘product’s’ specificities with the public’s specific demands. from a theoretical standpoint, the findings differ from those detected by previous research. then, too, some contradictions representing rich material to foster new research directions have been highlighted as follows. firstly, as expected by the researchers, a considerable number of younger respondents (18–29 years) (32.2%) were detected in the area and this differs from the general notion and results by other researchers that mostly older people visit national parks. the low percentages of respondents older than 50 years (50–64 years, 7.2% and 65–74 years, 12.1%) were therefore unexpected, compared to previous findings (beh & bruyere 2007; galloway 2002; kibicho 2006; marques et al. 2010; kruger et al. 2010; saayman, kruger & fouche 2009; the international ecotourism society 2006). secondly, the finding related to spending behaviour highlights the presence of a new and potentially lucrative market when compared to those detected by previous research (kruger et al. 2010; mehmetoglu 2007); that is, a younger segment of respondents aged 18–29 years. this group was characterised by the highest expenditure per person and a higher expenditure per person when compared to those aged 30–49 years. then, younger respondents seem to represent a rather lucrative market, which would be a crucial variable for the maintenance and the sustainable growth of the area and is in line with the worldwide trends of youth tourism. the youth tourist sector, which accounts for 20% of the international tourism market, is growing faster than most other travel segments, with a global volume growth of 3% – 5% a year and is valued at approximately $165 billion per year. estimates show that youth and student travel market will reach 300 million arrivals by 2020 and represent $320 billion in market values (world youth student and educational travel confederation 2011). furthermore, the reasons for visiting a national park showed some differences in terms of age groups. specifically, significant differences were found for the motivational dimension ‘social interactions’: respondents aged 18–29 years and those aged 30–49 years revealed themselves to be more likely to be motivated by a possibility to spend time with family or friends, compared to the oldest group aged 65–74 years. unlike past research (e.g. moscardo & green 1999) that found differences concerning the activities participated in by respondents’ age, no statistically significant differences were found here in regard to the motivational dimension ‘activities and adventure sports performance’ by age group. in addition, unlike past research (kim, lee & klenosky 2003), age groups reported similarities for most of the motivational factors detected. finally, it is worth noting that amongst those respondents who were not aware of being in a national park, the majority were younger than 50 years old (i.e. in the 18–29 years and 30–49 years age groups). in brief, visitors participating in the research and aged 18–29 years were the second largest group and the heaviest spending segment per person. they were more motivated to visit for the day, when compared to respondents aged 65–74 years, because of the possibility of spending time with family (or someone special) and friends, they were less aware of being in a national park when compared to respondents aged 65–74 years, they mainly held a postgraduate qualification and largely came from europe or south africa. those aged 30–49 were the largest group but the lowest spending segment per person; the rest of their profile is similar to that of the youngest group. no statistically significant results were detected for the older groups concerning their expenditure patterns. practical implications based on the findings of this research, some practical implications for the sustainable development and financing of the tmnp have been outlined. the promotion of a mere increase in the number of visitors to capture economic benefits is to be avoided because it may have a cost to the natural environment (see eagles et al. 2002). therefore, a de-marketing strategy is considered necessary, but directed at the segment that showed the strongest propensity to spend (aged 18–29 years) and whose growth will help sustain the park. however, because the younger market at national parks is generally less present and known compared to the older segments, marketing to the younger groups should be conducted by first investigating the impacts of their activities. this relates also to park visitor management capacity. by contrast, no marketing strategy should be considered for the next oldest age respondents (aged 30–49 years) who were found to be lower-spenders and whose numbers are already significant and stable in this particular national park, as well as in worldwide national parks (e.g. saayman, kruger & fouche 2009). recommendations the following are some ideas for a specific marketing strategy. increasing the frequency of visits and attracting the younger market aged 18–29 years could be achieved by offering, for instance, a youth loyalty card, especially to those visitors residing in south africa, of whom there are considerable numbers amongst the younger respondents. hosting youth days might be another successful strategy to attract this market. in addition, a marketing strategy could include social-related content, as these respondents were found to be more likely to be motivated by the possibility of spending time with family and friends. for example, picnic and braai areas, such as the buffels bay and bordjiesdrif, or the tokai picnic and braai area and its wide space for large groups, can be promoted. as an urban park offering several activities and adventure sports, a marketing strategy could be used also to target additional younger segments such as the ‘harder eco-tourists’ identified by weaver and lawton (2002) or the ‘self-centred visitors’ and the ‘occasional visitors’ identified by marques et al. (2010), who reported a high propensity to take part in these types of activities. finally, as mostly younger participants seemed to be unaware they were in a national park, the management should provide more information about this. educational programmes could represent another marketing tool for promoting repeated visits to the park and, in general, to other national parks. in fact, several research findings have highlighted how practical experiences in outdoor settings, such as hands-on conservation activities and on-site educational programmes at an early age, shape and promote positive interests and associated behaviours towards the environment, thus ensuring nature conservation and preservation (mcduff & jacobson 2000; palmer 1993; sward 1996; voordouw 1987). limitations a limitation of the current research is the difficulty in generalising these results to a wider population of the park. to test the general applicability of these findings to other samples, this study should be replicated. secondly, previous research highlighted the importance of trip-related characteristics (e.g. type of accommodation, length of stay) for the spending segments characterisation. however, these variables could not be included in the present research because the tmnp, compared to other south african national parks, has a high percentage of day visitors and low percentage of repeat day visitors and overnight visitors (saayman, kruger & fouche 2009). this may be due, in part, to the fact that the majority of the visitors across all ages were foreigners from overseas visiting the tmnp as one of the stops on their tour. therefore, the tmnp management should do more, in terms of marketing plans, to target the domestic visitor market, for example, by providing discounts and/or loyalty cards to south africans. thirdly, unlike most past research that found motivational differences by age, here the characterisation of the age groups may be considered as limited because just one factor of five motivational factors significantly differentiated the age groups. future research is required to shed more light on this point. fourthly, although past research highlighted the importance of the activities participated in, in terms of increasing expenditure patterns (mehmetoglu 2007), this study was unable to find any differences in terms of the activities as reasons for the visit by the age (spending) segments. further research should again be conducted to shed more light on this point. finally, this study was unable to find any differences amongst the age groups in terms of expenditures per person for each of the categories considered (e.g. restaurants, food, etc.). however, it would be interesting to know what younger groups spent the extra money on. future research should be conducted to shed more light on this point. conclusion top ↑ this research adds to existing literature by shedding more light on the expenditure patterns (behaviour) of a sample of visitors to the tmnp, grouped by age range, and contributes to the topic of visitor behaviours to nature-based attractions. the respondents’ profile was found to differ significantly from the profiles of visitors to other parks, as the tmnp respondents are younger and most of them are foreigners. as parks are generally visited by older people, this study highlights the importance of a younger market. future research in national parks should pay particular attention to this market, which was found as an emerging but very relevant economic market and is anticipated as playing a key role in supporting sustainable lifestyles in the future (coetzee & saayman 2009; council for game and wildlife conservation 2003). acknowledgements top ↑ the authors want to acknowledge the following people and institutions. firstly, sanparks for funding and allowing the research, especially, mr glenn phillips and bheki zwane. secondly, the national research foundation for funding this research. thirdly, all the respondents for completing the questionnaires and, lastly, the reviewers for their valuable comments. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions m.s. (north-west university) was the project leader in developing the questionnaire and conducting the survey. f.c. (north-west university) was the main author and also responsible for the statistical analysis. both authors reviewed and finalised the article. references top ↑ beh, a.w. & bruyere, b.l., 2007, ‘segmentation by visitor motivation in threen kenyan national reserves’, tourism management 28, 1464–1471. http://dx.doi.org/10.1016/j.tourman.2007.01.010buckley, r., 1994, ‘a framework for ecotourism’, annals of tourism research 21, 661–669. http://dx.doi.org/10.1016/0160-7383(94)90126-0 buckley, r., 2002a, a global triple-bottom-line evaluation for ecotourism, world ecotourism summit, quebec. buckley, r., 2002b, ‘managing tourism in parks: research priorities of industry associations and protected area agencies in australia’, journal of ecotourism 1, 162–172. http://dx.doi.org/10.1080/14724040208668122 bushell, r. & mccool, s.f., 2007, ‘tourism as a tool for conservation and support of protected areas’, in r. bushell & p.f.j. eagles (eds.), tourism and protected areas: benefits beyond boundaries, pp. 12–26, cabi international, wallingford. bushell, r., staiff, r. & eagles, p., 2007, ‘tourism and protected areas: benefits beyond boundaries’, in r. bushell & p.f.j. eagles (eds.), tourism and protected areas: benefits beyond boundaries, pp. 1–11, cabi international, wallingford. butler, r.w. & boyd, s.w., 2000, tourism and national parks: issues and implications, john wiley and sons, new york. cha, s., mccleary, k.w. & uysal, m., 1995, ‘travel motivations of japanese overseas travellers: a factor-cluster segmentation approach’, journal of travel research 34, 33–39. http://dx.doi.org/10.1177/004728759503400104 chhabra, d., 2007, ‘estimating benefits and costs of casino gambling in iowa, united states’, journal of travel research 46, 173–182. child, b. & dalal-clayton, b., 2004, ‘transforming approached to cbnrm: learning from the luangwa experience, zambia’, in t.o. mcshane & m.p. wells (eds.), getting biodiversity projects to work: towards more effective conservation and development, pp. 256–289, columbia university press, new york. coetzee, w. & saayman, m., 2009, ‘sustainability and ecotourism’, in m. saayman (ed.), ecotourism – getting back to basics, pp. 124–146, leisure publications, potchefstroom. pmid:19290941 council for game and wildlife conservation, 2003, ‘young people: the biggest lobby group for sustainable use’, african indaba 1, 4–6. crompton, j.l., 1979, ‘motivations for pleasure vacation’, annals of tourism research 6, 408–424. http://dx.doi.org/10.1016/0160-7383(79)90004-5 crompton, j.l. & mckay, s.l., 1997, ‘motives of visitors attending festival events’, annals of tourism research 24, 425–439. http://dx.doi.org/10.1016/s0160-7383(97)80010-2 down, d., 2000, family spending: a report on the 1999–2000 family expenditure survey, the national statistics office, london. pmcid:pmc3695199 downward, p. & lumsdon, l., 2004, ‘tourism transport and visitor spending’, journal of travel research 42, 415–420. http://dx.doi.org/10.1177/0047287504263038 eagles, p.f.j., mccool, s.p. & haynes, c.d., 2002, sustainable tourism in protected areas. guidelines for planning and management, iucn publications services unit, cambridge. frochot, i. & morrison, a.m., 2000, ‘benefit segmentation: a review of its applications to travel and tourism research’, journal of travel & tourism marketing 9, 21–45. http://dx.doi.org/10.1300/j073v09n04_02 galloway, g., 2002, ‘psychographic segmentation of park visitor markets: evidence for the utility of sensation seeking’, tourism management 23, 581–596. http://dx.doi.org/10.1016/s0261-5177(02)00025-0 geldenhuys, s. & saayman, m., 2009, ‘ecotourism venues’, in m. saayman (ed.), ecotourism – getting back to basics, pp. 26–42, leisure publications, potchefstroom. gibson, h., attle, s.p. & yiannakis, a., 1998, ‘segmenting the active sport tourist market: a life span perspective’, journal of vacation marketing 4, 52–64. http://dx.doi.org/10.1177/135676679800400105 iso-ahola, s.e., 1999, ‘motivational foundations of leisure’, in e.l. jackson & t.l. burton (eds.), understanding leisure recreation, mapping the past, charting the future, pp. 35–51, venture publishing, state college. israel, g.d., 2009, determining sample size, florida state university, cooperative extension service, gainesville. international union for conservation of nature, 2004a, 2003 durban world park congress, protected areas programme, gland. international union for conservation of nature, 2004b, can protected areas contribute to poverty reduction?, iucn, gland. jang, s., ismail, j.a. & ham, s., 2002, ‘heavy spenders, medium spenders, and light spenders of japanese outbound pleasure travelers’, journal of hospitality & leisure marketing 9, 83–106. http://dx.doi.org/10.1300/j150v09n03_07 jang, s.c., bai, b., hong, g.s. & o’leary, j.t., 2004, ‘understanding travel expenditure patterns: a study of japanese pleasure travelers to the united states by income level’, tourism management 25, 331–341. http://dx.doi.org/10.1016/s0261-5177(03)00141-9 johns, n. & gymothy, s., 2002, ‘market segmentation and the prediction of tourist behavior: the case of bornholm, denmark’, journal of travel research 40, 316–327. http://dx.doi.org/10.1177/0047287502040003009 jönsson, c. & devonish, d., 2008, ‘does nationality, gender and age affect travel motivation? a case of visitors to the caribbean island of barbados’, journal of travel and tourism marketing 25, 398–408. http://dx.doi.org/10.1080/10548400802508499 kibicho, w., 2006, ‘tourists to amboseli national park: a factor-cluster segmentation analysis’, journal of vacation marketing 12, 218–231. http://dx.doi.org/10.1177/1356766706064618 kim, k., jogaratnam, g. & noh, j., 2006, ‘travel decisions of students at a us university: segmenting the international market’, journal of vacation marketing 12, 345–357. http://dx.doi.org/10.1177/1356766706067606 kim, s.s., lee, c. & klenosky, d.b., 2003, ‘the influence of push and pull factors at korean national parks’, tourism management 24(2), 169–180. kotler, p., 1992, ‘total marketing’, business week advance, executive brief 2. kruger, m. & saayman, m., in review, ‘segmenting visitors to the addo elephant national park based on education level’, journal of tourism research and hospitality, submitted 2013. kruger, m., saayman, m. & saayman, a., 2010, ‘expenditure-based segmentation of visitors to tsitsikamma national park’, acta commercii 10(1), 137–149. mabunda, d.m. & wilson, d., 2009, ‘commercialization of national parks: south africa’s kruger national park as an example’, in j. saarinen, f. becker, h. manwa & d. wilson (eds.), sustainable tourism in southern africa, pp. 116–133, channel view publications, bristol. marques, c., reis, e. & menezes, j., 2010, ‘profiling the segments of visitors in portuguese protected areas’, journal of sustainable tourism 18, 971–996. http://dx.doi.org/10.1080/09669582.2010.497222 mcduff, m. & jacobson, s., 2000, ‘impacts and future directions of youth conservation organizations: wildlife clubs in africa’, wildlife society bulletin 28, 414–425. mehmetoglu, m., 2007, ‘typologising nature-based tourists by activity – theoretical and practical implications’, tourism management 28, 651–660. http://dx.doi.org/10.1016/j.tourman.2006.02.006 mok, c. & iverson, t.j., 2000, ‘expenditure-base segmentation: taiwanese tourists to guam’, tourism management 21, 299–305. http://dx.doi.org/10.1016/s0261-5177(99)00060-6 moscardo, g. & green, d., 1999, ‘age and activity participation on the great barrier reef’, tourism recreation research 24, 57–68. moutinho, l., 1987, ‘consumer behaviour in tourism’, european journal of marketing 21, 5–44. http://dx.doi.org/10.1108/eum0000000004718 myburgh, e. & saayman, m., 1999, ecotourism in action: practical guidelines and principles, 1st edn., leisure consultants and publications, potchefstroom. oberholzer, s., saayman, m., saayman, a. & slabbert, e., 2010, ‘the socio-economic impact of africa’s oldest marine park’, koedoe 52(1), 1–9. oh, y.j., cheng, c.k., lehto, x.y. & o’leary, j.t., 2004, ‘predictors of tourists’ shopping behaviour: examination of socio-demographic characteristics and trip typologies’, journal of vacation marketing 10, 308–319. http://dx.doi.org/10.1177/135676670401000403 palmer, j., 1993, ‘development of concern for the environment and formative experiences of educators’, journal of environmental education 24, 26–30. http://dx.doi.org/10.1080/00958964.1993.9943500 patterson, i. & pegg, s., 2009, ‘marketing the leisure experience to baby boomers and older tourists’, journal of hospitality marketing & management 18, 254–272. http://dx.doi.org/10.1080/19368620802594136 richards, g. & wilson, j.y., 2003, new horizons in independent youth and student travel, a report for the international student travel confederation and the association of tourism and leisure education, istc, amsterdam. saayman, m., 2009, ‘community tourism’, in m. saayman (ed.), ecotourism – getting back to basics, pp. 67–96, leisure publications, potchefstroom. saayman, m. & slabbert, e., 2004, ‘a profile of tourists visiting the kruger national park’, koedoe 47, 1–8. http://dx.doi.org/10.4102/koedoe.v47i1.74 saayman, m., saayman, a. & du plessis, c., 2005, ‘analysis of spending patterns of visitors of three world cup cricket matches in potchefstroom, south africa’, journal of sport tourism 10, 211–221. http://dx.doi.org/10.1080/14775080600611491 saayman, m., kruger, m. & fouche, m., 2009, executive summary of research profiles of selected national parks in south africa, institute for tourism and leisure studies, potchefstroom. saayman, m., saayman, a. & ferreira, m., 2009, ‘the socio-economic impact of the karoo national park’, koedoe 51(1), 158–168. saayman, m., van der merwe, p. & pienaar, j., 2009, ‘expenditure-based segmentation of tourists to the kruger national park’, acta academica 41, 107–127. seaton, a.v., 1996, ‘the analysis of tourism demand: market segmentation’, in a.v. seaton & m.m. bennett (eds.), marketing tourism products, pp. 28–54, international thomson business press, london. sherman, e. & schiffman, l.g., 1991, ‘quality-of-life (qol) assessment of older consumers: a retrospective review’, journal of business and psychology 6, 107–119. http://dx.doi.org/10.1007/bf01013687 south african national parks, 2010, annual report 2009–2010, viewed 20 january 2011, from http://www.sanparks.org/about/annual/default.php south african national parks, 2012, annual report 2011–2012, viewed 20 november 2012, http://www.sanparks.org/about/annual/default.php spotts, d.m. & mahoney, e.m., 1991, ‘segmenting visitors to a destination region based on the volume of their expenditure’, journal of travel research 29, 24–31. http://dx.doi.org/10.1177/004728759102900405 spss version 18.0 for windows, 2009, computer software, spss inc., chicago. sward, l., 1996, ‘experiential variables affecting the environmental sensitivity of el salvadoran environmental professionals’, paper presented at the annual conference of the north american association of environmental education, burlingame, california, 01–02 november. the international ecotourism society, 2006, global ecotourism fact sheet, viewed 21 january 2011, from http://mekongtourism.org/website/wp-content/uploads/downloads/2011/02/fact-sheet-global-ecotourism-iets.pdf thrane, c. & farstad, e., 2011, ‘domestic tourism expenditures: the non-linear effects of length of stay and travel party size’, tourism management 32, 46–52. voordouw, j.j., 1987, youth in environmental action: an international survey, joint publication by international youth foundation, international union for the conservation of nature and united nations environment programme, gland. waite, c., n.d., travel features: south africa – cape town, viewed 21 february 2014, from http://www.boardseekermag.com/travel_features/southafrica/south_africa.htm weaver, d.b. & lawton, l.j., 2002, ‘overnight ecotourist market segmentation in the gold coast hinterland of australia’, journal of travel research 40, 270–280. world commission on protected areas, 1998, economic values of protected areas: guidelines for protected area managers, joint publication by task force on economic benefits of protected areas of the world commission on protected areas and economics service unit of international union for the conservation of nature, gland. world tourism organization, 2002, sustainable development of ecotourism: a compilation of good practices, wto, madrid. world wildlife fund international, 2001, guidelines for community-based development, wwfi, ledbury. world youth student and educational travel confederation, 2011, the power of youth travel, viewed 21 february 2014, from http://wysetc.org/publications/power-of-youth-travel/ abstract introduction methods results discussion conclusion acknowledgements references about the author(s) cathy m. dzerefos school of animal, plant and environmental sciences, university of the witwatersrand, johannesburg, south africa ed t.f. witkowski school of animal, plant and environmental sciences, university of the witwatersrand, johannesburg, south africa citation dzerefos, c.m. & witkowski, e.t.f., 2016, ‘bridging the knowing–doing gap in south africa and the role of environmental volunteer groups’, koedoe 58(1), a1394. http://dx.doi.org/10.4102/koedoe.v58i1.1394 note: this article is dedicated to the friends of the haenertsburg grassland (frohg) for acting on the words of archbishop desmond tutu ‘do your little bit of good where you are; it’s those little bits of good put together that overwhelm the world’. original research bridging the knowing–doing gap in south africa and the role of environmental volunteer groups cathy m. dzerefos, ed t.f. witkowski received: 12 apr. 2016; accepted: 24 aug. 2016; published: 31 oct. 2016 copyright: © 2016. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the implementation gap between science, policy and practice has led to loss of biodiversity and ecosystem services throughout africa and is described in a case study from limpopo province, south africa. in 2006, the south african national biodiversity institute first highlighted the woodbush granite grassland (wgg) in the greater tzaneen local municipality as the only critically endangered ecosystem in limpopo province. five years later (2011), the critically endangered listing was published in the government gazette no. 34809. after repeated and sustained efforts for many years from volunteers of a local environmental group – currently known as the friends of the haenertsburg grassland (frohg) – in 2015 the intent to formally protect 126 ha was published in the government gazette no. 2609. unfortunately, the proposed protected area accounts for only 66% of the largest remaining fragment of wgg, which excludes an important colony of medicinal plants. considering that only 6% of the original extent of wgg remains in an untransformed state the whole fragment should be conserved. non-alignment of municipal spatial priorities, as in the haenertsburg town plan from 1896, to provincial and national environmental priorities has resulted in numerous incidents that have degraded what little remains of the wgg ecosystem. failure of the provincial authorities to act timeously to enforce environmental regulations resulted in the frohg successfully involving national authorities to stop illegal land occupation while another incident involving an illegal fence was resolved 9 years after erection. a strengthened relationship with lepelle northern water has resulted in better planning of activities in relation to an existing pipeline. this case study shows various avenues available to environmental volunteer groups in south africa and suggests that long-term lobbying can yield positive results. conservation implications: formal conservation of wgg through the intended nature reserve proclamation represents application of environmental legislation (notably listing notice 3, national environmental management act 107 of 1998: environmental impact assessment regulations, 2014), scientific recommendations and policy. better cooperation between provincial administration and frohg will benefit the protection and management of wgg. introduction in many parts of africa, biodiversity (noss 1990) and functioning ecosystems remain intact (opio-odongo 2013), providing beneficial plants and animals (dzerefos, witkowski & kremer-köhne 2016; novellie, biggs & freitag-ronaldson 2013; sango & godwell 2015), water storage and flood retention services (opio-odongo 2013), soil carbon and nitrogen (tesfaye et al. 2016) and areas for tourism or spiritual significance (cer 2015; dzerefos 2004). to ensure sustainable development necessary to alleviate poverty, unemployment and illiteracy, environmental legislation and implementation are essential to maintain ecological infrastructure (dea 2013). the translation of science into the decision-making process and practical application by government administration has been identified as obstacles to securing environmental sustainability in africa (cowling & wilhelm-rechmann 2007) even in a relatively politically and economically secure country like south africa, which has holistic and sound environmental legislation (cbd 2014). climate change research for sub-saharan africa indicates increased frequencies of droughts and flooding impacting negatively on biodiversity and ecosystem services (dea 2013; sango & godwell 2015). the grassland biome is predicted to be most at risk (carbutt & martindale 2014; dea 2013; thompson & swemmer 2014) with species responding negatively to habitat fragmentation through anthropogenic activities over a 50to 100-year time lag (fourie, rouget & lötter 2015). in response to these predictions, the south african adaptation strategy to climate change recommends securing upper water catchment grasslands to safeguard water storage, flood retention capacity and biodiversity (dea 2013). indeed in the last decade, 60 000 ha of grasslands have been proclaimed as protected areas (carbutt & martindale 2014). the limpopo province has one critically endangered ecosystem, the woodbush granite grassland (wgg) (desmet et al. 2013; mucina et al. 2006; rsa 2011), located in the greater tzaneen local municipality (gtm), in the foothills of the northern drakensberg escarpment (figure 1). wgg is a source of scarce plants for traditional healing and hand brushes that have regional cultural relevance (dzerefos et al. 2016). as a mere 1.8% was conserved (desmet et al. 2013) and 6% of the original extent is untransformed (niemandt 2015), there is a high risk that it will not persist into the future. the largest continuous fragment of wgg, having an area of 192 ha, lies adjacent to the small town of haenertsburg. it contributes to the headwaters of the ebenezer dam (seshoka, de lange & faysse 2004) through the kantoor and wilge rivers and two natural springs (moreroa 2014). the second largest wgg patch measures 60 ha and constitutes a peninsular jutting into ebenezer dam (figure 2) and land tenure rests with government. several much smaller wgg remnants, ranging in size from 5 ha to 25 ha, occur next to the magoebaskloof hotel, on privately owned farms or on communal grazing lands. up until october 2010, the haenertsburg grassland was under the custodianship of the national department of public works (dpw); thereafter, it was transferred to limpopo province. the intention to declare 126 ha (66%) of the haenertsburg grassland a provincial nature reserve was recently published (lpg 2015). the state-owned grasslands are continuous in the south with a private property known as danallen, which is characterised by three small patches of northern mistbelt forest (mucina & geldenhuys 2006) and drops steeply towards the wilge river. the government and private-owned land have no fences between them, and the louis changuion hiking trail traverses both. sections of wgg adjoin pine plantations and there is an active fire management programme spearheaded by the timber industry (dzerefos et al. 2016), which has resulted in a network of firebreaks and roads stretching from remote areas in the drakensberg escarpment to the lowlands. the wgg and the ebenezer dam are separated by an important arterial route (the r528) between polokwane and tzaneen. the ebenezer dam is a significant water source, supplying the provincial capital, polokwane, 60 km away, as well as rural settlements along the r528 with potable water (seshoka et al. 2004). haenertsburg town and most outlying residential plots and tourist developments in the area do not have adequate sewage management and use septic tanks and french drains (spatial development framework [sdf] 2009), which could pollute groundwater. the letaba river is the natural outflow conduit of ebenezer dam and serves numerous downstream users in the water-limited lowland areas including rural communities, industry and agriculture. over a decade ago, the water demands on the letaba river were found to exceed the capacity of the river (seshoka et al. 2004), which suggests that downstream water users who are further from the ebenezer dam, like the kruger national park and rural communities in mozambique, may not receive sufficient water. figure 1: haenertsburg town with woodbush granite grassland patches (in green) located in the foothills of wolkberg wilderness area, northern drakensberg escarpment, mopani district, limpopo province, south africa. figure 2: google earth pro image (2015) of the two largest remaining woodbush grassland fragments under public tenure. within the broader study area, steep altitudinal gradients that influence temperatures, frost and precipitation in conjunction with variable soil types lead to juxtaposition of habitats suitable for subtropical (avocado) and temperate (kiwi and blueberry) crops. this distinction is mirrored in the timber plantations, where pine grows optimally in the wetter areas and eucalyptus in the drier areas. towards polokwane in the west, the climate is hotter, drier and frost free and supports savannah vegetation. in contrast, a grassland–forest mosaic occurs in cooler, high-rainfall areas in the drakensberg escarpment and associated hills, such as where wgg occurs. the highest point in the escarpment, known as the iron crown (2126 m) is a mere 6 km from haenertsburg, which is located in undulating hills. a 40-km north-east descent along the r528 ends at an altitude of 752 m in tzaneen. tzaneen is known for its warm, humid, frost-free climate, which supports citrus, mango and litchi orchards. a distinct easterly rainfall gradient exists from the escarpment to the drier lowveld area and kruger national park. the milder climate of haenertsburg and immediate surroundings contributes to a tourism industry specialising in adventure sport, mountain biking, hiking and boating activities. the national environmental management: biodiversity act (2004), provides for listing threatened ecosystems, in order to reduce ecosystem and species extinction, by preventing further degradation in areas with exceptionally high conservation value (rsa 2011). biodiversity targets, to prevent loss of threatened species, have been calculated for south african ecosystems and are necessary for biodiversity planning, monitoring and management (cbd 2014). in south africa, biodiversity targets are determined using the species–area relationship, which results in a higher biodiversity target for species-rich ecosystems relative to species-poor ones (desmet & cowling 2004). hence, for wgg, a higher biodiversity target of 27% was set, relative to the 23% average for other ecosystems in limpopo (desmet et al. 2013; mucina et al. 2006) and 20% internationally. however, already 10 years ago, mucina et al. (2006) stated that wgg was critically endangered and this target cannot be met as only about 10% remained in a natural state. furthermore, wgg was prone to bush encroachment arising from both sour bushveld and scrub forest, which is compounded by fire exclusion (mucina et al. 2006) and climate change (thompson & swemmer 2014). large parts of wgg had been planted to exotic timber since the early 1900s (dzerefos 2004) or developed for agriculture, housing or tourism (moreroa 2014). more recently, a study of aerial photographs from 1948, 1977 and 2008, together with some ground truthing of land cover, where access to land was possible, indicated that only 6% of the original wgg remains untransformed (niemandt 2015). using remote sensing to assess wgg, desmet et al. (2013) calculated 45% as still intact, but this includes the whole forest–scrub–grassland mosaic as a functional management unit, whereas niemandt (2015) only considered the grassland areas. furthermore, the 6% was likely to be an over-estimate as some wetland and degraded areas may have been included as they could only be distinguished through site visits, which were not possible for the whole extent (niemandt 2015). the protection of grassland biodiversity has the simultaneous benefit of securing ecosystem services by increasing water capture and storage for lower catchments (carbutt & martindale 2014). the national environmental management: protected areas act (2003) and the national protected area expansion strategy (cbd 2014) are existing mechanisms that could have formally been used to secure the wgg. unfortunately, prior to the study by mucina et al. (2006), the description of wgg did not exist and it was included in the widespread north-eastern mountain grassland occurring from limpopo southwards to kwa-zulu natal and the eastern cape (low & rebelo 1996). even if wgg was recognised at the time, it may have been excluded because it is highly fragmented. south africa has a parliamentary system, operating at national, provincial (nine provinces) and local (district and local municipalities) government levels with an independent judiciary (rossouw & wiseman 2004). national, provincial and local levels of government have distinct but interrelated legislative and executive authority (figure 3; novellie et al. 2013) governed by values enshrined in the constitution. provincial environmental legislation, such as the limpopo environmental management act (lema) (act 7 of 2003) calls for sustainable and managed use of important biodiversity areas where natural resources are collected. government officials working in the environmental field through national or provincial legislation give effect to the international commitments, which south africa has signed, such as the convention on biological diversity (cbd 2014). hence, performance and delivery on environmental mandates at the provincial level are of national relevance (cer 2015; moreroa 2014; rossouw & wiseman 2004; wilhelm-rechmann & cowling 2013). figure 3: the framework within which environmental departments fulfil their mandates to protect biodiversity and ecosystem services, which is guided by policies and legislation which arise from the constitution, 1996 and inform national and provincial legislation. provincial authorities are mandated to advise local municipalities on ecological and conservation priorities required in the land-use planning process (wilhelm-rechmann & cowling 2013) and infrastructure provision. for instance, the spatial planning and land use management act (no. 16 of 2013) requires local government to prepare a spatial development plan (sdp) (rsa 2013). legislation such as lema as well as specialist sector reports, such as the limpopo conservation plan (desmet et al. 2013), should inform decision-making processes and guide the compilation of the sdf every 5 years (rsa 2013; sdf 2009; wilhelm-rechmann & cowling 2013). the limpopo province department of economic development, environment and tourism (ledet) is an organ of state accountable for the conservation of provincial natural areas (anthony, scott & antypas 2010). the provincial premier appoints a member of the executive, from among the provincial legislature, to head provincial departments. typically, the term of office is 5 years. since 2000, a community-based volunteer group, initially known as haenertsburg environmental monitoring and action group (hemag) and currently known as friends of the haenertsburg grassland (frohg), have actively engaged with ledet to ensure that biodiversity and ecosystem services of wgg are not compromised and environmental regulations enforced. as of march 2016, frohg had 81 paid-up members, who are environmentally aware residents with skills such as ecotourism, horticulture, farming, mapping and education. traditional healers and hand brush harvesters from a wider area have also been shown to greatly value this grassland’s biodiversity as it has cultural and economic value (dzerefos et al. 2016). this case study, of limpopo province’s only critically endangered ecosystem, wgg, is used to show how current legislation offers the required framework for specialist knowledge to be realised and the stewardship role that environmental volunteer groups can offer. methods historical records using the archives of the community-based, environmental group frohg, letters and emails to authorities, as well as memos, reports and government gazettes pertaining to wgg were consulted. individuals and groups that had a known interest in lobbying on behalf of wgg were contacted to contribute records of correspondence and the subsequent replies that were received from provincial authorities. letters collected date back to 1988 but the historical record peaks from 2005 when email communication predominated and hemag (now frohg) lodged a submission to ledet to have the haenertsburg grassland proclaimed a site of ecological importance through lema. records reporting illegal activities on wgg have also been kept. the most significant was the fencing of public land by an adjacent landowner in 2007 (figure 2) and the demarcation and selling of small plots in 2011 (troskie 2011). the email correspondence was arranged in terms of date, designation of recipient, organisation, subject and whether the matter was resolved. if the email was forwarded by the recipient to another person, this person was added to the database and because the recipient was copied on the email, it was logged that the initial person had replied. keeping a detailed record of correspondence with authorities identifies where and when there was a breakdown in communication and service delivery. these results are conservative estimates of communications as telephonic discussions and site inspections were unrecorded. moreover, the regional manager of the wildlife and environment society of south africa, a national non-profit organisation involved in biodiversity protection, had also corresponded with provincial and local authorities from 2008 to 2011, but unfortunately these records were deleted when the post became vacant. nonetheless, overall these records provide an important historical accounting of the various efforts made and responses received. important role players key events pertaining to the haenertsburg grassland were arranged in a timeline so that the performance of key role players could be easily viewed. to quantify efficiency and accountability, outstanding responses on issues pertaining to wgg were followed up with the three tiers of government, namely the national environmental crimes and incidents hotline (env-hotline), the limpopo legislature (public participation and petitions unit) and ledet, between june 2014 and march 2016. the env-hotline, operated by the national department of environmental affairs, is an avenue for the public to report non-compliance with environmental legislation (cer 2014). replies or non-replies were documented. all parties were informed of this research project and copies of past emails detailing relevant information were sent to recipients for ease of reference. ledet was invited to discuss the research method but there was a 4-month time delay before they responded, by which time data collection and analysis of available historical information had been completed. nonetheless, as the paper was written and reviewed, ledet officials were provided with opportunity to comment and provide input. historical town plan in mid-2014, images appearing on google maps of haenertsburg village and the adjacent grassland indicated settlement demarcations envisaged in the 1890s in areas of pristine wgg. the findings of the limpopo conservation plan (desmet et al. 2013) and land cover assessment of wgg (niemandt 2015), as well as inputs from traditional plant users (dzerefos et al. 2016), suggest that the original town plans should be aligned with national and provincial environmental priorities. this process requires the intervention of ledet. the outdated information was reported to google maps on 22 july 2014, by clicking on the ‘report a problem’ icon located at the bottom right of the image and also to the gtm town planners. results historical timeline high levels of transformation of wgg are evident from the 1970s (niemandt 2015) and coincide with the first letters sent to the authorities highlighting the importance of wgg. these letters were written by community members, the curators of two herbaria, the botanical society of southern africa and biodiversity specialists employed by the provincial government (table 1). financial support from the national lottery distribution trust fund allowed hemag to commission environmental education resources, biodiversity studies and an environmental management plan (emp) in 2004, according to national environmental management act 107 of 1998: environmental impact assessment regulations, 2014 (nema). a ledet ecologist commended the emp and recommended the inclusion of more areas. in 2005, hemag lodged a formal submission to ledet to declare the wgg around haenertsburg town a site of ecological importance. ledet officials, specialising in botany and herpetology, conducted investigations and recommended upscaling the proclamation to a nature reserve. table 1: chronology of events pertaining to the largest remaining continuous section of woodbush granite grassland, adjacent to haenertsburg village, limpopo province. from 2006 to 2013 (table 1), a number of degrading events occurred in the haenertsburg grassland, which led to loss of biodiversity and public frustration with environmental law enforcement. in an attempt to stop damage to the grassland, hemag erected no entry signage, indicating that it was a site of ecological importance. although this may have reduced off-road vehicle use, other misdemeanours continued. for example, a landowner having a residential property next to the grassland and the haenertsburg primary school bulldozed the grassland–forest margin to erect a fence on public property. although frohg (previously hemag) raised objections with the gtm (having a land-use planning responsibility) (wilhelm-rechmann & cowling 2013), ledet (having the responsibility to uphold environmental legislation and advise gtm on biodiversity priorities) and the dpw (land custodian until october 2010), each administrative body passed responsibility to the other. frohg sent 33 emails regarding the fence, between october 2007 and december 2008 to ledet and had a response rate of 42%. because of the failure of provincial department intervention in the fence matter, it was reported to the public participation and petitions unit for limpopo province by means of a petition signed by 165 individuals. the matter was heard by the petitions committee in july 2015, which ruled the fence illegal and instructed dpw to ensure that the fence be removed. the matter took 8 years to be heard during which time illegal sections of the fence were vandalised twice and antelope kept on the property escaped. the petitions committee ruling was implemented in september 2016 (table 2). table 2: follow up of outstanding issues with the national environmental crimes and incidents hotline, the limpopo legislature public participation and petitions unit and limpopo department of economic development, environment and tourism from june 2014 to september 2016. another incident, reported on in provincial newspapers, involved the illegal staking out and selling of 25 plots on the haenertsburg grassland by an individual claiming ownership of the area below the haenertsburg primary school (troskie 2011). the local and provincial authorities were immediately notified and provided with photographic evidence of unearthed grassland plants. the environmental compliance and enforcement division of ledet responded that it was not possible to issue a directive to the south african police service (saps) to stop the sub-division of land because the reserve application was not finalised and referred the frohg to the custodian of the land, the dpw. because of the lack of environmental capacity in dpw, the loss of protected and threatened biodiversity was not viewed as a matter of urgency and there was no indication that the matter would be attended to. consequently, the frohg alerted the south african national biodiversity institute (sanbi), a national parastatal involved in biodiversity research, planning and policy. sanbi urgently referred the matter to the select committee land and environmental affairs, parliament of south africa (operational from may 2009 to march 2014), which along with the public protector and the land claims office persuaded ledet that it was their responsibility to intervene. ledet then issued an official directive to the saps indicating that the demarcations were illegal and should be stopped. the saps and the gtm dismantled one of the shacks that had been erected and the stakes, marked with ‘buyers’ surnames, were removed. the office of the chief land claims commissioner, in a letter to the select committee dated december 2011, confirmed that the area was being investigated as part of a land claim made by the mmamabolo community. the letter confirmed that the staking of land had been done illegally, by an opportunist not of the mmamabolo community. because of the discriminatory legislation of the past, many communities had lost ancestral lands throughout south africa. a process was started in 1994 to research land claims and compensate dispossessed people. the staking out of plots was repeated in february 2013 by the same person, but was promptly stopped by the saps who remembered the directive from 2011. yester year and current maps the original town plan of haenertsburg, as surveyed in september 1887 (figure 4), was approved by the acting surveyor-general in march 1888. most of the town plan has been realised with the exception of the eastern section, below the haenertsburg primary school. a few houses were built for dpw employees near the r528 but most of the section was not developed as the biotite granite extrusions proved too costly to build on. the rocks have acted as grassland fire refugia and a large colony of the medicinal plants, merwilla plumbea (blue squill) (dzerefos 2004) as well as boophone disticha (bushman poison bulb), occur there. the eastern section is likely to hold the highest concentration of iucn red list category plants in the haenertsburg grassland and was also the area where land was illegally demarcated and sold (troskie 2011). the town plan that appeared on google maps in 2014 for the eastern section was removed by google when they were alerted to the status of the ecosystem. gtm indicated that the existing town plan could only be formally aligned with biodiversity priorities through provincial government intervention, notably the proclamation of the proposed reserve. figure 4: the original town plan of haenertsburg, limpopo province, south africa, as surveyed in 1887. the arrows indicate water drainage lines. the solid line indicates an undeveloped area with important medicinal plants that has been excluded by provincial government from the proposed nature reserve. good news in half-measure notice of the intention to declare 66% of the largest remaining fragment of wgg, a nature reserve, was published in the government gazette no. 2609 (lpg 2015) and no objections were raised by the public. frohg submitted an application to ledet to be designated as the management authority of the nature reserve to be declared, which has been received favourably by ledet. although these are signs of positive development, it seems a half-measure to not declare the full extent of the largest continuous fragment, which is under state tenure and which spatial configuration analysis has identified as the main source area for the other island patches of wgg (niemandt 2015). the inclusion of the contested area (figure 2), which is currently zoned for housing development into the nature reserve, is necessary if ledet is serious about fulfilling the national biodiversity target for wgg because less than 6% of the original extent remains (niemandt 2015). moreover, the status of wgg in the limpopo conservation plan commissioned by ledet suggests that all wgg should be conserved (desmet et al. 2013). discussion ledet has a conservation plan in place (desmet et al. 2013) and has published the intention to declare the wgg nature reserve (lpg 2015). a decade has elapsed since the local environmental group’s submission to have 192 ha of wgg proclaimed a protected area, but only 66% has been included in the proposed reserve (lpg 2015). notably, the unprotected 34%, is the area where illegal land sales had taken place (troskie 2011) and where important medicinal plants are located (dzerefos et al. 2016; sdf 2009); hence, it is the area that is most in need of protection. as two ledet biodiversity specialists recommended a wgg reserve in 2005 and the significance of wgg resources to the poor has been highlighted (dzerefos et al. 2016), senior government officials have no excuse but to recognise the significance of the area unless economic development is being pursued with no regard for socio-environmental issues. this could have been the reason why the illegal squatters (troskie 2011) were not stopped immediately by ledet or gtm until the matter was taken to the public protector and the select committee through sanbi. if the land-use change had not been challenged, an informal settlement could have put the water quality of ebenezer dam at risk, threatened numerous protected and beneficial plants and increased vulnerability of the ecosystem to alien invasive plants and soil erosion. the importance of precedence was illustrated in 2013 when illegal demarcation of plots by the same opportunist was promptly stopped by the saps who recalled the directive from 2011. despite provisions in terms of the promotion of access to information act (no. 2 of 2000), whereby the public can, within reason, request information and thereby hold public officials to account (mfene 2013), government officials simply ignored requests for feedback on minutes of meetings or the reasons for leaving out an important, contiguous section of the proposed wgg reserve. it seems that responding to the public, which has been touted as key to realising a local conservation agenda (cowling & wilhelm-rechmann 2007; swemmer & taljaard 2011), has not been implemented by ledet, reinforcing the general impression that government officials are unreliable and unaccountable for poor service delivery (mfene 2013). nevertheless, it is very encouraging that ledet has begun discussion with the frohg in 2016 to manage the new reserve. official recognition and support of the frohg could result in cost-effective, 24/7 on the ground biodiversity protection. other south african environmental groups have raised objections over the application of environmental legislation (dea 2014). objections follow nema appeals process on environmental authorisations at provincial level or escalate to national level in some instances (figure 5). we perceive that the env-hotline, although operating as a national monitor and providing a reference number, has limited influence as it merely requests feedback from the provincial administration on the allegation of environmental non-compliance. as resolution and feedback to the complainant is not provided by the env-hotline, it appears to have limited impact. however, as complaints are counted and published, these may impact on resource allocation (cer 2014), improved service delivery and appointment of environmental management inspectors (emis) (dea 2014). indeed, in recent years the number of emis in limpopo province has increased substantially more than any other province, from 75 (in 2011/2012) to 237 (in 2013/2014) (dea 2014) and it is likely that implementation, compliance and enforcement of policy and legislation will improve in future. figure 5: a diagrammatic summary of escalating an environmental problem to the three tiers of government (local, provincial and national) to reach a resolution. where matters remain unresolved, a court application could be made, but these could be compromised if the judiciary is not familiar with the complexity of environmental threats posed by the breach of environmental law (moreroa 2014; young 2012). environmental law clinics at universities have undergraduate students requiring experiential learning who can take on cases at reduced cost. alternatively, non-profit organisations, such as the centre for environmental rights, offer affordable advice and assistance for those in financial need. for example, the traditional healers organisation of africa were dissatisfied with how ledet handled damage to a sacred area near thabazimbi and laid criminal charges against the steel mining company’s directors (cer 2015). numerous successful court actions have taken place in recent years such as the kloof conservancy’s landmark victory in the durban high court (case number 12667/2012), which holds all levels of government accountable to the constitution and the law for discharge of environmental duties. monitoring and evaluation of provincial authorities’ outputs and responses to the public should be prompt, transparent and comprehensive so that environmental issues need not be escalated to national level, result in court action or even vandalism. relationships between frohg and other wgg stakeholders have improved over time. discussion and planning have proven better for public relations than mitigating after biodiversity has been lost. most notable is the relationship with lepelle northern water (lnw), which post-2013 were involved with maintenance and excavations (brown & cook 2012; table 1) occurring without an attempt to save threatened or protected plants or plan rehabilitation. in contrast in 2015, lnw appointed msw engineering and project management consultants and srk consulting to design and plan in consultation with frohg, how the existing pipeline running through wgg could be repaired and maintained with minimal environmental damage. the agreed approach was to do more than the regulations called for and to engage a botanist to locate and rescue threatened and protected plant species in the area where excavations were planned. the mitigation actions will set a strong precedent of environmental best practice in threatened ecosystems. unmonitored plant collection in wgg offers an opportunity to engage with traditional healers and hand brush harvesters who earn a living and fulfil an important socio-economic niche in limpopo province (dzerefos et al. 2016). the gauteng directorate of nature conservation has demonstrated how a provincial authority can engage constructively with plant users to benefit conservation (dzerefos & witkowski 2001; taylor & atkinson 2012). an adaptive management approach, such as community-based natural resources management (opio-odongo 2013) which includes a medicinal plant nursery, an environmental education centre and a fire management plan (dzerefos et al. 2016), would contribute to improved environmental awareness and securing water quality of the upper catchment of ebenezer dam. the wgg around haenertsburg could be expanded through the biodiversity stewardship programme (bsp). a co-management agreement between ledet and the owner of the property danallen, which is continuous with the pending nature reserve, could result in a win–win situation, in terms of removal of alien vegetation, resources to manage fire threat and the existing hiking trail. since 2002, bsp has been implemented throughout south africa except for limpopo province (carbutt & martindale 2014). bsp assists land owners adjacent to protected areas to take advantage of the local government municipal property rates act (2004), which states that a municipality may not levy a rate on a special nature reserve as long as it is not developed or used for commercial, business, agricultural or residential purposes. conclusion the identification of areas (mucina et al. 2006; niemandt 2015) that require protection are aligned with national legislation on critically endangered ecosystems (rsa 2011) and the limpopo province conservation plan (desmet et al. 2013). implementation of the conservation plan may enhance connectivity between habitat patches, which is essential for the conservation of viable grasslands (desmet et al. 2013; fourie et al. 2015). effective land-use planning is achievable as good habitat connectivity exists through water drainage lines and firebreaks, which during the wet season are left to grow wild. these stretch from the backbone of the drakensberg escarpment (including the wolkberg wilderness) to the 192 ha of wgg around haenertsburg and a 60 ha wgg peninsular at ebenezer dam. the intent to proclaim the haenertsburg nature reserve, albeit only 126 ha, shows progress by ledet to uphold environmental priorities but more effort is needed in advising gtm and the private sector on the requirements of lema and nema and the importance of wgg. environmental groups, similar to frohg throughout south africa, should see this case study as a success, even though lobbying for wgg is ongoing. this study provides an example of the various avenues that could be followed to help ensure better compliance and implementation of environmental legislation. an understanding of the appropriate legislation that applies to a situation and which tier of government is responsible is essential. detailed record keeping of communications and raising awareness within civil society through the media can also be helpful. as the wheels of government move slowly, it is prudent to prepare for the long haul, to act as a group rather than as individuals and to choose environmental issues to lobby with careful research and specialist input. the intent to proclaim a wgg reserve and negotiations to designate frohg as the management authority of the reserve are positive milestones for a public–private partnership and shows that civil society, scientists and provincial government can work together to achieve important conservation outcomes. furthermore, the inclusion and recognition of environmental volunteer groups brings a diversity of skills and local knowledge to support important ongoing stewardship roles and is highly cost effective as it is based on volunteerism and community service. acknowledgements a postdoctoral research fellowship from the university of the witwatersrand provided financial support. the friends of the haenertsburg grassland are acknowledged for compiling an archive of information dating back to 1986 and pursuing the protection of these grasslands. pieter winter (south african national biodiversity institute), dr ute schwaibold and sylvie kremer-köhne (school of animal, plant and environmental sciences, university of the witwatersrand), advocate jonathan heher, christo rheeders attorneys, the centre for environmental rights and tsholo matlou, department of environmental affairs, gave valued inputs. cilla wildgoose is thanked for assistance with figures 1 and 2. prof. louis changuion, previously in the history department of the university of limpopo, gave useful comments on the final manuscript and sourced the original town plan. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions both authors were responsible for conception and writing while c.m.d. collected and analysed data. references anthony, b.p., scott, p. & antypas, a., 2010, ‘sitting on the fence? policies and practices in managing human-wildlife conflict in limpopo province, south africa’, conservation and society 8(3), 225–240. http://dx.doi.org/10.4103/0972-4923.73812 brown, l.r. & cook, c.l., 2012, ‘preliminary ecological survey for the proposed bulk water pipeline, haenertsburg-polokwane, limpopo province’, royal haskoning dhv, lepelle northern water, unpublished. carbutt, c. & martindale, g., 2014, ‘temperate indigenous grassland gains in south africa: lessons being learned in a developing country’, parks 20(1), 104–125. http://dx.doi.org/10.2305/iucn.ch.2014.parks-20-1.cc.en centre for environmental rights (cer), 2014, numbers matter: join us in reporting number of environmental rights violations, viewed 24 august 2015, from www.cer.org.za/news/numbers-matter-join-us-in-reporting-violations-of-environmental-rights centre for environmental rights (cer), 2015, traditional healers lay criminal charges against mining company and its directors for environmental crimes, viewed 24 august 2015, from www.cer.org.za/news/traditional-healers-lay-criminal-charges-against-mining-company-and-its-directors-for-environmental-crimes changuion, l., 2007, thar’s gold in them thar hills: haenertsburg 1887–1907, review printers, polokwane. convention on biological diversity (cbd), 2014, ‘fifth national report for south africa’, viewed 24 august 2015, from http://www.cbd.int/doc/world/za/za-nr-05-en.pdf cowling, r.m. & wilhelm-rechmann, a., 2007, ‘social assessment as a key to conservation success’, oryx 41, 135–136. department of environmental affairs (dea), 2013, long-term adaptation scenarios flagship research programme (ltas) for south africa, viewed 24 august 2015, from https://www.environment.gov.za/sites/default/files/docs/summary_policymakers_bookv3.pdf department of environmental affairs (dea), 2014, national environmental compliance and enforcement report 2013/14, environmental management inspectorate capacity development and support directorate, viewed 24 august 2015, from https://www.environment.gov.za/sites/default/files/docs/nationalenvironmental_complianceandenforcement_report2013_14.pdf desmet, p. & cowling, r., 2004, ‘using the species–area relationship to set baseline targets for conservation’, ecology and society 9(2), 11, viewed 24 august 2015, from http://www.ecologyandsociety.org/vol9/iss2/art11 desmet, p.g., holness, s., skowno, a. & egan, v.t., 2013, limpopo conservation plan, v.2: technical report, viewed 24 august 2015, from http://bgis.sanbi.org/limpopo/lcpv2_technicalreport_hires.pdf dzerefos, c.m., 2004, ‘today, yesterday and tomorrow: the story of an african grassland’, veld and flora 90(1), 18–19. dzerefos, c.m. & witkowski, e.t.f., 2001, ‘density and potential utilisation of medicinal grassland plants from abe bailey nature reserve, south africa’, biodiversity & conservation 10, 1–22. http://dx.doi.org/10.1023/a:1013177628331 dzerefos, c.m., witkowski, e.t.f. & kremer-köhne, s., 2016, ‘aiming for the biodiversity target with the social welfare arrow: medicinal and other useful plants from a critically endangered grassland ecosystem, limpopo province, south africa’, international journal of sustainable development & world ecology. http://dx.doi.org/10.1080/13504509.2016.1174963 fourie, l., rouget, m. & lötter, m., 2015, ‘landscape connectivity of the grassland biome in mpumalanga, south africa’, austral ecology 40, 67–76. http://dx.doi.org/10.1111/aec.12169 google earth pro v 7.1. (google earth pro), 2014, haenertsburg, limpopo province, 23º56’49” s 29º56’54” e, eye alt 10240 feet, afrigis pty (ltd) cnes astrium 2016, viewed 26 january 2016. low, a.b. & rebelo, a.g. (eds.), 1996, vegetation of south africa, lesotho and swaziland. a companion to the vegetation map of south africa, lesotho and swaziland, department of environmental affairs and tourism, pretoria. limpopo provincial gazette (lpg), 2015, ‘intention to declare attached respective land parcels as protected areas’, 22(2609), viewed 08 march 2016, from http://www.gpwonline.co.za/gazettes/pages/provincial-gazettes-limpopo.aspx?p=4 mfene, p.n., 2013, ‘public accountability: a professional basis for the south african public service’, administratio publica 21(1), 6–23. moreroa, m.p., 2014, ‘the legal implications of grassland as a threatened treasure: analysis of haenertsburg plains in south africa’, msc thesis, department of environmental law and management, university of limpopo. mucina, l. & geldenhuys, c.j., 2006, ‘afrotemperate, subtropical and azonal forests’, in l. mucina & m.c. rutherford (eds.), the vegetation of south africa, lesotho and swaziland, pp. 601–602, strelizia 19, south african biodiversity institute, pretoria. mucina, l., hoare, d.b., lotter, m.c., du preez, p.j., rutherford, m.c., scott-shaw, c.r. et al., 2006, ‘grassland biome’, in l. mucina & m.c. rutherford (eds.), the vegetation of south africa, lesotho and swaziland, pp. 349–412, strelizia 19, south african biodiversity institute, pretoria. niemandt, c., 2015, ‘anthropogenic impacts on the highly threatened woodbush granite grasslands in limpopo, south africa’, msc thesis, department of plant science, university of pretoria. noss, r.f., 1990, ‘indicators for monitoring biodiversity: a hierarchical approach’, conservation biology 4, 355–364. http://dx.doi.org/10.1111/j.1523-1739.1990.tb00309.x novellie, p., biggs, h. & freitag-ronaldson, s., 2013, research and monitoring: interface with legislation, policy and management, sanparks, viewed 24 august 2015, from www.sanparks.org/assets/docs/conservation/reports/2013_research_report.pdf opio-odongo, j., 2013, state of the environment in africa. unep. sustainable development services, viewed 24 august 2015, from http://www.unep.org/pdf/aeo3.pdf republic of south africa (rsa), 2011, national list of ecosystems listed as threatened and in need of protection, gazette no. 34809, notice no. 1002, viewed 09 march 2016, from http://bgis.sanbi.org/ecosystems/summary_%20listed_ecosystems_province.pdf republic of south africa (rsa), 2013, the spatial planning and land use management act, no. 16 of 2013, gazette no. 36730, notice no. 559, viewed 15 march 2016, from http://www.gov.za/sites/www.gov.za/files/36730_5-8_act16of20.pdf rossouw, n. & wiseman, k., 2004, ‘learning from the implementation of environmental public policy instruments after the first ten years of democracy in south africa’, impact assessment and project appraisal 22(2), 131–140. http://dx.doi.org/10.3152/147154604781766012 sango, i. & godwell, n., 2015, ’climate change trends and environmental impacts in the makonde. communal lands, zimbabwe’, south african journal of science 111(7/8). http://dx.doi.org/10.17159/sajs.2015/20140266 seshoka, j., de lange, w. & faysse, n., 2004, ‘the transformation of irrigation boards into water user associations in south africa: case studies of the lower olifants, great letaba and vaalharts water user associations’, international water management institute, viewed 24 august 2015, from http://www.iwmi.cgiar.org/publications/working_papers/working/wor72.pdf spatial development framework (sdf), 2009, greater tzaneen local municipality, viewed 24 august 2015, from www.tzaneen.gov.za/tzaneen/docs/.../spatia%20framework.pdf surveyor-general office, 1887, haenertsburg, houtboschberg, goudvelden, approved by the acting surveyor-general, pretoria, 05 march 1888. swemmer, l.k. & taljaard, s., 2011, ‘sanparks, people and adaptive management: understanding a diverse field of practice during changing times’, koedoe 53(2). http://dx.doi.org/10.4102/koedoe.v53i2.1017 taylor, s.j. & atkinson, d., 2012, ‘delivering community benefits acts as insurance for the survival of small protected areas such as the abe bailey nature reserve, south africa’, koedoe 54(1). http://dx.doi.org/10.4102/koedoe.v54i1.1043 tesfaye, m.a., bravo, f., ruiz-peinado, r., pando, v. & bravo-oviedo, a., 2016, ‘impact of changes in land use, species and ele on soil organic carbon and total nitrogen in ethiopian central highlands’, geoderma 261, 70–79. http://dx.doi.org/10.1016/j.geoderma.2015.06.022 thompson, d.i. & swemmer, a.m., 2014, ‘are grasslands doomed under global change? triennial burning cannot prevent shrubland transformation of mid-altitude south african grasslands’, scripta botanica belgica 52, 437. troskie, a., 2011, ‘squatters threaten scarce ecosystem’, letaba herald 40(36), 1. wilhelm-rechmann, a. & cowling, r.m., 2013, ‘local land use planning and the role of conservation: an example analyzing opportunities’, south african journal of science, viewed 24 august 2015, from http://www.sajs.co.za/sites/default/files/publications/pdf/willhelm-rechmannl_research%20art.pdf young, m., 2012, ‘are we greening our court rooms? an analysis of the recent decisions, in magaliesburg protection association v mec: department of agriculture, conservation, environment and rural development’, south african journal of environmental law and law policy 19(1), 63–73. filelist convert a pdf file! supplement to koedoe. 1977: 181-185. is there a place for nature conservation in the transkei? m m ntloko director of forestry and nature conservation dept. of forestry and agriculture private bag x5002 umtata republic of transkei the republic of the transkei is situated in the south-east of the republic of south africa and is bounded by the ..cape province, natal, lesotho and the indian ocean. the area of the country is some 43 200 km 2 , almost as large as denmark and twice the size of wales. it is a country made up of mountainous regions along the lesotho border and the rest is made up mostly of rolling hills with deep river valleys. it is a summer rainfall region ranging between 500 mm and 1 300 mm per year and thus the country may be termed well watered. climatic conditions may be described as temperate with occasional frosts during the worst winter months. according to statistics, communal pastorage covers over 70% of the transkei area, arable land 24%, with forest plantations and residential areas 3% each. the 1971 population census revealed population figures as follows: 1 650 825 blacks, 9 556 whites and 7 655 coloured. the people are mostly pastoral farmers with maize, whilst sorghum crops playa lesser but important role in the daily needs of the people. u mtata is the administrative capital of the transkei where all the government departments are situated. in passing it is interesting to note that the name u mtata is derived from the sneezewood (ptaeroxylon obliquum) tree which in xhosa is named the umthathi tree. according to acocks (1953) the transkei is divided roughly into six veld types each with conservation and/or recreational potential. these veld types are: 1. pondoland coastal plateau sourveld 2. coastal forest 3. mangrove woodlands 4. n gongoni veld 5. mountain forests of the highland sourveld and 6. valley bushveld 181 pondoland coastal plateau sourveld this area starts north of port st. johns and co-incides with the outcrop of table mountain sandstone and takes in the districts of lusikisiki, flagstaff and bizana. the forests in this area are characterised by the following trees: strelitzia nicolai, (wild banana), croton sylvaticus and ficus natalensis while the sourveld which is the densest veld in the republic, is so dense that the grasses grow as single shoots rather than as tufts, and is characterised by them eda triandra, tristachya hispida and diheteropogon filifolius . coastal forest coastal forests occur along most of the transkei coast and good examples being the manubi, dwesa and egoso forests. these forests are characterised by trees such as heywoodia lucens, milletia grandis and vepris undulata . mangrove woodlands mangrove forests occur on many rivers which have suitable muddy flats. exceptionally good examples are found at the mouths of the mtafufu, umgazana and umtata rivers. the three species of mangroves that occur are avicennia marina, bruguiera gymnorhiza and rhizophora mucronata. ngongoni veld this veld type occupies a narrow and irregular belt of rolling country just above coastal forest belt, and is characterised by aristida junciformis. mountain forests of the highland sourveld these forests have been slowly eroded and now consist of very small scattered examples with the best being seen near umtata and at langeni, mhlahlane, baziya and also at gomo near mount ayliff. valley bushveld this veld type is found in most of the southern river valleys and a particularly good example is found along the kei river. due to the topography and abundant rainfall the transkei is well endowed with water-rich rivers, most of which have their catchment areas within the borders of the transkei. the rivers were stocked with bass, carp and trout in the past by early settlers. all the species have done well and trout seem to have established themselves in the upper reaches of most of the larger rivers. the traditional way oflife has both been beneficial and detrimental to the fauna and flora of the country. the beliefs of the people have helped 182 in the protection of certain species of trees, birds and animals as well as the forests, the latter being areas of escape during faction fights. the traditional ways of farming have been detrimental to the soil as each farmer wants to be able to supply his household with grain and this often leads to them ploughing areas on steep gradients which are not suitable. having the community spread over large areas brings the people and more especially the herd boys with their dogs into daily contact with what wildlife there is in the area, and all their leisure time is spent hunting. the fuel needs of the people take a heavy toll on the forests and where no trees are available even dung is used for fires which means nothing goes back into the soil. the constant need for grazing has led to overburning and this in turn has led to soil erosion. the department of agriculture is busy with the planning oflocations which consist of a living area as well as indicating where ploughing, grazing and other activities may be undertaken while other areas are left where wildlife can re-establish itself again. the needs of the people are brought closer to their homes e.g. woodlots are planted in these areas as fuel for the population. the transkei has its own nature conservation act (act no.6 of 1971) and through this the division of nature conservation has come into being. the division is situated within the department of forestry which in turn is part of the larger department of agriculture and forestry. its present staff is very small and consists of four nature conservation officers, one senior nature conservation officer and one professional officer, but will soon be expanded to include eight professional officers as well as a number of nature conservation officers posts that will be filled from students completing the nature conservation course at cwaka college in zululand . in addition to these posts the forestry department have many forest guards posts who help in the administration of the nature conservation act. to ensure that all aspects of nature conservation are properly researched, a conservation advisory board has been formed to advise the transkei government. it is hoped that by the formation of this board any projects which can have an effect on the environment will be properly researched at the vital planning stage. all reserves will be planned by suitably qualified ecological planners as well as having management plans drawn up by qualified persons. following six nature reserves are in different stages of proclamation, planning and development: 1. dwesa nature reserve 2. hluleka nature reserve 3. mount thesiger nature reserve 4. mkambati nature reserve 5. swartberg nature reserve and the 6. nduli nature reserve 183 dwesa nature reserve dwesa was proclaimed in 1974 and consists of an area of 5 000 ha and is divided roughly into one-third of rolling grassveld, and two-thirds of indigenous coastal forest. the tourist facilities have been planned by a firm of ecological/landscape planners and the first bungalows are at present being erected. buffalo, eland and blesbok have been reintroduced to supplement the bushbuck and blue duiker, grey duiker, genets, mongeese and vervet and samango monkeys already occurring there. it is hoped that in the near future that one of the conservation departments who have hippo under their control will be able to let this department have a few as there are three very large estuaries ideal for hippo the last hippo being shot in this area in the early 1890's. other animals that used to occur there are reedbuck, zebra and hartebeest, all which seem to be very difficult to obtain at present. an easy way of locating dwesa on a map is to find the bashee river and the reserve is situated to the south of it. hluleka nature reserve hluleka is an area of i 000 ha and situated about 40 km south of port st. johns and was proclaimed during 1976. it is at present being planned by an ecological/landscape firm and is being fenced at the moment. mount thesiger nature reserve this area is situated directly inland from port st. johns and is at present under the control of the forestry department of the republic of south africa. on the eve of independence this area will be proclaimed a nature reserve. there are steep rises from the coast and it consists of forest and sour grassland interspersed with heaths and proteas. mkambati nature reserve this area of 7000 ha is situated between the mtentu and msikaba rivers. it can be classified under the coastal sourveld type and the very rarejubaeopsis caffra communities occur within its borders. this area will be proclaimed as a reserve in the very near future. swartberg nature reserve this reserve of 9000 ha is situated on the borders of the transkei and natal and is well within the snowline as it lies 2 000 m above sea level. in the summer large numbers of waterfowl descend on vleis left behind by the melting snow. the estate of a single part owner is all that is withholding the proclamation of this reserve. 184 nduli nature reserve nduli is an existing reserve at u mtata which is being administered by the local municipality and will be taken over and enlarged to include a national botanical garden on th e eve of independence . in addition the division is constantly on th e lookout for viable reserves within the vegetation types as well as th e establishment of a mangrove reserve which is a priority. while we are discussing reserves i would like to mention that the traditional erection of fences you stay on your side and we stay on our side attitude does not work in the transkei. we in the transkei make it a point of involving the people around the reserve in as many of the facets of the day to day running of the reserve as possible. we draw all our labour from the surrounding location and this includes the staff in the tourist camps. in conjunction with the department of agriculture we plan the settlement areas in such as way as to be able to draw our labour from them . we also encourage market gardening and traditional home crafts amongst the people so that they can sell these directly to the tourists . in addition regular get-togethers are held so that every person on the reserve and around it knows exactly what is being done. this tends to minimise the problems usually encountered with other but similar land use projects in the transkei. extension work is hitherto confined to a weekly 30 minutes radio program, lectures to the agricultural college at tsolo and schools in and around umtata as well as schools in the vicinity of the reserves . a start was made during 1976 on introducing a course for biology teachers at dwesa and this seemed to be a success. however, it is difficult to gauge this success as for some reason or other, there is no follow up to questionnaires sent to these teachers after the course. even though we are very limited by staff at the moment we have a large bass breeding station at tsolo, a trout hatchery at mhlahlane (26 km from umtata) and a carp breeding station at tsomo which will be moved to tsolo in the very n ear future . to take the pressure off our cycads we have a nursery combined with the trout hatchery from which at least 600 seedlings were sold during 1976 most of these being bought by visiting officials from pretoria! the question placed before me was "whether there is a place for nature conservation in the transkei" to which i reply yes . there is a tremendous need for nature conservation in the transkei to preserve its natural assets and to boost its national economy through tourism. reference acocks , j p h 1953. veld types of south africa. mem. bot . surv . s. afr. 28. 185 page 1 page 2 page 3 page 4 page 5 article information authors: robert f. brand1 nacelle collins2 p. johann du preez3 affiliations: 1applied behavioural ecology & ecosystem research unit, university of south africa, south africa 2free state department of economic development, tourism and environmental affairs, free state, south africa 3department of plant sciences, university of the free state, south africa correspondence to: robert brand email: rbrand@mweb.co.za postal address: private bag x6, florida 1710, south africa dates: received: 15 apr. 2014 accepted: 10 sept. 2014 published: 16 apr. 2015 how to cite this article: brand, r.f., collins, n. & du preez, p.j., 2015, ‘a phytosociology survey and vegetation description of inselbergs in the ukhahlamba-drakensberg park world heritage site, south africa’, koedoe 57(1), art. #1233, 12 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1233 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/koedoe.v57i1.1233-1. copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. a phytosociology survey and vegetation description of inselbergs in the ukhahlamba-drakensberg park world heritage site, south africa in this short communication... open access • abstract • introduction • study area • materials and methods    • data collection    • data processing    • classification    • naming of plant communities    • gradient analysis • results    • species composition    • gradient analysis    • describing the plant communities • discussion    • classification       • major community 1, rhodohypoxis rubella, wetland grass and forblands       • major community 2, crassula peploides – polevansia rigida, sheet rock grass and forblands       • major community 3, pentaschistis exserta – merxmuellera stricta, high-altitude alpine grassland       • major community 4, merxmuellera drakensbergensis – helichrysum trilineatum, high-altitude alpine fynbos grassland    • syntaxonomical classification • conservation and climate change • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ no previous scientific surveys have been conducted on inselbergs in the drakensberg. the aim of this study was to collect specimens, identify, describe and name the vegetation clusters and assess biogeographical connections with other afromontane regions. a total of 103 relevés where sampled from six inselbergs. the plant sampling was carried out according to the braun-blanquet method with the plant and environmental data entered in turboveg and exported as a cornell condensed format file (cc!) into juice. classification was completed using twinspan (two-way indicator species analysis) (modified), resulting in 4 major communities, 11 communities, 13 sub-communities and 18 variants. ordination (indirect) was carried out using canoco (version 4.5) to investigate the relationship between species. the four major communities identified are rhodohypoxis rubella (wetland grass and forblands), scirpus ficinioides – crassula peploides (sheet rock grass and forblands), pentaschistis exserta (high-altitude alpine grassland), previously undescribed, and merxmuellera drakensbergensis – helichrysum trilineatum (high-altitude alpine fynbos grassland), described in other vegetation and floristic studies. four habitats were identified, namely wetlands, sheet rock shallow soil, high-altitude alpine grassland and deep soil high-altitude fynbos grasslands. substrate and moisture availability appeared to be the defining micro-climatic conditions determining the different vegetation clusters whilst altitude is the overriding environmental factor influencing all vegetation. conservation implications: rising temperatures as a result of carbon dioxide increase is predicted to drastically decrease the number of endemic and near-endemic montane species, whilst altering the composition of vegetation units which comprise the alpine vegetation. introduction top ↑ studies of high-altitude inselbergs within the east and southern afromontane region are few. whilst studies of inselbergs in africa have been concentrated more in west africa (barthlott, gröger & porembski 1993; parmentier et al. 2006; porembski & brown 1995; porembski et al. 1996, 1997; seine et al. 1998), several have been undertaken in namibia (burke 2001, 2002, 2004) and south africa, such as with the korannaberg (du preez 1991) and platberg, at 2394.5 m, some 60 km north of the drakensberg (brand, du preez & brown 2010). other inselberg surveys have been conducted in moçambique (van noort, gardiner & tolley 2007) which document the high biological diversity and establishe inselbergs as sites of high endemism. many are denoted as biodiversity ‘hotspots’ (van noort et al. 2007), which also represent relictual plant populations of past climate conditions. climate change and global warming, with its rising carbon dioxide levels, will have a profound impact on the grassland biome and the structure of c3 and c4 grasses (mucina & rutherford 2006), in particular the alpine grasses embedded in the grassland biome. inselbergs have also been assessed for their high conservation value and biotic diversity (ed. huntley 1989). this article is the first phytosociological survey of the high-altitude inselbergs in the drakensberg. its aims are to (1) classify the inselberg alpine vegetation into communities and present the species composition, (2) examine environmental gradients to identify habitats and (3) identify and examine biogeographical affinities with previously described or identified plant communities. this new knowledge will contribute to the understanding and conservation management of high-altitude alpine vegetation and species unique to the drakensberg. study area top ↑ the study sites are located in the drakensberg, south africa, and consist of six inselbergs, all over 3000 m in altitude (figure 1). the most northerly, sentinel, is located at 28°44’25.97” s and 28°53’27.42” e, with four inselbergs – eastern buttress, inner horn, outer horn and dragon's back – sequentially between it and the most southerly of the inselbergs surveyed, namely cathkin peak at 29°4’35.03” s and 29°20’59.38” e. all six peaks fall within the kwazulu-natal ukhahlamba-drakensberg park, a world heritage site and part of the drakensberg alpine centre (dac) biodiversity hotspot. all inselbergs with the exception of sentinel are relatively flat-topped, with boulders, shallow soils occurring on sheet rock, gravel plains, seeps, interconnected seasonally inundated wetlands and ephemeral pools (figures 2 and 3). they are all sheer sided, separated by varying distances from the main escarpment by cliffs of 1000 m or more, and form an archipelago of island-like peaks stretching some 60 km north to south. figure 1: maps of study sites: (a) study area with kwazulu-natal ukhahlamba-drakensberg park (shaded) and (b) south africa indicating drakensberg alpine centre. figure 2: photographs of inselberg vegetation in the drakensberg: (a) typical inselberg vegetation of low, cushion-forming fynbos shrubs and bunch grass on outer buttress, with the sloping peak of sentinel in the background, (b) the flat top of inner buttress showing three of the four habitats: sheet rock outcrops/gravel plains, alpine grasslands and deep soil, helichrysum/erica fynbos grassland. devils tooth in the foreground, (c) tall, montane grass community: merxmuellera drakensbergensis, erica dominans and the yellow-flowering moraea alticola, (d) deep soil habitat of the yellow-flowering helichrysum trilineatum, erica species and passerina drakensbergensis with typical caespitose habitat, indicative of alpine conditions, (e) wetland habitat, with aponogeton ranunculiflorus the drakensberg alpine centre endemic, hydrophyte, an ephemeral-pool specialistn and (f) mixed habitats on cathkin of wetlands, gravel seeps, sheet rock shallow soils, embedded in rocky terrain with alpine grasslands and the helichrysum/erica fynbos. figure 3: uncommon vernal pools on cathkin and outer buttress, habitat for specialist aquatic plants aponogeton junceus, crassula dependens, crassula gemmifera, limosella grandiflora and limosella vesiculosa. materials and methods top ↑ data collection a total of 103 plots where sampled on the six inselbergs. plot sizes varied according to substrate, and were subjectively estimated at either 3 m x 3 m for sheet rock, seeps or gravel plains, or 6 m x 5 m for grass or dwarf shrub vegetation as per theoretical criteria (westhoff & van der maarel 1980) and established field practice in south africa (brown et al. 2013). in all sample plots each species was recorded, all plants counted and cover estimated using the modified braun-blanquet cover/abundance scale: r, +, 1, 2a, 2b, 3, 4, 5 (mueller-dombois & ellenberg 1974; whitaker 1980). selection and sampling of vegetation was carried out where a visible difference could be seen in vegetation or where it occurred in clearly different habitats such as rocky outcrops, wetlands, sheet rock or seeps. fieldwork was carried out in november 2005, with initial identification of known plants carried out in the field and later conducted in the donald killick kzn herbarium (cfr) and the geo potts herbarium (blfu) at the university of the free state. more problematic material was identified at the south african national biodiversity institute herbarium (pre) in pretoria. plant species nomenclature was according to germishuizen et al. (2006), and updated with the november 2009 precis database at the south african national biodiversity institute (sanbi), pretoria. habitat as well as floristic data was captured using turboveg (hennekens & schaminée 2001), with the subsequent relevés generated exported as a cornell condensed format file (cc!) into juice version 7.0.28 (tichý & holt 2006). data processing a first approximation at clustering was carried out using the twinspan (two-way indicator species analysis) algorithm of hill (1979) which is incorporated as part of the juice program. to produce the syntaxonomic table using juice, separators were defined at six hierarchical levels, with group size standardised. using juice version 7.0.28 (tichý & holt 2006), the diagnostic species were identified by applying a statistical fidelity measurement which is useful for assessment of species concentration in vegetation units, and for comparing diagnostic values amongst species in a particular vegetation unit, or amongst vegetation units for a particular species (tichý & holt 2006). fidelity values were calculated using the phi coefficient, which considers only presence/absence data, so that the fidelity values are not influenced by the cover/abundance values of the species (lepš & hadincová 1992). the fisher's exact test was employed along with the phi coefficient fidelity measure to calculate the true probability of obtaining the observed number of occurrences of the species in the vegetation unit under the null hypothesis of independence. using the fisher's exact test along with the phi coefficient measure of fidelity causes values that are not statistically significant at the predefined p-value to be assigned a fidelity value of 0, where p was chosen to be < 0.001. species are therefore only identified as diagnostic if their fidelity values exceed the subjectively chosen lower threshold. the braun-blanquet normal scale was used, and a combination of frequency, fidelity and cover was selected, using the default settings of 67% frequency and 45.3% fidelity. despite the subjectivity and inaccuracy of the braun-blanquet method and the use of non-numerical scores ‘r’ and ‘+’, which pose computation problems discussed in detail by podani (2006), this method of field data collection was used to conform with and make this survey's data compatible with the thousands of relevés already sampled in south africa. the continued use of the braun-blanquet method in south africa is also suggested by brown et al. (2013). classification within juice the lower threshold values for the diagnostic, constant and dominant species when applying the ‘analysis of columns of syntaxonomic tables’ (tichý & holt 2006) function were set to 70, 60 and 50 respectively, whilst the upper threshold values were set to 80, 70 and 60 respectively. species that exceed the lower threshold are listed whilst those that exceed the upper threshold are printed in bold. naming of plant communities the naming of plant communities was carried out according to guidelines suggested by brown et al. (2013). the syntaxonomic names for the major communities, communities, sub-communities and variants were derived according to diagnostic, dominant and constant species obtained from floristic and environmental data processed in juice (tichý & holt 2006). gradient analysis the skewness and kurtosis calculations performed with pc-ord version 5.0 revealed the non-unimodal distribution of the species data (also confirmed by the disjunct nature of the dataset as indicated by the dca eigenvalue of one for the first axis) (gauch 1982a, 1982b). to achieve a normal distribution the species data were log-transformed during ordination (legendre & legendre 1998). a final manipulation of relevé columns and species rows was carried out in juice to fine-tune the phytosociological table, which was exported into excel and refined for presentation by moving rows containing species and adding alphabetic letters to denote species groups (online appendix 1). groups of similar ecological characteristics were identified and related to environmental gradients. the relationship of the identified plant communities with the environmental variables is presented in figures 4 and 5. figure 4: correspondence analysis of all relevés showing major communities 1, 2, 3 and 4. figure 5: correspondence analysis with wetlands (group 1) removed to show more clearly the gradients typifying groups 2, 3 and 4. results top ↑ species composition a total of 103 relevés and 189 species where incorporated in the classification; 2.6% of these were pteridophytes, 31% were monocotyledons and 66% were dicotyledons. there are 79 species that occur in ≥ 4 relevés and which do not form appreciable clusters, and have been left out of the formal phytosociological description. however, all species are included in online appendix 1. the average number of species per plot is 14.16 (s.d. 6.56), minimum 4, maximum 32. there are seven sub-communities and variants with the same numbers of relevés, diagnostic, constant and dominant species. accordingly, only the sub-communities have been described and named whilst ignoring the variants. there are 18 variants which have different diagnostic, constant and dominant species, which have been named. subsequently, there are 13 unique sub-communities, 18 variants, 11 communities and four major communities which form the syntaxonomical classification presented in online appendix 1. gradient analysis twinspan produced four major communities representative of four different habitats (figures 4 and 5). these are: major community 1: wetland grass and forblands. major community 2: sheet rock grass and forblands. major community 3: high-altitude alpine grassland. major community 4: high-altitude alpine fynbos grassland. figure 4 shows outliers representing 11 relevés, all of which comprise major community 1, wetlands. relevés 94, 91, 92 and 90 have the highest values and form major community 3, which is the wettest of all habitats and is comprised of obligate wetland species: aster erucifolius, crassula dependens, crassula gemmifera, limosella grandiflora, limosella vesiculosa, pentaschistis airoides and rhodohypoxis rubella (online appendix 1), which includes aponogeton junceus found in deep, water-filled pools. relevés 101, 102 and 103 represent major community 2, a mix of forbs favouring moist, shallow, wet rock or gravel areas, whilst relevés 97, 98, 99 and 100, which constitute major community 1, have the dominant grass pentaschistis airoides and the wetland forb limosella vesiculosa, which is found in ephemeral or temporary wetlands and can withstand desiccation, high salinity and alkalinity and extreme temperatures of freezing and thawing (heilmeier et al. 2005). major community 1 ephemeral wetlands possibly dry up during june, july and august, the 3 driest months in the alpine region of the drakensberg (mucina & rutherford 2006). major community 1 wetlands represent a gradation of wetland habitats, from deep pools (community 1; species group b, online appendix 1) to seasonal, shallow rocky seeps (community 2; species group c, online appendix 1) to semi-permanent or ephemeral pools (community 3; species group d, online appendix 1). limosella grandiflora (species group d, online appendix 1) is an angiosperm, poikilohydric plant indicative of vernal or ephemeral pools (figure 3), similar to inselbergs in namibia and zimbabwe (heilmeier et al. 2005; mokuku 1991) and described by hillard and burtt (1987) in the northern drakensberg. in figure 4, the tight clustering of relevés 79–61 constitutes major community 2, and forms a decreasing gradient overlaid by relevés 81 through 96 to 61. the habitat of sheet rock shallow soils is confirmed by the dominant species scirpus ficinioides, a facultative wetland species (marneweck & kotze 1999), and the diagnostic species crassula peploides. grasses forming major community 2 are the two facultative wetland species festuca caprina and koeleria capensis (marneweck & kotze 1999). additionally, within major community 2, the three grasses favouring seeps or seasonally inundated wetland habitats are pentaschistis exserta, polevansia rigida and styppeiochloa gynoglossa (gibbs-russell et al. 1991). these sheet rock shallow soil habitats are found on all inselbergs, as shown in figures 4 and 5, where group 2 represents major community 2, group 3 represents major community 3 (high-altitude alpine grasses) and group 4 represents major community 4 (high-altitude fynbos grasslands). the results from figures 4 and 5 show habitat and plant affinities associated throughout the six inselbergs whilst also showing separation into discrete clusters centred on each inselberg. describing the plant communities all clusters are named in the discussion and fully listed in the online appendix 1. major community 1, rhodohypoxis rubella (wetland grass and forblands), which consists of 11 relevés, contains nine species (five monocots, four dicots) of which eight are endemic or near-endemic, high-latitude afromontane/dac species with rhodohypoxis rubella the diagnostic species (75.8% abundance) as well as providing 100% constancy. it is comprised of three communities. community 1 has the diagnostic species limosella vesiculosa (100%) and the constant species pentaschistis airoides and rhodohypoxis rubella. community 2 has no diagnostic species whilst constant species are aster erucifolius, crassula dependens (the only non-endemic dac taxon), rhodohypoxis rubella and the shrub clutia nana (67% constancy). community 3 has the diagnostic species aponogeton junceus (86.2%) and limosella grandiflora (80.9%), with constant species aponogeton junceus and crassula gemmifera (75%) and limosella grandiflora and rhodohypoxis rubella (100%). all species provide limited cover/abundance with the shallow, seasonally inundated wetlands on gravel or sheet rock outcrop substrate. major community 2, scirpus ficinioides – crassula peploides (sheet rock grass and forblands) is comprised of two communities, five sub-communities and eight variants. this major community has limited cover/abundance from all species with the diagnostic species crassula peploides present in 54% of 28 relevés. the vegetation is comprised of 61% monocots (18 species) and 13 dicots of which 3 are crassulaceae (online appendix 1). the sedge scirpus ficinioides is the dominant species. the two communities are community 2.1 (22 relevés) with scirpus ficinioides the dominant species (71% presence), and community 2.2 (5 relevés) which has no dominant species. the diagnostic species is ficinia cinnamomea (88.8% presence), with constant species clutia nana, felicia linearis, and oxalis obliquifolia. species composition and habitat may suggest that major community 2 represents a gradient of vegetation types in transition between wetland habitat (major community 1) and alpine grassland (major community 3). sub-community 2.1.1, crassula peploides – rhodohypoxis rubella (wet grassland), is composed of two facultative wetland species. it has three variants (online appendix 1), is comprised of 9 relevés, and has 78% constancy of both wetland species. it has limited cover/abundance of all species and consists of scattered, low plants including the alpine grasses polevansia rigida, pentaschistis exserta, koeleria capensis (variant 2.1.1.1) and pentaschistis airoides (variant 2.1.1.3). forbs and geophytes include endemic and near-endemic dac taxa (online appendix 1). sub-community 2.1.2, scirpus ficinioides – styppeiochloa gynoglossa (alpine grassland), consists of five relevés, dominated by the sedge scirpus ficinioides and defined by the dac-endemic grass styppeiochloa gynoglossa. in comparison with the previous sub-community it has a reduced presence of wetland species (online appendix 1), with variant 2.1.2.2 defined by the endemic forbs glumicalyx montanus, and variant 2.1.2.3 defined by helichrysum pagophilum. sub-community 2.1.3, erica dominans – polevansia rigida (erica alpine grassland), consists of eight relevés and has three variants (online appendix 1), with erica dominans having 88% consistency, and the first appearance of any erica species. there are only two wetland species, crassula peploides and rhodohypoxis baurii, both of which have low presence. three additional medium to tall alpine bunch-grasses, festuca caprina (variant 2.1.3.2), eragrostis caesia and merxmuellera drakensbergensis (variant 2.1.3.3), occur. sub-community 2.2.1, clutia nana – hesperantha baurii (forbland), with three relevés, and sub-community 2.2.2, erica algida – erica glaphyra (heathland), with two relevés, contain only 16 species, 12 of which comprise the phytosociological classification. the graminoids, forbs and geophytes are small, low species, including the two erica species erica algida (dac near-endemic) and erica glaphyra (dac endemic). major community 3 has three communities, two sub-communities and no variants. it is named the pentaschistis exserta – merxmuellera stricta (high-altitude alpine grassland), defined and dominated by pentaschistis exserta, and includes pentaschistis airoides (community 3.1), cyperus schlechteri, festuca caprina and merxmuellera stricta (community 3.2). community 3.3 is dominated and defined by ericaceous fynbos shrubs erica dominans, passerina montana, cliffortia nitidula and helichrysum trilineatum, with the alpine grass merxmuellera stricta having limited cover/abundance. there are a total of 45 species of which 35 form the syntaxonomical classification; 11 of those are endemic and 10 near-endemic dac taxa. of the six grasses, one is endemic (pentaschistis exserta), three are near-endemic (eragrostis caesia, pentaschistis airoides and pentaschistis basutorum), with festuca caprina and merxmuellera stricta afromontane species. all species are montane, using c3 metabolism, with only eragrostis caesia (near-endemic) using the c4 pathway. of the three sedges, cyperus schlechteri and schoenoxiphium schweickerdtii are near-endemic. of the 14 forbs, 6 are endemic, 4 near-endemic and 4 widespread. of the seven geophytes, four are endemic (eucomis schijffii, moraea alticola, ornithogalum sephtonii and sebaea spathulata). of the five shrubs, one is endemic (erica dominans), two are near-endemic (helichrysum trilineatum, macowania sororis) and the other two are montane fynbos. major community 4, merxmuellera drakensbergensis – helichrysum trilineatum (high-altitude alpine fynbos grassland), is the most widespread vegetation type and comprises 46% of the vegetation found on all six inselbergs. there are three diagnostic species: the sedge scirpus falsus, the shrub helichrysum trilineatum (the tallest plant on the inselbergs) and the montane bunch grass merxmuellera drakensbergensis. constant species include helichrysum trilineatum, scirpus falsus, merxmuellera drakensbergensis and schoenoxiphium schweickerdtii. dominant species are erica dominans, erica glaphyra, merxmuellera disticha, merxmuellera drakensbergensis, passerina drakensbergensis and scirpus falsus. major community 4 has three communities, five sub-communities and eight variants (online appendix 1). ten erica species were identified: erica aestiva, erica dissimulans, erica dominans, erica flanaganii, erica glaphyra, erica thodei, all endemic to the drakensberg, erica algida and erica alopecurus, both near-endemic, erica maesta and erica reenensis, both montane species also occurring in the eastern cape (carbutt & edwards 2006). however, only erica dominans and erica glaphyra occurred with significant presence (±45% – 49% respectively), with erica thodei providing 29% presence and 69% constancy (online appendix 1). excluding erica, a total of 19 other fynbos shrubs or dwarf shrub species were recorded, of which helichrysum trilineatum (81.6% cover/abundance) is the dominant shrub, growing up to 2.5 m tall on cathkin peak. other fynbos shrubs include euryops decumbens (16%), erica montanus, macowania glandulosa, macowania sororis (asteraceae), muraltia alticola, muraltia flanaganii, muraltia montana (22.4%), polygala hottentotta (polygalaceae), cliffortia browniana, cliffortia filicaulis, cliffortia filicauloides, cliffortia nitidula (rosaceae), anthospermum basuticum (55.5%), anthospermum hispidulum, anthospermum monticola (rubiaceae), gnidia aberrans, passerina drakensbergensis (16%) and passerina montana (thymelaceae). the following three communities comprise major community 4: community 4.1, scirpus falsus – anthospermum basuticum (fynbos shrubland), with 11 relevés, two sub-communities and two variants, community 4.2, erica glaphyra – helichrysum trilineatum (erica alpine grassland), with 24 relevés, two sub-communities and eight variants, and community 4.3, merxmuellera drakensbergensis – schoenoxiphium schweickerdtii (alpine grassland), with 14 relevés and two sub-communities (online appendix 1, species group r). community 4.1, scirpus falsus – anthospermum basuticum (fynbos shrubland), has the sedge scirpus falsus as the dominant species and the dac-endemic shrub anthospermum basuticum as the defining species. it has two sub-communities: sub-community 4.1.1, macowania glandulosa – disa crassicornis (fynbos shrubland) and sub-community 4.1.2, scirpus falsus – anthospermum basuticum, which is defined by a high cover/abundance of the common sedge scirpus falsus and has a lack of a unique species clustering. community 4.2, erica glaphyra – helichrysum trilineatum (alpine grassland), is the largest cluster (22 relevés) with 90% presence of erica glaphyra, which is also the dominant species. it is defined by helichrysum trilineatum (82% constancy) and represents the widespread, dominant vegetation type on all inselbergs. it has two sub-communities: sub-community 4.2.1, schoenoxiphium filiforme – euryops montanus (fynbos), with three variants, and sub-community 4.2.2, erica dominans – schoenoxiphium schweickerdtii (heathland), with five variants. sub-community 4.2.1 is dominated by the endemic sedge schoenoxiphium filiforme, and the cluster is defined by euryops montanus, a dac endemic. erica glaphyra has 100% constancy and high cover/abundance values associated with this sub-community in contrast with sub-community 4.2.2., which is dominated by erica dominans (96% constancy) and a concomitant reduced presence of erica glaphyra. this mutual exclusion is also seen with the two sedges, schoenoxiphium filiforme (67% constancy in sub-community 4.2.1 and 0% in sub-community 4.2.2) and schoenoxiphium schweickerdtii (44% constancy in sub-community 4.2.1 and 90% in sub-community 4.2.2). the last community to the right on the syntaxonomic table (online appendix 1), community 4.3, merxmuellera drakensbergensis – schoenoxiphium schweickerdtii (alpine grassland), is dominated by the tall, rhizomatous, endemic grass merxmuellera drakensbergensis with 100% constancy and cover/abundance of 30% – > 70% of relevés. it is so dominant that it has almost totally excluded all other alpine grasses, fynbos families, for example ericaceae, and the tall fynbos shrubs, for example passerina, cliffortia, euryops and helichrysum trilineatum. this could be a combination of two factors; it is found on deep soil and is a large, tufted grass which would out-compete smaller species. some plants were found to be present throughout many of the relevés in major community 4, but with limited cover/abundance. they include six forbs, namely othonna burttii, scabiosa columbaria, craterocapsa tarsodes, helichrysum infaustum, helichrysum bellidiastrum and alepidea pusilla. other widespread forbs occurring with limited cover/abundance found in major communities 2, 3 and 4 include rhodohypoxis baurii (absent from major community 1), psammotropha obtusa and oxalis obliquifolia. the facultative wetland species rhodohypoxis rubella, which dominates and defines major community 1, also occurs scattered throughout other relevés in major communities 2 and 4. discussion top ↑ classification the results from the phytosociological analysis indicate the inselberg vegetation has four major communities, 11 communities, 13 sub-communities and 18 variants (online appendix 1). overlap exists with sub-communities and variants. the name designated for each cluster is for the highest order unit, whilst ignoring those of lower rank (online appendix 1). the names are: major community 1, rhodohypoxis rubella, wetland grass and forblands community 1.1, limosella vesiculosa – pentaschistis airoides, wetland community 1.2, crassula dependens – aster erucifolius, wetland community 1.3, limosella grandiflora – aponogeton junceus, wetland major community 2, crassula peploides – polevansia rigida, sheet rock grass and forblands community 2.1, scirpus ficinioides – pentaschistis exserta, grassland sub-community 2.1.1, crassula peploides – rhodohypoxis rubella, wet grassland variant 2.1.1.1 koeleria capensis variant 2.1.1.2 eucomis schijffii variant 2.1.1.3 pentaschistis airoides sub-community 2.1.2, scirpus ficinioides – styppeiochloa gynoglossa, alpine grassland variant 2.1.2.1 glumicalyx montanus variant 2.1.2.2 helichrysum pagophilum sub-community 2.1.3, erica dominans – polevansia rigida, alpine grassland variant 2.1.3.1 clutia nana variant 2.1.3.2 festuca caprina variant 2.1.3.3 styppeiochloa gynoglossa community 2.2, ficinia cinnamomea – felicia linearis, heathland sub-community 2.2.1, clutia nana – hesperantha baurii, forbland sub-community 2.2.2, erica algida – erica glaphyra, heathland major community 3, pentaschistis exserta – merxmuellera stricta, high-altitude alpine grassland community 3.1, pentaschistis airoides – helichrysum pagophilum, alpine grassland community 3.2, festuca caprina – erica dominans, alpine grassland sub-community 3.2.1, cyperus schlechteri – glumicalyx goseloides, alpine grassland sub-community 3.2.2, pentaschistis basutorum – hebenstretia dura, alpine grassland community 3.3, passerina montana – cliffortia nitidula, fynbos shrubland major community 4, merxmuellera drakensbergensis – helichrysum trilineatum, high-altitude alpine fynbos grassland community 4.1, scirpus falsus – anthospermum basuticum, fynbos shrubland sub-community 4.1.1, macowania glandulosa – disa crassicornis, fynbos shrubland variant 4.1.1.1 merxmuellera disticha variant 4.1.1.2 erica reenensis sub-community 4.1.2, scirpus falsus – anthospermum basuticum, fynbos shrubland community 4.2, erica glaphyra – helichrysum trilineatum, alpine grassland sub-community 4.2.1, schoenoxiphium filiforme – euryops montanus, fynbos variant 4.2.1.1 helichrysum albo-brunneum variant 4.2.1.2 erica dissimulans variant 4.2.1.3 cliffortia filicauloides sub-community 4.2.2, erica dominans – schoenoxiphium schweickerdtii, heathland variant 4.2.2.1 passerina drakensbergensis variant 4.2.2.2 erica maesta variant 4.2.2.3 dimorphotheca jucunda variant 4.2.2.4 moraea alticola variant 4.2.2.5 merxmuellera stricta community 4.3, merxmuellera drakensbergensis – schoenoxiphium schweickerdtii, alpine grassland sub-community 4.3.1, helichrysum montanum – sebaea spathulata, forbland sub-community 4.3.2, glumicalyx goseloides – oxalis obliquifolia, wet meadow major community 1 only has a single diagnostic species, rhodohypoxis rubella, with constancy of 75 and no dominant species. there are no other species which occur with >± 50% dominance or constancy (online appendix 1), thus a single species is used to name it. the wetland vegetation consists of 11 relevés with nine species of which only pentaschistis airoides occurs with any significant presence. it constitutes largely aquatic forbs and contributes 10% to the total vegetation surveyed. the high-altitude bogs, mires and peatlands described by killick (1963, 1978a, 1978b, 1990), hillard and burtt (1987) and hill (1996) and found on the escarpment top in lesotho and the mont-aux-source/tugela falls area show floristic affinities with major community 1, rhodohypoxis rubella (wetland) and to a lesser extent major community 2, scirpus ficinioides – polevansia rigida (alpine grassland). aquatic taxa include aponogeton, crassula, limosella and rhodohypoxis on wet gravel plains and seeps. major community 2, crassula peploides – polevansia ridida (sheet rock grass and forblands), occurring on gravel seeps, has limited cover/abundance of most species and, after major community 1, is lowest in species richness and contributes 39% of the total relevés. the shallow soils, gravel plains and sheet rock would appear to be the limiting factor responsible for the low growth form and limited species numbers and presence. the scattered, open nature vegetation is problematic in naming major community 2. polevansia rigida was selected as the primary name as it had the highest cover/abundance and constancy out of all other species for all 28 relevés (online appendix 1). monocots comprise 61% of the diagnostic, constant and dominant species for major community 2, constituted by 31 species, 18 are monocots and 13 are dicots of which 3 are crassulaceae. no carnivorous plants were found, unlike at platberg, where urticaceae and droseraceae species formed distinct communities (brand, du preez & brown 2013). carnivorous plants from both these families are reported to be significant components of shallow soil and ephemeral flush vegetation communities occurring on inselbergs in the ivory coast (porembski & barthlott 1997). major community 3, pentaschistis exserta – merxmuellera stricta (high-altitude alpine grassland), is dominated by and contains the majority of the c3 high-altitude, medium to small bunch grasses, pentaschistis airoides, pentaschistis basutorum, festuca caprina and merxmuellera stricta, which was chosen as the second syntaxonomic name (online appendix 1) as it has a 54% presence and cover/abundance, greater than the other five co-dominant species. the fynbos shrubs, best represented in community 3.3, include cliffortia nitidula, erica dominans, macowania sororis, passerina montana with the forbs glumicalyx goseloides, hebenstretia dura, helichrysum album, helichrysum sutherlandii, helichrysum pagophilum and helichrysum trilineatum (26%, cover/abundance) the only non-fynbos shrub. geophytes include eucomis schijffii, gladiolus longicollis, and ornithogalum sephtonii; all high-altitude dac-endemic taxa. of significance is that no themeda triandra was found whilst both festuca costata and festuca caprina have very limited occurrence, unlike the themeda-festuca alpine veld of acocks (1988). additionally, chrysocoma ciliate, described as common and with increased occurrence on overgrazed areas (acocks 1988), was found in only one relevé with ‘+’ cover. major community 4, merxmuellera drakensbergensis – helichrysum trilineatum (high-altitude alpine fynbos grassland) represents the vegetation common to and found on all inselbergs. the dominant plants are alpine c3 grasses combined with helichrysum trilineatum, fynbos shrubs and the genus erica which form heathlands. major community 4 has species and physiognomic affinities with the ukhahlamba basalt grassland and the drakensberg afroalpine heathland (mucina & rutherford 2006). this cluster shares affinities with the lower-altitude drakensberg-amathole afromontane fynbos and the lesotho highlands basalt grassland (mucina & rutherford 2006). common dominant and prominent shrubs include the fynbos genera cliffortia, erica, euryops, helichrysum and macowania. the altitude precludes any trees or shrubs over 2.5 m tall, which do occur at lower, more protected altitudes. this major community contains most of the endemic forbs and geophytes, which constitute numerous rare, threatened and endangered taxa. sub-communities 4.1.1 and 4.2.1 are good representatives of this, as all three orchidaceae, disa crassicornis, dracomonticola virginea and satyrium longicauda lindl. var. jacottetianum, occur in them. other endemic geophytes include albuca humilis, eucomis schijffii, moraea alpine, moraea alticola, ornithogalum sephtonii and rhodohypoxis rubella. all inselberg vegetation has a grass component, including most of the wetlands, rocky outcrops and gravel seeps, with 89 out of 103 relevés complying. alpine grasses provide 98% cover/abundance and consistency throughout all four major communities, with the high-altitude vegetation dominated by c3 grasses. there are two c4 grasses: polevansia rigida, 14 of the 28 relevés forming community 2.1, and eragrostis caesia, occurring in 4 relevés and with low cover/abundance. although there were only 27 graminoids (grasses and sedges) compared with 162 non-graminoid shrubs and forbs, they contribute 81% (82 plots) of the total cover of the 103 plots. the sedge scirpus ficinioides was especially dominant in sub-communities 2.1.1 and 2.1.2, with the grass pentaschistis exserta providing 93.5% presence in major community 3 (14 of 15 relevés). dominant graminoid species varied from community to community, with merxmuellera drakensbergensis, scirpus falsus and schoenoxiphium schweickerdtii the most dominant (major community 4, 98% average cover). the monotypic alpine grass styppeiochloa gynoglossa, endemic to south africa, is dominant in community 2.2 and sub-community 4.1.2. community 2.2 is defined by ficinia cinnamomea, and ficinia gracilis is dominant for variants 4.2.2.2 and 4.2.2.3 (80% cover). festuca costata occurs in only nine plots with only a ‘+’ value. this is contrary to its reported occurrence and dominance used by acocks (1988) to define the themeda-festuca alpine veld and which mucina and rutherford (2006) state defines the core of the dac of endemism. of the 16 grass species (from nine genera), all are c3 except for polevansia rigida, the only mat-forming, stoloniferous, rhizomatous grass, and eragrostis caesia. the presence of eragrostis caesia, which is a short to medium height (450 mm – 600 mm) densely tufted grass, unlike other eragrostis species, may be explained by the same temperature protection – thermal microclimate created by dense or large tussock growth form of merxmuellera drakensbergensis and merxmuellera stricta (körner 2003; tieszen et al. 1979). this same physiognomic, large-tussock grass is exhibited by another c4 grass, andropogon amethystinus, which forms large tussocks, found on the aberdares and mt. kenya above 3000 m, and which has the same growth characteristic as eragrostis caesia (tieszen et al. 1979). plant physiological adaptation to alpine conditions can be seen in the large tufted grasses merxmuellera, pentaschistis, festuca, eragrostis caesia, styppeiochloa gynoglossa; the filiform leaves of the sedges and forbs cyprus schlechteri, ficinia cinnamomea, scirpus ficinioides, schoenoxiphium filiforme, schoenoxiphium schweickerdtii; the forbs felicia linearis and thesium pallidum; both tall and low cushion-shaped shrubs helichrysum trilineatum, cliffortia, erica, macowania sororis, passerina; low tufts anthospermum basuticum and muraltia. it is also displayed by the pubescence of many forbs, arctotis arctotoides, berkheya rhapontica, craterocapsa tarsodes, dimorphotheca jucunda, gazania krebsiana, geranium multisectum, glumicalyx goseloides, helichrysum album, helichrysum sutherlandii, helichrysum pagophilum, and senecio othonniflorus. succulents and geophytes, plants with below-ground bulbs or corms, are two other adaptations to alpine conditions. succulent plants found are delosperma sphalmanthoides and othonna burttii and geophytes include albuca polyphylla, disa crassicornis, dracomonticola virginea, eucomis schijffii, hesperantha baurii, ledebouria ovatifolia, moraea alticola, both species of rhodohypoxis and sebaea spathulata sebaea thodeana (hedberg 1964; körner 2003; masao et al. 2013). the high-altitude wetland community dominated by sedges and described by hill (1996) has affinities with major communities 3 and 4. the common sedges are scirpus ficinioides and schoenoxiphium filiforme. two dominant grasses of hill's high-altitude wetland community are merxmuellera drakensbergensis and pentaschistis oreodoxa, not found in the inselberg wetlands, but components of major community 4. from hill's (1996) description of the high-altitude wetland community, it shows more species and vegetation affinities with major community 4, the alpine grasslands, than with wetlands on the inselbergs. of all regions in the drakensberg, the inselbergs may be regarded as showing no anthropogenic influence from grazing, ploughing or fire, which does occur but is lighting induced. there is limited human presence: climbers have built cairns on most of the peaks, there are log books on sentinel and cathkin peak, a bovine skull – probably a cow placed as vulture food – and the metal top of a smoke grenade was found on cathkin, and tourists visit inner and outer horn via helicopter; however, human impact is negligible, unlike the heavily impacted, overgrazed, hoof-eroded adjacent escarpment in lesotho. syntaxonomical classification despite the broad treatment of the southern africa vegetation by mucina and rutherford (2006), large parts of south africa still remain to be surveyed in detail using phytosociological methods. there is still much work to be done before south africa reaches the level of vegetation analysis, classification and description which currently exists in europe. consequently, the syntaxonomical plant associations presented use the standard system in current use in south africa (brown et al. 2013), that is, community, sub-community and variant, which are analogous with alliance, association and sub-association, the original designations used by braun-blanquet (1964), and discussed by westhoff and van der maarel (1980). this article does not attempt to place the plant communities into a formal syntaxonomical classification of the established hierarchy of nomenclature and existing abstract categorisations of the european zurich-montpellier system (mueller-dombois & ellenberg 1974; weber, moravec & theurillat 2000). this survey encompasses 103 relevés, sampled on six inselbergs with a combined surface area of only 31.9 ha. it is felt that this survey does not give sufficient representation to cover the vegetation of the extensive alpine region of the drakensberg. it would be appropriate for a larger-scale survey, which incorporates plots on the escarpment adjacent as well as further north and south of the inselbergs, to be conducted. this would provide sufficient data to assign formal syntaxonomic ranks and names to the floristic clusters. the descriptions presented in this survey should be considered as a starting point. conservation and climate change top ↑ papers on inselbergs in the free state (brand et al. 2010; du preez 1991) have provided suggestions to assist decision-makers to design policies and provide protection for vegetation, relict plant populations and endemic taxa found only on inselbergs. as part of the biodiversity plan for free state, the department of economic development, tourism and environmental affairs (detea) has started the process of identifying inselbergs and making suggestions on a provincial scale to best manage and protect them (n. collins [detea], pers. comm., 03 may 2012). in kwazulu-natal (kzn), which shares a common provincial boundary with the free state, the protection of inselbergs is assured as part of the ukhahlamba-drakensberg park world heritage site management plan (ezemvelo kzn wildlife 2011). the management plan also stipulates on-going research to fill gaps in ecosystem function and reduce the risk of making incorrect management decisions. all world heritage sites require a protected area management plan as defined in article 1 of the world heritage convention act (no. 49 of 1999) (see https://www.environment.gov.za/sites/default/files/legislations/world_heritage_conventionact49.pdf), and which is incorporated in south african national legislation to protect biodiversity. kzn is also in the process of developing its buffer policy (forster 2007) to further protect the high mountains and their peaks (ezemvelo kzn wildlife 2011). as the first survey of these inselbergs, this article can make a contribution to the practical application of the kzn protected areas management plan. rising carbon dioxide levels with the concomitant increase in temperature will affect the composition and structure of alpine plant communities (körner 2003), and particularly grasses which use the c3 metabolic pathway, which are the most vulnerable to increase in temperature. the predicted effects are a drastic decrease in c3 grasses, as well as a decrease in the drakensberg alpine vegetation in both area and number of vegetation units (mucina & rutherford 2006). the alpine vegetation is embedded in the grassland biome and is defined as high-altitude drakensberg grassland, a subdivision of the grassland biome (mucina & rutherford 2006). a detailed phytosociological map has not been made of the vegetation above 2900 m. predictions are thus broad based and are extrapolations, which suggest an increase in woody components and a decrease in numbers of frost days and rainfall (mucina & rutherford 2006). peters (1992) predicted that a 3 ° c increase in global temperature would be equivalent to a 500 m upward shift in altitudinal zones. the threat posed by global warming to the world's alpine regions, including the drakensberg, will cause a significant reduction in the distribution of plants, and change their structure and composition, forcing some taxa to higher altitudes (taylor 1996), whilst those dac endemics and c3 grasses already found at altitude will face the threat of total loss of habitat. for inselbergs – such as platberg, which at its highest point is 2394.4 m, below the critical altitude of 2500 m (taylor 1996) –the loss of the taxa could be potentially catastrophic. conclusion top ↑ this article is a first attempt at a phytosociological analysis and vegetation description of high-altitude alpine plant communities in the drakensberg, which, using ordination methods, identified four different habitats. it is unknown how species numbers and cover in these habitats may change seasonally as a result of decrease in rainfall during the drier season. the overriding ecological factor is prolonged freezing, the effect of high altitude, followed by high rainfall and, undoubtedly, soil composition and its depth, which play an important part in lower amplitude ecological factors responsible for species richness and diversity. this also confirms within community plant affinities as well as the broader landscape level vegetation clustering. legislation is in place to protect the inselbergs on the drakensberg escarpment as the free state province is in the process of drafting a provincial biodiversity strategy to specifically protect inselbergs. acknowledgements top ↑ the fieldwork was supported by the national geographic committee for research and exploration (grant # 7920-05), without which funds the study would not have been possible. the principle investigator thanks the maluti drakensberg transfrontier project and the kwazulu/natal conservation staff for assistance with the fieldwork and plant identification. photographs are courtesy of richard lechemere-oertel, boyd escort and rob scott-shaw, who also helped with plant identification in the field and at the donald killick herbarium (cpf). the excellent maps of the study sites where composed by heidi snyman of ezemvelo kzn wildlife. competing interests the authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article. authors’ contributions r.f.b. (university of south africa) was the project leader and principle author, n.c. (free state department of economic, tourism and environmental affairs) provided theoretical input and processed the data in juice, and p.j.d.p. (university of the free state) contributed 53 relevés and edited the syntaxonomic table presented as online appendix 1. references top ↑ acocks, j.p.h., 1988, veld types of south africa, 3rd edn., memoirs of the botanical survey of south africa no. 57, national botanic gardens/botanical research institute, pretoria. barthlott, w., gröger, a. & porembski, s., 1993, ‘some remarks on the vegetation of tropical inselbergs: diversity and ecological differentiation’, biogéographica 69, 105–124. brand, r.f., du preez, p.j. & brown, l.r., 2010, ‘a floristic analysis of the vegetation of platberg, eastern free state, south africa’, koedoe 52(1), art. #710, http://dx.doi.org/10.4102/koedoe.v52i1.710 brand, r.f., du preez, p.j. & brown, l.r., 2013, ‘high altitude montane wetland vegetation classification of the eastern free state, south africa’, south african journal of botany 88, 223–236. http://dx.doi.org/10.1016/j.sajb.2013.07.011 braun-blanquet, j., 1964, pflanzensoziologie, grundzüge der vegetationskunde, 3rd edn., springer, vienna. http://dx.doi.org/10.1007/978-3-7091-8110-2 brown, l.r., du preez, p.j., bezuidenhout, h., bredenkamp, g.j., mostert, t.h.c. & collins, n.b., 2013, ‘guidelines for phytosociological classification and description of vegetation in southern africa’, koedoe 55(1), http://dx.doi.org/10.4102/koedoe.v55i1.1103 burke, a., 2001, ‘determinants of inselberg floras in arid nama karoo landscapes’, journal of biogeography 28, 1211–1220. http://dx.doi.org/10.1046/j.1365-2699.2001.00623.x burke, a., 2002, ‘plant communities of a central namib inselberg landscape’, journal of biogeography 13(4), 483–492. burke, a., 2004, ‘from plains to inselbergs: species in special habitats as indicators of climate change?’, journal of biogeography 31, 831–841. http://dx.doi.org/10.1046/j.1365-2699.2003.00984.x carbutt, c. & edwards, t.j., 2006, ‘the endemic and near-endemic angiosperms of the drakensberg alpine centre’, south african journal of botany 72, 105–132. http://dx.doi.org/10.1016/j.sajb.2005.06.001 du preez, p.j., 1991, a syntaxonomical and synecological study of the vegetation of the south-eastern orange free state and related areas with special reference to korannaberg, phd thesis, university of the orange free state, bloemfontein. ezemvelo kwazulu-natal wildlife, 2011, concept development plan: ukhahlamba-drakensberg park world heritage site, ezemvelo kwazulu-natal wildlife, pietermaritzburg. forster, c.w., 2007, ‘specifications and methodology for the establishment of the buffer zone to the ukhahlamba drakensberg world heritage site park’, draft final report for kzn wildlife, pietermarizburg. gauch, h.g., 1982a, multivariate analysis in community ecology, cambridge university press, cambridge. http://dx.doi.org/10.1017/cbo9780511623332 gauch, h.g., 1982b, ‘noise reduction by eigenvector ordinations’, ecology 63(6), 1643–1649. http://dx.doi.org/10.2307/1940105 germishuizen, g., meyer, n.l., steenkamp, y. & keith, m. (eds.), 2006, a checklist of south african plants, south african botanical diversity network, report no. 41, sabonet, pretoria. gibbs-russell, g.e., watson, l., koekemoer, m., smook, l., barker, n.p., anderson, h.m. et al., 1991, grasses of southern africa, memoirs of the botanical survey of south africa no. 58, national botanic gardens/botanical research institute, pretoria. hedberg, o., 1964, ‘features of afroalpine plant ecology’, acta phytogeographica suecica 49, 1–144. heilmeier, h., durka, w., woitke, m. & hartung, w., 2005, ‘ephemeral pools as stressful and isolated habitats for the endemic aquatic resurrection plant chamaegigasintrepidus’, phytocoenologia 35(2–3), 449–468. http://dx.doi.org/10.1127/0340-269x/2005/0035-0449 hennekens, s.m. & schaminée, j.h., 2001, ‘turboveg: a comprehensive database management system for vegetation data’, journal of vegetation science 12, 589–591. http://dx.doi.org/10.2307/3237010 hill, m.o., 1979, twinspan – a fortran program for arranging multivariate data in an ordered two-way table by classification of the individuals and attributes, section of ecology and systematics, cornell university, ithaca, new york, usa. hill, t.r., 1996, ‘description, classification and ordination of the dominant vegetation communities, cathedral peak, kwazulu-natal drakensberg’, south african journal of botany 62, 263–269. hillard, o.m. & burtt, b.l., 1987, the botany of southern natal drakensberg, national botanical gardens, cape town. huntley, b.j. (ed.), 1989, biotic diversity in southern africa: concepts and conservation, oxford university press, cape town. killick, d.j.b., 1963, an account of the plant ecology of the cathedral peak area of the natal drakensberg, memoirs of the botanical survey of south africa no.34, government printer, pretoria. killick, d.j.b., 1978a, ‘the afro-alpine region’, in m.j.a. werger (ed.), biogeography and ecology of southern africa, pp. 515–560, junk, the hague. http://dx.doi.org/10.1007/978-94-009-9951-0_12 killick, d.j.b., 1978b, ‘notes on the vegetation of the sani pass area of the southern drakensberg’, bothalia 12, 537–542. killick, d.j.b., 1990, a field guide to the flora of the natal drakensberg, jonathan ball, johannesburg. körner, c., 2003, alpine plant life: functional plant ecology of high mountain ecosystems, springer, berlin. http://dx.doi.org/10.1007/978-3-642-18970-8 legendre, p. & legendre, l., 1998, numerical ecology, elsevier science, amsterdam. lepš, j. & hadincová, v., 1992, ‘how reliable are our vegetation analyses?’, journal of vegetation science 3(1), 119–124. http://dx.doi.org/10.2307/3236006 marneweck, g.c. & kotze, d., 1999, ‘appendix w6: guidelines for delineation of wetland boundary and wetland zones’, resource directed measures for protection of water resources: wetland ecosystems, department of water affairs and forestry, viewed 12 november 2013, from https://www.dwa.gov.za/documents/policies/wrpp/wetland%20ecosystems.htm masao, c.a., gizaw, a., piñeiro, r., tusiime, f.m., wondimu, t., abdi, a.a. et al., 2013, ‘phylogeographic history and taxonomy of some afro-alpine grasses assessed based on aflps and morphometry: deschampsia cespitosa, d. angusta and koeleria capensis’, alpine botany 123(2), 107–122. mokuku, c., 1991, ‘classification of the alpine plant communities of the mjika-lisiu’, msc thesis, department of biological sciences, university of zimbabwe, harare. mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, strelitzia 19, south african national biodiversity institute, pretoria. mueller-dombois, d. & ellenberg, h., 1974, aims and methods of vegetation ecology, wiley, new york. parmentier, i., oumorou, m., porembski, s., lejoly, j. & decocq, g., 2006, ‘ecology, distribution, and classification of xeric monocotyledonous mats on inselbergs in west africa and atlantic central africa’, phytocoenology 36(4), 547–564. http://dx.doi.org/10.1127/0340-269x/2006/0036-0547 peters, r.l., 1992, ‘conservation of biological diversity in the face of climatic change’, in r.l. peters & t.e. lovejoy (eds.), global warming and biological diversity, pp. 15–30, yale university press, new haven, connecticut. podani, j., 2006, ‘braun-blanquet's legacy and the data analysis in vegetation science’, journal of vegetation science 17, 113–117. http://dx.doi.org/10.1111/j.1654-1103.2006.tb02429.x porembski, s. & barthlott, g.b., 1997, ‘a species poor tropical sedge community afrotrilepis pilosa mats on inselbergs in west africa’, nordic journal of botany 16(3), 239–246. http://dx.doi.org/10.1111/j.1756-1051.1996.tb00222.x porembski, s. & brown, g., 1995, ‘the vegetation of inselbergs in the comoé national park (ivory coast)’, conservatoire et jardin botanicues de geneve 50(2), 351–365. porembski, s., szarzynske, j., mund, j-p. & barthlott, w., 1996, ‘biodiversity and vegetation of small-size inselbergs in a west african rain forest (taï, ivory coast)’, journal of biogeography 23, 47–55. http://dx.doi.org/10.1046/j.1365-2699.1996.00982.x porembski, s., fisher, e. & biedenger, n., 1997, ‘vegetation of inselbergs, quartzitic outcrops and ferricretes in rwanda and eastern zaire (kivu)’, bulletin du jardin botanique de l’etat, bruxelles 66, 81–95. seine, r., becker, u., porembski, s., follman, g. & barthlott, w., 1998, ‘vegetation of inselbergs in zimbabwe’, edinburgh journal of botany 55, 267–293. http://dx.doi.org/10.1017/s0960428600002195 taylor, d., 1996, ‘mountains’, in w.m. adams, a.s. goudie & a.r. orme (eds.), the physical geography of africa, pp. 287–306, oxford university press, oxford. tichý, l. & holt, j., 2006, juice: program for management, analysis and classification of ecological data, program manual, vegetation science group, masaryk university, brno. tieszen, l.l., senyimba, m.m., imbamba, s.k. & troughton, j.h., 1979, ‘the distribution of c3 and c4 grasses and carbon isotope discrimination along an altitudinal and moisture gradient in kenya’, oecologia 37(3), 337–350. http://dx.doi.org/10.1007/bf00347910 van noort, s., gardiner, a.j. & tolley, k.a., 2007, ‘new records of ficus (moraceae) species emphasize the conservation of significance of inselbergs in mozambique’, south african journal of botany 73, 642–649. http://dx.doi.org/10.1016/j.sajb.2007.04.063 weber, h.e., moravec, j. & theurillat, j.p., 2000, ‘international code of phytosociological nomenclature’, journal of vegetation science 11, 739–768. http://dx.doi.org/10.2307/3236580 westhoff, v. & van der maarel, e., 1980, ‘the braun-blanquet approach’, in r.h. whitaker (ed.), classification of plant communities, pp. 287–378, kluwer academic, the hague. whitaker, r.h., 1980, classification of plant communities, kluwer academic, the hague. article information author: ben j. strohbach1 affiliation: 1school of natural resources and spatial sciences, polytechnic of namibia, namibia correspondence to: ben strohbach postal address: private bag 13388, windhoek, namibia dates: received: 30 oct. 2012 accepted: 23 may 2014 published: 28 nov. 2014 how to cite this article: strohbach, b.j., 2014, ‘vegetation of the eastern communal conservancies in namibia: i. phytosociological descriptions’, koedoe 56(1), art. #1116, 18 pages. http://dx.doi.org/10.4102/ koedoe.v56i1.1116 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/ koedoe.v56i1.1116 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. vegetation of the eastern communal conservancies in namibia: i. phytosociological descriptions in this original research... open access • abstract • introduction • research method and design    • study area       • climate       • landscapes and geology    • procedure • results    • 1. acacio fleckii–terminalietum prunioidis ass. nov.    • 2. acacio tortilis–combretetum imberbis ass. nov.    • 3. ptycholobio biflori–acacietum luederitzii ass. nov.    • 4. eragrostio echinocloideae–eriocephaloetum luederitziani ass. nov.    • 5. acacio melliferae–catophractetum alexandri ass. nov.    • 6. acacio melliferae–hyphaenetum petersianae ass. nov.    • 7. the panicum gilvum–marsilea vlei community    • 8. eragrostio rigidioris–urochloaetum brachyurae ass. nov.    • 9. tarchonantho camphorati–acacietum eriolobae ass. nov.    • 10. stipagrostio uniplumis–acacietum melliferae ass. nov.       • 10.1. stipagrostio uniplumis–acacietum melliferae rhigozetosum brevispinosi subass. nov.       • 10.2. stipagrostio uniplumis–acacietum melliferae typicum subass. nov.    • 11. terminalio sericeae–acacietum eriolobae ass. nov.    • 12. combreto collini–terminalietum sericeae ass. nov.       • 12.1. combreto collini–terminalietum sericeae acacietosum melliferae subass. nov.       • 12.2. combreto collini–terminalietum sericeae grewietosum flavae subass. nov.       • 12.3. combreto collini–terminalietum sericeae typicum subass. nov.       • 12.4. combreto collini–terminalietum sericeae burkeaetosum africanae subass. nov.    • 13. terminalio sericeae–schinziophytetum rautanenii ass. nov.    • 14. burkeo africanae–pterocarpetum angolensis ass. nov. • discussion    • higher syntaxonomic groupings and relations to similar vegetation in southern africa    • vegetation mapping    • land use and land degradation • conclusion • acknowledgements    • competing interests • references abstract top ↑ the establishment of communal conservancies aims to have the local communities share in the benefits especially of wildlife resources, in this way spearheading the conservation of the environment. the desert margins programme in namibia aimed to develop vegetation resource data for the otjituuo, okamatapati, ozonahi, african wild dog, otjinene, epukiro, otjombinde, omuramba ua mbinda, eiseb and ondjou communal conservancies, in order to assist with natural resource planning. for this purpose, a phytosociological survey of this area, with 422 relevés, was conducted during 2004. the data was captured in turboveg and forms part of the namibian phytosociological database (givd af-na-001). the data was split into two, representing two major land forms, the ‘hardeveld’ and the ‘sandveld’, respectively. a classification was undertaken using the modified two-way indicator species analysis (twinspan) procedure. further refinements, based on field observations and literature sources, were performed using cocktail procedures. thirteen vegetation associations were formally described in this article, of which two were subdivided into subassociations. these associations can broadly be grouped into broad-leaved savanna types typical of the central and northern kalahari of namibia and microphyll savannas found on the transitions to the central plateau. one association, the burkeo africanae–pterocarpetum angolensis, forms the southern fringe of the zambesian baikiaea woodlands ecoregion of the world wildlife fund, whilst all the other associations fall within the kalahari acacia–baikiaea woodlands ecoregion. the combreto collini–terminalietum sericeae is the most widespread association and dominates the landscape. threats to the vegetation include overutilisation and regular fires, both of which could easily lead to desertification. this threat is aggravated by global climate change.conservation implications: this article described 13 plant associations of the central kalahari in eastern namibia, an area hitherto virtually unknown to science. the information presented in this article forms a baseline description, which can be used for future monitoring of the vegetation under communal land use. introduction top ↑ since namibia’s independence there has been a substantial increase in efforts to develop the country’s communal farming areas. many of these areas have been, and still are, subjected to poor and indifferent management practices resulting from a lack of sound ecological (baseline) data and hence a poor understanding especially of the vegetation resource base – its biodiversity, dynamics, possibilities of utilisation and its limitations (hines 1992; mendelsohn et al. 2002). this is also the case when looking at the communal lands in eastern namibia, which consist of what was formerly known as bushmanland, hereroland west and hereroland east. a phytosociological study on the vegetation of bushmanland (part of the present tsumkwe district) was conducted by hines (1992). information regarding the vegetation of the former hereroland could thus far only be approximated from the preliminary vegetation map of namibia (giess 1998), as well as the relative homogenous farming areas report (department landbou tegniese dienste 1979). the above report lists farming possibilities and constraints of the commercial farming area and includes a grazing-capacity map based on expert opinion. this map has been found to be outdated (espach, lubbe & ganzin 2006; lubbe 2005) mainly because the concept of carrying capacity has since changed from a fixed rate per area to a biomass concept, which is an (annually changing) estimate of the amount of fodder produced during the previous season’s rainfall (bester 1988; espach et al. 2006; espach, lubbe & ganzin 2010). the preliminary vegetation map of namibia is regarded as relatively accurate as far as the delineation of vegetation types is concerned, but some anomalies have been found. furthermore, its descriptions of namibian vegetation types are very basic. both publications were based on limited observations and some photographic records, but no actual ecological surveys were conducted when they were compiled. few attempts have since been undertaken to describe the vegetation of namibia further (burke & strohbach 2000; strohbach & jürgens 2010).the target area selected for this study was the former hereroland, which now consists of communal and emerging commercial farms within the omaheke and otjozondjupa regions of central-eastern namibia. in order to empower the farmers to sustainably utilise their natural resources, especially wildlife resources, the establishment of communal conservancies is promoted in namibia (weaver & petersen 2008; weaver & skyer 2005). at the start of the study (2004), nine communal conservancies were planned; by 2005, the following conservancies had already been proclaimed: ozonahi (okakarara), african wild dog (okandjatu), otjituuo and okamatapati. the ondjou conservancy (gam) followed in 2006, whilst the eiseb, omuramba ua mbinda and otjombinde conservancies (the former two forming the ‘eiseb block’ and the latter the ‘rietfontein block’) were only proclaimed in 2011 (nambian association of community based natural resource management [cbnrm] support organisations 2011; steynberg 2011). of those included in this study, only the epukiro and otjinene farming areas are not yet considered as emerging conservancies (figure 1). figure 1: overview map of the communal conservancies within the omaheke and otjozondjupa regions in namibia. the average annual rainfall is indicated by the isohyets, with their values in millimetres, indicated on the right. the goal of the desert margins programme (namibia) was the conservation and restoration of biodiversity in namibia’s desert margins. one of the major outputs was the establishment of baseline information on natural resources available to local communities in selected areas. in order to establish such baseline information, a survey of the vegetation types was commissioned for these communal conservancies as a baseline of the ecosystems to be managed. this is in line with generally proposed uses of such vegetation descriptions and maps (brown et al. 2013; strohbach 2012). the nomenclature throughout this article follows klaassen and kwembeya (2013). research method and design top ↑ study area the study area consisted of the farming areas of okakarara, okondjatu, otjituuo, okamatapati, gam, eiseb, rietfontein, epukiro and otjinene in the otjozondjupa and omaheke regions (figure 1), covering 57 414.72 km2. climate very little meteorological data has officially been recorded within the study area. climate variables for the area have been extrapolated from records collected in gobabis, grootfontein, tsumkwe and ghanzi (in botswana). average long-term annual rainfall ranges from about 325 mm to 350 mm in the south-eastern corner (rietfontein block) and the westernmost part (around okakarara) and from about 425 mm to 450 mm in the central part of the area – roughly from okondjatu and otjinene to the mangetti dune (figure 1). an important aspect of this rainfall is its often erratic nature within the rainfall season and from year to year. droughts are regarded as common (botha 1998; leser 1972; mendelsohn & el obeid 2002). during winter, mean daily temperatures range from 2.5 °c (minimum) to 22.2 °c (maximum) (with 8 frost days) in the southernmost parts of the study area and from about 4.3 °c (minimum) to 23.6 °c (maximum) (with 2 frost days) in the northern parts. during summer, mean daily temperatures range from 17.4 °c (minimum) to 31.8 °c (maximum) in the southern parts and from 17.9 °c (minimum) to 30.2 °c (maximum) in the northern parts. however, daytime temperatures can soar to just below 40.0 °c, especially during november and december (mendelsohn & el obeid 2002). landscapes and geology the study area is situated on the vast kalahari system (geological survey 1980; thomas 1988; mendelsohn et al. 2002; mendelsohn & el obeid 2003). this ancient basin has been gradually filled up with silt and clay sediments and windblown sands during the tertiary to quaternary periods (32–39 ma) (geological survey 1980). the windblown sands dominate the surface soils of the study area. occasionally, exposed rock formations can be found – these are mainly sandstones, limestones, schists and dolomites of the much older karoo and damara sequences.the sandy landscapes are generally flat to rolling (with 6° – 9° slopes); these plains are incised by omuramba valleys or alternated with vegetated fossil (no longer actively moving) dunes. an omuramba (plural: omiramba) can best be described as a shallow watercourse with no visible gradient (king 1963). leser (1972) differentiates between coarse-grained yellow sands, which were found mostly on plains, and finer-grained reddish sands, present especially on the fossil dunes. apart from the omuramba valleys, which often have exposed or underlying calcretes and limestone ridges, occasional pans formed by the limited runoff in the area have grey clays to grey sands. a more thorough study of the soils and landscapes is presented by strohbach et al. (2004). procedure surveying followed the general method employed for the vegetation survey of namibia project (strohbach 2001). sampling of 422 plots throughout the study area took place during april and may 2004. owing to the vastness of the area, and the limited access to it, emphasis was placed on the sampling of dominant vegetation types (rather than rare niche types) along roads, tracks and cutlines, following a stratified systematic approach. false colour landsat thermatic mapper (tm) imagery was used for stratification. at each survey plot of 20 m × 50 m (1000 m2), a braun-blanquet relevé was compiled. all occurring vascular plant species were noted down, as well as their typical growth forms and estimated percentage crown cover. habitat descriptors included: the position by way of a gps-reading (referenced to wgs84), the landscape, local topography, slope and aspect, lithology, degradation indicators, as well as a photograph. unknown species and reference specimens were collected, identified and deposited at the national herbarium of namibia (wind).the relevé data was captured on turboveg (hennekens & schaminée 2001) and forms part of the namibian phytosociological database (givd id: af-na-001) (strohbach & kangombe 2012; see dengler et al. 2011). for the original report (strohbach et al. 2004), this data was classified with phytotab (westfall et al. 1982). the initial study identified three associations, which were represented by a limited number of relevés, but which are known also from other studies (hines 1992, own unpublished data). for this reason, relevant relevé data available in the namibian phytosociological database (givd id: af-na-001) from the tsumkwe district (25 relevés from hines 1992 surveyed on 400 m2 plots) and from the southern mangetti area near tsintsabis in the grootfontein district (five relevés from own unpublished data surveyed on 1000 m2 plots) was extracted and included in the dataset. the final dataset consists of 451 relevés, with 448 species. for the present study the classification was repeated using juice 7 (tichý 2002). total inertia was used as diversity measure, whilst pseudospecies cut levels set at 0%, 2%, 5% and 20%, were utilised because the gradients were often very gradual, resulting in strong overlapping of species distribution. for convenience, the dataset was split into two, using the first division of a modified two-way indicator species analysis (twinspan) classification (roleček et al. 2009). the two relevé groups were further classified, again using modified twinspan with the same settings as before. the highest level of divisions still yielding floristically well-characterised clusters was used. after inspection of the results, it was realised that further refinements were necessary on some of the clusters, based on both field observations and previous vegetation descriptions, in particular cole and brown (1976), hines (1992), leser (1972) and strohbach et al. (2004). this was performed with cocktail (bruelheide 1997; bruelheide & flintrop 1994) using known character species for selection. in several cases, recombinations of clusters as subassociations or even associations were necessary to produce ecological significant results. the fact that such clusters could not be differentiated with diagnostic species but had similar habitat, structure and dominant species composition was taken as criteria for recombination. diagnostic species were determined using the phi co-efficient of association (chytrý et al. 2002). for this calculation the relevé numbers were standardised following tichý and chytrý (2006). species with phi ≥ 40 were considered as diagnostic and with phi ≥ 60 as highly diagnostic; however, species with a non-significant fidelity at α = 0.05 using fisher’s exact test were omitted. species occurring at least with a 60% frequency were regarded as constant and with at least an 80% frequency as highly constant. further calculations to describe the structure and species richness were performed using the available growth form data and species richness and density. for the calculation of species density (number of species per 1000 m2), the relevé data from hines (1992) was excluded, as these were sampled on 400 m2 plots, not 1000 m2 plots as all other relevés. results top ↑ the classification results are depicted in figure 2 as a dendrogram. the first division of the dataset represents two distinct habitat and physiognomic groupings: the ‘sandveld’ or broad-leafed savannas on very sandy habitats (265 relevés with 305 species) and the ‘hardeveld’ (‘hard veld’) on slightly to very loamy soils, occasionally even shallow rocky substrates with microphyll savannas (186 relevés with 402 species). figure 2: dendrograms of the classification results of, (a) microphyll savannas and (b) broad-leaved savannas, also indicating refinements to certain clusters performed during data processing. the numbers correspond to the association numbers used in the text. within the hardeveld (figure 2 top), the second cluster was split into two associations with the aid of cocktail (bruelheide 1997; bruelheide & flintrop 1994), using the presence of felicia clavipilosa subsp. clavipilosa, lycium bosciifolium, ptycholobium biflorum subsp. angolensis and xerophyta humilis versus combretum imberbe. in addition to differences in composition, these two associations were recognised in the field as different in structure and habitat. a third association (no. 5), based on the dominance (> 5% cover) of catophractes alexandri, but the absence of indicator species eriocephalus luederitzianus and hyphaene petersiana, was defined also using cocktail. this is in line with various literature sources, in particular cole and brown (1976), hines (1992), leser (1972) and strohbach et al. (2004). the initial clusters 8, 9 and 10 could not be differentiated through diagnostic species, only through subtle differences in constant and dominant species. therefore clusters 9 and 10 were combined as a subassociation (10.2), which, in turn, was combined with cluster 8 (subassocaition 10.1) to form a larger association (10) because of the high similarity in permanent species composition, structure and habitat. within the group of relevés representing the broad-leaved savannas, again, cocktail had to be employed to strengthen cluster 1, based on the presence of pterocarpus angolensis and burkea africana. this cluster was also split into two, based on the presence of schinziophyton rautanenii and strychnos pungens. these refinements were based on findings from the original study (strohbach et al. 2004) as well as hines (1992). clusters 2 to 5 were recognised as subassociations of a single association, because of the very similar structure, dominance by terminalia sericea and various combretum species and similar habitat. in effect, these four subassociations represent a precipitation gradient (strohbach & kutuahuripa submitted). the full phytosociological table is presented in downloadable form in online appendix 1 and, similarly, the synoptic table in online appendix 2. the classification yielded 11 communities, of which 10 are formally described associations, following the international code for phytosociological nomenclature (weber, moravec & theurillat 2000). species with high fidelity (phi ≥ 60) and high constancy (frequency ≥ 80%) are indicated in bold. structural description follows edwards (1983), whilst habitat descriptions follow the soil and terrain database (soter) terminology (food and agricultural organization [fao] 1995). 1. acacio fleckii–terminalietum prunioidis ass. nov. the type for this association is relevé 1336, sampled on 29 april 2004 at 20°12’40” s, 20°46’22” e. only two relevés of this association were sampled in the present study area, but this association occurs extensively within the nyae-nyae conservancy further north (hines 1992). an additional 15 relevés from the hines (1992) dataset were included in the present dataset. this association is characterised by the diagnostic species croton gratissimus, grewia bicolor, terminalia prunioides, hibiscus calyphyllus, talinum arnotii, searsia marlothii, combretum apiculatum subsp. apiculatum, grewia villosa, barleria senensis, achyranthes aspera var. sicula, panicum maximum, solanum kwebense and crotalaria flavicarinata. these are constantly associated with pupalia lappacea, grewia flavescens, dichrostachys cinerea, brachiaria deflexa and urochloa brachyura. a total of 63 species have been observed in this association, with, on average, 20 species per 1000 m2.hines (1992) refers to these thickets as terminalia prunioides association and describes them as occurring on some sub-outcropping geological features, specifically fold structures (e.g. near cin qo) throughout the nyae-nyae conservancy. bearing out hines’s (1992) findings, these thickets were found on low limestone ridges, mostly covered by shallow sand deposits with few stones on the surface (< 1% cover), north of gam. here the altitude ranges between 1110 m.a.s.l. and 1130 m.a.s.l., with generally rolling slopes. the soils are mostly shallow. as a result of the loose sand cover, few or no signs of crusting, but slight to moderate signs of erosion were observed. the thickets sampled in this study are situated near the resource-poor settlements of gam and showed severe signs of degradation through overgrazing and felling of trees for timber. terminalia prunioides dominated the tree layer at between 40% and 60%. the shrub understorey was, on average, less than 10%, making this association a tall, closed woodland. grass cover was generally variable, dominated by annual grasses (figure 3a and figure 4a). only two perennial grass species were observed, these being panicum maximum and cenchrus ciliaris, indicative of the poor grazing resource these thickets represent. although this could be attributed to overgrazing in the area, heavy shading by the overstorey would also prevent the growth of good grazing. figure 3: box-and-whisker plots of the typical structure of the various associations and subassociations, (a) acacio fleckii-terminalietum prunioidis, (b) acacio tortilis-combretetum imberbis, (c) ptycholobio biflori-acacietum luederitzii, (d) eragrostio echinocloideae-eriocephaloetum luederitziani, (e) acacio melliferae-catophractetum alexandri, (f) acacio melliferae-hyphaenetum petersianae, (g) eragrostio rigidioris-urochloaetum brachyurae, (h) tarchonantho camphorate-acacietum eriolobae, (i) stipagrostio uniplumis-acacietum melliferae rhigozetosum brevispinosi, (j) stipagrostio uniplumis-acacietum melliferae typicum, (k) terminalio sericeae-acacietum eriolobae, (l) combreto collini-terminalietum sericeae acacietosum melliferae, (m) combreto collini-terminalietum sericeae grewietosum flavae, (n) combreto collini-terminalietum sericeae typicum, (o) combreto collini-terminalietum sericeae burkeaetosum africanae, (p) terminalio sericeae-schinziophytetum rautanenii and (q) burkeo africanae-pterocarpetum angolensis. figure 4: typical examples of various associations and subassociations, (a) acacio fleckii–terminalietum prunioidis, (b) acacio tortilis–combretetum imberbis, (c) ptycholobio biflori–acacietum luederitzii, (d) eragrostio echinocloideae–eriocephaloetum luederitziani, (e) acacio melliferae–catophractetum alexandri, (f) acacio melliferae–hyphaenetum petersianae, (g) panicum gilvum–marsilea vlei, (h) eragrostio rigidioris–urochloaetum brachyurae, (i) tarchonantho camphorate–acacietum eriolobae, (j) stipagrostio uniplumis–acacietum melliferae rhigozetosum brevispinosi, (k) stipagrostio uniplumis–acacietum melliferae typicum, (l) terminalio sericeae–acacietum eriolobae, (m) degraded combreto collini–terminalietum sericeae acacietosum melliferae, (n) combreto collini–terminalietum sericeae acacietosum melliferae in better condition, (o) combreto collini–terminalietum sericeae grewietosum flavae, (p) combreto collini–terminalietum sericeae typicum, (q) combreto collini–terminalietum sericeae burkeaetosum africanae, (r) terminalio sericeae–schinziophytetum rautanenii and (s) burkeo africanae–pterocarpetum angolensis. figure 4 (continues...): typical examples of various associations and subassociations, (a) acacio fleckii–terminalietum prunioidis, (b) acacio tortilis–combretetum imberbis, (c) ptycholobio biflori–acacietum luederitzii, (d) eragrostio echinocloideae–eriocephaloetum luederitziani, (e) acacio melliferae–catophractetum alexandri, (f) acacio melliferae–hyphaenetum petersianae, (g) panicum gilvum–marsilea vlei, (h) eragrostio rigidioris–urochloaetum brachyurae, (i) tarchonantho camphorate– acacietum eriolobae, (j) stipagrostio uniplumis–acacietum melliferae rhigozetosum brevispinosi, (k) stipagrostio uniplumis–acacietum melliferae typicum, (l) terminalio sericeae–acacietum eriolobae, (m) degraded combreto collini–terminalietum sericeae acacietosum melliferae, (n) combreto collini–terminalietum sericeae acacietosum melliferae in better condition, (o) combreto collini–terminalietum sericeae grewietosum flavae, (p) combreto collini–terminalietum sericeae typicum, (q) combreto collini–terminalietum sericeae burkeaetosum africanae, (r) terminalio sericeae–schinziophytetum rautanenii and (s) burkeo africanae–pterocarpetum angolensis. figure 4 (continues...): typical examples of various associations and subassociations, (a) acacio fleckii–terminalietum prunioidis, (b) acacio tortilis– combretetum imberbis, (c) ptycholobio biflori–acacietum luederitzii, (d) eragrostio echinocloideae–eriocephaloetum luederitziani, (e) acacio melliferae– catophractetum alexandri, (f) acacio melliferae–hyphaenetum petersianae, (g) panicum gilvum–marsilea vlei, (h) eragrostio rigidioris–urochloaetum brachyurae, (i) tarchonantho camphorate–acacietum eriolobae, (j) stipagrostio uniplumis–acacietum melliferae rhigozetosum brevispinosi, (k) stipagrostio uniplumis–acacietum melliferae typicum, (l) terminalio sericeae–acacietum eriolobae, (m) degraded combreto collini–terminalietum sericeae acacietosum melliferae, (n) combreto collini–terminalietum sericeae acacietosum melliferae in better condition, (o) combreto collini–terminalietum sericeae grewietosum flavae, (p) combreto collini–terminalietum sericeae typicum, (q) combreto collini–terminalietum sericeae burkeaetosum africanae, (r) terminalio sericeae– schinziophytetum rautanenii and (s) burkeo africanae–pterocarpetum angolensis. 2. acacio tortilis–combretetum imberbis ass. nov. the type for this association is relevé 1240, sampled on 23 april 2004 at 20°40’57” s, 17°42’53” e. six relevés were classified into this association. it is characterised by the diagnostic species guilleminea densa, mollugo nudicaulis, lotononis listii, combretum imberbe, dactyloctenium aegyptium, alternanthera pungens, kalanchoe rotundifolia, corchorus tridens, zehneria marlothii, portulaca hereroensis, aptosimum glandulosum and chloris virgata. these are constantly associated with eragrostis porosa, ziziphus mucronata, tragus berteronianus, eragrostis trichophora, chamaesyce inaequilatera, acrotome species and acacia mellifera subsp. detinens. a total of 91 species were observed here, with, on average, 37 species per 1000 m2.very striking are the tall combretum imberbe (leadwood) trees interspersed with smaller trees of acacia tortilis in this association. the understorey of shrubs is variable, mostly less than 10% cover, making this a tall, semi-open bushland (figure 3b and figure 4b). the omuramba omatako near okakarara has a distinct wetland characteristic, with eragrostis rotifer present in two, and schoenoplectus senegalensis and marsilea macrocarpa present in one, of the sites – all species common in areas with seasonal standing water. although some of the patches are drier and have a higher cover of grasses, including some perennial grasses, this syntaxon is to be regarded as having a low grazing capacity. if left ungrazed during the rainy season (which it was evidently not), this could, like the oshanas in the central north, be valuable winter grazing. the omuramba omatako is at an altitude of between 1290 m.a.s.l. and 1310 m.a.s.l. south of okakarara and it flows in a north-easterly direction towards otjituuo. the slopes are flat to gently undulating, whilst the soils are strongly crusted. at one of the sites, signs of selective wood harvesting of combretum imberbe was observed. 3. ptycholobio biflori–acacietum luederitzii ass. nov. the type for this association is relevé 1233, sampled on 23 april 2004 at 20°42’01” s, 17°35’49” e. six relevés were classified into this association. it is characterised by the diagnostic species felicia clavipilosa subsp. clavipilosa, lycium bosciifolium, hibiscus micranthus, xerophyta humilis, ptycholobium biflorum subsp. angolensis, barleria lancifolia, pavonia burchellii, chloris virgata, hibiscus vitifolius, kyphocarpa angustifolia, calostephane divaricata, ocimum americanum var. americanum, solanum delagoense, barleria senensis, aristida rhiniochloa and dactylotenium aegyptium. these are constantly associated with tragus berteronianus, eragrostis trichophora, acacia mellifera subsp. detinens, acacia luederitzii, ziziphus mucronata, kleinia longiflora, grewia flava, gisekia africana, eragrostis porosa, eragrostis dinteri, brachiaria deflexa, boscia albitrunca, aristida adscensionis, pupalia lappacea, pogonarthria fleckii, grewia flavescens, dicoma tomentosa, cleome rubella, asparagus cooperi and aristida congesta subsp. congesta. a total of 89 species were observed here, with, on average, 48 species per 1000 m2.this association has a consistent tree layer of about 6 m – 8 m in height with a cover of about 12%. the shrub layer reaches a cover of about 24%, thus making the structure a short, semi-open thicket (figure 3c and figure 4c). the ptycholobio biflori–acacietum luederitzii occurs typically on the floodplains around the omuramba omatako, south of okakarara in the ozonahi conservancy, characterised by the erosion plains around the omatako omuramba throughout its catchment. these floodplains are gently undulating and generally at an altitude of 1290 m.a.s.l. – 1330 m.a.s.l. the soils are shallow, with a hardpan layer at a depth of 30 cm, and generally strongly crusted. only slight sheet erosion was observed. these floodplains are (relatively) undisturbed; although, a few show signs of bush encroachment. on one occasion severe overgrazing was observed near a village. grazing value is low, with only two perennial grasses (eragrostis trichophora and aristida congesta) being found consistently. both are low producers of biomass, whilst aristida congesta is also known as generally hard and unpalatable (müller 2007). the annual grass layer is dominated by eragrostis porosa and aristida rhiniochloa. the latter can produce fairly high amounts of biomass, but is a very hard grass with a strong, pungent seed. both these species are regarded as unpalatable. the poor grazing condition is aggravated by the consistent occurrence of the poisonous geigeria ornativa. 4. eragrostio echinocloideae–eriocephaloetum luederitziani ass. nov. the type for this association is relevé 1180, sampled on 16 april 2004 at 21°34’16” s, 20°36’14” e. ten relevés were classified into this association. diagnostic species for this association are oropetium capense, eragrostis echinochloidea, eriocephalus luederitzianus, enneapogon desvauxii, salsola tuberculata, panicum coloratum, cynodon dactylon, jamesbrittenia atropurpurea subsp. atropurpurea, eragrostis truncate and chenopodium petiolariforme. these are constantly associated with eragrostis porosa, hermannia modesta, asparagus cooperi, acacia hebeclada subsp. hebeclada, tragus berteronianus, stipagrostis uniplumis var. uniplumis and grewia flava. a total of 86 species were observed here, with, on average, 27 species per 1000 m2.with a variable, but very low, tree cover and low shrubs between 40 cm and 120 cm in height dominating the landscape, the structure can best be described as a low, semi-open shrubland (figure 3d and figure 4d). this association occurs in omiramba with heavier loamy and clay soils, as well as a pan near otjutuuo in the far north. the soils are often strewn, up to 15%, with calcrete fragments of sizes ranging from gravel to large stones. rocks were even found on one site. the soil surface is moderately to strongly crusted, showing signs of slight sheet erosion. the slopes are generally flat to undulating. these omiramba occur at altitudes of 1180 m.a.s.l. – 1330 m.a.s.l. although the eragrostio echinocloideae–eriocephaloetum luederitziani occasionally has a very high cover of grasses, this is not the norm. the dominant grass species are eragrostis echinochloidea, eragrostis porosa, enneapogon desvauxii, tragus racemosus and stipagrostis uniplumis var. uniplumis. however, only the latter species is a bulk biomass producer. enneapogon desvauxii in particular is a very short grass, about 5 cm high, often forming a fairly dense cover. it is a fast grower, producing leaves and seed within a short period of time (thus the vernacular name ‘agt dae gras’ [eight day grass]). the combination of species – enneapogon desvauxii, eragrostis echinochloidea, eriocephalus luederitzianus and salsola tuberculata – is evident of a harsh microclimate dictated by soil conditions (high ph and/or sodic). the grazing production in this habitat is thus low and the grazing capacity must equally be regarded as low. 5. acacio melliferae–catophractetum alexandri ass. nov. the type for this association is relevé 1260, sampled on 24 april 2004 at 20°35’44” s, 18°05’26” e. twenty-one relevés were classified into this association, which is characterised by the occurrence of catophractes alexandri as diagnostic species. this species also dominates the landscape with an average 20% – 40% cover. it is constantly associated with grewia flava, acacia mellifera subsp. detinens, eragrostis porosa, clerodendrum ternatum, urochloa brachyura, stipagrostis uniplumis var. uniplumis, rhigozum brevispinosum, tragus berteronianus, melinis repens subsp. grandiflora, otoptera burchellii, kleinia longiflora, boscia albitrunca, gisekia africana and aristida congesta subsp. congesta. a total of 132 species were observed in this association, with, on average, 36 species per 1000 m2.the acacio melliferae–catophractetum alexandri is strikingly dominated by more or less dense stands of catophractes alexandri. trees are scarce to totally absent, whilst an average of 40% – 60% shrub cover of between 1 m and 3 m in height is characteristic, thus it can be described as a tall, closed shrubland (figure 3e and figure 4e). hines (1992) described a similar vegetation unit (the catophractes alexandri subassociation of the croton gratissimus–grewia flava association) for the nyae-nyae conservancy. this unit is described as occuring on soils with shallow underlying calcretes. as localised stands of dense catophractes alexandri can be found throughout northern namibia, always on relatively shallow calcrete soils, this community is recognised as a separate association, even though it is closely related to the stipagrostio uniplumis–acacietum melliferae. catophractes alexandri is regarded a typical indicator for petric calcisols (i.e. shallow loamy soils with calcretes) (cole & brown 1976; hines 1992). such soils occur in some of the omiramba, in interdunal valleys and around larger pans and depressions. generally, the slopes are flat to gently undulating, at altitudes of between 1220 m.a.s.l. and 1450 m.a.s.l. the soil surface is often covered, but never more than 2%, with fragments of the underlying calcretes that vary in size from small (gravel) to large (stones). leser (1972) describes similar soil conditions for his catophractes alexandri shrublands along the black nossob. the soil surface was also observed to be moderately to strongly crusted, showing signs of slight sheet erosion. in one case, slight rill erosion was observed. bush encroachment is a common disturbance. the most prominent perennial grasses are stipagrostis uniplumis, aristida congesta, eragrostis rigidior, eragrostis trichophora and cenchrus cilliaris. these are all known to be neither highly palatable, nor indicators of good veld (müller 2007). most grass species are annuals and also typical pioneer or subclimax species. it is, however, commonly believed that the grazing on these calcareous soils is sweeter and more nutritional, thus the quality of grazing is considered moderate. 6. acacio melliferae–hyphaenetum petersianae ass. nov. the type for the acacio melliferae–hyphaenetum petersianae is relevé 1403, sampled on 26 april 2004 at 19°37’50” s, 18°31’58” e. ten relevés were classified into this association. it is characterised by the diagnostic species hyphaene petersiana, enneapogon scoparius, triraphis purpurea, combretum hereroense, peltophorum africanum, sporobolus ioclados, acalypha segetalis, enneapogon desvauxii, limeum arenicolum, aristida stipoides, tragus racemosus, eragrostis annulata, elaeodendron transvaalense, brachiaria deflexa, terminalia prunioides and seddera suffruticosa. these are constantly associated with acacia mellifera subsp. detinens, eragrostis porosa, stipagrostis uniplumis var. uniplumis, pogonarthria fleckii, eragrostis trichophora, aristida adscensionis, urochloa brachyura, grewia flavescens, grewia flava, gisekia africana, geigeria ornativa, commiphora pyracanthoides, asparagus cooperi, aristida rhiniochloa and acacia luederitzii. a total of 112 species were observed here, with, on average, 44 species per 1000 m2.the typical structure is a tall, moderately closed bushland, with a tree layer varying between 7 m and 12 m in height, reaching a cover of 5%, with shrubs covering about 45% of the landscape (figure 3f and figure 4f). the acacio melliferae–hyphaenetum petersianae forms the extensive palmvlakte [‘palm plains’] of the southern grootfontein district, extending into the north-western otjituuo conservancy. but here, in the vicinity of the omuramba omatako, it has a distinct wetland character, thus resembling the parkiesveld found near maroelaboom. the presence of elaeodendron transvaalense, eragrostis aspera and sporobolus iocladus are typical indicators of this wetland character. the plains are gently undulating to undulating, at an altitude of between 1220 m.a.s.l. and 1260 m.a.s.l. the soils are generally shallow to very shallow on calcrete and/or limestone, with a calcrete fragment cover of up to 40% on the soil surface. the surface is also moderately to strongly crusted, occasionally with signs of slight sheet erosion. bush encroachment, often severe, is a common problem here, as it is in the karstveld in general. the grazing condition is low, with only two perennial grasses (eragrostis trichophora and stipagrostis uniplumis) occurring consistently, but at a very low abundance, at the sample sites. some of the annual grasses found here are known to be palatable, but all are low biomass producers, whilst many of them are also shade grasses and difficult to reach under the trees and shrubs where they grow. 7. the panicum gilvum–marsilea vlei community the vleis are represented only by one relevé, which is regarded as inadequate to warrant a formal description of these wetland grasslands as an association. however, for the record, the following is presented: characteristic for this association are the species vahlia capensis, schoenoplectus muricinux, pycreus macrostachyos, panicum gilvum, marsilea nubica var. gymnocarpa, lindernia parviflora, diandrochloa pusilla, cyperus species, portulaca kermesina, dactyloctenium giganteum, kohautia virgata, lindneria clavata, schoenoplectus senegalensis, marsilea macrocarpa, ipomoea hochstetteri, eragrostis rotifer, panicum maximum, ipomoea coptica and eragrostis biflora.the general structure of these vleis is as follows (figure 4g): within the matrix of surrounding vegetation (mostly the stipagrostio uniplumis–acacietum melliferae) is a fringe of dense acacia luederitzii trees, in combination with tall shrubs such as acacia mellifera subsp. detinens, acacia hebeclada subsp. hebeclada and ziziphus mucronata surrounding the water-holding depression. this fringe generally has the nature of a thicket, with a number of shade-loving grasses (e.g. panicum maximum) and various mesophylic to hydrophylic forbs growing in the herb layer. at a point towards the centre of the depression (point of semi-permanent waterlogging?) the thicket fringe stops entirely, making way for a grassland dominated by eragrostis rotifer. various hydrophylic forbs grow in the substratum, for example lindernia parvifolia. as the soil gets moister, various cyperaceae (schoenoplectus, pycreus species), some marsilea species and diandrochloa pusillia occur. these reach well into the standing water. in the shallow standing water, marsilea species dominate, being replaced by panicum gilvum in the deep standing water (figure 4g). occasionally water plants such as aponogeton desertorum, lasiopogon species and even nymphaea species occur in the deeper pans (personal observations in the grootfontein district). here, panicum gilvum is also generally replaced by echionochloa species. in the surveyed pan, however, these were not present. similar vleis have been reported to be widespread throughout the northern kalahari (hines 1993), without any detailed floristic description. these vleis are important temporal water sources and centres of biodiversity (especially faunal diversity) in the landscape. in the dry season, the remains of panicum gilvum and other grasses should form valuable, although very limited, grazing resources. 8. eragrostio rigidioris–urochloaetum brachyurae ass. nov. the type for this association is relevé 1322, sampled on 28 april 2004 at 20°25’10” s, 19°49’18” e. seventeen relevés were classified into this association. diagnostic of this is the occurrence of brachiaria nigropedata, eragrostis rigidior, schmidtia pappophoroides, diospyros lycioides and cynanchum orangeanum. these are constantly associated with urochloa brachyura, stipagrostis uniplumis var. uniplumis, vernonia poskeana, bulbostylis hispidula, aristida congesta subsp. congesta, grewia flava, sesamum capense, otoptera burchellii, melinis repens subsp. grandiflora, eragrostis trichophora, cleome rubella, boscia albitrunca, ziziphus mucronata, pogonarthria fleckii, oxygonum alatum, lonchocarpus nelsii, heliotropium steudneri, gisekia africana, dichrostachys cinerea, clerodendrum ternatum and acacia hebeclada subsp. hebeclada. a total of 125 species were observed here, with, on average, 40 species per 1000 m2.tree cover is virtually absent, whilst the shrub cover is generally absent or very low. if shrubs are present, their cover is below 20%. small dense patches of tarchonanthus camphoratus or acacia hebeclada subsp. hebeclada in particular may occasionally be found, usually where water accumulates during the rainy season. accordingly, this association is best described as a tall, semi-open shrubland with a strong grassland character (figure 3g and figure 4h). the eragrostio rigidioris–urochloaetum brachyurae typically occurs in sandy omiramba crossing through the study area, if not disturbed by overgrazing. the altitude ranges between 1330 m.a.s.l. in the west to 1150 m.a.s.l. in the east. the slopes are generally flat to undulating. soil depth is moderately deep, with moderate to strong soil crusting present. slight sheet erosion is widespread. this association forms a potentially good source of grazing especially because few poisonous plants are present. the biggest threats to these pastures are settlements along the linear pathways through the sandy plains. generally, settlements lead to localised overgrazing in the omiramba, often resulting in the almost exclusive presence of weedy species such as sida cordifolia. 9. tarchonantho camphorati–acacietum eriolobae ass. nov. the type for this association is relevé 1558, sampled on 06 may 2004 at 21°14’14” s, 19°01’16” e. twenty-two relevés were classified into this association, which is characterised by the diagnostic species tylosema esculentum, elephantorrhiza elephantina, ozoroa paniculosa, ipomoea oblongata, dicoma schinzii and limeum fenestratum. these are constanly associated with stipagrostis uniplumis var. uniplumis, grewia flava, dichrostachys cinerea, acacia mellifera subsp. detinens, acacia erioloba, vernonia poskeana, gisekia africana, pogonarthria fleckii, melinis repens subsp. grandiflora, acacia fleckii, urochloa brachyura, eragrostis rigidior, aristida congesta subsp. congesta, otoptera burchellii, bulbostylis hispidula, oxygonum alatum, eragrostis porosa and eragrostis dinteri. a total of 111 species were observed in this association, with, on average, 39 species per 1000 m2.the characteristic acacia erioloba occurs here mainly as tall shrub and occasionally as short to tall trees. thus the typical structure is a short, moderately closed shrubland within the study area (figure 3h and figure 4i). leser (1972) describes an acacia giraffae–terminalia sericea–grewia flava–tarchonanthus camphoratus association for the south-western part of the sandveld – that is, the area south and west of otjinene towards summerdown and steinhausen. here, acacia erioloba (syn. = acacia giraffae) occurs as a dominant tree, typical for the camelthorn savanna sensu giess (1998). it is clear that the relevés sampled from this association within the present study are along the northern fringe, thus not displaying the characteristic structure. the grazing is dominated by the two relatively hard, bulk-producing species, stipagrostis uniplumis var. uniplumis and eragrostis rigidior. schmidtia pappophoroides, digitaria seriata, anthephora pubescens and brachiaria nigropedata are occasionally present as palatable species. grazing trails at the sandveld research station near drimiopsis indicate that the latter three species could contribute a far higher percentage to the grazing production (bester & reed 2003; bester et al. 2003a, 2003b, 2003c). the poisonous species elephantorrhiza elephantina and gnidia polycephala are commonly found, indicating, together with the relatively low occurrence of palatable species, that the grazing within the study area has been degraded and, at present, produces only moderately as subclimax. 10. stipagrostio uniplumis–acacietum melliferae ass. nov. the type for this association is a subassociation, the stipagrostio uniplumis–acacietum melliferae typicum. fifty-nine relevés were classified within this association. this association has no diagnostic species, but is typified by the constant occurrence of acacia mellifera subsp. detinens, grewia flava, urochloa brachyura, boscia albitrunca, stipagrostis uniplumis var. uniplumis, grewia flavescens, tephrosia burchellii, dichrostachys cinerea, rhigozum brevispinosum, ehretia alba, talinum crispatulum, evolvulus alsinoides, acacia luederitzii, melinis repens subsp. grandiflora, eragrostis rigidior, gisekia africana, clerodendrum ternatum, tragus berteronianus, kleinia longiflora and bulbostylis hispidula. based on habitat differences, this association can be subdivided into two subassociations. 10.1. stipagrostio uniplumis–acacietum melliferae rhigozetosum brevispinosi subass. nov. the type for this subassociation is relevé 1377, sampled on 18 april 2004 at 20°53’14” s, 20°03’36” e. thirty relevés were classified into this subassociation, which is characterised by the diagnostic species eragrostis lehmanniana, crotalaria sphaerocarpa and ipomoea magnusiana. these are constantly associated with boscia albitrunca, urochloa brachyura, acacia mellifera subsp. detinens, grewia flava, rhigozum brevispinosum, vernonia poskeana, stipagrostis uniplumis var. uniplumis, tephrosia burchellii, grewia flavescens, eragrostis rigidior, dichrostachys cinerea, oxygonum alatum, ehretia alba, bulbostylis hispidula, melinis repens subsp. grandiflora, evolvulus alsinoides and acacia luederitzii. a total of 141 species were observed here, with, on average, 46 species occurring per 1000 m2.trees do occur in this subassociation, but generally at very low cover. with a high shrub cover, this subassociation is best described as a short, moderately closed bushland (figure 3i and figure 4j). with a highly variable grass cover, this subassociation has a moderate grazing capacity. stipagrostio uniplumis–acacietum melliferae rhigozetosum brevispinosi is a typical ecotonal vegetation type towards sandy soils, fringing the stipagrostio uniplumis–acacietum melliferae typicum on more loamy soils. this is also evident from the species composition, with its numerous psammophyll species. this subassociation occurs on sand drift or ‘hardeveld’ plains (i.e. plains with more loamy soils), with moderately deep to very deep soils and weak surface crusting. bush encroachment is a common disturbance. the altitude of these plains ranges between 1030 m.a.s.l. and 1470 m.a.s.l. slopes vary between flat and undulating, whilst the average annual rainfall varies between 400 mm and 500 mm. 10.2. stipagrostio uniplumis–acacietum melliferae typicum subass. nov. the type for this subassociation is relevé 1285, sampled on 25 april 2004 at 20°06’22” s, 18°31’18” e. twenty-nine relevés were classified into this subassociation, which features no diagnostic species, but can be recognised according to the following constant species: grewia flava, acacia mellifera subsp. detinens, dichrostachys cinerea, stipagrostis uniplumis var. uniplumis, grewia flavescens, tephrosia burchellii, urochloa brachyura, kleinia longiflora, boscia albitrunca, tragus berteronianus, eragrostis porosa, ehretia alba, talinum crispatulum, rhigozum brevispinosum and pogonarthria fleckii. a total of 128 species were observed in this subassociation, with, on average, 38 species per 1000 m2. this subassociaton is the type for stipagrostio uniplumis–acacietum melliferae.although some trees are present, their cover was found to be very variable. however, the shrub component is very consistent, with a cover of 40% – 50% and a height of 3 m – 5 m, thus forming a high, moderately closed bushland sensu edwards (1983) (figure 3j and figure 4k). the stipagrostio uniplumis–acacietum melliferae typicum occurs on plains, interdunal valleys (‘streets’) and sand drift plains throughout the study area. slopes range from flat to undulating. the altitude of these plains ranges from 1120 m.a.s.l. to 1460 m.a.s.l. the soils are mostly sandy and deep, only occasionally with signs of calcretes. the soil surface is normally crusted, ranging from moderate to strong. as in stipagrostio uniplumis–acacietum melliferae rhigozetosum brevispinosi, bush encroachment is also a widespread problem. this subassociation has a very high diversity of grass species.the most consistent and abundant grass species is stipagrostis uniplumis var. uniplumis (average cover 12%). all other grasses with a high abundance are annual species, all regarded as pioneer or subclimax species. a large variety of poisonous plants occur here, with the biggest threat being geigeria ornativa, occurring on 24% of all sites. with this very variable, very pioneer-like grass cover and high number of poisonous plants, grazing is regarded as being moderate. 11. terminalio sericeae–acacietum eriolobae ass. nov. the type for this association is relevé 1544, sampled on 05 may 2004 at 21°02’03” s, 18°20’13” e. seventeen relevés were classified into this association, which is characterised by the diagnostic species bauhinia petersiana subsp. macrantha, terminalia sericea, megaloprotachne albescens and eragrostis dinteri. these are constantly associated with melinis repens subsp. grandiflora, grewia flavescens, acacia mellifera subsp. detinens, urochloa brachyura, stipagrostis uniplumis var. uniplumis, acacia fleckii, tephrosia burchellii, dichrostachys cinerea, boscia albitrunca, grewia flava, acrotome spp., acacia erioloba, rhigozum brevispinosum, pogonarthria fleckii, gisekia africana, eragrostis trichophora, pollichia campestris, kyphocarpa angustifolia, evolvulus alsinoides, eragrostis porosa, dicoma schinzii and commiphora angolensis. a total of 120 species were observed here, with, on average, 42 species per 1000 m2.with acacia erioloba trees being the most prominent species, this association can best be described as a short, moderately closed bushland (figure 3k and figure 4l). the terminalio sericeae–acacietum eriolobae resembles the terminalia sericea–acacia giraffae–grewia flava association described for the north-eastern sandveld area (leser 1972) and is thus a typical example of the camelthorn savanna of the central kalahari sensu giess (1998). the dominance of aristida stipitata and terminalia sericea demonstrates the nearness of this association to that of combreto collini–terminalietum sericeo. these bushlands are to be seen as an ecotone between the broad-leaved savannas to the north and east and the microphyll thornbush savannas to the south and west. this bushland association occurs on gently undulating to undulating plains and sand drift plains and even on interdunal streets at an altitude of between 1020 m.a.s.l. and 1490 m.a.s.l. soil depth varies, but is mostly moderately deep to very deep. surface crusting is variable – from uncrusted to moderately and even strongly crusted. likewise, erosion varies between slight to moderate wind erosion and slight to moderate sheet erosion. the typical poisonous plants for this association are elephantorrhiza elephantina and gnidia polycephala. both are more or less equally frequent. occasionally, both dichapetalum cymosum and geigeria ornativa also occur – the former as remnant from the more mesic broad-leaved savanna, the latter as an indicator of degradation. the medicinal plant harpagophytum procumbens occurs in small patches throughout this vegetation type, but densities are too low to warrant economic exploitation. 12. combreto collini–terminalietum sericeae ass. nov. the type for this association is a subassociation, combreto collini–terminalietum sericeae typicum. a total of 243 relevés were classified into this association. the combreto collini–terminalietum sericeae is characterised by the diagnostic species grewia flava, urochloa brachyura, rhigozum brevispinosum, eragrostis rigidior, acacia mellifera subsp. detinens, merremia verecunda, rhynchosia totta, dicoma schinzii, cleome rubella, acacia fleckii, aristida congesta subsp. congesta, requienia sphaerosperma and indigofera bainesii. these are constantely associated with terminalia sericea, stipagrostis uniplumis var. uniplumis, grewia flavescens, melinis repens subsp. grandiflora, tephrosia burchellii, bauhinia petersiana subsp. macrantha, vernonia poskeana, combretum collinum, lonchocarpus nelsii, xenostegia tridentata subsp. angustifolia, bulbostylis hispidula, megaloprotachne albescens, phyllanthus pentandrus, gisekia africana, commiphora angolensis, acrotome spp., dichrostachys cinerea, oxygonum alatum and digitaria seriata; in absence of the diagnostic species of the burkeo africanae–pterocarpetum angolensis and terminalio sericeae–schinziophytetum rautanenii.this association covers most of the landscape of the eastern communal areas. owing to slight differences in habitat (mainly rainfall variability but also, to a lesser extent, soil depth and loam content of the sand), four subassociations can be recognised. livestock is equally dependant on grazing as on browse. a widespread threat to livestock farming in this area is the poisonous plant dichapetalum cymosum (‘magou’, ‘gifblaar’), which is known to increase under conditions of overgrazing and disturbance (bester 1989; de sousa correia & van rensburg 2000; van eck 2000). 12.1. combreto collini–terminalietum sericeae acacietosum melliferae subass. nov. the type for this subassociation is relevé 1455, sampled on 01 may 2004 at 21°31’43” s, 20°59’55” e. ninety relevés were classified into this subassociation. combreto collini–terminalietum sericeae acacietosum melliferae is characterised by the diagnostic species acacia mellifera subsp. detinens, dicoma schinzii, grewia flava, aristida congesta subsp. congesta and requienia sphaerosperma. these are constantly associated with terminalia sericea, urochloa brachyura, tephrosia burchellii, grewia flavescens, stipagrostis uniplumis var. uniplumis, melinis repens subsp. grandiflora, acacia fleckii, vernonia poskeana, dichrostachys cinerea, bauhinia petersiana subsp. macrantha, bulbostylis hispidula, phyllanthus pentandrus, combretum collinum, gisekia africana, xenostegia tridentata subsp. angustifolia, ipomoea hackeliana, oxygonum alatum, commiphora angolensis, acrotome spp. and phylenoptera nelsii. a total of 153 species were observed here, with, on average, 40 species per 1000 m2.the structure is best described as a short, moderately closed bushland (figure 3l and figures 4m and 4n). the variable grass cover often observed in this subassociation is most likely the result of the severe overgrazing, often resulting in bush encroachment. the grass layer is dominated by the annual megaloprotachne albescens and urochloa brachyura, with stipagrostis uniplumis var. uniplumis and eragrostis trichophora being the main perennial species. at best, the grazing capacity is poor to mediocre. this subassociation forms a patchy ecotone to the thornbush savannas to the south and west, probably because of localised, less favourable soil-moisture regimes (either from higher runoff on the slopes of omiramba, or more xeric soils). the altitude ranges between 1200 m.a.s.l. in the east and north to 1500 m.a.s.l. in the south-west, as this vegetation type gradually changes to stipagrostio uniplumis–acacietum melliferae. 12.2. combreto collini–terminalietum sericeae grewietosum flavae subass. nov. the type for this subassociation is relevé 1439, sampled on 29 april 2004 at 20°05’05” s, 20°29’08” e. forty-five relevés were classified into this subassociation, which is characterised by the diagnostic species grewia flava, clerodendrum ternatum, rhigozum brevispinosum, neorautanenia amboensis, ipomoea bolusiana, talinum crispatulum, indigofera bainesii, eragrostis rigidior, tylosema esculentum, anthephora pubescens and boscia albitrunca. these are constantly associated with urochloa brachyura, stipagrostis uniplumis var. uniplumis, vernonia poskeana, terminalia sericea, melinis repens subsp. grandiflora, oxygonum alatum, phylenoptera nelsii, bulbostylis hispidula, acanthosicyos naudinianus, xenostegia tridentata subsp. angustifolia, tephrosia burchellii, dichrostachys cinerea, commiphora angolensis, grewia flavescens, acrotome spp., schmidtia pappophoroides, megaloprotachne albescens, limeum fenestratum, acacia fleckii and merremia verecunda. a total of 135 species were observed here, with, on average, 41 species per 1000 m2.tree cover is virtually absent, making this subassociation a high, moderately closed shrubland (figure 3m and figure 4o). it occurs on sand drift plains and occasionally in sandy omiramba at an altitude of between 1040 m.a.s.l. and 1450 m.a.s.l. the sands are generally deep to very deep. the most common disturbances observed are bush encroachment and sometimes severe overgrazing. next to dichapetalum cymosum, elephantorrhiza elephantina also occurs occasionally. although this suffrutex is less poisonous to cattle, it is still a threat to grazers. grass cover in this variation is in general low. the annual species megaloprotachne albescens, urochloa brachyura and melinis repens subsp. grandiflora dominate the grass sward. important perennial species are stipagrostis uniplumis var. uniplumis, eragrostis rigidior, eragrostis lehmanniana and eragrostis trichophora. in general, the grazing is regarded as poor to mediocre. 12.3. combreto collini–terminalietum sericeae typicum subass. nov. the type for this subassociation is relevé 1288, sampled on 25 april 2004 at 19°48’14” s, 18°35’45” e. sixty-one relevés were classified into this subassociation, which is characterised by the constant occurrence of terminalia sericea, combretum collinum, grewia flavescens, tephrosia burchellii, stipagrostis uniplumis var. uniplumis, melinis repens subsp. grandiflora, xenostegia tridentata subsp. angustifolia, commiphora angolensis, bauhinia petersiana subsp. macrantha, megaloprotachne albescens, bulbostylis hispidula, acrotome species, vernonia poskeana, phyllanthus pentandrus, phylenoptera nelsii, gisekia africana, digitaria seriata, ochna pulchra, urochloa brachyura, acacia ataxacantha, hibiscus meeusei, tephrosia lupinifolia, dichrostachys cinerea and ipomoea hackeliana. a total of 98 species were observed here, with, on average, 37 species per 1000 m2.combreto collini–terminalietum sericeae typicum only occasionally has trees, making it a short, moderately closed bushland with a strong shrubland character, dominated by terminalia sericea and various combretum species – of which combretum collinum is the most prominent (figure 3n and figure 4p). the sandy habitat occurs on extensive plains ranging between 1020 m.a.s.l. and 1500 m.a.s.l. altitude. the grazing is dominated by the annual megaloprotachne albescens, followed by aristida stipitata. common perennial grass species are stipagrostis uniplumis var. uniplumis and eragrostis lehmanniana. the grazing is thus relatively poor, especially in the dry season, and cattle have to rely on browse for fodder. 12.4. combreto collini–terminalietum sericeae burkeaetosum africanae subass. nov. the type for this subassociation is relevé 1411, sampled on 27 april 2004 at 19°54’48” s, 19°03’59” e. forty-seven relevés were classified into this subassociation, which is characterised by the constant occurrence of terminalia sericea, ochna pulchra, combretum collinum, bauhinia petersiana subsp. macrantha, lonchocarpus nelsii, stipagrostis uniplumis var. uniplumis, megaloprotachne albescens, digitaria seriata, xenostegia tridentata subsp. angustifolia, combretum psidioides, burkea africana, eragrostis pallens, grewia flavescens, phyllanthus pentandrus, gisekia africana, tephrosia burchellii, vernonia poskeana, melinis repens subsp. grandiflora and bulbostylis hispidula. a total of 72 species were observed here, with, on average, 31 species per 1000 m2.characteristic of this subassociation are burkea africana trees amidst a shrubland dominated by terminalia sericea and various combretum species. the structure is thus a typically short, moderately closed bushland (figure 3o and figure 4q), found mostly on sandy plains, occasionally also on dunes, between 1030 m.a.s.l. and 1460 m.a.s.l. altitude. grazing conditions approximate those found in combreto collini–terminalietum sericeae typicum. 13. terminalio sericeae–schinziophytetum rautanenii ass. nov. the type for this association is relevé 4013, sampled on 26 march 1992 at 19°25’17” s, 18°23’20” e. six relevés were classified into this association, which is characterised by the diagnostic species strychnos pungens, schinziophyton rautanenii, erlangea misera, phyllanthus maderaspatensis, bidens spp., croton gratissimus, tricholaena monachne, thunbergia aurea, rhynchosia venulosa, monechma debile, hypoestes forskaolii, rhynchosia sublobata, ximenia americana, hibiscus vitifolius and vangueria infausta. these are constantly associated with terminalia sericea, stipagrostis uniplumis var. uniplumis, ochna pulchra, combretum collinum, grewia flavescens, burkea africana, bauhinia petersiana subsp. macrantha, aristida stipitata and melinis repens subsp. grandiflora. a total of 70 species were observed here, with, on average, 34 species per 1000 m2.terminalio sericeae–schinziophytetum rautanenii has been found on the crests and upper slopes of the high longitudinal dunes just north of the rooiboklaagte omuramba (21°00’ s). additional schinziophyton rautanenii (manketti tree) groves could be identified from a distance at a few locations in the gam and eiseb area. the dunes on which the manketti groves occur vary in altitude between 1240 m.a.s.l. in the south and only 1110 m.a.s.l. further north. long-term average annual rainfall ranges between 400 mm and 500 mm. the deep sands found here (> 150 cm) show only few signs of wind erosion and weak surface crusting, with almost no disturbances. the tall manketti trees associated with a dense understorey shrub layer result in a tall, moderately closed thicket (figure 3p and figure 4r). 14. burkeo africanae–pterocarpetum angolensis ass. nov. the type for this association is relevé 1312, sampled on 27 april 2004 at 19°50’54” s, 18°57’08” e. sixteen relevés were classified within this association, which is characterised by the diagnostic species pterocarpus angolensis, burkea africana, crotalaria flavicarinata, hermannia eenii and rotheca myricoides. these are constantly associated with terminalia sericea, bauhinia petersiana subsp. macrantha, ochna pulchra, combretum collinum, digitaria seriata, phylenoptera nelsii, combretum psidioides, stipagrostis uniplumis var. uniplumis, megaloprotachne albescens, gisekia africana, xenostegia tridentata subsp. angustifolia, panicum kalaharense, aristida stipitata, vernonia poskeana, tephrosia lupinifolia, melinis repens subsp. repens, eragrostis pallens, commelina africana var. krebsiana and bulbostylis hispidula. a total of 61 species were observed here, with, on average, 32 species per 1000 m2.this association occurs mostly in the northern sandy plains of the otjituuo and gam conservancies. patches of this vegetation type can also be found in the northern okamatapati and otjinene conservancies. it is fairly widespread further north and has been described as ‘burkea africana tall closed woodland’ by hines (1992). typically, the habitat consists of extensive sandy plains ranging at altitudes of between 1050 m.a.s.l. and 1340 m.a.s.l. the depth of the sands is generally more than 150 cm, with weak or no surface crusting and signs of only slight wind erosion. typical disturbance factors in these bushlands are frequent fires and bush encroachment. although pterocarpus angolensis (teak tree) is a very prominent species, reaching a height of 12 m – 15 m, neither its density nor its trunk diameter warrants commercial exploitation of this timber resource. few of the other woody species reach tree height, the only ones being burkea africana and, occasionally, terminalia sericea and combretum collinum. owing to the relatively high cover of understorey shrubs, the structure is typically a tall, moderately closed thicket (figure 3q and figure 4s). discussion top ↑ higher syntaxonomic groupings and relations to similar vegetation in southern africa the initial splitting of the dataset between the sandveld and the hardeveld – that is, between the deep sands of the central kalahari basin and the surrounding transitions to the central plateau (de pauw et al. 1998), with more mesic conditions on shallower soils and/or soils with higher loam content – is reflected as between mostly the broad-leaved savannas (associations 12–14) and the microphyll savannas (associations 1–11).the broad-leafed savannas in turn can be split into two major groupings: strohbach and petersen (2007) suggest the recognition of a vegetation class burkeo–pterocarpetea, which is considered representative of the zambesian–baikiaea woodlands ecoregion of the world wildlife fund (wwf) (vetter 2001). this class is typically a semi-open woodland characterised by, ‘… various caesalpinioideae (burkea africana, bauhinia petersiana, occasionally baikiaea plurijuga and guibourtia coleosperma) and papilionoideae (pterocarpus angolensis), as well as combretaceae (combretum spp., terminalia sericea)’ (vetter 2001:n.p). this applies, to a large extent, to both the terminalio sericeae–schinziophytetum rautanenii and the burkeo africanae–pterocarpetum angolensis, although both are relatively species-poor in terms of the caesalpinioideae, compared to the communities described for the central kavango (strohbach & petersen 2007). these two associations thus form the southernmost, most xeric extent of burkeo–pterocarpetea and the zambesian–baikiaea woodlands ecoregion of the wwf (vetter 2001), extending into north-western botswana. bekker and de wit (1991) make no mention of such species combinations in their vegetation description, though. in contrast, the combreto collini–terminalietum sericeae has no woodland character and generally forms a shrubland, at best a bushland. the caesalpinioidae are represented only by bauhinia petersiana subsp. macrantha (a shrub) and occasionally burkea africana (as a short tree). owing to, especially, these structural, but also to compositional differences, this association is to be recognised as a separate higher syntaxon, possibly a class combreto–terminalietea sericeae. the fact that the subdivisions of combreto collini–terminalietum sericeae are recognised only as subassociations is because of the very close compositional relationship between them and this needs to be confirmed through further studies. this association covers most of the landscape of the eastern communal areas and is expected to make a significant contribution to the landscape of the kalahari acacia–baikiaea woodlands as defined by the wwf (spriggs 2001a). bekker and de wit (1991) mention such vegetation occurs west of the ghanzi ridge, whilst cole and brown (1976) describe such vegetation for the kalahari sand plain south and west of the ghanzi ridge. similar vegetation has also been found elsewhere in namibia, in particular in the maroelaboom area (e.g. at the sonop observatory – jürgens et al. 2010) and in north-central namibia (kangombe 2010). this vegetation type seems to have resemblances to the central sandy bushveld (svcb 12) of south africa (mucina & rutherford 2006). the central sandy bushveld includes the nylsvley study area, which was subject to extensive ecosystem research (scholes & walker 1993). the subdivision of the hardeveld is not as clear-cut. it is speculated that the acacio fleckii–terminalietum prunioidis and the acacio melliferae–hyphaenetum petersianae could potentially form a separate class. this is also suggested by giess (1998), who recognised the karsteveld as separate from the thornbush savanna. similar vegetation, especially to acacio fleckii–terminalietum prunioidis, has been described by cole and brown (1976) for the ghanzi ridge area north-east of ghanzi towards the ngamiland border. the eragrostis echinocloideae–eriocephaloetum luederitziani and the acacio melliferae–catophractetum alexandri form typical niche vegetation related to shallow carbonate-rich soils. there are obvious resemblances between these two associations, warranting their own higher syntaxonomic unit. acacio melliferae–catophractetum alexandri, especially, occurs widespread throughout eastern and north-eastern namibia (hines 1992; leser 1972), but also in north-western botswana (cole & brown 1976). it could not be established whether similar vegetation exists within the kalahari bushveld types of south africa (mucina & rutherford 2006). the remainder of the microphyll savannas are typical for the acacietea (volk & leippert 1971); that is, they are dominated by various species of the genus acacia. the acacietea are divided by both giess (1998) and the wwf into two larger units, these being the thornbush savanna (roughly equivalent to the namibian savanna woodlands – spriggs 2001b) and the camelthorn savanna, which forms the northern part of the kalahari xeric savanna (seymour 2001). typical camelthorn savanna elements are the terminalio sericeae–acacietum eriolobae and the tarchonantho camphorate–acacietum eriolobae. whereas the former association seems to occur widespread in western botswana (cole & brown 1976) and has been described also from the sandveld research station north of gobabis in namibia (jürgens et al. 2010; september 2006), tarchonantho camphorate–acacietum eriolobae has resemblances to the kimberley thornveld (svk 4) or kuruman thornveld (svk 9) of south africa (mucina & rutherford 2006). vegetation mapping the initial vegetation map prepared by strohbach et al. (2004) was refined by hüttich et al. (2009) and is presented in that publication. for the mapping of the vegetation types, a number of associations had to be combined, in a similar way as was undertaken for the soter methodology (fao 1995) or the landscapes of the kalahari gemsbok national park (van rooyen et al. 2008). for purposes of comparison, the mapping units described by hüttich et al. (2009) are related to the vegetation associations described in table 1 in this article. table 1: association names as used in the present study, as related to the community names used in the original 2004 study and the 2009 vegetation mapping units. land use and land degradation large parts of the vegetation have been found to be either over-utilised, subject to regular heavy fires, or bush encroached. in general, bush encroachment in these areas is regarded as high (de klerk 2004), thus further threatening the grazing resources. with this in mind, many farmers resort to the harvesting of droppers (fence posts) from terminalia sericea as an extra source of income. overgrazing, especially if combined with overutilisation of this woody resource and/or frequent fires in the area, will lead to an increased exposure of the soil surface to both wind erosion and denitrification, the latter as a result of increased heating of the surface during the day (schlesinger & peterjohn 1991; schlesinger et al. 1996). the kalahari sands are, in any case, extremely nutrient-poor (dougill & thomas 2004; wang et al. 2007) – such a denitrification cycle will lead to further reduction in the soil nutrient status and thus to further reduction in the vegetation cover. this is a classic example of ‘biological feedback’ into desertification (schlesinger et al. 1990), in an ecosystem which is, in any case, predicted to aridify and turn into a mobile sand sea over the next half century as a result of global climate change (midgley et al. 2005; thomas, knight & wiggs 2005). this will mean an even more rapid resource loss over the entire area, resulting in major socioeconomic problems for the country. to ensure sustainable land use in the area, it is essential that good vegetative cover be maintained, whether dominated by grass, shrub or tree. conclusion top ↑ the vegetation within the eastern communal conservancies of namibia forms a large expanse of transitional vegetation between the xeric southern kalahari and the woodlands of the northern kalahari, with adjacent transitions to the central plateau with its stonier and/or loamier soils. as the soils are generally nutrient-poor (dougill & thomas 2004; wang et al. 2007), the entire ecosystem is extremely fragile and susceptible to desertification (thomas et al. 2005). in order to prevent this, proper land-use planning is essential for the emerging and established communal conservancies. this article provides important baseline information needed for proper integrated land-use planning, also in line with the national land reform programme of the government of namibia (haub 2009).this article also provides an important contribution towards the inventorying of vegetation in namibia under the vegetation survey of namibia project (strohbach 2001; strohbach & jürgens 2010). the newly described vegetation associations represent four important ecoregions in namibia, being the forest savannas and woodlands, the karstveld, the thornbush savanna and the camelthorn savannah sensu giess (1998). the higher order syntaxonomy of these associations is still unclear, mainly because only limited similar studies exist within namibia. this makes an assignment to a specific syntaxonomic rank impossible. future studies should concentrate on the description of adjacent areas especially within the karstveld, thornbush and camelthorn savannas, to confirm and expand the classification results of the hardeveld. as more such studies become available, a better understanding of the higher order syntaxa will become available. a phytosociological study to the seasonal wetlands of the kalahari, building on pioneer work by hines (1993), will assist to deepen the understanding of these azonal features, which also form a major habitat in this ecoregion. there is an urgent need for an online register of vegetation associations for southern africa, as has been suggested by brown et al. (2013). this will facilitate the proper description and comparison of vegetation types in relation to similar types and ecozones in southern africa. acknowledgements top ↑ thanks are due to ms marianne strohbach for her help with the fieldwork. the assistance of the staff of the national herbarium of namibia in the identification of plant specimens is also gratefully acknowledged. this project was co-funded by the global environment facility through the desert margins programme and the government of namibia, through the recurrent budget of the directorate agriculture research and training. competing interests the author declares that he has no financial or personal relationships which may inappropriately have influenced him in writing this article. references top ↑ bekker, r.p. & de wit, p.v., 1991, ‘contribution to the vegetation classification of botswana, field document 34’, food and agriculture organization of the united nations, united nations development programme and government of botswana, gaborone.bester, f.v., 1988, ‘die bepaling van die grasproduksie van natuurlike veld [the determination of grass production off natural veld]’, agricola 6, 26–30. bester, f.v., 1989, ‘navorsing gedoen met betrekking tot gifblaar dichapetalum cymosum [research done in regards of gifblaar dichapetalum cymosum]’, agricola 7, 60–64. bester, f.v. & reed, e.r., 2003, ‘grass yield in the sandveld camel thorn savanna: rainfall vs stocking rate’, spotlight on agriculture 71, ministry of agriculture, water and rural devlopment, windhoek. bester, f.v., van eck, j.a.j., kolling, h. & van rooyen, b., 2003a, ‘the influence of stocking rate on the grass yield in the camel thorn savanna’, spotlight on agriculture 70, ministry of agriculture, water and rural devlopment, windhoek. bester, f.v., van eck, j.a.j., kolling, h. & van rooyen, b., 2003b, ‘grazing capacity in the sandveld camel thorn savanna of namibia’, spotlight on agriculture 72, ministry of agriculture, water and rural devlopment, windhoek. bester, f.v., van eck, j.a.j., kolling, h. & van rooyen, b., 2003c, ‘the influence of stocking rate on the distribution of individual grass species in the sward’, spotlight on agriculture 74, ministry of agriculture, water and rural devlopment, windhoek. botha, l., 1998, ‘history of drought in namibia’, agricola 10, 5–12. brown, l.r., du preez, p.j., bezuidenhout, h., bredenkamp, g.j., mostert, t.h., collins, n.b. et al., 2013, ‘guidelines for phytosociological classifications and descriptions of vegetation in southern africa’, koedoe 55, 1103. http://dx.doi.org/10.4102/koedoe.v55i1.1103 bruelheide, h., 1997, ‘using formal logic to classify vegetation’, folia geobotanica phytotaxonomia 32, 41–46. http://dx.doi.org/10.1007/bf02803883 bruelheide, h. & flintrop, t., 1994, ‘arranging phytosociological tables by species-relevé groups’, journal of vegetation science 5, 311–316. http://dx.doi.org/10.2307/3235854 burke, a. & strohbach, b.j., 2000, ‘review: vegetation studies in namibia’, dinteria 26, 1–24. chytrý, m., tichý, l., holt, j. & botta-dukat, z., 2002, ‘determination of diagnostic species with statistic fidelity measures’, journal of vegetation science 13, 79–90. http://dx.doi.org/10.1111/j.1654-1103.2002.tb02025.x cole, m.m. & brown, r.c., 1976, ‘the vegetation of the ghanzi area of western botswana’, journal of biogeography 3, 169–196. http://dx.doi.org/10.2307/3038009 de klerk, j.n., 2004, ‘bush encroachment in namibia. report on phase 1 of the bush encroachment research, monitoring and management project’, ministry of environment and tourism, windhoek. de pauw, e., coetzee, m.e., calitz, a.j., beukes, h. & vits, c., 1998, ‘production of an agro-ecological zones map of namibia (first approximation). part ii: results’, agricola 10, 33–43. de sousa correia, r.j. & van rensburg, l., 2000, ‘geographical distribution and local occurence of dichapetalum cymosum (hook) engl. (gifblaar) in namibia and surrounding areas’, agricola 11, 19–28. department landbou tegniese dienste, 1979, ‘die afbakening van redelike homogene boerderygebiede van die noordelike en sentrale substreke van s.w.a. met die heersende knelpunte en beoogde ontwikkleingsprogramme vir die verskillende bedryfstakke [the delimitation of relative homogenous farming areas in the northern and central subregions of swa with their constraints and planned development programmes for the different branches]’, department landbou tegniese dienste, windhoek. dengler, j., jansen, f., glöckler, f., peet, r.k., de cáceres, m., chytrý, m. et al., 2011, ’the global index of vegetation-plot databases (givd): a new resource for vegetation science’, journal of vegetation science 22, 582–597. http://dx.doi.org/10.1111/j.1654-1103.2011.01265.x dougill, a.j. & thomas, d.s.g., 2004, ‘kalahari sand soils: spatial heterogeneity, biological soil crusts and land degradation’, land degradation & development 15, 233–244. http://dx.doi.org/10.1002/ldr.611 edwards, d., 1983, ‘a broad-scale structural classification of vegetation for practical purposes’, bothalia 14, 705–712. espach, c., lubbe, l.g. & ganzin, n., 2006, ‘determining grazing capacity in namibia: approaches and methodologies’, agricola 16, 28–39. espach, c., lubbe, l.g. & ganzin, n., 2010, ’determining grazing capacity in namibia with the aid of remote sensing’, agricola 20, 41–61. food and agricultural organization (fao), 1995, ‘global and national soils and terrain digital databases (soter)’, world soil resources reports 74, rev. 1, land and water development division, food and agriculture organization of the united nations, rome. geological survey, 1980, south west africa/namibia geological map 1:1000000, geological survey of the republic of south africa and south west africa/namibia, windhoek. giess, w., 1998, ‘a preliminary vegetation map of namibia’, dinteria 4, 1–112. haub, o., 2009, ‘understanding of land-use planning and its relevance in namibia’, namibia land management ser. 1, namibia institute for democracy and ministry of lands and resettlement, windhoek. hennekens, s.m. & schaminée, j.h.j., 2001, ‘turboveg, a comprehensive data base management system for vegetation data’, journal of vegetation science 12, 589–591. http://dx.doi.org/10.2307/3237010 hines, c.j.h., 1992, ‘an ecological study of the vegetation of eastern bushmanland (namibia) and its implications for development’, msc thesis, institute of natural resources, university of natal, pietermaritzburg. hines, c.j.h., 1993, ‘temporary wetlands of bushmanland and kavango; northeast namibia’, madoqua 18, 57–69. hüttich, c., gessner, u., herold, m., strohbach, b.j., schmidt, m., keil, m. et al., 2009, ‘on the suitability of modis time series metrics to map vegetation types in dry savanna ecosystems: a case study in the kalahari of ne namibia’, remote sensing 1, 620–643. http://dx.doi.org/10.3390/rs1040620 jürgens, n., haarmeyer, d.h., luther-mosebach, j., dengler, j., finckh, m. & schmiedel, u. (eds.), 2010, patterns at local scale: the biota observatories, biodiversity in southern africa, klaus hess publishers, göttingen & windhoek. kangombe, f.n., 2010, ‘the vegetation of omusati and oshana regions, central-northern namibia’, msc thesis, department of plant sciences, university of pretoria. king, l.c., 1963, south african scenery, hafner publishing company, new york. klaassen, e.s. & kwembeya, e.g. (eds.), 2013, ‘a checklist of namibian indigenous and naturalised plants’, occasional contributions 5, national botanical research institute, windhoek. leser, h., 1972, ’geoökologische verhältnisse der pflanzengesellschaften in den savannen des sandveldes um den schwarzen nossob und epukiro [geo-ecological relationships of different plant communities in the sandveld savannas at the black nossob and epukiro]’, dinteria 6, 1–44. lubbe, l.g., 2005, ‘towards an updated carrying capacity map for namibia: a review of the methodologies currently used to determine carrying capacity in namibia’, agricola 15, 33–39. mendelsohn, j. & el obeid, s., 2003, sand and water. a profile of the kavango region, struik publishers, cape town. mendelsohn, j., jarvis, a., roberts, c. & robertson, t., 2002, atlas of namibia, david phillips publishers, cape town. mendelsohn, j.m. & el obeid, s., 2002, communal lands of eastern namibia, raison and nnf, windhoek. midgley, g., hughes, g., thuiller, w., drew, g. & foden, w., 2005, ‘assessment of potential climate change impacts on namibia’s floristic diversity, ecosystem structure and function’, climate change research group, south african national biodiversity institute for the namibian national biodiversity programme, directorate of environmental affairs, cape town. mucina, l. & rutherford, m.c. (eds.), 2006, ‘the vegetation of south africa, lesotho and swaziland’, strelitzia 19, south african national biodiversity institute, pretoria. müller, m.a.n., 2007, grasses of namibia, 2nd edn., ministry of agriculture, water and forestry, windhoek. namibian agricultural resources information system, 2001, ‘namibian agricultural resources information system (naris), agro-ecological zoning program’, ministry of agriculture, water and rural development, windhoek. nambian association of community based natural resource management (cbnrm) support organisations, 2011, nacso: conservancies – by name, viewed 18 december 2011, from http://www.nacso.org.na/soc_profiles/conservancylist.php roleček, j., tichý, l., zelený, d. & chytrý, m., 2009, ‘modified twinspan classification in which the hierarchy respects cluster heterogeneity’, journal of vegetation science 20, 596–602. http://dx.doi.org/10.1111/j.1654-1103.2009.01062.x schlesinger, w.h. & peterjohn, w.t., 1991, ‘processes controlling ammonia volatilization from chihuahuan desert soils’, soil biological biochemistry 23, 637–642. http://dx.doi.org/10.1016/0038-0717(91)90076-v schlesinger, w.h., raikes, j.a., hartley, a.e. & cross, a.f., 1996, ‘on the spatial pattern of soil nutrients in desert ecosystems’, ecology 77, 364–374. http://dx.doi.org/10.2307/2265615 schlesinger, w.h., reynolds, j.f., cunningham, g.l., huenneke, l.f., jarell, w.m., virginia, r.a. et al., 1990, ‘biological feedbacks in global desertification’, science 247, 1043–1048. http://dx.doi.org/10.1126/science.247.4946.1043 scholes, r.j. & walker, b.h., 1993, an african savanna. synthesis of the nylsvley study, cambridge university press, cambridge. http://dx.doi.org/10.1017/cbo9780511565472 september, z.m., 2006, mapping the vegetation of the sandveld research station in namibia with the aid of remote sensing, bsc hons mini dissertation, department of plant sciences, university of pretoria. seymour, c., 2001, kalahari xeric savanna (ecoregions/wwf), viewed 23 april 2010, from http://www.worldwildlife.org/ecoregions/at1309 spriggs, a., 2001a, kalahari acacia–baikiaea woodlands (ecoregions/wwf), viewed 23 april 2010, from http://www.worldwildlife.org/ecoregions/at0709 spriggs, a., 2001b, namibian savanna woodlands (ecoregions/wwf), viewed 23 april 2010, from http://www.worldwildlife.org/ecoregions/at1316 steynberg, f., 2011, ‘nuwe bewaringsgebiede open in omahekestreek [new conservation areas open in omaheke region]’, republikein, 01 july, viewed 17 december 2011, from  http://www.republikein.com.na/politiek-en-nasionale/ toerisme-en-omgewingsake/nuwe-bewaringsgebiede-open-in-omahekestreek.130727 strohbach, b.j., 2001, ‘vegetation survey of namibia’, journal of the namibia scientific society 49, 93–124. strohbach, b.j., 2012, ‘providing relevant, useful information on namibian vegetation types’, agricola 22, 7–39. strohbach, b.j. & jürgens, n., 2010, ‘towards a user-friendly vegetation map of namibia: ground truthing approach to vegetation mapping’, in u. schmiedel & n. jürgens (eds.), patterns and processes at regional scale, biodiversity in southern africa, pp. 46–56, klaus hess publishers, göttingen. strohbach, b.j. & kangombe, f., 2012, ‘national phytosociological database of namibia’, biodiversity & ecology 4, 298. http://dx.doi.org/10.7809/b-e.00095 strohbach, b.j. & petersen, a., 2007, ‘vegetation of the central kavango woodlands in namibia: an example from the mile 46 livestock development centre’, south african journal of botany 37, 391–401. http://dx.doi.org/10.1016/j.sajb.2007.03.002 strohbach, b.j., strohbach, m., kutuahuripa, j.t. & mouton, h.d., 2004, ‘a reconnaissance survey of the landscapes, soils and vegetation of the eastern communal areas (otjiozondjupa and omaheke regions), namibia’, unpublished report for the desert research foundation of namibia and the desert margins programme, national botanical research institute, windhoek. strohbach, b.j. & kutuahuripa, j.t., submitted, ‘vegetation of the eastern communal conservancies in namibia: ii. environmental drivers’, koedoe. thomas, d.s.g., 1988, ‘the nature and depositional setting of arid and semi-arid kalahari sediments, southern africa’, journal of applied ecology 14, 17–26. thomas, d.s.g., knight, m. & wiggs, g.f.s., 2005, ‘remobilization of southern african desert dune systems by twenty-first century global warming’, nature 435, 1218–1221. http://dx.doi.org/10.1038/nature03717 tichý, l., 2002, ‘juice, software for vegetation classification’, journal of vegetation science 13, 451–453. http://dx.doi.org/10.1111/j.1654-1103.2002.tb02069.x tichý, l. & chytrý, m., 2006, ‘statistical determination of diagnostic species for site groups of unequal size’, journal of vegetation science 17, 809–818. http://dx.doi.org/10.1111/j.1654-1103.2006.tb02504.x van eck, j.a.j., 2000, ‘control of gifblaar (dichapetalum cymosum) at oshimati’, agri-info 5, 14–16. van rooyen, m.w., van rooyen, n., bothma, j. du p. & van den berg, h.m., 2008, ‘landscapes in the kalahari gemsbok national park, south africa’, koedoe 50, 99. http://dx.doi.org/10.4102/koedoe.v50i1.154 vetter, s., 2001, zambezian–baikiaea woodlands (ecoregions/wwf), viewed 23 april 2010, from http://www.worldwildlife.org/ecoregions/at0726 volk, o.h. & leippert, h., 1971, ‘vegetationsverhältnisse im windhoeker bergland, südwestafrika [vegetation relations in the windhoek mountainveld]’, journal der s.w.a. wissenschaftliche gesellschaft xxv, 5–44. wang, l., d’odorico, p., ringrose, s., coetzee, s. & macko, s.a., 2007, ‘biogeochemistry of kalahari sands’, journal of arid environments 71, 259–272. http://dx.doi.org/10.1016/j.jaridenv.2007.03.016 weaver, l.c. & petersen, t., 2008, ‘namibia communal area conservancies’, in best practises in sustainable hunting. a guide to best practices from around the world, cic technical series, pp. 48–52, cic–international council for game and wildlife conservation and fao, budakeszi. weaver, l.c. & skyer, p., 2005, ‘conservancies: integrating wildlife land-use options into the livelihood, development, and conservation strategies of namibian communities’, in conservation and development interventions at the wildlife/livestock interface: implications for wildlife, livestock and human health. iucn, gland, switzerland, paper presented at the fifth world parks congress, durban, 08–17 september 2003. weber, h.e., moravec, j. & theurillat, j.p., 2000, ‘international code of phytosociological nomenclature. 3rd edition’, journal of vegetation science 11, 739–768. westfall, r.h., dednam, g., van rooyen, n. & theron, g.k., 1982, ‘phytotab: a program package for braun-blanquet tables’, vegetatio 49, 35–37. http://dx.doi.org/10.1007/bf00051562 article information authors: robert f. brand1 leslie r. brown1 pieter j. du preez2 affiliations: 1applied behavioural ecology and ecosystem research unit, university of south africa, south africa 2department of plant sciences, university of the free state, south africa correspondence to: leslie brown email: lrbrown@unisa.ac.za postal address: private bag x6, florida 1710, south africa dates: received: 28 sept. 2010 accecpted: 07 sept. 2011 published: 10 nov. 2011 how to cite this article: brand, r.f., brown, l.r. & du preez, p.j., 2011, ‘the grassland vegetation of platberg, eastern free state, south africa’, koedoe 53(1), art. #1027, 13 pages. http://dx.doi. org/10.4102/ koedoe.v53i1.1027 copyright notice: © 2011. the authors. licensee: aosis openjournals. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) the grassland vegetation of platberg, eastern free state, south africa in this original research... open access • abstract • introduction    • study area • materials and methods • results    • vegetation classification and description       • 1. themeda triandra – cyanotis speciosa major community       • 1.1 cheilantes ecklonia – crassula sarcocaulis community       • 1.2 hyparrhenia hirta – tristachya leucothrix community       • 1.2.1 hyparrhenia hirta – leonotis ocymifolia subcommunity       • 1.2.2 selago galpinii – hyparrhenia hirta subcommunity       • 1.2.3 felicia filifolia – diospyros austro-africana subcommunity       • 1.2.4 hyparrhenia hirta – polygala hottentotta subcommunity       • 1.3 eragrostis caesia – ipomoea crassipes community       • 1.4 cyanotis speciosa – leonotis ocymifolia community       • 2. agapanthus campanulatus – aristida monticola major community       • 2.1 andropogon appendiculatus – helictotrichon longifolium community       • 2.1.1 cineraria eriodioides – senecio inornatus subcommunity       • 2.1.2 tristachya leucothrix – koeleria capensis subcommunity       • 2.1.3 andropogon appendiculatus – pennisetum sphacelatum subcommunity       • 2.1.4 watsonia lepida – helichrysum pallidum subcommunity       • 2.1.5 ledebouria cooperi – selago galpinii subcommunity       • 2.1.6 andropogon appendiculatus – senecio pauciculatus subcommunity       • 2.1.7 lotononis lotononoides – hesperantha baurii subcommunity       • 2.1.8 habenaria lithophyla – eragrostis racemosa subcommunity       • 2.2 muraltia saxicola – helichrysum chionosphaerum community       • 2.2.1 muraltia saxicola – aristea woodii subcommunity       • 2.2.2 muraltia saxicola – helichrysum aureum subcommunity       • 2.2.3 ficinia stolonifera – muraltia saxicola subcommunity       • 2.3 andropogon distachyos – helichrysum aureum community       • 2.4 passerina montana – melinis nerviglumis community       • 2.5 cyperus semitrifidus – digitaria monodactyla community       • 2.5.1 digitaria monodactyla – selaginella caffrorum subcommunity       • 2.5.2 eriospermum ornithogaloides – digitaria monodactyla subcommunity       • 2.5.3 digitaria monodactyla – crassula sarcocaulis subcommunity    • species richness and structure • discussion    • floristic affinities    • conservation • conclusion • acknowledgements    • authors’ contributions • references abstract (back to top) platberg is an inselberg that presents a refuge for indigenous plants and animals. uncontrolled human access to this area threatens this sensitive ecosystem. the vegetation of platberg was investigated to obtain an inventory of the different plant species and communities present in this area. a hierarchical classification, a description and an ecological interpretation of the grassland communities of platberg are presented. a total of 169 sample plots were placed on a stratified random basis within the study area. from a twinspan classification a total of 27 different plant communities, which can be grouped into two major community types, nine communities, 18 sub-communities and six variants, were identified. a significant difference in species richness was found between the two major communities, with the higher-altitude communities having a higher species richness than the communities on the lower-lying slopes. a total of 26 endemic or near endemic drakensberg alpine centre species were recorded. conservation implications: anthropogenic influences are felt globally on ecosystems. high-altitude habitats and organisms will be affected first. inselbergs have high levels of endemic organisms and are reservoirs and refugia for unique genetic material. this grassland plant community survey of platberg provides valuable information on inselberg ecology for conservation planning. introduction (back to top) the grassland biome of southern africa is continually under threat from anthropoidal activities such as crop production, overgrazing, residential and industrial development, and invasion by alien plant species. these pressures threaten the biodiversity of this sensitive biome (van wyk & smith 2001). the prevention of large-scale loss of biodiversity is a daunting challenge facing conservationists worldwide (ed. huntley 1994). in order to conserve our natural resources it is important that ecosystems are studied to compile an inventory of the different plant and animal species present. platberg (near harrismith in the eastern free state) is an inselberg located within the grassland biome. the vegetation comprises a mosaic of woody and grass-dominated vegetation units with short to tall sour grasses constituting the majority of the species composition (mucina & rutherford 2006). the korannabergnear excelsior (also in the eastern free state) is the only other inselberg where a phytosociological study was performed, by du preez (1991, 1992), du preez and bredenkamp (1991) and du preez, bredenkamp and venter (1991). studies of the grassland vegetation of inselbergs in africa are limited to studies in west africa by porembski and barthlott (1995), porembski and brown (1995), porembski et al. (1996) and porembski et al. (1998), in central atlantic africa by parmentier et al. (2006),and in namibia by burke (2001, 2004). the lack of data of inselberg vegetation can be attributed to the physical difficulty of accessing inselbergs. this inaccessibility also contributes to the unique biodiversity found on these high-altitude areas (van wyk & smith 2001). owing to the proposed commercial development on and around platberg, as well as the aims of the maloti drakensberg transfrontier project, it was considered necessary to undertake a more detailed plant ecology survey to describe and classify the vegetation of platberg. this paper aims to classify and describe the grassland vegetation present on the platberg plateau and slopes. study area platberg (29°10’s, 28°16’e) is located along the n3 highway between johannesburg and durban. it forms a distinctive flat-topped, l-shaped plateau with clear-cut contact between the dolerite plateau and the cave sandstone of the clarens formation (figure 1). the soil is generally shallow and weakly developed lithosols, with plains soil being mostly sandy loam containing boulders and gravel (hoare & bredenkamp 2001; mucina & rutherford 2006). altitudes range from 1900 m.a.s.l. to 2394 m.a.s.l. with a surface area of approximately 3000 ha for the study site. the slopes are steep, with numerous vegetated gullies and boulder scree slopes below the vertical cliffs. a permanent stream drains off the southern side and flows off the escarpment in a steeply incised gully (figure 2). rainfall is seasonally, with the most rain falling between november and march. mean annual precipitation for the drakensberg is 739 mm, with platberg’s eastern summit receiving up to 1200 mm (sa weather services 2007). the area has a cold temperature regime (june and july being the coldest months), with a mean annual temperature of 9.6 °c. minimum and maximum temperatures range from –2 °c (july) to 24 °c (january), respectively. figure 1: location of the study area. figure 2: topography of platberg. materials and methods (back to top) the scope of the study was to sample above the 1800-m contour in order to work within the limits set by killick (1963, 1978), who regarded the region in the drakensberg above 1800 m as a distinct floristic region called the afro-alpine region. plot sizes were fixed at 16 m² (du preez & bredenkamp 1991; malan 1998). after a recognisance of the study area, a total of 169 sample plots were placed on a stratified-random basis within representative stands of vegetation to exclude as much heterogeneity as possible with regard to floristic composition, structure and habitat. the general position of the sample plots was pre-determined according to aerial photographs but each section within the identified stand had the same chance of being selected. chosen sample plots found not to be representative of the general vegetation were moved to be more representative, in accordance with the requirements for braun-blanquet surveys (mueller-dombois & ellenberg 1974; westhoff & van der maarel 1980). all species present in sample plots, except bryophytes and lichens, were recorded. taxon names conform to those of germishuizen and meyer (2003). over 1500floristic collections were made and identified at either the geo-potts herbarium (university of the free state, bloemfontein) or the national herbarium in pretoria. the specimens are currently housed at the geo-potts herbarium at the university of the free state. the cover or abundance of each species was estimated using the modified 9-degree braun-blanquet sampling scale (keddy 2007; kent & coker 1992). the surveys were conducted over a period of 24 months (february 2005 to january 2007). environmental data collected included geology (johnson et al. 2006), soil type (soil classification working group 1991), aspect, exposure, size of rocks, altitude, percentage of the area covered by rock, and topography. erosion and slope were estimated as described by brand, du preez and brown (2008). additional notes were made on management and utilisation of the area. floristic data were captured on the floristic database turboveg (hennekens 1996a, 1998). the data were then exported to megatab (hennekens 1996b), followed by the use of twinspan (a multivariate classification technique) to classify the different plant communities. further refinement was achieved by the repositioning of species and relevés. plant communities were recognised by using diagnostic and/or characteristic species as defined by westhoff and van der maarel (1980) andkent and coker (1992). species were considered to be diagnostic if they were present in at least 40% of the relevés of a specific plant community, with a constancy of 20% or less in other communities. in exceptional cases, species with a constancy of between 30% and 40% were included if they had a constancy of less than 5% in other communities. the results are presented in a phytosociological table (see online supplement). in order to compare the mean species abundance of the two major communities, a parametric t-test for homogeneous variances was used as the community species variances were not significantly different (f = 1.111, p > 0.05). the nonparametric kruskal–wallis analysis of variance (anova) by ranks was used to compare species abundance within major communities. data were analysed using statistica version 9.0 (statistica 2009). figure 3: species abundance of the communities within major community 1. figure 4: species abundance of the communities within major community 2. results (back to top) vegetation classification and description the hierarchical classification compiled from 169 relevés resulted in the identification of 27 different plant communities that can be grouped into two major community types, nine communities, eighteen sub-communities and six variants, as listed below. the data set contains 6 trees, 62 shrubs, 55 grasses, 23 succulents,19 sedges, 85 geophytes, 16 ferns, and 310 forb plant species. a total of 26 drakensberg alpine centre (dac) endemics and 22 alien plant species were found to be present (brand, du preez & brown 2010). the variants are discussed in detail in brand (2007) and are only listed in this paper. all phytosociological descriptions in this paper refer to the table provided the online supplement mentioned earlier. 1. themeda triandra – cyanotis speciosa major community    1.1 cheilanthes eckloniana – crassula sarcocaulis community    1.2 hyparrhenia hirta – tristachya leucothrix community       1.2.1 hyparrhenia hirta – leonotis ocymifolia subcommunity       1.2.2 selago galpinii – hyparrhenia hirta subcommunity       1.2.3 felicia filifolia – diospyros austro-africana subcommunity       1.2.4 hyparrhenia hirta – polygala hottentotta subcommunity    1.3 eragrostis caesia – ipomoea crassipes community    1.4 cyanotis speciosa – leonotis ocymifolia community 2. agapanthus campanulatus – aristida monticola major community    2.1 andropogon appendiculatus – helictotrichon longifolium community       2.1.1 cineraria eriodioides – senecio inornatus subcommunity           2.1.1.1 senecio inornatus variant           2.1.1.2 ficinia stolonifera variant       2.1.2 tristachya leucothrix – koeleria capensis subcommunity       2.1.3 andropogon appendiculatus –pennisetum sphacelatum subcommunity           2.1.3.1 nemesia caerulea variant           2.1.3.2 tristachya leucothrix variant       2.1.4 watsonia lepida – helichrysum pallidum subcommunity       2.1.5 ledebouria cooperi – selago galpinii subcommunity       2.1.6 andropogon appendiculatus –senecio paucicalyculatus subcommunity       2.1.7 lotononis lotononoides – hesperantha baurii subcommunity           2.1.7.1 aster bakeranus variant           2.1.7.2 erica alopecurus variant       2.1.8 habenaria lithophyla – eragrostis racemosa subcommunity    2.2 muraltia saxicola – helichrysum chionosphaerum community       2.2.1 muraltia saxicola – aristea woodii subcommunity       2.2.2 muraltia saxicola – helichrysum aureum subcommunity       2.2.3 ficinia stolonifera – muraltia saxicola subcommunity    2.3 andropogon distachyos – helichrysum aureum community    2.4 passerina montana – melinis nerviglumis community    2.5 cyperus semitrifidus – digitaria monodactyla community       2.5.1 digitaria monodactyla – selaginella caffrorum subcommunity       2.5.2 eriospermum ornithogalum – digitaria monodactyla subcommunity       2.5.3 digitaria monodactyla – crassula sarcocaulis subcommunity the grassland vegetation is characterised by the presence of the perennial bunch grasses themeda triandra, heteropogon contortus and eragrostis curvula and the forbs commelina africana and oxalis obliquifolia (species group am), with constancies higher than 50% throughout the study area. other prominent species include the shortperennial grasses tristachya leucothrix (species group ad), eragrostis racemosa and harpochloa falx, and the geophyte watsonia lepida (species group ak). 1. themeda triandra – cyanotis speciosa major community this major community is found on the hot, dry northern and western aspects of platberg, at relatively low altitudes (1939 m.a.s.l. – 2078 m.a.s.l.). this community is characterised by the presence of species from species group i (online supplement) and is dominated by the grasses eragrostis curvula and themeda triandra (species group am). other prominent species include the grass tristachya leucothrix (species group ad) and the forb cyanotis speciosa (species group i). 1.1 cheilantes ecklonia – crassula sarcocaulis community this community is found on a north-western aspect on very steep slopes ( > 26°) in full sun at an altitude of approximately 2000 m.a.s.l. rock cover varies from 5% to 50% with moist, shallow soils (20 mm – 180 mm deep). the geology is sheet rock outcrops on clarens formation sandstone and dolerite dykes. the communityshows either no or moderate levels of trampling or erosion. the community is characterised by species from speciesgroup a and includes the succulents crassula sarcocaulis and crassula lanceolata ssp. transvaalensis, the sedge pycreus nigricans, the forb pelargonium bowkerii and the geophytes hypoxis filiformis and ornithogalum tenuifolium. the community is poorly represented by species from other groups with a paucity of grasses. it has one dominant species – the xerophytic fern cheilanthes eckloniana (species group h) – and significant cover abundance from the grass dwarf succulent shrub c. sarcocaulis and the sedge p. nigricans (species group a), the grass melinis nerviglumis (species group h), the forb bidens pilosa (species group i) and the geophyte hypoxis costata (species group y). the endemic geophyte merwilla plumbea (species group ad) is also present. there is an average of 20.6 species per 30 m². 1.2 hyparrhenia hirta – tristachya leucothrix community this community is located at relatively low altitudes (1926 m – 2078 m) on the warm northern or north-western aspect in full sun. the slope varies from moderate to very steep (8° – 45°). rock cover varies from 0% to 40%, with an average of between 5% and 10%. it is characterised by species from species group e and consists of only two grasses, namely hyparrhenia hirta and brachiaria serrata. the vegetation is dominated bythe perennial grasses tristachya leucothrix (species group ad) and themeda triandra (species group am), with significant cover abundance of heteropogon contortus, eragrostis curvula (species group am), eragrostis racemosa and harpochloa falx (species group al), the forb felicia filifolia (species group aj), the dwarf shrub searsia discolor (species group ad), and the fynbos sub-shrub selago galpinii (species group ad). the community has 32.41 species per 30 m². the hyparrhenia hirta – tristachya leucothrix community has four sub-communities: 1.2.1 hyparrhenia hirta – leonotis ocymifolia subcommunity this subcommunity is located on the warm, dry northern side of platberg on very steep (26° – 45°) slopes. soils are dry and shallow (30 mm – 120 mm), with little rock cover (< 10%). moderate to high levels of trampling and erosion observed are due to vegetation use by cattle. this subcommunity is characterised by the forbs leonotis ocymifolia and the geophyte asparagus cooperi from species group b. it is dominated by the perennial grasseshyparrhenia hirta (species group e), themeda triandra and eragrostis curvula (species group am), and harpochloa falx (species group al). 1.2.2 selago galpinii – hyparrhenia hirta subcommunity this subcommunity is located on the northern moderate to steep (8° – 26°) stony hill slopes on the sandstone of the clarens formation. soils are shallow (40 mm –100 mm) and dry and display moderate to high levels of trampling and erosion. low sclerophyllous shrubs (< 2 m) provide limited canopy cover (< 25%). there is no diagnostic species group for this subcommunity and it is defined by the bunch grass hyparrhenia hirta (species group e) and the sub-shrub selago galpinii (species group ad). it is dominated locally by the perennial grasses tristachya leucothrix (species group ad), themeda triandra and heteropogon contortus (species group am). other species that provide significant cover abundance include felicia filifolia (species group aj) and the forb acalypha punctata (species group ad). 1.2.3 felicia filifolia – diospyros austro-africana subcommunity the felicia filifolia – diospyros austro-africana subcommunity is located at an altitude of 2000 m on the hot, dry northern aspect. the slope is between 16° and 45°. the subcommunity is located on cave sandstone sheet rock, with dry shallow soil (40 mm – 100 mm deep). no trampling or erosion was recorded. canopy cover ofbetween 5% and 15% was observed, provided by the low (< 2 m) shrub diospyros austro-africana. species group c defines this subcommunity and includes the shrubs d. austro-africana, euclea crispa, osyris lanceolata and searsia dentata, the grass trachypogonspicatus, the forb helichrysum aureonitens and the fynbos shrub cliffortia ramosissima. the subcommunity is dominated by the perennial grasses themda triandra, eragrostis curvula (species group am) and eragrostis racemosa (species group al), the low sedge cyperus semitrifidus (species group aj), and the dwarf shrub d. austro-africana (species group c). other species locally dominant include the fynbos shrub c. ramosissima, the tree e. crispa (species group c) and the short grass brachiaria serrata (species group e). 1.2.4 hyparrhenia hirta – polygala hottentotta subcommunity the subcommunity is located between 1992 m.a.s.l. and 2078 m.a.s.l. on the hot, dry northern or north-eastern aspect. the slope varies from moderate to very steep (16° – 45°). rock size varies from stones to boulders (50mm to more than 500 mm). soils are dry, mixed ib type and of medium depth (120 mm – 200 mm) on cave sandstone. levels of erosion and trampling are moderate to high. only one relevé had canopy cover (5%) for shrubs lower than 2 m. the vegetation comprises mostly tall, perennial grasses, with the shrub diospyros austro africana present in some areas. two plants from species group d define this community. they are the erect forb polygala hottentotta and the uncommon succulent aloe maculata. there is a well-developed grass layer that dominates the community and includes the perennial grasses hyparrhenia hirta (species group e), tristachya leucothrix (species group ad),eragrostis racemosa (species group al), themeda triandra and heteropogon contortus (species group am). other species of significance are the geophyte hypoxis rigidula (species group f) and the grasses cymbopogon dieterlenii (species group aj) and eragrostis curvula (species group am). 1.3 eragrostis caesia – ipomoea crassipes community this community is found on the farm platberg on most aspects on the northern flank of platberg. the slope is steep to very steep (16° – 45°) on scree or boulder beds, below the cave sandstone at an altitude of approximately 2000 m.a.s.l. rock size exceeds 500 mm, with exposure between 10% and 30%. soils are dry and shallow(30 mm – 120 mm) and of mixed ib type. no or only moderate trampling and erosion were observed. this community is poorly developed and is characterised by the single grass species eragrostis caesia (species group d). the community has no dominant species,but the grasses themeda triandra, eragrostis curvula, heteropogon contortus (species group am) and tristachya leucothrix (species group ad) provide significant cover abundance. an average of 27.57 species per 30 m² was recorded. 1.4 cyanotis speciosa–leonotis ocymifolia community this community is located on basalt scree and boulder rubble on and below cave sandstone outcrops at altitudes between 2000 m.a.s.l. and 2100 m.a.s.l. it is located on the farm platberg on the hot, dry northern or north-western grassy slopes (16° – 45°). soils are shallow (40 mm – 120 mm) with good drainage and not more than 15% rock cover. no or only moderate erosion and trampling due to livestock usage were observed. this community has no single defining species group; it is characterised by the forbs leonotis ocymifolia (species group b) and cyanotis speciosa (species group i). vegetativley it is poorly developed and dominated by a single grass, themeda triandra, with significant presence of the perennial grasses eragrostis curvula (species group am), harpochloa falx (species group al) and tristachya leucothrix, the dwarf shrub searsia discolor and the forb acalypha punctata (species group ad). anaverage of 24.88 species per 30 m² was recorded. 2. agapanthus campanulatus – aristida monticola major community this grassland community is found on all aspects on the summit area of platberg. it consists of relatively flat terrain on the plateau region or rocky outcrops on the summit or steep sides. this major community is characterised by species group ai (online supplement) and is dominated by the forb watsonia lepida (species group ak) and short, perennial grasses tristachya leucothrix (species group ad), heteropogon contortus, themeda triandra (species group am), eragrostis racemosa and harpochloa falx (species group al). within this major grassland community, two distinct vegetation divisions occur: a moist zone with deep soils, dominated by the tall maroon grass andropogon appendiculatus (species group s), and a series of progressively rocky regions with shallow, dry soils dominated by three short grasses, namely t. leucothrix (speciesgroup ad), elionurus muticus and digitaria monodactyla (species group aj). 2.1 andropogon appendiculatus – helictotrichon longifolium community this community is located on all aspects of the summit area on flat or gently rolling terrain with damp soils (> 300 mm deep). no trampling or erosion is evident. there are no rock sheets, outcrops or boulders; the soil is fine grained (< 5 mm). species group s defines this community and includes the perennial bunch grasses andropogon appendiculatus and helictotrichon longifolium, the forbs senecio inornatus and zaluzianskya microsiphon. vegetation is dominated by a well-developed grass layer, which includes a. appendiculatus, h. longifolium (species group s), tristachya leucothrix (species group ad), themeda triandra and heteropogon contortus (species group am), and the bulb watsonia lepida (species group ak). other important species include the grasses eragrostis racemosa, harpochloa falx (species group al) and the forb helichrysum pilosellum (species group ab). this community has anaverage of 31.67 species per relevé. this community has eight sub-communities and six variants. 2.1.1 cineraria eriodioides–senecio inornatus subcommunity the cineraria eriodioides – senecio inornatus subcommunity has two variants. it is located on steep to very steep grassy slopes on the cool southern or south-westernside of platberg. soils are gravelly mixed ib type, with a depth of more than 300 mm, and are damp or waterlogged. rock exposure is either absent or between 5% and 20%, with no trampling or erosion present. there are two distinct altitudes where the two variants occur; the senecio inornatus variant at 2144 m.a.s.l. – 2297 m.a.s.l. and the ficinia stolonifera variant lower down at 1950 m.a.s.l. – 2196 m.a.s.l. the species clutia affinis, asparagus microraphis, cineraria erodioides, bupleurum mundii and the shrub euclea coriacea (species group l) define the community. grasses, forbs, succulents and herbs are all poorly developed. the subcommunity is dominated by only two species: the forb s. inornatus (species group s) and the grass themeda triandra (species group am). other species of importance include the perennial grasses andropogon appendiculatus, helictotrichon longifolium (species group s), cymbopogon dieterlenii and eragrostis chloromelas (species group aj), the forb c. affinis (species group l) and the geophyte watsonia lepida (species group ak). 2.1.2 tristachya leucothrix – koeleria capensis subcommunity this subcommunity variant is located on the hot and dry western, south-western and north-eastern moderately sloping (3° – 26°) footslopes of platberg at altitudesof between 1950 m.a.s.l. and 2100 m.a.s.l. soils are shallow to deep (40 mm – 300 mm) and well drained, with rock exposure ranging between 0% and 20%. the subcommunity is found on basalt scree rubble below the cave sandstone. no erosion or trampling is evident; however, the grasses are moribund. there is an interesting community association of defining species, which includes the short, high-altitude grass koeleria capensis, the sedge cyperus obtusiflorus var. flavissimus and the thymelaeaceae shrub struthiola angustiloba. this shrub species usually occurs in an area between the eastern cape and southern kwazulu-natal and hence it is the first record of the species in the free state. the community also includes two low fabaceae shrubs, namely argyrolobium harveyanum and eriosema kraussianum (species group m). the dominant species of the community form an extremely well-developed herbaceous layer and include the high-altitude, sour short grasses k. capensis (speciesgroup m), tristachya leucothrix (species group ad), elionurus muticus (species group aj), eragrostis racemosa, harpochloa falx (species group al) and heteropogon contortus (species group am). another species of significance is the perennial grass andropogon appendiculatus (species group s). herbs, forbs and geophytes are poorly developed, with the exception of helichrysum pilosellum (species group ab). 2.1.3 andropogon appendiculatus – pennisetum sphacelatum subcommunity the andropogon appendiculatus – pennisetum sphacelatum subcommunity is located on all aspects of platberg, except for the hot, dry northern and north-western sides, on the moderately flat (3° – 16°) summit plateau area and on the steep to very steep (16° – 45°) midslope. the geology is either summit basalt or mixed cavesandstone and basalt scree slopes. soils are all moist or waterlogged and generally deep (> 300 mm), although shallow (40 mm – 120 mm) soil is present on the midslope. rockiness is either absent or between 5% and 25%. this well-developed, tall (600 mm – 1200 mm) vegetation with its moist grassland community is moribund, indicating the lack of fire or grazing. it has two variants. the andropogon appendiculatus – pennisetum sphacelatum subcommunity is characterised by species group n and is poorly represented by species other than the perennial grass pennisetum sphacelatum. other species include the tall forbs senecio isatideus and cynoglossum hispidum, the geophyte moraea alticola and the prostrateherbs trifolium burchellianum, silene bellidioides and silene undulata. a well-developed grass layer dominates this community and includes p. sphacelatum (species group n), andropogon appendiculatus (species group s), tristachyaleucothrix (species group ad), aristida monticola (species group ai), themeda triandra and heteropogon contortus (species group am), and the bulb watsonia lepida (species group ak). other species common throughout this subcommunity include the grasses helictotrichon longifolium (species group s) and eragrostis curvula (species group am), and the forbs senecio inornatus (species group s) and helichrysum pilosellum (species group ab). there are two variants, namely nemesia caerulea and t. leucothrix. 2.1.4 watsonia lepida – helichrysum pallidum subcommunity this subcommunity is located on all cool aspects of platberg, at midslope altitudes of 1920 m.a.s.l. – 2100 m.a.s.l. and just below the summit plateau at 2248 m – 2295 m. the geology varies: the summit is pure basalt, whilst at lower altitudes there is a mix of clarens formation sandstone, dolerite dykes and basalt scree. soils are deep (> 300 mm) and seasonally damp, with only four communities having rock exposure of between 5% and 15%. no erosion or trampling was evident. thewatsonia lepida – helichrysum pallidum subcommunity is a short-to-medium height (300 mm – 500 mm) community with moribund grasses. there is no single defining species group, with the community being characterised by the bulb watsonia lepida (species group ak) and the forb helichrysum pallidum (species group ab). dominating the subcommunity are the grasses tristachya leucothrix (species group ad) and themeda triandra (species group am), and w. lepida (species group ak). species of importance include the grasses andropogon appendiculatus, aristida monticola (species group ai) and heteropogon contortus, and the forbs senecio inornatus (species group s) and commelina africana (species group am). 2.1.5 ledebouria cooperi – selago galpinii subcommunity the ledebouria cooperi – selago galpinii subcommunity is found on moderate rolling slopes (3° – 16°) of the summit plateau. this subcommunity is a high-altitude one,found at 2282 m.a.s.l. – 2336 m.a.s.l., with seasonally moist soils that are generally more than 300 mm deep. however, some sample plots have shallow soils (40 mm – 120 mm) with rock outcroppings of either 0% or between 5% and 15%. geology is summit basalt and shows no evidence of trampling or erosion. the geophyte ledebouria cooperi (species group aj) and the small shrub selago galpinii (species group ad) define this subcommunity. the subcommunity has a well-developed grass layer and is dominated by a single species, the perennial grass tristachya leucothrix (species group ad). other common species include the perennial grasses andropogon appendiculatus (species group s), aristida monticola (species group ai), eragrostis racemosa, harpochloa falx, themeda triandra and heteropogon contortus (species group am), the sedge carex zuluensis (species group ag) and the geophyte watsonia lepida (species group ak). 2.1.6 andropogon appendiculatus – senecio pauciculatus subcommunity the andropogon appendiculatus – senecio pauciculatus subcommunity is found on cool southern aspects of the plateau at 2307 m.a.s.l. – 2337 m.a.s.l. the terrain has a moderate slope (3° – 16°) with seasonally damp soils of medium depth (80 mm – 180 mm) on basalt or cave sandstone. the subcommunity is located on rocky terrain with 5% – 25% outcrop cover. there is no trampling or erosion. this subcommunity has no single defining species. it is characterised by andropogon appendiculatus (species group s) and senecio pauciculatus (species group x). the community has a well-developed grass layer, which includes the perennial grasses a. appendiculatus, helictotrichon longifolium (species group s) and tristachyaleucothrix (species group ad), the sour mountain grass aristida monticola (species group ai), elionurus muticus (species group aj), the short sour grasses eragrostis racemosa (species group al), heteropogon contortus and themeda triandra (species group am), the grass-like dianthus mooiensis (species group ab) and the geophytewatsonia lepida (species group ak). 2.1.7 lotononis lotononoides – hesperantha baurii subcommunity this lotononis lotononoides – hesperantha baurii subcommunity is a low-grass (300 mm – 400 mm), sub-shrub community located on all aspects of the cool, southernside of platberg. altitudes vary from 2198 m.a.s.l. to 2324 m.a.s.l. geology is summit basalt or cave sandstone at slightly lower altitudes. the slope is either gently rolling (3° – 8°) or mild to steep (16° – 45°). soils are seasonally damp and of medium depth (100 mm – 200 mm) with some rock exposure (5% – 15%). no or moderate erosion was observed but no trampling was evident. the lotononis lotononoides – hesperantha baurii subcommunity is characterised by members of species group q and includes the invasive alien tree pinus patula, the geophyte hesperantha baurii, the herbs haplocarpha scaposa and alepidea natalensis, the succulent crassula vaginata and the grass panicum coloratum. the subcommunity is dominated by only two species, namely the sour grass tristachya leucothrix (species group ad) and the geophyte watsonia lepida (species group am). other species with significant presence include the perennial grass heteropogon contortus (species group am) and the forb lotononis lotononoides (species groupaa). there are two variants, namely aster bakeranus and erica alopecurus. detailed descriptions of all variants, including these two, can be found in brand (2007). 2.1.8 habenaria lithophyla – eragrostis racemosa subcommunity this subcommunity is located on moderate to steep slopes (8° – 26°) on southern aspects of the stony summit basalt plateau area. altitudes are high and range from2254 m.a.s.l. to 2321 m.a.s.l. rock exposure varies between 10% and 40% on very shallow, seasonally wet soils of 20 mm to 120 mm deep. no erosion or trampling is evident. the small, erect orchid habenaria lithophyla (species group r) and the short grass eragrostis racemosa (species group al) define the subcommunity. it has poorly developed grammanoid geophytes and forbs. species with significant presence include the grasses tristachya leucotrix (species group ad), eragrostis racemosa (species group al) and heteropogon contortus (species group am), and the geophyte watsonia lepida (species group ak). 2.2 muraltia saxicola – helichrysum chionosphaerum community this community has three sub-communities and is located either on the summit basalt or the cave sandstone. this community occurs at either lower altitudes, atabout 2100 m, or high altitudes on stony, rocky summit areas between 2250 m.a.s.l. – 2378 m.a.s.l. outcrop exposure varies from 5% to 25% along moderate slopes (3° – 16°). soils are dry and shallow (60 mm – 180 mm) or seasonally damp(180 mm – 220 mm deep). vegetation varies between 200 mm and 400 mm high and shows little or no signs of erosion or trampling. the succulent delosperma sutherlandii, the two asteraceous forbs helichrysum chionosphaerum and helichrysum argentissimum, and xysmalobium involucratum from species group v define this community. this community has a well-developed grass layer, which includes the perennial grasses tristachya leucothrix (species group ad), eragrostis racemosa, harpochloa falx (species group al) and heteropogon contortus (species group am), and the dominant geophyte watsonia lepida (species group ak). other species of significance include the prostrate fynbos dwarf shrub muraltia saxicola (species group w), the sedge ficinia stolonifera (species group ai) and the forb h. chionosphaerum (species group v). this community has an average of 34.41 species per relevé. 2.2.1 muraltia saxicola – aristea woodii subcommunity this is a short-grass, prostrate forb community, located on stony, gradually sloping (3° 16°) basalt terrain on the warm, northern aspects of platberg. soils are dry and shallow (60 mm – 180 mm) on rocky outcrops showing 5% – 20% exposure. no trampling or erosion was recorded. species group t defines this community and includes the geophyte aristea woodii, the forb senecio inequidens and the orchid satyrium longicaula. the subcommunity is dominated by the short perennial grasses tristachya leucothrix (species group ad), eragrostis racemosa, harpochloa falx (species group al) andheteropogon contortus (species group am), and the bulb watsonia lepida (species group ak). 2.2.2 muraltia saxicola – helichrysum aureum subcommunity this prostrate sclerophyllerous forb subcommunity of medium height (400 mm) is located on all southern aspects on grassed, rocky basalt outcrops on the plateau area. it occurs at altitudes between 2250 m.a.s.l. and 2330 m.a.s.l., with 10% – 25% rock outcrop. soils are seasonally damp, shallow to medium deep (40 mm – 220mm) and show no evidence of erosion or trampling. the subcommunity is characterised by species from species group u and includes the forbs helichrysum aureum and wahlenbergia lycopodioides, the sedges eleocharis dregeana and scleria welwitschii, and the fynbos sub-shrub gnidia fastigata. the short grasses tristachya leucothrix (species group ad), eragrostis racemosa, harpochloa falx (species group al) and heteropogon contortus (species group am) and the geophyte watsonia lepida (species group ak) dominate this subcommunity. there is limited presence of other species; the only ones of significance are the fynbosprostrate dwarf shrub muraltia saxicola (species group w) and the forb helichrysum chionosphaerum (species group v). 2.2.3 ficinia stolonifera – muraltia saxicola subcommunity this sedge and prostrate sclerophyllous forb subcommunity is located on all aspects of platberg on the summit basalt and the cave sandstone. altitudes range from low (2100 m.a.s.l.) to high (2259 m.a.s.l. – 2378 m.a.s.l.). the slope is gradual (3° – 8°), with rock exposure of 10% – 20% on seasonally damp soil of variable depths (between 20 mm and 200 mm). vegetation varies between 200 mm and 400 mm high and shows no signs of erosion or trampling. there is no defining species group for this subcommunity, which is characterised by the common sedge ficinia stolonifera (species group ai) and the prostrate fynbossub-shrub muraltia saxicola (species group w). it is dominated by a well-developed grammanoid layer, which includes the perennial grasses tristachya leucotrix (species group ad), eragrostis racemosa (species group al) and heteropogon contortus (species group am), and the sedge ficinia stolonifera (species group ai). the only forb and geophyte of significance are helichrysum chionosphaerum (species group v) and watsonia lepida (species group ak), respectively. 2.3 andropogon distachyos – helichrysum aureum community the andropogon distachyos – helichrysum aureum community has a limited extent and is located on only three relevés. it comprises a total of 133 species. this community has the highest species average (43.33 species per relevé). it is found on rocky outcrops (5% – 15% exposure) on moderate to steep (16° – 26°) midslopes on the northern aspects between1970 m.a.s.l. and 1993 m.a.s.l. the community is located on a dolerite dyke, which intrudes the clarens formation cave sandstone. the soil is a mixed ib type. no or only moderate erosion and trampling are evident on well-drained soils that range from shallow to medium deep (40 mm – 200 mm). members of species group z define this community and include the the low fynbos sub-shrub gnidia caffra, the perennial grasses andropogon distachyos and monocymbium ceresiiforme, the forbs helichrysum aureum, acalypha schinzii, argyrolobium stipulaceum, senecio coronatus, xysmalobium parviflorus and dicoma anomala, and the geophytic forb pentanisia prunelloides. there is a paucity of species from most other groups, with a poorly developed grass layer. species that dominate this community include the perennial, short sour grasses a. distachyos (species group z), tristachya leucothrix (species group ad) and eragrostis racemosa (species group al), and the bulb watsonia lepida (species groupak). other important species include the grasses m. ceresiiforme (species group z) and themeda triandra (species group am), and the forb h. aureum (species group z). 2.4 passerina montana – melinis nerviglumis community the passerina montana – melinis nerviglumis community comprises a mixture of woody and grass species with 5% – 70% canopy cover provided by shrubs of between 2 m and 6 m high. it is located on the northern, north-western and southern aspects on mobile scree or the rocky (15% – 45%) contact between the basalt and cave sandstone on steep to very steep (16° – 45°) slopes. soils are shallow to medium deep (10 mm – 200 mm) and moderate to high erosion levels were recorded. the passerina montana – melinis nerviglumis community is characterised by species from species group ac, which comprises the low sclerophyllous shrub passerina montana, the geophyte ledebouria ovatifolia, the fern asplenium aethiopicum, the caespitose forb helichrysum sutherlandii, and the restionaceae ischyrolepis schoenoides. it is dominated by only two species, namely the fynbos shrub p. montana (species group ac), and the bulb watsonia. lepida (species group ak). it has a poorly developed grammanoid layer that includes panicum ecklonii, tristachya leucothrix (species group ad), i. schoenoides (species group ac), the sedge ficinia stolonifera (species group ai) and the grasses themeda triandra, heteropogon contortus and eragrostis curvula (species group am). this community has an average of 35.40 species per relevé. 2.5 cyperus semitrifidus – digitaria monodactyla community the cyperus semitrifidus – digitaria monodactyla community is located on all aspects and cardinal points on flat to moderate (3° – 16°) rocky terrain on the summitplateau basalt. soils are shallow (20 mm – 120 mm) and damp. rock cover varies between 5% and 50%, but averages 25%. the vegetation is low (10 cm – 30 cm), with poorly developed grass, forb, herb and succulent layers. the flat, wet, loose gravel and sheet rock nature of the community results in no or very little erosion and trampling. there is no defining species group for this community; however, the sedge cyperus semitrifidus and the short perennial grass digitaria monodactyla, both from species group aj, define this community. these two species also dominate the community, which has a poorly developed grass and forb layer. additional species ofimportance include the mountain grass aristida monticola and the two forbs agapanthus campanulatus and helichrysum dasycephalum (species group ai). the cyperus semitrifidus – digitaria monodactyla community has three sub-communities, namely the digitaria monodactyla – selaginella caffrorum, eriospermumornithogaloides – digitaria monodactyla and the digitaria monodactyla – crassula sarcocaulis sub-communities. the three sub-communities are defined by short grasses and prostrate ferns, minute geophytes, and karroid succulent shrubs, respectively. the community is characterised by 14 relevés, with an average of 25.14 species per relevé. 2.5.1 digitaria monodactyla – selaginella caffrorum subcommunity this is a low (5 cm – 25 cm) subcommunity located on rock sheets with 25% – 50% outcrop on basalt, dolerite and cave sandstone, on all aspects of platberg. soils are moist and very shallow (20 mm – 60 mm), without any erosion or trampling. most grasses and forbs are poorly developed, with the community characterised by the single, fine-leaved prostrate fern selaginella caffrorum (species group ae). only two species dominate the community, namely the short grass digitaria monodactyla (species group aj) and the fern s. caffrorum (species group ae). other species of significance include the succulent sub-shrub crassula sarcocaulis (species group a) and the geophyte schizocarphus nervosa (species group ag). 2.5.2 eriospermum ornithogaloides–digitaria monodactyla subcommunity this is subcommunity comprising low grasses (100 mm – 200 mm) and geophytes, located on the cool, southern side on flat basalt rock sheets with 10% – 30% exposure on the summit plateau area. it occurs between 2196 m.a.s.l. and 2328 m.a.s.l. soils are shallow (20 mm – 120 mm), damp to seasonally waterlogged and consist of degraded gravel (grain < 10 mm). no erosion or trampling is evident. it has no defining species group; rather, it is characterised by the geophyte eriospermum ornithogaloides (species group ag) and the monotypic grass digitaria monodactyla (species group aj). it has a very poorly developed grass and forb layer, without any single species clearly dominating the subcommunity. species ofimportance include the grass d. monodactyla and the forb helichrysum dasycephalum (species group aj) and the geophyte agapanthus campanulatus (species group ai). 2.5.3 digitaria monodactyla – crassula sarcocaulis subcommunity the digitaria monodactyla – crassula sarcocaulis subcommunity is found on rocky outcrops, gravel plains and rock sheets (exposure 5% – 15%) on the eastern aspect on all sides of the summit basalt at 2100 m.a.s.l. – 2350 m.a.s.l. soils are shallow (20 mm – 100 mm) and seasonally damp, without any erosion or trampling. this subcommunity is characterised by succulent shrubs and low grasses, specifically filipendula vulgaris and the small succulent aloe aristata (species group ah). the herbaceous layer and species numbers are poorly developed throughout the community. the community is dominated by the succulent shrubs crassula sarcocaulis (species group a) and ruschia putterillii and the short sour grass digitaria monodactyla, both from species group aj. other species of significance include two grasses, cymbopogon dieterlenii and eragrostis chloromelas, the sedge cyperus semitrifudus (species group aj) and the exotic species f. vulgaris (species group ah). the consistency of cyanothis speciosa (species group i), which favours habitats with shallow soils on rocky terrain, should be noted. species richness and structure platberg has an average species richness of 30.98 species per 30 m2. the mean number of species (species abundance) in the two major communities are significantly different (t(167) = 2.020, p < 0.05). the mean number of species is significantly higher in major community 2 than in major community 1 (31.7 vs. 28.3), with a 95% confidence interval for the difference between means (0.08 – 6.85). the descriptive statistics for species abundance are provided in table 1. for major community 1, the sample plots were located at relatively low altitudes (< 2000 m.a.s.l.), with an average species richness of 28.27 species per 30 m2. for major community 2, the relevés were spread over the plateau area of platberg at altitudes higher than 2000 m.a.s.l., with an average species richness of 31.73 species per 30 m2. thus, the data indicate that more species occur at higher altitudes. owing to a significant difference in species abundance between the major communities, the next step in the analysis process was to examine differences between the species abundance of the respective communities within each of the major communities. no significant difference was found for the species abundance of the communities comprising major community 1, with kruskal–wallis h(3, n = 37) = 7.14, p > 0.05. a significant difference in species abundance was found between the communities within major community 2, with kruskal–wallis h(4, n = 132) = 15.75, p < 0.01. moreover, multiple comparisons of the mean ranks for all groups within major community 2 revealed significant differences between the species abundance of communities 2.2 and 2.5, and also between communities 2.3 and 2.5 (p < 0.05 for both comparisons). figures 3 and 4 provide graphic representations of these comparisons in the form of box plots, with 95% confidence intervals for median values of the species abundance of the communities within major communities 1 and 2, respectively. (the descriptive statistics for species abundance for the communities within each of the major communities are provided in table 2.) in both these comparisons, the species abundance of community 2.5 was the lower of the respective pairs. community 2.5 occurs on shallow, moist soil that is underlain by basalt rock sheets. these conditions are not conducive to plant growth, which could explain the low species richness of this high-altitude community. both communities 2.2 and 2.3 are located on rocky terrain with deeper soil and a combination of grasses, forbs and low shrubs. this higher species richness of rocky terrains may be ascribed to rocks protecting seedlings and young plants from grazing and fire, as well as to reduced effects of shading from tall woody vegetation (everson, everson & tainton 1988; heini, silva & tacheba 2004; seabloom & richards 2003). table 1: species richness for the major plant communities on platberg. table 2: species abundance for the different communities within each of the major communities. discussion (back to top) with regard to species composition, the grassland communities along the slopes and footslopes (major community 1) and those on the plateau (major community 2) on platberg show distinct dissimilarities. this can be attributed to different geological substrates, altitude differences and percentage surface rock. the themeda triandra – cyanotis speciosa major community (1) occurs on shallow soils mostly on the clarens, elliot and molteno formations and is composed, generally, of short grasses and low forbs on rocky terrain or outcrops. theagapanthus campanulatus – aristida monticola major community (2) is a medium-to-tall grassland restricted to the plateau (basalts and dolerite) with deeper soil and higher clay content, which has greater moisture retention ability. floristic affinities some floristic associations exist between platberg and korannaberg, which is located approximately 300 km west of platberg in the drier interior of the eastern free state. korannaberg is the only other inselberg on which a similar study has been conducted. it is at a lower altitude (1500 m – 1881 m) (du preez 1991; du preez & bredenkamp 1991) and has no basaltic cap. common species shared by these two sites include the grasses cymbopogon dieterlenii, digitaria monodactyla, elionurus muticus, eragrostis capensis, eragrostis chloromelas, eragrostis curvula, eragrostis racemosa, harpochloa falx, pentaschistis setifolia, themeda triandra and tristachya leucothrix, and the forbs helichrysum nudifolium, selaginella caffrorum and crassula lanceolata. the festuca caprina grassland of the golden gate highlands national park and qwa-qwa national park (mucina & rutherford 2006) also has similarities with that of platberg. grasses prominent in both areas include c. dieterlenii, e. muticus, e. curvula, t. triandra and t. leucothrix. floristic links also exist with the cathedral peak area of the drakensberg, where the themeda–festuca sub-alpine sour grassland dominates the main montane region. these genera are also dominant on platberg. numerous species recorded at platberg and in the drakensberg at a high altitude have been identified for primary grassland communities in gauteng (behr & bredenkamp 1988) and for the bankenveld (bredenkamp & brown 2003; grobler, bredenkamp & brown 2006). they include ajuga ophrydis, schistostephium crataegifolium, senecio isatideus, senecio inornatus, sida dregei, the woody gnidia capitata and searsia discolor. grasses, which are either characteristic or dominant species in platberg and have also been recorded by grobler et al. (2006) and bredenkamp and brown (2003), include brachiaria serrata, d. monodactyla, e. muticus, e. racemosa, e. chloromelas, e. curvula, heteropogon contortus, hyparrhenia hirta, melinis nerviglumis, monocymbium ceresiiforme, panicum natalensis, setaria sphacelata, t. triandra, trachypogon spicatus and t. leucothrix. species recorded at platberg that emphasise the floristic relationships of the dac, the karoo and the fynbos (acocks 1988; hill 1996; hilliard & burtt 1987; kay, bredenkamp & theron 1993; moffett et al. 2001; roberts 1969) include: a. ophrydis, andropogon appendiculatis, aristida junciformis, aristida monticola, b. serrata, cheilanthes quadripinnata, cheilanthes hirta, commelina africana, crassula alba, d. monodactyla, e. racemosa, e. chloromelas, e. curvula, e. muticus, euclea crispa, euphorbiastriata, g. capitata, h. falx, helichrysum aureonitens, helichrysum polycladum, helichrysum rugulosum, helichrysum cephaloideum, hermannia depressa, h. contortus, h. hirta, ipomoea crassipes, leucosidea sericea, m. nerviglumis, m. ceressiforme, myrsine africana, pollichia campestris, searsia rigida, s. discolor, s. isatideus, pennisetum sphacelatum, t. triandra, t. spicatus and vernonia natalensis. according to mucina and rutherford (2006) three major temperate grassland genera, namely festuca, bromus and koeleria, represent a northern affinity, showing links with east africa and eurasia, whilst genera such as pentaschistis and merxmuellera have southern affinities with the cape region. these genera are also presenton platberg, indicating floristic affinity with these areas. conservation very little of the dac grassland is preserved, with the majority in kwazulu-natal and a smaller portion in the golden gate highlands national park and the qwa-qwa national park (now incorporated into the golden gate highlands national park). only 5.5% (2194 km²) of the entire dac is protected (carbutt & edwards 2004, 2006). these habitats, with their complex abiotic and biotic factors, are responsible for generating high species richness (cowling, holmes & rebelo 1992; ed.huntley 1989; van wyk & smith 2001). the platberg grasslands as found in this study are mostly natural with limited human impact and a relatively high species richness. various affinities with other high-altitude grasslands exist and as such the conservation of this area will contribute to the conservation of the larger grassland biome in southern africa. conclusion (back to top) prior to this study, very little was known about the vegetation of platberg. this phytosociological classification and description of the vegetation of platberg serves as an inventory of the natural ecosystems and plant species present in the area. this study resulted in the identification 26 plant communities, which can be grouped into two major communities. a total of 551 vascular plants, of which 26 are endemic or near-endemic dac species, were identified. floristic affinities with various other high-lying grasslands were found. the species richness of the study area, together with the threat of declining genetic diversity (ed. anderson 2001), global warming and subsequent climate change (peters 1992), underlines the importance of mountains and the associated inselbergs as refuge areas for different plant species and the need for their urgent protection. as a unit the grassland biome requires more protected areas, identified according to ecological criteria based on phytosociological studies that are able to identify vegetation units on which rare, threatened, endemic or near-endemic plants grow. it is important that platberg specifically, but inselbergs generally, be given attention as conservation sites with the development of a management plan by the department of environment, tourism and economic affairs of the free state. the results of this study could serve as a basis for future conservation planning and management of the area. the presence of various alien plant species poses a threat to the natural vegetation. it is recommended that an eradication plan is implemented as soon as possible so as to prevent the degradation of this natural grassland ecosystem. acknowledgements (back to top) the authors acknowledge the national geographic society (grant number 7920-05) for the generous funds without which the fieldwork and the study would not have been possible. authors’ contributions r.f.b., l.r.b. and p.j.d.p. all contributed equally to data collection and analysis and subsequent writing of the manuscript. references (back to top) acocks, j.p.h., 1988, veld types of south africa, botanical research institute, pretoria. (memoirs of the botanical survey of south africa, no. 57) anderson, j.m. (ed.), 2001, towards gondwana alive, national botanical institute, pretoria. behr, c.m. & bredenkamp, g.j., 1988, ‘an analysis of the flowering plants and ferns of the witwatersrand national botanic garden’, south african journal of botany 54(6), 515–524. brand, r.f., 2007, ‘phytosociology of platberg mountain, eastern free state, south africa’, phd thesis, dept. of plant sciences, university of the free state. brand, r.f., du preez, p.j. & brown, l.r., 2008, ‘a floristic description of the afromontane fynbos communities on platberg, eastern free state, south africa’, koedoe 50(1), 202–213. brand, r.f., du preez, p.j. & brown, l.r., 2010, ‘a floristic analysis of the vegetation of platberg, eastern free state, south africa’, koedoe 52(1), 1–11. bredenkamp, g.j. & brown, l.r., 2003, ‘a reappraisal of acocks’ bankenveld: origin and diversity of vegetation types’, south african journal of botany 69(1), 7–26. burke, a., 2001, ‘determinants of inselberg floras in arid nama karoo landscape’, journal of biogeography 22, 1211–1220. burke, a., 2004, ‘from plains to inselbergs: species in special habitats as indicators of climate change?’, journal of biogeography 31, 831–841. http://dx.doi.org/10.1046/j.13652699.2003.00984.x carbutt, c. & edwards, t.j., 2004, ‘the flora of the drakensberg alpine centre’, edinburgh journal of botany 60(3), 581–607. carbutt, c. & edwards, t.j., 2006, ‘the endemic and near-endemic angiosperms of the drakensberg alpine centre’, south african journal of botany 72, 105–132. cowling, r.m., holmes, p.m. & rebelo, a.g., 1992, ‘plant diversity and endemism’, in r.m. cowling (ed.), the ecology of fynbos: nutrients, fire and diversity, pp. 4–21, oxford university press, cape town. du preez, p.j., 1991, ‘a syntaxonomical and synecological study of the vegetation of the south-eastern orange free state and related areas with special reference to korannaberg’, phd thesis, dept. of plant sciences, university of the orange free state. du preez, p.j., 1992, ‘the classification of the vegetation of korannaberg, eastern orange free state, i, afromontane fynbos communities’, south african journal of botany 58(3),165–172. du preez, p.j. & bredenkamp, g.j., 1991, ‘vegetation classes of the southern and eastern orange free state (republic of south africa) and the highlands of lesotho’, navorsing nasionale museum, bloemfontein 7, 477–526. du preez, p.j., bredenkamp, g.j. & venter, h.j.t., 1991, ‘the syntaxonomy and synecology of the forests in the eastern orange free state, south africa’, south african journal of botany 57, 198–206. everson, c.s., everson, m.t. & tainton, n.m., 1988, ‘effects of intensity and height of shading on the tiller initiation of six grass species from the highland sourveld of natal’, south african journal of botany 54(4), 315–318. germishuizen, g. & meyer, n.l. (eds.), 2003, plants of southern africa: an annotated checklist, national botanical institute, pretoria (strelitzia, no. 14). grobler, c.h., bredenkamp, g.j. & brown, l.r., 2006, ‘primary grassland communities of the urban open spaces in gauteng, south africa’, south african journal of botany 72, 367– 377. http://dx.doi.org/10.1016/j.sajb.2005.10.008 heini, m., silva, j. & tacheba, b., 2004, ‘vegetation changes after a single fire-event in the okavango delta wetland, botswana’, south african journal of botany 70(5), 695–704. hennekens, s.m., 1996a, turboveg: a software package for input, processing, and presentation of phytosociological data,computer software, university of lancaster, ibn-dlo. hennekens, s.m., 1996b, megatab: a visual editor for phytosociological tables, computer software, giesen & geurts, ulft. hennekens, s.m., 1998, turboveg: clipper database management software for storage, selection, and export of vegetation data (relevés), computer software, green world research, wageningen. hill, t.r., 1996, ‘description, classification and ordination of the dominant vegetation communities, cathedral peak, kwazulu-natal drakensberg’, south african journal of botany 62, 263–269. hilliard, o.m. & burtt, b.l., 1987, the botany of southern natal drakensberg, national botanical gardens, cape town. hoare, d.b. & bredenkamp, g.j., 2001, ‘syntaxonomy and environmental gradients of the grasslands of the stormberg/drakensberg mountain region of the eastern cape, south africa’, south african journal of botany 67, 595–608. huntley, b.j. (ed.), 1989, biotic diversity in southern africa, concepts and conservation, oxford university press, cape town. huntley, b.j. (ed.), 1994, botanical diversity in southern africa, national botanical institute press, pretoria (strelitzia, no. 1). johnson, m.r., van vuuren, c.j., visser, j.n.j., cole, d.i., wickens, h. de v., christie, a.d.m. et al., 2006, ‘sedimentary rocks of the karoo supergroup’, in m.r. johnson, c.r. anhaeusser & r.j. thomas (eds.), the geology of south africa, pp. 461–499, geological society of south africa, johannesburg. kay, c., bredenkamp, g.j. & theron, g.k., 1993, ‘the plant communities of the golden gate highlands national park in the north-eastern orange free state’, south african journal of botany 59(4), 442–449. keddy, p.a., 2007, plants and vegetation: origins, processes and consequences, cambridge university press, cambridge. kent, m. & coker, p., 1992, vegetation description and analysis: a practical approach, john wiley & sons, chichester. killick, d.j.b., 1963, an account of the plant ecology of the cathedral peak area of the natal drakensberg, government printer, pretoria (memoirs of the botanical survey of south africa, no. 34). killick, d.j.b., 1978, ‘the afro-alpine region’, in m.j.a. werger (ed.), biogeography and ecology of southern africa, pp. 515–560, junk, the hague. malan, p.w., 1998, ‘vegetation ecology of the southern free state’, phd thesis, dept. of plant sciences, university of the orange free state. moffett, r.o., daemane, m.e., pitso, t.r., lentsoane, r. & taoana, t.r.n., 2001, ‘a checklist of the vascular plants of qwa-qwa and notes on the flora and vegetation of the area’, uniqwa research chronicles 3(2), 32–83. mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, south african national biodiversity institute, pretoria. (strelitzia, no 19) mueller-dombois, d. & ellenberg, h., 1974, aims and methods of vegetation ecology, john wiley & sons, new york. parmentier, i., oumorou, m., porembski, s., lejoly, j. & decocq, g., 2006, ‘ecology, distribution, and classification of xeric monocotyledonous mats on inselbergs in west africa and atlantic central africa’, phytocoenology 36(4), 547–564. peters, r.l., 1992, ‘conservation of biological diversity in the face of climatic change’, in r.l. peters & t.e. lovejoy (eds.), global warming and biological diversity, pp. 15–30, yale university press, new haven. porembski, s. & barthlott, g.b., 1995, ‘a species poor tropical sedge community: afrotrilepis pilosa mats on inselbergs in west africa’, nordic journal of botany 16(3), 239–246. porembski, s. & brown, g., 1995, ‘the vegetation of inselbergs in the comoé national park (ivory coast)’, conservatoire et jardin botanicues de geneve 50(2), 351–365. porembski, s., martinelli, g., ghlemüller, r. & barthlott, w., 1998, ‘diversity and ecology of saxicolous vegetation mats on inselbergs in the brazilian atlantic rainforest’, diversity and distribution 4, 107–119. porembski, s., szarzynske, j., mund, j-p. & barthlott, w., 1996, ‘biodiversity and vegetation of small-size inselbergs in a west african rain forest (taï, ivory coast)’, journal of biogeography 23, 47–55. roberts, b.r., 1969, ‘the vegetation of the golden gate highlands national park’, koedoe 12, 15–28. seabloom, e.w. & richards, s.a., 2003, ‘multiple stable equilibrium in grassland mediated by herbivore population dynamics and foraging behaviour’, ecology 84(11), 2891–2904. http://dx.doi.org/10.1890/02-0020 south african weather services, 2007, climate of south africa, climate statistics for bethlehem from 1961–1990, government printers, pretoria. soil classification working group, 1991, soil classification: a taxonomic system for south africa, department of agricultural development, soil and irrigation research institute, pretoria. (memoirs on the agricultural natural resources of south africa, no. 15) statistica version 9.0, 2009, computer software, statsoft inc., new york. van wyk, a.e. & smith, g.f., 2001, regions of floristic endemism in southern africa: a review with emphasis on succulents, umdaus press, pretoria. westhoff, v. & van der maarel, e., 1980, ‘the braun-blanquet approach’, in r.h. whittaker (ed.), classification of plant communities, pp. 287–378, kluwer academic publishers, the hague. abstract introduction materials and methods results discussion conclusion acknowledgements references appendix 1 about the author(s) keoikantse sianga okavango research institute, university of botswana, botswana richard fynn okavango research institute, university of botswana, botswana citation sianga, k. & fynn, r., 2017, ‘the vegetation and wildlife habitats of the savuti-mababe-linyanti ecosystem, northern botswana’, koedoe 59(2), a1406. https://doi.org/10.4102/koedoe.v59i2.1406 original research the vegetation and wildlife habitats of the savuti-mababe-linyanti ecosystem, northern botswana keoikantse sianga, richard fynn received: 12 may 2016; accepted: 06 mar. 2017; published: 23 may 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract this study classified the vegetation of the savuti-mababe-linyanti ecosystem (smle), northern botswana and developed a detailed map that provides a reliable habitat template of the smle for future wildlife habitat use studies. the major vegetation units of the smle were determined from satellite imagery and field visits and then mapped using landsat 8 and rapideye imagery and maximum likelihood classifier. these units were sampled using 40 m x 20 m (800 m²) plots in which the coverage of all plant species was estimated. non-metric multidimensional scaling (nms) demonstrated that plant communities were determined by gradients in soil texture or fertility and wetness. nms 1 represented a gradient of soil texture with seven woodland communities on sandy soils (sandveld communities and baikiaea forest) dominated by baikiaea plurijuga in baikiaea forest and terminalia sericea and philenoptera nelsii in sandveld, with various indicator species differentiating the various sandveld community types. mopane woodland further from and riparian woodland adjacent to permanent water was common on less sandy alluvial soils. mineral-rich, heavy clay soils in the sump of a large paleolake system support open grassland and mixed senegalia/vachellia (acacia) savanna, with the mineral-rich soils supporting grasses high in minerals such as phosphorus, calcium, sodium and potassium, and thus this region is a critical wet season range for migratory zebra. taller, high-quality grasses in the mosaic of sandveld and mopane woodland communities provide critical grazing for taller grass grazers such as buffalo, roan and sable antelope, whereas wetland communities provide reliable green forage during the dry season for a variety of herbivores, including elephant. this study has demonstrated how large-scale environmental gradients determine functional habitat heterogeneity for wildlife. conservation implications: our study demonstrated that the functionality of protected areas is determined by large-scale environmental gradients. thus conservation science must aim to ensure that protected areas cover the full range of key environmental gradients in a region (soil texture and wetness in our study). our habitat map provides a data base for wildlife habitat use studies in the region. introduction there is growing recognition of the importance of spatial heterogeneity in determining biodiversity (macfayden et al. 2016) and adaptive foraging options for herbivores (hobbs et al. 2008; hopcraft, olff & sinclair 2010; owen-smith 2004) and predators (hopcraft, sinclair & packer 2005) and for facilitating predation avoidance and avoidance of competition among guilds of herbivores or carnivores (mills & gorman 1997; rettie & messier 2000). spatial heterogeneity is strongly associated with vegetation heterogeneity, and detailed vegetation maps derived from remote-sensed spectral variation are likely to represent a large proportion of spatial heterogeneity (macfayden et al. 2016). plant community variation on environmental gradients is associated with complex combinations of environmental factors and associated plant species composition, richness and physiognomy, leading to distinct habitat attributes for animals such as seasonal resources, predation avoidance and shelter (fynn, chase & roder 2014; hopcraft et al. 2010). functional differences among different plant communities in meeting the seasonally varying needs of animals have been referred to as functional habitat heterogeneity and play a key role in determining the viability of herbivore populations (hopcraft et al. 2010; owen-smith 2004). environmental factors such as soil fertility, forage quality and water availability, which covary strongly with vegetation, may also influence human settlement, crop field and livestock distribution patterns. development of detailed vegetation maps is, therefore, essential for (1) understanding the distribution of biodiversity across a region, (2) research on wildlife species’ home range and habitat use, (3) conservation planning and (4) understanding socio-ecological interactions. vegetation maps are especially useful for determining seasonal habitat use of collared wildlife species where thousands of global positioning systems (gps) locations may be obtained from a collared animal, which cannot easily be followed up on, especially in vast wilderness regions with little road access, such as in northern botswana. currently, no detailed vegetation map exists for northern botswana, with the best available product being the elephant habitat map of northern botswana, which lacks the detail and accuracy needed for comprehensive wildlife habitat use studies (ringrose 2006). a key region for wildlife in northern botswana is the regional-scale contrast of extensive floodplains of the okavango delta and the linyanti swamps, with the vast woodland systems adjacent to these wetland systems. the savuti-mababe-linyanti ecosystem (smle) with its extensive wetlands and woodlands and the open grasslands of the mababe depression (md) has excellent functional habitat heterogeneity for wildlife (fynn et al. 2014), as well as great aesthetic features for tourism, providing the basis for it forming the core region for wildlife and tourism in botswana. thus, the smle is a region of key conservation importance, being one of the few remaining relatively unfragmented ecosystems in africa with important heterogeneity of seasonal habitats that support a wide diversity of wildlife species, including important populations of rare species such as wild dog, eland, sable and roan antelope, as well as long-range migrations of several herbivore species (fynn et al. 2014; naidoo et al. 2014; sianga 2014). thus, apart from the need for a detailed vegetation classification and habitat map for conservation planning in the smle and for future wildlife habitat use studies in the region, this article provides baseline ecosystem-level data on the smle, which allows the various herbivore and carnivore studies to be analysed in relation to the same habitat map. the objectives of this study were to (1) classify the vegetation of the smle of northern botswana, (2) develop a detailed vegetation map that provides a reliable habitat template of the smle for wildlife habitat use studies and for future studies and (3) discuss how the heterogeneity in vegetation and soils of the smle contributes to functional habitat heterogeneity for wildlife. materials and methods study area this study was done in the smle, northern botswana (figure 1), an extensive region of woodlands and open grasslands wedged between two major wetland systems – the okavango delta and the linyanti swamps. the smle is characterised by a semi-arid climate with an annual rainfall of around 500 mm on the western side (okavango delta) and increasing eastwards to around 600 mm in the chobe enclave region (botswana meteorological services). the climate is characterised by a wet season (december–april), cool, early dry season (may–august) and a hot, late dry season (september–november), where the daily maximum temperatures are regularly between 35 °c and 40 °c during this time (september–november) (fynn et al. 2014). water originating from the angolan highlands into the ecosystem through the okavango delta, linyanti swamps, selinda spillway and savuti channel leads to the development of extensive floodplain grasslands and swamps adjacent to the extensive woodland systems (mendelsohn et al. 2010). alluvial clays and aeolian sands in the ecosystem are occupied by colophospermum mopane and philenoptera nelsii–terminalia sericea woodlands (dryland woodlands), respectively (wolski & murray-hudson 2006), and low-lying drainage systems are dominated by woodlands of mixed communities (combretum imberbe and others) (mendelsohn et al. 2010). a large paleolake system known as the md (figure 1), characterised by clays of lacustrine origin, occurs between the okavango delta and the chobe region (teter 2007). the extremely high clay soils of the md provide key habitat heterogeneity in the general landscape of the smle, which is dominated by the kalahari sands. vegetation on the clay soils of the md is characterised by open senegalia/vachellia (acacia) spp. savanna grasslands and higher forage quality (fynn et al. 2014). figure 1: study area depicting the savuti-mababe-linyanti ecosystem and locations of sampling positions. vegetation sampling vegetation composition was sampled using 40 m × 20 m (800 m²) plots during the wet seasons of 2014 and 2015, with sampling conducted from january to the end of march to ensure that plants had attained inflorescences for easier identification. certain remote habitats of the md on water-logged heavy clay soils were inaccessible during the wet season and could only be sampled during the early dry season (april to mid-may) once the soils had dried out. plots were stratified within homogenous vegetation units (determined visually by extensive field surveys of the region) to ensure adequate sampling of all key vegetation types within the study area (figure 1). a total of 801 plots were located using two random numbers between 20 and 100, with the first random number taken along an access route such as a road and the second perpendicular from the road into the plant community. plots were generally spaced at least 500 m apart in each plant community. the gps coordinates of each plot were recorded using a garmin gps map 62s. all plants (trees, grasses and forbs) rooted in the plots were identified, recorded and their percentage cover estimated. unknown plants were pressed in the field and brought to the peter smith herbarium (psub) collection at the okavango research institute (maun, botswana) for identification, with nomenclature for all species following the plant list (http://www.theplantlist.org). five soil samples collected from each plot were mixed to form a composite sample, which was brought to the okavango research institute laboratory for texture and nutrient analysis. vegetation classification the data for all 801 vegetation plots (% cover abundance of species) were standardised using relativisations by maximum in principal component ordination 6 (pcord) (mccune, grace & urban 2002). these data were subjected to hierarchical cluster analysis (β linkage, β = -0.25, sorensen distance) in pcord 6 (mccune et al. 2002), comprising species (505 species) occurring at more than one site based on species distribution across 801 plots. indicator species analysis (isa) (dufrene & legendre 1997) was used to identify levels to define ecologically meaningful communities, and indicator values (ivs) were examined for statistical significance using monte carlo technique in pcord 6 (mccune et al. 2002). differences between communities were examined using multiresponse permutation procedure (mrpp) using sorensen distance measure (mccune et al. 2002). in addition, non-metric multidimensional scaling (nms) was used to plot the communities in ordination space in relation to key orthogonal gradients in the vegetation data using pcord 6 (mccune et al. 2002). such gradients are likely related to key environmental drivers and thus provide insights into how plant communities are structured in the ecosystem. vegetation mapping owing to the vast size of the smle, we were unable to purchase detailed rapideye imagery for the entire ecosystem and thus purchased rapideye imagery to map the core area of the ecosystem, where we needed most detail for several herbivore studies currently being conducted there and then used landsat 8 imagery to map the remainder of the ecosystem. a rapideye composite image composed of 34 tiles and wet season landsat enhanced thematic mapper scenes from earth explorer usgs of the study area were radiometrically and atmospherically corrected in envi 4.8 (envi 2010). the landsat images were mosaicked using seamless mosaic, and maximum likelihood classifier (supervised classification) in envi 4.8 (envi 2010) was used to map a subset of the 15 vegetation classes identified in the vegetation classification (reasons for mapping of a subset rather than all 15 vegetation classes is given in the results). a corrected rapideye image was also mapped using the maximum likelihood classifier in envi 4.8 (envi 2010). the two classified images were seamlessly mosaicked together, and an area of interest was extracted by mask in arcmap 10.2 (esri 2010). the area of each habitat was computed and converted into a minimum mapping unit in arcmap 10.2 (esri 2010). results mapping for mapping purposes, we required a habitat map that presented functionally distinct classes for herbivores. thus, we grouped functionally similar classes (which, in addition, were generally difficult to distinguish from each other through remote sensing and thus difficult to map as separate communities), described by the cluster analysis (figure 1-a1) and shown in the nms (figure 3). for example, the six sandveld types are not easily distinguished from each other through remote sensing and are functionally similar for herbivores, thus they can be mapped as one unit. by contrast, the communities on alluvial soils were very different functionally (e.g. wetland vs. mopane or riparian forests vs. dry floodplains) and could be easily distinguished through remote sensing and thus were mapped separately. with regard to the communities on heavy lacustrine clay soils, two distinct, functionally different communities occur, (1) the short grasslands with acacia spp. on silty soils and (2) the tall, open grasslands on vertisols, thus, functionally for herbivores, we chose to map them as two classes. classification a coincident minimum of isa p-value and maximum number of significant indicator species were found in seven and nine communities, respectively. another coincident minimum of isa p-value and maximum number of significant indicator species were found in 15 communities and were considered a meaningful ecological level of communities for a detailed vegetation classification. thus, hierarchical cluster analysis done on all 801 vegetation plots recognised 15 main vegetation communities within which many sub-communities occurred (figure 1-a1). mrpp tests of 15 communities suggested significant differences between communities (p < 0.000, table 1-a1) with chance-corrected within group agreement, a = 0.265. pairwise comparisons between communities suggested significant differences (p < 0.000, table 1-a1). the first axis of the nms1 analysis appears to be a gradient of soil texture and fertility, with communities on the most sandy soils having the most negative values, communities on loam soils having intermediate values and communities on extremely high clay soils having the most positive values on nms1, respectively (figure 3; table 1). the second nms axis (nms2) appears to be a weak gradient of wetness, with communities having the most positive values on nms2 being those that receive some sort of seasonal flooding from the annual flood pulse into the okavango and linyanti systems (setaria sphacelata – gomphocarpus fruticosus community) or seasonal rainfall (setaria incrassata – dichanthium annulatum community) or occur near permanent water with perhaps a shallow water table (tribulus terrestris – senna obtusifolia community) (authorities for plant species names are according to http://www.theplantlist.org). table 1: soil texture and nutrient characteristics in the savuti-mababe-linyanti ecosystem, northern botswana. vegetation on kalahari sands the nms (figure 3) delineated several vegetation communities in the smle that occur on deep kalahari sands, and they appear to be differentiated according to subtle variation in the silt and clay content of the soil (table 1). in the vegetation or habitat map, we refer to these communities on deep sands as sandveld communities (figure 2). although baikiaea plurijuga – baphia massaiensis is also found on deep sands (table 1), we refer to it in the map as baikiaea forest, not as sandveld (figure 2). figure 2: map of savuti-mababe-linyanti ecosystem vegetation (northern botswana). figure 3: non-metric multidimensional scaling ordination of plant communities on 801 plots. baikiaea plurijuga – baphia massaiensis this community is associated with high sand and low clay content of the soil (table 1). the community is extensive and occurs from northern edge of the beachhead of the md near ghoha hills and extends east all the way north-east into zimbabwe (figure 2). it is also found in small patches near the eastern edge of the okavango delta from the tsum tsum floodplains to the vumbura–motswiri region and in the woodlands west of the kwando system (figure 2). indicator species include baikiaea plurijuga, baphia massaiensis, croton gratissimus, hibiscus lobatum, panicum maximum, thunbergia reticulata and combretum apiculatum (table 2). baikiaea forests, while not supporting a high density of herbivores, are critical habitats for rare herbivores such as roan and sable antelope, and thus play a key role in the functional heterogeneity of the region. table 2: indicator species characteristics for 15 class divisions of plant communities in the savuti-mababe-linyanti ecosystem (northern botswana). table 2(continues…): indicator species characteristics for 15 class divisions of plant communities in the savuti-mababe-linyanti ecosystem (northern botswana). table 2(continues…): indicator species characteristics for 15 class divisions of plant communities in the savuti-mababe-linyanti ecosystem (northern botswana). apart from the baikiaea forest on deep sands, there are six communities which we have collectively mapped as sandveld (figure 2), and these are discussed below (in order of increasing nms axis 1 value). ipomoea chloroneura – oxygonum alatum this is one of the most extensive communities occurring on deep sands, where aeolian sands have infilled ancient river channels among the alluvial soils (supporting mopane), in the extensive woodlands between the okavango delta and the linyanti swamps (figure 2). this community is associated with soils among the highest sand and lowest silt and clay contents, as well as the lowest phosphorus, potassium, sodium and magnesium contents (table 1), and is recognised by the dominance of terminalia sericea in the woody layer. it is one of the most important wet season habitats for several tall grass grazers because of the abundance of high-quality tall grasses such as digitaria eriantha. indicator species in this community are ipomoea chloroneura, oxygonum alatum, hibiscus mastersianus, chamaecrista stricta, erlangea misera, chamaecrista absus, ceratotheca sesmoides, philenoptera nelsii, pavonia senegalensis and basananthe pedata (table 2). eragrostis pallens – ochna pulchra similar to the ipomoea chloroneura – oxygonum alatum community, the eragrostis pallens – ochna pulchra community is associated with soils among the highest sand content and lowest phosphorus, potassium, sodium and magnesium contents (table 1). like the previous community, it is recognised by the dominance of t. sericea in the woody layer; however, eragrostis pallens, ochna pulchra, burkea africana, aristida stipitata, phyllanthus burchellii, d. eriantha, dicoma schinzii, euphorbia critonoides and phyllanthus mendesii are some of the characteristic species which separate it from the previous community (table 2). commiphora angolensis – combretum collinum this community is spatially separated from other sandveld communities, being found only south-east of the md and appears to be an extensive kalahari community, extending towards nxai pan and the central kalahari game reserve (figure 2). it is associated with soils of the lowest clay contents and highest sand contents, as well as the lowest phosphorus, potassium, sodium and magnesium contents (table 1), and and is characterised by woody species not seen in typical sandveld communities west of the md, such as commiphora angolensis, combretum collinum and senegalia (acacia) ataxacantha. this community’s position in the nms (figure 3), however, suggests that its plant composition is more similar to the communities with soils of intermediate clay content, and therefore, we suspect that either there is more clay in the subsoil (we only sampled the topsoil) or the sands are not as deep as in the previous two communities. the structure is also different from the other sandveld types, which are relatively tall woodlands, whereas commiphora angolensis – combretum collinum sandveld is more of a shrubland, which may also suggest that the soils are not as deep. indicator species include c. angolensis, c. collinum, s. ataxacantha, combretum molle, evolvulus alsinoides, neorautanenia amboensis, xenostegia tridentata, waltheria indica, ochna serrulata, bauhinia petersiana, duosperma crenatum, acanthosicyos naudinianus and commiphora africana (table 2). it serves as a key wet season habitat for tall grass grazers such as buffalo, eland and roan antelope, probably because it occurs far from any permanent water sources (figure 2) and, therefore, has among the highest cover of the high-quality grass d. eriantha. boscia albitrunca – dactyloctenium giganteum this community, although still a sandveld community on sandy soils, was associated with higher clay and calcium and phosphorus contents than other sandveld types (similar to brachiaria nigropedata – combretum hereroense community in clay content) (table 1). it is mainly found around the peripheries of paleolake mababe (md) (figure 2), where there was some moderate deposition of clay and silt by the paleolake waters. indicator species are boscia albitrunca, dactyloctenium giganteum, indigofera flavicans, sida chrysantha, mundulea sericea, ocimum gratissimum, digitaria ternata and cleome hirta (table 2). there is an abundance of the high-quality grazing grass d. giganteum which, together with the important browse provided by b. albitrunca, results in this community type being a key wet season habitat for herbivores such as buffalo and a dry season habitat for eland and other browsers in the smle. brachiaria nigropedata – combretum hereroense the brachiaria nigropedata – combretum hereroense community, similar to the boscia albitrunca – dactyloctenium giganteum community, was associated with some moderate degree of ancient sediment deposition in the peripheries of paleolake mababe and around the kwando-linyanti system, resulting in more clay than in the other sandveld community types (table 1). it serves as a key wet season habitat for buffalo because of an abundance of one of their most favoured grasses, b. nigropedata. indicator species include b. nigropedata, c. hereroense, lantana angolensis, andropogon gayanus, dalbergia melanoxylon, schmidtia pappophoroides and polydora poskeana (table 2). tribulus terrestris – senna obtusifolia the tribulus terrestris – senna obtusifolia community occurs on deep sands but always adjacent to riparian vegetation (croton megalobotrys – setaria verticillata and justicea divaricata – eragrostis superba communities), found near water bodies of the okavango delta, linyanti swamps, selinda spillway and the savuti channel. although the sands of this community are deep, they have higher clay content (table 1), which again is likely associated with ancient sediment deposition from previous water bodies or extreme floods. this community is recognised by the dominance of the woody layer by tall p. nelsii (kalahari appleleaf) and with very little t. sericea occurrence (as opposed to other sandveld communities where t. sericea is common), which is likely because of the higher clay content in these sandy soils. indicator species included t. terrestris, s. obtusifolia, sida cordifolia, dichrostachys cinerea and zehneria marlothii (table 2). vegetation on alluvial loam soils these communities occur at intermediate levels of silt and clay (as compared with the lower levels of silt and clay of the sandveld communities) as a result of ancient alluvial deposition of sediments. colophospermum mopane – jasminum stenolobum this community is recognised by the dominance of the woody layer by c. mopane, which appears to dominate once the clay content of the soil reaches a critical level but not too high where other communities exist (table 1). this is an extensive community that occurs all over the ecosystem, near and further from permanent water (figure 2). seasonal waterholes occur in this community and serve as drinking points for various herbivores during the wet season and grasses, while sparse, are often palatable. indicator species in mopane woodland include c. mopane, j. stenolobum, tragus berteronianus, zornia glochidiata, kyllinga buchananii, brachiaria deflexa and aristida adscensionis (table 2). setaria sphacelata – gomphocarpus fruticosus this community was associated with high silt–clay and intermediate sand contents of the soil (table 1), but the key feature driving species composition is that it is seasonally inundated by floodwaters from the major wetland systems of the region. the community is common in moderately-flooded zones within the overall wetland system shown in the map (figure 2). however, our focus was on the dryland communities, and our sampling regime in wetlands was limited and not designed to capture the variation in wetland community composition on flooding depth and duration gradients; hence, this community represents one of many different wetland community types. characteristic species of this community type included setaria sphacelata, gomphocarpus fruticosus, cyperus longus, crotalaria platysepala, momordica balsamina, sesamum triphyllum and bulbostylis hispidula (table 2). wetlands provide critical dry season grazing for herbivores because of their shallow water tables and the ability to provide green forage for herbivores during the dry season. justicea divaricata – eragrostis superba this is an open grassland community found directly adjacent to floodplains (mapped as dry floodplains in the map) along the major water bodies (figure 2) and is associated with moderate silt–clay, and by intermediate sand, phosphorus, potassium and calcium contents of the soil (table 1). although this community is rarely (if ever) subject to flooding, it is probably maintained in a treeless state by high water tables from the adjacent floodplains. indicator species include justicea divaricata, eragrostis superba, eragrostis trichophora, geigeria schinzii, orthanthera jasminiflora and justicia betonica (table 2). croton megalobotrys – setaria verticillata this is a riparian woodland community (figure 2), associated with major water bodies in the region (table 1). species including croton megalobotrys, setaria verticillata, diospyros mespiliformis, philenoptera violacea, senegalia (acacia) nigrescens and asystasia gangetica characterised this community (table 2). riparian woodlands can occur as open or closed woodlands. open riparian woodlands are structurally characterised by open canopy woodlands dominated by p. violacea, s. nigrescens and c. imberbe, whereas closed riparian woodlands are characterised by tall, closed canopy woodlands dominated by c. megalobotrys, d. mespiliformis, p. violacea, s. nigrescens and c. imberbe. in the map, these two riparian woodland communities were mapped as riparian woodlands (figure 2). vegetation on heavy clay lacustrine deposits of the sump of the mababe depression chloris virgata – boerhavia coccinea the chloris virgata – boerhavia coccinea community occurs on the more silty, intermediate clay soils on the edge of the sump of the md (as well as in areas adjacent to the linyanti swamps and kwando river) and was associated with high silt, phosphorus and calcium (table 1). it is included under acacia grasslands in the vegetation map (figure 2), named as such because these communities are characterised by short grasslands with acacia species in the woody layer (of varying tree density). this is a widespread community that develops where the clay content reaches higher levels than that found in alluvial soils. thus, this community type is associated with water bodies that could have deposited those clays such as the paleolake mababe or the kwando/linyanti swamps (figure 2). this community was characterised by chloris virgata, boerhavia coccinea, vachellia (acacia) tortilis, hermannia kirkii, vachellia (acacia) hebeclada, cucumis anguria, dicoma tomentosa and aerva leucura (table 2). the community is dominated by the highly palatable annual grasses, c. virgata and urochloa trichopus, and forms a critical wet season grazing resource on the edge of the sump of the md for migratory zebra, as well as for tsessebe, impala and wildebeest. one of the key functional features of this community for herbivores is that it offers the highest phosphorus levels in soils and grasses of all community types in the ecosystem (table 1). as one moves off the silty soils on the edge of the sump of the md, which support the chloris virgata – boerhavia coccinea community, towards the centre of the md, soils become deep, heavy black vertisols dominated by taller grasses such as cenchrus ciliaris and bothriochloa insculpta. the vegetation on these vertisols is mapped as tall, open grasslands (figure 2) and consists of two main communities, (1) the cenchrus ciliaris – senegalia (acacia) mellifera community and (2) the bothriochloa insculpta – rhynchosia minima community. cenchrus ciliaris – senegalia mellifera this open savanna grassland community occurs on deep black clay soils (vertisols) deeper into the md and was associated with soils among the lowest sand content and highest silt–clay content and importantly had by far the highest soil calcium and potassium content in the smle (table 1). indicator species are c. ciliaris, s. mellifera, indigofera scaberrima and euphorbia polycephala (table 2). the very high soil fertility of the cenchrus ciliaris – senegalia mellifera community (table 1) makes this habitat, together with the chloris virgata – boerhavia coccinea community, which are directly adjacent to each other, an extremely important wet season range for the large migratory zebra population. thus, these two md vegetation communities make up a critical part of the functional heterogeneity of the region. bothriochloa insculpta – rhynchosia minima this open savanna grassland community occurs on similar soils to the previous community (table 1), but it makes up much less area on the md. bothriochloa insculpta, rhynchosia minima, cyathula orthacantha, leonotis nepetifolia and dinebra retroflexa characterised this community (table 2). dominance by the unpalatable b. insculpta makes this community type less important for wildlife. setaria incrassata – dichanthium annulatum this open grassland community also occurs on the md, being found as vast, extremely open grasslands in the far north-eastern part of the md, and is characterised by seasonal flooding from rainfall. soils in this community are not as high clay as others on the md (table 1); however, from a structural point of view, the vegetation is mapped as tall, open grassland because it is a tall grass community although very different in composition to the other tall grass communities on the md (figure 2). setaria incrassata, dichanthium annulatum, enicostema axillare, panicum coloratum, cynodon dactylon, digitaria milanjiana and aristida scrabrivalvis are some of the species that characterised this community (table 2). the community appears to be utilised by many roan antelope over the dry season. discussion vegetation classification and mapping of the savuti-mababe-linyanti ecosystem the vegetation of the smle was clustered into 15 major vegetation communities characterised by different herbaceous and woody species (figure 3), with the primary axis of variation in plant community composition (nms1) appearing to be driven by variation in soil texture and fertility (figure 3; table 1). the secondary axis of variation in plant community composition (nms2) appeared to be driven by wetness, although it was not a clear effect as with the texture gradient probably because we focused mainly on dryland communities. the landscape template, which provides the basis of the vegetation heterogeneity of the region, was formed by a variety of processes. the mosaic of sand-filled paleoriver channels among alluvial deposits that supports the mopane–sandveld woodland mosaic between the okavango delta and the linyanti swamps was formed by ancient wetlands similar to the current delta, with the river channels subsequently becoming infilled by kalahari sands of aeolian origin (mccarthy, gumbricht & mccarthy 2005). similarly, the deep sands supporting the baikiaea forests are of aeolian origin (mccarthy et al. 2005). the vast 3000 km2 md originates from paleolake mababe (teter 2007), which has a central sump of about 70 km × 20 km, where lacustrine clays and sediments deposited, giving rise to the deep vertisols of the md. between the beachhead of the lake and the sump zone is a zone of soils where it appears that deposition of lacustrine clays declined with increasing distance from the sump zone. thus, the inner sump has soils of the highest clay content supporting the bothriochloa insculpta – rhynchosia minima and cenchrus ciliaris – senegalia mellifera communities, followed by the edges of the sump with more silty soils supporting the chloris virgata – boerhavia coccinea community, then a zone of silty sands supporting mopane woodland and between the mopane and the beachhead of the lake are sandy soils but with higher clay and silt than the typical aeolian sands, which support the boscia albitrunca – dactyloctenium giganteum and brachiaria nigropedata – combretum hereroense sandveld communities (figure 2). outside the md, the typical aeolian sands have the lowest clay and silt contents because they received no lacustrine sediment deposition. the extremely low clay sands support the ipomoea chloroneura – oxygonum alatum, eragrostis pallens – ochna pulchra and commiphora angolensis – combretum collinum sandveld communities. species such as c. angolensis, c. collinum, b. plurijuga, b. massaiensis, p. nelsii, o. pulchra, b. africana and t. sericea which dominated communities growing on sandy soils in this study were also found as indicator species for sandy regions in other studies (coetzee et al. 1976; gertenbach & potgieter 1978; tedder 2012; tedder et al. 2013). the mopane community was associated with alluvial soils of slightly higher clay than the sandveld communities (table 1), as observed elsewhere (tedder 2012; tedder et al. 2013; van voorthuizen 1976; wolski & murray-hudson 2006). the mopane and sandveld communities are widely distributed across the ecosystem, forming a woodland mosaic of mopane woodland on alluvial soils alternating with sandveld woodland on paleoriver channels infilled with kalahari sands (figure 2). the mopane–sandveld mosiac (as well as baikiaea forest) provides key habitat for herbivore species favouring medium and tall grasses, such as buffalo, roan and sable antelope and elephant (bennitt, bonyongo & harris 2014, 2015; fynn et al. 2014; sianga 2014; taolo 2003), because of the abundance of digestible, leafy forage of high-quality grass species dominant in sandveld woodland (e.g. d. eriantha, panicum maximum and schmidtia papophoroides), as well as d. milanjiana and panicum maximum in mopane woodland (sianga et al. 2017b). importantly, these high-quality grasses are most abundant far from permanent water (>15 km – 20 km) (sianga et al. 2017b), which explains why buffalo tend to favour the woodlands furthest from permanent water during the wet season (bennitt et al. 2014; sianga 2014; sianga, fynn & bonyongo 2017a; sianga et al. 2017b), as do roan and sable antelope (havemann 2014; hensman et al. 2014). in addition, the numerous ephemeral waterholes of the mopane woodland allow herbivores to remain far out from permanent water in these woodlands during the wet season where they are able to avoid high concentrations of predators (harrington et al. 1999). once the waterholes dry up during the dry season, herbivores are forced to move closer to the permanent water sources of the okavango delta and linyanti swamps, where buffalo forage mainly in the wetlands (bennitt et al. 2014, 2015; sianga 2014; sianga et al. 2017a), whereas roan and sable antelope visit the wetlands only every 3–4 days to drink and then return back to the safety of the woodlands far from water (havemann 2014; hensman et al. 2014). thus, these vast woodland systems provide key functional habitat heterogeneity for provision of high-quality forage far from water during the wet season and low predation risk all year round. the croton megalobotrys – setaria verticillata community, a riparian woodland (riverine) community, was correlated with silt–clay soils and occurred along edges of watercourses (witkowski & o’connor 1996). this community was dominated by species such as croton megalobotrys, p. violacea, combretum mossambicense, d. mespiliformis and s. nigrescens, which are adapted to obtaining soil moisture through lateral ground-water discharge from higher water tables (ellery, ellery & mccarthy 1993; hamandawana 2011; ringrose et al. 2007). the setaria sphacelata – gomphocarpus fruticosus community, a floodplain grassland type found along watercourses, had grass (s. sphacelata) and sedge (c. longus) or forbs (g. fruticosus) as indicator species suggesting extensive wetness, as these species are mostly abundant in wetlands or swamp margins (heath & heath 2009). this community probably experiences periods of dryness over the annual cycle as indicated by the presence of opportunistic dryland species (c. platysepala, m. balsamina, s. triphyllum and b. hispidula). variation in wetland community composition is driven by gradients of flood depth and duration, with c. dactylon often characterising the parts of the gradient with the lowest depth and duration of flooding, panicum repens and s. sphacelata often characterising areas with intermediate depth and duration of flooding and tall sedges and grasses such as oryza longistaminata and vossia cuspidata characterising areas of the gradient with the largest depth and duration of flooding (fynn et al. 2015; murray-hudson et al. 2011, 2014). this variation in composition and phenology on flooding gradients provides important variation in green forage supply for herbivores from the early to late dry season, owing to variation in availability of soil moisture for growth, allowing for adaptive foraging over the dry season (fynn et al. 2015). our sampling focused on the dryland communities, and we did not attempt to sample across the flood depth and duration gradient in the wetlands; thus, the setaria sphacelata – gomphocarpus fruticosus community represents only a small part of the variation in plant community composition that would occur in the region mapped as wetland in figure 2. the spatial location of various wetland community types is not a constant and will shift location within the bounds of the area mapped as wetland (figure 2) according to variation in flooding regimes over time. from a conservation management perspective, however, it should be recognised that the overall wetland community shown in the map (figure 2) represents gradients of wetness and composition that provide critically important forage and adaptive foraging options for many herbivore species from the early to late dry season right across africa (fynn et al. 2015) and in the smle (bartlam-brooks, bonyongo & harris 2013; bennitt et al. 2014; fynn et al. 2014; sianga et al. 2017a). consequently, linkages between large wetland systems and adjacent dryland systems must be maintained to ensure that functional habitat heterogeneity is maintained (fynn et al. 2015; hopcraft et al. 2010). communities on the high clay soils of the md, especially the chloris virgata – boerhavia coccinea and cenchrus ciliaris – senegalia mellifera communities, are critical wet season habitats for the large zebra migration in the region, as well as for wildebeest, tsessebe and impala (fynn et al. 2014; joos-vandewalle 2000; sianga 2014; sianga et al. 2017a). this is because of the high clay soils and accumulation of a high concentration of minerals in the soil when it was a lake system (teter 2007). the p-rich soils give rise to high p content in grass leaves (as well as other minerals) (fynn et al. 2014; joos-vandewalle 2000; sianga 2014). thus, pregnant and lactating herbivores can obtain sufficient intake of nutrients to meet their high demands for nutrients during the wet season, a key functional aspect of wet season ranges for herbivores (hopcraft et al. 2010; owen-smith 2004). in this regard, the chloris virgata – boerhavia coccinea community on the edge of the md appears to be particularly important for p, having the highest soil p levels (table 1), while the vertisols of the cenchrus ciliaris – senegalia mellifera communities deeper into the md have the highest concentrations of potassium and calcium. this may explain why zebra are observed to switch their foraging bouts between chloris virgata – boerhavia coccinea community and the cenchrus ciliaris – senegalia mellifera community over the day (sianga 2014), which may be a mechanism to maximise overall intake of key minerals, protein and energy during the wet season (owen-smith 2002). also, the open grasslands of the md provide better visibility, which reduces predation risk and is thus suitable as a calving ground. in fact, selection for low predation-risk habitat for the calving period may dominate the hierarchy of habitat selection decisions made by ungulates (rettie & messier 2000). during the day, zebra appeared to make use of short, open grasslands with high visibility (mainly the chloris virgata – boerhavia coccinea community) but at night, they moved further into the md to the s. mellifera woodlands perhaps as an adaptive strategy to elude predators – reliance on sighting predators during the day and hiding from them at night (sianga 2014). dominance of much of the southern half of the md by c. ciliaris is informative as this species is often found in areas with elevated levels of p in soils (blackmore, mentis & scholes 1990), which demonstrates why the md is an important wet season range for herbivores. the bothriochloa insculpta – rhynchosia minima community is dominated by b. insculpta, which is adapted to seasonally flooded vertisols which form hard surfaces during winter (cook & clem 2000). the well-developed root systems of these large perennial grasses (b. insculpta and c. ciliaris) also likely promote access to soil moisture from deeper water tables when surface water dries out during the dry season (cook & clem 2000; hamandawana 2011). the setaria incrassata – dichanthium annulatum community was dominated mainly by grass species such as s. incrassata, d. annulatum, p. coloratum, c. dactylon and d. milanjiana and occurred on seasonally flooded vertisols, with seasonal flooding maintaining this community as an open grassland with no trees (cook & clem 2000). species such as s. incrassata and d. annulatum are well-known dominants of seasonally flooded, heavy clay soils in southern africa (cook & clem 2000). this community type was found only in the north-eastern part of the md and occurs in one of the most remote and inaccessible areas of northern botswana, rarely ever accessed by people because there are no roads there and seasonal flooding of the heavy clay soils makes access impossible during the wet season. our many sightings of roan antelope in this vegetation type while we were sampling suggest that it is regularly used by roan antelope, which is a species known to favour seasonally flooded grasslands and prefers areas with little disturbance by people. conclusion this study demonstrated that large heterogeneity of plant communities driven by gradients in soil texture or fertility and wetness plays a key role in providing critical functional resource and habitat heterogeneity that allows (1) herbivores to adapt to seasonal variation in resources and (2) allows niche diversity to support a diverse guild of herbivores. floodplains and seasonally flooded grasslands provide a reliable source of green forage during the dry season for herbivores, whereas the vast woodland mosaic of mopane and sandveld on alluvial soils and kalahari sands provides cover, low predation risk and medium height, leafy grasses for rare herbivores all year round and for buffalo and elephant during the wet season. the fertile heavy clay soils of the md provide additional functional resource heterogeneity in an ecosystem otherwise largely dominated by sandy soils, where elevated levels of minerals such as ca and p in grasses on the md enable pregnant and lactating females to meet their elevated requirements for these resources. thus, the regional-scale separation of large wetland systems (a functional dry season habitat) from large woodland systems and the fertile clay soils of the md (functional wet season habitats) is an underlying driver of both buffalo (wetlands to woodlands) and zebra (wetlands to paleolake systems) migrations in the smle (bennitt et al. 2014; naidoo et al. 2014; sianga 2014). certain communities such as the boscia albitrunca – dactyloctenium giganteum community provide a high density of trees with green leaves of acceptable quality during the dry season (e.g. b. albitrunca), which provide an important reserve resource for browsers at this time of the year (owen-smith 2002). thus, the heterogeneous mix of extensive sandveld, mopane and baikiaea woodlands, open fertile grasslands and extensive wetland systems provides excellent functional habitat and resource heterogeneity in the ecosystem, which enables herbivores to adapt to variable resources needs and avoid predation. in addition, this heterogeneity creates niche diversity, which enables coexistence of a high diversity of large mammals (herbivores and carnivores), including one of the largest roan and sable antelope and eland populations in southern africa. thus, our results demonstrate that a key point of general significance for conservation science is that protected areas need to cover the main large-scale regional environmental gradients in a region (e.g. the full texture and wetness gradients in this study region) if they are to provide sufficient habitat heterogeneity needed to provide appropriate seasonal adaptive foraging options for wildlife and to support a diverse guild of herbivores and their associated predators. in addition, this vegetation classification and vegetation map will provide a critical database for wildlife habitat selection studies in the region and will be useful for environmental and conservation policy-makers in the assessment and monitoring of plant communities as well as for developing conservation strategies and management plans for the ecosystem. acknowledgements the ministry of environment, wildlife and tourism and the department of wildlife and national parks (botswana) are highlighted for permitting us to conduct this research in northern botswana (ewt 8/36/4 xxv [16]). the authors finally acknowledge mrs. sibonile sibanda of hatfield consultants (gaborone, botswana) and mrs. marion stelles and dr achim roder (university of trier, germany) for their invaluable guidance with the remote sensing and vegetation mapping work. great plains safaris (selinda reserve) and okavango wilderness safaris (botswana) are acknowledged for their logistic support. the authors acknowledge the southern african science service centre for climate change and the adaptive land management (sasscal) for funding this research. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions all authors directly participated in the planning, execution and analysis of this study. references bartlam-brooks, h.l.a., bonyongo, m.c. & harris, s., 2013, ‘how landscape scale changes affect ecological processes in conservation areas: external factors influence land use by zebra (equus burchellii) in the okavango delta’, ecology and evolution 3(9), 2795–2805. https://doi.org/10.1002/ece3.676 bennitt, e., bonyongo, m.c. & harris, s., 2014, ‘habitat selection by african buffalo (syncerus caffer) in response to landscape-level fluctuations in water availability on two temporal scales’, plos one 9(7), e101346. https://doi.org/10.1371/journal.pone.0101346 bennitt, e., bonyongo, m.c. & harris, s., 2015, ‘behaviour-related scalar habitat use by cape buffalo (syncerus caffer)’, plos one 10(12), e0145145. https://doi.org/10.1371/journal.pone.0145145 blackmore, a.c., mentis, m.t. & scholes, r.j., 1990, ‘the origin and extent of nutrient-enriched patches within a nutrient-poor savanna in south africa’, journal of biogeography 17(4/5), 463–470. https://doi.org/10.2307/2845378 coetzee, b., van der meulen, f., zwanziger, s., gonsalves, p. & weisser, p., 1976, ‘a phytosociological classification of the nylsvley nature reserve’, bothalia 12(1), 137–160. https://doi.org/10.4102/abc.v12i1.1388 cook, b. & clem, r., 2000, ‘which grass for where?’, tropical grasslands 34, 156–161. dufrene, m. & legendre, p., 1997, ‘species assemblages and indicator species: the need for a flexible asymmetrical approach’, ecological monographs 67(3), 345–366. https://doi.org/10.2307/2963459 ellery, w.n., ellery, k. & mccarthy, t.s., 1993, ‘plant distribution in islands of the okavango delta, botswana: determinants and feedback interactions’, african journal of ecology 31(2), 118–134. https://doi.org/10.1111/j.1365-2028.1993.tb00526.x envi version 4.8, 2010, computer software, itt corporation, boulder, co. esri version 10.2, 2010, computer software, environmental systems research institute, redlands, ca. fynn, r.w.s., chase, m. & roder, a., 2014, ‘functional habitat heterogeneity and large herbivore seasonal habitat selection in northern botswana’, south african journal of wildlife research 44(1), 1–15. https://doi.org/10.3957/056.044.0103 fynn, r.w.s., murray-hudson, m., dhliwayo, m. & scholte, p., 2015, ‘african wetlands and their seasonal use by wild and domestic herbivores’, wetlands ecology and management 23(4), 559–581. https://doi.org/10.1007/s11273-015-9430-6 gertenbach, w. & potgieter, a., 1978, ‘a phytosociological classification of the hlane wildlife sanctuary, swaziland’, koedoe 21(1), 47–65. https://doi.org/10.4102/koedoe.v21i1.960 hamandawana, h., 2011, ‘methodology to detect long-term trends in groundwater by monitoring changes in vegetation distribution’, international journal of remote sensing 32(12), 3329–3343. https://doi.org/10.1080/01431161003747497 harrington, r., owen-smith, n., viljoen, p.c., biggs, h.c., mason, d.r. & funston, p., 1999, ‘establishing the causes of the roan antelope decline in the kruger national park, south africa’, biological conservation 90, 69–78. https://doi.org/10.1016/s0006-3207(98)00120-7 havemann, c.p., 2014, ‘population dynamics and foraging ecology of roan antelope in northern botswana’, msc thesis, university of pretoria, pretoria, south africa. heath, r. & heath, a., 2009, field guide to the plants of northern botswana including the okavango delta, royal botanic gardens, richmond. hensman, m.c., owen-smith, n., parrini, f. & bonyongo, c.m., 2014, ‘home range occupation and habitat use of sable antelope in the okavango delta region of northern botswana’, african journal of ecology 52(2), 237–245. https://doi.org/10.1111/aje.12111 hobbs, n.t., galvin, k.a., stokes, c.j., lackett, j.m., ash, a.j., boone, r.b. et al., 2008, ‘fragmentation of rangelands: implications for humans, animals, and landscapes’, global environmental change-human and policy dimensions 18, 776–785. https://doi.org/10.1016/j.gloenvcha.2008.07.011 hopcraft, j.g.c., olff, h. & sinclair, a., 2010, ‘herbivores, resources and risks: alternating regulation along primary environmental gradients in savannas’, trends in ecology & evolution 25(2), 119–128. https://doi.org/10.1016/j.tree.2009.08.001 hopcraft, j.g.c., sinclair, a.r.e. & packer, c., 2005, ‘planning for success: serengeti lions seek prey accessibility rather than abundance’, journal of animal ecology 74, 559–566. https://doi.org/10.1111/j.1365-2656.2005.00955.x joos-vandewalle, m.e.j., 2000, ‘movements of migratory zebra and wildebeest in northern botswana’, phd thesis, university of the witwatersrand, johannesburg, south africa. macfayden, s., hui, c., verberg, p.h. & van teeffelen, a.j.a., 2016, ‘quantifying spatiotemporal drivers of environmental heterogeneity in kruger national park, south africa’, landscape ecology. early view 31, 2013–2029. mccarthy, j., gumbricht, t. & mccarthy, t.s., 2005, ‘ecoregion classification in the okavango delta, botswana from multitemporal remote sensing’, international journal of remote sensing 26(19), 4339–4357. https://doi.org/10.1080/01431160500113583 mccune, b., grace, j.b. & urban, d.l., 2002, analysis of ecological communities, mjm software design, gleneden beach, or. mendelsohn, j., vanderpost, c., ramberg, l., murray-hudson, m., wolski, p. & mosepele, k., 2010, okavango delta: floods of life, raison, windhoek. mills, m.g.l. & gorman, m., 1997, ‘factors affecting the density and distribution of wild dogs in the kruger national park’, conservation biology 11, 1397–1406. https://doi.org/10.1046/j.1523-1739.1997.96252.x murray-hudson, m., combs, f., wolski, p. & brown, m.t., 2011, ‘a vegetation-based hierarchical classification for seasonally pulsed floodplains in the okavango delta, botswana’, african journal of aquatic science 36, 223–234. https://doi.org/10.2989/16085914.2011.636904 murray-hudson, m., wolski, p., murray-hudson, f., brown, m.t. & kashe, k., 2014, ‘disaggregating hydroperiod: components of the seasonal flood pulse as drivers of plant species distribution in floodplains of a tropical wetland’, wetlands 34, 927–942. https://doi.org/10.1007/s13157-014-0554-x naidoo, r., du preez, p., stuart-hill, g., beytell, p. & taylor, r., 2014, ‘long-range migrations and dispersals of african buffalo (syncerus caffer) in the kavango–zambezi transfrontier conservation area’, african journal of ecology 52, 581–584. https://doi.org/10.1111/aje.12163 owen-smith, n., 2002, adaptive herbivore ecology, cambridge university press, cambridge. owen-smith, n., 2004, ‘functional heterogeneity in resources within landscapes and herbivore population dynamics’, landscape ecology 19(7), 761–771. https://doi.org/10.1007/s10980-005-0247-2 rettie, w.j. & messier, f., 2000, ‘hierarchical habitat selection by woodland caribou: its relationship to limiting factors’, ecography 23(4), 466–478. https://doi.org/10.1111/j.1600-0587.2000.tb00303.x ringrose, s., 2006, report: results and struggles of the elephant habitat mapping project, university of botswana, botswana. ringrose, s., vanderpost, c., matheson, w., wolski, p., huntsman-mapila, p., murray-hudson, m. et al., 2007, ‘indicators of desiccation-driven change in the distal okavango delta, botswana’, journal of arid environments 68(1), 88–112. https://doi.org/10.1016/j.jaridenv.2006.03.030 sianga, k., 2014, ‘habitat use by buffalo and zebra in relation to spatial and temporal variability of resources in the savuti-mababe-linyanti ecosystem of northern botswana’, mphil thesis, university of botswana, botswana. sianga, k., fynn, r.w.s. & bonyongo, m.c., 2017a, ‘seasonal habitat selection by african buffalo syncerus caffer in the savuti–mababe–linyanti ecosystem of northern botswana’, koedoe 59(2), a1382. https://doi.org/10.4102/koedoe.v59i2.1382 sianga, k., van telgen, m., vrooman, j., fynn, r.w.s. & van langevelde, f., 2017b, ‘spatial refuges buffer landscapes against homogenisation and degradation by large herbivore populations and facilitate vegetation heterogeneity’, koedoe 59(2), a1434. https://doi.org/10.4102/koedoe.v59i2.1434. taolo, c., 2003, ‘population ecology, seasonal movement and habitat use of the african buffalo (syncerus caffer) in chobe national park, botswana’, phd thesis, norwegian university of science and technology, trondheim, norway. tedder, m.j., 2012, ‘dry woodland and savanna vegetation dynamics in the eastern okavango delta, botswana’, phd thesis, university of kwazulu-natal, pietermaritzburg. tedder, m.j., kirkman, k.p., morris, c.d., trollope, w.s.w. & bonyongo, m.c., 2013, ‘classification and mapping of the composition and structure of dry woodland and savanna in the eastern okavango delta’, koedoe 55(1), 1–8. https://doi.org/10.4102/koedoe.v55i1.1100 teter, k.l., 2007, ‘paleoenvironmental reconstruction of paleolake mababe, northwestern botswana from sediment chemistry and biological productivity data’, msc thesis, oklahoma state university, stillwater, ok. the plant list, version 1.1. viewed 13 january 2017, from http://www.theplantlist.org van voorthuizen, e.g., 1976, ‘the mopane tree’, botswana notes and records 8, 223–230. witkowski, e. & o’connor, t., 1996, ‘topo-edaphic, floristic and physiognomic gradients of woody plants in a semi-arid african savanna woodland’, vegetatio 124(1), 9–23. https://doi.org/10.1007/bf00045140 wolski, p. & murray-hudson, m., 2006, ‘flooding dynamics in a large low-gradient alluvial fan, the okavango delta, botswana, from analysis and interpretation of a 30-year hydrometric record’, hydrology and earth system sciences discussions 10(1), 127–137. https://doi.org/10.5194/hess-10-127-2006 appendix 1 table 1-a1: multiresponse permutation procedure pairwise comparisons between identified classes for plant communities in the savuti-mababe-linyanti ecosystem (northern botswana). table 1-a1 (continues…): multiresponse permutation procedure pairwise comparisons between identified classes for plant communities in the savuti-mababe-linyanti ecosystem (northern botswana). figure 1-a1: cluster analysis showing the 15 plant communities identified in the savuti-mababe-linyanti ecosystem, northern botswana. article information authors: armin h. seydack1 cornelia c. grant2 izak p. smit2 wessel j. vermeulen1 johan baard1 nick zambatis2 affiliations: 1scientific services, garden route national park, south african national parks, south africa 2scientific services, kruger national park, south african national parks, south africa correspondence to: armin seydack postal address: po box 3542, knysna 6570, south africa dates: received: 08 mar. 2011 accepted: 05 oct. 2011 published: 08 feb. 2012 how to cite this article: seydack, a.h., grant, c.c., smit, i.p., vermeulen, w.j., baard, j. & zambatis, n., 2012, ‘large herbivore population performance and climate in a south african semi-arid savanna’, koedoe 54(1), art. #1047, 20 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1047 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. large herbivore population performance and climate in a south african semi-arid savanna in this original research... open access • abstract • introduction    • study area • materials and methods    • pattern/process matching    • patterns in herbivore population performance       • population performance trends       • spatiotemporal patterns       • climate and herbivore population performance    • climate-vegetation response model (process)       • development of an explanatory framework       • key forage resource indicators       • herbivore species guilds of nutritional requirements       • plant metabolic performance trade-off constraints       • spatial distribution of key forage resources (cvrm-1)       • climate-driven temporal trends in forage availability (cvrm-2)       • spatiotemporal interaction in forage availability (cvrm-3) • results    • patterns in long-term population performance trends       • historical population trends       • extant population trends    • landscape-scale spatial patterns in population performance    • climate and herbivore population performance       • buffalo and waterbuck       • blue wildebeest and zebra       • selectively feeding antelope species • discussion    • linking climate–vegetation responses with large herbivore population performance    • bottom-up and top-down impacts on large herbivore population dynamics • conclusion • acknowledgements    • competing interests    • authors’ contributions • references • footnote abstract top ↑ long-term population performance trends of eight large herbivore species belonging to groups of disparate foraging styles were studied in the semi-arid savanna of the kruger national park, south africa. over the past century the number of bulk feeders (buffalo, waterbuck, blue wildebeest and plains zebra) had increased towards comparatively high population densities, whereas population numbers of selectively feeding antelope species (sable antelope, roan antelope, tsessebe and eland) declined progressively. detailed analyses revealed that population numbers of buffalo and waterbuck fluctuated in association with food quantity determined by rainfall. population performance ratings (1944–2003) of the species for which forage quality was important (blue wildebeest, zebra and selective grazers) were correlated negatively with minimum temperature and positively with dry-season rainfall.interpretation according to a climate–vegetation response model suggested that acclimation of forage plants to increasing temperature had resulted in temperature-enhanced plant productivity, initially increasing food availability and supporting transient synchronous increases in population abundance of both blue wildebeest and zebra, and selective grazers. as acclimation of plants to concurrently rising minimum (nocturnal) temperature (tmin) took effect, adjustments in metabolic functionality occurred involving accelerated growth activity at the cost of storage-based metabolism. growth-linked nitrogen dilution and reduced carbon-nutrient quality of forage then resulted in phases of subsequently declining herbivore populations. over the long term (1910–2010), progressive plant functionality shifts towards accelerated metabolic growth rather than storage priority occurred in response to tmin rising faster than maximum temperature (tmax), thereby cumulatively compromising the carbon-nutrient quality of forage, a key resource for selective grazers. the results of analyses thus revealed consistency between herbivore population trends and levels of forage quantity and quality congruent with expected plant metabolic responses to climate effects. thus, according to the climate-vegetation response model, climate effects were implicated as the ultimate cause of large herbivore population performance in space and over time. conservation implications: in its broadest sense, the objective of this study was to contribute towards the enhanced understanding of landscape-scale functioning of savanna systems with regard to the interplay between climate, vegetation and herbivore population dynamics. introduction top ↑ the abundance of herbivore communities in african savannas is determined principally by the availability of food resources of adequate quality (east 1984; fritz & duncan 1994; sinclair, dublin & borner 1985). the influence of rainfall, especially during the dry season, on the availability of forage of adequate quality and large herbivore population performance has been recorded in numerous studies (inter alia de bie 1991; dunham, robertson & grant 2004; mduma, sinclair & hilborn 1999; mills, biggs & whyte 1995; owen-smith & ogutu 2003). however, in many studies the emphasis is placed on top-down control of herbivore populations through predation (owen-smith & mills 2006; owen-smith & mills 2008a). large herbivore population trends in the kruger national park (knp) over the past century have been characterised by a notable dichotomy: a progressive increase of the three main bulk-feeding large herbivore species, namely blue wildebeest (connochaetes taurinus), zebra (equus burchelli) and buffalo (syncerus caffer), stabilising at relatively high population densities, and the population decline of four selectively feeding species, namely sable antelope (hippotragus niger), roan antelope (hippotragus equinus), tsessebe (damaliscus lunatus lunatus) and eland (taurotragus oryx). initial population increases of bulk-feeding antelope and browsing species during the first half of the twentieth century were attributed largely to population recovery from decimation by hunting and epizootics (joubert 2007a; pienaar 1963). impacts by boundary fencing of the park, culling operations and water provisioning further shaped population abundance and spatial distributions of some species (joubert 2007a, 2007b). furthermore, ongoing impacts by predation and climate (rainfall) on large herbivore population abundance in the knp have been documented (dunham et al. 2004; harrington et al. 1999; joubert 2007a, 2007b; owen-smith & ogutu 2003; owen-smith, mason & ogutu 2005; owen-smith 2008; whyte & joubert 1988). the long-term and spatially widespread nature of population trends, however, suggested that over-arching landscape-scale influences may not as yet have been accounted for in currently available studies. a greater role of climate effects as a long-term landscape-scale factor was therefore implicated. in this context, dry-season rainfall and ambient temperature were considered important climate variables. ambient temperatures have been increasing progressively over the past century (figure 1), with minimum (nocturnal) temperature (tmin) having increased more steeply than maximum (daytime) temperatures (tmax) (houghton et al. 1996). accordingly, in this study we explored to what extent climate–vegetation responses could plausibly explain diverse and divergent spatiotemporal patterns in population performance of eight large herbivore species with divergent forage requirements in a semi-arid savanna system. preliminary assessments had revealed large herbivore species groups with similar within-group population trends over time. these appeared to be matched by commonalities of habitat preferences (with regard to altitude, underlying geology, northern compared with southern knp) and features of nutritional ecology [seydack et al. 2012; for roughage bulk feeders such as buffalo and waterbuck, see hofmann (1973, 1989); for short-grass preference grazers such as blue wildebeest and zebra, see bodenstein, meissner and van hoven (2000) and grange and duncan (2006); for selectively feeding antelope species such as sable, roan, tsessebe and eland, see skinner and chimimba (2005)]. the study involved three components: • establishing correlations (patterns) between herbivore population performance trends and climate features (rainfall, temperature) • application of the climate–vegetation response model (seydack et al. 2012), which defined the expected effects of climate on key indices of forage availability for herbivores (process) towards the interpretation of herbivore population performance patterns in space and over time • exploring the degree of consistency with which climate-controlled forage quantity and quality can explain correlations between climate and herbivore population trends. figure 1: southern hemisphere surface air temperature anomalies between 1850 and 2009. through such pattern/process matching the output of this study is intended to represent an explanatory framework that may effectively serve to direct further research in pursuit of rigorous verification of inferred causal linkages. the results of this study were interpreted and discussed in the context of making a contribution towards the ongoing endeavour to disentangle the relative effects of bottom-up control (nutrition) compared with top-down regulation (predation) of large herbivore abundance in african savannas. in its broadest sense, the objective of this study was to contribute towards the enhanced understanding of landscape-scale functioning of savanna systems with regard to the interplay between climate, vegetation and herbivore population dynamics. study area the kruger national park (knp) is situated in north-eastern south africa and represents a large (approximately 2 000 000 ha) semi-arid savanna system, covering an altitudinal range of between 200 m a.s.l. and 700 m a.s.l. the knp south of the olifants river falls within the lowveld bushveld climatic zone (500 mm – 700 mm rain per year), whilst northern areas fall within the northern arid bushveld zone (300 mm – 500 mm rain per year) (venter, scholes & eckhardt 2003). rainfall is concentrated in the austral summer, between october and april. high temperatures are experienced (mean annual temperature = 21.9 °c), with winters generally being frost free (venter et al. 2003).the park is longitudinally subdivided into granitic substrates forming relatively nutrient-poor sandy soils in the west and more nutrient-rich basalt-derived clay soils in the east (venter et al. 2003). the savanna vegetation on nutrient-poor substrates tends to be dominated by trees of combretaceae (e.g. combretum and terminalia species) and caesalpineaceae, whereas those on more nutrient-rich clay soils are dominated by trees of mimosaceae – especially acacia species (gertenbach 1983; venter et al. 2003). the vegetation of the northern knp is characteristically dominated by mopane (colophospermum mopane) in tree or shrub form (gertenbach 1983; venter et al. 2003). the herbaceous layer of the knp is dominated by c4 grass species (kennedy, biggs & zambatis 2003) and the more nutrient-rich savanna vegetation types on clay soils carry dense stands of nutritious, high-bulk grasses (venter et al. 2003). materials and methods top ↑ pattern/process matching the meaningful assessment of climate effects on long-term herbivore population performance requires suitable data spanning time periods containing sufficient replication of population abundance to climate variability. the nature, quality and comparability of such data are generally variable, ranging from mere population performance trend ratings to reasonably accurate census estimates. accordingly, there is limited scope for sophisticated statistical analyses that directly link herbivore population numbers and climate. in this study we explored correlations between herbivore population performance trends and climate through pattern/process matching; that is, through establishing to what extent climate–herbivore population trend correlations (pattern) were consistent with expected climate effects on forage availability over time and space (process). the expected climate effects were made explicit through the development of a climate–vegetation response model, where existing knowledge, inter alia through literature review and local research, was interpreted and synthesised to produce a framework that outlined how vegetation response to climate was expected to shape forage quantity and quality (seydack et al. 2012). in accordance with this approach, this study involved three main components. firstly, spatiotemporal patterns of herbivore population performance trends were established and associations with climate variables assessed (pattern). the impact of vegetation responses to climate on indicators of forage availability parameters and species-specific requirements of the herbivores in respect of forage quantity and quality were then defined through the climate–vegetation response model (process). we then inferred causality of climate–herbivore population patterns from the consistency with which parameters of forage availability, as shaped by vegetation response to climate, could explain correlations between climate and herbivore population trends. patterns in herbivore population performance population performance trends the eight large herbivore species selected for this study were chosen to represent contrasting demographic patterns and feeding styles. bulk-grazing species included the african buffalo, blue wildebeest and plains zebra. less common than the three principal bulk-feeding species, the waterbuck, was selected to represent a habitat specialist with a contrasting feeding style to the selective grazers. the four antelope species grouped as selectively feeding species were sable antelope, roan antelope, tsessebe and eland. archival annual reports were perused for population trend observations and estimates [ludorf 1918; archives (skukuza) 1946–1952; steyn 1953–1961; archives (skukuza) 1902–1945]. population estimates for the period 1961–1976 were furthermore retrieved from scientific services records (see joubert 1976; joubert & bronkhorst 1977; pienaar 1963; whyte & joubert 1988). population counts conducted by fixed-wing aircraft of most of the larger ungulates in the knp were undertaken irregularly between 1965 and 1975 and annually from 1977 to 1997. the full extent of the knp was covered only between 1980 and 1993, with total aerial coverage being pursued every year between may and august using four observers. parallel strips of about 800 m wide (400 m on each side) were flown at 65 m – 70 m above ground level (viljoen 1996). since 1998 aerial census counts were undertaken at relatively low sampling intensities (kruger, reilly & whyte 2008), using the distance sampling method (buckland et al. 1993). after 1993 population estimates of the four rare antelope species for the years 2005/2006 were extracted from whyte (2007). african buffalo counts had been conducted separately by using a helicopter to split herds into smaller groups and then photographing them. animals on the photographs were then counted with visual aids. spatiotemporal patterns to gain greater understanding of factors that controlled population performance of the species studied, an analysis of habitat preferences in respect of altitude and soil nutrient status was carried out. between one and three burn blocks in the park were combined to form 123 compartments or spatial units for analyses (northern knp: 1–65; southern knp: 66–123). each unit was created to encompass only a restricted range in altitude and was relatively homogeneous with regard to underlying geology. the average size of units was 154 km2, generally ranging between 70 km2 and 370 km2. for each unit, representative variable means were determined in respects of kernel population densities of the eight study species in 1977, 1980, 1986, 1990 and 1993 (for buffalo: 1986, 1990, 1993, 1999 and 2004), long-term rainfall, altitude and soil nutrient status.animal density distribution patterns were smoothed by converting the point-based census data to a 0.5-km grid using kernel densities with a 5-km search radius. the kernel function was based on the quadratic kernel function described in silverman (1986). mean altitudes for units of analysis were derived from a digital elevation model (environmental systems research institute 2006). long-term rainfall was based on spline interpolation of long-term average rainfall of 23 ranger posts distributed across the park. mean nutrient status ratings for spatial units of analysis were computed based on geology (venter et al. 2003) and vegetation indicators of nutrient availability according to the landscape vegetation units of gertenbach (1983). for the same geological substrate soil nutrient status was further differentiated according to the predominance of the following tree species in order of decreasing nutrient availability indication: fine-leaved acacia species > colophospermum mopane (clay soil) > combretum species (scholes & walker 1993; venter et al. 2003). weighted means according to proportional spatial representation of soil nutrient availability ratings were calculated for each unit of analysis. the following soil nutrient availability ratings were applied: • 1: sandveld communities • 2: granites with prominence of combretum species • 3: granites with prominence of colophospermum mopane; vegetation associated with lebombo north and south • 4: thickets of sabi and crocodile rivers • 5: vegetation communities on gabbro, shales and amphibolites • 6: basalts with dominance of colophospernum mopane • 7: basalts with prominence of acacia species. multiple regression analyses of population densities on altitude and soil nutrient status were performed for the northern and southern knp separately, corrected for spatial autocorrelation as required. three types of regression analysis were applied, namely ordinary least squares, spatial lag model and spatial error model (anselin 2005). the spatial neighbourhood had to be defined for each of the two spatial regression models: neighbouring polygons of all those sharing sides (rook contingency) or neighbouring polygons sharing a side or only a vertex (queen contingency). the spatial regression decision process based on the lagrange multiplier test statistic was used to identify which of the models produced the most reliable results (anselin 2005). whenever one of the spatial models was selected, owing to the presence of spatial autocorrelation, akaike information criteria were used to identify whether the rook or queen contingency models gave the best result. climate and herbivore population performance two types of analysis were conducted to explore any associations between climate variables and herbivore population performance. one was based on the population trends as reconstructed from historical records and extant census data over the period 1944–2003 and the other involved repeated measures analysis of variance (anova) of actual census data from 1977 to 2005/2008, differentiated according to north–south subregions and altitude.for the analysis of association of climate and herbivore population performance between 1944 and 2003 we reconstructed the temporal population trends for the groups of wildebeest/zebra, selective grazer species, buffalo and waterbuck from historical observations or population estimates (1944–1976) and census data (1977–2003). the temporal population performance trends were interpreted according to the following four categories, with assigned rank variables given in parentheses: decreasing (1); remaining low (2); high or sustained at relatively high levels (3); increasing (4). spearman rank correlation analyses were performed between these rank variables and tmax, tmin, annual rainfall and mean monthly dry-season rainfall (april–september). variable means for groups of two consecutive years were used, resulting in n = 30 for the period of analysis spanning 60 years. analyses were differentiated according to northern and southern knp in respect of the blue wildebeest and zebra group and performed in respect of three time lag periods (zero-, twoand four-year time lags). since the spearman rank analyses implicated that temporal population trends were linked to climate, we further performed a series of repeated measures anovas to explore space–time (climate) interactions. for this purpose we defined strata formed by grouping neighbouring units of analyses (compartments as described above). these strata, defined with regard to subregions on a north-to-south gradient and altitude groupings, thus contained replicates represented by neighbouring units of sampling (legendre & mcardle 1997). partly depending on data availability, three separate repeated measures anovas were performed in respect of population density estimates determined for the relevant compartments and species or species groups from aerial survey census data: • for blue wildebeest and zebra, data were used from the subregions north (150 m a.s.l. – 550 m a.s.l.), central lower (150 m – 350 m), central upper (350 m – 550 m), southern lower (150 m – 350 m) and southern upper (350 m – 650 m), for the years 1980, 1986, 1990, 1993, 2002 and 2008. • further analyses of population density data from the northern knp for the four selective grazer species, blue wildebeest and zebra involved three altitudinal groupings (low, middle and upper altitude) and the years 1977, 1980, 1986, 1990, 1993 and 2005. because granites and basalts stretched over divergent altitudinal ranges and the distributions of certain species were concentrated on specific substrates, analyses specific for species–substrate combinations were performed: sable antelope on granites, roan and tsessebe on basalts, and blue wildebeest, zebra and eland over both substrates combined (altitudinal ranges for granites or basalts combined: low: 150 m a.s.l. – 300 m a.s.l.; mid-altitude: 300 m – 400 m; upper: 400 m – 550 m). • of the selective grazers only sable antelope occurred at sufficient population densities in both the northern and the southern knp. analyses were performed for the years 1980, 1986, 1990, 1993 and 2005, and the following spatial units: upper, middle and lower altitudes in the northern knp, upper and lower altitudes in the central knp, and upper south-facing as well as north-facing, middle and lower altitudes in the southern knp. subregions along the north-to-south gradient represented areas of increasing rainfall (venter et al. 2003) and altitudinal groupings have implications for the prevailing temperature regimes according to lapse rates of –0.777 °c.100 m-1 (tmax) and –0.465 °c.100 m-1 (tmin), respectively (schulze 1995). given the extent of global warming over the past century (figure 1), these lapse rates imply the equivalent to a lowering of the average altitude of the knp of between 100 m and 200 m. the analyses described in this section were aimed at revealing patterns in population performance of the relevant herbivore species in respect of climate variables in space (north–south gradient: rainfall; altitude: temperature) and over time (climate data: south african weather service), as well as spatiotemporal interactions. climate-vegetation response model (process) development of an explanatory framework exploration of spatiotemporal patterns in population performance of eight large herbivore species had revealed three population performance response groups (i.e. groups of large herbivore species which shared broad-scale habitat preferences and similar population performance trends over time) and group-specific correlations with climate variables. a climate–vegetation response model had been developed representing a framework for the interpretation of these spatiotemporal patterns of herbivore population performance in relation to climate effects (seydack et al. 2012). as part of the climate–vegetation response model, we (1) defined indicators of forage availability (quantity and quality), (2) established forage selection requirements characteristic of and unique to the three herbivore population performance response groups (herbivore species guilds of nutritional requirements), and (3) developed a plant–climate response process model. the resulting climate–vegetation response model defined the expected effects of climate on indices of forage availability (process effects), allowing us to make inferences that linked spatiotemporal patterns in herbivore population performance and climate (seydack et al. 2012). key forage resource indicators the climate–vegetation response model differentiates between three key indicators affecting forage availability to herbivores, namely nitrogen productivity, nitrogen quality and carbon-nutrient quality. nitrogen productivity refers to the quantitative availability of forage items for herbivores of a given nitrogen quality (plant protein content) as a result of biomass production. high nitrogen quality of forage implies high nitrogen concentrations of plant tissues; that is, plant protein content not unduly diluted by structural or non-structural carbon and of relatively high digestibility. carbon-nutrient quality forage items are characterised by relatively high concentrations of non-structural carbohydrates (tnc) and ash minerals resulting from metabolic allocation to storage under circumstances promoted by constrained growth (low temperatures, moisture stress: tw conditions), but sustained photosynthesis. at the plant species level, andropogonoid grass species are physiologically predisposed to cope with pronounced seasonality (osborne 2008). the associated physiological capacity of species of andropogoneae for sustained metabolic performance at variable and lower resource levels (dry-season growth) underpins their metabolic scope for storage of metabolites (tnc and minerals). andropogonoid grass species accordingly tend to produce forage of relatively high carbon-nutrient quality (seydack et al. 2012). herbivore species guilds of nutritional requirements as inferred from the nature of the diets of eight large savanna herbivore species (bodenstein et al. 2000; grange & duncan 2006; heitkönig & owen-smith 1998; macandza, owen-smith & cross 2004; magome et al. 2008; parrini 2006; watson & owen-smith 2000), three herbivore species guilds of nutritional requirements were identified (seydack et al. 2012): bulk feeders with tolerance to fibrous herbage (buffalo and waterbuck), bulk feeders with preference for high nitrogen-quality forage (short-grass preference grazers: blue wildebeest and zebra) and selective feeders for which dietary items of relatively high carbon-nutrient quality, notably in the dry season, represent key forage resources (selective grazers: sable antelope, roan antelope, tsessebe and eland). plant metabolic performance trade-off constraints the principal trade-off in the context of the metabolic performance trade-off model (seydack et al. 2012) is between maximised peak performance closely tracking and responsive to conditions of resource surplus (rmp: resource-responsive metabolic performance mode) and sustained performance when subject to conditions of varying resource levels or deficits (smp: sustained metabolic performance mode). the capacity for maximum metabolic performance under conditions of resource surplus is at the cost of the capacity for sustained performance under conditions of resource deficits or fluctuations. in line with this reasoning, we expect a propensity towards the rmp mode under conditions of high and less fluctuating levels of temperature and water availability (tw conditions: combined relatively high and sustained levels of water and temperature during the growing season), whereas under conditions of fluctuating resource levels and pronounced deficits (with reference to temperature and water: tw or tw conditions), the smp mode would be expected to predominate. the relative expression of smp to rmp mode functionality or acclimation is thus a function of the combined availability levels of temperature and water (tw resource levels or conditions). the higher the concurrent and uninterrupted levels of temperature and water available to plants, the more pronounced the intraspecific metabolic adjustment towards higher rmp mode functionality settings. depending on the spatiotemporal nature of resource availability for plants (water and temperature resource levels and variability or seasonality), the relative expression of sustained to resource responsive metabolic performance is determined, shaping forage quantity and quality as a consequence of the relative prevalence of smp-tw/tw to rmp-tw plant functionality.plants in relative smp mode are comparatively stress tolerant in order to cope with resource level fluctuations. as such their metabolism is capable of functioning at relatively low temperature and water availability/optima (smp-tw functional efficiency). such metabolic capacity is, however, associated with relatively high storage-to-growth allocation priority and comparatively high maintenance respiration costs (i.e. they are temperature sensitive). in contrast, plants in relative rmp mode are geared to the maximum use of surplus resource conditions, with functionality at relatively high temperature and water resource level optima and high growth-to-storage priority (seydack et al. 2012). thus, abundance of resources such as water, nutrients and temperature enhances rmp mode productivity, but increasing productivity (increased rmp-tw functionality) progressively reduces plant tissue quality for herbivores through nutrient dilution (an et al. 2005; dwyer et al. 2007; ’t mannetje 1982, 1984; mutanga et al. 2004; van soest, mertens & deinum 1978; wilson 1982; wilson & ford 1971), reduction in digestibility (’t mannetje 1984; van soest et al. 1978; wilson 1982; wilson & ford 1973) and reduced allocation to storage (low tnc content; estiarte & peñuelas 1999; herms & mattson 1992; van soest et al. 1978). three main groups of process effects are involved in the climate–vegetation response model (cvrm): adjustments in plant metabolic performance (smp-rmp functionality) settings shaping the spatial distribution of forage resources (cvrm-1), smp-to-rmp temperature acclimation responses (cvrm-2), and spatiotemporal interactions in these effects (cvrm-3). spatial distribution of key forage resources (cvrm-1) following the climate–vegetation response model, plant metabolic adjustments to the north–south water availability and altitudinal temperature gradients are expected to determine the spatial distribution of smp-to-rmp functionality settings across the knp (seydack et al. 2012). the general pattern is one of higher smp mode functional expression in the drier, northern knp and at higher altitudes, and advanced rmp functionality settings towards the south. as the smp mode involves relatively growth-curbed, storage-based metabolism, the availability of forage of enhanced carbon-nutrient quality is accordingly expected to be relatively higher in the northern knp and at upper altitudes. plant metabolic performance at intermediate smp–rmp functionality favours lower levels of nitrogen dilution and is associated with forage of relatively high nitrogen quality, especially of plants growing on nutrient-rich geological substrates (central knp). climate-driven temporal trends in forage availability (cvrm-2) increasing tmin is expected to stimulate nocturnal maintenance respiration (wan et al. 2009), promoting temperature acclimation in order to restore the original steady-state respiration-to-photosynthesis ratio (dewar, medlyn & mcmurtrie 1999). as temperature increases, leaf metabolic pools adjust until the original steady-state respiration-to-photosynthesis ratio is re-established. leaf labile carbon, starch and protein pools decrease in response to increased temperature, eventually reaching a new, lower steady state, which is consistent with the general pattern of plant carbohydrate and soluble protein content being negatively correlated with growth temperature (dewar et al. 1999; wan et al. 2009). these acclimation responses represent intraspecific smp to more rmp functionality shifts (smp-to-rmp temperature acclimation). according to the climate–vegetation response model both shorter-term transient temperature acclimation pulses (figure 1: increasing temperature towards 1940; increasing towards 1960; increasing after 1975) and longer-term shifts towards advanced rmp functionality settings are predicted to have occurred in response to temperature trends over the past century (figure 1; seydack et al. 2012). • cvrm-2.1: pulses of increased forage availability due to increased productivity of quality forage induced by pseudo-cyclic phases of temperature increases (figure 1: increasing towards 1940; increasing towards 1960; increasing post-1975). in response to rising tmax, transient pulses of smp productivity (smp-p temperature acclimation phase state) occur when the temperature optima for plant functionality are still relatively low (tw metabolic activity). however, as acclimation to tmin takes effect, increased rmp-tw metabolic activity of plants results in reductions in forage quality (low metabolic allocation to storage and increased nitrogen dilution) and thus forage availability. the complete smp–rmp–smp temperature acclimation sequence involves four temperature acclimation phase states: 1: smp-t (smp functionality at relatively low temperatures; np ↓, nq ↓, cnq ↑↑); 2: smp-p (enhanced smp productivity due to rising tmax; np ↑, nq ↑, cnq ↑, smp-tw efficiency ↑); 3: smp-t (smp functionality at relatively higher/increased temperatures; np ↑, nq ↑, cnq ↓); 4: rmp-t (rmp functionality at high temperatures; np ↑, nq ↓, cnq ↓↓, smp-tw efficiency ↓), reverting to phase states 3: smp-t (lower altitudes) or 1: smp-t (upper altitudes) as ambient temperatures decrease again.1 • cvrm-2.2: long-term progressive shifts towards relatively advanced rmp functionality settings in response to cumulative acclimation to increasing tmin and declining diurnal temperature ranges. trends of progressively deteriorating carbon-nutrient quality and increasing nitrogen productivity to nitrogen quality ratios (np/nq ↑) over the past century were accordingly anticipated (figure 1). these temperature acclimation responses were expected to have resulted in transient pulses of increased forage availability (increased landscape-scale availability of forage items of favourable nitrogen and carbon-nutrient quality), as well as the progressive long-term decline of carbon-nutrient quality of forage and an increasing nitrogen productivity/nitrogen quality ratio associated with advanced rmp functionality settings (seydack et al. 2012). spatiotemporal interaction in forage availability (cvrm-3) accentuated temperature acclimation responses were predicted for circumstances where low and unpredictable water availability favoured the prevalence of smp mode settings; that is, in northern relative to southern knp. more pronounced associated effects on nitrogen quality and carbon-nutrient quality are thus expected towards the northern knp, with attenuation or absence of such responses as relatively high rmp functionality settings already predominate towards the southern knp. as plants are already functioning at higher rmp settings in the southern knp, smp–rmp–smp temperature response pulses (smp–rmp temperature acclimation sequences) are expected to be attenuated relative to those of the northern knp, where pronounced associated pulses of forage availability are expected to be encountered (seydack et al. 2012). results top ↑ patterns in long-term population performance trends historical population trends pertinent features of historical trends (1900–1970) in population performance regarding the eight species under study can be summarised as follows:• from a situation prevailing before about 1920–1930, where population abundance of selective grazers and bulk feeders was of similar orders of magnitude, the former generally declined progressively over subsequent decades, whereas the latter had generally increased over this time span (figure 2: following ludorf 1918; pienaar 1963). • a pronounced declining phase occurred for 10 or more years preceding the mid-1950s; involving, particularly, sable antelope, roan antelope, tsessebe, wildebeest and eland. the only grazers not to show this trend were buffalo. population decreases were more severe in the southern and central knp (steyn 1953–1961; steyn 1954). after this decline, only sable antelope sustained reasonable population numbers south of the olifants river (figure 2). main source habitat centres and prospering populations of selectively feeding antelope species were consistently recorded in respect of higher-lying areas in the western boundary zone and further west of the park [joubert 1976; archives (skukuza) 1902–1945]. • zebra population numbers surpassed those of wildebeest during 1955–1970 and stabilised at higher levels than the latter, especially in the northern knp (joubert 2007a; steyn 1954; figure 2). figure 2: population estimates for six of the eight species studied to show long–term population trends for (a) selective grazers and (b) bulk feeders in the northern and the southern kruger national park. extant population trends blue wildebeest and zebra had largely similar population performance trends over the time span, as depicted in figures 3a and 3b. clear divergence in performance trends between northern and southern populations was evident. in the south, population numbers of both species increased (1978–1992) and then seemed to have stabilised, fluctuating around some asymptotic carrying capacity level. the seemingly high year-to-year fluctuations since 1998 were taken to be largely the result of sampling inefficiencies (kruger et al. 2008), partly associated with relatively low sampling intensities of the censuses during that time period (figures 3a and 3b). after exhibiting parallel population increases and stabilisation as for southern populations, northern populations of both species were subject to prolonged decreases in their population numbers. it is also evident that wildebeest population performance in the north had been substantially poorer than south of the olifants river (figure 3a). population trends of buffalo and waterbuck were broadly similar and showed no fundamental north–south divergence (figures 3c and 3d). population numbers of both species declined substantially subsequent to the 1991/1992 drought and increased in the years thereafter (figures 3c and 3d). figure 3: population numbers of (a) blue wildebeest, (b) zebra, (c) buffalo and (d) waterbuck based on censuses carried out between 1978 and 2006 (northern and southern kruger national park). relatively superior demographic performance was clearly evident in northern populations of selectively feeding antelope species, especially regarding roan antelope, tsessebe and eland (figures 4a–d), contrasting the performance of populations in the south, which had been reduced to low numbers earlier in the first half of the previous century (historical population trends). sable antelope, tsessebe and eland had pronounced population increases in the northern knp, followed by steep population declines for all four selectively feeding antelope species manifesting around 1986/1987. southern populations of sable antelope, roan and tsessebe, already at relatively low levels, further decreased to exceptionally low numbers between 1990 and 1994 (figures 4a–c). overall, population performance trends amongst the four selectively feeding antelope species were broadly similar (figures 4a–d). these trends differed from those of the four bulk-feeding species in that the population performance of selectively feeding antelope species progressively deteriorated after 1986 without significant recovery. in contrast, buffalo and waterbuck numbers recovered in the years after the 1991/1992 drought (figures 3c and 3d). in conclusion, it transpired that the eight species under study could be assigned to three groups of broadly similar population performance trends: buffalo/waterbuck (figures 3c and 3d), blue wildebeest/zebra (figures 3a and 3b), and the four selectively feeding antelope species (figures 4a-d). figure 4: population numbers of (a) sable antelope, (b) roan antelope, (c) tsessebe and (d) eland based on censuses carried out between 1954 and 2006 (northern and southern kruger national park). landscape-scale spatial patterns in population performance three species groups of similar spatiotemporal population preferences were confirmed. selectively feeding antelope species characteristically preferred upper altitude habitat (table 1 and figure 5) and northern populations seemed to perform better than southern populations (figure 4). roan antelope and tsessebe were found in higher-lying parts of their preferred habitat of higher nutrient status (table 1), as is evidenced by higher population concentrations on basalts in north-eastern parts of the park (figures 5c–d). bulk-feeding species did not exhibit preferences for upper altitude habitat and population performance was either higher in the southern (central) knp (blue wildebeest and zebra: figures 3a–b) or indifferent in this respect (buffalo and waterbuck: figures 3c–d). blue wildebeest and zebra, as expected for high nitrogen quality short-grass grazers, preferred nutrient-rich areas (table 1), in contrast to buffalo and waterbuck (table 1) with their capacity to tolerate forage of lower quality (hofmann 1973, 1989). the species of these emerging population performance groups exhibited within-group congruence regarding their spatiotemporal habitat preferences and also of their population trends over time, resulting in three consistent population performance response groups (buffalo/waterbuck; blue wildebeest/zebra, and the four selectively feeding antelope species). figure 5: distribution of census observation localities in relation to altitude for sable antelope (black dots) and eland (white dots) during (a) 1980/1986 combined and (b) 2005, and for roan antelope (black dots) and tsessebe (white dots) during (c) 1980/1986 combined and (d) 2005. table 1: multiple regression analyses of population densities on habitat variables of nutrient status and altitude (northern kruger national park). the spatial distribution of buffalo and waterbuck was not consistently related to the landscape-scale habitat niche factors explored (altitude, north–south plant smp-to-rmp metabolic performance mode gradient, geological substrate), presumably because of the less exacting forage quality requirements of these two species. comparatively stable population performance of blue wildebeest and zebra was encountered in the southern (central) knp (figures 3a–b), which coincides with the expected landscape-scale prevalence of high nitrogen quality forage items (cvrm-1: intermediate rmp-smp-tw functionality on nutrient-rich sites, associated with reduced incidence of nitrogen dilution of forage items with structural and non-structural carbon). the preference of selective grazers for upper altitude or northern knp sites was congruent with smp-tw functionality of plants, which is conducive to relatively high carbon-nutrient quality of forage items (cvrm-1). these results confirm that populations of the species belonging to the identified population performance response groups are spatially distributed in the landscape as expected and in accordance with group-specific nutritional requirements. in the next section, we explore whether group-specific herbivore population performance trends were consistent with expectation of climate effects on the relevant key indices of forage availability (nitrogen productivity, nitrogen quality, carbon-nutrient quality), thereby linking landscape-scale temporal patterns of herbivore population performance to climate according to the climate–vegetation response model. climate and herbivore population performance buffalo and waterbuck population trends of buffalo (figure 3c) and waterbuck (figure 3d) broadly followed inter-annually declining and increasing rainfall trends from 1981 to 1992 and 1993 to 2006, respectively. a positive association of population performance trend ratings of these species with periodic rainfall confirmed this (table 2), as also found by mills et al. (1995) and owen-smith and ogutu (2003). because buffalo and waterbuck do not have high forage quality requirements (hofmann 1973, 1989), population densities appeared to be tracking forage quantity largely determined by rainfall. both these species are water dependent and their need to remain close to or have sufficient access to significant open water sources may further have constrained forage availability during drought years (smit 2011). table 2: spearman rank correlations of population performance trend ratings with climate variables for blue wildebeest/zebra, selectively feeding antelope species groups (figure 6), buffalo and waterbuck. blue wildebeest and zebra the long-term population trends of wildebeest and zebra combined, as reconstructed from historical records and extant census data (figures 3a–b), are shown in figure 6 (line a) against climate variables. population performance trend ratings over the relevant time span for these two species were found to be negatively correlated with tmin (four-year lag) and positively with dry-season rainfall (two-year lag; table 2). wildebeest and zebra population increases had occurred (figure 6, line a: c.1955–1963, 1977–1986) following time periods of below-average tmax (figure 6: broken line a) and associated phases of temperature increases (figure 7a: 1953–1959, mid-1970s onwards). these periods of population increases also coincided with phases of sustained, relatively high diurnal temperature ranges (figure 7a), whereas years of population decline (c.1944–1953, 1967–1974, 1991–2002) were partially preceded and associated with elevated tmin [figure 6: line b (middle panel)] and phases of reduced diurnal temperature ranges [figure 6: shaded rectangles, line g (bottom panel)]. such conditions were expected to initiate the smp-rmp temperature acclimation sequence in accordance with the climate–vegetation response model (cvrm-2.1). with an increased availability of forage with high nitrogen quality as key resource, blue wildebeest and zebra populations initially increased during the smp-p phase state and then declined as temperature acclimation progressed into the rmp-t phase state and nitrogen quality of forage was compromised (cvrm-2.1). three such temperature acclimation sequences and associated population cycles (i.e. increasing and then declining populations) seem to have occurred during the past century (approximately bounded by periods 1910–1955, 1956–1975 and 1976–2002; see figures 6 and 7). the first of the three sequences would have been predominated by the prolonged initial occurrence of the smp-t phase state (cvrm-2.1) owing to comparatively low atmospheric temperatures (figure 1). according to the climate–vegetation response model, the associated conditions of reduced nitrogen productivity and relatively low nitrogen quality of available forage items (due to nitrogen dilution by non-structural carbohydrates) were expected to be unfavourable for the population performance of wildebeest and zebra. their comparatively low population numbers at the time (1918: figure 2) are consistent with expectations in this regard. figure 6: long-term population trends reconstructed from historical records and extant census data for (a) blue wildebeest and zebra in southern kruger national park (bold arrows) and trend deviation for northern populations (broken line arrow), and (b) selectively feeding antelope species: sable, roan, tsessebe and eland (bold arrows; thinner arrows for sable and eland only). figure 7: annual means of mean monthly minimum temperatures and maximum-minimum temperature differences (tmax–tmin = diurnal temperature range) at (a) messina, northwest of the kruger national park and (b) pretoriuskop (southern knp at upper altitudes; south african weather service). spatiotemporal interactions in population performance along a north–south gradient were detected (figure 8; time period: f = 12.87, df = 5, p < 0.000001; time period × species: f = 0.49, df = 5, p = 0.78; time period × subregions: f = 6.42, df = 20, p < 0.000001). in the northern knp the population cycle was accentuated in that relatively high population peaks were achieved (1986), with population numbers declining to particularly low numbers later (2002/2008). in the central lower knp population densities stabilised at pre-increase levels (1980) but in southern lower knp no increase or decline cycle was evident (figure 8). also, the population cycle in southern upper knp was attenuated in comparison with that seen in central upper knp. these patterns are consistent with predictions of the climate–vegetation response model. smp-to-rmp acclimation sequences (cvrm-2.1) underpinning the associated herbivore population cycles are expected to be accentuated where smp mode settings are most pronounced (northern knp) and much attenuated or absent with relative prevalence of rmp mode settings of the vegetation southwards (cvrm-3: reduced temperature acclimation response in plants with prevalence of rmp mode settings). figure 8: population densities of blue wildebeest and zebra (1980–2008: kruger national park), differentiated according to subregions of divergent metabolic mode settings (cvrm-1). in the knp overall, blue wildebeest numbers were higher than those of zebra during the first of the three reconstructed population cycles (steyn 1954) and also at peak numbers of the second cycle (1963: figure 2b). thereafter, zebra population numbers remained higher than those of blue wildebeest (figures 2 and 3a–b). this is in agreement with the greater tolerance of zebra to declining nitrogen quality of suitable forage (bodenstein et al. 2000; smuts 1972). with generally rising atmospheric temperatures since about 1910 (figure 1) nitrogen productivity of especially c4 grasses would have increased, but at the cost of nitrogen quality (climate–vegetation response model). the ratio of nitrogen productivity to nitrogen quality thus increased progressively (cvrm-2.2), resulting in more and spatially extended availability of forage with comparatively reduced nitrogen quality. this disproportionately favoured the increase of zebra relative to blue wildebeest abundance. selectively feeding antelope species elevated tmin and reduced levels of dry-season rainfall were found to be associated with poor population performance of selectively feeding antelope species [table 2; figure 6: line b (top panel), with reference to lines b and c (middle panel)]. the negative impact manifested in reduced population densities with some time lag (table 2). reconstructed population trends for the four selective grazer species manifesting in the northern knp were crudely synchronous with those of blue wildebeest and zebra, exhibiting three cycles of increasing/declining population numbers over the past century (figure 6: line b). these periods of peak abundance followed phases of increasing temperature and reduced diurnal temperature ranges (dtrs: tmax– tmin; figure 7a). the population cycles were thus in temporal concordance with pulses of temperature increases (figure 7a), which were expected to initiate smp-rmp temperature acclimation sequences (cvrm-2.1). during the initial stage of increasing temperature, tmax is comparatively high relative to tmin (increased dtr: figure 7a). the associated smp-p acclimation phase state (enhanced smp mode productivity) supports increasing nitrogen productivity of forage of sustained nitrogen quality, and notably of increased carbon-nutrient quality of dry-season forage (cvrm-2.1), thereby forming the basis for increases of selective grazer and blue wildebeest/zebra population numbers alike (figure 6; 1978–1986: figures 4a-d). as temperature acclimation to nocturnal warming (rising tmin) takes effect, cn-quality of forages declines during the smp-t phase state (cvrm-2.1), resulting in population declines of selective grazers (figure 6, line b: after mid 1940s, mid 1960s and 1986; figures 9a-d). this phase state is, however, still favourable for nitrogen quality forage (cvrm-2.1), the key resource for blue wildebeest and zebra, and thus their populations continued to prosper, expanding into upper-altitude habitat (figures 9e–f, 1986–1990/1993). as temperature acclimation progressed to the final rmp-t phase state, both nitrogen quality and carbon-nutrient quality of forage were substantially compromised, resulting in declining populations also of blue wildebeest and zebra (figures 9e–f). forage of higher carbon-nutrient quality, a key resource for selective grazers, is expected under conditions of lower temperature, that is, higher altitudes (cvrm-1), which explains the preference of selective grazers species for upper-altitude habitat (table 1, figures 5, 9a–d and 10). population declines of these species were, accordingly, also less pronounced at higher altitudes (figures 9a–d; time period × altitude class interactions: p = 0.15; 0.0002; 0.03; 0.62; values indicate that population performance over time is not independent of altitude in the case of sable antelope, tsessebe and roan antelope). figure 9: population densities of four selective grazers, (a) sable antelope, (b) tsessebe, (c) roan antelope, (d) eland and two large herbivore species selecting for nitrogen–quality patches, (e) blue wildebeest and (f) zebra in the kruger national park (1977–2005 or 2008) differentiated over three altitude classes. figure 10: population densities of sable antelope in subregions of the kruger national park (1980–2005), differentiated into broad classes of altitude. of the four selectively feeding antelope species, only sable antelope occurred in substantial numbers in the southern knp during the time period representing the third population cycle. whereas northern population abundance of these species exhibited a pronounced population cycle between 1978 and 1994 (figures 4a–d), a similar pattern in sable antelope numbers in the southern knp was comparatively attenuated and somewhat delayed (figure 4a; for tsessebe, see figure 4c). the relatively attenuated nature of population cycles in the southern knp (cvrm-3), as also exhibited by blue wildebeest and zebra populations, is attributed to a reduced smp-to-rmp temperature acclimation response owing to the relative prevalence of rmp mode settings (already) applicable to the area (cvrm-1). sable antelope population densities in the southern knp at upper altitudes (pretoriuskop area) (figure 10: upper south-facing and north-facing; 450 m a.s.l. – 650 m a.s.l.) were apparently sustained at levels similar to those historically encountered over larger areas towards lower altitudes [figure 2a: 1918; archives (skukuza) 1902–1945]. the comparatively favourable population performance of sable antelope populations in the pretoriuskop area is attributed to conditions supporting carbon-nutrient quality of forage items: comparatively high altitude, increased incidence of dry-season rainfall (mean monthly dry-season rainfall between 1982 and 2005: letaba = 59 mm, satara = 70 mm, skukuza = 91 mm, pretoriuskop = 104 mm), reduced smp-to-rmp temperature acclimation response and the occurrence of andropogonoid grass species (cf. le roux 2010: heteropogon contortus/hyperthelia dissoluta). however, population densities progressively declined after 1990 (figure 10), apparently more so in predominantly north-facing areas. temperature conditions had been deteriorating progressively towards and beyond 1990. increasing tmin and decreasing diurnal temperature ranges (figure 7b) and further smp-to-rmp temperature acclimation (more pronounced rmt-t phase states) would have particularly compromised carbon-nutrient quality of forage plants. accordingly, the associated observed decline in sable antelope population numbers was consistent with predictions of the climate–vegetation response model.the crudely synchronous population cycles of blue wildebeest and zebra and the selective grazers (figure 6), but long-term inverse population trends of these two groups (figures 2a–b) represented a seemingly paradox result. however, the long-term increase of blue wildebeest and zebra, as nitrogen quality patch selectors, and the concurrent decline of antelope species selecting for carbon-nutrient quality plant parts (figure 2a–b) were consistent with the predictions of the climate–vegetation response model. according to this model, increasing temperatures over the last eight to nine decades (with pronounced phases of temperature increases during approximately 1920–1940 and after about 1975: figures 1 and 7) would have progressively enhanced nitrogen productivity and resulted in higher availability of nitrogen quality forage in the landscape for blue wildebeest and zebra owing to the predominance of the smp-t phase state. (cvrm-2.1: nitrogen not diluted by high levels of storage-based carbohydrates, which is postulated to have applied before 1940 when the smp-t acclimation phase state was expected to have predominated.) concurrently, however, congruent with predictions of the climate–vegetation response model, acclimation of plants to progressively rising tmin (nocturnal warming) since the 1910s (figure 1) resulted in cumulative shifts of smp-to-rmp functionality settings (cvrm-2.2). the associated metabolic allocation adjustments from storage towards growth priority were associated with reductions in the storage-based carbon-nutrient quality of forage in the landscape. accordingly, the landscape-scale availability of carbon-nutrient quality forage, a key resource for selective grazers, would have declined (ongoing smp-to-rmp temperature acclimation: cvrm-2.2), resulting in the long-term decline of their population numbers in the knp. discussion top ↑ linking climate–vegetation responses with large herbivore population performance initial assessments revealed the existence of three population performance response groups; that is, groups of large herbivore species which shared similar population performance trends over time (figures 3 and 4: buffalo and waterbuck; blue wildebeest and zebra; sable antelope, roan antelope, tsessebe and eland). member species of these groups also shared landscape-scale distribution patterns in relation to subregion on a north-to-south (rainfall) gradient, altitude and nutrient status based on the underlying geology. perusal of relevant studies (herbivore species guilds of key nutritional requirements) further revealed that the species of the same population performance response groups had common key nutritional resource requirements (seydack et al. 2012). this permitted the grouping of the eight species under study into congruent guilds: bulk feeders with tolerance to fibrous herbage (roughage bulk feeders: buffalo and waterbuck), bulk feeders with preference for high nitrogen quality forage (short-grass preference grazers: blue wildebeest and zebra) and selective feeders for which dietary items of high carbon-nutrient quality are of particular importance (selective grazers: sable antelope, roan antelope, tsessebe and eland). accordingly, key nutritional requirements for these three population performance response groups related to the availability of forage quanity, nitrogen quality and carbon-nutrient quality, respectively. a climate–vegetation response model was developed, which defined the expected impact of climate on these key resources (seydack et al. 2012).according to the climate-vegetation response model forage quality is determined by the position of plant functionality on the smp–rmp metabolic performance mode gradient (spatially: metabolic mode settings: cvrm-1; temporally: temperature acclimation sequences: cvrm-2). the smp mode permits plant functionality at relatively low temperature and water availability levels, which supports storage-based nutrient accumulation and growth-curbed metabolic activity, resulting in comparatively high-quality forage. plants in the rmp mode perform under conditions of high water and temperature levels, which results in low-quality forage. with the climate–vegetation response model being central to an explanatory framework, we proceeded to apply an approach of pattern/process matching where we explored the extent of consistency with which climate impacts on key nutritional resources (process) could explain spatiotemporal correlations between climate and herbivore population trends (pattern). whereas buffalo and waterbuck, grouped as roughage bulk feeders, seemed to be tracking forage quantity largely determined by rainfall (table 2; mills et al. 1995; owen-smith & ogutu 2003), temperature and dry-season rainfall were identified as influential in determining the availability of quality forage in the landscape important for selective grazers and those that prefer short grasses. the importance of dry-season rainfall for these species has been identified in previous studies (dunham et al. 2004; owen-smith & ogutu 2003). however, the effectiveness of dry-season rainfall in determining the availability of quality forage in the landscape appears to be interdependent with the prevailing plant metabolic performance mode settings (spatial scale) and temperature acclimation responses (temporal scale), as controlled spatiotemporally by gradients of water availability and temperature effects. according to the climate–vegetation response model, prevalence of the smp mode in the arid northern knp results in pronounced smp-to-rmp temperature acclimation sequences (cvrm-2.1) in response to pulses of temperature increases [figure 7a; broadly synchronous with temperature trends of the southern hemisphere (figure 1)], thereby inducing accentuated population cycles of short-grass preference grazers (figures 3a, b and 6) and selective grazers (figures 4 and 6). such pronounced temperature responses are expected since smp mode plant functionality is sensitive to the impact of nocturnal warming on respiration (metabolic performance trade-off relationships) and pronounced phase states of temperature acclimation are associated with disproportionately enhanced (in smp-p phase state) or reduced (rmp-t phase state) smp-tw efficiency in responding to dry-season rainfall and producing high-quality forage in general. in the central knp, with intermediate rmp–smp mode settings and higher actual dry-season rainfall, a postulated favourable product of smp-tw efficiency and dry-season rainfall seems to support the relatively sustained population performance of blue wildebeest and zebra. the higher rainfall in the southern knp underpinned relative rmp mode settings (cvrm-1), which are inherently associated with poorer forage quality (lower carbon-nutrient quality, due to reduced storage allocation, and lowered nitrogen quality, due to dilution by structural carbon resulting from high tw growth priority). here selective grazers, mainly sable antelope, were sustained only at the highest altitudes and because of reasonable amounts of dry-season rainfall and the availability of carbon-nutrient quality forage procurable from andropogonoid grass species. extending the emerging pattern to near tropical conditions (rmp-tw metabolic mode settings), it follows that selective grazers can persist under such circumstances only if substantial dry-season rainfall occurs and suitable andropogonoid grass species are commonly available as forage plants. this seems to be the basis for the stable persistence of roan antelope in moist dystrophic west african savannas receiving 800 mm rainfall or more per year (de bie 1991). blue wildebeest and zebra do not occur under such conditions where extreme nitrogen dilution of forage items is encountered (rmp-tw functionality). whereas blue wildebeest and zebra populations performed poorly at extremes of the rainfall gradient (low rainfall: nitrogen dilution by storage products associated with smp functionality and nitrogen dilution by structural carbon at higher rainfall: rmp-tw functionality), selective grazer species may cope with such conditions. at the arid extreme, smp modes of plant functionality support forage with carbon-nutrient quality (given sufficiently low ambient temperatures or high altitude) and browsing then seems to be particularly advantageous (eland; roan antelope in more arid parts of west africa; de bie 1991). at the mesic end of the gradient, sufficient amounts of dry-season rainfall and andropogonoid grass species form the basis of their persistence. blue wildebeest and zebra populations perform well under intermediate smp–rmp plant functionality (central knp), where nitrogen quality of forage is favoured by relative smp-tw efficiency of plants, reasonable amounts of dry-season rainfall, growth activity curbed through moisture stress (smp-tw functionality) and nutrient-rich substrates. however, over the long term, populations of selective grazers declined (figure 2a) because of plants’ progressive acclimation to rising temperatures (since about 1910: figure 1), which compromised storage-based carbon-nutrient quality of available forage (cvrm-2.2). in contrast, rising ambient temperature, by enhancing nitrogen productivity under conditions of smp-tw plant functionality, increased the availability of forage with adequate nitrogen quality, expectedly thereby favouring increased population densities of blue wildebeest and zebra (figure 2b). bottom-up and top-down impacts on large herbivore population dynamics owen-smith and mills (2006) presented results of comprehensive analyses indicating that changes in population abundance within the multi-prey, general predator system in the knp arose from a complex interplay between changing climatic conditions, variable food production, shifting habitat conditions, varying vulnerability to predation and spill-over effects on other species. findings by owen-smith and mason (2005) highlighted the importance of changes in adult survival rates in shaping population trends, implicating an interaction with nutritional shortfalls and the central involvement of predation by lions (panthera leo), which prey largely on the adult segment of most of the larger herbivore species (owen-smith et al. 2005). patterns of shifting prey selection depending on changing relative abundance and vulnerability of the three principal prey species (wildebeest, zebra and buffalo) further implicated predation as the main cause of the decline of alternative prey species such as waterbuck and the selectively feeding antelope species (owen-smith & mills 2008a). however, owen-smith and mills (2006) caution that attempting to separate the relative importance of top-down influences through predation from bottom-up processes operating through food resources is difficult, as susceptibility to predation depends on changing nutritional conditions and other habitat conditions (concealment), particularly as these factors also affect predator populations. the results of this study permit an interpretation of these complex interacting patterns, placing greater emphasis on bottom-up, climatically linked nutritional factors relative to predation impacts.prey selection for both buffalo and waterbuck was inversely dependent on rainfall averaged over three preceding years (owen-smith & mills 2008a). both buffalo and waterbuck seemed more vulnerable to malnutrition, and thus to predation, during periods of low rainfall (owen-smith & mills 2006; owen-smith 2008; owen-smith & mills 2008a). despite heavy predation by lions on waterbuck (mills et al. 1995), the resultant mortality did not seem to be additive and more important than the negative effects of drought, and populations increased during normal years despite heavy predation on the species by lions and other predators (pienaar 1969). this also appears to be the case for buffalo. as forage resources improved after the 1991–1992 drought, both species showed recovery in abundance (figures 3c-d). prey selection on these species is thus considered rather to be a consequence of malnutrition and not the ultimate cause of declines in abundance. wildebeest and zebra were found to be more susceptible to predation under conditions of above-average rainfall. this was attributed to prey selection shifting towards these species owing to their susceptibility to predation being increased through the effect of higher vegetation cover facilitating hunting success of lions (owen-smith & mills 2008a). reduced exposure to predation was apparently associated with increasing abundance of these two species. also, higher predation pressure on adult wildebeest than on zebra (owen-smith & mills 2008b) was believed to have resulted in higher population numbers for zebra than for wildebeest in recent decades (joubert 2007b). these patterns were interpreted to support the contention that wildebeest and zebra, the primary prey species selected by lions in the knp (owen-smith & mills 2008b), were held below the food ceiling through additive predation (owen-smith 2008). however, following from the results of this study, some of these patterns can be interpreted alternatively with reference to the dynamics of nitrogen quality of the forage selected by these two species. accordingly, the reduced population performance during periods of above-average rainfall is attributed to reduced nitrogen quality due to nitrogen dilution associated with high grass productivity. between 1955 and 1970 the abundance of zebra commenced to exceed that of wildebeest and stabilised at higher population levels. according to our study, this was interpreted as resulting from zebra being tolerant to relatively lower nitrogen quality forage than wildebeest, as such quality was expected to decline with increasing nitrogen productivity and the associated progressively increasing tw growth priority over this time span (climate–vegetation response model). the provision of numerous artificial water points has been implicated to have benefited common water-dependent species, resulting in the influx of blue wildebeest and zebra into the range of the less water-dependent selective grazer species (gaylard, owen-smith & redfern 2003). population declines of selective grazer species were consequently attributed to increased predation owing to the associated build-up of lion numbers in their ranges (harrington et al. 1999). however, in spite of extensive water provisioning, the populations of blue wildebeest and zebra continued to fluctuate and the prolonged period of relatively low population densities of these species since the early 1990s in the northern knp (figures 3a–b) did not result in the recovery of selective grazer populations. evidence of a marked shift in prey selection by lions towards the selectively feeding antelope species around 1987 was reported by owen-smith and mills (2008a). the 2–3-fold increase in relative prey selection for these species was considered adequate to explain a similar rise in adult mortality observed after 1986 (owen-smith & mason 2005; owen-smith & mills 2008a). this shift in predation towards the selectively feeding antelope species was apparently induced by (1) lowered abundance and hence lower availability of buffalo as prey following the 1982–1983 drought and (2) coupled with reduced susceptibility of wildebeest and zebra over the period of low rainfall that set in during 1987. with all three principal prey species less available, lions apparently turned towards alternative prey species, including the selectively feeding antelope species and waterbuck (owen-smith & mills 2008a). however, although the involvement of predation in the decline of selectively feeding antelope species after 1986 is implicated, it is difficult to decouple the interactive predator–prey dynamics from the extrinsically driven context provided by changing food resource conditions (owen-smith et al. 2005; owen-smith & mills 2006). the results of this study support an interpretation of the decline of these antelope species involving an interaction between compromised forage quality and a resulting increased vulnerability to predation as a consequence. as accelerated rising temperatures since about 1980 (figures 1 and 7) increasingly compromised carbon-nutrient quality of forage selected by selectively feeding antelope species (climate–vegetation response model), their nutritionally linked condition apparently declined and had involved most adults of the populations by 1986/1987. rapid population declines ensued, with no signs of recovery to date (figures 4a-d). in contrast, as food resource conditions (rainfall-based forage quantity) improved for waterbuck – a species heavily predated on by lions – their population numbers progressively increased (figure 3d), whilst the population numbers of the selectively feeding antelope species remained low. according to the interpretation of this study the absence of recovery of these antelope populations is the result of continued low levels of carbon-nutrient quality of available forage due to sustained elevated ambient temperatures (figures 6 and 7). although impacts of predation on the small remaining populations of selectively feeding antelope species cannot be ignored (grant et al. 2002), we conclude that even though proximate impacts of predation are considered of significance, the ultimate cause of population performance trends of these species appears to be related to the climatically compromised carbon-nutrient quality of their forage. conclusion top ↑ from a situation apparently prevailing during the first two to three decades of the 1900s, during which the abundance of selective grazers (sable antelope, roan antelope and tsessebe) was of similar orders of magnitude to that of bulk feeding species (wildebeest, zebra and buffalo), the former declined progressively, whilst the latter increased over subsequent decades. these herbivore population trends were paralleled by progressively enhanced nitrogen productivity, increasing the availability of nitrogen quality forage and declining carbon-nutrient quality (storage-based plant metabolism), apparently linked to increasing temperature over time. further analysis revealed that population numbers of buffalo and waterbuck, as roughage bulk feeders, tracked forage quantity largely determined by rainfall. wildebeest and zebra, with their preference for grazing short grasses, were identified as nitrogen quality patch selectors. as such their population performance was favoured in habitats of high nutrient status with sustained nitrogen productivity and nitrogen quality, mainly found in the southern (central) knp. population performance of these species, as more clearly manifesting in the northern knp, was negatively associated with reduced dry-season rainfall and elevated temperatures, a syndrome of climate effects expected to reduce forage nitrogen quality through nitrogen dilution. the population performance of antelope species selecting for carbon-nutrient quality plant parts (sable antelope, roan antelope, eland and tsessebe) was relatively enhanced at higher altitude sites and in northern knp. relatively lower temperatures associated with upper altitudes were expected to conserve carbon-nutrient quality of forage and the prevalence of the sustained metabolic performance mode of plants in the northern knp to support relative plant carbon excess (climate–vegetation response model). population performance of these antelope species was found to be spatiotemporally negatively associated with high temperature conditions, consistent with predictions of the climate–vegetation response model, according to which carbon-nutrient quality is compromised by elevated ambient temperature. accordingly, the long-term population decline and the absence of population recovery of these species to date were attributed to progressively increasing and sustained elevated ambient temperature conditions as progressive smp-to-rmp temperature acclimation reduced the scope for storage-based metabolism in favour of biomass productivity (growth).the results of pattern/process matching conducted in this study revealed remarkable consistency between herbivore population trends and key indices of forage quantity and quality, shaped in accordance with expectations of plant responses to climate (climate–vegetation response model). with regard to all eight species studied, their targeting by lion predation through shifting prey selection, as implicated by owen-smith and mills (2008a), apparently coincided with time periods of compromised forage quantity or quality as predicted by climate–vegetation response processes in this study. population trends of these species were found to be in apparent synchrony with features of a changing climate (rainfall, dry-season rainfall, and maximum and minimum ambient temperatures). furthermore, some of the herbivore population trends, such as the lack of recovery of populations of the selectively feeding antelope species over recent years but population increases for waterbuck over the same period, cannot be readily explained with reference to predation, but conform to climatically driven forage availability as outlined in this study. we conclude that ultimate causality of the observed population trends of the species studied is linked to landscape-scale climate–vegetation responses controlling the availability of suitable forage (bottom-up regulation). proximate impacts of predation additionally shaped the basic population dynamics in a complex and interactive manner (top-down impacts). acknowledgements top ↑ we acknowledge the effort of rangers, other field staff and scientists with regard to collecting diverse data sets (e.g. animal population censuses and grassland monitoring) over many decades, without which this study would not have been possible. mss judith kruger, sandra macfadyen and sharon thomson of scientific services, knp are acknowledged for providing data used in this study. we also thank prof. herbert prins and ignas heitkönig for valuable discussions (both from wageningen university). the south african weather service provided temperature and rainfall data. this study represents an output of the specialist scientist research programme in systems ecology conducted by members of the conservation services division (knysna and skukuza) of south african national parks. the main author, in his capacity as honorary research associate at the botany department, university of cape town thankfully acknowledges general support received from this institution. competing interests the authors declare that they have no financial or personal relationship(s) which may have inappropriately influenced them in writing this paper. authors’ contributions a.h.s. (garden route national park) was responsible for the development of the climate-vegetation response model and the interpretation of large herbivore population performance patterns and trends in accordance with this model. c.c.g. (kruger national park), i.p.s. (kruger national park), w.j.v. (garden route national park) and n.z. (kruger national park) contributed during in-depth discussions by sharing their knowledge of plant and animal ecology. i.p.s. (kruger national park) also performed some statistical analyses and j.b. (garden route national park) was involved in the processing of spatial data used for analyses and the production of figures. references top ↑ an, y., wan, s., zhou, z., subedar, a.a., wallace, l.l. & luo, y., 2005, ‘plant nitrogen concentration, use efficiency, and contents in a tallgrass prairie ecosystem under experimental warming’, global change biology 11, 1733–1744. http://dx.doi.org/10.1111/j.1365-2486.2005.01030.xanselin, l., 2005, exploring spatial data with geodatm: a workbook, university of illinois, urbana-champaign. archives, kruger national park (skukuza), annual reports, sabi and shingwitsi game reserves, 1902–1925; kruger national park, 1926–1945. archives, skukuza, warden’s annual reports: kruger national park, unpublished reports, 1946–1952. bodenstein, v., meissner, h.h. & van hoven, w., 2000, ‘food selection by burchell’s zebra and blue wildebeest in the timbavati area of the northern province lowveld’, south african journal of wildlife research 30(2), 6–72. buckland, s.t., anderson, d.r., burnham, k.p. & laake, j.l., 1993, distance sampling estimating abundance of biological populations, chapman, london. de bie, s., 1991, wildlife resources of the west african savanna, wageningen agricultural university papers 91–92, wageningen. dewar, r.c., medlyn, b.e. & mcmurtrie, r.e., 1999, ‘acclimation of the respiration/ photosynthesis ratio to temperature: insights from a model’, global change biology 5, 615–622. http://dx.doi.org/10.1046/j.1365-2486.1999.00253.x dunham, k.m., robertson, e.f. & grant, c.c., 2004, ‘rainfall and the decline of a rare antelope, the tsessebe (damaliscus lunatus lunatus), in kruger national park, south africa’, biological conservation 117, 83–94. http://dx.doi.org/10.1016/s0006-3207(03)00267-2 dwyer, s.a., ghannoum, o., nicotra, a. & von caemmerer, s., 2007, ‘high temperature acclimation of c4 photosynthesis is linked to changes in photosynthetic biochemistry’, plant, cell and environment 30, 53–66. http://dx.doi.org/10.1111/j.1365-3040.2006.01605.x, pmid:17177876 east, r., 1984, ‘rainfall, soil nutrient status and biomass of large african savanna mammals’, african journal of ecology 22, 245–270. http://dx.doi.org/10.1111/j.1365-2028.1984.tb00700.x environmental systems research institute (esri), 2006, esri data and maps: elevation and image data, esri, redlands, california, usa. estiarte, m. & peñuelas, j., 1999, ‘excess carbon: the relationship with phenotypical plasticity in storage and defense functions of plants’, orsis 14, 159–203. fritz, h. & duncan, p., 1994, ‘on the carrying capacity for large ungulates of african savanna ecosystems’, proceedings of the royal society of london b 256, 77–82. http://dx.doi.org/10.1098/rspb.1994.0052, pmid:8008761 gaylard, a., owen-smith, n. & redfern, j., 2003, ‘surface water availability: implications for heterogeneity and ecosystem processes’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 171–188, island press, washington dc. gertenbach, w., 1983, ‘landscapes of the kruger national park’, koedoe 26, 9–121. grange, s. & duncan, p., 2006, ‘bottom-up and top-down processes in african ungulate communities: resources and predation acting on the relative abundance of zebra and grazing bovids’, ecography 29, 899–907. http://dx.doi.org/10.1111/j.2006.0906-7590.04684.x grant, c.c., davidson, t., funston, p.j. & pienaar, d.j., 2002, ‘challenges faced in the conservation of rare antelope: a case study on the northern basalt plains of the kruger national park’, koedoe 45(2), 45–62. harrington, r., owen-smith, n., viljoen, p.c., mason, d.r. & funston, p.j., 1999, ‘establishing the causes of the roan antelope decline in the kruger national park, south africa’, biological conservation 90, 69–78. http://dx.doi.org/10.1016/s0006-3207(98)00120-7 heitkönig, i.m.a. & owen-smith, n., 1998, ‘seasonal selection of soil types and grass swards by roan antelope in a south african savanna’, african journal of ecology 36, 57–70. http://dx.doi.org/10.1046/j.1365-2028.1998.114-89114.x herms, d.a. & mattson, w.j., 1992, ‘the dilemma of plants: to grow or defend’, quarterly review of biology 67(3), 283–335. http://dx.doi.org/10.1086/417659 hofmann, r.r., 1973, the ruminant stomach. stomach structure and feeding habits of east african game ruminants, vol. ii, east african literature bureau, nairobi. hofmann, r.r., 1989, ‘evolutionary steps of ecophysiological adaptation and diversification of ruminants: a comparative view of their digestive systems’, oecologia 78, 443–457. http://dx.doi.org/10.1007/bf00378733 houghton, j.t., meira filho, l.g., callander, b.a., harris, n., kattenberg a. & maskell, k., (eds.), 1996, climate change 1995: the science of climate change, cambridge university press, cambridge. joubert, s.c.j., 1976, ‘the population ecology of the roan antelope (hippotragus equinus equinus) (desmarest, 1804) in the kruger national park’, dsc thesis, faculty of science, university of pretoria. joubert, s.c.j. & bronkhorst, p.h.l., 1977, ‘some aspects of the history and population ecology of the tsessebe damaliscus lunatus lunatus in the kruger national park’, koedoe 20, 125–145. joubert, s., 2007a, the kruger national park: a history, vol. i., high branching, johannesburg. joubert, s., 2007b, the kruger national park. a history, vol. ii., high branching, johannesburg. kennedy, a.d., biggs, h. & zambatis, n., 2003, ‘relationship between grass species richness and ecosystem stability in kruger national park, south africa’, african journal of ecology 41, 131–140. http://dx.doi.org/10.1046/j.1365-2028.2003.00391.x kruger, j.m., reilly, b.k. & whyte, i.j., 2008, ‘application of distance sampling to estimate population densities of large herbivores in kruger national park’, wildlife research 35, 371–376. http://dx.doi.org/10.1071/wr07084 le roux, e., 2010, ‘habitat and forage dependency of sable antelope (hippotragus niger) in the pretoriuskop region of the kruger national park’, msc thesis, school of animal, plant & environmental sciences, university of the witwatersrand. legendre, b. & mcardle, b.h., 1997, ‘comparison of surfaces’, oceanologica acta 20, 27–41. ludorf, j.f., 1918, report of the game reserve commission, government printer, pretoria. macandza, v.a., owen-smith, n. & cross, p.c., 2004, ‘forage selection by african buffalo in the late dry season in two landscapes’, south african journal of wildlife research 34(2), 113–121. magome, h., cain, j.w.i., owen-smith, n. & henley, s.r., 2008, ‘forage selection of sable antelope in pilanesberg game reserve, south africa’, south african journal of wildlife research 38(1), 35–41. http://dx.doi.org/10.3957/0379-4369-38.1.35 ‘t mannetje, l., 1982, ‘problems of animal production from tropical pastures’, in j.b. hacker (ed.), nutritional limits to animal production from pastures, pp. 67–85, commonwealth agricultural bureaux, farnham royal. ‘t mannetje, l., 1984, ‘nutritive value of tropical and subtropical pastures, with special reference to protein and energy deficiency in relation to animal production’, in f.m.c. gilchrist & r.i. mackie (eds.), herbivore nutrition in the subtropics and tropics, pp. 56–66, the science press, craighall. mduma, s.a.r., sinclair, a.r.e. & hilborn, r., 1999, ‘food regulates the serengeti wildebeest: a 40-year record’, journal of animal ecology 68, 1101–1122. http://dx.doi.org/10.1046/j.1365-2656.1999.00352.x mills, m.g.l., biggs, h.c. & whyte, i.j., 1995, ‘the relationship between rainfall, lion predation and population trends in african herbivores’, wildlife research 22, 75–88. http://dx.doi.org/10.1071/wr9950075 mutanga, o., prins, h.h.t., skidmore, a.k., van wieren, s., huizing, h., grant, r. et al., 2004, ‘explaining grass-nutrient patterns in a savanna rangeland of southern africa’, journal of biogeography 31, 819–829. http://dx.doi.org/10.1111/j.1365-2699.2004.01072.x osborne, c.p., 2008, ‘atmosphere, ecology and evolution: what drove the miocene expansion of c4 grasslands?’, journal of ecology 96, 35–45. pmid:18784799, pmid:2517376 owen-smith, n., 2008, ‘changing vulnerability to predation related to season and sex in an african ungulate assemblage’, oikos 114, 602–610. http://dx.doi.org/10.1111/j.0030-1299.2008.16309.x owen-smith, n. & ogutu, j.o., 2003, ‘rainfall influences on ungulate population dynamics’, in j.t. du toit, h.c. biggs & k.h. rogers (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 310–331, island press, washington dc. owen-smith, n. & mason, d.r., 2005, ‘comparative changes in adult versus juvenile survival affecting population trends of african ungulates’, journal of animal ecology 74, 762–773. http://dx.doi.org/10.1111/j.1365-2656.2005.00973.x owen-smith, n., mason, d.r. & ogutu, j.o., 2005, ‘correlates of survival rates of ten african ungulate populations: density, rainfall and predation’, journal of animal ecology 74, 774–778. http://dx.doi.org/10.1111/j.1365-2656.2005.00974.x owen-smith, n. & mills, m.g.l., 2006, ‘manifold interactive influences on the population dynamics of a multispecies ungulate assemblage’, ecological monographs 76, 73–92. http://dx.doi.org/10.1890/04-1101 owen-smith, n. & mills, m.g.l., 2008a, ‘shifting prey selection generates contrasting herbivore dynamics within a large-mammal predator-prey web’, ecology 89(4), 1120–1133. http://dx.doi.org/10.1890/07-0970.1, pmid:18481536 owen-smith, n. & mills, m.g.l., 2008b, ‘predator-prey size relationships in an african large-mammal food web’, journal of animal ecology 77, 173–183. http://dx.doi.org/10.1111/j.1365-2656.2007.01314.x, pmid:18177336 parrini, f., 2006, ‘nutritional and social ecology of the sable antelope in a magaliesberg nature reserve’, phd thesis, school of animal, plant & environmental sciences, university of the witwatersrand. pienaar, u de v., 1963, ‘the large mammals of the kruger national park: their distribution and present-day status’, koedoe 6, 1–37. pienaar, u de v., 1969, ‘predator-prey relationships among the larger mammals of the kruger national park’, koedoe 12, 108–176. scholes, r.j. & walker, b.h., 1993, an african savanna: synthesis of the nylsvley study, cambridge university press, cambridge. http://dx.doi.org/10.1017/cbo9780511565472 schulze, r.e., 1995, hydrology and agrohydrology: a text to accompany the acru 3.00 agro-hydrological modeling system, university of natal (department of agricultural engineering), pietermartizburg. seydack, a.h., grant, c.c., smit, i.p., vermeulen, w.j., baard, j. & zambatis, n., 2012, ‘climate and vegetation in a semi-arid savanna: development of a climate-vegetation response model linking plant metabolic performance to climate and the effects on forage availability for large herbivores’, koedoe 54(1), art. #1046, 12 pages. http://dx.doi.org/10.4102/koedoe.v54i1.1046 silverman, b.w., 1986, density estimation for statistics and data analysis, chapman and hall, new york. sinclair, a.r.e., dublin, h. & borner, m., 1985, ‘population regulation of serengeti wildebeest: a test of the food hypothesis’, oecologia 65, 266–268. http://dx.doi.org/10.1007/bf00379227 skinner, j.d. & chimimba, c.t., 2005, the mammals of the southern african subregion, 3rd edn., cambridge university press, cambridge/cape town. smit, i.p.j., 2011, ‘resources driving landscape-scale distibution patterns of grazers in an african savanna’, ecography 34, 67–74. http://dx.doi.org/10.1111/j.1600-0587.2010.06029.x smuts, g.l., 1972, ‘seasonal movements, migration and age determination of burchell’s zebra (equus burchelli antiquorum, h. smith, 1841) in the kruger national park’, msc thesis, faculty of science, university of pretoria. steyn, l.b., 1953–1961, ‘annual reports: kruger national park’, unpublished reports, skukuza, south africa. steyn, l.b., 1954, report of the supervisor of game censuses in 1954 in the kruger national park, south african national parks, south africa. van soest, p.h., mertens, d.r. & deinum, b., 1978, ‘preharvest factors influencing quality of conserved forage’, journal of animal science 47, 712–720. venter, f.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation patterns’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83–129, island press, washington dc. viljoen, p.c., 1996, ecological aerial surveys in the kruger national park, report 04/1995, kruger national park (scientific services), skukuza. wan, s., xia, j., liu, w. & niu, s., 2009, ‘photosynthetic overcompensation under nocturnal warming enhances grassland carbon sequestration’, ecology 90(10), 2700–2710. http://dx.doi.org/10.1890/08-2026.1, pmid:19886480 watson, l.h. & owen-smith, n., 2000, ‘diet composition and habitat selection of eland in semi-arid shrubland’, african journal of ecology 38, 130–137. http://dx.doi.org/10.1046/j.1365-2028.2000.00229.x whyte, i.j., 2007, rare antelope in the kruger national park: trends, current status, and options for future management, report 03/2007, south african national parks, skukuza. whyte, i.j. & joubert, s.c.j., 1988, ‘blue wildebeest population trends in the kruger national park and the effect of fencing’, south african journal of wildlife research 18(3), 78–87. wilson, j.r., 1982, ‘environmental and nutritional factors affecting herbage quality’, in j.b. hacker (ed.), nutritional limits to animal production from pastures, pp. 111–131, commonwealth agricultural bureaux, farnham royal. wilson, j.r. & ford, c.w., 1971, ‘temperature influences on the growth, digestibility, and carbohydrate composition of two tropical grasses, panicum maximum var. trichoglume and setaria sphacelata, and two cultivars of the temperate grass lolium perenne’, australian journal of agricultural research 22, 563–571. http://dx.doi.org/10.1071/ar9710563 wilson, j.r. & ford, c.w., 1973, ‘temperature influences on the in vitro digestibility and soluble carbohydrate accumulation of tropical and temperate grasses’, australian journal of agricultural research 24, 187–198. http://dx.doi.org/10.1071/ar9730187 footnote top ↑ 1.for ease of reading, abbreviations and symbols are used to describe factors as follows: np, nitrogen productivity; nq, nitrogen quality, cnq, carbon-nutrient quality; ↑, increasing; ↓, decreasing. a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe original research vol. 50 no. 1 pp. 61 71 vegetation of the hantam-tanqua-roggeveld subregion, south africa. part 1: fynbos biome related vegetation abstract the succulent karoo hotspot stretches along the western side of the republic of south africa and namibia. a lack of botanical information on the hantam-tanqua-roggeveld area of the succulent karoo hotspot was identified during the skep (succulent karoo ecosystem plan) process. a grant from cepf (critical ecosystem partnership fund) funded a study to produce a vegetation map of the area to serve as baseline for ecosystem management. vegetation surveys were conducted over an area of more than three million hectares from august to october 2004. two major floristic units were identified, namely the fynbos biome related (mountain renosterveld) and succulent karoo biome related units. an analysis of the floristic data of the predominantly mountain renosterveld vegetation unit is presented in this paper. three associations were identified, which were subdivided into nine subassociations, one of which contains four variants. the vegetation units are described in terms of their species composition and their relationships with the physical environment. a vegetation map is provided depicting the geographical distribution of the different vegetation types. the main threat to the vegetation of the region identified by the farming community was a lack of infrastructure. keywords: mountain renosterveld, phytosociology, succulent karoo, vegetation classification, vegetation map the succulent karoo, which stretches along the western side of south africa and namibia, is recognised by the iucn as one of the global hotspots of biodiversity (myers et al. 2000, cepf 2003) and one of only two hotspots that are entirely arid (conservation international – website). in 2002 the succulent karoo ecosystem plan (skep) was launched to identify and generate consensus for a 20-year conservation and sustainable land-use strategy for the succulent karoo hotspot. skep aims to meet the quantitative targets for the conservation of vegetation and globally threatened and endemic species at particular sites, as well as critical ecological and evolutionary processes that must be conserved to ensure the persistence of these species (conservation international – website). for management purposes, the skep initiative subdivided the succulent karoo into four subregions, of which the hantamtanqua-roggeveld constituted one. in common with the rest of the succulent karoo, the vegetation of the hantam-tanquaroggeveld subregion includes a wide range of succulents, geophytes and annuals. after good rains, the spectacular autumn and spring displays of wild flowers in parts of the region attract large numbers of tourists. unlike many parts of namaqualand, such brilliant shows of annuals and geophytes are not only a feature of fallow fields, but also occur in the undisturbed natural vegetation in the hantam and roggeveld (van wyk & smith 2001). the identification, description and classification of vegetation units across the landscape comprise the critical first steps in building a framework for ecosystem management planning. information on the spatial, temporal and ecological properties of the vegetation units can lead to the improved understanding, protection and management of natural resources. progression of the skep initiative soon showed the paucity of data available on the biodiversity of the hantam-tanqua-roggeveld subregion, which was key to future planning, conservation and development. the critical ecosystem partnership fund (cepf), which is a joint initiative of conservation international, the global environmental facility, the government of japan, the macarthur foundation and the world bank, therefore granted funding for botanical studies in the subregion. the first step of the botanical study was to undertake a systematic broad-scale vegetation survey of the entire subregion of approximately three million hectares, which could be used as the basis for further detailed botanical investigations. the survey revealed two distinct vegetation groups, i.e. the fynbos biome related mountain renosterveld vegetation group and the succulent karoo biome related vegetation group. the aim of the present article is to report on the mountain renosterveld vegetation, depicting its component vegetation units on a map. a second article (van der merwe et al. in press) will report on the latter vegetation group. study area the hantam-tanqua-roggeveld subregion (fig. 1), as defined in the current study, lies in the predominantly winter rainfall region of the northern and western cape provinces of south africa, and covers an area of approximately three million hectares. in the west it stretches from east of the cederberg mountains helga van der merwe margaretha w. van rooyen noel van rooyen department of plant science university of pretoria south africa correspondence to: helga van der merwe e-mail: soekop@hantam.co.za postal address: department of plant science, university of pretoria, pretoria, 0002 61 2008 original research van der merwe, van der merwe & van rooyen koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 61 71 in the southwestern corner, northwards along the bokkeveld mountains to just north of loeriesfontein. the eastern border includes the roggeveld and nuweveld mountain ranges to just southwest of fraserburg, while the southern limit includes the tanqua and ceres karoo to where the swartrug mountains and the bontberg mountains meet north of ceres. the mountain renosterveld discussed in the current article is found on the roggeveld, nuweveld, komsberg, klein roggeveld, koedoesberg and hantam mountains. in general, this is the higher-lying part of the larger subregion that is actually situated in the fynbos biome (rutherford & westfall 1986). this area includes acocks’s (1988) mountain renosterveld (veld type 43), which is equivalent to the escarpment mountain renosterveld (unit 60) of low and rebelo (1998). according to mucina et al. (2005), six vegetation types are represented in the area, namely the nieuwoudtville shale renosterveld (frs 2); the roggeveld shale renosterveld (frs 3); the central mountain shale renosterveld (frs 5); the nieuwoudtville roggeveld dolerite renosterveld (frd 1); the hantam plateau dolerite renosterveld (frd 2); and the roggeveld karoo (skt 3). the earliest references to the botanical wealth of the hantamroggeveld date from the early 1900s. diels (1909 in van wyk & smith 2001) mentioned the high levels of endemism in the hantam-roggeveld and provided a useful floristic analysis of the region. he concurred with marloth (1908) that the region is floristically more closely related to the succulent karoo and the great karoo than to the cape floristic region, although cape floristic elements are clearly present, especially on the hantamsberg (van wyk & smith 2001). the roggeveld was also one of the three centres of endemism that hilton-taylor (1994) identified within the western cape domain, the other two centres being the western mountain karoo and tanqua karoo, which also fall within the hantam-tanqua-roggeveld subregion. van wyk and smith (2001) combined the hantamroggeveld into one of their 13 principal centres of plant endemism in southern africa and stressed the unique botanical importance of this area. the rainfall ranges from 132–467 mm per year (weather bureau 1998), which, although falling mainly in winter, includes a few summer thunderstorms. in 2004 the rainfall season was poor and the usual winter snowfalls on the high-lying areas were limited to the light snow that fell on one occasion, compared with the mean of six snow days recorded over a 24-year period by the weather bureau (1998). at sutherland the mean maximum for the warmest month, january, is 27.1°c, while, the extreme maximum recorded was 35.5°c in january 1980 (weather bureau 1998). the mean minimum for the coldest month, july, is –2.4°c, while the extreme minimum, -13.6°c, was recorded in july 1970 and august 1978 (weather bureau 1998). rocks of the ecca group cover most of this area with dwyka (consisting of tillite, sandstone, mudstone and shale) cropping out in the west and the beaufort group in the east (council for geoscience 1973, 1983, 1989, 1991, 1997, 2001). the ecca group includes sediments of the koedoesberg formation (consisting of sandstone and shale) and the tierberg formation (consisting of shale) (council for geoscience 1973, 1983, 1989, 1991, 1997, 2001). mudstones of the beaufort group are found on the eastern side of the study area (council for geoscience 1973, 1983, 1989, 1991, 1997, 2001). igneous intrusions of dolerite occur throughout the region, being easily recognisable as very hard, dark grey to nearly black rocks (van wyk & smith 2001). the soils of the roggeveld consist primarily of clays and silts derived from the karoo sequence shales (low & rebelo 1998) and are found on the slopes and foothills of the great escarpment along the various mountain ranges. methods and materials satellite images (bands: 4, 5 and 3 (r,g,b)) of the study area were visually stratified into relatively homogeneous units on the basis of colour, texture and topography. this stratification was used to select the sites at which sample plots were surveyed from august until october 2004. at each site gps (global positioning system) coordinates were taken and each species present in the plot was noted and assigned a cover-abundance value according to the braun blanquet cover-abundance scale (werger 1974). various environmental characteristics, such as altitude, topography, aspect, slope, an estimation of rock cover, the size of the rocks, soil type and colour, and the degree of erosion were noted at each sampling point. biotic effects, such as trampling, small mammal activity, or invasion by alien plants, were also recorded. a total of 390 sample plots covering the entire hantam-tanquaroggeveld subregion were surveyed in 2004. an analysis of the floristic data was undertaken using the turboveg and megatab computer package (hennekens & schaminée 2001). vegetation data were captured with the turboveg software and the data were classified with the aid of megatab. as a first step to the classification of the floristic data, a two-way indicator species analysis (twinspan) (hill 1979) was run in megatab. the result of the twinspan on the entire data set confirmed the presence of two distinct floristic groups, which enabled the data set to be split into two. a twinspan was then run separately on each data set, with the resulting tables being further refined to obtain clear species assemblages. the first phytosociological table, which characterises the vegetation of the predominately mountain renosterveld as defined by acocks (1988), is discussed in the current article. the major vegetation units distinguished in the mountain renosterveld were termed associations following the use as defined by nelder et al. (2005). associations are produced on the basis of the presence and abundance of species, vegetation structure and the spatial distribution of individuals in the dominant layer. subassociations are generally distinguished on the basis of elements in the subdominant layers. the subassociations are described in terms of a list of species featuring each structural layer, together with its canopy cover. figure 1 subregions in the skep planning domain (cepf 2003). 62 fynbos biome related vegetation original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 61 71 fi g u r e 2 * v eg et at io n m ap o f t he m ou nt ai n r en os te rv el d ve ge ta ti on o f t he h an ta m -t an qu ar og ge ve ld s ub re gi on . * e nl ar ge d fi gu re is a va ila bl e on lin e. 63 original research van der merwe, van der merwe & van rooyen koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 61 71 r e l e v é n u m b e r 3 3 3 3 3 3 3 3 3 3 3 1 2 3 2 3 1 2 1 2 2 2 3 1 3 3 2 3 3 3 3 3 3 2 2 3 2 3 3 3 3 2 2 2 1 1 1 1 1 1 1 2 3 3 1 3 3 3 2 2 2 2 3 2 3 1 2 1 1 3 1 1 1 1 1 3 3 3 3 2 3 3 1 1 1 3 3 1 1 3 3 3 3 1 1 1 2 3 1 1 1 1 1 2 4 6 4 9 0 4 8 3 8 7 7 4 5 0 0 7 0 3 3 2 2 2 8 7 3 9 2 2 3 4 7 8 8 9 7 9 4 3 9 5 3 9 2 3 3 3 4 4 3 8 3 3 6 6 7 8 5 8 8 3 8 3 2 5 2 8 8 8 8 4 4 6 2 0 4 5 8 9 6 6 6 3 7 4 4 5 9 9 4 6 0 9 6 7 6 7 3 4 5 1 2 0 1 2 6 7 0 5 9 1 2 7 4 4 8 3 1 5 3 0 5 6 4 9 0 3 3 4 9 2 2 5 1 5 6 6 2 3 5 8 5 2 9 6 1 6 3 2 8 5 4 0 3 4 5 1 2 4 9 3 5 5 9 2 1 6 7 0 8 9 2 2 7 5 1 3 7 9 4 4 0 8 5 2 3 1 2 7 4 9 8 6 4 4 2 0 6 1 0 8 0 9 1 1. 1 1. 2 1. 3 1 .4 1 .5 2. 1. 1 2 .1 .2 2. 1. 3 2. 1. 4 2 .2 2. 3 3 s p e c ie s g r o u p a e ri oc ep ha lu s m ic ro ph yl lu s 1 . a a . . . . 1 . a . . . . . . . . . + + . . + . . . + . . . . . . . . . . . . . . . . . . . + . . . . . . + . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . p en tz ia ( h r p3 17 ) sp . . + + . . + + + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e ur yo ps im br ic at us 1 1 . . 1 . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . d io sp yr os a us tr o -a fr ic an a r . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . + s p e c ie s g r o u p b a nt im im a cf . g ra ni tic a (h r 24 8) . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . + . . . . . . . p el ar go ni um s p. . . . . . . . . . . . + . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . s en ec io e ro su s . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . lo to no ni s sp . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . a sp ar ag us a sp ar ag oi de s . . . . . . . . . . . + . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p c s al vi a di se rm as . . . . . . . . . . . . . . . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . c hr ys an th em oi de s in ca na . . . . . . . . . . . . . . . . . + . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a pt os im um in di vi su m . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s tip ag ro st is n am aq ue ns is . . . . . . . . . . . . . . . . . . a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b ra un si a sp . . . . . . . . . . . . . . . . . . a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p d p en tz ia in ca na . . . . . . . . . . . . + . . . . + + . + + . + + + . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . + . . . . . . . m es em br ya nt he m um g ue ri ch ia nu m . . . . . . . . . . . . . . . . + . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a nd ro cy m bi um v ol ut ar e . . . . . . . . . . . . . . . . + . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p e e ur yo ps m ul tifi du s . . . . + + . . . . . . . . . . . + . . . . . . . . . + 1 + + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p hy llo bo lu s te nu ifl or us . . . . . . . . . . . + . . + . . . . . . . . . . + + . + + . . + . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p f r os en ia o pp os iti fo lia . . . + + 1 + + b . . . + + a 1 . . + . . + + 1 1 1 . + + + + + a . 3 . a 3 . . a + 3 3 . . 1 . . . . . . . . . . . . . . . . . . . . + . . . + . . . . . . . . . . . . + . . . . . . . + . . . . . . . . . . . . . . p te ro ni a gl om er at a . . . + + + . + + + . + + + . + . + . . + + . + + + . . + . + + . . + 1 a + . + . . . + + + + + . . . . + . . . . . . . . . . . . + . + . . + . . . . . . . . . . . . . + . . . + . . . + + + . . . . . . . . . . . . ly ci um s pp . . + + + . + + + . + + + . . + + + . + + . . + . . + . . . . + r . . . . + . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . k ar ro oc hl oa s ch is m oi de s . . . + + + . . . . . . . . . . + . + . . . . . + + + . 1 + . . . . . + . + + + . + + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . + . . . . . . . . . e ri oc ep ha lu s de cu ss at us . . . . . . b . . . . . . . . . . . + + . + . . . . + + . . . . . . . . . . . . . . . . + . . + . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p en tz ia c f. sp ha er oc ep ha la . . . . . . . . . . . + . . + . . . . . . + . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r us ch ia c ra do ck en si s . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . 1 . . . . . . . . . . 1 . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p g g al en ia a fr ic an a . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . r . r . + r + + . . . . . . . . . . . . . . . . + . . . + . . . . . . + . . + + . . . . . . . . + . . . r . . . . . m ed ic ag o sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . a + + . . . . . . . . . + . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p h o ed er a ge ni st ifo lia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . 1 . . + + . 1 + . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . + + . . . . . . . . . . u rs in ia p ili fe ra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . + + r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r us ch ia in tr ic at a . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . a . + . . . . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . + . . . . . . . u rs in ia c al en du lifl or a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . 1 . . . + + . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p i p la nt ag o ca fr a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . p oa b ul bo sa . . . . . . . . . . . + . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . + + + . . . . + . . + . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . g al en ia s ar co ph yl la . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . c ro m id on v ar ic al yx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . + . . . . . . . . . . . . . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e ri oc ep ha lu s pa ni cu la tu s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . + . . . . . . . . . . . + . . . . . . . . . . . . . . . lo to no ni s hi rs ut a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p j le ys er a gn ap ha lo de s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r + . . . . + . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . + . . s el ag o sp . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . + + + . . . . . . . . . . . . . + + . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ta b l e 1 ph yt os oc io lo gi ca l t ab le o f t he p re do m in an tly m ou nt ai n r en os te rv el d ve ge ta ti on o f t he h an ta m -t an qu ar og ge ve ld s ub re gi on 64 fynbos biome related vegetation original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 61 71 r e l e v é n u m b e r 3 3 3 3 3 3 3 3 3 3 3 1 2 3 2 3 1 2 1 2 2 2 3 1 3 3 2 3 3 3 3 3 3 2 2 3 2 3 3 3 3 2 2 2 1 1 1 1 1 1 1 2 3 3 1 3 3 3 2 2 2 2 3 2 3 1 2 1 1 3 1 1 1 1 1 3 3 3 3 2 3 3 1 1 1 3 3 1 1 3 3 3 3 1 1 1 2 3 1 1 1 1 1 2 4 6 4 9 0 4 8 3 8 7 7 4 5 0 0 7 0 3 3 2 2 2 8 7 3 9 2 2 3 4 7 8 8 9 7 9 4 3 9 5 3 9 2 3 3 3 4 4 3 8 3 3 6 6 7 8 5 8 8 3 8 3 2 5 2 8 8 8 8 4 4 6 2 0 4 5 8 9 6 6 6 3 7 4 4 5 9 9 4 6 0 9 6 7 6 7 3 4 5 1 2 0 1 2 6 7 0 5 9 1 2 7 4 4 8 3 1 5 3 0 5 6 4 9 0 3 3 4 9 2 2 5 1 5 6 6 2 3 5 8 5 2 9 6 1 6 3 2 8 5 4 0 3 4 5 1 2 4 9 3 5 5 9 2 1 6 7 0 8 9 2 2 7 5 1 3 7 9 4 4 0 8 5 2 3 1 2 7 4 9 8 6 4 4 2 0 6 1 0 8 0 9 1 1. 1 1. 2 1. 3 1 .4 1. 5 2 .1 .1 2. 1. 2 2 .1 .3 2 .1 .4 2 .2 2. 3 3 tr ib ol iu m h is pi du m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . + . . + . . . . + . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . a lb uc a sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e hr ha rt a ca ly ci na . . . . . . . . . . . . + . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p k c ot ul a nu di ca ul is . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . + . . . + . + . . + . . + . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . p ol yc ar en a au re a . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . + + . + . . . . + + . . . . . + . + . . + . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . n em es ia s pp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . + . . . . + . . . . . . . . . + . . . . . . . r . . . . . . . . . . . . . . . . . . . . p ha rn ac eu m a ur an tiu m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . + . . . . . . . + . . . + . . . . . . . . + . . . . . . . . . . . . . + . . . . . . . . . . . . . b er kh ey a sp . . . . . . . . . . . . . . . + . . . . + . . . . . . . . . . . . . . . . . . + + . . . . . . . + . . . . + . . . . . . + . . . + + . + . + + + . . . . . . . . . . + . . . r . . . . . . . . . . + . . . . . . . . . . s p e c ie s g r o u p l e ro di um c ic ut ar iu m . . . . . . . . . . . + . . . . . + . + . . . . . . + . . + . . . . . . . 1 3 . . . . + + . . . . . . . . + + . . . . + + + + + + + . . . + . + . + + + . . . . . . . + . . . . . . . . . . . . . . . . . . . . + + . b ro m us p ec tin at us . . . . . . . . . . . + . + . . . . . . + . . + . . . . . . . . . . + . + + . . . + . . . . . . + + + . . + . + + . . . . . . . + . . + . . . . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s en ec io c ak ile fo liu s . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . + . + + . + . + . . . + . + . + + . + . . . . + + + + . . + + + + . + . . + . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . + + . f el ic ia a us tr al is . . . . . . . . . . . . . + . . . + . + . . . . . + . . . . . . . . + . . . + + + . . . . . . . . . + . . . . . + . + + . . + . + . . + . + . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . h el io ph ila s pp . . . . . . . . . . . . . . + . + . . . . . . . . . . . . . . . . + . . . . . . . + . + . . . . . . . . . . . . . . . . . . . . + + . . . + . + . + . + . . . . . . . + . . . . . . . . . . . . . . . . . . . . . + + . a rc to th ec a ca le nd ul a . . . . . . . . . . . . . . . . + . . + . . . . . . + . . . . . . + . . . . . . . . + . . + . . . . . . . . . + . . . . . . . . . + . . . . . r . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h el ic hr ys um lu ci lio id es . . . . + . . . . . . . + . . + . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . + . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . e ri oc ep ha lu s pa up er ri m us . . . . . . . . . . . . . . a 1 . . . . . . . . . . . . . . . . . . . . + . . . . . . . . b . . . . . . . . . . . . . . . . . . . 1 . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . le ys er a te ne lla . . . . . . . . . . . . . . . . + . . . . . . . . . + . . + . . . + . . . . . . . . . . . . + . 1 . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . g az an ia r ig id a . . . . + . . . . . . . . . . . . . . . . . + . + . + . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . + . . . . + . . . . . . . . . . . . + . a de no gr am m a gl om er at a . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . + . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . s p e c ie s g r o u p m h er m an ni a cu ne ifo lia . . . . . . . . . . . . . . . . . . r . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . + . . . . . . . . . . . r . + . . . + + . r . . . . . . . . . . . . . . . . . . . . . . d or ot he an th us s p. . . . . . . . . . . . . . . . . + . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . + . . . + . . . . . . . . . . . . . . . . . . . . + . . . + + . . + . + . . . . . . . . . . . . . + . . . . . . . . . h el ic hr ys um h am ul os um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + + . . r . . . . . . . . . . . . . . . . . . . . . o ed er a se di fo lia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . tr ip te ri s ag hi lla na . . . . . . . . . . . . . . . + . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . + . + . . . . . . . . . . . + . . . . . . . u rs in ia n an a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . + . . . . + . . . . . . . . . . . . + . . . . . + . + . . . . . . . . . . . . . . . . . . . . c yp er ac ea e sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . + . . . . . . . . . . . . . . . . . . . . + . . . te tr ag on ia m ic ro pt er a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . + . . . . . . . . . . . . . . . . . . . . . . . + . . . . . + . . . . . . . . . . . . . . . . . . . . . . . ty le co do n w al lic hi i . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . + . . . . . . . . . . . . . . . . . . . . d or ot he an th us m au gh an ii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . s cl er an th us a nn uu s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . + . + . . . . . . . . . + . + . + . + . . . . . . . . . . . . . . . . . . . . . . + . p el ar go ni um g ro ss ul ar io id es . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . + . . r . . . . . . . . . . . . . . . . . . . + . f es tu ca s ca br a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . r . . . . r . . . + . . . . . . . . . . . . . . . + . s p e c ie s g r o u p n f el ic ia fi lif ol ia . . . . . . r . . a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . + + . 1 r . . + + + + + + . . + . . . . . . . . . . . p ol yg al a sc ab ra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . r . . . + . . . . . . . . . . . . . + . . . . . . . . r . . + + . . . . . . . . . . . . . . . . e hr ha rt a m el ic oi de s . . . + r . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . + . . r . . . + + . + + . + . . . + . . . . . + . s p e c ie s g r o u p o p te ro ni a gl au ca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . 1 . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . + . + + . + b . . . . . . h el ic hr ys um a sp er um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . + . + . . . . . . . . a nd ro cy m bi um s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . + . . . . . . + s ta ch ys r ug os a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . + . . . . . . . . . . le ip ol dt ia ( h r p3 47 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . s p e c ie s g r o u p p p as se ri na tr un ca ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b + + + c lif fo rt ia a rb or ea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + a ga th os m a sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + o th on na a ur ic ul ifo lia . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . r . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + r . h es pe ra nt ha s p. . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . ta bl e 1 (c on t.. .) 65 original research van der merwe, van der merwe & van rooyen koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 61 71 r e l e v é n u m b e r 3 3 3 3 3 3 3 3 3 3 3 1 2 3 2 3 1 2 1 2 2 2 3 1 3 3 2 3 3 3 3 3 3 2 2 3 2 3 3 3 3 2 2 2 1 1 1 1 1 1 1 2 3 3 1 3 3 3 2 2 2 2 3 2 3 1 2 1 1 3 1 1 1 1 1 3 3 3 3 2 3 3 1 1 1 3 3 1 1 3 3 3 3 1 1 1 2 3 1 1 1 1 1 2 4 6 4 9 0 4 8 3 8 7 7 4 5 0 0 7 0 3 3 2 2 2 8 7 3 9 2 2 3 4 7 8 8 9 7 9 4 3 9 5 3 9 2 3 3 3 4 4 3 8 3 3 6 6 7 8 5 8 8 3 8 3 2 5 2 8 8 8 8 4 4 6 2 0 4 5 8 9 6 6 6 3 7 4 4 5 9 9 4 6 0 9 6 7 6 7 3 4 5 1 2 0 1 2 6 7 0 5 9 1 2 7 4 4 8 3 1 5 3 0 5 6 4 9 0 3 3 4 9 2 2 5 1 5 6 6 2 3 5 8 5 2 9 6 1 6 3 2 8 5 4 0 3 4 5 1 2 4 9 3 5 5 9 2 1 6 7 0 8 9 2 2 7 5 1 3 7 9 4 4 0 8 5 2 3 1 2 7 4 9 8 6 4 4 2 0 6 1 0 8 0 9 1 1. 1 1. 2 1. 3 1 .4 1. 5 2 .1 .1 2 .1 .2 2 .1 .3 2 .1 .4 2. 2. 2 .3 3 s p e c ie s g r o u p q d ic er ot ha m nu s rh in oc er ot is + 3 . . . . . . r + . . . . . . . . . . . . . . . . a . . . . . . . . . . . r . . + + . r . . . 1 . a + 1 . 4 a + a . a + 1 + + . . a r a 1 a + . 4 4 . + . b 1 + + a . 1 a . 1 b b 1 a 1 3 1 a . . b . . 4 . b + 3 . m er xm ue lle ra s tr ic ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . 3 . . . . + . . . . 1 1 + + 3 . a . . . . + . . 3 . + . . a 1 + + + + + . + a . . r 1 3 + . + . . a b . 4 + . a + . + . g eo ph yt ic s pp . . . . + . . . . . . . + . + + + . . . + . . . . . . . . . . . + . . . . . . . + + . + + + + + 1 + . . + . + + . + . + + + + . + + + + . . + + . + + . + . . + . + . + + . + . . . . + . . . . + + + . + . . + + + + . o xa lis s pp . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . + + + + + + + + + + + . + + . + + + + + + + + + + . . . . + + . + + + + + + + . + + . + . . . + . + . . . + + + . . + . . . + . d im or ph ot he ca c un ea ta . . . . . . . . . . . + . . . . . . . . + . . . . . . . . . . . . . . . . . . + . + + + . + . + . . . . + . . . . . . . + + . . + . a 1 + + + . + . . . 1 1 + + + r . . . + + . . . + . . . . . . . . . . . . + + . . h el io ph ila c ri th m ifo lia . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . + . . . . . . + . + . + + . . . + . . . . . . . . . . + . . . . . + . + . . . . . . . + + . + . . . + . . . . . . . . . . . . . . . . . . . . . . z al uz ia ns ky a be nt ha m ia na . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . + . . . . . . . . . . + . + . + . + . . . . . . . . . . + + . + . + . . . . . . . . . . . . . . . . . . . . . . . . g ni di a sc ab ra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a . . . + . + . . . . . . . + . + . . . . . . . . + . . . . . . . . . . . . . . . . . s p e c ie s g r o u p r c hr ys oc om a ci lia ta + . + . + + + + + + + . . . + . 1 + + . . + + . + + + a + + . . . a . + + . . . . + . + + + + . . . + . . + . + . . . . . . . + + . + a . . . + . . . . + + . + + + + . r + . . 1 . + + + . r . 1 . 1 . + r . . . r . a sp ar ag us c ap en si s + . . . . + + . . + . . + + + . . . . . . . + . . . . . . . . . . . . . . . . . + + . . + . + + + . . . . . . r + . . + . . . + + . . + . + . + . . . r . r . . . . . . r . . . . . + . . + r . + . . + r . . . . . . m or ae a sp p. . . . . . . . . . + . 1 + . + . + + + + + + . + . . + + . . . . . + + . + . . . . + + . + . + + . + . . . . + + . . + r . . + + + + + . . . . + . . . + + . . . . . . . . + . . . . + . . . . + . + + . + . . + . + . e ur yo ps la te ri flo ru s . . a 1 . . + . + . . . . . . . . 1 . + a . . . . . . . . . . . . . . . . . . a + 1 1 . . . . . . . . . . . . . . + + . 1 . . . . . + + 1 a + 1 . . . . 1 . + + 1 1 . . 1 . 1 . . . . 1 + + . + 1 . . + + 1 1 . . . . e ri oc ep ha lu s er ic oi de s . r . . + 1 . b . . . 1 a 1 . . + . . . . . a . . 1 . . . + a b . . . + . . . + . . . . . . . . . . . . . . . . . . . . . . a . + . . . . . . . . . . . . . + . . . . . + . + 1 . . . . + . 1 . . . . + . . . + + . . h el ic hr ys um o bt us um . . . . . . . . . . . . . . . . + + . . . . . . . . + . r + . . . . . . . . . . . . . . . . + + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . + . . ta bl e 1 (c on t.. .) * n on -d ia gn os tic s pe ci es a re e xc lu de d * h r c ol le ct io n co de a nd n um be rs a re in cl ud ed fo r fu tu re r ef er en ce , i f n ec es sa ry * s pe ci m en s in ad eq ua te fo r id en tifi ca tio n ye t d iff er en t f ro m s pe ci es th at c ou ld b e id en tifi ed a re in di ca te d w ith a p lo t n um be r (p … ) * sp . = o ne s pe ci es in a ta xo no m ic g ro up * sp p. = m or e th an o ne s pe ci es in a ta xo no m ic g ro up . t he se s pe ci es , e ve n al th ou gh th ey a re g ro up ed to ge th er , a re in cl ud ed in th e ta bl e si nc e th ey o cc ur in d iff er en t s pe ci es g ro up s. h ow ev er , t he y ar e no t u se d in th e de sc ri pt io ns in th e te xt . 66 using the distribution of the sample plots, supported by 1:250 000 topocadastral maps, land type maps (agricultural research council 1986a, 1986b, 1995, 1999a, 1999b, 2002, 2003), geology maps (council for geoscience 1973, 1983, 1989, 1991, 1997, 2001) and satellite images, the stratified units were assigned to a vegetation unit in the floristic classification. where two or more subassociations were present in a mapping unit, but it was not possible to map them separately as a result of high spatial diversity, they were mapped as mosaics of specific vegetation types. species that were unidentifiable during the field surveys were collected and the herbarium specimens sent to the compton herbarium, kirstenbosch, for identification. the collection code (hr) and numbers of the specimens were kept throughout the process as not all the species, especially within the mesembryanthemaceae, have yet been positively identified. all voucher specimens are lodged at the schweickerdt herbarium (pru), university of pretoria, pretoria. nomenclature follows that of germishuizen and meyer (2003). results the floristic data analysis resulted in two phytosociological tables. the first table (table 1) contains the predominantly mountain renosterveld veld type, as defined by acocks (1988), or the escarpment mountain renosterveld vegetation type, as defined by low and rebelo (1998), and is described in this article. three associations were identified, which were subdivided into nine subassociations, one of which contains four variants, as set out in the following scheme: 1. rosenia oppositifolia mountain renosterveld 1.1 eriocephalus microphyllus – rosenia oppositifolia mountain renosterveld 1.2 antimima cf. granitica (hr248) – rosenia oppositifolia mountain renosterveld 1.3 pentzia incana – rosenia oppositifolia mountain renosterveld 1.4 euryops multifidus – rosenia oppositifolia mountain renosterveld 1.5 pteronia glomerata – rosenia oppositifolia mountain renosterveld 2. dicerothamnus rhinocerotis mountain renosterveld 2.1 erodium cicutarium – dicerothamnus rhinocerotis mountain renosterveld 2.1.1 galenia africana – dicerothamnus rhinocerotis mountain renosterveld 2.1.2 oedera genistifolia – dicerothamnus rhinocerotis mountain renosterveld 2.1.3 senecio cakilefolius – dicerothamnus rhinocerotis mountain renosterveld 2.1.4 euryops lateriflorus – dicerothamnus rhinocerotis mountain renosterveld 2.2 dimorphotheca cuneata – dicerothamnus rhinocerotis mountain renosterveld 2.3 merxmuellera stricta – dicerothamnus rhinocerotis mountain renosterveld 3. passerina truncata mountain renosterveld with the exception of unit 1.2, all vegetation units could be mapped (fig. 2). three additional mosaics were also mapped: the nieuwoudtville mosaic, consisting of vegetation units 2.1.1, 2.1.4, 2.2 and 5.1 (van der merwe et al. in press); the soekop mosaic, consisting of units 2.1.3 and 2.2; and the welgemoed mosaic, consisting of units 1.2, 1.3 and 1.5. a large number of different land types are found in the study area and therefore only the predominant types are listed for each vegetation unit (agricultural research council 1986a, 1986b, 1995, 1999a, 1999b, 2002, 2003). table 2 summarises the most important features of the various land type symbols that have been used in the text. • • • fynbos biome related vegetation original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 61 71 1.3 pentzia incana – rosenia oppositifolia mountain renosterveld subassociation 1.3 is located around sutherland and east of sutherland on mudstones of the beaufort group. it also occurs further south in combination with subassociations 1.2 and 1.5 in the vicinity of the farm welgemoed, at the foot of the komsberg mountains. this subassociation, excluding the mosaic unit, covers an area of 44 499 ha (5.5% of the total area covered by mountain renosterveld vegetation). land types include fc and da and altitude ranges from approximately 1300–1500 m above sea level. the high-lying ridges on level terrain to moderate slopes are usually covered with brown or light brown sandy soils. the high shrub cover is attributed to species such as pentzia incana (species group d), rosenia oppositifolia and pteronia glomerata (species group f) as well as chrysocoma ciliata and euryops lateriflorus (species group r). this subassociation shows local variations resulting from a low constancy of such species as stipagrostis namaquensis, braunsia sp. and chrysanthemoides incana (species group c). when the perennial shrub cover is high, species in group c do not occur, however, when the shrub cover is lower, species in group c can dominate. generally, the cover of the grass and non-grassy herbaceous layer is limited, except in the case of the grass species stipagrostis namaquensis that occurred in a single relevé sampled in a drainage line. 1.4 euryops multifidus – rosenia oppositifolia mountain renosterveld located just north of the komsberg, subassociation 1.4 is found predominantly on mudstones of the beaufort group and covers an area of 106 189 ha (13.1% of the total area covered by mountain renosterveld vegetation). land types present include fc and da, with, occasionally, deep deposits of the ia land type. this subassociation is found at an altitude of between 1400–1500 m above sea level. the level to gently sloped ridges and light brown soils in this subassociation support a high shrub canopy cover of between 60 and 90%. the diagnostic species euryops multifidus and phyllobolus tenuiflorus (species group e), together with rosenia oppositifolia (species group f), chrysocoma ciliata and eriocephalus ericoides (species group r) characterise this subassociation. the cover of the herbaceous component (including grasses) is usually limited description of plant communities (table 1, fig. 2) the 2004 winter season was extremely dry, resulting in annuals and geophytes being poorly represented in the survey. the following description will therefore focus on the perennial plant species with permanent above-ground organs. 1. rosenia oppositifolia mountain renosterveld this plant association is located at the southern end of the roggeveld and nuweveld mountains as well as in the vicinity of the farms onderplaas and droëkloof further north and occurs predominantly on land types fc and da. mudstones of the beaufort group as well as dolerites are found underlying this association. the association is generally found on level terrain, gentle or moderate sloping ridges with a low rock cover from 0 to 10% or a high rock cover of 70 to 90%. brown or light brown sandy soils are prevalent in this high-lying plant association. although a high shrub cover is present, the grass and annual component only sometimes feature, usually with less than 5% cover. the vegetation of this association is characterised by species group f with a high cover of rosenia oppositifolia and includes species such as pteronia glomerata and karroochloa schismoides. common species include chrysocoma ciliata, euryops lateriflorus and eriocephalus ericoides (species group r). this association has been subdivided into five subassociations. 1.1 eriocephalus microphyllus – rosenia oppositifolia mountain renosterveld this unit is found in the region of the nuweveld mountains and covers an area of 106 454 ha (13.2% of the total area covered by mountain renosterveld vegetation). geologically, this subassociation is found on mudstones of the beaufort group and predominantly on land type fc, indicating that there is lime present in the entire landscape. this high-lying vegetation occurs at an altitude of > 1400 m above sea level on ridges with level terraces to gentle slopes. the rock cover varies from zero to > 85%, and is usually comprised of stones (> 50–200 mm) and boulders (> 200 mm). shrub cover is generally high (mean cover 66%) and is characterised by species such as rosenia oppositifolia and pteronia glomerata (species group f) as well as the diagnostic species eriocephalus microphyllus, pentzia sp. (hrp317) and euryops imbricatus (species group a). other shrubs also present include chrysocoma ciliata, euryops lateriflorus and eriocephalus ericoides of species group r. grasses are either absent or their cover is limited to less than 5%, while annuals are seldom present. this phenomenon could, however, be a result of the drought conditions experienced during the time in which the surveys were conducted. 1.2 antimima cf. granitica (hr248) – rosenia oppositifolia mountain renosterveld dolerite-derived b horizon soils on land types da and fc characterise this subassociation that is scattered throughout the roggeveld mountains and has not been mapped as a separate unit. the altitude generally ranges from 1200 to 1350 m above sea level. the high-lying ridges with level terrain to moderate slopes on brown to red brown sandy soils are usually covered with stones (> 50–200 mm in size). shrub cover in this unit is high (with a mean value of 72%), while the herbaceous component is generally < 5%. almost no grasses contribute to the herbaceous cover. the shrub layer is characterised by species such as rosenia oppositifolia, pteronia glomerata (species group f), eriocephalus pauperrimus (species group l) as well as asparagus capensis and eriocephalus ericoides (species group r). diagnostic perennial species include antimima cf. granitica (hr248) and pelargonium sp. (species group b). land type meaning of symbol d prismacutanic and/or pedocutanic diagnostic horizons dominate. after subtracting exposed rock, stones or boulders, more than half of the remaining land must consist of duplex soils. da refers to land where duplex soils with red b horizons comprise more than half of the area covered by duplex soils. db refers to land where duplex soils with non-red b horizons comprise more than half of the area covered by duplex soils. f glenrosa and/or mispah forms (though other soils may occur). the group accommodates pedologically young landscapes that are not predominantly rock, alluvial or aeolian and in which the dominant soil-forming processes have been rock weathering, the formation of orthic topsoil horizons and clay illuviation. fa refers to land in which lime is rare or absent from the entire landscape. fb indicates land where lime occurs regularly (though possibly in small quantities) in one or more valley bottom soils. fc refers to land where lime is generally present throughout the entire landscape. i miscellaneous land classes. ia refers to land types with a soil pattern difficult to accommodate elsewhere, at least 60% of which comprises pedologically youthful, deep (more than 1 m to underlying rock) unconsolidated deposits. ib indicates land types with exposed rock (country rock, stones or boulders) covering 60–80% of the area. table 2 land type symbols and their meaning within the text (du plessis 1987) 67 original research van der merwe, van der merwe & van rooyen koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 61 71 as well as euryops lateriflorus (species group r). common annual species occurring in the unit include erodium cicutarium, bromus pectinatus, senecio cakilefolius and felicia australis (species group l). this subassociation has been subdivided into four variants. 2.1.1 galenia africana – dicerothamnus rhinocerotis mountain renosterveld this variant is floristically very diverse and occurs on the mudstones of the beaufort group and the shales of the ecca group. it is located in the region of the farms m’vera and vondelingsfontein at the northern extreme of the roggeveld mountains and kareebos and rooiwal west of the klein roggeveld mountains. it also forms a mosaic in combination with variant 2.1.4, subassociation 2.2, and subassociation 5.1 (van der merwe et al. in press) in the nieuwoudtville area. excluding mosaics, this variant covers an area of 25 369 ha (3.1% of the total area of the mountain renosterveld vegetation). various land types are present, predominantly of the da and fb types. the altitude is notably lower than for the other vegetation units, and varies from 600 to 1300 m above sea level. this variant occurs on undulating terrain. the light brown to brown coloured sandy soils are usually not covered by much rock, however, boulders (> 200 mm) do occur locally. a high shrub cover results primarily from the presence of dicerothamnus rhinocerotis (species group q) as well as the diagnostic species galenia africana (species group g). various annual species, such as cotula nudicaulis and polycarena aurea (species group k) and erodium cicutarium, senecio cakilefolius, felicia australis and leysera tenella (species group l), are present. the annual grass bromus pectinatus (species group l) contributes to the low cover of the grass component in the variant. the absence of species group j in this variant differentiates it from variant 2.1.2. the presence of galenia africana and various annuals indicates the increased disturbance that has taken place in this variant in the past. 2.1.2. oedera genistifolia – dicerothamnus rhinocerotis mountain renosterveld variant 2.1.2 occurs on the mudstones of the beaufort group in the klein roggeveld mountains, and covers an area of 46 797 ha (5.8% of the total area of the mountain renosterveld vegetation). it is found at an altitude between 1000 and 1300 m above sea level on level terrain to gentle slopes. the light brown sandy soils of this variant are covered with gravel (< 10 mm), small stones (> 10–50 mm) and boulders (> 200 mm), which are typical of land type ib. the high shrub cover (more than 70%) is primarily a result of the presence of dicerothamnus rhinocerotis (species group q) as well as oedera genistifolia (species group h) and euryops lateriflorus (species group r). merxmuellera stricta (species group q), a perennial grass, dominates the grass component of this variant. annual species are consistently present, however, their cover is low due to the drought conditions in the year in which the surveys were conducted. the presence of species groups g and j distinguishes variant 2.1.1 from variant 2.1.2, whereas the absence of species group i distinguishes variant 2.1.2 from variant 2.1.3. 2.1.3 senecio cakilefolius – dicerothamnus rhinocerotis mountain renosterveld this variant, excluding mosaics, covers an area of 13 654 ha (1.7% of the total area covered by mountain renosterveld vegetation) and is found on mudstones of the beaufort group and shales of the ecca group in the region of the farms botuin, blomfontein and de hoop in the roggeveld mountains, predominantly on land types da and fc. in combination with subassociation 2.2 in the region of the farm soekop, it is found in a mosaic to < 5%, which could be the result of the drought conditions in the year of survey. 1.5 pteronia glomerata – rosenia oppositifolia mountain renosterveld geologically, this subassociation occurs predominantly on mudstones of the beaufort group and is similar to subassociations 1.3 and 1.4. it is found on land types fc, da and db on the southwestern extreme of the roggeveld mountains. it also occurs in a mosaic with subassociations 1.2 and 1.3 in the vicinity of the farm welgemoed at the foot of the komsberg mountains. this subassociation covers an area of 69 233 ha (8.6% of the total area of the mountain renosterveld vegetation), excluding the mosaic unit. this high-lying (1200–1600m above sea level) subassociation is found on level terrain and gentle slopes on a range of different rock sizes, varying from gravel (< 10 mm) to boulders (> 200 mm). the soil colour also varies substantially from light brown to brown to red brown. shrub cover is generally high (> 60%) and the grassy component has a higher presence and cover compared with the previous subassociations. likewise, annual species are present in all relevés, with their cover generally being higher than in the previous subassociations. this was probably the case due to the local rain showers received in the area during the year of survey. no diagnostic species group separates this subassociation. the most prominent species present include rosenia oppositifolia, pteronia glomerata (species group f), erodium cicutarium (species group l) and euryops lateriflorus (species group r). the grass component is represented by karroochloa schismoides (species group f) and bromus pectinatus (species group l) with, in one relevé, a high cover of merxmuellera stricta (species group q). erodium cicutarium and felicia australis (species group l) as well as heliophila crithmifolia (species group q) represent some of the annual species. 2. dicerothamnus rhinocerotis mountain renosterveld this plant association is located in the roggeveld, klein roggeveld, koedoesberg and komsberg mountains and has been further subdivided into three subassociations. generally it can be found on the mudstones of the beaufort group or the shales of the ecca group on land types da, fb, fc, ib and fa. the level terrain and gentle slopes of subassociations 2.1 and 2.2 as well as the gentle to moderate slopes of subassociation 2.3 are usually comprised of light brown or brown sandy soils. the shrub cover is high (50–95%) and a grass and annual component are generally present throughout the association. the very high cover of dicerothamnus rhinocerotis, merxmuellera stricta and dimorphotheca cuneata (species group q) distinguishes this association from the rosenia oppositifolia mountain renosterveld (association 1). 2.1 erodium cicutarium – dicerothamnus rhinocerotis mountain renosterveld subassociation 2.1 generally occurs on mudstones of the beaufort group in the roggeveld, klein roggeveld and komsberg mountains on land types da, fb, fc and ib, and excluding mosaic units, covers an area of 213 410 ha (26.4% of the total area of mountain renosterveld vegetation). the altitude varies from 600 to about 1600 m above sea level and the landscape is gently undulating. soils are light brown sandy soils with a varying rock cover consisting predominantly of boulders (> 200 mm). the shrub cover is generally high (50–95%), with grass and other herbaceous species consistently occurring across all the surveyed sites. prominent perennial species in this subassociation include dicerothamnus rhinocerotis, merxmuellera stricta (species group q) 68 fynbos biome related vegetation original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 61 71 vegetation unit. it occurs at altitudes higher than 1200 m above sea level on level terrain to gently sloping landscapes with light brown to brown coloured soils. rocks are mostly absent, although boulders do occasionally occur. shrub cover varies considerably, with the main contributors being dicerothamnus rhinocerotis and dimorphotheca cuneata (species group q) as well as chrysocoma ciliata, asparagus capensis, euryops lateriflorus and eriocephalus ericoides (species group r). the grass component varies considerably depending on the presence or absence of the perennial grass merxmuellera stricta (species group q). the most prominent annual species include cromidon varicalyx and plantago cafra (species group i), cotula nudicaulis and polycarena aurea (species group k) as well as erodium cicutarium and senecio cakilefolius (species group l). the cover of this component is highly variable, depending on the amount of rainfall received locally during the season. variant 2.1.3 has a close affinity with variant 2.1.2 due to their sharing species group j, however, they differ as a result of the presence of species group i that is confined to variant 2.1.3. 2.1.4 euryops lateriflorus – dicerothamnus rhinocerotis mountain renosterveld variant 2.1.4 occurs on land types da and fc in the komsberg mountains and southwest of the basterberg mountains and covers an area of 127 590 ha (15.8% of the total mountain renosterveld vegetation), excluding the mosaic vegetation unit. the mosaic is found in combination with variant 2.1.1, subassociation 2.2, and subassociation 5.1 in the nieuwoudtville area (van der merwe et al. in press). this variant is generally found at high altitudes on level terrain to gentle slopes. the light brown soils are derived from mudstones of the beaufort group. rocks are generally absent, although boulders (> 200 mm) may occur locally. the high shrub cover is due primarily to dicerothamnus rhinocerotis and dimorphotheca cuneata (species group q) as well as chrysocoma ciliata, asparagus capensis and euryops lateriflorus (species group r). the grass cover is generally low, except where merxmuellera stricta (species group q) dominates. the cover of the annual component is generally low. two forms of variant 2.1.4 occur as a result of the presence or absence of species group k, which mainly consists of annual species. such species might have occurred throughout the region in a normal rainfall year. 2.2 dimorphotheca cuneata – dicerothamnus rhinocerotis mountain renosterveld this high-lying subassociation can be found in the keiskie mountains, at the northern extreme of the roggeveld mountains, and excluding mosaics, covers an area of 20 196 ha (2.5% of the total area covered by mountain renosterveld vegetation). it also occurs in combination with variant 2.1.3 in the region of the farm soekop and additionally, it forms a mosaic in the nieuwoudtville area in combination with variants 2.1.1 and 2.1.4 and subassociation 5.1 (van der merwe et al. in press). the land types present include da, fa and fc and the altitude varies from 700–1400 m above sea level. the undulating terrain is usually covered by a high percentage of boulders (>200 mm). the light brown to brown coloured sandy soils are derived from ecca shales. the shrub cover varies greatly (20–80%), whereas the cover of both the grass and annual species remains low. diagnostic species include hermannia cuneifolia, helichrysum hamulosum and oedera sedifolia (species group m). felicia filifolia, polygala scabra and ehrharta melicoides (species group n) are common to both subassociation 2.2 and 2.3, although subassociation 2.3 lacks species group m. the dominant species is dicerothamnus rhinocerotis and other prominent species include merxmuellera stricta, dimorphotheca cuneata (species group q) and chrysocoma ciliata, euryops lateriflorus and eriocephalus ericoides (species group r). 2.3 merxmuellera stricta – dicerothamnus rhinocerotis mountain renosterveld this subassociation is located in the region of the farms piet se nuplaas, droëberg, nuwepos, soekop and vaalhoek in the roggeveld mountains and includes the higher-lying vegetation of the koedoesberg and basterberg mountains. it covers an area of 230 838 ha (28.5% of the total mountain renosterveld vegetation). geologically, it occurs on the mudstones of the beaufort group, the shales of the ecca group and even, occasionally, on dolerites occurring within the mudstones and shales. land types include fc, da and occasionally ib at an altitude of 900 to 1600 m above sea level. the high-lying gentle to moderately steep slopes are usually covered with stones (> 50–200 mm) or boulders (> 200 mm). the soils are generally brown sandy soils. shrub and grass cover vary considerably, whereas the annual component is either absent or covers less than 1% of the area. three variations are distinguished in this subassociation. the first variation is differentiated by the presence of species group n, which is shared with subassociation 2.2. the second variation is characterised by the perennial shrub pteronia glauca (species group o), whereas the third variation does not include species group n or o. in all of these variations dicerothamnus rhinocerotis, merxmuellera stricta (species group q) and chrysocoma ciliata (species group r) dominate with a very high cover (60–95%). other species present include asparagus capensis, euryops lateriflorus and eriocephalus ericoides (species group r). 3. passerina truncata mountain renosterveld the third plant association, which is found exclusively on dolerites on land type ia, occurs at high altitudes (approximately 1500 m above sea level and higher) on the hantam mountain as well as at various locations scattered throughout high-lying areas in the roggeveld mountains. it covers an approximate area of 17 982 ha (2.2% of the total area of the mountain renosterveld vegetation). the high-lying terraces and plateaux consist of red brown sandy clay soils, with the rock cover varying from 20– 80%. the shrub cover is very high, except where a high cover of exposed rocks occurs. compared to the high shrub cover, the cover of the grass and annual species is generally very low. this association is differentiated by species group p, which includes diagnostic species such as passerina truncata and othonna auriculifolia. other common species present include dicerothamnus rhinocerotis, merxmuellera stricta and dimorphotheca cuneata (species group q) and eriocephalus ericoides (species group r). discussion according to rutherford and westfall (1986), low and rebelo (1998) and mucina et al. (2005) the vegetation of the subregion, as discussed in the present article, is situated predominantly in the fynbos biome. however, diels (1909 in van wyk & smith 2001) concurred with marloth (1908) that the region is floristically more closely related to the succulent karoo than to the cape floristic region. this area was also included in the skep initiative and not in the cape (cape action plan for the environment) initiative. the clear split between table 1 and the table presented on the succulent karoo related vegetation (van der merwe et al. in press) reveals that most of the species in species group f (table 1) 69 original research van der merwe, van der merwe & van rooyen koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 61 71 are found in the general species group in the upper portion of the succulent karoo table, whereas most of the species in species group q (table 1) are not found in the succulent karoo table. such a finding indicates association 1’s affinity with the succulent karoo biome vegetation of the escarpment karoo, roggeveld karoo and hantam karoo, as described in van der merwe et al. (in press). the true renosterveld of associations 2 and 3, as defined by species group q (table 1), is, however, lacking from the succulent karoo table and belongs to the fynbos biome related vegetation. this mountain renosterveld is probably distinct from other renosterveld vegetation types in any case and could be studied in the future. grazing and cropping are the main land-uses in the mountain renosterveld. sustainable land management tries to minimise the risk of veld degradation or species extinction by managing populations of plants and animals within an area to ensure that they can continue to reproduce and function normally, even after stressful conditions such as drought (esler et al. 2006). although damage can happen fast, recovery in the karoo is very slow, as it depends mainly upon unpredictable rainfall events (esler et al. 2006). sustainable farm management planning is critical for ensuring a productive, profitable future in the region. inadequate farming practices, resulting from a severe lack of infrastructure, especially fencing, pose a serious threat to the vegetation. farms in the region yield a low income as a result of the harsh environmental conditions and the unpalatable grazing caused by the dominance of dicerothamnus rhinocerotis. because of the low monetary value of the land and the high cost of infrastructure it is not financially viable for a farmer to invest too much in infrastructure, as the ability to recover such costs is limited. although the farmers are generally willing to implement improved veld management and infrastructure development, their financial means hinder their doing so. according to low and rebelo (1996) the degree of transformation in the escarpment mountain renosterveld, which closely corresponds to the mountain renosterveld as described in the current article, is unknown. however, many large tracts of land cultivated in the past are still cultivated due to the higher rainfall in the region compared with that experienced in the surrounding areas of the hantam and tanqua karoo. invasive species in the vegetation type described are predominantly annuals that were brought into the region with fodder from other parts of the world, and of which many have been naturalised over the centuries. the isolated individuals of prosopis species present are usually limited to highly disturbed areas alongside water points and feeding areas. the unpalatable renosterbos, dicerothamnus rhinocerotis, which dominates large sections of the mountain renosterveld is considered an encroacher by most farmers with its dominance being blamed on centuries of incorrect management practices in the region. also, overgrazing is thought to have substantially reduced the grassy component in the vegetation. the protected area network for the mountain renosterveld is severely under-represented. two local municipal nature reserves, namely the nieuwoudtville wildflower reserve (115 ha) and the akkerendam nature reserve (230 ha), fall within the region. a natural heritage site at banksgate, near sutherland, protects the rare sterboom, cliffortia arborea. the tanqua national park has substantially expanded during the last 3–5 years, with the latest land acquisitions including a section of mountain renosterveld vegetation. in conclusion, the aims of the project described in this article were to classify and describe the various vegetation units present in the mountain renosterveld part of the hantam-tanquaroggeveld subregion in terms of their species composition, environmental parameters and relationships to one another as well as to map their geographical distribution. such an inventory of vegetation types should aid future planning, resource management and biodiversity conservation, which should encourage sustainable land use practices, reducing the negative impact on the environment. acknowledgements the authors would like to thank the critical ecosystem partnership fund (cepf) for funding the project by way of the skep (succulent karoo ecosystem plan/program) initiative. the critical ecosystem partnership fund is a joint initiative of conservation international, the global environmental facility, the government of japan, the macarthur foundation and the world bank. a fundamental goal is to ensure that civil society is engaged in biodiversity conservation. the assistance of hennie van den berg of iris international in compiling the vegetation map is gratefully acknowledged. capenature, the department of tourism, environment and conservation (northern cape) and sanparks are thanked for the necessary permits and permission to conduct this research. references acocks, j.p.h. 1988. veld types of south africa. 3rd ed. memoirs of the botanical survey of south africa, 57: 1–146. agricultural research council 1986a. land type map 3220 sutherland. pretoria: institute for soil, climate and water. agricultural research council 1986b. land type map 3018 loeriesfontein. pretoria: institute for soil, climate and water. agricultural research council 1995. land type map 3118 calvinia. pretoria: institute for soil, climate and water. agricultural research council 1999a. land type map 3120 williston. pretoria: institute for soil, climate and water. agricultural research council 1999b. land type map 3218 clanwilliam. pretoria: institute for soil, climate and water. agricultural research council 2002. land type map 3319 worcester. pretoria: institute for soil, climate and water. agricultural research council 2003. land type map 3320 ladismith. pretoria: institute for soil, climate and water. cepf, 2003. ecosystem profile: the succulent karoo hotspot, namibia and south africa. critical ecosystem partnership fund report. conservation international: http://www.biodiversityhotspots. org (accessed july 2006). council for geoscience 1973. geological map 3218 clanwillliam. pretoria: council for geoscience. council for geoscience 1983. geological map 3220 sutherland. pretoria: council for geoscience. council for geoscience 1989. geological map 3120 williston. pretoria: council for geoscience. council for geoscience 1991. geological map 3220 ladismith. pretoria: council for geoscience. council for geoscience 1997. geological map 3319 worcester. pretoria: council for geoscience. council for geoscience 2001. geological map 3118 calvinia. pretoria: council for geoscience. du plessis, h.m. 1987. land types of the maps 2816 alexander bay, 2818 warmbad, 2916 springbok, 2918 pofadder, 3017 garies, 3018 loeriesfontein. memoirs on the agricultural natural resources of south africa, 9: 1–538. esler, k.j., milton, s.j. & dean, w.r.j. 2006. karoo veld ecology and management. pretoria: briza publications. germishuizen, g. & meyer, n.l. (eds). 2003. plants of southern africa: an annotated checklist. strelitzia, 14. pretoria: national botanical institute. 70 fynbos biome related vegetation original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 61 71 myers, n., mittermeir, r.a., mittermeir, c.g., de fonseca, g.a.b. & kent, j. 2000. biodiversity hotspots for conservation priorities. nature, 403: 853–858. nelder v.j., wilson, b.a., thompson, e.j. & dillewaard, h.a. 2005. methodology for survey and mapping of regional ecosystems and vegetation communities in queensland. version 3.1. updated september 2005. brisbane: queensland herbarium, environmental protection agency. rutherford, m.c. & westfall, r.h. 1986. biomes of southern africa. an objective characterisation. memoirs of the botanical survey of south africa, 54: 1–98. van der merwe, h., van rooyen, m.w. & van rooyen, n. in press. vegetation of the hantam-tanqua-roggeveld subregion, south africa. part 2. succulent karoo biome related vegetation. koedoe. van wyk, a.e. & smith, g.f . 2001. regions of floristic endemism in southern africa: a review with emphasis on succulents. pretoria: umdaus press. weather bureau 1998. climate of south africa: climate statistics up to 1990. wb 42. pretoria: government printer. werger, m.j.a. 1974. on concepts and techniques applied in the zürich-montpellier method of vegetation survey. bothalia, 11(3): 309–323. hennekens, s.m. & schaminée, j.h.j. 2001. turboveg, a comprehensive database management system for vegetation data. journal of vegetation science, 12: 589–591. hill, m.o. 1979. twinspan – a fortran program for arranging multivariate data in an ordered two-way table by classification of the individuals and attributes. ithaca: ecology & systematics, cornell university. hilton-taylor, c. 1994. western cape domain (succulent karoo). in davis, s.d., heywood, v.h. & hamilton, a.c. (eds), centres of plant diversity: a guide and strategy for their conservation, 1. cambridge: iucn publications unit. pp. 201– 203. low, a.b. & rebelo, a.g. 1998. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. marloth, r. 1908. das kapland, insonderheit das reich der kapflora, das waldgebiet und die karoo, pflanzengeografisch dargestellt. wiss. ergebn. deutsch. tiefsee-exped. ‘waldivia’, 1898 – 1899. 2, t. 3, jena: fischer. mucina, l., rutherford, m.c. & powrie, l.w. (eds). 2005. vegetation map of south africa, lesotho and swaziland, 1 : 1 000 000 scale sheet maps. pretoria: south african biodiversity institute. 71 filelist convert a pdf file! koedoe 19: 17-26 (1976) geographical distribution and population structure of springbok antidorcas marsupialis rumen protozoa in southern africa r. c. wilkinson and w. van hoven mammal research institute department ojzoology university oj pretoria pretoria 0002 abstract springbok were sampled in angola, south west africa, kalahari gemsbok national park, the cape province and eastern transvaal between july 1973 and july 1974. all the ciliates found in the 20 rumens investigated belonged to a single genus entodinium (family ophryoscolecidae) and they varied in number from 0,3 x 106 to 2,5 x 106 per ems of rumen fluid. the different species were found and a positive correlation is seen between total numbers and protein content of the feed. it has been concluded that the numbers of e. parvum are closely related to the protein concentration of the feed. introduction very little information is available on the ciliate fauna of wild ruminants residing in southern africa. since nothing is known in this respect from the springbok a series of rumen samples were collected in angola, south west africa, the kalahari gemsbok national park, the cape province and eastern transvaal betweenjuly 1973 andjuly 1974. the intention of this investigation was to obtain quantitative and qualitative (taxonomic) information about the rumen ciliate fauna of springbok living under natural conditions. the taxonomic results and descriptions of two new species has been submitted for publication separately. all the ciliates found in the 20 rumens investigated from the different geographical areas, belonged to a single genus entodinium (family ophryoscolecidae). this is unique since the only other ruminants that have ever been found to show an entodinium only rumen fauna, are the alaskan moose alces americana (dehority 1974) and the whitetailed deer of texas odocoileus virginianus (pearson 1965). the situation in the moose is uncertain since dogiel (927) found three other genera in its rumen as well. ruminants in general usually contain 8-12 genera 17 and a large number of different species that vary from simple types such as the entodinia to large anatomically complicated specimens such as has been found in the tsessebe damaliscus lunatus lunatus from northern transvaal (van hoven 1975). the 10 different species are listed in table 2. the two species entodinium kalaharicus and e. lucii were found for the first time in the springbok whilst e. jyferi (van hoven 1975) was described from the south african tsessebe. an attempt is further made to relate the ciliate data to the nutritional composition of the feed, the geographical distribution and other facts known about the ecology of the host animal. m alerial and m elhods the methods of collecting the material and the staining of the protozoa is described in van hoven (1974). extraction and counting of the ciliates was done by first washing the rumen fluid samples through fine-meshed sieves of 16 and 44 cm' with 100 cm 3 of water whilst being agitated with a fine paint brush. the resultant f1uid was thoroughly shaken up and 1 cm 2 of it was diluted with 10 cm 3 of water. to revent rapid settling of the organisms and to facilitate even dispersal in the counting chamber 1,25 cm3 of glycerine was added. this mixture was well shaken up and a small quantity of it placed in a mcmaster counting chamber to determine the total number of ciliates present. counts were done at 160 x magnification, four from each sample and the average of these was used in determining the numbers per cubic centimeter. for population structure determination random distribution on a slide was assumed because the sample of rumen f1uid was shaken up well immediately before a few drops were spread on the surface. part of the solid digesta was collected and sun dried. these were later analysed for starch, nitrogen, protein, fibre and ash. hosts and regions sampled: i . samples i, 2 and 3 were obtained from springbok shot in the district of karasburg in southern south west africa on 1973.06.08 and 1973.06.09. the vegetation in this area is desert-like with scattered clumps of grass and shrubs. 2. samples 4, 5 and 6 were obtained from springbok shot in the mocamedes district of angola on 1973.07.03. the vegetation is similar to the above. 3. samples 7 to 16 were obtained from springbok shot in the kalahari gemsbok national park on 1974.04.16. the vegetation is savanna in the cases of animals 7 to 12 and open grassland for 13 to 16. 18 4. samples 17, 18 and 19 were obtained from the ermelo district of the transvaal. the animals were shot on 1974.06.16. the vegetation in this area is open grassland. 5. sample 20 was from a springbok shot near graaff reinet in the cape province on 1974 .07 . 17 . results total numbers a considerable amount of variation was evident between the samples with regard to the total number of ciliate protozoa per cm 3 original rumen fluid counted. the number of cells varied from 301 300 in sample 19 to 2425 100 in sample 12. table 1 total number oj ciliate protoz.oa jound in the springbok jrom the different areas sampled area sample number total count/ml karasburg, s.w.a . 1 1 501 920 2 1 118 820 3 1 256 220 mocamedes, angola 4 1 891 080 5 5836800 6 1 791 240 kalahari gemsbok 7 1 171 620 national park, 8 1 094580 r.s.a . 9 2 138 160 10 1 860480 11 1 683 120 12 2425080 13 2 172 120 14 9 672 600 15 1 949400 16 1 625 940 errnelo, r.s .a. 1 7 452280 18 960 120 19 3 01 380 graaff reinet 20 2429040 19 there is evidence to prove that the numbers of cells in the rumen vary significantly with the seasons (westerling 1970; pearson 1965). this would, however, not appear to be the reason for the above mentioned discrepancy as these two samples were obtained within a month of each other. food composition as a factor influencing cell numbers has been found to be very important by a number of workers (westerling 1970). this is a more feasible explanation for the variations found in this study because the above extremes are samples from different areas having different vegetation types. there is a similarity in total numbers between animals from the same area which is not necessarily evident between areas. all animals sampled from the ermelo district had considerable lower numbers of entodinia than those from other areas. this indication that areas, i.e. vegetation types, influence the protozoan population will be expanded on in the section on food composition . total counts for each animal sampled are given intable 1. population composition the population composition of ciliate protozoa found in ruminants varies considerably both on the interspecific and intraspecific levels. in all the springbok examined, cases were found of one or more species totally lacking or represented by only a few individuals. this is despite the fact that other animals from the same area may show large numbers of the same species in their rumen populations. according to westerling (1970) a ciliate species, which commonly occurs in the rumen of a certain host, may occasionally be lacking in some individuals in the herd. this phenomenon may depend on chance or on actual fluctuations in the numbers of the species involved. it can be assumed that competition or other ecological factors at times cause the disappearance of some species from the rumen fauna and that re-inoculation from other host individuals in the herd, or a change in the biotope before the species is exterminated, enable the species to reattain normal numbers. the interval can be looked on as a period when this particular species is not adapted to the biotope. the most important factor in the ciliate population structure in the rumen of the springbok is that, as was found in the white-tailed deer (pearson 1965) and the alaskan moose (dehority 1974), only the genus entodinium of the family ophryoscolecidae was represented. the percentage representation of the different species is given in table 2 and fig. 1. from this it can be seen that e. parvum is the dominant species in all the populations with the exception of two animals from the kalahari where e. kalaharicus n. sp. dominated (samples 11 and 13 lan interesting characteristic of these population structures is that, al20 fig. 1: population composition of the rumen ciliate fauna of each sample/ key: e. parvum e. dubardi e. alces e. kalaharicus e.caudatum elucii e.anteronucleatum e. fyferi e. bovis , e.longinucleatum fig. i. population composition of the rumen ciliate fauna of each sample. though some of the less numerous species are sometimes absent trom a sample, the composition of each population is relatively constant, regardless of geographical area. the different areas studied showed different degrees of domination by e. parvum and a variation in the species which are sub-dominate (table 3). 21 table 2 percentage of total ciliate population oj each entodinium as found in all the springbok sampled ciliate sample number ( entodinium) i 2 3 4 5 6 7 8 9 10 ii 12 13 14 15 e. parvum 54,9 54,5 63,3 70,0 68,3 55,2 30,0 47,5 53,7 15,2 40,6 23,3 52,8 46,3 e. dubardii dubardi 6,6 9,3 7,9 11,2 12,3 8,4 11,6 9,6 2,8 4,9 7,2 4,3 16, i 5,3 e. alces 20,9 12,6 13,8 2,8 3,7 12,6 9,4 9,6 6,9 3,2 1,4 0,4 2,3 \,7 e. kalaharicus 2,2 3,7 1,3 2,4 2,4 4,2 14,4 7,3 4,6 37,9 27,6 43,6 12, i 23,4 e. caudatumf dubardi 0,5 3,2 2,5 3,3 18,4 9,2 11,9 27,0 13,7 19,9 4, i 13,9 e.lucii 13,7 14,4 8,9 8,4 4,1 13,0 3,6 14,6 18,3 10,3 7,2 5,7 3,6 7,4 e. antero nucleatum 0,4 3,7 3,5 0,4 3,3 0,5 3,6 1,4 0,9 0,3 1,4 1,4 7,6 0,8 e·fyferi \,3 \,4 \,3 1,6 2,9 2,8 0,4 0,9 0,9 1,4 1,2 e. bovis 9,0 0,4 0,9 0,4 e . longinucleatum 0,3 16 17 18 34,0 65,6 73,9 6,2 2,8 1,2 16,3 21, i 17,3 5, i 0,4 30,2 2,8 0,4 6,2 3,7 0,4 3,3 1,4 0,4 \,2 2,3 3,4 0,4 19 74,9 19,8 0,4 0,4 4,5 20 60,9 5, i 11,2 8,8 4,7 3,7 1,9 3,7 c'-{ c'-{ table 3 dominance status of ciliate protozoa found in different areas status area angola karasburg kalahari ermelo graaff reinet dominant e. parvum e.parvum e.parvum e.parvum e.parvum e. kalaha ricus sube. dubardi e. dubardi e. dubardi e. alces e. dubardi dominant e. alces e. lucii e. alces e. alces e.lucii e. caudatum e. kalaharicus e.lucii e. caudatum present in e. kalahae. caudatum e·fyferi e. dubardi e. lucii low ncus numbers e. anteroe. kalahae. anteroe. kalahae. anteronucleatum ricus nucleatum ricus nucleatum e·fyferi e. antetoe. bov/.s e. caudatum e·fyferi nucleatum e. anteroe·fyferi nucleatum e. alces e·fyferi e. lucii absent e. caudatum e. bovis e. bovis e. bovis e. bovis food composition the composition of the food ingested by the host animal plays an important role in the population structure and numbers of the rumen ciliate fauna . diets rich in protein have been shown to increase the ciliate populations considerably. similarly entodinium becomes particularly numerous in ruminants fed large quantities of grain because of their ability to utilize starch (hungate 1966). according to westerling (1970), this is because entodinium is generally believed to specialize on starch for its supply of energy. however, hungate (1966) cites sugden (1953) as stating that some very small species of entodinium are not able to utilize starch to a significan degree, due to their inability to ingest the large starch grains. the ability of ciliates to ingest and digest cellulose has been shown (hungate 1966) but their importance in cellulose digestion is not known. hungate (1966) cites a number of authors who believe that this is principally the function of rumen bacteria. the entodiniomorphs generally appear to have a very limited ability to utilize soluble substrates (abou akkada and howard 1960). thus 23 sugars should have little effect on the population structure where only entodinia are present. preservation of rumen digesta in formalin does not affect the analyses of it significantly as the residue on ignition of a 35% formaldehyde solution consists of: sulphates 0,002% free acid 0,03% chlo"ride 0,0001% heavy metals 0,0005% iron 0,00015% this gives a total of 0,03675% inorganic substance which will be shown in the ash determination. this is, however, so small that it can be ignored. the results of the rumen digesta analyses found in this study are given in table 4. table 4 analyses of rumen digesta from all the springbok in each area given as a percentage of the total food composition f ood rotein p f a s ibre sh tarch discussion angola karasburg 14,30 16,00 25,33 34,88 15,98 17,82 0,84 0,85 area kalahari ermelo graaff reinet 16,22 13,31 15,71 34,89 29,43 29,61 16,07 17,59 10,55 0,75 0,61 0,79 the total numbers of rumen ciliates in a host show a good correlation with the protein content of the food. although the differences in protein content of samples from different areas were not great, the lower protein diet of the springbok from ermelo coincides with a marked drop in numbers. similarly, the highest protein content, found in the kalahari, coincides with the highest numbers. this concurs with the observations of earlier workers (hungate 1966; westerling 1970). the ash content, or inorganic component of the food shows a slight negative correlation with the numbers. this is most evident in ermelo and graaff reinet. fibre content does not appear to be correlated with numbers at all as it shows a positive correlation with the drop in numbers in ermelo but a negative correlation in angola. this is consistant with the idea 24 that the digestion of cellulose is principally the function of the rumen bacteria. with respect to this hungate (1966) states, "entodinium, with the possible exception of entodinium bursa, does not digest cellulose, but has been observed to ingest it." although some correlation has been made in the past between the food composition and the genera present, little or nothing is known about its connection with the species variation. it has, however, been evident in this study that there is a connection between species composition and protein availability. in the relatively low protein diet of the springbok e. parvum dominates all populations with sub-dominant species forming low percentages of the population. this is particularly evident in ermelo where the lowest protein values were found coinciding with an average e. parvum population of 71 per cent. in angola the protein value is slightly higher and e. parvum forms 64,5% of the population. in graaff reinet and karasburg which have respectively higher protein values 60% and 57% of the ciliate population's consisted of e. parvum. the highest protein values were found from the kalahari and here e. parvum performed about 38% of the population. population. this suggests that e. parvum is at an ecological advantage in a lowprotein environment while those species which were sub-dominant or only present in low numbers, began to compete successfully with it as protein availability increased. this possibility must at this stage remain speculative and requires further intensive study to be verified. both dogiel (1927) and westerling (1970) found that the ciliate fauna in the reindeer rangifer tarandus consists largely of species which are specific to this host. this species specificity is not evident in the springbok as most species have been identified in other ruminants. this is because there is a decisive dietical difference between the reindeers diet and that of other ruminants. the bulk of the reindeer diet is lichen, the carbohydrates of which are comparitively unique. the diet of the springbok, on the other hand, is of a far more general nature. acknowledgements we thank prof. j. d. skinner and mr. t. robinson for collecting some of the samples and the national parks board of trustees and the owners of private nature reserves for their kind cooperation in obtaining the material. references abou akkada, a. r. and b. h. howard 1960. the biochemistry of rumen protozoa, 3. the carbohydrate metabolism of entodinium. biochem.j. 76:445-551. dehority, b. a. 1974. rumen ciliate fauna of alaskan moose (alces americana), musk-ox (ovibos moschatus) and dall mountain sheep (ovis dalli).j. protol.ool. 21 :26-32. 25 dogiel, v. a. 1927. monographie der familie ophryoscolecidae. arch. protistenk. 59: 1-288. hungate, r. e. 1966 . the rumen and its microbes. new york: academic pro pearson, h. a. 1965. rumen organisms in white-tailed deer from south texas.]. wildl. mgmt . 29 :493-496. van hoven, w. 1974. ciliate protozoa and aspects of the nutrition of the hippopotamus in the kruger national park . . s. afr. j. sci. 70 : 107-109. van hoven, w. 1975. rumen ciliate fauna of the tsessebe (damaliscus lunatus lunatus) in south africa. j . protozool. 22 :457-462. westerling, b. 1970. rumen ciliate fauna of semi-domestic reindeer (rangifer tarandus l) in finland: composition, volume and . some seasonal variations. acta zoo/. fennica . 127: 1-75. 26 page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 filelist convert a pdf file! supplement to koedoe. 1977: 68-78. die erkenning v an die noodsaaklikheid van natuurbewaring ten opsigte van basiese grondgebruiksbeplanning in die republiek v an suid-afrika j j la grange direkteur van omgewingsaangeleenthede dept. van 8eplanning en die omgewing privaatsak x213 pretoria 0001 abstract the concept of a natural resource is explained and nature conservation as a form of land use is discussed in some detail. special reference is made to the national physical development of planning and the environment on the role played by nature conservation in basic usage of the soil as planned by the state in the republic of south africa. indien onder die begrip natuurbewaring verstaan word dat dit die bewaring en gebruik van natuurlike hulpbronne is, dan moet duidelikheid verkry word oor wat natuurlike hulpbronne presies is. daar bestaan talle definisies, en ek wil slegs na twee verwys. chapman (1969) definieer natuurlike hulpbronne, by wyse van opsomming, soos volg: "in the human ecosystem, man assigns utility to various elements of his environment and thus confers upon them the role of resources. resources then are neither wholly of the physical world nor wholly of the world of man but are the result of the interaction between the two. the environmental elements that man calls upon to serve as resources, and the nature and size of the requirements he places upon them, depend on his numbers, his needs and desires, on his values and skills." isard, chogwill, kissen, seyforth en tatlock (1972) definieer natuurlike hulpbronne as " ... those natural conditions and raw materials which man uses to meet his needs and improve his net welfare." in die algemeen moet die mens dan nog een of ander aksie uitvoer om hierdie natuurlike toestande en grondstowwe te verander na werklike goedere en dienste. andersom gestel, sal iets in die natuur 'n hulpbron word sodra die mens dit benodig en dit kan benut. hieruit kan afgelei word dat die tipe en benutting van natuurlike hulpbronne 'n funksie is van die mens se vermoe om dit te ontwikkel. gevolglik sal daar alternatiewe beleidsrigtings vir sodanige ontwikkeling 68 wees aangesien sekeres verkieslik bo ander sal wees. van die faktore wat hier bepalende roile kan speel is byvoorbeeld omgewingsfaktore. benutting van natuurlike hulpbronne is 'n vorm van ontwikkeling en die invloede van ontwikkeling veral op die ekologie moet reeds in die beplanningstadium in ag geneem word. dit is dan uit 'n beplanningsoogpunt dat die natuurlike hulpbronne, soos wat hulle onder die begrip van natuurbewaring verstaan word, in hierdie referaat beskou sal word. soos hierbo aangetoon, kan feitlik enigiets teoreties as 'n natuurlike hulpbron beskou word. in 'n poging om egter prakties te wees, word vir die doel van hierdie bespreking op die klassifikasie van isard et al. (1972) gelet. hiervolgens word natuurlike hulpbronne volgens hul hernubaarheid geklassifiseer ten einde die moontlike invloede van die mens se aktiwiteite op die natuurlike omgewing te verstaan. hierdie klassifikasie van hernubaarheid (hernubaarheid kan ook beskou word as potensiaal van vervanging), sluit drie tipes natuurlike hulpbronne in: (i) nie-hernubare hulpbronne by. fossielbrandstof en mineraalafsettings, of hulpbronne wat so stadig hernu word dat die hernuwing buite rekening gelaat kan word; (ii) vloeihulpbronne by. sonlig, wind en getye, en water in die hidrologiese siklus. hierdie hulpbronne word voortdurend in die natuur voorsien; en (iii) natuurlike hernubare hulpbronne aile lewende organismes wat in staat is om te groei en te reproduseer word hierby ingesluit. hierdie hulpbronne se graad van hernuwing hang af van die fisiese omgewing en die omvang van die voortbrengende voorraad. die afleiding word dus gemaak dat die natuurlike hernubare hulpbronne die mees sensitiewe vir veranderinge in die omgewing is en dus die meeste deur fisiese beplanning en ontwikkeling, geraak sal word. huidige grondgebruik ridkor (1972) dui aan dat die grondgebruik in sommige lande in 1967 soos volg was: verenigde state van amerika frankryk w esdui tsland verenigde koninkryk europa as geheel suid-afrika 1 ha per 0,22 persone; 1 ha per 0,81 persone; 1 ha per 2,15 persone; 1 ha per 2,02 persone; 1 ha per 0,82 persone; 1 ha per 0,18 persone of 5,7 ha per persoon. die posisie in die vsa dui op 'n verdere verdeling van die 4,45 ha per persoon (3,64 ha indien alaska en hawaii uitgesluit word) soos volg: 69 0,81 ha vir gewasverbouing; 0,20 ha vir konstruksiewerke soos huise, besighede, paale en open bare installasies; 1,21 ha vir weiding, bosbou en ontspanning; 0,16 ha vir parke, monumente, damme en wildernisgebiede; 0,70 ha vir ontspanningsterreine (bv. staatsbosse); die res van die 4,45 ha per persoon is moeilik om te klassifiseer aangesien dit meerdoelig kan wees . ongelukkig is direkte vergelykbare statistiek vir die republiek van suid-afrika (rsa) nie beskikbaar nie maar die volgende gegewens vir 1970 is insiggewend (t abel i): tabel 1 grondgebruik in die rsa (1970) % van totale oppervlakte grondgebruik oppervlakte landsper persoon (ha) oppervlakte (ha) (i ) bewerk t e landbougrond 18119000 (a) jaa rgewasse 12 171 000 (b) permanente gewasse 862000 14,83 0,85 (c) aangeplan te wei ding 932000 (d) ander 4 154000 (2) l\'atuurlike weiding 83 260000 68,19 3,88 (3) nasionale parke 2 938 201 2,40 0,14 (4) ander natuurreservate 296034 0,25 0,01 (5) bosbougebiede 1 558,960 1,28 0,07 (6) beboude g ebiede (dorpe) 3 237368 ± 2,60 0,15 (7) ander 12692637 10,39 0,59 totaal 122 104200 100,00 5,69 (gegewens volgens handelinge van simposium vir omgewingsbewaring, 1973.) direkte vergelykbare gegewens vir ander lande is nie beskikbaar nie, maar statistiek vir die vsa in 1969 toon byvoorbeeld dat landbou in die vsa 56,7% van die landsoppervlakte beslaan het teenoor 83,02% in die republiek van suid-afrika. die probleem as gevolg van beskawingsdruk is ons besig om al hoe vinniger hoer eise aan ons natuurlike hulpbronne of ons omgewing, te stel of om dit selfs te 70 "gebruik". ons het al hoe meer van ons oop ruimtes nodig vir stadsbou vir groeiende stedelike bevolkings, waarin voorsiening gemaak moet word vir bewoning, werk en ontspanning. hiervoor het ons klip, sand en beton nodig en daarom 'n gedeelte van ons bodem. om van ons woonplek tot by ons werkplek of ontspanningsplek te kom, het ons meer ruimte nodig vir paaie, spoorwee en vliegvelde. al hoe meer natuurlike gebiede word gebruik om voedsel te produseer. meer bykomende werkgeleenthede word verlang en daarvoor word tot 'n groot mate meer grondstowwe benodig dus meer grond vir mynbou. waar mynbou vroeer grootliks onder die grond plaasgevind het, word oopgroefmynbou in 'n toenemende mate toegepas en hierdeur word nog meer bogrond gebruik. hiervoor word die infrastruktuur van krag, water, en vervoer verhoog en gevolglik vermindering van bruikbare grondoppervlakte. grond word ook afgestaan vir die doeleindes van ontspanning soos byvoorbeeld om ons damterreine en daar is versoeke vir meer. in bogenoemde gebruike ontstaan daar op sekere plekke so 'n mate van gekonsentreerde afval dat die natuurlike omgewing dit nie kan absorbeer nie. besoedeling eis gevolglik 'n verdere gedeelte van ons omgewing. dit is dus duidelik dat die verskillende aansprake op gedeeltes van ons bod em vinnig vermeerder. daarenteen vermeerder die bodem nie. ten einde die toedeling van ons beperkte bod em aan die onderskeie behoeftes te maak, is dit nodig om met in ag neming van sekere f~ktore, prioriteite te bepaal en daaruit sekere riglyne vir optrede, neer te ie dit word vir die doel van hierdie bespreking grondgebruiksbeplanning genoem. dit moet in ag geneem word dat daar vroeer genoeg ruimte was vir die behoeftes van elke instansie op sy eie, maar vandag qioet daar na die aangeleentheid op 'n gekoordineerde grondslag gekyk word. indien ons verstandig te werk gaan dan sal daar genoeg ruimte vir aile gebruike wees . beleid vir die doel van hierdie bespreking kan die volgende as regeringsbeleid aangehaal word uit african wildlife (1976): "die land se hulpbronne is die erfenis van sy mense. party van ons hulpbronne is beperk en die regering beskerm hulle teen wanbenutting. dit is trouens die regering se plig om te verseker dat sodanige hulpbronne oordeelkundig tot voordeel van die land as 'n geheel gebruik word. dit is die regeringsbeleid om gebiede van besondere natuurskoon of unieke natuurverskynsels of gebiede waar ongewone plantof diersoorte voorkom, teen wanbenutting en omgewingsbesoedeling te beskerm en te verseker dat sodanige gebiede tot voordeel en genot van ons land se mense gebruik word. die algemene beskouing in suid-afrika is om nie teen die mens nie, maar vir die mens te bewaar." van besondere belang is die beleid ten opsigte van landbougrond (dept. van beplanning en die omgewing 1975a). by geleentheid van die 71 habitatraad se konferensie oor kusgebiede op 3 april 1975, te durban, het sy edele j j loots, die toenmalige minister van beplanning en die omgewing in sy openingsrede onder andere na die bewaring van landbougrond verwys: "sonder om daarvan 'n lang storie te maak noem ek vir u dat ek en my departement, asook die minister van landbou en sy hoofamptenare, al meer ernstig bekommerd begin raak het oor die tempo waarteen goeie landbougrond deur ontwikkeling soos dorpstigting, mynbou, padbou" en ' n groot aantal ander doeleindes in beslag geneem word. suid-afrika het maar altesaam 15% bewerkbare grond, en ons hoe-potensiaal gronde beloop slegs 3,3% van die totaal. hierdie 3,3% lewer 40% van ons landbouproduksie. my departement is van plan om in samewerking met die departement van landbou, deur middel van die opstel van gidsen streekplanne, hierdie waardevolle en hoe-produserende landbougronde, sover enigsins moontlik , te beskerm en vir landbou te behou. om ons daarin te help het ons nou 'n besluit van die kabinet gekry wat: o bevestig dat die behoud van goeie landbougrond 'n saak van nasionale belang is; odie departement van landbou-tegniese dienste versoek om as 'n saak van voorkeur die goeie gehalte landbougrond te identifiseer; o aan die departement van beplanning en die omgewing opdrag gee, in samewerking met aile ander ins tansies betrokke by die opstel van gidsplanne en streekplanne, om in aile beplanning die hoogste prioriteit te verleen aan die behoud van hoe-potensiaallandbougrond vir voedselproduksie." daar is nog ander voorbeelde van beleid oor bewaring maar by die twee aanhalings kan volstaan word. wat gedoen word grondgebruiksbeplanning is 'n konsep wat aktiwiteite van onder andere die volgende staatsdepartemente en ander liggame insluit: waterwese, landbou, bosbou, vervoer, verdediging, mynwese, bantoeadministrasie en -ontwikkeling, die nasionale parkeraad, elektrisiteitsvoorsieningskommissie (evkom), die provinsiale administrasies, plaaslike besture, die meergebiede-ontwikkelingsraad, die raad vir nasionale gedenkwaardighede, en die sa natuurstigting. hierdie liggame pleeg dikwels oorleg met die departement van beplanning en die omgewing. daar is geen wetlike verpligting daarvoor nie. al hoe meer van die betrokke liggame pak egter hul beplanning op 'n gekoordineerde grondslag in medewerking met andere aan. die volgende gevalle word as voorbeelde genoem van beplanning waar veral ook na natuurbewaring omgesien word: 72 (i) departement van beplanning en die omgewing die nasionale fisiese ontwikkelingsplan (nfo), (dept van beplanning en die omgewing, 1975b) hierdie plan maak oorhoofs voorsiening vir die doeltreffende benutting van die land se beskikbare hulpbronne met inagneming van die bodemgesteldheid en die bevolkingsverspreiding. bestaande belange en aansprake op die gebruiksmoontlikhede van beskikbare oppervlaktes word in ag geneem. in die plan word getrag om 'n fisiese raamwerk te skep waarbinne die onderskeie bedrywighede van landbou-, bosbou-, mynbou-, behuisings-, onderwys-, gesondheids-, vervoer-, ontspanningen ander maatskaplike dienste onderneem kan word. die nfo bepaal dus riglyne vir optimale aanwending van landsoppervlakte vir effektiewe beplanning deur die onderskeie ins tansies, soos byvoorbeeld natuurbewanng. die beplanningsadviesraad van die eerste minister in opdrag moet die adviesraad die eerste minister adviseer oor waar die mense van ons land teen die jaar 2000 moet woon"werk en ontspan. in sy ad vies aan die eerste minister, en die kabinet, word aandag geskenk aan grondgebruiksbeplanning en natuurbewaring. 'n groot gedeelte van die adviesraad se werk word deur hulpkomitees gedoen. enkele voorbeelde is: hulpkomitee vir die beskikking van staatsgrond vir ontspanning en ander doeleindes, en by damterreine (dept. vim beplanning en die omgewing 1975c) hierdie komi tee bedien die ministers van landbou en waterwese van advies. die behoeftes van natuurbewaring ontvang hier die nodige aandag. hulpkomitee vir die kusgebied (dept. van m~planning en die omgewing 197 5a) planne word opgestel m.b.t. grondgebruike wat veral aandag vestig op bewaringswaardige gebiede soos riviermonde, strandmere en vleie. hulpkomitee vir nasionale berggebiede (la grange 1976) die opdrag aan hierdie komi tee is: (a) om ondersoek in te stel na die bewaring van berggebiede op 'n nasionale grondslag; en (b) om aanbevelings aan die onderskeie ins tansies te maak oor hoe die onderskeie berggebiede op ' n gekoordineerde grondslag benut en bewaar kan word. 73 'n vroeere verslag oor die grondgebruik in die drakensbergopvanggebied in natal is reeds in 1972 gepubliseer. ondersoeke oor die bewaring van die magaliesberg en die kaapse skiereiland is tans aan die gang. hulpkomitee vir buitelugontspanning (dept. van beplanning en die omgewing 1976a) aangesien buitelugontspanning en natuurbewaring verband met mekaar hou, word daar na natuurbewaring in die werksaamhede van hierdie komi tee omgesien. raad vir die omgewing (ro) hierdie raad is deur die minister van beplanning en die omgewing ingestel en is 'n adviserende en koordinerende liggaam. die ro wat 'n geskikte forum bied vir die bespreking van natuurbewaring teen die bree agtergrond van grondgebruiksbeplanning, het die volgende ins tansies as lede: departemente van beplanning en die omgewing, landboutegniese dienste, bantoe-administrasie en -ontwikkeling, gemeenskapsbou, bosbou, gesondheid, nyw~rheidswese, arbeid, mynwese, nasionale opvoeding, vervoer, waterwese, die vier provinsiale administrasies, administrasie van suidwes-afrika, raad op atoomkrag, buro vir standaarde, wetenskaplike en nywerheidsnavorsingsraad, elektrisiteitsvoorsieningskommissie, nasionale parkeraad, habitatraad, georganiseerde nywerheid, en die verenigde munisipale bestuur. die minister van beplanning en die omgewing, is voorsitter van die kabinetskomitee vir die omgewing waarvan die ministers van vervoer, gesondheid, landbou, ekonomiese sake en waterwese lede is. nasionale komitee vir natuurbewaring (nakor) in nakor is die volgende instansies wat met natuurbewaring gemoeid is verteenwoordig: die departement van natuurbewaring van die provinsiale administrasies en die administrasie van suidwes-afrika, die nasionale parkeraad, die nasionale botaniese tuine, die departemente bosbou, waterwese, landboukrediet en -grondbesit, bantoeadministrasie en -ontwikkeling, verdediging, beplanning en die omgewing, vervoer, landbou-tegniese dienste, mynwese, asook die wnnr. die komitee skakel ten nouste met die raad vir die omgewing. een van die belangrikste huidige take is om 'n nasionale natuurbewaringsplan op te stel waarna dan gepoog sal word om hierdie behoeftes in die nfo in te werk. navorsing navorsing het 'n belangrike bydrae te make in verband met grondgebruiksbeplanning. die ro het die nouste skakeling met die nasionale program vir omgewingsnavorsing. hierdie program word op advies van die komitee vir omgewingswetenskappe, deur die wnnr vir die departement van beplanning en die omgewing asook die staatsinstansies, administreer. die werk van hierdie program is van toegepaste aard en hou direkte verband met grondgebruik. beheermaatreehs bo en behalwe die feit dat die departement die voorsitters asook die sekretariaat verskaf vir die beplanningsadviesraad van die eerste minister en die raad vir die omgewing, speel dit 'n verdere belangrike rol ten opsigte van grondgebruiksbeplanning deur middel van sekere beheermaatreels. die belangrikste beheermaatreels wat die departement op sy eie ten opsigte van grondgebruik het word vervat in die wet op omgewingsbeplanning, 1967. artikel4(1) van die wet kan soos volg opgesom word: die minister kan na raadpleging met die minister van landbou en die administrateur van die betrokke provinsie by kennisgewing in die staatskoerant grond, wat in daardie kennisgewing vermeld word, vir sekere doeleindes voorbehou. van die doeleindes wat vir hierdie bespreking ter sake is, is natuurgebiede. in die wet word die begrip 'natuurgebied' soos volg beskryf: '''n gebied wat gebruik sou kan word in belang van en tot voordeel en vir die genot van die publiek in die algemeen en vir die voortplanting, beskerming of behoud van wilde dierelewe, wilde plantegroei of voorwerpe van geologiese, etnologiese, geskiedkundige of ander wetenskaplike belang". die voorwerpe wat in hierdie beskrywing van natuurgebiede genoem word, kan vir praktiese doeleindes biologies en nie-biologies ingedeel word. oor die biologiese gedeelte, dit wil se ". . . voortplanting, beskerming of behoud van wilde dierelewe, wilde plantegroei ... " word die departement ook deur die ro en nakor van advies bedien. vir die tweede gedeelte naamlik " ... voorwerpe van geologiese, etnologiese, geskiedkundige of ander wetenskaplike belang", het die ro 'n werkgroep ingestel waarin die betrokke ins tansies verteenwoordig is. dit word beklemtoon dat die nouste samewerking met die raad vir nasionale gedenkwaardighede bestaan. gevolglik kan die departement in oorlegpleging met die minister van landbou en die betrokke administrateur, in sekere gevalle 'n gebied as 'n natuurgebied voorbehou. in wese beteken dit dat die eienaar van sodanige gebied met sy bestaande gebruik van die gebied kan voortgaan maar indien hy die gebruik sal wil verander, dan kan dit net vir 'n natuurgebied wees. hierdie saak kan sy direkte toepassing in berggebiede vind. die departement kan ook langs 'n ander weg beheer uitoefen oor grondgebruik, en vir die doel van hierdie konferensie dus ook oor grondgebruik in berggebiede. die hele rsa, uitgesonderd grond wat in 'n regsgebied van 'n plaaslike owerheid voorkom waar 'n beplanning75 skema bestaan, ofwat in bantoegebied gelee is, is in terme van artikel5 van bogenoemde wet, op 5 oktober 1973 as 'n beheerde gebied verklaar vir die doeleindes van die verandering in gebruik van grond. voor hierdie datum was daar reeds 'n aantal beheerde gebiede waar dieselfde beheer gegeld het. dit beteken dat as grond vir 'n ander doel gebruik wil word as die doel waarvoor dit gebruik was op die datum van beheerdverklaring (dit beteken verskillende datums afhangende van die datum van beheerdverklaring van die gebied) dan moet die applikant vir sodanige verandering toestemming van die departement verkry. voorbeelde is sake-ondernemings soos winkels, restaurante, padkafees, vulstasies en slaghuise. beheer in sulke gebiede word kragtens artikel 6(1)(c) van die wet uitgeoefen . die departement kon ook op 'n derde manier beheer oor verandering van grondgebruik uitoefen. ingevolge 'n wysiging van bogenoemde wet (artikei6b( i)), kan die departement nou ook beheer uitoefen oor groewe onder andere van klip, klei, sand, gruis en grond vir byvoorbeeld steenmakerye, gruis vir konstruksiewerk (ander dan paaie en spoorwee) en bogronds vir grasperke. ingevolge artikel6b(2) kan die departement nou ook na raadpleging met die departement van mynwese, beheer uitoefen oor mineraalverwerkingsaanlegte soos byvoorbeeld steenkoolwassery, sandwassery, smelterye van byvoorbeeld grond, koper, yster en chroom. die bepalings van artikel 6b(1) en (2) is geldig op enige plek in die republiek van suid-afrika. die departement het geen beheer oor die verandering van grondgebruik vir mynbou as sodanig nie. onder artikel 6(2) (d) van die wet op omgewingsbeplanning, 1967, oefen die departement egter wel beheer uit oor die verandering van grondgebruik vir behuising en sakeondernemings by myne. gidsplanne gidsplanne kan vir beide stedelike en omstedelike gebiede opgestel word. die prosedure vir die opstel van 'n gidsplan deur 'n gidsplankomitee, en die goedkeuring daarvan al dan nie, deur die minister van beplanning en die omgewing, word uiteengesit in die wet op omgewingsbeplanning, 1967. wysigings tot 'n goedgekeurde gidsplan, of die terugtrekking daarvan, kan net bewerkstellig word deur die hele prosedure soos vir die opstel van die gidsplan te herhaal. enige belangstellende persoon kan voorstelle vir insluiting in 'n gidsplan maak. kortliks saamgevat word in so 'n gidsplan die toekomstige ontwikkelingsbehoeftes en grondgebruik in die gidsplangebied aangedui. ander planne wat reeds voltooi is of byna voltooi is, maar nie volgens die voorskrifte van die wet opgestel is nie en dus nie statutere dokumente is nie, word genoem "riglyne vir t oekomstige ontwikkeling." 76 (ii) ander instansies enkele voorbeelde van wat ander ins tansies doen betreffende natuurbewaring in hul grondgebruiksbeplanning is : departement van waterwese deur middel van sy drakensbergprojek-omgewingskomitee (dept. van beplanning en die omgewing 1974):hierdie komitee het aan die departement van waterwese en evkom advies gelewer oor stappe om die ongewenste effekte van die tweede fase van die tugela-vaal staatswaterprojek en die drakensbergpompopgaar-ontwikkeling soos in die witpapier wpr-'74 beskryf, te voorkom of te verminder. die departement van waterwese het verder 'n komitee vir ondersoek van die ekologie van die umfolozirivier ingestel (dept. van beplanning en die omgewing 1975a) in verband met die keuse van ' n beoogde damterrein in die u mfolozirivier. die d epartement van mynwese:'n werkskomitee ingestel om hom te advis ee r oor gebiede vir mynbou aan die zoeloelandse kus asook die nodige voorsorgmaatreels vir behoud of die herstel van die natuurlik e omgewing tydens mynbou-aktiwiteite . voorts het die departement van mynwese 'n komitee ingestel om hom te ad vi seer in verband met oopgroefmynbou (dept. van beplanning en die omgewing 1976b) en veral oor die inslag wat sodanige mynbou op landbougrond sal he. siot-opmerkings ons bodem is nie groot genoeg vir ongekoordineerde gebruiksaansprake nie . instansies wat met gesag daartoe belas is, moet met die toedeling van die bodem die volgende faktore by besluitnemings in oorweging neem : ekologiese, kultuur-historiese, estetiese, politi eke, tegniese, ekonomiese, sosiale, en die wetlike. die bepaling van watter gewig aan elke faktor toegeken moet word, is debatteerbaar. verwysings anon , 1976. ons stem saam! hoe iyk dit met sames melting? afr. wildlife 30( i) : 41-43. chapman , j d 1969. interactions between m a n and hi s resources. in : resources and man. dept. van beplanning en die omgewing , 1974. omgewinglen vironment rsa oktober. dept. van beplanning en die omgewing, 197 5a . omgewingl environm ent rsa m ei. dept. van beplanning en die omgewing , 1975b. nasionale fisiese ontwikkelingsplan. dept. van beplanning en die omgewing , 1975c. omgewinglenvironment rsa juli e. dept. van beplanning en die omgewing , 1976a. omgewingl environment rsa junie. 77 dept. van beplanning en die omgewing, 1976b. omgewing/environment rsa april. isard, w c , l chogwill,] kiss in , r h seyforth and r tatlock, 1972. ecologic-economic ana(ysis for r egional development la grange,]] 1976 . bergamgewings van suid-afrika: die ra1 van die departement va n beplanning en die amgewing. kanferensie van die habitatraad. ridgor, r c 1972. resource and environmental consequences of population growth in the united states. 78 page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 abstract introduction methods and materials results discussion implications for conservation acknowledgements references about the author(s) claire n. gordon department of conservation ecology and entomology, stellenbosch university, south africa liesl eichenberger sanbona wildlife department, sanbona wildlife reserve, south africa paul vorster sanbona wildlife department, sanbona wildlife reserve, south africa alison j. leslie department of conservation ecology and entomology, stellenbosch university, south africa shayne m. jacobs department of conservation ecology and entomology, stellenbosch university, south africa citation gordon, c.n., eichenberger, l., vorster, p., leslie, a.j. & jacobs, s.m., 2016, ‘diet and seasonal dispersal of extralimital giraffe at sanbona wildlife reserve, little karoo, south africa’, koedoe 58(1), a1346. http://dx.doi.org/10.4102/koedoe.v58i1.1346 short communication diet and seasonal dispersal of extralimital giraffe at sanbona wildlife reserve, little karoo, south africa claire n. gordon, liesl eichenberger, paul vorster, alison j. leslie, shayne m. jacobs received: 12 aug. 2015; accepted: 03 june 2016; published: 31 aug. 2016 copyright: © 2016. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract south african giraffe (giraffa camelopardalis giraffa) have been introduced as an extralimital species to private farms in the little karoo on the basis of economic sustainability, and the need to create a competitive tourism product. however, little is known about the mediumto long-term impacts and ecological sustainability of such introductions. the diet of a population of giraffe on sanbona wildlife reserve, near the town of ladismith, was assessed via direct observations between january and october 2014, in order to determine their potential impact on the world’s most species-rich semi-desert, the succulent karoo. unlike giraffe in their native range, the sanbona population showed seasonal preference for browse species. acacia karroo (sweet thorn) appears to be the preferred browse species during autumn and spring, with schotia afra being the preferred species in winter, and no significant preference being shown in summer. giraffe also appeared to seasonally move between catchments where tree species other than a. karroo occurs, especially during winter and spring when the tributaries of the brak river, containing mixed acacia with s. afra (karoo boer-bean) and euclea undulata (small-leaved guarri), were visited with increasing frequency. these results largely confirm the importance of a. karroo as the main browse species in this environment but also suggest that other species may be important components of the diet of extralimital giraffe in the little karoo. on farms where a. karroo is dominant, supplementary feed may be needed when a. karroo browse is unavailable due to leaf drop. conservation implications: acacia karroo was the main browse species of extralimital g. c. giraffa at sanbona wildlife reserve, but it switched to s. afra during winter. this suggests that an assessment of alternative food species forms part of suitability assessments for the introduction of extralimital g. c. giraffa for areas similar to sanbona. introduction over the past two decades, large areas in south africa have seen shifts in land use from agriculture to game farming, for either commercial hunting or ecotourism and the photographic safari industry (vorster 2011). most of these land owners base their management decisions on economic sustainability (grant, peel & bezuidenhout 2011), often introducing extralimital species (castley, boshoff & kerley 2001). charismatic extralimital species, such as the south african giraffe, have been introduced to the western and eastern cape due to their popularity among foreign tourists (parker & bernard 2005). the browse preference of south african giraffe has been studied extensively in the savanna-dominated biomes of southern africa (bond & loffel 2001; fennessy 2004) and a few studies have been conducted in the mesic thicket-dominated eastern cape, where they are also extralimital (parker & bernard 2005). however, no literature is readily available which documents the species dietary preferences or potential impacts in areas further west, such as game farms of the little karoo, which straddle the fynbos, succulent karoo and thicket biomes. the little karoo falls within three unique biodiversity hotspot vegetation types, namely the succulent karoo, fynbos and subtropical thicket biomes (egoh et al. 2009). the lower lying areas of the little karoo are dominated by succulent karoo vegetation, which is the most species-rich semi-desert vegetation type in the world (mucina & rutherford 2006). the succulent karoo, therefore, is an area of high conservation priority, and as most of the land lies in the hands of private landowners, the growth and ecological sustainability of conservation in the area are dependent on adequately managing these private lands (pasquini et al. 2009). the recent introduction of certain extralimital species, such as giraffe (giraffa camelopardalis), elephant (loxodonta africana) and hippopotamus (hippopotamus amphibious), to the little karoo, is a potential cause for concern, as little research has been conducted into their impact on the region’s vegetation, or the suitability of local vegetation types to sustain these animals. even though 13% of south africa’s total land surface is used for game farming, very little research has been conducted into the impact indigenous herbivore species have on rangelands (hoffman et al. 2009). it is therefore imperative to determine the impacts such extralimital species will have on the surrounding vegetation or habitat. three key questions were asked: (1) what is the species composition of the south african giraffe populations’ diet, (2) what is the most preferentially browsed plant species and (3) are there seasonal changes in diet and movement across the reserve? the major aim of this study was to determine the major diet species of giraffe introduced to the little karoo. methods and materials study area sanbona wildlife reserve is located in the little karoo region of the western cape, south africa, 49 km northeast of the town of montagu and 27 km northwest of the town of barrydale. the reserve is approximately 54 000 ha in size and measures 25 km from north to south, and 30 km from east to west. it is the largest privately owned conservation area within the western cape and offers an ecotourism and photographic safari product. it hosts the only free-roaming big five safari experiences in the province (lynch, vorster & vorster 2013), which includes a small population of lions that occasionally predates on the giraffe population (a. hughes [sanbona wildlife department] pers. comm., september 2014). sanbona, established in 2002, consists of 19 formerly agricultural farms which were merged by removing internal fencing. the farms were previously utilised for wheat and lucerne crops, domestic animal production, small-scale commercial game farming for hunting and game sales, recreational farming and tourism (lynch et al. 2013). sanbona is divided into two main sections: sanbona north and sanbona south, which are divided by the warmwaterberg range. for management reasons, most of the large and dangerous game species are found in sanbona north; this includes the giraffe population. sanbona north consists of a more arid environment, dominated by succulent karoo vegetation, specifically the western little karoo and the little karoo quartz vygieveld vegetation types (lynch et al. 2013; mucina & rutherford 2006). the landscape is undulating and flat in parts, consisting of low-to-medium-height mosaic karoo shrublands with both succulent and non-succulent plants. the main drainage line in sanbona north, the brak river, is dominated by dense stands of acacia karroo, and there are several smaller tributaries which feed into it. the vegetation alongside these smaller tributaries’ consists of a mosaic of species, largely dominated by schotia afra and euclea undulata and interspersed with a. karroo. we recognised four seasons in the area, namely: summer (01 december – 28/29 february), autumn (01 march – 31 may), winter (01 june – 31 august) and spring (01 september – 30 november). sanbona lies in a transition zone between the summer and winter rainfall areas and experiences slightly higher rainfall averages in winter months. sanbona south (renosterveld) receives more rainfall than sanbona north (succulent karoo) due to the rainfall shadow created by the warmwaterberg range. the reserve is susceptible to sporadic drought and flooding events, and sanbona north experiences convectional thunderstorms in summer (lynch et al. 2013; mucina & rutherford 2006). in the hottest month, february, the mean daily temperature is 21 °c with midday temperatures regularly exceeding 35 °c. june is the coldest month with average daily temperatures of 6.2 °c, and early morning temperatures regularly falling below freezing (lynch et al. 2013). the study species giraffe are an iconic african mammal. adults can stand 4 m – 5.5 m tall, and weigh as much as 700 kg – 1100 kg (skinner & chimimba 2005). both males and females have ossicones, with the males often thicker, knobbed and hairless at the ends. giraffe are gregarious, associating in non-territorial, loose, open herds with no fixed social hierarchy. individuals rarely stay together for prolonged periods of time, with the exception of females with calves (ciofolo & le pendu 2013; estes 1992; leuthold 1979; tutchings et al. 2013). giraffe are generally observed to be exclusively browsers, although some records of grazing behaviour have been reported (estes 1992; seeber et al. 2012). giraffe are efficient feeders; not only are they able to browse above the level of most other species, but their tongue is able to navigate around the most aggressive thorns in order to consume up to 34 kg of foliage a day (estes 1992). giraffe are currently listed as ‘least concern’ by the iucn, as their population is estimated to number approximately 80 000 individuals across africa (fennessy & brown 2010; tutchings et al. 2013). the natural distribution range of giraffe includes the arid and dry savanna zones of sub-saharan africa, wherever trees occur (fennessy & brown 2010; tutchings et al. 2013). south african giraffe are extralimital to the western cape and the succulent karoo. sanbona initially introduced a founder population of six sub-adult giraffe in 2005. in 2008, a further eight individuals were introduced to the reserve (lynch et al. 2013). as of the 2014 game count, the total population stands at 28 individuals (l. vorster [sanbona wildlife department] pers. comm., september 2014). determining giraffe distribution since 2008, sanbona has been collecting distribution data on various species via field guide sighting sheets (standardised forms on which the field guides note the location of various animal species on their daily game drives). for the purpose of this study, sighting sheets from 2009 to the end of 2013 were analysed. sighting sheets recorded general giraffe sightings and omitted any demographic information. analysis involved recording all noted locations into an excel spreadsheet from the hard copy sighting sheets. gps co-ordinates were collected by comparing written records with google earth images of the reserve and a hard copy of the reserve map which shows road and landmark names. gps co-ordinates were then exported into qgis (qgis development team 2014), together with existing reserve shapefiles, in order to map the seasonal distribution of the giraffe for the period 2009–2013. direct observations of giraffe behaviour in order to determine the primary browse species for the giraffe population during the four seasons, direct observations were conducted. we used the interval scan method, as used by parker (2004) in a study on giraffe in the eastern cape. direct observations involved observing the first group of giraffe we found on a sampling day (lone males were excluded) for two consecutive hours, from a distance of no less than 200 m. observations were conducted daily, for four consecutive days. feeding records were recorded by scanning through the group using binoculars, and noting what species each individual was eating every 2 min. plant species which were not identified through binoculars were identified by following up on foot at the end of the two-hour period. data analysis for the direct observation data, initial analysis involved compiling graphs in excel in order to determine which species of plants were the most commonly browsed across all four seasons. shapiro–wilk tests were conducted on all four data sets to test for normality. the five most preferred browse species were then analysed against each other using the kruskal–wallis test, for each of the four seasons, as all four data sets were not normally distributed. all statistical analyses were conducted using statistica 12 (statsoft inc. 2013). results distribution mapping the results of the distribution mapping (figure 1) showed that in autumn and spring, the giraffe utilised very similar areas, with spring showing a slightly larger preference for the smaller tributaries, as well as the brak river and the main acacia thicket. autumn showed very little usage of the smaller tributaries, with most of the giraffe clustering around the brak river and an occasional few straying as far as the bellair dam. there was a more visible difference between summer (figure 1a) and winter (figure 1b) distribution, with giraffe showing a marked increase in movement into the smaller tributaries in winter. in both seasons, however, individuals were still found to move quite far from the brak river and main acacia thicket, but this appeared to be more frequent in winter. in both summer and winter, unlike the other two seasons, individuals were found to move as far south as tilney gorge, south of the bellair dam, a distance of approximately 20 km. figure 1: seasonal movement and distribution of giraffe on sanbona. (a) the summer movement pattern, (b) winter movement pattern. the summer and winter maps showed the largest visual difference, with summer showing a strong clustering around the brak river and winter showing movement into the smaller tributaries. direct diet observations the direct observations revealed that giraffe on sanbona browse on 17 plant species, including two alien invasive species, namely pink tamarisk (tamarix ramosissima) and old man saltbush (atriplex nummularia). it also revealed that bone forms a part of their diet in the winter months, although the frequency of this could not be quantified. for further analyses, the five most preferred plant species were selected. across all four seasons, a. karroo was browsed significantly more than any other species (h = 1552.41; n = 4836; p < 0.001; figure 2). although e. undulata and s. afra were not the most favoured browse species, they were significantly favoured over both the lycium species and salsola aphylla (h = 1552.41; n = 4836; p < 0.001). summer showed no species to be significantly favoured, although a. karroo, s. afra and e. undulata were browsed significantly more than either s. aphylla or the lycium species (h = 254.90; n = 1179; p < 0.001; figure 2a). in autumn, a. karroo was browsed significantly more than any of the other four species (h = 773.03; n = 1210; p < 0.001; figure 2b), while in winter, s. afra was browsed significantly more than any of the other four species (h = 280.08; n = 1225; p < 0.001; figure 2c). just as in autumn, in spring a. karroo was browsed significantly more than any of the other four species (h = 1060.70; n = 1215; p < 0.001; figure 2d). figure 2: box and whisker plots to show the results of the kruskal–wallis tests performed on the direct observation data of giraffe feeding behaviour. (a) the results for summer, where no individual was found to be significantly favoured, (b) the results for autumn, where acacia karroo was favoured the most, (c) the results for winter, where schotia afra was found to be the most favoured browse species, (d) the results for spring, where acacia karroo was by far the most preferentially browsed plant species. discussion the movement patterns of the south african giraffe population on sanbona appear to be highly seasonal. during summer and autumn, the giraffe cluster around the brak river, and the main acacia thicket. in winter, there seems to be visible movement out of the main river line and into the smaller tributaries. this coincides to an extent with the seasonal change in preferred browse, as these smaller tributaries are home to most of the reserve’s s. afra and e. undulata. as the acacia thicket in the brak river is affected by the first few frosts of winter, and the acacia spp. start to drop their leaves, the giraffe begin to actively disperse in search of alternate food sources. just like giraffe in other arid areas, the sanbona population seems to migrate and disperse seasonally, following food availability (fennessy 2009; le pendu & ciofolo 1999). the giraffe on sanbona showed strong seasonal preferences for different browse species. unlike in their native range and in kwa-zulu natal, where browse from the genus acacia was preferred throughout the year (bond & loffell 2001), the sanbona population consumed significantly less a. karroo in winter, most likely due to frost related foliage lost, with s. afra becoming the most browsed species. these results were similar to those found by parker and bernard (2005) in their study of extralimital giraffe in the eastern cape. the eastern cape study revealed that searsia longispina was the most commonly browsed species in winter. however, since this species is not prevalent on sanbona, the giraffe population used s. afra and e. undulata as alternative browse sources in the winter months. the sanbona giraffes’ switch in preference from a. karroo to s. afra and e. undulata as a food source in winter coincided with most of the a. karroo on the reserve dropping their foliage after the first few frost events. although not formally recorded, it is noteworthy that the giraffe were regularly observed feeding on plants below 1.5 m in height, particularly s. aphylla and a. karroo. several other studies have found giraffe to regularly feed below their maximum foraging height (leuthold 1978; leuthold & leuthold 1972). this feeding behaviour may indicate that available browse at mean foraging height may not be able to fulfil all dietary requirements. this is an important consideration, as little is known about the nutrient requirements of giraffe, and whether available browse in the little karoo is able to fulfil these requirements. implications for conservation as is the case for much of south africa, the majority of the little karoo lies in the hands of private landowners and so the challenge of conserving this highly species-rich region lies in finding a balance between managing for conservation and ensuring economic sustainability (pasquini et al. 2009). the introduction of giraffe onto sanbona, as in many parts of the western cape, was driven by the desire to create a competitive tourism product. anecdotal evidence suggests that giraffe have already been introduced to over 20 properties in the region (j. gird [living lands] pers. comm. september 2014). the results of this study also provide useful management suggestions for other landowners in surrounding areas who have introduced or intend to introduce giraffe. this study, carried out in the previously under researched little karoo, supports the importance of a. karroo in giraffe feeding behaviour found by parker and bernard (2005) in their eastern cape study. furthermore, it suggests that a. karroo phenology may be constraining browse availability during certain times of the year, most likely associated with frosts. the implications of this for any manager or property owner intending to introduce giraffe to their property in either the western or the eastern cape is that extralimital giraffe populations in these regions need sufficient populations of evergreen tree species to utilise as winter browse. if a property contains very few alternatives to a. karroo, it may result in managers being forced to provide supplementary feed for their giraffe or face the possibility of suffering fatalities in the winter months. before giraffe are introduced to any property, baseline vegetation or habitat assessments should be conducted to establish whether suitable sources of winter browse are available. acknowledgements we would like to thank the following sanbona staff members: andrew slater, ruan erasmus and andrew hughes. thanks to justin gird (living lands) for his suggestions and inspired debates about all things wildlife. thanks to matt wijers (su) and elsa bussier (uct) for their assistance with qgis, as well as gillian gordon (struik) for her assistance with editing, and dr justin harvey (su) for his assistance with the statistical analyses. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions the research being reported was carried out by c.n.g., when she was a 4th-year conservation ecology student at stellenbosch university, and l.e. and p.v. assisted with fieldwork and also carried out some of the fieldwork (one season). s.m.j. and a.j.l. supervised the student. the funding was supplied by the student and sanbona wildlife reserve. references bond, j.w. & loffell, d., 2001, ‘introduction of giraffe changes acacia distribution in a south african savannah’, african journal of ecology 39, 286–294. http://dx.doi.org/10.1046/j.1365-2028.2001.00319.x castley, j.g., boshoff, a.f. & kerley, g.i.h., 2001, ‘compromising south africa’s natural biodiversity – inappropriate herbivore introductions’, south african journal of science 97, 344–348. ciofolo, i. & le pendu, y., 2013, ‘subfamily giraffinae – giraffe’, in j. kingdon & m. hoffmann (eds.), mammals of africa, volume vi, pigs, hippopotamuses, chevrotain, giraffes, deer and bovids, pp. 96–110, bloomsbury publishing, london. egoh, b.n., reyers, b., carwardine, j., bode, m., o’farrell, p.j., wilson, k.a. et al., 2009, ‘safeguarding biodiversity and ecosystem services in the little karoo, south africa’, conservation biology 24(4), 1021–1030. http://dx.doi.org/10.111/j.1523-1739.2009.01442.x estes, r., 1992, the behaviour guide to african mammals, university of california press, los angeles, ca. fennessy, j., 2004, ecology of desert-dwelling giraffe giraffa camelopardalis angolensis in northwestern namibia, phd thesis, university of sydney, sydney. fennessy, j., 2009, ‘home range and seasonal movements of giraffa camelopardalis angolensis in the northern namib desert’, african journal of ecology 47, 318–327. http://dx.doi.org/10.1111/j.1365-2028.2008.00963.x fennessy, j. & brown, d., 2010, ‘giraffa camelopardalis’, in the iucn red list of threatened species, viewed 30 september 2014, from http://www.iucnredlist.org/details/9194/0 grant, r.c.c., peel, m.j.s. & bezuidenhout, h., 2011, ‘evaluating herbivore management outcomes and associated vegetation impacts’, koedoe 53(2), 1–15. http://dx.doi.org/10.4102/koedoe.v53i2.1008 hoffman, m.t., madden, c.f., saayman, n. & botha, j.c., 2009, ‘the impact of indigenous ungulate herbivory over five years (2004–2008) on the vegetation of the little karoo, south africa’, african journal of range and forage science 26(3), 169–179. http://dx.doi.org/10.2989/ajrf.2009.26.3.8.953 le pendu, y. & ciofolo, i., 1999, ‘seasonal movements of giraffes in niger’, journal of tropical ecology 15, 341–253. http://dx.doi.org/10.1017/s0266467499000863 leuthold, b.m., 1979, ‘social organization and behaviour of giraffe in tsavo east national park’, african journal of ecology 17, 19–34. http://dx.doi.org/10.1111/j.1365-2028.1979.tb00453.x leuthold, b.m. & leuthold, w., 1972, ‘food habits of giraffe in tsavo national park, kenya’, east african wildlife journal 10, 129–141. http://dx.doi.org/10.1111/j.1365-2028.1972.tb00867.x leuthold, w., 1978, ‘ecological separation among browsing ungulates in tsavo east national park, kenya’, oecologia 35, 241–252. http://dx.doi.org/10.1007/bf00344735 lynch, k., vorster, l. & vorster, p., 2013, environmental management plan. sanbona wildlife department, sanbona wildlife reserve, montagu. mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, strelitzia 19, south african national biodiversity institute, pretoria. parker, d.m., 2004, ‘the feeding biology and potential impact of introduced giraffe (giraffa camelopardalis) in the eastern cape province, south africa’, msc dissertation, rhodes university, south africa. parker, d.m. & bernard, r.t.f., 2005, ‘the diet and ecological role of giraffe (giraffe camelopardalis) introduced to the eastern cape, south africa’, journal of zoology 267, 203–210. http://dx.doi.org/10.1017/s0952836905007399 pasquini, l., cowling, r.m., twyman, c. & wainwright, j., 2009, ‘devising appropriate policies and instruments in support of private conservation areas: lessons learned from the klein karoo, south africa’, conservation biology 24(2), 470–478. http://dx.doi.org/10.1111/j.1523-1739.01344.x qgis development team, 2014, qgis geographic information system version 2.6.0. open source geospatial foundation project, http://qgis.osgeo.org seeber, p.a., ndlovu, h.t., duncan, p. & ganswindt, a., 2012, ‘grazing behaviour of giraffe in hwange national park, zimbabwe’, african journal of ecology 50, 247–250. http://dx.doi.org/10.1111/j.1365-2028.2011.01314.x skinner, j.d. & chimimba, c.t., 2005, the mammals of the southern african sub-region. cambridge university press, cambridge. statsoft, 2013, statistica version 12. statistica inc., http://www.statsoft.com tutchings, a., fennessy, s., marais, a. & fennessy, j., 2013, africa’s giraffe giraffa camelopardalis: a conservation guide, giraffe conservation foundation, viewed 03 december 2015, from http://www.giraffeconservation.org/booklets.php vorster, p.h., 2011, ‘the feeding and spatial ecology of cheetahs (acinonyx jubatus) and lions (panthera leo) in the little karoo’, msc dissertation, rhodes university, south africa. article information authors: hanlie malherbe1 michael samways1 affiliations: 1department of conservation ecology and entomology, stellenbosch university, south africa correspondence to: hanlie malherbe postal address: hochschule weihenstephan-triesdorf, hans-carl-von-carlowitz-platz 3, 85354 freising, germany dates: received: 21 oct. 2013 accepted: 07 jan. 2014 published: 23 may 2014 how to cite this article: malherbe, h. & samways, m., 2014, ‘rocky shores of a major southern african marine protected area are almost free from intertidal invertebrate alien species’, koedoe 56(1), art. #1206, 5 pages. http://dx.doi.org/10.4102/ koedoe.v56i1.1206 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. rocky shores of a major southern african marine protected area are almost free from intertidal invertebrate alien species in this short communication... open access • abstract • introduction • research method and design    • study area    • data collection and analyses • results and discussion • conclusion • acknowledgments    • competing interests    • authors' contributions • references abstract top ↑ a major threat to marine ecosystems is the establishment and proliferation of invasive alien species. this study addresses gaps in our knowledge regarding marine alien invertebrate species in the kogelberg biosphere reserve (kbr) and adjacent betty’s bay marine protected area (mpa) in the western cape of south africa, together a potentially important area for south-coast marine conservation. understanding the distribution and geographical expansion of these species is critical for conservation planning. a quantitative systematic survey of the intertidal rocky shore region was undertaken. the mytilid mediterranean mussel, mytilus galloprovincialis, and the bryozoan watersipora subtorquata were the only alien species recorded along the coastline, which included the mpa. the abundance of m. galloprovincialis was significantly higher outside the mpa, and the abundance of w. subtorquata was significantly higher inside the mpa. with only two alien species recorded, the betty’s bay mpa and its surroundings support relatively few marine alien species with regards to rocky shore invertebrate biodiversity. conservation implications: it is important that the betty’s bay mpa and its adjacent coastline maintain its current status as an area with relatively few marine alien species. the conservation implications on management require routine surveys of this region to detect early introductions of any additional species. introduction top ↑ the appearance of invasive alien species is a threat to marine reserves and marine protected areas (mpas). under favourable conditions, coastal invaders spread quickly over extensive geographical ranges, occasionally reaching marine reserves and mpas (grosholz 2002). once a species becomes established and spreads, it may have adverse effects on the indigenous marine biodiversity and marine communities (bax et al. 2003; molnar et al. 2008; padilla & williams 2004). for example, the collapse of the bivalve fisheries industry of eastern north america resulted from the introduction of the european shore crab carcinus maenas. a halt in the production of san francisco bay’s, usa phytoplankton arose from the invasion of the asian clam potamocorbula amurensis (grosholz 2002). furthermore, the introduction of the comb jellyfish mnemiopsis leidyi in the black sea resulted in a rapid decrease of phytoplankton blooms and consequently the collapse of various fish stocks (shiganova 1998). in south africa, the invasion of the mytilid mediterranean mussel, mytilus galloprovincialis, has substantially altered certain local mussel beds, increasing mussel biomass along the coastline (robinson et al. 2005) and displacing the indigenous mussels aulacomya ater and choromytilus meridionalis (hockey & van erkom schurink 1992). mytilus galloprovincialis is considered the most invasive marine invertebrate species in south africa. however, on a positive note, the endemic african black oystercatcher, haematopus moquini, has adapted its diet to mainly feed on m. galloprovincialis. this has enabled a significant recovery of the population of h. moquini (hockey & van erkom schurink 1992). also, local human communities along the west coast of south africa have reaped benefits from the introduction of m. galloprovincialis, with a small-scale mussel culturing industry now in operation along this coast. rocky shore invertebrate alien species have appeared along the south african coastline. some of these have established and spread in their new home range, invading marine reserves and mpas, and their surroundings (griffiths, mead & robinson 2009a; laird & griffiths 2008; robinson, griffiths & kruger 2004; robinson et al. 2005). the betty’s bay mpa and its surroundings are susceptible to such invasive species. it is therefore critical to have an understanding of the current status of this region. the aim of this study is to specifically fill gaps in our knowledge of the status of intertidal rocky shore invertebrate alien species of the betty’s bay mpa and surroundings, and to determine the extent of invasion of these species in the region. research method and design top ↑ study area this study was conducted in the kogelberg biosphere reserve (kbr) and the adjacent mpa, 90 km south-east of cape town in the western cape province, south africa. the kbr coastline is diverse, characterised by sandy beaches, rocky shores, estuary runoffs, subtidal kelp forests and reefs. the betty’s bay mpa was first declared in 1981 as the h.f. verwoerd marine reserve (tunley 2009). under the marine living resources act (act no. 18 of 1998), the marine reserve was proclaimed the betty’s bay mpa in 2000. seven principal rocky shore research sites were selected, and are referred to here as kogel bay (34º14’40”s; 18º51’11”e), rooi els (34º17’57”s; 18º48’59”e), pringle bay (34º20’52”s; 18º49’29”e), stony point (34º22’14”s; 18º53’41”e), betty’s bay main beach (34º22’01”s; 18º53’58”e), jock’s bay (34º21’26”s; 18º55’59”e) and kleinmond (34º20’44”s; 18º59’34”e). kogel bay, rooi els and pringle bay are located to the west of the mpa; stony point, betty’s bay main beach and jock’s bay are located within the mpa; and kleinmond is located to the east of the mpa (figure 1). distances between the study sites varied considerably due to interspersed sandy beaches and estuary runoffs. the longest distance, between kogel bay and rooi els, was 7 km, and the shortest distance, between stony point and betty’s bay main beach, was 800 m. it was evident that a wave exposure gradient exists along the kbr coastline. stony point, betty’s bay main beach and pringle bay were recognised as sheltered sites, jock’s bay and kleinmond were recognised as semi-exposed sites, and kogel bay and rooi els were recognised as exposed sites. it was also observed that the coastline of the mpa is sheltered compared to the coastline to the west and east of the mpa. figure 1: location of study sites along the kogelberg biosphere reserve coastline, western cape province, south africa. data collection and analyses transects were established from low to high shore at each of the seven selected study sites. the number of transects per site varied depending on the size of the site. the greatest number of transects at a specific site was nine, and the least was three. a total number of 42 transects were used, of which 20 were inside the mpa, and 22 outside. six intertidal 1 m2 sampling units (sus) were stratified per entire transect from low to high shore of the intertidal zone. zonation of rocky shores is subjective. however, the high-, midand low shore along the kbr supports a unique composition of rocky shore marine invertebrate species living in rock pools or on bare rock surfaces. sampling units were designated to rock pools and bare rock surfaces. the number of sus per shore region differed to include all rocky shore marine invertebrate species in the scope of the study. two rock pool sus and one rock surface su were allocated to the low shore. one rock pool su and one rock surface su were allocated to the mid shore. one rock pool was allocated to the high shore. the sus of the low shore were referred to as low tide pool seaward, low tide pool leeward and low tide rock surface. the sus of the mid shore were referred to as mid tide pool and mid tide rock surface. the su of the high shore was referred to as high tide pool.quantitative systematic sampling was done from october 2010 to december 2010. in order not to overlook any alien species, an in situ visual count of all focal rocky shore marine invertebrate species, except the phylum annelida, order isopoda, and the order amphipoda, was done at each su. in the case of colony-forming species, a count value was allocated to the area covered by the species. for each 1 cm2 a value of one was allocated. the counting method for each species was consistent. the time spent at each su depended on the time needed to count each species. some rock pool species prefer to reside under loose rocks. loose rocks were always present in the low tide pool seaward, low tide pool leeward and the mid tide pool. no more than three medium-sized rocks in each of the latter rock pools were overturned. in the case of smaller rocks, two rocks were overturned and counted as one medium-sized rock. generalized linear models (glms) were used to firstly test for significant differences in mean abundance of the identified marine alien species across the mpa and non-mpa (nmpa), and, secondly, to test for a significant difference across the study sites (o’hara 2009; zuur, ieno & elphick 2010). to test for a significant difference in mean abundance across the mpa and nmpa, all the sus in the mpa were pooled together, and all the sus in the nmpa were pooled together. to test for a significant difference in mean abundance across the study sites, all the sus per study site were pooled together. these glms were calculated with normal distribution (with identity link functions) for all data, as means were > 5, and the minimum number of successes and failures were < 5 (bolker et al. 2009). as these analyses showed no over-dispersion of variances compared to the models, wald χ2 (z), statistics were calculated using the penalised quasilikelihood technique (bolker et al. 2009). statistical analysis software (sas) package (statistical analysis software 2006) version 9.1.3 was used to run the analyses. results and discussion top ↑ only two marine alien invertebrate species, the alien mediterranean mussel, m. galloprovincialis, and the alien bryozoan watersipora subtorquata, were recorded along the rocky shores of the kbr. it is evident that m. galloprovincialis was not evenly distributed along the rocky shores of the kbr. mean abundance of m. galloprovincialis was significantly greater outside the mpa (18.3 m-2) than inside (no individuals recorded) (df = 1, z = 4.38, p ≤ 0.05, figure 2). figure 2: abundance of mytilus galloprovincialis in the marine protected area and non-marine protected area. there was a significant difference in mean abundance of m. galloprovincialis across the study sites, with a significantly greater abundance at kogel bay than at any other sites (df = 6, z = 25.13, p ≤ 0.001, figure 3). it has been documented that the level of wave exposure affects the distribution and abundance of m. galloprovincialis, which is characteristic at semi-exposed and exposed sites (blamey & branch 2009). this appears to be the case along the kbr coastline since the rocky shores, with the exception of pringle bay, is subjected to a greater wave exposure outside the mpa, particularly at kogel bay and rooi els. in all likelihood, the abundance of m. galloprovincialis could be higher along the coastal region to the west and east of the mpa and at the study site jock’s bay located at the western boarder of the mpa than was measured, since by chance the selected study sites and sampling units were placed at areas in which m. galloprovincialis was absent. in contrast to m. galloprovincialis, w. subtorquata was significantly more abundant inside the mpa (df = 1, z = 5.97, p ≤ 0.05) with a mean abundance of 17.3 m-2 compared to 5.5 m-2 outside the mpa (figure 4). there was no significant difference in mean abundance of w. subtorquata across the study sites (df = 6, z = 8.70, p > 0.05). from the former finding, it could be concluded that this species prefers the sheltered conditions of the mpa. however, there is no information on the effects of wave exposure on w. subtorquata in south africa. the difference in the abundance of w. subtorquata along the kogelberg biosphere reserve coastline was in all likelihood from the integrated functioning of various factors. figure 3: abundance of mytilus galloprovincialis at the different study sites. figure 4: abundance of watersipora subtorquata in the marine protected area and non-marine protected area. various introductions of marine alien invertebrate species and invasions along the coast of south africa have been documented. mytilus galloprovincialis, the european shore crab c. maenas, and the periwinkle littorina saxatilis have naturalised outside the boundaries of the saldanha bay harbour situated in the west coast national park (wcnp) marine reserve, south africa. the reserve forms part of the saldanha bay–langebaan lagoon system. a survey conducted by robinson et al. (2004) indicated that the abundance of m. galloprovincialis within the wcnp was low; l. saxatilis was confined to the lagoon and salt marshes, the alien anemone sagartia ornata was confined to the lagoon, and no live individuals of c. maenas were recorded. other identified marine alien invertebrate species such as the ascidians diplosoma listerianum, ciona intestinalis and ascidiella aspersa are confined to the harbour. it is also recognised that the saldanha bay–langebaan lagoon system supports the most marine alien species along the south african coast (robinson et al. 2004). a survey conducted by robinson et al. (2005) indicates that m. galloprovincialis is present along the west coast, south coast and east coast, almost reaching east london. significant populations of the pacific barnacle balanus glandula have been detected along the west coast (griffiths et al. 2009a, 2009b) from misty cliffs to elands bay (laird & griffiths 2008). the west coast supports a prime example of upwelling. pineda and lópez (2002) found that the settlement of b. glandula larvae increased with high-frequency internal motion. this may explain the presence of this species along the cool waters of west coast and its absence along the kbr coastline. watersipora subtorquata has been recorded along the coast of the western and eastern cape provinces (branch et al. 2010). the former and latter geographical regions are interspersed with marine reserves and mpas, which suggests that these coastal areas of conservation may be invaded by marine alien invertebrate species. since many marine and estuarine alien species in south africa have been overlooked (mead et al. 2011), the current status of marine alien invertebrate species along the shores of south africa, including marine reserves and mpas, may be different. this is a challenge that must be addressed. with only two alien species recorded along the rocky shores of the kbr, it is evident that the betty’s bay mpa supports relatively few marine alien species relating to rocky shore invertebrate biodiversity, particularly compared to the well-studied wcnp. the inclusion of the saldanha bay harbour in the wcnp makes this conservation region much more susceptible to the invasion of marine alien invertebrate species. in the case of the betty’s bay mpa, it has more sheltered shores, which do not support the proliferation of m. galloprovincialis. it is suggested that the sheltered nature of the betty’s bay mpa should be considered the primary reason for its good condition regarding m. galloprovincialis invasion. it is therefore not considered feasible to implement marine reserves and mpas to decrease the geographical extent of marine alien species. marine alien species will potentially invade any favourable region along the coast. identifying and eliminating pathways is considered to be the only practical method in preventing possible introductions and proliferation of marine alien species. prevention is therefore critical for effective conservation (bax et al. 2003). it is of concern that many mpas in south africa, including the betty’s bay mpa, function as isolated units. it is not viable to focus only on isolated regions of this highly integrated and vast open system. managing large, multi-purpose mpas is of great interest as it provides opportunities to conserve highly mobile species, and protects demarcated regions from pollution and unwanted negative impacts from human activities (hoegland, sumaila & farrow 2001). it is also suggested that habitat heterogeneity must be considered when selecting priority areas for coastal conservation. a greater diversity of rocky shore species are conserved when applying the habitat heterogeneity approach rather than the hotspot approach. furthermore, the level of wave exposure plays an important role in defining the habitat types of rocky shores (blamey & branch 2009). the sheltered nature of the betty’s bay mpa may lack specific habitat types, therefore decreasing the species richness being protected. this emphasizes the importance of taking a more holistic management approach (mora et al. 2006). in 2010, only 9% of all south africa’s mpas were declared as no-take mpas (griffiths et al. 2010), although shore angling is a resource utilization activity that is allowed in the betty’s bay mpa (tunley 2009). it is evident that the betty’s bay mpa will function more adequately if managed as a large, multi-purpose mpa. this protected coastal region unfortunately experiences much pressure from overexploitation, development (tunley 2009) and abalone poaching (stanvliet et al. 2004). a current study is underway to investigate the status of fish communities inside and outside the mpa. it should, however, be noted that an african penguin spheniscus demersus colony (a red list species), as well as other species, particularly the abalone haliotis midae and the west coast rock lobster, jasus lalandii, are protected within the mpa (tunley 2009). conclusion top ↑ with only two alien species, the mytilid mussel, m. galloprovincialis, and the bryozoan w. subtorquata, being recorded, this study revealed that the southern african coastline around the betty’s bay mpa has been invaded by relatively few marine alien species with regards to rocky shore invertebrate biodiversity. fortunately, the abundance of these two species in the betty’s bay mpa is relatively low. the mpa is therefore in good condition regarding intertidal invasive alien species. marine introductions and invasions are considered irreversible (bax et al. 2003) and eradication of these two species along this coastline is not viable. nevertheless, we recommend routine surveys of this region are conducted to detect early introductions of any additional species. acknowledgments top ↑ we thank capenature and the department of environmental affairs, marine coastal management, for the provision of research permits (danelle kleinhans, dkleinhans@capenature.co.za, ref. 1/2/1/6/5/f8, permit no. aaa-004-00481-0035). we also thank c.l. griffiths (department of zoology, university of cape town – marine research institute) for assistance in the identification of specimens, and j.s. pryke (department of conservation ecology and entomology, stellenbosch university) for assistance with statistical analyses. m.j. samways acknowledges funding from the national research foundation. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions h.m. (university of stellenbosch) designed the methods, carried out the field surveys, interpreted the results, and wrote the first draft of the manuscript. m.j.s. (university of stellenbosch) assisted with the design of the methods and co-prepared the manuscript. references top ↑ arcgis version 10.1, 2012, computer software, esri, redlands.bax, n., williamson, a., aguero, m., gonzales, e. & geeves, w., 2003, ‘marine invasive alien species: a threat to global biodiversity’, marine policy 27, 313–323. http://dx.doi.org/10.1016/s0308-597x(03)00041-1 blamey, l.k. & branch, g.m., 2009, ‘habitat diversity relative to wave action on rocky shores: implications for the selection of marine protected areas’, aquatic conservation marine and freshwater ecosystems 19, 645–657. http://dx.doi.org/10.1002/aqc.1014 bolker, b.m., brooks, m.e., clark, c.j., geange, s.w., poulsen, j.r., stevens, m.h. et al., 2009, ‘generalized linear mixed models: a practical guide for ecology and evolution’, trends in ecology & evolution 24, 127–135. http://dx.doi.org/10.1016/j.tree.2008.10.008 branch, g.m., griffiths, c.l., branch, m.l. & beckley, l.e., 2010, two oceans – a guide to the marine life of southern africa, struik nature, cape town. griffiths, c.l., mead, a. & robinson, t.b., 2009a, ‘a brief history of marine bio invasions in south africa’, african zoology 44, 241–247. http://dx.doi.org/10.3377/004.044.0212 griffiths, c.l., robinson, t.b. & mead, a., 2009b, ‘the status and distribution of marine alien species in south africa’, ecological studies 204, 393–408. griffiths, c.l., robinson, t.b., lange, l. & mead, a., 2010, ‘marine biodiversity in south africa: an evaluation of current states of knowledge’, plos one 5, 1–13. http://dx.doi.org/10.1371/journal.pone.0012008 grosholz, e., 2002, ‘ecological and evolutionary consequences of coastal invasions’, trends in ecology & evolution 17, 22–27. http://dx.doi.org/10.1016/s0169-5347(01)02358-8 hockey, p.a.r. & van erkom schurink, c., 1992, ‘the invasion biology of the mussel mytilus galloprovincialis on the southern-african coast’, transactions of the royal society of south africa 48, 123–139. http://dx.doi.org/10.1080/00359199209520258 hoegland, p., sumaila, u.r. & farrow, s., 2001, marine protected areas, academic press, massachusetts. laird, m.c. & griffiths, c.l., 2008, ‘present distribution and abundance of the introduced barnacle balanus glandula darwin in south africa’, african journal of marine science 30, 93–100. http://dx.doi.org/10.2989/ajms.2008.30.1.9.459 mead, a., carlton, j.t., griffiths, c.l. & rius, m., 2011, ‘introduced and cryptogenic marine and estuarine species of south africa’, journal of natural history 45, 2463–2524. http://dx.doi.org/10.1080/00222933.2011.595836 molnar, j.l., gamboa, r.l., revenga, c. & spalding, m.d., 2008, ‘assessing the global threat of invasive species to marine biodiversity’, frontiers in ecology and the environment 6, 485–492. http://dx.doi.org/10.1890/070064 mora, c., andrefouet, s., costello, m.j., kranenburg, c., rollo, a., veron, j. et al., 2006, ‘coral reefs and the global network of marine protected areas’, science 312, 1750–1751. http://dx.doi.org/10.1126/science.1125295 o’hara, r.b., 2009, ‘how to make models add up – a primer on glmms’, annales zoologici fennici 46, 124–137. http://dx.doi.org/10.5735/086.046.0205 padilla, d.k. & williams, s.l., 2004, ‘beyond ballast water: aquarium and ornamental trades as sources of invasive species in aquatic ecosystems’, frontiers in ecology and the environment 6, 485–492. pineda, j. & lópez, m., 2002, ‘temperature, stratification and barnacle larval settlement in two californian sites’, continental shelf research 22, 1183–1198. http://dx.doi.org/10.1016/s0278-4343(01)00098-x robinson, t.b., griffiths, c.l. & kruger, n., 2004, ‘distribution and status of marine invasive species in and bordering the west coast national park’, koedoe 47, 79–87. http://dx.doi.org/10.4102/koedoe.v47i1.73 robinson, t.b., griffiths, c.l., mcquiad, c. & rius, m., 2005, ‘marine alien species of south africa – status and impacts’, african journal of marine science 27, 297–306. http://dx.doi.org/10.2989/18142320509504088 statistical analysis software version 9.1., 2006, computer software, sas institute inc., cary, north california. shiganova, t.a., 1998, ‘invasion of the black sea by the ctenophore mnemiopsis leidyi and the recent changes in pelagic community structure’, fisheries oceanography 7, 305–310. http://dx.doi.org/10.1046/j.1365-2419.1998.00080.x stanvliet, r., jackson, j., davis, g., de swardt, c., mokhoele, j., thom, q. et al., 2004, ‘the unesco biosphere reserve concept as a tool for urban sustainability – the cubes cape town case-study’, urban biosphere and society: partnership of cities 1023, 80–104. tunley, k., 2009, ‘state of management of south africa’s marine protected areas’, in wwf south africa report series, viewed december 2013, from http://assets.wwfza.panda.org/downloads/mpastateofmanagementreport04nov2009weblowerdpi.pdf zuur, a.f., ieno, e.n. & elphick, c.s., 2010, ‘a protocol for data exploration to avoid common statistical problems’, methods in ecology and evolution 1, 3–14. http://dx.doi.org/10.1111/j.2041-210x.2009.00001.x article information authors: nicola j. van wilgen1 mbulelo dopolo1 alexis symonds2 wessel vermeulen3 elzette bester4 kyle smith5 melodie a. mcgeoch1,6 affiliations: 1cape research centre, south african national parks, south africa2conservation services, south african national parks, south africa 3knysna scientific services, south african national parks, south africa 4storms river village conservation services, south african national parks, south africa 5wilderness scientific services, south african national parks, south africa 6centre for invasion biology, south african national parks, south africa correspondence to: nicola van wilgen postal address: po box 216, steenberg 7947, south africa dates: received: 12 june 2012 accepted: 20 nov. 2012 published: 30 may 2013 how to cite this article: van wilgen, n.j., dopolo, m., symonds, a., vermeulen, w., bester, e., smith, k. & mcgeoch, m.a., 2013, ‘an inventory of natural resources harvested from national parks in south africa’, koedoe 55(1), art. #1096, 5 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1096 note: additional supporting information may be found in the online version of this article as online appendixes: http://dx.doi.org/10.4102/ koedoe.v55i1.1096-1. copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. an inventory of natural resources harvested from national parks in south africa in this checklist... open access • abstract • introduction • methods • results and discussion • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ resource harvesting is permissible within south african protected areas under certain conditions as part of benefit sharing that seeks to strengthen relationships with communities living adjacent to parks. however, not all resource use is authorised and little is currently known about what is harvested, or the extent and impacts of harvesting in parks. this limits capacity to monitor and set the boundaries for such use. this paper provides a checklist of resources harvested within each of 19 national parks managed by south african national parks. data were gathered by means of a question-based survey of park staff. a database detailing the parks from which each resource was harvested and its purpose(s) was compiled, representing the most comprehensive list of resources harvested from parks to date. a total of 382 harvested biological and abiotic resources (284 terrestrial and 98 aquatic), used for a wide range of purposes, were identified across parks. many of the resources, especially animals (96%), were harvested destructively. the strongest motivation for harvest was subsistence, although most resources were also used for financial gain through informal business. although current data are not sufficient to determine harvest sustainability for most resources, better data and increased awareness of resource use activities will enable future research to this end. conservation implications: the checklist of harvested resources provides critical baseline data for parks, which will facilitate assessment of park-specific priorities for research, monitoring and management action. introduction top ↑ to be effective, protected areas need to succeed in conserving biodiversity whilst providing livelihood opportunities that safeguard continued socio-economic benefits (mcneely 1993; salomon et al. 2011). the overharvesting of resources was identified through the millennium ecosystem assessment (millennium ecosystem assessment 2005) as a driver of global environmental change, which poses a threat to biodiversity and the people dependent on it. overharvesting is not limited to areas outside of formal protection, but can also happen within protected areas, especially when resources become scarce outside of such areas (agardy et al. 2003; castley & kerley 1996; naughton-treves et al. 2006; waite 2007). increasing pressures on both local and global natural resources, stakeholder rights and the need for development in many countries have led to people’s needs becoming a core consideration in conservation, with the recognition that without the support of local people, conservation efforts are likely to fail (naughton-treves, buck holland & brandon 2005; zaccagnini et al. 2001). this is true also in south africa, where many people rely directly on natural resources for their livelihoods and even survival (dovie, shackleton & witkowski 2007; shackleton & shackleton 2004; shackleton et al. 2007). however, how best to ensure the balance between sustainable resource use and conservation of biodiversity remains uncertain (mcshane et al. 2011; robinson 2011; salafsky 2011). south african national parks (sanparks) have a resource use policy aimed at ensuring sustainable use of biological resources for the benefit of local communities (sanparks csd policy unit 2010). the policy is guided by both south african national legislation (the national environmental management: protected areas act [republic of south africa 2003] and its accompanying regulations) and international best practice (secretariat of the convention on biological diversity 2004). however, managing resource use and determining (or even defining) whether or not use is sustainable is not straightforward in practice (callicott & mumford 1997; chapman, de lacey & whitmore 2006; hardin 1968). although challenges range from the development of sustainable harvest systems (e.g. hall & bawa 1993; seydack 1995; vermeulen 2009) to unrealistic expectations with regard to benefits (especially commercial, as discussed by belcher and schreckenberg [2007]) or unrealistic promotion of access to resources by policy makers, one of the most prevalent problems is the lack of, or insufficient, data to estimate factors necessary for determining sustainable use thresholds. these factors include the species harvested, their distribution and life histories, harvest practices and quantities extracted (ndanyalasi, bitariho & dovie 2007; ticktin 2004; williams & kepe 2008). to begin to overcome this challenge, several sets of baseline data are required. in this paper, we detail a checklist of resources that are harvested from each of the 19 south african national parks managed by sanparks as one of the first steps towards addressing these knowledge gaps. this first checklist of harvested species will provide (1) a basis for identifying those resources that may be threatened and that should not be harvested, or that require stricter control or alternative harvest practices, (2) a list from which to prioritise species for further study and (3) an important baseline for monitoring resource use in national parks (mcgeoch et al. 2011). methods top ↑ data were collected using a questionnaire that was completed by park management personnel and sanparks scientists in each park. the questionnaire was developed by scientists and policy makers in the conservation services division of sanparks, with the aim of gathering information on natural resources currently harvested from the sanparks estate. the questionnaire covered only harvestable resources, or consumptive resource use; that is, all activities where a tangible benefit (economic, social, cultural or ecological) is gained from a resource, with all or part of the resource removed. non-consumptive use, which includes a range of tourism activities, was not considered. the questionnaire consisted of four sections, one each for terrestrial, aquatic and abiotic resources and one section for resources harvested and made available for use through management actions (online appendix 1). within each section a number of potential resource use purposes (e.g. animals used for medicinal or ritual purposes, plants used for medicinal or ritual purposes, or animals used as a food source) were listed to ensure that those tasked with data compilation had a common understanding of the full range of resource uses of interest. an open-ended option (‘other’) was included in each section to accommodate information not considered during the design of the questionnaire. for each resource listed, the respondents were further required to answer a series of questions pertaining to the purpose, authorisation, user group and quantity of the resource used. for the questions where data availability was likely to be limited, a range of possible answers were provided to allow comparable estimates of use (online appendix 1). use of these categorical ranks to score quantities was reserved for when no other information was available and their use was not encouraged otherwise. blank questionnaires were sent to each of the 19 national parks and instructions for their completion were communicated telephonically and via email. responses were collated and a follow-up questionnaire (re-survey) was sent to all parks one year later to allow for revision or addition of further information. where possible (n = 7), this was followed up with face-to-face meetings with park managers and rangers where the listed resources and possible omissions were discussed. these meetings took the form of round-table discussions between park managers, section rangers and scientists, with data being displayed and updated during the meeting. the completed questionnaire for each park was therefore the outcome of collective rather than individual contributions. once all data had been cleaned and checked, a single database containing a full list of resources used in sanparks was compiled. the database included the parks from which each resource was reported as being harvested, the parts of the resource harvested, and the motivation(s) and purpose(s) for its harvest. where relevant and possible, species names were assigned to the resource used. scientific names were verified using relevant literature and official species databases such as the plant list (the plant list 2010), the world register of marine species (costello et al. 2012) and the encyclopedia of life (encyclopedia of life 2008). because multiple common names are often used for a single species, especially in different languages, the common names used by those contributing to questionnaire completion were retained. these common names are those likely to be most widely used in park-specific contexts. purposes of resource use, as detailed in the responses from each park, were grouped into ten categories as follows: • bait • construction • food • fuel • grazing • handcrafts or decorations • medicinal or ritual use • ornamental plants (including cut flowers) or pets • thatching or weaving material • timber. in addition, four motivations for harvesting resources were identified, namely financial gain, subsistence (including all personal use), park management (including alien species removal and population management through culling or live sale) and recreation (muth & bowe 1998). results and discussion top ↑ results showed that a wide range of species and resources are harvested from parks for a variety of purposes. the final checklist of harvested resources comprised 382 resources, of which 289 were identified to species level, 46 to genus level and 40 to family level or higher (e.g. blood worms, a polychaete species). the remainder were non-specific (six abiotic resources and grazing; see online appendix 2). of the 382 resources used, 284 (74%) were terrestrial and 98 (26%) were aquatic (90 marine and 8 freshwater; see online appendix 2). the abiotic resources harvested included gravel, rocks, salt, soil or sand, water and diamonds, with the latter being from the richtersveld national park. the biological resources included 194 plants, 179 animals, mushrooms (not identified to species level) and two seaweed species. these species belonged to 152 families, of which 92 families had only one species harvested, whilst 27 families had more than two species harvested and six families had 10 or more species harvested. the family from which the most species were harvested (22 species) was the bovidae (even-toed ruminant ungulates), followed by the sparidae (a group of marine fish), with 19 species harvested. the most commonly used plant families were the fabaceae (17 species) and the asteraceae (16 species). in addition, 26 alien species were also noted as being harvested (6.8% of all resources). products derived from alien species harvest are considered positive spin-offs, where, unlike other harvested species, the aim is not sustainable harvest but rather eradication the over medium to long term. although survey respondents did not always specify which parts of a resource were used, 95% of resources in the final list had at least one ‘part’ specified (online appendix 3). based on the parts reported to be harvested, nearly all cases where animals were harvested would have resulted in the death or removal of the animal from the national park. for 95.5% of listed animal resources, the whole animal was harvested, whilst specific parts were listed for 7% of these resources. these parts, as well as the parts of the remaining 4.5% of animal resources, included body parts, bones, dung, eggs, fat, feathers, honey, horns, quills, shells, skin, talons and tusks. harvesting of some of these parts (e.g. bones and horns) may require that the animal be killed, but they could be harvested from animals that are already dead, whilst other parts (such as dung, feathers or quills) would not require the death of the animal. only 24.2% of the listed plant species were harvested as whole plants or in a manner that could result in the death of the plant (e.g. timber harvest where the entire above-ground portion of the plant is cut down to obtain the useable parts, although for some species coppicing does allow for regeneration; see kaschula, twine & scholes 2005). it was more difficult to determine whether the other uses of plants were destructive. use of plants for medicinal purposes often included harvest of the roots, corms or bulbs of plants, which, depending on the method and extent of harvesting, could result in the death of the plants. harvest of bark, branches, stems and leaves may also result in the death of plants, but in many instances stems and branches were harvested as fuel, which may have been from dead specimens and often included use of non-native species. use of other plant parts, for example the use of flowers and seeds and even foliage, may not result in the death of plants, but could affect their reproductive output (gaoue & ticktin 2008; peters 1999). recreation (a motivation for 19% of harvested resources) and park management (13%) were less frequent motivators than subsistence or harvest for financial gain. however, the way in which the survey was designed required that financial gain versus subsistence (or both) be specified as a motivation for all harvested resources. therefore, resource harvest motivated by park management or recreation would also have been counted as having a subsistence or financial motivation, albeit secondary rather than primary. most harvested resources (49%) were used for both subsistence and financial gain (usually in the form of small informal businesses, especially traditional medicines), whilst nearly a third (30%) were used only for subsistence (online appendix 2). although 21% of resources were used solely for financial gain, large-scale commercial harvesting was limited to the marine sector and timber in two parks (garden route and table mountain), which included the harvest of non-native pines and blackwood. commercial harvesting was mostly authorised, with the exception of abalone and rhino poaching. unauthorised resource use was fairly common, with 42% of reported resources harvested without authorisation (across all 19 parks) and a further 36% of resources harvested (1) with authorisation only some of the time, (2) in some parks but not others, or (3) in contravention of authorised limits. despite these figures, seven of the 19 parks reported that no unauthorised resource use took place within the park. whilst this is likely to be true for the very isolated parks, the levels of unauthorised resource use reported from the other 12 parks (44.2% of all resources used in these parks) suggest that further investigation may reveal additional resource use activities. although the checklist presented here represents a valuable first step in documenting resource use across national parks, it is likely to be incomplete. the current list is based on knowledge supplied only by the management and scientists of protected areas. caveats exist particularly for unauthorised cases of resource use, which are known to be substantial in some parks, particularly those with open access (petersen et al. 2012; van wilgen & mcgeoch in preparation). the level of effort associated with data collection varied across the country, with surveys of parks in the cape provinces being more comprehensive than those in other provinces (due to the feasibility of re-visiting some parks for follow-up). nonetheless, all parks (with the exception of table mountain and tankwa karoo) were surveyed twice, approximately one year apart. very little information emerged on the harvest quantities of each resource (also see van wilgen & mcgeoch [in preparation]) and additional information on life stage (for biological resources), quantity, frequency and extent of harvesting will be critical to the future assessment of sustainable yields (botsford, castilla & peterson 1997; goodland & daly 1996; waite 2007). in general there is a paucity of published research on resource extraction from protected areas in south africa (petersen et al. 2012; shackleton 2009) and the outcome of the current survey reveals that this is also true of national parks. this checklist will contribute not only as a baseline for further work but also to increasing awareness within south african protected area agencies about the role that resource use plays and the information and actions required to sustainably manage resources. more comprehensive and detailed information will improve the effectiveness with which parks are able to manage authorised use and promote strategies to limit unauthorised activities. acknowledgements top ↑ we would like to thank the park management staff for their interest and participation in data collection and the sanparks regional ecologists (angela gaylard, carly cowell, cathy greaver, marna herbst and michael radzilani) who assisted in contacting parks and following up on questionnaire completion. in addition, we thank louise swemmer and especially wendy annecke for discussion on resource use in protected areas in the social context and referral to key papers. the work was funded by the andrew w. mellon foundation and the sanparks park development fund, and was conducted under the auspices of the sanparks global environmental change project. two anonymous referees provided useful comments on an earlier version of the manuscript. competing interests the authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article. authors’ contributions n.j.v.w. (sanparks) was responsible for data collation, analysis and compilation of results. m.d. (sanparks), w.v. (sanparks), a.s. (sanparks) and m.a.m. (sanparks) were responsible for the design of the original survey and data collection. e.b. (sanparks) and k.s. (sanparks) contributed research results from the garden route national park. m.a.m. also made conceptual contributions to the manuscript. references top ↑ agardy, t., bridgewater, p., crosby, m.p., day, j., dayton, p.k.r., laffoley, k. et al., 2003, ‘dangerous targets? unresolved issues and ideological clashes around marine protected areas’, aquatic conservation: marine and freshwater ecosystems 3, 353–367.belcher, b. & schreckenberg, k., 2007, ‘commercialisation of non-timber forest products: a reality check’, development policy review 25, 355–377. http://dx.doi.org/10.1111/j.1467-7679.2007.00374.x botsford, l.w., castilla, j.c. & peterson, c.h., 1997, ‘the management of fisheries and marine ecosystems’, science 277, 509–515. http://dx.doi.org/10.1126/science.277.5325.509 callicott, j.b. & mumford, k., 1997, ‘ecological sustainability as a conservation concept’, conservation biology 11, 32–40. http://dx.doi.org/10.1046/j.1523-1739.1997.95468.x castley, j.g. & kerley, g.i.h., 1996, ‘the paradox of forest conservation in south africa’, forest ecology and management 85, 35–46. http://dx.doi.org/10.1016/s0378-1127(96)03748-6 chapman, j., de lacey, t. & whitmore, m., 2006, ‘sustainability practice and sustainable use’, in m. lockwood, g.l. worboys & a. kotharis (eds.), managing protected areas: a global guide, pp. 377–405, earthsan, uk and usa. costello, m.j., vandepitte, l., appeltans, w., bouchet, p., boxshall, g., gordon, d. et al. (eds), 2012, world register of marine species, viewed 12 june 2012, from http://www.marinespecies.org dovie, d.b.k., shackleton, c.m. & witkowski, e.t.f., 2007, ‘conceptualizing the human use of wild edible herbs for conservation in south african communal areas’, journal of environmental management 84, 146–156. http://dx.doi.org/10.1016/j.jenvman.2006.05.017, pmid:17045732 encyclopedia of life, 2008, encyclopedia of life, viewed 12 june 2012, from http://www.eol.org gaoue, o.g. & ticktin, t., 2008, ‘impacts of bark and foliage harvest on khaya senegalensis (meliaceae) reproductive performance in benin’, journal of applied ecology 45, 34–40. http://dx.doi.org/10.1111/j.1365-2664.2007.01381.x goodland, r. & daly, h., 1996, ‘environmental sustainability: universal and non-negotiable’, ecological applications 6, 1002–1017. http://dx.doi.org/10.2307/2269583 hall, p. & bawa, k., 1993, ‘methods to assess the impact of extraction of non-timber tropical forest products on plant-populations’, economic botany 47, 234–247. http://dx.doi.org/10.1007/bf02862289 hardin, g., 1968, ‘the tragedy of the commons. the population problem has no technical solution; it requires a fundamental extension in morality’, science 162, 1243–1248. kaschula, s.a., twine, w.c. & scholes, m.c., 2005, ‘the effect of catena position and stump characteristics on the coppice response of three savannah fuelwood species’, environmental conservation 32, 76–84. http://dx.doi.org/10.1017/s0376892905001980 mcgeoch, m.a., dopolo, m., novellie, p., hendricks, h., freitag, s., ferreira, s. et al., 2011, ‘a strategic framework for biodiversity monitoring in south african national parks’, koedoe 53, art. #991, 10 pages. mcneely, j.a. (ed), 1993, parks for life: report of the 4th world congress on national parks and protected areas, iucn, gland. mcshane, t.o., hirsch, p.d., trung, t.c., songorwa, a.n., kinzig, a., monteferri, b. et al., 2011, ‘hard choices: making trade-offs between biodiversity conservation and human well-being’, biological conservation 144, 966–972. http://dx.doi.org/10.1016/j.biocon.2010.04.038 millennium ecosystem assessment, 2005, ‘ecosystems and human well-being: synthesis’, island press, washington dc. muth, r.m. & bowe, j.f., 1998, ‘illegal harvest of renewable natural resources in north america: toward a typology of the motivations for poaching’, society & natural resources 11, 9–24. http://dx.doi.org/10.1080/08941929809381058 naughton-treves, l., alvarez-berr, n., brandon, k., bruner, a., holland, m., ponce, c., et al., 2006, ‘expanding protected areas and incorporating human resource use: a study of 15 forest parks in ecuador and peru’, sustainability: science, practice, and policy 2, 32–44. naughton-treves, l., buck holland, m. & brandon, k., 2005, ‘the role of protected areas in conserving biodiversity and sustaining local livelihoods’, annual review of environment and resources 30, 219–252. http://dx.doi.org/10.1146/annurev.energy.30.050504.164507 ndanyalasi, h.j., bitariho, r. & dovie, d.b.k., 2007, ‘harvesting of non-timber forest products and implications for conservation in two montane forests of east africa’, biological conservation 134, 242–250. http://dx.doi.org/10.1016/j.biocon.2006.06.020 peters, c.m., 1999, ‘ecological research for sustainable non-wood forest product exploitation: an overview’, in t.c. sunderland, l.e. clark & p. vantommes (eds.), non-wood forest products of central africa: current research issues and prospects for conservation and development, pp. 19–363, fao, rome. petersen, l.m., moll, e.j., collins, r. & hockings, m., 2012, ‘development of a compendium of local, wild-harvested species used in the informal economy trade, cape town, south africa’, ecology & society 17, 1–26. republic of south africa, 2003, national environmental management: protected areas act, 57 of 2003, government printer, pretoria. robinson, j.g., 2011, ‘ethical pluralism, pragmatism, and sustainability in conservation practice’, biological conservation 144, 958–965. http://dx.doi.org/10.1016/j.biocon.2010.04.017 salafsky, n., 2011, ‘integrating development with conservation a means to a conservation end, or a mean end to conservation?’, biological conservation 144, 973–978. http://dx.doi.org/10.1016/j.biocon.2010.06.003 salomon, a.k., gaichas, s.k., jensen, o.p., agostini, v.n., sloan, n.a., rice, j. et al., 2011, ‘bridging the divide between fisheries and marine conservation science’, bulletin of marine science 87, 251–274. http://dx.doi.org/10.5343/bms.2010.1089 sanparks csd policy unit, 2010, ‘resource use policy’, policy document 17/p – csd/pol/resource use/03-10/vs1, csd policy unit, sanparks. secretariat of the convention on biological diversity, 2004, ‘addis ababa principles and guidelines for the sustainable use of biodiversity (cbd guidelines)’, secretariat of the convention on biological diversity, montreal. seydack, a.h.w., 1995, ‘an unconventional approach to timber yield regulation for multi-aged, multispecies forests. i. fundamental considerations’, forest ecology and management 77, (1–3), 139–153. http://dx.doi.org/10.1016/0378-1127(95)03577-w shackleton, c. & shackleton, s., 2004, ‘the importance of non-timber forest products in rural livelihood security and as safety nets: a review of evidence from south africa’, south african journal of science 100, 658–664. shackleton, c.m., 2009, ‘will the real custodian of natural resource management please stand up’, south african journal of science 105, 91–93. shackleton, c.m., shackleton, s.e., buiten, e. & bird, n., 2007, ‘the importance of dry woodlands and forests in rural livelihoods and poverty alleviation in south africa’, forest policy and economics 9, 558–577. http://dx.doi.org/10.1016/j.forpol.2006.03.004 the plant list, 2010, version 1. published on the internet, viewed 12 june 2012, from http://www.theplantlist.org ticktin, t., 2004, ‘the ecological implications of harvesting non-timber forest products’, journal of applied ecology 41(1), 11–21. http://dx.doi.org/10.1111/j.1365-2664.2004.00859.x van wilgen, n.j. & mcgeoch, m.a., in preparation, ‘ignorance as a risk to achieving the dual goals of resource use and conservation in protected areas’. vermeulen, w.j., 2009, ‘the sustainable harvesting of non-timber forest products from natural forests in the southern cape, south africa: development of harvest systems and management prescriptions’, phd thesis, department of conservation ecology and entomology, stellenbosch university. waite, t.a., 2007, ‘revisiting evidence for sustainability of bushmeat hunting in west africa’, environmental management 40, 476–480. http://dx.doi.org/10.1007/s00267-006-0207-9, pmid:17638049 williams, s. & kepe, t., 2008, ‘discordant harvest: debating the harvesting and commercialization of wild buchu (agathosma betulina) in elandskloof, south africa’, mountain research and development 28, 58–64. http://dx.doi.org/10.1659/mrd.0813 zaccagnini, m.e., cloquell, s., fernandez, e., gonzález, c., lichtenstein, g., novaro, a. et al., 2001, ‘analytic framework for assessing factors that influence sustainability of uses of wild living natural resources’, iucn susg technical advisory committee of the iucn species survival commission, washington dc. pmid:11407650 filelist convert a pdf file! koedoe 19: 43-48 (1976) sex reversal in the dageraad chrysoblephus cristiceps (pisces, sparidae) c . a. robinson tsitsikama coastal national park p.o . stormsrivier 6308 abstract this paper reports protogynous hermaphroditism in the dageraad chrysoblephus cristiceps. mass-length relationships indicate growth changes in male and female fish . seasonal maturity in dageraad, and hermaphroditism in south afrit:an spari<l fish are discu~ed. a regulatory meastlre for the conservation of dageraad is proposed. introduction the occurrence of natural sex reversal in south african sparid fish was first recorded by lucks (1970) in west coast steenbras lithognathus aureti and penrith (1972) in roman chrysoblephus laticeps. mehl (974) in a study on the white steenbras lithognathus lithognathus found this sparid to be permanently hermaphroditic. he postulated that it was a rudimentary hermaphrodite with only one of the two sexes functioning throughout the individual's life span and concluded that it could be a transitionary stage towards eventual gonochorism. chrysoblephus laticeps was shown to be a case of protogynous hermaphroditism while lithognathus aureti proved to be protandrous hermaphroditic. penrith (972) stated that although he did not have large fish in his samples, the data available suggested that sexual reversal in all probability took place. during a study of two submerged reefs in the inshore waters of the tsitsikama coastal national park this supposition was confirmed. the reefs currently being studied are known locally as middelbank and rheeder se knol. due to environmental factors presently being investigated chrysoblephus cristiceps are seldom i:¥en at middelbank although the depth of the reef is approximately the same (10-15 fathoms). almost exclusively the data on which this report is based came from rheeder se knol, situated six km east of the storms river mouth. material and methods samples were collected by hand-line fishing from an 5,5 m vessel. fish were mass measured to the nearest milligram and measured for total and caudal lengths to the nearest millimeter. macroscopic examination of the gonads was conducted after dissection. gonads were re43 corded on a 1 to 7 maturity index scale : 1 being inactive, 6 being ripe and running and 7 being spent. there appeared to be no external sexual dimorphism in chrysoblephus cristiceps, during the macroscopic examination of the gonads individuals of ten were found in a transformation stage i.e. with the male gonad developing and the female gonad degenerating. these fish were recorded according to the dominating gonad. the records included in the analyses number 510 fish and the majority were collected during 1973 and 1974. the standard exponential relationship between fish mass and length were cal<::ulated. this is expressed by the function: w=ali> where l = length in millimeters a = intercept of the y axis w = mass in milligrams b = exponent logarithmic transformations yield a linear relationship from which a and b can be calculated by the method ofleast-squares. ... '" ~ z 100~--------------~~--------------~~----------------~ 75r----------1~--------~------~------_*--------------~ ~ ~~--------~------------~--~----------~~----------~ ii: ... ... 25r--------.~-------------.~._--------------~._------~ 11 zi .. zt ii n " .5 '0 l length cem) fig. i. size composition of a dageraad population as obtained from handline catches in tsitsikama coastal national park. 44 results (i) by plotting the relative frequency of the size groups it is clearly seen that sex reversal is taking place (fig. 1). the majority seem to undergo this sex change during the lengths 33-37 em (caudal length), although there are some of both sexes between 29 and 41 em due to differential growth rates. the mode of the female individuals in the population, peaks at 26-30 em whereas the male individuals in the populations peak at 39-43 centimeter. the conclusion is that chrysoblephus cristiceps is protogynous hermaphroditic. (ii) the mass-length relationships were calculated: w = 0,0324 u·877 w = 0,0052 u·m it can be seen from fig. 2 that in the smaller female fish this relationship is curvi-linear whereas in the larger, male fish, this reiationship is linear i.e. growth changes between sexes. (iii) by grouping all the maturity index data by month, as inactive (maturity indices 1 and 2) and active (maturity indices 3 to 7), the relative proportion of active to inactive indicates a peak activity in november and a minimum in july-august, suggesting a spawning period in october to december (fig. 3). discussion chan ( 1970) reviewed the phenomenon of sex reversal found in vertebrates and concluded that sexuality in these animals may be classified into two main groups i.e. gonochorism and hermaphroditism . he notes that in the sparids there exists a diversified expression of sexuality and hermaphroditism. the basic structure of the gonads of sparids are reported to consist of a paired ovotestes fused posteriorly while the cavities of the two lobes form a common duct. internally each lobe consists of ovarium lamellae, where ovarium follicles develop and mature. posteriorly on the ventro-iateral part of each lobe, testicular tissues are visible as two bands adjacent to the lumen of the gonad with connective tissue separating male and female zones. macroscopically the development of the testicular part were seen to develop on the ventrolateral portion of the gonads in chrysoblephus cristiceps as well as the roman chrysoblephus laticeps and another sparid, the red steenbras petrus rupestris. this latter species was reported by penrith (1972) to be bisexual or gonochoristic. the evolution of hermaphroditism is extremely scanty but it is obvious that simultaneous hermaphroditism would be a disadvantage due to the possibility of self-fertilization and inbreeding. however, protogynous hermaphrodites could provide a mechanism as good as gonochorists. ' penrith (1972) presents an intriguing theory for the presence of pro45 e ~om~----------------------------~~------~ fi) w .... ~ z: ui ... ~ fig. 2. 10 .to 50 fi) w .... ooc 2 w "" ooc 2 mass-length relationships for chrysoblephus cristiceps 00 and 99· togynous hermaphroditism in chrysoblephus laticeps. he believes that this fish with its large heavy head, flabby body, and soft slightly concave caudal fin, becomes hydrodynamically a poorer swimmer as it increases in size. the advantage of sex reversal being that the younger reproductive sector of the population could avoid their natural predators, while the larger slower males would have the potential of fertilizing several 46 . , , • , '0 ii , 2. i' i. a + fig. 3. monthly relative frequency of maturity indices 3 to 7 fish in the population. females. in this way survival of the species under natural conditions is guaranteed. chrysoblephus cristiceps is similar in body dimensions to chtysoblephus laticeps and is therefore also prone to becoming a poorer swimmer as it increases with size and age. dageraad were relatively abundant along the south coast between cape point and algoa bay but their numbers and size in catches have decreased rapidly during recent times. f. spalmer (pets . comm.) noted that at arniston this change has been most dramatic, whereas the fishermen caught many, large dageraad they now return with fewer and smaller fish. the tsitsikama coast is treacherous and the population of dageraad have not been exposed to the same pressure of exploitation as other areas along the coast. it is suggested that the reason for good catches still to be obtained at tsitsikama, is due to the protection these fish are afforded in the park. at present the conservation of this species is based on a minimum length size of 20 centimeters. it is proposed that due to the protogynous hermaphroditic nature of dageraad that a regulatory measure be imposed on the maximum size and that all fish over 40 cm (lc) should be released when caught. (a similar regulation exists in the case of the lobsters along the northwestern shore of america). 47 a cknowledgemenls i would like to extend my thanks to messrs. t. dearlove, d. f. bower, j. a. fourie, p. van rooyen, p. scott and p. ngubu for assisting in the catching and recording of these fish. references chan, s. t. h . 1970. natural sex reversal in vertebrates. phil. trans . roy . sqc. lond. b. 259 :59-71. lucks, d. k. 1970. aspekte van die biologie van die witsteenbras (lilhognalhus aureli smith, 1962) in die sandwichhawe-strandmeer. m .sc. thesis, university of stellenbosch. (unpublished). mehl, j. a. p. 1974. ecology, osmoregulation-and reproductive biology of the white steenbras, lithognathus lithognalhus (teleostei: sparidae). zool. afro 8(2): 157-230. penrith, m. j. 1972. sex reversal in the sparid fish chrysoblephus laliceps. koedoe 15: 135-139. 48 page 1 page 2 page 3 page 4 page 5 page 6 reviewer acknowledgement koedoe 2011; 53(1)http://www.koedoe.co.za koedoe african protected area conservation and science recognises the value and importance of the peer reviewer in the overall publication process – not only in shaping the individual manuscript, but also in shaping the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely, productive peer review process. we would like to take this opportunity to thank all reviewers who participated in shaping this volume of koedoe: we appreciate the time taken to perform your review successfully. page 1 of 1 in an effort to facilitate the selection of appropriate peer reviewers for koedoe, we ask that you take a moment to update your electronic portfolio on www.koedoe.co.za for our files, allowing us better access to your areas of interest and expertise, in order to match reviewers with submitted manuscripts. if you would like to become a reviewer, please visit the journal website and register as a reviewer. to access your details on the website, you will need to follow these steps: 1. log into the online journal at http://www. koedoe.co.za 2. in your ‘user home’ [http://www.koedoe. co.za/index.php/koedoe/ user] select ‘edit my profile’ under the heading ‘my account’ and insert all relevant details, bio statement and reviewing interest. 3. it is good practice as reviewer to update your personal details regularly to ensure contact with you throughout your professional term as reviewer to koedoe. please do not hesitate to contact me if you require assistance in performing this task. margo martens submissions@koedoe.co.za tel: +27 (0)21 975 2602 fax: +27 (0)21 975 4635 koedoe african protected area conservation and science andrew deacon barend erasmus brian van wilgen cang hui charmaine uys clive webber corli coetsee darrell townsend deon furstenburg enoabasi anwana freek venter georgette lagendijk gyan sharma helen suich holly copeland ian craigie ian whyte jay barton john herrmann john wilson jozef šibík jürgen dengler lixin wang mariska te beest mark robertson martin vavra mary cadenasso rachel mostert robert westfall ryan emanuel sam ferreira shayne jacobs stefan foord stefan schindler stefan siebert stephen woodley vassiliki kati wayne twine willie boonzaaier yu-lung hsieh filelist convert a pdf file! participants to the symposium d j ackerman rsa n h fick lebowa d p ackerman rsa b g fourie rsa g m adendorff rsa j l s fourie rsa mrs c allcock rsa p f fourie rsa n allcock rsa j fowkes rsa r h andrag rsa g f van l froneman rsa g f barkhuizen rsa dr f fuggle rsa mrs a bean rsa i garland rsa cd beaumont rhod esia s gcumisa zululand mrs 0 berrange rsa j t geddes page rsa d j beuster bophuthatswana w p d gertenbach rsa b bezuidenhout rhodesia n h gilfillan rsa dr bhnengu zululand tgili rsa prof r bigalke rsa r girdwood zululand dr r bigalke rsa h w j grobler rsa e g h h blohm rsa h j grove rsa r bolten rhod esia n gumede swaziland j h bosch rsa m h gunukelo qwaqwa h. botha rsa dr a j hall-martin rsa l botha rsa dr h hamburger rsa prof r botha rsa dr j hanks rsa prof j du p bothma rsa dr sue hart rsa h h braack rsa w hartman rsa dr 0 j brand rsa the hon o brightman rhodesia w w b havemann rsa p j l bronkhorst rsa dr 0 hey rsa b r bryden rsa mrs c hugh es rhodesia am brynard rsa b j huntley rsa o butler rsa l hurry rsa o chapman rsa dr j a hurter rsa dr g f t child rhod esia ow immelman rsa j clarke rsa rj immink rsa b j coetzee rsa r jones rsa cooper lesotho dr s c j joub ert rsa k cooper rsa j j kloppers rsa r cross rsa dr r knobel rsa a s d e beer vendaland korsten rsa mj de beer rsa b j krohn rsa w a de beer rsa a e kuschke rsa dr g de graaff rsa mrs s m labuschagne rsa j h de kock rsa j j la grange rsa b de la bat swa b p lamprecht rsa dr v de vos rsa m landsman rsa chief j m olamini swaziland c f s a ie riche rsa his royal highness prince p j ie roux rsa m d1amini swaziland p r mabusa swaziland s d1amini swaziland chief a n madzivhandila vendala nd mrs h downing rsa chief b malatji lebowa o h c du plessis rsa dr 0 martiny rsa j w j c du plessis rsa w massyn rsa dr s s du plessis rsa e t matenge botswana dr 0 edwards rsa i a w mcdonald rsa prof f c eloff rsa i mclelland rhodesia dr i l engelbrecht rsa k t mgwamotsoko botswana mrs i l engelbrecht rsa j millar rsa m english rsa m g l mills rsa the hon w h fa urie rsa t minnie rsa p ferreira rsa m g mohale qwaqwa vi dr e moll rsa cd stainbank rsa dr h 0 monnig rsa mrs j stoltz rsa dr g de v morrison mp rsa chief s v suping bophuthatswana w 0 morsbach rsa dc swart rsa the h on p k taylor rhod esia dr lap a munnik rsa a e g trollip rsa chief n e ngomane swaziland the hon d c h uys rsa dr r g noble rsa p w a van der merwe gazank ulu m ntloko transkei a w b van der walt rsa h a oosthuizen lebowa dr p t van der walt rsa j g pansegrouw rsa dr r van der wcst huiz en rsa h c payne rsa adv h van dyk rsa mrs sparker rhodesia e p van niekerk rsa b m phillips lesotho the hon dr u de v pienaar rsa s g j van niekerk rsa m rs e a reill y swaziland w van riet rsa t e reilly swaziland levan rooyen rsa dr e m rh oodie rsa p f van straaten rsa d richards rsa t he hon p zj van vuurenrsa dr g a robinson rs a t he hon a c van wyk rsa m robson rh odesia p van wyk rsa dr a e rupert rsa lj van zyl rsa d rushworth rhodesia k yeary rsa dr h b rycroft rsa a l versfeld ciskei r salmon rsa dr f vollmar switserland dr m schweppenhauser rhod esia a h vosloo rsa j l serfontein rsa g n wagener rsa n j shaw australia c walker rsa k r shuter rsa swell s rsa m l sifelani rhodesia d whitespunner rhod esia a 0 simpson rsa mrs j woodl ey rsa chief 0 sithole zululand m iss j wright rsa z p siwane ciskei dr e young rsa n smit zululand d yssel rsa lj smith rsa d y zimu zululand dr g l smuts rsa chief z p ziwane ciskei vb page 1 page 2 abstract introduction materials and methods results discussion and recommendations conclusion acknowledgements references appendix 1 about the author(s) carina coetzer research unit: environmental sciences and management, north-west university, south africa hindrik bouwman research unit: environmental sciences and management, north-west university, south africa citation coetzer, c. & bouwman, h., 2017, ‘waterbird flight initiation distances at barberspan bird sanctuary, south africa’, koedoe 59(1), a1419. https://doi.org/10.4102/koedoe.v59i1.1419 original research waterbird flight initiation distances at barberspan bird sanctuary, south africa carina coetzer, hindrik bouwman received: 27 june 2016; accepted: 11 apr. 2017; published: 30 may 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract with tourism in south africa expanding, the number of avitourists increases. the increase in infrastructure and human activities in protected areas, if not managed properly, can be harmful to birds. flight initiation distances (fid) can be used as a method to monitor habituation to disturbances. this study was performed at the barberspan bird sanctuary, north west province, south africa, to determine the levels of habituation among waterbirds and make appropriate recommendations regarding the management of the reserve. our results indicated a 0.29 m increase in fid per gram reported mean biomass. compared with conspecific or congeneric birds from australia, europe and north america, south african birds have relatively larger fids to human disturbance, which may indicate lower habituation. we also calculated buffer zones based on the maximum fid of the waterbirds for three mass groups. these buffer zones were then matched with the spatial distribution of the birds along the shoreline. we recommend that the mean fid for the blacksmith lapwing, vanellus armatus (62 m), can be used as approach distance outside the breeding season in areas where the birds are sparsely distributed and 104 m during the breeding season in breeding areas. a large buffer of 200 m is suggested for areas with threatened, sensitive and skittish species. however, it is still preferable for avitourists to use the bird hides along the shores. conservation implications: this study provides information for conservation management at barberspan, based on typical birder activity. smaller birds would need smaller buffer zones, while larger birds need much greater distances from observers to minimise disturbance. similar studies can be applied elsewhere. introduction south africa is an ecotourism hotspot, with large numbers of international and local tourists visiting its numerous parks and reserves. when the number of tourists visiting south africa increases, avitourists (birdwatchers) will also increase, as south africa hosts a large variety of species, habitats, and botanical centres of endemism (biggs et al. 2011). this influx of human activity into protected natural areas, if not managed properly, can be harmful to the resident wildlife and have far-reaching effects on the future of that area as an ecotourism destination. flight initiation distances (fid) can be defined as the distance at which an individual bird approached by a predator or threat initiates flight, and can be used to determine the degree of habituation of avian species in a protected area (blumstein 2003, 2006). there are various terminologies used in the literature on distances that birds may be approached at, such as buffer zone, set-back distance and approach distance (blumstein et al. 2003; madsen 1998; mcleod et al. 2013; rodgers & smith 1995). here, we consider the use of approach distances (for individual birds) and buffer zones (for specific areas) to be appropriate. fleeing generally occurs when the costs and benefits are optimised. this is known as the optimal escape theory (møller & tryjanowski 2014; stankowich & blumstein 2005). costs may include decreased time for nest building or foraging (møller et al. 2013a; møller & tryjanowski 2014) and consequently poorer health in certain species if the situation is not remedied (kerbiriou et al. 2009; møller 2010; møller & liang 2012). fid is a species-specific trait (blumstein et al. 2003) and can, therefore, be used to determine the effect of disturbances on species, including rare or vulnerable species. it should, however, be mentioned that fid is also dependent on other factors such as flock size, speed of approach, distance to and availability of safe structures, starting distance (the distance from which the predators or humans begin to approach the individual or group) and type of disturbance (blumstein 2003, 2006; cooper 2005; dugatkin 2009; geist et al. 2005; kullberg & lafrenz 2007; mcleod et al. 2013). as birds habituate to human activities in specific areas such as urban and recreational areas, their fids will shorten and allow them to coexist with humans (møller et al. 2013a). however, in a bird sanctuary or reserve, it is generally deemed undesirable for birds to habituate, as the disruption in foraging time and the increased stress levels in the birds can lead to decreased biomass (borgmann 2011; kerbiriou et al. 2009; rodgers & smith 1995). some birds, like the goliath heron ardea goliath, also generally avoid any man-made structures (hockey, dean & ryan 2005), limiting them to sanctuaries and mostly natural areas. some species numbers are, although globally listed as least concerned, declining because of human disturbances, including pollution. these species include the yellow-billed stork mycteria ibis (which is near threatened in south africa) and the african darter anhinga rufa (hockey et al. 2005). it should be noted that there are opposing opinions concerning habituation. goering and cherry (1971) found disturbance to have no significant effects on breeding success, and nisbet (2000) suggests that habituation to humans should be promoted in waterbirds, as he found no evidence of human disturbances causing substantial harm to gulls (larus spp.), terns (sterna spp.) and herons (ardeidae). hockey et al. (2005) also mention that the numbers of several species have increased because of man-made structures creating more habitat areas for them to occupy. some of these species include the three-banded plover charadrius tricollaris and hadeda ibis bostrychia hagedash. gill, norris and sutherland (2001) suggest that if birds avoid disturbance in an area, it does not necessarily have a negative consequence at the population level. they might only be avoiding the disturbance because they have access to a better site in the near vicinity. anthropogenic disturbances should, therefore, be well managed to allow sufficient time for foraging and nest building by birds, but it is not completely necessary to terminate all human activity in order to preserve the natural quality of the area. for that reason, it is necessary to study the fids of birds to determine their level of habituation to disturbances (mcleod et al. 2013). once the fids of species are determined in a reserve or sanctuary, buffer zones for recreation, approaches, and other human activities can be established for better conservation of the habitats (blumstein et al. 2003; mcleod et al. 2013). the aim of this study was to determine the fids of the waterbirds at barberspan bird sanctuary and propose buffer zones accordingly to ensure better management of the site as a bird sanctuary. materials and methods study site this study was conducted at barberspan bird sanctuary (26° 35̕ s 25° 35̕ e), 307 km west of johannesburg, between sannieshof and delareyville. this is a summer rainfall area with annual rainfall averaging around 500 mm. the daily mean minimum and maximum temperatures during summer are 15 °c and 30 °c, respectively, and during winter 0 °c and 20 °c (south african weather service data). barberspan was one of the first (1975) wetlands in south africa to be designated a wetland of international importance (ramsar site) and represents a rare and unusual type of perennial wetland, that is, a large grassland pan. the water surface covers approximately 2000 ha and only has an overflow during high water levels, making the water of the pan alkaline because of evaporation. it functions as an important seasonal stopover site for migrating waterbirds and provides sanctuary for moulting waterbirds (milstein 1975; ndlovu et al. 2013). of great importance is its provision of perennial water in an area characterised by seasonal wetlands (milstein 1975). there is no hunting allowed in the reserve. a mean of 3052 tourists visit barberspan annually but less than 10% are strictly birdwatchers, while the remaining 90% are anglers or tourists passing through the area to another destination (van der merwe, s., 2016, email, 08 september, barbersp@lantic.net). between april 2013 and march 2014, 3243 people in 1393 vehicles visited the reserve. more than 350 aquatic and terrestrial bird species have been recorded. the pan sometimes holds over 20 000 birds (bouwman & hoffman 2007). during the july 2014 coordinated waterbird counts (cwac), 6858 individuals representing 44 waterbird species were observed (cwac report, july 2014, van der merwe, s., 2016, email, 05 september, barbersp@lantic.net). fids were measured in all accessible areas around the pan, including the frequently used general angling area on the eastern shore, the infrequently used competition angling area and the nature reserve covering the whole of the western shore. the latter area is a no-angling area and is strictly for the use of hikers and birders. these localities with varying numbers of visitors enabled us to get a general view of the birds’ habituation at barberspan. data collection and analysis flight initiation distances fieldwork was conducted in 5-day sessions, once in a month, for 3 months (march, april, and july 2014) as these autumn months include a high number of both resident and some migratory waterbird species. observations were made, and fids were recorded from an hour after sunrise for 3 h (06:30–09:30), and again 2 h before sunset until sunset (16:00–18:00). fids were measured by identifying the bird from as far away as possible, and taking a starting distance with a leica rangemaster 1200 laser range finder. two or three observers approached the birds, wearing dark-coloured clothing for uniformity and consistency, and walking abreast so as to not appear as only one person (geist et al. 2005; mcleod et al. 2013). the mean height of the observers was 1.8 m. if birds were in a group, a focal, visible individual at the edge of the group was selected to measure starting distance and fid. the bird was approached head-on and in a straight line at a leisurely walking pace (± 2 km/h). the birds were not stalked but approached in the open, which the flat and open terrain of barberspan allowed for. fid was taken as soon as the bird took flight because of the observers. in our case, we assumed that the obvious interest shown by the approaching humans by looking and pointing things at them (heads, binoculars, fingers, and the laser range finder; see also bateman & fleming 2011) was perceived as threats by the birds being approached. any flight because of obvious relocation for foraging or other disturbances was not recorded. mean body mass of each species was used as a reference to fid as it was previously found to correlate with fid (møller & erritzøe 2010; møller, vagasi, & pap 2013b). the mean mass of the species was obtained from del hoyo, elliot and sargatal (1992, 1996, 2004). data for species with more than five fid observations were used (33 species, for a total of 525 records) for univariate analyses. the untransformed data were analysed in graphpad prism v5.04 (http://www.graphpad.com) using linear regression and column statistics. linear regression was used to investigate the correlation between mass and fid. one-phase decay was used to plot fid versus starting distance, with outliers eliminated (1% and 99% percentiles). a non-linear, one-phase decay model was chosen assuming that the rate at which fid increases is proportional to an increase in starting distance (http://www.graphpad.com). buffer zones were calculated by using the maximum fid for species in arbitrarily chosen mass classes, namely small (less than 200 g), medium (201 g – 1000 g), and large birds (more than 1000 g). although several authors (borgmann 2011; rodgers & smith 1995) suggest using only or mostly the most sensitive or skittish species to establish buffer zones (or set-back distances), we combined data for all species as this represents the actual situation on the ground, as barberspan may be merely a stopover site for some of the skittish and rare birds. using this combination allowed for better accommodation of all species when proposing a buffer zone. results we generated 525 individual fids over 30 sessions divided equally between dawn and dusk sessions. the biomass of the different species varied over two orders of magnitude (21 g – 4330 g). the mean fids for both families that were the most common and the different mass classes of the birds at barberspan are presented in table 1. generally, larger bird species had greater fids (table 1). species with mean fids less than 40 m were waders (families charadriidae, recurvirostridae, scolopacidae, motacillidae and podicipedidae), while species with mean fids greater than 100 m included five ardeidae species and the lesser flamingo phoenicopterus minor. the shortest fid was for the three-banded plover at 7 m, while the greatest fid was for the goliath heron at 300 m. a summary of the minimum, maximum, mean, and median fids, including the starting distance and coefficient of variation (cv), for 33 waterbird species at barberspan is presented in table 2. table 2 also shows comparable fid data (conspecific or congeneric) from blumstein (2006). table 1: mean flight initiation distance (m) with standard deviation for the most common waterbird families and mass classes found at barberspan. table 2: flight initiation distances for 33 waterbird species at barberspan. there was no normal distribution of all fids combined (figure 1a), neither for the log-transformed data, so we used untransformed data as it describes a normal situation to be faced in a water body such as barberspan. a mean fid of 73.5 m was derived with a standard deviation of 43.5 m; the upper standard deviation (s.d.) was at 125 m. the 95% confidence intervals of the mean were very narrow around the mean, between 70 m and 77 m. figure 1: (a) all recorded flight initiation distances, irrespective of species. (b) linear regression of flight initiation distances relative to bird mass. (c) coefficient of variation (%) for all species. (d) non-linear (one-phase decay) regression of mass and flight initiation distances and starting distances. because most species-specific fid data were normally distributed according to the shapiro–wilk normality test, they were not transformed for regression analyses. various non-linear models were fitted, but linear regression produced the best results (figure 1b and d). one-phase decay gave the best results for regressing fid versus starting distances (figure 1d). mass influenced fid positively (figure 1b) using linear regression (r2 = 0.3928; p < 0.0001). for every gram of mass increase, there was a corresponding 0.29 m increase in fid. the cvs of the fids were normally distributed (figure 1c). the mean cv for all species was 37.5%, with a standard deviation of 10.3%. the starting distances influenced the fid markedly (r2 = 0.6581) using a one-phase decay model (figure 1d). nine outliers were excluded as they exceeded the 1% percentile. the plateau was reached at 246 m, after which the increase in starting distances did not affect fid. discussion and recommendations flight initiation distance for the relatively small area sampled and the good visibility, we assumed that our sample (consisting of the 33 species recorded more than five times) was representative of the occurrence of the species at barberspan. however, larger, more conspicuous birds, such as the goliath heron, would attract more attention per individual than smaller birds per sampling effort and there may be a bias towards the larger birds. the smaller waders and ducks are not restricted to open habitat, as some (such as the red-billed teal anas erythrorhyncha and three-banded plover) can also be found in or near areas with more vegetation along the shoreline. a shorter fid, allowed for by the ability to quickly escape, makes it safer for them to venture here. the taller birds would be less restricted by obstructive vegetation, because a higher eye height would allow the farther detection of bigger prey items and threats (blumstein 2006). some of the species we observed, especially those favouring the frequently used angling area, seemed to be partially habituated to humans. fids reported for australian, european and american conspecifics and congenerics (blumstein 2006) were consistently shorter than at barberspan, and some by a quite a large margin (table 2). the large fids measured at barberspan bird sanctuary may be accounted for by two interacting reasons. firstly, black-backed jackal canis mesomelas, cape clawless otter aonyx capensis, caracal felis caracal, small-spotted genet genetta genetta, a variety of mongooses, and many raptors (including african fish eagle haliaeetus vocifer, taking ducks and flamingos; h. bouwman, pers. obs.) also occur at the barberspan bird sanctuary. these are all known predators of birds. this occurrence of fast-moving predators might contribute to the longer fids when compared with congenerics and conspecifics elsewhere (blumstein 2006; table 2). møller and liang (2012) predicted and found that species in tropical areas take smaller risks (therefore longer fids) than the same or closely related species in temperate areas. according to them, this effect was not only related to predation pressure, possibly being greater in tropical areas, but also to life-history traits, especially clutch size. tropical birds live longer, produce smaller clutches and start breeding at a later age. therefore, they should be more risk-averse than their temperate counterparts (møller & liang 2012). the species cvs were normally distributed (figure 1c), and the regression between cv and mass was not significant (p = 0.0711), indicating a set of common factors that birds, irrespective of species, consider when perceiving and reacting to threats. describing these factors will need more research. our results confirm those of blumstein (2003) and cooper (2005), namely that fid and starting distances are not linearly related (see figure 1d, r2 = 0.6581). beyond a certain distance, an increase in mean starting distances does not seem to affect fid, and therefore, one-phase decay was an appropriate model. distances longer than 246 m (the plateau) is very long and visual acuity and observation skills of the observers may become a factor. conservation management nature reserves dedicated to mostly birdwatching such as barberspan should be managed with least possible disturbance to the birds, while allowing birdwatchers good and unobstructed views and minimising disturbance to birds. figure 1a shows that the upper standard deviation of the mean of all observations was 125 m. not approaching any bird by less than 125 m would protect against most disturbances, but would still affect the larger birds such as the goliath heron. however, birds are not distributed evenly around barberspan because of large differences in vegetation, shallows, wind and bottom conditions (mud, gravel, sand, etc.). certain shallower areas with muddy bottoms have mainly smaller birds where the minimum approach distance may be shortened. smith and bouwman (unpublished) conducted a study in 2004 to establish distribution patterns along the edge of the pan for conservation purposes. their findings are presented in appendix 1 (figure 1-a1) and used to derive appropriate buffer zones. the shores of barberspan are not homogenous. combined with seasonal changes in water levels, a single approach to minimising disturbances will not be feasible. we propose the use of different approach distances for differing bird sizes, and buffer zones based on the average water level following the wet-season (september–march), assuming that most birds congregate at this interface. buffer zones are set at the maximum fid per mass group, which is 104 m for small, 213 m for medium and 300 m for large birds. rodgers and smith (1995) suggested the buffer zones (or set-back distances) in florida be set at 100 m for wading-bird colonies, and 180 m for tern and skimmer colonies. their distances were calculated by adding the standard deviation to the mean from their sample, and then adding another 40 m to allow for the distances where the birds usually become agitated by a disturbance. using a similar formula, erwin (1989) suggested roughly the same distances. these distances were, however, for breeding colonies. around foraging and loafing sites for most waterbirds in florida, rodgers and smith (1997) suggested a buffer zone of about 100 m. borgmann (2011) suggested a buffer zone of 250 m for waterfowl, diving ducks, wading birds and shorebirds, to lessen the impact of human disturbance on sensitive or skittish species. considering what smith and bouwman (unpublished) found at barberspan in 2004 (appendix 1), we suggest that the mediumand large-bird buffer zones (213 m and 300 m, respectively) be used in areas with greater species richness, such as hotspots a, b and e (figure 1-a1). elsewhere, the small-bird buffer zone can be applied. it might, however, be somewhat impractical as birders might not be able to see the plovers and sandpipers at 104 m. therefore, we suggest that the small-bird buffer be set at the highest species mean fid for the group smaller than 200 g, which is 62 m for the blacksmith lapwing vanellus armatus. during the breeding season, the original buffer zone of 104 m may still be implemented in areas surrounding the breeding colonies. for practical reasons, the 300 m buffer for large birds is only applicable in areas where threatened and sensitive species occur regularly, as the highest mean fid for birds larger than 1000 g is still accounted for under the medium buffer. for the goliath heron and flamingos, being the largest species at barberspan, we propose an individual minimum approach distance of the median fid (127 m) plus one standard deviation (56 m) be used for a more practical buffer of 183 m. there are several bird hides along the western shore of barberspan and it is still preferable for tourists to use these facilities. walking along the shore of the pan is allowed if size-class buffer zones are adhered to, while not disturbing the birds. the practical implementation of our proposals will need integration with the barberspan management plan following consultations with the management and stakeholders. information on appropriate buffer zones and sensitive areas can be provided at reception, and the distances can be indicated by representation of bird cut-outs at appropriate distances for birders to practice their distance estimations. the effectiveness of the memory retention of distances should, however, be tested. several factors in our study may be perceived as constraints. firstly, we approached the birds directly on foot (simulating a threat), while a more oblique approach might result in shorter fids (bateman & fleming 2011). approach by vehicle, which may often be the case for birdwatchers in open habitat, would presumably have an influence on fid (mcleod et al. 2013). another constraint may be that our observations were made outside the breeding season. species that forage close to their nests (such as the kittlitz’s plovers charadrius pecuarius and blacksmith lapwings) may adjust their anti-predator behaviours and fids, while birds that forage away from their nests, such as the herons, presumably would not or very little. for example, møller et al. (2013a) found that fid was shortened for european bird species following harsh winter. we are also not aware of the effects of drought on fids, which could be a valuable future research area. the species distribution and biomass study (smith & bouwman unpublished) was conducted 10 years prior to the flight initiation study. distribution patterns and biomass could have changed over time. however, we believe the changes are not so profound as to disregard the information, as the conditions around the pan have not drastically changed. conclusion this study has shown how flight initiation distances (fids) of waterbirds can be used to suggest management of bird sanctuaries such as barberspan. based on a combination of fids and spatial distribution around the pan, we propose approach distances for individual birds and buffer zones to be implemented around the pan according to the size of the birds (62 m and 183 m for small and larger species, respectively), and species diversity and richness of the area. practical methods for implementation can be considered by the management and stakeholders of barberspan. future studies may focus on addressing the constraints mentioned in this study, such as repeating the fid measurements during the breeding season, the effects of the climate, and changes thereof, on the fid, as well as finding one or more predictive factors which birds, irrespective of species, may consider when perceiving and reacting to threats in their environment. acknowledgements the authors thank mr s. van der merwe, manager of barberspan bird sanctuary, as well as andrew mvundle, amos koloti and lebo moeti, monitoring and research assistants from the reserve. the north west parks and tourism board gave permission for this study. opinions expressed and conclusions arrived at are those of the authors and are not necessarily to be attributed to the north west parks and tourism board, or national research foundation. the authors thank the national research foundation of south africa for funding. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions the study was conceived by h.b. and c.c. fieldwork was done by c.c. c.c. and h.b. analysed the data and drafted the manuscript. references bateman, p.w. & fleming, p.a., 2011, ‘who are you looking at? hadeda ibises use direction of gaze, head orientation, and approach speed in their risk assessment of a potential predator’, journal of zoology 285, 316–323. https://doi.org/10.1111/j.1469-7998.2011.00846.x biggs, d., turpie, j., fabricius, c. & spenceley, a., 2011, ‘the value of avitourism for conservation and job creation – an analysis from south africa’, conservation and society 9, 80–90. https://doi.org/10.4103/0972-4923.79198 blumstein, d.t., 2003, ‘flight initiation distance in bird is dependent on intruder starting distance’, the journal of wildlife management 67, 852–857. https://doi.org/10.2307/3802692 blumstein, d.t., 2006, ‘developing an evolutionary ecology of fear: how life history and natural history traits affect disturbance tolerance in birds’, animal behaviour 71, 389–399. https://doi.org/10.1016/j.anbehav.2005.05.010 blumstein, d.t., anthony, l.l., harcourt, r. & ross, g., 2003, ‘testing a key assumption of wildlife buffer zones: is flight initiation distance a species specific trait?’, biological conservation 110, 97–100. https://doi.org/10.1016/s0006-3207(02)00180-5 borgmann, k.l., 2011, a review of human disturbance impacts on waterbirds, audubon, ca, viewed 01 june 2016, from http://www.sfbayjv.org/news-general.php bouwman, h. & hoffman, r., 2007, ‘the effects of fire on grassland bird communities of barberspan, north west province, south africa’, ostrich 78, 591–608. https://doi.org/10.2989/ostrich.2007.78.3.6.317 cooper, w., 2005, ‘when and how do predator starting distances affect flight initiation distances?’, canadian journal of zoology 83, 1045–1050. https://doi.org/10.1139/z05-104 del hoyo, j., elliot, a. & sargatal, j., 1992, handbook of the birds of the world, vol. 1, lynx editions, barcelona. del hoyo, j., elliot, a. & sargatal, j., 1996, handbook of the birds of the world, vol. 3, lynx editions, barcelona. del hoyo, j., elliot, a. & sargatal, j., 2004, handbook of the birds of the world, vol. 9, lynx editions, barcelona. dugatkin, l.a., 2009, principles of animal behaviour, 2nd edn., w.w. norton & company, new york. erwin, r.m., 1989, ‘responses to human intruders by birds nesting in colonies: experimental results and management guidelines’, colonial waterbirds 12, 104–108. https://doi.org/10.2307/1521318 geist, c., liao, j., libby, s. & blumstein, d.t., 2005, ‘does intruder group size and orientation affect flight initiation distance in birds?’, animal biodiversity and conservation 28, 69–73. gill, j.a., norris, k. & sutherland, w.j., 2001, ‘why behavioural responses may not reflect the population consequences of human disturbance’, biological conservation 97, 265–268. https://doi.org/10.1016/s0006-3207(00)00002-1 goering, d.k. & cherry, r., 1971, ‘nestling mortality in a texas heronry’, wilson bulletin 83, 303–305. hockey, p.a.r., dean, w.r.j. & ryan, p.g. (eds.), 2005, roberts birds of southern africa, 7th edn., trustees of the john voelcker bird book fund, cape town. kerbiriou, c., le viol, i., robert, a., porchar, e., gourmelon, f. & julliard, r., 2009, ‘tourism in protected areas can threaten wild populations: from individual response to population viability of the chough pyrrhocorax pyrrhocorax’, journal of applied ecology 46, 657–665. https://doi.org/10.1111/j.1365-2664.2009.01646.x kullberg, c. & lafrenz, m., 2007, ‘escape take-off strategies in birds: the significance of protective cover’, behavioural ecology and sociobiology 61, 1555–1560. https://doi.org/10.1007/s00265-007-0387-1 madsen, j., 1998, ‘experimental refuges for migratory waterfowl in danish wetlands. i. baseline assessment of the disturbance effects of recreational activities’, journal of applied ecology 35(3), 386–397. https://doi.org/10.1046/j.1365-2664.1998.00314.x mcleod, e., guay, p.j., taysom, a.j., robinson, r.w. & weston, m.a., 2013, ‘buses, cars, bicycles, and walkers: the influence of the type of human transport on the flight responses of waterbirds’, public library of science 8, 1–12. https://doi.org/10.1371/journal.pone.0082008 milstein, p.l., 1975, ‘the biology of barberspan, with special reference to the avifauna’, ostrich supplement 10, 1–74. møller, a.p., 2010, ‘up, up and away: relative importance of horizontal and vertical escape from predators for survival and senescence’, journal of evolutionary biology 23, 1689–1698. https://doi.org/10.1111/j.1420-9101.2010.02034.x møller, a.p. & erritzøe, j., 2010, ‘flight distance and eye size in birds’, international journal of behavioural biology 116, 458–465. https://doi.org/10.1111/j.1439-0310.2010.01754.x møller, a.p., grim, t., ibáñez-álamo, j.d., markó, g. & tryjanowski, p., 2013a, ‘change in flight initiation distance between urban and rural habitats following a cold winter’, behavioural ecology 24(5), 1211–1217. https://doi.org/10.1093/beheco/art054 møller, a.p. & liang, w., 2012, ‘tropical birds take small risks’, behavioural ecology 24(1), 267–272. https://doi.org/10.1093/beheco/ars163 møller, a.p. & tryjanowski, p., 2014, ‘direction of approach by predators and flight initiation distance of urban and rural populations of birds’, behavioural ecology 25(4), 1–7. https://doi.org/10.1093/beheco/aru073 møller, a.p., vagasi, c.i. & pap, p.l., 2013b, ‘risk-taking and the evolution of mechanisms for rapid escape from predators’, journal of evolutionary biology 26, 1143–1150. https://doi.org/10.1111/jeb.12147 ndlovu, m., cumming, g.s., hockey, p.a.r., nkosi, m.d. & mutumi, g.l., 2013, ‘a study of moult-site fidelity in egyptian geese, alopochen aegyptiaca, in south africa’, african zoology 48(2), 240–249. https://doi.org/10.1080/15627020.2013.11407589 nisbet, i.c.t., 2000, ‘disturbance, habituation and management of waterbird colonies’, waterbirds: the international journal of waterbird biology 23(2), 312–332. rodgers, j.a., jr. & smith, h.t., 1995, ‘set-back distances to protect nesting bird colonies from human disturbance in florida’, conservation biology 9(1), 89–99. https://doi.org/10.1046/j.1523-1739.1995.09010089.x rodgers, j.a., jr. & smith, h.t., 1997, ‘buffer zone distances to protect foraging and loafing waterbirds from human disturbance in florida’, wildlife society bulletin 25(1), 139–145. stankowich, t. & blumstein, d.t., 2005, ‘fear in animals: a meta-analysis and review of risk assessment’, proceedings of the royal society b 272, 2627–2634. https://doi.org/10.1098/rspb.2005.3251 appendix 1 figure 1-a1 (presented here for illustration purposes) shows the mean species richness (a) and the mean biomass (b) found along the western shores of the pan, measured over 3 months in 2004. the eastern shores were not measured, as this is the frequently disturbed, public angling area. the information in figure 1-a1 allows for deriving appropriate protective buffer zones. figure 1-a1: mean avian species richness (a) and biomass (b) along the western shore of barberspan. the species richness hotspot marked a (figure 1-a1) is rarely disturbed by tourists and the bird hide is only accessible to visitors, while angling is prohibited in this area. the reeds and tall grass found here also make it a good breeding spot. large areas of shallow water and exposed mud banks occur here. this is also where the highest biomass was found (figure 1-a1b) as it is home to several duck and heron species, as well as spoonbills. hotspot b on figure 1-a1 is a seasonal wetland and island (goose point), depending on water levels, restricting access at high water. during our study, the area could be approached on foot during march 2014, but not during the other 2 months. both smaller waders and larger birds occur here. hotspots c–e (figure 1-a1) are shallow bays and have some dense reed clumps growing on the banks, making it favourable for a large number of species to feed, breed and find shelter. the shores between b and e are often visited by birders, as well as resident antelope for drinking. these areas are prone to flooding and during flood events, it hosts higher species richness. the shores are overgrown with tall grass, limiting the number of waders. hotspot e is the least accessible part of the pan, despite the national road passing not far from there. it has dense reed stands and trees. this will account for the high preference of the birds to feed, breed and find shelter here, as can be seen in figure 1-a1. article information authors: david newsome1 shannon hassell1 affiliations: 1school of environmental science, murdoch university, australia correspondence to: david newsome postal address: murdoch university, 60 south street, murdoch, western australia wa 6150, australia dates: received: 06 feb. 2013 accepted: 17 july 2013 published: 24 june 2014 how to cite this article: newsome, d. & hassell, s., 2014, ‘tourism and conservation in madagascar: the importance of andasibe national park’, koedoe 56(2), art. #1144, 8 pages. http://dx.doi.org/10.4102/ koedoe.v56i2.1144 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. tourism and conservation in madagascar: the importance of andasibe national park in this original research... open access • abstract • introduction • tourism and environmental conservation • madagascar as a natural-area tourism destination    • protected areas in madagascar    • tourism and environmental conservation in madagascar • threats to conservation in madagascar • the success of andasibe through local community engagement and conservation support • conclusion • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ madagascar is renowned for high levels of biodiversity and endemism. as a result of its unique flora and fauna, as well as the high levels of human threat to the environment, such as illegal clearing, hunting and political instability, it is a critical global conservation priority. andasibe–mantadia national park in eastern madagascar is one of the most popular protected areas visited by tourists. observations carried out in 2011 showed that even though there were some negative impacts associated with natural-area tourism, the benefits to both the local communities and associated biological conservation outweighed the negatives. natural-area tourism at andasibe is well organised, with many local guide associations having partnerships with international organisations and 50% of park fees going directly to local communities. forest loss is a widespread problem in madagascar, but at andasibe the forest is valued for its ecological function and as a generator of profits from natural-area tourism. exploitation of the park was not observed. andasibe is an example of how conservation and natural-area tourism can work together in madagascar for the benefit of local communities and the environment. however, with the current unstable political climate and lack of adequate wider tourism and conservation planning frameworks, awakening to its potential as a leading conservation tourism destination will not be a simple task.conservation implications: this research demonstrated that ecotourism can be an effective means of achieving conservation objectives, whilst, at the same time, improving the livelihoods of local people. we caution, however, that governments can do a lot more to encourage and support the nexus between tourism and conservation. introduction top ↑ ecological sustainability is a concept that has gained global attention in recent times, with landmark publications such as the bruntland report (world commission on environment and development 1987) and the rio declaration on environment and development (united nations 1992). in response to the increasing pressure being placed upon the natural environment, aspects of the tourism industry and system have progressed to incorporate this concept. tourism is a complex array of activities (e.g. see mill & morrison 1992; weaver & lawton 2006); however, for the purposes of exploring the context of tourism in madagascar, it is defined as natural-area tourism which also encompasses ecotourism and wildlife tourism. ecotourism is where attractions are nature-based, learning activities and outcomes are incorporated and attractions are managed for sustainability. wildlife tourism is where the focus is centred on viewing and interacting with wildlife under natural conditions. where such tourism aids in the protection of natural areas it is termed conservation tourism (newsome, moore & dowling 2013).conservation tourism (ct) can thus be viewed as a subset of ecotourism, where the activity acts as an instrument for biological conservation. buckley (2010:2) defined it more specifically as ‘commercial tourism which makes an ecologically significant net positive contribution to the effective conservation of biological diversity’. whilst ecotourism has been defined in various ways (blamey 1997; ceballos-lascuráin 1996; donohoe & needham 2006; fennell 2008; harrison 1997; weaver 2005, 2008), ct, however, has a more specific biological conservation focus. conservation of the natural assets (e.g. birds, mammals and flora) on which it depends is often a defining tenet of ecotourism (fennell 2008; newsome et al. 2013; weaver & lawton 2007) but it is not always considered essential (buckley 2010), nor has tourism’s actual contribution to conservation been scientifically studied and monitored in much depth in the literature (kruger 2005; weaver & lawton 2007). as the need for biological conservation and ecological sustainability becomes greater due to habitat loss, hunting wildlife, population increases and poverty, using a number of tools or instruments, such as tourism, can assist in achieving conservation goals. the aim of this article is to explore the potential utility and role of natural-area tourism in conservation, with a specific focus on madagascar. in particular, some recent personal observations from andasibe–mantadia national park and surrounds are highlighted as evidence that natural-area tourism does work in conserving wildlife and their habitats. it is cautioned, however, that although andasibe is a ct success story, much work remains to be done to realise the benefits evidenced at the national level in madagascar. tourism and environmental conservation top ↑ natural-area tourism, including its subsets ecotourism and ct, holds the potential for a number of positive impacts to occur on a variety of levels. it generates foreign currency and creates opportunities for local employment, which, in less developed countries, is a key limiting factor in development (mbaiwa 2008; sinclair 1998). kruger (2005) studied global cases of ecotourism and found that in those cases deemed sustainable, four main effects were present. these were increased conservation of the natural resource base, revenue creation by non-consumptive means increased for local communities, revenue increased at the regional and national level and positive changes of local communities towards conservation. using natural resources, such as landscapes and focal species, as the primary tourism attraction can encourage greater appreciation and conservation attitudes amongst local communities who need an attractive natural environment for such tourism to be successful. for example, this has been applied successfully with tigers in india (karanth et al. 2012; sinha et al. 2012), sea turtles and whales in australia (wilson & tisdell 2003) and african wild dogs in south africa (lindsey et al. 2005). in places where there are competing economic interests, such as mining and logging, tourism can be a more sustainable use of natural resources in the long term (briassoulis 2002; curtin sustainable tourism centre 2010). however, ct has its limitations and negative cases have been reported (buckley 2010). green and higginbottom (2001) grouped the potential impacts of humans on wildlife through natural-area tourism into three categories: disruption of activity, direct killing or injury and habitat alteration (including provision of food). kruger (2005) and lansing and de vries (2007) also questioned ecotourism’s role in conservation and whether it actually contributes towards conservation or if it is only a marketing ploy used by the tourism industry to capitalise on its popularity. cases of abusing the ‘eco’ label (‘green washing’) are common (fennell 2008; self, self & bell-haynes 2010). this is further exacerbated by the lack of common knowledge of what exactly ecotourism entails, lack of a single global eco-certification system and the lack of political will to embrace sustainability principles (kruger 2005; self et al. 2010). exploiting the very thing that tourists come to see is a concern and increased popularity can attract more tourists thereby increasing the chance of direct and indirect disturbances (newsome, dowling & moore 2005; stem et al. 2003). other examples include proliferations of unsustainable tourism infrastructure development (sinha et al. 2012), removing other forms of wildlife (predators, forest cover) to more easily see focus wildlife (tisdell & wilson 2012) and disturbances to wildlife such as disrupted courtship and mating, and finding food (newsome et al. 2005; newsome et al. 2013) which varies between species (blumstein et al. 2005). finding a balance between conserving the resource base on which ct depends and contributing to the economic, financial and social sustainability of the host community is an ongoing concern and requires careful, holistic management. madagascar is considered one of the world’s biodiversity hotspots (conservation international 2012). yet, it has had a tumultuous political, social and environmental history which we now turn to discuss in the context and tourism and conservation. madagascar as a natural-area tourism destination top ↑ madagascar is located off the east coast of africa in the indian ocean and is the fourth largest island in the world. owing to its long period of geographical isolation, it is characterised by extremely high wildlife endemism; of those that have been discovered, 100% of terrestrial mammals, 92% of reptiles and 100% of amphibians are unique to the island, with new species being continually discovered (goodman & benstead 2003, 2005). the natural environment of madagascar has been detrimentally impacted by humans as a result of habitat destruction, hunting and live capture of animals, with major negative flow on effects such as erosion and wide spread species extinctions (goodman & benstead 2003, 2005; grosvenor 1987; mittermeier et al. 2010). because of its high biodiversity, species endemism and level of human threat, madagascar is a critical global conservation priority (goodman & benstead 2003, 2005). the remaining natural environment is of great interest to tourists, with wildlife tourism being the most popular subset of natural-area tourism interests (christie & crompton 2003), especially with appealing species such as lemurs, chameleons, birds and baobab trees (mittermeier et al. 2010; pawliczek & mehta 2008). the remaining protected areas (pas) are extremely important in both biological conservation and supporting wildlife for natural-area tourism. niche wildlife tourism providers in the usa and europe run organised tours to see wildlife in madagascar, focusing on lemurs, birds and endemic plant life (figure 1). the international profile of madagascar’s endemic wildlife has resulted in the development of accommodation, tourism services and tour guiding at wildlife hotspots such as nosy mangabe, masoala national park, kirindy forest, berenty private reserve and andasibe–mantadia national park. figure 1: small group of international tourists (niche tourism) viewing and photographing indri at andasibe national park. protected areas in madagascar as of 2011, there were 45 terrestrial pas in madagascar, including 19 national parks, 5 strict nature reserves and 21 special reserves (wollenberg et al. 2011) (figure 2). this was a considerable increase to the existing pa network, which can be attributed to the malagasy government committing in 2003 to a tripling of pas to approximately 5.5 million ha by 2008 (pawliczek & mehta 2008; wollenberg et al. 2011). private and community-owned pas add to this public network (buckley 2010). the then president, marc ravalomanana, saw the potential of the unique remaining natural environment to assist in his government’s plans to further develop one of the poorest countries in the world and made conservation an important pillar of his madagascar action plan (map) (anon 2008; presidency of madagascar 2006). despite the high biodiversity values of madagascar and the natural-area tourism potential, its conservation implications have not yet been fully realised (christie & crompton 2003; pawliczek & mehta 2008). there are, however, examples of positive ecotourism development, which may contribute to biological conservation. figure 2: map of the protected areas (national parks, nature reserves and special reserves) of madagascar. note that a significant portion of the protected area network is inadequately protected. tourism and environmental conservation in madagascar without endemic wildlife as an attractor, the natural-area tourism industry of madagascar would struggle to survive. the impetus to conserve what remains is strong, at least with those who directly benefit from wildlife tourism, such as local communities adjacent to pas and local guides, up to the government level (ormsby & manni 2006; presidency of madagascar 2006). even though many pas are small in area, they support highly significant ecological communities that often occur nowhere else (buckley 2010; goodman & benstead 2003). buckley (2010) stated that:without the income from natural-area tourism, there is a much greater risk that the small areas of remnant vegetation will be cut for firewood and timber and used to graze cattle. (p. 47) such tourism focusing on viewing both flora and fauna shifts the attention from immediate unsustainable resource consumption to a potentially more sustainable non-consumptive pattern of behaviour (tisdell & wilson 2012). this direct form of conserving the natural environment through natural-area tourism is complemented by other, more indirect methods. ormsby and mannie (2006) studied the role of ecotourism in providing an economic justification for conservation and also the role of guides in raising awareness and support for conservation in masoala national park, in the north-east of madagascar. they found that local guides played a key role in communicating to tourists the importance of the environment, with 73% of surveyed guides stating that they wanted to pursue a career in guiding to help protect the environment. training is rigorous and involvement with a guide association is compulsory, as is knowledge of a secondary language other than malagasy or french (newsome et al. 2013; ormsby & manni 2006). in addition to educating tourists, the guides, in association with a local environmental youth group, broadcast a weekly hour-long radio programme at a local station. the aim is to communicate to local communities the importance of natural-area tourism and conservation of the natural environment. one guide explained that many local people do not realise the biological and economic importance of madagascar’s environment. masoala national park also allocates half of park visitor entrance fees to local management committees to use for development projects of their choice (ormsby & manni 2006). this further encourages the protection of the environment at the local level. international commercial tour companies can also make a positive contribution to conservation through tourism. buckley (2010) studied the company natural habitat adventures (nha) and their wildlife viewing tour of madagascar (natural habitat adventures 2012). nha is an american ecotourism travel and tour company with a strong sustainability focus and a unique partnership with the world wide fund for nature. it hires local guides at destinations, contributing to community social and economic development as well as ensuring that the tourists receive superior local knowledge. nha utilises a variety of pas for their tours, including private reserves. similar economic incentives for conservation occur, such as developing markets for forest products in buffer zones between agricultural land and protected areas; however, landowners are not restricted by public land-clearing laws. buckley (2010:50) stated that private land will remain undisturbed only if, (1) local land owners have no need or incentive to cut them or (2) if they gain greater net benefits by protecting them. according to pawliczek and mehta (2008:41), madagascar is like a ‘sleeping beauty’ who is finally awakening to its natural-area tourism potential. wildlife tourism in this unique natural environment does not, however, guarantee a positive contribution towards conservation. stephenson (1993) studied the impacts of tourism at perinet (now andasibe–mantadia national park) and observed that despite the presence of designated walking paths, there was a proliferation of informal trails created by visitors and guides to get closer for wildlife viewing. this was causing an increase in erosion, trampling of native vegetation and favouring invasions of herbs, weeds and rats. stephenson (1993) also reported that daily forest searches for animals such as the lemur indri (indri indri) were also causing animals to become habituated to humans and show no alarm in their presence. in a later study, buckley (2010:51) reported incidences where wildlife was being exploited for profit by local people. to ensure sightings, some guides had illegally captured wildlife and both charged tourists to see these caged animals or had placed them in an easy to see spot so as to mislead tourists into believing they had ‘discovered’ the animal, for example a rare tomato frog, dyscophus antongili that had been refrigerated to slow it down. mittermeier et al. (2010) also reported the illegal capture and touristic exhibition of a fossa (cryptoprocta ferox), a native mammal made famous by the film madagascar. what was most concerning was that the visitors to perinet and the ‘conservation tourists’ on the organised trip did not seem to notice anything awry and seemed pleased to have photographic opportunities; potentially perpetuating an increase in both negative impacts and illegal actions which are counterproductive to biological conservation. ill conceived and unmanaged wildlife tourism can have negative impacts on conservation; in madagascar, however, the threats run much deeper than this. threats to conservation in madagascar top ↑ the threats to conservation in madagascar run deep and wide. since humans colonised the island around 2000 years ago, it is estimated that only 10% – 20% of the original forest cover remains (conservation international 2012; goodman & benstead 2003, 2005). this loss of habitat has caused widespread extinctions, erosion, loss of arable land and invasions of aggressive pest species (durbin & ratrimoarisaona 1996; mittermeier et al. 2010). systemic issues such as poverty, traditional unsustainable farming methods (primarily slash and burn), a lack of conservation and ecotourism education and repeated political turmoil have played major roles in this loss of habitat and in the battle for biological conservation (keane et al. 2011; mittermeier et al. 2010; ormsby & manni 2006; pawliczek & mehta 2008). this is exacerbated by the variety of different ethnic groups, basic education levels and differences in values placed upon the natural environment (scales 2012).in the period prior to 2009, the malagasy government made several attempts to increase the use of ecotourism as a development and conservation tool with various policy statements, with varying levels of success (pawliczek & mehta 2008). these steps towards utilising natural-area tourism as a positive instrument for development were shattered following a coup in 2009, with tensions between the government and the then mayor of the capital, antananarivo, leading to violent opposition and the presidents’ eventual forced resignation from office (ploch & cook 2012). current tourism and conservation policies such as the map and the national environmental action plan were dismissed with the upheaval and tourism arrivals plunged from 375 000 in 2008 to 163 000 in 2009 (mittermeier et al. 2010). international outrage at the unconstitutional taking of power froze international development aid, which was instrumental in the battle for sustainable development and biological conservation (ploch & cook 2012; united states of america: department of state 2012). in addition to this, looting and poaching has been a major problem since the 2009 coup, with a poverty stricken public taking advantage of political instability by cutting down valuable rosewood (dalbergia baronii) and others for export, trampling fragile habitats and hunting and eating bush meat, including endangered lemurs (braun 2009; mittermeier et al. 2010). those employed in the natural-area tourism industry as guides and support were suddenly out of work and living in fear for their livelihoods (braun 2009; mittermeier et al. 2010). this was not the first time political instability has impacted both madagascar’s natural-area tourism industry as well as its biological conservation; in 1991 tourist arrivals dropped to approximately 35 000 from 53 000 in 1990 and illegal deforestation and poaching occurred as a direct result of political instability. another political crisis in 2002 also abruptly halted tourism development (durbin & ratrimoarisaona 1996; pawliczek & mehta 2008; united states of america: department of state 2012). it is widely understood that without the support of the government through policy, both the conservation tourism industry in madagascar and the resource it depends on, as well as biological conservation in its own right, will be threatened (buckley 2010; durbin & ratrimoarisaona 1996; mittermeier et al. 2010; pawliczek & mehta 2008). in the closing section of pawliczek and mehta’s (2008) paper, they saw madagascar’s natural-area tourism industry (and associated conservation efforts) at a crossroads between: unstrategic tourism development for a few with exploitation of natural resources and pursuing non-sustainable and rapid results to boost the country’s development; or a harmonised mix of different forms of tourism (of which ecotourism is an important one) to develop the economy of the country by integrating the local population and improving their living standards, targeting a responsible and long-term vision in which nature resources will be exploited in a sustainable way. (p. 63) this was a reflection of what grosvenor, the then president of the national geographic society, said in the late 1980s (grosvenor 1987). at that point, extinctions were rife, natural-area tourism was in its infancy and foreign scientists were only just being allowed onto the island to undertake research. grosvenor (1987) closed his article by asking the question: ‘can madagascar avoid a new wave of extinctions?’ as an answer, he proposed: maybe. severe poverty still grips the island’s ten million people, and projections of continued forest losses are gloomy. but if there is a turn around, it will happen because of a commitment by the malagasy government to protect the country’s natural resources. (p. 5) it has been 25 years since this statement was made and it seems as though this question remains largely unanswered. both pawliczek and mehta (2008) and grosvenor (1987) understood that without political support, the conservation of the environment, and consequently the industries of ecotourism and ct that support it, will fail. without public spending on pas, enforced regulations on illegal natural resource consumption, international aid for conservation and development projects and with diminished trust in investing in madagascan tourism, the future of conservation tourism in madagascar is again in danger (ormsby & manni 2006; pawliczek & mehta 2008). in 2008, when pawliczek and mehta’s paper was written, the future was looking brighter for madagascar. ‘sleeping beauty’ was finally awakening to its ecotourism and ct potential and the positive step towards development was occurring. unfortunately for the rich biological diversity, the remaining fragile flora and fauna communities, those species yet to be discovered and those communities who depend on natural-area tourism for their livelihood, due to political instability, the darker path at the crossroads seems to have been taken. can this situation be turned around in the current political climate? is ct still a viable tool for biological conservation? recent observations made in andasibe–mantadia national park point to a positive turn being made in the tumultuous history of natural-area tourism and conservation in madagascar. the success of andasibe through local community engagement and conservation support top ↑ andasibe–mantadia national park is 155 km2 in extent and comprises a range of vegetation types. the andasibe section (formerly perinet reserve, 810 ha) comprises mid-altitude montane rainforest occurring at around 1000 m.a.s.l., whilst the mantadia section consists of mid-altitude (800 m.a.s.l. – 1300 m.a.s.l.) and lowland rainforest with an area of some 10 000+ ha. faunal components include a rich assemblage of mammals (10 species of lemur), birds (couas, vangas, ground rollers and two species of asity), reptiles (five species of chameleons, including parsons chameleon, leaf tailed gecko and madagascar tree boa) and amphibians. nature-based tourism activity and access is by guide-led excursions into the park along an extensive network of formed paths where bird watching and observations of lemurs and chameleons takes place. night-spotting activities are not allowed in the park but are led by guides in adjacent forested areas. the park has historically received about 20 000 visitors a year, with the analamazaotra section of the park (where habituated indri occur) being one of the most visited sites in madagascar (mittermeier et al. 2010). foreign visitors mostly occur in tour groups (numbers range from 5 or 10, up to 15) and at present there are few free independent travellers. domestic tourism is increasing in importance with groups of up to 40 people at a time from schools and universities also visiting the park.as already noted, stephenson (1993) previously found significant problems in connection with unsustainable natural-area tourism practices at perinet. a recent appraisal of the situation, however, found that many improvements have been made since the early 1990s and that most of the specific impacts observed by stephenson (1993) were not readily detectable (newsome et al. 2013). at the same time, as highlighted in other contexts earlier, there are some potential tourism impacts. based on observations conducted in 2011, newsome et al. (2013) additionally reported tourists leaving trails in order to locate and observe indri and other species of lemur. this tourist activity has caused a proliferation of informal trails and trampling of vegetation. some tourists were observed to call out at the indri in order to gain the animals attention for photographic purposes. sometimes there were up to 50 people at an indri sighting along with considerable turnover of visitors as people left and others joined the sighting during the viewing period that lasts from around 07:00 until around 12:30. despite these impacts and the presence of humans, the habituated indri groups do not show any disturbance behaviour (a. rakotonoely, pers. comm., 01 october 2011). in terms of the delivery of information and interpretation, the quality of guiding is variable and the main function of the guide appears to be in regard to locating and identifying species, but guides are also willing and able to answer questions. nevertheless, the situation at andasibe today must be viewed in the positive context of increased rainforest tourism globally. furthermore, and given the situation of reduced impacts overall as described above, there is now more emphasis on the importance of tourism in conserving remnant patches of rainforest and increased stakeholder understanding of the principles of ecotourism and the professionalisation of tour guiding over the last 10 years. andasibe is managed according to a park management plan, but the actual details contained in the plan are difficult to obtain. currently, the park has 40+ rangers whose duties include permit delivery, protection via enforcement of regulations, support for scientists and tour guiding. guides who are not rangers need approval to work in the park (figure 3) and the park runs training courses for guides. the guides association of madagascar (natural tour guide federation of madagascar) is now 20 years old. its main objective is to join tour guides into one association with the aim of training during the low season. the association works with experts, for example botanists from kew gardens and experts from birdlife international, and generally conducts courses lasting 3 days. however, courses can last for 1–2 weeks so that aspiring and junior guides can gain skills in the identification and ecology of plants, birds and reptiles with the aim of gaining a certificate or qualification which will aid them in gaining employment as tour guides. figure 3: cabinet located at andasibe national park with records of approved and registered tour guides working in the park. the majority of guides live in the village of andasibe. at andasibe, the local people, who are members of organised groups such as the association des guides d’ andasibe, the tambata guide association and the mitsingo local community, supply the majority of tour guiding services. the mitsingo local community is particularly involved in conservation work such as native tree planting and the implementation of a corridor project between mantadia national park and andasibe national park. the mitsingo forest station is a community reserve created by german conservationist, mr rainer in 1992–1993. the local community surrounding the park, such as mitsingo, works with international non-governmental organisations and the guides association that was originally created to help protect the forest. in 2009, vanga was developed as an english-speaking guide association with a focus on wildlife interpretation and conservation. the plan was to share knowledge especially between local guides with particular emphasis on the delivery of good interpretation. training is conducted during the low season and there is collaboration with experts such as botanists from the missouri botanical gardens and herpertologists from the capital city, antananarivo. it is estimated that there are at least 70, and as many as 100, guides working in andasibe–mantadia national park, most of which are from the local community of andasibe village (mittermeier et al. 2010).the economic and positive livelihood benefits of tourism to the local community living in and around andasibe village (population c. 14 000) are considerable (d. newsome, pers. obs., 01 october 2011). although no actual amount has been directly calculated, there are tangible indicators (protection of community forest, improved road conditions and village services) that demonstrate positive financial benefits that are gained from a percentage (50%) of park fees that flows from the park to the local community and employment of local people as guides, as well as the creation of ecotourism-related jobs. at andasibe, this has resulted in funding for road upgrades (figure 4) and other infrastructural developments such as schools, market place development, a dispensary and piped water. there is employment as guides (70 from andasibe village) and employment opportunities for the local community to work in hotels (in 1995 there was one hotel, compared with 10 hotels in 2011), in retail and as drivers. given these opportunities to earn a living directly associated with natural-area tourism there is a major incentive to protect the forest and wildlife. forest loss is a widespread problem (figure 5) in madagascar, but at andasibe the forest is valued for its ecological function as a generator of profits from natural-area tourism and exploitation of the park was not observed (d. newsome, pers. obs., 01 october 2011). in addition to international tourists (figure 1), the local community, school groups and university students are now part of the visitor profile at andasibe and such local tourism further engages malagasy to realise the importance of wildlife in their lives and the need to conserve habitat for tourism purposes. figure 4: main access road in andasibe village which has been improved via funding from tourism profits. roads in madagascar can be extremely rutted with many pot holes, which become waterlogged when wet and shedding dust when dry. hardening and sealing main access roads is a considerable improvement to village access and functionality. figure 5: slash and burn agriculture in the vicinity of andasibe national park. conclusion top ↑ natural-area tourism, comprising ecotourism and wildlife tourism, has the potential to contribute towards biological conservation through directly benefitting adjacent local communities (alexander 2000; buckley 2010; burns & sofield 2001). this can be achieved through increased economic exchange via employment and funding for infrastructure and buildings, as observed at andasibe–mantadia national park in eastern madagascar. the promotion, operation of ecotourism and the quality of guiding are fostered by mitsinjo and the local andasibe guides association who work in both parks. the role and value of the guides associations that are derived from the local community in promoting community-based conservation cannot be underestimated. not only are guides working within the borders of andasibe, they are also conducting night tours on the perimeter of the pa. profits from this provide a strong incentive to protect the forest not only within but outside designated borders, which expands the reach of conservation for both itself and for tourism purposes. although there were some observed environmental impacts associated with locating and viewing lemurs at andasibe, as similarly observed by stephenson (1993), the benefits of nature-based tourism in biological conservation far outweigh any of its impacts, such as trampling the undergrowth. because ecotourism is firmly established at andasibe, this, in itself, provides opportunities for ecotourism-related impacts to be studied, monitored and managed in the longer term, which should be an objective in all forms of sustainable tourism, including ct.although local people can see the benefits of wildlife tourism, they still need to be engaged and educated for biological conservation to be more fully realised. this extension of tourism into educating the local community about the need for biological conservation was shown by ormsby and manni (2006) to be occurring with park guides broadcasting weekly radio programmes to the local communities adjacent to masoala national park. this was not observed at andasibe; however, this extension of the guide’s responsibilities could potentially be applied to further the positive outcomes of biological conservation at andasibe. with the support of international partners, such as kew gardens (uk) and missouri botanical gardens (usa), further programmes such as this could be put into place, extending the reach of natural-area tourism to assist with biological conservation. larger, systemic issues such as poverty, the need for land for housing and agriculture and the reliance on forest products remain and have been made even greater by the recent political crisis in 2009. however, the case of andasibe demonstrates that natural-area tourism can be effective in contributing towards biological conservation and poverty alleviation for people who live near important biodiversity and protected areas. it thus seems that madagascar is now repeating the history of increasing tourism arrivals and development along with the associated recognition of the importance of the environment. at the same time, however, it appears that the hard work that has been put in place is being jeopardised through policies such the map being forgotten and unimplemented. buckley (2010) made the point that an adequate conservation framework needs to be in place for tourism to assist with ct goals. this is true in that conservation frameworks in madagascar have been created but have not had appropriate political space to be implemented. the previous government had attempted to do this by expanding the pa network in madagascar, but large portions of this remain poorly protected (goodman & benstead 2003). at the government policy level, much work remains to be done and the ecotourism potential of madagascar’s wildlife needs to be valued further. andasibe national park is an example of how conservation and tourism can work together in madagascar for the benefit of both local communities and the environment. however, as pawliczek and mehta (2008) have indicated, madagascar is like a ‘sleeping beauty’ with great potential for ct, but with so many ongoing threats, awakening to this potential is unfortunately not a simple task. acknowledgements top ↑ we thank alain rakotonoely for helpful discussions and assistance in the field. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions d.n. (murdoch university) conceived the article, conducted the fieldwork, wrote 50% of the text and supplied the diagrams. s.h. (murdoch university) conducted a literature review of conservation tourism and investigated the issues surrounding tourism and conservation in madagascar. references top ↑ anon., 2008, ‘environment: madagascar’, africa research bulletin: economic, financial and technical series 45(7), 17914–17915. http://dx.doi.org/10.1111/j.1467-6346.2008.01828.xalexander, s.e., 2000, ‘resident attitudes towards conservation and black howler monkeys in belize: the community baboon sanctuary’, environmental conservation 27(4), 341–350. http://dx.doi.org/10.1017/s0376892900000394 blumstein, d.t., fernandez-juricic, e., zollner, p.a. & garity, s.c., 2005, ‘inter-specific variation in avian responses to human disturbance’, journal of applied ecology 42(5), 943–953. http://dx.doi.org/10.1111/j.1365-2664.2005.01071.x blamey, r.k, 1997, ‘ecotourism: the search for operational definition’, journal of sustainable tourism 5(2), 109–130. braun, d., 2009, lemurs, rare forests threatened by madagascar strife, viewed 10 december 2012, from http://newswatch.nationalgeographic.com/2009/03/23/lemurs_threatened_by_madagascar_strife/ briassoulis, h., 2002, ‘sustainable tourism and the question of the commons’, annals of tourism research 29(4), 1065–1085. http://dx.doi.org/10.1016/s0160-7383(02)00021-x buckley, r., 2010, conservation tourism, cab international, wallingford. http://dx.doi.org/10.1079/9781845936655.0000 burns, g.l. & sofield, t.h.b., 2001, the host community: social and cultural issues concerning wildlife tourism, wildlife tourism research report series 4, cooperative research centre for sustainable tourism, gold coast. ceballos-lascuráin, h., 1996, tourism, ecotourism and protected areas: the state of nature-based tourism around the world and guidelines for its development, iucn, cambridge. christie, i.t. & crompton, d.e., 2003, republic of madagascar: tourism sector study, viewed 03 december 2012, from http://www.worldbank.org/afr/wps/wp63.pdf conservation international, 2012, where we work: madagascar and the indian ocean islands, viewed 06 december 2012, from http://www.conservation.org/where/priority_areas/hotspots/africa/ madagascar_and_the_indian_ocean_islands/pages/default.aspx curtin sustainable tourism centre, 2010, kimberely whale coast tourism: opportunities and threats, viewed 20 november 2012, from http://www.wilderness.org.au/pdf/kimberley_whalecoastreport.pdf donohoe, h.m. & needham, r.d., 2006, ‘ecotourism: the evolving contemporary definition’, journal of ecotourism 5(3), 192–210. http://dx.doi.org/10.2167/joe152.0 durbin, j.c. & ratrimoarisaona, s.-n., 1996, ‘can tourism make a major contribution to the conservation of protected areas in madagascar?’, biodiversity & conservation 5, 345–353. http://dx.doi.org/10.1007/bf00051778 fennell, d., 2008, ecotourism, 3rd edn., routledge, abingdon. goodman, s.m. & benstead, j.p., 2003, the natural history of madagascar, chicago university press, chicago. goodman, s.m. & benstead, j.p., 2005, ‘updated estimates of biodiversity and endemism for madagascar’, oryx 39(1), 73–77. http://dx.doi.org/10.1017/s0030605305000128 green, r. & higginbottom, k., 2001, the negative effects of wildlife tourism on wildlife, wildlife tourism research report series 5, viewed 06 december 2012, from http://www.crctourism.com.au/wms/upload/images/disc%20of%20images%20and%20pdfs/ for%20bookshop/documents/5green3.pdf grosvenor, g.m., 1987, ‘madagascar: a lot to lose’, national geographic 172, 5. harrison, d., 1997, ‘ecotourism in the south pacific: the case of fiji’, in world ecotour ’97 abstracts volume, p. 75, biosfera, rio de janiero. karanth, k.k., defries, r., srivathsa, a. & sankaraman, v., 2012, ‘wildlife tourists in india’s emerging economy: potential for a conservation constituency?’, oryx 46(3), 382–390. http://dx.doi.org/10.1017/s003060531100086x keane, a., ramarolahy, a.a., jones, j.p.g. & milner-gulland, e.j., 2011, ‘evidence for the effects of environmental engagement and education on knowledge of wildlife laws in madagascar’, conservation letters 4(1), 55–63. http://dx.doi.org/10.1111/j.1755-263x.2010.00144.x kruger, o., 2005, ‘the role of ecotourism in conservation: panacea or pandora’s box?’, biodiversity & conservation 14(3), 579–600. http://dx.doi.org/10.1007/s10531-004-3917-4 lansing, p. & de vries, p., 2007, ‘sustainable tourism or marketing ploy?’, journal of business ethics 72(1), 77–85. http://dx.doi.org/10.1007/s10551-006-9157-7 lindsey, p.a., alexander, r.r., du toit, j.t. & mills, m.g.l., 2005, ‘the potential contribution of ecotourism to african wild dog lycaon pictus conservation in south africa’, biological conservation 123(3), 339–348. http://dx.doi.org/10.1016/j.biocon.2004.12.002 mbaiwa, j.e., 2008, ‘the realities of ecotourism development in botswana’, in a. spenceley (ed.), responsible tourism: critical issues for conservation and development, pp. 205–223, earthscan, london. mill, r.c. & morrison, a.m., 1992, the tourism system, prentice hall, englewood cliffs. mittermeier, r.a., louis, e.e. jnr, richardson, m., schwitzer, c., langrand, o., rylands, a.b. et al., 2010, lemurs of madagascar, 3rd edn., conservation international, arlington. natural habitat adventures, 2012, ultimate madagascar adventure, viewed 10 december 2012, from http://www.nathab.com/africa/wild-madagascar/ newsome, d., dowling, r.k. & moore, s.a., 2005, wildlife tourism, channel view publications, clevedon. newsome, d., moore, s.a. & dowling, r.k., 2013, natural area tourism: ecology, impacts and management, 2nd edn., channel view publications, clevedon. ormsby, a. & manni, k., 2006, ‘ecotourism benefits and the role of local guides at masoala national park, madagascar’, journal of sustainable tourism 14(3), 271–287. http://dx.doi.org/10.1080/09669580608669059 pawliczek, m. & mehta, h., 2008, ‘ecotourism in madagascar: how a sleeping beauty is finally awakening’, in a. spenceley (ed.), responsible tourism: critical issues for conservation and development, pp. 41–68, earthscan, london. ploch, l. & cook, n., 2012, madagascar’s political crisis, viewed 30 november 2012, from http://www.fas.org/sgp/crs/row/r40448.pdf presidency of madagascar, 2006, madagascar action plan, viewed 03 december 2012, from http://siteresources.worldbank.org/madagascarextn/resources/map_madagascar.pdf royal botanic gardens: kew, 2012, map 1: protected areas of madagascar, viewed 30 november 2012, from http://www.kew.org/gis/projects/madagascar/mad_parks.html scales, i.r., 2012, ‘lost in translation: conflicting views of deforestation, land use and identity in western madagascar’, the geographical journal 178(1), 67–79. http://dx.doi.org/10.1111/j.1475-4959.2011.00432.x, pmid:22413174 self, r.m., self, d.r. & bell-haynes, j., 2010, ‘marketing tourism in the galapagos islands: ecotourism or greenwashing?’, international business and economics research journal 9(6), 111–126. sinclair, m.t., 1998, ‘tourism and economic development: a survey’, the journal of development studies 34(5), 1–51. http://dx.doi.org/10.1080/00220389808422535 sinha, b.c., qureshi, q., uniyal, v.k. & sen, s., 2012, ‘economics of wildlife tourism: contribution to livelihoods of communities around kanha tiger reserve, india’, journal of ecotourism 11(3), 207–218. http://dx.doi.org/10.1080/14724049.2012.721785 stem, c.j., lassoie, j.p., lee, d.r. & deshler, d.j., 2003, ‘how “eco” is ecotourism? a comparative case study of ecotourism in costa rica’, journal of sustainable tourism 11(4), 322–347. http://dx.doi.org/10.1080/09669580308667210 stephenson, p.j., 1993, ‘the impacts of tourism on nature reserves in madagascar: perinet, a case study’, environmental conservation 20, 262–265. http://dx.doi.org/10.1017/s0376892900023067 tisdell, c. & wilson, c., 2012, nature-based tourism and conservation: new economic insights and case studies, edward elgar, cheltenham. united nations, 1992, report of the united nations conference on environment and development, viewed 25 november 2012, from http://www.un.org/documents/ga/conf151/aconf15126-1annex1.htm united states of america: department of state, 2012, background note: madagascar, viewed 30 november 2012, from http://www.state.gov/r/pa/ei/bgn/5460.htm weaver, d.b., 2005, ‘comprehensive and minimalist dimensions of ecotourism’, annals of tourism research 32(2), 439–455. http://dx.doi.org/10.1016/j.annals.2004.08.003 weaver, d.b., 2008, ecotourism, 2nd edn., wiley australia, milton. weaver, d.b. & lawton, l.j., 2006, tourism management, wiley australia, milton. weaver, d.b. & lawton, l.j., 2007, ‘twenty years on: the state of contemporary ecotourism research’, tourism management 28(5), 1168–1179. http://dx.doi.org/10.1016/j.tourman.2007.03.004 wilson, c. & tisdell, c., 2003, ‘conservation and economic benefits of wildlife-based marine tourism: sea turtles and whales as case studies’, human dimensions of wildlife 8(1), 49–58. http://dx.doi.org/10.1080/10871200390180145 wollenberg, k.c., jenkins, r.k.b., randrianavelona, r., ramilamanana, r., ralisata, m., ramanandraibe, a. et al., 2011, ‘on the shoulders of lemurs: pinpointing the ecotouristic potential of madagascar’s unique herpetofauna’, journal of ecotourism 10(2), 101–117. http://dx.doi.org/10.1080/14724049.2010.511229 world commission on environment and development, 1987, our common future, oxford university press, new york. article information authors: judith m. kruger1 sandra macfadyen1 affiliations: 1south african national parks, skukuza, south africa correspondence to: judith kruger email: judith.botha@sanparks.org postal address: po box 178, skukuza 1350, south africa dates: received: 21 june 2010 accepted: 08 mar. 2011 published: 13 may 2011 how to cite this article: kruger, j.m. & macfadyen, s., 2011, ‘science support within the south african national parks adaptive management framework’, koedoe 53(2), art. #1010, 7 pages. doi:10.4102/koedoe.v53i2.1010 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) science support within the south african national parks adaptive management framework in this essay... open access • abstract • introduction • sanparks science support process    • data collection    • data storage and documentation    • data dissemination and sharing    • data analysis • conclusion • acknowledgements • references abstract (back to top) ‘behind all good science is good science support.’ implementing a successful strategic adaptive management (sam) framework requires an effective science support structure. this structure must be effective in all areas of data management, starting with data collection and ending with the dissemination of knowledge, to facilitate timeous management decisions and associated actions. accordingly, south african national parks has embraced the use of various technologies to enable the effective implementation of a functional support structure. this paper described these technologies and discussed how they benefit the implementation of the sam framework. conservation implications: the importance of functional support structures in science and conservation management is frequently undervalued in a system where emphasis is placed on scientific products. in order to promote research and facilitate analysis, sound data management practices are essential to integrating knowledge into an organisation’s institutional memory. introduction (back to top) south african national parks’ (sanparks) network of 19 protected areas traverse extensive environmental gradients and biodiversity and are exposed to ever-changing and intensifying global population pressure (balmford et al. 2001). in response, sanparks has adopted an adaptive management approach to biodiversity conservation. this style of management recognises the heterogeneous nature of ecological systems (pickett, cadenasso & benning 2003) by ensuring management actions are spatially dynamic over time. if scientists and managers do not adapt and learn from their management actions, conservation will fail, to the detriment of the species and ecosystems being protected. moreover, if adaptive management is to be effective in sanparks, science support structures must be in place to sustain the research needed to assess the outcomes of monitoring and management interventions. these structures should be aimed at capturing the learning that individuals develop, and integrating it into the collective or institutional memory of the group (salafsky, margoluis & redford 2001). the process of science support (figure 1) is rooted in the development of systems for storing, organising and accessing information. this process includes, (1) the ongoing collection of new data, (2) the storage and documentation of data, (3) the dissemination and sharing of data, (4) data analysis, (5) further research and (6) the promotion of knowledge generation. the ideal result is a distributed network centre with data structures and standards in place to allow scientists and managers to apply and adjust management strategies appropriately (louisiana protection and restoration 2009). however, sanparks represents a diverse range of ecosystems, which is mirrored by the diversity of environmental data that must be collected to facilitate research in sanparks. as a result, data collection methods must be dynamic as they may include large ranges of biotic and abiotic variables, which could produce vast and heterogeneous datasets. not surprisingly, it has proven difficult to build data management and analytical systems to accommodate these data in general (jones et al. 2006). traditionally, each type of data is stored in customised databases that are not connected to one another, so the data are largely inaccessible for crosscutting analysis and modelling. figure 1: the south african national parks (sanparks) science support process indicating the relationship between the six process steps. in the case of sanparks’ thresholds of potential concern (tpcs; see roux & foxcroft 2011), which form an integral part of the strategic adaptive management (sam) framework, a number of different datasets are required for individual analyses. for this reason, creating a streamlined system of data collection, analysis, presentation of results and knowledge feedback, will ensure timeous and accurate outputs. in addition to maintaining this technology, organisations also need to develop human capital and institutional memory to ensure continuity in the event that a key individual leaves the organisation. in most cases, people are the primary driving forces behind change or the adoption of new technology within an organisation. if one or more of these drivers, or change agents, disappear from the system, the programmes previously under their supervision may be at risk of collapse if the proper structures are not in place to absorb the impact. we describe each step of the science support process in figure 1, indicating the innovations and experience to be gained from it. sanparks science support process (back to top) data collection sanparks has a wide variety of data that must be collected to monitor the state of the environment successfully. to ensure the accuracy of the data is preserved and the collection process standardised, a hand-held computer application (cybertracker) is customised for each monitoring programme. this icon-based system allows both literate and nonliterate field workers to record observations, with latitude and longitude coordinates, using a personal digital assistant (pda) device that features an integrated global positioning system (gps). in this way, data collected in the field are simultaneously captured electronically using graphic check lists, which enhance data accuracy. the cybertracker’s icon-based system also allows computer illiterate field workers to record data with minimal training, thereby enhancing organisational skills development and capacity building. as a result, this data collection process can enable field rangers to contribute directly to the management and research of each park by simply collecting basic environmental data during their regular daily patrols (e.g. distribution of rare and endangered species, availability of surface water and disease outbreaks). in addition, the integrated gps ensures that separate gps skills are not necessary to record latitude and longitude coordinates of each observation, whilst the moving map functionality allows the user to pinpoint his position on a 1:50 000 or 1:250 000 topographical map or aerial photograph at any time (see macfadyen 2007 for operational details). the value of the cybertracker system was first recognised by mr douw (‘swannie’) swanepoel in 2000, when 44 palm iiie organisers and 44 garmin 12xl gps devices interfaced via a cable were purchased for the kruger national park (knp). in 2003, a further 120 handspring visor deluxe pdas with integrated magellan gps companions were purchased. as a result, each of the 22 management sections in the knp were issued with five pdas for deployment with field rangers and five additional pdas were issued to the scientific services division in skukuza for various research projects. since then, collective fundraising initiatives have raised over r200 000.00 and enabled sanparks to purchase new pocket pcs (with integrated gps) for many other parks. as a result, cybertracker is operational in all south african national parks (figure 2a). established databases include ranger patrols, vegetation condition assessments, animal behaviour monitoring, rare and endangered species listings, and invasive species distribution mapping. each database has been customised to facilitate conservation management, research and monitoring (figure 2b). the largest of these databases is the ranger patrol system, which is managed by a network of section rangers and operated by a core of field rangers across all parks. the system is customised in-house with an icon-based interface that features english and local language descriptions (e.g. shangaan) for the collection of daily patrol information and basic environmental data. according to macfadyen (2009), the information gathered during these ranger patrols is used by sanparks management to: • plan section patrols for area-integrity mapping • provide an early warning system for disease outbreaks • identify trends in illegal exit and entry points • enable the detection and control of invasive alien species • report fence-breaks to the state veterinary department for animal health purposes. figure 2: (a) country-wide distribution of the south african national parks’ (sanparks) cybertracker program, with operational databases (b) in 18 of 21 sanparks customised to facilitate conservation management, research and monitoring. in addition, the programme supports local community projects such as the xilongana craft programme in the thulamahashe region, which designs and produces protective canvas cases for the pocket pcs. in general, the ranger patrol system aims to enhance park-specific environmental management and research through field monitoring (see macfadyen 2007 for more detail about the sanparks cybertracker program). scientists use the collected data to facilitate research and assist in making informed management decisions regarding rare species monitoring (endangered wildlife trust 2011; murn 2009), invasive species research (dietemann, lubbe & crewe 2006; foxcroft et al. 2009; foxcroft et al. 2010; hui et al. 2011), fire mapping, archaeological inventorying, species distribution mapping, ecosystem interactions (burkepile et al. 2008; somers & hayward 2009), and veterinary–wildlife interfaces (brahmbhatt et al. in press; ferguson & hanks 2010; jori et al. 2009). data storage and documentation once data has been collected it needs to be correctly archived and, more importantly, clearly documented in order to maintain its long-term usability. sanparks uses a flexible, structured metadata standard called ecological metadata language (eml), which is a metadata standard developed by the ecological community in the usa (fegraus et al. 2005). eml was developed to address the lack of dataset documentation and to provide structure to traditionally unstructured information (i.e. tabular data with no metadata; michener et al. 1997). the eml standard evolved by incorporating various elements from other metadata language standards and is supported by a wide range of users ensuring continual improvement. as the eml structure is modular and easily extensible it can be used for the documentation of both metadata and datasets in a number of different data formats. eml requires multiple levels of documentation to be specified. the first level includes dataset identification, which is documented in the title, abstract and keyword fields. the next level, known as the discovery level, includes information on both the geographic extent and the temporal coverage of the datasets. the geographic extent is captured with spatial coordinates that can then be used to plot coordinates on a map. applications can leverage both the geographical and date–time fields for dataset discovery. the evaluation level metadata includes information about the methods used, as well as the project level information. these methods indicate how the data was collected and may be described as a set of hierarchical processes with substeps. the third level contains access information, which indicates who may change and read the data and metadata. furthermore, this information also describes where the data can be obtained and the required data format. the final level is the logical model information that describes the structure of data tables and their variables. each variable requires a name, description and a measurement type to be captured. once the eml has been created for a particular data package, which includes the metadata and the data tables, it can be uploaded into a data repository where it can be browsed, downloaded and archived. the knowledge network for biocomplexity (knb; http://knb.ecoinformatics.org/index.jsp) now has 16 partner organisations who contribute eml data and metadata to a global data repository, including sanparks. an added advantage of being a part of this global network is the replication of data between sites, which reduces the risk of data loss. the access rules do not change with replication, which means that the dataset owner can still determine who is allowed to access the data without needing to be concerned that the information will be amended once replication occurs. collating accurate metadata about heterogeneous datasets can be time consuming and therefore difficult without supporting software tools. as a result, sanparks adopted the free software product morpho (higgins, berkley & jones 2002) to create eml metadata to be uploaded into the sanparks data repository (http://dataknp.sanparks.org/sanparks). morpho creates eml using a wizard front end for the simpler metadata, with an eml tree for the more complex metadata. morpho is then used to upload the data to the sanparks data repository, which can then be searched through a web interface. an extension to this system allows spatial layers, with standard federal geographic data committee metadata created through arcgis, to be incorporated into the repository. this additional functionality allows searches for spatial and nonspatial data simultaneously. data dissemination and sharing following the capture of data (and associated metadata) in the data repository, software tools can be automated to access this information and perform various analyses. the network of data repositories allows users to search for data from any of the participating sites using keywords or localities of interest (http://dataknp.sanparks.org/sanparks). this promotes integration of global ecological data and allows for collaboration across scales, as well as transcontinental analysis. data analysis although data can be searched and shared across all sites globally, as indicated earlier, data stored in the sanparks database are used in standardised routine analyses to assess whether the thresholds of specific ecological variables have been exceeded. these thresholds provide upper and lower limits of acceptable change in specific ecological indicators (roux & foxcroft 2011). the tpcs are therefore developed to provide early warning signs of impending unacceptable changes. for example, one of the key concerns and focus areas for tpc development has been on river management (for a full treatise see mcloughlin et al. 2011 and pollard, du toit & biggs 2011). excessive abstraction of water upstream of the knp has lowered flow volumes below, inhibiting the functional processes of the river ecosystems. these thresholds, or in-stream flow requirements (ifr), have been determined for all perennial rivers. the monitoring data is analysed for each river in order to determine whether these thresholds (ifrs) have been exceeded. to accomplish these kinds of cross-scale and cross-disciplinary analyses, an analytical tool is required which can interface with a variety of data sources and provide access to commonly used analytical tools. one such approach is to use a scientific workflow system, such as kepler (ludäscher et al. 2006), to orchestrate the analysis (figure 3). scientific workflows in general and kepler, in particular, have the capability to access multiple analytical systems such as r and matlab, as well as other statistical and modelling systems. each workflow can consist of a series of analytical steps, starting from data cleaning and concluding with complex analytical procedures to produce reports and visual outputs. for example, figure 4 shows the results of the river-flow analysis, indicating periods when the threshold was exceeded (i.e. where river flow was below minimum requirements, indicated in red). figure 3: the kepler scientific workflow application uses river-flow (water volume) data from the data repository to analyse whether predefined thresholds have been exceeded. figure 4: river flow analyses for the letaba river for the period 01 january 2007 – 31 december 2007, where the minimum in-stream flow requirement (ifr) is determined for all the main rivers in the kruger national park, providing a threshold against which to examine daily flow rates. the ifr is measured in m3/second. using a workflow approach brings with it a number of advantages. firstly, it enables users to determine clearly the dataset version used for a particular analysis, thereby reducing errors. this allows analyses to be rerun on various versions of the datasets. secondly, the workflow procedure and the analytical outputs can also be uploaded to the data repository, allowing the same workflow procedure to be rerun later and the outputs archived. sanparks has adopted kepler to analyse the data collected through routine monitoring programmes (figure 3) and thus detect any breaches of the tpcs. however, park managers need these analyses to be run on a regular basis to enable rapid decision making. additionally, the reports and figures generated by the analyses need to be shared broadly with a variety of management stakeholders. as such, development has begun on a web-based execution and reporting system that will make all outputs broadly accessible to the necessary stakeholders. conclusion (back to top) adaptive management is a systematic, rigorous and scientifically defensible approach to learning based on outcomes of management actions (walters & holling 1990). strategic adaptive management, as practiced in sanparks, is more pragmatic and goal-orientated and is reliant on an effective science and management partnership (rogers 1998). as a result, the criteria for the success of sam include both biological and social measures, encompassing learning and the application of new knowledge (kleiman et al. 2000). however, a gap exists at both the social and technological levels of sam, including institutionalisation and implementation (coleman et al. 2010; rogers 1998). although adaptive management practices should include the documentation of the research processes followed and the results achieved, on which management decisions can be based, a technological divide often prevents competent data management. the science support tools we described here aim to harness current developments in technology to facilitate the implementation of the adaptive management process. acknowledgements (back to top) we would like to acknowledge mr douw (‘swannie’) swanepoel posthumously for introducing the cybertracker program to sanparks and for his unyielding dedication to the knp and conservation in general. we would further like to acknowledge the support of various funders who have supported the sanparks cybertracker program over the years, namely, deutsche gesellschaft für internationale zusammenarbeit, sappi limited, mrs erna goodwin estate, thubelisha homes, richtersveld poverty relief unit, unitrans, medscheme, west rand honorary rangers, sanparks forum moderators and members (http://www.sanparks.org/forums/), and over 50 members of the general public (http://www.sanparks.org/parks/kruger/conservation/scientific/ donations/cybertracker). finally, we would like to acknowledge the staff at the national centre for ecological analysis and synthesis for development and implementation of these tools, as well as the a.w. mellon foundation for funding. references (back to top) balmford, a., moore, j.l., brooks, t., burgess, n., hansen, l.a., williams, p., et al., 2001, ‘conservation conflicts across africa’, science 291, 2616−2619. doi:10.1126/science.291.5513.2616, pmid:1128337 brahmbhatt, d.p., fosgate, g.t., dyason, e., budke, c., gummow, b., jori, f., et al., in press, ‘contacts between domestic livestock and wildlife at the kruger national park interface of the republic of south africa’, preventive veterinary medicine. burkepile, d., smith, m.d., fynn, r.w.s. & knapp, a.k., 2008, ‘interactions between herbivore size and fire regime on plant community structure’, in south african national parks (eds.), reports on scientific research projects undertaken in the savanna parks of the south african national parks during 2008. project reports knp 15/1/2–2008, p. 23, sanparks, pretoria. coleman, a.m., wigmosta, m.s., lane, l.j., tagestad, j.d. & roberts, d., 2010, ‘gis-based adaptive management decision support system to develop a multi-objective framework: a case study utilizing gis technologies and physically-based models to achieve improved decision making for site management’, journal of map and geography libraries 4(2), 269−284. dietemann, v., lubbe, a. & crewe, r.m., 2006, ‘human factors facilitating the spread of a parasitic honeybee in south africa’, journal of economical entomology 99, 7−13. doi:10.1603/0022-0493(2006)099[0007:hfftso]2.0.co;2 endangered wildlife trust, 2011, ‘the endangered wildlife trust: birds of prey programme. registered sanparks research project’, viewed 04 february 2011, from https://www.ewt.org.za/whatwedo/ourprogrammes/birdsofpreyprogramme.aspx fegraus, e., andelman, s.j., jones, m.b. & schildhauer, m., 2005, ‘maximizing the value of ecological data with structured metadata: an introduction to ecological metadata language (eml) and principles for metadata creation’, bulletin of the ecological society of america 86(3), 158−168. doi:10.1890/0012-9623(2005)86[158:mtvoed]2.0.co;2 ferguson, k. & hanks, j. (eds.), 2010, ‘fencing impacts: a review of the environmental, social and economic impacts of game and veterinary fencing in africa with particular reference to the great limpopo and kavango-zambezi transfrontier conservation areas’, mammal research institute, pretoria, viewed 04 february 2011, from http://www.wcs-ahead.org/gltfca_grants/grants.html foxcroft, l.c., richardson, d.m., rouget, m. & macfadyen, s., 2009, ‘patterns of alien plant distribution at multiple spatial scales in a large national park: implications for ecology, management and monitoring’, diversity and distributions 15, 367−378. doi:10.1111/j.1472-4642.2008.00544.x foxcroft, l.c., jarošík, v., pyšek, p., richardson, d.m. & rouget, m., 2010, ‘protected area boundaries as a natural filter of plant invasions from surrounding landscapes’, conservation biology 25(2), 400–405. doi:10.1111/j.1523-1739.2010.01617.x higgins, d., berkley, c. & jones, m.b., 2002, ‘managing heterogeneous ecological data using morpho’, in j. kennedy (ed.), proceedings of the 14th international conference on scientific and statistical database management, washington dc, usa, july 24–26, 2002. hui, c., foxcroft, l.c., richardson, d.m. & macfadyen, s., 2011, ‘defining optimal sampling effort for large-scale monitoring of invasive alien plants: a bayesian method for estimating abundance and distribution’, journal of applied ecology, viewed n.d., from http://onlinelibrary.wiley.com/doi/10.1111/j.1365-2664.2011.01974.x/abstract, doi:10.1111/j.1365-2664.2011.01974.x jones, m.b., schildhauer, m., reichman, o.j. & bowers, s., 2006, ‘the new bioinformatics: integrating ecological data from the gene to the biosphere’, annual review of ecology, evolution, and systematics 37, 519−544. doi:10.1146/annurev.ecolsys.37.091305.110031 jori, f., vosloo, w., du plessis, b., bengis, r., brahmbatt, d., gummow, b., et al., 2009, ‘a qualitative risk assessment of factors contributing to foot and mouth disease outbreaks on the western boundary of kruger national park’, revue scientifique et technique oie 28(3), 917−933. kleiman, d.g., reading, r.p., miller, b.j., clark, t.w., scott, j.m., robinson, j., et al., 2000, ‘improving the evaluation of conservation programs’, conservation biology 14(2), 356−365. doi:10.1046/j.1523-1739.2000.98553.x louisiana protection and restoration, 2009, ‘final technical report: adaptive management appendix’, lacpr, us army corps of engineers, new orleans district, mississippi valley division, new orleans, pp. 1−14, viewed n.d., from http://lacpr.usace.army.mil/default.aspx?p=s&t=15&i=27 ludäscher, b., altintas, i., berkley, c., higgins, d., jaeger-frank, e., jones, m., et al., 2006, ‘scientific workflow management and the kepler system’, workflow in grid systems. concurrency and computation: practice & experience 18(10, special issue), 1039−1065. macfadyen, s., 2007, ‘cybertracker: conservation in the palm of our hands’, viewed 04 february 2011, from http://www.sanparks.org/parks/kruger/conservation/scientific/gis/cybertracker_poster.pdf macfadyen, s., 2009, ‘south african national parks cybertracker programme’, viewed 04 february 2011, from http://www.sanparks.org/parks/kruger/conservation/scientific/gis/cybertracker.php matlab, 2004, computer software, the mathworks inc., natick. mcloughlin, c.a., deacon, d., sithole, h. & gyedu-ababio, t., 2011, ‘history, rationale, and lessons learned: thresholds of potential concern in kruger national park river adaptive management’, koedoe 53(2), art. #996, 27 pages. doi:10.4102/koedoe.v53i2.996 michener, w.k., brunt, j.w., helly, j.j., kirchner, t.b. & stafford, s.g., 1997, ‘non-geospatial metadata for the ecological sciences’, ecological applications 7, 330−342. doi:10.1890/1051-0761(1997)007[0330:nmftes]2.0.co;2 murn, c., 2009, ‘ecological characteristics of the white-headed vulture trigonoceps occipitalis’ in southern africa, in south african national parks (eds.), reports on scientific research projects undertaken in the savannah national parks of south african national parks during the year 2009 and projects that were completed during 2008. research project reports 2009, knp 15/1/2-2009, p. 157, sanparks, pretoria. pickett, s.t.a., cadenasso, m.l. & benning, t.l., 2003, ‘biotic and abiotic variability as key determinants of savanna heterogeneity at multiple spatiotemporal scales’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 22−40, island press, washington dc. pollard, s., du toit, d. & biggs, h.c., in press, ‘river management under transformation:the emergence of strategic adaptive management of river systems in the kruger national park’, koedoe. r, version 2.9.0, 2009, computer software, the r foundation for statistical computing, s.l. rogers, k., 1998, ‘managing science/management partnerships: a challenge of adaptive management’, conservation ecology 2(2), 1−4. roux, d.j. & foxcroft, l.c., 2011, ‘the development and application of strategic adaptive management within south african national parks’, koedoe 53(2), art. #1049, 5 pages. doi:10.4102/koedoe.v53i2.1049 salafsky, n., margoluis, r. & redford, k., 2001, ‘adaptive management: a tool for conservation practitioners’, biodiversity support program, pp. 1−99, viewed n.d., from http://www.rmportal.net/library/content/tools/biodiversity-conservation-tools/putting-conservation-in-context-cd/adaptive-management-resources/5-5-a.pdf salafsky, n., margoluis, r., redford, k.h. & robinson, j.g., 2002, ‘improving the practice of conservation: a conceptual framework and research agenda for conservation ccience’, conservation biology 16(6), 1469−1479. doi:10.1046/j.1523-1739.2002.01232.x somers, m. & hayward, m., 2009, ‘impact of climate change on predator prey preferences’, in south african national parks (eds.), reports on scientific research projects undertaken in the savannah national parks of south african national parks during the year 2009 and projects that were completed during 2008. research project reports 2009, knp 15/1/2–2009, p. 215, sanparks, pretoria. walters, c.j. & holling, c.s., 1990, ‘large-scale management experiments and learning by doing’, ecology 71(6), 2060−2068. doi:10.2307/1938620 reviewer acknowledgement http://www.koedoe.co.za open access koedoe recognises and acknowledges the value and importance of reviewers in the publication process – not only in shaping the individual manuscript, but also in upholding the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely, productive peer review process. we would like to take this opportunity to thank all reviewers who participated in shaping this volume of koedoe: we appreciate the time taken to perform your review successfully. page 1 of 1 in an effort to facilitate the selection of appropriate peer reviewers for koedoe, we ask that you take a moment to update your electronic portfolio at www.koedoe.co.za for our records. this will allow us better access to your areas of interest and expertise, in order to match reviewers with submitted manuscripts. if you would like to become a reviewer, please visit the journal website and register as a reviewer. to access your details on the website, you will need to follow these steps: 1. log into the online journal at http://www. koedoe.co.za 2. in your ‘user home’ [http://www.koedoe. co.za/index.php/koedoe/ user] select ‘edit my profile’ under the heading ‘my account’ and insert all relevant details, bio statement and reviewing interest. 3. it is good practice as a reviewer to update your personal details regularly, to ensure contact with you throughout your professional term as reviewer to koedoe. please do not hesitate to contact me if you require assistance in performing this task. margo martens submissions@koedoe.co.za tel: +27 (0)21 975 2602 fax: +27 (0)21 975 4635 koedoe african protected area conservation and science anna songhurst arne witt astrid von kotze brian taylor bruce mann charmaine uys colin attwood darragh woodford dian spear dianne scott geoffrey howard gretel van rooyen gus mills guy castley hein stapelberg hendrik sithole hervé fritz ian russell isaac mapaure jacques delabie jean-louis janeau jeff hughes jürgen dengler ladislav mucina lesley henderson lucy king mark robertson megan griffiths mervyn lötter niels dreber olivier bonnet paul cross paul funston philip desmet piet-louis grundling robyn bushell samson chimphango simon power stefan siebert steve mccool theo mostert ute schmiedel wendy annecke 140 article information authors: jan a. venter1,2 bruce q. mann3 affiliations: 1department of biodiversity conservation, eastern cape parks and tourism agency, south africa 2school of life sciences, university of kwazulu-natal, westville campus, south africa 3oceanographic research institute, marine parade, south africa correspondence to: jan venter postal address: po box 11235, east london 5213, south africa dates: received: 30 june 2011 accepted: 02 june 2012 published: 10 sept. 2012 how to cite this article: venter, j.a. & mann, b.q., 2012, ‘preliminary assessment of surf-zone and estuarine line-fish species of the dwesa-cwebe marine protected area, eastern cape, south africa’, koedoe 54(1), art. #1059, 10 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1059 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. preliminary assessment of surf-zone and estuarine line-fish species of the dwesa-cwebe marine protected area, eastern cape, south africa in this checklist... open access • abstract • introduction • study area • research method and design    • study sites    • catch and effort data • results    • species diversity, status and composition    • size composition    • catch per unit effort       • sample site catch per unit effort       • species-specific catch per unit effort • ethical considerations • discussion • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ a preliminary assessment of surf-zone and estuarine line fish was carried out in the dwesa-cwebe marine protected area (mpa), on the wild coast, south africa. the purpose was to provide baseline data on inshore line-fish stocks in the mpa. a total of 28 species was recorded, of which 53% have a conservation status reflecting some concern and 43% are endemic to southern africa. this highlights the value of the mpa for protection of important line-fish species. within the mpa, localised differences were detected in species diversity, size frequency and catch per unit effort between unexploited and illegally exploited areas. these differences were more prominent in slow growing, long-lived species. it thus appears that illegal exploitation is negatively affecting fish populations within the mpa, which counteract and potentially could eliminate the benefits of fish protection typically associated with no-take mpas. these results highlight the need for improved law enforcement and better communication with neighbouring communities to increase awareness. it is further recommended that the current no-take status of the mpa should be maintained. in addition, baseline fisheries information was collected on certain fish species that could be used to inform future conservation management of the mpa.conservation implications: the dwesa-cwebe marine protected area is unique and important for the conservation of key surf zone and estuarine fish species. however there is a significant risk to the fish populations due to illegal exploitation. key interventions should include enhanced law enforcement but, more important, the creation of alternative livelihoods and long term sustainable benefits to local communities. introduction top ↑ research and monitoring of marine biodiversity features is an essential component of the management of marine protected areas (mpas). this is particularly true for inshore line-fish species, as they are predominantly over-exploited (cowley, brouwer & tilney 2001). marine protected areas can play an important role in providing a refuge for over-exploited line-fish stocks and the ’spillover effect‘ is an important ecosystem service provided to adjacent exploited areas (bennett & attwood 1991; cowley et al. 2001; mcclanahan & mangi 2000; kerwath et al. 2008). many surf-zone line-fish species have both recreational and subsistence value and could thus be considered to have significant socio-economic importance along the wild coast (fielding et al. 1994; pradervand 2004). the dwesa-cwebe mpa was formally declared as a marine reserve in 1991 (transkei environmental conservation decree no. 9 of 1992), and re-proclaimed in terms of the marine living resources act (act no. 18 of 1998) in 2000. the entire reserve is currently zoned as a no-take mpa, where no consumptive marine resource use is permitted. however, enforcement has been problematic since the mpa’s proclamation and certain sections of the mpa continue to be utilised illegally by local community members, hotel guests and holiday cottage owners residing within the dwesa-cwebe nature reserve. currently, considerable pressure is being placed on management authorities to rezone certain sections of the mpa to accommodate subsistence and recreational fishing. however, a recent report strongly advised against such action, due to the potential negative impacts on line fish populations (fielding 2010). little data exist on the surf-zone and estuarine line-fish stocks within the mpa, and thus a monitoring project was initiated at the beginning of 2009 with the following objectives, (1) to provide baseline data on the surf-zone and estuarine line-fish species assemblage, (2) to document size composition and relative abundance of key inshore line-fish species and (3) to gain an understanding of the impacts of current illegal fishing on inshore line-fish species within the mpa. this paper presents the preliminary results of this project for the years 2009–2010. study area top ↑ the dwesa-cwebe marine protected area is located on the wild coast, approximately 120 km northeast of east london on the east coast of south africa (figure 1). the mpa incorporates approximately 16 km of mainly rocky shore coastline and extends 6 nautical miles (10.8 km) out to sea. figure 1: the location of the dwesa-cwebe marine protected area with the dwesa, cwebe and mbashe sampling sites indicated. the mpa is located in a transition zone between the agulhas and natal bioregions (lombard et al. 2004; maree, booth & whitfield 2000). because of its location within this region of transition, the mpa is considered to be in a sensitive area that is not replicated elsewhere on the coast of south africa (fielding 2010). the mbashe estuary, situated centrally within the mpa, ranks 28th in terms of the conservation importance of estuaries in south africa (turpie et al. 2002). the surf-zone adjacent to the mbashe mouth is also considered to be a key spawning aggregation site for white steenbras (lithognathus lithognathus), the stock of which has collapsed (bennett 1993; mann 2000). research method and design top ↑ study sites two sample sites were initially identified where research fishing took place. these were the cwebe sample site and the dwesa sample site, located to the north-east and south–west of the mbashe estuary mouth respectively (figure 1). each sample site was 3.6 km long and incorporated representative portions of the available inshore habitats. it was known from field ranger patrol data, law enforcement records and from physical signs left by shore fishers, that the cwebe sample site received more illegal fishing pressure than the dwesa sample site (ecpta, unpublished data). at the end of 2010, a third sample site, namely the mbashe estuary, which stretches from the estuary mouth up to the old pont ± 5 km upstream, was also incorporated into the study area (figure 1). catch and effort data detailed catch and effort data were collected by means of research fishing, similar to the methods described by attwood (2003). four-day sampling trips took place in the months january, may, july and october 2009 as well as february, august and october 2010. the number of anglers used during sampling trips varied between 4 and 8. fishing took place during daylight hours (between 07:00 and 17:00) using a variety of baits (mainly pilchards, squid, prawns and red bait). an equal amount of time, in terms of the number of days, was spent fishing in the different sampling areas. all fish caught were carefully handled, measured to the nearest millimetre and released unharmed. barbless hooks were used to reduce release mortality. catch per unit effort (cpue) was calculated as the number of fish caught per angler per hour. it was assumed that cpue was directly related to relative abundance and takes the form cpue = nq, where n is abundance and q is the catch ability factor that mediates the relationship (bennett & attwood 1991). the catch per unit effort cpue was calculated as: where e was calculated as ∑hi per day with h as the hours fished by angler i, c was calculated as ∑fi per day where f is the number of fish caught by angler i . sampling effort was spread over the sampling sites by setting time constraints. once an angler started fishing at an area within the sample site, he had to stay there for at least one hour but could not stay longer than two hours. when an angler moved, he had to move a distance of more than 50 m. trimble geoxm global positioning system (gps) handheld computers (trimble navigation limited, sunnyvale, california, usa) with cybertracker software (liebenberg 2003) were used to collect data in the field. at the end of each sampling day, all data were transferred to a laptop computer. fish species status was described according to their stock status (griffiths, attwood & thomson 1999) and/or according to their international union for conservation of nature (iucn) red list category if they had been assessed (iucn 2011). differences in size structure, site-based cpue and species-based cpue between the dwesa and cwebe sample sites were tested using t-tests. results from the mbashe sample site were not included in the statistical analysis because of insufficient data. results top ↑ species diversity, status and composition a total of 28 species was recorded during the study period (table 1). the dwesa site produced more species than the cwebe site and the mbashe estuary (table 2). of the total of 28 species, five were cartilaginous fish and 23 were bony fish species. line fish species diversity varied between the sites, with 17 fish families being sampled in total and the dwesa sample site producing 15, the cwebe sample site 13 and the mbashe only four families (table 2). twelve of the species sampled were southern african endemics, with the dwesa sample site producing 11, the cwebe sample site 10 and the mbashe site zero (table 2). the stock status of the species sampled included one optimally exploited, three over-exploited, six collapsed, 14 unknown and three probably over-exploited or collapsed species (table 2; mann 2000). one species was classified as endangered, two were classified as vulnerable, two near-threatened and two least concern, according to the iucn red list for threatened species (table 2; iucn 2011). these included two serranid species and the rest were cartilaginous species. the other 21 species, which were all bony fish species, have not had their conservation status evaluated except for l. lithognathus, which was listed as ’lower risk/conservation dependent‘, by older iucn criteria (table 2). table 1: a list of the fish species sampled in the dwesa-cwebe marine protected area with the percentage composition indicated. table 2: a summary of the species caught (number of families and species) and their stock status according to mann (2000) and their international union for conservation of nature red list categorisation. the most abundant fish species caught in the mpa during the study period were from the families sparidae, sciaenidae, serranidae and rhinobatidae (table 1). species such as austroglossus sp., gymnothorax undulatus and amblyrhynchotes honckenii, which technically are not typical line-fish species, were occasionally sampled, but were included in the results for species inventory purposes (table 1). size composition only a few species were sampled in sufficient numbers in order to compare size composition between the different sample sites. these included pachymetopon grande, diplodus sargus capensis, argyrosomus japonicus, epinephelus andersoni and rhinobatos annulatus. although less abundant, p. grande in the dwesa sample site were significantly larger than those in the cwebe sample site (figure 2a and table 3). figure 2: the length frequency distribution of (a) pachymetopon grande (cwebe n = 148; dwesa n = 70), (b) diplodus sargus capensis (cwebe n = 88; dwesa n = 233), (c) argyrosomus japonicus (cwebe n = 46; dwesa n = 94; mbashe n = 30), (d) epinephelus andersoni (cwebe n = 23; dwesa n = 55) and (e) rhinobatos annulatus (cwebe n = 54; dwesa n = 51; mbashe n = 11) in the dwesa, cwebe and mbashe sample areas. table 3: results of the statistical comparison of mean fish sizes between the dwesa and cwebe sample sites for five dominant species sampled in the dwesa-cwebe marine protected area. differences in sizes of d. s. capensis between those caught in the dwesa and cwebe sites were not significant (figure 2b and table 3). argyrosomus japonicus sampled in the mbashe were smaller than those caught in the surf-zone (figure 2c). however, there was no significant difference between the size of a. japonicus sampled in the dwesa and cwebe sample sites (figure 2c and table 3). epinephelus andersoni showed similar size frequency distributions within the two sample sites (figure 2d and table 3). there was also no significant difference between the sizes of r. annulatus caught in the dwesa versus the cwebe sample sites (figure 2e and table 3). catch per unit effort sample site catch per unit effort the overall cpue for all sites sampled in the dwesa-cwebe mpa was 0.71 fish.angler-1.hour-1 + 0.48 sd; n = 191. the cpue in the dwesa sample site (0.84 fish.angler-1.hour-1 + 0.49 sd; n = 94) was significantly higher (p = < 0.0001) than that in the cwebe sample area (0.57 fish.angler-1.hour-1 + 0.43 sd; n = 89) (figure 3 and table 4). the highest cpue was achieved during the july 2009 sampling trip and the lowest during the february 2010 sampling trip (figure 3). the first mbashe estuary sampling trip produced a similar cpue (0.83 fish.angler-1. hour-1 + 0.43 sd; n = 8) to the dwesa sample site (figure 3 and table 4). figure 3: the catch per unit effort recorded at the three different sampling sites during seven field trips conducted between january 2009 and october 2010. table 4: the results of the comparison of catch per unit effort between the dwesa and cwebe sample sites for five dominant species sampled in the dwesa-cwebe marine protected area. age at 50% maturity and maximum age are also indicated. species-specific catch per unit effort the cpue for p. grande was similar during the different sampling trips, with a peak in the cwebe sampling site during july 2009 (figure 4a). the cpue for this species was significantly lower at the dwesa sample site compared to the cwebe sampling site (table 4). figure 4: the catch per unit effort for five dominant species (a) pachymetopon grande, (b) diplodus sargus capensis, (c) argyrosomus japonicus, (d) epinephelus andersoni and (e) rhinobatos annulatus, in the different sample sites in the dwesa-cwebe marine protected area during seven sampling trips conducted between january 2009 and october 2010. catches of d. s. capensis peaked in july 2009 at both sites and the cpue for d. s. capensis was significantly lower in the cwebe sample site compared to the dwesa sample site (figure 4b and table 4). the cpue of a. japonicus was highly variable (figure 4c). the cpue for this important species was significantly higher in the dwesa sample site than in the cwebe sample site (figure 4c and table 4). a high cpue (0.44 fish.angler-1.hour-1 ± 0.33 sd; n = 8) of small a. japonicus was recorded in the mbashe estuary during the first sampling trip conducted there in october 2010. catches of e. andersoni stayed relatively stable over the first six sampling trips (figure 4d). the cpue for e. andersoni was significantly lower in the cwebe sample site in comparison with the dwesa sample site (figure 4d and table 4). there was no significant difference in the cpue for r. annulatus between the dwesa and cwebe sample sites (figure 4e and table 4). ethical considerations top ↑ ecpta have adopted the standard saiab ethical procedures for handling aquatic organisms and research was conducted under permits issued through the department of environmental affairs and the department of agriculture, forestry and fisheries. discussion top ↑ cognisance must be taken of the preliminary nature of the results presented in this paper, which span a period of only two years and seven field trips. götz, cowley and winkler (2008) recommend a minimum of four years’ sampling. however, the pressure currently being placed on the authorities responsible for the management of the dwesa-cwebe mpa to open up parts of the reserve to subsistence and recreational shore-angling warranted analysis and publication of these results in order to help guide the decision-making process.the 28 species sampled during the current study period represent only 72% of the possible 39 inshore line-fish species likely to occur in the dwesa-cwebe mpa, based on the results of an earlier survey of the lower transkei coast (mann et al. 2003). other studies in the former transkei area also produced more species and families (table 5) (mann et al. 2003; pradervand 2004). these studies were based on roving-creel as well as shore-angling competition data and spanned a large proportion of the wild coast shoreline. in the case of the competition data (pradervand 2004), these were collected over a substantially longer time period (1977–2000), from the umtamvuna to the mbashe river. further afield, the port of ngqura (dicken 2010), tsitsikamma mpa (götz et al. 2008) and the goukamma mpa (dicken 2010; götz et al. 2008; pradervand & hiseman 2006) were also more species diverse than the dwesa-cwebe mpa (table 5). however, as only one sampling method was used during the current study (i.e. shore-angling), the number of fish species sampled was limited. a variety of sampling methods, which include line-fishing, underwater visual surveys, seine netting and possibly rotenone collections (for small cryptic species), over a long time period, are needed to enable compilation of a comprehensive fish species list for an mpa (wood et al. 2000). further sampling is thus needed in the dwesa-cwebe mpa to establish a more comprehensive fish species list. table 5: a comparison of fishery statistics between dwesa-cwebe marine protected area and a number of other areas in south africa. several overexploited and collapsed line-fish species were present in the dwesa-cwebe mpa as a whole. some were relatively abundant (e.g. a. japonicus, e. andersoni and p. grande), whilst other species were less abundant (e.g. dichistius capensis, pomatomus saltatrix, umbrina robinsoni, epinephelus marginatus, cymatoceps nasutus, rhabdosargus sarba, sparodon durbanensis and l. lithognathus). the red-listed species for which iucn assessments currently exist included only serranids and a few elasmobranch species. these included one endangered species (e. marginatus), two vulnerable species (carcharius taurus and haploblepharus fuscus) and two near-threatened species (e. andersoni and triakis megalopterus). the only other species with red-listed status was the endemic sparid l. lithognathus (skelton 1996). the assessment for l. lithognathus was done using older red-listing criteria and needs to be re-assessed (iucn 2011). the process of red-listing south african sparids and sciaenids is currently underway and the results should be available by the end of 2012 (b. mann, march 2012 ori, pers. obs.). a high proportion of the species sampled (53%) had some level of concern in terms of their stock status (or red-list status) and 43% were southern african endemics. this highlights the importance of the dwesa-cwebe mpa for the protection of threatened line-fish stocks. these results are not surprising considering that the mpa falls within an important transition zone between the agulhas and natal bio-regions (lombard et al. 2004; maree et al. 2000). the mpa can thus be considered to be of high conservation importance in terms of these criteria (bond 1989; wood et al. 2000).the three most abundant species caught in the dwesa-cwebe mpa were d. s. capensis, p. grande and a. japonicus. in other areas of the former transkei, outside proclaimed mpas, the most abundant species were p. saltatrix, d. s. capensis, r. annulatus, p. grande and sphyrna sp. (table 5) (mann et al. 2003; pradervand 2004). further south, in the goukamma mpa, the most abundant species were d. s. capensis, d. capensis and sarpa salpa (table 5) (pradervand & hiseman 2006), whilst in the tsitsikamma mpa, d. s. capensis, d. capensis and boopsoidea inornata were the most abundant species caught from the shore (götz et al. 2008). the main difference between the above areas and the dwesa-cwebe mpa was the abundance of a. japonicus and p. grande. only the port of ngqura, which is not a formally protected area, had similar abundances of a. japonicus (dicken 2010). inclusion of the mbashe estuary as a third sampling site in october 2010 revealed a high catch rate of juvenile a. japonicus suggesting that this estuary provides an important nursery area for this species. this was confirmed by current research being conducted in the mbashe estuary (n. james, saiab, april 2011 pers. comm.). this emphasises the importance of the dwesa-cwebe mpa for the protection of this heavily overexploited species. within the mpa, the dwesa sample site had a higher number of species than the cwebe sample site (table 2) and a significantly higher cpue, suggesting higher fish abundance (table 4). two major factors could be influencing this comparison. one is the difference in surf-zone habitat structure between the two sample sites and the other is the level of illegal exploitation. no surveys have been done to quantify the differences in habitat structure, but from on-site observation, both sites appear to be very similar in nature. within the surf-zone both sites have sandy substrata, broken rock and sand, rocky reefs and rocky points. with regard to illegal exploitation, there was clear evidence of higher levels of illegal exploitation of fish in the cwebe sample site. this is not surprising, considering its closer proximity to the hotel, holiday cottages and community areas. this evidence is further supported by the higher number of illegal activities reported by the mpa’s field rangers in the cwebe area (ecpta unpublished data). these preliminary results thus suggest that illegal fishing on the cwebe side of the mpa may be having an impact on the surf-zone line-fish community. in addition to this, several species-specific observations were made during the current survey. there was a higher relative abundance of d. s. capensis, e. andersoni and a. japonicus in the dwesa sample area compared to the cwebe sample area. all these species are relatively slow growing and reach a maximum age of > 10 years (cowley et al. 2001; fennessy 1998). species with these life history characteristics typically produce a lower yield per unit stock and have a slower rate of recovery following overexploitation (buxton & clarke 1989; cowley et al. 2001). no-take mpas are therefore frequently recommended as an important tool (in addition to conventional fisheries management regulations) to ensure effective protection and rebuilding of overexploited stocks of species of this nature (bennett & attwood 1991; attwood & bennett 1995; cowley et al. 2001; mann et al. 2006). no-take mpas help to allow exploited fish stocks to recover by placing a portion of the population off-limits to users. setting aside a no-take mpa is similar to limiting fishermen in terms of catch and/or effort, or any similar management strategy that effectively decreases fish mortality (byers & noonburg 2007; gell & roberts 2003; goni et al. 2010). illegal exploitation could potentially eliminate the positive effect of fish dispersal (known as spillover) from mpas (byers & noonburg 2007). byers and noonburg (2007) also predict that an initial large investment in enforcement efforts would provide the greatest return on maintaining the benefits of the mpa to the fishery of an area. the comparison of length frequencies between the two study areas was less revealing, with only p. grande showing evidence of significantly larger size classes present in the dwesa sample site compared to the cwebe sample site. surprisingly, very few l. lithognathus (0.47% of the total catch) were sampled during the study. this is cause for concern, as the beaches adjacent to the mbashe mouth were historically well-known for catches of this species, particularly during late winter (july–september) when adults formed large spawning aggregations (bennett 1993). although preliminary, the results of this study show that there are significant localised differences within the mpa that are probably caused by illegal exploitation. significant improvements in law enforcement and education are therefore needed in the dwesa-cwebe mpa, in order to decrease the current negative effects of illegal exploitation of fish. furthermore, it is also recommended that the current no-take status of the mpa should not be changed and that alternative options should be explored to improve socio-economic conditions within the local communities living adjacent to the dwesa-cwebe mpa. conclusion top ↑ this is the first baseline species list that has been developed for the dwesa-cwebe mpa. the 28 line-fish species recorded in this study include a significant number of nationally and even globally important species from a conservation perspective. important information regarding the biology of various key species (i.e. length frequency and relative abundance) was collected during this study, which could help inform future conservation management of the mpa. this study has also highlighted the regional importance of the dwesa-cwebe mpa by comparing it to other areas along the wild coast and further afield. however due to the preliminary nature of this study it is strongly advised that this monitoring project should be continued for a minimum of at least another three years to enable a more reliable evaluation to be made of the effectiveness of this mpa. acknowledgements top ↑ we would like to thank the volunteers, especially daniel fourie, keith funnel, gerhard barnard, harry steyn and schalk steyn, who give up their valuable time to assist with field sampling. paul cowley provided valuable assistance during the conception of the project. colin attwood, chantelle radue and dave balfour provided useful comments on earlier drafts of this manuscript. funding for the study was provided by the eastern cape parks and tourism agency, as well as the department of environmental affairs. competing interests the authors declare that they have no financial or personal relationship(s) which may have inappropriately influenced them in writing this paper. authors’ contributions j.a.v. (eastern cape parks and tourism agency) was the project leader, was responsible for experimental design, conducted all field work and performed all data analysis. b.q.m. (oceanographic research institute) was involved in the conceptualisation of the project, helped train field workers and gave substantial input in preparing the manuscript. references top ↑ attwood, c.g., 2003, ‘dynamics of the fishery for galjoen dischistius capensis, with assessment of monitoring methods’, african journal of marine science 25, 311–330. http://dx.doi.org/10.2989/18142320309504020 attwood, c.g. & bennett, b.a., 1995, ‘modelling the effect of marine reserves on the recreational shore-fishery of the south-western cape, south africa’, south african journal of marine science 16, 227–240. http://dx.doi.org/10.2989/025776195784156458 bennett, b.a., 1993, ‘aspects of the biology and life history of white steenbras lithognathus lithognathus in southern africa’, south african journal of marine science 13, 83–96. http://dx.doi.org/10.2989/025776193784287257 bennett, b.a. & attwood, c.g., 1991, ‘evidence for recovery of a surf-zone fish assemblage following the establishment of a marine reserve on the southern coast of south africa’, marine ecology progress series 75, 173–181. http://dx.doi.org/10.3354/meps075173 bond, w.j., 1989, ‘describing and conserving biotic diversity’, in b.j. huntley (ed.), biotic diversity in southern africa: concepts and conservation, pp. 2–18, oxford university press, cape town. buxton, c.d. & clarke, j.r., 1989, ‘the growth of cymatoceps nasutus (teleostei: sparidae), with comments on diet and reproduction’, south african journal of marine science 8, 57–65. http://dx.doi.org/10.2989/02577618909504551 byers, j.e. & noonburg, e.g., 2007, ‘poaching, enforcement, and the efficacy of marine reserves’, ecological applications 17, 1851–1856. http://dx.doi.org/10.1890/07-0067.1, pmid:17974326 cowley, p.d., brouwer, s.l. & tilney, r.l., 2001, ‘the role of the tsitsikamma national park in the management of four shore-angling fish along the south-eastern cape coast of south africa’, south african journal of marine science 24, 1–9. dicken, m.l., 2010, ‘the ichthyofauna in the port of ngqura, south africa’, african journal of marine science 32, 491–499. http://dx.doi.org/10.2989/1814232x.2010.538139 ecpta, 2010, unpublished field ranger records containing data on law enforcement activities, dwesa-cwebe nature reserve, willowvale. fennessy, s.t., 1998, biology and stock assessment of serranidae, oceanographic research institute, durban. fielding, p., 2010, dwesa-cwebe status report and scientific recommendations, department of enivironmental affairs: oceans and coast, cape town. fielding, p., robertson, w.d., dye, a.h., tomalin, b.j., van der elst, r.p., beckley, l.e. et al., 1994, transkei coastal fisheries resources, oceanographic research institute, durban. gell, f.r. & roberts, c.m., 2003, ‘benefits beyond boundaries: the fishery effects of marine reserves’, trends in ecology and evolution 18, 448–455. http://dx.doi.org/10.1016/s0169-5347%2803%2900189-7 goni, r., hilborn, r., diaz, d., mallol, s. & adlerstein, s., 2010, ‘net contribution of spillover from a marine reserve to fishery catches’, marine ecology progress series 400, 233–243. http://dx.doi.org/10.3354/meps08419 götz, a., cowley, p.d. & winkler, h., 2008, ‘selected fishery and population parameters of eight shore-angling species in the tsitsikamma national park no-take marine reserve’, african journal of marine science 30, 519–532. http://dx.doi.org/10.2989/ajms.2008.30.3.7.641 griffiths, m.h., attwood, c.g. & thomson, r., 1999, ‘new management protocol for the south african linefishery’, in b.q. mann (ed.), third southern african marine linefish symposium, pp. 145–156 , sancor, arniston. iucn, 2011, iucn red list of threatened species version 2011.2, retrieved april 2012, from http://www.iucnredlist.org/ kerwath, s.e., götz, a., attwood, c.g. & sauer, w.h.h., 2008, ‘the effect of marine protected areas on an exploited population of sex-changing temperate reef fish: an individual-based model’, african journal of marine science 30, 337–350. http://dx.doi.org/10.2989/ajms.2008.30.2.10.560 liebenberg, l., 2003, towards a worldwide environmental monitoring network, cybertracker, cape town. retrieved may 19, 2012, from http://www.cybertracker.org/index.php?option=com_content&view=article&id=140&itemid=125 lombard, a.t., strauss, t., harris, j., sink, k., attwood, c. & hutchings, l., 2004, ‘south african national spatial biodiversity assessment: marine component’, nsba technical reports, south african national biodiversity institute, pretoria. mann, b.q., 2000, south african linefish status reports, oceanographic research institute durban. mann, b.q., celliers, l., fennessy, s.t., bailey, s. & wood, a.d., 2006, ‘towards the declaration of a large marine protected area: a subtidal ichthyo-faunal survey of the pondoland coast in the eastern cape, south africa’, african journal of marine science 28, 535–551. http://dx.doi.org/10.2989/18142320609504204 mann, b.q., macdonald, a.m., sauer, w.h.h. & hecht, t., 2003, ‘evaluation of participation in and management of the transkei shore linefishery’, south african journal of marine science 25, 79–97. http://dx.doi.org/10.2989/18142320309504002 maree, r.c., booth, a.j. & whitfield, a.k., 2000, ‘effect of water temperature on the biogeography of south african estuarine fishes associated with the subtropical/warm temperate subtraction zone’, south african journal of science 96, 184–188. mcclanahan, t.r. & mangi, s., 2000, ‘spillover of exploitable fishes from a marine park and its effect on the adjacent fishery’, ecological applications 10, 1792–1805. http://dx.doi.org/10.1890/1051-0761%282000%29010%5b1792:soeffa%5d2.0.co;2 pradervand, p., 2004, ‘long-term trends in the shore fishery of the transkei coast, south africa’, african zoology 39, 247–261. pradervand, p. & hiseman, r., 2006, ‘an analysis of the recreational shore fishery in the goukamma marine protected area’, african zoology 41, 275–289. http://dx.doi.org/10.3377/1562-7020%282006%2941%5b275:aaotrs%5d2.0.co;2 skelton, p., 1996, lithognathus lithognathus, iucn redlist, retrieved 08 june 2011, from http://www.iucnredlist.org/ turpie, j.k., adams, j.b., joubert, a., harrison, t.d., collothy, b.m., maree, r.c. et al., 2002, ‘assessment of the conservation priority status of south african estuaries for use in management and water allocation’, water sa 28, 16. http://dx.doi.org/10.4314/wsa.v28i2.4885 wood, a.d., brouwer, s.l., cowley, p.d. & harrison, t.d., 2000, ‘an updated check list of the ichthyofaunal assemblage of the tsitsikamma national park, south africa’, koedoe 43, 83–95. abstract introduction methods results discussion acknowledgements references about the author(s) hindrik bouwman unit for environmental sciences and management, north-west university, potchefstroom campus, south africa enrico cronje unit for environmental sciences and management, north-west university, potchefstroom campus, south africa citation bouwman, h. & cronje, e., 2016, ‘an 11-digit identification system for individual nile crocodiles using natural markings’, koedoe 58(1), a1351. http://dx.doi.org/10.4102/koedoe.v58i1.1351 short communication an 11-digit identification system for individual nile crocodiles using natural markings hindrik bouwman, enrico cronje received: 28 aug. 2015; accepted: 15 jan. 2016; published: 20 may 2016 copyright: © 2016. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract research and conservation of wild crocodiles and husbandry of captive crocodiles requires the reliable identification of individuals. we present a method using the individual colour markings on the first 10 single-crest scutes on the tails of nile crocodiles (crocodylus niloticus). the scutes are scored by number for colour, with a prefix for left or right providing a binary 11-digit identification number (identification numbers [ids]; e.g. 12232232242 and 22333233232) per crocodile. a survey of 359 captive nile crocodiles showed no duplication. however, 42% had asymmetrical scute markings requiring a binary approach. there does not seem to be a change in patterns with age, except that the number of missing scutes increased. a small trial showed that this method can be applied in the field, although more work is needed to determine observer bias and establish parameters for observability in the field. it is unlikely that both left and right ids would be obtainable for each individual, but other distinctive markings such as scute shape and damage can be used to register the two ids to one individual. having two independent ids for each crocodile provides the possibility of two independent population estimates for equal effort without having to link left and right ids to individuals. our proposed method would be useful in conservation, individual tracking and husbandry. conservation implications: a non-invasive marking and recapture method for nile crocodile is presented whereby the first 10 single-crest scutes are scored for colour, allowing conservation practitioners to count and monitor crocodile populations and individuals. this method provides two equal-effort estimations of population size, as left and right hand sides are scored independently. introduction the reasons for accurate counts of crocodiles in any system can vary, but most likely mainly involves research and conservation. an impetus to count nile crocodiles (crocodylus niloticus) in the kruger national park (knp) occurred during the 2008/2009 mortality episodes in the olifants, letaba and sabie rivers. the numbers of animals that died remain uncertain but at least 170 carcasses were found (ferreira & pienaar 2011), with smaller numbers of carcasses in subsequent years. near the confluence of the olifants and letaba rivers and downstream into the olifants river gorge, crocodile numbers (using various counting techniques) declined from 780 in 2008 to 505 in 2009. elsewhere in southern africa, crocodile numbers have also declined in loskop dam (oberholster et al. 2010) and flag boshielo dam (dabrowski, oberholster & dabrowski 2014), ndumu game reserve (calverly & downs 2014), lake sibaya (combrink et al. 2011), and the panhandle of the okavango delta (bourquin & leslie 2011), or remained constant or increased slightly in the gonarezhou national park (zisadza-gandiwa et al. 2013) and some rivers in the knp (ferreira & pienaar 2011). there are a number of constraints with deriving crocodile population estimates. the fraction of the actual number of crocodiles that are available for sampling is the detectability bias. the detectability bias is dependent on many factors, including time of day (or night), season, vantage point (height, aerial platform, flat terrain, etc.), proportion of animals under water and not detected, and size and age of the crocodile, etc. (ferreira & pienaar 2011; calverley & downs 2014; zisadza-gandiwa et al. 2013 and their references). fixed-wing and helicopter counts consistently yielded lower estimates than night-time spotlight surveys (ferreira & pienaar 2011; combrink et al. 2011). however, correction factors or estimates can be applied to any counting technique to derive population estimates (ferreira & pienaar 2011; combrink et al. 2011; calverley & downs 2014; bayliss et al. 1986). estimating populations can also be done using mark-recapture (e.g. bayliss et al. 1986; nair et al. 2012). the constraints here are dependent largely on equal catchability of the crocodiles in the system and marking a large enough proportion of the crocodiles. this did not prove feasible in the olifants river gorge area in the knp, using plastic tags attached to tail scutes and/or scute clipping (ferreira & pienaar 2011). however, a similar method was used successfully in the panhandle of the okavango delta, where 224 of 1717 marked individuals were recaptured over 4 years (bourquin & leslie 2011). however, capturing and marking individuals is time-consuming, may alter the behaviour of the individuals (bourquin & leslie 2011; underhill & fraser 1989), there could be marker loss, and there may be catchability and re-catchability bias in size or age classes and sex (bayliss et al. 1986), amongst a number of factors. several of these factors can be reduced when using natural markings on the animal, not least of which are avoiding the direct and lingering effects of capture and handling, and the relative permanency of the marks. crocodile surveys using natural mark schemes have been used successfully (swanepoel 1996; nair et al. 2012). both used variations of the method developed by singh & bustard (1976). these methods use unique marking (dark blotches) on the sides of the tail and the single crest of scutes on the tail. however, none of these have tested the possibility of duplication between animals, comparisons between left and right aspects and the scoring system employed does not easily lend itself to computer comparisons. we describe and test a binary 11-digit, identification system for individuals based on the colour and presence or absence of the first 10 proximal scutes of the single crest. methods from proximal to distal on the dorsal posterior of the nile crocodile, there is a double-row or crest of scutes that changes to a single row where the tail narrows. each of these scutes has a colour pattern or is absent because of injury. we visited a crocodile farm to compile a photographic database of as many crocodiles as possible. using a high-definition camera, photographs were taken of the tail sections of the 10 proximal single-crest scutes of nile crocodiles of various age groups, ranging from a few months to over 20 years of age (figures 1a and 1b). each of the single-crest scutes has a colour pattern consisting of white or light to dark or black, or absent. the first 10 single-crest scutes were scored proximal to distal. figure 1: scute colouration as used for deriving identification numbers. (a and b) left and right views of the same juvenile crocodile from castlekop crocodile farm. respective ids are 12232232242 and 22333233232. (c and d) views of two different crocodiles photographed in botswana, with respective ids of 13233233223 and 23222322332. (e and f) ‘tourist’ photographs, taken in south africa (24212323433 with 18× optical zoom) and botswana (11221022142 with 20× optical zoom). initially, we assumed that the scute colouration would be symmetrical, that is, the left and right sides of the scutes would be identical, therefore enabling the identification number (id) to be the same regardless of which side the crocodile is viewed from. however, this was not the case (see results). this means that to accurately identify a crocodile, the researcher must be able to view both sides of the tail. if only the right side of a crocodile’s tail has been recorded in the past and the left side of that same crocodile’s tail is now recorded without being able to see the right side, the database might register it as a new crocodile whilst the crocodile has already been recorded, leading to inaccurate population estimates. the scoring system accommodates this by allocating the first digit to orientation. it is therefore possible over time to complete a database for a certain area with both the left and right sides of all the crocodiles recorded, as certain crocodiles are scored again when both sides of the tail are visible. the first digit of the 11-digit id donates the aspect of the tail that is scored; left (1), right (2), or uncertain or unknown (3). scoring of the 10 scutes is done by assigning a number to each scute based on the particular colouration: first digit – left = 1, right = 2, unknown = 3. a missing or damaged scute = 0. a white or light scute = 1. a scute less than 50% dark or black = 2. a scute more than 50% dark or black = 3. a completely dark or black scute = 4. a scute that cannot be scored = 5. each crocodile therefore, has two, linked, 11-digit id numbers with 9 765 625 possible unique id numbers per side, excluding the missing scute category. we assigned ids to 359 nile crocodiles of the castlekop crocodile farm near barberton, mpumalanga. the ids were entered into a microsoft excel 2007 database and scanned for duplicates. as mentioned above, we also tested for asymmetry. except for frequencies (table 1; figure 2), further statistics including reporting of means is not possible as the data per colour score are categorical and not normally distributed because of many zeros in some colour categories. figure 2: cumulative distribution of scute colouration per crocodile age group. (a) cumulative distribution of light/white-coloured scutes per crocodile age group. (b) cumulative distribution of scutes that are less than 50% dark/black per age group. (c) cumulative distribution of scutes that are more than 50% dark/black per age group. (d) cumulative distribution of scutes that are completely dark/black per age group. (e) cumulative distribution of missing scutes per age group. table 1: data of scores per age group from the castlekop crocodile farm. we trialled this method in the field to see if nile crocodile scutes can be scored with binoculars and super-zoom cameras. we also scanned our photograph collections for scorable photos to see if opportunistic ‘tourist’ photographs may be useful. results at castlekop, we scored 49 nile crocodile individuals in the 0to 1-year age group, 140 in the 1–7-year age group, 159 in the 7–20-year age group and 11 in the group older than 20 years. these categories were dictated by the housing of the crocodiles in age-pens. figures 1a and 1b show the left and right views of the same juvenile crocodile (0–1 year) with respective ids of 12232232242 and 22333233232, which is asymmetrical. of 56 crocodiles that were photographed from both sides, 62% of the left or right scute colouration patterns were asymmetrical. figures 1c and 1d scored 13233233223 and 23222322332 for nile crocodiles photographed in the wild in botswana. both have missing tails tips, but both have 10 scorable scutes. the scoring of the last scute of figure 1c was difficult as it was in the shade, but could be done by zooming in. figure 1e, a ‘tourist’ photo from the knp (18× optical zoom), scored as 24212323433 after zooming in. figure 1f (a tourist photo from botswana with 20× zoom) scored 11221022142. zooming in allowed the finding of the first single-crest scute that is not immediately apparent from the print. the sixth scute is partially missing (0). this characteristic could be used in registering the right id with the left when this crocodile is sighted later from the other side. most of the nile crocodile scutes scored in the < 50% or > 50% categories (table 1), followed by light or white, dark or black and missing scutes. no consistent pattern of change in colour categories with age is apparent. with most of the scutes in the > 50% and < 50% categories, small differences in the light or white and dark or black categories will result in relatively large variable percentages between age groups. the consistent maximum number of scutes in any crocodile (third number in the columns) remained remarkably constant with age, except for missing scutes. here, as can be expected, the percentages increased with age. the number of crocodiles with at least one missing scute increased from 22.4% in the youngest group to 72.7% in the oldest. the cumulative distribution of the number of scutes per crocodile per colour score showed no crocodiles with all 10 scutes exclusively in one colour category (figure 2). no crocodile had more than five dark or black scutes and none more than seven light or white scutes. especially for the light or white (figure 2a) and dark or black (figure 2d) categories, up to 45% of the animals had none of either. none or almost none had no scutes with < 50% dark colouration (figure 2b). missing scutes (scored as 0) is likely to increase with age because of handling injury and fighting. this is apparent in table 1 and in figure 2. a small study on the banks of the crocodile river (knp) showed that 48 nile crocodiles were repeatedly identified everyday over a period of a week. discussion the chance of two nile crocodiles in a population of 500 having the same id is less than 0.0001%, assuming that the colouration patterns are expressed randomly. because it can be assumed that a reasonable proportion of the crocodiles that were photographed at the crocodile farm must be related, our findings show that there were no duplicates in id numbers in a relatively isolated interbreeding group. in 48% of cases, the patterns were symmetrical, meaning that it cannot be presumed that a left and right id would coincide. because it is unlikely that both sides of the tail would be observable in the field, left and right ids cannot be directly linked. there may be cases though, where scute wear patterns and/or missing scutes would be visible from both sides, as in figure 2f, where the sixth scute is partially missing. visual comparisons of photographs would eventually register left and right ids with a reasonable measure of confidence. having two id numbers per crocodile is not ideal. however, for population studies, separate left and right id numbers (even if linkable to registered individuals) would provide two independent estimations, probably providing more confidence in upper and lower population limits. table 1 and figure 2 imply that, with the exception of missing scutes, colouration patterns and therefore id numbers would remain relatively stable with age. this observation would need substantiation, probably best done in a captive population of nile crocodiles, where a substantial number of individuals can be followed over a long period. for short-term studies though, such as individual movement tracking and population estimates, stability of the ids would most likely be sufficient. id numbers that would change with age because of damage or missing scutes would not present an insurmountable problem, as identification through partial ids in many circumstances may still be possible. in many cases, not all 10 scutes will be visible. often, the last portion of the tail would still be in the water, curved away from the observer, or covered in mud. here, identification using a partial id score would in some cases still be possible if the scorable scutes, coupled with other distinctive markings such as scute shapes, have a unique combination. although this method was trailed successfully on a small scale on nile crocodiles, the utility needs to be shown by larger and longer term studies. it will also be necessary to investigate observer bias in scoring crocodile scutes and to determine the parameters whereby identifications can be made with confidence, such as distance between the observer and crocodiles, approachability in the wild, light conditions and magnification needed for cameras and telescopes. however, sighting and re-sighting issues remain, likely biased against smaller individuals. other limitations would be scutes covered in mud or algae. assigning and recognising ids could also be trailed using hd photographs taken from aerial platforms, or even using infrared optics at night. ‘tourist’ photos might also prove useful in tracking individuals over time, especially if the cameras have geolocation built in. this method may also be applicable to other crocodilian species where the tails have distinctive and stable colour patterns. this method may therefore prove useful in conservation (e.g. population estimates) and research (e.g. tracking of movements) that may also involve citizen participation, or keeping track of the ages, sex and conditions of individuals in the wild and captivity. acknowledgements we thank danie pienaar and xander combrink, and suria ellis for help with some of the statistics. we also thank the castlekop crocodile farm near barberton, south africa, in particular charles ferreira. funding was provided by the national research foundation of south africa. opinions expressed and conclusions arrived at are those of the authors and are not necessarily to be attributed to the national research foundation (nrf). competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions e.c. and h.b. conceived and planned the project. e.c. performed all the fieldwork. e.c. and h.b. analysed the data and prepared the manuscript. references bayliss, p., webb, g.j.w., whitehead, p.j., dempsey, k. & smith, a., 1986, ‘estimation of the abundance of saltwater crocodiles, crocodylus porosus schneider, in tidal wetlands of the northern territory: a mark-recapture experiment to correct spotlight counts to absolute numbers, and the calibration of helicopter and spotlight counts’, australian wildlife research 13, 309–320. bourquin, s.l. & leslie, a.j., 2011, ‘estimating demographics of the nile crocodile (crocodylus niloticus laurenti) in the panhandle region of the okavango delta, botswana’, african journal of ecology 50, 1–8. calverley, p.m. & downs, c.t., 2014, ‘population status of nile crocodiles in ndumu game reserve, kwazulu-natal, south africa (1971–2012)’, herpetologia 70, 417–425. combrink, x., korrübel, j.j., kyle, r., taylor, r. & ross, p., 2011, ‘evidence of a declining nile crocodile (crocodylus niloticus) population at lake sibaya, south africa’, south african journal of wildlife research 41, 145–157. dabrowski, j., oberholster, p.j. & dabrowski, j.m., 2014, ‘water quality of flag boshielo dam, olifants river, south africa: historical trends and the impact of drought’, water sa 40, 345–357. ferreira, s.m. & pienaar, d., 2011, ‘degradation of the crocodile population in the olifants river gorge of kruger national park’, aquatic conservation: marine and freshwater ecosystems 21, 155–164. nair, t., thorbjarnarson, j.b., aust, p. & krishnaswamy, j., 2012, ‘rigorous gharial population estimation in the chambal: implications for conservation and management of a globally threatened crocodilian’, journal of applied ecology 49, 1046–1054. oberholster, p.j., myburgh, j.g., ashton, p.j. & botha, a.-m., 2010, ‘responses of phytoplankton upon exposure to a mixture of acid mine drainage and high levels of nutrient pollution in lake loskop, south africa’, ecotoxicology and environmental safety 73, 326–335. singh, l.a.k. & bustard, h.r., 1976, ‘a method to identify individual young gharial (gavialis gangeticus)’, british journal of herpetology 5, 669–671. swanepoel, d.g.j., 1996, ‘identification of the nile crocodile crocodylus niloticus by the use of natural tail marks’, koedoe 39, 113. underhill, l.g. & fraser, m.w., 1989, ‘bayesian estimate of the number of sunbirds feeding at an isolated and transient nectar resource’, journal of field ornithology 60, 382–387. zisadza-gandiwa, p., gandiwa, e., jakarasi, j., van der westhuizen, h. & muvengwi, j., 2013, ‘abundance, distribution and population trends of nile crocodile (crocodylus niloticus) in gonarezhou national park, zimbabwe’, water sa 39, 165–169. filelist convert a pdf file! supplement to koedoe. 1977: 243-247. nature conservation as a resource for tourism t cowen director south african tourist corporation private bag x164 pretoria 0001 the spectacular development in air technology during and since the second world war, and a parallel economic growth, have been responsible for a tremendous increase in international tourist statistics, making tourism the world's largest industry. for the sake of clear international definition, a tourist is regarded as a person who visits a country other than his country of normal residence, for any reason other than being gainfully employed within the country he visits. during 1975 such tourist arrivals numbered 213 million, and the receipts from international tourism amounted to us $32000 million. domestic tourism also plays an important role in most countries . it is the rule rather than the exception that the local tourist creates the demand for the development of amenities, which can then cater for the tourist from abroad. tourism today is highly organised. the private sector provides hotels and transport. governments, however, have a major responsibility in financing communication links, including air and surface transport, and the building of roads. governments also play a leading part in the organisation and planning of tourism, which could and should include setting aside areas for nature conservation. the south african tourist corporation (satour), responsible for the promotion of tourism to the republic of south africa (rsa) from abroad for the purpose of earning foreign exchange, is a statutory body established in 1947. in its internatlonal marketing operation sato~h is supported and assisted by the south african airways and its pool partners, other air and shipping lines; travel wholesalers, tour operators, approximately 20000 travel agents overseas, motoring clubs and south african missions abroad. satour relies on the co-operation of the department of tourism and its national tourist bureaux, other state departments, the south african railways, provincial and local authorities, the hotel board, publicity associations, hoteliers, ground operators and -last but certainly not least the national parks board of trustees of the republic of south africa. 243 the rsa as a tourism product has many components. to list just a few of the main components, i would immediately think of our country's scenery, climate, mineral wealth and its national parks as they are so well described in satour's tourist literature: geographically, we have been endowed in this small tip of africa with many climates. the northernmost reaches of the republic lie just within the tropic of capricorn; in the south, the city of cape town faces out on to the atlantic ocean, which sweeps to meet the indian ocean at cape agulhas, in a wild union of force and beauty. if one could orbit the earth in a space ship and look down on southern africa, it would seem that at some moment in the heat of creation the entire continent tipped forward and the hot wrinkled skin of the land slid down towards the sea, but before it could disappear hissing and bubbling under the waves, the continent bobbed up, righting itself, and there at the edge the great mass remained solidified into towering escarpments, fold upon fold, forming an enormous mountain barrier between the interior and the two oceans. on the north-west coast the mountains are scoured by the spuming fury of the atlantic which carries in its restless fists, the cold benguela current washing up from the antarctic wastes . it brings no gift of rain to that infamous stretch ofland between the mountains and the sea known as the skeleton coast. beyond the windswept barrier, stretching inland to the north and east, are the arid lands of namaqualand and the karoo dry land that merges with the kalahari desert, huge, silent and curiously beautiful. prevailing winds bring about a fine mediterranean climate at the south west cape, and from there around the coast the scene mellows. mountains are green-clad with natural forests. inland, protected by the mountains, the land rises to great plateaus. the highveld soars to 2000 m, where winter nights are frosty and the days bright with sun. to the east, by way of a great escarpment, the land plunges down to the warm lowveld area, thick with bush, and habitat of the greatest variety of african animals. i would point out that there is probably not a region in the world where the variety of mineral wealth is so vast. the rsa has a soil so varied in mineral content and a terrain so varied in physical characteristics and climate, that it has given rise to an overwhelming variety of plants. of the world's 300 or so families of flowering plants, 190 are represented in the rsa 18 000 species in all. the plants nurture a wide variety of herbivores grazing for the hippo, white rhino, zebra and buffalo, treetop browsing for the giraffe, leaves for the kudu, shrubs for the eland, bark for the porcupine, berries for monkeys, baboons and birds, branches for the elephant, and thorn trees for the black rhino. these in turn provide food for a spectacular variety of carnivores jackals of many kinds, foxes, hyeanas, seven different felines, including the lion and the spotted cats. in recent years, as it becomes obvious that people are no longer 244 content to travel the world viewing cities and the works of man, the rsa has endeavoured to find a happy compromise between continuing the development of one of the world's materially very wealthy regions, and fulfilling her obligations as custodian to a uniqu e concentration of wildlife by the es tablishment and subsequent developm ent of th e national park board of trustees. to be really attractive to the international tourist, a country does not only need amenities and facilities such as sophisticated hotels and transport, but must have features which are unique unto itself. europe, where mediaeval architecture and other antiquities abound, is at present our main source of tourism. the european tourist who is surrounded and feels stifled perhaps by people and buildings, yearns for wide open spaces , a sunny climate, a smogless environment, and, most of all, responds to the "return to nature" call. a strong motivation for travel would be the need for man's temporary flight from artificiality towards unsullied nature . to find peace of mind, how could he spend his savings and leisure time better than in a nature conservation area? the rsa has such a variety of tourist attractions that satour in its marketing concentrates on the all-incorporating advertising slogan "a world tour in one country". of all the attractions, the nature and game reserves surpass them all. a recent survey conducted by satour in 10 countries established that 23% of the total sample chose to visit the rsa's game reserves; the only higher percentage being that of overseas visitors wanting to visit friends and relatives. they would, once here, also wish to have the opportunity of experiencing our national parks. the national parks board of trustees, through its avowed policy of conservation, created a tourist product which satour, as a tourist marketing organisation, exploits to its fullest advantage. in 1657, on the 26th october,jan van riebeeck as commander of the cape of good hope, made an entry in his journal. part of it reads: " the commander today went to inspect a forest ... the free carpenter, leendert cornelissen of zevenhuisjen, wished to have the sole right to obtain timber from it, in such a way that the forest would suffer no damage, but would be improved ... ". these words inspired one of satour's highly successful advertising campaigns, embodied in the slogan " conservation is a 300-year-oldword in the rsa". satour's marketing overseas has been notably successful. the total visitor arrival figure for 1975 was 730368, an increase of 20% on the figure for 1974. tourists from our neighbouring territories comprise 45% of total arrivals, namely 327 380, which represents an increase of 12% on that of 1974. most countries with large tourist intakes rely heavily on over border marketing areas for their main supply of tourists. the rsa does not have that potential nearby, but has to look towards overseas countries such as the united kingdom, united states of america (usa) and germany for tourists . in view of this, it is encouraging to note that 10 245 years ago a meagre 27% of our visitor arrivals came from overseas, whereas in 1975 the numbers of overseas visitors increased to 55% of all tourist arrivals. main sources of tourism from overseas were: the united kingdom 140324, (this is an increase of 34% on the 1974 figure) . the usa with 48712 was on 14%, germany 41506 (+26%), australia 20654 (+ 15%) and holland 18 238 (+ 27%). the average length of stay for these tourists was 19 days, and the estimated earnings of r274 million in foreign exchange during 1975 was an all time record. although it will require all our marketing skills this year to improve on the record 1975 figures due in part to a slight and, we believe, temporary hesitancy about travelling to the rsa after the unrest in black townships received such wide coverage abroad the overall picture is good. satour has set itself the marketing target of a million visitors by 1980 and is confident this target will be reached a year or two ahead of schedule. it has, however, been pointed out that tourism, with the multitudes that it brings with it, harbours the seeds of destruction. visitors in ever growing numbers flock to the national parks in the usa some 50 million in 1955, more than 110 million in 1965 and according to the latest estimate, more than 200 million in 1976. duringjuly 1976, visitors to the yellowstone national park were caught in traffic jams. it is reported that similar scenes are being played out every day this season in america's 285 national parks, recreation areas and historic sites. by the end of may 1976, for example, yellowstone had 70% more visitors than in the first five months oflast year. by the end of june 1976, the nearby grand teton national park reported an increase of 38 %. yosemite national park in california registered a 63% increase through may 1976. the national parks and conservation association has published a stark report detailing manpower deficiencies and deteriorating conditions throughout th e system. although we are many years away from these staggering numbers and acute problems, it is emphatically not too early to support the chief director of national parks in the rsa in his call for the conservation of many natural wonders, ecosystems and plant and animal species that still exist without protection, on a national level. new conservation areas will not only serve the cause of wild life protection but will spread the flow of tourism in the future. dr knobel said in 1972 that our generation is the last one to have the opportunity of setting aside space for conservation; no other country on this earth has the existing possibilities and versatility as the rsa has for the establishment of nature areas, and thus enhancing the diversity of our tourist product. the demonstrable fact that it is possible to use agricultural land more gainfully for touristic purposes should not be forgotten . it was estimated by the department of agricultural economics and marketing that the annual agricultural income of the addo elephant national park would have been r15 000 while its tourist income was 246 r45000. similarly the agricultural income of the mountain zebra national park and the augrabi es falls national park would have been ril 000 and r2 500 respectively whereas from tourism it was actually rl8 000 and r28 200. in the kruger national park the annual agricultural income was estimated at approximately r3 million compared to an amount of nearly r6 million derived from tourism. it is apparent therefore that tourism does have its blessings, even in the field of nature conservation; moreover the valuable influx of foreign exchange draws the government's attention to the value of nature cons ervation , not merely for nature' s sake alone, but also for the sake of the country's economics as well as its foreign image. one cannot deny the interdependence of nature cons ervation and tourism. to carry out nature conservation money is needed, and this tourism can supply. the same tourists who need and seek wild life, are the ones who provide the necessary funds for its upkeep and expansion . a zulu who had completed a course at th e wilderness leadership school aptly summed up this interdependence with the words: "we were very impressed. w e were told that the game r eserves the zulu is about to inherit don't belong to us, they belong to the world. but if th ey belong to the world, then let the world pay for their upkeep , because we can ' t afford to". these wise words bring two things to our attention: that our heritage of abundant nature is to be shared with the rest of the world, and also, that the r est of the world, in the form of tourism, is th e medium by which this can b e financed . let us, in this age in which so many values seem to be in a precarious balance, appreciate the work done by our nature conservationists for, as carl linnaeus wrote 200 years ago: "there is no place on earth with such a variety of rare plants, animals, insects and other marvels of nature as africa". even with our open spaces, sunny climate and different cultures, we find ourselves on a very competitive basis with other countries of the world. however, in th e sphere of wildlife parks, the rsa is rated among the best in the world. other competing countries, which also have similar potentials, seem to be losing out on their tourist business because of malpractices contrary to nature conservation in their parks, and because they do not have the conservation-orientated policy which our national parks board of trustees so strictly applies. we must rememb er that all natural resources can be exhausted, but nature, which has been conserved, has no limits; it can only grow and grow into abundance. the more the world expands industrially, the more our nature conserving projects will come to be valued and sought after so that it is certain that the next 50 vears of nature conservation in the rsa will be more prosperous than the 50 fruitful years that have passed. 247 page 1 page 2 page 3 page 4 page 5 article information authors: mariette marais1 antoinette swart1 affiliations: 1agricultural research council, plant protection research institute, south africa correspondence to: mariette marais postal address: private bag x134, queenswood 0121, south africa dates: received: 11 dec. 2013 accepted: 27 aug. 2014 published: 17 nov. 2014 how to cite this article: marais, m. & swart, a., 2014, ‘plant nematodes in south africa. 12. checklist of plant nematodes of the protected areas of the eastern cape province’, koedoe 56(1), art. #1220, 3 pages. http://dx.doi.org/10.4102/ koedoe.v56i1.1220 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/ koedoe.v56i1.1220-1. copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. plant nematodes in south africa. 12. checklist of plant nematodes of the protected areas of the eastern cape province in this checklist... open access • abstract • introduction • materials and methods • results and discussion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ soil-inhabiting nematodes, including plant-parasitic nematodes, are considered to be the most abundant multicellular organisms in the soil, and of particular interest since they are an integral part of the interlocking chain of nutrient conversions. because of their abundance and relative susceptibility to both physical and chemical changes, these organisms are used as indicator organisms. the national collection of nematodes (ncn) consists of a core collection, the meloidogyne collection and the juan heyns collection, which are housed at the plant protection research institute of the agricultural research council in pretoria. vast amounts of biodiversity data are contained in ncn, and the digitising of the collection from 2007 to 2014 yielded unpublished locality information, especially datasets of plant nematodes reported from protected areas of the eastern cape. two hundred and thirty plant nematode species belonging to 36 genera were reported from the eastern cape. of these, only 80 were from protected areas, whilst 163 were from uncultivated areas (outside protected areas) and 148 from cultivated areas. ten species were described from protected areas, namely criconemoides silvicola, meloinema silvicola, ogma tuberculatum, paralongidorus cebensis, paralongidorus hanliae, scutellonema tsitsikamense, trichodorus vandenbergae, xiphinema erriae, xiphinema ornatizulu and xiphinema simplex. only m. silvicola, o. tuberculatum, p. cebensis and s. tsitsikamense were not reported from other provinces, suggesting endemism.conservation implications: the diversity of nematode fauna is not adequately protected as most nematode biodiversity in the eastern cape lies outside protected areas, with only 80 of the 230 plant-feeding nematode species in the province being reported from protected areas. introduction top ↑ the national collection of nematodes (ncn), one of the four national collections housed in the biosystematics programme of the plant protection research institute, agricultural research council, pretoria, was founded in the mid-1950s, with the first type specimens deposited in 1958. the national collection of nematodes consists of a core collection, the meloidogyne collection and the juan heyns collection. the national collection of nematodes currently consists of 180 000 specimens, of which 7140 are type specimens, mostly from southern africa, but also from such diverse localities as antarctica and the amazon.the type specimens of 510 species are currently deposited in ncn and we now know that 455 plant nematode or plant-feeding nematode species are reported from south africa. the south african plant-parasitic nematode survey (sappns) programme was initiated in 1987 to produce a comprehensive assessment of the nematode biodiversity resources of south africa. one of the four objectives of sappns is to compile an inventory of the plant nematodes of south africa (marais 2006). as part of this initiative, surveys were undertaken in various areas, including some of the protected areas of the eastern cape, in order to establish the incidence and distribution of plant nematodes in these areas (van den berg 1996). in this article, plant nematodes are seen as nematodes that feed on plants and therefore always have a tylenchoid stomatostyle, dorylaimoid odontostyle or trichodorid onchiostyle. this includes nematodes that have an ectoparasitic, endoparasitic or semi-endoparasitic life cycle (yeates et al. 1993). as recommended in the international code of zoological nomenclature, locality data and host plant data are usually published as part of species descriptions. apart from a checklist of the tsitsikamma national park (van den berg 1996), the nematode biodiversity information of the eastern cape had not been consolidated into a single publication. during the digitising of the specimens of ncn, a number of unpublished records of plant nematodes reported from protected areas in the eastern cape were discovered. materials and methods top ↑ the datasets reflected in the checklist consist of nematodes collected by various nematologists over a period of 30 years. in some cases the collection and extraction methods are not known. the standard procedure of ncn is that soil samples are taken with a garden trowel, spade or soil auger, depending on the terrain in the rhizosphere of the plant. the plants are collected at the same time and the soil and accompanying plant are sealed in plastic bags, placed in a cool box containing ice bricks and transported to the laboratory of ncn where the samples are stored in a cold room at 12 °c. the relevant collection data, including locality, habitat, substrate, moisture regime, soil type, exposure slope, aspect and host plant, are noted in the field.the nematodes are extracted from the soil using the sugar centrifugal-flotation method (jenkins 1964). specimens are killed in water by the gradual application of heat, then fixed and preserved in distilled water, 4% formaldehyde and 1% propionic acid (known as fpg), dehydrated to anhydrous glycerine and permanently mounted in anhydrous glycerine on cobb aluminium slides (hooper & evans 1993; netscher & seinhorst 1969). nematodes are extracted from plant material using the method described by kleynhans (1997), and are killed, preserved and mounted in the same way as the nematodes extracted from soil. the classification of south african plant nematodes followed here is a synthesis of the classification by maggenti et al. (1988) for tylenchina; and decraemer (1995) and duarte et al. (2010) for trichodoridae. authorities for genera regarded as valid here are hunt, luc and manzanilla-lópez (2005) for criconemoides; marais (2001) for helicotylenchus; escuer and arias (1997) for paralongidorus; brzeski (1998) for paratylenchus; and handoo (2000) for tylenchorhynchus. results and discussion top ↑ the checklist reflects specimens collected between 1968 and 1998 from seven protected areas in the eastern cape. the sappns database contains 311 records of nematodes sampled in uncultivated areas in the eastern cape, of which 98 records are from localities in protected areas (online appendix 1). the database also contains 505 records of localities sampled in cultivated areas, which include crop fields, plantations, gardens and sports fields. it was found that 80 plant nematode species were reported from protected areas (table 1), 163 species from uncultivated areas (outside protected areas) and 150 species from cultivated areas.in total, 230 (51%) of the 455 currently known plant nematode species from south africa have been recorded from the eastern cape. of these, 29 species were described from the province; 10 of these were described from the protected areas in the province: criconemoides silvicola van den berg 1996, meloinema silvicola kleynhans 1988, ogma tuberculatum van den berg 1996, paralongidorus cebensis (heyns & coomans 1989) escuer & arias 1997, paralongidorus hanliae liebenberg, heyns & swart 1993, scutellonema tsitsikamense van den berg 1976, trichodorus vandenbergae de waele & kilian 1992, xiphinema erriae hutsebaut, heyns & coomans 1988, xiphinema ornatizulu hutsebaut, heyns & coomans 1989 and xiphinema simplex hutsebaut, heyns & coomans 1989. only m. silvicola, o. tuberculatum, p. cebensis and s. tsitsikamense were not reported from other provinces, suggesting endemism. acknowledgements top ↑ we thank various colleagues and clients for collecting the samples, the taxonomists at arc-ppri and (the former) rand afrikaans university for species identification and the south african biodiversity information facility for financial assistance in order to digitise a major portion of ncn. competing interests the authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article. authors’ contributions m.m. (plant protection research institute) is the coordinator of sappns, project leader of digitising the national collection and taxonomist responsible for helicotylenchus, belonolaimidae, dolichoridae, trichodoridae and the subfamily meloidogyninae, and was responsible for writing the draft concept of the manuscript. a.s. (plant protection research institute) is the taxonomist responsible for longidoridae, anguinidae, aphelenchoididae and the subfamily heteroderinae, and was responsible for the final review of the manuscript. m.m. and a.s. were both responsible for writing this article. references top ↑ brzeski, m., 1998, nematodes of tylenchina in poland and temperate europe, museum i instytut zoologii polska akademia nauk, warzawa.decraemer, w., 1995, the family trichodoridae: stubby root and virus vector nematodes, kluwer academic press, dordrecht. http://dx.doi.org/10.1007/978-94-015-8482. duarte, i.m., de almeida, m.t., brown, d.j.f., marques, i., nielsen, r. & decraemer, w., 2010, ‘phylogenetic relationships, based on ssu rdna sequences, among the didelphic genera of the family trichodoridae from portugal’, nematology 12(2), 171–180. http://dx.doi.org/10.1163/156854109x461721 escuer, m. & arias, m., 1997, ‘paralongidorus iberiss p.n. and p. monegrensiss p.n. from spain with a polytomous key to the species of the genus paralongidorus siddiqi, hooper & khan, 1963 (nematoda: longidoridae)’, fundamental and applied nematology 20(2), 135–148 viewed n.d., from http://horizon.documentation.ird.fr/exl-doc/pleins_textes/fan/010009433.pdf handoo, z., 2000, ‘a key and diagnostic compendium to the species of the genus tylenchorhynchus cobb, 1913 (nematoda: belonolaimidae)’, journal of nematology 32(1), 20–34. pmid:19270946. hooper, d.j. & evans, k., 1993, ‘extraction, identification and control of plant nematodes’, in k. evans, d.l. trudgill & j.m. webster (eds.), plant parasitic nematodes in temperate agriculture, pp. 31–59, cab international, wallingford. hunt, d.j., luc, m. & manzanilla-lópez, r.h., 2005, ‘identification, morphology and biology of plant parasitic nematodes’, in m. luc, r.a. sikora & j. bridge (eds.), plant parasitic nematodes in subtropical and tropical agriculture, 2nd edn., pp. 11–52, cabi publishing, wallingford. http://dx.doi.org/10.1079/9780851997278.0011 jenkins, w.r., 1964, ‘a rapid centrifugal-flotation technique for separating nematodes from soil’, plant disease reporter 48, 692. kleynhans, k.p.n., 1997, collecting and preserving nematodes. manual for a safrinet course in practical nematology, safrinet, the southern african (sadc) loop of bionet-international, pretoria. maggenti, a.r., luc, m., raski, d.j., fortuner, r. & geraert, e., 1988, ‘a reappraisal of tylenchina (nemata). 11. list of generic and supra-generic taxa, with their junior synonyms’, revue de nématologie 11, 177–188. marais, m., 2001, ‘a monograph of the genus helicotylenchus steiner, 1945 (nemata: hoplolaimidae)’, phd dissertation, department of entomology and nematology, university of stellenbosch. marais, m., 2006, ‘south african plant-parasitic nematode survey (sappns)’, plant protection news 67, 6. netscher, c. & seinhorst, j.w., 1969, ‘propionic acid better than acetic acid for killing nematodes’, nematologica 15, 286. van den berg, e., 1996, ‘a first list of plant-parasitic nematodes from the tsitsikamma national park, with descriptions of two new species of the subfamily criconematinae’, koedoe 39(1), 43–54. http://dx.doi.org/10.4102/koedoe.v39i1.281 yeates, g.w., bongers, t., de goede, r.g.m., freckman, d.w. & georgieva, s.s., 1993, ‘feeding habits in nematode families and genera – an outline for soil ecologists’, journal of nematology 25, 315–331. filelist convert a pdf file! supplement to koedoe. 1977: 203-209. the conservation role of forestry in south africa d p ackerman secretary, department of forestry private bag x93 pretoria 0001 in september 1965, at the signing ce remony of a national park bill, united states presid ent johnson remarked: " we are living in th e century of change. but if future gen erations are to rem ember us more with gratitude than with sorrow, we must achieve more than just th e miracles of technology. we must also leave them a glimpse of th e world as god really mad e it, not just as it looked when we got through with it." in th e same spirit, forestry in the republic of south africa (rsa) shares the responsibility of conserving and restoring as far as possible the environm ent in which we and succeeding generations must live. the rol e which forestry plays in the economic developm ent of the rsa has long been recognised, but its other major function the stewardship if green landscapes may be overlooked by those who are obsessed with immediate material gain. apart from their task of assuring the supply of timber and other forest produce for their country, foresters hav e a duty to preserve vegetation and oth er natural ass ets on all th e land entrusted to their care which is not devoted to commercial plantations. these natural assets are not preserved purely for their own sake. conservation is for people. conservation costs money and the taxpayer who pays for it naturally wants to see where his money goes and enjoy the fruits of conservation, and for this reason there is a close tie between conservation and outdoor recreation. in som e parts of southern africa nature conservation is mainly orientated to the preservation of indigenous wildlife in gam e parks. many wild animals are dangerous, and visitors to the parks can view them only from motor vehicles or protected lookouts. there is clearly an imperative need for nature reserves in which the public can move without restriction beyond that which is necessary to pres erve the natural wildness. in europe and america forests constitute a significant proportion of nature r eserves and they are managed by foresters to preserve and increase their scientific, aesthetic and recreational valu es. the establishment offorest res erves in the rsa in which the accent is on flora rather than fauna , where visitors can walk and ride, picnic and 203 camp, has to date perhaps not received the attention it deserves because foresters have to some extent been pre-occupied with timber production. nature conservation combined with outdoor recreation and the promotion of tourism, is an aspect of land utilisation that is receiving increasing attention. in natural forests, controlled exploitation need not be excluded and open areas can even be made available to farmers for grazing in times of drought. in this way effective multiple use of the land can be achieved while still retaining public enjoyment of recreation and amenity values as an important object of management. the recreational use of state forest land in the rsa has been sporadic and unplanned for many years. there was a tendency to regard it as a thing to be discouraged. the formulation of a definite policy for facilitating and encouraging outdoor recreation in the state forests was to a large extent the outcome of a visit to the rsa in 1962 by prof c f brockman of the school of forestry and natural resources of the university of washington, who expressed the following views in reporting on his tour: " ... it is quite evident that in addition to wood production your forests , too , can become of increasing importance in outdoor recreation for the following reasons: (i) their scenic beauty and predominantly mountain settings. (ii) accessibility to motorists along a number of main roads and the possibility of opening forest roads to the public. (iii) the possibility of developing a wide variety of interests and activities for visitors in certain areas." the cons ervation role offorestry can be best appreciated if we go back to the original meaning of the word forest. it is derived from the latin ''foris'' meaning " outside" andforestis in the latin of the middle ages came to mean an unenclosed area lying outside the boundaries of villages or parks. originally, therefore, forest referred to all uncultivated or untended land, which in europe would in the main have had trees growing on it, but would also have included scrub, wasteland and what we call veld and savannah. nowadays we think of a forest as a plant community in which a fairly dense growth of trees is dominant. terms such as "high forest", "scrub forest" and " savannah" are used, but those who think of forestry primarily as a commercial undertaking are inclined to envisage a forest essentially as a regimented "plantation" with trees "lined up and lopped and drilled" to create a mechanistic, artifically-maintained sawlog factory, rather than an independently viable association of plants and animals a biocoenose adapted to the environment a forest ecosystem. the direct value of forests as a source of forest produce and the indirect benefits flowing from their amenity values, are both important, 204 but the emphasis on one or other type of benefit has varied because of differences in the socio-economic conditions of the country in the historical period concerned. in ancient times the forest was a refuge for people from their enemies and a shelter from inclement weather. it may also have harboured rogues and criminals. at a later stage many forests were reserved for hunting and pleasure. the direct value of forests as sources of timber and other products did not exceed the indirect values until the industrial revolution sharply increased the demand for timber from the end of the 18th century onwards. the continuous growth of industry since then and the pressure of expanding population are still causing increased demands for timber and other forest products, but in those areas such as western europe and the eastern united states where industrial civilisation has attained its greatest development, the accessory benifits of forest land have come to be accepted as of paramount importance. governments have realised that there is much more to forestry than the establishment and management of plantations for immediate profits, the exploitation of natural forests and the operation of sawmills, pulpmills and other wood-processing plants. the practice of forestry also includes the preservation and management of water catchments, the conservation of nature reserves in natural forests and other categories of land and the development of outdoor recreation facilities. foresters stabilise drifting sands and give expert advice on the choice of trees for woodlots, shade, shelter and ornament on farms and in parks and gardens and along streets in urban areas. the maintenance of adequate green areas is also an essential aspect of the forester's task. you may ask what the department of forestry has done in the rsa to conserve the natural vegetation. the department of forestry is probably best known to the public for the plantations of exotics, mainly pines, which it has established for the production of timber. it is perhaps not so widely realised that the department was the first body in the rsa to concern itself with conservation of the country's indigenous flora. the first step taken to protect the indigenous forests of the southern cape from the destructive exploitation then taking place, was the closure of these forests by the colonial administration of the cape in 1846 at the instance of its forest service. these forests were subsequently re-opened to controlled exploitation but with the passage of time control was intensified by successive legal enactments culminating in 1939 with the de-registration of the remaining woodcutters who were granted state pensions in lieu of their right to fell and market timber from these forests. since that time all indigenous forests belonging to the state have been managed strictly on a conservation basis. active research is being undertaken with the object of rehabilitating and consolidating these forests. legislation aimed at the acquisition of suitable privately-owned indigenous forests has also been enacted. ifforest officers had not fought with dedicated determination to preserve the indigenous forests and if 205 alternative timber resources in the form of plantations had not been established, it can be accepted that very little of these forests would remain today. provision is made in the forest act for the protection of any tree species or forest by proclamation, and a list of protected trees, which may not be felled without the authority of the minister, has been drawn up and published. several private forests in the southern cape have been likewise protected. included in the area controlled by the department are 30 nature reserves comprising some 7 500 ha, which are managed for the conservation of specific rare plant species or ecosystems. the department's main conservation operations, however, are conducted in the mountain catchment reserves which exceed a million hectares in extent. they are managed primarily for the optimum yield of clear water, but this objective is fully compatible with the conservation of the flora and the maintenance of a favourable habitat for wild life. the area of mountain ca tchments is being steadily increased by the acquisition of suitable land . an expanding staff of conservation officers which includes plant and animal ecologists, is employed on research aimed at the establishment of the most effective methods of management for maximum yield of pure, silt-free water in conjunction with protection of the fauna and flora . rare and endangered plant and animal species receive special attention. res earch is also being undertaken into the re-introduction of animals, such as eland, and other antelope into state forests where they formerly lived but have long since disappeared. special conservation research projects on state forest land are being conducted on behalf of the d epartment by universities and provincial nature conservation departmen ts. in recent years there has been a growing awareness, in the rsa as in other western countries, of the need to conserve primitive environments where people of this and future generations can study ecosystems virtually unaltered by man; where they can enjoy landscapes unscarred by roads, railways or power lines; where they can find peace and solitude; and where they can pit their own unaided resources against the forces of nature. although all demarcated forests enjoy a degree of entrenchment under the forest act of 1968, there was a need for the special preservation of selected areas which had retained their original wild character before they too would be changed by technological progress. the forest act was accordingly amended in 1971 to empower the minister, on the recommendation of the national monuments council, to set aside state forests or portions of state forests as wilderness areas for the preservation of forests and natural scenery. areas so proclaimed cannot be alienated nor can any rights be granted over them except with the approval of the senate and the house of assembly. five wilderness areas, totalling an area of nearly 185 000 ha, have already been 206 proclaimed and a number of others, making up an even larger area, are under consideration . to make conservation areas accessible to people, th e department is engaged in the d evelopment of a system of hiking trails on the lines of the appalachian trail in the eastern united states and similar trails in other countries. the south african trail with its supplementary routes is intended ultimately to stretch continuously from the cedarberg in the western cape along the mountains following the southern and eastern coastlines to terminate in the soutpansberg in the northern transvaal. the trail is not confined to state forest land and it is controlled by the national hiking way board on which state departments, provincial administrations, the national parks board and mountain and other outdoor clubs are represented. the system consists basically of hiking trails and walks. hiking trails are longer than walks and include overnight facilities such as huts, shelters and sites for tents. hiking trails are more strenuous than walks and cater for those who wish to carry a backpack or rucksack through relatively undeveloped lands with overnight shelters. walks provide leisurely outings through selected environments lasting a few hours and do not include overnight facilities. the following sections have been completed and are open for public use: the fanie botha hiking trail from near sabie to beyond graskop; the soutpansberg hiking trail along the soutpansberg eastwards from above louis trichardt; and the elephant walk in the diepwalle state forest near knysna. three other sections are due for completion by f ebruary 1977. the national parks board ' s spectacular otter hiking trail along the tsitsikama coast from stormsrivier to grootrivier will also form a link in the system. since the fanie botha hiking trail was opened in 1973, usage has increased and it is expected that there will be regular growth in the number of hikers using the already completed sections. it can be confidently expected that the time will come when a significant proportion of the active population in the more thickly populated areas will be attracted to these gateways to the un trammelled outdoors. the trails and walks can bring large numbers of the public, and particularly the impressionable younger segment, into closer contact with nature and all her wonders. this cannot fail to foster the cause of conservation in the rsa and to bring into many lives a deep appreciation of the need to preserve the green heritage which is their birthright. the department makes a major contribution to the conservation of a number of the country's veld types on its mountain catchment reserves. one of its most important functions is the permanent protection of the cape fynbos which comprises one of the richest and most divers e floras in the world. rapid agricultural and industrial development has resulted in the sacrifice of many hectares offynbos. were it not for the protection 207 by the department of its mountain catchments, the flora in these areas would be sadly depleted. the mountain catchments provide a last refuge for many of south africa's most beautiful flowering plants and several rare and endangered species. through d edicated research and farsighted conservation planning, excellent progress has been made with the rehabilitation of many endangered species . conservation offauna also has a high priority on all state forests, and good progress has been made in this direction. it is not generally realised that plantations of exotic species also provide excellent cover for game, which has increased spectacularly in numbers on many state forests. the periodic burning of firebreaks provides grazing for the game which finds sanctuary in the plantations. large sums of money are spent on the eradication of undesirable alien vegetation growing on state forest land. the problem presented by the intrusion of aggressive exotic species is daunting in its magnitude. it can be tackled successfully only by the application of co-ordinated national effort backed by adequate finance. the matter is receiving attention at the highest levels and continuous research is directed at finding cheaper and more effective methods of control. the valuable work being done by forestry bodies in the reclamation of driftsands along the south african coastline is well known. in this we are following the example of denis the first of portugal, who pioneered the reclamation of coastal dunes 500 years ago by planting them with the maritime pine. the timber from these plantations was used in shipbuilding. reclaimed driftsand has made valuable residential and industrial development possible, especially in the expansion of the cities of cape town and port elizabeth. casuarina species have also been used in the reclamation of coastal drift sands in the summer rainfall area. although progress made by the department in all aspects of nature conservation in the rsa is impressive, much still remains to be done. the scale and scope of the work is being expanded as rapidly as the recruitment of trained conservation staff and availability of funds permit. although the rsa is fortunate in having large areas of open veld and mountain, the threat posed by the encroachment of the growing population and expanding industries is very real. pollution of the atmosphere and water supplies, and indeed of our whole environment, is a danger we dare not ignore simply because it is not as immediately threatening to us as it is in some of the more intensively industrialised countries. the fact must be faced pollution is no longer a problem peculiar to the densely-populated, highly-industrialised countries. it has become a universal problem and we in the rsa must join in the fight against it now. we all have a responsibility to playa full part in exercising the stewardship if the green landscape. let us take timely and effective action to preserve the land entrusted to our care by meticulously planning the conservation of water sources, soil and vegetation, and providing 208 facilities for outdoor recreation to attract tourists as well as our own people out into the open to enjoy pure air, sparkling waters, undisturbed vegetation and protected wildlife. may i conclude by asserting that the planting of trees has a rightful place in these plans. trees are essential to satisfy the national demand for forest products but they can also beautify and conserve the south african environment. 209 page 1 page 2 page 3 page 4 page 5 page 6 page 7 article information authors: dian spear1 melodie a. mcgeoch2 llewellyn c. foxcroft3 hugo bezuidenhout4 affiliations: 1centre for invasion biology, department of botany and zoology, stellenbosch university, south africa 2centre for invasion biology and cape research centre, south african national parks, steenberg, south africa 3centre for invasion biology and conservation services, south african national parks, skukuza, south africa 4scientific services, south african national parks, kimberley, south africa correspondence to: dian spear email: dspear@sun.ac.za postal address: private bag x1, matieland 7602, stellenbosch, south africa dates: received: 01 nov. 2010 accepted: 04 apr. 2011 published: 11 aug. 2011 how to cite this article: spear, d., mcgeoch, m.a., foxcroft, l.c. & bezuidenhout, h., 2011, ‘alien species in south africa’s national parks’, koedoe 53(1), art. #1032, 4 pages. doi:10.4102/koedoe.v53i1.1032 copyright notice: © 2011. the authors. licensee: aosis openjournals. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) alien species in south africa’s national parks in this checklist... open access • acknowledgements • supporting information • references invasive alien species (ias) are one of the major threats to biodiversity in protected areas and pose a significant management challenge (see allen, brown & stohlgren 2009; pyšek, jarošík & kučera 2002). one of the first steps towards managing ias in protected areas is establishing which alien species are present, followed by ongoing surveillance and prevention efforts to combat new introductions (foxcroft et al. 2009). information on the identity and traits of alien species is needed for conducting risk assessments and prioritising species for control, as well as for monitoring management effectiveness in preventing new introductions (mcgeoch et al. 2010). it also provides a first step towards monitoring the extent of occurrence of alien species in national parks.the online checklist that accompanies this summary provides a taxonomic list of alien plant and animal species for south africa’s 19 national parks (including marine protected areas). an online index with common names is also provided. the checklist is intended to serve, (1) as a baseline against which future improvements in knowledge of the alien fauna and flora in south african national parks (sanparks) may be compared and (2) for future monitoring of the success of alien species prevention and control (foxcroft 2009; mcgeoch et al. 2011). the checklist was compiled using a range of information sources, including scientific publications, reference books, the working for water information system (wims), the birds in reserves project database (http://birp.adu.org.za/), sanparks invasive species control unit biocontrol database, park management, lower level and operational plans, kimberley sanparks herbarium (ksan) records, species listed for monitoring with handheld computers loaded with cybertracker software (http://www.cybertracker.org; hereafter referred to as cybertracker data) and communication from specialists and sanparks staff. sources for literature searches included content pages of koedoe (initially the research journal of sanparks, published since 1958; see http://www.koedoe.co.za) and isi web of science and google scholar searches using the names of national parks and the search terms ‘alien’, ‘introduced’ and ‘exotic’. cybertracker data (256 records) were obtained for the addo elephant, agulhas, augrabies falls, camdeboo, golden gate highlands, kalahari gemsbok, karoo, mapungubwe, marakele, mokala, mountain zebra, namaqua and richtersveld national parks (currently held at the geographic information system laboratory at skukuza). thirty-three records were obtained from the ksan for nine parks. in addition to contributions from the authors, data were obtained from seven specialists (on plants, mammals, the harlequin ladybird, painted reed frog and invertebrates). data were also obtained from at least 20 in situ national parks staff in 15 of the national parks, largely via email. plant species names were checked for validity using http://www.theplantlist.org and animal names were checked using a number of databases, including http://www.gbif.org/, http://www.biolib.cz, http://www.itis.gov/ and http://www.fishwise.co.za. taxonomy was assigned according to the angiosperm phylogeny group (2003, 2009) for plants and multiple sources for animals. alien species here refer to species that occur outside their historical distribution ranges. literature and online database searches were conducted to determine indigenous ranges of species and to designate species as alien. the alien species listed include, (1) domestic and livestock animals (see campos et al. 2007 for predation by cats and dogs), (2) extralimital species, that is species that are indigenous to south africa but that have been introduced into national parks outside their historical ranges (see spear & chown 2009a, 2009b for ungulates and their impacts), (3) bird species that have expanded their geographic ranges in response to human modified habitats, for example hadeda ibis, bostrychia hagedash (macdonald, richardson & powrie 1986) and (4) biological control agents released to control invasive alien plants. a substantial number of records (54%) were obtained from sources other than primary literature, such as from databases, specialists, park management plans, park management staff and rangers (figure 1). alien plants, freshwater fish, marine organisms and snails are the best studied taxa in national parks (based on the number of publications) and some national parks have been relatively well studied compared with others (table 1; online checklist). plants contribute most to alien species richness in national parks and a similar dominance of alien species lists by plants has been shown globally (delivering alien invasive species inventory for europe 2009; mcgeoch et al. 2010). there are particular taxa that are considered most likely to be under-represented on these lists (see table 2 for representation of taxonomic groups). commensal species (species benefiting from human habitation and food) associated with buildings and dwellings in national parks are one such group, such as cockroaches (these are reported for kruger national park but not for other national parks) and mice and rats (mus musculus and rattus rattus are reported for only three national parks) (online checklist). domestic animals (e.g. cats, felis catus and dogs, canis familiaris) and livestock are also likely to be in the vicinity of most national parks and, if they are not resident in national parks, may be transient visitors. national parks that are surrounded by game farms are also likely to be subject to extralimital and alien game species intrusions (spear & chown 2009a) and these mammal species may be missing from lists for some national parks. there are likely to be many more alien invertebrates (e.g. insects, springtails and earthworms) in sanparks than recorded here. the insect taxa currently listed are largely biological control agents, with little record of other alien insects. insects that may be present but yet unrecorded in some national parks include the european wasp, vespula germanica, the harlequin ladybug, harmonia axyridis and the argentine ant, linepithema humile. vespula germanica is found in table mountain national park and could be in other national parks in the western cape province (see tribe & richardson 1994), whereas harmonia axyridis has been recorded in the vicinity of addo elephant, bontebok, mountain zebra and west coast national parks (lambert smith pers. comm.). other groups that are likely to be under-represented on alien species lists for sanparks include grasses (milton 2004), garden plants (although well studied for the kruger national park; foxcroft, richardson & wilson 2008) and marine organisms (griffiths et al. 2010). as with most alien species lists, those provided here are thus almost certainly incomplete (see pyšek et al. 2008) and may also include inaccuracies as a result of one or more of the following factors, (1) inadequate surveys, inventories and knowledge, (2) taxonomic uncertainty and species misidentification (including cryptogenic species), (3) outdated historical records that may no longer be accurate, (4) unpublished or grey literature information is not always accessible and (5) inadequate information on species historical ranges. nonetheless, over time and with the planned increase in research, surveillance and monitoring of ias in national parks (mcgeoch et al. 2011), the list accuracy and completeness may be incrementally improved. the publication of these baseline lists will also contribute to future assessments of the relative contribution of increased knowledge versus new invasions and extirpations (see costello & solow 2003) to the status of, and trends in, alien species in sanparks. figure 1: percentage of species records from different data sources, namely peer-reviewed publications and reference books (publication), park management plans and lower level alien and rehabilitation plans (man. plan.), working for water information management system (wims), ex-situ specialists and sanparks herbarium (experts), and sanparks in situ staff and cybertracker data (staff). table 1: number of alien species records per national park for different taxa, as collated from peer-reviewed publications. table 2: summary of alien species in south african national parks (sanparks) by taxonomic group. the number of species per group and number of national parks in which each taxonomic group occurs are shown, as well as the environmental associations of the species. acknowledgements (back to top) this research was funded by the dst-nrf centre for invasion biology and the global invasive species programme. the following people are thanked for assistance with information on alien species in sanparks: charmaine uys, james pryke, marna herbst, peter novellie, riaan stals, sarah davies, tony rebelo, johan baard, leighan mossop, abel ramavhale, andrew deacon, bernard van lente, carli venter, conrad strauss, elton le roux, ernest daemane, ettienne fourie, geoff nichols, giel de kock, guin zambatis, hendrik sithole, ilse welgemoed, johan de klerk, johan taljaard, lesley henderson, letsie coetzee, mphadeni nthangeni, nollie bosman, paddy gordon, peter burdett, pierre nel, riaan nel, robyn wood, ruth-mary fisher, samantha schroder, steven khosa and thys ahrends. isaiah moyo and working for water are thanked for providing data from the wims. sandra macfadyen is thanked for providing cybertracker data. two anonymous reviewers are thanked for commenting on the manuscript. supporting information (back to top) additional supporting information may be found in the online version of this article, namely:• online checklist: lists of alien plant and animal taxa in south africa’s 19 national parks. • online index: lists of animal and plant species and their common names. references (back to top) allen, j.a., brown, c.s. & stohlgren, t.j., 2009, ‘non-native plant invasions of united states national parks’, biological invasions 11, 2195−2207. doi:10.1007/s10530-008-9376-1angiosperm phylogeny group, 2003, ‘an update of the angiosperm phylogeny group classification for the orders and families of the flowering plants: apg iii’, botanical journal of the linnean society 141, 399−436. doi:10.1046/j.1095-8339.2003.t01-1-00158.x angiosperm phylogeny group, 2009, ‘an update of the angiosperm phylogeny group classification for the orders and families of flowering plants: apg iii’, botanical journal of the linnean society 161, 105−121. doi:10.1111/j.1095-8339.2009.00996.x campos, c.b., esteves, c.f., ferraz, k.m.p.m.b., crawshaw jr., p.g. & verdade, l.m., 2007, ‘diet of free-ranging cats and dogs in a suburban and rural environment, south-eastern brazil’, journal of zoology 273, 14−20. doi:10.1111/j.1469-7998.2007.00291.x costello, c.j. & solow, a.r., 2003, ‘on the pattern of discovery of introduced species’, proceedings of the national academy of sciences, united states of america 100, 3321−3323. doi:10.1073/pnas.0636536100, pmid:12615995, pmcid:152290 delivering alien invasive species inventory for europe (eds.), 2009, handbook of alien species in europe, springer, dordrecht. foxcroft, l.c., 2009, ‘developing thresholds of potential concern for invasive alien species: hypotheses and concepts’, koedoe 51, 1−6. doi:10.4102/koedoe.v51i1.157 foxcroft, l.c., richardson, d.m., rouget, m. & macfadyen, s., 2009, ‘patterns of alien plant distribution at multiple spatial scales in a large national park: implications for ecology, management and monitoring’, diversity and distributions 15, 367−378. doi:10.1111/j.1472-4642.2008.00544.x foxcroft, l.c., richardson, d.m. & wilson, j.r.u., 2008, ‘ornamental plants as invasive aliens: problems and solutions in kruger national park, south africa’, environmental management 41, 32−51. doi:10.1007/s00267-007-9027-9, pmid:17943344 griffiths, c.l., robinson, t.b., lange, l. & mead, a., 2010, ‘marine biodiversity in south africa: an evaluation of current states of knowledge’, plos one 5, e12008. doi:10.1371/journal.pone.0012008, pmid:20689849, pmcid:2914023 macdonald, i.a.w., richardson, d.m. & powrie, f.j., 1986, ‘range expansion of the hadeda ibis bostrychia hagedash in southern africa’, south african journal of zoology 21, 331−342. mcgeoch, m.a., butchart, s.h.m., spear, d., marais, e., kleynhans, e.j., symes, a. et al., 2010, ‘global indicators of alien species invasion: threats, biodiversity impact and responses’, diversity and distributions 16, 95−108. doi:10.1111/j.1472-4642.2009.00633.x mcgeoch, m.a., dopolo, m., novellie, p., hendricks, h., freitag, s., ferreira, s. et al., 2011, ‘a strategic framework for biodiversity monitoring in sanparks’, koedoe 53(2), art. #991, 10 pages. doi:10.4102/koedoe.v53i2.991 milton, s.j., 2004, ‘grasses as invasive alien plants in south africa’, south african journal of science 100, 69−75. pyšek, p., jarošík, v. & kučera, t., 2002, ‘patterns of invasion in temperate nature reserves’, biological conservation 104, 13−24. doi:10.1016/s0006-3207(01)00150-1 pyšek, p., richardson, d.m., pergl, j., jarošik, v., sixtová, z. & weber. e., 2008, ‘geographical and taxonomic biases in invasion ecology’, trends in ecology and evolution 23, 237−244. doi:10.1016/j.tree.2008.02.002 spear, d. & chown, s.l., 2009a, ‘the extent and impacts of ungulate translocations: south africa in a global context’, biological conservation 142, 353−363. doi:10.1016/j.biocon.2008.10.031 spear, d. & chown, s.l., 2009b, ‘non-indigenous ungulates as a threat to biodiversity’, journal of zoology 279, 1−17. doi:10.1111/j.1469-7998.2009.00604.x tribe, g.d. & richardson, d.m., 1994, ‘the european wasp, vespula germanica (fabricius) (hymenoptera: vespidae) in southern africa and its potential distribution as predicted by ecoclimatic matching’, african entomology 2, 1−6. article information authors: helga van coller1 frances siebert1 stefan j. siebert1 affiliations: 1school of biological sciences, north-west university, south africa correspondence to: helga van coller postal address: private bag x6001, potchefstroom 2520, south africa dates: received: 12 sept. 2012 accepted: 21 dec. 2012 published: 18 mar. 2013 how to cite this article: van coller, h., siebert, f. & siebert, s.j., 2013, ‘herbaceous species diversity patterns across various treatments of herbivory and fire along the sodic zone of the nkuhlu exclosures, kruger national park’, koedoe 55(1), art. #1112, 6 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1112 copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. herbaceous species diversity patterns across various treatments of herbivory and fire along the sodic zone of the nkuhlu exclosures, kruger national park in this original research... open access • abstract • introduction    • study area • methods    • experimental design and sampling • results • discussion • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ understanding relationships between large herbivores and plant species diversity in dynamic riparian zones is critical to biodiversity conservation. the nkuhlu exclosures in the kruger national park (knp) provided opportunity to investigate spatial heterogeneity patterns within riparian zones, as well as how these patterns are affected by fire and herbivory. a monitoring project was initiated to answer questions about the dynamics of the herbaceous layer and was aimed at determining, (1) whether there exists meaningful variance in herbaceous plant species richness and diversity across different treatments in the ecologically sensitive sodic zone and (2) whether an increase in herbaceous biomass, an artefact of herbivory and fire exclusion, suppresses herbaceous plant species diversity and richness. herbaceous vegetation was sampled in two 1 m2 circular sub-plots in the eastern and western corners of 81 fixed plots. the biomass of each plot was estimated with a disc pasture meter (dpm) diagonally with the plot. dpm-readings were converted to kg/ha, according to the latest conversions for the lowveld savanna. species richness and biomass showed significant variance across treatments, whereas no significant variation in herbaceous species diversity was perceived. combined treatment of fire absence and herbivore presence contributed to higher forb species richness in the sodic zone. biomass is significantly higher in fully fenced areas where herbivores are excluded, as opposed to the open and partially fenced areas. although no significant variation was recorded for diversity across treatments, lowest diversity was recorded in the absence of all herbivores, especially in combination with fire treatment. therefore herbivores are essential in sustaining herbaceous plant species richness in the sodic zone, whilst no significant results were found with regard to their effect on species diversity. although statistically non-significant, fire seems to suppress species richness.conservation implications: this study could be used as framework to advance and develop science-based management strategies for, at least, the sodic zones of the knp. research in these exclosures will create better understanding of these landscapes, benefit ecosystem conservation planning of national parks and also provide valuable long-term information on key ecological processes. introduction top ↑ sodic patches are often associated with footslopes of undulating granitic landscapes (venter 1990) and are commonly referred to as ‘nutrient hotspots’ as they produce high quality forage (grant & scholes 2006). deep, duplex soil containing dispersed clay with a low infiltration capacity and high concentrations of nitrogen, phosphorus and sodium characterise sodic patches (dye & walker 1980; khomo & rogers 2005). sodic soil is associated with reduced hydraulic conductivity as a result of the hyper-accumulation of exchangeable sodium and is prominent in landscapes derived from sodium-releasing parent material such as granite (khomo & rogers 2005). soil is considered ‘sodic’ when high concentrations of soluble salts prevail, which are dominated by sodium, the ph is greater than 8.5 and electrical conductivity is less than 4.0 ds/m (tarasoff, mallory-smith & ball 2007). the level of sodium in soil is often reported as the sodium adsorption ratio (sar). soil is classified as sodic when the sar is above 13; at levels below 13, sodium can cause deterioration of soil structure and problems with water infiltration (davis, waskom & bauder 2012). sodic patches form an integral part of savanna ecosystems because of the ecosystem services and functioning they provide, such as, (1) the accumulation of nutrients, (2) the provision of open spaces for predator vigilance and (3) the formation of wet season wallowing points (khomo & rogers 2005). furthermore, these key resource areas sustain animal body condition for dry season survival and support reproduction through nutritional benefits (grant & scholes 2006). despite their ecological significance, sodic patches are often considered as desolate land because of their disturbed appearance and low aesthetic value (khomo & rogers 2005) caused by overgrazing, which, in turn, impacts on herbaceous species diversity, richness and biomass (jacobs & naiman 2008). fire and herbivory are regarded as important ecosystem modifiers, especially in semi-arid southern african savannas, often acting together (archibald et al. 2005). fire and herbivory are considered as dominant mediators of vegetation change in the kruger national park (knp) and interact strongly with rainfall unpredictability and prolonged droughts (du toit, rogers & biggs 2003). the impact of fire and herbivory on herbaceous biomass dynamics is, however, poorly understood (archibald et al. 2005; jacobs & naiman 2008). the establishment of the nkuhlu exclosures in the knp in 2001 provided an opportunity to study the spatial and temporal heterogeneity patterns within and around riparian zones, as well as how these patterns are affected by herbivory and fire. by excluding these drivers from the system it is believed that their effects could be determined over time (o’keefe & alard 2002). jacobs and naiman (2008) were the first to report on the relationship between herbivory and species richness at the nkuhlu exclosures. after 2 years of herbivory exclusion, a marked increase in standing biomass was recorded, along with lower species richness in the absence of herbivores. these patterns were, however, less significant along the sodic zone (jacobs & naiman 2008). this article is therefore aimed at reporting on herbaceous plant species richness, diversity and biomass specifically along the sodic zone of the nkuhlu exclosures after an additional 5-year cycle of herbivory and fire exclusion. the objectives of this study were to test, (1) whether significant variation in herbaceous plant species diversity, richness and biomass could be detected across different combined treatments of herbivory and fire and (2) whether increasing biomass, an artefact of herbivory and fire exclusion, suppresses diversity and richness of herbaceous plant species in the sodic zone of the nkuhlu exclosures. study area the nkuhlu large-scale long-term exclusion experiment (24°58′s, 31°46′e) is situated on the northern bank of the sabie river, approximately 18 km downstream from skukuza (siebert & eckhardt 2008). this semi-arid, subtropical savanna has two distinct seasons: a hot, sporadically wet growth season from october to april and a warm, dry, non-growing season (scogings et al. 2012). the mean annual rainfall is 561 mm, whilst temperatures range from a mean minimum of 5.6 ºc in winter to a mean maximum of 32.6 ºc in the summer months. crests and midslopes are characterised by coarse, shallow, sandy soil overlaying weathered rock, whilst footslopes below the seep-line are characterised by deep, duplex soil (scogings et al. 2012; siebert & eckhardt 2008). the duplex soil on footslopes is referred to as the sodic zone in this article. on a national scale, the nkuhlu exclosures are located within the granite lowveld vegetation unit (svi 3) (mucina & rutherford 2006). it is characterised by tall shrubland with few trees to moderately dense low woodland on deep sandy uplands. the dense herbaceous layer is dominated by the graminoids aristida congesta, digitaria eriantha and panicum maximum. the riparian zone of the exclosures forms part of the subtropical alluvial vegetation unit (aza 7), which is described by mucina and rutherford (2006) as having flat alluvial riverine terraces that supports a complex channel of flowing rivers and river-fed pans, marginal reed belts, as well as flooded grasslands, ephemeral herblands and riverine thickets. the sodic zone plant community was described by siebert and eckhardt (2008) as sporobolus nitens–euclea divinorum dry sodic savanna, covering roughly 23% of the total sampling area of the nkuhlu exclosures. the sodic plant community supports contrasting vegetation and herbivore activities that differ from those found on upland savanna soil (du toit et al. 2003). it stretches across the exclosure fence lines and shows noticeable visual differences amongst herbivory and fire treatments. the diagnostic and dominant herbaceous species of the sodic community include forbs, such as abutilon austro-africanum, portulaca kermesina and ocimum americanum, and graminoids, such as chloris virgata, enteropogon monostachyus and s. nitens (siebert & eckhardt 2008). methods top ↑ experimental design and sampling the nkuhlu exclosures comprise 139 ha of semi-arid savanna in the riparian zone of the sabie river and consist of three herbivory treatments, each divided into a fire and no-fire treatment, hence six treatment combinations overall (figure 1). the herbivory treatments consist of, (1) a partially fenced area of 44 ha that was designed to specifically exclude elephants (giraffes are also excluded because their body size), (2) an open, unfenced area of 25 ha (referred to as the ‘control site’) and (3) a fully fenced area of 70 ha, which was designed to exclude all herbivores larger than a hare (siebert & eckhardt 2008). figure 1: experimental layout of 12 transects within six different fire and herbivory treatments of the nkuhlu exclosures, kruger national park. treatment 6 was excluded from this study to include sodic sites only. vegetation of sodic soil is considered to be more palatable by large herbivores, particularly grazers and mixed feeders (intermediate feeders) than crest vegetation (scogings 2011). mammalian herbivores residing in the study area include: browsers – such as giraffe (giraffa camelopardalis), black rhino (diceros bicornis), greater kudu (tragelaphus strepsiceros) and the scrub hare (lepus saxatilis), grazers – such as cape buffalo (syncerus caffer), hippo (hippopotamus amphibius), blue wildebeest (connochaetes taurinus) and plains zebra (equus quagga), and mixed or intermediate feeders – such as impala (aepyceros melampus), african elephant (loxodonta africana) and steenbok (raphicerus campestris) (scogings et al. 2012). the density of elephants in the study area fluctuates, but has been estimated and considered high at 0.5 km-2 – 2.0 km-2 (scogings et al. 2012).fire treatment of the nkuhlu exclosures was dependant on whether the larger knp burn blocks (fire management units) in which the exclosures lie were burned irrespective of the source and the actual extent of the fire. fires occurred in october 2002 and august 2007. floristic sampling, which conformed to the guidelines stipulated in the field operations manual for herbivore and fire exclosures (o’keefe & alard 2002), was conducted during the 2010 rainy season (i.e. october–march) (van wyk & fairall 1969) when plant cover was at its maximum and most plant species were present (keller 2002). all treatments were sampled, although only data from the sodic zone were analysed for this study. as the sodic zone did not extend into the control site with fire, two transects could not be considered for floristic analyses. each treatment of fire and/or herbivory consisted of two permanent transects (figure 1). transects stretch from the river to the crest of the catena and run perpendicular to the river. permanent transects were marked with metal droppers. fixed plots were situated 5 m downstream from each transect dropper. the position of transects and plots were plotted on the vegetation map of the nkuhlu exclosures (siebert & eckhardt 2008) to identify plots that are representative of the sodic zone. the data of 81 fixed plots of 10 m × 20 m (long side parallel to the river channel) within the sodic zone (figure 2) were analysed. herbaceous vegetation was sampled in two 1 m2 circular sub-plots at the eastern (upland downstream) and western (river upstream) corners of each fixed plot respectively (figure 2). only herbaceous individuals rooted within the circle were recorded. each living individual (forbs and grasses) was counted and all species names conform to germishuizen and meyer (2003). the biomass of each plot was estimated, by taking 10 readings along the diagonal of the plot, with a lowveld savanna-calibrated disc pasture meter (dpm) (figure 2). these readings were converted to kg/ha according to the latest conversion of dpm-readings to biomass for the lowveld savanna (zambatis et al. 2006). two equations for dpm height groups (≤ 26 cm and > 26 cm) were proposed by zambatis et al. 2006: kg.ha-1 = [31.7176 (0.32181/x) x0.2834]2 r = 0.9796; r2 = 0.951; p < 0.0005 [eqn 1] kg.ha-1 = [17.3543 (0.9893x) x0.5413]2 r = 0.948; r2 = 0.882; p < 0.0005 [eqn 2] where x is the mean dpm height (in cm) of a site. species richness is the number of species present in a defined area (begon, townsend & harper 2006). in this study, richness was considered as the number of species per circular sub-plot (1 m2). species diversity was calculated for the plots with the shannon-wiener index of diversity (h’), which is sensitive to both species richness and the number of individuals (abundance) per species (keller 2002). the greater the number of species and the higher the evenness (proportion of each species), the higher the h’-value, which suggests that the diversity in the plot is high (begon et al. 2006; keller 2002). the function h’ used in this study is: h’ = -∑ (pi.ln pi) … [eqn 3] where pi is the relative abundance of the ith species. this index provides a rough measure of diversity, which is much less biased by sample size than species richness. to test for significant variation in species diversity, richness and biomass of the herbaceous layer amongst the five different treatments of fire and herbivory, one-way analysis of variance (anova) was applied to the data set in statistica version 10 (2010). where statistically significant results were obtained, post-hoc pair-wise comparisons of the treatments – using the tukey unequal n hsd (honestly significant difference) test – were conducted. results top ↑ analysis of variance (table 1) for all variables revealed significant effects of herbivory, with or without fire, on both biomass (p < 0.0001) and species richness (p = 0.0002) across the different treatments, whilst the effects of herbivory, with or without fire, on species diversity was non-significant (p = 0.4). figure 2: positioning of the 1 m2 sub-plots for herbaceous species sampling and the sampling line for disc pasture meter readings within a typical fixed plot along a transect at the nkuhlu exclosures, kruger national park. the corners of each plot are permanently marked with metal droppers which represent a different position parallel to the sabie river. table 1: one-way analysis of variance of the mean values for herbaceous biomass and species richness and diversity across treatments along the sodic zone of the nkuhlu exclosures, kruger national park. figure 3: mean biomass (p < 0.0001) across fire and herbivory treatments in the sodic zone of the kruger national park. figure 4: mean species richness (p = 0.0002) across fire and herbivory treatments in the sodic zone of the kruger national park. mean biomass (kg/ha) varied significantly across treatments (p = 0.0001) (table 1; figure 3). as would be expected, mean biomass was highest in the fully fenced plots as a result of herbivore exclusion. in pair-wise comparisons of biomass means, significant differences were revealed between the control (i.e. open, no fire) site with unrestricted herbivory and the fully fenced area, both with (tukey: p < 0.05) and without fire (tukey: p < 0.05) treatment. furthermore, both of the partial herbivore treatments showed significant differences compared to both fully fenced herbivore exclusion treatments. although the effect of fire seems secondary to herbivory, mean biomass was consistently lower in the fire treatments (figure 3).a total of 187 herbaceous plant species were recorded in the sodic zone of the nkuhlu exclosures. species richness varied significantly across the combinations of herbivory and fire (p = 0.0002) (table 1). highest species richness was associated with areas exposed to herbivory (figure 4), although species richness in the control site (i.e. open, no fire) and the partially fenced treatment without fire was significantly higher compared to the fully fenced treatment with fire. species richness in the partially fenced site with fire was, however, not significantly higher than the full exclosure treatments (tukey: p > 0.05). shannon-wiener diversity index values showed no significant variation across the five combinations of herbivory and fire (p = 0.4) (table 1). discussion top ↑ the effect of herbivore exclusion on total herbaceous biomass has been shown to be lower in the sodic zone than elsewhere in the riparian zone (jacobs & naiman 2008). this phenomenon could be attributed to the drier conditions prevailing in the sodic zone throughout the year, as well as the shallow surface soils and relative impenetrability, making herbaceous species establishment and growth even more challenging. as would be expected, the exclusion of herbivores from the sodic zone lead to herbaceous biomass increases. the biomass range of 2500 kg/ha – 2900 kg/ha (figure 3) in the full exclosure is unusually high for sodic sites in semi-arid savanna ecosystems, because sodic soil is usually sparsely vegetated and therefore associated with land degradation and gully formation that can become self-reinforcing as a result of its susceptibility to sheet erosion (khomo & rogers 2005). the exclusion of herbivores, however, brings forth a gradual improvement in soil surface conditions, which re-establishes the predominance of topsoil water and allows for an increase in the biomass of grasses (walker et al. 1981).furthermore, herbivory reduces biomass and canopy cover of certain plants and promotes spatial heterogeneity (jacobs & naiman 2008). sodic zones are favoured as forage patches by large herbivores, which gain essential nutrients from the herbaceous species growing there. the total exclusion of these herbivores from savanna riparian zones leads to a loss of game paths, feeding patches and wallows, causing a reduction in plant community heterogeneity (jacobs & naiman 2008). according to bakker, blair and knapp (2003), this heterogeneity, caused by different patches with long and short biomass, is thought to enhance the coexistence of a greater number of herbaceous species, hence boosting species richness. although jacobs and naiman (2008) found no consistent effects of increasing biomass on species richness in the drier sodic zone after 4 years of herbivory exclusion (i.e. 2004/2005), our results indicated that the significantly higher biomass in the absence of herbivory seemed to suppress species richness after 10 years of herbivory exclusion. highest species richness was associated with areas exposed to herbivory (figure 4), thus areas where plant community heterogeneity was highest. no significant variation in species diversity could be confirmed. as the shannon-wiener index of species diversity acknowledges the abundance of individual species, it could be expected that variation in species diversity is less perceptible than species richness. the sodic zone, in particular, is often dominated by few forb species that are well adapted to harsh conditions. jacobs and naiman (2008) reported that the succulent forb, trianthema salsoides, contributed to a substantial proportion of the forb biomass in 2003–2004. although t. salsoides was still prevalent in the 2010 data set, it was replaced as a dominant forb species by achyranthes aspera. dominance by a single species reduces the evenness of the sodic zone ecosystem, hence suppressing diversity. lowest species richness was recorded in areas where herbivores were excluded. this is in accordance with oba, vetaas and stenseth (2001) who suggested that long-term exclusion of herbivores may not necessarily have an increasing effect on species diversity and richness. the dynamic character of the vegetation in the sodic zone ecosystem therefore results in less predictable temporal changes in herbaceous species diversity. conclusion top ↑ in contradiction with the ‘intermediate disturbance hypothesis’, which suggests that highest diversity is maintained at intermediate levels of disturbance (begon et al. 2006), highest species richness was associated with highest disturbance (i.e. open treatment). increasing biomass as a result of the absence of herbivory suppresses the richness of herbaceous species in the sodic zone, although species richness is even lower in its combined effect of fire treatment. however, whilst species diversity patterns showed no significant differences across the treatments, lowest diversity was also measured in the absence of herbivory, but with fire treatment. it is therefore inferred that, although sodic patches are often associated with overgrazed, trampled vegetation, herbivores play an important role in the maintenance of species richness and diversity in these nutrient hotspots. although statistically not significant, fire seems to suppress diversity and richness of the herbaceous layer in the sodic riparian zone. floristic monitoring over a larger time scale under similar conditions is however required to support these premature results. figure 5: mean shannon-wiener diversity index values (p = 0.4) across fire and herbivory treatments in the sodic zone of the kruger national park. the combined assemblage of herbivores with (open treatment), or without (partial treatment) elephants, has the largest reduction in herbaceous standing biomass within the sodic zone of the nkuhlu exclosures. herbivory is suggested to be the primary driver of herbaceous vegetation dynamics in the sodic zone along the sabie river, whether elephants are being included (open treatment) or excluded (partial treatment). whilst suppressing standing biomass, herbivory maintains higher levels of herbaceous species richness. fire, however, is suggested to be a secondary driver within the herbivory treatments, because it seems to suppress both herbaceous species richness and standing biomass. despite clear patterns, no significant variation or differences with regard to herbaceous species diversity could be detected across treatments, which necessitates further long-term monitoring of herbaceous vegetation at nutrient hotspots along the sabie river. as the conservation of total biodiversity is the mandate of south african national parks (sanparks), these findings have direct implications for the management of sodic zones in the knp. the exclusion of herbivores may alter ecosystem functioning, especially at smaller scales, by increasing biomass at the cost of herbaceous species richness. herbivores play an essential role in the reduction of herbaceous plant competition, in that herbivores decrease grass biomass which, in turn, promotes herbaceous species richness because fast-growing grasses can no longer overtop and shade out forbs. herbivores are therefore considered imperative in the maintenance of species richness and diversity and ecosystem functioning in african savanna ecosystems, especially in sodic zones. acknowledgements top ↑ the authors would like to acknowledge sanparks for general logistical support. we are also grateful to the research unit: environmental sciences and management, north-west university for financial support. in addition, our appreciation goes to thomas rikonibe for assistance and protection in the field, as well as philip ayres, hannes myburgh and pieter kloppers, who provided valuable assistance with field surveys. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions h.v.c. (north-west university) was a post-graduate student who collected, analysed and reported the data, whilst f.s. (north-west university) was the project leader and supervisor to h.v.c. and was responsible for project design, data collection, analyses and reporting of data. s.j.s. (north-west university) made contributions in data sampling and reporting, whilst also fulfilling the role of co-supervisor. h.v.c., f.s. and s.j.s. wrote the manuscript. references top ↑ archibald, s., bond, w.j., stock, w.d. & fairbanks, d.h.k., 2005, ‘shaping the landscape: fire-grazer interactions in an african savanna’, ecological applications 15, 96–109. http://dx.doi.org/10.1890/03-5210 bakker, c., blair, j.m. & knapp, a.k., 2003, ‘does resource availability, resource heterogeneity or species turnover mediate changes in plant species richness in grazed grasslands?’, oecologia 137, 685–391. http://dx.doi.org/10.1007/s00442-003-1360-y, pmid:12955488 begon, m., townsend, c.r. & harper, j.l., 2006, ecology from individuals to ecosystems, blackwell publishing, melbourne. davis, j.g., waskom, r.m. & bauder, t.a., 2012, managing sodic soils, viewed 11 september 2012, from http://www.ext.colostate.edu/pubs/crops/00504.html du toit, j., rogers, k.h. & biggs, h.c., 2003, the kruger experience: ecology and management of savanna heterogeneity, island press, washington, dc. dye, p.j. & walker, b.h., 1980, ‘vegetation-environment relations on sodic soils of zimbabwe rhodesia’, journal of ecology 68, 589–606. http://dx.doi.org/10.2307/2259424 germishuizen, g. & meyer, n.l. (eds.), 2003, ‘plants of southern africa: an annotated checklist’, strelitzia 14, south african national botanical institute, pretoria. grant, c.c. & scholes, m.c., 2006, ‘the importance of nutrient hot-spots in the conservation and management of large wild mammalian herbivores in semi-arid savannas’, biological conservation 130, 426–437. http://dx.doi.org/10.1016/j.biocon.2006.01.004 jacobs, s.m. & naiman, r.j., 2008, ‘large african herbivores decrease herbaceous plant biomass while increasing plant species richness in a semi-arid savanna toposequence’, journal of arid environments 72, 891–903. http://dx.doi.org/10.1016/j.jaridenv.2007.11.015 keller, b.e.m., 2002, ‘plant diversity in lythrum, phragmites, and typha marshes, massachusetts, usa’, wetlands ecology and management 8, 391–401. http://dx.doi.org/10.1023/a:1026505817409 khomo, l.m. & rogers, k.h., 2005, ‘proposed mechanism for the origin of sodic patches in kruger national park, south africa’, african journal of ecology 43, 29–34. http://dx.doi.org/10.1111/j.1365-2028.2004.00532.x mucina, l. & rutherford, m.c. (eds.), 2006, the vegetation of south africa, lesotho and swaziland, south african national biodiversity institute, pretoria. oba, g., vetaas, o.r. & stenseth, n.c., 2001, ‘relationships between biomass and plant species richness in arid-zone grazing lands’, journal of applied ecology 38, 836–845. http://dx.doi.org/10.1046/j.1365-2664.2001.00638.x o’keefe, t. & alard, g., 2002, ‘effects of herbivores and fire on riparian and upland savanna ecosystems’, unpublished report on field operations manual for herbivore and fire exclosures on the sabie and letaba rivers in the kruger national park. scogings, p.f., 2011, ‘stem growth of woody species at the nkuhlu exclosures, kruger national park: 2006–2010’, koedoe 53(1), art. #1035, 8 pages. http://dx.doi.org/10.4102/koedoe.v53i1.1035 scogings, p.f., johansson, t., hijáltén, j. & kruger, j., 2012, ‘response of woody vegetation to exclusion of large herbivores in semi-arid savannas’, austral ecology 37, 56–66. http://dx.doi.org/10.1111/j.1442-9993.2011.02249.x siebert, f. & eckhardt, h.c., 2008, ‘the vegetation and floristics of the nkuhlu exclosures, kruger national park’, koedoe 50, 126–144. http://dx.doi.org/10.4102/koedoe.v50i1.138 statistica version 9.1, 2010, statsoft, inc., tulsa. tarasoff, c.s., mallory-smith, c.a. & ball, d.a., 2007, ‘comparative plant responses of puccinellia distans and puccinnellia nuttalliana to sodic versus normal soil types’, journal of arid environments 70, 403–417. http://dx.doi.org/10.1016/j.jaridenv.2007.01.008 van wyk, p. & fairall, n., 1969, ‘the influence of the african elephant on the vegetation of the kruger national park’, koedoe 12(1), 57–76. http://dx.doi.org/10.4102/koedoe.v12i1.747 venter, f.j., 1990, ‘a classification of land use for management planning in the kruger national park’, phd thesis, department of geography, university of south africa. walker, b.h., ludwig, d., holling, c.s. & peterman, r.m., 1981, ‘stability of semi-arid savanna grazing systems’, journal of ecology 69, 473–498. http://dx.doi.org/10.2307/2259679 zambatis, n., zacharias, p.j.k., morris, c.d. & derry, j.f., 2006, ‘re-evaluation of the disc pasture meter calibration for the kruger national park, south africa’, african journal of range and forage science 23(2), 85–97. http://dx.doi.org/10.2989/10220110609485891 filelist convert a pdf file! koedoe 19: 153-164 (1976) population characteristics and recent history of lions in 1wo parts of the kruger national park g. l. smuts dept. of nature conservation private bag x404 skukuza 1350 abstract a survey entailing the capture and marking of lions was undertaken in two parts of the kruger national park, republic of south africa, during 1974 and 1975. a total of 347 lions were captured and marked while 62 were destroyed as part of an experimental cropping operation. sixty out of 66 prides contained marked lions . the central district (5 560 km 2) had a minimum population of 708 lions (60 prides) while the crocodile bridge area (463 km2) had 67 lions . adult sex ratios differed between areas with a range of 10:1,79 to 10:2,49 and was density dependent. sex ratios for all age classes together varied between 10: 1,49 and 10: 1,79 (n = 591 lions). age composition for the central district was: adults 53,3%; sub-adults 17,1%; large cubs 8,8%; small cubs 20,9% (n = 708 lions). lion density (all ages) ranged h-om 10,5 to 15,5 lions/loo km 2 while lion biomass ranged from 11,0 to 18,2 kg/km2. pride sizes ranged from four to 21 with a mean of 11,8 for the whole study area. average number of adult males per pride was 2,1 with a maximum of five. ten marked lions were destroyed outside the park and part of one pride was migratory within the park. the past and present distribution of prides and their sizes, indicated that the lion population had increased considerably and this increase could be correlated with an increase in the number of artificial waterholes and the stabilization of existing waterholes. since 1933, 29 dams were built and 64 windmills erected in the study areas, with the number of lions and their prey populations more than doubling since this date. introduction with the construction of the 500 km long western boundary fence of the kruger national park (knp), republic of south africa, in 1961 and the new game proof fence on the east (completed early in 1976), it has become increasingly necessary to assess the status of the now enclosed 153 game populations. although most medium-sized and large herbivores are censused annually, comparable data for the larger carnivores are almost non existent. at the same time the status of certain ungulate populations has indicated that predation is a major factor influencing the growth of these populations. to gain a better insight into predatorprey relationships, it was decided to initially undertake a survey of the lions panthera leo krugeri in the central district and the crocodile bridgelower sabi area of the knp (fig. 1). this survey was subsequently designed to take the form of a census while at the same time data could also be collected on characteristics relevant to the population dynamics, general distribution and movements of lions. due to the practical problems involved in censusing lions, there is relatively little published data on aspects such as density, biomass, sex and age composition of lion populations in large areas. the only really reliable data are those for small study areas such as the nairobi national park (rudnai 1973) and the ngorongoro crater (schaller 1972, kruuk 1972), where individuals or groups could be identified. other than these, population sizes have largely been estimated by a variety of unsatisfactory methods. more recently three rough survey methods were described for the eastern selous game reserve by rodgers ( 1974). material and methods the survey method employed is described in detail by smuts, whyte and dearlove (1976), and entailed the capture and marking oflions from each group encountered. the areas surveyed were the central district and the crocodile bridge-lower sabi area east of the gomondwane tar road in the southern district (fig. 1). two independent teams started the survey in the mnondozi-rietpan area (fig. 1) during july 1974, gradually working northwards until the eastern half of the central district had been covered . the western half was then done starting again in the south and working northwards towards the olifants river. the crocodile bridge-lower sabi area was surveyed during march and april, the entire operation being completed in august 1975. in the case of groups of between one and five lions, it was endeavoured to mark all the individuals, while in larger groups between 50% and 80% were ear tagged. to facilitate data analysis the central district was divided into four areas as indicated in fig. i. these were chosen on the basis that considerable differences occur in each area with respect to both vegetation and game composition, especially the western and eastern halves of the central district, while area (1) in the northwest differs from area (2) in that a large part of area (i) is dominated by colophospermum mopane shrubs and trees . zebra equus burchelli antiquorum and wildebeest connochaetes taurinus taurinus also migrate from area (1) to (2), spending the wet season in area (1) and the dry season in area (2) (smuts 1974). on the eastern half of the central district, zebra and wildebeest in area (3) are largely sedentary, while those in area (4) are migratory, undertaking 154 a north-south movement each year (smuts 1974). the crocodile bridgelower sabi area is actually a continuation of area (4) and the eastern lebombo flats, but lies on the southern side of the perennial sabi river where it forms a fairly closed unit for the game populations (smuts 1974). the boundaries of each area were fitted after the approximate ranges of all the prides encountered had been plotted onto a map, thereby grouping the prides associated with each particular area. since entire prides were seldom captured on one night, different groups captured at the same locality on successive nights or in the same general area, were grouped and regarded as being one pride. marked lions which were recaptured or resighted during the survey also assisted in the allocation of individuals or groups to a pride. age composition was analysed by dividing all lions into the following age classes. ages were assigned to classes by utlizing data published by schaller (972), bertram (1973), rudnai (973) and personal observations on age determination and growth in the lion (smuts, anderson and austin 1976) : (i) adult males (4+ years old) -lions with a well developed mane and a body mass in excess of 170 kg (except in unusual cases or emaciated specimens) . adult males can also be identified to some extent by the greyer colour of their coat, while scars are often more numerous on the face. behaviour at the carcass and particularly in the presence of other males also helps to distinguish adult males from large sub-adults. in in this group the longest cusps of the second upper (p5) and lower (p.) permanent premolars are worn (smuts, anderson and austin 1976). adult females (4+ years old) females with a body mass in excess of 110 kg (except in some old and/or emaciated specimens). only lionesses which had had one or more litters were regarded as adult. the prominently enlarged nipples distinguished these females from their nulliparous counterparts. tooth wear is similar to that for adult males . (ii) sub-adult males (2-4 years old) lions varying in mass from about 100 kg to almost 180 kilogram. here the mane is incompletely developed while the incisors and the second upper premolar (p5) start wearing at about four years of age (smuts, anderson and austin 1976). sub-adult females (2-4 years old) females varying in mass from about 75 kg to 120 kilogram. tooth wear is similar to that for sub-adult males . (iii) large cubs (1-2 years old) lions varying in mass from about 50 to 90 kilogram . between 12 and 14 months of age the permanent canines erupt (smuts, anderson and austin 1976). these lions, both male and female, were separated from the small cubs either by estimating or actually measuring body mass and by examining their dentition. (iv) small cubs (0-1 year old) lions with a body mass of less than 50 kilogram. using measurements of mass taken from some 89 lions during the study, together with those of five known-age specimens, the following average body masses were calculated for their corresponding age classes. 155 these measurements were used to calculate lion biomass for each part of the study area. in all instances average body mass includes stomach fill: adult males (181 kg; n = 14); adult females (126 kg; n = 25); sub-adult males (146 kg; n = 25) and sub-adult females (103 kg; n = 11). for cubs, rough growth curves were plotted and the following average body masses used for each age class; male cubs (1-2 years) (77 kg); female cubs 0-2 years) (60 kg); cubs (8-12 months) (44 kg); cubs (6-8 months) (30 kg); cubs (4-6 months) (22 kg); cubs (0-2 months) (5 kg). the recent history( 1902-1956) of the two lion populations was studied by utilizing the following information: annual reports compiled since 1903 by the respective wardens in the sabi game reserve and later the kruger national park; maps compiled by rangers during 1956 illustrating the distribution and sizes of prides in the two study areas. since the provision of artificial waterholes in the form of dams and windmills as well as the stabilization of natural waterholes by man appeared to be important in influencing the distribution and numerical status of game populations in the areas surveyed, these programs, initiated in 1933, were studied in relation to the colonisation of new areas by herbivore and predator populations (smuts 1975). results during the survey a total of 34 7 lions were captured and marked while 141 of the marked lions were also recaptured. of the 347, 319 were marked in the central district (45% of the population) while 62 were additionally destroyed as part of an experimental cropping operation (fig. 1). in the crocodile bridge-lower sabi area 28 were marked (42% of the population). all but three of the 60 prides in the central district contained marked lions while three out of six in the crocodile bridge-lower sabi area were marked. final survey results indicated that the central district had a minimum population of 708 lions (fig. 1), while the crocodile bridge-lower sabi area had 67 lions. the composition of individual prides is described in a separate memorandum (smuts 1975), while table 1 summarises relevant data on sex ratio, age composition, lion density and lion biomass per area. mean pride sizes and average number of adult females per pride and area are given in tables 2 and 3 respectively. modal pride size for the central district was nine. the maximum number of adult males in one pride and definitely associating together, was five for pride 26 in the central district. modal number of adult males per pride for the central district was two, the average being 2,1. all sex ratios differed significantly from the expected 1: 1 ratio with chi-square values increasing from 4,0 (area 1 all ages) to 22,0 (central district all ages). by the time the survey had been completed it was noted that the females and young of one pride were migratory (fig. 1), following the 156 -u-. ...., [ { [ area as per fig. i i 2 3 4 i and 2 (western boundary 3 and 4 (eastern boundarv crocodil~ bridgelowersabi 1&2&3&4 (central district ) adults 10: 1,79 25:43 10: 2, 19 24:50 10: 1,99 32:61 10 :2,49 41 :97 10: 1,99 49,94 10: 2,29 73: 158 10 :2,29 13:29 10: 2,19 122:251 table i population characteristic;, density and biomass for lions from the central district and the crocodile bridge-lower sabi are of the kruger national park sex ratio age composition of population (%) lion density per 100 km' size of lion adults area adults biomass suband suball adults sublarge small and subcubs (km' ) all and sub(kg/km' ) adults adults ages adults cubs cubs adults ages adults 10: 1,59 10: 1,79 10 :1,59 49,3 10,9 13,8 26, i 60,1 39,9 1310,0 10,5 6,3 6:9 31 :52 40:60 n=68 n=15 n=19 n=36 n=83 n=55 n=138 n=83 11,0 10:0 ,89 10: 1,7 9 10: 1,79 55,1 14,5 5,8 24 ,6 69,6 30,4 i 176,0 11,7 8,2 ii :9 35:59 35:61 n=76 n=20 n=8 n=34 n=96 n=42 n=138 n=96 12,5 10 :0,39 10: 1,49 10 :1,5 9 60 ,4 13,0 3,3 23,4 73,4 26,6 1276,0 12,1 8,9 15:5 47 :66 47 :70 n=20 n=5 n=36 n=113 n=1l3 n=41 n=154 n=113 13,9 10:0 ,59 10: 1,49 50,4 23,7 10,8 15,1 74, i 25,9 i 798,0 15,5 11,5 45:21 86:118 91: 130 n=140 n=66 n=30 n=42 n=206 n=72 n=278 n=206 17,7 10: 1,19 10: 1,79 10 :1,69 52,2 12,7 9,8 25,4 64,9 35,1 2486,0 11,1 7,2 17: 18 66 : 111 75: 121 n=144 n=35 n=27 n=70 n=179 n=97 n=276 n=179 11,7 10 :0,49 10: 1,49 53,9 19 ,9 8,1 18 , 1 73,9 26,2 3074,0 14, i 10,4 60:26 133: 184 138 :200 n=233 n=86 n=35 n=78 n=319 n=113 n=432 n=319 16,1 10 :0,69 10 :1,89 10: 1,59 62 , 7 17,9 16,4 3,0 80 ,6 19,4 463,0 14 ,5 11,7 5:3 18:32 24:37 n=42 n=12 n=11 n=2 n=54 n=13 n=67 n=54 18,2 !o :0,69 10 :1,59 10: 1,59 53,3 17, i 8,8 20,9 70,3 29,7 5560,0 12,7 9,0 77 :44 199:295 211 :319 n=377 n=121 n = 62 n=148 1l=498 n=210 n=708 n=498 14,1 --.table 2 mean pride sizes for lions from the central district and the crocodile bridgelower sabi area rifthe kruger national park number of number of mean pride range area as per fig. 1 prides lions size 1 13 138 10,6 4 to 18 2 12 138 11,5 9 to 18 3 12 154 12,8 7 to 21 4 23 278 12,1 7 to 21 1 and 2 (western boundary 25 276 11,0 4 to 18 3 and 4 (eastern boundary) 35 432 12,3 7 to 21 crocodile bridgelower sabi 6 67 11,2 6 to 16 1 and 2 and 3 and 4 (central district) 60 708 11,8 4 to 21 table 3 average number of adult females per pride for lions from the central district and crocodile bridge-lower sabi area rifthe kruger national park average number of number of area as per fig. 1 number of adult adult range prides females females 1 13 43 3,3 2 to 5 2 12 51 4,3 2to6 3 12 61 5,1 2 to 8 4 23 98 4,3 2 to 9 1 and 2 (western boundary) 25 94 3,8 2to6 3 and 4 (eastern boundary) 35 159 4,5 2to9 crocodile bridgelower sabi 6 29 4,8 3 to 8 1 and 2 and 3 and 4 (central district) 60 253 4,2 2to9 158 herds of zebra and wildebeest onto the lindanda plains, a distance of some 45 km . the males, by contrast, stayed behind in the mnondozi area. excluding the migratory group and a few individuals from pride 5 (fig. i), all prides have so far remained in the same general area. the movements of a few sub-adult males and sub-adult females were rather erratic and they moved distances of up to 20 km in one or two days. a number of marked lions have moved out of the knp (fig. i) and of these 10 are known to have been shot three sub-adult males and two adult females on cattle farms in mocambique, two adult females on a game farm near komatipoort, two adult females on a farm near crocodile bridge and one adult female on a cattle farm near klaseri . last mentioned had moved a distance of 70 km from her original site of capture at ngwenyeni dam in the central district. annual reports (stevensonhamilton 1903-1945) indicate that since the first warden entered the sabi game reserve (a section of the now kruger national park) in 1902, there have been marked increases in the numbers of most medium and large-sized herbivores. prior to this time, however, and in particular towards the end of the 19th century, game populations in the eastern transvaal lowveld were in very poor state. this condition could largely be ascribed to the appalling slaughter of game perpetrated by indigenous bantu tribes, pioneer white hunters as well as members of the armed forces during the anglo boer war of 1899-1902. in addition, the rinderpest epizootic of 1896-98 dealt a very decisive blow, severly reducing the numbers of most animals, but in particular of species such as the buffalo syncerus coffer, kudu tragelaphus strepsiceros and eland taurotragus oryx (stevenson-hamilton 1925). during 1903 col. stevenson-hamilton estimated that there were only a few lions in the reserve. however, from that year onwards game animals were afforded considerable protection from hunters, with the result that populations gradually increased. in 1925 the lion population for the central district was estimated at 250 and by 1956 the estimate was 424. the history of the artificial water for game program (smuts 1975) indicated that between 1933 and 1974 a total of 25 dams were built and 62 windmills erected in the central district. over the same period four dams were built and two windmills erected in the crocodile bridgelower sabi area. although the early lion surveys were carried out less intensively than the present ones, it could be established that at least 12 new prides (n = 159 lions) had been formed since 1956 in the central district. these were prides 7, 8, 10, 20, 21, 22, 23, 26,32, 49, '50 and 53 (fig. i). eight of these (prides 8, 21, 22, 23, 26, 49, 50 and 53) could be associated directly with the provision of new waterholes or the stabilization of existing ones. in the crocodile bridge-lower sabi area two prides (n = 27 lions) were formed after 1956 in the vicinity of mhlanganzwane and mpanamana dams which were completed during 1956 and 1957 respectively. 159 discussion general although six of the 66 prides in the two study areas did not contain marked individuals, only three of these were not seen by the capture teams. these prides were, however, located audibly and their approximate sizes obtained from the respective game rangers living in each area. the estimated activity area for each pride was based on a limited number of resightings, information obtained from game rangers and on the relative distribution of surrounding prides. these areas overlap considerably and fig. 1 should be regarded as a schematic rather than a true representation of each pride's activity area. although the survey is regarded as a reliable total count, the populations of 708 and 67 lions for each area are minimum figures since the possibility exists that a few small groups and females with small cubs could have been overlooked. it is not possible to estimate the magnitude of the error although this is believed to be insignificant for the following reasons: numerous ground observations have failed to report on large groups without marked individuals; in areas where the survey overlapped with itself and in one area (bangu-balule) which was re-surveyed, new results were in very close agreement with those of the initial surveys . the greatest source of error is due to "edge-effect" since certain of the peripheral prides may regularly move further out of the knp than is presently known. similarly other groups may also move into the park more frequently than is realized. movements towards the east will now, however, be almost impossible unless the lions scale the new 2,4 m high game proof fence, or leave via a few deep ravines which could not be adequately fenced. on the west lions can move fairly freely to and from the knp by passing underneath or between strands of barbed wire of the game deterrent fence. all lions which were destroyed outside the eastern border of the park left before the new fence was erected. population characteristics, density and biomass sex ratios (table 1) indicate that in all areas there is a preponderance offemales with the most distorted adult sex ratio being found in area (4) 00: 2,4(9). areas with the most distorted adult sex ratios also had the lowest percentages of cubs. this was particularly so in area (4) and in the crocodile bridge-lower sabi area. since a distorted sex ratio indicates a differential sex specific mortality rate, data in table 1 indicate that lions on the eastern half of the central district (areas (3) and (4)) as well as those in the crocodile bridge-lower sabi area ( a similar habitat to area (4)), have experienced the highest mortality rates. because areas with the highest density of adult lions (table 1) have the most distorted adult sex ratios, it would appear that adult sex ratio is density dependent and probably the result of intra-specific competition. unfortunately insufficient cubs were sexed (2000 :269 9) to be able to make reliable conclu160 sions on the age at which sex ratio starts to become distorted. in captivity, however, sex ratio at birth is parity (brand 1963 1316'6': 130 « «). schaller (1972) also found that the sex ratio in cubs under one year of age was equal. it is very likely therefore that a differential mortality rate starts operating at about two years of age, when lions become independent (rudnai 1973). it may, however, start even earlier since signs of growing independence and the beginnings of the male pattern of behaviour begin to show at about nine months of age (rudnai op cit.). when studying sub-adult sex ratios (table 1) and principally those for the western and eastern halves of the central district the above type of pattern does not seem to apply. here, for example, the western boundary area (areas (1) and (2)) had a sub-adult sex ratio of 16': 1,1 « (n = 35 lions) while that for the eastern boundary area (areas (3) and (4)) was 16':0,4« (n = 86 lions). these differences seem to indicate that there may not be a gradual reduction in the number of young males relative to females after about nine months of age, but rather that there is a sudden reduction at a certain age and under specific conditions. assuming that the distorted sub-adult sex ratio for the eastern boundary area is due to an elevated mortality rate of sub-adult females relative to males, it is possible that the males will show a similar high mortality when they start competing with adult males and that this will continue until the sex ratio is eventually distorted in favour of adult females as was found in all the areas studied (table 1). the possibility that sub-adult lions disperse at different rates to and from the western and eastern halves of the central district (due to conditions outside the park) and the small size of each sample, however, precludes further speculation. sex ratios in the present study were more distorted than those found by schaller (1972) for the serengeti and particularly so for the adult plus sub-adult group where he obtained figures of between 16': 1,1 « and 16':1,3«« (16':1,4«« and 16':1,8«« in the present study table 1). here the lower lion density for the serengeti ecological unit (± 7,8 lions/ 100 sq km schaller 1972) when compared with the central district (12,7 lions/l 00 sq km) supports the previous idea that sex ratio and therefore mortality rate is density dependent. the age composition of the lions in the various areas also shows important differences (table 1). here the proportion of cubs decreases as density increases, the lowest percentage being found in area (4) and the crocodile bridgelower sabi area. area (1) had the highest percentage of cubs. these differences indicate that lions in the western half of the central district and particularly in area (1), were increasing relative to those in the savannah areas on the east. on a long-term basis; however, lions on the east are more successful than those on the west as is indicated by data on density and biomass (table 1), mean pride sizes (table 2) and the number of adult females per pride (table 3). the fact that area (1) had 14% more cubs than area (4) is striking since both areas were surveyed during the month of july although in two suc161 cessive years. the low percentage of cubs in area (4) and the crocodile bridge-lower sabi area, together with the emigration of lions from these areas, indicate that food supply is possibly becoming a limiting factor. six of the nine lions which have left the knp via the eastern boundary have been adult females. since it was not always possible to ascertain the exact social status of all lions allocated to a particular pride, except when the entire pride was captured together on one night, pride sizes are slightly suspect. in certain cases resightings of marked lions and those recaptured did, however, help in the allocation of individuals to a particular pride. for these reasons the word "pride" has been used in preference to "group" although the limitations are realized. mean pride sizes (table 2) indicate that the prides on the eastern half of the central district are larger than those on the west, the largest mean being for area (3) and the smallest for area (1). the two largest prides (n = 21 lions each) were also recorded on the east (fig. 1). schaller (1972) obtained a mean pride size of 15,1 (range = 4 to 37) for 14 prides whose size was determined accurately i.e., prides which were known to have retained their identity for at least one year. data collected in both study areas (tables 1 and 2) indicate that mean pride size is not density dependent. it is quite likely therefore, that food availability in the long term, is the regulating force. this is supported by the higher density of prey animals on the eastern half of the central district than on the west (smuts 1975). since the number of cubs in a pride may cause its size to change considerably from year to year, bertram (1973) found that the most useful measure of pride size was the number of adult females. using this criterion it can be seen (table 3) that the largest mean pride size was recorded for area (3) on the eastern half of the central district, followed by the crocodile bridge-lower sabi area. the largest number of adult females per pride came from areas (3), (4) and crocodile bridge-lower sabi. these data indicate that lions in the eastern areas have a greater reproductive potential than those on the west. when comparing mean pride sizes using adult females only, area (3) plus (4) has a reproductive potential 15,6% greater than area (1) plus (2). data presented in table 1 indicate that by retaining the present population of 251 adult females in the central district, but increasing cub percentages in areas (2), (3) and (4) to that of area (1), where 40% of the population are under two years of age, would cause the overall lion population in the central district to rise from 708 to 819 . compared to the nairobi national park where 60% of the lion population is under two years of age (rudnai 1973), it is clear that the present population is below its potential maximum. the high percentage of adults and subadults in area (4), the distorted sex ratio of adults together with the fact that two marked males have been killed in fights and a number of adults shot outside the knp in a relatively short time, indicate that in some areas at least, the population is on the decline. in conclusion therefore, 162 most evidence, scanty as it may be, points to a lion population which has already reached its asymptote. this is supported by the results obtained from a series of intensive vehicle counts conducted since 1968. these . counts (smuts 1975) indicate that the lion population in the central district reached its peak between 1971 and 1973 when there were some 825 lions. history of the lion population in the central district bearing in mind that regulation of a lion population is influenced both by food and sociai behaviour (bertram 1973 and 1975) and that each pride occupies a fairly exclusive territory, the boundaries of which change little in the long term (bertram 1973), it seems safe to say that a vast and lasting increase (50% or more) in the size of a lion population in a large area, would require the settlement of unoccupied areas by new prides, with or without the increase in size of existing prides. this is apparently what has occurred in the central district where in all probability the lion population has more than doubled during the past 50 years. by studying the maps on pride distribution and sizes compiled during the 1956 lion survey in the central district and crocodile bridgelower sabi area and comparing these with fig. 1, it was possible to correlate upward trends in the lion population with the provision of artificial waterholes and with stabilization of permanent drinking spots. the distribution of prides indicated that new prides had colonised areas where water had been provided and where sedentary prey species had consequently settled. it was concluded (smuts 1975) that the increase in the lion population since 1925 had taken place gradually as prey populations increased and despite moderate cropping of lions up until 1958. the most dramatic increase, however, probably took place after 1933 when the first borehole was sunk and as the artificial water for game program gradually gained momentum, thereby opening up more and more areas for herbivore and carnivore colonisation. acknowledgements the national parks board of trustees are thanked for facilities provided and for permission to publish this paper. mr i. j. whyte and mr t. w. dearlove are thanked for their enthusiastic field assistance as are messrs. a. j. espag, j. steyn, j. clarke, j. p. ackerman, b. p. lamprechts, j. h. de kock, p.chauke, l. mangane, m. singwane, j. sithole and p. nkuna. mr s. c. j. joubert is thanked for advice and indirect assistance received during the study. 163 references bertram, b. c. r. 1973. lion population regulation e. afr. wild!.}., 11 :215-225. bertram, b. c. r. 1975. the social system of lions. scientific american, april: 54-65. brand, d. 1963. records of mammals bred in the national zoological gardens of south africa during the period of 1908 to 1960. proc. zool . soc. lond., 140:617-59. kruuk, h. 1972. the spotted hyena. a study of predation and social behaviour. ed . g. b. schaller, chicago and london: the univ., of chicago press. rodgers, w. a. 1974. the lion (panthera leo, linn) population of the eastern selous game reserve. e. afr. wildl.}.} 12:313-317. rudnai, judith. 1973. reproductive biology of lions (panthera leo massaica neumann) in nairobi national park. e. afr. wildl. ]., 11 :241253. schaller, g. b. 1972. the serengeti lion. a study of predator prey relations. ed. g. b. schaller. chicago and london: the univ. of chicago press. smuts, g. l. 1974. game movements in the kruger national park and their relationship to the segregation of sub-populations and the allocation of culling compartments.}. 5th. afr. wildl. mgmt ass., 4: 51-58. smuts, g. l. 1975. predator prey relationships in the central district of the kruger national park with emphasis on wildebeest and zebra populations. memorandum, national parks board of trustees, pretoria. 74 pp. smuts, g . l.,j. l. anderson andj. c. austin. 1976. age determination and growth in the african lion (panthera leo) (in prep.). smuts, g. l., i.j. whyte and t. w. dearlove. 1976. a mass capture technique for lions. e. afr. wildl.}., (in press). stevenson-hamilton, j. 1903-1945. annual reports of the warden of the kruger national park, national parks board of trustees, pretoria. 164 page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 article information authors: bruce r. ellender1,2,3 olaf l.f. weyl1,3 affiliations: 1south african institute for aquatic biodiversity, grahamstown, south africa 2department of ichthyology and fisheries science, rhodes university, south africa 3centre for invasion biology, south african institute for aquatic biodiversity, grahamstown, south africa correspondence to: bruce ellender email: bru.ellender@gmail.com postal address: private bag 1015, grahamstown 6140, south africa dates: received: 04 nov. 2014 accepted: 27 jan. 2015 published: 22 may 2015 how to cite this article: ellender, b.r. & weyl, o.l.f., 2015, ‘resilience of imperilled headwater stream fish to an unpredictable high-magnitude flood’, koedoe 57(1), art. #1258, 8 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1258 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. resilience of imperilled headwater stream fish to an unpredictable high-magnitude flood in this original research... open access • abstract • introduction • research method and design    • study site    • field surveys    • data analysis       • rainfall and flow       • fish population dynamics • results    • rainfall and flow       • fish responses • discussion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ headwater stream fish communities are increasingly becoming isolated in headwater refugia that are often cut off from other metapopulations within a river network as a result of non-native fish invasions, pollution, water abstraction and habitat degradation downstream. this range restriction and isolation therefore makes them vulnerable to extinction. understanding threats to isolated fish populations is consequently important for their conservation. following a base-flow survey, a high-magnitude flood (peak flow of 1245 m−3s−1) provided an opportunity to investigate the response of endangered eastern cape redfin pseudobarbus afer populations to a natural disturbance in the waterkloof and fernkloof streams, two relatively pristine headwater tributaries of the swartkops river system within the groendal wilderness area, eastern cape province, south africa. pseudobarbus afer had limited distributions, occupying 3 km in both the fernkloof and waterkloof streams. fish population assessments before and after the flood event indicated that there were no longitudinal trends in p. afer abundance before or after the flood, but overall abundance post-flooding in the fernkloof stream was higher. there were no noticeable changes in p. afer size structure preand post-flood. pseudobarbus afer showed resilience to a major flooding event most likely related to evolution in river systems characterised by environmental stochasticity. conservation implications: this research provides insight into the population level responses of native headwater stream fishes to unpredictable natural disturbance. of particular relevance is information on their ability to withstand natural disturbances, which provides novel information essential for their conservation and management especially as these fishes are already impacted by multiple anthropogenic stressors. introduction top ↑ headwater stream fishes in south africa are increasingly being isolated in small fragmented headwater refuges as a result of competition with and predation by non-native fishes, downstream water abstraction, pollution and habitat degradation (marr et al. 2010; tweddle et al. 2009; weyl et al. 2014). protected areas play an important role in conserving headwater fishes by preserving habitats and preventing non-native fish introductions (russell 2011). it is increasingly recognised, however, that the persistence of fishes in these environments may be dependent on dispersal between complementary habitats for reproduction, feeding, rearing and disturbance avoidance (franssen et al. 2006; labbe & fausch 2000; schlosser & angermeier 1995). isolation may therefore compromise their long-term persistence (fausch et al. 2009). understanding the vulnerability of isolated refuge populations to natural catastrophic events such as flooding is therefore important for conservation planning. headwater streams are considered particularly susceptible to floods because they have small catchments and are easily influenced by relatively minor changes in local conditions (meyer et al. 2007). unpredictable, infrequent and catastrophic floods can result in slope failures, bank erosion, substrate scouring, and loss of habitat and biota (resh et al. 1988). in fish populations, floods can result in downstream displacements (cambray 1994), reduced abundance (magalhaes, schlosser & collares-pereira 2003; matthews 1986; nislow et al. 2002; pires et al. 2008), recruitment failure (letcher & terrick 1998) and alteration in community composition (matthews 1986; nislow et al. 2002). despite the documented impacts of floods on stream fish communities (letcher & terrick 1998; matthews 1986; nislow et al. 2002; pires et al. 2008; resh et al. 1988), fishes also display long-term resilience and have been shown to return to equilibrium fairly rapidly following catastrophic flood events (dolloff, flebbe & owen 1994; matthews 1986). under natural conditions, fishes are able to recolonise disturbed stream reaches from metapopulations in undisturbed refuge habitats (dolloff et al. 1994; matthews 1986; medeiros & maltchik 2001). isolation is a concern for freshwater fish conservation in south africa, where many headwater stream fishes are endemic to individual river systems and populations are increasingly fragmented by man-made (dams, weirs) or biological (invasive fish predation) barriers (ellender, weyl & swartz 2011; tweddle et al. 2009). one such species is the mandela lineage of the endangered eastern cape redfin minnow, pseudobarbus afer (peters, 1864) (swartz & impson 2007). its natural distribution is limited to only three river systems (sundays, swartkops and baakens) in the eastern cape province of south africa (swartz, skelton & bloomer 2007). populations are heavily impacted by non-native fish predation (ellender et al. 2011) and only persist in isolated headwater refugia (ellender 2013). fortunately, a considerable portion of these headwater refugia are located in two protected areas, the addo elephant national park (sundays river system) and the groendal wilderness area (swartkops river system). whilst these protected areas provide protection from non-native fishes and habitat destruction, the impact of natural catastrophic floods on these isolated populations are unknown. this article provides insight into the population level responses of p. afer to an unpredictable high-magnitude flood event by comparing the distributions, relative abundance and population structure of p. afer in two headwater streams before and after an unpredictable high-magnitude flood. it was hypothesised that the flood would (1) result in displacement and mortality of headwater fishes, resulting in changes in abundance and distribution in headwater streams and (2) that juvenile life history stages of p. afer would be more vulnerable to flood disturbance. research method and design top ↑ study site the study was undertaken within the groendal wilderness area on two second-order, episodic headwater tributaries (fernkloof stream, waterkloof stream) of the kwa-zunga river, a major tributary of the swartkops river system, eastern cape, south africa (figure 1). as a result of their entire catchments being situated within the groendal wilderness area (a protected area proclaimed for the protection of indigenous forest and water resources), the fernkloof and waterkloof streams are relatively pristine with undammed catchments and intact riparian zones characterised by closed canopy (fernkloof: 46% ± 23% canopy cover; waterkloof: 33% ± 24% canopy cover) tropical coastal forest (acocks 1975) and mesic grassy fynbos (campbell 1985). the character of the two streams is typical for episodic headwater streams in the eastern cape. both streams have steep, high profile banks within narrow valleys. in-stream habitat was characterised by isolated pools (fernkloof [mean ± s.d.] length: 10.8 m ± 3.2 m; width: 3.6 m ± 0.9 m; depth: 0.7 m ± 0.4 m; waterkloof [mean ± s.d.] length: 11.9 m ± 3.7 m; width: 5.5 m ± 4.8 m; depth: 0.7 m ± 0.6 m) connected by subsurface flows, with bedrock, large unconsolidated boulder, cobble and pebble substrates. surface flow only connects pools for a few days after heavy sustained rainfall. figure 1: the situation and sites sampled on the fernkloof and waterkloof streams, headwater tributaries of the swartkops river system within the groendal wilderness area, eastern cape, south africa. the position of the wincanton gauging weir (m1h004) on the elands river and the weather station (m1e001) at groendal dam are also noted. field surveys ten sites on each stream were sampled before (may 2010) and two months after (august 2012) a major flooding event (june 2012) by electrofishing using a samus 725g backpack electrofisher, attached to a 12 v battery with settings standardised at the duration of 0.3 ms and a frequency of 80 hz. because flood scouring and filling changed the position of pools within the two streams in the post-flood survey, the nearest pool to the pre-flood sample site was electrofished. single-pass electrofishing was conducted from the downstream side (tail) of the pool in an upstream direction, covering the entire length of each pool. upon completion of the pass, fish were identified to species level, measured, counted and released. electrofishing data were used as an index of relative abundance and catch per unit effort (cpue) was expressed as fish m−3 using volume estimates from the habitat data collected for each sampled pool. block nets were not used as pools were primarily isolated on the surface because of the episodic nature of the sampled streams. at each sampling site, temperature, conductivity and ph were measured using a hanna hi98129 combo ph and electrical conductivity meter (hanna instruments inc., woonsocket, usa). turbidity (ntu) was measured using a hanna hi 98703 turbidimeter (hanna instruments inc., woonsocket, usa). to estimate pool volume and habitat diversity, the length (± 0.1 m) of the pool was measured, followed by equally spaced width measurements (± 0.1 m). on each width transect, three depths (± 0.1 m) were measured; the outer two were 0.2 m each from the left-hand and right-hand stream bank and the third measurement taken midstream. at each depth measurement, the habitat type was recorded. canopy cover was estimated as a percentage of total cover and bankside vegetation type was recorded. data analysis rainfall and flow long-term hydrological and meteorological data were obtained from the department of water affairs hydrology section (dwaf 2012); the locations of the rainfall and gauging stations are presented in figure 1. these data were used to illustrate rainfall, flow variability and the magnitude of the flood in the headwaters of the swartkops river system. because of an absence of long-term flow data for the fernkloof and waterkloof streams, flow data were obtained from a representative swartkops river system headwater tributary, the gauging weir at wincanton on the elands river (m1h004) for the period 06 april 1965 – 05 january 2012. rainfall data were obtained from station m1e001 at groendal dam (period: 16 february 1950 – 11 january 2006) and supplemented with data from the nearby station m1e002 at uitenhage (period: 30 november 2006 – 05 january 2012). spearman rank order correlation was used to test for a relationship between rainfall and flow in the swartkops river system headwater streams at a significance of p < 0.05. the following criteria, outlined by resh et al. (1988), were used to classify the magnitude of the flood disturbance as: any relatively discrete event in time that is characterised by a frequency, intensity, and severity outside a predictable range, and that disrupts ecosystem, community, or population structure and changes resources or the physical environment. (p. 434) these criteria were applied to peak flow data for the swartkops river system to verify the magnitude of the flood in june 2012. fish population dynamics pre-flood and post-flood comparisons were limited to a single fish species, p. afer, as they were ubiquitously distributed from the limit of fish distributions (p. afer distribution: waterkloof = 3 km; fernkloof = 3 km) in the upper reaches to the confluence with the mainstem kwa-zunga river before the flood, allowing longitudinal analyses. to investigate the impact of a major flooding event on p. afer in headwater streams, pre-flood and post-flood electrofishing catch per unit effort (cpue, fish m−3) estimates were compared within the two streams using a main-effects analysis of variance (anova) at a significance level of p ≤ 0.05. specifically, cpue was tested for longitudinal trends (upper versus lower reaches), as well as flooding impact (pre-flood versus post-flood) and an interaction between the two factors. because of complications resulting from differences in sampling months, with the pre-flood survey occurring at the end of the spawning season and the post-flood survey at the beginning of the spawning season (cambray 1994), p. afer juveniles (< 40 mm) and adults (> 40 mm) were analysed separately. to test the vulnerability of the two p. afer size classes to the flooding event, the frequency of occurrence (number of sites per stream reach where each size class was present expressed as a percentage of all sites in that reach) by stream reach (upper and lower reaches) was tested for significance by reach and pre-flooding and post-flooding using a chi-squared test of independence. all analyses were undertaken using ms excel 2007, microsoft® and statistica 10.0, statsoft®. results top ↑ rainfall and flow mean annual rainfall (mean ± s.d.) for the period 16 february 1950 – 05 january 2012 was 620.7 ± 212.3 mm yr−1. rainfall was variable, falling in an unpredictable, erratic pattern (figure 2). there was a highly significant correlation between rainfall (station m1e001 and m1e002) and peak flow (gauging weir m1h004) (spearman r = 0.389; z = 9.45; p = 0.000), highlighting the system's episodic nature. peak flows between april 1965 and may 2012 were generally low (mean ± s.d.: 30.3 m−3s−1 ± 202.3 m−3s−1) and were predominantly < 25 m−3s−1 (89%), with a further 8.7% < 500 m−3s−1. during the 47 years on record, there were only four peak flows exceeding 1000 m−3s−1 (figure 2). the flooding event with peak flows of 1245 m−3s−1 (a measure of instantaneous flow at peak discharge) in june 2012 was of a frequency, intensity and severity outside of the predictable range and resulted in changes to the physical environment, which, according to resh et al. (1988), classified the flood as a major disturbance. figure 2: monthly peak flows (black) at the wincanton gauging weir on the elands river and total monthly rainfall (grey) at groendal dam (16 february – 11 january 2006) and at uitenhage (30 november 2006 – 05 january 2012) illustrating rainfall and flow variability in the headwater tributaries of the swartkops river system (flow data are missing for the period april 1981 – september 1986). fish responses the distribution and abundance of fishes from the fernkloof and waterkloof streams pre-flood and post-flood are summarised in table 1. four species were recorded from the fernkloof and waterkloof streams, of which three were shared (native: p. afer and cape kurper sandelia capensis [cuvier, 1831]; non-native: african sharptooth catfish clarias gariepinus [burchell, 1822]) and an additional non-native, banded tilapia tilapia sparrmanii a. smith 1840 was recorded in the fernkloof stream only. pseudobarbus afer distributions were limited to a 3 km stretch in both the fernkloof and waterkloof streams. table 1: the stream zones (lower reaches, upper reaches), sites (10 sites), distribution, catch per unit effort (fish m−3), relative abundance (%) of fishes by site (%), including water-quality variables from site 1 in the lower reaches to site 10 in the upper reaches of the fernkloof and waterkloof streams, headwater tributaries of the kwa-zunga river within the groendal wilderness area. there were no significant pre-flood and post-flood longitudinal or interactive effects (stream reach: upper and lower reaches; disturbance: pre-flood and post-flood) in cpue for p. afer in the waterkloof stream (pre-flood mean ± s.e.: 1.24 ± 0.33 fish m−3; post-flood mean ± s.e.: 2.43 ± 0.40 fish m−3) (table 2). there was, however, a significant increase in post-flood p. afer cpue in the fernkloof stream (pre-flood mean ± s.e.: 1.33 ± 0.49 fish m−3; post-flood mean ± s.e.: 6.53 ± 2.12 fish m−3) (figure 3; table 2). the frequency of occurrence and length distributions for p. afer pre-flood and post-flood are summarised in table 3 and figure 4. table 2: a summary of pre-flood and post-flood longitudinal (upper and lower stream reaches) and disturbance (pre-flood and post-flood) main-effects anova results for pseudobarbus afer cpue (fish m−3) from the waterkloof and fernkloof streams, headwater tributaries of the kwa-zunga river within the groendal wilderness area. figure 3: pre-flood and post-flood longitudinal abundance trends for pseudobarbus afer catch per unit effort (fish m−3) from the waterkloof and fernkloof streams, headwater tributaries of the kwa-zunga river within the groendal wilderness area, eastern cape, south africa. table 3: the frequency of occurrence (% of sites containing pseudobarbus afer) of juvenile (< 40 mm) and adult (> 40 mm) pseudobarbus afer pre-flood and post-flood separated by stream reach (lower and upper) for the fernkloof and waterkloof streams headwater tributaries of the kwa-zunga river within the groendal wilderness area. figure 4: pre-flood and post-flood length frequency distribution for pseudobarbus afer from the upper and lower reaches of the fernkloof (a = lower; c = upper) and waterkloof (b = lower; c = upper) streams, headwater tributaries of the kwa-zunga river within the groendal wilderness area, eastern cape, south africa. the frequency of occurrence for juvenile (< 40 mm) and adult p. afer was independent of stream reach and were subsequently grouped by stream (table 3). the frequency of occurrence of juvenile and adult p. afer was independent of flooding for the waterkloof (pre-flood versus post-flood: χ2 = 0.027, df = 1, p = 0.87) and fernkloof streams (pre-flood versus post-flood: χ2 = 0.044, df = 1, p = 0.83) (table 3). in the waterkloof stream, p. afer size distributions were similar between stream reaches and pre-flooding and post-flooding (pre-flood lower mean, range: 43.7, 22 mm – 74 mm fork length (fl); pre-flood upper mean, range: 40.7, 16 mm – 71 mm fl; post-flood lower mean, range: 42.8, 21 mm – 67 mm fl; post-flood upper mean, range: 47.2, 28 mm – 81 mm fl) (figure 4). in the fernkloof stream, the mean length of p. afer was larger in the lower reaches but similar before and after the flood (pre-flood lower mean, range: 53.0, 11 mm – 96 mm fl; pre-flood upper mean, range: 43.0, 24 mm – 72 mm fl; post-flood lower mean, range: 52.0, 30 mm – 87 mm fl; post-flood upper mean, range: 46.3, 21 mm – 100 mm fl). discussion top ↑ flood-related disturbances typically elicit a variety of responses from fishes in stream environments, the nature of which are related to the predictability, magnitude/intensity and duration of the event (lytle & poff 2004) as well as the specific adaptations of the fishes to the abiotic conditions of that environment (franssen et al. 2006; lytle & poff 2004). in the fernkloof and waterkloof streams, p. afer demonstrated resilience to a major flooding event. in both small streams, there were no longitudinal trends in p. afer abundance before or after the flood, but overall abundance post-flooding in the fernkloof stream was higher. this may have been the result of the inability to sample the exact sites before and after the flood as some pools were filled up by flood deposition. the nearest pool to the pre-flood site was sampled, which may have resulted in sampling a refuge pool where post-flood abundances were high as a result of collection of fishes from that stream segment. good recruitment during the 2010/2011 spawning season might also have led to the observed increase in abundance as during this period rainfall was higher than the preceding two spawning seasons. increased recruitment during periods of good flow was recorded by franssen et al. (2006) for intermittent prairie stream fishes in north-eastern kansas, usa. however, the p. afer length frequency data from the waterkloof and fernkloof streams do not support this, as size structuring before and after the flood was similar. the vulnerability of stream fishes to flood disturbance is not always clearly evident from overall abundance trends, but in some cases certain life history stages of particular species may be more vulnerable to disturbance than others (gasith & resh 1999; letcher & terrick 1998; lytle & poff 2004). in new england streams in the usa, a massive localised flood caused an age-0 year-class failure in brook trout salvelinus fontinalis (mitchill, 1815) and brown trout salmo trutta linnaeus 1758 and a large decrease in abundance of atlantic salmon salmo salar linnaeus 1758 (letcher & terrick 1998). similar results have been recorded for other species after catastrophic flood events (dolloff et al. 1994; harvey 1987; matthews 1986; nislow et al. 2002). this was not the case for p. afer as there were no significant changes in the occurrence of either juvenile or adult p. afer following the flood. both life history stages were distributed throughout both streams from their confluence with the mainstream kwa-zunga in the lower reaches, to the limit of p. afer distribution in the upper reaches. the susceptibility of juvenile fishes to flood displacement and mortality has been documented to decrease rapidly with a small increase in length (harvey 1987) and p. afer may show similar trends in changing vulnerability with size. in the nearby wit river, a headwater tributary of the gamtoos river system, cambray (1994) reported that late free embryos and early larvae drift out of the areas where they were spawned. during the current study, the flooding event took place 3–5 months after the peak spawning season of p. afer (cambray 1994), perhaps allowing young-of-the-year fish time to grow out of the most vulnerable life history stages. distribution and movement of non-native species indicated species-specific and stream-specific flood response. for example, after the same flood, ellender, woodford and weyl (2015) noted that the flood event appeared to have flushed c. gariepinus out of the fernkloof stream but facilitated its penetration into the waterkloof stream. temporal longitudinal movement was also noted in the blindekloof stream, another kwa-zunga river tributary, during a period of prolonged flow (± 5 months) following the june 2012 flood, when c. gariepinus penetrated 5 km upstream from the mainstem (ellender et al. 2015). in conclusion, the observed resilience of p. afer populations to a major flooding event is most likely related to their evolution in river systems characterised by environmental stochasticity (dolloff et al. 1994; magalhaes et al. 2003; pires et al. 2008). the long-term persistence of p. afer in relatively pristine isolated headwater refugia may therefore not be threatened by flood disturbances of the observed magnitude and intensity. the biggest long-term threat may be the flood-facilitated penetration of headwater refugia by non-native species. acknowledgements top ↑ b.r.e. received financial support from: the south africa–netherlands research programme on alternatives in development (sanpad project 10/06), the national research foundation of south africa (nrf), rhodes university, the dst/nrf centre of excellence for invasion biology and the water research commission (wrc project no. k5/1957/4, k5/2039 and k5/2261). eastern cape parks board and staff of the groendal wilderness area are thanked for access and for their logistical assistance. geraldine taylor and patrick gourley are thanked for field assistance. research was conducted following saiab animal ethics guidelines. research permits were issues by the eastern cape department of economic development and environmental affairs (dedea) and eastern cape parks. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions b.r.e. (south african institute for aquatic biodiversity) was responsible for study conceptualisation, data collection, analyses, writing and editing the manuscript. o.l.f.w. (south african institute for aquatic biodiversity) contributed to conceptualising the study, writing and editing the manuscript. references top ↑ acocks, j.p.h., 1975, ‘veld types of south africa’, memoirs of the botanical survey of south africa 40, 1–128. cambray, j.a., 1994, ‘the comparative reproductive styles of two closely related african minnows (pseudobarbus afer and p. asper) inhabiting two different sections of the gamtoos river system’, environmental biology of fishes 41, 247–268. http://dx.doi.org/10.1007/bf02197848 campbell, b.m., 1985, ‘a classification of the mountain vegetation of the fynbos biome’, memoirs of the botanical survey of south africa 50, 1–21. dolloff, c.a., flebbe, p.a. & owen, m.d., 1994, ‘fish habitat and fish populations in a southern appalachian watershed before and after hurricane hugo’, transactions of the american fisheries society 123, 668–678. http://dx.doi.org/10.1577/1548-8659(1994)123<0668:fhafpi>2.3.co;2 dwaf, 2012, department of water affairs and forestry river station database, viewed 28 june 2012, from http://www.dwaf.gov.za/hydrology/ ellender, b.r., 2013, ‘ecological consequences of non-native fish invasions in eastern cape headwater streams’, phd thesis, department of ichthyology & fisheries science, rhodes university. ellender, b.r., weyl, o.l.f. & swartz, e.r., 2011, ‘invasion of a headwater stream by non-native fishes in the swartkops river system, south africa’, african zoology 46, 39–46. http://dx.doi.org/10.3377/004.046.0116 ellender, b.r., woodford, d.j. & weyl, o.l.f., 2015, ‘the invasibility of small headwater streams by an emerging invader, clarias gariepinus’, biological invasions 17, 57–61. http://dx.doi.org/10.1007/s10530-014-0744-8 fausch, k.d., rieman, b.e., dunham, j.b., young, m.k. & peterson, d.p., 2009, ‘invasion versus isolation: trade-offs in managing native salmonids with barriers to upstream movement’, conservation biology 23, 859–870. http://dx.doi.org/10.1111/j.1365-2427.2006.01640.x franssen, n.r., gido, k.b., guy, c.s., tripe, j.a., shrank, s.j. & strakosh, t.r. et al., 2006, ‘effects of floods on fish assemblages in an intermittent prairie stream’, freshwater biology, 51, 2072–2086. http://dx.doi.org/10.1111/j.1365-2427.2006.01640.x gasith, a. & resh, v.h., 1999, ‘streams in mediterranean climate regions: abiotic influences and biotic responses to predictable seasonal events’, annual review of ecology and systematics 30, 51–81. http://www.jstor.org/stable/221679. http://dx.doi.org/10.1146/annurev.ecolsys.30.1.51 harvey, b.c., 1987, ‘susceptibility of young-of-the-year fishes to downstream displacement by flooding’, transactions of the american fisheries society 116, 851–855. http://dx.doi.org/10.1577/1548-8659(1987)116<851:soyftd>2.0.co;2 labbe, t.r. & fausch, k.d., 2000, ‘dynamics of intermittent stream habitat regulate persistence of a threatened fish at multiple scales’, ecological applications 10, 1774–1791. http://dx.doi.org/10.1890/1051-0761(2000)010[1774:doishr]2.0.co;2 letcher, b.h. & terrick, t.d., 1998, ‘maturation of male age-0 atlantic salmon following a massive, localized flood’, journal of fish biology 53, 1243–1252. http://dx.doi.org/10.1111/j.1095-8649.1998.tb00245.x lytle, d.a. & poff, n.l., 2004, ‘adaptation to natural flow regimes’, trends in ecology & evolution 19, 94–100. http://dx.doi.org/10.1016/j.tree.2003.10.002 magalhaes, m.f., schlosser, i.j. & collares-pereira, m.j., 2003, ‘the role of life history in the relationship between population dynamics and environmental variability in two mediterranean stream fishes’, journal of fish biology 63, 300–317. http://dx.doi.org/10.1046/j.1095-8649.2003.00148.x marr, s.m., marchetti, m.p., olden, j.d., garcía-berthou, e., morgan, d.l., arismendi, i. et al., 2010, ‘freshwater fish introductions in mediterranean-climate regions: are there commonalities in the conservation problem?’, diversity and distributions, 16, 606–619. http://dx.doi.org/10.1111/j.1472-4642.2010.00669.x matthews, w.j., 1986, ‘fish faunal structure in an ozark stream: stability, persistence and a catastrophic flood’, copeia 2, 388–397. http://www.jstor.org/stable/1444997 medeiros, e.s.f. & maltchik, l., 2001, ‘fish assemblage stability in an intermittently flowing stream from the brazilian semiarid region’, austral ecology 26, 156–164. http://dx.doi.org/10.1046/j.1442-9993.2001.01099.x meyer, j.l., strayer, d.l., wallace, j.b., eggert, s.l., helfman, g.s. & leonard, n.e., 2007, ‘the contribution of headwater streams to biodiversity in river networks’, journal of the american water resources association 43, 86–103. http://dx.doi.org/10.1111/j.1752-1688.2007.00008.x nislow, k.h., magilligan, f.j., folt, c.l. & kennedy, b.p., 2002, ‘within-basin variation in the short-term effects of a major flood on stream fishes and invertebrates’, journal of freshwater ecology 17, 305–318. http://dx.doi.org/10.1080/02705060.2002.9663899 pires, a.m., magalhães, m.f., moreira da costa, l., alves, m.j. & coelho, m.m., 2008, ‘effects of an extreme flash flood on the native fish assemblages across a mediterranean catchment’, fisheries management and ecology 15, 49–58. http://dx.doi.org/10.1111/j.1365-2400.2007.00570.x resh, v.h., brown, a. v, covich, a.p., gurtz, m.e., li h.w., minshall, g.w. et al., 1988, ‘the role of disturbance in stream ecology’, journal of the north american benthological society 7, 433–455. http://www.jstor.org/stable/1467300 russell, i.a., 2011, ‘conservation status and distribution of freshwater fishes in south african national parks’, african zoology 46, 117–132. http://dx.doi.org/10.3377/004.046.0103 schlosser, i.j. & angermeier, p.l., 1995, ‘spatial variation in demographic processes of lotic fishes: conceptual models, empirical evidence, and implications for conservation’, american fisheries society symposium 17, 392–401. swartz, e. & impson, d., 2007, ‘pseudobarbus afer’, in iucn red list of threatened species, version 2014.2, viewed 13 october 2014, from http://www.iucnredlist.org swartz, e.r., skelton, p.h. & bloomer, p., 2007, ‘sea-level changes, river capture and the evolution of populations of the eastern cape and fiery redfins (pseudobarbus afer and pseudobarbus phlegethon, cyprinidae) across multiple river systems in south africa’, journal of biogeography 2086–2099. http://dx.doi.org/10.1111/j.1365-2699.2007.01768.x tweddle, d., bills, r., swartz, e., coetzer, w., da costa, l., engelbrecht, j. et al., 2009, ‘the status and distribution of freshwater fishes’, in w.r.t. darwall, k.g. smith, d. tweddle & p. skelton, (eds.), the status and distribution of freshwater biodiversity in southern africa, pp. 21–37, international union for conservation of nature, gland, switzerland, and south african institute for aquatic biodiversity, grahamstown, south africa. weyl, o.l.f., finlayson, b., impson, n.d., woodford, d.j. & steinkjer, j., 2014, ‘threatened endemic fishes in south africa's cape floristic region: a new beginning for the rondegat river’, fisheries 39, 270–279. http://dx.doi.org/10.1080/03632415.2014.914924 article information author: peter novellie1 angela gaylard1 affiliation: 1conservation services division, south african national parks, south africa correspondence to: peter novellie postal address: po box 787, pretoria 0001, south africa dates: received: 15 aug. 2013 accepted: 07 june 2013 published: 13 aug. 2013 how to cite this article: novellie, p. & gaylard, a., 2013, ‘long-term stability of grazing lawns in a small protected area, the mountain zebra national park’, koedoe 55(1), art. #1108, 7 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1108 copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. long-term stability of grazing lawns in a small protected area, the mountain zebra national park in this original research... open access • abstract • introduction • research method and design    • setting    • procedure • results • discussion • conclusion • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ we examined a heavily grazed plant community dominated by creeping grass species with the aim of, (1) determining its response to the exclusion of grazing and (2) its long-term persistence. this plant community was particularly favoured by wild ungulate species that prefer short grasses – blesbok (damaliscus dorcas phillipsi), springbok (antidorcas marsupialis) and black wildebeest (connochaetes gnou). exclusion of grazing by large herbivores by means of fencing resulted in the virtual disappearance of the creeping grasses and their replacement by tall tufted species. on plots that remained unfenced, the plant species composition was found to be little changed after an interval of more than 20 years. the number large stock unit equivalents (lsu) per ha carried by the plant community was used as a proxy for grazing intensity. monitored for approximately 2 years at the start of the study, lsu per ha was found to greatly exceed levels recommended for commercial livestock production. this plant community conforms to a recently published definition of a grazing lawn, in that intense grazing promotes palatable, grazing-tolerant grass species. conservation implications: the positive association between grazers and grazing-tolerant grass species evidently persisted for more than 20 years and there was no evidence of an increase in abundance of unpalatable plant species. despite the small size of the park, which limited the extent of large herbivore movements, localised heavy grazing did not lead to range degradation. introduction top ↑ the term ‘grazing lawn’ has been widely used to describe patches that, as a result of frequent, intense grazing, become dominated by short, grazing-tolerant grass species (archibald 2008; archibald et al. 2005; cromsigt & olff 2008; mcnaughton 1984; stock, bond & van de vijver 2009; waldram, bond & stock 2007). in a detailed review of this phenomenon, cromsigt and kuijper (2011:212) defined a grazing lawn as ‘a grazed or browsed patch in which intense foraging leads to increased resource availability and a proportional increase in palatable species in the patch’. thus, grazing lawns are characterised by a positive association between grazers and grazing-tolerant grasses; intensive grazing has the effect of increasing the productivity and quality of the food resource (cromsigt & kuijper 2011). it has been demonstrated (archibald 2008; cromsigt & olff 2008; stock et al. 2009; waldram et al. 2007) that grazing lawns do not derive solely from existing matrices of abiotic factors, such as soil heterogeneity (although their distribution may be influenced by such factors), but are animal driven. consequently, they can increase or decrease depending on fire, rainfall and grazing regime.the formation of a shifting mosaic of grazing lawns together with areas of taller grass appears to be a key determinant of spatial heterogeneity of food resources that is necessary to maintain a diversity herbivore species (cromsigt & olff 2008; fuhlendorf & engle 2001; pickett, cadenasso & benning 2003) and is important for maintaining biological diversity and stability in grasslands (tilman & downing 1994). from this perspective, patch-selective grazing and grazing lawn formation should be regarded as a desirable phenomenon in grassland systems that are managed with the objective of maintaining biodiversity and natural ecological processes. however, concern has been expressed about possible negative effects of patch-selective grazing (fuls 1992; kellner & bosch 1992). it is conceivable that, under certain conditions, protracted use of individual patches may reach a point where the positive association between grazers and palatable grasses breaks down and the lawn transforms to a state that can no longer support grazing. this may be a particular risk in small protected areas, where fences restrict the spatial scale over which grazing dynamics can play out (owen-smith 1983). under these conditions, patch-selective grazing may not result in a stable (or at least resilient) mosaic of grazing lawns, but may instead cause progressive depletion of the grazing resource. to avoid range deterioration in small protected areas, it has been proposed that stocking rates of wild ungulate species should be maintained in accordance with norms set for commercial livestock production (meissner 1982). however, if grazing lawns are prone to transform to a different state, then the setting of stocking rates for the area as a whole may be ineffective in preventing such change (kellner & bosch 1992). it is therefore important to understand the drivers of grazing lawn formation in small protected areas and to monitor the stability of the grazing lawn mosaic. the key question is whether the lawns retain their attractiveness to grazing ungulates over the long term, or whether they eventually transform to a state, either through soil loss or encroachment by unpalatable species, which no longer provides grazing. the mountain zebra national park (mznp) is a relatively small protected area, which carries localised high densities of grazing ungulates, some of which strongly favour grazing lawns (novellie 1990). this study was conducted on some of the most heavily used grazing lawns in the mznp, examining (1) their long-term persistence and (2) response to the exclusion of grazing by means of fenced exclosures. research method and design top ↑ setting the mznp is situated 24 km west of the town of cradock in the eastern cape province, south africa. at the time when the observations commenced (1987), the park was only 6536 ha. in the late 1990s it was greatly expanded in size, extending over 28 412 ha at the conclusion of the study. monthly minimum and maximum temperatures vary from 6 °c to 28 °c in summer (from september to march) and from 0 °c – 20 °c in winter (from april to august) (brown & bezuidenhout 2000). rainfall averages about 400 mm, with most (70%) falling in the summer months. periodic light snow occurs during the winter months and frost is common between may and october. the park lies in a transition zone, incorporating elements of three biomes, the nama-karoo, grassland and albany thicket. it includes three of the vegetation types recognised by mucina and rutherford (2006): the eastern upper karoo, karoo escarpment grassland and eastern cape escarpment thicket. this study was conducted in karoo escarpment grassland in a valley near the western boundary of the park, near two small impoundments called the oudamme. although fires occur periodically in the higher karoo escarpment grassland of the park, the areas of grazing lawn seldom, if ever, burn. no fires occurred in the oudamme valley between 1987 and 2010. data on habitat use by grazing ungulates (novellie 1990, 1991) revealed that the oudamme valley was one of the most heavily grazed areas of the park during the late 1980s. this area was therefore selected for a more detailed assessment of the long-term stability of the grazing lawns. procedure in march and april of 1987, four matched pairs of plots, each roughly 30 m by 30 m in size, were laid out in the heavily grazed areas of the oudamme valley. in each plot a 200-point survey for canopy spread cover was conducted (see novellie & strydom 1987 for further details). the points were arranged in eight rows of 25 each, with 1-m spacing between points and 1-m or 2-m spacing between rows. a canopy spread strike was recorded if a point fell within an imaginary line drawn around the perimeter of the canopy of a plant. for each strike the species of plant was recorded. one member of each pair of plots was then selected at random and a fence approximately 1.5 m high was erected around it to exclude grazing by large herbivores. the other member of each pair remained exposed to grazing. using the same method, the surveys of two of the pairs of plots were repeated in june 2009. the surveys of the remaining two pairs were completed in june 2010. the plant species names reported in this article follow klopper et al. (2006). canonical correspondence analysis was used to investigate changes from 1987 to 2009–2010 in plant species composition in the fenced and unfenced plots (ter braak 1987).between 1987 and 2009–2010 no formal surveys were conducted, but the plots were visited every second or third year (by the first author) to get a general impression of the state of the vegetation. an indication of the intensity of use of the habitat by large herbivores was obtained by conducting five to ten counts per month from 1987 to 1989. ideally, the density of animals should have been sampled exactly within the four 30 m by 30 m plots that were left unfenced, but at this small scale a very large number of counts would have been needed to get a reliable indication of average animal density. it was therefore necessary to conduct the animal counts in larger-sized plots. two plots, each 100 m by 100 m (1 ha) were identified in similar habitat, situated about 200 m apart from one another and adjacent to the four matched pairs of plots used for the exclosure study. the corners were marked with 1.8 m high steel fencing standards which could be seen through binoculars by an observer on the ridge overlooking the oudamme valley. thus, the animals within the boundaries of the plots could be counted without disturbing them. the canopy spread cover of plant species within the two 1 ha plots was determined by 200-point surveys, as described above. the 200-point surveys were replicated at randomly chosen locations within each 1 ha plot, ten surveys in one plot and eight in the other. (the results of these surveys are also reported by novellie 1990). in total, one of the 1 ha plots was checked 199 times and the other 255 times. for comparison with department of agriculture stocking rate standards as applied during the 1980s, the average numbers of ungulates in each plot were converted to large stock unit equivalents (lsu), according to the method of meissner (1982), as published in the government gazette (republic of south africa 1985). animal counts in the 1 ha plots were not continued after 1989, but the locations of animals plotted during the annual helicopter census (gaylard 2011) provided a rough indication of use of the oudamme valley by large herbivores between 1997 and 2011. results top ↑ table 1 shows the mean canopy spread cover of grass species in the four matched pairs of plots and each of the two 1 ha plots at the start of observations in 1987. the species composition was broadly similar across all plots confirming that they were representative of the same plant community. the most abundant grasses in terms of canopy spread cover were the creeping species cynodon incompletus and tragus koelerioides. the former grew in dense lawns around accumulations of dung but elsewhere occurred generally as sparse runners. in contrast, t. koelerioides was not closely associated with dung patches and occurred as sparse runners rather than lawns. the bunch grasses merxmuellera disticha, digitaria eriantha and eragrostis curvula were next most abundant in terms of cover. merxmuellera disticha was very patchy in distribution and the tall, dense tufts showed relatively little signs of grazing. in contrast, d. eriantha and e. curvula were widely distributed and virtually all tufts of these species were cropped to a height of around 5 cm. the annual grass aristida congesta was widespread but its cover was generally low. the taller bunch grasses: themeda triandra, cymbopogon pospischilii and heteropogon contortus were rare throughout the oudamme valley, the only specimens seen at that time were growing within the canopy of shrubs where they were protected against grazing. table 1: percentage canopy spread cover (mean ± standard error) of grass species compared between the study plots (four matching pairs of plots roughly 30 m by 30 m each) and plots 1 and 2 (each 100 m by 100 m) which were used to determine the density of grazing ungulates using the area. figure 1 and figure 2 compare the fenced and matched unfenced plots in the years 1987 and 2009–2010. only the most abundant and widespread grass (figure 1) and dwarf shrub (figure 2) species are shown, as the others were either too uncommon or too localised to draw conclusions as to their response to the exclusion of grazing. figure 1: mean percentage canopy cover of the most common grass species in four pairs of plots compared between 1987 and 2009–2010 for fenced and unfenced plots: (a) aristida congesta (annual grass), (b) cynodon incompletus (creeping grass), (c) tragus koelerioides (creeping grass), (d) eragrostis curvula (bunch grass), (e) digitaria eriantha (bunch grass), (f) themeda triandra (bunch grass), (g) cymbopogon pospischilii (bunch grass) and (h) heteropogon contortus (bunch grass). figure 2: mean percentage canopy cover of some dwarf shrub species in four pairs of plots compared between 1987 and 2009–2010 in fenced and unfenced plots: (a) felicia fascicularis (dwarf shrub), (b) helichrysum dregeanum (dwarf shrub) and (c) chrysocoma ciliata (dwarf shrub). the canonical correspondence analysis showed that species composition was significantly associated with treatments, with the strongest association for the fenced plots in 2009–2010 (f = 9.77; p = 0.002). the ordination diagram (figure 3) shows how the species were clustered with reference to the treatments. of the grass species, t. triandra, c. pospischilii and h. contortus formed a group that was very strongly associated with the exclosures as surveyed during 2009–2010. in clear contrast to this group, the exclosure plots were characterised by a lack of the annual grass a. congesta and the creeping grasses c. incompletus and t. koelerioides. the bunch grasses e. curvula and d. eriantha were in an intermediate position in that they were associated with all plots, although they both showed a relative increase in cover in the fenced plots in 2009–2010 relative to the other plots (figure 1). figure 3: ordination diagram of the canonical correspondence analysis. in summary, the fenced plots in 2009–2010 showed a complete transformation in grass species composition; the creeping and annual species all but disappeared and taller bunch grass species were dominant (figure 1). on the other hand, the unfenced plots showed relatively minor changes in grass species composition from 1987 to 2009–2010 (figure 1). in contrast to the grasses, none of the common dwarf shrub species showed a consistent response to fencing, instead differences are apparent between the survey years (figure 2 and figure 3). chrysocoma ciliata is associated with both the fenced and unfenced plots in 1987, but in 2009–2010 it declined in all plots. helichrysum dregeanum is associated with unfenced and fenced plots, but only in 2009–2010. felicia fascicularis increased in the unfenced plots, but there was considerable between-plot variation in the case of the fenced plots. the reason for the changes in abundance in these dwarf shrub species from 1987 to 2009–2010 is unknown. over the period 1987–1989, both of the 1 ha plots were utilised mainly by short grass-grazers, blesbok (damaliscus dorcas phillipsi), black wildebeest (connochaetes gnou) and springbok (antidorcas marsupialis). mountain zebra (equus zebra zebra) and ostrich (struthio camelus) were observed there comparatively infrequently (table 2). red hartebeest (acelaphus buselaphus), also common in the park at the time, were never observed in these plots. the stocking rate carried by plot 1 was estimated at 9.7 ha per lsu, whilst that of plot 2 was 1.8 ha per lsu. the difference in animal density between the two may have been caused by the fact that there was more surface rock in plot 1; hence, the cover of short grass favoured by grazers was lower in plot 1 (mean ± standard error: 16.0% ± 2.2%) than in plot 2 (28.0% ± 2.7%). for both plots, the stocking rates are considerably higher than the 14 ha per lsu recommended for the area by the grootfontein agricultural development institute (1986). table 2: mean numbers of large herbivore species counted in two 1 ha sized plots from 1987 to 1989, with large stock unit equivalents and stocking rate expressed as ha per large stock unit equivalent. the census results (table 3) indicated that the oudamme valley continued to support fair numbers of large herbivores from 1997 to 2011. it was, however, not possible to be sure from the census results of the exact plant community being utilised, as the animals are likely to have moved around in response to the helicopter. this is probably the reason why table 3 indicates a relatively greater abundance of mountain zebra and mountain reedbuck, species which tend to favour the ridges around the valley rather than the valley bottom habitat in the 1 ha plots. for this reason, we did not attempt to estimate animal densities from the census results. table 3: numbers of different species counted during the annual helicopter census in the oudamme valley from 1997 to 2011. discussion top ↑ the results described above are in accordance with evidence in the literature (archibald 2008; cromsigt & olff 2008; stock et al. 2009; waldram et al. 2007) that grazing lawns are animal driven; if grazing is removed the creeping grasses give way to taller bunch grasses. the change in species composition in the exclosures follows the successional sequence predicted by vorster’s (1982) veld condition index for the karoo region, which classifies t. triandra, c. pospischilii, h. contortus and d. eriantha as ‘decreaser’ species, which decrease in abundance if subjected to heavy grazing, whereas e. curvula, c. incompletus and t. koelerioides are ‘increaser’ species which are typically promoted by grazing. in terms of vorster’s index, which is designed as a guide to commercial pastoralists, the proliferation of the increaser species at the expense of decreaser species is regarded as a sign of deterioration of veld condition, something to be rectified by reducing stocking densities. this is obviously not true for natural systems, where grazing lawns of increaser species are the product of a positive association between consumers and the resource. this is evident from the fact that blesbok, springbok and black wildebeest showed a strong preference for communities dominated by increaser species (novellie 1990) and occupied these habitat patches at relatively high densities. moreover, the increaser species are highly palatable to the short-grass grazers, as is evident from the extent to which they were defoliated in the study area. it is noteworthy that, although all the fenced plots were dominated by one or other bunch grass species, there were differences between the plots in the particular grass species that eventually came to dominate; in some it was t. triandra, in others c. pospischilii or d. eriantha. given that the tufted grasses were extremely rare in the area before fencing, it seems likely that this was caused by chance variation between plots in the nearest sources of colonising seeds. for this plant community (described as lower slope degraded dwarf shrubland by van der walt 1980), the term ‘grazing lawn’ is perhaps a misnomer, because it is only c. incompletus that forms actual lawns and these are generally no more than a few square metres in area immediately surrounding dense patches of ungulate dung. nevertheless, we regard it as appropriate to refer to the c. incompletus – t. koelerioides community as a whole – rather than specifically the cynodon lawns – as a grazing lawn, as it conforms to cromsigt and kuijper’s (2011:212) definition of a lawn in that ‘intense foraging leads to increased resource availability and a proportional increase in palatable species in the patch’. the most common dwarf shrub species associated with the grazing lawns, f. fascicularis and h. dregeanum, are regarded as being palatable to domestic stock by vorster (1982) and, as noted, frequently showed signs of browsing in the oudamme valley. it is interesting to consider whether this community may qualify as a browsing as well as a grazing lawn (as defined by cromsigt & kuijper 2011). however, these dwarf shrubs persisted in the exclosures (figure 2) so, unlike the creeping grasses, their presence does not appear to be dependent on grazing. we did not quantitatively measure the rate at which the transformation in plant species composition within the exclosures took place, but the periodic inspections during the study period suggested that it occurred very slowly. little change in species composition could be discerned during the first 5 years after construction of the exclosures (p. novellie, pers. obs.). as noted, the tall grass species that came to dominate the exclosures were initially very uncommon in the area, so the slow transition may have been due to scarcity of seed. in the mznp, it appears that the mechanism driving the formation of these grazing lawns is the patch-selective grazing of blesbok, springbok and black wildebeest. the habit of the territorial males of these species of occupying on ‘stamping grounds’ and of defecating and urinating at specific localities (du plessis 1972; lynch 1971; novellie 1975, 1990; von richter 1972), results in accumulations of dung and associated cynodon lawns. termite mounds are present in the oudamme valley and may also play a facilitating role in influencing the formation of grazing lawns (see cromsigt & olff 2008). du plessis (1972) describes how a blesbok male, introduced into a paddock, initially focused his grazing around the margins of a termite mound, eventually creating a patch of closely grazed grass. megaherbivores, such as white rhino, have been demonstrated to play a key role in creating grazing lawns in hluhluwe-imfolozi park (cromsigt & olff 2008; waldram et al. 2007), but megaherbivores are absent from the mznp. conclusion top ↑ although there has been concern around the possibility that the grazing lawns may transform into an ungrazable state, it appears that the plant species composition on the grazed plots showed little change 20 years after the initial surveys, despite continued use of the area by the short-grass grazers. not having conducted interim descending point surveys we are uncertain as to fluctuations in species composition that may have occurred during the 20 year interval. however, no sign of transformation could be detected during the visits to the study area made about every 2–3 years; in general, the appearance the plant species composition appeared to be very stable. there was never any indication of dominance by the species of annual grasses and unpalatable dwarf shrubs (e.g. a. congesta and c. ciliata) that were identified by vorster (1982) as characterising the worst state of veld degradation in the karoo region. thus, no evidence could be detected of a breakdown of the positive association between grazers and the creeping grasses. however, this is not to say that it could not occur under certain circumstances. in small protected areas it is highly desirable to develop a predictive understanding of the relationship between the overall stocking rate maintained, on the one hand, and grazing lawn stability and/or resilience on the other. acknowledgements top ↑ we are grateful to park manager, ettienne fourie, for constructing and maintaining the exclosures in good condition over many years, to section ranger, robyn wood, and student ranger, nonkazimlo baca, for able assistance with the vegetation surveys and to animal ecologist, guy castley, for historical air census data. an earlier draft of the article was greatly improved by the comments of two anonymous reviewers. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions p.n. (south african national parks) conducted the on-the-ground field surveys and wrote the manuscript, whilst a.g. (south african national parks) collected air census data, conducted the statistical analysis and made conceptual contributions. references top ↑ archibald, s., 2008, ‘african grazing lawns – how fire, rainfall, and grazer numbers interact to affect grass community states’, journal of wildlife management 72(2), 492–501. http://dx.doi.org/10.2193/2007-045archibald, s., bond, w.j., stock, w.d. & fairbanks, d.h.k., 2005, ‘shaping the landscape: fire-grazer interactions in an african savanna’, ecological applications 15, 96–109. http://dx.doi.org/10.1890/03-5210 brown, l.r. & bezuidenhout, h., 2000, ‘the phytosociology of the farm de rust in the mountain zebra national park, eastern cape’, koedoe 43(1), 1–18. http://dx.doi.org/10.4102/koedoe.v43i1.204 cromsigt, j.p.g.m. & kuijper, d.p.j., 2011, ‘revisiting the browsing lawn concept: evolutionary interactions or pruning herbivores?’, perspectives in plant ecology, evolution & systematics 13, 207–215. http://dx.doi.org/10.1016/j.ppees.2011.04.004 cromsigt, j.p.g.m. & olff, h., 2008, ‘dynamics of grazing lawn formation: an experimental test of the role of scale-dependent processes’, oikos 117, 1444–1452. http://dx.doi.org/10.1111/j.0030-1299.2008.16651.x du plessis, s.s., 1972, ‘ecology of blesbok with special reference to productivity’, wildlife monographs 30, wildlife society, inc., bethesda. fuhlendorf, s.d. & engle, d.m., 2001, ‘restoring heterogeneity on rangelands: ecosystem management based on evolutionary grazing patterns’, bioscience 51(8), 625–632. http://dx.doi.org/10.1641/0006-3568(2001)051[0625:rhorem]2.0.co;2 fuls, e.r., 1992, ‘semi-arid and arid rangelands: a resource under siege due to patch-selective grazing’, journal of arid environments 22, 191–193. gaylard, a., 2011, ‘aerial survey report for the mountain zebra national park’, unpublished report, south african national parks. grootfontein agricultural development institute, 1986, ‘grazing capacity zones for the karoo region’, grootfontein agricultural development institute, middelburg. kellner, k. & bosch, o.j.h., 1992, ‘influence of patch formation in determining the stocking rate for southern african grasslands’, journal of arid environments 22, 99–105. klopper, r.r., chatelain, c., banninger, v., habashi, c., steyn, h.m., de wet, b.c. et al., 2006, ‘checklist of the flowering plants of sub-saharan africa. an index of accepted names and synonyms’, south african botanical diversity network report 42, sabonet, pretoria. lynch, c.d., 1971, ‘a behavioural study of the blesbok, damaliscus dorcas phillipsi, with special reference to territoriality’, unpublished msc. thesis, department of zoology, university of pretoria. mcnaughton, s.j., 1984, ‘grazing lawns: animals in herds, plant form, and coevolution’, the american naturalist 124(6), 863–886. http://dx.doi.org/10.1086/284321 meissner, h.h., 1982, ‘theory and application of a method to calculate forage intake of wild southern african ungulates for purposes of estimating carrying capacity’, south african journal of wildlife research 12, 41–47. mucina, l. & rutherford, m.c. (eds.) 2006, ‘the vegetation of south africa, lesotho and swaziland’, strelitzia 19, south african national biodiversity institute, pretoria. novellie, p.a., 1975, ‘comparative social behaviour of springbok and blesbok on the jack scott nature reserve, transvaal’, unpublished msc. thesis, department of zoology, university of pretoria. novellie, p.a., 1990, ‘habitat use by indigenous grazing ungulates in relation to sward structure and veld condition’, journal of the grassland society of southern africa 7, 16–23. novellie, p.a., 1991, ‘seasonal movements and habitat use by african grazing ungulates in a small conservation area’, in proceedings of the ivth international rangeland congress, vol. 2, pp. 709–713, association française de pastoralisme, montpellier, france. novellie, p.a. & strydom, g., 1987, ‘monitoring the response of vegetation to use by large herbivores: an assessment of some techniques’, south african journal of wildlife research 17, 109–117. owen-smith, r.n., 1983, ‘dispersal and the dynamics of large herbivores in enclosed areas: implications for management’, in r.n. owen-smith (ed.), management of large mammals in african conservation areas, pp. 127–143, haum educational publishers, pretoria. pickett, s.t.a., cadenasso, m.l. & benning, t., 2003, ‘biotic and abiotic variability as key determinants of savanna heterogeneity at multiple temporal scales’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 23–40, island press, washington, dc. republic of south africa, 1985, ‘government notice r. 2687’, government gazette 10029 (06 december)’, government printers, pretoria. stock, w.d., bond, w.j. & van de vijver, c.a.d.m., 2009, ‘herbivore and nutrient control of lawn and bunch grass distributions in a southern african savanna’, plant ecology 206(1), 15–27. http://dx.doi.org/10.1007/s11258-009-9621-4 ter braak, c.j.f., 1987, ‘the analysis of vegetation–environment relationships by canonical correspondence analysis’, plant ecology 69, 69–77. http://dx.doi.org/10.1007/bf00038688 tilman, d. & downing, j.a., 1994, ‘biodiversity and stability in grasslands’, nature 367, 363–365. http://dx.doi.org/10.1038/367363a0 van der walt, p.t., 1980, ‘a phyto-sociological reconnaissance of the mountain zebra national park’, koedoe 23, 1–32. http://dx.doi.org/10.4102/koedoe.v23i1.632 von richter, w., 1972, ‘territorial behaviour of the black wildebeest connochaetes gnou’, zoologica africana 7, 207–231. vorster, m., 1982, ‘the development of the ecological index method for assessing veld condition in the karoo’, proceedings of the grassland society of southern africa 17, 84–89. waldram, m., bond., w. & stock, w., 2007, ‘ecological engineering by a mega-grazer: white rhino impacts on a south african savanna’, ecosystems 11, 101–112. http://dx.doi.org/10.1007/s10021-007-9109-9 article information authors: richard j. stirzaker1 dirk j. roux2 harry c. biggs3 affiliations: 1csiro land and water, canberra, australia 2south african national parks, george, south africa 3south african national parks, skukuza, south africa correspondence to: richard stirzaker email: richard.stirzaker@csiro.au postal address: po box 1666, act 2601, australia dates: received: 04 june 2010 accepted: 28 nov. 2010 published: 13 may 2011 how to cite this article: stirzaker, r.j., roux, d.j. & biggs, h.c., 2011, ‘learning to bridge the gap between adaptive management and organisational culture’, koedoe 53(2), art. #1007, 6 pages. doi:10.4102/koedoe.v53i2.1007 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) learning to bridge the gap between adaptive management and organisational culture in this essay... open access • abstract • introduction • science and certainty • complex and ‘knowable’ systems • adaptive management • adaptive management and organisational culture • learning • conclusion • references abstract (back to top) adaptive management is the problem-solving approach of choice proposed for complex and multistakeholder environments, which are, at best, only partly predictable. we discuss the implications of this approach as applicable to scientists, who have to overcome certain entrained behaviour patterns in order to participate effectively in an adaptive management process. the challenge does not end there. scientists and managers soon discover that an adaptive management approach does not only challenge conventional scientific and management behaviour but also clashes with contemporary organisational culture. we explore the shortcomings and requirements of organisations with regard to enabling adaptive management. our overall conclusion relates to whether organisations are learning-centred or not. do we continue to filter out unfamiliar information which does not fit our world view and avoid situations where we might fail, or do we use new and challenging situations to reframe the question and prepare ourselves for continued learning? conservation implications: for an organisation to effectively embrace adaptive management, its mangers and scientists may first have to adapt their own beliefs regarding their respective roles. instead of seeking certainty for guiding decisions, managers and scientists should acknowledge a degree of uncertainty inherent to complex social and ecological systems and seek to learn from the patterns emerging from every decision and action. the required organisational culture is one of ongoing and purposeful learning with all relevant stakeholders. such a learning culture is often talked about but rarely practised in the organisational environment. introduction (back to top) pursuing the life sciences is as much a calling as it is a career. we are drawn to a science course at university through some combination of a fascination with the living world and a desire to use natural resources wisely. the renowned biologist edward o. wilson says that at age eighteen he saw science as the study of ants, frogs and snakes and a wonderful excuse to stay outdoors (wilson 1998). for many of us, the objective view of the world that science provides appealed more than the literature and history classes we took at school. during university-level practicals, students have to solve problems to which objective answers exist. for example, how much chitin does a crab shell contain? or, what is the identity of the bacteria in the broth? what is the change in momentum when two bodies collide? the tutor already knows the correct answer and if the experiment is performed properly, the students should get it right too. however, there may be several different interpretations to the meaning of a poem, which can all be apparently acceptable. the scientific method gives one correct result, which is independent of the observer. modern science has built this predictive capability on four essential components. firstly, one states a clear hypothesis about cause and effect that is testable by experiment. secondly, one designs an experiment that tests a prediction emanating from the hypothesis in a controlled environment. thirdly, one replicates the experiment to show that the observations are not the result of a chance event. fourthly, the work is documented and subjected to peer review before the new knowledge can serve as building blocks for further investigations. the ability to tease apart a system into its constituent components and study it systematically allows scientists to infer cause and effect. such reductionism is said to be the primary and essential activity of scientific research (wilson 1998). at the outset of this essay, we offer a very brief description of organisational culture as it relates to science, our interest in adaptive management, and why we foresee an uneasy relationship. culture consists of bundles of shared norms, which are behaviours common to a group. norms give a group a sense of cohesion and protection against undesirable change, but can simultaneously cause a group to resist new, potentially beneficial ideas (ehrlich & levin 2005). many scientists share a culture centred around the robust methodology that underpins their craft: the production of reliable information and the ability to make predictions, quantify uncertainty and expose error. scientific thinking has produced knowledge and products that have contributed enormously to economic development and social upliftment over the last 300 years. scientists not exposed to further study of the humanities, or even an introductory course on the philosophy of science, are often surprised by growing criticism of the very norms most of them regard as self-evident. even if it were valid to pass off some of these critics as postmodern deconstructionists, scientists still have to heed a call from within their own community about the need to think differently about contemporary issues facing society (lubchenko 2008; steffen, crutzen & mcneill 2007; walker & salt 2006). for example, ulanowicz (2009:93) writes that ‘it is indeed feasible to march directly into the jaws of oblivion on the tacit assumptions that support conventional science’. holling (1995:29) warns of the ‘pathology’ that emanates from scientists’ belief that they fully understand cause and effect – that success in managing one target variable in isolation leads to ‘less resilient and more vulnerable ecosystems, more rigid and unresponsive management agencies, and more dependent societies’. it is not our purpose here to provide a critique of reductionist science; suffice it to say that we sympathise with the view of ulanowicz (2009) that biology and ecology are not entirely reducible to physics. life scientists cannot fully explain the world from the bottom up and therefore are interested in finding approaches to managing ecological systems that accept and act on this understanding. moreover, since human systems interact with biological systems in diverse ways we propose that adaptive management is the problem-solving approach of choice for environments that are highly changeable, heterogeneous and often unpredictable, and usually involve multistakeholder interest. but we also anticipate that its implementation will frequently conflict with aspects of our own science culture and that of the organisations where we work. science and certainty (back to top) modern society’s knowledge of the physical sciences is so well developed in many areas that we put our total trust in its products. most of us expect an aeroplane to get us safely to our destination even if we do not understand all the intricacies involved in the process. one reason for this certainty is that we can be reasonably sure about the boundaries of the system. we believe that the engineers, pilots and air traffic controllers know which factors they need to understand well, and which they can ignore. the biological sciences present more of a challenge because the boundaries of a system are not always clear. an experiment is performed in ‘controlled’ conditions, yet it is impossible to exclude all factors extraneous to the hypothesis owing to practical constraints often associated with larger-scale experiments. not all of us can fumigate small mangrove islands to study the re-colonisation process (simberloff & wilson 1969). statistical techniques provide an objective means for identifying cause and effect in controlled experiments and authors of scientific publications know that reviewers start to object when one extrapolates findings too far from the experimental conditions under which the data were collected. the new knowledge holds only when other factors are excluded; that is, scientists’ claims are valid within certain defined boundary conditions. by speculating outside the narrow range of the measured data or the conditions in which they were collected, one can no longer claim the same certainty. in other words, our knowledge is obtained by framing or constraining the problem. what does this mean for understanding cause and effect in biological or ecological research? it is tempting to believe that the problem of boundary conditions can be overcome just by doing more experiments until all the combinations and permutations have been exhausted. this view is problematic. as already discussed, replicating experiments under controlled conditions is extremely difficult at ecosystem level, and has even less relevance for the action research of the social disciplines (rogers 2006). but the problem may lie even deeper. walters and holling (1990) state that our ecological knowledge is not only incomplete but also elusive, particularly when we consider the range of values held by different groups in society and the political constraints to action. gallopin et al. (2001) refer to a degree of ‘irreducible uncertainty’ associated with complex social–ecological systems. complex and ‘knowable’ systems (back to top) in order to test the claim of ‘irreducible uncertainty’, one needs to differentiate between different types of p roblem and be able to infer cause and effect accordingly. snowden (2002) categorises problems into those that may be difficult or complicated but are ultimately ‘knowable’ through reductionist scientific method, and problems that are complex and always characterised by an inherent degree of uncertainty. for example, we can categorise an aeroplane as a complicated but knowable machine. although it is made of thousands of different parts, the function of each in relation to another is understood. apart from the most extreme conditions, aeroplanes behave in a predictable way, and we trust them with our lives. human systems tend to be complex: the people that make up the system change and the way they relate to one another is highly context dependent and therefore not entirely predictable. snowden gives the following example: consider what happens in an organisation when a rumour of reorganisation surfaces: the complex human system starts to mutate and change in unknowable ways; new patterns form in anticipation of the event. on the other hand, if you walk up to an aircraft with a box of tools in your hand, nothing changes. (snowden 2005:105) one can accept the notion of an aeroplane being complicated but knowable, while a human system is very unpredictable, but what about biology and ecology? surely biological systems are subject to natural laws that give the system some form of deterministic behaviour? ulanowicz (2009) cautions that the idea of tight cause and effect in open systems should be set aside in preference for poppers’ view of ‘propensities’. a propensity downgrades cause and effect to a more general likelihood of one factor influencing another. more importantly, it is the combined effect of several propensities acting together on the whole system that facilitates unique and sometimes surprising behaviour of a specific system. in table 1 we distinguish between problems that are complicated but knowable and those that are complex (informed partly by snowden and boone 2007). the complexity of the problems in the right-hand column of table 1 is due to feedbacks, thresholds and, often, nonlinear interactions within the system, together with lags and cross-scale effects. such factors combine to give the system a degree of uncertainty. an example of feedback is the reinforcement of terrestrial warming as ice sheets melt, because there is less ice to reflect the sun’s energy. an example of a threshold effect is the rapid switch from savannah vegetation to woodland thicket once perennial grasses that support fire reach a critically low density (walker & meyers 2004). nonlinear interaction means that a small change in one factor can have a big effect somewhere else. when there are multiple such interactions it becomes practically impossible to keep track of all the causal relationships. table 1: some characteristics of knowable and complex problems. complex systems also have so-called emergent properties. an analogy might be ‘team spirit’. the statistical attributes of each member of the team may be known, but the joint interaction of the team sets up a dynamic that strongly affects how the team performs as a unit. in other words, there are mutually beneficial interactions between players that give the team its unique character. if we disassemble the team to study each player’s attributes in detail, the team spirit, or emergent property, disappears. adaptive management (back to top) feedbacks, thresholds, multiple nonlinear interactions, lags, chance events and emergent properties contribute to a general uncertainty about cause and effect, and, consequently, the impact of our management actions. this realisation led to the development of the field of adaptive management (holling 1978; lee 1993; walters 1986). its fundamental premise is that the puzzle of a social–ecological system can never be fully solved by studying the pieces. we have to use real-life management of the system as a whole and turn it into an experiment by asking the right questions, implementing decisions, collecting the right data and learning from the experience. the emphasis is on formulating an explicit mental model, however imperfect, and then acting accordingly by managing and monitoring to see how our understanding can be improved as we gain further insight of the system. furthermore, many of the ecological problems we face are as much a controversy over values as disputes about cause and effect. this challenges the positivist view of science, which regards science as the principal producer of reliable knowledge that should be passed on to those with a management responsibility (ziman 2000). broader society now demands that their local and experiential knowledge, as well as their values, be considered in management plans. therefore, when both the facts are uncertain and their interpretation is contested, we need an approach that can integrate knowledge from different sources and treat management activities as experiments from which we can learn. adaptive management is a way of getting around the dilemma of delaying decisions until we fully understand all the potential consequences of our actions. the act of management is itself an experiment, but clearly not in the traditional sense of controls and replication. to distinguish adaptive management from simple trial and error, considerable effort should be put into integrating existing information from different disciplines and perspectives. appropriate models should be used to frame the questions, eliminate the least likely solutions and identify the knowledge gaps (stankey, clark & bormann 2005; walters 1997). monitoring is a central issue. adaptive management needs an intellectual paper trail to show that reasoning underlies the actions – we cannot learn without this (lee 1993; venter et al. 2008). the conceptual framework, whether represented by a simple diagram or sophisticated model, should be matched by the amount of effort put into monitoring. unfortunately, monitoring is much more expensive than modelling, which can create tension between researchers and funders or managers. often the true value of monitoring will be reaped only by the next generation of scientists; a problem for those responsible for paying the bills now (walters 1997). conversely, it is the responsibility of scientists to identify variables that are most likely to be indicative of system behaviour. this may involve identification of integral measures that remove much of the ‘noise’ and variables that warn in advance of an approaching threshold (stirzaker et al. 2010). we simply cannot measure everything. according to stankey et al. (2005) the principles of adaptive management are widely acclaimed, but remain more an ideal than a demonstrated reality. one of the several reasons rogers, roux and biggs (2000) cite for this is that the new way of operating does not comply with the old organisational culture with an authoritarian structure. moreover, when investigating large multidisciplinary problems we get overloaded with information and often experience ‘turf protection’ among scientists and between scientists and managers. this raises a new question: if issues that we once saw as ‘knowable’ are in fact complex and demand a radically different problem-solving approach, do we also need to think through the ways our organisations operate? adaptive management and organisational culture (back to top) scientists responding to the challenge of living with more uncertainty can find their organisations moving in the opposite direction. the conventional view of curiosity-driven research that leads to new findings, beneficial applications and tools for the improvement of human welfare (ravetz 2004) has given way to formal methods of planning and accountability. science no longer has special status in a government’s budget. the case for investmentin science must be carefully argued with explicit costs and benefits, with timelines to show what will be delivered when and by whom. organisational culture and adaptive management are likely to clash, at least initially, on several fronts. firstly, scientists are expected to produce the knowledge that managers need to make informed decisions. a focus on inherent unpredictability seems to be undermining the foundation of this social contract. secondly, organisations spend considerable time streamlining their portfolio of work into ‘manageable’ units aligned to corporate goals, whereas adaptive management can be a messy web of relationships encompassing scientific, social and political perspectives. thirdly, adaptive management requires us to be open to learning things that may be counter to the way we normally operate. it requires a level of flexibility that challenges the way that things have always been done. as scientific organisations strain under the pressure to adopt a more overt ‘business principles’ approach, there is a greater focus on specifying outputs of programmes well in advance and minimising risk at all stages of the project. reporting on project deliverables and milestones is required on shorter timescales, which makes it easier to follow the contract than to follow up surprises. whereas accountability and risk management are obviously important, rigid management systems run counter to the nature of adaptive management. organisations certainly need to balance order with creativity, but the desire for certainty and control can overwhelm the desire to nurture the flexible learning approach required for adaptive management. according to wheatley (2005), organisations should resist the notion that there is some optimum structure that will deliver results: if a system becomes too homogenous, it becomes vulnerable to environmental shifts. if one form is dominant, and that form no longer works when the environment shifts, the entire system can collapse. … if leaders fail to encourage diverse ways of doings things, they destroy the system’s capacity to adapt. (wheatley 2005:78) if, for example, we lose the balance between flexibility and responsibility and opt for an overly planned and rigid system, we may find we spend more time interacting with the organisation and less with the real world. at the other end of the spectrum the organisation may become too chaotic when everyone follows their own plans. people do not learn from their mistakes and bad behaviour is not brought to account. both extremes are counter-productive. wheatley (2003:39) writes: ’if a system has too much order, it atrophies and dies. yet if it lives in chaos, it has no memory.’ when science is seen as just another business, goals such as improving efficiency are accepted without question. this sounds sensible at first, but there is evidence that targeting efficiency as the prime goal can destroy the very thing we are trying to manage (walker & salt 2006). rogers et al. (2000) cite the example of a ‘command-and-control’ approach to managing water resources. what starts out as efficient delivery of services easily spills over into exploitation when the focus is too narrow and agencies take too long to respond to feedbacks. cilliers (2006:109) argues that the value of organisations lies in their ability to be ’stable enough not to be buffeted around by every fluctuation, [and] … flexible enough to be able to adapt when necessary’. he advises against a culture where ’speed is linked with efficiency, and has become a virtue in itself.’ our real identity is forged when we are able to reflect adequately on our experience, and based on that reflection, to resist certain change (cilliers 2006). yet, when the pressure is on to deliver results too quickly, we are more likely to react than to reflect. learning (back to top) if an organisation is going to embrace adaptive management, it will have to learn to do things differently. stankey et al. (2005) propose that we normally learn by accumulating new facts, but our understanding moves ahead in leaps when we ask new questions and see the old facts in a new light. this is critical for adaptive management. instead of filtering out information that is unfamiliar or does not fit our world view, or avoiding situations where we might fail, we use challenging situations to reframe the question and prepare ourselves for learning new things. garvin (1993) defines a learning organisation as one ‘skilled at creating, acquiring, and transferring knowledge, and at modifying its behaviour to reflect new knowledge and insights.’ modifying behaviour is difficult and should involve three overlapping phases. • cognitive: members of the organisation are exposed to new ideas, expand their knowledge, and begin to think differently. • behavioural: employees begin to internalise new knowledge and alter their behaviour (as manifested, for example, in the use of new vocabulary). • performance improvement: changes in behaviour lead to measurable improvements in results or outcomes. thus far we have dealt with the cognitive aspects of adaptive management: the idea that we should distinguish complex systems from those that may be difficult but ultimately able to be fully understood (table 1). we have also addressed the early behavioural changes: adopting a new language that goes beyond the use of new vocabulary to determine what the new concepts really mean in a specific context and how they can inform our approach to science. the third step – applying the new concepts in the real world – is, however, the major stumbling block. of course, if there is no performance improvement the ‘new’ behaviour will be challenged and will most likely be replaced by a next wave of ideas, or the organisation will default to its old ways. learning is the mechanism through which we change our individual and collective understanding of our world. new knowledge enables us to respond differently to new circumstances and challenges. the rate and relevance of our learning will, in effect, determine our ability to respond to external changes effectively. in this sense, learning proficiency relates to what we should learn about (and what we should forget), who we should learn with, and how we should learn. to accommodate new knowledge, previous learning and beliefs sometimes have to be left behind. selectively ‘forgetting’ outdated knowledge is referred to as unlearning (becker 2005). however, unlearning may not be a straightforward or easily manageable activity. individuals (often unknowingly) protect existing knowledge by actively disregarding conflicting information (lyndon 1989). it appears that more recently acquired knowledge is easier to relinquish than knowledge that was acquired and reinforced over an extended period of time. experts may be especially susceptible to ‘trained incapacity’ (miller & morris 1999): the more someone’s knowledge is shaped by learning within a defined field, the harder it becomes to associate with knowledge that emerges from other fields. environmental issues inevitably imply the involvement of multiple stakeholders. therefore, life scientists need to be prepared to learn together from diverse sources (keen, brown & dyball 2005; pahl-wostl & hare 2004). we should not only settle for compromise but strive for a consensus that can distribute the benefits and costs of our interventions in a way that is (1) equitable and (2) within the ecological limits (rogers 2006). compromise involves trading off conflicting demands against those who hold a contrary position, creating winners and losers. the approach of seeking consensus is about moving beyond the problem and ‘developing a set of shared values that guide future decision making’ (rogers 2006). ongoing learning is uncomfortable. it is much easier to believe we already know most of what we need to know. if we feel overwhelmed by new information our dominant learning mode is reactive and we tend to reinforce pre-established knowledge and frames of reference. scientists need to perceive their working environments as safe to envisage alternative futures and to learn along new and dynamic trajectories towards such futures (senge et al. 2005). furthermore, to learn with other parties who may hold very different world views, requires us to learn with empathy and humility. to be empathetic means to consider different perspectives and assumptions, temporarily suspending our own in the process, so that we can inquire into the reasons for people’s views (senge et al. 1999). in this sense, humility means acknowledging that the knowledge base in any given field is too vast for a single person to master. even the expert’s knowledge is only a partial reflection of what is known. however, by combining one’s partial knowledge with that of others, one can, in practice, use more knowledge than one’s own (wenger 2005). conclusion (back to top) successful adaptive management in a multistakeholder context rests on three pillars, namely the ability to form a robust, shared conceptualisation, the ability to monitor key variables that will shed light upon this conceptualisation, and the ability to learn from the experience. if any of these are compromised, the structure will collapse. it is easy to be so enamoured by the conceptualisation of the problem that we fail to invest in thorough monitoring, and equally easy to keep collecting data without knowing how the knowledge will be used. the test is: are we still learning and can we document our learning journey (venter et al. 2008)? learning is never quick or easy, and involves travelling along detours and going down blind alleys. when disillusioned, scientists should avoid the trap of falling back into the old pattern of over-promising and under-delivering as they proffer ‘silver bullet’ solutions for complex problems to those who control the purse strings. similarly, organisations should resist defaulting to command-and-control systems that appear to have delivered some certainty in the past. organisations will have to find and foster the champions of adaptive learning, including the visionary activist, the respected integrator and the rebel bureaucrat (gunderson, holling & light 1995). there will need to be a shift from the view that benefits come from power or withholding information and ideas, to one where benefits come from sharing, and there are clear incentives to reflect this. learning together rather than competing against one another is absolutely central. science culture has been forged in a competitive environment – competition for best ideas to secure limited funding, competition for space in top journals, and even a league table of citation metrics that purport to show how useful our work has been. learning should involve exploring, discovering, reflecting, listening and sharing frustrations and surprises. managers, scientists and stakeholders need to see themselves as part of the same community, where benefits and risks are shared within the context of a shared vision. rogers and breen leave us with the core challenge: perhaps the most important lesson ecologists should learn is not to enter the new social theatre as ’experts’ (ludwig 2001), but as co-learners, interactive players seeking consensus on stage. for some ecologists, and for ecology as a science, this transition will certainly be difficult. we will judge success by a shift from research outputs that impress peers to outcomes that allow society to better respond to environmental challenges. (rogers & breen 2003:50) references (back to top) becker, k., 2005, ‘individual and organisational unlearning: directions for future research’, international journal of organisational behaviour 9, 659−670. cilliers, p., 2006, ‘on the importance of a certain slowness’, emergence: complexity and organizations 8(3), 106–113. ehrlich, p.r. & levin, s.a., 2005, ‘the evolution of norms’, plos biology 3, 943−948. doi:10.1371/journal.pbio.0030194, pmid:15941355, pmid:1149491 gallopin, g.c., funtowicz, s., o’connor, m. & ravetz, j., 2001, ‘science for the twenty first century: from social contract to the scientific core’, international journal of social science 168, 219−229. doi:10.1111/1468-2451.00311 garvin, d.a., 1993, ‘building a learning organisation’, harvard business review july-august, 78−91. gunderson, l.h., holling, c.s. & light, s.s., 1995, ‘barriers broken and bridges built: a synthesis’, in l.h. gunderson, c.s. holling & s.s. light (eds.), barriers and bridges to the renewal of ecosystems and institutions, pp. 489–532, columbia university press, new york. holling, c.s., 1978, adaptive environmental assessment and management, john wiley, london. holling, c.s., 1995, ‘what barriers? what bridges?’, in l.h. gunderson, c.s. holling & s.s. light (eds.), barriers and bridges to the renewal of ecosystems and institutions, pp. 3– 34, columbia university press, new york. keen, m., brown, v.a. & dyball, r., 2005, ‘social learning: a new approach to environmental management’, in m. keen, v.a. brown & r. dyball (eds.), social learning in environmental management: towards a sustainable future, pp. 1−21, earthscan, london. lee, k.n., 1993, compass and gyroscope. integrating science and politics for the environment, island press, london. lubchenco, j., 2008, ‘entering the century of the environment: a new social contract for science’, science 279, 491−496. doi:10.1126/science.279.5350.491 ludwig, d., 2001, ‘the era of management is over’, ecosystems 4, 758–764. doi:10.1007/s10021-001-0044-x lyndon, h., 1989, ‘i did it my way! an introduction to “old way/new way” methodology’, australasian journal of special education 13, 32−37. doi:10.1080/1030011890130107 miller, w.l. & morris, l., 1999, fourth generation r&d: managing knowledge, technology, and innovation, john wiley & sons, new york. pahl-wostl, c. & hare, m., 2004, ‘processes of social learning in integrated resources management’, journal of community and applied social psychology 14, 193−206. doi:10.1002/casp.774 ravetz, j., 2004, ‘the post-normal science of precaution’, futures 36, 347–357. doi:10.1016/s0016-3287(03)00160-5 rogers, k.h., 2006, ‘the real river management challenge: integrating scientists, stakeholders and service agencies’, river research and applications 22, 269–280. doi:10.1002/rra.910 rogers, k.h. & breen, c.m., 2003, ‘the ecology-policy interface’, frontiers in ecology and the environment 1, 49–50. doi:10.1890/1540-9295(2003)001[0050:tepi]2.0.co;2, doi:10.1890/1540-9295(2003)001[0049:tepi]2.0.co;2 rogers, k., roux, d. & biggs, h., 2000, ‘challenges for catchment management agencies: lessons from bureaucracies, business and resource management’, water sa 26, 505–511. senge, p., kleiner, a., roberts, c., ross, r. & smith, b., 1999, a fifth disciple resource: the dance of change, nicolas brealey, london. senge, p., scharmer, c.o., jaworski, j. & flowers, b.s., 2005, presence: exploring profound change in people, organizations and society, nicholas brealey, london. simberloff, d. & wilson, e.o., 1969, ‘experimental zoogeography of islands: the colonization of empty islands’, ecology 50, 278−296. doi:10.2307/1934856 snowden, d., 2002, ‘complex acts of knowing: paradox and descriptive self-awareness’, journal of knowledge management 6, 100−111. doi:10.1108/13673270210424639 snowden, d.j. & boone, m.e., 2007, ‘a leader’s framework for decision-making’, harvard business review november, 69–76. stankey, g.h., clark, r.n. & bormann, b.t., 2005, adaptive management of natural resources: theory, concepts, and management institutions, department of agriculture, forest service, pacific northwest research station, portland, or. steffen, w., crutzen, p.j. & mcneill, j.r., 2007, ‘the anthropocene: are humans now overwhelming the great forces of nature?’, ambio 36, 614−621. doi:10.1579/0044-7447(2007)36[614:taahno]2.0.co;2 stirzaker, r.j., biggs, h.c., roux, d.j. & cilliers, p., 2010, ‘requisite simplicities to help negotiate complex problems’, ambio 39, 600−607. doi:10.1007/s13280-010-0075-7, pmid:21141779 ulanowicz, r.e., 2009, a third window: natural life beyond newton and darwin, templeton foundation press, indiana university. venter, f.j., naiman, r.j., biggs, h.c. & pienaar, d.j., 2008, ‘the evolution of conservation management philosophy: science, environmental change and social adjustments in kruger national park’, ecosystems 11, 173–192. doi:10.1007/s10021-007-9116-x walker, b.h. & meyers, j.a., 2004, ‘thresholds in ecological and social–ecological systems: a developing database’, ecology and society 9(2), viewed 18 february 2011, from http://www.ecologyandsociety.org/vol9/iss2/art3 walker, b.h. & salt, d.a., 2006, resilience thinking: sustaining ecosystems and people in a changing world, island press, washington dc. walters, c., 1986, adaptive management of renewable resources, macmillan, new york. walters, c., 1997, ‘challenges in adaptive management of riparian and coastal ecosystems’, conservation ecology 1(2), 1, viewed 01 november 2010, from http://www.consecol.org/vol1/iss2/art1/ walters, c.j. & holling, c.s., 1990, ‘large-scale management experiments and learning by doing’, ecology 71, 2060–2068. doi:10.2307/1938620 wheatley, m.j., 2005, finding our way: leadership for an uncertain time, berrett-koehler, san francisco. wenger, e., 2005, learning for a small planet: a research agenda, viewed 15 october 2010, from http://www.ewenger.com/ wilson, e.o., 1998, consilience: the unity of knowledge, vintage books, new york. ziman, j., 2000, real science: what it is, and what it means, cambridge university press, cambridge. doi:10.1017/cbo9780511541391 article information authors: thinandavha c. munyai1 stefan h. foord2 affiliations: 1centre for invasion biology, department of ecology and resource management, university of venda, south africa 2centre for invasion biology, and south african research chair for biodiversity value and change, department of zoology, university of venda, south africa correspondence to: thinandavha munyai email: caswell.munyai@univen.ac.za postal address: private bag x5050, thohoyandou 0950, south africa dates: received: 22 july 2014 accepted: 20 nov. 2014 published: 30 mar. 2015 how to cite this article: munyai, t.c. & foord, s.h., 2015, ‘an inventory of epigeal ants of the western soutpansberg mountain range, south africa’, koedoe 57(1), art. #1244, 12 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1244 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. an inventory of epigeal ants of the western soutpansberg mountain range, south africa in this original research... open access • abstract • introduction • materials and methods    • study area       • ant sampling    • data analysis • results • discussion • conclusion • acknowledgments    • competing interests    • authors’ contributions • references • appendix 1 • appendix 2 • appendix 3 abstract top ↑ the distribution, abundance and sensitivity of invertebrates to habitat change are largely unknown. long-term monitoring of ecological gradients with standardised and comparable protocols can form the basis of a better understanding. altitudinal gradients are particularly relevant within this context. here we provide a check list and baseline data for ant species collected over a 5-year period across the soutpansberg mountain range, south africa. standardised pitfall surveys across 11 sites yielded a total of 133 species in 38 genera and 6 subfamilies. sample coverage of epigeal ants was 0.98 for the transect as a whole. of these species, 21% were restricted to the southern slope of the mountain and 14% to the northern slope. extrapolated richness estimates reached an asymptote for all, except for three sites. these were the only sites impacted by bush encroachment. observed richness was the highest at a low-altitude mesic site that is exposed to considerable disturbance by megaherbivores and mechanical clearing of woody vegetation. structural classification of vegetation was predictive of a broad-scale ant assemblage structure. on a smaller scale, however, structure was a function of elevation, space and temperature. conservation implications: future monitoring should target indicator taxa associated with bush encroachment, particularly with reference to their impacts on grasslands. bush encroachment could endanger several ant species associated with mesic grasslands and woodlands on the mountain, as well as ant diversity, as these were the habitats with the highest ant diversity. introduction top ↑ although invertebrates comprise the bulk of diversity in an ecosystem, they are often excluded from conservation initiatives. cardoso et al. (2011) identified seven reasons for this. three of the shortfalls had to do with: (1) the distribution of invertebrate species is unknown (wallacean), (2) invertebrate abundance and their changes in space and time are unknown (prestonian) and (3) invertebrates’ way of life and sensitivity to habitat types are largely unknown (hutchinsonian). these three shortfalls are particularly relevant to how ecologists practise their science, as the distribution, abundance in time and space, and sensitivities to habitat change are largely unknown for most species. these constraints can be resolved through inventories that follow standardised and comparable protocols (cardoso et al. 2011). ants respond to the environment at small scales and considerable support exists for their use as indicators of disturbance and habitat degradation (see andersen & majer [2004] for review). ants are easily collected and species or morpho-species level determination are aided through readily available taxonomic expertise (ward 2010). in addition to being diverse, abundant and a dominant component of animal biomass in terrestrial ecosystems (hölldobler & wilson 1990), they also perform important functions in ecosystems, including pollination, myrmechory and nutrient cycling, among others (wielgoss et al. 2014). the soutpansberg mountain range is a centre of endemism (hahn 2002) and is the focal point for the newly proclaimed vhembe biosphere reserve, one of the united nations educational, scientific and cultural organisation (unesco) world network of biosphere reserves (www.unesco.org. it has the highest plant generic and family level diversity among the 18 recognised centres of plant endemism in southern africa (van wyk & smith 2001). recent results suggest that this diversity and endemism is mirrored by spiders (foord 2008) and reptiles (kirchhof et al. 2010). as an inselberg that rises 1000 m above the surrounding plains, the soutpansberg mountain range has provided a refuge for several taxa during periods of climate change and will once more play a biogeographic role in the region as regional climate changes take effect in response to elevated global greenhouse gases (tshiala et al. 2011). transects can provide valuable diversity, distribution and abundance information as well as population trends. altitudinal transects are particularly relevant within the context of contemporary climate changes. here we summarise the results of a standardised inventory of epigeal ants over a 5-year period (2009–2013), along a transect across the western soutpansberg mountain range. we provide a checklist of epigeal ants with a sample coverage of 0.98 for the whole transect. a posteriori defines distinct ant assemblages across the transect and tests for differences between these groups. materials and methods top ↑ study area the 16.5 km altitudinal transect extends over the highest point of the soutpansberg mountain range, lajuma (1748 m), in the luvhondo nature reserve. it includes the northern and southern slopes of the mountain and consists of 11 sites, spaced at 200 m altitudinal intervals. it starts at 800 m a.s.l. in the limpopo valley to the north of the mountain, ascends 900 m to lajuma, and descends another 800 m to the plains south of the range (figure 1). figure 1: location of the soutpansberg transect in south africa (inset) and the position of the 44 replicates with the 5 structural habitat types. the study area stretches across five broad structural vegetation types, namely woodland, sedgeland-herbland, forest, thicket and shrubland. all the sites on the northern aspect are woodland (total tree cover 0.1% – 75%; shrub cover < 10%; mean crown-gap ratio > 0.1). they can be characterised as low (1 m – 5 m-tall trees), open (crown-gap ratio of 2–8.5) and woodland (figures 2a–2d), mostly on shallow quartzitic soils (figures 2b–2d). one site is characterised by deep sandy soils (figure 2a). dominant trees species include burkea africana, ochna pulchra, diospyros dichrophylla, rhus magalismontanum, boscia albitrunca, adansonia digitata, acacia nigrescens and terminalia sericea. figure 2: (a) 08n and (b) 12n deep sandy soils characterised by trees such as adansonia digitata, acacia nigrescens and terminalia sericea. (c) 10n and (d) 14n are on shallow quartzitic soils and include burkea africana, ochna pulchra, diospyros dichrophylla, rhus magalismontanum, boscia albitrunca and terminalia sericea. higher elevation sites (e) 17n and (f) 16s low shrubland patches with shallow rocky soils on quartzite, dominated by coleochloa species. vegetation types along the plains and lower southern foothills are largely thicket (figure 3d) and shrubland (figure 3e), located on red loamy-clay soils dominated by woody species such as dichrostachys cinerea, acacia caffra and olea europaea. figure 3: (a) low shrubland patches (14s) with shallow rocky soils on quartzite, dominated by coleochloa species. (b) tall forest (12s) dominated by woody species of croton sylvaticus and ekebergia capensis. (c) short forest (12s2). low elevation sites: (d) thicket site (10s) on clay soils dominated by woody species such as acacia caffra and olea europaea. (e) shrubland (09s) on clay soils dominated by woody species such as dichrostachys cinerea. structurally, the lowest site on the southern aspect (figure 3e) is tall, closed shrubland (< 0.1% tree cover). it is dominated by a d. cinerea matrix with patches that are mechanically cleared of all woody plants. site 10s (figure 3d) is low thicket (10% – 100% tree cover and > 10% shrub cover). the forests are found at mid-elevation, between 1200 m a.s.l. and 1400 m a.s.l. on the southern slope and include both short (figure 3c) and tall (figure 3b) forest sites with a crown-gap ratio of < 0.1. they are dominated by woody species such as croton sylvaticus and ekebergia capensis. the higher elevational zones of the southern aspect and summit consist of a sedgeland-herbland matrix with low shrubland patches that grow on shallow rocky soils on quartzite, dominated by coleochloa species (figures 2e–3a).soil temperature extremes were the highest on the summit of the mountain (mean annual range 10 °c – 55 °c) and the lowest in the tall forests of the southern aspect (14 °c – 32 °c). ant sampling ants were sampled through standardised pitfall trapping (munyai & foord 2012) at each of the 11 sites. sample units consisted of 10 pitfall traps (ø62 mm) laid out in a sampling grid (2 x 5) with 10 m spacing between traps. pitfall traps were left open for 5 days, since it is considered representative and does not sample excessive ants (see munyai & foord [2012] and references therein). from september 2009 onwards (appendix 1), sampling units were spatially replicated four times at each site. replicates were separated by < 300 m to avoid pseudo-replication (ness et al. 2004). the traps contained a 50% solution of propylene glycol because it neither attracts nor repels ants (abensperg-traun & steven 1995). the ant samples were washed in a laboratory and stored in 70% ethanol. they were sorted to morpho-species and, where possible, identified to species by comparison with reference collections housed in the kruger national park in south africa and the commonwealth scientific and industrial research organisation (csiro) tropical ecosystems research centre in darwin, australia. bolton (1994) was used to identify some specimens to genus level, while valid names were confirmed using antcat (http://www.antcat.org) and antweb (http://www.antweb.org). the unidentified (morpho) species were given the following number codes: university of venda collection_01, 02, etc. (i.e. uvoc_01, 02, etc.). bothroponera, mesoponera, ophthalmopone and pseudoponera were considered valid genera rather than being subsumed under pachycondyla (schmidt & shattuck 2014). subfamilies aenictinae and cerapachyinae were both considered invalid and therefore subsumed under dorylinae, as treated by brady et al. (2014). reference collections are housed at the university of venda and csiro tropical ecosystems research centre (darwin, australia). data analysis chao 2 and ice, in estimates version 9.1.0 (colwell 2013), were used to estimate species richness. sample coverage at each site and the transect as a whole was done with sample-based rarefaction and extrapolation of the incidence matrix. sample coverage represents the proportion of the total number of individuals in the ant assemblage along the transect that belong to the species represented in our sample (chao & jost 2012). both interpolation and extrapolation of samples was based on the bernoulli product model as well as unconditional variance estimates for the rarefaction (interpolation) (chao & jost 2012) and extrapolation curves. this sample-based estimation provides both unconditional variance abundance data of this study and species richness predicted by a 1000 pitfall samples and were considered to be a fair reflection of the richness at a site. samples within each replicate were pooled over the period of the study for multivariate analysis. ant assemblage structure was mapped using non-metric multidimensional scaling of bray–curtis similarity measures (clarke & gorley 2006). a posteriori groups of ant assemblages were defined based on hierarchical clustering with group average linking. support for these groups was tested with the permutational procedure simprof in primer version 6.1.6 (clarke & gorley 2006). results top ↑ in total, 85 759 ant specimens were collected during the 13 sampling periods (appendix 2), comprising 133 species in 38 genera and 6 subfamilies. myrmicinae (with 72 species, 86% of the total abundance and 15 genera) was the most diverse and abundant subfamily, followed by formicinae (27 species and 6 genera). ponerinae (12 genera) had the second highest generic diversity. the most specious genera were tetramorium (22 species), camponotus (19 species), monomorium (13 species), pheidole (9 species), crematogaster (6 species), meranoplus (4 species), and ocymyrmex, anochectus, bothroponera and plectroctena, each with 3 species (appendix 2). sample-based species rarefaction and extrapolation curves for the transect as a whole approximated an asymptote, indicating that most of the epigeal species were collected with a sample coverage of 0.98. sample coverage for all sites was > 0.95, while the forest site 12s coverage was the lowest, at 0.955. based on richness estimations (table 1) and species richness extrapolations (figures 4 and 5), ant richness peaked at mid-elevations on the northern aspect. only at woodland site 14n was species richness significantly greater that of 17n, the summit (figure 4). figure 4: extrapolated species richness for each site and 95% confidence intervals based on unconditional variance, from 1000 pitfall samples for each of the 11 sites across the soutpansberg mountain range. figure 5: species accumulation curves and their 95% confidence intervals for 1000 samples per site. the solid circles represent reference samples; solid lines are rarefied samples and dashed lines are extrapolation based on mau tau estimation. (a) northern aspect, (b) higher elevational sites and (c) low to mid-elevation southern aspect. sbt, accumulation curve for the whole transect. table 1: observed number of species, abundance, richness estimates (chao 2 and ice, available in estimates 9.1.0 package) and sample coverage for broad structural vegetation types and 11 sites with their geographical coordinates along the soutpansberg transect. the pattern on the southern aspect was more complex, with richness decreasing significantly from shrubland site 9s to forest site 12s (figure 4) and then increasing at sedgeland sites 14s and 16s before decreasing again towards the summit (figure 4). based on predicted richness of sites (figure 5), the ranking of sites changes only for 14n. notably, the predicted curve for 16s also suggests a considerable increase in richness with increased sampling. the sites with the highest and lowest variation in mean annual temperature variation (17n and 12s) also had the lowest species richness. in total, 104 species were collected along the northern slope and 117 species on the southern slope. twenty-eight species were restricted to the southern slope and nineteen species to the northern slope. ten species, camponotus uovc_18 (maculatus gp.), lepisiota uovc_01 (capensis gp.), monomorium albopilosum emery, monomorium uovc_07, monomorium_08 (monomorium gp.), pheidole uovc_01 (megacephala gp.), tetramorium cf. setigerum mayri, tetramorium uovc_01 (squaminode gp.), tetramorium uovc_08 (simillimum gp.) and technomyrmex pallipes smith, occurred at all the sites (appendix 2). a total of 20 ponerine species were sampled from both the southern and the northern aspects. fifty per cent of all ponerine species, namely anochectus uovc_01, bothroponera ?strigulosa emery, bothroponera uovc_02, bothroponera uovc_03, hypoponera uovc_01, leptogenys intermedia emery, leptogenys schwabi forel, ophthalmopone uovc_01, plectroctena uovc_02 and pseudoponera uovc_01, were restricted to the southern slope of the mountain. the other 10 species occurred on both slopes of the mountain. the non-metric multidimensional scaling plot identified three major groupings: the high elevation sedgeland-herbland sites (14s – summit); woodlands (08n – 14n); and shrubland, thicket and forest sites (09s – 12s) (figure 6). at bray–curtis similarity measures of 50, each of the 3 groups divides into 2 (figure 6). the groupings reveal distinct assemblages associated with the summit (17n) at higher elevations, while 14n was distinct from all the lower elevation sites on the northern aspect. low elevation (shrubland and thicket sites 09s – 10s) and mid-elevation (forest sites 12s2 – 12s) of the southern aspect formed a distinct grouping. figure 6: non-metric multidimensional scaling of ant assemblages for samples pooled within each of the 44 replicates across 11 sampling sites in the soutpansberg mountain range. discussion top ↑ the level and higher taxon diversity of this study compares favourably with that of similar studies in the cederberg and drakensberg mountains, both of which employ a protocol similar to this study. the cederberg transect located in the cape floristic kingdom includes 17 sites over a distance of 160 km. that study recorded 86 species in 24 genera (botes et al. 2006). the transect in the grasslands of the drakensberg, with 9 elevational sites over a distance of 104 km along the sani pass, collected 98 species in 28 genera and 7 subfamilies (bishop et al. 2014). only one other study of ants along an altitude exists for the savanna biome in south africa, where schoeman & foord (2012) found 104 species in 29 genera along a 1000 m elevational range in the marakele national park. with a higher generic plant diversity than the cape floristic kingdom (hahn 2006) and 90% of all spider families found in the savanna biome (foord et al. 2011), the soutpansberg is characterised by rich, higher taxon diversity. myrmicinae and formicinae were the most diverse subfamilies. the most speciose genera were tetramorium, camponotus, pheidole and monomorium. this conforms to results from other studies in south africa (botes et al. 2006). monomorium is very diverse in australia, while camponotus and pheidole are the most diverse in brazil (campos et al. 2011), although the highly diverse genus in australia, polyrhachis, is only represented by two species along this transect. except for lepisiota, genera found throughout the transect belong to cosmopolitan genera (camponotus, monomorium, pheidole and tetramorium). most of the species accumulation curves of predicted richness based on 1000 pitfall samples reached an asymptote. there were two exceptions: a woodland site 14n and sedgeland site 16s, both of which also had the lowest coverage. although 14n is in the woodland vegetation type, it is near the ecotone between sedgeland and woodland. the large turnover of taxa at this site could be the result of mass effects from taxa found in sedgeland. the steep slope of the 16s accumulation curve might suggest a temporal change in the ant assemblages of this site. based on photographic records at each of the pitfalls, the coleochloa species that dominate this site are being invaded by short woody shrubs. turnover at this site might be linked to this encroachment and could probably lead to increased abundance of taxa associated with sites that are structurally more complex, such as forest sites 12s and 12s2 and shrubland site 9s. this has particularly important implications for the management of the few remaining grassland habitats in the soutpansberg. the high diversity of ponerinae species at the base of southern aspect might be due to more productive ecosystems associated with the basalt intrusions, and higher rainfall. disturbance also seems to positively impact ponerine richness, which peaks in shrubland site 9s. this site is inside a wildlife sanctuary that is stocked with buffalo and where the bush is mechanically cleared. the three major groupings of ant assemblages across the study area conform to the broad structural classification of the vegetation used in this study (edwards 1983). conclusion top ↑ this study provides a thorough inventory of ground-dwelling ants, with a sample coverage of 0.98. the sample coverage represents the proportion of the total number of individuals in the ant assemblage along the transect that belongs to the species represented in our sample (chao & jost 2012). extrapolated richness for most sites reached an asymptote, except for three sites. all three of these sites can be considered to be sites that have experienced increased levels of bush encroachment. this is particularly concerning at 16s, which represents some of the last remaining grassland habitats in the soutpansberg. ant taxa that act as indicators of bush encroachment should be identified in order to monitor the rate of these impacts. a single site on the lowest elevation of the southern aspect (09s) yielded 57% of the total ant species. this site experiences higher levels of disturbance through herbivory and bush clearing and provides support for the important role of disturbance as a driver of ant diversity in the savanna biome. the structural classification of vegetation in this study is a good predictor of broad-scale ant assemblage structure. finer scale distinctions were linked to elevation, structural elements and temperature. acknowledgments top ↑ special thanks to alan andersen and hendrick sithole for verifying ant identifications. we are grateful to ian gaigher (lajuma research centre), steven fick (koedoesvlei farm) and dave dewsnap (goro nature reserve) for providing access to their farms. several volunteers from the university of venda and lajuma research centre are thanked for their assistance in the field. this study was funded by the dst-nrf center of excellence for invasion biology and the university of venda. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions t.c.m. (university of venda) and s.h.f. (university of venda) were responsible for the project design. t.c.m. identified the specimens. t.c.m. and s.h.f. performed the analysis and wrote the manuscript. references top ↑ abensperg-traun, m. & steven, d., 1995, ‘the effects of pitfall trap diameter on ant species richness (hymenoptera: formicidae) and species composition of the catch in a semi-arid eucalypt woodland’, journal of ecology 20(2), 282–287. andersen, a.n. & majer, j.d., 2004, ‘ants show the way down under: invertebrates as bioindicators in land management’, frontiers in ecology and the environment 2(6), 291–298. http://dx.doi.org/10.1890/1540-9295(2004)002[0292:astwdu]2.0.co;2 bishop, t.r., robertson, m.p., van rensburg, b.j. & parr, c.l., 2014, ‘elevation–diversity patterns through space and time: ant communities of the maloti-drakensberg mountains of southern africa’, journal of biogeography 41, 2256–2268. http://dx.doi.org/10.1111/jbi.12368 bolton, b., 1994, identification guide to the ant genera of the world, harvard university press, cambridge, massachusetts. botes, a., mcgeoch, m.a., robertson, h.g., van niekerk, a., davids, h.p. & chown, s.l., 2006, ‘ants, altitude and change in the northern cape floristic region’, journal of biogeography 33, 71–90. http://dx.doi.org/10.1111/j.1365-2699.2005.01336.x brady, s.g., fisher, b.l., schultz, t.r. & ward, p.s., 2014, ‘the rise of army ants and their relatives: diversification of specialized predatory doryline ants’, bmc evolutionary biology 14(5), 93. http://dx.doi.org/10.1186/1471-2148-14-93 campos, r.i., vasconcelos, h.l., andersen, a.n., frizzo, t.l.m. & spena, k.c., 2011, ‘multi-scale ant diversity in savanna woodlands: an intercontinental comparison’, austral ecology 36, 983–992. http://dx.doi.org/10.1111/j.1442-9993.2011.02255.x cardoso, p., erwin, t.l., borges, p.a.v. & new, t.r., 2011, ‘the seven impediments in invertebrate conservation and how to overcome them’, biological conservation 144, 2647–2655. http://dx.doi.org/10.1016/j.biocon.2011.07.024 chao, a. & jost, l., 2012, ‘coverage-based rarefaction and extrapolation: standardizing samples by completeness rather than size’, ecology 93, 2533–2547. http://dx.doi.org/10.1890/11-1952.1 clarke, k.r. & gorley, r.n., 2006, primer v6: user manual/tutorial. primer-e, plymouth. colwell, r.k., 2013, estimates: statistical estimation of species richness and shared species from samples. version 9. persistent url <purl.ocls.org/estimates>. edwards, d., 1983, ‘a broad-scale structural classification of vegetation for practical purposes’, bothalia 14, 705–712. foord, s.h., 2008, ‘cladistic analysis of the afrotropical hersiliidae (arachnida: araneae) with the first records of murricia and the description of a new genus from madagascar’, journal of afrotropical zoology 4, 111–142. foord, s.h., dippenaar-schoeman, a.s., haddad, c.r., lotz, l.n. & lyle, r., 2011, ‘the faunistic diversity of spiders (arachnida: araneae) of the savanna biome in south africa’, transactions of the royal society of south africa 66, 170–201. http://dx.doi.org/10.1080/0035919x.2011.639406 hahn, n., 2002, ‘endemic flora of the soutpansberg’, msc thesis, faculty of science and agriculture, university of kwazulu-natal. hahn, n., 2006, ‘floristic diversity of the soutpansberg, limpopo province, south africa’, phd thesis, faculty of natural and agricultural sciences, university of pretoria. hölldobler, b. & wilson, e.o., 1990, the ants, harvard university press, cambridge, massachusetts. http://dx.doi.org/10.1007/978-3-662-10306-7 kirchhof, s., kramer, m., linden, j. & richter, k., 2010, ‘the reptile species assemblage of the soutpansberg (limpopo province, south africa) and its characteristics’, salamandra 46, 147–166. munyai, t.c. & foord, s.h., 2012, ‘ants on a mountain: spatial, environmental and habitat associations along an altitudinal transect in a centre of endemism’, journal of insect conservation 16(5), 677–695. http://dx.doi.org/10.1007/s10841-011-9449-9 ness, j.h., bronstein, j.l., andersen, a.n. & holland, j.n., 2004, ‘ant body size predicts dispersal distance of ant-adapted seeds: implications of small-ant invasions’, ecology 85, 1244–1250. http://dx.doi.org/10.1890/03-0364 schmidt, c.a. & shattuck, s.o., 2014, ‘the higher classification of the ant subfamily ponerinae (hymenoptera: formicidae), with a review of ponerine ecology and behavior’, zootaxa 3817(1), 001–242. http://dx.doi.org/10.11646/zootaxa.3817.1.1 schoeman, c.s. & foord, s.h., 2012, ‘a checklist of epigaeic ants from marakele national park, limpopo province, south africa (hymenoptera: formicidae)’, koedoe 54(1), 1–7. http://dx.doi.org/10.4102/koedoe.v54i1.1030 tshiala, m.f., olwoch, j.m. & engelbrecht, f.a., 2011, ‘analysis of temperature trends over limpopo province, south africa’, journal of geography and geology 3, 13–21. http://dx.doi.org/10.5539/jgg.v3n1p13 van wyk, a.e. & smith, g.f., 2001, regions of floristic endemism in southern africa, umdaus press, pretoria. ward, p.s., 2010, ‘taxonomy, phylogenetics, and evolution’, in l. lach, c.l. parr & k.l. abbott (eds.), ant ecology, pp. 3–17, oxford university press, new york. wielgoss, a., tscharntke, t., rumede, a., fiala, b., seidel, h., shahabuddin, s. et al., 2014, ‘interaction complexity matters: disentangling services and disservices of ant communities driving yield in tropical agroecosystems’, proceedings of the royal society b: biological sciences 281, 2013–2144. appendix 1 top ↑ table a1: summary of the number of replicates set out at each of the 11 sites during each of the 13 surveys along the soutpansberg mountain range. a replicate within a site consisted of 10 pitfalls in a 2 × 5 grid. pitfalls were separated by 10 m and replicates within a site were separated by at least 300 m to ensure independence. appendix 2 top ↑ table a2: checklist of subfamilies and ant species collected during 13 surveys in different broad structural vegetation types across an elevational transect in the western soutpansberg mountain range in south africa. appendix 3 top ↑ figure a1: images of some notable and interesting species collected during 13 surveys: (a) introduced species ponera uovc_01, (b) camponotus fulvopilosus de geer, (c) ocymyrmex flaviventris santschi and (d) platythyrea lamellosa roger. reviewer acknowledgement http://www.koedoe.co.za open access koedoe african protected area conservation and science koedoe recognises and acknowledges the value and importance of reviewers in the publication process – not only in shaping each individual manuscript, but also in upholding the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely and productive peer-review process. we would like to take this opportunity to thank all reviewers who participated in shaping this volume of koedoe: page 1 of 1 in an effort to facilitate the selection of appropriate peer reviewers for koedoe, we ask that you take a moment to update your electronic portfolio at http://www.koedoe. co.za for our records. this will allow us better access to your areas of interest and expertise, in order to match reviewers with submitted manuscripts. if you would like to become a reviewer, please visit the journal website and register as a reviewer. to access your details on the website, you will need to follow these steps: 1. log into the online journal at http://www. koedoe.co.za 2. in your ‘user home’ [http://www.koedoe. co.za/index.php/koedoe/ user] select ‘edit my profile’ under the heading ‘my account’ and insert all relevant details, bio statement and reviewing interest. 3. it is good practice as a reviewer to update your personal details regularly, to ensure contact with you throughout your professional term as reviewer to koedoe. please do not hesitate to contact me if you require assistance in performing this task. margo van blerk submissions@koedoe.co.za tel: +27 (0)21 975 2602 fax: +27 (0)21 975 4635 abraham nkhata albert strydom andy drumm anna spenceley art pedersen cornelius van heerden elmarie slabbert engela de crom glen hvenegaard helen suich jacob mwitwa jasmine moreira jeff marion jon kohl keith bosak laura becerra lisa king marcella daye peet van der merwe rick rollins steven martin sue snyman susan moore willie boonzaaier willie coetzee willie hood yi-chung hsu yu-fai leung 86 article information authors: mamokete n.v. dingaan1,2 pieter j. du preez1 affiliations: 1department of plant sciences, university of the free state, south africa2department of soil, crop and climate sciences, university of the free state, south africa correspondence to: mamokete dingaan postal address: po box 339, bloemfontein 9300, south africa dates: received: 15 feb. 2012 accepted: 20 sept. 2012 published: 14 mar. 2013 how to cite this article: dingaan, m.n.v. & du preez, p.j., 2013, ‘grassland communities of urban open spaces in bloemfontein, free state, south africa’, koedoe 55(1), art. #1075, 8 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1075 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/ koedoe.v55i1.1075-1 copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. grassland communities of urban open spaces in bloemfontein, free state, south africa in this original research... open access • abstract • introduction • research method and design    • study area    • vegetation sampling and data analysis • results    • vegetation classification    • vegetation description       • 1. felicia muricata – themeda triandra major community       • 1.1. panicum coloratum – digitaria eriantha community       • 1.1.1. cyperus usitatus – digitaria eriantha sub-community       • 1.2. eragrostis obtusa – sporobolus fimbriatus community       • 1.2.1. panicum schinzii – themeda triandra sub-community       • 1.3. eragrostis biflora – themeda triandra community       • 2. aristida congesta – themeda triandra major community       • 2.1. trichoneura grandiglumis – rhynchosia nervosa community       • 2.1.1. conyza bonariensis – eragrostis curvula sub-community       • 2.1.2. antephora pubescens – digitaria argyrograpta sub-community       • 2.2. eragrostis trichophora – cyperus capensis community       • 2.3. hyparrhenia hirta – themeda triandra community       • 2.4. melinis repens – eragrostis lehmanniana community    • ordination • trustworthiness    • reliability    • validity • discussion • conclusion • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ natural vegetation in urban environments is greatly impacted by human activities and it is in constant threat of degradation and destruction as a result of urbanisation. this vegetation, although fragmented, serves an important ecological function and needs to be properly managed and conserved. studies on urban vegetation are lacking in south africa, with only a handful having been carried out since the end of the last century. this study was initiated to identify, classify and describe the grassland communities of the urban open spaces in bloemfontein. relevés were compiled in 61 sample plots, where species present and habitat information were recorded. care was taken to restrict sample plots to vegetation in pristine condition, wherever possible, and severely degraded stands were avoided. a two-way indicator species analysis (twinspan) classification, refined by braun-blanquet procedures, revealed two distinct major communities, seven communities and four sub-communities. both detrended and canonical correspondence analyses indicated the vegetation units to be associated with soil texture and ph, although biotic factors such as overgrazing, burning and mowing also influence the composition of the vegetation. the proper management and conservation of urban open spaces requires in-depth knowledge of the spatial distribution, floristic, structural and functional compositions within the major vegetation types in this environment. the present study further contributed towards formulating ways for the proper management, utilisation and functioning of the open spaces within the bloemfontein area.conservation implications: the grassland biome of south africa is poorly conserved, mainly because of its status as an agricultural hub of the country. the preservation of natural and semi-natural forms of urban vegetation is important because such vegetation, although often disturbed and degraded, could form dispersal corridors between peri-urban and rural vegetation. introduction top ↑ bloemfontein lies within the grassland biome (rutherford & westfall 1994) and is classified by acocks (1988) as part of the dry cymbopogon – themeda veld (vt 50). here themeda triandra is ecologically the most important grass, both to wild and domestic animals, and is the most widely distributed of the climax grasses. one of the key environmental variables affecting the functioning of the grassland ecosystem globally is rainfall variability (tilman & el haddi 1992). bloemfontein is situated in a semi-arid region, where precipitation is low and highly variable (noy-meir 1973; sala & lauenroth 1982). although semi-arid regions typically receive substantial precipitation for at least a few months of the year, lack of adequate rainfall for many months may also prevail (bailey 1979).the grassland biome of south africa is an important agricultural region, especially for the intensive production of crops such as maize and wheat (department of agriculture, forestry and fisheries 2010). it is also important for extensive stock farming; mainly for dairy, beef and wool production. however, the intensive crop production and livestock grazing pressure have resulted in the destruction or degradation of large portions of pristine vegetation in the biome. in addition, natural and semi-natural vegetation is constantly impacted by urban development and, according to low and rebelo (1996), many of the natural areas in the biome have been lost as a result of urbanisation. in recognition of the poor conservation status of the biome, cohen and hugo (1986) emphasise the importance of conservation outside officially designated nature reserves, such as in urban open spaces. in the vicinity of bloemfontein, as in many areas of the biome, natural vegetation is represented by small fragmented areas. this fragmentation of previously intact natural environments may cause the potential loss of habitat for some species, the isolation of other species on natural remnants within the urban environment and even the total extinction of certain plant species (wood et al. 1994). studies of urban vegetation in south africa are lacking and only a limited number of vegetation surveys have been carried out in these areas: for example, in durban, kwazulu-natal province (roberts 1993); potchefstroom, north west province (cilliers, van wyk & bredenkamp 1999); klerksdorp, north west province (van wyk, cilliers & bredenkamp 2000); and pretoria, midrand, johannesburg and parts of the west rand, gauteng province (grobler, bredenkamp & brown 2006). the aim of this study was therefore to identify, classify and describe the grassland communities of the natural open areas in bloemfontein. research method and design top ↑ study area the study was carried out in natural open spaces within the bloemfontein metropolitan area, which include the inner city area, as well as the surrounding farms and smallholdings. the study area extends from approximately 29°00′s to 29°15′s and 26°07′e to 26°21′e, at an altitude of 1350 m a.s.l. – 1450 m a.s.l.there are various large open spaces in the area, but, for this study, special emphasis was placed on the natural and semi-natural areas. these areas are fragmented and are most prominent towards the periphery of the city. they comprise numerous hills and ridges, areas along drainage lines and uncultivated patches on smallholdings and farms. other open spaces studied were vacant residential lots and the yet-to-be-developed areas as indicated in the spatial development framework of the mangaung metropolitan municipality (2011). green belts adjacent to railway lines and main roads were also included in the study because they form part of the natural open space system within the bloemfontein area, but the open spaces in the inner city area, such as parks and sports grounds, were excluded from the study as they were fully transformed. according to the köppen climate classification system, bloemfontein falls under the bsk climate zone (schulze 1947). this is a steppe climate in which the winters are dry and the mean annual temperature is below 18.0 °c (le houérou, poppov & see 1993; schulze & mcgee 1978). maximum temperatures are recorded in december and january, averaging 30.1 °c and 30.6 °c, respectively (figure 1). below zero temperatures are often recorded for june (-1.0 °c) and july (-1.2 °c). the annual mean maximum temperature is 24.6 °c and the annual mean minimum temperature is 7.6 °c, with the average annual temperature of 16.1 °c. rain mainly falls during the summer months, often in the form of thunderstorms. the average annual rainfall is 550 mm. figure 1: long-term (1951.2000) meteorological data of the bloemfontein airport (29°06′s, 26°18′e, 1351 m a.s.l.), sourced from the south african weather service. three different land types are distinguished in the study area, namely the ca, dc and ea land types. within each of these land types, soils vary from sandy to clayey, as a result of the variation in parent material. the ca land type, occurring as ca8 and ca22 subdivisions, is a mixture of duplex and plinthic soils. the ca8 subdivision is found on the western part of the study area and consists mainly of sandy hutton–bainsvlei soils and more clayey valsrivier–swartland soils; the ca22 subdivision occurs in the south and is mainly dominated by valsrivier soils. the dc land type occurs as subdivision dc13, which is dominated by duplex soils of the valsrivier–swartland–sterkspruit forms. it is only present in small pockets in the eastern part of the study area. the ea land type is found in the central and northern parts and occurs as subdivision ea39, the most extensive in the study area. it consists mainly of sandy oakleaf soils, although a mixture of the clayey milkwood, arcadia and valsrivier soils are also prominent. the dominant soil profiles are described according to land type survey staff (1993) and the soil forms and series are used according to the soil classification working group (1991).the geologic formations of the study area belong to the karoo supergroup and consist of the tierberg formation of the ecca group and the adelaide sub-group of the beaufort group, as well as dolerite intrusions of the post-karoo age (johnson et al. 2006). vegetation sampling and data analysis relevés were compiled in 61 sample plots, each fixed at 16 m2. care was taken to restrict sample plots to vegetation in pristine condition, wherever possible, and severely degraded stands were avoided. in each sample plot, all vascular plants present were recorded and all plant nomenclature was used according to germishuizen and meyer (2003). the cover-abundance values of all species present were allocated according to the braun-blanquet scale, as given by mueller-dombois and ellenberg (1974), as well as kent and coker (1996). habitat parameters recorded included soil depth, rockiness of the soil surface and the presence of biotic influences such as trampling and grazing. a total of 25 soil samples was collected and analysed for ph, organic matter and texture. no soil samples were collected at one part of the study area, the kwaggafontein hills, as this is a conservation area and no permission was granted to collect such samples.the vegetation and habitat data were first captured into the turboveg database (hennekens 1996a) and then imported to megatab (hennekens 1996b), a visual editor for phytosociological tables, where the first approximation of the vegetation classification was obtained by the application of the two-way indicator species analysis (twinspan) (hill 1979a). further refinement of the initial classification results was subsequently conducted within the megatab programme, resulting in a phytosociological table. the relationships between the plant communities and environmental variables were determined by an ordination algorithm, detrended correspondence analysis (dca) (hill 1979b) and also by an extension of dca, the canonical correspondence analysis (cca) (ter braak & šmilauer 2009). results top ↑ vegetation classification the grassland communities of the bloemfontein area are classified as bloemfontein dry grassland by mucina et al. (2006) the vegetation is dominated by t. triandra, a companion species with a wide ecological amplitude and eragrostis lehmanniana as a typical associate. other constantly present grasses are eragrostis curvula and digitaria eriantha. the forbs oxalis depressa, nidorella resedifolia and hibiscus pusillus are of widespread occurrence, although they are hardly prominent (see online appendix). the habitats occupied by each of the communities are, by and large, distinguished by differences in soil depth, texture and ph (table 1). biotic factors, especially livestock grazing practices, also influence the composition of the communities. table 1: soil characteristics of the grassland communities of the bloemfontein area (mean values). the classification resulted in two major communities, which are grouped into seven communities and four sub-communities. the hierarchical classification of the plant communities is as follows:1. felicia muricata – themeda triandra major community 1.1. panicum coloratum – digitaria eriantha community 1.1.1. cyperus usitatus – digitaria eriantha sub-community 1.2. eragrostis obtusa – sporobolus fimbriatus community 1.2.1. panicum schinzii – themeda triandra sub-community 1.3. eragrostis biflora – themeda triandra community 2. aristida congesta – themeda triandra major community 2.1. trichoneura grandiglumis – rhynchosia nervosa community 2.1.1. conyza bonariensis – eragrostis curvula subcommunity 2.1.2. antephora pubescens – digitaria argyrograpta subcommunity 2.2. eragrostis trichophora – cyperus capensis community 2.3. hyparrhenia hirta – themeda triandra community 2.4. melinis repens – eragrostis lehmanniana community. vegetation description 1. felicia muricata – themeda triandra major community this widespread grassland community is mainly associated with low-lying, flat open plains where the soils are generally sandy clay loam, with the clay content ranging between 18.5% and 27.0% (table 1). the vegetation has a uniform representation of grasses, with forbs and rarely shrubs interspersed between them. the two climax grasses t. triandra and d. eriantha (species group n) have a characteristically high presence, indicating good veld condition. this is further indicated by the notable poor occurrence of the pioneer grass a. congesta and the sub-climax grass eragrostis superba (species group g), which are known to occur in disturbed veld. other well represented species include eragrostis chloromelas, f. muricata (species group a) and e. lehmanniana (species group n). this major community has six diagnostic species: namely, indigofera alternans, pentzia globosa, tripteris aghillana, schkuhria pinnata, e. chloromelas and f. muricata (species group a). three communities are recognised within this major community. 1.1. panicum coloratum – digitaria eriantha community this grassland community is situated on flat plains on the city margins and also on fragmented natural areas between and within residential areas. it is differentiated and dominated by the grass species p. coloratum and eragrostis plana (species group b). other equally prominent grasses are e. chloromelas (species group a), t. triandra, e. lehmanniana, aristida adscensionis and d. eriantha (species group n). this community has a low number of species, with an average of only 11 recorded per sample plot. one sub-community is recognised within the community, which is differentiated by its relatively high soil moisture. 1.1.1. cyperus usitatus – digitaria eriantha sub-community this grassland community can be found occurring on flat open plains on the southern part of the study area. it is differentiated by species from group c, namely c. usitatus and anthospermum rigidum subsp. pumilum. this sub-community is dominated entirely by grasses, with only a handful of forbs showing a rare occurrence. the most prominent grasses are e. chloromelas (species group a), p. coloratum (species group b), t. triandra, a. adscensionis and d. eriantha (species group n). the only forb with a strong presence is c. usitatus (species group c), whilst t. aghillana (species group a) and o. depressa (species group n) are constantly present, but never prominent. an average of 13 species per sample plot was recorded. 1.2. eragrostis obtusa – sporobolus fimbriatus community this community is found in pasture areas in the south of bloemfontein, as well as towards the west of the city. it is dominated entirely by grasses, most notably eragrostis obtusa and s. fimbriatus, which are also the diagnostic species (species group d). other equally dominant grasses are t. triandra and d. eriantha (species group n), with f. muricata (species group a), h. pusillus and selago densiflora (species group n) found occurring consistently within the community, but never prominently. an average of 13 species per sample plot was recorded, and one sub-community is recognised within this community. 1.2.1. panicum schinzii – themeda triandra sub-community this is a sub-community found on the grassland plains and footslopes of the gently sloping small hills of the southern part of the study area. grasses are dominant, the most prominent being e. obtusa, s. fimbriatus (species group d), p. schinzii (species group e), t. triandra, e. lehmanniana, e. curvula and d. eriantha (species group n). the only forbs of prominent occurrence are f. muricata (species group a), conyza podocephala (species group e) and helichrysum dregeanum (species group n). the diagnostic species for the sub-community are p. schinzii, argemone ochroleuca, monsonia angustifolia, c. podocephala, chloris virgata and nenax microphylla (species group e). an average of 14 species per sample plot was recorded. 1.3. eragrostis biflora – themeda triandra community this community is associated with open grasslands on the periphery of the western suburbs of bloemfontein. the diagnostic species are the shade-loving grass eragrostis biflora, the erect perennial herb cyperus obtusiflorus, the karroid shrub lycium cinereum and the highly palatable grass digitaria tricholaenoides (species group f). the grassland is dominated by i. alternans, p. globosa (species group a), e. biflora (species group f), t. triandra and e. lehmanniana (species group n). this community has the lowest species richness of all the grassland communities of the bloemfontein area, with an average of only eight species per sample plot recorded. 2. aristida congesta – themeda triandra major community this widespread community represents grasslands that are characterised by sandy loam soils. the sand content ranges between 67.1% and 85.6% (table 1). it is mainly found in the northern and western homesteads and farming areas and, to a smaller extent, on the smallholdings to the east of bloemfontein. the most dominant species in the community is t. triandra (species group n), with other grasses such as a. congesta, e. superba (species group g) and e. lehmanniana (species group n) also prominent. the diagnostic species for the community are a. congesta and e. superba (species group g), grasses which are also indicators of veld disturbance. four communities are recognised within this major grassland. 2.1. trichoneura grandiglumis – rhynchosia nervosa community this is a community situated on the western part of the study area. it occurs on the sides of the kwaggafontein hills and also on the grass plains at the tempe airfield. seven diagnostic species are identified for the community: the grasses t. grandiglumis, pogonarthria squarrosa and heteropogon contortus and the forbs r. nervosa, pollichia campestris, moraea pallida and senecio burchellii (species group h). this community is partially dominated by the highly palatable grasses e. superba (species group g) and t. triandra (species group n) and the less palatable a. congesta (species group g), t. grandiglumis, p. squarrosa, h. contortus (species group h) and e. lehmanniana (species group n). other constantly present species are r. nervosa (species group h), o. depressa, s. densiflora and h. dregeanum (species group n). two sub-communities are recognised within this grassland community. 2.1.1. conyza bonariensis – eragrostis curvula sub-community this is a sub-community occurring on the north-western part of the study area, particularly at the tempe airfield and the neighbouring skydiving ground. the perennial grasses a. congesta (species group g), t. grandiglumis (species group h), t. triandra and e. curvula (species group n) are dominant. other species of notable occurrence include r. nervosa (species group h), c. bonariensis (species group i) and s. densiflora (species group n). the diagnostic species are salvia verbenaca, aristida canescens and c. bonariensis (species group i), whilst the strong presence of e. curvula and d. eriantha (species group n) further distinguishes the sub-community from the a. pubescens – d. argyrograpta sub-community. an average of 13 species per sample plot was recorded. 2.1.2. antephora pubescens – digitaria argyrograpta sub-community this sub-community is situated at the kwaggafontein hills, a conservation area on the western part of the study area. it occurs on the gently undulating sides of the hills, as well as on the adjacent bottomland area on deep, red sandy soils, with little or no surface rock. because this vegetation unit occurs in a conservation area, it is in primary condition and protected from the effects of grazing. it also has the highest species richness, with an average of 30 species per sample plot and the highest number of diagnostic species (20). some diagnostic species include: the grasses a. pubescens, d. argyrograpta, aristida stipitata, tragus koeleroides and cynodon dactylon, the herbs gazania krebsiana, commelina africana, harpagophytum procumbens and dicoma macrocephala and the karroid shrub chrysocoma ciliata (species group j). the vegetation consists of a wide variety of grass species, of which the most dominant is a. pubescens (species group j). its abundance is regarded as an indicator of good veld condition (van oudtshoorn 1999). other prominent grasses are a. congesta and e. superba (species group g), t. grandiglumis, p. squarrosa, and h. contortus (species group h), d. argyrograpta and a. stipitata (species group j), t. triandra and e. lehmanniana (species group n), whilst p. campestris (species group h), s. densiflora and h. dregeanum (species group n) are amongst the dominant forbs. 2.2. eragrostis trichophora – cyperus capensis community this plant community occurs on the smallholdings of the eastern city margin. species from group k differentiate the community: namely, the grasses e. trichophora and eragrotis gummiflua, the sedge c. capensis, the perennial dwarf shrub selago albida and the herb crotalaria sphaerocarpa. these diagnostic species are also the most dominant, with a. congesta (species group g) being the only other spe cies of prominent occurrence. it replaces t. triandra (species group n) in some overgrazed patches. only a limited number of species is present in this community and an average of only 11 species per sample plot was recorded. 2.3. hyparrhenia hirta – themeda triandra community this is a community occurring along the roadsides at the northern margin of the city, where the soil is slightly gravelly. the tall, tufted perennial h. hirta (species group l) is the only diagnostic species and entirely dominates, sometimes growing to heights exceeding 1 m. the other prominent species in this community are the grasses t. triandra, e. lehmanniana and e. curvula (species group n). an average of 11 species per sample plot was recorded. 2.4. melinis repens – eragrostis lehmanniana community this is a slightly disturbed community occurring on roadsides on the western and north-western part of the study area. the soil along the roads where this community is found is, in general, gravelly, at times with small stones also found. this is largely a grass community, with rhus lancea (species group m) the only woody species and t. aghillana (species group a) being the only forb of notable occurrence. the most dominant grasses include m. repens and enneapogon cenchroides (species group m), t. triandra and e. lehmanniana (species group n). e. cenchroides characteristically grows in clumps and, together with m. repens, dominates in disturbed patches, especially where the soils are exceptionally deep. the diagnostic species for this community are m. repens, e. cenchroides, r. lancea and fingerhuthia africana (species group m). an average of 12 species per sample plot was recorded. ordination the dca analysis complements the classification in the online appendix and reveals distinct discontinuities between the communities (figure 2). the cca ordination of sample plots and soil variables (figure 3) illustrates that communities of the f. muricata – t. triandra major community (community 1) occur in soils with relatively high organic matter and clay content and slightly lower ph (i.e. neutral soils). communities of the a. congesta – t. triandra major community (community 2), on the other hand, are associated with more sandy soils that have less organic matter, but slightly higher ph (table 1). there is a correlation between soil texture and organic matter content; organic matter is positively correlated with the clay content (r = 0.757; p < 0.001) and negatively correlated with the sand content (r = 0.666; p < 0.001). figure 2: a detrended correspondence analysis ordination of the grassland vegetation of bloemfontein, showing the relative positions of the relevés along the first two axes (axis 1 and axis 2). figure 3: a canonical correspondence analysis biplot of 25 sample relevés and soil environment variables for the first two axes: axis 1 (horizontal) and axis 2 (vertical). the biplot of the species and soil environmental variables (figure 4) further supports the assertion that species of the f. muricata – t. triandra major community (species groups a, b, c, d, e) have a strong association with higher clay content and organic matter in the soil. species from group f, together with the character species of the communities of a. congesta – t. triandra major community (species groups g, h, i, k) are strongly associated with the soil depth and sand gradients. in contrast, the other character species of the a. congesta – t. triandra major community, namely h. hirta (species group l), m. repens, e. cenchroides and f. africana (species group m) are strongly associated with the ph gradient. these species are commonly found on gravelly soil (van oudtshoorn 1999) and, in the study area, they are often located along roadsides and in road reserves. the only woody character species, r. lancea (species group m), is also strongly associated with the ph gradient and it often prefers calcareous substrates (van wyk & van wyk 1997). figure 4: a canonical correspondence analysis biplot of 59 plant species and soil environment variables for the first two axes: axis 1 (horizontal) and axis 2 (vertical). trustworthiness top ↑ the classification of the grassland communities of bloemfontein is based on data acquired through human observations and requires consideration of the trustworthiness of the data in the contexts below. reliability the nature of this study does not require repeated surveys and therefore the authors were able to satisfactorily draw conclusions from their research findings because they believe they have utilised the most widely used and arguably the most efficient of all types of vegetation sampling techniques. a potential bias, however, can arise during the vegetation classification but this is compensated for by the use of computer programmes such as twinspan, decorana and canoco. validity the authors believe the methodologies employed were successful at achieving the objectives of the study. discussion top ↑ the classification of the grasslands of the urban open spaces in bloemfontein revealed the existence of seven identifiable plant communities and four sub-communities, all represented within two major communities. the vegetation can be broadly classified as the t. triandra – e. lehmanniana grassland, where t. triandra is the dominant sweet grass. other major grasses include e. lehmanniana, e. curvula, d. eriantha and e. chloromelas. out of a total of 119 species recorded for the study area, 64 play a diagnostic role, 10 are companion species and a further 45 are either localised or of very rare occurrence. the distribution and composition of the plant communities are linked with habitat factors such as soil depth and texture, rockiness of the soil surface and habitat disturbance (especially utilisation by livestock).the results of the soil analysis indicate a correlation between soil texture and organic matter content, whereby organic matter is positively correlated with the clay content and negatively correlated with the sand content. these correlations can be explained in two ways. firstly, soils with high clay content have a high water holding capacity and therefore high moisture content. communities in such soils are, as a result, more productive; hence the high organic matter content in soil. secondly, according to foth (1990), clay plays a role in protecting organic matter decomposition, thus contributing towards the high organic matter content in the f. muricata – t. triandra major community. communities of the a. congesta – t. triandra major community, on the other hand, occupy areas of higher sand content that are characterised by low soil moisture content as a result of low water holding capacity. these communities are therefore less productive and hence have less organic matter. the distribution of species commonly found on gravelly soil along roadsides and in road reserves, such as h. hirta, m. repens, e. cenchroides and f. africana is strongly affected by ph. the soils near and along roads are known to have higher ph levels; this is attributed to calcareous road dust and leachate (auerbach, walker & walker 1997; johnston & johnston 2004). communities occurring on pasture land in farming areas show notable signs of overgrazing and trampling. there is also a tendency of selective grazing by livestock, through which animals over-utilise the palatable grasses. this results in the replacement of nutritious perennial grasses by annual grasses and weeds of inferior value. the invasion of the veld by the unpalatable species can have serious ecological implications because most of these invaders are a threat to the indigenous species. it is therefore necessary to manage the vegetation according to ecological principles in order to maintain optimum floristic composition and diversity. furthermore, there is a common practice of mowing or burning vegetation that grows along roads and railway tracks in the vicinity of residential and commercial areas. this vegetation, though disturbed and unstable, is vital simply because most of the natural vegetation in urban areas occurs in fragmented forms as a result of urbanisation (florgård 2007). the disturbed areas could therefore form dispersal corridors between the natural fragments and, in this way, act as stepping-stones for a variety of plant species (davis & glick 1978). the preservation of these disturbed areas may, in the end, offset the biological isolation of the natural areas and, in so doing, provide ecological continuity throughout the urban environment (millard 2004; poynton & roberts 1985). conclusion top ↑ the distribution and composition of the grassland communities of the bloemfontein area are linked with habitat factors such as soil depth and texture, with the f. muricata – t. triandra major community associated with clayey soils and the a. congesta – t. triandra major community found on sandy soils. the communities are also impacted by habitat disturbance, mainly in the form of overgrazing, mowing and burning. this phenomenon is not unique to these grasslands, because large portions of pristine vegetation in the grassland biome of south africa have been degraded and destroyed due to livestock grazing pressure and the ploughing of arable land. the proper management and conservation of urban open spaces requires in-depth knowledge of the spatial distribution, floristic, structural and functional compositions within the major vegetation types in the urban environment. the present vegetation study was initiated to contribute towards formulating ways for the proper functioning and management of the open spaces within the bloemfontein metropolitan area. the benefits of conserving urban vegetation are immense; scientifically, socially, and economically. open space within urban areas has value upon urban microclimate, hydrology, biodiversity, and ecological processes (nicholson-lord 1987; platt, rowntree & muick 1994). this phytosociological study provides valuable information on the structure and composition of the vegetation of the bloemfontein metropolitan area, which can be utilised by the relevant local and national authorities for environmental planning and conservation strategies for the area. acknowledgements top ↑ the assistance of the following persons is sincerely appreciated: dr pieter le roux (university of the free state) for his help in interpreting the soils and land types of the study area, ms yvonne dessels and the soil science laboratory staff of the university of the free state for the soil analysis, as well as the two anonymous reviewers for their useful comments and suggestions. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions m.n.v.d. (university of the free state) conducted the vegetation survey, data analysis and wrote the manuscript, p.j.d.p. (university of the free state) conceptualised the project, conducted part of the vegetation survey and assisted with the vegetation classification. references top ↑ acocks, j.p.h., 1988, ‘veld types of south africa’, 3rd edn., memoirs of the botanical survey of south africa 57, 1–146.auerbach, n.a., walker, m.d. & walker, d.a., 1997, ‘effects of roadside disturbance on substrate and vegetation properties in arctic tundra’, ecological applications 7(1), 218–235. http://dx.doi.org/10.1890/1051-0761(1997)007[0218:eordos]2.0.co;2 bailey, h.p. , 1979, ‘semi-arid climates: their definition and distribution’, in a.e. hall, g.h. cannell & h.w. lawton (eds.), agriculture in semi-arid environments, pp. 73–97, springer-verlag, berlin. http://dx.doi.org/10.1007/978-3-642-67328-3_3 cilliers, s.s., van wyk, e. & bredenkamp, g.j., 1999, ‘urban nature conservation: vegetation of natural areas in the potchefstroom municipal area, north west province, south africa’, koedoe 42, 1–30. http://dx.doi.org/10.4102/koedoe.v42i1.218 cohen, m. & hugo, m.l., 1986, ‘conservation outside of officially designated areas – with emphasis on the urban environment’, parks and grounds 38, 15–23. davis, a.m. & glick, t.f., 1978, ‘urban ecosystems and island biogeography’, environmental conservation 5, 299–304. http://dx.doi.org/10.1017/s037689290000638x department of agriculture, forestry and fisheries, 2010, abstract of agricultural statistics, daff directorate, agricultural statistics, pretoria. florgård, c., 2007, ‘preserved and remnant natural vegetation in cities: a geographically divided field of research’, landscape research 32, 79–94. http://dx.doi.org/10.1080/01426390601097750 foth, h.d., 1990, fundamentals of soil science, 8th edn., john wiley & sons, new york. germishuizen, g. & meyer, n.l. (eds.), 2003, ‘plants of southern africa: an annotated checklist’, strelitzia 14, 1–1231. grobler, c.h., bredenkamp, g.j. & brown, l.r., 2006, ‘primary grassland communities of urban open spaces in gauteng, south africa’, south african journal of botany 72, 367–377. http://dx.doi.org/10.1016/j.sajb.2005.10.008 hennekens, s.m., 1996a, turboveg: software package for input, processing and presentation of phytosociological data, ibn-dlo, university of lancaster. hennekens, s.m., 1996b, megatab: a visual editor for phytosociological tables, giesen & geurts, ulft. hill, m.o., 1979a, twinspan: a fortran program for arranging multivariate data in an ordered two-way table by classification of individuals and attributes, cornell university, new york. hill, m.o., 1979b, decorana: a fortran program for detrended correspondence analysis and reciprocal averaging, cornell university, new york. johnson, m.r., van vuuren, c.j., visser, j.n.j., cole, d.i., wickens, h. de v., christie, a.m.d. et al., 2006, ‘sedimentary rocks of the karoo super group’, in m.r. johnson, c.r. anhaeusser & r.j. thomas (eds.), the geology of south africa, 2nd edn., the geological society of south africa, johannesburg. johnston, f.m. & johnston, s.w., 2004, ‘impacts of road disturbance on soil properties and on exotic plant occurrence in subalpine areas of the australian alps’, arctic, antarctic, and alpine research 36(2), 201–207. http://dx.doi.org/10.1657/1523-0430(2004)036[0201:iordos]2.0.co;2 kent, m. & coker, p., 1996, vegetation description and analysis, 2nd edn., john wiley & sons, london. land type survey staff, 1993, ‘land types of the maps 2926 aa-bloemfontein and 2926 ab-maselspoort’, memoirs on the agricultural natural resources of south africa 14. le houérou, h.n., poppov, g.f. & see, l., 1993, agro-bioclimatic classification of africa, agrometereology series working paper no. 6, food and agricultural organization, rome. low, a.b. & rebelo, a.g., 1996, vegetation of south africa, lesotho and swaziland, department of environmental affairs and tourism, pretoria. mangaung metropolitan municipality, 2011, integrated development plan review for 2011–2012, mangaung metropolitan municipality, bloemfontein. millard, a., 2004, ‘indigenous and spontaneous vegetation: their relationship to urban development in the city of leeds, uk’, urban forestry & urban greening 3, 39–47. http://dx.doi.org/10.1016/j.ufug.2004.04.004 mucina, l., hoare, d.b., lötter, m.c., du preez, p.j., rutherford, m.c, scott-shaw, c.r. et al., 2006, ‘grassland biome’, in l. mucina & m.c. rutherford (eds.), the vegetation of south africa, lesotho and swaziland, strelitzia 19, pp. 348–437, south african national biodiversity institute, pretoria. mueller-dombois, d. & ellenberg, h., 1974, aims and methods of vegetation ecology, wiley, new york. nicholson-lord, d., 1987, the greening of the cities, routledge & kegan paul, london. http://dx.doi.org/10.4324/9780203392614 noy-meir, i., 1973, ‘desert ecosystems: environment and producers’, annual review of ecology and systematics 4, 25–51. http://dx.doi.org/10.1146/annurev.es.04.110173.000325 platt, r.h., rowntree, r.a. & muick, p.c., 1994, the ecological city: preserving and restoring urban biodiversity, the university of massachusetts press, amherst. poynton, j.c. & roberts, d.c., 1985, ‘urban open space planning in south africa: a biogeographical perspective’, south african journal of botany 81, 33–37. roberts, d.c., 1993, ‘vegetation ecology of municipal durban, natal: floristic classification’, bothalia 23, 271–326. rutherford, m.c. & westfall, r.h., 1994, ‘biomes of southern africa. an objective categorisation’, 2nd edn., memoirs of the botanical survey of south africa 63, 1–94. sala, o.e. & lauenroth, w.k., 1982, ‘small rainfall events: an ecological role in semiarid regions’, oecologia 53, 301–304. http://dx.doi.org/10.1007/bf00389004 schulze, b.r., 1947, ‘the climates of south africa according to the classifications of köppen and thornwaite’, south african geographical journal 29, 32–42. http://dx.doi.org/10.1080/03736245.1947.10559253 schulze, b.r. & mcgee, o.s., 1978, ‘climatic indices and classifications in relation to the biogeography of southern africa’, in m.j.a. werger (ed.), biogeography and ecology of southern africa, pp. 19–52, w junk, the hague. http://dx.doi.org/10.1007/978-94-009-9951-0_2 soil classification working group, 1991, ‘soil classification: a taxonomic system for south africa’, memoirs on the agricultural natural resources of south africa 15. ter braak, c.j.f. & šmilauer, p., 2009, canoco reference manual and user’s guide to canoco for windows: software for canonical community ordination (version 4.5), microcomputer power, new york. tilman, d. & el haddi, e., 1992, ‘drought and biodiversity in grasslands’, oecologia 89, 257–264. van oudtshoorn, f., 1999, guide to grasses of south africa, briza publications, pretoria. van wyk, b. & van wyk, p., 1997, field guide to trees of southern africa, struik publishers, cape town. van wyk, e., cilliers, s.s. & bredenkamp, g.j., 2000, ‘vegetation analysis of wetlands in the klerksdorp municipal area, north west province, south africa’, south african journal of botany 66, 52–62. wood, j., low, a.b., donaldson, j.s. & rebelo, a.g., 1994, ‘threats to plant species diversity through urbanisation and habitat fragmentation in the cape metropolitan area, south africa’, in b.j. huntley (ed.), botanical diversity in southern africa, pp. 259–274, creda press, cape town. pmid:7941307 a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe original research vol. 50 no. 1 pp. 82 92 vegetation description of the doornhoek section of the mountain zebra national park (mznp), south africa abstract the mountain zebra national park (mznp) has been extended over the last couple of years. one of the newly procured areas is the doornhoek section, which had been adjacent to the park. to develop scientifically sound management programmes for conservation areas, it is essential that an inventory of their natural resources be undertaken. the aim of this study was to classify, describe and map the vegetation of the doornhoek section of the park. the floristic data were analysed in accordance with the braun-blanquet procedures using the bbpc suite. the data analysis resulted in the identification of eight communities, which can be grouped into seven major community types (rhus lucida–buddleja glomerata shrubland, rhigozum obovatum–rhus longispina shrubland, helichrysum dregeanum–aristida diffusa grassland, pentzia globosa–enneapogon scoparius grassland, aristida adscensionus–pentzia globosa grassland, cadaba aphylla–acacia karroo woodland and lycium oxycarpum–acacia karroo woodland). four of these communities occur on the higher-lying plateau, mid-slope and crest areas, while the other four communities are located on the lower-lying mid-plateau and foot slope, along drainage lines and in valley-bottom areas. the description of the plant communities, together with the vegetation map, can serve as a basis for formulating a management programme for the larger park. although sections of doornhoek have been overgrazed and degraded in the past, its recent addition to the mznp contributes to the available habitat preferred by large herbivores, such as valley bottoms, foot-slopes and plateaux. keywords: vegetation classification, inventory, braun-blanquet, mountain zebra national park (mznp), doornhoek south africa has an entrenched system of national parks situated within various biomes to protect and manage the ecological integrity of the different ecosystems for present and future generations. one such park, the mountain zebra national park (mznp), located in the eastern cape province of south africa, was established in 1937 to preserve the last cape mountain zebra (equus zebra zebra) population in the bankberg area of the park. in order to sustain a viable population of these animals, the park area was extended in 1960 and again in 1996, when various farms adjacent to the park were also procured. these newly incorporated areas resulted in the park more than doubling its size from its previous 6 536 ha (1960) to approximately 21 000 ha. the park is visited by more than 14 000 people annually and has considerable potential to serve as an education centre for environmental education and wildlife management. for any natural area to be managed effectively, it is important that scientifically sound management programmes be developed and implemented. this necessitates an inventory of its natural resources to be undertaken, which enables full quantitative measures of biodiversity to be used to make conservation decisions. a detailed vegetation study of the newly acquired areas of the park was therefore undertaken. although extensive studies on various aspects of the vegetation of the old park area were conducted by van der walt (1980), novellie (1990a; 1990b) and novellie and bezuidenhout (1994) and also of the de rust (brown & bezuidenhout, 2000), ebenaeser (de klerk, brown & bezuidenhout, 2003) and ingleside and welgedacht (brown & bezuidenhout, 2005) areas, little is known about the vegetation and habitat status of the doornhoek section. the aim of this study was therefore to classify, describe and map the vegetation of the doornhoek section of the park in order to serve as an inventory of the representative ecosystems and their biota. these relatively homogeneous plant communities should form the basis for the compilation of wildlife and ecotourism management plans. this study forms part of a larger and long-term research project undertaken by south african national parks (sanparks) and the university of south africa (unisa) to describe and map the vegetation of each of the newly acquired farms and assess the habitats of these areas for the establishment of large herbivores. study area the mznp is situated approximately 25 km west of the town of cradock in the eastern cape province. it extends from latitude 320 05’ to 320 20’s and longitude 250 23’ to 250 32’e (see fig. 1). the doornhoek section comprises 4 029 ha and lies approximately 4 km north of the mznp. the vegetation consists of a mixture of dwarf shrubs and grasses with acacia karroo trees in the lower-lying areas and river beds (van der walt 1980). according to acocks (1988), the hugo bezuidenhout conservation sciences south african national parks south africa leslie r. brown department of environmental sciences university of south africa south africa correspondence to: leslie r. brown e-mail: lrbrown@unisa.ac.za postal address: applied behavioural ecology and ecosystem research unit, department of environmental sciences, university of south africa, private bag x6, florida, 1710, republic of south africa 32 82 2008 original research bezuidenhout & brown koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 82 92 area can be classified as false karroid broken veld (37), while hoffman (1996) classifies it as eastern mixed nama karoo (52). some isolated patches of the south-eastern mountain grassland (grassland biome) are also found in the study area (lubke, bredenkamp & van rooyen, 1996). according to mucina, rutherford and powrie (2005) and mucina and rutherford (2006), this vegetation is classified as eastern upper karoo. the terrain is mountainous and comprises lower-lying valleys, drainage lines and river beds, together with steep to moderately steep mountain slopes and relatively flat to undulating plateaux. land types with reference to physiography, geology and soil three land types, namely da, ib and fc, occur in the doornhoek section of the mznp. according to the land type survey staff (1986), “a land type denotes an area that can be shown at 1:250 000 scale and that displays a marked degree of uniformity with respect to terrain form, soil pattern and climate”. a remarkable association between the major plant communities and the different land types of the area has been observed (bezuidenhout 1993). according to the land type survey staff (1986), the d-land type refers to a soil pattern where duplex soils, such as swartland and valsrivier forms, are dominant. without exposed rocks, stones or boulders, more than half of the remaining land must consist of duplex soils. unit da refers to land where duplex soils with red b horizons comprise more than half the area covered by duplex soils. these plains are dominated by plateau midslopes. the geology of this land type is mudstone, shale and sandstone of the balfour formation, beaufort group of the karoo supergroup. the f-land type refers to pedologically young landscapes, which are predominantly rocky and not alluvial or aeolian. the fc unit refers to land where lime occurs regularly in upland and valley-bottom soils (land type survey staff, 1986). soil forms that epitomise this land type are glenrosa and oakleaf. the geology consists of mudstone, shale and sandstone of the beaufort group of the karoo sequence with rare dolerite intrusions (land type survey staff, 1999). two topographical positions are prominent in this terraced landscape, namely the foot slopes and valley bottoms. the ib-land type unit refers to exposed rocks, which cover 60 to 80% of the area. the rocky portions of ib may be underlain by soil that would qualify the unit for inclusion in another broad soil pattern had it not been for the surface rockiness. the mid-slopes are the most prominent topographical position in this mountainous landscape, with the dominant soil-rock complex consisting of rock, while the glenrosa soil form is subdominant. the geology consists of dolerite with mudstone, shale and sandstone of the balfour formation, beaufort group of the karoo supergroup (land type survey staff, 1999). climate the average annual rainfall for the area, as measured at the mznp weather station from july 1962 to june 2004, is 405 mm, with the highest rainfall of 651 mm and lowest rainfall of 153 mm being recorded in 1977 and 1966 respectively. during the previous 10 years, the lowest rainfall recorded was 213,4 mm (in 1992) and the highest 565,5 mm (in 1991). the average monthly rainfall varies from 11,3 mm in the dry winter season to 60,8 mm 33 figure 1 location of the mountain zebra national park (mznp) 83 mountain zebra national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 82 92 in the wet summer season (see fig. 2). the average maximum temperature varies between 23,11 and 28,40 c in summer (from september to march) and 16,2 and 22,70 c in winter (from april to august), while the average minimum temperature varies between 5,6 and 13,60 c in summer and 0,05 and 7,80 c in winter. the mean average maximum and minimum temperatures are also indicated in fig. 2. methods by using 1:50 000 stereo aerial photographs, the study area was stratified into physiognomicphysiographic units. after a reconnaissance of the area, a total number of 66 sample plots was located on a randomly stratified basis within the different homogeneous units identified from the aerial photographs. the number of sample plots allocated within each homogeneous unit depended on the size of the area, the larger the area the higher the number of sampling plots being allocated to the unit. plot sizes were fixed at approximately 400 m2 according to brown (1997). in each sample plot, all species were recorded and the cover abundance of each species was assessed in accordance with the braun-blanquet cover-abundance scale (mueller-dombois & ellenberg, 1974). plant species identification was done according to germishuizen and meyer (2003). structural terminology was done according to edwards (1983). the percentage cover of the herbaceous layer (grasses and herbaceous plants), the shrub layer (woody species varying in height between > 0 to 3 m) and the tree layer (woody species higher than 3 m) was also estimated. the habitat information noted, including geology, soil forms, soil depth (shallow < 0,3 m, medium 0,3 to 0,9 m and deep > 0,9 m) and other edaphic factors, slope, aspect and the rockiness of the soil surface of each relevé was qualitatively described. rock sizes were estimated according to their diameter using the following scale: small (10 to 50 mm), medium (50 to 200 mm) and large (> 200 mm). the floristic data were analysed according to braun-blanquet procedures using the bbpc suite (bezuidenhout, biggs & bredenkamp, 1996). the numerical classification program twinspan (hill, 1979), which several phytosociologists regard as a successful approach for classification (mucina & van der maarel, 1989; bredenkamp & bezuidenhout, 1995; cilliers, 1998), was used to derive a first approximation of the floristic data. further refinement of the classification was achieved by braun-blanquet procedures (bredenkamp, joubert & bezuidenhout, 1989; kooij, bredenkamp & theron, 1990; bezuidenhout, 1993; eckhardt, 1993; brown & bezuidenhout, 2005). using the phytosociological table and the habitat information gathered during the sampling period, the different plant communities were identified and described according to their dominant species. dominant species are those that are 34 figure 2 average monthly rainfall and mean average minimum and maximum temperatures as recorded at the mountain zebra national park (mznp) weather station (rainfall covers the period from july 1962 to june 2005, while temperatures reflect the period from january 1992 to december 1998) figure 3 vegetation map for the doornhoek section of the mountain zebra national park (mznp) 84 0 5 10 15 20 25 30 jan feb mar apr may jun jul aug sep oct nov dec months te m pe ra tu re (0 c ) 0 10 20 30 40 50 60 70 r ai nf al l ( m m ) max temp min temp rainfall (mm) original research bezuidenhout & brown koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 82 92 most conspicuous in the community and are high in one or more of the importance values (whittaker, 1978), in this case cover and frequency. no attempt was made to fix syntaxa names formally, as this is normally avoided in detailed local studies (coetzee, 1983). soil nomenclature followed the classification of the soil classification working group (1991). results classification the data analysis resulted in the identification of eight different communities, which can be grouped into seven major community types. four of these communities occur on the higher-lying plateau, mid-slope and crest areas, while the other four communities are located on the lower-lying mid-plateau and foot slope, along drainage lines and in valley-bottom areas: a. relatively higher-lying vegetation communities on crests and steep slopes: 1. rhus lucida–buddleja glomerata shrubland 2. rhigozum obovatum-rhus longispina shrubland 2.1 carissa macrocarpa-rhus longispina shrubland 2.2 crabbea angustifolia-rhus longispina shrubland 3. helichrysum dregeanum-aristida diffusa grassland b. relatively lower-lying vegetation communities on foot slopes and in bottomland areas: 4. enneapogon scoparius-pentzia globosa grassland 5. aristida adscensionus-pentzia globosa grassland 6. cadaba aphylla-acacia karroo woodland 7. lycium oxycarpum-acacia karroo woodland description of the plant communities a. relatively higher-lying vegetation communities on crests and steep slopes: 1. rhus lucida–buddleja glomerata shrubland this shrubland is restricted to the higher mid-slopes. it is characterised by steep slopes with a gradient ranging between 250 and 400 and steep cliffs in some areas. rock cover is, as expected, high and varies between 55% and 90% with shallow, rocky soils sometimes covered by sparse vegetation. this community is associated with the ib-land type and is the smallest community in the study area, comprising only 53 ha. species from species group a (table 1) are diagnostic for this shrubland and include the trees olea europaea subsp. africana and cussonia paniculata, the shrubs buddleja glomerata, tarchonanthus camphoratus and myrsine africana, the dwarf shrub stachys linearis, the grasses melinis nerviglumis and andropogon schirensis, and the forbs achyranthes spp., clematis brachiata, datura stramonium and pellaea calomelanos. the woody layer comprising mostly large shrubs (1 m tall) ranges in cover between 25% and 35%, while the herbaceous layer (0,2 m tall) has a patchy appearance ranging in cover from 8% to 50% in some localities. the vegetation is dominated by the large shrubs buddleja glomerata and rhus lucida (species groups a & d respectively table 1), which typically occurs on rocks and hills (coates palgrave 2002). the shrubs tarchonanthus camphoratus (species group a – table 1), gymnosporia buxifolia and grewia occidentalis (species group f – table 1) are also prominent although they have relatively low canopy cover. the grasses melinis repens (species group a – table 1), digitaria eriantha, sporobolus fimbriatus (species group k – table 1) and enneapogon scoparius (species group k – table 1) together with the declared weed datura stramonium (species group a – table 1) are locally prominent within this community. similar communities have been described in the mznp by brown and bezuidenhout (2000; 2005) and de klerk et al. (2003). these communities are all associated with steep cliffs, large rocks and boulders, shallow soil, and sparse vegetation cover. 2. rhigozum obovatum-rhus longispina shrubland the rhigozum obovatum-rhus longispina shrubland encompassing 1 078 ha in size, is located on the mountain foot slopes and steep mid-slopes mainly within the southern and eastern sections of the doornhoek section. the gradient on the mountain mid-slopes is mild to steep, while the foot slopes are generally flat with a slight gradient. the presence of species from species group b (table 1) is characteristic, while the vegetation is dominated by the shrub rhus longispina (species group b – table 1) and the grass aristida diffusa (species group i – table 1). this community can be divided into two sub-communities, namely the carissa macrocarpa-rhus longispina shrubland and the crabbea angustifolia-rhus longispina shrubland. 2.1 carissa macrocarpa-rhus longispina shrubland this sub-community (763 ha) is restricted to the mid-slopes and some mid-plateau areas of the study area and is associated with the ib-land type. the dominant soil-rock complex consists of rock, while the mispah-soil form is subdominant with a 30% to 55% rock cover. the mild to steep gradient varies between 100 and 210. the sub-community is characterised by the presence of species from species group c (see table 1) and includes the shrubs carissa macrocarpa, boscia oleoides, ehretia rigida and pappea capensis, the dwarf shrub lantana rugosum, the grasses cenchrus ciliaris, eustachys paspaloides and eragrostis trichophora, and the forbs boophane disticha, cheilanthes hirta, phyllanthus madagascariensis, hermannia cernua, asparagus burchellii and opuntia ficus-indica. the woody layer consists mainly of shrubs between 2 m and 3 m tall with a canopy cover of 10% to 25%, while the herbaceous layer covers between 35% and 50% of the area with an average height of 0,45 m. the vegetation is dominated by the shrub rhus longispina (species group b – table 1) and the grass aristida diffusa (species group i – table 1), while the shrub rhus lucida (species group d – table 1) and the grasses cymbopogon pospischilii (species group h – table 1), setaria sphacelata, heteropogon contortus (species group i), enneapogon scoparius and aristida adscensionis (species group k) are prominent. other species present include the tree acacia karroo (species group p – table 1), the shrub grewia occidentalis (species group f – table 1) and the dwarf shrub eriocephalus ericoides (species group i – table 1), which are locally prominent throughout the community. the community is similar to the enneapogon scoparius-ehretia rigida shrubland in the de rust section (brown & bezuidenhout, 2000), the setaria sphacelata-rhus longispina sub-community in the ebenaeser section (de klerk et al., 2003) and the rhus lucida-carissa macrocarpa shrubland in the ingleside section (brown & bezuidenhout, 2005) of the park. the sub-community comprises medium to large boulders with shallow soil and was prone to overgrazing by angora goats previously present in the doornhoek section. 2.2 crabbea angustifolia-rhus longispina shrubland located on the foot slopes of the mountains in the southern section of the study area, this shrubland (315 ha) occurs in deeper mispah soil than the cymbopogon pospischilii-rhus longispina shrubland (sub-community 2.1). rock cover is lower ranging, between 20% and 40%, and consists of small to medium-sized rocks. this sub-community is associated with the ib-land type. the vegetation is characterised by the presence of the forbs crabbea angustifolia, sutera halimifolia and hibiscus trionum 3585 mountain zebra national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 82 9236 p l a n t c o m m u n it y n u m b e r 1 2 3 4 5 6 7 2. 1 2. 2 r el ev é 10 17 18 1 2 8 16 20 23 25 30 29 26 36 37 41 42 58 57 21 22 24 31 28 3 5 7 13 11 12 14 15 38 39 40 43 4 4 45 46 52 56 35 63 6 4 65 49 51 5 4 55 61 62 50 6 3 4 60 53 4 27 9 19 32 47 59 33 48 66 s p ec ie s g ro u p a d ia g n o st ic s p ec ie s fo r th e r h u s lu ci d ab u d d le ja g lo m er at a s h ru b la n d (1 ) b ud dl ej a gl om er at a b 3 b ta rc ho na nt hu s ca m ph or at us + + s ta ch ys li ne ar is + + + o le a eu ro pa ea su bs p af ri ca na + + + a ch yr an th es s pe ci es + + c le m at is b ra ch ia ta + c us so ni a pa ni cu la ta + *d at ur a st ra m on iu m a m el in is n er vi gl um is 1 m yr si ne a fr ic an a + p el la ea c al om el an os + + a nd ro po go n sc hi re ns is + + s p ec ie s g ro u p b d ia g n o st ic s p ec ie s fo r th e r h ig iz u m o b ov at u m -r h u s lo n g is p in a s h ru b la n d (2 ) r hu s lo ng is pi na a a 1 a a a a b b b 1 a a + a + 1 a a 1 a 1 + + 1 + r hi go zu m o bo va tu m + + + b 1 a + a + 1 + + + a + a sp ar ag us th un be rg ia nu s + + + + + + + + + + + s po ro bo lu s af ri ca nu s + + + + + + + + + + + + + + + a ri st id a su bs p ba rb ic ol lis + + + + + + + + a sp ar ag us m uc ro na ta + + + + s p ec ie s g ro u p c d ia g n o st ic s p ec ie s fo r th e c ar is sa m ac ul at ar hu s lo ng is pi na s h ru b la n d (2 .1 ) c ar is sa m ac ro ca rp a + + 1 + + + + + + + + + b os ci a ol eo id es + + + + + + + + + c en ch ru s ci lia ri s + + 3 + + + + + + + + b oo ph an e di st ic ha + + + + + + e us ta ch ys pa sp al oi de s + + + + + + c he ila nt he s hi rt a + + + + + p hy lla nt hu s m ad ag as ca ri en si s + + + + la nt an a ru go su m r + + e hr et ia r ig id a + + + + h er m an ni a ce rn ua + + + + a sp ar ag us b ur ch el lii + + + + *o pu nt ia fi cu sin di ca + + + + + + ta b l e 1 ph yt os oc io lo gi ca l t ab le o f t he v eg et at io n of th e d oo rn ho ek s ec ti on o f t he m ou nt ai n z eb ra n at io na l p ar k (m z n p) (s pe ci es in di ca te d w it h an a st er is k [* ] a re d ec la re d in va de rs ) 86 original research bezuidenhout & brown koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 82 92 37 p l a n t c o m m u n it y n u m b e r 1 2 3 4 5 6 7 2. 1 2. 2 r el ev é 10 17 18 1 2 8 16 20 23 25 30 29 26 36 37 41 42 58 57 21 22 24 31 28 3 5 7 13 11 12 14 15 38 39 40 43 4 4 45 46 52 56 35 63 6 4 65 49 51 5 4 55 61 62 50 6 3 4 60 53 4 27 9 19 32 47 59 33 48 66 p ap pe a ca pe ns is + + e ra gr os tis tr ic ho ph or a + + + + + s p ec ie s g ro u p d c o m m o n sp ec ie s fo r co m m u n it ie s 1 & 2 .1 r hu s lu ci da 1 a a 1 1 + + 1 + + + s ol an um s up in um + + + + + + + + + s ta ch ys c ym ba la ri a + + + + s p ec ie s g ro u p e d ia g n o st ic s p ec ie s fo r th e c ra b b ea a n g u st if o lia -r h u s lo n g is p in a s h ru b la n d (2 .2 ) c ra bb ea a ng us tif ol ia + + + s ut er a ha lim ifo lia + + + h ib is cu s tr io nu m + + + + + s p ec ie s g ro u p f c o m m o n sp ec ie s fo r co m m u n it ie s 1, 2 .1 & 2 .2 g ym no sp or ia bu xi fo lia + + + + + 1 + + + + + + + + g re w ia o cc id en ta lis + + + + + + + + + + + + + + + + + s p ec ie s g ro u p g d ia g n o st ic s p ec ie s fo r th e h el ic h ry su m d re g ea n u m -a ri st id a d if fu sa g ra ss la n d (3 ) h el ic hr ys um dr eg ea nu m + + + + + + + + + + + + + a nt ho sp er m um hi sp id ul um + + + + + + b ec iu m bu rc he lli an um + + + + + b s al so la tu be rc ul at a + + + p eg ol et tia r et ro fr ac ta + + + + + s p ec ie s g ro u p h c o m m o n sp ec ie s fo r co m m u n it ie s2 .1 , 2 .2 & 3 c ym bo po go n po sp is ch ili i a 1 b + 1 + + + + + + + + + + + d ig ita ri a er ia nt ha a + a + + 1 + + + + + + + + + + + + + + 1 + te ph ro si a sp ec ie s + + + + + + + + + + + + + + + in di go fe ra a lte rn an s + + + + + + + + + + + + s po ro bo lu s fim br ia tu s a + + + + + + + 1 1 + + + d io sp yr os a us tr o af ri ca nu m + + + + c hr ys oc om a ci lia ta + + + + + + + + + a sp ar ag us s tr ia tu s + + + + + + + s p ec ie s g ro u p i c o m m o n sp ec ie s fo r co m m u n it ie s 2. 1, 2 .2 , 3 & 4 a ri st id a di ff us a + + a 1 3 3 3 + b b 3 b + b + + b + + b + a a 3 + + 3 + + b + a + tr ag us k oe lle ri oi de s + + + + + + + + + + + + + + + + + + + + + + + + 1 + + + + + + + 1 + + + e ri oc ep ha lu s er ic oi de s + 1 + + 1 + 1 + + + + 1 + + + + + + + + + + + b + s et ar ia s ph ac el at a + + + 1 + a a 1 + 1 + + 1 + + + 1 + 1 + + ta bl e 1 (c on t.. .) 87 mountain zebra national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 82 9238 p l a n t c o m m u n it y n u m b e r 1 2 3 4 5 6 7 2. 1 2. 2 r el ev é 10 17 18 1 2 8 16 20 23 25 30 29 26 36 37 41 42 58 57 21 22 24 31 28 3 5 7 13 11 12 14 15 38 39 40 43 4 4 45 46 52 56 35 63 6 4 65 49 51 5 4 55 61 62 50 6 3 4 60 53 4 27 9 19 32 47 59 33 48 66 h et er op og on co nt or tu s + 1 + + a 1 1 + + + a a + + + 1 + 1 a a a 1 + 1 1 + 1 + + s ut er a sp ec ie s + + + + + + + + + + + + + + + + + o ro pe tiu m c ap en se + + + + + + + 1 + + + + + a g al en ia s pe ci es + + + + + + + + + + + + + + + + li m eu m v is co su m + + + + + + + + + + + + + + + + + + + + + + + + + + + o xa lis s pe ci es + + + + + + + + + + + + + + + + a pt os im um pr oc um be ns + + 1 + + + + + + + b le ph ar is m itr at a + + + + + + + + + + 1 1 + + + + s p ec ie s g ro u p j d ia g n o st ic s p ec ie s fo r th e c ad ab a ap hy lla -a ca ci a ka rr o o w o o d la n d (6 ) c ad ab a ap hy lla + + + + + + + + 1 + + + d io sp yr os ly ci oi de s + + + + + + + + + + + + + a tr ip le x se m ib ac ca ta + + + + + z yg op hy llu m in cr as sa ta + + + + + 1 + + *c ir si um v ul ga re + + + s al so la k al i + + a + s p ec ie s g ro u p k c o m m o n sp ec ie s fo r co m m u n it ie s 2. 1, 2 .2 , 3 , 4 , 5 & 6 p en tz ia g lo bo sa + + + 1 + + + + a + + + + b b + 1 a 3 + + + a + + + + 3 3 3 3 a a a 3 b + b 1 1 3 3 3 3 1 3 + a a a + + 1 + + e nn ea po go n sc op ar iu s 1 + + 1 + 1 a 1 1 + + + + + + + a 3 b 3 3 + 3 1 a + 1 + + a + 1 + + a a 1 + 1 a b + + 1 + + + 1 1 e ra gr os tis le hm an ni an a + + a + b + + + + + + + a 1 + + + 1 b + + 4 3 b + 1 + + + + 3 + 1 + + + + e ra gr os tis o bt us a + + + + + 1 + + + + + + + + + 1 a + + a + + + + + + + + + + + + + + a ri st id a su bs p co ng es ta + + + + + + + 1 a + + + 1 + 1 a + 1 + + b + + + 3 b + + 4 b b + + + a ri st id a ad sc en si on is + + + 1 + 3 + b 1 + + + + 1 + + 1 a + a b + b a 3 a 3 a 1 + + + + + p an ic um c ol or at um + + + + + + + + + + + + t he si um li ne at um + + + + + + + + + + + d el os pe rm um fr ut es ce ns + + + + + + + + + + + + + + + + + + + + + + + + + + + f el ic ia fi lif ol ia + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + f el ic ia m ur ic at a + + + + + + + + + + + + + + + + + + + + + + tr ag us b er te ro ni an us + + + + + + + + + + + + + + + + + + + + + + + + + + + + + tr ib ul us te rr es tr is + + + + + + + + + + + + + + + + ly ci um c in er eu m + + + + + + + + + + + + + + + 1 + 1 + + + + 1 + + + + + 1 s p ec ie s g ro u p l d ia g n o st ic s p ec ie f o r th e ly ci u m o xy ca rp u m -a ca ci a ka rr o o w o o d la n d (7 ) *d at ur a fe ro x + + s p ec ie s g ro u p m c o m m o n sp ec ie s fo r co m m u n it ie s 6 & 7 ta ge te s m in ut a + + + + + + + b id en s bi pi nn at a + + + + + ta bl e 1 (c on t.. .) 88 original research bezuidenhout & brown koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 82 92 (species group e – table 1). there are no trees present and the shrub layer, which is between 1,5 m and 2 m tall, ranges in canopy cover between 5% and 10%. the herbaceous layer is dominant and has a canopy cover of 20% to 45% and is approximately 0,4 m tall. this sub-community is dominated by the shrub rhus longispina (species group b – table 1) and the grass enneapogon scoparius (species group k – table 1). the tree acacia karroo (species group p – table 1), the dwarf shrub pentzia globosa (species group k – table 1) and the grass heteropogon contortus are codominants within this sub-community. other species also present include the shrub rhigozum obovatum (species group b – table 1), the dwarf shrub eriocephalus ericoides (species group i – table 1) and the grasses aristida diffusa, setaria sphacelata (species group i – table 1) and chloris virgata (species group o – table 1). the sub-community shows similarities with the enneapogon scoparius-ehretia rigida shrubland of the de rust section (brown & bezuidenhout, 2000), although the latter is present on plateau mid-slopes, and the setaria sphacelata-rhus longispina subcommunity in the ebenaeser section (de klerk et al., 2003). 3. helichrysum dregeanum-aristida diffusa grassland this grassland community, which is only 305 ha in size, is found on the higher-lying plateau areas located in the southwestern section of the study area. the soil is shallow, with small to medium-sized rocks, covering between 10% and 20% of the area and occurs on the da-land type. smaller sections with slightly deeper soil are also present, although localised in distribution. the community is characterised by the presence of species from species group g (table 1), namely the dwarf shrubs helichrysum dregeanum, becium burchellianum, salsola tuberculata and pegolettia retrofracta and the forb anthospermum hispidulum. the tree and shrub layers are not well developed, with only a few single individuals present and a total canopy cover of approximately 1% to 3%. the grass layer has an approximately 35% canopy cover and an average height of 0,4 m. the vegetation is dominated by the grasses aristida diffusa and heteropogon contortus (species group i – table 1). prominent species include the dwarf shrub pentzia globosa and the grasses enneapogon scoparius, eragrostis lehmanniana and aristida congesta subsp. congesta (species group k – table 1). the absence of species from species group o, which is present in almost all the other communities, is also a feature of this community, while the dwarf shrub salsola tuberculata (species group g – table 1) occurs only within this community. the dominant grasses aristida diffusa and heteropogon contortus form dense stands that are evenly dispersed throughout the community. the grasses oropetium capense and tragus koellerioides (species group i – table 1) are also present and are prominent in patches locally, while the dwarf shrub helichrysum dregeanum (species group g – table 1) is present throughout the community. no similar communities have been described in previous studies in the de rust, ebenaeser, ingleside and welgedacht sections of the park. b. relatively lower-lying vegetation communities on foot slopes and bottom-land areas: 4. pentzia globosa-enneapogon scoparius grassland the pentzia globosa-enneapogon scoparius grassland occurs on the mid-plateau and foot-slope areas adjacent to the drainage lines located in the north-eastern section of the study area. it is associated with the fc-land type and the dominant soil form is shallow mispah, while the medium-deep oakleaf soil form occurs in the lower-lying areas. the soil is gravelly and 39 p l a n t c o m m u n it y n u m b e r 1 2 3 4 5 6 7 2. 1 2. 2 r el ev é 10 17 18 1 2 8 16 20 23 25 30 29 26 36 37 41 42 58 57 21 22 24 31 28 3 5 7 13 11 12 14 15 38 39 40 43 4 4 45 46 52 56 35 63 6 4 65 49 51 5 4 55 61 62 50 6 3 4 60 53 4 27 9 19 32 47 59 33 48 66 s p ec ie s g ro u p n c o m m o n sp ec ie s fo r co m m u n it ie s 4, 5 , 6 & 7 p si lo ca ul on a bs im ile + + + + 1 1 + + + + + + + + + 1 + 1 + 1 1 + + s p ec ie s g ro u p o c o m m o n sp ec ie s fo r co m m u n it ie s 2. 1, 2 .2 , 3 , 4 , 5 , 6 & 7 ly ci um o xy ca rp um + + + + + + + + + r + r + + 1 a + 1 + 1 a a b p an ic um m ax im um + + + + + + + + + + + + + + 3 3 1 u ro ch lo a pa ni co id es + + + + + + + + + + + + + + + + + + + + + + + + + + + + c hl or is v ir ga ta + + + + + a + + r 1 + + 1 + + 1 1 1 a 3 1 1 3 3 3 4 3 b a 1 a + 1 s p ec ie s g ro u p p c o m m o n sp ec ie s fo r al l o f t h e co m m u n it ie s o f d o o rn h o ek f ar m a ca ci a ka rr oo + + 1 + a + + + + 1 + + + + 1 + + 1 1 a 1 + r + + 1 r a 1 a a a b 3 b b b a 3 c yn od on in co m pl et us + + + + + + + + + + + + + + + + + + + + + + + + a + + 1 b + + 1 w al af ri da g en ic ul at a + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + a sp ar ag us su av eo le ns + + + + + + + + + + + + + + + + 1 + + + + + + + + + + + + + 1 + + + + + + + + + *o pu nt ia a ur an tia ca + + + + + + + + + + + + + + + + + + + + + + + m or ae a po ly st ac hy a + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + ta bl e 1 (c on t.. .) 89 mountain zebra national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 82 92 small rocks cover between 5% and 50% of the surface. this community, which has a total size of 786 ha, comprises 20% of the study area. the community has no distinguishing species group but the presence of both species groups i and k (see table 1) is characteristic. no trees are present and the herbaceous layer, which has an average height of 0,3 m, has a canopy cover ranging between 30% and 60%. the vegetation is dominated by the dwarf shrub pentzia globosa and the grasses enneapogon scoparius and eragrostis lehmanniana (species group k – table 1). the grasses eragrostis obtusa and aristida adcensionis (species group k – table 1) and the grass aristida diffusa (species group i – table 1) are prominent in large sections of the community. the small shrub lycium cinereum (species group k – table 1) is prominent locally. the community shows similarities with the eragrostis obtusapentzia globosa sub-community of the ebenaeser section (de klerk et al., 2003) and the eragrostis obtusa-pentzia globosa shrubland on the farms ingleside and welgedacht (brown & bezuidenhout, 2005). locally, some patches are dominated by the increaser 2 grasses aristida congesta subsp. congesta (species group k – table 1) and chloris virgata (species group o – table 1) (van oudtshoorn, 2004). these patches were heavily grazed in the past by goats and sheep, leading to the presence of these pioneer grasses. 5. aristida adscensionus-pentzia globosa grassland this open grassland is found in the central and western sections of the study area in the lower-lying valley bottoms and floodplain areas associated with the fc-land type. the community is 589 ha in size and has low rock cover varying between 0% and 10%. the dominant soil forms are hutton and oakleaf. this community is characterised by the presence of species from species group k and the absence of species from species group i (table 1). only a few trees (2 m to 3 m tall) are present and they have a total canopy cover of approximately 1%, while the shrub layer (1 m to 2 m tall) has a cover of approximately 5%. the herbaceous layer between 0,2 m and 0,45 m tall has an average canopy cover of 43% and ranges between 35% and 70%. the vegetation is dominated by the dwarf shrub pentzia globosa (species group k – table 1) and the grass chloris virgata (species group o – table 1). prominent species include the grasses aristida adscensionis, enneapogon scoparius, aristida congesta subsp. congesta and eragrostis lehmanniana (species group k – table 1). the spreading shrub psilocaulon absimile (species group n – table 1) is present in some localities within this community. similar communities have been described in the de rust section (psilocaulon junceum-eragrostis lehmanniana grassland) and ingleside and welgedacht sections (chloris virgata-cynodon incompletus grassland) by brown and bezuidenhout (2000; 2005). this community forms part of the larger drainage lines within the study area. various soil barriers, however, were constructed within the community, probably to prevent erosion during floods. this, together with overgrazing, has resulted in the area becoming degraded and dominated by a large number of prominent pioneer-plant species, such as aristida congesta subsp. congesta, chloris virgata, psilocaulon absimile and cynodon incompletus (species groups k, o, n and p – table 1). termite mounds were also noted in some localities of the community. 6. cadaba aphylla-acacia karroo woodland this community (439 ha in extent) is located in the narrow drainage lines between the mountains in the southern and eastern sections of the study area. it is associated with the daland type and the dominant soil form is oakleaf. not much rock is present, small rocks covering approximately 3% of the area. the vegetation is characterised by the presence of the shrubs diospyros lycioides and cadaba aphylla, the dwarf shrubs atriplex semibaccata, zygophyllum incrassata and salsoli kali and the forb cirsium vulgare (species group j – table 1). the tree layer, which is taller than 3 m, has a canopy cover of between 1% and 50%, the shrub layer (1 m to 3 m tall) between 5% and 25% and the grass layer (0,2 m to 0,5 m tall) between 15% and 50%. the vegetation is dominated by the trees acacia karroo and lycium oxycarpum (species groups p and o respectively – table 1), the grass chloris virgata (species group o – table 1) and the spreading shrublet psilocaulon absimile (species group n – table 1). the shrub cadaba aphylla (species group j – table 1), although present in sparse stands in communities 1 and 2 (table 1), has a high constancy – 71% – within this woodland. cadaba aphylla is a preferred shrub and is utilised frequently by ungulates, which adds to this community being preferred by herbivores. it is also an important source of nectar for various bird and insect species (le roux, kotzé, nel & glen, 1994). the community shows similarities with the sporobolus africanus-acacia karroo woodland (brown & bezuidenhout, 2000), setaria sphacelata-cadaba aphylla shrubland (de klerk et al., 2003) and panicum maximum-acacia karroo woodland (brown & bezuidenhout, 2005) of the de rust, ebenaeser, and ingleside and welgedacht sections of the park. none of these three communities, however, is as degraded as this woodland, having more climax and fewer pioneer species. 7. lycium oxycarpum-acacia karroo woodland the lycium oxycarpum-acacia karroo woodland is located in large, open drainage lines and the valley bottom in the central part of the study area. this large community is 779 ha in size and is associated with the fc-land type. the dominant soil form is oakleaf and no rocks are present on the surface. the herbaceous annual and declared weed datura ferox (species group l – table 1) is characteristic of this community and typically occurs in drainage channels and river beds (henderson, 2001). the tree layer has a canopy cover of 20% and is taller than 4 m, while the shrub layer covers between 20% and 40% of the area and is between 2 m and 3 m tall. the welldeveloped herbaceous layer, dominated by grasses up to 0,7 m tall, has an 80% canopy cover. the vegetation is dominated by the trees acacia karroo (species group p – table 1) and lycium oxycarpum and the grass panicum maximum (species group o – table 1). the grasses chloris virgata (species group o – table 1) and cynodon incompletus (species group p – table 1) are prominent. this community is similar to community 6 but differs in that the spreading shrub psilocaulon absimile (species group n – table 1) is not as prominent and the shrub cadaba aphylla (species group j – table 1) together with the dwarf shrub pentzia globosa (species group k – table 1) are absent. the climax grass panicum maximum (species group o – table 1) is also dominant within this woodland, while it is merely present and even absent in community 6. thus, although similar to community 6, which also occurs in drainage lines, this community is not as degraded and therefore has a greater similarity with the sporobolus africanusacacia karroo woodland (brown & bezuidenhout, 2000), the setaria sphacelata-cadaba aphylla shrubland (de klerk et al., 2003) and the panicum maximum-acacia karroo woodland (brown & bezuidenhout, 2005) of the de rust, ebenaeser, and ingleside and welgedacht sections of the park. 40 90 original research bezuidenhout & brown koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 82 92 discussion the general vegetation of the study area is characterised by the presence and, in some areas, the dominance of the tree acacia karroo and the dwarf shrub pentzia globosa, while the grasses cynodon dactylon and enneapogon scoparius are also present in most of the plant communities. large sections of the vegetation, especially in the lower-lying areas, were overgrazed due to previous farming practices with angora goats and sheep. the higher-lying areas were less prone to overgrazing, although patches do occur within the plateau areas where overgrazing did occur. the different plant communities identified and described are clearly distinguishable from each other in the field as separate vegetation units. plant communities 1, 2 and 3 are characteristic of the higher-lying areas, which have high rock cover and shallow soil. in contrast, communities 4, 5 and 6 are found on the lower-lying areas, which have low rock cover and deeper soil. plant community 1 is characteristic of the higher-lying areas on the escarpment, which has very steep (250 to 280) upper slopes and high rock cover, ranging between 50% and 90%, and shallow soil. the woody layer is the most prominent layer and is dominated by the shrub buddleja glomerata. plant community 2 is located on the steep mid-slopes, which have high rock cover and shallow soil. the woody layer is still prominent and is dominated by the tall shrub rhus longispina, which is conspicuous on all the mid-slopes in the study area. the herbaceous layer is more prominent, covering up to 50% of the area, and is dominated by the grass aristida diffusa. located on the higher-lying plateau areas in shallow to medium-deep soil with a 10% to 20% rock cover is the helichrysum dregeanum – aristida diffusa grassland (plant community 3). this grassland was prone to overgrazing, which is evident in the plant species composition, where the vegetation is dominated by the grasses aristida diffusa and heteropogon contortus. plant community 4 is located on the lower-lying mid-plateau and rocky foot-slope areas next to the drainage lines. the soil is deeper than that of the higher-lying areas (communities 1, 2 and 3) and has smaller rocks and lower rock cover. the vegetation is dominated by the dwarf shrub pentzia globosa and grass enneapogon scoparius. both communities 6 and 7 are associated with the drainage lines, which have little to no rock cover and deep alluvial soil. these areas were intensively grazed in the past and many pioneer species are therefore present. plant community 7 has better vegetation cover and composition, although sections were also overutilised in the past. the presence of dense stands of the declared weed opuntia aurantiaca throughout the study area is a reason for concern. this species is not present in such large numbers in the old section of the park and the possibility exists that it could spread from here into the old section. management of the park, in collaboration with working for water, has been actively involved in the eradication of this invasive species but it is recommended that monitoring plots be marked to monitor the recovery and/ or spread of this species. this species poses a huge threat to the environment in that it can replace the natural vegetation, thereby altering the functioning of the natural ecosystem. communities 5, 6 and 7 are all degraded due to previous overgrazing as a result of angora goat and sheep-farming practices. the degraded condition of these communities is indicated by the presence of a high number of pioneer and weedy species, such as psilocaulon absimile, cynodon incompletus, aristida congesta subsp. congesta, chloris virgata, urochloa panicoides, tagetes minuta, bidens bipinnata and cirsium vulgare. due to communities 6 and 7 being located in the drainage lines, they are also prone to disturbance from floods after heavy rainfall. the deeper and more fertile soil of these areas is responsible for the plants occurring here being more palatable. this, in turn, results in animals congregating in these areas, which causes further degradation. these areas will therefore have to be monitored regularly once game is relocated here to ensure that they are not overutilised and the stocking rates will have to be such that they allow the vegetation to recover. concluding remarks the braun-blanquet approach proved to be an accurate and effective way whereby floristically defined plant communities could be classified and identified in the field. a total of eight different, clearly recognisable plant communities, which can be grouped into seven major communities, were identified and described for the study area. the doornhoek section, although overgrazed and degraded in sections, increases the area available for large herbivores, especially the valley bottom, foot slope and plateau areas. an understanding of the plant communities and their associated habitats is of fundamental importance for devising sound management and conservation strategies. the description of the plant communities, together with the vegetation map, can serve as a basis for formulating a management programme for this section of the park as well as the larger park. the data obtained from this study will be incorporated into the existing vegetation map and management plan for the mznp as part of a larger project undertaken by unisa and sanparks. acknowledgements the following individuals and institutions are sincerely thanked: national research foundation (nrf) and unisa for financing this study; sanparks, especially the management of the mznp, for allowing and encouraging the research; mr johan de klerk for his dedication and support of this project; dr stephen holness for assistance with the maps; mr ernest daemane (kimberley south african national parks herbarium) for verifying the plant species names; and gn dithlale (unisa) for assistance with the species data analysis. references acocks, j.p.h. 1988. veld types of south africa. 3rd ed. memoirs of the botanical survey of south africa, 57: 1–146. bezuidenhout, h. 1993. syntaxonomy and synecology of western transvaal grasslands. phd dissertation, university of pretoria, pretoria. bezuidenhout, h., biggs, h.c. & bredenkamp, g.j. 1996. a process supported by the utility bbpc for analysing braun-blanquet data on a personal computer. koedoe, 39(1): 107– 112. bredenkamp, g.j. & bezuidenhout, h. 1995. a proposed procedure for the analysis of large data sets in the classification of south african grasslands. koedoe, 38(1): 33– 39. bredenkamp, g.j., joubert, a.f. & bezuidenhout, h. 1989. a reconnaissance survey of the vegetation of the plains in the potchefstroom-fochville-parys area. south african journal of botany, 55(2): 199–206. brown, l.r. 1997. a plant ecological and wildlife management plan for the borakalalo nature reserve, north west province. phd dissertation, university of pretoria, pretoria. brown, l.r & bezuidenhout, h. 2000. the phytosociology of the farm de rust in the mountain zebra national park, eastern cape. koedoe, 43(1): 1–18. brown, l.r & bezuidenhout, h. 2005. a vegetation description and classification of the farms ingleside and welgedacht in 4191 mountain zebra national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 82 92 the mountain zebra national park, eastern cape. koedoe, 48(2): 23–42. coates palgrave, m.c. 2002. keith coates palgrave trees of southern africa. cape town: struik. coetzee, b.j. 1983. phytosociology, vegetation structure and landscapes of the central district, kruger national park. dissertationes botanicae, 69: 1–456. cilliers, s.s. 1998. phytosociological studies of urban open spaces in potchefstroom, north west province, south africa. phd thesis, potchefstroom university for che, potchefstroom. de klerk, j., brown, l.r. & bezuidenhout, h. 2003. plant communities of the ebenaeser section of the mountain zebra national park. koedoe, 46(2): 1–13. eckhardt, h.c. 1993. a synecological study of the vegetation of the north-eastern orange free state. msc thesis, university of pretoria, pretoria. edwards, d. 1983. a broad-scale structural classification of vegetation for practical purposes. bothalia, 14: 705–712. germishuizen, g. & meyer n.l. (eds.). 2003. plants of southern africa: an annotated checklist. strelitzia, 14. henderson, l. 2001. alien weeds and invasive plants. plant protection research institute handbook no. 12. cape town: agricultural research council. hill, m.o. 1979. twinspan: a fortran program for arranging multivariate data in an ordered two-way table by classification of the individuals and attributes. new york: cornell university. hoffman, t. 1996. eastern mixed nama karoo, in a.b. low & a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. kooij, m.s., bredenkamp, g.j. & theron, g.k. 1990. classification of the vegetation of the b land type in the north-western orange free state. south african journal of botany, 56: 309– 318. land type survey staff. 1986. land types of the maps se27/20 witdraai, 2720 noenieput, 2722 kuruman, 2724 christiana, 2820 upington, 2822 postmasburg. memoirs of the agricultural natural resources of southern africa, 3: 1–185. land type survey staff. 1999. land types occurrence (maps) and areas of 3224 graaff-reinet. unpublished information from the institute for soil, climate and water, pretoria. le roux, p.m., kotzé, c.d., nel, g.p. & glen, h.f. 1994. bossieveld, grazing plants of the karoo and karoo-like areas. bulletin, 428. lubke, r., bredenkamp, g.j. & van rooyen, n. 1996. southeastern mountain grassland, in a.b. low & a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. mucina, l. & rutherford, m.c. (eds.). 2006. vegetation of south africa, lesotho and swaziland. strelitzia, 19. mucina, l., rutherford, m.c. & powrie, l.w. (eds.). 2005. vegetation map of south africa, lesotho and swaziland, 1:1 000 000 scale sheet maps. pretoria: south african national biodiversity institute. mucina, l. & van der maarel, e. 1989. twenty years of numerical syntaxonomy. vegetation, 81: 1–15. mueller-dombois, d. & ellenberg, h. 1974. aims and methods of vegetation ecology. new york: wiley. novellie, p. 1990a. habitat use by indigenous grazing ungulates in relation to sward structure and veld condition. south african journal of wildlife research, 7(1): 16–23. novellie, p. 1990b. the impact of a controlled burn on karroid merxmuellera mountain veld in the mountain zebra national park. south african journal of ecology, 1: 33–37. novellie, p.a. & bezuidenhout, h. 1994. the influence of rainfall and grazing on vegetation changes in the mountain zebra national park. south african journal of wildlife research, 24(3): 60–71. soil classification working group. 1991. soil classification: a taxonomic system for south africa. memoirs on the agricultural natural resources of south africa, 15: 1–262. van der walt, p.t. 1980. a phytosociological reconnaissance of the mountain zebra national park. koedoe, 23: 1–32. van oudtshoorn, f. 2004. guide to grasses of southern africa. pretoria: briza publications. whittaker, r.h. 1978. dominance types, in r.h. whittaker (ed.). classification of plant communities. the hague. 92 filelist convert a pdf file! koedoe 19: 27-30 (1976) vitamin a storage in the liver of certain african wild ruminants and elephant c. bartelmus, d. giesecke institut jur tierphysiologie, universitiit m unchen 8 munchen 22 veteriniirstrasse 13 germany and p. hoppe department oj animal physiology university oj nairobi p. 0.30197 nairobi kenya abstract the levels of vitamin a were determined in liver samples from 40 animals including ruminant spesies feeding on grass (4 buffalo, 6 blue wildebeest, 5 gemsbok) or mainly on plant material other than grass (10 impala, 3 springbok, 7 kudu and 5 elephants). the mean values obtained for the ruminant species ranged from 212 to 1217 lu. vitamin a/g liver fresh weight mass and showed no relationship to the percentage of grass in rumen contents. an unusually low level of 6 7 i. u. was found in elephant. introduction the role of vitamin a as an essential precursor of the visual purple is well known. more recent reports have emphasized its importance for growth, fertility and resistance against infections (tiews and hoppe 1972). within certain limits the dietary supply of ~-carotene, the main precursor of vitamin a, is reflected by the storage of vitamin a in the liver (moore 1957). however, definite differences between species are obvious. in a study of the physiological levels of vitamin a in european wild ruminants the question was raised whether liver storage is related to the feeding habits of the various species (briiggemann, drescher-kaden, hoppe and walser-karst 1970). a co-operative investigation of rumen digestive physiology in south african wild ruminants (van gylswyk and giesecke 1973), offered an opportunity to examine liver samples of species feeding either strictly on grass or on plant material other than grass. samples of elephant liver were also made available. 27 material and methods a total of 40 adult animals representing 7 species (4 buffalo syncerus caffer, 6 blue wildebeest connochaetes taurinus, 5 gemsbok oryx gazella, 10 impala aepyceros melampus, 3 springbok antidorcas marsupialis, 7 kudu tragelaphus strepsiceros and 5 elephant loxodonta african a) were sampled during the dry season from the end of july to the end of september 1971 in localities shown in table 1. most liver samples were excised within one hour (in exceptional cases within two hours) after death. the samples were hydrolysed immediately in koh (10% w/v in ethanol) under oz-free conditions, or preserved in liquid nitrogen. after hydrolysis the ether-extraction and spectrophotometric measurement at 325 nm was performed as described elsewhere (briiggemann et al. 1970). results and discussion the results are given in table 1 as mean values with standard errors of the various species. the ruminant species are grouped according to their feeding habits. as indicated by cursory botanical analyses of rumen contents (van gylswyk et ai. 1973), buffalo, blue wildebeest and gemsbok could be termed true grazers, grass representing almost 100% of the diet. impala, springbok and kudu were essentially browsers at this time of the year, as grass constituted less than 10% of the total mass of samples. table i characteri,tio oj anima/; ,ampled and vitamin a content o/liver number percentage of vitamin a animal content ofliver species lo cality 'of sample of grass in rumen (i.u. vitamin animals contents " ai'!!, tl-csh mass) buffalo kruger national park 3 100 365 ± 195"" addo elephant park i 15 blue wilde kruger national park 6 100 i 217 ± 145 beest gemsbok kalahari gemsbok park 3 96 sandh eu wel, s.w.a. 2 212 ± 212 1 rnpala kruger national park 10 4 574 ± 234 springbok kalahari gernsbok park 3 10 619 ± 96 kudu kruger national park 2 7 516±295 thornkloof, eastern cape 5 2 elephant kruger national park 5 67 ± 10 per cent of identifiable plant material on dry mass basis (after van gylswyk and giesecke 1973 ) ':":' mean values ± standard errors 28 the concentration of vitamin a in liver samples varied greatly between species. no obvious relation to the feeding type existed as, with the exception of elephant, both the highest and the lowest levels were found in two of the grazing species viz.. blue wildebeest and gemsbok. the browsers, impala, springbok and kudu showed relatively uniform values of vitamin a storage. it would thus appear that the levels of vitamin a in the liver are mainly governed by factors characteristic of the animal species rather than of the feeding habits. in this context the extremely low content of vitamin a in the liver of elephant, a species mainly feeding on green leaves, is worth nothing. as in cattle, sheep and horses, the mass of the liver of elephants constitutes about 1-1,5% of total body mass (sikes 1971). thus, the low level of vitamin a is not compensated by the mass of the liver relative to body mass. vitamin a storage in the liver of buffalo is of the same order of magnitude as in domestic cattle but considerably lower than in the european wild bovids, mouflon ovis musimon and chamois rupicapra rupicapra with respective mean values of about 1 600 and 800 to 1 500 i. u. per g liver fresh mass (bnlggemann et al.1970). ~-carotene was only found in trace amounts in the livers of species investigated. this is in accordance with domestic ruminants in which the liver storage of ~-carotene is minimal as compared to vitamin a. as indicated by the standard errors of the mean values given in table 1, the variation within species was considerable. a similar degree of variability observed in european wild ruminants could be attributed in part to differences in age, sex and season (briiggemann et al. 1970). the present data, including mostly both sexes and only one season, would not lend themselves for such comparison because of the small numbers of animals examined. in european species an additional important factor was infection with ectoor endoparasites, which was shown to lower vitamin a levels to roughly one third of the normal values depending on the degree of infection. even though the animals sampled in the present study were not examined systematically for parasites, all the elephants were found heavily infected with intestinal parasites as well as the gemsbok obtained from the kalahari gemsbok national park. this may explain in part the low mean values of vitamin a storage in the livers of these species, and a more thorough investigation may be rewarding. acknowledgements the authors are indebted to the national parks board of trustees for permission to undertake the study and to dr. g . de graaff for organizing the excursions. two of the authors (c. b. and d. g.) would like to thank dr. helen m. schwartz and the national chemical research laboratory of the c.s.i.r. for support and encouragement and all colleagues of the d.m.r. unit at onderstepoort for advice and helpful assistance. 29 references bruggemann, j. , u. drescher-kaden, p. hoppe and k. walser-karst. 1970. untersuchungen tiber den vitamin agehalt in der leber von wildtieren. i. mitteilung: vitamin aleberspeicher bei wildlebenden wiederkauem. internat. j. vito res . 40 :249-259. moore, t . 1957. vitamin a, amsterdam : elsevier. sikes, s. k. 1971. the natural history of the african elephant. london: weidenfeld and nicholson. tiews, j. and p. hoppe . 1972. vitamine. world congress of animal nutrition iv :27 5-295. van gylswyk, n. o. and d. giesecke . 1973 . a summary of preliminary findings in a rumen microbiological investigation on wild ruminants . koedoe 16 :191-194. 30 page 1 page 2 page 3 page 4 filelist convert a pdf file! supplement to koedoe. 1977: 186-189. the role of botanic and zoological gardens in national conservation planning h b rycroft director: national botanic gardens of south africa kirstenbosch claremont cape 7735 i am to talk about botanic and zoological gardens and the role they play, or should play, in the conservation of our wild life in its broadest sense. to get everything straight, let us see in the first place what we mean by a botanic and a zoological garden. the simplest definition of a botanic garden is that of the international directory of botanic gardens (2nd ed. fletcher, henderson, and prentice) which simply requires a garden to be open to the public and in which the plants are labelled. c s gager, a former director of the brooklyn botanical garden, new york, defined and described a botanic garden as: "originally a garden with the plants arranged according to some system of botanical classification. a botanic garden is properly defined as a scientific and educational institution, whose purpose is the advancement and diffusion of a knowledge and love of plants . modern gardens usually possess living collections, herbaria, library, experimental gardens for plant breeding, laboratories, lecture halls and in some cases, botanical museums, together with scientific, educational and administrative staffs." the ideal of the national botanic gardens of south africa is primarily an authenticated living collection of indigenous plants, cultivated in gardens. material and facilities for research are provided, knowledge is disseminated to scientists and the general public and visitors are made aware of this floral heritage and encouraged to conserve the south african flora. the broad policy of the institution is scientific and educational and it is concerned with the cultivation, display, scientific study and conservation of the native flora of the republic of south africa, through its efforts and activities, both in the gardens and in the country generally. zoological gardens can be defined in vaguely similar terms, the accent of course being on animals, although plants also playa very significant role in zoos. whatever planning is undertaken for the conservation of our natural resources or any other resource for the benefit of man, the role of plants 186 is paramount. plants, it must be remembered, form the basis of all animal life, including that of man himself. without plants, without vegetation, without green organisms, there can be no animals. we are aware that the ultimate source of all energy on our planet earth, is the sun, and that the principal means by which this energy is harnessed is through plants. approximately 0,1% of the solar energy which reaches us, is fixed by the wonderful and still incompletely understood process of photosynthesis. this fraction, small as it is, results in the production on a world-wide basis, of between 150 and 200 thousand million (150 000 000 000-200 000 000 000) tons of dry organic matter every year. the rapid, even frightening, increase in human population requires simply that an ever larger fraction of the en ergy fixed by plant life be diverted to the support of a single species man himself. the long-term trend of evolution towards building up complex, integrated, stable ecosystems is being reversed before our very eyes, with consequences which we can but dimly foresee, or perhaps fear to an ticipate. man in his efforts to increase his own food supply and to provide living space, resorts to the destruction of natural ecosystems and their replacement with artificial ones and with deserts of concrete, brick, mortar and plastics. the veld is being cleared to make way for biologically sterile industry, townships and sprawling cities connected by equally barren networks of roads and railways. another more forbidding effect of man's activities is that an ever-increasing fraction of the earth's surface is being withdrawn, wholly or partially, from energy fixation as a result of building development and soil deterioration through exhaustion or erosion. the plant life on which we all ultimately depend for our very existence, is being threatened, not only on land but also in lakes and in the oceans. it is imperative that measures be found to halt, and, if possible, to reverse this trend. clearly, research on plants, their metabolism, their ecological relationships and the maintenance of a healthy vegetation cover have become matters of vital necessity to the human race. upon their successful application, man's continuing dominance of the biosphere may ultimately depend. several general principles apply equally to botanic and to zoological gardens. the benefits flowing from them can be comparable and, in this respect, i consider particularly (i) public service, (ii) education, (iii) research and (iv) conservation. uk moore: botanic gardens and arboreta). these various aspects are closely interlinked, but can be viewed separately. (i) public service. let us be clear about this in our own minds. gardens and zoos are established and maintained largely because the man in the street, the general public, you and i want them for the joy of having them. the scientific, educational and cultural benefits, most important 187 though they may be, are not always the main reasons why we have these establishments. gardens of both types offer social, cultural, economic, recreational and aesthetic facilities. (ii) education. many gardens are adjunct to, or actual parts of, educational institutions. as such the organisation provides courses, programmes or facilities in plantand/or animal-related subjects, varying from formal graduate level courses to popular one-day study excursions. these may be: (a) graduate and undergraduate level biology, botany, horticulture and zoology; (b) short courses and workshops on many phases of plant and animal study for the general public and professionals from other disciplines; (c) nature study programmes for school groups; (d) environmental awareness programmes for groups from pre-school age to retired persons; (e) special workshops for teachers of natural sciences; and (f) classes and demonstrations for professional botanists, horticulturists and zoologists. (iii) conservation. with the increased pressure of population and development upsetting or destroying natural areas, gardens are in a position of public and political influence not yet fully realised or exploited . they can playa significant role by: (a) acquisition of natural areas; (b) planting of representative native plants, cultivation of rare and endangered species of plants, breeding of rare and endangered species of animals and preserving portions of the garden estate in their natural state; (c) various courses offered to the public, which stress the importance of natural areas; (d) preservation of various ornamental forms and varieties of plants which may be neglected by nurseries; and (e) co-operative efforts with other conservation groups to lobby for particular legislation or governmental protection. (iv) research. the field of research which can be undertaken is almost limitless. gardens should have the unique advantage of offering indoor laboratory facilities similar to those in most universities and research establishments and, in addition, field conditions to supplement such facilities. in planning conservation on a national basis, gardens and zoos do not provide the ultimate goal or aim in the broadest sense because they present only a fraction or small sample of the whole, in what might be not completely natural conditions. they do, however, provide the essential stepping stones towards the realisation of the most desirable conservation ideals. 188 the individual ecological components must be studied and understood before the total mosaic of the ecosystem can be tackled. for example, the autecology of as many species as possible, both plant and animal, must be investigated. this type of work can be largely undertaken in botanic gardens and zoos, where adequate facilities are available. in building up the mosaic it is essential to call in the assistance and co-operation of disciplines in philosophies other than biology, such as those of the engineer, town planner, geologist, sociologist, meteorologist, economist and even the politician. let's face it, the last named can either make or break any conservation plan or programme! in many instances, the best way to preserve a species is to preserve the habitat in which it occurs naturally. we must decide, however, whether we wish to preserve an individual species or specified number of species or whether we think it desirable to preserve or conserve the community in its entirety. if the latter is the case, we must further decide in what particular condition the community or ecosystem must be maintained . is it the climax condition where we understand that all the species of both the plant and animal kingdom exist and will continue to exist in perfect and stable equilibrium, or is it some seral stage of the succession which can be maintained at that level only by special treatment, manipulation and management? botanic and zoological gardens can provide the facilities for much of the research that is required in attempting to make the decisions and to find the answers. 189 page 1 page 2 page 3 186.pdf page 1 article information authors: leslie r. brown1 pieter j. du preez2 hugo bezuidenhout1,3 george j. bredenkamp4 theo h.c. mostert5 nacelle b. collins6 affiliations: 1applied behavioural ecology and ecosystem research unit, university of south africa, south africa2department of plant sciences, university of the free state, south africa 3south african national parks scientific services, hadison park, kimberley, south africa 4department of plant science, university of pretoria, south africa 5department of botany, university of zululand, south africa 6free state department of economic development, tourism & environmental affairs, bloemfontein, south africa correspondence to: leslie brown postal address: private bag x6, florida 1710, south africa dates: received: 25 july 2012 accepted: 29 apr. 2013 published: 23 july 2013 how to cite this article: brown, l.r., du preez, p.j., bezuidenhout, h., bredenkamp, g.j., mostert, t.h.c. & collins, n.b., 2013, ‘guidelines for phytosociological classifications and descriptions of vegetation in southern africa’, koedoe 55(1), art. #1103, 10 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1103 note: additional supporting information may be found in the online version of this article as an online appendix 1 http://dx.doi.org/10.4102/ koedoe.v55i1.1103-1 and online appendix 2 http://dx.doi.org/10.4102/ koedoe.v55i1.1103-2. copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. guidelines for phytosociological classifications and descriptions of vegetation in southern africa in this checklist... open access • abstract • introduction • purpose of this article • history of phytosociology in south africa • recommended minimum requirements for phytosociological studies in south africa    • approach    • principles    • field survey methods    • data analysis       • vegetation classification       • environmental gradient analysis or ordinations       • other statistical analyses, calculations and comments       • tables       • vegetation maps    • classification and description of plant communities       • classification of plant communities       • naming of plant communities       • description of plant communities       • syntaxonomic descriptions       • recommendations for concluding remarks • concluding remarks and recommendations • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ changes in the environment are first observed in changes in the vegetation. vegetation survey, classification and mapping form the basis on which informed and scientifically defendable decisions on the environment can be taken. the classification and mapping of vegetation is one of the most widely used tools for interpreting complex ecosystems. by identifying different plant communities we are essentially identifying different ecosystems at a particular hierarchical level. phytosociologists in europe have been involved in such studies following, in particular, the braun-blanquet approach since the early 1900s. in south africa, such studies were undertaken on a limited basis from the early 1970s and have since then steadily increased. the surveying of the enormous diversity of south african vegetation is one of the objectives of phytosociological studies. the demand for such data has steadily increased over the past few years to guide conservation policies, biodiversity studies and ecosystem management. in south africa, numerous publications on the vegetation of conservation and other areas in the different biomes have been produced over the last few decades. however, vegetation scientists in south africa experience unique problems. the purpose of this article is therefore to provide an overview of the history and the specific focus of phytosociological studies in south africa and to recommend minimum requirements and methods to be followed when conducting such studies. it is believed that the incorporation of these requirements will result in scientifically justifiable research of high quality by phytosociologists in south africa.conservation implications: effective conservation cannot be obtained without a thorough knowledge of the ecosystems present in an area. consistent vegetation classifications and descriptions form the basis of conservation and monitoring exercises to maintain biodiversity. the incorporation of these guidelines and requirements will facilitate quality phytosociological research in south africa. introduction top ↑ one of the earliest examples of an informal description of southern african vegetation dates back to the late 1400s. in december 1497, vasco da gama sailed from mossel bay in an easterly direction, ensuring that he always had sight of the terrestrial land. interestingly he made no mention of the different wild animals, but instead he continually referred to the vegetation they observed on the land (skead 2011). since then people have used vegetation to assist with finding their way in and around the african continent. vegetation is the most physical representation of the environment (kent 2012). any spatial and temporal changes in habitats are first observed in the vegetation. as a result, the mapping and description of vegetation has either informally or formally played an important role as a tool to classify and interpret different ecosystems. these assessments led to the development of the discipline of vegetation science (plant ecology), and its various fields of specialisation, of which synecology (the classification, description and mapping of vegetation, succession and dynamics) (barbour, burk & pitts 1987) is an important field. synecology stems from observations of the great variety of form and structure of plants that repeat themselves in similar environmental conditions. vegetation scientists started to explore the world to describe and map the diverse plant life of the planet. in the early 20th century most vegetation scientists in europe were occupied with the phytosociological (phyto = plant; sociology = groupings of species) classification, description and mapping of the continents’ vegetation. professor v. westhoff once commented that phytosociology is the science of recognising and identifying the stepping stones in vegetation within the overwhelming swamp of variation (bredenkamp 1982; mostert 2006). currently, vegetation classification and mapping is one of the most widely used tools to assist in the interpretation of complex ecosystems and to simplify the spatial and temporal complexity of these ecosystems (doing 1970; mucina & rutherford 2006). vegetation also fulfils an important function in that it provides food and shelter for wildlife (mucina & rutherford 2006; skead 2011). modern scientific society uses phytosociology either to form the background of scientific studies on animals (e.g. henzi et al. 2011; hirst 1975; pasternak et al. 2013) or to define different plant communities, which could be unique in the sense of rare or endangered plant species (e.g. du preez & brown 2011; edge, cilliers & terblanche 2008; gotze et al. 2008; janecke, du preez & venter 2003). results of phytosociological studies should be the foundation for informed decisions on wildlife management and nature conservation. phytosociology not only serves as the backbone of ecosystem studies but also forms part of the planning for monitoring plant species or communities and rare or endangered ecosystems. vegetation is a collective term for all the plant communities in the same way that flora is a collective term for all the plant species. we describe plant communities, not ‘vegetation communities’. with climate change, phytosociological studies will become more important because, in most cases, only vegetation data are available to use for comparisons. data collected according to accepted quantitative methods, can be used to give detailed information about plant species abundance and also a description of the structure of the vegetation. such data can also be used for gradient analyses, measuring plant species diversity, the study of successional changes and measuring plant production of different ecosystems. vegetation classification is also useful to assist in making informed decisions on the habitat that is available for wildlife, as well as making informed decisions on fire policy and programmes aimed at clearing alien plant species. detailed vegetation classification, mapping and description also form the basis from which informed and scientifically defendable decisions can be taken for infrastructure and other development in an area. it also assists with impact assessment for development purposes. when the expansion of conservation areas is considered, the results of phytosociological studies should be used to assist with planning of these expansions to guide conservation decisions about important, scarce or rare plant communities. at an appropriate scale (association level), plant communities can thus be used as surrogates for ecosystem delineations with very high accuracy. this ability of vegetation to act as a surrogate for ecosystem descriptions or delineations is seated in the fact that vegetation is a highly visible and measurable biological manifestation of all the other environmental factors shaping and driving a specific ecosystem (barbour et al. 1987; kent 2012). phytosociology is used in many different research fields in nature conservation, (e.g. mammalogy, ornithology, herpetology, entomology, geology, soil science, landscape ecology, limnology, etc.) to describe the habitat and give the reader an idea of the (expected) species composition and vegetation structure. it is therefore important that vegetation description studies should be as comprehensive as possible. with the current pressure on the environment, and especially natural vegetation, as a result of mining and agricultural activities. as well as urban and rural development, the need for proper planning cannot be underestimated. although the need for mining or agricultural development sometimes supersedes conservation importance, vegetation classification and description is important to identify ecologically sensitive areas. these studies are currently compulsory in south africa in terms of the national environmental management: biodiversity act (act no. 10 of 2004) (south african government 2004). by identifying different plant communities, we are essentially identifying different ecosystems at a particular hierarchical level. plant communities and their associated vegetation maps are therefore regarded as reliable surrogates for the demarcation of macro-ecosystems. describing, monitoring and managing the surrogate (plant community) are the first steps in effectively monitoring and managing the entire ecosystem, without trying to understand and manage the bewildering detail of all the different components and interactions of the ecosystem. vegetation surveys, classification and mapping provide a framework for understanding the differences between ecosystems (chytrý, schaminee & schwabe 2011). although this approach to ecosystem management is not fail safe or without its shortcomings, it provides ecologists with a sensible, tangible tool and first approximation for the management of ecosystem patterns and processes. purpose of this article top ↑ phytosociological research has a long history. however, vegetation scientists in south africa are experiencing various problems that do not occur elsewhere in the world (see later). the purpose of this article is to provide an overview of the history of phytosociological studies in south africa and to recommend broad guidelines on the minimum requirements and methods to be followed when conducting vegetation classification, description, habitat interpretation studies and mapping following the braun-blanquet approach. history of phytosociology in south africa top ↑ we attempt to present a brief history of the development of phytosociology in south africa using a few examples for explanation, rather than giving a complete review of all the phytosociological research conducted or publications that have appeared on south african vegetation during the last century. most of the earlier national and regional studies were mainly of a non-formal descriptive nature (e.g. bayer 1955; bews 1918; dyer 1937; edwards 1967; killick 1963; louw 1951; muir 1929; pole evans 1922), often providing only species lists of a particular area. acocks (1953) classified the south african vegetation into 70 veld types and 75 variations based on comparison of floristics from stand data. he recorded the abundances of all species. consequently, many south african vegetation scientists used a more flexible approach. statistical numerical classification methods were applied as a first approximation of the plant communities in a particular area, but with the option to ‘refine’ the classification by applying zurich–montpellier methods. relevés could be moved to other clusters (if deemed necessary) by considering more factors (especially total floristic composition, diagnostic species and habitat interpretation) than only those used by the particular numerical algorithm (bredenkamp 1982). application of the twinspan classification algorithm (hill 1979) made a major contribution to obtain more objectivity and repeatability in the classification whilst simultaneously retaining the advantages of a phytosociological table. this provided not only the hierarchical classification and total floristic composition of each plant community at different levels, but also a valuable overview of species cover and abundance, constancy, fidelity and the habitat. the phytosociological table thus became a necessity in every phytosociological study. as computer software became more readily available, the turboveg programme was developed to serve as a standardised format for the storage and management of vegetation data (hennekens 1996; hennekens & schaminee 2001). apart from the pioneering studies of werger (1973) and van der meulen (1979), which were considered to be regional studies, the majority of the earlier studies by south african researchers were more local in extent. the goals of sustainable utilisation linked with effective conservation cannot be achieved without a thorough knowledge of the ecology of a particular area (edwards 1972; werger 1974). it was therefore recommended that conservation policies and environmental management plans should be based on this knowledge (edwards 1972). this resulted in many phytosociological projects initiated in nature conservation areas (e.g. national parks, provincial nature reserves or privately owned and managed game reserves), which resulted in numerous publications on the vegetation of conservation areas in the different biomes of south africa. selected examples include: • grassland: bloem (1988), bredenkamp (1975), coetzee (1972), kay, bredenkamp and theron (1993) and swanepoel (2006). • savanna: bezuidenhout (1994), brown (1997), gertenbach (1978), mostert (2006), stalmans and peel (2010), van rooyen (1983) and van staden (2002). • nama-karoo: brown and bezuidenhout (2000), cleaver, brown and bredenkamp (2005), rubin and palmer (1996), van der walt (1980) and werger and coetzee (1977). • succulent karoo: jurgens (2004) and le roux (1984). • fynbos: boucher (1997), mcdonald (1988), zietsman (2003) and zietsman and bredenkamp (2006). • forest: geldenhuys and murray (1993), grobler (2009) and matthews et al. (2001). • thicket: palmer (1981). • azonal units: collins (2011), du preez and brown (2011) and pretorius (2012). • additional regional studies: bezuidenhout (1993), du preez (1991), eckhardt (1993), fuls (1993), hoare (1997) and kooij (1990). recommended minimum requirements for phytosociological studies in south africa top ↑ to achieve the goal of syntaxonomic synthesis of southern african vegetation and to attain internationally acceptable standards for local studies in, for example, national parks, nature reserves, private game farms and other conservation areas, we strongly recommend the following minimum requirements for phytosociological studies in southern africa. approach we recommend that the zurich–montpellier (braun-blanquet) school of total floristic compositions (braun-blanquet 1932; kent 2012; mueller-dombois & ellenberg 1974; werger 1973; westhoff & van der maarel 1978) should be followed. the main benefit of this approach is that much of the world’s vegetation has been and is continued to be surveyed and classified according to a relatively uniform protocol (chytrý et al. 2011). principles • sample plot placement is not subjective as originally proposed by braun-blanquet (1932), but placed in a stratified random manner within floristically uniform units because it is accepted that each of these units represents a single plant community.• the vegetation in a sample plot must be representative of a single plant community. therefore, a minimum plot size for the particular plant community is used. • the sample plot must be uniform with regard to its biophysical make-up. independent of the scale of the study, floristically homogeneous units that can physically be identified and managed in the field must be used to define plant communities. large scale-related heterogeneous sample plots are avoided as they will result in mixed relevés containing species from different plant communities and a range of biophysical features. these mixed relevés are not according to the basic zurich–montpellier principle of sampling within homogeneous units. mixed relevés are difficult to interpret ecologically and to assign to a specific recognisable plant community. the classification of mixed relevés groups unrelated communities together and may ignore smaller and sometimes threatened or endangered ecosystems. • ideally, all species present in the sample plot should be identified and recorded during the time of survey. owing to seasonality and natural impacts (e.g. grazing, fire, irregular rainfall, erosion, flooding, droughts, etc.), especially in arid and semi-arid areas, some plant species are not easily identifiable (werger 1973). in addition, the rich floristic diversity of south africa and the many cryptic species result in taxonomical problems owing to several undescribed and often unidentifiable species (e.g. some mesembs, orchids, geophytes and annuals). field survey methods the following field survey methods are recommended:• stratified random placement should be determined during the desktop phase whilst preparing for fieldwork. stratification of the vegetation can be performed by means of a geographic information system (gis) using a number of environmental data overlays in combination with satellite and aerial imagery. applicable spatial environmental data overlays include landscapes, land types, terrain units, topography, altitude, climate, geology, soil characteristics, surface rock cover, land use, land cover, or any other relevant spatial data set. overlaying these data sets onto available aerial and satellite imagery will aid in the accurate delineation of relatively homogeneous vegetation units. placement of sample plots within each homogeneous vegetation-cum-habitat unit should be random. in the field, however, the randomly determined location of each sampling plot should be critically evaluated according to the first rule of the zurich–montpellier sampling method (placement of the sampling plot should be within a homogeneous vegetation patch representative of the perceived plant community). if the sampling plot does not fall within a homogeneous representative vegetation stand, it should be moved to the nearest locality that does fulfil this criterion. • a minimum plot size should be determined based on species–area curves produced for each physiognomic–physiographic unit sampled (kent 2012). it is important to note that when gathering data to determine such species–area curves, the first rule of the braun-blanquet method (placement of the nested set of progressively larger plots within a homogeneous vegetation stand, representative of the perceived plant community) should be observed very strictly. alternatively, the researcher may use plot sizes similar to those used in other successful braun-blanquet surveys in similar physiognomic–physiographic units. the researcher should refer to available literature and clearly state the aspects of similarity between the vegetation described with the various plant communities occurring in the study area. the plot sizes given in table 1 may be used as a guideline for the relevant vegetation to be sampled. • for formal syntaxonomic classification, weber, moravec and theurillat (2000) states that only one relevé is required for a formal description of a syntaxon (plant community) although a minimum of ten plots are recommended. to comply with statistical requirements in local phytosociological studies, a minimum of three sampling plots for each of the stratified physiognomic–physiographic units is recommended. exceptions to this suggestion may be in the case of azonal vegetation covering so little of the study area that the placement of more than two plots may lead to pseudo-replication. the required minimum number of plots will depend largely on the size of the plant community in question, being less in smaller plant communities. the description of plant communities based on one or two relevés should be strongly discouraged and such communities should be regarded as doubtful units (weber et al. 2000). • apart from compiling all existing environmental data during the preparation for fieldwork (desktop gis), as many as possible of the following environmental parameters should be measured and determined in the field at each sampling plot during the actual fieldwork phase (recommended but not limited to): geology, surface rock cover, land type, terrain form, soil depth, soil form, soil texture, soil structure, altitude, coordinates, aspect and slope. • soil samples should preferably be collected at the average rooting depth utilised by the vegetation that is being studied. where the soil is classified, the diagnostic horizon from which the sample was taken should be noted. • south africa’s vegetation is highly dependent on rainfall, which is, in most cases, seasonal and erratic. it is recommended that a survey be conducted during the optimal growth period. the period of field survey should be stated clearly. • cover–abundance values for each species recorded within a sample plot should be estimated using one of the many compatible cover–abundance scales used in current phytosociological studies (e.g. modified braun-blanquet cover scale, domin cover scale, plant-number scale) (kent 2012; westfall et al. 1996). in current southern african surveys, the modified braun-blanquet cover–abundance scale is preferred for vegetation classification studies. table 1: suggested minimal area values (m2) for various plant communities. data analysis vegetation classification • it is recommended that phytosociological data are captured in a format that can be imported into programs such as juice (tichý & holt 2006). • algorithms assume that the data have a normal distribution. data should be tested for normality (e.g. using pc-ord [mccune & mefford 2006]) and transformed if the assumption is not satisfied. • either divisive or agglomerative clustering can be used for classification. when using divisive clustering, the modified twinspan (two-way-indicator species analysis [hill 1979]) algorithm proposed by roleček et al. (2009) as contained in juice (tichý 2002) is recommended. unlike the original version, the modified twinspan algorithm does not enforce a dichotomy of classification but instead, at each step, divides only the most heterogeneous cluster of the previous hierarchical level. thus, the application of the modified twinspan algorithm results in vegetation units of similar internal heterogeneity. pseudospecies cut levels used in the classification must be indicated (hotanen 1990). pseudospecies cut levels should consider the scale used during data collection (e.g. where the modified braun-blanquet scale was used, the following pseudospecies cut levels are recommended: 0, 5, 15, 25, 50, 75). • for fidelity measures the use of the phi coefficient of association (chytrý et al. 2002) of each species should be calculated to determine their fidelity to each plant community as an indication of their suitability as diagnostic species. equalisation of relevé cluster sizes (tichý & chytrý 2006) is recommended before calculating the phi coefficient. one can also weigh the relative importance of common and rare species by changing the equalised size of the site groups. this can easily be achieved in the juice program (tichý 2002). it is further recommended that fisher’s exact test be performed simultaneously with the calculation of the phi coefficient to exclude species with non-significant fidelity from the groups of diagnostic species. • description of clusters must include diagnostic, constant and dominant species. it is recommended that these are derived using juice, with 75, 60 and 50 recommended for lower threshold values and 80, 80 and 60 for upper threshold values for diagnostic, constant and dominant species, respectively. in addition, the jaccard and/or sorensen values should be reported. • manual rearrangement of species after automated clustering is permissible. however, rearrangement of clusters and relevés is strongly discouraged. if relevés or clusters were rearranged, it must be accompanied by a detailed justification. the dendrogram should still reflect the original clustering. the modified twinspan classification technique has been described and recommended here owing to its wide use in classification research. it is, however, important to note that other classification tools also exist. optimclass is one of the latest methods used to determine optimal partition between the different data sets of a study area and can also be computed in the juice program (tichý et al. 2010). this method is based on species with a high fidelity, which can subsequently be used as diagnostic species. environmental gradient analysis or ordinations • should the researcher wish to illustrate floristic gradients within and between plant communities or link these gradients with habitat variables, a suitable gradient analysis algorithm should be used, for example canoco, pc-ord and primer.• detrending should be used only where ordination without detrending produces uninterpretable results (e.g. where the objects are clustered together). • it is recommended that papers report on the outcome of procedures followed to determine whether the most important environmental variables were measured (for correspondence analysis and detrended correspondence analysis). it is also recommended that if the number of environmental variables was reduced, the amount of explanatory power lost should be reported. • the decision for the underlying model used during ordination (linear or unimodal) should be noted or explained. other statistical analyses, calculations and comments we encourage further analyses of the data using sound statistical methods to investigate some of the following aspects, where relevant:• diversity • richness • evenness • medicinal and economic value • endemism • conservation status and value • production, grazing or browsing value. tables diagnostic species (see later) are used to arrange plant communities (associations) into a hierarchical classification and are presented in a phytosociological or a synoptic table. the table is the hallmark of the phytosociological study and the interpretation of the environment and its different plant communities. the table is the manifestation of a reliable and accurate research method. the arrangement of the different species groups within the table is therefore of utmost importance (westhoff & van der maarel 1980; werger 1974):• phytosociological table: the phytososiological table should be arranged such that the diagnostic species group for the plant community (association) is above the diagnostic groups of the sub-associations and variants. the species groups are labelled alphabetically with an indication of diagnostic groups for easy reference. species commonly shared amongst communities, probably representing alliances, orders and classes, are normally placed towards the bottom of the table. an example of a typical phytosociological table is included in online appendix 1. • synoptic table: once plant communities have been identified in the phytosociological table, a synoptic table can be produced to summarise the data for each plant community (kent 2012). in a synoptic table, each plant community is summarised in a single column containing values such as fidelity and constancy for each species. an example of a typical synoptic table is included as online appendix 2. one or both of these tables must be used, as long as they contribute to a better understanding of the plant communities identified. it should, however, be noted that phi-coefficient values, used for the synoptic table, are determined from absence/presence data and not cover–abundance data. it therefore means that pseudospecies are not used in the determination of diagnostic species. thus, one cannot solely use these results in the description of the communities, as dominant species with low phi-coefficient values will be omitted from the table. it is therefore important that the researchers use the phytosociological table to describe the dominant and co-dominant species of the community. vegetation maps effective knowledge of the different plant communities and their potential to provide habitats for animals are essential to make scientifically based management decisions for natural areas. the spatial representation of the different plant communities is therefore important. in the past, technologies to provide accurate and detailed information on the location and distribution of plant communities did not exist. owing to technological advances floristically based classifications provide excellent information for the construction of accurate high-resolution vegetation maps (clegg & o’connor 2012; dias, elias & nunes 2004). vegetation maps of natural areas have become indispensable for managers of natural areas. these maps not only indicate the location, distribution and abundance of the different plant communities, together with rare and endemic species, but are also used to monitor changes in cover, structure and composition of the vegetation. it is therefore strongly recommended that vegetation classification and description manuscripts of conservation areas are all accompanied by vegetation maps. classification and description of plant communities plant species names used should follow the latest comprehensive south african plant species list (i.e. currently germishuizen & meyer [2003], with online updates from plants of south africa [posa], which can be accessed via the sanbi website, or the african plants database [version 3.3.4] at http://www.ville-ge.ch/musinfo/bd/cjb/africa). if the authors are of the opinion that the names of certain plant taxa are outdated or inappropriate, the use of newer names for those should be indicated clearly and referenced from the relevant published taxonomic literature.when describing plant communities in normal local habitat classifications (not formal syntaxonomy) the following is recommended. classification of plant communities the classification of different plant communities are based on total floristic composition, although they are recognised by their diagnostic species (character species and differential species). character species are species that are mostly restricted to a specific plant community. they thus characterise the community by their occurrence in one community and by being absent or less frequent in other communities. sub-communities are, in many cases, characterised by the presence or absence of certain species and these are referred to as differential species. these species are used mostly to define the lower syntaxa (westhoff & van der maarel 1980). all diagnostic species identified using ‘objective’ statistics (phi coefficient) should be evaluated with regard to their ‘robustness’ as reliable and predictable indicators of a given plant community. the fewer relevés used to describe a plant community, the more subjective the interpretation of diagnostic species will be. naming of plant communities according to weber et al. (2000) names are only labels to assist in the classification of plant communities and, as such, they will never be wholly adequate. it is, however, more important to understand what is meant by a name than to find one that is characteristic in every respect (weber et al. 2000). it is therefore important that basic rules are followed when naming plant communities to avoid confusion and to enable consistency. the following is therefore proposed:• plant community names are assigned following the same guidelines as presented in the international code of phytosociological nomenclature (weber et al. 2000) for formal syntaxonomical classification, but do not use the specified taxon epithets. according to this rule, the dominant plant name or the one that dominates the structure is second (weber et al. 2000:749 [article 10b]). the first name can be a diagnostic or a co-dominant species. a sub-community will start with the community name followed by a characteristic or dominant species for that sub-community (weber et al. 2000:749 [article 13]). variants can have only one name, usually referring to a diagnostic or dominant species for that variant. only an en-dash (–) is acceptable between the named taxa. example: plant community 1: themeda triandra–acacia karroo community; sub-community 1.1: themeda triandra–acacia karroo–diospyros lycioides sub-community and variant 1.1.1: selaginella dregei variant. • plant community names can have physiognomic descriptors at the end, but the style should be applied consistently. various combinations can be used, with, for example, only the major communities having descriptors. the descriptors are part of the name and should start with capital letters. environmental attributes are not used in the plant community name. example: themeda triandra–acacia karroo woodland or themeda triandra–acacia karroo short open woodland. • in the naming of plant communities, subjective emphasis may be placed on perennial (‘non-fleeting’) species that can be identified and found reliably during most years (not only during unusually good rainfall cycles). such robust species should preferably be used in the names of plant communities whilst short-lived annuals should be avoided. • it is also recommended that existing names for similar communities in related vegetation are retained to prevent several different names for similar communities. this should also assist when formal syntaxonomic classification of vegetation is undertaken. an attempt should therefore be made to avoid unnecessary synonomy. this implies that the researcher must be aware of the relevant literature. referees of papers should strive to identify such synonomy. description of plant communities although a plant community is known by its dominant species, its total floristic composition is characteristic where some species have a greater diagnostic value than others (beard 1980). it is recommended that the description of a plant community follows the standard format, namely starting with the locality and habitat (e.g. geology, land type, soil, topography, rock cover, altitude, erosion). this should be followed by the diagnostic species, which can be either listed or referred to in a table (preferred). the description is continued by listing the prominent (high-cover or abundance) and conspicuous species, their cover, growth form or any other relevant information pertaining to the community that would be useful in identifying and understanding the dynamics within the community. it is very useful to include colour photographs that illustrate typical examples of each plant community. it is highly recommended that the different communities are mapped using gis technology and that the map is included in the paper. syntaxonomic descriptions the need for formal syntaxonomic studies is recognised. smaller and localised data sets are, however, not suitable for formal syntaxonomic descriptions. these data sets need to be combined with other compatible data sets for a particular bioregion (mucina & rutherford 2006) to cover the variation of the various associations present. formal syntaxonomic classification and descriptions are discussed in detail by weber et al. (2000) and their recommendations should be followed for this purpose. recommendations for concluding remarks it has been found that readers of scientific papers pay most attention to the first and last paragraphs of a discussion (wenderoth 2012). it is therefore important that the first paragraph summarises the main findings of the research. in the last paragraph, the significance of the findings should be clearly explained.the researchers can decide whether they want to have separate discussion and conclusion sections (preferred by most journals). however, it is important that the discussion should not repeat what has already been stated in the plant community descriptions; rather, the results (i.e. plant community descriptions) should be discussed in relation to the original objective or problem stated at the beginning of the paper. the results are new knowledge and should be compared critically with similar vegetation studies elsewhere. similarities and differences should be highlighted and possible reasons given and discussed. it is important to refer continually to the results of the study and not to discuss unrelated aspects. other important aspects that can be discussed include endemic species, rare and endangered species, medicinal species, production, veld condition, environmental gradients within and between communities, and biodiversity of different plant communities. to emphasise the importance of phytosociological studies we cannot merely list the different plant communities and sub-communities anymore. therefore, an effort should be made in the discussion to discuss the conservation or biodiversity value and practical implications for the different ecosystems. concluding with applied recommendations increases the possibility that readers from other disciplines would find these studies worthwhile and interesting. it is, however, important that all recommendations be based on sound scientific theory. the conclusion should give a precise statement of the main findings and what they mean. the findings should also be mentioned within the context of previous findings. implications of the study’s findings, shortcomings of the study and further research or research questions should be mentioned. concluding remarks and recommendations top ↑ this article intended to provide broad guidelines for local phytosociological studies in conservation areas in southern africa. we believe that by developing guidelines for vegetation classification and descriptions in the southern african context, the quality and value of papers published will increase whilst, at the same time, preventing confusion between vegetation scientists in southern africa. although older datasets and studies are still recognised and valid, the use of modern analytical software and techniques is strongly encouraged. this will also align southern african vegetation studies with current international trends in vegetation science. local phytosociological studies in south africa are essential for efficient wildlife management programmes and conservation policies for ecosystems and biodiversity within national parks, nature reserves, private game farms and other natural areas. vegetation classification and descriptions provide information to interpret spatial variation between species as well as an understanding of vegetation–environment relationships (clegg & o’connor 2012). in addition, the predicted impacts of climate change on the environment makes the description of the vegetation of an area even more important, as it offers the only record of the current state of the environment. thus, classifications and descriptions offer baseline information for assessing the potential changes in the environment that may result from climate change from a plant and animal perspective (clegg & o’connor 2012). these data sets also play an important role in and contribute largely to formal syntaxonomic studies (luther-mosebach et al. 2012). this is an important reason for recommending a more standardised approach for local phytosociological studies, as it will produce compatible data sets that will assist with compiling a much-needed formal syntaxonomic classification for southern africa. for future development and growth of phytosociology in south africa, the curation of datasets and literature of all phytosociological research should be administered centrally (e.g. by the south african national biodiversity institute or the south african environmental observation network). the organisation should set up, facilitate and manage a system that will enable all researchers in this field to submit and extract electronic data and a list of publications and other literature for particular regions. such an information system will provide researchers with relevant literature and data, which could prevent unnecessary duplication of plant community names or research, and will encourage better communication and collaboration between researchers. ideally, south africa should move closer to a system such as synbiosys europe (http://www.synbiosys.alterra.nl/synbiosyseu) and synbiosys netherlands (schaminée, hennekens & ozinga 2012) to act as a national database. although south africa is currently developing synbiosys fynbos (http://www.synbiosysfynbos.org/program.html) and synbiosys kruger, these are only on a regional scale. acknowledgements top ↑ competing interests the authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article. authors’ contributions l.r.b. (university of south africa) was the project leader and p.j.d.p. (university of the free state), h.b. (sanparks scientific services), g.j.b. (university of pretoria), t.h.c.m. (university of zululand) and n.b.c. (free state department of economic development, tourism & environmental affairs) made conceptual contributions. references top ↑ acocks, j.p.h., 1953, ‘veld types of south africa’, memoirs of the botanical survey of south africa 28, 1–92. barbour, m.g., burk, j.h. & pitts, w.d., 1987, terrestrial plant ecology, benjamin cummings, menlo park, ca. bayer, a.w., 1955, ‘the ecology of grasslands’, in d. meredith (ed.), the grasses and pastures of south africa, pp. 539–550, central news agency, parow. beard, j.s., 1980, ‘the physiognomic approach’, in r.h. whittaker (ed.), classification of plant communities, pp. 33–64, w junk, london. bews, j.w., 1918, the grasses and grassland of south africa, davis, pietermaritzburg. bezuidenhout, h., 1993, ‘syntaxonomy and synecology of western transvaal grasslands, south africa’, phd thesis, department of botany, university of pretoria. bezuidenhout, h., 1994, ‘an ecological study of the major vegetation communities of the vaalbos national park, northern cape i. the than-droogeveld section’, koedoe 37, 19–42. http://dx.doi.org/10.4102/koedoe.v37i2.335 bloem, k.j., 1988, ‘’n plantsosiologiese studie van die verlorenvallei natuurreservaat, transvaal [a phytosociological study of the verlorenvallei nature reserve, transvaal]’, msc thesis, department of botany, university of pretoria. boucher, c., 1997, ‘vegetation description and management guidelines for the blombos private nature reserve, riversdale district’, ecological research report 26, department of botany, university of stellenbosch. braun-blanquet, j., 1932, plant sociology, transl. g.d. fuller & h.s. conard, mcgraw-hill, new york. bredenkamp, g.j., 1975, ‘’n plantsosiologiese studie van die suikerbosrandnatuurreservaat [a phytosociological study of the suikerbosrand nature reserve]’, msc dissertation, department of botany, university of pretoria. bredenkamp, g.j., 1982, ‘’n plantekologiese studie van die manyeleti-wildtuin [a plant ecological study of the manyeleti game reserve]’, dsc thesis, department of botany, university of pretoria. brown, l.r., 1997, ‘a plant ecological and wildlife management plan of the borakalalo nature reserve, north-west province’, phd thesis, department of botany, university of pretoria. brown, l.r. & bezuidenhout, h., 2000, ‘the phytosociology of the farm de rust on the mountain zebra national park, eastern cape’, koedoe 43(1), 1–18. http://dx.doi.org/10.4102/koedoe.v43i1.204 chytrý, m., schaminee, h.j. & schwabe, a., 2011, ‘vegetation survey: a new focus for applied vegetetation science’, applied vegetation science 14, 435–439. http://dx.doi.org/10.1111/j.1654-109x.2011.01154.x chytrý, m., tichý, l., holt, j. & botta-dukát, z., 2002, ‘determination of diagnostic species with statistical fidelity measures’, journal of vegetation science 13, 79–90. http://dx.doi.org/10.1111/j.1654-1103.2002.tb02025.x cleaver, g., brown, l.r. & bredenkamp, g.j., 2005, ‘the phytosociology of the vermaaks, marnewicks and buffelsklip valleys of the kammanassie nature reserve, western cape’, koedoe 48(1), 1–16. http://dx.doi.org/10.4102/koedoe.v48i1.162 clegg, b.w. & o’connor, t.g., 2012, ‘the vegetation of malilangwe wildlife reserve, south-eastern zimbabwe’, african journal of range & forage science 29(3), 109–131. http://dx.doi.org/10.2989/10220119.2012.744352 coetzee, b.j., 1972, ‘’n plantsosiologiese studie van die jack scott-natuurrreservaat [a phytosociological study of the jack scott nature reserve]’, msc dissertation, department of botany, university of pretoria. collins, n.b., 2011, ‘vegetation classification of pans and wetlands, free state province’, phd thesis, department of botany, university of the free state. dias, e., elias, r.b. & nunes, v., 2004, ‘vegetation mapping and nature conservation: a case study in terceira island (azores)’, biodiversity & conservation 13(8), 1519–1539. http://dx.doi.org/10.1023/b:bioc.0000021326.50170.66 doing, h., 1970, ‘the use of phytosociological methods in various parts of the world’, belmontia 4, 53–61. du preez, p.j., 1991, ‘a syntaxonomical and synecological study of the vegetation of the south-eastern orange free state and related areas with special reference to korannaberg’, phd thesis, department of botany, university of the orange free state. du preez, p.j. & brown, l.r., 2011, ‘impact of domestic animals on ecosystem integrity of lesotho high altitude peatlands’, in o. grillo & g. venora (eds.), ecosystems biodiversity, pp. 249–270, intech, n.p. dyer, r.a., 1937, ‘the vegetation of the divisions of albany and bathurst’, memoirs of the botanical surveys of south africa 17, 1–138. eckhardt, h.c., 1993, ‘a synecological study of the vegetation of the north-eastern orange free state’, msc dissertation, department of botany, university of pretoria. edge, d.a., cilliers, s.s. & terblanche, r.f., 2008, ‘vegetation associated with the occurrence of the brenton blue butterfly’, south african journal of science 104, 505–510. http://dx.doi.org/10.1590/s0038-23532008000600028 edwards, d., 1967, ‘a plant ecological survey of the tugela river basin, natal’, memoirs of the botanical surveys of south africa 36, 1–285. edwards, d., 1972, ‘botanical survey and agriculture’, proceedings of the grassland society of south africa 7, 15–19. http://dx.doi.org/10.1080/00725560.1972.9648660 fuls, e.r., 1993, ‘vegetation ecology of the northern orange free state’, phd thesis, department of botany, university of pretoria. geldenhuys, c.j. & murray, b., 1993, ‘floristic and structural composition of hanglip forest in the soutpansberg, northern transvaal’, south african forestry journal 165, 9–20. http://dx.doi.org/10.1080/00382167.1993.9629384 germishuizen, g. & meyer, n.l. (eds.), 2003, ‘plants of southern africa: an annotated checklist’, strelitzia 14, national botanical institute, pretoria. pmid:16468931 gertenbach, w.p.d., 1978, ‘plantgemeenskappe van die gabbro-kompleks in die noordweste van die sentrale distrik van die nasionale krugerwildtuin [plant communities of the gabbro-complex in the north-western section of the central district of the kruger national park]’, msc dissertation, department of botany, potchefstroom university for christian higher education. gotze, a.r., cilliers, s.s., bezuidenhout, h. & kellner, k., 2008, ‘analysis of the vegetation of the sandstone ridges (ib land type) of the north-eastern parts of the mapungubwe national park, limpopo province, south africa’, koedoe 50(1), 72–81. http://dx.doi.org/10.4102/koedoe.v50i1.136 grobler, l.e.r., 2009, ‘a phytosociological study of peat swamp forests in the kosi bay lake system, maputaland, south africa’, msc thesis, department of botany, university of pretoria. hennekens, s.m., 1996, turboveg: software package for input, processing and presentation of phytosociological data, ibn-dlo, wageningen, university of lancaster. hennekens, s.m. & schaminee, j.h.j., 2001, ‘turboveg: a comprehensive database management system for vegetation data’, journal of vegetation science 12, 589–591. http://dx.doi.org/10.2307/3237010 henzi, s.p., brown, l.r., barrett, l. & marais, a.j., 2011, ‘troop size, habitat use, and diet of chacma baboons (papio hamadyas urisinus) in commercial pine plantations: implications for management’, international journal of primatology 32, 1020–1032. http://dx.doi.org/10.1007/s10764-011-9519-6 hill, m.o., 1979, twinspan: a fortran program for arranging multivariate data in an ordered two-way table by classification of individuals and attributes, cornell university, new york. hirst, s.m., 1975, ‘ungulate-habitat relationships in a south african woodland/savanna ecosystem’, wildlife monographs 44, 1–40. hoare, d.b., 1997, ‘syntaxonomy and synecology of the grasslands of the southern parts of the eastern cape’, msc thesis, department of botany, university of pretoria. hotanen, j.p., 1990, ‘the effect of pseudospecies cut level settings on the results of twinspan classification’, suo 41(2), 43–53. janecke, b.b., du preez, p.j. & venter, h.j.t., 2003, ‘vegetation ecology of the pans (playas) of soetdoring nature reserve, free state’, south african journal of botany 69(3), 401–409. jurgens, n., 2004, ‘a first classification of the vegetation of the richtersveld (rsa) and directly adjacent regions in in namibia and south africa’, schumannia 4, 149–180. kay, c., bredenkamp, g.j. & theron, g.k., 1993, ‘the plant communities of the golden gate highlands national park in the north-eastern orange free state’, south african journal of botany 59(4), 442–449. kent, m., 2012, vegetation description and analysis: a practical approach, wiley-blackwell, oxford. kershaw, k.a., 1964, ‘quantitative and dynamic ecology’, edward arnold, london. killick, d.b.h., 1963, ‘an account of the plant ecology of the cathedral peak areas of the natal drakensberg’, memoirs of the botanical surveys of south africa 34, 1–178. kooij, m.s., 1990, ‘a phytosociological study of the north-western orange free state’, msc dissertation, department of botany, university of pretoria. le roux, a., 1984, ‘’n fitososiologiese studie van die hester malan natuurreservaat [a phytosociological study of the hester malan nature reserve]’, msc thesis, department of botany, university of pretoria. louw, w.j., 1951, ‘an ecological account of the vegetation of the potchefstroom area’, memoirs of the botanical surveys of south africa 24, 1–105. luther-mosebach, j., dengler, j., schmiedel, u., röwer, i.u., labitzky, t. & gröngröft, a., 2012, ‘a first formal classification of the hardeveld vegetation in namaqualand, south africa’, applied vegetation science 15(3), 401–431. http://dx.doi.org/10.1111/j.1654-109x.2011.01173.x matthews, w.s., van wyk, a.e., van rooyen, n. & botha, g.a., 2001, ‘vegetation of the tembe elephant park, maputaland, south africa’, south african journal of botany 67, 573–594. mccune, b. & mefford, m.j., 2006, pc-ord. multivariate analysis of ecological data, version 6, mjm software, gleneden beach, oregon. mcdonald, d.j., 1988, ‘a synopsis of the plant communities of the swartboschkloof, jonkershoek, cape province’, bothalia 18, 233–260. mostert, t.h.c., 2006, ‘vegetation ecology of the soutpansbergand blouberg area in the limpopo province’, phd thesis, department of botany, university of pretoria. mucina, l. & rutherford, m.c., 2006, ‘the vegetation of south africa, lesotho and swaziland’, strelitzia 19, south african national biodiversity institute, pretoria. mueller-dombois, d. & ellenberg, h., 1974, aims and methods of vegetation ecology, wiley, new york. muir, j., 1929, ‘the vegetation of the riversdale area, cape province’, memoirs of the botanical surveys of south africa 13, 1–82. palmer, a.r., 1981, ‘a study of the vegetation of the andries vosloo kudu reserve, cape province’, msc thesis, department of botany, rhodes university. pasternak, g., brown, l.r., kienzle, s., fuller, a., barrett, l. & henzi, s.p., 2013, ‘population ecology of vervet monkeys in a high latitude, semi-arid riparian woodland’, koedoe 54(1), art. #1078, 9 pages. http://dx.doi.org/10.4102/koedoe.v54i1.1078 pole evans, i.b., 1922, ‘the main botanical regions of south africa’, memoirs of the botanical survey of south africa 4, 49–53. poore, m.e.d., 1956, ‘the use of phytosociological methods in ecological investigations iv: general discussion of phytosociological problems’, journal of ecology 44, 28–50. http://dx.doi.org/10.2307/2257153 pretorius, m.l., 2012, ‘a vegetation classification and description of five wetland systems and their respective zones on the maputaland coastal plain’, msc dissertation, department of environmental sciences, university of south africa. roleček, j., tichý, l., zelený, d. & chytrý, m., 2009, ‘modified twinspan 986 classification in which the hierarchy respects cluster heterogeneity’, journal of vegetation science 20, 596–602. http://dx.doi.org/10.1111/j.1654-1103.2009.01062.x rubin, f. & palmer, a.r., 1996, ‘the physical environment and major plant communities of the karoo national park, south africa’, koedoe 39, 25–52. http://dx.doi.org/10.4102/koedoe.v39i2.293 schaminée, j.h.j., hennekens, s.m. & ozinga, w.a., 2012, ‘the dutch national vegetation database’, in j. dengler, j. oldeland, f. jansen, m. chytrý, j. ewald, m. finckh et al. (eds.), vegetation databases for the 21st century, biodiversity & ecology 4, 201–209. http://dx.doi.org/10.7809/b-e.00077 skead, j.c., 2011, historical incidences of larger land mammals in the broader western and northern cape, centre for african conservation ecology, nelson mandela metropolitan university. south african government, 2004, national environmental management: biodiversity act no. 10, south african government, pretoria. stalmans, m. & peel, m., 2010, ‘plant communities and landscapes of the parque nacional de zinave, mozambique’, koedoe 52(1), 1–11. http://dx.doi.org/10.4102/koedoe.v52i1.703 swanepoel, b.a., 2006, ‘the vegetation ecology of ezemvelo nature reserve bronkhorstspruit south africa’, msc dissertation, university of pretoria. tichý, l., 2002, ‘juice, software for vegetation classification’, journal of vegetation science 13(3), 451–453. http://dx.doi.org/10.1111/j.1654-1103.2002.tb02069.x tichý, l. & chytrý, m., 2006, ‘statistical determination of diagnostic species for site groups of unequal size’, journal of vegetation science 17, 809–818. tichý, l. & holt, j., 2006, juice. program for management, analysis and classification of ecological data, masaryk university, brno. tichý, l., chytrý, m., hájek, m., talbot, s.s. & botta-dukát, z., 2010, ’optimclass: using species-to-cluster fidelity to determine the optimal partition in classification of ecological communities’, journal of vegetation science 21, 287–299. http://dx.doi.org/10.1111/j.1654-1103.2009.01143.x van der meulen, f., 1979, ‘plant sociology of the western transvaal bushveld, south africa, a syntaxonomic and synecological study’, dissertiones botanicae 49, 1–191. van der walt, p.t., 1980, ‘a phytosociological reconnaissance of the mountain zebra national park’, koedoe 23, 1–32. http://dx.doi.org/10.4102/koedoe.v23i1.632 van rooyen, n., 1983, ‘die plantegroei van die roodeplaatdam natuurreservaat ii. die plantgemeenskappe [the vegetation of the roodeplaatdam nature reserve ii. the plant communities]’, south african journal of botany 2, 115–125. van staden, p.j., 2002, ‘an ecological study of the plant communities of the marakele national park’, msc thesis, department of botany, university of pretoria. weber, h.e., moravec, j. & theurillat, j.p., 2000, ‘international code of phytosociological nomenclature. 3rd edition’, journal of vegetation science 1, 739–768. wenderoth, p., 2012, how to write a scientific research paper, viewed 10 may 2012, from http://vision.psy.mq.edu.au/~peterw werger, m.j.a., 1973, ‘phytosociology of the upper orange river valley, south africa: a syntaxonomical and synecological study’, dsc thesis, science department, catholic university of nijmegen. werger, m.j.a., 1974, ‘on concepts and techniques applied in the zurich-montpellier method of vegetation survey’, bothalia 11, 309–323. werger, m.j.a. & coetzee, b.j., 1977, ‘a phytosociological and phytogeographical study of the augrabies national park, republic of south africa’, koedoe 20, 11–51. http://dx.doi.org/10.4102/koedoe.v20i1.929 westfall, r.h., van staden, j.m., panagos, m.d., breytenbach, p.j.j. & greeff, a., 1996, ‘scale related vegetation sampling’, arc-range and forage institute, pretoria. westhoff, v. & van der maarel, e., 1978, ‘the braun-blanquet approach’, in r.h. whittaker (ed.), classification of plant communities, pp. 289–399, w junk, the hague. http://dx.doi.org/10.1007/978-94-009-9183-5_9 westhoff, v. & van der maarel, e., 1980, ‘the braun-blanquet approach’, in r.h. whittaker (ed.), classification of plant communities, pp. 287–399, w junk, the hague. zietsman, m.m., 2003, ‘phytosociological study of andrews field and tsaba-tsaba nature reserve, bredasdorp district, western cape’, msc dissertation, department of botany, university of pretoria. zietsman, m.m. & bredenkamp, g.j., 2006, ‘dune vegetation and coastal thicket communities in threatened limestone fynbos of andrews field and tsaba-tsaba nature reserve struisbaai, western cape’, koedoe 49, 33–47. http://dx.doi.org/10.4102/koedoe.v49i1.110 nel.qxd habitat, food, and small mammal community structure in namaqualand m. van deventer and j.a.j. nel van deventer, m. and j.a.j. nel. 2006. habitat, food, and small mammal community structure in namaqualand. koedoe 49(1): 99–109. pretoria. issn 0075-6458. the effect of habitat differences and food availability on small mammal (rodent and elephant shrew) species richness, diversity, density and biomass was investigated in namaqualand, south africa. species richness in the three habitats sampled, namely upland succulent karoo, dry riverine shrub and north-western mountain renosterveld was low, with only 2–4 species per habitat. rodents trapped were predominantly gerbillurus paeba and aethomys namaquensis, with fewer mus minutoides and petromyscus sp. the only non-rodent was the elephant shrew elephantulus edwardii. ten habitat features, the percentage of total plant cover, tree cover, shrub cover, grass cover, plant litter, total basal cover, sand, gravel or rock cover, and the dominant plant height were recorded at 30 randomly chosen points on five sampling grids in each habitat. small mammal density and biomass was significantly correlated with food availability (green foliage cover, seeds, and relative density and biomass of insects). species richness and diversity of small mammals were significantly correlated with shrub cover. numbers and biomass of specific species correlated significantly with different habitat features in each case. key words: habitat features, small mammals, species richness, diversity, namaqualand. m. van deventer and j.a.j. nel , department of botany and zoology, university of stellenbosch, private bag x1, matieland, 7602 republic of south africa (jan@sun.ac.za). issn 0075-6458 99 koedoe 49/1 (2006) introduction small mammals, especially rodents, are often abundant, widely distributed and important components of nearly all terrestrial ecosystems, especially in semi-arid or arid areas (brown et al. 1986; price & jenkins 1986; kerley & erasmus 1992). rodents are consumers of both seeds and herbage (price & jenkins 1986; kam et al. 1997) and also serves as prey to carnivores. although often omnivorous (landry 1970; kerley 1992a), they can also be predominantly granivorous, especially in semi-arid and arid ecosystems (brown 1975; mares 1976; brown et al. 1979; bar et al. 1984). the degree of granivory differs between deserts, being higher in north american and israeli deserts than in south american, southern african and australian deserts (kerley 1992a, 1992b). coexistence of granivorous rodent species can be facilitated by size differences (bowers & brown 1982), although in some cases overlap in resource use can be low (brown & lieberman 1973). where seed consumption by different-sized species is high, selective granivory can affect the composition of the plant community (brown et al. 1986). small mammals, especially rodents, may be reliable indicator species because they are abundant enough to allow meaningful statistical analysis of data, belong to different trophic levels and functional guilds, and are easily studied (brown 1975). in addition, their community characteristics can reflect ecological disturbance in, e.g., conservation areas (avenant 2000). in spite of this, small mammals are still often overlooked in the planning and conservation of an area (els & kerley 1996). moreover, in nel.qxd 2006/04/17 11:00 pm page 99 south africa the ecological factors that influence the structure of small mammal communities in geographically separate areas, even with broadly similar habitats, have hardly been investigated (kerley 1992c). rodent species diversity in deserts has been linked to habitat heterogeneity (rozenzweig & winakur 1969), productivity (brown 1973) or both (abramsky 1978, 1988). however, species diversity can vary depending on mean annual rainfall, with lower precipitation depressing species numbers and density, and vice versa (nel 1983; abramsky 1988). this would be a reflection of changes in primary productivity (rutherford 1980). in addition, area of habitat has been shown to affect species numbers (ricklefs & lovette 1999; nee & cotgreave 2002; allen & white 2003). williams et al. (2002) state that within limitations set by the regional species pool, local diversity is determined by the habitat capacity (the product of productivity and both spatial and temporal habitat diversity), the habitat itself and the length of time over which the environment has been relatively stable. ricklefs & lovette (1999) concluded that habitat-specialised, largepopulation species are strongly affected by habitat diversity while, in contrast, species with weak habitat specialisation and lowpopulation densities show a strong area effect. habitat preference is a reflection of a species’s physiological (bowers & brown 1982) and nutritional needs (kerley & erasmus 1992; kerr & packer 1997) set by social interactions (kotler & brown 1988), antipredation factors, competitive ability (hughes & ward 1993; hughes et al. 1994; kotler 1984; lancaster 1996) and successional roles of the animals (els & kerley 1996). it has an adaptive basis (martin 1998) and ultimately leads to particular species being more abundant in some habitats than others so as to maximise their lifetime reproductive success (cramer & willig 2002). in addition, a number of studies have demonstrated that small mammal community structure is a function of plant architecture (rosenzweig & winakur 1969; kerley 1992a). as rodents must adapt or became extirpated because they are non-vagile (mares & hulse 1997) a relationship between rodent density and spatial variations, and other demographic parameters on the one hand and habitat heterogeneity on the other can exist (cueto et al.1995). namaqualand covers ~ 55 000 km² (van rooyen et al.1992) and is part of the succulent karoo biome in south africa (cowling & pierce 1999). only 1.6 % of this semidesert region has been set aside for formal conservation (boonzaaier et al. 2002). this area is rich in succulent flora (van rooyen et al. 1991; esler & rundel 1997) but few data are available on its small mammals (dieckmann 1979; joubert & ryan 1999; schradin 2005; nel unpubl. data) with even less on small mammal habitat relations and community characteristics. here we investigate the effect of habitat features on the numbers and species richness of small mammals, and the effect of food availability on their densities and biomass. we quantify habitat use by small mammals (rodents and elephant shrews of < 200 g) in three vegetation types (≅ habitats). variables quantified were habitat features, food availability, and small mammal community structure. we determined: a) small mammal species richness, density and biomass in each of the three habitats; b) the numerical contribution of each species to the community in each habitat (species diversity and evenness of representation); obtained c) a measure of the variation between habitats; and d) estimated food availability (relative abundance of green foliage, seed and insect numbers, as well as insect biomass). areas of the chosen vegetation types (≅ habitats) present within the park were estimated, and the relationship between small mammal community characteristics and habitat and food availability, respectively, determined. in addition, the relative contribution of habitat variation and food availability to species richness, density and biomass was determined. koedoe 49/1 (2006) 100 issn 0075-6458 nel.qxd 2006/04/17 11:00 pm page 100 study area our study area was the namaqua national park (nnp), about 495 km north of cape town and 22 km northwest of kamieskroon. the park was proclaimed in 1988, with the farm skilpad (30°09'57"s, 17º47'53"e) as the nucleus. the park was recently enlarged to ca 73619 ha (a. rossouw pers. comm.). this semi-desert area is hot with dry summers somewhat tempered by onshore movements of cold air from benguela current of the south atlantic. winters are cold but rainfall sparse and erratic (schulze 1947), with the highest precipitation—predominantly fog—from may to august. precipitation increases from west (~ 50 mm p.a.) to east (> 400 mm in the kamiesberg) (cowling et al. 1999). at skilpad on the eastern edge of the park, rainfall varied from 603.5 mm in 1986 to 142 mm in 2000, with an average of 332.8 mm per year. winter temperatures vary between 7–19 ºc, and summer temperatures between 20–32 ºc (schulze 1947). there is also a west to east altitude gradient in the park, from 180 m a.s.l in the far west, rising to ca 300 m a.s.l. at melkboom and then to 750 m a.s.l. at skilpad in the foothills of the kamiesberg. the geology of the namaqua rocky hills is characterised by granite gneiss of the namaqualand metamorphic province (cowling et al. 1999), and the patchy topography is characterised by broken granite kopjes and plateaux separated by sandy or gravel plains (dieckmann 1979). the soil is lime rich, weakly developed on a rock substrate (low & rebelo 1996), and a light reddish-brown sand to light sandy loam covered, in some parts, by a thin layer of white quartz pebbles or gravel (dieckmann 1979). dwarf, succulent shrubs and ephemeral species dominate the vegetation (low & rebelo 1996). the numerous ephemeral species (van rooyen et al.1990) are able to complete their life cycles rapidly during periods when temperature and moisture conditions are favourable for growth (van rooyen et al. 1991). their leaves, flowers and seeds are important food sources for the resident mammals (dieckmann 1979). methods three sites, each situated in a distinct habitat (vegetation type), were sampled, i.e., in upland succulent karoo, dry riverine shrub and north-western mountain renosterveld. the upland succulent karoo or namaqua broken veld is characterised by dome-shaped granite hills. succulent species, particularly of the family mesembryanthemaceae dominate the granite and gneiss-rich soils; grasses are uncommon (low & rebelo 1996) although many species occur. this habitat covers 25767 ha in the park, and grid altitude varied from 308–343 m a.s.l., with aspects facing w, nw, n, ne and e. dry riverine shrub and sweet thorn (acacia karroo) borders seasonal streams (theron et al. 1993). this habitat, part of the upland succulent karoo of low & rebelo(1996), is here regarded as a distinct habitat for small mammals. it is characterised by deep sandy soils with grasses, woody shrubs and a. karroo. it covers an area of 3681 ha in the park. grids were at altitudes from 260–285 m a.s.l., with aspects facing s-ne. the north-western mountain renosterveld or mountain renosterveld is characterised by deep sandy loam soil with renosterbos elytropappus rhinocerotis the main plant growth form (low & rebelo 1996). this habitat covers 7362 ha in the park, with grids at 707–751 m a.s.l. aspects were e, se and nw. at each site, small mammals, habitat features and food availability were sampled in five grids: a large, main one (a) with 70 trap stations, and three subsidiary, smaller ones (b–d) with 20 trap stations each, while a similar grid (e) also sampled small mammals to determine reproductive status and diet. at each site (i.e., habitat), sampling grids were separated by 100 m to gauge intra-habitat variation, while sites were 10 km apart and probably have a shared biogeographic history (vide williams et al. 2002). all three sites were sampled between 2–19 september 2003 to minimise temporal variation. small mammals were sampled by means of the capturemark-release-recapture method. aluminium sherman live traps (230 x 75 x 90 mm) were used, with traps spaced 15 m apart in seven rows (grid a), or two rows (grid b–e), and with ten stations per row, i.e. 150 trapping stations per site. in addition, habitat characteristics were measured at randomly selected issn 0075-6458 101 koedoe 49/1 (2006) nel.qxd 2006/04/17 11:00 pm page 101 trapping stations within each grid. voucher specimens for stomach analyses and an accurate determination of reproductive condition (nel et al. 1984; rautenbach & nel 1980) were collected on grid e. trap stations were 15 m apart, giving a sampling plot of 135 x 90m (1.215 ha) for grid a, and 135 x 15 m (0.2025 ha) each for grids b–e. traps were numbered, and baited with a mixture of rolled oats, peanut butter, golden syrup and bovril (meat and vegetable extract) (atkinson 1997; els & kerley 1996; joubert & ryan 1999). traps were set for five consecutive nights and four days, except for grid b (four nights) in the upland succulent karoo and all the grids in the dry riverine shrub (four nights and three days), to minimise bias due to variability in the relative trapability of each species (williams et al. 2002). trapping sessions avoided the night of full moon or nights before and after it (jensen & honess 1995). traps were checked, rebaited and reset early each morning and during the late afternoons. captured small mammals were identified, marked by an unique toe-clipped number (kotler 1984), weighed, sexed, reproductive condition noted and released at the point of capture. trapping protocol was accepted by south african national parks. one taxon, petromyscus, could be identified only to genus level. nomenclature follows skinner & smithers (1990). habitat structure in relation to vegetation structure was described at sampling points randomly selected (els & kerley 1996; williams et al. 2002). a robust wire square (1 m²) (kent & coker 1992) was dropped with one corner in a northerly direction from the trap at the 30 selected trap stations in each habitat. the latitude, longitude, and elevation (using gps), aspect, and visual estimates of percentage vegetation cover by trees, shrubs, and grass; plant litter, edaphic composition (percentage of sand/ gravel/ rock), as well as the height of the dominant plants (i.e. plant species covering most of the area) were recorded. this yielded a measurement of habitat features at different sampling points and combined for the grids in a habitat allowed between-habitat differences to be recognised. food availability, taken as foliage, seed and insect availability (kerley 1992b) was recorded at the same points as the environmental features. a measure of crude seed density within the 1 m² sampling plots was obtained using five categories: 0 = no seeds/m²; 1 = 1–25 seeds/m²; 2 = 26–75 seeds/m²; 3 = 76– 150 seeds/m²; and 4 = >150 seeds/m². foliage availability represented the percentage green foliage cover within the random squares. insect availability at sampling points was measured using pitfall traps (8 cm diameter x 15 cm deep plastic containers) halffilled filled with water and dishwashing liquid, and set at the randomly selected points for a five-day period (cueto et al. 1995). the insects were then removed, preserved in 70 % ethanol, dried, counted, and weighed. as duration of sampling differed between habitats, insect abundance was calculated using a relative density index (rdi = number of caught individuals/number of traps x number of days involved). combining values for the three categories gave a crude indication of food availability in each habitat. measurements of small mammal populations included trapping success, species richness, species diversity and evenness of representation, abundance, density and biomass. trapping success (or percentage capture rate) was taken as the number of small mammals captured per 100 trap nights (one trap set for a 12 h period = one trap night (nel 1978)). species richness expressed the number of trapped species. species diversity (number of species and numerical contribution of each to the community) was calculated using the shannon-wiener diversity index (krebs 1999). for a measure of population size the schumachereschmeyer method of abundance (a modification of the lincoln index which measures the ratio of marked and unmarked individuals in the population) was used (krebs 1999). this assumes that the population size remains constant over the sampling period, that sampling is random and that all individuals have an equal chance of capture. for our short study period these assumptions were probably justified. in addition, this method takes into account non-random mixing of marked and unmarked individuals. small mammal density (individuals captured/ha) and biomass (total mass (g) of all individuals/ ha) were also calculated. diet was deduced from stomach analysis, which provide only a relative measure of a species’ diet by estimating the contribution (expressed as %) of seeds (white material), foliage (green material) and insects (black material). contents of stomachs were mixed and evenly spread in water in a petri dish placed on a grid. the percentage white, green and brown material within six squares were estimated using a stereomicroscope, and the means calculated. although this is a crude approximation of the actual diet, some idea of the food resources utilised by the different species present at each locality sampled can be gained (rautenbach & nel 1980). correlation analyses (sokal & rohlf 1995) helped investigate relations between small mammal community characteristics and environmental features and between features. the 95 % level (p < 0.05) was koedoe 49/1 (2006) 102 issn 0075-6458 nel.qxd 2006/04/17 11:00 pm page 102 regarded as statistically significant, and applies wherever the term ‘significant’ is used below. results trapping success for 4230 trap nights averaged 4.1 % (range 0–12.2 % within habitats). during 258 capture events 151 individuals (excluding recaptures) of five small mammal species (four rodent and one macroscelidid) were captured. rodents dominated the catch at all sites in terms of numbers, trap success and biomass (tables 1 & 2). species richness ranged from 2–4 per habitat, and 0–3 on any one grid within a habitat. the hairy-footed gerbil, gerbillurus paeba (n = 84, 55.6 %) dominated the community, followed by the namaqua rock mouse aethomys namaquensis (48, 31.8 %). other species captured were petromyscus sp. (n = 5), mus indutus (n =1) and elephantulus edwardii (n =13). the recapture rate (number of recaptures/100 trap nights) was 3.6 %. population size or abundance estimates (mean n = 10; range 0–27), densities (mean = 24.9/ha; range 0–54.3/ha) and biomass (mean = 1314.3 g; range 155– 3859 g/ha) of small mammals captured varied between and within habitats (tables 1 & 2). only g. paeba occurred in all three habitats; a. namaquensis and e. edwardii in upland succulent karoo and north-westissn 0075-6458 103 koedoe 49/1 (2006) table 1 small mammal community attributes in upland succulent karoo (usk); dry riverine shrub (drs); and north western mountain renosterveld (nwmr) in the namaqua national park. south africa. mean (and range values, where applicable) reflect sampling in five grids, in each habitat usk drs nwmr trap nights (total) 1510 1190 1530 trapping success (%) 4.4 (3.7–12.2) 1.9 (0–6.4) 4.5 (3.9–11.7) no. individuals caught 60 (8–20) 22 (0–8) 69 (8–26) density (n/ha) 29.6 (12–49) 10.9 (0–20) 34.1 (21–54) biomass (g/ha) 1991 (399-3859) 459 (155-1071) 1493 (740-2210) population size (n) 11 (8–17) 5 (2–10) 15 (8–27) species richness 4 2 3 species diversity h' 1.78 0.658 0.971 evenness e 0.888 0.415 0.612 table 2 density (n/ha) and biomass (g/ha) of small mammal species in upland succulent karoo (usk), dry riverine shrub (drs), and north-western mountain renosterveld (nwmr) in the namaqua national park, south africa. figures reflect means from 5 grids in each habitat. total numbers caught in parentheses species habitat usk drs nwmr aethomys namaquensis (28) (20) density 13.87 9.88 biomass 1012.4 584.2 gerbillurus paeba (15) (21) (48) density 7.41 10.37 23.7 biomass 296.1 517.8 851.8 mus minutoides (1) density 0.49 biomass 0.65 petromyscus sp. (5) density 2.47 biomass 64.2 elephantulus edwardii (12) (1) density 5.93 0.49 biomass 618.3 57.3 ern mountain renosterveld, while petromyscus sp. and mus indutus were only recorded in the upland succulent karoo and dry riverine shrub habitats respectively. tables 1 & 2 give relative capture frequencies and small mammal nel.qxd 2006/04/17 11:00 pm page 103 community characteristics in the different habitats. total plant cover was the highest in the north-western mountain renosterveld and lowest in upland succulent karoo (table 3). shrubs dominated in all habitats. plant litter cover and visible edaphic cover (sand, gravel or rock) was highest in the upland succulent karoo, while the latter was lowest in north-western mountain renosterveld, with gravel predominating the non-vegetated areas. the highest dominant plants occurred in the renosterveld, and the lowest in the dry riverine shrub. there was a significant positive correlation between total plant cover and the height of the dominant plants (r = 0.762), and total visible rock cover (r = 0.707). the means and range of green foliage cover were of 46.6 % (34–57), 69.6 % (62–78) and 66.4 % (47–76) for upland succulent karoo, dry riverine shrub and north-western mountain renosterveld, respectively. the relative density index (rdi) for insects for the same habitats were (means and ranges) 15.77 (2.6–27.95), 16.9 (12–25.28) and 9.37 koedoe 49/1 (2006) 104 issn 0075-6458 table 3 mean values of each habitat variable for upland succulent karoo (usk); dry riverine shrub (drs); and north-western mountain renosterveld (nwmr) in the namaqua national park, south africa. values reflect the mean of 30 sampling points in each habitat usk drs nwmr total cover (%) 24.8 25.0 49.6 tree cover (%) 1.0 11.8 12.4 shrub cover (%) (succulents included) 86.2 49.0 69.6 grass cover (%) 12.8 39.2 18.0 plant litter (%) 53.4 30.4 33.6 total basal cover (%) 43.0 25.2 24.8 sand (%) 6.4 9.8 27.6 gravel (%) 65.4 81.6 68.0 rock (%) 28.2 8.6 4.4 dominant plant growth height (m) 0.58 0.46 1.1 (5.6–16.6), while mean mass (and ranges) were 0.3938 g (0.0406–0.5415), 0.0981 g (0.0297–0.2629) and 0.0880 g (0.0108– 0.2514). no seeds were present in the sampling squares in any of the habitats. species diversity and species richness were both positively and significantly correlated with shrub cover (r = 0.848 and r = 0.702, respectively) while elephant shrew biomass was positively and significantly correlated with both total visible edaphic cover and rock cover (r = 0.857 and r = 0.744, respectively). a number of the small mammal community-environment features were negatively correlated, but significantly so only for species diversity and grass, evenness and grass, density and grass, species richness and grass and total rodent biomass and grass. only g. paeba, a. namaquensis and e. edwardii were caught in sufficient numbers to analyse the relation between numbers and biomass with environmental features. gerbillurus paeba numbers and biomass correlated significantly with total plant cover (r = 0.605 and r = 0.845, respectively), and biomass also with height of the dominant plants (r = 0.829). aethomys namaquensis numbers correlated significantly with shrub cover (r = 0.787), and their biomass with rock cover (r = 0.593). in addition, numbers and biomass of e. edwardii correlated significantly with total visible edaphic cover (r = 0.942 and r = 0.857, respectively), rock cover (r = 0.737 and r = 0.744, respectively) and insect mass (r = 0.714 and r = 0.615, respectively). the areas of the three habitats in the park varied from 3681 ha to 25767 ha. all three extended widely outside the park so that the island biography theory cannot be invoked; specific small mammal species occurred in a variety of habitats. comparing species richness, density and biomass of the small mammals with habitat characteristics and food availability showed different patterns. food availability had a strong effect on density (r = 0.747, 0.01>p>0.001) and biomass (0.714, 0.01>p>0.001) ). habitat characteristics did not show any significant correlanel.qxd 2006/04/17 11:00 pm page 104 tion with density or biomass, but shrub cover was significantly correlated with species richness and diversity (r = 0.702 and r = 0.848, respectively). food availability therefore contributed mostly to density and biomass while habitat characteristics did not contribute significantly to small mammal community structure. all species trapped were omnivorous, although the rodents were predominantly granivorous and the macroscelid insectivorous. although no seeds were recorded in the sampling squares, substantial amounts were eaten by g. paeba, a. namaquesis, and petromyscus sp. gerbillurus paeba diet included seeds (60–80 %), green plant material (20–40%) and insects (≤ 20 %); that of aethomys namaquesis ≤ 90 % seeds, ≤ 60 % green plant material and 20 % insects; while petromuscys sp. took ≤ 90 % seeds, ≤ 10% green plant material and ≤ 20 % insects. elephantulus. edwardii took mostly insects (≤ 70%), and less seed and green plant material (≤ 20% and 30%, respectively) (table 4). discussion our study recorded fewer species than others in namaqualand and other arid or semi-arid parts of south africa (dieckman 1979; nel 1983; kerley & erasmus 1992; joubert & ryan 1999), and far fewer than the diverse small mammal fauna of north american deserts (brown & lieberman 1973), but not that of the argentinian monte desert (mares 1976). this could be due to previous disturbance brought about by intensive smallstock grazing before the park was promulgated (avenant 2000), or trapping during a trough in small mammal density resulting from the preceding extensive drought. during 2003 rainfall at the springbok weather station ca. 65 km to the north of the nnp was the lowest since 1984, with 5 mm recorded in autumn and winter (april–july), whereas the long-term average (since 1960) is 102 mm p.a. this resulted in the local extinction of the bush karoo rat, otomys unisulcatus, in the goegap nature reserve some 20 km east of springbok (schradin issn 0075-6458 105 koedoe 49/1 (2006) table 4 contribution (mean % volume) of seeds (white material), foliage (green material) and insects (brown material) to the diet of small mammals in the namaqua national park, south africa species n seeds foliage insects gerbillurus paeba 18 57 22 21 aethomys namaquensis 10 64.6 26.1 9.3 petromyscus 3 90.67 1.67 7.67 elephantulus k edwardii 4 27.25 28.5 44.25 2005). at skilpad in the east of the nnp, average annual rainfall from 1986–2002 was 333 mm, but only 184 mm from jan–sept 2003. long-term trapping in the sw kalahari (nel 1983, nel unpubl.) have shown that species richness and density of small mammals fluctuate in concert with precipitation, a feature perhaps common to deserts (abramsky 1988). such fluctuations could be a secondary response tracking fluctuating primary productivity (rutherford 1980), which would be mirrored by seed availability (reichman 1984; price & reichman 1987) and insect biomass. while seed banks in north american deserts can be large (price & joyner 1997), seed availability, where recorded, is much lower in south african arid and semi-arid regions (milton & dean 1990; kerley 1992b). this could account for the low species richness and low numbers found in our study, as the four rodent species recorded were all primarily granivorous. in addition, seeds of different size tend to be partitioned between differentsized rodent species (brown & lieberman 1973; bowers & brown 1982). seed density in our study grids was too low for seeds to be separated into different-sized samples, but size differences between the coexisting granivorous species were large: m. indutus = 6.1 g, petromyscus sp.= 21 g, g. paeba = 26 g and a. namaquensis = 47 g (i.e. more than the 1.5 x differences in mass suggested to allow coexistence (brown 1973)); while petromyscus also showed some habitat separation from the other rodents. although species richness was low, the capture success of 6.1 % was somewhat higher than that nel.qxd 2006/04/17 11:00 pm page 105 recorded in the karoo (kerley 1992b; kerley & erasmus 1992), where capture successes of 4.8 % and 2.9 %, respectively, were obtained. our higher figure, however, could be an artifact of the prevailing low seed availability which possibly enhanced use of artificially-provided food sources, such as bait in traps. all species captured were omnivorous to a greater or lesser extent, but the rodents were primarily granivorous and the macroscelidid primarily insectivorous, corresponding with results from other studies in arid southern africa (nel 1978; kerley 1992a, 1992b); kerley (1992a) has argued that strictly granivorous rodents are less well represented in arid and semi-arid southern africa than in north american and israeli deserts. in contrast to results of kerley (1992a) from the nama karoo, in our study g. paeba in succulent karoo had a predominantly granivorous diet, as was also found in the sw kalahari (nel 1978). as in other studies (brown & kurzius 1987; kotler & brown 1988; morris 1996), common species (g. paeba and a. namaquensis) were distributed through all or most vegetation types while the other species (m. indutus, petromyscus and e. edwardii) had lower densities and were confined to a particular habitat. in common with results from similar studies in southern africa (nel 1983; kerley & erasmus 1992) which recorded fluctuating species richness and density of small mammals, emigration from refugia rather than dispersal from a source to a sink (johnson et al. 2005) probably account for this phenomenon. microhabitat features such as vegetation structure, cover and height, relatively humidity, litter depth and foliage height diversity are related to the life form and growth form of plant species within a plant community (els & kerley 1996) and affect the small mammal community structure (price 1978; rosenzweig & winakur 1967). similar correlations emerged in our study. small mammal diversity and richness correlated significantly with shrub cover (the dominant plant growth form), with macroscelidid biomass correlating significantly with total basal and rock cover. such correlations suggest that shrub cover in our study area and probably most of the succulent karoo biome is a dominant factor affecting small mammal species diversity, while basal and rock cover determine macroscelidid biomass. therefore selecting shrub cover would presumably reflect both physiological and anti-predator adaptations of small mammals, as well as being a measure of forage availability. along with habitat characteristics correlates, food availability also influences small mammal densities and biomass (kotler & brown 1988). as expected, macroscelidid biomass correlated significantly with insect mass. when the different species were contrasted with environmental features, each showed a significant correlation with some form of environmental feature. this clearly reflects the different physiological, nutritional, social and anti-predator requirements of the small mammal species in question. the linear relationship between species richness and habitat area (allen & white 2003; bruun 2001; nee & cotgreave 2002), as well as species richness and habitat diversity, has been extensively studied (brown & kurzius 1987; cramer & willig 2002; cueto et al. 1995; kerr & packer 1997; kotler & brown 1988; williams et al. 2002). there is some debate about the relationship between habitat diversity and species-area relationship (spar) and their effect on species richness (ricklefs & lovette 1999; mcguinness 2000; van rensburg et al. 2002). in our study the habitats sampled, although varying in size, extended far outside the park so that the species-area relationship cannot be invoked. our study found no correlation between species richness and habitat characteristics. this could be the result of the very low species richness of small mammals recorded due to long-term low primary productivity, especially so in the preceding year (brown & davidson 1977), all species being habitat generalists (morris 1996), low spatial heterogeneity within and between the sample plots in a specific habitat, a lack of habitatspecificity in most species occurring, the koedoe 49/1 (2006) 106 issn 0075-6458 nel.qxd 2006/04/17 11:00 pm page 106 habitats sampled being non-island or virtual island habitats, or a combination of these factors. small mammal densities and biomass correlated with food availability, which supports our hypothesis that density and biomass are dependent on food availability. however, investigations that studied the simultaneous effect of area, habitat diversity and food availability on species characteristics, including small mammals, are limited. longterm studies elsewhere in arid southern africa (nel 1978, 1983) have emphasized that once-off sampling of small mammal populations in arid and semi-arid regions in south africa can bring about misleading results, as numbers of constituent species in a given community fluctuate dramatically in response to rainfall fluctuations. this leads to changes in inter-specific interactions, changes in habitat utilisation and probably also changes in the diet of particular species over time. the results of our survey, undertaken during a long-term drought and at an unknown point in the fluctuating cycle of the community as a whole may therefore not be an accurate reflection of long-term small mammal community composition and dynamics at our study site in the succulent karoo. acknowledgements we thank sanparks for allowing the study and providing facilities and accommodation, and especially andre rossouw for providing accommodation at melkboom, and other members of the namaqua national park for their assistance. we are greatly indebted to g.i.h. kerley and r.j. van aarde for providing detailed critical comments and many helpful suggestions on earlier drafts. references abramsky, z, 1978. small mammal community ecology. changes in species diversity in response to manipulated productivity. oecologia 34: 113–123. abramsky, z. 1988. the role of habitat and productivity in structuring desert rodent communities. oikos 52: 107–114. allen, a. p. & e.p. white. 2003. effects of range size on species-area relationships. evolutionary ecology research 5: 493–499. atkinson, r.p.d. 1997. practical aspects of trapping small mammals in the tropics. journal of zoology 242: 390–394. avenant, n.l. 2000. small mammal community characteristics as indicators of ecological disturbance in the willem pretorius nature reserve, free state, south africa. south african journal of wildlife research 30: 26–33. bar, y., z. abramsky & y. gutterman. 1984. diet of gerbilline rodents in the israeli desert. journal of arid environments 7: 371–376. boonzaaier, m., p. margerman, r. smart. & m. norval. 2002. succulent karoo ecosystem plan. sub-regional report, namaqualand. bowers, m. a. & j.h. brown. 1982. body size and coexistence in desert rodents: chance or community structure. ecology 63: 391–400. brown, j.h. 1973. species diversity of seed-eating desert rodents in sand dune habitats. ecology 54: 775-787. brown, j. h. 1975. geographical ecology of desert rodents. pp. 315-341. in: cody, m. l. & j.m. diamond (eds.). ecology and evolution of communities. new haven, connecticut: belknap press. brown, j.h. & g.a. lieberman. 1973. resource utilization and coexistence of seed-eating rodents in sand dune habitats. ecology 54: 788-797. brown, j.h. & d.w. davidson. 1977. competition between seed-eating rodents and ants in desert ecosystems. science 196: 880-882. brown, j.h., o.j. reichman. & d.w. davidson. 1979. granivory on desert ecosystems. annual review of ecology and systematics 10: 201-207. brown, j.h., d.w. davidson, j.c. munger & r.s. inouye. 1986. experimental community ecology: the desert granivore system. pp. 41-61 in: diamond, j. & t.j. case (eds.). community ecology. new york: harper & row. brown, j. h. & m.a. kurzius. 1987. composition of desert rodent faunas: combinations of coexisting species. annales zoologica fennica 24: 227–237. bruun, h.h. 2001. determinants of species richness in patches of grassland and heathland in himmerland (denmark). nordic journal of botany 21: 607–613. cowling, r.m., k.j. esler & p.w. rundel. 1999. namaqualand, south africa—an overview of a unique winter-rainfall desert ecosystem. plant ecology 142: 3–21. cowling, r. m. & s. pierce. 1999. namaqualand: a succulent desert. cape town: fernwood press. issn 0075-6458 107 koedoe 49/1 (2006) nel.qxd 2006/04/17 11:00 pm page 107 cramer, m. j. & m.r. willig. 2002. habitat heterogeneity, habitat associations, and rodent species diversity in a sand-shinnery-oak landscape. journal of mammalogy 83: 743–753. cueto, v. r., m. cagnoni & m.j. piantanida. 1995. habitat use of scapteromys tumidus (rodentia: cricetidae) in the delta of the paranà river, argentina. mammalia 59: 25–34. dieckmann, r. c. 1979. note on the smaller mammals of the hester malan nature reserve, springbok, namaqualand. south african journal of zoology 14: 85–89. els, l. m. & g.i.h. kerley. 1996. biotic and abiotic correlates of small mammal community structure in the groendal wilderness area, eastern cape, south africa. koedoe 39: 121–130 esler, k. j. & p.w. rundel. 1997. so what is so special about namaqualand? south african journal of science 93:488. hughes, j. j. & d. ward. 1993. predation risk and distance to cover affect foraging behaviour in namib desert gerbils. animal behaviour 46: 1243–1245. hughes, j. j., d. ward & m.r. perrin. 1994. predation risk and competition affect habitat selection and activity of namib desert gerbils. ecology 75: 1397–1405. jensen, s. p. & p. honess. 1995. the influence of moonlight on vegetation height preference and trappability of small mammals. mammalia 59: 35–42. johnson, c.n., k. vernes. & a. payne. 2005. demography in relation to population density in two herbivorous marsupials: testing for source-sink dynamics versus independent regulation of population size. oecologia 143: 70-76. joubert, d. f. & p.g. ryan. 1999. differences in mammal and bird assemblages between commercial and communal rangelands in the succulent karoo, south africa. journal of arid environments 43: 287–299. kam, m., i.s. khokhlova. & a.a. degen. 1997. granivory and plant selection by desert gerbils of different body size. ecology 78: 2218-2229. kent, m. & p. coker. 1992. vegetation description and analysis. london: belhaven press. kerley, g.i.h. 1992a. trophic status of small mammals in the semi-arid karoo, south africa. journal of the zoological society, london 226: 563572. kerley, g.i.h. 1992b. small mammal seed consumption in the karoo, south africa: further evidence for divergence in desert biotic processes. oecologia 89: 471–475. kerley, g.i.h. 1992c. ecological correlates of small mammal community structure in the semi-arid karoo, south africa. journal of the zoological society, london 227: 17-27. kerley, g.i.h. & t. erasmus. 1992. small mammals in the semi-arid karoo, south africa: biomass and energy requirements. journal of arid environments 22: 251-260. kerr, j. t. & l. packer. 1997. habitat heterogeneity as a determinant of mammal species richness in high-energy regions. nature 385: 252–254. kotler, b.p. 1984. risk of predation and the structure of desert rodent communities. ecology 65: 689–701. kotler, b.p. & j.s. brown. 1988. environmental heterogeneity and the coexistence of desert rodents. annual review of ecology and systematics 19: 281–307. krebs, c. j. 1999. ecological methodology. menlo park, california: longman. lancaster, j. 1996. scaling the effects of predation and disturbance in a patchy environment. oecologia 107: 321–331. landry, s.l. 1970. the rodentia as omnivores. quarterly review of biology 45:35–372. low, a. b. & a.g. rebelo. 1996. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. mares, m.a. 1976. convergent evolution of desert rodents: multivariate analyses and zoogeographic implications. paleobiology 2:39–63. mares, m. a. & a.c. hulse. 1977. patterns of some vertebrate communities in creosote bush deserts. pp. 209–226. in: mabry, t. j., j.h. hunziker & d.r. difeo (eds.). creosote bush: biology and chemistry of larrea in new world deserts. stroudsburg, pennsylvania: dowden, hutchinson & ross. martin, t. e. 1998. are microhabitat preferences of coexisting species under selection and adaptive? ecology 79: 656–670. mcguinness, k.a. 2000. distinguishing area and habitat heterogeneity effects: a simulation test of the macnally and watson (1997) protocol. austral ecology 25: 8–15. milton, s. & w.j.r. dean. 1990. seed production in rangelands of the karoo. south african journal of science 86: 231–233. morris, d.w. 1996. coexistence of specialist and generalist rodents via habitat selection. ecology 77: 2352–2364. nee, s. & p. cotgreave. 2002. does the species/ area relationship account for the density/ area relationship. oikos 99: 545–551. nel, j.a.j. 1978. habitat heterogeneity and changes in small mammal community structure and resource utilization in the southern kalahari. bulletin of the carnegie museum of natural history 6: 118–131. nel, j.a.j. 1983. changes in population structure of kalahari rodents over a decade: 1970–1980. koedoe 49/1 (2006) 108 issn 0075-6458 nel.qxd 2006/04/17 11:00 pm page 108 annale musee royal de l’afrique centrale sciences zoologiques 237: 173–178. nel, j.a.j., i.l. rautenbach, d.a. els & g. de graaff. 1985. rodents and other small mammals of the kalahari gemsbok national park. koedoe 27 (supplement): 229–235. price, m.v. 1978. the role of microhabitat in structuring desert rodent communities. ecology 59: 910–921. price, m.v. & s.h. jenkins. 1986. rodents as seed consumers and dispersers. pp. 191–235. in: murray, d.r. (ed.). seed dispersal. sydney: academic press. price, m.v. & o.j. reichman. 1987. spatial and temporal heterogeneity in sonoran desert seed pools, and implications for heteromyid rodent foraging. ecology 68: 1797–1811. price, m. v. & j.w. joyner. 1997. what resources are available to desert granivores: seed rain or soil bank? ecology 78: 764–773. rautenbach, i.l. & j.a.j. nel. 1980. mammal diversity and ecology in the cedarberg wilderness area, cape province. annals of the transvaal museum 32: 101–124. reichman, o.j. 1984. spatial and temporal variation in seed distributions in desert soils. journal of biogeography 11: 1–11. ricklefs, r. e. & i.j. lovette. 1999. the roles of island area per se and habitat diversity in the species-area relationships of four lesser antillean faunal groups. journal of animal ecology 68: 1142–1160. rosenzweig, m. l. & j. winakur. 1969. population ecology of desert rodent communities: habitats and environmental complexity. ecology 50: 558–572. rutherford, m.c. 1980. annual plant productionprecipitation relations in arid and semi-arid regions. south african journal of science 76: 53–56. schradin, c. 2005. nest-site competition in two diurnal rodents from the succulent karoo of south africa. journal of mammalogy 86: 757–762. schulze, b. r. 1947. the climates of south africa according to the classifications of koppen and thornthwaite. south african geographical journal 29: 32–42. skinner, j. d. & r.h.n. smithers. 1990. the mammals of the southern african subregion. pretoria: university of pretoria. sokal, r.r. & f.j. rohlf. 1995. biometry: the principles and practice of statistics in biological research. new york: freeman. theron, g.k., m.w. van rooyen & n. van rooyen. 1993. skilpad wildflower reserve: botanical survey, department of botany, university of pretoria, pretoria. van rensburg, b. j., s.l. chown & k.j. gaston. 2002. species richness, environmental correlates and spatial scale: a test using south african birds. the american naturalist 159: 566–577. van rooyen, m. w., n. grobbelaar, g.k. theron. & n. van rooyen. 1991. the ephemerals of namaqualand: effects of photoperiod, temperature and moisture stress on development and flowering of three species. journal of arid environments 20: 15–29. van rooyen, m. w., n. grobbelaar, g.k. theron & n. van rooyen. 1992. the ephemerals of namaqualand: effect of germination date on development of three species. journal of arid environments 22: 51–66. van rooyen, m. w., g.k. theron & n. grobbelaar. 1990. life form and dispersal spectra of the flora of namaqualand, south africa. journal of arid environments 19: 133–145. williams, s. e., h. marsh & j. winter. 2002. spatial scale, species diversity and habitat structure: small mammals in australian tropical rain forest. ecology 83: 1317–1329. issn 0075-6458 109 koedoe 49/1 (2006) nel.qxd 2006/04/17 11:00 pm page 109 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice article information authors: duan biggs1,2 louise swemmer2 glen phillips3 joep stevens3 stefanie freitag2 rina grant2,4 affiliations: 1centre for biodiversity and conservation science, university of queensland, australia2south african national parks scientific services, kruger national park, skukuza, south africa 3south african national parks tourism managing executive, pretoria, south africa 4department of zoology, nelson mandela metropolitan university, south africa correspondence to: duan biggs postal address: centre of excellence for environmental decisions and the nerp environmental decisions hub, centre for biodiversity and conservation science, university of queensland, st lucia qld 4072, australia dates: received: 19 apr. 2013 accepted: 06 nov. 2013 published: 24 june 2014 republished: 30 july 2014 how to cite this article: biggs, d., swemmer, l., phillips, g., stevens, j., freitag, s. & grant, r., 2014, ‘the development of a tourism research framework by south african national parks to inform management’, koedoe 56(2), art. #1164, 9 pages. http://dx.doi.org/10.4102/ koedoe.v56i2.1164 note: rina grant is a research associate for the department of zoology at nelson mandela metropolitan university, south africa. this article was republished with the added affiliation for rina grant and note. copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. the development of a tourism research framework by south african national parks to inform management in this original research... open access • abstract • introduction • the context of tourism in south african national parks    • overview and history    • strategic adaptive management and tourism research • development of a tourism research framework    • initial stakeholder workshop for tourism research    • furthering the research framework    • identifying knowledge gaps in the implementation of the responsible tourism strategy    • a social-ecological systems perspective       • tourist demand       • relationship with neighbouring communities and broader society       • global environmental change    • conducting an integrated park management planning exercise • operationalising and funding tourism research    • research project facilitation and management    • funding options • discussion    • lessons learnt and challenges    • different cultures of operating, objectives and timeframes    • legal structures and organisational procedures    • entrenched ways of thinking and capacity constraints    • research and the strategic adaptive management of tourism • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ tourism is critical source of financing for conservation in africa. south african national parks (sanparks) raises in excess of 80% of their own funds through tourism revenue. sanparks has a culture of co-learning between scientists and conservation managers through a process known as strategic adaptive management (sam). despite the critical role that tourism plays in sanparks, it has, until recently, not been formally incorporated in the sam process. moreover, sanparks recently adopted a new responsible tourism policy to guide the development and management of tourism across all national parks. the new policy calls for tourism that supports biodiversity conservation, is environmentally efficient and socially responsible. in 2011, sanparks initiated a tourism research programme to support the incorporation of tourism in sam and to provide enabling information for the implementation of the responsible tourism policy. this article summarised the development of the tourism research programme in sanparks and its key research themes. conservation implications: an active tourism research programme that integrates science and management is necessary for tourism to play a stronger role in delivering outcomes for conservation, neighbouring communities and broader society. introduction top ↑ tourism is an important and growing source of financing for protected areas (pas) and conservation globally (dharmaratne, yee sang & walling 2000; naughton-treves, buck holland & brandon 2005; thur 2010). in africa, the financing provided by tourism is particularly important (south african national parks [sanparks] 2012). yet, financing pas and conservation through tourism presents a range of challenges. attempts to maximise tourism revenue may have negative impacts on biodiversity conservation objectives (grant, peel & bezuidenhout 2011). many pas in africa are surrounded by low income communities and challenges, trade-offs and potential synergies exist between optimising for income from visitors and delivering benefits and building and maintaining a healthy relationship with neighbouring communities (barrett, travis & dasgupta 2011; biggs, turpie, fabricius & spenceley 2011; botha, witkofski & cock 2007; naidoo & adamowicz 2005; spenceley 2008). indeed, the relationship between tourism, conservation and broader society forms part of a broader social-ecological system (a joint system of humans and nature) (ban et al. 2013).sanparks raises in excess of 80% of its annual revenue through tourism and is therefore considered one of the more successful pa agencies globally (licht, slotow & millspaugh 2008; sanparks 2012; varghese 2008). sanparks is responsible for managing 19 national parks in south africa, including adjoining marine pas. the sanparks’ estate includes globally renowned parks, such as kruger national park and table mountain national park, and core protected areas in two of the planet’s 25 biodiversity hotspots (myers et al. 2000). of the 19 pas managed by sanparks, only a small number of iconic, internationally known pas, such as kruger national park and table mountain national park, operate at a profit and finance the establishment and management of many other pas in south africa managed by sanparks (sanparks 2012). sanparks is guided by a management philosophy and process known as strategic adaptive management (sam) which has received widespread international exposure (du toit, rogers & biggs 2003; gaylard & ferreira 2011; van wilgen & biggs 2011; venter et al. 2008). this has been supported by the creation of a culture of co-learning between scientific services and conservation managers (biggs, ferreira, freitag-ronaldson & grant-biggs 2011; roux & foxcroft 2011; roux et al. 2006; stirzaker, roux & biggs 2011; venter et al. 2008). until recently, tourism research in sanparks has taken two different forms. firstly, is the research coordinated by the scientific services department, which is often conducted by outside researchers and facilitated and reviewed by sanparks scientific services staff (biggs & rogers 2003; gaylard & ferreira 2011). secondly, is shorter-term research that addresses specific market issues and which is commissioned by the tourism development and marketing executive at the sanparks head office in pretoria (hereafter the tourism executive). research coordinated by scientific services feeds into management through bimonthly conservation meetings, forums between science and management and through scientific inputs on specific management issues. although the conservation meetings have been conducted at regular intervals for over a decade, these typically have not involved the discussion of tourism or tourism research. the development of a tourism research framework was seen to be a way in which tourism research could strengthen how tourism is managed within the framework of sam. thus, in 2011, sanparks initiated the development of a tourism research framework to better integrate tourism research and decision-making with management within the framework of sam. in this article, we describe the development of the tourism research framework in sanparks. firstly, we describe the context in which tourism operates in sanparks; next we describe the process that was followed to develop the tourism research framework. this is followed by a description of the research framework and a discussion of the challenges experienced in implementing the framework, as well as the lessons the sanparks’ experience holds for other conservation agencies with an interest in developing tourism research. the context of tourism in south african national parks top ↑ overview and history sanparks (previously the national parks board) was created in 1926 through an act of parliament as a para-statal organisation (varghese 2008). the core mandate of sanparks is the conservation and management of cultural and natural heritage through a system of national parks. from its inception, sanparks was created in a way in which it was able to generate and use revenue from tourism to fulfil its mandate. tourism in sanparks has grown exponentially since the first three tourist cars entered the kruger national park in 1927. this was followed by 180 cars in 1928 and 850 cars in 1929 (stevenson-hamilton 1937). by the 1950s, over 10 000 people visited the park each year. in the 2011–2012 financial year, national parks received a total of 4.7 million visitors, of which 77% were south african residents. table mountain national park accounted for 2.3 million (49%) guests, mainly day visitors and the kruger national park accounted for 1.4 million (30%). the total revenue from tourism income, including entry fees and retail, amounted to 84% of the total income to sanparks in the 2011–2012 financial year (sanparks 2012). following the political change in south africa in 1994, there has been a strong drive to increase the number of previously excluded or marginalised visitors (non-whites, defined as people of indian, coloured, or black races) to national parks (sanparks 2012). the number of non-white visitors has increased since 1995, but not as much as desired by sanparks. in 2011–2012, 24% of all visitors were non-white. there is a continued strong drive to increase the number of non-white visitors to national parks through developing a more diverse tourism product. in addition, around and after the time of the political change in 1994, there was a realisation of the need to increase and diversify the income stream to national parks (varghese 2008). one strategy that was pursued entailed the development of public–private partnerships (ppps) and concessions in national parks. as of 2012, there are 40 active ppps in national parks that represent 7% of total tourism revenue (sanparks 2012). the ppps are developed and managed with a focus on achieving black economic empowerment (bee) and poverty alleviation objectives as outlined by the national government (sanparks 2012). in 2011, sanparks adopted a responsible tourism policy to guide the development and management of tourism across all national parks (phillips 2012). the policy aligns with new government policies and calls for tourism that supports biodiversity conservation, is environmentally efficient and socially responsible (phillips 2012). the implementation of this policy will be based on satisfying evolving market needs, through service excellence, and the maintenance of high quality standards and infrastructure. sound business principles will be used to generate revenue from tourism initiatives to support the sanparks conservation and constituency-building mandates (phillips 2012). strategic adaptive management and tourism research sanparks adopted sam as the approach of choice to address the complexity inherent in managing its pas (venter et al. 2008). sam consists of three components: adaptive planning (where a vision is defined together with stakeholders and the objectives that have to be met to achieve the vision are developed), adaptive implementation and, lastly, adaptive evaluation (the important step of evaluating and learning from the outcomes of the implementation) (roux & foxcroft 2011). the application of sam aims to contribute to the achievement of the sanparks vision: ‘south african national parks connecting to society’ and mission: ‘to develop, manage and promote a system of national parks that represents the biodiversity and heritage assets by applying best practice, environmental justice, benefit sharing and sustainable use’ (sanparks 2012). tourism has an important role to play in achieving the people’s objective which is, ‘to provide human benefits and build a strong constituency, preserving as far as possible the wilderness qualities and cultural resources associated with national parks’ (swemmer & taljaard 2011). in addition, there is a specific tourism objective under the people’s objective which aims to develop, manage and enhance a range of sustainable tourism products in synergy with the sanparks conservation ethic. the purpose of the tourism research framework is to highlight the required research to support the achievement of conservation, tourism and people’s objectives and to integrate tourism decision-making in the implementation of sam in sanparks. research guided by the tourism research framework can play an important role in enhancing the evaluation of the achievement of sanparks’ objectives, which is necessary as part of the adaptive planning, implementation and evaluation steps of sam. studies that evaluate the importance of iconic mammals, such as lion and elephant, to provide tourists with a high quality, nature-based, value-for-money experience (di minin et al. 2013; lindsey et al. 2007), the socio-economic impact of national parks (saayman & saayman 2010), or the impact of tourism on adjacent communities (anthony 2007; strickland-munro, allison & moore 2010) can provide valuable insights into tourism management when incorporated in the adaptive evaluation process. the intention of the tourism research framework is not only to guide the integration of information for management outcomes but also to be adapted over time as new management needs and objectives emerge. development of a tourism research framework top ↑ the development of the tourism research framework followed a number of iterative steps and processes. these included an initial tourism research workshop, the identification of knowledge gaps associated with sanparks’ responsible tourism policy, the development of a social-ecological systems understanding of tourism to national parks and conducting an integrated park management planning exercise (figure 1). in accordance with the principles of sam, the thinking during the development of the research framework was that it would be evaluated, reviewed and adapted over time. figure 1: process followed for the development of the tourism research framework. initial stakeholder workshop for tourism research the development of the tourism research programme was initiated through engaging a variety of stakeholders, including tertiary research institutions, representatives from district municipalities and private companies that were in ppps with sanparks, in a tourism research workshop held in february 2011.a wide range of sanparks staff also participated. these included members of the tourism executive, policy development, corporate social investment, scientific services, people and conservation and the trans-frontier conservation areas division. the themes covered in the workshop ranged from managing the trade-offs between biodiversity and economics in tourism, neighbouring community benefits from tourism, cultural tourism, aligning tourism product development with the principles of responsible tourism and meeting tourist demands. the working groups discussed the themes in the context of objectives and prioritised management challenges which were then used to identify information and knowledge gaps. this formed the foundation for the drafting of an expansive set of tourism research needs and a draft tourism research framework was established (spenceley 2011). furthering the research framework the further development of the framework took a transdisciplinary approach to research, which recognises that knowledge is co-created between scientists, practitioners and policymakers and that the continual questioning of different disciplinary perspectives is essential (max-neef 2005). in addition, such an approach based on complexity thinking integrates biodiversity conservation, ecological, social and economic approaches. furthermore, the tourism executive expressed the need for tourism decision-making in sanparks to become strategic, structured and evidence-based. from august 2011, extensive discussions took place within scientific services, and with the tourism executive, to further develop a clear agenda for tourism research by advancing the original framework. with this aim, the following three processes were followed: • the identification of knowledge gaps and research needs that stem from the sanparks responsible tourism policy adopted by sanparks (phillips 2012). • the development of a social-ecological systems diagram of tourism in national parks and the questions that this system’s perspective raises. • conducting a tourism management planning exercise for a national park, together with the park and regional management and tourism staff. identifying knowledge gaps in the implementation of the responsible tourism strategy not only is baseline data required to measure sanpark’s progress towards achieving the aims of responsible tourism, but the structures and processes to enable the successful implementation of the policy require understanding (table 1). the key research areas that were identified during this process were: • monitor, evaluate and research the drivers of visitor satisfaction and how it can be improved. • assess the levels of, and research the determinants of successful constituency-building for conservation through tourism and how this can be strengthened. • evaluate and understand the synergies and trade-offs between income from tourism in sanparks and the achievement of conservation and constituency-building objectives and identify opportunities to synergise tourist income with conservation and constituency-building objectives. in addition, the discussion between scientific services and the tourism executive generated a range of additional research needs: • human capacity and human capital is critical for the functioning of the entire tourism and conservation system and research is needed on how it can be strengthened (hall 2009). • the legal and regularity framework governs the way in which tourism products are managed and can be delivered; in other words, management processes, such as procurement procedures and human resource procedures, may affect sanparks’ ability to achieve the objectives of the responsible tourism policy. the interaction between these legal and organisation frameworks and the strategic adaptive management of tourism needs to be understood. • bee is a key national policy imperative in south africa and research is needed to understand how the development and management of tourism in sanparks currently contributes to south africa’s imperative for bee and societal transformation and how this can be improved (sanparks 2012). this research focus transcends the entire organisation, including the structure and management of ppps. sanparks should enable economic empowerment which empowers marginalised and poor communities (ashley 2006). management processes and structures that enable successful transformation but ensure the delivery of top quality tourism services and products need to be researched and developed. an important component of this issue is the research and development of principles and models that can guide partnerships with communities neighbouring parks and other stakeholders so that they can receive more benefits from tourism to national parks. processes and principles to enable the just, transparent and sustainable distribution of such benefits also require research. • the role of national parks in forging a common national identity and a sense of national pride (pretes 2003). table 1: selected principles and associated knowledge gaps and research needs of the south african national parks responsible tourism strategy. a social-ecological systems perspective the role of tourism to pas in conservation and in generating societal benefits plays out in broader interactive social-ecological systems (ban et al. 2013; strickland-munro et al. 2010). an understanding of tourism’s relation to national parks needs to therefore account for these broader social-ecological interactions. the development of social-ecological systems diagrams are widely used as a tool to illuminate the dynamics of a system that require understanding for its management (anderies, janssen & ostrom 2004; wilkinson 2012). a social-ecological systems diagram was developed by scientific services and the tourism executive to establish whether the process of developing the diagram raised additional considerations and research questions that did not emerge in the stakeholder workshop, or in the analysis of the responsible tourism policy research needs. the social-ecological systems diagram that we developed (figure 2) provides an outline of the joint understanding of the tourism system in sanparks by scientific services and the tourism development and marketing executive. below, the key components of the social-ecological systems diagram (figure 2) and the research needs that stemmed from each are discussed. figure 2: a social-ecological systems diagram of tourism to national parks. tourist demand tourist demand and therefore revenue depends on the propensity to travel, which is associated with factors such as, (1) the trendiness of the destination and (2) the value attributed to the travel experience (biggs 2011; hjalager 2010). factors such as concern over carbon emissions from long haul travel may reduce the perceived value of travel to national parks in south africa and therefore reduce demand (tol 2007). affordability is another important factor and depends on issues such as the exchange rate for international visitors and the real cost of visiting national parks (smeral 2010). security concerns can also play a powerful role in reducing demand for travel to national parks (hall, timothy & duval 2004). the tourists’ perceptions of the state of the natural or cultural attractions in parks will depend on their expectations, the interpretation and visitor information provided by sanparks and also their knowledge of a system. for example, research on tourists to coral reefs has shown that visitor perception of coral reef condition can be very different from ecological metrics (andersson 2007; biggs 2011; uyarra, watkinson & côté 2009). different market segments will also differ in their perception – more knowledgeable and experienced tourists are more likely to detect degradation in the environment. the above factors contribute to the expectations of an experience that will result from visiting the destination, which is also an important determinant of tourist demand (fenton, young & johnson 1998). the expectations of what an experience may hold depends on a range of factors that, for the purposes of the sanparks tourism research framework, can be summarised into three groups, (1) experiences of past visits, levels of satisfaction (gnoth 1997), (2) expectations generated by discussions with the social network (wang, yu & fesenmaier 2002) and (3) exposure and interaction with marketing materials and interaction with travel agents (middleton et al. 2009). levels of visitor satisfaction depend on the interaction between these expectations and the visitor’s experience of the pa and its natural and cultural attractions. this experience is mediated by the condition of the tourist infrastructure and facilities and interactions with pa staff. the development of the social-ecological systems understanding led to the identification of additional research priorities to understand the key risks to continued tourism demand and the tourism revenue stream: • perceived security, which partly depends on relationships with neighbouring communities. • affordability of sanparks as a destination relative to competitors. • expectations of experience and levels of visitor satisfaction. • quality of service, infrastructure and facilities. • perceived state of natural and cultural attractions. relationship with neighbouring communities and broader society the relationship between a national park and its neighbouring communities can have an important impact on both the perception of the pa and security concerns (anthony 2007; berkes 2007). engagement by sanparks with neighbouring communities and also broader society will influence the perceptions of these stakeholders and potentially affect their actions and therefore the human impacts that they have on parks (botha et al. 2007). the impacts of neighbouring communities and sectors such as industry and agriculture in broader society include issues such as the illegal harvesting of resources and water extraction from, and pollution of rivers (figure 2). importantly, improving the relationship between pas and neighbouring communities requires the use of a complexity perspective, the ability to deal with multiple objectives through deliberative, participatory processes (berkes 2007). for sanparks to achieve its constituency-building objective, it needs to understand and manage the perception of, and the relationship between, neighbouring communities, broader society and national parks. furthermore, building a constituency for conservation requires the strenghtening the relationship between sanparks, neighbouring communities and broader society. this requires the promotion of mutual benefits with key stakeholders, as well as opportunities for growth and development of neighbouring communities, whilst acknowledging that sanparks is not a development agency (phillips 2012). key research priorities that can support sanparks in achieving these objectives are: • what are the job creation and economic benefits of national parks and, more broadly, of sanparks? • how can the employment and economic contribution of national parks be strengthened? • how can the non-economic benefits of national parks be strengthened, both to visitors and neighbouring communities? • what are the key barriers to enhancing the socio-economic contribution of national parks? • which opportunities exist to involve neighbouring communities and other stakeholders to enhance benefits from tourism to national parks (e.g. the development of cultural tours)? • how can the funds and opportunities from sanparks to neighbouring communities and broader stakeholders be distributed to ensure fairer access to all? global environmental change global environmental change can influence tourism to pas in a number of ways (gossling & hall 2006). climate change receives a lot of attention, but it is the interaction of climate change with pressures, such as invasive species (e.g. see letnic, webb & shine 2008), bush encroachment (e.g. blaum et al. 2007; hudak & wessman 1998) and droughts (e.g. wall & badke 1994), that may be of more importance. since 2000, the kruger national park has been affected by a number of substantial floods which have led to a loss of revenue as a result of a lack of access to tourist and accommodation facilities, but which also have had a significant financial burden in the replacement and repair of damaged infrastructure (sanparks 2012). key research priorities identified with respect to environmental change are:• what are the key potential risks from environmental change on visitor satisfaction and tourist demand as a result of impacts such as bush encroachment, more frequent and severe flooding and/or droughts? • what are the repair and maintenance cost and revenue implications of increased frequency and severity of natural disasters such as floods? • what are the potential risks that environmental change poses to the livelihoods of neighbouring communities (e.g. from more frequent or extreme floods or droughts) and how may these impact on constituency-building efforts and tourism? conducting an integrated park management planning exercise in addition to the theoretical development and discussions between scientific services and the tourism executive to strengthen the transdisciplinary nature of the tourism research framework, the revision of an integrated park management plan was part of the process. the management plan for augrabies falls national park (hereafter augrabies) was due for revision and this was used as an opportunity for a two-day workshop. representatives of scientific services and the tourism executive participated in a two-day management planning exercise together with conservation management, people and conservation, and tourism management and marketing staff from sanparks based at augrabies and in the broader region. the objective of the workshop was to integrate tourism planning with the biodiversity conservation plan and the plan for engaging and working with local communities. the process highlighted the value of a broader sanparks tourism research agenda which can inform (and be informed by) tourism planning at the park level. every national park will have their own specific research needs and therefore the tourism research framework should be implemented in consideration of the different contexts of each national park. however, a few of the tourism research needs that emerged in augrabies are likely to be relevant to many national parks. for example, a key issue in augrabies is the need to understand the market for new products to increase visitor stay and visitor income. in addition, there are numerous local communities in close proximity to augrabies that can potentially partner with sanparks to deliver cultural tours and products to visitors. this would simultaneously deliver benefits to local communities and increase the diversity of products offered to tourists. the market demand for and the potential partnership models through which such products can be managed needs to be explored. as the tourism management planning process is conducted for other national parks, other park-specific needs may emerge. operationalising and funding tourism research top ↑ research project facilitation and management sanparks has a strong track record over the past decade and a half of coordinating and managing research with scientific partners at universities and research institutions in south africa and internationally (du toit et al. 2003; gaylard & ferreira 2011). to date, much of this research has been focused on ecological or biological issues. the aim of the tourism research programme is to foster and conduct high quality research of international standard through partnerships with research institutions to support the sam of tourism in sanparks. tourism research projects will be managed through the same research project management process that is in place for all research projects. individuals from universities and research agencies that are interested in collaborating with sanparks in conducting tourism research submit a research proposal. the sanparks staff member responsible for coordinating tourism research will work together with the respective individuals to ensure that the research proposal is aligned with sanparks objectives and information needs. sometimes, commercially oriented research with tight deadlines is required to inform urgent management decisions. such rapid assessments will follow an alternative faster review system that still aims to ensure the rigour and the value of the research. funding options through partnerships with scientific institutions, the co-funding of tourism research is possible. related funding options include joint applications for industry and research linkage grants (with south african and international institutions) and joint funding of graduate student or research fellowships. importantly, tourism research expertise and capacity is required within sanparks to coordinate, facilitate and integrate research to ensure that it aligns with sanpark’s information needs and can effectively support the practice of sam within the tourism domain. discussion top ↑ lessons learnt and challenges our article has described one of the first attempts, to our knowledge, of integrating tourism research into the adaptive management policies of a pa agency. the integration of tourism research into the sam of tourism in sanparks remains a challenge. despite the development of a tourism research framework jointly between scientific services and the tourism executive, there is still little integration between tourism management and science and limited progress towards managing tourism within the framework of sam. through the process of developing the tourism research framework and its implementation, we faced a number of challenges and learnt numerous lessons that may be of value to other pa agencies and conservation organisations that attempt to integrate tourism research and decision-making and conservation management in a transdisciplinary fashion. the key issues that we identified are, (1) different objectives and different timeframes for tourism and conservation objectives, (2) auditing and organisational reporting systems that are not congruent with an adaptive approach to management and (3) human and organisational capacity constraints and entrenched ways of thinking about managing tourism. we discuss each of these in turn. different cultures of operating, objectives and timeframes tourism managers have to report on short-term financial performance as part of the evaluation of whether annual financial objectives have been met. their requirements can lead to a very different way of thinking about the type of tourism research that is needed. in essence, there is a tension between a more business-oriented operating culture focused on short-term financial objectives and a conservation management and science culture focused on longer-term, more holistic objectives such as biodiversity conservation and achieving community benefits. these objectives are not necessarily mutually exclusive and they can be aligned (e.g. see biggs, ban & hall 2012). however, this alignment requires regular interaction between scientific services, tourism managers and national park managers. the implementation of the tourism research framework through these regular interactions represents an attempt to implement a novel transdisciplinary, science-based approach to management – a challenging task (gunderson, holling & light 1995). closer cooperation between scientific services, the tourism executive and management both within parks and at national level will be essential to implement sam and the findings of tourism research effectively and through regular communication and collaboration this cooperation can be achieved. legal structures and organisational procedures legal structures and auditing procedures in organisations originate from a perspective of command and control and not one of adapting and learning in a complex, adaptive system (ebbesson 2010). although the theory and science of legal practices and auditing is adapting to a more complex world, these changes will take time to filter through to individual pa agencies and conservation organisations. research and experimentation is required to find ways of managing tourism and conservation more adaptively within existing auditing and legal procedures, as well as to develop new processes that are more adaptive but still satisfy the need for control and regulations of auditors and regulators (ebbesson 2010). entrenched ways of thinking and capacity constraints the cognitive frameworks that people use to interpret and the world and make decisions play a critical role in managing the complex interface between tourism and conservation (biggs, abel, knight, leitch, langston & ban 2011). entrenched perspectives about what tourists want, how to increase the number of tourists to a pa and how tourism should be managed act as potential constraints to managing the tourism and conservation interface more creatively, with more room for the emergence of synergies between tourism and conservation. however, there are numerous participatory social processes (e.g. barnaud, bousquet & trebuil 2008; biggs, abel, knight, leitch, langston & ban 2011; cundill et al. 2012; voinov & bousquet 2010) which can be used to address this challenge and develop a shared vision for action for tourism and conservation based on the co-construction of a shared understanding of a system and the alternative options available. research and the strategic adaptive management of tourism the aim of the development of the tourism research framework was to strengthen the sam of tourism in national parks and to integrate research coordinated and guided by scientific services into the decisions and policies of the tourism executive and individual park managers. the benefits of this integration would be a more holistic consideration of the interactions between biodiversity conservation and tourism in policy and management decisions. however, because of the challenges described above, the collaboration between scientific services and the tourism executive to achieve this integration has not been sustained. these challenges were not completely unexpected, but perhaps insufficient attention was paid to how these challenges could be managed earlier on in the dialogue between scientific services and the tourism executive. implementing a novel transdisciplinary, science-based approach to the adaptive management of tourism through sam was always going to be challenging (gunderson et al. 1995). a commitment from both scientific services and the tourism executive to regular communication, collaboration and the commitment of sufficient resources and human capacity for this collaboration will be necessary. moreover, both departments need to acknowledge and respect that they operate according to different cultural norms and perspectives with sometimes differing objectives (biggs, abel, knight, leitch, langston & ban 2011; cundill et al. 2012). through such collaborative processes, co-learning between the departments can emerge which together with closer cooperation between scientific services and the tourism executive and management will be essential to integrate the findings of tourism research effectively within a sam framework in sanparks. a further important lesson learnt is that the implementation of tourism research in management decision-making could possibly have been strengthened by collaborating more closely with the managers in individual national parks as well as the tourism executive. many decisions or components are taken at the level of individual national parks and developing the research framework in a more participatory and decentralised way would have enabled the implementation of the research framework to take place at the level of individual national parks as well. the benefits of decentralisation for conservation and natural resource management are widely discussed in the literature (andersson & ostrom 2008; bohensky 2008). conclusion top ↑ in this article, we discussed the development of a transdisciplinary tourism research framework by sanparks. the development of the framework was led by scientific services and included extensive interactions with managers at an individual national park, with the tourism executive and other stakeholders. the research suggested by this framework is slowly getting underway and sanparks is adapting the framework in accordance with the lessons learnt. we believe that our experiences in integrating tourism research with conservation management can hold valuable lessons for other conservation agencies. acknowledgements top ↑ the authors would like to thank the kruger park marathon club for providing financial support to the development of a tourism research framework. in addition, we would like to thank harry biggs, giju varghese, marna herbst and danie pienaar who provided their time and insight during the development of the tourism research framework and this article. in addition, we would like to thank two reviewers who provided us with comments to substantially improve this article. stefan gelcich at the pontifica universidad catolica de chile provided office space and resources during the revision stage of this article. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions d.b. (university of queensland) was the project leader and led the writing of the manuscript. g.p. (sanparks tourism managing executive) and j.s. (sanparks tourism managing executive) participated in workshop discussions and made conceptual contributions. l.s. (sanparks scientific services), s.f. (sanparks scientific services) and r.g. (sanparks scientific services) made conceptual contributions and assisted with writing and editing. references top ↑ anderies, j.m., janssen, m.a. & ostrom, e., 2004, ‘a framework to analyze the robustness of social-ecological systems from an institutional perspective’, ecology and society 9, 18.andersson, j.e.c., 2007, ‘the recreational cost of coral bleaching – a stated and revealed preference study of international tourists’, ecological economics 62(3–4), 704–715. http://dx.doi.org/10.1016/j.ecolecon.2006.09.001 andersson, k. & ostrom, e., 2008, ‘analyzing decentralized resource regimes from a polycentric perspective’, policy sciences 41, 71–93. http://dx.doi.org/10.1007/s11077-007-9055-6 anthony, b., 2007, ‘the dual nature of parks: attitudes of neighbouring communities towards kruger national park, south africa’, environmental conservation 34(3), 236–245. http://dx.doi.org/10.1017/s0376892907004018 ashley, c., 2006, facilitating pro-poor tourism with the private sector. lessons learned from pro-poor tourism pilots in southern africa, overseas development institute, london. ban, n.c., mills, m., tam, j., hicks, c.c., klain, s., stoeckl, n. et al., 2013, ‘a social-ecological approach to conservation planning: embedding social considerations’, frontiers in ecology and the environment 11(4), 194–202. barnaud, c., bousquet, f. & trebuil, g., 2008, ‘multi-agent simulations to explore rules for rural credit in a highland farming community of northern thailand’, ecological economics 66(4), 615–627. http://dx.doi.org/10.1016/j.ecolecon.2007.10.022 barrett, c.b., travis, a.j. & dasgupta, p., 2011 ‘on biodiversity conservation and poverty traps’, proceedings of the national academy of sciences 108(34), 13907–13912. http://dx.doi.org/10.1073/pnas.1011521108, pmid:21873176, pmcid:pmc3161563 berkes, f., 2007, ‘community-based conservation in a globalized world’, proceedings of the national academy of sciences 104(39), 15188–15193. http://dx.doi.org/10.1073/pnas.0702098104, pmid:17881580, pmcid:pmc2000555 biggs, d., 2011, ‘understanding resilience in a vulnerable industry – the case of reef tourism on australia’s great barrier reef’, ecology & society 16(1), 30. biggs, h.c. & rogers, k., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59–80, island press, washington, dc. biggs, h., ferreira, s., freitag-ronaldson, s. & grant-biggs, r., 2011, ‘taking stock after a decade: does the “thresholds of potential concern” concept need a socio-ecological revamp?’, koedoe 53(2), 9 pages. http://dx.doi.org/10.4102/koedoe.v53i2.1002 biggs, d., turpie, j., fabricius, c. & spenceley, a., 2011, ‘the value of avitourism for community-based conservation – an analysis from south africa’, conservation and society 9(1), 80–90. http://dx.doi.org/10.4103/0972-4923.79198 biggs, d., abel, n., knight, a.t., leitch, a., langston, a. & ban, n.c., 2011, ‘the implementation crisis in conservation planning could “mental models” help?’, conservation letters 4, 169–183. http://dx.doi.org/10.1111/j.1755-263x.2011.00170.x biggs, d., ban, n.c. & hall, c.m., 2012, ‘lifestyle values, resilience, and nature-based tourism’s contribution to conservation on australia’s great barrier reef’, environmental conservation 39(4), 370–379. http://dx.doi.org/10.1017/s0376892912000239 blaum, n., rossmanith, e., popp, a. & jeltsch, f., 2007, ’shrub encroachment affects mammalian carnivore abundance and species richness in semiarid rangelands’, acta oecologica 31(1), 86–92. http://dx.doi.org/10.1016/j.actao.2006.10.004 bohensky, e.l., 2008, ‘discovering resilient pathways for south african water management: two frameworks for a vision’, ecology and society 13, artcile 19. botha, j., witkofski, e.t.f. & cock, j., 2007, ‘is “something better than nothing”? the impacts of ten outreach nurseries on south african community stakeholders’, forests, trees and livelihoods 17, 199–221. http://dx.doi.org/10.1080/14728028.2007.9752597 cundill, g., cumming, g.s., biggs, d. & fabricius, c., 2012, ‘soft systems thinking and social learning for adaptive management’, conservation biology 26(1), 13–20. http://dx.doi.org/10.1111/j.1523-1739.2011.01755.x, pmid:22010884 dharmaratne, g.s., yee sang, f. & walling, l.j., 2000, ‘tourism potentials for financing protected areas’, annals of tourism research 27(3), 590–610. http://dx.doi.org/10.1016/s0160-7383(99)00109-7 di minin, e., fraser, i., slotow, r. & macmillan, d.c., 2013, ‘understanding heterogeneous preference of tourists for big game species: implications for conservation and management’, animal conservation 16, 249–258. http://dx.doi.org/10.1111/j.1469-1795.2012.00595.x du toit, j.t., rogers, k.h. & biggs, h.c., 2003, the kruger experience: ecology and management of savanna heterogeneity, island press, washington, dc. ebbesson, j., 2010, ‘the rule of law in governance of complex socio-ecological changes’, global environmental change 20, 414–422. http://dx.doi.org/10.1016/j.gloenvcha.2009.10.009 fenton, d.m., young, m. & johnson, v.y., 1998, ‘re-presenting the great barrier reef to tourists: implications for tourist experience and evaluation of coral reef environments’, leisure sciences: an interdisciplinary journal 20(3), 177–192. gaylard, a. & ferreira, s., 2011, ‘advances and challenges in the implementation of strategic adaptive management beyond the kruger national park – making linkages between science and biodiversity management’, koedoe 53(2), 8 pages. http://dx.doi.org/10.4102/koedoe.v53i2.1005 gnoth, j., 1997, ‘tourism motivation and expectation formation’, annals of tourism research 24(2), 283–304. http://dx.doi.org/10.1016/s0160-7383(97)80002-3 gossling, s. & hall, c.m., 2006, tourism and global environmental change, routledge, abingdon. grant, r.c.c., peel, m.j.s. & bezuidenhout, h., 2011, ‘evaluating herbivore management outcomes and associated vegetation impacts’, koedoe 53(2), 15 pages. gunderson, l.h., holling, c. & light, s.s., 1995, barriers and bridges to the renewal of ecosystems and institutions, columbia university press, new york. hall, c.m., 2009, ‘tourism firm innovation and sustainability’, in s. gössling, c.m. hall & d. weaver (eds.), sustainable tourism futures: perspectives on systems, restructuring and innovations, pp. 282–298, routledge, new york. pmcid:pmc2677279 hall, c.m., timothy, d. & duval, d., 2004, safety and security in tourism: relationships, management and marketing, haworth press, new york. hjalager, a.-m., 2010, ‘a review of innovation research in tourism’, tourism management 31(1), 1–12. http://dx.doi.org/10.1016/j.tourman.2009.08.012 hudak, a.t. & wessman, c.a., 1998, ‘textural analysis of historical aerial photography to characterize woody plant encroachment in south african savanna’, remote sensing of environment 66(3), 317–330. http://dx.doi.org/10.1016/s0034-4257(98)00078-9 letnic, m., webb, j.k. & shine, r., 2008, ‘invasive cane toads (bufo marinus) cause mass mortality of freshwater crocodiles (crocodylus johnstoni) in tropical australia’, biological conservation 141(7), 1773–1782. http://dx.doi.org/10.1016/j.biocon.2008.04.031 licht, d.s., slotow, r. & millspaugh, j., 2008, ‘out of africa: lessons from park management in south africa’, the george wright forum 25, 20–29. lindsey, p.a., alexander, r., mills, m.g.l., romanach, s. & woodroffe, r., 2007, ‘wildlife viewing preferences of visitors to protected areas in south africa: implications for the role of ecotourism in conservation’, journal of ecotourism 6, 19–33. http://dx.doi.org/10.2167/joe133.0 max-neef, m.a., 2005, ‘foundations of transdisciplinarity’, ecological economics 53(1), 5–16. http://dx.doi.org/10.1016/j.ecolecon.2005.01.014 middleton, v.t.c., fyall, a., morgan, m. & ranchod, a., 2009, marketing in travel and tourism, eslevier, oxford. myers, n., mittermeier, r.a., mittermeier, c.g., da fonseca, g.a.b. & kent, j., 2000, ‘biodiversity hotspots for conservation priorities’, nature 403, 853–858. http://dx.doi.org/10.1038/35002501, pmid:10706275 naidoo, r. & adamowicz, w.l., 2005, ’biodiversity and nature-based tourism at forest reserves in uganda’, environment and development economics 10, 159–178. http://dx.doi.org/10.1017/s1355770x0400186x naughton-treves, l., buck holland, m. & brandon, k., 2005, ‘the role of protected areas in conserving biodiversity and sustaining local livelihoods’, annual review of environmental resources 30, 219–252. http://dx.doi.org/10.1146/annurev.energy.30.050504.164507 phillips, g., 2012, responsible tourism in sanparks: 2012–2022 strategy, sanparks, pretoria. pretes, m., 2003, ’tourism and nationalism’, annals of tourism research 30(1), 125–142. http://dx.doi.org/10.1016/s0160-7383(02)00035-x south african national parks, 2012, south african national parks annual report 2010–2011, sanparks, pretoria. saayman, m. & saayman, a., 2010, ‘regional development and national parks in south africa: lessons learned’, tourism economics 16, 1037–1064. http://dx.doi.org/10.5367/te.2010.0007 roux, d.j. & foxcroft, l.c., 2011, ‘the development and application of adaptive management in south african national parks’, koedoe 53(2), 5 pages. http://dx.doi.org/10.4102/koedoe.v53i2.1049 roux, d.j., rogers, k.h., biggs, h.c., ashton, p.j. & sergeant, a., 2006, ‘bridging the science-management divide: moving from unidirectional knowledge transfer to knowledge interfacing and sharing’, ecology and society 11(1), article 4. smeral, e., 2010, ‘impacts of the world recession and economic crisis on tourism: forecasts and potential risks’, journal of travel research 49(1), 31–38. http://dx.doi.org/10.1177/0047287509353192 spenceley, a., 2008, responsible tourism: critical issues for conservation and development, earthscan, london. spenceley, a., 2011, ‘a tourism research workshop for south african national parks, 01–02 february 2011’, south african national parks, pretoria. stevenson-hamilton, j., 1937, south african eden, penguin, johannesburg. stirzaker, r.j., roux, d. & biggs, h.c., 2011, ‘learning to bridge the gap between adaptive management and organisational culture’, koedoe 53(2), 6 pages. http://dx.doi.org/10.4102/koedoe.v53i2.1007 swemmer, l.k. & taljaard, s., 2011, ‘sanparks, people and adaptive management: understanding a diverse field of practice during changing times’, koedoe 53(2), 7 pages. http://dx.doi.org/10.4102/koedoe.v53i2.1017 strickland-munro, j.k., allison, h.e. & moore, s.a., 2010, ‘using resilience concepts to investigate the impacts of protected area tourism on communities’, annals of tourism research 37, 499–519. http://dx.doi.org/10.1016/j.annals.2009.11.001 thur, s.m., 2010, ‘user fees as sustainable financing mechanisms for marine protected areas: an application to the bonaire national marine park’, marine policy 34(1), 63–69. http://dx.doi.org/10.1016/j.marpol.2009.04.008 tol, r.s.j., 2007, ‘the impact of a carbon tax on international tourism’, transportation research part d: transport and environment 12(2), 129–142. http://dx.doi.org/10.1016/j.trd.2007.01.004 uyarra, m., watkinson, a. & côté, i., 2009, ‘managing dive tourism for the sustainable use of coral reefs: validating diver perceptions of attractive site features’, environmental management 43(1), 1–16. http://dx.doi.org/10.1007/s00267-008-9198-z, pmid:18810525 van wilgen, b.w. & biggs, h.c., 2011, ’a critical assessment of adaptive ecosystem management in a large savanna protected area in south africa’, biological conservation 144, 1179–1187. http://dx.doi.org/10.1016/j.biocon.2010.05.006 varghese, g., 2008, ‘public–private partnerships in south african national parks: the rationale, benefits, and lessons learnt’, in a. spenceley (ed.), responsible tourism: critical issues for conservation and development, pp. 69–84, earthscan, london. venter, f.j., naiman, r.j., biggs, h.c. & pienaar, d.j., 2008, ‘the evolution of conservation management philosophy: science, environmental change and social adjustments in kruger national park’, ecosystems 11(2), 173–192. http://dx.doi.org/10.1007/s10021-007-9116-x voinov, a. & bousquet, f., 2010, ‘modelling with stakeholders’, environmental modelling and software 25(11), 1268–1281. http://dx.doi.org/10.1016/j.envsoft.2010.03.007 wall, g. & badke, c., 1994, ‘tourism and climate change: an international perspective’, journal of sustainable tourism 2(4), 193–203. http://dx.doi.org/10.1080/0966958 9409510696 wang, y., yu, q. & fesenmaier, d.r., 2002, ‘defining the virtual tourist community: implications for tourism marketing’, tourism management 23(4), 407–417. http://dx.doi.org/10.1016/s0261-5177(01)00093-0 wilkinson, c., 2012, ‘social-ecological resilience: insights and issues for planning theory’, planning theory 11(2), 148–169. http://dx.doi.org/10.1177/1473095211426274 abstract introduction study area methods results discussion conclusion acknowledgements references about the author(s) francesco martini department of environmental biology, sapienza university of rome, italy robert cunliffe department of environmental biology, sapienza university of rome, italy alessio farcomeni department of public health and infectious diseases, sapienza university of rome, italy michele de sanctis department of environmental biology, sapienza university of rome, italy giacomo d’ammando school of animal, plant and environmental sciences, university of the witwatersrand, south africa fabio attorre department of environmental biology, sapienza university of rome, italy citation martini, f., cunliffe, r., farcomeni, a., de sanctis, m., d’ammando, g. & attorre, f., 2016, ‘classification and mapping of the woody vegetation of gonarezhou national park, zimbabwe’, koedoe 58(1), a1388. http://dx.doi.org/10.4102/koedoe.v58i1.1388 note: additional supporting information may be found in the online version of this article as online appendix 1: http://dx.doi.org/10.4102/koedoe.v58i1.1388-1 and online appendix 2: http://dx.doi.org/10.4102/koedoe.v58i1.1388-2 original research classification and mapping of the woody vegetation of gonarezhou national park, zimbabwe francesco martini, robert cunliffe, alessio farcomeni, michele de sanctis, giacomo d’ammando, fabio attorre received: 07 mar. 2016; accepted: 06 july 2016; published: 29 sept. 2016 copyright: © 2016. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract within the framework of the great limpopo transfrontier conservation area (gltfca), the purpose of this study was to produce a classification of the woody vegetation of the gonarezhou national park, zimbabwe, and a map of its potential distribution. cover-abundance data of woody species were collected in 330 georeferenced relevés across the park. these data were used to produce two matrices: the first one using the cover-abundance values as collected in five height layers and the second one based on merging the layers into a single cover value for each species. automatic classifications were produced for both matrices to determine the optimal number of vegetation types. the two classification approaches both produced 14 types belonging to three macro-groups: mopane, miombo and alluvial woodlands. the results of the two classifications were compared looking at the constant, dominant and diagnostic species of each type. the classification based on separate layers was considered more effective and retained. a high-resolution map of the potential distribution of vegetation types for the whole study area was produced using random forest. in the model, the relationship between bioclimatic and topographic variables, known to be correlated to vegetation types, and the classified relevés was used. identified vegetation types were compared with those of other national parks within the gltfca, and an evaluation of the main threats and pressures was conducted. conservation implications: vegetation classification and mapping are useful tools for multiple purposes including: surveying and monitoring plant and animal populations, communities and their habitats, and development of management and conservation strategies. filling the knowledge gap for the gonarezhou national park provides a basis for standardised and homogeneous vegetation classification and mapping for the entire great limpopo transfrontier conservation area. introduction gonarezhou national park (gnp) was established in 1975, in the southeast of zimbabwe alongside the border with mozambique, stretching between the mwenezi and save rivers, 21°00’–22°15’ s and 30°15’–32°30’ e (figure 1). it comprises a roughly rectangular strip of land some 35 km – 45 km in width and some 135 km long. together with the adjacent malapati safari area, the total size of the study area is about 5000 km2. in conjunction with the kruger national park (knp) of south africa and the limpopo (lnp), banhine (bnp) and zinave (znp) national parks of mozambique, the gnp forms part of the great limpopo transfrontier conservation area (gltfca). to support the establishment of the gltfca, a standardised vegetation survey of the three parks in mozambique has been carried out (stalmans, gertenbach & carvalho-serfontein 2004; stalmans & peel 2010; stalmans & wishart 2005), while comprehensive studies of vegetation in the knp go back to the 1980s of the last century (gertenbach 1983; venter & gertenbach 1986). a number of vegetation studies have been carried out for the gnp (gandiwa et al. 2011a, 2011b, 2012; o’connor & campbell 1986; tafangenyasha 1997, 2001; wild 1955; zisadza-gandiwa et al. 2013), but these focused only on certain areas or species. only one study produced a comprehensive vegetation map with a description of 15 vegetation types (sherry 1970), but was largely based on work carried out in 1959–1960 in connection with tsetse fly control operations (farrell 1968). the aim of this paper is to provide an updated and comprehensive woody vegetation survey of the gnp, to analyse the relationships between vegetation types and environmental factors and to produce a map of potential natural vegetation. an evaluation of the main threats to the vegetation types was also conducted. such a study is a fundamental requirement in order to develop appropriate conservation and management measures and to monitor landscape changes in the future within the framework of the gltfca. figure 1: location of gonarezhou national park within the great limpopo transfrontier conservation area. study area the climate of the gnp can be regarded as semi-arid (gandiwa et al. 2011b). mean annual rainfall was 499 mm over the period from 1970 to 2009 (gandiwa et al. 2016). the bulk of the rain falls between november and march, with may – september being the driest months. annual rainfall is highly variable. droughts are relatively common with occasional severe occurrences, while parts of the park are prone to flooding, particularly around the save-runde junction. temperatures are relatively high, with daily temperatures exceeding 30 °c occurring in all months of the year. elevation ranges from 160 m a.s.l. at the save-runde junction to a maximum of 578 m a.s.l. on the chiwonja hills. morphologically, the gnp forms part of the limpopo-save lowlands of zimbabwe. these extend across the southernmost part of the country between the shashe and the save rivers, in the form of a relatively flat plain that rises gently to the north from the limpopo river, and form part of the pliocene erosion cycle (lister 1987). within the park, there are minor localised exposures of the earlier post-african surface on the highest parts of the chiwonja hills, and the more recent quaternary surface along the major wide sand-bed drainages of the lower reaches of the mwenezi, runde and save rivers. geologically, the gnp forms part of a down-faulted basin that has subsequently been filled by various igneous intrusions and sedimentary deposits. the central and major part of the park is covered by cretaceous sedimentary deposits, known locally as the malvernia beds, comprising a succession of gently dipping red and white sandstones, grits and conglomerates, variably cemented by calcite. portions of the park to both the north and south are underlain by deep basalt beds, intruded in places by more recent intermediate acid rocks. small areas of more recent quaternary alluvial deposits are found in association with the major drainages. given the relatively young geology and prevailing semi-arid climatic conditions, the soils tend to be relatively shallow and unleached and with appreciable reserves of weatherable materials such that they are inherently fertile (nyamapfene 1991; thompson & purves 1978). methods vegetation data set we classified the vegetation of gnp through the analysis of woody species composition. woody plants represent one of the most distinguishing features of the vegetation in southern africa (germishuizen & meyer 2003; o’brien, field & whittaker 2000), and have been widely used for modelling vegetation structure and physiognomy in the region and elsewhere (daru et al. 2015; kubota et al. 2014; o’brien 1993). a total of 330 relevés were conducted between march and july 2010 (figure 2), following the methodology of timberlake, nobanda and mapaure (1993). sampling was targeted to distinct vegetation communities that were clearly recognisable on satellite imagery (landsat and google earth) and/or in the field. a plotless method was used for each relevé, whereby all woody species present within an area of 0.5 ha – 2 ha were recorded and classified in five height classes: seedlings, less than 0.5 m (mainly regeneration), 0.5 m – 3 m (shrubs and young trees), trees taller than 3 m (tall shrubs and trees); and mature trees (canopy height trees). each species was allocated to a cover-abundance value in each layer using a six-point modified braun–blanquet scale: + = < 2%, 1 = 2% – 10%, 2 = 11% – 25%, 3 = 26% – 50%, 4 = 51% – 75% and 5 = 76% – 100%. any species that could not be reliably identified in the field was collected and later compared with the collection available at the national herbarium in harare. figure 2: location of the 330 vegetation relevés. environmental parameters considered relevant to the interpretation of plant distribution were also recorded. these included landscape type, lithology, soil type, slope, aspect, presence of termitaria, proportion of bare ground and litter cover, presence of surface capping, gulley and sheet erosion. a database was created in turboveg (hennekens & schaminee 2001). the cover values in different layers for each species were subsequently combined, in order to obtain a single value of cover per species per relevé. calculations were performed under the assumption of random species distribution within a plot. environmental data set a digital elevation model (dem) with a spatial resolution of 90 m, representing the geographic area of interest, was downloaded from the data set of the cgiar consortium for spatial information (www.cgiar-csi.org). the dem cell size was re-scaled to a resolution of 100 m. eighteen bioclimatic variables were downloaded from worldclim (www.worldclim.org) at a spatial resolution of 1 km, and subsequently downscaled to a resolution of 100 m according to the methodology of zimmermann et al. (2007). a linear regression presenting altitude as the response variable and the bioclimatic variables as explanatory covariates was performed in statistica (version 8, statsoft inc., tulsa), after testing for the normality of the covariates using the kolmogorov–smirnov test. in case normality was rejected, an appropriate box–cox transformation was used. the intercept and angular coefficient values of each bioclimatic variable were then used in arcgis (version 10.1, esri inc., redlands, california) to produce bioclimatic raster maps at a spatial resolution of 100 m. four additional topographic variables were calculated: slope, curvature, aspect and incoming solar radiation. data analyses two classifications were carried out: a traditional one based on the percentage cover for each species summed across all layers and one based on the individual cover values of each species in each layer, which can be considered similar to the integrated synusial approach developed in switzerland and france (gillet & gallandat 1996), where the classification scheme is built based on the classifications of separate vegetation layers. in order to identify the optimal number of vegetation types for the 330 relevés, a dissimilarity matrix was created using the kulczynsky distance method (faith, minchin & belbin 1987) to calculate dissimilarities between them. for each matrix, classifications based on 2, 3, 4, and continuing up to 15 types were calculated. the clustering technique used was partitioning around medoids. for each of these classifications, the gap statistic was obtained, and the optimal number of types was set as the minimum number of types no lower than one standard error of the maximum gap statistic observed (tibshirani, walther & hastie 2001). the modified rand index was used in order to quantify the similarity between the two selected classifications. all statistical analyses were performed in r (www.r-project.org). for the two classifications, for each type, diagnostic, constant and dominant species were identified in juice (tichy 2002) according to the following selection criteria: constant species: frequency threshold of 50%. dominant species: cover threshold of 25%. diagnostic species: fidelity threshold of 15 (phi coefficient; chytrý et al. 2002). for the identification of diagnostic species, in order to remove dependence from the size of the vegetation types, a standardisation was first performed. the fisher’s exact test was calculated, and a zero fidelity value was given to species with significance p > 0.001. after selection of the best classification, and after verifying the normality of the environmental data, a supervised classification was performed using the observed data (i.e. the classified relevés) as a training set and the bioclimatic and topographic variables as covariates in order to produce a map of the potential natural vegetation. this procedure has its theoretical background in predictive vegetation modelling, which has been defined as ‘predicting the distribution of vegetation across the landscape based on the relationship between the spatial distribution of vegetation and environmental variables’ (franklin 1995). based on this assumption, several methods have been used to model the spatial distribution of vegetation types (attorre et al. 2014; brzeziecki, kienast & wildi 1993; maggini et al. 2006). in our study, random forests (rf) (breiman 2001) was used as the classifying method, and the probability of each background cell being assigned to one of the types was estimated using their climatic and topographic data. rf was chosen because of its widely recognised efficacy in such vegetation studies (attorre et al. 2011; bradter et al. 2011; mellor et al. 2013; rodriguez-valiano et al. 2012). the optimal type for each cell was decided through a maximum a-posteriori criterion. to confirm the validity of the spatialisation procedure, we used two methods: cramer’s v index was calculated by comparing the percentage of relevés for each vegetation type expressed as a percentage of the total number of relevés with the percentage of background cells attributed to each vegetation type. cramer’s v index is a measure of association for cross-classification tables. additionally, we computed the out-of-bag classification error, which is an unbiased measure of misclassification directly obtained as a by-product of rf estimation. the out-of-bag-classification error is based on repeatedly sampling subsets of the available data for estimation; the data not sampled are then used as a test set for evaluating the misclassification probability. a non-metric multidimensional scaling (nmds) was performed in r in order to investigate the relationship between the resulting vegetation types and the environmental variables (bioclimatic, topographic and geological features). results woody species a total of 333 woody species were identified. this included three new records for zimbabwe (commiphora schlechteri, croton steenkampianus and putterlickia verrucosa) and 12 new records for southern zimbabwe. fifteen of the recorded species, within zimbabwe, are confined to the south of the country, with nine of them being recorded only from the gnp. twenty four of the recorded species are classified as red data species within zimbabwe, including four species classified as critically endangered (commiphora neglecta, periploca nigrescens, rinorea elliptica and schotia capitata), two endangered species (adenium multiflorum and milicia excelsa) and five vulnerable species (acacia exuvialis, adenia fruticosa subsp. simplicifolia, manilkara concolor, pachypodium saundersii and suregada zanzibariensis). thirteen alien species were recorded comprising four woody species (lantana camara, senna occidentalis, senna septemtrionalis and withania somnifera) and nine herbaceous species (acanthospermum hispidum, alternanthera pungens, corchorus trilocularis, datura stramonium, euphorbia heterophylla, ocimum americanum var. americanum, sida cordifolia, sorghum halepense and xanthium strumarium). one or more exotic species were recorded from 28 out of the 330 samples (8%), with an overall total of 31 observations. vegetation classification for both classifications, based on separate or merged cover values, the results of the gap statistic indicated that 14 was the optimum number of vegetation types. in both cases, the resulting 14 vegetation types were allocated to three major groups corresponding to biogeographical regions: mopane, miombo and azonal alluvial woodlands. while the differences in classification were relatively minor, as confirmed by the relatively high value obtained for the modified rand index (58%), through comparing diagnostic, constant and dominant species, and in conjunction with expert knowledge, the classification using ‘separate layers’ was considered more effective and representative than the ‘merged layers’ classification. for instance, in the former classification, only one type was considered intermediate or ecotonal between mopane and miombo woodlands (type 12), while for the latter classification two types were ecotonal between mopane and miombo woodlands: one comprised a rather fuzzy type of spirostachys africana woodland and the other had similarities with the surrounding combretaceae mixed woodlands. three major groups were identified consisting of miombo vegetation (covering approximately 46.97% of the gnp), mopane vegetation (38.19%) and alluvial woodlands (6.67%). approximately 8.18% of the relevés (type 12) could not be unequivocally assigned to any of these three main groups and have been classified as ‘intermediate’. the vegetation types identified through the ‘separate layers’ classification, together with the number of relevés for each type, are shown in table 1. although there was considerable variation in the number of relevés per type (range = 12–40, table 1), none of the vegetation types were represented by either a very small or excessively large proportion of relevés (range = 3.6% – 12.1%, table 2). descriptions of the 14 vegetation types based on their dominant, constant and diagnostic species, and a synoptic table with the frequencies of all woody species in each of the 14 types, are presented in online appendixes 1 and 2, respectively. table 1: vegetation types, names and associated numbers of relevés and correspondence to previous vegetation classifications for the gonarezhou national park and other great limpopo transfrontier conservation area parks. table 2: numbers and percentages of relevés (from the field survey) and areas (cells from the spatialisation model) assigned to each of the 14 vegetation types, together with rf out-of-bag error predictions for each type. in the miombo woodlands group (types 2, 5, 6, 9, 10 and 14), dominant species included brachystegia tamarindoides subsp. torrei, combretum celastroides subsp. celastroides, combretum collinum subsp. collinum, guibourtia conjugata, julbernardia globiflora, millettia usaramensis subsp. australis, pteleopsis myrtifolia and terminalia sericea. constant species included xeroderris stuhlmannii (five types), cassia abbreviata subsp. beareana, combretum apiculatum, hugonia orientalis and strychnos madagascariensis (each four types), and combretum collinum subsp. collinum, guibourtia conjugata, monodora junodii v. junodii, pteleopsis myrtifolia and senna petersiana (each three types). the dominant species in the mopane woodlands group (types 1, 4, 7, 8, 11 and 13) were colophospermum mopane and combretum apiculatum, followed by acacia exuvialis, combretum zeyheri and mimusops zeyheri. colophospermum mopane and combretum apiculatum were constant throughout all seven mopane types. type 12 was somewhat intermediary in nature: combretum zeyheri achieved dominance in 4% of the samples, and the presence of colophospermum mopane and combretum apiculatum was 70% and 67%, respectively, which was generally lower than for the other mopane types. while the group included a number of typical mopane constant species (cassia abbreviata subsp. beareana, cissus cornifolia, etc.), it also shared a number of constant species that were characteristic of the miombo woodlands (strychnos madagascariensis and xeroderris stuhlmannii, etc.). the alluvial woodlands were quite distinctive. the only dominant species was cordyla africana, but there were a high number of constant species: acacia tortilis subsp. heteracantha, berchemia discolor, boscia mossambicensis, et cetera. most of these were unique to the alluvial woodlands, other than combretum imberbe, combretum mossambicense, diospyros loureiriana subsp. loureiriana, drypetes mossambicensis, flueggea virosa subsp. virosa and philenoptera violacea, which were shared predominantly with mopane types, but also a few species with the miombo types (combretum mossambicense and drypetes mossambicensis). the results of the nmds are shown in figure 3a for the 14 vegetation types and figure 3b for the three macro-groups. the correlation between the climatic gradient and the mopane and miombo macro-groups is evident (figure 3b). the mopane macro-group is centred on the left of the graphic; these plots were generally located in ‘warmer, more arid and steeper’ areas, while the miombo macro-group is centred on the right of the graphic representing ‘cooler, moister, and higher altitude’ locations. geology does not appear to be important for the segregation of vegetation types, except for the alluvial group. figure 3: non-metric multidimensional scaling of (a) the 14 vegetation types and (b) the three macro-groups. the segregation between the 14 vegetation types is less clear (figure 3a). vegetation types are more readily identifiable within the miombo macro-group (e.g. types 2, 6, 9, 10 and 14). within the mopane macro-group it is difficult to identify single vegetation types except for types 11 and 13. of note is the intermediate position of type 12 which is located at the centre of the diagram. vegetation mapping potential natural vegetation maps for the 14 types and the three biogeographical regions are presented in figure 4. the most potentially widespread types are types 9 and 10 of the miombo region, comprising mixed combretaceae woodland, respectively, with burkea africana (type 9) or guiburtia conjugata (type 10), generally found on sand-loam soils on flat upland areas (figure 4a). type 3 mixed woodland on alluvium is mainly localised on the clay-loam soils along the runde river. the mopane types are mainly restricted to the intervening lowlands, where they occur interspersed with one another (figure 4b). figure 4: map of potential distribution of (a) vegetation types and (b) biogeographical regions. the latter is superimposed on topography to create a 3d effect. assessment procedures highlighted the reliability of the map. in particular, cramer’s index is very low (< 0.007), confirming a very small (and not significant) difference between the proportion of relevés assigned to each vegetation type based on species composition and that of cells classified in the same type based on the rf model (table 2). this result is confirmed by the low out-of-bag prediction error values obtained for each type (range = 0.000–0.048, table 2). discussion vegetation types the paper was aimed at providing an updated and comprehensive analysis of the woody vegetation of the gnp, within the framework of the gltfca. from a methodological point of view, we demonstrated that a classification based on layers can be as effective as a traditional one where species abundance measures are merged in one value. we speculate that in averaging some information might be lost; hence, we recommend clustering based on layers measurements, which give a more direct assessment. the analysis of the 330 relevés allowed the identification of 14 vegetation types belonging to three macro-groups: alluvial (one type), mopane (six types) and miombo (six types) woodlands and one intermediate type between the last two (table 1, online appendixes 1 and 2). the first two correspond to well-known biogeographical regions and confirm the findings of daru et al. (2015) that the gltfca belongs to a specific phyloregion ‘zambesian transition zone’ at the interface of mopane or miombo. we also confirmed that the mopane and miombo, even though phylogenetically similar, differ in respect to environmental parameters, with mopane vegetation being located in the lower lying, drier and warmer parts of the gnp and miombo types in moister and cooler upland areas (figures 3b and 4b). specifically, within the gltfca, beside in the gnp, true miombo (brachystegia or julbernardia) vegetation has been found only in the znp (stalmans & peel 2010), which is also located in the northern part of the area, while it is absent to the south (bnp, lnp and knp). when comparing the results of our analysis with the only previous comprehensive vegetation classification for the gnp (sherry 1970), which found 15 vegetation types, we recorded only 8 types in common (table 1). among those that did not match well are, for instance, sherry’s spirostachys africana-terminalia prunioides woodland that can be generically associated with the mopane macro-group, but not to any one of our types, as spirostachys africana had significant presence in types 4 and 12, and terminalia prunioides in types 11 and 13 (online appendix 2). moreover, sherry’s kirkia, commiphora, adansonia open woodland occupies an intermediate position; kirkia acuminata had a significant presence in type 6 (miombo macro-group), commiphora mollis was predominantly recorded from relevés classified under the mopane macro-group but was not significant for any type, and adansonia digitata was significant for both type 6 and for type 3 mixed woodland on alluvium. similarly, for sherry’s acacia nigrescens-acacia welwitschii tree savanna, acacia nigrescens was significant for type 1 while acacia welwitschii was not a significant component for any of the 14 types here and was mainly recorded from relevés classified under the miombo macro-group. other types of sherry (1970) are difficult to match with any types from the present study. none of their characteristic species, combretum fragrans, terminalia stenostachya, baphia obovata and millettia stuhlmannii, were identified as constant, dominant or diagnostic species in any of the 14 vegetation types described here. these differences could be because of differences in terms of data collection and classification methods. additionally, they may relate to vegetation changes that have occurred during the past 40 years, for example, as described by tafangenyasha (1997, 2001), gandiwa and kativu (2009) and gandiwa et al. (2011a, 2011b), who investigated the decline of several tree species in the gnp. methodological sampling differences also influenced the comparison between the current gnp classification and those of the other parks of the gltfca. however, based on our results, we were able to identify several vegetation types in common. colophospermum mopane is the dominant species in three types that differentiate structurally as woodland (type 7), open woodland (type 4) and bushland (type 13). together with similar vegetation types recognised for the three mozambican parks (bnp, znp and lnp), they can all be compared to the wide and diversified cissus cornifolia–colophospermum mopane community, which was described with a large number of relevés mainly for the knp in the revision of mopaneveld vegetation by siebert, bredenkamp and siebert. (2003). in the gnp, the structural differences among the three types is hypothesised to be because of the combined effect of environmental factors, with type 7 mopane woodland found mainly on clay soils and the other two types on various loamy soils, from clay to sandy loams (online appendix 1), and the impacts of elephants and fire (gandiwa & kativu 2009). colophospermum mopane–acacia nigrescens woodland (type 1) is mainly found on clay soils and similar communities have been found in the lnp, as colophospermum mopane–combretum imberbe woodland (stalmans et al. 2004). colophospermum mopane–combretum apiculatum woodland (type 8) occupies hills and ridges similarly to the vegetation found on the central-western part of the knp (gertenbach 1983) as well as in protected area adjacent to its western boundaries (peel, kruger & macfadyen 2007) and on the rugged veld of the lnp (stalmans et al. 2004). the last vegetation type belonging to the mopane group seems not to have any corresponding type in the other gltfca parks. type 11 (colophospermum mopane woodland with combretum hereroense and terminalia prunioides) occupies the drier areas of the park on shallow soils and the only similar vegetation can be the terminalia prunioides–grewia bicolor thicket of the lnp. type 12 (mixed woodland with colophospermum mopane, combretum apiculatum and combretum zeyheri) is an intermediate type between mopane and miombo as is indicated by its central position in the nmds diagram of figure 3a. in particular, type 12 shares species from the two macro-groups: cassia abbreviata subsp. beareana, cissus cornifolia, combretum mossambicense, dichrostachys cinerea subsp. africana, flueggea virosa subsp. virosa, maerua parvifolia and markhamia zanzibarica from the mopane group, and strychnos madagascariensis, combretum zeyheri, lannea schweinfurthii v. stuhlmannii, pseudolachnostylis maprouneifolia and xeroderris stuhlmannii from the miombo group. however, considering the significant frequency of colophospermum mopane, we decided to include this type in the corresponding group when elaborating the potential natural vegetation maps (figure 4). similar mixed communities are recorded from the lnp (stalmans et al. 2004) and the bnp (stalmans & wishart 2005). mopane vegetation types identified in the gnp are very similar in term of species composition, as can be observed in the nmds diagram (figure 3a) where several types are strongly intermixed (types 1, 4, 7 and 8), and in the map of the potential natural vegetation (figure 4a), which is characterised by a marked salt-and-pepper effect. in this last case, the use of a soil map, still lacking for the entire gnp, would have increased the predictive capacity of the rf model. of the six miombo types identified in the present study, four correspond to types recognised by sherry (1970), one of which is brachystegia tamarindoides woodland (table 1). the continued presence of this type is noteworthy as brachystegia tamarindoides is one of the tree species considered to be in marked decline, according to tafangenyasha (1997, 2001) being almost completely eliminated from some parts of the park. other common types with sherry (1970) include guibourtia conjugata woodland (type 1) and dry deciduous sandveld woodland and scrub, which corresponds to types 9 and10 of the present study (mixed combretaceae woodland with burkea africana or guibourtia conjugata, respectively). the latter two types are the most widespread ones in the gnp, particularly in the central part of the park on cretaceous sedimentary upland areas (figure 4a). guibourtia-characterised vegetation types have also been identified in the other four gltfca parks. within the miombo group, we also included two vegetation types (types 5 and 6) characterised by the presence of androstachys johnsonii. type 5 is similar to the androstachys johnsonii–croton pseudopulchellus woodland that was described by sherry (1970) and also van rooyen, theron and grobbelaar (1981) in the northern part of the knp, and subsequently recognised also for the lnp (stalmans et al. 2004) and bnp (stalmans & wishart 2005). in the case of the gnp, the significant presence of miombo associated species such as brachystegia tamarindoides subsp. torrei, combretum celastroides subsp. celastroides, combretum collinum subsp. collinum, guibourtia conjugata, hugonia orientalis, hymenocardia ulmoides, monodora junodii var. junodii, pseudolachnostylis maprouneifolia, pteleopsis myrtifolia, senna petersiana, strophanthus kombe, strychnos madagascariensis, terminalia sericea, vitex mombassae and xeroderris stuhlmannii, allowed its inclusion in this group, together with type 6, which is very similar, as shown in figure 3a. vegetation on alluvium (type 3) was principally found along the runde river in the north and the mwenezi river in the south (figure 4a). it occurred in fringing ribbons or pockets along water courses, on alluvial terraces and on flood plains. at the save-runde junction, it constitutes one of the largest expanses of this vegetation type in zimbabwe. although riparian woodlands occur widely throughout the country and adjacent regions, the portion around the save-runde junction comprises one of the largest remaining and relatively intact portions and is already formally recognised as comprising one of 20 important bird areas in zimbabwe, on the basis of regularly supporting a significant number of range-restricted bird species (childes & mundy 1997). although alluvial vegetation was found in all the gltfca parks, the most similar one is that from znp along the banks of the save river, with many species in common (e.g. cordyla africana, philenoptera violacea and trichilia emetica). land-use patterns and their influence on the vegetation of the gonarezhou national park direct human disturbances to the vegetation have resulted through clearing for agriculture and bush clearing for the control of tsetse fly. prior to declaration as a park, resident communities cultivated scattered fields on alluvial deposits along parts of the lower runde river and in the vicinity of the save-runde junction. more recently, there has been extensive settlement and cultivation within the northern basalt plain in association with the gulugi drainage (dunham et al. 2010). moreover, the bulk of the vegetation throughout the park is highly degraded. there has been a massive reduction of trees, with upper canopy trees, in particular, having been virtually eliminated or greatly reduced over large areas, as documented in specific studies focused on brachystegia tamarindoides (tafangenyasha 1997, 2001), colophospermum mopane and combretum apiculatum in the northern part of the park (gandiwa & kativu 2009), and androstachys johnsonii and acacia tortilis (gandiwa et al. 2011a, 2011b). many of the remaining large trees have serious bark damage and/or have been reduced to the level of the sub-canopy or even the shrub layer, while many of the younger trees are multi-trunked from having been knocked or burned down to ground level, probably repeatedly, and then resprouted. in many stands, relatively high numbers of dead trees were observed. despite high levels of degradation of vegetation resources, the physical environment has remained relatively intact, with only localised occurrences of sheet and gully erosion, presumably because most of the terrain is virtually flat or gently undulating. as for the neighbouring knp (eckhardt, van wilgen & biggs 2000), the cause of the tree reduction appears to be because of elephants acting in concert with fire (gandiwa & kativu 2009; magadza, coulson & tafangenyasha 1993; tafangenyasha 2001). during the present survey, evidence of elephant damage to woody vegetation and of burning was widespread throughout the park. elephant densities have been sustained at relatively high levels since about 1970, although prior to 1992/1993, elephant populations were kept in check largely through culling. since then, there has been steady growth, to the extent that the total elephant population is now higher than ever recorded, with the overall density approaching two animals per km2, according to results of the most recent aerial survey carried out in september 2009 (dunham et al. 2010). all the four woody alien plant species, recorded during the field campaign, occur in the type 3 mixed woodland on alluvium (online appendix 2). this confirms the findings of foxcroft and richardson (2003), who for the knp indicated riverbanks and floodplains as ideal habitats for the establishment of invasive plants, because of disturbances caused by floods and the competitive capacity of alien species being able to germinate and to establish quicker than indigenous species after floods and to produce allelopathic agents to inhibit the growth of other species. beside alien species, the principal river systems of the gnp have been seriously impacted through upstream dam construction leading to reduced flow regimes and upstream agricultural activities leading to marked siltation and water pollution in the form of agricultural chemicals. conclusion in order to effectively manage protected areas, it is important to have a comprehensive description of the vegetation at a scale that is relevant to management. our work was aimed at updating the knowledge on the vegetation of the gnp in the context of the gltfca where previous vegetation studies concerning the other parks (kruger, limpopo, banhine and zinave) have been recently conducted. moreover, because appropriate management and conservation strategies rely on a better understanding of the functioning of these savanna systems, the predictive method proposed for the elaboration of the natural potential vegetation map of the park is a useful tool for a better understanding the functioning of these savanna systems. its application can be further extended to the other gltfca parks in order to provide a reference model for the elaboration of appropriate and effective transboundary management and conservation strategies. acknowledgements we gratefully acknowledge the assistance of anthony mapaura, tom muller and julius shimbani and staff of the zimbabwe parks and wildlife management authority in collecting the field data, as well as logistical and financial support provided by the frankfurt zoological society. the work has been conducted within the framework of the secosud ii project supported by the italian development cooperation. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions f.a. planned the research; r.c. conducted the field work and built the classification; f.m. performed the arcgis and vegetation analyses; f.m. and f.a. led the writing; a.f. and m.d.s. performed the statistical analyses; and g.d’a. performed the analyses for the validation of the spatialisation model. all authors critically revised the manuscript. references attorre, f., alfò, m., de sanctis, m., francesconi, f., valenti, r., vitale, m. et al., 2011, ‘evaluating the effects of climate change on tree species abundance and distribution in the italian peninsula’, applied vegetation science 14(2), 242–255. http://dx.doi.org/10.1111/j.1654-109x.2010.01114.x attorre, f., francesconi, f., de sanctis, m., alfò, m., martella, f., valenti, r. et al., 2014, ‘classifying and mapping potential distribution of forest types using a finite mixture model’, folia geobotanica 49(3), 313–335. http://dx.doi.org/10.1007/s12224-012-9139-8 bradter, u., thom, t.j., altringham, j.d., kunin, w.e. & benton, t.g., 2011, ‘prediction of national vegetation classification communities in the british uplands using environmental data at multiple spatial scales, aerial images and the classifier random forest’, journal of applied ecology 48, 1057–1065. http://dx.doi.org/10.1111/j.1365-2664.2011.02010.x breiman, l., 2001, ‘random forests’, machine learning 45, 5–32. http://dx.doi.org/10.1023/a:1010933404324 brzeziecki, b., kienast, f. & wildi, o., 1993, ‘a simulated map of the potential natural forest vegetation of switzerland’, journal of vegetation science 4, 499–508. http://dx.doi.org/10.2307/3236077 childes, s.l. & mundy, p.j., 1997, important bird areas of zimbabwe, birdlife international, cambridge, united kingdom. chytrý, m., tichý, l., holt, j. & botta-dukát, z., 2002, ‘determination of diagnostic species with statistical fidelity measures’, journal of vegetation science 13, 79–90. http://dx.doi.org/10.1111/j.1654-1103.2002.tb02025.x daru, b.h., van der bank, m., maurin, o., yessoufou, k., schaefer, h., slingsby, j.a. et al., 2015, ‘a novel phylogenetic regionalization of phytogeographical zones of southern africa reveals their hidden evolutionary affinities’, journal of biogeography 43(1), 155–166. http://dx.doi.org/10.1111/jbi.12619 dunham, k.m., van der westhuizen, e., van der westhuizen, h.f. & gandiwa, e., 2010, aerial survey of elephants and other large herbivores in gonarezhou national park (zimbabwe), zinave national park (mozambique) and surrounds: 2009. parks and wildlife management authority, harare. eckhardt, h.c., van wilgen, b.w. & biggs, h.c., 2000, ‘trends in woody vegetation cover in the kruger national park, south africa between 1940 and 1998’, african journal of ecology 38, 108–115. http://dx.doi.org/10.1046/j.1365-2028.2000.00217.x faith, d.p., minchin, p.r. & belbin, l., 1987, ‘compositional dissimilarity as a robust measure of ecological distance’, vegetatio 69, 57–68. http://dx.doi.org/10.1007/bf00038687 farrell, j.a.k., 1968, ‘preliminary notes on the vegetation of the lower sabi-lundi basin, rhodesia’, kirkia 6, 223–248. foxcroft, l.c. & richardson, d.m., 2003, ‘managing alien plant invasions in the kruger national park, south africa’, in l.e. child, j.h. brock, g. brundu, k. prach, p. pyšek, p.m. wade et al. (eds.), plant invasions: ecological threats and management solutions, pp. 385–403, backhuys publishers, leiden, the netherlands. franklin, j., 1995, ‘predictive vegetation mapping: geographic modeling of biospatial patterns in relation to environmental gradients’, progress in physical geography 19, 474–499. http://dx.doi.org/10.1177/030913339501900403 gandiwa, e., chikorowondo, g., zisadza-gandiwa, p. & muvengwi, j., 2011a, ‘structure and composition of androstachys johnsonii woodland across various strata in gonarezhou national park, southeast zimbabwe’, tropical conservation science 4(2), 218–229. gandiwa, e., heitkonig, i.m.a., eilers, p.h.c. & prins, h.h.t., 2016, ‘rainfall variability and its impact on large mammal populations in a complex of semi-arid african savanna protected areas’, tropical ecology 57(2), 163–180. gandiwa, e. & kativu, s., 2009, ‘influence of fire frequency on colophospermum mopane and combretum apiculatum woodland structure and composition in northern gonarezhou national park, zimbabwe’, koedoe 51(1), 36–41. http://dx.doi.org/10.4102/koedoe.v51i1.685 gandiwa, e., magwati, t., zisadza, p., chinuwo, t. & tafangenyasha, c., 2011b, ‘the impact of african elephants on acacia tortilis woodland in northern gonarezhou national park, zimbabwe’, journal of arid environments 75, 809–814. http://dx.doi.org/10.1016/j.jaridenv.2011.04.017 gandiwa, e., tupulu, n., zisadza-gandiwa, p. & muvengwi, j., 2012, ‘structure and composition of woody vegetation around permanent-artificial and ephemeral-natural water points in northern gonarezhou national park, zimbabwe’, tropical ecology 53, 169–175. germishuizen, g. & meyer, n.l., 2003, plants of southern africa: an annotated checklist. strelitzia 14, national botanical institute, pretoria, south africa. gertenbach, w.p.d., 1983, ‘landscapes of the kruger national park’, koedoe 26, 9–121. http://dx.doi.org/10.4102/koedoe.v26i1.591 gillet, f. & gallandat, j.d., 1996, ‘integrated synusial phytosociology: some notes on a new, multiscalar approach to vegetation analysis’, journal of vegetation science 7, 13–18. http://dx.doi.org/10.2307/3236411 hennekens, s.m. & schaminée, j.h.j., 2001, ‘turboveg, a comprehensive data base management system for vegetation data’, journal of vegetation science 12, 589–591. http://dx.doi.org/10.2307/3237010 kubota, y., hirao, t., fujii, s.j., shiono, t. & kusumoto, b., 2014, ‘beta diversity of woody plants in the japanese archipelago: the roles of geo-historical and ecological processes’, journal of biogeography 41, 1267–1276. http://dx.doi.org/10.1111/jbi.12290 lister, l.a., 1987, the erosion surfaces of zimbabwe. bulletin no. 90, zimbabwe geological survey, geological survey department, harare. magadza, c., coulson, i. & tafangenyasha, c., 1993, ‘ecology of gonarezhou national park’, in b.k. downie (ed.), gonarezhou national park management planning programme, phase i, march 1993: background data reports. department of national parks and wild life management, harare. maggini, r., lehmann, a., zimmermann, n.e. & guisan, a., 2006, ‘improving generalised regression analysis for the spatial prediction of forest communities’, journal of biogeography 33, 1729–1749. http://dx.doi.org/10.1111/j.1365-2699.2006.01465.x mellor, a., haywood, a., stone, c. & jones, s., 2013, ‘the performance of random forests in an operational setting for large area sclerophyll forest classification’, remote sensing 5, 2838–2856. http://dx.doi.org/10.3390/rs5062838 nyamapfene, k., 1991, soils of zimbabwe, nehanda publishers, harare. o’brien, e.m., 1993, ‘climatic gradients in woody plant species richness: towards an explanation based on an analysis of southern africa’s woody flora’, journal of biogeography 20, 181–198. http://dx.doi.org/10.2307/2845670 o’brien, e.m., field, r. & whittaker, r.j., 2000, ‘climatic gradients in woody plant (tree and shrub) diversity: water energy dynamics, residual variation, and topography’, oikos 89, 588–600. http://dx.doi.org/10.1034/j.1600-0706.2000.890319.x o’connor, t.g. & campbell, b.m., 1986, ‘classification and condition of the vegetation types of the nyahungwe area on the lundi river, gonarezhou national park, zimbabwe’, south african journal of botany 52, 117–123. http://dx.doi.org/10.1016/s0254-6299(16)31575-7 peel, m.j.s., kruger, j.m. & macfadyen, s., 2007, ‘woody vegetation of a mosaic of protected areas adjacent to the kruger national park, south africa’, journal of vegetation science 18, 807–814. http://dx.doi.org/10.1111/j.1654-1103.2007.tb02597.x rodriguez-valiano, v.f., ghimire, b., rogan, j., chica-olmo, m. & rigol-sanchez, j.p., 2012, ‘an assessment of the effectiveness of a random forest classifier for land-cover classification’, isprs journal of photogrammetry and remote sensing 67, 93–104. http://dx.doi.org/10.1016/j.isprsjprs.2011.11.002 sherry, b.y., 1970, vegetation map of gonarezhou national park, unpublished map, department of national parks, harare. siebert, f., bredenkamp, g.j. & siebert, s.j., 2003, ‘a comparison of mopaneveld vegetation in south africa, namibia and zimbabwe’, bothalia 33, 121–134. http://dx.doi.org/10.4102/abc.v33i1.439 stalmans, m., gertenbach, w.p.d. & carvalho-serfontein, f., 2004, ‘plant communities and landscapes of the parque nacional do limpopo, mocambique’, koedoe 47, 61–81. http://dx.doi.org/10.4102/koedoe.v47i2.83 stalmans, m. & peel, m., 2010, ‘plant communities and landscapes of the parque nacional de zinave, mozambique’, koedoe 52(1), 1–11. http://dx.doi.org/10.4102/koedoe.v52i1.703 stalmans, m. & wishart, m., 2005, ‘plant communities, wetlands and landscapes of the parque nacional de banhine, moçambique’, koedoe 48, 43–58. http://dx.doi.org/10.4102/koedoe.v48i2.99 tafangenyasha, c., 1997, ‘tree loss in the gonarezhou national park (zimbabwe) between 1970 and 1983’, journal of environmental management 49, 355–366. http://dx.doi.org/10.1006/jema.1996.9987 tafangenyasha, c., 2001, ‘decline of the mountain acacia, brachystegia glaucescens in gonarezhou national park, southeast zimbabwe’, journal of environmental management 63, 37–50. http://dx.doi.org/10.1006/jema.2001.0458 thompson, j.g. & purves, w.d., 1978, a guide to the soils of zimbabwe, zimbabwe agricultural journal technical handbook no. 3, government printer, harare. tibshirani, r., walther, g. & hastie, t., 2001, ‘estimating the number of data clusters via the gap statistic’, journal of the royal statistical society 63, 411–423. http://dx.doi.org/10.1111/1467-9868.00293 tichy, l., 2002, ‘juice, software for vegetation classification’, journal of vegetation science 13, 451–453. http://dx.doi.org/10.1111/j.1654-1103.2002.tb02069.x timberlake, j., nobanda, n. & mapaure, i., 1993, ‘vegetation survey of the communal lands – north and west of zimbabwe’, kirkia 14, 171–270. van rooyen, n., theron, g.k. & grobbelaar, n., 1981, ‘a floristic description and structural analysis of the plant communities of the punda milia-pafuri-wambiya area in the kruger national park’, journal of south african botany 47, 585–626. venter, f.j. & gertenbach, w.p.d., 1986, ‘a cursory review on the climate and vegetation of the kruger national park’, koedoe 29, 139–148. http://dx.doi.org/10.4102/koedoe.v29i1.526 wild, h., 1955, ‘observations on the vegetation of the sabi-lundi junction area’, rhodesia agricultural journal 52, 533–546. zimmermann, n.e., edwards, t.c., moisen, g.g., frescino, t.s. & blackard, j.a., 2007, ‘remote sensing-based predictors improve distribution models of rare, early successional and broadleaf tree species in utah’, journal of applied ecology 44, 1057–1067. http://dx.doi.org/10.1111/j.1365-2664.2007.01348.x zisadza-gandiwa, p., parakasingwa, c., mashapa, c., muboko, n. & gandiwa, e., 2013, ‘status of woody vegetation along riparian areas in gonarezhou national park, zimbabwe’, greener journal of agricultural sciences 3, 592–597. article information authors: tony g. rebelo1 stefanie freitag2 chad cheney3 melodie a. mcgeoch4 affiliations: 1threatened species research programme, south african national biodiversity institute, south africa 2kruger national park, south african national parks, south africa 3table mountain national park, south african national parks, south africa 4cape research centre, south african national parks, south africa correspondence to: tony rebelo email: rebelo@sanbi.org postal address: private bag x1, claremont 7735, south africa dates: received: 11 aug. 2010 accepted: 02 mar. 2011 published: 02 mar. 2011 how to cite this article: rebelo, t.g., freitag, s., cheney, c. & mcgeoch, m.a., 2011, ‘prioritising species of special concern for monitoring in table mountain national park: the challenge of a species-rich, threatened ecosystem’, koedoe 53(2), art. #1019, 14 pages. doi:10.4102/koedoe.v53i2.1019 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) prioritising species of special concern for monitoring in table mountain national park: the challenge of a species-rich, threatened ecosystem in this essay... open access • abstract • introduction    • the scope of the problem    • systems for determining conservation priorities    • the sanparks context • methods    • study area, scale of assessment and baseline information    • definition of species of special concern    • selection of prioritisation variables    • summing versus categorisation methods • results    • comparison of summing versus categorisation methods    • independence of variables    • is a biotic score appropriate?    • can priority species of special concern be delimited? • discussion    • towards a uniform system across taxa    • problems: representation of lowland species • conclusion • acknowledgements • references • appendix 1 abstract (back to top) conservation requires that species are monitored to ensure the persistence of species and ecosystem processes. in areas with large numbers of threatened species, this can be a major challenge. here we explore prioritising species of special concern on the cape peninsula, south africa, conserved primarily in the table mountain national park. with 307 terrestrial plant and animal species listed as threatened on the iucn red list (plus 208 as non-least concern) and 332 endemic to the peninsula, it is impossible to monitor and manage all species with current resources. at a workshop of conservation managers and ecosystem and taxonomical specialists, 14 variables were incorporated into a simple scoring scheme to develop a priority listing of these species. despite care to ensure that variables were independent, there was strong autocorrelation amongst biotic versus management variables. there was concern that biotic variables would be masked by management criteria, but this was not the case. we propose that monitoring should focus on as many top-scoring species as resources allow (including volunteers) and that setting a cut-off value for delimiting sensitive species should be eschewed. a major challenge is that many species are typical of lowland ecosystems, which are poorly represented in the national park. although priority species for monitoring have been identified, this will need to be tempered with the monitoring costs and logistics of implementing the programme. conservation implications: owing to the large number of threatened and endemic species in the cape peninsula, it is impossible to monitor all species with current resources. management must focus on ecosystem maintenance as species-focused management will inevitably result in conflict with other threatened species. monitoring should focus on as many top-scoring species as resources allow. the costs and logistics of a monitoring programme still need to be worked out. introduction (back to top) one of the principle roles of protected areas, and in many instances the motivation for their establishment, is to prevent the extinction of species (brooks et al. 2002; vellak et al. 2009). the risk for species extinction, locally and globally, is negatively correlated with both distributional extent (range) and abundance (or population size) (gaston et al. 2008). these variables thus form the basis for the measurement and assessment of extinction risk and the management of threatened species (bakker & doak 2009). the internatinal union for conservation of nature red list provides a system for classifying the extinction threat of species at a global scale (iucn 2006). however, global extinction risk does not always reflect regional conservation priorities (bubb et al. 2009) and it is necessary to consider other factors when planning finer-scale conservation strategies (freitag & van jaarsveld 1997; gardenfors et al. 2001; given & norton 1993; hansen et al. 1999; miller et al. 2007; mace et al. 2008; schnittler & gunther 1999). conservation action takes place at local scales, such as within the bounds of protected areas or protected area networks within biomes, and prioritising species for monitoring and conservation action must necessarily take place at this scale. this prioritisation process has three components (which are not necessarily sequential or mutually exclusive), namely: • identifying which species in the species pool are to be considered species of special concern • prioritising (ranking) the species on this list using a selected range of criteria • deciding which of these species are to be monitored and which species require conservation action. here we report on the outcome of such a process (specifically the first two points) for the initial prioritisation of species in the table mountain national park (tmnp) in the biodiversity-rich fynbos biome. the decision about the species selected for monitoring is constrained by the financial and human resources available and will be the subject of a forthcoming analysis (rebelo, freitag-ronaldson, cheney & mcgeoch, unpublished results). south african national parks (sanparks) is mandated to conserve plant and animal species within parks as well as the ecosystems they represent. to this end the organisation must monitor and, where necessary, take conservation action to ensure that species in national parks do not become extinct. the cape floristic region (cfr) is particularly rich in plant species and also in rare and threatened species (goldblatt & manning 2000; raimondo et al. 2009). as a result there are an inordinately large number of candidate species for monitoring and managing – a significant challenge given data inadequacies and resource shortages. the tmnp was thus chosen as a pilot study for developing a system for selecting species for monitoring, particularly in species-rich situations. the scope of the problem the cape peninsula is one of the richer centres of endemism in the fynbos biome of the cfr (goldblatt & manning 2000), with 2500 of the 9000 plant species of the region; more than 194 species are not found elsewhere on earth. furthermore, the flora of the cfr is exceptionally threatened: more than half (1736) of all threatened red list plant species in south africa occur here (raimondo et al. 2009). the distribution of these threatened species is also focussed on the cape peninsula, with 319 plant species recorded from the city of cape town magisterial district and 182 of these historically recorded on the peninsula (rebelo et al. 2011). the distribution should be seen in context of the small size of the area, with one threatened plant species for each 3 km2 of the peninsula. in addition to the high native diversity, some 424 alien plant species are naturalised on the peninsula (rebelo, unpublished), thereby constituting 14% of the total flora. therefore, although the peninsula is species rich, alien invasive species pose a significant threat to its native flora and fauna (richardson & van wilgen 2004). although species monitoring remains essential, the presence of 2500 indigenous plant species on the peninsula requires that the conservation approach focus on ecosystem management, as species-specific interventions will inevitably negatively impact other species. however, species monitoring is also essential for other reasons. fynbos is noted for its high beta and gamma diversity (goldblatt & manning 2000). as a result, monitoring a few ’indicator’ variables or species will unlikely reflect population and species-specific changes across the peninsula. this situation also poses significant challenges to the development of thresholds of potential concern for rare species, described as monitoring endpoints that define the upper and lower levels along a continuum of change, beyond which action is necessary (biggs & rogers 2003; foxcroft 2009). the high spatial and temporal dynamics for fynbos plant communities (bond, maze & desmet 1995; privett, cowling & taylor 2001; rebelo et al. in press; thuiller et al. 2007) is another particular challenge to designing appropriate monitoring systems for relevant species. where there is conflict between ecosystem and species management, the long-term survival of species in the wild – as well as that of their many associated competitors, symbionts and other species – is possible only within the context of a natural ecosystem. short-term emergency interventions to save particular species are often best managed ex situ, but long-term conservation must necessarily take place in situ. ideally all threatened taxa and all locally endemic taxa should be monitored, but even this is clearly a substantial challenge. at the very least all critically endangered taxa – those most threatened with imminent extinction – have to be monitored within the tmnp. however, it is unclear whether this is a realistic goal given available resources. because no prioritisation exercise has been attempted to date, there is little information available to inform planning for monitoring and subsequent management actions to be triggered by monitoring for species of special concern. systems for determining conservation priorities the ideal system for prioritising and guiding a monitoring programme is one that considers the actual value of the species with regard to their intrinsic values (e.g. diversity, phylogeny and function) (bengtsson, jones & setala 1997; turpie 2003), costs of monitoring, managing and saving the species, as well as the costs incurred by delaying an intervention (joseph, maloney & possingham 2009). this would allow a budget of costs to be compiled and different strategies to be evaluated against available funds, or a particular budget to be justified against predetermined goals (mace, possingham & leader-williams 2006). unfortunately, such data do not exist for even the most common species, let alone all the species anticipated for a species-rich region such as the cape peninsula. consequently, the consideration of a range of other prioritisation variables is unavoidable. the options for prioritising species for monitoring and management involve either scoring and summing, or categorising criteria and assigning scores, followed by ranking. this process is based on a series of selected variables of conservation priority against which each species is assessed (de grammont & cuaron 2006). for example, regan et al. (2008) used an approach in which species were scored according to known threats, the spatial and temporal extent of threats, and species and population characteristics. species are subsequently ranked by these variables and for each class (or group) formed, a focal species is selected for monitoring. alternatively, species may be given values for variables of interest, which are then aggregated (e.g. summed) in some manner to provide a total score for each species. although both schemes require variables to be selected, and each species subsequently to be scored according to predetermined criteria, in reality, both the assigning of variables and their weighting are influenced by the chosen approach. therefore it is not straightforward to compare the outcomes of the various approaches. such approaches are nonetheless simple to implement, provided that comparative data are available across all species. the sanparks context the mandate of the tmnp (as for other national parks) derives from the protected areas act (act 57 of 2003) and the biodiversity act (act 10 of 2004). the specific biodiversity objectives (as outlined in the park management plan) are predicated by institutional biodiversity values. briefly, these are to: • adopt a complex systems view to ensure natural functioning and long-term persistence of ecosystems • conserve representative and complementary components of biodiversity to ensure resilience and ecosystem integrity • use responsible and sustainable management interventions under a ‘minimum interference’ philosophy, which recognises that natural and social systems change over time. the aim of biodiversity management in the tmnp is to maintain the natural patterns and processes of the land and seascapes of the tmnp (sanparks 2008). this includes all representative ecosystems on the cape peninsula, which the tmnp represents as the only sizeable conservation area. as part of this goal, understanding and knowledge of species population levels and trends are essential. these data are also required to update the iucn red list and national red lists, required nationally by the department of environmental affairs. appropriate monitoring and action thresholds (martin et al. 2009) for all red-listed species of conservation concern (raimondo et al. 2009) will be required, without excluding species for which there is insufficient information for formal red-listing. collaboration with other conservation agencies and initiatives is essential to secure the future of all species over their historical distribution ranges. part of the mandate is to prevent the extinction of any species within the cape peninsula. management for ecosystem integrity and biodiversity should nevertheless take precedence over single species management, except perhaps in the case of keystone species. a species of special concern monitoring programme has been identified as one of 10 monitoring programmes constituting the sanparks biodiversity monitoring system (see mcgeoch et al. 2011). monitoring species of special concern, for example by means of the red list index (butchart et al. 2005), is essential to quantify changes in the conservation status of a species and, in cases where extinction risk has increased, to take conservation action. here the outcome of a pilot study for the tmnp is presented as an approach that may be considered for more widespread adoption within sanparks. the results for the tmnp are presented with reference to (1) the conservation priority variables selected, (2) a description of the scoring and ranking processes applied to species, and (3) the implications of the outcome for species conservation and monitoring on the cape peninsula. methods (back to top) a one-day workshop was hosted by the south african national biodiversity institute (kirstenbosch research centre) in november 2007 (younge-hayes, rebelo & cheney 2007). the workshop focussed on selecting the prioritisation variables, developing a procedure for scoring and ranking, and evaluating the output of the system with candidate species. the objective was to reach agreement and obtain ideas for determining which species require monitoring, given available resources and existing threats. a series of questions was posed to guide discussions at the workshop, based on the outcome of species scoring and ranking conducted by a.g. rebelo (unpublished) prior to the workshop. • which ‘obvious’ species are missing for the park? these could not be based on theory or principle; specific species names were required. • do you agree with the prioritisation variables? which variables are missing? • do you agree with the scoring and scoring criteria? can they be refined? • does the ranking of the species make sense? if not, what needs to be done? the outcome of the workshop (also presented in younge-hayes et al. [2007]) is summarised in tables 1 and 2 and the discussion section of this paper. based on the refinement of variables and scoring of species for each of the variables indentified during the workshop, especially those related to monitoring, the database was repopulated. final values for species and rankings are provided in supplementary table 1. table 1: variables used to establish the conservation priority of species of special concern for determining monitoring priorities on the cape peninsula, south africa. priorities were established by scoring each species (0–3) based on criteria for each variable and the percentages of species (n = 776) allocated to each score are shown for each variable. study area, scale of assessment and baseline information clearly establishing the extent of the area under consideration is essential for evaluating priorities for species conservation and the species scores attributed are necessarily context specific. traditionally, the cape peninsula is the area west of a line from rietvlei to sandvlei (470 km2) on the south-western end of africa (see figure 1; also adamson & salter 1950). the tmnp (250 km2) represents 53% of the area and is the only large reserve on the cape peninsula. a buffer zone of 5 km around the borders of the tmnp covers the entire peninsula as defined above, or up to 18º30’ e. the entire tmnp is embedded within the city of cape town. this area thus covers the peninsula mountain chain as well as lowland areas to the east, traditionally known as the cape flats. although only a small proportion of the cape flats occurs in the cape peninsula as defined above, several critical conservation areas occur there. figure 1: table mountain national park and other conservation areas on the cape peninsula. the black line is the classical boundary of the cape peninsula from rietvlei to sandvlei. the edge of the map approximates an alternative definition, viz. 18o30’ e. smaller reserves within the peninsula (all managed by the city of cape town) include raapenberg (8 ha), rondebosch common (38 ha), kenilworth race course (42 ha), sandvlei (50 ha plus 160 ha open water) and rondevlei (220 ha plus 70 ha open water). these are all on the cape flats, yet this veld type is represented by only 177 ha in the tmnp at tokai. however, most of the latter is still under plantations, which will be incrementally harvested and restored by 2025 (sanparks 2009). the only other remaining areas of this habitat, albeit significantly degraded, are rondebosch east (15 ha), wingfield airfield (65 ha), meadowridge (5 ha), princessvlei (70 ha) and capricorn park (5 ha). with the exception of princessvlei, sandvlei, rondevlei and capricorn park, which are largely cape flats dune strandveld, the other cape flats reserves are the species-rich cape flats sand fynbos (rebelo et al. 2011). the tmnp thus contains most of cape flats sand fynbos already being conserved(66%) or available for conservation (51%) in the cape peninsula and presents, arguably, the only area capable of being managed as a viable natural ecosystem. with the exception of cape flats dune strandveld, all other vegetation types on the peninsula are conserved exclusively in the tmnp. furthermore, four vegetation types, namely peninsula granite fynbos, peninsula sandstone fynbos, peninsula shale fynbos and peninsula shale renosterveld, are endemic to the peninsula and conserved exclusivelyin the tmnp (rebelo et al. 2006, 2011). the natural history of the peninsula is relatively well known. an earlier botanical handbook (now dated) gives distribution data of indigenous and naturalised species (adamson & salter 1950). the guthrie herbarium, which forms part of the bolus herbarium, deals specifically with peninsula species. various local field guides also exist (e.g. trinder-smith 2006), including some for specific groups such as ferns (roux 1979), trees (moll & scott 1981), ericaceae (oliver & oliver 2000), restionaceae (haaksma & linder 2000), and proteaceae (rebelo 2000). in 1996, the cape peninsula was the focus of a special issue of the journal biodiversity and conservation (vol. 5), where, amongst others, threats and endemics of both animals and plants were reviewed. conservation issues for the city itself are considered in rebelo et al. (2011). local red lists have been compiled for almost 25 years (e.g. hall & ashton 1983), and the scale of urbanisation in the area has meant that extinctions and near-extinctions on the peninsula are often explicitly recorded in the national red list of plants (raimondo et al. 2009). all scientific names used in this manuscript are supplied in supplementary table 1 and botanical nomenclature follows goldblatt and manning (2000). definition of species of special concern the following categories of taxa were considered to fall within the definition of species of special concern: • red list taxa in threatened (cr, en and vu) or conservation concern (ex, ew, nt, lc-rare, dd) categories (e.g. silvertree, with an iucn red list status of en (see table 1 and iucn (2006) for definitions of status abbreviations) endemic taxa (e.g. table mountain ghost frog), defined as taxa with more than 80% of their range, populations or individuals confined to the peninsula • locally threatened taxa (e.g. spinning conebush, known from a single extant and two extinct populations on the peninsula, but common and widespread elsewhere) • re-introduced taxa that are extinct (e.g. whorled heath), threatened (e.g. flats conebush), cannot be adequately conserved elsewhere (e.g. bontebok), or that have been re-introduced recently and still are regarded technically extinct on the peninsula (i.e. have not maintained their populations for three generations, such as the klipspringer) • taxa of special management concern (e.g. baboon, which is an iconic, yet damage-causing animal on the urban edge and tourism nodes) • taxa that were monitored in the past (e.g. false heath, which is no longer considered threatened under iucn criteria but was previously ranked as threatened under local criteria) • taxa that were previously considered as one of the above or could be confused with one of the above; these were included for cross-referencing, database maintenance and system evaluation. these include taxa that have been split into peninsula-based and non-peninsula-based taxa: taxa that were erroneously thought to occur on the peninsula in the past or that were endemic to the peninsula, but are now considered part of a more widespread taxon. a few benchmark species were also included to determine how species, which were agreed to be unimportant, would rank on the system. ideally all taxa of animals, plants, fungi and microorganisms of the peninsula should be monitored and assessed for conservation status. in reality, however, funding and time are limited, expertise and data are lacking, and it is not feasible to monitor even all species of special concern as defined above. as a result, including low priority species in the prioritisation process is of little benefit, other than to check that the system does not overly score such species at the expense of more obviously important ones. in addition, although all populations of all species should be evaluated (and any published subspecies or variants), this is, unfortunately, beyond the scope of current resources and information. consequently, taxa associated with other monitoring processes were explicitly excluded, as their monitoring will not necessarily be compatible with monitoring for threats and declines. these include: • alien species and their biocontrol agents • species subject to harvesting • keystone and indicator species • species for monitoring fire dynamics (mainly serotinous species, especially proteaceae), water extraction, climate change and other ecosystem processes. marine and shoreline taxa were also excluded, although species breeding above the high water mark were included. it is not clear whether having a single listing for terrestrial and marine systems would benefit management and this aspect was not explored further. another issue is whether only species of special concern should be considered for evaluation to begin with (as described above), or whether all taxa should potentially be considered and those exceeding a specific score should be labeled as species of special concern. in other words, should inclusion be determined a priori based on pre-selected variables, or should initial inclusion be determined by some cut-off value independent of the individual variables themselves, i.e. those species scoring over 30, or the top 100 scoring species? a priori inclusion criteria may even be independent of variables being considered, for example, only non-flying mammals on the peninsula less than 50 kg in mass. for purposes of monitoring species on the peninsula, the prioritisation system adopted allows for continuous evaluation of all species (i.e. inclusive), both with regard to continued monitoring of a particular species or determining the next species that requires monitoring, depending on resource availability. selection of prioritisation variables clearly, chosen prioritisation variables should be applicable across all relevant taxa that have available data. where hard data values are not available, they should be inferred, or alternatively, default values should be meaningful. where data are consistently lacking, care should be taken that missing data are not scored too low, otherwise these species may never feature in the scheme. this is especially important for invertebrates and other groups that tend not to have as much data as plants and vertebrates. most importantly, criteria should be explicitly defined to prevent manipulation by increasing the scores of favourite or iconic species. variable values should be constant for a species and should not vary from one location to another; this can be resolved by selecting a mean or extreme value. variables should be independent of one another and, even if used as a proxy for some other variable that is less tangible or harder to measure, should not be strongly correlated to other variables used. for example, ’iucn red list status’, ‘rate of population decline’ and ’degree of threat’ are equivalents and cannot justifiably be considered independent variables in the assessment. it stands to reason that the variables and their scoring should be carefully considered. one of the most important principles is that values should not be double counted. similarly, if variables are independent and their scores ranked so as to be equivalent, then no weighting or complicated summing procedures should be required. in reality, however, it is almost impossible to obtain variables that are not correlated as there are, invariably, some taxa or attributes that are co-linear. for example, taxa with large individuals tend to be threatened with higher rates of extinction because they take longer to mature, produce less progeny per parent, require larger foraging areas, are less able to hide or escape, are preferentially utilised by humans, etc. whereas size is an easy metric to recognise in obvious correlations, other less apparent relationships undoubtedly exist. therefore, although some degree of inclusion of correlated variables may be unavoidable, the inclusion of obvious and strongly correlated ones should be well justified, or the consequence of their joint inclusion at least explored prior to use. prior to the workshop, all variables used nationally and locally for monitoring and evaluating were compiled, including those from the kruger national park (freitag 2011), capenature (g. palmer, pers. comm.) and gauteng (pfab 2002). the variables used are defined in table 1, while those that proved problematic and therefore were not used are shown in table 2. the variables used can be grouped as those referring to: • the taxon in relation to the park and its ecology and threats – the biotic score • variables that influence management of the taxon and monitoring specifically – the management score. table 2: variables not inclued for ranking of species of special concern for determining monitoring priorities on the cape peninsula, south africa. problems with these variables include variations between population, lack of data, vague and complicated circumscriptions, and subjective threats that do not hold under detailed scrutiny. the variables selected for this exercise are not to be confused with variables that may be measured when the selected species are actually monitored on the ground; that is, only a subset of variables are shown in table 1 and are likely to be measured in the process of monitoring the selected species. summing versus categorisation methods the proposed aggregation method was used simply to assign and then add the scores for each species across all the variables. the maximum score would thus be 42 points and the minimum 2 (table 1). taxa with the same score are equal in rank and this method thus places each species along a continuum of monitoring priority (for clarity we refer to this summing method as ‘method 1’). for comparison, an alternative method based on a categorisation system (see de grammont & cuaron 2006) was used, referred to here as ‘method 2’. this method ranks species according to four variables, namely (1) red list status, followed by (2) urgency, and then (3) endemism and (4) importance of park (see table 1). with method 2, the species are scored based on the four mentioned variables, with the highest possible category of species assigned a ranking of 3, 3, 3, 3 (or 3333 for convenience) and the lowest category of species assigned a ranking of 0000 (see table 1). results (back to top) some 776 taxa (including plants, mammals, birds, frogs, reptiles, fish and some groups of invertebrates) were included in the list for assessment, with 28 being ’extraneous’ (i.e. no longer considered to occur on the peninsula [appendix]). the peninsula has 245 plant and 62 animal taxa listed as threatened on the iucn red list, and a further 154 and 24 taxa, respectively, that are of ’conservation concern’ (i.e. not ‘least concern’), as shown in table 3. originally 192 plant taxa were listed as endemic to the peninsula, but on completion of the red list these were reduced to 183, with an additional 149 animal taxa considered to be endemic to the peninsula at present (tables 3 and 4). although 89 taxa are on the local peninsula red list (table 4), most of these (80%) also have global red list status that exceeds ‘least concern’. not all locally extinct animals were included, but most of those with a threatened iucn red list status were. seven animal species currently monitored by the tmnp and associates (african penguin, black oystercatcher, bontebok, chacma baboon, grey rhebuck, klipspringer and the western leopard toad) were included for evaluation. table 3: numbers of species of special concern, including taxa on the iucn red list, local red list or labelled as endemica taxa, on the cape peninsula, south africa. table 4: endemic or threatened plant taxa (according to global iucn red list and local red list) on the cape peninsula, south africa. beetles, frogs, spiders and scorpions are listed as threatened on the final list of species of special concern (supplementary table 1), whilst millipedes, centipedes, crustaceans, earthworms and bugs (insecta, hemiptera) are listed as endemic (see table 3). amongst the plants, nine families each had more than 10 threatened taxa, one had more than 10 locally threatened taxa, and eight families had 10 or more endemic taxa (table 4). the 10 top-scoring taxa (table 5) were accepted by participants at the workshop. these include a mix of plants and animals, with a strong emphasis on taxa listed as extinct in the wild or critically endangered. the silvertree, an enigmatic emblem on table mountain, features on the list primarily owing to its public profile and past conservation efforts. two locally extinct antelope that have been re-introduced to the peninsula and are currently being monitored, also feature. comparison of summing versus categorisation methods as shown in table 5, comparison of the top-scoring species according to method 1 (as described in the previous section) versus the outcome of method 2 yielded 59 taxa in the top category (viz. 3333). these featured seven of the top 10 species in table 5 as well as species scoring as low as 17 (appendix and figure 2). on its own, method 2 is inadequate because there are far too many species and its discriminatory power is weak. it is thus not obvious how to rank the top 59 taxa (appendix), other than by using method 1. the top category taxa include a butterfly, a velvet worm, an amphipod, a frog, 15 beetles and 40 plants (including nine ericas, seven proteas and four sedges). it is also not obvious which further variables should be included next to refine categorisation for determining prioritisation. further refinement is thus arbitrary. similarly, it is not clear a priori how to contrive a ranking by weighting criteria in any particular way. more could possibly be achieved with a sophisticated categorisation procedure, but having decided not to construct complicated summing procedures, using a complicated categorisation procedure seemed equally contrived and was not explored further. figure 2: top categorisation (method 2) based on variables ‘red list’, ‘urgency’, ‘endemism’ and ‘importance of park’ versus summing for species of special concern on the cape peninsula, south africa. the dashed line indicates the 10 top-scoring taxa using the summing method, and the solid line the top score possible for extraneous species with the variables and criteria used. independence of variables despite careful selection of variables, in reality, the variables were found to be strongly, and always positively, correlated. the variables ‘importance of park’, ‘relative endemicity’ and ‘available habitat’ were orthogonal to most other variables, but there was no tight clustering (figure 3). management attributes tend to cluster together, but biotic attributes were widely dispersed. redundancy is suggested only for ‘importance of park’ and ‘available habitat’. figure 3: principle components analysis of variables for species of special concern on the cape peninsula, south africa. components 1 and 2 accounted for 18% and 17% of the variance, respectively. is a biotic score appropriate? participants at the workshop were worried that the biotic score would be overwhelmed by the management criteria and that taxa important for monitoring would be neglected in favour of taxa that were easier to monitor. however, the 10 top species based on the biotic score are similar to those based on the total score (figure 4). six species (plain painted lady, blue-eye uintjie, erica-leaf climbers-friend, table mountain copper, peninsula bonnet and cloud disa) tie for the tenth position with regard to biodiversity ranking, instead of the three lower-ranking species (table 5). figure 4: comparison of the biotic and total score for species of special concern on the cape peninsula, south africa. species below the line have a disproportionately higher management score, whereas those above the line have a management score below the average, r2 = 0.85. table 5: top-scoring species of special concern for monitoring in the table mountain national park (and cape peninsula). the strong relationship between the biotic and total scores (figure 4) suggests that the effect of management criteria is not sufficiently large to warrant special attention or any weighting of biodiversity variables. the species elevated into the top 10 over and above their pure biotic score (silvertree, klipspringer and grey rhebuck) are generally agreed to merit special attention. can priority species of special concern be delimited? based on the decision to consider any species for evaluation (i.e. rather than using earlier criteria to predetermine species of special concern), determining which species qualify for monitoring based on their scores remains an issue. a fluctuating cut-off score is not acceptable as a target number of species are required to evaluate monitoring progress, available budget and the desired goal. scores for taxa potentially range between 2 and 42, although the highest recorded was 36 and the lowest 6 (appendix). the highest score for an ‘extralimital’ or ‘extraneous’ species is 16 (at rank 502) for the outeniqua yellowwood, a highly charismatic species that qualifies as alien to the peninsula and an ecosystem-transforming invasive species in this context (richardson et al. 2000). bats spread it from gardens into indigenous forests. (a possible solution is to allow only male plants to be grown in urban areas within 1 km of the tmnp.) the next highest score was 12 (at rank 725). the lowest score for an extinct taxon was 20 (peninsula water sedge at rank 173), 18 for a critically endangered taxon (the common button daisy, at rank 286), 15 for an endangered taxon (for four species of beetles, at rank 591), and 13 for a vulnerable taxon (ochthebius capicola [coleoptera: hydraenidae], at rank 704). the lowest score for an endemic taxon, the cape pepperweed (brassicaceae), was 14 at rank 653. the highest scoring taxon of least concern is the leopard with 27 points (at rank 13; locally extinct), followed by the river blacktip with 26 points (at rank 28; locally endangered owing to its extreme rarity). as for other extinct charismatic taxa, the leopard will not require monitoring unless it is re-introduced onto the peninsula. if an alternative cut-off score is considered for monitoring, 118 red list and endemic taxa would be excluded at a score of 16; 46, 12 and 1 taxa would be exluded at scores 15, 14 and 13, respectively (table 6). we therefore propose that a score of 15 be used as the cut-off for a species of special concern, resulting in 653 taxa that require monitoring under ideal circumstances. by comparison, monitoring only the top 100 taxa would result in 481 red list and endemic taxa not being monitored (table 6), whilst monitoring the top 200 would omit 371 taxa. if the top 300 and top 400 were monitored, 258 and 132 taxa would be omitted, respectively. all these levels would result in a large proportion of red list and endemic species not being monitored and every attempt should be made to obtain funding to monitor the top 650 taxa as a minimum goal. it is not possible to ascertain how many species could feasibly be monitored, as it will depend on both the logistics involved and the intensity of monitoring required. this will be explored in a subsequent paper. table 6: taxa not monitored according to red list status and endemicity, with different cut-off scores and ranks for determining special species on the cape peninsula, south africa. endemics exclude those species with threatened red list status. however, it should be noted that only 543 of 2500 plant taxa on the peninsula (approximately 22%) were scored, and only 171 taxa out of thousands of arthropods (pryke & samways 2008). the frequency distribution of taxa by score suggests that incorporating a score below 18 would result in many additional taxa being included as species of special concern (figure 5). the status of many of these species is ‘near threatened’, ‘data deficient’, or ’least concern: rare’. it could be argued that these should also be monitored, but adequate resources do not exist. figure 5: distribution of scores for taxa evaluated for species of special concern on the cape peninsula, south africa (n = 776), in relation to iucn red list status and endemic status. discussion (back to top) towards a uniform system across taxa it is essential to progress from a prioritisation system that targets specific taxonomic groups to, as we did here, one that considers all taxa in one system. in the past emphasis on, for example, large mammals and birds led to unjustified competition for resources and funds (mace et al. 2008; sitas, baillie & isaac 2009). all taxa, from various ecosystems, need to be evaluated in a comparative system, using a common currency across all groups. this has been achieved by the iucn red listing process., which assesses extinction risk (iucn 2006). a similar goal is strived for here, although the criteria for inclusion and priority are more varied. providing managers with a single list of species for prioritisation and monitoring is likely to be more effective. although the rankings are park-specific, the framework can be used to compare parks and problems between parks. compiling a single, agglomerative listing across different management units (nature reserves or national parks) currently serves little purpose, as budgets are usually park specific. however, such a list could be used for justifying funding allocations for monitoring across park clusters. we do not suggest that all criteria used here should be used across parks, but rather that the approach outlined in this paper should be tested on different parks prior to any widespread adoption. furthermore, the values assigned to species for the variables in tables 1 are dynamic and should regularly be assessed against available information for each species, preferably in an open forum every 5 years. it is also important to keep in mind that the approach outlined here is for the purpose of identifying and prioritising species for monitoring and conservation action and not for assessing their conservation status per se (the latter being achieved using the iucn red listing process). taxa that are extinct in the wild or locally extinct should, wherever feasible, be re-introduced and subsequently become a particular monitoring concern. however, historical sites for extinct species that have not been totally destroyed should be monitored occasionally to ensure that the species has not re-emerged. for example, although mimetes stokoei in the kogelberg did not emerge for three fire cycles (45 years), presumably because the block burning system resulted in low intensity fires, seeds germinated from the seed bank after a high-intensity summer fire in 1999 (rebelo et al., in press). however, the resources allocated to their monitoring will be minimal, although it may vary with potentially greater investment for species with underground seed banks (e.g. erica versus protea), for smaller rather than bigger organisms (e.g. earthworm versus rhino), and for species more recently extinct (e.g. leucadendron grandiflorum (1805) versus isolepis bulbifera [1950]). therefore, extinct and locally extinct taxa should be maintained on the list as a high priority. this all-inclusive approach is slightly different from the approach developed in the savanna of the kruger national park where all red list animals were considered for monitoring (freitag-ronaldson 2006), whilst other species were prioritised for monitoring. however, on the cape peninsula, with over 600 threatened red list and endemic species, it is not feasible to monitor species of special concern using such an approach. problems: representation of lowland species many of the red list taxa on the peninsula are cape flats species that have been heavily impacted by urbanisation. however, only the tokai section of the tmnp occurs on the cape flats, the remainder of the national park being largely mountain habitat. nevertheless, the tokai area of the park is the largest area of this veld type currently conserved and represents the most important conservation contribution to the veld type in the southern half of cape town (rebelo et al. 2011). furthermore, although the tokai section is representative of this veld type, little more remains; that which does remain is in city of cape town nature reserves. this is further complicated by large areas of the tokai area of the park still being under pines. this area thus requires restoration, which depends on fire for the regeneration of fynbos from the seed banks. as the last area is currently scheduled for restoration in 2024, it will not be possible to determine the population sizes and species complement of the tokai section of the park, compared to the original sand fynbos species pool, for many years. how much effort should the tmnp spend in monitoring cape flats species, given the small area of lowland fynbos in the tmnp? obviously, close cooperation with the city of cape town and amateur botanists will be required. ecosystem management in the area should be considered within the context of its unique situation as the final outpost of the cape flats ecosystem, and species should be managed accordingly with due care to population sizes, genetic integrity and symbiont needs. a high-priority, threatened ecosystem approach to its conservation is thus critical until (and indeed after) further species-level information becomes available. conclusion (back to top) a species of special concern monitoring programme has been identified as one of 10 monitoring programmes constituting the sanparks biodiversity monitoring system (see mcgeoch et al. 2011). the challenge for protected areas generally, and in this instance specifically for the tmnp, is to establish a realistic monitoring programme for species of special concern based on an agreed set of variables for establishing conservation and monitoring priority. this is necessary given limited resources, accelerating pressure on species and the enormity of the task at hand, particularly in species-rich regions such as the cape flora (and most specifically the table mountain and agulhas national parks). specific monitoring programmes for such species also need to be robust and enable separation of natural population fluctuations from anthropogenically induced declines (spellerberg 2005). this, along with an assessment of the associated monitoring costs and implications, as well as an outline of theoretical thresholds of potential concern for such species, will be described in a subsequent publication. we have shown that a prioritised species list for monitoring species of special concern can be compiled across diverse taxa, within diverse ecosystems, and that experts can both contribute and agree to the ranking so obtained. although the priority species for monitoring have been identified, this will need to be tempered with the monitoring costs and logistics of implementing such a programme. acknowledgements (back to top) we thank jonathan ball for information on butterflies and chafers, paul brock for stick insects, james pryke for dragonflies and leaf hoppers, and clive turner for water beetles from diverse coleopteran families. we thank participants to the one-day workshop that was held at the kirstenbosch research centre on 15 november 2007. we thank sanbi, especially tilla raimondo and lize von staden for access to detailed information on the red list taxa. we thank pat holmes for useful discussion on topics. references (back to top) adamson, r.s. & salter, t.m., 1950, flora of the cape peninsula, juta, cape town. bakker, v.j. & doak, d.f., 2009, ‘population viability management: ecological standards to guide adaptive managment for rare species’, frontiers in ecology and the environment 7, 158−165. doi:10.1890/070220 bengtsson, j., jones, h. & setala, h., 1997, ‘the value of biodiversity’, trends in ecology and evolution 12, 334−336. doi:10.1016/s0169-5347(97)01135-x biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59−80, island press, washington dc. bond, w.j., maze, k. & desmet, p., 1995, ‘fire life histories and the seeds of chaos’, ecoscience 2, 252–60. brooks, t.m., mittermeier, r.a., mittermeier, c.g., da fonseca, g.a.b., rylands, a.b., konstant, w.r., et al., 2002, ‘habitat loss and extinction in the hotspots of biodiversity’, conservation biology 16, 909−923. doi:10.1046/j.1523-1739.2002.00530.x bubb, p.j., butchart, s.h.m., collen, b., dublin, h., kapos, v., pollock, c., et al., 2009, iucn red list index: guidance for national and regional use, iucn, gland. butchart, s.h.m., stattersfield, a.j., baillie, j., bennun, l.a., stuart, s.n., akcakaya, h.r., et al., 2005, ‘using red list indices to measure progress towards the 2010 target and beyond’, philosophical transactions of the royal society b 360, 255−268. doi:10.1098/rstb.2004.1583, pmid:15814344, pmcid:1569445 de grammont, p.c. & cuaron, a.d., 2006, ‘an evaluation of threatened species categorization systems used on the american continent’, conservation biology 20, 14−27. doi:10.1111/j.1523-1739.2006.00352.x, pmid:16909655 foxcroft, l.c., 2009, ‘developing thresholds of potential concern for invasive alien species: hypotheses and concepts’, koedoe 50(1), art. #157, 6 pages. doi:10.4102/koedoe.v51i1.157 freitag, s. & van jaarsveld, a.s., 1997, ‘relative occupancy, endemism, taxonomic distinctiveness and vulnerability: prioritizing regional conservation actions’, biodiversity and conservation 6, 211−232. doi:10.1023/a:1018392019594 freitag-ronaldson, s., 2006, ‘sanparks framework for species of conservation concern’, unpublished internal report, south african national parks. gardenfors, u., hilton-taylor, c., mace, g.m. & rodriguez, j.p., 2001, ‘the application of iucn red list criteria at regional levels’, conservation biology 15, 1206−1212. doi:10.1111/j.1523-1739.2001.00112.x, doi:10.1046/j.1523-1739.2001.00112.x gaston, k.j., jackson, s.f., cantu-salazar, l. & cruz-pinon, g., 2008, ‘the ecological performance of protected areas’, annual review of ecology evolution and systematics 39, 93−119. doi:10.1146/annurev.ecolsys.39.110707.173529 given, d.r. & norton, d.a., 1993, ‘a multivariate approach to assessing threat and for priority setting in threatened species conservation’, biological conservation 64, 57−66. doi:10.1016/0006-3207(93)90383-c goldblatt, p. & manning, j.c., 2000, ‘cape plants: a conspectus of the cape flora of south africa’, strelitzia 9. haaksma, e.d. & linder, h.p., 2000, restios of fynbos, botanical society of south africa, cape town. hall, a.v. & ashton, e.r., 1983, threatened plants of the cape peninsula, threatened plants research group, cape town. hansen, a.j., rotella, j.j., kraska, m.p.v. & brown, d., 1999, ‘dynamic habitat and population analysis: an approach to resolve the biodiversity manager’s dilemma’, ecological applications 9, 1459−1476. doi:10.2307/2641410 doi:10.1890/1051-0761(1999)009[1459:dhapaa]2.0.co;2 international union for conservation of nature, 2006, guidelines for using the iucn red list categories and criteria, prepared by the standards and petitions working group of the iucn ssc biodiversity assessments sub-committee in december 2006. iucn. see international union for conservation of nature. joseph, l.n., maloney, r.f. & possingham, h.p., 2009, ‘optimal allocation of resources among threatened species: a project prioritization protocol’, conservation biology 23, 328−338. doi:10.1111/j.1523-1739.2008.01124.x, pmid:19183202 mace, g.m, possingham, h.p. & leader-williams, n., 2006, prioritizing choices in conservation, in macdonald, d.w. & service, k. (eds.), key topics in conservation biology, pp. 17–34, blackwell, oxford, available from , http://www.kent.ac.uk/anthropology/dice/resources/nlw-macdonaldch2.pdf mace, g.m., collar, n.j., gaston, k.j., hilton-taylor, c., akcakayaz, h.r., leader-williams, n., et al., 2008, ‘quantification of extinction risk: iucn’s system for classifying threatened species’, conservation biology 22, 1424−1442. doi:10.1111/j.1523-1739.2008.01044.x, pmid:18847444 martin, j., runge, m.c., nichols, j.d., lubow, b.c. & kendall, w.l., 2009, ‘structured decision making as a conceptual framework to identify thresholds for conservation and management’, ecological applications 19, 1079−1090. doi:10.1890/08-0255.1, pmid:19688917 mcgeoch, m.a., dopolo, m., novellie, p., hendriks, h., freitag, s., ferreira, s., et al., 2011, ‘a strategic framework for biodiversity monitoring in south african national parks’, koedoe 53(2), art. #991, 10 pages. doi:10.4102/koedoe.v53i2.991 miller, r.m., rodríguez, j.p., aniskowicz-fowler, t., bambaradeniya, c., boles, r., eaton, m.a., et al., 2007, ‘national threatened species listing based on iucn criteria and regional guidelines: current status and future perspectives’, conservation biology 21, 684−696. doi:10.1111/j.1523-1739.2007.00656.x, pmid:17531047 moll, e. & scott, l., 1981, trees and shrubs of the cape peninsula, ecolab, cape town. oliver, i. & oliver, t., 2000, ericas of the cape peninsula, protea atlas project, cape town. pfab, m.f., 2002, ‘priority ranking scheme for red data plants in gauteng, south africa’, south african journal of botany 68, 299−303. privett, s.d.j., cowling, r.m. & taylor, h.c., 2001, ‘thirty years of change in the fynbos vegetation of the cape of good hope nature reserve, south africa’, bothalia 31, 99−115. pryke, j.s. & samways, m.j., 2008, ‘conservation of invertebrate biodiversity on a mountain in a global biodiversity hotspot, cape floral region’, biodiversity and conservation 17, 3027−3043. doi:10.1007/s10531-008-9414-4 raimondo, d., von staden, l., foden, w., victor, j.e., helme, n.a., turner, r.c., et al., 2009, ‘red list of south african plants’, strelitzia 25. rebelo, t., 2000, proteas of the cape peninsula, protea atlas project, cape town. rebelo, a.g., boucher, c., helme, n., mucina, l., rutherford m.c., et al., 2006, ‘fynbos biome’, in l. mucina & m.c. rutherford (eds.), the vegetation of south africa, lesotho and swaziland, pp. 52–219, strelitzia 19. rebelo, a.g., holmes, p.m., dorse, c. & wood, j., 2011, ‘cape town: averting a biodiversity megadisaster?’, south african journal of botany 77, 20-35. rebelo, a.g., helme, n., holmes, p.m., forshaw, c.n, von staden, l. et al., in press, ‘african proteaceae red data list’, strelitzia. regan, h.m., hierl, l.a., franklin, j., deutschman, d.h., schmalbach, h.l., winchell, c.s., et al., 2008, ‘species prioritization for monitoring and management in regional multiple species conservation plans’, diversity and distributions 14, 462−471. doi:10.1111/j.1472-4642.2007.00447.x richardson, d.m., pyšek, p., rejmánek, m., barbour, m.g., panetta, f.d. & west, c.j., 2000, ‘naturalization and invasion of alien plants: concepts and definitions’, diversity and distributions 6, 93−107. doi:10.1046/j.1472-4642.2000.00083.x richardson, d.m. & van wilgen, b.w., 2004, ‘invasive alien plants in south africa: how well do we understand the ecological impacts?’, south african journal of science 100, 45−52. roux, j.p., 1979, cape peninsula ferns, national botanical gardens, kirstenbosch. sanparks. see south african nantional parks. south african national parks., 2008, table mountain national park: park management plan, sanparks, pretoria. south african national parks., 2009, tokai and cecilia management framework 2005-2025, table mountain national park, cape town. schnittler, m. & günther, k.f., 1999, ‘central european vascular plants requiring priority conservation measures — an analysis from national red lists and distribution maps’, biodiversity and conservation 8, 891−925. doi:10.1023/a:1008828704456 sitas, n., baillie, j.e.m. & isaac, n.j.b., 2009, ‘what are we saving? developing standardized approach for conservation action’, animal conservation 12, 231−237. doi:10.1111/j.1469-1795.2009.00244.x spellerberg, i.f., 2005, monitoring ecological change, cambridge university press, cambridge. doi:10.1017/cbo9780511614699 thuiller, w., slingsby, j.a., privett, d.j. & cowling, r.m., 2007, ‘stochastic species turnover and stable coexistence in a species-rich, fire-prone plant community’, plos one 9, 1−8. trinder-smith, t., 2006, wild flowers of the table mountain national park, botanical society of south africa, cape town. turpie, j.k., 2003, ‘the existence value of biodiversity in south africa: how interest, experience, knowledge, income and perceived level of threat influence local willingness to pay’, ecological economics 46, 199−216. doi:10.1016/s0921-8009(03)00122-8 vellak, a., tuvi, e., reier, u., kalamees, r., roosaluste, e., zobel, m., et al., 2009, ‘past and present effectiveness of protected areas for conservation of naturally and anthropogenically rare plant species’, conservation biology 23, 750−757. doi:10.1111/j.1523-1739.2008.01127.x pmid:19128324 younge-hayes, a., rebelo, t. & cheney, c., 2007, ‘workshop on monitoring priorities for rare and endangered species on the cape peninsula’, kirstenbosch research centre, tmnp, cape town, 5 november. appendix 1 (back to top) online appendix table 1: list of species of specialconcern in the table mountain national park and associated scores. refer to online appendix http://koedoe.co.za/index.php/koedoe/article/view/1019/1255 abstract introduction research method and design results discussion conclusion acknowledgements references about the author(s) guy hausler veterinary wildlife services, south african national parks, skukuza, south africa kerry slater department of environmental sciences, university of south africa, south africa citation hausler, g. & slater, k., 2016, ‘the composition of mixed-species foraging flocks of birds in kruger national park, south africa’, koedoe 58(1), a1314. http://dx.doi.org/10.4102/koedoe.v58i1.1314 short communication the composition of mixed-species foraging flocks of birds in kruger national park, south africa guy hausler, kerry slater received: 01 apr. 2015; accepted: 06 jan. 2016; published: 09 june 2016 copyright: © 2016. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract mixed-species foraging flocks (msffs) of birds can be defined as aggregations of more than two species that actively initiate and continue their association while foraging, without being drawn to a single resource. msffs have been well documented for terrestrial habitats globally, but rarely in southern africa. this study describes the composition of msffs in two habitat types (acacia and combretum) within the southern kruger national park, south africa during the late dry season. thirty-one msffs were recorded in each of the two habitat types, with 1251 individuals of 74 different species being observed. we found that compared to combretum, (mean: 10.7 ± 5.2 s.d.) acacia had significantly more individuals per msffs (mean: 21.5 ± 12.6 s.d.) and more species per msff (acacia mean: 8.7 ± 3.5 s.d.; combretum mean: 5.9 ± 1.7 s.d.). the mean number of individuals per species per 31 msffs was 9.3 (± 4.5 s.d.) and 7.6 (± 5.6 s.d.) in the acacia and combretum habitat types respectively. the most frequently occurring species in both habitat types was the fork-tailed drongo (dicrurus adsimilis). there was a significant association between certain species pairs in both habitats. future studies in this area could be done to investigate the reasons behind the differences in msff sizes and species numbers between habitats. the season during which this study was performed excluded all summer migrants and a similar investigation in the wet season may reveal a different msff composition. conservation implications: understanding the dynamics and compositions of msffs, could form a valuable component of avian biodiversity monitoring both in and outside of protected areas. within a given area, changes in the composition and behaviour of msffs over time could potentially be used as early indicator of threats to biodiversity. introduction mixed-species foraging flocks (msffs) of birds can be defined as aggregations of more than two species that actively initiate and continue their association (harrison & whitehouse 2011) while foraging in an area or habitat (greig-smith 1978). despite the global prevalence of msffs among terrestrial bird communities (harrison & whitehouse 2011; morse 1970) msffs have rarely been characterised in southern africa. the presence and role of each species in a flock reflects its ecological niche as well as the various factors leading to flock formation (morse 1970). in temperate areas, mssfs tend to form more readily in non-breeding seasons (morse 1970) and differ from other aggregations of birds in their cohesion, rather than being a gathering of different species drawn to a particular resource (such as a fruiting tree); these flocks remain together as they progress through the habitat (goodale & beauchamp 2010). only those individuals that lead the flock or make a concerted effort to follow and remain in the flock should be considered as msff members (greig-smith 1978). msffs are typically led by a particular species, a role typically fulfilled by a so-called ‘nucleus species’. a nucleus species is usually a species with high flocking propensity, encountered in a high percentage of msffs, often occurs in intra-specific flocks, and usually displays conspicuous behaviour such as vocalising or making conspicuous movements (hutto 1994). the individuals of a nucleus species are wholly or partially responsible for the formation and continued cohesion of the msff (goodale & beauchamp 2010; greig-smith 1978; morse 1970) with other species in the msff organising their various activities around the nucleus species. although various feeding guilds are present, msffs are usually dominated by insectivorous species (croxall 1976; greig-smth 1978). in some habitats multiple types of msffs may be distinguished, based on differences in foraging height (zou et al. 2011) or the presence of particular species or guilds (king & rappole 2001). although some authors have suggested that the regular occurrence of a given species in msffs is directly related to the abundance of the species (morse 1978; zou et al. 2011), this is not always the case in savanna ecosystems (greig-smith 1978). furthermore, some species may be recorded in msffs only when the flock happens to move through the territory of that species (greig-smith 1978). the mean number of species per msff ranges from 3.8 in forested areas in china (zou et al. 2011) to 7.7 and 9.9 in the former eastern provinces and barotse of northern rhodesia respectively (winterbottom 1943). furthermore, although differences between msffs observed in different vegetation types within one geographical area may be minimal (croxall 1976; winterbottom 1943), msffs in adjacent acacia and mixed woodland habitats in northern south africa do show significant differences in composition and mean number of species (thomson & ferguson 2007). to date, msffs occurring in any of the various savanna habitats of the kruger national park (knp), south africa have not been investigated despite the major role that birds play in savanna ecosystems and the maintenance of savanna heterogeneity (kemp et al. 2003). the main aim of this study was to investigate the composition of msffs in a southern section of knp. research method and design study area the study area comprises the sabie/crocodile thorn thickets landscape type (gertenbach 1983) located in the southern knp. this entire landscape is underlain by granites (venter, scholes & eckhardt 2003) with crests and mid-slopes supporting broad-leaved combretum bushveld, whereas foot-slopes are dominated by fine-leaved tree species (particularly acacia spp.). the foot-slopes display two principle forms of woodland: one has a dense, tangled understory, while the other, found in brackish (sodic) areas, has minimal understory vegetation and large bare patches of soil, typically beneath acacia grandicornuta (horned-thorn) trees (gertenbach 1983). the sites surveyed for msffs in this study fall within this landscape type and lie in the area bound by the s65 gravel road in the west and nkuhlu picnic spot in the east (figure 1). for the purpose of this study the following habitats were investigated: combretum-dominated bushveld on crests and acacia-dominated woodland in the lowlands, hereafter referred to as combretum and acacia respectively. the study area falls within a summer rainfall region with an average of between 511 mm and 566 mm of rain annually. the average maximum and minimum daily temperatures are 32.6 °c and 20.6 °c in summer (november – february), dropping to 25.9 °c and 5.6 °c in winter (may – august) respectively (venter et al. 2003). figure 1: location of study area, southern kruger national park, showing landscapes. procedure surveys were conducted on 16 days during the late dry season (july – early september 2013) and restricted to 06:00 – 10:00 and 16:00 – 17:00 each sampling day to coincide with the greatest levels of bird activity (pers. obs.). data were collected by walking or driving through suitable areas of the landscape until a msff was encountered. the msff was then observed from a fairly fixed point for as long as the msff remained cohesive and/or as long as new species continued to join the msff (typically > 12 min). recording of data stopped when the msff dispersed or if visual contact of the msff was lost. to ensure that no msff was sampled more than once we tried to avoid sampling the same area multiple times. however, some areas were sampled on more than one occasion, but this was only done after an interval of more than 10 days to minimise repetitive sampling of particular msffs. unfortunately wider sampling within the study area was not possible because of restrictions on human movement enforced by park management as a result of anti-poaching operations at the time of the study. both acacia and combretum were surveyed on each sampling day to eliminate as far as possible, effects of weather conditions. the slope position (i.e. crest, foot-slope, etc.), dominant woody vegetation, and a knp vegetation map (derived from venter 1990) were used for classifying sites. site specific information recorded for each msff included vegetation type, gps position, date, time, cloud cover, and any interesting msff behaviour. observations were made with the aid of a pair of zeiss 10x40 binoculars and a hand-held electronic range finder was used for the estimation of 20-m distances. the species within each msff were identified as per hockey, dean and ryan (2005) and recorded along with the numbers of individuals of each species. species foraging in all vegetation strata as well as on the ground were recorded as long as they showed clear tendencies to join and follow the msff. species using a sit-and-wait hunting technique, which made no effort to follow the msff as it traversed their position, were ignored, as were aerial species such as swifts (apodidae). only bird species observed within a 20-meter radius of the core of the msff’s activities (such as a particular tree) were recorded, as the dense nature of some of the areas surveyed prevented observations over a wider area. we calculated the number of individuals per species across all msff’s and the frequency of occurrence of each species in msffs. we then calculated the following: the mean number of individuals per msff, the number of species per msff, and the number of individuals per species per msff. a wilcoxon rank sum test was used to investigate whether there were any significant differences in the number of individuals and species per msff between the two habitat types. associations between species in msffs were examined by scoring each species as either present (1) or absent (0) in each msff. this presence–absence data was then analysed using a pearson’s correlation to give a measure of association between two species. results during the study period, 62 msffs were encountered, equally divided between the two habitat types (i.e. 31 msffs in acacia and 31 msffs in combretum). in total 1251 individuals of 74 different species were observed in these msffs. of these 74 species, 29 species in acacia and 24 species in combretum were regarded as regular flocking species (species that occurred in > 10% of flocks; king & rappole 2001). only these species were used for further analysis (table 1). of the 32 (43%) species that were regarded as regular flocking species, 21 species (28%) were recorded in both the acacia and the combretum, 8 (11%) in the acacia only, and 3 (4%) in the combretum only. table 1: bird species observed in more than 10% of mixed-species foraging flocks in acacia (n = 31 flocks) and combretum (n = 31 flocks) habitat types in kruger national park, south africa. across flocks the mean number of individuals per species was 9.3 (± 4.5 s.d.) and 7.6 (± 5.6 s.d.) in the acacia and combretum habitat types respectively. results from the wilcoxon rank sum test indicated that msffs in acacia had significantly (p < 0.0001, n = 31, z = 4.2) more individuals per msffs (mean: 21.5 ± 12.6 s.d.) than those in combretum (mean: 10.7 ± 5.2 s.d.). a significant difference (t = 3.91; p = 0.0003) in the number of species per msff between the acacia and combretum was also found, with the acacia having a mean number of species per msff of 8.7 (± 3.5 s.d.) and the combretum 5.9 (± 1.7 s.d.). the frequency of occurrence of a recorded species in the msffs often varied according to habitat type. the three most frequently occurring species in acacia msffs were fork-tailed drongo (dicrurus adsimilis), southern black tit (melaniparus viger), and chinspot batis (batis molitor); whereas in combretum they were fork-tailed drongo, rattling cisticola (cisticola chiniana), and chinspot batis (table 1). there were 26 significant correlations among species pairs in the acacia habitat of which 20 were positive (table 2). of these, the strongest associations are between southern yellow-billed hornbills (tockus leucomelas), fiscal flycatchers (melaenornis silens), and southern red-billed hornbills (tockus rufirostris); black-backed puffbacks (dryoscopus cubla) and tawny-flanked prinias (prinia subflava); southern grey-headed sparrows (passer diffuses) with fiscal flycatchers; and southern masked weavers (ploceus velatus), with arrow-marked babblers (turdoides jardineii). table 2: correlations among bird species observed in more than 10% of mixed-species foraging flocks in the acacia (n = 31 flocks) study site in kruger national park, south africa. in the combretum habitat there were nine significant correlations among species pairs of which five were positive (table 3). the following positive correlations were found in the combretum habitat: long-billed crombecs (sylvietta rufescens) with rattling cisticola, emerald-spotted wood doves (turtur chalcospilos) and blue waxbills (uraeginthus angolensis) with arrow-marked babblers, southern grey-headed sparrows with yellow-fronted canaries (crithagra mozambica), green wood hoopoes (phoeniculus purpureus) with black-backed puffbacks. negative correlations in the combretum habitat included those between chinspot batis’ and arrow-marked babblers; red-billed firefinches (lagonosticta senegala) with arrow-marked babblers; greater blue-eared starlings (lamprotornis chalybaeus) with fork-tailed drongos; and cardinal woodpeckers (dendropicos fuscescens) with rattling cisticolas. table 3: correlations among bird species observed in more than 10% of mixed-species foraging flocks in the combretum (n = 31 flocks) study site in kruger national park, south africa. discussion the msffs observed during this study had mean msff sizes similar to the mean msff size (16.9–19.3) recorded in tropical forests in myanmar, southeast asia (croxall 1976; king & rappole 2001) and west african savannas (23.2) (greig-smith 1978). the mean number of species per msff was comparable to msffs studied in other african savannas (winterbottom 1943). acacia supported larger flocks and a greater number of species per flock when compared to the combretum. msffs in adjacent acacia and mixed woodland habitats in northern south africa also showed significant differences in composition and mean number of species (thomson & ferguson 2007). acacia also exhibited a higher mean number of species per msff, possibly because of having greater structural complexity than combretum. species diversity typically increases with structural complexity of habitat (skwono & bond 2003). testing this hypothesis is however beyond the scope of this study, and reasons for these differences in msff sizes and species numbers between habitats would make for interesting future study. having said this, the majority of species recorded in msffs in this study were recorded in both habitat types surveyed. in some cases, inconspicuous msff participants such as stierling’s wren-warbler (calamonastes stierlingi) may have been under-recorded because of the dense nature of the habitats in which they occurred. this would lead to underrepresentation of these species in the results. although such species could have been actively revealed by flushing them, this would have led to msff dispersal and was thus deemed unsuitable. in both vegetation types, fork-tailed drongos occurred in 71% of msffs, a similar finding to that of many other studies where drongos were recorded as common msff participants (croxall 1976; greig-smith 1978; swynnerton 1915; winterbottom 1943; zou et al. 2011). in the msffs recorded in nyslvley nature reserve, south africa drongos however occurred in up to 92% of msffs (thompson & ferguson 2007). drongos have been recorded as a nucleus species of msffs in various studies (cordeiro et al. 2015; goodale & kotagama 2006; nefdt 1989), and because of their prevalence in msffs in this study, drongos are therefore a very likely nucleus species in msff of the southern knp. other studies have however found that despite drongos occurring in msff they do not always take on the role of a nucleus species within a msff (greig-smith 1978). of the other frequently encountered species, the chinspot batis and southern black tit are notable because both batises (batis spp.) and tits (parus spp.) are frequent participants of msffs in savannas and forests elsewhere in africa (greig-smith 1978; nefdt 1989; thomson & ferguson 2007; winterbottom 1943). in both acacia and combretum there was a clear tendency of certain species to be present in the majority of msffs. the chinspot batis, southern black tit, and fork-tailed drongo are frequent participants in both habitat types. the regular occurrence of pairs of fiscal flycatchers in msffs in the acacia habitat is noteworthy as this species is an uncommon winter visitor to the knp (newman 1991) and this vegetation type probably represents a favoured habitat for these flycatchers when they occur temporarily in this region. the season during which this study was conducted excluded all summer migrants and a similar investigation in the wet season may therefore reveal a different msff composition. the non-random patterns of positive associations among observed msffs in this study support findings by king and rappole (2001) that members of some species tend to associate with other species. king and rappole (2001) suggested that the reasons for these positive associations between species in msffs was to facilitate more efficient foraging by flushing prey from the vegetation through which they move. the reasons for the positive association between msff members in this study are likely to be similar, but need further investigation to verify this. conclusion this study is the first of its kind to be undertaken in the knp and forms the baseline for ongoing work investigating the composition of msffs occurring in the broader landscapes of this region. the analysis of msffs could form a valuable component of the biodiversity monitoring currently being undertaken in the knp, and any future change in the composition and behaviour of msffs may be an early indicator of threats to the biodiversity of this region. acknowledgements we would like to thank south african national parks and in particular dr c.c. grant for her assistance and specialist advice during the course of this project. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions g.h. was responsible for project design, data collection, and initial data analysis. k.s. was the project supervisor, statistically analysed data for final manuscript, and critically reviewed the final article. references cordeiro, n.j, borghesio, l., joho, m.p., monoski, t.j., mkongewa, v.j. & dampf, c.j., 2015, ‘forest fragmentation in an african biodiversity hotspot impacts mixed-species bird flocks’, biological conservation 188, 61–71. croxall, j.p., 1976, ‘the composition and behaviour of some mixed-species bird flocks in sarawak’, ibis 118(3), 333–346. gertenbach, w.p.d., 1983, ‘landscapes of the kruger national park’, koedoe 26, 9–121. goodale, e. & beauchamp, g., 2010, ‘the relationships between leadership and gregariousness in mixed-species bird flocks’, journal of avian biology 41, 99–103. goodale, e. & kotagama, s.w., 2006, ‘vocal mimicry by a passerine bird attracts other species involved in mixed-species flocks’, animal behaviour 72, 471–477. greig-smith, p.w., 1978, ‘the formation, structure and function of mixed-species insectivorous bird flocks in west african savanna woodland’, ibis 120(3), 284–297. harrison, n.m. & whitehouse, m.j, 2011, ‘mixed-species flocks: an example of niche construction?’, animal behaviour 81, 675–682. hockey, p.a.r., dean, w.r.j. & ryan, p.g. (eds.), 2005, robert’s birds of southern africa, 7th edn., the trustees of the john voelcker bird book fund, cape town. hutto, r.l., 1994, ‘the composition and social organization of mixed-species flocks in a tropical deciduous forest in western mexico’, condor 96, 105–118. kemp, a.c., dean, w.j.r., whyle, i.j., milton, s.j. & benson, p.c., 2003, ‘birds: responders and contributions to ecological heterogeneity’, in j. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience – ecology and management of savanna heterogeneity, pp. 276–291, island press, washington, dc. king, d.i. & rappole, j.h., 2001, ‘mixed-species bird flocks in dipterocarp forest of north-central burma (myanmar)’, ibis 143(4), 380–390. morse, d.h., 1970, ‘ecological aspects of some mixed species foraging flocks of birds’, ecological monographs 40(1), 119–168. morse, d.h., 1978, ‘structure and foraging patterns of tits and associated species in english woodland during the winter’, ibis 120(3), 298–312. nefdt, r.j.c., 1989, ‘a study of some bird parties in zambia’, ostrich 14, 27–32. newman, k., 1991, birds of the kruger national park, rev. edn., southern book publishers, halfway house. skwono, a.l. & bond, w.j., 2003, ‘bird community composition in an actively managed savanna reserve, importance of vegetation structure and vegetation composition’, biodiversity and conservation 12, 2279–2294. swynnerton, c.f.m., 1915, ‘mixed bird parties’, ibis 57(2), 346–354. thomson, r.l. & ferguson, j.w.h., 2007, ‘composition and foraging behaviour of mixed species flocks in two adjacent woodland habitats: a special and temporal perspective’, ostrich 78(1), 65–73. venter, f.j., 1990, ‘a classification of land for management planning in the kruger national park’, phd thesis, university of south africa, pretoria. venter, f.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation pattern’, in j. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience – ecology and management of savanna heterogeneity, pp. 83–129, island press, washington, dc. winterbottom, j.m., 1943, ‘on woodland bird parties in northern rhodesia’, ibis 85(4), 437–442. zou, f., chen, g., yang, q. & fellows, j.r., 2011, ‘composition of mixed-species flocks and shifts in foraging location of flocking species on hainan island, china’, ibis 153(2), 269–278. vol. 50 no. 1 pp. 18 21koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research jozua j. viljoen 1department of nature conservation tshwane university of technology south africa 2mammal research institute department of zoology and entomology university of pretoria south africa andre ganswindt section of veterinary wildlife studies faculty of veterinary science university of pretoria south africa german primate centre department of reproductive biology germany rupert palme institute of biochemistry department of natural sciences university of veterinary medicine austria hendrik c. reynecke1 johan t. du toit 2 department of wildland resources utah state university usa william r. langbauer jr pittsburgh zoo & ppg aquarium usa correspondence to: jozua j. viljoen e-mail: viljoenjj@tut.ac.za postal address: tshwane university of technology, private bag x680, pretoria, 0001, south africa abstract during the past several years, non-invasive monitoring of steroid metabolites in faeces of elephants has become an increasingly popular technique to generate more information about the causal relationship between hormones and behaviour in both living elephant species. this is important knowledge which can be used to optimise local conservation and wildlife management by finding new strategies for better elephant population management and control. in this context, however, information about an actual involvement of the hypothalamic-pituitary-adrenal axis during assumable stressful events is still limited, especially for wildlife populations. one difficulty in discovering such information is often the lack of reliable data for hormone baseline levels. therefore, the aim of this study was to determine baseline concentrations of faecal glucocorticoid metabolites that could be expected within age classes and between seasons in african elephants (loxodonta africana) in the kruger national park (knp). a total of 374 faecal samples were collected from randomly located family herds in the southern knp between may 2002 and august 2005. the samples were analysed for immunoreactive concentrations of faecal glucocorticoid metabolites using a validated enzyme immunoassay for 3α,11oxo-cortisol metabolites (3α,11oxo-cm). all samples were grouped according to the estimated age class of the subject using a field method based on bolus diameter, and regarding the ecological season collected. no significant differences in faecal 3α,11oxo-cm concentrations were found across age classes (h 3 = 7.54; p = 0.057), but the mean 3α,11oxo-cm concentration of samples collected in the dry season (n = 196) was significantly higher than in the wet season (n = 178) (u = 15206.50; p = 0.032), which indicates a possible physiological stress situation due to a decline in food quantity and quality. the information generated in this study represents a reliable data set for baseline concentrations of faecal glucocorticoid metabolites for elephants within the knp and can be used to measure the stressrelated effects of translocations, management actions and the impact of chosen land use activities. keywords: glucocorticoid metabolites, faecal, baseline, african elephant, enzyme immunoassay measurement of concentrations of faecal glucocorticoid metabolites in free-ranging african elephants within the kruger national park 18 2008 faecal glucocorticoid metabolites in african elephants original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 18 21 a free-ranging animal is in a state of stress if it is required to make abnormal or extreme adjustments in its physiology or behaviour in order to cope with adverse aspects of its environment or management (friend 1980). from the physiological perspective, several endocrine responses are, amongst others, involved during stress situations. the frontline hormones to overcome these situations are the glucocorticoids, which are indicators of adrenal activity and thus of stress (möstl & palme 2002). as a physiological mechanism, stress is not inherently negative (moberg 2000), however, prolonged high concentrations of glucocorticoids may decrease individual fitness (munck et al. 1984) and reproductive success (liptrap 1993), which may cascade into long-term behavioural changes. the assessment of adrenal endocrine function in vertebrates via blood samples was an often used tool in the past, but the procedure in itself can elicit a glucocorticoid response (möstl & palme 2002). in order to avoid such self-induced stress responses, the use of non-invasive methodologies for endocrine assessment is necessary. the collection of urine or faeces as basic material for endocrine analysis seems to be a reliable alternative to blood hormone measurements, which is demonstrated by the large number of established non-invasive methodologies for monitoring adrenal function in a variety of different mammalian species (goymann et al. 1999; wasser et al. 2000, sands & creel 2004), including elephants (brown et al. 1995, ganswindt et al. 2003). brown et al. (1995) successfully developed and validated a method to assess adrenal function non-invasively through the analysis of cortisol in urine from captive elephants, but due to the practical difficulties involved with collecting urine samples in areas with absorptive ground, this method would not be an option for regular sampling from elephants in the wild. faecal samples offer the advantage that they can easily be collected, and a few non-invasive methods have already been developed and successfully tested to enable the measurement of glucocorticoid metabolites in elephant faeces (stead et al. 2000, wasser et al. 2000, foley et al. 2001, ganswindt et al. 2003; 2005). in this respect, it could be shown that a pharmacologically induced physiological stress response, induced via administration of adrenocorticotropic hormone (acth), could be monitored in captive african elephants. although work on free-ranging african elephants has been limited, ganswindt et al. (2005) found that elevated androgen levels in free-ranging male african elephants could not be related to the age of the individual. seasonal effects however do appear to influence concentrations of faecal glucocorticoid metabolites, as foley et al. (2001) found the highest concentrations during the dry season. however, a regular sample collection at a certain time before and after a predictable stressful event is necessary to create a reliable baseline for comparison, which already underlines the importance of known baseline levels to reliably determine if any specific situation is more or less stressful than the norm. we suggest, for example, a stressful situation can be triggered in elephants by several stimuli, including environmental factors (food quality and quantity): behavioural (courtship and mating); and psychosocial causes (translocation, culling). more information on the endocrine regulation of potential stressrelated events is needed, as the effect of human and natural disturbances on elephant populations has been restricted to behavioural indicators (whyte 2003). it is still not known what constitutes a naturally occurring stressful situation for freeranging elephants, or which management actions or land use practices will result in elevated stress indicators. in this paper we provide a quantitative baseline measure of concentrations of faecal glucocorticoid metabolites, after taking age and seasonal effects into account, in order to start establishing baseline levels for elephants in the kruger national park (knp). materials and methods study area and population the study area is located in the knp and falls within the lebombo arid mountain bushveld, sweet lowveld bushveld and mixed lowveld bushveld vegetation types (low & rebelo 1996). the knp covers an area of approximately 19 000 km² and can be longitudinally divided into resistant granites in the west, succeeded by ecca shales, basalt and rhyolites in the east, that give rise to different soil types and the associated flora and fauna (venter et al. 2003). data collection was restricted to the southern knp and extended from the sabie and crocodile river thickets, sclerocarya birrea subsp. caffra/acacia nigrescens savanna, mixed combretum to the lebombo south landscapes. sampling a matriarch of a family herd (median size 12) was fitted with a vhf radio collar and this group was followed on foot so as not to bias sampling close to roads. samples were collected as soon as possible after an individual had defecated, from individuals in the collared group as well as from family herds (median size 13) located within the study area. we tried to exclude individual bias by collecting samples throughout each feeding patch used by a family herd. when there were obvious signs that a herd had detected our presence, such as a definite orientation towards or away from us, no samples were collected, in order to avoid the collection of stress-induced samples. all samples were collected between 07:00 and 11:00 am. rubber gloves were used to collect approximately 100 grams of faecal material that was then placed in a labelled plastic bag. the time lapsed between defecation and the freezing of a sample was standardised to a maximum of two hours. each sample was marked with the date of collection, a gps coordinate, and the estimated age class of the subject using a field method based on the bolus diameter as described by wimberger (2001). faecal samples were collected over the four ecological seasons as classified by zambatis (2002), namely early dry (may–july); late dry (aug–oct), early wet (nov–jan) and late wet (feb–april). this allowed us to determine, through concentrations of glucocorticoid metabolites, whether environmental factors, which change between seasons, have any possible influence on physiological stress. samples (n = 374) were collected between may 2002 and august 2005. the average annual rainfall during the study was 572 mm, which is in the known average annual rainfall range of 500–700 mm for this region, and therefore would not have unduly influenced environmentally induced feeding stress (venter et al. 2003). faecal extraction and hormone assays faecal samples were extracted according to the procedure described by merl et al. (2000). in brief, 0.5 g faeces plus 1 ml water and 4 ml methanol were vortexed for 30 minutes and centrifuged at 2000 g for 15 minutes. a quantity of 1 ml of the supernatant was mixed with 5 ml ethyl ether and 0.25 µl of a 5% nahco 3 solution, and centrifuged at 2000 g for 15 min. the aqueous phase was frozen at -20 °c overnight and then the ether was decanted and dried down under a stream of n 2 . following evaporation, the samples were reconstituted in an assay buffer and taken to assay. faecal extracts were measured for immunoreactive glucocorticoid metabolites using an enzyme immunoassay for 3α,11oxo-cortisol metabolites (3α,11oxo-cm) (möstl et al. 2002), which has previously been shown to provide reliable information on adrenocortical function in the african elephant (ganswindt et al. 2003; 2005). briefly, 50 µl aliquots of standards, quality controls, and diluted faecal extracts were pipetted in duplicate into microtiterplate wells. a total of 100 µl of biotinylated label and antiserum (raised in a rabbit against 5βandrostane-3α-ol-11-one-17-cmo) were added and the plates incubated overnight at 4 °c. following incubation, the plates 19 original research viljoen, ganswindt, palme, reynecke, du toit & langbauer jr koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 18 21 were washed four times and 250 µl (4.2 mu) of streptavidin horseradish peroxidase conjugate added to each well. following incubation in the dark for 45 min at 4 °c, plates were washed again, before 250 µl (69.4 nmol) tetramethylbenzidine was added and plates further incubated (45 min; 4 °c). the reaction was terminated by adding 50 µl of 2 m h 2 so 4 and the absorbance measured at 450 nm (reference filter: 620 nm) with an automated plate reader. sensitivities of the assays at 90 % binding were 3.0 pg/well and intraand interassay coefficients of variation, determined by repeated measurements of high and low value quality controls ranged between 3.0 % and 12.5 %, respectively. statistical analysis the age class data, as well as the data from the ecological season, was tested for normality using the shapiro-wilks w test. the data sets were not normally distributed and subsequently subjected to non-parametric statistical methods. an anova was performed to test for the possible effect of age and season on concentrations of faecal glucocorticoid metabolite levels. the computer program statistica (stasoft. 1995) was used for all statistical analyses. all tests were two-tailed, with the αlevel of significance set at 0.05. in cases of all pair-wise multiple comparison procedures, the α-level was adjusted by applying the procedure described by holm (1979). results no significant variation in concentrations of faecal glucocorticoid metabolites (h 3 = 7.54; n = 374; p = 0.057) was found across age classes (table 1). therefore, the age classes were combined for further analysis. although no significant differences in concentrations of faecal glucocorticoid metabolites were found between the four ecological seasons (table 2), there was a statistically significant difference (u = 15206.50; p = 0.032) between the wet season (n = 178) and dry season (n = 196), after the early and late period of both the dry and wet seasons were combined (figure 1). discussion this study provides new information on baseline concentrations of faecal glucocorticoid metabolites that could be expected within age classes and between seasons in african elephants in the southern kruger national park. our results show that the variability in baseline concentrations of faecal glucocorticoid metabolites (3α,11oxo-cm levels) in african elephant faeces is dependent on seasonal changes, rather than on the age class of the subject. in the present study faecal 3α,11oxo-cm levels differ significantly between seasons, but no differences were found across age classes. although the seasonal effect found seems to be rather small, future studies using methods of faecal hormone analysis to determine the effect of a potential stressor should account for seasonal effects, especially between the wet and dry season. the fact that no significant variation in faecal 3α,11oxo-cm levels was found across age classes confirm findings by ganswindt et al. (2005), who also reported no age effects on 3α,11oxo-cm levels in a group of african elephant bulls (n = 52, age range: 18–49 years) living in the samburu and buffalo springs national reserves, kenya. in the same study, ganswindt and colleagues described a small but clear elevation in 3α,11oxo-cm levels in longitudinal hormone profiles of african elephant bulls at the end of a long dry period (ganswindt et al. 2005). a correlation between season-dependent rainfall and adrenal endocrine function (highest concentrations of faecal glucocorticoid metabolites in the dry season) was further described for female elephants by foley et al. (2001). the present study confirms a possible influence of a dry period on increased glucocorticoid excretion, because the mean concentration of faecal glucocorticoid metabolites of samples collected in the dry season was significantly higher than in the wet season. the elevation of concentrations of faecal glucocorticoid metabolites during the dry season could be an indication of increased physiological stress due to a decline in food quantity and quality. this was a suggestion already made by codron et al. (2006), who reported that the percentage of nitrogen in elephant faeces from the southern knp showed a dramatic increase from the dry to the wet season. an elevation of nitrogen in faeces is known as a useful indicator of nutritional status. we could unfortunately not investigate differences in concentrations of faecal glucocorticoid metabolites between sexes due to safety considerations for the observers (distance away from the herd), and visual impairment caused by vegetation structure. however, further studies should examine the difference in 3α,11oxo-cm levels between the sexes after intense sampling from cows and bulls within both seasons, because sex could be a co-factor for the variability in baseline concentrations of faecal glucocorticoid metabolites. greyling (2004) found that during periods of resource limitation, males showed significantly lower levels of faecal minerals, together me a n me a n ±s e me a n ±s d d ry w e t w e t a n d d ry s e a s o n 2 0 4 0 6 0 8 0 1 0 0 1 2 0 1 4 0 1 6 0 fa ec al g lu co co rti co id (n g/ g) figure 1 box-plots of grouped concentrations of immunoreactive faecal glucocorticoid metabolites (ng/g) for free-ranging family herds of elephants in knp in the dry and wet seasons from may 2002 to august 2005. table 1 n, mean, standard deviation (concentrations of faecal glucocorticoid metabolites (ng/g ww)) in the four ecological seasons (age classes separate) for free-ranging family herds of elephants in knp from may 2002 to august 2005. season adults sub-adults juveniles n 3α,11oxo-cm conc. (mean±std) n 3α,11oxo-cm conc. (mean±std) n 3α,11oxo-cm conc. (mean±std) early dry 68 85.19, 47.08 50 94.04, 44.23 28 91.29, 56.22 late dry 23 79.70, 70.68 17 88.71, 69.90 10 80.10, 46.53 early wet 39 72.62, 43.39 23 68.48, 38.71 14 101.79, 54.27 late wet 41 67.61, 39.48 36 82.25, 48.51 25 81.36, 59.68 table 2 n, mean, standard deviation (concentrations of faecal glucocorticoid metabolites (ng/g ww)) in the four ecological seasons (age classes combined) for free-ranging family herds of elephants in knp from may 2002 to august 2005. season individuals n 3α,11oxo-cm conc.(mean±std) early dry 146 89.39, 47.86 late dry 50 82.84, 65.23 early wet 76 76.74, 45.29 late wet 102 76.15, 48.26 20 faecal glucocorticoid metabolites in african elephants original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 18 21 with higher levels of fibre, than adult females. these results indicate that males could be maintaining diets of poorer quality than females and consequently be nutritionally more stressed than females during the dry season. further studies should also investigate the difference in concentrations of faecal glucocorticoid metabolites between elephants from different areas, which would finally allow a comparison of mean concentrations of faecal glucocorticoid metabolites across elephant management regions within the knp (whyte et al. 2003). this would aid in creating a baseline for the entire park; that would be per season, per region and between different elephant densities. finally, this information could also be applied more universally, e.g. if a significant correlation with average monthly rainfall can be shown, or whether a direct correlation exists between increasing physiological stress and vegetation damage. conclusion a validated method (eia) was applied in assessing the level of physiological stress in free-ranging elephant herds. the method has been shown to be practical and enables long-term monitoring of ethological or environmental factors without interfering with the result. acknowledgements south african national parks is thanked for allowing fieldwork in the kruger national park. dr i. whyte and prof. d. g. a. meltzer for their assistance. we are grateful to mrs a. human for the laboratory work. we thank the field rangers of the lower sabie section for providing protection during data collection. the south african national research foundation and pittsburgh zoological society provided funding. references brown, j.l., c.m. wemmer & j. lehnhardt. 1995. urinary cortisol analysis for monitoring adrenal activity in elephants. zoo biology 14: 533–542. codron, j. j., lee-thorp, a., sponheimer, m., codron, d., grant, r.c & de ruiter, d.j. 2006. elephant (loxodonta africana) diets in kruger national park, south africa: spatial and landscape differences. journal of mammalogy 87(1): 27–34. friend, t.h. 1980. stress: what is it and how can it be quantified? int j stud anim prob 1 (6): 366–373. foley, c.a.h., papageorge,s.& wasser, s.k. 2001. non-invasive stress and reproductive measures of social and ecological pressures in free-ranging african elephants (loxodonta africana). conservation biology 15: 1134–1142. ganswindt a., heistermann, m., palme, r., borragan, s. & hodges, j.k. 2003. non-invasive assessment of adrenal function in the male african elephant (loxodonta africana) and its relation to musth. gen comp endocrinal 134: 156–166. ganswindt, a., rasmussen, h.b., heistermann, m. & hodges, j.k. 2005. the sexually active states of free-ranging male african elephants (loxodonta africana): defining musth and non-musth using endocrinology, physical signals, and behaviour. horm behav 47 (1): 83–91. goymann, w., möstl, e., van’t hof, t, east, m.l. & hofer, h. 1999. noninvasive fecal monitoring of glucocorticoids in spotted hyenas, crocuta crocuta. gen. comp. endocrinal 114: 340-348. greyling, m.d. 2004. sex and age related feeding distinctions in the feeding ecology of the african elephant, loxodonta africana. ph.d thesis, university of the witwatersrand, johannesburg. holm, s. 1979. a simple sequentially rejective multiple test procedure. scandinavian journal of statistics 6:65–70. liptrap, r.m. 1993. stress and reproduction in domestic animals. ann ny acad sci 697: 275–84. low, a. b. & rebelo, a. g. 1996. vegetation of south africa, lesotho and swaziland. department of environmental affairs & tourism, pretoria. merl, s., scherzer, s., palme, r. & möstl, e. 2000. pain causes increased concentrations of glucocorticoid metabolites in horse faeces. j equine vet sci 20: 586–90. moberg, g.p. 2000. biological response to stress: implications for animal welfare. in: moberg, g.p. & mench, j.a. (eds.) the biology of animal stress. pp. 123–46, cabi publishing, 2000. möstl, e., maggs, j.l., schrötter, g., besenfelder, u.& palme, r. 2002. measurement of cortisol metabolites in faeces of ruminants. vet res commun 26: 127–39. möstl, e. & palme, r. 2002. hormones as indicators of stress. domestic animal endocrinology 23: 67–74. munck, a., guyre, p.m. & holbrook, n.i. 1984. physiological function of glucocorticoids in stress and their relationship to pharmacological actions. endocr. rev. 5: 25–44. sands, j. & creel, s. 2004. social dominance, aggression and faecal glucocorticoid levels in a wild population of wolves, canis lupus. anim. behav. 67, 387–396. stasoft. 1995. statistica for windows. stasoft, inc., 2300 east 14th street, tulsa, ok u.s.a. stead, s.k., meltzer, d.g.a. & palme, r. 2000. the measurement of concentrations of faecal glucocorticoid metabolites in the serum and faeces of captive african elephants (loxodonta africana) after acth stimulation. tydskr. s.afr. vet 71(3): 192–195. venter, f. j, r., scholes, j.& eckhardt, h. c. 2003. the abiotic template and its associated vegetation pattern. in: du toit, j. t., rogers, k. h. & biggs, h. c. (eds.) pp.83–129, island press, washington, d.c. wasser, s. k., hunt, k.e., brown, j.l., cooper, k., crockett, c.m., bechert, u., millspaugh, j.j., larson, s.& monford, s.l. 2000. a generalized faecal glucocorticoid assay for use in a diverse array of non-domestic mammalian and avian species. gen comp endocrinol 120: 260–75. whyte, i. 2003. the movement patterns of elephant in the kruger national park in response to culling and environmental stimuli. pachyderm 16:72–80. whyte, i. j., van aarde, r.j., pimm, s.l. 2003. kruger’s elephant population: its size and consequences for ecosystem heterogeneity. in: j. t. du toit, rogers, k. h. & biggs, h. c., (eds.) pp.332-348, island press, washington, d.c. wimberger, k. 2001. aging free-ranging african elephants (loxodonta africana) using faecal bolus diameter. recitation for bsc honours. department of zoology and entomology, university of pretoria. (unpublished) zambatis, n. 2002. annual section ranger reports. knp vegetation monitoring. scientific services knp. (unpublished) 21 article information authors: adrian j. armstrong1 robert f. brand2 affiliations: 1conservation planning division, ezemvelo kzn wildlife, south africa 2environmental, water and earth sciences, tshwane university of technology, south africa correspondence to: adrian armstrong postal address: po box 13053, cascades 3202, south africa dates: received: 30 apr. 2012 accepted: 09 july 2012 published: 30 oct. 2012 how to cite this article: armstrong, a.j. & brand, r.f., 2012, ‘invertebrates on isolated peaks in the ukhahlamba-drakensberg park world heritage site, south africa’, koedoe 54(1), art. #1082, 10 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1082 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/ koedoe.v54i1.1082-1 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. invertebrates on isolated peaks in the ukhahlamba-drakensberg park world heritage site, south africa in this original research... open access • abstract • introduction    • aims • research method and design    • invertebrate species targeted    • study sites    • statistical analysis • results    • invertebrates    • vegetation and invertebrate association    • invertebrate data analysis • ethical considerations • discussion    • endemic invertebrates of the drakensberg alpine centre    • factors responsible for the diversity and composition of invertebrate communities on the inselbergs    • limitations of the study    • further research    • biodiversity conservation • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ a survey to document and describe the alpine flora and various focal faunal taxa on six isolated inselberg-like peaks (total area of 31.9 ha), all 3000 m or higher, located in the ukhahlamba-drakensberg park world heritage site, south africa, was undertaken in early summer in 2005. study of the fauna of these peaks should be informative because the impacts of controllable anthropogenic threats on the invertebrate communities on them should be minimal or absent in comparison with those on the main massif. a total of 341 invertebrate individuals representing 61 species were recorded from the focal taxa (oligochaeta, gastropoda and certain groups of insecta, i.e. focal taxa within the blattoidea, dermaptera, orthoptera, hemiptera, diptera, lepidoptera, coleoptera and hymenoptera). the 61 species recorded consisted of two species from the oligochaeta, one species from the gastropoda and 58 species from the insecta. eleven species (one from the oligochaeta, ten from the insecta) are endemic and 11 species (one from the oligochaeta, ten from the insecta) are probably endemic to the drakensberg alpine centre, constituting 36.1% of the total species recorded. the results suggest that the drakensberg alpine centre (dac), as for plants, is a centre of endemism for invertebrates. cluster analysis showed that the species composition of the two northern peaks, sentinel and eastern buttress, clustered together, separate from a cluster formed by the outer horn, inner horn and dragon’s back and from the cluster formed by the southernmost peak, cathkin. non-metric multi-dimensional scaling results indicated that distance from the sentinel, the most northerly peak sampled, and mean minimum temperature for july had the strongest correlations with the species data, reflecting change over a straight-line distance of nearly 60 km in a south-easterly direction.conservation implications: only a small proportion (ca. 5.5%) of the dac is conserved, the majority of which lies in the ukhahlamba-drakensberg park world heritage site. conservation of more of the dac, including more of its latitudinal extent, is required to adequately conserve its unique plant and invertebrate communities. introduction top ↑ the invertebrate fauna of the drakensberg alpine centre (dac), including the alpine region of the ukhahlamba-drakensberg park world heritage site in kwazulu-natal with its escarpment and peaks over 3000 m, has not been as well studied as some areas at lower altitudes in the province, for example, the world heritage site foothills and some of the midlands, coastal and savanna regions (bazelet & samways 2011; lovell et al. 2010; pryke & samways 2001, 2003, 2012; uys, hamer & slotow 2006, 2009, 2010; see also clark, barker & mucina 2011). consequently little is known regarding invertebrate distributions and invertebrate community composition and structure on high-altitude inselbergs in the drakensberg. the dac is located in south africa – in parts of kwazulu-natal (kzn), the eastern cape and free state – and in lesotho. it contains a unique assemblage of plants that is rich in endemics, and is an important centre of plant diversity in southern africa (killick 1978; van wyk & smith 2001). approximately 2520 angiosperm (higher plant) taxa, of which 334 (13%) are endemics, occur in the dac (carbutt & edwards 2004, 2006; van wyk & smith 2001). the dac is defined by floristic criteria (killick 1990), but was designated as ‘alpine’ based on climatological criteria (van wyk & smith 2001). the centre can be divided into a subalpine zone (c. from 1800 m a.s.l. to 2800 m a.s.l.) and an alpine zone (c. from 2800 m a.s.l. to 3482 m a.s.l.; killick 1978; van wyk & smith 2001). the peaks were chosen for several reasons: their altitude (situated within the alpine zone), the assumed infrequent nature or exclusion of the two major disturbance factors along the drakensberg escarpment, namely, fire and herbivory by domestic livestock, and the isolation of the peaks, or inselbergs, which can be regarded as an archipelago of ‘islands’, which theoretically could house unique, high-altitude endemics. fire frequency, extent and intensity, as well as season of burn and time since last burn can influence the composition of invertebrate assemblages at lower altitudes in the ukhahlamba-drakensberg park world heritage site (uys 2006; uys et al. 2006, 2010). invertebrate diversity is lower in grassland under communal grazing in comparison to grassland under nature conservation in a region south-east of the ukhahlamba-drakensberg park world heritage site (robertson & van noort 2003; wildy 2003). the results of an invertebrate survey carried out between 28 november and 05 december 2005 are presented here. the goal of the invertebrate survey was to sample six out of a possible 11 isolated peaks at altitudes of 3000 m or more in the ukhahlamba-drakensberg park world heritage site in kwazulu-natal. the peaks were chosen to ensure that a sufficient latitudinal and longitudinal range was incorporated by the sampling and that the peaks were large enough to safely land a helicopter and to establish sample plots adequate for invertebrate and vegetation sampling. aims the aims of this survey were to identify invertebrate species of focal groups that occur on the isolated peaks in the alpine zone of the kwazulu-natal section of the dac and to give some indication of whether:• the dac is a centre of endemism for invertebrates, where a relatively high proportion of invertebrate species are endemic. • invertebrate communities change from isolated peak to isolated peak, and possible reasons for this change. research method and design top ↑ invertebrate species targeted the invertebrate groups targeted during sampling included those known to have numbers of species endemic to mountainous areas of south africa (e.g. armstrong, van hensbergen & geertsema 1994), for some groups owing to a proportion or all of them being flightless or for which taxonomic expertise and/or aids to identification of species were available in south africa. these focal animal groups were as follows (other reasons for their inclusion in the study are given in parentheses): earthworms (soil engineers), snails (herbivores), large cockroaches (plant detritivores), wingless grasshoppers (herbivorous on forbs), leafhoppers and other bugs (herbivores, predators), beetles (herbivorous on various plant parts, predators of invertebrates, detritivores, decomposers), crane flies and robber flies (decomposers, insect predators), satyrid butterflies (herbivorous on grasses) and bees (pollinators).only adults of each species were included in the analyses because juveniles do not necessarily have the specific diagnostic identification characters. specimens were identified to species through: (1) reference to identified specimens and sometimes also to unidentified specimens housed in several collections, that is, the kwazulu-natal museum in pietermaritzburg (nmsa), and the south african national collection of insects (sanc) and the ditsong national museum of natural history, formerly the transvaal museum (tmsa), in pretoria, (2) identification keys, published taxonomic papers and a regional fauna guide (alexander 1964; brown 1962; eardley 1988; eardley, kuhlmann & pauly 2010; fennah 1967; iwan 2001; londt 1981; marshall 1904; princis 1963; pringle, henning & ball 1994; slater 1964; straneo 1958; synave 1956; theron 1986; wittmer 1949) and (3) specialist taxonomists (see acknowledgements). the distribution ranges and endemicity of the species were ascertained or estimated from specimen label data and distribution records contained in the publications listed above and in the biodiversity database of ezemvelo kzn wildlife, and from armstrong et al. (1994), armstrong and van hensbergen (1997a, 1997b, 1999), arnott (2006), hamer and slotow (2007) and uys et al. (2006, 2010). study sites six isolated peaks, all greater than 3000 m in elevation, were sampled during the survey, respectively occurring from north to south: the sentinel, eastern buttress, inner horn, outer horn, dragon’s back and cathkin (figures 1a and 1b). a 50 m by 50 m (0.25 ha) plot was demarcated using a tape-measure-calibrated ranging tlr 75 rangefinder and four orange-dyed, 1.5 m high, wooden dowels on a northerly-facing slope (where possible, one side was aligned to an approximately 59º magnetic bearing) of similar topographic profile on each peak. the top of the profile was a rocky slope and the bottom of the profile was more or less flat, generally with small shallow pools of standing water. topographic constraints (only precipitous northerly slopes) on the outer horn meant that the plot was predominantly flat with some of it on a southerly-facing slope. the co-ordinates of the centre of the plot at each site were recorded using a global positioning system (gps) (except for the sentinel plot, where the co-ordinates were obtained from a 1:50 000 scale topographic map). the surface area in hectares of each peak was calculated from the track log of a gps used to record these areas, and from a 1:30 000 orthophoto for the dragon’s back peak. a brief description of each plot and various parameters associated with each are given in table 1. figure 1: (a) map of south africa showing the extent of the drakensberg alpine centre (grey) and the study region (box). (b) enlargement of the study region indicating the study sites (dots). each plot was sampled by adrian j. armstrong for 15 minutes of search time (i.e. excluding time used to record or capture and store specimens), in three phases, by means of a slow meandering walk through the plot. one period of five minutes was concentrated on searching bare ground and rocks and also under rocks for endogaeic, epigaeic and other stationary invertebrates. the countdown timer was stopped when an area of suitable rocks for turning was encountered, and all turnable rocks within an area of approximately 4 m2 were turned over to reveal invertebrates hiding under the rocks. the rocks were then replaced in their original positions. the timer was started again when walking resumed. another period of five minutes of search time was spent searching flowering plants for pollinators. apart from bees, only insects that were feeding or sitting on the pollen-producing parts of the flowers were considered potential pollinators. a third period of five minutes search time was spent looking for low-flying insects, insects sitting on vegetation and insects disturbed from vegetation by the observer’s feet and net. specimens were collected for identification in the laboratory and in museums. sampling was done between 06h25 and 11h45, to make the most of the generally clearer early morning weather. the plot data were used in the comparison of sites. the comparison is relative and indicates differences in species density; it is not a comparison of total species complement. statistical analysis the true number of focal species on each plot was estimated from the sampling data in estimates (colwell 2005) using the chao1 estimator with the classic setting if there were no singletons or doubletons, or with the bias-corrected setting otherwise, and with the default settings for the other parameters. analysis of the plots by species matrix was carried out using pc-ord5 (mccune & mefford 1999). mccune and grace (2002) and the pc-ord5 advisor wizard were used to select an appropriate data relativisation method and analysis type. the percentage of cells empty was 74.317%. the co-efficient of variation (cv) of plot totals was 88.48% and the cv of species totals was 240.30%. species abundances were relativised by maximum to counter the effect that a few species that were relatively abundant at the time of sampling would have on the analysis and because the sampling time was restricted to a short period, which led to most species being rare in the samples. after relativisation, no plots or species were recognised as outliers using the sorensen (bray-curtis) distance measure and a cutoff of two standard deviations from the grand mean. a cluster analysis dendrogramme was produced using the sorensen (bray-curtis) distance measure and the flexible beta (beta set to −0.25) group linkage method. non-metric multidimensional scaling (nmds) was carried out using the autopilot mode and the sorensen (bray-curtis) distance measure (mccune & grace 2002). a random starting configuration and 250 runs with real data and 250 runs with randomised data for comparison were utilised. table 1a: brief description of the 50 m x 50 m plot at each site and parameters considered in the analysis. table 1b: brief description of the 50 m x 50 m plot at each site. the variables included in the environmental matrix in the nmds are indicated in table 1. the expectations were that:• the further away the peaks from the sentinel in straight-line distance, the greater the turnover of species and therefore the greater the difference in observed species complement. • species composition should be more similar between closer peaks than between peaks further apart. • peaks further away from the main massif were expected to have depauperate faunas compared to peaks closer to the main massif, owing to stochastic events that may have eliminated certain species that would not have recolonised the peak easily or rapidly from the main massif. • the larger the area of the peak, the greater the number of species that would be recorded. • differences in extreme temperatures (represented by the means of daily minimum temperature for the coldest month, july, and the means of daily maximum temperature for the hottest month, january) or else in mean annual temperature would influence the distribution of species on the peaks (climate data obtained from schulze 2007). • differences in mean annual precipitation between the peaks would cause differences in the faunal communities on the peaks. • differences in the altitude of the peaks are small and would not influence the composition of the faunal communities. the six peaks were grouped into a northern pair (sentinel and eastern buttress), a central pair (inner and outer horns) and a southern pair (dragon’s back and cathkin). results top ↑ invertebrates a total of 341 individuals of 61 species, comprising 26 families, of the focal taxa were recorded in all plots sampled (appendix 1). the total of 61 species of invertebrates included two orders, two families and two species of oligochaeta, a single order, single family and single species of gastropoda and eight orders, 23 families and 58 species (95% of the total invertebrate species sampled) of insecta. within the focal groups sampled, between 18% and 36% of the total invertebrate species recorded are endemic to the dac (known endemics and known + possible endemics, respectively; appendix 1), although the proportion varied between families, ranging from 0% to 100% (figure 2). twenty-nine species of the total 61 were precisely identified (three of which were described after the survey), four were undescribed, and eight others were probably undescribed (appendix 1). many of the remaining species may also be undescribed. eleven species are not known outside the dac, with a further 11 possibly endemic to the dac (appendix 1; figure 2). the outer horn had the most species (36) and the eastern buttress the fewest (4; appendix 1). the outer horn and dragon’s back peaks shared the most species (ten), followed by the inner horn and outer horn peaks with seven shared species (table 2). five species were shared between the outer horn and cathkin and between the outer horn and sentinel. the estimated true number of focal species for most plots was higher than that actually recorded (table 3). table 2: number of species shared between plots. figure 2: number of species recorded of each endemicity status (not endemic to the drakensberg alpine centre, endemic or probably endemic to the drakensberg alpine centre, and status unknown) in the invertebrate families sampled. table 3: estimated numbers (to nearest integer) of focal species present on each plot according to the chao1 estimator. table 4: pearson and kendall correlations between the environmental variables and the non-metric multidimensional scaling ordination axes. vegetation and invertebrate association the phytosociological survey done simultaneously with the invertebrate survey on the same inselbergs (brand, collins & du preez, in prep.) found the vegetation structured into four main communities consisting of shrubs, graminoids (grasses and sedges), forbs and geophytes. the first main vegetation community consists of shrubs and graminoids found on all peaks and consists of low shrubs (from 50 cm to 200 cm height) – erica glyphra, erica dominans, helichrysum trilineatum and anthospermum basuticum – plus tall bunch grass, merxmullera drakensbergensis, and the sedge scirpus falsus. the second main vegetation type consists mostly of high-altitude endemic grasses, pentaschistis exerta, pentaschistis basutorum and festuca caprina. this community predominates on the eastern buttress and cathkin peaks. the third community type is not well defined and consists of short, scattered forbs, geophytes and high-altitude, endemic grasses. the fourth main vegetation community is confined to wetlands or rocky seeps and consists of limited numbers of small, scattered, low geophytes, forbs and succulents most of which are obligate wetland plants, having a > 99% probability of occurring in wetlands (tiner 1999). these habitats with t heir associated vegetation and invertebrates occur on all the inselbergs, portions of which were included in the invertebrate sampling plots (figure 3a). as a result, no clear association between invertebrates and specific vegetation communities could be discerned. figure 3: (a) invertebrate sampling on cathkin peak. the unique high altitude vegetation is a mix of bunch grasses, merxmuellera and pentaschistis, and low cushion shrubs, erica, euryops and helichrysum, habitat for herbivore and detritivore invertebrates. (b) drakensberg escarpment, showing the island-like archipelago of inselbergs. summits in the distance from the right: inner tower, devil’s tooth and eastern buttress; sentinel in the foreground. (c) the flat surface of cathkin peak allows for low terrain heterogeneity, but houses high numbers of endemic invertebrates and plants. (d) the dac-endemic inflated-front wingless grasshopper, eremidium inflatifrons (female). (e) montane tail-wagger snail, sheldonia fuscicolor, on cathkin peak. invertebrate data analysis the cluster analysis indicated that the two northern peaks clustered separately from the cathkin peak cluster and the cluster formed by the other three peaks in terms of species composition, but that the closest peaks in straight-line distance did not necessarily cluster together the most closely (figure 4, table 1). figure 4: cluster analysis dendrogramme showing relationship between peaks in species space. the preliminary runs of the nmds ordination recommended a two-dimensional solution. the best two-dimensional solution had a final stress of 0.00 after 97 iterations. the monte carlo test result indicated that the final stress could not have been obtained by chance (p < 0.0478). the plot of stress versus iteration number for the final run was flat after iteration ten, indicating that a stable solution had been found. no good correlations between axis one of the nmds and the environmental variables were obtained, whereas some good correlations between axis two and the environmental variables were evident (table 4). the latter results indicated that the distance from the sentinel and the mean daily minimum temperature for july had the highest correlations with the species data (table 4), with distance to the closest peak having the next highest correlation. the correlations between the species data and the other environmental variables generally were low (table 4). ethical considerations top ↑ all work was done under ezemvelo kzn wildlife permit no. 2762/2004 (see the acknowledgements section). discussion top ↑ the study set out to sample focal taxa on a number of inselbergs in the alpine region of kzn to give an indication of whether the dac is a centre of endemism for invertebrates, whether invertebrate communities change from isolated peak to isolated peak, and possible reasons for any detected change. the inselbergs were chosen as study sites because of their relative isolation from the main drakensberg massif, on the assumption that the plant and animal communities have been little changed over the past few millennia by anthropogenic pressures such as livestock grazing and altered fire regimes. grazing by cattle and other domestic livestock is absent on the inselbergs and it is assumed that some uncontrolled fires started by humans do not reach the inselbergs because of their relative isolation. the main herbivores on the inselbergs are invertebrates and small mammals. rodent faeces, burrows and cuttings in the grass were found on the outer horn and cathkin peaks as well as small bones also suggestive of rodents (brand 2006, submitted). however, this herbivory is likely to have a positive effect on the vegetation in contrast to that of medium-sized to large domestic livestock. consequently, the vegetation can be regarded as being ‘pristine’ and not exhibiting the effects of over-grazing by cattle and sheep. this study of the plant and invertebrate communities on the inselbergs of the alpine region of the ukhahlamba-drakensberg park world heritage site appears to be the first of its kind. the conservation context of the study is the planned increase in area under protection in the maloti drakensberg transfrontier project planning region (maloti drakensberg transfrontier project 2008). endemic invertebrates of the drakensberg alpine centre between 18% and 36% of the invertebrate species recorded are endemic to the dac, a higher percentage than that for angiosperms as a whole (carbutt & edwards 2006). this is suggestive that the dac is also a centre of endemism for invertebrates, although much more field collecting and taxonomic work on the invertebrate fauna is required (clark et al. 2011) before this suggestion can be substantiated. a firm conclusion on the exact percentage of the invertebrates recorded during this study that are endemic to the dac is hampered by the fact that a large proportion of the species could not be identified. only a proportion of the invertebrate taxa on the inselbergs (the focal taxa) could be sampled in the short time available. to compile for the dac as comprehensive a list of insects and other focal invertebrates as that compiled for the angiosperms, so as to determine amongst other things the exact percentage of endemism for these invertebrates, will require much more intensive and extensive sampling in the various seasons throughout the dac.the number of undescribed species indicates that the invertebrate fauna of the dac is still not well-described. certain species, however, do appear to have restricted ranges even within the dac (appendix 1). the fact that some endemic microchaetid earthworms and wingless bugs have restricted distribution ranges is well known (plisko 2003; stiller 2009, 2010; theron 1986). these species may only occur on certain peaks as a result of the spatial distribution of the environmental factors that influence the species’ distributions and the relative immobility of the species. these environmental factors may include soil attributes such as ph, depth, and clay and water content, and vegetation attributes such as physiognomy and host-plant distributions (e.g. edwards & bohlen 1996 for earthworms). species associated with grasses (because they or their larvae are grass-feeders or else they are associated with forbs growing in grassland) would be more likely to be found on the peaks with a greater proportion of the area under grasses, these being the sentinel, dragon’s back, and cathkin (main plant communities one and two comprised of shrubs and grasses). lepidopteran and hemipteran grass-feeders such as pseudonympha gaika, pravistylus interdiscus, and drakensbergena bisulca (pringle et al. 1994; stiller 2009, 2010), and an orthopteran that feeds on leaves of bushes or forbs in grassland, basutacris inflatifrons (brown 1962; figure 3d), tend to show this type of distribution (appendix 1). other possibilities are that specific plant species with which certain wingless species are associated may not occur on some inselbergs, or finally, some wingless species may not be present on certain peaks because they have become extinct on those peaks through the impact of environmental stochasticity. factors responsible for the diversity and composition of invertebrate communities on the inselbergs inselbergs are defined as isolated, freestanding topographic features, clearly visible in the landscape (gröger & barthlott 1996; parmentier et al. 2006). they are composed either of igneous or sedimentary rock and have sheer, or near sheer sides (gröger & barthlott 1996). they constitute sites of high biodiversity with large numbers of endemic taxa and represent relictual populations of previous episodes of climate change (parmentier et al. 2006; porembski et al. 1996). the six inselbergs surveyed are isolated from the escarpment by several hundred metres and form island-like archipelago strings of peaks with sheer basalt sides up to 1000 m high. they are all inaccessible to humans except by climbing or helicopter. the length of time the individual peaks have been separated could not be inferred from geological or geomorphological information, but king (1963) accounts for a retreat of the original escarpment rim near the current day kzn coast of 161 km in 123 million years, which killick (1990) reduces to an average rate of 1.3 mm per year.the most important factors influencing invertebrate community composition on the inselbergs was distance from the sentinel and mean minimum temperature for july, according to the nmds analysis (table 4). distance from the sentinel reflects how much further away in a south-easterly direction the next peak is from the sentinel (figure 1b). turnover of species in this south-easterly direction is reflected. that the two northern peaks clustered together, the most southerly peak, cathkin, formed its own cluster, and the other three peaks formed a cluster according to the cluster analysis (figure 4) gives support to this contention. mean annual rainfall tends to increase in this direction, as do the maximum and minimum temperature figures to a lesser extent (table 1). change in species identities and taxon richness occurs with increasing altitude in the drakensberg mountains and, at least for some taxa, fewer species are encountered at higher altitudes (e.g. armstrong & van hensbergen 1997a, 1997b; samways 1989; stuckenberg 1962). pryke and samways (2008, 2010) found that invertebrate species richness, abundance and composition changed with altitude on table mountain. epigaeic and aerial invertebrates decreased in richness and abundance with increasing altitude but the converse was true for foliage invertebrates. therefore, beta diversity might be expected to be lower at higher altitudes in the dac. only one species was recorded in all three clusters (figure 4; appendix 1). more intensive sampling may reveal that other species occur in all the clusters. uys et al. (2009) determined high turnover values for flightless forest invertebrates at scales varying from a few hundred metres to tens of kilometres at lower altitudes in the same general region of the ukhahlamba-drakensberg park world heritage site as this study. mean minimum temperature for july (the coldest month) was found to influence invertebrate community composition. the temperature data were at a one minute by one minute scale that was possibly too large to give precise temperatures for each peak. the south african weather service has 17 stations in the drakensberg, of which two, sani pass (2865 m) and witsieshoek (2500 m, below the sentinel), are at high altitude, yet still below the altitudes of the study sites (nel 2007). the influence of low temperatures, frost, and low oxygen partial pressure on insects has been documented (chapman 2002), and these play an appreciable part as limiting factors responsible for insect endemism and diversity at high altitude. altitude and vegetation structure are two of the most important environmental variables that influence invertebrate diversity on table mountain (pryke & samways 2008, 2010). the texture and structure of alpine vegetation, regardless of slope, aspect and soils, is dependent on the maximum and minimum temperatures (körner 2003), which might thus influence the relationship between invertebrates and vegetation type. insects and other invertebrates, like lizards, are poikilotherms; their temperature depends largely on the amount of incident solar radiation and on the ambient temperature (chapman 2002; clusella-trullas et al. 2008). clusella-trullas et al. (2008) found that skin reflectance was more strongly related to mean annual radiation than to mean annual temperature in their sample of 68 and 66 species of lizards, respectively. dark beetles absorb more direct shortwave and visible radiation than light-coloured ones, which would enable them to use more limited amounts of sunlight to warm up and become active, providing wind is absent (e.g. turner & lombard 1990). alpine and arctic insects are often dark, or have darker scales and wing undersides to take advantage of low ambient light and temperature levels (chapman 2002). about 60% of the insect species sampled during the present study are black or very dark brown or grey, and may be darker than congeners at lower altitudes. the montane tail-wagger snail sheldonia fuscicolor (figure 3e), for example, is darker than its congeners at lower altitudes in kzn (herbert & kilburn 2004). insect taxa that have species known to be frost tolerant include blattoidea, hemiptera, diptera, lepidoptera, coleoptera and hymenoptera (chapman 2002; cf. appendix 1). insects living in the dac have to endure temperatures close to and below freezing at various times of the year, and especially in winter, and so are likely to be freeze tolerant (sinclair & chown 2005). two species from the kzn part of the dac, a cockroach (probably perisphaeria guillarmodi; s. van noort, pers. comm., 23 april 2012) and a ground beetle, were included in the study of sinclair and chown (2005). both these species are moderately freeze-tolerant, with a lethal temperature lower than the temperature of crystallisation. the low temperature at which 50% mortality (lt50) of the cockroach was recorded was −9.7 ºc and the lt50 for the ground beetle was −4.4 ºc (sinclair & chown 2005). therefore, many of the species recorded are likely to be adapted to the low solar radiation incidence and ambient temperatures experienced during the winter season and the frequently cloudy and wet summer period in the dac (killick 1978). the ability to endure low solar radiation and temperatures is likely a major factor that influences the composition of invertebrate communities in the dac. expectations one, two, five and seven were the only ones supported or partially supported by the data (see research methods and design: statistical analysis; figure 4; tables 1, 2 & 4). surface area of the inselbergs was not important in terms of structuring the invertebrate communities nor in terms of the number of species present (tables 1 & 4). the eastern buttress yielded only four species from the focal groups, even though it was the second-largest peak, possibly as a result of the cool and damp conditions following the storm the evening before sampling, which may have inhibited some invertebrate activity and therefore detection of certain species. the outer horn and the inner horn were expected to have the most similar faunal community in terms of invertebrate species identities, because they were the pair of peaks closest together (table 1). they shared seven species (as recorded during the plot counts), and this pair of peaks was ranked second in number of shared species. one of the more important factors allowing for high biodiversity is terrain heterogeneity (cowling & lombard 2002), that is, mountains tend to be topographically complex with variations of soil, rocky slopes, and aspect and have more species than flat areas due to the greater number of ecological niches available. the relatively flat topography on the inselbergs (figure 3b & 3c) does not allow for significant habitat differentiation and only four main vegetation communities (habitats) are found. despite this, of the total 61 invertebrate taxa found, 11 are dac endemics, with a further 11 possibly endemic to the dac (figure 2). this represents 36.1% endemic and possibly endemic species out of the total fauna sampled and is comparable with that found in a study of invertebrates on table mountain, cape town, where habitat heterogeneity was higher (total of 175 species, with 6 surface-endemic invertebrate species and 28 cape peninsula endemics, giving 16% endemicity; pryke & samways 2008, 2010). endemicity varied from 0% to 100% amongst the various families recorded in the present study (figure 2). picker and samways (1996) recorded percentage endemicity of between 5% and 67% for various faunal taxa on the cape peninsula in a far more comprehensive study. limitations of the study the fact that a high percentage of invertebrate species were only represented by a single specimen (appendix 1) and that the true numbers of focal species on the plots is likely to be higher than those recorded (table 3) indicates that more time needs to be spent sampling on the peaks in all four of the main vegetation communities of the study plots to gain a fuller picture of the invertebrate communities and the associations between the invertebrates and the plant communities. this was not possible during the present study owing to logistical constraints, including the inaccessibility of the peaks, which could only be reached by helicopter and in good flying conditions, and the mountain weather conditions, which are very variable in early summer in the drakensberg. the weather conditions, particularly with respect to the formation of clouds, can and did change rapidly at the sampling sites. in particular, the periodic overcast conditions that pertained during sampling may have suppressed insect activity to some extent and, therefore, detection of insects. further research further sampling to elucidate the impacts that unmanaged burning and livestock grazing have on the invertebrate fauna of the dac would be valuable. a survey design that used paired plots, on the main drakensberg massif that is subject to human activity, especially burning and livestock grazing, and on neighbouring isolated peaks that are not, or seldom, subject to these impacts, would be most appropriate. this may be difficult to achieve, as access to the peaks is conditional on sufficient budget for helicopter transport, and the prevailing weather conditions.the results of the present study suggest that the focal invertebrate taxa for sampling should be groups with range-restricted dac-endemic species, such as (other reasons in parentheses): earthworms (detritivores and soil engineers that are likely to be sensitive to soil water changes caused by trampling and excessive removal of vegetation and plant litter), wingless grasshoppers (herbivores that may be sensitive to changes in vegetation structure and composition), flightless bugs (herbivores that are likely to be sensitive to vegetation changes caused by excessive fire and livestock herbivory), crane flies (detritivores that are likely to be sensitive to changes in vegetation and surface water), ground beetles (predators of other invertebrates), darkling beetles (detritivores) and hair beetles (decomposers of vertebrate remains that may be sensitive to changes in abundance of mammals and birds caused by hunting pressure and changes in vegetation structure caused by altered fire and herbivory regimes). a relatively large proportion of insect species inhabiting high-altitude areas are wingless (e.g. endrődy-younga 1989; roff 1990). these insects are most likely to be vulnerable to large-scale changes in vegetation as their mobility is reduced compared to fully-winged species. the sampling methods used would have to be appropriate to these groups and the sampling would have to be intensive enough to discern changes in the abundance and identities of species occurring in the treatment area relative to the control. the precise relationship between plant community structure and alpine invertebrate use of, dependence on, and insect pollination of plants is not clear from this survey and requires further investigation. a vegetation-based approach to insect conservation may be fruitful (e.g. panzer & schwartz 1998). biodiversity conservation the diversity of invertebrates and plants found on the six inselbergs in the drakensberg is exceptional, especially considering that 61 focal invertebrate species and 189 plant species were recorded on the 31.9 ha studied. the high number of endemic and possibly endemic invertebrate species (22), which constitutes about 36% of the invertebrate species sampled, coupled with the 112 endemic and near endemic plant species, adds further prominence to the unique faunal and floral value of this region of the dac. the inselbergs represent refugia for plants and animals because these are sites relatively untouched by direct anthropogenic influence. climate change is likely to start influencing the diversity of plant species, vegetation structure and invertebrate species on the inselbergs. however, this should not prevent these high-altitude, alpine areas from receiving statutory protection. carbutt and edwards (2006) calculated that only about 5.5% of the dac is formally conserved, and about 97% of this conserved area occurs within the ukhahlamba-drakensberg park world heritage site in kwazulu-natal. the latter authors have already noted the need for the continued protection of the biota of the dac. the present study provides evidence that it is not only the unique and threatened flora that needs continued protection and conservation, but also the relatively large proportion of invertebrate species that are endemic to the dac. conservation of more of the dac should form the basis for inter-institution and international (lesotho and south african) collaboration to conserve its unique plants and animals, and also to obtain a better understanding of this alpine ecosystem through research in various and distant areas of the dac where anthropogenic impacts are minimal. conclusion top ↑ this study is the first to have sampled invertebrate taxa in a standardised way (timed counts on a 50 m by 50 m plot) on the top of each of six high inselbergs of the ukhahlamba-drakensberg park world heritage site, which are free of most anthropogenic impacts. the results suggest that, as for plants, the dac is a centre of endemism for invertebrates, with a substantial number of undescribed species and species endemic to the dac being recorded. more sampling both on the inselbergs and on the main massif is required to enable an accurate estimate of the proportion of invertebrate species that are endemic to the dac to be made, as well as to give an indication of how various anthropogenic threats impact the invertebrate fauna. invertebrate species turnover was shown to occur in a south-easterly direction. the results indicate that more of the latitudinal and longitudinal extent of the dac should be given statutory protection, as there are likely to be invertebrate species endemic to the dac that are not yet protected. this would be best achieved through the ongoing collaboration between south africa and lesotho through the maloti-drakensberg transfrontier park programme. acknowledgements top ↑ the national geographic committee for research and exploration supported the fieldwork (grant # 7920-05), as without these funds the study would not have been possible. we thank the following people for identifying specimens or for assisting with their identification: danuta plisko (earthworms), michael stiller (bugs), david rider (aegaleus), dai herbert (snails), beth grobbelaar (beetles), riaan stals (beetles), ruth muller (beetles) and james harrison (beetles). we also thank nunku mkhize and keenan stears for setting specimens. richard lechmere-oertel is thanked for providing the areas of the larger peaks calculated from the track logs of a gps and for photographs, debbie jewitt for advice on using pc-ord5 and comments on a previous version of the manuscript, and heidi snyman for assistance with the production of some of the figures. specimens were collected under ezemvelo kzn wildlife permit no. 2762/2004 and are deposited in the south african national collection (sanc), the ditsong national museum of natural history (tmsa), the durban natural science museum (dmsa) and the kwazulu-natal museum (nmsa). the photographs in figures 3a, 3b and 3c are courtesy of richard lechmere-oertel of the maloti-drakensberg transfrontier park project. editorial comments by ray jansen of tshwane university of technology (tut) and the comments of two anonymous reviewers were very useful and improved the manuscript. competing interests the authors declare that they have no financial or personal relationship(s) which may have inappropriately influenced them in writing this paper. authors’ contributions a.j.a. (ezemvelo kzn wildlife) was responsible for the sampling techniques, collection of field data, processing of specimens and deposition in museums, identification of specimens through collaboration with taxonomic experts, collation of ancillary data such as distribution ranges from specimens and the literature, analysis of the data, and writing up of the paper. r.f.b. (tshwane university of technology) was the project leader and developed the concept of sampling on the isolated peaks, secured the grant that enabled the fieldwork to be undertaken, contributed ideas as to how to further develop the paper, and contributed to the botanical and other sections of the paper. references top ↑ alexander, c.p., 1964, ‘diptera (nematocera): tanyderidae, ptychopteridae, tipulidae’, in b. hanstrom, p. brink & g. rudebeck (eds.), south african animal life, vol. 10, pp. 229–441, swedish natural science research council, stockholm. pmid:21540865 armstrong, a.j. & van hensbergen, h.j., 1997a, ‘evaluation of afforestable montane grasslands for wildlife conservation in the north-eastern cape, south africa’, biological conservation 81, 179–190. pmid:17538161 armstrong, a.j. & van hensbergen, h.j., 1997b, ‘a test of the applicability of a wildlife conservation evaluation method to an afforestable region of mpumalanga’, south african journal of wildlife research 27, 73–77. pmid:17960012 armstrong, a.j. & van hensbergen, h.j., 1999, ‘identification of priority regions for animal conservation in afforestable montane grasslands of the northern eastern cape province, south africa’, biological conservation 87, 93–103. pmid:18784101 armstrong, a.j., van hensbergen, h.j. & geertsema, h., 1994, ‘evaluation of afforestable montane grasslands for wildlife conservation in the north-eastern cape, south africa. part 1. methods’, south african forestry journal 171, 7–20. pmid:19197326 arnott, w.l., 2006, ‘the effect of burning frequency on invertebrate and indigenous flowering forb diversity in a drakensberg grassland ecosystem’, msc thesis, school of biological and conservation sciences, university of kwazulu-natal, pietermaritzburg. bazelet, c.s. & samways, m.j., 2011, ‘relative importance of management vs. design for implementation of large-scale ecological networks’, landscape ecology 26, 341–353. brand, r.f., 2006, ‘the ecology and biogeography of pre and post gondwana inselbergs, drakensberg alpine centre, south africa’, report to the national geographic society, committee for science and exploration, washington, dc. brand, r.f., submitted, ‘a floristic, biogeographic and growth form assessment of afroalpine inselberg vegetation in the drakensberg alpine centre’, south african journal of botany. pmid:8136946 brand, r.f., collins, n. & du preez, p.j., in prep., ‘a phytosociology survey and biogeographical description of inselberg vegetation in the drakensberg alpine centre, south africa’. brown, h.d., 1962, ‘new and interesting grasshoppers from southern africa – 3 (othoptera: acridoidea)’, journal of the entomological society of south africa 25, 198–229. carbutt, c. & edwards, t.j., 2004, ‘the flora of the drakensberg alpine centre’, edinburgh journal of botany 60, 581–607. carbutt, c. & edwards, t.j., 2006, ‘the endemic and near-endemic angiosperms of the drakensberg alpine centre’, south african journal of botany 72, 105–132. chapman, r.f., 2002, ‘thermal relations’, in r.f. chapman (ed.), the insects: structure and function, pp. 509–530, cambridge university press, cambridge. clark, v.r., barker, n.p. & mucina, l., 2011, ‘the great escarpment of southern africa: a new frontier for biodiversity exploration’, biodiversity and conservation 20, 2543–2561. clusella-trullas, s., terblanche, j.s., blackburn, t.m. & chown, s.l., 2008, ‘testing the thermal melanism hypothesis: a macrophysiological approach’, functional ecology 22, 232–238. colwell, r.k., 2005, estimates: statistical estimation of species richness and shared species from samples, version 7.5, user’s guide and application published, viewed 11 june 2012, from http://purl.oclc.org/estimates cowling, r.m. & lombard, a.t., 2002, ‘heterogeneity, speciation/extinction history and climate: explaining regional plant diversity patterns in the cape floristic region’, diversity and distributions 8, 163–179. eardley, c.d., 1988, ‘a revision of the genus lithurge latreille (hymenoptera: megachilidae) of sub-saharan africa’, journal of the entomological society of south africa 51, 251–263. eardley, c., kuhlmann, m. & pauly, a., 2010, ‘the bee genera and subgenera of sub-saharan africa’, abc taxa 7, 1–139. edwards, c.a. & bohlen, p.j., 1996, biology and ecology of earthworms, chapman & hall, london. endrődy-younga, s., 1989, ‘the evolution of alternative life styles in coleoptera’, in m.n. bruton (ed.), alternative life-history styles of animals, pp. 317–327, kluwer academic publishers, dordrecht. fennah, r.g., 1967, ‘new and little-known fulgoroidea from south africa (homoptera)’, annals of the natal museum 18, 655–714. gröger, a. & barthlott, w., 1996, ‘biogeography and diversity of the inselbergs (laja) vegetation of southern venezuela’, biodiversity letters 3, 165–179. hamer, m. & slotow, r., 2007, ‘an assessment and ecological interpretation of the terrestrial invertebrate fauna of the south african portion of the maloti-drakensberg bioregion’, unpublished report to the maloti-drakensberg transfrontier park project, inland invertebrate initiative, university of kwazulu-natal, pietermaritzburg. herbert, d. & kilburn, d., 2004, field guide to the land snails and slugs of eastern south africa, natal museum, pietermaritzburg. iwan, d., 2001, ‘two new species of bantodemus koch, 1955 (coleoptera: tenebrionidae: platynotini) from south africa’, annales zoologici (warszawa) 51, 505–510. killick, d.j.b., 1978, ‘the afro-alpine region’, in m.j.a. werger (ed.), biogeography and ecology of southern africa, pp. 515–560, junk, the hague. killick, d.j.b., 1990, a field guide to the flora of the natal drakensberg, jonathan ball and ad. donker, johannesburg. king, l.c., 1963, south african scenery: a textbook of geomorphology, oliver and boyd, edinburgh. körner, c., 2003, alpine plant life: functional plant ecology of high mountain ecosystems, springer-verlag, berlin. londt, j.g.h., 1981, ‘afrotropical asilidae (diptera) 5: the genus dasophrys loew, 1858 (= hobbyus bromley, 1952) (asilinae: asilini)’, annals of the natal museum 24, 635–699. lovell, s.j., hamer, m.l., slotow, r.h. & herbert, d., 2010, ‘assessment of sampling approaches for a multi-taxa invertebrate survey in a south african savanna-mosaic ecosystem’, austral ecology 35, 357–370. maloti drakensberg transfrontier project, 2008, ‘conservation and development strategy (2008–2028) for the maloti drakensberg transfrontier conservation area’, maloti drakensberg transfrontier project south africa, howick. marshall, g.a.k., 1904, ‘a monograph on the coleoptera of the genus hipporrhinus, schh. (curculionidae)’, proceedings of the zoological society of london, 6–141. mccune, b. & grace, j.b., 2002, analysis of ecological communities, mjm software design, gleneden beach, oregon. mccune, b. & mefford, m.j., 1999, pc-ord: multivariate analysis of ecological data, version 4, mjm software design, gleneden beach, oregon. nel, w., 2007, ‘on the climate of the drakensburg: rainfall and surface-temperature attributes, and associated geomorphic effects’, phd dissertation, department of geography, geo-informatics and meteorology, university of pretoria, pretoria. panzer, r. & schwartz, m.w., 1998, ‘effectiveness of a vegetation-based approach to insect conservation’, conservation biology 12, 693–702. parmentier, i., oumorou, m., porembski, s., lejoly, j. & decocq, g., 2006, ‘ecology, distribution, and classification of xeric monocotyledonous mats on inselbergs in west africa and atlantic central africa’, phytocoenology 36, 547–564. picker, m.d. & samways, m.j., 1996, ‘faunal diversity and endemicity of the cape peninsula, south africa – a first assessment’, biodiversity and conservation 5, 591–606. plisko, j.d., 1994, ‘new and little-known species of proandricus plisko, 1992 from lesotho and transkei (southern africa) (oligochaeta: microchaetidae)’, annals of the natal museum 35, 229–235. plisko, j.d., 2003, ‘eleven new south african earthworms (oligochaeta: microchaetidae) with new information on some known species, and an inventory of the microchaetids of kwazulu-natal’, african invertebrates 44, 279–325. porembski, s., szarzynske, j., mund, j-p. & barthlott, w., 1996, ‘biodiversity and vegetation of small-size inselbergs in a west african rain forest (taï, ivory coast)’, journal of biogeography 23, 47–55. princis, k., 1963, ‘blattariae’, in b. hanstrom, p. brinck & g. rudebeck (eds.), south african animal life, vol. 9, pp. 9–318, swedish natural science research council, stockholm. pringle, e.l.l, henning, g.a. & ball, j.b. (eds.), 1994, pennington’s butterflies of southern africa, struik, cape town. pryke, j.s. & samways, m.j., 2008, ‘conservation of invertebrate biodiversity on a mountain in a global biodiversity hotspot, cape floral region’, biodiversity and conservation 17, 3027–3043. pryke, j.s. & samways, m.j., 2010, ‘significant variables for the conservation of mountain invertebrates’, journal of insect conservation 14, 247−256. pryke, j.s. & samways, m.j., 2012, ‘ecological networks act as extensions of protected areas for arthropod biodiversity conservation’, journal of applied ecology 49, 591−600. pryke, s.r. & samways, m.j., 2001, ‘width of grassland linkages for the conservation of butterflies in south african afforested areas’, biological conservation 101, 85–96. pryke, s.r. & samways, m.j., 2003, ‘quality of remnant indigenous grassland linkages for adult butterflies (lepidoptera) in an afforested african landscape’, biodiversity and conservation 12, 1985–2004. robertson, h.g. & van noort, s., 2003, ‘effects of agricultural practices on ants and other hymenoptera in four farming areas of south africa’, unpublished conservation farming project report to the national botanical institute, pretoria. roff, d.a., 1990, ‘the evolution of flightlessness in insects’, ecological monographs 60, 389–421. samways, m.j., 1989, ‘taxon turnover in odonata across a 3000 m altitudinal gradient in southern africa’, odonatologica 18, 263–274. schulze, r.e. (ed.), 2007, ‘south african atlas of climatology and agrohydrology’, wrc report 1489/1/06, water research commission, pretoria. sinclair, b.j. & chown, s.l., 2005, ‘climatic variability and hemispheric differences in insect cold tolerance: support from southern africa’, functional ecology 19, 214–221. slater, j.a., 1964, ‘hemiptera (heteroptera). lygaeidae’, in b. hanstrom, p. brinck & g. rudebeck (eds.), south african animal life, vol. 10, pp. 15–228, swedish natural science research council, stockholm. stiller, m., 2009, ‘a revision of the south african leafhopper genus drakensbergena linnavuori (hemiptera: cicadellidae: drakensbergeninae)’, zootaxa 2132, 1–39. stiller, m., 2010, ‘revision of the southern african leafhopper genus pravistylus (hemiptera: cicadellidae: deltocephalinae)’, zootaxa 2468, 1–81. straneo, s.l., 1958, ‘carabidae pterostichinae’, in b. hanstrom, p. brinck & g. rudebeck (eds.), south african animal life, vol. 5, pp. 9–318, swedish natural science research council, stockholm. stuckenberg, b.r., 1962, ‘the distribution of the montane palaeogenic element in the south african invertebrate fauna’, annals of the cape provincial museums 2, 190–205. synave, h., 1956, ‘contribution a l’étude des issidae africains (homoptera – fulgoroidea)’, bulletin de institute royal des sciences naturelles de belgique 32, 1–22. theron, j.g., 1986, ‘new genera and species of southern african coelidiinae (homoptera: cicadellidae), with description of the new tribe equeefini’, phytophylactica 18, 153–163. tiner, r.w., 1999, wetland indicators: a guide to wetland identification, delineation, classification and mapping, lewis publishers, washington, d.c. turner, j.s. & lombard, a.t., 1990, ‘body color and body temperature in white and black namib desert beetles’, journal of arid environments 19, 303–315. uys, r., 2006, ‘fire effects on the fauna and flora of the maloti-drakensberg bioregion: a review’, unpublished report, maloti-drakensberg transfrontier project, ezemvelo kzn wildlife, pietermaritzburg. uys, c., hamer, m. & slotow, r., 2006, ‘effect of burn area on invertebrate recolonization in grasslands in the drakensberg, south africa’, african zoology 41, 51–65. uys, c., hamer, m. & slotow, r., 2009, ‘turnover in flightless invertebrate species composition over different spatial scales in afrotemperate forest in the drakensberg, south africa’, african journal of ecology 47, 341–351. uys, c., hamer, m. & slotow, r., 2010, ‘step process for selecting and testing surrogates and indicators of afrotemperate forest invertebrate diversity’, plos one 5(2), e9100. pmid:20161757 van wyk, a.e. & smith g.f., 2001, regions of floristic endemism in southern africa: a review with emphasis on succulents, umdaus press, hatfield. wildy, e.j., 2003, ‘effect of different land use practices on invertebrate diversity in underberg (kwazulu-natal, south africa)’, unpublished conservation farming project report to the national botanical institute, pretoria. wittmer, von w., 1949, ‘neue malacodermata aus der sammlung des british museum, london (2. beitrag zur kenntnis der malacodermata afrikas)’, annals and magazine of natural history series 12 2, 746–764. article information authors: dirk j. roux1,2 llewellyn c. foxcroft3,4 affiliations: 1south african national parks, george, south africa 2sustainability research unit, nelson mandela metropolitan university, george, south africa 3south african national parks, skukuza, south africa 4centre for invasion biology, department of botany and zoology, stellenbosch university, south africa correspondence to: dirk roux email: dirkr@sanparks.org postal address: private bag x 6531, george 6530, south africa how to cite this article: roux, d.j. & foxcroft, l.c., 2011, ‘the development and application of strategic adaptive management within south african national parks’, koedoe 53(2), art. #1049, 5 pages. doi:10.4102/koedoe.v53i2.1049 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) the development and application of strategic adaptive management within south african national parks in this editorial... open access • introduction • strategic adaptive management in sanparks    • philosophical foundation       • adaptive planning       • adaptive implementation       • adaptive evaluation • roadmap through this special issue • conclusion • acknowledgements • references introduction (back to top) adaptive management is an appealing approach to deal with inherent uncertainty in complex and interactive social-ecological systems (holling 2001; rogers 2003). in short, adaptive management is about learning-by-doing in a scientific way, adapting behaviour and overall direction as new information becomes available. it provides a structured way for improving our incomplete understanding through an iterative process of setting objectives, implementing policy decisions and evaluating the implications of their outcomes for future decision making. in essence, adaptive management is: the process of treating natural resource management as an experiment such that the practicality of trial and error is added to the rigour and explicitness of the scientific experiment, producing learning that is both relevant and valid. (meffe et al. 2002) when adaptive management is practiced, policies become hypotheses and management actions become the experiments to test those hypotheses (folke et al. 2005). first referred to as adaptive environmental assessment and management (aeam) (holling 1978; walters 1986), adaptive management has grown into an established field of research and practice (allan & stankey 2009; armitage, berkes & doubleday 2008; meffe et al. 2002; oglethorpe 2002; walters 2002). whilst there is a rich literature on the philosophical merits of adaptive management, the actual day-to-day implementation had faced many obstacles (berkes, colding & folke 2003; johnson 1999; lee 1993; mclain & lee 1996; rogers 2003; shea et al. 2002; walters 1997). yet a version of adaptive management that developed in south africa has grown to become an integral part of the thinking, planning and decision-making within south african national parks (sanparks). this version is referred to as strategic adaptive management (sam) and this special issue is dedicated to reflecting on the development and implementation of sam within sanparks and its stakeholder community over a 10-year period. in this editorial we present a brief introduction to the main components and vocabulary of sam as practiced by sanparks, as well as a roadmap through the papers that constitute the two parts of the special issue. the papers that follow in this special issue, and the references therein, will provide the reader with a rich source of literature and in-depth treatise of sam and its development within sanparks. strategic adaptive management in sanparks (back to top) philosophical foundation the century-long evolution of management practices in the kruger national park is chronicled in venter et al. (2008). an optimisation approach in the early years (c.1902–1980), as well as command-and-control (1955–1985) and laissez-faire (1985–1995) approaches were embraced before the adoption of adaptive management in 1995. the appropriateness of adaptive management for natural resource management in general, and its adoption by kruger national park, stems from a growing awareness of two critical challenges, (1) the existence of ecological complexity and social complexity and hence social-ecological complexity and (2) the existence of multiple stakeholders with diverse (and often divergent) perceptions, values and expectations. a defining characteristic of complex systems is that patterns emerge or self-organise from the local interactions between components of the system. the interactions and feedbacks between components can be ‘nonlinear’, resulting in an inherent degree of unpredictability in cause-and-effect relationships and making them ‘knowable’ only in retrospect. an emergent property, for example patterns in organism distribution, is not a property of any single agent but of the system as a whole (levin 1998, 1999; snowden & stanbridge 2004). the second challenge relates to the multistakeholder nature of common-property natural resources; even fenced-off protected areas are increasingly influenced by external social issues (venter et al. 2008). these stakeholders may subscribe to widely varying world views, based on different values and knowledge forms, with expectations that play out over different time horizons and spatial scales. under these circumstances, management should probably avoid targeting an optimal solution for ‘the (single) problem’, but should adopt an ongoing learning and negotiation process where mutual sense-making and adaptation are prioritised (pahl-wostl & hare 2004). this reality has lead some authors and practitioners to coin the term ‘adaptive co-management’, as a descriptor of adaptive management that explicitly caters for mutual learning and cooperation between stakeholders such as conservation agencies, researchers and local communities (armitage et al. 2007; borrini-feyerabend et al. 2000; olsson, folke & berkes 2004; ruitenbeek & cartier 2001). the presence of limited predictability (or a certain level of irreducible uncertainty), as well as multiple stakeholders with frequent conflicting interests, suggests that there are two fundamental conditions necessary for effective management of natural resources, (1) to learn and adapt and (2) to do so purposefully with relevant partners. sam, which was initially developed in the context of managing rivers and their catchment areas (biggs & rogers 2003; rogers & bestbier 1997; rogers & biggs 1999), provides a framework for facilitating such learning. it incorporates the iterative learning dimension of adaptive management and the mutual learning dimension of co-management. in addition, it emphasises a forward-looking dimension, hence the reference to ‘strategic’. in summary, as grant et al. (2008) point out, sam is designed to be strategic (facilitate action with foresight and purpose), adaptive (facilitate learning whilst we are doing) and participatory (facilitate engagement and empowerment of stakeholders). sam is a modular process that allows practitioners to start with any of the five broad steps in the adaptive cycle and to expand their efforts from there. the five steps can be grouped into three interrelated subprocesses or components that have taken shape in the application of sam, namely adaptive planning, adaptive implementation and adaptive evaluation (figure 1). in the following sections we introduce the basic steps of sam in more detail. figure 1: schematic summary of the steps in the strategic adaptive management process, based on the work of biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience. ecology and management of savanna heterogeneity, pp. 59-80, island press, washington dc; pollard, s.r. & du toit, d.r., 2007, guidelines for strategic adaptive management – experiences from managing the rivers of the kruger national park. guidelines of unep/gef project no. gf/27-13-03-4679. ecosystems, protected areas and people project; and scholes, r.j. & mennell, k.g., 2008, ‘summary for policymakers’, in r.j. scholes & k.g. mennell (eds.), elephant management: a scientific assessment for south africa, pp. 1-21, wits university press, johannesburg. adaptive planning the adaptive planning process of sam is seen increasingly as a critical condition for achieving its successful implementation (rogers pers. comm., 14 september 2010). the aim of this process is to build a sense of common purpose amongst all relevant stakeholders and to develop a collective roadmap for getting from a current (usually undesirable) reality to a more desirable social-ecological system. stakeholder inclusivity is vital to the success of an adaptive planning process. in the case of sanparks, stakeholders include park managers, scientists, government policymakers, agency managers, wildlife activists, traditional communities neighbouring parks, farmers that share catchments of rivers that flow through parks, ngos and ecotourists. successful adaptive planning depends on the facilitation of a constructive dialogue amongst these stakeholders with their diverse and often divergent values, expectations, professional norms and reward systems (rogers & breen 2003). the first step in adaptive planning is to create a common vision in which stakeholders agree on the social, technical, economic, ecological and political contexts of the system to be managed. a critical part of this visioning exercise is to reach agreement on values, or operating principles, which should guide management decision making in the future. the v-steep (values, social, technical, economic, ecological and political) framework that emerges provides an approach to describing the context as comprehensively as possible (see pollard & du toit 2007). a further part of visioning is to deliberate and reach consensus on the vital attributes of the system to be managed and their determinants. vital attributes, as perceived by stakeholders, are the distinctive and special features of the social-ecological system of concern that are the key to its management (rogers & bestbier 1997). a vision statement is formulated on the basis of this understanding of the context and values. the vision, together with the vital attributes of the system to be managed, informs the setting of objectives. a nested hierarchy of objectives starts with high-level objectives that are set, firstly, to ensure the maintenance of the identified vital attributes of the system to be managed and, secondly, to overcome the constraints and threats to meeting the vision. through a step-by-step process, these high-level objectives (which are largely based on stakeholder values) are deconstructed into a series of objectives of increasing detail until they represent measurable, scientifically credible endpoints. the result is referred to as an objectives hierarchy (figure 2). acknowledging the dynamic nature of ecosystems, the measurable targets (figure 2) describe the boundaries of the desired state (as opposed to an optimal value). these boundaries are also referred to as thresholds of potential concern (tpcs) and are essentially hypotheses of the outer limits of acceptable change. these ‘boundaries’ are acceptable changes in the attribute of interest, embedded within specific temporal and spatial scales (see foxcroft & mcgeoch 2011). as such, their validity and appropriateness remains open to challenge and tpcs are revised as understanding of the system improves (pollard & du toit 2007). tpcs are developed in collaboration between managers, scientists and field staff responsible for monitoring. the last step that forms part of the adaptive planning process is to scope or analyse various options for achieving the objectives that were derived in the previous step. importantly, this step is still conducted in cooperation with stakeholders. different options are identified, their likely consequences predicted and the acceptability of those consequences assessed. finally, a combination of management options that provide the best potential social-ecological system outcomes and learning opportunities is selected for implementation (grant et al. 2008; pollard & du toit 2007). figure 2: generic objectives hierarchy that links the vision to scientific endpoints, based on the work of pollard, s.r. & du toit, d.r., 2007, guidelines for strategic adaptive management – experiences from managing the rivers of the kruger national park. guidelines of unep/gef project no. gf/27-13-034679. ecosystems, protected areas and people project; rogers, k. & bestbier, r., 1997, development of a protocol for the definition of the desired state of riverine systems in south africa, department of environmental affairs and tourism, pretoria; and rogers, k.h. & biggs, h.c., 1999, ‘integrating indicators, endpoints and value systems in strategic management of the kruger national park’, freshwater biology 41, 439-451. doi:10.1046/j.1365-2427.1999.00441.x. adaptive implementation adaptive implementation entails incorporating the options that were selected in the previous step as part of the operating procedures and business routines of the relevant organisation(s). this requires the development of detailed action plans, allocation of the necessary resources and the implementation of those plans. a key component of the adaptive implementation of action plans is to develop monitoring protocols to describe the subject and focus of what to monitor and establish the frequency at which to do so. monitoring endpoints are linked to the measurable targets (or tpcs) in figure 2. part of the ‘new’ management procedures is to establish a forum for the regular evaluation of monitoring results against set tpcs, as well as standard procedures for dealing with tpc excedance and for capturing and sharing learning (pollard & du toit 2007). adaptive evaluation one of the main purposes of sam (and adaptive management in general) is to purposefully learn and adapt over time. therefore, it is essential that learning becomes an explicit step in the strategic adaptive management process (figure 1). however, learning should not be seen as a mere step to be taken at the end of the process, but should rather occur throughout the planning and implementation phases via a series of feedback loops. continuous evaluation and learning is facilitated by reflecting on the following questions (figure 3): • is the monitoring adequate, cost effective and feasible? • has the intended plan of operation materialised? • were the selected options appropriate? • were the predicted consequences correct and, if not, why? • were the consequences actually acceptable? • even if the predicted consequences were correct and are acceptable, are the objectives and vision being met? figure 3: the adaptive management process with feedback loops for ongoing reflective learning at multiple points during the process. roadmap through this special issue (back to top) the papers contained in this special issue were solicited to capture and share as much as possible of the experience gained by sanparks during the development and implementation of sam to date. the authors were allowed a fair degree of freedom to ensure that a broad spectrum of perspectives and diverse lessons are captured in the special issue. we refer to the papers in this issue as essays rather than research papers as some of the authors use a narrative writing style to convey their understanding of sam. the papers in the special issue can be divided into two parts. part one consists of papers of a more systemic, generic or philosophical nature, whilst the papers in part two focus largely on thresholds of potential concern and how these thresholds were derived and applied for specific target indicators. whilst each paper can be read on its own, the papers from part one and part two are meant to be complementary. part one starts off with a detailed contextual setting of the kruger national park as the main study area for the development of sam (pollard et al. 2011). the authors of this essay provide an historical overview of how changes in management paradigms eventually led to the adoption of sam as the approach of choice, as well as how this approach was pioneered in the sphere of river management. in the next essay, biggs, breen, slotow, freitag-ronaldson and hockings (2011) examine the relationship between assessment and reflection, focusing on how these processes can be used in a complementary way to catalyse learning for adaptive management. stirzaker et al. (2011) allude to the often contradicting nature of the entrained behaviour of many scientists and managers alike, as well as the behaviour required to participate effectively in an adaptive management process. they explore the shortcomings and requirements of organisations in terms of enabling adaptive management. in the fourth essay, holness and biggs (2011) address a question that is of critical importance to a conservation agency: are systematic conservation planning and adaptive management compatible processes? these authors argue that systematic conservation planning should be practiced as an intrinsic part of a broader adaptive management approach and suggest how such a marriage can be achieved. systematic monitoring is a key factor in our ability to learn and adapt. to this end, mcgeoch et al. (2011) propose a framework for biodiversity monitoring which would address the biodiversity objectives as outlined in the management plans of south african national parks. gaylard and ferreira (2011) reflect on how the process of sam itself has been adapted in response to various implementation challenges. the final essay in part one of this issue provides a transition to the part two papers. biggs, ferreira, freitag-ronaldson and grant-biggs (2011) provide a critical assessment of the usefulness of the concept of thresholds of potential concern. these authors propose a reconceptualisation of the tpc concept, based on learning over a period of one decade, to increase its utility within the sam process. part two contains a collection of 11 essays, each focusing on a specific theme. these essays aim to summarise the development of tpcs for the particular theme and assess how this fits into the broader strategic adaptive management approach. these areas have not received the same amount of attention over the last decade. therefore, whilst some papers will provide an in-depth discussion of their evolution, others will present recent developments and provide suggestions for future directions. as the pioneering work on sam in the kruger national park was conducted on river management, mcloughlin et al. (2011) take us through a journey explaining the history and rationale behind the tpc concept. furthermore, they show how the tpcs for river management were used and how they evolved as learning progressed. fire research and management in sanparks has been an ongoing process for a number of decades, with a number of different approaches being implemented, both within and across parks nationwide. van wilgen et al. (2011) discuss the changes from ‘trial and error’ through to ‘active adaptive management’, indicating the development of various forms of thresholds. another area where adaptive management approaches and tpcs have developed over a number of years is with invasive alien species. here foxcroft and mcgeoch (2011) aim to link the management actions, monitoring programme, research efforts and tpcs in order to develop functional feedback mechanisms and enable improved management and learning. whilst not discussing the development of tpcs specifically, a number of essays propose approaches to adopting an adaptive management strategy. they also suggest tpcs and aim to test the use of these in particular cases. for example, grant et al. (2011) evaluate herbivore–vegetation interactions and how determining thresholds for these can prevent unacceptable changes in desired vegetation states and patterns. mcgeoch et al. (2011) deal with approaches to monitoring the often neglected terrestrial and freshwater biodiversity. similarly, except for some specific case studies, resource use management has been given little attention until recently. in their essay, scheepers et al. (2011) provide an overall framework for applying adaptive approaches to resource use management. they illustrate this with three case studies, covering a range of approaches and timeframes, and conclude with opportunities for future expansion. as global environmental change intensifies, one of the most pressing issues that conservation and protected area managers face is ensuring the persistence of rare species (rebelo et al. 2011). however, in areas with high endemicity, and which are facing a number of potentially negative impacts, assessing which species are most in need of special attention is problematic. for example, table mountain national park has 307 threatened iucn red list (plus 208 non-least concern) and 332 endemic terrestrial plant and animal species. rebelo et al. (2011) present an approach to dealing with this challenging problem.four essays probe some underlying philosophical or technical issues, including the idea that behind all good science lies good science support. this is an essential but underrated part of the overall science management and monitoring partnership. kruger and macfadyen (2011) discuss a number of innovations that deal with these issues, from collecting, managing and automating data management, to developing systems to report back on tpcs. a challenge with implementing tpcs is determining at what stage the breach of a tpc is triggered and how the lag effects of this breach are handled. scholes and kruger (2011) present a potential approach to this, illustrating it with an example from the kruger national park. owing to the different needs and analytical approaches necessary to implementing tpcs and management across the sanparks estate, ferreira et al. (2011) use conceptual linkages between objectives, indicators, mechanisms and modulators to help identify key concerns in relation to management objectives. based on these linkages, the underlying mechanisms responsible for the management concern may be evaluated. conclusion (back to top) the dominant message that emerges from the papers in this special issue is that adaptive management is about structured learning. the authors reflect on various mechanisms that are used to make current assumptions and understanding explicit so that relevant stakeholders can learn in a structured way. these mechanisms include co-creating a desired state or vision, setting objectives, formulating thresholds of potential concern, and monitoring and evaluating the consequences of management decisions. this volume of papers represents a comprehensive documentation and reflection of this process, after applying the principles of sam across an entire conservation agency for over 10 years. acknowledgements (back to top) we thank dr richard stirzaker, prof. kevin rogers and dr harry biggs for discussions and comments during the development of this editorial. references (back to top) armitage, d., berkes, f. & doubleday, n., 2007, adaptive co-management: collaboration, learning, and multi-level governance, ubc press, vancouver. berkes, f., colding, j. & folke, c. (eds.), 2003, navigating social-ecological systems: building resilience for complexity and change, cambridge university press, cambridge. biggs, h., ferreira, s., freitag-ronaldson, s. & grant-biggs, r., 2011, ‘taking stock after a decade: does the ‘thresholds of potential concern’ concept need a socio-ecological revamp?’ koedoe 53(2), art. #1002, 9 pages. doi:10.4102/koedoe.v53i2.1002 biggs, h.c., breen, c., slotow, r., freitag, s. & hockings, m., 2011, ‘how assessment and reflection relate to more effective learning in adaptive management’, koedoe 53(2), art. #1001, 13 pages.doi:10.4102/koedoe.v53i2.1001 biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience. ecology and management of savanna heterogeneity, pp. 59−80, island press, washington dc. borrini-feyerabend, g., farvar, m.t., nguinguiri, j.c. & ndangang, v., 2000, co-management of natural resources: organizing negotiation and learning by doing, kasparek zverlag, heidelberg. ferreira, s., deacon, a., sithole, h., bezuidenhout, h., daemane, m. & herbst, m., 2011, ‘from numbers to ecosystems and biodiversity: a mechanistic approach to monitoring’, koedoe 53(2), art. #998, 12 pages. doi:10.4102/koedoe.v53i2.998 folke, c., hahn, t., olsson, p. & norberg, j., 2005, ‘adaptive governance of social-ecological systems’, annual review of environmental resources 30, 441−473. doi:10.1146/annurev.energy.30.050504.144511 foxcroft, l.c. & mcgeoch, m., 2011, ‘implementing invasive species management in an adaptive management framework’, koedoe 53(2), art. #1006, 11 pages. doi:10.4102/koedoe.v53i2.1006 gaylard, a. & ferreira, s., 2011, ‘advances and challenges in the implementation of strategic adaptive management beyond the kruger national park – making linkages between science and biodiversity management’, koedoe 53(2), art. #1005, 8 pages. doi:10.4102/koedoe.v53i2.1005 grant, r., sherwill, t., rogers, k., biggs, h., freitag, s., hofmeyr, m., et al., 2008, a framework for developing and implementing management plans for south african national parks, south african national parks, pretoria. grant, r.c.c., peel, m.j.s. & bezuidenhout, h., 2011, ‘evaluating herbivore management outcomes and associated vegetation impacts’, koedoe 53(2), art. #1008, 15 pages. doi:10.4102/koedoe.v53i2.1008 holling, c.s., 2001, ‘understanding the complexity of economic, ecological and social systems’, ecosystems 4, 390−405. doi:10.1007/s10021-001-0101-5 holness, s.d. & biggs, h.c., 2011, ‘systematic conservation planning and adaptive management’, koedoe 53(2), art. #1029, 9 pages. doi:10.4102/koedoe.v53i2.1029 johnson, b.l., 1999, ‘introduction to the special feature: adaptive management – scientifically sound, socially challenged?’, conservation ecology 3(1), 10, viewed n.d., from http://www.consecol.org/vol3/iss1/art10/ kruger, j.m. & macfadyen, s., 2011, ‘science support within the south african national parks adaptive management framework’, koedoe 53(2), art. #1010, 7 pages. doi:10.4102/koedoe.v53i2.1010 lee, k., 1993, compass and gyroscope: integrating science and politics for the environment, island press, washington dc. levin, s.a., 1998, ‘ecosystems and the biosphere as complex adaptive systems’, ecosystems 1, 431−436. doi:10.1007/s100219900037 levin, s.a., 1999, fragile dominion: complexity and the commons, perseus publishing, cambridge. mcgeoch, m.a., dopolo, m., novellie, p., hendriks, h., freitag, s., ferreira, s., et al., 2011, ‘a strategic framework for biodiversity monitoring in south african national parks’, koedoe 53(2), art. #991, 10 pages. doi:10.4102/koedoe.v53i2.991 mcgeoch, m.a., sithole, h., samways, m.j., simaika, j.p., pryke, j.s., picker, m., et al., 2011, ‘conservation and monitoring of invertebrates in terrestrial protected areas’, koedoe 53(2), art. #1000, 13 pages. doi:10.4102/koedoe.v53i2.1000 mclain, r.j. & lee, r.g., 1996, ‘adaptive management: promises and pitfalls’, environmental management 20(4), 437−448. doi:10.1007/bf01474647, pmid:8661620 mcloughlin, c.a., deacon, a., sithole, h. & gyeduababio, t., 2011, ‘history, rationale, and lessons learned: thresholds of potential concern in kruger national park river adaptive management’, koedoe 53(2), art. #996, 27 pages. doi:10.4102/koedoe.v53i2.996 meffe, g.k., nielsen, l.a., knight, r.l. & schenborn, d.a., 2002, ecosystem management: adaptive community-based conservation, island press, washington dc. oglethorpe, j., 2002, adaptive management: from theory to practice, world conservation union, gland. ohlsson, l., 1999, environment, scarcity and conflict: a study of malthusian concerns, phd thesis, university of gothenburg. olsson, p., folke, c. & berkes, f., 2004, ‘adaptive co-management for building resilience in social-ecological systems’, environmental management 34(1), 75−90. doi:10.1007/s00267-003-0101-7,pmid: 15383875 pahl-wostl, c. & hare, m., 2004, ‘processes of social learning in integrated resources management’, journal of community and applied social psychology 14, 193−206. doi:10.1002/casp.774 pollard, s.r. & du toit, d.r., 2007, guidelines for strategic adaptive management – experiences from managing the rivers of the kruger national park. guidelines of unep/gef project no. gf/27-13-03-4679. ecosystems, protected areas and people project, pp. 1–35. pollard, s., du toit, d. & biggs, h., 2011, ‘river management under transformation: the emergence of strategic adaptive management of river systems in the kruger national park’, koedoe 53 (2), art. #1011, 14 pages. doi:10.4102/koedoe.v53i2.1011 rebelo, t.g., freitag, s., cheney, c. & mcgeoch, m.a., 2011, ‘prioritising species of special concern for monitoring in table mountain national park: the challenge of a species-rich, threatened ecosystem’, koedoe 53(2), art. #1019, 14 pages. doi:10.4102/koedoe.v53i2.1019 rogers, k., 2003, ‘adopting a heterogeneity paradigm: implications for management of protected savannas’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience. ecology and management of savanna heterogeneity, pp. 41−58, island press, washington dc. rogers, k.h., roux, d.j. & biggs, h.c., 2000, ‘challenges for catchment management agencies. lessons from bureaucracies, business and resource management’, water sa 26, 505−512. rogers, k. & bestbier, r., 1997, development of a protocol for the definition of the desired state of riverine systems in south africa, department of environmental affairs and tourism, pretoria. rogers, k.h. & biggs, h.c., 1999, ‘integrating indicators, endpoints and value systems in strategic management of the kruger national park’, freshwater biology 41, 439−451. doi:10.1046/j.1365-2427.1999.00441.x ruitenbeek, j. & cartier, c., 2001, the invisible wand: adaptive co-management as an emergent strategy in complex bio-economic systems. occasional paper no. 34. center for international forestry research, bogor. scheepers, k., swemmer, l. & vermeulen, w.j., 2011, ‘applying adaptive management in resource use in south african national parks: a case study approach’, koedoe 53(2), art. #999, 14 pages. doi:10.4102/koedoe.v53i2.999 scholes, r.j. & kruger, j.m., 2011, ‘a framework for deriving and triggering thresholds for management intervention in uncertain, varying and time-lagged systems’, koedoe 53(2), art. #987, 8 pages. doi:10.4102/koedoe.v53i2.987 scholes, r.j. & mennell, k.g., 2008, ‘summary for policymakers’, in r.j. scholes & k.g. mennell (eds.), elephant management: a scientific assessment for south africa, pp. 1−21, wits university press, johannesburg. shea, k., possingham, h.p., murdoch, w.w. & roush, r., 2002, ‘active adaptive management in insect pest and weed control: intervention with a plan for learning’, ecological applications 12(3), 927−936. doi:10.1890/1051-0761(2002)012[0927:aamiip]2.0.co;2 snowden, d. & stanbridge, p., 2004, ‘the landscape of management: creating the context for understanding social complexity’, emergence: complexity and organization 6(1–2), 140−148. stirzaker, r.j., roux, d.j. & biggs, h.c., 2011, ‘learning to bridge the gap between adaptive management and organisational culture’, koedoe 53(2), art. #1007, 6 pages. doi:10.4102/koedoe.v53i2.1007 swemmer, l.k. & taljaard, s., 2011, ‘sanparks, people and adaptive management: understanding a diverse field of practice during changing times’, koedoe 53(2), art. #1017, 7 pages. doi:10.4102/koedoe.v53i2.1017 van wilgen, b.w., govender, n., forsyth, g.g. & kraaij, t., 2011, ‘towards adaptive fire management for biodiversity conservation: experience in south african national parks’, koedoe 53(2), art. #982, 9 pages. doi:10.4102/koedoe.v53i2.982 venter, f.j., naiman, r.j., biggs, h.c. & pienaar, d.j., 2008, ‘the evolution of conservation management philosophy: science, environmental change and social adjustments in kruger national park’, ecosystems 11, 173−192. doi:10.1007/s10021-007-9116-x walters, c., 1997, ‘challenges in adaptive management of riparian and coastal ecosystems’, conservation ecology 1(2), 1, viewed n.d., from http://www.consecol.org/vol1/iss2/art1/ vol. 50 no. 1 pp. 154 159koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research travel motivations of tourists visiting kruger national park abstract the kruger national park (knp) one of the largest conservation areas in south africa, attracts in excess of one million visitors a year and is regarded as an icon destination in international tourism. since this park attracts more tourists than any other park in south africa, the purpose of this article is to determine the reasons (the travel motives) why tourists visit the park. little research has been done on travel motives to national parks and this was the first of its kind in south africa. the research was conducted by means of questionnaires. a factor analysis was used to determine the travel motives. six factors were identified, namely nature, activities, attractions, nostalgia, novelty and escape from routine. some of these motives were confirmed by similar research in other countries, although the similarities are not significant. this research confirmed that different attractions and destinations fed different travel motives, hence the need for more studies of this nature to be conducted. keywords: travel motives, ecotourism, wildlife tourism, national parks, destinations. the kruger national park (knp) was formally established in 1926, with the amalgamation of the sabi and the singwitsi game reserves. the reason behind the conception of the park was to stop uncontrolled hunting in this area. therefore, the main purpose was that of conservation. however, at the time of the proclamation of the kruger national park in 1926, the idea of tourism was already well established (joubert 2007; pienaar 2007). since then, tourism has developed and currently the park attracts in excess of one million tourists or visitors per year. it is one of the top five international tourist destinations in south africa. even though the park is a top international attraction, most tourists (approximately 80%) are local tourists. this confirms that the park has successfully completed the transition from a pure conservation area to a tourist attraction of international significance that both benefits tourism and makes a contribution to the local community and its economy. in highlighting the importance of national parks, uysal, mcdonald and martin (1994) state that national parks and natural areas are powerful magnets to tourists, and that these attractions are major export earners. in support of the latter, saayman and saayman (2006) found that the kruger national park (figure 1) generated approximately r1,5 billion for the region annually. eagles and mccool (2002) stated that the kruger national park had developed a significant international profile over time and is now the anchor of the wildlife-tourism industry in south africa. scenic beauty and wildlife remain major tourism attractions (ecotourism) for both international and national tourists, who make up 80% of the tourist numbers in or to south africa (gcis 1998; gcis 1999). however, south africa is one of many countries or destinations worldwide that offers this type of tourism product. as countries and destinations strive to increase their share of the international and national tourism market, it becomes important to understand why people travel and why they choose a specific destination (oh, uysal and weaver 1995). therefore, the purpose of this study is to determine the reasons (the travel motives) why tourists visit the kruger national park. by determining the motives, according to saayman (2006), marketing can be undertaken more effectively and specific factors can be taken into account when marketing strategies are being planned. this article is organised as follows: the literature review follows the introduction, then the method of research, which is followed by the results of the research, after which certain conclusions are drawn and recommendations are made. literature review and problem statement in 1994, fodness (1994) stated that effective tourism marketing is impossible without an understanding of the consumers’ motivations, or, to put it differently, to know the answer to the question of what motivates people to travel. the aim of marketing, according to saayman (2006), is the effective and efficient use of resources in the changing environment of today in order to ensure a profit, survival, and growth of the tourism organisation destination. fodness (1994) adds that further insights into tourists’ travel motivation can benefit tourism marketing, specifically with regard to product development, service quality evaluation, image development and promotional activities. mill and morrison (1985) support the notion that motivation plays a very important role in the process of travelling, vacation, and when visiting friends and relatives. motivation comes into play when a person wants to satisfy a need and must take action to do so. mill and morrison (1985) state further that the behaviour of tourists is influenced by a small number of factors, and a person can be motivated by more than one motive at a time. a literature review on travel motivation revealed a wide variety of motivations and a great number of publications, the following of which are merely a few examples: • jang and wu (2006) studied the travel motivations of taiwanese seniors • oh et al. (1995) investigated australian tourists’ travel motivations • tao, eagles and smith (2004) looked into motivations of asian tourists travelling to taiwan’s taroko national park • awaritefe (2004) looked into tourists’ travel motivations to nigeria • fodness (1994) looked into travel motivations for leisure travel peet van der merwe melville saayman institute for tourism and leisure studies north-west university south africa correspondence to: peet van der merwe e-mail: peet.vandermerwe@nwu.ac.za postal address: private bag x6001, north-west university, potchefstroom 2520, south africa 154 2008 travel motivations of tourists original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 154 159 • correia, oom do valle and moço (2007) looked into tourists’ travel motivations to exotic places • uysal et al. (1994) looked into travel motivations of australian tourists to us national parks and nature areas • bansal and eiselt (2004) looked into tourists’ travel motivations to canadian maritime province • swanson and horridge (2006) looked into travel motivations influencing the type of souvenirs tourists purchase • kim, borges and chon (2006) investigated travel motivations of visitors to festivals • backman, backman, uysal and sunshine (1995) examined dimensions of event motivations • schneider and backman (1996) examined the crosscultural equivalence of a motivation scale • lee, lee and wicks (2004) identified the underlying dimensions of motivations of visitors attending the 2000 world culture expo • yoon and uysal (2005) investigated tourists’ travel motivations in general based on an analysis of the findings of the above authors, it is clear that tourists have different motives for visiting different attractions and/or destinations, as can be seen from table 1. the literature review also clearly revealed that very little research has been done on travel motives to national parks. studies focusing on the latter are those of uysal et al. (1994), tao et al. (2004), and awaritefe (2004). uysal et al. (1994) conducted research on australian tourists visiting national parks and natural areas in the united states. five factors were identified, namely, “relaxation/hobbies,” “novelty,” “enhancement of kinship relations,” “escape” and “prestige”. tao et al. (2004) focused on asian tourists visiting taroko national park in taiwan. two factors have been identified, namely, “learning about nature” and “participating in recreation activities”. awaritefe (2004) conducted research on tourists visiting parks in nigeria. the most significant motives were self-actualisation, an educational or cultural context, and recreational or leisure pursuits. after the three studies conducted on visitors to national parks as indicated above have been compared, the results are still not comparable. this shows that different visitors to different parks have quite different motives. this could be because of (1) what these parks offer, (2) where they are located, (3) the type of market and (4), the type of activities, to name but a few. therefore, the lack of research done at national parks in general and the fact that different parks revealed different results was the motivation underlying this research. added to this is the fact that never before has this type of research been conducted at national parks in south africa. uysal et al. (1994) add to this understanding that, in order to market effectively, it is essential to generate more specific knowledge about visitors to parks and natural areas. oh et al. (1995) conclude that if countries strive to increase their share of visitors, it becomes essential to understand why people travel and why they choose a specific destination. method of research since the data used in the analysis were gathered by means of consumer-based questionnaires over a period of seven years, namely 2001 to 2007, the methodology used will be discussed under the following headings: (i) the questionnaire, (ii) the samples, and (iii) the method. the questionnaire the questionnaire used to survey visitors to the kruger national park remained similar throughout the period (2001–2007) of data collection and consisted of three sections. in section a, demographic details were surveyed while section b focused on spending behaviour (marital status, age, province of origin, number of persons paid for, frequency of visits, length of stay and amount spent) and motivational factors. the section on travel motivations was based on the work of crompton (1977) and was adapted for national parks. section c of the questionnaire consisted of more detailed information about the consumer’s general behaviour (type of magazines or newspapers they read and their catering preferences, for example). for the purposes of this article, the information obtained from sections a and b is predominantly used. the samples surveys at the kruger national park were conducted annually between 2001 and 2005. since 2006, two surveys were conducted annually, in winter and in summer. in table 2, the figure 1 kruger national park (source: african safari travel, 2007) 155 original research van der merwe & saayman koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 154 159 sample sizes and the different camps where the surveys were conducted are given, and it is evident that the sample size has grown significantly over the past years. all visitors to the camp received a questionnaire that they completed in their own time. field workers collected the questionnaires again during the evenings and early mornings. note that the definition of a tourist is applied for the purpose of this research and therefore only overnight visitors have been considered in the analyses. a total of 2 899 questionnaires were administrated and, according to saayman and fouché (2007), the profile of visitors has remained similar since 2001. hence it is believed that the sampling is representative of the population. the method microsoft excel was used for data capturing and basic data analysis. the factor analysis was conducted by means of spss (2006) and oblique promax with a rotation oblimin method was used. to explain the variance-covariance structure of a set of variables through a few linear combinations of these variables, a principal component analysis can be applied. the aim of such an analysis is to (i) reduce the data and (ii) to assist in the interpretation of the data. although p components are required to explain the total variability in the system, much of this variability can normally be accounted for by a small number, k, of principal components. there is almost as much information in k components as there is in the initial p variables. the k components can thus replace the p variables, thereby reducing the data set. the analysis often reveals relationships that were not previously suspected and thereby allows interpretations that would not ordinarily result (johnson & wichern, 2002: 426). factor analysis extends a principal component analysis, since both can be viewed as attempts to approximate the covariance matrix. however, the approximation based on the factor analysis model is much more elaborate than that of a principal component analysis. although factor analysis has its origin in the measurement of intelligence, it can be applied to most disciplines with success. the purpose of a factor analysis is to describe the covariance relationships among many variables in terms of a few underlying, but unobservable, random quantities called factors. the factor model can be motivated by the following argument: suppose that variables can be grouped according to their correlations. that is, all variables in a particular group are highly correlated among themselves, but small correlations with variables in a different group. if this is researcher/s travel motives backman, backman, uysal and sunshine (1995) • excitement • external • family • socialising • relaxation schneider and backman (1996) • family togetherness • socialisation • social/leisure • festival attributes • escape • event excitement lee, lee and wicks (2004) • cultural exploration • family togetherness • novelty • escape • event attractions • socialisation fodness (1994) • knowledge function • utilitarian function (punishment, minimisation) • value expressive function (self-esteem) • value expressive function (self-enhancement) • utilitarian function (reward maximisation) jang and wu (2006) pull factors: • cleanliness and safety • facilities, events and costs • natural and historic sites push factors: • ego-enhancement • self-esteem • knowledge seeking • relaxation • socialisation yoon and uysal (2005) • excitement • knowledge and learning experience • relaxation • achievement • family togetherness • escape • safety • fun oh et al. (1995) • safety/comfort seekers • culture/history seekers • novelty/adventure seekers • luxury seekers kim et al. (2006) • family togetherness • socialisation • site attraction • festival attraction • escape from routine swanson and horridge (2006) internal motivators: • desire for escape • rest • relaxation • prestige • health and fitness • adventure • social interaction external motivators: • attractiveness of the destination • tangible resources (beaches, recreational activities and cultural attractions) • travellers’ perceptions and expectations (novelty, benefit expectations and marketing image) bansal and eiselt (2004) • climate • relaxation • adventure • personal • education • sites and festivals correia, oom de valle and moço . (2007) the research revealed three push factors/motives: • knowledge • leisure • socialisation the pull factors that were revealed were: • facilities • core attractions • landscape features loker and perdue (1992) • excitement and escape • adrenalin, excitement seeking • family and friends-oriented • naturalist (those who enjoyed nature surroundings) • escape (those who valued the escape by itself) table 1 analysis of research on travel motives researcher/s travel motives goeldner, ritchie and mcintosh (2000) • spirituality • social status • escape • cultural enrichment crompton (1977) identified seven socio-psychological motives for travelling: • escaping from the everyday environment • discovering and evaluating of oneself • recreation and travelling • status • regression • strengthening of family ties • facilitation of social interaction weaver and oppermann (2000) identified three categories: • leisure • visiting friends and relatives • business they can be subdivided into the following four categories: • relaxation and recreation • sport and health • religion • education table 1 (cont...) 156 travel motivations of tourists original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 154 159 the case, it is conceivable that each group of variables represents a single underlying factor that is responsible for the observed correlations. it is this type of structure that a factor analysis seeks to confirm (johnson & wichern 2002). results the results will be discussed in two sections. firstly, an overview of the profile for visitors to the kruger national park will be given, and, secondly, the results of the factor analysis (visitor motives) will be discussed. four surveys were conducted during the summer season (november/december) and four during the winter season (may/july). due to this, there were minor differences between the summer and winter profile of visitors such as province of origin (see table 3) and length of stay – tourists tend to stay longer during winter season, which leads to increased spending during the winter seasons. based on the results captured in table 3, tourists visit national parks at least twice a year. these tourists are loyal to this particular park and they enjoy the wildlife that the park offers. in addition, they want to relax. these tourists are predominantly afrikaans speaking (one of south africa’s eleven official languages), and are approximately 45 years old, coming mainly from gauteng and the western cape province. visitors to this park are well educated and they stay for an average of one week. the descriptive results showed that “to relax” was regarded as the most important reason for visiting the park. the next section will deal with the motives for travelling as indicated by the factor analysis. the factor analysis (pattern matrix) identified six factors. based on the items included in the identified factors, these factors can be named accordingly from table 4: nature, activities, attractions, nostalgia, novelty, and escape from routine. • factor 1: nature nature included aspects such as “to see endangered species,” “to see animals,” “to see plants,” “for educational reasons,” “to take photos of animals” and “to take photos of plants”. this factor is confirmed by oh et al. (1995) as well as swanson and horridge (2006) as a motive for travel. it should be noted, however, that none of those studies were conducted at national parks. nature as a factor has a mean value of 3.0062, which is the third highest of the six factors. • factor 2: activities activities include sub-categories such as “to attend conferences,” “to attend events” and “hiking”. oh et al. (1995) also found that activities such as nightlife, exotic atmospheres, and amusement or theme parks play an important role in the travel motivations of tourists. of the six factors, this one scored the lowest mean value of 1.5392. • factor 3: attractions attractions include “accommodation,” “brand of the park,” “climate of location” and “grew up with the park”. kim et al. (2006) also indicated that “attractions” is an important travel motivator. this factor has a mean value of 2.8072. • factor 4: nostalgia nostalgia consists of “family time”, “park is visited from childhood days”, “to experience wildlife”, “family time”, and “different species”. this aspect achieved the second highest mean value of 3.4352. none of the other research projects consulted during the literature review identified this as a motive for travelling. • factor 5: novelty novelty consists of “explore new destinations” and “socialising with friends”. the same travel motive was identified by uysal et al. (1994) for australian tourists visiting national parks in the united states of america. this motive has also been identified in research by oh et al. (1995) and lee et al. (2004) and has a mean value of 2.7724. • factor 6: escape from routine factor 6 consists of “routine vacation” and “to relax”. uysal et al. (1994) and kim et al. (2006) found that “escape from daily routine” was an important motive for tourists to travel. kim et al. (2006) further indicated that “escape” includes aspects such as “to get away from demands of life” and “daily routine”. swanson and horridge (2006) also identified “escape” as an important travel motivator, and included aspects such as “seeing spectacular scenery,” year 2001 2002 2003 2004 2005 2006 2006 2007 survey month may july december december december july november june # of questionnaires 220 323 246 400 450 476 171 613 camps 78 berg en dal 68 satara 40 olifants 34 shingwedzi 62 berg en dal 87 satara 93 olifants 81 shingwedzi 20 berg en dal 75 satara 21 olifants 66 lower sabie 64 skukuza 70 berg en dal 84 satara 39 olifants 72 lower sabie 135 skukuza 57 berg en dal 128 satara 79 letaba 63 lower sabie 128 skukuza 19 malelane 74 pretoriuskop 249 skukuza 49 olifants 85 letaba 36 letaba 55 skukuza 80 satara 161 berg en dal 173 satara 191 skukuza 88 letaba table 2 total number of questionnaires completed – 2001 to 2007 category profile home language afrikaans (70%) age 35–49 years of age (average:44,5) marital status married (84%) province of residence gauteng and western cape predominantly during winter months mpumalanga and gauteng during summer months level of education diploma/degree (80%) number of people paid for 3–4 people mode of transport sedan and 4x4 vehicle number of visits to national parks over 3 years 6 times (2 per year) length of stay 4–14 days during winter and 2-7 days during summer preference of the park wildlife reasons for visiting the park to relax expenditure r6 000 per trip (winter months) r5 000 per trip (summer months) visitor references self-catering preferred newspaper rapport and beeld preferred magazines weg, huisgenoot/you, getaway/wegbreek preferred radio stations jacaranda and highveld stereo preferred tv programmes sport, nature and news heard about the park word-of-mouth preferred accommodation chalets table 3 visitors profile: kruger national park 2001–2007 157 original research van der merwe & saayman koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 154 159 “visiting places i’ve never seen before” and “visiting exiting places”. this factor scored the highest mean value of the six factors, namely 4.2133, and is the most common motive for travel if one compares the various research projects in table 1. based on the results of the component correlation matrix as captured in table 5, the low correlation between the different factors shows that the factors can be clearly distinguished. the motives of tourists visiting the kruger national park are thus very specific and well defined. the table indicates that tourists visiting the park to experience nature have similar motives to those of tourists visiting the park for activities or to escape from routine, for example. based on this research, the next section will discuss the findings and conclusions. findings and conclusions the aim of this article was to determine the travel motives of tourists visiting the kruger national park in south africa. this was the first time that this type of research was conducted in a national park in south africa. the results of the survey revealed six travel motives, namely nature, activities, attractions, nostalgia, novelty, and escape. from the results of the literature review, it became clear that these findings support the notion that different attractions and destinations feed different motives to travel. therefore, it was expected that motives for visiting the kruger national park would also be different or at least similar to some of the studies conducted at other national parks. research by uysal et al. (1994), kim et al. (2006), swanson and horridge (2006) and oh et al. (1995) confirmed some of the motives in their research, although the similarities are not noticeably significant. compared to research by uysal et al. (1994) where the latter had five motives, only two, namely novelty and escape, were similar. none of the motives from the research by tao et al. (2004) or awaritefe (2004) were similar, although those particular studies were also conducted in national parks. the motives that were comparable with other research in other fields of tourism remain novelty and escape. it therefore seems that people travel to escape from their everyday routine as well as to experience something new. research in the kruger national park confirmed that the motives with the highest mean value were to escape, followed by nostalgia and nature. the latter is very important information for marketers of national parks, since the marketing campaign could be based on these findings. therefore, the concept of a “place to escape” can be used successfully, as this is a major motive. coupled to that is the aspect of nostalgia, and then the aspect of nature. in the past, marketers focused primarily on the wildlife that the kruger national park has to offer, especially the big 5. although this focus was successful, very little was done with regard to the escape motive. combining the nature and escape motives, for example “kruger national park, a place to escape while enjoying the big 5”, could add value to the marketing campaign. it is interesting to note that nostalgia as a motive has a higher rating than nature in the case of kruger national park. this is the first time that this motive was identified in research on travel motivations. a possible reason for this could be that most of the tourists visit this park regularly and could be regarded as brand loyal. they therefore become “attached” to the park. another possible reason could be that most of these brand-loyal tourists grew up with the park. component travel reasons nature activities attractions nostalgia novelty escape mean values 3.0062 1.5392 2.8072 3.4352 2.7724 4.2133 to see endangered species to see animals to see plants for educational reasons to take photos of animals to take photos of plants .854 .809 .741 .671 .560 .498 to attend conferences to attend events hiking .856 .780 .763 accommodation brand of the park climate of location grew up with the park .805 .637 .629 .482 park is visited since childhood to experience wildlife family time different species -.780 -.695 -.653 -.624 explore new destinations socialising with friends .794 .555 routine vacation relaxation .825 .787 table 4 pattern matrix component nature activities attractions nostalgia novelty escape nature 1.000 .206 .228 -.219 .115 .110 activities .206 1.000 .227 -.013 .072 -.096 attractions .228 .227 1.000 -.194 .048 .171 nostalgia -.219 -.013 -.194 1.000 -.141 -.094 novelty .115 .072 .048 -.141 1.000 -.030 escape .110 -.096 .171 -.094 -.030 1.000 table 5 component correlation matrix 158 travel motivations of tourists original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 154 159 jang, s. & wu, c.e. 2006. seniors’ travel motivations and the influential factors: an examination of taiwanese seniors. tourism management, 27: 306–316. johnson, r.a. & wichern, d.w. 2002. applied multivariate statistical analysis, 5th ed. new jersey: prentice hall. joubert, s.c.j. 2007. the history on the development of the sabie and shingwedzi reserves and the kruger national park, 1898 to 1946. available at: www.sanparks.org. [accessed 12 august 2007]. kim, h., borges, m.c. & chon, j. 2006. impacts of environmental values on tourism motivations: the case of fica, brazil. tourism management, 27: 957–967. lee, c.k., lee, y.k. & wicks, b.e. 2004. segmentation of festival motivation by nationality and satisfaction. tourism management, 25: 61–70. loker, l. & perdue, r. 1992. a benefit segmentation of a nonresident summer travel market. journal of travel research, 31(1): 31–35. mill, r.c. & morrison, a.m. 1985. the tourism system: an introductory text. englewood cliffs: prentice hall. oh, h.c., uysal, m., weaver, p.a. 1995. product bundles and market segmentation based on travel motivations: a canonical correlation approach. hospitality management, 14(2): 123–137. pienaar, u de v. 2007. neem uit die verlede. available at: www. sanparks.org. [accessed 12 august 2007]. saayman, m. 2006. marketing tourism, in products & destinations: getting back to basics. 2nd ed. potchefstroom: leisure consultants and publications, p. 234 saayman, m. & saayman, a. 2006. estimating the economic contribution of visitor spending in the kruger national park to the regional economy. journal of sustainable tourism, 14(1): 67–81. saayman, m. & fouché, m. 2007. executive summary of research profiles for selected national parks in south africa. potchefstroom: institute for tourism and leisure studies. schneider, i.e. & backman, s.j. 1996. cross-cultural equivalence of festival motivations: a study in jordan. festival management and event tourism, 4(3/4): 139–144. spss inc. 2006. spss® 15.0 for windows, release 15.0.0, copyright© by spss inc. chicago: illinois. swanson, k.k. & horridge, p.e. 2006. travel motivations as souvenir purchase indicators. tourism management, 27: 671– 683. toa , c.h., eagles, p.f.j. & smith, s.l.j. 2004. profiling taiwanese ecotourists using a self-definition approach. journal of sustainable tourism, 12(2): 149–168. uysal, m., mcdonald, c.d. & martin, b.s. 1994. australian visitors to us national parks and natural areas. international journal of contemporary hospitality management, 6(3): 18–24. weaver, d.b. & oppermann, m. 2000. tourism management. brisbane: wiley australia. yoon, y. & uysal, m. 2005. an examination of the effects of motivation and satisfaction on destination loyalty: a structural model. tourism management, 26: 45–56. the contribution of this research lies in the confirmation of “escape” as the most important motive for visiting the kruger national park, as well as the role that nostalgia plays. added to this, the study conducted by uysal et al. (1994) in national parks in the usa showed a slight similarity in terms of novelty and escape as travel motives. similar research in other national parks, however, showed no similarities. it would, therefore, be interesting to establish the motives for travelling to other national parks in south africa. added to this could be the desire to classify the distinguishing motives between national and international tourists visiting national parks, and particularly the kruger national park. however, it could be anticipated that nostalgia may not be such an important role motivation to the international tourist. acknowledgements the authors would like to thank sanparks for financial assistance (especially mr glenn phillips). we also would like to thank all the referees for their input in improving the document. references african safari travel. 2007. kruger national park map. available at: www.go2africa.com/south-africa/kruger-national-park/ map. [accessed 3 december 2007]. awaritefe, o.d. 2004. motivation and other considerations in tourist destination choice: a case study of nigeria. tourism geographies, 6(3): 303–330. bansal, h. & eiselt, h.a. 2004. exploratory research of tourists’ motivations and planning. tourism management, 25: 387– 396. backman, k.f., backman, s.j., uysal, m. & sunshine, k.m. 1995. event tourism: an examination of motivations and activities. festival management and event tourism, 3(1): 15–24. correia, a., oom do valle, p. & moço, c. 2007. why people travel to exotic places. international journal of culture, tourism and hospitality research, 1(1): 45–61. crompton, j.l. 1977. motives for pleasure vacation. annals of tourism research, 1(4): 408–424. eagles, p.f.j. & mccool, s.f. 2002. tourism in national parks and protected areas: planning and management. wallingford: cabi publishing. fodness, d. 1994. measuring tourist motivation. annals of tourism research, 21(3): 555–581. gcis (government communication and information system). 1998. south africa official yearbook, cape town: rustica. gcis (government communication and information system). 1999. south africa official yearbook, cape town: rustica. goeldner, c.r., ritchie, j.r. & mcintosh, r.w. 2000. tourism principles: practices and philosophies. 8th ed. new york: wiley. 159 article information authors: m. kyle s. smith1 nerina kruger1 taryn s. murray2,3 affiliations: 1rondevlei scientific services, south african national parks, south africa 2south african institute for aquatic biodiversity, grahamstown, south africa 3department of ichthyology and fisheries science, rhodes university, south africa correspondence to: kyle smith email: kyle.smith@sanparks.org postal address: po box 176, sedgefield 6573, south africa dates: received: 02 july 2014 accepted: 08 jan. 2015 published: 01 july 2015 how to cite this article: smith, m.k.s., kruger, n. & murray, t.s., 2015, ‘aerial surveys conducted along the garden route coastline, south africa, to determine patterns in shore fishing effort’, koedoe 57(1), art. #1236, 9 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1236 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. aerial surveys conducted along the garden route coastline, south africa, to determine patterns in shore fishing effort in this original research... open access • abstract • introduction • research method and design    • study area    • methods    • analyses       • aerial survey data maps       • effort • results    • spatial patterns in fishing effort    • temporal patterns in fishing effort    • total angling effort • discussion    • spatio-temporal patterns in fishing activity    • non-compliance with spatial closures       • limitations of the study • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ coastal environments provide a wide range of leisure opportunities, including recreational fishing. understanding spatial and temporal fishing patterns is important in ensuring wise management and sustainable use. to provide information on shore angler effort and distribution, randomised aerial surveys of the garden route coast between the eastern border of the tsitsikamma marine protected area and the kaaimans river mouth in the west were undertaken between december 2008 and november 2009. a total of 15 flights were conducted, with six flights taking place over weekends, two on public holidays and the balance on normal week days. angler effort was not uniformly distributed along the coastline, and spatial analysis highlighted coastal areas both inside and outside marine protected areas that had increased angler effort. in general, fishing effort was highest around more densely populated areas and concentrated in areas with easy access. although angler counts were highly variable, the seasonality of shore angling effort showed a slight increase during autumn and winter and angling effort was significantly higher on weekends. conservation implications: data obtained during these surveys can assist management with future conservation planning exercises, whilst also guiding daily law enforcement patrols to maximise angler encounters. introduction top ↑ coastal environments provide a wide range of recreational opportunities and are highly valued amongst various user groups (james 2000). understanding recreational use patterns is often necessary for adequate coastal management (smallwood et al. 2011). in particular, the spatial and temporal distribution of activities can be used in future coastal zoning, whilst also elucidating compliance with current zonation plans. similarly, a knowledge of resource use patterns can be used by management authorities in planning and optimising when and where law enforcement patrols should take place in order to maximise outputs and minimise costs. impacts arising from recreational activities depend on several factors, such as the type of activity, the number of participants, and the resilience of different species and habitats to the pressure (davenport & davenport 2006; meyer & holland 2008). recreational fishing is a popular activity and can arguably have a large impact through over-exploitation and harvesting of select species. although it is unlikely that a single angler's catch can have a measurable impact on fish populations, the cumulative impacts of (1) the widespread nature of recreational angling, (2) the total number of anglers involved and (3) the defined habitats in which fishing occurs are important factors to consider (cooke & cowx 2004; mcphee, leadbitter & skilleter 2002). within the south african shore-based linefishery, overfishing by recreational anglers is seen as a contributing factor to changes that have occurred in the composition of species catch (bennett 1991; brouwer & buxton 2002; brouwer et al. 1997; dunlop & mann 2012) and decreases in the abundance of target species (attwood & farquhar 1999; bennett 1991; cowley, brouwer & tilney 2002). this has resulted in increased emphasis on marine protected areas (mpas) as the basis for marine conservation, an essential component for fisheries management in south africa (attwood et al. 1997). the benefits of mpas typically include the restoration of marine ecosystems and natural ecosystem functioning, increased species diversity, and natural size and age structure of protected fish populations (halpern 2003). further fishery benefits include a spillover of subadult and adult fish, and the production of eggs and larvae that can be distributed over a large area (brouwer et al. 2003; kerwath et al. 2013; tilney et al. 1996). however, the effectiveness of mpas in meeting their conservation and fishery objectives is dependent on user compliance, with illegal fishing reducing their efficacy and negatively impacting fishery benefits (arias & sutton 2013). unfortunately, quantification of non-compliance by recreational anglers is rare, in particular with spatial zonation and over large areas. during this study, aerial surveys, as an alternative method to traditional on-the-ground or water-based surveys (brouwer 1997; mann et al. 2008; pollock et al. 1994), were used to (1) test the hypothesis that shore angling effort along the garden route coastline shows spatial and temporal variation, (2) assess compliance with spatial zoning and (3) provide recommendations to improve the efficiency of law enforcement activities within this area. research method and design top ↑ study area this study was conducted along the garden route coastline of south africa between the groot river in the east and the western border of the kaaimans river mouth (figure 1). the study area consisted of predominantly rocky coastline (78%) and sandy beaches (22%) and was 248 km long. the area falls under the jurisdiction of four local municipalities, with several towns and villages scattered throughout the adjacent interior. coastal access points are plentiful. the majority of people living in the area are situated in or near the towns of george, knysna and plettenberg bay (table 1). figure 1: a map of south africa showing the location of the study area along the south coast, and the location of marine protected areas, municipal wards and towns within the study area. table 1: population numbers per ward for each of the local municipalities within the study area. three mpas are situated within the study area: the tsitsikamma mpa consists of 68 km of closed (no resource use) coastline and forms part of the garden route national park, managed by south african national parks, whilst both the robberg and goukamma mpas, managed by capenature, are open to shore angling, but no boat angling is permitted. methods monthly aerial surveys covering the survey area were conducted between december 2008 and november 2009 in order to obtain ‘instantaneous’ counts of shore angling effort. survey days and flight times were randomly selected, but depended on the weather, and pilot and observer availability. all flights were conducted during daylight hours, with the earliest flight taking place at 09:10 and the latest at 15:55. all counts were done in an east–west direction, as the observers were seated on the starboard side of the plane. the first four surveys were undertaken in a four-seater robinson helicopter, after which a four-seater cessna light aircraft was used. flight parameters, including air speed and altitude, were standardised, but depended on weather conditions and the pilot's discretion. flight times ranged between 70 min and 80 min, with an average air speed of 189 km.hr−1 for both aircraft. altitude ranged between 30 m a.s.l. and 80 m a.s.l. for the robinson and between 150 m a.s.l. and 250 m a.s.l. for the cessna. on each flight, two observers counted shore anglers and one scribe entered the data into a global positioning system-enabled pocket pc. each spotter was equipped with a pair of binoculars (10 x 42 magnification) and continuously scanned the coastline, verifying the other spotter's counts. when large groups of anglers were encountered, a second flyby was made and anglers were re-counted by both spotters. information recorded during the aerial surveys included the date, time, weather, sea conditions, number and location of anglers or boats and habitat where shore anglers were fishing (rock or sand). data were also obtained from three aerial surveys conducted in december 2008 as part of a study researching the ecology, value and management of the garden route (chalmers et al. 2009). methods were similar, with the same cessna plane and pilot being used, two observers and similar standardised flight speed and altitude. this information was used in temporal, but not spatial (coastal effort maps), analyses. analyses aerial survey data maps the southern cape coastline that falls within the study area was buffered by 1 km to create a polygon representing the area of interest. spatial mapping of angler effort was plotted using three different coastal breaks. the first method (equal breaks) split the coastline into 10 km sections. the second method compared the status (mpa or non-mpa) using the layer of the mpas of the national spatial biodiversity assessment (lombard et al. 2004). the final method used the coastal breaks described in the coastal sensitivity atlas of southern africa (jackson & lipschitz 1984). for this analysis, exposed rocky headlands and wave-cut rocky platforms were combined into one category (rocky). aerial survey point data containing angler counts were overlaid onto each coastal break layer and the density of anglers per coastal section for each of the different breaks was calculated as anglers.km−1. angler effort was depicted in graduated colours for seven natural break classes. kruskal–wallis one-way analyses of variance (anovas) were performed to determine if there was a difference in angling effort spatially for each of the three different coastal breaks, and temporally between week and weekend days and seasons. effort the unit of fishing effort chosen was angler outings, with data pooled seasonally due to unequal sampling effort. total seasonal fishing effort (eseasonal outings) was calculated using the following formulae: where ew1 and ew2 are the weekday and weekend estimates respectively, given by: where j is week days or weekends, ei is the number of anglers on the ith day, d is the number of days sampled and p is the potential number of sampling days. total annual fishing effort (etoutings) was calculated as the sum of the estimated seasonal effort (eseasonal outings) multiplied by a correction factor (2.48) to account for angler turnover. total annual fishing effort (etoutings) was then multiplied by the average fishing trip duration of anglers interviewed during two ground-based roving creel studies (smith 2012; van zyl 2011). a kruskal–wallis one-way anova was performed to determine if there was a difference in angling effort between seasons, and a mann–whitney u test was used to establish if there was a difference in angling effort between week and weekend days. results top ↑ a total of 15 aerial surveys were conducted between december 2008 and november 2009 (table 2), with 6 flights occurring during summer months, 4 in autumn, 3 during winter and 2 in spring. most flights were conducted in the morning, with 3 weekend flights having being completed during the afternoon. table 2: summary of aerial surveys conducted each month, grouped seasonally and stratified according to week or weekend day. spatial patterns in fishing effort angling effort showed a heterogeneous distribution throughout the study area and although variability in count data was high, a significant difference in angling effort occurred between mpas and non-mpas (kruskal–wallis one-way anova, df = 6, p-value < 0.05) (figure 2) and equal area breaks (kruskal–wallis one-way anova, df = 24, p-value < 0.05) (figure 3). figure 2: the south african garden route coastal section broken into equal areas (a) and the spatial distribution of total angling effort (anglers.km−1) between kaaimans river in the west and the eastern border of the tsitsikamma marine protected area at the groot river (b). up–down bars indicate the 95% confidence interval. figure 3: the south african garden route coastal section divided into marine protected areas and open areas (a) and the spatial distribution of total angling effort (anglers.km−1) within marine protected areas and open areas between the kaaimans river in the west and the eastern border of the tsitsikamma marine protected area at the groot river (b). up–down bars indicate the 95% confidence interval. the majority of anglers (72.50%) fished in open access areas outside formal conservation borders (including no-take and controlled zones) and a bonferroni post-hoc test showed that the tsitsikamma mpa had significantly less fishing effort than all other sections (p-value < 0.05) with the exception of zone 9, in which no anglers were encountered. although the tsitsikamma mpa is zoned for no-take (i.e. no consumptive use), a total of 32 illegal fishermen were counted during all the surveys, with an overall angler density of 0.03 anglers.km−1 (figure 3). outside of formal protected areas, the highest density of anglers occurred in section one (wilderness) with 1.35 anglers.km−1, followed by section 15 (plettenberg bay) and 17 (nature's valley) with overall angler densities of 0.78 anglers.km−1 and 0.58 anglers.km−1 respectively (figure 2). these three sections contributed to the overall angler densities being highest (0.73 anglers.km−1) along the larger section of coastline between the kaaimans river mouth and the western border of the goukamma mpa. this was followed by the plettenberg bay coastline (robberg to de vasselot) with 0.65 anglers.km−1 and the de vasselot section of the garden route national park with the third highest density of 0.51 anglers.km−1 (figure 3). the goukamma and robberg mpas had overall angler densities of 0.41 anglers.km−1 and 0.14 anglers.km−1 respectively (figure 3), whilst the coastal section between the goukamma and robberg mpas, consisting of large stretches of exposed rocky coastline with few access points, had a relatively low overall angler density of 0.16 anglers.km−1. angling effort occurred on both rocky and sandy substrates throughout the study area with both high densities (2.23 anglers.km−1 – 3.94 anglers.km−1) occurring over rocky (figure 4a) and sandy substrates (figure 4b). although total angler density on rocky substrates was lower (0.31 anglers.km−1) than on sandy substrates (0.47 anglers.km−1), fishing effort between substrates was not significantly different (mann–whitney u test, p-value 0.221). figure 4: the distribution of total angling effort (anglers.km−1) between the kaaimans river in the west and the western border of the no-take tsitsikamma marine protected area between rocky areas (a), sandy areas (b) and both rocky and sandy areas combined (c). temporal patterns in fishing effort although more anglers were encountered during the autumn (0.37 anglers.km−1) and winter (0.38 anglers.km−1) months (figure 5a), with fewer anglers fishing in spring (0.20 anglers.km−1) and summer (0.21 anglers.km−1), no significant difference was observed (kruskal–wallis one-way anova, df = 3, p-value 0.266). this was due to the high variability within the data, exacerbated by the low sample size. fishing pressure was significantly higher during weekends and public holidays (0.38 anglers.km−1) than on week days (0.18 anglers.km−1) (mann–whitney u test, df = 1, p-value 0.028) (figure 5b). figure 5: mean shore angling effort (anglers.km−1) (a) seasonally and (b) during week and weekend days along the south african garden route coast between december 2008 and november 2009 (public holiday counts were included as weekend days). total angling effort total annual shore angling effort was estimated at 49 812 angler outings. recent angler interviews conducted along portions of this coastline indicate that anglers spend on average between 5 hours (smith 2012) and 6 hours (van zyl 2011) fishing per outing. using these figures, a total annual fishing effort estimate was estimated to be between 226 645 angler hours and 298 872 angler hours. discussion top ↑ assessing spatio-temporal resource use patterns is important when implementing fishery and conservation management practices. this information can help with future conservation and spatial planning exercises by highlighting areas from which anglers may be displaced or, conversely, areas where no fishing activity occurs and which may act as natural spatial harvest refugia for target species (smallwood & beckley 2012). furthermore, identifying popular fishing locations can allow management authorities to maximise resource allocation and streamline daily law enforcement operations (smallwood & beckley 2012). spatio-temporal patterns in fishing activity angler distribution is unlikely to be uniformly distributed along any coastline and may be impacted by access points and infrastructure that have a clustering effect (smallwood et al. 2012). similarly, habitat preferences of target fish species can influence the spatial distribution of anglers attracted to particular sites and different species (smallwood et al. 2013). recreational fishing along the garden route coastline showed a heterogeneous spatial distribution, with some localised areas having high angler densities. these areas were generally associated with ease of access and proximity to the more heavily populated urban areas of george and plettenberg bay. a similar pattern was shown along the kwazulu-natal coast of south africa (mann et al. 2008), where angler effort was largely concentrated around access points, with increased effort along the more developed and highly populated sections of the coastline. the concentration of anglers around easy access points along south africa's coastline has more than likely been heightened by the 2002 ban (schedule 44 of the national environmental management act [act no. 107 of 1998]) on beach driving, which limits anglers’ access to the more remote sections of coastline (mann & tyldesley 2012; mann et al. 2008). in fisheries where angling effort is widely dispersed (e.g. coastlines), instantaneous estimates of total fishing effort obtained from aerial surveys are more accurate and preferable to land-based roving creel surveys (pollock et al. 1994). although no previous aerial surveys have been undertaken along this section of coastline for comparative purposes, a number of land-based roving creel surveys covering sections of the coastline have been completed, including plettenberg bay (king 2006), the goukamma mpa (van zyl 2011) and the wilderness to sedgefield coastline (smith 2012). these projects, conducted at finer spatial scales, show similar patterns and general trends to those recorded during the aerial surveys conducted in this study. greater angling effort (1.03 anglers.km−1) was recorded along the wilderness to sedgefield (smith 2012) and plettenberg bay coastlines (0.71 anglers.km−1 [king 2006]) during land-based surveys, in comparison to 0.84 anglers.km−1 and 0.69 anglers.km−1 recorded during the aerial surveys of this study. due to the generally shorter distance and slower speed of land-based creel surveys, a higher estimation of effort in comparison to aerial surveys would be expected (pollock et al. 1994). the goukamma study did not report on angler densities in terms of anglers.km−1 of coastline, but estimated total effort relating to angler hours (21 428 [van zyl 2011]), approximately half the 48 167 angler hours estimated in the wilderness study (smith 2012). this is similar to the trend seen in the effort estimations of the aerial surveys, with an overall angler density of 0.46 anglers.km−1 recorded for the goukamma mpa. the distribution and density of recreational users in general is known to be impacted by large temporal factors such as seasons and holiday periods (hingham & hinch 2002). large-scale patterns in recreational shore fishing along the south african east coast have also been shown to fluctuate seasonally, with an increase in summer and a decrease in winter (brouwer et al. 1997). both king (2006) and smith (2012) have shown that recreational shore fishing within plettenberg bay and the wilderness coastline followed this trend, attributing the increase in effort to an influx of visitors during school holiday periods. however, results from the aerial surveys are more consistent with those from goukamma (van zyl 2011) and the kwazulu-natal shore fishery (dunlop & mann 2012; mann et al. 2008), which both showed an increase in shore angling density over the winter months. two popular targeted shore angling species, galjoen (dichistius capensis) and elf (pomatomus saltatrix), show strong seasonal abundance and have closed seasons between 15 october and the last day in february, and 01 october and 30 november respectively. it is thus likely that those anglers who specifically target these species are temporarily absent from the fishery during these closed periods (mann et al. 2008; van zyl 2011). given that no previous aerial surveys have been conducted along the garden route coastline and shore-based surveys have only covered portions of the coastline, it is impossible to say whether total fishing effort is increasing or not. a decrease in shore-based angling effort along the kwazulu-natal coastline (dunlop & mann 2012) has been attributed to security concerns, declining catches, an increase in the cost of fishing and stricter linefish regulations. as catch rates within the shore fishery have decreased (brouwer & buxton 2002), the duration and frequency of fishing may have altered, which in turn would affect the turnover rate used to calculate the total estimated effort. however, the current average fishing duration of between 5 hours (smith 2012) and 6 hours (van zyl 2011) is very similar to the 5 hours calculated 15 years earlier (brouwer 1997). independent effort calculations are seen as a crucial part of the management and monitoring of the south african shore fishery (dunlop & mann 2012) and the total angling effort estimated from this study provides a baseline for future work. non-compliance with spatial closures with greater biodiversity and abundance of fish, mpas where fishing is allowed are attractive locations for recreational anglers (cooke et al. 2006; götz, cowley & winker 2008; parker, booth & mann 2013). however, non-compliance by anglers fishing in closed, no-take areas can undermine conservation and fishery management objectives. along the garden route, shore fishing is allowed in all mpas except tsitsikamma. however, illegal shore fishing (poaching) was observed on several occasions during this study within the no-take tsitsikamma mpa. although fishing levels inside the park were low compared to outside adjacent areas, illegal fishing within this important mpa is cause for concern. within the ningaloo marine park in north west australia, anglers were found to be concentrated in localised areas with easy access, and limited non-compliance was observed within no-take zones (smallwood et al. 2011). the low levels of non-compliance were attributed to a number of factors, including (1) multiple educational tools (signboards and brochures), (2) visual zonation reference points, (3) various enforcement patrols (boatand land-based) and (4) a willingness by anglers to fish at alternative sites (smallwood & beckley 2012). more research is required regarding the illegal fishing taking place within the tsitsikamma mpa to gain a greater understanding of the extent of this activity and the underlying drivers, which could then be used by management to address and resolve this issue. although the goukamma and robberg mpas are open to shore-based fishing, these areas had comparatively lower angler densities than other sections of coastline outside the formal conservation areas. fishing locations within robberg are limited due to a large seal colony on the eastern side and the exposed rocky coastline on the west, restricting fishing to a few well-known locations (king 2005). the goukamma mpa has a combination of habitat types and land-based roving creel surveys conducted within this reserve (van zyl 2011) showed that anglers were clustered around easily accessible areas, favouring rocky and mixed shores over sandy beaches. similar to the aerial survey data presented here, angler densities were greater around the buffalo bay section of the reserve, which may be due to ease of access, with anglers in the goukamma section having to walk up to 6 km to reach some fishing locations. limitations of the study surveys to provide information on spatial and temporal patterns of resource use need to be designed and implemented at appropriate scales. numerous access points, the longitudinal and rugged nature of coastlines and the size of management areas complicate survey design. aerial surveys can be an effective technique for obtaining spatial and temporal data on recreational activities, including fishing, over a broad landscape (pollock et al. 1994). such surveys are cost-effective and efficient, with minimal personnel involved and can cover large geographic areas in a relatively short time (malvestuto 1983). to avoid sampling biases, flights must be random in terms of direction, time of day and type of day (week or weekend) (pollock et al. 1994). in practice this becomes difficult, with weather conditions in particular playing an important role in determining flight days. the greatest limitation of this study was the inability to keep sampling effort equal between months, with flights conducted on both week and weekend days. subsequently, monthly estimates of angler effort and comparisons were not possible. analysing the data by season to estimate total angler effort attempts to alleviate this issue to some extent, but an unknown degree of error is being introduced and needs to be acknowledged. spatial accuracy of observed data points was improved through the use of data loggers that automatically recorded time and positional information, enabling the observers to simply record numbers and substrate. prominent landmarks were also geo-referenced prior to the aerial flights, providing a known position that could be recorded, eliminating sampling errors associated with fishing at these points. however, visibility bias and in particular difficulty in seeing anglers on rocky sections of coastline may have led to an underestimation of total angling effort. ground truthing of counts would be important for future surveys. conclusion top ↑ aerial surveys are a useful and well-established method of collecting data across large spatial scales. these data can be used in both future planning exercises and also in determining the success of current management plans and policies (e.g. non-compliance with spatial zoning). although our ability to estimate total angler effort from the current data set is limited due to unequal monthly sampling effort, results from this study showed temporal trends with seasonal fluctuations for recreational fisheries. an increase in effort over weekends and public holidays was evident, whilst broad-scale spatial patterns were similar to results from more in-depth, land-based projects. in general, fishing effort was highest around the more heavily populated areas of george and plettenberg bay and concentrated in areas with easy access. should aerial surveys be implemented again, care should be taken to ensure equal monthly effort (i.e. flights should be conducted at least bi-monthly – on one weekday and one weekend day or public holiday), with flight scheduling incorporating back-up days for bad weather and mechanical problems. the value of surveys could be further increased by collecting data on other types of coastal resource use, such as bait harvesting. the findings of this study have relevance to the management of the fishery along the garden route coastline in guiding the deployment of fisheries management field personnel, both spatially and temporally, to maximise contact with fishers and therefore improve levels of compliance. acknowledgements top ↑ funding for the flights was made available through the national marine protected area budget, allocated to the south african national parks by the department of environmental affairs. we thank the volunteers and pilots who assisted during the aerial surveys and two anonymous reviewers for their positive contributions towards this manuscript. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions m.k.s.s. (south african national parks) was responsible for the study conceptualisation, data collection, analyses, report writing and editing. n.k. (south african national parks) contributed to the study design, data collection, gis analyses, report writing and editing, and t.s.m. (south african institute for aquatic biodiversity) contributed to the study design, data collection, report writing and editing. references top ↑ arias, a. & sutton, s.g., 2013, ‘understanding recreational fishers’ compliance with no-take zones in the great barrier reef marine park’, ecology and society 18(4), 18. http://dx.doi.org/10.5751/es-05872-180418 attwood, c.g. & farquhar, m., 1999, ‘collapse of linefish stocks between cape hangklip and walker bay, south africa’, south african journal of marine science 24, 151–160. http://dx.doi.org/10.2989/025776102784528448 attwood, c.g., mann, b.q., beaumont, j. & harris, j.m., 1997, ‘review of the state of marine protected areas in south africa’, south african journal of marine science 18, 341–367. http://dx.doi.org/10.2989/025776197784160910 bennett, b.a., 1991, ‘conservation in the marine environment: some problems with management of shore-angling in the southwestern cape’, south african journal of aquatic science 17, 12–18. http://dx.doi.org/10.1080/10183469.1991.9631309 brouwer, s.l. & buxton, c.d., 2002, ‘catch and effort of the shore and skiboat linefisheries along the south african eastern cape coast’, south african journal of marine science 24, 341–354. http://dx.doi.org/10.2989/025776102784528286 brouwer, s.l., 1997, ‘an assessment of the south african east coast linefishery from kei mouth to stil [sic] bay’, msc thesis, department of ichthyology and fisheries science, rhodes university, grahamstown. brouwer, s.l., griffiths, m.h. & roberts, m.j., 2003, ‘adult movement and larval dispersal of argyrozona argyrozona (pisces: sparidae) from a temperate marine protected area’, african journal of marine science 25, 395–402. http://dx.doi.org/10.2989/18142320309504028 brouwer, s.l., mann, b.q., lamberth, s.j., sauer, w.h.h. & erasmus, c., 1997, ‘a survey of the south african shore-angling fishery’, south african journal of marine science 18, 165–177. http://dx.doi.org/10.2989/025776197784161126 chalmers, r., bennett, r.h., turpie, j.k., andrew, m., andrew, t., clark, b.m. et al., 2009, ‘ecology, value and management of the garden route coast’, report prepared for wwf-cape marine programme, south africa. cooke, s.j. & cowx, i.g., 2004, ‘the role of recreational fishing in global fish crises’, bioscience 54, 857–859. http://dx.doi.org/10.1641/0006-3568(2004)054[0857:trorfi]2.0.co;2 cooke, s.j., danylchuk, a.j., danylchuk, s.e., suski, c.d. & goldberg, t.l., 2006, ‘is catch-and-release recreational angling compatible with no-take marine protected areas?’, ocean and coastal management 49, 342–354. http://dx.doi.org/10.1016/j.ocecoaman.2006.03.003 cowley, p.d., brouwer, s.l. & tilney, r.l., 2002, ‘the role of the tsitsikamma national park in the management of four shore-angling fish along the south-eastern cape coast of south africa’, south african journal of marine science 24, 27–35. http://dx.doi.org/10.2989/025776102784528664 davenport, j. & davenport, j.l., 2006, ‘the impact of tourism and personal leisure transport on coastal environments: a review’, estuarine, coastal and shelf science 67, 280–292. http://dx.doi.org/10.1016/j.ecss.2005.11.026 dunlop, s.w. & mann, b.q., 2012, ‘an assessment of participation, catch and effort in the kwazulu-natal shore-based marine linefishery, with comments on management effectiveness’, african journal of marine science 34, 479–496. http://dx.doi.org/10.2989/1814232x.2012.725526 götz, a., cowley, p.d. & winker, h., 2008, ‘selected fishery and population parameters of eight shore-angling species in the tsitsikamma national park no-take marine reserve’, african journal of marine science 30, 519–532. http://dx.doi.org/10.2989/ajms.2008.30.3.7.641 halpern, b.s., 2003, ‘the impact of marine reserves: do reserves work and does reserve size matter?’, ecological applications 13(1), 117–137. http://dx.doi.org/10.1890/1051-0761(2003)013[0117:tiomrd]2.0.co;2 hingham, j. & hinch, t., 2002, ‘tourism, sport and seasons: the challenges and potential of overcoming seasonality in the sport and tourism sectors’, tourism management 23(2), 175–185. http://dx.doi.org/10.1016/s0261-5177(01)00046-2 jackson, l.f. & lipschitz, s., 1984, ‘coastal sensitivity atlas of southern africa’, compiled for the department of transport, charts 16–18, national book printers, goodwood, cape town. james, r.j., 2000, ‘from beaches to beach environments: linking the ecology, human-use and management of beaches in australia’, ocean & coastal management 43(6), 495–514. http://dx.doi.org/10.1016/s0964-5691(00)00040-5 kerwath, s.e., winker, h., götz, a. & attwood, c.g., 2013, ‘marine protected area improves yield without disadvantaging fishers’, nature communications 4, 2347. http://dx.doi.org/10.1038/ncomms3347 king, c.m., 2005, ‘towards a new approach for coastal governance with an assessment of the plettenberg bay shore-based linefishery’, msc thesis, department of ichthyology and fisheries science, rhodes university, grahamstown. lombard, a.t., strauss, t., harris, j., sink, k., attwood, c. & hutchings, l., 2004, south african national spatial biodiversity assessment 2004: technical report, volume 4: marine component, south african national biodiversity institute, pretoria. malvestuto, s.p., 1983, ‘sampling the recreational fishery’, in l.a. nielsen & d.l. johnson (eds.), fisheries techniques, pp. 397–419, american fisheries society, bethesda, maryland. mann, b.q. & tyldesley, m., 2012, ‘monitoring the recovery of a previously exploited surf-zone habitat in the st lucia marine reserve using a no-take sanctuary area as a benchmark’, in c. attwood, t. booth, s. kerwath, b. mann, s. marr, j. duncan et al. (eds.), ‘a decade after the emergency: the proceedings of the 4th linefish symposium’, pp. 115–127, wwf south africa report series – 2013/marine/001. mann, b.q., nanni, g. & pradervand, p., 2008, ‘a monthly aerial survey of the kwazulu-natal marine shore fishery’, unpublished report, oceanographic research institute, durban. mcphee, d.p., leadbitter, d. & skilleter, g.a., 2002, ‘swallowing the bait: is recreational fishing in australia ecologically sustainable?’, pacific conservation biology 8(1), 40–51. meyer, c.g. & holland, k.n., 2008, ‘spatial dynamics and substrate impacts of recreational snorkelers and scuba divers in hawaiian marine protected areas’, journal of coastal conservation 12, 209–216. http://dx.doi.org/10.1007/s11852-009-0043-1 parker, d., booth, a.j. & mann, b.q., 2013, ‘a spatio-temporal assessment of the trachinotus botla shore-fishery in kwazulu-natal, south africa’, african journal of marine science 35, 35–46. http://dx.doi.org/10.2989/1814232x.2013.769908 pollock, k.h., jones, c.m. & brown, t.l., 1994, ‘angler survey methods and their applications in fisheries management’, american fisheries society special publication 25, 135–164. smallwood, c.b. & beckley, l.e., 2012, ‘spatial distribution and zoning compliance of recreational fishing in ningaloo marine park, north-western australia’, fisheries research 125–126, 40–50. http://dx.doi.org/10.1016/j.fishres.2012.01.019 smallwood, c.b., beckley, l.e. & moore, s.a., 2013, ‘influence of zoning and habitats on the spatial distribution of recreational activities in a multiple-use marine park’, coastal management 40(4), 381–400. http://dx.doi.org/10.1080/08920753.2012.692312 smallwood, c.b., beckley, l.e., moore, s.a. & kobryn, h.t., 2011, ‘assessing patterns of recreational use in large marine parks: a case study from ningaloo marine park, australia’, ocean & coastal management 54, 330–340. http://dx.doi.org/10.1016/j.ocecoaman.2010.11.007 smith, m.k.s., 2012, ‘garden route national park – wilderness section coastal marine resource use monitoring programme’, scientific report number 05/2012, south african national parks, skukuza. superweb by space time research, n.d., viewed april 2014, from http://interactive.statssa.gov.za/superweb/login.do tilney, r.l., nelson, g., radloff, s.e. & buxton, c.d., 1996, ‘ichthyoplankton distribution and dispersal in the tsitsikamma national park marine reserve, south africa’, south african journal of marine science 17, 1–14. http://dx.doi.org/10.2989/025776196784158482 van zyl, c., 2011, ‘the use of a roving creel survey to monitor exploited coastal fish species in the goukamma marine protected area, south africa’, mtech thesis, nelson mandela metropolitan university, port elizabeth. article information author: robert e. manning1 affiliation: 1park studies laboratory, university of vermont, united states of america correspondence to: robert manning postal address: 313a aiken center, 81 carrigan drive, university of vermont, burlington, vt 05405, united states of america dates: received: 27 mar. 2013 accepted: 13 nov. 2013 published: 24 june 2014 how to cite this article: manning, r.e., 2014, ‘research to guide management of outdoor recreation and tourism in parks and protected areas’, koedoe 56(2), art. #1159, 7 pages. http://dx.doi.org/10.4102/ koedoe.v56i2.1159 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. research to guide management of outdoor recreation and tourism in parks and protected areas in this original research... open access • abstract • introduction • a management framework • research to support management    • formulating indicators and standards of quality    • monitoring indicators of quality    • implementing management actions to ensure standards of quality are maintained • a growing body of research and associated literature • ethical considerations • trustworthiness • conclusion • acknowledgements    • competing interests • references abstract top ↑ a framework for managing outdoor recreation and tourism in parks and protected areas was presented in this article. this management-by-objectives framework includes, (1) formulating indicators and standards of quality, (2) monitoring indicators of quality and (3) implementing management actions designed to maintain standards of quality. this management framework can be used to help balance the demand for outdoor recreation and tourism and the need to protect park resources and the quality of the visitor experience. a programme of research to help guide application of this management framework was described and illustrated. this research is part of a growing body of scientific and professional literature on outdoor recreation and tourism that can be used to build the capacity of park and protected area management agencies. conservation implications: the management framework described in this article, and the associated programme of research, can be used by conservation practitioners to balance use and protection of national parks and protected areas. introduction top ↑ parks and protected areas are becoming increasingly important in contemporary society. parks are vital to people in many ways: they offer open, green spaces in our ever-developing world, they are retreats from the hectic lives that many of us lead and they protect wildlife and other elements of biodiversity, as well as historical and cultural resources that are important markers of society. and, of course, tourism and outdoor recreation are also important, offering healthy and satisfying leisure activity, intimate contact with the out-of-doors, opportunities to build family solidarity, enjoyment and appreciation of our natural environment and cultural heritage, employment and economic development to local people and communities and a myriad of other benefits. because of their importance, parks and protected areas have been established across the globe. formally designated national and international protected areas now number over 150 000, include more than 24 million km2of land and water and cover more than 12% of the surface of the earth (international union for conservation of nature & united nations environment programme world conservation monitoring centre 2011, 2012). tourism and outdoor recreation have also grown exponentially (balmford et al. 2009). worldwide tourism has been estimated to account for 10% of gross domestic product and outdoor recreation and wildlife viewing make up one of tourism’s fastest growing sectors (davenport et al. 2002; goodwin 1996; mastny 2001; world travel and tourism council 2007). but how can parks and protected areas be managed to maximise their effectiveness? in particular, parks and protected areas are generally established to meet two objectives, (1) protect park resources and the quality of the visitor experience and (2) accommodate visitor use and enjoyment. these objectives can be complementary when visitor use leads to widespread support for parks and protected areas, including economic development opportunities for local communities and needed revenues to manage parks. however, these objectives can often come into conflict when the amount and type of visitor use leads to unacceptable impacts on park resources and the quality of the visitor experience. this paper addresses a programme of research and management designed to help maximise attainment of management objectives and minimise potential conflict between these objectives. a management framework top ↑ a number of frameworks have been developed to help guide management of outdoor recreation and tourism in parks and protected areas. prominent examples include limits of acceptable change (lac) developed by the us forest service (mccool 1994; mccool & cole 1997; stankey et al. 1985) and visitor experience and resource protection (verp) developed by the us national park service (manning 2001, 2007, 2009; national park service 1997). whilst there are some differences in terminology and sequencing of steps, these and related management-by-objectives frameworks rely on the three basic steps as shown in figure 1 and described below (manning 2004).firstly, management objectives and associated indicators and standards of quality are formulated for a park or site within a park. management objectives describe desired conditions – the level of resource protection and the type and quality of recreation experiences – to be maintained. associated indicators and standards of quality define these objectives in quantitative, measurable form. secondly, indicators of quality are monitored to determine if standards of quality are being maintained. thirdly, if standards of quality are violated, or are in danger of being violated, then management action is required. this management framework represents a long-term commitment to management that requires maintaining standards of quality, periodic monitoring of indicators of quality and reconsideration of management practices based on monitoring data. when circumstances warrant, for example when a management plan has reached the end of its useful life and needs to be revised, management objectives and associated indicators and standards of quality can be reconsidered. the management framework outlined above is built on foundational concepts that have emerged in the environmental literature. for example, it represents a form of ‘adaptive management’; that is, it is implemented based on the best information available, but new information is gathered in an ongoing programme of monitoring and research and management practices are revised in keeping with this new information (christensen et al. 1996; holling 1978; lee 1993; stankey, clark & bormann 2005; walters 1986). it uses management objectives and associated indicators and standards of quality as quantitative expressions of the ‘carrying capacity’ of parks and protected areas: the type and amount of use that can be accommodated without unacceptable impacts to park resources and the quality of the visitor experience (graefe, vaske & kuss 1984; manning 2007, 2011a; shelby & heberlein 1986; stankey & manning 1986; wagar 1964; whittaker et al. 2011). the framework requires that management actions be implemented when monitoring shows that standards of quality are in danger of being violated and this is keeping with the call for deliberative management as demanded in the context of common property resources (of which parks and protected areas are classic examples) (hardin 1968). management objectives and associated indicators and standards of quality can and should be derived from ecological research and knowledge, along with public participation and engagement as suggested by the concept of ‘ecosystem management’ (agee & johnson 1987; grumbine 1994; society of american foresters 1993). figure 1: a framework for managing outdoor recreation and tourism in parks and protected areas. research to support management top ↑ application of the management framework described above can and should be supported by a programme of research. this section of the article outlines examples of a series of studies conducted in national parks to help inform each of three basic steps of the management framework described in the previous section. this programme of research, (1) uses surveys of park visitors and other stakeholders and visual simulations of a range of park and outdoor recreation conditions to help guide formulation of indicators and standards of quality, (2) develops a computer simulation model of recreation use to monitor crowding-related standards of quality and (3) tests the efficacy of management practices designed to maintain standards of quality. formulating indicators and standards of quality a programme of research was designed to help support formulation of indicators and standards of quality at arches national park (manning & freimund 2004; manning et al. 1996). arches covers 29 500 ha of high-elevation desert in south-east utah and is named after the nearly 2000 distinctive natural sandstone arches in this area. a road connects many of the most iconic features of the park and this contributes to rapidly growing visitation that now numbers more than a million visits per year. the programme of research was conducted in two phases.phase i explored potential indicators of quality for the visitor experience. interviews were conducted with park visitors and 10 focus group sessions were held with park staff, local community residents and representatives of key stakeholder groups. interviews and focus groups asked participants what they considered to be the most important factors that defined the quality of the visitor experience at arches. study findings suggested that the most important indicators were, (1) number of visitors seen at attraction sites and along trails, (2) the presence of social trails (visitor-made informal trails) and associated soil and vegetation impacts and (3) visitor knowledge of park regulations about hiking off official, maintained trails. phase ii research was designed to help formulate standards of quality for the indicator variables noted above. a survey of park visitors was conducted using both personal interviews and mail-back questionnaires. visual research methods were used to illustrate a range of conditions for indicator variables. for example, a series of 16 computer-generated images was created to represent a range of visitor use levels at delicate arch, a principal visitor attraction. these images were created using photo-editing software and examples of these images are shown in figure 2. these images were presented to a representative sample of visitors who had just completed a hike to delicate arch. respondents were asked to judge the acceptability of each image on a scale of -4 (‘very unacceptable’) to +4 (‘very acceptable’). analogous sets of images were created for the number of hikers along developed trails and environmental impacts caused by off-trail hiking. study findings for the acceptable number of people at delicate arch are shown in figure 3. this figure represents the average (mean) acceptability ratings for each of the 16 study images. it is clear from the graph that acceptability declines with increasing use. average acceptability ratings fall out of the acceptable range and into the unacceptable range at about 30 people at one time (paot) at delicate arch and park staff selected this number as the minimum acceptable standard of quality. in a similar manner, standards of quality were formulated for other indicator variables and other features and areas in the park (national park service 1995). the park is now monitoring indicator variables to ensure that standards of quality are being maintained. figure 2: sample computer-generated photographs illustrating a range of use levels in terms of people at one time at delicate arch in arches national park, utah, with (a) 0 people, (b) 12 people, (c) 52 people and (d) 108 people. figure 3: average acceptability ratings for the 16 photographs illustrating a range of people at one time at delicate arch in arches national park, utah. monitoring indicators of quality the management framework described above includes monitoring indicators of quality. however, monitoring some indicators, such as the number of groups encountered along a trail network, can be difficult to directly observe. moreover, it would be useful to be able to estimate the maximum amount of visitor use that can be accommodated in a park without violating the standards of quality that have been set (a form of ‘proactive monitoring’) (lawson et al. 2003). computer-based simulation modelling can be used to address both of these issues. simulation modelling has become an important tool in many fields of study and practice and is increasingly being applied in management of parks and outdoor recreation (cole 2005). an example of its usefulness as a monitoring tool is manifested in a study at yosemite national park, california (manning et al. 2002). much of the recreational use of the park is focused on yosemite valley and its iconic attractions, including sheer granite walls of up to 1500 m and several waterfalls that are amongst the highest in the world. the park accommodates over 4 million visits per year and its carrying capacity is a perennial and contentious issue. research at yosemite was conducted in two phases. the first phase used a series of visitor surveys to support formulation of indicators and standards of quality for the visitor experience (much like the study at arches described above). many visitors reported that crowding was an important indicator of quality at attraction sites (i.e. the number of paot at an attraction site) and along trails (i.e. the number of people per viewscape along trails or persons-per-viewscape [ppv]). visitors also reported standards of quality – maximum acceptable paot and ppv – for these indicator variables. the second phase of research focused on developing simulations of visitor use at important attraction sites and trails. these simulation models were designed to estimate the maximum number of visitors that could be accommodated at these sites and trails without violating the paot and ppv standards of quality found in the first phase of research. observations and counts of visitor use at several sites and trails were conducted, including number of visitors arriving per hour, visitor group size, length of time visitors stop at attraction sites, the speed at which visitors hike and the length of typical trail viewscapes (how far ahead hikers can typically see along trails). simulation models of visitor use were developed using these and other data. models were constructed using the commercially available, object-oriented simulation software, extend. the object orientation of the software does not require code-writing and enables programming through graphic display of objects and connections. the model was constructed to provide output in both graphic and numeric formats. figure 4, for example, provides a graphic display of minute-by-minute ppv levels along the trail to bridalveil fall over the duration of a day. this output was derived from a model run using an average summer day total use level of 1415 visitors (derived from the counts of visitor use). the primary use of the study models was to estimate the maximum total daily use that could be accommodated at each study site without violating crowding-related standards of quality. models were run multiple times to estimate these numbers. the simulation models for bridalveil fall suggest that a maximum of 3200 visitors a day can be accommodated on the trail to bridalveil fall without violating the ppv standard of quality, but that only 1700 visitors a day can be accommodated at the base of bridalveil fall without violating the paot standard of quality. figure 4: persons-per-viewscape along the trail to bridalveil fall in yosemite national park, california. implementing management actions to ensure standards of quality are maintained the management framework described above requires that management action be taken when monitoring demonstrates that standards of quality have been violated or are in danger of being violated. but how effective are alternative management actions? research can help evaluate the potential efficacy of selected management practices.a study of recreation at the summit of cadillac mountain in acadia national park, maine is an example of the way in which research can help guide management (park et al. 2008). acadia attracts nearly 2.5 million visits per year and the park is less than 20 000 ha; this makes acadia a very intensively used national park. cadillac mountain is an especially popular visitor attraction as its summit offer expansive views and is accessible by both road and trail. more than 5000 visitors per day may ascend the mountain on peak summer days (baldwin & lapage 2003; turner & lapage 2001). a short, paved loop trail is provided for visitor use. unfortunately, many visitors walk off the maintained trail and this has caused extensive damage to the summit’s fragile soils and vegetation (evan 2002; turner & lapage 2001). a research project was administered to test the efficacy of alternative management practices designed to encourage visitors to stay on the maintained trail. the study employed a series of experimental treatments – five management practices from ‘indirect’ (e.g. information and education) to ‘direct’ (e.g. a fence bordering the trail) and associated controls. during all treatment and control periods, researchers unobtrusively observed visitors and recorded the number that walked off the maintained trail. in addition, a survey of visitors was conducted to explore why the experimental management treatments worked or not. study findings suggest that all of the experimental management treatments reduced the percentage of visitors who walked off the maintained trail. more aggressive applications of indirect management practices were more effective than less aggressive applications and the direct practice of fencing was the most effective. however, none of the indirect management practices reduced off-trail walking enough to effectively limit damage to soils and vegetation. implications of the study are that a coordinated programme of indirect and direct management practices be implemented at cadillac mountain (and similar sites) that includes, (1) a rule requiring that visitors stay on maintained trails, (2) enforcement of this rule as necessary, (3) unobtrusive fencing along the margins of the trail, (4) redesigning of the maintained trail to extend it, widen it, and to provide spur trails to key photo points and (5) an aggressive education programme informing visitors of the regulation to stay on maintained trails and the reason for this regulation. a growing body of research and associated literature top ↑ the studies outlined above are a part of a growing body of research and associated scientific and professional literature that can help guide management of parks and protected areas for outdoor recreation and tourism. these advances can help build the capacity of park and protected area management agencies in several ways. firstly, research has supported development of a number of conceptual and management frameworks (manning 2011b; mccool, clark & stankey 2007). conceptual frameworks – such as carrying capacity (manning 2007; shelby & heberlein 1986; wagar 1964; whittaker et al. 2011), common property resources (hardin 1968), limits of acceptable change (frissell & stankey 1972), the recreation opportunity spectrum (brown, driver & mcconnell 1978; clark & stankey 1979; driver & brown 1978), sustainable tourism and ecotourism (buckley 2009; honey 2008), ecosystem management (agee & johnson 1987; grumbine 1994), adaptive management (holling 1978; lee 1993; stankey et al. 2005; walters 1986) and social and environmental justice (floyd 1999; solop, hagen & ostergren 2003; stanfield mccown et al. 2012) – identify and analyse issues vital to management of parks and protected areas and offer managers organisational structures to think through these important issues. moreover, these conceptual frameworks have formed the foundation for a number of management frameworks such as lac and verp, as discussed in this article. a number of related management frameworks have been developed and applied in parks and protected areas around the world, offering a choice of management frameworks from which to choose (belokurov et al. 2009; brown et al. 2006; environment canada and park service 1991; graefe, kuss & vaske 1990; hockings 2003; hockings, carter & leverington 1998; hockings et al. 2006; jacobson et al. 2009; mccool et al. 2007; moore, smith & newsome 2003; nilsen & tayler 1997; pavese, leverington & hockings 2007; shelby & heberlein 1986). the similarity of many of these frameworks suggests that it may be more important to choose and apply any one of the frameworks than to struggle with the choice of which framework is ‘best’ (manning 2004).secondly, there is a growing body of research on indicators and standards of quality for managing outdoor recreation and tourism in parks and protected areas. these studies address both park resources (hammitt & cole 1998) and the quality of the visitor experience (manning 2011a) and use a diverse set of natural and social science research methods. indicators and standards of quality have begun to be compiled in several synthetic sources (manning 2011a; national park service 2013). thirdly, monitoring protocols have been developed and applied in diverse settings and these can guide development and application of monitoring of visitor use of parks and protected areas, as well as the ecological and experiential impacts of this use (newsome, moore & dowling 2013). for example, handbooks for monitoring visitor use and associated encounters amongst visitors in wilderness areas have been developed (watson, cronn & christensen 1998; watson et al. 2000). similarly, guidelines for monitoring visitor impacts on trails, campsites and related areas have been developed (leung & monz 2006; marion & leung 2011; monz & d’luhosch 2010). monitoring incorporates a range of techniques, including automated counters, visitor surveys, remote sensing, global positioning systems and simulation modelling (cole 2005; gimblett & skov-peterson 2008; kim & daigle 2011; lawson 2006; newsome et al. 2013). fourthly, a substantial body of research is addressing the effectiveness of alternative management practices applied to parks and protected areas (buckley 2004; manning 2011a). this body of work has identified a range of management strategies and practices, tested the effectiveness of these practices in diverse contexts and has begun to develop a series of principles to guide the application of management practices (e.g. manning 2003, 2011a). moreover, a recent book has developed a matrix-based approach to considering alternative strategies and practices, along with a series of 20 case studies of effective management of outdoor recreation and tourism in national parks (manning & anderson 2012). fifthly, the research illustrated above has been conducted in a number of parks and protected areas around the world and this has contributed to the growing body of scientific and professional literature on management of parks and protected areas in diverse environmental, social and managerial contexts. emergence of a number of books that integrate and synthesise this body of work is an encouraging sign of the growing maturity of the field of park and protected area management for outdoor recreation and tourism (buckley 2004, 2009; eagles & mccool 2002; hammitt & cole 1998; hsu et al. 2010; manning 2007, 2011a; mccool & moisey 2008; newsome et al. 2013). this growing body of scientific and professional work is helping build the capacity of park and protected area management. close collaboration between managers and scientists will aid in successful management of parks and protected areas for outdoor recreation and tourism (manning 2011a; mccool et al. 2007). ethical considerations top ↑ some of the data reported in this article were collected as part of surveys of visitors to several units of the us national park system. these surveys were approved by institutional review committees of the principal investigator’s university. these surveys were administered on a voluntary basis and potential respondents were told the purpose of the surveys and approximately how long they would take. response rates were very high as most visitors to the national parks are inherently interested in how these areas are managed and are pleased to report on their experiences and attitudes toward park management issues. respondents’ names are not recorded to assure anonymity and confidentiality. trustworthiness top ↑ the reliability and validity of the data reported in this article are supported by publication of findings from these and related studies in many papers published in the peer-reviewed scientific literature. all of these papers have been reviewed by a minimum of three external reviewers who have examined all facets of these studies, including reliability and validity. conclusion top ↑ parks and protected areas have seen substantial growth in recent decades. however, management of these areas requires careful attention to maximise their potential environmental and societal values and to minimise potential conflict between public use of parks and protected areas and protection of park resources and the quality of the visitor experience. a recent study suggests that there is considerable room for improvement in management effectiveness (leverington et al. 2010). based on compilation and analysis of over 8000 assessments of park and protected area management across the world, the authors conclude that 42% of cases were judged to have major deficiencies. however, application of research was found to be positively correlated with management effectiveness. this suggests that management should be based on the growing body of scientific and professional literature.outdoor recreation and tourism are increasingly important uses of parks and protected areas. however, they can also challenge park managers to minimise the potential impacts of tourism and outdoor recreation on park resources and the quality of the visitor experience. management-by-objectives frameworks have emerged from the scientific and professional literature that can guide planning and management of parks and protected areas for outdoor recreation and tourism. these management frameworks draw on the broader literature in environmental planning and management and can be supported by a programme of research. the examples of this type of research outlined in this article suggest ways in which research can help inform each of the three primary steps of the management-by-objectives framework, (1) formulating indicators and standards of quality, (2) monitoring indicators of quality and (3) implementing management actions to help ensure that standards of quality are maintained. these and related programmes of research around the world can help inform management of parks and protected areas for outdoor recreation and tourism. acknowledgements top ↑ competing interests the author declares that he has no financial or personal relationships that may have inappropriately influenced him in writing this article. references top ↑ agee, j. & johnson, d., 1987, ecosystem management for parks and wilderness, university of washington press, seattle.baldwin, e. & lapage, w., 2003, visitor behaviors and resource impacts at cadillac mountain acadia national park, part ii: sign comparison study summer 2002, technical report for acadia national park natural resources division. balmford, a., beresford, j., green, j., naidoo, r., walpole, m. & manica, a., 2009, ‘a global perspective on trends in nature-based tourism’, plos biol 7(6), e1000144. http://dx.doi.org/10.1371/journal.pbio.1000144 belokurov, a., besancon, c., pavese, h., burgess, n.d., dudley, n., stolton, s. et al., 2009, ‘new resources for assessing the effectiveness of management in protected areas’, oryx 43(1), 14. brown, p., driver, b. & mcconnell, c., 1978, ‘the opportunity spectrum concept in outdoor recreation supply inventories: background and application’, proceedings of the integrated renewable resource inventories workshop, usda forest service general technical report rm-55, pp. 73–84. brown, g., koth, b., kreag, g. & weber, d., 2006, managing australia’s protected areas: a review of visitor anagement models, frameworks, and processes, sustainable tourism cooperative research center, gold coast. buckley, r. (ed.), 2004, environmental impacts of ecotourism, cabi publishing, wallingford. http://dx.doi.org/10.1079/9780851998107.0000 buckley, r., 2009, ecotourism principles and practices, cambridge university press, cambridge. christensen, n., bartuska, a., brown, j., carpenter, s., d’antonio, c., francis, r. et al., 1996, ‘report of the ecological society of america committee on the scientific basis for ecosystem management’, ecological applications 6, 665–691. http://dx.doi.org/10.2307/2269460 clark, r. & stankey, g., 1979, the recreation opportunity spectrum: a framework for planning, management, and research, usda forest service research paper pnw-98. cole, d.n. (compiler), 2005, computer simulation modeling of recreation use: current status, case studies, and future directions, general technical report rmrs-gtr-143, us department of agriculture forest service, rocky mountain research station, fort collins. davenport, l., brockelman, w.y., wright, p.c., ruf, k. & rubio del valle, f.b., 2002, ‘ecotourism tools for parks’, in j. terborgh, c. van schaik, l. davenport & m. rao (eds.), making parks work, pp. 279–306, island press, washington, dc. driver, b. & brown, p., 1978, ‘the opportunity spectrum concept in outdoor recreation supply inventories: a rationale’, proceedings of the integrated renewable resource inventories workshop, usda forest service general technical report rm-55, pp. 73–84. eagles, p.f.j. & mccool, s.f., 2002, tourism in national parks and protected areas: planning and management, cabi publishing, wallingford. http://dx.doi.org/10.1079/9780851995892.0000 environment canada and park service, 1991, selected readings on the visitor activity management process, environment canada, ottawa. evans, c.b., 2002, summit steward 2002 season report, acadia national park, maine. floyd, m., 1999, ‘race, ethnicity and use of the national park system’, social science research review 1(2), 1–23. frissell, s. & stankey, g., 1972, ‘wilderness environmental quality: search for social and ecological harmony’, proceedings of the society of american foresters annual conference, hot springs, 01–05 october 1972, pp. 170–183. gimblett, r. & skov-peterson, h. (eds.), 2008, monitoring, simulation, and management of visitor landscapes, university of arizona press, tucson. goodwin, h.j., 1996, ‘in pursuit of ecotourism’, biodiversity conservation 5, 277–291. http://dx.doi.org/10.1007/bf00051774 graefe, a., vaske, j. & kuss, f., 1984, ‘social carrying capacity: an integration and synthesis of twenty years of research’, leisure sciences 8, 275–295. http://dx.doi.org/10.1080/01490408609513076 graefe, a.r., kuss, f.r. & vaske, j.j., 1990, visitor impact management: the planning framework, vol. 2, national parks and conservation association, washington, dc. grumbine, r., 1994, ‘what is ecosystem management?’, conservation biology 8, 27–38. http://dx.doi.org/10.1046/j.1523-1739.1994.08010027.x hammitt, w.e. & cole, d.n., 1998, wildland recreation: ecology and management, wiley, new york. hardin, g., 1968, ‘the tragedy of the commons’, science 162, 1243–1248. http://dx.doi.org/10.1126/science.162.3859.1243, pmid:17756331 hockings, m., carter, b. & leverington, f., 1998, ‘an integrated model of public contact planning for conservation management’, environmental management 22(5), 643–654. http://dx.doi.org/10.1007/s002679900136, pmid:9680534 hockings, m., 2003, ‘systems for assessing the effectiveness of management in protected areas’, bioscience 53(9), 823–832. http://dx.doi.org/10.1641/0006-3568(2003)053%5b0823:sfateo%5d2.0.co;2 hockings, m., stolton, s., dudley, n., leverington, f. & courrau, j., 2006, evaluating effectiveness: a framework for assessing the management of protected areas, 2nd edn., iucn, gland. http://dx.doi.org/10.2305/iucn.ch.2006.pag.14.en holling, c., 1978, adaptive environmental assessment and management, john wiley & sons, london. honey, m., 2008, ecotourism and sustainable development, 2nd edn., island press, washington, dc. hsu, y-c., leung, y-f. li, y-h., wang, c-p. & lue, c-c., 2010, an assessment of frameworks useful for public land recreation planning, hua li publishing, taipei. international union for conservation of nature & united nations environment programme world conservation monitoring centre, 2011, the world database on protected areas (wdpa): january 2011, unep-wcmc, cambridge. international union for conservation of nature & united nations environment programme world conservation monitoring centre, 2012, the world database on protected areas (wdpa): february 2012, unep-wcmc, cambridge. jacobson, c., hughey, k.f.d., allen, w.j., rixecker, s. & carter, r.w., 2009, ‘toward more reflexive use of adaptive management’, society & natural resources: an international journal 22(5), 484–495. http://dx.doi.org/10.1080/08941920902762321 kim, m-k. & daigle, j.j., 2011, ‘detecting vegetation cover change on the summit of cadillac mountain using multi-temporal remote sensing datasets: 1979, 2001, and 2007’, environmental monitoring and assessment 180, 63–75. http://dx.doi.org/10.1007/s10661-010-1772-1, pmid:21082342 lawson, s., manning, r., valliere, w. & wang, b., 2003, ‘proactive monitoring and adaptive management of social carrying capacity in arches national park: an application of computer simulation modeling’, journal of environmental management 68, 305–313. http://dx.doi.org/10.1016/s0301-4797(03)00094-x lawson, s.r., 2006, ‘computer simulation as a tool for planning and management of visitor use in protected natural areas’, journal of sustainable tourism 14(6), 600–617. http:// http://dx.doi.org/10.1007/s00267-010-9564-5, pmid:20859627 manning, r., 2001, ‘visitor experience and resource protection: a framework for managing the carrying capacity of national parks’, journal of park and recreation administration 19, 93–108. manning, r., 2003, ‘emerging principles for using information/education in wilderness management’, international journal of wilderness 9(1), 20–27. manning, r., 2004, ‘recreation planning frameworks’, in j.j. vaske (ed.), society and natural resources: a summary of knowledge, pp. 83–96, modern litho, jefferson. manning, r., 2007, parks and carrying capacity: commons without tragedy, island press, washington, dc. manning, r., 2009, parks and people: managing outdoor recreation at acadia national park, university of vermont press, burlington. manning, r., 2011a, studies in outdoor recreation, oregon state university press, corvallis. manning, r., 2011b, ‘frameworks for defining and managing the wilderness experience’, in d.n. cole (compiler), wilderness visitor experiences: progress in research and management, pp. 158–176, proceedings rmrs-p-66, us department of agriculture forest service rocky mountain research station, fort collins. manning, r. & anderson, l., 2012, managing outdoor recreation: case studies in the national parks, cabi publishing, wallingford. http://dx.doi.org/10.1079/9781845939311.0000 manning, r. & freimund, w., 2004, ‘use of visual research methods to measure standards of quality for parks and outdoor recreation’, journal of leisure research 36(4), 557–579. manning, r., freimund, w., lime, d. & pitt, d., 1996, ‘crowding norms at frontcountry sites: a visual approach to setting standards of quality’, leisure sciences 18, 39–59. http://dx.doi.org/10.1080/01490409609513271 manning, r., valliere, w., wang, b., lawson, s. & newman, p., 2002, ‘estimating day use social carrying capacity in yosemite national park’, leisure 27(1–2), 77–102. marion, j. & leung, y-f., 2011, ‘indicators and protocols for monitoring impacts of formal and informal trails in protected areas’, journal of tourism & leisure studies 17(2), 215–236. mastny, l., 2001, treading lightly: new paths for international tourism, worldwatch institute, washington, dc. mccool, s., 1994, ‘planning for sustainable nature dependent tourism development: the limits of acceptable change system’, tourism recreation research 19, 51–55. mccool, s. & cole, d. (eds.), 1997, proceedings – limits of acceptable change and related planning processes: progress and future direction, missoula, montana, 20–22 may 1997, general technical report int-371, us department of agriculture forest service intermountain research station, ogden. mccool, s.f., clark, r.n. & stankey, g.h., 2007, an assessment of frameworks useful for public land recreation planning, general technical report pnw-gtr-705, us department of agriculture forest service, pacific northwest research station, portland. mccool, s. & moisey, r.n., 2008, tourism, recreation and sustainability: linking culture and environment, 2nd edn., cabi, cambridge. http://dx.doi.org/10.1079/9781845934705.0000 moore, s.a., smith, a.j. & newsome, d.n., 2003, ‘environmental performance reporting for natural area tourism: contributions by visitor impact management frameworks and their indicators’, journal of sustainable tourism 11(4), 348–375. http://dx.doi.org/10.1080/09669580308667211 monz, c. & d’luhosch, p., 2010, ‘a simplified image analysis method for monitoring wilderness campsites’, international journal of wilderness 16(1), 26–31. national park service, 1995, visitor experience and resource protection implementation plan: arches national park, denver service center, denver. national park service, 1997, verp: the visitor experience and resource protection (verp) framework – a handbook for planners and managers, denver service center, denver. national park service, 2013, national park service park planning, viewed 17 september 2013, from http://parkplanning.nps.gov/planninghome.cfm newsome, d., moore, s.a. & dowling, r.k., 2013, natural area tourism: ecology, impacts, and management, 2nd edn., channel view publications, bristol. nilsen, p. & tayler, g., 1997, ‘a comparative analysis of protected area planning and management frameworks’, in s.f. mccool & d.n. cole (eds.), proceedings – limits of acceptable change and related planning processes: progress and future direction, missoula, montana, 20–22 may 1997, general technical report int-371, us department of agriculture forest service intermountain research station, ogden, pp. 49–57. park, l., manning, r., marion, j., lawson, s. & jacobi, c., 2008, ‘managing visitor impacts in parks: a multi-method study of the effectiveness of alternative management practices’, journal of park and recreation administration 26(1), 97–121. pavese, h.b., leverington, f. & hockings, m., 2007, ‘global study of protected areas management effectiveness: the brazilian perspective’, natureza & conservacao 5(1), 152–162. shelby, b. & heberlein, t., 1986, carrying capacity in recreation settings, oregon state university press, corvallis. society of american foresters, 1993, sustaining long-term forest health and productivity, society of american foresters, bethesda. solop, f., hagen, k. & ostergren, d., 2003, ethnic and racial diversity of national park system visitors and non-visitors, national park service technical report, viewed 16 august 2010, from http://www.nature.nps.gov/socialscience/docs/archive/ethnicandracialdiversity.pdf stanfield mccown, r., laven, d., manning, r. & mitchell, n., 2012, ‘engaging new and diverse audiences in the national parks: an exploratory study of current knowledge and learning needs’, the george wright forum 29(2), 129–138. stankey, g., clark, r. & bormann, b., 2005, adaptive management of natural resources: theory, concepts and management institutions, usda forest service general technical report pnw-gtr-654. stankey, g., cole, d., lucas, r., peterson, m., frissell, s. & washburne, r., 1985, the limits of acceptable change (lac) system for wilderness planning, usda forest service general technical report int-176. stankey, g. & manning, r., 1986, ‘carrying capacity of recreation settings’, a literature review: the president’s commission on americans outdoors, pp. m-47–m-57, us government printing office, washington, dc. turner, r. & lapage, w., 2001, visitor behaviors and resource impacts at cadillac mountain, acadia national park, cooperative agreement 1443ca4520-a-0015, us department of the interior, washington, dc. wagar, j.a., 1964, ‘the carrying capacity of wild lands for recreation’, forest science monograph 7, n.p. walters, c., 1986, adaptive management of renewable natural resources, macmillan, new york. watson, a.e., cronn, r. & christensen, n.a., 1998, monitoring inter-group encounters in wilderness, research paper rmrs-rp-14, us department of agriculture forest service, rocky mountain research station, fort collins. watson, a.e., cole, d.n., turner, d.l. & reynolds, p.s., 2000, wilderness recreation use estimation: a handbook of methods and systems, general technical report rmrs-gtr-56, us department of agriculture forest service rocky mountains research station, ogden. whittaker, d., shelby, b., manning, r., cole, d. & haas, g., 2011, ‘capacity reconsidered: finding consensus and clarifying differences’, journal of park and recreation administration 29(1), 1–20. world travel and tourism council, 2007, the global travel and tourism summit, world travel and tourism council, london. abstract introduction material and methods results discussion acknowledgements references about the author(s) francois roux department of zoology, university of johannesburg, south africa scientific services, mpumalanga tourism and parks agency, south africa gert steyn department of zoology, university of johannesburg, south africa clinton hay department of biological sciences, university of namibia, namibia ina wagenaar department of zoology, university of johannesburg, south africa citation roux, f., steyn, g., hay, c. & wagenaar, i., 2018, ‘movement patterns and home range size of tigerfish (hydrocynus vittatus) in the incomati river system, south africa ’, koedoe 60(1), a1397. https://doi.org/10.4102/koedoe.v60i1.1397 original research movement patterns and home range size of tigerfish (hydrocynus vittatus) in the incomati river system, south africa francois roux, gert steyn, clinton hay, ina wagenaar received: 21 apr. 2016; accepted: 05 apr. 2018; published: 27 june 2018 copyright: © 2018. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract historical data suggested that the tigerfish (hydrocynus vittatus) of the incomati river migrates upstream and downstream as part of their life history. it has been suggested that this movement was a prerequisite for successful spawning in inundated floodplains in mozambique. recent advances in aquatic radio telemetry provided a reliable mechanism to monitor fish movement and increase knowledge of the ecology of tigerfish. from 04 january 2003 to 22 december 2003, 41 tigerfish in the incomati river system were fitted with radio transmitters to record movement patterns and estimate home range size. on average, each fish was tracked 72 times, and the total number of fixes was 2971 over the study period, including 1322 summer fixes and 1649 winter fixes. the mean longest distance travelled by tigerfish was 730 m (range = 75 m to 3200 m). the home range size varied between individual fish, but on average fish stayed within a defined home range of 48 846 m2. tigerfish showed high site fidelity to specific habitats within specific activity zones and movement occurred primarily within these defined zones. differences in movement pattern, longest distance travelled and home range size could not be attributed to the sex or size of the fish. no large-scale movement patterns associated with specific life history activity were observed; thus, previous reports of large-scale downstream migrations and spawning migrations appear to be invalid. the presence of weirs in the study area impedes free fish movement as these weirs create migration obstructions. conservation implications: river regulation such as damming, water abstraction, obstructive barriers and channel modification may have a detrimental impact on the survival strategy of this species. implementation of these results in a management policy will provide a reliable basis for species specific requirements such as upstream reservoir release management; minimum flow volumes required for downstream ecosystem maintenance and management and planning of structures obstructing natural flow. introduction the freshwater fish genus hydrocynus is represented by six species, all endemic to africa. they are pikelike predators, commonly termed ‘tigerfishes’ for their prominent dentition and dark lateral stripes (gery 1977). in southern africa, one of these species, hydrocynus vittatus (commonly known as tigerfish), occurs in the zambezi and okavango rivers and in the lowveld reaches of coastal systems (skelton 2001). the southern african tigerfish (h. vittatus) has a limited distribution in south africa, where it is restricted to the lowveld reaches of the limpopo river system, mainly within the kruger national park (knp), and further south in the lower reaches of the usutho and phongolo rivers (gaigher 1967). the incomati river system (south africa) is a marginal area in the distribution range of tigerfish where they occur in relatively low abundance. being essentially a lowveld species in south africa, it is intolerant to cold water and migrates downstream to lower lying reaches of these rivers during winter where water temperatures are higher and more stable (pienaar 1978; steyn et al. 1996; van loggerenberg 1983; skelton 2001). mortalities caused by a sudden drop in temperature (< 16.0 °c) related to cold water in the incomati river were reported on several occasions (deacon 1991; gagiano 1997; personal observation by authors; van loggerenberg 1983). gagiano (1997) reported mortalities in the piet grobler dam in the knp at a temperature of 14.5 °c during the winter period. the habitat and environmental conditions in the incomati river system differ considerably from the favourable conditions present in the larger northern tropical river systems such as the zambezi river. tigerfish are inhabitants of open, well-oxygenated waters such as found in the larger rivers and lakes (pienaar 1978). in contrast to the larger rivers and lakes in the north of south africa, the rivers of the knp are relatively small, highly regulated because of anthropogenic impacts and subject to extreme seasonal variations (du preez & steyn 1992; gertenbach 1991). variation and flow volumes, especially in the presence of instream damming structures such as weirs, can severely impact the ability of fish species to migrate in accordance with their life history requirement (baras & lucas 2002). furthermore, all the major rivers of the knp are subjected to high silt loads which can severely reduce dissolved oxygen concentrations of the water and may be lethal to fish (buermann et al. 1995). there has been a long history of fish mortalities in the knp caused by large amounts of suspended particles present in the water (knp annual reports 1946–1992). the negative impact of increased silt loads on the aquatic macro-invertebrate diversity in the major rivers of the knp was reported by moore and chutter (1988). sub-lethal effects of suspended solids on fish are varied and include negative impacts on reproduction, egg survival, growth, oxygen consumption, haematology, feeding and social behaviour (crouse, callahan & malaug 1981; wilber 1983). indirect effects include reduced food availability, clogging of gillrakes and filaments, reduced growth rate, reduced resistance to disease and disturbances of natural movements and migrations of fish (albaster & lloyd 1980; bruton 1985). tigerfish has a prominent ecological status as top predator, sharing the same trophic level as crocodiles in the knp riverine ecosystems. their limited presence in the knp and their vulnerability to impacts described above served as motivation for several studies since the work of gaigher (1967). in south africa, research on tigerfish concentrated on ecological aspects (gaigher 1970, 1973; 1975; gagiano 1997; van loggerenberg 1983), reproduction (steyn 1993; steyn & van vuren, 1992; steyn et al. 1996), tooth replacement (gagiano, steyn & du preez 1996), age estimation and maturity (gerber et al. 2009) and genetics (kotze et al. 1998). recent advances in aquatic radio telemetry provided a reliable means to acquire further information on the behaviour ecology of fish species and to improve our knowledge on tigerfish. despite several comprehensive studies as mentioned above, conservationists and river managers were still left with key questions on the (1) migrational requirements, (2) movement patterns and (3) ability to overcome obstructions in order to maintain functionality of a viable tigerfish population in the incomati river system. the objective of this study was to use biotelemetry to answer these key questions. material and methods description of the study area the incomati river drains parts of mpumalanga, swaziland and mozambique between the limpopo river system in the north and the phongolo river system in the south. it is economically one of the most important river basins in south africa, and it consists of three adjacent sub-basins: the komati, crocodile and sabie (darwall et al. 2009). the main river descends from the highland plateau in mpumalanga and swaziland and flows through the coastal plains of mozambique to the indian ocean just north of maputo at villa laisa. the total basin area is about 46 800 km2 of which 63% is in south africa, 5% in swaziland and 32% in mozambique. the average discharge of the incomati river basin at the estuary is about 100 m3/s to 200 m3/s, corresponding to about 3600 million m3 per year, to which south africa contributes 82%, swaziland about 13% and mozambique about 4% (darwall et al. 2009). the study area includes two rivers, namely the crocodile river and the komati river, which join to form the incomati river below the border town of komatipoort. the crocodile river flows along the boundary of the knp, and at the confluence, the border extends across the river to also include the lower reach of the komati river (figures 1 and 2). below the confluence, the incomati river can be described as a meandering river, incised into a wide sandy river bed, and in some sections, it flows through multiple bedrock channels. the river varies between 40 m and 50 m wide, with mostly large sandy pools and occasional rapids and a few riffles (roux et al. 1990). collection and tagging were done upstream and downstream of the confluence between knp and tenbosch weirs and the low-water bridge in the komati river. the choice of the collection and tagging area was motivated by the relative abundance of tigerfish in this river reach. the ability of tigerfish to overcome obstructions and their various home ranges later defined the extent of the study area. historically, tigerfish distribution data would indicate that tigerfish occur up to an altitude of 300 m in the incomati river system. gaigher (1967) previously collected tigerfish in the crocodile river gauge close to the town of nelspruit and in the komati river close to the town of tonga on the border between south africa and swaziland. consequently, the experimental design made provision for long-distance tracking in relation to historical distribution in the incomati river system. figure 1: map indicating the location of the study area in the mpumalanga province of south africa, in close proximity to mozambique. the applicable rivers are illustrated and marked with the numerous weirs. figure 2: map of the study area indicating tagging sites of all 41 radio-tagged fish. collection and handling of the species collection and handling of fish were performed in such a manner as to minimise physical and physiological stress to the specimens (spedicato, lembo & marmulla 2005). tigerfish were caught using two techniques: rod and reel with artificial lures and fly-fishing, both using barbless hooks to reduce injury to fish and to facilitate quick release, thereby reducing lactic acid stress and ensuring survival after handling and release (gerber et al. 2017). tagging of fish in total, 41 sexually mature tigerfish were tagged with radio transmitters (advanced telemetric systems inc. ats, usa, 142 mhz–144 mhz) in 2003. as the sexing of h. vittatus is relatively difficult based on external characteristics, males were only positively identified if they were ripe and running and producing semen. large females in or close to the spawning season were easily sexed as they displayed characteristic body size, form and weight (gaigher 1967; gagiano 1997; g.j. steyn pers. comm., 2003). the standard length (sl) was measured (mm), and mass (g) of each specimen collected was determined using a measuring tape and a bogagrip (scale). following capture, fish were anaesthetised with 2-phenoxyethanol (0.3 ml/l), minimising hyperactivity and stress. the radio transmitters were selected from ats models f2040, f2130 and f2010 with trailing whip antennae and were externally attached to fish with two strands of orthopaedic wire (0.65 mm diameter) below the dorsal fin following thorstad, økland and heggeberget (2001). to facilitate rapid healing of the needle wounds, the tagged fish were placed in a terramycin bath (25 mg/l water) for 10 min prior to release. the deployment of the small f2040 transmitters made it possible to tag smaller fish because of the relatively low weight of the transmitter, but remaining within the 2% rule (brown et al. 1999; peake & mckinley 1997). all radio-tagged fish were released at their respective sampling points, and staggered deployment over several months allowed for continuous data retrieval over a full year period, consequently covering all seasons (table 1; figure 2). staggered deployment was necessary because of the limited lifespan of the transmitters. table 1: individual fish collection, release and radio-tagging data (home range and longest distances). table 1 (continues...): individual fish collection, release and radio-tagging data (home range and longest distances). fish tracking procedures fish were tracked using a challenger r2100 receiver and a four-element yagi antenna (ats inc.) over a 12-month period (04 january to 22 december 2003) on average every second day, covering both summer and winter periods. care was taken to minimise behavioural side-effects by keeping a reasonable distance from tagged fish (hocutt, seibold & jesien 1994). tracking was done on foot from the banks of the river by using the homing-in technique (jick 1979). if there was any uncertainty regarding the position of the fish, the triangulation method was then applied (jick 1979). in instances where fish were lost, aerial surveys were conducted using a micro-light aircraft to relocate a specific fish. for all tracking surveys, location was determined using a handheld global positioning system receiver (garmin etrax). upon detection, the global position system (gps) coordinates of the fish’s location were noted (accuracy ± 5 m). hydrology, water quality and meteorological data flow levels in the incomati river system were determined from daily readings at the knp gauging weir in the incomati river. water temperature, ph and conductivity were recorded daily in the crocodile river, komati river and below the confluence of the two rivers (in the incomati river) using eutech portable microprocessor-based water quality instruments. meteorological data were gathered from a nearby weather station (transvaal sugar board, komatipoort), including rainfall, minimum and maximum air temperatures. data analysis two fish that moved out of the study area into mozambique shortly after tagging were excluded from the analysis. in addition, a third fish showed no movement for an extended period after tagging and was presumed dead and excluded from the analysis. descriptive statistics for the entire study period (summer and winter) were based on more than 10 fixes per fish for 38 fish. gps coordinates of the radio-tracked tigerfish were used to calculate longest distances travelled and to determine home range sizes. bi-variate gaussian or normal distribution kernel methods (seaman & powell 1996; silwerman 1986; worton 1989) were used to plot home ranges. this group of methods is part of a more general group of parametric kernel methods that employ distributions other than the normal distributions as the kernel elements which are associated with each point in the set of location data. because of the meandering nature and relatively small width and limited available habitat within the incomati river system during low flow periods at specific sites, an adaptation of the simplified minimum convex polygon (mcp) (baker 2002; creel & creel 2002; meulman & klomp 1999) was used. boundaries of home ranges were drawn using different sets of location data (planet gis). this method of using the shoreline as a boundary of the home range is a widely accepted and commonly used method in fish telemetry experiments (hocutt et al. 1994). for ease of statistical analysis, a binning algorithm was implemented in which the longest distance travelled, home range size and the radio-tagged fish were grouped in classes according to their magnitude. for longest distance travelled (økland et al. 2005), fish were organised in classes ranging from 100 m to 500 m, 501 m to 1000 m, 1001 m to 1500 m, 1501 m to 2000 m and > 2000 m travelled. the home range size were classed in groups ranging from 0 m2 to 10 000 m2, 10 001 m2 to 20 000 m2, 20 001 m2 to 50 000 m2, 50 001 m2 to 100 000 m2 and > 100 000 m2. the ibm spss statistics 18 program was used for basic and inferential statistics which include frequencies, normality, correlations and comparisons (spss 2009). ethical consideration the project proposal was approved with ethical clearance by the faculty of science, university of johannesburg and mpumalanga parks and tourism (permit number mpb 8553.). results water quality, hydrology and meteorological data mean water temperature results in the incomati river system indicate that the minimum is reached in july (18.02 °c) after which temperatures gradually increase to a mean temperature of 24 °c during october. the highest mean monthly river water temperature during this study (30.61 °c) was recorded in the crocodile river during january (figure 3). the highest mean monthly river water temperature in the komati river (30.17 °c) was recorded during february. temperatures in the incomati river, below the confluence, were influenced by both tributaries, and consequently, the highest mean monthly temperature for the incomati river (28.88 °c) was recorded during february. figure 3: graph indicating water and air temperature for the months january–december 2003 in the incomati river. for the tigerfish active summer period, november to april, the mean monthly ph values varied between 8.1 and 8.5, whereas the conductivity fluctuated between 274 µs/cm and 622 µs/cm in the incomati river. summer conductivity values were lower than winter values, but summer ph values were higher. during summer, the turbidity levels increased as a result of the higher summer flows. although not measured, turbidity was observed to be closely associated with rainfall events in the catchment during the summer period. the highest rainfall recorded was during the months of november (115.4 mm) and february (191.7 mm). the mean monthly flow (figure 4) for the winter period (may–october) in the incomati river, when tigerfish are less active, varied between 0.44 m3/s and 1.89 m3/s compared to 0.82 m3/s and 13.12 m3/s for the summer period (november–april), when tigerfish are active. the highest flow spikes were recorded during the spawning season (october–february) in the summer period (steyn 1993; steyn & van vuren 1991; steyn et al. 1996). on three occasions, flow spikes in excess of 25.00 m3/s, with the largest of 51.76 m3/s, occurred in january (figure 4). figure 4: water flow in the incomati river over the period january–december 2003. radio-tagged fish in total, 41 fish were radio-tagged with a mean length (sl) of 62.7 cm (range 47 cm – 76 cm) and a mean weight of 2418 g (range 910 g – 4990 g) (table 1; figure 2). of the 41 radio-tagged fish, 11 (26.8%) were males and 30 (73.2%) were females in a 1:3 sex ratio. for the radio-tagged males, the length (sl) varied between 47 cm and 60 cm (mean = 55.4 cm) and the weight varied between 910 g and 2040 g (mean = 1605.5 g). for the radio-tagged females, the length (sl) ranged from 57 cm to 76 cm (mean = 65.4 cm) and the weight ranged from 1810 g to 4990 g (mean = 2717 g) (table 2). table 2: summary of average fish length, number of fixes, longest distance travelled and home range. movement the total distance of the river where adult fish were captured and equipped with radio tags measured 5.2 km. after capture, tagging and the associated disturbance to a fish when released, the fish normally moved upstream or downstream and normally only returned 2 to 5 days later to the original tagging site, thereby suggesting site fidelity. the distance moved directly after tagging varied over the 2to 5-day period from 48 m to 1038 m. in total, 35 (85.4%) of the fish tagged returned to the original tagging site within the mentioned period, but 6 (14.6%) never returned, 3 of which moved downstream into mozambique and were not recorded again. this showed angling in the form of catch and release may be a major disturbance, but this also confirmed site fidelity of tigerfish to a specific home range. the gps coordinates of each sample or release site, tag number, type of tag and size, weight and sex of each fish are presented in table 1. over time, a movement pattern emerged for each of the 41 radio-tagged fish, and the longest distances travelled and home ranges could be determined (table 1). on average, fish were tracked 72.5 times (table 2) and the total number of fixes was 2971 for the period 04 january 2003 to 22 december 2003. some individuals were tracked up to 161 times. the maximum total of fixes (n = 161) per individual was associated with a tag life of 10 months. for the summer period (january–april, november and december 2003), there were 1322 fixes, and for the winter period (may–october 2003), there were 1649 fixes. for the summer period (or part thereof), there were 40 active radio-tagged fish, but only 32 active radio-tagged fish for the winter period (or part thereof). the mean number of fixes for females was 78.4 (n = 30) per fish with a range of 6–161. the mean number of fixes for males was 56.2 (n = 11) per fish with a range of 7–110. the reason for the lower amount of fixes for males (56.2 fixes) in comparison with females (78.4 fixes) can be ascribed to the differences in radio tag types used. as males are generally smaller than females, smaller f2040 radio tags, with a much shorter lifespan (94 days), were used to stay within the 2% rule. longest distance travelled for the statistical analysis, data were obtained from 38 radio-tagged tigerfish with more than 10 fixes. the mean longest distance travelled (n = 38) was 729.66 m (table 2) with a range from 74.5 m to 3200 m. when analysing the longest distance travelled by the different radio-tagged fish, 47.4% (18 out of the 38 fish) travelled between 100 m and 500 m, 34.2% (13 fish) between 501 m and 1000 m, 10.5% (4 fish) between 1001 m and 1500 m, 5.3% (2 fish) between 1501 m and 2000 m and 2.6% (1 fish) travelled more than 2000 m (table 2; figure 5). figure 5: clustering of longest distance travelled of radio-tagged tigerfish for the period january–december 2003. a, 0 m–500 m; b, 501 m–1000 m; c, 1001 m–1500 m; d, 1501 m–2000 m; e, > 2000 m. when distinguishing between the different sexes and longest distance travelled, 46.4% of females (13 out of 28 fish) travelled between 100 m and 500 m, 39.3% (11 fish) between 501 m and 1000 m, 7.1% (2 fish) between 1001 m and 1500 m, 3.6% (1 fish) between 1501 m and 2000 m and 3.6% (1 fish) travelled more than 2000 m. for the males, 50% (5 out of 10 fish) travelled between 100 m and 500 m, 20% (2 fish) between 501 m and 1000 m, 20% (2 fish) between 1001 m and 1500 m and 10% (1 fish) between 1501 m and 2000 m (table 2). the furthest movement recorded was 3200 m over a 3-day period. this female moved out of its known home range (18 fixes) and established a new home range approximately 3018 m upstream (table 2). for females, the mean longest distance travelled was 734.4 m (n = 28) with a range of 74.5 m to 3200 m, and for males, the mean longest distance travelled was 716.3 m (n = 10) with a range of 148.9 m to 1800 m. no significant differences were found between males and females for longest distances travelled (mann–whitney u test, mean ranking males 19.8 and females 18.6, p = 0.753). three different tigerfish movement patterns were recorded (figure 6). movement patterns were obtained from a combined effect of distance travelled and home range sizes (figure 7). although all the fish displayed some degree of site fidelity within a specific activity zone, movement pattern 1 represents fish that moved 100 m to 500 m within a well-defined home range, and movement occurred only within this specific home range. movements of fish number 8 serves as example for this type of movement pattern (figure 8). the majority (47.37%) of the radio-tagged fish displayed characteristics of movement pattern 1 (figure 6, cluster a). movement pattern 2 represents fish that displayed site fidelity for two or more areas within a larger well-defined home range, spanning a distance of 501 m to 1500 m. movements of fish number 15 serve as example for this type of movement pattern (figure 9). this group was represented by 44.7% of radio-tagged fish (figure 6, cluster b). movement pattern 3 represents fish that showed little site fidelity and would temporarily occupy small areas within a large undefined home range that spans more than 1500 m. movements of fish number 23 serve as example for this type of movement (figure 10). fish within the latter group can be seen as vagrants without established home ranges for a specific period. most of these fish were also later lost as they moved out of the study area and could not be relocated. fish in this group were large females of weight ranging between 2720 g and 3580 g and represented 7.89% of the radio-tagged fish (figure 6, cluster c). figure 6: scatter graph depicting three different movement patterns by radio-tagged tigerfish related to home range size and longest distance travelled for the study period january–december 2003. figure 7: clustering of home range sizes of radio-tagged fish. a, 0 m2–10 000 m2; b, 10 001 m2–20 000 m2; c, 20 001 m2–50 000 m2; d, 50 001 m2–100 000 m2; e, > 100 000 m2. figure 8: map of radio-tagged fish number 8 indicating home range and longest distance travelled (type 1 movement pattern). figure 9: map of radio-tagged fish number 15 indicating home range and longest distance travelled (type 2 movement pattern). figure 10: map of radio-tagged fish number 23 indicating home range and longest distance travelled (type 3 movement pattern) for a detailed account of the movement patterns and demarcation of the home ranges of each of the 41 radio-tagged fish, see roux (2013). the dots indicate individual fixes during tracking and the contours around the fixes indicate the defined home range. home range sizes the home range size varied between individual fish with 38.2% (13 fish) localising within an area between 0 m2 and 10 000 m2 (mean = 5567.95 m2) and 18.42% (7 fish) localising within an area between 10 001 m2 and 20 000 m2 (mean = 14 435.53 m2). furthermore, 18.42% (7 fish) occupied a home range area between 20 001 m2 and 50 000 m2 (mean = 31 092.2 m2), whereas 10.5% (4 fish) occupied an area between 50 001 m2 and 100 000 m2 (mean = 79 809.55 m2) and 18.42% (7 fish) utilised an area > 100 000 m2 (mean = 163 692.90 m2) (table 2; figure 7). on average, the fish (n = 38) stayed within a defined home range of 48 846.36 m2. the home range size for males and females compared favourably with a mean of 53 296.52 m2 (n = 28) and a range from 331.6 m2 to 236 496 m2 for females and a mean of 36 385.9 m2 (n = 10) and a range from 1338.8 m2 to 135 982.6 m2 for males. no statistically significant differences were found between the sexes for their home range size (mann–whitney u test, mean ranking females = 20.71 and males = 16.10, p = 0.260). migration obstructions none of the 41 tagged fish crossed the tenbosch weir. three individuals, namely numbers 7, 12 and 18, moved upstream in the crocodile river to be briefly recorded in the vicinity of this weir. the tenbosch weir has a crest height of 2 m and a fish ladder constructed at the side of the weir. this ladder is of the vertical slot type and appears to be non-functional to fish migration in general. only two radio-tagged fish (fish numbers 15 and 39) ventured into the lower komati river, close to the confluence with the crocodile river, where they were confined in a pool below the low-water bridge for a few days. they were not able to overcome this obstacle. this low-water bridge at komatipoort was constructed on a natural dolerite intrusion that stretches across the river. contrary to the above, a total number of 16 crossings, both upstream and downstream, were recorded at the knp weir. this gauging weir has a crest height of approximately 1.2 m with a well-designed fish way to facilitate fish movement at medium to high flow conditions. tagged fish with allocated numbers 1, 4, 6, 20, 27 and 37 crossed the knp weir downstream and upstream over the period january to march, whereas fish 19 crossed the knp weir downstream during july and returned upstream three days later. fish numbers 1 and 27 each crossed on three occasions, whereas fish number 20 crossed the knp weir on four occasions with only a few day intervals between upstream and downstream crossings. numerous visual observations were made of untagged tigerfish jumping over this weir over the duration of this study. successful crossing at the knp weir occurred at flow velocities between 1.78 m³/s and 16.2 m³/s (table 3). table 3: successful crossings of radio-tagged tigerfish at the kruger national park weir. discussion this study confirmed that external tagging attachment protocol (thorstad et al. 2001) was suitable for the study of tigerfish behavioural ecology through biotelemetry in that only a single mortality was recorded from the 41 radio-tagged fish. furthermore, visual observations of radio-tagged fish swimming just below the surface were made on numerous occasions and fouling of radio tags appeared to be minimal, thus having no significant effect on the swimming capabilities or movement patterns of tagged fish. after capture, tagging and the associated disturbance to a fish, it normally moved either upstream or downstream and returned 2–5 days later to the original tagging site, thereby confirming site fidelity. the distance moved directly after tagging varied over the 2to 5-day period from 48 m to 1038 m. in total, 35 of the tagged fish returned to the original tagging site within the mentioned time frame. six fish never returned; three of these moved downstream into mozambique and were lost. in general, tigerfish displayed high site fidelity to specific habitats within specific activity zones, and movement occurred primarily within these defined home ranges. the longest distance travelled by fish was during summer and early winter, when water temperatures exceeded 24 °c. these periods coincided with high water levels in the study area, which probably facilitated movement between different habitats. some degree of site fidelity of h. vittatus was also reported by økland et al. (2005) for the upper zambezi, whereas consistent fidelity to an activity core was reported by baras et al. (2002) for hydrocynus brevis in the niger river, mali. during our study, little to no movement was recorded in the winter months when water temperatures were below 24 °c. the mean lowest temperature recorded in the incomati river system of 18 °c is close to the minimum temperature range for the survival of tigerfish. during a tigerfish translocation exercise when laboratory-induced breeding was attempted, prior to successful breeding at skukuza (steyn et al. 1996), a temperature drop from 27 °c to 18 °c during a 4-hour transport period killed almost all of the fish. the mean longest distance travelled during this investigation was relatively short (729.66 m). in the zambezi, two movement patterns were distinguished where approximately 50% of the fish moved < 1000 m among tracking surveys. the remaining fish showed consistent site fidelity for periods with long-distance movements (> 1000 m) to new areas among residency periods. in the incomati river system, only 18% of the fish displayed long-distance movement > 1000 m and the longest distance was 3200 m. the longest distance travelled in the incomati river system was relatively short in comparison with the longest distance of 18.8 km travelled in the zambezi river (økland et al. 2005). irrespective of the shorter distances travelled in the incomati river system, the total unobstructed river upstream to tenbosch was not utilised by all tagged individuals and the option to migrate downstream was available but not utilised. nevertheless, three movement patterns demonstrated by incomati river system tigerfish do not describe the dependency on upstream or downstream migration behaviour expected for this species in the study area. implicit of the relative short distances travelled, they are crucial for the survival of h. vittatus in the lower incomati river system. site fidelity and restricted mean home range (48 846 m2) in comparison with the much larger home range of zambezi tigerfish (276 978 m2), supported by various historical observations of their vulnerability to environmental stressors such as low temperature, low flow and high silt loads, are indicative of a population that does not function optimally on the edge of its distribution, in accordance with the law of tolerance (odum 1971). sub-optimal functionality of another tigerfish population in the knp is also reflected in the results of gagiano (1997), during an ecological investigation on tigerfish in the olifants and letaba rivers. tigerfish of all sizes in these rivers were found to feed almost exclusively on invertebrates. this finding is in contrast with the tigerfish from other systems, where fish play a major role in their diet (jackson 1961). differences in movement patterns, longest distance travelled and home range size could not be explained by sex or the size of the fish. tigerfish show opportunistic movement patterns, and home ranges can change in size and location as a result of seasonal shifts, prey availability, habitat availability and cover as well as life history requirements. no large-scale movement pattern or specific activity-related migrations were observed. thus, reports of large-scale migrations of tigerfish downstream into mozambique during winter in the incomati river (van loggerenberg 1982) seem to be no longer relevant, probably because of their limited numbers and because of suitable habitat created by the damming of the knp weir and subsequent deeper water bodies where the temperature is more stable to find refuge during winter. there was no evidence of upstream congregation of tigerfish at the tenbosch weir or large-scale downstream crossings at the knp weir. from the pattern of crossings at the knp weir, it is inferred that some of the marked fish that successfully crossed this weir responded to the stress associated with the tagging procedure and returned later to demonstrate site fidelity. these fish were tagged either just downstream or upstream of the knp weir, followed by a fleeing response over the weir (fish numbers 1, 6, 19 and 20). some of the crossings could be associated with higher flow conditions (fish numbers 4 and 37), whereas fish number 27 probably displayed natural behaviour as the crossing occurred more than a month after tagging. irrespective of the motivation for crossing the weir, in context with the life span of the tags for above fish, these events were limited to only a few occasions during a period of several months, which again displayed site fidelity. flow volumes that varied between 1.94 m³/s and 16.22 m³/s during successful crossings suggest that the knp weir is not a restrictive barrier to tigerfish and the population is open to gene flow from mozambique. contrary to this, our results suggest that tigerfish in the crocodile river, upstream from the tenbosch weir, is isolated; consequently, the upstream population cannot be replenished after mortalities because of extreme environmental conditions such as influx of cold water, low flow and high silt loads and will most probably disappear in this part of its historical distribution range. in the komati river, upstream movement is restricted close to the confluence at the low-water bridge, consequently isolating the upstream population in the komati river which currently is heavily subjected to water abstraction and agricultural activities. the isolation of upstream tigerfish populations in the incomati river system and their vulnerability to environmental impacts emphasise the ecological significance and inclusion of this river reach into the borders of the knp as well as the functionality and importance of the knp weir. based on the knowledge gained during this study on the behaviour of tigerfish, recommendations on the instream flow requirements (ifrs) of this species need to be adopted into the ecological flow requirements for the incomati river system and setting of the ecological reserve to ensure the ecological maintenance and functioning of the instream habitats utilised by tigerfish (kleynhans & engelbrecht 2000). environmental flow allocations and maintenance of ecological requirements of aquatic ecosystems are entrenched in the national water act (no 36 of 1989) and specified as components of the ecological reserve. within the framework of resource directed measures for protection of water resources, established by the department water affairs and sanitation (dwa), the implemented ecological reserve needs to be monitored and can be adjusted to meet the targets and resource quality objectives (king, tharme & de villiers 2000). acknowledgements the authors acknowledge de beers, venetia mines and barloworld for funding this project. they thank the incomati tigerfish action group (itag), domien and bart van buynder for their logistical support and for their enthusiasm for the tigerfish species and its conservation. they also thank peter kimberg and michael mashaba for their assistance with fieldwork and data collection. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions f.r. was the researcher who performed most of the sampling and data analysis. g.s. was the researcher who was responsible for the experimental design. c.h. was the co-worker who is a fish expert with experience in biotelemetry. i.w. was the promoter of the phd study, made conceptual contributions and proofread the manuscript. f.r. and g.s. wrote the manuscript. references albaster, j.s. & lloyd, r., 1980, water quality criteria for freshwater fish, fao, united nations, london. baker, j., 2002, ‘population density and home range estimates for the eastern bristlebird at jervis bay, south-eastern australia’, corella 25, 62–67. baras, e., togola, b., sicard, b. & benech, v., 2002, ‘behaviour of tigerfish hydrocynus brevis in the river niger, mali, as revealed by simultaneous telemetry of activity and swimming depth’, hydrobiologia 483, 103–110. https://doi.org/10.1023/a:1021359008246 baras, e. & lucas, m.c., 2002, ‘impacts of man’s modification of river hydrology on the migration of freshwater fishes: a mechanistic perspective’, ecohydrology and hydrobiology 146, 431–448. buermann, y., du preez, h.h., steyn, g.j., harmse, j.t. & deacon, a., 1995, ‘suspended silt concentrations in the lower olifants river (mpumalanga) and the impact of silt releases from the phalaborwa barrage on water quality and fish survival’, koedoe 38(2), 11–33. https://doi.org/10.4102/koedoe.v38i2.312 brown, r.s., cooke, s.j., anderson, w.g. & mckinley, r.s., 1999, ‘evidence to challenge the “2% rule” for biotelemetry’, north american journal of fisheries management 19, 867–871. https://doi.org/10.1577/1548-8675(1999)019<0867:etctrf>2.0.co;2 bruton, m.n., 1985, ‘the effects on suspensoids on fish’, hydrobiologia 125, 221–241. https://doi.org/10.1007/bf00045937 creel, s. & creel, n.m., 2002, the african wild dog: behaviour, ecology and conservation, princeton university press, new jersey. crouse, m.r., callahan, h.f. & malaug, k.w., 1981, ‘effects of fine sediments on growth of juvenile coho salmon in laboratory streams’, transactions american fisheries society 110, 281–286. https://doi.org/10.1577/1548-8659(1981)110<281:eofsog>2.0.co;2 darwall, w.r.t., smith, k.g., tweddle, d. & skelton, p.h., 2009, the status and distribution of freshwater biodiversity in southern africa, iucn and grahams town, south africa: saiab, gland, switzerland, p. 120. deacon, a., 1991, ‘“visvrektes in olifantsrivier” (fish mortality in olifants river)’, custos 18, 31. du preez, h.h. & steyn, g.j., 1992, ‘a preliminary investigation of the concentration of selected metals in the tissues and organs of tigerfish (hydrocynus vittatus) from the olifants river, kruger national park, south africa’, water sa 18(2), 131–136. gagiano, c.l., 1997, ‘an ecological study on the tigerfish hydrocynus vittatus in the olifants and letaba rivers with special reference to artificial reproduction’, unpublished master’s dissertation, rand afrikaans university, johannesburg, viewed 05 september 2014 from http://hdl.handle.net/10210/7031 gagiano, c.l., steyn, g.j. & du preez, h.h., 1996, ‘tooth replacement of tigerfish hydrocynus vittatus from the kruger national park’, koedoe 38(1), 1–6. gaigher, i.g., 1967, ‘aspects of the ecology of the tigerfish, hydrocynus vittatus castlenau in the incomati river system’, unpublished master’s dissertation, university of pretoria, pretoria. gaigher, i.g., 1970, ‘ecology of the tigerfish (hydrocynus vittatus) in the incomati river system, south africa’, zoologica africana 5(2), 211–227. https://doi.org/10.1080/00445096.1970.11447393 gaigher, i.g., 1973, ‘the habitat preferences of fishes from the limpopo river system, transvaal and mozambique’, koedoe 16, 103–116. https://doi.org/10.4102/koedoe.v16i1.888 gaigher i.g., 1975, ‘evidence for tooth replacement in the tigerfish hydrocynus vittatus’, arnoldia rhodesia 7(1), 1–4. gery, j., 1977, characoids of the world, t.f.h. publications, new york, p. 672. gerber, r., howatson, g., greenfield, r., wagenaar, i. & smit, n., 2017, ‘physiological response to angling of africa’s premier angling species, the tigerfish hydrocynus vittatus’, journal of african zoology 52(2), 91–98. https://doi.org/10.1080/15627020.2017.1300069 gerber, r., smit, n.j., pieterse, g.m. & durholtz, d., 2009, ‘age estimation, growth and size at sexual maturity of tigerfish hydrocynus vittatus from the okavango delta, botswana’, african journal of aquatic science 34(3), 239–247. gertenbach, w.p.d., 1991, problems facing kruger national park: first annual research meeting, 19–20 march 1991, programme report 2, 1–4, kruger national park, skukuza. hocutt, c.h., seibold, s.e. & jesien, r.v., 1994, ‘potential use of biotelemetry in tropical continental waters’, revue d’hydrobiologie tropicale, paris 27, 77–95. jackson, p.b.n., 1961, ‘the impact of predation, especially by tigerfish, hydrocynus vittatus, on african freshwater fishes’, proceedings of the zoological society london 136(4), 603–622. https://doi.org/10.1111/j.1469-7998.1961.tb05895.x jick, t.d., 1979, ‘mixing qualitative and quantitative methods: triangulation in action’, administrative science quarterly 24, 602–611. https://doi.org/10.2307/2392366 king, j.m., tharme, r.e. & de villiers, m.s., 2000, environmental flow assessments for rivers: manual for the building block methodology, wrc report no: tt 131/60, water research commission, pretoria, p. 340. kruger national park, annual reports, 1946–1992, unpublished report, national parks board. meulman, e.p. & klomp, n.i., 1999, ‘is the home range of the heath mouse pseudomys shortridgei an anomaly in the pseudomys genus?’, victorian naturalist 116, 196–201. moore, c.a. & chutter, e.m., 1988, a survey of the conservation status and benthic biota of the major rivers of the kruger national park, contract report national institute of water research, csir. odum, e.p., 1971, ‘fundamentals of ecology’, saunders, london. økland, f., thorstad, e.b., hay, c.j., naesje, t.f. & chanda, b., 2005, ‘patterns of movement and habitat use by tigerfish (hydrocynus vittatus) in the upper zambezi river, namibia’, ecology of freshwater fish 14, 79–86. https://doi.org/10.1111/j.1600-0633.2004.00080.x peake, s. & mckinley, r.s., 1997, ‘influence of transmitter attachment procedures on swimming performance of wild and hatchery-reared atlantic salmon smolt’, transactions of the american fisheries society 126, 707–714. https://doi.org/10.1577/1548-8659(1997)126<0707:iotapo>2.3.co;2 pienaar, u.v., 1978, the freshwater fishes of the kruger national park, national parks board of trustees, pretoria. roux, f., 2013, ‘a study on the behaviour of tigerfish (hydrocynus vitattus) using biotelemetry, to determine habitat utilisation and survival strategies in the lower incomati river system’, phd dissertation, university of johannesburg, johannesburg, south africa, viewed 29 january 2018 from http://hdl.handle.net/10210/12356 roux, f., kleynhans, c.j., thirion, c., hill, l., engelbrecht, j.s., deacon, a.r. et al., 1990, ‘adaptive assessment and management of riverine ecosystems; the crocodile/elands river case study’, water sa 25(4), 501–507. seaman, d.e. & powell, r.a., 1996, ‘an evaluation of the accuracy of kernel density estimators for home range analysis’, ecology 77, 2075–2085. https://doi.org/10.2307/2265701 silwerman, b.w., 1986, density estimation for statistics and data analysis, chapman and hall, london, uk. skelton, p.h., 2001, a complete guide to the freshwater fishes of southern africa, struik publishers, south africa. spedicato, m.s., lembo, g. & marmulla, g. (eds), 2005, aquatic telemetry: advances and applications, food and agricultural organisation of the united nations (fao), rome. spss inc., 2009, pasw statistics for windows, version 18.0, spss inc., chicago, il. steyn, g.j., 1993, ‘physico-chemical characteristics of tigerfish semen’, southern african journal of wildlife research 23, 44–47. steyn, g.j., gagiano, c.l., deacon, a.r. & du preez, h.h., 1996, ‘notes on the induced reproduction and development of tigerfish, hydrocynus vittatus (characidae), embryos and larvae’, environmental biology of fishes 47, 387–398. https://doi.org/10.1007/bf00005052 steyn, g.j. & van vuren, j.h.j., 1991, ‘cryopreservation of the spermatozoa of two african fishes (characidae)’, southern african journal of wildlife research 21, 76–81. thorstad, e.b., økland, f. & heggeberget, t.g., 2001, ‘are long term negative effects from external tags underestimated. fouling of an externally attached telemetry transmitter’, journal of fish biology 59, 1092–1094. https://doi.org/10.1111/j.1095-8649.2001.tb00174.x van loggerenberg, n.p., 1980, kunsmatige teelt van die tiervis hydrocynus vittatus castelnau, en die biologiese aspekte wat daarmee verband hou, projek tn 6/4/2/2/1/18, transvaal provinsiale vissery instituut, lydenburg. van loggerenberg, n.p., 1982, kunsmatige teelt van die tiervis hydrocynus vittatus castelnau, in transvaal, vierde projekverslag, projek tn 6/4/2/2/1/18, transvaal provinsiale vissery instituut, lydenburg. van loggerenberg, n.p., 1983, ‘conservation of tigerfish and fish farming techniques’, fauna and flora 40, 30–31. wilber, c.g., 1983, turbidity in the aquatic environment. an environmental factor in fresh and oceanic waters, cc thomas, springfield, illinois, usa. worton, b.j., 1989, ‘kernel methods for estimating the utilisation distribution in home range studies’, ecology 70, 164–168. https://doi.org/10.2307/1938423 article information authors: harry biggs1 sam ferreira1 stefanie freitag-ronaldson1 rina grant-biggs1 affiliations: 1scientific services, south african national parks, skukuza, south africa correspondence to: harry biggs email: biggs@sanparks.org postal address: private bag x402, skukuza 1350, south africa dates: received: 10 may 2010 accepted: 01 oct. 2010 published: 13 may 2011 how to cite this article: biggs, h., ferreira, s., freitag-ronaldson, s. & grant-biggs, r., 2011, ‘taking stock after a decade: does the ‘thresholds of potential concern’ concept need a socio-ecological revamp?’, koedoe 53(2), art. #1002, 9 pages. doi:10.4102/koedoe.v53i2.1002 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) taking stock after a decade: does the ‘thresholds of potential concern’ concept need a socio-ecological revamp? in this original research... open access • abstract • introduction • dealing with the barriers    • taking decisions and action in spite of uncertainty about a threshold or a limit    • can thresholds of potential concern be used to describe, or can they include, societal preferences?    • can thresholds of potential concern be adjusted to take embedded human behaviour, especially differences in the rate of institutional responses, into account? • towards a broader definition • conclusion • acknowledgements • references adstract (back to top) the concept of thresholds of potential concern (tpcs) as implemented for the last decade in strategic adaptive management in south african national parks (sanparks), has proved workable in practice in a number of instances, but in others appears beset by conceptual and practical limitations or barriers. three common challenges relate to (1) situations where there is uncertainty about whether and where real thresholds exist, (2) whether and how preferences and other social constructs, as opposed to what were seen as objective biophysical variables only, can be used for tpcs and (3) whether it is admissible to adjust tpcs to allow for variations in societal behaviour, in particular rate of management response. all three challenges arise in the face of tpc objectivity implied by the original definition, and in the light of the original view that tpcs be set some distance prior to a presumed ecological threshold. this paper suggests that the three challenges can be partly or largely dealt with by the use of a wider socio-ecological view, rather than seeing tpcs in isolation or as being only biophysical. also, while detection of abrupt changes is helpful, it makes little practical difference if some tpcs happen to describe linear processes. the very decision to intervene can induce an abrupt change. once a wider socio-ecological approach is employed, it becomes necessary for the user to specify the particular usage envisaged for the tpc, for instance, whether it is considered a preference and whether that preference is believed in any way to be related to an ecological threshold. in all cases, it is recommended that some form of explicit representation of the socio-ecological view is constructed – we suggest a cause-and-effect diagram (and give an example generated through a thought experiment) which describes presumed relationships in the subsystem of interest. this provides a broader systemic context and a shared understanding, and has implications for considering scenarios and management alternatives. for practical reasons, from the several states and processes in such a subsystem, only a few links can be chosen on which to base particular tpcs. if we have understood the subsystem well enough, these few links, at each of which a tpc is developed, will act as diagnostic points at which we can monitor the performance of the subsystem adequately. a broadened definition of a tpc is presented, supporting this approach. conservation implications: the concept of thresholds (initially ecological thresholds) has started influencing conservation management practice, a commonly-used formulation for management decision-making being the threshold of potential concern (tpc). practical tpc usage can often be improved by moving away from its initially pure ecological outlook, rather framing understanding through an interlinked socio-ecological view. introduction (back to top) strategic adaptive management as an operating philosophy (see roux & foxcroft 2011) has attracted wide interest in south africa. it spread from its original usage on river issues in and around the kruger national park (biggs & rogers 2003) to a wide range of other biophysical themes associated with national parks elsewhere in the country and more generally, to natural resource management issues such as integrated water resources management (freitag et al. in press). one central construct which has attracted interest is that of thresholds of potential concern (tpcs; biggs & rogers 2003). the current application of tpcs equates to ‘decision thresholds’ sensu martin et al. (2009), but ones which specifically presage predicted ecosystem thresholds. these authors also define such ‘ecological thresholds’, as well as another category, ‘utility thresholds’, which are value-based ones, giving high leverage if acted on. the strategic adaptive management approach in which tpcs are embedded assumes an often-changing context in which resource management is taking place. the approach thus strives to take a modest position, and to inculcate assertive learning-by-doing in what is seen as a complex world (sensu levin 1999) in which surprises occur. the original concept of tpcs (rogers & biggs 1999) hinged on upper and lower limits of a variable of interest in the ecosystem, being set with the idea of allowing management some time to respond, before the expected ecosystem threshold was crossed. this therefore carries the challenge of predicting or anticipating an ecological outcome via knowing, or at least being able to hypothesise, an approximate ecological threshold (martin et al. 2009). nevertheless, even if a state change is predicted, the approach does not always link the tpc to the hypothesised mechanisms of change explicitly (see ferreira et al. 2011) and may thus be less useful to management than if the cause of change is known. tpcs were originally seen as flowing from high-level vision and objectives, forming ‘low-level goals … provid[ing] managers on the ground with targets of ecosystem condition … scientifically rigorous, spatially and temporally bounded… act[ing] as amber lights to warn managers of possible unacceptable environmental change’; in practice being ‘upper and lower levels of change in selected biotic and abiotic variables which act as indicators of the acceptability of ecosystem condition’ (rogers & biggs 1999). it should be noted that this definition obviously does not preclude ongoing management, although imminent or actual tpc exceedance will always lead to consideration of an important additional management response. this usage has, apart from a few attempts to recommend it in other contexts (e.g. when wilderness qualities were perceived to be dropping below an acceptable level [south african national parks (sanparks) internal communication, freek venter] and for tourism standards and resource use guidelines, in a few postgraduate studies), been limited to biophysical issues, and mostly these have been viewed one at a time. this has led to some progress, some frustration and some learning (van wilgen & biggs in press; gaylard & ferreira 2011). the progress in certain theme areas, such as river management and fire management, has been held up as an example for possible use in other theme areas, according to biggs et al. (2011). however, these authors point out that there is too little deeper reflection generally associated with the use of tpcs, and that when such reflection does take place (as it did in the fire and river examples they describe), it enables a fuller and more productive adaptive cycle to occur. even though tpcs play this catalytic role in enabling these adaptive loops to be closed, there are several conceptual obstacles now evident after more than a decade of use of tpcs in the institutionalised way described by biggs and rogers (2003). ferreira et al. (2011) and gaylard and ferreira (2011) address several issues around emergent tpc usage, such as the fact that they are less successful if the underlying hypothesised mechanisms are not explicitly articulated. for instance, it hardly helps to have a tpc for unacceptably reduced river flow if one cannot understand what drivers (e.g. increased abstraction) are leading to this, or what worrying consequences (e.g. increased sedimentation) are likely. such explicit articulation is now undertaken in joint sciencemanagement discussions, which slows down the process, but results in the building of a more strongly shared rationality and thus carries wider organisational support than before. the joint hypothesising of such mechanisms, within the context of a socio-ecological system, also helps to link the objectives with the actual monitoring decisions more effectively, especially since it is not possible to monitor all the links and there is seldom one single ‘correct’ variable, or group of variables, to monitor in a complex system (stirzaker et al. 2010). also, many parks are, because of their need for rehabilitation, lying largely outside the desired state, sensu roux and foxcroft (2011). a wider formulation of the systemic context in which the tpcs are embedded (i.e. a richer and broader picture of the way the system appears to function) assists in helping understand the manner and pathway through which restoration might best be applied. designing tpcs to focus on processes and/or on rate changes, and not only on outcomes, would be make them even more useful, but this is very challenging, partly because of lack of detailed understanding of these processes. in this paper, we attempt to respond to several recurring conceptual issues that appear to act as dampers to the use of tpcs: • there is often uncertainty as to whether a real threshold even exists, and if so, where exactly it lies. this makes tpc developers and users hesitant, given the certainty they may originally have expected tpcs to deliver. hesitation also occurs when they suspect that a process is linear, or for any other reason, is believed not to have a clear threshold. • there is a feeling that even if a functional biophysical threshold exists (say a point after which so many large trees, ostensibly acting as nutrient pumps, disappear from a landscape, that its nutrient status quickly changes), it is perceived to be easier in some cases to set tpcs as human preferences (e.g. there are fairly clear preferences for large trees in open savannas [herbert prins, unpublished data]). similarly, scientists and managers in sanparks, although often suggesting that the threshold concept would be useful for pure economic or sociological (say tourist perceptional) phenomena, have in practice invariably not gone ahead and developed and implemented these in the way that has happened with biophysical ones. we speculate that this hesitation is partly because of perceived vagueness of these compared to the perception of ‘hard biophysical’ tpcs, even though preferences (such as for large trees in savannas) may often be expressed rather precisely. this hesitation to develop social tpcs may be related to the popular, yet flawed, perception that ecology is largely an exact science and/or to the uncertainty and associated lags resulting from the biophysical interacting with the social domain (such as delays in decision making or implementation). shortage of capacity to formulate these social and economic tpcs (as opposed to the more capacitated biophysical science division) has also been cited as a contributing reason for their absence in sanparks management plans. • there is uncertainty, if levels of tpcs are adjusted to give for example, slower responding management systems enough chance to still react in time, whether this constitutes a breach of their objectivity. in an attempt to find a basis from which to deal with the above obstacles, we will use the term ‘limit’ to refer to what is generally considered an unambiguous change, such as cessation of surface flow in a perennial river. we sometimes use ‘boundary’ (rather than threshold, since this paper deals with a potential widened meaning for tpcs) to refer to the transition at which a generally clearly acknowledged change of state occurs, such as from an open grassland to a thicket, or from a state-operated model to a concessioned model (for example, a tourism facility in a park). we recognise what we will call ‘ preferences’ along a gradient of continuous change, often not associated with a boundary or biophysical limit. we distinguish ‘preference’ from, or at least recognise that it grades into, ‘acceptability’ the latter implying something closer to non-negotiable, a feature some users originally may have implicitly attributed to tpcs, even though clear processes of tpc revision (rogers & biggs 1999) were described because it was anticipated they would be required often. we address these barriers explicitly and thus ask three questions, (1) how can we overcome the damper which arises because of uncertainty about whether a threshold exists and where it lies, (2) can tpcs be used for preferences and (3) can tpcs be adjusted to take embedded human behaviours, especially in the rate of institutional responses, into account? we then propose a tpc revamp that easily accommodates several biological and cultural concerns and perceptions inherent in socio-economic-ecological systems that are an intrinsic part of conservationists’ mandate as expected by society. dealing with the barriers (back to top) taking decisions and action in spite of uncertainty about a threshold or a limit managers cannot afford to be paralysed by uncertainties. in fact, adaptive management makes much of moving forward by conceptualising and probing the system intelligently and thus ‘reframing the problem in a way that gives a place to stand and take action while we learn more about how the system is really operating’ (stirzaker et al. 2010). rather than following the cue arising from the very common cautiousness of scientists (who often recommend waiting until more evidence is available before making a decision), generally adaptive management encourages us to move forward by making a decision, though this should be balanced with some slowness to enable appropriate stability and reflection (cilliers 2006). this decision-making imperative should include the possibility of an explicit, well-considered, ‘no-action-now’ management decision being taken. importantly, this differs from the conventional ‘wait-until-more-evidence-is-available’ approach, in that it and all other options and their anticipated consequences have been concurrently evaluated and the ‘ no-action-now’ decision taken as the best option. since such a decision is then as valid as any well-reasoned decision that drives immediate action, decision-makers should not feel they have not taken a decision simply because there has been no immediate resultant physical action on the ground. current biophysical tpcs can be seen as lying on a continuum of ‘empirically well or fairly well understood’ through an intermediate position ‘informed by expert opinion’ right down to ‘intelligent early guesswork’ (see also gaylard & ferreira 2011). this spectrum would likely also apply when tpcs are more widely used for social and economic variables. when a large proportion of tpcs are associated with intermediate and higher levels of inherent uncertainty, our observation is that this tends to disturb and even paralyse potential adaptive management practitioners. this caution is understandable, but also needs to be overcome if progress is to be made in adaptive management. this is particularly so because of the trade-off between encouraging and allowing systems to vary a sufficient amount within the ‘tent of acceptability’, that is, the envelope representing the desired state, to promote resilience, and the need to yet respond in time, so as not to land up on an undesired trajectory outside this envelope (see figure 1). the level of risk tolerance as opposed to risk aversion, (the so-called ‘risk appetite’), therefore plays an important role in how conservationists constrain that envelope of acceptable variability. the level at which they decide on their attitude to risk in different situations is thus pivotal. for instance, practitioners who regularly invoke the precautionary principle (cooney 2004) but in a traditional sense, run the risk of reacting too soon and attempting to constrain the ecosystem by not allowing it to vary and develop resilience. much of the critique of earlier management in the kruger park (in du toit et al. 2003) rests on the idea that managers had tried to hold the system relatively constant in time and space because of a belief that allowing great fluctuations was inappropriate. applied with differing sets of values in isolation, the precautionary principle can also provide fuel for seemingly well justified but opposing management recommendations, as documented in an australian example (http://www.acat.act.gov.au/decisions.php?action=decision&id=29). however, modern application of the precautionary principle (cooney 2004) suggests acknowledgement of full societal context including societal values guiding objectives, which can include a belief in the value of allowing ‘healthy’ amounts of variation in the ecosystem and concurrent use of adaptive management feedbacks. the application of tpcs is consistent with this newer approach, which we suggest needs to be even more cognisant of development of tools such as tpcs. tpcs together define the outline of a desired state (figure 1), derived through an adaptive planning process (see roux & foxcroft 2011) which defines a vision and hierarchy of objectives which explicitly capture and express societal values and expectations. in essence, thus, ecological tpcs are determined by human values and expectations, but guided by known or presumed biophysical limits. using tpcs, the full set of adaptive management steps are designed to then help take the most appropriate decision for implementation in this milieu of uncertainty, and to learn and change effectively as one progresses. figure 1: two-dimensional schematic of the ‘tent of acceptability’ or ‘desired state envelope’. we have thus argued that adaptive management methodology places a premium on making decisions at various levels of uncertainty, usually then eliciting probing actions which help develop understanding of complex systems, but that there is also sometimes place for a wellreasoned ‘no-action-now’ decision. modern interpretations of the precautionary principle support this complex systems view, and depending on context, can address greater levels of risk than previously. strategic adaptive management effectively incorporates value considerations in its fundamental planning phase, recognising that these, together with understanding of biophysical limits, underlie most tpcs in current use. arguments presented below on incorporation of social preferences constitute important further reasons why decision-makers should not be paralysed by uncertainty concerning actual limits. can thresholds of potential concern be used to describe, or can they include, societal preferences? we could argue that this is a non-question, given that the biophysical tpcs, although influenced by absolute limits, are also a form of societal preference. however, the question is so often asked in this form (and the expression to date of tpcs, has been in terms of biophysical variables) that we feel that it has been useful to retain it as the obvious entry point for those asking it. a better formulation for future use might be ‘can social or biophysical descriptors qualify equally validly for societal preferences as embodied in tpcs?’ strategic adaptive management is rooted in human values as a basis for deciding on the desired state (roux & foxcroft 2011) which is described in joint social and ecological terms (walker & salt 2006). one component is a ‘healthy’ biophysical system, where ‘healthy’ is deemed in accordance with these values. a commonly depicted intersection of social (including economic) and biophysical facets (figure 2) suggests that the desired state can most productively be seen as the socially and biophysically desirable and sustainable overlapping area. again, ‘biophysically sustainable’ refers to absolute physical limits, but also to considerations of human acceptability and even preference. so again, in spite of the fact that both circles involve human choice, we find the figure useful as an entry point, the target of the biophysical one being expressed in biophysical terms. the overlap in figure 2 requires the context, if not the definition, of a tpc to be expanded beyond only addressing the biophysical subsystem. it can still be useful to have definitions for tpcs in pure biophysical or social subsystems, but these must then be viewed as parts of the joint system. figure 2: desirable and sustainable biophysical and social domains, and the overall ‘desired state’ in their overlap zone. a common debate in sanparks is about the introduction of the ‘big 5’ (elephant, rhino, buffalo, lion and leopard) into smaller, usually recently proclaimed, parks. this is usually justified on the basis of both restoring natural biota and processes, and presumed attractiveness to tourists. statements which can be quoted as supporting this are often available in the vision and objectives of the specific park’s management plan, which will have been drawn up, in line with the national environmental management: protected areas act (act no. 57 of 2003), in a stakeholder-centred way. contention that has arisen around such introductions (for instance, damage to adjacent livestock from escaped predators) is in several cases not well resolved, and flags the need for more holistic and nuanced guidance in the higher-level processes (such as derivation or prioritisation of the park mission and objectives) which provide guidance for such decisions. without going into how this is being addressed by sanparks, and without specifying any particular park, we create here a plausible but fairly generic ‘thought experiment’ in which one set of possible outcomes of such decisions, as they might currently be made, could play out when viewed as an interacting socio-ecological system. the purpose of doing this is to illustrate that social and ecological tpcs interact, and are usefully viewed as a joint system, actually necessitating the incorporation of social tpcs. we use the kind of cause-effect diagram that might be produced by a park science-management forum at sanparks, without adhering to the full rigour and strict conventions of a formal influence diagram. the particular cause-effect diagram in figure 3 has three vertical threads, depicting outcomes for tourism, ecological condition and neighbour relations. the heart of the diagrammatic narrative shows a positive or reinforcing feedback loop between increasing possibilities for elephant viewing in the park, and tourism-derived income. in this particular story, only those negative (dampening) feedbacks which later on influence tourism income in some way (resultant problems with trail camp accommodation, unacceptable tourist-elephant conflict and ultimately, loss of viewing opportunities due to a mass decline of elephant following a severe drought) manage to start slowing down the major reinforcing loop which initially governs park decision-making in this hypothetical narrative. potential feedbacks about other concerns covered in sanparks’ mandate (habitat condition, access to tourism by lower/middle income locals, and costs borne by neighbouring communities) are, in this particular scenario, all insufficiently strong to reverse that central trend. obviously we could continue by exploring what might happen if those currently ‘dead-end’ options became stronger (producing their own effective feedbacks), and develop by similar thought experimentation further illustrative narratives with their own cause-effect diagrams. in fact, the elephant ecological impact tpc, which is treated, for reasons of clarity of the overall story, as a single item in the figure, unpacks in practice into a whole diagram of its own, currently in use in various forms in sanparks. the point is that whatever diagrams are deemed necessary by a science-management forum (the normal vehicle to discuss such issues in the sanparks case) should be drawn. figure 3: an illustrative cause-and-effect diagram showing how inter-related social and biophysical themes are a typical issue in park management. figure 3 should be adequate to illustrate why such an approach is useful, if not essential. it provides the backdrop of systemic understanding for the justification and positioning of tpcs. that systemic understanding, represented by the diagram as a group mental model, is thus jointly agreed on, at least by the participants, bearing in mind that they also bring with them wider stakeholder impressions from frequent interactions with these groups, some of these interactions being mandatory for park plans. figure 3 also hypothetically lists the most likely tpcs which might be used, and illustrates why both social and biophysical tpcs are essential if a holistic understanding is to be obtained. it also lists a host of other potential tpcs which could be used in a world with limitless capacity. there is an art to selecting a few particular ones, from all the possible ones which could be thought of, enough to give a fair chance of an overall understanding of the system, but not so many that the associated development, monitoring and interpretation activities become overwhelming. we need to remember that a diagram of the kind shown in figure 3 would constitute only one of several, and possibly in a large or complex park, up to perhaps 15 cases. each diagram will have associated tpcs. even given that several of the same tpcs occur in different diagrams (in other words, in different theme areas), the overall capacity to develop tpcs, monitor, manage and reflect, remains finite, and a parsimonious but effective set needs to be chosen. while there is some evidence that ecosystem changes might often (but certainly not always) take the form of thresholds (schröder et al. 2005), these are invariably connected to social changes (resilience alliance & santa fe institute 2004) which are underlaid by values, the results of these value shifts often leading to fairly abrupt change (as in a ban on smoking). we thus need to become equally comfortable with the fact that many of these social (including economic) changes can also have a threshold nature – such as changes in fashion, and financial collapses (scheffer 2009). however, the issue of linearity compared to nonlinearity has probably caused unnecessary confusion in the implementation of tpcs to date. in any system with feedbacks causing movement across a boundary, even if some of the changes are linear, there could be a state change of interest to natural resource or park managers. if there is no state change, or if the change is linear, predictable, and especially if it is reversible, then this (unusual) set of circumstances will be relatively straightforward for management to handle. our interest therefore focuses on the more frequently occurring (often non-linear) tenacious state changes in which one threshold (or more often, a series of interlinked thresholds in a coupled sequence as in figure 3) can change. this is loosely called a regime shift (scheffer 2009): a relatively sudden jump from one dynamic regime to another, in which the interactions between the controlling variables (and therefore the explanatory diagram) usually alter. certain of these regimes are desirable as judged by operative values, and others undesirable. when in a desirable one, management aims to maintain its ‘healthy’ state by allowing variation within it, but not allowing an undesirable trajectory to take the system outside of the limits. one important sign of the system moving outside of the desired state is a consistent directionality until the next state is reached (ferreira et al. 2011). said another way, allowing this desired variation (inside the envelope, figure 1) usually implies that net overall movement hovers around zero (i.e. in the ‘ball-in-the-basin’ metaphor, walker & salt 2006). conversely, management initiatives to move the system from an undesired state to get back within it, also implies directionality to achieve this (figure 1). this was often referred to as ‘back inside’ or ‘back to natural’ in conservation circles and is formally restoration (society for ecological restoration 2004), although recently there is increasing recognition of the need to sometimes create acceptable ‘novel ecosystems’ (http://herenthereneverywhere.wordpress.com/2010/02/04/novel-ecosystems/). human needs and values are often central underlying drivers of these changes, with the biophysical changes, such as global temperature increase, often being inadvertent outcomes which then lead to many of the changes. the adaptive management framework (see roux & foxcroft 2011) should thus clarify how exactly tpcs might work as both ‘retainers’ of a particular state and ‘ targets’ to which to return (freitag et al. in press). whether a boundary crossing is threshold or non-threshold in underlying nature, the tpcs should be reflective of the desired state and its inherent values. although sanparks has made good progress (see roux & foxcroft 2011) in making this connection, through use of adaptive planning, it is still grappling with this when underlying objectives requiring to be balanced, are only partly compatible, as in the example in figure 3. in fact, the situation in the generic example in figure 3 (and alternative scenarios which might arise if the ‘dead-ends’ become more influential) will in the opinion of the authors be highly influenced by determination of effective synergies and trade-offs at the level of organisational and societal values. until this happens effectively, and even afterwards, we consider such cause-effect diagrams indispensible in the formulation of tpcs and in the management of such a situation. often an emergent slow variable, say poorer education over many years, can lead to gradually poorer service delivery. in natural resource management, this may manifest as a sudden shortfall in a crisis period, for example, during a cyclone or a drought. an important challenge resides in setting thresholds which link the focused sub-system to the slower variables operating on the overall socio-ecological system. this is important because slow variables may be difficult or impossible to measure, yet may irrevocably change the system. slow variables may also be biophysical, sedimentation of rivers being a good example in rivers flowing through the kruger national park. sedimentation is relatively difficult to measure, and its consequences could easily have taken kruger park management by surprise were it not for an intensive research programme which highlighted it by drawing systemic diagrams of the type we now use and recommend in this paper, and were it not for the linkage made between sedimentation and some faster-acting, more easily measured variables. in figure 3, investigation may show that profitability tpcs could be used as an early warning system to detect if, and perhaps at what scales, slower variables (slow changes in ecological condition which lead, over decades, to a sudden major change) are changing in an undesirable direction. the well-known history of elephant culling in kruger national park (whyte et al. 1998; van aarde et al. 1999) serves as a further example of the interactions between biophysical aspects and societal preferences. the history details, from the date of its inception until it was stopped, what was perceived as a mechanistic effort at capping populations. this was to avoid passing a perceived biophysical threshold of ecological degradation (van wyk & fairall 1969), derived through use of the precautionary principle within the rangeland ecological paradigms in use at that time. this capping of populations had other unexpected biophysical effects, such as higher calving rates during the years of culling (van aarde et al. 2008). the overall system had in fact changed in a step fashion (sensu scheffer 2009) as a consequence of the intervention, even if protagonists of culling felt that such a change was acceptable given what they were attempting to avoid. in broader perspective, the culling campaign led to an increase in social unacceptability over time, a de facto threshold, and to suspension of culling (whyte et al. 1998) with consequent further elephant population growth (young et al. 2009). the socio-ecological system had passed a boundary, having moved from a state of ‘culling by technocrats, acceptable to society’ to a state of ‘no longer necessarily acceptable, suspend culling and rethink overall issue’. in retrospect, at park-level, the elephant impasse resulted in decision paralysis while parks authorities were required to collect more information and conclusive proof of habitat destruction. while this caused significant frustration among managers, the resultant ‘slowness’ (cilliers 2006) has enabled a rethink of ecological paradigms (gaylard & ferreira 2011). nevertheless, now, 16 years after culling stopped in 1994, the originally predicted ecological thresholds have not been passed, except for concern around the tpc concerning reduction of large trees that has been tabled (sanparks 2009) and may lead to management aimed at reducing effects, possibly even culling (now legally allowable again; deat 2008), in at least some parts of the kruger national park. we have stated that while a broader view is essential for contextualisation, it is not feasible to track the full suite of variables and links, and some focus on a key part is needed in the interests of practical expediency. gaylard and ferreira’s (2011) unpacking of ecological concerns extended into the social realm, coupled with scenario evaluation and comparative risk assessment (see box in gaylard & ferreira 2011), may actually help overcome parts of this challenge. an example is how conservationists may wish to manage elephant effects that range from ecological to societal (scholes & mennell 2008). unpacking these effects could help converge onto a common mechanism and driver of how elephants can cause ecological change, induce human conflict, enhance tourism experience and generate revenue (gaylard & ferreira 2011) – in their analysis, spatial use by elephants turns out to be central. tpcs as originally formulated, akin to decision thresholds (martin et al. 2009), will thus reflect modulators of elephant spatial use, while allowing the inclusion of utility-based tpcs (martin et al. 2009) and will embody a range of ecological, social, economic and financial contexts. although it is useful to link these to actual ecological thresholds, we have learnt that restricting tpc usage to only this latter class is very limiting and has proved frustrating. extending their usage to include societal preferences and take into account organisational and human behaviour in the ways described, and unpacking the real meaning of each, better supports how society influences how conservationists should act in their interest. the narrative in figure 3, and the potential other narratives that could unfold if the currently ‘dead-end arguments’ in figure 3 were to gain sufficient momentum, demonstrate that even innocent-looking social choices can have very real consequences for the desired biophysical state in a conservation agenda. the viewpoint that humans can choose to carry out any of a range of activities on an ecosystem template that is generally in good shape, is correct. however, zooming in slightly, there are different possibilities in the specific mix of ecosystem services available, and these are of explicit management interest. for instance, predictions that the mopani vegetation type will expand under climate change (rutherford et al. 1999) and perhaps later embrace all except the southernmost corner of kruger national park, will almost certainly lead to different tourist outcomes and experiences. we could argue that both mopani and what is left of the other habitats are fundamentally healthy, and similar social activities can be practised on both. but the way human values and park management might intersect with this projected outcome, leads to varying social trajectories (for instance, game-viewing tourism is popular along rivers in the mopani areas, but far less so away from rivers) and potential thought experiments which, like those in figure 3, will illustrate how coupled and interdependent the subsystems are, and help avoid or navigate such change. it is clear that much hinges on what is considered acceptable or preferred, or unacceptable and not preferred. this is broadly circumscribed during the adaptive planning process by the desired state (roux & foxcroft 2011) which should have more appropriately been called the ‘desired set of varying conditions’ to reinforce the idea that systems tend to change all the time. what articulated desired states turn out to look like, and what is acceptable, are both fundamentally influenced by a variety of values, some of which are explicit and/or shared between stakeholder groupings (see sanparks [2008] for a set of explicit sanparks conservation values). it is well known that where there is a relatively high degree of sharing of values, societal preferences can more easily be defined and articulated, as for tpcs. we thus feel that social tpcs should not only be allowed and encouraged, but that they are in fact essential (along with biophysical tpcs) if a full socio-ecological understanding is to be available as a basis for decision-making. can thresholds of potential concern be adjusted to take embedded human behaviour, especially differences in the rate of institutional responses, into account? while remote ecosystems, almost uninfluenced by human events, can demonstrate naturally generated threshold events, almost all natural resource management situations in south africa have fundamentally important social drivers and social outcomes. in virtually every case, various legislative and management authority mandates may also require contrasting outcomes, and a somewhat different culture and behaviour characterises each such authority. for instance, conservation and animal health legislation in south africa may not have common outcomes (in the context of elephant management, see scholes & mennell 2008). even small differences in the biophysical situation can often reflect in some way the range of possibilities or outcomes in the social domain and vice versa. resource management is typified by such partly competing, partly overlapping goals, and as holistic as possible an understanding is required to deal with this. tpcs developed in such a joint situation will need to reflect both cultures, or increasingly, a newly forged joint understanding and behaviour. tpcs were originally defined as being some time before a ‘real threshold’ and so the question arises whether it would then be reasonable to set them more conservatively (i.e. to be exceeded even earlier) in a context where management responses might, for whatever reason, take longer. for instance, slower anticipated institutional responses in for example, one country compared to another. a rational empiricist might say this detracts from the objective setting of the tpc, because it takes into account the shortcomings of institutions and people, even though most will agree it is easier to change before, rather than after, a threshold is exceeded. for instance, degraded rivers are very difficult to restore; it is easier to keep a healthy river in a good state, irrespective of the institutional capacity that this requires in one catchment or country compared to another. however, making such an adjustment for capacity would seem to be consistent with, and even in support of, the logic describing the ‘buffer ’ timing that should be considered in refining tpcs, according to scholes and kruger (2011). towards a broader definition (back to top) a re-orientation of the kind recommended in this paper may necessitate a revised definition of a tpc. we suggest ‘any practical decision prompt based on a particular level (or upper and lower levels) of a social or biophysical variable which assists managers to avoid (or to transition into) a different state which is considered undesirable (or desirable). tpcs should not be seen in isolation, but as part of dealing with a complex socio-ecological system, and should be informed by biophysical limits in conjunction with human values’. conclusion (back to top) all three barriers described can be tackled by viewing the system in a wider context. tpcs are not isolated entities. they should relate to the broader subsystems, which often have social and biophysical interactions, in which they are embedded, and reflect the values and objectives comprising the desired state. the wider view is usually represented by a cause-effect diagram akin to the thought experiments presented here, and similar to what gaylard and ferreira (2011) use to help enhance tpc implementation. if it does not do so already, this should include the social domain, often manifest as preferences or as net economic effects which can be stated as acceptable thresholds. that will frame tpc usage explicitly in a coupled socio-economic-ecological context, where change in a single driver, or often a set of related cascading thresholds, leads to crossing of one or various overlapping boundaries, and thus to a regime shift (scheffer 2009). until recently, most usage of tpcs has been geared towards a setting at least slightly before perceived ecological thresholds. practitioners should be as plain as their knowledge allows them to be, about whether any of these are linked to thresholds or boundaries. if so, it seems entirely reasonable that the decision on how far before, might depend on the likely speed of response mechanisms, guided by principles described in scholes and kruger (2011) and that clearer management responses are enabled if the mechanisms causing change are explicitly hypothesised (gaylard & ferreira 2011). nevertheless, if it appears that the tpc is purely a preference, this should be stated in the supporting documentation, and where the link between a particular preference and any supposed real biophysical threshold is unclear, this should be noted and commented on, to assist future evolving development of the management system. whilst we encourage wider socio-ecological contextualisation, it seems sensible when it comes to monitoring to rather choose key focus areas and track just those segments of the wider understanding. if objectives or drivers change, or if the variables followed are not adequate for the task, it may be necessary to expand or adjust these, probably within the framing of an adjusted cause-effect diagram. biophysical and social sciences are indispensable in helping understand the whole picture required for tpc formulation, and practitioners cannot be expected to understand the management meaning clearly, without some understanding of the pervasive inter-linkages between these. such joint thinking should be encouraged right down to the operationally detailed components of park management plans. the commonality of joint mind-maps of socio-economic-ecological systems thus stretches across hierarchies of governance locally, provincially, nationally and internationally. the setting of thresholds is a means of checking to see if the originally jointly defined desired state (with its various objectives and goals) is maintained. this may include a variety of both qualitative (generally at entry-level and then followed by increasingly building the rigour in the understanding) and quantitative tpcs (see also gaylard & ferreira 2011). tpcs can only be practical if constructed, evaluated, adjusted and reacted upon within the context of human preference and behaviour. if the changes suggested in this paper could be combined with refinements through approaches that capture a generality of tpc definitions as being process-orientated and recognising variability as a key outcome (ferreira et al. 2011), and approaches that unpack key management concerns into cause-and-effect diagrams (gaylard & ferreira 2011), then we may be entering the second and, it is hoped, more fruitful generation of tpc usage, and not restricted to only a moderate number of ‘special’ cases which appear to work well. a good opportunity therefore exists to write a unified set of guidelines collating these changes in approach and practical use. acknowledgements (back to top) we thank the many scientific collaborators, managers and rangers within sanparks for fruitful discussions over the years. these discussions have prompted ideas about conceptual and practical problems concerning tpcs and have led to the writing of this paper. thanks also to oonsie biggs for insightful discussions and comments on an earlier draft of this paper, as well as to two anonymous reviewers whose comments and inputs have led to a more focused and tightened manuscript. references (back to top) biggs, h.c., breen, c., slotow, r., freitag-ronaldson, s. & hockings, m., 2011, ‘how assessment and reflection relate to more effective learning in adaptive management’, koedoe 53(2), art. #1001, 13 pages. doi:10.4102/koedoe.v53i2.1001 biggs, h.c. & rogers, k.m., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59−80, island press, washington. cilliers, f.p., 2006, ‘on the importance of a certain slowness. stability, memory and hysteresis in complex systems’, emergence: complexity and organisation 8(3), 106–113. cooney, r., 2004, the precautionary principle in biodiversity conservation and natural resource management: an issues paper for policy-makers, researchers and practitioners iucn, gland, switzerland and cambridge, uk. deat, 2008, national environmental management: biodiversity act, 2004 (act 10 of 2004): national norms and standards for the management of elephants in south africa, department of environment affairs & tourism, pretoria, south africa. du toit, j.t., rogers, k.h. & biggs, h.c. (eds.), 2003, the kruger experience: ecology and management of savanna heterogeneity, island press, washington. ferreira, s.m., deacon, a., sithole, h., bezuidenhout, h., daemane, m. & herbst, m., 2011, ‘from numbers to ecosystems and biodiversity – a mechanism approach to monitoring’, koedoe 53(2), art. 998, 12 pages. doi:10.4102/koedoe.v53i2.998 freitag, s., biggs, h. & breen, c.m., (in press), ‘fifteen years of the spread and maturation of adaptive management in south african national parks: organisational learning in systems perspective’, in w. freimund, s. mccool & c.m. breen (eds.), engaging complexity in protected area management: challenging occam’s razor, university of kwazulu-natal press, pietermaritzburg, south africa. gaylard, a. & ferreira, s.m., 2011, ‘modification and maturation of sanpark’s adaptive planning process making critical linkages between conservation objectives and actions’, koedoe 53(2), art. 1005, 8 pages. doi:10.4102/koedoe.v53i2.1005 levin, s., 1999, fragile dominion: complexity and the commons, helix books, boston. martin, j., runge, m.c., nichols, j.d., lubow, b.c. & kendall, w.l., 2009, ‘structured decision making as a conceptual framework to identify thresholds for conservation and management’, ecological applications 19(5), 1079–1090. doi:10.1890/08-0255.1, pmid:19688917 resilience alliance & santa fe institute, 2004, thresholds and alternate states in ecological and social-ecological systems, ‘resilience alliance (online)’, viewed 15 april 2009, from http://www.resalliance.org/index.php?id=183 rogers, k. & biggs, h.c., 1999, ‘integrating indicators, endpoints and value systems in strategic management of the kruger national park’, freshwater biology 41, 439−451. doi:10.1046/j.1365-2427.1999.00441.x roux, d. & foxcroft, l.c., 2011, ‘the development and application of strategic adaptive management (sam) within south african national parks’, koedoe 53(2), art. 1049, 5 pages. doi:10.4102/koedoe.v53i2.1049 rutherford, m.c., midgley, g.f., bond, w.j., powrie, l.w., robert, r. & allsopp, j., 1999, south african country study of climate change. plant biodiversity: vulnerability and adaptation assessment, pp. 1–59, national botanical institute climate change group, cape town. sanparks, 2008, policy context: sanparks’ mandate and values, chapter 2 of ‘a framework for developing and implementing management plans for south african national parks’ viewed 15 april 2009, from http://www.sanparks.org/parks/kruger/conservation/scientific/key_issues/plans/adaptive/pdfs/chapter_02.pdf sanparks, 2009, minutes of the conservation services management committee meeting, south african national parks, kruger national park archives, skukuza on 02 april, 2009 scheffer, m., 2009, critical transitions in nature and society, princeton university press, princeton. scholes, r.j. & kruger, j.m., 2011, ‘a framework for deriving and triggering thresholds for management intervention in uncertain, varying and time-lagged systems’, koedoe 53(2), art. #987, 8 pages. doi:10.4102/koedoe.v53i2.987 scholes, r.j. & mennell, k.g. (eds.), 2008, elephant management: a scientific assessment for south africa, wits university press, johannesburg, south africa. schröder, a., persson, l. & de roos, a.m., 2005, ‘direct experimental evidence for alternative stable states: a review ’, oikos 110, 3–19. doi:10.1111/j.0030-1299.2005.13962.x society for ecological restoration international science and policy working group, 2004, the ser international primer on ecological restoration. version 2. stirzaker, r., biggs, h., roux, d. & cilliers, p., 2010, ‘requisite simplicities to help negotiate complex problems’, ambio 39, 600–607. doi:10.1007/s13280-010-0075-7, pmid:21141779 walker, b. & salt, d., 2006, resilience thinking: sustaining ecosystems and people in a changing world, island press, covelo. van aarde, r.j., ferreira, s.m., jackson, t., page, b., de beer, y., gough, k., et al., 2008, ‘elephant population biology and ecology’, in r.j. scholes & k.g. mennell (eds.), elephant management: a scientific assessment for south africa, pp. 84−145, wits university press, johannesburg, south africa. doi:10.1111/j.1469-1795.1999.tb00075.x van aarde, r.j., whyte, i.j. & pimm, s.l., 1999, ‘culling and the dynamics of the kruger national park elephant population ’, animal conservation 2, 287−294. van wilgen, b.w. & biggs, h.c., (in press), ‘a critical assessment of adaptive ecosystem management in a large savanna protected area in south africa’, biological conservation. doi:10.1016/j.biocon.2010.05.006 van wyk, p. & fairall, n., 1969, ‘the influence of the african elephant on the vegetation of the kruger national park’, koedoe 12, 66−75. whyte, i.j., van aarde, r.j. & pimm, s.l., 1998, ‘managing the elephants of kruger national park’, animal conservation 1, 77−83. doi:10.1111/j.1469-1795.1998.tb00014.x young, k.d., ferreira, s.m. & van aarde, r.j., 2009, ‘the influence of increasing population size and vegetation productivity on elephant distribution in the kruger national park’, austral ecology 34, 329−342. doi:10.1111/j.1442-9993.2009.01934.x article information authors: ben j. strohbach1 josephat t. kutuahuripa2 affiliations: 1school of natural resources and spatial sciences, polytechnic of namibia, namibia 2plant production research, ministry of agriculture, water and forestry, namibia correspondence to: ben strohbach email: bstrohbach @polytechnic.edu.na postal address: private bag 13388, windhoek, namibia dates: received: 30 oct. 2012 accepted: 26 aug. 2014 published: 28 nov. 2014 how to cite this article: strohbach, b.j. & kutuahuripa, j.t., 2014, ‘vegetation of the eastern communal conservancies in namibia: ii. environmental drivers’, koedoe 56(1), art. #1117, 12 pages. http://dx.doi.org/10.4102/koedoe.v56i1.1117 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/koedoe.v56i1.1117-1. copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. vegetation of the eastern communal conservancies in namibia: ii. environmental drivers in this original research... open access • abstract • introduction • research method and design    • study area    • procedure       • available soil moisture       • climatic data       • environmental gradients • results • discussion    • climatic gradients versus changes in soil types    • plant adaptations to environmental drivers    • shortcomings in data analysis • conclusion • acknowledgements    • competing interests    • authors’ contributions • references • footnotes abstract top ↑ the eastern communal conservancies are situated along the western fringe of the kalahari basin. under a very short rainfall gradient, the vegetation abruptly changes from microphyllous acacia-dominated savannas to mesophyll savannas, dominated by terminalia sericea and combretum spp. we hypothesise that this is caused by changes in soil moisture availability brought about by changes in soil texture from loamy soils to deep sands (the ‘inverse texture effect’). for this analysis, we used vegetation and soils data derived from a recognisance survey of the natural resources of the study area. as the sites in the soil and vegetation surveys did not overlap, it was decided to use only synoptic data for the plant associations in the analysis. non-metric multidimesional scaling ordination was utilised as ordination technique of the vegetation data and various environmental parameters, including soil texture, soil hydraulic parameters, climatic and fire regime parameters, were overlaid as biplots onto the resulting graph, as were various plant functional attributes particularly related to climatic conditions. the main environmental gradient identified within the study area is the rainfall gradient. this relatively short gradient, however, does not explain the marked change in vegetation observed within the study area. this change is attributed to the change in soil type, in particular, the soil texture and the associated soil hydraulic parameters of the soil. this gradient is closely correlated to leaf size, explaining the change from microphyll savannas to mesophyll savannas along the change from loamy to sandy soils. one of the lesser understood mechanisms for the survival of these mesophyll plants on sandy soils seems to be a deep root system, which is actively involved in water redistribution within the soil profile – by hydraulic lift, inverse hydraulic lift and stem flow. conservation implications: understanding these mechanisms will greatly assist in understanding savanna dynamics. with the threat of global climate change, we postulate that the vegetation will gradually change from the present mesophyll to a microphyll savanna, but that the grass sward will probably not develop very well. shrub and tree removal (‘bush harvesting’) is likely to speed up the desertification process within this area. introduction top ↑ in the first article of this series (strohbach 2014), the vegetation of eight communal conservancies, as well as two farming areas in east-central namibia, are described. these conservancies are situated within the camelthorn savanna in the south and the tree savanna and woodlands described by giess (1998) in the north. here, under similar climatic conditions, the vegetation abruptly changes from a microphyll, acacia-dominated savanna to a mesophyll savanna dominated by terminalia sericea and various combretum species, as the soils change from relative fine-grained soils of the central plateau to deep, coarse sands of the kalahari basin (leser 1972). on an aridity gradient stretching over roughly 1100 km from north-northwest to south-southwest within the kalahari desert of botswana (on uniformly sandy soils), macrophyllous trees are gradually replaced by microphyllous trees as the rainfall decreases and the aridity increases (skarpe 1996). this rainfall gradient is estimated to range from 600 mm in the north-northeast to less than 250 mm in the south-southwest. on a similar aridity gradient from the lesotho highlands to the west coast of southern africa (with a rainfall gradient between 522 mm and 41 mm per annum), macrophyll plants were replaced by microphyll plants within a riverine forest as aridity increased (werger & ellenbroek 1978). in the same study, an increasing number of plants with sclerophyllous leaves (as adaptations to arid conditions) were found in the more arid environments. more abrupt changes in the composition of moist savannas of central mozambique (ranging from grasslands to savannas, to woodlands, forests and swamp forests) have, in detail, been demonstrated by tinley (1982) to be related to differences in soil profiles, especially with regard to soil texture, soil depth and the presence and depth of constricting layers within the profile, despite a uniform climate. however, little is known on the causes for the abrupt change from microphyll to macrophyll savannas in the arid western kalahari basin region. water availability is the main driving force of vegetation in arid environments (noy-meir 1973; tinley 1982; walter & breckle 1984). tree roots are known to utilise an expansive soil volume to source nutrients and water. record depths of 60 m and 68 m have been documented for acacia erioloba and boscia albitrunca, respectively, in the kalahari (canadell et al. 1996). pterocarpus angolensis tree roots reach a depth of between 15 m and 20 m (mwitwa, munthali & van wyk 2008), whilst baphia massaiensis subsp. obovata has been reported to have a rooting depth of about 2.5 m – 2.7 m (savory 1963). terminalia sericea, one of the most abundant species in the study area, is known to have a shallow root system less than 1 m deep under arid conditions (350 mm mean annual rainfall) (hipondoka & versfeld 2006; hipondoka et al. 2003). under more mesic conditions (450 mm mean annual rainfall), however, this species tends to retain its tap root, reaching beyond the explored depth of 1.2 m. similar findings are known also from the banksia woodlands in australia, where some deep-rooted species obtain their water from a depth of up to 9 m. other evidence from australian eucalyptus savannas suggest that deep-rooting species are better capable of surviving drought conditions than shallow-rooting species (fensham & fairfax 2009). it is generally assumed that fine-textured soils have greater porosity, and thus a greater water-holding capacity, than coarse-textured (sandy) soils (brady & weil 2002). finer soils have a larger pore volume, albeit smaller pores, allowing more water to be stored in the profile. added to this is the fact that water flows faster through larger pores – a sandy soil loses water to deep soil layers more rapidly than a loamy or clay soil. in arid and semi-arid areas, however, the inverse texture effect is observed – vegetation is lusher and better developed, often with a higher net primary productivity, on sandy soils than on finer clay or silt soils (austin et al. 2004; noy-meir 1973). this is a result of the fact that soil water is held by adhesion to the fine-grained particles and is, in effect, not available to plants. in sandy soils, water infiltrates to deeper soil layers during the rainy season, out of reach of fast-growing, shallow-rooted plants such as grasses. during the dry season, this deep soil moisture reservoir becomes available to deep-rooted plants (brady & weil 2002; noy-meir 1973; walter & breckle 1984). an in-depth analysis of this phenomenon has been provided by laio, porporato, fernandez-illescas and rodriguez-iturbe (2001), laio, porporato, ridolfi and rodriguez-iturbe (2001), porporato et al. (2001) and rodriguez-iturbe et al. (2001). with this article, we want to illustrate the effects of environmental factors influencing the formation and distribution of various vegetation associations found within the study area. in particular, we hypothesise that the ‘sudden’ or marked change from microphyll acacia-dominated savannas to mesophyll savannas (terminalia sericea, combretum spp.) in the eastern communal areas is the result of a combination of increased rainfall and especially increased water availability in the deep, sandy soils of the kalahari. research method and design top ↑ study area the omaheke [‘great sand desert’] forms the western fringe of the kalahari basin (mendelsohn & el obeid 2002, 2003; mendelsohn et al. 2002) (figure 1). the central, major part of the landscape is formed by an extensive sand plateau, sloping from the southern, western and north-western fringe towards the central east. the topography of the kalahari sand plains is flat to almost flat. these kalahari sands, with weak to little development in the profiles, are classified as haplic or ferralic arenosols.1 these soils are generally deep to very deep and well drained. they have a low agricultural potential as they are leached, have a low water-holding capacity and a low nutrient status (coetzee 2003; mendelsohn et al. 2002; strohbach et al. 2004). figure 1: the study area (indicated in bold lines along the east-central border of namibia) in relation to the kalahari sand basin in southern africa (grey shading). within this sandy plain, occasional dune fields occur (de pauw et al. 1998; thomas 1984). the sand dunes reach a height of up to 20 m, forming an undulating to rolling landscape. the dune crest, dune slopes and foot of the dunes consist of ferralic arenosols with haplic arenosols mostly on the lower footslopes of the dunes. soils on the interdunal depressions and isolated small pans of the kalahari vary in depth from moderately deep to very deep sandy soils. the two major soil types that occur within this soil-mapping unit are petric calcisols (table 1) and haplic arenosols. isolated pans have shallow haplic calcisols with calcrete horizons surfacing and some gleying was occasionally found (strohbach et al. 2004) (figure 2). figure 2: dominant landscape units within the study area, indicating also the dominant soils for these landscape units (compare with table 1). table 1: vegetation associations and their habitats from the study area, as extracted from global information system layers. the sand plateau is drained by a number of deeply incised drainage channels, commonly referred to as omiramba (singular: omuramba) (king 1963; strohbach 2008). these fossil drainage lines were formed during the wetter phase predominating the early to mid-pleistocene age. the depths and widths of the omiramba depend on the prevailing winds and water flows experienced throughout the ages. very often, calcrete outcrops are exposed at the edges of the major omiramba, especially the otjozondjou, epukiro, eiseb and rietfontein omiramba. the major soil types of the omiramba are arenic or calcic fluvisols, occasionally associated with petric or haplic calcisols (strohbach et al. 2004) (table 1) (figure 2). along the south-western fringes of the kalahari sand basin, but also south and east of gam, the sand cover is shallow, often overlaying calcretes, with occasional calcrete pans and reworked whitish, loose, unstructured kalahari sands. the topography is flat to almost flat. typical here are haplic calcisols, whilst in the far north-east some eutric cambisols (table 1) have also been found (strohbach et al. 2004) (figure 2). floodplains occur between the klein omatako omuramba and the omuramba omatako and stretch up to okanguindi village, south-east of okakarara. they are subjected to repeated seasonal flooding as a result of the relatively low relief. the continuous erosion of the surface horizon by water, results in the formation of many pans and watercourses throughout the area. owing to continuous reshaping and removal of the parent material and topsoil, the soils of this mapping unit are much younger in terms of development and age. the dominant soils of this mapping unit, arenic fluvisols (table 1), are weakly developed and poorly structured (figure 2). soils within the lower positions are subjected to flooding and show signs of mottling caused by poor drainage and water-logging. in the far north-west of the study area (north-west of otjituuo towards grootfontein), but also north of gam, is a plateau with limestone related to the karstveld. hard and soft calcrete and/or limestone outcrops are exposed at the soil surface. the dominant soils are lithic and molic leptosols, with some skeletic leptosols being associated (strohbach et al. 2004) (figure 2). the climate is relatively uniform, with a rainfall gradient ranging, on average, from just under 400 mm in the south to just under 500 mm in the north, annually (figure 1). the variation in rainfall (cv) is about 60% for the entire study area (botha 1996; mendelsohn et al. 2002; namibian agricultural resources information system [naris] 2001). this typical summer rainfall area has a dry winter and a dry and hot spring season (figure 3). frost is not unheard of in winter, with between 5 days of frost in the north and 30 days or more in the south (mendelsohn et al. 2002; southern african science service centre for climate change and adaptive land use [sasscal] weathernet 2012) (figure 3a–3c). the average relative humidity drops to below 10% during the dry months, but rises to 70% – 80% during the rainfall months (mendelsohn et al. 2002). figure 3: climate diagrams for the stations, (a) tsumkwe and (b) sonop (both to the north of the study area) and (c) sandveld to the south of the study area. regular veld fires are known to occur within the study area (le roux 2011; mendelsohn & el obeid 2002). the fire return period ranges between less than 2 years and 10 years and longer, with especially the north-eastern part of the study area being burned very frequently. as a combination of a variety of factors, including the fire return period, the seasonality of fires, et cetera, le roux (2011) determined a ‘fire regime’ for various areas in namibia. most of the study area is classed as having a ‘mild’ to ‘moderate’ fire regime. procedure at the same time as the vegetation survey in april 2004, a survey of the major landscape types and associated soil types was conducted. in addition to describing the landscapes based on field observations and topographic map interpretation, soil profiles were exposed by digging soil pits on representative sites throughout the study area (strohbach et al. 2004). however, because of the fairly labour-intensive sampling procedure, only 71 sample pits were dug. very few of these were at the same localities as the vegetation sample plots (strohbach 2014). this means that for this particular study, whilst also relying on existing geographic information system (gis) sources (le roux 2011; mendelsohn & el obeid 2002; mendelsohn et al. 2002; naris 2001), only extrapolated environmental data were available. it thus made sense to use synoptic composition data of the associations and subassociations for all analyses, rather than the individual relevé data, for comparison to the available environmental data; see the summarised data used in table 1. for all analyses, the panicum gilvum–marsilea vlei community was omitted as a potential outlier, also because only one relevé was collected in this community. analytical soil data from the soil profile descriptions were augmented with standard soil parameters developed for the world soil reference base (batjes 2002). available soil moisture soil hydrological parameters, specifically saturated hydraulic conductivity, saturation matric potential and porosity, can be directly linked to the soil texture, that is, the percentage of sand, silt and clay in the soil (cosby et al. 1984). we followed the convention of these authors and used the composition of the midpoint of the soil-textural class on the soil texture triangle. we used the soil characteristics of the dominant soil horizon in the case of soils with widely varying soil properties in their horizons; that is, we would ignore a shallow, sandy a-horizon and rather use the characteristics of the deeper, loamy b-horizon. soil hydrological parameters were derived from the textural classes, using data provided by cosby et al. (1984) and laio, porporato, ridolfi and rodriguez-iturbe (2001). no attempt was made to estimate the actual soil matric potential based on annual rainfall, soil texture and soil depth, because, in many cases (especially within the various arenosols), the depth of the soil profile is unknown, but it could reach depths of up to 250 m or deeper (mendelsohn & el obeid 2002). climatic data data on mean annual rainfall and frost days were extracted by overlaying plot positions onto the relevant gis layers. all other relevant climatic data (cv of rainfall, average minimum temperatures, relative humidity in both wet and dry seasons, etc.) were too uniform across the study area to warrant their consideration as significant environmental gradients (botha 1996; mendelsohn et al. 2002; naris 2001). the extracted data were averaged for the associations and these averages used in the analysis. in a similar way, the fire return period and fire regime (based on the data of le roux 2011) were extracted for plots and summarised for the associations. environmental gradients in order to demonstrate the main environmental drivers, the vegetation data were processed in two ways: by means of a non-metric multidimensional scaling (nms) ordination using pc-ord 6 (mccune, grace & urban 2002), as well as a classification of plant functional attributes using vegclass (gillison 2002; gillison & carpenter 1997). reciprocal averaging (hill 1973) was not used because of a severe arch effect. as the data set was relatively small, the use of nms, using the sørensen distance measure, was regarded as more feasible. the ordination procedure was completed by using pc-ord 5 (mccune et al. 2002) and repeated twice. the first run was based on a random starting configuration with 50 real data runs and 249 runs on randomised data. after 107 iterations a final solution was reached, with a stress of 7.196 and a final instability of 0.00001. the results were saved and used as base configuration for the second run. this time, only 10 runs with real data were allowed, also requesting a varimax rotation of the results. the environmental data were overlain as a biplot, to indicate the environmental gradients of interest. a second biplot was created, using plant functional attributes (see below) as variables. as water availability is hypothesised to be the main driving force in the composition of the vegetation, plant functional attributes (pfas) as related to water availability were utilised to illuminate these gradients. the pfas were chosen according to criteria described by gillison and carpenter (1997), in order that they might be used in the vegclass software for analysis. in the context of this study, all pinnately compound leaves have been classified as ‘filicoid’ (‘filicoid’ leaves are, according to the description of gillison & carpenter 1997, fern-like leaves). the pfas were identified from a variety of sources, including the tree atlas of namibia (curtis & mannheimer 2005), the field guide to trees and shrubs of namibia (mannheimer & curtis 2009), the field guide to the plants of northern botswana (heath & heath 2009), grasses of namibia (müller 2007), the prodromus zur flora von swa (merxmüller 1966) and a number of internet sites such as jstor plant science (http://plants.jstor.org), flora of zimbabwe (http://www.zimbabweflora.co.zw/index.php) and photo guide to plants of southern africa (http://www.southernafricanplants.net/index.php). the data were captured in vegclass, ensuring that individual species had the same functional attributes across syntaxa. this is contrary to the recommendations of gillison and carpenter (1997), but necessary as these pfas were collected in retrospect; furthermore, this is acceptable as these pfas represent the typical attributes of the species within the broad forest savanna and woodland of the northern kalahari (sensu giess 1998). the distance between all associations, based on changing pfa combinations, was calculated using the wald-wolfowitz distance measure as recommended by gillison and carpenter (1997). as the software lacks an option to reduce the resulting distance matrix to two dimensions, the resulting distances were plotted using the distance measures from association 14 (burkeo africanae–pterocarpetum angolensis) and association 1 (acacio fleckii–terminalietum prunioidis) and as axes 1 and 2, respectively. biplots of both the habitat and key pfa data were overlain onto this graph, again to elucidate the gradients. in all cases, the biplots were limited to those environmental variables or pfas with an r2 > 0.3. as a total of 31 pfas were identified (resulting in 123 combinations as plant functional types), it was decided to use only key pfas in biplot overlays onto ordination diagrams. these key pfas included the following: leaf size, leaf and photosynthetic adaptations, growth form (sensu raunkiaer – mueller-dombois & ellenberg 1974) and the number of perennial plants (opposing the growth form ‘therophytes’, which are annual plants). the leaf sizes have been combined as follows: microphyll leaves are all leaf sizes ≤ microphyll, sensu gillison and carpenter (1997), or ≤ 2025 mm2. macrophyll leaves are all leaf sizes ≥ notophyll, sensu gillison and carpenter (1997), or ≥ 2025 mm2. these key pfa data are represented in table 2, whilst the detailed pfa data are presented in online appendix 2. table 2: simplified key plant functional attributes, expressed as a percentage of all species found in the relevant associations. once the main environmental drivers were determined, the significance of these in affecting the compositional and plant functional changes between various vegetation associations was determined using a multiple linear regression analysis. for this, two runs of the procedure were completed, using the percentage occurrence of macrophyll and microphyll plants in each association as dependent variables and the mean annual rainfall, mean number of frost days, mean soil depth as well as soil texture, expressed as mean percentage sand, silt and clay, as independent variables. soil ph and electric conductivity was not used in this calculation, as these factors were assumed to be closely related to the soil texture. the f-value for variables to be entered in the regression was set to 0.5 for forward selection. for the calculation, the software statistica (2013) was used. results top ↑ the nms ordination results and the vegclass results are depicted in figure 3 and figure 4, respectively. the nms results represent 91.5% of the variation between the various associations, with axis 1 representing 52.8% and axis 2 representing 38.7%. figure 4: non-metric multidimensional scaling results of the synoptic vegetation data, depicting, (a) biplot overlaid with the habitat factors and (b) biplot overlaid with the key plant functional attributes. rainfall, soil depth and sandy soils are positively correlated with ordination axis 1, whilst the clay and silt content, combined with increasing ph, are negatively correlated with this axis (figure 4a). as expected, hydraulic conductivity is co-correlated with sandy soils along the main axis, whilst soil porosity, soil matric potential and soil pore volume are closely associated with the finer textured soils. these tendencies are partially confirmed by the vegclass results, with the soil texture also influencing the distribution of associations in relation to association 14 (burkeo africanae–pterocarpetum angolensis) (figure 5a). however, the soil hydraulic parameters do not seem to be of importance in the vegclass scatter plot. figure 5: scatter diagrams of the distances between various vegetation associations based on their plant functional attributes (using the distances from associations 14 and 1 on the x-axis and y-axis respectively), depicting, (a) biplot overlaid by the habitat data and (b) biplot overlaid with the key plant functional attributes. the main gradient manifests itself in a gradual change from mesophyll to microphyll plants – the mesophyll plants, as expected, being associated with the deep, sandy soils under higher rainfall regimes, whilst the microphyll plants are more restricted to the loamy soils (figure 4b). together with the microphylls, the loamy soils support a strong annual flora (therophytes) and plants with succulent leaves, whereas the sandy soils support macrophyll plants, perennials, especially phanerophytes, but also plants with filicoid leaves and deciduous plants. exactly the same combinations of pfas are found to correlate with the main gradient in the vegclass scatter diagram (figure 5b). the two results differ in the reaction to the second axis – within the nms ordination, cryptophytes (e.g. geophytes, geoxylic suffrutices sensu white 1976) are negatively correlated to axis 2, whilst chamaphytes (dwarf shrubs) react to the secondary axis of the vegclass results. no indication of a sensitivity to frost could be identified, whilst sclerophyll leaves are only very weakly correlated with the secondary axis of the nms (r2 < 0.2). fire regime did feature in the vegclass scatter plot. the fire regime in the study area generally varies between mild to moderate (le roux 2011) and does not show any tendency on the ordination diagram (figure 4a). it is more likely that with certain associations (especially the widespread combreto collini–terminalietum sericeae on the kalahari sand plains), fire is primarily responsible for the structure and finer compositional changes (figure 6). this, however, needs to be studied in greater depth. figure 6: fires have a severe structure-altering effect on bushlands and shrublands of especially the combreto collini–terminalietum sericeae typicum (subassociation 12.3), as seen from, (a) unburned bushland and (b) vegetation reduced to a low shrubland after a severe fire. the results of the multiple linear regressions indicate that, for both microphyll and macrophyll plants, rainfall is a significant driver in the occurrence of the plants. soil depth and soil texture (sand and clay content) were only insignificantly associated with this change in composition, whilst other factors did not obtain the necessary f-value to be taken up in the regression (table 3). table 3: multiple linear regression results for the dependent variables ‘macrophyll plants’ and ‘microphyll plants’, respectively. discussion top ↑ climatic gradients versus changes in soil types rainfall has, by far, the strongest and most consistent effect on the vegetation in arid to semi-arid climates. this also holds true for the vegetation within the present study area, which was proven to be the only significant driver in the change from microphyll to macrophyll vegetation (table 3). our present gradient is roughly 200 km long, with a rainfall gradient of less than 150 mm. skarpe (1996) described a similar change in vegetation as it occurs across botswana along a rainfall gradient over 1100 km on uniformly sandy soils; she unfortunately does not provide rainfall differences in her paper. from the geographical description, it is assumed to be a gradient from bokspits in far south-west botswana (177 mm mean annual precipitation – bhattachan et al. 2012) to the four-country corner with zimbabwe, namibia and zambia in north-east botswana. for nearby katima mulilo in north-east namibia, a long-term mean annual precipitation of 758 mm has been recorded (namibia meteorological services 1997). from this, a rainfall gradient of approximately 580 mm is assumed for skarpe's study. her main finding is that broad-leafed trees and grasses are positively correlated with increasing precipitation, whilst fine-leafed phanerophytic species and chamaphytic species were positively correlated with increasing aridity along the gradient. whereas skarpe (1996) describes a gradual change in vegetation over a rather lengthy gradient, the change from microphyll-dominant vegetation (associations a1 – a11) to macrophyll-dominant vegetation (associations a12 – a14) within the study area is rather abrupt over a very short geographical distance (cf. hüttich et al. 2009; strohbach 2014). these dramatic changes occur over virtually no rainfall gradient at all and are related to the change in soil type, from a fine-textured, loamy soil to the coarse-textured, deep sands of the kalahari basin, as suggested by leser (1972). tinley (1982) describes similar dramatic changes resulting from changes in soil texture for mozambique. these changes are caused by the difference in water availability as a result of differences in soil moisture relationships (in particular, a higher soil matric potential and higher porosity, but also higher adhesion, in loamy soils versus a high hydraulic conductivity in sandy soils), causing the inverse texture effect (figure 4) (austin et al. 2004; noy-meir 1973; porporato et al. 2001). the gradual change between the various subassociations of the combreto collini–terminalietum sericeae (a12) and the burekeo africanae–pterocarpetum angolensis (a14) from south to north happens on similar sandy soils and can be compared to the changes in pfas across a climatic gradient described by skarpe (1996). plant adaptations to environmental drivers owing to the higher hydraulic conductivity of sandy soils, rainfall infiltrates faster, but, at the same time, water percolates to deep soil layers. this means that water is available in the upper soil layers for a comparatively short period during the rainy season, whereas deeper soil layers (below 2 m) act as a reservoir for deeper-rooted plants (b. strohbach, unpublished data). therefore, deep-rootedness is a prerequisite for the survival of phanerophytes growing on such sandy soils. this holds true for most of these species (hipondoka & versfeld 2006; hipondopka et al. 2003; mwita et al. 2008; savory 1963; see also timberlake & calvert 1993). deep-rooted geoxylic suffrutices, for example entada arenaria and dichapetalum cymosum, better known from moister savannas and woodlands of the sudano-zambezian ecoregions (500 mm – 700 mm mean annual rainfall) (white 1976), thus also succeed in growing in this environment. the latter species (gifblaar or magou) is problematic throughout the study area, where it can increase rapidly in density by taking advantage of disturbances caused by high numbers of cattle (bester 1989). another adaptation to the temporary high availability of water in sandy soils during the rainy season (because of high infiltration rate) can be seen in stem succulents, specifically various commiphora species (figure 4b). these are able to photosynthesise, also during the dry season, by means of a photosynthetic layer under the translucent bark – an adaptation to xeric environments (referred to as ‘cortic photosynthesis’ by gillison & carpenter 1997). whether pinnately compound leaves (‘filicoid leaves’) point to any specific adaptation, or are rather a coincidence related to the species composition, could not be established. the grass sward of the sandy soils is dominated by the annual species aristida stipitata and megaloprotachne albescens. most grass roots tend to be limited to the upper 40 cm to 60 cm of the soil profile; they are therefore unable to access water from deeper water resources (hipondoka et al. 2003). with this annual character, the grass sward utilises a strategy of growing rapidly during the rainy season to exploit available resources to the fullest in the shortest possible time, but does not store water in underground organs or access the water that is available in deeper soil layers during adverse seasons. the few perennial grass tufts either have hairy leaves (e.g. digitaria seriata), or are extremely fine-leaved (e.g. stipagrostis uniplumis var. uniplumis) as an adaptation to this harsh environment. digitaria seriata is also often found within bush clumps, making use of both shade and increased water infiltration as a result of stem flow under these canopies. similar patterns were found in other arid environments (martinez-meza & whitford 1996; pressland 1973). perennial grasses within the kalahari are also known to utilise inverse hydraulic lift to channel water down into the soil profile (to a depth of between 50 cm and 100 cm). this is postulated as a potential mechanism (not only by grasses, but also by trees) to create an own ‘reservoir’ of soil water for use during the dry season (schulze et al. 1998). most of the phanerophytes growing on sandy soils are deciduous, shedding their leaves during the dry winter and hot, dry spring months (table 2, own observations). this is a typical adaptation to avoid water loss (fensham & fairfax 2009; seghieri, floret & pontanier 1995). many phanerophytes, especially terminalia sericea and various combretum spp., have hairy leaves, an adaptation typically associated with psammophil plants (warming 1909). within this study, deciduous plants were not limited to the sandy soils. many species growing on loamy soils exhibit the same adaptation; however, several species are distinctly evergreen (e.g. boscia albitrunca) or at least semi-deciduous, that is, shedding their leaves just before the next leaf flush (e.g. acacia erioloba, acacia hebeclada subsp. hebeclada). these evergreen to semi-deciduous species are also deep-rooted (canadell et al. 1996; moustakas et al. 2006; schulze et al. 1998), enabling them to obtain water from deep soil layers during the dry season. overall, however, the tendency is for the leaves to become distinctly smaller on the more loamy soils, an indication that the soil moisture availability is far lower on loamy soils than on sandy soils. an extreme case of edaphic aridity is presented by the shallow calcrete pans of the eragrostio echinochloideae–eriocephaloetum luederitziani (a4). in addition to the very shallow profile, the soil is characterised by a high skeletal content, a high ph (> 8) and a relatively high electric conductivity (> 100 µs/cm). moreover, the low soil volume can only store limited quantities of water, resulting in this water being saturated with ions, which means that it is not readily available to plants. the nature of these soils is manifested in the vegetation structure; the association being dominated by dwarf shrubs and, furthermore, within this association, 14% of all species had sclerophyll leaves (compared to an average of 6% – 9% for all other associations) and a relatively high microphyll leaf component (89%) (table 2). although fire is known to have a severe influence on the vegetation composition in the kavango and zambezi regions (geldenhuys 1977; le roux 2011; mendelsohn & el obeid 2003; mendelsohn & roberts 1997; strohbach & petersen 2007), only structural alterations were evident during the present study (figure 6). it is thus unlikely that fire is a driving factor in the discrimination of vegetation associations within the study area. shortcomings in data analysis a major shortcoming of this study is the fact that only synoptical data (both for vegetation composition as well as for environmental variables) could be employed. because of this, the clear soil gradients illustrated by the ordination techniques were not conclusive in the multiple linear regression. future studies concerning such moisture relations should take this problem into consideration. conclusion top ↑ the study supports the hypothesis that the broad-leafed savanna of the kalahari is limited in distribution by the deep sands, as a result of better soil moisture conditions. the improved soil moisture conditions in the kalahari sands are a result of the inverse texture effect (noy-meir 1973). in combination with increased rainfall to the north, this even allows tall trees to grow and replace the dominating shrublands with woodlands, as is evident from the transition between the combreto collini–terminalietum sericeae (a12) and the burkeo africanae–pterocarpetum angolensis (a14) – roughly coinciding with the 500 mm rainfall isohyet. deep-rootedness of phanerophytic species is one of the major adaptations of psammophil plants to utilise available water resources, but also to recharge and redistribute soil moisture resources in the profile (burgess et al. 1998; kizito et al. 2012; martinez-meza & whitford 1996; schulze et al. 1998). a better understanding of these mechanisms is essential if one wishes to gain a clear grasp of savanna dynamics. furthermore, predictions regarding the effects of global climate change on the vegetation in the kalahari environment can be improved by understanding the strategies employed by different plant species in the utilisation of water. it is predicted that, over the next 50 years, rainfall will decrease over namibia, resulting in less available grazing for extensive livestock production (midgley et al. 2005). in another study, it is predicted that the sands of the kalahari basin will be remobilised during the next 50 years, resulting in largely unvegetated mobile dunes, as a result of global climate change (thomas, knight & wiggs 2005). we postulate that climate change will manifest itself initially through a change in composition, from the present mesophyll savanna to a microphyll (acacia-dominated) savanna, very similar to that of the present camelthorn savanna sensu giess (1998). the camelthorn savanna forms part of the greater kalahari landscape on sandy soils, between about 250 mm and 400 mm mean annual rainfall, to the south of the present study area. it is, however, doubtful whether the grass sward will, under conditions of prolonged drought, develop to the present, relatively productive state. an added danger to productive savannas is the ongoing practice of harvesting terminalia sericea for fencing poles. already planned are projects to harvest ‘invader’ bush within the study area, to be used as fuel for power generation. such large-scale removal of shrubs and small trees from the already poor landscape will inevitably lead to reduced water infiltration into the soil (because of reduced stem flow) and reduced water redistribution within the soil profile, in turn leading to less favourable conditions for grasses. the net effect will be an accelerated desertification through a ‘man-made drought’. acknowledgements top ↑ thanks are due to ms marianne strohbach for assistance with the vegetation survey, as well as to mr heiner mouton for helping with the soil survey. the soil samples were analysed by the agriculture laboratory, directorate of agricultural research and training, in windhoek. this project was co-funded by the global environment facility through the desert margins programme and the government of namibia, through the recurrent budget of the directorate of agricultural research and training. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions b.j.s. (polytechnic of namibia) was the project leader. he was involved in the vegetation survey and was responsible for analysing the data and writing up the article. j.t.k. (ministry of agriculture, water and forestry) was responsible for the soil survey and description of the soil types. both contributed to the interpretation of the results. references top ↑ austin, a.t., yahdjian, l., stark, j.m., belnap, j., porporato, a., norton, u. et al., 2004, ‘water pulses and biogeochemical cycles in arid and semiarid ecosystems’, oecologia 141, 221–235. http://dx.doi.org/10.1007/s00442-004-1519-1 batjes, n.h., 2002, ‘soil parameter estimates for the soil types of the world for use in global and regional modelling’ (version 2.1), isric report 2002/02c, international food policy research insititute and international soil reference and information centre, wageningen. bester, f.v., 1989, ‘navorsing gedoen met betrekking tot gifblaar dichapetalum cymosum [research done concerning poison leaf dichapetalum cymosum]’, agricola 7, 60–64. bhattachan, a., tatlhego, m., dintwe, k., o'donnell, f., caylor, k.k., okin, g.s. et al., 2012, ‘evaluating ecohydrological theories of woody root distribution in the kalahari’, plos one 7, e33996. http://dx.doi.org/10.1371/journal.pone.0033996 botha, l., 1996, rainfall as an indicator of the agroclimate of namibia, mdiptech thesis, department of meteorology, technikon pretoria. brady, n.c. & weil, r.r., 2002, the nature and properties of soils, 13th edn., prentice hall, upper saddle river. burgess, s.s.o., adams, m.a., turner, n.c. & ong, c.k., 1998, ‘the redistribution of soil water by tree root systems’, oecologia 115, 306–311. http://dx.doi.org/10.1007/s004420050521 canadell, j., jackson, r.b., ehleringer, j.b., mooney, h.a., sala, o.e. & schulze, e.-d., 1996, ‘maximum rooting depth of vegetation types at the global scale’, oecologia 108, 583–595. http://dx.doi.org/10.1007/bf00329030 coetzee, m.e., 2003, ‘characteristics of namibian soils in a nutshell’, spotlight on agriculture 65. cosby, b.j., hornberger, g.m., clapp, r.b. & ginn, t.r., 1984, ‘a statistical exploration of the relationships of soil moisture characteristics to the physical properties of soils’, water resources research 20, 682–690. http://dx.doi.org/10.1029/wr020i006p00682 curtis, b. & mannheimer, c., 2005, tree atlas of namibia, national botanical research institute, windhoek. de pauw, e., coetzee, m.e., calitz, a.j., beukes, h. & vits, c., 1998, ‘production of an agro-ecological zones map of namibia (first approximation). part ii: results’, agricola 10, 33–43. food and agricultural organization & un educational, scientific and cultural organization, 1988, ‘soil map of the world – revised legend’, world soil resources report 60, fao, rome. fensham, r.j. & fairfax, r.j., 2009, ‘drought-related tree death of savanna eucalypts: species susceptibility, soil conditions and root architecture’, journal of vegetation science 18, 71–80. http://dx.doi.org/10.1111/j.1654-1103.2007.tb02517.x geldenhuys, c.j., 1977, ‘the effect of different regimes of annual burning on two woodland communities in kavango’, south african forestry journal 103, 32–42. http://dx.doi.org/10.1080/00382167.1977.9629467 giess, w., 1998, ‘a preliminary vegetation map of namibia’, dinteria 4, 1–112. gillison, a., 2002, ‘a generic, computer-assisted method for rapid vegetation classification and survey: tropical and temperate case studies’, conservation ecology 6, 3. gillison, a.n. & carpenter, g., 1997, ‘a generic plant functional attribute set and grammar for dynamic vegetation description and analysis’, functional ecology 11, 775–783. http://dx.doi.org/10.1046/j.1365-2435.1997.00157.x heath, a. & heath, r., 2009, field guide to the plants of northern botswana including the okavango delta, kew publishing, royal botanic gardens, kew. hill, m.o., 1973, ‘reciprocal averaging: an eigenvector method of ordination’, journal of ecology 61, 237–249. http://dx.doi.org/10.2307/2258931 hipondoka, m.h.t. & versfeld, w.d., 2006, ‘root system of terminalia sericea shrubs across rainfall gradient in a semi-arid environment of etosha national park, namibia’, ecological indicators 6, 516–524. http://dx.doi.org/10.1016/j.ecolind.2005.07.004 hipondoka, m.h.t., aranibar, j.n., chirara, c., lihavha, m. & macko, s.a., 2003, ‘vertical distribution of grass and tree roots in arid ecosystems of southern africa: niche differentiation or competition?’, journal of arid environments 54, 319–325. http://dx.doi.org/10.1006/jare.2002.1093 hüttich, c., gessner, u., herold, m., strohbach, b.j., schmidt, m., keil, m. et al., 2009, ‘on the suitability of modis time series metrics to map vegetation types in dry savanna ecosystems: a case study in the kalahari of ne namibia’, remote sensing 1, 620–643. http://dx.doi.org/10.3390/rs1040620 institut cartogràfic de catalunya, namibian ministry of agriculture, water and rural development & spanish agency for international co-operation, 2000, project to support the agro-ecological zoning programme (aez) in namibia. main report, icc, mawrd & aeci, windhoek. king, l.c., 1963, south african scenery, hafner publishing company, new york. kizito, f., dragila, m.i., senè, m., brooks, j.r., meinzer, f.c., diedhiou, i. et al., 2012, ‘hydraulic redistribution by two semi-arid shrub species: implications for sahelian agro-ecosystems’, journal of arid environments 83, 69–77. http://dx.doi.org/10.1016/j.jaridenv.2012.03.010 laio, f., porporato, a., fernandez-illescas, c.p. & rodriguez-iturbe, i., 2001, ‘plants in water-controlled ecosystems: active role in hydrologic processes and response to water stress. iv. discussion of real cases’, advances in water resources 24, 745–762. http://dx.doi.org/10.1016/s0309-1708(01)00007-0 laio, f., porporato, a., ridolfi, l. & rodriguez-iturbe, i., 2001, ‘plants in water-controlled ecosystems: active role in hydrologic processes and response to water stress. ii. probabilistic soil moisture dynamics’, advances in water resources 24, 707–723. http://dx.doi.org/10.1016/s0309-1708(01)00005-7 le roux, j., 2011, ‘the effect of land use practices on the spatial and temporal characteristics of savanna fires in namibia’, phd thesis, faculty of sciences, universitätsbibliothek der universität erlangen-nürnberg. leser, h., 1972, ‘geoökologische verhältnisse der pflanzengesellschaften in den savannen des sandveldes um den schwarzen nossob und epukiro [geo-ecological relationships in the savannas of the sandveld at the black nossob and epukiro]‘, dinteria 6, 1–44. mannheimer, c.a. & curtis, b.a. (eds.), 2009, le roux and müller's field guide to the trees and shrubs of namibia, macmillan education namibia, windhoek. martinez-meza, e. & whitford, w.g., 1996, ‘stemflow, throughfall and channelization of stemflow by roots in three chihuahuan desert shrubs’, journal of arid environments 32, 271–287. http://dx.doi.org/10.1006/jare.1996.0023 mccune, b., grace, j.b. & urban, d.l., 2002, analysis of ecological communities, mjm software, gleneden beach. mendelsohn, j.m. & el obeid, s., 2002, communal lands of eastern namibia, raison, windhoek. mendelsohn, j. & el obeid, s., 2003, sand and water. a profile of the kavango region, struik publishers, cape town. mendelsohn, j. & roberts, c., 1997, an environmental profile and atlas of caprivi, directorate of environmental affairs, namibia. mendelsohn, j., jarvis, a., roberts, c. & robertson, t., 2002, atlas of namibia, david phillips publishers, cape town. merxmüller, h. (ed.), 1966, prodromus zur flora von swa [prodromus of the flora of swa], j. cramer verlag, lehre. midgley, g., hughes, g., thuiller, w., drew, g. & foden, w., 2005, ‘assessment of potential climate change impacts on namibia's floristic diversity, ecosystem structure and function’, climate change research group, south african national biodiversity institute for the namibian national biodiversity programme, directorate of environmental affairs, cape town. moustakas, a., guenther, m., wiegand, k., mueller, k.-h., ward, d., meyer, k.m. et al., 2006, ‘long-term mortality patterns of the deep-rooted acacia erioloba: the middle class shall die!’, journal of vegetation science 17, 473–480. mueller-dombois, d. & ellenberg, h., 1974, aims and methods of vegetation ecology, john wiley & sons, new york. müller, m.a.n., 2007, grasses of namibia, 2nd edn., ministry of agriculture, water and forestry, windhoek. mwitwa, j.p., munthali, c.r.y. & van wyk, g., 2008, ‘heritability of shoot die-back and root biomass in sixteen pterocarpus angolensis (fabaceae) half-sib families from malawi, namibia and zambia’, southern forests: a journal of forest science 70, 221–226. namibia meteorological services, 1997, ‘monthly and annual rainfall normals for selected stations in namibia which are used in the 10-day rainfall bulletins’, working document from the namibia meteorological services, windhoek. namibian agricultural resources information system (naris), 2001, ‘naris agro-ecological zoning programme’, ministry of agriculture, water and rural development, windhoek. noy-meir, i., 1973, ‘desert ecosystems: environment and producers’, annual review of ecology & systematics 54, 25–51. http://dx.doi.org/10.1146/annurev.es.04.110173.000325 porporato, a., laio, f., ridolfi, l. & rodriguez-iturbe, i., 2001, ‘plants in water-controlled ecosystems: active role in hydrologic processes and response to water stress. iii. vegetation water stress’, advances in water resources 24, 725–744. http://dx.doi.org/10.1016/s0309-1708(01)00006-9 pressland, a.j., 1973, ‘rainfall partitioning by an arid woodland (acacia aneura f. muell.) in south-western queensland’, australian journal of botany 21, 235–245. http://dx.doi.org/10.1071/bt9730235 rodriguez-iturbe, i., porporato, a., laio, f. & ridolfi, l., 2001, ‘plants in water-controlled ecosystems: active role in hydrologic processes and response to water stress. i. scope and general outline’, advances in water resources 24, 695–705. http://dx.doi.org/10.1016/s0309-1708(01)00004-5 savory, b.m., 1963, ‘site quality and tree root morphology in northern rhodesia’, rhodesian journal of agricultural research 1, 55–64. schulze, e.-d., caldwell, m.m., canadell, j., mooney, h.a., jackson, r.b., parson, d. et al., 1998, ‘downward flux of water through roots (i.e. inverse hydraulic lift) in dry kalahari sands’, oecologia 115, 460–462. http://dx.doi.org/10.1007/s004420050541 seghieri, j., floret, c. & pontanier, r., 1995, ‘plant phenology in relation to water availability: herbaceous and woody species in the savannas of northern cameroon’, journal of tropical ecology 11, 237–254. http://dx.doi.org/10.1017/s0266467400008713 skarpe, c., 1996, ‘plant functional types and climate in a southern african savanna’, journal of vegetation science 7, 397–404. http://dx.doi.org/10.2307/3236283 southern african science service centre for climate change and adaptive land use (sasscal) weathernet, 2012, welcome to the weather stations of angola, botswana, namibia and zambia, viewed 30 october 2012, from http://www.biota-africa.org/weather/index.php statistica version 12, 2013, computer software, statsoft inc., tulsa. strohbach, b.j., 2008, ‘mapping the major catchments of namibia’, agricola 18, 63–73. strohbach, b.j., 2014, ‘vegetation of the eastern communal conservancies in namibia: i. phytosociological descriptions’, koedoe 56(1), art. #1116, 18 pages. http://dx.doi.org/10.4102/koedoe.v56i1.1116 strohbach, b.j. & petersen, a., 2007, ‘vegetation of the central kavango woodlands in namibia: an example from the mile 46 livestock development centre’, south african journal of botany 37, 391–401. http://dx.doi.org/10.1016/j.sajb.2007.03.002 strohbach, b.j., strohbach, m., kutuahuripa, j.t. & mouton, h.d., 2004, ‘a reconnaissance survey of the landscapes, soils and vegetation of the eastern communal areas (otjiozondjupa and omaheke regions), namibia’, unpublished report for the desert research foundation of namibia and the desert margins programme, national botanical research institute, windhoek. thomas, d.s.g., 1984, ‘ancient ergs of the former arid zones of zimbabwe, zambia and angola’, transactions of the institute of british geographers 9, 75–88. http://dx.doi.org/10.2307/621868 thomas, d.s.g., knight, m. & wiggs, g.f.s., 2005, ‘remobilization of southern african desert dune systems by twenty-first century global warming’, nature 435, 1218–1221. http://dx.doi.org/10.1038/nature03717 timberlake, j.r. & calvert, g.m., 1993, ‘preliminary root atlas for zimbabwe and zambia’, the zimbabwe bulletin of forestry research 10, forestry commission, harare. tinley, k.l., 1982, ‘the influence of soil moisture balance on ecosystem patterns in southern africa’, in b.j. huntley & b.h. walker (eds.), ecology of tropical savannas, ecological studies ser. 42, pp. 175–192, springer verlag, berlin. http://dx.doi.org/10.1007/978-3-642-68786-0_9 walter, h. & breckle, s.-w., 1984, spezielle ökologie der tropischen und subtropischen zonen, ökologie der erde [special ecology of the tropical and subtropical zones, ecology of the earth], vol. 2, gustav fischer verlag, stuttgart. walter, h., harnickell, e. & mueller-dombois, d., 1975, climate-diagram maps 1–36, springer verlag, berlin. warming, e., 1909, oecology of plants. an introduction to the study of plant-communities, oxford university press, london. werger, m.j.a. & ellenbroek, g.a., 1978, ‘leaf size and leaf consistence of a riverine forest formation along a climatic gradient’, oecologia 34, 297–308. http://dx.doi.org/10.1007/bf00344908 white, f., 1976, ‘the underground forests of africa: a preliminary review’, gardens’ bulletin singapore 29, 57–71. footnotes top ↑ 1.soil definitions follow the world soils reference base (food and agricultural organization & un educational, scientific and cultural organization 1988). a full description of the soil types found is presented in online appendix 1. article information author: jane carruthers1 affiliation: 1department of history, university of south africa, south africa correspondence to: jane carruthers email: carruej@unisa.ac.za postal address: po box 392, unisa 0003, south africa dates: received: 05 oct. 2010 accepted: 07 mar. 2011 published: 30 june 2011 how to cite this article: carruthers, j., 2011, ‘pilanesberg national park, north west province, south africa: uniting economic development with ecological design – a history, 1960s to 1984’, koedoe 53(1), art. #1028, 10 pages. doi:10.4102/koedoe.v53i1.1028 copyright notice: © 2011. the authors. licensee: aosis openjournals. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) pilanesberg national park, north west province, south africa: uniting economic development with ecological design – a history, 1960s to 1984 in this original research... open access • abstract • introduction • pilanesberg landscape and early politics • origins of the pilanesberg national park – the 1960s and 1970s • pilanesberg national park 1978–1983 • the process of revision 1983–1984 • conclusion • acknowledgements • references abstract (back to top) in the late 1970s, a ground-breaking project began in the pilanesberg district in what is now the north west province of south africa to create a wildlife conservation and eco-tourism venture from degraded marginal farmland in an aesthetically attractive extinct volcanic crater. the establishment of this national park was innovative in a number of respects, including a partnership between landscape and ecological designers, local community development and participation, regional tourist satisfaction, trophy hunting, environmental education, ecological restoration, and wildlife conservation and management. this paper briefly explored the park’s early history, explaining its landscape, its early peopling and historical land use. the narrative then concentrated on the first five years of the park’s existence, from its inception in 1977, under the aegis of agricor, bophuthatswana’s rural development agency, to 1984, when responsibility for the park was given over to bophuthatswana national parks, a parastatal agency, and a new era began. the article contended that 1984 is an appropriate date on which to conclude the early history of the pilanesberg national park (pnp) because it was then that the experimental phase of the park ended: its infrastructure was sufficiently developed to offer a satisfactory visitor experience, the management plan was revised, its bureaucratic structures were consolidated and an attitude survey amongst the local community was undertaken. embedding the originating period of the pnp in its historical, political and socio-economic context, the paper foregrounded those elements in the park’s beginnings that were new in the southern african protected area arena. thus, elements that relate to socio-politics, landscape and ecological design and restoration, and early relations with neighbouring communities were emphasised. this paper has been written by an historian and is therefore conceptual and historical, conforming in language and structure to the humanities style (environmental history). it relies on published and unpublished literature and oral information and the critical evaluation of these sources. conservation implications: the pioneering example of the pnp as a protected area is relevant to the field of conservation science because, as human population densities increase, as the tourism sector develops, as marginal farmland becomes available for new uses, and as it becomes important to include neighbouring communities in conservation activities, a study of this park’s early history and socio-political and economic context may be of assistance in the development of similar projects elsewhere in south africa and beyond. introduction (back to top) conservation managers and wildlife biologists in southern africa are familiar with the fact that national parks and other protected areas are often advertised and marketed as ‘unspoilt nature’, although they are, in fact, manipulated to meet objectives such as tourist satisfaction, carrying capacity, pasture and biodiversity management together with a variety of other key goals that may change over time. it is also true that many protected areas are far from being ‘pristine wilderness’ unaffected by past human activities, but are the consequence of ‘fortress conservation’ (brockington 2002) and the people (or their descendants) who were removed, often forcibly, in the interests of wildlife conservation have grievances that play out in the political arena. because of the human dimension involved in land use and present management, protected areas are not neutral spaces or landscapes without history; they are definitively shaped by their pasts.the pilanesberg national park (pnp), situated in what is today the north west province of south africa, was established in 1977. at the time, the area formed part of the western transvaal (a province of south africa until 1994) in an african reserve that was about to transform from an apartheid bantustan consisting of a number of ‘tswana homelands’ into bophuthatswana, nominally an independent state, an enclave within white south africa. it is an unusual national park from both points of view mentioned earlier. this particular area did not appear ‘natural’ at the time of its foundation. it was fully recognised to be property that had been heavily utilised and altered by many groups of people over the preceding centuries. it was deliberately and carefully redesigned as a national park, being restored ecologically from farmland and into which a wide variety of indigenous animals were reintroduced. once the white farmers had been expropriated (as will be explained at a later stage), the african people who returned to live in the pilanesberg were not to be forcibly removed as had been the case elsewhere; rather, their consent was sought and they were promised that the establishment of the national park would not be to their detriment but to their economic advantage. indeed, the very rationale of the pnp was that it was to become an engine of regional economic development. pilanesberg landscape and early politics (back to top) some 50 km north of the town of rustenburg lies the root zone of an extinct volcano – roughly circular, some 572 km2 (c.50 000 ha) in extent and measuring between 23 km and 28 km in diameter. it appears as a complex series of eroded rings of low mountains and hills that rise approximately 300 m – 600 m above the surrounding land. there is one perennial river and a number of freshwater and saline springs; however, the largest permanent body of water is mankwe dam (covering an area of approximately 2 km2), which was constructed by white farmers in the late 1950s. the climate is benign; the average rainfall is 600 mm – 700 mm per year, although there are regular droughts (farrell, van riet & tinley 1978; mccarthy & rubidge 2005; mucina & rutherford 2006). in terms of vegetation, the pilanesberg is significant because it is a transition zone between the arid savanna and the moist savanna biome. owing to the complex substrate, there is a wide variety of landscapes and habitats for both plants and animals (farrell et al. 1978; mucina & rutherford 2006). this landscape within the crater is aesthetically attractive and was the subject of comment by many 19th century travellers and explorers, amongst them thomas baines, who painted the ring of hills in 1869 as he journeyed into what is now botswana.the region had been inhabited continuously by tswana-speaking people probably for many centuries. at the time of permanent white settlement in the mid-19th century, the bakgatla bakgafela clan lived in the pilanesberg area under their chief pilane (d. 1850), who gave his name to the modern district. according to makgala (2009) and mbenga (1996), this community can also be referred to as the ‘kgafela-kgatla’, ‘bakgatla-baga-kgafela’ or ‘bakgatla-ba-ga-kgafela’ – meaning the ‘kgatla people of kafela’ in various forms of the tswana language. kgafela was the kgosi (chief) who originally gave his name to the community. the current head of the clan is kgosi kgafela ii, who resides in mochudi, botswana, whilst kgosi nyalala pilane leads the group at saulspoort. the bakgatla was one of the few groups that did not resist the arrival of the boers but became their allies, assisting them in exploits of war and ivory-hunting. their association with white settlers and access to firearms enhanced the bakgatla powerbase and enabled them to increase their cattle herds and vanquish some of their local rivals and enemies. in later decades, however, the bakgatla lands were commandeered and carved up into settler farms on which the bakgatla became rent-paying or labour tenants. many of the community settled at saulspoort (breutz 1953; makgala 2009; manson & mbenga 2009; mbenga 1996, 1997; mbenga & morton 1997; morton 1992, 1995; schapera 1953). in 1913, the natives land act (act no. 27 of 1913; union of south africa) confined black south africans to very limited areas of the country and, in 1936, the native trust and land act (act no. 18 of 1936; union of south africa) attempted to provide more land for africans by designating ‘released areas’ that were to be purchased by the south african native trust and added to the african reserves. the pilanesberg was one of these released areas and the white-occupied farms were very slowly expropriated until the exercise was complete in the early 1960s. the bakgatla were thus allowed to return to their ancestral land. in 1961, the growing severity and oppression of apartheid politics affected the pilanesberg directly when it became a designated ‘tswana homeland’. during the 1970s, these various tswana homelands were consolidated into a number of islands within white south africa and became the ‘independent nation’ of bophuthatswana in 1977. regional politics were fraught. tidimane pilane, kgosi of the bakgatla, and lucas mangope, head of the bahurutshe clan, were rivals, pilane supporting the african national congress (anc) and mangope the apartheid state (butler, rotberg & adams 1977; jones 1999; lawrence & manson 1994). origins of the pilanesberg national park – the 1960s and 1970s (back to top) the principles underlying wildlife and conservation management in southern africa were changing during the 1960s and 1970s (carruthers 2007a, 2007b, 2008). in terms of philosophy, the idea of utilising wildlife sustainably by cropping and translocation gained ground in parts of africa, replacing an older tradition of strict preservation. at the same time, there were technical improvements in the transporting and immobilisation of wild animals that led to the easier movement and sale of wildlife (dasmann 1959, 1964; dasmann & mossman 1960, 1961; eltringham 1984; johnson et al. 2008; mossman & mossman 1976). in conjunction with bophuthatswana politics, these developments were relevant to the formation of the pnp. indeed, the park could not have come into being in a pre-translocation and pre-game sales era.apartheid social planning gained momentum during the 1960s and there were government initiatives to make the bantustans more self-sufficient economically and thus able to sustain a larger number of black africans outside of ‘white’ south africa, with a view to separating the homelands permanently from the other ‘white’ parts of the country. in 1969, there was a recommendation from ‘apartheid’s social engineers’ – a ‘potchefstroom-based team of “development experts” commissioned to find ways of enhancing the economic viability of an “independent state”’ (van onselen 1996:477) – that the crater be made into a recreation resort and nature reserve. however, for reasons that are unclear, nothing came of the idea at the time, but it was raised again in 1973. the following year mangope established a feasibility study (brett 1989; johnson et al. 2008; magome & collinson 1998). the matter received a boost when the southern sun hotel group – which, through managing director sol kerzner, had close ties with the bophuthatswana president and his government – finalised a plan to build a casino and hotel resort in the pilanesberg (to be named ‘sun city’) that would bring revenue into the region. at a time of strict petrol rationing and thus the curtailment of long-distance motor car travelling, it was expected that a game reserve adjacent to the hotel would provide an added attraction for tourists from johannesburg and pretoria, who would flock to sun city for the kinds of entertainment not available in white south africa, such as multiracial mingling, soft pornography and gambling. having first considered the location of mankwe dam for the hotel, the facility was relocated to its present site and construction began in 1978 (bureau for economic research re bantu development [south africa] 1978; boonzaaier pers. comm., 01 march 2010). after gaining independence, bophuthatswana established a number of organs of state. one of these was a parastatal development body tasked with promoting rural self-sufficiency. named the agricultural development corporation (agricor), this body fell under the bophuthatswana department of agriculture and was to play a decisive role in the establishment of the pnp. through its managing director, david beuster, agricor raised the funds for the game reserve and, despite the fact that many bakgatla people and livestock lived in and used the crater, and that it contained numerous farm houses, roads, dams and fences, the pilanesberg was formally proclaimed a nature reserve in 1977. it is worth emphasising that agricor, a parastatal body specifically responsible for economic and community development in the rural sector, was given the administration of this future national park rather than the department of nature conservation, because it was regarded as a rural improvement project and not a nature conservation exercise. what was audacious for the period was that beuster and mangope employed landscape architects to design this game reserve adjacent to sun city. the firm that was instructed to act as consultants to draw up a management plan was farrell and van riet, landscape architects and ecological planners, then a recently established pretoria-based company, and it was instructed to act as consultants and to draw up a management plan. by the time he established the partnership of farrell and van riet, willem van riet was a leader in the field of landscape architecture in south africa and he was primarily responsible for linking landscape architecture with ecological planning in the country. van riet had initially qualified as an architect at the university of cape town but thereafter, from 1972 to 1975, he had benefited from studying at the university of pennsylvania under ian mcharg, the renowned landscape architect and author of design with nature (1971), a book that is widely regarded as one of the most influential of the 20th century (schnadelbach 2001). in his autobiography, mcharg explained that the genesis of design with nature lay in a meeting between himself, russell train, the president of the conservation foundation and ray dasmann, a noted ecologist and the foundation’s chief scientist. apparently, train said, ‘ian, ray and i have decided that the time has come for a book on ecology and planning’ and mcharg agreed to write it (mcharg 1996). train and dasmann were correct: the book was perfectly timed and widely used and quoted. in a quest for life (1996), mcharg describes landscape architecture as a discipline very close to nature and its preservation and he was particularly keen to encourage planning that was appropriate to specific environments. mcharg sought out trained architects such as van riet for his postgraduate landscape architecture programme, providing not only a stimulating academic environment but substantial financial subsidies (mcharg 1981). in this way, and through van riet, ideas around ecologically apt planning and design from the usa made their way into southern africa. sharon kingsland argues that the science of nature reserve design emerged in tandem with the interdisciplinary field of conservation biology. she explains that the basic rationale for such design is to protect biodiversity, using ideas from island biogeography, prioritising conservation of the indigenous species of plants and animals of the area, and employing operational research and mathematical techniques for linear programming (kingsland 2002a, 2002b). if this is the norm, then the pilanesberg was highly unusual because the biodiversity had been totally compromised by farming activities and there were extremely few remaining indigenous plants and animals – certainly large mammals had become locally extinct. the creation of the pilanesberg involved little conservation biology and focused, at first, entirely on ecological restoration and landscape design. the national park emerged from the collaboration between van riet and ken tinley, a young ecologist who had also been inspired by mcharg. as a university student, tinley advocated mcharg’s design with nature to his contemporaries (huntley 2010). tinley was one of an emerging new generation of wildlife ecologists in south africa (many of whom were educated at the university of natal). in partnership with van riet, he worked on a number of nature reserves, particularly in the ‘homelands’, where there was scope for new ideas because the reserves in these localities were not in the control of the various philosophically and bureaucratically entrenched provincial nature conservation authorities or the national parks board. these included locations in pondoland (including mkambati), in maputaland and in the gorongosa national park in mozambique, as well as private game reserves (farrell & van riet landscape architects and ecological planners 1975; tinley 1978; tinley 2010; van riet 2010). van riet and tinley were employed to design the pnp and they presented their report in 1978. what they suggested was somewhat revolutionary in the context of southern african national park and game reserve planning and it marked a strong contrast to the fortress conservation and wildlife management practices that then held sway. entitled ‘pilanesberg national park: planning and management proposals for department of agriculture, republic of bophuthatswana’ (farrell et al. 1978), this is an important document and worth summarising in some detail. the report began with what was a provocative premise in an era of fortress conservation: that any conservation measure would ultimately be futile unless wildlife and nature could deliver tangible, visible benefits to humans within a particular socio-economic and geographical milieu. in other words, the survival of wildlife in africa was dependent on rural african people. van riet and tinley (farrell et al. 1978) argued that protected areas should not be viewed in isolation, but in their regional ecological and economic contexts as productive primary (ecological services) and secondary (tourism, education and wealth-creation) landscapes. the report paid particular attention to wildlife as a source of protein, as well as of traditional medicine and other natural products that might be sustainably harvested by local people, together with wildlife tourism being a source of employment and income. these principles were in sharp contrast with those espoused by, for example, the national parks board (now sanparks) that were focused on settler values that emphasised white, middle class tourist recreation and created places in which: lessons in tidiness, adherence to and acceptance of rules and regulations … [were taught] [where] people can be disciplined not to litter, not to pick flowers … [and that generate] tranquillity … so desperately needed in a world where people are caught up in the tensions of city life. (knobel 1979:233) in contradistinction to this viewpoint that urban visitors and romantic ideas of wilderness were the focus of nature conservation initiatives, tinley (1979) had expounded the philosophy that the regional context of any conservation project was decisive. he strongly believed that the needs and aspirations of rural people were paramount and argued on the basis that many conservation departments throughout africa: have based their activities on the dogma that tourism and wildlife conservation are two sides of the same coin. thus staff and funds are used mostly for catering and tourist facilities and the natural areas become filled with urban nuclei to justify the existence of parks. in this way conservation departments continue to be directly responsible for despoiling the last wild places for which they are custodians. (tinley 1979:33) the pilanesberg was to be different and the aim was to reflect a new vision of conservation practice in africa. not only did they introduce the radical idea of using national parks sustainably as engines of regional development sensitive to local community needs, van riet and tinley also introduced a novel concept of planning and design (farrell et al. 1978). after surveying the geomorphology and other aspects of the landscape, habitats and vegetation cover, their report proposed that in order to maximise wildlife viewing in the small area and utilise it to the best advantage, all major tourism facilities should be located on the boundaries of the park, thus preserving the interior of the crater from unsightly camps, restaurants and other amenities. this peripheral development was different from other national parks and game reserves which had normally sited major visitor accommodation within the protected area itself. moreover, using the internal watersheds as ecological borders, van riet and tinley suggested dividing the crater into seven (later reduced to five) distinct activity zones. there would be zones for trails, hunting, visitors, wilderness, special use, multiple use and peripheral development. there would be no roads or amenities in the wilderness and special use zones and a buffer zone would separate these two from the resource utilisation areas, the rest camps and the intensive use areas. there were specific recommendations for planning and using each of these zones so that activities would harmonise with each other (farrell et al. 1978). the pilanesberg project was highly unusual in that van riet and tinley had a free hand and a flexible institutional, bureaucratic and policy environment within in which to work. they were not burdened with an entrenched public service, hidebound politicians, an historical legacy of preservation philosophy, or outdated or ill-sited roads and other tourist amenities (child 2008). moreover, the pilanesberg crater, with its rings of hills that hid the plains and human developments beyond from view, was the ideal topography in which to recreate a natural-looking environment. whilst at the start there were cultivated lands, evidence of stock grazing and farmsteads, alien vegetation, roads, and so on, once these – and the people – were removed, the crater presented an almost clean slate for design. a list of appropriate mammals that should be introduced was provided in the section on ‘management proposals’ (farrell et al. 1978) and appropriate herd sizes given. also departing from the then accepted norm in managing protected areas, van riet and tinley made provision for trophy hunting and prioritised environmental education. in short, the report designed a 50 000 ha national park, literally from the bottom up. pilanesberg national park 1978–1983 (back to top) in the opening chapter of their report, van riet and tinley had made strong statements about the holistic philosophy of sustainable national parks and the role of local people within them, but, on the whole, they focused on planning the future ecological management of the pnp (farrell et al. 1978). it appears that they assumed that politicians and sociologists would take care of the human and community dimensions of the enterprise. unlike other national parks and protected areas in south africa that had involved forced removals, the idea for the pnp was that the bakgatla would participate in decisions about the new national park, vacate the crater area voluntarily, contribute to its social and economic planning, and to its management thereafter. in this regard, the fact that the new bophuthatswana ‘nation’ planned a national park was important in terms of nation-building, and the pilanesberg was to be the public demonstration of these ‘civilised’, modern and international values (carruthers 1997). early in 1979, negotiations were completed between van riet and tidimane pilane and, in turn, between pilane and the other bakgatla chiefs (keenan 1984). the bakgatla, through their kgosi, agreed to surrender their grazing and land rights in the crater. just how ‘voluntary’ this agreement was, has later been hotly contested in a land restitution claim instigated by the bakgatla who aver that they were strongly coerced by the ‘strong-arm tactics’ employed by mangope and his officials (mbenga 2011) who were determined to steamroller the game reserve through for the benefit of sun city and its supporters. in any event, the bakgatla did not control the entire pilanesberg crater. the national park consisted of freehold land of 8500 ha, obtained directly from the resident bakgatla (viz. schaapkraal, welgeval and portions of legkraal, koedoesfontein, kruidfontein, saulspoort, rooderand and doornpoort), whilst some 4500 ha came from a newly arrived group, the bakubung, namely wydhoek and portions of ledig and koedoesfontein. in addition, 1000 ha was obtained from private owners and the rest – the majority of the property – consisted of 46 000 ha of state land that had been expropriated from white people by the department of bantu affairs (as explained earlier) to augment the paucity of land allocated to africans (keenan 1984:14). the bakgatla apparently acquiesced in their relocation from the pilanesberg crater on the basis that they would be allocated two nearby state farms to replace their lost communal grazing and that they would be fully compensated for the land and structures that were required by the national park. in addition, they would be recompensed for the full costs of removal and also retain the right to enter the reserve in order to visit graves and to collect firewood, thatching material and medicinal plants. moreover, they were promised an (unspecified) portion of entry ticket sales and tidimane pilane was to be appointed onto the national park’s governing and management board of trustees, thus ensuring a bakgatla a voice in a form of joint management (magome & collinson 1998; makgala 2009). it was unfortunate that these agreements were both informal and verbal and thus neither effectively witnessed nor formally contractual (van riet 2010). subsequently, it emerged that pilane, a political opponent as has been previously explained, had been threatened by mangope with eviction from other state land if he did not agree to the pnp proposal (keenan 1984:16–17). moreover, in later years, complaints surfaced that pilane, who was not a universally popular leader, had not adequately consulted with the rest of the community (makgala 2009:33−335) and thus did not speak for everyone. today, there is considerable sophistication in all quarters when consulting communities affected by national parks and other protected areas, but at the time of the pnp’s establishment there were no protocols to follow and no mechanisms for predicting or resolving disputes or difficulties that might arise between the various parties involved. there were no such examples to follow in the 1970s and in the absence of experience on the part of van riet and the bakgatla – and in the context of high apartheid and mangope’s dictatorship – many details were left vague or unrecorded. one needs to recall, however, that the bakgatla were not the only people affected by the establishment of the pnp. whilst tidimane pilane, as a traditional kgosi, was in ostensible control of bakgatla ‘communal land’, the farm welgeval inside the crater was inhabited by a community of long standing in the area, who owned part of the property in their own right. they agreed to relocation, provided they were fully compensated and this agreement was formally documented. it is on the basis of this written evidence that the welgeval community has subsequently been awarded a land restitution claim on this farm and the land has been leased back to the pnp (manson & mbenga 2009). many other local people were also not consulted, presumably either because they were considered to be fractured groups without leaders to give them voice, or perhaps because they lived on the borders of the pilanesberg rather than within it, or, even, perhaps because they were mangope dissidents and were thus ignored. thus many people were dissatisfied with, and disadvantaged by, these arrangements. for example, the farm ledig (south of the pilanesberg) was occupied by the bakubung, part of a disunited tswana-speaking group that had been forcibly removed from the outskirts of the small town of boons where they had formed a ‘black spot’ within white south africa and were therefore obliged to relocate in the late 1960s. they used the crater for grazing their cattle and goats. apparently, as far as these people were concerned, ‘care was taken to ensure that all talk of the project was kept away from the bakubung notables and strictly confined to official circles’ (van onselen 1996: 477). soon, without warning, there were reports of ‘a giant game fence snaking across the pilanesberg’, and excluding them and their livestock (van onselen 1996: 498). there were also large numbers of non-tswana nguni-speakers, many of whom had also been forced into a homeland from urban areas or white farms, who also had to make use of the pilanesberg for their survival. it seems evident from oral sources and later comments that, whilst negotiations and participation had occurred at top political levels, the views of ordinary people had neither been sought nor taken into account. this particularly included those who were not part of the formal ‘tribal’ structures of the district, newcomers and outsiders and who perhaps would pay the heaviest price in terms of losing access to land and livelihood opportunities (keenan 1984:39–43). owing to its bold conception and future plan, the pilanesberg project received considerable local and international publicity. local people were negative about the creation of the pnp because it impacted directly on their lives, whilst many scientists and conservationists were critical of its ambitious ‘operation genesis’ – the mass reintroduction of many species of wildlife. added to the difficulties (and the adverse publicity) was that development and administration did not proceed smoothly. the personnel was generally incompetent and many officials of agricor lacked experience and, for this and other reasons, they were either removed from their posts or encouraged to resign (brett 1989:112). it was only in october 1980 that jeremy anderson, whose doctoral research in zululand and subsequent study on lion management in the umfolozi game reserve (natal) had gained him a reputation as a capable and knowledgeable wildlife manager, was employed by beuster as director. anderson – with his scientific expertise, energy, enthusiasm and familiarity with the new scientific thinking emanating from east africa – was tailor-made for the job (boonzaaier pers. comm., 01 march 2010). anderson was joined by willem boonzaaier, previously employed in the private sector, as chief administrative officer to handle the financial side of the operation. between them, they appointed qualified and appropriate staff, including ecologists roger collinson and peter goodman in 1981. those involved in these early days recall the magnetism of being associated with what was then an experimental nature reserve, of working in a multiracial environment, and of encountering the dominant personalities and interesting characters who sought to put pilanesberg on its feet (owen-smith, magome & grossman pers. comm., 01 january 2011). the success of the pnp was predicated on that of sun city, and the resort prospered to the extent that it was expanded in 1981 and again in 1984. the close friendship between kerzner and mangope meant that the bophuthatswana political elite in the mangope government were extremely supportive of the sun city development, as was the south african regime; however, tidimane pilane was less enthusiastic for a number of reasons. firstly, he was a member of the opposition party, an anc supporter with a different vision for the country’s future, and he disagreed with many of the policies of the mangope government. secondly, he believed that sun city offered no benefit to local people because employment in the resort went to outsiders to the district (makgala 2009:322–323), a grievance shared by the bakubung. thirdly, tourist revenue, one of the major reasons why the bakgatla had agreed to vacate the crater in favour of a national park, was minimal, despite the promises that visitors from sun city would flock to the reserve, and no financial gain came their way. the pnp was also criticised because wildlife introductions had not gone as smoothly as planned. owing to the fact that some species were being given away freely by south african conservation bodies that had problems of over-stocking, whilst others were inexpensive to acquire, the pilanesberg became stocked with an incorrect balance of wild animals. this resulted in some habitats being inappropriately modified because of over-grazing, leading to even further reduction in biodiversity and condemnation from local and international scientists (van aarde 2010). for reasons of veterinary and disease control, it was difficult to obtain appropriate species because wildlife movements were curtailed from places in which cattle diseases were endemic. wildlife therefore had to be sourced from disease-free populations a long distance away, such as namibia or the eastern cape, and transport was thus extremely costly. additionally, because adult elephant males can pose a danger during the capture and transport process, fewer of them were translocated in comparison with females and young and thus subsequent breeding success was low and herd sizes and composition were skewed (garaï et al. 2004; hancock 1983). the mixing of gene pools was also a matter of scientific concern. some introductions were made before the fencing was complete and animals were kept in a holding camp that was too small, which resulted in many animal deaths once the grazing inside the camp was depleted (collinson & anderson 1984:169–70). these introductions of large mammals were extremely costly and the bophuthatswana government was not able to fund them. the money came from the south african wildlife foundation (sawf), founded in 1968 by afrikaner business magnate anton rupert, who was a trustee of south africa’s national parks board and who had close ties with the national party. in 1979, rupert presented the pnp as a project to the world wildlife fund international and the international union for the conservation of nature and received approval to fund it (schwarzenbach 2010). given rupert’s personal belief in cultural and linguistic ethnicity, assistance to the ‘nation’ of bophuthatswana through the sawf was entirely in character. indeed, bophuthatswana may have held special significance for the rupert family, for his wife, huberte (neé goote), came from the western transvaal and she spent some years at derdepoort on the botswana border (domisse 2005:353). as can be appreciated from the aforementioned explanations, the early years of the pnp presented huge challenges to those involved. there was no existing institutional policy framework – either scientific or bureaucratic – within which to operate. whilst this meant freedom from precedent, it also meant that the enterprise moved slowly. owing to the limited number of rangers and labourers that could be employed, the process of restoring a natural environment – a ‘mammoth task’ that hancock (1984) referred to as ‘renaturalisation’ – was extremely slow. anderson (pers. comm., 23 february 2010) recalled that in terms of basic infrastructure virtually nothing in existed and a perimeter fence, off-loading ramps, translocation stations, bomas, feedlots and pens had all to be constructed by the small staff. more than 1000 km of internal farm fencing had to be removed, as had the many solid concrete cattle dips, farm reservoirs and windmills, about 30 large farmhouses and over 100 smaller houses, outbuildings and huts. these were bulldozed and the rubble was used to fill large dongas (hancock 1984). borrow pits, landscape scars and old lands had to be rehabilitated and general farming detritus (e.g. old vehicles and heavy, rusted implements) cleared. invasive alien plants were abundant, not only jointed cactus (opuntia spp.), but huge old trees, especially australian eucalypts – all these had to go, some being felled, others poisoned (anderson pers. comm., 23 february 2010; hancock 1984). in addition, there was no living or office accommodation for game guards, management or labourer staff or for stores. prefabricated buildings were erected to meet some of these needs but, for a few years, all management personnel and their families lived in caravans. workshops and vehicle maintenance points were also needed. the construction of permanent buildings, game-viewing hides and visitor amenities such as camps and entrance gates proceeded slowly, as did re-siting of roads to make them suitable for game drives (old straight farm roads had to be obliterated). the local staff was unskilled and thus training, education and mentorship had to be provided. at this time there was no regular telephone communication in the park and only radio phones could be used – a scarce and expensive resource (anderson pers. comm., 23 february 2010). veld and wildlife monitoring systems also had to begin anew and proceed in tandem with applied management tools such as a burning regime. jules turnbull-kemp, a senior game ranger recruited by agricor from rhodesia, and who later became warden of the pnp, was responsible for receiving the wildlife introductions. at the time, there were no formal studies to assist with determining the ability of different species to survive or thrive on old farmlands and only by observation and experience did it emerge how animal populations coped with, and altered with, the recovery and restoration of the habitat (turnbull-kemp pers. comm., 01 march 2010). because large-scale models (such as the kruger national park) could not be applied to a small area like the pnp, fire policy also had to be determined from scratch. an innovation in terms of management philosophy was that stocking was carried out at a high rate (i.e. many animals at one time) so that the take-off rate (reducing numbers through hunting) was optimised as soon as possible (turnbull-kemp pers. comm., 01 march 2010). not surprisingly given the dearth of wildlife and abundant unsightly evidence of former fields, houses and roads in the crater, tourism did not take off quickly, indeed tourists were not encouraged for a number of years until a ‘satisfactory game viewing experience’ could be guaranteed (hancock 1984). anderson had doubted that gate revenue from day visitors (which was to be the major source of income for the bakgatla) was ever likely to produce any substantial income (boonzaaier pers. comm., 01 march 2010) and so took the decision to introduce trophy hunting to generate some immediate income. because the pnp was an agricor initiative, and was the only project of this nature in its stable, the administrative and financial arrangements were as independent as – and could be as experimental as – those of the conservation management. as administrative officer, boonzaaier adapted commercial systems to the park’s requirements. there was no model to follow: the park had to be up and running as quickly and profitably as possible. the first question was: where was an initial income to come from? in this regard, wildlife management and administration were able to dovetail. anderson’s idea of revenue-producing trophy hunting could only take place if there were surplus animals to shoot. to determine the optimum stocking rate that would be needed to manipulate species numbers to obtain the best returns, anderson, collinson and boonzaaier designed a complex model to determine how many (and which) species were required to profit most from game sales, hunting, meat production or tourist viewing. wildlife populations were therefore predicated on formulas that demonstrated the best return on investment, per land unit (anderson pers. comm., 23 february 2010; boonzaaier pers. comm., 01 march 2010). the pnp’s management was innovative because, instead of a few wild animals of various species being introduced and then allowed slowly to build up their numbers, large populations were introduced at the start and thus numbers increased very quickly, providing a surplus after only a year or two in the case of some species (anderson 1986). very careful records of net production versus utilisation were maintained. these calculations were novel because they were being made for the first time in a protected area: wildlife was being taken into account as a financial asset, not merely a ‘nature conservation’ ethical good. just as cattle and game farmers entered their herds into their accounting books and measured the profit from them, so too did this national park (boonzaaier pers. comm., 01 march 2010; a more detailed history of wildlife management and conservation science in the pnp will be the focus of a later paper by the present author.) however, all these developments took time. moreover, the early 1980s saw one of the subcontinent’s worst droughts of the century. this meant that the rehabilitation and restoration of the pnp grasslands and vegetation took far longer than anticipated; it also meant that the displaced bakgatla and others were short of grazing on the farms to which they had been relocated and many looked longingly at the recovering (albeit slowly) veld in the pnp that had been free of grazing cattle for a few years (manson & mbenga 2009). during the drought, mobile pnp animals wrought havoc on properties outside the reserve. baboons, in particular, climbed over the perimeter fences and ravaged the maize fields of neighbouring black communities in ledig and elsewhere (van onselen 1996:510). the process of revision 1983–1984 (back to top) by 1984, despite obstacles and slow progress, it could be said that the pnp was maturing and that it had come to a stage when an overall review to guide its future strategic direction was required. in that year, a new management plan was devised, the bophuthatswana national parks board was founded and a community relations survey was conducted. considerable experience had been gained by park management during the five years since the park’s opening and the pnp began to meet some of its objectives. as hancock (1984) described in response to those ‘wondering exactly what, if anything, has been happening in the pilanesberg’, during this time the park had been fenced, the landscape rehabilitated, buildings razed and obliterated, tourist roads constructed, wildlife introduced, two visitor camps constructed and foot safaris and trophy hunting operated satisfactorily. it seemed that the experimental phase was ending and that consolidation and review was needed (collinson & goodman 1982). tinley was not involved in the re-planning process as he had left south africa by that time. the new report, ‘a five year development plan for pilanesberg national park as requested by the bophuthatswana government and the bophuthatswana national parks board, september 1983’, was authored by willem boonzaaier, roger collinson and willem van riet. because the construction of tourist facilities within the national park had been so slow and the project had been so costly, these managers feared that the investment of the previous five years in management, rehabilitation and wildlife introductions might be wasted unless clearer objectives were re-established. the primary goal of the pnp was stated to be to ‘maintain and where necessary create an ecosystem comprising a biota of as wide a variety of indigenous plant and animal species’ as possible (boonzaaier, collinson & van riet 1983). the secondary objective was defined as ‘to utilise the area and its natural resources in ways that will yield the greatest benefits to bophuthatswana and its people, both now and in the future’ (boonzaaier et al. 1983). the multiple zoning of the initial plan had proved to be too complex to manage effectively. boonzaaier, collinson and van riet thus recommended re-zoning the park into two overarching types of areas, a ‘managed natural area’ and a ‘natural environment recreation area,’ each subdivided into zones. within a ‘managed natural area’ (i.e. well within the park’s boundaries), accommodation would be limited and cater for very small groups, with the only permissible activities being walking on designated trails and trophy hunting. the ‘natural environment recreation area’ would be devoted to general visitor and multiple uses and would include peripheral development at the manyane, bakubung and bakgatla gates. however, these plans would be extremely expensive and the park would probably continue to run at a loss whilst development proceeded through various phases (boonzaaier et al. 1983). the year 1984 was also significant because the overall managerial and bureaucratic structures of the park were altered when a national parks and wildlife management board for bophuthatswana was created along the lines of a parastatal to manage the pnp. this new structure, which was formalised in 1987 with the national parks act (act no. 24 of 1987; bophuthatswana government), resulted from the merger of the division of nature conservation of the department of agriculture and forestry and agricor, thus ending their somewhat competitive and even acrimonious relationship. the new national parks organisation, nicknamed bopparks, was to be managed by an appointed board. this move was significant because it meant that the pnp was now freed from its roots in a rural development organisation and was provided with a more conventional home within a parks board. this was to impact on its later trajectory. the need to review, and if necessary to change, the objectives and management of the pilanesberg in 1984 may also have been related to political and economic factors inherent in the bophuthatswana state. at this time bophuthatswana was economically stressed and, in fact, was experiencing ‘an acute financial crisis’ (jones 1999, 2001). there was also growing political dissension and even violence as a rupture developed between the democratic and inclusionist policies of anc supporters in bophuthatswana, including tidimane pilane, and the mangope faction, with its ideal of an ethnic tswana nation in an artificially segregated south africa (jones 1999). within the broader south (and southern) african political landscape, violence, revolutionary activities, harsh repression and military intervention were endemic and an atmosphere of tension was the order of the day. many of the flashpoints thereof were in the so-called ‘independent bantu states’. in this atmosphere of political turmoil it was clear that relations between the pilanesberg’s managers and the bakgatla had deteriorated. the matter was aggravated by the fact that tidimane pilane was sidelined when he was not appointed to the bopparks board as he should have been in terms of the ‘agreement’ with van riet (magome & collinson 1998; makgala 2009:321). this affront marginalised and offended the bakgatla, who were anc allies and thus opposed to mangope’s regime. together with the fact that no monetary compensation was accumulating for the community (which is what they had been promised) because there were few visitors and thus little by way of gate fees, relations between the pnp and the bakgatla were tense because of ‘broken promises’. perhaps the initial undertakings of beneficiation had been over-generous, but the bophuthatswana government had apparently reneged on agreements about land and financial compensation to people the state regarded as political opponents. in terms of the breakdown in communication between neighbours and the pnp, magome and collinson (1998) believe that it owed much to the heavy demand for very rapid development and effective wildlife and administrative management, which meant that park authorities had little time to devote to nurturing community relationships. in order to identify and address issues of concern, it was decided to conduct a formal survey of the attitudes of the local people to the pnp. in 1984, jeremy keenan, a sociologist then employed at the university of the witwatersrand (and who did not disguise his anti-bophuthatswana views), was tasked to report on community relations (keenan 1984:5). keenan and his researchers uncovered seriously negative perceptions of the national park at many levels. there was discontent over the verbal initial arrangements regarding the evacuation of the crater and inadequate financial and property compensation, particularly as cattle-rustling and other theft of property occurred during the removals. the bakgatla perceived the administration of bophuthatswana and its officials as ‘dictatorial and deceitful’ (keenan 1984) and they alleged that farms intended for compensation had been given away to government ministers and mangope cronies. to demonstrate their discontent, they had decided that they wanted to take back their land in the pnp and, to this end, had begun a court action (keenan 1984). the fact that tidimane pilane was a leading figure in opposition politics exacerbated the situation further. people involved at the time believe that the pnp was used to score political points in these oppositional politics at a time of unrest in south africa, creating divisions even amongst bopparks staff members (boonzaaier pers. comm., 01 march 2010). however, despite the grievances and the misunderstandings about the function of a national park that were reflected, keenan’s report also indicated that there was some local support for the pnp and for its educational outreach programme in particular (keenan 1984:67–74). because the report was leaked to the media by keenan himself, it attracted a great deal of attention that resulted in the managers of the pilanesberg being caught in the middle of the fracas between the government and the bakgatla (magome & collinson 1998). conclusion (back to top) it is now nearly 35 years since the establishment of the pnp and if the argument here has been that circumstances had shifted so much in the first five years of its existence that a review was necessary, then how much more has the context changed in the 30 succeeding years. not only did bophuthatswana itself undergo violent revolutions in 1988 and 1994, which saw lucas mangope and all that he had stood for overthrown, but the republic of south africa itself underwent a peaceful democratic revolution in 1994 that totally transformed the political environment of the country. this had enormous repercussions through every organ of government, as well as impacting greatly on the socio-economic environment, including in the conservation arena. in the post-1994 ‘new’ south africa, the management philosophy, objectives and style have altered considerably in both the pnp and in other game reserves in what is now north west province.nature conservation – although some argue that tourism income and economic beneficiation determines policy more than biodiversity conservation (johnson et al. 2009) – has expanded in the north west province and it has retained the use of ‘national park’ even though the region is no longer a separate ‘nation’. whilst governance of the protected areas in north west has shifted with the changing responsibilities of various provincial departments, thanks to the original mission of the pnp to assist rural development and upliftment, natural resource management as an income-generating, employment-creating and capacity-building exercise remains a high priority in the region. despite the political uncertainties, the pnp had become successful by 1991 and the bophuthatswana government then took over marginal farmland on the botswana border, reclaiming and stocking it in the same way as pnp to create the upmarket madikwe game reserve. after 1994, and the establishment of the north west parks and tourism board (note the inclusion of tourism in the name of this body) plans were to form a substantial heritage corridor that would link the protected areas. events subsequent to 1984 were extremely important and warrant further study. moreover, the lessons learnt around ecological restoration and wildlife and conservation management require careful research and evaluation, as do their influence in other parts of south africa. nonetheless, it is instructive to reflect on how the creation and management of the pilanesberg national park during its early years introduced a number of fresh developments into south african natural resource design, management and conservation, some of which have become more important in the current protected area estate. whilst not adopted universally, some of the experimental aspects of the pilanesberg, including peripheral development, the consumptive use of wildlife in protected areas, the provision of a variety of visitor accommodation, community engagement and local empowerment, reclamation of farmland, translocation of wildlife, trained african senior personnel, environmental education, and a commercialisation and concession policy, have now become a part of modern conservation practice in both state-owned and private protected areas. research relating to how the innovations in landscape design, wildlife management and community issues later influenced developments in other protected areas is currently being undertaken by the present author and it is anticipated that another, more focused publication in this regard will result. acknowledgements (back to top) i would like to thank the following for their generous assistance in providing inspiration, guidance, information, documentation, discussion and for correcting factual and editorial errors: jeremy anderson, willem boonzaaier, bruce brockett, vincent carruthers, roger collinson, david grossman, brian huntley, cynthia kemp, hector magome, bernard mbenga, eugene moll, archie mossman, sue mossman, norman owen-smith, alexis schwarzenbach, ken tinley, rudi van aarde and willem van riet. financial support from the university of south africa, the national research foundation, and the garden and landscape studies department of harvard university is also gratefully acknowledged. adapted from a paper delivered at the 2010 dumbarton oaks garden and landscape studies symposium, 'designing wildlife habitats' (14 and 15 may, 2010), and included in the forthcoming volume wildlife habitats (2013) published by dumbarton oaks research library and collection. references (back to top) anderson, j.l., 1986, ‘restoring a wilderness: the reintroduction of wildlife to an african national park’, international zoological yearbook 24/25, 192–199. doi:10.1111/j.1748-1090.1985.tb02538.xbureau for economic research re bantu development (south africa), 1978, bophuthatswana at independence, bureau for economic research re bantu development (south africa), babelegi. boonzaaier, w.v., collinson, r.f.v. & van riet, w.f., 1983, ‘a five year development plan for pilanesberg national park as requested by the bophuthatswana government and the bophuthatswana national parks board, september 1983’, unpublished report. bophuthatswana government, 1987, national parks act, bophuthatswana government, bophuthatswana. brett, m.r., 1989, pilanesberg: jewel of bophuthatswana, frandsen, sandton. breutz, p.-l., 1953, the tribes of rustenburg and pilansberg districts. ethnological publications, no. 28, department of native affairs, pretoria. brockington, d., 2002, fortress conservation: the preservation of the mkomazi game reserve, tanzania, james currey, oxford. butler, j., rotberg, r.i. & adams, j., 1977, the black homelands of south africa: the political and economic development of bophuthatswana and kwazulu, university of california press, berkeley. carruthers, j., 1997, ‘nationhood and national parks: comparative examples from the post-imperial experience’, in t. griffiths & l. robin (eds.), ecology and empire: environmental history of settler societies, pp. 125–138, keele university press, edinburgh. carruthers, j., 2007a, ‘conservation and wildlife management in south african national parks 1930s–1960s’, journal of the history of biology 412, 203–236. carruthers, j., 2007b, ‘influences on wildlife management and conservation biology in south africa c. 1900 to c. 1940’, south african historical journal 58, 65–90. doi:10.1080/02582470709464745 carruthers, j., 2008, ‘“wilding the farm or farming the wild”: the evolution of scientific game ranching in south africa from the 1960s to the present’, transactions of the royal society of south africa 63(2), 160–181. doi:10.1080/00359190809519220 child, b., 2008, ‘recent innovations in conservation’, in h. suich & b. child (eds.), parks in transition: evolution and innovation in wildlife conservation: parks and game ranches in transfrontier conservation areas, pp. 277–287, island press, washington dc. collinson, r.f. &. anderson, j.l., 1984, ‘problems, principles, and policy in the reintroduction of large mammals in conservation areas’, acta zoologica fennica 172, 169–170. collinson, r.f.h. & goodman, p.s., 1982, ‘an assessment of range condition and large herbivore carrying capacity of the pilanesberg game reserve, with guidelines and recommendations for management’, inkwe 1, 1–55. dasmann, r.f., 1959, environmental conservation, john wiley & sons, new york. dasmann, r.f., 1964, african game ranching, pergamon, oxford. dasmann, r.f. & mossman, a.s., 1960, ‘the economic value of rhodesian game’, rhodesian farmer 30(51), 17–20. dasmann, r.f. & mossman, a.s., 1961, ‘commercial use of game animals on a rhodesian ranch’, wild life 3(3), 7–14. domisse, e., 2005, anton rupert: a biography, tafelberg, cape town. drummond, j.h., 1990, ‘rural land use and agricultural production in dinokana village, bophuthatswana’, geojournal 22(3), 335–343. doi:10.1007/bf00711346, doi:10.1007/bf00192833 eltringham, s.k., 1984, wildlife resources and economic development, john wiley & sons, chichester. farrell, van riet & tinley, k.l., 1978, ‘pilanesberg national park, bophuthatswana: planning and management proposals for department of agriculture, republic of bophuthatswana, august 1978’, unpublished report. farrell & van riet landscape architects and ecological planners, 1975, ‘planning and management proposals for the dwesa forest reserve, transkei’, prepared for the department of forestry and nature conservation, umtata, unpublished report. garaï, m.e., slotow, r., carr, r.d. & reilly, b., 2004, ‘elephant reintroductions to small fenced reserves in south africa’, pachyderm 37, 28–36. hancock, p., 1983, ‘elephant restocking at pilanesberg’, african wildlife 37(6), 227. hancock, p., 1984, ‘pilanesberg game reserve: four years later’, african wildlife 37(1), 47−51. huntley, b.j., 2010, email, 24 march, brianjhuntley@googlemail.com johnson, s.r., boonzaaier, w., collinson, r. & davies, r., 2008, ‘changing institutions to respond to challenges: north west parks, south africa’, in h. suich & b. child (eds.), parks in transition: evolution and innovation in wildlife conservation: parks and game ranches in transfrontier conservation areas, pp. 289–305, island press, washington dc. jones, p.s., 1999, ‘“to come together for progress”: modernisation and nation-building in south africa’s bantustan periphery – the case of bophuthatswana’, journal of southern african studies 25(4), 579–605. doi:10.1080/030570799108489 jones, p.s., 2001, ‘the etiquette of state-building and modernization in dependent states: performing stateness and the normalization of separate development in south africa’, geoforum 33, 25–40. doi:10.1016/s0016-7185(01)00010-0 keenan, j., 1984, ‘report on the socio-economic effects of the pilanesberg game reserve on the surrounding population and the attitudes of the surrounding population to the game reserve, university of the witwatersrand, on behalf of the pilanesberg game reserve, south africa, 1984’, unpublished report. kingsland, s., 2002a, ‘creating a science of nature reserve design: perspectives from history’, environmental modeling and assessment 7, 61–69. doi:10.1023/a:1015633830223 kingsland, s., 2002b, ‘designing nature reserves: adapting ecology to real-world problems’, endeavour 26(1), 9–14. doi:10.1016/s0160-9327(00)01396-x knobel, r., 1979, ‘the economic and cultural values of south african national parks’, in i. player (ed.), voices of the wilderness, pp. 230–239, jonathan ball, johannesburg. lawrence, m. & manson, a., 1994, ‘the “dog of the boers”: the rise and fall of mangope in bophuthatswana’, journal of southern african studies 20(3), 447–461. doi:10.1080/03057079408708413 magome, d.t. & collinson, r.f.h., 1998, ‘from protest to pride: a case study of pilanesberg national park, south africa’, the world bank/wbi’s cbnrm initiative, viewed 16 july 2007, from http://srdis.ciesin.columbia.edu/cases/south_africa-0003.html makgala, c.j., 2009, history of the bakgatla-baga-kgafela in botswana and south africa, bakgatla-baga-kgafela tribal administration, moruleng/saulspoort. manson, a. & mbenga, b., 2009, ‘the evolution and destruction of oorlam communities in the rustenburg district of south africa: the cases of welgeval and bethlehem, 1850s–1980’, african historical review 41(2), 85–115. mbenga, b.k., 1996, ‘the bakgatla-baga-kgafela in the pilanesberg district of the western transvaal from 1899 to 1931’, d. litt et phil thesis, department of history, university of south africa. mbenga, b.k., 1997, ‘forced labour in the pilanesberg: the flogging of chief kgamanyane by commandant paul kruger, saulspoort, april 1870’, journal of southern african studies 23(1), 127–140. doi:10.1080/03057079708708526 mbenga, b.k., 2011, email, 10 january, bernard.mbenga@nwu.ac.za mbenga, b. & morton, f., 1997, ‘the missionary as land broker: henri gonin, saulspoort 269 and the bakgatla of rustenburg district, 1862–1922’, south african historical journal 36(1), 145–167. doi:10.1080/02582479708671273 mccarthy, t. & rubidge, b., 2005, the story of earth and life, struik, cape town. mcharg, i.l., 1971, design with nature, doubleday, new york. mcharg, i.l., 1981, ‘human ecological planning at pennsylvania’, landscape planning 8, 109–120. doi:10.1016/0304-3924(81)90029-0 mcharg, i.l., 1996, a quest for life: an autobiography, john wiley & sons, new york. morton, f., 1992, ‘slave-raiding and slavery in the western transvaal after the sand river convention’, african economic history 20, 99–118. doi:10.2307/3601632 morton, f., 1995, ‘land, cattle and ethnicity: creating linchwe’s bakgatla, 1875–1920’, south african historical journal 33(1), 131–154. doi:10.1080/02582479508671851 mossman, s.l. & mossman, a.s., 1976, wildlife utilization and game ranching: report on a study of recent progress in this field in southern africa, iucn, morges. mucina, l. & rutherford, m.c. (eds.), 2006, ‘the vegetation of south africa, lesotho and swaziland’, strelitzia 19, 1–807. schapera, i., 1953, the tswana, international african institute, london. schnadelbach, r.t, 2001, ‘ian mcharg’, in j.a. palmer (ed.), fifty key thinkers on the environment, pp. 228–241, routledge, london. schwarzenbach, a., 2010, email, 04 may, alexis.swarzenbach@bluewin.ch tinley, k.l., 1978, ‘mkambati nature reserve: an ecological and planning study, 1978’, unpublished report. tinley, k.l., 1979, ‘the maintenance of wilderness diversity in africa’, in i. player (ed.), voices of the wilderness, pp. 230–243, jonathan ball, johannesburg. tinley, k.l., 2010, email, 20 september, kentinley@optusnet.com.au union of south africa, 1913, natives land act, union of south africa, pretoria. union of south africa, 1936, natives trust and land act, union of south africa, pretoria. van aarde, r.j., 2010, email, 11 march, rjvaarde@zoology.up.za van onselen, c., 1996, the seed is mine: the life of kas maine, a south african sharecropper 1894–1985, david philip, cape town. van riet, w., 2010, email, 26 march, wvanriet@ppf.org.za abstract introduction methods results ethical considerations discussion conclusion acknowledgements references about the author(s) leigh combrink school of life sciences, university of kwazulu-natal, south africa the endangered wildlife trust, johannesburg, south africa hendrik j. combrink department of veterinary tropical diseases, university of pretoria, south africa andré j. botha the endangered wildlife trust, johannesburg, south africa colleen t. downs school of life sciences, university of kwazulu-natal, south africa citation combrink, l., combrink, h.j., botha, a.j. & downs, c.t., 2017, ‘habitat structure and diversity influence the nesting success of an endangered large cavity-nesting bird, the southern ground-hornbill’, koedoe 59(1), a1438. https://doi.org/10.4102/koedoe.v59i1.1438 original research habitat structure and diversity influence the nesting success of an endangered large cavity-nesting bird, the southern ground-hornbill leigh combrink, hendrik j. combrink, andré j. botha, colleen t. downs received: 29 aug. 2016; accepted: 10 july 2017; published: 17 nov. 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract habitat features can have a profound effect on the nesting success of birds. savannas are often managed with predators and large herbivores as priority species, with little thought to the many bird species that management decisions could affect. using a data set spanning seven breeding seasons, we examined how nesting success of southern ground-hornbills (sghs) bucorvus leadbeateri in the kruger national park varied as a result of various environmental and habitat factors within a radius of 3 km surrounding the nest site. identifying which factors affect nesting success will allow for targeted management efforts to ensure the long-term survival of sghs both within and outside of protected areas. habitat structure and diversity of the vegetation surrounding the nest were the most influential factors on sgh nesting success. sghs require open grassy areas for foraging and areas with large trees for nesting. savanna habitat drivers such as elephants and fire should be managed to ensure that sufficient large trees are able to establish in the landscape and to control for bush encroachment. this is especially important in areas earmarked for sgh reintroductions. nest sites of sghs should be monitored to mitigate any structural changes in the habitat surrounding the nests. nests should be modified or artificial nest sites provided, where nests have been damaged or lost, to ensure the continued presence of these birds in african savannas. conservation implications: habitat structure and diversity surrounding southern ground-hornbill nests has a significant impact on their nesting success. this highlights the importance of monitoring vegetation change in savanna habitats where they occur. management of savanna areas should take factors that influence bush encroachment, such as fire and elephants, into account to ensure the long-term persistence of these birds. introduction determining nesting success is fundamental to understanding the breeding biology of birds (dinsmore, white & knopp 2002). there are many factors that affect nesting success, such as the availability of high-quality habitat and nesting sites (loegering & anthony 2006; paredes & zavalaga 2001; vickery, hunter & wells 1992), food availability (boulton, richard & armstrong 2008; oro, pradel & lebreton 1999; rastogi, zanette & clinchy 2006), predation pressure (boulton et al. 2008; oro et al. 1999; rastogi et al. 2006; robinson et al. 2000; zarones et al. 2014), landscape effects (habitat loss and fragmentation) (boulton et al. 2008; reidy, thompson & peak 2009; winter et al. 2006) and human disturbance (blackmer, ackerman & nevitt 2004; woolaver et al. 2014). identifying the suite of habitat characteristics that influence particular species and populations is important for management decisions for the conservation of these species (martin 2014). the southern ground-hornbill (sgh) bucorvus leadbeateri inhabits savannas throughout much of africa and is currently listed as vulnerable, mainly as a result of habitat loss, land-use change and persecution (birdlife international 2016). in south africa, this species is considered endangered and has suffered significant declines throughout its range (a.c. kemp & r. webster unpubl. data; taylor, peacock & wanless 2015), with around 50% of the national population residing within the kruger national park (kemp 1995). sghs are terrestrial, carnivorous and co-operative breeders. they generally occur in groups of between 2 and 11 birds (kemp, joubert & kemp 1989), consisting of an alpha breeding pair and related subordinate helpers (kemp 1995). they are secondary cavity nesters and occupy territories within the kruger national park ranging from approximately 3000 ha – 12 000 ha (l. combrink unpubl. data). nests are most often located in natural cavities in large trees, but are occasionally in cliffs or earth banks (kemp 1988). the majority of known nests within the kruger national park are in trees, with only 1 of the 38 nests referred to in this study being a nest in a cliff face. females lay two eggs, 3–7 days apart. only the first-hatched chick is provided for, while the second-hatched chick mostly perishes as a result of starvation (kemp 1995). the sgh species recovery plan (jordan 2011) highlights the use of captive-reared birds and reintroductions as conservation interventions for this species. however, the exact habitat requirements for sghs are considered a knowledge gap (jordan 2011). before reintroductions of sghs can occur, it is essential to first understand the factors that affect sgh nest site selection and that influence their nesting success. kemp and kemp (1991) suggest that suitably sized nest cavities for sgh are limited within savannas. however, in their study from 1967 to 1995, kemp and begg (1996) did not find any of the nest site characteristics tested to have an influence on sgh nesting success. recent changes in management strategies in the kruger national park, in particular those relating to fire and elephants (van wilgen et al. 2008; young, ferreira & van aarde 2009), could have significantly affected the survival and recruitment of large trees. this in turn could affect the availability of suitably sized cavities in the landscape. wilson and hockey (2013) found that sgh groups nesting in natural cavities were more successful when there was 3 km of open woodland surrounding the nest. they also found that sgh groups using artificial nests were more successful than those occupying natural nests. their study was conducted in a private nature reserve adjacent to the central kruger national park which is 180 000 ha in extent (around 9% the size of the kruger national park). using data from seven complete breeding seasons of nest monitoring (2008–2015), we investigated what factors affected the nesting success of sgh groups in the kruger national park. specifically, we asked (1) whether the harvesting of the second-hatched chicks impacted on the survival to fledging age of first-hatched chicks and (2) what nest characteristics, habitat and environmental factors affected overall nesting success of sghs. we also tested whether the habitat structure within 3 km surrounding the nest and aspects of the nest site itself affected sgh nesting success for all vegetation types with known sgh nests throughout the kruger national park. using our results, we suggest possible management implications and interventions needed to ensure the long-term sustainability of the sgh throughout its range. methods study area this study was conducted within the kruger national park, south africa (22–26°s, 30–32°e), which comprises around 2 million ha with an average annual rainfall of 350 mm – 750 mm (gertenbach 1980). the park is largely divided longitudinally with more granitic soils in the west and basaltic soils in the east (venter 1990). the habitat consists mainly of savanna, with pockets of dense woody vegetation within broader grasslands (gertenbach 1983). nest surveys we monitored all known and accessible sgh nesting sites within kruger national park (n = 38) from the 2008/2009 breeding season until 2014/2015 (figure 1). the breeding season for sghs coincides with the wet season, from october to april each year. initial nest checks were usually conducted in november, with active nests being revisited throughout the breeding season to determine the outcome of the nesting attempt. in some instances, the second-hatched chick was removed or harvested from the nest for the purposes of being included in the captive rearing and reintroduction programme (jordan 2011). characteristics and measurements of each nest site were also collected. the nest site characteristics that we measured were: diameter of the base of the cavity (length and breadth), depth of the cavity (nest lip to base), volume of the nest (length × breadth × height), height of the nest entrance from the ground and the diameter at breast height of the nest tree. we were only able to collect chick survival and nesting success data from one cliff nest, which is completely inaccessible for measurements to be taken. only measurements from nest tree cavities were included in this analysis (n = 37). figure 1: the locations of southern ground-hornbill nesting sites within the kruger national park, south africa. chick survival and nesting success the nestling period in sghs is around 86 days in length. nestlings were fitted with alphanumeric metal rings and colour rings at 60–70 days for future individual identification. we considered the nestling to have fledged and the nest successful if the nestling reached ringing age. reports of sightings of the fledged ringed chicks have confirmed that this is a reasonable assumption (l. combrink unpubl. data). chick survival was recorded as 0 if the nest failed and 1 if the nest was successful. nests were also considered to have failed if the eggs were infertile or the eggs or chicks were predated. as a result of the large distances and travel times between sghs nests in the kruger national park, it was not possible to calculate daily nest survival rate. we calculated nesting success per group over the entire study period using the following formula: nesting success is equal to the number of years where the chick survived divided by the number of years where the outcome was known. years, where the outcome was not known, were excluded from the analyses. this yielded a proportion between 0 and 1. models were weighted using the number of years of known outcome. nest habitat gis layers obtained from the south african national parks (sanparks) were used to extract environmental variables related to each sgh nest. sghs are thought to be central-place foragers, concentrating their breeding season activity around the nest site. we, therefore, included a circular buffer with a radius of 3 km around each nest for the purpose of extracting the related environmental variables and to test wilson and hockey’s (2013) proposal about the area of open woodland surrounding each nest across the extent of the kruger national park. habitat was classified according to the 2013–2014 south african national land cover data set (geoterraimage 2014). for each 3 km buffer zone, the proportions of the various land types were calculated. these data were then used to calculate the shannon-wiener diversity index (shannon & weaver 1949; spellerberg & fedor 2003) as a measure of vegetation diversity, using the r package vegan (oksanen et al. 2016). insect species richness has been shown to be positively correlated with the normalised difference vegetation index (ndvi – measure of vegetation greenness) (pettorelli et al. 2011). in the kruger national park, the ndvi calculated for the growing or wet season is highly correlated with above-ground biomass and rainfall (wessels et al. 2006). for the nesting success models, the mean seasonal ndvi and mean ndvi yield (seasonal biomass increase) values were averaged for the seasons where the outcome or fate of the nest attempt was known. mean seasonal ndvi and mean ndvi yield layers were obtained from the mod13q1 product of version 6 of the modis 16 day terra vegetation indices data set (didan 2015) via the modistools package (tuck & philips 2015). soil type was classified as the proportion of granite, basalt, gabbro, ecca shale and rhyolite within the buffer zone and expressed as a percentage. the length (total number of kilometres) of all streams and rivers and the distance from the nest to the nearest stream were calculated using quantum gis (qgis). similarly, length (total number of kilometres) of all tourist and management roads, and distance from the nest to the nearest road (tourist or management) were included. rainfall for the current breeding season (mm), the previous breeding season (mm) and the previous non-breeding season (mm) were determined using sanparks’ rainfall data, collected at weather stations throughout the kruger national park. the rainfall data from the weather station closest to each nest site were used as a proxy for rainfall within the buffer zone for the particular time period. the mean annual rainfall (percentage of the buffer zone within the various rainfall categories – see table 1) was calculated using the most current continuous vector layer of interpolated rainfall data provided by sanparks. table 1: fixed effects included in the southern ground-hornbill nesting success land and water generalised linear mixed models. data analyses we conducted all statistical analyses using r (version 3.2.2; r core team 2015). a generalised linear mixed model with a binomial distribution and logit link function was applied using the lme4 package (bates et al. 2015) for both chick survival and nesting success. in the chick survival model, chick survival was the response variable and treatment (whether the nest was harvested or not) included as a fixed effect. group (defined using the nest name) and year were included as random effects, to account for any effects on chick survival related to the individual group or season, which were not explicitly measured or modelled here. for the nesting success models, nesting success was the response variable with group included as a random effect. some of the predictor variables were found to be highly correlated (r ≥ 0.7) and these were examined separately against the response variables to determine which produced the model with the lowest akaike information criteria (aic) ranking. only those variables were kept in the model. where two correlated variables had the same effect in the model, the variable that was present for the larger number of samples was retained. owing to rank deficiency, we could not include all predictor variables in one model for nesting success. for this reason, similar variables (water related, habitat related and nest related) were grouped together and these models were compared with the final selected models having the lowest aic rankings. for sgh chick survival, we had a treatment model looking at the effect that removing the second-hatched chick from the nest had on the first-hatched chick. for nesting success, we had three models namely, a nest model looking at nest dimensions, tree species, et cetera; a land model focusing on habitat related features; and a water model including streams, rainfall, et cetera. further variables in the land and water models were excluded, owing to rank deficiency, as their contribution to the buffer zone areas were negligible or only present for a small number of nests. for the nest models, nest cavity width, breadth, depth and height of cavity entrance were all highly correlated with nest volume (r ≥ 0.7). as a result, only nest volume and diameter at breast height were included in the nest models as fixed effects. all models with significant variables were checked for over dispersion and drop1 (bates et al. 2015) was then used to determine the model with the best fit and lowest aic value. if significant results were produced, the p-values for the final models were then adjusted using false discovery rate (fdr) to account for multiple comparisons on a small data set (pike 2011) using the base package in r (r core team 2015). table 1 details the fixed effects that were included in the respective land and water models for nesting success. results chick survival we monitored 100 sgh nesting attempts in the kruger national park during the study, for which the outcome of the nesting attempt was known. of these 100 nesting attempts, 76 were successful and 24 failed. second-hatched chicks were harvested from 32 of these nesting attempts (l. combrink unpub. data). we tested the effect of the removal of the second-hatched chick on the survival to fledging age of the first-hatched chick (treatment model), but found no significant impact of the nest treatment (estimate = 0.8538, standard error = 0.7704, z = 1.108, p = 0.2677). although not significant, the log odds of the first-hatched chick surviving to fledging age when harvesting is conducted, were 92% and when the nests were only monitored, the log odds of the first-hatched chick surviving to fledging age were 83%. this indicated that removing the second-hatched chick had no significant effect on the survival to fledging age of the first-hatched chick, although there does seem to be some benefit to the first-hatched chick when the second-hatched chick is removed. nesting success of the 37 sgh tree nests in our study, cavities had a mean length of 48.7 cm ± 2.0 cm (30 cm – 71 cm), a mean breadth of 43.1 cm ± 2.6 cm (25 cm – 102 cm) and a mean depth of 49.3 cm ± 4.2 cm (11 cm – 160 cm). all nests were situated in trees with a mean diameter at breast height of 129.6 cm ± 17.0 cm (64 cm – 544 cm) and cavities were located at a mean height of 5.6 m ± 0.4 m (3 m – 12 m) from the ground. we modelled group nesting success as a factor of the various nest site parameters, but did not find that any played a significant role (nest volume [p = 0.948], dbh [p = 0.179]). nesting success was calculated for 38 sgh nests (37 tree nests and 1 cliff nest) across the kruger national park. the best of the land models relating habitat features to nesting success (tables 2 and 3) included all four of the major habitat classes (thicket and dense bush, grassland, low shrubland and bare ground), latitude, longitude, percentage of gabbro soils, distance from the nest to the nearest road and habitat diversity. although longitude and the low shrubland habitat type did not have significant effects on nesting success, removing them from the model increased the aic, possibly owing to them being almost significant after adjusting the p-values. for the water model, none of the variables included showed any significant effect on overall nesting success (table 4). table 2: output of generalised linear mixed model selection comparing nesting success of southern ground-hornbills. table 3: parameter estimates, standard errors, z values, p-values and adjusted p-values (using false discovery rate) for variables in the best land model for southern ground-hornbill nesting success (with the lowest akaike information criteria). table 4: parameter estimates, standard errors, z values, p-values and adjusted p-values (using false discovery rate) for variables in the best water model for southern ground-hornbill nesting success (with the lowest akaike information criteria). latitude had a significant effect on sghs’ nesting success, with nests in the north being less successful than those in the south. an increase in amount of thicket and dense bush, grassland or bare ground surrounding the nest caused nesting success to decrease. similarly, the greater the percentage of gabbro soils within the buffer zone around each nest, the lower the overall group nesting success. nests with a higher diversity of habitats within the buffer zone had a higher nesting success than those with more homogenous surrounding habitat. the proximity of nests to roads improved overall nesting success, with more successful nests being situated closer to road networks (table 3). ethical considerations this work was part of a registered research project with sanparks (pottl988) and was approved by the relevant animal ethics committee. discussion in this study, we analysed data spanning seven breeding seasons to determine what factors contribute to sgh nesting success in the kruger national park. in particular, we investigated whether harvesting of second-hatched redundant chicks impacts on the survival to fledging of first-hatched chicks. our results showed that this practice had no detrimental effects and could even benefit the first-hatched chick in some ways (although this potential benefit was not significant). this was an important finding for the conservation of sgh, as harvesting of second-hatched chicks is crucial to the established captive-rearing programme and the subsequent formulation of groups for future reintroductions. we found that the habitat features in the area immediately surrounding sgh nest sites will greatly impact their nesting success. shifts in habitat structure can often result in changes in species assemblages which may in turn impact the reproductive output of the associated bird species through loss of potential nesting habitat (martin 2014), changes in prey base (burke & nol 1998), changes in foraging ability (butler & gillings 2004) and increased predation risk (badyaev 1995; haensly, crawford & meyers 1987; yurizharikov & cooke 2007). for territorial species, which are already restricted to finding nest sites and sufficient prey within their territory, these impacts can be exacerbated, as changes in prey availability or accessibility or the availability of suitable nesting sites within their territory will lead to a reduction in productivity. sghs require a territory with delicate balance between more open grassy areas for foraging and woodland areas for nesting. the dominance or increase in density of any one of the vegetation types (thicket and dense bush, grassland, low shrubland and bare ground) around the nest site will negatively affect sgh nesting success. dense grass was associated with both vegetation types found on gabbro soils (venter 1986), another factor found to negatively influence nesting success. the diversity index results supported these findings in that sgh groups with a greater diversity of vegetation types surrounding the nest had a significantly greater nesting success than those with more homogenous vegetation around the nest site. one interesting result was the increase in nesting success with an increase in the proximity of a nest site to the road. this was the only feature of the actual nest, in terms of characteristics, that showed any influence on sgh nesting success. this result may again reinforce the positive influence of a diversity of habitats, as road verges are often more vegetated as a result of increased precipitation runoff (smit & asner 2012), which could make these good foraging areas. some sgh groups in the kruger national park beg from vehicles, mostly during the winter months (combrink pers. obs.). in winter, when the abundance of available prey is at its lowest, sghs are known to concentrate their foraging in areas around sources where prey abundance is higher (kemp et al. 1989). thus, a steady food source associated with roads in the winter months could increase the health and fitness of the alpha pair and thereby increase their nesting success. sghs spend around 70% of the day walking (kemp 1995), and have been known to cover distances of around 7 km in a day (wilson & hockey 2013). knight (1990) showed that sghs favour areas where the grass is less than 50 cm in height. a change in habitat structure, such as an increase in grassland, and thicket or dense bush would likely impact sgh foraging efficiency. not only would prey detection and acquisition be more difficult, but also the denser habitat structure could increase the risk of ambush by predators (butler & gillings 2004; wilson & hockey 2013). over the past century, there has been an increase in woody plants in grasslands and savannas (buitenwerf et al. 2012; o’connor, puttick & hoffman 2014; wigley et al. 2010). this woody or bush encroachment can significantly alter the biodiversity in the area (buitenwerf et al. 2012) and have large consequences for conservation and protected areas (wigley et al. 2010). bush encroachment is often considered the result of changes in disturbance regimes (buitenwerf et al. 2012; parr et al. 2014; wigley et al. 2010). disturbance, such as fire and herbivory (with african elephants loxodonta africana in particular), is known to engineer savanna vegetation (o’connor et al. 2014; sankaran, ratnam & hanan 2008). with the increasing elephant densities in the kruger national park (young et al. 2009) and the occurrence of frequent fires, many trees will have difficulty establishing (helm et al. 2011). without recruitment, the loss of woody species and canopy trees can significantly alter the habitat composition with a potential change in habitat structure through bush encroachment (baxter & getz 2005; smallie & o’connor 2000) from more open savannas to dense woodland (hibbard et al. 2001; parr et al. 2014). in addition, the loss of tall canopy trees in the landscape will reduce the number of potential nesting trees for sgh and many other large tree nesting raptors, such as white-backed vultures gyps africanus (murn et al. 2013) and martial eagles polemaetus bellicosus (van eeden et al. 2017). nest sites for sghs are thought to be limited in the kruger national park (kemp et al. 1989) owing to a scarcity of suitably sized cavities. the loss of a sgh nest site within a territory could be detrimental to the group’s productivity. following nest loss or collapse, some groups in the kruger national park have failed to breed for the remainder of our study period – in one case, for a total of four seasons. wilson and hockey (2013) found sgh nesting success for natural nests to be significantly correlated with the amount of open woodland within 3 km of the nest site. in our models, open woodland and grassland habitat types were negatively correlated. the grassland habitat type was retained in our final land model, as of the two habitat types, it produced the model with the lowest aic value. we found that an increase in the grassland habitat type had a negative effect on nesting success. therefore, we can deduce that an increase in open woodland around the nest will be beneficial to sgh. this means that the amount of open woodland surrounding the nest site is an important factor to consider when deciding on the placement of nest boxes or when creating nest sites for sghs, regardless of the vegetation type in which the nest is located. authorities responsible for vegetation management should take into account the possible influence that management decisions and actions can have on habitat composition and structure and the influence this has on overall biodiversity (skowno & bond 2003). the influence of climate change on bush encroachment into grasslands (wigley et al. 2010) and the subsequent changes to the structure of savanna vegetation (buitenwerf et al. 2012; parr et al. 2014) cannot be actively controlled. however, management authorities can control and even mitigate other drivers, such as the influence of frequent fires and elephants. in the kruger national park, where the elephant population is seemingly on the increase (young et al. 2009), the potential impact of high densities of elephants on the vegetation structure and subsequent impacts on overall species diversity should not be ignored. elephant impacts on large trees in african savannas should therefore be monitored and managed where necessary to allow for the establishment of sufficient large trees. similarly, in areas where sgh nests are located, and in particular, those known to be successful, fires should be managed so as not to damage the large nest trees and alter the habitat structure towards a more homogenous surrounding habitat. conclusion our data support the harvesting of second-hatched chicks as a viable conservation initiative, as set out in the sgh species recovery plan (jordan 2011). removal of second-hatched chicks from sgh nests was found to have no significant effect on the survival to fledging of the first-hatched chick. we therefore recommend that harvesting of second-hatched chicks from wild sgh nests continues to support the captive rearing and reintroduction programme (jordan 2011). our results showed that habitat structure and diversity are critical when deciding on a suitable reintroduction site for sghs. it is also imperative that nest site placement maximise the amount of open woodland and a diversity of habitats surrounding the nest. annual monitoring of sgh nest sites will allow for the early detection and possible mitigation of bush encroachment or changes in habitat structure surrounding nest sites. in addition, monitoring would detect when natural nest sites collapse or are no longer suitable. maintenance of these natural nest sites and the erection of artificial nest boxes, in cases where natural nests collapse, should be considered as a conservation intervention. sghs readily take to nest boxes and groups nesting in artificial nests have been shown to have a significantly higher breeding success when compared with groups using natural nest sites (wilson & hockey 2013). sghs, as with many other large bird species, are considered to be safe within protected areas, with most of the threats to the birds affecting those populations occurring in unprotected areas. we have shown that even within protected areas, without sufficient management interventions to control the drivers of bush encroachment and nest site losses, the nesting success and population status of sghs, and potentially all large tree canopy nesting birds, will decline. acknowledgements we thank sanparks for allowing access to the kruger national park and providing the relevant data and gis layers. we would also like to thank the numerous staff members, researchers and volunteers who assisted with field work over the years. we thank the ford wildlife foundation and the first rand foundation for providing financial support for this research. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions l.c. and c.t.d. conceptualised the article. l.c., h.j.c. and a.j.b. conducted the field work. l.c. and h.j.c. analysed the data. l.c. wrote the article. all authors provided editorial input for the article. references badyaev, a.v., 1995, ‘nesting habitat and nesting success of eastern wild turkeys in the arkansas ozark highlands’, condor 97, 221–232. https://doi.org/10.2307/1368998 bates, d., maechler, m., bolker, b. & walker, s., 2015, ‘fitting linear mixed-effects models using lme4’, journal of statistical software 67, 1–48. https://doi.org/10.18637/jss.v067.i01 baxter, p.w.j. & getz, w.m., 2005, ‘a model-framed evaluation of elephant effects on tree and fire dynamics in african savannas’, ecological applications 15, 1331–1341. https://doi.org/10.1890/02-5382 birdlife international, 2016, ‘species factsheet: bucorvus leadbeateri’, viewed 18 april 2016, from http://www.birdlife.org/datazone/speciesfactsheet.php?id=983 blackmer, a.l., ackerman, j.t. & nevitt, g.a., 2004, ‘effects of investigator disturbance on hatching success and nest-site fidelity in a long-lived seabird, leach’s storm-petrel’, biological conservation 116, 141–148. https://doi.org/10.1016/s0006-3207(03)00185-x boulton, r.l., richard, y. & armstrong, d.p., 2008, ‘influence of food availability, predator density and forest fragmentation on nest survival of new zealand robins’, biological conservation 14, 580–589. https://doi.org/10.1016/j.biocon.2007.12.007 buitenwerf, r., bond, w.j., stevens, n. & trollope, w.s.w., 2012, ‘increased tree densities in south african savannas: >50 years of data suggests co2 as a driver’, global change biology 18, 675–684. https://doi.org/10.1111/j.1365-2486.2011.02561.x burke, d.m. & nol, e., 1998, ‘influence of food abundance, nest-site habitat and forest fragmentation on breeding ovenbirds’, auk 115, 96–104. https://doi.org/10.2307/4089115 butler, s.j. & gillings, s., 2004, ‘quantifying the effects of habitat structure on prey detectability an accessibility to farmland birds’, ibis 146, 123–130. https://doi.org/10.1111/j.1474-919x.2004.00352.x didan, k., 2015, mod13q1 modis/terra vegetation indices 16-day l3 global 250m sin grid v006, nasa eosdis land processes daac. https://doi.org/10.5067/modis/mod13q1.006 dinsmore, s.j., white, g.c. & knopp, f.l., 2002, ‘advanced techniques for modelling avian nest survival’, ecology 83, 1476–1488. https://doi.org/10.1890/0012-9658(2002)083[3476:atfman]2.0.co;2 geoterraimage, 2014, 2013–2014 south african national land-cover dataset, data user report and metadata, february 2015, version 05, geoterra image (gti) pty ltd, south africa. gertenbach, w.p.d., 1980, ‘rainfall patterns in the kruger national park’, koedoe 23, 35–43. https://doi.org/10.4102/koedoe.v23i1.634 gertenbach, w.p.d., 1983, ‘landscapes of the kruger national park’, koedoe 26, 9–122. https://doi.org/10.4102/koedoe.v26i1.591 haensly, t.f., crawford, j.a. & meyers, m., 1987, ‘relationships of habitat structure to nest success of ring-necked pheasants’, journal of wildlife management 52, 421–425. https://doi.org/10.2307/3801029 helm, c., wilson, g., midgley, j., kruger, l. & witowski, e.t.f., 2011, ‘investigating the vulnerability of an african savanna tree (sclerocarya birrea spp. affra) to fire and herbivory’, austral ecology 36, 964–973. https://doi.org/10.1111/j.1442-9993.2010.02232.x hibbard, k.a., archer, s., schimel, d.s. & valentine, d.a., 2001, ‘biochemical changes accompanying woody plant encroachment in a subtropical savannah’, ecology 82, 1999–2011. https://doi.org/10.1890/0012-9658(2001)082[1999:bcawpe]2.0.co;2 jordan, m. (ed.), 2011, southern ground hornbill (bucorvus leadbeateri) species recovery plan for south africa, conservation breeding specialist group (ssc/iucn)/cbsg south africa/johannesburg zoo/endangered wildlife trust, johannesburg. kemp, a.c., 1988, ‘the behavioural ecology of the southern ground hornbill: are competitive offspring at a premium?’, in r. van den elzen, k.-l. schuchmann & k. schmidt-koenig (eds.), proceedings of the international centennial meeting of the deutsche ornithologen-gesellschaft: current topics in avian biology, pp. 261–271, 1988, bonn, germany. kemp, a.c., 1995, the hornbills, oxford university press, oxford. kemp, a.c. & begg, k.s., 1996, ‘nest sites of the southern ground hornbill bucorvus leadbeateri in the kruger national park, south africa, and conservation implications’, ostrich 67, 9–14. https://doi.org/10.1080/00306525.1996.9633773 kemp, a.c., joubert, s.c.j. & kemp, m.i., 1989, ‘distribution of southern ground-hornbills in the kruger national park in relation to some environmental features’, south african journal of wildlife research 19, 93–98. kemp, a.c. & kemp, m.i., 1991, ‘timing of egg laying by southern ground-hornbills bucorvus leadbeateri in the central kruger national park, south africa’, ostrich 62, 80–82. knight, g.m., 1990, ‘status, distribution and foraging ecology of the southern ground hornbill (bucorvus leadbeateri)’, in natal’, msc thesis, university of natal, pietermaritzburg. loegering, j.p. & anthony, r.g., 2006, ‘nest-site selection and productivity of american dippers in the oregon coast range’, the wilson journal of ornithology 111, 281–438. https://doi.org/10.1676/04-100.1 martin, t.e., 2014, ‘consequences of habitat change and resource selection specialization for population limitation in cavity-nesting birds’, journal of applied ecology 52, 475–485. https://doi.org/10.1111/1365-2664.12375 murn, c., combrink, l., ronaldson, g.s., thompson, c., & botha, a.j., 2013, ‘population estimates of three vulture species in kruger national park, south africa’, ostrich 84, 1–9. https://doi.org/10.2989/00306525.2012.757253 o’connor, t.g., puttick, j.r. & hoffman, m.t., 2014, ‘bush encroachment in southern africa: changes and causes’, african journal of range and forage science 31, 67–88. https://doi.org/10.2989/10220119.2014.939996 oksanen, j., blanchet, f.g., kindt, r., legendre, p., minchin, p.r., o’hara, r.b. et al., 2016, ‘vegan: community ecology package, r package version 2.3-3’, viewed 15 march 2016, from http://cran.r-project.org/package=vegan oro, d., pradel, r. & lebreton, j-d., 1999, ‘food availability and nest predation influence life history traits in audouin’s gull, larus audouinii’, oecologia 118, 438–445. https://doi.org/10.1007/s004420050746 paredes, r. & zavalaga, c.b., 2001, ‘nesting sites and nest types as important factors for the conservation of humboldt penguins (sphensicus humboldti)’, biological conservation 100, 199–205. https://doi.org/10.1016/s0006-3207(01)00023-4 parr, c.l., lehmann, c.e.r., bond, w.j., hoffman, w.a. & andersen, a.n., 2014, ‘tropical grassy biomes: misunderstood, neglected, and under threat’, trends in ecology and evolution 29, 205–213. https://doi.org/10.1016/j.tree.2014.02.004 pettorelli, n., ryan, s., mueller, t., bunnefeld, n., jedrzejewska, b., lima, m. & kausrud, k., 2011, ‘the normalised difference vegetation index (ndvi): unforeseen successes in animal ecology’, climate research 46, 15–27. https://doi.org/10.3354/cr00936 pike, n., 2011, ‘using false discovery rates for multiple comparisons in ecology and evolution’, methods in ecology and evolution 2, 278–282. https://doi.org/10.1111/j.2041-210x.2010.00061.x rastogi, a.d., zanette, l. & clinchy, m., 2006, ‘food availability affects diurnal nest predation and adult antipredator behaviour in song sparrows, melospiza melodia’, animal behaviour 72, 933–940. https://doi.org/10.1016/j.anbehav.2006.03.006 r core team, 2015, r: a language and environment for statistical computing, r foundation for statistical computing, vienna, austria, viewed 02 february 2015, from https://www.r-project.org reidy, j.l., thompson, f.r. iii & peak, r.g., 2009, ‘factors affecting golden-cheeked warbler nest survival in urban and rural landscapes’, journal of wildlife management 73, 407–413. https://doi.org/10.2193/2007-516 robinson, w.d., robinson, t.r., robinson, s.k. & brawn, j.d., 2000, ‘nesting success of understory forest birds in central panama’, journal of avian biology 31, 151–164. https://doi.org/10.1034/j.1600-048x.2000.310207.x sankaran, m., ratnam, j. & hanan, n.p., 2008, ‘woody cover in african savannas: the role of resources, fire and herbivory’, global ecology and biogeography 17, 236–245. https://doi.org/10.1111/j.1466-8238.2007.00360.xs shannon, c.e. & weaver, w., 1949, the mathematical theory of communication, university of illinois press, urbana. skowno, a.l. & bond, w.j., 2003, ‘bird community composition in an actively managed savanna reserve, importance of vegetation structure and vegetation composition’, biodiversity and conservation 12, 2279–2294. https://doi.org/10.1023/a:1024545531463 smallie, j.j. & o’connor, t.g., 2000, ‘elephant utilization of colophospermum mopane: possible benefits of hedging’, african journal of ecology 38, 1–9. https://doi.org/10.1046/j.1365-2028.2000.00258.x smit, i.p.j. & asner, g.p., 2012, ‘roads increase woody cover under varying geological, rainfall and fire regimes in african savannah’, journal of arid environments 80, 74–80. https://doi.org/10.1016/j.jaridenv.2011.11.026 spellerberg, i.f. & fedor, p.j., 2003, ‘a tribute to claude shannon (1916 –2001) and a plea for more rigorous use of species richness, species diversity and the ‘shannon–wiener’ index’, global ecology and biogeography 12, 177–179. https://doi.org/10.1046/j.1466-822x.2003.00015.xs taylor, m.r., peacock, f. & wanless, r.m., 2015, the eskom red data book of birds of south africa, lesotho and swaziland, birdlife south africa, johannesburg. tuck, s. & phillips, h., 2015, ‘modistools: modis subsetting tools’, r package version 0.94.6, viewed 29 april 2015, from http://cran.r-project.org/package=modistools van eeden, r., whitfield, d.p., botha, a. & amar, a., 2017, ‘ranging behaviour and habitat preferences of the martial eagle: implications for the conservation of a declining apex predator’, plos one 12:e0173956. https://doi.org/10.1371/journal.pone.0173956 van wilgen, b.w., govender, n. & macfadyen, s., 2008, ‘an assessment of the implementation and outcomes of recent changes to fire management in the kruger national park’, koedoe 50, 22–31. https://doi.org/10.4102/koedoe.v50i1.135 venter, f.j., 1986, ‘soil patterns associated with the major geological units of the kruger national park’, koedoe 29, 125–138. https://doi.org/10.4102/koedoe.v29i1.525 venter, f.j., 1990, ‘a classification of land for management planning in the kruger national park’, ph.d. thesis, university of south africa, pretoria. vickery, p.d., hunter, m.l., jr. & wells, j.v., 1992, ‘use of a new reproductive index to evaluate relationship between habitat quality and breeding success’, auk 109, 697–705. https://doi.org/10.2307/4088145 wessels, k.j., prince, s.d., zambatis, n., macfadyen, s., frost, p.e. & van zyl, d., 2006, ‘relationship between herbaceous biomass and 1-km2 advanced very high resolution readiometer (avhrr) ndvi in kruger national park, south africa’, international journal of remote sensing 27, 951–973. https://doi.org/10.1080/01431160500169098 wigley, b.j., bond, w.j. & hoffman, t., 2010, ‘thicket expansion in a south african savanna under divergent land use: local vs. global drivers?’, global change biology 16, 964–976. https://doi.org/10.1111/j.1365-2486.2009.02030.x wilson, g. & hockey, p.a.r., 2013, ‘causes of variable reproductive performance by southern ground-hornbill bucorvus leadbeateri and implications for management’, ibis 155, 476–484. https://doi.org/10.1111/ibi.12042 winter, m., johnson, d.h., shaffer, j.a., donovan, t.m. & svedarsky, w.d., 2006, ‘patch size and landscape effects on density and nesting success of grassland birds’, usgs northern prairie wildlife research center paper 235, http://digitalcommons.unl.edu/usgsnpwrc/235 woolaver, l.g., nichols, r.k., morton, e.s. & stutchbury, b.j.m., 2014, ‘breeding ecology and predictors of nest success in the critically endangered ridgway’s hawk buteo ridgwayii’, bird conservation international 25, 385–398. https://doi.org/10.1017/s0959270914000318 young, k.d., ferreira, s.m. & van aarde, r.j., 2009, ‘the influence of increasing population size and vegetation productivity on elephant distribution in the kruger national park’, austral ecology 34, 329–342. https://doi.org/10.1111/j.1442-9993.2009.01934.x yurizharikov, d.b.l. & cooke, f., 2007, ‘influence of landscape pattern on breeding distribution and success in a threatened alcid, the marbled murrelet: model transferability and management implications’, journal of applied ecology 44, 748–759. https://doi.org/10.1111/j.1365-2664.2007.01319.x zarones, l., sussman, a., morton, j.m., plentovich, s., faegre, s., aguon, c. et al., 2014, ‘population status and nest success of the critically endangered mariana crow corvus kubaryi on rota, northern mariana islands’, bird conservation international 25, 220–233. https://doi.org/10.1017/s0959270914000045 article information authors: nicholas theron1,2 raymond jansen3 paul grobler1 antoinette kotze1,4 affiliations: 1department of genetics, university of the free state, south africa2mabula ground hornbill research and conservation project, bela-bela, south africa 3department of environmental, water and earth sciences, tshwane university of technology, south africa 4national zoological gardens of south africa, pretoria, south africa correspondence to: nicholas theron postal address: po box 864, underberg 3257, south africa dates: received: 09 jan. 2013 accepted: 08 aug. 2013 published: 11 oct. 2013 how to cite this article: theron, n., jansen, r., grobler, p. & kotze, a., 2013, ‘the home range of a recently established group of southern ground-hornbill (bucorvus leadbeateri) in the limpopo valley, south africa’, koedoe 55(1), art. #1135, 8 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1135 copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. the home range of a recently established group of southern ground-hornbill (bucorvus leadbeateri) in the limpopo valley, south africa in this original research... open access • abstract • introduction • research method and design    • study area    • study sites    • vegetation sampling    • invertebrate sampling    • home range analysis    • data analysis • results    • vegetation    • invertebrates    • home range size and habitat utilisation • ethical considerations • discussion    • seasonal abundance of invertebrates    • seasonal home range size and habitat utilisation    • southern ground-hornbill territory size    • southern ground-hornbill nesting requirements • conclusion • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ little is known about southern ground-hornbill (sgh) population ecology outside of large, formally protected areas where the largest declines in numbers have been recorded. the sgh has started re-colonising, establishing group territories and breeding successfully in the limpopo valley on the northern border of south africa, following localised extinction from the 1950s to the 1970s. a group of sgh was monitored over a period of 14 months by means of radio telemetry across privately owned land in order to investigate their seasonal habitat movements in this semi-arid, predominantly livestock-based environment. we also investigated seasonal fluctuations in invertebrate prevalence, as an indication of food availability and its influence on seasonal sgh group movements and foraging activity patterns. there was a clear increase in food availability during the summer rainfall period allowing the group to forage over a wider area, whilst winter foraging remained localised within their range. kernel home range analysis indicated a marked difference in size between the summer (13 409 ha) and winter (5280 ha) home ranges, with an overall home range of 19 372 ha, which is approximately double that of home ranges recorded that fall within formally and informally protected reserves. in this article, we proposed that food availability is the driving force for home range size and seasonal activity patterns in a semi-arid livestock-ranching habitat. conservation implications: the limpopo valley sgh population is one of the most significant outside protected areas in south africa. this population is especially vulnerable to threats such as poisoning, persecution for window breaking and drought, as shown by their near extirpation from the area. conservation efforts need to focus on awareness amongst local farmers, provision of artificial nests and continued monitoring of groups. introduction top ↑ the southern ground-hornbill (sgh) (bucorvus leadbeateri) is the largest hornbill in the world and the largest cooperatively breeding bird species (kemp & kemp 1980). it is a territorial, sedentary species and occurs in groups averaging 3–4 individuals consisting of an alpha pair and a mixture of juveniles and adult helpers (kemp & kemp 1980; knight 1990). the species is officially listed as vulnerable in south africa (kemp 2000), whilst a recent review of the international union for conservation of nature (iucn) global status has precipitated a change in the species’ status from least concern to vulnerable by birdlife international (2010). the south african population is estimated at between 1500 and 2000 individuals, with almost half of this population (600–700 individuals) found in the kruger national park (knp) (kemp 2005). the reduction in numbers has primarily been associated with habitat degradation, persecution for window breaking, secondary poisoning and the loss of suitable large trees for nesting. in order to reverse these negative population trends, a critical component of the species’ conservation strategy will be to understand what constitutes ideal sgh habitat. suitability of habitat can be seen to contribute to the overall fitness and survival of an individual (block & brennan 1993) and the resources and conditions necessary to allow an organism to survive, reproduce and persist (hall, krausman & morrison 1997).according to kemp (1988), nesting sites are the primary resource associated with sgh territories, whilst food resources are secondary. data from ground and aerial counts in the knp spanning 20 years indicated that there exists no relationship between the frequency groups were encountered and group size with the distribution of rainfall, geology, geomorphology, drainage lines, soil types, vegetation types or vegetation structure (kemp, joubert & kemp 1989). the importance of food resources to the species in african savannas is shown by the influence it has on sgh behaviour. during periods of lowest food abundance in winter, sgh adapt their behaviour by concentrating in areas around waterholes that have higher densities of ungulates and where food abundance will be higher (kemp et al. 1989). furthermore, observations indicate sgh dig more during the drier months of the year when surface prey is less abundant, often in and around piles of elephant dung (kemp 1995) and in rhino middens. the onset of breeding is also restricted to the summer months (october–march) and is triggered by the start of the summer rains. it is this summer rainfall that is believed to be the proximate factor influencing the seasonal availability of food for the species, primarily invertebrates (kemp 1976; kemp & kemp 1991) which form an important component of sgh diet during the wetter summer months (kemp & kemp 1978; knight 1990). in south africa, densities of one group per 100 km² are recorded in the knp (kemp & kemp 1980), kwazulu-natal midlands (knight 1990) and the eastern cape (vernon 1984), whilst observations indicate densities of approximately one group per 20 km² in the mana pools region of zimbabwe (kemp 2005) and the masai mara in kenya (n. theron, pers. obs.). southern ground-hornbills disappeared from the limpopo valley during the 1950s–1970s (pers. comm. land owners), likely the result of extreme drought conditions and overgrazing aggravated by threats such as poisoning, electrocutions and persecution for window breaking. re-colonisation since the early 1990s has subsequently occurred in the region with the establishment of successful breeding groups. only 10 groups are known north of the soutpansberg mountains, between the platjan border post and the towns of alldays and musina, with an estimated density of approximately one group per 600 km² (theron et al. 2013). one group, which is the focus of this study, has been present in the area since the mid-1990s and has bred in an adansonia digitata (baobab tree) at least twice from the summer of 2008 until the summer of 2011 (n. theron, pers. obs.). home range, habitat use, foraging behaviour and breeding success of sgh outside formally protected areas has not been investigated. furthermore, behavioural studies on re-emerging sgh populations have yet to be undertaken. this is especially important in terms of developing a management plan for the conservation of this threatened species particularly for those areas they historically occupied. moreover, quantitative seasonal data on the general availability of prey resources that include invertebrate abundance and the influences this has on foraging behaviour patterns of sgh, especially during the winter months (march–august), are still poorly understood. this study is a first attempt at quantifying the seasonal movement patterns of a group of sgh within a predominantly agricultural landscape outside of a protected area. furthermore, this study aims to investigate the seasonal availability of invertebrates in relation to rainfall, temperature and changes in vegetation cover and the possible consequence this may have on the habitat utilisation, movement and seasonal home range size of a group of sgh in the limpopo valley. research method and design top ↑ study area the study was undertaken from july 2008 to october 2009 in the limpopo valley, a semi-arid landscape that forms part of the savanna biome at an altitude ranging between 300 m.a.s.l. and 1000 m.a.s.l. (mucina & rutherford 2006) on the northern border of south africa with zimbabwe (figure 1) at the north-western extent of the sgh’s range in south africa. the main vegetation types, classified by mucina and rutherford (2006), include musina mopane bushveld making up the majority of the plains and limpopo ridge bushveld covering the hills and ridges scattered throughout the area. colophospermum mopane (mopani) trees, broad-leaved deciduous species such as a. digitata, terminalia prunioides (lowveld cluster leaf) and various species of commiphora dominate the area (n. theron, pers. obs.). the limpopo valley is a low rainfall area, with a yearly average of 341.6 mm (jordaan et al. 2004) and a mean monthly temperature ranging between 15.5 °c and 29.9 °c. rainfall is seasonal and sporadic, with 79.7% of the total precipitation occurring in summer (data from 1980–2009 supplied on request by the south african weather service). commercial cattle and game ranching dominate the land-use, with an estimated 3.0% of musina mopane bushveld and 1.0% of limpopo ridge bushveld transformed mainly for agriculture (mucina & rutherford 2006). the area is further characterised by low average rainfall, shallow nutrient poor and severely eroded soils with low moisture retention (jordaan et al. 2004). figure 1: map of the study area in south africa (inset) with a detailed map of the limpopo valley and the location of stoke safaris where invertebrate and vegetation data were collected. study sites four study sites within the group’s territory were randomly chosen on the stokes farm (242 ms; 22°28′s, 29°52′e) to collect vegetation and invertebrate data from october 2008 to september 2009. these study sites were targeted because of the constant presence of sgh on the farm over a number of months. sites 2 and 3 were historically grazed by cattle and are now stocked with indigenous game species. site 1 is currently a breeding camp for nyala (tragelaphus angasii) and sable antelope (hippotragus niger). site 4 is old agricultural land that was abandoned more than 15 years ago. vegetation sampling a visual estimate of cover to describe temporal changes in vegetation was conducted monthly at 10 geo-referenced points per site. canopy cover of grass species formed the focus of this estimate by using a 1 m² metal quadrant at each point and the percentage grass cover estimated according to the following categories: < 10%, 10% – 25%, 25% – 50%, 50% – 75% and 75% – 100%. each category was allocated a score, with one being the lowest and five the highest, to compile a monthly score ranging between 10 and 50. the overall condition score for each replicate was ascribed by those developed for this habitat by jordaan et al. (2004), where three categories were described depending on seasonal precipitation levels, namely: extremely bad (bare soil and forbs dominate), bad (annual grasses such as aristida spp. dominate) and good (annual grasses such as aristida spp. and perennial grasses such as eragrostis lehmanniana dominate). invertebrate sampling both pitfall and sweep net methods were employed to target invertebrates that would mostly be encountered by a sghs mode of foraging behaviour. pitfall samples would include species that are mainly ground-dwelling, whilst sweep netting would sample invertebrates occurring predominantly on surface vegetation (standen 2000). pitfall traps and sweep netting methods were employed monthly from october 2008 to september 2009 to determine seasonal availability at the same geo-referenced site the vegetation was sampled. at each site, 13 pitfall traps were set out using the nested cross array method (perner & schueler 2004). plastic tubes with a 10 cm diameter and 15 cm in length were buried flush with the soil surface and cups dropped into each and filled with propylene glycol up to 3 cm deep. propylene glycol is non-toxic and safe for animals that may feed on invertebrates caught in traps. pitfalls were left out for a sampling period of 4 days each month. pitfalls disturbed by animals were not included in the analysis as selected invertebrates may have been removed. invertebrates were washed and stored in polytop vials with a 75% ethanol solution for later identification up to order level. sweep netting was performed monthly by walking the same geo-referenced 200 m transect at each of the four sites within the same week pitfalls were sampled. a 45-cm diameter net was used to take sweep net samples where 200 sweeps were performed per line transect (one sweep for each step taken) before 10 am. all invertebrates were identified up to the level of order, counted and measured volumetrically using the volumetric water displacement method (jansen & crowe 2006). home range analysis a group of five sgh consisting of an alpha male, alpha female, one sub-adult male and female and one juvenile was captured by luring them into a netting trap with a fibreglass sgh model and a vocal territorial recording. a holohil® tail transmitter was fitted onto the main tail deck-feather of the alpha female. after a 30-day settling period, the group was tracked using a handheld yagi antennae and an aor® receiver. locality data were collected every 3–4 h where possible, including roost sites, 5 days a month from august 2008 to september 2009. nest sites were identified in order to assess their possible influence on the movement of groups in the landscape. gps readings were recorded using a garmin® gps. other factors such as the influence of human structures (buildings, roads and highways) on the group’s movements were also observed and recorded. data analysis ranges vii software (south, kenward & walls 2008) was used to analyse home range data where the harmonic mean and kernel home range calculations were used in estimating seasonal home range size. harmonic mean home range estimates are highly sensitive to outlying observations and thus force the inclusion of many grid points. as such, the outcome of the home range size is an overestimate of true size, whereas kernel estimators are well defined and tractable (seaman & powell 1996) and are presented in this study. furthermore, gps co-ordinates were loaded into qgis 1.8.0 lisboa (open source geospatial foundation project) and overlayed on 1:10 000 high resolution orthophotos and spatial layers representing rivers, roads and vegetation types to further note if any associations exists with group movements and structural habitat features. statistical data analysis was undertaken using the software program statistica (2009). kruskal-wallis one way analysis of variance (anova) was used to test for any variations between sites with regards to invertebrate prevalence and vegetation cover. a correlation analyses was performed (spearman’s) to determine if there exists any association between invertebrate numbers and grass cover. correlations were performed against invertebrate data and relevant meteorological data (rainfall as well as maximum and minimum temperatures) where p < 0.05 (95%) denotes significance. a two tailed t-test was further employed to test whether there was any significant difference between invertebrate sampling methods. results top ↑ vegetation monthly estimates of cover were very similar across all four sites (anova: kw = 2.12, df = 3, p = 0.5470) and the average overall monthly score for all sites was very low at 17.7, representing an average cover of between 10.0% and 25.0%. the veld condition using the jordaan et al. (2004) criteria ranks each site as extremely bad throughout the study period. graphical representation of grass cover and rainfall revealed the close relationship between precipitation and the response of vegetation growth (figure 2). grass growth was only stimulated following the december–january rains (272.6 mm), resulting in the highest grass cover scores during february. the total annual rainfall during the study was 40% above average at 477.7 mm, of which 460 mm (94.6%) fell during summer (november–march). figure 2: seasonal comparison of rainfall and grass cover scores. invertebrates the two sampling methods differed significantly with the pitfall traps being the most successful (t = 6.25, df = 21, p < 0.001). sweep netting yielded a total of 1193 individuals in comparison to the pitfalls with a total of 18 272 (table 1). the pitfall data showed positive correlations between the number of invertebrates with both mean monthly maximum temperatures (r2 = 0.531, p < 0.05) and minimum temperatures (r2 = 0.612, p < 0.05) and the volume of invertebrates with mean monthly rainfall (r2 = 0.563, p < 0.05). no associations were found between meteorological variables and sweep net data. as such, sweep net data proved unreliable and pitfall data seemed a more appropriate method of collecting invertebrates in this very arid environment. all statistical procedures made use of pitfall invertebrate data and sweep net data was not used. invertebrate numbers and volume peaked during summer (november–april) and were lowest in mid-winter (may–july; table 1). there were no significant differences between sites with regards to the number or volume of invertebrates sampled using pitfall traps (anova: kw = 0.539, df = 3, p = 0.910). furthermore, no association was found between invertebrate numbers (prevalence) and relative grass cover (spearman r = 0.07, p = 0.834). table 1: total numbers and volume of invertebrates captured from october 2008 to september 2009 using two sampling methods. no invertebrates were captured using sweep nets during the months of october and december differences between the compositions of orders between the two sampling methods were apparent (figures 3a–3d). coleopterans dominated pitfall sampling and made up 62% of the numbers and 76% of the total volume. orthopterans were the most abundant order sampled with sweep nets for both numbers and volume, representing 62% and 76% respectively. orthoptera were the second-most abundant order in terms of pitfall volume, making up 6% of the total volume. other notable differences in the most dominant orders collected for the two sampling methods were hemiptera, which represent a significant proportion of the invertebrates captured by sweep netting but not in pitfall traps for both numbers and volume, whilst the converse was true for the order of hymenoptera; strongly represented in the pitfalls but not the sweep net sample set. figure 3: comparison of the number and volume (determined using the volumetric water displacement method) of invertebrate orders captured by pitfall and sweep net sampling at four sites and expressed as a percentage according to, (a) numbers of invertebrates from pitfall traps categorised per order, (b) volume of invertebrates from pitfall traps categorised per order, (c) numbers of invertebrates from sweep nets categorised per order and (d) volume of invertebrates from sweep nets categorised per order. home range size and habitat utilisation a total of 201 fixes were recorded. the territory size of the group varied between 19 372 ha and 22 731 ha by the kernel and harmonic mean home range analysis, respectively. the results of the kernel home range indicated a summer (september–february) and winter (march–august) size of 13 409 ha (figure 4a) and 5280 ha (figure 4b), respectively. although the group would generally avoid areas with human activity, roads and highways did not seem to restrict movement of the group as these were easily traversed, often regularly. a total of three baobab trees with suitable nesting cavities were recorded, with the female spending at least one day in each nest. the group did not breed during the study period and the female was never observed in the primary nest tree which was used historically by the group. the group subsequently bred in this nest during the 2009–2010 season. finally, this study highlighted the limitations of using radio telemetry to follow sgh groups in this challenging environment where farms are all privately owned and acquiring permission to access a property can be difficult. ideally, a number of emerging sgh groups should be fitted with satellite transmitters to suitably quantify home range and habitat use in this environment. figure 4: the (a) summer and (b) winter seasonal kernel home ranges of a southern ground-hornbill group during the study period in relation to environmental features. contour lines indicate kernel mean analyses using 95% fixes at 5% intervals. ethical considerations top ↑ the project was approved by the ethics and scientific committee of the national zoological gardens of south africa. all relevant permits and permission from landowners were acquired before capture of the sgh group and during fieldwork. discussion top ↑ seasonal abundance of invertebrates this study reveals positive correlations between insect abundance and temperature and rainfall across seasons. this is similar to studies in other areas where wet and dry seasons alternate and insect abundance is influenced by temperature and rainfall (da silva, frizzas & de oliveira 2011; poulin, gaetan & mcneil 1992). in these variable environments, the seasonal abundance of invertebrates is related to the presence of food, which for herbivorous insects would be the flush of new leaves and grass brought about by the wet season, in turn influencing the abundance of predatory invertebrates (wolda 1979). the limpopo valley experiences a distinct rainy season from november to march with grass growth immediately responding to the onset of rains, thus stimulating invertebrate activity and abundance. an extreme case of this response is the large scale and unpredictable outbreaks of the mopani worm (lepidoptera: imbrasia belins) which is known to occur in arid habitats dominated by c. mopane (oberprieler 1986). such an outbreak occurred within the sgh territory during the study but unfortunately the dramatic increase of invertebrate biomass was not reflected in our results because the outbreak did not occur in the invertebrate sampling site. however, the spectacular influence rainfall plays in this ecosystem is reflected by the sudden rainfall-induced emergence of a species of monster tiger beetle (coleoptera: manticora sp.) resulting in the volume of insects caught in pitfalls increasing six-fold in november, the third highest volume of insects caught for any particular month over the study period. if future similar studies are undertaken, it is recommended that a more direct method of sampling mopani worms and other invertebrates is employed. seasonal home range size and habitat utilisation seasonal variations in home range size and the seasonal movements of the group are closely related to fluctuations in the surface abundance and availability of prey. the winter home range coincides with the period of lowest invertebrate abundance, is around 60% smaller than the summer home range and includes areas of core usage with large tracts of habitat where the group was not recorded. at the onset of summer in 2008, a sporadic and isolated thundershower occurred on the northern edge of the groups territory and the group immediately responded by moving to this area most likely in anticipation of the emergence of mopani moths and other invertebrates. mopani moths and mopani worms are an important food resource for birds in this harsh environment, as shown by gaston, chown and styles (1997) who found sgh feeding on mopani worms only during the fifth instar stage. indeed, the group was again found concentrating in this area during march 2009 whilst the outbreak of mopani worms (n. theron, pers. obs.) were in the fourth and fifth instar stage. although observations of the study group could not be made, it is likely that the group was feeding on the profusion of mopani worms found throughout this area. winter foraging was characterised by bouts of foraging focused within very specific locations over a number of days, with sudden, deliberate long distance movements (of up to 20 km) to other parts of the territory where localised foraging would again occur. these forays seemed to be purposeful rather than occurring in a random fashion and were likely undertaken to test other feeding areas known to the group before then returning to more productive foraging areas. during the last few months of winter (july–september), shortly prior to the first rains in both 2008 and 2009, invertebrate numbers and grass cover were at their lowest. during this time, the movement of the group was restricted at times to less than 500 m per day, which, in 2008, was a camp with a high density of ungulates. kemp et al. (1989) also recorded similar behaviour in the knp, with sgh most often observed during the drier winter months in areas associated with a high density of ungulates and resultant invertebrate prey resources found in the dung of invertebrates. furthermore, in the knp, kemp and kemp (1980) noted that during winter sgh foraging behaviour changes and the group will switch from foraging behaviour that focuses on invertebrates above the ground to strategies involving turning over dung, logs and other items as well as subterranean digging for invertebrates – even up to depths of up to 40 cm (kemp & kemp 1978). the ability of sgh to adapt and access invertebrates during winter is a key survival strategy in this harsh unpredictable habitat. it is interesting to note that species such as secretary birds (sagittarius serpentarius) and kori bustards (ardeotis kori), which have similar feeding and habitat requirements to sgh but cannot access subterranean prey, are uncommon in the area. the survival and fitness of a long-lived k-selected, territorial species such as the sgh would be dependent on the ability of the alpha pair to react to, locate and access available food resources particularly during the drier winter months when food is the most limited. southern ground-hornbill territory size the limpopo valley occurs at the south-western extent of the range of sgh and is a semi-arid environment. semi-arid environments are generally less productive because the scarcity of rainfall affects the production of vegetation (guttal & jayaprakash 2007). as such, the area may be considered marginal for sgh in terms of their habitat requirements, where sgh are recorded to prefer savanna habitats and being absent from largely treeless arid regions such as the kalahari (kemp 2005). the territory size of this group of sgh is almost double the size observed in the knp (kemp & kemp 1980) and kwazulu-natal (knight 1990) and 10 times the size reported in the mana pools region of zimbabwe (kemp 2005) and the masai mara reserve in kenya (n. theron, pers. obs.) which was calculated according to the density of groups. this corresponds with many studies that show a documented significant negative relationship between food abundance and territory size (adams 2001). the relatively drier rainfall patterns found in south africa in comparison to other parts of africa provides an explanation for the extreme variations found in sgh territory size across the species range. many studies have shown that the abundance of food and the costs of expelling intruders are important determinants of territory size (adams 2001). in this study, the effects of density and pressure from other groups are not expected to be an important factor because of the low density of sgh groups in the limpopo valley (theron et al. 2013). therefore, in the absence of density-dependant factors and with the observed sensitive interaction between rainfall, vegetation growth and invertebrate abundance, a large territory would in effect act as an insurance buffer increasing the chance that sporadic rainfall events occur in a group’s territory. in addition, large territories are required to provide sufficient food resources in a nutrient-poor environment. it is likely therefore, that in the limpopo valley food availability is the primary factor determining territory size. southern ground-hornbill nesting requirements although nests are not a limiting factor for the sgh group studied, all seven known nests utilised by groups in the limpopo valley are found in a. digitata trees and there seems to be a pattern with sgh distribution related to the presence of large a. digitata trees (n. theron, pers. obs.). this observation, however, needs to be verified by future ongoing monitoring of groups in the limpopo valley. in this semi-arid landscape, with few rivers, the conditions for the growth of suitably large trees containing nest holes is severely restricted, with a. digitata being the only common tree, not restricted to water courses, that can reach large enough dimensions for sgh nesting requirements. it is therefore plausible that the lack of a. digitata trees and the possible nesting cavities these trees may contain is the primary reason for the absence of sghs in certain parts of the limpopo valley. in the future, the provision of artificial nests in these areas may prove an effective management intervention to encourage new groups to re-colonise these areas. conclusion top ↑ owing to the sensitive nature of the musina mopane bushveld and limpopo ridge bushveld vegetation types and the unpredictable availability of food resources, a relatively large area (200 km² per group) needs to be secured in order to support a viable population of sgh. this is especially relevant when considering that the only statutorily conserved portion of the study site falls within the mapangubwe national park. little of these vegetation types have been transformed (mucina & rutherford 2006), although mining is a looming threat and this highlights the important role private landowners need to play in maintaining the ecological integrity of vast tracts of this area and the conservation of sgh habitat in south africa. moreover, long-term ecological monitoring of this population of sgh is further required to fully understand the home range and habitat requirements within the limpopo valley. of special interest is the competitive influence other groups may have as they re-colonise the area and the adjustment of territory size as population densities increase. acknowledgements top ↑ we are grateful to the farmers and landowners of the limpopo valley who granted us access to their farms, particularly the staff and management of stokes safaris. we would like to acknowledge financial and equipment support from the tshwane university of technology, the mabula ground hornbill research and conservation project, the university of the free state and the national research foundation. we are also grateful to the south african weather service for providing climate data. finally, we acknowledge the helpful inputs of three anonymous reviewers. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions n.t. (university of the free state) was responsible for project design, field work and data collection, data analyses and preparation of the manuscript. r.j. (tshwane university of technology) was the project leader and responsible for project design, statistical and home-range analyses and preparation of the manuscript. p.g. (university of the free state) contributed towards project funding, project design and preparation of the manuscript. a.k. (university of the free state) contributed towards project funding, project design and preparation of the manuscript. references top ↑ adams, e.s., 2001, ‘approaches to the study of territory size and shape’, annual review of ecology and systematics 32, 277–303. http://dx.doi.org/10.1146/annurev.ecolsys.32.081501.114034birdlife international, 2010, the birdlife data zone, viewed 01 march 2010, from http://www.birdlife.org/datazone/species block, w.m. & brennan, l.a., 1993, ‘the habitat concept in ornithology: theory and applications’, current ornithology 11, 35–91. http://dx.doi.org/10.1007/978-1-4757-9912-5_2 da silva, n.a.p., frizzaz, m.r. & de oliveira, c.m., 2011, ‘seasonality in insect abundance in the “cerrado” of goiás state, brazil’, revista brasileira de entomologia 55(1), 79–87. http://dx.doi.org/10.1590/s0085-56262011000100013 gaston, k.j., chown, s.l. & styles, c.v., 1997, ‘changing size and changing enemies: the case of the mopane worm’, acta oecologica 18(1), 21–26. http://dx.doi.org/10.1016/s1146-609x(97)80077-x guttal, v. & jayaprakash, c., 2007, ‘self-organization and productivity in semi-arid ecosystems: implications of seasonality in rainfall’, journal of theoretical biology 248, 490–500. http://dx.doi.org/10.1016/j.jtbi.2007.05.020, pmid:17645895 hall, l.s., krausman, p.r. & morrison, m.l., 1997, ‘the habitat concept and a plea for standard terminology’, wildlife society bulletin 25(1), 173–182. jansen, r. & crowe, t.m., 2006, ‘food preferences of swainson’s spurfowl, pternistis swainsonii, in a diverse agricultural landscape’, south african journal of wildlife research 36(2), 113–121. jordaan, j.j., wessels, d.c.j., dannhauser, c.s. & rootman, g.t., 2004, ‘secondary succession in the mopani veld of the limpopo valley, south africa’, african journal of range and forage science 21(3), 205–210. http://dx.doi.org/10.2989/10220110409485853 kemp, a.c., 1976, ‘factors affecting the onset of breeding in african hornbills’, proceedings of the 16th international ornithological congress, canberra, australia, august 12–17, 1974, pp. 248–257. kemp, a.c., 1988, ‘the behavioural ecology of the southern ground hornbill: are competitive offspring at a premium?’, proceedings of the international 100 do-g meeting, current topics in avian biology, bonn, germany, 1988, pp. 267–271. kemp, a.c., 1995, the hornbills, bucerotiformes, oxford university press, oxford. kemp, a.c., 2000, ‘southern ground hornbill’, in k.n. barnes (ed.), the eskom red data book of birds of south africa, lesotho and swaziland, pp. 117–119, birdlife south africa, johannesburg. kemp, a.c., 2005, ‘ground hornbills’, in p.a.r. hockey, w.a.j. dean & p.g. ryan (eds.), roberts birds of southern africa, 7th edn., pp. 158–159, john voelcker bird book fund, cape town. kemp, a.c. & kemp, m.i., 1980, ‘the biology of the southern ground hornbill bucorvus leadbeateri (vigors) (aves: bucerotidae)’, annals of the transvaal museum 32(4), 65–100. kemp, a.c. & kemp, m.i., 1991, ‘timing of egg laying by southern ground hornbills bucorvus leadbeateri in the central kruger national park, south africa’, ostrich 62, 80–82. kemp, a.c., joubert, s.c.j. & kemp, m.i., 1989, ‘distribution of southern ground hornbills in the kruger national park in relation to some environmental features’, southern african journal of wildlife research 19(3), 93–98. kemp, m.i. & kemp, a.c., 1978, ‘bucorvus and sagittarius: two modes of terrestrial predation’, in a.c. kemp (ed.), proceedings of the symposium on african predatory birds, northern transvaal ornithological society, pretoria. knight, g.m., 1990, ‘status, distribution and foraging ecology of the southern ground hornbill (bucorvus cafer) in natal’, unpublished msc thesis, department of zoology and entomology, university of natal, durban. mucina, l. & rutherford, m.c., 2006, ‘the vegetation of south africa, lesotho and swaziland’, strelitzia 19, south african botanical institute, pretoria. oberprieler, r., 1986, ‘pinus radiata. don: a new foodplant record for lmbrasia (gonimbrasia) belina (westwood) (lepidoptera: saturniidae)’, journal of the entomological society of southern africa 49, 168–170. perner, j. & schueler, s., 2004, ‘estimating the density of ground-dwelling arthropods with pitfall traps using a nested-cross array’, journal of animal ecology 73, 469–477. http://dx.doi.org/10.1111/j.0021-8790.2004.00821.x poulin, b., gaetan, l. & mcneil, r., 1992, ‘tropical avian phenology in relation to abundance and exploitation of food resources’, ecology 73(6), 2295–2309. http://dx.doi.org/10.2307/1941476 seaman, a.d. & powell, r.a., 1996, ‘an evaluation of the accuracy of kernel density estimators for home range analysis’, ecology 77(7), 2075–2085. http://dx.doi.org/10.2307/2265701 south, a.b., kenward, r.e. & walls, s.s., 2008, ranges 7: for the analysis of tracking and location data, online manual, anatrack ltd., wareham. standen, v., 2000, ‘the adequacy of collecting techniques for estimating species richness of grassland invertebrates’, journal of applied ecology 37(5), 884–893. http://dx.doi.org/10.1046/j.1365-2664.2000.00532.x statistica version 9, 2009, computer software, statsoft inc., tulsa. theron, n.t., dalton, d., grobler, j.p., jansen, r. & kotze, a., 2013, ‘molecular insights on the re-colonization of the limpopo valley, south africa, by southern ground-hornbills’, journal of ornithology 154(3), 727–737. http://dx.doi.org/10.1007/s10336-013-0937-4 vernon, c.j., 1984, ‘the ground hornbill at the southern extremity of its range’, ostrich 57, 16–24. http://dx.doi.org/10.1080/00306525.1986.9633633 wolda, h., 1979, ‘seasonality parameters for insect populations’, researches on population ecology 20(2), 247–256. http://dx.doi.org/10.1007/bf02512630 abstract introduction research method and design results discussion conclusion acknowledgements references appendix 1: environmental data description for addo elephant national park appendix 2: elephant location data summary appendix 3: availability sample size sensitivity analysis appendix 4: elephant predicted use animations about the author(s) timothy j. fullman department of geography, university of florida, united states gregory a. kiker department of agricultural and biological engineering, university of florida, united states school of mathematics, statistics and computer science, university of kwazulu-natal, south africa angela gaylard conservation services, south african national parks, garden route regional office, south africa jane southworth department of geography, university of florida, united states peter waylen department of geography, university of florida, united states graham i.h. kerley department of zoology, nelson mandela metropolitan university, south africa citation fullman, t.j., kiker, g.a., gaylard, a., southworth, j., waylen, p. & kerley, g.i.h., 2017, ‘elephants respond to resource trade-offs in an aseasonal system through daily and annual variability in resource selection’, koedoe 59(1), a1326. https://doi.org/10.4102/koedoe.v59i1.1326 original research elephants respond to resource trade-offs in an aseasonal system through daily and annual variability in resource selection timothy j. fullman, gregory a. kiker, angela gaylard, jane southworth, peter waylen, graham i.h. kerley received: 10 june 2015; accepted: 15 nov. 2016; published: 28 mar. 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract animals and humans regularly make trade-offs between competing objectives. in addo elephant national park (aenp), elephants (loxodonta africana) trade off selection of resources, while managers balance tourist desires with conservation of elephants and rare plants. elephant resource selection has been examined in seasonal savannas, but is understudied in aseasonal systems like aenp. understanding elephant selection may suggest ways to minimise management trade-offs. we evaluated how elephants select vegetation productivity, distance to water, slope and terrain ruggedness across time in aenp and used this information to suggest management strategies that balance the needs of tourists and biodiversity. resource selection functions with time-interacted covariates were developed for female elephants, using three data sets of daily movement to capture circadian and annual patterns of resource use. results were predicted in areas of aenp currently unavailable to elephants to explore potential effects of future elephant access. elephants displayed dynamic resource selection at daily and annual scales to meet competing requirements for resources. in summer, selection patterns generally conformed to those seen in savannas, but these relationships became weaker or reversed in winter. at daily scales, resource selection in the morning differed from that of midday and afternoon, likely reflecting trade-offs between acquiring sufficient forage and water. dynamic selection strategies exist even in an aseasonal system, with both daily and annual patterns. this reinforces the importance of considering changing resource availability and trade-offs in studies of animal selection. conservation implications: guiding tourism based on knowledge of elephant habitat selection may improve viewing success without requiring increased elephant numbers. if aenp managers expand elephant habitat to reduce density, our model predicts where elephant use may concentrate and where botanical reserves may be needed to protect rare plants from elephant impacts. introduction conservation often involves trade-offs between competing goals and objectives. while trade-offs between biodiversity conservation and human needs have been emphasised (e.g. hirsch et al. 2011; mcshane et al. 2011), it is also important to take into account trade-offs between conserving different aspects of biodiversity. in addo elephant national park (aenp), south africa, managers seek to balance the local conservation of african elephants (loxodonta africana) and the ecological processes they provide with protecting rare plants and providing for tourism. the park was founded in 1931 to protect the remnant population of elephants in the eastern cape province following depletion by hunting (hall-martin 1980). fences were erected in 1954 to prevent conflict with farmers, and the population rapidly grew to its current level of over 500 elephants (hall-martin 1980; sanparks 2011). although elephants comprise around 80% of the herbivore biomass in aenp (landman et al. 2012), the park is also home to 19 other large mammal species (sanparks 2008). this diversity of species attracts national and international tourists, with over 90% of tourists rating wildlife viewing as a ‘very important’ motivation for visiting (boshoff et al. 2007). demand by tourists for wildlife viewing opportunities sometimes leads to stocking high densities of tourist-favoured species, such as elephants. such actions, however, may fail to meet tourism objectives and may conflict with the south african national parks (sanparks) mandate to preserve diverse, healthy ecosystems (maciejewski & kerley 2014). this mismatch is especially crucial in aenp, which contains a wide variety of rare and important plant species in addition to animals (lombard et al. 2001). the park contains portions of five of the nine south african biomes, comprising 43 different vegetation units (sanparks 2008). concerns about disturbance of vegetation by elephants have been raised by several studies that documented loss of plant biodiversity and biomass (kerley & landman 2006; landman, kerley & schoeman 2008; lombard et al. 2001). a better understanding of the factors influencing elephant movement is needed to inform conservation decisions by park managers regarding wildlife, rare plants and tourism (ferreira et al. 2011). such information can be used to identify where high elephant use coincides with rare plant hotspots, suggesting locations for enclosure with fences to protect rare plants from elephants and other herbivores. it can also help guide tourist viewing to maintain a balance between ecological health and visitor satisfaction. currently elephants are fenced into three discrete sections of aenp, occupying about 60 000 ha of the total 160 000 ha area of the park (ferreira et al. 2011). understanding how elephants use resources in the currently accessible areas of aenp will clarify possible effects of future expansion of access to other areas of the park. we investigate elephant resource selection through satellite imagery and movement data to evaluate how elephant space use responds to vegetation productivity (greenness), artificial water points, slope and terrain ruggedness across space and time in aenp. while numerous studies consider the influence of vegetation, water and other factors on elephant movement and distribution (e.g. chamaillé-jammes et al. 2013; loarie, van aarde & pimm 2009b; marshal et al. 2011; roever, van aarde & leggett 2012), they are generally conducted in savanna systems, featuring distinct wet and dry seasons, with differing factors influencing elephant habitat selection in each season (loarie et al. 2009b; roever et al. 2012). aenp, on the contrary, exhibits no strong seasonal rainfall pattern (ferreira et al. 2011; gough & kerley 2006) and features evergreen succulent thicket in the main elephant areas (landman et al. 2008), rather than the savanna vegetation of trees and grasses typical of elephant populations elsewhere. it is unclear how elephant resource use in aseasonal thicket will differ from resource use in savanna systems. as a set of non-exclusive hypotheses, we predict that elephant resource selection in the aseasonal thicket of aenp will correspond to that observed in savanna environments. while selection patterns may well differ in an aseasonal thicket ecosystem, these hypotheses provide a basis for comparison. elephants must consume great quantities of forage to sustain their large body size and high absolute metabolic requirements (hopcraft, olff & sinclair 2010). positive selection for green vegetation has been indicated in several studies of elephant habitat use (loarie et al. 2009b; pittiglio et al. 2012; wall et al. 2013; but see boettiger et al. 2011). elephants are water-dependent herbivores and typically require access to drinking water every day or two (chamaillé-jammes et al. 2013; owen-smith et al. 2006). water availability alters elephant movement and distribution patterns (chamaillé-jammes, valeix & fritz 2007; loarie, van aarde & pimm 2009a; shannon et al. 2009; smit, grant & whyte 2007a), with elephants typically selecting for areas near water. steep slopes require high energy expenditures from elephants because of their large body size, and thus tend to be avoided (roever et al. 2012; wall, douglas-hamilton & vollrath 2006). rugged terrain is preferentially used by elephants because of the presence of greater nutrient concentrations and forage density compared with less rugged areas (nellemann, moe & rutina 2002). in summary, if elephants in aseasonal aenp follow the patterns exhibited by elephants in seasonal systems, we expect them to show (1) a positive association with vegetation greenness, (2) a negative association with distance to water, (3) a negative association with steeper slopes and (4) a positive association with more rugged terrain. we use step selection functions (forester, im & rathouz 2009; fortin et al. 2005) to evaluate the hypotheses identified above, measuring elephant resource selection at a daily scale in aenp. environmental covariates are interacted with functions of time to investigate annual variability in selection patterns, while three data sets of daily movement steps recorded in the morning (06:00), midday (12:00) and afternoon (16:00) are analysed to reflect circadian patterns of resource selection. we use the results of these analyses to suggest implications for management that balance the needs of elephants, the demands of tourists and the potential of future elephant expansion to other areas of aenp. research method and design study area aenp lies along the southern coast of south africa (figure 1) and features an aseasonal climate (figure 1-a1), with about 400 mm – 450 mm of year-round rainfall (gough & kerley 2006; lombard et al. 2001). there are no permanent natural water sources in the main camp and colchester sections of the park (sanparks 2008), aside from a single brackish spring in colchester. instead, water is provided through artificial water points fed by pumped groundwater (landman et al. 2012). figure 1: (a) addo elephant national park exists in an aseasonal climatic system on the south african coast. (b) the elephant-accessible areas evaluated in this study lie at the heart of the larger addo elephant national park. (c) primary elephant-accessible areas of addo elephant national park. the fence separating main camp and colchester was dropped in august 2010, making both sections available to elephants. the nyathi section remained isolated by fences from main camp for the duration of the study. botanical reserves and other fenced areas are inaccessible to elephants. a smaller population of elephants occupies the kuzuko section of addo elephant national park (not pictured) and is not included in this study. for most of aenp’s history, elephants were confined primarily within the main camp section of the park (figure 1c). as the elephant population grew, this section was enlarged, increasing from 22.7 km2 at the time of fencing to 120.0 km2 in 2008 (gough & kerley 2006; hall-martin 1980; landman et al. 2012). several areas within the elephant-accessible portion of the park were fenced off as botanical reserves to protect vulnerable plants from elephants and other herbivores (figure 1c; lombard et al. 2001). in august 2010, the fence separating the main camp and colchester sections of aenp (figure 1c) was removed, providing elephants with access to both areas. at the time of this study, most of the elephants in the park occupied the main camp/colchester section, with a smaller population separated by a fence in the nyathi section (figure 1c; sanparks 2011). a third, even smaller population was established in the kuzuko section of aenp in 2005. other sections not available to elephants have also been added to aenp over time to expand protection of the diverse biomes in the region (figure 1b). in the late 1990s, a ‘greater addo elephant national park’ was proposed to offer a combination of terrestrial and marine areas that would promote both conservation and development (kerley & boshoff 1997). while elephants currently do not have access to these areas, there is interest by park management in expanding elephant access to other sections. data description elephant telemetry data geographic positioning system (gps) collar data (africa wildlife tracking, pretoria, south africa) were obtained for seven female elephants, each representing a different family group. elephants were collared by sanparks veterinarians following established animal care protocols. six of the collared elephants occupied the main camp – colchester section of aenp, while one occupied the nyathi section (figure 1c). collars recorded location data at frequencies ranging from hourly to three records per day between march 2010 and march 2013. records were filtered to create three data sets of daily observations: daily locations at 06:00, 12:00 and 16:00. positional dilution of precision (pdop) records for each collar indicated high levels of accuracy (table 1-a2; d’eon & delparte 2005). remotely sensed vegetation data vegetation data were obtained from the moderate resolution imaging spectroradiometer (modis). the mod13q1 product provided vegetation indices at a 250 m spatial resolution in 16-day composites. each pixel in a composite contained the maximum observation across the 16-day period to reduce issues associated with clouds, aerosol loading and shadows (huete et al. 2011). normalised difference vegetation index (ndvi) layers from modis tile h20v12 were obtained from the usgs land processes distributed active archive center (https://www.lpdaac.usgs.gov/) for the period spanning from 06 march 2010 to 21 march 2013. ndvi indicates the ‘greenness’ of plants across the landscape (huete, justice & van leeuwen 1999), reflecting vegetation productivity (pettorelli et al. 2005) and nutritional quality (loarie et al. 2009b). it is commonly used in wildlife ecology and management (pettorelli et al. 2011) and in studies of elephant movement (e.g. boettiger et al. 2011; marshal et al. 2011). preprocessing of modis composites was conducted using the modis reprojection tool (https://www.lpdaac.usgs.gov/tools/modis_reprojection_tool) and arcgis (version 9.3, esri, redlands, ca). this included projecting modis composites to universal transverse mercator (utm) zone 35s and resampling using nearest neighbour sampling to ensure pixels were exactly 250 m × 250 m. all data with pixel qualities of 0 and 1 were retained for analysis, based on the pixel reliability summary provided with the mod13q1 data. other covariate data in addition to ndvi, covariate data were obtained for the distance to artificial water points (hereafter distance to water), slope and terrain ruggedness (figure 2-a1). distance to water was calculated in arcgis at a 250 m spatial resolution using shapefiles of water point locations obtained from sanparks. elevation data were obtained at a 30 m × 30 m resolution from the aster global digital elevation model v2 (aster gdem is a product of meti and nasa). data were downloaded from nasa reverb (http://www.reverb.echo.nasa.gov/) and then projected to utm zone 35s in arcgis. block means were calculated on the elevation data using a 250 m × 250 m rectangular neighbourhood to match the resolution of the ndvi and distance to water data. the resulting raster was resampled to a 250 m resolution using nearest neighbour sampling. slope was calculated from the elevation raster using the spatial analyst toolbox in arcgis. terrain ruggedness was calculated using the vector ruggedness measure (vrm) developed by sappington, longshore and thompson (2007). this measure takes into account heterogeneity in both slope and aspect and yet has a low correlation with slope, allowing both measures to be included in analyses to represent different components of selection by animals (sappington et al. 2007). terrain ruggedness was calculated in arcgis using the vrm tool (vrm 2012) with a 3 × 3 pixel window on the 250 m elevation raster. resource selection analysis step selection functions (ssfs; forester et al. 2009; fortin et al. 2005) were developed using conditional logistic regression (clr) models to investigate resource selection by elephants at a daily scale. these models, alternatively called discrete choice models, evaluate ‘choice sets’ contrasting characteristics of the choice made at a given time (the elephant location) with those that were available (other nearby locations; duchesne, fortin & courbin 2010; manly et al. 2002). how ‘available’ locations are defined in such studies can strongly influence findings (beyer et al. 2010). when used to evaluate resource selection functions, clr models typically involve a matched design in which animal presence points are associated with ‘available’ locations drawn within a limited area (boyce 2006; duchesne et al. 2010). we sampled the empirical distribution of daily step lengths and turning angles for all individuals to determine our available locations (forester et al. 2009). step lengths and turning angles were calculated using the ‘adehabitatlt’ package in r (calenge 2006; r core team 2016). twenty available points were drawn for each observed elephant location, based on a sensitivity analysis (appendix 3). available points drawn from areas inaccessible to elephants (i.e. outside the park or within botanical reserves) were discarded and replacement points were drawn. we developed a set of candidate models representing potential resource selection by elephants at a daily scale (table 1) for locations recorded in the morning (06:00), midday (12:00) and afternoon (16:00). all models contained a spline for distance to the previous used point to help reduce bias in ssf estimation (forester et al. 2009). distance splines were included using the pspline function of the ‘survival’ package in r (therneau 2015; therneau & grambsch 2000) with two degrees of freedom, following the approach of panzacchi et al. (2016). variance inflation factors (vif) were used to check for the presence of collinearity among covariates in the full model using r code from zuur et al. (2009). all vif values were less than 1.5; therefore, all covariates were retained in the candidate model set. to allow for changing selection over time, candidate models were also run with each covariate interacted with a function of time. time was included following equation 3 in wilson et al. (2014), using a function f(t), defined as: where t is the julian day of an elephant location and its associated available locations. population-level ssf analyses under each candidate model were run for the six elephants occupying the main camp – colchester section of aenp. covariate values were scaled by subtracting the mean and dividing by the standard deviation, following the approach of northrup et al. (2013). models were run using the ‘survival’ package in r. selection between models was performed with the ‘aiccmodavg’ package in r (mazerolle 2016), using akaike’s information criterion corrected for small sample size (aicc; burnham & anderson 2002) to select the most parsimonious model. table 1: candidate models for elephant resource selection in addo elephant national park, south africa. we followed the approach of forester et al. (2009) to calculate robust standard errors controlling for multiple observations per individual. this involved calculating deviance residuals for each top model identified through the model selection process, fitting an intercept-only mixed-effects model to the residuals with a random intercept for individual elephants, using the autocorrelation function of this model to determine the lag of correlation, and assigning the data into independent clusters based on the identified lag. the data were subset into two independent models using the clusters and a clr model was fit on each subset. the resulting covariance matrices from each subset were then averaged to provide adjusted standard errors. predictive performance of the top model for each daily data set was assessed both for interpolative ability within the main camp – colchester section of aenp and extrapolative ability using independent data from the collared female elephant in the nyathi section. interpolative predictive ability was determined using k-fold cross-validation (boyce et al. 2002; johnson et al. 2006), with the used and available data from the six main camp – colchester elephants split 90% – 10% across 10 folds. differences in covariate values in the nyathi section of aenp compared with the main camp – colchester section (figures 2-a1 and 3-a1) led to exclusion of the nyathi elephant from the population-level resource selection model, but provided an opportunity to test model predictive performance in an area with differing availability. extrapolative predictive ability was evaluated with an adapted version of the boyce et al. (2002) approach. the top clr model for each of the three daily data sets was predicted to the nyathi section of aenp and the resulting values were binned into 10 quantiles. the spearman’s rank correlation between binned values at locations used by the nyathi elephant and the expected number of observations per bin based on the clr predictions was used to indicate predictive performance. to visually represent habitat use over time by elephants in aenp, relative habitat suitability was predicted for the main camp – colchester section at a 250 m resolution following equation 2 in wilson et al. (2014): where w(s, t) is the relative habitat use prediction for pixel s at time t, β is the vector of selection coefficients estimated using the procedure above, f(t) follows equation 1 above and x(s, t) is the vector of environmental covariates measured for pixel s at time t. predicted values were then rescaled using a linear stretch to range between 0 and 1 following equation 7 in decesare et al. (2012): predicted maps of relative elephant use within the main camp – colchester area were estimated for 70 sixteen-day periods, corresponding to the modis ndvi composites described above. an aggregated view of the relative probability of elephant use was obtained by averaging each of the 70 predicted maps and applying a linear stretch following equation 3. the elephant-accessible areas of aenp are situated within a larger network of sections that together make up the proclaimed national park (figure 1b). previous management of elephant numbers in aenp has involved opening new sections of the park to elephants (kerley & landman 2006), as witnessed in the colchester section during this study. managers are thus not only interested in how elephants use habitats within currently accessible areas but also in likely movements of elephants if additional fences are removed. to provide a preliminary answer to this question, we used the midday ssf model to predict relative probability of use by elephants across the greater aenp. only the midday model was used in this extrapolation because it was the only model to show robust predictions when extrapolated to the nyathi area (see the results section). results the seven collared elephants in this study (six in the main camp – colchester section of aenp plus one in the nyathi section) each contributed between 504 and 1046 locations to the ssf analyses (table 1-a2). depending on the time of the day, elephants exhibited daily steps that averaged between 2151.5 m and 2267.5 m (table 1-a2). results were similar when the elephant occupying the nyathi section of aenp was excluded (table 2-a2). all three daily movement data sets had clearly supported top models (table 2). the full model including all covariates and time-varying selection coefficients (model 30 in table 1) was the best for all three data sets, with akaike weights between 0.98 and 1.00. selection patterns varied across the daily data sets (table 3), indicating both daily and annual variation in resource selection by female elephants in aenp. predictive performance of the daily models varied across data sets for both interpolation and extrapolation (table 3). interpolative predictive performance, as indicated by k-fold cross-validation, was high for both the 12:00 and 16:00 daily selection models, while the 06:00 model performed poorly. extrapolative predictive performance, predicting use for the nyathi elephant from main camp – colchester resource selection models, showed strong predictive ability at 12:00, but weak performance at 06:00 and 16:00. table 2: model selection results for elephant resource selection at a daily scale in addo elephant national park, south africa. daily selection was considered in the morning (06:00), midday (12:00) and afternoon (16:00) to reflect within-day variation in selection patterns. table 3: step selection function regression coefficients and associated standard errors for elephants in addo elephant national park at a daily scale, recorded in the morning (06:00), midday (12:00) and afternoon (16:00). we tested four hypotheses of elephant selection based on patterns observed in savanna systems, namely that elephants would show (1) a positive association with vegetation greenness, (2) a negative association with distance to water, (3) a negative association with steeper slopes and (4) a positive association with more rugged terrain. elephant selection of ndvi varied across the year, with the strongest selection in the winter (july–september) and the least selection in the summer (november–february; figure 2a). daily selection of greener vegetation (high ndvi values) in the morning (06:00) was consistently strong across the year, with the exception of february, aligning with the first hypothesis. at midday (12:00), however, selection patterns were more nuanced, with elephants avoiding greener vegetation from mid-november to march and selecting for greener vegetation from late may to september. elephants showed little selection or avoidance of ndvi in the afternoon (16:00) throughout the year, contrasting with the first hypothesis. consistent with the second hypothesis, female elephants in aenp tended to select areas closer to water (negative selection coefficient for distance to water), though this varied both diurnally and annually (figure 2b). elephants showed a strong selection for areas near water points in the summer (november–mid-february) at midday and in the afternoon, and the weakest relationship in the morning. annual patterns in selection coefficients for slope at midday and in the afternoon (figure 2c) generally supported the third hypothesis, with a predominantly negative relationship that was strongest from october to february and weakest from may to august. conversely, daily selection patterns in the morning showed little relationship with slope in the summer, but a positive selection for steeper slopes in mid-winter (mid-may–mid-july). variability in selection coefficient values across the year for terrain ruggedness was similar in the morning, midday and afternoon (figure 2d). while some annual fluctuations in selection were apparent, 95% confidence intervals for all 3 hours always overlapped zero, indicating little support for selection or avoidance of rugged terrain and contrasting with the fourth hypothesis. figure 2: plots of annual coefficient estimates (lines) and their 95% confidence intervals (shaded regions) derived from step selection functions of daily elephant movements recorded at 06:00 (red), 12:00 (blue) and 16:00 (green) for (a) the normalised difference vegetation index, (b) distance to artificial water, (c) slope and (d) terrain ruggedness. the relative probability of use predicted across the main camp – colchester area using the ssf models varied throughout the day and across the year (figure 3; appendix 4). predicted use in the morning typically emphasised higher probabilities of use in different parts of the park than at midday and noon. on average across the study period, elephant use was predicted to be higher in the southern portions of the elephant-accessible area in the morning and in the northern parts at midday (figure 4). use in the afternoon was more evenly distributed across the elephant-accessible area (figure 4). when extrapolated to the greater aenp, the midday model predicted the highest probabilities of use in the currently available areas of main camp and colchester as well as the botanical reserves and southern portion of nyathi (figure 5). predicted use was also relatively high in the northernmost portion of the park around darlington dam. the central areas of aenp tended to exhibit lower relative probabilities of use according to the midday model. figure 3: predicted use by elephants in addo elephant national park on three representative dates throughout the study period (rows) and for the three times of day considered (columns). (a) 01 january 2011, 06:00, (b) 01 january 2011, 12:00, (c) 01 january 2011, 16:00, (d) 23 april 2011, 06:00, (e) 23 april 2011, 12:00, (f) 23 april 2011, 16:00, (g) 30 september 2011, 06:00, (h) 30 september 2011, 12:00, (i) 30 september 2011, 16:00. figure 4: average relative probability of use by elephants in addo elephant national park. maps reflect the predicted relative probability of use in (a) the morning (06:00), (b) midday (12:00) and (c) afternoon (16:00), averaged across the 70 sixteen-day periods corresponding to the moderate resolution imaging spectroradiometer composites. figure 5: relative probability of use by elephants across addo elephant national park. the map reflects the predicted relative probability of use based on the midday (12:00) step selection function model, averaged across the 70 sixteen-day periods corresponding to the moderate resolution imaging spectroradiometer composites used in this study. discussion our findings are consistent with our first three hypotheses in that elephants in subtropical thicket responded positively to feeding opportunities (vegetation greenness) and water and negatively to slope. this provides some confidence in the ability to extrapolate findings between biomes in terms of resource selection by elephants. the fourth hypothesis of a positive association of elephants with more rugged terrain was not supported. the latter hypothesis was predicated upon patterns of increased nutrient distribution being associated with rugged terrain in savanna landscapes (nellemann et al. 2002). the lack of support for this hypothesis was expected on the basis of the generally high nutrients in subtropical thicket soils (kerley & landman 2006). hence, given the oligotrophic nature of many savanna soils (east 1984), the lack of success in extrapolating this pattern between biomes likely reflects a biome-scale difference rather than a change in elephant resource selection. there is increasing recognition of the importance of understanding dynamic patterns both in animal movement (bowler & benton 2005; mueller & fagan 2008) and resource selection (mcloughlin et al. 2010). animals occupying dynamic environments face substantial variability at multiple spatial and temporal scales to which they must respond if they are to obtain the necessary resources to survive and reproduce. the interaction of these changes in environmental conditions and individual behavioural responses results in larger patterns of population-level movement and distribution (mueller & fagan 2008). understanding the drivers of these space use patterns enables managers to incorporate spatiotemporal patterns of species distribution and impacts into their conservation decisions seeking to balance multiple objectives. our results indicate that female elephants in aenp exhibit dynamic patterns of resource selection at both daily and annual scales. analysis of elephant selection for green vegetation, distance to water and slope indicates variable selection patterns over time both annually and daily. annual variability in resource selection (evans & harris 2012; jachowski, slotow & millspaugh 2012; shannon et al. 2006), movement patterns (leggett 2010; loarie et al. 2009a; wittemyer et al. 2008) and home range size (osborn 2003; shannon et al. 2006) has been previously reported for elephants, though typically in the context of seasonal systems. our study shows that in the aseasonal context of aenp, elephants also show variable selection patterns across the year. for ndvi, distance to water and slope, the most notable divergences from the hypotheses of selection based on patterns seen in savanna systems occur during the winter (approximately may–october; figure 2). while rainfall patterns are aseasonal in aenp, temperature follows much more consistent patterns, with the winter generally being cooler than the summer (figure 1-a1). ambient temperature influences habitat selection in elephants (kinahan, pimm & van aarde 2007). the cooler temperatures in the winter may relax the need for elephants to remain close to water, allowing them to select for areas of greener vegetation as seen in the positive midday selection coefficients for ndvi during this period. daily variation in elephant movement and resource selection is apparent in many systems (e.g. cook, henley & parrini 2015; galanti et al. 2006; graham et al. 2009). these patterns, however, are typically associated with human activity and perceptions of risk by elephants (but see loarie et al. 2009a). while previous studies often break daily patterns into day–night comparisons, we record finer levels of variability, with selection patterns in the morning often differing from those observed at midday and in the afternoon. this is most pronounced in the selection of green vegetation, but is also somewhat apparent for distance to water and slope. boettiger et al. (2011) suggest that elephant foraging strategies in seasonal savannas of northern kenya are more complex than simply selecting areas with the highest productivity, as has been indicated elsewhere (e.g. loarie et al. 2009b; wall et al. 2013). our study finds similar complexity of selection in the aseasonal system of aenp, where elephants consistently select for greener vegetation (higher ndvi values) in the morning, but exhibit negative or neutral selection patterns later in the day. a trade-off has been noted for many herbivores between balancing water requirements and nutritional needs (e.g. chamaillé-jammes et al. 2013; redfern et al. 2003). for elephants, the need to regularly access drinking water (chamaillé-jammes et al. 2013) constrains habitat use options (redfern et al. 2003), limiting the ability of elephants to consistently select for green vegetation. while the small size of aenp and relative frequency of water points in the elephant-accessible areas mean that water is not limiting in the traditional sense of a lack of sufficient water, the localisation of available water interacting with the regular need to access water by elephants may still constrain movement decisions. indeed, water can still serve as a constraining factor even in systems with very high water availability (redfern et al. 2003). in aenp, we propose that the concentration of available water at borehole-fed artificial water points leads elephants to function like central-place foragers, as has been seen for other species (rozen-rechels et al. 2015). elephants may venture away from water points in the morning, seeking to maximise their forage intake and meet their nutritional requirements, before returning to water points in the middle of the day. this could explain the generally positive selection for greener vegetation and weaker relationship with distance to water observed in the morning data. as a generalist forager (kerley & landman 2006), elephants may use a variety of habitats to meet their nutritional requirements. however, the finite water availability in aenp leads to similar patterns of use when drinking, as common targets constrain resource selection strategies (boettiger et al. 2011). this diversity of options for elephant use in the morning may explain why the morning ssf model had substantially poorer interpolative prediction ability compared with the midday and afternoon models. in addition to elephants balancing forage–water trade-offs through daily variability in habitat use, they may be able to further minimise trade-offs through habitat modification. high elephant utilisation around artificial water points leads to changes in vegetation structure and composition, with succulent thicket vegetation being largely replaced by grass (landman et al. 2012). o’connor, goodman and clegg (2007) suggested that elephants should favour green grasses and forbs as a food source because of their high potential intake rate. dietary studies indicate this is the case in aenp, as cynodon dactylon, the main grass growing around highly utilised water points (landman et al. 2012), comprises a significant proportion of elephant diet (landman et al. 2008; paley & kerley 1998). elephants thus may be creating favourable conditions that enhance their forage opportunities, similar to patterns seen in other systems (du toit & olff 2014). these modified areas may allow elephants to meet their water requirement while minimising their cost in reduced forage opportunities. all location data in this study are for female elephants. patterns may differ for male elephants, as studies from various parts of southern africa have found behavioural, diet and movement differences between sexes (evans & harris 2012; shannon, mackey & slotow 2013; smit, grant & whyte 2007b). similarly, whitehouse and schoeman (2003) reported differences between female and male elephant movement patterns in aenp. in addition, duffy et al. (2011) suggested that sexual activity may alter elephant movement patterns, even overriding the importance of factors such as water. it is unclear what influence male elephants in aenp may have had on female movement and habitat selection or how the patterns described here for female herds reflect habitat selection decisions made by males. however, gps collars were recently placed on several male elephants in aenp (a. gaylard, unpublished data), enabling future comparison of habitat selection patterns for male elephants with those observed here. management implications balancing the objectives of protecting biodiversity and enhancing tourist experiences and outcomes is a key challenge for conservation in protected areas worldwide. in some cases, a desire to improve tourist outcomes leads to managers increasing numbers of charismatic species. in aenp, this has meant introducing lions (panthera leo; sanparks 2008), while other south african parks and reserves stock high densities of lions, cheetahs (acinonyx jubatus) and elephants (clements, cumming & kerley 2016; maciejewski & kerley 2014). at high densities, however, elephants have negative impacts on the vegetation and animals of aenp (kerley & landman 2006; landman & kerley 2014; landman et al. 2008; tambling et al. 2013), threatening the rare plants contained within the park and compromising the mandate of sanparks to protect biodiversity. fortunately, recent analyses suggest that high elephant densities may not be necessary to produce positive tourist experiences (maciejewski & kerley 2014). this provides an opportunity to promote ‘smarter tourism’, improving viewing success by guiding tourists towards areas where they are likely to see elephants, rather than by increasing elephant numbers artificially. the results of this and other similar studies can inform such efforts, providing daily and annual guidance on elephant space use patterns that can help visitors’ planning. for example, our ssf models suggest that areas around water points may not provide the best viewing opportunities for early morning tourism. tourists should visit other areas of the park in the morning or focus on other species during this period. by midday, the probability of viewing elephants around water points should increase and should be maintained into the afternoon, though with greater variability. these models can also be used to inform plans to develop tourism infrastructure such as roads that will improve access to elephant-viewing opportunities. it is important to note, however, that these are general trends and that specific patterns of habitat use by elephants may vary at different times of the year as elephants respond to other factors such as ambient temperature or rainfall (birkett et al. 2012). in aenp, as in many other small fenced reserves in south africa (mackey et al. 2006; slotow et al. 2005), the elephant population has grown rapidly over time, often exceeding the estimated carrying capacity of the elephant-accessible areas (gough & kerley 2006; hayward & kerley 2009). one way managers have addressed increasing elephant numbers in aenp is by opening new areas to elephants, temporarily reducing overall density (gough & kerley 2006). our projections of the relative probability of elephant use across the greater aenp (figure 5) offer insights into where elephant use may concentrate if given access to new areas. as expected, the areas currently available to elephants in main camp, colchester and the southern portion of nyathi show high likelihood of continued use by elephants. the use of these areas by elephants is likely to remain high even if other areas are made available because of the long-term spatiotemporal memory of elephants (van aarde et al. 2008), which enables social transmission of site fidelity across generations (fishlock, caldwell & lee 2016). the botanical reserves and other fenced areas in the main camp and colchester sections are explicitly included in our forecasts and, as expected, show an overall high likelihood of use by elephants, emphasising their importance in protecting rare plants from elephant impacts (lombard et al. 2001). other areas that are included in the greater aenp but currently lack permanent water (shown in purple in figure 5) could be made available to elephants if artificial water points are established, as they have been in main camp, colchester and nyathi. managers can use our resource selection model to test the effects of potential water point placement on predicted probabilities of elephant use in these disconnected areas to assess their suitability for elephant introduction. we note that although the extrapolative validation of the midday ssf model showed it to be robust to making predictions under different resource availability in the nyathi section, this does not guarantee that the observed relationships will be maintained elsewhere across the greater aenp. this may especially be true if elephants modify existing habitats as noted above, which could alter the probability of elephant use in some areas. in addition, our analyses show that elephant resource selection and habitat use patterns in aenp vary throughout the day; thus, maps generated based on midday selection relationships may not reflect the relative probability of use by elephants at all hours of the day. our predictions should be treated as hypotheses of future use by elephants. field verification is required to determine how elephant use actually conforms to model predictions as access is expanded. these observations will reveal how vegetation communities change as elephants move into areas from which they have long been absent and may suggest whether additional botanical reserves need to be established to protect rare plants from elephant impacts. conclusion behavioural trade-offs are common in nature. we find this to be true for female elephants in the aseasonal system of aenp, which employ dynamic patterns of resource selection at daily and annual scales to meet their competing requirements for forage, water and other resources. in the summer, selection patterns for vegetation productivity (ndvi), distance to water and slope generally conform to those exhibited by elephants in savanna systems, but these relationships become weaker or reversed in the winter. at daily scales, resource selection by elephants varies in the morning from patterns seen at midday and in the afternoon, likely reflecting temporal reactions of elephants to the trade-off between acquiring sufficient forage and water. these results reinforce the importance of considering dynamic resources in studies of animal selection (mcloughlin et al. 2010). for elephants, future research should expand consideration of daily variability in movement and resource selection from situations relating to human–elephant conflict to consider other aspects of resource use. protected area managers also seek to balance trade-offs between competing objectives. in aenp, managers must weigh trade-offs between the desires of tourists and the conservation needs of elephants, rare plants and other elements of biodiversity. our findings provide an opportunity to promote ‘smarter tourism’, improving viewing success by guiding tourists towards areas where they are likely to see elephants and improving tourist access infrastructure in such areas, rather than by increasing elephant numbers. in general, areas near waterholes are likely to be highly utilised by elephants at midday and in the afternoon, offering rewarding tourist viewing opportunities, but are less likely to be used by elephants in the morning. the results also offer insights into possible effects of future expansion of elephant access in aenp, predicting where elephant use may concentrate if elephants are given access to new areas. such information can suggest priorities for detailed assessment of vegetation communities to determine where key areas of rare plants occur prior to elephant access. such information may allow establishment of new botanical reserves, helping protect plants from elephant impacts and improving the likelihood of sustainable outcomes in the park. acknowledgements the authors would like to thank south african national parks (sanparks) for providing permission, data and support for this project, especially to z. silcock. m. landman provided helpful background about aenp and its elephants. p. bradshaw and i. smit provided gis data for aenp. r. wilson and d. gregovich offered helpful suggestions and r code that improved the analysis. funding was provided by the cleveland metroparks zoo and cleveland zoological society, national science foundation grant no. 0801544 and nasa project nnx09ai25g. the wilderness society provided support to t.j.f. for the writing of this article. the funders of this work provided financial support but had no role in study design, data collection, analysis or interpretation of results. these actions and the decision to publish were solely at the discretion of the authors. r. fletcher and three anonymous reviewers provided helpful comments on previous versions of this article. j. steele facilitated formation of collaborative research between sanparks, the university of florida and nelson mandela metropolitan university. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions t.j.f. was the project leader and designed and conducted the analysis. a.g. and g.i.h.k. provided elephant telemetry data and offered insightful revisions of the article. p.w. provided guidance with analysis and a thorough revision of the article. g.a.k. and j.s. supervised the project and contributed to the writing and revision of the article. references beyer, h.l., haydon, d.t., morales, j.m., frair, j.l., hebblewhite, m., mitchell, m. et al., 2010, ‘the interpretation of habitat preference metrics under use-availability designs’, philosophical transactions of the royal society b 365, 2245–2254. https://doi.org/10.1098/rstb.2010.0083 birkett, p.j., vanak, a.t., muggeo, v.m.r., ferreira, s.m. & slotow, r., 2012, ‘animal perception of seasonal thresholds: changes in elephant movement in relation to rainfall patterns’, plos one 7(6), e38363. https://doi.org/10.1371/journal.pone.0038363 boettiger, a.n., wittemyer, g., starfield, r., volrath, f., douglas-hamilton, i. & getz, w.m., 2011, ‘inferring ecological and behavioral drivers of african elephant movement using a linear filtering approach’, ecology 92(8), 1648–1657. https://doi.org/10.1890/10-0106.1 boshoff, a.f., landman, m., kerley, g.i.h. & bradfield, m., 2007, ‘profiles, views and observations of visitors to the addo elephant national park, eastern cape, south africa’, south african journal of wildlife research 37, 189–196. https://doi.org/10.3957/0379-4369-37.2.189 bowler, d.e. & benton, t.g., 2005, ‘causes and consequences of animal dispersal strategies: relating individual behaviour to spatial dynamics’, biological review 80, 205–225. https://doi.org/10.1017/s1464793104006645 boyce, m.s., 2006, ‘scale for resource selection functions’, diversity and distributions 12, 269–276. https://doi.org/10.1111/j.1366-9516.2006.00243.x boyce, m.s., vernier, p.a., nielsen, s.e. & schmiegelow, f.k.a., 2002, ‘evaluating resource selection functions’, ecological modelling 157, 281–300. https://doi.org/10.1016/s0304-3800(02)00200-4 burnham, k.p. & anderson, d.r., 2002, model selection and multimodel inference: a practical information-theoretic approach, 2nd edn., springer-verlag, new york. calenge, c., 2006, ‘the package adehabitat for the r software: a tool for the analysis of space and habitat use by animals’, ecological modelling 197, 516–519. https://doi.org/10.1016/j.ecolmodel.2006.03.017 chamaillé-jammes, s., mtare, g., makuwe, e. & fritz, h., 2013, ‘african elephants adjust speed in response to surface-water constraint on foraging during the dry-season’, plos one 8(3), e59164. https://doi.org/10.1371/journal.pone.0059164 chamaillé-jammes, s., valeix, m. & fritz, h., 2007, ‘managing heterogeneity in elephant distribution: interactions between elephant population density and surface-water availability’, journal of applied ecology 44, 625–633. https://doi.org/10.1111/j.1365-2664.2007.01300.x clements, h.s., cumming, g.s. & kerley, g.i.h., 2016, ‘predators on private land: broad-scale socioeconomic interactions influence large predator management’, ecology and society 21(2), 45. https://doi.org/10.5751/es-08607-210245 cook, r.m., henley, m.d. & parrini, f., 2015, ‘elephant movement patterns in relation to human inhabitants in and around the great limpopo transfrontier park’, koedoe 57(1), art. #1298, 1–7. d’eon, r.g. & delparte, d., 2005, ‘effects of radio-collar position and orientation on gps radio-collar performance, and the implications of pdop in data screening’, journal of applied ecology 42, 383–388. https://doi.org/10.1111/j.1365-2664.2005.01010.x decesare, n.j., hebblewhite, m., schmiegelow, f., hervieux, d., mcdermid, g.j., neufeld, l. et al., 2012, ‘transcending scale dependence in identifying habitat with resource selection functions’, ecological applications 22, 1068–1083. https://doi.org/10.1890/11-1610.1 duchesne, t., fortin, d. & courbin, n., 2010, ‘mixed conditional logistic regression for habitat selection studies’, journal of animal ecology 79, 548–555. https://doi.org/10.1111/j.1365-2656.2010.01670.x duffy, k.j., dai, x., shannon, g., slotow, r. & page, b., 2011, ‘movement patterns of african elephants (loxodonta africana) in different habitat types’, south african journal of wildlife research 41, 21–28. https://doi.org/10.3957/056.041.0107 du toit, j. & olff, h., 2014, ‘generalities in grazing and browsing ecology: using across-guild comparisons to control contingencies’, oecologia 174, 1075–1083. https://doi.org/10.1007/s00442-013-2864-8 east, r., 1984, ‘rainfall, soil nutrient status and biomass of large african savanna mammals’, african journal of ecology 22, 245–270. https://doi.org/10.1111/j.1365-2028.1984.tb00700.x evans, k. & harris, s., 2012, ‘sex differences in habitat use by african elephants (loxodonta africana) in the okavango delta, botswana: is size really the deciding factor?’, african journal of ecology 50, 277–284. https://doi.org/10.1111/j.1365-2028.2012.01322.x ferreira, s., freitag-ronaldson, s., pienaar, d. & hendriks, h., 2011, elephant management plan: addo elephant national park 2011–2020, scientific services, south african national parks, skukuza, south africa. fishlock, v., caldwell, c. & lee, p.c., 2016, ‘elephant resource-use traditions’, animal cognition 19, 429–433. https://doi.org/10.1007/s10071-015-0921-x forester, j.d., im, h.k. & rathouz, p.j., 2009, ‘accounting for animal movement in estimation of resource selection functions: sampling and data analysis’, ecology 90, 3554–3565. https://doi.org/10.1890/08-0874.1 fortin, d., beyer, h.l., boyce, m.s., smith, d.w., duchesne, t. & mao, j.s., 2005, ‘wolves influence elk movements: behaviour shapes a trophic cascade in yellowstone national park’, ecology 86, 1320–1330. https://doi.org/10.1890/04-0953 galanti, v., preatoni, d., martinoli, a., wauters, l.a. & tosi, g., 2006, ‘space and habitat use of the african elephant in the tarangire-manyara ecosystem, tanzania: implications for conservation’, mammalian biology 7, 99–114. https://doi.org/10.1016/j.mambio.2005.10.001 gough, k.f. & kerley, g.i.h., 2006, ‘demography and population dynamics in the elephants loxodonta africana of addo elephant national park, south africa: is there evidence of density dependent regulation?’, oryx 40, 434–441. https://doi.org/10.1017/s0030605306001189 graham, m.d., douglas-hamilton, i., adams, w.m. & lee, p.c., 2009, ‘the movement of african elephants in a human-dominated land-use mosaic’, animal conservation 12, 445–455. https://doi.org/10.1111/j.1469-1795.2009.00272.x hall-martin, a., 1980, ‘elephant survivors’, oryx 15, 355–362. https://doi.org/10.1017/s0030605300028830 hayward, m.w. & kerley, g.i.h., 2009, ‘fencing for conservation: restriction of evolutionary potential or a riposte to threatening processes?’, biological conservation 142, 1–13. https://doi.org/10.1016/j.biocon.2008.09.022 hirsch, p.d., adams, w.m., brosius, j.p., zia, a., bariola, n. & dammert, j.l., 2011, ‘acknowledging conservation trade-offs and embracing complexity’, conservation biology 25, 259–264. hopcraft, j.g.c., olff, h. & sinclair, a.r.e., 2010, ‘herbivores, resources and risks: alternating regulation along primary environmental gradients in savannas’, trends in ecology and evolution 25, 119–128. https://doi.org/10.1016/j.tree.2009.08.001 huete, a., didan, k., van leeuwen, w., miura, t. & glenn, e., 2011, ‘modis vegetation indices’, in b. ramachandran, c.o. justice & m.j. abrams (eds.), land remote sensing and global environmental change: nasa’s earth observing system and the science of aster and modis, pp. 579–602, springer, new york. huete, a., justice, c. & van leeuwen, w., 1999, modis vegetation index (mod 13): algorithm theoretical basis document, version 3, usgs land process distributed active archive center, viewed 04 december 2012, from http://www.modis.gsfc.nasa.gov/data/atbd/atbd_mod13.pdf jachowski, d.s., slotow, r. & millspaugh, j.j., 2012, ‘physiological stress and refuge behavior by african elephants’, plos one 7(2), e31818. https://doi.org/10.1371/journal.pone.0031818 johnson, c.j., nielsen, s.e., merrill, e.h., mcdonald, t.l. & boyce, m.s., 2006, ‘resource selection functions based on use-availability data: theoretical motivation and evaluation methods’, journal of wildlife management 70, 347–357. https://doi.org/10.2193/0022-541x(2006)70[347:rsfbou]2.0.co;2 kerley, g.i.h. & boshoff, a., 1997, ‘a proposal for a greater addo national park: a regional and national conservation and development opportunity’, terrestrial ecology research unit report 17, university of port elizabeth, port elizabeth, south africa. kerley, g.i.h. & landman, m., 2006, ‘the impacts of elephants on biodiversity in the eastern cape subtropical thickets’, south african journal of science 102, 395–402. kinahan, a.a., pimm, s.l. & van aarde, r.j., 2007, ‘ambient temperature as a determinant of landscape use in the savanna elephant, loxodonta africana’, journal of thermal biology 32, 47–58. https://doi.org/10.1016/j.jtherbio.2006.09.002 landman, m. & kerley, g.i.h., 2014, ‘elephant both increase and decrease availability of browse resources for black rhinoceros’, biotropica 46, 42–49. https://doi.org/10.1111/btp.12066 landman, m., kerley, g.i.h. & schoeman, d.s., 2008, ‘relevance of elephant herbivory as a threat to important plants in the addo elephant national park, south africa’, journal of zoology 274, 51–58. landman, m., schoeman, d.s., hall-martin, a.j. & kerley, g.i.h., 2012, ‘understanding long-term variations in an elephant piosphere effect to manage impacts’, plos one 7(9), e45334. leggett, k., 2010, ‘daily and hourly movement of male desert-dwelling elephants’, african journal of ecology 48, 197–205. https://doi.org/10.1111/j.1365-2028.2009.01101.x loarie, s.r., van aarde, r.j. & pimm, s.l., 2009a, ‘fences and artificial water affect african savannah elephant movement patterns’, biological conservation 142, 3086–3098. https://doi.org/10.1016/j.biocon.2009.08.008 loarie, s.r., van aarde, r.j. & pimm, s.l., 2009b, ‘elephant seasonal vegetation preferences across dry and wet savannas’, biological conservation 142, 3099–3107. https://doi.org/10.1016/j.biocon.2009.08.021 lombard, a.t., johnson, c.f., cowling, r.m. & pressey, r.l., 2001, ‘protecting plants from elephants: botanical reserve scenarios within the addo elephant national park, south africa’, biological conservation 102, 191–203. https://doi.org/10.1016/s0006-3207(01)00056-8 maciejewski, k. & kerley, g.i.h., 2014, ‘elevated elephant density does not improve ecotourism opportunities, convergence in social and ecological objectives’, ecological applications 24, 920–926. https://doi.org/10.1890/13-0935.1 mackey, r.l., page, b.r., duffy, k.j. & slotow, r., 2006, ‘modelling elephant population growth in small, fenced, south african reserves’, south african journal of wildlife research 36, 33–43. manly, b.f.j., mcdonald, l.l., thomas, d.l., mcdonald, t.l. & erickson, w.p., 2002, resources selection by animals: statistical design and analysis for field studies, 2nd edn., kluwer academic publishers, boston, ma. marshal, j.p., rajah, a., parrini, f., henley, m., henley, s.r. & erasmus, b.f.n., 2011, ‘scale-dependent selection of greenness by african elephants in the kruger-private reserve transboundary region, south africa’, european journal of wildlife research 57, 537–548. https://doi.org/10.1007/s10344-010-0462-1 mazerolle, m.j., 2016, aiccmodavg: model selection and multimodel inference based on (q)aic(c), r package version 2.0-4, viewed 27 april 2016, from http://www.cran.r-project.org/package=aiccmodavg mcloughlin, p.d., morris, d.w., fortin, d., vander wal, e. & contasti, a.l., 2010, ‘considering ecological dynamics in resource selection functions’, journal of animal ecology 79, 4–12. https://doi.org/10.1111/j.1365-2656.2009.01613.x mcshane, t.o., hirsch, p.d., trung, t.c., songorwa, a.n., kinzig, a., monteferri, b. et al., 2011, ‘hard choices: making trade-offs between biodiversity conservation and human well-being’, biological conservation 144, 966–972. https://doi.org/10.1016/j.biocon.2010.04.038 mueller, t. & fagan, w.f., 2008, ‘search and navigation in dynamic environments – from individual behaviors to population distributions’, oikos 117, 654–664. https://doi.org/10.1111/j.0030-1299.2008.16291.x nellemann, c., moe, s.r. & rutina, l.p., 2002, ‘links between terrain characteristics and forage patterns of elephants (loxodonta africana) in northern botswana’, journal of tropical ecology 18, 835–844. https://doi.org/10.1017/s0266467402002547 northrup, j.m., hooten, m.b., anderson, c.r., jr. & wittemyer, g., 2013, ‘practical guidance on characterizing availability in resource selection functions under a use-availability design’, ecology 94, 1456–1463. https://doi.org/10.1890/12-1688.1 o’connor, t.g., goodman, p.s. & clegg, b., 2007, ‘a functional hypothesis of the threat of local extirpation of woody plant species by elephant in africa’, biological conservation 136, 329–345. https://doi.org/10.1016/j.biocon.2006.12.014 osborn, f.v., 2003, ‘seasonal influence of rainfall and crops on home-range expansion by bull elephants’, pachyderm 35, 53–59. owen-smith, n., kerley, g.i.h., page, b., slotow, r. & van aarde, r.j., 2006, ‘a scientific perspective on the management of elephants in the kruger national park and elsewhere’, south african journal of science 102, 389–394. paley, r.g.t. & kerley, g.i.h., 1998, ‘the winter diet of elephant in eastern cape subtropical thicket, addo elephant national park’, koedoe 41, 37–45. https://doi.org/10.4102/koedoe.v41i1.244 panzacchi, m., van moorter, b., strand, o., saerens, m., kivimäki, i., st. clair, c.c. et al., 2016, ‘predicting the continuum between corridors and barriers to animal movements using step selection functions and randomized shortest paths’, journal of animal ecology 85, 32–42. https://doi.org/10.1111/1365-2656.12386 pettorelli, n., ryan, s., mueller, t., bunnefeld, n., jędrzejewska, b., lima, m. et al., 2011, ‘the normalized difference vegetation index (ndvi): unforeseen successes in animal ecology’, climate research 46, 15–27. https://doi.org/10.3354/cr00936 pettorelli, n., vik, j.o., mysterud, a., gaillard, j.-m., tucker, c.j. & stenseth, n.c., 2005, ‘using the satellite-derived ndvi to assess ecological responses to environmental change’, trends in ecology and evolution 20, 503–510. https://doi.org/10.1016/j.tree.2005.05.011 pittiglio, c., skidmore, a.k., van gils, h.a.m.j. & prins, h.h.t., 2012, ‘identifying transit corridors for elephant using a long time-series’, international journal of applied earth observation and geoinformation 14, 61–72. https://doi.org/10.1016/j.jag.2011.08.006 r core team, 2016, r: a language and environment for statistical computing, r foundation for statistical computing, vienna, austria, version 3.2.4, viewed 13 april 2016, from http://www.r-project.org redfern, j.v., grant, r., biggs, h. & getz, w.m., 2003, ‘surface-water constraints on herbivore foraging in the kruger national park, south africa’, ecology 84, 2092–2107. https://doi.org/10.1890/01-0625 roever, c.l., van aarde, r.j. & leggett, k., 2012, ‘functional responses in the habitat selection of a generalist megaherbivore, the african savannah elephant’, ecography 35(11), 972–982. https://doi.org/10.1111/j.1600-0587.2012.07359.x rozen-rechels, d., van beest, f.m., richard, e., uzal, a., medill, s.a. & mcloughlin, p.d., 2015, ‘density-dependent, central-place foraging in a grazing herbivore: competition and tradeoffs in time allocation near water’, oikos 124, 1142–1150. https://doi.org/10.1111/oik.02207 sanparks, 2008, addo elephant national park management plan, sanparks, skukuza, south africa. sanparks, 2011, animal abundances in parks – 2010/2011, sanparks, skukuza, south africa. sappington, j.m., longshore, k.m. & thompson, d.b., 2007, ‘quantifying landscape ruggedness for animal habitat analysis: a case study using bighorn sheep in the mojave desert’, journal of wildlife management 71, 1419–1426. https://doi.org/10.2193/2005-723 shannon, g., mackey, r.l. & slotow, r., 2013, ‘diet selection and seasonal dietary switch of a large sexually dimorphic herbivore’, acta oecologica 46, 48–55. https://doi.org/10.1016/j.actao.2012.10.013 shannon, g., matthews, w.s., page, b.r., parker, g.e. & smith, r.j., 2009, ‘the affects of artificial water availability on large herbivore ranging patterns in savanna habitats: a new approach based on modelling elephant path distributions’, diversity and distributions 15, 776–783. https://doi.org/10.1111/j.1472-4642.2009.00581.x shannon, g., page, b., slotow, r. & duffy, k., 2006, ‘african elephant home range and habitat selection in pongola game reserve, south africa’, african zoology 41, 37–44. https://doi.org/10.3377/1562-7020(2006)41[37:aehrah]2.0.co;2 slotow, r., garaï, m.e., reilly, b., page, b. & carr, r.d., 2005, ‘population dynamics of elephants re-introduced to small fenced reserves in south africa’, south african journal of wildlife research 35, 23–32. smit, i.p.j., grant, c.c. & whyte, i.j., 2007a, ‘do artificial water holes influence the way herbivores use the landscape? herbivore distribution patterns around rivers and artificial surface water sources in large african savanna park’, biological conservation 136, 85–99. https://doi.org/10.1016/j.biocon.2006.11.009 smit, i.p.j., grant, c.c. & whyte, i.j., 2007b, ‘landscape-scale sexual segregation in the dry season distribution and resource utilization of elephants in kruger national park, south africa’, diversity and distributions 13, 225–236. https://doi.org/10.1111/j.1472-4642.2007.00318.x tambling, c.j., minnie, l., adendorff, j. & kerley, g.i.h., 2013, ‘elephants facilitate impact of large predators on small ungulate prey species’, basic and applied ecology 14, 694–701. https://doi.org/10.1016/j.baae.2013.09.010 therneau, t., 2015, ‘a package for survival analysis in s. r package version 2.38’, viewed 13 april 2016, from http://www.cran.r-project.org/package=survival therneau, t.m. & grambsch, p.m., 2000, modeling survival data: extending the cox model, springer, new york. van aarde, r., ferreira, s., jackson, t., page, b., de beer, y., gough, k. et al., 2008, ‘elephant population biology and ecology’, in r.j. scholes & k.g. mennell (eds.), elephant management: a scientific assessment of south africa, pp. 84–145, witwatersrand university press, johannesburg, south africa. vrm, 2012, vector ruggedness measure (terrain ruggedness) tool, viewed 22 june 2015, from http://www.arcgis.com/home/item.html?id=9e4210b3ee7b413bbb1f98fb9c5b22d4 wall, j., douglas-hamilton, i. & vollrath, f., 2006, ‘elephants avoid costly mountaineering’, current biology 16, 527–529. https://doi.org/10.1016/j.cub.2006.06.049 wall, j., wittemyer, g., klinkenberg, b., lemay, v. & douglas-hamilton, i., 2013, ‘characterizing properties and drivers of long distance movements by elephants (loxodonta africana) in the gourma, mali’, biological conservation 157, 60–68. https://doi.org/10.1016/j.biocon.2012.07.019 whitehouse, a.m. & schoeman, d.s., 2003, ‘ranging behaviour of elephants within a small, fenced area in addo elephant national park, south africa’, african zoology 38, 95–108. wilson, r.r., horne, j.s., rode, k.d., regehr, e.v. & durner, g.m., 2014, ‘identifying polar bear resource selection patterns to inform offshore development in a dynamic and changing arctic’, ecosphere 5(10), 136. https://doi.org/10.1890/es14-00193.1 wittemyer, g., polansky, l., douglas-hamilton, i. & getz, w.m., 2008, ‘disentangling the effects of forage, social rank, and risk on movement autocorrelation of elephants using fourier and wavelet analyses’, proceedings of the national academy of sciences of the united states 109, 19108–19113. https://doi.org/10.1073/pnas.0801744105 zuur, a.f., ieno, e.n., walker, n.j., saveliev, a.a. & smith, g.m., 2009, mixed effects models and extensions in ecology with r, springer, new york. appendix 1: environmental data description for addo elephant national park figure 1-a1: addo elephant national park features an aseasonal climate. (a) rainfall (mm) during the study period (06 march 2010 – 21 march 2013). rainfall records are aggregated to 16-day periods matching the normalised difference vegetation index moderate resolution imaging spectroradiometer composites. (b) mean normalised difference vegetation index (solid line) ± 1 standard deviation (dashed lines) during the study period. the mean and standard deviation of normalised difference vegetation index were calculated across the three elephant-accessible sections of addo elephant national park: main camp, colchester and nyathi. (c) while the rainfall and normalised difference vegetation index patterns are aseasonal, temperature follows more regular annual patterns. figure 2-a1: covariate data used as inputs into step selection function models for resource selection by female elephants in addo elephant national park, south africa. covariates modelled included (a) normalised difference vegetation index, (b) distance to artificial water, (c) slope and (d) terrain ruggedness. figure 3-a1: covariate values differed between the main camp – colchester and nyathi sections for (a) normalised difference vegetation index (wilcoxon rank-sum test w = 3 517 400, p < 0.001), (b) distance to artificial water (w = 1 270 300, p < 0.001), (c) slope (w = 2 636 900, p < 0.001) and (d) terrain ruggedness (w = 1 871 300, p < 0.001). appendix 2: elephant location data summary table 1-a2: descriptive statistics of daily elephant location data recorded in the morning (06:00), midday (12:00) and afternoon (16:00). for each parameter, the mean ± s.d. is reported for seven female elephants in addo elephant national park, south africa. table 2-a2: descriptive statistics of daily elephant location data recorded in the morning (06:00), midday (12:00) and afternoon (16:00). for each parameter, the mean ± s.d. is reported for the six female elephants occupying the main camp – colchester section of addo elephant national park, south africa. appendix 3: availability sample size sensitivity analysis following the guidance of northrup et al. (2013), we conducted a sensitivity analysis on the effects of the size of the availability sample on our conditional logistic regression (clr) coefficient estimates. for each used elephant location, a candidate set of 100 potentially available locations was drawn as described in the main text. we tested seven availability ratios ranging from 1 to 50 available locations per used location (availability ratios = 1, 3, 5, 10, 20, 35 and 50). we fit clr models using the full time-interacted model (model 30 in table 1) with available points randomly selected from the candidate set. this was repeated 100 times for each availability ratio, and the mean coefficient estimate and 95% simulation envelope were calculated. the sensitivity analysis was repeated for each of the three daily elephant data sets (morning [06:00], midday [12:00] and afternoon [16:00]). simulation results indicated relatively small changes in regression coefficient estimates across varying availability ratios (figure 1-a3 – figure 3-a3). simulation envelopes grew narrower as the availability ratio increased and coefficient estimates tended to stabilise at around 20 available locations per used location. coefficient estimates for the time-interacted covariates (not shown here) showed similar patterns to the time-independent coefficients. in light of these results, we used an availability ratio of 20 available locations for each used location in subsequent analyses. figure 1-a3: coefficient estimates (black points) and 95% simulation envelopes (solid lines) for (a) the normalised difference vegetation index, (b) distance to water, (c) slope and (d) terrain ruggedness, calculated from 100 conditional logistic regression models fit to varying ratios of available to used elephant locations recorded at 06:00. figure 2-a3: coefficient estimates (black points) and 95% simulation envelopes (solid lines) for (a) normalised difference vegetation index, (b) distance to water, (c) slope and (d) terrain ruggedness, calculated from 100 conditional logistic regression models fit to varying ratios of available to used elephant locations recorded at 12:00. figure 3-a3: coefficient estimates (black points) and 95% simulation envelopes (solid lines) for (a) normalised difference vegetation index, (b) distance to water, (c) slope and (d) terrain ruggedness, calculated from 100 conditional logistic regression models fit to varying ratios of available to used elephant locations recorded at 16:00. appendix 4: elephant predicted use animations figure 1-a4: animation of predicted space use by female elephants in the morning (06:00) in addo elephant national park, south africa. predicted use reflects step selection function model predictions to the elephant-accessible main camp – colchester section of addo elephant national park for 16-day periods from 06 march 2010 to 21 march 2013, corresponding to the moderate resolution imaging spectroradiometer composites used for analysis (see the main text for details). figure 2-a4: animation of predicted space use by female elephants at midday (12:00) in addo elephant national park, south africa. predicted use reflects step selection function model predictions to the elephant-accessible main camp – colchester section of addo elephant national park for 16-day periods from 06 march 2010 to 21 march 2013, corresponding to the moderate resolution imaging spectroradiometer composites used for analysis (see the main text for details). figure 3-a4: animation of predicted space use by female elephants in the afternoon (16:00) in addo elephant national park, south africa. predicted use reflects step selection function model predictions to the elephant-accessible main camp – colchester section of addo elephant national park for 16-day periods from 06 march 2010 to 21 march 2013, corresponding to the moderate resolution imaging spectroradiometer composites used for analysis (see the main text for details). figure 4-a4: animation of predicted space use by female elephants across addo elephant national park, south africa. predicted use reflects midday (12:00) step selection function model extrapolations to the full addo elephant national park, including areas that are currently not accessible to elephants. predictions represent 16-day periods from 06 march 2010 to 21 march 2013, corresponding to the moderate resolution imaging spectroradiometer composites used for analysis (see the main text for details). abstract introduction protected areas in tanzania tourism in tanzania methodology results discussion conclusion acknowledgements references about the author(s) anna spenceley school of tourism & hospitality, university of johannesburg, south africa andrew rylance school of tourism & hospitality, university of johannesburg, south africa sadiki l. laiser tanzania wildlife management authority, ministry of natural resources and tourism, tanzania citation spenceley, a., rylance, a. & laiser, s.l., 2017, ‘protected area entrance fees in tanzania: the search for competitiveness and value for money’, koedoe 59(1), a1442. https://doi.org/10.4102/koedoe.v59i1.1442 original research protected area entrance fees in tanzania: the search for competitiveness and value for money anna spenceley, andrew rylance, sadiki l. laiser received: 20 sept. 2016; accepted: 09 jan. 2017; published: 30 mar. 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract user fees charged by tanzania’s game reserves (gr) and wildlife management areas (wmas) have not changed since 2008. although previous research has been done on visitors’ willingness-to-pay to enter national parks in tanzania, none has been conducted on grs and wmas. this article assesses the entrance fees in grs and wmas, by comparing them with equivalent fees charged in tanzania (at national parks and the ngorongoro crater) and also with regional protected areas in botswana, kenya, namibia, south africa and zimbabwe. based on 28 semi-structured interviews with key stakeholder institutions working on tourism and conservation and more than 50 online survey responses from tanzanian tourism operators, the research reviews local opinion and issues relating to adjusting current entrance fees. the article considers that while one objective for generating revenue from entrance fees is for conservation management, it is difficult to establish appropriate fees where there are gaps in knowledge about existing levels of visitation, tourism revenue and associated management costs. conservation implications: this article has implications for protected area management practices, as it provides information on processes by which managers can review and revise entrance fee values. introduction in some countries, there has been a determined shift from using government funding for the capital and operational needs of protected areas (pas), towards charging fees for using services and facilities provided to visitors (e.g. canada: van sickel & eagles 1998), tour operators and investors (e.g. south africa: spenceley 2004). in general, this shift has been caused by changes in government priorities for the use of public funds. user fees provide a mechanism for pa authorities to capture financial benefits of tourism that often accrue primarily to the private sector and which can be used to manage over-used or damaged areas (kibira 2014) and also for the general management of a pa. approaches to establishing user fees in order to finance the management of pas have been extensively documented. perhaps one of the most extensively used approaches to researching entrance fees is contingent valuation approaches with willingness-to-pay (wtp) surveys (e.g. asafu-adjaye & tapsuwan 2008; baral, stern & bhattarai 2008; barnes, schier & van rooy 1997; bruner et al. 2015; greiner & rolfe 2004; kahn 2009; kibira 2014; mmopwlwa, kgathi & molefhe 2007; moran 1994; reyisdottir, song & agrusa 2008; richer & christensen 1999; schultz, pinazzo & cifuentes 1998; thur 2010; walpole, goodwin & ward 2001; wang & jia 2012). other approaches to establishing recreational value and fees include the travel cost method (e.g. chen et al 2004; flemming & cook 2008; herath & kennedy 2004; turpie & joubert 2001) and price differentiation (e.g. chase et al. 1998; krug, suich & haimbodi 2002). more often than not, the wtp studies have found travellers are (1) willing to pay to visit pas and (2) are willing to pay more than the established fee. for example, in annapurna conservation area, nepal, a wtp study found that on average visitors were willing to pay usd69, rather than the actual entrance fee of usd27 (baral et al. 2008). similarly, in komodo national park in indonesia, an analysis of financial data and a wtp study with visitors found that although only 6.9% of park fees were recovered, tourists were willing to pay more than 10 times the current entrance fee (walpole et al. 2001). wtp analysis was done by dikgang and muchapondwa (2016) found that the conservation fees in the kgalagadi transfrontier park could be increased by up to 115% – which would almost double the current revenue. the pattern continues in marine pas too: in bonaire national marine park in the netherlands antilles, a survey found the average wtp for annual access ranged from usd61 to usd134, although the actual fee at that time was only usd10 (thur 2010). furthermore, it was predicted that doubling the usd10 user fee in bonaire would have virtually no impact on visitation rates (thur 2010). in tanzania, previous studies have been conducted on visitors wtp to visit national parks (e.g. bruner et al. 2015; kibira 2014). based on interviews with 3263 visitors, bruner et al. (2015) found that non-residents would be willing to pay substantially higher fees in the serengeti national park and that this would not seriously reduce the level of visitation. it was predicted that phasing-in a usd60 increase in the serengeti conservation fee over several years could raise an additional usd14.8 million by 2020 (equivalent to increasing the park’s revenue by 57%). however, the study also established that price increases were park specific. increased fees were not, for example, recommended at kilimanjaro national park (because of a perception that fees were already too high) and only modest increases be applied to lake manyara and arusha national parks to keep pace with inflation (e.g. usd5). bruner et al.’s (2015) assessment supports earlier work by kibira (2014) on the serengeti. kibira (2014) used contingent valuation to generate experimental data assessing the effects of changes in entrance fees to the serengeti national park. kibira (2014) interviewed a random sample of 539 international tourists who visited the serengeti national park in 2013 and 2014 during the peak tourist season. the study found that visitation demand to the serengeti is highly inelastic, which he attributes to the uniqueness of the park. kibira (2014) suggests that tanzanian national parks (tanapa) should be charging at least usd67 per day and up to usd185 if they want to maximise their revenues for park management. although these studies exist for national parks, no study was identified that had reviewed pricing for game reserves (grs) or wildlife management areas (wmas) in tanzania. lindberg (2001) highlights that although there is no single ‘correct’ system for charging fees, often management decisions are only based on the objective of revenue generation. sometimes the decisions are made with little or no consultation with affected stakeholders, notably the tourism industry and local communities. lindberg (2001) warns that having narrow objectives and weak consultation can lead to unintended effects, such as sometimes a reversal of fee decisions by the pa authority, in response to stakeholder pressure. protected areas in tanzania protected wildlife areas in tanzania cover a total area of 246 260.6 km², covering 26.6% of the country’s land area. these include 16 national parks, 28 grs, 44 game-controlled areas (gcas) and 38 wmas. historically, the ministry of natural resources and tourism (mnrt)’s wildlife division (wd) has been responsible for the management of wildlife resources outside tanapa and the ngorongoro conservation area authority (ncaa). however, the wildlife conservation act of 2009 directed the mnrt to establish the tanzanian wildlife management authority (tawa), which would assume responsibility from the wd for wildlife management in the country’s grs, wmas and gcas. following the official launch of tawa in may 2014, its organisational framework was outlined in the wildlife conservation (the tanzania wildlife management authority) establishment order, 2014. the order describes tawa’s proposed directorate of wildlife utilisation, which would be responsible for activities including issuing permits and licenses in conservation areas, the development of tourism products, identification and diversification of economic potential from wildlife, and also for managing a business with the private sector. by contrast to the wd, tawa is to be a self-financed autonomous organisation, with the mandate to hold back 80% of its revenues to be administered and used according to own demands and priorities. it is important to highlight that financing this substantial pa system is extremely challenging. to illustrate, it is estimated that the financial requirements for both recurrent and capital budgets for the pa system costs usd49 million in 2016/2017 (mnrt 2015). furthermore, predictions suggest that tawa’s financing gap will be in the region of usd12.6 million in the 2016/2017 financial year, with photographic tourism predicted to contribute usd5.2 million in revenue, trophy hunting and trade in wild animals to contribute usd18.4 million and usd12.5 million to be provided by government for salaries (mnrt 2015). a reorientation of the pa entrance and activity fees had been identified by the government of tanzania as an option for ensuring that their pas are competitively priced relative to their competitors. boosting revenues through tourism could help close the financing gap required for continued active conservation management. to support the mnrt in its process of developing tawa, a study was commissioned by the deutsche gesellschaft für internationale zusammenarbeit (giz) programme ‘sustainable management of natural resources’ to evaluate the potential for non-consumptive tourism to generate revenue for conservation and management. a key focus of this study was to review potential changes in the fee structure of the pas. notably, the fees charged for grs, gcas and wmas fees have remained unchanged since the wildlife conservation (non-consumptive wildlife utilisation) regulations, 2008. just in terms of inflation, usd1 in 2008 had the same buying power as usd1.11 did in 2015 (cpi, undated). therefore, this research presented a strategic opportunity to review prices of entrance and activity fees that would be competitive with other pas in tanzania and across southern africa. tourism in tanzania the number of international arrivals in tanzania has grown dramatically from about 500 000 in 2000 to over 1.14 million visitors in 2014 (world bank 2015). the tourism sector generates the bulk of export revenues for the country, typically exceeding minerals. it provides a reliable source of revenue to the government and generates direct employment to over 400 000 people (world bank 2015). in 2013 tourism was estimated to account for about 9.9% of gdp (equivalent to an amount of usd4 billion in direct and indirect contributions) (world bank 2015). the majority of international arrivals are travelling for leisure, recreation and holiday (81%), followed by visiting friends and relatives (8%), business (6%), transit (2%) and other reasons (3%) (mnrt 2015). the average length of stay in tanzania in 2014 was 10 days, a duration that has been stable since 2012 (mnrt 2015). the average per night expenditure for a participant on a package tour is more than double that of a non-package tourist, at usd277 in 2014. however, there has been a dramatic decline in the level of expenditure by both types of travellers since 2012: by 27% for package tourists and 50% for non-package tourists (mnrt 2015). the reasons for this decline are not known. visitation to national parks and to the ngorongoro crater is reported in the tourism statistical bulletin, but visitation to grs and wmas is not. the most visited pas in the country are the ngorongoro crater (37.8% of recorded entries), followed by serengeti national park (24.9% of entries), tarangire (11.2% of entries) and arusha (10.8% of entries) (adapted from mnrt tourism division 2015). it is estimated the 46% of international tourists experience a wildlife activity, compared to 26% with a beach holiday component and 11% doing hunting and fishing (sanches-pereira & muwanga 2015). the purpose of this article is to describe the use of regional price benchmarking and market research with tourism stakeholders to investigate options to adapt fees charged at grs and wmas. this article focuses particularly on entrance fees and a discussion of other activity and service fees (i.e. camping fees, activity fees, commercial operator fees, concession fees, motor vehicle or boat or aircraft and balloon fees) can be found elsewhere (spenceley 2016). methodology a literature review was undertaken relating to previous research on tourism, conservation and pas in tanzania. documents were obtained from donor and development agencies working in tanzania, from stakeholders consulted and from an online search for academic research and grey literature. during this process, it was realised that no detailed information was available on existing levels of visitation, revenues accrued or the management costs of grs or wmas in tanzania. for the stakeholder consultation process on fees for grs and wmas, two approaches were used: a semi-structured questionnaire used for face-to-face and telephone interviews and also a structured online survey tool. the semi-structured questionnaire tool was developed and implemented through a series of face-to-face and telephone interviews with representatives of 28 institutions working on tourism and conservation during october 2015. these included government departments (3 interviews), pa authorities (5 interviews), donor agencies (6 interviews), non-governmental organisations (2 interviews), private sector associations (3 interviews) and accommodation facilities in pas (8 interviews). the stakeholders were selected through a combination of suggestions made by giz and a review of relevant institutions in tanzania and referrals by interviewees (i.e. snowball sampling). the questionnaire included 37 questions that enquired about which pa systems were most competitive and comparable to tanzania’s, user fee options for grs and wmas and processes that should be applied to adjust them. the online survey tool was developed to collate opinions on entrance fees within tanzania’s grs and wmas. the timeframe and resources available for the study meant that it was not feasible to interview visitors at each pa, and therefore, the survey was designed for tour operators working in pas. given their knowledge of their client base and the tourism sector as a whole, it was considered that their opinions provided a useful proxy for visitor preferences, though this is a limitation of the present study. this questionnaire simply presented the current prices charged by grs and wmas and asked respondents to indicate whether the price (which was shown) should remain the same or be changed to a different value. this approach was judged to be more practical than a dichotomous choice approach because such a diverse range of fees was listed in the wildlife conservation (non-consumptive wildlife utilisation) regulations, 2008 that needed to be investigated for grs and wmas separately (i.e. entry fees; commercial photography fees; camping fees; vehicle fees [e.g. cars, boats, rafts, canoes]; night drives; fishing; professional guide fees; aircraft and balloon fees). providing an option for open responses (as by greiner & rolfe 2004) ensured that the online questionnaire was not excessively long or complicated and improved the likelihood of a good response rate. the survey was distributed to operators by the country’s three main tourism associations: tanzanian confederation of tourism, the hotels association of tanzania and the tanzanian association of tour operators to their members. the questionnaire was available online between 23 october and 08 november 2015, and 50 responses were received during that time. for the regional price benchmarking process, information on current pa fees were collected from pa authority websites. these were converted into usd equivalent values for comparison. prices were collated for pas in tanzania (i.e. national parks, the ngorongoro conservation area, grs and wmas). prices were also reviewed for a selection of southern african pa agencies in east and southern africa that have similar and competitive wildlife products to tanzania, as suggested by stakeholders during the stakeholder consultation process (see above). these included national parks and grs, but there was no publically available data identified for wma-equivalent areas. the countries reviewed comprised botswana, kenya, namibia, south africa and zimbabwe. in south africa, information was also used by two provincial conservation agencies that manage grs (i.e. ezemvelo kwazulu-natal wildlife and cape nature). results national and regional benchmarking of entrance fees at the time of this study, the entrance fees charged by the wd for non-citizen adults at selous, ikorogo, grumeti and maswa grs was usd50, plus an additional usd15 conservation fee for tourists staying outside the reserves. for tourists staying inside the grs, the entrance fee is the same, but the conservation fee increased to usd25. this is the same entrance fee charged by tanapa for the serengeti. by contrast, the entrance fees charged in wmas are the lowest in the country, at usd10 for adults (see table 1). entrance fees are valid for 24 h in the wd and tanapa’s pas, and tanapa’s and the ncaa fees are valid for a single entry. table 1: entrance fees charged by different protected areas in tanzania (usd equivalent). comparison with entrance fees charged for grs and wmas with pas in other southern african countries demonstrates that entrance fees vary both in price and also the way in which they are charged. variations include charging a flat fee for the entrance and activities, to different rates depending on where the visitor is domiciled, to a fee that depends on the means of travel. table 2 shows that the value of entrance fees charged for non-citizen adults at grs are the second highest in the region, surpassed only by kenya’s national parks, at usd80. the non-citizen adult entrance fee for wmas is the same as the cheapest national park in zimbabwe and more than the fee for adult non-citizens in namibia (table 3). all but one of the park agencies reviewed charged different entrance fees for different reserves, while botswana charges the same entrance fee for all pas. table 2: entrance fees charged by different protected areas in africa (usd equivalent). table 3: current and proposed entrance fees for game reserves and wildlife management areas in usd equivalent values. the fee categories relating to origin used by the wd are the same as applied by tanapa and the ncaa (i.e. adult, child, citizen, non-citizen). however, while tawa’s citizen prices apply to citizens of tanzania, tanapa’s citizen rates are applied more broadly, to east african expatriates and residents. similar categorisations are used by other pa authorities in east and southern africa: south african national parks (sanparks) uses rates for their citizens and residents, southern african development community (sadc) nationals and other nationalities. kwazulu-natal (kzn) wildlife in south africa has rates for south african and sadc residents. kenya wildlife service charges the same rates for citizens of kenya, uganda, tanzania, rwanda and burundi and higher rates for residents and non-citizens. botswana has rates for citizens of botswana, residents of botswana and non-residents. namibia has categories for citizen and residents, sadc residents and foreign residents. cape nature in south africa does not have different rates for people of different origins. instead, they have higher ‘peak period’ rates and lower rates for ‘off-peak’ times. the wd and ncaa charge adult rates for people aged 18 years and over, while tanapa charges adult fees for people aged 16 years and over. the wd’s children’s category is from 5 to 17 years, while tanapa’s are from 5 to 16 years. in all cases, infants below the age of 5 years are free of charge. the categories used for different age classes in other countries in east and southern africa vary as to whether adult rate are charged from 12 years (sanparks), 13 years (kzn wildlife, cape nature, zimbabwe), 16 years (namibia) or 18 years (kenya, botswana). consultation process feedback on changes to existing fees from the online survey, on average, those proposing a change in fees in selous, ikorogno, grumeti and maswa grs suggested a decrease in the rates for non-citizen adults (to usd41) and children (to usd21) compared to a slight increase for citizen adults (usd2.3) and children (to usd1.6). for other grs, the average suggested fees were not very different from the current fees. in wmas, the average entrance fee proposed was roughly half the current rate charged for non-citizens. however, in most cases, responses implied that respondents were satisfied with the existing fee (i.e. an alternative value was not suggested). further suggestions from stakeholders from the semi-structured interviews and online survey in relation to entrance fees included suggestions to: establish an all-inclusive entrance fee that covers entry and all activities (e.g. game drives, walking, camping, night drives) to make them better value for money. a single fee is also much easier to implement and monitor than multiple fees, particularly where there are few field personnel to check permits. standardise fees with tanapa, including discounted rates for east african residents, with tanzanian residents paying the same as citizens. explore options for seasonal rates, with discounted rates for low season periods and higher rates for peak times to maximise revenue and encourage more visitation in quite periods. maintain entrance fees as valid for 24 h, but with the option for multiple entries during that time. this provides greater flexibility and value for money for visitors. make the fees in grs more competitive. the grs are currently very expensive (e.g. particularly selous, with an additional conservation fee) given the lack of infrastructure, poaching levels, weak services in selous and lower prices in competing destinations in tanzania’s national parks, the ngorongoro crater and also in pas across southern africa. many wmas have degraded wildlife environments and cannot compete with national parks. they should have a single conservation fees to cover the entrance fee and activities undertaken in the areas. discussion the study found that current non-citizen adult entrance fees are usd30 – usd50 in grs and usd10 in wmas (excluding conservation fees). the assessment identified that gr entrance fees are high when compared to those charged in tanapa and the ngorongoro crater. furthermore, when compared with entrance fees in other east and southern african countries, the gr entrance fees are second highest in the region and are exceeded only by kenya’s entrance fee of usd80 per day. even the entrance fees for wmas, which have few tourism assets and services, are more expensive than some national parks in namibia and zimbabwe. as a number of grs and wmas are a greater distance from popular tourism routes, any increase in entrance fees may act as a negative incentive as travel costs are already relatively higher than other tanzanian wildlife destinations. the categories of entrance fees applied by the wd, tanapa and ncaa are different. they each have different age categories for adult, child and infant entrances and have different ways of charging for citizens, residents and non-residents (e.g. tanapa has a rate for east african residents), which creates a complicated overall picture and adds complexity to the revenue collection and reporting system. in destinations where there are clusters of pas operated by different pa authorities and villages, tourists may be charged multiple entrance fees on the same days. this escalates the cost for tourists who wish to explore natural and cultural attractions in a particular area and constrains their movement. there are no discounted rates or loyalty cards available for multiple entries to the grs and wmas over longer periods of time (e.g. weekly, monthly or annual passes), as there are in other african countries. for example, in south africa, sanparks has an annual ‘wild card’ fee for individuals, couples and families, with different rates for different ‘clusters’ of pas and for citizens or residents, residents of the sadc and international visitors. similarly, kenya wildlife service has annual passes for adults and children, marine annual passes, tour driver and corporate passes. to illustrate the benefit of the loyalty cards, sanparks generated usd5.4 million in revenue from its wild card in the financial year ending march 2015 (sanparks annual report, 2014/2015). an important element to review in relation to user fees is value for money. in comparing the quality of the tourism experience in grs and wmas with national parks in the country or the ngorongoro crater, some of the disadvantages identified by stakeholders include that they tend to have less wildlife, are often remote and more difficult to access, have weaker supporting infrastructure, and have fewer, lower capacity employees. however, advantages of grs and wmas include that they have more flexibility in terms of the type of tourism activities they can offer (e.g. night drives), low tourist numbers and also that they provide the only opportunity for expansion of accommodation within pas in tanzania, as the national parks and crater are at capacity. anecdotally, the only gr that was highlighted by stakeholders as having the potential to generate equivalent revenues to a national park was the selous (however, this could not be confirmed, because revenues for specific pas were not available). one of the key challenges faced in this analysis was that it was not possible to establish how much additional revenue fee changes would generate for tawa because data on existing levels of visitation and tourism revenue were not available. furthermore, data on associated management costs for each gr and wma were not available to establish the extent to which existing or proposed revenues would meet them. these data challenges are not unique to tanzania and have also been described for conservation areas in mozambique (rylance 2017). before making policy decisions to adjust entrance fees in grs and wmas in tanzania, it is recommended that further research be undertaken. this should include (1) a comprehensive wtp study for an all-inclusive entrance fee to cover entry and all activities, (2) collaborate with tanapa and ncaa to design and establish single destination-based entrance fees for clusters of pas and consistent age fee categories for adults, children and infants, (3) research the viability of a loyalty card for multiple entries over longer periods of time (e.g. a 1–2 week pass for international arrivals and monthly or annual passes for citizens and residents), (4) extend the citizen rate to citizens of the east african community (as done in kenya and by tanapa) and establish a discounted rate for residents of tanzania, (5) fees should be reviewed every 2 years and informed by market research with tourists and operators (including wtp studies and regional benchmarking on pa fees). because the facilities and wildlife within grs and wmas are not as attractive to visitors as in national parks and the ngorongoro crater, it is unlikely that moderate changes to the prices of entrance fees for grs and wmas would affect visitation to national parks and the crater. conclusion at this stage, it should be recognised that the acceptance and effects of fees may depend on the historical, political, legal, economic and social context – including the control and use of funds raised (buckley 2003). for example, research has found that tourists are willing to pay more for park entrances if that money is used to protect the environment (baral et al. 2008). there are also differences in the types of services that people are willing to pay for, or not. to illustrate, bowker, cordell and johnson (1999) found that 80% of visitors surveyed in the united states felt that user fees should be charged to cover at least part of the cost of camping areas, exhibits and boat jetties, but not for picnic areas, historic sites or ablution facilities. our study found that tourism operators were particularly resistant to increases in entrance fees because there was no transparency about the re-investment of existing fees in the parks. it is clear that any decision to adjust fees should be mindful of the implications for tourists and tour operators. on a practical note, tour operators should be advised of future fee changes 2 years in advance, to ensure that they have time to integrate changes into their brochures and travel budgets. it should also be noted that since the benchmarking analysis and survey took place, a value added tax of 18% has been introduced on entrance fees in tanzania. therefore, the charge for the entrance fee and conservation fee for grs has increased to usd76.70 and usd88.50 for a tourist staying outside and inside the gr, respectively. as yet, there is no information on how the market will react to this change, particularly in terms of the price competitiveness with other destinations. lindberg (2001) suggests that if a decision is made to adjust fees following a review of the advantages and disadvantages, fee objectives should be reviewed to determine the fee amount and type. the following potential fee objectives should be considered (lindberg 2001:1–2): cost recovery, which involves generation of sufficient revenue to cover part or all of tourism’s financial costs (e.g. construction and maintenance of a visitor centre) and possibly tourism’s other costs (e.g. ecological damage). generation of ‘profit’, with the excess of revenue over cost being used to finance traditional conservation activities (at the destination or at other sites) or to achieve other objectives. generation of local business opportunities, which may involve low fees in an effort to maximise the number of visitors and/or the earmarking of fees to enhance site or experience quality. provision of maximum opportunities for learning and appreciation of the natural resource, which may also involve low fees. visitor management to reduce congestion and/or ecological damage, which would involve fees high enough to influence visitor behaviour. at the time of writing, this type of review of user fee objectives has not been undertaken in tanzania. the adjustment of fees has been primarily based on the objective of raising revenues to pay for management costs. therefore, this analysis should be used as part of the decision-making process for fee adjustments for the new tawa, but not in isolation from other important data and comprehensive consultative processes. acknowledgements competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions a.s. worked on the research design; undertook the field research, benchmarking research, stakeholder interviews, and analysis; and wrote part of the literature review, discussion and conclusions. a.r. contributed to the research design, literature review, conclusions and discussion. s.l.l. was involved in the assignment design and some stakeholder interviews. references asafu-adjaye, j. & tapsuwan, s., 2008, ‘a contingent valuation study of scuba diving benefits: case study in mu ko similan marine national park, thailand’, tourism management 29(6), 1122–1130. https://doi.org/10.1016/j.tourman.2008.02.005 baral, n., stern, m.j. & bhattarai, r., 2008, ‘contingent valuation of ecotourism in annapurna conservation area, nepal: implications for sustainable park finance and local development’, ecological economics 66(2–3), 218–227. https://doi.org/10.1016/j.ecolecon.2008.02.004 barnes, j.j., schier, c. & van rooy, g., 1997, tourists’ willingness to pay for wildlife viewing and wildlife conservation in namibia, dea research discussion paper, no. 15, march 1997, government of namibia, windhoek. bowker, j.m., cordell, h.k. & johnson, c.y., 1999, ‘user fees for recreation services on public lands: a national assessment’, journal of park and recreation administration 17(3), 1–14. bruner, a., kessey, b., mnaya, j., wakibara, j. & maldonado, j., 2015, tourists willingness to pay to visit tanzania’s national parks: a contingent valuation study, conservation strategy fund discussion paper no. 9, september 2015, conservation strategy fund, washington, dc. buckley, r., 2003, ‘pay to play in parks: an australian policy perspective on visitor fees in public pas’, journal of sustainable tourism 11(1), 56–73. https://doi.org/10.1080/09669580308667193 cape nature, 2013, ‘tariffs’, viewed 10 november 2016, from http://www.capenature.co.za/tariffs/ chase, l.c., lee, d.r., schultze, w.d. & anderson, d.j., 1998, ‘ecotourism demand and differential pricing of national park access in costa rica’, land economics 74(4), 466–482. https://doi.org/10.2307/3146879 chen, w., hong, h., liu, y., zhang, l., hou, x. & raymond, m., 2004, ‘recreation demand and economic value: an application of travel cost method for xiamen island’, china economic review 15(4), 398–406. https://doi.org/10.1016/j.chieco.2003.11.001 cpi, undated, ‘cpi inflation calculator’, viewed 10 november 2016, from http://data.bls.gov/cgi-bin/cpicalc.pl?cost1=1&year1=2008&year2=2015 department of wildlife and national parks, botswana (dwnp botswana), undated, ‘park fees’, viewed 10 november 2015, from http://www.mewt.gov.bw/dwnp/article.php?id_mnu=214 dikgang, j. & muchapondwa, e., 2016, ‘the determination of park fees in support of benefit sharing in southern africa’, tourism economics. https://doi.org/10.1177/1354816616655254 flemming, c.m. & cook, a., 2008, ‘the recreational value of lake mckenzie, fraser island: an application of the travel cost method’, tourism management 29(6), 1197–1205. https://doi.org/10.1016/j.tourman.2008.02.022 government of zimbabwe, 2014, parks and wildlife management authority (tariff of fees) bylaws, 2014, cap, harare, p. 201. greiner, r. & rolfe, j., 2004, ‘estimating consumer surplus and elasticity of demand of tourist visitation to a region in north queensland using contingent valuation’, tourism economics: the business and finance of tourism and recreation 10(3), 317–328. https://doi.org/10.5367/0000000041895076 herath, g. & kennedy, j., 2004, ‘estimating the economic value of mount buffalo national park with the travel cost and contingent valuation models’, tourism economics 10(1), 63–78. https://doi.org/10.5367/000000004773166529 imagine tanzania tourism bureau, 2014, new park fees in the ngorongoro conservation area, viewed 13 august 2014, from https://www.facebook.com/ittbtz/posts/758013970930170 kahn, h., 2009, ‘willingness to pay and demand elasticities for two national parks: empirical evidence from two surveys in pakistan’, environment, development and sustainability 11(2), 293–305. https://doi.org/10.1007/s10668-007-9111-6 kenya wildlife service, undated, conservation fees from 1st january 2015, kenya wildlife service, nairobi. kibira, g., 2014, ‘the economic implications of conservation in serengeti national park on adjacent local communities in tanzania’, phd progress report, school of economics, university of cape town. krug, w., suich, h. & haimbodi, n., 2002, park pricing and economic efficiency in namibia, dea research discussion paper, no. 45, june 2002, government of namibia, windhoek. kzn wildlife, 2014, important information (tariffs 2014–15), kzn wildlife, pietermaritzburg. lindberg, k., 2001, pa visitor fees, overview, cooperative research centre for sustainable tourism, griffith university, nathan, australia. mmopwlwa, g., kgathi, d.l. & molefhe, l., 2007, ‘tourists’ perceptions and their willingness to pay for park fees: a case study of self-drive tourists and clients for mobile tour operators in moremi gr, botswana’, tourism management 28(4), 1044–1056. https://doi.org/10.1016/j.tourman.2006.08.014 moran, d., 1994, ‘contingent valuation and biodiversity: measuring the user surplus of kenyan pas’, biodiversity & conservation 3(8), 663–684. https://doi.org/10.1007/bf00126859 namibia wildlife resorts ltd (nwr) 1998–2015, ‘namibian parks’ daily fees’, viewed 10 november 2015, from http://www.namibweb.com/nwrdailyfees.htm ngorongoro conservation area authority (ncaa), 2013, rules and regulations, ngorongoro conservation area, viewed 20 november 2015, from http://www.ngorongorocrater.go.tz/index.php/2012-09-05-18-04-07 reyisdottir, m., song, h. & agrusa, j., 2008, ‘willingness to pay entrance fees to natural attractions: an icelandic case study’, tourism management 29(6), 1076–1083. https://doi.org/10.1016/j.tourman.2008.02.016 richer, j.r. & christensen, n.a., 1999, ‘appropriate fees for wilderness day use: pricing decisions for recreation on public land’, journal of leisure research 31(3), 269–280. rylance, a., 2017, ‘estimating tourism’s contribution to conservation area financing in mozambique’, tourism and hospitality research 17(1), 24–33. https://doi.org/10.1177/1467358415613119 sanches-pereira, a. & muwanga, m.k., 2015, enhancing linkages between tourism and sustainable agriculture sectors in the united republic of tanzania, united nations conference on trade and development (unctad), geneva, switzerland. schultz, s., pinazzo, j. & cifuentes, m., 1998, ‘opportunities and limitations of contingent valuation surveys to determine national park entrance fees: evidence from costa rica’, environment and development economics 3, 131–149. https://doi.org/10.1017/s1355770x98000072 south african national parks (sanparks), 2015a, ‘conservation fees, 1 november 2015 to 31 october 2016’, viewed 10 november 2015, from http://www.sanparks.org/tourism/tariffs/ south african national parks (sanparks), 2015b, ‘tariffs, 1 november 2015 to 31 october 2016’, viewed 10 november 2015, from http://www.sanparks.org/tourism/tariffs/ spenceley, a., 2004, ‘responsible nature-based tourism planning in south africa and the commercialisation of kruger national park’, in d. diamantis (ed.), ecotourism: management and assessment, pp. 267–280, thomson learning, london. spenceley, a., 2016, assessment of the development of non-consumptive tourism activities under tanzania wildlife management authority (tawa), report to giz: part 0: executive summary, part 1: background and context, part 2: results and analysis. giz, dar es salaam, tanzania. tanzania national parks (tanapa), undated, ‘tariffs: july 2015 – june 2016’, viewed 10 november 2015, from http://www.tanzaniaparks.com/regulations_and_park_fees.html the united republic of tanzania, 2008, the wildlife conservation (non-consumptive wildlife utilization) regulations, 2008, [g.n no. 357 of 28/11/2008]. government of tanzania, dar es salaam, tanzania. thur, s.m., 2010, ‘user fees as sustainable financing mechanisms for marine pas: an application to the bonaire national marine park’, marine policy 34(1), 63–69. https://doi.org/10.1016/j.marpol.2009.04.008 turpie, j. & joubert, a., 2001, ‘estimating potential impacts of a change in river quality on the tourism value of kruger national park: an application of travel cost, contingent, and conjoint valuation methods’, water sa 27(3), 387–398. https://doi.org/10.4314/wsa.v27i3.4983 united republic of tanzania ministry of natural resources and tourism (mnrt), 2015, tanzania wildlife management authority (tawa) financing plan 2015–2020. mnrt, dar es salaam, tanzania. united republic of tanzania ministry of natural resources and tourism (mnrt), tourism division, 2015, the 2014 tourism statistical bulletin, draft. mnrt, dar es salaam, tanzania. van sickel, k. & eagles, p., 1998, ‘budgets, pricing policies and user fees in canadian parks’ tourism’, tourism management 19(3), 225–235. https://doi.org/10.1016/s0261-5177(98)00017-x walpole, m., goodwin, h.j. & ward, k.g.r., 2001, ‘pricing policy for tourism in pas: lessons from komodo national park, indonesia’, conservation biology 15(1), 218–227. https://doi.org/10.1111/j.1523-1739.2001.99231.x wang, p. & jia, j., 2012, ‘tourists willingness to pay for biodiversity conservation and environment protection dalai lake protected area: implications for entrance fee and sustainable management’, ocean and coastal management 62, 24–33. https://doi.org/10.1016/j.ocecoaman.2012.03.001 wikitravel, 2015, ‘ngorongoro conservation area’, viewed 10 november 2015, from http://wikitravel.org/en/ngorongoro_conservation_area world bank, 2015, tanzania’s tourism futures: harnessing natural assets, world bank group report number 96150-tz, september 2015. world bank group, washington, dc. filelist convert a pdf file! koedoe 19: 89-132(1976) reproduction in the zebra mare equus burchell/ ant/quorum from the kruger national park c. l. smuts national parks board of trustees private bag x404 skukuw jj50 abstract reproductive characteristics of burchell's zebra mares are described using data collected from captive and free ranging animals and the reproductive tracts of 310 mares shot during a game cropping campaign. the pubertal interval in zebra mares ranges from age 16 to 22 months, succesful mating occurring for the first time at 23 months of age. full reproductive capacity is attained at three years. zebra mares are seasonally polyoestrous, with an average of 85% of all mating and foaling occurring during the wet summer months (october to march) . the gestation period is about 375 days and peak mating and foaling occurred in december or january for four successive years. the reproductive cycle of the zebra mare is synchronised with photoperiod. rainfall and grass growth flushes, however, have a profound effect on the initiation of mating activity and on irregularities in relative monthly conception percentages. mares have their first postpartum oestrus at eight to nine days with heat lasting approximately six days. foals are normally weaned at 11 months of age (range = nine to 16 months). conception rate varied between 64 and 91% (mean = 79%). rainfall and the concomitant increase in green vegetation was found to affect annual conception rate, with regular summer rainfall being more important than total summer precipitation. graafian follicles attain diameters exceeding 30 mm during prooestrous, all ovulations eventually taking place via the ovarian fossa. multiple ovulations are rare but secondary ovulations occur between the 70th and 1l0th day of pregnancy. the left ovary is more active than the right and trans uterine migrations of ova are common. no twin conceptions or instances of reproductive senescence were recorded. five ovarian abnormalities were found, all incapable of causing permanent sterility. 89 introduction the study on reproduction in the zebra mare forms part of an investigation involving growth, reproduction and population characteristics of burchell's zebra (equus burchelli antiquorum) in the kruger national park (knp) (smuts 1974a). the purpose of the study was to collect general information on reproduction in the mare but particularly on aspects such as puberty and sexual maturity, the seasonal reproductive cycle and factors influencing conception rate. analysis of data has indicated that, as with the stallion (smuts 1976), reproductive characteristics of the burchell's zebra mare are similar to the domestic equids. zebra mares are seasonally polyoestrous with an average of 85% of all mating and foaling occurring during the summer months. material and methods the collection of reproductive data was confined to the central district (5 560 sq. km) of the knp. here free ranging zebra were observed with ii (40'0' + 799) also being captured and retained in captivity for varying periods of time. captive animals provided data on gestation period, the heat cycle and lactation. field observations were limited to counts designed to determine the foaling season while certain behavioural aspects were noted whenever the opportunity presented itself. from july 1969 to june 1973,48 monthly foaling season counts were undertaken between the first and fifth day of each month. these counts were conducted from a vehicle and the routes were designed so as to sample representative sections of the range of both the satara (sedentary) and the eastern boundary (migratory) zebra sub-populations (smuts 1972). during each count 300 to 600 km were covered until a random sample of i 000 zebra had been counted and classified into the following age classes: newly born foals; foals i week to i month old; foals 2 to 12 months old; immatures and sub-adults plus adults. between may 1969 and november 1972 the reproductive tracts of 310 mares, shot during a cropping campaign, were collected. generally the entire reproductive tract was removed for examination with each ovary being marked left or right, excised and placed into bottles containing 10% formalin. fallopian tubes were dissected out and their convolutions freed from the mesosalpinx. their total length, from the uterine to the abdominal ostium, was then determined. in non-pregnant mares uterine and vaginal measurements (fig. 1) were taken. other data routinely collected included: (i) mass of the udder and a description of the state of activity, (ii) whether the mare had a foal at foot and its age, (iii) the side of implantation (pregnant mares), (iv) foetal measurements and sex (smuts 1975), (v) mass of the uterus and its contents. embryonic and foetal gonads were also collected and fixed together with the ovaries of the corresponding mare. once the ovaries and foetal gonads were properly fixed all attachments were dissected away (ovarian 90 ovary fallopian tube uteri ne horn broad ligament body of uterus ---------------. ----\\\\11111111111,\,----------------------l 1____ _ _ ______________ ___ \. \ ~1c ____________________ j':'_ ce:::rnal os o 5 10 !we wi , em " ' supraurethral part of vagi na -·:·~~h~.-----------------------urethral orifice infra-urethral part of vagina (vulva) '----------------------cli tori 5 fig. 1. scale diagram of the reproductive tract of a burchell's zebra mare illustrating the measurements (1 to 8) routinely made on the tracts of non-pregnant mares. these measurements were taken by laying the tract on a smooth, wet surface and allowing it to settle into position. ligament and mesovarium or epididymis) and the mass of each gland determined. ovaries were later sectioned macroscopically so that follicles and corpora lutea could be counted. cutting was done using a large scalpel and sections 2 to 4 mm thick were run in a vertical plane from the free to the attached border of each ovary. the maximum 91 diameter of all visible follicles or corpora lutea were measured and, to avoid double counts, the coagulated liquor folliculi of each follicle was scraped out as it was counted . the ages of all zebra were determined as described by smuts (1972 and 1974b). results social organisation socially burchell's zebra populations consist of coherent family and stallion groups with family groups comprising one adult stallion (leader stallion) and one to five mares and their foals. the aspects are described in more detail by klingel (1965) and smuts (197 4a and 1976). the oestrous cycle the oestrous cycle of burchell's zebra may be divided into the following phases: pro-oestrus the phase immediately preceding oestrus or the preliminary period of refusal and delay. this is characterised by a marked increase in the activity of the reproductive organs (ovaries, uterus and vagina). oestrus the period of acceptance of the male. the mare "stands" for the stallion. metoestrus the phase succeeding oestrus when reproductive activity subsides (the luteal phase). dioestrus the quiescent period between heat cycles. anoestrus the prolonged period of sexual rest during which ovarian function is largely in abeyance (the non-breeding season) and the uterus is small and anaemic (benesch and wright 1957). this is also the normal state of the filly until at adolescence the primary sex organs begin to function. in captive animals oestrus lasted for about six days with copulation occurring on numerous occasions during each day. the foal heat (first postpartum oestrus) occurred eight and nine days after birth (n = 2). supporting these observations was the ovary of an adult mare with a two week old foal at foot, which contained a very recent corpus haemorrhagicum. gestation period, pregnancy and birth one gestation period, recorded from the last observed copulation, was obtained from a captive mare. this gave a period of 396 days. at six to seven months before birth, pregnancy is manifest in the external app earance of the mare, the abdomen being slightly enlarged when viewed posteriorly. after this period foetal growth is rapid (smuts 1975), with the mass of the pregnant uterus and its contents doubling during the last two months of pregnancy (29 to 58 kg). at this stage a mare's 92 movements become ungainly due to the unstabilizing and swaymg motion of the heavy abdominal region. having given birth, mares frequently eat part of the afterbirth . the afterbirth for two mares which foaled in captivity had masses of 3,2 and 3,3 kilogram. mother-joal relationships in burchell's zebra the mother-foal bond is close, particularly during the first few weeks of life. at this early stage the mare assumes a protective role and will not allow any other zebra to approach her foal too closely. during their first few weeks of life, foals are not yet fully imprinted with the mother image and when forcibly separated from their mothers, will follow most large moving objects, even a vehicle. during this period foals characteristically stay very close to their mothers, usually right up against their sides. mares with young foals tend to be more wary than other members of the group and will move off at the first sign of danger. lactation and weaning. in three different captive pregnant mares, without foals at foot, it was found that a visible increase in size of the mammary gland (udder) is initiated at an average of 35 days before parturition, actual results being 31, 36 and 38 days. milk can only be expressed from the mammary gland after parturition. just prior to parturition the udder attains a mass of 2,5 kg while the mass of the inactive udder during the first trimester of the pregnancy varies between about 200 and 500 g in mares without a foal at foot. during the first week after parturition and if the foal is alive and suckling, the mass of the udder decreases to about 2,2 kilogram. as the foal gets older it drinks less and the gland gradually subsides to its involuted state. should a mare lose her foal soon after birth, the udder increases in size as it becomes engorged with milk and in two cases udders were found to have a mass of 4,5 kilogram. in similar cases with captive mares, the udder was found to involute within about five days after loss of the foal. in cases where it was possible to examine the udder of a mare and accurately determine the age of her foal, it was noted that the earliest weaning occurred when the foal was 11 months old. comparing the activity of the mammary gland with the stage of a subsequent pregnancy indicates that lactation and suckling continue, if the foal remains alive, until one or two months prior to parturition. should a mare not conceive soon after foaling she normally continues to lactate until her foal is over a year old. one captive mare continued to lactate until her foal was 16 months old. in this mare, lactation ended six months prior to the birth of her next foal. 93 development of the uterus and vagina uterine and vaginal measurements taken from a series of maiden mares and non-pregnant adult mares are given in table 1 and 2 respectively. last-mentioned measurements were only taken from mares in which complete involution of the uterus had taken place. the actual measurements recorded are illustrated in fig. 1. the length of the fallopian tubes in adult mares varied from 15,5 to 24,5 cm with an average of 19,8 cm (n = 19). the adult dimension is usually attained by 2,5 years of age (n = 7 immature and sub-adult mares). table i ulrrillr and vaginal dimensions jor seven young maiden burchell's zebra mares collected in the central district of the kruger national park ref. no. age (years) on measurement (em) fig. i i j!. 1-j--2 1-j--2 21 3 3+ i le ngth of left 9,3 8,8 7,0 7,0 10,7 9,6 9,8 uterine horn right 9,7 7,6 6,8 6,8 12,5 11,5 11,2 2 width of left 5,0 4,9 4,8 uterin e horn right 3,5 4,8 4,9 3 spread of uterine 16,4 16,0 14,0 12,8 22,5 23,3 20,7 horn 4 width of body of 3,2 5,8 5,1 5,7 6,8 6,6 6,5 uterus 5 length of body of 11 ,6 12,1 6,3 7,6 8,9 17,8 14,1 uterus 6 length of cervix 3,6 4,0 5,4 4,7 6,3 4,2 4,0 7 length of vagina 28,1 17, i 28,3 25,6 30,5 23,8 24,0 and vulva 8 total length (vagina 43,8 37,8 42 ,0 40 ,0 45,8 50,0 45,8 and body of uterus) development of the ovaries as in the domestic horse (hammond and wodzicki 1941), the ovary of adult zebra mares is characteristically bean-shaped. however, during both prenatal and postnatal life a number of clear developmental changes take place in the ovary. in particular these changes concern the gross morphology of the gland with simultaneous histological changes. to illustrate the growth in mass process as it occurs in the foetal gonad, 94 table 2 uterine and vaginal dimensions for adult burchell's z.ebra mares collected in the central district of the kruger national park ref. no on measurement (em) n max. min. mean s.d. fig. i i length of uterine left 10 14,5 8,0 12, i 2,1 horn right 10 17, i 8,6 12,2 2,5 2 width of uterine left 9 7,0 4, i 5,3 0,9 horn right 9 6,2 4, i 5,3 0,7 3 spread of uterine horn 10 33,0 18,9 25,7 5, i 4 width of body uterus 10 8,5 3,9 7,0 1,5 5 length of body of uterus 10 20,6 13,3 17,5 2,2 6 length of cervix 10 5,8 4,0 4,9 0,2 7 length of vagina and vulva 9 30,9 20,5 24,4 2,9 8 total length (vagina and 9 54,6 47,8 51,5 2,4 body of uterus) fig. 2 has been included. both foetal ovaries and testes have been used as the pattern of growth is basically the same. fig. 3 on the other hand, illustrates postnatal growth of the ovary. in fig. 4 ovaries have been arranged in a series from a 70-day foetus (1) to a fully mature adult with a large growing corpus luteum (0). the arrangement clearly shows how the foetal ovary initially increases in size (2) and then regresses (3) until after birth (4) and how, in the postnatal ovary, developing follicles deform the gland as they increase in size (5 to 10). growth of follicles in the adult ovary, however, will be discussed in a later section. gross changes in the size and histology of zebra foetal ovaries are discussed by smuts (197 4a). the shape and size of the postnatal ovary is largely determined by follicular development in the stroma (smuts 1974a). this development in turn varies considerably from animal to animal but in general the characteristic almost round shape observed at birth changes to flat and then to the bean-shape of the mature adult. the shape at birth (fig. 4), with the characteristic area of germinal tissue (later ovarian fossa), changes within the first few months of life so that at approximately one to two months of age the ovary assumes a flat shape, i.e. the free border is no longer the apex of a hump but is almost flat and is gradually assuming the characteristic features of the ovarian fossa. as follicles 95 -q en uj -0:: ~ 0 a uj 0:: ~ ll o 0 0 80 0 60 o os o o 0 • 00 o ........ 0---.... : 0 0 00 0 •••• •• ~ _ 00 •··• ••. 000 0 .' 00 " o. ~ ./ 0000.··".0 .... 0 0 __ / 0 0 0 0··...... 00 0 ~ 0 0 0 0 if> .. ,... 0 00 0 0 0 0 00 0 000 0 o 'l. 0 ... q,o 0 ~~..g, .. ~~..."..,..j''&--!!a.£.at;~ __ o -<do 0 "cf. #0 0 0 -cit".,.. ~odo il 8 o 0 00,,00 oocid08 '1> o 0 40 20 o o~--~~~--r-~~~r-~-.-,--~~~--r-~~--r-~ 30 70 110 o 190 230 270 310 3 0 phase of pregnancy (days) fig. 2. growth in mass with age of foetal gonads from burchell's zebra. (n = 43 pairs of foetal ovaries and 41 pairs of foetal testes). the curve was litted by eye. (/) w ~ 4 > o o w it: ~ u. o 90 80 $()cual maturity pueertal 70 immature .t.. ... i .. full bruding capac ity 60 so '0 30 20 10 .. i.y .j t .. ,... _ ..... . .."'···1·'" .... --... 1 .......... . ----1---............ ' .. -......... , 5 6 7 • i k) 11 12 13 " 1s" 1 19 years fig. 3. po stnatal growth in mass with age of burchell's zebra ovaries. the sample includes ovaries from 257 mares between birth and 19 years of age. the vertical lines indicate total variation within each age class and the two almost horizontal broken lines, the average maximum and average minimum mass for paired ovaries from mature mares. the broken lines thus illustrate average mass of the inactive ovary (lower line) and average mass of the ovary when at maximum activity (upper line). average data on puberty and sexual maturity are also illustrated . the curve was fitted by eye. 96 fig. 4. growth and development of the burchell's zebra ovary from the prenatal through the early postnatal to the adult stage. development proceeds in a clockwise direction from (1) to (10). with the exception of(1) and (2) all ovaries are arranged in pairs (each pair being from the same animal) those in the outer circle having been sliced down the centre from the free to the attached border and then opened out. with the exception of (5) and (6) the ovaries of the inner circle are arranged with their attached (mesovarial) borders in the same direction e.g. the edge of (2) closest to the "scale" is the attached border while the edge of (10) furthest from the "scale" is the attached border. in the case of (5) and (6) the free borders are dorsally situated. the small arrows present on ovaries (2) of the inner circle illustrate development of the ovarian fossa. (scale em). legend: (i) = 7o-day foetus; (2) = 255-day foetus; (3) = 375-day (full term) foetus; (4) = newly born foal; (5) = 2-month old foal; (6) = 6-month old foal; (7) = 9-month old foal; (8) = l!-year old mare; (9) = 2t-year old (sexually mature) mare; (10) = 3!-year old adult mare; (a) = attached border of ovary; (f) = free border of ovary. start enlarging below the germinal epithelium, the ovary is slowly inverted due to the relatively greater development of the medullary region in comparison to the cortical area. the latter area, now partially surrounded by, but not easily distinguishable from the medulla, forms the ovarian fossa. this fossa has normally been formed by the time the mare is three to six months old but becomes more pronounced in older mares with growing follicles. 97 in burchell's zebra there is undoubtedly a very clear morphological and histological difference between the free border with its contained ovarian fossa and the rest of the ovary. this is especially the case in young mares where the difference is visible even externally with the naked eye. i n all mares ovulation was found to occur from the ovarian fossa and never from any other region of the free surface of the ovary as was found by kupfer (928) in young horse and donkey mares. in the zebra there definitely appears to be a predestined area, the ovarian fossa, for the ovulation of follicles . . in adult zebra mares the involuted anoestrous ovary has an approximate mass of 109 (range = 5 to 18 g) while the active ovary has a mass of 30 g (range = 10 to 55 g) (fig. 3). the size and shape of the ovary is as variable as its mass and during the late stages of pregnancy it is generally long and narrow, and tapers from the tubal to the uterine extremity (fig. 8). during pro-oestrus, oestrus, metoestrus and during pregnancy the shape and size varies according to the size of the developing follicles and corpora lutea. fig. 5. difference in size between the tubular genitalia of a sexually mature (2!year old) burchell's zebra mare (upper) and an immature yearling. both tracts are viewed ventrally. legend: (1) = right ovary; (2) = fimbriated end offallopian tube; (3) = fallopian tube enclosed by mesosalpinx; (4) = ovarian ligament; (5) = right uterine horn; (6) = uterine horn slit open; (7) = body of uterus; (8) = cervix; (9) = external os of cervix; (10) = supra-urethral part of vagina (vagina proper); (1 j) = infra-urethral part of vagina (vulva); (12) = labium of vulva; (13) = clitoris in ventral commissure of vulva; (14) = bladder. 98 puberty and sexual maturity figure 5 illustrates the differences in size between the tubular genitalia of a sexually mature (2,5 year old) zebra mare and an immature yearling while development of the ovaries is illustrated in fig. 3. accepting that ovulation is an indication of puberty (donovan 1966), sections of ovaries from 40 knp mares between the ages of one and three years proved that the first ovulations occur at 1,5 to two years of age (2 mares out of a sample of8). in one case a mare estimated to be 21 months old, was found to be in the process of aborting a 473 g foetus. since the foetus was estimated to be four months old, the mare must have ovulated and conceived when approximately 17 months old. in the older age classes (2 to 3 years) (n = 24 mares) it was found that there was a gradual increase in the number of ovulating mares and by 3,25 to 3,5 years of age (n = 12 mares) all had ovulated as indicated by the presence of a developing or regressing corpus luteum. the conception rates for a sample of 58 young mares from a number of age classes are given in table 3. considering the two pregnant mares in the twoto 2,5 year old group and the four in the 2,5 to three-year old group and calculating when they conceived (method of huggett and widdas 1951 see smuts 1975) the following ages of first conception were obtained: 23, 24,27,29,31 and 32 months respectively. averaging these gives a first conception age of 28 months. table 3 age-specific conception rates for a sample of 58 young burchell's zebra mares collected in the central district of the kruger national park between 1969 and 1972 age class no. in no . conception (years) sample pregnant rate of age pregnancy class (%) 1 3 0 0 i-it 8 0 0 it-2 8 1 12,5 aborting mare 2-2t 9 2 22,2 normal pregnancies 2t-3 15 4 26,7 normal pregnancies 3-3t 15 11 73,3 normal pregnancies 99 the mating and foaling season the mating season is that period of the year when mammals mate and has often been referred to as the time when the females of a species come into oestrus. the foaling season on the other hand is that portion of the year during which the majority of young are born. the knp which forms part of the eastern transvaal lowveld, is a semi-arid region of the southern temperate zone. within the park annual rainfall varies considerably from place to place and even within an area, from one year to the next. over 80% of the rain, however, falls during the summer months (october to march) with a concomitant increase in green vegetation. the portion of the central district from which reproductive data were collected, i.e., the tshokwanesa tara area receives a mean annual precipitation of 529 mm (n = 32 consecutive years, 1941 to 1972) with a range of 232 to 942 millimetres. mean monthly precipition as registered at these two stations during the study period, are shown in fig. 6. mean monthly daylight hours for the study area are also illustrated. these were computed using data on times of sunrise and sunset at latitude 250 south and longitude 300 east of greenwich as supplied by the south african astronomical observatory. average daily temperatures for skukuza, which is situated on the southern edge of the study area, were calculated from maximum and minimum daily temperatures. by averaging the monthly data for 1969 to 1972, the temperature curves illustrated in fig. 6 were plotted . to obtain quantitative data on seasonal mating and foaling activity and also the annual spread of these activities, two basic methods were employed. firstly the foaling season for the central district was delimited by doing counts to determine the percentage of newly-born foals surviving to be counted each month. these results are summarised in tables 4, and 5 and fig. 6. all these data represent the average condition for the sa tara and eastern boundary sub-populations and for all practical purposes the condition for the entire central district. the second method used to delimit the mating and foaling season was based on a sample of 180 embryos and foetuses collected from shot mares . by measuring the mass of these embryos and foetuses and employing the foetal growth formula of huggett and widdas (1951) (smuts 1975) it was possible to calculate both the month of conception and the projected month of birth of each embryo or foetus (table 5). since the collection of this material was undertaken on a monthly basis and the sample was distributed over a period from may 1969 to november 1972, between three and six foetuses could be collected per month. after calculating both the month and year of birth and conception for each embryo or foetus they were then distributed into the 12-monthly intervals (july to june) and the relative percentage conceived or to be born during each month of each 12-month interval was calculated (table 5). in fig. 6 relative monthly conception percentages as given in table 5 have been illustrated. these can be compared directly with the illustrated 100 i t,~ z 0 ~ u e 160 z 0 0 .§ u j 12ij ~ ~ '" >-z z ~ 80 0 :e 8 >;:: ",z ~ ~ 8 4 o:e~ ~ " z 0 b 3 ",,,>~1.i.i!..l. 00 ~~~ w:>w zou u~ o \ / ~ jl \ / . , __ i ~. .---../ '-. jasondj rmamj j a so ndj fmamjj a s ono 1969 1970 1971 , , 1972 noj f mamj 30 1973 25 average daily 20 temperature ee) ~~ daylight hours _~ 333mm ~4ol _ 414mm _-j71mm 6 --· 10 ~q ·----6j,6~~ -91.3 % [ ' ~~ at latitude 25 " 5, " 9 longitude 30 e u :f! <ii ~ 20 ii' :e o ~ j a 1968 j , 1968 1969 month and year fig. 6. the foaling season (upper graph) and mating season (lower graph) for zebra from the central district of the kruger national park. rainfall histograms as well as curves for average monthly daylight hours and average daily temperature are also included in the figure . total summer precipitation (october to march) and conception rates for mares shot during the winter months of each year are illustrated above and between the conception curves. i@ ~ f!! ~ ~ ~ m ., ~ f!! ~ !fa o .j f 1972 1913 table 4 foaling activity lor burchell ',! ubra counted at monthly intervals in the central district oj the kmgrr national park between july 1969 and february 1973, counts have been divided intojour 12-monlhly interval! so that comparisons can be made between the joaling peaks oj successive 12-monthly intervals perce ntage relative percentage relative of new ivpercentage of newlypercentage month born lo~ls of newlymonth born foals of newlyand in each born foals and in each born foals year monthly counted year monthly counted sample during sample during each month each month .lui\' 1969 0,34 2,59 july 1970 0, 15 1,66 aug-, 0,35 2,67 aug, 0,24 2,66 sept. 0,39 2,97 sept. 0,05 0 ,55 oct. 0 ,2 9 2,21 oct. 0,31 3,43 no\', 0 , 19 1,45 nov, 1,02 11,30 dec, 1,2 9 9,83 dec. 1,80 19,93 jan, 1970 4,18 31,86 jan, 1971 1,83 20,27 feh, 3,24 24,70 feb . 1,66 18,38 mar. 1,65 12,58 mar. 1,15 12,74 apr. 0,70 5,34 apr. 0,46 5,09 mav 0,39 2,97 may 0,27 2,99 june 0 , 11 0,84 june 0,09 1,00 julv 1971 0,09 0,6 5 july 1972'" 0,19 1,98 aug. 0,00 0,00 aug. 0,20 2,08 sept. 0 ,00 0,00 sept. ':: 0,15 1,56 oct. 0,05 0,36 oct. 0,38 3,95 nov. 0,97 7,03 nov. 0,26 2,71 dec. 3,67 26,61 dec. 3,62 37,67 jan. 1972 3,72 26,98 jan. 1973 0,98 10,20 feb, 1,62 11,68 feb. 1,00 10,41 mar. 1,3 1 9,50 mar. ::: 1,37 14,26 apr. 1,05 7,61 apr. ':' 0,74 7,70 may 0,82 5,95 may 0,49 5,10 june 0,50 3,62 june .;, 0,23 2,39 ,', during these seven months no co unts were undertaken and their results were estimated by averag-ingthe data ii-om th e corresponding months of the preceding three years. climatic variables. average foaling and mating activity are given in table 6 which also indicates that the two methods of determining the foaling season provide similar results. further evidence for strong seasonal reproductive activity is supplied by analysis of ovaries from 48 non-pregnant adult mares. of this group 17 were definitely in anoestrus, the rest having either large corpora lutea and/or many developing follicles. of the 17, one was shot during december and the other 16 between april and july (5 in june). this means that over 94% of all anoestrous mares were collected during the winter (non-breeding season) and indicates that should a mare fail to 102 o e"o z 0 '" -( i.-j '" 0 z :l -( 0 '" month and year of birth july 1969 aug. sept. oct. nov. dec. jan. 1970 feb. mar. april may june nt.' 0 0 0 i 3 to 8 8 6 2 0 4 table 5 sea.lonal distribution of births (i) and conceptions (ii) for burchell's z.ebra collected in the central di.ltrict of the kruger national park between may 1969 and november 1972. (n = 180 embryos and foetuses) relative relative relative percentage month percentage month percentage month of foals to and n of foals to and n of foals to and be born year of be born year of be born year of during each birth during each birth during each birth month month month 0 july 1970 1 3,85 july 1971 0 0 july 1972 0 aug. 0 0 aug. 2 2,94 aug. 0 sept. 2 7,69 sept. 0 0 sept. 2,38 oct. 8 30,77 oct. 0 0 oct. 7,14 nov. 3 11,54 nov. ii 16,18 nov. 23,81 dec. 3 11,54 dec. 12 17,65 dec. 19,05 jan . 1971 7 26,92 jan. 1972 13 19,12 jan. 1973 19,05 feb. i 3,85 feb. 13 19,12 feb. 14,29 mar. 0 0 mar. ii 16,18 mar. 4,76 april 0 0 april 3 4,41 april 0 may i 3,85 may 1 1,47 may 9,52 june 0 0 june 2 2,94 june ------------relative percentage n of foals to be born during each month i 2,27 1 2,27 i 2,27 3 6,82 7 15,91 17 38,64 7 15,91 5 11,36 i 2,27 0 0 0 0 i 2,27 o oj:>. table 5. (continued) month and relative month and relative month and year of n" monthly year of n monthly year of concepconception concepconception conception percentage tion percentage tion july 1968 0 0 july 1969 1 3,85 july 1970 z aug. 0 0 aug. 0 0 aug. 0 sept. 0 0 sept. 4 15,38 sept. '" « oct. 2 4,76 oct. 7 26,92 oct. "'-l '" nov. 6 14,29 nov. 2 7 ,69 nov. c) dec. 8 19,05 dec. 5 19,23 dec. z jan. 1969 9 21,43 jan. 1970 6 23,08 jan. 1971 f: « feb. 6 14,29 feb. 0 0 feb. ::e mar. 4 9,52 mar. 0 0 mar. april 3 7,14 april 0 0 april may 2 4,76 may i 3,85 may june 2 4,76 june 0 0 june ':' n = the number of embryos and foetuses used to calculate the relative percentages. relative month and relative n monthly year of n monthly conception concepconception percentage tion percentage i 1,43 july 1971 0 0 i 1,43 aug. 1 2,38 0 0 sept. 4 9,52 4 5,71 oct. 3 7,14 14 20,00 nov. 6 14,29 9 12,86 dec. 17 40,48 16 22,86 jan. 1972 6 14,29 11 15,71 feb. 3 7,14 9 12,86 mar. i 2,38 2 2,86 april 0 0 0 0 may 0 0 3 4,29 june 1 2,38 -o '" table 6 average foaling and mating activity for burchell's zebra in the central district of the kruger national park for the years 1969 to 1973 . this table represents the averages for the data given in tables 4 and 5 foaling season mating season average ( 1) field count method (2) foetal mass method between method (j) and (2) average average relative relative average average percentage percentage , relative relative month of newly per cent no offoals percent percentage per cent month n" monthly per cent born foals of total to be born of total of foals of total conception of total counted during each produced percentage during each month of per month month of the year the year july 1,47 2 i,ll 1,29 july 2 i,ll aug. 1,74 3 1,67 1, 71 aug. 2 i,ll sept. 1,30 3 1,67 1,49 sept. 8 4,44 oct. 2,27 12 6,67 4,47 oct. 16 8,89 nov. 5,37 } 46,4 24 13,33 9,35 nov. 28 15,56 dec. 22,83 82,6 42 23,33 } 42,8 87,8 23,08 }44,6 85,2 dec. 39 21,67 } 42,2 85,6 jan. 23,56 35 19,44 21,50 jan. 37 20,56 feb. 16,54 27 15,00 15 , 77 feb. 20 11,11 march 12,06 18 10,00 11,03 march 14 7,78 april 6,49 5 2,78 4 ,64 april 5 2,78 may 4,33 2 i,ll 2,72 may 3 1,67 june 2,05 7 3,89 2,97 june 6 3,33 on = the number of embryos and foetuses used to calculate the relative percentages conceive during the summer, she is likely to remain in anoestrus during the ensuing winter. analysis of these ovaries is discussed in a later section. conception rate and the factors affecting it between 1969 .05.16 and 1972.11.13 the reproductive tracts of 236 adult (3 to 20-year old) mares were examined to ascertain their reproductive status . of this sample, 184 were confirmed pregnant (78%) as indicated by the presence of a conceptus in the uterus. in addition to these, 35 adult mares were shot by various staff members during 1966. of this sample 31 were visibly pregnant (88,6%). the conception rate for the entire sample of 271 adult mares of which 215 were visibly pregnant, is 79,3% and covers a period of five years. breakdown of conception data into the various adult age classes is given in table 7. table 7 age-specific conception rates for 279 burchell's zebra mares collected in the central district ojthe kruger national park between 1969 and 1972 age class (years) 1-2 2-3 3-4 4-5 5-6 6-7 7-8 8-9 9-10 10-11 11-12 12-13 13-14 14-15 15-16 16-17 17-18 n 19 24 15 21 30 22 26 14 22 18 27 14 7 10 8 o 2 no . visibly pregnant 1 6 11 19 20 16 18 12 15 15 25 14 5 9 4 2 conception rate (%) 5,3* 25,0 73,3 90,5 66,7 72,7 69,2 85,7 68,2 83,3 92,6 100,0 71,4 90,0 50,0 100,0 ", the single pregnant mare in this group was in the process of aborting when shot 106 by dividing the collection of adult mares from the central district into two groups representing those from the sedentary satara subpopulation and those from the migratory eastern boundary sub-population, the following conception rates were obtained: satara sub-population: 80,2% (n = iii mares) eastern boundary sub-population: 78,8% (n = 160 mares). these differences are not significant at the 5% level (chi-square = 0,01). since conception rate is relatively constant throughout the central district it would be interesting to determine whether rates differ significantly from one year to the next. however, to compare the conception rate of one breeding season with that of another it is necessary, in the case of the zebra, to separate the sample of adult mares into those collected during the breeding (mating) season and those collected during the non-breeding season. this is essential because only those collected during the non-breeding seasons of individual years can be compared. the reason is that mares shot during the mating season and carrying large foetuses are mares that have reacted to conditions from the previous season, while those carrying small foetusses or embryos have fallen pregnant during the current season and therefore reflect the conditions of this season. conception data collected during both these times of the year would thus represent the average condition for two successive breeding seasons and it would subsequently be difficult to assess the effects of a particular season on conception rate. the same would apply to non-pregnant mares, i.e. one would not know which season was responsible for their condition. bearing the above-mentioned in mind and looking at the graph of relative monthly conception percentages illustrated in fig. 6, it is possible to divide the period of collection into non-mating seasons as given in table 8. these seasons represent intervals of time which start when monthly conception percentage drops drastically and extend up to the time when monthly conception percentage shows its first marked intable 8 comp aris on of non-breeding season conception rates for burchell's zebra for each oj/our colljecutive yean. only adult mares ( 3 to 20 years old) collected in the central district oj the kruger park were us ed approximate no. of no . non-breeding seaso n length of adult confirmed conception (non-mating season ) season mares pregnant rate (%) (months ) shot april ' 69-s ept. ' 69 5 20 14 70,0 feb. '70-0 ct. '70 8 44 28 63 ,6 april ' 7 i-oct. '71 6 46 42 91,3 march '72-0ct. ' 72 7 41 31 75,6 107 crease. mares collected during these periods would, in the majority of cases, have reacted to the conditions of the preceding summer season and their conception rate would thus be representative of that season. conception rates for the four non-breeding seasons experienced during the study period are also given in table 8 and in fig. 6. ovarian activity during the oestrous cycle for convenience the ovarian cycle of the adult mare has been divided into two sections, namely ovarian activity during the oestrous cycle and ovarian activity during pregnancy. these cycles, however, follow each other should fertilization and implantation be effected. fig. 7. four pairs of burchell's zebra ovaries cut from their free to attached b.orders and opened out to illustrate their internal appearance at four phases of the oestrous cycle. (i) pro-oestrus one follicle in the lower ovary has increased in size, while the upper ovary contains two corpora lutea from previous oestrous cycles; (2) metoestrus note the large corpus haemorrhagicum and the absence of large and medium-sized follicles; (3) dioestrus the corpus luteum has started to regress and follicular development is apparent. this is probably a late dioestrus specimen and most of the ovarian stroma is taken up with follicles of various sizes; (4) anoestrus the typical anoestrus ovary assumes a cashew nut shape and lacks medium-sized and large follicles. dark brown corpora lutea in the final stages of regression (see arrows) are frequently present while stroma is very obvious. notice that in all specimens ovulation has taken place via the ovarian fossa. 108 the phases of the oestrous cycle in the zebra have been outlined in a previous section. these phases are, however, primarily related to the activity of the ovaries and in this context they may be summarised as follows: pro-oestrus there is a marked increase in follicular activity with one follicle generally considerably larger than the others. regressing corpora lutea from preceding cycles may be present (fig. 7). oestrus similar to pro-oestrus except that there is no doubt as to which follicle is going to rupture and the mare now accepts the stallion. unfortunately none of the cropped mares could be placed in this category. table 9 characteristics of the ovaries of 42 adult, non-pregnant burchell's zebra mares shot in the central district ojthe kruger national park feature of ovary pro-oestrus metoestrus dioestrus anoestrus n=3 n=6 n= 18 n= 15 mean 25,1 16,0 18,0 9,9 mass (g) max. 36,8 30,0 39,8 18 , i min . 15,8 6,0 5,4 4,8 mean 4,4 4,3 4,0 3,5 length (e m ) max . 4,7 5,5 6,0 4,6 min . 3.8 3,5 2,7 2,8 mean 3,0 2,2 2,6 2,1 width (em ) max. 4,2 2,8 3,2 2,8 min. 2,2 1,6 1,2 1,5 diameter (mm) range in number 1-10 10-57 1~4 6-41 11-41 of follicles per pair 11-20 2-9 0-3 1-17 0-2 of 0 varies 21-30 0-1 0 0-2 0 31-40 0-1 0 0 0 greatest diameler of follicle (mm) 31 20 25 13 diameter (m m) range in number 1-10 1-6 0-1 0-4 0-6 of corpora lutea 11-20 0-2 0-1 0-1 0 per pair of ovaries 21-30 0 0-1 0-1 0 31-40 0 0-1 0 0 grealest diameter of corpus luteum 20 32 27 10 (mm) 109 metoestrus the ripe graafian follicle has ruptured and a characteristic corpus haemorrhagicum formed. at this stage the cavity of the follicle is filled with a lymph-like fluid which usually has a considerable amount of extravasated blood in it (fig. 7). as the cells lining the ruptured follicle undergo rapid hypertrophy and become luteinized, an orange coloured corpus luteum is formed. follicular activity is largely in abeyance. dioestrus should the mare fail to conceive, she goes into dioestrus or anoestrus. during dioestrus graafian follicles start developing slowly (fig. 7) while the corpus luteum starts to regress, becoming smaller and lighter (ye llow ) in colour. this phase is generally followed by the prooestrus phase. anoestrus mares which have failed to conceive towards the end of the mating season may cease to cycle and go into anoestrus. the ovaries of anoestrous mares (fig. 7) are relatively inactive and contain a few small follicles and usually a number of tiny dark brown corpora lutea in advanced stages of regression (corpora albicantia). changes occurring in the ovaries of 42 adult non-pregnant mares are given in table 9 and should be compared with fig. 7 which illustrates examples of four of the phases of the oestrous cycle. in the sample of 42 adult non-pregnant mares, there was only one specimen in which a multiple (double) ovulation may have taken place. this was a pro-oestrous mare with a regressing corpus luteum in each ovary. these corpora lutea measured 19 x 19 and 20 x 19 millimetres. the sample of 42 was a bit small to satisfactorily gauge comparative ovarian activity but in 64% of the specimens the left ovary had a greater mass than the right, while in 60% the corpus luteum was situated in the left ovary. the sample sizes precluded statistical evaluation. in none of the ovaries sectioned was there any evidence that ovulation had taken place from any region of the ovary other than the ovarian fossa. i n a number of cases there was protrusion of the corpus luteum or corpus haemorrhagicum (fig. 7) from the ovarian fossa. ovarian activity during pregnancy ovarian activity during the 37 5-day long gestation period of the zebra mare has been divided into 19 arbitrary phases (table 10). unfortunately there were only four specimens available for mares in the early stages of pregnancy (phases i to 3), this largely being due to the fact that mares are easily classified non-pregnant during the first week or two after conceiving and that hunters were reluctant to shoot mares with young foals at foot. for the other 16 phases, each of which (except the last one) covers a period of 20 days, larger samples were available. altogether 172 pairs of ovaries from pregnant mares were sectioned, the more important data being summarised in table 10. representative pairs of 110 data collect ed _from "lacroscopical sections made on j 7:2 pairs €if ovaries from burchell ~s z.ebra a t various _' tages of pregnancy. data have been grouped into 19 arbitrary pregnancy phases and means, maxima and minima for various p arameters of each phase calculajed. all mares were shot in the central district of the kruger national park between 1969 and 1972 p has e s 0 f" ' pre g nan c y (, to 19) 10 11 12 1] ,. '" '6 '7 18 '9 ronception age in oays ~e size ' 5 1 20 1 23 161_8° 181_10° 1'0'_'2°1'2 1_'401'4'_'601'6'_'8°1 181 2uo i20 1-220 1221-240 1241-2ffi i 26 1-280 \ 28 1_joo i 301-320 1 321_j40 1 341_jgj \,'6 1 _37~j n . 1 " .1 " .. 2 ". 12 n-e n.1 1 noo8 n _l ) n a4 n .1 3 n .1 j ".10 ",.1 1 ., .4 .,.1:? "",5 " .11 n . 16 n .1 s taa.ss of" pa ired ovaries (9) length (c.) width (c») t:l 01arrleter of follicle meij" "", mi., . .. en l.\3 x, min , .. ,,.n .... , ui n, mean and r3ng e in number of fall ides 87,9 1 16,6 148,0 140 ,5 139 ,a 1 42,5 jo ,7 _ 52,a 89 ,1 71,7 58,7 45 ,9 _ 43 , 5 19,6 21 , 5 21,1 ii , ::) 5,21 2,91·,5 i ",' i·" 6, i j, e '),7 5 , 6 5,2 5 ,' 2,7 j ," 2 ,7 3,1 3 , 3 1 2, 0 1 3 , 0 1 2 , 7 1 2 , 6 3,4 2,1 3 , 1 3 , 9 j ,~ .1,3 l,a 2 ,8 1, 2 1,7 4 , 3 5,' 3,' 2,6 j ,6 1 ,s' 3,9 5 ,0 3, 0 2 , 4 j,5 ' , 8 34 ,2 24,0 2 1, 8 ':)6, 9 43,3 38,0 19 , 8 19,0 1j,4 . , 3 5 , 8 3,2 2,. 3,6 1,8 c.'.,8 3 , 3 2 , 0 2 , 8 ',6 3,9 :. ,2 3 , 2 2 , ' 3 , j , , 5 19,1 tl~,:' 7," 1 , 8 . , 8 '? , 6 1 , 9 2 , 6 1,2 i , _ 5 5 1 8 1 6 \ 11 1 17 , 1 1a 13 4§ 123 ~12 j-:15 0-38 ~11 9 17 :3 .3: q2:; mean i 17 range l'; g; ~ ffi .. cl s g 6 10 11 is 16 20 21 25 26 3d me a n i 12 fie",e mea' i 1? ra'''ge ""an range ... an ra nge " an range greatest oia!.je~ 15 i " f"olucle :" nwrd ~ q.~ ~i §2; ~l'; §~ di i'ltneter of" co rpus luteul"l ;a ,;;;e in numbe r of ccrp8 r!l l l;tea 1 _ § iiwn ira..-.ge 6 10 ""'" i range " 15 "'" i range 16 za rtvn i q,a ,'ge 2' 25 mm ' range 26 jo '!i<n i hango:;: 0-' (]..., 16 1 13 1 12 1 '" 1 1? 13 1 7 '~21 3-23 2-27 5-3& 1?8 3-27 922 '~'e i 1:9 4 3 7_1~1 0-20 1 3-16 i (l.,1j 0-10 0-9 1 (l..? i :j-1 ~2 i 02 :i i ;: 1 1 0, i c-, i j...4 o 0' i i 0-2 2c i 211 17 0-1 0-7 i :j....5 0-, j..., i 1)....2 i 0-' 1>-1 i 01 i 0-1 0-1 i 0-1 o 0-, 28 0-6 >-2 0-, 0-2 0-, 17 20 0-. 2_4 >-, q..., i 0-1 1>-' i 0, q...2 i (l.,1 0-, 11 " 0-2 ~5 0-4 >-, 0-' oj 0-' 0-, 0, 0-, ~j 0, [l; c1 31 .33 mm i rarv;lc i , i _ ' 0 -1 28,3 19,3 jo,3 3 :'\4 13 , 6 13,1 " , 5, , 2 . ':1 ',9 2 , : '1 , 11 a,. 6 ,' 3,0 ',9 3 , 7 ' , s 19 12 539 223 )....11 i 019 0-2 " ·~5 0-5 0-' 2-2 0-, 0-2 3-2 0-3 cl 18,9 i ~o.3 1 17 , 9 lb ,a 23 , 9 25 ,3 22 , 6 26,9 '5 , 3 11,6 12,0 1~,2 -l,u .,8 3 , 3 1 , 8 2,' 1,5 a,':> 6,0 3,0 2,0 2,a 1,3 a,7 6,' 3, , 1,8 2 , , 1,3 a,a 6,a 3,2 1, 9 2 , 8 ',' 19,2 28,7 9,0 a,8 7,a 3,6 1, 9 2 , 3 ', 2 1~25 ~;1 ~~o ~~ i 4~' 5 012 ' 3-18' 2-14 "0 ' 1 ~<! 01 i i 8-1 11 10 11 0-2 3-s 24 ' 7 1_ 3 0-1 0, 0-, 0-1 0, 0-' 0-' 01 c~~~;i~ largest (xjnpus 1 2"7><22 121k 19 i z?""9 1j4x29 t 29x2'3 i 29><25 i 27,..21 i 25x22 i 22,,1 7 i 25><17 i 2: ,, 1j i 18 x' s i 2 1,, 19 i 11><7 '''',9 1(::)1(2 i :;,,6 ' 2x'7 18,9 n,s 11,5 il , ,1 5 ,6 3,0 ',9 j, • ',a '" '-'0 o'e o c.. , " ,; 1-3 0-, 9.,<3 ovaries h-om mares at various stages of pregnancy appear in fig. 8. the ovaries of zebra mares present a number of interesting features. firstly follicular development is stimulated during certain phases of pregnancy fig. 8. eleven pairs of ovaries from burchell's zebra mares at various stages of pregnancy. the ovaries have been cut from their free to attached borders and opened out to illustrate the internal appearance. all ovarian pairs are arranged vertically with the tubal extremity uppermost. on the photograph the coagulated liquor folliculi of most follicles appears as a shiny surface, while the corpora lutea are duller, one half of each being pin-pointed by a small black arrow. legend: (1) = 15 days pregnant. notice many small follicles and the large developing corpus luteum with blood stained liquor folliculi still in the centre. (2) = 20 days pregnant. at this stage the corpus luteum of pregnancy has already reached its maximum size and there was a reduction in follicular activity; (3) = 74 days. at this stage there was an increase in follicular activity while the corpus luteum of pregnancy was gradually regressing; (4) = 102 days. in this specimen as in (5), follicular activity is still obvious. it also has two corpora lutea both being approximately the same size; (5) = 113 days; (6) = 135 days. this specimen shows reduced follicular activity in both ovaries, while a luteal scar (corpus albicans) is clearly visible as a dark narrow streak in the lower ovary. this is a structure from a previous oestrous cycle; (7) = 145 days. here ovarian activity is still obvious in both ovaries; (8) = 165 days. in this specimen follicular activity is reduced, as is the size of the corpus luteum of pregnancy; (9) = 239 days . strongly reduced follicular activity and a small corpus luteum and one corpus albicans; (10) = 343 days. little follicular activity and no visible corpora lutea; (1 i) = 375 days of full-term. two small follicles are visible in the lower ovary. both ovaries have become long and thin a typical shape for the late stages of pregnancy. 112 and secondly, this activity may be followed by ovulation resulting in the fonnation of so called secondary corpora lutea. using average data in table 10 together with raw data on individual pairs of ovaries, figs. 9 and 10 have been prepared. these figures illustrate ovarian activity during pregnancy and use the mass and width of the gland respectively, as indices of activity. " _80 2 ii) 070 ~ z 0 0" ...j :! 50 w lr 40 0 w ~ 30 t il 20 0 ii) •• ii) ~ ~ 0 70 •• foetal ovaries • • foetal testes . . : ", .-." . ........••..•. • • ,ii' • . ". • • .......... • • .. ....•..• • • • .,. • • .. • • • • • .. • ...• • • " ' .. • · . • • • .0 no 1)(1 150 170 ., 2'10 230 250 27'0 lto "0 s30 350 175 conception age (days) fig. 9. scatter diagram illustrating changes in the mass of paired burchell's zebra ovaries during pregnancy. average results for the 19 arbitrary phases of pregnancy given in table 10 are also illustrated (closed circles). the curve was fitted by eye. e 40 e +u 0 >30 0 q: # ~ ..•......•.... .--.....•• tt 0 20 § ll 0 j: 10 .... e -~ 0 30 70 110 150 190 230 270 310 3 phase of pregnancy (days) fig. 10. width ot burchell's zebra ovaries (smooth surface to smooth surface) as an index of follicular activity. the open circles give the average width for each of the 19 arbitrary phases of pregnancy given in table 10 and the vertical lines the total variation of each. the curve was fitted by eye. 113 in addition to the above-mentioned indirect methods of assessing ovarian activity, activity can be gauged directly from the size of developing or regressing follicles. follicles with the greatest diameter are found between a conception age of 23 and 160 days (table 10). in one instance of a mare, estimated 102 days pregnant, there was a graafian follicle with a diameter of 28 millimetres. this follicle may be compared to prooestrous follicles (table 9) . in addition the only two instances of recent ovulations (corpora haemorrhagica) during pregnancy were found in mares 72 and 98 days pregnant. since good samples were available for the rest of pregnancy, it is obvious that follicular activity reaches a maximum between about 70 and 110 days and that secondary corpora lutea resulting from ovulation may be found during these stages. after this stage of increased follicular development there is a gradual regression of follicles until the relatively inactive 190 to 375 day stage is achieved . in contrast to follicular activity, the corpus luteum of pregnancy shows less spectacular changes. in mares with single ovulations and no secondary corpora lutea, the corpus luteum of pregnancy regresses fairly slowly and remains clearly visible in most full and near-term mares. here it appears as a dark yellow-brown corpus albicans with dimensions of up to 10 x 2 millimetres. in mares with secondary corpora lutea, it is impossible to gauge what happens to the corpus luteum of pregnancy but it is likely that it regresses more rapidly than in mares without secondary corpora lutea. in mares with more than one corpus luteum, the largest may still measure 21 x 19 mm at 249 days (table 10). in fig. 8 the upper ovary of the number "9" pair shows a fairly large regressing corpus luteum. this animal was 239 days pregnant. figure 8 also illustrates the gradual regression in size of the corpus luteum of pregnancy. no corpora lutea are visible in the last two pairs of ovaries (1o and i i) although sections further down produced the characteristic scar tissue of corpora albicantia. by counting and measuring corpora lutea in the 172 pairs of ovaries sectioned, it was found that 26 05,1%) animals had secondary corpora lutea, i.e., two or more corpora lutea which definitely originated during the current pregnancy. as mentioned, two of these were fresh corpora haemorrhagica, the rest being corpora lutea in various stages of regression. all secondary corpora lutea appeared to have resulted from ovulations in contrast to the possibility that luteinization of an unrupted follicle may have occurred. in this respect all corpora lutea had the typical pear shape with the narrow core region abutting on the ovarian fossa (fig. 8, ovary pairs numbers 3 to 8). of the 26 individuals showing secondary corpora lutea, 23 had one secondary corpus luteum, two had two secondary corpora lutea and one (a mare 249 days pregnant) may have had three or more. in the latter mare a total of eight corpora lutea with diameters of between six and 21 mm were identifiedall were produced from follicles which had ovulated. it is, however, unlikely that all the corpora lutea had been produced during the current lj4 pregnancy, i.e. two or three could possibly have been from oestrous cycles prior to the pregnancy. the distribution of the 23 single secondary corpora lutea and the corpora lutea of pregnancy were as follows: one in the left ovary and one in the right ovary = 13 both in the left ovary = 7 both in the right ovary = 3 in the three pairs of ovaries with two or more secondary corpora lutea, the left ovary again showed the greater activity. with regard to diflerential activity of the ovaries, a number of deductions can be made. as in the case of the stallion (smuts 1976), the left gland of the mare appears to be more active than the right. excluding the tiny corpora albicantia (1 to 5 mm diameter) which are easily missed when sectioning ovaries macroscopically, a total of 109 corpora lutea were counted in all left ovaries and 73 in all right ovaries. this disparity (60% left and 40% right) indicates a diflerential ovulation rate and is a highly significant deviation from the expectation that ovaries are equally active (chi-square = 7,12; p<o,oi). where the corpora lutea of pregnancy could be positively identified, 32 were in the left and 16 in the right ovary. similarly the mass of the left and right ovaries differed significantly from one another. out of 340 ovaries (170 pairs) the left ovary had a greater mass than the right in 120 instances (70,6% left> right) (chi-square = 28,82; p < 0,005). since the left ovary is more active than the right, it would be interesting to see how this compares with implantation. in a sample of 63 pregnant mares in which the side of implantation was ascertained without any doubt, 35 (55,6%) were left and 28 (44,4%) right. however, in 48 of these 63, the corpus luteum of pregnancy could also be identified and it was found that trans-uterine migration had occurred in 20 instances (41,7%). in the other 28 cases ovulation and implantation occurred on the same side. of the 20 migrations 12 were from the left horn of the uterus to the right. ovarian abnormalities during the present study 262 pairs of ovaries from mares between one and 18 years of age were examined and macroscopically sectioned. of this sample, 219 pairs were from adult mares (over 3 years of age). gross examination of these ovaries indicated that five pairs displayed bior unilateral abnormalities, all five being from mares in the adult group. histological aspects of these ovaries are described in detail by rmes and smuts (1975). discussion short term studies on aspects of reproduction in the zebra mare (e.b. boehmi) have been undertaken by king (1965) in east africa, while some comparative aspects of reproduction in the equidae have been discussed 115 by king (1966). other studies on burchell's zebra have also been undertaken on the subspecies boehmi and here the studies of klingel merit mention (see klingel 1965, 1967 and 1969 a and b and klingel and klingel 1965) as do those oftrumler (1958) and wackernagel (1965). the oestrous cycle according to wackernagel (1965), oestrus in burchell's zebra lasts from two to nine days (six days in the present study). the first postpartum oestrus which occurred at eight to nine days is in agreement with the nine to 10 days of wacke rna gel (1965) and the seven days given by klingel (i969a). wackernagel (op cit.) also found the length of the dioestrous period to be between 17 and 24 days (range = 11 to 28 days) . these observations are similar to the domestic horse where oestrus varies from five to nine days (asdell 1946), depending on the breed. in south africa, foal heats of the horse have been recorded commencing at 8,8 days (du plessis 1964) and eight to 11 days (belonje 1949). the average length of the oestrous cycle has been recorded at 23,7 days (van niekerk 1967) (south africa) and 22,8 days (sato and hoshi 1934) (korean mares in japan) while the length of dioestrus was from 11 to 16 days (day 1940) and 17 to 19 days (belonje 1949). gestation period pregnancy and birth as is the case with the various breeds of domestic horse, duration of gestation in zebras is extremely variable. kenneth and ritchie (1953), for example, quote periods ranging between 336 and 375 days, for equus burchelli. more recently wackernagel (1965) was able to determine the gestation period for 28 grant's zebra (equus burchelli boehmi) conceived and born in the basle zoo, switzerland. here the average for both male and female foals was 371,2 days. males were carried longer, on an average 373,2 days (range = 361-390 days) while females were carried on an average for 369,5 days (range = 361 to 378 days). comparative results are given by klingel (1969a), i.e. 378 and 385 days while in the present study it was 398 days. aspects of the birth process as it occurs in burchell's zebra have been described by klingel and klingel (i965) and wackernagel (1965). wackernagel (op cit.) also measured the mass of two afterbirths (2,7 and 2,9 kg) which were similar to those in the present study (3,2 and 3,3 kg). m otherjoal relationships the close bond between a mare and her young foal must be of considerable survival value during the critical period of imprinting. similarly the protective role of the mare must be of value since klingel and klingel (1965) have seen it operate in the presence of predators such as spotted hyenas (crocuta crocuta). here mares with young foals would hide behind other members of the family group. 116 lactation and weaning although most foals are weaned at 11 months of age they may be separated from their mothers at an earlier age without ill effects. in one instance, for example, a foal had already become a member of a stallion group at nine months of age. this animal had probably lost its mother a short while before . captive foals forcibly separated from their mothers have been successfully weaned at six to eight months of age, while king (1965) gives the weaning age for e.b. boehmi as seven months. development oj the uterus and vagina as is the case with the stallion (smuts 1976) the general morphology and location of the female reproductive tract and its associated structures are very similar to those of the domestic horse (eckstein and zuckerman 1962) and have been described by king (1965). by comparing the dimensions of the tubular genitalia (table 1) with the minimum values given for adult mares in table 2, it can be seen that in most instances the adult dimensions have been attained by 2,5 years of age. development oj the ovaries in fig. 2 it can be seen that the mass of the foetal gonad increases very slowly up to a conception age of approximately 125 days after which there is a rapid increase in mass until about 230 days. after this age they gradually decrease in size until just after birth when a minimum mass of approximately four grams is attained. the fluctuations are strikingly similar to those reported for foetal horse gonads by cole, hart, lyons and catchpole (1933) and amoroso and rowlands (1951). the large variation in ovary mass for the different age classes of adults shown in fig. 3 is mainly due to the reproductive status of each mare. average data, however, indicates that ovarian growth in young mares is rapid and that the average mass of the active adult ovary is attained at about 20 months of age. this contrasts with the stallion where the minimum mass of adult testes is attained for the first time by 3,5 year old stallions (smuts 1976). fig. 4 ((6) to (io)) clearly shows the extent to which growth of the post natal ovary is dependent on follicular and luteal development. the observation that ovulation in the zebra mare occurs via the ovarian fossa has been verified in studies on the domestic horse. here witherspoon and talbot (1970) found that the fimbria were displaced in a position where they covered the areas of the ovarian fossa alone, but not the entire ovarian surface. their research involved the use of fiberoptics, cinematography, india ink translocation and histologic evaluation . puberty and sexual maturity in the present study no direct field observations were made with respect to mating activity in young mares. wackernagel (1965), however, 117 found that young captive zebra mares displayed the first signs of oestrus at 15,5 months while king (1965) gives an age of 22 months . klingel (1969a) states in his study that the youngest mares that were recorded being served were estimated to be 18 months old. with rapid ovarian development commencing at about 16 months of age (fig. 3) and since one mare had conceived at 17 months of age and later aborted her foetus, it would seem that the pubertal interval extends from about 16 to 22 months with the first successful conceptions occurring at 23 months. this is supported by ovulations, the first of which were noted in mares 1,5 to two years old . although mares may conceive successfully at 23 months of age the average age of first conception (28 months ) indicates that most mares only become fully mature during their third year of life. these conclusions are supported by data given by king (1965 ) where the average age of first conception was also 28 months (range = 22 to 33 months) . wackernagel on the other hand obtained ages of 30 and 31 months for captive mares with another of 20 months for a mare which aborted five months later. klingel (1969a) found that the first successful conceptions occurred at about 30 months of age . in addition to chronological age, body mass is known to be related to puberty uoubert 1963) and in the zebra mare the body mass range displayed by adult mares is attained for the first time by two-year olds (smuts 1975) . finally, full reproductive capacity is only attained at about three years of age (fig. 3) as exemplified by an elevated (73,3%, table 3) conception rate, comparable to that for adult mares (79,3%). in contrast to the zebra stallion with its long pubertal interval and late age at sexual maturity when compared to the domestic horse (smuts 1976), puberty and sexual maturity in the zebra mare occur at roughly the same age as they do in horse mares. according to cupps, anderson and cole (1969) the pubertal interval in the domestic horse mare extends from eight to 20 months. nishikawa and hafez (1962) state that puberty is attained at between 15 and 18 months, while sugie and nishikawa (1954) give 16 to 17 months and belonje (1958) 12 to 24 months. in the case of donkey mares, kupfer (1928) found that ovulation commenced at one year of age. sexual maturity on the other hand is reported to occur at 20 months (sugie and nishikawa 1954) and 25 to 28 months (nishikawa and hafez 1962) in the horse mare, and at 24 to 30 months in the donkey mare (kupfer 1928). the mating and foaling season as is the case with the zebra stallion (smuts 1976), sexual activity in the . mare is largely seasonal (fig. 6) with the majority of conceptions and births occurring in the summer (wet) season. the zebra is thus similar to most seasonal breeding species in that the mating season is initiated at such a time that environmental conditions are optimal at parturition, when both mother and offspring are most susceptible to detrimental effects (sadleir 1969). in this species mating takes place very lis soon after parturition (8-9 days) and for all practical purposes they could be regarded as occurring at the same time (bearing in mind that the gestation period is just over a year or ± 375 days). from table 4 it can be seen that peak foaling normally occurs in january (3 years out of 4) followed by december and then february. a similar pattern is obtained when studying the foaling season as obtained using embryos and foetuses (table 5). in addition to these, an average of 44,6% of the total foal crop is produced during december and january (table 6) and an average of 85,2% during the sixmonth summer season (october to march), i.e. only 14,8% of foals are born during the dry winter months. table 6 further indicates that on average 85,6% of all conceptions take place during the summer. this again correlates well with the previously-mentioned foaling activity and proves that both mating and foaling are synchronised with the summer season when green food is normally freely available. in the case of the domestic horse it has also been proved that reproductive activity is largely confined to the wet season (quinlan, van rensburg and steyn 1951; van niekerk 1967). from fig. 6 it is clear that conception and foaling closely follow daylight hours, temperature and rainfall. daylight hours and temperature are, however, very constant from one year to the next, while rainfall may vary considerably. since relative monthly conception percentages over the four-year period varied considerably, it is obvious that some factor other than daylight hours (photoperiod) or temperature must be causing the variation. when viewing the rainfall histograms it can be seen that relative monthly conception percentages follow a similar erratic trend. what in fact appears to happen may be summarised as follows: the first spring rains (usually october to november) are followed by a marked increase in conceptions. this is especially obvious when conception rate during the previous summer was low and many anoestrous mares enter the winter season (e.g. the winter for 1970 was eight months long extending from march to october and the conception rate was only 63,6%) . here the first rains which fell in october 1970, were followed by a rapid increase in breeding activity and consequently an increased relative monthly conception percentage. this then remained practically constant for the next four months so that 42 out of 46 adult mares collected during the ensuing winter were pregnant (conception rate = 91,3%). the fact that relative monthly conception percentages remained so consistent between january and march is believed to be due to the good rainfall recorded during december and january and to the large numbers of non-pregnant mares which entered the 1970/71 breeding season. the dependence of high relative monthly conception percentages on rainfall is best illustrated by the 1969/70 summer season. during this season rainfall was extremely variable from one month to the next with only 36 and six mm of rain being recorded during november and january respectively. the total rainfall was also unusually low (total = 119 370,5 mm for october 1969 to march 1970). when viewing the corresponding relative monthly conception percentages, one finds a similar pattern in that the initially high conception for october is followed by a marked drop during november with an increase during december and january. the increase during january is probably due to the good rains (90 mm) experienced during december, while the drop during february is due to the poor rains (6 mm) of january. the erratic rainfall for this summer season also appears to have resulted in an unusually low conception rate only 63,6%. erratic relative monthly conception percentages are probably largely due to the fact that foaling mares fail to show the normal foal heat and only do so when conditions are favourable. ovaries taken from a sample of 11 non-pregnant mares collected during the winter of 1970 proved that 10 were in deep anoestrus, the largest follicle being 12 millimetres in diameter. additionally there were no fresh corpora lutea (except for one specimen) but numerous very small corpora albicantia. most of these mares had young, twoto six-month old, foals at foot. unfortunately the monthly foaling season counts were not sensitive enough to show up the erratic state of the previous year's conception. this is partly due to the fact that field counts depict foal survival, i.e. births minus early postnatal mortality, and not merely births. the early start of conceptions during september 1969 also demonstrates that anoestrous mares or those foaling early in the season will come on heat and conceive early if conditions are favourable. since the foaling season peaks illustrated in fig. 6 have been drawn above their corresponding months and rainfall it can be seen that the foaling season also approximates rainfall. this, however, is due to the fact that the gestation period is just over 12 months long and that through the process of natural selection, mating has been geared and genetically fixed to take place during the summer months. throughout this time the animals probably respond to a variety of environmental stimuli of which the most important are daylength, temperature, rainfall, vegetation and possibly certain social factors. rainfall does, however, either hasten or delay the onset of breeding activity and causes a certain amount of in-season irregularity. although rainfall may be an important stimulus, increased nutritional intake due to the growth flush following the first spring showers, is more likely to be the cause of the reproductive activity. in horses, for example, nutritional factors are considered to be responsible for irregularities in the oestrous cycle and for low fertility (quinlan et al. 1951). under the environmental conditions prevailing in the karoo midlands of south africa, belonje (1958) has also demonstrated that fluctuations in the commencement and duration of the breeding of thoroughbred mares was due to the condition of the mare which in turn could be ascribed to nutritional intake. in any natural area the seasonal condition of the veld is largely due to the three climatic variables, rainfall, daylight hours and temperature. however, since temperature and in particular day length are practically 120 seasonal constants, only rainfall and the concomitant increase in green vegetation can cause the summer irregularities in breeding activity. at this stage it would be impossible to decide which vegetation factors were responsible for the onset of mating activity and in-season irregularities. these reactions could, for example, be due either to increased and improved intake, to substances present in green grass, or to direct stimulation of the central nervous system via the eyes. since any, and possibly all, of these could playa role in initiating reproductive activity, only carefully controlled experiments would be able to provide a possible solution. although the inference has been made that oestrogenic substances present in freshly sprouting grasses stimulate breeding in the red kangaroo (megaleia tufa) (newsome 1966) and that some factor present in green grass is necessary for successful reproduction of rabbits (oryctolagus cuniculus) (myers and poole 1962), there is little evidence to suggest that a similar state of affairs may exist in other medium-sized and large mammals. in fact oestrogenic pastures are generally held to have an adverse effect on reproduction, impairing fertility in both ewes and cows (moule, braden and lamond 1963). the actions and effects of plant oestrogens and the experimental problems underlying research in this field are reviewed by samuel (1967). he also mentions that not only is there a variation in the oestrogenic activity of plants under different environmental conditions but that larger doses of oestrogen can inhibit follicular growth by suppressing the secretion of pituitary gonadotrophins, whereas small doses may enhance follicular development. these matters are nevertheless still very much in the experimental phase and valid conclusions would be difficult to make. the importance of photo-stimulation and temperature on the sexual periodicity of the zebra cannot, however, be overlooked. in mammals (amoroso 1955) these two factors are known to have important influences on the regulation of the oestrous cycle and sexual behaviour. in the case of the domestic horse, burkhardt (1947), for example, has shown in a series of controlled experiments that oestrus and ovulation followed by pregnancy could be induced in anoestrous mares by general irradiation with strong artificial light. nishikawa, sugie and harada (1952) similarly were able to stimulate ovarian activity earlier in korean ponies subjected to five hours of extra light added at the end of the day. the ovaries of mares given this treatment began to function some 65 to 80 days earlier than unlighted controls. they were also able to stimulate ovarian function in the normal non-breeding season and in both experiments induced ovarian activity was accompanied by normal fertility. nishikawa and horie (1952) who used similar light regimes on the stallion, reported that extra light accelerated the normal increase in semen volume and brought it forward by about two months. the importance of season on the breeding habits of the horse is aptly demonstrated when mares transported from the northern to the southern hemisphere change over their times of breeding to fit the seasons in their new surroundings (belonje 1960). 121 in his study on zebra in the ngorongoro crater, klingel (1965 and 1969a) states that zebra foals are born during any month of the year, the majority from october to march, with a peak iiljanuary. his distribution of foaling is also very similar to that found in the knp and is likewise correlated with rainfall. fairall (1968) found that zebra in the knp foal from september to march with a peak during the period november to february. on the basis of the findings of the present study it would, however, be extremely difficult to justify the findings of ansell (i 960) whereby it is stated that july to september is the period of peak foaling in zambia. in summary it may be said that zebra in the central district of the knp are seasonal breeders with between 85 and 88% of all births and conceptions occurring during the summer months which extend from october to march. within this interval, december and january are the peak months. in agreement with domestic horse breeds which have a defined breeding season, the zebra displays a reproductive cycle which is synchronised with the photoperiod. in this respect conception percentages rise and fall as does the length of daylight hours (and average daily temperature). of particular importance, however, is rainfall which has a profound effect on the initiation of breeding activity and irregularities in relative monthly conception percentages within a particular season. the effect of rainfall may either be direct, or indirect via increased summer nutritional intake. it may similarly also be said that the breeding cycle of the zebra mare is most probably regulated by the interplay of a variety of environmental and possibly social factors. conception rate and the factors affecting it since the fallopian tubes of visibly non-pregnant mares were not washed out to recover any possible blastocysts, the conception rate of 79,3% must be regarded as the minimum. it is unlikely, nowever, that very many mares in early pregnancy were classified as non-pregnant since the conceptus should be visible by 10 to 15 days after fertilization. in the domestic horse, for example, pregnancy can be diagnosed rectally by palpating the uterine horns and by actually feeling the embryonic mass at 14 days (van niekerk 1965). assuming that all embryos less than 15 days old were overlooked in the present study, it was calculated (smuts 1974a) that the average conception rate would rise from 79,3 to 82,7 percent. a conception rate of 79,3% to 82,7% for the four study years would appear fairly average as a wide range of rainfall and other climatic conditions were experienced during this period. in this respect the 1968/69 summer season may be regarded as being normal, while the 1969/70 and 1970/7 1 summer seasons were dry with expecially erratic conditions prevailing during the 1969/70 season. in contrast to these, the summer of 1971/72 was extremely wet with the highest rainfall in 32 years being experienced in the tshokwanelsatara area (fig. 6) where all material was collected. 122 the conception rates for young mares given in table 3 indicate that the adult rate is achieved just after three years of age. further breakdown of conception data into the various adult age classes (table 7) proves that there are no adult age-specific fertility rate differences and that mares remain fertile into old age. the conception rate differences for the adult age classes in table 7 are due to unequal sampling of the age classes during the different years of collection. conception rates for the four non-breeding seasons experienced during the study period (tabk 8) indicate that conception rate varies considerably from one year to the next. these variations represent the reaction of zebra to their environment. when viewing the data on summer rainfall and the conception rates for the non-breeding season (fig. 6) it can be seen that poor summer rains are not necessarily followed by a low conception rate. in fact the highest conception rates of 88,6% (1966) and 91,3% (1971) were recorded after the driest summers. during these summers (october to march) 350 and 333 mm of rain were recorded respectively (average summer rainfall = 425 mm) . similarly the high summer rainfall experienced during 1971/72 (759 mm highest in 32 years) was followed by a below average conception rate while the almost average rainfall for the 1968/69 summer (414 mm) also produced a sub-average conception rate. lastly the 1969/7 0 summer season with its erratic total rainfall of 37 1 mm produced the lowest ever conception rate of 63,6%. conception success must obviously be due to some factor(s) other than total summer precipitation. the results of the present study indicate that in addition to summer precipitation, conception rate may be influenced by numerous and frequently interrelated factors. firstly the actual months during which the greatest proportion of the summer rains fall, appears to be important. in this respect poor or erratic rains experienced during the months of peak mating activity, namely november to february, may precipitate low conception rates. this was particularly obvious during the 1969/70 breeding season when very little rain was recorded during november and january. the 70% conception rate recorded during the nonbreeding season of 1969 may also be partly ascribed to the relatively poor rainfall of january and february 1969 which are normally the months of peak breeding. in contrast, good and regular summer rains do not necessarily produce a high conception rate. during the 1971/72 summer, for example, high rainfall (759 mm) was followed by a relatively low conception rate (75,6%). this could either be due to excessively wet conditions or it may be linked with the unusually high conception rate of the previous year. the high conception rate (91,3%) recorded during the nonbreeding season of 19 71 warrants special discussion since quite a different factor, namely the effective length of the preceding non-breeding season (winter), appears to have been operative. from fig. 6 it can be seen that 1970 was made up of an unusually long winter extending from february 123 to october (8 months). conception rate recorded during this period was very low (63,6%) with the result that a large number of nonpregnant mares were being subjected to an extended season of sexual inactivity. the below average rainfall registered between october 1970 and april 1971 was, however, sufficient to stimulate mating activity to the extent that 91,3% of all recently foaled and non-pregnant mares were mated. these results may be explained on the basis that anoestrous mares subjected to extended winters are more likely to come into oestrus and conceive during the ensuing mating season (summer) than those subjected to short winters. the same condition may apply to pregnant mares . it thus appears that animals deprived of optimal conditions for extended periods are more likely to respond to these when they do occur. one may therefore postulate that this type of deprivation causes a lowered sexual threshold and consequently more vigorous summer mating activity and increased conception rates. when compared to other associated herbivore species, and in particular potential competitors such as the wildebeest, the average adult conception rate of 79,3% for zebra is decidedly low. the same applies to the adult conception rate range of 63,6 to 91,3 percent. in the central district of the knp adult wildebeest cows (two to 16 + years old) have an average conception rate of 92% (n = 125) while that for one to two-year olds is 32% (n = 31) (braack 1973). in the serengeti wildebeest, watson (1969) reports an average adult conception rate of 96% with an average of 37% for cows in their first reproductive year and 83% for cows in their second year. it is thus obvious that the wildebeest which can conceive successfully during its second year (one year and four months) (watson 1969), generally has a greater breeding potential than the zebra which only conceives successfully during its third year (tables 3 and 7). an average adult conception rate diflerence of 12% for the two species in the knp is further evidence for the greater breeding potential of the wildebeest. when comparing zebra and impala, for which adult fertility rates of 97% (dasmann and mossman 1962) and adult lambing rates of 90% (fairall 1969) have been registered, it becomes clear that in addition to conception rate, success of the zebra population must also depend on increased postnatal survival. it must, however, be borne in mind that successive conceptions in the zebra will tend to occur approximately 20 days later during each successive year. this is due to an average gestation period of 375 days, followed by a foal heat and possible conception about 10 days after parturition. this would eventually put a mare out of phase with the normal mating season (october to march) and since mares stand a reduced chance of conceiving during the winter (april to september), they would only come on heat during the following summer (approximately six months later). since one would expect a certain percentage of mares to be out of phase each year, it would undoubtedly have an effect on the overall conception rate of the population. this pheno124 to october (8 months). conception rate recorded during this period was very low (63,6%) with the result that a large number of nonpregnant mares were being subjected to an extended season of sexual inactivity. the below average rainfall registered between october 1970 and april 1971 was, however, sufficient to stimulate mating activity to the extent that 91,3% of all recently foaled and non-pregnant mares were mated. these results may be explained on the basis that anoestrous mares subjected to extended winters are more likely to come into oestrus and conceive during the ensuing mating season (summer) than those subjected to short winters. the same condition may apply to pregnant mares . it thus appears that animals deprived of optimal conditions for extended periods are more likely to respond to these when they do occur. one may therefore postulate that this type of deprivation causes a lowered sexual threshold and consequently more vigorous summer mating activity and increased conception rates. when compared to other associated herbivore species, and in particular potential competitors such as the wildebeest, the average adult conception rate of 79,3% for zebra is decidedly low. the same applies to the adult conception rate range of 63,6 to 91,3 percent. in the central district of the knp adult wildebeest cows (two to 16 + years old) have an average conception rate of 92% (n = 125) while that for one to two-year olds is 32% (n = 31) (braack 1973). in the serengeti wildebeest, watson (1969) reports an average adult conception rate of 96% with an average of 37% for cows in their first reproductive year and 83% for cows in their second year. it is thus obvious that the wildebeest which can conceive successfully during its second year (one year and four months) (watson 1969), generally has a greater breeding potential than the zebra which only conceives successfully during its third year (tables 3 and 7). an average adult conception rate diflerence of 12% for the two species in the knp is further evidence for the greater breeding potential of the wildebeest. when comparing zebra and impala, for which adult fertility rates of 97% (dasmann and mossman 1962) and adult lambing rates of 90% (fairall 1969) have been registered, it becomes clear that in addition to conception rate, success of the zebra population must also depend on increased postnatal survival. it must, however, be borne in mind that successive conceptions in the zebra will tend to occur approximately 20 days later during each successive year. this is due to an average gestation period of 375 days, followed by a foal heat and possible conception about 10 days after parturition. this would eventually put a mare out of phase with the normal mating season (october to march) and since mares stand a reduced chance of conceiving during the winter (april to september), they would only come on heat during the following summer (approximately six months later). since one would expect a certain percentage of mares to be out of phase each year, it would undoubtedly have an effect on the overall conception rate of the population. this pheno124 menon would also partly account for the low average conception rate found in zebra when compared to other herbivores with gestation periods shorter than one year. unfortunately it is difficult to compare conception rates in the zebra with the domestic horse where a variety of additional factors may influence conception. in certain areas, however, horses are allowed free range and association with stallions and in these cases conception rate is decidedly higher than in the zebra. day (1939), for example, mentions that ponies running wild have. a conception rate of 95% in britain. in south africa, van rensburg and van heerden (1953) obtained a conception rate of 87,5% for free roaming mares. these higher rates could partly be due to the shorter gestation period in domestic horses (329-345 days; asdell 1946), with the result that animals are less likely to become out of phase with their normal mating season than are zebra. additionally supplementary feeds during times of hardship would favour the domestic equine over the wild zebra. ovarian activity during the oestrus cycle from table 9 it can be seen that the mean mass and dimensions of the pro-oestrous ovary exceed those of the other three phases. similarly this ovary contains the follicle with the greatest diameter. occasionally late dioestrous ovaries are difficult to distinguish from pro-oestrous specimens and they may attain a mass of almost 40 g due to follicular development. the important difference between the various phases of the oestrous cycle, however, lies in the development of the corpus luteum. in pro-oestrous ovaries there mayor may not be a corpus luteum with a diameter in excess of 10 mm, while those with diameters of less than 10 mm are generally very old and quite possibly the remains of corpora lutea from the previous breeding season. regressing corpora lutea generally have a very characteristic pear shape, the thin tip being in apposition to the ovarian fossa. as regression progresses the body becomes darker in colour (ochre in formalin-fixed specimens) and thinner, until a very narrow streak of scar tissue remains (the corpus albicans) . during metoestrus, the corpus luteum attains its maximum size (32 mm) after which it gradually regresses. during dioestrus regression continues, the corpus luteum, however, still being present by the time the succeeding cycle commences. in this respect the zebra mare is similar to the domestic horse (hammond and wodzicki 1941) in that the corpus luteum of a previous generation (cycle) can exist, although reduced in size, side by side with one of the last generation and a third from the new generation. in addition, corpora btea may continue to regress for at least one year and during these stages they assume the thin dark tapering shape (± 10 x 2 mm) illustrated in the anoestrous ovaries (fig. 7l. similar structures are, however, also present in the other phases of the cycle. 125 ovarian activity during pregnancy according to rowlands (in litt.), equids differ characteristically from other mammals in that during early pregnancy a new crop of follicles is produced which ovulate to form secondary corpora lutea as the main source of progesterone, following the regression of the primary corpus luteum. this state of affairs differs from hystricomorph rodents and some primates where follicles luteinise at the same time, or after other follicl es ovulate to form the primary corpus luteum of pregnancy. these bodies persist as accessory corpora lutea (rowlands in litt.). the word "secondary" is therefore used to describe corpora lutea which have formed during pregnancy through luteinisation of ruptured follicles, while accessory corpora lutea describe the bodies formed through luteinisation of unruptured follicles. by referring to fig's 8, 9 and 10 it can be seen that ovarian activity is largely in abeyance between approximately 190 days and full term pregnancy. during the latter half of pregnancy the ovary is typified by the absence of follicles with diameters greater than 12 millimetres. small regressing follicles may, however, be quite abundant (table 10), th eir numbers generally varying from one animal to the next. in some near-term specimens over 50 tiny (one to 10 mm) follicles were counted, while in others there were only four or five . in contrast to the latter half of pregnancy, the first 190 days show a number of interesting phases of activity. unfortunately few specimens were available for the first 60 days and these additionally indicated that there was considerable variation between individual mares. although the small sample precludes reliable conclusions, it would appear that ovarian activity increases gradually after conception to reach a peak between the 40th and iloth days of pregnancy (fig's 9 and 10). this peak in turn appears to be correlated with the secretion of serum gonadotrophin (smuts 1974a). data collected from my series of pregnant mares are very similar to those obtained by king (1965) from 20 pregnant grant's zebra mares shot in east africa . he found that large (20 mm) follicles were present up to 119 days of pregnancy after which their diameter decreased. the only secondary corpus luteum he could positively identify was at 98 days (as was one of the cases in the present study). he also mentions that the corpus luteum of pregnancy begins to regress at 22 weeks of pregnancy and that the presence of a large corpus luteum for more than five months was probably a result of the formation of fresh corpora lutea (secondary corpora luteal during early pregnancy. in the present study, however, mares lacking secondary corpora lutea frequently still had the corpus luteum of pregnancy visible after 200 days of pregnancy and in a few instances up to 300 days of pregnancy. when compared to mares of various domestic horse breeds, it is obvious that the ovarian cycle is very similar in the zebra mare. differences, however, firstly include the dimensions and mass of the glands in the two species. in the horse, single ovaries frequently have a mass 126 of up to 80 g (sisson and grossman 1953) with the normal mass ranging between 40 and 70 g (hammond and wodzicki 1941). during early pregnancy when follicular activity is high, the ovaries may attain a mass (paired) of over 150 g (van rensburg and van niekerk 1968), while in the zebra mare the greatest mass (paired) recorded was 89,1 g in a 69-day pregnant specimen. similarly the length of the ovaries in the horse is greater, being from 5,5 to eight cm (hammond and wodzicki 1941), while in the zebra they range from 2,5 to 7,4 cm in adult individuals (tables 9 and 10). in addition to these, ovarian activity in burchell's zebra and the horse also differs in certain other respects. the stimulus for follicular development and secondary ovulations during early pregnancy appears to be greater in the horse (cole, howell and hart 1931; king 1966). secondary ovulations and the formation of secondary corpora lutea also appear to occur at an earlier date in the horse mare. day (1940), for example, observed the occurrence of ovulation on the 23rd day of pregnancy, while allen (1971) found that the average time of the first ovulation was 52,5 days and that for second ovulations, 66 days. van rensburg and van niekerk (1968) also found that the rise in follicular activity after the 40th day resulted in secondary ovulations but that corpora lutea formed after the 50th day were produced by luteinization of unruptured follicles. rowlands (1949), however, feels that early luteal structures in his material were all formed from follicles that had ovulated. the early formation of secondary corpora lutea in the horse is, however, by no means a rule, as allen (1971) recorded two relatively late in gestation (after 80 days). all these observations indicate that not only is there a tremendous variation between breeds, but even within breeds from the same area. development of the corpus luteum of pregnancy is similar in both species, the body reaching its maximum size at 14 days of pregnancy in the horse (harrison 1946). according to van rensburg and van niekerk (1968), the primary corpus luteum of pregnancy in the horse remains quite active at least up to the third month of pregnancy. in grevy's zebra, the corpus luteum of pregnancy was found to remain functional for at least 60 days according to king (1966) who also felt that the relatively few secondary corpora lutea produced in zebra species (burchell's and grevy's) was compensated for by an extended period of activity of the corpus luteum of pregnancy. the more recent chemical assays undertaken by van rensburg and van niekerk (1968), however, tend to invalidate these observations since they found that activity continued up to three months and concluded that secondary corpora lutea in the horse were essentially supplementary sources of progesterone. as is the case with the zebra mare in the present study, the horse appears to display differential ovarian activity. hancock (1948) and arthur (1958), for example, found that 52,2 and 63,0% respectiv~ly of ovulations were from the left ovary in cycling mares (60% in the present study). in pregnant horse mares similar results have been obtained and here allen (1971) reports a percentage of 53 for ovulations 127 from the left ovary, while osborne (1966) recorded a significant 55,6% from a total of 3 056 single and multiple ovulations. in contrast to the zebra mare where slightly more implantations were recorded from the left: uterine horn (55,6%; n = 63 mares), more implantations appear to occur in the right horn of the horse (see amoroso, hancock and rowlands 1948, and hancock 1948). transuterine migration, however, occurs at approximately the same rate in both species 41,7% in the zebra and 40,0 and 50,0% in the horse (hancock 1948 and amoroso et al. 1948, respectively). as is the case with the zebra, most migrations in the horse are from the left horn of the uterus to the right. twinning in contrast to the domestic horse which occasionally produces viable twins (asdell 1946), a zebra mare has not yet been observed with twins. i n fact, the only record of a twin conception is given by wackernagel (1965). these were aborted in the eighth month. in the domestic horse the incidence of twin pregnancies varies between one and five per cent (nishikawa and hafez 1962). usually, however, one of the twins dies before birth and twin births occur only in very few cases. hancock (1948) has also shown that although twin conceptions may occur in up to 20,8% of pregnancies, foetal resorption may be high, especially before the 70th day of pregnancy. in this respect he quotes live twin birth rates of between 0,5 and 1,1 %. klingel (1965 and 1969a) also mentions that no twin births have been recorded from zebra in the wild, while the absence of twin conceptions in the sample of 215 pregnant mares examined during the present study supports his observations. king (1965) also found no evidence of twins in the 20 pregnancies he examined. the zebra, in common with other large african herbivores, is thus a monotocous species. reproductive senescence in the sample of 236 adult mares examined, the oldest two were estimated to be 17 to 18 years old (table 7). both these specimens were pregnant. since free-ranging zebra in the knp seldom survive to over 20 years of age (smuts 1974b) it is very unlikely that reproductive senescence ever occurs in the population. this may, however, be a factor in zoological gardens where authenticated ages of up to 28 years have been recorded (flower 1931). acknowledgements the author is particularly indebted to the following people and instances: the chief director of national parks, dr. r. knobel and the national parks board of trustees under whose auspices this study was undertaken. professor j. du p. bothma, dr. u. de v. pienaar, dr. j. 128 hanks, dr. j. f. w. grosskopf and dr. w. h. gerneke who reviewed parts of my original d.sc. dissertation. members of the research and management sections in the knp who assisted with various aspects of the field work. my parents and my wife for assistance and encouragement throughout the duration of the project. references allen, w. e. 1971. the occurrence of ovulation during pregnancy in the mare. vet. rec. 88 :508-509. amoroso, e. c. 1955. hormone control of the oestrous cycle. vet. rec. 67: 1072-1084. amoroso, e. c., j. l. hancock and i. w. rowlands. 1948. ovarian activity in the pregnant mare. nature, lond. 161 :355-356. amoroso, e. c. and i. w. rowlands. 1951. honnonal effects in the pregnant mare and foetal foal.]. endocrinol. 7 :50-53. ansell, w. f. h. 1960. the breeding of some larger mammals in northern rhodesia. proc. 7.o0l. soc. lond. 134:251-274. arthur, g. h. 1958. an analysis of the reproductive function of mares based on post-mortem examination. vet. rec. 70:682-686. asdell, s. a. 1946 . patterns of mammalian reproduction. new york: comstock publishing company inc. belonje, c. w. a. 1949. observations on reproduction in the thoroughbred mare.]. s. afr. vet. med. ass. 20:21-33. belonje, c. w. a. 1958. fertility and infertility of thoroughbred mares under environmental conditions prevailing in the karoo midlands of south africa. d.y.sc. thesis, faculty of yet. sc., univ. of pretoria. (unpublished). belonje, c. w. a. 1960. the physiology of reproduction in the mare with special reference to the thoroughbred.]. s. afr. vet . med. ass. 31:115-128. benesch, f. and j. wright. 1957. veterinary obstetrics. london: bailliers, tindall and cox. braack, h. h. 1973. population dynamics of the blue wildebeest, connochaetes taurinus taurinus (burchell, 1823) in the central district of the kruger national park. project report submitted to the univ. of rhodesia. 85 pp. (unpublished). burkhardt,.j. 1947. transition from anoestrus in the mare and the effects of artificial lighting.}. agric. sci. 37:64-68. cole, h. h., g. h. hart, w. r. lyons and h. r. catchpole. 1933. the development and hormonal content of foetal horse gonads. anat. rec. 56 :275-292. cole, h. h ., c. e. howell and g. h. hart. 1931. the changes occurring in the ovary of the mare during pregnancy. anat. rec. 49: 199-209. 129 cupps, p. t., l. l. anderson and h. h. cole. 1969. the estrous cycle. pages 217-250 in "reproduction in domestic animals", 2nd ed. eds. h. h. cole and p. t. cupps. new york: academic press. dasmann, r. f. and a. a. mossman. 1962. population studies of impala in southern rhodesia.j. wildl. mgmt 26 :262-268. day, f. t. 1939. some observations on the causes of infertility in horse breeding. vet. rec. 51 :581-587. day, f. t. 1940. clinical and experimental observations on reproduction in the mare.j. agric. sci. 30 :244-261. donovan, b. t. 1966. control of follicular growth and ovulation. pages 3-26 in "reproduction in the female mammal". eds. g. e. lamming and e. c. amoroso. london: butterworth and co. du plessis, j. l. 1964. some observations and data in thoroughbred breeding.}. s. afr. vet. med. ass. 35 :215--221. eckstein, p. and s. zuckerman. 1962. morphology of the reproductive tracts. pages 43-155 in "marshall's physiology of the reproduction", vol. 1(1). 3rd ed. ed. a. s. parkes. london: longmans. fairall, n. 1968. the reproductive seasons of some mammals in the kruger national park. zool . afr. 3: 189-210. fairall, n. 1969. the use of the eye lens technique in deriving the age structure and life table of an impala (aepyceros melampus) population. koedoe 12: 90-96. flower, s. s. 1931. contribution to our knowledge of the duration of life in vertebrate animals. v. mammals. proc. 1.ool. soc . lond. 10 1 : 145--235. hammond, j. and k. wodzicki. 1941. anatomical and histological changes during the oestrous cycle in the mare. proc. r . soc. series b. 858.130:1-23. .. hancock, j. l. 1948. notes on oestrus, ovulation and pregnancy in the mare. vet . rec. 60:679-682. harrison, r.j. 1946. the early development of the corpus luteum in themare.j. anat. lond. 80:160-166. huggett, a. st. g. and w. f. widdas. 1951. the relationship between mammalian foetal weight and conception age.j. physiol. 114: 306-317. imes, g. d. and g. l. smuts. 1975. gross and microscopic observations of ovarian abnormalities from five burchell's zebra, equus burchelli antiquorum, smith 1841. onderstepoortj. vet res. 42: 109-116. joubert, d. m. 1963. puberty infemale farm animals. anim. breed. abstr. 31 :295-306. kenneth, j. h. and g. r. ritchie. 1953. gestation periods: a table and bibliography. technical communications no.5 of the commonwealth bureau of animal breeding and genetics, edinburgh. king,j. m. 1965. a field guide to the reproduction of the grant's zebra and grevy's zebra. e. afr. wildl.j. 3 :99-117. 130 king, j. m. 1966. comparative aspects of reproduction in the equidae. ph.d. thesis, university of cambridge. klingel, h. 1965. notes on the biology of the plains zebra (equus quagga boehmi matschie). e. afr. wildl.}. 3 :86-88. klingel, h. 1967. soziale organisation und verhalten freilebender steppenzebras. z. tierpsychol. 24 :580-624. klingel, h. 1969a. reproduction in the plains zebra (equus burchelli boehmi): behaviour and ecological factors. }. reprod. fert. 6: 339-345. klingel, h. 1969b. the social organisation and population ecology of the plains zebra (equus quagga). zool. afro 4 :249-263. klingel, h. and u. klingel. 1965. die geburt eines zebras (equus quagga boehmie matschie). z. tierpsychol. 23:72-76. kupfer, m. 1928. the sexual cycle of female domesticated mammals. dept. of agric. 13th and 14th reports of the director of vet. education and research, part. i.: 1211-1270. moule, g. r., a. w. h. braden and d. r. lamond. 1963. the significance of oestrogens in pasture plants in relation to animal production. anim. breed. abstr. 31: 139-157. myers, k. and w. e. poole. 1962. a study of the biology of the wild rabbit, oryctolagus cuniculus (l), in confined populations iii. reproduction. austr.}. zoo!. 10:225-267. newsome, a. e. 1966. the influence of food on breeding in the red kangaroo in central australia. csiro wildl. res. 11: 187-196. nishikawa, y. and e. s. e. hafez. 1962. the reproduction of horses. pages 266-276 in "reproduction in farm animals". ed. e.s.e. hafez. philadelphia: lea and febiger. nishikawa, y. and t. horie. 1952. studies on the effect of daylength on the function of the testes. bull. natn. inst. agric. sci. {japan} ser. g. 3: 45. from: anim. breed. abstr. 22: abs. no. 439. nishikawa, y., t. sugie and n. harada. 1952. studies on the effect of daylength on reproductive function in horses. i. effect of daylength on the function of the ovary. bull. natn. inst. agric. sci., japan ser. g. 3: 35-44. from: anim. breed. abstr. 22: abs. no. 436. osborne, v. e. 1966. an analysis of the pattern of ovulation as it occurs in the annual reproductive cycle of the mare in australia. aust. vet.}. 42:149-154. quinlan, j., s. w. j. van rensburg and h. p. steyn. 1951. the oestrous cycle of the mare when maintained under stable conditions with restricted exercise at onderstepoort. onderstepoort}. vet. res. 25: 105-119. rowlands, i. w. 1949. serum gonadotrophin and ovarian activity in the pregnant mare.}. endocrin. 6: 184-191. sadleir, r. m. f. s. 1969. the ecology oj reproduction in wild and domestic mammals. london: methuen and co. ltd. samuel, d. e. 1967. a review of the effects of plant estrogenic substances on animal reproduction. ohio}. sci. 67 :308-312. 131 sato, s. and s. hoshi. 1934. studies on reproduction in the mare. iii. the oestrous duration and oestrous cycle.}. jap. soc. vet. sci . 13:237-251. sisson, s. and j. d. grossman. 1953. pages 581-594 in "the anatomy of the domestic animals". fourth ed. revised . philadelphia: w. b. saunders company. smuts, g. l. 1972. seasonal movements, migration and age determination of burchell's zebra (equus burchelli antiquo rum, h. smith, 1841) in the kruger national park. m.se. wildl. mgmt. thesis, university of pretoria. 170 pp. smuts, g. l. 1974a. growth, reproduction and population characteristics of burchell's zebra (equus burchelli antiquorum, h. smith, 1841) in the kruger national park. d.se. wildl. mgmt. dissertation . university of pretoria. 268 pp. smuts, g. l. 1974b. age determination in burchell's zebra (equus burchelli antiquorum) from the kruger national park.}. sth afr. wildl . mgmt ass., 4: 103-115. smuts, g. l. 1975. preand post-natal growth phenomena of burchell's zebra (equus burchelli antiquorum). koedoe 18 :69-102. smuts, g. l. 1976 . reproduction in the zebra stallion (equus burchelli antiquorum) from the kruger national park. zoo i. afro (in press). sugie, t. and y. nishikawa. 1954. studies on sexual maturity in the mare. bull. natn. inst. agric. sci. tokyo (chiba). g. 8: 151-159. from anim. breed. abs. 23. abs. no. 1602. trumler, e. 1958. beobachtungen an den b6hmzebras des george von opel freigeheges fur tierforschunge. v: 1. das paarungsverhalten. saugetierk. mitt. 6:1-48. van niekerk, c. h. 1965. the early diagnosis of pregnancy, the development of the foetal membranes and nidation in the mare. jl. s. afr. vet. med. ass. 36 :483-488. van niekerk, c. h. 1967. pattern of the oestrous cycle ofmares.jl. s. afr. vet. med. ass . 38 :299-307. van rensburg, s. w . j., andj. s. van heerden. 1953. infertility in mares caused by ovarian dysfunction. onderstepoort}. vet. res. 26 :285-313. van rensburg, s. j. and c. h. van niekerk. 1968. ovarian function, follicular oestradiol-17b and luteal progesterone and 20 hydroxy-pregn-4-en-3-one in cycling and pregnant equines. onderstepoort}. vet. res. 35:301-318. wackernagel, h. 1965. grant's zebra (equus burchelli boehmi) at basle zoo a contribution to breeding biology. int. zoo yb. 5 :38-41. watson, r. m. 1969. reproduction of wildebeest (connochaetes taurinus albojubatus thomas) in the serengeti region, and its significance to conservation.}. reprod. fert. suppl. 6 :287-310. witherspoon, d. m. and r. b. talbot. 1970. ovulation site in the mare .}. am. vet. med. assn. 157: 1452-1459. 132 page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 page 14 page 15 page 16 page 17 page 18 page 19 page 20 page 21 page 22 last.pdf page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 page 14 page 15 page 16 page 17 page 18 page 19 page 20 page 21 page 22 page 23 turner.qxd micromorphology of the springbok louse damalinia (tricholipeurus) antidorcus found at the rietvlei nature reserve near pretoria, south africa m.l. turner turner, m.l. 2006. micromorphology of the springbok louse damalinia (tricholipeurus) antidorcus found at the rietvlei nature reserve near pretoria, south africa. koedoe 49(1): 79–84. pretoria. issn 0075-6458. this report provides additional information on the morphology of damalinia (tricholipeurus) antidorcus, previously described only by light microscopy. live lice were collected and prepared for viewing by sem. micromorphological features investigated included aspects of the forehead, mandibles and spiracles. the average length of the males was 2.0 mm and females 2.04 mm. the anterior hyalin region of the head was notably emarginated. a well-developed medial groove on the ventral surface of the head between the mandibles was noted. the surface of the epipharynx was smooth. the labrum with its typical scale-like appearance gradually changed into the pulvinal area of the clypeus. both genders had notched left mandibles. the mandibles were highly serrated on the oral surfaces. situated ventrally below and caudal to the mandibles were a pair of labial palpi. the immediate area between, and surrounding the labial palpi was littered with numerous spiked setae and two well developed larger setae raised at their bases. the tips of the palpi bore three larger and longer sensory setae. palpi of this nature are described here for the first time in d. antidorcus. key words: ectoparasites, lice, springbok. m.l. turner, electron microscope unit, medunsa campus, university of limpopo, 0204 republic of south africa. issn 0075-6458 79 koedoe 49/1 (2006) introduction the springbok (antidorcas marsupialis) is found in many reserves and game farms in south africa. ledger (1980) recorded damalinia (=tricholipeurus) (bedford1931) antidorcus as a common ectoparasite (mallophagean) of the springbok (fig. 1). few of the parasitic lice species have been described using the scanning electron microscope (sem) and only line diagrams of the first description of d. antidorcus by bedford (1931) and an insert of a second sketch after werneck (1950) as recorded by ledger (1980) were found in an extensive literature search. these diagrams were drawn from light microscope (lm) images obtained from the technology of the early twentieth century during 1931. this study was performed to serve as an aid in the identification of d. antidorcus and to add to the existing detail found on its morphology. the superorder psocodea comprises two orders, one of which is the phthiraptera. phthiraptera have no free-living stages and are all obligate ectoparasites found on almost all avians and approximately a quarter of all mammals (smith 2000; soler cruz 1995). lice are further divided into four suborders namely amblycera, ischnocera, rhyncophthirina (a monogeneric group found on elephants and warthog only) and anoplura. amblycera and ischnocera are of the biting (mallophaga) type whilst the latter (anoplura) are commonly known as the sucking lice (smith 2000). the genus damalinia comprises three subgenera (soler cruz & martin mateo 1998,): d. tricholipeurus, d. cervicola and d. damalinia (bovicola, ledger 1980). turner.qxd 2006/04/17 10:34 pm page 79 materials and methods live lice were collected from springbok in the rietvlei nature reserve situated to the south east of pretoria during an organised culling session. the springbok were healthy and not heavily infested, thus only six female lice, in varying stages of maturity, and two males were found in the area around the base of the ear. the specimens were processed for scanning electron microscopy as described by turner et al. (2004) and viewed in a leica 420 stereoscan. results the sem investigation revealed several micromorphological specialisations not previously demonstrated in bedford’s (1931) light microscopically derived line diagrams or description of d. antidorcus. the following specialisations were revealed when the lice were viewed in the scanning electron microscope: the total average length of the males was 2.0 mm versus 2.04 mm for females, a deviation (0.02 mm) that was barely notable to the above-mentioned measurements. this slight deviation could be ascribed to the difference in the resolving power of the lightand electron microscopes and likely due to the small number of lice collected. the ventral surface of the head clearly demonstrated the structures of the preantennal regions such as ventral carina, pulvinus, labrum, mandibles and clypeus (turner 2003) (fig. 2). the anterior hyalin region of the head was notably emarginated. a welldeveloped medial groove on the ventral surface of the head between the mandibles was noted (this groove fits snuggly over the host’s hairs and probably also facilitates attachment) (turner et al. 2002). the labrum that lies within the medial groove, appeared to be well developed and was found anterior to, and above the mandibles. its scale-like surface flowed with a gradual changeover into the smooth pulvinal area of the clypeus (fig. 3). in both genders, the left mandibles were notched on the cutting edges (fig. 4). this could aid in rasping the skin and scraping up epidermal exfoliation during feeding. the left and right mandibles were serrated on the oral surfaces (fig. 5) especially koedoe 49/1 (2006) 80 issn 0075-6458 fig. 1. a low magnification sem micrograph of the lateral view d. antidorcus. turner.qxd 2006/04/17 10:34 pm page 80 notable when viewed laterally. the ridged serrations probably facilitate attachment to the host’s hair, especially in extreme circumstances such as when the springbok seeks refuge whilst running in dense tall grassy areas, during life threatening situations. situated ventrally below and caudally to the mandibles were a pair of labial palpi. the immediate area between, and surrounding the labial palpi contained numerous spinelike, spiked setae and two well developed larger, flexible setae, which were raised at their bases. the spiked setae were similar to the epipharyngeal dentate structures found in the blesbuck louse damalinia crenelata issn 0075-6458 81 koedoe 49/1 (2006) fig. 2. the ventral aspect of the head, (a) first antennal segment, (c) clypeus, (la) labrum, (m) mandibles, (p) pulvinus, (t) trabecular region, (arrow) epipharynx, (*) labial palpi. fig. 3. an enlarged view of the labrum (l) showing the gradual structural changeover to the pulvinus (p). (c) = clypeal border. fig. 4. sem of the mandible showing the notched cutting edge (arrow). fig. 5. a high magnification of the oral surface of the right mandible where (s) represents the serrated surface. fig. 6. the pair of labial palpi with numerous spiked or spinous setae (arrows) and two well developed larger sensory setae (s), which are raised at their bases. the tips of the palpi bore three larger and longer sensory setae (*). turner.qxd 2006/04/17 10:34 pm page 81 koedoe 49/1 (2006) 82 issn 0075-6458 fig. 8. the distal tip of the antennae constituted 10 apical setae (s). fig. 9. the 2 pore organs (p) with three plate organs showing the characteristic radiating pattern of slitlike openings. fig. 7. the triangular trabeculum (t). (a) = antenna. (turner 2003) whereas the surface of the epipharynx was smooth in d. antidorcus. the tips of the palpi bore three larger and longer sensory setae. palpi of this nature were described here for the first time in d. antidorcus (fig. 6). the three segmented antennae were directed caudally and showed thick, short, robust first segments in the male whilst the first segments in the female were much shorter and thinner. the first segment was well protected by a protruding trabeculum (fig. 7). the shape of the trabeculum was sexually dimorphic, as that of the male was more acutely angled and more caudally directed than that of the female. the well-developed peg organ (sensilla basoconica) on the distal tip of the antennae consisted of 10 apical setae which varied in length (fig. 8). antennal tuft organs (sensillae coeloconica) were absent within the two pore organs. adjacent to the pore organs were three plate organs, with their characteristic radiating pattern of slitlike openings (fig. 9). unlike the plate organs found on b. caprae (sebei et al, 2004), a louse infesting goats, which are surrounded by individually raised skeletal plates, those of d. antidorcus were each encompassed in a single plate. this could be of taxonomic importance during sem investigations as these characteristics are not visible under the light microscope. the prothorax and mesothorax were clearly distinguishable. the posterior edge of the mesothorax showed some sexual dimorphism in that the male (fig. 10), had a regular transverse row or fringe of seta on, and overlapping, the posterior margin whereas the row of setae in the female (fig. 11) was irregularly arranged and situated notably further from the margin. the edge was straight in the male whilst more rounded and pointed in the female. added dimorphism in the same figures, was the slight variation in the posterior parietal area of the edge of the dorsoventrally flattened head. these sexual dimorphic features are recorded here for the first time in d. antidorcus. two thoracic spiracles were noted (fig. 12). each was situated laterally on the left and right side of the mesothorax. the spiracles were well turner.qxd 2006/04/17 10:34 pm page 82 issn 0075-6458 83 koedoe 49/1 (2006) fig. 10 & 11. the posterior edge of the meso-thorax (mt) of the male with its regular transverse fringe (rf) of seta and the dimorphic difference in the female respectively. (p) = parietal area, (pt) = pro-thorax, (if) = irregular fringe. fig. 13. the spiracular lumen was lined with regular circular disc-shaped (d) structures, intersected by trabecular (t) divisions. developed. the spiracular lumen was lined with regular circular disc-shaped structures intersected with trabecular divisions that could probably be likened to the mammal pharynx with its cartilage rings (fig. 13). previous literature suggests that these ‘pharyngeal’ discs and ‘trabeculae’ filter the air during discontinuous respiration and prevent dust particles and other debris from entering (turner 2004). literature reviews did not reveal any terminology for the filtering apparatus as described above. the typical damalinean legs showed extended claws which closed towards fixed, appositional but shorter, thumbs on the distal ends of the tibiae. according to bedford (1931), the mid tibia was narrower and longer than the fore or hind tibiae. six pairs of abdominal spiracles were present whilst the second segment only showed a stigmatal scar with no spiracle. (fig. 14 a & b). the female genitalia were underdeveloped and not worthy of discussion in this study whilst the male parameres only protruded slightly (fig. 15). a comparative investigation on different species of the same genus will be performed in the near future. no other remarkable micromorphology was noted. this investigation does not purport to be taxonomic in fig. 12. the thoracic spriracle (arrow). turner.qxd 2006/04/17 10:34 pm page 83 nature but was rather intended to assist biologists and veterinarians in the identification of the species and in comparative studies. acknowledgements riaan marais the manager of the reserve, cecil labuschagne for his unselfish contribution during specimen collection. toekie kleynhans at the reception office of rietvlei nature reserve, for always being so friendly and polite. the friends of rietvlei for the use of their facilities. references bedford, g.a.h. 1931. new genera and species of mallophaga. report of the director of veterinary services and animal industry 17: 283–297. ledger, j.a. 1980. publications of the south african institute for medical research: the arthropod parasites of vertebrates in africa south of the sahara, phthiraptera (insecta). johannesburg: south african institute for medical research. (publications of the south african institute for medical research no. 56) funny place for a bracket to close 4: 196–296. sebei, p.j., c.m.e. mccrindle, e.d. green & m.l. turner. 2004. use of sem to confirm the identity of lice infesting communally grazed goat herds. onderstepoort journal of veterinary research 71: 87–92. smith, v.s. 2000. avian louse phylogeny (phthiraptera: ischnocera): a cladistic study based on morphology. phd thesis, university of glasgow. soler cruz, m.d. 1995. antennal sense organs of phthiraptera (insecta). scanning electron microscopy of several species of anoplura. micron 26(1): 7–14. soler cruz, m.d. & m. mateo. 1998. sensory equipment of the antennal flagellum of several species of damalinia (phthiraptera: thrichodectidae). micron 29(6): 431–438. turner, m.l., c. baker & r. marais. 2002. a scanning electron microscope investigation of the waterbuck louse bovicola hilli found at the reitvlei nature reserve. koedoe 45(1): 59–63. turner, m.l. 2003. the micromorphology of the blesbuck louse damalinia (damalinia) crenelata as observed under the sem. koedoe 46(1): 65–71 turner, m.l., c. labuschagne & e.d. green. 2004 the morphology of the african buffalo louse haematopinus bufali as observed under the sem. koedoe 47(2): 83–90. usda, 1970. chewing and sucking lice as parasites of mammals and birds. united states department of agriculture. (technical bulletin; no. 1849.) koedoe 49/1 (2006) 84 issn 0075-6458 fig. 14 a & b. six pairs of abdominal spiracles (fig. b) (arrow) were present whilst the second segment only showed a stigmatal scar (fig. a) (arrow). fig. 15. the male genital area (arrow) = parameres, (a) = abdomen. turner.qxd 2006/04/17 10:34 pm page 84 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <feff004f007000740069006f006e007300200070006f0075007200200063007200e900650072002000640065007300200064006f00630075006d0065006e00740073002000500044004600200064006f007400e900730020006400270075006e00650020007200e90073006f006c007500740069006f006e002000e9006c0065007600e9006500200070006f0075007200200075006e00650020007100750061006c0069007400e90020006400270069006d007000720065007300730069006f006e00200070007200e9007000720065007300730065002e0020005500740069006c006900730065007a0020004100630072006f0062006100740020006f00750020005200650061006400650072002c002000760065007200730069006f006e00200035002e00300020006f007500200075006c007400e9007200690065007500720065002c00200070006f007500720020006c006500730020006f00750076007200690072002e0020004c00270069006e0063006f00720070006f0072006100740069006f006e002000640065007300200070006f006c0069006300650073002000650073007400200072006500710075006900730065002e> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! koedoe 19: 49-62(1976) a revised check-list of birds in the kalahari gemsbok national park m. g . l. mills kalahari gemsbok national park private bag x890 p.o. gemsbokpark 8815 abstract a more complete list of the birds in the kalahari gemsbok national park is given, including the results of three and a half years of observations. a total of 214 species have now been identified for the park, among which 75 are resident throughout the year, 37 are migrants, 14 are nomads and 88 are vagrants. introduction several lists of the birds in the kalahari gemsbok national park, republic of south africa, have been published in recent years (de villiers 1958; prozesky and haagner 1962; broekhuysen, broekhuysen, martin, martin, martin and morgan 1968; maclean 1970). with the exception of maclean (1970) these lists have been drawn up by the authors during short visits to the park. however, maclean only worked in the southern area for 19 months. this list represents three and a half years observations over the entire park, with additional records from liversidge (pen. comm.). several groups of birds, notably the rap tors, pipits, larks and warblers, are difficult to identify in the field. where any doubt exists as to the identity of a species, it has been omitted unless a specimen has been taken. systematic list an indication of the status of each species in the park is given according to the following key (after kemp 1974); r resident (found in the park throughout the year) m migrant (a species known to come to south africa on a regular annual migration) n nomad (a regular, temporary visitor to the area) v vagrant (an irregular, temporary visitor to the area) . the number of each species has been taken from mclachlan and liversidge (1966) and the nomenclature follows the check list of the birds of south africa (s.a.o.s . list committee 1969). 49 ostrich (r) volstruis 6 cape dabchick (v) kaapse duikertjie struthio camelus linnaeus podiceps ruficollis (pallas) 47 white-breasted cormorant (v) witbo rsdu i ker phalacrocorax lucidus (lichtenstein) 50 reed cormorant (v) rietduiker 54 grey heron (v) bloureier 55 black-headed heron (v) phalacrocorax ajricanus (gmelin) ardea cinerea linnaeus swartkopreier ardea melanocephala vigors & children 58 great white egret (v) groot witreier 60 yellow-billed egret (v) geelbek-witreier 61 cattle eg ret (v) bosluisvoel 62 squacco heron (v) ralreier 67 little bittern (v) woudapie 69 night heron (v) nagreier 72 hamerkop (v) egretta alba (linnaeus) egretta intermedia (wagler) ardeola ibis (linnaeus) ardeola ralloides (scopoli) ixobrychus minutus (linnaeus) nycticorax nycticorax (linnaeus) scopus umbretta gmelin 73 marabou stork (v) leptoptilos crumeniferus (lesson) maraboe 75 saddlebill (v) ephippiorhynchus senegalensis (shaw) saalbekooivaar 77 wooly -necked stork (v) ciconia episcopus (boddaert) wolnekooievaar 50 78 white-bellied stork (m) klein swart sprinkaanvoel 79 black stork (v) ciconia abdimii (lichtenstein) ciconia nigra (linnaeus) groot swart sprinkaanvoel 80 white stork (m) groot wit sprinkaanvoel 123 rock kestrel (r) rooivalkie ciconia ciconia (linnaeus) falco tinnunculus linnaeus 126 pigmy falcon (r) dwergvalkie poliohierax semitorquatus a. smith 128 black kite (m) swartwou 129 yellow-billed kite (m) geelbekwou milvus migrans (boddaert) milvus migrans parasitus (daudin) 130 black-shouldered kite (r) elanus caeruleus (desfontaines) blouvalkie 134 tawny eagle (r) kouvoel 135 steppe eagle (m) steppe-arend 139 booted eagle (m) dwergarend 141 african hawk eagle (v) afrikaanse jagarend 142 martial eagle (r) breekoparend 145 brown snake-eagle (r) bruin slangarend aquila rapax (temminck) aquila nipalensis (hodgson) aquila pennata (gmdin) aquila fasciata vieillot polemaetus bellicosus (daudin) circaetus cinereus vieillot 146 black-breasted snake-eagle (r) swartbos slangarend ciraetus pectoralis a. smith 149 fish eagle (v) visarend 51 haliaetus vocifer (daudin) 151 bataleur (r) berghaan 152 jackal buzzard (v) jakkalsvoel 154 buzzard (m) jakkalsvoel 157 ovambo sparrowhawk (v) ov ambosperwer 161 little-banded goshawk (v) klein gebande sperwer 162 gabar goshawk (r) klei n witv alki e 165 chanting goshawk (r) groot witv alk 168 pallid harrier (m) vaal paddavreter 169 black harrier (v) witkruisv alk 170 montagu's harrier (m) blou paddavreter 171 banded harrier hawk (v) gymnogene 189 african quail (n) afrikaanse kwartel terathopius ecaudatus (daudin) buteo rujofuscus (forster) buteo buteo (linnaeus) accipiter ovampensis gurney accipiter badius (gmelin) micronisus gabar (daudin) melierax musicus (daudin) circus macrourus (gmelin) circus maurus (temminck) circus pygargus (linnaeus) polyboroides radiatus (scopoli) cortumix cotumix (linnaeus) 190 harlequin quail (v) bontkwartel cotumix delegorguei delegorgue 192 crownded guinea-fowl (v) numida meleagris (linnaeus) tarentaal 196 kurrichane button-quail (n) bosveldkwartel tjie tumix sylvatica (desfontains) 52 208 purple gallinule (v) porphyrio porphyrio (linnaeus) koningriethaan 212 red-knobbed coot (v) fulica cristata gmelin bleshoender 217 kori bustard (r) otis kori burchell gompou 218 ludwig's bustard (v) otisludwigii (ruppel) ludwigse pou 219 stanley bustard (v) otis denhami children veldpou 224 red-crested korhaan (r) eupodotis rujicrista (a. smith) boskorhaan 225 black korhaan (r) afrotis ajra (linnaeus) swartkorhaan 228 african jacana (v) actophilomis africanus (gmelin) langtoon 238 three-banded sand plover (v) charadrius tricollaris vieillot drieband-strandlopertjie 240 caspian plover (m) asiastiese strandloper 242 crowned plover (r) kiewietjie 245 blacksmith plover (v) bontkiewietjie 251 curlew sandpiper (m) kro mbek-strandloper 253 little stint (m) klein strandloper 256 ruff (m) kemphaan 258 common sandpiper (m) gewone ruiter 53 charadrius asiaticus pallas vanellus coronatus (boddaen) vanellus armatus (burchell) calidris minuta (pontoppidan) calidris minuta (leislerl philomachus pugnax (linnaeus) tringa hypoleucos (linnaeus) 262 marsh sandpiper (m) tringa stagnatilis (bechstein) m 0 erasruiter 263 greenshank (m) tringa nebularia (gunnerus) groenpoot-ruiter 264 wpod sandpiper (m) tringaglareola linnaeus bosruiter 268 whimbrel (m) numenius phaeopus (linnaeus) klein wulp 270 black-winged stilt (v) himantopus himantopus (linnaeus) rooipoot-elsie 275 cape dikkop (r) burhinus capensis (lichtenstein) dikkop 276 burchell's courser (n) cursorius rufus gould bloukop-drawwertjie 277 temminck's courser (v) cursorius temminckii swainson trekdrawwertjie 278 double-banded courser (r) dubbelband-drawwertjie rhinoptilus ajricanus (temminck) 280 bronze-wing courser (v) rhinoptilus chalcopterus (heuglin) bronsvlerkdrawwertjie 304 white -winged lake tern (m) witvlerkmeerswawel childonias leucoptera (temminck) 307 namaq.ua sandgrouse (r) pterocles namaqua (gmdin) kelkiewyn 308 spotted sandgrouse (r) pterocles burchelli w. sclater sandpatrys 311 rock pigeon (v) columba guinea linnaeus bosduif 316 cape turtle dove (r) streptopelia capicola (sundevalj) tortelduif 317 laughing dove (r) rooibors-duifie streptopelia senegalensis (linnaeus) 54 318 namaqua dove (r) namakwaduifie 340 cuckoo (m) koekoek oena capensis (linnaeus) cuculus canorus linnaeus 346 great spotted cuckoo (v) clamator glandarius (linnaeus) groot gevlektekoekoek 348 jacobin cuckoo (m) clamator jacobinus (boddaert) nuwejaarsvoel 352 diederik cuckoo (m) chrysococcyx caprius (boddaert) diedrikkie 359 barn owl (r) tyto alba (scopoli) nonnetjie-uil 360 grass owl (v) tyto capensis (a. smith) grasuil 363 scops owl (v) otus scops (linnaeus) klein ooruil '''364 white-faced owl (r) otusleucotis (temminck) witwang-ooruiltjie 365 pearl-spotted owlet (r) glaucidium perlatum (vieillot) dwerg-uiltjie 368 spotted eagle owl (r) bubo ajricanus (temminck) gevlekte ooruil 369 giant eagle owl (r) bubo lacteus (temminck) reuse-ooruil 371 european nightjar (m) camprimulgus europaeus linnaeus europese naguil 372 rufous-cheeked nightjar (m) rooiwang-naguil caprimulgus rufigena a. smith 378 european swift (m) europese windswawel 380 black swift (m) swart windswawel 55 apus apus (linnaeus) apus barbatus (p. sclater) 383 white-rumped swift (v) apus caffer (lichtenstein) witkrui swi ndswa wel 385 little swift (v) apus ajjinis (gray) klein windswawel 391 white-backed mousebird (v) colius colius (linnaeus) witkruis-muisvot.l 392 red-faced mousebird (v) urocolius indicus (latham) rooiwang-muisvot.l 403 striped kingfisher (v) halcyon chelicuti (stanley) gestreepte visvanger 404 european bee-eater (m) merops apiaster linnaeus europese byvreter 411 swallow-tailed bee-eater (r) miksterbyvreter merops hirundineus (lichtenstein) 412 european roller (m) coracias garrulus linnaeus europese troupant 413 lilac-breasted roller (r) coraciascaudata linnaeus gewone troupant 415 purple roller coracias naevia daudin groot troupant 418 african hoopoe (r) upupa epops bechstein hoephoep 419 red-billed hoopoe (v) phoeniculus purpureus (miller) kakelaar 421 scimitar-bill hoopoe (r) swartbekkakelaar rhinopomastus cyanomelas (vieillot) 424 grey hornbill (v) grys neushoringvot.l tockus nasutus linnaeus 426 yellow-billed hornbill (r) tockusflavirostris (ruppel!) geelbekneushoringvot.l 432 pied barbet (r) lybius leucomelas (boddaert) bont houtkapper 56 450 cardinal woodpecker (r) dendropicosfuscescens (vieillot) kardinaal-spegt 457 monotonous lark (v) mirofrajavanica horsfield bosveldlewerkie 459 fawn-coloured lark (r) mirajra ajricanoides a. smith vaalbruin lewerkie 460 sabota lark (r) mirajrasabotaa. smith sabota-lewerkie 464 dusky lark (v) pinarocorys nigricans (sundevall) donker lewerkie 466 clapper lark (r) mirajra apiata (vieillot) klappertj i e 474 spike-heeled lark (r) vlakvoeltjie certhilauda albojasciata lafresnaye 485 grey-backed finch-lark (n) grys kaffertjie eremopterix verticalis (a. smith) 486 black-eared finch-lark (v) egte kaffertjie eremopterix australis (a. smith) 488 red-capped lark (v) rooikoplewerkie tephrocorys cinerea (gmelin) 490 pink-billed lark (v) rooibeklewerkie calandrella conirostris (sundevall) 492 stark's lark (n) starkse lewerkie 493 european swallow (m) europese swawel 495 white-throated swallow (v) calandrella slarki shelley hirundo rustica linnaeus witkeelswa wel hirundo albigularis strickland 498 pearl-breasted swallow (v) hirundo dimidiata sundevall p~relbo rsswa wel tj i e 502 larger striped swallow (v) hirundo culcullata (boddaert) groot streepswawel 57 504 cliff swallow (r) familieswawel 506 rock martin (r) kransswawel 517 fork-tailed drongo (r) m i kstertbyv an g er 519 golden oriole (m) europese wielewaal 522 pied crow (v) witborskraai 523 black crow (r) swartkraai 525 grey tit (r) piettjoutjou hirundo spilodera (sundevall) hirundo rupestris scopoli dierurus adsimilis (bechstein) oriolus oriolus (linnaeus) corvus albus (miiller) corvus eapensis lichtenstein parus ajer gmelin 531 penduline tit (r) anthoseopus minutus (shaw and nodder) grys kapokvoel 536 pied babbler (v) turdoides bieolor (jardine) wit ka tlagter 544 red-eyed bulbul (n) pyenonotus nigrieans (vieillot) rooioog-tiptol 557 groundscraper thrush (v) turduslitsitsirupa (a. smith) gevlekte lyster 561 short-toed rock thrush (v) korttoon-klipwagter montieola brevi pes (waterhouse) 564 mountain chat (v) bergtapult 566 grey-rumped sickle wing chat (v) bleekt apuit 568 capped wheatear (m) skaapwagter 570 familiar chat (r) spekvreter 58 oenanthe montieola vieillot cereomela sehlegelii (wahlberg) oenanthe pileala (gmelin) cereomela jamiliaris (stephens) 571 layard ' s chat (v) cercomela tractrac (wilkes) klein spekvreter 575 ant-eating chat (r) myrmecocichlaformicivora (vieillot) swartpiek 586 kalahari scrub robin (r) erythropygia paena a. smith wipstert 599 willow warbler (m) phylloscopus trochilus (linnaeus) hofzanger 600 yellow-bellied bush warbler (r) geelbuikbossanger eremomela icteropygialis (lafresnaye) 619 rufous-eared warbler (r) roo i 0 0 rklei njantji e 621 crombek (v) krombek 629 common fantail cisticola (n) prinia pectoralis a. smith sylvietta rujescens (vieillot) gewone veldtingtinkie cisticolajuncidis (rafinesque) 630 desert cisticola (r) cisticola aridula whiterby bleekveld-tinktinkie 650 black-chested prinia (r) priniajlavicans (vieillot) swartbors-langsterttinktinkie 654 spotted flycatcher (m) muscicapa striata (pallas) europese vlieevanger 658 tit-babbler (r) parisoma subcaerulem (vieillot) tjeriktik 661 marl co flycatcher (r) melaenomis mariquensis a . smith marl co vlieevanger 663 chat flycatcher (r) melaenomis injuscatus (a . smith) groot vlieevanger 674 pririt flycatcher (r) batispririt(vieillot) pririt-bonbontrokkie 686 cape wagtail (v) motacilla capensis linnaeus kwikkie 59 689 yellow wagtail (v) geel kwikkie 692 richard's pipit (n) gewone koester 695 buffy pipit (n) vaalkoester 706 lesser grey shrike (m) europese grys laksman 707 fiscal shrike (r) laksman 708 red-backed shrike (m) rooiruglaksman motacilla lutea (gmelin) anthus novaeseelandiae (gmelin) anthus vaalensis shelley lanius minor gmelin lanius collaris linnaeus lanius collurio linnaeus 711 crimson-breasted shrike (r) rooiborsfiskaal liniarius atrococcineus burchell 714 three-streaked tchagra (v) tchagra australis (a. smith) klein rooivlerklaksman 722 bokmakierie (r) malaconotus z.eylonus (linnaeus) 730 white-crowned shrike (v) kremetartlaksman eurocephalus anguitimens a. smith 731 brubru shrike (r) bontroklaksman 735 wattled starling (n) sprinkaanvoel 737 cape glossy starling (r) klein glansspreeu nilaus afer (latham) creatophora cinerea (menschen) lamprotornis nitens (linnaeus) 743 burchell's glossy starling (r) groot glansspreeu lamprotornis australis (a. smith) 764 dusky sunbird (v) namakwa-suikerbekkie 775 cape white-eye (v) kaapse glasogie 60 nectarinia fusca (vieillot) zosterops pallidus swainson 779 buffalo weaver (v) buffelwewer bubalomis albirostris (vieillot) 780 white-browed sparrow-weaver (r) koringvoel plocepasser mahali a. smith 783 sociable weaver (r) familievoel 784 house sparrow (r) engelse mossie 785 great sparrow (n) groot mossie 786 cape sparrow (r) mossie 787 grey-headed sparrow (r) gryskopmossie philetairus socius (latham) passer domesticus (linnaeus) passer motitensis (smith) passer melanurus (muller) passer griseus (vieillot) 789 scaly-feathered finch (r) sporopipes squamifrons (a. smith) baardmannetjie 803 masked weaver (r) ploceus velatus vieillot swartkeel-g eel vin k 805 red-billed quelea (n) quelea quelea (linnaeus) rooibekvink 808 red bishop bird (v) euplectes orix (linnaeus) rooi kaffervink 820 red-headed finch (n) amadina erythrocephala (linnaeus) rooikopmossie 840 violet-eared waxbill (v) uraeginthus granatina (linnaeus) koningrooibekkie 843 common waxbill (v) estrilda astrild (linnaeus) rooibekkie 847 shaft-tailed whydah (r) vidua regia (linnaeus) pylstert 861 blackhead canary (v) serinus alario (linnaeus) swartkop-kanarie 61 866 yellow canary (r) geelsysie 871 lark-like bunting (n) vaal streepkoppie references serinus flaviventris (swainson) emberiza impetuani a. smith. broekhuysen, g. j., m. h. broekhuysen, j. martin e. martin, r. martin and h. k. morgan. 1968. observations on the birdlife in the kalahari gemsbok national park. koedoe ii: 145-160. de villiers, j. s. 1958. a report on the bird life of the kalahari gemsbok national park. koedoe 1: 143-161. kemp, a. c. 1974. the distribution and status of the birds of the kruger national park. koedoe monograph 2. mclachlan, g. r. and r. liversidge. 1966. roberts birds oj south africa. central news agency, south africa. maclean, g. l. 1970. an analysis of the avifauna of the southern kalahari gemsbok national park. zool. afro 5(2):249-273. prozesky, o. p. m. and c. h. haagner. 1962. a check-list of the birds of the kalahari gemsbok park. koedoe 5: 171-182. s.a.o.s . list committee. 1969. check. list oj the birds of south africa. cape town: south african ornithological society. 62 page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 page 14 article information authors: anna hübner1 lý t. phong2 trương s.h. châu1 affiliations: 1german corporation for international development, dong hoi, vietnam2school of hotel and tourism management, hong kong polytechnic university, hong kong correspondence to: anna hübner postal address: kapitän-borgwardt-weg 15, 18119 rostock-hohe düne, germany dates: received: 17 feb. 2013 accepted: 06 nov. 2013 published: 24 june 2014 how to cite this article: hübner, a., phong, l.t. & châu, t.s.h., 2014, ‘good governance and tourism development in protected areas: the case of phong nha-ke bang national park, central vietnam’, koedoe 56(2), art. #1146, 10 pages. http://dx.doi.org/10.4102/ koedoe.v56i2.1146 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. good governance and tourism development in protected areas: the case of phong nha-ke bang national park, central vietnam in this original research... open access • abstract • introduction    • protected area governance and governance principles • research method and design • results    • governance of protected areas in vietnam    • phong nha-ke bang national park       • ownership of land resources and types of tourism site management       • income sources       • management body       • management model       • governance criteria and the public–private for-profit national park model • ethical considerations    • recruitment procedures    • informed consent    • data protection • trustworthiness    • reliability    • validity • discussion • conclusion • acknowledgements    • competing interests    • authors’ contributions • references • footnote abstract top ↑ protected areas are increasingly expected to serve as a natural income-producing resource via the exploitation of recreational and touristic activities. whilst tourism is often considered a viable option for generating income which benefits the conservation of a protected area, there are many cases in which insufficient and opaque planning hinder sustainable development, thereby reducing local benefit sharing and, ultimately, nature conservation. this article delineated and examined factors in governance which may underlie tourism development in protected areas. based on graham, amos and plumptre’s five good governance principles, a specific analysis was made of the phong nha-ke bang national park in central vietnam, which highlighted challenges in the practical implementation of governing principles arising for nature conservation, sustainable tourism development and complex stakeholder environments. despite the limited opportunity of this study to examine the wider national and international context, the discussion facilitated an overview of the factors necessary to understand governance principles and tourism development. this article could serve as a basis for future research, especially with respect to comparative analyses of different management structures existing in vietnam and in other contested centrally steered protected area spaces.conservation implications: this research has shown that tourism and its development, despite a more market-oriented and decentralised policymaking, is a fragmented concept impacted by bureaucratic burden, lack of institutional capacities, top-down processes and little benefit-sharing. there is urgent need for stakeholders – public and private – to reconcile the means of protected areas for the ends (conservation) by clarifying responsibilities as well as structures and processes which determine decision-making. introduction top ↑ parks and protected areas (pas) serve a multiplicity of functions and are confronted by an array of social, political and economic expectations. at the same time, pas stand increasingly in conflict with the preservation of natural resources, rising socio-economic expectations and their productive importance for local populations. there are also conflicts with local government and forest enterprises. land-use issues and suppressed viewpoints of local communities, the depletion of natural resources, illegal activities such as poaching, logging, agricultural use and global environmental change further fuel challenges for pa conservation and management (dearden 2000; eagles, mccool & haynes 2002; gössling 2003; larsen 2008).recreational and touristic activities are increasingly used as a justification for the creation of pas or as an additional income source for their maintenance (tang & jang 2009). tourism, particularly in evolving destinations in developing countries, may not be compatible with the standards of long-term sustainable development, but it generates important income which benefits pa conservation (tosun 2001). many cases have shown that uncoordinated management and lack of transparency in planning hinder sustainable development, thereby reducing local benefit sharing and, ultimately, nature conservation. others caution against overrating the role of tourism in conservation, citing over-dependency on tourism for local populations living in and around pas and against economic valuation of pas as natural ecosystems continue to deteriorate (gössling 2003; semone et al. 2011; united nations development programme [undp] 2010). the call for contributions to this special issue of koedoe highlights that both positive and negative consequences have to be dealt with when managing and planning for tourism and its increasing economic role for pas. it was pointed out during the 5th world’s park congress held in durban in 2003 that different forms of management and governance make a significant impact on the status and evolution of ecosystem services. the performance of governance is dependent on decision-making and power-relationships based on objectives, accountability and capacities (including authority, legitimacy and finances) of a variety of stakeholders. conceptual papers which analyse the impact of governance on management structure and practices and the degree to which ‘good governance’ criteria may influence management and planning for tourism development in pas have been limited (i.e. buteau-duitschaever et al. 2010; eagles 2008; eagles et al. 2010, 2013; hannah 2006; su, wall & eagles 2007; su & xiao 2009; suntikul 2010). this manuscript will not close this conceptual gap, but it will attempt to help guide practice and future research for the governance of pas. some of the relevant questions which are pursued here address the challenges for (good) governance and discuss which ‘good governance’ criteria are critical for pa management, such as: how are local interests balanced with national interests? sub-questions which are pertinent for parks and pas in vietnam generally are: how may the tourism system fit into the pa system in vietnam, what is the degree of independence from government and the impact of decentralised governance on tourism development? to what extent are stakeholder networks and engagement impacted by pa governance issues or, vice versa, how does pa governance impact on stakeholder engagement? are there values which are not obvious that can be revealed to underlie the methods and structures of governance? this article will not be able to cover all these topics in detail, nor does discussion take place on the macro level of policymaking or governance. nonetheless, it may be a first step in advancing knowledge on aggregated issues which are reflected at the ‘nested concepts of government’, that is, at the local level (eagles 2009:245). at the same time, topics addressed above remain under-researched, especially in south-east asia in general and vietnam in particular. based on the case study of the phong nha-ke bang national park (pnkb np) in central vietnam, this article initially describes the relationships which exist between planning, management and governance. subsequently, specific factors relating to the creation of visitor opportunities, and which may exert more or less influence on the governance and planning of the pnkb np, and of pas in general, are discussed. graham, amos and plumptre (2003) outlined five main principles of good governance for a undp list of characteristics for pas. acknowledging that these principles may not be equally applied in different contexts, eagles (2009) examined these criteria in respect of eight common management models for tourism management in pas and therewith provided an overview of the implications that a variety of partnerships have on the way tourism in pas is governed and managed. following his analysis, eagles (2009:245) encouraged discussion on the ‘contentious issues of criterion fulfilment with each model’. the second objective of this article is to examine to what extent proposed ‘good governance’ criteria apply to the pnkb np. pas in vietnam are nestled within complex settings, which include a range of actors and a mix of management forms. protected area governance and governance principles governance and planning are widely, sometimes controversially, used terms, particularly within political environments. planning develops long-term goals, whilst governance is defined as ‘the interactions among structures, processes and traditions that determine how power and responsibilities are exercised, how decisions are taken, and how citizens or other stakeholders have their say’ (graham et al. 2003:ii; cf. also abrams et al. 2003). governance does not follow a fixed progress, but remains a rather loose process which involves a variety of stakeholders (graham et al. 2003). to increase visitation, enhance visitor experiences and, often as a response to their own capability and capacity limitations, many parks connect to other stakeholders and enter into partnerships. whilst partnerships have become forms of governance in many protected areas, the complexity of stakeholders involved in the planning process of pas, as well as the dynamic socio-political and economic environments that pas sit in, challenge tourism management and governance (eagles 2009; gössling 2003).‘good governance’ is defined as a ‘fair’ and effective way of exercising governing powers (means) in order to meet the objectives (ends) of pas (abrams et al. 2003). based on characteristics which have been delineated by the undp, graham et al. (2003) delineate five principles of ‘good governance’ for pas, whilst considering that these principles might not be equally applicable in every context (table 1). these principles are generally accepted as necessary requisites to the success of pas (abrams et al. 2003; graham et al. 2003). eagles (2009) examined a variety of partnership models for the management of tourism services and activities in pas. he illustrates a wide variety of possible partnerships involving governmental institutions only, public and for-profit or non-profit institutions, for-profit or non-profit organisations alone providing public services, a state in which service activities in tourism are sold or leased out to non-profit or for-profit agencies, or involving public entities which might function like a private operator, a statal or provincial entity or a delegated management to some other body, the community or a single individual from the community (abrams et al. 2003; graham et al. 2003). in addition, the different types of management bodies, ownership of land and conservation resources and income of sources take an important impact on pa governance (eagles 2008, 2009). to encourage discussion for their practical application, eagles (2008, 2009) applied the five criteria for good governance (table 1) to the eight most common management models which underpin tourism partnerships in pas on the criteria of management body, income source and land ownership constellation. using a five-point likert scale (from ‘very weak’ to ‘very strong’) based on secondary literature and personal experience, eagles (2009) illustrated the extent to which the single governance criteria apply to the various management models. a precondition for analysis was that equal importance was given to the criteria and that the management models examined were not context-bound. efficiency, public participation, strategic vision and responsiveness were the most common, whilst accountability and transparency were the least common criteria receiving attention. financial effectiveness and equity were the most highly valued. these criteria were more likely strongly linked to management models in which a non-profit organisation rather than a for-profit organisation were involved (eagles 2008, 2009). furthermore, eagles (2009) found that specific models, such as the ‘national park model’ (government ownership of resources, tax funded and management by local government), would involve more partners than other models if implemented (glover & burton 1998; graham et al. 2003; lockwood 2010; more 2005). jamal and stronza (2009:185) examined the ‘complex planning domain’ in a broader pa context which evolves from the variety of actors present in pa management. they found that ‘systems’, namely the park, tourism, ecological and community-resident system, each with its own stakeholders and their interests and values, stand in an interdependent (not symbiotic) nested, but fragmented relationship to each other. an examination of these relationships, which was based on a community-based tourism concept implemented in bolivia, revealed that only collaborative approaches (such as co-management) which consider powers, values and, above all, local knowledge, are likely to succeed in the long term. at the same time, jamal and stronza (2009) illustrated a major gap for tourism planning and management in pas: the use and conservation gap. tourism organisations, planners or businesses largely focus on marketing and promotion rather than on conservation and the sustainable use of resources, demanding for more commitment on ‘process-related factors, such as trust, commitment, open communication, flexibility and the ability to manage conflict’ (moore & weiler 2009:131). table 1: good governance principles for national parks and protected areas management. research method and design top ↑ the pnkb np in vietnam was designated a united nations educational, scientific and cultural organization (unesco) natural world heritage site (whs) in 2003 and represents a typical provincially managed pa, as is the case with many pas in vietnam. organisational ethnography was used to gather data for this study (schwartzman 1993). two authors of this article currently work for the german corporation for international development (giz) project ‘nature conservation and sustainable development of the natural resources in pnkb np region’, which has been active in the park region since 2007. descriptions of governance and planning for tourism development in the pnkb np are based on their daily participation and observation of park management, as well as on their working experience with relevant stakeholders. numerous formal and informal interviews and discussions with local stakeholders were collected over three years of fieldwork. they cover a variety of issues, ranging from public–private partnership and product development to marketing and hospitality service improvement. secondary sources relate to grey literature, as well as to internal studies and reports on, for example, responsible tourism, payments for ecosystem services through tourism, handicraft production, alternative livelihood development and value chain analysis. at the same time, data and knowledge have been integrated from the second author of this article, who is currently undertaking his phd on the pnkb np, examining the influence of governance principles on tourism development.to investigate how international best practice governance principles were applied at the park, the researchers used semi-structured interviews to collect first-hand data. key questions included, (1) whether these good governance principles apply to the park at the moment, (2) to what extent they were applied and (3) if any further underlying governance principles can be uncovered. data were collected in two phases, from july to september 2012 and from february to april 2013. data were analysed using content analysis (yin 2003a, 2003b). results top ↑ governance of protected areas in vietnam vietnam is one of the most bio-diverse nations in the world (undp 2010), with 7.6% of its terrestrial land considered pas. vietnam features around 30 national parks (most of which are legally classified as ‘special-used forests’ [sufs]), 48 nature reserves, 11 species management areas and 39 landscape protection sites (government of vietnam 2012).different ministries govern pas in the country. the ministry of agriculture and rural development (mard) holds management responsibility for terrestrial sufs, marine and wetland pas are under the ministry of fisheries and the ministry of natural resources and environment, the ministry of planning and investment is concerned with financial issues, the vietnam national administration of tourism develops and implements the national tourism strategy and is responsible for promoting many of the pas. currently, mard finances seven out of 128 sufs, others depend on provincial budgets provided either by the respective provincial people’s committee (ppc) or by other line agencies such as the provincial forest protection department. the past two decades have seen drastic changes in tourism development in the country. the political reforms of doi moi1 significantly influenced the way pas are managed and governed. suntikul, butler and airey (2010) examined the socio-economic, cultural and political issues which have been posed for tourism development and its stakeholders in pas with the economic transition processes. they found that: withdrawal of control … has enabled and encouraged change in tourism patterns in national parks. international organizations and private enterprises have taken advantage of unprecedented possibilities opened up by the removal of government restrictions and monopolies. international ngos generally are interested in only some aspects of park development. the decentralization of control has also given individual parks new powers of self-determination, but there is little evidence of vision or innovation from any specific park management authority. currently, no single entity or interest can be said to have a complete vision for, or complete control of, the development of vietnam’s national parks. (suntikul 2010:215; see also larsen 2008) phong nha-ke bang national park the phong nha-ke bang national park is located in the central vietnamese province of quang binh. it covers an area of 123 326 ha, which has been divided into three administrative areas, each operating under a different management regime: a strictly protected area, an ecological restoration area and an administration and services area, with further sub-zones existing for the former two areas. the pnkb np has received worldwide acclaim with the inscription onto the unesco natural world heritage list for its outstanding geological and geomorphological values (unesco 2013). the number of visitors has exponentially risen in the past decade from several tens of thousands to more than 400 000 visitors per year in 2012. the provincially approved sustainable tourism development plan 2010–2020 for the pnkb np region (including the national park and its buffer zone) is, at least officially, the main guiding document for tourism development for provincial and district government as well as for planners and investors (people’s committee of quang binh province 2010). ownership of land resources and types of tourism site management resource ownership of the pnkb np, including the np area and the world heritage property, lies with the national park management board (npmb) and thereby with the national government. the land of the np is managed as suf land and is protected under vietnamese forest law. the npmb, with a provincial status similar to that of a provincial line department, is responsible for (managing) the protection of these resources. the npmb can also make sustainable use of the resources for tourism activities, given that relevant regulations set out by the respective responsible ministries are adhered to. cooperation with line agencies on the implementation of these activities is encouraged. two villages with a combined population of around 78 inhabitants are located within the borders of the np. table 2 outlines the different types of management which apply to the resources that are currently exploited for tourism activities in the pnkb np.in june 2012, a prime ministerial decision (decision 24/qđ-ttg) (prime minister 2012) was released on piloting investment opportunities for private tourism development in vietnam’s nps. the decision proposes the conversion of tourism centres or other ‘ecotourism business units’ into joint stock companies (whereby the npmb will hold at least 51% of capital), if annual revenues exceed 3000 million vnd. rental fees of operated sites and activities within the np are subject to negotiation every 5 years, with rental periods of a maximum of 50 years to apply. at the same time, the decision also encourages lease agreements between suf management boards and tourism businesses, organisations or individuals. leasing periods for up to 50 years can be agreed upon, with the possibility of extensions of up to 20 years. whilst the pnkb np has indicated interest in becoming a pilot site for conversion of the pnkb tourism centre into a joint stock company, so far no serious follow-up has taken place. table 2: examples of management types of tourism activities in the phong nha-ke bang national park. income sources income for the np derives from profits made by the pnkb tourism centre from a single lease contract with a private cave operator (2% of the yearly revenue), as well as from an environmental fee, which needs to be paid by every visitor to touristic sites within the strictly protected zone. both the tourism centre and the administration unit handle, and are responsible for, financing activities carried out by the pnkb np. revenues are transferred to the treasury house of the department of finance. both the np and the pnkb tourism centre have access to this money, according to budget estimations submitted for approval to the treasury house at the beginning of each year. salaries and administration, as well as fixed costs for the park, derive from a separate provincial budget which derives from provincial revenues and societal taxes (table 2). the state also makes smaller budgets available for co-management protection practices of forest resources. eco-touristic development is generally encouraged to ‘create revenue sources to cover incurred costs, generate income for staff and officers as well as become an alternate financial source replacing investments from the state’s budget’ (government of vietnam 2012). management body the pnkb np is under government management. it is under the authority of the ppc of quang binh. a management board (npmb) with one director and three vice directors is in place, appointed by the ppc which directly reports back to the said authority. this board is responsible for the operational management of the np with management procedures documented and approved by the ppc. the np is organised into four units and two functional offices (administration and organisation, planning and finance). the line departments most relevant to tourism development in the np include the department of culture, sports and tourism (docst), the department of agriculture and rural development and the department of planning and investment. the former has responsibilities for monitoring activities relevant to cultural conservation and development, as well as for tourism in both the core and buffer zones of the park. the latter two hold responsibility for directing the investment and implementation of infrastructural plans and activities delivered at the district and commune level. figure 1 provides an overview of the institutional framework in which the park is set.the pnkb tourism centre officially assists the npmb in the management and planning of tourism and in the organisation of tourism activities and services in the np. it has to report directly to the np. the pnkb tourism centre is a ‘quasi-parastatal’ and autonomously functioning unit and considered a profit-making business. whilst the tourism centre has to apply, as do other tourism businesses, for permissions to operate tours within the strictly protected zone of the np, it also owns and operates caves and other tourism sites in the np’s ecological restoration and service and administration areas. figure 1: institutional framework of the phong nha-ke bang national park. management model figure 2 illustrates the ideal management model according to legislation and policy documents. the npmb holds the authority to manage and protect, as well as to develop, the park’s resources. the tourism centre advises on tourism development and is responsible for the management and organisation of tourism activities which have been approved for operations both by the npmb and the ppc. the management can be conducted ‘in house’, or by co-management or concession agreements with third parties.private sector actors that are approved to manage specific tourism sites (e.g. through a concession type or forest lease agreement) report back to the npmb. with the development of new sites and routes, the npmb has to consult the docst for revision of the activity or site development applications. this can be done directly or indirectly via the ppc of quang binh. district and commune authorities support activities by, for example, planning for public security. the asian development bank, the kreditanstalt für wiederaufbau and giz provide financial and technical assistance in the implementation processes. the ppc holds the final decision-making power on the management and development of sites within the np. figure 2: ideal management model in the phong nha-ke bang national park. governance criteria and the public–private for-profit national park model this ideal management model for pnkb np is subject to a mix of different management types as outlined by eagles (2008, 2009), namely the national park, the parastatal and the public and for-profit combination model. resource ownership lies with the government, funding is obtained through both provincial taxes and revenues generated by the tourism centre and a private-run company, and the government-appointed tourism centre as well as the private company manage sites in the np. in the following discussion, this model will be termed the public–private for-profit national park (pppnp) model (see section above entitled ‘protected area governance and governance principles’). the pppnp model demonstrates features which apply to the single forms of management, including the national park, parastatal and public and for-profit combination model (eagles 2008, 2009). the focal points of this ‘ideal’ revolve around equity, participation and direction. however, a number of contextual issues which guide governance in practice underlie these factors.the provision of a strong strategic vision is one of the strengths of this model. although diverging in their nature, both the national park (through the park management plan) (people’s committee of quang binh province 2012a), and the pnkb tourism centre set forth clear visions on the future direction for park protection and tourism development. the park is environmentally driven, whilst the objectives of the tourism centre and of private stakeholders are clearly driven by economic concerns. at the same time, the longevity of strategic visions poses challenges. one interviewee from the npmb states that: ‘our park has announced a new strategy last year. after one year, another strategic vision was decided upon. the connection between stakeholders and projects [e.g. which support the formulation of plans] is not good … when we have a strategic vision, we need to maintain it in ten or twenty years time and seek for expected outcomes. for example, we have just bought new clothes today. suddenly, we find these not to be fashionable anymore the next day and we tend to throw it away. it is waste of resources and public money.’ (npmb employee 1, male, 55 years old) at the same time, one interviewee fears that the imbalance of strategic vision may lead to a ‘harvesting of the fruits while destroying the plant at the same time’ (npmb employee 2, female, 38 years old). nature conservation objectives are increasingly being pushed to the background. vietnam is currently undergoing a transition period and has recently been upgraded to a lower middle-income country (the world bank 2014). this poses an increasing challenge to donors and current investors, as well as publicly involved project implementers in pas. although the socio-economic development plan until 2015 (people’s committee of quang binh province 2008) acknowledges environmental protection, donors are increasingly challenged to retain a conservation focus due to changes in public authority commitment. in contrast, donors have had a great deal of impact on strategic visions, most often with ‘good governance’ intentions. the set principles however may not always apply to provincial or pa conservation visions. in addition to differences in visions, larsen (2008:441) highlights that there is at times ‘institutional [and juridical] confusion with overlapping sites involving different ecosystems or overlapping categories (such as pnkb being both a national park and a whs)’. the current pppnp model only minimally reflects equity concerns, other than that there should be no entrance fees for entering the np. access to services and facilities is restricted in a number of ways. services offered by both the tourism centre and the private sector are bound to the supply–demand market mechanism and profits often far outweigh the coverage of incurred costs. this poses difficulties, particularly to domestic visitors, and specifically to those who arrive from the surroundings of the national park. many locals who have grown up and lived in and around the buffer zone of the park can simply not afford to visit the world heritage site. the pppnp model suggests open revision processes of applications for touristic development sites, as well as for permissions to access specific sites. yet, accountability is undermined by lobbying activities, initiated from both the public sector and private enterprises. the latter seek to avoid, as far as possible, laws and regulations, and wish to create an environment in which public sector ‘interference’ is minimal. these attitudes also lead to conflict amongst private sector stakeholders who are interested in ‘genuine’ collaboration. application procedures, contract selection and monitoring are constrained, to an unknown extent, by a lack of transparency. at the same time, allegations of negligence in drafting a contract between the park (and parties involved in the revision process) and a private operator have been made, involving a 50-year lease agreement in which the operator can deny the monitoring of visitor numbers. the park holds insufficient funds, authority and capacity to monitor tourism activities in the park and has difficulties in communicating relevant information on budgets and operations. an npmb interviewee points out that, despite a ‘knowledge pool’, there are still constraints in human resource capacity in tourism development. however, there are sometimes opportunities to seek help from ngos or other counterparts, though it is often not known how and with whom to seek help from (npmb employee 3, male, 40 years old). participation and benefit-sharing remain problematic outside the ideal pppnp management model. although both the recently introduced operational and management plans (people’s committee of quang binh province 2012b) highlight ecotourism as one of the major incentives for the involvement of local people, communities do not currently benefit from the touristic developments that take place. in 2009, an eco-trail was developed on the basis of a co-management concept which would allow relevant stakeholders, especially local communities, to be substantially integrated into the management of the trail, especially through the employment of locals. after 2 years, it became evident that local staff, when they left, were being increasingly replaced by non-local staff. financial compensations are sometimes demanded of job seekers by staff working in and around the national park to secure a position. there are some downsides to the practical implementation of the management model: there are too many boat operators and, in fact, too many photographers and souvenir vendors for whom tourism serves as an additional income source. earnings are not sufficient to make a living, so that income is topped up with forest resources by families. this is often also the case for those locals who have sold their lands to government or to private (tourism) investors. estimated income figures do not take into account the income from selling land and from labour migration. touristic sites are state-controlled (through the np or the pnkb tourism centre) or leased to private operators. both entities are profit-oriented, and benefit-sharing takes place only on a small scale, for example, through the employment of local porters. participation is difficult around the main touristic sites in the park. there is currently no participation of ethnic minority people living in the core zone of the park and their involvement in future activities can be seen as a form of cultural commodification (larsen 2008). there is also increasing pressure on ethnic minorities’ customary land rights and resources from the growing demand for touristic activity development inside the np. the financial flexibilities of the park, as well as of the tourism centre, have been reduced. revenues of the tourism centre are considered as revenue for the province. staff, administration and other fixed costs are largely covered from the profits, or are funded from provincial budgets. at the beginning of every year the npmb, as well as the tourism centre, are obliged to provide the ppc with an annual working budget. throughout the year, both organisations can then apply for funds to be made available. this leads to restrictions in both responsiveness to urgent events and investments made by the tourism centre in nature conservation. ethical considerations top ↑ application for ethical review for research involving the human participants of this study was approved in the scope of the phd thesis by the hong kong polytechnic university for a period from 25 october 2010 to 25 december 2013, with the reference number hsears20120627002. recruitment procedures in qualitative research, many ethical issues can arise during a data collection period. when the researchers enter a community, it is considered good protocol to seek permission to conduct the research from potential stakeholders or groups within the community. in this study, although the researchers had officially been accepted by the top management of the park, and had received support from the relevant gatekeepers, it was considered ethical not to coerce potential stakeholders to join the study (buchanan, boddy & mccalman 1988). it is the researchers’ obligation to ensure that potential participants do not join the study because of the power relationships between employee and employer, as well as guaranteeing minimisation of harm. informed consent the researchers took the necessary time to establish rapport with all interviewees (buchanan et al. 1988). it was made sure that they received informed consent from each participant personally, which takes into account the right to refuse and withdraw without any report to the gatekeeper. gaining ethical accessibility to all levels of a community is something out of the control of the researchers. fortunately, in this case, the prolonged engagement with the park has shown to work in addressing this issue. during the data collection period, the researcher had to provide enough information about the study to the participants. as the result of the study needs to be written down as a report, the researcher needed to mention this information at the beginning of every interview, in order to inform and obtain allowance from the participants. data protection the researcher needs to ensure anonymity and confidentiality not only of the recorded data, but also of how the researcher discusses information gathered from participants in public areas (hennink, hutter & bailey 2011). during the data collection process in this study, the researcher took various steps to ensure the interview situations remained confidential. most of the interviews were conducted in private rooms or offices. if the setting had to be in an open air (e.g. at the ticket booth or cave entrance), the researcher asked the interviewees to move to a quiet part of the place or another location. all information identifying the respondents was removed from the interview transcripts and/or further quotations. therefore, no individual participants can be identified personally. the interview recordings were stored on a password-protected pc to which only the researcher and the supervisor had access. trustworthiness top ↑ in this study, trustworthiness was established through prolonged engagement, triangulations and member checks (padgett 1998). reliability for the prolonged engagement, the researchers spent approximately 3 years (for the giz fieldwork) and 4 months (for the phd study) in the np or surrounding areas. the aim of the prolonged engagement is to build relationships and rapport with relevant stakeholders within the park. in order to fully intermingle with the stakeholders, the researchers lived in a rented guest house in the park. furthermore, collecting additional data and spending time with the participants also increases the ability for a qualitative researcher to reach data saturation (lietz, langer & furman 2006). validity this study used data triangulation and methodological triangulation to maintain the necessary rigour of case study method (patton 1987). multiple data sources from different methods (in-depth interview, documentary and observation) were used in the data collection and data analysis processes. an important technique in dealing with trustworthiness is member checking (creswell 1998, 2003; horsburgh 2003; johnson & waterfield 2004; lincoln & guba 1985; sandelowski 1993). apart from npmb employees, the researchers invited the giz and academic researchers to join the member checking of the good governance result. the researchers believed that these professional external stakeholders (i.e. staff of the giz, academic researchers) would be able to make valuable contributions during the member checking process (horsburgh 2003; johnson & waterfield 2004). discussion top ↑ this article reviewed ‘good’ governance criteria based on the specific case study of the phong nha-ke bang national park, located in central vietnam. the criteria helped to pinpoint some of the contentious factors which may be embedded in the ‘ideal’ management model (with ‘ideal’ meaning here the officially proposed pa management and structure) which governs the visitor and tourism management of the np. the management model which guides the pnkb np was termed the pppnp model, with government-owned resources, public and private funding sources and co-management practices (between one private tourism business and the np and/or provincial authorities) in place.the discussion of the governance criteria revealed several contextual factors in the practical implementation of this model, which has implications for partnerships and planning, as well as for communication between the public and private tourism sectors in the park. in theory, the ‘ideal’ pppnp model assumes the high value of (financial) efficiency, strategic vision and, to some extent, equity. our research, however, shows that, in practice, the pppnp model is significantly governed by opaque structures and processes and underlain by cultural values which define responsibilities, decision-making and the degree of involvement of a variety of tourism actors. eagles (2009) highlighted financial burden as one of the major, if not as the major constraints, for nps to act. it is interesting to note that, although the np receives considerable income from tourism, there is a bureaucratic burden to access these financial resources, which is problematic for the pnkb np. the fact that finances are centralised also has major impacts on tourism management and planning. despite the de-centralisation processes which has taken place in governance, fragmentation, little information sharing and slow progress have not only been caused by a lack of institutional capacities, but also because top-down processes still apply. top-down processes, with the province holding control over investment applications in tourism, vitally determine tourism planning and management in the park. actors are most often still bound by ‘the formal roles of responsibilities [as well as by] a complex set of power relationships and vested interests often cutting across formal roles and responsibilities’ (larsen 2008:457). interests and relationships are defended to an extent that leads to a lack of control, unofficial communication flows and erratic commitment by any stakeholder (group). burdens are not only created ‘inter-systematically’ (jamal & stronza 2009) (e.g. between different actors and public and/or private partners), but also ‘intra-systematically’ (e.g. amongst public partners or amongst private sector businesses). whilst top-down and decentralised processes have the ability to also guide equity, there is little evidence in the pnkb np which would justify this. local, national and international tour operators reap most of the benefits. tourism development is not only concerned with cave and walking route leases, but new regulations such as decision 24 (prime minister 2012) further facilitate private sector investment. international organisations and private enterprises have taken advantage of unprecedented possibilities opened up by the removal of government restrictions and monopolies; ngos or donors are generally interested in only some aspects of park development. in late 2012, a private tourism stakeholder explained during an informal talk on public–private partnerships in pnkb that ‘national parks are managed by politics, not by nature lovers or tourism experts’ (tour operator 1, male, 42 years old). tourism is inevitably linked to local and national politics. the ‘tourism system’ is not standing in a ‘nested’ symbiotic relationship, neither with the park, the community-resident, nor least with the ecological system (jamal & stronza 2009). but how can the means (responsibilities in exercising powers according to the ideal pppnp model) be better guided to suit the ends (conservation objective) in the pnkb np? a reconciliation of the current model to ensure improvements in benefit-sharing mechanisms, policymaking and enforcement should be considered (truong 2013). conclusion top ↑ this article has suggested further questions on governance issues for tourism and visitor management in vietnam’s pas. first of all, a comparison of different management models prevalent in vietnam should be undertaken. a specific comparison could take place not only amongst provincially governed pas, but also between those provincially and centrally governed. to what extent do decentralised levels (national and provincial) exercise their powers on direction and equity of pa management in general? to what extent have political change and the socio-economic transition period in vietnam influenced governance principles of pas? what relationship is there between different powers of planning, regulations or control over finances? what is the (national and sub-national) government’s relation with its citizens? tourism in vietnam is developing controversially. it is a balancing act of reconciling socio-economic benefits and environmental conservation (including potential pressure on pas to generate more income to fund pa management). concepts such as responsible or eco-tourism development receive increased attention, but very often this attention is limited to promotional activities. pas increasingly suffer from the one-sided direction of tourism development. this article does not deliver a conceptual response, but it may be an initial step to further consider the importance of governance criteria for tourism and visitor management of pas in vietnam. it may furthermore guide future analysis of management models through its application to the ‘real-world’ case study of phong nha-ke bang national park. acknowledgements top ↑ competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions a.h. (giz) was responsible for compiling secondary and primary data and prepared the research article. l.t.p. (hong kong polytechnic university) collected large parts of the primary data and provided important input on drafts. t.s.h.c. (giz) provided important information on the pnkb context as a result of many years of working experience in the area and gave input on drafts. references top ↑ abrams, p., borrini-feyerabend, g., gardner, j. & heylings, p., 2003, evaluating governance: a handbook to accompany a participatory process for a protected area, joint publication of parks canada, ottawa and tilcepa – theme on indigenous and local communities, equity and protected areas of the international union for conservation of nature iucn, commission on environmental, economic and social policy and world commission on protected areas, gland.buchanan, d., boddy, d. & mccalman, j., 1988, ‘getting in, getting on, getting out, and getting back’, in a. bryman (ed.), doing research in organizations, pp. 53–67, taylor & francis, london. buteau-duitschaever, w.c., mccutcheon, b., eagles, p.f.j., havitz, m.e. & glover, t.d., 2010, ‘park visitors’ perceptions of governance: a comparison between ontario and british columbia provincial parks management models’, tourism review 65(4), 31–50. http://dx.doi.org/10.1108/16605371011093854 creswell, j., 1998, qualitative inquiry and research design: choosing among five traditions, sage, london. creswell, j., 2003, research design: qualitative, quantitative, and mixed methods approaches, sage, london. dearden, p., 2000, ‘tourism, national parks and resource conflicts’, in s.w. boyd & r.w. butler (eds.), tourism and national parks: issues and implications, pp. 187–201, john wiley & sons, chichester. eagles, p.f.j., 2008, ‘governance models for parks, recreation and tourism’, in k.s. hanna, d.a. clark & d.s. slocombe (eds.), transforming parks and protected areas: management and governance in a changing world, pp. 39–61, routledge, london. eagles, p.f.j., 2009, ‘governance of recreation and tourism partnerships in parks and protected areas’, journal of sustainable tourism 17(2), 231–248. http://dx.doi.org/10.1080/09669580802495725 eagles, p.f.j., havitz, m., mccutcheon, b., buteau-duitschaever, w. & glover, t., 2010, ‘the perceived implications of an outsourcing model on governance within british columbia provincial parks in canada: a quantitative study’, environmental management 45(6), 1244–1256. http://dx.doi.org/10.1007/s00267-010-9477-3, pmid:20364254 eagles, p.f.j., mccool, s.f. & haynes, c.d., 2002, sustainable tourism in protected areas: guidelines for planning and management, iucn, gland. eagles, p.f.j., romagosa, f., buteau-duitschaever, w.c., havitz, m., glover, t.d. & mccutcheon, b., 2013, ‘good governance in protected areas: an evaluation of stakeholders’ perceptions in british columbia and ontario provincial parks’, journal of sustainable tourism 21(1), 60–79. http://dx.doi.org/10.1080/09669582.2012.671331 glover, t.d. & burton, t., 1998, ‘a model of alternative forms of public leisure services delivery’, in m.f. collins & i. cooper (eds.), leisure management: issues and applications, pp. 139–155, cabi, london. gössling, s., 2003, ‘tourism and development in tropical islands: political ecology perspectives’, in s. gössling (ed.), tourism and development in tropical islands: political ecology perspectives, pp. 1–37, edward elgar publishing, cheltenham. government of vietnam, 2012, implementation of sustainable development, national report at the united nations conference on sustainable development (rio +20), 20–22 june 2012, hanoi, vietnam. graham, j., amos, b. & plumptre, t., 2003, governance principles for protected areas in the 21st century: a discussion paper, viewed 26 july 2013, from http://dspace.cigilibrary.org/jspui/handle/123456789/11190 hannah, l.e., 2006, ‘governance of private protected areas in canada: advancing the public interest?’, phd thesis, department of geography, university of victoria, viewed 26 july 2013, from https://dspace.library.uvic.ca:8443//handle/1828/1886 hennink, m., hutter, i. & bailey, a., 2011, qualitative research methods, sage, london. horsburgh, d., 2003, ‘evaluation of qualitative research’, journal of clinical nursing 12(2), 307–312. jamal, t. & stronza, a., 2009, ‘collaboration theory and tourism practice in protected areas: stakeholders, structuring and sustainability’, journal of sustainable tourism 17(2), 169–189. http://dx.doi.org/10.1080/09669580802495741 johnson, r. & waterfield, j., 2004, ‘making words count: the value of qualitative research’, physiotherapy research international 9(3), 121–131. larsen, p.b., 2008, ‘linking livelihoods and protected area conservation in vietnam: phong nha-ke bang world heritage, local futures?’, in m. galvin & t. haller (eds.), people, protected areas and global change: participatory conservation in latin america, africa, asia and europe, vol. 3, pp. 431–470, geogaphica bernensia, bern. lietz, c., langer, c. & furman, r., 2006, ‘establishing trustworthiness in qualitative research in social work: implications from a study regarding spirituality’, qualitative social work 5(4), 441–458. lincoln, y. & guba, e., 1985, naturalistic inquiry, sage, beverly. lockwood, m., 2010, ‘good governance for terrestrial protected areas: a framework, principles and performance outcomes’, journal of environmental management 91(3), 754–766. http://dx.doi.org/10.1016/j.jenvman.2009.10.005, pmid:20364254 pmid:19896262 moore, s.a. & weiler, b., 2009, ‘tourism-protected area partnerships: stoking the fires of innovation’, journal of sustainable tourism 17(2), 129–132. http://dx.doi.org/10.1080/09669580802582506 more, t., 2005, ‘from public to private: five concepts of park management and their consequences’, the george wright forum 22(2), 12–20. padgett, d., 1998, qualitative methods in social work research: challenges and rewards, sage, thousand oaks. patton, m., 1987, creative evaluation, 2nd edn., sage, newbury park. people’s committee of quang binh province, 2008, an overall report on socio-economic master plan of quang binh province until 2020. people’s committee of quang binh province, 2010, sustainable tourism development plan 2010–2020: phong nha-ke bang national park region, tourism resource consultant, giz, dong hoi. people’s committee of quang binh province, 2012a, strategic management plan 2013–2025 phong nha-ke bang national park, kfw, dong hoi. people’s committee of quang binh province, 2012b, operational management plan 2013–2023 phong nha-ke bang national park, kfw, dong hoi. prime minister, 2012, ‘decision no 24/2012/qd-ttg hanoi on investment policy for development of special-use forests in 2011–2020’, 01 june, government of vietnam, hanoi. sandelowski, m., 1993, ‘rigor or rigor mortis: the problem of rigor in qualitative research revisited’, advanced in nursing science 16(2), 1–8. schwartzman, h.b., 1993, ethnography in organizations, vol. 27, sage, newbury park. semone, p., laws, e., ruhanen, l., wang, z. & scott, n., 2011, ‘governance approaches in new tourist destination countries: introducing tourism law in cambodia, laos and vietnam’, in e. laws, h. richins, j. agrusa & n. scott (eds.), tourist destination governance: practice, theory and issues, pp. 25–38, cabi, wallingford. http://dx.doi.org/10.1079/9781845937942.0025 su, d., wall, g. & eagles, p., 2007, ‘emerging governance approaches for tourism in the protected areas of china’, environmental management 39(6), 749–759. http://dx.doi.org/10.1007/s00267-006-0185-y, pmid:17453276 su, d. & xiao, h., 2009, ‘the governance of nature-based tourism in china: issues and research perspectives’, journal of china tourism research 5(4), 318–338. http://dx.doi.org/10.1080/19388160903382517 suntikul, w., 2010, ‘tourism and political transition in reform-era vietnam’, in r. butler & w. suntikul (eds.), tourism and political change, pp. 133–143, goodfellow publishing, oxford. suntikul, w., butler, r. & airey, d., 2010, ‘implications of political change on national park operations: doi moi and tourism to vietnam’s national parks’, journal of ecotourism 9(3), 201–218. http://dx.doi.org/10.1080/14724040903144360 tang, c.-h. & jang, s., 2009, ‘the tourism–economy causality in the united states: a sub-industry level examination’, tourism management 30(4), 553–558. http://dx.doi.org/10.1016/j.tourman.2008.09.009 the world bank, 2014, world development indicators vietnam, viewed 05 january 2014, from http://data.worldbank.org/country/vietnam#cp_wdi tosun, c., 2001, ‘challenges of sustainable tourism development in the developing world: the case of turkey’, tourism management 22(3), 289–303. http://dx.doi.org/10.1016/s0261-5177(00)00060-1 truong, v.d., 2013, ‘tourism policy development in vietnam: a pro-poor perspective’, journal of policy research in tourism, leisure and events 5(1), 28–45. http://dx.doi.org/10.1080/19407963.2012.760224 unesco, 2013, phong nha-ke bang national park – unesco world heritage centre, viewed 27 july 2013, from http://whc.unesco.org/en/list/951 united nations development programme, 2010, removing barriers hindering protected area management effectiveness in vietnam, undp, hanoi, viewed 29 july 2013, from http://www.undp.org/content/dam/vietnam/docs/project documents/31386_protected_area_prodoc110124x.pdf yin, r.k., 2003a, applications of case study research, 2nd edn., sage, thousand oaks. yin, r.k., 2003b, case study research: design and methods, 3rd edn., sage, thousand oaks. footnote top ↑ 1. doi moi [renovation] was a period of reforms beginning in 1986 which introduced ‘open-door’ policies that encouraged a shift from a socialist-oriented to a market-oriented economy. article information authors: clifton s. meek1,2 david m. richardson2 ladislav mucina2,3 affiliations: 1percy fitzpatrick institute, dst/nrf centre of excellence, university of cape town, south africa2centre for invasion biology, department of botany and zoology, stellenbosch university, south africa 3department of environment and agriculture, curtin university, australia correspondence to: david richardson postal address: private bag x1, matieland 7602, south africa dates: received: 20 june 2012 accepted: 25 oct. 2012 published: 15 mar. 2013 how to cite this article: meek, c.s., richardson, d.m. & mucina, l., 2013, ‘plant communities along the eerste river, western cape, south africa: community descriptions and implications for restoration’, koedoe 55(1), art. #1099, 14 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1099 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/ koedoe.v55i1.1099-1 copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. plant communities along the eerste river, western cape, south africa: community descriptions and implications for restoration in this checklist... open access • abstract • introduction • research method and design    • study area    • field sampling    • vegetation analysis and presentation • results    • plant communities       • cunonia capensis–brachylaena neriifolia community       • brabejum stellatifolium–metrosideros angustifolia community       • sporobolus africanus–stoebe plumosa community       • quercus robur–cinnamomum camphora community       • cliffortia odorata–ageratina adenophora community       • populus x canescens–tropaeolum majus community       • bromus diandrus–erodium moschatum community       • arundo donax–tradescantia fluminensis community       • phragmites australis–persicaria lapathifolia community       • salix mucronata subsp. mucronata–cyperus textilis community    • ordination • discussion • conclusion • acknowledgments    • competing interests    • authors' contributions • references abstract top ↑ riparian plant communities fulfil many functions, including the provision of corridors linking protected areas and other zones of high conservation value. these habitats across much of south africa’s cape floristic region, especially in the lowlands, have been heavily impacted and degraded by human activities. there is increasing interest in the restoration of degraded riparian zones and the ecosystem services they provide to enhance the conservation value of landscapes. previous studies of riparian vegetation in the cape floristic region focused on pristine headwater systems, and little is known about human-impacted communities that make up most of the riparian vegetation in downstream areas. more information is needed on the composition of these plant communities to establish a baseline for management intervention. the riparian zone of the eerste river in south africa’s western cape province provides a good opportunity to study the features of riparian vegetation along the entire gradient, from pristine vegetation in a protected area through different levels of human-mediated degradation. riparian vegetation was surveyed in 150 plots along the entire length of the eerste river (ca. 40 km). data were analysed using the vegetation classification and analysis software package juice. final groupings were plotted onto a two-dimensional detrended correspondence analysis plane to check the position of the communities in the reduced multidimensional space. ten distinct plant communities were identified, including several novel communities dominated by alien plant species. descriptions of each plant community are presented. diagnostic, constant and dominant species are listed and the major structural and ecological characteristics of each community are described.conservation implications: major changes to hydrological and soil properties, nutrient dynamics and disturbance regimes and plant species composition along sections of the riparian zone mean that restoration of many of these habitats to their historic condition is not feasible. however, several native plant species that provide key ecosystem services persist in and adjacent to transformed communities, offering substantial opportunities for restoration to achieve certain goals. introduction top ↑ the flora of south africa’s cape floristic region (cfr) is exceptionally diverse, with one of the highest levels of species diversity endemism of any flora in the world. research on vegetation ecology in the fynbos biome of the cfr has focused largely on aspects of the iconic cape shrublands – mainly fynbos and renosterveld (e.g. cowling 1992; cowling, richardson & mustart 1997; linder 2003). riparian vegetation in the fynbos biome is, however, structurally and compositionally distinct from the surrounding cape shrublands. the predominant vegetation type of riparian zones in the western cape has been variously named closed scrub fynbos (campbell 1986; cowling & holmes 1992; cowling et al. 1997), hygrophilous mountain fynbos (taylor 1978) and broad sclerophyllous closed scrub (holmes et al. 2005; kruger 1978). other types, ranging from tall herblands to forests, may also occur in the riparian zone (kruger 1978). true afrotemperate forest may develop in areas of steep topography that afford protection from fires (manders & richardson 1992). despite being recognised as a special vegetation type in the biome, riparian vegetation has been paid very little attention in the regional literature. where it is discussed, it is usually in the context of the larger terrestrial matrix. very few formal classifications of riparian vegetation in the fynbos biome have been published (but see prins, holmes & richardson 2004; sieben, mucina & boucher 2009; sieben & reinecke 2008) and none have looked at riparian communities outside relatively pristine headwater systems, which occur mostly in protected areas.riparian communities in the cfr, especially those in the south-western part of the region, have been heavily impacted recently by human activities. intensive land use in the lowlands has led to increased soil erosion and sedimentation, increased nutrient inputs, and altered hydrological regimes due to water impoundment and abstraction. in many cases, vegetation clearance for agriculture and urbanisation has included riparian vegetation. in addition, riverine corridors provide ideal conditions for the establishment, proliferation and spread of the many alien plants that were introduced to the region. the constant disturbance along rivers as a result of their dynamic hydrological nature and their ability to function as conduits for dispersal of propagules means that riparian zones are severely invaded worldwide (foxcroft, rouget & richardson 2007; hood & naiman 2000; planty-tabacchi et al. 1996). positive feedback mechanisms promote rapid spread of aliens along riparian corridors through human-induced habitat alteration and increased propagule pressure (richardson et al. 2007). this combination of pressures on the natural environment has led to the establishment of novel ecosystems (sensu hobbs et al. 2006) along the riparian corridors of the cfr. no corridors remain undisturbed in their lower reaches and it is unclear what the composition of historical plant communities in these downstream systems would have been (brown 1998; prins et al. 2004). these riparian ecosystems fulfil a range of useful functions in the western cape and there is increasing interest in conserving or restoring key ecosystem services in these zones. intact riparian corridors in the cfr have also been identified as being important for migration and exchange of inland and coastal biotas in a human-dominated landscape in the face of global change (rouget et al. 2003). before sensible restoration efforts can be undertaken, more information is required on the composition of these ecosystems. the eerste river provides a superb natural laboratory for the study of the changing dynamics of a river and its associated riparian habitats. this study aimed to describe the plant communities along the entire length of the riparian corridor of the eerste river in the western cape province to establish a baseline for management and future research. the 150 permanent plots established in this study (see also meek, richardson & mucina 2010) form a permanent transect with considerable potential for other studies of biotic and abiotic changes along a well-studied river situated close to a university. another objective of this work was to identify native species that could be used in restoration efforts in disturbed riparian areas in the fynbos biome. research method and design top ↑ study area the eerste river originates in the jonkershoek mountains in the western cape, south africa. it has a catchment area of 420 km2, and it is approximately 40 km long. the source of the river lies 60 km east of cape town at an altitude of 530 m, from where it flows in a north-westerly direction towards the town of stellenbosch. at stellenbosch, the river takes an abrupt turn, flowing southwards until it meets the atlantic ocean in false bay near the town of macassar (figure 1). the river is influenced by flows from several tributaries along its route, with the bank-to-bank width averaging 5 m near the headwaters and 14 m in the lower river zone. the average gradient of the river ranges from 24 m/km in the mountain stream zone to as little as 2 m/km in the lower reaches of the river (king 1981). figure 1: the location of the eerste river in the western cape, south africa. the geology of the eerste river catchment is composed of table mountain sandstone in the upper reaches and malmesbury shale and cape granite of the bokkeveld group in the lower reaches. low-lying coastal plain areas are composed of aeolian sands (brown & dallas 1995). the catchment of the river lies entirely within a mediterranean-like climate zone, which has cool, wet winters and hot, dry summers. annual rainfall averages range from more than 3000 mm (sieben et al. 2009) in the upper reaches of the river to around 570 mm in the lower reaches. about 80% of this precipitation falls in winter downpours, with only 7% of the annual rain occurring between december and march (king 1981).as with most rivers in areas of concentrated development, the eerste river has undergone dramatic changes in the years since european settlement. intensive human use of the river dates back to as early as 1697, with over-abstraction becoming a problem as early as 1862. indigenous trees were used for firewood and timber in the early days of european settlement and were later replaced by plantings of english oaks (quercus robur) (brown & magoba 2009). currently, the landscape surrounding the eerste river supports a number of land uses, including nature conservation, commercial forestry, residential use, various forms of agriculture (vineyards, orchards and crop production) and communal grazing. approximately the first 6 km of the upper river reaches are bordered by natural vegetation within the hottentots holland and jonkershoek nature reserves, and is relatively unaffected by human influence. although forestry plantations of pinus radiata exist in portions of the landscape in the jonkershoek nature reserve, ‘a wide buffer zone of natural fynbos vegetation has been maintained between the river and plantation areas’ (meek et al. 2010). since 1981, the upper reach of the river has been regulated by the kleinplaas dam, above which the eerste river is perennial (department of water affairs and forestry [dwaf] 2004). during summer months, a municipal weir above the dam diverts the river’s flow to the ida’s valley dam, which supplies drinking water and water for domestic use to the town of stellenbosch (brown & dallas 1995). in order to provide compensation releases for downstream users, this flow is replaced with water from the theewaterskloof dam through an interbasin transfer scheme. water from the theewaterskloof dam contains high sediment loads, significantly reducing the water quality and negatively affecting downstream biotas in the eerste river (brown & magoba 2009). ‘stellenbosch (human population ca. 117,000) is the main urban area along the river, with additional urban development present in macassar’ (meek et al. 2010). historically, the river through stellenbosch was notably shallower and more flood prone than it is today, containing heavily wooded banks and numerous islands. considerable canalisation of the river through the town of stellenbosch has all but eliminated the river’s historic flood regime through this area (brown & magoba 2009). the banks of the river through stellenbosch are currently both deep and heavily confined. just south of stellenbosch, the eerste river is joined by the plankenburg river. water is abstracted at this point, near the confluence with the blouklip river, and (at lower levels by by owners of riparian properties) along most of its length. treated municipal effluent enters the river through the veldwagters tributary. below the confluence of the veldwagters and eerste rivers, treated effluent is an important component of river flow in summer. this makes the river largely perennial in downstream areas. field sampling vegetation of the riparian zone was surveyed along the entire length of the eerste river, from the headwaters to just upstream of the estuary. all fieldwork was performed from mid-september through mid-december (i.e. in spring) 2008 in order to capture the greatest number of herbaceous species. vegetation plots were paired at intervals of 500 m along both banks of the river and plot edges were extended to bankfull margins. a total of 150 plots were surveyed and locations were recorded using a garmin gps map76 handheld global positioning system. plot sizes were adjusted depending upon structure of the plant community at each plot location in order to capture all regularly occurring species. herb-dominated communities were sampled with plots of 20 m2, shrublands/thicket with 50 m2 plots, and forest communities with 100 m2 plots. the decision on the plot size was based upon values suggested in westhoff and van der maarel (1973).all plant species within a plot were recorded, and height and absolute cover for each species was estimated. species were determined to be native or alien following the criteria of pyšek et al. (2004) and using published floras including goldblatt and manning (2000), henderson (2001) and bromilow (2005). species that could not be identified in the field were collected and labelled, and identifications were later made at the bolus herbarium, university of cape town. all species determinations were verified by the herbarium curator (meek et al. 2010), and collections have been archived at the centre for invasion biology at stellenbosch university (http://academic.sun.ac.za/cib/). the nomenclature of the plant names follows goldblatt and manning (2000), except for searsia (see moffett 2007). surface soil samples collected from each plot were air dried and the ph was measured using a calibrated ph meter in 10-g samples of soil suspended in 25 ml of potassium chloride solution. soil cation concentrations of sodium, calcium, magnesium, and potassium were measured using ammonium acetate extraction, and phosphorous was measured using the bray ii method for samples with a ph ≤ 6.9 (bray & kurtz 1945) and the olsen method (olsen et al. 1954) for samples with a ph > 7. total carbon content (walkley–black method; walkley 1935) and electrical resistance were also measured. vegetation analysis and presentation vegetation data comprising 150 sample plots were classified by beta-flexible clustering (β = –0.25), using square root transformation and manhattan distances as the resemblance measure to calculate distance between plots. the result of the clustering was translated into a structured relevé table using the software package juice version 7.0 (tichý, chytrý & zelený 2009). vegetation data recorded per vertical layer were used to assure that both floristic and structural components of vegetation composition would be considered by the classification. fidelity values were calculated for each of the resulting clusters, and the species in each of the columns of the synoptic table constructed by juice were sorted by fidelity using the phi coefficient of association (cut level of 0.35; l. tichý, pers. comm., 16 mar. 2009). the structured synoptic table was then translated back into a relevé table and inspected for plausibility (interpretability) of the resulting clusters. clusters were further split or combined until desired resemblance and homogeneity criteria were reached. interpretability of clusters with respect to field knowledge of the plant communities factored heavily in the decision on the final make-up of clusters. the final 10 clusters were plotted on a two-dimensional ordination plot of the detrended correspondence analysis (dca; hill & gauch 1980) using the ecological software package canoco 4.5 (ter braak & šmilauer 2002) to visualise the level of ‘spatial’ separation (relative to the ordination plane) of the classification.descriptions of each plant community were compiled. these list diagnostic, constant and dominant species (alien species are marked using an asterisk), and provide a narrative describing the major structural and ecological characteristics of the community. diagnostic species were determined using the phi coefficient of association between species and vegetation units as a measure of fidelity (chytrý et al. 2002). species were considered to be diagnostic if they had a phi coefficient of 0.35 or higher. constant taxa are those that occurred in at least 35% of relevés for a given community, whilst dominant species are those that had a cover value of 45% or greater in at least 5% of the relevés. a species binomial system was used in naming the communities, with one dominant and one diagnostic species being selected for the names. results top ↑ plant communities ten distinct plant communities were identified. the floristic and structural composition of the communities is represented in the structured relevé table (table 1). a list of all species encountered in the vegetation plots appears as online appendix 1. table 1a: structured relevé table containing taxa grouped according to their fidelity to particular communities. table 1b: structured relevé table containing taxa grouped according to their fidelity to particular communities. cunonia capensis–brachylaena neriifolia community abbreviated name: cunonia community (figure 2a). figure 2: plant communities along the upper reaches of the eerste river (e.g. primarily upstream of the plankenburg river confluence): (a) cunonia community showing tall canopy of metrosideros angustifolia with brachylaena neriifolia and prionium serratum in the undergrowth; (b) brabejum community showing shrubby brabejum stellatifolium and metrosideros angustifolia with prionium serratum growing along the river bank; (c) sporobolus community with sporobolus africana in the foreground and cliffortia strobilifera in the background next to the kleinplaas dam; (d) quercus community. brabejum stellatifolium mixes with pittosporum undulatum and eucalyptus camaldulensis in the shrub layer with a canopy of quercus robur and ilex mitis and (e) cliffortia community showing dense growth of cliffortia odorata with freylinia lanceolata above. diagnostic species: cunonia capensis, brachylaena neriifolia, ilex mitis, schoenoxiphium lanceum, todea barbara, blechnum capense, cassine schinoides, maytenus acuminata, aristea capitata, rapanea melanophloeos, metrosideros angustifolia, podalyria calyptrata, blechnum punctulatum, brabejum stellatifolium, erica caffra, ischyrolepis subverticillata, pteridium aquilinum, pellaea calomelanos, elegia capensis, cliffortia cuneata, diospyros glabra, asparagus scandens, cassytha ciliolate.constant taxa: m. angustifolia, i. mitis. dominant species: c. capensis, r. melanophloeos, p. aquilinum, b. stellatifolium. the cunonia community occurs at altitudes of 262 m – 496 m, primarily where there has been no disturbance on lands immediately adjacent to the riparian zone. rock cover estimates range from 10% – 80%, with an average of 42%. very little soil is present amidst the rock and tree roots. where soils are present, they are sand or gravel with a low ph (average = 4.5), low nutrient content, and high resistivity (i.e. low electrical conductivity). this community is characterised by a tall tree stratum (6 m – 13 m) with an average cover of 72%. this stratum is dominated by c. capensis, i. mitis, m. angustifolia and b. stellatifolium. less common tree species in this stratum include c. schinoides, platylophus trifoliatus, maytenus oleoides and r. melanophloeos. a tall shrub stratum (3 m – 6 m) grows below this, and has an average cover of 61%. it is dominated by b. neriifolia and b. stellatifolium, although m. acuminata and m. angustifolia are also common in this stratum. this combination of tall tree and shrub strata creates a fully closed canopy above the forest floor. a herb stratum (< 1.5 m), with an average cover of 29% and dominated by ferns, is the typical understorey of this community. common understorey species include b. capense, b. punctulatum, t. barbara and s. lanceum. structurally and floristically, this community is similar to the r. melanophloeos – c. capensis community described by mcdonald (1988). it should be noted that this community was completely devoid of alien vegetation, with the exception of a single acacia melanoxylon sapling present in one relevé. this relevé is adjacent to a bridge used by forestry vehicles, suggesting that the a. melanoxylon seed may have been introduced into the habitat by a passing vehicle. brabejum stellatifolium–metrosideros angustifolia community abbreviated name: brabejum community (figure 2b).diagnostic species: m. angustifolia, b. stellatifolium, virgilia oroboides, i. mitis, p. aquilinum, freylinia lanceolata, e. caffra, halleria elliptica, rubus pinnatus, psoralea aphylla, phylica pubescens, pentameris thuarii, p. calyptrate. constant taxa: searsia angustifolia, olea europaea subsp. africana, kiggelaria africana, b. neriifolia, prionium serratum, pittosporum undulatum, d. glabra. dominant species: b. stellatifolium, f. lanceolata, p. serratum. the brabejum community can be found at altitudes from 228 m to 400 m, on sandy soils with low ph (average = 4.3), low resistivity and low nutrient content. this community tends to be interspersed with the cunonia community where it occurs at altitudes between 228 m and 262 m, above which the cunonia community dominates. the amount of exposed rock ranges from 2% to 90%, with an average of 35%. the vegetation is composed primarily of an almost impenetrable layer of shrubs (up to 5 m), with an average cover of 96%. this stratum is dominated by b. stellatifolium, which occurs in every relevé. interspersed with the b. stellatifolium in this layer are m. angustifolia, f. lanceolata, i. mitis, e. caffra and p. calyptrata, amongst others. p. serratum often grows along the water’s edge. above this stratum there is sometimes an emergent tree layer (up to14 m) with an average cover of 11%. the dominant species in this stratum is v. oroboides. a herb stratum dominated by p. aquilinum provides the understorey in this community, with an average cover of 15%. whilst this community is composed primarily of native species, a number of alien species have managed to invade, including p. undulatum and acacia saligna. structurally and floristically, this community has many similarities to the k. africana – b. stellatifolium community described by prins et al. (2004), the h. elliptica – b. stellatifolium community described by mcdonald (1988) and the restio gaudichaudianus – m. angustifolia community described by van wilgen and kruger (1985). sporobolus africanus–stoebe plumosa community abbreviated name: sporobolus community (figure 2c).diagnostic species: stoebe plumosa, r. pinnatus, sporobolus africanus, juncus lomatophyllus, paspalum urvillei*, passerina corymbosa, searsia tomentosa, cliffortia strobilifera, a. saligna*, juncus effuses. constant taxa: s. angustifolia, o. europaea subsp. africana, d. glabra. dominant species: s. africanus. the sporobolus community occurs along the banks of the kleinplaas dam at an altitude of 277 m. it is represented here by only two relevés. this community is dominated by s. africanus, which forms its own stratum, 1.0 m high and with an average cover of 53%. s. angustifolia and c. strobilifera emerge from this stratum, reaching 2 m in height, whilst s. plumosa and r. pinnatus form a low stratum up to 0.5 m. less prominent species found within this community include p. urvillei, j. lomatophyllus and d. glabra, amongst others. whilst p. serratum grows in dense patches along some areas of the kleinplaas dam, it was not captured in either of these relevés. quercus robur–cinnamomum camphora community abbreviated name: quercus community (figure 2d).diagnostic species: cinnamomum camphora*, ligustrum ovalifolium*, acacia longifolia*, syzygium australe*, acacia elata*, q. robur*, p. undulatum*, quercus palustris*, agapanthus africanus (native, but here not in original habitat). constant taxa: b. stellatifolium, commelina benghalensis*, acacia mearnsii*, tradescantia fluminensis*, o. europaea subsp. africana, s. angustifolia, k. africana, p. serratum, m. angustifolia, bromus diandrus*, briza maxima*, brachiaria deflexa, ageratina adenophora*. dominant species: q. robur*, p. serratum, b. stellatifolium. the quercus community is the most prominent riparian plant community of urban areas in and around stellenbosch, occurring at altitudes between 81 m and 225 m. soils are sandy and mildly acidic (average ph = 5.08), with low resistivity and moderate nutrient content. exposed rock ranges from 0% to 80%, with an average of 18%. this is the most diverse of the eerste river plant communities, thanks in part to the extraordinary number of alien species that have been introduced into the area. however, this community also maintains high native-species diversity relative to the other alien-dominated communities. absolute cover of alien species averages 123%, whilst the absolute cover of native species averages 73%. dominating the tree stratum (5 m – 21 m) in this community is q. robur. the alien species c. camphora, eucalyptus camaldulensis and a. mearnsii play a secondary role in the canopy, as do the native species o. europaea subsp. africana, i. mitis and m. angustifolia. average cover in this stratum is 67%. below the tree stratum is a diverse shrub stratum (1.5 m – 5 m) dominated by the natives p. serratum and b. stellatifolium, with an average cover of 68%. other native species that commonly occur in this stratum are o. europaea subsp. africana, s. angustifolia, and k. africana. alien species common to this stratum include c. camphora, l. ovalifolium, a. mearnsii, a. longifolia and p. undulatum. dominating the understorey are alien t. fluminensis and pennisetum clandestinum. other common understorey alien species include b. maxima, c. benghalensis and a. adenophora. this stratum is sparser than the shrub and tree strata, with an average cover of 49%. cliffortia odorata–ageratina adenophora community abbreviated name: cliffortia community (figure 2e).diagnostic species: cliffortia odorata, sesbania punicea*, a. adenophora*, cynodon dactylon, paraserianthes lophantha*. constant taxa: p. clandestinum*, c. benghalensis*, f. lanceolata, b. diandrus*. dominant species: c. odorata. a dense growth of c. odorata (up to 1.5 m) is the defining component of this community, which occurs at altitudes from 114 m to 225 m on sandy soils with moderate resistivity, low nutrient content and an average ph of 5.2. often interspersed with c. odorata are the alien species a. adenophora, c. benghalensis, p. clandestinum and in several relevés, small saplings of s. punicea. emergent through this ground cover are shrubs (up to 4 m) such as f. lanceolata and p. lophantha, providing a cover ranging from 0% to 74%, with an average of 35%. this community generally lacks a tree stratum, although in two relevés it contained a relatively open canopy of alien trees, including e. camaldulensis, q. robur and a. mearnsii. this community is possibly identical to the c. odorata shrublands described by sieben (2003). populus x canescens–tropaeolum majus community abbreviated name: populus community (figure 3a). figure 3: plant communities along the lower reaches of the eerste river (e.g. primarily downstream of the plankenburg river confluence): (a) populus community. tall populus x canescens towers above a herbaceous understorey of tropaeolum majus and vinca major, with olea europea subsp. africana and kiggelaria africana in the middle shrub layer; (b) bromus community with tropaeolum majus and bromus diandrus in the foreground; (c) arundo community showing dense thicket of arundo donax with tropaeolum majus in the understory and anredera cordifolia climbing the arundo stalks; (d) phragmites community showing dense thicket of phragmites australis, with salix mucronata subsp. mucronata and pennisetum clandestinum present on the left and (e) salix community showing bushy patches of salix mucronata subsp. mucronata. diagnostic species: populus x canescens*, tropaeolum majus*.constant taxa: t. fluminensis*, galium spurium, p. clandestinum*, b. deflexa, zantedeschia aethiopica, sonchus oleraceus*, q. robur*. dominant species: t. fluminensis*, t. majus*, p. x canescens*, q. robur*, o. europaea subsp. africana, e. camaldulensis*, c. benghalensis*. of all the communities described here, the populus community is by far the most widespread along the eerste river, accounting for close to a third of all relevés. it is found primarily adjacent to agricultural lands at altitudes between 6 m and 162 m. soils are sandy and mildly acidic (average ph = 5.48), with low resistivity and high nutrient content. this community has the highest absolute cover of alien species of all the eerste river plant communities, whilst the average absolute cover of native species is only 27%. dominating the tree stratum (5 m – 22 m) in this community is p. x canescens, although e. camaldulensis or q. robur replace p. x canescens as the dominant species in some relevés. other prominent tree species in this community include a. mearnsii and salix babylonica. the only native tree species that seem to have maintained a foothold in some areas are o. europaea subsp. africana and, to a lesser extent, k. africana and salix mucronata subsp. mucronata. the tree canopy is almost never closed in this community (63% cover on average), allowing for enough light to promote growth of a herb stratum (< 1 m) with an average cover of 97%. dominating this understorey are t. fluminensis and t. majus, both of which form dense mats of vegetation along the forest floor, which makes it difficult for competing plants to germinate. other common understorey species include b. deflexa, c. benghalensis and p. clandestinum, as well as the native species g. spurium and z. aethiopica. the shrub stratum (1 m – 4 m) is generally sparse (average cover of 22%) and is dominated by young p. x canescens. bromus diandrus–erodium moschatum community abbreviated name: bromus community (figure 3b).diagnostic species: erodium moschatum*, ehrharta longiflora, b. diandrus*, fumaria muralis*, medicago polymorpha*, hordeum murinum*, geranium molle*. constant taxa: p. clandestinum*, g. spurium, t. majus*, z. aethiopica, echium plantagineum*. dominant species: b. diandrus*, p. clandestinum*, t. majus*, rapistrum rugosum*. the bromus alien grassland community occurs at altitudes from 6 m to 114 m in heavily disturbed areas along the river, primarily downstream of stellenbosch and adjacent to agricultural or communal grazing lands. it generally does not occur as a continuous unit, but rather in isolated patches interspersed amongst other communities. soils are sandy, with low resistivity, neutral ph (average = 6.11), and high nutrient content. this plant community is dominated by non-native grasses and herbs and is unique amongst the riparian communities represented here for its lack of a shrub or tree stratum. common species found within this community include the annual grasses b. diandrus, e. longiflora and h. murinum, as well as the perennial grass p. clandestinum. herbs common to this community include the aliens e. moschatum, t. majus and e. plantagineum, as well as the native species g. spurium and z. aethiopica. dwarf versions of many of these species can be found in heavily grazed areas along the river, where a majority of the species have been pruned to within a few centimetres off the ground. p. clandestinum does particularly well in these areas owing to its creeping, mat-forming nature. very little bare ground is present within this community, which has an average cover of 96%. arundo donax–tradescantia fluminensis community abbreviated name: arundo community (figure 3c).diagnostic species: arundo donax*, t. fluminensis*, rubus fruticosus*, q. robur*, anredera cordifolia*. constant taxa: t. majus*, vinca major*, g. spurium. dominant species: a. donax*, t. fluminensis*, q. robur*. the arundo community occurs within an altitudinal range of 15 m – 53 m, adjacent to agricultural lands downstream of stellenbosch. soils are sandy, with low resistivity, low ph (average = 4.45) and unusually high phosphorus and potassium content. species diversity is particularly low, with a mean of nine species per 50 m2 quadrat. as with the bromus community, this community occurs primarily in isolated patches distributed amongst other downstream communities. the defining component of this community is the dense stands of the alien species a. donax (up to 6 m), within which very few, if any, native species are able to survive. the thicket of a. donax emerges above a herbaceous stratum (up to 0.6 m) of other alien species, including v. major, t. majus and t. fluminensis, as well as the weedy native species g. spurium. the tree k. africana was the only other native species found growing within this community. the alien herbaceous vine a. cordifolia is often found twining up the a. donax stalks, whilst q. robur creates a canopy up to 13 m tall above the a. donax thicket. the tree stratum has a relatively low cover (average 26%) compared to the herb (44%) and shrub strata (95%). phragmites australis–persicaria lapathifolia community abbreviated name: phragmites community (figure 3d).diagnostic species: phragmites australis, persicaria lapathifolia*, rumex crispus*, helminthotheca echioides*, r. rugosum*, tetragonia fruticosa, lolium perenne*, sonchus asper*, lycium ferocissimum, malva arborea*, raphanus raphanistrum*, c. strobilifera. constant taxa: z. aethiopica, s. mucronata subsp. mucronata, p. clandestinum*, g. spurium, f. muralis*, c. benghalensis*, xanthium strumarium*, t. majus*, s. oleraceus*, e. plantagineum*, c. dactylon. dominant species: p. australis, p. lapathifolia*. the phragmites community occurs within an altitudinal range between 5 m and 10 m, primarily adjacent to agricultural and grazing lands in the lower reaches of the river, as well as some untransformed lands close to the river estuary. soils are sandy with low resistivity, neutral ph (average = 6.80), and high nutrient content. similar to the arundo community, the most prominent feature of this community is the dense thickets (3 m – 5 m tall) formed by the dominant species, in this case p. australis. this community differs from the arundo community, however, in that it contains a greater cover percentage of native species and is found further downstream. the tall herb p. lapathifolia (up to 1 m) is often interspersed with p. australis, particularly in relevés further from the estuary. in some areas, portions of the p. australis thicket have been grazed down and p. clandestinum has taken hold, along with other non-native herbaceous species such as f. muralis, c. benghalensis and x. strumarium. this community often abuts the salix community (see later) and, as such, s. mucronata subsp. mucronata occasionally overhangs the p. australis, providing minimal tree cover. salix mucronata subsp. mucronata–cyperus textilis community abbreviated name: salix community (figure 3e).diagnostic species: s. mucronata subsp. mucronata, z. aethiopica, cynanchum obtusifolium, cyperus textilis. constant taxa: t. fluminensis*, r. crispus*, r. rugosum*, p. clandestinum*, c. benghalensis*. dominant species: s. mucronata subsp. mucronata, c. textilis, cyperus dives, t. fluminensis*, t. majus*. as mentioned earlier, this community often abuts the phragmites community and is found in similar areas along the lower reaches of the river, at altitudes from 5 m to 15 m. soils are similarly sandy, with low resistivity, neutral ph (average = 6.83), and high nutrient content. native species dominate in this community, with an average absolute cover of 95% as compared with an average of 65% for alien species. the native s. mucronata subsp. mucronata forms a moderately tall (6 m – 11 m) canopy above an otherwise heavily herbaceous community. prominent understorey herbs include the alien t. fluminensis and native z. aethiopica, with native c. obtusifolium occasionally twining through the branches of the s. mucronata subsp. mucronata. in some areas, s. mucronata subsp. mucronata tends to grow in dense, shrub-like patches. thick clumps of c. textilis often intermingle with shrubby s. mucronata subsp. mucronata, creating a tangle of impenetrable vegetation. ordination the results of the dca ordination are graphically illustrated in figure 4. the plant communities show very clear ‘spatial’ separation (with reference to the reduced multivariate space) along axes 1, 2 and 3, suggesting that the floristic-based classification was robust. figure 4: detrended correspondence analysis ordination of the riparian vegetation plots showing positions of the 10 plant communities along axes (a) 1 and 2 and (b) 1 and 3. discussion top ↑ this study provides the first description of riparian plant communities along the entire length of a river system in the cfr, including the many novel communities that now occur in the human-impacted riparian areas in this region. a study of this nature poses a particular challenge owing to the continually changing nature of many of these plant communities. this is especially true in urban and agricultural stretches of the river, which continue to be transformed by the proliferation of already present alien plant species and the incursion of new alien species.the distribution of plant communities along the eerste river is not determined by any single factor, but by the interaction of several environmental factors. it is difficult to separate the effects of environmental factors in a system that has been, and continues to be, so heavily transformed by human intervention. no single environmental component can be treated in isolation, although it appears that land use has played an important role in shaping the composition of plant communities. those riparian areas where the landscape surrounding the river has been most heavily transformed by human factors are home to many of the novel plant communities dominated by alien species that are described in this paper, including the bromus, populus, arundo, and quercus communities. the relationship between alien plant species and surrounding land use was addressed in a separate study (meek et al. 2010). owing to the extensive changes in soil nutrients and species composition that have occurred, complete restoration of these areas to their historic species assemblages (where this is known) is not feasible, and may even be detrimental (potentially leading to further degradation) in some cases. steps can, however, be taken to remove some of the more aggressive alien species that are well established, whilst also re-introducing key native species into some communities (galatowitsch & richardson 2005). whilst it is impossible to know with certainty which native species dominated in these communities prior to human intervention, the native species that persist provide some clues. for instance, the populus community, which now occurs along 30% of the eerste river, has a number of native species that maintain a tenuous foothold in this novel community. these include the native trees s. mucronata subsp. mucronata, o. europaea subsp. africana and k. africana, all of which provide valuable ecosystem services to both humans and wildlife (holmes et al. 2008; richardson & fraser 1995; venter & venter 1996). it is likely that the dense understorey of t. majus and t. fluminensis within the populus community has suppressed the germination of native seedlings of these species, but they seem able to thrive below a canopy of p. x canescens. consequently, active planting will be necessary to establish denser stands of native vegetation, but little maintenance other than managing the invasive species may be necessary following these initial restoration activities. once native species have grown large enough to provide the necessary soil stabilisation, p. x canescens and other alien trees could be removed in order to provide a greater opportunity for native species to thrive. there is an even greater opportunity for restoration along portions of the river dominated by the quercus community, as a diverse array of native plant species persist amidst the alien species here. in fact, there is considerable overlap between native species in the brabejum community and those in the quercus community, including species such as b. stellatifolium, m. angustifolia, p. serratum, and f. lanceolata. this, combined with the observation that these two communities exist both adjacent to and interspersed with each other, suggests that the quercus community may have historically looked very similar to the brabejum community. as such, land managers interested in restoring portions of the riparian habitat dominated by the quercus community can consider the composition of the brabejum community to discover suitable species for restoration activities. the most appropriate inventory of native species for restoration along a particular stretch of river could further be confirmed by scanning upstream and downstream vegetation for key native species in these areas. the quercus community exists as far downstream as the confluence with the plankenburg river, below which the populus community becomes dominant. interspersed with the populus community is the arundo community, which is relatively rare along the river. the reed species a. donax is highly invasive and has major impacts in many mediterranean-type ecosystems (brooks et al. 2004; coffman, ambrose & rundel 2010; gaertner et al. 2009). efforts should be made to remove this species whilst it still occurs along only a small section of the eerste river. arundo donax tends to crowd out all other species, creating monospecific stands and providing few, if any, of the ecosystem services that other alien species may provide. interspersed with both the populus and salix communities is the bromus community. as with the arundo community, this community is in need of active restoration along the eerste river. the bromus community exists as a result of extensive clearing of woody vegetation by landowners and intensive grazing along portions of the lower river. this has left the river in these areas extremely vulnerable to erosion and sedimentation, adding to the river’s already poor water quality in downstream areas. where this community occurs in communal grazing lands, the appropriate species for restoration can be found amongst the native species in the salix community, primarily s. mucronata subsp. mucronata and c. textilis. in these cases, the riparian area will need to be fenced or otherwise protected from grazing until the plants are of such a size that grazing will not disturb them. this could probably be achieved over one or two growing seasons, given the rapid growth of these species. once these species have established themselves along the riverbank, they will provide not only numerous ecosystem services but also additional forage for cattle being grazed in the area. in other areas of the river where this community exists, the appropriate species for restoration will depend on adjacent communities. where it exists interspersed with the populus community, the appropriate species for restoration will be those listed earlier for the populus community; where it exists adjacent to the quercus community, the appropriate species for restoration can be found amongst the native species in the brabejum or cliffortia communities. no other study from south africa has shown such distinct differences amongst riparian plant communities as we have revealed in the current study. in fact, other studies of riparian communities (e.g. prins et al. 2004) tended to show an overlapping continuum of plant communities rather than a series of unique communities with distinct differences. this is largely because previous studies were confined to riparian communities of headwater systems, where most native vegetation is intact and there is little human interference. invasion by alien plant species and a range of human-mediated habitat modifications in downstream areas of the eerste river have radically changed vegetation structure and composition (meek et al. 2010). this has created an extensive array of novel plant communities, effectively disrupting the natural continuous vegetation patterns through increased local dominance and associated disassembly processes. however, even the two primarily native plant communities (those dominated by brabejum and cunonia), limited to the upper reaches of the river, were quite distinct on the axes of the dca. whilst these communities share many floristic similarities, they are structurally quite different. had the species structural data layers been combined into a single layer, these two communities would likely have overlapped to a much greater extent in the diagram, and would not have been as distinct in the initial cluster analyses. of the 10 plant communities, the brabejum and cunonia communities were also the most tightly clustered in the ordination diagram, whilst the compositional attributes of other plant communities were quite diffuse. this attests to the ever evolving nature of the downstream plant communities, which continue to be changed and shaped by human activities in the surrounding landscape, as well as the major influence of humans on the hydrological regime of the river. conclusion top ↑ an important priority for biodiversity conservation in south africa is the protection and management of riparian habitats, which have become increasingly valuable owing to the multitude of ecosystem services they supply. appropriate management of these riparian areas depends largely upon knowledge of riparian plant communities so that informed decisions can be made regarding their management and restoration. this study details the extensive changes in plant species composition that have occurred in many portions of the eerste river’s riparian zone, and helps to further an understanding of how riparian plant communities in the western cape are changing as a result of anthropogenic activities. furthermore, it provides key data on those native species that continue to persist within and adjacent to alien-dominated communities and offers insights with regard to options for restoration. additional research is needed to determine long-term goals for the riparian vegetation along the eerste river. all stakeholders need to be consulted to establish achievable goals. the results of this study provide important baseline information to guide management aimed at delivering sustainable ecosystem services. acknowledgments top ↑ support for this work was provided by the dst-nrf centre of excellence for invasion biology. we thank suzaan kritzinger-klopper and donald iponga for assistance and companionship in the field, terry trinder-smith for help with identifying plant specimens, christy momberg and mathilda van der vyver for administrative support, lubomír tichý for help with the juice software program, and the many landowners along the eerste river, who kindly allowed us access to their land. c.s.m. also thanks the staff and students at the percy fitzpatrick institute for their support during the study. capenature kindly provided the plant collection permits. competing interests the authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article. authors’ contributions c.s.m. (university of cape town) and d.m.r. (stellenbosch university) were responsible for writing of the manuscript, c.s.m. was responsible for vegetation sampling and data analysis and l.m. (stellenbosch university) assisted with the ordination analysis and classification of plant communities. d.m.r. provided overall project guidance. references top ↑ bray, r.h. & kurtz, l.t., 1945, ‘determination of total, organic, and available forms of phosphorus in soils’, soil science 59, 39–45. http://dx.doi.org/10.1097/00010694-194501000-00006bromilow, c., 2005, problem plants of south africa, briza publications, pretoria. brooks, m.l., d’antonio, c.m., richardson, d.m., grace, j.b., keeley, j.e., ditomaso, j.m. et al., 2004, ‘effects of invasive alien plants on fire regimes’, bioscience 54, 677–688. http://dx.doi.org/10.1641/0006-3568(2004)054[0677:eoiapo]2.0.co;2 brown, c., 1998, ‘the ecological status of western cape rivers investigated: shock survey findings’, african wildlife 52, 27–28. brown, c.a. & dallas, h.f., 1995, eerste river, western cape: situation assessment of the riverine ecosystem. final report, june 1995, southern waters ecological research and consultancy, csir, stellenbosch. brown, c.a. & magoba, r., 2009, ‘rivers and wetlands of cape town’, report no. tt 376/08, water research commission, pretoria. campbell, b.m., 1986, ‘montane plant communities of the fynbos biome’, vegetatio 66, 3–16. http://dx.doi.org/10.1007/bf00044078 chytrý, m., tichý, l., holt, j. & botta-dukát, z. 2002, ‘determination of diagnostic species with statistical fidelity measures’, journal of vegetation science 13, 79–90. http://dx.doi.org/10.1111/j.1654-1103.2002.tb02025.x coffman, g.c., ambrose, r.f. & rundel, p.w., 2010, ‘wildfire promotes dominance of invasive giant reed (arundo donax) in riparian ecosystems of mediterranean-type climate regions’, biological invasions 12, 2723–2734. http://dx.doi.org/10.1007/s10530-009-9677-z cowling, r.m. (ed.), 1992, the ecology of fynbos. nutrients, fire and diversity, oxford university press, cape town. cowling, r.m. & holmes, p.m., 1992, ‘flora and vegetation’, in r.m. cowling (ed.), the ecology of fynbos: nutrients, fire and diversity, pp. 23–61, oxford university press, cape town. cowling, r.m., richardson, d.m. & mustart, p.j., 1997, ‘fynbos’, in r.m. cowling, d.m. richardson & s.m. pierce (eds.), vegetation of southern africa, pp. 99–130, cambridge university press, cambridge. department of water affairs and forestry (dwaf), 1993, ‘western cape system analysis. water quality, volume 1: general’, report no. p g000/00/2891, cape town. department of water affairs and forestry (dwaf), 2004. ‘berg water management area: internal strategic perspective’, report no. p wma 19/000/00/0304, pretoria. foxcroft, l.c., rouget, m. & richardson, d.m., 2007, ‘risk assessment of riparian plant invasions into protected areas’, conservation biology 21, 412–421. http://dx.doi.org/10.1111/j.1523-1739.2007.00673.x, pmid:17391191 gaertner, m., den breeÿen, a., hui, c. & richardson, d.m., 2009, ‘impacts of alien plant invasions on species richness in mediterranean-type ecosystems: a meta-analysis’, progress in physical geography 33, 319–338. http://dx.doi.org/10.1177/0309133309341607 galatowitsch, s.m. & richardson, d.m., 2005, ‘riparian scrub recovery after clearing of invasive alien trees in headwater streams of the western cape, south africa’, biological conservation 122, 509–521. http://dx.doi.org/10.1016/j.biocon.2004.09.008 goldblatt, p. & manning, j.c., 2000, cape plants: a conspectus of the cape flora of south africa, national botanical institute, pretoria. henderson, l., 2001, alien weeds and invasive plants: a complete guide to declared weeds and invaders in south africa, agricultural research council, pretoria (arc-ppri, ppri handbook no. 12). hill, m.o. & gauch, jnr., h.g., 1980, ‘detrended correspondence analysis: an improved ordination technique’, vegetatio 42, 47–58. http://dx.doi.org/10.1007/bf00048870 hobbs, r.j., arico, s., aronson, j., baron, j.s., bridgewater, p., cramer, v.a. et al., 2006, ‘novel ecosystems: theoretical and management aspects of the new ecological world order’, global ecology and biogeography 15, 1–7. http://dx.doi.org/10.1111/j.1466-822x.2006.00212.x holmes, p.m., esler, k.j., richardson, d.m. & witkowski, e.t.f., 2008, ‘guidelines for improved management of riparian zones invaded by alien plants in south africa’, south african journal of botany 74, 538–552. http://dx.doi.org/10.1016/j.sajb.2008.01.182 holmes, p.m., richardson, d.m., esler, k.j., witkowski, e.t.f. & fourie, s., 2005, ‘a decision-making framework for restoring riparian zones degraded by invasive alien plants in south africa’, south african journal of science 101, 553–564. hood, w.g. & naiman, r.j., 2000, ‘vulnerability of riparian zones to invasion by exotic vascular plants’, plant ecology 148, 105–114. http://dx.doi.org/10.1023/a:1009800327334 king, j.m., 1981, ‘the distribution of invertebrate communities in a small south african river’, hydrobiologia 83, 43–65. http://dx.doi.org/10.1007/bf02187150 kruger, f.j., 1978, ‘a description of the fynbos biome project’, south african national scientific programmes report 28. csir, pretoria. linder, h.p., 2003, ‘the radiation of the cape flora, southern africa’, biological reviews 78, 597–638. http://dx.doi.org/10.1017/s1464793103006171, pmid:14700393 linder, h.p., meadows, m.e. & cowling, r.m., 1992, ‘history of the cape flora’, in r.m. cowling, (ed.), the ecology of fynbos: nutrients, fire and diversity, pp. 113–134, oxford university press, cape town. manders, p.t. & richardson, d.m., 1992, ‘colonization of cape fynbos communities by forest species’, forest ecology and management 48, 277–293. http://dx.doi.org/10.1016/0378-1127(92)90150-8 mcdonald, d.j., 1988, ‘a synopsis of the plant communities of swartboschkloof, jonkershoek, cape province’, bothalia 18, 233–260. meek, c.s., richardson, d.m. & mucina, l., 2010, ‘a river runs through it: land-use and the composition of vegetation along a riparian corridor in the cape floristic region, south africa’, biological conservation 143, 156–164. http://dx.doi.org/10.1016/j.biocon.2009.09.021 moffett, r.o., 2007, ‘name changes in the old world rhus and recognition of searsia (anacardiaceae)’, bothalia 37, 165–175. olsen, s.r., cole, c.v., watanabe, f.s. & dean, l.a., 1954, ‘estimation of available phosphorus in soils by extraction with sodium bicarbonate’, usda circular 939, 1–19. planty-tabacchi, a., tabacchi, e., naiman, r.j., deferrari, c. & decamps, h., 1996, ‘invasibility of species-rich communities in riparian zones’, conservation biology 10, 598–607. http://dx.doi.org/10.1046/j.1523-1739.1996.10020598.x prins, n., holmes, p.m. & richardson, d.m., 2004, ‘a reference framework for the restoration of riparian vegetation in the western cape, south africa, degraded by invasive australian acacias’, south african journal of botany 70, 767–776. pyšek, p., richardson, d.m., rejmánek, m., webster, g.l., williamson, m. & kirschner, j., 2004, ‘alien plants in checklists and floras: towards better communication between taxonomists and ecologists’, taxon 53, 131–143. http://dx.doi.org/10.2307/4135498 richardson, d.m. & fraser, m.w., 1995, ‘birds of swartboskloof, jonkershoek valley, stellenbosch: a mountain fynbos habitat’, southern birds 18, 1–44. richardson, d.m., holmes, p.m., esler, k.j., galatowitsch, s.m., stromberg, j.c., kirkman, s.p. et al., 2007, ‘riparian vegetation: degradation, alien plant invasions, and restoration prospects’, diversity and distributions 13, 126–139. http://dx.doi.org/10.1111/j.1366-9516.2006.00314.x rouget, m., cowling, r.m., pressey, r.l. & richardson, d.m., 2003, ‘identifying spatial components of ecological and evolutionary processes for regional conservation planning in the cape floristic region, south africa’, diversity and distributions 9, 191–210. http://dx.doi.org/10.1046/j.1472-4642.2003.00025.x sieben, e.j.j., 2003, ‘the riparian vegetation of the hottentots holland mountains’, phd dissertation, university of stellenbosch. sieben, e.j.j., mucina, l. & boucher, c., 2009, ‘scaling hierarchy of factors controlling riparian vegetation patterns of the fynbos biome at the western cape, south africa’, journal of vegetation science 20, 17–26. http://dx.doi.org/10.1111/j.1654-1103.2009.03989.x sieben, e.j.j. & reinecke, m.k., 2008, ‘description of reference conditions for restoration projects of riparian vegetation from the species-rich fynbos biome’, south african journal of botany 74, 401–411. http://dx.doi.org/10.1016/j.sajb.2008.01.176 taylor, h.c., 1978, ‘capensis’, in m.j.a. werger (ed.), the biogeography and ecology of southern africa, pp. 171–230, w. junk, the hague. http://dx.doi.org/10.1007/978-94-009-9951-0_8 ter braak, c.j.f. & šmilauer, p., 2002, canoco 4.5 reference manual and canodraw for windows user’s guide: software for canonical community ordination, microcomputer power, ithaca, ny. tichý, l., chytrý, m. & zelený, d., juice version 7.0, computer software, viewed n.d. from http://www.sci.muni.cz/botany/juice.htm van wilgen, b.w. & kruger, f.j., 1985, ‘the physiography and fynbos vegetation communities of the zachariashoek catchments, southwestern cape province’, south african journal of botany 51, 379–399. venter, f. & venter, j., 1996, making the most of indigenous trees, briza publications, pretoria. walkley, a., 1935, ‘an examination of methods for determining organic carbon and nitrogen in soils’, journal of agricultural science 25, 598–609. http://dx.doi.org/10.1017/s0021859600019687 westhoff, v. & van der maarel, e., 1973, ‘the braun-blanquet approach’, in r.h. whittaker (ed.), ordination and classification of communities, pp. 617–726, w. junk, the hague. http://dx.doi.org/10.1007/978-94-010-2701-4_20 a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe original research vol. 50 no. 1 pp. 184 201 the plant communities of the andover game reserve, south africa abstract floristic characteristics of the andover game reserve (agr) were surveyed using an area-based survey technique and classified according to the data recorded from 88 relevés, using the phytotab-pc software package. three plant communities, of which two each contain two variants, were described and mapped. the plant communities and their causative environmental factors were validated through detrendedand canonical correspondence multivariate analyses. the plant communities of the agr were found to typify the floristics associated with the catenal sequences located in undulating areas on granite. broad-leaved savanna is located at the crest and upper mid-slopes while fine-leaved savanna occurs along the footslopes of the agr. seeplines, a characteristic occurrence along catenas, are found at the transitional zone between the upper broadand lower fine-leaved savanna plant communities. this study forms the basis for the compilation of a revised ecological management plan for the andover game reserve. keywords: plant communities, phytotab-pc, catena, savanna, granitic soils protected areas have been set aside for the protection of natural resources and to perpetuate the natural conditions (visser et al. 1996) that are necessary for the continued existence of these systems. management, in this context, is the practice by which that purpose is realised (pyle 1980). for the effective management of any natural area a comprehensive description or base line study is of paramount importance (barrett et  al. 2006; brown & bezuidenhout 2000; brown et  al. 2005; coombes & mentis 1992; reilly & macfadyen 1992) as management decisions are based on a comprehensive understanding of the potential of an area. management of a conservation area starts once the area has been defined on a map (spinage 1979). bredenkamp and brown (2001) emphasised the use of plant communities as a reliable basis for ecological planning and management. this paper aims to describe the plant communities of the andover game reserve (agr), south africa, and forms a critical part in the compilation of a revised ecological management plan for the reserve. study area the agr is situated between the southern latitudes 24˚ 33’ and 24˚ 38’ and eastern longitudes 31˚ 10’ and 31˚ 17’, and encompasses an area of 7 100 ha. the agr is approximately 20 km southwest of the orpen gate of the kruger national park. the physiography is characterised by an undulating landscape with interspersed drainages and upper landscapes arranged predominantly from a westerly to an easterly direction. the agr is located within the bushveld lowveld region of the revised köppen climate classification (kruger 2004; schulze 1994) and receives a mean annual precipitation of 782.9 mm (agromet 1996). mean daily maximum temperatures in january and july are 31.7 ˚c and 24 ˚c, with the mean daily minimum being 20.1 ˚c and 9.2 ˚c for the same periods, respectively (agromet 1996). the geology of the pilgrim’s rest area, which includes the lowveld, has been described by walraven (1989). the agr is situated on the swazian basement complex, which consists predominantly of gneisses and granites. the north-western section of the agr comprises grey to pale-brown, mediumto course-grained quartz-feldsparbiotite gneiss with subordinate mafic to ultramafic xenoliths. the south-eastern section of the agr consists of makhutswi gneiss, which is light-grey, medium-grained biotite gneiss with course-grained quartz-feldspar leucosomes (walraven 1986). the reserve incorporates a combination of two veld types, namely veld type 10 or lowveld and veld type 11, also known as arid lowveld (acocks 1988), and collectively described by driver et al. (2005) as granite lowveld bushveld. recently this area has been described as the granite lowveld by mucina and rutherford (2006). methods the level of detail required from a particular study, determines the working scale or associated sampling scale, which in turn determines the smallest mappable unit area both in terms of field procedures and cartography (rutherford & westfall 1986). pressey and bedward (1991) and panagos (1995) illustrated the effect of different scales on the same data set, where at socalled coarse scales the data set showed fewer groups than when classified for finer scales. panagos (1995) and westfall et  al. (1996) recommend a sampling scale of 1:12 000 for areas less than 10 000 ha in extent. a recent (1996) monochrome 1:60 000 aerial photograph was used to construct a photographic mosaic that was then enlarged to a scale of approximately 1:13 000. the photographic mosaic was stratified into eight relatively homogeneous areas using a method that involved the recognition of pattern based on texture and grey values (cronje 2004; panagos et  al. 1998). the number of sampling quadrats positioned in each homogeneous unit depended on the size of heath p. cronje doornkloof nature reserve south africa mike d. panagos brian k. reilly department of nature conservation tshwane university of technology south africa correspondence to: heath p. cronje e-mail: cronjehp@lantic.net postal address: doornkloof nature reserve, po box 94, colesberg, 9795, south africa 184 2008 original research cronje, panagos & reilly koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 184 201 the area: the larger the area the higher the number of sampling quadrats allocated to the unit. a minimum of four sampling quadrats was allocated to the smaller homogeneous units to maximise detection of variability. the sampling quadrat area was set at 200 m2 (10 m x 20 m), relative to the sampling scale (westfall et al. 1997). this quadrat size has become established as an optimised area for savanna (boucher & jarman 1977; bredenkamp 1975; panagos et  al. 1998; panagos & reilly 2006; theron 1973; van rooyen 1983; werger 1974). a total of 88 quadrats were subjectively placed throughout the homogeneous strata as identified on the aerial photographic mosaic. relevés were located in the field using a global positioning system (gps), the aerial photographic mosaic and a topographic map (cronje 2004). each relevé was geo-referenced using a gps and the environmental factors recorded at each quadrat included: topographic unit (i.e. crest, mid-slope, footslope, riverine); slope in degrees (estimated); aspect in degrees, with the aid of a compass; soil form, with diagnostic horizons identified up to a depth of 1.2 m according to macvicar et  al. (1991); soil characteristics were classified (sandy, loam or clay) based on a texture analysis using the ‘sausage test’ (brady 1984); and the visual occurrence of fire and any other noticeable disturbances. the floristic parameters recorded at each quadrat were species composition, growth forms (tree, shrub, dwarf shrub, grass, and forb) according to edwards (1983) and westfall (1992), and canopy cover determined with the use of the plant number scale (westfall & panagos 1988; westfall et al. 1996.) a community composition analysis (panagos et al. 1998; westfall et al. 1996) was carried out for each community and variant. it entailed calculating the projected canopy cover-to-frequency ratios of species in five growth form classes (tree, shrub, dwarf shrub, grass, and forb) using phytotab-pc (westfall 1992). a linear relation between cover and frequency is assumed and the expected cover for the actual frequency of each species in a community is calculated according to the linear regressions. three distinct groups are formed i.e. those with a higher cover and those with a lower cover than the standard error of the mean. the third group of species falls between the first two groups. the first group is referred to as strong competitor species because of the high individual resource-space requirements and the second group is called weak competitor species because of the low individual resource-space requirements. the third group is termed normal competitor species. plant species nomenclature followed that of the national herbarium, pretoria (germishuizen & meyer 2003). the floristic data were classified using the phytotab-pc program package (westfall 1992, westfall et  al. 1997) and the resultant plant communities were validated using a detrended correspondence analysis (dca) using the canoco software package (ter braak & šmilauer 1998). a canonical correspondence analysis (cca) (ter braak & šmilauer 1998) was used to validate floristic and environmental relations. results classification a total of 333 plant taxa were recorded in three main plant communities, with the first two plant communities each having 0 2 41 kilometers strychnos madagascariensis aristida stipitata subsp. graciliflora short closed woodland (community 1) strychnos madagascariensis indigofera inhambanensis (variant 1a) strychnos madagascariensis parinari curatellifolia (variant 1b) acacia gerrardii setaria sphacelata var. sericea short closed woodland (community 2) acacia gerrardii fuirena pachyrrhiza var. pachyrrhiza (variant 2a) acacia gerrardii themeda triandra (variant 2b) gymnosporia senegalensis phragmites mauritianus riparian low thicket (community 3) legend 0 2 41 kilometers strychnos madagascariensis aristida stipitata subsp. graciliflora short closed woodland (community 1) strychnos madagascariensis indigofera inhambanensis (variant 1a) strychnos madagascariensis parinari curatellifolia (variant 1b) acacia gerrardii setaria sphacelata var. sericea short closed woodland (community 2) acacia gerrardii fuirena pachyrrhiza var. pachyrrhiza (variant 2a) acacia gerrardii themeda triandra (variant 2b) gymnosporia senegalensis phragmites mauritianus riparian low thicket (community 3) legend figure 1 plant communities of the andover game reserve 185 plant communities original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 184 201 c o m m u n it ie s 1 2 3 v a r ia n t s 1a 1b 2a 2b r e l e v é n u m b e r 4 6 2 1 1 8 7 1 2 5 3 1 2 1 2 1 1 4 3 2 2 5 7 3 6 2 3 7 6 5 3 1 7 4 4 3 7 6 2 3 3 4 5 7 2 3 2 6 1 4 4 4 4 1 3 6 6 7 4 7 8 8 5 5 8 7 5 5 5 6 8 7 6 5 6 8 8 8 8 5 3 4 8 7 4 7 1 2 8 3 3 9 3 6 8 6 2 4 0 4 2 5 6 5 0 1 8 1 7 3 6 7 2 0 8 1 2 3 6 7 4 9 9 0 9 9 2 5 7 5 1 5 2 9 0 2 8 4 7 5 1 4 3 8 6 0 1 6 3 0 8 5 1 9 4 7 6 0 2 3 1 9 4 8 7 5 6 s p ec ie s g ro u p a : d ia g n o st ic s p ec ie s o f t h e s tr yc h n o s m ad ag as ca ri en si s a ri st id a st ip it at a va r. g ra ci lifl o ra s h o rt c lo se d w o o d la n d (c o m m u n it y 1) s tr yc hn os m ad ag as ca ri en si s 1 1 1 2 + 1 1 1 + 1 + + 4 1 + + + 4 + 1 + + 1 + + 1 1 1 1 1 + 3 1 + li m eu m v is co su m 2 5 1 1 3 1 1 + 3 1 2 1 1 2 1 + 1 1 1 1 5 2 2 2 + + 2 2 1 1 1 1 1 a ri st id a st ip ita ta s ub sp . g ra ci lifl or a 1 2 1 2 2 + 2 1 1 1 1 1 1 1 3 2 2 1 2 1 2 2 1 1 m ac ro ty lo m a ax ill ar e va r. ax ill ar e + + 3 1 2 1 1 + 2 1 + + 1 1 1 1 + + + 2 2 p hi le no pt er a vi ol ac ea 1 + + + 1 + 1 1 1 + 1 + 1 1 + 1 1 1 + + h er m bs ta ed tia o do ra ta + + + + + + + + 1 1 1 1 1 + 1 + + 1 s tr yc hn os s pi no sa + + 1 1 1 1 + + 4 2 + 1 + + 1 + + c ro ta la ri a sc hi nz ii 1 1 + 2 1 1 1 1 1 + + 1 + 1 1 2 c le om e m on op hy lla + + 2 1 1 2 + + 1 + 1 + 1 3 1 1 1 r hy nc ho si a to tt a va r. to tt a 1 1 + 1 1 1 + 1 3 + 1 te ph ro si a lo ng ip es 3 2 3 + 6 1 + 1 1 + s es am um a la tu m + + 1 1 + + 1 1 1 1 h er m an ni a gl an du lig er a + 2 + 1 1 + 1 1 + 1 2 in di go fe ra d al eo id es v ar . d al ei od es 1 3 1 1 1 5 1 1 1 1 c ro ta la ri a bu rk ea na + + 2 1 + + + 1 e ra gr os tis le hm an ni an a 1 + 1 1 2 1 1 1 w ah le nb er gi a ba nk si an a 2 4 2 1 j 2 c om m el in a fo rs ka ol ii + 1 2 1 1 2 + o xy go nu m d re ge an um s ub sp . c an es ce ns 2 1 + 1 1 1 o ld en la nd ia a ffi ni s su bs p. fu ga x + 2 + 1 1 + p se ud og na ph al iu m o lig an dr um 1 + + 1 + s p ec ie s g ro u p b : d ia g n o st ic s p ec ie s o f t h e s tr yc h n o s m ad ag as ca ri en si s i n d ig o fe ra in h am b an en si s v ar ia n t 1 a in di go fe ra in ha m ba ne ns is 2 2 1 1 2 2 1 + + 6 1 2 2 + 3 1 1 + + + c ro to la ri a sp ha er oc ar pa 1 1 + 1 1 1 + 1 + 1 + + 2 1 1 1 + + le uc as n eu fli ze an a + 1 1 5 1 1 2 + 1 1 1 1 + + 1 1 1 1 1 in di go fe ra a st ra ga lin a 3 1 2 1 + 1 2 3 4 1 2 2 + + 1 2 1 k oh au tia c ae sp ito sa s ub sp . b ra ch yl ob a 1 1 + 1 1 1 2 1 2 1 1 1 in di go fe ra c ha rl ie ri an a va r. ch ar lie ri an a 1 1 1 2 + 1 2 + + 1 1 2 + le on ot is n ep et ifo lia 1 1 1 1 1 2 2 + 1 1 1 c ha m ae cr is ta b ie ns is + + 1 1 + + + 2 h er m an ni a qu ar tin ia na 2 + + + 1 + 1 1 + z or ni a m iln ea na 1 + 1 + 2 + 2 + 1 h el ic hr ys um c an do lle an um + + + 1 + + d ic om a m ac ro ce ph al a + 1 1 + h em iz yg ia p et re ns is 2 1 2 1 e ra gr os tis a sp er sa 1 1 1 + h el ic hr ys um s p. + + 1 + a ri st id a m er id io na lis + 3 2 1 1 f ic us th on ni ng ii + + + c om m el in a be ng ha le ns is 2 1 1 te ph ro si a vi llo sa s ub sp . e hr en be rg ia na + + 2 ta b l e 1 ph yt os oc io lo gi ca l c la ss ifi ca ti on o f t he v eg et at io n of th e a nd ov er g am e r es er ve 186 original research cronje, panagos & reilly koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 184 201 c o m m u n it ie s 1 2 3 v a r ia n t s 1a 1b 2a 2b r e l e v é n u m b e r 4 6 2 1 1 8 7 1 2 5 3 1 2 1 2 1 1 4 3 2 2 5 7 3 6 2 3 7 6 5 3 1 7 4 4 3 7 6 2 3 3 4 5 7 2 3 2 6 1 4 4 4 4 1 3 6 6 7 4 7 8 8 5 5 8 7 5 5 5 6 8 7 6 5 6 8 8 8 8 5 3 4 8 7 4 7 1 2 8 3 3 9 3 6 8 6 2 4 0 4 2 5 6 5 0 1 8 1 7 3 6 7 2 0 8 1 2 3 6 7 4 9 9 0 9 9 2 5 7 5 1 5 2 9 0 2 8 4 7 5 1 4 3 8 6 0 1 6 3 0 8 5 1 9 4 7 6 0 2 3 1 9 4 8 7 5 6 s tig a as ia tic a 1 + + a ca nt ho sp er m um h is pi du m + + 1 a nn on a se ne ga le ns is + 1 1 f ic us s tu hl m an ni i 1 1 + + h ib is cu s ca nn ab in us + + 1 + ja tr op ha s pi ca ta + + m on ec hm a de bi le 2 2 c uc um is z ey he ri + + s ch iz ac hy ri um je ff re ys ii 2 2 + p te ro ca rp us a ng ol en si s 1 + s id a al ba 1 1 s p ec ie s g ro u p c : d ia g n o st ic s p ec ie s o f t h e s tr yc h n o s m ad ag as ca ri en si s p ar in ar i c u ra te lli fo lia v ar ia n t 1 b p te ro ca rp us r ot un di fo lia + + + 1 1 + h yp er ic um la la nd ii 1 3 1 a nt id es m a ve no su m + + + p ar in ar i c ur at el lif ol ia 1 + s et ar ia s ph ac el at a va r. se ri ce a 1 2 m om or di ca b al sa m in a + 1 1 te ph ro si a rh od es ic a va r. rh od es ic a 3 1 z in ni a pe ru vi an a + 1 p as pa lu m s cr ob ic ul at um 4 1 c ro ta la ri a la nc eo la ta s ub sp . l an ce ol at a + 2 s p ec ie s g ro u p d : d ia g n o st ic s p ec ie s o f t h e a ca ci a g er ra rd ii s et ar ia s p h ec el at a va r. s er ic ea s h o rt c lo se d w o o d la n d (c o m m u n it y 2) a ca ci a ge rr ar di i + + 1 1 5 5 1 + 1 1 + 3 9 a 1 5 5 6 6 4 8 7 6 a 5 4 c 8 3 + c om br et um h er er oe ns e 4 + 1 2 + + g 3 5 5 b 3 6 4 l 1 5 4 4 a 6 1 7 + 6 + + s et ar ia s ph ac el at a va r. sp ha ce la ta 9 3 2 8 3 e 2 9 a 5 7 8 4 9 3 4 a 7 1 1 c ha et ac an th us s et ig er 3 3 5 3 1 4 + 2 6 1 8 3 2 4 5 5 7 2 a 1 ju st ic ia p ro tr ac ta s ub sp . r od es ia na 2 1 1 1 5 2 1 2 2 1 3 + 2 + v er no ni a ol ig oc ep ha la 1 1 + + 2 + 3 1 + 2 + + d ih et er op og on a m pl ec te ns 4 + 2 3 + 2 2 3 2 1 8 h el ic hr ys um o xy ph yl lu m 1 + 2 + 2 1 + 1 + + lo ud iti a fla vi da 1 1 3 1 4 1 9 1 b ot hr io ch lo a in sc ul pt a 1 1 + + + 2 1 1 b ra ch ia ri a xa nt ho le uc a 2 6 1 2 + 1 1 m ur da nn ia s im pl ex 1 1 1 2 1 1 k yl lin ga a lb a + 1 + 1 1 1 2 a bu til on a us tr o -a fr ic an um 1 2 + 1 c hl or is v ir ga ta 1 1 1 1 + s ol an um c at om be le ns e + 1 + + + s p ec ie s g ro u p e : d ia g n o st ic s p ec ie s o f t h e a ca ci a g er ra rd ii f u ir en a p ac hy rr h iz a va r. p ac hy rr h iz a v ar ia n t 2 a f ui re na p ac hy rr hi za v ar . p ac hy rr hi za + 2 a 9 2 e 4 1 1 1 4 3 1 3 4 v ah lia c ap en si s su bs p. v al ga ri s va r. vu lg ar is 4 1 4 1 6 4 2 + 2 1 t a b l e 1 (c o n t. ..) 187 plant communities original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 184 201 c o m m u n it ie s 1 2 3 v a r ia n t s 1a 1b 2a 2b r e l e v é n u m b e r 4 6 2 1 1 8 7 1 2 5 3 1 2 1 2 1 1 4 3 2 2 5 7 3 6 2 3 7 6 5 3 1 7 4 4 3 7 6 2 3 3 4 5 7 2 3 2 6 1 4 4 4 4 1 3 6 6 7 4 7 8 8 5 5 8 7 5 5 5 6 8 7 6 5 6 8 8 8 8 5 3 4 8 7 4 7 1 2 8 3 3 9 3 6 8 6 2 4 0 4 2 5 6 5 0 1 8 1 7 3 6 7 2 0 8 1 2 3 6 7 4 9 9 0 9 9 2 5 7 5 1 5 2 9 0 2 8 4 7 5 1 4 3 8 6 0 1 6 3 0 8 5 1 9 4 7 6 0 2 3 1 9 4 8 7 5 6 b ra ch ia ri a br iz an th a 1 5 2 2 1 a ly si ca rp us v ag in al is v ar . v ag in al is + 1 1 3 1 s p ec ie s g ro u p f: d ia g n o st ic s p ec ie s o f t h e a ca ci a g er ra rd ii t h em ed a tr ia n d ra v ar ia n t 2 b b le ph ar is in te gr ifo lia v ar . i nt eg ri fo lia 3 + + + 2 1 1 1 + + 1 1 + t he m ed a tr ia nd ra 2 2 1 + 4 1 + 2 3 + 1 3 g re w ia b ic ol or 1 1 1 1 1 + + + + + 1 + o rm oc ar pu m tr ic ho ca rp um 1 + + + + 1 1 + + + + 1 + m ic ro ch lo a ca ff ra 1 1 1 2 2 2 6 p le ct ro ni el la a rm at a 1 1 2 + + + + + + p ap pe a ca pe ns is + + + + + a sp ar ag us b uc ha na ni i + + + + + tr ag ia r up es tr is 1 1 1 1 1 + g ar de ni a vo lk en si i + + 1 + + p an ic um c ol or at um + 2 2 + 2 2 ip om oe a ob sc ur a va r. ob sc ur a + 1 + + + s po ro bo lu s ni te ns 2 3 + 2 2 2 a ca ci a bu rk ei + 6 + + 1 a bi ld ga ar di a tr ifo lia 3 6 1 2 c ym bo po go n ex ca va tu s + 3 2 + 2 b uc hn er a sp . + 1 + in di go fe ra s p. 3 + 1 1 b ar le ri a sa xa til is 1 + 1 1 e ri oc hl oa s ta pfi an a 1 2 6 le de bo ur ia s p. 1 + 1 a lo e m ar lo th ii + + 1 c om m ip ho ra s ch im pe ri + 1 1 + h ib is cu s pu si llu s 2 1 1 2 a ca ci a ca ff ra 1 + e nt er op og on m on os ta ch yu s su bs p. a fr ic an us 4 8 z an th ox yl um c ap en se 1 + + o rt ho si ph on s uf fr ut es ce ns 1 5 5 k al an ch oe r ot un di fo lia 1 + t he si um p al lid um 1 + s p ec ie s g ro u p g : s p ec ie s co m m o n to c o m m u n it y 1 an d v ar ia n t 2 a h yp er th el ia d is so lu ta h h 2 a f 4 + 8 9 5 2 b + c e 8 8 4 h 1 1 1 g 8 7 e 2 c 1 6 l 9 1 + i f 9 7 1 b d 8 9 6 j 5 d g 6 f d ic er oc ar yu m s en ec io id es 2 + 2 1 2 2 1 1 + 1 1 1 1 1 1 1 1 1 + 1 2 + 1 2 3 1 + + + 2 2 1 2 1 1 1 2 + 1 + 2 1 1 1 h ib is cu s sp . 1 1 9 1 2 1 2 2 1 2 + 7 3 2 2 2 1 8 1 + 1 1 4 2 a 1 1 + c 1 1 3 1 3 1 2 2 1 1 3 3 4 2 c ha m ae cr is ta a bs us 5 4 3 9 1 1 1 5 7 6 1 4 + 1 9 1 c 1 1 3 9 2 + 3 6 7 7 1 + 8 + 1 1 2 1 7 3 + + 1 + 1 tr ic lic er as la ce ra tu m 2 1 1 1 3 1 1 1 + 1 + + 1 1 1 + 1 1 1 + + 1 + 1 2 + 1 1 + 1 1 1 s en na p et er si an a + + + 1 1 + 1 + 2 1 + 1 1 + 1 1 1 + 1 1 + 1 1 + + + 2 2 + 1 c yn od on d ac ty lo n 2 4 1 1 1 2 2 3 9 8 1 2 2 2 1 1 1 e 1 5 2 3 c 1 1 2 g 5 6 lo be lia e ri nu s 1 1 1 + 2 1 1 + 2 2 + 1 1 1 1 1 1 1 2 1 + + t a b l e 1 (c o n t. ..) 188 original research cronje, panagos & reilly koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 184 201 c o m m u n it ie s 1 2 3 v a r ia n t s 1a 1b 2a 2b r e l e v é n u m b e r 4 6 2 1 1 8 7 1 2 5 3 1 2 1 2 1 1 4 3 2 2 5 7 3 6 2 3 7 6 5 3 1 7 4 4 3 7 6 2 3 3 4 5 7 2 3 2 6 1 4 4 4 4 1 3 6 6 7 4 7 8 8 5 5 8 7 5 5 5 6 8 7 6 5 6 8 8 8 8 5 3 4 8 7 4 7 1 2 8 3 3 9 3 6 8 6 2 4 0 4 2 5 6 5 0 1 8 1 7 3 6 7 2 0 8 1 2 3 6 7 4 9 9 0 9 9 2 5 7 5 1 5 2 9 0 2 8 4 7 5 1 4 3 8 6 0 1 6 3 0 8 5 1 9 4 7 6 0 2 3 1 9 4 8 7 5 6 p hy lla nt hu s re tic ul at us 1 + + 1 1 + + 1 + + 1 1 + 1 2 + 1 2 1 3 c yp er us in de co ru s va r. in de co ru s 1 + 1 1 1 1 1 1 1 1 2 + 1 1 1 1 1 1 p ili os tig m a th on ni ng ii + + + + 1 1 + + 2 1 1 + + 1 + g om ph oc ar pu s fr ut ic os us 1 2 + 1 1 1 + + 1 s p ec ie s g ro u p h : s p ec ie s co m m o n to v ar ia n t 1 b an d c o m m u n it y 2 e ra gr os tis in am oe na 1 4 + 6 3 2 1 1 2 b 1 4 + 1 + 2 c yp er us d en ud at us 1 2 1 6 7 1 2 5 6 2 8 1 3 + 2 a ca ci a sw az ic a + + + + 1 + 9 1 2 2 4 g 1 p yr os tr ia h ys tr ix + + + 1 1 + + + + + 1 + 1 c ar is sa e du lis + 1 + 1 + + + + + + + + + c ar is sa b is pi no sa + + + + + + 1 + + + 1 h el io tr op iu m o va lif ol iu m 1 + 1 2 1 + + + 1 1 e ra gr os tis tr ic ho ph or a 1 1 + + + + 1 s p ec ie s g ro u p i: s p ec ie s co m m o n to c o m m u n it y 1 an d 2 w al th er ia in di ca 2 1 1 3 2 3 3 4 1 1 1 8 1 2 2 1 1 1 1 + 3 4 1 + 1 4 2 + 8 7 4 1 1 3 2 2 1 2 3 2 2 2 1 1 4 1 4 + 2 2 2 2 1 2 2 1 3 5 1 1 2 2 2 1 1 1 4 2 1 3 1 1 + 1 2 1 + 1 + 1 1 2 + + p og on ar th ri a sq ua rr os a 1 1 3 4 1 4 1 1 2 3 1 3 1 2 2 1 1 2 2 2 3 3 2 1 3 4 2 2 5 5 2 4 2 8 1 3 4 1 2 2 1 3 1 1 2 3 4 1 2 2 2 2 1 1 2 3 1 2 2 2 2 + 1 5 5 4 2 1 1 1 1 1 1 + 1 + 2 1 + te rm in al ia s er ic ea 6 a e 5 6 2 i 9 4 9 + h b 9 5 7 2 l 3 4 7 k 9 9 9 7 c q d 6 5 1 1 3 c + 4 4 o j 1 a 6 6 c o 4 i f b 4 1 + 2 4 + 2 1 1 1 1 1 1 1 5 3 + + 2 + + + + d ig ita ri a er ia nt ha 2 2 4 3 2 3 1 3 5 3 3 1 2 1 6 2 5 2 3 1 3 5 1 2 + 2 5 j 9 2 1 4 4 3 3 1 5 5 2 3 1 4 1 2 3 8 3 5 1 3 1 3 1 3 2 3 1 2 3 9 8 1 b 4 4 6 3 1 2 1 1 2 e ra gr os tis r ig id io r 1 + 2 1 3 + 1 1 3 2 2 + 1 2 1 1 1 5 2 + 2 + 1 2 1 2 1 1 1 2 1 1 1 2 1 1 3 2 1 1 2 1 1 1 1 2 2 3 2 + 1 + 1 2 1 2 + 1 2 2 1 1 3 3 3 1 + 2 + 1 + p er ot is p at en s 2 2 8 a a i 1 1 2 8 1 5 2 1 6 1 1 2 1 3 9 2 1 3 4 2 2 6 3 1 5 3 1 a 2 6 2 1 9 1 1 3 2 2 2 1 1 1 2 3 1 1 1 3 4 1 1 1 1 1 2 1 2 + + 1 c ha m ae cr is ta m im os oi de s 1 1 1 1 1 1 1 2 1 2 3 2 1 + 1 1 1 1 1 1 1 1 + 1 + 1 1 1 2 2 1 1 1 7 2 1 1 1 1 1 3 1 + 1 1 + 1 1 3 + 1 1 + 2 1 3 3 1 1 1 1 1 + 1 1 b ul bo st yl is c on te xt a 5 6 2 a 6 2 6 3 1 3 3 3 2 1 4 1 2 2 2 6 4 3 3 7 4 2 2 2 1 + 2 7 7 3 2 4 1 3 2 3 + 2 1 1 4 5 f 1 5 f 3 k 3 1 3 4 2 8 3 4 1 1 1 1 2 a ga th is an th em um b oj er i s ub sp . b oj er i 5 + 1 + 1 + + + + 2 1 6 + 1 1 1 7 1 1 + 1 g 7 2 1 1 9 1 2 2 1 2 2 2 5 5 7 8 2 1 + 3 a 3 1 2 8 3 + 2 1 6 4 2 3 2 5 + 1 1 2 3 2 1 tr ic hn eu ra g ra nd ig lu m is 2 1 2 3 2 2 1 + 1 2 2 2 2 2 3 1 1 2 3 5 1 + 1 4 1 2 2 2 1 3 2 6 1 6 1 1 4 1 3 1 1 1 1 2 1 1 1 1 1 1 3 + 1 1 3 1 1 2 s cl er oc ar ya b ir re a su bs p. c af fr a 1 1 1 + 4 3 7 3 + 1 4 2 3 1 2 + 1 1 1 + 1 1 1 + 1 + 1 + + + 1 + + 1 + 1 1 + 1 1 + + + + + 1 + + 1 + + + 1 + + + + + + v er no ni a st ee tz ia na 2 1 d 8 c 5 3 5 1 3 2 5 2 4 5 2 1 f 6 b 7 2 6 + 3 2 3 f 5 b 8 + 2 + 7 1 1 1 3 2 4 3 1 1 + 1 2 1 + 2 1 1 7 1 + 1 + 1 m er re m ia tr id en ta ta s ub sp . a ng us tif ol ia 6 1 2 2 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 1 1 1 1 3 1 3 1 1 1 + 1 1 2 2 + 3 2 1 + 1 1 1 1 1 2 3 1 1 + + s ty lo sa nt hu s fr ut ic os a 1 1 1 1 + + + 2 1 2 1 2 + 1 2 1 3 3 1 3 1 1 1 3 1 2 2 2 2 1 1 2 4 1 2 + 1 1 2 1 2 + 2 + + + + 2 1 + 2 3 1 1 1 + g ym no sp or ia g la uc op hy lla + + + 1 + + 2 + 1 + + + + 1 + 1 1 1 + 1 + + + + + + 1 1 + + 1 1 + 2 1 2 1 1 1 1 + + + + + + 1 + 2 1 + + 1 a ri st id a co ng es ta s ub sp . b ar bi co lli s 2 2 1 2 + 1 1 1 1 1 3 1 2 1 1 1 1 + 1 2 1 2 1 1 2 2 2 1 1 2 1 + 1 + 2 1 1 1 2 1 1 + + 1 1 + 2 1 1 1 1 1 1 in di go fe ra fi lip es 2 + 4 1 4 1 1 2 1 2 3 1 2 1 2 1 1 3 2 2 2 1 + 1 1 2 1 1 2 1 1 4 + 2 2 1 1 1 1 2 1 2 + 1 1 2 1 1 2 1 1 o ld en la nd ia h er ba ce a va r. he rb ac ea + + 1 1 4 + + 4 2 + 2 + 2 1 2 1 2 2 1 1 1 3 1 2 3 1 2 2 5 1 3 + 2 1 3 2 1 3 9 2 1 3 3 3 + 5 1 2 2 + + te ph ro si a pu rp ur ea s ub sp . l ep to st ac hy a va r. le pt os ta ch ya 1 1 + 1 3 1 4 1 1 + 1 1 3 1 1 1 1 1 1 1 1 1 1 6 2 + 2 1 2 1 + 2 + 1 1 + + + 1 1 1 + 2 + 2 + + + + 1 u ro ch lo a m os am bi ce ns is 1 2 1 7 2 4 1 + 1 1 3 3 3 2 + 1 1 1 1 1 + 1 5 1 1 b 1 5 1 7 1 3 1 + 1 3 1 1 1 2 1 + 3 + 2 1 1 2 1 + s pe rm ac oc e se ne ns is 2 3 1 2 1 2 2 1 1 1 3 2 8 1 3 2 2 1 2 + 4 1 3 1 5 4 4 6 2 2 1 2 1 2 3 2 2 + 1 2 1 1 1 2 3 1 + s id a co rd ifo lia 1 + 1 + + 1 1 2 1 1 1 + 2 1 + 1 1 1 1 1 2 1 2 1 2 1 2 1 1 1 2 1 + 2 2 1 2 + + + 1 + + c yp er us o bt us ifl or us v ar . o bt us ifl or us + 2 + 2 1 1 1 1 + 1 + 1 1 1 2 3 1 1 2 1 1 3 1 2 1 + 1 2 2 1 1 + 1 1 1 1 1 1 1 1 1 1 d io sp yr os m es pi lif or m is 1 + + + + + + + + + + + 1 + + 1 1 + + 1 + + + 3 1 + + + + 1 2 1 + + + + + + + 2 c or ch or us c on fu su s 1 1 1 1 1 1 2 1 1 2 1 2 2 1 1 1 1 + 1 1 2 + 1 2 2 + 2 1 + 1 1 + 1 1 1 1 + + c om br et um c ol lin um 1 2 9 + 4 3 2 + 3 3 1 + 1 + + + + + 5 + 1 1 + 5 + + + 3 1 2 7 1 2 + + 8 2 h et er op og on c on to rt us + 1 1 + 1 1 + + 2 2 + 2 1 2 2 3 + 1 1 1 7 2 1 1 4 4 3 2 3 5 2 6 1 2 1 1 2 t a b l e 1 (c o n t. ..) 189 plant communities original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 184 201 c o m m u n it ie s 1 2 3 v a r ia n t s 1a 1b 2a 2b r e l e v é n u m b e r 4 6 2 1 1 8 7 1 2 5 3 1 2 1 2 1 1 4 3 2 2 5 7 3 6 2 3 7 6 5 3 1 7 4 4 3 7 6 2 3 3 4 5 7 2 3 2 6 1 4 4 4 4 1 3 6 6 7 4 7 8 8 5 5 8 7 5 5 5 6 8 7 6 5 6 8 8 8 8 5 3 4 8 7 4 7 1 2 8 3 3 9 3 6 8 6 2 4 0 4 2 5 6 5 0 1 8 1 7 3 6 7 2 0 8 1 2 3 6 7 4 9 9 0 9 9 2 5 7 5 1 5 2 9 0 2 8 4 7 5 1 4 3 8 6 0 1 6 3 0 8 5 1 9 4 7 6 0 2 3 1 9 4 8 7 5 6 p hy lla nt hu s m ad er as pa te ns is 6 + 1 1 + 1 1 1 1 1 2 + + 1 1 2 1 1 1 1 1 1 1 + 1 1 1 + + 2 2 2 1 1 1 e uc le a di vi no ru m + + 1 + 1 + 1 + + + + a 2 + 3 1 + 2 5 1 7 4 + + 4 5 + 3 2 4 e 7 6 + k oh au tia v ir ga ta 1 2 1 + 1 1 1 1 2 + 5 1 1 4 2 1 1 6 + 1 3 1 1 3 2 1 3 + 1 + 1 1 + c od di a ru di s + + + + + + + + + + 1 + + 1 + + + 1 + 1 + 2 1 + + + 2 + 1 + 1 1 + m el in is n er vi gl um is 1 + 1 + + + + 1 + + + + 1 2 4 + 1 + 1 + 1 1 1 1 1 1 1 1 + + 1 a lb iz ia h ar ve yi + + + + + + + + + 1 1 + 6 1 2 + 6 1 2 + 1 + + + + 1 1 8 3 1 1 la nt an a ru go sa + + + + + 1 + + 1 + 1 + 1 + + 2 1 1 1 1 + + 1 + 2 + 1 1 + 1 c on yz a al bi da 1 + 1 1 3 3 + 1 + 1 1 1 2 1 + 1 + 5 + 1 1 1 2 1 1 + 1 1 + e vo lv ul us a ls in oi de s + + 1 + + 2 1 + 1 1 2 1 + 1 + + 1 1 1 1 1 + 2 2 1 + + c he ila nt he s vi ri di s va r. gl au ca + + + 1 + + + + 1 3 1 + 1 1 1 5 + 2 1 2 1 1 1 + 1 + 1 e ra gr os tis g um m ifl ua 1 6 + 1 2 3 5 2 5 1 + 4 d + 1 1 5 9 1 7 1 5 3 3 8 7 c om m el in a af ri ca na v ar . l an ci sp at ha + 1 2 + 1 3 2 1 1 2 2 + 2 1 + 1 2 1 + 1 + 1 2 + 1 + e la eo de nd ro n tr an sv aa le ns e + + 1 + 1 1 + + + + + + 1 + + + + + + + + 1 + + + p ol lic hi a ca m pe st ri s + + 2 + 1 + + 1 1 1 1 1 + + + + 1 + + + 1 + 1 e ra gr os tis s up er ba + 1 1 1 4 1 1 1 1 1 1 2 + 2 1 1 1 1 2 2 + o ci m um g ra tis si m um s ub sp . g ra tis si m um v ar . gr at is si m um + + 1 + + 1 1 + + + + 1 1 1 2 1 1 + + o ci m um a m er ic an um v ar . a m er ic an um 1 + + 1 + 1 + + 1 7 1 1 1 2 1 2 + 2 + m el ha ni a pr os tr at a + + 1 + 3 + 1 1 1 1 + 1 1 1 + 1 + 1 1 + d al be rg ia m el an ox yl on + + + + + 1 2 + + + 1 + + 1 3 1 + + + w ah le nb er gi a un du la ta + + + + 1 1 2 1 1 1 1 1 1 1 3 + 1 + + p ri va c or di fo lia v ar . a by ss in ic a + + 1 + 1 + + + + 1 1 + + 1 + + + 1 tr iu m fe tt a pe nt an dr a 1 1 2 6 3 2 2 3 2 1 1 5 1 1 4 1 1 m on so ni a an gu st ifo lia 1 2 1 1 + + + 3 1 5 1 1 1 2 1 + + te ph ro si a pu rp ur ea s ub sp . l ep to st ac hy a va r. pu be ns ce ns 1 + 1 1 1 1 1 1 1 1 1 1 + + + 1 + h ib is cu s m ic ra nt hu s + + + + + + 1 + + + 1 tr ic ho la en a m on ac hn e + 1 + + 1 + 1 1 1 + + h el io tr op iu m s tr ig os um 1 2 1 + + 1 + + + + a ca ci a ex uv ia lis 1 + 1 + 1 + 1 + 2 1 2 c om br et um m ol le + 1 1 + + + 1 2 + s id a dr eg ei 1 + 2 + + + 1 + 1 3 c om br et um z ey he ri + + 1 + + 1 + + s p ec ie s g ro u p j: d ia g n o st ic s p ec ie s o f t h e g ym n o sp o ri a se n eg al en si s p h ra g m it es m au ri ti an u s ri p ar ia n lo w t h ic ke t ( c o m m u n it y 3) e us ta ch ys p as pa lo id es 1 4 2 i 8 c uc um is m et ul ife ru s 2 1 + + h yp oe st es fo rs ka ol ii ‘fo rm b ’ a c i + p er si ca ri a co ny zo id es 2 1 4 3 e ri oc hl oa m ey er ia na s ub sp . m ey er ia na a 5 2 9 g ym no sp or ia s en eg al en si s k 3 1 + t hu nb er gi a ne gl ec ta 3 + 1 + le uc a gl ab ra ta v ar . g la br at a 1 1 1 p hr ag m ite s m au ri tia nu s 8 n h t a b l e 1 (c o n t. ..) 190 original research cronje, panagos & reilly koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 184 201 c o m m u n it ie s 1 2 3 v a r ia n t s 1a 1b 2a 2b r e l e v é n u m b e r 4 6 2 1 1 8 7 1 2 5 3 1 2 1 2 1 1 4 3 2 2 5 7 3 6 2 3 7 6 5 3 1 7 4 4 3 7 6 2 3 3 4 5 7 2 3 2 6 1 4 4 4 4 1 3 6 6 7 4 7 8 8 5 5 8 7 5 5 5 6 8 7 6 5 6 8 8 8 8 5 3 4 8 7 4 7 1 2 8 3 3 9 3 6 8 6 2 4 0 4 2 5 6 5 0 1 8 1 7 3 6 7 2 0 8 1 2 3 6 7 4 9 9 0 9 9 2 5 7 5 1 5 2 9 0 2 8 4 7 5 1 4 3 8 6 0 1 6 3 0 8 5 1 9 4 7 6 0 2 3 1 9 4 8 7 5 6 a ge ra tu m c on yz oi de s 5 4 1 p an ic um d es te um 2 2 4 3 1 h et er om or ph a tr an sv aa le ns is + 1 c yp er us s ex an gu la ri s 2 2 s pi ro st ac hy s af ri ca na 1 7 2 s p ec ie s g ro u p k : s p ec ie s co m m o n to v ar ia n t 2 b an d c o m m u n it y 3 a sp ar ag us s et ac eu s 1 + + + + + + + + + + 1 + 1 + + + r ho ic is su s tr id en ta ta 1 + + + + 1 + + + s p ec ie s g ro u p l: c o m m o n an d ra re s p ec ie s ar ra n g ed a cc o rd in g to c o n st an cy v al u es p an ic um m ax im um 2 1 2 2 1 2 b a c c g 3 4 d 5 d 5 8 g a 5 a 6 2 a 5 3 7 a 9 6 6 6 a 5 2 l j 2 9 a 7 g 8 b 6 9 2 2 2 g 3 2 2 a 7 3 5 1 1 2 n k 2 b 3 b 8 8 3 2 4 3 6 d 4 d k 5 a 8 6 e 6 7 d ic hr os ta ch ys c in er ea + 7 a 9 b 9 5 e b 5 7 f k 7 7 c 2 6 e a a 3 6 1 b a 5 2 5 5 7 5 h 9 6 2 e 7 6 a d 9 1 6 3 8 7 1 g c 5 b 5 7 8 i l 7 1 a 4 5 3 5 b f 1 a 8 + 8 4 + + + 4 + b 9 4 7 + 7 1 1 p hy lla nt hu s pa rv ul us v ar . g ar ip en si s 3 1 1 2 1 1 5 3 1 2 3 3 4 7 1 2 5 3 4 4 1 1 2 5 2 1 2 1 2 5 4 1 2 2 1 7 1 3 1 2 1 2 3 1 2 2 1 2 1 1 1 + 1 1 1 1 3 1 1 1 1 1 1 1 1 2 2 1 2 5 1 1 1 3 2 3 2 1 1 2 1 + 1 1 1 p ol yg al a sp he no pt er a 2 + 1 + + 1 1 1 1 1 2 1 2 1 1 2 1 1 + 1 1 1 1 1 + 2 + 1 2 1 1 + 1 1 2 1 1 1 1 1 + 1 1 1 1 1 1 1 1 1 1 2 1 1 3 1 2 1 + 3 + + a ca ci a ni lo tic a 2 + 1 4 + 1 + + + 6 4 3 7 5 1 + 4 2 + 8 6 5 c 1 1 + 8 + + + 1 + 9 2 4 1 + 4 3 7 6 + + 1 + + + + 1 + + 6 + 7 p + + + b id en s bi pi nn at a 1 1 + 2 + 1 + 1 + 2 + 1 1 1 1 2 8 1 1 1 1 + 3 6 8 1 + d 1 2 1 2 1 1 5 3 + 1 1 1 1 1 1 1 1 5 + 1 1 + + 1 + ja sm in um fl um in en se 1 1 + + + 1 + 2 + 1 1 2 1 + 1 1 1 3 1 1 4 + 1 + 1 1 + 1 + 7 1 2 + 5 1 1 2 + 1 + 2 + 2 + + + 4 4 2 + e uc le a na ta le ns is s ub sp . a ng us tifl ia + + + + + + 2 + + 1 1 + + + + + 1 2 + + + + + + 1 1 + + 6 1 1 4 1 + + + + + + 1 + + + 1 + + + z iz ip hu s m uc ro na ta + + + + + + + + + + + + 1 + + 1 1 + + + 1 1 + + 6 1 + + + + 1 + 3 2 1 + + 2 + + 1 2 e hr et ia a m oe na + 1 + 1 + + 1 + 1 1 + 1 + + + + 1 + + + + + 1 + 1 1 2 + + + + + + + 1 + + 1 1 + + e pa lte s ga ri ep in a 5 3 + + + + 1 1 + + + 2 6 + 1 + 1 8 d l 3 6 4 5 d 6 e 2 2 a a 8 1 a 4 1 1 7 c 1 + f lu eg ge a vi ro sa 1 + 1 + + + + + 1 + + + 1 + + + + + + + 1 + + 1 1 + + + + 1 + + + + + + 1 5 3 + + tr iu m fe tt a sp . 2 1 5 1 1 + 1 1 3 1 6 5 1 1 8 2 1 1 + 1 2 1 1 1 1 + + 1 + 1 1 1 1 + + + 2 + 1 1 p el to ph or um a fr ic an um 1 + + + + + + + 1 + 1 + + + 2 + + 3 1 + 6 3 + 3 1 1 + + + 1 + 1 + 4 1 1 + + g re w ia fl av es ce ns v ar . fl av es ce ns + + + 1 + 1 l + + + + + 1 + 1 + 1 + + 3 2 + 1 + + + + 1 1 + 1 2 + + + c om m el in a ec kl on ia na 1 + 2 + + 1 1 1 + 1 + + 2 + 2 1 1 + 2 1 1 1 + 1 1 + 1 1 + + 1 2 2 1 e ra gr os tis c ur vu la + 1 + 1 1 9 2 2 2 1 e 5 2 2 + 1 3 3 2 5 9 1 6 2 3 5 3 a 6 1 1 + h os lu nd ia o pp os ita 1 + + + 1 2 3 + 1 + + 1 4 3 + + 1 1 + 2 1 + 1 1 + ta ge te s m im ut a 1 4 1 6 2 + d 4 d 3 + 2 + 2 + 1 1 1 a + 2 1 + + a ch yr op si s le pt os ta ch ya 2 1 1 + 1 1 1 1 + + 1 1 1 3 2 + 1 3 1 3 2 + s ch ku hr ia p in na ta 1 1 + 2 1 2 2 + 1 + + + 2 6 1 1 1 + + 1 + s po ro bo lu s af ri ca nu s + + 1 2 2 2 2 5 1 2 1 1 2 4 1 1 3 2 1 + + c om br et um im be rb e 1 4 + + + + + 3 + 5 + 1 3 + 1 + + 2 n 2 2 c yp er us s ph ae ro sp er m us 2 2 1 + 1 + 1 1 1 1 2 3 3 1 1 3 1 e 1 1 a ca ly ph a vi lli ca ul is 2 3 1 3 + 2 2 1 2 5 5 + 3 + + 1 + + + s ol an um p an du ri fo rm e 2 + 1 + 2 1 7 1 + 1 2 1 + + 1 3 + 1 + ip om oe a pl eb ei a su bs p. a fr ic an a + + 1 2 1 1 + 1 2 1 s pe ci es a (u ni de nt ifi ed ) + + 3 3 1 + 1 + 2 li pp ia ja va ni ca 1 + + + + + + 1 r hu s pe nt he ri + + 1 + + 1 + la nn ea s ch w ei nf ur th ii va r. st uh lm an ni i + + + + + + + k al an ch oe c re na ta s ub sp . c re na ta v ar . c re na ta 1 + + 1 1 1 + g re w ia m on tic ol a + + + + + 1 1 t a b l e 1 (c o n t. ..) 191 plant communities original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 184 201 c o m m u n it ie s 1 2 3 v a r ia n t s 1a 1b 2a 2b r e l e v é n u m b e r 4 6 2 1 1 8 7 1 2 5 3 1 2 1 2 1 1 4 3 2 2 5 7 3 6 2 3 7 6 5 3 1 7 4 4 3 7 6 2 3 3 4 5 7 2 3 2 6 1 4 4 4 4 1 3 6 6 7 4 7 8 8 5 5 8 7 5 5 5 6 8 7 6 5 6 8 8 8 8 5 3 4 8 7 4 7 1 2 8 3 3 9 3 6 8 6 2 4 0 4 2 5 6 5 0 1 8 1 7 3 6 7 2 0 8 1 2 3 6 7 4 9 9 0 9 9 2 5 7 5 1 5 2 9 0 2 8 4 7 5 1 4 3 8 6 0 1 6 3 0 8 5 1 9 4 7 6 0 2 3 1 9 4 8 7 5 6 a br us p re ca to ri us s ub sp . a fr ic an us 1 1 + 1 + + e hr et ia o bt us ifo lia + + + + + + d om be ya r ot un di fo lia + + + + 1 p hy lla nt hu s in cu rv us 1 1 c om br et um a pi cu la tu m 1 1 3 + m el ha ni a fo rb es ii + 1 3 1 b er ch em ia z ey he ri + 1 + + ju st ic ia fl av a 1 1 2 + tu rb in a ob lo ng at a + 1 1 1 la gg er a cr is pa ta + + 1 1 c yp ho st em m a su bc ili at um 1 1 + + g om ph re na c el os io id es 1 + 2 1 r hy nc ho si a m in im a va r. m in im a + + 3 + h yp ar rh en ia fi lip en du la 1 1 3 1 r ic ha rd ia s ca br a 4 + 1 + o xa lis o bl iq ui fo lia + 1 + c yp er us k ir ki i + 1 tr ag us b er te ro ni an us 1 1 p el la ea c al om el an os v ar . c al om el an os + + d ip ca di v ir id e 1 1 m un du le a se ri ce a + + tr oc ho m er ia m ac ro ca rp a su bs p. m ac ro ca rp a 1 1 h yp ox is v ill os a + + d ig ita ri a lo ng ifl or a 1 + c om m ip ho ra m ol lis + + h ib is cu s sc hi nz ii 1 + s ch ot ia b ra ch yp et al a 1 + p av et ta s ch um an ni an a + + a ri st id a ad sc en si on is + + p yr en ac an th a ka ur ab as sa na + + li pp ia s p. 2 1 a ca ci a ro bu st a su bs p. c la vi ge ra + 6 m ys tr ox yl on a et hi op ic um + + c ro ta la ri a la bu rn ifo lia s ub sp . a us tr al is 1 1 g re w ia c af fr a + + c ra ss ul a la nc eo la ta s ub sp . t ra ns va al en si s 1 te ph ro si a lu pi ni fo lia 1 a es ch yn om en e m ic ra nt ha 2 p up al ia la pp ac ea v ar . l ap pa ce a 1 h yp tis p ec tin at a 2 g la di ol us d en si flo ru s 1 c er op eg ia r ac em os a su bs p. s et ife ra + t a b l e 1 (c o n t. ..) 192 original research cronje, panagos & reilly koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 184 201 t a b l e 1 (c o n t. ..) c o m m u n it ie s 1 2 3 v a r ia n t s 1a 1b 2a 2b r e l e v é n u m b e r 4 6 2 1 1 8 7 1 2 5 3 1 2 1 2 1 1 4 3 2 2 5 7 3 6 2 3 7 6 5 3 1 7 4 4 3 7 6 2 3 3 4 5 7 2 3 2 6 1 4 4 4 4 1 3 6 6 7 4 7 8 8 5 5 8 7 5 5 5 6 8 7 6 5 6 8 8 8 8 5 3 4 8 7 4 7 1 2 8 3 3 9 3 6 8 6 2 4 0 4 2 5 6 5 0 1 8 1 7 3 6 7 2 0 8 1 2 3 6 7 4 9 9 0 9 9 2 5 7 5 1 5 2 9 0 2 8 4 7 5 1 4 3 8 6 0 1 6 3 0 8 5 1 9 4 7 6 0 2 3 1 9 4 8 7 5 6 d ac ty lo ct en iu m a us tr al e 1 d ac ty lo ct en iu m a eg yp tiu m 1 z or ni a gl oc hi di at a 1 tr ib ul us te rr es tr is 1 tr id ax p ro cu m be ns + te ph ro si a fo rb es ii su bs p. in te ri or + o zo ro a sp ha er oc ar pa + g is ek ia p ha rn ac io id es 2 tr ic hi lia e m et ic a + d io sp yr os ly ci oi de s su bs p. g ue rk ei + a lb iz ia v er si co lo r + s ac ci ol ep is c ur va ta 2 v er no ni a fa st ig ia ta + d ic om a te m en to sa 1 s pe ci es d (u ni de nt ifi ed ) 2 m on op si s de ci pi en s a c ro ta la ri a sp ha er oc ar pa s ub sp . s ph ae ro ca rp a 2 la nn ea d is co lo r + s pe ci es b (u ni de nt ifi ed ) + o rn ith og al um te nu ifo liu m s ub sp . t en ui fo liu m 1 s et ar ia in cr as sa ta 3 ip om oe a co pt ic a va r. co pt ic a + a sp ar ag us c oo pe ri + b ot hr io ch lo a bl ad hi i 1 a ca ci a si eb er ia na v ar . w oo di i + in di go fe ra ly de nb ur ge ns is 3 m on so ni a gl au ca 1 in di go fe ra d re ge an a + s pe ci es c (u ni de nt ifi ed ) 1 e ra gr os tis b ar bi no di s 1 s ch m id tia p ap po ph or oi de s 6 lu dw ig a oc to va lv is s ub sp . o ct ov al vi s + h el io tr op iu m z ey la ni cu m + p av on ia b ur ch el lii + v er no ni a gl ab ra v ar . l ax a 4 v er be ni a bo na ri en si s + s id a rh om bi fo lia 2 b au hi ni a ga lp in ii + d io sp yr os v ill os a va r. pa rv ifo lia 1 c is sa m pe lo s m uc ro na ta 4 a br us la ev ig at us 2 b ra ch ia ri a se rr at a 3 193 plant communities original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 184 201 t a b l e 1 (c o n t. ..) c o m m u n it ie s 1 2 3 v a r ia n t s 1a 1b 2a 2b r e l e v é n u m b e r 4 6 2 1 1 8 7 1 2 5 3 1 2 1 2 1 1 4 3 2 2 5 7 3 6 2 3 7 6 5 3 1 7 4 4 3 7 6 2 3 3 4 5 7 2 3 2 6 1 4 4 4 4 1 3 6 6 7 4 7 8 8 5 5 8 7 5 5 5 6 8 7 6 5 6 8 8 8 8 5 3 4 8 7 4 7 1 2 8 3 3 9 3 6 8 6 2 4 0 4 2 5 6 5 0 1 8 1 7 3 6 7 2 0 8 1 2 3 6 7 4 9 9 0 9 9 2 5 7 5 1 5 2 9 0 2 8 4 7 5 1 4 3 8 6 0 1 6 3 0 8 5 1 9 4 7 6 0 2 3 1 9 4 8 7 5 6 d ic om a ze yh er i 1 x im en ia c af fr a + g er be ra p ilo se llo id es 1 h ib is cu s al tis si m us 2 b on at ea s p. 2 v er no ni a ad oe ns is 1 s ol an um n ig ru m + u re ly tr um a gr op yr oi de s 3 e ra gr os tis c ili ar is + s ol an um in ca nu m 1 c hl or op hy tu m s p. + two variants (table 1). the identified plant communities were as follows: 1. the  strychnos  madagascariensis  -  aristida  stipitata  var.  graciliflora short closed woodland 1a the strychnos  madagascariensis  -  indigofera  inhambanensis  variant 1b the strychnos  madagascariensis parinari  curatellifolia  variant 2. the acacia  gerrardii  -  setaria  sphacelata  var. sphacelata  short closed woodland 2a the acacia  gerrardii fuirena  pachyrrhiza  var.  pachyrrhiza  variant 2b the acacia gerrardii themeda triandra variant 3. the gymnosporia  senegalensis  -  phragmites  mauritianus riparian low thicket the species groups were arranged to highlight the environmental gradients associated with savanna sandveld ecosystems. the phytosociological classification consists of 12 species groups (a–l) of which 205 (61.7%) and 128 (38.4%) species constitute the diagnostic and non-diagnostic portions, respectively (table 1). a vegetation map of the agr providing the spatial occurrence of the plant communities is presented in fig. 1. description of plant communities the strychnos madagascariensis aristida stipitata var.  graciliflora short closed woodland (community 1) community 1 (fig. 1; table 1, species group a) occurs throughout the reserve, predominating on the crests and upper mid-slopes of the landscape with a slope ranging between 2˚ and 5 .̊ the soils found in these physiographic units are deep sandy soils (mean clay > 5%), with a mean depth of 1 m. soil forms associated with this plant community are hutton, clovelly and cartref (macvicar et al. 1991). this community contributes the most to the overall species richness of the agr, with a total of 258 plant species (table 2), of which the majority, 15.7% and 45.4%, of the species richness is within the grass and forb growth forms, respectively (table 3). the woody component of the community is dominated (in terms of cover) primarily by terminalia  sericea and dichrostachys  cinerea  with 12.2% and 8.2% cover, respectively (table 4). terminalia sericea forms dense fringes of varying degrees toward the lower portions of the mid-slopes. strychnos  madagascariensis is a weak competitor with a density of three trees per hectare (table 4). digitaria  eriantha, hyperthelia dissoluta, panicum maximum (mainly under tree canopies) and  perotis  patens are dominant (in terms of 0 10 20 30 40 50 1 1a 1b 2 2a 2b 3 60 communities and variants c an op y co ve r ( % ) forbs grasses shrubs dwarf shrubs trees figure 2 histogram indicating the percentage canopy cover for the growth forms recorded in the main plant communities and their associated variants in the andover game reserve 194 original research cronje, panagos & reilly koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 184 201 cover) in the herbaceous layer (table 4). aristida stipitata subsp.  graciflora occurs as a normal competitor (table 4). the forbs, agathisanthemum bojeri subsp. bojeri, chamaecrista absus, vernonia  steetziana, tagetes  minuta  and wahlenbergia  banksiana  occur as strong competitors (table 4). the actual cover contribution of each growth form in this community is shown in fig. 2, with a high total percentage cover of 67.8%. the strychnos madagascariensis indigofera inhambanensis variant 1a variant 1a occurs mainly in the eastern half of the reserve (fig. 1; table 1, species group b), where the predominant underlying geology is grey to pale brown, mediumto coursegrained quartz-feldspar-biotite gneiss with subordinate mafic to ultramafic xenoliths (walraven 1986). the species richness consists of 197 plant species (table 2), with the dwarf shrub (14.5%), grass (14.9%) and forb (49%) growth forms contributing the most to the species richness of this variant (table 3). dominant species (in terms of cover) include the trees dichrostachys cinerea, terminalia  sericea, the grasses perotis  patens, digitaria  eriantha, bulbostylis contexta, hyperthelia dissoluta, panicum maximum, and the forbs  tagetes  minuta, chamaecrista  absus, vernonia  steetziana and agathisanthemum  bojeri  subsp.  bojeri (table 5).  indigofera  inhambanensis  is within a normal competition range (table 5). the vegetation structure in terms of the actual percentage cover relative to each growth form is illustrated in fig. 2, with a total cover of 64.9% the strychnos madagascariensis - parinari curatellifolia variant 1b this variant (1b) of community 1 occurs within the western portion of the reserve (fig. 1; table 1, species group c). the underlying geology consists of makhutswi gneiss, which is light-grey, medium-grained biotite gneiss with coursegrained quartz-feldspar leucosomes (walraven 1986). the occurrences of frequent fires (annually) from the bordering rural communities may be an additional causative factor of this variant. the species richness of this variant is slightly higher (207 species) than that of variant 1a (table 2), with the grass (17.8%) and forb growth forms (43.9%) contributing the most to the variant’s species richness (table 3). dichrostachys  cinerea and terminalia  sericea remain strong competitors in the tree growth form (table 6). euclea  divinorum and phyllanthus  reticulatus are strong competitors in the shrub and dwarf shrub growth forms, respectively (table 6). fuirena  pachyrrhiza var.  pachyrrhiza, perotis patens, cynodon dactylon, hyperthelia dissoluta and panicum maximum are dominant grasses in terms of cover, of which the latter two are also strong competitors (table 6). strong forb competitors include bidens  bipinnata,  wahlenbergia  banksiana and agathisanthemum bojeri subsp. bojeri (table 6). the growth forms depicting community structure relative to their actual percentage canopy cover are shown in fig. 2, with a high community cover of 72.8%. the acacia gerrardii setaria sphacelata var. sphacelata short  closed woodland (community 2)  community 2 (fig. 1; table 1, species group d) occurs on the lower mid-slopes and footslopes of the agr, with a slope ranging between 5˚ and 10 .̊ the soils are sandy-loam (mean clay 5–10%) towards the lower mid-slopes, becoming more clayey (mean clay > 50%) on the footslopes. soils occurring in the plant community are: cartref, fernwood, katspruit, kroonstad, sepane, valsrivier and sterkspruit (macvicar et  al. 1991). the mean soil depth is 1 m along the lower mid-slopes, decreasing to 0.5 m along the footslopes. a total of 222 plant species (table 2) occur in the community, with the highest species richness being in the forb (39.7%) growth form (table 3). the dominant trees (in terms of cover) are aristida  stipitata  subsp. graciliflora, acacia  nilotica, combretum  hereroense, dichrostachys  cinerea  and the shrub euclea divinorum, of which the latter three species are also strong competitors (table 7). dominant grasses (in terms of cover) include eragrostis gummiflua, eragrostis curvula, digitaria  eriantha,  setaria  sphacelata  var.  sphacelata,  bulbostylis  contexta,  hyperthelia dissoluta and panicum maximum, with the four latter species also being strong competitors  (table 7).  although hyperthelia dissoluta occurs as a strong competitor, its dominance in terms of cover (3.9%) is substantially lower in contrast to its dominance in community 1 and its variants 1a and 1b (table 4, table 5 and table 6). noticeably strong forb competitors are  chaetacanthus  setiger and epaltes  gariepina  (table 7). the actual canopy cover contribution of each growth form within the plant community is shown in fig. 2, with the total percentage cover for the community being 65.6%. measurements communities and variants 1 1a 1b 2 2a 2b 3 total no. relevés. 53 33 20 31 12 19 4 total no. species. 258 197* 207* 222 147* 193* 118 total diagnostic species. 181 132* 139* 160 97* 128* 72 diagnostic proportion. 70.2% 67.1% 67.2% 72.1% 66% 66.3% 61% species richness in terms of mean species per relevé. 56 57 54 56 49 61 53 community variation minimum species per relevé. 37 37 40 33 33 47 41 maximum species per relevé. 70 70 66 73 61 73 75 range of species per relevé. 33 33 26 40 28 26 34 mean species per relevé. 56 57 54 56 49 61 53 standard error of the mean of species per relevé. 1.01 1.3 1.5 1.7 2.4 1.5 7.6 community variation (proportion spp./relevé/ community). 21.6% 28.9% 25.9% 25.4% 33.4% 31.6% 44.9% community variation in terms of the standard deviation. 2.9% 3.8% 3.2% 4.2% 5.5% 3.4% 12.8% * several of the same species occurring in the main plant communities also occur in the variants and thus the species totals of the variants will not equate to the sum of the species in the main plant communities growth form communities and variants 1 1a 1b 2 2a 2b 3 no % no % no % no % no % no % no % trees 42 13.0 30 12.5 30 11.9 35 12.2 21 12.0 32 12.8 17 13.1 shrubs 39 12.0 22 9.1 33 13.0 42 14.6 18 10.3 37 14.8 23 17.7 dwarf shrubs 45 13.9 35 14.5 34 13.4 46 16.0 27 15.4 38 15.2 9 6.9 grass 51 15.7 36 14.9 45 17.8 50 17.4 34 19.4 45 18.0 23 17.7 forbs 147 45.4 118 49.0 111 43.9 114 39.7 75 42.9 98 39.2 58 44.6 table 2 community and variant statistics for the andover game reserve table 3 species richness within the communities and their associated variants according to each growth form * it is possible for more than one species to occurr in one or more growth forms in the woody plant categories 195 plant communities original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 184 201 cyperus  denudatus  and  cyperus  sphaerospermus  occurring  as normal competitors in this hydromorphic vegetation unit (table 8). species growth relations within this  variant are presented in  table 8.  hyperthelia  dissoluta  remains a strong grass competitor (table 8), primarily occurring towards the upper regions of this lower mid-slope vegetation unit.  epaltes  gariepina  is a strong forb competitor indicative of wet or seepline areas (table 8).  dichrostachys  cinerea  remains  a strong competitor  within the woody component of this vegetation unit (table 8). the total percentage cover for the variant is 71.1%, with proportions within the woody, herbaceous and forb components being 15%, 39.6% and 15.7% (fig. 2), respectively, clearly demonstrating the hydromorphic characteristic of the species growth form* comp. status** canopy cover (%) crown diameter (m) ind./ ha*** canopy/ canopy gap (m) terminalia sericea t strong 11.3 3.3 135 6.4 dichrostachys cinerea t strong 7.6 3.8 67 9.9 strychnos madagascariensis t weak 0.1 2.2 3 62.2 hyperthelia dissoluta g strong 7.1 0.3 14 960 0.7 panicum maximum g strong 6.8 0.2 25 241 0.5 digitaria eriantha g normal 2.1 0.1 51 377 0.4 perotis patens g normal 2.9 0.2 17 282 0.7 agathisanthemum bojeri subsp. bojeri f strong 1.3 0.3 179 2.5 chamaecrista absus f strong 2.2 0.2 9 145 1.0 vernonia steetziana f strong 4.5 0.2 22 850 0.6 indigofera inhambanensis f normal 0.2 0.3 334 5.8 tagetes minuta f strong 1.3 0.1 9 229 1.0 * t = tree; g = grass; f = forb ** competitor status *** individuals per hectre table 5 species and growth form relations in variant 1a species growth form* comp. status** canopy cover (%) crown diameter (m) ind./ ha*** canopy/ canopy gap (m) terminalia sericea t strong 13.5 3.0 178 5.3 dichrostachys cinerea t strong 8.8 2.9 128 7.0 strychnos madagascariensis t weak 0.1 1.6 3 60.0 euclea divinorum s strong 1.0 1.9 36 16.7 phyllanthus reticulatus ds strong 0.02 0.4 19 25.0 perotis patens g weak 1.4 0.1 12 108 0.9 hyperthelia dissoluta g strong 10.0 0.3 16 578 0.6 panicum maximum g strong 10.2 0.2 32 380 0.4 cynodon dactylon g normal 1.9 0.1 115 500 0.3 fuirena pachyrrhiza var. pachyrrhiza g normal 1.9 0.2 7 614 1.1 bidens bipinnata f strong 1.1 0.1 7 114 1.2 wahlenbergia banksiana f strong 1.9 0.1 20129 0.7 agathisanthemum bojeri subsp. bojeri f strong 1.4 0.3 2 228 2.1 * t = tree; ds = dwarf shrub; g = grass; f = forb ** competitor status *** individuals per hectre table 6 species and growth form relations in variant 1b the acacia gerrardii - fuirena pachyrrhiza var. pachyrrhiza variant 2a this variant (fig. 1 table 1, species group e) is indicative of the seeplines and the transition areas between the broadand fineleaved savanna. soils occurring in this vegetation unit have a characteristic g-horizon, namely katspruit and kroonstad (macvicar et  al. 1991). as a result of the g-horizon, seasonal water emerges to the surface forming wet-spots or seeplines, with characteristic hydromorphic species (i.e. within the grass and forb growth forms) dominating these areas. the floristic richness consists of 147 individual species (table 2) with the grass and forb layers contributing 19.4% and 42.9% (table 3), respectively, to the species richness of this variant. diagnostic species are fuirena  pachyrrhiza  var.  pachyrrhiza and  vahlia  capensis subsp. vulgaris var. vulgaris (table 1, species group e), with eragrostis inamoena, eragrostis gummiflua, eragrostis curvula, species growth form* comp. status** canopy cover (%) crown diameter (m) ind./ ha*** canopy/ canopy gap (m) acacia gerrardii t normal 2.9 3.50 29 17.3 acacia nilotica t normal 2.7 4.60 27.9 23.3 acacia swazica t normal 1.2 1.40 76 11.5 combretum hereroense t strong 3.9 2.70 67 11.0 dichrostachys cinerea t strong 6.5 3.30 76 9.6 euclea divinorum s strong 1.5 2.00 36 16.7 panicum maximum g strong 8.2 0.20 29 199 0.5 hyperthelia dissoluta g strong 3.9 0.30 7629 1.0 bulbostylis contexta g strong 3.5 0.10 179 581 0.2 setaria sphacelata var. sphacelata g strong 3.1 0.24 7 207 1.1 eragrostis gummiflua g normal 1.8 0.30 3703 1.6 eragrostis curvula g normal 1.9 0.10 15871 0.8 digitaria eriantha g normal 1.4 0.10 20113 0.7 chaetacanthus setiger f strong 1.3 0.10 2 289 2.1 epaltes gariepina f strong 5.5 0.30 55 418 0.4 table 7 species and growth form relations in community 2 species growth form* comp. status** canopy cover (%) crown diameter (m) ind./ ha*** canopy/ canopy gap (m) terminalia sericea t strong 12.2 3.2 150 6.0 dichrostachys cinerea t strong 8.2 3.5 85 8.7 strychnos madagascariensis t weak 0.1 1.9 3 60.3 hyperthelia dissoluta g strong 8.2 0.3 15 525 0.7 panicum maximum g strong 8.1 0.2 28 278 0.5 digitaria eriantha g weak 1.7 0.1 39 612 0.8 perotis patens g weak 2.3 0.1 15 872 0.8 aristida stipitata subsp. graciliflora g normal 0.1 0.1 4 334 1.7 agathisanthemum bojeri subsp. bojeri f strong 1.3 0.3 1 807 2.4 chamaecrista absus f strong 1.6 0.2 7 834 1.1 vernonia steetziana f strong 3.0 0.2 17 146 0.7 tagetes minuta f strong 1.0 0.1 7 441 1.2 wahlanbergia banksiana f strong 0.7 0.1 9 345 1.1 hibiscus sp. f strong 1.0 0.1 6 715 1.2 * t = tree; g = grass; f = forb ** competitor status *** individuals per hectre table 4 species and growth form relations in community 1 196 * t = tree; ds = dwarf shrub; g = grass; f = forb ** competitor status *** individuals per hectre original research cronje, panagos & reilly koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 184 201 seepline with a clear reduction in the occurrence of the woody components in these areas. the acacia gerrardii themeda triandra variant 2b this variant (fig. 1; table 1, species group f) is restricted to the footslopes of the study area and follows the drainage lines originating in the south and flowing in a northerly direction. the variant is also located along the drainage lines in the eastern parts of the reserve. soil forms in this vegetation unit are predominantly sterkspruit, sepane and valsrivier, which are duplex soils (macvicar et al. 1991). duplex soils are characterised by an abrupt textural or structural transition between the surface horizon and the subsoil (matthee 1996). the glenrosa soil form (macvicar et  al. 1991) also occurs where the parent granite rock lies shallow below the surface. the species richness in this variant amounts to 193 individual species (table 2), with the forbs contributing the most (39.2%) to this vegetation unit (table 3). dominant species (in terms of cover) include the trees dichrostachys cinerea, combretum hereroense, acacia nilotica,  acacia gerrardii, the shrub euclea divinorum, the grasses panicum  maximum, setaria sphacelata var. sphacelata, digitaria eriantha, and the forbs chaetacanthus setiger and epaltes gariepina (table 9). the total canopy cover for this variant is 62.1% (fig. 2). the gymnosporia senegalensis phragmites mauritianus  riparian low thicket (community 3) this plant community (fig. 1; table 1, species group j) is found along the seasonal rivers occurring in the northern and north-western corner of the reserve. alluvial soils are the predominant soils in the community. this community has the lowest species richness with a total of 118 species (table 2), with the shrub (17.7%), grass (17.7%) and forb (44.6%) growth forms contributing the most to the species richness of this vegetation unit (table 3). combretum  imberbe  and  gymnosporia  senegalensis  are strong woody competitors (table 10). dominant herbaceous species (in terms of cover) include eriochloa  meyeriana  subsp.  meyeriana,  eustachys  paspaloides,  phragmites  mauritianus  and  panicum  maximum,  of which  phragmites  mauritianus  is a strong competitor (table 10). hypoestes  forskaolii  ‘form b’  is a strong forb competitor (table 10). the community structure expressed as a percentage actual cover contributed by each growth form is illustrated in figure 2. the total class cover is 99.4% and is indicative of dense river vegetation, particularly within the grass (48.9%) and forb (21%) growth forms (fig. 2). the woody proportion constitutes 18.4% of the total class cover (fig. 2) species growth form* comp. status** canopy cover (%) crown diameter (m) ind./ ha*** canopy/ canopy gap (m) dichrostachys cinerea t strong 9.5 3.4 105 7.6 bulbostylis contexta g strong 7.9 0.1 232 603 0.2 hyperthelia dissoluta g strong 10.0 0.3 19 710 0.6 panicum maximum g strong 6.2 0.2 19 737 0.6 fuirena pachyrrhiza var. pachyrrhiza g normal 0.5 0.3 593 4.6 eragrostis inamoena g normal 1.1 0.2 2539 2.0 cyperus denudatus g normal 1.1 0.1 10 141 1.0 cyperus sphaerospermus g normal 1.7 0.1 27 508 0.6 eragrostis gummiflua g normal 2.0 0.2 5 021 1.4 eragrostis curvula g normal 2.0 0.1 18 321 0.7 vahlia capensis subsp. vulgaris var. vulgaris f normal 0.5 0.2 8 654 1.1 epaltes gariepina f strong 7.3 0.1 61 276 0.3 * t = tree; g = grass; f = forb ** competitor status *** individuals per hectre table 8 species and growth form relations in variant 2a species growth form* comp. status** canopy cover (%) crown diameter (m) ind./ ha*** canopy/ canopy gap (m) dichrostachys cinerea t strong 4.6 3.2 57 11.80 combretum hereroense t strong 4.9 2.7 88 9.30 acacia nilotica t strong 4.2 4.7 24 18.20 acacia gerrardii t strong 4.4 3.6 42 13.70 euclea divinorum s strong 2.4 2.0 53 13.50 panicum maximum g strong 9.6 0.2 35 989 0.41 setaria sphacelata var. sphacelata g strong 4.3 0.2 9 094 0.90 digitaria eriantha g normal 2.0 0.1 25 719 0.60 epaltes gariepina f strong 4.3 0.1 48 453 0.40 chaetacanthus setiger f strong 2.1 0.3 3 434 1.70 table 9 species and growth form relations in variant 2b species growth form* comp. status** canopy cover (%) crown diameter (m) ind./ ha*** canopy/ canopy gap (m) combretum imberbe t strong 13.5 4.1 101 7.1 spirostachys africana t normal 1.3 5.1 6 40.6 gymnosporia senegalensis s strong 10.3 3.1 133 6.6 flueggea virosa ds strong 0.9 2.2 22 21.8 phragmites mauritianus g strong 22.2 0.5 11 640 0.6 eriochloa meyeriana subsp. meyeriana g normal 5.3 0.2 13 199 0.8 eustachys paspaloides g normal 10.3 0.3 20 282 0.5 hypoestes forskaolii ‘form b’ f strong 14.3 0.2 349 798 0.4 table 10 species and growth form relations in community 3 axes 1 2 3 4 total inertia eigenvalues 0.86 0.59 0.43 0.29 9.72 lengths of gradient (sd’s) 4.77* 4.38* 3.95 3.23 cumulative percentage variance of species data 8.90 15.00 19.4 22.4 * significant gradients table 11 statistical significance of a detrended correspondence analysis presenting gradient lengths on four axes. gradient lengths with standard deviations (sds) of four or greater have statistical significance (ter braak & šmilauer 1998) environmental variables variable no. lambda-a p-value f-value soil 3 0.57 0.001 5.38 slope 2 0.54 0.001 5.29 physiographical units 1 0.20 0.001 2.05 seeplines 4 0.21 0.001 2.07 table 12 influence of environmental variables on the community variation on the agr computed by a canonical correspondence analysis monte carlo test. (full model. p-values ≤ 0.05 are significant (ter braak & šmilauer 1998)) 197 * t = tree; ds = dwarf shrub; g = grass; f = forb ** competitor status *** individuals per hectre * t = tree; ds = dwarf shrub; g = grass; f = forb ** competitor status *** individuals per hectre plant communities original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 184 201 figure 3 detrended correspondence analysis (ter braak & šmilauer 1998) ordination of the complete data set of the andover game reserve with a standard deviation (sd) of 4.77. relevés highlighted in red represent communities 1 and 2 and relevés highlighted in blue represent community 3 figure 4 detrended correspondence analysis (ter braak & šmilauer 1998) ordination of the data set of the andover game reserve with a sd of 4.75 depicting the terrestrial communities (i.e. relevés representing community 3 removed). relevés highlighted in blue and red represent communities 1 and 2. relevés highlighted in black and green represent variant 2a and outliers, respectively 198 original research cronje, panagos & reilly koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 184 201 and the high percentage grass cover found in this riparian community is due to the inclusion of the reed phragmites  mauritianus in the grass growth form, contributing 22.2% cover on its own (table 10). verification of the plant communities a dca of the agr data set using canoco (ter braak & šmilauer 1998) provided a significant standard deviation gradient of 4.7 on the first axis with further significant and modest gradients on axes 2, 3, and 4 (table 11). the cluster diagrams generated by canoco (fig. 3 & fig. 4) reflect similar groupings to the relevé groupings generated by the phytotab-pc (westfall 1992) classification programme (table 1). a separation is evident between community 3 (relevés highlighted in blue) and the terrestrial communities (1 & 2 plus their variants) (relevés highlighted in red) (fig. 3). excluding the riparian community (3) from the data set and performing another secondary ordination expands the terrestrial communities with relevés highlighted in blue, black and red, representing community 1, variant 2a and community 2, respectively (fig. 4). outliers in terms of seepline (variant 2a) relevés are highlighted in green. the lack of clustering of the seepline relevés (fig. 4) can be attributed to the catenal phenomenon in terms of the variable width of the seeplines and their similarities to the adjacent plant communities with regard to being transitional between the upper and lower plant communities. of the environmental factors recorded, the factor most likely to have had the greatest influence computed by a cca (table 12) on the distribution of plant communities within the agr was soil in terms of moisture and clay content. discussion the agr’s undulating granitic physiography resulted in the identification of a catenal sequence similar to that described by scholes (1986, 1997). the plant communities of the agr are thus a function of the catenal sequence in the granite lowevld bushveld (cronje 2004; driver et  al. 2005), reflecting a mosaic of units having a ‘tortoise shell’ appearance. although plant species composition is influenced by such soil properties as nutrient status, ph, salinity and texture, the overwhelming factor determining the spatial distribution and productivity of forest, savanna and grassland is soil moisture balance (tinley 1982; walker 1985). at a scale of hundreds of meters to kilometres, the migration of fine soil particles and ions from ridge crests to valley floors, under the influence of water movement and gravity, establishes a topo-sequence of soils and associated vegetation known as a catena (scholes 1997). the availability of this soil moisture to plants is dictated by the soil type, in terms of clay content. fine-textured soils along the foot-slopes (high clay) are more xeric and thus water in these soils is limiting for much of the year. under the same climatic conditions in sandy soils (i.e. along the crests and upper midslopes), moisture is much less of a limiting factor (knoop & walker 1985) although it is quickly lost to the bottomlands. the sandy soils generally have an acidic ph, with alkaline soils on the footslopes (bredenkamp 1982; scholes 1986). therefore the differences in soil constituency reflect plant productivity, vegetation structure and species composition (o’ connor 1985). floristic richness is also influenced by climate and management factors such as grazing and fire (o’connor 1991; o’connor 1995; scholes 1997; teague & smit 1992). the vegetation of the agr relative to the catenal sequence described by scholes (1997) has the broad-leaved savanna (community 1) occurring on the crests and upper mid-slopes with sandy soils and the fine-leaved savanna (community 2) on lower mid-slopes and footslopes with clay soils. another characteristic feature of the catenal sequence is the formation of seeplines (i.e. an intermittent perched water table) that have characteristic hydromorphic or dambo grassland species (scholes 1986; 1997) and is represented on the agr by the community variant 2a. the formation of this hydromorphic grassland is as a result of the interface between the sandy upland and the clayey bottomland (scholes 1997). the influence of the interface of the sand and clay is not necessarily limited to a specific area but can expand from the moist seepline areas into the broad-leaved savanna particularly with regard to hydromorphic sedges and forbs, which are clearly noticeable in the phytosociological table (table 1), as indicated by the bold broken line between variants 2a and 1b (species group e). a possible reason for this could be a damming-up effect causing moisture to move upwards from the seepline, resulting in a higher moisture content in the lower section of variant 1b and thus creating hydromorphic conditions similar to that of a seepline. water moving downslope towards the evident interface would then result in a higher soil moisture content up to the seepline, where the water emerges, creating a hydromorphic grassland. variant 2a (acacia gerrardii fuirena  pachyrrhiza  var.  pachyrrhiza) is clearly recognisable by the predominance of sedges and hydromorphic grasses and the sparseness of trees, as reflected by the proportional cover values of the woody, grass and forb growth forms being 15%, 39.6% and 15.7%, respectively (fig. 2). this hydromorphic community is more moist than the areas above and below it and therefore remains green and palatable for longer periods of time, thus making it a valuable habitat for wildlife, especially during the dry winter season (scholes 1986). floristic and habitat correlation floristic and habitat correlations not only aid in the verification of the vegetation classification but also provide an understanding of the causes of community differentiation (westfall 1992). finding a correlation between habitat and plant communities does not necessarily validate a classification, because with all the habitat factors available it should be possible to find some or other correlation with a specific group of plants. what is important however is that habitat factors shown to correlate with the plant communities should form some sort of environmental gradient (panagos et  al. 1998). the initial canoco dca (ter braak & šmilauer 1998) ordination of all relevés provided a significant gradient (> 4 sd, table 11). scatter diagrams (fig. 3 & fig. 4) reflect relevé groupings similar to that produced by phytotab-pc (westfall 1992). a cca (ter braak & šmilauer 1998) conducted on the environmental factors (table 12) recorded in this study indicated soil texture (i.e. sandy, sandy loam, loam and clay), and hence soil moisture, to be the environmental factor that had the greatest influence on the plant community differentiation. the relevé grouping produced both by phytotab-pc (westfall 1992) and canoco (ter braak & šmilauer 1998) reflects a gradient relating to soil fertility and moisture clearly evident on catenas of granitic origin. thus the dca and the cca supported and confirmed the vegetation classification of the agr obtained in this study as analysed by phytotab-pc (westfall 1992). conclusions with the exception of the riverine work of myburgh (1999), this is the first non-riverine application of a scale-related vegetation technique (westfall et  al. 1996) used in conjunction with phytotab-pc (westfall 1992) in the arid or granitic lowveld (acocks 1988; mucina & rutherford 2006). the stratification and classification of the vegetation of the agr at a scale of 1:13 000 has provided a map of the reserve with three main plant communities, two of which have two variants. the plant communities of the agr are indicative of vegetation on deep sandy soils of the granite lowveld bushveld veld type (driver et al. 2005) with affinity to the catenal phenomenon. the 199 plant communities original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 184 201 use of a multivariate data analysis technique on the agr data set confirmed the phytosociological classification of agr. the use of the scale-related vegetation survey technique (westfall et al. 1996; westfall & panagos 1988) has provided quantitative data together with a vegetation map of the agr, which can be used to formulate a management plan for the andover game reserve. an integral component of reserve management is the implementation of a vegetation monitoring programme for the andover game reserve. the data collected in this study was objective with regard to the systematic manner (cronje 2004) in which it was collected with the use of the plant number scale (westfall & panagos 1988). it is recommended that an area-based monitoring technique that incorporates the data collected in this survey be used so that this baseline data can be expanded upon to benefit management decision support of the andover game reserve. acknowledgements the authors thank mr a.j. botha and mr i.c. sharp for logistical support and valued comments; the tshwane university of technology for funding; dr w.j. myburgh, mr j.j. kotze and dr r.h. westfall for assistance with data analysis; and mrs i. cronje for dedicated assistance and motivation. references acocks, j.p.h. 1988. veld types of south africa. memoirs of the  botanical survey of south africa, 57: 1-146 agromet database. 1996. pretoria: agricultural research council, institute for soil, climate and water. barrett, a.s., brown, l.r., barrett, l. &. henzi, s.p. 2006. phytosociology and the plant community utilization by vervet monkeys of the blydeberg conservancy, limpopo province. koedoe, 49(1): 49–68. boucher, c. & jarman, m.l. 1977. the vegetation of the langebaan area, south africa. transactions  of  the  royal  society of south africa, 42: 241–272. brady, n.c. 1984. the  nature  and  properties  of  soils (9th ed.). london: collins macmillan publishers. bredenkamp, g.j. 1975. plantsosiologiese  studie  van  die  suikerbosrand  natuurreservaaat. msc thesis, pretoria: university of pretoria. bredenkamp, g.j. 1982. ‘n plantekologiese studie van die manyeletiwildtuin. dsc thesis, pretoria: university of pretoria. bredenkamp, g.j. & brown, l.r. 2001. a reliable ecological basis for environmental planning. urban  green  file, november/ december 2001: 38–39. brown, l.r. & h. bezuidenhout, h.b. 2000. the phytosociology of the de rust section of the mountain zebra national park, eastern cape. koedoe, 43(1): 1–18. brown, l. r., marais, h. , henzi, s.p. & barrett, l. 2005. vegetation classification as the basis for baboon management in the bourke’s luck section of the blyde canyon nature reserve, mpumalanga. koedoe, 48(2): 71–92. coombes, p.j. & mentis, m.t. 1992. a procedure for defining conservation management objectives and goals. in:  ebedes, h (ed.). proceedings  of  the  3rd  international  wildlife  ranching symposium. pretoria. cronje, h.p. 2004. rangeland  management  implications  of  a  scalerelated  vegetation  survey  in  the  arid  lowveld,  south  africa. mtech thesis. pretoria: tshwane university of technology. driver, a., maze, k., rouget, m., lombard, a.t., nel, j., turpie, j.k., cowling, r.m., desmet, p., goodman, p., harris, j., jonas, z. reyers, b. sink, k. & strauss, t. 2005. national spatial biodiversity assessment 2004: priorities for biodiversity conservation in south africa. strelitzia, 17. pretoria: south african national biodiversity institute. edwards, d. 1983. a broad-scale structural classification of vegetation for practical purposes. bothalia, 14: 705–712. germishuizen, g. & meyer, n.l. (eds.). 2003. the plants of southern africa: an annotated checklist. strelitzia, 14. pretoria: south african national biodiversity institute. knoop, w.t. & b.h. walker, b.h. 1985. interactions of woody and herbaceous vegetation in a southern african savanna. journal of ecology, 73: 235–253. kruger, a.c. 2004. climate of south africa. climate region. ws45. pretoria: south african weather service. macvicar, c.n., bennie, a.t.p., villiers, j.m., ellis, j., fey, m.v., harmse, h.j., von, m., hensley, m., lambrechts, j.j.n., bruce, r.w., dohse, t.e., eloff, j.f., grey, d.c., hartmann, m.o., idema, s.w.j., laker, m.c., merryweather, f.r., michael, d., schloms, b.h.a., schönau, a.p.g., snyman, k., van niekerk, b.j., verster, e., loxon, r.f., meyer, j.h., paterson, d.g., schoeman, j.l., scotney, d.m., turner, d.p., van rooyen, t.h. & yager. t.u. 1991. soil classification: a taxonomic system for south africa. memoirs on the agricultural natural  resources of south africa, no.15: 1–257. matthee, j.f.g. 1996. a  primer  on  soil  conservation. pretoria: government printer. pp. 1–124. mucina, l. & rutherford, m.c. (eds.). 2006. the vegetation of south africa, lesotho and swaziland. strelitzia, 19. pretoria: south african national biodiversity institute. myburgh, w.j. 1999. oewerplantegroei van die olifantsriviersisteem  –  ‘n  ekologiese  perspektief. pretoria: water research commission, report no. 663/1/99. isbn 1868455688. o’ connor, t.g. 1985. a synthesis of field experiments concerning the grass layer in the savanna regions of southern africa. south african national scientific programmes report, 114: 1–26. o’connor, t.g. 1991. influence of rainfall and grazing on the compositional change of herbaceous layer of a sandveld savanna. journal  of  the  grassland  society  of  southern  africa,  8(3): 103–109. o’connor, t.g. 1995. transformation of savanna grassland by drought and grazing. african  journal  of  range  and  forage  science, 12(2): 53–60. panagos, m.d. 1995. a comparative classification of the sourishmixed bushveld on the farm roodeplaat (293) using quadrat and  point  methods. msc thesis. petermaritzburg: university of natal. panagos, m.d. & reilly, b.k. 2006. use of an area-based survey technique to detect vegetation changes in sour bushveld. koedoe, 49(1): 69–77. panagos, m.d., westfall, r.h., van staden, j.m. & zacharias, p.j.k. 1998. the plant communities of the roodelpaat experimental farm, gauteng, south africa and the importance of classification verification. south  african  journal of botany, 64(1): 44–46. pressey, r.l. & bedward, m. 1991. mapping the environment at different scales: benefits and costs for nature conservation. in: margules, c.r. & austin, m.p. (eds.) nature conservation:  cost  effective  biological  surveys  and  data  analysis.  canberra: csiro. pyle, r.m. 1980. management in nature reserves. in: soulé, m.e. & wilcox, b.a. (eds.). conservation  biology,  an  evolutionary  ecological  perspective. sunderland: massachusett: sinauer. pp. 319–328. reilly, b.k. & macfadyen, i.d. 1992. an eco-systematic procedural model for the ecological management of letaba ranch game reserve, gazankulu. in: ebedes, h (ed). proceedings of the 3rd  international wildlife ranching symposium. pretoria. rutherford, m.c. & westfall, r.h. 1986. biomes of southern africa: an objective categorisation. memoirs of the botanical  survey of south africa, 54: 1–98. scholes, r.j. 1986. a  guide  to  bush  clearing  in  the  eastern  transvaal  lowveld, 2nd ed. johannesburg: university of the witwatersrand. scholes, r.j. 1997. savanna. in: cowling, r.m., richardsom, d. m. & pierce, s.m. eds. vegetation of southern africa. cambridge: cambridge university press. pp. 258–277. 200 original research cronje, panagos & reilly koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 184 201 schulze, b.r. 1994. climate  of  south  africa  (wb  28).  part  8.  general  survey. pretoria: weather bureau; department of environmental affairs and tourism. spinage, c.a.. 1979. what is wildlife management? in: ajaya, s.s. & halstead, l.b. (eds.). wildlife  management  in  savanna  woodland. london: taylor & francis, pp. 186–193. teague, w.r. & smit, g.n. 1992. relations between woody and herbaceous components and the effects of bush-clearing in southern african savannas. journal of the grassland society of  southern africa, 9(2): 60–71. ter braak, c.j.f. & p. šmilauer, p. 1998. canoco  reference  manual  and  user’s  guide  to  canoco  for  windows:  software  for  canonical  community  ordination  (version  4). ithaca: new york. microcomputer power. theron, g.k. 1973. ‘n ekologiese studie van die plantegroei van die  loskopdamnatuurreservaat. dsc thesis. pretoria: university of pretoria. tinley, k.l. 1982. the influence of soil moisture balance on the ecosystem patterns on southern africa. in: huntley, b.j. & walker, b.h. (eds). ecology of tropical savannas. berlin: springer-verlag. van rooyen, n. 1983. die plantegroei van die roodeplaatdamnatuurreservaat ii. die plantgemeenskappe. south  african  journal of botany, 2: 115–125. visser, n., van hoven, w. & theron, g.k. 1996. the vegetation and identification of management units of the honnet nature reserve, northern province, south africa. koedoe, 39(1): 25–42. walker, b.h. 1985. structure and function of savannas: an overview. in: tothill, j.h. & mott, t.t. (eds). ecology  and  management  of  the  worlds  savannas.  canberra: australian academy of science, pp. 350–353. walraven, f. 1986. 2430 pilgrim’s rest 1: 250 000 geological series. pretoria: government printer. walraven, f. 1989. the geology of the pilgrim’s rest area. pretoria: government printer. werger, m.j.a. 1974. on concepts and techniques applied in the zürich-montpellier method of vegetation survey. bothalia, 11: 309–323. westfall, r.h. 1992. objectivity in stratification, sampling and classification of vegetation. phd thesis. pretoria: university of pretoria. westfall, r.h. & panagos, m. d. 1988. the plant number scale – an improved method of cover estimation using variablesized belt transects. bothalia, 18(2): 289–291. westfall, r.h., theron, g.k. & van rooyen, n. 1997. objective classification and analysis of vegetation data. plant ecology, 132: 137–154. westfall, r.h., van staden, j.m., panagos, m.d., breytenbach, p.j.j. & greef, a. 1996. scale-related vegetation sampling. arc – range and forage institute. private bag x05, lynn east 0039. 201 filelist convert a pdf file! supplement to koedoe. 1977: 106-115. present day status and future of nature conservation in the republic of malawi o g anstey and a j hall-martin department of national parks and wildlife ulongwe and wildlife research unit kasungu malawi introduction when malawi (formerly nyasaland) became independent in 1964 the status and future of its game reserves were in jeopardy. the former administration had adopted a policy of benign neglect towards the country's wildlife areas (anon 1963), and the remnants of the department of game, fish and tsetse control had been absorbed by the forestry department in 1963. fortunately the life president of malawi, dr h kamuzu banda, took a strong interest in wildlife conservation and it was only his personal intervention, and the advent of independence, that saved the former lengwe game reserve from deproclamation (hayes 1967) as planned by the colonial administration. with the life president's encouragement and the dedicated efforts of the staff responsible for wildlife, the tide which had been running strongly against nature conservation wa.s turned, culminating in the establishment of a separate department of national parks and wildlife only a decade after independence. physiography and climate the republic of malawi (rm) lies in south-east central africa between 9°25' and 17°08' sand 33° and 36°e. its total area is 121 600 km2 which includes 28 416 km2 of surface water (lakes malawi, chilwa and malombe). the country has borders with mocambique, tanzania and zambia (fig. 1) and is totally landlocked. the great rift valley in which lakes malawi and malombe lie traverses the country from north, to south. these lakes are drained by the shire river which follows the rift valley to the zambesi river. the rift valley plains lie at altitudes of 100-450 m and are separated from the central african miocene peneplain surface by the escarpments which form the walls of the rift valley. the peneplain surface lies at an altitude of 600-1 500 m and forms the greater part of the country. isolated granite inselbergs and mountains rise above the plain to altitudes of3 001 m (mlanje), 2 087 m 106 10 12 14 16 zambia mocambique scale o 35 70 105 140 175 km -l[mlrft\~ game reserve _ national pa.rk ....... ... proposed extension tanzania ~r~i-.-. . -.-.-.mocambique • lilongwe fig. 1. republic of malawi showing the distribution of national parks and game reserves. 107 (zomba) and 2260 m (dedza) while in the north of the country the high montane plateaux of the nyika (2606 m), vipya (2 082 m) and misuku (1 859 m) cover large areas. these montane areas are believed to be the remnants of the late jurassic peneplain (chapman 1968). these three major physiographic regions exert a major influence on the vegetation types. the climate of the rm is partly continental in character (griffiths 1972) but the physiographic regions and the proximity of the indian ocean have a great effect on rainfall and temperature and there is considerable local variation. the rift valley areas are generally hot with maximum temperatures commonly reaching 45°c (hall-martin 1975), but with a mean annual temperature of 18°c-24°c, whilst the mountain plateaux have a mean annual temperature below 19°c with frequent fros ts during the winter mon ths ( chapman 1968). rainfall varies from 500-1 125 mm per annum over most of the country but is much higher where topographic features playa role, such as the northern shores of lake malawi and on mountain slopes which intercept rain bearing winds. on mount mlanje, for example, the rainfall reaches 2 000 millimeters. there is a clearly defined seasonal rhythm in rainfall and temperature resulting in a wet season from november to april, and a dry season from may to october. a feature of the dry season climate is the regular invasion of moist maritime air from the mocambique channel resulting in periods of cool, overcast and drizzling weather. vegetation the vegetation of the rm has been described by various authors (wild and barbosa 1967; brass 1953; chapman 1962, 1968; chapman and white 1970; jackson 1954; hall-martin 1972, 1975). as a result the distribution of the most important vegetation types is now known, as also the occurrence of many rare and endangered species. the greater part of the miocene surface is covered in dense to open miombo woodland composed largely of members of the broad-leaved decid uous genera brachystegia, julbernardia and isoberlinia. drainage lines in these woodlands are usually edaphic grasslands, known locally as "dambos". in the lower lying areas of the peneplain surface the miombo gives way to a deciduous tree savanna in which the principal woody plants are pterocarpus angolensis, combretum spp, pericopsis angolensis, terminalia sericea, burkea africana and ostryoderris stuhlmanii and several acacia species. on the escarpment areas and the upper rift valley plains dry deciduous tree savanna and woodland dominated by colophospermum mopane with a grass layer composed largely of the genera eragrostis, cenchrus, schmidtia and aristida occurs. the floor of the rift valley is covered by a deciduous tree savanna dominated by adansonia digitata, sterculia appendiculata, kirkia acuminata, cordyla africana and acacia nigrescens. the most important grasses are panicum maximum, urochloa mossam108 bicensis and digitaria milanjiana. in most areas the river banks and termite mounds support forest or thickets. areas along the lake shores and major river valleys support swamp grasslands; cyperus papyrus and phragmites mauritianus beds; and the palms hyphaene and borassus are common in wetter areas. the other major plant formations of the rm are those associated with the montane areas. broad-leaved evergreen forest is distributed in patches on all the higher inselbergs and mountains and consists of many types often dominated by ocotea usambarensis, chrysophyllum gorungosanum, strombosia scheffieri, drypetes gerrardii and aningeria adofji-jriederici. at high altitudes conifer forest is also found but is confined to the nyika plateau where the dominant species is juniperus procera and on mt mlanje where widdringtonia whytei is dominant and stands 50 m tall. montane grasslands dominated by exotheca abyssinica, loudetia simplex and monorymbium ceresiiforme are present on all high mountain plateaux. there are also many other vegetation types in malawi which are less widespread such as lowland evergreen forest, lowland dry deciduous forest and thicket and many lesser savanna and woodland types. human population the population of the rm in 1976 is estimated at 5 000 000, living at a density of67 per km2 with local densities of 309 per km 2 (anon 1963). the people are mostly agriculturalists but a transition to an industrialised society is taking place rapidly. nature conservation legislation national parks and game reserves in the rm are authorised and managed largely in compliance with three government acts. the land act authorizes acquisition of customary land and its declaration as public land which can then be gazetted as a national park. the game act (1954) prescribes a means for establishing game reserves and identifies activities which are prohibited in game reserves. the national park act (1969) provides the most comprehensive control order over activities within any area of the department of national parks and wildlife. a significant inclusion of the national park bill is that no national park may be reduced in area without a specific resolution of parliament. a new comprehensive flora and fauna act is in draft stage. this law will supercede the game act, and will provide the authority for stricter protection outside the parks. parks and reserve administration from 1963 to 1973, the national parks and game reserves were the responsibility of the game division of the department of forestry and 109 game. in april 1973, at the direction of the life president of malawi, a separate department of national parks and wildlife was established within the ministry of agriculture and natural resources. this department now manages four national parks, three established game reserves and one proposed game reserve. crop protection and control of other problem animals outside the parks and game reserves is also the responsibility of the department. the department of national parks and wildlife is headed by a principal warden who is responsible to the secretary for agriculture and natural resources. consistent with the three administrative divisions of the rm, three regional wardens supervise department activities in these regions. a senior warden is in charge of each of the four national parks. the conservation force presently consists of 17 rangers, 15 head scouts and 108 scouts distributed throughout the eight areas. there is also a crop protection unit within the department h eaded by a crop protection officer. this unit deals with marauding wild animals and has 20 supporting staff. a wildlife research unit is situated in the kasungu national park. a conservation sanctuary and a research and monitoring unit has been set up in the nyika national park. a conservation sanctuary and education centre has been set up on 100 ha of old forest res erve within the new lilongwe capital city. this will be the focus of conservation education activities of the department. three of the national parks (lengwe, kasungu and nyika) have overnight visitor facilities and are open to the public. the fourth park (liwonde), gazetted in 1972, is expected to be opened in 1977. the game reserves are being managed on a care and maintenance basis. however, their potential for eventual utilization is being investigated. while making provisions for visitor use in the national parks, the government's first concern is to perpetuate the parks and game reserves as biotic reserves. the department's management of each area reflects this emphasis. the department maintains contact with relevant organisations at the national and international level and in particular is an active participant in the activities of the southern african regional commission for the conservation and utilization of the soil (sarccus) and the eastern african wildlife and national parks annual meetings. much of the impetus for national park development and wildlife conservation in the rm has been due to the personal interest demonstrated by h.e. the life president. material assistance has also been received from the national fauna preservation society of malawi (formerly the nyasaland fauna preservation society) whose crucial role in many early developments is chronicled by hayes 1974, the world wildlife fund, the frankfurt zoological society, the african wildlife leadership foundation and the sa nature foundation. the assistance of the latter organization was of particular importance to the establishment of the wildlife research unit in 1969. the research and 110 monitoring unit in the nyika national park has been set up with canadian funds. however, the main donor has been the british government which has provided k670 000 (ki = ri) to date, while the malawi government provides k270 000 each year. training of malawian rangers has been undertaken at mweka college in tanzania with the support of fao and more recently at cwaka college in k wa zulu , south africa. the national parks and game reserves the total area falling under the protection of the national parks act (1969) or the game act (1954) is 10426 km 2 or roughly 8,6% of th e total surface area of the rm. in addition to this a further 7 770 km 2 (6,4% of total surface area) is protected or partly protected as forest reserve . some details of national parks and game reserves ·are as follows : nyika national park 304 385 ha (figure includ es extensions to the park which have been approved but are not yet gazetted) this park includes virtually the whol e of the famous nyika plateau ( i 830-2606 m) and its escarpment slopes. the main vegetation types protected are extensive montan e grasslands , broad-leaved evergreen forest on the plateau and escarpment including entandophragma excelsum montan e forest and ocotea/ ficalhoa montane forest , phillipia benguelensis communities, protea communities, transitional brachystegia woodlands on the escarpment slopes and conifer forest dominated by junip erus procera . these small patches of junipers and hagenia abyssinica are here at the southern limits of their range, ternstroemia polypetala is only known from the nyika in the flora zambesiaca area (chapman 1968). the nyika is also the type locality of many plants , birds, reptiles and mammals. among the most interesting mammals occurring in the park are the roan antelope hippotragus equinusj crawshay's zebra equus burchelli crawshayiij livingstone's eland taurotragus oryx livingstonii) common reedbuck redunca arundinum) grey duiker sylvicapra grimmia) bush buck tragelaphus scriptus) leopard panthera pardus) side-striped jackal canis adustus, chequered elephant shrew rhyncocyon cirnei. nyasa red and black squirrel heliosciurus lucifer, tree hyrax dendrohyrax arboreus, klipspringer oreotragus oreotragus, spotted hyaena crocuta crocuta and nchima monkey cercopithecus mitis. planned extensions to the park will include greater areas of escarpment slopes and brachystegia woodlands resulting in increases in numbers of those migratory species which move off the plateau annually. buffalo syncerus caffer and elephant loxodonta africana which occur in the northern extension will then be added to the fauna of th e park. iii kasungu national park 220 320 ha this park lies on the miocene surface of central malawi and consists of generally flat landscape with inselbergs. the predominant vegetation is miombo woodland (brachystegia julbernardia isoberlinia) with dambo grasslands. deciduous tree savanna with terminalia sericea) combretum spp, strychnos spp and acacia spp dominant, and echinochloa pyramidalis, imperata cylindrica grasslands in suitable localities are also important. the most common mammals are elephant, buffalo, lichtenstein's hartebeest alcelaphus lichtensteinii} sable hippotragus niger} zebra e.b. selousi) roan, reedbuck, oribi ourebia ourebi) lion panthera leo) leopard, spotted hyaena, grey duiker, sharpe's grysbok raphicerus sharpei) waterbuck kobus ellipsip. rymnus and warthog phacochoerus aethiopicus. there are also small populations of black rhinoceros diceros bicornis) cheetah acinorryx jubatus and johnstone's impala aepyceros melampus johnstonii. lengwe national park 90 720 ha (figure includes extensions to the park which have been approved but are not yet gazetted). situated on the floor of the rift valley this park supports representatives of all the rift valley vegetation types except aquatic and hygrophilous types. the most important communities are newtonia hildebrandtiilpterocarpus antunesii dry deciduous forest and thickets, acacia nigrescens savanna and woodland, dalbergia melanoxylon tree savanna, hyphaene ventricosa palm savanna, combretum apiculatum savanna and colophospermum mopane woodland. the approved extension will add brachystegia and pterocarpus angolensis woodlands to the park. a total of 36 plant species collected in the park have not yet been found elsewhere in malawi, and includes endemics of the shire/zambesi lowlands such as cola mossambicensis. the dry forest and thickets are particularly important as the habitat of the northern-most population of nyala tragelaphus angasi in africa (numbering i 500-2000), and the park also supports suni nesotragus moschatus, bushbuck, bushpig potamochoerus porcus, nchima monkey, chequered elephant shrew, four-toed elephant shrew petrodromus tetradactylus, kudu tragelaphus strepsiceros, impala, buffalo, spotted hyaena, leopard, warthog, sable and roan antelope. this park is also the only known locality of the recently described mollusc rachis cunctatoris (van bruggen 1975) which was collected for the first time in 1970 by one of us (aj.h-m). liwonde national park 58 616 ha this is the newest park in the rm and contains representative vegetation types of the rift valley including reedbeds, papyrus swamp, palm savanna, colophospermum mopane woodland (with individual trees taller than 30 m), piliostigma savanna, brachystegia woodland and dry forest. the most important large mammals are elephant, hippopotamus hip112 popotamus amphibius. sable, kudu and duiker. this is the only park in malawi which contains crocodiles crocodylus niloticus, and which lies within the historical range of johnstone's wildebeest connochaetes taurinus johnstonii. which has been extinct in malawi since 1925. this park includes both banks of the shire river and is therefore of great importance as protected examples of this biome are rare in malawi. vwaza marsh (proposed) game reserve 103680 ha the vwaza marsh lies at the foot of the nyika plateau from where it derives most of its water. the predominant vegetation of the marsh itself is a saline swamp grassland with woody plants restricted to thickets on termite mounds perched above seasonal flood levels. surrounding the marsh is extensive brachystegia woodlands, colophospermum mopane woodlands (at the extreme northern limit of its range), deciduous thicket and piliostigmalacacia tree savanna. this area still supports viable populations of elephant, buffalo, hippo and several species of antelopes . nkhotakota game reserve 174960 ha the reserve occupies the escarpment between the miocene plain and the rift and the vegetation is predominantly miombo woodland. on the slopes of chipata, an inselberg on the edge of the escarpment a small patch of montane forest is protected. the reserve has numbers of elephant, zebra, sable, bush buck and small animals. black rhino were exterminated in the area as recently as the 1950's. however, with its great scenic beauty this reserve is ideally suited for walking trail u tiliza tion . majete game reserve 64 000 ha this reserve also occupies the transitional area extending from the miocene surface down to the shire river on the floor of the rift valley. the vegetation types are varied and include brachystegia woodlands, terminalia sericea tree savanna, pterocarpus angolensis savanna, colophospermum mopane woodland and riverine thicket and forest, small populations of elephant, eland, waterbuck and other antelopes still inhabit the area. mwabvi game reserve 25 920 ha this is the smallest of the game reserves but perhaps one of the most important in the rm and occupies the edge of the rift valley. most of its vegetation types are also found in the lengwe national park and these include colophospermum mopane woodland, acacia nigrescens savanna and extensive dry forest and thickets. this reserve is most important as a sanctuary for black rhinoceros, nyala and suni and as part of the lower shire valley eastern escarpment conservation plan of the i.d.a. development project. 113 as is evident from the above review, most of the major vegetation types of the rm are adequately represented in the national parks and game reserves. several types of broad-leaved evergreen forest as well as widdringtonia whytei forest and several miombo types, and a few endemic or endangered species such as burtdavya nyassica do not occur in game reserves or parks. however, most of these occur in forest reserves where they are presently adequately protected. there are few, if any, vegetation types which are not therefore protected in some way a situation which is probably unique in africa. all the extant large mammals are also found in the sanctuaries. only one large mammal has become extinct in malawi during this century namely the johnstone's wildebeest. the puku kobus vardoni occurred in large numbers on the bua and rusa floodplains in central malawi. excessive hunting led to its disappearance by 1958. however, since 1971 several individuals, originating from zambia , have on occasions been sighted in kasungu national park. there are also records of the occurence of cookson' s wildebeest c.t. cooksonii in the kasungu area this century and isolated individuals have also recently been sighted in the park. future prospects under the enlightened leadership ofh.e. the life president great strides have been made in nature conservation in the 12 years since independence. during this time the total land area under protection (other than forestry lands) has been increased from 5 732 km2 to 10 426 km2 or by 82 per cent. seeing that most biotic communities are now represented in parks or reserves, the tempo of increase of these areas will probably decline in the future. the major systems still requiring representative protection are some of the vegetation types which occur north of the nyika and the aquatic biomes . however, a national park to protect a sample of lacustrine environment has been approved in principle and will be established in 1977. this park will play a vital role in the conservation of the large number of species of cichlids used in the tropical fish trade. it will also be the only biotic reserve covering a deep water lake environment in africa. the park will also be of particular interest as lake malawi has the highest rate of endemism among its fishes of any lake in the world with 223 out of250 species (89%) endemic (beadle 1974). of the larger mammals which can be regarded as endangered are the black rhino (with small populations in kasungu and mwabvi), the cheetah (small population in kasungu), the wild dog lycaon pictus (possibly only vagrants from zambia occur in the kasungu national park), the puku and cookson's wildebeest (isolated individuals in kasungu) and the pangolin manis temmincki which is rare in malawi. in the case of the rhino and cheetah adequate areas of suitable habitat are now protected, but probably none of the parks are large enough to 114 support a viable wild dog population. the crocodile is also endangered outside the liwonde national park. the flying squirrel anomalurus derbianus is the only mammal known to occur outside protected wildlife areas but it is found in forest reserves in northern malawi. having created a viable department of national parks and wildlife which can cater for most of the conservation needs of the country the priority of the malawi government is now to consolidate the department and to develop the existing parks and reserves to their best potential for both conservation and utilization on the basis of practical applied research coupled with a constant environmental monitoring programme. references anon, 1963. note on the problems surrounding the implementation of a wild life utilization policy in a d ensely populated country. in : conservation of nature and natural resources in modern african states. i. u.c.n. publications. new series no. i: 250-252. beadle, l c 1974. the inland wate rs of tropical africa. london: longman. brass , l ] 1953. vegetation of nyasaland. r eport on the vernay-nyasaland expedition of 1946. mem. n.y. bot. card. 8(3). chapman , ] d 1962 . the vegetation of the mlanje mountains. nyasaland. zomba: govt. pro chapman, ] d 1968. malawi. in : conse rvation of vegetation in africa so uth of the sahara. acta. phytogeogr. suec. 54: 215-224. chapman] d and f white. 1970. the evergreen forests of malawi. comm. forestry inst. , univ. of oxford. griffiths , y f 1972 (ed). climates of africa. world survey of climatology. vol. 10. hall-martin , a] 1972. aspects of the plant ecology of the lengwe national park. mala wi. m.sc. thesis, univ. of pretoria. hall-martin , a] 197 5. classification and ordination of forest and thi cket vegetation of th e l engwe national pa rk , malawi. k irkia 10( i) : 131-184. hayes , g d 1967. how independence saved an african reserve . oryx 9: 24-27. hayes , g d 1974. conservation in malawi old and new . oryx 12: 334-340. ]ackson , g 1954. preliminary ecological survey of nyasaland. proc. 2nd. inter. afr. soils coif. leopoldville: 679-690. van bruggen, a c 1975. new taxa of streptaxidae and enidae (mollusca gastropoda pulmonata) from south africa and malawi. zool. m eded. leiden. 49 : 207-223. wild, hand l a grandvaux barbosa . 1967 . vegetation map of the flora zamb esiaca area. salisbury: collins. lis page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 abstract introduction methods results discussion acknowledgements references about the author(s) arne b.r. witt centre for agriculture and biosciences international, nairobi, kenya centre for invasion biology, department of botany and zoology, stellenbosch university, south africa sospeter kiambi kenya wildlife service, nairobi, kenya tim beale centre for agriculture and biosciences international, oxfordshire, united kingdom brian w. van wilgen centre for invasion biology, department of botany and zoology, stellenbosch university, south africa citation witt, a.b.r., kiambi, s., beale, t. & van wilgen, b.w., 2017, ‘a preliminary assessment of the extent and potential impacts of alien plant invasions in the serengeti-mara ecosystem, east africa’, koedoe 59(1), a1426. https://doi.org/10.4102/koedoe.v59i1.1426 original research a preliminary assessment of the extent and potential impacts of alien plant invasions in the serengeti-mara ecosystem, east africa arne b.r. witt, sospeter kiambi, tim beale, brian w. van wilgen received: 26 july 2016; accepted: 03 mar. 2017; published: 22 may 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract this article provides a preliminary list of alien plant species in the serengeti-mara ecosystem in east africa. the list is based on broad-scale roadside surveys in the area and is supplemented by more detailed surveys of tourist facilities in the masai-mara national reserve and adjoining conservancies. we encountered 245 alien plant species; significantly more than previous studies, of which 62 (25%) were considered to have established self-perpetuating populations in areas away from human habitation. these included species which had either been intentionally or accidentally introduced. of the 245 alien plants, 212 (including four species considered to be native to the region) were intentionally introduced into gardens in the national reserve and 51 (24%) had established naturalised populations within the boundaries of these tourism facilities. of the 51 naturalised species, 23 (11% of the 212 alien species) were recorded as being invasive within the ecosystem, outside of lodges and away from other human habitation. currently, the serengeti-mara ecosystem is relatively free of widespread and abundant invasive alien plants, with a few exceptions, but there are extensive populations outside of the ecosystem, particularly to the west, from where they could spread. we address the potential impacts of six species that we consider to pose the highest risks (parthenium hysterophorus, opuntia stricta, tithonia diversifolia, lantana camara, chromolaena odorata and prosopis juliflora). although invasive alien plants pose substantial threats to the integrity of the ecosystem, this has not yet been widely recognised. we predict that in the absence of efforts to contain, or reverse the spread of invasive alien plants, the condition of rangelands will deteriorate, with severe negative impacts on migrating large mammals, especially wildebeest, zebra and gazelles. this will, in turn, have a substantial negative impact on tourism, which is a major economic activity in the area. conservation implications: invasive alien plants pose significant threats to the integrity of the serengeti-mara ecosystem and steps will need to be taken to prevent these impacts. the most important of these would be the removal of alien species from tourist facilities, especially those which are known to be naturalised or invasive, the introduction of control programmes aimed at eliminating outlier invasive plant populations to slow down the spread, and the widespread use of biological control wherever possible. introduction the establishment and management of a network of protected areas is a key component of global strategies to protect biodiversity and to conserve a representative sample of the earth’s ecosystems. proclamation of protected areas is in itself no guarantee that the ecosystems therein will not become degraded, as they face a host of threats, including chronic shortages of management funds, legal and illegal resource use, climate change, pollution and invasion by alien species. ongoing active management will therefore be needed to address these threats. invasive alien species can pose significant threats to protected area ecosystems worldwide (foxcroft et al. 2013a), and one report (de poorter 2007) identified 487 protected areas where invasive alien species were recorded as a threat. in africa (with the notable exception of south africa), very little is known about invasive alien species across the continent’s protected areas (foxcroft, witt & lotter 2013b). a lack of information on the extent of these invasions, and the problems that they cause, ultimately translates into a failure to adequately provide for their management. as a starting point, therefore, it would be important to record the extent of the threats posed by invasive alien species to individual protected areas and to assess the options for achieving effective control (van wilgen et al. 2016). in this article, we report on the findings of surveys of invasive alien plants in the serengeti-mara ecosystem in east africa. prior to our survey, the only available information on alien plants in the area was from henderson (2002) who found 41 ‘problem’ plants in the ngorongoro crater, 8 of which were considered to be native; a report by bukombe et al. (2012) who recorded 13 alien plant species in the serengeti national park, including two species, senna didymobotrya (fresen.) h.s. irwin & barneby (fabaceae) and ricinus communis l. (euphorbiaceae), which are often considered to be native; and a report by clark, lotter and runyoro (2010) who listed 147 invasive alien and indigenous ‘weedy’ species in the ngorongoro conservation area. our surveys subsequently revealed that many of the alien species listed by clark et al. (2010) are not invasive. here, we provide an updated list of the introduced, naturalised and invasive alien plant species present. we then focus on six taxa that are expected to generate the largest impacts and review the degree to which they may be expected to impact the integrity of the serengeti-mara ecosystem. we also make recommendations regarding the management interventions that would be required to prevent or reduce these impacts. methods study site the serengeti-mara ecosystem is a trans-border area that covers ± 100 000 km2 in the northwest of tanzania and southwest of kenya (figure 1). the core conservation areas consist of the masai-mara national reserve in kenya which covers ± 1500 km2, while the serengeti national park in tanzania covers 14 750 km2. both of these protected areas are surrounded by buffer zones such as inner and outer group ranches or conservancies in kenya and the ngorongoro conservation area, loliondo game controlled area and the maswa, grumeti and ikorongo game reserves in tanzania. the area has varied vegetation and topography, including savanna, grassland, riverine forests, inselbergs (characteristic rock outcrops which rise suddenly above a plain) and wetlands. the mean annual rainfall is around 1000 mm, ranging from 508 mm on the plains to 1200 mm near lake victoria in the west, with a short rainy season in october and november and a longer rainy season from march to may. the ecosystem is well known for the migration of 2 million animals (mainly wildebeest, zebras and gazelles), which range between the serengeti national park and surrounding conservation areas and the masai-mara national reserve and the adjoining conservancies, and the area is an important tourist destination. the area has a relatively large number of tourist roads and tracks, especially in the masai-mara national reserve, and several lodges have been built, starting in the 1960s; these lodges and other tourist facilities, such as camping sites, can accommodate thousands of tourists, and many more staff and community members reside permanently within the ecosystem. figure 1: location of the masai-mara national reserve, the serengeti national park, and surrounding protected areas in east africa. species surveys species surveys were undertaken in the serengeti-mara ecosystem during or just after the rains, in order to facilitate plant identification, because actively growing and flowering plants are easier to identify. an initial series of field surveys was carried out in the masai-mara national reserve between 04 and 11 april 2011. during this time, we drove along most of the roads and some of the jeep tracks in an area known as the ‘mara triangle’, the national reserve central plains, the national reserve east and ol derikesi and conservancies to the north (mara north, lemek, ol choro, enonkishu) and along the periphery of conservancies to the east (isaaten and siana) (figure 2), with one observer recording the species seen, its status and approximate location. coordinates, at or within 1 km, of each locality where an alien species was found to be present or naturalised, or invasive and spreading (as defined by pyšek et al. 2004), were recorded using a hand-held global positioning system (gps) receiver. the methodology used was similar to that described by henderson (2007) and rejmánek et al. (2017). where we could not immediately identify a species, herbarium specimens were collected or photographs taken for later identification by taxonomists. in august 2016, we undertook a further road survey to record the extent of parthenium hysterophorus l. (asteraceae) (a species that is currently spreading rapidly, but that is also actively being managed in the ‘mara triangle’) to assess both the degree of spread between 2011 and 2016 and the effectiveness of management interventions. figure 2: (a) location of tourist nodes (lodges and other accommodation) in the masai-mara national reserve and adjacent conservation areas that were surveyed for the occurrence of alien plant species and (b) roads in the area that were surveyed for the occurrence of alien plant species. similar vehicle-based observations were undertaken during two trips en transit through the serengeti national park and ngorongoro crater in 2012. in addition, we conducted vehicle-based surveys in land adjacent to the protected areas, especially to the west of the serengeti national park, where small-scale farming and pastoralism are the main land-use practices. finally, we consolidated our list with those of bukombe et al. (2012), clark et al. (2010) and henderson (2002), based on the surveys they undertook in the ngorongoro conservation area and the serengeti national park. tourism facilities as pathways of alien plant introductions tourist facilities in protected areas can be an important source of invasive alien species. for example, foxcroft, richardson and wilson (2008) surveyed 36 tourist camps and staff villages in the kruger national park, south africa, and identified 258 alien plant species, several of which subsequently became important invaders of the surrounding ecosystem. we therefore surveyed the grounds of 24 tourist facilities (lodges or other accommodation sites, which often included relatively large, fenced grounds covered by natural vegetation) in the masai-mara national reserve (figure 2). we noted all alien plant species present, and recorded whether the species had established naturalised populations within the tourism facilities and whether they had become invasive. species were regarded as naturalised if they were considered to have been reproducing consistently, sustaining populations over many life cycles without direct intervention by humans (richardson et al. 2000). species were recorded as invasive if they were found to be spreading beyond the fences or outside of the tourism facilities or other human habitation. evidence of impact of selected important species although many alien species have established in the serengeti-mara ecosystem, a relatively small number poses disproportionate threats, as they spread rapidly, have the potential to invade large areas and can generate large impacts. according to clark et al. (2010), it has been suggested that acacia mearnsii de wild, leucaena leucocephala (lam.) de wit (fabaceae), caesalpinia decapetala (roth) alston (fabaceae), parkinsonia aculeata l. (fabaceae), melia azedarach l. (meliaceae), jacaranda mimosifolia d. don (bignoniaceae), eucalyptus spp. (myrtaceae), lantana camara (l.) (verbenaceae), datura stramonium l. (solanaceae), lonicera japonica thunb. (caprifoliaceae) and azolla filiculoides lam. (possibly azolla cristata kaulf.) (salviniaceae) pose the biggest threat to the ngorongoro conservation area. however, we are of the opinion that of these 11 species, only l. camara and five others, not listed by any previous studies, pose an even bigger threat to the whole ecosystem based on their recorded impacts elsewhere in the region or on the continent and their current and potential distribution in the region, especially in the serengeti-mara ecosystem (illori et al. 2010; maundu et al. 2009; mcconnachie et al. 2011; shackleton et al. 2017; shackleton et al. in press). all of these are known to be aggressively invasive and have the potential to substantially reduce the ability of rangelands to support grazing mammals, and several have other impacts, being allelopathic or toxic, or having an ability to affect the health of livestock or wildlife. the species were: chromolaena odorata (l.) r.m. king & h.rob. (asteraceae), a shrub from central america lantana camara, a shrub from central and south america opuntia stricta (haw.) haw. (cactaceae), a spinescent stem succulent from central america parthenium hysterophorus l. (asteraceae), an annual herb from tropical america prosopis species and hybrids (fabaceae), trees and shrubs from north, south and central america tithonia diversifolia (hemsl.) a. gray (asteraceae), a shrub from central america. for each species, we located published accounts on the nature and extent of impacts associated with the species, and summarised this information in brief accounts intended to illustrate the potential of the species to impact on the conservation value of the serengeti-mara ecosystem. results species surveys during our roadside surveys, we encountered 62 alien plant species that had established adventive populations in the serengeti-mara ecosystem (table 1). these included ruderal weeds such as bidens pilosa l. (asteraceae), alternanthera pungens kunth (amaranthaceae), gomphrena celosioides mart (amaranthaceae), cirsium vulgare (savi) ten. (asteraceae), flaveria bidentis (l.) kuntze (asteraceae), conyza spp. (asteraceae) and others, which were probably all accidentally introduced or had moved unaided into the ecosystem from adjoining areas. these species were mainly confined to roadsides or other man-made disturbances and were not regarded as being transformers as described by richardson et al. (2000), in that they probably have negligible impacts on biodiversity or ecosystems over a large area. however, other species such as p. hysterophorus, which were probably also accidentally introduced or moved into the ecosystem unaided, are already widespread and abundant and pose a significant threat to biodiversity. a large number of alien species have also been intentionally introduced. of the 245 alien plant species seen and recorded in the ecosystem (including ipomoea hildebrandtii vatke [convolvulaceae], ipomoea cairica [l.] sweet, s. didymobotrya and r. communis which are native to the region but may have been intentionally introduced into the ecosystem as ornamentals, and excluding the additional species recorded by clark et al. [2010] and henderson [2002]), 212 species were considered to have been intentionally introduced, mainly as ornamentals, in tourism accommodation facilities (table 1). of those 212 alien ornamental plant species seen in tourist facilities, 51 were considered to be naturalised in that they had spread and established self-perpetuating populations within the tourist compounds, often in natural vegetation. species such as callisia repens (lindl.) woodson (commelinaceae) had established large populations in woodland understoreys of some lodges together with tradescantia zebrina (rose) d.r. hunt (commelinaceae), while climbers or creepers such as pyrostegia venusta (ker gawl.) miers (bignoniaceae), tecoma capensis (thunb.) lindl. (bignoniaceae), solanum seaforthianum andrews (solanaceae) and others had invaded woodlands, scrambling or climbing over native trees and contributing to canopy collapse. of these naturalised ornamentals, 23 species appeared to have spread beyond the facility fences or other human habitation, where they were also being cultivated, and established populations in the adjacent natural vegetation (table 1). these included species such as catharanthus roseus (l.) g. don (apocynaceae), zinnia peruviana (l.) l. (asteraceae), t. diversifolia, solanum mauritianum scop. (solanaceae), senna spectabilis (dc.) h.s. irwin & barneby (fabaceae), l. camara and tecoma stans (l.) juss. ex kunth. (bignoniaceae). although the cactus species opuntia monacantha (willd.) haw. (cactaceae) and austrocylindropuntia subalata (muehlenpf.) backeb. were present in lodge gardens in the masai-mara national reserve, and adjoining conservancies, they were not found to be invasive, unlike the situation within the serengeti national park and surrounding conservation areas where the former had escaped cultivation and established populations in the wild. opuntia stricta and c. decapetala were not seen in any tourism facilities but were widely grown, especially as living fences, in villages within and immediately adjacent to the serengeti-mara ecosystem from where they have established populations in the natural vegetation. another species that is probably not cultivated but nevertheless abundant on the edge of the ecosystem is c. odorata. table 1: alien plant species, including some native plant species which may be growing outside of their natural range, recorded in the serengeti-mara ecosystem and immediate surrounds (excluding vegetable crops and grains). table 1:(continues…) alien plant species, including some native plant species which may be growing outside of their natural range, recorded in the serengeti-mara ecosystem and immediate surrounds (excluding vegetable crops and grains). table 1:(continues…) alien plant species, including some native plant species which may be growing outside of their natural range, recorded in the serengeti-mara ecosystem and immediate surrounds (excluding vegetable crops and grains). table 1:(continues…) alien plant species, including some native plant species which may be growing outside of their natural range, recorded in the serengeti-mara ecosystem and immediate surrounds (excluding vegetable crops and grains). table 1:(continues…) alien plant species, including some native plant species which may be growing outside of their natural range, recorded in the serengeti-mara ecosystem and immediate surrounds (excluding vegetable crops and grains). table 1:(continues…) alien plant species, including some native plant species which may be growing outside of their natural range, recorded in the serengeti-mara ecosystem and immediate surrounds (excluding vegetable crops and grains). our second roadside survey of the extent of p. hysterophorus in the masai-mara national reserve revealed a significant densification of p. hysterophorus infestations within the national reserve central plains and the establishment of numerous new infestations to the east (figure 3). however, intensive management in the form of ongoing control of p. hysterophorus over several years, using manual (hand-pulling) and chemical (tordon 101 containing the active ingredients picloram and 2,4-d) control interventions in the adjacent ‘mara triangle’ resulted in a reduction in the distribution and abundance of this noxious weed. this provides an illustration of what can potentially be achieved with focussed management programmes, especially when infestations are still small and localised. these efforts need to be expanded because much of this conservation area is under threat from further p. hysterophorus invasions. figure 3: successive surveys in (a) 2011 and (b) 2016 of parthenium hysterophorus in the masai-mara national reserve and surrounding areas in kenya, showing the establishment of new populations to the east of the reserve by 2016. overall, our extensive roadside surveys revealed that the serengeti-mara ecosystem is currently relatively free of transforming invasive alien plants, with the exception of relatively small localised populations of l. camara, t. diversifolia, o. stricta, opuntia ficus-indica, and o. monacantha, with more extensive and widespread infestations of p. hysterophorus in the masai-mara national reserve and adjoining conservancies (figure 4). unless they are managed, these infestations are likely to expand, as the results of our second survey on p. hysterophorus in 2016 clearly demonstrated. this is very likely considering that climate suitability maps, developed by kriticos et al. (2015) and mcconnachie et al. (2011), indicate that most of the serengeti-mara ecosystem is climatically suitable for the establishment of p. hysterophorus, especially the northwestern section (figure 5). other invasive plant species such as c. odorata and prosopis juliflora, which are currently abundant outside of the ecosystem, within predominantly communal lands to the west (figure 4), are also likely to spread into the ecosystem in the foreseeable future, if steps are not taken to prevent this. figure 4: distribution of six invasive alien plant species, (a) chromolaena odorata, (b) lantana camara, (c) opuntia stricta, (d) parthenium hysterophorus, (e) prosopis species and (f) tithonia diversifolia, in and around the serengeti-mara ecosystem, east africa. figure 5: climex generated map of the relative climatic suitability of the serengeti-mara ecosystem and surrounding areas for parthenium hysterophorus based on a model developed by mcconnachie et al. 2011. evidence of impact of selected important species our review of available literature on the impacts of six selected taxa revealed that each poses substantial threats to the serengeti-mara ecosystem, should they be allowed to spread and densify within the ecosystem or invade it from adjacent areas. evidence for the main impacts are summarised below. opuntia stricta is a succulent shrub that was introduced from central america, and it is regarded as a transformer species in savannas and arid grasslands (henderson 2001). the species is highly invasive and forms dense stands, impeding movement and access across the landscape. in madagascar, o. stricta has invaded land used for crop and pasture production, and has encroached on villages and roads, impeding human mobility (larsson 2004). here, the cactus has had a negative impact on native grasses and herbs, and affects trees by inhibiting their growth and regeneration (larsson 2004). the small spines (known as glochids) on the fruit, when consumed by livestock, lodge in their gums, on their tongues or in their gastrointestinal tracts, causing bacterial infections, while the hard seeds may cause rumen impaction, which can be fatal and which often leads to excessive, enforced culling of affected animals (ueckert et al. 1990). similar impacts have been recorded in laikipia county, kenya, where pastoralists have lost significant numbers of livestock (shackleton et al. in press). people who consume the fruits develop diarrhoea and may suffer from serious infections caused by the spines (larsson 2004). in kenya, o. stricta infestations have resulted in the abandonment of land (a.b.r. witt pers. observ.). lantana camara is an invasive shrub or scrambling plant introduced from central and south america. the species invades forest edges, savannas and degraded rangelands, where it forms dense, impenetrable thickets (henderson 2001). the species reduces biodiversity and threatens a host of rare and endangered species. in australia, turner and downey (2010) identified 275 native plant species and 24 native animal species that were threatened by l. camara. in crop production systems in southeast asia, lantana has both reduced yields and increased management costs incurred by growers of durian, pineapples, bananas and rubber (waterhouse 1993). lantana camara is also toxic to livestock, causing pastoral losses that were estimated at aus$ 7.7 million in queensland, australia, in 1985, and which included 1500 animal deaths, reduced productivity, loss of pasture and higher control costs (van oosterhout 2004). in south africa, lantana poisoning accounts for about 25% of all reported cases of livestock poisoning by plants (wells & stirton 1988). there have even been recorded fatalities in people, especially children, after consuming the green fruit (cabi 2016; sharma 2007). lantana camara can also alter fire regimes, allowing fires to penetrate into forests and woodlands that are normally resistant to fire (berry, wevill & curran 2011; day et al. 2003). parthenium hysterophorus is an annual herb native to tropical america, which has become a widespread invader of rangelands and cropping fields in at least 34 countries in africa, asia, australia and the middle east (adkins & shabbir 2014). the species is allelopathic, which enables it to suppress natural vegetation in a wide range of habitats (aggarwal & kohli 1992; evans 1997; mcfadyen 1992; van der laan 2006), including native grasses in the kruger national park (van der laan 2006). the weed was estimated to reduce stocking rates in queensland, australia, by 25% for light to medium infestations, and by as much as 80% for heavy infestations (mcfadyen 1992) and by as much as 90% in india (jayachandra 1971). parthenium hysterophorus also causes severe allergenic reactions (dermatitis, hay fever and asthma) in a large proportion of people who come into contact with it, as well as in livestock and wildlife (patel 2011; towers & mitchell 1983). the weed is now considered, by 90% of the farmers in the lowlands of ethiopia, to be the most serious weed of croplands and grazing areas (tamado & milberg 2004). shrubs/trees in the genus prosopis were introduced into east africa in the 1980s, with some species and hybrids having already invaded over 1 million ha in kenya, where they have the potential to invade nearly half of kenya’s surface area (maundu et al. 2009; witt 2010). the species, including p. juliflora, were originally introduced to kenya, and have recently invaded tanzania. invasive prosopis spp. are associated with many negative impacts, thereby reducing grazing capacity (ndhlovu, milton-dean & esler 2011), eliminating many species from invaded ecosystems (dean et al. 2002; schachtschneider & february 2013; shackleton et al. 2015; steenkamp & chown 1996) and reducing water resources (dzikiti et al. 2013). thus, despite some benefits in the form of fuelwood and edible pods (livestock fodder), their overall net economic contribution is negative, and set to worsen further as the species continues to spread (wise, van wilgen & le maitre 2012). in ethiopia, p. juliflora has reduced understorey cover for perennial grasses from 68% to 2%, and has reduced the number of grass species from seven to two (kebede & coppock 2015); in south africa, a relatively light prosopis spp. invasion (15% cover) led to a 34% reduction in the grazing capacity (ndhlovu et al. 2011). by transforming habitats and eliminating pasture species, p. juliflora is threatening the survival of grévy’s zebra (equus grevyi) in invaded areas (kebede & coppock 2015). dense stands reduce access and impede the movement of people and animals, while the thorns frequently cause injury. local communities in kenya, sudan, eritrea, malawi, south africa and pakistan have all reported negative consequences of these invasions (brown, boudjelas & de poorter 2004; pasiecznik et al. 2001). in semi-arid parts of africa, prosopis trees have depleted the natural resources on which many thousands of people depend, spawning conflict between communities over the diminishing resources. chromolaena odorata is a scrambling shrub introduced from central america. it is an aggressive invader of savanna ecosystems, where it has the potential to transform the vegetation (henderson 2001). one mature plant can produce roughly 1 million seeds per year, which enables rapid spread and the establishment of large populations over a relatively short period (witkowski & wilson 2001). its ability to form dense, impenetrable thickets leads to the displacement of native plant species (te beest, esler & richardson 2015a), while the dry stems and leaves, which are rich in oils, also increase fire intensities (mcfadyen 2004), contributing to additional biodiversity loss. in south africa, infestations are impacting negatively on the breeding biology of the nile crocodile (leslie & spotila 2001), while in cameroon it is displacing native species in the family zingiberaceae, a major food source for the endangered western lowland gorilla (van der hoeven & prins 2007). in southeast asia, it is a serious weed affecting oil palm, rubber, coffee, cashew, fruit and forestry (waterhouse 1993). some agricultural areas in southeast asia ‘have been abandoned because siam weed [i.e. c. odorata] has taken over pasture and crops’ (crc for weed management 2003:1). it also causes serious health problems in livestock and people (aterrado & talatala-sanico 1988; sajise, palis & lales 1972; soerohaldoko 1971), while significantly reducing the livestock-carrying capacities of pastures. tithonia diversifolia is an annual or perennial shrub introduced from central america, which invades savannas and grasslands (henderson 2001). the species forms dense stands which can displace native plants and the animals associated with them. its production of numerous small, light seeds, coupled with its ability to spread vegetatively, allows it to invade and to establish readily and rapidly in new locations (muoghalu & chuba 2005). in nigeria, it was shown to reduce species diversity in invaded plots by 25% (oludare & muoghalu 2014), displacing native vegetation in wetlands (borokini 2011), and contributing to the local extinction of valued native species, including some important medicinal plants (oludare & muoghalu 2014). it is even reported to be out-competing the formidable invasive shrub c. odorata (olubode, awodoyin & ogunyemi 2011). as such it is now considered to be one of the most damaging of all invasive plant species in nigeria (borokini 2011). tithonia diversifolia also competes with agricultural crops (illori et al. 2010) and invasions have reportedly led to the abandonment of some farms in the copperbelt region of zambia (a.b.r. witt pers. observ., 2010). discussion potential impacts on the serengeti-mara ecosystem many exotic species, that have the potential to become invasive, do not always spread rapidly when first introduced to a new environment, but the rate of spread often increases once the species has naturalised and becomes invasive. many of the species we recorded have not been found in the serengeti-mara area until relatively recently, so we can reasonably expect the rate of spread into currently uninvaded areas to increase in the near future. although it is sometimes assumed that relatively unmodified ecosystems (such as the protected areas of the masai-mara and serengeti) will be resistant to invasions, there is also evidence to the contrary. for example, the establishment of c. odorata can be facilitated in savannas and grasslands by small-scale disturbances that create micro-sites for establishment, ultimately aiding their long-term persistence in grass-dominated areas (te beest, mpandza & olff 2015b). this is borne out by the fact that c. odorata has aggressively invaded savanna vegetation in the hluhluwe-imfolozi park in south africa (dumalisile 2009; howison 2009). it cannot be assumed that these protected ecosystems will remain uninvaded. parthenium hysterophorus is also invasive in many other protected areas in africa, and we have observed that this species, which is often regarded as only being able to invade disturbed or over-grazed areas, establishes readily on termite mounds that are scattered across the serengeti-mara landscape, providing widespread foci from which further invasions can commence. although invasive alien plants pose substantial threats to the integrity of the serengeti-mara ecosystem, this has not yet been widely recognised. for example, in a recent comprehensive treatment of the ecology and conservation of the serengeti (sinclair et al. 2015), invasive alien plants are only mentioned once, where it is stated that: although the exact extent of these invasions is not known, these species have taken over grasslands in other ecosystems and could exclude wildebeest from accessing critical areas in the future. (p. 168) given that the large numbers of grazing mammals are dependent on good-quality forage, and given further that invasions by the alien plant species currently establishing in the area can reduce carrying capacities by up to 90% (jayachandra 1971; mcfadyen 1992; ndlovu et al. 2011; yapi 2014), large impacts can be expected. for example, ogutu et al. (2009) found that the abundance of six large grazing mammal species declined markedly and persistently throughout the masai-mara national reserve between 1989 and 2003, and that the declines were contemporaneous with progressive habitat deterioration because of a range of factors, although invasion by alien plants was not considered. the serengeti wildebeest population is regulated by food supply, and the main cause of mortality (75% of cases) was found to be under-nutrition (mduma, sinclair & hilborn 1999). rampant invasion would almost certainly result in large losses of rangeland fodder, leading to drastic declines in populations of wildebeest and other large grazing mammals. a drastic decline in wildebeest numbers would trigger many changes, as wildebeest are currently remarkably abundant, and influence virtually every dynamic of the ecosystem (grant et al. 2015). thus, plant invasion could potentially have big effects not only on ecosystem integrity and productivity but also on tourism, which is a very important contributor to the economies of both kenya and tanzania. appropriate management responses given that alien plant invasions pose large threats to the serengeti-mara ecosystem, it would seem prudent to develop and implement control programmes to reduce the severity of these threats. we propose that three key interventions should be implemented as a matter of urgency. firstly, all alien plant species, especially those that are known to be naturalised, invasive or potentially invasive, should be removed from the grounds of tourist facilities. secondly, control programmes aimed at eliminating outlier populations should be implemented to slow spread. finally, biological control solutions should be implemented wherever possible. we discuss each of these in the following sections. removal of alien plants around tourist facilities: all alien plants, whether invasive or not, should ideally be removed from the grounds of any developed parts of the protected areas. tourist and staff facilities can be a major source of invasive species, and this would be best addressed by removing all alien plants while populations are still small. in the kruger national park, south africa, foxcroft and freitag-ronaldson (2007) found that the park staff played a major role in facilitating alien plant invasions. staff members unwittingly introduced alien species into the gardens of tourist camps as well as into their own gardens, for ornamental and other uses. many species subsequently escaped and became invasive. attempts to remove these species began in the mid-1980s, but there was significant resistance from the residents towards the alien plant control team for many years (foxcroft et al. 2008). this was overcome as an understanding of the problem grew, and strategies employed by the kruger national park team included an initial focus on high-risk species (leaving lower-risk species in place in the meantime), and clearing gardens whenever there was staff turnover that resulted in temporary vacating of houses. similar problems can be expected in the serengeti-mara ecosystem. however, there is legislative and other support for invasive species interventions in and outside of protected areas in africa, and specifically in kenya, which would provide strong justification for the removal of alien species in the face of resistance (table 2). table 2: some national, regional and african legislation, policies, conventions and treaties that make reference to invasive alien plants. implementation of control programmes: control programmes should be initiated as soon as possible, with priority being given to lightly invaded areas, isolated populations or the edges of invading populations. higgins, richardson and cowling (2000) demonstrated that clearing strategies that prioritised low-density sites dominated by juvenile alien plants proved to be significantly more cost-effective than strategies that targeted densely invaded areas. these authors also found that delaying the initiation of clearing operations considerably increased the eventual costs of control and the risks to native biodiversity. such early interventions can be very effective, as shown by the results of clearing of p. hysterophorus in the ‘mara triangle’ (figure 3). implementation of biological control: we strongly advocate the fullest possible use of biological control, which should be integrated with other control practices, wherever possible. biological control is a safe, inexpensive and sustainable option for gaining control of many invasive alien plant species. the main benefits of biological control are that the agents establish self-perpetuating populations and often establish throughout the range of the target weed, including areas that are not accessible for chemical or mechanical control; control of the weed is permanent; there are no negative impacts on the environment; the cost of biological control programmes is low relative to other approaches, and in most cases only requires a once-off investment; and benefits can be reaped by many stakeholders independent of their financial status and irrespective of whether they contributed to the initial research (greathead 1995). studies in south africa have demonstrated phenomenal returns on investment in economic terms from biological control projects, where estimated benefit:cost ratios ranged from 8:1 up to 3726:1 (van wilgen & de lange 2011). biological control is arguably an indispensable element of any integrated programme to control invasive alien plants, as most other interventions will fail in the long term if used in isolation. despite this, concerns over the safety of the practice often prevent its implementation in many countries. much of the resistance to the use of biological control arises from ignorance, but the track record of the practice suggests that this should not be the case. it was estimated that by the end of 2012, there were 1555 separate and intentional releases of 469 species of weed biological control agents against 175 species of non-native target weeds (when related taxa of unidentified plant species, such as some opuntia species, are counted as single target weeds) (winston et al. 2014). these so-called ‘classical’ biological control projects have been conducted in a total of 90 countries (winston et al. 2014), with an excellent record of safety and success (van wilgen, moran & hoffmann 2013). there are already several agents that would be available for use against major weed species that currently threaten the serengeti-mara ecosystem. for example, infestations of o. stricta have been brought under control in the kruger national park, south africa, by the introduction and establishment of a sap-sucking bug, dactylopius opuntiae (cockerell) (dactylopiidae) (foxcroft & hoffmann 2000; hoffmann, moran & zeller 1998). this classical biological control agent was recently released on o. stricta in laikipia county, kenya, where it has established and is reducing the density and spread of this invasive cactus. in australia, the ability of p. hysterophorous to form tall and dense invasive stands has been considerably reduced through the release of multiple biological control agents, thus substantially increasing the effectiveness of other control interventions (dhileepan & mcfadyen 2012). work on developing suitable biological control for p. hysterophorus has been initiated in south africa, building on the work in australia (strathie, mcconnachie & retief 2011), with a number of agents already having been released in the field. a chrysomelid beetle, zygogramma bicolorata pallister (chrysomelidae) has also been released in ethiopia and around arusha, tanzania, although establishment in the field is yet to be confirmed. biological control research in south africa has also addressed l. camara (urban et al. 2011), with a number of these agents already present in east africa. however, some newly released and established agents in south africa could complement those agents already present in kenya and tanzania, such as the flowerbud-galling mite aceria lantanae (cook) (eriophyidae) and the root-feeding flea beetle longitarsus bethae (chrysomelidae) (urban et al. 2011). attempts have been made to establish the gall-forming fly, cecidochares connexa (macquart) (tephritidae), on c. odorata in northwestern tanzania, but we will only be able to confirm establishment at the end of 2017. a number of other agents for the control of c. odorata have also been released and have subsequently been established in south africa (zachariades et al. 2011b). a number of seed-feeding beetles have been released in south africa and elsewhere for the control of invasive prosopis species (zachariades et al. 2011a), and research is currently underway to develop agents that feed on the vegetative parts of this invasive tree. research has also been initiated in south africa to develop agents for the control of t. diversifolia (simelane, mawela & fourie 2011). it is imperative that the kenyan and tanzanian authorities work together to facilitate the introduction, mass rearing and release of additional classical biological control agents to complement any current and future invasive plant management strategies in the serengeti-mara ecosystem. failure to do so would result in the possible demise of one of the natural wonders of the world, the annual wildebeest migration. acknowledgements the authors are grateful to the jrs biodiversity foundation for providing resources for surveys throughout the east african region and for the production of distribution maps. grumeti fund provided additional information on the distribution of chromolaena odorata. the authors extend their thanks to staff from kenya wildlife service, the masai-mara national reserve, including brian heath, and others for logistical and other support. the authors would also like to thank the lodge owners and managers, who kindly allowed them access to lodge gardens and other tourist facilities. the authors thank the dst-nrf centre of excellence for invasion biology and the national research foundation (grant 87550 to b.v.w.). they would also like to thank the australian high commission, kenya, for providing funding for initial surveys and other activities in the masai-mara national reserve. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions a.b.r.w. was responsible for initiating the field surveys, which he conducted with the assistance of s.k. t.b. undertook spatial analyses and compiled the maps. the interpretation of results and writing of the article was shared between b.w.v.w. and a.b.r.w. references adkins, s. & shabbir, a., 2014, ‘biology, ecology and management of the invasive parthenium weed (parthenium hysterophorus l.)’, pest management science 70, 1023–1029. https://doi.org/10.1002/ps.3708 african union, 2016, african convention on nature conservation and natural resources, viewed 23 may 2016, from https://www.au.int/web/sites/default/files/treaties/7782-file-african_convention_conservation_nature_natural_resources.pdf aggarwal, a. & kohli, r.k., 1992, ‘screening of crops for seed germination against parthenium hysterophorus l. leachates’, in p. tauro & s.s. narwal (eds.), proceedings of the 1st national symposium on allelopathy in agroecosystems, hisar, india, february 1992, pp. 66–68. aterrado, e.d. & talatala-sanico, r.l., 1988, ‘status of chromolaena odorata research in the philippines’, in r. muniappan (ed.), proceedings of the first international workshop on biological control of chromolaena odorata, bangkok, thailand, february 19–march 04, 1988, pp. 53–55. berry, z.c., wevill, k. & curran, t.j., 2011, ‘the invasive weed lantana camara increases fire risk in dry rainforest by altering fuel beds’, weed research 51, 525–533. https://doi.org/10.1111/j.1365-3180.2011.00869.x borokini, t.i., 2011, ‘invasive alien plant species in nigeria and their effects on biodiversity conservation’, tropical conservation science 4, 103–110. https://doi.org/10.1177/194008291100400110 brown, l.s., boudjelas, s. & de poorter, m., 2004, 100 of the world’s worst invasive alien species: a selection from the global invasive species database, the invasive species specialist group (issg), species survival commission (ssc) of the world conservation union (iucn), auckland, new zealand. bukombe, j.k., mwita, m., kittle, a. & kija, h., 2012, invasion of alien plants in serengeti: a call for management action, tanzania wildlife research institute, arusha. cabi, 2016, lantana camara, in invasive species compendium, cab international, wallingford, uk, viewed 20 june 2016, from http://www.cabi.org/isc clark, k., lotter, w.d. & runyoro, v., 2010, ngorongoro conservation area: invasive alien plant strategic management plan 2010, ngorongoro conservation area authority, arusha, tanzania. crc for weed management, 2003, weed management guide: siam weed or chromolaena (chromolaena odorata), crc for australian weed management, commonwealth department of environment and heritage, australia. day, m.d., wiley, c.j., playford, j. & zalucki, m.p., 2003, lantana: current management, status and future prospects, status and future prospects, aciar monograph 102, canberra, australia. dean, w.r.j., anderson, m.d., milton, s.j. & anderson, t.a., 2002, ‘avian assemblages in native acacia and alien prosopis drainage line woodland in the kalahari, south africa’, journal of arid environments 51 1–19. https://doi.org/10.1006/jare.2001.0910 de poorter, m., 2007, invasive alien species and protected areas: a scoping report. part 1. scoping the scale and nature of invasive alien species threats to protected areas, impediments to invasive alien species management and means to address those impediments, global invasive species programme, invasive species specialist group, viewed 23 july 2016, from http://www.issg.org/gisp_publications_reports.htm dhileepan, k. & mcfadyen, r.c., 2012, ‘parthenium hysterophorus l. – parthenium’, in j. cullen, m. julien & r. mcfadyen (eds.), biological control of weeds in australia, pp. 448–462, csiro publishing, melbourne, australia. dumalisile, l., 2009, ‘effects of chromolaena odorata on mammalian biodiversity in hluhluwe-imfolozi park, south africa’, msc thesis, university of pretoria, south africa. dzikiti, s., schachtschneider, k., naiken, v., gush, m., moses, g. & le maitre, d.c., 2013, ‘water relations and the effects of clearing invasive prosopis trees on groundwater in an arid environment in the northern cape, south africa’, journal of arid environments 90, 103–113. https://doi.org/10.1016/j.jaridenv.2012.10.015 environmental treaties and resource indicators, 2016, protocol concerning protected areas and wild fauna and flora in the eastern african region, viewed 03 may 2016, from http://sedac.ciesin.org/entri/texts/eastafrpro.html evans, h.c., 1997, ‘parthenium hysterophorus: a review of its weed status and the possibilities for biological control’, biocontrol news and information 18, 89n–98n. foxcroft, l.c. & freitag-ronaldson, s., 2007, ‘seven decades of institutional learning: managing alien plant invasions in the kruger national park, south africa’, oryx 41, 160–167. https://doi.org/10.1017/s0030605307001871 foxcroft, l.c. & hoffmann, j.h., 2000, ‘dispersal of dactylopius opuntiae cockerell (homoptera: dactylopiidae), a biological control agent of opuntia stricta (haworth.) haworth. (cactaceae) in the kruger national park’, koedoe 43, 1–5. https://doi.org/10.4102/koedoe.v43i2.194 foxcroft, l.c., pyšek, p., richardson, d.m. & genovesi, p., 2013a, ‘plant invasions in protected areas: patterns, problems and challenges’, springer, dordrecht. foxcroft, l.c., richardson, d.m. & wilson, j.r., 2008, ‘ornamental plants as invasive aliens: problems and solutions in kruger national park, south africa’, environmental management 41, 32–51. https://doi.org/10.1007/s00267-007-9027-9 foxcroft, l.c., witt, a. & lotter, w.d., 2013b, ‘icons in peril: invasive alien plants in african protected areas’, in l.c. foxcroft, p. pyšek, d.m. richardson & p. genovesi (eds.), plant invasions in protected areas: patterns, problems and challenges, pp. 117–143, springer, dordrecht. grant, j., hopcraft, c., holdo, r.m., mwangomo, e., mduma, s.a.r., thirgood, s.j. et al., 2015, ‘why are wildebeest the most abundant herbivore in the serengeti ecosystem?’, in a.r.e. sinclair, k.l. metzger, s.a.r. mduma & j.m. fryxell (eds.), serengeti iv: sustaining biodiversity in a coupled human-natural system, pp. 125–174, university of chicago press, chicago, il. greathead, d.j., 1995, ‘benefits and risks of classical biological control’, in h.m.t. hokkanen & j.m. lynch (eds.), biological control: benefits and risks, pp. 53–63, cambridge university press, cambridge. henderson, l., 2001, alien weeds and invasive plants, plant protection research institute handbook no. 12, agricultural research council, pretoria. henderson, l., 2002, ‘problem plants in ngorongoro conservation area, tanzania’, unpublished report submitted to the nca in november 2002, 15p. henderson, l., 2007, ‘invasive, naturalised and casual alien plants in southern africa: a summary based on the south african plant invaders atlas (sapia)’, bothalia 37, 215–248. https://doi.org/10.4102/abc.v37i2.322 higgins, s.i., richardson, d.m. & cowling, r.m., 2000, ‘using a dynamic landscape model for planning the management of alien plant invasions’, ecological applications 10, 1833–1848. https://doi.org/10.1890/1051-0761(2000)010[1833:uadlmf]2.0.co;2 hoffmann, j.h., moran, v.c. & zeller, d.a., 1998, ‘long-term population studies and the development of an integrated management programme for control of opuntia stricta in kruger national park, south africa’, journal of applied ecology 35, 156–160. https://doi.org/10.1046/j.1365-2664.1998.00283.x howison, o.e., 2009, ‘the historical spread and potential distribution of the invasive alien plant chromolaena odorata in hluhluwe-imfolozi park’, msc thesis, university of kwazulu-natal, durban, south africa. illori, o.j., otusanya, o.o., adelusi, a.a. & sanni, r.o., 2010, ‘allelopathic activities of some weeds in the asteraceae family’, international journal of botany 61, 161–163. https://doi.org/10.3923/ijb.2010.161.163 jayachandra, j., 1971, ‘parthenium weed in mysore state and its control’, current science 40, 568–569. kanga, e., kenana, l., ngoru, b. & lala, f. (eds.), 2013, national strategy and action plan for the management of invasive species in kenya’s protected areas 2013–2018, kenya wildlife service (kws), nairobi, kenya. kebede, a.t. & coppock, d.l., 2015, ‘livestock-mediated dispersal of prosopis juliflora imperils grasslands and the endangered grevy’s zebra in northeastern ethiopia’, rangeland ecology & management 68, 402–407. https://doi.org/10.1016/j.rama.2015.07.002 kenya forest service, 2016, special issue: kenya gazette supplement no. 88 (acts no. 7), the forests act, 2005, government printer, nairobi, kenya, viewed 23 may 2016, from http://www.kenyaforestservice.org/images/mmmb/forests%20act%20no.7%20of%202005.pdf kenya wildlife service, 2016, special issue: kenya gazette supplement no. 181 (acts no. 47), the wildlife and conservation management act, 2013, government printer, nairobi, kenya, viewed 03 june 2016, from http://www.kws.go.ke/downloads kriticos, d.j., brunel, s., ota, n., fried, g., oude lansink, a.g.j.m., panetta, f.d. et al., 2015, ‘downscaling pest risk analyses: identifying current and future potentially suitable habitats for parthenium hysterophorus with particular reference to europe and north africa’, plos one 10(9), e0132807. https://doi.org/10.1371/journal.pone.0132807 larsson, p., 2004, introduced opuntia spp. in southern madagascar: problems and opportunities, minor field studies no 285, swedish university of agricultural sciences, uppsala. leslie, a.j. & spotila, j.r., 2001, ‘alien plant threatens nile crocodile (crocodylus niloticus) breeding in lake st. lucia, south africa’, biological conservation 98, 347–355. https://doi.org/10.1016/s0006-3207(00)00177-4 maundu, p., kibet, s., morimoto, y., imbumi, m. & adeka, r., 2009, ‘impacts of prosopis juliflora on kenya’s semi-arid and arid ecosystems and local livelihoods’, biodiversity 10, 33–50. https://doi.org/10.1080/14888386.2009.9712842 mcconnachie, a.j., strathie, l.w., mersie, w., gebrehiwot, l., zewdie, k., abdurehim, a. et al., 2011, ‘current and potential geographical distribution of the invasive plant parthenium hysterophorous (asteraceae) in eastern and southern africa’, weed research 51, 71–84. https://doi.org/10.1111/j.1365-3180.2010.00820.x mcfadyen, r.c., 2004, ‘chromolaena in east timor: history, extent and control’, in m.d. day & r.e. mcfadyen (eds.), chromolaena in the asia-pacific region, proceedings of the 6th international workshop on biological control and management of chromolaena, cairns, australia, may 06–09, 2003, pp. 8. mcfadyen, r.e., 1992, ‘biological control against parthenium weed in australia’, crop protection 11, 400–407. https://doi.org/10.1016/0261-2194(92)90021-v mduma, s.a., sinclair, a.r.e. & hilborn, r., 1999, ‘food regulates the serengeti wildebeest: a 40-year record’, journal of animal ecology 68, 1101–1122. https://doi.org/10.1046/j.1365-2656.1999.00352.x muoghalu, j.i. & chuba, d.k., 2005, ‘seed germination and reproductive strategies of tithonia diversifolia (hemsl.) gray and tithonia rotundifolia (pm) blake’, applied ecology and environmental research 3, 39–46. https://doi.org/10.15666/aeer/0301_039046 national council for law reporting, 2012, laws of kenya, suppression of noxious weeds act, chapter 325, viewed 23 may 2016, from http://kenyalaw.org/kl/fileadmin/pdfdownloads/acts/suppressionofnoxiousweedsact_cap325.pdf national environment management authority, 2016, environmental management and coordination act of 1999, viewed 23 may 2016, from http://www.nema.go.ke/images/docs/legislation%20and%20policies/environmental%20act%20(emca1999)%20.pdf ndhlovu, t., milton-dean, s.j. & esler, k.j., 2011, ‘impact of prosopis (mesquite) invasion and clearing on the grazing capacity of semiarid nama karoo rangeland, south africa’, african journal of range & forage science 28, 129–137. https://doi.org/10.2989/10220119.2011.642095 ogutu, j.o., piepho, h.p., dublin, h.t., bhola, n. & reid, r.s., 2009, ‘dynamics of mara–serengeti ungulates in relation to land use changes’, journal of zoology 278, 1–14. https://doi.org/10.1111/j.1469-7998.2008.00536.x olubode, o.s., awodoyin, r.o. & ogunyemi, s., 2011, ‘floral diversity in the wetlands of apete river, eleyele lake and oba dam in ibadan, nigeria: its implication for biodiversity erosion’, west african journal of applied ecology 18, 109–119. https://doi.org/10.4314/wajae.v18i1.70319 oludare, a. & muoghalu, j.i., 2014, ‘impact of tithonia diversifolia (hemsly) a. gray on the soil, species diversity and composition of vegetation in ile-ife (southwestern nigeria), nigeria’, international journal of biodiversity and conservation 6, 555–562. https://doi.org/10.5897/ijbc2013.0634 pasiecznik, n.m., felker, p., harris, p.j.c., harsh, l.n., cruz, g., tewari, j.c. et al., 2001, ‘the prosopis juliflora–prosopis pallida complex: a monograph’, hdra, coventry. patel, s., 2011, ‘harmful and beneficial aspects of parthenium hysterophorus: an update’, biotech 1, 1–9. https://doi.org/10.1007/s13205-011-0007-7 pyšek, p., richardson, d.m., rejmánek, m., webster, g.l, williamson, m. & kirschner, j., 2004, ‘alien plants in checklists and floras: towards better communication between taxonomists and ecologists’, taxon 53, 131–143. https://doi.org/10.2307/4135498 rejmánek, m., huntley, b.j., le roux, j.j. & richardson, d.m., 2017, ‘a rapid survey of the invasive plant species in western angola’, african journal of ecology 55, 56–69. https://doi.org/10.1111/aje.12315 richardson, d.m., pyšek, p., rejmanek, m., barbour, m.g, panetta, f.d. & west, c.j., 2000, ‘naturalization and invasion of alien plants: concepts and definitions’, diversity & distributions 6, 93–107. https://doi.org/10.1046/j.1472-4642.2000.00083.x sajise, p.e., palis, r.k. & lales, j.s., 1972, ‘chromolaena odorata imperils grassland’, pasture newsletter 1, 1–2. schachtschneider, k. & february, e.c., 2013, ‘impact of prosopis invasion on a keystone tree species in the kalahari desert, plant ecology 214, 597–605. https://doi.org/10.1007/s11258-013-0192-z shackleton, r.t., le maitre, d.c., van wilgen, b.w. & richardson, d.m., 2015, ‘the impact of invasive alien prosopis species (mesquite) on native plants in different environments in south africa’, south african journal of botany 97, 25–31. https://doi.org/10.1016/j.sajb.2014.12.008 shackleton, r.t., witt, a.b.r., nunda, w. & richardson, d.m., 2017, ‘chromolaena odorata (siam weed) in eastern africa: distribution and socioecological impacts’, biological invasions 19, 1285–1298. https://doi.org/10.1007/s10530-016-1338-4 shackleton, r.t., witt, a.b.r., merinyi, f. & van wilgen, b.w., in press, ‘distribution and socio-ecological impacts of the invasive alien cactus opuntia stricta in eastern africa’, biological invasions. sharma, o.p., 2007, ‘a review of the hepatotoxic plant lantana camara’, critical reviews in toxicology 37, 313–352. https://doi.org/10.1080/10408440601177863 simelane, d.o., mawela, k.v. & fourie, a., 2011, ‘prospective agents for the biological control of tithonia rotundifolia (mill.) s.f. blake and tithonia diversifolia (hemsl.) a. gray (asteraceae) in south africa’, african entomology 19, 443–450. https://doi.org/10.4001/003.019.0223 sinclair, a.r.e., metzger, k.l., mduma, s.a.r. & fryxell, j.m., 2015, serengeti iv: sustaining biodiversity in a coupled human-natural system, university of chicago press, chicago, il. soerohaldoko, s., 1971, ‘on the occurrence of eupatorium odoratum at the game reserve penanjung, west java, indonesia’, weeds in indonesia 2, 9. steenkamp, h.e. & chown, s.l., 1996, ‘influence of dense stands of an exotic tree, prosopis glandulosa benson, on a savanna dung beetle (coleoptera: scarabaeinae) assemblage in southern africa’, biological conservation 78, 305–311. https://doi.org/10.1016/s0006-3207(96)00047-x strathie, l.w., mcconnachie, a.j. & retief, e., 2011, ‘initiation of biological control against parthenium hysterophorus l. (asteraceae) in south africa’, african entomology 19, 378–392. https://doi.org/10.4001/003.019.0224 tamado, t. & milberg, p., 2004, ‘control of parthenium (parthenium hysterophorus) in grain sorghum (sorghum bicolour) in the smallholder farming system in eastern ethiopia’, weed technology 18, 100–105. https://doi.org/10.1614/wt-03-033r te beest, m., esler, k.j. & richardson, d.m., 2015a, ‘linking functional traits to impacts of invasive plant species: a case study’, plant ecology 216(2), 293–305. https://doi.org/10.1007/s11258-014-0437-5 te beest, m., mpandza, n.j. & olff, h., 2015b, ‘fire and simulated herbivory have antagonistic effects on resistance of savanna grasslands to alien shrub invasion’, journal of vegetation science 26, 114–122. https://doi.org/10.1111/jvs.12208 towers, g.h.n. & mitchell, j.c., 1983, ‘the current status of the weed parthenium hysterophorus l. as a cause of allergic contact dermatitis’, contact dermatitis 9, 465–469. https://doi.org/10.1111/j.1600-0536.1983.tb04465.x turner, p.j. & downey, p.o., 2010, ‘ensuring invasive alien plant management delivers biodiversity conservation: insights from an assessment of lantana camara in australia’, plant protection quarterly, 25, 102–110. ueckert, d.n., petersen, j.l., potter, r.l., whipple, j.d. & wagner, m.w., 1990, range and sheep management for reducing pearmouth and other pricklypear-related health problems in sheep flocks, texas agricultural experimental station program report 4782, 40–41, texas a&m university, college station, san angelo, texas, usa. urban, a.j., simelane, d.o., retief, e., heystek, f., williams, h.e. & madire, l.g., 2011, ‘the invasive “lantana camara l.” hybrid complex (verbenaceae): a review of research into its identity and biological control in south africa’, african entomology 19, 315–348. https://doi.org/10.4001/003.019.0225 van der hoeven, c.a. & prins, h.h.t., 2007, ‘invasive plant species threatens gorilla in equatorial africa’, phd thesis, department of environmental sciences, resource ecology group, wageningen university, wageningen, the netherlands. van der laan, m., 2006, ‘allelopathic interference potential of the alien invader plant parthenium hysterophorus l.’, msc thesis, university of pretoria, south africa. van oosterhout, e., 2004, lantana control manual: current management and control options for lantana (lantana camara) in australia, department of natural resources, mines and energy, brisbane, australia. van wilgen, b.w. & de lange, w.d., 2011, ‘the costs and benefits of biological control of invasive alien plants in south africa’, african entomology 19, 504–514. https://doi.org/10.4001/003.019.0228 van wilgen, b.w., fill, j.m., baard, j., cheney, c., forsyth, a.t. & kraaij, t., 2016, ‘historical costs and projected future scenarios for the management of invasive alien plants in protected areas in the cape floristic region’, biological conservation 200, 168–177. van wilgen, b.w., moran, v.c. & hoffmann, j.h., 2013, ‘some perspectives on the risks and benefits of biological control of invasive alien plants in the management of natural ecosystems’, environmental management 52, 531–540. https://doi.org/10.1007/s00267-013-0099-4 waterhouse, d.f., 1993, the major arthropod pests and weeds of agriculture in southeast asia: distribution, importance and origin, australian centre for international agricultural research, canberra, 141p. wells, m.j. & stirton, c.h., 1988, lantana camara: a poisonous declared weed, department of agriculture and water supply, pretoria, south africa. winston, r.l., schwarzländer, m., hinz, h.l., day, m.d., cock, m.j.w. & julien, m.h., 2014, biological control of weeds: a world catalogue of agents and their target weeds, 5th edn., usda forest service, forest health technology enterprise team, morgantown, wv. wise, r.m., van wilgen, b.w. & le maitre, d.c., 2012, ‘costs, benefits and management options for an invasive alien tree species: the case of mesquite in the northern cape’, journal of arid environments 84, 80–90. https://doi.org/10.1016/j.jaridenv.2012.03.001 witkowski, e.t.f. & wilson, m., 2001, ‘changes in density, biomass, seed production and soil seed banks of the non-native invasive plant, chromolaena odorata, along a 15 year chronosequence’, plant ecology 152(1), 13–27. https://doi.org/10.1023/a:1011409004004 witt, a.b.r., 2010, ‘biofuels and invasive species from an african perspective – a review’, gcb bioenergy 2, 321–329. https://doi.org/10.1111/j.1757-1707.2010.01063.x yapi, t.s., 2014, ‘an assessment of the impacts of invasive australian wattle species on grazing provision and livestock production in south africa’, msc thesis, stellenbosch university, south africa. zachariades, c., hoffmann, j.h. & roberts, a.p., 2011a, ‘biological control of mesquite (prosopis species) (fabaceae) in south africa’, african entomology 19, 402–415. https://doi.org/10.4001/003.019.0230 zachariades, c., strathie, l.w., retief, e. & dube, n., 2011b, ‘progress towards the biological control of chromolaena odorata (l.) r.m. king & h. rob. (asteraceae) in south africa’, african entomology 19, 282–302. https://doi.org/10.4001/003.019.0229 panagos & reilly.qxd use of an area-based survey technique to detect vegetation changes in sour bushveld m.d. panagos and b.k. reilly panagos, m.d. and b.k. reilly. 2006. use of an area-based survey technique to detect vegetation changes in sour bushveld. koedoe 49(1): 69–78. pretoria. issn 0075-6458. most strategies for monitoring vegetation change on reserves and game ranches are based on point methods. area-based methods form the basis of initial floristic classifications from which vegetation maps are constructed. the question arises whether or not these area-based methods can be used for monitoring or not. this paper compares two area-based data sets from the same sites with an intervening period of five years. data were collected on a sour bushveld game farm, in the limpopo province of south africa. the structure of the woody vegetation, particularly in two communities reflected a plant successional progression over time. all six re-surveyed sites reflected an increase in plant species richness and this increase was most substantial in one community where the total number of plant species increased from 17 in 1996 to 34 in 2001 at one sampling site and from 26 in 1996 to 45 in 2001 at the other. this study has shown that change in species composition can be detected using area-based sampling techniques but that absolute measures, such as density, should be employed rather than estimates, especially with regard to plant cover. keywords: area-based vegetation sampling, species composition, structure, canopy cover m.d. panagos and b.k. reilly, department of nature conservation, tshwane university of technology, private bag x680, pretoria, 0001 republic of south africa. issn 0075-6458 69 koedoe 49/1 (2006) introduction monitoring has been defined as “the maintenance of regular surveillance to test the null hypothesis of no change in predefined properties of a system which is vulnerable to impacts, the nature and timing of which are not necessarily known” (mentis 1984). ferrar (1983) states that monitoring and evaluation of vegetation is the key to natural resource management and vegetation monitoring involves three fundamental steps: a) quantification of the vegetation characteristics; b) assessment of any change to these characteristics over time; and c) projection of trends in these characteristics in the future. despite vegetation monitoring enjoying considerable attention in the 1970s and 1980s, this vital veld management action has unfortunately not been sustained due to inappropriate and time consuming techniques being used (short et al. 2003). point-based techniques originally proposed by levy & madden (1933) and later dyksterhuis (1949) have been adopted and variously modified in south africa, starting with the development of the wheel-point technique (tidmarsh & havenga 1955), the nearest plant method (foran et al. 1978), the benchmark method (tainton et al. 1980) and the foot-point method (mentis 1981). more recently, a so-called key species technique was developed (trollope 1990) and implemented in the kruger national park. they have been shown to be efficient in terms of time when compared to area-based sampling (panagos & zacharias 1995) and lend themselves readily to statistical analyses. however, since these techniques make use of only 100 or 200 points for data collection, their ability to show change with high precision has been questioned in terms of basal cover (mentis et al. 1980), less frequently encountered plants (everson et al. 1990; o'connor panagos & reilly.qxd 2006/04/17 10:32 pm page 69 & roux 1995; panagos 1995; short et al. 2003), and statistical power (reilly & panagos 2002). also, all of these techniques are used primarily for measuring the condition of the grass sward and seldom include nongraminoid plants (when these plants are included they are lumped into two broad categories termed ‘forbs’ and ‘woodies’). more recently, techniques such as becvol (smit 1989) and the revised procedure for conducting veld condition assessments (vcas) in the kruger national park (zambatis 2002) have been included with herbaceous layer techniques to include some form of woody stratum evaluation. furthermore, problems arise when attempts are made to show change over time using relative measures of single species. the increase or decrease in a particular species’ relative frequency is masked as an artifact of changes in other species. this situation is improved by lumping species into indicator guilds but the situation is not ideal in terms of showing change over time. area-based field sampling techniques, on the other hand, have historically been employed in the mapping and classification of vegetation and this is well documented. these survey techniques have involved recording all rooted plants present within a square or rectangular unit of varying sizes (termed a quadrat) and then assigning some estimation of percentage canopy cover to each species recorded in the quadrat using a cover class value. density, biomass, and frequency were not recorded in these units when the end product was a vegetation classification and map. what is not well documented is the use of this technique for the detection of vegetation change although the authors are aware of a number of ecologists who advocate and use this technique to monitor vegetation as part of a managerial action on game reserves and game ranches. the lack of the largescale use of an area-based sampling technique to detect vegetation change is because it is time consuming since all species (woody plants, forbs and grasses) within a demarcated sampling unit have to be identified, recorded and quantified. thus, fewer sites are completed for a study than would be the case using a point-based technique. also, historically, many of these sampling sites were chosen subjectively, not geo-referenced and the measure of cover-abundance used to quantify the species present in the sampling plot was, at best, a subjective estimate. areabased sampling has been historically difficult to repeat and does not readily lend itself to statistical analyses. data collected following the braun-blanquet approach, has been shown to be of use in assessing veld condition (westfall et al. 1983; eckhardt et al. 1993) and, more recently, a concerted effort has been made to make area-based sampling techniques more objective and thus of possible use in vegetation monitoring (westfall et al. 1996, westfall et al. 1997, panagos et al. 1998). this paper explores the ability (or lack thereof) of an area-based vegetation survey technique to detect change in the structure, composition and cover of the vegetation at the same sites with a five-year intervening period. study area the study area comprises the farm weltevreden 135 kr situated in the limpopo province, approximately 30 km east of vaalwater at 24º18's–24º23's, 28º08'e–28º13'e, and is approximately 700 ha in extent. the vegetation of the farm forms part of the savanna biome (rutherford & westfall 1986) and occurs in veld type 20 or sour bushveld (acocks 1988). the main physiographic features of the study area are: the southern edge of the sandriviersberge (2041 m a.s.l.) in the south; the groot wolwefontein river on the eastern boundary draining the farm in a northerly direction; and an undulating plateau (1080–1190 m a.s.l.) in the west and north. the geology of the farm forms part of the kransberg sub group of the waterberg group (2428 nylstroom 1:250 000 geological map). no detailed soil survey exists for the study area. however, the farm is included on the 1:250 000 nylstroom (2428) land type map koedoe 49/1 (2006) 70 issn 0075-6458 panagos & reilly.qxd 2006/04/17 10:32 pm page 70 and consists of two land types namely the bb and fa land types described by the land type survey staff (1988). the bb land type consists mainly of hutton and clovelly soil forms (deep sandy soils), while the fa land type consists mainly of mispah and glenrosa soil forms (shallow soils) (macvicar et al. 1977). climatic data for weltevreden, obtained from the agromet (1996) database, based on data recorded at the vaalwater weather station for 18 years, show that the area has a mean annual rainfall of 606 mm and a mean monthly maximum temperature of 29.7 ºc and a mean monthly minimum temperature of 3.6 ºc. methods the quantification of the vegetation characteristics commenced in 1996 when a base-line survey of the farm weltevreden 135kr was completed. three plant communities were identified using floristic data recorded in 12 quadrats (10 m x 20 m) that were placed in stratified units delineated using aerial photographs at a scale of 1:12 000. in february 2001, a monitoring programme, as defined above, was implemented with the re-survey of six of the original twelve sampling sites. the same fieldworker (1st author) solely collected field data during both time periods. these six sites were randomly chosen (two in each plant community) and located using previously recorded gps readings. site orientation was replicated with the use of a permanent marker and a compass bearing of the long side of the quadrat. floristic parameters recorded during the baseline survey were recorded again at each of the six sample sites, viz., the presence or absence of plant species within the quadrats, the assignment of growth forms to each species recorded (tree, shrub, dwarf-shrub, grass & forb) following westfall et al. (1996) and an estimate of canopy cover for each species recorded, in variably sized transects, based on the average crown to gap ratio (westfall & panagos 1988). plant taxa recorded in a sample site were identified using existing knowledge and field guides such as van oudtshoorn (1992), van wyk & malan (1988) and coates-palgrave (1988). plant species names or nomenclature is that used by the national herbarium, pretoria as described in germishuizen & meyer (2003). where a positive field identification was not possible, plants were assigned a field ‘nickname’, collected and later identified by a credible plant taxonomist. vegetation structure was classified according to edwards (1983) and is based on the cover and the height of the plants present in each community. data were analysed per community for change over time by comparing: net gain/loss of plant species in each quadrat within each growth form to detect an increase/decrease in plant species richness; change in canopy cover within each growth form in each qaudrat to detect changes in vegetation structure; and ratios of herbaceous plants (grasses and forbs) to woody growth forms in each quadrat. all three of these parameters can be used to determine the condition of the vegetation. in terms of statistical analyses for percentage canopy cover, a chi-square statistic was computed for each site in each community with the 1996 as the expected value and 2001 as the observed value (h0: no change over time). similarly, chi-square statistics were computed for all the species in all growth forms for the six re-surveyed sites from 1996 to 2001. although chi-square is idealy suited to use with frequencies it is applied in this case as a test of association or homogeneity. all analyses were done using statistica (stasoft inc. 1995). results the results of the follow-up survey are presented for each of the three plant communities identified on the farm weltevreden. 1. lannea discolor aristida meridionalis low closed woodland (community 1) there was a significant change in this community in terms of percentage canopy cover from 1996 to 2001 (χ2 = 15.63; df = 4; p = 0.0035 and χ2 = 89.16; df = 4; p = 0.00 for sites 1 and 3 respectively) (table 1). the number of woody species in community 1 remained more or less constant over the fiveyear period (table 2). however, the woody plant structure in terms of growth forms and canopy cover had changed considerably in site 3 from an open woodland with more shrubs and dwarf shrubs in 1996, to a closed woodland having fewer shrubs and dwarf shrubs and more trees in 2001 (table 1). site 1 shows a similar trend with an increase in the tree growth form and canopy cover. these observations are borne out by the test issn 0075-6458 71 koedoe 49/1 (2006) panagos & reilly.qxd 2006/04/17 10:32 pm page 71 koedoe 49/1 (2006) 72 issn 0075-6458 table 1 percentage canopy cover recorded for community 1 on the farm weltevreden in 1996 and 2001 sample site 1 1996 2001 sample site 3 1996 2001 trees 6.5 10.1 trees 12.3 36.4 shrubs 6.5 6.8 shrubs 20.5 0.5 dwarf shrubs 8.8 1.0 dwarf shrubs 2.9 0.2 grasses 28.6 15.4 grasses 7.3 16.6 forbs 0.4 0.9 forbs 0.6 2.8 total 50.2 34.2 total 43.5 56.5 table 2 change in species numbers per growth form in community 1 on the farm weltevreden between 1996 and 2001 sample site 1 sample site 3 number of woody plant species 1996 2001 trees 1 2 shrubs 2 5 dwarf shrubs 13 7 total number of woody 16 14 species & species turnover 3 ds 1 ds missing new number of grass species 1996 2001 annuals 1 1 perennials 10 11 total number of grass 11 12 species & species turnover 1 missing 2 new total number of forb 1996 2001 species & species turnover 7 16 1 missing 10 new total species for all 34 42 growth forms number of woody plant species 1996 2001 trees 3 11 shrubs 3 2 dwarf shrubs 10 2 total number of woody 16 15 species & species turnover 5 ds 3 t, 1 s, missing 1 ds new number of grass species 1996 2001 annuals 1 2 perennials 7 10 total number of grass 8 12 species & species turnover 0 missing 4 new total number of forb species 1996 2001 & species turnover 10 10 5 missing 5 new total species for all 34 37 growth forms statistics of each growth form in the significance test. the number of grass species remained relatively constant in site 1 but increased overall in site 3 by four grasses (one annual and three perennials) (table 2). grass cover declined by 13 % in site 1 but increased by 7 % in site 3 (table 1). forb species increased considerably in site 1 from seven species in 1996 to 16 species in 2001 but remained constant in site 3 (table 2). the percentage cover for forbs did not change much over the five-year period in either of the sample sites (table 1). this growth form (forbs) was mainly responsible for an increase in species richness in site 1 from 34 species in 1996 to 42 species in 2001. the total number of species in site 3 increased from 34 to 37. the ratio of forbs to other panagos & reilly.qxd 2006/04/17 10:32 pm page 72 species for these two sites was 38% (16/42) in site 1 and 27% (10/37) in site 3. the proportion of grasses to other species was 29 % and 32 % in sites 1 and 3 respectively. there was a significant change in this community in terms of number of plant species from 1996 to 2001 (χ2 = 19.94; df = 5; p = 0.0013 and χ2 = 30.35; df = 5; p = 0.00 for sites 1 and 3 respectively). 2. the rhus gracillima dichapetalum cymosusm low closed woodland (community 2) the woody species richness in community 2 remained relatively constant with an overall species decrease in site 5 and an increase in species in site 9 (table 4). change from 1996 to 2001 was significant in terms of perissn 0075-6458 73 koedoe 49/1 (2006) table 3 percentage canopy cover in community 2 on the farm weltevreden in 1996 and 2001 sample site 5 1996 2001 sample site 9 1996 2001 trees 1.0 12.7 trees 27.9 33.8 shrubs 11.0 0.0 shrubs 0.0 0.1 dwarf shrubs 0.5 1.4 dwarf shrubs 0.4 1.9 grasses 9.0 13.7 grasses 7.6 12.3 forbs 0.1 0.2 forbs 0.4 0.8 total 21.6 28.0 total 36.3 48.9 table 4 change in species numbers per growth form in community 2 on the farm weltevreden between 1996 and 2001 sample site 5 sample site 9 number of woody 1996 2001 plant species trees 3 4 shrubs 3 0 dwarf shrubs 6 6 total number of woody 12 10 species & species turnover 1 t, 1 s, 2 ds 2 ds missing new number of grass species 1996 2001 annuals 2 3 perennials 7 10 total number of grass 9 13 species & species turnover 2 missing 6 new total number of forb 1996 2001 species & species turnover 6 5 5 missing 4 new total species for all growth forms 27 28 number of woody 1996 2001 plant species trees 3 5 shrubs 1 2 dwarf shrubs 11 13 total number of woody 15 20 species & species turnover 2 ds 1 t, 1 s, missing 5 ds new number of grass species 1996 2001 annuals 1 4 perennials 7 9 total number of grass 8 13 species & species turnover 2 missing 7 new total number of forb 1996 2001 species & species turnover 3 12 1 missing 10 new total species for all growth forms 17 32 panagos & reilly.qxd 2006/04/17 10:32 pm page 73 centage canopy cover (χ2 = 152.06; df = 4; p = 0.00 and χ2 = 10.18; df = 4; p = 0.038 for sites 5 and 9 respectively) (table 3). scrutiny of the statistics shows that the change in the structure is disproportionate in the shrub layer; in essence the significant change can be almost entirely attributed to changes in this layer. tree cover increased in both sites, shrub cover declined in site 5 and remained constant in site 9. dwarf shrub cover increased in both sites (table 3). the number of grass species showed an overall increase in richness with four new grasses in site 5 (one annual and three perennials) and five new grasses in site 9 (three annuals and two perennials) (table 4). grass cover showed an overall improvement in both sites (table 3). forb cover increased slightly in both sites (table 3) and species richness in this growth form decreased in site 5 but increased in site 9 (table 4). the increase in grass and forb species in site 9 resulted in the total species richness for this site increasing from 17 species in 1996 to 32 in 2001 (table 4). the ratio of forbs to other species in site 5 was 18 % (5/28) and in site 9 was 38 % (12/32). the proportion of grasses to other species was 46 % and 41 % in sites 5 and 9 respectively. there was an insignificant change in this community in terms of number of species from 1996 to 2001 in site 5 (χ2 = 5.29; df = 5; p = 0.38) and a significant change in site 9 (χ2 = 39.27; df = 5; p = 0.00). 3. lannea edulis eragrostis pallens low closed woodland (community 3) species composition changed significantly in this community from 1996 to 2001 (χ2 = 153.3; df = 4; p = 0.00; and χ2 = 83.56; df = 4; p = 0.00) (table 6). the canopy cover measured for all five growth forms in this community did not change significantly during the five-year period (table 5). tree cover decreased slightly in both sites, shrub cover remained the same, dwarf shrub cover increased slightly and grass and forb cover remained relatively constant (table 5). however, the change in species richness was considerable in both re-surveyed sites (table 6). in site 11, species richness increased from 13 species in 1996 to 34 species in 2001, mainly in the grass and forb growth forms. grasses increased from three to 13 species and forbs increased from one to nine species. site 12 showed increases in all growth form categories: woody species increased from seven to 16 mainly due to eight new dwarf shrubs; grasses increased from six to nine species and forbs increased from four to seven species. the forb component in site 11 comprised 26 % (9/34) and in site 12, 16 % (7/45) of the species present were forbs. the proportion of grasses to other plant species was 38 % and 20 % in sites 11 & 12 respectively. once again the there was a significant change in this community in terms of number of species from 1996 to 2001 (χ2 = 71.13; df = 5; p = 0.00 and χ2 = 24.63; df = 5; p = 0.00 for sites 11 and 12 respectively). discussion the structure of the woody vegetation, particularly in communities 1 & 2 reflected a plant successional progression. in 1996, the structure of the vegetation in these two communities was dominated by plants in the shrub and dwarf shrub strata. these two communities had become tall closed woodlands dominated by more plants in the tree stratum i.e. the shrubs and dwarf shrubs had grown into trees. the tree cover in these two communities increased appreciably, in some cases more than double its original amount and the shrub and dwarf shrub strata decreased or remained constant, i.e. although these two growth forms were present in these two communities, their cover had been reduced. the tree cover in community 3 decreased slightly, while the cover for shrubs and dwarf shrubs increased in this community. from a management perspective, the increase in tree cover and decrease in shrub and dwarf shrub cover tends to ‘open up’ the woody vegetation. the maintenance of plants in the shrub or dwarf shrub strata often leads to an increase in density and ultimately bush thickening. bush thickening was not evident at the time of re-surveying, save for a small area in community 2 where koedoe 49/1 (2006) 74 issn 0075-6458 panagos & reilly.qxd 2006/04/17 10:32 pm page 74 terminalia sericea occurred at high densities. with the exception of one site (site 1, community 1) the grass cover increased throughout the farm, as did forb cover. the changes in plant cover and structure can probably be explained by the change in landuse (from cattle farming to game farming) as well as the high precipitation occurring during the period under review (edwards pers. comm.—edwards, charles, p.o. box 26739 monument park, pretoria, 0105). the increase in grass and forb cover can probably be attributed to the removal of cattle at high stocking rates and the introduction of game issn 0075-6458 75 koedoe 49/1 (2006) table 5 percentage canopy cover in community 3 on the farm weltevreden in 1996 and 2001 sample site 11 1996 2001 sample site 12 1996 2001 trees 31.8 29.6 trees 19.5 18.3 shrubs 0 0 shrubs 0 0.9 dwarf shrubs 0.1 2.7 dwarf shrubs 0.1 2.9 grasses 5.0 6.9 grasses 6.2 10.4 forbs 0.1 3.0 forbs 0.1 0.3 total 37.0 42.2 total 25.9 32.8 table 6 change in species numbers per growth form in community 3 on the farm weltevreden between 1996 and 2001 sample site 11 sample site 12 number of woody plant species 1996 2001 trees 4 4 shrubs 0 0 dwarf shrubs 5 8 total number of woody 9 12 species & species turnover 1 t, 4 ds 1 ds missing new number of grass species 1996 2001 annuals 1 3 perennials 2 10 total number of grass 3 13 species & species turnover 1 missing 11 new total number of forb species & species turnover 1996 2001 1 9 0 missing 8 new total species for all growth forms 13 34 number of woody plant species 1996 2001 trees 4 4 shrubs 0 1 dwarf shrubs 3 11 total number of woody 7 16 species & species turnover 0 ds 1 t, 7 ds missing new number of grass species 1996 2001 annuals 1 2 perennials 5 7 total number of grass 6 9 species & species turnover 0 missing 3 new total number of forb species & species turnover 1996 2001 4 7 1 missing 4 new total species for all growth forms 26 45 panagos & reilly.qxd 2006/04/17 10:32 pm page 75 including grazers, browsers and mixed feeders, at relatively low stocking rates. all six re-surveyed sites reflected an increase in plant species richness and this increase was most substantial in community 3 where the total number of plant species increased from 17 in 1996 to 34 in 2001 (site 11) and from 26 in 1996 to 45 in 2001 (site 12). species in the grass, forb and to a lesser extent dwarf shrub growth form categories were responsible for this large increase. the species in the trees and shrubs growth form categories remained constant or increased slightly. this trend, i.e. grasses, forbs and dwarf shrubs being responsible for most of the nett gain in species richness, was reflected in the other two communities as well but not as marked as in community 3. the proportion of forb species to other plant species in a given area is often an indication of the condition of the veld since most of these plants are opportunistic by nature. in other words, they occupy the space left by more permanent and often desirable plant species, which have been reduced in cover and frequency usually due to some environmental impact, either climatic or herbivory. the mean proportion of forbs to other plants (in terms of species richness) on weltevreden in 2001 was relatively high at 27 % and in combination with the low mean grass cover of 13 % (for the same year), taken in isolation this would be cause for concern. however, the mean grass cover for 1996 was only 10 % and when consideration is given to the low mean cover for forbs at 1.3 % (for 2001) and that the proportion of grass plants to the other species is high at 34 %, then it becomes evident that the veld is recovering from historical overgrazing. conclusions this study has shown that traditional areabased vegetation survey techniques (such as a braun blanquet approach) can be used for the detection of change in species composition if great care is taken with sample site relocation and species identification. these problems can be alleviated by the use of permanent demarcation of sample sites and the collection of voucher plant specimens. the canopy cover estimations using the plant number scale of westfall and panagos (1988) are an improvement compared to other frequently used cover-abundance scales, such as the braun-blanquet or the domin krajina scales (mueller-dombois & ellenberg 1974). this is largely due to the use of 33 cover classes in the plant number scale as well as a physical count of species in variable-sized transects based on average crown size and average gaps between individuals of the same species. the braun-blanquet cover scale makes use of only six classes and the domin krajina cover scale makes use of ten classes and observations are visually estimated usually from a sedentary position in the middle of the quadrat. nevertheless, the quantification of canopy cover using the plant number scale remains an estimation. this is due to problems associated with the choice of direction of the variable-sized transect in which the count of species is conducted and the definition of what constitutes an individual plant. future monitoring efforts using area-based techniques should make use of absolute abundances (such as a density count) rather than cover estimations. although this application shows some changes in the vegetation of the area, the technique’s analysis excludes estimation of data dispersion, hence precision cannot be calculated. much as point methods are susceptible to shortcomings in terms of adequate sample size (reilly & panagos 2001) area-based methods run the risk of pseudoreplication (hulbert 1984) and autocorrelation (fortin et al. 1989) and survey designs need to be scrutinised in this regard if they are to achieve their potential in terms of monitoring change over time. acknowledgements professor gerhard malan of the tshwane university of technology, for reviewing the manuscript prior to submission. koedoe 49/1 (2006) 76 issn 0075-6458 panagos & reilly.qxd 2006/04/17 10:32 pm page 76 references acocks, j.p.h. 1988. veld types of south africa. memoirs of the botanical survey of south africa 57: 1-146. agromet database. 1996. agricultural research council, institute for soil, climate and water, pretoria. coates palgrave, k. 1988. trees of southern africa. cape town: struik. dyksterhuis, e.j. 1949. condition and management of rangeland based on quantitative ecology. journal of range management 2: 104-114. eckhardt, h.c., n. van rooyen & g.j. bredenkamp. 1993. use of braun-blanquet data for the assessment of veld condition and grazing capacity in grassland. african journal of range and forage science 10(1): 41-46. edwards, d. 1983. a broad scale structural classification of vegetation for practical purposes. bothalia 14:705-712. everson, t.m., g.p.y. clarke & c.s. everson. 1990. precision in monitoring plant species composition in montane grasslands. vegetatio 88: 135-141. ferrar, a.a. 1983. guidelines for the management of large mammals in african conservation areas. pretoria: council for scientific and industrial research, national scientific programmes unit. (south african national scientific programmes report; no. 69.) foran, b.d., n.m. tainton & p. de v. booysen. 1978. the development of a method for assessing veld condition in three grassveld types in natal. proceedings of the grassland society of southern africa 13:27-33. fortin, m, p. drapeau & p. legendre. 1989. spatial autocorrelation and sampling design in plant ecology. vegetatio 83:209-222. germishuizen, g. & n.l. meyer (eds.). 2003. the plants of southern africa: an annotated checklist. strelitzia 14. hulbert, s.h. 1984. pseudoreplication and the design of ecological field experiments. ecological monographs 54(2): 187-211. land type survey staff. 1988. land types of the maps 2426 thabazimbi and 2428 nylstroom. memoirs of the agricultural natural resources of south africa no. 10. levy, e.b. & e.a. madden. 1933. the point method of pasture analysis. new zealand journal of agriculture 6:267-279. macvicar, c.n., j.m. de villiers, r.f. loxton, e. verster, j.j.n. lambrechts, f.r. merryweather, j. le roux, t.h. van rooyen & h.j. von m. harmse. 1977. soil classification: a binomial system for south africa. pretoria: soil and irrigation research institute, department of agricultural technical services. mentis, m.t. 1981. evaluation of the wheel-point and step-point methods of veld condition assessment. proceedings of the grassland society of southern africa 16:89-94. mentis, m.t. 1984. monitoring in south african grasslands. pretoria: council for scientific and industrial research, national scientific programmes unit. (south african national scientific programmes report; no. 91.) mentis, m.t., r.h.f. collinson & m.g. wright. 1980. the precision of assessing components of the condition of moist tall grassveld. proceedings of the grassland society of southern africa 15: 43 46. mueller-dombois, d. & h. ellenberg (eds.). 1974. aims and methods of vegetation ecology. new york: wiley. o’connor, t.g. & p.w. roux. 1995. vegetation changes (1949-71) in a semi-arid, grassy dwarf shrubland in the karoo, south africa: influence of rainfall variability and grazing by sheep. journal of applied ecology 32 (3): 612-626. panagos, m.d. 1995. a comparative classification of the sourish mixed bushveld on the farm roodeplaat (293 jr) using quadrat and point methods. msc thesis, university of natal, pietermaritzburg. panagos, m.d. & p.j.k. zacharias. 1995. research note: a time and motion study for two vegetation survey techniques. african journal of range and forage science 12(1): 81-84. panagos, m.d., k.m. van staden, r.h. westfall & p.j.k. zacharias. 1998. the plant communities of the roodeplaat experimental farm, gauteng, south africa and the importance of classification verification. south african journal of botany 64(1): 44-61. reilly, b.k. & m.d. panagos. 2002. statistical power of commonly used point methods in grassland monitoring. african journal of range and forage science 19:117-122. rutherford, m.c. & r.h. westfall. 1986. biomes of southern africa an objective categorization. memoirs of the botanical survey of south africa 54:1 98 short, a.d., t.g. o’connor & c.r. hurt. 2003. medium-term changes in grass composition and diversity of highland sourveld in the southern drakensberg in response to fire and grazing management. african journal of range and forage science 20(1):1-10. smit, g.n. 1989. quantitative description of woody plant communities: part i. an approach. journal of the grassland society of southern africa 6(4): 186-194. stasoft inc. 1995. statistica for windows. tulsa, ok: stasoft, inc. (http://www.stasoft.com) tainton, n.m., p.j. edwards & m.t. mentis. 1980. a revised method for assessing veld condition. issn 0075-6458 77 koedoe 49/1 (2006) panagos & reilly.qxd 2006/04/17 10:32 pm page 77 proceedings of the grassland society of southern africa 15: 37-42. trollope, w.s.w. 1990. development of a technique for assessing veld condition in the kruger national park using key grass species. journal of the grassland society of southern africa 7(1): 46-51. van oudtshoorn, f.p. 1992. guide to grasses of south africa. pretoria: briza. van wyk, a.e. & s.j. malan. 1988. veldgids tot die veldblome van die witwatersrand en pretoria gebied, insluitend magaliesberg en suikerbosrand. cape town: struik. westfall, r.h. & m.d. panagos. 1988. the plant number scale: an improved method of cover estimation using variable sized belt transects. bothalia 18:289 291. westfall, r.h., n. van rooyen & g.k. theron. 1983. veld condition assessments in sour bushveld. proceedings of the grassland society of southern africa 18:73-76. westfall, r.h., g.k. theron & n. van rooyen. 1997. objective classification and analysis of vegetation data. plant ecology 132:137-154. westfall, r.h., j.m. van staden, m.d. panagos, p.j.j. breytenbach & a. greeff. 1996. scale related vegetation sampling. pretoria: agricultural research council, range & forage institute (roodeplaat). zambatis, n. 2002. revised procedures for veld condition assessment in the kruger national park. unpublished report, scientific services, kruger national park, skukuza. koedoe 49/1 (2006) 78 issn 0075-6458 panagos & reilly.qxd 2006/04/17 10:32 pm page 78 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research vol. 50 no. 1 pp. 72 81 analysis of the vegetation of the sandstone ridges (ib land type) of the north-eastern parts of the mapungubwe national park, limpopo province, south africa abstract the establishment of the mapungubwe national park has been an objective of several conservationists for many years. the ultimate objective is that this park should become a major component of a transfrontier national park shared by botswana, zimbabwe and south africa. the aim of this study was to identify, classify and describe the plant communities present in the ib land type of the park. sampling was done by means of the braun-blanquet method. a total of 48 stratified random relevés were sampled in the ib land type. all relevé data were imported into a turboveg database, after which the numerical classification technique twinspan was used as a first approximation. subsequently, braun-blanquet procedures were used to refine data and a phytosociological table was constructed, using the visual editor, megatab. two plant communities and several subcommunities and variants were identified and described from the phytosociological table. keywords: braun-blanquet procedures, mapungubwe national park, phytosociology, savanna biome, vegetation classification the establishment of the mapungubwe national park (initially the vhembe-dongola national park) has been an objective of south african national parks for many years. the main motivation has been the rich biodiversity of the area, its great scenic beauty and the cultural importance of the archaeological treasures of mapungubwe. the artefacts discovered at this site rank among the most important pieces of ancient history yet found in sub-saharan africa (willcox 1966). the significance of this proposed park and its surrounding areas is further enhanced by the potential role of the area as a sanctuary for some of the most endangered mammals on earth, such as the black rhinoceros and the african wild dog (robinson 1996). one of the primary goals of nature conservation should be focused on preserving diversity and the persistence of species (walker 1989). when the natural habitats of plants have been destroyed, it is possible to cultivate and conserve plants in botanical gardens, but there is little of the genetic variation, as found between plants in their natural habitats, in botanical gardens (mcnaughton 1989). therefore, in order to conserve nature and its resources, it is important that nature reserves/ national parks are upgraded and managed well (brown 1997). nature reserves and national parks provide reservoirs of a country’s fauna and flora (greyling & huntley 1984; brown 1997). a sound knowledge of the vegetation ecology of areas of conservation significance is essential for the establishment of efficient wildlife and environmental management programmes and the compilation of conservation policies (bredenkamp & theron 1978; bredenkamp et al. 1993; bezuidenhout 1996). edwards (1972) stated that the optimal use of natural resources cannot be taken care of without adequate knowledge of the ecosystems involved. in the mapungubwe national park, the ib land type (figure 1) has the second highest plant diversity of all the land types in the park, with 219 plant species identified during the course of this study. there are a large number of microhabitats within each of the larger identifiable habitats, and these contribute to the high species diversity and complex nature of the ib land type. these microhabitats range from depressions in the rock surface, where soil that is transported by wind and water is trapped, to vertical face-rock habitats and cracks and seams in the mother rock. because of the complexity of the habitats within this land type, the classification of the resulting complex vegetation was a great challenge. the ib land type is scattered with signs of the ancient civilisation of the world-renowned mapungubwe hill. the mapungubwe cultural landscape is one of the unesco world heritage sites of south africa (huffman 2005). rock paintings and petroglyphs, primeval building sites and age-old potsherds are commonly found in the area. this heritage site is an extremely important site in the early history of southern africa, as it was here that social, cultural and political developments led to the first complex society in southern africa (huffman 2005, kuman et al. 2005). notwithstanding the unique vegetation of the albie r. götze sarel s. cilliers 1school of environmental sciences and development north-west university potchefstroom south africa hugo bezuidenhout arid ecosystem research unit conservation services south african national parks kimberley south africa klaus kellner1 correspondence to: sarel s. cilliers e-mail: sarel.cilliers@nwu.ac.za postal address: school of environmental sciences and development, section botany, private bag x6001, north-west university, potchefstroom, 2531, south africa 72 2008 vegetation of sandstone ridges original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 72 81 29°10'0"e 29°10'0"e 29°20'0"e 29°20'0"e 29°30'0"e 29°30'0"e 22 °2 0' 0" s 22 °2 0' 0" s 22 °1 0' 0" s 22 °1 0' 0" s ± 0 2 4 6 81 km farm boundaries studyarea land type: lb395 shashe lim po po limpopo province botswana zimbabwe botswana zimbabwe limpopo province den staat den staat riedel riedel regina greefswald locality map area, the archaeological and cultural heritage is an important reason for the proclamation of the mapungubwe national park (robinson 1996). the aims of this study were to identify, classify and describe the plant communities of the study area based on plant species composition, environmental variables and anthropogenic influences. information generated by this study could be further used in conjunction with vegetation studies on all the other land types in the mapungubwe national park (götze 2002; götze et al. 2003) to identify and map ecologically sound management units on which the optimal management of the park could be based, including recommendations on restoration practices for degraded areas (götze 2002). study area the mapungubwe national park is centred on the confluence of the limpopo and shashi rivers in the limpopo valley in the limpopo province of south africa. it lies on the international borders between botswana, zimbabwe and south africa (robinson 1996) (figure 1). sandstone ridges were studied on the farms greefswald and riedel, because entrance was restricted to the other farms (figure 1), as they were not formally included in the park at the time the survey was done (götze 2002). the climate is semi-arid, with a long-term mean annual rainfall of 350 mm per annum (willcox 1966; robinson 1996). rainfall is highly variable and usually falls during the summer months between october and march. surface drainage is mostly in a northerly direction towards the limpopo river. none of the rivers in the area, including the limpopo, are perennial. the study area is between 300 m and 780 m above sea level. south of the limpopo river the topography tends to be flat with sandstone and conglomerate ridges and koppies. nearer the limpopo river the plains give way to a rugged, hilly terrain. according to van rooyen and bredenkamp (1996), the study area is situated in mopane bushveld, while acocks (1988) placed it in veld type no.15, also called mopane veld. the most recent vegetation map of south africa, lesotho and swaziland (vegmap) places the ib land type in the limpopo ridge bushveld in the mopane bioregion of the savanna biome (mucina et al. 2005, mucina & rutherford 2006). other vegetation studies done in the mapungubwe national park and the surrounding areas include a terrain and vegetation evaluation by the former south african defence force (south african defence force 1986a, 1986b), a study by o’connor (1991) of vegetation types in the limpopo-venetia nature reserve, and a vegetation survey by timberlake et al. (1999) of the maramani/tuli area, across the limpopo river in south-western zimbabwe. other studies in the musina area that included parts of the limpopo ridge bushveld were done at the messina experimental farm (dekker & van rooyen 1995) and the honnet nature reserve (visser et al. 1996). however, none of the previous studies focused specifically on the sandstone ridges of the ib land type. robinson (1996) stated that the park has numerous archaeological sites dating from the early stone age to the present. many of these sites, which are concentrated in the area of the confluence of the limpopo and shashi rivers, are of major importance and scientific value. of particular interest is the zhizo site (ad 700–900) on the farm schroda, and mapungubwe hill and the adjoining bambandyanalo (ad 1100–1250), situated on the farm greefswald. according to voigt and plug (1981), the mapungubwe site is considered to be of major importance in sub-saharan africa and is the most remarkable iron age site in south africa. additional features of importance are the numerous rock paintings and petroglyphs found in the area. methods with the aid of different maps of the area, such as topographical, geological, soil and existing vegetation maps and aerial figure 1 location of the ib land type in the mapungubwe national park 73 original research götze, cilliers, bezuidenhout & kellner vol. 50 no. 1 pp. 72 81koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za photographs, a preliminary evaluation of the study area was made. in conjunction with this, a detailed reconnaissance of the area was done to determine homogeneous areas in the vegetation before sampling commenced. using visual estimates, sampling was done by means of the braun-blanquet method (muellerdombois & ellenberg 1974). land types were used to stratify the study area. other land types that were studied included the ia (götze et al. 2003), db, fb and ae land types (götze 2002). a total of 48 stratified random relevés, each covering an area of 50 x 50 m, were sampled. plant nomenclature follows germishuizen et al. (2006) and soil classification was done in accordance with the soil classification work group (1991). trees and shrubs were classified using the guidelines set by edwards (1983). trees were classified as rooted, woody, self-supporting plants over 2 m high and with one or a few definite trunks, and shrubs were classified as rooted, woody, self-supporting, multi-stemmed or singlestemmed plants less than 2 m in height (edwards 1983). the habitat conditions, including geology, soil forms and other edaphic factors, slope, aspect and the rockiness of the soil surface were qualitatively described for each relevé. this information was used in the description of the different plant communities. the coordinates of each relevé were determined with the use of a gps and noted. all relevé data were imported into the turboveg database (hennekens 1996a). the numerical classification technique twinspan (hill 1979a), which is regarded by several phytosociologists (mucina & van der maarel 1989; bredenkamp & bezuidenhout 1995; cilliers 1998) as a successful approach for classification, was used as a first approximation to classify the floristic data. subsequently, braun-blanquet procedures (bezuidenhout et al. 1996) were used to refine data and construct phytosociological tables (table 1) using a visual editor, megatab (hennekens 1996b). using the phytosociological tables and the habitat information gathered during the sampling period, the different plant communities were identified and described. results and discussion classification the ib395 land type (figure 1) contains a unique habitat that consists of rocky koppies with shallow lithosols of the mispah and glenrosa soil forms. the terrain occupied by this land type is made up of rugged crest areas (terrain unit 1 = tu1), vertical cliffs (tu2), foot slopes (tu3) and drainage areas at the very foot of the sandstone koppies (tu5). the geology is made up of sandstone and siltstone of the clarens formation of the karoo sequence (land type survey staff 2000). the following vegetation types were identified: 1. the hexalobus monopetalus – croton gratissimus var. subgratissimus community 1.1 the psydrax livida – oropetium capense subcommunity 1.1.1 the mundulea sericea – zornia glochidiata variant 1.1.2 the strychnos madagascariensis – coleochloa pallidior variant 1.2 the ficus tettensis – waltheria indica subcommunity 1.2.1 the stipagrostis uniplumis – pavonia dentata variant 1.2.2 the hibiscus coddii – tragia rupestris variant 2. the terminalia prunioides – grewia bicolor community 2.1 the acacia senegal – barleria senensis subcommunity 2.2 the colophospermum mopane – vernonia cinerascens subcommunity 2.3 the hermbstaedtia odorata – heliotropium ovalifolium subcommunity description of the plant communities 1. the hexalobus monopetalus – croton gratissimus var. subgratissimus community the hexalobus monopetalus – croton gratissimus var. subgratissimus community has a high plant species diversity, with more than 152 species recorded during this study. the community consists of a complex mosaic of microhabitats on rocky outcrops in shallow sandy mispah and glenrosa soil forms. this community is found mainly on the farm greefswald in the mapungubwe national park. large areas of bare sandstone are common and constitute between 30 and 70% of the total surface area. the community is moderately grazed, mostly by small wildlife types. this is due to the fact that the largest part of this rocky community is inaccessible to most large mammals. the community is home to a healthy population of the nimble klipspringer (oreotragus oreotragus), which feeds on herbs and shrubs on or at the base of rocky outcrops (wynne-jones 1982). apart from natural weathering of sandstone and natural erosion of the resulting sand particles, no other erosion of a serious nature was observed in the area. the diagnostic plant species of the hexalobus monopetalus – croton gratissimus var. subgratissimus community are found in species group a (table 1). these include the tree species hexalobus monopetalus var. monopetalus, croton gratissimus var. subgratissimus, ficus abutilifolia and philenoptera violacea, and the shrub species abutilon grandiflorum and ochna inermis. the diagnostic grasses are a trichoneura sp., schmidtia pappophoroides, aristida scabrivalvis subsp. scabrivalvis and brachiaria nigropedata, and the other herbaceous species include the sedge species bulbostylis hispidula subsp. pyriformis and the forbs evolvulus alsinoides, cleome hirta and barleria affinis (species group h, table 1). this community is further differentiated from the terminalia prunioides – grewia bicolor community (2) by the absence of the species of species group i (table 1). the average tree height of this community is 3.3 m and the canopy cover varies from 10 to 50%. the shrub layer has an average height of 1.3 m and a canopy cover of between 6 and 60%. the herbaceous layer varies in canopy cover from 30 to 71%, with an average height of 0.5 m. two subcommunities and four variants were identified in this community. the subcommunities are as follows: 1.1 the psydrax livida – oropetium capense subcommunity 1.2 the ficus tettensis – waltheria indica subcommunity 1.1 the psydrax livida – oropetium capense subcommunity this subcommunity is found almost exclusively on the mispah soil form and, in a few cases, on the glenrosa soil form, mostly on the crest (tu1) and to a lesser extent on cliff sides (tu2) and foot slopes (tu3) of the ib land type. shallow soils with a soil depth of up to 20 cm are found in depressions in the rock surface. herbaceous species and small shrubs are found in these depression microhabitats. deeper soils occur in run-off areas, crevices and seams in the rock, where a larger amount of soil has accumulated with a soil depth of up to 50 cm, which is more suitable for plant growth. grazing in the area is concentrated in the run-off areas, which are more accessible to game. it was also observed that these latter areas are utilised by wildlife for shelter. a common microhabitat occurring in this subcommunity is small depressions in the mother rock, which are filled by wind and water-transported soil and in which small herbaceous species have become established. the grass species oropetium capense (species group b, table 1) and the sedge bulbostylis hispidula subsp. pyriformis (species group a, table 1), together with some other small herbs and grasses, occur in these depressions. 74 vegetation of sandstone ridges original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 72 81 table 1 phytosociological table of the vegetation of the ib land type in the north-eastern parts of the mapungubwe national park relevé no. 1 1 1 2 2 1 2 2 1 1 1 2 1 2 1 1 2 2 2 0 0 0 0 0 0 2 0 2 0 0 2 1 0 1 1 0 2 0 0 0 1 1 1 1 0 1 0 0 8 9 9 0 1 9 0 0 9 9 9 0 9 0 8 8 1 0 1 6 7 6 6 5 7 0 6 0 7 0 0 8 7 8 9 6 0 0 0 0 9 8 8 9 5 3 0 6 8 6 9 1 2 7 0 2 2 3 0 3 8 4 7 3 1 6 0 3 0 5 8 7 4 5 6 8 6 2 7 4 5 9 5 9 9 7 8 9 1 5 6 4 1 9 1 7 communities 1 2 sub-communities 1.1 1.2 2.1 2.2 2.3 variants 1.1.1 1.1.2 1.2.1 1.2.2 species group a diagnostic species of the hexalobus monopetalus-croton gratissimus var. subgratissimus community (1) trichoneura species + + 1 1 a 1 1 a + + 1 a 1 1 a 1 1 1 a a a 1 b + b 1 b a . 1 a . . . . . . . . . . . . . . . . . bulbostylis hispidula subsp. pyriformis + + + + + + + 1 + + + 1 + + + + + + 1 1 a a a + . + 1 + 1 a + . . . . . . . . . . . . . . . . . hexalobus monopetalus var. monopetalus . + 1 + 1 1 + a + 1 a 1 a 1 + 1 . 1 1 a b 1 1 a . a b 1 1 . 1 . . . . 1 . . . . . . . . . . . . croton gratissimus var.subgratissimus . 1 a . a a . a . a a a . a 1 1 a a b a + + a . b a b a b a a . . . . . . . . . a . . . . . . . schmidtia pappophoroides + . a + b . + 1 1 . + 1 . . + + . . . + 1 b + a 1 1 1 a . . a . . . . + . . . . . . . . . 1 . . aristida scabrivalvis subsp. scabrivalvis + + . . 1 1 + . . + + b + 1 . 1 a a . . + . + 1 + a 1 . . a 1 . . . . . . . . . . . . . . . . . brachiaria nigropedata . 1 . a . a 1 + a . 1 . + 1 . . 1 . 1 + + . + + + 1 . . . + + . . . . . . . . . . . . . . . . . ficus abutilifolia . a 1 1 a a 1 . . a a . 1 1 1 1 . a 1 . 1 1 1 1 . . r 1 . . a . . . . . . . . . . . . . . . . . evolvulus alsinoides + . . . + + + + . . + . . . + + + + . + . . + + + + . + + . . . . + . . . . . . + . + . . + . . cleome hirta . . + . . . . 1 + . . . . + . . . . + . 1 1 . . 1 . + . . . + . . . . + . . . . + . . . . . . . abutilon grandiflorum 1 . a . 1 . . a a . . + . . . . + . . + + b . . + + . + + . 1 . . . + . . . . . 1 . . . . . . . ochna inermis . + . . 1 . . . . + . . . + + + . 1 1 . + . + . 1 + + . . + . . . 1 . . . . . . . + . a . . . . philenoptera violacea + . . . 1 . . . . . 1 . . . . 1 . . . . . + . + . a . . . . a . . . . 1 . . . . . . . . . . . . species group b diagnostic species of the psydrax livida-oropetium capense subcommunity (1.1) oropetium capense + 1 + + + + + + + + + + 1 . 1 + + + + . . . . . . + . + . . . . . . . . . . . . . . . . . . . . hibiscus engleri + 1 1 1 1 + a 1 1 1 1 . + + . + 1 . 1 . . . . . . 1 . 1 . . a + . + . . . . . . . . . 1 . . . . psydrax livida . 1 a 1 + a 1 1 a a 1 . 1 . 1 . . a 1 . . . . . 1 . + . . . . . . . . . . . . . . . . . . . . . sida cordifolia . + 1 . + + . 1 + 1 . 1 + + . . + 1 + . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . tephrosia euprepes + 1 . a . . . . 1 a 1 a 1 a + 1 + a . . . . + 1 . . . . . . . + . . . . . . . . . . . . . . . . croton pseudopulchellus . . . a . . a + . a . . 1 a + + . . . . 1 . . . . . . a . . . . . . . . . . . . . . . . . . . . species group c diagnostic species of the mundulea sericea-zornia glochidiata variant (1.1.1) mundulea sericea a b a a b a a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . zornia glochidiata + + . . + 1 + + . . . . . . . . . . . + + . . + + + . . . . . . . . . . . . . . . . . . . . . . brachylaena huillensis . . 1 + . a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . sclerocarya birrea subsp. caffra . . . 1 . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . 1 . . . . . . b . . + . eragrostis viscosa 1 . . + . . . . . . . . . . . . . + . . + . . . . . . . . . . . . . . . . . . . 1 . . . . . . . species group d diagnostic species of the strychnos madagascariensis-colechloa pallidior variant (1.1.2) coleochloa pallidior . 1 . + . . . 1 a + 1 + a a + + b a 1 . . . . . . . . . . . . . . + . . . . . . . . . . . . . . corchorus kirkii 1 1 . . . . . + + + . + + . 1 + 1 + + + . . . a . + + . + . . . . . . . . . . . . . . . . . . . indigofera laxeracemosa . . . . . + + + + + + . . + 1 + + . . . + . + . . 1 . . . 1 . . . . . . . . + . . . . . . . . . digitaria eriantha + . . . + . 1 . + 1 . . 1 1 . . 1 1 . . . . . 1 . 1 . . . . . . . . . . . . . . . 1 . . . a . . euclea natalensis subsp. angustifolia . . . . . . . . . + + . . + . . + + + . + . . + . . . . . . . . . . . . . . . . . . . . . . . . xerophyta retinervis . . . a . . + . . + + . + . . 1 . + . . a . . . . . . . . . . . . . . . . . . . . . . . . . . . pappea capensis . . . . . 1 . a a . . . . . + . . . a . . . . 1 . . . . 1 . . . . . . . . . . . . . . . . . . . pogonarthria squarrosa . 1 . . . . . . . . 1 . . . . 1 . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . species group e common species of subcommunity 1.1 and variant 1.2.1 eragrostis lehmanniana var. lehmanniana a a 1 + . + + + 1 1 1 + a + a b + a + 1 + 1 1 . . + . . . . . + . . . . . . . . a . . . . . . . aristida junciformis subsp. junciformis . a . a 1 . a a + 1 1 + + a + + 1 + a 1 + . + . . 1 . . . . . . . . . . . . . . . . . . . 1 . . melinis repens subsp. repens 1 + b 1 . 1 a . 1 1 + + . . + 1 . + . + 1 + a a . + + . . . . . . . . . . . . . 1 1 + . . 1 . . strychnos madagascariensis 1 . . . . . 1 + . 1 a a 1 + a a . + . . 1 . 1 b . . . . . . . . . . . . . . . . . . . . . . . . tephrosia rhodesica var. rhodesica . 1 . 1 . a 1 . . . 1 . 1 + . . 1 + + . + . a + . + . . . . . . . . . . . . . . . . . . . . . . hemizygia petrensis . 1 . + . a + . . . + + + + . + 1 + . + + . . . . . . . . . . . . . . . + . . . . . . . . . . . tephrosia virgata . . . . + . . . . 1 . . . . + . . 1 + 1 + . . . . . . + . . . . . . . . . . . . . . . . . . . . species group f diagnostic species of the ficus tettensis-waltheria indica subcommunity (1.2) ficus tettensis . . . . . 1 1 1 + . + . . . . . . . 1 1 + 1 . + 1 a . a a . a . . . . . . . . . . . . . . . + . waltheria indica 1 . . . + . + + . . . . . + . . . . + 1 + + . + + + . + + + . . . . . + . . . . 1 . . . . . . . sida ovata 1 . . . . . . . . . . . . . 1 + . . . + + 1 + . + . 1 . . 1 . . + . . + . . . . . . . . . + . . hibiscus species . . . . . . . . . . . . . . . . . . . + 1 . . . 1 . a . + 1 . . . . . . . . . . . . . . . . . . momordica balsamina . . . . . . . . . . . . . . 1 + . . . 1 . 1 . . . + . . . + + . + + . . . . . . . . . . . . . . species group g diagnostic species of the stipagrostis uniplumis-pavonia dentata variant (1.2.1) stipagrostis uniplumis var. uniplumis . . + + . . + . . . . 1 + + . . . . . 1 a + + + . + . . . . . . . . . . . . . . + . . . . . . + pavonia dentata . 1 . . . . . . . . . . . . . . . . . + + . 1 1 . . . . . . . . . . . . . . . . . . . . . . . . acrotome inflata + . . . . + . . . . . . . . . . . + . . + . + + . . . . . + . . . . . . . . . . + . 1 . . . . . combretum microphyllum . . . . . . . . . . . . . . . . . . . 1 . b . . 1 . . . . . . . . . . 1 . . . . . . . . a . . . lannea schweinfurthii var. stuhlmannii . . . . . . . . . . 1 . . . . . . . . . + 3 . . . . . . . . . . . . . . . . . . . . . . . . . . blepharis subvolubilis . . . . . + . . . . . . . . . . . . . . + . . a . . . . . . . . . . . . . . . . . . . . . . . . species group h diagnostic species of the hibiscus coddii-tragia rupestris variant (1.2.2) hibiscus coddii . . . . a . . . + . . . . . . . . . a . . . . . . a + a a 1 1 . . . . . . . . . . . . . . . . . tragia rupestris . + + . . . 1 + . . . . . . . 1 . . . . . . . + + + . + + + + . . . . . . . . . . . . . . . . . barleria affinis . . . . . . . + . + . . . . + . . . . . . . . . . a a . 1 . . . . . . . . . . . . . . . . . . . sterculia rogersii . . . . . . . . . . . . . . . . . . . . . . . . + . . + 1 . . . . . . . . . + . . . . + . . . . commelina africana . . . . . . . . . . . . . . . . . . . . . + . . + . . . a . . . 1 . . . . . . . . . . . . + . . species group i diagnostic species of the terminalia prunioides-grewia bicolor community (2.) grewia bicolor . . . . . . . . 1 1 1 . . . . + . . + . + . . . + . . + 1 . . 1 a 1 a 1 a . 3 1 a a 1 1 1 1 + a terminalia prunioides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b + a a + 1 + a a . b . a a 1 a 1 commicarpus pentandrus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + + 1 1 . + . + . + . + . 1 . a commiphora tenuipetiolata . . + . . . . . . . . . . . . . + + . . . . . . . . . . . . . 1 . 1 b . a . 1 1 . 1 . + + a . b commiphora glandulosa . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . a . 1 . . . b a . 1 . 1 . . . 1 ptycholobium contortum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b . + + . . . + . . 1 . . . 1 . 1 aristida congesta subsp. congesta . . . . . . . . . . . . . . . . . . . . . . . + . . . + . . . + . . + . + . . . . a . . . . + b acacia nigrescens . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . 3 . . . + . . + . . . . b species group j common species of subcommunities 2.1 and 2.2 tephrosia purpurea . . . . . . . . . + . . . . 1 . . . . . . . . . . . . . . . . a . a a . a . . 1 1 a a a . b . . dicoma tomentosa . . + . . . . + . . . . . . + . . . . . . . . . + . . . . . . a . 1 1 . a . a 1 a a a . . 1 . . leucas sexdentata . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . + + 1 1 . 3 1 1 . + . 1 . . . . 75 original research götze, cilliers, bezuidenhout & kellner vol. 50 no. 1 pp. 72 81koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za relevé no. 1 1 1 2 2 1 2 2 1 1 1 2 1 2 1 1 2 2 2 0 0 0 0 0 0 2 0 2 0 0 2 1 0 1 1 0 2 0 0 0 1 1 1 1 0 1 0 0 8 9 9 0 1 9 0 0 9 9 9 0 9 0 8 8 1 0 1 6 7 6 6 5 7 0 6 0 7 0 0 8 7 8 9 6 0 0 0 0 9 8 8 9 5 3 0 6 8 6 9 1 2 7 0 2 2 3 0 3 8 4 7 3 1 6 0 3 0 5 8 7 4 5 6 8 6 2 7 4 5 9 5 9 9 7 8 9 1 5 6 4 1 9 1 7 communities 1 2 sub-communities 1.1 1.2 2.1 2.2 2.3 variants 1.1.1 1.1.2 1.2.1 1.2.2 kyphocarpa angustifolia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . + + . + . . . . + . + 1 1 . . aristida adscensionis . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . a . 1 . . . . + + . + . + . . . . hibiscus micranthus var. micranthus + . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . + . . + . . . + + + . . + . . acacia senegal var. rostrata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . + . . . . . 1 . . . 1 . . abutilon sonneratianum . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . + . . + + + + . . . . . . . . kirkia acuminata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b . . . 1 . . . . a . . 1 . . . species group k diagnostic species of the acacia senegal-barleria senensis subcommunity (2.1) acacia senegal var. leiorhachis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + 1 + 1 + . . . . . . . . . . . barleria senensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + + + 1 + . . . . . . . . . . + cordia grandicalyx . . . . . . . . + . . . . . + . . . . . . . . . . . . . . . . . + . 1 + . . . . . . . . . . . . abutilon austro-africanum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . + + . . + . . . + . . . . . + aristida stipitata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1 . . . . . . . . a . . . . megalochlamys hamata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1 . . . . . . . . . . . . petalidium aromaticum var. aromaticum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . 1 . . . . . . . . . . . . commiphora mollis . . . . . + . . . . . . + . . . . . . . . . . . . . . . . . . . 1 . . . 1 . . + . . . . . . . . species group l diagnostic species of the colophospermum mopane-vernonia cinerascens subcommunity (2.2) colophospermum mopane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a a a . a b . . a . . vernonia cinerascens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . + + . 1 . . . rhigozum zambesiacum . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . + . . . . . + . . . 1 1 . . 1 . . corchorus asplenifolius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . + + . . . + . . cenchrus ciliaris . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . a . 1 . . . boscia foetida . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . a + . . . . . . + . . species group m common species of subcommunities 1.2, 2.1 and 2.2 ceratotheca triloba . . . . . . . . . . . . . . . + + . . 1 + + + + + + 1 . . . . + . + + . 1 . + + . + . 1 + a . . gisekia pharnacioides var. pharnacioides . . . . . . . . . + . . . . . . . . . + . + + + 1 . a . . + . . a . . 1 . + + + . . . . + . 1 . brachiaria deflexa . . . . . . . + . . . . . . + . . . . . . + . . . . + . 1 . + . + . . + . . + 1 . + + . a . + . phyllanthus loandensis . . . . . . . . . . . . . . + . . . . . . + + . . + . . . . . . . . + . . . + . . + . + . + . . tephrosia polystachya var. polystachya . . . . . . . . . . . . . . . . . . . 1 . + . . + . + . . . . . a . . 1 . . . . . . . . 1 . . . species group n common species of community 1 and subcommunities 2.1 and 2.2 combretum apiculatum subsp. apiculatum 1 + 1 . 1 a a a a a . a 1 a + a a a 1 . + . a b . 1 . 1 . + 1 a b b + 1 a . a 1 a 1 a b + 3 . . hermannia glanduligera + . 1 . 1 + + . + a . 1 . . + + . + . + + 1 1 + 1 + + 1 1 . a + + + 1 + 1 . . . + + . 1 . . . . acalypha indica + + 1 . + . . + . + . . . + + . + . 1 . + 1 . . . + . 1 + . + 1 . 1 + + 1 + . . . + + + a + . . panicum maximum b + . + . a 1 + 1 + 1 + . 1 1 1 . 1 + 1 1 a 1 . . 1 1 1 . . 1 . . 1 . a 1 + + . a . a 1 . 1 . . albizia harveyi + 1 . 1 . . a . . 1 1 b + 1 . + + . . 1 a . . b + . b . . . . 1 . + 1 + + . . . . . . 1 b . . . grewia flavescens . . + . + . . 1 a + . . 1 + . + + . . . 1 a . . . . . + a . 1 + . + + a 1 . . + 1 + . . . + . . cucumis zeyheri + . + . . + + + + + . 1 . . + + + . . + . . . . + . 1 + . . . + 1 + + a . . + . + 1 + + . + . . commelina benghalensis + . . . 1 . . 1 + . . . . . + . . . + . 1 a . . a + . 1 . . 1 . . + + 1 . . + . . 1 + + . . . . achyranthes aspera var. aspera + . . . . . . + . . . . . . + . . + + . . . . . . . . . 1 . . . + + . 1 + . . . + . . + . + . . dichrostachys cinerea subsp. africana a . . . . + + . . . . a . . . . . . . + . a . + . . . . . . . . . . . 1 . . + . a + . . . . . . commiphora edulis subsp. edulis + . . . . + . . . + + . . . . . . . . . . . 1 . a + . . . . . . . + . . . . . . . . . a . . . . calostephane divaricata + . + . . 1 . . . + + . . . . . . . . . . + . . . . . . . . . . . . . + . . . . . + . + . + . . acacia erubescens b . . . . . . . . a . . . . a + . . . . . . . . . . . . . . . a . . + . . . . . . . a . . 1 . . species group o common species of subcommunity 1.2 and community 2 hermbstaedtia odorata + . . . . . . . . . . 1 . . . + . . . . . 1 . . . . . . . + . . + . . + . . . . + + . . . . + + heliotropium ovalifolium . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . 1 + . . 1 + + urochloa mosambicensis . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . 1 . . . . . . . . . . . . . b . cordia monoica . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . b tribulus zeyheri subsp. zeyheri . . . . . . . . . . . . . . . . . . . . . + . . + . . . . . . . + . . . . + + + . . . . + . a . cleome angustifolia subsp. petersiana . . . . . . . . . . . . . . . . . . . . . + . + . . + . . . . . 1 . . + . + + . . . . . . . . 1 species group p common species of all the communities enneapogon cenchroides + + 1 . 1 + + 1 1 + + 1 + + + + + + 1 + + a + + + + a 1 a 1 1 a 3 + 3 a 3 3 3 3 + b . + a b a a boscia albitrunca 1 . + . 1 1 1 a a 1 . . + + 1 . . + + 1 + + a a a 1 a 1 1 + + 1 1 1 1 + 1 . + a . . . + . 1 1 1 solanum tettense var. renschii + 1 1 . . 1 1 + 1 1 . + . . + + + + . + . + . + . . + . + . + 1 a 1 + a . 1 a + + . + 1 a 1 . + cadaba termitaria . . . . . + . . . . . . . . + + . . + . . . . 1 1 . . . . . . . . . 1 a . . . . . . 1 . + a . + seddera suffruticosa . + . . . . . . . + + . . . + . . . . . + . . . . . . . . . . . . . . + . + . + + + . . . + . 1 eragrostis trichophora . 1 . . + . . . . . . . . . 1 a . . . . + . . . 1 . . . . . . . . . . . . . 1 + . . . . . . 1 . companion species *xanthium strumarium + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pellaea calomelanos var. calomelanos . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . combretum mossambicense . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pupalia lappacea var. lappacea + . . . + . . . + . + . . . + . . . . . . . . . . + . . . . . . . . . + . . . . . + . . . . . . blepharis aspera . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . aloe chabaudii var. chabaudii . . . + . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . flueggea virosa subsp. virosa . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xerophyta humilis . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . barleria species . . . . . . . . . + . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . plicosepalus kalachariensis . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pycreus pelophilus . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . euphorbia cooperi var. cooperi . . . + . . . . . . + . . . . . . 1 . . . . . . . . . . . . . . . + . . + . . . . . . . . . . . ficus salicifolia . . . . . . . . + . . . . . . . . a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . sericorema sericea . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . xerophyta viscosa . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . leonotis ocymifolia . . . . . . . + . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . abrus precatorius subsp. africanus . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . maerua angolensis . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . schizachyrium jeffreysii . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . oxygonum delagoense . . . . . + . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cleome monophylla . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . geigeria burkei subsp. fruticulosa . . . . . + . . . . . . . . . 1 . + . . . . . . . . . . . . . . . . + . . . + . . . . . . . . 1 76 table 1 (cont...) vegetation of sandstone ridges original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 72 81 relevé no. 1 1 1 2 2 1 2 2 1 1 1 2 1 2 1 1 2 2 2 0 0 0 0 0 0 2 0 2 0 0 2 1 0 1 1 0 2 0 0 0 1 1 1 1 0 1 0 0 8 9 9 0 1 9 0 0 9 9 9 0 9 0 8 8 1 0 1 6 7 6 6 5 7 0 6 0 7 0 0 8 7 8 9 6 0 0 0 0 9 8 8 9 5 3 0 6 8 6 9 1 2 7 0 2 2 3 0 3 8 4 7 3 1 6 0 3 0 5 8 7 4 5 6 8 6 2 7 4 5 9 5 9 9 7 8 9 1 5 6 4 1 9 1 7 communities 1 2 sub-communities 1.1 1.2 2.1 2.2 2.3 variants 1.1.1 1.1.2 1.2.1 1.2.2 merremia pinnata + . . . . . . . . . . . . + . . + . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . vigna species . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vigna frutescens subsp. frutescens + . . . . . . . . . . . . . . . . . . . . 1 . . . + . . . . . . . . . + . . . . . . . . . . . . sesamum triphyllum var. triphyllum . . . . . . . . . . . . . . . . . . . . . + + . + . . . . . . . + . . + . . . . . . . . . . . . cyperus rupestris var. rupestris . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . marsdenia sylvestris . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . indigofera charlieriana var. charlieriana . . . . . . . . . . . . . . . . . . . r + . . . . . . . . . . . . . . . . . . . . . . . . + . . vangueria infausta subsp. infausta . . . . . . . . . . . + . . . + . . . . . . + a . . . . . . . . . . . . . . . . . . . . . . . . cassia abbreviata subsp. beareana . . . . . . . . . . . . . . + . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . monechma divaricatum . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . panicum schinzii . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . bridelia mollis . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . steganotaenia araliacea var. araliacea . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . ctenolepis cerasiformis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . monsonia senegalensis . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . 1 . . + . . . . . . . . . . . . . nemesia fruticans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . ocimum americanum var. americanum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . kohautia aspera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . hermannia modesta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . grewia tenax var. tenax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a . . . . . . . . . . . . . hibiscus sabiensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . melhania rehmannii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . + . . . . . . + . . . . . . setaria sagittifolia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b . . . . . . . . . . . . panicum deustum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a . . . . . . . . . . . . justicia protracta subsp. rhodesiana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . decorsea schlechteri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . + . . . . . . . . + . . . asparagus africanus . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + 1 . . . . . . . . . . . grewia occidentalis var. occidentalis . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . + . . . . + . . . . . . . . . + ipomoea sinensis subsp. blepharosepala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . acacia mellifera subsp. detinens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + a . . . . . . . . loudetia simplex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . commiphora viminea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . euphorbia species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . neuracanthus africanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . tinnea rhodesiana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . chamaecrista absus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + + . . . . . . . adansonia digitata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b . . . . b . . . . . . . vahlia capensis subsp. vulgaris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a . . . . . . . acacia tortilis subsp. heteracantha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . + . . . cucumis metuliferus . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . crotalaria species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . + . . . . . . . *bidens pilosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . indigofera heterotricha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . pavonia burchellii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . grewia villosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . + + . . + . . + . . . . . . euphorbia eylesii . . . . . . . . . . . . . . . . . + . . . . . . . + . . . . . . . . . . . . . . . + + . + . . . crotalaria virgulata subsp. grantiana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b . . . . . indigofera trita subsp. subulata . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . 1 . . . . . merremia kentrocaulos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . ximenia americana var. microphylla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . ipomoea magnusiana . . . . . + . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . + . . . . . melhania acuminata var. agnosta + . . . . . . . . . . . . . . + . . . . + . . . . . . . . . . . . . . . . . . . . . + . . . . . dichilus lebeckioides + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . monsonia glauca . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . asparagus suaveolens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . 1 . . . balanites pedicellaris subsp. pedicellaris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a . . . asparagus laricinus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . barleria transvaalens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . berchemia zeyheri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . cadaba aphylla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . aristida rhiniochloa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . becium filamentosum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . bothriochloa insculpta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . *plumbago zeylanica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . aptosimum lineare var. lineare + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . amaranthus species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r + . . . . + . + . *alternanthera pungens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . *chenopodium carinatum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . chloris virgata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . dactyloctenium aegyptium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . eleusine coracana subsp. africana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . indigastrum costatum subsp. macrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . + . *schkuhria pinnata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . setaria verticillata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . tragus berteronianus . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . + . . . . . . + . enneapogon desvauxii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + indigofera circinnata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . + . . . . . . . . + indigofera species . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . + . . . . . . . . . . . . * exotic species 77 original research götze, cilliers, bezuidenhout & kellner vol. 50 no. 1 pp. 72 81koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za the diagnostic species of this subcommunity are found in species group b (table 1) and include the tree species psydrax livida and the shrubs croton pseudopulchellus and hibiscus engleri. the diagnostic species also include the grass species oropetium capense and the forbs sida cordifolia and tephrosia euprepes. this subcommunity is differentiated from the ficus tettensis – waltheria indica subcommunity (1.2) by the absence or infrequent occurrence of the species of species groups f and m (table 1). the tree species cover between 10 and 35% of the area, and have an average height of 3.0 m. the shrub layer has an average height of 1.3 m and the canopy cover varies between 6 and 26%. the average height of the herbaceous layer is 0.5 m and the canopy cover varies between 30 and 60% of the subcommunity area. two variants were identified in the psydrax livida – oropetium capense subcommunity: 1.1.1 the mundulea sericea – zornia glochidiata variant 1.1.2 the strychnos madagascariensis – coleochloa pallidior variant 1.1.1 the mundulea sericea – zornia glochidiata variant this variant is most commonly found on crests (tu1) and less commonly on foot slopes (tu3) in the psydrax livida – oropetium capense subcommunity (1.1). the dominant soil form is mispah, while glenrosa occurs to a lesser extent. the soils are shallow and sandy in texture. approximately 35% of the area is overlain by solid sandstone and the rockiness of the available soil surfaces is between 55 and 70%. this variant shows signs of moderate grazing because it is accessible to larger mammals in some instances. this variant occasionally was observed in the vicinity of natural seeps and rock pools filled with water during the rainy period. the diagnostic species of the mundulea sericea – zornia glochidiata variant are found in species group c (table 1). these include the tree species mundulea sericea, brachylaena huillensis and sclerocarya birrea subsp. caffra, the grass species eragrostis viscosa and the forb zornia glochidiata. this variant is differentiated from the strychnos madagascariensis – coleochloa pallidior variant (1.1.2) by the infrequent occurrence or absence of the species of species group d (table 1). large specimens of the tree species ficus abutilifolia are also common in this variant. the average height of the tree layer is 3.0 m and the canopy cover ranges from 10 to 35%. the shrub layer has an average height of 1.3 m and canopy cover varying between 10 and 25%. the herbaceous layer covers between 40 and 60% with an average height of 0.5 m. 1.1.2 the strychnos madagascariensis – coleochloa pallidior variant this variant makes up approximately 60% of the surface area of the psydrax livida – oropetium capense subcommunity (1.1). it occurs mainly on the undulating foot slopes (tu3) of the ib land type, but also to a lesser extent on the crests of the terrain (tu1). the main soil forms are sandy mispah and glenrosa soils. grazing levels seemed to be lower than in the mundulea sericea – zornia glochidiata variant (1.1.1), as this area is less accessible to large mammals. the plant diversity of this variant is marginally higher than that of the mundulea sericea – zornia glochidiata variant (1.1.1), probably due to the larger number of microhabitats found in this variant because of the more undulating nature of the terrain, which provides more microclimates than the open, flat terrain of the mundulea sericea – zornia glochidiata variant (1.1.1). it was noted that most of the larger tree species were found growing in seams and crevices in the sandstone. the rockiness of the soil surface varies between 50 and 75% and roughly 30% of the area consists of bare rock. diagnostic species include the tree species euclea natalensis subsp. angustifolia, which also occurs in shrub form, and pappea capensis, the grasses digitaria eriantha and pogonarthria squarrosa, the sedge coleochloa pallidior and the forbs corchorus kirkii, indigofera laxeracemosa and xerophyta retinervis (species group d, table 1). species with high cover abundance values are the tree species strychnos madagascariensis (species group e, table 1) and the shrubs croton pseudopulchellus (species group b, table 1) and grewia flavescens (species group n, table 1). the tree layer has an average height of 3.0 m and a canopy cover varying between 10 and 35%. the shrub layer covers between 6 and 26% of the variant area, with an average height of 1.2 m. the herbaceous layer reaches an average height of 0.5 m and the canopy cover varies between 30 and 60%. 1.2 the ficus tettensis – waltheria indica subcommunity this subcommunity of the hexalobus monopetalus – croton gratissimus var. subgratissimus community (1) occurs on deeper glenrosa and mispah soil forms than the psydrax livida – oropetium capense subcommunity (1.1). this subcommunity is situated on crests (tu1), steep cliff sides (tu2) and to a certain extent on the upper parts of the foot slopes (tu3) of the ib land type. the so-called cliff sides on these sandstone ridges seldom have a vertical drop. the drop usually forms small, sometimes narrow terraces. on the ledges formed by the terrace-like terrain, shallow to moderately deep soils have accumulated and serve as a growth medium for tree species (especially the wild fig species such as ficus abutilifolia, f. tettensis and f. salicifolia). the diagnostic species of this subcommunity are found in species group f (table 1). they include the tree species ficus tettensis and the forb species waltheria indica and sida ovata, hibiscus species and the climber momordica balsamina. the absence or infrequent occurrence of species group b (table 1), along with higher cover abundance values for the sedge species bulbostylis hispidula subsp. pyriformis (species group a, table 1), differentiates this subcommunity from the psydrax livida – oropetium capense subcommunity (1.1). the presence of the species of species group m (table 1) further differentiates this subcommunity from the other subcommunity (1.1) of the hexalobus monopetalus – croton gratissimus var. subgratissimus community (1). the tree layer has an average height of 3.8 m. this is on average 80 cm higher than the tree layer of the ficus tettensis – waltheria indica subcommunity. the tree cover of this subcommunity varies between 5 and 50%. the shrub layer reaches an average height of 1.5 m and has a varying canopy cover of 22 to 60%. the herbaceous layer covers between 35 and 70% of the area and has an average height of 0.6 m. two variants were identified in this subcommunity: 1.2.1 the stipagrostis uniplumis – pavonia dentata variant 1.2.2 the hibiscus coddii – tragia rupestris variant 1.2.1 the stipagrostis uniplumis – pavonia dentata variant this variant is most commonly situated on the crest (tu1) and foot slope (tu3) of the ib land type. the glenrosa soil form dominates the area occupied by the variant and the mispah soil form occurs less regularly. the areas in which this variant is found have deeper soils than those of the hibiscus coddii – tragia rupestris variant (1.2.2). the variant is minimally grazed and no severe signs of vegetation and soil degradation were observed. large portions of bare rock surfaces are present and cover approximately 35% of the area. the rockiness of the soil surface is between 30 and 40%. the diagnostic species of the stipagrostis uniplumis – pavonia dentata variant are found in species group g (table 1). they include the tree species lannea schweinfurthii var. stuhlmannii, the woody creeper combretum microphyllum, the grass stipagrostis uniplumis and the forbs pavonia dentata, acrotome inflata and blepharis subvolubilis var. subvolubilis. pavonia dentata is regarded as an endemic species to the limpopo ridge bushveld (mucina 78 vegetation of sandstone ridges original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 72 81 & rutherford 2006). other species that differentiate this variant are the shrub dichrostachys cinerea (species group n, table 1) and the forb ceratotheca triloba (species group m, table 1). the presence of the species of species group e (table 1) further differentiates this variant from the hibiscus coddii – tragia rupestris variant (1.2.2) the tree layer is well developed, with an average height of 4.0 m and a canopy cover of 15 to 50%. the shrub layer has a canopy cover ranging from 22 to 60% and an average height of 1.5 m. the herbaceous layer covers 35 to71% of the area and reaches an average height of 0.6 m. 1.2.2 the hibiscus coddii – tragia rupestris variant this variant is found on moderately deep and shallow soils on the crests (tu1) and cliff sides (tu2) of the ib land type. the main soil forms are glenrosa and mispah. the variant is often found on the verges of cliff sides and also on ledges and crevices on steep inclines of the sandstone ridges. grazing levels are low and the area as a whole is not seriously degraded. open rock surfaces cover about 40 to 50% of the area of the variant and the rockiness of the soil surface is between 20 and 30%. diagnostic species of this variant are found in species group h (table 1) and include the tree species sterculia rogersii, the shrubby hibiscus coddii and the forbs tragia rupestris and commelina africana. the absence of the shrub dichrostachys cinerea (species group n, table 1), as well as the species of species groups e and g (table 1) further differentiate this variant from the stipagrostis uniplumis – pavonia dentata variant (1.2.1). the tree layer has a canopy cover varying between 5 and 30%, with an average height of 3.5 m. the shrub layer reaches an average height of 1.3 m and has a canopy cover varying from 5 to 50%. the canopy cover of the herbaceous layer varies from 20 to 66%, with an average height of 0.5 m. 2. the terminalia prunioides – grewia bicolor community this community is found on shallow and moderately deep soils on the crests (tu1) and foot slopes (tu3) of the ib land type and occurs on the two farms greefswald and riedel in the mapungubwe national park. the common soil forms found in this community are shallow mispah and deeper glenrosa lithosols, as well as moderately deep hutton soils. the soils generally have a sandy texture and, as a result, are well drained. the largest portion of this community is underlain by sandstone as the mother rock, and a smaller portion is underlain by intrusions of dolerite. these dolerite areas are in the form of long, narrow intrusions, often cropping out in jagged hilly formations or rocky flats. approximately 10 to 30% of the area of the community is covered by exposed bedrock and between 20 and 50% of the available soil surface is covered by rocks. the area is moderately grazed and, in some places, is trampled by game. with the exception of the hermbstaedtia odorata – heliotropium ovalifolium subcommunity, the terminalia prunioides – grewia bicolor community is generally in a good condition. the diagnostic species of this community are found in species group i (table 1). these species include the tree species terminalia prunioides, commiphora tenuipetiolata, c. glandulosa and grewia bicolor, with the latter occurring mostly in shrub form. other diagnostic species are the grass species aristida congesta subsp. congesta and the bushy forb commicarpus pentandrus. the terminalia prunioides – grewia bicolor community (2) is differentiated from the hexalobus monopetalus – croton gratissimus var. subgratissimus community (1) through the absence or infrequent occurrence of the species of species group a (table 1). these two communities only have a few species in common, namely the general species of species group p (table 1). the tree layer of this community has a canopy cover of between 10 and 57% and an average height of 4.8 m. the shrub layer covers between 5 and 45% of the area, with an average height of 1.4 m, and the herbaceous layer has a varying canopy cover of 32 to 90% and an average height of 0.5 m. three subcommunities were identified in the terminalia prunioides – grewia bicolor community (2), namely: 2.1 the acacia senegal – barleria senensis subcommunity 2.2 the colophospermum mopane – vernonia cinerascens subcommunity 2.3 the hermbstaedtia odorata – heliotropium ovalifolium subcommunity 2.1 the acacia senegal – barleria senensis subcommunity this subcommunity is found on dolomite outcrops and on areas with a mixture of sandstone and dolomite on the crests (tu1) and foot slopes (tu3) of the ib land type on the farm greefswald in the mapungubwe national park. the soil forms that dominate this variant are glenrosa and mispah. the soils are shallow and have a sandy to loamy texture. approximately 30% of the surface area of the variant is covered by solid rock and the rockiness of the soil surface is between 30 and 45%. the area is moderately grazed and animals utilise bush clumps for shelter. diagnostic species are found in species group k (table 1) and include the woody species acacia senegal var. leiorachis, commiphora mollis and cordia grandicalyx, with the latter often occurring in shrub form. the diagnostic species further include the shrubby abutilon austro-africanum, the grass species aristida stipitata and the forbs barleria senensis, megalochlamys hamata and petalidium aromaticum var. aromaticum. other species that differentiate this variant from the colophospermum mopane – vernonia cinerascens subcommunity (2.2) and the hermbstaedtia odorata – heliotropium ovalifolium subcommunity (2.3) are the tree species albizia harveyi and the forbs hermannia glanduligera (species group n, table 1) and to a lesser extent tephrosia polystachya var. polystachya (species group m, table 1). the height of the tree layer of the acacia senegal – barleria senensis subcommunity averages 4.4 m, with a canopy cover of 30 to 57%. the shrub layer has an average height of 1.3 m and the canopy cover varies between 8 and 45%. the herbaceous layer has a canopy cover ranging from 48 to 90% and an average height of 0.5 m. 2.2 the colophospermum mopane – vernonia cinerascens subcommunity this subcommunity is found on the farms greefswald and riedel in the mapungubwe national park. as in the case of the acacia senegal – barleria senensis subcommunity (2.1), it occurs on the crests (tu1) and foot slopes (tu3) of the ib395 land type. the main soil forms found in this variant are shallow mispah and moderately deep glenrosa and hutton soils. the soil has a sandy to loamy texture and in the case of hutton soils the subsoil is rocky. the rockiness of the soil surface is between 20 and 50% and approximately 10 to 20% of the area is covered by exposed bedrock. grazing levels are moderate to low, depending on the accessibility of the landscape. the diagnostic species of this subcommunity are found in species group l (table 1). they include the tree species colophospermum mopane and boscia foetida, the shrub species rhigozum zambesiacum, the grass cenchrus ciliaris and the forbs vernonia cinerascens and corchorus asplenifolius. similarities between the colophospermum mopane – vernonia cinerascens subcommunity (2.2) and the acacia senegal – barleria senensis subcommunity (2.1) come in the form of the species of species group j (table 1). the tree layer has an average height of 4.2 m and a canopy cover varying between 10 and 45%. the shrub layer has a canopy 79 original research götze, cilliers, bezuidenhout & kellner vol. 50 no. 1 pp. 72 81koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za identified. according to the south african defence force (1986b), a feature of the landscape in which these communities are found is the steep irregular slopes, the shallow lithosols and the large number of large rocks (up to 30 cm in diameter) found on the soil surface. the ficus tettensis – aristida meriodinalis tall closed woodland described by dekker & van rooyen (1995) shows some similarities with the ficus tettensis – waltheria indica subcommunity of the hexalobus monopetalus – croton gratissimus var. subgratissimus community, but was associated with domeshaped granite hills in that study. the albizia – combretaceae open woodland/shrubland was identified and described in a study conducted by timberlake et al. (1999) across the limpopo river in the south-western parts of zimbabwe. regarding habitat and floristic composition, this vegetation unit shows similarities with the hexalobus monopetalus – croton gratissimus var. subgratissimus (1) and terminalia prunioides – grewia bicolor (2) communities, which were identified on the ib395 land type during the course of this study. this study made an important contribution to our knowledge of the vegetation on sandstone hills in the limpopo ridge bushveld, because little was known about them previously. due to the fragmentary nature of the ib land type, a vegetation map was not produced. a vegetation map of the mapungubwe national park will be compiled after a synthesis of the current study and studies on other land types, such as the ia land type (götze et al. 2003) and the db, fb and ae land types (götze 2002, götze et al. in prep.). the synthesis will also put the communities described in this study into context with other communities in the park (interpretation of relationships between different communities), making it possible to compare the major vegetation groups and subgroups in the park with previously described units such as the different types of mopane veld (siebert et al. 2003). the major vegetation groups and subgroups of the mapungubwe national park will be used as a basis to compile and map management units (götze 2002, götze et al. in prep.). acknowledgements south african national parks and the peace parks foundation are thanked for technical and financial assistance, and the south african defence force for providing soil and vegetation maps of the area. this study forms part of an msc thesis completed at the potchefstroom university for christian higher education (now potchefstroom campus of north-west university). references acocks, j.p.h. 1988. veld types of south africa, 3rd ed. memoirs of the botanical survey of south africa 57: 1–146. bezuidenhout, h. 1996. the major vegetation communities of the augrabies falls national park, northern cape. 1. the southern section. koedoe, 39(2): 7–24. bezuidenhout, h., biggs, h.c. & bredenkamp, g.j. 1996. a process supported by the utility bbpc for analyzing braunblanquet data on a personal computer. koedoe, 39(1): 107– 112. bredenkamp, g.j. & bezuidenhout, h. 1995. a proposed procedure for the analysis of large data sets in the classification of south african grasslands. koedoe, 38(1): 33–39. bredenkamp, g.j. & theron, g.k. 1978. a synecological account of the suikerbosrand nature reserve 1. the phytosociology of the witwatersrand geological system. bothalia, 12: 513– 529. bredenkamp, g.j., deutschländer, m.s. & theron, g.k. 1993. a phytosociological analysis of the albizio harveyi eucleetum divinorum from sodic bottom land clay soils of the manyeleti game reserve, gazankulu, south africa. south african journal of botany, 59(1): 57–64. cover ranging from 5 to 40% and an average height of 1.4 m. the canopy cover of the herbaceous layer varies between 45 and 68%, with an average canopy height of 0.6 m. 2.3 the hermbstaedtia odorata – heliotropium ovalifolium subcommunity this highly disturbed subcommunity occurs mainly on the farm greefswald in the mapungubwe national park. the degradation was caused primarily by the actions of the former south african defence force. the subcommunity is commonly found on old campsites, which vary from permanent to temporary camps. a large degree of deforestation and landscaping has taken place in most of these areas. it was also observed that specimens of the majestic adansonia digitata, which were removed from the surrounding areas, were transplanted in some of these areas. as a whole, these areas are degraded severely, with large areas devoid of vegetation. an old dumping site and a case of serious soil pollution at a former fuelling station were recorded as being among the most severely degraded sites in the area. the main soil forms vary from shallow mispah to moderately deep glenrosa and even some patches of witbank soils of varying depths. the witbank soil form comprises soils deposited by humans, usually for landfills or other landscaping works (soil classification work group 1991). the rockiness of the soil surface varies from 15 to 25%. grazing levels are moderate, but the impact of the grazing in these areas is severe due to the large degree of anthropogenic-induced degradation. no diagnostic species were identified in this degraded subcommunity, but it was characterised by the constant occurrence and high abundance of the species of species group o (table 1), including the shrubby cordia monoica, the grass species urochloa mosambicensis and the forbs hermbstaedtia odorata, heliotropium ovalifolium, tribulus zeyheri subsp. zeyheri and cleome angustifolia subsp. petersiana. this subcommunity is differentiated from the acacia senegal – barleria senensis subcommunity (2.1) and the colophospermum mopane – vernonia cinerascens subcommunity (2.2) by the absence or infrequent occurrence of the species of species groups j to n (table 1). the poor conditions of the veld are confirmed further by the presence of the weed species amaranthus, alternanthera pungens, chenopodium carinatum and schkuhria pinnata, and the grass species tragus berteronianus (other species, table 1). the tree layer reaches an average height of 6.0 m and a canopy cover of between 12 and 45%. the shrub layer has an average height of 1.6 m and the canopy cover varies between 1 and 30%, while the herbaceous layer reaches an average height of only 0.3 m and a poor canopy cover of between 30 and 45%. conclusion the high plant diversity, the great scenic beauty and uniqueness of the landscape, the cultural importance of the archaeological treasures of the mapungubwe cultural landscape in the ib land type in the mapungubwe national park, and a rich population of mammals, birds and reptiles, give this area a large claim to conservation. some of the vegetation types described in the ib land type are similar to those previously described in other studies. the south african defence force (1986b) previously described the vegetation and habitat of the sandstone ridges of the ib395 land type in a report on the vegetation of the maps of the areas mapungubwe (2229ab) and coila (2229ad). the mixed colophospermum mopane, combretum apiculatum, commiphora spp. and terminalia prunioides community identified by the south african defence force (1986b) was described on the same sandstone ridges of the study area in which the hexalobus monopetalus – croton gratissimus var. subgratissimus (1) and terminalia prunioides – grewia bicolor (2) communities were 80 vegetation of sandstone ridges original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 72 81 mucina, l. & rutherford, m.c. (eds.). 2006. the vegetation of south africa, lesotho and swaziland. strelitzia 19. pretoria: south african national biodiversity institute. mucina, l. & van der maarel, e. 1989. twenty years of numerical syntaxonomy. vegetatio, 81: 1–15. mucina, l., rutherford, m.c. & powrie, l.w. (eds.). 2005. vegetation map of south africa, lesotho and swaziland, 1:1 000 000 scale sheet maps. pretoria: south african national biodiversity institute. mueller-dombois, d. & ellenberg, h. 1974. aims and methods of vegetation ecology. new york: wiley. o’connor, t.g. 1991. the vegetation types of the limpopovenetia nature reserve. internal report, de beers consolidated mines, pretoria. robinson, g.a. 1996. limpopo valley national park: towards trans-frontier conservation in south africa. unpublished report, south african national parks. siebert, f., bredenkamp, g.j. & siebert, s.j. 2003. a comparison of mopaneveld vegetation in south africa, namibia and zimbabwe. bothalia, 33(1): 121–134. soil classification work group, 1991. grondklassifikasie: ‘n taksonomiese sisteem vir suid-afrika. memoirs on the natural agricultural resources of south africa 15. pretoria: department of agricultural development. south african defence force. 1986a. report on the vegetation of the map areas 2229aa, pontdrif and 2229ac, evangelina. unpublished report, south african defence force. south african defence force. 1986b. report on the vegetation of the map areas 2229ab, mapungubwe and 2229ad, coila. unpublished report, south african defence force. timberlake, j., mapaure, i. & chapano, c. 1999. vegetation survey of parts of the southern lowveld, zimbabwe. consultancy report for cesvi lowveld project. occasional paper in biodiversity no. 6. biodiversity foundation for africa, bulawayo. van rooyen, n. & bredenkamp, g.j. 1996. mopane bushveld. in: low, a.b. & rebelo, a.g. (eds.). vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. pp. 20–21 visser, n., van hoven, w. & theron, g.k. 1996. the vegetation and identification of management units of the honnet nature reserve, northern province, south africa. koedoe, 39(1): 25–42. voigt, e.a. & plug, i. 1981. early iron-age herders of the limpopo valley. internal report, transvaal museum, pretoria. walker, b.h. 1989. diversity and stability in ecosystem conservation. in: warren, a. & goldsmith, f.b. (eds.). conservation for the twenty-first century. new york: john wiley. pp. 121–130. willcox, a.r. 1966. memorandum concerning a suggested nature reserve and archaeological reserve in the limpopo valley. unpublished memo, national parks board. wynne-jones, a. 1982. hunting, on safari in east and southern africa. johannesburg: macmillan. brown, l.r. 1997. a plant ecological study and wildlife management plan of the borakalo nature reserve, northwest province. phd dissertation, university of pretoria, pretoria. cilliers, s.s. 1998. phytosociological studies of urban open spaces in potchefstroom, north west province, south africa. phd dissertation, potchefstroom university for che, potchefstroom. dekker, b. & van rooyen, n. 1995. the physical environment and plant communities of the messina experimental farm. south african journal of botany, 61(3): 158–167. edwards, d. 1972. botanical survey and agriculture. proceedings of the grassland society of south africa, 7: 15–19. edwards, d. 1983. a broad-scale structural classification of vegetation for practical purposes. bothalia, 14(3/4): 705–712. germishuizen, g., meyer, n.l., steenkamp, y. & keith, m. (eds.). 2006. a checklist of south african plants. south african botanical diversity network report no. 41. pretoria: sabonet. götze, a.r. 2002. the classification of plant communities and the evaluation of restoration technologies in different land-use areas in the vhembe-dongola national park, northern province, south africa. msc thesis. potchefstroom university for c.h.e., potchefstroom. götze, a.r., cilliers, s.s., bezuidenhout, h. & kellner, k. 2003. analysis of the riparian vegetation (ia land type) of the proposed vhembe-dongola national park, limpopo province, south africa. koedoe, 46(2): 45–64. götze, a.r., cilliers, s.s., bezuidenhout, h. & kellner, k. in prep. vegetation analysis of the db, fb and ae land types of selected areas of the mapungubwe national park, limpopo province, south africa. greyling, t. & huntley, b.j. 1984. directory of southern african conservation areas. south african national scientific programmes report, 98: 1–310. hennekens, s.m. 1996a. turboveg: software package for input, processing and presentation of phytosociological data. university of lancaster: ibn-dlo. hennekens, s.m. 1996b. megatab: a visual editor for phytosociological tables. ulft: giesen & gertus. hill, m.o. 1979a. twinspan: a fortran program for arranging multivariate data in an ordered two-way table by classification of individuals and attributes. cornell university, ithaca, new york. huffman, t.n. 2005. mapungubwe, ancient african civilisation on the limpopo. johannesburg: wits university press. kuman, k., le baron, j.c. & gibbon r.j. 2005. earlier stone age archaeology of the vhembe-dongola national park (south africa) and vicinity. quaternary international, 129: 23–32. land type survey staff, 2000. information on the land types of the map 2228 – alldays. pretoria: directorate of agricultural information. mcnaughton, s.j. 1989. ecosystems and conservation in the twenty-first century. in: western, d. & pearl, m.c. (eds.). conservation for the twenty-first century. oxford: oxford university press. pp. 109–120 81 filelist convert a pdf file! koedoe 19 : 167-168 (1976) the distribution of tomopterna krugerensis (anura: ranidae) in the republic of south africa n. i. passmore department ojzoology university oj the witwaterstand j ohanne sburg 2000 a new and cryptic species of tomoptema, t. krugerenis, has recently been described from the kruger national park, republic of south africa, (passmore and carruthers 1975), where it appears to be confined to the wambiya sandveld region in the north. the closely related and very similar t. delalandei cryptotis has not been recorded from this region . on 1976.01.06 and 1976.01.16 the author encountered a large breeding aggregation of t. krugerensis in a temporary rain pool on the farm mosdene, (naboomspruit, transvaal). the number of calling males was estimated at about 50 (± 10%). the mating calls heard and recorded were indistinguishable from those previously recorded in the kruger national park. t. delalandei cryptototis was also present and calling at this locality, together with t. krugerensis, on both occasions. t. krugerensis is morphologically very similar to t. delalandei cryptotis and the distinction of these two species is difficult in preserved material. the only reliable morphological differences are size, the extent of webbing on the hind feet and the presence of a doubled proximal subarticular tubercle on the first digit of the fore feet in t. krugerensis (passmore et al. 1975) . these differences are equally evident in the naboomspruit material as they are in the material from the kruger national park. the sympatric occurrence of these two species at this locality, accompanied by the maintenance of morphological and other differences, is a further clear indication of their distinctness at the species level. acknowledgement the author is grateful for the hospitality of mr. e. a. galpin, and mr. r. galpin of naboomspruit, transvaal. reference passmore, n. i. and v. c. carruthers, 1975 . a new species of tomoptema (anura: ranidae) from the kruger national park, with notes on related species. koedoe 18: 31-50. 167 page 1 a new benchmark in biodiversity monitoring book title: biodiversity in southern africa. volume 1–3 isbn: 978-99916-57-30-1 publisher: klaus hess publishers: göttingen & windhoek; 2010, r600* *book price at time of review reviewer: robert j. scholes1 affiliation: 1csir natural resources and environment, pretoria, south africa postal adress: po box 395, pretoria 0001, south africa how to cite this book review: scholes, r.j., 2012, ‘a new benchmark in biodiversity monitoring’, koedoe 54(1), art. #1109, 2 pages. http://dx.doi.org/10.4102/koedoe.v54i1.1109 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. book review open access the biota project began in south africa and namibia in the year 2000. it recently published a three-volume synthesis of its first decade of work. unusually, ‘biota’ is not an acronym for anything – it is the name of a major german-funded study into biodiversity patterns, initially focusing on the west side of southern africa. biota now has transects in north, west and east africa as well.biota (http://www.biota-africa.org) has been a pioneer in the field of ‘biodiversity observatories’ in other words, locations where the biodiversity in all its forms and aspects is monitored over large scales and long periods of time. conceptually, the biota approach represents a large step up from the simple checklists taken in small plots that characterised the field previously. in a rough outline, the biota network consists of 1 km x 1 km squares, distributed strategically along subcontinental-scale environmental gradients. in the case of southern africa, the network consists of 37 such squares; stretching from the high winter rainfall of the cape peninsula, through the drylands of namaqualand and the namib, to the high summer rainfall of northern namibia. in some cases, the squares are paired at a location, representing contrasting land uses. the sampling design within the 1 km squares is elaborate. each square is divided into 100 plots of 1 ha; nested within the plots are quadrats of 50 m x 20 m and 10 m x 10 m for vegetation sampling. other parts of the plot are reserved for activities that cause disturbances (such as soil pits or trapping of small animals) or areas that must be protected from disturbance, such as the mapping of soil crusts. the plots are pre-classified using vegetation and geomorphological information from aerial photos or high-resolution remote sensing. a stratified-random sample of 20 plots is taken, with weighting to ensure that rare habitat types are also featured. each square has an automatic weather station. the detail of the information gathered has a mesmerising thoroughness. no stone is left unturned and no taxonomic group is too obscure to attract attention. pedodiversity, biological soil crusts, lichens, vascular plants, millipedes, termites, bugs, beetles, butterflies, moths and vertebrates all have their own sample protocols. even humans and their use of the land come under the microscope. it is perhaps not surprising that three hefty volumes, in colour, were needed to summarise this wealth of information. the real long-term value of this project probably esides in the primary observations themselves, and how they change over time. the original data is in the public domain, which is conveniently included as a cd at the back of volume 3. what emerges from this prodigious and costly effort? volume 1 (801 pages) describes the methods and observed biodiversity patterns, site by site, within broad ecosystem clusters such as ‘woodland savanna’, ‘thornbush savanna’, ‘nama karoo’, ‘namib desert’, ‘succulent karoo’ and ‘fynbos’. it is beautifully laid out and illustrated, but the sheer volume of data is likely to overwhelm all but the most dedicated reader. volume 2 is a more approachable 348 pages. it extracts the patterns and processes at regional scale, in the form of eight chapters, each with multi-authored subsections, covering topics ranging from land use history, through biogeography, contemporary human impacts, the use of experiments and models and capacity building. i suspect most readers will dip into the papers that most attract them, rather than reading cover-to-cover. some interesting (but not completely unanticipated) patterns emerge. for instance, plant species richness at all scales from 10 m x 10 m to 1000 m x 1000 m declines with declining (summer) rainfall, is lowest in the desert and rises steeply with increasing winter rainfall to a maximum in the moist fynbos, at about double the score recorded in the wettest savannas. this pattern is much less obvious if the richness of plant families is considered. deserts have substantially less than half the family richness of the other biomes, which differ little between them. about a third of the variation in plant species richness is accounted for by soil-type diversity. moth diversity followed a similar relationship to precipitation as plants did, and reptile diversity increased with increasing grass cover. even a decade-long study is too short to say much about changes over time. there was a small increase in plant species richness over this period – so consistently across sites that i agree with the authors that it most probably reflects the growing observational skills of the botanists, rather than a fundamental change in the landscape. interestingly, in this study land use intensity has no consistent impact on species richness. volume 3, a comparatively slim 226 pages in length, covers management implications. it emphasises the interdisciplinary nature of the biota programme and includes modelled results from climate change studies as well as discussions of land use systems; in both cases only loosely linked to the biota observatories and their data. i was slightly disappointed to find little that was really new or could only have been derived from a biota-style system. it is perhaps unreasonable to expect novel insights from a young programme; nevertheless the pressure was obviously there to demonstrate how all that effort helped to guide decision-making, so something had to be said. the biota team have set an ambitious standard for biodiversity monitoring. the german research authorities continue to support the programme in its second decade, but have already signalled a desire that the within-region countries need to take over the long-term funding. herein lies the rub: despite paying a laudable amount of attention to building local partnerships and capacity, it is not clear that this is the system that south africa and namibia (or the various other countries in africa) would have built for their own purposes, and can afford to sustain. will it survive to fulfil its long-term promise, or is this a luxury german vehicle in an environment where a bakkie would be more appropriate? these volumes deserve a place in many institutional libraries in the region, especially those that deal with the ecosystems of the western part of the subcontinent. koedoe_58(1).indb reviewer acknowledgement http://www.koedoe.co.za open access koedoe recognises and acknowledges the value and importance of reviewers in the publication process – not only in shaping the individual manuscript, but also in upholding the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely, productive peer review process. we would like to take this opportunity to thank all reviewers who participated in shaping this volume of koedoe: page 1 of 1 in an effort to facilitate the selection of appropriate peer reviewers for koedoe, we ask that you take a moment to update your electronic portfolio at www.koedoe.co.za for our records. this will allow us better access to your areas of interest and expertise, in order to match reviewers with submitted manuscripts. if you would like to become a reviewer, please visit the journal website and register as a reviewer. to access your details on the website, you will need to follow these steps: 1. log into the online journal at http://www. koedoe.co.za 2. in your ‘user home’ [http://www.koedoe. co.za/index.php/koedoe/ user] select ‘edit my profile’ under the heading ‘my account’ and insert all relevant details, bio statement and reviewing interest. 3. it is good practice as a reviewer to update your personal details regularly, to ensure contact with you throughout your professional term as reviewer to koedoe. please do not hesitate to contact us if you require assistance in performing this task. publisher: publishing@aosis.co.za tel: +27 21 975 2602 fax: +27 21 975 4635 koedoe african protected area conservation and science anastasios legakis andrew rogers angela gaylard angelika wilhelm-rechmann anne i. dagg arne b.r. witt ben j. strohbach benjamin j. wigley chris harris colleen t. downs danny govender enoabasi d. anwana ernita van wyk georgette lagendijk gordon o’brien gretel van rooyen hari sridhar james brazill-boast jan a. venter jeremy simmonds joana bezerra jose a. corronca julian fennessy kate c. fagan llewellyn c. foxcroft louise swemmer magdalena remisiewicz marc e. stalmans martin k.s. smith mduduzi ndlovu mervyn c. lötter michael mills mike peel neville d. impson norbert cordeiro olaotswe e. kgosikoma peter b. moyle ram c. sharma robert buitenwerf ryan emanuel sven bourquin theo h.c. mostert tim o’connor tony palmer yolanda pretorius yuri m. marusik article information authors: mariette marais1 antoinette swart1 affiliations: 1national collection of nematodes (biosystematics programme), arc-plant protection research institute, south africa correspondence to: mariette marais postal address: private bag x134, queenswood 0121, south africa dates: received: 02 may 2012 accepted: 03 oct. 2012 published: 08 mar. 2013 how to cite this article: marais, m. & swart, a., 2013, ‘plant nematodes in south africa. 11. checklist of plant nematodes of the protected areas of kwazulu-natal’, koedoe 55(1), art. #1086, 2 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1086 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/ koedoe.v55i1.1086-1 copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. plant nematodes in south africa. 11. checklist of plant nematodes of the protected areas of kwazulu-natal in this checklist... open access • abstract • introduction • results and discussion • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ nematodes are some of the most abundant soil organisms and are an essential part of soil ecology. these organisms are used as indicator organisms and can be linked to soil health. biological collections contain vast amounts of data, with the national collection of nematodes housed at the plant protection research institute, agricultural research council being no different. during the digitising of the collection a number of unpublished records of plant nematodes reported from protected areas in kwazulu-natal were found in the south african plant-parasitic nematode survey database. a total of 222 plant nematode species belonging to 39 genera were reported from the province, with only 94 of these species reported from the protected areas and 172 and 159 species reported from uncultivated (outside the protected areas) and cultivated areas, respectively. only nine species, criconema silvum, criconema talanum, helicotylenchus marethae, ogma dracomontanum, ogma louisi, ogma ueckermanni, paralongidorus deborae, trichodorus rinae and xiphinemella marindae were described from protected areas, whilst o. dracomontanum, p. deborae and t. rinae were subsequently also reported from other provinces. conservation implications: a higher degree of diversity of nematodes was observed in the unprotected areas of the province. the observation suggests that nematode fauna, and by implication also other invertebrates, are not adequately protected. introduction top ↑ since the late 1950s nematodes were collected, identified and deposited in the national collection of nematodes at the plant protection research institute, agricultural research council, pretoria. descriptions and locality data of new species were published, but the incidence of plant nematodes in the protected areas of kwazulu-natal was never consolidated in a single document as was done for a number of protected areas in other provinces (botha & heyns 1990; van den berg, marais & tiedt 2007; van der vegte & heyns 1963). in 1987 the national collection of nematodes initiated the south african plant-parasitic nematode survey (sappns) to record the biodiversity of plant nematodes in south africa (marais 2006). as part of this initiative, surveys were undertaken in various protected areas, including those in kwazulu-natal (van den berg 1992). during digitising the specimens of the national collection of nematodes, a number of unpublished records of plant nematodes reported from protected areas in kwazulu-natal were discovered. results and discussion top ↑ the checklist reflects collections between 1963 and 2006 from 18 protected areas in kwazulu-natal. the sappns database contains 450 records of nematodes sampled in uncultivated areas in kwazulu-natal, of which 161 are from localities in protected areas (online appendix). the database also contains 580 records of localities sampled in cultivated areas, which include crop fields, plantations, gardens and sports fields. it was found that 94 plant nematode species were reported from protected areas (table 1), 172 species from uncultivated areas (outside the protected areas) and 159 species from cultivated areas. in total, 222 different plant nematode species are currently reported for kwazulu-natal. of these, 54 species were described for the province but only nine species were described for the protected areas in the province. table 1: number of plant nematode species recorded from kwazulu-natal. the classification of south african plant-parasitic nematodes followed here is a synthesis of the classification by maggenti et al. (1988) for tylenchina and decraemer (1995) and duarte et al. (2010) for trichodoridae. the synonymisation of the genera longidoroides and siddiqia with paralongidorus, as proposed by escuer and arias (1997), is accepted. authorities for genera regarded as valid here are marais (2001) for helicotylenchus, escuer and arias (1997) for paralongidorus, brzeski (1998) for paratylenchus, handoo (2000) for tylenchorhynchus and hunt, luc and manzanilla-lópez (2005) for criconemoides. acknowledgements top ↑ we thank various colleagues for collecting the samples, the taxonomists at the arc-ppri and (the former) rand afrikaans university for species identification and the south african biodiversity information facility for financial assistance to digitise a portion of the national collection of nematodes. competing interests the authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article. authors’ contributions m.m. (arc-plant protection research institute) is the co-ordinator of the sappns, project leader of digitising the national collection of nematodes and taxonomist responsible for helicotylenchus and belonolaimidae, dolichoridae, trichodoridae and the subfamily meloidogyninae. m.m. was responsible for writing the draft concept of the manuscript. a.s. (arc-plant protection research institute) is the taxonomist responsible for the longidoridae, anguinidae, aphelenchoididae and the subfamily heteroderinae at the collection and was responsible for the final review of the manuscript. m.m. and a.s. both contributed to writing the article. references top ↑ botha, a. & heyns, j., 1990, ‘a preliminary check list of the free-living and plant-parasitic nematodes of the kruger national park’, koedoe 33(2), 73–76. http://dx.doi.org/10.4102/koedoe.v33i2.442brzeski, m., 1998, nematodes of tylenchina in poland and temperate europe, museum i instytut zoologii polska akademia nauk, warzawa. decraemer, w., 1995, the family trichodoridae: stubby root and virus vector nematodes, kluwer academic press, dordrecht. http://dx.doi.org/10.1007/978-94-015-8482-1 duarte, i.m., de almeida, m.t., brown, d.j.f., marques, i. nielsen, r. & decraemer, w., 2010, ‘phylogenetic relationships, based on ssu rdna sequences, among the didelphic genera of the family trichodoridae from portugal’, nematology 12(2), 171–180. http://dx.doi.org/10.1163/156854109x461721 escuer, m. & arias, m., 1997, ‘paralongidorus iberis sp.n. and p. monegrensis sp.n. from spain with a polytomous key to the species of the genus paralongidorus siddiqi, hooper & khan, 1963 (nematoda: longidoridae)’, fundamental and applied nematology 20(2), 135–148. handoo, z., 2000, ‘a key and diagnostic compendium to the species of the genus tylenchorhynchus cobb, 1913 (nematoda: belonolaimidae)’, journal of nematology 32(1), 20–34. pmid:19270946 hunt, d.j., luc, m. & manzanilla-lópez, r.h., 2005, ‘identification, morphology and biology of plant parasitic nematodes’, in m. luc, r.a. sikora & j. bridge (eds.), plant parasitic nematodes in subtropical and tropical agriculture, 2nd edn., pp. 11–52, cabi publishing, wallingford. http://dx.doi.org/10.1079/9780851997278.0011 maggenti, a.r., luc, m., raski, d.j., fortuner, r. & geraert, e., 1988, ‘a reappraisal of tylenchina (nemata). 11. list of generic and supra-generic taxa, with their junior synonyms’, revue de nématologie 11(2), 177–188. marais, m., 2001, ‘a monograph of the genus helicotylenchus steiner, 1945 (nemata: hoplolaimidae)’, phd dissertation, department of entomology and nematology, university of stellenbosch. marais, m., 2006, ‘south african plant-parasitic nematode survey (sappns)’, plant protection news 67, 6. van den berg, e., 1992, ‘description of two new ogma southern, 1914 species (nemata: criconematoidea) with a list of plant-parasitic nematode species from natal’, koedoe 35(1), 9–18. http://dx.doi.org/10.4102/koedoe.v35i1.385 van den berg, e., marais, m. & tiedt, l.r., 2007, ‘plant nematodes in south africa. 10. check-list of plant nematodes from the goegap and witsand nature reserves, northern cape province, with a description of a new rotylenchus species (hoplolaimidae: nematoda)’, african plant protection 13, 28–35, viewed 22 september 2011, from http://journals.sabinet.co.za/webz/advancedquery?sessionid=01-64234-250806035 van der vegte, f.a. & heyns, j., 1963, ‘a report on nematodes found in soil and root samples from the kruger national park’, koedoe 6(1), 134–136. http://dx.doi.org/10.4102/koedoe.v6i1.820 article information authors: robert buitenwerf1 andrew kulmatiski2 steven i. higgins3,4 affiliations: 1institut für physische geographie, goethe universität frankfurt, germany 2department of plants, soils and climate and the ecology center, utah state university, united states 3department of botany, university of otago, new zealand 4biodiversity and climate research centre, senckenberg gesellschaft für naturforschung, germany correspondence to: robert buitenwerf postal address: altenhöferallee 1, 60438 frankfurt am main, germany dates: received: 19 mar. 2014 accepted: 04 aug. 2014 published: 10 nov. 2014 how to cite this article: buitenwerf, r., kulmatiski, a. & higgins, s.i., 2014, ‘soil water retention curves for the major soil types of the kruger national park’, koedoe 56(1), art. #1228, 9 pages. http://dx.doi.org/10.4102/ koedoe.v56i1.1228 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. soil water retention curves for the major soil types of the kruger national park in this short communication... open access • abstract • introduction • methods    • study site    • approach    • water retention curves    • soil texture • results • discussion • acknowledgements    • competing interests    • authors’ contributions • references • appendix 1 • appendix 2 abstract top ↑ soil water potential is crucial to plant transpiration and thus to carbon cycling and biosphere–atmosphere interactions, yet it is difficult to measure in the field. volumetric and gravimetric water contents are easy and cheap to measure in the field, but can be a poor proxy of plant-available water. soil water content can be transformed to water potential using soil moisture retention curves. we provide empirically derived soil moisture retention curves for seven soil types in the kruger national park, south africa. site-specific curves produced excellent estimates of soil water potential from soil water content values. curves from soils derived from the same geological substrate were similar, potentially allowing for the use of one curve for basalt soils and another for granite soils. it is anticipated that this dataset will help hydrologists and ecophysiologists understand water dynamics, carbon cycling and biosphere–atmosphere interactions under current and changing climatic conditions in the region. introduction top ↑ soil moisture is a key driver of plant productivity in many ecosystems, since water-stressed plants close their stomata to curb water loss, resulting in reduced carbon dioxide (co2) assimilation rates and, thus, growth (lambers et al. 2008). in order to understand water availability, ecologists, agronomists and land managers often use measurements of gravimetric (g of water g-1 soil) or volumetric (cm3 of water cm-3 soil) soil water to estimate water availability to plants. these measurements can easily be made by a wide array of sensors or simply by weighing and drying soil samples. the problem with these measurements is that they do not necessarily provide information about whether or not water is available to plants. this is because surface tension can bind large amounts of water to soils with a high internal surface area (i.e. clays) at such large negative pressures that plants cannot oppose them and are therefore unable to take up the water (vogel 2012). the negative pressure with which water is bound to the soil is the soil water potential. in non-saline soils, the relationship between soil water content and soil water potential largely depends on texture. clay particles have a large surface area relative to their volume and therefore have the ability to bind large amounts of water. as a result, a clay soil with, for example, 10% moisture, may have a highly negative water potential, making water uptake impossible for most plants. sand particles are larger and more spherical and therefore have a lower surface-to-volume ratio. consequently, a sandy soil with 10% moisture may have a water potential that is close to zero, allowing for easier water uptake by plants. temperate crops can transpire water through their stomata, and can therefore photosynthesize, down to a soil water potential of about -1.5 mpa. whilst this is called the permanent wilting point, some plants in arid systems are able to transpire water at a soil water potential as low as -5 mpa (baldocchi et al. 2004; rodriquez-iturbe & porporato 2004). ecologists, vegetation modellers and those modelling biosphere–atmosphere interactions therefore need accurate estimates of soil water potential in the range from -5 mpa to 0 mpa to predict and explain stomatal responses, carbon dynamics, and water and energy budgets. pedo-transfer functions transform variables that are easy and cheap to measure into more informative variables that are too difficult or expensive to measure directly (bouma 1989). one such function is the soil water retention curve, which transforms soil water content into soil water potential. the water retention curve is often estimated from information on soil texture, but can be determined precisely by measuring both variables simultaneously on samples under controlled conditions. here we provide empirically derived soil water characteristic curves for seven soils representing the major soil types of the kruger national park (knp). we test whether the same curve can be used for soils with a similar texture. methods top ↑ study site the kruger national park is situated in the north-east of south africa between 30.9–32.0 °e and 22.3–25.5 °s. the park receives between 450 mm and 750 mm yr-1 of rain, most of which falls between october and march. average temperatures are approximately 25 °c in summer and 20 °c in winter. most of knp is underlain by either basaltic rock that weathers into clay-rich soils or granitic rock that weathers into sandy soils. both of these dominant parent materials are old: the basaltic rock was formed ~200 ma, whilst the granite was formed ~2050 ma (venter et al. 2003). seven dominant soil types from across the park (venter et al. 2003) were sampled (figure 1; table 1). the sampled soil types cover approximately 65% of knp. figure 1: dominant soil types of the kruger national park following venter et al. (2003), and the sampling locations of this study. table 1: coordinates of the sampling sites and description of soils at those sites, following venter et al. (2003). the sand, silt and clay fractions are the mean of all samples from a site in percentages ± the standard error. approach using a soil auger, samples were collected at three depths: at the top of the profile (0 cm – 10 cm), at the bottom (110 cm or degraded bedrock, whichever was shallower) and in the middle (table 2). soils were air dried and sieved in a 2 mm-aperture sieve to remove large roots and rocks. the proportion of rock with dimensions greater than 2 mm was generally negligible (< 3%). table 2: parameter estimates, standard deviations of the parameter estimates, and goodness of fit for the power functions fitted as water retention curves for soils in the kruger national park. water retention curves soil moisture retention curves were derived empirically using an instrument that exploits the chilled-mirror technique (wp4t, decagon devices inc., pullman). a soil sample is inserted into the device and equilibrated with the headspace of a sealed chamber, so that the water potential of the air in the chamber is the same as the water potential of the sample. the point of condensation on a mirror is detected by shining a beam of light onto the mirror and recording its reflectance with a light sensor. at the point of condensation, the temperature of the mirror is recorded, allowing the water potential of the air, and therefore of the sample, to be calculated. the instrument measures the water potential with an accuracy of 0.05 mpa over the range of -0.1 mpa – -0.5 mpa and an accuracy of 0.1 mpa over the range of -0.5 mpa – -300 mpa. therefore, it adequately covers the range relevant to plant growth. the water content of the sample at the time of measuring water potential is determined gravimetrically (i.e. it is weighed and reweighed after oven-drying at 60 °c to a constant weight). since the relationship between water potential and water content is often hysteretic (affected by the initial state of the system), we constructed soil moisture retention curves from both drying and wetting soils. soil from each sampling location and depth was subdivided into 15 samples of 5 g – 6 g each. these samples were used to construct a single drying curve and a single wetting curve for each depth at each sampling location. for the drying curve, the samples were initially saturated with distilled water, sealed, and left to equilibrate overnight. water potential and moisture content were then measured repeatedly as the samples air dried. a typical curve was completed within a day. for the wetting curve, incremental amounts of moisture were added to initially dry samples to achieve a range of saturation levels. samples were then sealed and left to equilibrate overnight, after which the water potential and moisture content were measured. empirical models for the water retention curve are typically written to be solved for water content (see assouline et al. [1998] for an overview). we fitted two widely used models, proposed by van genuchten (1980), and fredlund and xing (1994), which were rewritten to solve for water potential given a known water content using non-linear regression in r (r core team 2013). we compared the performance of these models to simpler power and exponential functions, which were fitted by maximising the likelihood using iterative methods (plummer 2013). these functions were fitted to both the entire measured range of water potentials (0 mpa – -100 mpa) and to a subset that is relevant to plant growth (0 mpa – -8 mpa). to assess the variability amongst sites of the same geology, we compared site-specific curves to curves for which the data were aggregated by geology. this was done for four levels of water content (0.03 g g-1, 0.05 g g-1, 0.1 g g-1 and 0.15 g g-1). for this analysis, soils from punda, which are derived from quartzite (venter et al. 2003), were included with granite-derived soils as they had a very similar texture. soil texture particle size affects the total surface area for a given mass or volume of soil, and therefore the shape of the water retention curve. to quantify sand, silt and clay fractions, the hydrometer method (bouyoucos 1962) was used. for each sample, between 40 g and 50 g of soil was dispersed with an electrical mixer in 100 ml of a 5% sodium hexametaphosphate ([napo3]6) solution, in order to break down clay aggregates. this mixture was allowed to soak overnight and mixed again the following morning before being transferred to a 1 l cylinder filled up to 1 l with distilled water. sediment was dispersed with a plunger and hydrometer readings were taken after 40 s and 6 h 52 min. fractions were calculated as: where h is the hydrometer reading at t seconds after inserting the hydrometer, and w is the sample weight in grams. hydrometer readings were corrected for the added (napo3)6 by subtracting the hydrometer reading of a cylinder with 100 ml (napo3)6 solution filled up to 1 l with distilled water. the temperature was measured at the time of each hydrometer reading and 0.4 g l-1 of solute concentration was added to the hydrometer reading for every degree c above 20 °c ( http://uwlab.soils.wisc.edu/files/procedures/particle_size.pdf). results top ↑ as expected, soils from the basalt substrate had a much higher clay content than soils from the granitic substrate (table 1; table 2). the water potential and water content data is given in online appendix 1. the fredlund and xing model (1994) appeared to fit the data well, with r2 values > 0.99 for most curves. however, the confidence intervals on the parameter estimates could not be estimated using standard techniques, suggesting the parameters were not identifiable. in other words, multiple combinations of parameter values can result in an identical fit. this outcome is likely to be a symptom of over-parameterisation of the model. the van genuchten model (1980) did not converge well, also hinting at over-parameterisation. therefore only results from the simpler power and exponential functions were reported. the best fit to individual curves in the -0.5 mpa – -8 mpa range was provided by a power function of the form (table 2; appendix 1): where ψ is water potential in mpa, θ is gravimetric water content in g g-1, and a and c are estimated coefficients. for a given water content, water potentials from drying curves were generally lower than for wetting curves (appendix 1). when combining data from all sites per geological substrate, the residuals of the power function fits showed systematic bias. an exponential model to describe the data aggregated by geology was therefore used. the exponential model fits these grouped data reasonably well in the -0.5 mpa – -8 mpa range: basalt r2 = 0.59; granite r2 = 0.66 (figure 2; see appendix 2 for 0 mpa – -100 mpa range). it should be noted that r2 values should be interpreted with caution in non-linear regression, and that the standard error of the regression (ser) is a better measure of goodness of fit: basalt ser = 1.42 mpa; granite ser = 1.13 mpa. the equation to convert soil water content to water potential for basaltic (clayey) soils in the -0.5 mpa – -8 mpa range is: for granitic (sandy) soils in the -0.5 mpa – -8 mpa range, the equation is: to assess the accuracy of these geology-specific curves, we compared the range in water potential for four water contents (0.03, 0.05, 0.10 and 0.15), as predicted by the site-specific models to the predicted water potential, using the geology-specific curve (figure 3). the geology-specific curves predicted water potentials close to the median value of site-specific curves, suggesting that they might be useful for modelling applications. figure 2: the relationship between soil moisture and water potential in the range relevant to plant growth, for (a) basalt-derived and (b) granite-derived soils in the kruger national park. note that the pressures on the y-axis are negative pressures. the water retention curves were fitted using an exponential model. figure 3: comparing site-specific and geology-specific curves for (a) basalt and (b) granite. each box shows the range of predicted water potentials from all site-specific curves per geological substrate for a given water content. the solid horizontal line in the box denotes the median of predicted values. the horizontal dotted lines indicate the wilting range (-1.5 mpa – -5 mpa). the * symbols show the predicted values from the geology-specific curves. the water potentials at water contents of 0.03 g g-1 and 0.05 g g-1 in basaltic soils are not shown as they are < -25 mpa and not relevant for plant physiological processes. discussion top ↑ ecophysiological and hydrological interpretation of soil water content – an affordable measure of soil moisture – requires the use of pedo-transfer functions that transform water content into water potential. we generated such water retention curves with a wp4t instrument (decagon devices inc., pullman) using empirical data, and provided curves for common soil types in knp, south africa. a power function provided a good fit for individual samples, and, depending on the required accuracy, a single exponential function per geological substrate may be used. at low water contents (i.e. ~5% for granitic and ~15% for basaltic soils), water potentials estimated from our pedo-transfer functions begin to vary widely between sites, reflecting differences in clay content. thus, researchers interested in precise estimates of soil water potentials in dry soils should use our site-specific functions. however, these water potentials are below the permanent wilting range (-1.5 mpa – -5 mpa; figure 3) and represent small volumes of soil water. researchers using the equations provided here should take care to avoid several potential problems. first, our analyses relied on the chilled-mirror technique. this method is highly accurate in the dry range (-0.5 mpa – -300 mpa), but is less accurate in the wet range of soils (-0.001 mpa – -0.5 mpa). it should be noted that some crop species may become water limited at -0.03 mpa (rodriquez-iturbe & porporato 2004). second, the functions presented here are based on gravimetric soil water measurements, whilst some field sensors measure soil moisture volumetrically. to use the functions provided in this study, volumetric soil moisture should be divided by the bulk density of the soil. bulk density in knp varies with texture, spatially, and with soil depth (wigley et al. 2013). when converting gravimetric or volumetric contents to water potentials, researchers should be aware of the potential role of coarse rock fragments. a soil sensor that provides volumetric water content in a rocky soil will underestimate soil water potentials because the volumetric sensor reports the water content of a volume comprised of both rock and soil. in conclusion, the presented soil water retention curves will improve estimates of plant-available water from measurements of volumetric or gravimetric soil moisture in knp and surrounding areas with similar geological substrates. acknowledgements top ↑ the authors would like to thank the management of the kruger national park for enabling this study. ben wigley kindly made bulk density data available; tercia strydom made equipment available; alexander zizka, julia fischer and jenny toivio assisted with data collection; and stephen doucette-riise and his team collected soil samples. this study was funded by the deutsche forschungsgemeinschaft (dfg). competing interests the authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article. authors’ contributions r.b. (goethe universität frankfurt) collected and analysed data, and wrote the article. a.k. (utah state university) conceived the study and wrote the article. s.i.h. (university of otago) analysed data and wrote the article. references top ↑ assouline, s., tessier, d. & bruand, a., 1998, ‘a conceptual model of the soil water retention curve’, water resources research 34(2), 223–231. http://dx.doi.org/10.1029/97wr03039 baldocchi, d.d., xu, l. & kiang, n., 2004, ‘how plant functional-type, weather, seasonal drought, and soil physical properties alter water and energy fluxes of an oak–grass savanna and an annual grassland’, agricultural and forest meteorology 123(1 & 2), 13–39. http://dx.doi.org/10.1016/j.agrformet.2003.11.006 bouma, j., 1989, ‘using soil survey data for quantitative land evaluation’, in b.a. stewart (ed.), advances in soil science, pp. 177–213, springer-verlag, new york. http://dx.doi.org/10.1007/978-1-4612-3532-3_4 bouyoucos, g.j., 1962, ‘hydrometer method improved for making particle size analyses of soils 1’, agronomy journal 54(5), 464–465. http://dx.doi.org/10.2134/agronj1962.00021962005400050028x fredlund, d. & xing, a., 1994, ‘equations for the soil-water characteristic curve’, canadian geotechnical journal 31(3), 521–532. http://dx.doi.org/10.1139/t94-061 lambers, h., chapin iii, f.s. & pons, t.l., 2008, plant physiological ecology, springer, new york. http://dx.doi.org/10.1007/978-0-387-78341-3 plummer, m., 2013, jags – just another gibbs sampler, viewed 2013, from http://mcmc-jags.sourceforge.net/ r core team, 2013, r: a language and environment for statistical computing, r foundation for statistical computing, vienna, austria, viewed 2013, from http://www.r-project.org/ rodriquez-iturbe, i. & porporato, a., 2004, ecohydrology of water-controlled ecosystems: soil moisture and plant dynamics, cambridge university press, cambridge. van genuchten, m.t., 1980, ‘a closed-form equation for predicting the hydraulic conductivity of unsaturated soils’, soil science society of america journal 44(5), 892–898. http://dx.doi.org/10.2136/sssaj1980.03615995004400050002x venter, f.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation pattern’, in j.t. du toit, k.v. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83–129, island press, washington dc. vogel, s., 2012, the life of a leaf, university of chicago press, chicago. http://dx.doi.org/10.7208/chicago/9780226859422.001.0001 wigley, b.j., coetsee, c., hartshorn, a.s. & bond, w.j, 2013, ‘what do ecologists miss by not digging deep enough? insights and methodological guidelines for assessing soil fertility status in ecological studies’, acta oecologica 51, 17–27. http://dx.doi.org/10.1016/j.actao.2013.05.007 appendix 1 top ↑ figure a1: the relationship between soil moisture and water potential in the range relevant to plant growth for each of the sampled soils in the kruger national park, letaba, lower sabie, phalaborwa, pretoriuskop, punda, satara, skukuza. the pressures on the y-axis are negative pressures. the water retention curves were fitted using a power function. the parameter values for the curves are given in table 2. figure a1 (continues...): the relationship between soil moisture and water potential in the range relevant to plant growth for each of the sampled soils in the kruger national park, letaba, lower sabie, phalaborwa, pretoriuskop, punda, satara, skukuza. the pressures on the y-axis are negative pressures. the water retention curves were fitted using a power function. the parameter values for the curves are given in table 2. appendix 2 top ↑ figure a2: the relationship between soil moisture and water potential in the 0 mpa – -100 mpa range for (a) basalt-derived and (b) granite-derived soils in the kruger national park. the pressures on the y-axis are negative pressures. the water retention curves were fitted using an exponential model. filelist convert a pdf file! supplement to koedoe. 1977: 224-229. deep sea fisheries implications in conservation l botha chief professional officer department of industries sea fisheries branch private bag sea point 8060 i stand before you as what i would reluctantly like to call a crisis conservationist. in marine fisheries conservation we are exposed to the full blast of human greed. dr hey has referred to the unfortunate human attitudes to nature conservation. i fully share his concern. fisheries conservation finds itself in the unenviable position of having largely to apply conservation principles to species which are more often than not in advanced stages of commercial exploitation. in this situation ethics, aesthetics and science cuts little or no ice with decision-makers. the only language which they understand is economics. unfortunately fisheries conservation has also been hampered by the well-meant but thoroughly unscientific efforts of certain lay so-called "conservation societies". through their admirable but purely humanistic approach to conservation, they have antagonized economically-orientated decision-makers and thereby unfortunately harmed the cause of conservation. in this regard i would like to associate myself strongly with dr vollmar, mr de la bat and numerous other speakers who have emphasized that we should put all our efforts into convincing decision-makers and others that conservation is in fact an economic proposition. in the deep-sea fisheries conservation situation, mr chairman, i feel that we should take special note of the emphasis placed on "utilization" or "wise use" by many previous speakers, rather than laying too much emphasis on the traditional "safeguarding" interpretation of conservation. in this regard, mr chairman, i have followed with interest the evolution of the concept of nature conservation over the last few years. more recent definitions have given increasing prominence to terms such as "utilization" and "wise use" of wildlife resources. i do feel, mr chairman, that the concept of "nature utilization" may be easier to sell to decision-makers and the public at large, rather than the concept of "nature conservation" with its more traditional implications of protection purely for pleasure. especially in developing countries, and southern africa is a developing area mr chairman, wise utilization of natural 224 resources makes more sense at least initially to the ordinary people, than the comparative easthetic luxury of pure traditional conservation . however, i do not wish to digress from my topic of marine conservation. my main purpose in drawing attention to the "new" definition of nature conservation is that my and other similar institutes can now for the first time consider themselves as having come in from the cold, so to speak. for many years marine fisheries conservation was not considered by many professionals as true conservation in the strict sense of the old definition because of the overriding importance of the commercial utilization component thereof. marine fisheries conservation will probably always be considered as a somewhat special type of conservation since it is invariably applied to populations which are already in a state of varying degrees of exploitation. nevertheless, i do feel that the new definition now provides some justification for my institute to consider itself as probably the largest specialized nature conservation institute of its type in southern africa . with a staff of more than 500, five laboratories and field stations, 12 research and fisheries patrol vessels and an operational area stretching from the cunene river right around the coast to the southern mocambique border, this is no hollow claim. at stake is the well-being of an industry which harvests more than i million metric tons of marine life valued at close on r200 million per year and an earner of valuable foreign exchange. the wise utilization of these vast living marine resources is the main purpose and function of my organization. the sea fisheries institute is a branch of the south african department of industries, which in turn advises the minister of economic affairs on fisheries management policy. the ultimate responsibility for the conservation of marine life beyond the highwater-mark rests with the minister. such responsibility is exercised through the sea fisheries act and regulations and various other relevant acts, which are in turn administered by the sea fisheries branch . the all important research function of the branch is carried out by a team of 43 professional scientists qualified in the fields of physical and chemical oceanography, ,fisheries biology, planktology, marine acoustics, algology, marine mammology, electronic data processing, engineering and other highly specialized fields. research projects logically relate in priority to the relative commercial importance of the various components of the fishing industry. current priority projects in the branch consequently entail work on the population dynamics, biology and the fisheries, of in the first instance, species such as pilchards, anchovies, mackerel and maasbanker, which form the basis of the massive pelagic sector of the fishing industry. this sector of the industry alone accounts for more than 80% of south africa's annual harvest from the sea and has secured our coveted, although in some quarters dubious, position as the second largest fish meal exporter in the world after peru, and one of the major fishing nations . the catastrophic collapse of the californian pilchard 225 fishery because of overfishing during the nineteen fifties, and the recent virtual collapse of the world's largest single species fishery for the pcruvian anchovy because ofa combination of heavy fishing and adverse environmental conditions, have underlined the tremendous importance of a sound ecological approach to such animals . in fact, despite energetic efforts to save it, the south african pilchard fishery also collapsed during the late nineteen sixties. the vast pilchard resource off the south west african coast is now the last major fishery of its kind in the world and no effort is spared to keep it that way. research staff of the branch have pioneered population abundance estimates by means of inter alia airborne survey methods, which have yielded much quicker results than the traditional somewhat more accurate but time-consuming methods of population estimates by means of recruitment, mortality, growth and fishing effort studies. particular attention is paid to environmental parameters which have a profound influence on stock availability and abundance. the fishery is currently managed by means of a catch quota system, limits on the number of processing factories, hold capacity and number of vessels, a closed season and closed areas. early results seem to confirm that the r ece nt decline in the swa pilchard fishery has been arrested in this way and provide considerable optimism for the future. other projects entail work on south africa's important rock lobster resources, including an assessment of the abundance of the newlydiscovered rock lobster resource off port elizabeth. early indications are that this new stock has an extremely slow growth rate and that its potential catch may therefore be considerably lower than the optimistic expectations created by initial high catch rates. pending more concrete results about potential yields, this new fishery is currently managed on a permit basis and foreign vessels have been ousted from the fishery by extending south africa's jurisdiction over this sedentary continental shelf species which occurs outside our present fishing zone, in terms of an existing geneva convention which provides for such action. south africa's traditional coastal rock lobster fishery is in a good state and is managed successfully by means of a catch quota, sanctuaries, a minimum permissible fish size and a ban on the catching of soft-shelled animals and berried females. the rock lobster industry itself is also subjected to license limitation and strict landing, transport, processing and quality control restrictions. other research projects entail work on various bait organisms such as mussels, oysters, red-bait, scallops and abalone, while considerable attention has lately also been focussed on whales and particularly on seals. you may know that the last south african whaling company has recently ceased operations altogether and a long south african tradition as a whale-catching country has thus come to an end. the reason for this closure has mainly resulted from a very strict catch quota system now introduced by the international whaling commission (iwc) as a result of years of overexploitation of certain whales by various countries. south 226 africa is a member of the iwc and scientists from the sea fisheries branch have and are still taking an active part in the scientific work of the commission. while no tuna fishery as such exists in south africa, the country is an active member of the international commission for the conservation of atlantic tunas (iccat). scientists from the branch attend regular meetings of this commission and contribute data to the population assessments of the stocks of the various highly-migratory tunas of the atlantic ocean. apart from the foregoing, the sea fisheries branch also undertakes studies on the physical! chemical aspects of our oceans including work on pesticides and marine pollution. however, since it is not my intention at the present time to inform you fully of all the various and wide ranging projects in the field of marine conservation in current progress at the branch, the foregoing brief yet inadequate outline will have to suffice. i would, however, now like to come to the main topic of my talk namely the implications of conservation in deepsea fisheries. you may be aware of the fact that some fifteen different foreign countries are most actively removing almost a million tons of fish from the waters around southand south west africa each year. this development started on a modest scale in 1962 when spanish and japanese trawlers, driven thereto by extremely low catch rates resulting from years of heavy fishing on their home grounds, first probed and then arrived in force on the then rich southeast atlantic fishing grounds. their phenomenal catch successes soon became known in fishing circles and at present some 300 huge, modern stern-trawlers drag up tons and tons of fish from the cold and fertile waters of the southern african westcoast. their main target is the cape hake (or stockfish) which is the dominant fish species in the area. estimates of maximum sustainable yield for hake provide a figure of some 950000 tons per year, but catches have in recent years exceeded 1 million metric tons. these high catches in combination with declining catch rates have led to serious national and international concern. this concern culminated in a meeting of plenipotentiaries which was held under the auspices of the food and agriculture organization of the un in rome during 1969. the result of this meeting was a convention on the conservation of the living resources of the southeast atlantic ocean , in which the participating nations agreed to take certain prescribed steps to conserve the marine populations in the area. the convention became a reality in 1972 when the newly-established international commission for the southeast atlantic fisheries (icseaf) met for the first time at its headquarters in madrid to give effect to the terms of the convention. members of icseaf currently comprise south africa, spain, the ussr, portugal, japan, cuba, poland, israel, bulgaria, france, belgium, italy and the german democratic republic. the commission meets every second year in madrid where it considers the scientific 227 advice of its permanent scientific advisory council. scientists from member countries, including scientists from the sea fisheries branch, participate actively in the work of the council. the scientific work entails the usual biological and population dynamics studies with the main aim of establishing optimum catch levels. the difficulty of course with deepsea fisheries conservation is that operations are conducted in international waters beyond the jurisdiction of coastal state territorial or fisheries waters. for this reason it is not yet possible for a coastal state such as south africa to unilaterally prescribe and enforce adequate conservation measures for the living marine resources far off its own coasts. i say this is not yet possible, because certain recent developments will in due course in fact make this a possibility. this brings us to the core of our present discussion on the implications in conservation of deepsea fisheries. the existing set-up of international deepsea fisheries conservation provides no adequate protection of the exploited stocks. it is left to the lengthy, cumbersome and often undecisive deliberations and actions of international commissions such as icseaf, iwc and others to agree to conservation measures. these international fisheries commissions have no legal powers and can only offer advice to governments, which mayor may not be endorsed and accepted in practice by the governments concerned, depending on their respective interests. the result is that, despite the very active existence of a multitude of such fisheries commissions throughout the world, few if any of them have achieved real conservational successes. many conservation measures have in fact been formally accepted, but the practical implementation and enforcement thereof remains a largely unsolved problem. hence we witness the so-called "fish war" between the united kingdom and iceland for example, which has had serious repercussions even in jeopardizing their future participation in the north atlantic treaty organization. it is precisely this dissatisfaction with existing international deep-sea fisheries commissions which has prompted many countries to press for exclusive coastal state control of fisheries. you may know that such deliberations are at present in progress at the un law of the sea conference. as far as fisheries conservation is concerned, wide consensus exists at the conference that the conservation and management of living marine resources will in the future be the exclusive responsibility of coastal states up to 200 nautical miles off their respective coasts. while some countries have doubts about the competence and expertise with which certain countries may exercise such a responsibility, most are convinced that it will for the first time in history provide for the full and adequate protection of coastal fisheries. highly migratory and wideranging species such as tuna and whales will still, however, by their very nature be the responsibility of international commissions. despite these developments and despite the dissatisfaction with existing international fisheries commission, an organization such as 228 icseaf has nevertheless taken significant steps to conserve fish stocks. so for example the member countries of icseaf recently unanimously adopted and implemented a minimum mesh-size in the trawlnets of their vessels fishing in the southeast atlantic ocean . it furthermore implemented a joint international inspection scheme to enforce this regulation. south africa, through the sea fisheries branch, is a leading and active participant in this scheme which, with 13 countries participating, must surely be one of the largest conservation schemes in operation at this moment in our area. south african fisheries inspectors of the sea fisheries branch have during the past year boarded some 180 japanese, russian, bulgarian, polish and south african fishing vessels on the high seas to inspect their nets in terms of the scheme of joint international enforcement. only 19 (or 10%) contraventions have been reported, which is considered as very good for a conservation scheme of this nature, scope and complexity. clearly the will to succeed is there, which augurs well for the future. it is this kind of co-operative and positive development in international deep-sea fisheries conservation which gives some hope for the continued existence of international fisheries commissions, despite the looming exclusive coastal state jurisdiction of the future. this is then, in summary, how i see the implications in deep-sea conservation in the future, namely the continued existence of international commissions, which would act mainly as a forum for the collective sharing of scientific problems and the transfer of marine scientific technology, coupled with and perhaps supplementing full and exclusive coastal state jurisdiction over the vast and valuable living resources of the world oceans. there is a considerable amount of urgency to protect many already overexploited stocks, but as far as a deep-sea conservation is concerned, i am optimistic that man will rise to the challenge and that the new order will ensure adequate protection of our marine wildlife. 229 page 1 page 2 page 3 page 4 page 5 page 6 book information title: game ranch management book cover game ranch management editors: j. du p. bothma j.g. du toit isbn: 9780627027154 publisher: van schaik publishers, pretoria; 2010, p. 979, hardcover: r769.95* *book price at time of review book review information reviewer title: meeting the challenges of game ranch management reviewer: roger collinson1 affiliation: 1change management advisor, etosha national park, namibia email: rcollinson@met.na postal address: po box 86819, eros park, windhoek, namibia how to cite this article: collinson, r., 2011, ‘meeting the challenges of game ranch management’, koedoe 53(1), art. #1050, 2 pages. doi:10.4102/koedoe.v53i1.1050 copyright notice: ©2011. the authors. licensee: aosis openjournals. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) meeting the challenges of game ranch management since the publication of the afrikaans version of game ranch management in 1986 and the appearance of the first english edition in 1989, the game ranching industry in south africa, and in neighbouring states such as namibia, zimbabwe and zambia, has grown in leaps and bounds. the role this book has played in facilitating this phenomenal growth has been undoubtedly significant. accordingly, the book’s editors, contributing authors and publishers need to be highly commended for this – especially with regard to their efforts over the past 25 years in providing regularly updated afrikaans and english editions in response to an ever-increasing knowledge base. at first sight, the most striking feature of the recently published fifth english edition of game ranch management is its large size. in comparison with the first and the other earlier editions, this striking feature gives the impression that the latest edition has more than ably kept abreast of the rapid growth of the industry it addresses. on closer examination, this first impression is largely confirmed by the wide range of topics covered by the book contents – inclusive of some theoretical but mainly very practical matters. more specifically, the topics covered include inter alia research, ecological principles and monitoring, vegetation types, game ranch planning, the biology and diseases of game ranch species, capture, hunting, captive breeding, as well as other aspects of commercial wildlife utilisation such as habitat management and rural development in the game ranching context. despite its voluminous content, the editors mention in the book’s preface that the knowledge base of game ranch management has grown exponentially over the last few decades, to the extent that it now exceeds the maximum page volume for a single book. one way of dealing with a challenge of this nature would have been to split the book into two volumes; instead, the current editors and publishers wisely elected to meet this challenge by excluding material which is readily available in separate, more specialised books on game ranch management topics. accordingly, the book often refers the reader to other publications currently available, such as intensive wildlife production in southern africa (edited by j. du p. bothma and n. van rooyen and published by van schaik publishers in 2005) and the capture, care and management of wildlife (authored by mike la grange and published by van schaik publishers in 2005). whilst the original intention of this book was to produce a text for the benefit of wildlife management students and wildlife producers, its content is also of some relevance to people with either a professional or armchair interest in wildlife conservation and/or ecotourism. book sales of previous editions support this contention, as the latter group of people has been a major market source over the years – not only in south africa but also elsewhere in africa and the world. thus, the relevance of the book to this wide ranging readership is yet another one of its strengths. however, these strengths are countered to some extent by a weakness that cannot go unmentioned here. the essence of this weakness is encapsulated in the ‘acknowledgements’ on page vii of the book, where the editors state at the start of the first paragraph ‘… the extensive production of wildlife, or wildlife ranching as it is also known, has moved from being based mainly on experience to being based mainly on fact’. this statement holds only for what may be termed ‘hard-edged’ technology such as that developed and refined over the years for game fencing, game capture, disease treatment, other veterinary technology, intensive game breeding, remote tracking, road construction, water point infrastructure, et cetera. however, with regard to what may be termed ‘soft-edged’ technology or applied ecologically based science, i believe that the editors’ statement, as quoted above, is somewhat misleading. important examples of where applied ecological science is relevant are the optimisation of multispecies game stocking rates and the management of natural habitats for maximising financial returns from a game ranch. given the many complexities and variables that exist within and between individual game ranches, such as rainfall, soils, vegetation and other landscape features, there are no (and probably never will be any) readily available textbook recipes, solutions or facts for reliably informing the game rancher on these applied ecological issues. accordingly, it is now recognised that the most useful approach to this seeming dilemma is for game ranchers to determine individualised (i.e. their own tailor-made) solutions for their specific set of objectives and environmental circumstances, ultimately through a process of trial and error. for the successful application of such an approach, the role of applied science should be to provide game ranchers with an objective decision-making framework for guiding them accordingly – whilst, at the same time, equipping them with an understanding of the many ecological and monitoring principles and techniques involved. this type of objective decision-making framework or process is known more formally as adaptive management. unfortunately, no elaboration of the concept of adaptive management could be found when reviewing this book and the term is absent from both the table of content and the index at the end of the book. in the absence of an understanding of this concept and a framework for applying it, game ranchers involved with extensive wildlife production in multispecies ‘natural systems’ are left groping in the dark with a kaleidoscope of untested and misunderstood ecological variables and principles. in these circumstances it is not surprising that game ranchers are expediently moving more and more towards single species intensive production systems (e.g. sable, roan and buffalo breeding schemes and ‘put and take’ production systems), where the environment can be simplified and rigorously controlled by way of ‘hard-edged’ technology that has been well developed, refined and documented and, hence, where there is greater certainty and less risk. thus, whilst the fifth english edition of game ranch management provides an expanded, robust source of information for game ranch managers, additional emphasis on these ‘soft’ issues would provide further value. on the basis of the aforementioned, it may be said that whilst building on this substantial volume, the future challenge is to produce and publish a specialist book that addresses the concept of adaptive management specifically and also provides game ranch managers with decision-making frameworks for dealing with a variety of applied management topics. imperative topics in this regard are optimum stocking rates, sex and age structures, sustained yield harvesting, fire management, bush encroachment, predator prey relationships and water distribution – to name but a few. a publication of this nature would not only be of benefit to game ranching systems where commercial production is the primary objective, but also to wildlife ranches, private game reserves and formally protected areas where biodiversity conservation and/or ecotourism is of prime importance. issn 0075-6458 45 koedoe 49/2 (2006)issn 0075-6458 45 koedoe 49/2 (2006) introduction the agulhas plain at the southern tip of africa has been recognised as an area of high conservation importance for both terrestrial plants (rebello & siegfried 1992; cowling & mustart 1994; cowling et al. 1999) and wetland biota (de villiers 1988; ryan et al. 1988; rebelo 1992). the heuningnes river system is recognised as a priority system for freshwater fish conservation in the cape floristic region (impson et al. 1999). formal conservation of the area commenced in september 1999 with the proclamation of the agulhas national park (anp). there is an ongoing park expansion program with the objective of including representative examples of both terrestrial and aquatic ecosystems. where the terrestrial vegetation of the region has been the subject of intensive research (e.g., cowling et al. 1988; cowling & holmes 1992a; willis et al. 1996; cole et al. 2000), studies of aquatic systems have mostly been once-off assessments (silberhauer & king 1991; jones et al. 2000) and concentrated predominantly on water chemistry. there are few accounts of aquatic biota, though the area is recognised as supporting a diverse aquatic macro-invertebrate fauna (j. king pers. comm.). the objective of this study was to improve understanding of the aquatic biota of the region, specifically by describing the diversity and distribution of fishes in and adjacent to the park. the in-park surveys also form a component of a long-term research program to assess the contribution national parks in south africa make towards the conservation of south aquatic systems in and adjacent to agulhas national park with particular reference to the fish fauna i.a. russell and n.d. impson russell, i.a. and n.d. impson. 2006. aquatic systems in and adjacent to agulhas national park with particular reference to the fish fauna. koedoe 49(2): 45–57. pretoria. issn 0075-6458. the study aimed to determine the distribution and relative abundance of freshwater fishes in and adjacent to agulhas national park. fourteen fish species were collected during surveys carried out in the heuningnes and ratel river systems and a variety of endorheic pans from 2003 to 2005. seven of the species are marine fishes tolerant of low salinity (mugil cephalus, liza richardsonii, monodactylus falciformis, caffrogobius gilchristi, rhabdosargus holubi, solea bleekeri, gilchristella aestuaria) with the remaining seven species being primary freshwater fishes. three are indigenous, including pseudobarbus burchelli, sandelia capensis, and galaxias zebratus of which two morphological forms were recorded within agulhas national park. the remaining four species are invasive aliens (cyprinus carpio, lepomis macrochirus, micropterus salmoides, micropterus punctulatus). classification using physico-chemical variables indicated four major wetland groups, with fishes occurring predominantly in the group comprising palustrine, lacustrine and riverine wetlands, and in one instance in a brackish endorheic pan. management actions which should facilitate conservation of indigenous fishes in the agulhas region are suggested. key words: diversity, freshwater fish, conservation, wetlands, agulhas national park, soetendalsvlei, heuningnes river, nuwejaars river, ratel river. i.a. russell, south african national parks, p.o. box 176, sedgefield, 6573. south africa; n.d. impson, scientific services, western cape nature conservation board, jonkers­ hoek, stellenbosch, 7600 south africa. russell.indd 45 2006/10/15 10:58:32 pm koedoe 49/2 (2006) 46 issn 0075-6458 african freshwater fishes. the off-park surveys formed component of the overberg river-health survey which is part of the south african river health program, the main purpose of which is to serve as a source of information regarding the overall ecological status of river ecosystems in south africa. study area the anp is situated between latitude 34º40's– 34º50's and longitude 19º38'e–20º02'e in the western cape province of south africa, and covers an area of approximately 16 807 ha. typical rainfall is 450–480 mm/y (cowling & witkowski 1994), with more than 65 % falling in winter between may and october (thwaites & cowling 1988). three different geological substrates occur in the agulhas area, namely bokkeveld shale (voëlvlei) malmesbury shale (soutpan and buffeljacht pan) and bredasdorp limestone (die pan) (carr 2005). soil types in the agulhas area are highly varied, ranging from coarse sands along the coast, alluvial or colluvial topsoils over residual or transported clays in the east, grey calcareous sands in the central areas, and acidic, highly leached, soils in western areas (thwaites & cowling 1988). terrestrial vegetation is extremely diverse (> 1 750 species) with pronounced edaphic endemism, and consists of stands of restioid fynbos, dune fynbos, limestone fynbos, acid sand proteoid fynbos, ericaceous fynbos, and renosterveld (cowling & holmes 1992b; low & rebelo 1996). wet restioid fynbos is frequently found on the low-lying, seasonally waterlogged areas, and around larger pans phragmites reeds dominate (mustart et al. 1997). the main river systems in and adjacent to the anp are the heuningnes and ratel. the heuningnes river has two major tributaries, the nuwejaars and kars rivers. the nuwejaars river flows through the north-eastern reaches of the park into soetendalsvlei. mean annual runoff (mar) is 37.6 x 106 m3 (raimondo & barker 1988). large seasonally inundated wetlands are associated with its valley, though most occur outside the park. soetenfig. 1. map of the agulhas national park indicating sites used to sample fish assemblages. 19°45' 19°50' 19°55' 20°00'19°40' 34°50' 34°45' 34°40' 7 atlantic ocean 2 3 4 5 1 8 9 6 12 13 14 15 11 17 18 19 16 10 n port elizabeth durban cape town south africa heuningnes river rivers park boundary endorheic wetland sample site marsh lacustrine wetland 3 0 5 km soetendalsvlei voëlvlei melkbos pan die pan soutpan renosterkop pan palustrine wetland nuwejaars river ratel river drievleitjies springfield pans kars river russell.indd 46 2006/10/15 10:58:32 pm issn 0075-6458 47 koedoe 49/2 (2006) dalsvlei is connected to the indian ocean via the heuningnes river. the kars river arises on the northern slopes of the soetmuisberg and bredasdorp mountains before flowing southwards and entering karsriviervlei and the heuningnes river, near the park. the ratel river in the western reaches of anp is a much smaller system, with a mar of 7.05 x 106 m3 (raimondo & barker 1988). extensive wetlands occur along its lower reaches before it flows into the atlantic ocean. several endorheic pans occur within the park. they have been formed and are maintained by the interplay of a variety of geomorphological processes, including drainage disruption, aeolian activity, and the solution and weathering of the substrate (goudie 1991; marshall & harmse 1992; shaw & thomas 1997). the larger bodies of water such as soetendalsvlei and voëlvlei, which are directly connected to the nuwejaars river, are lake-like in that they remain inundated year round. many of the smaller pans are not connected to any fluvial system or only have small locally sourced channels entering them. they are ephemeral with an annual inundation/drying regime—a function of winter rainfall and subsequent summer evaporation. soetendalsvlei (sample site 1) is described by silberbauer & king (1991) as moderately saline (2–5 g/kg), whereas several of the smaller pans (for example, sample site 12 soutpan, sample site 7 melkbospan, and sample site 6 vispan) are strongly saline (16– 86 g/kg). river waters are brackish (< 2 g/kg salts) and alkaline as a result of passage through limestone-bearing strandvlei sands (noble & hemens 1978). the ph in most of the lowland wetlands varies between 6 and 8 (silberbauer & king 1991), though in soetendalsvlei ph of up to 8.5 has been recorded (noble & hemens 1978). methods surveys were undertaken by sanparks personnel within agulhas national park, forming the main component of this assessment, and by western cape river health practitioners in rivers outside of anp as part of a broad survey of rivers of the overberg region. for the sanparks assessment, 19 sample sites were used of which three were in the nuwejaars river system, two in the ratel river system, and the remaining 14 in pans (fig. 1). fish sampling and water quality assessments were undertaken from 1–5 december 2003 (sample sites 1–18) and on 11 march 2004 (sample site 19). for the river-health assessment, seven sample sites were surveyed in the heuningnes river system (fig. 2) on 12 october 2005. consecutive numbers after a hash symbol were assigned to different pans with the same name, for example melkbospan # 1 and melkbospan # 2, to differentiate between sample sites. water quality parameters measured in the field at the time of fish sampling in anp were: water temperature (°c) and salinity (g/kg) using a ysi model 33 s-c-t meter; dissolved oxygen (mg/l) using a ysi 550a o2 meter; and ph (ysi 60 ph meter). water samples were transported to a laboratory where turbidity (ntu) was measured with a hach 16800 turbidimeter. parameters measured in the heuningnes river system during river-health assessments were conductivity (μ mhos), ph and dissolved oxygen (mg/l) using portable ysi meters. sampling for fishes in soetendalsvlei (sample site 1) was done with two brown multifilament gillnets (40 m x 2 m x 55 mm and 80 mm) and one monofilament gillnet (20 m x 2 m x 100 mm), a beach seine net (40 m x 3 m x 3 mm), with 50 m warps, and baited longlines (two 20 m lines each with 20 hooks). gillnets and longlines were set overnight from 16:00 to 09:00. sampling in the nuwejaars river (sample sites 9 & 10) and ratel river (sample sites 2 & 3) was undertaken with a hand-held electro-fishing apparatus, powered by a 220v ac, 1.8 kva portable generator. anchovy seine nets operated by two persons were used to collect fishes in pans within the park (10 m x 1.5 m x 3 mm) as well as in the heuningnes river (5 m x 2 m x 3 mm) during the river-health assessment. a dipnet (sass net) was used to sample fishes in the pietersielieskloof and kleinpietersielies rivers as the water was too shallow to use a seine net. fishes collected were identified using keys in skelton (2001) and whitfield (1998), and counted. identification of species collected during river-health assessments was based entirely on field observations, whereas identification from sanparks surveys were based predominantly, and in the case of galaxias zebratus castelnau, 1861 entirely on assessment of museum voucher specimens now housed at south african institute for aquatic biodiversity (saiab) in grahamstown. (voucher numbers for specimens collected in anp: sandelia capensis = 77372, 77379, 77383, 77384, 77387; galaxias zebratus (form a) = 77373, 77378, 77386; galaxias zebratus (form b) = 77385; mugil cephalus = 77375; liza richardsonii = 77376; gilchristella aestuaria = 77382; caffrogobius gilchristi = 77377; cyprinus carpio = 77380; microp­ terus punctulatus = 77374, 77381). gilchristella aes­ tuaria (gilchrist, 1913) collected in soetendalsvlei russell.indd 47 2006/10/15 10:58:32 pm koedoe 49/2 (2006) 48 issn 0075-6458 atlantic ocean n 20 21 22 23 24 26 25 0 21 km 7 14 heuningnes river kars rivergrashoek river nuwejaars river ratel river pietersielieskloof klein pietersielies kersgat rooidraaibrug schietpad brakpan riverside soutkloof kastaiinskloof river coetzeeskraal rivers park boundary sample site lacustrine wetland 3 catchment boundary 19°50'19°40' 19°70'19°60'19°30' 34°50' 34°40' 34°30' 34°20' fig. 2. map of the heuningnes catchment showing fish survey sites of the river health assessment. were sub-sampled, with live individuals counted in lots of approximately 100 and released. at the time of writing, galaxias zebratus is still recognised as a single species, though anatomical differences within the meta population indicate the occurrence of different forms that, with closer investigation, may be accorded full species status (e. swartz pers. comm.). two forms were described from specimens collected in anp, defined here as “form a” and “form b”. galaxias zebratus (form a) is characterised by having an elongated body, small eyes, broad caudal peduncle and distinctive striped body markings, and g. zebratus (form b) characterised by having a deeper body form, large eyes, narrow caudal peduncle, and body markings being either very light or absent (e. swartz pers. comm.). as species collected outside of anp were assessed only in the field, no distinction was made between different forms and are referred to here as g. zebratus. for g. zebratus within anp, distinction is made between the two forms. similarly, at the time of writing, pseudobarbus burchelli smith, 1841 is still recognised as a single species, though genetic work by swartz (2005) on the pseudobarbus group indicates that the population from the grashoek river in the heuningnes river system is divergent from other p. burchelli populations in the breede system. although the heuningnes population could well, following closer investigation, be accorded full species status (e. swartz pers. comm.) for consistency this species has continued to be described here as p. burchelli. observations were made at sample sites within anp of the distribution of emergent aquatic plants. they were considered to be wide-spread or abundant if established stands occurred on > 50 % of wetland margin, of intermediate abundance if the margin colonised was between 20 % and 50 %, and sparse if < 20 % of the margin was colonised. the same proportions were applied to submerged macrophytes and epipsammic algae with respect to the percentage of substratum colonised in sample areas. cluster analysis was performed on water quality parameters at the different sites within anp, using ward’s method and euclidean metric as distance measure in statgraphics, version 4.0. russell.indd 48 2006/10/15 10:58:33 pm issn 0075-6458 49 koedoe 49/2 (2006) results water quality the physical and chemical parameters of the wetlands sampled within agulhas national park showed wide variability (table 1). daytime water temperature varied 11.9 °c between waterbodies, with the coolest water (< 22 °c) occurring in the deep vegetated palustrine wetlands of the ratel river system, and the warmest (> 30 °c) in shallow saline pans such as springfield pan # 1, renosterkop pan and melkbospan # 1. there was large variability in salinity between waterbodies. the most saline water sampled (agulhas soutpan) was 990 times that of the least saline (ratel river # 2). the range in ph between waterbodies was also wide, with waters in the ratel river being slightly acidic (6.4–6.8), and pans such as springfield pan # 1 and die vleitjies # 2 being strongly alkaline (ph = 10). water in all endorheic pans was alkaline, with highly saline pans (> 100 g/kg) being weakly alkaline (ph 7.3– 7.8) whereas in moderately saline and brackish pans (< 60 g/kg) alkalinity was generally higher (ph 8.1–10.0). dissolved oxygen was consistently low (5.2–3.7 mg/l) in the highly saline waterbodies. a wider range in dissolved oxygen values were recorded in moderately saline to fresh waterbodies (5.2–10.7 mg/l), with dissolved oxygen in most waterbodies being high (> 7 mg/l). variability in turbidity between waterbodies was very high, with the most turbid waterbody (die pan) being 4213 times more turbid than the nearby and least turbid waterbody (melkbospan # 2). ichthyofauna eleven fish species were collected in anp (table 1). the species composition in soetendalsvlei showed a strong marine influence with the collection of two estuarine species (caffrogobius gilchristi (boulenger, 1900), liza richardsonii (smith, 1846)), and three species (g. aestuaria, monodactylus fal­ ciformis lacepède, 1800, mugil cephalus linnaeus, 1758) which, although occasionally found in coastal rivers (skelton 2001), occur predominantly in estuaries or inshore coastal waters (whitfield 1998). no indigenous freshwater fishes were collected in soetendalsvlei, though four alien freshwater fishes (cyprinus carpio linnaeus, 1758, lepomis macrochirus rafinesque, 1819, micropterus salmoides (lacepède, 1802), micropterus punctulatus (rafinesque, 1819)) have established populations. in the nuwejaars river which flows into soetendalsvlei, both estuarine (m. cephalus, g. aestuaria) and alien (c. carpio, m. punctulatus) fishes persist, along with the indigenous freshwater sandelia capensis (cuvier, 1831). the ratel river contained two indigenous freshwater fishes (s. capensis, g. zebratus) with s. cap­ ensis populations also occurring in the lakelike pans (buffelsjacht and ratel river pan) associated with this river system. an isolated population of g. zebratus (form a) occurs in melkbospan # 2. in the rivers adjacent to the anp, 11 species were collected during the river health assessment (table 2). all three species of indigenous primary freshwater fishes expected for the region were recorded, including p. burchelli. sandelia capensis and g. zebratus were widespread and fairly common. both forms (a and b) of cape galaxias were recorded, notably in the pietersielieskloof catchment. three invasive alien freshwater fishes were recorded, namely c. carpio, l. macrochirus and m. punctulatus. these species are probably more widespread than indicated in the survey, as several sites had deep wide pools where use of the small seine was ineffective. a single site near the heuningnes estuary at the struisbaai road bridge was sampled which yielded an unusual catch comprising two primary freshwater fishes, one alien (c. carpio) and one indigenous (s. capensis), and several estuarine-marine species (g. aestuaria, m. falciformis, c. gilchristi, rhabdosargus holubi (steindachner, 1881) and solea bleekeri (boulenger, 1898)). wetland classification: cluster analysis revealed four distinctive wetland groups (fig. 3): russell.indd 49 2006/10/15 10:58:33 pm koedoe 49/2 (2006) 50 issn 0075-6458 fish sampling fish species site description water quality methods marine freshwater alien s ite n um be r s ite n am e l at itu de ( so ut h) l on gi tu de ( e as t) t em pe ra tu re ( ºc ) s al in ity ( m g/ kg ) p h ( ph u ni ts ) d is so lv ed o xy ge n (m g/ l) t ur bi di ty ( n t u ) g ill n et tin g (1 2 ho ur s et s) s ei ne n et tin g (e ff ec tiv e ha ul s) e le ct ro -f is hi ng ( m in ut es ) l on glin es ( 12 h ou r se ts ) m ug il ce ph al us l iz a ri ch ar ds on ii g ilc hr is te lla a es tu ar ia m on od ac ty lu s fa lc ifo rm is c af fr og ob iu s gi lc hr is ti s an de lia c ap en si s g al ax ia s ze br at us f or m a g al ax ia s ze br at us f or m s a & b c yp ri nu s ca rp io l ep om is m ac ro ch ir us m ic ro pt er us p un ct ul at us m ic ro pt er us s al m oi de s group 1: riverine, palustrine & lacustrine wetlands 3 ratel river # 2 34°43’58” 19°43’10” 21.4 0.2 6.8 7.9 8.6 15 214 3 2 rater river # 1 34°45’15” 19°43’48” 24.4 0.3 6.4 7.6 7.9 60 40 58 19 ratel river pan 34°44’35” 19°40’30” 20.5 0.9 7.5 8.3 264 3 78 17 buffelsjag 34°44’30” 19°39’30” 21.5 5.3 8.1 8.4 9.5 1 31 10 nuwejaars river # 2 34°43’28” 19°57’40” 22.7 1.2 7.8 6.5 2.5 60 1 1 15 1 9 nuwejaars river # 1 34°43’18” 19°57’05” 21.4 1.8 7.7 7.2 36 60 2 35 24 33 11 1 soetendalsvlei 34°44’20” 19°59’40” 23.1 1.0 7.6 7.8 13 1 2 1 44 15 4750 81 1 9 1 25 5 group 2: brackish endorheic wetlands 15 springfield pan # 2 34°44’30” 19°54’50” 25.4 3.9 8.7 10.7 3.8 1 18 drie vleitjies # 2 34°45’05” 19°45’10” 22.3 9.5 10.0 9.4 3.6 1 8 melkbos pan # 2 34°43’20” 19°45’30” 25.5 3.0 9.3 8.2 1.5 2 61 14 springfield pan # 1 34°44’15” 19°54’40” 32.4 7.1 10.0 9.5 5.6 1 13 soutpan extension 34°43’35” 19°54’40” 28.4 5.8 9.6 9.6 2.1 1 6 drie vleitjies # 1 34°45’00” 19°45’20” 27.8 17.5 9.9 10.3 3.4 1 group 3: saline endorheic wetlands 11 soutbos pan 34°43’23” 19°56’35” 27.6 41.4 8.6 5.2 83 1 16 renostekop pan 34°45’35” 19°55’30” 30.7 52.8 8.8 7.2 6.0 1 4 rietfontein pan 34°44’20” 19°44’20” 28.9 58.5 9.2 6.4 9.2 1 group 4: highly saline endorheic wetlands 7 melkbos pan # 1 34°43’15” 19°45’20” 30.1 162.0 7.4 3.7 55 1 12 agulhas soutpan 34°43’20” 19°55’20” 29.7 198.0 7.3 4.8 3843 1 5 die pan 34°44’40” 19°45’20” 27.0 112.8 7.8 5.2 6319 1 table 1 number of fish specimens per species collected in the agulhas national park during december 2003 and march 2004, and physico­chemical parameters at the time of fish sampling. site numbers as per fig. 1 russell.indd 50 2006/10/15 10:58:34 pm issn 0075-6458 51 koedoe 49/2 (2006) fish sampling fish species site description water quality methods marine freshwater alien s ite n um be r s ite n am e l at itu de ( so ut h) l on gi tu de ( e as t) t em pe ra tu re ( ºc ) s al in ity ( m g/ kg ) p h ( ph u ni ts ) d is so lv ed o xy ge n (m g/ l) t ur bi di ty ( n t u ) g ill n et tin g (1 2 ho ur s et s) s ei ne n et tin g (e ff ec tiv e ha ul s) e le ct ro -f is hi ng ( m in ut es ) l on glin es ( 12 h ou r se ts ) m ug il ce ph al us l iz a ri ch ar ds on ii g ilc hr is te lla a es tu ar ia m on od ac ty lu s fa lc ifo rm is c af fr og ob iu s gi lc hr is ti s an de lia c ap en si s g al ax ia s ze br at us f or m a g al ax ia s ze br at us f or m s a & b c yp ri nu s ca rp io l ep om is m ac ro ch ir us m ic ro pt er us p un ct ul at us m ic ro pt er us s al m oi de s group 1: riverine, palustrine & lacustrine wetlands 3 ratel river # 2 34°43’58” 19°43’10” 21.4 0.2 6.8 7.9 8.6 15 214 3 2 rater river # 1 34°45’15” 19°43’48” 24.4 0.3 6.4 7.6 7.9 60 40 58 19 ratel river pan 34°44’35” 19°40’30” 20.5 0.9 7.5 8.3 264 3 78 17 buffelsjag 34°44’30” 19°39’30” 21.5 5.3 8.1 8.4 9.5 1 31 10 nuwejaars river # 2 34°43’28” 19°57’40” 22.7 1.2 7.8 6.5 2.5 60 1 1 15 1 9 nuwejaars river # 1 34°43’18” 19°57’05” 21.4 1.8 7.7 7.2 36 60 2 35 24 33 11 1 soetendalsvlei 34°44’20” 19°59’40” 23.1 1.0 7.6 7.8 13 1 2 1 44 15 4750 81 1 9 1 25 5 group 2: brackish endorheic wetlands 15 springfield pan # 2 34°44’30” 19°54’50” 25.4 3.9 8.7 10.7 3.8 1 18 drie vleitjies # 2 34°45’05” 19°45’10” 22.3 9.5 10.0 9.4 3.6 1 8 melkbos pan # 2 34°43’20” 19°45’30” 25.5 3.0 9.3 8.2 1.5 2 61 14 springfield pan # 1 34°44’15” 19°54’40” 32.4 7.1 10.0 9.5 5.6 1 13 soutpan extension 34°43’35” 19°54’40” 28.4 5.8 9.6 9.6 2.1 1 6 drie vleitjies # 1 34°45’00” 19°45’20” 27.8 17.5 9.9 10.3 3.4 1 group 3: saline endorheic wetlands 11 soutbos pan 34°43’23” 19°56’35” 27.6 41.4 8.6 5.2 83 1 16 renostekop pan 34°45’35” 19°55’30” 30.7 52.8 8.8 7.2 6.0 1 4 rietfontein pan 34°44’20” 19°44’20” 28.9 58.5 9.2 6.4 9.2 1 group 4: highly saline endorheic wetlands 7 melkbos pan # 1 34°43’15” 19°45’20” 30.1 162.0 7.4 3.7 55 1 12 agulhas soutpan 34°43’20” 19°55’20” 29.7 198.0 7.3 4.8 3843 1 5 die pan 34°44’40” 19°45’20” 27.0 112.8 7.8 5.2 6319 1 russell.indd 51 2006/10/15 10:58:34 pm koedoe 49/2 (2006) 52 issn 0075-6458 fish sampling fish species site description water quality methods marine freshwater alien s ite n um be r s ite n am e l at itu de ( so ut h) l on gi tu de ( e as t) c on du ct iv ity ( μ m ho s) p h ( ph u ni ts ) d is so lv ed o xy ge n (m g/ l) d ip n et ( m in ut es s am pl in g) s ei ne n et tin g (e ff ec tiv e ha ul s) r ha bd os ar gu s ho lu bi c af fr og ob iu s gi lc hr is ti g ilc hr is te lla a es tu ar ia m on od ac ty lu s fa lc ifo rm is s ol ea b le ek er i s an de lia c ap en si s p se od ob ar bu s bu rc he lli s p. g al ax ia s ze br at us c yp ri nu s ca rp io l ep om is m ac ro ch ir us m ic ro pt er us p un ct ul at us 20 kars river # 1 34°41’33” 19°82’06” 62 5.7 8.8 1 7 >100 17 21 kars river # 2 34°42’97” 19°91’53” 7.0 7.4 2 8 8 1 22 nuwejaars river # 1 34°57’83” 19°70’79” 60 6.7 6.3 2 3 >50 8 23 nuwejaars river # 2 34°63’40” 19°86’50” 91 6.7 3.0 2 11 10 24 klein pietersielies 34°54’73” 19°80’73” 43 6.1 8.8 5 5 25 pietersielieskloof 34°54’23” 19°81’87” 38 5.3 6.7 5 12 26 heuningness 34°68’86” 20°03’36” 7.9 7.8 3 5 >10 >50 >10 >10 6 2 table 2 fish species caught at seven sites on the heuningness catchment during the october 2005 river­health assessment. included are average scores for three water quality parameters taken during the seasonal assessment. site numbers as per fig. 2 group 1. palustrine, lacustrine and riverine wetlands waterbodies in this group include the ratel and nuwejaars rivers, and the large (1 853 ha) permanently inundated lacustrine soetendalsvlei, and the deep (> 1 m) palustrine waterbodies of buffeljacht pan and ratel river pan in the ratel river system. salinity in these waterbodies is low (0.2–5.3 g/kg), ph ranges from slightly acid to moderately alkaline (ph 6.4–8.1), dissolved oxygen is moderate (6.5–8.4 mg/kg), and turbidity is highly variable (2–264 ntu) though is low in most waterbodies. water temperature (20.5–24.4 °c) is substantially lower (approx. 7 °c) than that recorded in large, shallow, endorheic wetlands in the region. these wetlands are associated with major drainage lines. most wetlands support submerged macrophytes, which in some cases (e.g. ratel river pan) are abundant, and all have extensive stands of emergent macrophytes, principally phragmites australis (cavanilles) trinius ex steudel and typha capensis (rohrbach). all waterbodies support fish communities. the ratel river sites form a subgroup by virtue of waters being acidic (ph < 6.8) and having very low salinity (< 0.3 g/kg). group 2. brackish endorheic wetlands this group of six wetlands consists predominantly of smaller endorheic pans located on the periphery of large highly saline pans, and presumably forming part of the drainage system of these waterbodies. there is large variability in salinity (3.0–17.5 g/kg) though in most waterbodies is below 10.0 g/kg. waters are characterised by being strongly alkaline (ph 8.7–10.7), having high dissolved oxygen (8.2–10.7 mg/kg) and very low turbidity (2–6 ntu). these wetlands form two subgroups (fig. 2) which differ principally in terms of water temperature, with drie vleitjies # 1, soutpan extension and springrussell.indd 52 2006/10/15 10:58:34 pm issn 0075-6458 53 koedoe 49/2 (2006) field pan # 1 being warmer (27.8–32.4 °c) at the time of sampling than were melkbos pan # 2, drie vleitjies # 2, and springfield pan # 2 (22.3–25.5 °c). submerged rooted macrophytes and/or filamentous algae occur in most waterbodies though are generally not abundant. emergent macrophytes occur on the margins of most waterbodies, though stands are generally sparse. fishes occur in one waterbody, namely an isolated population of g. zebratus (form a) in the brackish (3.0 g/kg) well-vegetated melkbos pan # 2. group 3. saline endorheic wetlands this wetland group consists of three, small, shallow pans which are geographically removed from the larger pan clusters, and are not obviously associated with any of the major drainage lines. the salinity of waters is high (41.4–58.5 g/kg), ph is strongly alkaline (8.6–9.2), dissolved oxygen is moderate (5.2–7.2 mg/kg), and turbidity is moderate to fish sampling fish species site description water quality methods marine freshwater alien s ite n um be r s ite n am e l at itu de ( so ut h) l on gi tu de ( e as t) c on du ct iv ity ( μ m ho s) p h ( ph u ni ts ) d is so lv ed o xy ge n (m g/ l) d ip n et ( m in ut es s am pl in g) s ei ne n et tin g (e ff ec tiv e ha ul s) r ha bd os ar gu s ho lu bi c af fr og ob iu s gi lc hr is ti g ilc hr is te lla a es tu ar ia m on od ac ty lu s fa lc ifo rm is s ol ea b le ek er i s an de lia c ap en si s p se od ob ar bu s bu rc he lli s p. g al ax ia s ze br at us c yp ri nu s ca rp io l ep om is m ac ro ch ir us m ic ro pt er us p un ct ul at us 20 kars river # 1 34°41’33” 19°82’06” 62 5.7 8.8 1 7 >100 17 21 kars river # 2 34°42’97” 19°91’53” 7.0 7.4 2 8 8 1 22 nuwejaars river # 1 34°57’83” 19°70’79” 60 6.7 6.3 2 3 >50 8 23 nuwejaars river # 2 34°63’40” 19°86’50” 91 6.7 3.0 2 11 10 24 klein pietersielies 34°54’73” 19°80’73” 43 6.1 8.8 5 5 25 pietersielieskloof 34°54’23” 19°81’87” 38 5.3 6.7 5 12 26 heuningness 34°68’86” 20°03’36” 7.9 7.8 3 5 >10 >50 >10 >10 6 2 1 9 10 17 19 2 3 4 16 11 6 13 14 8 18 15 5 12 7 0 5 10 15 20 25 30 highly saline endorheic brackish endorheic saline endorheic palustrine, lacustrine & riverine nuwejaars system ratel river ratel system lakes cool deep pans warm shallow pans salt pans large, turbid salt pans large, clear salt pan euclidean distance sample site number fig. 3. classification of wetland systems within agulhas national park based physico-chemical variables (water temperature, salinity, ph, dissolved oxygen, turbidity) as determined during fish surveys undertaken in during december 2003 and march 2004. russell.indd 53 2006/10/15 10:58:34 pm koedoe 49/2 (2006) 54 issn 0075-6458 high (6–83 ntu). rooted submerged macrophytes are absent, though filamentous algae is abundant. marginal wetland plants are sparse, consisting predominantly of juncus and other rush species. fishes are absent. group 4. highly saline endorheic wetlands three waterbodies fall into this group, namely die pan, agulhas soutpan, and melkbos pan # 1. these are large, shallow (< 1 m) pans characterised by waters which are highly saline (112.8–198.0 g/kg), alkaline (ph 7.3– 7.8), with dissolved oxygen levels that are very low (3.7–5.2 mg/kg) and turbidity very high (55–6319 ntu). the substratum consists of fine, grey-white sediments which are easily disturbed and possibly contribute to periodic high turbidity levels in the shallow waters. both submerged and emergent aquatic plants appear to be largely absent, as are fishes. discussion comparative water quality assessments (salinity, ph, turbidity) undertaken in 13 agulhas wetlands in earlier studies (silberbauer & king 1991; jones et al. 2000) demonstrate, in most cases, similar ranges for wetland groups despite seasonal differences in sampling. substantial differences with this study were that jones et al. (2000) reported renosterkop pan to be both highly saline (> 100 g/kg) and turbid (203 ntu), and silberbauer & king (1991) reported the three pans (melkbos pan # 1; agulhas soutpan; die pan) categorised in this study as highly saline (> 100 g/kg) to be only moderately saline (16, 39 & 25 g/kg respectively). furthermore, these same three pans were found by jones et al. (2000) to be moderately turbid (31–52 ntu) compared to the extremely high turbidity recorded in 2003. high variability in water chemistry, particularly salinity, in saline endorheic wetlands is likely to be a result primarily of the annual inundation/drying regime, and suggests that the distinction between saline (group 3) and highly saline (group 4) wetlands (fig. 2) may be an artefact of seasonspecific sampling rather than reflecting a fundamental difference in physico-chemical characteristics. jones et al. (2000) grouped agulhas wetlands primarily on geographical similarities, suggesting that wetlands in the same area frequently share characteristics. the present data indicate that hydrological characteristics may be a more informative means of classifying agulhas wetlands, particularly with respect to the occurrence of biota such as fishes. however, longer term assessment of the physico-chemical properties of agulhas wetlands, incorporating both seasonal and annual variability, is needed for more informative classification and characterisation of wetland types. few published data exist on ichthyofauna of the agulhas area. bickerton (1984) cites unpublished reports on fishes previously collected in soetendalsvlei (cf. cape piscatorial society 1937; barham 1968; louw 1968; mcveigh 1980), with the resultant species count differing from current surveys in respect of lithognathus lithognathus (cuvier, 1830) being previously recorded, and the new record of c. gilchristi. both l. lithog­ nathus and c. gilchristi have been recorded in low salinity waters (< 3 g/kg) (whitfield 1998) though they are not species commonly associated with the lower reaches of freshwater systems, as are the other marine fishes recorded in soetendalsvlei (skelton 2001). it is thus not expected that either species would be common in the predominantly freshwater soetendalsvlei. all three indigenous freshwater fish species expected to occur in the agulhas plain, namely p. burchelli, g. zebratus and s. capensis were recorded, with populations of the latter two species occurring within anp. published records indicate g. zebratus as the only fish species in the ratel river system (cape department of nature and environmental conservation 1984). however, the occurrence of s. capensis within the ratel river and associated palustrine pans is not unexpected given that is has been recorded in adjacent systems (saiab species distribution database). the comparatively high abundance of s. capensis in the ratel system can possibly be ascribed to the wide habitat tolerance russell.indd 54 2006/10/15 10:58:34 pm issn 0075-6458 55 koedoe 49/2 (2006) of this species, particularly with respect to the seasonality of waterbodies (jubb 1967), combined with the apparent absence of alien predators in the ratel river. the occurrence of g. zebratus (form a) in melkbos pan # 2 is unusual, being the only fish population in an endorheic pan in agulhas national park. there is no information on how this species came to occur in this wetland. in the absence of any records suggesting that this unusual distribution is the result of introduction by humans, the conservation of this isolated population should be afforded high priority. the distribution of p. burchelli is given by cape department of nature and environmental conservation (1984) and skelton (1988) to include the heuningnes river, which concurs with a species distribution records from the 1940s given by gaigher (1978). cambray & stuart (1985) failed to collect specimens during 1980, though recent surveys in 2004 yielded specimens from the uppermost reaches of the grashoek river in the heuningnes river system (swartz 2005). this assessment yielded new distribution records from the upper kars river. furthermore, in february 2006, a local farmer on the property coetzeeskraal, next to the kastaiingskloof river (fig. 2), requested that capenature remove indigenous fish from a small offstream dam that was drying up. this dam receives flow from the river via a furrow. seine netting yielded a thriving community of p. burchelli, g. zebratus and s. capensis, suggesting that these species may still be present in the kastaiinskloof river. the absence of p. burchelli from survey sites in the remainder of this system is a major cause for concern and appears associated with the presence of the piscivorous m. punctulatus, and major habitat damage caused by the floods of february 2005. three invasive alien freshwater fishes were recorded during the survey, namely c. car­ pio, l. macrochirus and m. punctulatus. the latter two species are widespread and common, especially in the larger rivers. the date and reason for the introduction of c. carpio is uncertain, although it is known that this species was extensively stocked as an angling and food fish into rivers across the south africa in the late 1800s and early 1900s (de moor & bruton 1988). cyprinus carpio can be detrimental to indigenous fishes when present in large numbers in that it consumes the eggs of other fishes, and can cause habitat degradation by removing aquatic plants and increasing turbidity (de moor & bruton 1988). cyprinus carpio may be more widespread than these surveys reflect and future fish surveys should utilise gill nets in the bigger deeper pools. micropterus punctulatus was introduced into the nuwejaars river at elim in 1940 (harrison 1977). de moor & bruton (1988), however, suggest that its preference for large, turbid rivers and intolerance of low ph waters will exclude it from establishing populations in south coastal drainage rivers, where environmental conditions would, for the most part, be unfavourable. distribution maps in skelton (2001) do not indicate m. punctulatus as occurring in the agulhas region, though its continued occurrence and now relatively high abundance in soetendalsvlei and associated rivers indicate that this invasive alien is well established in the lower nuwejaars and kars rivers. it is likely that l. macrochirus were stocked into the heuningnes system shortly after the introduction of m. punctulatus. lepomis macrochirus were introduced into south africa as a fodder fish for bass, when it became apparent that bass had eliminated the smaller indigenous fish species, and thus required an alternative prey source. this species, especially when abundant, can have a major impact on smaller indigenous fishes through competition and predation (de moor 1988). it is regarded as a pest species as it has negligible angling value while causing considerable ecological damage in certain areas. where these alien species occur, their impact on p. burchelli and s. capensis has likely been severe, resulting in poor recruitment and localised extirpations. galaxias zebratus russell.indd 55 2006/10/15 10:58:35 pm koedoe 49/2 (2006) 56 issn 0075-6458 seems to have fared better than the other two species, especially in rivers with abundant submerged aquatic plants, where it could escape predation due to its cryptic colouration and small size. one additional freshwater fish species, namely anguilla mossambica peters, 1852 could potentially occur in anp and surrounds. the natural range of a. mossambica on the african continent includes east coast rivers from kenya to cape agulhas (skelton 2001). the anp rivers would consequently be at the southernmost range of a. mossambica which could possibly result in it being uncommon, and hence not yet recorded in the study area. recommendations the following recommendations are suggested to improve conservation of freshwater fish and wetlands in the agulhas area: conservation agencies should endeavour to improve public awareness of the detrimental consequences of introducing alien fishes into dams and rivers. as an alternative to alien fishes, local indigenous fishes can be stocked into dams in the catchment. this action should form part of a documented recovery program for these fishes. efforts should be made to improve the conservation status of areas where p. burchelli still occurs. these could include land purchase and stewardship arrangements. land-owner awareness of these fishes is a critical factor. the eradication of invasive alien vegetation from riparian zones and wetlands is a priority for restoring flow and rehabilitating severely and moderately invaded systems. fish populations should be monitored on a regular basis, and where required information collected, or specimens retained, that would facilitate taxonomic identification and reassessment of indigenous freshwater species. acknowledgements thanks are due to eddy kivett, suzaan kritzinger and bronwyn botha for assistance with the field surveys in agulhas national park. verification of taxa by dr. e. swartz of saiab is gratefully acknowledged. riki de villiers of capenature prepared fig. 2. we are also grateful to dr. j. cambray, dr. e. swartz and one anonymous reviewer for their critical comments on an earlier version of the manuscript. references barham, w.t. 1968. report on survey of zoetendalsvlei lake, district of bredasdorp. cpa department of nature and environmental conservation. unpublished report. bickerton, i.b. 1984. heuningnes (csw 19). report no. 25. in: heydorn, a.e.f. & j.r. grindley (eds.). estuaries of the cape. part ii: synopsis of available information on individual systems. stellenbosch: council for scientific and industrial research (csir research report; no. 424). cape department of nature and environmental conservation. 1984. freshwater fishes of the cape. cape town: cape department of nature and environmental conservation. (cape conservation series; no. 5). cape piscatorial society. 1937. the most southerly lake in africa: zoetendals vlei, about five miles from cape agulhas, bredasdorp division, cape province. circular 27. carr, a. 2005. late quarternary environmental change on the agulhas plain, winter rainfall zone, south africa. phd thesis, university of sheffield, sheffield. cole, n.s., a.t. lombard, r.m. cowling, d. euston-brown, d.m. richardson & c.e. heijnis. 2000. framework for a conservation plan for the agulhas plain, cape floristic region, south africa. cape town: university of cape town, institute for plant conservation. (ipc report 0001). cowling, r.m. & p.m. holmes 1992a. endemism and speciation in a lowland flora from the cape floristic region. biological journal of the linnaean society 47:367–383. cowling, r.m. & p.m. holmes. 1992b. flora and vegetation. pp. 23–63. in: cowling, r.m. (ed.). the ecology of the fynbos: nutrients, fire and diversity. cape town: oxford university press. cowling, r.m. & p.j. mustart. 1994. vegetation and conservation report for the southern overberg structure plan. university of cape town, institute for plant conservation. unpublished report. cowling, r.m. & e.t.f. witkowski. 1994. convergence and non-convergence of plant traits in climatically and edaphically matched sites in medirussell.indd 56 2006/10/15 10:58:35 pm issn 0075-6458 57 koedoe 49/2 (2006) terranean australia and south africa. australian journal of ecology 19: 220–232. cowling, r.m., b.m. campbell, p. mustart, d.j. mcdonald, m.l. jarman & e.j. moll. 1988. vegetation classification in a floristically complex area: the agulhas plain. south african journal of botany 54:290–300. cowling, r.m., r.l. pressey, a.t. lombard, p.g. desmet & a.g. ellis. 1999. from representation to persistence: requirement for a sustainable reserve system in the species rich mediterraneanclimate deserts of southern africa. diversity and distributions 5(1):51–71 de moor, j. & m.n. bruton. 1988. atlas of alien and translocated indigenous aquatic animals in southern africa. pretoria: council for scientific and industrial research. (south african national scientific programmes report 144). de villiers, a.l. 1988. micro frog. pp. 29–32. in: branch, w.r. (ed.). south african red data book—reptiles and amphibians. pretoria: council for scientific and industrial research (south african national scientific programmes; report no. 151). gaigher, i.g. 1978. the distribution, status and factors affecting the survival of indigenous freshwater fishes in the cape province. cape department of nature and environmental conservation. unpublished report. goudie, a.s. 1991. pans. progress in physical geog­ raphy 15: 221–237. harrison, a.c. 1977. the early transactions of the cape piscatorial society. part vi. piscator 100: 66–68. impson, n.d., i.r. bills, j.a. cambray & a. le roux. 1999. the primary freshwater fishes of the cape floristic region: conservation needs for a unique and highly threatened fauna. capenature internal report. 26pp. jones, m.g.w., g.d.p. van nieuwenhuizen & j.a. day. 2000. selecting priority wetlands for conservation measures. the agulhas plain as a case study. university of cape town. unpublished report. jubb, r.a. 1967. freshwater fishes of southern africa. cape town: balkema. low, a.b. & a.g. rebelo. 1996. vegetation of southern africa, lesotho, and swaziland. pretoria: department of environmental affairs and tourism. louw, r.j. 1968. survey of fish population. soetendals vlei, voël vlei. district bredasdorp. cpa department of nature and environmental conservation. unpublished report. marshall, t.r. & j.t. harmse. 1992. a review of the origin and propagation of pans. south african geographer 19: 9–21. mcveigh, s.j. 1980. inspection at soetendalsvlei, bredasdorp district. cpa department of nature and environmental conservation. unpublished report. 6pp. mustart, p., r.m. cowling & j. albertyn. 1997. southern overberg. cape town: botanical society of south africa (south african wild flower guide; no. 8). noble, r.g. & j. hemens. 1978. inland water ecosys­ tems in south africa – a review of research needs. pretoria: council for scientific and industrial research. (south african national scientific programmes; report no. 34). raimondo, j.p. & j.a. barker. 1988. eskom nuclear site investigation: southern cape region, regional study, gansbaai to agulhas. university of cape town, environmental evaluation unit. unpublished report. rebelo, a.g. 1992. red data book species in the cape floristic region: threats, priorities and target species. transactions of the royal society of south africa 48:55–86. rebelo, a.g. & w.r. siegfried. 1992. where should nature reserves be located in the cape floristic region, south africa? models for the spatial configuration of a reserve network aimed at maximizing the protection of floral diversity. conservation biology 6:243–252. ryan, p.g., l.g. underhill, j. cooper & m. waltner. 1988. waders (charadrii) and other waterbirds on the coast, adjacent wetlands and offshore islands of the southwestern cape province, south africa. bontebok 6:1–19 shaw, p.a. & d.s.g. thomas. 1997. playas, pans and salt lakes. pp. 293–317. in: thomas d.s.g. (ed.). arid zone geomorphology. london: wiley. silberhauer, m.j. & j.m. king. 1991. geographical trends in the water chemistry of wetlands in the south-western cape province, south africa. south african journal of aquatic science 17(1/2): 82–88. skelton, p.h. 1988. a taxonomic revision of the redfin minnows (pisces, cyprinidae) from southern africa. annals of the cape provincial museum 16(10): 201–307. skelton, p.h. 2001. a complete guide to the freshwa­ ter fishes of southern africa. cape town: struik. swartz, e.r. 2005. phylogeography, phylogenetics and evolution of the redfins (teleostei, cyprinidae, pseudobarbus) in southern africa. phd thesis, university of pretoria, pretoria. thwaites, r.n. & r.m. cowling. 1988. soil-vegetation relationships on the agulhas plain, south africa. catena 15: 333–345. whitfield, a.k. 1998. biology and ecology of fish­ es in southern african estuaries. grahamstown: j.l.b. smith institute of ichthyology (ichthyological monographs of the j.l.b. smith institute of ichthyology; no. 2). willis, c.k., r.m. cowling & a.t. lombard. 1996. patterns of endemism in the limestone flora of south african lowland fynbos. biodiversity & conservation 5:55–73. russell.indd 57 2006/10/15 10:58:35 pm abstract introduction journal ethical requirements for publishing primary fish research ethical considerations for field research on fishes conclusion acknowledgements references appendix 1 appendix 2 about the author(s) rhett h. bennett south african institute for aquatic biodiversity, grahamstown, south africa bruce r. ellender south african institute for aquatic biodiversity, grahamstown, south africa centre for invasion biology, south african institute for aquatic biodiversity, south africa tuuli mäkinen south african institute for aquatic biodiversity, grahamstown, south africa tshoanelo miya south african institute for aquatic biodiversity, grahamstown, south africa school of life sciences, university of kwazulu-natal, south africa paula pattrick south african institute for aquatic biodiversity, grahamstown, south africa ryan j. wasserman south african institute for aquatic biodiversity, grahamstown, south africa centre for invasion biology, south african institute for aquatic biodiversity, south africa darragh j. woodford south african institute for aquatic biodiversity, grahamstown, south africa centre for invasion biology, south african institute for aquatic biodiversity, south africa school of animal, plant and environmental sciences, university of the witwatersrand, south africa olaf l.f. weyl south african institute for aquatic biodiversity, grahamstown, south africa centre for invasion biology, south african institute for aquatic biodiversity, south africa citation bennett, r.h., ellender, b.r., mäkinen, t., miya, t., pattrick, p., wasserman, r.j. et al., 2016, ‘ethical considerations for field research on fishes’, koedoe 58(1), a1353. http://dx.doi.org/10.4102/koedoe.v58i1.1353 review article ethical considerations for field research on fishes rhett h. bennett, bruce r. ellender, tuuli mäkinen, tshoanelo miya, paula pattrick, ryan j. wasserman, darragh j. woodford, olaf l.f. weyl received: 27 aug. 2015; accepted: 08 aug. 2016; published: 02 dec. 2016 copyright: © 2016. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract collection of data from animals for research purposes can negatively impact target or by-catch species if suitable animal ethics practices are not followed. this study aimed to assess the ethical requirements of peer-reviewed scientific journals that publish primary literature on fishes, and review the ethical considerations and animal care guidelines of national and international documents on the ethical treatment of animals for research, to provide an overview of the general ethical considerations for field research on fishes. a review of 250 peer-reviewed, isi-rated journals publishing primary research on fishes revealed that nearly half (46%) had no mention of ethics, treatment of animals or ethical requirements for publication in their author guidelines or publication policies. however, 18% of the journals reviewed identify a specific set of ethical guidelines to be followed before publishing research involving animals. ethical considerations for investigators undertaking field research on fishes, common to most animal care policies, legislation and guiding documents, include adhering to relevant legislation, minimising sample sizes, reducing or mitigating pain and distress, employing the most appropriate and least invasive techniques and accurately reporting methods and findings. this information will provide potential investigators with a useful starting point for designing and conducting ethical field research. application of ethical best practices in field sampling studies will improve the welfare of study animals and the conservation of rare and endangered species. conservation implications: this article provides a list of ethical considerations for designing and conducting field research on fishes. by reviewing sampling techniques and processes that are frequently used in field research on fishes and by highlighting the potential negative impacts of these sampling techniques, this article is intended to assist researchers in planning field research to minimise impacts on fish populations. it is envisaged that this review will be a useful resource for journal editorial committees intending to introduce ethical requirements for publication and for researchers, managers, conservation practitioners and research organisations when designing field studies on fishes, applying for ethical clearance and developing institutional ethical guidelines. introduction studies on wild populations of animals are integral to scientific research (basel declaration society 2010). sampling for such research may be non-invasive (e.g. simple observation, visual census and video surveys), intrusive (e.g. external marking, fin-clipping, blood extraction, internal transmitter implantation) or lethal (e.g. voucher specimens, specimens sacrificed for biological and ecological data collection). while intrusive techniques may provide information that is not obtainable via non-invasive methods (heupel & simpfendorfer 2010), such sampling can negatively impact target or non-target (by-catch) species at individual or population levels or damage the surrounding habitat (animal behaviour editorial 2012). investigators, therefore, need to weigh up the costs and benefits in each case and select sampling methods and experimental procedures that minimise the potential disturbances to target and non-target organisms and their habitat (canadian council on animal care 2005; south african bureau of standards 2008; use of fishes in research committee 2014). in much of the world, the humane care and use of animals for scientific purposes is guided by the ethical framework of the three r’s: replacement of animals by non-animal models where possible; reduction in the number of animals used to the minimum number required for valid scientific results and refinement of procedures and animal care standards to minimise pain, suffering, distress or lasting harm (russell & burch 1959). studies employing sampling of wild populations should therefore eliminate, mitigate or minimise the potential for pain and distress and the duration of exposure to pain (ccac 2005; murray & fuller 2000; sabs 2008). accordingly, numerous guiding documents have been produced, such as the care and use of animals for scientific purposes (sabs 2008) and guide for the care and use of laboratory animals (national research council 2011), which identify important factors for consideration before and during field sampling studies. furthermore, certain peer-reviewed scientific journals (e.g. fisheries research, fisheries management and ecology and canadian journal of fisheries and aquatic sciences) prescribe ethical considerations that need to be met for research involving live animals to be published. however, many journals do not require information on the ethical treatment of study animals, while those that do, lack consistency. pain can be defined as ‘an unpleasant sensory and emotional experience associated with actual or potential tissue damage’ and nociception as ‘unconscious detection of potentially injurious stimuli by peripheral, spinal, and subcortical levels of the nervous system’ (american fisheries society 2014:20–21). recent research suggests that fishes have highly developed cognitive capabilities and the necessary physiological capacity for pain reception, nociception and suffering, resembling those found in other vertebrate groups (bshary & brown 2014; chandroo, duncan & moccia 2004). ethical considerations for research on sentient animals should therefore be applied to fish as they are to other vertebrates (metcalfe & craig 2011). however, many of the guiding documents are largely or entirely directed at the ethical use and treatment of mammalian animals, particularly captive animals (cooke et al. 2016), with few addressing the ethical use and treatment of fishes, particularly in the wild. furthermore, because of the diversity of animal taxa under research and the multitude of techniques available, a single set of guidelines is neither practical nor possible (ufrc 2014). this article aimed to assess the ethical requirements of peer-reviewed scientific journals that publish primary literature on fishes and identify the guiding documents most commonly cited in instructions of these journals to submitting authors. the article also aimed to highlight the common ethical considerations for designing and undertaking field research on fishes, with particular relevance to the commonly used techniques in ichthyological and fisheries research in southern africa. these considerations are based on international and national animal care and use policies and animal ethics guiding documents and published literature. it is envisaged that this review will be a useful resource for journal editorial committees intending to introduce ethical requirements for publication and for researchers, managers, conservation practitioners and research organisations when designing field studies on fishes, applying for ethical clearance and developing institutional ethical guidelines. journal ethical requirements for publishing primary fish research scientific journals that publish primary research on fishes were identified through a structured literature search, using the online citation indexing service web of science, maintained by thomson reuters. a text search for ‘fish*’, under the research areas of ‘fish biology’ and ‘fish ecology’ performed in february 2015 returned articles from more than 500 english-language, isi-rated journals. the instructions to authors and publication policies of the 250 journals with the most articles relating to fish research were scrutinised to identify their editorial policies relating to the publication of primary research involving the use of animals. of the 250 journals reviewed (appendix 1), 54.0% (135) have some mention of animal ethics requirements: 18.0% (45) stipulate a specific guiding document, to which submitted work must adhere for publication (category 1), 21.6% (54) require formal approval by an institutional, national or international animal ethics committee (aec) (category 2) and 14.4% (36) require only a statement within the manuscript on the welfare of study animals (category 3). the instructions to authors of the remaining 46.0% (115 journals), however, make no reference to animal ethics requirements (category 4), suggesting that the treatment of animals in almost half of these articles had not been questioned. a similar review, of 288 randomly selected english-language journals publishing original research on animals, revealed that nearly half the journals assessed had no editorial policy relating to the use of animals in research (osborne, payne & newman 2009). these two searches therefore highlight the large discrepancies in journal ethical requirements, with almost half not requiring adherence to ethical standards, while journals that do, lack consistency. the 45 journals that require authors to adhere to a specific set of ethical guidelines (category 1) together identify 15 guiding documents (table 1). the scope of these documents varies considerably, from highly prescriptive, for example, guide for the care and use of laboratory animals (nrc 2011), to providing a general tone for investigators to follow, for example, basel declaration (bds 2010). these included guidelines relating specifically to reporting, such as the arrive guidelines (animal research: reporting of in vivo experiments; kilkenny et al. 2010); national legislation, such as the united kingdom’s animals (scientific procedures) act 1986; animal use guidelines published specifically by the respective journal (i.e. animal behaviour and journal of fish biology) and guiding documents prepared specifically for the ethical treatment of fishes, such as guidelines on the care and use of fish in research, teaching and testing (ccac 2005) and guidelines for the use of fishes in research (afs 2014) (table 1). table 1: guiding documents for the ethical treatment of animals in research, stipulated by 45 isi-rated, peer-reviewed journals that publish primary research on fishes, which must be followed by authors submitting research for publication in these journals. these guiding and policy documents (for web links, see appendix 2) provide a useful starting point for investigators wishing to explore each of the common aspects for ethical consideration in more detail. while the documents range broadly in their scope, background and applicability, many share several common aspects for ethical consideration (table 2). these aspects appear repeatedly in most comprehensive animal care and use guiding documents and several countries’ legislation relating to the ethical treatment of animals in research. table 2: common aspects for consideration for the ethical treatment of animals in research, mentioned in the guiding documents identified in the author guidelines of 45 isi-rated journals that publish primary research on fishes, as well as in numerous animal care and use guiding documents and national policies. the clear and unified theme across almost all guiding documents is the need to maintain the welfare of study animals (table 2). to this end, many of these guidelines advocate adopting the principle of the three r’s – replace, reduce, refine (russell & burch 1959). most guiding documents also highlight the importance of adhering to regional, national or international legislation and gaining approval from at least a local or institutional aec, and many identify the requirements and considerations for holding (i.e. keeping in captivity), breeding and husbandry of animals. specific procedures, such as the sampling of blood or tagging of fishes, receive relatively less focus in these documents, although most documents identify the need for standardised, approved procedures and prescribe that investigators familiarise themselves with the adverse effects associated with different methods and choose the least invasive techniques to suit the aims of the study and the information required (e.g. behavioural observations, genetic studies or specimen collections) (ccac 2005; ufrc 2014). the following subsections of this review expand on these common ethical considerations, providing information (with citations to relevant literature) for researchers, managers and conservation practitioners, when undertaking sampling or field research on fishes and for journals intending to develop or specify ethical requirements for publication. particular focus is placed on the considerations for capture, handling and processing of fishes, which are relatively under-represented in many guiding documents. ethical considerations for field research on fishes based on the guiding documents identified in the previous section and numerous institutional, provincial, national and international guidelines for the care and use of animals in research, we expand on the key ethical aspects for consideration before, during and after undertaking field research on fishes, which we group into five categories: regulations, experimental design, handling and holding of study animals, experimental procedures and reporting of findings. regulations prior to any study involving live animals, investigators should familiarise themselves with the regulations and legislation pertaining to the proposed species, techniques and study area and obtain all necessary permits (murray & fuller 2000). all research activities and field sampling should follow accepted policies, standard operating procedures and international guidelines for the care and use of animals in research (ccac 2005). in south africa, animal research is governed by the veterinary and para-veterinary professions act (south african government, act no. 19 of 1982), with guidelines provided by the south african national standard (sans) 10386: the care and use of animals for scientific purposes (sabs 2008). investigators should also ensure that all methods, sample sizes, target species and study locations are approved by at least an institutional or preferably national or international aec. all persons involved in the capture, handling, holding, treatment and processing of experimental animals must be appropriately trained for their specific roles during an experimental procedure (bradford, korman & higgins 2005; murray & fuller 2000). experimental design animal welfare, sample size and the principle of the three r’s the most repeated consideration, in most guiding documents, is to maintain the welfare of the study animals and minimise or mitigate pain, suffering and distress (afs 2014; ccac 2005). this may be achieved through less invasive techniques and refining procedures in ways that reduce the intensity and durations of exposure to pain, suffering and distress (mohr 2013; nrc 2011). the number of individuals captured, manipulated or killed for the purpose of research is a fundamental consideration of any scientific study involving live animals. the appropriate sample size is contingent on the nature and objectives of the proposed research. most guiding documents therefore advocate the principle of the three r’s (russell & burch 1959). for experimental research, particularly field research in which animal subjects cannot be replaced by computer or other surrogate models, the number of study animals should be minimised as far as practicably possible, but sufficient to provide representative information (ufrc 2014). cost–benefit analysis a key principle of justifying animal experimentation is that the cost incurred to the individual or population (e.g. stress, deaths) is outweighed by the benefit of the research (e.g. advancement of knowledge) to society (knight 2011). thus, in proposed studies for collecting or manipulating fishes, a cost–benefit analysis should underlie decisions regarding sample size (animal behaviour editorial 2012). for biodiversity studies, a sample should be sufficient to capture phenotypic or genotypic variation, although this may be difficult to determine without prior knowledge of a taxon’s natural genetic variation (miyamoto et al. 2008). in experimental biology, a minimum sample size may be determined to satisfy the required statistical power (knight 2011). voucher specimen collections should sacrifice the minimum number of animals necessary for effective species confirmation, taking into account sexual dimorphism, ontogenetic change in identifiable features and similarity to related species (animal research review panel n.d.). species and sampling localities during field collection, vulnerable life-history stages or life-cycle events, such as aggregations of breeding fish, and sensitive habitats should be avoided (afs 2014). species regarded as imperilled or threatened should not be subjected to invasive sampling studies, unless deemed necessary for the specific purpose of gaining information for improved conservation (afs 2014). by-catch of non-target species should be minimised, but if captured these should be released promptly and with minimal injury (afs 2014). the number of sampling sites, depending on habitat variation, distribution range and the purpose of collection (e.g. confirming range shifts), should be minimised. investigators should also select a sampling method that has the least impact on the fishes and the local ecosystem (ccac 2005). recent debate around the conservation ethics of killing endangered species for scientific research highlights the ethical challenge of justifying the lethal sampling of fishes in field work (heupel & simpfendorfer 2010; minteer et al. 2014). for lethal sampling, particularly of threatened species, the proportion of the population affected by sampling (with implications for the viability of the sampled population) and the purpose of collecting, as it applies to the conservation management of the species, should be considered. certain fisheries journals, such as journal of fish biology and indian journal of fisheries, have explicit policies on the sizes and justification of lethal collections reported on in submitted manuscripts and will not accept manuscripts that involve the unmotivated killing or damage of iucn red list threatened or endangered species. handling, holding and release of experimental fishes captured fish should be handled in a manner that minimises pain, distress, suffering and unnecessary loss of external mucus or scales (ccac 2005). the duration of handling and the experimental procedure should be kept to a minimum, to avoid unnecessary stress and exposure time (afs 2014). investigators should assess the suitability of holding study animals prior to, during or after experimentation. if holding tanks are to be used, they must allow fish to rest comfortably, minimise risk of escape or injury, be adequately aerated, maintain constant temperature and minimise the risk of disease transmission (barker et al. 2002). holding tanks should be disinfected between uses and holding areas must be safe, quiet and hygienic (barker et al. 2002). while post-experimentation captivity may allow recovery monitoring, this may be undesirable for species vulnerable to stress from captivity, and holding of animals in field research, particularly large-bodied animals, may be impractical (jepsen et al. 2002). where appropriate, fish should be released as soon as possible after completion of processing (e.g. measuring, tagging or collection of tissue). all fish having undergone an experimental procedure must be in good health when released and able to return to their natural environment with ‘normal’ physiological and behavioural functioning (bradford et al. 2005; ccac 2005). all necessary steps must be taken to prevent predation and injury upon release and the introduction of pathogens or harmful chemicals into the environment (sabs 2008). experimental procedures sampling methods for fish larvae high levels of natural mortalities and rapid recoveries from population reductions are characteristic events in the early life histories of many fish species (ufrc 2014). it is therefore generally regarded by most ethical standards that the sampling of fish larvae that have not developed beyond exclusive reliance on their own yolk nutrients does not require ethical clearance (canadian council on animal care 2007). however, according to the sabs (2008), live, sentient non-human vertebrates, including eggs, foetuses and embryos of fish, are to be cared for and used in ways that are judged to be scientifically, technically and humanely appropriate. furthermore, investigations involving the early life stages of fishes typically require the sacrifice of large numbers of study organisms. therefore, a precautionary approach should be followed, in which sample sizes and by-catch are minimised, particularly when threatened species constitute the target or possible by-catch species (ccac 2005). for fish larvae, killing by immersion in ice or a fixative such as ethanol or formalin is generally acceptable; however, this should be preceded by an overdose of an appropriate anaesthetic (craig 2006). any proposed non-use of anaesthesia on the grounds of compromising the aim of the study should be fully justified a priori (metcalfe & craig 2011). when fish larvae become independently feeding forms, ethical authorisation must be obtained (ccac 2007). non-invasive fish sampling techniques visual sampling, such as underwater visual census (by snorkelling or scuba) and underwater video surveys, is common in marine, estuarine and freshwater environments (ellender et al. 2012; willis, millar & babcock 2000) and provides a valuable tool for collecting qualitative and quantitative fishery-independent data on fish density, diversity, community structure and behaviour (brock 1982). underwater videography is increasingly used for assessing and monitoring fish communities, as digital video provides standardised methodology and permanent records (langlois et al. 2010). the non-destructive nature of these methods makes them suitable for sensitive habitats and protected areas (mallet & pelletier 2014), and in most cases, ethical justification for their use is not necessary. in some instances, however, divers conducting visual or video observations or the deployment of remote equipment may cause habitat damage. investigators should thus attempt to minimise any potential impact on the habitat. capture sampling techniques numerous sampling techniques involve the actual capture (such as trapping, netting or hooking) of target fishes and are thus inherently invasive. investigators therefore need to understand the uses, limitations and threats of these methods, in order that the most appropriate method be employed, to provide the most satisfactory result, while minimising pain, suffering and distress (eu 2010). netting: there are a variety of netting methods for the capture of fishes, some passive and some active. gill nets are passive nets, hung vertically in the water column. the gills or fins of fish swimming into the net become entangled in the mesh, preventing their escape. mesh size is an important consideration, as larger mesh sizes usually result in the capture of larger fish, while providing some form of exclusion for smaller species or individuals. the use of gill nets for sampling is generally accepted, although gill nets can result in cuts on the body from the filament of the net or the drowning of obligate air-breathing fishes, amphibians, reptiles and mammals (ellender et al. 2016). fyke nets are also passive nets, comprising a vertical wall of net that guides fishes into an enclosed cod-end or bag at either end. they are generally considered non-destructive, as fish are free to swim in the cod-end once captured, allowing them to be released relatively unharmed. however, fyke net placement is an important consideration, as nets set in areas of low oxygen concentration can result in elevated fish mortality. furthermore, fyke nets are not species selective and mortality of by-catch, particularly drowning of obligate air-breathing animals, can occur (larocque, cooke & blouin-demers 2012). mortality can also result from predatory fishes, frogs or crabs that are caught in the cod-end or from external sources, such as otters attacking fishes through the net (ellender et al. 2016). given the associated predation risk to captured fishes, fyke nets may be unsuitable in certain environments, particularly if they pose a high capture threat to endangered species. simple gear modifications, such as creating air spaces using floats to allow captured animals to breathe (larocque et al. 2012) or selective mesh placed over the entrance to exclude mammals or reptiles, can however reduce mortality and by-catch. reducing soak time or checking nets at shorter intervals enables by-catch and target species to be processed and released more rapidly. owing to the potential injuries, by-catch and mortalities associated with gill and fyke netting, motivation for their use is warranted and identification of potential by-catch species and an assessment of their conservation status should form an integral part of the planning phase of any project using these gears. active net gears include, inter alia, seine nets and various forms of trawl nets. seine nets are vertical nets, with a floating top line and weighted bottom line, actively hauled to encircle a set area of water and pull the fish within that area to shore. they are a preferred gear for sampling fish communities in shallow littoral environments such as beaches, estuaries and artificial impoundments (lapointe, corkum & mandrak 2006). a cod-end at the centre of the net acts as a corral point for fish as they are brought to shore. seine netting is generally regarded as non-destructive, although mortality of small-bodied species and post-release mortality of discarded by-catch species have been recorded (kennelly & gray 2000). fish mortality can be minimised by handling fish within the submerged cod-end before bringing the net onshore. in some cases, seine netting can damage submerged macrophyte habitat in haul regions, although the impact is usually minor (bayley & herendeen 2000). trawling refers to nets towed from a boat, which usually sample hyper-benthic environments. the use of dragged or towed gears, which may damage the habitat, benthic environment or animals sampled (feyrer et al. 2013), should be avoided or minimised (ufrc 2014). alternative methods should also be sought where possible; for example, feyrer et al. (2013) investigated the use of attaching a camera to the open cod-end of a trawl net, allowing fish to be recorded but to pass through the net unharmed, greatly reducing mortality. electrofishing: electrofishing is one of the most effective and commonly employed sampling methods in shallow freshwater environments (snyder 2003). while generally considered non-destructive, it can have negative impacts (snyder 2003). in extreme cases, severe spinal injuries, internal haemorrhages, bleeding at the gills, physiological stress, asphyxiation and harmful effects on embryos have been recorded (snyder 2003). it is therefore recommended that researchers familiarise themselves with, and apply, recommendations for international best practises regarding electrofishing (e.g. beaumont et al. 2002; goodchild 1991). current strengths and settings and the electrofishing gear itself should be suited to the target species and conditions in the study area, such as conductivity, water depth and survey area (bohlin et al. 1989). when sampling threatened or endangered fishes, electrofishing should be used with caution and alternative methods employed where possible (ellender et al. 2012). if multiple electrofishing passes are intended, all fish captured should immediately be removed from the net and placed in aerated buckets, filled with ambient water, to avoid prolonged exposure. bohlin et al. (1989) and snyder (2003) provide comprehensive reviews on standardising electrofishing protocols and harmful effects of electrofishing. hook-and-line fishing: hook-and-line fishing is commonly used in a range of research applications, such as quantifying abundance, assessing community structure and for tagging studies (bennett & attwood 1993; dunlop, mann & van der elst 2013; willis et al. 2000). catch-and-release is often employed, whereby fishes are released to the capture environment after undergoing procedures such as measuring or tagging, with minimal impact to the fishes and environment (cooke et al. 2013a). however, excessive fight times or post-capture air exposure and high water temperatures have been shown to induce physiological stress responses in hook-and-line caught fishes, with implications for their successful release (cooke & suski 2004). in addition, fish captured by hook-and-line fishing may be subjected to stress as a result of barotrauma injuries if rapidly brought to the surface (keniry et al. 1996), or damage to the gills or viscera through hook ingestion (domeier, dewar & nasby-lucas 2003). the potential impacts of hook-and-line fishing should thus be thoroughly considered before using this method to capture endangered species or in excessive water depths where barotrauma is likely. relevant mitigation measures, such as choice of hook design and size, optimal fishing times and minimised handling times need to be considered (see cooke & suski 2004). the use of circle hooks has been shown to decrease the incidence of gutor deep-hooking (cooke & suski 2004) and post-release mortality (prince et al. 2007) and the use of barbless hooks can reduce injury and facilitate hook removal (schaeffer & hoffman 2002). brownscombe et al. (2016) provide guidelines for best practice in catch-and-release recreational fisheries. ichthyocides ichthyocides (poisons) and anaesthetics have been used extensively to determine fish species composition, particularly in freshwater and shallow reef environments (ackerman & bellwood 2000; willis 2001). rotenone, for example, is a commonly used ichthyocide, highly effective at killing most fish species (afs 2014; willis 2001). although such techniques are effective for assessing cryptic species and provide greater species and family counts than non-destructive methods (willis 2001), they can cause excessive mortality for sensitive species, are non-selective, vary widely in effectiveness, provide inefficient sampling of highly mobile species and cannot be used in sensitive areas or where sampling is to be repeated (brock 1982). furthermore, recent studies have highlighted that many non-target organisms are susceptible to ichthyocide concentrations used for fishes (e.g. dalu et al. 2016). careful consideration, especially of collateral impacts on non-target biota, is therefore necessary, prior to using ichthyocides. clearwater, hickey and martin (2008) discuss several ichthyocides for pest control; however, a comprehensive review of ichthyocides and their use in fish research is lacking. anaesthesia and analgesia sedatives or anaesthetics are commonly prescribed and used during prolonged or invasive procedures, to immobilise fish and reduce pain and physiological stress related to capture, handling and processing, through depression of the nervous system (ccac 2005; ufrc 2014). minor procedures usually require mild sedation, whereas intrusive (major) procedures, such as laparotomy, usually require deep anaesthesia (thorsteinsson 2002). where anaesthetics or sedatives are used, only approved drugs should be used and administered at a concentration and duration appropriate to the size and species of fish and in accordance with established guidelines and relevant legislation (afs 2014; ufrc 2014). the sans 10386 (sabs 2008) recommends four anaesthetics for use on fishes in south africa, including tricaine methanesulfonate (ms-222), benzocaine (and benzocaine hydrochloride), metomidate and ketamine hydrochloride. however, none of these is registered in australia, new zealand, the united states, canada or europe (javahery & moradlu 2012). other anaesthetics common in fish research include 2-phenoxyethanol, clove oil, aqui-stm (active ingredient isoeugenol) and quinaldine sulphate (javahery & moradlu 2012; wagner & cooke 2005). however, many of these are regarded as hazardous (javahery & moradlu 2012), particularly if fish are released and subsequently consumed by humans (thorsteinsson 2002). of these anaesthetics, only ms-222 is approved by the united states food and drug administration (fda) for use on animals in the united states (afs 2014), but has a 21-day withdrawal period before fish can be released into the wild, negating its use in field studies. eugenol (a derivative of clove oil) and carbon dioxide are the only immediate-release drugs approved by the fda (afs 2014) – yet neither is recommended for use in south africa (sabs 2008), and carbon dioxide allows considerable trauma at shallow sedation and is not suitable for heavy sedation required in invasive procedures (javahery & moradlu 2012). therefore, identifying an appropriate drug and concentration, particularly for fish intended for immediate release, is often a challenging task and should be based on thorough research of contemporary literature and legislation prior to any implementation (see thorsteinsson 2002 for comprehensive review of anaesthetics). anaesthetics can induce prolonged negative metabolic effects and stress responses in fishes (ccac 2005), as well as hypoxia from reduced respiration and vascular activity (thorsteinsson 2002). in certain cases, this may lead to post-release mortality or increased predation risk (cooke et al. 2005). numerous studies on fishes, involving handling and even invasive procedures such as laparotomy, have deemed the use of anaesthesia inappropriate (wagner & cooke 2005). for large elasmobranchs, anaesthesia would require specialised lifting equipment, considerable risk to the animals and excessive quantities of chemicals; therefore, such studies usually rely on tonic immobility (e.g. holland et al. 1999). therefore, the use of anaesthetics should be evaluated prior to any study (thorsteinsson 2002), particularly when fish are planned for release into the wild and in some cases may not be appropriate at all (cooke et al. 2013b, 2016). collecting tissue and blood samples tissue collection: a variety of methods have been used to successfully collect tissue or blood samples from anaesthetised fish without significant effects on their survival. non-lethal tissue sampling includes fin-clipping, excision of scales and gill, muscle or skin biopsies (mccormick 1993; ufrc 2014). fin-clipping is common for genetic sampling and its deleterious effects have been extensively studied. many studies reported no negative effects and complete regeneration of the affected fins in weeks (woodall, koldewey & shaw 2011), while others reported some decrease in survival or growth of fin-clipped fish (o’grady 1984). the appropriate site to sample is species specific. for example, partial clipping of the dorsal fin was successfully used for genetic sampling of seahorses with no significant effect on mortality (woodall et al. 2011). blood collection: fish blood is collected for various reasons, such as haematology and clinical chemistry tests (satheeshkumar et al. 2012), parasitological investigations (shahi et al. 2013) and investigations on adaptations to subzero temperatures (miya et al. 2014). such studies have demonstrated that blood samples can be successfully obtained, even from fish less than 100 g in mass, without compromising their survival. plastic syringes with a small volume of anticoagulant such as sodium or ammonium heparin or sodium citrate can be used to avoid the rapid clotting that can occur with the use of glass syringes (stoskopf 1993). the size of the syringe and diameter of needle should be appropriate to the size of fish. the preferred site for blood sampling from anaesthetised fish is the caudal vein running beneath the vertebrae, using a lateral or ventral approach. study objectives will determine the required blood volume and concentration of anticoagulant. other methods used to collect blood from fishes include cardiac puncture and caudal bleeding (ufrc 2014). these methods, however, can compromise the survival of the fish. needles are most effective when new and sharp and should be replaced or sterilised after each sample to avoid contamination and unnecessary injuries to the fish. marking and tagging of fishes marking (using dyes or fin-clipping) and tagging of fishes with external or internal tags has been used extensively to estimate population sizes, discriminate stocks, identify migration rates and movement patterns and assess fish growth and survival (mcfarlane, wydoski & prince 1990). numerous marking and tagging methods are used, although most are intrusive to some degree (thorsteinsson 2002). the capture, handling and marking/tagging process can affect the behaviour, physiology, swimming capacity, predator avoidance, feeding, growth rate and ultimate survival of tagged organisms (naef-daenzer et al. 2005; thorsteinsson 2002). studies should therefore aim to minimise these impacts, for the welfare of the study animal, to ensure that the process provides scientifically valuable data, and to maximise the value from every individual that has been subjected to the stress of capture, handling and processing (cooke et al. 2013b). marking: marking techniques for fishes, such as external dyes, fin-clipping, hot branding (e.g. lasers) and freeze branding (e.g. liquid nitrogen), do not breach the skin or musculature and are thus generally considered to have minimal impact on fishes (murray & fuller 2000). however, such marks usually exhibit short-term retention, and brightly coloured external marks or dyes (and any external part of a tag) should be avoided, as these can affect social structure or interactions of the fish or reduce its predator avoidance or hunting capabilities (guy, blankenship & nielsen 1996; murray & fuller 2000). chemical tags are another form of marking and may be natural (i.e. accumulation from the local environment) or induced, through immersion, injection or ingestion of chemicals, which become incorporated into tissue or calcified structures (nielsen 1992). chemical tagging is long lasting and allows large numbers of individuals to be tagged and does not necessarily require anaesthetic or even handling (guy et al. 1996), but often requires sacrifice of the animal at a later stage. general tagging considerations: to minimise the effects of tags on fish and maximise data collection and tag retention, tag size and weight should be appropriate for the size, weight and species of fish (cooke et al. 2011; stasko & pincock 1977). the tag placement and attachment method must also be appropriate for the study species (naef-daenzer et al. 2005; sabs 2008) and have minimal influence on the posture, buoyancy, locomotion (stasko & pincock 1977), behaviour, growth and survival of the fish (bradford et al. 2009). where necessary, the suitability of a chosen tag type, size and attachment method should be assessed prior to tagging studies on a new species (ccac 2005; murray & fuller 2000). external tag attachment: external attachment is used for simple plastic or wire tags and electronic transmitters and requires that an external or trailing part of the tag be anchored somehow within the musculature or skeletal structure of the fish. a common type of external tagging, particularly in southern africa, is dart tagging (maggs & cowley 2016). dart tags are inserted using a sharp, sterile tag applicator, usually at the base of the dorsal fin and anchored behind the pterygiophores (attwood 1998). electronic tags can also be attached externally in a similar manner or with wires or nylon cords inserted through the dorsal musculature of the fish and secured with a plastic or metal plate (liedtke & rub 2012; thorstad et al. 2013). sharks and large pelagic fishes are often tagged in situ in the water, by means of a dart tag or electronic tag attached to the end of a sharpened pole. for pelagic sharks, tags (particularly satellite tags) are also commonly attached by means of plastic or metal bolts through the dorsal fin (thorsteinsson 2002). external tagging is simple and does not necessarily require animals to be anaesthetised, or even brought out of the water (thorsteinsson 2002). external tags, however, can have negative impacts, as they may increase drag and impede normal swimming ability and reduce growth or survival (murray & fuller 2000). the attachment mechanisms breach the skin and musculature and could lead to infection, tissue damage or necrosis, reduced growth or reduced reproductive capacity (thorsteinsson 2002). external tags, particularly larger volume electronic tags, should thus be shaped and attached in such a way that minimises drag, entanglement in aquatic vegetation, irritation and constriction (ccac 2005; murray & fuller 2000). owing to the disadvantages of external tag attachment, a number of internal tag implantation techniques have been used. gastric insertion: gastric insertion of devices involves voluntary ingestion of the device embedded in bait or the forcing of the device down the pharynx, past the cardiac sphincter and into the stomach by means of a glass or plastic rod (liedtke & rub 2012; thorsteinsson 2002). the process is less invasive than surgical implantation (winger & walsh 2001), and the internal device overcomes many of the problems associated with external tags, such as entanglement, drag or wound development (stasko & pincock 1977). however, gastric insertion can only be used when fish are not feeding and there are several drawbacks to the technique. there is a risk of tag loss through regurgitation, the presence of the device may affect the ability of the fish to feed and injury may be caused to the oesophagus or stomach (thorstad et al. 2013; winter 1996). furthermore, gastric insertion cannot be used on fishes with a food-crushing pharyngeal apparatus, and the process does not allow for long-term tag retention (stasko & pincock 1977). surgical implantation of tags or devices: laparotomy (the surgical implantation of a device through an incision in the body wall) is becoming recognised as a superior technique to external attachment and gastric insertion, particularly for long-term tag retention, reduced long-term physiological stress and data that may be more representative of normal behaviour (cooke et al. 2011; thorstad et al. 2013). the technique is considered to have little or no methodological bias in the long-term, and the internal placement of the tags removes the potential entanglement and hydrodynamic interference that may result from external tags (jepsen et al. 2002; thorstad et al. 2013). however, the recovery time after surgical implantation may be greater than that for external attachment or gastric insertion (thorsteinsson 2002). furthermore, this process requires suitably trained or qualified personnel and is governed in certain states by formal legislation. in south africa, surgical procedures are regulated under the veterinary and para-veterinary professions act (sa government, act no. 19 of 1982). for all tagging procedures, aseptic technique should be maintained, although this is difficult in most field settings. antibiotic powder (topical) or liquid (topical or by injection) can be applied post-tagging to prevent secondary infection. before releasing tagged fish back into their natural environment, they should be monitored until having recovered from the effects of anaesthesia and surgery. euthanasia where the sacrifice of study fish cannot be avoided (e.g. voucher specimen collection or physiological studies), humane euthanasia techniques should be used (sabs 2008). animals may be euthanised by chemical or physical methods, and the selected methods should be predictable, minimise pain and stress, produce rapid loss of consciousness and be compatible with the scientific aims (american veterinary medical association 2013; sabs 2008). fishes to be preserved should be euthanised prior to immersion in formalin or other preservatives (ccac 2005). where feasible, euthanasia should consist of a two-step process with initial anaesthesia until loss of equilibrium, followed by a physical or chemical method to cause brain death (ccac 2005). rapid cooling, followed by prolonged exposure in ice-cooled water, is recommended for small-bodied fishes (avma 2013). overdosing with anaesthetics, such as ms-222, is an accepted method (neiffer & stamper 2009), whereas the use of carbon dioxide, suffocation (removal from water) and decapitation alone are generally not. the avma (2013) provides comprehensive guidelines on euthanasia of animals. reporting equally important as study design, animal welfare and appropriate methodologies in animal research, are the accurate, comprehensive reporting and communication of procedures, protocols and results, to facilitate knowledge transfer (filipecki et al. 2011). to this end, the arrive guidelines were developed to ensure that all relevant information is included in publications, to maximise the value of publications and render them useful for scientific advancement or policy making (kilkenny et al. 2010). such information may include (inter alia) numbers, demographics and sources of study subjects; sampling localities and times; detailed processing methods; reagent concentrations; holding conditions; statistical tests; positive and negative outcomes; limitations; new discoveries; and applicability of results beyond the specific study (kilkenny et al. 2010). conclusion the review of journal ethical requirements revealed considerable variability in the level of ethical reporting for research involving animals to be accepted for publication. while publishers, research institutions and government agencies may vary in their requirements for demonstrated ethical behaviour, such inconsistency should not deter investigators from taking individual responsibility for their ethical duties in sampling or manipulating fishes in the field. the principles of animal welfare and biodiversity conservation should always guide the development of research project methods, regardless of whether they are required by the journal in which an investigator wishes to publish. to aid this principle, this review highlights both the common guiding documents on the ethical treatment of animals (particularly fishes) and common, important ethical considerations for managers, researchers, technicians and students, as they apply to commonly used methods for field-based research on fishes. primary ethical considerations are that (1) all field, capture, sampling and processing activities must adhere to relevant legislation and should be approved by at least an institutional aec, (2) the welfare of the study animals should be of primary concern and all efforts should be made to prevent or reduce pain, suffering and distress, (3) sample sizes should be kept to the minimum, but sufficient to achieve the objectives of the study, (4) the conservation status of the target organism should be considered, (5) destructive or invasive gears and processes should be applied with caution and (6) standard and accepted procedures should be followed. the review will provide a useful resource for journal editorial committees developing ethical guidelines for publication and for potential investigators designing and conducting field research on fishes. application of these ethical requirements in field sampling studies will improve fish welfare and the conservation of fishes, especially rare and critically endangered species. acknowledgements the authors thank mrs sally schramm, saiab senior librarian, for assisting with the journals search protocol. two anonymous reviewers are also thanked for considerable constructive input, and taryn murray is thanked for commenting on and proofreading the final version. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions r.h.b. was the lead author and o.l.f.w. was the project leader. r.h.b., b.r.e., p.p., t. mäkinen, t. miya, r.j.w., d.j.w. and o.l.f.w. contributed to conceptualisation, analyses, writing and editing the manuscript. references ackerman, j.l. & bellwood, d.r., 2000, ‘reef fish assemblages: a re-evaluation using enclosed rotenone stations’, marine ecology progress series 206, 227–237. http://dx.doi.org/10.3354/meps206227 american fisheries society, 2014, ‘guidelines for the use of fishes in research’, american fisheries society, bethesda, md. american veterinary medical association, 2013, ‘avma guidelines for the euthanasia of animals: 2013 edition’, american veterinary medical association, schaumberg. animal research review panel of the new south wales provincial government, n.d., ‘collection of voucher specimens’, viewed 25 august 2015, from http://www.animalethics.org.au/policies-and-guidelines/wildlife-research/voucher-specimens attwood, c.g., 1998, ‘tagging the correct way’, the fishing journal 1, 23–26. barker, d., allan, g.l., rowland, s.j. & pickles, j.m., 2002, a guide to acceptable procedures and practises for aquaculture and fisheries research, nsw fisheries animal care and ethics committee, orange. basel declaration society, 2010, basel declaration: a call for more trust, transparency and communication on animal research, basel declaration committee, basel. bayley, p.b. & herendeen, r.a., 2000, ‘the efficiency of a seine net’, transactions of the american fisheries society 129, 901–923. http://dx.doi.org/10.1577/1548-8659(2000)129<0901:teoasn>2.3.co;2 beaumont, w.r.c., taylor, a.a.a., lee, m.j. & welton, j.s., 2002, guidelines for electric fishing best practice, r&d technical report w2-054/tr, environment agency, bristol. bennett, b.a. & attwood, c.g., 1993, ‘shore-angling catches in the de hoop nature reserve, south africa, and further evidence for the protected value of marine reserves’, south african journal of marine science 13, 213–222. http://dx.doi.org/10.2989/025776193784287202 bohlin, t., hamrin, s., heggberget, t.g., rasmussen, g. & saltveit, s.j., 1989, ‘electrofishing – theory and practice with special emphasis on salmonids’, hydrobiologia 173, 9–43. http://dx.doi.org/10.1007/bf00008596 bradford, m.j., korman, j. & higgins, p.s., 2005, ‘using confidence intervals to estimate the response of salmon populations (oncorhynchus spp.) to experimental habitat alterations’, canadian journal of fisheries and aquatic science 62, 2716–2726. http://dx.doi.org/10.1139/f05-179 bradford, r.w., hobday, a.j., evans, k. & lansdell, m., 2009, cmar code of practice for tagging marine animals, csiro marine and atmospheric research paper 028, csiro, hobart. brock, r.e., 1982, ‘a critique of the visual census method for assessing coral reef populations’, bulletin of marine science 32(1), 269–276. brownscombe, j.w., danylchuk, a.j., chapman, j.m., gutowsky, l.f.g. & cooke, s.j., 2016, ‘best practices for catch-and-release recreational fisheries – angling tools and tactics’, fisheries science. http://dx.doi.org/10.1016/j.fishres.2016.04.018 bshary, r. & brown, c., 2014, ‘fish cognition’, current biology 24, r947–r950. http://dx.doi.org/10.1016/j.cub.2014.08.043 canadian council on animal care, 2005, guidelines on: the care and use of fish in research, teaching and testing, canadian council on animal care, ottawa. canadian council on animal care, 2007, guidelines on: procurement of animals used in science, canadian council on animal care, ottawa. chandroo, k.p., duncan, i.j.h. & moccia, r.d., 2004, ‘can fish suffer?: perspectives on sentience, pain, fear and stress’, applied animal behaviour science 86, 225–250. http://dx.doi.org/10.1016/j.applanim.2004.02.004 clearwater, s.j., hickey, c.w. & martin, m.l., 2008, ‘overview of potential piscicides and molluscicides for controlling aquatic pest species in new zealand’, science for conservation 283, 5–74. cooke, s.j., crossin, g.t., patterson, d.a., english, k.k., hinch, s.g., young, j.l. et al., 2005, ‘coupling non-invasive physiological assessments with telemetry to understand inter-individual variation in behaviour and survivorship of sockeye salmon: development and validation of a technique’, journal of fish biology 67, 1342–1358. http://dx.doi.org/10.1111/j.1095-8649.2005.00830.x cooke, s.j., donaldson, m.r., o’connor, c.m., raby, g.d., arlinghaus, r., danylchuk, a.j. et al., 2013a, ‘the physiological consequences of catch-and-release angling: perspectives on experimental design, interpretation, extrapolation and relevance to stakeholders’, fisheries management and ecology 20, 268–287. http://dx.doi.org/10.1111/j.1365-2400.2012.00867.x cooke, s.j., nguyen, v.m., murchie, k.j., thiem, j.d., donaldson, m.r., hinch, s.g. et al., 2013b, ‘to tag or not to tag: animal welfare, conservation, and stakeholder considerations in fish tracking studies that use electronic tags’, journal of international wildlife law and policy 16(4), 352–374. http://dx.doi.org/10.1080/13880292.2013.805075 cooke s.j. & suski, c.d., 2004, ‘are circle hooks an effective tool for conserving marine and freshwater recreational catch-and-release fisheries?’, aquatic conservation: marine and freshwater ecosystems 14, 299–326. http://dx.doi.org/10.1002/aqc.614 cooke, s.j., wilson, a.d.m., elvidge, c.k., lennoz, r.j., jepsen, n., colotelo, a.h. et al., 2016, ‘ten practical realities for institutional animal care and use committees when evaluating protocols dealing with fish in the field’, reviews in fish biology and fisheries 26, 123–133. http://dx.doi.org/10.1007/s11160-015-9413-y cooke, s.j., woodley, c.m., eppard, m.b., brown, r.s. & nielsen, j.l., 2011, ‘advancing the surgical implantation of electronic tags in fish: a gap analysis and research agenda based on a review of trends in intracoelomic tagging effects studies’, reviews in fish biology and fisheries 21, 127–151. http://dx.doi.org/10.1007/s11160-010-9193-3 council for international organization of medical sciences/international council for laboratory animal science, 2012, international guiding principles for biomedical research involving animals, council for international organization of medical sciences/international council for laboratory animal science, geneva. council of europe, 1986, european convention for the protection of vertebrate animals used for experimental and other scientific purposes, european treaty series 123, council of europe, strasbourg. craig, j.f., 2006, ‘ethical justification for the use and treatment of fishes in research’, journal of fish biology 68, 1–2. http://dx.doi.org/10.1111/j.0022-1112.2006.01035.x dalu, t., wasserman, r.j., jordaan, m., froneman, w.p. & weyl, o.l., 2015, ‘an assessment of the effect of rotenone on selected non-target aquatic fauna’, plos one 10, e0142140, viewed 24 june 2016, from http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0142140, http://dx.doi.org/10.1371/journal.pone.0142140 domeier, m.l., dewar, h. & nasby-lucas, n., 2003, ‘mortality rate of striped marlin (tetrapturus audax) caught with recreational tackle’, marine and freshwater research 54, 435–445. http://dx.doi.org/10.1071/mf01270 dunlop, s.w., mann, b.q. & van der elst, r.p., 2013, ‘a review of the oceanographic research institute’s cooperative fish tagging project: 27 years down the line’, african journal of marine science 35, 209–221. http://dx.doi.org/10.2989/1814232x.2013.769909 editorial, 2011, ‘ethical justification for the use and treatment of fishes in research: an update’, journal of fish biology 78, 393–394. editorial, 2012, ‘guidelines for the treatment of animals in behavioural research and teaching’, animal behaviour 83, 301–309. ellender, b.r., becker, a., weyl, o.l.f. & swartz, e.r., 2012, ‘underwater video analysis as a non-destructive alternative to electrofishing for sampling imperilled headwater stream fishes’, aquatic conservation: marine and freshwater ecosystems 22(1), 58–65. http://dx.doi.org/10.1002/aqc.1236 ellender, b.r., wasserman, r.j., ndaleni, p.m., mofu, l., wu, y. & weyl, o.l.f., 2016, ‘reporting by-catch and mortalities in targeted biological field surveys on fishes’, aquatic conservation: marine and freshwater ecosystems 26(3), 596–601. http://dx.doi.org/10.1002/aqc.2613 european union, 2010, ‘directive 2010/63/eu of the european parliament and of the council of 22 september 2010 on the protection of animals used for scientific purposes’, official journal of the european union 276, 33–79. feyrer, f., portz, d., odum, d., newman, k.b., sommer, t., contreras, d. et al., 2013, ‘smeltcam: underwater video codend for trawled nets with an application to the distribution of the imperiled delta smelt’, plos one 8, e67829. http://dx.doi.org/10.1371/journal.pone.0067829 filipecki, a.t.p., machado, c.j.s., valle, s. & de oliveira teixeira, m., 2011, ‘the brazilian legal framework on the scientific use of animals’, ilar e-journal 52, e8–e15, viewed 06 june 2016, from http://ilarjournal.oxfordjournals.org/content/52/1/e8.full.pdf+html goodchild, g.a., 1991, code of practice and guidelines for safety with electric fishing, eifac occasional paper 24, food and agriculture organisation, rome. guy, c.s., blankenship, h.l. & nielsen, l.a., 1996, ‘tagging and marking’, in b.r. murphy & d.w. willis (eds.), fisheries techniques, pp 353–383, american fisheries society, bethesda, md. heupel, m.r. & simpfendorfer, c., 2010, ‘science or slaughter: need for lethal sampling of sharks’, the journal of the society for conservation biology 24, 1212–1218. http://dx.doi.org/10.1111/j.1523-1739.2010.01491.x holland, k.n., wetherbee, b.m., lowe, c.g. & meyer, c.g., 1999, ‘movements of tiger sharks (galeocerdo cuvier) in coastal hawaiian waters’, marine biology 134, 665–673. http://dx.doi.org/10.1007/s002270050582 javahery, s. & moradlu, a.h., 2012, ‘aqui-s, a new anesthetic for use in fish propagation’, global veterinaria 9(2), 205–210. jepsen, n., koed, a., thorstad, e.b. & baras, e., 2002, ‘surgical implantation of telemetry transmitters in fish: how much have we learned?’, hydrobiologia 483, 239–248. http://dx.doi.org/10.1023/a:1021356302311 keniry, j., brofka, w.a., horns, w.h. & marsden, j.e., 1996, ‘effects of decompression and puncturing the gas bladder on survival of tagged yellow perch’, north american journal of fisheries management 16(1), 201–206. http://dx.doi.org/10.1577/1548-8675(1996)016<0201:eodapt>2.3.co;2 kennelly, s.j. & gray, c.a., 2000, ‘reducing the mortality of discarded undersize sand whiting sillago ciliata in an estuarine seine fishery’, marine and freshwater research 51(8), 749–753. http://dx.doi.org/10.1071/mf00014 kilkenny, c., browne, w.j., cuthill, i.c., emerson, m. & altman, d.g., 2010, ‘improving bioscience research reporting: the arrive guidelines for reporting animal research’, plos biology 8(6), e1000412, viewed 06 june 2016, from https://www.nc3rs.org.uk/arrive-guidelines, http://dx.doi.org/10.1371/journal.pbio.1000412 knight, a., 2011, ‘weighing the costs and benefits of animal experiments’, in eighth world congress on alternatives and animal use in the life sciences, alternatives to animal experimentation, proceedings, montreal, canada, 21–25 august, 2011, pp. 289–294. langlois, t.j., harvey, e.s., fitzpatrick, b., meeuwig, j.j., shedrawi, g. & watson, d.l., 2010, ‘cost-efficient sampling of fish assemblages: comparison of baited video stations and diver video transects’, aquatic biology 9, 155–168. http://dx.doi.org/10.3354/ab00235 lapointe, n.w.r., corkum, l.d. & mandrak, n.e., 2006, ‘a comparison of methods for sampling fish diversity in shallow offshore waters of large rivers’, north american journal of fisheries management 26, 503–513. http://dx.doi.org/10.1577/m05-091.1 larocque, s.m., cooke, s.j. & blouin-demers, g., 2012, ‘a breath of fresh air: avoiding anoxia and mortality of freshwater turtles in fyke nets by the use of floats’, aquatic conservation: marine and freshwater ecosystems 22(2), 198–205. http://dx.doi.org/10.1002/aqc.1247 liedtke, t.l. & rub, m.w., 2012, ‘techniques for telemetry transmitter attachment and evaluation of transmitter effects on fish performance’, in n.s. adams, j.w. beeman & j.h. eiler (eds.), telemetry techniques: a user guide for fisheries research, pp. 45–87, american fisheries society, bethesda, md. maggs, j.q. & cowley, p.d., 2016, ‘nine decades of fish movement research in southern africa: a synthesis of research and findings from 1928 to 2014’, reviews in fish biology and fisheries 26, 287–302. http://dx.doi.org/10.1007/s11160-016-9425-2 mallet, d. & pelletier, d., 2014, ‘underwater video techniques for observing coastal marine biodiversity: a review of sixty years of publications (1952–2012)’, fisheries research 154, 44–62. http://dx.doi.org/10.1016/j.fishres.2014.01.019 mccormick, s.b., 1993, ‘methods for nonlethal gill biopsy and measurement of na*, k* -atpase activity’, canadian journal of fisheries and aquatic sciences 50, 656–658. http://dx.doi.org/10.1139/f93-075 mcfarlane, g.a., wydoski, r.s. & prince, e.o., 1990, ‘a historical review of the development of external tags and marks’, american fisheries science symposium 7, 9–29. metcalfe, j.d. & craig, j.f., 2011, ‘ethical justification for the use and treatment of fishes in research: an update’, journal of fish biology 78(2), 393–394. http://dx.doi.org/10.1111/j.1095-8649.2010.02900.x minteer, b.a, collins, j.p., love, k.e. & puschendorf, r., 2014, ‘avoiding (re)extinction’, science 344, 260–261. http://dx.doi.org/10.1126/science.1250953 miya, t., gon, o., mwale, m. & cheng, c.-h.c., 2014, ‘the effect of habitat temperature on serum antifreeze glycoprotein (afgp) activity in notothenia rossii (pisces: nototheniidae) in the southern ocean’, polar biology 37, 367–373. http://dx.doi.org/10.1007/s00300-013-1437-y miyamoto, n., fernández-manjarrés, j.f., morand-prieur, m.-e., bertolino, p. & frascaria-lacoste, n., 2008, ‘what sampling is needed for reliable estimations of genetic diversity in fraxinus excelsior l. (oleaceae)?’, annals of forest science 65, 403–403. http://dx.doi.org/10.1051/forest:2008014 mohr, b., 2013, ‘the current status of laboratory animal ethics in south africa’, alternatives to laboratory animals 41(4), 48–51. murray, d.l. & fuller, m.r., 2000, ‘a critical review of the effects of marking on the biology of vertebrates’, in l. boitani & t.k. fuller (eds.), research techniques in animal ecology, pp. 15–64, columbia university press, new york. naef-daenzer, b., fruh, d., stalder, m., wetli, p. & weise, e., 2005, ‘miniaturization (0.2 grams) and evaluation of attachment techniques of telemetry transmitters’, journal of experimental biology 208, 4063–4068. http://dx.doi.org/10.1242/jeb.01870 national cancer institute, n.d., laboratory animal services programme, frederick national laboratory for cancer research: animal care and use committee guidelines, viewed 06 june 2016, from https://ncifrederick.cancer.gov/lasp/acuc/frederick/guidelinesfnl.aspx national research council, 2011, guide for the care and use of laboratory animals: eighth edition, national academic press, washington, dc. neiffer, d.l. & stamper, n.a., 2009, ‘fish sedation, anesthesia, analgesia, and euthanasia: considerations, methods, and types of drugs’, ilar journal 50(4), 343–360. http://dx.doi.org/10.1093/ilar.50.4.343 nielsen, l.a., 1992, methods of marking fish and shellfish, american fisheries society special publication 23, bethesda, md. o’grady, m.f., 1984, ‘the effects of fin-clipping, floy-tagging, and fin-damage on the survival and growth of brown trout (salmo trutta l.) stocked in irish lakes’, fish management 15(2), 49–58. http://dx.doi.org/10.1111/j.1365-2109.1984.tb00408.x osborne, n.j., payne, d. & newman, m.l., 2009, ‘journal editorial policies, animal welfare, and the 3rs’, the american journal of bioethics 9(12), 55–59. http://dx.doi.org/10.1080/15265160903318343 prince, e.d., snodgrass, d., orbesen, e.s., hoolihan, j.p., serafy, j.e. & schratweiser, j.e., 2007, ‘circle hooks, “j” hooks and drop-back time: a hook performance study of the south florida recreational live-bait fishery for sailfish’, istiophorus platypterus’, fisheries management and ecology 14, 173–182. russell, w.m.s. & burch, r.l., 1959, the principles of humane experimental technique, methuen, london. satheeshkumar, p., ananthan, g., senthilkumar, d., khan, a.b. & jeevanantham, k., 2012, ‘comparative investigation on haematological and biochemical studies on wild marine teleost fishes from vellar estuary, southeast coast of india’, comparative clinical pathology 21, 275–281. schaeffer, j. & hoffman, e., 2002, ‘performance of barbless hooks in a marine recreational fishery’, north american journal of fisheries management 22, 229–235. http://dx.doi.org/10.1577/1548-8675(2002)022<0229:pobabh>2.0.co;2 shahi, n., yousuf, a.r., rather, m.i., ahmad, f. & yaseen, t., 2013, ‘first report of blood parasites in fishes from kashmir and their effect on the haematological profile’, open veterinary journal 3(2), 89–95. snyder, d.e., 2003, electrofishing and its harmful effects on fish, information and technology report 2003-0002, u.s. geological survey, denver, co. society of environmental toxicology and chemistry, n.d., setac code of ethics, viewed 06 june 2016, from https://www.setac.org/?page=setacethics south african bureau of standards, 2008, the care and use of animals for scientific purposes, south african national standard 10386, sabs standards division, groenkloof. south african government, 1982, veterinary and para-veterinary professions act (act no. 19 of 1982), south african government, pretoria. stasko, a.b. & pincock, d.g., 1977, ‘review of underwater biotelemetry, with emphasis on ultrasonic techniques’, journal of the fisheries research board of canada 34(9), 1261–1285. http://dx.doi.org/10.1139/f77-189 stoskopf, m.k., 1993, ‘fish medicine’, saunders, philadelphia, pa. thorstad, e.b., rikardsen, a.h., alp, a. & ökland, f., 2013, ‘the use of electronic tags in fish research’, turkish journal of fisheries and aquatic sciences 13, 881–896. thorsteinsson, v., 2002, tagging methods for stock assessment and research in fisheries, report of concerted action fair ct.96.1394 (catag), marine research institute technical report 79, reykjavik. united kingdom government, 1986, ‘animals (scientific procedures) act’, her majesty’s stationery office, london. use of fishes in research committee (joint committee of the american fisheries society, the american institute of fishery research biologists, and the american society of ichthyologists and herpetologists), 2014, guidelines of the use of fishes in research, american fisheries society, bethesda, md. wagner, g.n. & cooke, s.j., 2005, ‘methodological approaches and opinions of researchers involved in the surgical implantation of telemetry transmitters in fish’, journal of aquatic animal health 17(2), 160–169. http://dx.doi.org/10.1577/h04-037.1 willis, t.j., 2001, ‘visual census methods underestimate density and diversity of cryptic reef fishes’, journal of fish biology 59, 1408–1411. http://dx.doi.org/10.1111/j.1095-8649.2001.tb00202.x willis, t.j., millar, r.b. & babcock, r.c., 2000, ‘detection of spatial variability in relative density of fishes: comparison of visual census, angling and baited underwater video’, marine ecology progress series 198, 249–260. http://dx.doi.org/10.3354/meps198249 winger, p.d. & walsh, s.j., 2001, ‘tagging of atlantic cod (gadus morhua) with intragastric transmitters: effects of forced insertion and voluntary ingestion on retention, food consumption and survival’, journal of applied ichthyology 17, 234–239. http://dx.doi.org/10.1046/j.1439-0426.2001.00280.x winter, j., 1996, ‘advances in underwater biotelemetry’, in b.r. murphy & d.w. willis (eds.), fisheries techniques, pp. 555–590, american fisheries society, bethesda, md. woodall, l.c., koldewey, h.j. & shaw, p.w., 2011, ‘serial monogamy in the european long-snouted seahorse hippocampus guttulatus’, conservation genetics 12(6), 1645–1649. http://dx.doi.org/10.1007/s10592-011-0253-6 world medical association, 2008, declaration of helsinki: ethical principles for medical research involving human subjects, world medical association, helsinki. appendix 1 table 1-a1: categorisation, based on animal ethical requirements, of 250 isi-rated, peer-reviewed, scientific journals with the most articles relating to fish research, revealed through a web of science (thomson reuters) search. table 1-a1 (continues…): categorisation, based on animal ethical requirements, of 250 isi-rated, peer-reviewed, scientific journals with the most articles relating to fish research, revealed through a web of science (thomson reuters) search. table 1-a1 (continues…): categorisation, based on animal ethical requirements, of 250 isi-rated, peer-reviewed, scientific journals with the most articles relating to fish research, revealed through a web of science (thomson reuters) search. table 1-a1 (continues…): categorisation, based on animal ethical requirements, of 250 isi-rated, peer-reviewed, scientific journals with the most articles relating to fish research, revealed through a web of science (thomson reuters) search. table 1-a1 (continues…): categorisation, based on animal ethical requirements, of 250 isi-rated, peer-reviewed, scientific journals with the most articles relating to fish research, revealed through a web of science (thomson reuters) search. appendix 2 table 1-a2: web links to guiding documents, policies and legislation for the ethical treatment of animals in research, identified by 45 isi-rated journals, in their instructions to authors. filelist convert a pdf file! supplement to koedoe. 1977: 248-254. summary of the proceedings of the symposium "the state of nature conserv a tion in southern africa" o martiny president wildlife society of southern africa pobox 44189 un den 2104 ut me start by saying that the symposium has, to a very large extent, been a report back from many departments and from many sections within and without the republic of south africa, and we are very grateful to the neighbouring states for the valuable contribution they made, in telling us what they are doing. i think this is a birthday celebration and not a funeral celebration, and therefore i hope that we can go away from here with some positive thoughts, rather than to bow down and get scared off by the heavy clouds that are hanging over the whole question, not only of conservation, but of the whole human population explosion survival. i must, however, without going into details regarding each person and what they have given us, mention just one or two. the main one, of course, as far as i am concerned, is dr r knobel, whose tireless energy over the many years has really meant building up the kruger national park. the second part is to thank his staff, and all the people who work with him. as you can see, they give him grey hairs but when you look at some of them, they also have grey hairs. it's a heavy task of organising and running this large organisation. thirdly, i would like to thank the speakers, without mentioning them individually. tome, the most important thing that emerged in this symposium was the absolute dedication with which the different speakers spoke, of where they were working in a field in which they thoroughly believed and they sold it to us with all the sincerity that was possible. i think that all of you in this field, are to be congratulated and we have to thank you that we have people like you to direct conservation in and around our borders. from the top of the ladder with the biggest, down to th e people with the smallest, it has been hard work and it has been very difficult. no reserve has just been created easily with the exception, of 248 course, that dr knobel goes up in a plane, looks out of the window and says, "that would be nice", and it was done. hocus, pocus. we hope he does it again some time. the other aspect which i found terribly interesting, was i don't know if all of you have experienced it the communication which took place at this symposium. as in so many cases, the real importance of a conference is afterwards, especially after one has had a couple of drinks and one's tongue loosens up thereby meeting each other and gaining a respect for each other and a mutual understanding. my plea at this symposium would be that scientists especially, see eye to eye, for they are always at each other because one is pinching the other's research, or the one does not agree with the other. may i make a plea that as we have got to know each other, and as we have got respect for each other, in future before we criticise, communicate, and maybe we will agree instead of fighting each other. i think that today is a very special day in which we are honouring the founder of the whole question of conservation in southern africa, namely president paul kruger. and when we think back to the days when he did this and even in later years when stevenson-hamilton had to fight the very difficult battle, we must remember that in those days man set out to conquer the wilderness. anybody who considered conserving it, was thought of as a crank. and yet, what they have persisted with, allows us to reap the benefits today. so if we encounter further battles and difficulties and seemingly hopeless situations, please just think back that if they could do it, we should be able to go further as well. despite difficulties, we have to win somewhere along the line. the outlook, of course, in those days was that the wilderness had to be tamed. the outlook today is that there is no more land to spare. and i do not think that this should dissuade us from attempting to get as much land conserved now and in the very near future, as we possibly can whatever the battles involved because if we do not do it now, maybe there will not be any further opportunity to do so. population explosion has popped up its head over and over again. it is something with which all of us are becoming more and more concerned. however, i wish to put in a word of warning: it is easy to say control population, but as you know, sex has come to stay and unfortunately, you cannot legislate on it and you cannot really legislate on birth control. that is the surest fuel that you can add to any revolution. you cannot do it by force, the only other way of doing it is by education. to implement birth control far more thought needs to be devoted to it and we must try to use every conceivable m~thod that we can think of, to limit this population explosion amongst all people. i think the security that we require, as i said, lies in the field of education. we do not have to indoctrinate people, but we have at least got to make them understand so that the decisions they make can be balanced. to this problem i would like to add, after the excellent talk that mr. 249 immelman gave us, (which many speakers said brought us back to reality), the immense problems that agriculture faces in this country. we are dealing with a land that is being despoiled, eroded and lost in encroaching desert -lack of water. the significant part of the paper was that by the year 2000, more or less, there will be no adequate water supply or land available. my question is: am i worried merely for the next twenty years, when i may still be alive, or should i worry about my child who will be alive beyond that? and if we are going to take measures similar to the population explosion, surely we don't wait until the over-used land is spoilt and degraded even further before we take action. when i said it was difficult to legislate on sex, it is just as difficult for any government politically to legislate on land. i think the time has come when one must think of land, not as belonging to yourself, bu t tha t you are the cus todian of tha t land for the generations to come. i would like to see action now, before we reach the situation where a farmer is told that a portion of his land should be rested and conserved by law. if this were to be the case, the food prices would go up, but does it really matter? if you take the price of food today, compared to the price of food ten years ago, we have had no control over that. it has just rocketed sky high and although i am probably one of the fattest here, none of you look starved you are all managing. but why must we wait until the year 2000, when so much more soil has gone into the sea, before we say, "now we are going to turn the screws on. now we must have a dictatorship to put it back into order". surely the time has come that if the farmer worked only part of his land and had to conserve part of it, that land might still recover and be conserved as part of the overall conservation policy. i put it to you as a thought. why wait and leave it to our children? are we too scared to tackle these problems now? apparently we have it so good and we are so selfish that we are not prepared to take action. we are leaving it for the future and we do not appreciate that we can do with less and not more. most of the speakers today and over the last few days were optimistic. i think this is right. it's a birthday and not a funeral. but this optimism is qualified by many "but's" and "irs". i think that reasons for conservation were conflicting, especially the one justifying it on economic grounds: not one of your speakers really believed in the economics of the matter. if you were mercenary, none of you would be in this job. you'd be out in business, you'd have earned the money and bought your own private game farm for your own selfish ends, for hunting or relaxation. but you're in the conservation game for idealism and belief in the conservation action. however, we have to justify this to others, to try to allow us to keep or conserve what we have got and to extend it. mr de la bat, of course, had a wonderful case. he proved the economics of it. i think that mr reilly in the swaziland setting, also proved that there are other reasons for conservation. but i would like to say to you, if conservation or game farming were an economic proposi2.')0 tion, why are so many of you farmers here not in game farming? because you do not believe it yet and you know it is not that easy. but there are glimmerings of hope in this. there are game farms that are in fact profitable. i was very glad recently to hear, and dr bigalke, of course, is involved in this in the northern cape, that with the change in prices, springbok farming is now becoming competitive with sheep farming in the same area; and obviously the utilisation of the veld is enhanced and the conservation of the land for the future is better. there are other examples where this approach works , and to go to extremes, go to the lion park it pays. but whether you call that conservation, of course, is a different story. but i think far more is needed and i know many people are trying to find and prove that conservation pays. but don't let us sell a concept which we ourselves are not sure of as yet. i would like to say to the farmer interested in conservation, that he should attempt to show that even part of his farm pays under conservation. nobody says that cattle and sheep and planted crops alone are the answer to our future food problems. there are many features that we are ignorant of let's try to provide the proof. to th e businessman i would say this: thos e of you who own game farms from money earned in other fields, think very seriously that twenty years from now, that ideal that you have conserved may not be there any longer unless you entrench it in clauses and give it to national parks , to provinces. that's the best chance that there is of conserving it. and i mean this very sincerely, that if you want that land which you've worked your life for as your am bi tion to have as a reserve , if you try to hang on to it and pass it on to your children, economics will come in and it will go under the plough. if it is given to the state, it will be conserved forever and that could be your contribution and future memorial. let me switch to another aspect. another reason why conservation is required is that we must not allow any species of plant or animal to become extinct. i am referring to the medical and related values. i'll try very briefly, without detailed knowledge, to give you a few examples. firstly, i want to mention that, as you all know, throughout the ages, plants have formed the basis of the medicines used by man. and even in modern times, uuggling with all the funny little organic chemistry molecules and particles of it), invariably the medicine is discovered from a plant before it is synthesized into the medicines that we use today. go back into history to a thing like the foxglove. if it were extinct we would not have digitalis. let's come nearer to modern times and take cortisone. the first cortisone was extracted from bulbs which came out of east africa and certainly provided a major trade. penicillin, streptomycin, which came out of measly little moles in the sewers under a city and because somebody observed that the bacteria in the vicinity were killed, we have these wonders of modern medicine. they, in part, are helping the population explosion the wrong way round, but let us 251 remember that it is of value to ourselves to keep healthy and strong, but it doesn't necessarily mean that we have got to add on many, many more individuals. the rh factor: this was discovered in blood while dealing with the question of jaundiced babies. the factor was found because of an inter-action between the blood of a rhesus monkey and that's where the "rh" comes from and human blood. if rhesus monkeys were extinct, we would possibly not have found this at all. furthermore, there is the question of smallpox. we all get vaccinated against it. if we didn't have cows, maybe we wouldn't have had smallpox. but let's go further and mention that a disease in cattle known as "elephant's skin disease" is also found in wild ebeest. a serum has been developed from wildebeest which is now used in cattle to stop this dis ease and thereby provides us with many valuable cattle hides that otherwise would have been lost. there are simple questions in this which are terribly fascinating. as you know, there is competition in nature. the dominant one takes over and there are likewise many diseases where you can cure on e dis ease by giving a person another disease. we have it in tuberculosis and in leprosy where th ere are many bacilli and the one gives you a cross immunity against the other. this is a field which onderstepoort and veterinarians around the world are far ahead of human medicine in researching these diseases, and much of what we have learned in medicine has come from a nimal research but mainly on domestic animals. when dr pienaar gave us that excellent outline of the vast research that is going on in the national parks and, of cours e, at the provincial level and elsewhere, it is clear that scientists are just scratching the surface. we have in the past us ed insects: honey bees to am erica , dung beetles to australia, lots of other controlled insects and pests like the cochineal on the prickly pear. there are so many of thes e unknown factors , of som e littl e creepy crawly thing sitting somewhere, that we know nothing about at this stage. can we afford to destroy it before we know whether this may be a possible aid that we require? at present they are res earching on plants to find anti-cancer agents. in laboratories they are still constantly researching on any plant to find medicines and other uses. comes our pollution, which plant can tak e it, which can survive , which can hold on to the soil before it goes, which can revive the soil to bring it back into use. all of these are questions that we do not know enough about and we are all amateurs or we are just scratching the surface to try to find out about it. destroy that plant or that animal and we will nev er get the answer. i think this is a very powerful aspect that we must think of. if we think of the worst, apart from the population explosion, one of the ways of curing it, of course, is the atomic bomb. but for those of us who might survive, which plants and which animals will be able to survive with us? how many are sensitive, how many are not sensitive and why? there are many angles. i do not think that we should 252 should be thought of, always accentuating conservation into the entire problem in as strong as possible a way. furthermore, such a department should become essential, not only internally in the republic of south africa, but also beyond our borders, where there should be co-operation, at the highest level. this is not the time for us to have so many different departments involved and mixed up, where so very often, a doesn't know what b is doing and through the lack of co-ordination it means time and waste of time and makes effective conservation very difficult. i would like a reassessment of this whole question of a department of conservation and environment in its own full right. i would like a reassessment on the whole question which was brought up by dr du plessis, of provincial reserves that really have the status of national parks, of even municipal reserves that really have the status of what should be provincial reserves, and of the question of what should be for tourism primarily and what should be wilderness or total conservation and, of course, the marine parks come into this what's in the sea and what's on the land nearby. should we not have a reassessment and a set of values drawn up soon, on deciding basic criteria by which people administer reserves, basic criteria which will be acceptable to all of you, but will lay down a set of rules by which people have to look after reserves. and should we not entrench as many of the real valuable reserves in this country under national, rather than allow it to be under provincial control? mr reilley mentioned very clearly that in swaziland, under their beautiful soil, lies coal. there is a potential for sugar and we all know that if anything of value to man for his immediate needs is found, that reserve or potential reserve may be doomed. but if such a reserve was granted national status, where its title was set up, then maybe we could entrench it so that if they remove something they have got to replace it with something next door ofa similar quality. but if it's done in bits and pieces here and there, you end up with different people going beyond what they really are aiming to do. and then, of course, the major plea before i end: all of you must have been acutely aware that a small country like malawi, since its independence, has taken the plunge and has put ii % of their land surface aside for conservation. we in south africa have something like 2,2% put aside. surely, we must aim to get at least 10% of our country conserved and the reason for it is going to be that such land may still be the most valuable entity to conserve in the year 2000 or to rebuild what may have been destroyed at that stage. ladies and gentlemen, i am sorry that i did not cover all the papers individually, but i think that the message that came out of the papers has been accentuated by me. may i, on behalf of the national parks board of trustees, thank all of you who have participated speakers and audience, for a very wonderful few days and once more thank the officials for what they have made of this occasion. thank you. 254 article information authors: jacobus du p. bothma1 marius d. bothma2 affiliation: 1centre for wildlife management, university of pretoria, south africa 2halls head, mandurah, australia correspondence to: jacobus bothma postal address: po box 2726, george 6530, south africa dates: received: 23 feb. 2012 accepted: 09 july 2012 published: 11 oct. 2012 how to cite this article: bothma, j. du p. & bothma, m.d., 2012, ‘leopard range size and conservation area size in the southern kalahari’, koedoe 54(1), art. #1076, 4 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1076 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. leopard range size and conservation area size in the southern kalahari in this short communication... open access • abstract • introduction • study area • research method and design • results • ethical considerations • trustworthiness    • reliability    • validity • discussion • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ the range use patterns of adult leopards were used to examine the impact of environmental quality on conservation area size in the arid south-western portion of the kgalagadi transfrontier park in southern africa. the ranges of the leopards are the largest recorded in the world, with a mean size of 2104.4 km2(sem995.95 km2) for males and 1258.5 km2 (sem 541.50 km2) for females. overlaps in range use within and between the sexes and the size of this conservation area make it possible to sustain a genetically viable population of leopards in this arid environment.conservation implications: when establishing conservation areas that contain large carnivores in arid and semi-arid regions, prey abundance and range use should be considered for the area to be able to sustain viable populations of such carnivores. the results emphasise the importance of establishing large transfrontier conservation areas where individual conservation areas are too small to do so. this study is the first to do so for leopards in southern africa. introduction top ↑ opportunistically hunting apex predators such as the large cats will hunt every prey animal that is encountered and their survival is therefore dependent on prey abundance and geographic distribution, which are a reflection of habitat quality through environmental conditions such as rainfall and ambient temperature (coe, cumming & phillipson 1976; sunquist & sunquist 2002). consequently, the size and location of a conservation area will affect its ability to support genetically viable populations of leopards, panthera pardus. range use by the females of large solitary felids depends on prey abundance and distribution whilst that of males depends on the geographic distribution of females and is a product of social behaviour (sunquist & sunquist 2002). it is therefore expected that the ranges of the leopard in semi-arid and arid regions will be larger than in mesic ones and that behavioural adaptations such as range use overlap between adult males may impact on viable leopard conservation area size, because if the ranges of the leopards did not overlap, as happens in many mesic regions, it would require larger conservation areas to sustain a genetically viable leopard population. in the prey-rich equatorial forests at the edge of the chitwan national park in nepal, for example, the ranges of adult male leopards are exclusive and small (mean size: 10 km2) (seidensticker, sunquist & mcdougal 1990), whilst in the savannas of the more semi-arid kruger national park the ranges are larger (mean size: 25 km2) and there is a mean overlap of 19% in range use by the adult males (bailey 1993). in the most arid, south-western part of the kgalagadi transfrontier park the ranges of leopards are the largest recorded globally, but range use overlaps and core area range use sizes were not indicated by bothma et al. (1997). because leopards are socially solitary animals, it is also expected that an adult male will only make contact with an adult female when she is in oestrus.according to prof. b. jansen van vuuren (genetics laboratory, department of zoology, university of johannesburg, pers. comm., 05 june 2012) it was previously believed that a minimum population size of 500 animals was required to maintain a genetically viable population. however, it is now recognised that this will depend on many aspects, including genetic heterogeneity and species-specific traits, although the absolute minimum population should preferably not be less than 50 breeding animals. nevertheless, at the time of its declaration in 1931, the addo elephant national park contained only 11 elephants, loxodonta africana, whilst this population now numbers in excess of 700 animals (preller 2008). three hypotheses were briefly studied here. the first was that the ranges of leopards in the most arid south-western portion of the kgalagadi transfrontier park will be larger than in mesic regions. the first was that the ranges of leopards in the arid south-western portion of the kgalagadi transfrontier park will be large. the second that behavioural adaptations such as range use overlap by adult males may impact on the viability of the size of this conservation area for leopards. the third was that an adult male and female would probably only make contact when she was in oestrus. study area top ↑ the kgalagadi transfrontier park is a combination of the kalahari gemsbok national park of south africa (9591 km2) and the gemsbok national park of botswana (28 400 km2) and hence has a surface area of 37 991 km2. the ambient temperature and annual rainfall vary greatly, with daytime temperatures as low as −10.3 °c and as high as 45.4 °c in the south-western region. the erratic annual rainfall has a mean of 185 mm in the south-western, and 450 mm in the north-eastern regions. the variable, and often localised, rainfall affects primary productivity and causes the larger herbivorous mammals (potential leopard prey) to be distributed unevenly in time and space (bothma & mills 1977; mills & retief 1984; van der walt et al. 1984; van rooyen et al. 1990). there are 20 landscapes in the kalahari gemsbok national park, with dunes featuring prominently in many of them (van rooyen et al. 2008). this study was conducted in the most arid south-western portion of the kgalagadi transfrontier park. research method and design top ↑ the range sizes that were calculated by bothma et al. (1997) were based on very high frequency (vhf) radio-collar locations that were analysed with early kernel methods for 95% (total range use) and 50% (core area of use) isopleths. to standardise these according to currently accepted approaches (börger et al. 2006), the original data were re-analysed with the arcview 3.2 version 1.1 and the animal movement extension to arcview software ( http://esri/software/arcgis/arcview) of hooge and eigenlaub (1997) and by using 90% and 50% isopleths respectively. the data are based on the geographic locations of three adult males from 10 june 1988 to 16 november 1989 and five adult females from 15 october 1987 to 11 october 1992 (bothma et al. 1997). in addition, global positioning system (gps) data on three adjacent adult males and two females later became available for range use overlap studies. the locations of an adult male and female relative to each other were then also recorded daily from 14 november 2000 to 03 april 2001 after they were collared on the same day, 15.8 km apart. these and all the more recent location fixes were based on the use of earthspan satellite gps radio-collars that were linked to an argos satellite. only those locations with a location fix accuracy of 350 m or less were used in these analyses. results top ↑ maps of the original localities of occurrence of the leopards studied over time appear in bothma et al. (1997) and will not be repeated here. the re-analysed range use data yielded a mean total range size of 2104.4 km2 (standard error of the mean 995.95 km2) for adult males and 1258.5 km2 (sem 541.50 km2) for females, as opposed to means of 2182.4 km2(sem 283.49 km2) and 1488.3 km2(sem 665.61 km2) respectively in the earlier study of bothma et al. (1997). the mean core area of range use for the adult males was 499.9 km2 (sem 297.40 km2 ) and 292.4 km2 (sem 145.59 km2) for the females, but core areas of range use were not calculated by bothma et al. (1997). there was a mean overlap of 48.3% (sem 18.67%) of range use by three adult males and one of 30.6% by two adult females. the three males used a mean of 30.0% (sem 8.15%) of the ranges of two females and the two females one of 51.6% of the ranges of the three males.the adult male and female that were gps-collared close to each other yielded 388 accurate location fixes for the male and 618 for the female. the collar of the male stopped functioning on 03 april 2001 and data gathering for the female ceased on 14 july 2001. on four of the 104 days of study, these locations were within 100 m of each other (well within the margin of error of the location fixes that were used) or concurred. these incidents were 33, 35, 35 and 37 days apart, with a mean interval of 35.0 days (sem 0.82 days). they all occurred within a 294.5 km2 portion (27.2%) of the total range of 1082.5 km2 that was used by the male during the 104 days of daily satellite tracking. three of the four incidents were within the area of overlap between the ranges of the male and female. in the exception, the female had moved deeper into the range of the male. one incident was in the core area of range used by the female and two in the core area of range used by the male. on the days before possible contact, the male and female were a mean of 3.1 km apart (sem 0.96 km), and on the days following such an incident, one of 8.2 km (sem 1.50 km) apart. on the other 98 days of the study, they were a mean of 17.4 km (sem 0.93 km) apart. the above-mentioned male and female leopards did not use all parts of their total ranges or the core areas of these ranges every month. for the four months that data were available, the male used a mean of 72.1% (sem 9.58%) of his total range and 77.4% (sem 6.30%) of his core area of range per month. for the eight months that data were available for the female, she used a mean of 43.4% (sem 11.18%) of her total range and 54.5% (sem 15.15 %) of her core area of range per month. ethical considerations top ↑ the leopards were collared by staff of south african national parks, south africa, according to national and international guidelines that existed there and at the university of pretoria at the time of the study. the project was formally approved by the then south african national parks board. no formal ethical protocols were applicable at the time of the research for the univeristy of pretoria or the south african national parks. no students or other field researchers were involved in collecting the data because the data were downloaded from orbiting satellites. the leopards were darted and fitted with gps radio collars according to accepted norms and were not approached or handled subsequently. the staff involved had the medical benefits of their organisation. the data used here were only used by the authors. trustworthiness top ↑ the results and conclusions drawn are based on the observations of the senior author who conducted the data interpretation following the geographic information systems (gis) analysis. reliability the techniques used are generally accepted for repetitive range use studies at a given time although it is recognised that changing environmental conditions can yield variations in it. this is a normal ecological occurrence. however, the same general pattern of range use will emerge even if the specific data were to change to some degree. it is the pattern and not the specifics which is important to generalise the theory. validity the hypotheses that were tested and accepted validate the generalised theories that were developed. discussion top ↑ whilst the use of kernel estimates for range use calculation was questioned by hemson et al. (2004), subsequent studies have confirmed that it is an efficient method to do so provided that 90% isopleths are used to estimate the total area of range use and 50% isopleths to estimate the core area of range use (börger et al. 2006). however, the minimum convex polygon method should rather not be used (nilsen, pedersen & linnell 2008). in the current study, the data that were derived from vhf and gps tracking were not combined because of possible biases when using two different methods to collect the data.the mean interval of 35 days between possible contacts between the male and female fits the known mean inter-oestrus period of 46 days (range: 20–55 days) in leopards (sadleir 1966). this seems to indicate that an adult male and female leopard will only make contact when the female is in oestrus. given the mean range use overlap that was found within the leopard sexes in this study, the kalahari gemsbok national park portion of the kgalagadi transfrontier park is expected to be able to support 10 adult males and 25 adult females (total population: 35) whilst the gemsbok national park is expected to be able to support 37 adult male and 99 adult female leopards (total population: 136). the combined conservation area therefore seems to be large enough to support 171 adult leopards, which is more than the proposed minimum population of 50 leopards to maintain a genetically viable population, but the kalahari gemsbok national park portion alone cannot do so. however, the leopard population in the kgalagadi transfrontier park may well be larger, because the study was done in the most arid south-western portion, whilst funston (2001) found 4.5 times more leopard tracks in the north-eastern part of this park than in the south-western portion. this difference could be linked to an expected higher primary productivity (coe et al. 1976) in the north-eastern portion because the rainfall in this park increases from the south-west to the north-east. therefore, the real population size of leopards in the kgalagadi transfrontier park may potentially be greater than when it is based only on the data from the south-western portion of it. moreover, the extent of range use overlap of possible uncollared leopards in the study is unknown. the large variations in range size and use that were found made valid statistical analyses of the means impossible to calculate because they led to large standard deviations and errors of the means. moreover, it is extremely difficult to find and collar all the possible leopards in a study of range use in the vast and largely trackless southern kalahari. because prey distribution and abundance appear to determine the range size of adult female leopards and their range sizes will influence those of the males, the range use by leopards in the kgalagadi transfrontier park must respond over time to the rainfall variations that affect the abundance and distribution of the larger herbivore prey (bothma & mills 1977, mills & retief 1984, van der walt et al. 1984; van rooyen et al. 1984). although leopards also prey on a wide range of other prey, their main prey animals are herbivores in the size range from 20 kg to 70 kg (bothma & le riche 1984, 1986, 1994). conclusion top ↑ the range sizes and range use overlaps of the leopards in the southern kalahari are the largest recorded in the world. because leopards occur in a wide range of habitats, the size of conservation areas that contain leopards should take the variation in their range size use in various habitats into consideration so as to ensure that they can contain genetically viable populations. adaptations in social behaviour such as the within-sex range use overlap that is found in prey-poor, arid regions make it possible for the prey-poor conservation areas such as the kgalagadi transfrontier park to support a larger leopard population than would be the case if no range use overlaps occurred. this study therefore confirms the value of creating large conservation areas, especially in arid regions, where large felids occur. the results support the hypotheses that the ranges of leopards in semi-arid and arid regions are larger than in mesic ones and that behavioural adaptations such as range use overlap between adult males may impact on viable conservation area size for leopards because they would have had to have been larger if no overlaps occurred. the admittedly small sample size of the distance and contact between an adult male and female within her oestrus cycle also confirms the extensive spoor tracking data, which showed that adult leopards in the southern kalahari are largely solitary and that they only make contact when a female is in oestrus (bothma & le riche 1995). the study opens up an interesting avenue of future research. a study of leopard range use at selected intervals along the rainfall gradient from the more arid south-western to the wetter north-eastern region of this park, combination with prey distribution and abundance studies and genetic analyses of the leopards, should more conclusively indicate the influence of this environment on leopard range use. it should also confirm whether the kgalagadi transfrontier park contains a genetically viable leopard population or not. a study of the range use by leopards at selected intervals along the more arid south-west to wetter north-east rainfall gradient will enable the more accurate calculation of the leopard population in this park. a similar study of prey abundance and distribution will explain the range use patterns more clearly. genetic analyses will be conservationally important. in combination, these studies will help to determine whether this transfrontier park can sustain a genetically viable leopard population or not. acknowledgements top ↑ the capture and radio-collaring of the leopards in this study occurred under strict supervision of trained staff of the south african national parks and followed the ethical code of that organisation and of the university of pretoria. the south african natural history unit of route 66 films allowed the use of some satellite radio-collar data. funding was received from the university of pretoria and shayamanzi game (pty) ltd (http://www.leopard.tv/index.php). the then warden e.a.n. le riche gave valuable field support and b. bothma typed numerous drafts of the manuscript. we sincerely thank them all. competing interests the authors declare that they have no financial or personal relationship(s) which may have inappropriately influenced them in writing this paper. authors’ contributions j. du p.b. (university of pretoria) was the project leader, was responsible for the project design, did the field research, collected the satellite gps location details, analysed and interpreted the results and wrote the manuscript, whilst m.d.b. (halls head) did the arcview and gis analyses. references top ↑ bailey, t.n., 1993, the african leopard, columbia university press, new york.bothma, j. du p. & le riche, e.a.n., 1984, ‘aspects of the ecology and behaviour of the leopard panthera pardus in the kalahari desert’, koedoe (supplement) 27, 259–279. http://dx.doi.org/10.4102/koedoe.v27i2.585 bothma, j. du p. & le riche, e.a.n., 1986, ‘prey preference and hunting efficiency of the kalahari desert leopard’, in s.d. miller & d.d. everett (eds.), cats of the world: biology, conservation and management, pp. 389–414, national wildlife federation, washington, d.c. bothma, j. du p. & le riche, e.a.n., 1994, ‘scat analysis and aspects of defecation in northern cape leopards’, south african journal of wildlife research 24(1 & 2), 21–25. bothma, j. du p. & le riche, e.a.n., 1995, ‘evidence of the use of rubbing, scent-marking and scratching posts by kalahari leopards’, journal of arid environments 29, 213–219. bothma, j. du p. & mills, m.g.l., 1977, ‘ungulate abundance in the nossob river valley, kalahari desert’, in t.j. peterle (ed.), proceedings of the xiiith international congress of game biologists, the wildlife institute, washington d.c., atlanta, march 10–15, 1977, pp. 90–102. bothma, j. du p., knight, m.h., le riche, e.a.n. & van hensbergen, h.j., 1997, ‘range size of southern kalahari leopards’, south african journal of wildlife research 27(3), 94–99. börger, l., franconi, n., de michele, c., gantz, a., meschi, f., manica, a. et al., 2006, ‘effect of sampling regime on the mean and variance of home range size’, journal of animal ecology 75, 1393–1405, viewed 15 may 2012, from http://www.zoo.com.uk/zoostaff/manica/ms/2006_borger_et_al_j_anim_ecol.pdf coe, m.j., cumming, d.h.m. & phillipson, j., 1976, ‘biomass production of large herbivores in relation to rainfall and primary production’, oecologia 22, 341–354, viewed 15 may 2012, from http://www.deepdyve.com/lp/springer-journals/biomass-and-production-of-large-african-herbivores-in relation-to-9gm4lg2dhc funston, p., 2001, kalahari transfrontier project: final report, endangered wildlife trust, parkview. hemson, g., johnson, p., south, a., kenward, r., ripley, r. & macdonald, d., 2004, ‘are kernels the mustard? data from global positioning system (gps) collars suggest problems for kernel home-range analyses with least squares cross validation’, journal of animal ecology 74, 455–462, viewed 15 may 2012, from http://www.onlinelibrary.wiley/com/doi/10.1111/j.1365-2656.2005.00944.x hooge, p.n. & eigenlaub, b., 1997, animal movement extension to arcview version 1.1, anchorage, alaska biological science center, u.s. geological survey. mills, m.g.l. & retief, p.f., 1984, ‘the response of ungulates to rainfall along the riverbeds of the southern kalahari’, koedoe (supplement) 27, 129–141. http://dx.doi.org/10.4102/koedoe.v27i2.574 nilsen, e.b., pedersen, s. & linnell, j.d.c., 2008, ‘can minimum convex polygon home ranges be used to draw biologically meaningful conclusions?’, ecological research 23, 635–639. http://dx.doi.org/10.1007/s11284-007-0421-9 preller, b., 2008, the silent giants of southern africa (loxodonta africana), 2nd edn., bob preller, wilderness. sadleir, r.m.f.s., 1966, ‘notes on reproduction in large felidae’, international zoo yearbook 6, 184–187, viewed 15 may 2012, from http://www.onlinelibrary.wiley.com/doi/10.1111/izy.1966.6.issue-1/issuetoc seidensticker, j., sunquist, m.e. & mcdougal, c., 1990, ‘leopards living at the edge of royal chitwan national park, nepal’, in j.d. daniel & j.s. serrao (eds.), conservation in developing countries: problems and prospects, pp. 415–423, bombay natural history society, bombay. sunquist, m. & sunquist, f., 2002, wild cats of the world, chicago university press, chicago. van der walt, p.t., retief, p.f., le riche, e.a.n., mills, m.g.l. & de graaff, g., 1984, ‘features of habitat selection by large herbivorous mammals and the ostrich in the southern kalahari conservation area’, koedoe (supplement) 27, 119–128. http://dx.doi.org/10.4102/koedoe.v27i2.573 van rooyen, n., bezuidenhout, d.j., theron, g.k. & bothma, j. du p., 1990, ‘monitoring of the vegetation around artificial watering points (windmills) in the kalahari gemsbok national park’, koedoe 33, 63–88. http://dx.doi.org/10.4102/koedoe.v33i1.453 van rooyen, m.w., van rooyen, n., bothma, j. du p. & van den berg, h.m., 2008, ‘landscapes in the kalahari gemsbok national park’, koedoe 50(1), 99–112. http://dx.doi.org/10.4102/koedoe.v50i1.154 article information authors: sharon pollard1 derick du toit1 harry biggs2 affiliations: 1association for water and rural development, acornhoek, south africa 2south african national parks, skukuza, south africa correspondence to: sharon pollard email: sharon@award.org.za postal address: private bag x420, acornhoek 1360, south africa dates: received: 22 june 2010 accepted: 22 feb. 2011 published: 13 may 2011 how to cite this article: pollard, s., du toit, d. & biggs, h., 2011, ‘river management under transformation: the emergence of strategic adaptive management of river systems in the kruger national park’, koedoe 53(2), art. #1011, 14 pages. doi:10.4102/koedoe.v53i2.1011 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) river management under transformation: the emergence of strategic adaptive management of river systems in the kruger national park in this essay... open access • abstract • introduction    • the broader context of the development of a management response approach in the kruger national park    • the worsening situation: a brief overview of the lowveld water resources    • water for game programme: attempts to dampen variability    • small sections of rivers and catchments in the kruger national park    • institutional arrangements • the formal adaptive response: developing a strategy to respond to the challenges posed by declining river system integrity    • the knp rivers research programme and adaptive management    • a new phase of research: linking outputs to management    • participation in wider catchment forums: the development of the catchment management strategy • lessons that emerged from learning in action • conclusion • references abstract (back to top) protected areas such as the kruger national park (knp) face many management challenges, of which ensuring a healthy flow of rivers into the park is one of the most important. although previous management policies isolated the knp from its neighbours, this position has changed as the knp seeks to negotiate a respected ‘place’ for water and conservation in a competitive environment. a major catalyst for this re-orientation has been the response from the knp to the growing water crisis where its position needed to be seen within the wider catchment and policy context in south africa. this paper presents an overview of the transforming management practices of the knp in a changing political, socio-economic and environmental context, through the lens of water resources. we show that the knp management model moved beyond inward-looking, isolationist policies to adopt responsivity to major change factors. the new approach was applied first in the sphere of river management in the knp after which it spread to other domains such as fire and game management. it explicitly incorporates an experimental–reflexive orientation and considers management as a process of learning-by-doing. this paper strives to review the transformation since the onset of explicit adaptive management of these rivers. the development of a new stewardship, based on a stakeholder-centred vision and on learning-focused management, has been a main achievement for the knp. a closer partnership between researchers, managers and field staff, supported with buy-in and co-learning, has led to a management framework based on a clear vision informed by stakeholder involvement, an objectives hierarchy, a scoping of management options, a monitoring system and a reflective evaluation process with feedback loops. although developed through a focus on rivers, the framework can be embraced for the management of ecosystems as a whole. conservation implications: the explicit adoption of strategic adaptive management for the rivers entering the knp has had considerable implications not only with regard to management practice within the park, but also for the relationships with neighbours. this has also meant setting and implementing new goals and priorities with managers and staff. introduction (back to top) the broader context of the development of a management response approach in the kruger national park protected areas such as the kruger national park (knp) face many daunting management challenges, of which ensuring a regular and healthy flow of rivers through the park is arguably one of the most important. although previous management policies isolated the knp from its neighbours, the park is not an ‘island’ (pollard, shackleton et al. 2003). the park’s responses to the growing water crisis (pollard & du toit 2011) are increasingly being referenced against the wider context of water management in south africa, which includes attention to the new water legislation and options for negotiating a respected ‘place’ for water and conservation in a competitive economic environment. the aim of this paper is to present an overview of the adaptive management practices of the knp in a changing political, socio-economic and environmental context, from a water resource perspective. it tracks the park’s approach to water management as it changed from inward-looking, isolationist policies to one focused on responsivity to major change factors (venter et al. 2008). water is an interesting lens through which to examine managerial transformation in the park because, as a fugitive resource, it ‘knows no boundaries’. for research and management to be meaningful, one should therefore look beyond the boundaries imposed by administrative and political systems. in this regard the knp is – within south africa, at least – the ‘downstream user’ of six major rivers that traverse or border the park. a wider view recognises that south africa and the knp lie upstream of or adjacent to international neighbours mozambique, zimbabwe and swaziland, with whom water must be shared and managed (figure 1). thus, no account would be complete without consideration to the wider context within which the knp is embedded. an in-depth account of the socio-economic and environmental profile of the this context has appeared in several publications (e.g. dovie, shackleton et al. 2006; niehaus 2001; pollard et al. 2003; pollard, biggs et al. 2008; ramutsindela 2002; stadler 1994) and hence we provide only an overview here to highlight major characteristics. the knp is located in the lowveld, an area that includes the knp and an additional stretch of land around it (50 km wide) on the park’s western border (see pollard et al. 2003). the rivers that traverse the knp cross into mozambique, where the lowveld extends to coastal floodplains and estuaries. the landscape includes a rich variety of landforms, climate and vegetation, as well as cultures and land uses. the climate is tropical to subtropical, with drought being endemic to the region (tyson 1986). the drakensberg escarpment, exceeding 1800 m a.s.l., descends rapidly eastward to the lowveld of the knp and mozambique, with average altitudes of 600 m and 400 m, respectively. similarly, yearly rainfall declines from more than 1200 mm along the escarpment to less than 450 mm at the eastern border with mozambique. at least two million people live within 50 km of the western border of the knp, with some 500 000 on the eastern border in the area of mozambique included in the great limpopo transfrontier conservation area (gltfca) (cumming and ahead 2004, although according to personal communication with d. cumming, demographic data are uncertain). there is a diversity of cultures and major groups include the tsonga, the vhavenda, the pedi and the swazi. in south africa these groups were segregated under the apartheid government into the former bantustans of gazankulu, venda, lebowa and kangwane, respectively, from the late 1960s until the first democratic government was elected in 1994. in contrast to the adjacent, sparsely settled commercial farms historically owned largely by whites (5–20 people per square kilometre), these former homelands experienced densities as high as 300 people per square kilometre as a result of forced relocations (pollard, mendiguren et al. 1998). the legacy of apartheid means that the former bantustan areas are still characterised by high levels of poverty today, with large disparities (especially between people living inside and outside these areas) in access to water, sanitation, education and employment opportunities as well as attendant environmental degradation as people tried to eke out a living on marginal land (pollard et al. 1998). since 2002, with new conservation ventures being launched, a part of mozambique was incorporated into what is the easterly component of the great limpopo transfrontier park (gltp) and the gltfca. as was the case with the ‘homelands’ in south africa people living here also experienced a series of re-settlements – in this case due to colonial rule, the civil war and now the gltp (refugee research 2002). the period between the 1960s and the 1980s was marked by economic expansion, mainly in the form of commercial agriculture, including afforestation, and some limited mining (see pollard, riddell. et al. 2010). during this time water was considered only insofar as it was needed. in keeping with global practices, water deficits were dealt with through increased infrastructure such as dams and interbasin transfers (pearce 1992). issues of sustainability or equity were of little concern and, indeed, this period of uncontrolled development was unconstrained by water resources. by the early 1990s, south africa faced some major socio-political changes, which were to have a direct impact on the knp (see carruthers 1995; pollard et al. 2003; pollard & du toit 2007). the high density of poor people on the western boundary lead to an environmental and economic situation that fostered conflict over land and resources (stadler 1994). the collapse of the apartheid regime and the establishment of a democratic government in 1994 heralded the transformation of the politico-legal environment in south africa. owing to changes to land ownership policies and land reform, the future of the park and the definition of its boundaries are now embedded in a different socio-political climate. protected area legislation underwent transformation, which redirected conservation endeavours to reflect the intentions of the south african constitution. the former legislation that goverened parks was replaced by the new protected areas act in 2003. legislation with regard to water resources underwent a fundamental re-orientation with the promulgation of the national water act in 1998 (south africa 1998), moving from a system of riparian rights to one in which there is no private ownership of water but rather state custodianship. underlying the transformation is the challenge of redressing past inequities and supporting development whilst ensuring the sustainability of the resource base upon which livelihoods and a healthy environment is founded. sustainability, together with equity, took centre stage with the recognition of the reserve as a constitutional right to water (department of water affairs and forestry 2004). the national water act of 1998 defines the reserve as a quantity of water to sustain basic human needs and to ensure the sustainability of the resource itself. a major change was that water was to be managed from a more holistic, catchment perspective with consideration to stakeholder participation. this signalled a recognition of linkages – not only between upstream and downstream use or between land and water, but also between people and resources – in water resource production and management. it is in this context that the democratic government of south africa has re-orientated the management of water resources, and the knp responded by adopting new policies and management approaches. this paper aims to provide a synthesis of a number of research initiatives that have been launched since formal adaptive management was first explicitly adopted in the early 1990s, most notably phases ii and iii of the knp rivers research programme (knprrp), ecosystems, protected areas and people project of the international union for conservation of nature (iucn-epp) and more recent work under the shared rivers initiative (sri) (pollard & du toit 2011). perhaps because of the key importance of freshwater, and the variability inherent in river function, adaptive management arose first in the river context in the knp and later spread to other areas of natural resource management in the park (freitag et al., in review). the knprrp ended in 2000 and has been fairly well documented as an entity (breen et al. 2000), but relatively little new development or documentation emerged since. although the knp continued to implement adaptive protocols catchment management agencies (cmas) were anticipated to be formed to allow the work of the knprrp to be put into practice. however, real-world lags led to delays, which eventually triggered a feeling that the knprrp should have continued. a new action research initiative was thought to be needed for further work, which led to the eventual formulation of the sri in 2007 (see pollard and du toit [2011] for a full description). however, between 2000 and 2005, a separate research programme, known as the river savanna boundaries programme, was run in the knp with collaborators from south africa and the united states. the programme focused on landscape ecology and on the links between riverine and terrestrial systems, which contributed greatly to the understanding of heterogeneity as an organising factor in ecosystems. the programme helped to mainstream the heterogeneity paradigm in the knp and also, generally, in the minds of savanna scientists, particularly because of a very successful annual meeting started under the auspices of the programme and the subsequent publication of the kruger experience: ecology and management of savanna heterogeneity (du toit et al. 2003) assisted by funding from the programme. an opportunity arose in 2004 under the iucn-epp to document the experience and lessons learned from the knprrp. the purpose of the iucn-epp was to enable managers of protected areas to implement, in the face of global change, adaptive management strategies through a number of mechanisms, one of which was to document lessons from key case studies such as that of the knp. the authors subsequently conducted research along with a range of park staff regarding the changing nature of river research and management (pollard & du toit 2006, 2007). by 2007, the need for consequences and lead-ons from the knprrp led to the start of the sri. the sri has since examined the complexities of achieving compliance with the environmental water requirements (ewr) of the lowveld rivers that flow through the park (pollard & du toit 2009). in south africa ewrs, also known as the ecological reserve, are used as benchmarks for assessing sustainability in freshwater resources (see, for example, pollard and du toit [2008]). sustainability is one of the cornerstones of the 1998 national water act. within the knp two key change factors frame contemporary management practices, which are fundamentally different from those of the past. the first is the alarming decline in surface water quality and quantity (pienaar 1970), together with the associated biodiversity changes of the rivers that flow through the knp (see later). the second is the recognition that lowveld savannas are not stable-state agricultural systems but rather that heterogeneity and flux are inherent characteristics (du toit, rogers et al. 2003; peel 1999; rogers & o’keeffe 2003). in retrospect, it is interesting to note how closely interlinked the histories of these factors have been and some suggest that their mutual influence has guided transformed management of the knp over the past 15 years (pollard & du toit 2005; van wilgen & biggs 2010). these factors had a number of implications. firstly, because all six perennial rivers originate outside of the park, the knp has had to broaden its areas of engagement to include a catchment-based perspective. this situation is not unique to the knp; many of the world’s protected areas rely on rivers of which the catchment areas are not co-incident with park boundaries. secondly, in recognition of the dynamic nature of ecosystems (and indeed the socio-ecological system, as described by pollard, biggs et al. [2008]), the knp embarked on an approach that explicitly incorporates an experimental orientation and which views management as adaptive, that is, learning by doing (kingsford et al. 2011). the co-evolution and influence of these two factors will be explored in the following sections. another notable aspect to this overview is how the practical outcomes of an adaptive approach – development of an objectives hierarchy and thresholds of potential concern (tpcs) – have been extended far beyond the management of water alone (freitag et al. in review) and now frame all aspects of ecosystem management (such as fire and herbivores) in the knp. what transpired was the complete overhaul of a management approach: a change from one of immutable goals and objectives based on stable-state ecosystem theory to one based on learning-management iterations designed to maintain variability, which is a fundamental attribute of semi-arid savannas and rivers (davies, o’keeffe et al. 1995). we wish to use the narrative of water resources management to elaborate on the adaptive management approach and its evolution within the knp. figure 1: tmhaep s ionudtihca atifnrigc athne n matiaojonra rli vpearr kssy s(tseamnsp anrkds )a sscoiecniactee sdu cpaptcohrtm pernotcse ossf tinhdei ceaatistnegr nth ees crearlaptimoennsth,i plo bwevtewlde ean dth ker usgixe pr rnoactieossn astl eppasr.k, south africa. anthropogenic changes have meant that of the six historically perennial systems, only the sabie river has remained so. the worsening situation: a brief overview of the lowveld water resources changes to the river systems have been evident since the 1960s (pienaar 1970) and since then most systems have experienced progressive degradation in quantity and quality and associated fauna and flora. of the six river systems of the park, five were perennial and one, the shingwedzi, was naturally seasonal (o’keeffe & davies 1991). the first deterioration was evident already some 45 years ago when the perennial letaba ceased flowing and subsequent cessations have transformed this river into a non-perennial system. a similar situation occurred in the luvuvhu river in the 1960s and later in the olifants river. the crocodile river has experienced flow constancy as well as a seasonal reversal as a result of regulation (see pollard & du toit 2010), and both the crocodile and olifants have suffered heavy pollution and invasion by alien plant species. the sabie river is regarded as the least perturbed of the major rivers of the knp, with relatively small distributional changes in fish species (russell & rogers 1988). in comparison, an (at least transient) loss of species has characterised the other knp rivers: the letaba, olifants and crocodile rivers appeared to have lost between four and six fish species, and the luvuvhu river has lost nine species. most of these have re-appeared at lower abundances. agricultural abstraction is regarded as the primary driver for the increasing demands on the water resources (o’keeffe & davies 1991; pollard, riddell et al. 2010). that, together with afforestation and in some areas mining, has been implicated in the hydrological modifications evident today (chunnett 1990; water research commission 2001). this situation has been exacerbated by the past allocation inequities between the various user sectors and has been accompanied by escalating tensions. for example, the sand river catchment boasts the highest percentage of afforestation of any catchment in south africa, and the associated reduction in streamflow (smith & scott 1992) has led to disputes between timber growers and other downstream users during past dry cycles (pollard et al. 1998). recent studies have indicated that all lowveld catchments, with the exception of the sabie, are in water deficit (demand exceeds availability) or will be once the requirements for the ecological reserve are met (dwaf 2004a,b,c; dwaf 2009; pollard, riddell. et al. 2010). the aridity of the lowveld and the frequency of drought, coupled with current and projected population densities and water demands, mean that there is insufficient water to meet current needs at the required assurance levels in most catchments (pollard et al. 1998). this situation provided a drive for further water resource developments, mainly in the form of dams, although the enthusiasm for impoundments has waned somewhat since 1994, and has been replaced by a more integrated approach to water resource management (pollard et al. 2007), in which the knp has been instrumental in setting the tone. indeed, in some cases this has meant reviewing and revising earlier policies and actions to the extent that dams within the park, such as the shimuwini dam (figure 2), have been destroyed figure 2: (a)the shimuwini dam and (b) its removal in 2004. water for game programme: attempts to dampen variability already in the 1970s the warden of the knp realised that upstream impacts on the river systems were being felt within the park and correctly surmised that these were likely to worsen. at that time his concerns focused primarily on the potential loss of herbivore species because of insufficient water for game. initially he prevailed upon government to act, but soon adopted an inward-looking approach, reasoning that the interests of irrigators and forestry were too pervasive and powerful to counteract. the resultant water for game programme was designed to support herbivore populations during drought through a network of reliable water points (such as drinking troughs supplied by boreholes) and the construction of weirs and sluices, which were to mitigate upstream impacts by ensuring the adequate flow of perennial rivers during droughts (pienaar 1970). in total, some 400 boreholes, dams, sluices and weirs were built, but as gaylard et al. (2003) note, the intended benefits did not materialise. as prudent as this policy may have seemed at the time, it was detrimental in two respects. firstly, the numbers of game, especially of rare antelope like roan, did not improve in the longer run – probably because of the competition with escalating zebra populations – and herbivore numbers continued to fluctuate (see e.g. owen-smith & ogutu 2003). secondly, the use of surface water influenced faunal, and hence vegetation, distribution patterns at multiple scales, because the water point distribution ensured that the majority of the land was within 5 km of a permanent water source (gaylard et al. 2003). although, with the benefit of hindsight, it is easy to see the shortcomings of this policy, it certainly was in keeping with the dominant management approaches of the time. all strongly interventionist in nature, the strategies were designed to reduce variability and unpredictability (mabunda et al. 2003). it is only in the past decade or so that new thinking, with heterogeneity at the forefront, has really come into its own. a detailed account is beyond the scope of this paper but it is instructive to note the breadth of this thinking detailed in the kruger experience: ecology and management of savanna heterogeneity (du toit, rogers et al. 2003). small sections of rivers and catchments in the kruger national park if the length of rivers or area of catchments that fall within the park are examined (figure 3), the linkages and vulnerability of the park to external influences are starkly apparent, with almost all the associated catchment areas falling largely outside of the knp boundaries. the most contained system is the sabie, which is relatively short, with 110 km of its total length of 190 km falling within the park. on the other hand, in the olifants, a severely degraded system, only a 100-km-long stretch of its total length of 840 km (or 11%) falls within the knp, accounting for a mere 8% of the catchment area. only 25% of each of the catchments of the crocodile and letaba rivers fall within the park, and some 36% and 18% of the river lengths, respectively. although 61% of the catchment area of the luvuvhu falls within the park, this accounts for only 34% of the river length. moreover, all the major river systems flowing eastwards through the knp ultimately feed mozambique (figure 1) and under international obligations south africa is required to honour certain flows through to that country. the knp thus sits between two realities: on the one hand, it is the victim of upstream use and abuse and, on the other, it acts as a buffer for mozambique downstream, through partly securing the water demands which will flow on into that country. figure 3: a comparison of (a) total catchment areas and (b) river lengths of the five perennial river systems found within the knp (from pollard, s.r. & du toit, d., 2007, ‘guidelines for strategic adaptive management: experiences from managing the rivers of the kruger national park, south africa’, iucn/ unep/gef project no. gf/ 271303-4679, ecosystems, protected areas and people project, planning and managing protected areas for global change). institutional arrangements against this precarious situation, we explore the response of the knp, which, as for the examples of fire and borehole provision, has transformed over time. what is distinctive about rivers is that owing to their fugitive nature, knp management was forced to look beyond the park borders for potential solutions (venter et al. 2008) and to devise monitoring and management responses that were embedded in a wider socio-political landscape. the knp has influenced institutional arrangements (biggs et al. 2008) and has undertaken, or at times initiated, engagement in wider water management actions such as catchment strategy development, international agreement revision, water quality monitoring and even legal action. mitigatory actions taken with respect to, for instance, maintaining the flow of the sabie during the 1992 drought (venter & deacon 1995) and in the letaba river (pollard & du toit 2008) would otherwise not have succeeded. indeed, the role of the knp as ‘watchdog’ (the first agency to alert water managers and society in general of a river problem) has been highlighted by pollard and du toit (2008) as essential to functional feedback loops. in the political climate that has prevailed since 1994, stakeholder involvement, transparency and accountability are regarded as key tools for achieving equity and sustainability. this means that the knp can no longer operate as a conservation island, because such policies compel it both to be involved and to partake in wider stakeholder discussions within its expanded, albeit ‘informal’, borders for water resources negotiations. the national water act (1998) outlines institutional arrangements for the management of water through the catchment management agencies operative for 19 water management areas (wmas). the knp straddles three wmas: • the inkomati (incorporating the sabie and crocodile rivers within the inkomati river basin, an international drainage basin shared by south africa, swaziland and mozambique) • the olifants • the luvuvhu–letaba. both the olifants and luvuvhu–letaba systems form part of the limpopo basin in mozambique. although only the inkomati wma has been gazetted thus far, it has set a precedent in that a seat for conservation is reserved in the composition of the board. this places the onus for participation on the conservation sector (and especially the knp) and affords a much stronger voice than in the past. the formal adaptive response: developing a strategy to respond to the challenges posed by declining river system integrity (back to top) the knp rivers research programme and adaptive management until the late 1980s, river management per se was not explicit as a park objective other than park authorities asking the (then) department of water affairs and forestry (dwaf) for special releases from upstream dams. nonetheless, by the late 1980s the development of water quality guidelines seemed to signal a resurgence of belief that the knp could exert constructive pressure on external agencies above ad hoc requests for releases, for instance, from the tzaneen dam. additionally, dwaf announced its intentions to allocate water for environmental flows in rivers as it became obvious that if demands were unchecked, the integrity of rivers would be threatened (dwaf 2004). despite these intentions, estimates were limited to preliminary calculations and based on absolute amounts of water. indeed, the first formal recognition of water for instream flow needs for south african rivers was by roberts (1983), who used an allocation for ‘conservation’ of 11% of the country’s mean annual runoff (later modified to 8% of the exploitable water resources by jezewski and roberts [1986]). roberts acknowledged that this figure was simplistic in that it was based on coarse, countrywide estimates of water for estuaries, lakes and nature reserves. as such, it could not be used for individual rivers (see also breen et al. [1994]), but nonetheless provided the catalyst for future work. researchers contested this figure and pointed to the paucity of understanding regarding lowveld river systems as a major challenge to management for sustainability of the rivers. ultimately a much more sophisticated and ground-breaking approach for calculating riverine water requirements was developed by a south african team (king et al. 2000) and prototyped through the knprrp and other research initiatives in south africa (see later). by 1988 the knprrp was conceptualised and initiated as a co-operative undertaking by managers or resource users, funding agencies and researchers (breen, dent et al. 2000). the knprrp consisted of three phases (appendix 1). phase i (1989–1993), which ran for four years, was largely limited to scientific research. the focus was on a range of research topics relating to environmental water requirements but was unstructured in detail (o’keeffe & coetzee 1996). moreover, managers within the knp were not convinced that, in practical terms, the research outputs supported the continuing management crises that they experienced (h. biggs, pers. comm.). in 1991, the newly constructed zoeknog dam in the upper sand catchment collapsed, delivering sediment for weeks into the river, a phenomenon that persisted through the knp and into mozambique (weeks, pollard et al. 1992). such a patent demonstration of undesirable consequences of poor design or construction compelled the park to respond through radio interviews and public platforms. it had not been customary before this for the park or their associated researchers to respond, signalling the start of a more public voice for the knp. it again highlighted the need for directed research that could support managers in their response to short-term crises. other research in the programme examined the potential fragility of the system, such as the effects on fish of being confined to shrinking pools (pollard, weeks et al. 1994) and vegetation changes associated with reed-bed colonisation (carter & rogers 1989). a comprehensive review of phase i recommended a second phase (1994–1996), with greater emphasis on predictive capabilities and management action, which was to be more intimately linked to a decision making system. it was during this phase that collaboration between managers and researchers improved with some co-learning. researchers began responding more explicitly to short-term crises experienced by managers, and managers benefited from the longer-term view provided by researchers. the political transformations that accompanied democratic transitions in 1994 were also major drivers for change, creating opportunity for far more effective international engagement. in 1995 the knprrp hosted an international conference on integrated catchment management in skukuza, a concept which was receiving increasing attention within the dwaf itself. this served to focus interests on holistic water resources management and, interestingly, raised the profile of international issues associated with water sharing across country borders. at about the same time, research interest in complexity theory and adaptive management within natural resource management started to grow. these ideas arose as a critique of approaches based on averages and the propensity to view nature as balanced, linear and predictable. variability, in fact, was highlighted as the key characteristic of semi-arid systems (davies, o’keeffe et al. 1995). even where ranges were recognised (e.g. introducing a variation of between 7000 and 9000 in knp elephant numbers), it was still not appreciated that savanna ecosystems need more extremes than these slight fluctuations to build resilience. this paradigm suggests that the recognition of variation and extreme events are fundamental for biodiversity management. this idea was central to the determination of environmental flows where variation in flow regimen was seen as a key driver of the system. the building-block approach (king et al. 2000) introduced the concept of incorporating freshes (small and intermediate floods) into a flow regimen, which were seen as essential linkages to certain key biotic or abiotic events such as spawning or sediment flushing. other concerns at the time centered on the entrenched and ‘command-and-control’ nature of management within the knp (biggs & rogers 2003; du toit, rogers et al. 2003). the imperative of political transformation necessitated change from one of an insular approach of managing the park, separate from its neighbours, to one which attempts to embed the knp within the socio-economic landscape and encourages wider participation, transparency and public ownership. moreover, the entrenched science–management activities such as monitoring were aligned with a facilitated, ‘learning-by-doing’ approach. the conservation of protected savannas in africa has been dominated by a focus on charismatic species and, as mentioned, influenced by stable-state concepts such as carrying capacity, with less emphatic regard for scale or the inherent dynamics of ecosystems. the previous approach has been challenged for its failure to embrace spatial heterogeneity and flux in ecosystems and for not always recognising a fuller array of compositional, structural and functional elements of biodiversity and ecosystems (noss 1990). indeed, a recent publication centred on the theme of heterogeneity in the knp (du toit et al. 2003) bears testimony to this fundamental shift in thinking. this raised a number of questions and challenges for the research and management community. firstly, what research was needed to elucidate the important characteristics of heterogeneity? secondly, how was management to embrace such variability and flux as the norm and when would the ‘variability norm’ be unacceptably exceeded? as noted by rogers (2005), strategic adaptive management (sam), and its associated objectives hierarchy, is one of the few recognised models for managing uncertainty in interactive social and ecological systems, whilst still aiming purposefully at a carefully articulated (but assumed to be shifting) desired state. as explained, river management was in crisis during the early 1990s and despite a vigorous initiative on the part of the knprrp and a few sanparks associates, most managers in the knp had not internalised that river management was an explicit part of their brief. interestingly, another important co-driver of the change in knp management was the ‘impasse’ on elephant culling, which came to a head at about the same time as the knprrp underwent a major reorientation (freitag et al. in review). heated public debate and scrutiny called for reforms to the culling programme and a moratorium on culling was introduced (van aarde, whyte et al. 1999). this essentially set the scene for other programmes to be influenced by the thinking that had developed in the knprrp. notably, a conference to discuss elephant management was held in skukuza and this provided an opportunity for participants to examine progress that had been made within the knp management framework. the key conclusions were that (1) the vision and objectives cascading from this vision needed to be improved and (2) elephants need to be managed as part of an ecosystem (braack 1997a). this led to the revision of the entire knp management plan (braack 1997a, b), starting with a visioning exercise of which the learnings and elements were already available through the knprrp. under the theme of accountability, the knp had to go public with its objectives. at the time, the knprrp was in the process of exploring and prototyping the concept of defining and operationalising the desired future state (dfs) of rivers (rogers & bestbier 1997). through collaborative efforts between the knp management and the knprrp, the application of this concept was explored for use beyond river management alone. much of the philosophy behind the dfs is that, as a public participation process which arrives at a joint agreement, much of the potential conflict can be reduced. after clear objectives were set as an objectives hierarchy, questions arose as to what needed to be monitored to achieve these objectives. a large collaborative meeting between managers and researchers in the knp heralded the start of measurable endpoints, known as tpcs (see, for example, braack et al. [1997], mcloughlin wrc 2011. and pollard and du toit [2007]). as described elsewhere, these tpcs are intimately embedded in an adaptive management framework. critically, they are set against the background of complex systems, representing spatio-temporal flux, often with lower and upper limits (see mcloughlin et al. [2011] for a comprehensive review of river-related tpcs). the third phase of the knprrp (1998–2000) was designed to enable the completion of first-generation procedures and technologies to support sam of rivers and to promote corrective action through the participation of stakeholders, especially those who had previously been marginalised. the need for a more holistic approach also prompted creative thinking around the issue of integrated catchment management and the role of sam in this regard. it could be argued that, given the leadership and close relationship between key individuals in the knprrp, dwaf and the water research commission (wrc), who funds water research in south africa, a strong learning alliance was formed, albeit informal. many of the ideas emerging from the knprrp and the process of water law reform were echoed in wrc research reports and were mutually reinforcing. for example, today notions of sam are embedded within certain strategic documents and guidelines of dwaf. moreover, the approach is now firmly embedded in scientific services within sanparks on an ongoing basis. although not named as such, a new phase of work was initiated in 2007 (after being discussed for several years) to build on that of the knprrp. it was driven by scientists’ and managers’ questions as to the apparent lack of improvement in the status of the rivers despite the advent of the national water act. the intention was to deepen the understanding surrounding the casues for the lack of improvement. about a year later the knp also recognised the need to strengthen its own adaptive management of the rivers and also initiated an associated project, both of which will now be discussed. a new phase of research: linking outputs to management the knprrp was followed by a hiatus in programmatic river research until the conceptualisation of the two initiatives, both quite distinct from the earlier rivers programme, and both strongly focused on action research and adaptive management processes. one of these, introduced earlier as the sri (pollard & du toit 2008), focused on understanding the factors that enable or constrain meeting the commitment to the ecological reserve in six river systems flowing through the knp (luvuvhu, letaba, olifants, sabie–sand, crocodile and komati). the intention was to build supportive programmes in phase ii due to commence in 2010. the other closely linked project aimed to consolidate the sam process for freshwater management in the park, mainly by operationalising the tpcs through effective science–management links (biggs et al. 2003; 2008; 2010). primarily as a result of findings from the sri, which identified case situations that were amenable to effective study, the current focus is on the letaba and crocodile rivers. in both cases there is strong evidence of feedback loops between key role players. feedback loops and self-organisation are considered to be essential components of resilient systems and adaptive management (biggs & rogers 2003a; holling 2001; holling & gunderson 2002). feedback is the basis for learning in a reflexive system. where systems often fail is where one or more of these steps fail such as in cases where the learning is not passed on or is passed to an inappropriate body. when functional, these loops set up a self-organising system that is responsive to change. as recognition for this approach grows, so does the interest in what makes the feedbacks work (see pollard et al. 2008). in the letaba catchment, for example, a number of key feedback loops of self-organisation and self-regulation are evident (figure 4). the knp monitors flows against the reserve requirements (which have until recently been static; that is, not actively dynamic in line with current exact rainfall) and, on detecting problems, the water affairs manager (who manages the tzaneen dam), in turn, alerts the groot letaba water users association to curtail use. they, in turn, inform users of curtailment rules and monitor adherence. although not always popular, the regulatory system is respected and adhered to by the members. there are a number of causal factors behind the success of these two loops, including the requirements of the law (the reserve), the availability of benchmarks against which to monitor (the reserve), the presence of a ‘watchdog’ (the knp in this case), the responsiveness of the manager and users, and the ability to self-organise. whilst an in-depth analysis of these is beyond the scope of this paper and is examined by pollard and du toit (in press), a key point is that the ‘watchdog’ role, which is as important as any of the other roles, is often overlooked and hence needs to be recognised as critical. the sam project is now refining, together with users, an adaptive monitoring management system. the essence of this system is that there are different levels of concern related to the status of a resource in question (e.g. river flow) and hence different management actions linked to each. the severity of the ‘worry level’ is given via an indicator or tpc, which is collaboratively determined (mcloughlin 2011). the important principle, therefore, is that there is an envelope of levels of concern – supported by a clear rationale – and each is linked to different management actions. the sri (phase i) has demonstrated that the requirements of the ecological reserve are not being met with regard to quantity in all six rivers, despite an improved policy environment and the initiation of integrated water resources management (pollard et al. 2010). this can, in part, be attributed to lags that are an inherent part of the process of reform in a complex environment; setting the reserve today does not mean that it will be met tomorrow. however, it is important to consider what makes certain delays unacceptable. in many cases, especially in the northern wmas, issues such as tardiness in authorisation, unlawful use, the lack of integration of water resources management and supply, weak monitoring and enforcement, and the dearth of skills and capacity all need to be addressed as a matter of urgency. in others, such as in the crocodile and komati rivers, recent advances in water resources management provide real possibilities for improvement. the knp has again been an important roleplayer in this regard, acting both as a catalyst for change and as a constructive stakeholder. figure 4: figure 4: functional feedback loops in the letaba catchment. feedback loops are evident when information that arises (discovered or learnt) as part of a management process is passed on to an (appropriate) body who takes (appropriate and effective) action and feeds this back. the self-regulation component is provided for by the water user association regulating its own members through internal agreements and practice. note the weak linkages to the regional and national department of water affairs. participation in wider catchment forums: the development of the catchment management strategy the commitment to manage water holistically is captured in the national water act, which requires that water resources are managed from a catchment perspective. ultimately, cmas will take over the management of the water resources, especially with regard to water allocation and protection of the resource. representation is secured through various structures such as catchment management committees or forums (cmcs or cmfs) and water user associations (wuas). in many cases, the knp has initiated forums that could be considered cmf precursors or prototypes. in the case of the crocodile river catchment, the knp has spearheaded the establishment of the crocodile river forum (crf) and for the sabie river, the sabie river working group, which started in 1991. today, the knp participates in fora that cover all major rivers entering the park and plays an important role in tabling its position and interests in water resource decisions. the relationship of the knp with and its influence on the agriculture or forestry sectors are also worth mentioning. an example is the sabie river working group, which managed to save the sabie river in the knp from a flow stoppage during the 1992 drought (biggs et al. 2010). the knp initiated this forum and was an active member but it was chaired by an irrigation farmer from the hazyview area for many years. another example is the marula weir, which was to be constructed in the crocodile river for irrigation purposes. the knp managed to stop the building of the weir even though the foundation had been started (venter et al. 1995). although the relationship between the irrigation farmers and the knp was strained as a result, the two parties have managed to build a good relationship subsequently as mutual understanding improved. more recently, the knp supported the efforts of a local initiative, namely the ‘save the sand’ programme, in advocating the withdrawal of poorly managed afforestation in the upper sand river catchment. the plantations, conceived as labour-creation schemes under the bantustan regime, covered excessively steep slopes, wetlands and riparian zones, which caused sediment problems and reduced base flows (figure 1). the support from the knp was not so much about actual effects (which were most heavily felt before the rivers enter the knp boundaries), but rather about the principle of wise use and management of natural resources. many more such examples are available in both peer-reviewed (e.g. kingsford et al. 2011; mcloughlin 2011) and grey literature (reports submitted to the wrc under project k5/1797). lessons that emerged from learning in action (back to top) this section deals with the emerging outcomes of the transformation described in this paper, which can now be reflected upon and used as guidance for a way forward. many important learnings in this field are already available, for example, descriptions and discussion of the: • transformation from a more closed to a more open style of management in the knp (venter et al. 1995) • general steps involved in sam and the related progress (kingsford et al. 2011) • spread of sam from a rivers application to a wide range of domains, not only in the knp but also across general conservation applications in south africa (freitag et al. in review) • mechanics of actual operationalisation of feedbacks built around thresholds (mcloughlin 2011). this paper has taken a more direct look at the philosophical and paradigmatic changes, and styles of management and research, that have characterised the transformation we describe. the overview considered the fuller range of initiatives relevant to water resource management in the park and its surrounds, both before and after the inception of explicit or formal adaptive management. the particular lessons elucidated by this review thus overlap with several lessons from each of the studies described, but also reinforce or complement those lessons with additional value. the observation that the sam approach has been widely accepted within the knp and that no fundamental alternatives for river management have appeared to date, may mean that the knp and other active collaborators are beginning to understand the complexity required to broker decisions effectively on a continuing (dynamic) basis. the fact that there have been ongoing structured research programmes on rivers seems to imply that, at least in the context of the knp, active levels of research involvement may be a prerequisite for coping in a fast-changing world with difficult resource management issues. the process of adopting sam was, as described earlier, a process of recursive action over time. although the sam approach began mainly with biophysical aspects within the park, over time, the recognition of wider socio-ecological systems (initially catchments) became central, with more active systemic connections being realised. this meant that, initially, management procedures became more inclusive of issues as they emerged and ultimately more complicated. there came a point where managers were overwhelmed, which resulted in a retraction to requisite simplicities (holling & gunderson 2002), so that the management process did not become untenable. what is the overall meaning of findings of this particular overview for the knp and for the wider community who are engaged in sam of rivers? the main achievement for the knp has been the development of a new way for approaching its conservation mandate based on complexity principles. this led to the emergence of a management framework over nearly 15 years (detailed in pollard and du toit [2007] and described earlier). the sam framework, although developed through a focus on rivers, can be embraced for the management of ecosystems as wholes. in summary, the framework requires that management be directed towards achieving a desired state (biggs & rogers 2003; breen at al. 2000). indeed, this has fundamentally re-orientated the management of the knp, staff and resource allocations. as explained earlier, once this higher-order statement had been debated and captured in a vision, it provided the basis for the development of objectives and endpoints that could be readily traced back to the vision. this process has allowed a much closer partnership to develop between researchers, managers and field staff, with a strong sense of buy-in and collective learning made possible (pollard & du toit 2005). an important shift in the management principle governing semi-arid savannas is that the desired future state is not a stable state but one that is based on a fundamental recognition of variability as an overarching characteristic to confer resilience. thus, judicious management is predicated on understanding the underlying ecosystem drivers and characteristics of the system in question. moreover, since river systems are dynamic and in a continual state of flux, it is necessary to monitor conditions and to revisit management objectives. system dynamics need to be understood in the broader context of events both inside and outside of the protected area. adopting the sam framework, with its key features being a clear vision informed by stakeholder involvement, an objectives hierarchy, a consideration of management options, an apparatus of tpcs and a reflective evaluation process consisting of feedback loops, has been challenging but important for knp staff. pollard and du toit (2007) noted that the collaborative role of researchers and management in developing tpcs and ensuring they are met has been cited as a powerful motivation for monitoring staff, such as rangers and wardens, who then become a key link in the iterative sam cycle. the value of involving field staff in setting management objectives cannot be underestimated with regard to developing commitment and buy-in. the tpcs are hypotheses and hence the tpcs and the associated ‘desired state’, should be audited and refined in a reflexive manner (mcloughlin 2011; pollard & du toit 2006). knowledge management is a challenge that needs to be addressed. biggs and rogers (2003) point out that after a tpc has been tabled, several unpredictable threads of information tend to emerge as implementation proceeds. these threads may or may not be documented at the appropriate level of quality (i.e. everything is taken to be equally relevant). these authors recommend a continual ‘roping together’ of the information so that the organisation benefits as a whole, thus averting disparate and isolated approaches. the sam approach is likely to generate a wealth of field data that need to be recorded, captured and made accessible. (mcloughlin 2011; pollard & du toit 2006). this is seen as one of the challenges for the knp. at present, the park is developing a knowledge management system based on a geographic information system as well as non-spatial databases. the intention is to draw science and management together by putting data to productive use rather than archiving for historical purposes only. once the challenging aspects of knowledge management have been negotiated they can lead to the need for shared learning. here the knp has experimented with the formation of ‘communities of practice’ (lave & wenger 1991) from, initially, a core of enthusiasts whose task it is to continually rework and improve the sam system and make it more accessible for use by others. experience shows that there is a need for programmes run by the knp to be integrated so that, by drawing on a wider variety of specialists and practitioner experiences, more realistic tpcs can be set in the future (pollard & du toit 2006). nonetheless, lessons for integrating new concepts, such as ecosystems services and social ecology, with more traditional approaches are yet to be learned. pollard and du toit (2008) argue that the legislative environment for water resources management and the approach of integrated water resources management afford a particularly strong basis and coherent currency for the adaptive management of river systems. the approach in the knp thus complements – and puts into practice – the spirit and intent of the national water act. in the case of river management, an additional challenge has been to broaden horizons and deal with the realities of conflicting drivers and objectives. river systems are common-property resources (pollard & cousins 2008). in south africa, there is no private ownership of water and flow through a portion of land does not confer inalienable rights on that land owner. moreover, because demand is viewed from a catchment-scale perspective of the total water resources, there will inevitably be tradeoffs and compromises in working towards more equitable and sustainable configurations for catchments (pollard & du toit 2008). these two factors necessitate that stakeholders, including protected area staff, participate in water resources management where different interests and demands on the water resources are used to negotiate water sharing. fortunately for the knp, the new legislative environment has provided strong support for the concept of sustainability through the provision of the reserve, which not only provides a benchmark for monitoring, but also carries legislative ‘clout’, strengthening the knp’s position as ‘watchdog’ (pollard & du toit 2008). this is critical given that infringements of the ecological reserve are evident in all rivers flowing through the park (pollard et al. 2010). importantly, the knp staff do not only monitor the rivers, but link outputs clearly to different actions according to the severity of the infringement (mcloughlin 2011), the transparency of which is important for monitoring staff (pollard & du toit 2006). although the systems are still being strengthened and successful responsive action nonetheless varies, the basis for building feedback loops is in place. indeed, as mentioned earlier, these feedbacks are essential for adaptive management, for without these, learning cannot happen (mcloughlin 2011; pollard & du toit 2006; pollard & du toit in press). conclusion (back to top) in conclusion, it is important to remember that adaptive management is not an end in itself, but a process that evolves as new learnings are brought to bear. as a result of the challenges confronted in addressing changes in rivers, the knp has charted new ground in management, research and outreach. the approach embraces the challenge of managing a sensitive, complex system in a context where uncertainty is always an underlying factor. it encourages the ‘first bold step forward’ where ‘implementation paralysis’ can often hamper decision making. by using the best available information to set tpcs, sam monitors trends and then demands reflection on collaboratively defined goals before jointly agreed action is initiated. the collaborative nature of implementing the sam system forges a partnership between science and management – an approach that is seen as a way forward for parks, conservation and science (folke, carpenter et al. 2002; van wilgen & biggs 2010). equally important is that such thinking has an institutional home for its eventual mainstreaming as management discourse, which the knp may well provide. references (back to top) biggs, h. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs, the kruger experience: ecology and management of savanna heterogeneity, pp. 59−80, island press, washington dc. biggs, h.c., breen, c.m. & palmer, c.g., 2008, ‘engaging a window of opportunity: synchronicity between a regional river conservation initiative and broader water law reform in south africa’, international journal of water resources development 24(3), 329−343. biggs, r., westley, f.r. & carpenter, s.r., 2010, ‘navigating the back loop: fostering social innovation and transformation in ecosystem management’, ecology and society 15(2), 9, viewed on 15 january 2011, from http://www.ecologyandsociety.org/vol15/iss2/art9/ braack, l., 1997a, a revision of parts of the management plan for the kruger national park. vol. vii: an objectives hierarchy for the kruger national park, south african national parks, skukuza. braack, l., 1997b, a revision of parts of the management plan for the kruger national park. vol. viii: policy proposals regarding issues relating to biodiversity maintenance, maintenance of wilderness qualities, and provision of human benefits, south african national parks, skukuza. breen, c., dent, m., jaganyi, j., madikizela, b., maganbeharie, j., ndlovu, j., et al., 2000, ‘the kruger national park rivers research programme’, final report, water research commission, pretoria. breen, c., quinn, n., deacon, a., 1994, ‘a description of the kruger park rivers research programme’, second phase: programme description: pp. 43. breen, c.m., bestbier, r., et al., 1997, ‘integrating socio-economic and governance systems with ecological knowledge of structure and function of riparian system. the ecology and management of riparian corridors in southern africa’, proc. international workshop, kruger national park, south africa. carruthers, j., 1995, the kruger national park: a social and political history, university of natal press, pietermaritzburg. chunnett, f., 1990, ‘water resources planning of the sabie river catchment’, water research commision, 1(10). cumming, d.h.m. & ahead great limpopo tfca working group, 2004, ‘sustaining animal health and ecosystem services in large landscapes-2nd draft-concept for a programme to address wildlife, livestock and related human and ecosystem health issues in the greater limpopo trans-frontier conservation area’, 24 pp., from http://www.wcs-ahead.org/workinggrps_limpopo.html davies, b.r., o’keeffe, j.h. & snaddon, c.d., 1995, ‘river and stream ecosystems in southern africa. predictably unpredictable’, in c.e. cushing, k.w. cummins & g.w. minshall, river and stream ecosystems, elsevier press, new york. department of water affairs and forestry, 2004a, internal strategic perspective. inkomati wma, pretoria. department of water affairs and forestry, 2004b, internal strategic perspective. luvuvhu/letaba wma, pretoria. department of water affairs and forestry, 2004c, internal strategic perspective, olifants river wma, pretoria. department of water affairs and forestry, 2004d, national water resources strategy, pretoria, pretoria. department of water affairs and forestry, 2009, inkomati water availability assessment: water requirements, pretoria. dovie, d.b.k., shackleton, c.m. & witkowski, e.t.f., 2006, ‘valuation of communal area livestock benefits, rural livelihoods and related policy issues’, land use policy 23, 260−271. du toit, j.t., rogers, k.h. & biggs, h.c., (eds.), 2003, the kruger experience. ecology and management of savanna heterogeneity, island press, washington dc. dwaf. see department of water affairs and forestry. folke, c., carpenter, s., elmqvist, t., gunderson, l., holling, c.s., walker, b., et al., 2002, ‘resilience and sustainable development building adaptive capacity in a world of transformations’, ambio 31(5), 437−440. freitag, s., biggs, h. & breen, c.m., in review, ‘fifteen years of the spread and maturation of adaptive management in south african national parks: organisational learning in systems perspective’, in w. freimund, s. mccool & c.m. breen (eds.), engaging complexity in protected area management: challenging occam’s razor, university of kwazulu-natal press, pietermaritzburg, south africa. gaylard, a., owen-smith, n. & redfern, j., 2003, ‘surface water availability: implications for heterogeneity and ecosystems processes’, in j.t. du toit, k.h. rogers & h.c. biggs, the kruger experience. ecology and management of savanna heterogeneity, pp. 171−188, island press, washington dc. holling, c.s. & gunderson, l.h., 2002, ‘resilience and adaptive cycles’, in l.h. gunderson & c.s. holling, panarchy: understanding transformations in human and natural systems, pp. 25−62, island press, washington dc. jezewski, j. & roberts c.p.r., 1986, ‘estuarine and lake freshwater requirements’, technical report tr129, department of water affairs. king, j.m., tharme, r.e. & de villers, m.s., 2000, ‘environmental flow assessments for rivers: manual for the building block methodology’, water resources commission report tt 131/100, pretoria, south africa. kingsford, r.t., biggs, h.c. & pollard, s.r., 2011, ‘strategic adaptive management in freshwater protected ares and their rivers’, biological conservation 144(4), 1194–1203. lave, j. & wenger, e., 1991, situated learning. legitimate peripheral participation, university of cambridge press, cambridge. mabunda, d., pienaar, d.j., et al., 2003, ‘the kruger national park: a century of management and research’, in j.t. du toit, k.h. rogers & h.c. biggs, the kruger experience. ecology and management of savanna heterogeneity, pp. 3−21, island press, washington dc. mcloughlin, c.a., deacon, d., sithole, h. & gyedu-ababio, t., 2011, ‘history, rationale, and lessons learned: thresholds of potential concern in kruger national park river adaptive management’, koedoe 53(2), art. #996, 27 pages. doi:10.4102/koedoe.v53i2.996 niehaus, i., 2001, witchcraft, power and politics. exploring the occult in the south african lowveld, pluto press, london. noss, r.f., 1990, ‘indicators of monitoring biodiversity: a hierarchical approach’, conservation biology 4, 355−364. o’keeffe, j. & coetzee, y., 1996, ‘status report of the kruger national park rivers research programme: a synthesis of results and assessment of progress to january 1996’, pretoria, water research commission: pp. 63. o’keeffe, j.h. & davies, b.r., 1991, ‘conservation and management of the rivers of the kruger national park: suggested methods for calculating instream flow needs’, aquat. conserv. mar. freshw. ecos. 1, 1−17. owen-smith, n. & ogutu, j., 2003, ‘rainfall influences on ungulate population dynamics’, in j.t. du toit, k.h. rogers & h.c. biggs, the kruger experience. ecology and management of savanna heterogeneity, pp. 422−446, island press, washington dc. pearce, f., 1992, dammed, bodley head, london. peel, m., 1999, ‘cattle grazing in the northern forestry area (dwaf): stocking densities guidelines’, report produced for the save the sand programme: pp. 31. pienaar, u.d.v., 1970, ‘water resources of the kruger park’, african wildlife 24, 180−191. pollard, s. & du toit, d., 2006, ‘recognizing heterogeneity and variability as key characteristics of savannah systems: the use of strategic adaptive management as an approach to river management within the kruger national park, south africa’, report for unep/gef project no. gf/2713-03-4679, ecosystems, protected areas and people project. pollard, s.r. & du toit, d., 2007, ‘guidelines for strategic adaptive management: experiences from managing the rivers of the kruger national park, south africa’, iucn/ unep/gef project no. gf/ 2713-03-4679, ecosystems, protected areas and people project, planning and managing protected areas for global change. pollard, s., du toit, d.r., reddy, y. & thlou, t., 2007, guidelines for the development of catchment management strategies: towards equity, efficiency and sustainability, department of water affairs and forestry, pretoria. pollard, s. & du toit, d., 2008, ‘the letaba catchment: contextual profile on factors that constrain or enable compliance with environmental flows’, shared river programme, draft report, project k5/1711. pollard, s. & du toit, d., 2009, ‘drawing environmental water allocations into the world of realpolitik: emerging experiences on achieving compliance with policy in the lowveld rivers, south africa. implementing environmental flow allocations’, port elizabeth, south africa. pollard, s. & du toit, d., in press, the importance of feedback loops in ensuring catchment resilience: an examination of six catchments in south africa, n.p. pollard, s., riddell, e., et al., 2010, compliance with the reserve: how do the lowveld rivers measure up?, report prepared for the water research commission: reserve assessment of lowveld rivers (del. 1), n.p. pollard, s., shackleton, c., et al., 2003, ‘beyond the fence: people and the lowveld landscape’, in j.t. du toit, k.h. rogers & h.c. biggs, the kruger experience. ecology and management of savanna heterogeneity, pp. 422−446, island press, washington dc. pollard, s.r., biggs, h. & du toit, d., 2008, ‘towards a socio-ecological systems view of the sand river catchment, south africa: an exploratory resilience analysis’, report to the water research commission, project k8/591, pretoria. pollard, s.r. & cousins, t., 2008, ‘towards integrating community-based governance of water resources with the statutory frameworks for integrated water resources management: a review of community–based governance of freshwater resources in four southern african countries to inform governance arrangements of communal wetlands’, water research commission report tt.328/08, pretoria, water research commission. pollard, s.r., mendiguren, j.c.p.d., et al., 1998, ‘save the sand phase i. feasibility study: the development of a proposal for a catchment plan for the sand river catchment’, pretoria, department of water affairs and forestry, department of agriculture and land affairs: 378. pollard, s.r. & du toit, d., 2008, ‘integrated water resources management in complex systems: how the catchment management strategies seek to achieve sustainability and equity in water resources in south africa’, water sa special edition iwrm 34(6), from http://www.wrc.org.za pollard, s.r., weeks, d.c., et al., 1994, ‘effects of the 1992 drought on the aquatic biota of the sabie and sand rivers’, in a pre-impoundment study of the sabie-sand river system, eastern transvaal, with special reference to predicted impacts on the kruger national park, vol. 2, pretoria, water research commission report: pp. 122. ramutsindela, m.f., 2002, ‘the perfect way to end a painful past? makuleke land deal in south africa’, geororum 33, 15−24. refugee, research, et al., 2002, ‘a park for the people? great limpopo transfrontier park – community consultation in coutada 16, mozambique’, preliminary research report (march), acornhoek, south africa. roberts, c.p.r., 1983, ‘environmental constraints of water resources developments’, proc s. afr. inst. civil engineers, n.p. rogers, k.h., 2005, ‘the real river management challenge: integrating scientists, stakeholders and service agencies’, river res. applic 22(2), 269–280. rogers, k.m. & bestbier, r., 1997, development of a protocol for the definition of the desired state of riverine systems in south africa, department of environmental affairs and tourism, pretoria. rsa, 1998, national water act, act 36 of 1998, republic of south africa government gazette, cape town, republic of south africa. russell, i.a. & rogers, k.h., 1988, ‘the distribution and composition of fish communities in major rivers of the kruger national park’, in s. kienzle and h. maaren, proceedings of the fourth south african national hydrological symposium, 20–22 november 1989, university of pretoria, university pretoria: 281–288. smith, r.e. & scott, d.f., 1992, ‘the effects of afforestation on low flows in various regions of south africa’, water sa 18(3), 185−194. stadler, j., 1994, ‘generational relationships in a lowveld village: questions of age, household and tradition’, msc thesis, university of witwatersrand. van aarde, r., whyte, i. & pimm, s., 1999, ‘culling and the dynamics of the kruger national park african elephant population’, animal conservation 2(4), 287−294. van wilgen, b.w. & biggs, h., 2010, ‘a critical assessment of adaptive ecosystem management in a large savanna protected area in south africa’, biological conservation, n.p. venter, f.j. & deacon, a.r., 1995, ‘managing rivers for conservation and ecotourism in the kruger national park’, water science and technology 32, 227−233. venter, f.j., gerber, f., deacon, a.r., viljoen, p.c. & zambatis, n., 1995, ‘ekologiese impakverslag: ‘n ondersoek na die voorgestelde maroelastuwal in die krokodilrivier’ [ecological impact report: an investigation of the proposed maroela weir in the crocodile river], report nr 1/95 scientific services, skukuza. venter, f.j., naiman, r.j., biggs, h.c. & pienaar, d.j., 2008, ‘the evolution of conservation management philosophy: science, environmental change and social adjustments in kruger national park’, ecosystems 11, 173−192. weeks, d.c., pollard, s.r., et al., 1992, ‘downstream effects on the aquatic biota of the sand river following the collapse of zoeknog dam’, report submitted to water research commission. water research commission, 2001, ‘state of the rivers report. crocodile, sabie-sand and olifants river systems’, pretoria, water research commission: pp. 40. table 1: timeline of the kruger national park rivers research programme outlining goals and focus (from breen, c., dent, m., jaganyi, j., madikizela, b., maganbeharie, j., ndlovu, j., et al., 2000, ‘the kruger national park rivers research programme’, final report, water research commission, pretoria; o’keeffe, j. & coetzee, y., 1996, ‘status report of the kruger national park rivers research programme: a synthesis of results and assessment of progress to january 1996’, pretoria, water research commission: pp. 63). book review bats and evolution book title: evolution’s chimera: bats and the marvel of evolutionary adaptation author: david jacobs isbn: 978-1775822127 publisher: uct press, 2016, r299.00* *book price at time of review review title: bats and evolution reviewer: m. brock fenton1 affiliation: 1department of biology, western university, canada corresponding author: brock fenton, bfenton@uwo.ca how to cite this book review: fenton, m.b., 2016, ‘bats and evolution’, koedoe 58(1), a1439. http://dx.doi.org/10.4102/koedoe.v58i1.1439 copyright notice: © 2016. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. in this short book, david jacobs uses bats to explore various aspects of evolution and evolutionary biology. in seven chapters, david goes from laying some ground work about evolution to considering how bats provide very good examples of evolution and evolutionary biology. evolution and bats are the focus of the book, which david addresses to students of biology. the book is not a detailed or encyclopaedic review of bats, rather a series of specific examples of how he uses bats to illustrate the fundamental aspects of evolutionary biology. it is important to keep the focus in mind, even though some of the examples explored in detail do not involve bats. in chapter 2, classification and diversification of bats are explored, from body size to flight and echolocation. chapter 3 deals mainly with the role that dna analyses have played in extending our understanding of the evolutionary relationships among bats. chapter 4 deals mainly with the possibility that competition (perhaps in the past) affected the diversification of bats. chapter 5 focuses on sound-mediated interactions between bats and their insect prey. chapter 6 considers the role that changes in echolocation signals might have played in bat evolution. chapter 7 approaches bats and conservation and finishes with an appeal for bats and their conservation. the book ends with a bibliography and an index, both of which are useful and appropriate. the book is well illustrated with a combination of drawings and photographs. together, the illustrations support the main themes of the book. some readers will be puzzled about topics not covered in the book, specifically the importance of white-nose syndrome (wns) and public health in the conservation of bats. wns has had a catastrophic impact on populations of some bats in the united states and canada where it is likely that some species of bats, including numerous common ones, will be extinct before too long. the role of bats in the spread of diseases that can afflict people (e.g. rabies, ebola and sars) may be real or imagined and surely strongly influence the public perception of bats. it is 10 years since dna barcode analysis was first used to document the details of bats’ insect prey. having this level of detailed information promises to revolutionise our view of bat ecology and behaviour. furthermore, details of prey selection offer the opportunity to move from rhetoric about competition (past or present) to actual details of how trophic interactions affect (or do not affect) communities of bats. i did not find mention of this approach in the book. i salute david for providing details of how bats can be used to advantage when studying evolution. bat biologists will find his examples interesting, showing once again that each of us has her or his own favourite stories about bat biology and evolution. article information authors: mduduzi ndlovu1,2 leigh combrink3 affiliations: 1school of animal, plant and environmental sciences, university of witwatersrand, south africa 2organisation for tropical studies, skukuza, south africa 3the endangered wildlife trust, gauteng, south africa correspondence to: mduduzi ndlovu email: mdu.ndlovu@wits.ac.za postal address: private bag x3, university of witwatersrand 2050, south africa dates: received: 22 apr. 2015 accepted: 30 aug. 2015 published: 27 nov. 2015 how to cite this article: ndlovu, m. & combrink, l., 2015, ‘feeding preferences of oxpeckers in kruger national park, south africa’, koedoe 57(1), art. #1316, 6 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1316 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. feeding preferences of oxpeckers in kruger national park, south africa in this original research... open access • abstract • introduction • materials and methods    • study area    • fieldwork    • data analysis • results • discussion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ oxpeckers reduce tick loads on ungulate hosts, but they are also known to feed on and exacerbate wounds. an understanding of the feeding behaviours and host preferences of these birds is important since they serve as agents of tick control on both domestic and wild ungulates. we conducted an observational study at two sites within the kruger national park in south africa, exploring the feeding preferences of both red-billed and yellow-billed oxpeckers. oxpeckers’ host preferences, body-location preferences on different hosts, prevalence of feeding and non-feeding behaviours, and frequency of tolerance versus rejection in different hosts were determined. it was found that yellow-billed oxpeckers had a smaller range of hosts – typically larger-sized ungulates – and that red-billed oxpeckers diversify to smaller-sized ungulate hosts when in competition with yellow-billed oxpeckers. body-location preferences were generally consistent across sites and across host species. tick feeding and other host-feeding behaviours (around the eyes, nose, mouth and ears, and anogenital areas) were fairly common. only six incidents of wound feeding, from a total of 855 observations, were recorded. tolerance by an ungulate host species was not related to oxpeckers’ host preferences, suggesting that other factors such as ungulate body size, tick species and tick stages on the host animal may play a significant role in the feeding preferences of oxpeckers. conservation implications: it is important to study oxpeckers’ behavioural feeding preferences so as to better understand their ecology and present distribution, and to determine where they can be reintroduced in future. reintroduction not only helps with the proliferation of oxpeckers, but also benefits ungulate hosts through ectoparasite removal and the subsequent control of tick-borne diseases. introduction top ↑ red-billed oxpeckers, buphagus erythrorhynchus, and yellow-billed oxpeckers, b. africanus, are known to take part in symbiotic relationships with african ungulates (dean & macdonald 1981, mundy 1983). oxpeckers reduce ectoparasite loads on host species and in return gain a major food source (mooring & mundy 1996). by removing ectoparasites, oxpeckers reduce host exposure to tick-borne diseases and the negative effects from ticks such as tick toxicosis, metabolic disturbances, anaemia and tick worry (zieger et al. 1998). in the wild, oxpeckers generally prefer to forage for ticks (and other ectoparasites) on large herbivorous mammals such as cape buffalo (syncerus caffer), giraffe (giraffa camelopardalis), hippopotamus (hippopotamus amphibius), kudu (tragelaphus strepsiceros), white rhinoceros (ceratotherium simum) and plains zebra (equus quagga burchelli) (see attwell 1966, grobler 1980, hustler 1987, koenig 1997, mooring & mundy 1996, stutterheim & stutterheim 1980). impala (aepyceros melampus) are the exception to this generalisation since they are considered to be relatively small ungulates, but appear to be highly preferred (e.g. grobler 1980, hart, hart & mooring 1990, stutterheim & stutterheim 1980). ticks, the birds main food source, are obtained by either pecking at the skin of the host or by sweeping the head along the host’s body whilst opening and closing the bill, a technique called ‘scissoring’ (attwell 1966; koenig 1997). these feeding techniques allow oxpeckers to remove almost all adult ticks from a host (bezuidenhout & stutterheim 1980). whilst ticks are the main food source for both oxpecker species, the birds are also known to feed on wound tissue, flies, mucous, saliva, earwax and blood (stutterheim 1981; weeks 2000). tick feeding is generally regarded as beneficial to the ungulate host (dean & macdonald 1981), whereas wound feeding can be detrimental as it prolongs the healing time of wounds and increases the chance of infection (mcelligott et al. 2004; weeks 1999). a host is more likely to tolerate an oxpecker performing a beneficial behaviour, and reject the bird that is wound feeding. accordingly, one would expect oxpeckers to feed more on ticks than on wounds. in south africa, both oxpecker species suffered population and range reductions during the first half of the 20th century, due to the treatment of livestock with poisonous acaricides, the over-hunting of large ungulates, and the rinderpest epidemic of 1896–1897 (mundy 1983; stutterheim 1982; stutterheim & brooke 1981). these factors significantly reduced the numbers of host species and ticks, which in turn caused a decline in oxpecker numbers and distribution. the remaining fragmented populations were restricted to protected areas and national parks (grobler 1980; stutterheim & brooke 1981). with the advent of new oxpecker-compatible ingredients in acaricides, the reintroduction of oxpeckers from national parks into areas within their historic range has been possible and remains a useful conservation tool for ensuring the long-term survival of this species in south africa. as part of conserving these birds, it is important to study oxpeckers’ behavioural preferences in their present natural habitat so as to better understand where they can be reintroduced in future. reintroduction not only helps with the proliferation of oxpeckers, but also benefits ungulate hosts through ectoparasite removal and the subsequent control of tick-borne diseases. research on the feeding ecology of oxpeckers has been carried out in various areas in east and southern africa. however, very few of these studies have actually explored interspecific feeding behaviours in sympatric cases (hustler 1987; koenig 1997; plantan 2009). yellow-billed oxpeckers have thicker bills and significantly heavier bodies compared to the slender-built red-billed oxpeckers (mundy 1983). in allopatric studies, both species appear to forage on similar host species (grobler 1980; grobler & charsley 1978; mooring & mundy 1996). however, given their differences in bill size and body mass, one would expect some level of interspecific competition, where an overlap of range distribution might occur. furthermore, previous results suggest that the relationship between oxpeckers and ungulates is primarily mutualistic (see nunn et al. 2011), but few studies have argued that the relationship may also be parasitic (mcelligott et al. 2004; plantan et al. 2013; weeks 1999). in this study, the feeding preferences of both red-billed and yellow-billed oxpeckers at two different locations (north versus south) within the kruger national park (knp) in south africa were examined. the differences in feeding behaviour between the species and between sites were evaluated. site differences were only considered for red-billed oxpeckers since they were found in both the north and south of knp, whereas yellow-billed oxpeckers were only sighted in the northern regions of the park. oxpeckers’ host preferences, body-location preferences on hosts, tolerance by host species and prevalence of feeding behaviours were recorded. materials and methods top ↑ study area this study was carried out between september 2013 and september 2014 at two sites within knp: skukuza (24°59′s, 31°46′e) and shingwedzi (23°6′s, 31°25′e). the survey area for each site covered a radius of approximately 50 km from each of these rest camps and data collection was done bi-annually (september 2013, february 2014 and september 2014). skukuza is located in the southern section of knp. the area sits on granitic rock and consists of slightly undulating plains. broad-leaved bushveld is found in the uplands and fine-leaved bushveld in the bottomlands. skukuza receives 500 mm – 750 mm of rainfall per year. shingwedzi is located in the northern regions of knp and consists of flat plains of broad-leaved shrubveld that are dominated by mopane (colophospermum mopane) woodlands. it receives 450 mm – 500 mm of rainfall per year and sits on basaltic rock (venter, scholes & eckhardt 2003). red-billed oxpeckers occur throughout knp, whilst yellow-billed oxpeckers are abundant in the north of the park (chittenden & whyte 2007). fieldwork data were collected early in the morning (07:00–10:00) and late in the afternoon (15:00–18:00), which coincides with oxpeckers’ peak feeding periods (plantan 2009). three game drive vehicles were used during each sampling period. each vehicle drove along non-overlapping routes at each study site area, looking for potential host species with or without oxpeckers. each vehicle carried between six and eight occupants. field observations were conducted using binoculars, and only oxpeckers observed within 200 m of the vehicle were included. observers in each vehicle recorded data using cybertracker software, accessed using smartphones. cybertracker enables the researcher to rapidly enter data electronically whilst still in the field. the observation techniques used were similar to that of grobler (1980), stutterheim and stutterheim (1980) and plantan (2009). each time a host species was encountered, data were collected on the following: (1) identity of the host species, (2) number of hosts available, (3) number of oxpeckers present, (4) oxpecker species, (5) age class of each oxpecker observed, (6) presence or absence of wounds on host, (7) location of oxpecker on host, (8) behaviour of each oxpecker, (9) host response (tolerance or rejection) and (10) gps coordinates and time of each observation. behaviours were divided into wound feeding behaviours, non-wound feeding behaviours (tick-, anogenital-, ear-, nose-, eyeand mouth feeding) and non-feeding behaviours (perching or preening). behavioural observations of birds were done within a 2 min period or until the bird was no longer visible or flew off the host. when estimating host numbers in instances where large numbers of animals were present and an accurate count was not possible, a probable underestimate of the total number was made. a host was considered to have a wound if the skin had a visible break, whether or not blood was present. oxpecker behaviour was considered to be rejected if the host animal actively tried to dislodge the bird through leg-stomping, nose-pushing, head-shaking, actively running away, etc. data analysis to determine oxpeckers’ host preferences, the method of stutterheim and stutterheim (1980) was used to obtain a preference index (pi) for each host species (number of birds or number of animals). chi-square tests were then used to compare preference index distributions between sites and between species. a similar analysis was also conducted to compare the results of the present study to historic data sets from stutterheim and stutterheim (1980) and grobler (1980). simple arithmetic was used to determine oxpeckers’ host-body-location preferences and the prevalence of different bird behaviours (i.e. non-feeding, tick feeding, wound feeding or other feeding) was also compared. the frequency of the hosts’ response to various oxpeckers’ behaviours (tolerance or rejection) was also explored. a linear regression model in rstudio (2013) was used to determine whether tolerance by a host species was related to the host preferences of oxpeckers. results top ↑ in skukuza, 6107 potential host animals and 329 red-billed oxpeckers were observed, with 511 behavioural instances recorded. in shingwedzi, 2254 potential hosts and 158 oxpeckers were observed, with 344 behavioural instances recorded. of the oxpeckers in shingwedzi, 120 were red-billed oxpeckers and only 38 were yellow-billed oxpeckers. overall, the most preferred host species for oxpeckers in knp was white rhinoceros (pi = 0.60) and giraffe (pi = 0.54). kudu and cape buffalo were also highly preferred, with pis of 0.18 and 0.17, respectively. oxpeckers were also present on hippopotamus, plains zebra and impala (figure 1). impala, whilst not highly preferred, were still important hosts due to their high abundance. more than 25% of all observed oxpeckers were found on impala, which also made up 74% of all observed host species. red-billed oxpeckers had different host animal preferences in skukuza compared to shingwedzi (figure 2). white rhinoceros, an important host animal in skukuza, was not recorded in shingwedzi. plains zebra was observed in both locations, but was only utilised as a host in shingwedzi. of the five host animals utilised in both areas, red-billed oxpeckers in shingwedzi showed a higher preference for kudu, hippopotamus and impala, compared to skukuza birds (x2(4) = 251.20, p < 0.001). yellow-billed oxpeckers had a smaller range of host animals than their red-billed counterparts (x2(5) = 30.59, p < 0.001). yellow-billed oxpeckers were only found on giraffe, cape buffalo and plains zebra, whilst red-billed oxpeckers in the same area were also found on kudu, hippopotamus and impala (figure 3). figure 1: host preferences of red-billed and yellow-billed oxpeckers from both skukuza and shingwedzi areas in the kruger national park. preference index is a ratio of the number of birds seen on a given host species divided by the number of hosts seen of that species. figure 2: a comparison of host preference indices for red-billed oxpeckers in the skukuza and shingwedzi areas of the kruger national park. figure 3: a comparison of host preference indices for red-billed and yellow-billed oxpeckers in the shingwedzi area of the kruger national park. a comparison of the present preference indices to those of grobler (1980) and stutterheim and stutterheim (1980) revealed differences in host preferences (table 1). oxpeckers in skukuza preferred white rhinoceros and, in contrast, was not a significant host animal in earlier studies. hippopotamus was not recorded as a host in the 1980 studies, but was a minor host in the findings of the present study. cape buffalo in skukuza was the preferred host, contrary to earlier findings from previous studies (grobler 1980) in the same region (x2(3) = 148.32, p < 0.001). compared to stutterheim and stutterheim (1980), data from shingwedzi (present study) indicate less preference for cape buffalo and giraffe, and more preference for impala, kudu and plains zebra (x2(4) = 83.26, p < 0.001). table 1: host preference indices from this study, compared to previous studies in the kruger national park. a total of 419 body-location preference instances were recorded. red-billed oxpeckers were most frequently observed on the back (41%, n = 170), head (20%, n = 83) and neck (16%, n = 65). the neck was particularly preferred on giraffe (40%, n = 25), whilst the flanks were also important for kudu (19%, n = 7). oxpeckers were also seen on the legs and anogenital regions of hosts (figure 4). figure 4: body-location preferences on the preferred hosts of red-billed oxpeckers in the skukuza and shingwedzi areas. a total of 386 unique behavioural incidences were recorded for red-billed oxpeckers at both sites. the most frequently recorded behaviour was non-feeding (44%, n = 169), but tick feeding (38%, n = 145) and other forms of host feeding (17%, n = 66) were also prevalent (figure 5). only 1.6% of all behavioural observations were of wound feeding (n = 6; red-billed oxpecker = 6, yellow-billed oxpecker = 0). yellow-billed oxpeckers were only seen feeding whilst on cape buffalo (n = 17); all observations of yellow-billed oxpeckers on other hosts were of non-feeding birds. figure 5: the frequency of red-billed oxpeckers’ behaviours on different ungulate host species in the skukuza and shingwedzi areas. a total of 80% of host animals observed throughout the study at both sites appeared to be tolerant of oxpeckers (table 2). the most tolerant host animals were giraffe, white rhinoceros and hippopotamus. cape buffalo, plains zebra, impala and kudu were slightly less tolerant. there was a non-significant correlation between the degree of tolerance of a host species and oxpeckers’ preference for that same host (r2 = 0.486, f = 4.723, p = 0.082). table 2: tolerance and rejection instances by host species in the skukuza and shingwedzi areas. discussion top ↑ the aim of the present study was to provide insight and understanding of both red-billed and yellow-billed oxpeckers’ feeding behaviours, particularly regarding host and body-location preferences. overall, red-billed and yellow-billed oxpeckers displayed a host preference for large ungulates (white rhinoceros, giraffe and cape buffalo). these findings are similar to previous studies (attwell 1966; grobler 1980; hustler 1987; stutterheim & stutterheim 1980). this preference could be a result of larger ungulates providing a greater surface area for ectoparasites, which are therefore able to carry larger tick loads (horak et al. 1983; koenig 1997; mooring & mundy 1996). generally, large ungulates have many ectoparasites, particularly ticks at various life cycle stages (grobler & charsley 1978; mooring & mundy 1996). large ungulates also have a limited ability to self-groom, which subsequently maintains, if not increases, their tick loads (mooring & mundy 1996). additionally, the three most preferred host species are often gregarious, travelling and feeding in large numbers, and therefore potentially increasing tick abundance, transmission and prevalence (koenig 1997). differences in host preferences were observed between populations of red-billed oxpecker in the shingwedzi and skukuza regions, with the shingwedzi population selecting for smaller ungulates (see figure 2). red-billed oxpeckers appear to be limited to medium and small-sized ungulates, such as kudu and impala, in the northern parts of knp. this is most likely because of a home range overlap with yellow-billed oxpeckers, resulting in interspecific competition (koenig 1997). the larger (in terms of body size) yellow-billed oxpecker is territorial and capable of outcompeting the smaller red-billed oxpecker (hall-martin 1987), permitting the former a preferential choice of ungulates. the study also shows that red-billed oxpeckers in the southern regions of knp utilised the preferred large ungulates in the absence of yellow-billed oxpeckers, whereas their preference shifted to smaller ungulates in the presence of yellow-billed oxpeckers. this further supports the notion of interspecific competition between red-billed and yellow-billed oxpeckers for access to larger ungulate hosts. contrary to the results from the present study, hustler (1987) and koenig (1997) in zimbabwe and kenya (respectively) did not find any differences in host ungulate preferences when both species occurred within the same geographic region. furthermore, koenig (1997) did not find any marked differences in the host species preferences of red-billed oxpeckers when comparing between areas of sympatry and areas of allopatry. perhaps these differences between the kenya study and the knp findings could be attributed to differences in ungulate densities between the two sites. one would assume that the kenya sites (masai mara reserve and lake nakuru national park) had a high abundance of large ungulates compared to knp, hence a marked host preference would only be apparent in lower ungulate densities where interspecific competition is unavoidable. however, that hypothesis cannot be tested without a measure of ungulate densities from all sites. surprisingly, the pi results in the present study differed from grobler (1980) and stutterheim and stutterheim (1980). the most plausible explanation is that these differences are related to changes in large ungulate numbers and distribution within knp in the past 30 years (chirima, owen-smith & erasmus 2012; sanparks 2012), coupled with a possible switch in oxpeckers’ host preferences in response to changing ungulate densities. optimal foraging strategy (pyke 1984) will predict that animals will concentrate on the most abundant and profitable food source. for example, there were fewer white rhinoceros and hippopotamus in knp in the 1980s compared to the present-day population (2014). it is therefore reasonable to conclude that as white rhinoceros and hippopotamus numbers increased, red-billed oxpeckers responded by selecting for these new abundant host species with potentially higher tick loads and less hair to hide the ticks. giraffe remained the most preferred host species in both the northern and southern regions of the park. this could also be a detection bias, given that it is possibly easier for flying birds to detect giraffe compared to other (shorter) species. nevertheless, it is reasonable to conclude that size plays a major role in oxpeckers’ host preferences (grobler 1980; grobler & charsley 1978; koenig 1997; mooring & mundy 1996; stutterheim 1982; stutterheim & brooke 1981). this is further supported by oxpeckers’ preference for white rhinoceros and hippopotamus (both large ungulates), recorded as host species in skukuza. interestingly, impala, an abundant, small-sized ungulate, was less preferred as a host species across studies. this surprising contradicts what has been reported by grobler (1980), stutterheim and stutterheim (1980) and hart et al. (1990). both red-billed and yellow-billed oxpeckers displayed a preference for the back and head regions of their respective host species. red-billed oxpeckers also preferred the necks of giraffe. these preferences coincide with body regions that are likely to exhibit higher tick abundance, given the degree of difficulty in tick-grooming access (mooring & mundy 1996; weeks 1999) and are often the areas that exhibit more wounds (pers. obs.). additionally, oxpeckers prefer feeding on the back regions of a host species since this is easily accessible and provides a stable perch (weeks 1999). the low prevalence of tick feeding by both oxpecker species on body regions that are easily self-groomed by the host species, such as the legs, further supports the possibility that oxpeckers show a preference for non-groomed body regions for their higher tick loads (koenig 1997; mooring & mundy 1996). the head is also preferred since it provides additional food resources other than ticks, i.e. saliva, mucus and earwax (stutterheim 1981). exceedingly low occurrences of wound feeding by red-billed oxpeckers and the absence of wound feeding in yellow-billed oxpeckers suggests that this feeding behaviour is not prevalent in knp as previously reported in cattle ranches in zimbabwe (weeks 2000). plantan (2009), in her study on both oxpecker species at shingwedzi in 2007, also found the prevalence of wound feeding behaviour to be very low (3.1% of 558 feeding observations). however, she did find that wound feeding was exhibited by yellow-billed oxpeckers more than red-billed oxpeckers. in the current study, there were instances where ungulates were observed with open wounds and yet oxpeckers did not tamper with the wounds, but rather continued tick feeding. this observation further suggests that oxpeckers pose no risk to free-roaming wildlife. large ungulates tolerated oxpeckers more so than medium-sized ungulates. this tolerance is inadvertently due to the reduced agility of large ungulates, providing a more stable perch and a greater feeding area (grobler 1980; hustler 1987; stutterheim & stutterheim 1980; weeks 1999). the preferential selection of larger hosts also indicates that reintroductions of oxpeckers should target areas with high tick loads and substantial, large ungulate populations of the host species that are preferred. however, conservationists should also consider that oxpeckers prefer certain tick species and tick stages (mooring & mundy 1996). therefore, even if a large ungulate has many ticks, this does not mean that the oxpecker will consume those ticks or utilise the host. however, the present study does provide additional insight and understanding of red-billed and yellow-billed oxpeckers’ feeding behaviours, particularly in knp and in terms of wild host species and body-location preferences. acknowledgements top ↑ the authors would like to thank the sanparks scientific services at knp, staff and students at the organization for tropical studies (ots) and the endangered wildlife trust for their support in this project. sanparks issued permits for this research project to ots south africa. competing interests the authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article. authors’ contributions m.n. (university of the witwatersrand) and l.c. (the endangered wildlife trust) were both responsible for the project design, data collection and management of the study. m.n. performed most of the data analysis and wrote the first draft of the article. l.c. made conceptual contributions to the article, did the literature review and revised the drafts. references top ↑ attwell, r.i.g., 1966, ‘oxpeckers and their associations with mammals in zambia’, puku 4, 17–48. bezuidenhout, j.d. & stutterheim, c.j., 1980, ‘a critical evaluation of the role played by the red-billed oxpecker buphagus erythrorhynchus in the biological control of ticks’, onderstepoort journal of veterinary research 55, 173–179. chirima, g.j., owen-smith, n. & erasmus, b.f.n., 2012, ‘changing distributions of larger ungulates in the kruger national park from ecological aerial survey data’, koedoe 54(1), art. #1009, 11 pages. http://dx.doi.org/10.4102/koedoe.v54i1.1009 chittenden, h. & whyte, i., 2007, roberts bird guide: kruger national park and adjacent lowveld, john voelcker bird book fund, cape town. dean, w.r.j. & macdonald, i.a.w., 1981, ‘a review of african birds feeding in association with mammals’, ostrich 52, 135–155. http://dx.doi.org/10.1080/00306525.1981.9633599 grobler, j.h., 1980, ‘host selection and species preference of the red-billed oxpecker buphagus erythrorhynchus in the kruger national park’, koedoe 23, 89–97. http://dx.doi.org/10.4102/koedoe.v23i1.637 grobler, j.h. & charsley, g.w., 1978, ’host preference of the yellow-billed oxpecker in the rhodes matopos national park, rhodesia’, south african journal of wildlife research 8, 169–170. hall-martin, a.j., 1987, ‘range expansion of the yellowbilled oxpecker buphagus africanus into the kruger national park, south africa’, koedoe 30, 121–132. http://dx.doi.org/10.4102/koedoe.v30i1.505 hart, b.l., hart, l.a. & mooring, m.s., 1990, ‘differential foraging of oxpeckers on impala in comparison with sympatric antelope species’, african journal of ecology 28, 240–249. http://dx.doi.org/10.1111/j.1365-2028.1990.tb01157.x horak, i.g., potgieter, f.t., walker, j.b., de vos, v. & boomker, j., 1983, ‘the ixodid tick burdens of various large ruminant species in south african nature reserves’, onderstepoort journal of veterinary research 50, 221–228. pmid: 6646664. hustler, k., 1987, ‘host preference of oxpeckers in the hwange national park, zimbabwe’, african journal of ecology 25(4), 241–245. http://dx.doi.org/10.1111/j.1365-2028.1987.tb01115.x koenig, w.d., 1997, ‘host preferences and behaviour of oxpeckers: co-existence of similar species in a fragmented landscape’, evolutionary ecology 11, 91–104. http://dx.doi.org/10.1023/a:1018439614008 mcelligott, a.g., maggini, i., hunziker, l. & könig, b., 2004, ‘interactions between red-billed oxpeckers and black rhinos in captivity’, zoo biology 23(4), 347–354. http://dx.doi.org/10.1002/zoo.20013 mooring, m.s. & mundy, p.j., 1996, ‘interactions between impala and oxpeckers at matobo national park, zimbabwe’, african journal of ecology 34, 54–65. http://dx.doi.org/10.1111/j.1365-2028.1996.tb00594.x mundy, p.j., 1983, ‘the oxpeckers of africa’, african wildlife 37, 111–116. nunn, c.l., ezenwa, v.o., arnold, c. & koenig, w.d., 2011, ‘mutualism or parasitism? using a phylogenetic approach to characterize the oxpecker-ungulate relationship’, evolution 65(5), 1297–1304. pmid: 21166791, http://dx.doi.org/10.1111/j.1558-5646.2010.01212.x plantan, t.b., 2009, ‘feeding behavior of wild and captive oxpeckers (buphagus spp.): a case of conditional mutualism’, phd thesis, university of miami, florida. plantan, t.b., howitt, m., kotze, a. & gaines, m.s., 2013, ‘feeding preferences of the red-billed oxpecker, buphagus erythrorhynchus: a parasitic mutualist?’, african journal of ecology 51, 325–336. http://dx.doi.org/10.1111/aje.12042 pyke, g.h., 1984, ‘optimal foraging theory: a critical review’, annual review of ecology and systematics 15, 523–575. http://dx.doi.org/10.1146/annurev.es.15.110184.002515 rstudio, version 0.98.1028, 2013, computer software, rstudio, inc., boston, ma. sanparks, 2012, ‘estimates for animal abundances in parks in the northern region’, south african national parks annual report 2011/2012, viewed 03 november 2014, from http://www.sanparks.co.za/assets/docs/general/annual-report-2012.pdf stutterheim, c.j., 1981, ‘the feeding behaviour of the redbilled oxpecker’, south african journal of zoology 16(4), 267–271. stutterheim, c.j., 1982, ‘past and present ecological distribution of the redbilled oxpecker (buphagus erythrorhynchus) in south africa’, south african journal of zoology 17(4), 190–196. http://dx.doi.org/10.1080/02541858.1982.11447802 stutterheim, c.j. & brooke, r.k., 1981, ‘past and present ecological distribution of the yellowbilled oxpecker in south africa’, south african journal of zoology 16(1), 44–49. stutterheim, c.j. & stutterheim, m., 1980, ‘evidence of an increase in a red-billed oxpecker population in the kruger national park’, south africa journal of zoology 15(4), 284. http://dx.doi.org/10.1080/02541858.1980.11447724 venter, f.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation pattern’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83–129, island press, washington, dc. weeks, p., 1999, ‘interactions between red-billed oxpeckers, buphagus erythrorhynchus, and domestic cattle, bos taurus, in zimbabwe’, animal behavior 58(6), 1253–1259. pmid: 10600147, http://dx.doi.org/10.1006/anbe.1999.1265 weeks, p., 2000, ‘red-billed oxpeckers: vampires or tickbirds?’, behavioral ecology 11(2), 154–160. http://dx.doi.org/10.1093/beheco/11.2.154 zieger, u., horak, i.g., cauldwell, a.e., uys, a.c. & bothma, j. du p., 1998, ‘the effect of chemical tick control on cattle on free-living ixodid ticks and on ticks parasitic on sympatric impala in the central province, zambia’, south african journal of wildlife research 28(1), 10–15. article information authors: tamara b. robinson1 cheruscha swart1 affiliations: 1department of botany and zoology, centre for invasion biology, stellenbosch university, south africa correspondence to: tamara robinson email: trobins@sun.ac.za postal address: private bag x1, matieland 7602, south africa dates: received: 05 aug. 2014 accepted: 23 sept. 2014 published: 25 mar. 2015 how to cite this article: robinson, t.b. & swart, c., 2015, ‘distribution and impact of the alien anemone sagartia ornata in the west coast national park’, koedoe 57(1), art. #1246, 8 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1246 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. distribution and impact of the alien anemone sagartia ornata in the west coast national park in this original research... open access • abstract • introduction • research method and design    • setting    • distribution and abundance of sagartia ornata    • coelenteron contents analysis    • community structure    • statistical analyses • results    • distribution and abundance of sagartia ornata    • coelenteron contents analysis    • community structure • discussion • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ sagartia ornata is an alien anemone that occurs intertidally within the west coast national park (wcnp). whilst baseline distributional data was gathered in 2001, the range and abundance of this alien has not been reassessed. the present study aimed to determine the current status and distribution of this anemone, to assess its diet so as to establish the role it may play as predator and to investigate its impact on sandy-shore communities. sagartia ornata was found to be restricted to the wcnp, where it occurred in densities of up to 508 ± 218 individuals per m2. within the park the distribution of this anemone had changed. populations were recorded in nanozostera capensis seagrass beds for the first time and this alien was absent from two areas in which it had previously occurred. diet analysis revealed indigenous polychaetes and amphipods as the dominant prey items consumed by s. ornata. this alien was found to significantly alter sandy-shore community structure, with differences caused primarily by increases in the abundance and biomass of the tanaid anatanais gracilis and the polychaete orbinia angrapequensis. additionally, invaded areas supported significantly greater invertebrate diversity, density and biomass. it is concluded that whilst this anemone negatively affects native biota, its current dependence on restricted habitats precludes widespread impacts with the park. conservation implications: with regard to conservation implications, this invasion should be routinely monitored outside the wcnp as in its native range s. ornata occurs on rocky shores and kelp holdfasts, suggesting a potential for spread along the west coast of south africa. introduction top ↑ the human-mediated spread of marine species from their natural ranges to new habitats is occurring with increasing frequency (seebens, gastner & blasius 2013) and resulting invasions have been reported from most regions (molnar et al. 2008). these invasions result primarily from shipping, the main vector being ship hull fouling (hewitt, gollasch & minchin 2009; mead et al. 2011a). other shipping-related vectors include ballast water (klein et al. 2010) and sediment retained in ballast tanks (briski, bailey & macisaac 2011). the increase in the number of ships and ballast tank size (carlton 1996), accompanied by an increase in transoceanic travel (ruiz et al. 1997) over the last century, has led to an increase in the introduction and rate of establishment of invasive marine species around the world (wonham et al. 2000). this is cause for concern as these alien species can pose a threat to indigenous biota (gurevitch & padilla 2004; molnar et al. 2008; strain & johnson 2013) and can have negative economic impacts (cinar 2013; pimentel, zuniga & morrison 2005). in south africa, 90 introduced and 39 cryptogenic species have been recognised to date (bolton, andreakis & anderson 2011; clark & griffiths 2012; mead et al. 2011b; peters et al. 2014), although this is likely to be an underestimation as alien species are continuously being introduced into the country. most introductions are recorded along the cool-temperate west coast (mead et al. 2011a); most occurring within harbours, a moderate number on rocky shores and in estuaries and very few offshore (mead et al. 2011a). following global trends, most species alien to south african waters have been introduced via hull fouling and ballast water (mead et al. 2013). the west coast national park (wcnp) is situated in the saldanha bay system on the cold west coast of south africa and includes langebaan lagoon (robinson, griffiths & kruger 2004). it was established in 1985 and is the only national park for the protection and conservation of marine species north of the cape peninsula (robinson et al. 2004). the wcnp faces numerous threats, including increased pollution within the bay owing to industrial development and urbanisation (kruger et al. 2005), expanding mariculture (olivier, heinecken & jackson 2013) and the introduction of marine alien species associated with the port (robinson et al. 2004). presently more alien marine species occur within the saldanha bay–langebaan lagoon system than in any other location along the south african coast (mead et al. 2011a). the alien anemone sagartia ornata was first recorded along langebaan lagoon's western shoreline in 2001 in the upper intertidal zone (acuña, excoffon & griffiths 2004). here, s. ornata was found intertidally on sandy shores partially covering the underlying rocks and in spartina maritima beds (robinson et al. 2004), where its densities reached up to 426 ± 81 (s.d.) individuals per m2 (robinson et al. 2005). sagartia ornata's introduction to the lagoon is thought to have occurred unintentionally through shipping via the harbour of saldanha bay (robinson et al. 2004). it is most likely that the introduction is recent, as extensive annual sampling of the lagoon failed to detect this anemone until 2001 (acuña et al. 2004). this species originates from western europe, the mediterranean and britain (robinson et al. 2004). there it is found widely occurring on kelp holdfasts and in crevices on rocky shores (gibson, hextall & rogers 2001). thus, the south african west coast, known for its cold water, rocky shores and extensive kelp beds, is very similar to s. ornata's native home range (acuña et al. 2004) and much potential exists for this species to spread along this coast. despite this, s. ornata's distribution and abundance was last assessed more than 10 years ago (robinson et al. 2004) and its impact on indigenous sandy-shore communities has not been considered. this is a notable gap in current knowledge as sea anemones can play an important role in marine communities through their role as predators (kruger & griffiths 1998). against this background, this study aimed firstly to determine the current status and distribution of s. ornata, secondly to assess the diet of this anemone so as to establish its role as an alien predator and thirdly to investigate its impact on the structure of the sandy-shore communities of invaded areas in langebaan lagoon. research method and design top ↑ setting this study was conducted within langebaan lagoon in the wcnp (figure 1). the lagoon forms a wetland of international importance and is registered under the ramsar and bonn conventions. the lagoon opens into the southern end of saldanha bay (pitcher & calder 1998) and consists of sandflats with clean, fine sand as well as salt marshes. the lagoon supports a great diversity of marine taxa, many of which are not found elsewhere along the west coast or are restricted to estuarine environments (robinson & griffiths 2002). figure 1: map of saldanha bay and langebaan lagoon, showing the 15 study areas which were sampled as well as the west coast national park boundaries. figure 2: mean density (+se) of sagartia ornata recorded per m2 in 2001 and 2013. distribution and abundance of sagartia ornata sampling was undertaken in april and may 2013. the lagoon was divided into areas of 3 km, following robinson et al. (2004) (figure 1). boundaries of areas used remain the same although the areas have been renumbered 1–15 for convenience. areas supporting suitable habitat (containing s. maritima beds or sand-covered rocks) were visited. in areas where the anemone was detected 10 quadrates of 0.25 m2 were randomly placed in each shore zone (high-, midand low-shore zones) and the number of anemones recorded. in addition, areas outside of the wcnp but within saldanha bay (i.e. marcus island, small bay and lynch point), as well as rocky shores 10 km north (i.e. jacobs bay) and south (i.e. tsaarsbank) of the mouth of the bay were searched to determine if this species had spread outside the park. coelenteron contents analysis to assess s. ornata's role as a predator, the coelenteron contents of 50 individuals that were collected randomly in various habitats around the lagoon were analysed. anemones were collected during low spring tide and each was placed into a separate jar. to each jar a mixture of one part sea water and one part 10% magnesium chloride (mgcl2) solution was added to relax and narcotise the anemones. these were then frozen at -4 °c. to collect any organisms egested during the period of transportation and preservation, the liquid from each jar was poured through a 15 µm mesh prior to processing. a dissecting microscope was used to examine the retrieved organisms and the anemones. the anemones were cut longitudinally to open their coelenterons and prey items were removed. prey items were identified to the lowest taxonomic group possible, counted and their wet-mass recorded. the shells of hard-shelled animals were also included in the weighing process. inedible items such as exoskeletons, empty shells and debris, as well as unidentifiable detritus were excluded from analysis, as it was unclear whether these items were ingested as debris or are parts of digested prey. the number of prey and the total mass of prey consumed by each anemone was expressed as a percentage. the percentage frequency of occurrence of each of the different types of prey organisms was determined. an index of relative importance (iri) was calculated for each prey species as an indication of the importance of each food item in the diet of s. ornata. these calculations were made using the following equation (berg 1979): where: %n = average numerical percentage occurrence of each item in the diet; %m = average percentage by mass of food item in all the guts examined; %f = frequency of occurrence of the food item. sample sizes were represented by the number of coelenterons examined that contained food items. occurrence is thus expressed as the percentage of coelenterons containing food in which each particular food category was found. community structure to determine the impact of s. ornata on community structure, community composition was compared between an invaded and uninvaded area. the areas were located in the mid-shore zone, as this was the zone where s. ornata was found. in each area, 10 randomly placed quadrats of 0.25 m2 were used to quantify community composition. all sediment within each quadrat was removed until the underlying rock to which the anemones were attached was reached (a depth of not more than 10 cm). this sediment was sieved through a 1 mm sieve to retain organisms occurring within the sediment. all biota recovered from above and below ground were identified to species level, counted and wet-weighed. statistical analyses prior to all univariate analysis, data were tested for homoscedasity and homogeneity of variances using statistica for windows (version 11). the abundance of the anemones recorded in each area was compared to that recorded in 2001 (robinson et al. 2004). a generalised least square (gls) model was used to determine the effect of area and year on the abundance of anemones. the dredge function in r studio (version 5.1) was used to select the model that best explained anemone abundance. the best model was selected based on the lowest akaike information criterion. invertebrate abundance (individuals per m2) and biomass (g per m2) were compared between invaded and uninvaded sites using a mann-whitney u test. the primer software package (plymouth marine laboratory, plymouth, uk) was used to consider community composition (based on numerical abundance and biomass) in invaded and uninvaded areas. to detect significant differences between the communities of the two sites, anosim (based on bray-curtis similarities) was used. a simper analysis identified the species which contributed most to any differences detected. non-metric multidimensional scaling (mds) plots and hierarchical cluster analysis was used to create a visual representation of any differences between communities. all multivariate analyses were carried out on fourth-root transformed non-standardised data. to complement the analysis of community composition, the shannon-wiener diversity index (h′) was used to assess the species diversity of each site (clarke & warwick 2001): where pi is the proportion of the total number of individuals arising from the ith species. this index was compared between invaded and uninvaded sites using a mann-whitney u test. results top ↑ distribution and abundance of sagartia ornata sagartia ornata occurred in the mid-shore zone. the abundance of s. ornata individuals differed significantly between areas (gls, f(7) = 13.67, p < 0.01) and years (gls, f(1) = 12.25, p < 0.01), with more anemones being recorded in areas 2, 10 and 14 in 2013 (t = 3.6, p < 0.01) ( 2). the presence of s. ornata was recorded in areas 3 (6.0 + 2.4 [se] individuals per m2) and 5 (3.6 + 2.2 [se] individuals per m2) for the first time in 2013 whilst it no longer occurred in areas 9 and 13. high densities of the alien anemone were maintained through time in area 14 (372.6 + 24.7 [se] and 508.4 + 69.1 [se] in 2001 and 2013, respectively). whilst s. ornata occurred in sandy areas covering underlying rock and in s. maritima beds in 2001, in 2013 it was no longer present in the s. maritima beds but was found instead in nanozostera capensis beds (areas 2 and 5) and attached to rocks resting on fossilised oyster beds (areas 3 and 10). no anemones were found outside the wcnp within saldanha bay or on the open coast. coelenteron contents analysis only 35 of the 50 anemones whose coelentera were inspected contained prey items. table 1 presents the percentages of numerical occurrence (%n), mass (%m) and frequency of occurrence (%f) as well as the iri of prey items, which were identified to taxonomic group. this index identified polychaetes as the most important prey, occurring most frequently (i.e. in 83% of anemones), accounting for 53% of prey by numbers and 31% by mass. amphipods were the second most important food item by mass (25%) and iri. leptostraca and tanaids both occurred in the same numbers (16%), although tanaids were recorded in more anemones (29%) than the tiny leptostraca (20%). crabs, bivalves, gastropods and hemichordata were present in the diet of s. ornata. table 1: the prey items found in the coelentera of sagartia ornata as well as their numerical percentage of occurrence (%n), percentage by mass (%m) and their frequency of occurrence (%f) in the diet. community structure invaded communities supported a significantly higher abundance (mann-whitney, u(18) = 56.5, p < 0.05) (figure 3a) and biomass (mann-whitney, u(18) = 55, p < 0.05) (figure 3b) of invertebrate individuals. additionally, multivariate comparisons of community structure revealed significant differences in communities in terms of both abundance (anosim, r = 0.799, p < 0.01) and biomass (anosim, r = 0.916, p < 0.01) (figure 4). these differences remained the same even when the contribution made by s. ornata was excluded. the simper revealed that differences in community structure were not caused by the loss or gain of any one species, but rather by changes in the abundance and biomass of numerous indigenous species. these included increases in the tanaid anatanais gracilis, the polychaete orbinia angrapequensis, the hermit crab diogenes brevirostris and gastropod clionella sinuata and concurrent decreases in the gastropod volvarina capensis and the crab hymenosoma orbiculare in invaded areas. invaded areas supported significantly higher species diversity (h′) (mann-whitney, u(18) = 78, p < 0.05) (figure 5). figure 3: sagartia ornata (a) abundance (individuals per m2) (note scale break) and (b) biomass (g per m2) in invaded and uninvaded sites. figure 4: non-metric multidimensional scaling plot of species (a) abundances and (b) biomass of areas invaded and uninvaded by sagartia ornata. dendogram based on group-average clustering for species (c) abundances and (d) biomass for these areas. all data was fourth-root transformed. the contribution made by sagartia ornata is included. figure 5: shannon-wiener species diversity (h′) for invaded and uninvaded sites. discussion top ↑ sagartia ornata has a wide native distribution that spans the mediterranean, britain and western europe (acuña et al. 2004). this species was first recorded in south africa in 2001, at which time it occurred only along langebaan lagoon's western shoreline within the wcnp (robinson et al. 2004). following this initial work, no follow-up studies have been conducted until now. this study found that s. ornata was still restricted to langebaan lagoon, although its distribution within the lagoon had changed. despite s. ornata living on rocky shores and in kelp holdfasts in its native habitat (gibson et al. 2001), it occurs in different habitats in south africa. in langebaan lagoon, s. ornata has been recorded in shallow sand partially covering underlying rocks, in s. maritima beds (robinson et al. 2004), attached to loose rocks on fossilised oyster beds and in beds of the cape eelgrass n. capensis. this shift in habitat is not uncommon for invading species. for example, the invasive zebra mussel dreissena polymorpha that occurs on hard substrates in rivers and lakes in its native range occurs on soft sediments in its invaded range (berkman et al. 2000). this exploitation of new habitats might reflect the adaptive ability often expressed by alien species (berkman et al. 2000). the spread of s. ornata to new areas within the park since 2001 is notable and is linked to its exploitation of new habitats (i.e. in n. capensis beds and the rather cryptic habitat of loose rocks resting on fossilised oyster beds). it is also possible that s. ornata might have been overlooked in the latter habitat in the 2001 survey, as it is not the type of habitat that was anticipated to support this anemone. it is unlikely, however, that this anemone will spread extensively within the lagoon because of the relatively restricted extent of the habitats in which it has been recorded. in addition to changes in distribution, the abundance of s. ornata has also changed, as more anemones were recorded during 2013 than during the 2001 survey. however, the reason for this change is not obvious. whilst the 2001 and 2013 surveys were conducted at the same time of the year (i.e. seasonal variation is unlikely to be of importance), routine monitoring of this anemone may reveal the drivers of the observed changes in abundance. previous studies have shown that anemones can be important predators in marine communities (kruger & griffiths 1997; posey & hines 1991) and as such the impact of s. ornata as a predator on the sandy-shore communities within langebaan lagoon is of interest. the coelenteron content analysis revealed that s. ornata ate a very limited diet, sand-dwelling polychaetes and amphipods being the most important prey items. the diet of an anemone normally reflects the faunal communities supported in its habitat (chintiroglou & koukouras 1992; kruger & griffiths 1998) and as the sandy-shore communities provide habitat for sand-dwelling and burrowing species (branch et al. 2010), it may explain their importance in the diet of s. ornata. sea anemones are sessile passive feeders mostly relying on prey to come to them (sebens 1981) and have various feeding mechanisms to meet their daily nutritional requirements. these include absorption of dissolved organic matter, forming of mutualisms with photosynthetic zooxanthellae (tsuchida & potts 1994), filter-feeding (lampitt & paterson 1987) and feeding macrophagously on either motile or wave-tumbled prey (sebens 1981). although the feeding mechanisms of s. ornata have not been investigated, it occurs in a sheltered habitat with little wave action, a habitat type where anemones rely to a great extent on mobile prey (kruger & griffiths 1998). it is therefore likely that s. ornata employs a prey catching strategy, as is common to anemones in temperate regions (kruger & griffiths 1996). the feeding mechanism of an anemone is an important determinant of its impact on the surrounding communities. anemones relying on wave action to supply them with prey would not have direct impacts on communities, as prey items would already have been removed from source populations (kruger & griffiths 1998). in contrast, anemones such as s. ornata that rely on mobile prey are more likely to affect their communities as prey are actively removed from communities. diet analyses can be affected by a variety of factors, such as poor preservation of specimens, egestion (lampitt & paterson 1987), digestion rate and turnover times (kruger & griffiths 1997), as they influence the retention time and quantities of prey items available for analysis (kruger & griffiths 1998). in addition, diet can vary on a seasonal and temporal basis (kruger & griffiths 1996). thus, the present study cannot be considered an exhaustive analysis of s. ornata's diet. however, it does provide an indication of the taxa consumed and insight into the predatory role of s. ornata. although the ecological role of s. ornata has not been documented in its native range, it was anticipated that this anemone would negatively affect native communities as this is a well-documented impact associated with many sandy-shore invaders (grosholz et al. 2000; robinson et al. 2007; ross et al. 2004). as predicted this pattern was observed for s. ornata, with invaded areas supporting a higher invertebrate abundance, biomass and diversity, as well as altered communities. as these changes were not driven by the loss of the species that were found to dominate s. ornata's diet, it is unlikely that this anemone is exerting its impact on native communities through its role as a predator. rather, these changes may be the result of impacts on the invaded habitat by s. ornata and indirect effects on native biota. this anemone appears to alter the surrounding habitat, consolidating sand and trapping coarse sediment (robinson pers. obs.). this has been observed elsewhere where alien species have changed soft sediment systems by altering abiotic factors such as water movement and sediment characteristics (berkman et al. 2000; mckinnon et al. 2009; ruiz et al. 1997). in turn, sediment type and structure influence the distribution, abundance and biomass of associated species (nel & branch 2013; thrush et al. 2003). this may explain the higher invertebrate abundance, biomass and elevated diversity observed in the sites where s. ornata has invaded. notably, the high abundance of tanaids recorded in the presence of s. ornata is likely a reflection of domination by coarser sediments, as these crustaceans are known to have reduced rates to tube building in the presence of sediments below 63 µm (krasnow & taghon 1997). in addition, tanaids themselves are known to change community structure by altering the habitat and preying on small and juvenile soft-bodied invertebrates (oliver & slattery 1985). it is thus suggested that a synergy between the primary impacts of s. ornata on habitat structure and the secondary impacts of these small crustaceans ultimately result in the observed differences between invaded and uninvaded areas. conclusion top ↑ this study aimed to provide an updated baseline on the status and distribution of s. ornata, to assess the diet of this alien anemone and to investigate its impact on the structure of the sandy-shore communities of the wcnp. a survey of the lagoon found that s. ornata remained restricted to the wcnp, although its distribution had changed. notably this anemone was recorded in n. capensis seagrass beds for the first time. nonetheless it is unlikely that s. ornata will spread extensively within this marine protected area because of the restricted nature of the habitats in which it occurs. diet analysis revealed the dominant prey items consumed by s. ornata were indigenous sand-dwelling polychaetes and amphipods. multivariate analysis showed that in invaded areas this alien altered sandy-shore community structure, increased diversity, density and biomass of native invertebrates. despite these impacts, the geographically restricted nature of the habitat types currently invaded by s. ornata limits the potential of this alien to negatively affect indigenous biota within the wcnp. however, routine monitoring of surrounding areas should be undertaken as the occurrence of s. ornata on rocky shores and in kelp beds in its native range suggests that this anemone could spread to outside the park. acknowledgements top ↑ the financial assistance of the national research foundation (nrf) and the nrf-dst centre of excellence for invasion biology are gratefully acknowledged. opinions expressed and conclusions arrived at are those of the authors and are not necessarily to be attributed to the nrf. we are also grateful to the west coast national park for supporting the study and granting us research permits. lastly, we would like to thank ben brooker, michelle jooste, haley pope and saachi sadchatheeswaran for assisting with fieldwork, sheree muller for laboratory assistance and koebraa peters and dr carol simon for assisting with species identification. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions t.b.r. (stellenbosch university) was project leader and responsible for the project design. she helped with data collection, guided statistical analyses and contributed to writing the manuscript. c.s. (stellenbosch university) undertook data collection and analysis and contributed to writing the manuscript. references top ↑ acuña, f.h., excoffon, a.c. & griffiths, c.l., 2004, ‘first record and redescription of the introduced sea anemone sagartia ornata (holdsworth, 1855) (cnidaria: actiniaria: sagartiidae) from south africa’, african journal of zoology 39, 314–318. berg, j., 1979, ‘discussion of methods of investigating the food of fishes, with reference to a preliminary study of the prey of gobiusculus flavescens (gobiidae)’, marine biology 50, 263–273. http://dx.doi.org/10.1007/bf00394208 berkman, p.a., garton, d.w., haltuch, m.a., kennedy, g.w. & febo, l.r., 2000, ‘habitat shift in invading species: zebra and quagga mussel population characteristics on shallow soft substrates’, biological invasions 2, 1–6. http://dx.doi.org/10.1023/a:1010088925713 bolton, j.j., andreakis, n. & anderson, r.j., 2011, ‘molecular evidence for three separate cryptic introductions of the red seaweed asparagopsis (bonnemaisoniales, rhodophyta) in south africa’, african journal of marine science 33, 263–271. http://dx.doi.org/10.2989/1814232x.2011.600339 branch, g.m., griffiths, c.l., branch, m.l. & beckley, l.e., 2010, two oceans: a guide to marine life of southern africa, struik nature, cape town. briski, e., bailey, s.a. & macisaac, h.j., 2011, ‘invertebrates and their dormant eggs transported in ballast sediments of ships arriving to the canadian coasts and the laurentian great lakes’, limnology and oceanography 56, 1929–1939. http://dx.doi.org/10.4319/lo.2011.56.5.1929 carlton, j.t., 1996, ‘pattern, process, and prediction in marine invasion ecology’, biological conservation 78, 97–106. http://dx.doi.org/10.1016/0006-3207(96)00020-1 chintiroglou, c. & koukouras, a., 1992, ‘the feeding habits of three mediterranean sea anemone species, anemonia viridis (forskål), actinia equina (linnaeus) and cereus pedunculatus (pennant)’, helgoländer meeresuntersuchungen 46, 53–68. cinar, m.e., 2013, ‘alien polychaete species worldwide: current status and their impacts’, journal of the marine biological association of the united kingdom 93, 1257–1278. http://dx.doi.org/10.1017/s0025315412001646 clark, b.m. & griffiths, c.l., 2012, ‘western pea crabs pinnixa occidentalis (rathbun, 1894) (brachyura: thoracotremata: pinnotheroidea) invade saldanha bay, south africa’, african journal of marine science 34, 153–156. http://dx.doi.org/10.2989/1814232x.2012.675128 clarke, k.r. & warwick, r.m., 2001, change in marine communities: an approach to statistical analysis and interpretation, 2nd edn., primer-e, plymouth. gibson, r., hextall, b. & rogers, a., 2001, photographic guide to the sea and shore life of britain and north-west europe, oxford university press, new york. grosholz, e.d., ruiz, g.m., dean, c.a., shirley, k.a., maron, j.l. & connors, p.g., 2000, ‘the impacts of a nonindigenous marine predator in a california bay’, ecology 81, 1206–1224. http://dx.doi.org/10.1890/0012-9658(2000)081[1206:tioanm]2.0.co;2 gurevitch, j. & padilla, d.k., 2004, ‘are invasive species a major cause of extinctions?’, trends in ecology and evolution 19, 470–474. http://dx.doi.org/10.1016/j.tree.2004.07.005 hewitt, c.l., gollasch, s. & minchin, d., 2009, ‘the vessel as a vector – biofouling, ballast water and sediments’, biological invasions in marine ecosystems ecological studies 204, 117–131. http://dx.doi.org/10.1007/978-3-540-79236-9_6 klein, g., macintosh, k., kaczmarska, i. & ehrman, j.m., 2010, ‘diatom survivorship in ballast water during trans-pacific crossings’, biological invasions 12, 1031–1044. http://dx.doi.org/10.1007/s10530-009-9520-6 krasnow, l.d. & taghon, g.l., 1997, ‘rate of tube building and sediment particle size selection during tube construction by the tanaid crustacean, leptochelia dubia’, estuaries 20, 534–546. http://dx.doi.org/10.2307/1352612 kruger, l.m. & griffiths, c.l., 1996, ‘sources of nutrition in intertidal sea anemones from the south-western cape, south africa’, south african journal of zoology 31, 110–119. kruger, l.m. & griffiths, c.l., 1997, ‘digestion rates of prey eaten by intertidal sea anemones from the south-western cape, south africa’, south african journal of zoology 32, 101–105. kruger, l.m. & griffiths, c.l., 1998, ‘sea anemones as secondary consumers on rocky shores in the south-western cape, south africa’, journal of natural history 32, 629–644. http://dx.doi.org/10.1080/00222939800770331 kruger, n., branch, g.m., griffiths, c.l. & field, j.g., 2005, ‘changes in the epibenthos of saldanha bay, south africa, between the 1960s and 2001: an analysis based on dredge samples’, african journal of marine science 27, 471–477. http://dx.doi.org/10.2989/18142320509504105 lampitt, r.s. & paterson, g.l.j., 1987, ‘the feeding-behavior of an abyssal sea-anemone from in situ time lapse photographs and trawl samples’, oceanologica acta 10, 455–461. mckinnon, j.g., gribben, p.e., davis, a.r., jolley, d.f. & wright, j.t., 2009, ‘differences in soft-sediment macrobenthic assemblages invaded by caulerpa taxifolia compared to uninvaded habitats’, marine ecology progress series 380, 59–71. http://dx.doi.org/10.3354/meps07926 mead, a., carlton, j.t., griffiths, c.l. & rius, m., 2011a, ‘revealing the scale of marine bioinvasions in developing regions: a south african re-assessment’, biological invasions 13, 1991–2008. http://dx.doi.org/10.1007/s10530-011-0016-9 mead, a., carlton, j.t., griffiths, c.l. & rius, m., 2011b, ‘introduced and cryptogenic marine and estuarine species of south africa’, journal of natural history 45, 2463–2524. http://dx.doi.org/10.1080/00222933.2011.595836 mead, a., griffiths, c.l., branch, g.m., mcquaid, c.d., blamey, l.k., bolton, j.j. et al., 2013, ‘human-mediated drivers of change: impacts on coastal ecosystems and marine biota of south africa’, african journal of marine science 35, 403–425. http://dx.doi.org/10.2989/1814232x.2013.830147 molnar, j.l., gamboa, r.l., revenga, c. & spalding, d., 2008, ‘assessing the global threat of invasive species to marine biodiversity’, frontiers in ecology and the environment 6, 485–492. http://dx.doi.org/10.1890/070064 nel, p. & branch, g.m., 2013, ‘assessment of the abundance and distribution of burrowing sandprawns and mudprawns (callichirus and upogebia species) in langebaan lagoon, south africa’, african journal of marine science 35, 1–14. http://dx.doi.org/10.2989/1814232x.2013.795914 oliver, j.s. & slattery, p.n., 1985, ‘effects of crustacean predators on species composition and population structure of soft-bodied infauna from mcmurdo sound, antarctica’, ophelia 24, 155–175. http://dx.doi.org/10.1080/00785326.1985.10429725 olivier, d., heinecken, l. & jackson, s., 2013, ‘mussel and oyster culture in saldanha bay, south africa: potential for sustainable growth, development and employment creation’, food security 5, 251–267. http://dx.doi.org/10.1007/s12571-013-0244-1 peters, k., griffiths, c. & robinson, t.b., 2014, ‘patterns and drivers of marine bioinvasions in eight western cape harbours, south africa’, african journal of marine science 36, 49–57. http://dx.doi.org/10.2989/1814232x.2014.890669 pimentel, d., zuniga, r. & morrison, d., 2005, ‘update on the environmental and economic costs associated with alien-invasive species in the united states’, ecological economics 52, 273–288. http://dx.doi.org/10.1016/j.ecolecon.2004.10.002 pitcher, g.c. & calder, d., 1998, ‘shellfish mariculture in the benguela system: phytoplankton and the availability of food for commercial mussel farms in saldanha bay, south africa’, journal of shellfish research 17, 15–24. posey, m.h. & hines, a.h., 1991, ‘complex predator-prey interactions within an estuarine benthic community’, ecology 72, 2155–2169. http://dx.doi.org/10.2307/1941567 robinson, t.b., branch, g.m., griffiths, c.l. & govender, a., 2007, ‘the invasion and subsequent die-off of mytilus galloprovincialis in langebaan lagoon, south africa: effects on natural communities’, marine biology 152, 225–232. http://dx.doi.org/10.1007/s00227-007-0697-x robinson, t.b. & griffiths, c.l., 2002, ‘invasion of langebaan lagoon, south africa, by mytilus galloprovincialis – effects on natural communities’, african zoology 37, 151–158. robinson, t.b., griffiths, c.l. & kruger, n., 2004, ‘distribution and status of marine invasive species in and bordering the west coast national park’, koedoe 47, 79–87. http://dx.doi.org/10.4102/koedoe.v47i1.73 robinson, t.b., griffiths, c.l., mcquaid, c.d. & rius, m., 2005, ‘marine alien species of south africa – status and impacts’, african journal of marine science 27, 297–306. http://dx.doi.org/10.2989/18142320509504088 ross, d.j., johnson, c.r., hewitt, c.l. & ruiz, g.m., 2004, ‘interaction and impacts of two introduced species on a soft-sediment marine assemblage in se tasmania’, marine biology 144, 747–756. http://dx.doi.org/10.1007/s00227-003-1223-4 ruiz, g.m., carlton, j.t., grosholz, e.d. & hines, a.n., 1997, ‘global invasion of marine and estuarine habitats by non-indigenous species: mechanisms, extent, and consequences’, american zoology 37, 621–632. sebens, k.p., 1981, ‘the allometry of feeding, energetics and body size in three sea anemone species’, biological bulletin 161, 152–171. http://dx.doi.org/10.2307/1541115 seebens, h., gastner, m.t. & blasius, b., 2013, ‘the risk of marine bioinvasion caused by global shipping’, ecology letters 16, 782–790. http://dx.doi.org/10.1111/ele.12111 strain, e.m.a & johnson, c.r., 2013, ‘the effects of an invasive habitat modifier on the biotic interactions between two native herbivorous species and benthic habitat in a subtidal rocky reef ecosystem’, biological invasions 15, 1391–1405. http://dx.doi.org/10.1007/s10530-012-0378-7 thrush, s.f., hewitt, j.e., norkko, a., nicholls, p.e., funnell, g.a. & ellis, j.i., 2003, ‘habitat change in estuaries: predicting broad-scale responses of intertidal macrofauna to sediment mud content’, marine ecology progress series 263, 101–112. http://dx.doi.org/10.3354/meps263101 tsuchida, c.b. & potts, d.c., 1994, ‘the effects of illumination, food and symbionts on growth of the sea anemone anthopleura elegantissima (brandt, 1835) i. ramet growth’, journal of experimental marine biology and ecology 183, 227–242. http://dx.doi.org/10.1016/0022-0981(94)90089-2 wonham, m.j., carlton, j.t., ruiz, g.m. & smith, l.d., 2000, ‘fish and ships: relating dispersal frequency to success in biological invasions’, marine biology 136, 1111–1121. http://dx.doi.org/10.1007/s002270000303 article information authors: rené fourie1 charles r. haddad1 anna s. dippenaar-schoeman2,3 anel grobler1 affiliations: 1department of zoology and entomology, university of the free state, south africa2agricultural research council – plant protection research institute, queenswood, south africa 3department of zoology and entomology, university of pretoria, south africa correspondence to: charles haddad postal address: po box 339, bloemfontein 9300, south africa dates: received: 18 sept. 2012 accepted: 29 jan. 2013 published: 09 apr. 2013 how to cite this article: fourie, r., haddad, c.r., dippenaar-schoeman, a.s. & grobler, a., 2013, ‘ecology of the plant-dwelling spiders (arachnida: araneae) of the erfenis dam nature reserve, south africa’, koedoe 55(1), art. #1113, 9 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1113 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/ koedoe.v55i1.1113-1. copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. ecology of the plant-dwelling spiders (arachnida: araneae) of the erfenis dam nature reserve, south africa in this original research... open access • abstract • introduction • materials and methods    • study area and period    • sampling methods    • guilds    • statistics • results    • grassland    • foliage • ethical considerations • discussion • conclusion • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ as part of the south african national survey of arachnida in the grassland biome, foliage-dwelling and grass-dwelling spiders (arachnida: araneae) were collected in the erfenis dam nature reserve in the central free state province from november 2005 to august 2007. foliage-dwelling spiders were collected from three common tree or shrub species (acacia karroo, searsia ciliata and searsia lancea) and grass-dwellers from four contrasting grasslands (uniform themeda triandra, mixed, weedy and woodland grasslands). from the grass layer, 1649 spiders were collected, representing 15 families and 82 species, whilst 496 tree-dwelling spiders were collected that represented 17 families and 52 species. there was some overlap in the fauna of the two strata, resulting in a total of 108 species from 18 families being collected. the araneidae, philodromidae, salticidae and thomisidae were consistently the most abundant in all grassland types and tree species, although salticidae were scarce on a. karroo. assemblage analysis indicates high similarity and overlap in the fauna of the four grassland types, suggesting that the structural complexity of grasslands has a limited effect on species composition. in contrast, the foliage-dwelling assemblages were more distinct, with only some overlap between the faunas of s. ciliata and a. karroo, suggesting a stronger vegetation structural effect in shaping arboreal spider assemblages. the isolation of trees and shrubs within the extensive grassy habitat may contribute to the more unique fauna and lower species richness of the woody vegetation. conservation implications: this study uncovered a rich diversity of plant-dwelling spiders from central south africa. grassland faunas show considerable temporal variation and some variability in microhabitat preferences, and sampling protocols should take this into account when spiders are considered in management plans and biodiversity surveys in reserves and private land. introduction top ↑ true grasslands were historically quite widespread throughout the southern parts of the african continent (bredenkamp, spada & kazmierczak 2002), but, at present, the grassland biome is largely limited to the central plateau of south africa, lesotho and parts of swaziland (carbutt et al. 2011; mucina & rutherford 2006; palmer & ainslie 2005). the grassland biome covers approximately a quarter of south africa’s land surface (palmer & ainslie 2005) and is considered to have an extremely high plant biodiversity, second only to that of the fynbos biome (low & rebelo 1996). a large proportion (approximately 40%) of the biome has been irreversibly transformed through agriculture, mining, urbanisation and industrialisation (mentis & huntley 1982; o’connor & kuyler 2005) and, as such, the remaining grassland habitat is of great conservation value and can be considered highly irreplaceable (egoh 2009). only an estimated 2.04% – 2.80% of the grassland area is conserved (carbutt et al. 2011; o’connor & kuyler 2005) and therefore effective management and conservation of private land is critical to protect the highly endemic fauna and flora of this biome (wessels et al. 2003). grasslands can be defined on the basis of a single-layered herbaceous plant community dominated by tufted graminoids (especially perennial poaceae), with perennial forbs that appear annually, and the rarity of woody plants, which are usually restricted to specific habitats that act as fire refugia, including drainage lines and rocky hilltops (carbutt et al. 2011). regarding spider biodiversity, data in the national collection of arachnida database at the agricultural research council – plant protection research institute, pretoria, indicates that nearly 910 point localities have been sampled in the grassland biome, but only 27 of these have more than 100 specimen records (foord, dippenaar-schoeman & haddad 2011). this shows that large parts of the biome remain very poorly studied. only as recently as three decades ago were the first ecological studies on spiders undertaken in the grassland biome, focusing on the ground-dwelling (jansen et al. in press; lotz, seaman & kok 1991; van den berg & dippenaar-schoeman 1991), termitophilous (haddad & dippenaar-schoeman 2002, 2006), grass-dwelling (dippenaar-schoeman, hamer & haddad 2011; haddad 2005) and litter-dwelling assemblages (butler & haddad 2011). consequently, there is considerable scope for research on spider biodiversity and ecological patterns in this unique vegetation type. a project was initiated in september 2005 to determine the arachnofauna of the erfenis dam nature reserve, located near theunissen in the free state province of south africa. initially, the primary focus of the study was to determine the impact of controlled burning on various faunal groups, which included arachnids, insects and small mammals. whilst the project was focused on the ground-dwelling fauna, the potential was later recognised to expand the project to the sampling of foliage-dwelling and grass-dwelling spiders from unburned areas as well. thus, the aim of this study was to determine the spider communities in four contrasting grassland types and three common tree species in the reserve over a 2-year period. this study also forms part of the south african national survey of arachnida (sansa) in the grassland biome. materials and methods top ↑ study area and period the erfenis dam nature reserve is located in the central part of the free state province, about 18 km south-east of theunissen. the dam that shares the same name as the reserve is part of the vet river system and extends over 3800 ha when full, with the reserve comprising about 400 ha of grassland surrounding the dam (stuart & stuart 1997). sampling was carried out during the last week of every third month from november 2005 until august 2007, coinciding with the end of each of the four seasons of the year. grass-dwelling spiders were collected from four contrasting types of grasslands (abbreviations and generalised coordinates given in parenthesis), which can be described as follows (plant classification follows germishuizen et al. 2006). uniform themeda triandra forssk. grassland (tg; 28°29.8′s, 26°47.5′e) that consisted primarily of homogenous t. triandra, with small scattered patches of eragrostis and cymbopogon grasses. mixed grassland (mg; 28°29.9′s, 26°48.5′e), which consisted of various eragrostis species, aristida congesta roem. & schult., enneapogon cenchroides (roem. & schult.) c.e.hubb, setaria pumila (poir.) roem. & schult., sporobolus fimbriatus (trin.) nees and tragus berteronianus schult., with very little t. triandra present. weedy grassland (weg; 28°29.7′s, 26°48.4′e) was structurally the most heterogenous, consisting of a. congesta, cymbopogon pospischilii (k.shumm.) c.e.hubb., eragrostis lehmanniana nees, e. tef (zucc.) trotter, setaria pumila and sporobolus fimbriatus grasses, with a variety of weeds, including nidorella resedifolia dc., and berkeya spp. lastly, woodland grassland (wog; 28°30.2′s, 26°47.5′e), which was dominated by t. triandra and eragrostis grasses, interspersed with acacia karroo hayne (mimosaceae), searsia lancea barkley (anacardiaceae) and ziziphus mucronata willd. (rhamnaceae) trees. foliage-dwelling spiders associated with woody plants were collected from three different tree or shrub species: the trees a. karroo (ak; 28°30.2′s, 26°47.5′e) and s. lancea (sl; 28°29.6′s, 26°47.6′e) and the low-growing shrub searsia ciliata licht. ex schult (sc; 28°29.9′s, 26°48.5′e). acacia karroo is widespread throughout south africa, except for the fynbos and succulent karoo biomes, and is found in a variety of habitats, from low-lying areas to the highveld. this tree species usually has a rounded crown, branching fairly low down on the trunk. it is variable in shape and size, reaching a maximum height of about 12 m where there is good water nearby. the leaves are finely textured and dark green, with flowers that appear in early summer in a mass of yellow pompoms. the thorns are usually paired, with a greyish to white colour and are long and straight (van wyk & van wyk 1997; venter & joubert 1984). searsia lancea is a small to medium sized evergreen tree that usually grows to a height of 7 m and a width of 7 m and is found throughout the free state. it is usually a single-stemmed, low branching tree that has a dense, soft, round canopy with compound leaves, possessing three narrowly lanceolate (lance shaped) leaflets. the small, inconspicuous flowers are greenish-yellow in colour and are produced from june until september (i.e. winter to early spring). searsia lancea occurs naturally in acacia woodland and along drainage lines, rivers and streams (van wyk & van wyk 1997; venter & joubert 1984). searsia ciliata forms an evergreen, thorny, thin and crooked-stemmed shrub that is 2 m – 4 m tall. the leaves are also trifoliolate, with minute flowers that are yellow-green in colour and insignificant. the fruits are drupes and are shiny light to dark brown in colour. it is widespread in the central drier parts of south africa (venter & joubert 1984). sampling methods grass-dwelling spiders were collected using a standard sweep net with a diameter of 35 cm. a total of 250 sweeps, comprising five subsamples of 50 sweeps each, were completed per grassland site during each sampling period. foliage-dwelling spiders were collected from the three woody plant species by holding a sweep net beneath vegetation and beating the same branch with a thick wooden stick three times per individual beat. a total of 100 beats, comprising two subsamples of 50 beats each, were taken from each tree species per sample date. the spiders were sorted in the field to avoid cannibalism and prevent specimens dying and drying out. all individuals were captured with a small glass vial and preserved in a larger bottle in 70% ethanol for each sample. the specimens were then sorted into morphospecies and then identified to family level and tallied. morphospecies were identified further to genus and species level by the second and third authors. owing to the poor taxonomic resolution of some families (e.g. linyphiidae, theridiidae and araneidae), some morphospecies could not be identified beyond family or genus level. guilds the spiders collected were grouped into guilds based on their foraging methods. spiders can be generally separated into two main groups, namely wandering spiders that actively search for prey and web-builders that construct silk structures to capture prey. the wandering spiders can be separated into ground wanderers (gw) and plant wanderers (pw), based on the substrates on which they are primarily active. web-builders can be separated based on the structure of their webs and the following groups can be recognised: funnel-web builders (fwb), gumfoot-web builders (gwb), hackle-web builders (hwb), orb-web builders (owb), sheet-web builders (swb) and space-web builders (spwb). statistics differences in monthly abundance and species richness between grassland types and between tree species were compared using repeated measures analysis of variance (anova) with a tukey-kramer multiple comparisons post-test. a linear regression analysis was performed to determine whether seasonal rainfall had an influence on the corresponding numbers of spiders present. all of these analyses were performed in graphpad instat version 3.00 (2008). the observed species richness for the grassland and tree species was calculated and the proportion of species collected per site was indicated as a percentage of the total collected for each method. chao estimated species richness (magurran 2004) was calculated using the equation: schao1 = sobs + f1² / 2f2. chao1 is based on the abundance data available, is a function of the ratio between the singletons and doubletons in the data and reaches an asymptote when each species is represented by at least two individuals. f1 equals the number of observed species represented by one individual and f2 equals the number of observed species represented by two individuals (magurran 2004). inventory completeness was analysed in paleontological statistics (past) version 2.07 (hammer, harper & ryan 2001) using the sample rarefaction function, which implements the ‘mao tau’ analytical procedure with standard errors indicated as 95% confidence intervals on the resulting plot. samples were arranged chronologically for the analyses. this type of analysis is based on a chao2-type estimator, as per colwell, mao and chang (2004). we conducted three separate analyses in past to assess levels of similarity in spider assemblages in the three tree species and four grassland types. firstly, we conducted a one-way analysis of similarity (anosim) procedure to determine similarity in spider assemblage structure based on the tree and grassland sample totals. anosim can be considered analogous to one-way anova and is performed on the underlying bray-curtis similarity matrix. spider abundance was transformed to absence or presence data prior to analysis. the results were presented as a global r-value that is analogous to the f-value calculated using anova. pair-wise comparisons between the tree or grassland types were presented as r-values that are analogous to the post hoc comparisons of anova (clarke & warwick 2001). a r-value close to 1 indicates high dissimilarity between the two compared assemblages, whilst a value close to 0 indicates high similarity in their composition. secondly, we performed a non-metric multidimensional scaling (mds) on the sample data from each grassland type or tree species based on a bray-curtis similarity matrix. the ordination diagrams produced group sites together based on similarities in their species composition, whilst also taking species abundance into account. samples that were most similar were grouped close to one another. we added convex hulls to each ordination diagram to group samples from each grassland type and tree species. lastly, we carried out a cluster analysis of the bray-curtis similarity matrix using the cluster function with a paired group algorithm. taking both mds and cluster analysis results into consideration allows for more accurate interpretation of the assemblage patterns (clarke & warwick 2001). results top ↑ grassland a total of 1649 spiders were collected from the four grassland types, representing 15 families and 82 species (see online appendix). the species richness of each of the grasslands varied between 38 and 56 and the number of unique species sampled from each (table 1) can largely be attributed to the sampling of singletons and doubletons. the rarefaction curve of the grassland samples did not reach an asymptote, indicating that further sampling would be necessary to sample all of the species present in the grasslands (figure 1a). table 1: species diversity and richness of grass-dwelling and tree-dwelling spiders collected by sweep-netting and foliage beating, respectively, in the erfenis dam nature reserve, free state province from november 2005 to august 2007. figure 1: sample rarefaction curves (mao tau) (in red) with 95% confidence intervals (in blue) for spiders collected from (a) four grassland types and (b) three tree or shrub species in the erfenis dam nature reserve, free state province, from november 2005 to august 2007. of the species sampled, 54 were hunting spiders (65.85%) whilst 28 species were web-builders (34.15%). the numerically dominant species collected were tibellus minor lessert, 1919 (philodromidae, 10.79%), misumenops rubrodecoratus millot, 1941 (thomisidae, 10.49%), thyene thyenioides (peckham & peckham, 1903) (salticidae, 9.16%), runcinia flavida (thomisidae, 6.19%) and tibellus hollidayi lawrence, 1952 (6.06%) (see online appendix). all of these species are active hunting spiders and contributed largely to the dominance of philodromidae, thomisidae and salticidae in the survey, whilst araneidae, which were also relatively abundant, had several species contributing to the overall abundance of the family (figure 2a). other families were not universally abundant but were only prominent in one, two or three types of grasslands. araneidae and salticidae were clearly more abundant in the mixed grassland, thomisidae were most prominent in the woodland grassland, whilst philodromidae were considerably lower in abundance in the woodland grassland compared to the three other grassland types (figure 2a). figure 2: proportional abundance of the dominant spider families collected from (a) four grassland types and (b) three tree or shrub species in the erfenis dam nature reserve, free state province from november 2005 to august 2007. monthly spider abundance did not differ significantly between the four grassland types (anova: f3,7 = 0.6461, p = 0.5940). however, monthly species richness differed significantly between sites (anova: f3,7 = 9.086, p = 0.0005). species richness was significantly higher in mixed grassland (p < 0.001) and themeda grassland (p < 0.01) than in weedy grassland and also significantly higher in mixed than in woodland grassland (p < 0.05).the anosim comparisons of the grass-dwelling spider assemblages (table 2) pointed to high levels of similarity between grassland spider assemblages (global r = 0.0685, p = 0.0786). the pair-wise comparisons between sites showed significant dissimilarity between the assemblages of the mixed and woodland grasslands, and between mixed and weedy grasslands, but none of the other compared assemblages differed significantly from one another (table 2). this was supported by the results of the mds ordination (figure 3a) and cluster analysis (figure 3b), which showed a large degree of overlap between the assemblages of the four grasslands. it is interesting to note that on several occasions the assemblages of the four grassland types clustered together based on the month of sampling (e.g. november 2005, may 2006 and february 2007) or were in adjacent groups within the analysis (february 2006, august 2006 and may 2007), rather than grouping together based on grassland type (figure 3b). table 2: analysis of similarity results comparing the grass-dwelling spider assemblages of four contrasting grassland types in the erfenis dam nature reserve, free state province from november 2005 to august 2007. figure 3: analysis of (a, b) grass-dwelling and (c, d) tree or shrub-dwelling spider assemblages in the erfenis dam nature reserve, free state province collected from november 2005 to august 2007, where (a, c) are non-metric multidimensional scaling based on bray-curtis similarity matrices and (b, d) are cluster analyses of the bray-curtis similarity matrices. regarding phenology, spider abundance appears not to be directly correlated to the rainfall of that particular season (r2 = 0.0231), but rather to preceding rainfall history, suggesting a lag effect (figure 4a). this pattern may have been compounded in february 2006 by the very high rainfall that month, which may have restricted spider activity during this period, accounting for the drastic drop in spider abundance. however, in november 2005, high numbers of grass-dwelling spiders were collected, even though rainfall was low in the previous three months. figure 4: seasonal abundance of spiders collected from (a) four grassland types and (b) three tree or shrub species in the erfenis dam nature reserve, free state province, compared to seasonal rainfall (mm) from november 2005 to august 2007. foliage a total of 496 spiders were collected by foliage-beating, representing 17 families and 52 species (online appendix). of the three species, s. ciliata had much higher species richness and many more unique species; a. karroo and s. lancea had a similar number of unique species (table 1). the rarefaction curve for the tree samples also did not reach an asymptote (figure 1b) and the slope was noticeably steeper than in the case of the grassland samples (figure 1a), indicating that a considerable number of species occurring on the woody plants were not sampled. compared to the grass-dwelling fauna, a slightly lower proportion of the foliage-dwellers were hunting spiders (32 spp., 61.54%); 20 species (38.46%) construct webs to capture prey. the most abundant foliage-dwelling species were philodromus browningi lawrence, 1952 (philodromidae, 13.91%), cheiracanthium furculatum karsch, 1879 (miturgidae, 9.88%), m. rubrodecoratus (8.47%), baryphas ahenus simon, 1902 (salticidae, 6.65%) and neoscona subfusca (c.l. koch, 1837) (araneidae, 5.47%). the abundance of these species, particularly p. browningi and m. rubrodecoratus, were largely responsible for shaping patterns of family dominance (figure 2b). philodromidae and thomisidae were most common on a. karroo, whilst salticidae, theridiidae and araneidae were more abundant on s. ciliata than the other woody species. whilst all of the families showed low abundance on s. lancea, theridiidae was totally absent from this species (figure 2b). differences in spider abundance between tree species were very significant (anova: f2,7 = 6.499, p = 0.0072). both a. karroo (n = 206) and s. ciliata (n = 220) had significantly more spiders than the 65 individuals collected from s. lancea (p < 0.05), but differences between a. karroo and s. ciliata were not significant (p > 0.05). monthly species richness also differed significantly between tree species (anova: f2,7 = 7.842, p = 0.0052), although significant differences in the pairwise comparisons were only detected between s. ciliata and s. lancea (p < 0.01). the anosim results indicated considerable overlap between assemblages of the tree species (global r = 0.3664, p < 0.0001). the pair-wise comparisons indicated the greatest dissimilarity between the assemblages of s. lancea and s. ciliata (r = 0.5145, p = 0.0003), with increasing similarity between the assemblages of a. karroo and s. lancea (r = 0.3078, p = 0.0036) and between a. karroo and s. ciliata (r = 0.2698, p = 0.0151). the mds ordination of the arboreal spider assemblages (figure 3c) supported these results and indicated that s. lancea has the most distinctive assemblage, whilst there was considerable overlap (and thus greater similarity) between the assemblages of a. karroo and s. ciliata. however, in the cluster analysis none of the trees’ samples clustered together in a single group, although six of the eight s. lancea samples fell within adjoining groups (figure 3d). this supported the mds results that indicated this tree as having the most distinct spider assemblage. as for the grass-dwelling fauna, rainfall in the season of sampling did not have a significant influence on the numbers of spiders in the tree foliage (r2 = 0.0976). similar depressions in spider abundance were observed during the summer of 2006 and during the winter months, with spider abundance peaking during the spring of the 2005–2006 year of sampling, and during the spring or summer of the 2006–2007 year of sampling, depending on the tree species (figure 4b). ethical considerations top ↑ this study was made possible through collecting permits from free state nature conservation. the study follows the ethical guidelines proposed for the sansa, approved by the sansa steering committee. discussion top ↑ this study provided the first comprehensive dataset on the grass-dwelling and foliage-dwelling spiders of the grassland biome of south africa within an ecological context. in part, the study aimed to assess the influence of vegetation diversity and structure on spider assemblages. as spiders are predators, they interact with the vegetation indirectly, which means that structural support available for webs and active hunting, prey availability and suitable microclimate may influence the composition of a spider community (e.g. dippenaar-schoeman & wassenaar 2006; gunnarsson 1990; harwood, sunderland & symondson 2001; haughton et al. 1999; spears & macmahon 2012; uetz & burgess 1979). spider abundance and diversity may be correlated with the specific vegetation characteristics, suggesting that the availability of habitats is important for spider colonisation and establishment (jŏgar, metspalu & hiiesaar 2004). free state grasslands consist of a high diversity of grasses and shrubs that create greater habitat complexity, which means that there are many niches in which spiders may live (haddad 2005). yet, as a habitat, grasses appear quite uniform in structure, which may limit the availability of structurally different niches in grasslands, despite high overall grass species richness. of the grasslands studied here, the mixed, woodland and weedy grasslands all had a more complex structure than the uniform themeda grassland. yet, the structurally complex weedy grassland has considerably lower spider species richness and projected species richness than the other habitats, despite quite similar abundance (table 1). according to mgobozi, somers and dippenaar-schoeman (2008), the progressive invasions of the invasive alien weed chromolaena odorata (l.) king & robinson (asteraceae) brings with it negative changes in native spider abundance, assemblage patterns, diversity and estimated species richness. it could be that the weeds present in the weedy grassland emitted allomones, which may have repelled many insects. this, coupled with a decrease in suitable plant species for phytophagous insects, may have caused a decrease in food resources for spiders, effectively contributing to the lower spider diversity. the estimated species richness predicted that the themeda grassland would have the highest species richness (table 1). yet, the mixed grassland was the most diverse habitat (structurally, as well as having the most plant species) and also had the most species rich spider assemblage. although the themeda grassland was very structurally uniform, it was very closely the second most species rich grassland. this is probably the result of the complex structure of the grass stems and inflorescences and the very dense tussocks that t. triandra form (less dense in the other grasses). in the present study, a similar number of species was collected in the themeda grassland (54 spp.) compared to another study conducted in similarly homogenous themeda grassland in the central free state (haddad 2005), in which 57 species were collected. the species richness in the woodland grassland was only slightly lower than the mixed and themeda grasslands and may have been expected to be higher as a result of the presence of woody plants, which could have created a more structurally complex habitat with more niches available for occupation by spiders. thus, expected migration between grasses and woody plants is apparently minimal and does not necessarily create more complex grass-dwelling assemblages. the estimated species richness for the three woody plants sampled predicted that s. lancea would have the highest species richness and a. karroo the least, but the s. ciliata shrubs had greater observed species richness and also more unique species than the two trees (table 1). this could be the result of the denser foliage of s. ciliata than that of the trees, which provides greater microhabitat complexity in the foliage of the plant. this can also be illustrated by the high abundance of web-building theridiidae in these shrubs, but their absence or scarcity in the two tree species. the two tree species also have thick trunks that are covered in coarse bark, which contributes to the structural complexity of these trees, but because only peripheral foliage was sampled the contribution that trunk-dwelling spiders could make to the assemblage structure of these trees was minimal. whilst the grassland fauna showed a higher species richness than the foliage-dwelling fauna, this may be an artefact of collecting effort, which is reflected in the total abundance collected by the two methods (number of foliage spiders was approximately one-third of the grassland spiders, with 40% of the number of sample units taken from grassland), as well as the slopes of the respective rarefaction curves. the steeper slope of the curve for foliage-dwellers indicates that considerably more species could still be sampled than in the case of the grassland fauna. considering the contrasting physical structures of the three woody species sampled, it was unusual to find that the two related species (s. lancea and s. ciliata) had the most different spider assemblages and that there was considerable overlap in the assemblages of s. ciliata with a. karroo. although a slightly larger number of unique species was collected from the woody vegetation, and the assemblages harboured by each tree or shrub species were generally more distinctive than in the case of the grass-dwelling fauna, it is likely that further sampling from the woody plants (to collect comparable numbers of individuals from the two strata) would have considerably reduced the number of singleton and doubleton species and effectively reduced the number of unique species apparently occurring on the woody plants. this may also have increased the species richness of the foliage fauna considerably, which may have produced higher levels of similarity between tree species. as such, the sampling discrepancies may mask the true levels of overlap in the assemblages of the woody plants. both hunting and web-building spiders are sensitive to changes in habitat structure and rainfall can affect habitat structure in subtle ways (langlands, brennan & pearson 2006). in their study, conducted in the great victoria desert in australia, sampling years with high spider abundance coincided with years of good rainfall that took place prior to sampling (langlands et al. 2006). this was also evident in the present study, with spider numbers increasing following months with good rainfall and decreasing following months with low rainfall. a bottom-up effect from rainfall might be responsible for the increases in spider abundance; however, the linear regression test results showed that rainfall of a particular season had no significant influence on spider abundance of that season in both the foliage and grass layers. our data would suggest that there is a lag between the times that rain falls and increased or decreased spider abundance; in the case of an increase this could be explained by the time needed for plants to grow and create more complex niches, which, in turn, attract a greater number of prey for spiders. at a finer temporal scale, rain falling immediately before or during the sampling day may also influence spider activity and hence capture rates (romero & vasconcellos-neto 2003). this may explain the low number of spiders captured in february 2006, following several days of heavy rainfall. according to langlands et al. (2006), some spiders tend to have strong seasonality, so this could also influence the number of spiders present during a year (e.g. van den berg & dippenaar-schoeman 1991). seasonal abundance of tree spiders sometimes differs from grass-dwelling spiders (dippenaar-schoeman, van den berg & van den berg 1989), although our study found a similar general pattern in both strata. most studies show peaks in temporal patterns of spider numbers and species during early to mid-summers (hatley & macmahon 1980; niemelä et al. 1994). more spiders were captured in the summer months than in the winter months for both strata, which reflect the high availability of prey and emergence of immature spiders from egg sacs during spring. lower prey availability during the winter is not problematic to spiders because they are very resistant to starvation and can lower their metabolic rates through inactivity, which enables them to survive harsh conditions (dippenaar-schoeman & jocqué 1997). according to aitchison (1987), there are winter-active spiders that utilise this season to gain an ecological advantage of little competition from other spiders or other predaceous invertebrates for food, or alternately are species that feed primarily on winter-active prey (e.g. dippenaar-schoeman, de jager & van den berg 1996). a good example from the present study is the archaeodictyna sp. (dictynidae), of which most specimens were collected in autumn and winter. however, most of the species appear to be strongly affected by the onset of winter, during which spider numbers were at their lowest. it is not surprising that the dominant species of both strata came from araneidae, salticidae and thomisidae, as these are some of the most diverse families of spiders (dippenaar-schoeman & jocqué 1997). these families have also dominated the plant-dwelling fauna in other south african studies in grassland (haddad 2005), savannah (dippenaar et al. 2008; dippenaar-schoeman, van den berg & prendini 2009; dippenaar-schoeman et al. 1989; haddad et al. 2010; muelelwa et al. 2010) and forest habitats (dippenaar-schoeman & wassenaar 2006). curiously, only a single species, m. rubrodecoratus, was abundant in both strata, indicating that this species is very flexible in its ecological requirements, whilst the other more abundant species are more particular in their microhabitat preferences. conclusion top ↑ the present study provided comprehensive data on the biodiversity of grass-dwelling and foliage-dwelling spiders in the grassland biome and tried to explain differences in assemblage structure in relation to vegetation structure and diversity. the dominant plant-dwelling families from both strata were the araneidae, philodromidae, salticidae and thomisidae, although only one species (m. rubrodecoratus) was abundant in both strata. grass-dwelling assemblages were richer in species than foliage-dwellers and this was likely an artefact of sampling intensity. sampling protocols should be adapted to have an equal number of sweeps and beats to collect more comparable numbers of specimens, which may better explain species richness and assemblage composition patterns. grassland assemblages are generally similar and show considerable overlap in species, indicating that the grass fauna is probably more flexible in their ecological requirements and can occupy more structurally variable plant communities within the much more extensive grassy habitat. this suggests limited effects of plant diversity and structural complexity in shaping assemblages. the foliage-dwelling spider communities differed somewhat from one another, suggesting that either the sampling intensity was inadequate or that the vegetation structures were indeed markedly different, affecting community structure through niche specialisation. estimations of total species richness (chao1) for grassland habitats and tree species suggest that grasslands consistently maintain much higher species richness than the isolated woody plants. as such, trees may function as islands within the large grassland matrix, supporting very different assemblages than the surrounding grassy habitat, but with their isolation contributing to lower levels of species richness than the surrounding continuous grassland habitat. acknowledgements top ↑ we thank the erfenis dam nature reserve staff, particularly the conservation manager robert lotze, for support during the study and permission to collect in the reserve. we would also like to thank robin lyle, dewald du plessis and joan adendorff for occasional assistance with fieldwork. andor venter (university of the free state) kindly identified some of the plant species. this study forms part of the first author’s msc at the university of the free state and was funded through a national research foundation of south africa (nrf) grant in the nrf thuthuka programme to the second author (grant number ttk2008050500003). the two anonymous referees provided useful comments and suggestions to improve the quality of the manuscript and their inputs are greatly appreciated. competing interests the authors declare that they have no financial or personal relationships that may be inappropriately affected through the production of this article. authors’ contributions this study formed part of r.f.’s (university of the free state) msc on spider ecology in the erfenis dam nature reserve. r.f. conducted field work, sorted and tallied specimens, performed some statistical analyses and wrote part of the manuscript. c.r.h. (university of the free state) supervised r.f., performed initial identifications, performed some statistical analyses and wrote part of the manuscript. a.s.d-s. (agricultural research council – plant protection research institute) conducted final identifications and contributed to writing the manuscript. a.g. (university of the free state) assisted with field work and sorting specimens. references top ↑ aitchison, c.w., 1987, ‘feeding ecology of winter-active spiders’, in w. nentwig (ed.), ecophysiology of spiders, pp. 264–273, springer-verlag, berlin. http://dx.doi.org/10.1007/978-3-642-71552-5_19bredenkamp, g.j., spada, f. & kazmierczak, e., 2002, ‘on the origin of northern and southern hemisphere grasslands’, plant ecology 163, 209–229. http://dx.doi.org/10.1023/a:1020957807971 butler, v.p. & haddad, c.r., 2011, ‘spider assemblages associated with leaf litter of three tree species in central south africa (arachnida: araneae)’, african journal of ecology 49, 301–310. http://dx.doi.org/10.1111/j.1365-2028.2011.01265.x carbutt, c., tau, m., stephens, a. & escott, b., 2011, ‘the conservation status of temperate grasslands in southern africa’, grassroots 11, 17–23. clarke, k.r. & warwick, r.m., 2001, change in marine communities: an approach to statistical analysis and interpretation, 2nd edn., primer-e, plymouth. colwell, r.k., mao, c.x. & chang, j., 2004, ‘interpolating, extrapolating, and comparing incidence-based species accumulation curves’, ecology 85, 2717–2727. http://dx.doi.org/10.1890/03-0557 dippenaar-schoeman, a.s. & jocqué, r., 1997, african spiders: an identification manual, plant protection research institute handbook 9, agricultural research institute, pretoria. dippenaar-schoeman, a.s. & wassenaar, t.d., 2006, ‘a checklist of spiders from the herbaceous layer of a coastal dune forest ecosystem at richards bay, kwazulu-natal, south africa (arachnida: araneae)’, african invertebrates 47, 63–70. dippenaar-schoeman, a.s., de jager, m. & van den berg, a., 1996, ‘ammoxenus species (araneae: ammoxenidae) – specialist predators of harvester termites in south africa’, african plant protection 2, 103–109. dippenaar-schoeman, a.s., hamer, m. & haddad, c.r., 2011, ‘an annotated checklist of the spiders (arachnida: araneae) of the mkambathi nature reserve, eastern cape, south africa’, koedoe 53, 1–10.  dippenaar-schoeman, a.s., van den berg, a. & prendini, l., 2009, ‘spiders and scorpions (arachnida: araneae, scorpiones) of the nylsvley nature reserve, south africa’, koedoe 51, 1–9. http://dx.doi.org/10.4102/koedoe.v51i1.161 dippenaar-schoeman, a.s., van den berg, a.m. & van den berg, a., 1989, ‘species composition and relative seasonal abundance of spiders from the field and tree layers of the roodeplaat dam nature reserve’, koedoe 32, 25–38. http://dx.doi.org/10.4102/koedoe.v32i1.462 dippenaar, s.m., modiba, m.a., khoza, t.t. & dippenaar-schoeman, a.s., 2008, ‘a checklist of the spiders (arachnida, araneae) of the polokwane nature reserve, limpopo province, south africa’, koedoe 50, 10–17. http://dx.doi.org/10.4102/koedoe.v50i1.128 egoh, b.n., 2009, integrating ecosystem services into conservation planning in south africa, unpublished phd thesis, department of botany and zoology, stellenbosch university. foord, s.h., dippenaar-schoeman, a.s. & haddad, c.r., 2011, ‘south african spider diversity: african perspectives on the conservation of a mega-diverse group’, in o. grillo & g. venora (eds.), changing diversity in changing environment, pp. 163–182, intech publishing, rijeka. germishuizen, g., meyer, n.l., steenkamp, y. & keith, m. (eds.), 2006, a checklist of south african plants, south african botanical diversity network report 41, sabonet, pretoria. graphpad instat version 3.00 for windows 95, 2008, computer software, graphpad software, san diego. gunnarsson, b., 1990, ‘vegetation structure and the abundance and size distribution of spruce-living spiders’, journal of animal ecology 59, 743–752. http://dx.doi.org/10.2307/4892 haddad, c.r., 2005, ‘ecology of spiders (arachnida: araneae) inhabiting themeda triandra forsskǻl grassland in semi-arid south africa’, navorsinge van die nasionale museum bloemfontein 21, 25–36. haddad, c.r. & dippenaar-schoeman, a.s., 2002, ‘the influence of mound structure on the diversity of spiders (araneae) inhabiting the abandoned mounds of the snouted harvester termite trinervitermes trinervoides’, journal of arachnology 30, 403–408. http://dx.doi.org/10.1636/0161-8202(2002)030[0403:tiomso]2.0.co;2 haddad, c.r. & dippenaar-schoeman, a.s., 2006, ‘spiders (araneae) inhabiting abandoned mounds of the snouted harvester termite trinervitermes trinervoides (sjöstedt) (isoptera: termitidae: nasutitermitinae) in the free state, south africa, with notes on their biology’, navorsinge van die nasionale museum bloemfontein 22, 1–15. haddad, c.r., honiball, a.s., dippenaar-schoeman, a.s., slotow, r. & van rensburg, b.j., 2010, ‘spiders (arachnida: araneae) as indicators of elephant-induced habitat changes in the maputaland centre of endemism, south africa’, african journal of ecology 48, 446–460. http://dx.doi.org/10.1111/j.1365-2028.2009.01133.x hammer, ø., harper, d.a.t. & ryan, p.d., 2001, ‘past – palaeontological statistics’, palaeo electronica 4, 1–33. harwood, j.d., sunderland, k.d. & symondson, w.o.c., 2001, ‘living where the food is: web location by linyphiid spiders in relation to prey availability in winter wheat’, journal of applied ecology 38, 88–99. http://dx.doi.org/10.1046/j.1365-2664.2001.00572.x hatley, c.l. & macmahon, j.a., 1980, ‘spider community organization: seasonal variation and the role of vegetation architecture’, environmental entomology 9, 632–639. haughton, a.l., bell, j.r., boatman, n.d. & wilcox, a., 1999, ‘the effects of different rates of the herbicide glyphosate on spiders in arable field margins’, journal of arachnology 27, 249–254. jansen, r., makaka, l., little, i.t. & dippenaar-schoeman, a.s., in press, ‘response of ground-dwelling spider assemblages (arachnida, araneae) to montane grassland management practices in south africa’, journal of insect conservation. jǒgar, k., metspalu, l. & hiiesaar, k., 2004, ‘abundance and dynamics of wolf spiders (lycosidae) in different plant communities’, agronomy research 2, 145–152. langlands, p.r., brennan, k.e.c. & pearson, d.j., 2006, ‘spiders, spinifex, rainfall and fire: long-term changes in an arid spider assemblage’, journal of arid environments 67, 36–59. http://dx.doi.org/10.1016/j.jaridenv.2006.01.018 lotz, l.n., seaman, m.t. & kok, d.j., 1991, ‘surface–active spiders (araneae) of a site in semi-arid central south africa’, navorsinge van die nasionale museum bloemfontein 7, 530–540. low, a.b. & rebelo, a.g., 1996, vegetation of south africa, lesotho and swaziland, department of environmental affairs and tourism, pretoria. magurran, a.e., 2004, measuring biological diversity, blackwell publications, bodmin. mentis, m.t. & huntley, b.l., 1982, a description of the grassland biome project, south african national scientific programmes report 62, graphic arts division of the centre for scientific and industrial research, pretoria. mgobozi, m.p., somers, m.j. & dippenaar-schoeman, a.s., 2008, ‘spider responses to alien plant invasion: the effect of shortand long-term chromolaena odorata invasion and management’, journal of applied ecology 45, 1189–1197. mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, strelitzia 19, south african national biodiversity institute, pretoria. muelelwa, m.i., foord, s.h., dippenaar-schoeman, a.s. & stam, e.m., 2010, ‘towards a standardized and optimized protocol for rapid assessments: spider species richness and assemblage composition in two savannah vegetation types’, african zoology 45, 273–290. http://dx.doi.org/10.3377/004.045.0206 niemelä, j., pajunen, t., haila, y., punttila, p. & halme, e., 1994, ‘seasonal activity of boreal forest-floor spiders (araneae)’, journal of arachnology 22, 23–31. o’connor, t.g. & kuyler, p., 2005, ‘national grasslands initiative: identification of compatible land uses for maintaining compatible biodiversity integrity’, unpublished report, south african national biodiversity institute, pretoria. palmer, a.r. & ainslie, a.m., 2005, ‘grasslands of south africa’, in j.m. suttie, s.g. reynolds & c. batello (eds.), grasslands of the world, pp. 77–120, plant production and protection series 34, food and agriculture organization of the united nations, rome. romero, g.o. & vasconcellos-neto, j., 2003, ‘natural history of misumenops argenteus (thomisidae): seasonality and diet on trichogoniopsis adenantha (asteraceae)’, journal of arachnology 31, 297–304. http://dx.doi.org/10.1636/02-19 spears, l.r. & macmahon, j.a., 2012, ‘an experimental study of spiders in a shrub-steppe ecosystem: the effects of prey availability and shrub architecture’, journal of arachnology 40, 218–227. http://dx.doi.org/10.1636/p11-87.1 stuart, c. & stuart, t., 1997, guide to southern african game and nature reserves, 4th edn., struik publishers, cape town. uetz, g.w. & burgess, j., 1979, ‘habitat structure and colonial behaviour in metepeira spinipes (araneae: araneidae), an orb-weaving spider from mexico’, psyche 86, 79–90. http://dx.doi.org/10.1155/1979/76206 van den berg, a. & dippenaar-schoeman, a.s., 1991, ‘ground-living spiders from an area where the harvester termite hodotermes mossambicus occurs in south africa’, phytophylactica 23, 247–253. van wyk, b. & van wyk, p., 1997, field guide to trees of southern africa, struik publishers, cape town. venter, h.j.t. & joubert, a.m., 1984, climbers, tree and shrubs of the orange free state, p.j. de villiers publishers, bloemfontein. wessels, k.j., reyers, b., van jaarsveld, a.s. & rutherford, m.c., 2003, ‘identification of potential conflict areas between land transformation and biodiversity conservation in north-eastern south africa’, agriculture, ecosystems and environment 95, 157–178. http://dx.doi.org/10.1016/s0167-8809(02)00102-0 article information author: peter f. scogings1 affiliation: 1department of agriculture, university of zululand, south africa correspondence to: peter scogings email: pscoging@pan.uzulu.ac.za postal address: private bag x1001, kwadlangezwa 3886, south africa dates: received: 04 nov. 2010 accepted: 07 apr. 2011 published: 08 july 2011 how to cite this article: scogings, p.f., 2011, ‘stem growth of woody species at the nkuhlu exclosures, kruger national park: 2006–2010’, koedoe 53(1), art. #1035, 8 pages. doi:10.4102/koedoe.v53i1.1035 copyright notice: © 2011. the authors. licensee: aosis openjournals. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) stem growth of woody species at the nkuhlu exclosures, kruger national park: 2006–2010 in this original research... open access • abstract • introduction • methods    • study area    • design and procedure • results • discussion and conclusion • acknowledgements • references abstract (back to top) an important aspect of managing african conservation areas involves understanding how large herbivores affect woody plant growth. yet, data on growth rates of woody species in savannas are scarce, despite its critical importance for developing models to guide ecosystem management. what effect do browsing and season have on woody stem growth? assuming no growth happens in the dry season, browsing should reduce stem growth in the wet season only. secondly, do functional species groups differ in stem growth? for example, assuming fine-leaved, spiny species’ growth is not compromised by carbon-based chemical defences, they should grow faster than broad-leaved, chemically defended species. dendrometers were fixed at 20 cm in height on the main stems of 244 random plants of six woody species in three plots (all large herbivores excluded, partial exclusion, and control) and observed from late 2006 to early 2010. average monthly increment (ami) per dendrometer and season (dry, wet) was calculated and the interaction between plot and season tested per species, controlling for initial stem girth. amis of combretum apiculatum, dichrostachys cinerea and grewia flavescens were zero in the dry season, whilst those of acacia exuvialis, acacia grandicornuta and euclea divinorum were either positive or negative in the dry season. wet-season ami of d. cinerea and dry-season ami of g. flavescens tended to be reduced by browser exclusion. net ami (sum of the seasonal amis) was tested per species, but results suggested that only d. cinerea tended to be affected by browser exclusion. the results also suggested that stem radial growth of some fast-growing species is more prone to reduction by browser exclusion than the growth of other species, potentially reducing their competitiveness and increasing their risk of extirpation. finally, the usefulness of grouping woody species into simple functional groups (e.g. fine-leaved vs. broad-leaved) for ecosystem management purposes in savannas requires further consideration. conservation implications: growth rates of woody plants are important parameters in savanna models, but data are scarce. monitoring dendrometers in manipulative situations over several years can help fill that gap. results of such studies can be used to identify species prone to high risk of extirpation. introduction (back to top) a major component of current research in savannas concerns the effects of large herbivores (especially elephants) on woody vegetation, with the ultimate objective being the development of improved management policies (levick & rogers 2008). growth rates of woody plants are important parameters in models developed for the purposes of understanding savanna ecosystems and ultimately guiding the management of extensive conservation areas comprising heterogeneous savanna landscapes and mega-herbivores, but critical data on growth are scarce (baxter & getz 2005). apart from the dearth of studies of woody plant growth rates under the influence of elephants, most studies of responses of woody plants to browsing in savannas have only recorded shoot growth, which generally responds positively to long-term severe browsing (engdahl 2008; fornara & du toit 2007; hrabar, hattas & du toit 2009; makhabu, skarpe & hytteborn 2006; riginos & young 2007). numerous short-term clipping studies have revealed the same trend (bergström, skarpe & danell 2000; fonara & du toit 2008a; rooke & bergström 2007; tsumele, mlambo & sebata 2006). the response of shoots is not necessarily a real response in total plant growth, but is the result of resources being reallocated to rectify browsing-induced shifts in the root : shoot ratio (teague 1985; teague & walker 1988). numerous studies have reported negative effects of long-term severe browsing on plant height (augustine & mcnaughton 2004; birkett & stevens-wood 2005; fornara & du toit 2008b; levick & rogers 2008; noumi et al. 2010). in contrast, very few studies have considered stem size, which has been found to either decrease or increase under long-term severe browsing (augustine & mcnaughton 2004; dharani et al. 2008; noumi et al. 2010; riginos & young 2007; rohner & ward 1999; scogings 1998). the significance of stem size is that it is closely related to plant mass and therefore is likely to reflect the whole plant’s response, which is important to understand because whole-plant under-compensation may indicate progressive carbon (c) exhaustion and reduced life expectancy (hester et al. 2006; nickless, scholes & archibald 2010; teague 1988). spatial and temporal variations in resource availability are important considerations for understanding plant responses to browsing in africa’s savannas (sankaran, ratnam & hanan 2008). savannas are seasonal systems characterised by alternating wet and dry seasons of several months each (kutsch et al. 2008). water, and hence nutrients, are most available when elevated soil water drives a pulse of nitrogen (n) mineralisation at the start of the wet season (jacobs et al. 2007; scholes, bond & eckhardt 2003). plant growth is therefore confined to the wet season, but is intermittent because of fluctuating resources that limit growth (williams et al. 2009). in addition to the temporal variation in resource availability, soil fertility varies spatially from sandy soils to clay-rich soils (hopcraft, olff & sinclair 2009; scholes et al. 2003). conventionally, clay-rich savannas are described as being dominated by ‘fine-leaved’, spiny, tannin-poor, ‘palatable’ species, whilst clay-poor savannas are said to be dominated by ‘broad-leaved’, spineless, tannin-rich, ‘unpalatable’ species (kutsch et al. 2008; scholes 1997). whether mineral n is always available in the soil (e.g. clay) or is only available when mineralised by soil organisms, water is needed in both cases to mobilise n for plants (jacobs et al. 2007; williams et al. 2009 and references therein). plants on both soil types could therefore experience temporary periods of ‘rich’ resource availability in otherwise ‘poor’ resource conditions, regardless of whether the soils are perceived to be ‘fertile’ or not (jacobs et al. 2007). hence, it is distinctly feasible that ‘fine-leaved’ and ‘broad-leaved’ species, which may be intuitively expected to respond differently to each other, may not differ consistently in their responses. the primary aim of this paper is to report stem girth growth of six ubiquitous woody species in the acacia nigrescens – combretum apiculatum savanna of the kruger national park during the period 2006–2010. in doing so, the paper aims to explore the question: what effect does browsing and season have on woody stem growth? the following null hypothesis was formulated to guide the analysis: browsing does not reduce stem growth in either the wet season or the dry season. the hypothesis was tested by analysing data compiled from dendrometer observations between the fourth and eighth years following the establishment of the nkuhlu large-scale long-term exclusion experiment (nllee). hence, it was not only possible to collect valuable data on woody stem growth in a semi-arid savanna, but other unanswered questions fundamental to understanding the functioning of savannas could be addressed in an experimental context. because contrasting soils existed at either end of the catena on which the experiment was situated, it was also possible to consider potential influences of soil types on woody stem growth. furthermore, because the suite of species used in the study comprised both ‘fine-leaved’ and ‘broad-leaved’ species, it was possible to explore whether or not patterns of stem growth reflected traditionally recognised functional groups of woody species. this paper represents the first documentation of data on stem growth rates of a suite of woody species in relation to large mammal herbivores in african savannas. methods (back to top) study area the nllee (24°58′s, 31°46′e) is adjacent to the sabie river, kruger national park, south africa (o’keefe & alard 2002). the climate is semi-arid subtropical with two broadly distinct seasons: a hot, occasionally wet, growth season (october–april) and a warm, dry, non-growing season (williams et al. 2009). mean annual rainfall (mar) at skukuza, 30 km west of nkuhlu, is approximately 550 mm. average daily temperatures at skukuza are 15.7 °c in june and 26.6 °c in january ( http://www.sanparks.org/parks/kruger/conservation/scientific/weather). average minimum temperature in june is 5.7 °c and average maximum temperature in january is 32.6 °c. the topography is an undulating landscape, 200 m – 230 m above mean sea level, derived from granite and covering the sequence of terrain morphology from footslope to crest. the crests and middle slopes are characterised by shallow, sandy, coarse soil overlying rock, whilst the footslopes below the seepline are characterised by deep, sodic, duplex soil, which is a typical pattern on catenas in granite-derived, semi-arid landscapes (grant & scholes 2006; khomo & rogers 2005). sodic soils in the study area comprise shallow (< 15 cm) sand overlying impermeable clay and have high ph (> 8.5) and reduced hydraulic conductivity (grant & scholes 2006; khomo & rogers 2005; tarasoff, mallory-smith & ball 2007). sodic soils are therefore regarded as stressful environments for vegetation, which is sparse, but regarded as more attractive than crest vegetation to large herbivores, especially grazers and mixed feeders (levick & rogers 2008; tarasoff et al. 2007). it is unclear if the attractiveness of sodic patches to herbivores is because the vegetation is inherently more nutritious, or because it offers other attractions such as predator vigilance, water, dietary salts or anti-acidosis minerals (abrahamson 1999; khomo & rogers 2005). nevertheless, trampling, excretion and defoliation are thought to maintain the vegetation in a nutritious vegetative state (grant & scholes 2006). plant communities were described by siebert and eckhardt (2008). abundant woody species (nomenclature follows schmidt, lötter & mccleland 2007) include acacia grandicornuta, euclea divinorum, spirostachys africana and pappea capensis on the sodic footslopes and dichrostachys cinerea, acacia exuvialis, acacia nigrescens, combretum apiculatum, combretum hereroense, combretum zeyheri, grewia flavescens, grewia bicolor, lannea schweinfurthii rhigozum zambesiacum, ormocarpum trichocarpum and philenoptera violacea on the sandy middle slopes and crests. most of the woody species are deciduous and shorter than 10 m. common mammal herbivores (nomenclature follows skinner & chimimba 2005) include impala (aepyceros melampus), african elephant (loxodonta africana), hippo (hippopotamus amphibius), black rhino (diceros bicornis), blue wildebeest (connochaetes taurinus), cape buffalo (syncerus caffer), plains zebra (equus quagga), greater kudu (tragelaphus strepsiceros), steenbok (raphicerus campestris), giraffe (giraffa camelopardalis) and scrub hare (lepus saxatilis). elephant, impala, kudu, giraffe, black rhino and steenbok include substantial woody vegetation in their diets. elephant density in the study area fluctuates, but has been estimated to be 0.5 km-2 – 2.0 km-2, which is considered to be high (grant et al. 2008). a breeding herd of 30–40 impala, several impala bachelors, two black rhino and an unknown number of steenbok reside in the study area, whilst low numbers of giraffe and kudu occur occasionally. design and procedure experimental exclusion plots were established in 2002 on the premise that exclusion of large herbivores (especially elephants) would allow their effects on components of the ecosystem to be determined (o’keefe & alard 2002). the experiment covers 139 ha and is intended to be maintained for at least 25 years. owing to the scale of the experiment, it was not practical to replicate treatments (jonsson et al. 2010; levick & rogers 2008). one exclusion plot (approximately 70 ha) excludes all herbivores the size of hares and larger, whilst a second exclusion plot (approximately 44 ha) excludes elephants (and also giraffe due to their height), but not other herbivores (o’keefe & alard 2002). the two exclusion plots are separated by a control area (approximately 25 ha) open to all large herbivores. therefore, if excluding either elephants or all large herbivores shows similar responses then it can be inferred that the main source of the effect is elephants (engdahl 2008; jonsson et al. 2010). if excluding elephants shows a different response to excluding all large herbivores then it can be inferred that the main source of the effect is not elephants. manual dendrometers (agricultural electronics corporation, tucson, usa) were used for the study. each dendrometer consisted of a metal band (3 mm × 0.75 mm) of user-defined length attached to a spring-loaded vernier gauge graduated to 0.1 mm precision. the dendrometers were attached at a height of 20 cm above ground on the main stems of 244 randomly selected individuals of three ‘fine-leaved’ species (a. exuvialis, a. grandicornuta, d. cinerea) and three ‘broad-leaved’ species (c. apiculatum, e. divinorum, g. flavescens) in both exclusion plots as well as in the control area. the fitted trees were in areas protected from fire. the selected species were amongst the most abundant woody species in the study area. several extra dendrometers were deployed in the control area compared to the exclusion plots to allow for anticipated losses over the anticipated years of monitoring (e.g. by animals uprooting or felling plants, or damaging the instruments by chewing or rubbing on them). a gps was used to randomly locate sites in a grid of 30 m × 30 m cells. an anomalous basaltic outcrop covering approximately 15% of the partial exclusion plot was excluded from sampling. one tree of each study species was located in each grid cell on condition that it was, (1) closest to the grid-cell centre, (2) taller than 2 m and (3) neither obviously stressed by disease, disturbance or neighbours, nor obviously growing in a nutrient-enriched patch such as a termite mound or hollow. the minimum height of selected g. flavescens plants was 1.5 m because it seldom grew taller than 2 m. if any condition was not met for the closest tree, then the next closest tree that met the conditions was sampled. girth (mm) of the dominant stem was measured before positioning a band on the stem. the vernier gauges were initially set at approximately 5 mm to allow for possible shrinkages to be recorded and were reset whenever they reached > 25 mm. the dendrometers were observed (to 0.1 mm) at intervals of one–three months, depending on logistical constraints, from september 2006 to february 2010. total rainfall at skukuza in 2006/2007 and 2007/2008 was below mar, but in 2008/2009 and 2009/2010 it was above mar (figure 1). furthermore, monthly rainfall during the second half of three of the wet seasons included in the study period was below the long-term average for each month. not all dendrometers were operational for the whole study period because of failure or damage. increments between successive observations were calculated for the operational period of each dendrometer. figure 1: monthly rainfall at skukuza for the duration of the study and two years prior to the study. for comparison, long-term average monthly rainfall is also shown. total annual rainfall for each year (july to june) is indicated above the bars. to test the interaction between plot and season, the wet season was simply defined as the six months from november to april, whilst the dry season comprised the remaining six months. monthly increments per dendrometer were calculated by dividing each individual increment by the number of months over which the increment had occurred. average monthly increments (amis) per season were then calculated across years for each dendrometer, giving a wet-season ami (amiwet) and a dry-season ami (amidry) for each dendrometer. net ami (nami) for each dendrometer was calculated as the sum of amiwet and amidry. each species’ data were analysed separately to test effects within species using systat 10 (spss 2000). two-factor anova was used to test the interaction effect of plot and season on either log10(ami + 1) or √(ami + 1), whilst one-factor anova was used to test the effect of plot on either log10 (nami + 1) or √(nami + 1), whichever transformation normalised the distribution of a species’ data. individual plants were assumed as replicates within each treatment. therefore, pseudo-replication constrained the analyses and estimates of model parameters were viewed with caution (underwood 1997:245). initial stem girth was included as a covariate in all analyses, but was dropped from the model if it had no effect. in all analyses, significant effects were declared when p < 0.05, but possible trends were not ignored when 0.05 < p < 0.10 indicated marginal significance. finally, to facilitate consideration of differences amongst functional groups of species, estimates of average annual growth per species were obtained by calculating average annual girth growth from mean nami × 12 and average annual radial growth though division of girth growth by π/2. because the dendrometers used in this study are intended to be observed for several more years, a thorough quantitative analysis of annual growth rates will be appropriate after a longer series of annual data has been completed. results (back to top) mean initial girth of each plant varied amongst the study species when it was fitted with a dendrometer (figure 2). initial girth was largest for a. grandicornuta, followed by c. apiculatum, but was smallest for g. flavescens. initial girth had no effect as a covariate in any of the analyses (p > 0.05). all species experienced significant effects of season on ami (p < 0.05), whilst marginally significant interactions between plot and season affected d. cinerea (f2,72 = 2.92; p = 0.060) and g. flavescens (f2,72 = 2.58; p = 0.082). wet-season ami of d. cinerea and dry-season ami of g. flavescens tended to be reduced by exclusion of all large herbivores (figure 3). reduction of the analysis to a simple comparison of amidry and amiwet within species revealed that, whilst amidry was zero for d. cinerea, g. flavescens and c. apiculatum, it was negative for a. exuvialis and positive for a. grandicornuta and e. divinorum (figure 4). when nami was tested per species, d. cinerea was affected only marginally by browser exclusion (f2,36 = 2.61; p = 0.087). mean nami of d. cinerea was 0.97 mm (sem = 0.226), 1.26 mm (sem = 0.236) and 1.77 mm (sem = 0.273) in the complete exclusion, partial exclusion and control plots, respectively. figure 2: mean initial girth (mm) of stems fitted with dendrometers at the nkuhlu exclusion experiment. error bars are standard errors of the means (from left to right, n = 39, 41, 41, 39, 39 and 44). species from left to right are: euclea divinorum, acacia exuvialis, acacia grandicornuta, dichrostachys cinerea, grewia flavescens and combretum apiculatum. figure 3: mean monthly increment in stem girth of six abundant woody species in the wet and dry seasons at the nkuhlu exclusion experiment, 2006–2010. error bars are standard errors of the means (for columns fe, ne and pe, n = 13, 15 and 14 for acacia exuvialis; 12, 15 and 12 for acacia grandicornuta, euclea divinorum, dichrostachys cinerea and combretum apiculatum; and 12, 18 and 12 for grewia flavescens). fe, exclusion of all large herbivores greater than hares; ne, control (no exclusion); pe, exclusion of elephants and giraffe only. figure 4: mean monthly increment in stem girth of six abundant woody species in the wet and dry seasons across all treatments at the nkuhlu exclusion experiment, 2006–2010. species from left to right are: grewia flavescens, combretum apiculatum, dichrostachys cinerea, acacia grandicornuta, acacia exuvialis and euclea divinorum. error bars are standard errors of the means (n = 44 for g. flavescens, 39 for d. cinerea, 39 for c. apiculatum, 42 for a. exuvialis, 41 for a. grandicornuta, and 39 for e. divinorum). average annual girth growth rates of the measured trees were estimated to be 10.6 mm, 8.2 mm and 8.0 mm for g. flavescens, d. cinerea and c. apiculatum, respectively, and 6.1 mm, 6.0 mm and 5.8 mm for a. exuvialis, a. grandicornuta and e. divinorum, respectively. correcting these estimates to account for differences in initial girth amongst the species gives values of 0.09 mm mm-1, 0.05 mm mm-1, 0.03 mm mm-1, 0.04 mm mm-1, 0.01 mm mm-1 and 0.03 mm mm-1, respectively. narrow-stemmed species (g. flavescens, d. cinerea and a. exuvialis) were relatively faster growing than wide-stemmed species (e. divinorum, c. apiculatum and a. grandicornuta). corresponding average annual radial growth rates (uncorrected) of the measured trees were 1.68 mm, 1.31 mm, 1.27 mm, 0.96 mm, 0.95 mm and 0.92 mm. therefore, the slower growing trees included in the study increased girth by approximately 5.9 mm per year, whilst the faster growing trees increased girth by approximately 8.9 mm per year, corresponding to approximately 0.94 mm and 1.42 mm radial growth per year. discussion and conclusion (back to top) the null hypothesis that browsing does not reduce the growth of woody stem girths in either the wet season or the dry season was not rejected for the particular suite of species observed over the duration of the study at the nllee. because excluding only elephants had no effect compared to the control treatment, it can be inferred that any trends associated with exclusion were not related to elephants. the most intuitive reason for not observing effects of browser exclusion is that the timing and duration of the study were not long enough to detect differences in semi-arid savannas, even though several studies across a wide range of rainfall (and soils) have shown otherwise (e.g. augustine & mcnaughton 2004; dharani et al. 2008; riginos & young 2007; sharam, sinclair & turkington 2006). on the other hand, no difference in stem girth growth suggests a positive effect of browsing on plant growth rate leading to compensation (renton, thornby & hanan 2007). however, the results did suggest that certain trends existed amongst the species during the study period, which are worth monitoring for some years to come as they may develop into real effects over a longer time period. for example, after several more years it is expected that negative effects of complete exclusion of large herbivores will become more pronounced (scogings et al. in press). the trends in the data also suggest partial exclusion may have negative effects on some species, for example, d. cinerea. the observed trends in d. cinerea complement observed changes in height of woody plants in the first five years of browser exclusion (scogings et al. in press). both the dendrometer and height data showed d. cinerea responding more strongly than other species to browser exclusion, where exclusion of all large herbivores increased height but tended to decrease stem girth, relative to no or partial exclusion. increasing height growth and decreasing girth may reflect increased competition for light, as is typically seen when shaded plants grow tall and thin (o’connor 1995). the results suggest stem radial growth of some fast-growing species is more prone to reduction by browser exclusion than the growth of other species, potentially as an indirect effect through increased competition (scogings et al. in press). after a few more years, the data will also give insights into mortality rates of each species monitored, as either direct or indirect effects of browser exclusion. the results suggested that exclusion of large herbivores may lead to increasingly adverse effects on growth rates of some species; therefore exclusion may influence drought related mortality in the long term because fast-growing plants are prone to drought stress (pollastrini et al. 2010). dharani et al. (2008) measured acacia xanthophloea trees of similar size to the ones measured at the nllee. growth was faster with than without browsing, which corroborates some of the trends observed amongst species studied at the nllee. riginos and young (2007) found faster stem growth in acacia drepanolobium with than without browsing, possibly due to elephants reducing competitive effects of grasses. in contrast, other studies have found reduced growth for several species under browsing compared to exclusion (augustine & mcnaughton 2004; scogings 1998). the overall stem growth rates observed at the nllee appear to be comparable to those observed in other studies. dharani et al. (2008) observed stem growth rates higher than the ones at the nllee, but scogings (1998) observed growth rates of six species that were generally lower than similar size trees at the nllee. stem growth of grewia tenax plants over three years following browser exclusion was found by augustine and mcnaughton (2004) to be similar to that of g. flavescens plants of similar height at the nllee, but acacia mellifera and acacia etbaica had much higher growth rates compared to any of the species studied at the nllee. none of the aforementioned studies observed either zero or negative rates. further research is needed before generalisations in stem growth amongst species and sites can be postulated. because of the complexity of savanna ecosystems, efforts to understand their functioning necessitate simplification by means of models. much is often generalised about the ecology of woody plants in savannas on the basis of functional groups that have been traditionally adopted to facilitate such efforts (e.g. see staver et al. 2009), although no clear definitions of the traditionally used terms exist in a savanna context. the leaf-size dichotomy usually forms the basis for further generalisations to be made about other aspects of woody species, such as the ‘palatability’ of woody plants in savannas. ‘fine-leaved’ species are perceived to be ‘palatable’ because they are deemed to be poorly endowed with c-based secondary metabolites (e.g. condensed tannins) in favour of thorns or spines (kutsch et al. 2008; scholes 1997). the perception derives from an assumed trade-off between growth and chemical defences (herms & mattson 1992). the observation that many ‘fine-leaved’ woody species are, (1) involved in bush encroachment and (2) display stimulated shoot growth in response to browsing, may be regarded as supporting the ultimate view that ‘fine-leaved’, spiny species are ‘palatable’, fast-growing and tolerant of severe browsing, compared to ‘broad-leaved’ species (moleele & perkins 1998; sweet & mphinyane 1986). the validity of such generalisations has not been tested. although the dendrometer study was not designed to test differences amongst functional groups of species, growth rates did not obviously correspond with leaf type because ‘fine-leaved’ species did not grow faster than ‘broad-leaved’ species. furthermore, analysis of low molecular weight phenolic compounds known to influence herbivores did not show clear distinction between ‘fine-leaved’ and ‘broad-leaved’ species at the nllee (hattas et al. in press). acacia exuvialis, in particular, is an example of ‘fine-leaved’ species that are well endowed with both c-based secondary metabolites and long thorns. however, growth rates did appear to be related to soil type and slope position. acacia grandicornuta and e. divinorum, which were abundant on the clay-rich sodic soils of the footslopes, maintained slightly positive growth in the dry season, whilst species on the sandy crests either did not grow or shrank in the dry season, which is likely related to differences in the timing of water availability across the catena (jacobs et al. 2007). in conclusion, this study provides evidence that stem growth of woody plants in african savannas is strongly seasonal, but the effects of large herbivores on woody plant growth remains unclear and requires further research. the results also suggest that catenal position and associated differences in soil type and hydrology generally influences stem growth. not all species within the broad functional groups of woody plants traditionally recognised in african savannas show the same stem growth patterns. long-term monitoring of dendrometers fitted to a variety of species in manipulative situations would provide much-needed insights into growth dynamics of woody plants, which could help managers of conservation areas to identify species prone to high risk of extirpation. acknowledgements (back to top) the zululand/sweden kruger browse project was funded by the national research foundation, university of zululand, agricultural research council, swedish research council and swedish international development agency. i would also like to acknowledge tuulikki rooke, dawood hattas, luthando dziba and alpheus zobolo for their enthusiastic contributions to development and implementation of the project, as well as thandeka mamashela, patricia shabangu, ntuthuko mkhize, julius tjelele, basanda nondlazi, gilbert pule, sicelo dludla, andries malate, ngcebo ngcobo, frederik engdahl and elin gunve for their help in the field. scientific services personnel, especially adolf manganyi, nokukhanya mpanza and noel nzima, also provided critical support for the fieldwork. finally, i would like to acknowledge the assistance provided by the two anonymous reviewers, who contributed to improving the manuscript. references (back to top) abrahamson, p.w., 1999, ‘the chemistry and mineralogy of three savanna lick soils’, journal of chemical ecology 25, 2215−2228. doi:10.1023/a:1020861505138 augustine, d.j. & mcnaughton, s.j., 2004, ‘regulation of shrub dynamics by native browsing ungulates on east african rangeland’, journal of applied ecology 41, 45−58. doi:10.1111/j.1365-2664.2004.00864.x baxter, p.w.j. & getz, w.m., 2005, ‘a model-framed evaluation of elephant effects on tree and fire dynamics in african savannas’, ecological applications 15, 1331−1341. doi:10.1890/02-5382 bergström, r., skarpe, c. & danell, k., 2000, ‘plant response and herbivory following simulated browsing and stem cutting of combretum apiculatum’, journal of vegetation science 11, 409−414. doi:10.2307/3236633 birkett, a. & stevens-wood, b., 2005, ‘effect of low rainfall and browsing by large herbivores on an enclosed savannah habitat in kenya’, african journal of ecology 43, 123−130. doi:10.1111/j.1365-2028.2005.00555.x dharani, n., kinyamario, j.i., wagacha, p.w. & rodrigues, a.j., 2008, ‘browsing impact of large herbivores on acacia xanthophloea benth in lake nakuru national park, kenya’, african journal of ecology 47, 184−191. doi:10.1111/j.1365-2028.2008.00954.x engdahl, f., 2008, ‘herbivory on woody plants and induced responses in two similar species of acacia in the kruger national park, south africa’, msc thesis, department of wildlife, fish and environmental studies, swedish university of agricultural sciences. fornara, d.a. & du toit, j.t., 2007, ‘browsing lawns? responses of acacia nigrescens to ungulate browsing in an african savanna’, ecology 88, 200−209. doi:10.1890/0012-9658(2007)88[200:blroan]2.0.co;2 fornara, d.a. & du toit, j.t., 2008a, ‘responses of woody saplings exposed to chronic mammalian herbivory in an african savanna’, ecoscience 15, 129−135. doi:10.2980/1195-6860(2008)15[129:rowset]2.0.co;2 fornara, d.a. & du toit, j.t., 2008b, ‘community-level interactions between ungulate browsers and woody plants in an african savanna dominated by palatable-spinescent acacia trees’, journal of arid environments 72, 534−545. doi:10.1016/j.jaridenv.2007.07.010 grant, c.c., bengis, r., balfour, d. & peel, m., 2008, ‘controlling the distribution of elephants’, in r.j. scholes & k.g. mennell (eds.), elephant management: a scientific assessment for south africa, pp. 329−369, wits university press, johannesburg. grant, c.c. & scholes, m.c., 2006, ‘the importance of nutrient hotspots in the conservation and management of large wild mammalian herbivores in semi-arid savannas’, biological conservation 130, 426−437. doi:10.1016/j.biocon.2006.01.004 hattas, d., hjältén, j., julkunen-tiitto, r., scogings, p.f. & rooke, t., in press, ‘differential phenolic profiles in six african savanna woody species in relation to antiherbivore defense’, phytochemistry. herms, d.a. & mattson, w.j., 1992, ‘the dilemma of plants: to grow or defend’, quarterly review of biology 67, 283−335. doi:10.1086/417659 hester, a.j., bergman, m., iason, g.r. & moen, j., 2006, ‘impacts of large herbivores on plant community structure and dynamics’, in k. danell, r. bergström, p. duncan & j. pastor (eds.), large herbivore ecology, ecosystem dynamics and conservation, pp. 97−141. cambridge university press, cambridge. doi:10.1017/cbo9780511617461.006 hopcraft, j.g.c., olff, h. & sinclair, a.r.e., 2009, ‘herbivores, resources and risks: alternating regulation along primary environmental gradients in savannas’, trends in ecology & evolution 25, 119−128. doi:10.1016/j.tree.2009.08.001 hrabar, h., hattas, d. & du toit, j.t., 2009, ‘differential effects of defoliation by mopane caterpillars and pruning by african elephants on the regrowth of colophospermum mopane foliage’, journal of tropical ecology 25, 301−309. doi:10.1017/s0266467409005872 jacobs, s.m., bechtold, j.s., biggs, h.c., grimm, n.b., lorentz, s., mcclain, m.e. et al., 2007, ‘nutrient vectors and riparian processing: a review with special reference to african semiarid savanna ecosystems’, ecosystems 10, 1231−1249. doi:10.1007/s10021-007-9092-1 jonsson, m., bell, d., hjältén, j., rooke, t. & scogings, p.f., 2010, ‘do mammalian herbivores influence invertebrate communities via changes in the vegetation? results from a preliminary survey in kruger national park, south africa’, african journal of range & forage science 27, 39−44. khomo, l.m. & rogers, k.h., 2005, ‘proposed mechanism for the origin of sodic patches in kruger national park, south africa’, african journal of ecology 43, 29−34. doi:10.1111/j.1365-2028.2004.00532.x kutsch, w.l., hanan, n., scholes, r.j., mchugh, i., kubheka, w., eckhardt, h. et al., 2008, ‘response of carbon fluxes to water relations in a savanna ecosystem in south africa’, biogeosciences 5, 1797−1808. doi:10.5194/bg-5-1797-2008 levick, s. & rogers, k.h., 2008, ‘patch and species specific responses of savanna woody vegetation to browser exclusion’, biological conservation 141, 489−498. makhabu, s.w., skarpe, c. & hytteborn, h., 2006, ‘elephant impact on shoot distribution on trees and on rebrowsing by smaller browsers’, acta oecologica 30, 136−146. doi:10.1016/j.actao.2006.02.005 moleele, n.m. & perkins, j.s., 1998, ‘encroaching woody species and boreholes: is cattle density the main driving factor in the olifants drift communal grazing lands, south-eastern botswana’, journal of arid environments 40, 245−253. doi:10.1006/jare.1998.0451 nickless, a., scholes, r.j. & archibald, s., 2010, ‘calculating the variance and prediction intervals for estimates obtained from allometric relationships’, paper presented at the 3rd biennial csir conference, pretoria, 30 august – 01 september. noumi, z., touzard, b., michalet, r. & chaieb, m., 2010, ‘the effects of browsing on the structure of acacia tortilis (forssk.) hayne ssp. raddiana (savi) brenan along a gradient of water availability in arid zones of tunisia’, journal of arid environments 74, 625−631. o’connor, t.g., 1995, ‘acacia karroo invasion of grassland: environmental and biotic effects influencing seedling emergence and establishment’, oecologia 103, 214−223. doi:10.1007/bf00329083 o’keefe, t. & alard, g., 2002, effects of herbivores and fire on riparian and upland savanna ecosystems: field operations manual for herbivore and fire exclosures on the sabie and letaba rivers in the kruger national park, south african national parks, skukuza. pollastrini, m., desotgiu, r., cascio, c., bussotti, f., cherubini, p., saurer, m. et al., 2010, ‘growth and physiological responses to ozone and mild drought stress of tree species with different ecological requirements’, trees 24, 695−704. doi:10.1007/s00468-010-0439-4 renton, m., thornby, d. & hanan, j., 2007, ‘canonical modelling: an approach for intermediate-level simulation of carbon allocation in functional-structural plant models’, in j. vos, l.f.m. marcelis, p.h.b de visser, p.c. struik & j.b. evers (eds.), functional-structural plant modelling in crop production, pp. 151−164, springer, dordrecht. riginos, c. & young, t.p., 2007, ‘positive and negative effects of grass, cattle, and wild herbivores on acacia saplings in an east african savanna’, oecologia 153, 985−995. doi:10.1007/s00442-007-0799-7, pmid:17661089 rohner, c. & ward, d., 1999, ‘large mammalian herbivores and the conservation of arid acacia stands in the middle-east’, conservation biology 13, 1162−1171. doi:10.1046/j.1523-1739.1999.97300.x rooke, t. & bergström, r., 2007, ‘growth, chemical responses and herbivory after simulated leaf browsing in combretum apiculatum’, plant ecology 189, 201−212. doi:10.1007/s11258-006-9177-5 sankaran, m., ratnam, j. & hanan, n., 2008, ‘woody cover in african savannas: the role of resources, fire and herbivory’, global ecology & biogeography 17, 236–245. schmidt, e., lötter, m. & mccleland, w., 2007, trees and shrubs of mpumalanga and kruger national park, jacana media, johannesburg. scholes, r.j., 1997, ‘savanna’, in r.m. cowling, d.m. richardson & s.m. pierce (eds.), vegetation of southern africa, pp. 258−277, cambridge university press, cambridge. scholes, r.j., bond, w.j. & eckhardt, h.c., 2003, ‘vegetation dynamics in the kruger ecosystem’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 242−246, island press, washington dc. scogings, p.f., 1998, ‘resistance of six savanna woody plant species to browsing by goats in the eastern cape province of south africa’, phd thesis, department of livestock and pasture science, university of fort hare. scogings, p.f., johansson, t., hjältén, j. & kruger, j., in press, ‘woody vegetation and large herbivores in semi-arid savannas’, austral ecology. sharam, g., sinclair, a.r.e. & turkington, r., 2006, ‘establishment of broad-leaved thickets in serengeti, tanzania: the influence of fire, browsers, grass competition, and elephants’, biotropica 38, 599−605. doi:10.1111/j.1744-7429.2006.00195.x siebert, f. & eckhardt, h.c., 2007, ‘the vegetation and floristics of the nkuhlu exclosures, kruger national park’, koedoe 50, 126−144. skinner, j.d. & chimimba, c.t., 2005, the mammals of the southern african subregion, cambridge university press, cape town. spss, 2000, systat 10, spss inc., chicago. staver, a.c., bond, w.j., stock, w.d., van rensburg, s. & waldram, m.s., 2009, ‘browsing and fire interact to suppress tree density in an african savanna’, ecological applications 19, 1909−1919. doi:10.1890/08-1907.1, pmid:19831079 sweet, r.j. & mphinyane, w., 1986, ‘preliminary observations on the ability of goats to control post-burning regrowth in acacia nigrescens/combretum apiculatum savanna in botswana’, journal of the grassland society of southern africa 3, 79−84. tarasoff, c.s., mallory-smith, c.a. & ball, d.a., 2007, ‘comparative plant responses of puccinellia distans and puccinellia nuttalliana to sodic versus normal soil types’, journal of arid environments 70, 403−417. doi:10.1016/j.jaridenv.2007.01.008 teague, w.r., 1985, ‘leaf growth of acacia karroo trees in response to frequency and intensity of defoliation’ in j.c. tothill & j.j. mott jj (eds.), ecology and management of the world’s savannas, pp. 220−222, australian academy of science, canberra, and cab, farnham royal. teague, w.r., 1988, ‘effect of the intensity and phenophase of defoliation and water stress on the rate of photosynthesis and the recovery of carbohydrate reserves in acacia karroo hayne’, journal of the grassland society of southern africa 5, 223−226. teague, w.r. & walker, b.h., 1988, ‘effect of intensity of defoliation by goats at different phenophases on leaf and shoot growth of acacia karroo hayne’, journal of the grassland society of southern africa 5, 197−206. tsumele, j., mlambo, d. & sebata, a., 2006, ’responses of three acacia species to simulated herbivory in a semi-arid southern african savanna’, african journal of ecology 145, 325−327. underwood, a.j., 1997, experiments in ecology, cambridge university press, cambridge. williams, c.a., hanan, n., scholes, r.j. & kutsch, w., 2009, ‘complexity in water and carbon dioxide fluxes following rain pulses in an african savanna’, oecologia 161, 469−480. doi:10.1007/s00442-009-1405-y, pmid:19582479, pmid:2757614 abstract introduction materials and methods results discussion conclusion acknowledgements references about the author(s) lucas p. rutina okavango research institute, university of botswana, botswana kefentse m. mogwera okavango research institute, university of botswana, botswana elford seonyatseng research division, department of wildlife and national parks, botswana charles mpofu research division, department of wildlife and national parks, botswana ditso ntloyathuto okavango research institute, university of botswana, botswana citation rutina, l.p., mogwera, k.m., seonyatseng, e., mpofu, c. & ntloyathuto, d., 2017, ‘herders’ ecological knowledge and carnivore predation on livestock investigations in makgadikgadi and nxai national parks, botswana’, koedoe 59(2), a1389. https://doi.org/10.4102/koedoe.v59i2.1389 original research herders’ ecological knowledge and carnivore predation on livestock investigations in makgadikgadi and nxai national parks, botswana lucas p. rutina, kefentse m. mogwera, elford seonyatseng, charles mpofu, ditso ntloyathuto received: 08 mar. 2016; accepted: 02 may 2017; published: 24 july 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract botswana is one of the countries in southern africa that pay compensation for human properties damaged by wildlife. before compensation is paid, a thorough investigation on determining wildlife species that have caused the damage is mandatory. because of insufficient resources by the department of wildlife and national parks, the initial investigation is carried out by herders. three basic indicators are used to determine carnivore predation; sighting the carnivore at the kill, tracks of the predator and examining the carcasses. in this study, we tested herders’ knowledge on the above three indicators. the study was conducted in a communal area around makgadikgadi and nxai national parks, botswana, where the main activities practiced by the local communities is pastoral farming. in general, there was a significant association between reported and perceived incidents of predation for all carnivores at all distances from protected areas. herders were able to identify the large carnivores visually. but they had difficulties in identifying carnivore tracks and kill characteristics. the results demonstrate the importance of involvement of local communities in human–wildlife conflict management. however, more education regarding identification of carnivore tracks and kill behaviour is needed for herders in the study area. conservation implications: based on the results of this study, this calls for a change in the management of human–wildlife conflict (hwc) and administration of the compensation scheme. decentralising hwc to local communities using existing government structures that exist at local level will not only supplement the inadequate resources by the department of wildlife and national parks (dwnp) to effectively mitigate the problem, but also empower local communities’ participation in wildlife management. introduction management of human–carnivore conflict using expert-based science (ebs) has been proven to be ineffective, and the conflicts have continued to have socio-economic implications on the livelihoods of pastoralists residing near protected areas globally (goodrich et al. 2011; inskip & zimmerman 2009). in response, pastoralists developed retaliatory behaviour by killing (both legal and illegal) or poisoning carnivores (ogada et al. 2003), leading to the observed decline of carnivore populations. information on distribution and intensity of human–carnivore conflict is essential in managing such conflicts around the world, and obtaining adequate information to come up with effective and efficient strategies requires extensive data on wildlife populations which is often insufficient in technical ebs (gilchrist, mallory & merkel 2005).techniques involved in ebs are usually expensive and demanding in terms of inputs, thus complicating research (anadón et al. 2009). however, it has been indicated that information from local knowledge and skills of communities could be of high quality at a relatively low cost (hartwig 2009a). despite appeals to integrate this knowledge in scientific research to enhance natural resource management, such efforts are seldom put into practice (sutherland et al. 2013). there is a widespread recognition that people residing in an area for long periods develop important and established relationships with and knowledge about those areas (prado & murrieta 2015). various terminologies have been proposed to describe this knowledge, such as local ecological knowledge (lek) (anadón et al. 2009), traditional ecological knowledge (tek) (phuthego & chanda 2004) and indigenous knowledge (ik) (roba & oba 2009). although many authors use this terminology interchangeably, gilchrist et al. (2005) and neagh et al. (2008) suggested that a distinction that comprehends the time scale from which this knowledge and skills has been acquired is required. the distinctions being that tek and ik are more of knowledge and practices acquired through adaptive management, by an ethnic group through generations interacting with nature and is believed to be transferrable from one generation to another (neagh et al. 2008), while lek, on the contrary, describes the skills and practices gained and developed by an individual’s annotations over his or her periods of interaction with nature, and this knowledge is not transferred from one generation to another (gilchrist et al. 2005). in many studies, lek is associated with the perception or skills possessed by individuals given their periods of interaction with nature. in this study, we assessed the competences of lek possessed by livestock herders to investigate carnivore species that predate on livestock around makgadikgadi and nxai national parks. we define lek in this study as the skills possessed by herders by virtue of experience from duties linking them with livestock depredation issues on a daily basis, and herders as individuals that are employed to look after livestock throughout the year, and this includes self-employed farmers. herding activities included general husbandry of keeping livestock (taking livestock to pasture and water in the morning and bringing them back to the kraal [boma – a place where they are kept safe during the night]), looking for lost and unhealthy animals and reporting any problem concerning the keeping of the livestock to the owner (in case of employed herders). livestock farming in botswana is very important because majority of the rural communities depend on it as a source of income and protein (statistics botswana 2009; statistics botswana 2013). as such, livestock production, especially cattle, plays a critical socio-economic role for farmers in botswana. depredation of livestock by carnivores therefore poses economic loss to farmers. although the botswana wildlife conservation and national park act of 1992 permits lethal control of wildlife in defence for human property damaged, botswana is still one of the countries in southern africa that pay compensation for human properties damaged by wildlife as a measure to balance the trade-off between wildlife conservation and the impact of conflict they bring to the farming communities. before compensation is paid, a thorough investigation on determining the wildlife species that has caused damage is mandatory. because of insufficient resources by the department of wildlife and national parks (dwnp), the initial investigation is carried out by herders. necessary skills are needed to best find out which carnivore species is causing damage. three basic indicators that are used to determine carnivore predation are sighting the carnivore at the kill (hereafter referred to as visual), tracks of the predator (hereafter referred to as tracks) and examining the carcasses (bite marks, body parts eaten and location of the carcasses, hereafter referred to as kill patterns). because information on livestock raiding carnivores in botswana is currently insufficient (hemson 2003; valeix et al. 2012), this study used lek possessed by herders to assess herders’ ability to identify the carnivore species causing damage using the three indicators mentioned above and mapped their perceptions on magnitude and spatiotemporal patterns of large carnivore predation on livestock. one of the significant aspects of using lek is to enhance the contribution of local people in policy decision-making in order to promote their participation in proper management of natural resources around them (calamia 1999). the objectives of this study were to (1) compare carnivore predation reported to and confirmed by the dwnp (hereafter refer to as pac) with reports made during interviews (hereafter referred to as lek) for lion, leopard and wild dog; (2) compare the carnivore predation risk at cattle post (an area where the main activity is livestock farming) as determined by data collected by pac and lek and (3) assess herders’ ability to identify carnivore species from visual photos, tracks and kill patterns. materials and methods study area the study was conducted in a communal area (ct 7) covering an area approximately 6543 km2 in the eastern side of makgadikgadi (ng 52 and ct 9) and nxai (ng 50) pans national parks, south of a wildlife management area (ct 1 and 2) and north of wildlife management area (ct 11), botswana (figure 1). a wildlife management area is an area designated for wildlife management, including sustainable utilisation of wildlife. the study area encompasses numerous cattle posts that practices subsistence mixed farming (arable and pastoral), with pastoral farming being the dominant farming type in the area. human population in the area and surroundings has increased significantly, by almost 50% per annum and on average each cattle post had 10–20 members (cso 2011). the study site composed of 63 cattle posts of which 9 (14.2%) were within 10 km from protected areas (near) and 54 (85.8%) were beyond 10 km from protected areas (far). the study site falls within a semi-arid zone, lying between the 250 mm and 450 mm rainfall isohyets, with rainfall predominately between november and april (shaw et al. 1997). however, during the study period, annual rainfall ranged from 338 mm to 412 mm (statistic botswana 2015). dominant tree species in the area includes baobab (adansonia digitata), some of which serve as land marks; leadwood (combretum imberbe), silver terminalia (terminalia sericea), mopane (colophospermum mopane) and real fan palm (hyphaene petersiana) (brooks & maude 2010). most of wildlife species are residents inside the parks, but several species seasonally migrate in and out in response to fluctuation of resources, especially surface water and forage. herbivore species include zebra (equus quagga), springbok (antidorcas marsupialis), blue wildebeest (connochaetes taurinus) and gemsbok (oryx gazella) (dwnp 2012; kgathi & kalikawe 1993). large carnivores include the african lion (panthera leo), leopard (panthera pardus), spotted hyena (crocuta crocuta), cheetah (acinonyx jubatus) and african wild dog (lycaon pictus) (hemson 2003; rutina et al. 2015; valeix et al. 2012). figure 1: map of the study area showing the communal area (ct 7) in relation to protected areas (national parks [nxai and makgadikgadi] and wildlife management areas [ct 1, ct 2 and ct 11]) and the three section division as used to group herders during the workshop. section a is gweta north, b is gweta south and c is nogatshaa. in the last century, the basarwa people were the dominant people in the area, depending mainly on hunting and gathering of wild resources (ferrar 1995). however, around the 1930s when technology for accessing groundwater in the area improved, other tribes and their livestock settled in the area. the arrival of these other ethnic groups, mainly bakalanga, banabiya and bangwato, in the area diluted the basarwa tek and ik. according to dwnp (dwnp, personal communication), the eastern boundary of makgadikgadi pans national park has been a centre of controversy since its proclamation in 1971 because people believed it was moved closer to them, thus taking their grazing land and crop fields. this has led to pastoralist’s argument that the eastern park boundary is too close to them. in addition, this has led to cattle moving and grazing in protected areas and wildlife management. the erection of the carnivore proof fence in the western boundary of the park, along the boteti river could have shifted wildlife movements towards the study area especially during the wet season when water is available. data collection and analysis incidents of carnivore predation on livestock data on incidents of carnivore predation on livestock were obtained from the dwnp. the incidents are investigated and documented by the department’s problem animal control (pac) unit for the purpose of financially compensating for damage by species that are considered to be too dangerous for farmers to confront or chase away from their property, or those that are considered to be of high protection priority. as such the records enable research on the spatial and temporal dynamics of human–wildlife conflict (hwc) in the country (schiess-meier et al. 2007). details involving type and number of livestock killed, predator species involved, date and place of attacks (name of cattle post) were captured. five years’ pac data from 2008 to 2012 were considered for analysis on the premise that more data improve accuracy of results. the reports lacked precise locations such as global positioning system (gps) coordinates of the actual location of damage, thereby compelling data to be analysed at the cattle post level. cattle posts were visited with the help of local community assistants, and their point locations were recorded using a handheld gps device. herders’ local ecological knowledge because it is important to gather data from people who have lived in the area long enough, and who have been closely associated with issues concerning livestock depredations, 62 herders, 51 (82%) from cattle posts far from protected areas and 11 (18%) from cattle posts near protected areas were randomly selected from the list of farmers who reported carnivore predation on their livestock to the dwnp. livestock herders (e.g. employed or owners) were targeted because they relatively spend more time with livestock. according to roba and oba (2009), livestock herder indicators are appropriate for pastoral farming because they are closely connected to human livelihoods as opposed to ecologists. a semi-structured questionnaire survey was conducted on the selected herders. the main objective of these interviews was to gain information regarding the herders’ understanding of wildlife–livestock interaction, with a focus on carnivore interactions with livestock (predation) and other human activities. specific objective was to assess how long the herders have been in the area, how many livestock did they lose because of each of the three carnivores (lion, leopard and wild dog) and how do they perceive predation risk (measured in terms of livestock lost to each carnivore per month). out of the 62 herders interviewed, 40 (65%) were self-employed (owners) and 22 (35%) were employed herders. all employed herders were males ranging between 25 and 40 years of age. of the self-employed herders, 18 (46%) were females and 22 (54%) were males. out of the 62 herders interviewed, 30 (48%), 10 females and 20 males were invited for a workshop to map carnivore predation hotspots. selection was based on their experience and period stayed in the area. the selection also considered the spatial distribution of the cattle posts. the study area was divided into three sections (northern, central and southern). ten herders were selected from each section. working in their sections, participants provided information based on their own experience or knowledge gained from other community residents about carnivore predation on livestock risk per each cattle post in their section. all information was recorded on the paper maps provided, by categorising cattle posts into one of the three risk categories (low = 1, medium = 2 and high = 3) for each carnivore. risk categories were based on the results from the interviews. herders considered predation incidents of zero to three per month as low risk, four to six as moderate and greater than six as high. to assess ecological knowledge of herders to identify carnivores using the three indicators used in compensation investigations, the 30 herders invited to the workshop were shown picture cards of carnivores and were asked to identify them (figure 2). similarly, herders were asked to identify tracks and examine carcasses from picture cards (figure 2). figure 2: examples of cards used to assess herders’ skills to identify carnivores. (a) visual, (b) track and (c) kill pattern. data analysis we used χ2 test to know whether there was a significant association between data obtained from confirmed predation incidents and data obtained from interviews, given the distance from protected areas and carnivore species. we also used the χ2 test to determine whether there were significant differences between the frequencies of herders who identified carnivores correctly to those who did not. kruskal–wallis test with dunn–bonferroni post hoc test was performed to test the hypothesis that risk categories of the three carnivores from lek and pac were the same across the distances from protected areas. we further calculated conditional probability that both lek and pac data will place a cattle post in the same category. the conditional probability was calculated as the chances that both lek and pac will place a cattle post in the same category divided by the total possible outcomes. finally, we use frequencies (percentages) to assess the ability of herders to identify carnivore species using the three indicators used in carnivore predation investigations by dwnp. results incidents of carnivore predation on livestock comparing data obtained from confirmed predation incidents and data obtained from interviews, there was no significant association between pac and lek incidence for all carnivores at all distances from protected areas (χ2 test, p > 0.05 for all tests). both pac and lek showed that more than 60%, 75% and 80% of the cattle posts in the study area experienced low predation by lion, leopard and wild dog, respectively. however, the association between pac and lek for all the carnivores was distance specific. lion pac incidents for cattle posts near protected areas were significantly higher than lek incidents, while there was no significant difference between pac and lek for cattle posts far from protected areas (figure 3a). for leopard and wild dog, lek incidents were higher than pac incidents for all the distance from protected areas (figure 3b and 3c, respectively). although no pac reports were available in the study area – because the species is not compensated, lek incidents showed that spotted hyena caused more damage than the three reported species, especially on cattle posts far from protected areas (figure 4). figure 3: comparison of carnivore predation on livestock incidents between department of wildlife and national parks–confirmed incidents (problem animal control) and herders interviewed (local ecological knowledge). (a) lion, (b) leopard and (c) wild dog. figure 4: comparison of carnivore predation in livestock risk areas between department of wildlife and national parks–confirmed incidents (problem animal control) and herders interviewed (local ecological knowledge). (a) lion, (b) leopard and (c) wild dog. carnivore predation risk after standardising reports to predation risk categories, a kruskal–wallis test showed that there was a statistically significant difference in lion predation risk categories between pac and lek for cattle posts beyond 10 km from protected areas (far) [χ2 (1) = 14.591, p < 0.001] (figure 4a). the dunn-bonferroni post hoc test indicated that the difference was significant for cattle posts near from protected areas (test statistic = 14.44, p = 0.004) but no significant difference was found for cattle posts far protected areas (test statistic = -9.11, p = 0.453) (figure 4a). both leopard and wild dog predation risk categories between pac and lek for all cattle posts were not statistically different (kruskal–wallis test, p > 0.05). predation risk categories between pac and lek for cattle posts beyond 10 km from protected areas were not statistically different (kruskal–wallis test, p > 0.05) for all three carnivores (figure 4a, b and c). the probability that both pac and lek will categorise a cattle post in the same predation risk category for lion was 70% and 56% for cattle posts far and near protected areas, respectively. for leopard, the probability that both pac and lek will categorise a cattle post in the same predation risk category was 67% and 56% for cattle posts far and near protected areas, respectively. for wild dog, the probability that both pac and lek will categorise a cattle post in the same predation risk category was 63% and 78% for cattle posts far and near protected areas, respectively. identification of carnivore species in general, herders were able to identify the large carnivores visually: near – 76% and far – 75% (figure 5). by contrast, herders had difficulties in identifying carnivore tracks and kill characteristics (tracks: near = 35%, far = 63%; kill characteristics: near = 35%, far = 34%, figure 5). results from the investigation diaries suggest that for all the cases where herders used their ecological knowledge to identify the carnivore that had caused the damage in 82%, 67% and 100% of the cases are approved by dwnp for lion, wild dog and leopard, respectively. figure 5: percentage of herders who identified carnivore species correctly from cards. (a) cheetah, (b) leopard, (c) lion, (d) spotted hyena and (e) wild dog. discussion although lek is not commonly used in mapping the spatial and temporal patterns of human–carnivore conflict, the results of this study show that confirmed carnivore predation incidents and reports gathered through interviewing herders did not significantly differ for all the three carnivores and distances from protected areas. this shows that carnivore predation information collected through lek and the standard protocols used by dwnp were complementary. similar results have been observed with other species such as tortoise distribution in south-eastern spain (anadón et al. 2009), fish and fisheries management in the beaufort sea, canada (hartwig 2009b), southern mexico (espinoza-tenorio et al. 2013), south-eastern brazil (teixeira et al. 2013) and wild mammals in northern gonarezhou national park, zimbabwe (gandiwa 2012). the results further show that both lek and pac data were in agreement over the fact that lion predation decreases with increasing distance from protected areas (figure 3a) and leopard incidents were evenly distributed throughout the study area (figure 3b). contrary to the other two carnivores, there was disagreement between lek and pac data on the distribution of wild dog predation incidents in the study area, with pac data indicating that wild dog predation incidents were evenly distributed and lek indicating that wild dog predation incidents increased with decreasing distance from protected areas. the results are partly comparable to the findings that stronger carnivores such as lion are more problematic in the periphery of protected areas, while outcompeted smaller ones are forced to forage deeper into human-dominated landscapes (gusset et al. 2009; mogwera & rutina 2015; rutina et al. 2015). it is also apparent that during the dry season, carnivore problems are more pronounced, particularly because the preferred wild prey has migrated to their dry season habitats, thus leaving livestock in high abundance and as alternative prey (valeix et al. 2012). in general, lek and pac were in agreement in categorising cattle posts for carnivore predation risk areas. in addition, the study shows that the chances that lek predictions for carnivore predation risk to be similar to confirmed incidents were high for both the three carnivore at all distances from protected areas. this is consistent with other studies that showed lek information complement scientific information in mapping environmental hazards (anh, phong & mulenga 2014; raungratanaamporn & pakdeeburee 2014; udmale, ichikawa & manandhar 2014), fisheries (silvano & begossi 2012), natural resource management (gilchrist et al. 2005; ruddle & calamia 1999), environmental planning (kujala et al. 2013), natural resources research (anadón et al. 2009; gandiwa 2012; silvano & begossi 2012) and hwc hotspots (calamia 1999; hartwig 2009a; rutina et al. 2015; strindberg et al. 2007). this significant contribution of lek to map carnivore predation areas indicates the potential impotence of lek in hwc management in the region. we expected that that herders, having grown up with wildlife could easily identify carnivores visually and their tracks. however, we found that herders can identify large carnivore species visually but had difficulties in identifying carnivore tracks and kills patterns. this could be attributed to the current herding practises that are carried in the study area. gontse et al. (2012) found that because of the paying schedule from the department of labour affairs, farmers are not able to pay herders for all the activities involved in the practice of herding. as such, herders are now taking livestock to water points and kraal that arrive back during the afternoon (gontse et al. 2012; rutina et al. 2015). contrary to the results of this study, information from the dwnp investigation diaries suggests confirmed 82% of lion-related incidents, 67% of wild dog-related incidents and 100% of leopard-related incidents identified by herders who used their ecological knowledge. furthermore, data from diaries suggest that herders use tracks and/or patterns of kill to identify carnivores causing damage. as we have shown above, herders had difficulties in identifying carnivore tracks and kill patterns (figure 5). we thus recommend that more education regarding identification of carnivore tracks and kill behaviour is needed for herders in the study area. nevertheless, the results of the study show that most of the carnivore predation on livestock information was from lek (e.g. number of cases [figure 3], risk areas [figure 4] and visual identification of carnivores [figure 5]) mentioned by the herders corresponded with information obtained from scientifically recognised methods. lack of resources by the dwnp to mitigate and attend to carnivore predation on livestock in the study area have been suggested as one factor contributing to human–carnivore conflict in the study area (gontse et al. 2012; hemson 2003; hemson et al. 2009; rutina et al. 2015; valeix et al. 2012). because of the inadequate resources by dwnp, most of hwc mitigation measures are conducted at a large scale and use a ‘one size fits all’ strategy. despite different opinions among scientists and policy-makers as to the use of lek in natural resource management (see mclain et al. 2013) and given the complexity and nature of hwc, mitigation strategies need to be area and species specific. the difficulties are mostly because of lack of mutual understanding among stakeholders, including local communities, towards wildlife management approaches, and this undermines the effectiveness of integrating lek in hwc. coordination among government, communities and scientists has been suggested as the best strategy to manage natural resources (espinoza-tenorio et al. 2013; gilchrist et al. 2005; mclain et al. 2013; neagh et al. 2008; phuthego & chanda 2004). based on the results of this study, we argue that decentralising hwc to local communities using existing government structures that exist at local level, such as district conservation committees, village development committees and village farmer committees will improve the problem. this will not only supplement the inadequate resources by the dwnp to effectively mitigate the problem, but also empower local communities in wildlife management and tap on the lek the community have. the implementation of community-based natural resources management goes along training communities to collect data for monitoring purposes and inform government strategies using a management-oriented monitoring system (moms). the use of moms in carnivore predation investigations will significantly improve information on management of human–carnivore conflict. this should include the recognition that in order to increase the effectiveness of reducing or mitigating carnivore predation on livestock, communities should play important roles by applying their lek understanding of the spatiotemporal dynamics of carnivore predation rather than focusing on increasing dwnp capability to mitigate hwc. over the past 10 years, southern african governments have been engaged in creating transfrontier conservation areas, consisting of a mosaic of different land uses. within this mosaic agricultural landscapes play an important role as habitat and corridors for wide-ranging wildlife, such as large carnivores, because the sizes of most protected wildlife areas are not large enough to adequately accommodate their home ranges. our study site forms part of the kavango–zambezi transfrontier conservation area (kaza tfca), a conservation area covering part of angola, botswana, namibia, zambia and zimbabwe. both countries, through revision of their national biodiversity strategy and action plans (nbsaps) to align them with cbd 2015 goals and 2020 aichi targets, recognise that hwc hinders the promotion of integration between biodiversity conservation and human well-being and have prioritised hwc as priority areas that need to be addressed in their integrated development plans (idps) for their kaza tfca components. the results of our study suggest that involving local communities in managing hwc could help the kaza tfca mandate to be realised. conclusion the results of the study showed that both pac and lek information were complementary in reporting incidents of carnivore predation on livestock and categorising cattle posts for carnivore predation risk for the three carnivores studied. this calls for involvement of local communities in carnivore predation investigations and management. this will not only complement the shortage resources by government to manage conflict, but also reduce the spatial scale at which conflict management is conducted. this can be done by decentralising hwc management to local communities using government structures that exist at local level. however, herders had difficulties in identifying carnivore tracks and kill patterns. this could be due to partly poor pictures used because information from the dwnp investigation diaries suggests that almost all confirmed incidents were identified by herders using tracks and/or patterns of kill. acknowledgements we acknowledge the southern africa science service centre for climate change and adaptive land-use (sasscal) and the university of botswana internal funding for supporting this project. the department of wildlife and national parks (serowe and nata offices) provided reported and confirmed data used in this study. gaoratwe and lungisane from dwnp nata office helped the research team in identifying cattleposts. thebe, sekotswe and toro, research assistants from the okavango research institute, and makgetho and moathodi, community assistants, helped the researcher in field data collection. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions l.p.r. was the principal investigator for the project, designed the project objectives, coordinated herders’ workshops, performed data analysis and write up. k.m.m. and d.n. conducted herders’ interviews, led group discussions during the herders’ workshops and contributed to the write up. e.s. and c.m. collected, collated and analysed carnivore predation on livestock reports from dwnp, participated in the herders’ workshops and write up. references anadón, j.d., giménez, a., ballestar, r. & pérez, i., 2009, ‘evaluation of local ecological knowledge as a method for collecting extensive data on animal abundance’, conservation biology: the journal of the society for conservation biology 23(3), 617–625. https://doi.org/10.1111/j.1523-1739.2008.01145.x anh, t.t., phong, t.v.g. & mulenga, m., 2014, ‘community consultation for climate resilient housing: a comparative case study in vietnam’, international journal of disaster risk reduction 10(pa), 201–212. https://doi.org/10.1016/j.ijdrr.2014.09.012 brooks, c. & maude, g., 2010, ‘wildlife resources and human wildlife conflict’, in center for applied resesearch (eds.), makgadikgadi framework management plan, p. 161, department of environmental affairs, botswana. calamia, m., 1999, ‘a methodology for incorporating traditional ecological knowledge with geographic information systems for marine resource management in the pacific’, resource management and knowledge information 16, 2–12. dwnp, 2012, aerial census of animals in botswana: 2012 dry. season, department of wildlife and national parks, gaborone. espinoza-tenorio, a., wolff, m., espejel, i. & montaño-moctezuma, g., 2013, ‘using traditional ecological knowledge to improve holistic fisheries management: transdisciplinary modeling of a lagoon ecosystem of southern mexico’, ecology and society 18(2), (6), 1–18 https://doi.org/10.5751/es-05369-180206 ferrar, t., 1995, makgadikgadi/nxai pan management plan, department of wildlife and national parks, gaborone. gandiwa, e., 2012, ‘local knowledge and perceptions of animal population abundances by communities adjacent to the northern gonarezhou national park, zimbabwe’, tropical conservation science 5(3) 255–269. https://doi.org/10.1177/194008291200500303 gilchrist, g., mallory, m. & merkel, f., 2005, ‘can local ecological knowledge contribute to wildlife management? case studies of migratory birds’, ecology and society 10(1), 12. https://doi.org/10.5751/es-01275-100120 gontse, k., mogwera, k., mmusi, m. & mhale, k., 2012, an investigatiojn of factors influencing persistance of human-carnivore conflict in western tutume sub-district, botswana, winter course report, university of botswana, maun. goodrich, j.m., seryodkin, i., miquelle, d.g. & bereznuk, s.l., 2011. conflicts between amur (siberian) tigers and humans in the russian far east. biological conservation 144, 584–592. gusset, m., swarner, m.j., mponwane, l., keletile, k. & mcnutt, j.w., 2009, ‘human–wildlife conflict in northern botswana: livestock predation by endangered african wild dog lycaon pictus and other carnivores’, oryx 43(1), 67–72. https://doi.org/10.1017/s0030605308990475 hartwig, l., 2009a, mapping traditional knowledge related to the identification of ecologically and biologically significant areas in the beaufort sea, canadian manuscript report of fisheries and aquatic sciences. hartwig, l., 2009b, ‘mapping traditional knowledge related to the identification of ecologically and biologically significant areas in the beaufort sea’, canadian manuscript report of fisheries and aquatic sciences 2895, 31. hemson, g., 2003, ‘the ecology and conservation of lions: human-wildlife conflict in semi-arid botswana’, phd thesis. lady magret hall, univrsity of oxford. hemson, g., maclennan, s., mills, g., johnson, p. & macdonald, d., 2009, ‘community, lions, livestock and money: a spatial and social analysis of attitudes to wildlife and the conservation value of tourism in a human-carnivore conflict in botswana’, biological conservation 142, 2718–2725. https://doi.org/10.1016/j.biocon.2009.06.024 inskip, c. & zimmermann, a., 2009, ‘human-felid conflict: a review of patterns and priorities worldwide’, oryx 43(1), 18–34. kgathi, d.k. & kalikawe, m.c., 1993, ‘seasonal distribution of zebra and wildebeest in makgadikgadi pans game reserve, botswana’, african journal of ecology 31, 210–219. https://doi.org/10.1111/j.1365-2028.1993.tb00534.x kujala, h., moilanen, a., araújo, m.b. & cabeza, m., 2013, ‘conservation planning with uncertain climate change projections’, plos one 8(2), 1–12. https://doi.org/10.1371/journal.pone.0053315 mclain, r., poe, m., cerveny, l., besser, d. & blahna, d., 2013, ‘making sense of human ecology mapping: an overview of approaches to integrating socio-spatial data into environmental planning’, human ecology 41(1), 17. https://doi.org/10.1007/s10745-013-9573-0 mogwera, k. & rutina, l., 2015, ‘predicting spatio-temporal distribution of human-carnivore conflict in response to environmental factors in makgadikgadi agro-ecosystem, northern botswana’, in l. rutina, m. nthomiwa & d. ntloyathuto (eds.), botswana symposium on wetlands and wildlife 2015, pp. 160–171, university of botswana, maun. neagh, l., ireland, n., quinn, r.j., donnelly, d.j. & cooper, j.a.g., 2008, ‘accurate mental maps as an aspect of local ecological knowledge (lek): a case study from’, ecology and society 13(1), 24. ogada, m.o., woodroffe, r., oguge, n.o. & frank, l.g., 2003, ‘limiting depredation by african carnivores: the role of livestock husbandry’, conservation biology 17, 1521–1530. https://doi.org/10.1111/j.1523-1739.2003.00061.x phuthego, t. & chanda, r., 2004, ‘traditional ecological knowledge and community-based natural resource management: lessons from a botswana wildlife management area’, applied geography 24(1), 57–76. https://doi.org/10.1016/j.apgeog.2003.10.001 prado, h.m. & murrieta, r.s., 2015, ‘ethnology in perception: the origins, interfaces and current trends of a growing field’, sao paulo xviii(4), 133–154. raungratanaamporn, i. & pakdeeburee, p., 2014, ‘government-communities collaboration in disaster management activity : investigation in the current flood disaster management policy in thailand’, procedia environmental sciences 20, 658–667. https://doi.org/10.1016/j.proenv.2014.03.079 roba, h.g. & oba, g., 2009, ‘community participatory landscape classification and biodiversity assessment and monitoring of grazing lands in northern kenya’, journal of environmental management 90, 673–682. https://doi.org/10.1016/j.jenvman.2007.12.017 ruddle, k. & calamia, m.a., 1999, ‘a methodology for incorporating traditional ecological knowledge with geographic information systems for marine resource management in the pacific, spc traditional marine resource management and knowledge information bulletin 10, 2–12. rutina, l., seonyatseng, e., mpofu, c. & mogwera, k., 2015, interactive influence of carnivore foraging behavior and farmers’ perception on human carnivore conflict in western tutume sub-district, botswana, technical report, okavango research institute, university of botswana, maun. schiess-meier, m., ramsauer, s., gabanapelo, t. & könig, b., 2007, ‘livestock predation – insights from problem animal control registers in botswana’, journal of wildlife management 71(4), 1267–1274. https://doi.org/10.2193/2006-177 shaw, p.a., stokes, s., thomas, d.s.g., davies, f.b.m. & holmgren, k., 1997, ‘palaeoecology and age of a quaternary high lake level in the makgadikgadi basin of the middle kalahari, botswana’, south african journal of science 93(6), 273–276. silvano, r.a.m. & begossi, a., 2012, ‘fishermen’s local ecological knowledge on southeastern brazilian coastal fishes: contributions to research, conservation, and management’, neotropical ichthyology 10(1), 133–147. https://doi.org/10.1590/s1679-62252012000100013 statistics botswana, 2009, population census, botswana government printers, gaborone. statistics botswana, 2012, agricltural sector report, botswana government printers, gaborone. statistics botswana, 2013, agricltural sector report, botswana government printers, gaborone. statistics botswana, 2015, agricltural sector report, botswana government printers, gaborone. strindberg, s., johnson, a.a.h., hallam, c., rasphone, a., van der helm, f., xiongyiadang, p. & sisavath, p., 2007, ‘recommendations for monitoring landscape species in the nam kading national protected area’, wildlife conservation society (wcs) and the integrated ecosystem and wildlife management project(iewmp) january-20(october 2015), 1–59. sutherland, w.j., gardner, t.a., haider, l.j. & dicks, l.v., 2013, ‘how can local and traditional knowledge be effectively incorporated into international assessments?’, oryx 48(1), 1–2. https://doi.org/10.1017/s0030605313001543 teixeira, j.b., martins, a.s., pinheiro, h.t., secchin, n.a., leão de moura, r. & bastos, a.c., 2013, ‘traditional ecological knowledge and the mapping of benthic marine habitats’, journal of environmental management 115, 241–250. https://doi.org/10.1016/j.jenvman.2012.11.020 udmale, p., ichikawa, y. & manandhar, s., 2014, ‘farmers â€tm perception of drought impacts, local adaptation and administrative mitigation measures in maharashtra’, international journal of disaster risk reduction 10, 250–269. https://doi.org/10.1016/j.ijdrr.2014.09.011 valeix, m., hemson, g., loveridge, a.j., mills, g. & macdonald, d.w., 2012, ‘behavioural adjustments of a large carnivore to access secondary prey in a human-dominated landscape’, journal of applied ecology 49, 73–81. https://doi.org/10.1111/j.1365-2664.2011.02099.x article information authors: benjamin j. wigley1,2 herve fritz2 corli coetsee1 william j. bond3 affiliations: 1school of natural resource management, nelson mandela metropolitan university, george campus, south africa2umr cnrs 5558 lbbe, university claude bernard lyon 1, france 3department of biological sciences, university of cape town, south africa correspondence to: benjamin wigley postal address: private bag x6531, george 6530, south africa dates: received: 07 may 2013 accepted: 18 sept. 2013 published: 11 mar. 2014 how to cite this article: wigley, b.j., fritz, h., coetsee, c. & bond, w.j., 2014, ‘herbivores shape woody plant communities in the kruger national park: lessons from three long-term exclosures’, koedoe 56(1), art. #1165, 12 pages. http://dx.doi.org/10.4102/ koedoe.v56i1.1165 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. herbivores shape woody plant communities in the kruger national park: lessons from three long-term exclosures in this original research... open access • abstract • introduction • research method and design    • study sites    • community composition    • trait sampling    • soil sampling    • statistical analyses • results    • community composition data    • population structure    • leaf traits of unique inside species versus common outside species    • chemical defence    • soil nutrients    • multivariate analyses • discussion • conclusion • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ the role of grazers in determining vegetation community compositions and structuring plant communities is well recognised in grassy systems. the role of browsers in affecting savanna woody plant communities is less clear. we used three long-term exclosures in the kruger national park to determine the effect of browsers on species compositions and population structures of woody communities. species assemblages, plant traits relating to browsing and soil nutrients were compared inside and outside of the exclosures. our results showed that browsers directly impact plant species distributions, densities and population structures by actively selecting for species with traits which make them desirable to browsers. species with high leaf nitrogen, low total phenolic content and low acid detergent lignin appeared to be favoured by herbivores and therefore tend to be rare outside of the exclosures. this study also suggested that browsers have important indirect effects on savanna functioning, as the reduction of woody cover can result in less litter of lower quality, which in turn can result in lower soil fertility. however, the magnitude of browser effects appeared to depend on inherent soil fertility and climate. conservation implications: browsers were shown to have significant impacts on plant communities. they have noticeable effects on local species diversity and population structure, as well as soil nutrients. these impacts are shown to be related to the underlying geology and climate. the effects of browsers on woody communities were shown to be greater in low rainfall, fertile areas compared to high rainfall, infertile soils. introduction top ↑ the role of herbivory in structuring plant communities and determining community compositions is well recognised (see reviews by augustine & mcnaughton 1998; huntly 1991). for example, native ungulate browsers have been found to have major impacts on shrub dynamics in east african savannas (augustine & mcnaughton 2004). different feeding behaviours and food preferences of browser species have major impacts on south african (owen-smith & cooper 1987) and west african savanna woody communities (jachmann & croes 1991). these impacts are often attributed to selective feeding related to food quality (owen-smith & cooper 1987). as in savannas (e.g. staver et al. 2009), herbivores have been found to regulate recruitment rate and species composition of trees in temperate forests (ammer 1996; kriebitzsch et al. 2000; van hees, kuiters & slim 1996) and grassland (anderson & briske 1995).many of the studies on browser–plant interactions in savannas have tended to focus on interactions between browsers and specific woody species. for example, coetzee et al. (2008), gadd (2002) and helm and witkowski (2012) looked at browser interactions with sclerocarya birrea, whilst fornara and du toit (2007) documented the response of acacia nigrescens to ungulate browsing. these two species are important browse species in southern african savannas. as savanna woody species tend to be highly variable in form and function, these studies usually do not provide a better understanding of savanna dynamics at the community level. augustine and mcnaughton (2004) emphasised the need for replicated experiments that assess the effects of native browsers on shrub dynamics in african savannas, as our understanding of the importance of browser effects on woody plant dynamics remains unclear. a better understanding of how browsers and woody plants interact at the community level and how these interactions affect savanna dynamics will be invaluable in managing browser populations. the existence of three long-term herbivore exclosures in the kruger national park, south africa, provides a good opportunity to empirically determine the role of browsers in determining woody species composition and population structures of woody communities growing at the three study sites. previous studies have shown evidence for taller canopies and higher tree and shrub densities inside exclosures at nhlangwini (asner et al. 2009; levick et al. 2010; trollope et al. 1998), makhohlola (levick et al. 2009; trollope et al. 1998) and n’waxitshumbe (asner et al. 2009; levick & rogers 2008; trollope et al. 1998). in these studies, the analyses were generally based on remote sensing and ignored species identity. differences in woody community compositions and plant traits between exclosure treatments are therefore still largely undocumented. differences in species compositions between treatments would be highly informative as the species found inside but not outside, are likely to be vulnerable to herbivory and therefore no longer found outside of the exclosures or found at reduced densities. the traits of these species can be used to provide us with clues as to what makes them vulnerable to herbivory. for example, díaz et al. (2007) found that at the global scale there is a spectrum of plant responses to herbivory with high levels of herbivory favouring annual over perennial plants, short plants over tall plants, prostrate over erect plants and stoloniferous plants over those with tussock architecture. this article aims to determine how browsers and plant traits interact and how these interactions affect woody plant population dynamics and community assemblages. we aim to determine if there are differences in species compositions and key plant functional traits between the woody communities found growing inside and outside the three long-term exclosures in the kruger national park (knp). plant traits are important in determining both the type of and degree of vertebrate herbivory, whilst they also can reflect plant responses to herbivory. we measured a number of plant traits that are likely to impact and respond to browsing; these included leaf nitrogen (n) and phosphorus (p), leaf carbon–nitrogen ratio (c:n), specific leaf area (sla), leaf size, leaf dry matter content (ldmc), leaf tensile strength, bite size index (bsi) and levels of secondary compounds. high leaf n and leaf p are associated with higher nutritional quality for herbivores (cornelissen et al. 2003). leaves with lower c:n ratios are much more attractive to herbivores as these leaves have higher n content (cornelissen et al. 2003). the sla of a species is usually a good correlate of its potential growth rate. lower sla values tend to correspond with relatively high investments in defences, both structural and chemical, and long leaf life span, all of which affect nutritional value (cornelissen et al. 2003). environmental nutrient stress and disturbance, as well as phylogenetic factors, play a role in determining leaf size (cornelissen et al. 2003). smaller leaves with higher sla are typically more attractive to herbivores. leaves with high ldmc values tend to be tough and therefore more resistant to disturbances such as herbivory or wind than leaves with low ldmc. species with low ldmc are usually associated with productive and highly disturbed areas (cornelissen et al. 2003). leaf tensile strength is a good indicator of the relative carbon investment in structural protection of the photosynthetic tissue. physically stronger leaves are better protected from abiotic (winds, storms) and biotic mechanical damage (herbivory). higher physical strength usually results in longer leaf span; however, this is usually combined with other physical and chemical defences such spines and tannins (cornelissen et al. 2003). bsi is a measure of how easily leaves can be browsed by a simulated browser (a human), whilst stem specific density broadly trades off against relative growth rate and stem defences against pathogens, herbivores and mechanical damage (cornelissen et al. 2003). condensed tannins and polyphenols are organic n-free chemical defences that bind with protein, reducing n availability to herbivores, thereby decreasing preference by mammal herbivores (bergvall & leimar 2005; haslam 1988; owen-smith 2005). the concentrations of the cell wall constituents (neutral detergent fibre [ndf], acid detergent fibre [adf] and acid detergent lignin [adl]) determine the digestibility of leaf material for herbivores. detergent analysis has been used to compare fibre digestibility across different plant types (e.g. see codron et al. 2007 and references therein). the following questions are specifically addressed using the three exclosures. firstly, has the long-term exclusion (> 40 years) of herbivores led to differences in woody plant species compositions at the three exclosures? secondly, do the structures, densities and abundances of the woody species populations differ inside and outside of the exclosures? thirdly, are there differences in plant leaf and stem traits between the species only found inside versus common species found both inside and outside? fourthly, has the long-term removal of herbivores from the exclosures had any effects on soil nutrient cycling? fifthly, do the exclosures help us to predict under what conditions herbivores play an important role in determining the structure and composition of woody plant communities in savannas? research method and design top ↑ study sites three long-term herbivore exclosures (> 40 years old) exist in the knp. these include the nhlangwini exclosure near pretoriuskop, the makhohlola exclosure near crocodile bridge and the n’waxitshumbe exclosure on the northern plains (table 1). at each exclosure two treatments were sampled, one inside the exclosure and one outside the exclosure at close proximity. table 1: site names, locations and descriptions of the exclosures in the kruger national park where sampling took place. the 220.0 ha nhlangwini exclosure was established in 1973 and occurs in the south-western portion of the knp near pretoriuskop rest camp. this is the wettest part of knp with mean average precipitation close to 750 mm. the exclosure is situated within the broad-leaved bushveld vegetation type that occurs on sandy soils derived from granitic rocks, common species include terminalia sericea and s. birrea (venter, scholes & eckhardt 2003). mean fire return interval is similar inside and out, with fire occurring on average every 3.6 years inside (asner et al. 2009) and less than every 4 years outside (smit et al. 2012). the makhohlola exclosure is a 2.4 ha exclosure constructed in the early 1970s. it is located in the south-eastern corner of the knp, just north of crocodile bridge. it occurs in the s. birrea–a. nigrescens savanna type (venter et al. 2003). the site occurs on soils derived from rocks of basaltic origin with mean annual rainfall of c. 600 mm. the mean fire return interval in the makhohlola exclosure (4.5 years) was slightly longer than outside (3.2 years), until 2004 when a new fire experiment was set up (see levick et al. 2009). since 2004, half of the exclosure has been protected from fire, whilst the other half is burnt in conjunction with the surrounding landscape (levick et al. 2009). from 2004 to the time of measurements in march 2011, the burn treatment and surrounding areas were burnt twice: in 2006 and 2010. the n’waxitshumbe exclosure is situated in the arid north-eastern section of the knp and was established in 1967. the original n’waxitshumbe exclosure was 230.0 ha, with an additional 72.0 ha added in 1986 (asner et al. 2009). mean annual precipitation is close to 400 mm and the soils here are derived from rocks of basaltic origin. the vegetation type is classified as colophospermum mopane shrubveld growing in broad-leaved bushveld (venter et al. 2003). the mean fire return interval for the n’waxitshumbe exclosure is approximately 4 years (asner et al. 2009), whilst smit et al. (2012) showed that the area surrounding the n’waxitshumbe exclosure has a mean fire return interval of less than 4 years. the faunal assemblage in the park includes all the major mammal species typical of the region. common browser species include kudu tragelaphus strepsiceros (pallas 1766), giraffe giraffa camelopardalis (linnaeus 1758), black rhinoceros diceros bicornis (linnaeus 1758), steenbok raphicerus campestris (thunberg 1811), grey duiker sylvicapra grimmia (linnaeus 1758) and bushbuck tragelaphus scriptus (pallas 1766). common mixed-feeders include impala aepyceros melampus (lichtenstein 1812) and the african elephant loxodonta africana (linnaeus 1758). community composition the distribution, abundance and density of the species growing both inside and outside of each of the exclosures was determined by walking four 100 m long by 4 m wide transects for each treatment at each site (1600 m2) in march 2011. the four transects in each treatment were taken parallel to each other and 50 m apart. inside and outside pairs were situated in as close a proximity as fences and firebreaks permitted. care was also taken to ensure that the paired transects were on similar positions along catenal gradients. at nhlangwini, the paired transects were taken perpendicular to the northern fence of the exclosure and parallel to the drainage in the area, outside transects were taken to the north-west of the exclosure. at makhohlola, the paired transects were taken perpendicular to the western fence and perpendicular to the drainage at the site, outside transects were taken to the north of the exclosure. owing to the limited size of the makhohlola exclosure (100 m × 240 m) two of the four transects were taken in the no-burn part of the fire experiment (since 2004) and two were taken in the part that burns in conjunction with the surrounding landscape. at n’waxitshumbe, the paired transects were taken perpendicular to the southern fence in the south-western portion of the exclosure and parallel to the drainage in that area. species identity was recorded for each woody plant encountered in each transect using a personal digital assistant (pda) with cybertracker software. the abundance data were used to create an inside abundance index, whereby the proportion of a species found in the inside compared to the outside treatment was calculated. this index was calculated by dividing the number of individuals of each species found inside by the total number of individuals found both inside and outside. thus a value of 1 indicates the species was only found inside, whilst a value of 0 indicates the species was only found outside. the inside abundance index was then plotted against a suite of leaf traits to test whether species growing on the inside had high values of these traits. this index can be used as a proxy to determine which plant species are preferentially selected for or ‘favoured’ by browsers relative to their abundance. trait sampling the dominant woody plant species making up 80% – 90% of the woody biomass inside and outside the three exclosures in the knp were determined (table 2). fourteen different plant traits relating to the leaves and stems were sampled for each species. all trait data (except bsi) were measured and recorded according to the methods explained in cornelissen et al. (2003). for each species, leaf material was collected from five individuals and analysed for n, p and carbon (c) content using a leco truspec cn analyser (leco corporation, st. joseph, mi). sla and average leaf area were measured using four healthy and complete sun-exposed leaves from five individuals (20 leaves per species). the bsi of a species is the total dry weight of the leaves removed from two human bites taken from five individuals (total of ten bites). an attempt was made to remove the maximum amount of leaf material with each bite. the bsi for all species was measured by the same person in order to control for potential differences between individual recorders. total condensed tannins were calculated according to the methods described by hattas and julkunen-tiitto (2012), whilst total polyphenols (tp) were calculated according to the methods described by hattas et al. (2005). cell wall constituents, that is, ndf and adf, were determined in an ankom fibre analyser (ankom technology corp., fairport, ny), whereas adl was determined in accordance with the acid detergent lignin in beakers method. ash was determined by incinerating the filter bag containing plant residue in a muffle furnace at 525 °c for 3 h. all cell wall constituents were determined consecutively as outlined in the ankom technology procedures (ankom technology 2010). table 2: dominant woody species accounting for > 80% of standing biomass for inside and outside treatments at the three kruger national park exclosures. soil sampling five replicate soil samples were collected both inside and outside of each exclosure at three depths (0 cm – 10 cm, 10 cm – 20 cm and 40 cm – 50 cm) using a soil auger. care was taken to sample away from the canopies of trees. soil bulk density measurements were taken at these same depths. the methods used for soil sampling and analyses are described in detail by wigley et al. (2013). statistical analyses all analyses were performed using r (r project 2012). chi-square tests were used to test for differences in tree densities between treatments. non-parametric wilcoxon tests were used to test for differences in trait measurements as the low replication resulting from the low species diversity at each site precluded the use of statistical tests founded on the normal distribution. analyses of variances were used to test for differences in soil nutrients between treatments using pooled data from all three depths, as the conditions for homoscedasticity were not violated (fligner test: p > 0.05; conover, johnson & johnson 1981). principle components analyses (pca) were used to explore the relationships between treatments, sites and measured plant traits. the function ‘pcasignificance’ available in the biodiversityr package (kindt & coe 2005) in r was used to test for significant pca axes using the broken-stick distribution. results top ↑ community composition data at the nhlangwini exclosure, there were a number of differences between the two communities. the four most common species for both treatments were t. sericea, dichrostachys cinerea, s. birrea and acacia gerrardii. however, t. sericea was found to be much more abundant (nearly 80% of all trees) inside compared to outside (c. 35%) of the exclosure (figure 1). dichrostachys cinerea showed the opposite pattern, with a higher relative abundance outside (30%) compared to inside (15%) the exclosure (figure 1). sclerocarya birrea and a. gerrardii showed similar abundances between treatments (figure 1). both treatments had a few unique species, with strychnos madagascariensis, catunaregam spinosa and searsia leptodictya only found inside, whilst antidesma venosum, combretum hereroense, albizia harveyi, philenoptera violacea and piliostigma thonningii were only found outside. species diversity was found to be higher on the outside of the exclosure, with a total of 16 species compared to 12 species inside. figure 1: the relative abundance of the species found inside and outside the three exclosures namely nhlangwini (a) inside and (b) outside, makhohlola (c) inside and (d) outside and n’waxitshumbe (e) inside and (f) outside, in the kruger national park. there were distinct differences in woody communities growing inside and outside of the makhohlola exclosure (figure 1). acacia nigrescens was found to be more abundant inside the exclosure (32%) but was still relatively abundant outside (22%). flueggea virosa was common inside (15%) but rare outside (4%). gymnosporia senegalensis was found to have a much higher abundance outside (33%) compared to inside (12%). lannea schweinfurthii and s. birrea had lower abundances outside (7% and 1%, respectively) compared to inside (12% and 7%, respectively). dichrostachys cinerea and a. harveyi were relatively common outside (13% and 9%, respectively) but rare inside (3% for both). dalbergia melanoxylon, ehretia rigida and euclea undulata were only found inside the exclosure, making species diversity higher on the inside (13 species) than outside (10 species). there were also distinct differences in the woody communities growing inside and outside of the n’waxitshumbe exclosure. at this site, c. mopane was found to be the most common species both inside (43%) and outside (40%, figure 1). the second most common species inside the exclosure (37%), d. melanoxylon, was not found outside of the exclosure. combretum imberbe and p. violacea were more common outside (14% for both) compared to inside (7% and 1%, respectively). sclerocarya birrea and g. senegalensis were relatively rare both inside and outside of the exclosures. dalbergia melanoxylon, grewia monticola and ozoroa obovata were unique to inside, whilst d. cinerea, a. harveyi and a. nigrescens were only found outside of the exclosure. thus species diversity was the same inside and outside of the exclosure, but species composition differed. population structure total woody plant densities were significantly higher for the inside treatments at all three sites (figure 2). these large differences were determined by the most common species growing at each site. the size class distributions for the two most common species growing at the nhlangwini exclosure (t. sericea and d. cinerea) show very different patterns for each treatment. the size class distribution of t. sericea inside the exclosure shows an inverse j-shaped curve, with many plants in the smaller size classes less than 3 m and fewer plants in the large size classes. the size class distribution for outside the exclosure shows much lower numbers of plants in the smaller size classes and fewer plants in the very large size classes (figure 3). a different pattern was evident for d. cinerea, with most plants falling into size classes between 2 m and 4 m inside the exclosure, whilst plants on the outside were more evenly distributed in a greater range of size classes (figure 3). figure 2: total tree densities for all woody plants greater than 0.5 m in height inside and outside the three long-term herbivore exclosures in the kruger national park. figure 3: size class distributions of terminalia sericea populations growing (a) inside and (b) outside and dichrostachys cinerea populations growing (c) inside and (d) outside of the nhlangwini exclosure in the kruger national park. at makhohlola, major differences were evident between treatments for each of the four most common species (figure 4). the size class distributions of l. schweinfurthii and a. nigrescens showed an approximately inverse j-shaped curve for the populations growing inside the exclosures but not for the outside populations. the inside populations of these two species both had a much higher number of individuals in the smaller (< 2 m) size classes than the outside populations (figure 4). the outside population of g. senegalensis was more uniform in height (1 m – 2 m) than the inside population (0 m – 3 m). there were many more and taller f. virosa plants growing on the inside of the exclosure compared to the outside (figure 4). figure 4: size class distributions of lannea schweinfurthii populations growing (a) inside and (b) outside, gymnosporia senegalensis populations growing (c) inside and (d) outside, flueggea virosa populations growing (e) inside and (f) outside and acacia nigrescens populations growing (g) inside and (h) outside of the makhohlola exclosure in the kruger national park. at n’waxitshumbe, the two most common species c. mopane and c. imberbe had similar size class distributions in each treatment (figure 5). however, there were higher densities in each size class for the inside treatment for c. mopane, whilst c. imberbe had similar numbers in each size class for both populations. the most striking difference was found for d. melanoxylon, where a population spanning the full range of size classes was found inside the exclosure, whilst not one individual was found outside of the exclosure in the sampling area (figure 5). figure 5: size class distributions of dalbergia melanoxylon populations growing (a) inside, combretum imberbe populations growing (b) inside and (c) outside and colophospermum mopane populations growing (d) inside and (e) outside of the n’waxitshumbe exclosure. no dalbergia melanoxylon plants were found growing on the outside of the exclosure in the kruger national park. leaf traits of unique inside species versus common outside species there were no significant differences (p > 0.05; wilcoxon test) in leaf nutrient concentrations – n, c, c:n, p, calcium (ca), magnesium (mg), sodium (na) and potassium (k) – between each of the inside and outside treatments (table 3). there were, however, some consistent trends; the lack of statistically significant differences was likely a result of the small sample sizes resulting from low woody species diversity at the sites. mean leaf n (%) was noticeably higher for the inside treatment at all three sites (table 3). the higher n and similar c concentrations inside of the nhlangwini exclosure resulted in a lower mean c:n ratio, whilst the other two sites had similar c:n ratios (table 3). leaf p (%) was similar between treatments at all three sites (table 3). table 3: mean (± s.e.) values of the measured leaf traits for the inside and outside treatments of three exclosures in the kruger national park. there were no significant differences (p > 0.05; wilcoxon test) in sla, leaf size, bsi and tensile strength between treatments. mean sla was consistently higher for species restricted to the inside treatments (table 3). average leaf area, which was highly correlated to bsi, were both higher inside the exclosures at the two nutrient rich sites on basalts (makhohlola and n’waxitshumbe), whilst the opposite pattern was true for the nhlangwini site which is situated on nutrient poor granite derived soils (table 3). mean leaf tensile strength was higher outside exclosures at the two basalt sites, but no difference was evident between treatments at the granite site (table 3).the pooled data showing the inside abundance index of each species plotted against leaf traits and bsi showed a weak positive relationship between the inside abundance index and leaf n and p (figures 6a and 6b). almost no relationship was evident between the inside abundance index and sla and bsi (figures 6c and 6d; p > 0.05 for all correlations). figure 6: data for, (a) leaf nitrogen, (b) leaf phosphorus, (c) specific leaf area and (d) bite size index plotted against the proportion of species found on the inside compared to the outside of three exclosures in the kruger national park. a value of 1 indicates that the species was only found inside, whilst a value of 0 indicates the species was only found outside the exclosures. chemical defence no significant differences (p > 0.05; wilcoxon test) in total phenolics, condensed tannins and fibre were evident between treatments. however, an interesting pattern emerged; mean leaf phenolic concentrations were consistently lower for species restricted to the inside treatments compared to those in the outside treatment, with the biggest difference evident at the nhlangwini site (table 3). mean condensed tannin concentrations showed no consistent patterns between treatments and sites (table 3). the three types of fibre analyses showed different patterns. ndf, which includes hemicellulose, cellulose and lignin, was slightly lower for the outside treatments at makhohlola and n’waxitshumbe, whilst the opposite pattern was found at nhlangwini (table 3). adf, which includes cellulose and lignin, showed no consistent patterns (table 3). adl, which only includes lignin, was consistently lower for the inside treatments at all three sites (table 3). soil nutrients although the pooled data for all depths of soil n and p showed a pattern of higher values for the three inside treatments, the only statistically significant differences were evident at n’waxitshumbe (p < 0.05; figures 7a and 7c). soil c and k also showed a pattern of being consistently higher for the inside treatments at makhohlola and n’waxitshumbe, with no differences evident at nhlangwini (figures 7b–7d). similar patterns were evident for n, c and p when total nutrient stocks were calculated using soil bulk density data (data not shown). figure 7: mean (± s.e.) for, (a) soil nitrogen, (b) carbon, (c) phosphorus and (d) potassium concentrations inside and outside of three exclosures in the kruger national park. plotted means are calculated from five replicates taken at three depths. multivariate analyses the pca performed on the data using 13 traits relating to the leaves and stem (figure 8) showed that the first two principal components together explained 44.0% of the total variation amongst species. the most important traits separating species along the first pca axis were ndf, adl and tp. the most important traits separating species along the second pca axis were leaf n, leaf c:n and stem density (figure 8). the pca showed that the species from the inside tended to more variable along pca axis 2 (higher leaf n and stem density), whilst the species from the outside tend to be more variable along pca axis 1 (higher fibre and tp values). the variance explained by the first pca axis was found to be insignificant, as the percentage of explained variance (24.3%) was not larger than the corresponding percentage of variance of the broken-stick distribution (24.5%), whilst the second pca axis was significant as the percentage of explained variance (20.3%) was larger than the corresponding percentage of variance of the broken-stick (16.8%) distribution (kindt & coe 2005). figure 8: principal components analysis of trait measurements from 20 species sampled inside and outside of three exclosures in the kruger national park. discussion top ↑ clear differences were found in both the species composition and population structure of the woody plant communities growing inside and outside of three herbivore exclosures in the knp. although the three exclosures were situated in very different vegetation types, some similarities exist in species’ responses to low herbivory. some species were frequently encountered both inside and outside of the exclosures (e.g. t. sericea and d. cinerea at nhlangwini, a. nigrescens and g. senegalensis at makhohlola, c. mopane at n’waxitshumbe). smaller size classes tended to be better represented inside, and larger size classes outside, for species which were found in large numbers both inside and outside of the exclosures. there were some exceptions though; g. senegalensis for instance, had similar population structures both inside and out. this could be because this species is not a preferred browsing species as a result of the high condensed tannin and adl concentrations in the leaves (see shrader et al. 2011). we argue that savanna woody species are either well adapted to browsing (such as d. cinerea), well defended (such as g. senegalensis) or both (such as a. nigrescens). for example, fornara and du toit (2007) showed that heavily browsed a. nigrescens trees develop tolerance traits, such as high regrowth rates and extensive branching, as well as resistance traits, such as close thorn spacing, suggesting that it is both well adapted and well defended. in fact, a meta-analysis by carmona, lajeunesse and johnson (2011) has shown that secondary metabolites are less important in plant defence when compared to other defensive plant traits such as physical resistance, gross morphology and growth rates. some species were found more frequently outside of the exclosures, such as d. cinerea, which was found in higher numbers outside of the exclosure at nhlangwini, and a. harveyi at all exclosures. these species are most likely stimulated by disturbance, for example d. cinerea shrubs rootsucker, and expand significantly when stimulated by fire and browsing (wakeling & bond 2007). more frequently, species were only found inside the exclosures (e.g. d. melanoxylon, g. monticola and o. obovata at n’waxitshumbe). dalbergia melanoxylon had the highest leaf n of all the species in this study and we assume that the species found only on the inside of the exclosures are mostly very palatable and targeted by browser species outside of the exclosures. dalbergia melanoxylon was found outside of the n’waxitshumbe exclosure in a previous study, but in lower lying areas than at our study area (levick & rogers 2008). it is uncertain whether d. melanoxylon has since disappeared outside the exclosure or whether this species may be preferentially targeted by browsers at specific positions along the catena. why do species become locally extinct outside of exclosures? impala and kudu are highly selective when foraging in nutrient-poor broadleaf savannas (cooper & owen-smith 1985; owen-smith & cooper 1987). this selectivity is attributed to differences amongst species in the level of defensive chemical compounds. alternatively, these species may also be predisposed to pollarding, uprooting and ringbarking and coppice poorly in response to elephant damage (o’connor, goodman & clegg 2007). a number of studies have documented the local extinction of plant species resulting from elephant impact (e.g. boundja & midgley 2010; o’connor et al. 2007 and references therein; penzhorn, robbertse & olivier 1974). several studies have shown that other herbivores can lead to the local extirpation of plant species. bond and loffel (2001) found that acacia davyi was no longer found in areas accessible to giraffe in ithala game reserve. total tree densities were significantly higher for the inside treatments at all three sites, usually with more individuals falling into the smaller size classes inside of the exclosures. these findings are in strong agreement with previous studies from the same study sites. trollope et al. (1998), levick and rogers (2008) and asner et al. (2009) reported higher tree densities and different population structures between treatments. moncrieff et al. (2011) reported different tree allometries between trees growing inside and outside of the same herbivore exclosures in the knp, with heavily browsed trees shorter for a given stem diameter. our results are therefore in agreement with previous studies and provide strong evidence that herbivory has a major impact on savanna woody plant diversity, species population structure and overall woody plant densities. the next step was to determine if there were differences between treatments for some of the key plant functional traits of each woody community. there was a trend in leaf n concentration, with higher mean n for the species unique to the inside treatments compared to the species found growing outside. no obvious patterns or trends were evident for leaf c:n or p. leaf structural traits, sla, average leaf size and bsi, which are all correlated, showed similar trends with higher values inside the exclosures at the two fertile sites and lower values inside compared to outside at the infertile site. the opposite pattern was evident for tensile strength where the leaves were generally tougher on the outside at the two fertile sites with no difference at the infertile site. apart from leaf n, total phenolic content and lignin content were the only traits that showed consistent patterns between treatments with higher values for species outside of the exclosures. increases of carbon-based structural metabolites with decreases in leaf n have been shown with high browsing intensity in previous work (bryant et al. 1991; wessels, van der waal & de boer 2007). an alternative theory for increases in lignin and leaf n inside the exclosures may be that herbivores preferentially select species with higher leaf n and that species with lower total phenolic and lignin content are also targeted by herbivores. in general, the removal of browsers led to higher soil nutrients, the response was the greatest at the two basalt sites. at n’waxitshumbe differences in soil n and p were higher with browsers absent, whilst soil c and k were also higher inside of the exclosures. the same pattern was evident at makhohlola, whilst the nutrient-poor site (nhlangwini) showed almost no differences between treatments for all measured soil nutrients. although herbivores have been found to increase soil nutrients (e.g. augustine & mcnaughton 2006; frank 2008), ritchie, tilman and knops (1998) found that herbivores can actually depress soil n by indirectly decelerating n cycling through decreasing the abundance of plant species with n-rich tissues. the increased soil fertility inside of the exclosures could possibly be explained by differing litter dynamics. the expectation would be that leaf litter with higher n and lower fibre content would result in faster cycling and improved soil fertility (e.g. reich et al. 2001). furthermore, the higher density of woody plants would increase the amount of leaf litter reaching the soil. thus the higher soil nutrients found inside of the exclosures could possibly be a result of the indirect effect of herbivores, as their removal resulted in higher tree densities. similarly, fire indirectly affects nitrogen cycling in the knp by decreasing woody densities (coetsee, bond & february 2010). the similar soil nutrient concentrations found inside and outside of the nhlangwini exclosure can most likely be attributed to the overall low litter quality and lower densities of herbivores resulting from the nutrient poor granitic soils at this site. conclusion top ↑ this study has demonstrated the important role browsers play in savanna dynamics. browsers directly impact species distributions, densities and population structures by actively selecting for species with favourable traits; this study suggests that species with high leaf n, low total phenolic content and low adl are favoured. forty years of no browsing impact was insufficient to allow the recruitment of the dominant species into larger size classes. this supports the idea that cohorts of trees recruit simultaneously, they would however need a break from both herbivory and fire. this study also suggests that browsers have important indirect effects on savanna functioning through their impact on soil nutrient cycling. the magnitude of these indirect effects on soil nutrient cycling appears to depend on inherent soil fertility and climate. the effect of browsers on both vegetation and soils was highest at the low rainfall site with high soil fertility, with a somewhat lesser effect at the higher rainfall basalt site, with almost no effect at the infertile high rainfall granite site. this work highlights the importance of herbivore exclosures in protected areas and some of the challenges faced by researchers when studying browser effects on savanna ecosystems because of the paucity of replicated long-term herbivory exclosures. future work should identify other possible sites in savannas from around the globe that could be used to corroborate the findings from this study. acknowledgements top ↑ this work formed part of the phd thesis of b.j. wigley and was made possible thanks to funding from a bdi-ped grant from the french national centre for scientific research, as well as the andrew w. mellon foundation. the authors are hugely grateful to: sanparks, kruger national park scientific services for allowing the study and logistical support, and scientific services game guards for protection in the field. thanks to two anonymous reviewers for their helpful comments which helped improve this manuscript. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions b.j.w. (nelson mandela metropolitan university) was responsible for the project design, data collection, analyses and reporting of data. c.c. (nelson mandela metropolitan university) assisted in collection of data and reporting, whilst w.j.b. (university of cape town) and h.f. (university claude bernard lyon 1) were both phd supervisors of b.j.w. and made valuable contributions to the refinement of the analyses and the manuscript. references top ↑ ammer, c., 1996, ‘impact of ungulates on structure and dynamics of natural regeneration of mixed mountain forests in the bavarian alps’, forest ecology and management 88, 43–53. http://dx.doi.org/10.1016/s0378-1127(96)03808-xanderson, v.j. & briske, d.d., 1995, ‘herbivore-induced species replacement in grasslands: is it driven by herbivory tolerance or avoidance?’, ecological applications 5, 1014–1024. http://dx.doi.org/10.2307/2269351 ankom technology, 2010, procedures, viewed 10 november 2013, from http://www.ankom.com/procedures.aspx asner, g.p., levick, s.r., kennedy-bowdoin, t., knapp, d.e., emerson, r., jacobson, j. et al., 2009, ‘large-scale impacts of herbivores on the structural diversity of african avannas’, proceedings of the national academy of sciences 106, 4947–4952. http://dx.doi.org/10.1073/pnas.0810637106, pmid:19258457, pmcid:pmc2650504 augustine, d.j. & mcnaughton, s.j., 1998, ‘ungulate effects on the functional species composition of plant communities: herbivore selectivity and plant tolerance’, the journal of wildlife management 62, 1165–1183. http://dx.doi.org/10.2307/3801981 augustine, d.j. & mcnaughton, s.j., 2004, ‘regulation of shrub dynamics by native browsing ungulates on east african rangeland’, journal of applied ecology 41, 45–58. http://dx.doi.org/10.1111/j.1365-2664.2004.00864.x augustine, d.j. & mcnaughton, s.j., 2006, ‘interactive effects of ungulate herbivores, soil fertility, and variable rainfall on ecosystem processes in a semi-arid savanna’, ecosystems 9, 1242–1256. http://dx.doi.org/10.1007/s10021-005-0020-y bergvall, u.a. & leimar, o., 2005, ‘plant secondary compounds and the frequency of food types affect food choice by mammalian herbivores’, ecology 86, 2450–2460. http://dx.doi.org/10.1890/04-0978 bond, w.j. & loffell, d., 2001, ‘introduction of giraffe changes acacia distribution in a south african savanna’, african journal of ecology 39, 286–294. http://dx.doi.org/10.1046/j.1365-2028.2001.00319.x boundja, r.p. & midgley, j.j., 2010, ‘patterns of elephant impact on woody plants in the hluhluwe‐imfolozi park, kwazulu‐natal, south africa’, african journal of ecology 48, 206–214. http://dx.doi.org/10.1111/j.1365-2028.2009.01104.x bryant, j.p., provenza, f.d., pastor, j., reichardt, p.b., clausen, t.p. & du toit, j.t., 1991, ‘interactions between woody plants and browsing mammals mediated by secondary metabolites’, annual review of ecology and systematics 22, 431–446. http://dx.doi.org/10.1146/annurev.es.22.110191.002243 carmona, d., lajeunesse, m.j. & johnson, m.t., 2011, ‘plant traits that predict resistance to herbivores’, functional ecology 25, 358–367. http://dx.doi.org/10.1111/j.1365-2435.2010.01794.x codron, d., lee-thorp, j.a., sponheimer, m., codron, j., de ruiter, d. & brink, j.s., 2007, ‘significance of diet type and diet quality for ecological diversity of african ungulates’, journal of animal ecology 76, 526–537. http://dx.doi.org/10.1111/j.1365-2656.2007.01222.x, pmid:17439469 coetsee, c., bond, w.j. & february, e.c., 2010, ‘frequent fire affects soil nitrogen and carbon in an african savanna by changing woody cover’, oecologia 162, 1027–1034. http://dx.doi.org/10.1007/s00442-009-1490-y, pmid:20213154 coetzee, b.j., engelbrecht, a.h., joubert, s.c.j. & retief, p.f., 2008, ‘elephant impact on sclerocarya caffra trees in acacia nigrescens tropical plains thornveld of the kruger national park’, koedoe 22, 39–60. conover, w.j., johnson, m.e. & johnson, m.m., 1981, ‘a comparative study of tests for homogeneity of variances, with applications to the outer continental shelf bidding data’, technometrics 23, 351–361. http://dx.doi.org/10.1080/00401706.1981.10487680 cooper, s.m. & owen-smith, n., 1985, ‘condensed tannins deter feeding by browsing ruminants in a south african savanna’, oecologia 67, 142–146. http://dx.doi.org/10.1007/bf00378466 cornelissen, j.h.c., lavorel, s., garnier, e., diaz, s., buchmann, n., gurvich, d.e. et al., 2003, ‘a handbook of protocols for standardised and easy measurement of plant functional traits worldwide’, australian journal of botany 51, 335–380. http://dx.doi.org/10.1071/bt02124 díaz, s., lavorel, s., mcintyre, s.u.e., falczuk, v., casanoves, f., milchunas, d.g. et al., 2007, ‘plant trait responses to grazing – a global synthesis’, global change biology 13, 313–341. http://dx.doi.org/10.1111/j.1365-2486.2006.01288.x fornara, d.a. & du toit, j.d., 2007, ‘browsing lawns? responses of acacia nigrescens to ungulate browsing in an african savanna’, ecology 88, 200–209. http://dx.doi.org/10.1890/0012-9658(2007)88[200:blroan]2.0.co;2 frank, d.a., 2008, ‘ungulate and topographic control of nitrogen: phosphorus stoichiometry in a temperate grassland; soils, plants and mineralization rates’, oikos 117, 591–601. gadd, m.e., 2002, ‘the impact of elephants on the marula tree sclerocarya birrea’, african journal of ecology 40, 328–336. http://dx.doi.org/10.1046/j.1365-2028. 2002.00385.x haslam, e., 1988, ‘plant polyphenols (syn. vegetable tannins) and chemical defense – a reappraisal’, journal of chemical ecology 14, 1789–1805. http://dx.doi.org/10.1007/bf01013477 hattas, d., stock, w.d., mabusela, w.t. & green, i.r., 2005, ‘phytochemical changes in leaves of subtropical grasses and fynbos shrubs at elevated atmospheric co2 concentrations’, global and planetary change 47, 181–192. http://dx.doi.org/10.1016/j.gloplacha.2004.10.010 hattas, d. & julkunen-tiitto, r., 2012, ‘the quantification of condensed tannins in african savanna tree species’, phytochemistry letters 5, 329–334. http://dx.doi.org/10.1016/j.phytol.2012.02.013 helm, c.v. & witkowski, e.t., 2012, ‘continuing decline of a keystone tree species in the kruger national park, south africa’, african journal of ecology 51(2), 270–279. http://dx.doi.org/10.1111/aje.12032 huntly, n., 1991, ‘herbivores and the dynamics of communities and ecosystems’, annual review of ecology and systematics 22, 477–503. http://dx.doi.org/10.1146/annurev.es.22.110191.002401 jachmann, h. & croes, t., 1991, ‘effects of browsing by elephants on the combretum/terminalia woodland at the nazinga game ranch, burkina faso, west africa’, biological conservation 57, 13–24. http://dx.doi.org/10.1016/0006-3207(91)90105-i kindt, r. & coe, r., 2005, tree diversity analysis: a manual and software for common statistical methods for ecological and biodiversity studies, world agroforestry centre, nairobi. pmcid:pmc1156951 kriebitzsch, w.u., oheimb, g.v., ellenberg, h., engelschall, b. & heuveldop, j., 2000, ‘development of woody plant species in fenced and unfenced plots in deciduous forests on glacial moraine soils in northern germany’, allgemeine forst-und jagdzeitung 171, 1–10. levick, s. & rogers, k., 2008, ‘patch and species specific responses of savanna woody vegetation to browser exclusion’, biological conservation 141, 489–498. http://dx.doi.org/10.1016/j.biocon.2007.11.001 levick, s.r., asner, g.p., kennedy-bowdoin, t. & knapp, d.e., 2009, ‘the relative influence of fire and herbivory on savanna three-dimensional vegetation structure’, biological conservation 142, 1693–1700. http://dx.doi.org/10.1016/j.biocon.2009.03.004 levick, s.r., asner, g.p., kennedy-bowdoin, t. & knapp, d.e., 2010, ‘the spatial extent of termite influences on herbivore browsing in an african savanna’, biological conservation 143, 2462–2467. http://dx.doi.org/10.1016/j.biocon.2010.06.012 moncrieff, g.r., chamaillé-jammes, s., higgins, s.i., o’hara, r.b. & bond, w.j., 2011, ‘tree allometries reflect a lifetime of herbivory in an african savanna’, ecology 92, 2310–2315. http://dx.doi.org/10.1890/11-0230.1, pmid:22352170 o’connor, t.g., goodman, p.s. & clegg, b., 2007, ‘a functional hypothesis of the threat of local extirpation of woody plant species by elephant in africa’, biological conservation 136, 329–345. http://dx.doi.org/10.1016/j.biocon.2006.12.014 owen-smith, n., 2005, ‘incorporating fundamental laws of biology and physics into population ecology: the metaphysiological approach’, oikos 111, 611–615. http://dx.doi.org/10.1111/j.1600-0706.2005.14603.x owen-smith, n. & cooper, s.m., 1987, ‘palatability of woody plants to browsing ruminants in a south african savanna’, ecology 68, 319–331. http://dx.doi.org/10.2307/1939263 penzhorn, b.l., robbertse, p.j. & olivier, m.c., 1974, ‘the influence of the african elephant on the vegetation of the addo elephant national park’, koedoe 17, 137–158. r project, 2012, ‘r: a language and environment for statistical computing’, r foundation for statistical computing, vienna, viewed 10 november 2013, from http://www.r-project.org/ reich, p.b., peterson, d.w., wedin, d.a. & wrage, k., 2001, ‘fire and vegetation effects on productivity and nitrogen cycling across a forest-grassland continuum’, ecology 82, 1703–1719. ritchie, m.e., tilman, d. & knops, j.m., 1998, ‘herbivore effects on plant and nitrogen dynamics in oak savanna’, ecology 79, 165–177. shrader, a.m., bell, c., bertolli, l. & ward, d., 2011, ‘forest or the trees: at what scale do elephants make foraging decisions?’, acta oecologica 42, 3–10. http://dx.doi.org/10.1016/j.actao.2011.09.009 smit, i.p., smit, c.f., govender, n., linde, m.v.d. & macfadyen, s., 2012, ‘rainfall, geology and landscape position generate large‐scale spatiotemporal fire pattern heterogeneity in an african savanna’, ecography 35, 1–13. staver, a.c., bond, w.j., stock, w.d., van rensburg, s.j. & waldram, m.s., 2009, ‘browsing and fire interact to suppress tree density in an african savanna’, ecological applications 19, 1909–1919. http://dx.doi.org/10.1890/08-1907.1, pmid:19831079 trollope, w.s.w., trollope, l.a., biggs, h.c., pienaar, d. & potgieter, a.l.f., 1998, ‘long-term changes in the woody vegetation of the kruger national park, with special reference to the effects of elephants and fire’, koedoe 41, 103–112. van hees, a.f.m., kuiters, a.t. & slim, p.a., 1996, ‘growth and development of silver birch, pedunculate oak and beech as affected by deer browsing’, forest ecology and management 88, 55–63. http://dx.doi.org/10.1016/s0378-1127(96)03809-1 venter, f.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation pattern’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83–129, island press, washington, dc. wakeling, j.l. & bond, w.j., 2007, ‘disturbance and the frequency of root suckering in an invasive savanna shrub, dichrostachys cinerea’, african journal of range and forage science 24, 73–76. http://dx.doi.org/10.2989/ajrfs.2007.24.2.3.157 wessels, d.c.j., van der waal, c. & de boer, w.f., 2007, ‘induced chemical defences in colophospermum mopane trees’, african journal of range and forage science 24, 141–147. http://dx.doi.org/10.2989/ajrfs.2007.24.3.4.297 wigley, b.j., coetsee, c., hartshorn, a.s. & bond, w.j., 2013, ‘what do ecologists miss by not digging deep enough? insights and methodological guidelines for assessing soil fertility status in ecological studies’, acta oecologica 51, 17–27. http://dx.doi.org/10.1016/j.actao.2013.05.007 filelist convert a pdf file! koedoe 19: 31-41 (i976) geologie van die tsitsikamakusstrook d. k. toerien departement van mynwese visagie straat pretoria 0001 same vatting die topografie, fisiografie en geologie van die tsitsikamakusstrook, wat die tsitsikamabos en -seekus nasionale parke insluit, word bespreek en toegelig met foto's en 'n geologiese kaart, skaall : 100000. abstract the topography, physiography and geology of the tsitsikama coastal area which includes the tsitsikama forest and coastal national parks are discussed and illustrated by photographs and a geological map, scale 1 : 100 000. inleiding die tsitsikamakusstrook strek langs die welbekende tuinroete en die gebied wat hier beskryf word, dek die 5 tot 8 km bree kusvlakte tussen die tsitsikamabergreeks en die indiese oseaan. die oosterlengtegrade 23°30' en 24° 15' vorm die wesen oosgrens respektiewelik. die tsitsikamabos nasionale park beslaan 478 ha van inheemse woud naby die paul sauerbrug oor die stormsrivier. die tsitsikama-seekus nasionale park is 'n smal aaneenlopende strook van 109 km 2, sowat 6.0 . km lank, langs die kuslyn tussen die monde van die twee grootrivlere. die hoe gemiddelde reenval van 1 050 mm per jaar stimuleer die weelderige plantegroei. inheemse woude en plantasies wat hoofsaaklik onder die beheer van die departement van bosbou val kenmerk die tsitsikama gebied . minder verweringsbestande rotsformasies is diep ingekerf en herkenbare dagsome is beperk tot die steil hange van die tsitsikamabergreeks, die diep, smal klowe van die riviere en die rotsagtige kuslyn . topografie en dreinering die rotsagtige kuslyn van die gebied vorm 'n eskarp, min of meer 180 m hoog. hiervandaan styg 'n kusvlak geleidelik na die binneland tot 'n hoogte van sowat 275m aan die voet van die tsitsikamabergreeks. dil is ongeveer 5 km breed in die weste en 8 km in die ooste. die berge skiet skerp die hoogte in met hoogste pieke tussen 1 300 m en 1 676 m net buite die noordgrens van die gebied. 'n aantal riviere soos die groot-, bloukrans-, elandsen storms31 rivicr dreineer vanuit die berge oar die kusvlakte na die see in diep en nou kransagtige klowe. ander, meestal jonger riviere, stort in watervalle oor die cskarp in die see. filiografie 'n uitstaande en opvallende kenmerk van die tsitsikamagebied is die verbascnde gelyk stoep land wat geleidelik styg van die eskarp by die kus lot aan die voet van die berge. hierdie is een van die mooiste brandersloepe in ons land. seker die beste waarnemingspunt is op die reguit stukkie oos-wes pad net voordat skerp suid gedraai word om in die grootrivierpas vanaf die weste af te daal (fig. 1 en fig. 2). oosen noordwaarts strek die landskap soos 'n tafel tot teen die tsitsikamabergreeks. branderaksie het dit geskuur oor weerstandbiedende steilhellende sandsteen en kwartsiet in die tersihe tydperk hens tussen 70 en 2 miljoen jaar gelede. met die daling van die land relatief tot seevlak is die kus geleidelik oorstroom tot 'n hoogte van 275m waar die see tot stilstand gekom het met die kuslyn teen die steil tsitsikamaberge as voorland. styging van die land het weer die branderstoep blootgele en die riviere het diep smal klowe daarin gekerf na die terugtrekkende see (fig. 2). hierdie klowe getuig van 'n relatief vinnige daling van die seevlak . die struktuur en samestelling van die onderliggende rotsformasies het bygedra tot die dreineringspatroon wat op die branderstoep ontwikkel het. so byvoorbeeld volg die vark-, 'n tak van die bloukransrivier, en gedeeltelik die elandsen oostelike grootrivier die strekking van die ingeplooide relatief sagte formasies van die groepe bokkeveld en tafelberg. die bloukransen stormsrivierpas is ook in hierdie gesteentes gesny. bogemelde stygings en daling van seevlak is sedertdien opgevolg deur soortgelyke herhaalde skommelings oor 'n vertikale hoogte van meer as 100 m sodat dit by tye heelwat laer gestaan het as tans. die bewegings was geleidelik, maar ook met rukke en stote sodat, soos verwag kan word, branderaksie ook ander waarneembare merke nagelaat het. daar is 'n mooi branderstoep op ongeveer 120 m by nature's valley wes van die grootrivier, en een tussen laasgenoemde en die bloukransriviermond (fig. 3). verder oos verskyn dit net as 'n knik in die eskarp. nog jonger stoepe op ongeveer 18 m, 6 m tot 7,6 m en andere is mooi ontwikkel buitekant hierdie gebied. tekens daarvan is egter wel te bespeur soos die van 7,6 m waarop die restaurant by stormsriviermond gelee is (fig. 4), en verskeie seegrotte langs die kus. soos in die geval by die sondagsriviermond en andere langs die suiden ooskus het boorgate getoon dat spoelgruis en sand die rivierkanale vul tot dieptes van meer as 30 m onderkant die huidige seevlak. opnames van die seebodem langs die kus het getoon dat hierdie rivierlope nog ver oor die vastelandplat voortstrek. seevlak moes dus eens op 'n tyd heelwat laer gestaan het as tans. die gedagte is dat ons vlak vastelandsplat gedurende hierdie tyd gekerf is. 32 met daaropvolgende styging(s) van seevlak is die rivierlope toe verdrink en gedeeltelik gevul met gruis en sand. sand is voor en in sommige se monde gespoel en hulle kuier nou lui-lui rond in hierdie opvulling. die grootrivier by nature's valley en die keurboomsrivier by plettenbergbaai, wes van hierdie gebied, is mooi voorbeelde. die besondere reguit tsitsikamakuslyn volg die oos-suidoostelike strekking van die geplooide formasies. 'n moontlike strekkingsverskuiwing in die see langs die kus kon ook hiertoe bygedra het. geologie algerneen die volgende tabel rangskik van onder na bo die geologiese formasies wat in die gebied voorkom vanaf die oudste tot die jongste (fig. 5). spoel en waaisand ] gruis en silkreet skalie sands teen, arkose sands teen, kwartsiet sandsteen, kwartsiet skalie sandsteen, kwartsiet sands teen, kwartsiet filliet laat tersier (?) tot kwaterner formasie gydo 1 baviaanskloof kouga tchando } cedarberg peninsula klein rivier'" groep bokkeveld tafelberg gamtoos supergroep kaap ':'al die formasies van die supergroep kaap en groep gamtoos is van marine oorsprong. die groep gamtoos word vooriopig gekorreleer met die groep malmesbury van die wes-kaap wat sowat 550 miljoen jaar oud is. die supergroep kaap se ouderdom is tussen 350 en 500 miljoen jaar. geologies vorm die tsitsikama deel van 'n groot geheel. die formasies van die supergroep kaap bou die kaapse plooigordel wat die suid-kaap om so om vanaf die suidelike namakwaland in die weste tot oos van grahamstad in die ooste. behalwe vir die kwaternere afsettings is die regionale strekking oossuidoos. die formasies is in hierdie gebied in 'n diep sinklien geplooi wat soos 'n lang wors oor die hele lengte van die tsitsikama strek. formasie kleinrivier soos op die geologiese kaart aangedui, dagsoom hierdie formasie in 'n sma i strook langs die kus. dit is mooi blootgele by die mond van die stormsrivier. 33 dit bestaan uit afwisselende dun bande van grys tot donkergrys sandsteen, kwartsitiese sandsteen en filliet, en is dan ook selektief verweer in weerstandbiedende uitstaande bande en riwwe van eersgenoemde en sloepe van filliet. die naaste volgende dagsome van die groep gamtoos is in die kouga-, baviaanskloofen gamtoosvalleie, en op grond van litologiese ooreenkoms word bogenoemde voorlopig met die formasie klein rivier gekorreleer. die strekking is soos die van die supergroep kaap. dis is ook saamgeplooi sodat 'n diskordansie nie waarneembaar is in hierdie gebied nie. die oorgang tot die massiewe ligter gekleurde sandsteen van die formasie peninsula kan egter in die tsitsikama binne kort afstand vasgestel word. formasie peninsula in die suiden oos-kaap vorm die peninsula die basis van die groep tafdberg. dit bestaan uit weerstandbiedende wit tot blougrys, hoofsaaklik grofkorrelige sandsteen en kwartsitiese sandsteen wat opvallend suiwer is. dit bou by uitstek die hoe bergreekse met skouspelagtige kranse om die suidpunt van suid-afrika o.a. tafelberg by kaapstad, asook die olifants-, hexrivier-, outeniekwa-, swarten tsitsikamaberge. die paul sauerbrug oar die starmsrivier is in kranse van die peninsula geanker en die vertikale massiewe bande van sandsteen is mooi in die hoe wande te sien (fig. 6). met verwering dra hierdie sandsteen grootliks by tot die opvallende wit sandstrande van die suidkus. die dagsome is in baie gevalle glasig wit, maar op plekke grys tot rooibruin gekleur deur ysteroksied. 'n opvallende kenmerk van die formasie is die wydverspreide afgeronde wit kwartsrolsteentjies tot en met duifeiergrootte. tussengelaagd met die sandsteen en kwartsitiese sands teen is wydgespasieerde swart skaliebande wat grys verweer en gewoonlik minder as 'n meter dik is. hul dagsoom egter selde daar hulle in die klowe weens die sagtheid van die skalie met bosstrepe bedek is. by die elandsrivier naby robbehoek waar die formasie steil hel, is die dikte sowat 1 480 meter. noordwaarts neem die dikte egter toe sodat dit noord van die kougarivier, buite hierdie gebied, 2 700 m is. op plekke langs die strekking, so os by die elandsrivier naby robbehoek, is die sandsteen donkergekleurd by die kontak met die formasie cedarberg. f ormasie c edarberg die formasie cedarberg volg konkardant op die peninsula. dit bestaan uit swart skalie wat sanderig kan wees en selfs dun sandsteenbande kan insluit naby die basis. weswaarts buite hierdie gebied gaan die sandsteenbande oor in 'n tilliet. 34 as 'n uiters inkompetente formasie is dit intens geplooi en diep verweer, en bou dit negatiewe landvorme. dagsome is baie skaars weens grondbedekking en plantegroei. waar sigbaar, is dit gewoonlik verweer tot veelkleurige kleie. die formasie se dikte kon nie hier gemeet word nie, maar verder noord is dit 35 m tot 55 meter. dit vertoon somtyds dunner as wat dit werklik is as gevolg van gedeeltelike uitknyping in plooie. naby die top word die forrnasie sanderig en die oorgang na die vol gende is konkordant en skerp. formasie tchando hierdie formasie bestaan ook uit sandstene, maar dit is nie so suiwer as die van die peninsula nie. dit is ietwat dunner geband en ook meer intens geplooi, en weens die laer weerstand teen verwering vertoon dit gewoonlik negatief in die topografie. die aanwesigheid van gedissemineerde ysteren mangaanminerale verleen 'n vuilbruin tot sjokoladebruin kleur aan die verweerde oppervlak. skaliebande kom ietwat meer voor as in die peninsula, maar is nog baie ondergeskik aan sandsteen. dagsome naby die kus toon 'n vermeerdering in dun skaliebandjies na die suide. die dikte by die kus is 280 m maar, soos in die geval van die vorige formasies, raak dit dikker noordwaarts, sodat oos van die baviaanskloof dit tot 330 m dik is. oorgang tot die volgende formasie is gewoonlik redelik skerp. formasie kouga bestaande uit skoner, witter en meer weerstandbiedende sandsteen en kwartsiet as sy voorganger, vertoon hierdie formasie topografies ook meer prominent. dit is egter nie so massief soos die peninsula nie. alhoewel die formasies peninsula en tchando ook kruisgelaagdheid vertoon, is dit geensins so opvallend soos in die kouga nie. laasgenoemde is ook sterk veldspaties naby die top. die voorkoms van skaliebande is min of meer soos in die tchando. die dikte in hierdie omgewing is sowat 300 m, maar verder noord tot soveel soos 400 meter. die formasie baviaanskloof volg konkordant en die oorgang is redelik skerp. formasie baviaansklorif hierdie formasie bestaan uit drie herkenbare eenhede. 'n middelste mediumkorrelrige opvallend veldspatiese sandsteen skei twee onsuiwer donkergekleurde sandsteeneenhede van mekaar. die middelste is blougrys, somtyds 'n ware arkose, en weens hoer weerstand teen verwering as die ander twee, bult dit gewoonlik uit in dagsome. dit is lig van kleur op die verweringsvlak selfs kenmerkend dofwit in droeer dele so as in die baviaanskloof, terwyl die ander twee 35 redelik maklik verweer tot 'n vuilbruin kleur. die formasie het glad nie die massiewe voorkoms van die vorige nie en bou dus negatiewe landvorme. ondergeskikte sliksteen en skaliebandjies kom wei voor, maar nie op regionale skaal nie. die totale dikte by die oostelike grootrivier is sowat 300 m, met die middelste sandsteen ongeveer 47 meter. die onderste lid is gewoonlik die dikste. aldrie varieer taamlik in dikte op regionale skaal. by die paul sauerdam in die kougavallei is die diktes van die drie lede (onderste eerste) 8"5 m, 53 m en 55 meter. wanneer daar in die bloukranspas vanaf die ooste afgedaal word, sny die pad 'n profiel deur hierdie formasie wat daar steil hel. die middelste lid is goed ontwikkel net voor die brug oorgesteek word onder in die pas en is daar ongeveer 43 m dik. die boonste lid van die formasie bevat fossiele wat ooreenkom met die in die formasies van die bokkeveld. hier is dit egter geheel en al vernietig deur deformasie. die basis van die konkordant-opvolgende formasie gydo van die groep bokkeveld word geneem waar die formasie baviaanskloof oorgaan in afwisselende skalie en sandsteen. f ormasie gydo die groep bokkeveld bestaan uit afwisselende skalieen sands teenformasies met diktes van 30 m tot 300 meter. net die onderste, formasie gydo, word in hierdie gebied gevind. dit is ingeknyp in die middel van 'n sinklien wat oor die lengte van die kaartgebied strek. aan die basis van die formasie volg 'n paar dun skalieen sandsteenbande mekaar afwisselend af, maar vir die res bestaan dit hoofsaaklik uit skalie met slegs 'n paar uiters dun sandsteenbandjies. die dikte van die formasie kan in hierdie gebied nie bepaal word nie weens die afwesigheid van die top daarvan en die intensiewe plooiing. dit is egter een van die dikste formasies in die bokkeveld. as 'n skalie is dit ook een van die formasies wat die minste weerstand bied in die gebied sodat sytakke van sowel die ouer, as die jonger riviere die strekking daarvan volg. dit is om die rede ook intens verkerf tot 'n uiterste heuwelen valleitopografie in die tersiere branderstoep. die vallei van die varkrivier, 'n sytak van die bloukrans, wat goed te sien is noord van die pad voor daar in die bloukranspas afgedaal word, is 'n mooi voorbeeld. in die bloukrans-, grootrivieren ou stormsrivierpas is hierdie skalie diep verweer. dit is spierwit geloog of geel-rooi en bruin gekleur deur ysteroksied. die oorspronklike swart kleur is nog te sien in die diepste dele van sommige passe en aan die kus vanaf nature's valley weswaarts. die skalie is intensief geplooi in die wande van die passe. die formasie dra baie dierefossiele en skulpe; sterk deformasie en latere verwering het dit egter verwoes. net op die kus by nature's valley en weswaarts is daar nog herkenbare tekens van hulle teenwoordigheid te bespeur in die vorm van geelbruin ysteroksiedryke vervormde reste. 36 oppervlakajsettings die tersiere branderstoep is bedek deur 'n bosbegroeide laag van leem en sanderige grond en klei. langs die paaie kan daar plek-plek in uitgrawings gesien word hoe sekondere ysteroksied dit geel, rooibruin en donkerbruin kleur. in die blueliliesbushbosreservaat suid en suidwes van pineview is knolle van gibbsiet in die wit kaolienagtige klei gevind. langs die valleie van die riviere is dun afgebroke lense van afgeronde rolstene ontbloot waar dit onder die bogemelde bogrond ie. dit mag alluviale of marine afsettings wees. klein kolle van silkreet dagsoom op die rant van die tersiere terras wes van die bloukransriviermond. die strandoord nature's valley is gelee op sand wat in bosbegroeide duine opgewaai is en die mond van die verdrinkte grootrivier gedeeltelik versper. struktuurgeologie die supergroep kaap en voor-kaapgesteentes is regionaal onderwerp aan sterk noord-suid georienteerde druk wat intensiewe plooiing op groot skaal veroorsaak het. die plooi-asse strek oos-suidoos en duik afwisselend redelik vlak oos en wes. oorplooiing na die noorde wat dikwels met oorskuiwing en breksiasie gepaard gaan is 'n algemene verskynsel. intensiewe kliewing en sku ifskeuring het plaasgevind en suidwaartshellende skuifskeurvlakke wat laagvlakke op groot skaal bykans uitgewis het, word maklik vir laasgenoemde aangesien. goedontwikkelende en opsigtelike diggespasieerde rekspanningsnate met hoe hellings na die weste en ooste, asook horisontale nate (fig. 6) dateer ook uit hierdie fase van plooiing. bogenoemde regionale verskynsels is feitlik almal in hierdie gebied waarneembaar. die profief a-b (fig. 5) loon die diep sinklien hierbo gemeld, wat soos 'n lang wors oor die hele lengte van die tersiere branderstoep strek en al die voor-tersiere formasies betrek. die suidelike been is effe oorgeplooi, en die effek op die topografie, van die ingeplooide formasies baviaanskloof en gydo wat minder weerstand bied, is reeds geskets. kleinskaalse plooie is ook prominent ontwikkel in hierdie inkompetente gesteentes. dit is duidelik waarneembaar in die b.e~gpasse en in die voor-kaapgesteentes by die mond van die stormsnvler. op die kus by die karavaanpark by slormsriviermond is die eifek van kliewing en skuifskeuring op verwering mooi uitgebeeld. differenside verwering langs hierdie vlakke het die vlakgeplooide strata omskep in skerp, feitlik loodregte, riwwe en sloepe. ooswaarts na en om die punt by die restaurant begin die geplooide bande in werklikheid hoe hellings aanneem met klein sinen antikliene ook sigbaar. die harde kwartsietiese bande vorm uitstaande riwwe terwyl die sagter sliksteen en skalies teruggekerf is in sloepe. by die westelike punt van die karavaanpark sny 'n aantal loodregte kwartsare dwarsoor die strekking (fig. 7). hulle vul die rekspannings37 nate hierbo genoem, en is tot 1,5 m dik. die kwartsare is swak gemineraliseer en in die verlede geprospekteer vir goud. die sulfides geleniet (loodkleurig), piriet en chakopiriet (goudkleurig) asook malgiet (liggroen) en die karbonaat ankeriet (bruin) is maklik herkenbaar. sekondere kwartsare, anders georienteer, is ook te sien soos byvoorbeeld in die uitstaande skilderkrans 'n paar kilometer wes van stormsriviermond. op die rotse in die sloepe voor die karavaanpark, wat maklik bereikbaar is met laagwater, kom taile langwerpige gaatjies met langasse parallel aan die nate voor. hul is blykbaar oplossingsholtes langs die nate, en is sowat 1 cm diep en tot 3 cm lank. die rotse is oor die algemeen hier sterk genaat. die diep worsvormige sinklien hierbo genoem vertak in twee naby die oosgrens as gevolg van 'n weswaartsduikende antiklien wat vanaf die ooste inkom. in die weste van die gebied kantel die sinklien na die sui de sodat naby die grootrivier normale en omgekeerde hellings so laag soos 300 en 40° na die noorde in die skalie en sands teen respektiewelik gemeet is. die oorsaak van hierdie verskynsel is moeilik te verklaar. dit mag veroorsaak wees deur weerstand teen druk van 'n bedekte koepel van voor-kaapgesteentes. die sinklien stuit teen 'n suidoosstrekkende skarnierverskuiwing wat deur die benede grootrivier gevolg word. die formasies is wes hiervan suid verplaas en keer verder weswaarts weer terug na die normale hellings. langs die verskuiwing is die gesteentes erg gebreksieer, veral die peninsula by 'n punt oos van die mond. die verskuiwing mag 'n tak wees van 'n moontlike strekkingsverskuiwing wat langs die kus in die see ie. die vermoede word versterk deur breksiasie op die kus op verskeie plekke net buite die gebied, oos sowel as wes. twee ander groot verskuiwings met reuse verplasings, wat waarskynlik hul oorsprong in rekking vanuit die suide het, loop noord van die gebied in die kougaen baviaanskloofriviervalleie. bibliografie die volgende publikasies kan vir bykomende inligting nageslaan word: haughton, s. h. 1969. geological history ofsouthem africa. cape town: geological society of south africa. king, l. c. 1963. south african scenery, a textbook of geomorphology. 3rd ed. edingburgh: 0 liver and boyd. krige, a. v. 1926. an examination of the tertiary and q.uarternary changes of sealevel in south africa, with special stress on the evidence in favour of a world-wide sinking of oceanlevel. ann. univ. stell. 5(a, 1):1-18. ruddock, a. 1968. cainozoic sea-levels and diastrophism in a region bordering algoa bay. trans. geol. soc. s. afr. 71(3):209-233. taljaard, m. s. 1948. oor berge en vlaktes. stellenbosch: die universiteitsuitgewers en -boekhandelaars (edms) beperk. 38 fig. i. 'n blik ooswaarts oor die tersiere branderstoep van die tsitsikama van ' n punt wes van die grootrivier, noord van nature's valley. fig. 2. ' n blik noordwaarts oor die diep nou kloof van die stormsrivier gesien vanaf die mond. let op hoe die tersiere branderstoep gekerf is tot teen die tsitsikamaberge. 39 fig. 3. 'n blik weswaarts oor die ongeveer 120 m branderstoep gesny oor die formasie peninsula wes van die bloukransrivier. die hoer tafelland op die horison is die tersiere branderstoep. fig. 4. die restaurant op die ongeveer 7 m terras by stormsriviermond. die hoer bosbegroeide vlak is dee! van die 120 m branderstoep. die suidwaartse helling van die strata is duidelik te sien op die voorgrond regs. 40 fig. 6. die paul sauerbrug oor die stormsrivier. let op die vertikale strata van die formasie peninsula sowel as die goedontwikkelde byna horisontale nate. fig. 7. die vertikale noord-suidstrekkende kwartsare op die kus wes van stormsriviermond. die sterk genaatheid van die gesteentes is opvallend. 41 page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 issn 0075-6458 77 koedoe 49/2 (2006)issn 0075-6458 77 koedoe 49/2 (2006) introduction wildlife viewing in protected areas as a form of recreation is steadily increasing in popularity. as a result, there has been a rapid increase in the number of small reserves (<1000 km²) during the last few years in south africa (druce et al. 2004; slotow et al. 2005). although some of these reserves have been developed to enhance biodiversity conservation, most of them exist purely for the eco-tourism industry. tourism entails the reintroduction of valuable viewing species into these reserves regardless of the ecology and the sustainability of these and other species. the african elephant (loxodonta africana) is one of the key species in the ecotourism industry. as a result, many state and private game reserves have reintroduced elephants, many of which were orphaned— surplus elephants originating from culling operations in kruger national park (garaï et al. 2004; slotow et al. 2005). as these small reserves are enclosed by electric fences which prevent any natural immigration & emigration, they need to be actively managed (slotow & van dyk 2004; slotow et al. 2005) to ensure that elephants can meet all of their ecological and social needs (garaï et al. 2004). the basic unit of elephant social organisation is the family group (laws & parker 1968; douglas-hamilton 1973). males born into the effect of mature elephant bull introductions on ranging patterns of resident bulls: phinda private game reserve, south africa heleen druce, k. pretorius, d. druce and r. slotow druce, heleen, k. pretorius, d. druce and r. slotow. 2006. the effect of mature elephant bull introductions on ranging patterns of resident bulls: phinda private game reserve, south africa. koedoe 49(2): 77–84. pretoria. issn 0075-6458. increasing popularity of wildlife viewing has resulted in a rapid increase in small, enclosed reserves in south africa. the african elephant is one of the many species that has been reintroduced into these reserves for eco-tourism. these elephant populations were established as young (<10 years old) orphans from prior kruger national park culling operations. consequently, this abnormal sex and age structure of these introduced populations has influenced their behavioural and spatial ecology. in pilanesberg national park, this abnormal behaviour was corrected by introducing older bulls and culling certain problem elephants. in july 2003, three older bulls (29–41 years old) were introduced into phinda private game reserve, kwazulu-natal, south africa in order to normalise the bull age structure. these introduced bulls were monitored intensively after release, as was the resident bull population, both before and after introduction of the older bulls. the introduced bulls settled into restricted ranges separate from the family groups. all the resident bulls decreased their home ranges at first, with most increasing their home ranges a year later. the resident bulls’ change in ranging patterns was due more to ecological factors than to the influence of the mature bull introduction. this study indicates that the introduction of older male elephants into small populations does not pose major risks or animal welfare concerns. key words: loxodonta africana, adaptive management, movement patterns, kernel ranges, gis. heleen druce, d. druce and r. slotow * (slotow@ukzn.ac.za), amarula elephant research programme, school of biological and conservation sciences, george campbell building, university of kwazulu-natal, durban, 4041 south africa; k. pretorius, phinda private game reserve, kwazulu-natal, south africa (kevinp@ccafrica.com). druce.indd 77 2006/10/15 10:53:43 pm koedoe 49/2 (2006) 78 issn 0075-6458 the family group will leave the group or are forcibly ejected after reaching puberty, which is usually between the age of 14 and 17 years (poole et al. 1984; poole 1989) within these bull groups male association is random and they do not form any long-term bonds with other individuals (laws & parker 1968; moss & poole 1983). phinda private game reserve was established in 1991, with an area of approximately 150 km² (fig. 1). the core of phinda’s elephant population was introduced from kruger national park between 1992 and 1994. these elephants were orphans from prior kruger culling operations, and because equipment to transport elephants was limited to animals shorter than 2 m, all the individuals were young and within the same age class (approximately 10 years old or younger at introduction). during 1994, four adult elephants, two bulls and two cows (approximately 20-25 years of age) were introduced from gonarezhou national park, zimbabwe. during june/july 2003, a total of 37 elephants of four different family groups were translocated from phinda to other reserves. three older bulls (see below) from sabi-sand in limpopo province, south africa were introduced into phinda between july and august 2003. in july 2004, the total elephant population comprised 78 individuals, including 19 adult bulls. prior to the bull introduction in 2003, phinda only had one 36-year-old bull (bull code ph1) and 15 young bulls aged between 15 and 26 that were independent of the breeding herds. this age structure was unnatural because of the large proportion of young males of approximately the same age. a natural population structure should have individuals spread through the different age groups. because the oldest resident bull was approximately ten years older than most of the rest of the phinda bulls, he dominated the other bulls, remaining in musth for long, continuous periods with abnormal displays of aggressive musth behaviour towards other bulls and humans (druce et al. 2006). this was possibly due to the absence of competition or conflict with peers or older bulls. as a result, it was decided to introduce three older bulls to create a more natural age structure n 32o %% % % % % % % % %% % % 27o 500 km 5 km0 0        fig. 1. phinda private game reserve. black dots indicate the position of sites where artificial feed was provided during the winter of 2003. druce.indd 78 2006/10/15 10:53:43 pm issn 0075-6458 79 koedoe 49/2 (2006) within the male population, by filling the age gap of bulls older than 25 years. it was expected they would have an influence on the behavioural, social and spatial ecology of the resident population. according to previous studies (slotow & van dyk 2004), african elephants are sensitive to changes in their social structure. the aims of this study were to determine (1) the effect of the introduction of three older elephant bulls on a young resident bull populations’ spatial ecology, by comparing ranging patterns for selected bulls before, during and after the introduction of new bulls; and (2) how the three new bulls settled into the reserve, by determining their ranging patterns immediately after release and comparing them to those of the resident bulls. methods this study was undertaken in phinda private game reserve, kwazulu-natal, south africa (27º51'30"s, 32º19'00"e) between march 2003 and june 2004. both the forest and savanna biomes are represented within phinda. the vegetation type within the forest biome at phinda is sand forest (low & rebelo 1996, type 3), while the savanna biome within phinda is described by three vegetation types; sweet lowveld bushveld (low & rebelo 1996, type 20), natal lowveld bushveld (low & rebelo 1996, type 26) and coastal bushveld-grassland (low & rebelo 1996, type 23). phinda has a summer rainfall regime and temperatures range from a minimum of 10 ºc in winter to a maximum of 35 ºc in summer. one perennial river, the mzinene river, flows from west to east through the southern section of phinda. during the rainy season, surface water is extensive and during the dry periods, six dams, distributed throughout the property, are supplied with borehole water. bull introductions and monitoring the first two bulls—ph32 aged 29, and ph33 aged 36—were introduced into the electrified phinda holding boma on 12 july 2003. these bulls remained in the boma until their release on 14 july 2003. a third bull—ph31 aged 41—was introduced into the boma on 1 august 2003 and broke out on the evening of 2 august 2003. on the evening of 3 august 2003 he broke out of the reserve though an un-electrified gate. the following day he was recaptured, using immobilisation, and returned to the boma for a further four days before his release into the reserve on 8 august 2003. before his second release, all gates were electrified and no elephants have since broken out of the reserve. all independent adult male and adult female elephants on phinda were identified through unique ear patterns, tusk size and shape as well as any other body characteristics. identity templates were drawn for each individual elephant, including the three older introduced bulls. two of these introduced bulls were fitted with gps collars. these collars download gps positions at set time intervals and transmit the gps points to a ground station using gsm cell-phones. the collars were set to download location points every two hours, which were then stored on a master computer. the phinda elephant population was monitored daily from a vehicle and at each sighting all elephants were identified using the master identification file. at every sighting, date, time, vehicle gps location, animal distance and bearing from the vehicle, total group size, number of males, females and young, habitat type, name of adult individuals present and behaviour codes were recorded. most observations were made from a vehicle on the existing road network, due to the difficulty of driving off road in dense woodland, especially in the north of phinda. with the extensive road network on the reserve, observations was regarded as being sufficient, with a total of 347 sightings for all five resident bulls for the duration of this study period. however, with only two bulls being collared, and located at will, some individual bulls may not have been seen for extended periods. the maximum period that an individual bull was not located was 29 days, but on average, the period between consecutive sightings for an uncollared bull was 3.9 days. ranging patterns in order to determine if the introduction of the three older bulls had any effect on the resident bulls’ ranging patterns, sightings data for the five resident bulls most frequently seen—ph1 age 36, ph2 age 26, ph4 age 26, ph6 age 24, and ph16 age 19—were compared with that of the two collared, introduced bulls (ph31 and ph32). for this study, location data from 1 april 2003 to 30 june 2004 were used. three time periods were selected for each individual bull, correlating with three months before the bull introduction (1 april to 30 june 2003), three months after the older bull introduction (1 august to 31 october 2003), and 9-12 months after the older bull introduction (1 april to 30 june 2004). as there may have been a number of locations recorded for a bull on a druce.indd 79 2006/10/15 10:53:43 pm koedoe 49/2 (2006) 80 issn 0075-6458 ph6 a b c c = 7.3 km² h = 44.5 km² n = 19 c = 8.6 km² h = 38.4 km² n = 17 c = 17.1 km² h = 64.8 km² n = 14 c = 3.9 km² h = 64.4 km² n = 23 c = 3.6 km² h = 31.5 km² n = 47 c = 14.7 km² h = 66.0 km² n = 16 ph16 c = 4.0 km² h = 47.0 km² n = 17 c = 3.3 km² h = 21.3 km² n = 41 c = 16.9 km² h = 69.8 km² n = 19 ph4 a b c c = 9.0 km² h = 45.8 km² n = 22 c = 3.3 km² h = 15.8 km² n = 54 c = 16.7 km² h = 91.0 km² n = 24 ph1 b c c = 3.1 km² h = 31.6 km² n = 18 c = 20.3 km² h = 111.5 km² n = 32 ph31 b c c = 7.4 km² h = 51.3 km² n = 92 c = 5.3 km² h = 45.6 km² n = 91 ph32 b c c = 10.3 km² h = 93.4 km² n = 93 c = 3.6 km² h = 44.2 km² n = 91 0 km a b ca ph2 c = 9.1 km² h = 83.5 km² n = 43 a b c 5 n fig. 2. ranging of bull elephants on phinda: the effect of the introduction of older elephant bulls on the ranging behaviour of younger, resident bulls. three maps are shown for each resident bull and two for the introduced bulls: (a) three months before the introduction of the new bulls, (b) three months directly after introduction, (c) three month period 9-12 months after introduction. c = core ranges (50 % kernel dark grey shading) and h = home ranges (95 % kernel light grey shading). n = number of sightings used to calculate ranges for that period. druce.indd 80 2006/10/15 10:53:44 pm issn 0075-6458 81 koedoe 49/2 (2006) particular day, only the first location after 06:00 each day for individual bulls was used in the analysis. the three-month period one year after the bull introduction was used in the analysis as it correlated with the same season as the previous year immediately before introduction. as a result, any confounding factor of season on bull ranging patterns could be accounted for. the period from june to october 2003 fell within an extremely dry winter, during which a combination of lucerne, eragrostis grass and sugarcane tops were provided at 10 waterholes and at the airstrip on the reserve (fig. 1). all the artificially provided food resources were positioned at water points. we assumed that the elephants would concentrate at or near these water sources due to the drought, with the result that the addition of artificial food would not have an influence on their movement patterns and corresponding home ranges. before analysis, the data set was checked for possible errors or duplicate records. each sighting record was checked on a master spreadsheet against the map for accuracy, with any outlying gps points or sightings that did not match the road name description being corrected and/or deleted. data were processed in microsoft excel and imported to arcview 3.2 (esri). the animal movement analysis arcview extension (hooge & eichenlaub 1997) was used in all gis analyses to estimate the ranges, with the 95 % kernel being used as the estimate of home range and the 50 % kernel as an estimate of core range (burt 1943; worton 1989; seaman & powell 1996). a least squares cross-validation (lscv) smoothing factor of 1000 m was used throughout all gis analyses. separate maps were produced for the various three-month periods for each individual bull. the core and home ranges of individuals were compared between the three month period before introduction and three months directly after introduction, as well as between the three months before introduction with the same three month period one year later. only ranges produced from more than 14 sightings were used in the statistical analysis. seaman et al. (1999) state that kernel home ranges constructed with less than 30 points result in larger home range estimates. however, for our data, there was no significant effect of sample size on home range size (linear regression: f1.17 = 0.16, p = 0.96. data normal: k-s: p = 0.695). we also confirmed no significant non-linear relationships. therefore the kernel analysis was used in this study as a comparison to assess influences specific to this study site and population over this time period. all sample sizes are presented with the ranges to allow readers to make independent assessments of the interpretations, and interpretations are cautious as sample size was relatively small. percentage overlap in ranges were calculated by overlaying maps of the prior three month period with maps of the later three month period and then dividing the area of overlap by the prior area (multiplied by 100), to give a percentage overlap area. results overall, the home ranges of the five resident bulls were concentrated mainly in the north of the reserve (fig. 2), although they also utilised the central areas in the south. during the months immediately after the introduction of the three bulls, the home ranges of resident bulls became much smaller and concentrated around water sources. during the three months prior to the introduction of the new bulls and a year after the introduction, ph1, the oldest resident bull, had a large home range (fig. 2) that covered a large proportion of the reserve. for the three-month period after the introduction of the older bulls (fig. 2), his home range included areas in the north and south. during all three of these study time periods, ph1 spent an average of 39 days in musth, while for the period immediately after the introduction, ph1 was only in musth for 19 days (druce et al. 2006). this may explain why his home ranges differed in size and distribution from that observed for both the other three-month periods, when he was in musth for almost the entire period. during musth periods, ph1 was mainly seen following breeding herds which tended to move throughout the reserve and have large home ranges (unpublished data). during times when ph1 was not in musth, he was on his own and did not follow herds. the resident bulls decreased their home ranges (95% kernel) significantly immediately after the introduction of the older bulls (wilcoxon signed ranks, t = -2.023, n = 5, p = 0.043, table 1). all core (50 % kernel) ranges, with the exception of ph6, decreased following the introduction of the older bulls, although this was not significant (wilcoxon signed ranks, t = -1.214, n = 5, druce.indd 81 2006/10/15 10:53:44 pm koedoe 49/2 (2006) 82 issn 0075-6458 p = 0.225). there was a significant increase in both core ranges (wilcoxon signed ranks, t = -2.02, n = 5, p = 0.043) and home ranges (wilcoxon signed ranks, t = -2.02, n = 5, p = 0.043) between the three-month period before the bull introductions with the three month period 9–12 months later (table 1). we assessed range shift by contrasting overlap of ranges relative to the period before the introduction. there was no significant difference in the overlap of the home (wilcoxon signed ranks: t = -0.674, n = 5, p = 0.50) or core (t = -1.753, n = 5, p = 0.080) ranges in the three months after introduction vs 9–12 months after introduction (table 1). this implies that the resident males did not shift their ranges in response to the introduction of the older bulls. the two introduced bulls utilised areas on the reserve not frequently utilised by the resident bulls. the oldest bull, ph31, patrolled the western boundary for the duration of the first three months (fig. 2). the second introduced bull, ph32, had a home range covering most of the reserve during the three-month periods after introduction (fig. 2). during the first three months after introduction, ph32 had two core ranges, one in the north and the other in the south. a year later, his core range had shifted to the far southern corner of the reserve (unpublished data). the third uncollared introduced bull (ph33) was not used in this analysis, but observational data shows that he associated closely with ph31, the oldest introduced bull (unpublished data). although these two bulls were introduced on different dates into the reserve, they joined up to form a bull coalition and were only seen apart during each other’s musth periods. discussion with the exception of pilanesberg national park (slotow et al. 2000; slotow & van dyk 2001; dickerson 2004; slotow & van dyk 2004), no studies have been done in small, enclosed reserves to determine the effect and success of older bull introductions into established elephant populations. this study was, therefore, important to determine the success of the introduction of the older bulls to phinda, their effect on the ranging patterns of the resident bulls and to allow comparison with the only other similar study that had been previously undertaken. the decrease in the resident bulls home ranges during the three-month period immediately after the introduction, may be due to a seasonal influence rather than the introduction per se. the home ranges of resident bulls were concentrated around water resources for table 1 effect of bull introductions on ranging patterns of resident bulls and changes in ranging patterns of introduced bulls % overlap of range 3 months after introduction with range 3 months before introduction range change % overlap of range 9-12 months after introduction with range 3 months before introduction range change bull corea homeb core home core home core home ph1 33 28 decreased decreased 6 81 increased increased ph2 75 31 decreased decreased 14 54 increased increased ph4 31 33 decreased decreased 44 75 increased increased ph6 36 46 increased decreased 35 43 increased increased ph16 83 35 decreased decreased 30 68 increased increased acore = 50% kernel bhome = 95% kernel druce.indd 82 2006/10/15 10:53:44 pm issn 0075-6458 83 koedoe 49/2 (2006) this three-month period as it fell within an extremely dry winter. the newly introduced bulls showed larger home ranges during this three-month period immediately after their release. these larger home ranges can possibly be explained as exploration of their new area (similar to findings in pilanesberg: slotow & van dyk 2004). however, it is interesting that the core ranges for the older introduced bulls during the three-month period immediately after introduction was also concentrated around water sources. the resident bulls knowledge of local resource distribution might have been a reason why they did not link up with the older bulls during the introduction period. seasonal factors surely seemed to have affected the resident bulls home ranges more than the introduction of the older bulls. interestingly enough, all the resident bulls avoided the older introduced bulls’ core ranges completely and showed only a small overlap with introduced bulls’ home ranges. therefore, the introduction of the older bulls could have influenced the resident bulls in that they avoided the older new bulls. however, the older introduced bulls may also have used spatially separate areas to other elephants, in setting up independent bull areas away from the female family groups. the older bulls successfully managed to establish themselves into the population. the introduction helped in normalising the age structure (slotow et al. 2005), although some of the younger bulls will still need to be removed in order to produce a more normal bull population. in phinda the resident bull population has remained unchanged until present, although in pilanesberg, all the problem animals were removed from the reserve (slotow et al. 2000; dickerson 2004). all the introduced bulls in pilanesberg were older than the resident bull population, while in phinda only one introduced bull was older than the oldest resident bull. both these factors appear to have contributed to pilanesberg’s immediate success. the introduction of older bulls into phinda resulted in no major disruptions to the resident bull behaviour, with only subtle effects on the resident bulls’ ranging patterns. the resident bulls’ ranging changes responded more to ecological factors than to the influence of the mature bull introduction. these results, in combination with those from the pilanesberg introduction (slotow & van dyk 2004), indicate that the introduction of older male elephants into small populations does not pose major risks or animal welfare concerns. acknowledgements this study was funded by a donation from distell (pty) ltd to the amarula elephant research programme, university of kwazulu-natal and by the conservation internship programme of the wildlands conservation trust. a special thanks to all the interns who helped with data collection. ccafrica, phinda private game reserve provided accommodation and logistical support, while clive and allan are thanked for maintaining the research vehicle. this work forms part of an nrf grant (gun number 2053623) to rs. references burt, w.h. 1943. territoriality and home range concepts as applied to mammals. journal of mammalogy 24: 346–352. dickerson, t. 2004. the effect of older elephant bull introduction on resident young orphan bull social dynamics: pilanesberg national park. bsc hons thesis, university of kwazulu-natal, durban. douglas-hamilton, i. 1973. on the ecology and behaviour of the manyara elephants. east african wildlife journal 11: 401–403. druce, d., h. genis, j. braak, s. greatwood, a. delsink, r. kettles, l. hunter & r. slotow. 2004. population demography and spatial ecology of a reintroduced lion population in the greater makalali conservancy, south africa. koedoe 47(1): 103–118. druce, h., k. pretorius, d. druce & r. slotow. 2006 (in press). the effect of mature elephant bull introduction on resident bull's group size and musth periods: phinda private game reserve. south african journal of wildlife research 36(2). garaï, m.e., r. slotow, r.d. carr & b. reilly. 2004. elephant reintroductions to small fenced reserves south africa. pachyderm 37: 28–36. hooge, p.n. & b. eichlenlaub. 1997. animal movement extension to arcview version 1.1. alaska biological science center. anchorage: u.s. geological survey. druce.indd 83 2006/10/15 10:53:44 pm koedoe 49/2 (2006) 84 issn 0075-6458 laws, r.m. & i.s.c. parker. 1968. recent studies on elephant populations in east africa. symposia of the zoological society of london. 21: 319–359. low, a.b. & a.g. rebelo (eds.). 1996. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. moss, c.j., & j.h. poole. 1983. relationship and social structure in african elephants. pp. 315– 325. in: r.a. hinde (ed.). primate social relationships: an intergrated approach. oxford: blackwell. poole, j.h. 1989. announcing intent: the aggressive state of musth in african elephant. animal behaviour 37: 140–152. poole, j.h., l.h. kasman, e.c. ramsay & b.l. lasley. 1984. musth and urinary testosterone concentrations in the african elephant (loxodonta africana). journal of reproduction and fertility 70: 255–260. seaman, d.r. & r.a. powell. 1996. an evaluation of the accuracy of kernel density estimators for home range analysis. ecology 77: 2075–2085. seaman, d.r., j.j. millspaugh, b.j. kernohan, g.c. brundige, k.j. raedeke & r.a. gitzen. 1999. effects of sample size on kernel home range estimates. journal of wildlife management 63: 739–747. slotow, r. & g. van dyk. 2001. role of delinquent young “orphan” male elephants in high mortality of white rhinoceros in pilanesberg national park, south africa. koedoe 44(1): 85–94. slotow, r. & g. van dyk. 2004. ranging of older elephants introduced to an existing small population without older males: pilanesberg national park. koedoe 47(2): 91–104. slotow, r., g. van dyk, j.h. poole, b. page & a. klocke. 2000. older bull elephants control young males. nature 408: 425–426. slotow, r., m.e. garaï, b. reilly, b. page & r.d. carr. 2005. population dynamics of elephants re-introduction to small fenced reserves in south africa. south african journal of wildlife research 35: 23–32. worton, b.j. 1989. kernel methods for estimating the utilization distribution in home-range studies. ecology 70: 164–168. druce.indd 84 2006/10/15 10:53:44 pm abstract introduction methods results discussion conclusion acknowledgements references about the author(s) phemelo gadimang department of wildlife and national parks, gaborone, botswana okavango research institute, university of botswana, botswana gaseitsiwe s. masunga okavango research institute, university of botswana, botswana citation gadimang, p. & masunga, g.s., 2017, ‘ground survey of red lechwe in the linyanti swamps and chobe floodplains, northern botswana’, koedoe 59(2), a1413. https://doi.org/10.4102/koedoe.v59i2.1413 research project no.: ewt 8/36/4 xx (76) original research ground survey of red lechwe in the linyanti swamps and chobe floodplains, northern botswana phemelo gadimang, gaseitsiwe s. masunga received: 08 june 2016; accepted: 14 mar. 2017; published: 23 may 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract a ground survey of red lechwe was carried out in the linyanti swamps and the chobe floodplains of northern botswana in the dry and wet seasons of 2012 and 2013, respectively. we documented numbers, sex ratio and age structure of red lechwe within the linear strips of 25 km × 300 m along the linyanti swamps and the chobe floodplains. results indicated a significant difference in the numbers of red lechwe between sites and seasons. about 66 and 755 red lechwe were estimated for chobe in the dry and wet season, respectively, with 343 and 261 of them estimated for linyanti in the dry and wet season, respectively. in chobe, the red lechwe densities varied widely between seasons (9 red lechwe/km2 – 101 red lechwe/km2) compared with linyanti, where the densities did not vary much between seasons (35 red lechwe/km2 – 46 red lechwe/km2). the lower densities of red lechwe in chobe in the dry season when compared with the wet season suggest a possible seasonal shift in the distribution of red lechwe to the nearby zambezi floodplains in namibia. conservation implications: the higher number of red lechwe in the chobe floodplains in the wet season indicates the potential of the floodplains as a habitat for this species in that season. the dry season shift in the distribution of red lechwe in chobe presents an opportunity for local communities in namibia to engage in tourism, whereas the return of the red lechwe to the floodplains in the wet season ensures protection of the animals as well as boosts the tourism potential of the chobe national park. introduction red lechwe (kobus leche leche gray) is one of the water-dependent terrestrial ungulates that have declined drastically over the past years over its range in southern africa (dipotso & skarpe 2006; iucn 2015). in botswana, it is reported to have declined by 57% – 59% in the last 15 years, and now fluctuates between 26 000 and 33 000 animals (chase 2011; dwnp 2012). although the population is stable in the wetlands of okavango delta, it has reached lowest numbers in decades in the linyanti swamps and the chobe floodplains which are a part of the upper zambezi river basin (chase 2011; dwnp 2012). the reduction in the red lechwe numbers in the chobe floodplains is attributed to human disturbance and competition for food and space from other ungulates that largely converge in large numbers along the chobe floodplains in the dry season for the year-long surface water provided by the river (dipotso & skarpe 2006; williamson 1990). in semi-arid environments, wetlands such as the okavango delta, the linyanti swamps and the chobe floodplains provide a lifeline to many aquatic, semi-aquatic and terrestrial animals. however, these wetlands and the other ones in semi-arid regions are particularly vulnerable to drought, climate change and human pressure (dugan 1992; von richter 1974). because of human pressure and climatic effects, wetlands are now among the most threatened ecosystems of the world (dugan 1992; williams 1993). populations of animal species that are dependent on these wetlands for water and forage resources also end up being affected by changes in the availability and quality of these resources (lin & batzli 2001). for example, non-migratory aquatic antelopes such as red lechwe may die in large numbers as depletion of surface water and food resources increases because of climate change effects and human pressure (bell et al. 1973; dobrouruka 1980; williamson 1990). red lechwe prefers nutrient-rich aquatic grasses and sedges, and its population size often reflects the quality of the wetlands (williamson 1990). therefore, interspecific competition for forage can restrict their population growth (von richter 1974; williamson 1990). poaching and human disturbance are also key factors whose effects in combination or singular can contribute to a decline in ungulate numbers (dipotso & skarpe 2006; hachileka 2003; o’shaughnessy 2010; spinage, williamson & williamson 1989; von richter 1974; williamson 1981). these disturbance factors can also cause biased sex ratios and age structures, and further suppress juvenile recruitment (dobrouruka 1980; milner, nilsen & andreassen 2007; milner-gulland et al. 2003). along the chobe floodplains in northern botswana, wetland antelopes are continually being disturbed by tourism activities with over 50 motorised boats and 100 trucks being driven daily in and along the chobe floodplains (hachileka 2003). in linyanti, the disturbance is relatively low as less than 25 vehicles a day are driven in the area (mcintyre 2014). the number of vehicles is limited by the long distance and deep sandy roads between linyanti and the nearest towns, as well as the poor visibility caused by thick reeds in the swamps (mcintyre 2014; williamson 1981). we, therefore, undertook a cross-sectional ground survey in both the linyanti swamps and the chobe floodplains to document seasonal differences in numbers, distribution, sex ratios and age structure of the red lechwe populations in these two wetlands. methods study area the study area comprises of the linyanti swamps, which are flooded all year round, and the chobe floodplains, which are sustained by the flood-pulsed backflow from the zambezi river. the two wetlands are located 200 km apart along the botswana-namibia border (hughes & hughes 1992; figure 1). figure 1: map of the study area, showing linyanti swamps (left) and chobe floodplains (top right). data collection data were collected during the period of low flood (july–sept.) in 2012 and of high flood (feb.–may) in 2013. these periods also overlapped with rainfall season (nov.–apr.) and no rainfall season (july–oct.), respectively. a distance sampling line transect method was used (thomas et al. 2010). along the chobe floodplains, the line transect was set between kabulabula and ihaha, and in the linyanti swamps it was set between king’s pool and zibadianja pool. the observation strip at each site covered an area of 7.5 km2 (i.e. 25 km long and 300 m wide). although the use of long and continuous transects may have violated the assumptions of the distance method which requires short and randomly placed transects, we believe the 25 km long transect we used at each site helped to reduce the chances of missing the isolated groups and individuals of this rare or uncommon red lechwe species. the surveys were done for 6 days in each wetland, in one dry season of 2012 from 17 august to 04 september and in one wet season of 2013 from 16 february to 03 march. each survey was carried out from 600 to 1000 h each day. there were two observers and one recorder. the vehicle was driven slowly at 20 km/h to minimise disturbance and was stopped at each sighting of the red lechwe. because of good visibility and low speed of the vehicle, we counted all individuals seen. we used binoculars to determine the distance to the animals, group size, sex and age group of all the individuals sighted, and used a global positioning system unit to record animal locations. juveniles were not sexed because of the difficulty in distinguishing between sexes in this group and were excluded in the calculations of the sex ratios. we classified juveniles as 0–1 year olds, sub-adults as 1–2 year olds and adults as over 2 years old with horns or body fully developed (mason 1990). data analysis the red lechwe population was estimated using distance sampling (ver. 6) set at 95% confidence interval (ci) (thomas et al. 2010). we assumed that red lechwe were randomly placed along the floodplains and swamps, with no limitation to visibility because of habitat cover. the data were further analysed using the generalised liner models (glm), poisson log-linear distribution and wald chi-square tests to determine significant differences in the overall number sighted, number of females, males, juveniles, sub-adults, adults, and sex ratio between the two wetlands, and between wet and dry season. the 6-day data for each survey were aggregated to derive means for the sites. means were considered significantly different at p < 0.05. we used glm because it does not require count data to be transformed as transformation can distort the real picture (o’hara & kotze 2010). results population size and distribution the population estimate for red lechwe in the linyanti swamps was 343 animals (distance sampling 95% ci: 186–634) in the dry season and 261 animals (distance sampling 95% ci: 153–443) in the wet season. this corresponds to a density of 46 red lechwe/km2 in dry season and 35 red lechwe/km2 in wet season. for the chobe floodplains, the red lechwe population was estimated at 66 (95% ci: 23–192) and 755 (95% ci: 257–2219) animals in dry and wet seasons, respectively. thus, the density was 9 red lechwe/km2 in the dry season and 101 red lechwe/km2 in the wet season. the cis of the population estimate for the chobe floodplains did not overlap, indicating no significant difference between dry and wet seasons (figure 2). the means of the observed numbers of red lechwe, however, differed significantly between wetlands (χ2 = 9.60, df = 1, p < 0.05) and seasons (χ2 = 3.91, df = 1, p < 0.05). within the chobe floodplains, the mean number of animals differed significantly between seasons (χ2 = 199.38, df = 1, p < 0.05) with a higher number of red lechwe seen in the wet season than in the dry season. within the wet season, the chobe floodplains had a significantly higher number of red lechwe compared with the linyanti swamps (χ2 = 16.68, df = 1, p < 0.05) (table 1). figure 2: population estimate of red lechwe in the chobe floodplains and the linyanti swamps in the wet and dry season. table 1: population estimate, sex and age ratios of red lechwe (kubus leche leche) in the chobe floodplains and the linyanti swamps, and between seasons. in terms of distribution, red lechwe in the linyanti swamps were evenly distributed across the swamps throughout the year, whereas in the chobe floodplains the animals were confined to one locality near kabulabula in dry season, and widespread in wet season from kabulabula to ngoma (figure 3). figure 3: transect layout (black bold line) and observations of red lechwe made along the (a) the chobe floodplains and (b) the linyanti swamps. sex and age structure the number of females differed significantly between wetlands (χ2 = 13.98, df = 1, p < 0.05) and seasons (χ2 = 4.51, df = 1, p < 0.05), with more females seen in the chobe floodplains than in the linyanti swamps, and in wet than in dry season. in the dry season, more males were sighted in the linyanti swamps than in the chobe floodplains (χ2 = 4.65, df = 1, p < 0.05). within the chobe floodplains, the number of males (χ2 = 49.79, df = 1, p < 0.05), females (χ2 = 149.57, df = 1, p < 0.05) and overall sex ratio (χ2 = 11.10, df = 1, p < 0.05) differed significantly between seasons. in the linyanti swamps, only the number of females (χ2 = 6.04, df = 1, p < 0.05) differed between seasons. there was significant difference in the number of adult red lechwe between wetlands (χ2 = 8.97, df = 1, p < 0.05) and seasons (χ2 = 4.28, df = 1, p < 0.05). number of sub-adults (χ2 = 4.72, df = 1, p < 0.05) only differed between wetlands, whereas that of juveniles differed only between seasons (χ2 = 4.56, df = 1, p < 0.05). there were more adults (χ2 = 135.08, df = 1, p < 0.05) and sub-adults (χ2 = 60.92, df = 1, p < 0.05) in the chobe floodplains than in the linyanti swamps. in the linyanti swamps, more juveniles were seen in the wet than dry season (χ2 = 16.10, df = 1, p < 0.05). between seasons, the change in number of juveniles in the linyanti swamps was 168%, whereas for the chobe floodplains it was only 28% and less significant. age ratios did not differ between the wetlands and seasons. discussion population size and distribution variations observed in the number of red lechwe were influenced by location and season. the flood-pulsed chobe floodplains experienced a larger variation in numbers between seasons when compared with the year-long flooded linyanti swamps. the population estimate of red lechwe within the surveyed 7.5 km2 area of the chobe floodplains ranged between 66 and 755 animals (9 red lechwe/km2 – 101 red lechwe/km2), whereas for the linyanti swamps the estimate ranged between 261 and 343 animals (35 red lechwe/km2 – 46 red lechwe/km2). the year round abundance and availability of surface water and aquatic vegetation around the linyanti swamps probably availed quality forage and habitat conditions needed by red lechwe (von richter 1974; williamson 1981), hence less variation in the densities and distribution between seasons. similarly, in the bangweulu swamps in zambia, when resources are plentiful especially in the wet season, black lechwe tends to be less mobile but evenly distributed with no discrete clusters across the swamps (thirgood et al. 1994). in the early 1980s, williamson (1981) carried out some aerial surveys in which he observed a much larger and more stable population of red lechwe in the linyanti swamps than in the chobe floodplains. therefore, we did not expect the linyanti swamps to have a lower number of red lechwe than the chobe floodplains. unlike ground surveys, aerial surveys can provide a much wider aerial view of the landscape and can pick up groups or individual animals which ground surveys may be limited in picking. thus, it is possible that the lower numbers we observed in our study could be a consequence of limited access and visibility over the linyanti swamps which much of them are on the namibia side. furthermore, anecdotal records indicate the area between selinda camp and the kwando lagoon camp on the western side of the linyanti swamps as another concentration area of red lechwe, a habitat which our transect did not cover. the large seasonal variation in red lechwe numbers in the chobe floodplains was also not expected in our study. the variations in the chobe floodplains could have been caused by a combination of factors that include competition with other ungulates over space and predation rates by lion (panthera leo) which are usually higher in the dry than wet season (halley & mari 2004). these factors have the potential to cause a temporary displacement of red lechwe from the floodplains (dipotso & skarpe 2006; halley & mari 2004; o’shaughnessy 2010; spinage et al. 1989). shortage of quality food can also cause many of the red lechwe to seasonally move to nearby sites which may be offering better quality food (mcintyre 2014). in zambia, black lechwe become highly mobile in the dry season when resources become scarce, and form discrete herds that move quickly across sites in search for green forage (thirgood et al. 1994). in botswana, there are currently no studies reporting on shifts in distribution of red lechwe along the chobe floodplains or nearby zambezi basin. however, observations made through aerial surveys indicate the presence of red lechwe in areas far from the floodplains and in the nearby zambezi floodplains (chase 2011; dwnp 2012), but these surveys did not attribute the sighting of isolated groups to migration or shift in the distribution away from the chobe floodplains. in comparison with aerial surveys, our ground surveys were limited in detecting where the red lechwe were in the dry season in the chobe floodplains. a longitudinal survey would have been much effective in providing a long-term coverage, in-depth understanding and convincing comparison of population dynamics of red lechwe between the two wetlands. despite this limitation, our findings indicate a possibility of dry season emigration of many red lechwe from the chobe floodplains to the nearby zambezi floodplains, and immigration into the chobe floodplains in the wet season. we doubt if red lechwe would move and stay for over a week in the nearby woodlands, given that it is a semi-aquatic species that prefers habitats with plenty of surface water and aquatic forage (bell et al. 1973; von richter 1974). our ground surveys were also able to locate an isolated and larger herd of red lechwe consisting of more than 119 animals in the wet season in the chobe floodplains which had not been detected by any survey done in the last decade (e.g. by chase 2011; dwnp 2012). herd sizes observed in the past surveys were lower (e.g. 41–80 animals) (chase 2011) when compared with those observed in our ground survey done 3 years later. a single group size of 119 red lechwe seen in our ground surveys gives hope that the numbers of red lechwe in the chobe floodplains may still be larger than previously thought, especially when one considers that ground surveys generally under-estimate population sizes. recent aerial surveys had estimated the population of red lechwe in the chobe floodplains to be around 19 animals (95% ci: 0–38) (dwnp 2012), from 399 animals estimated in 2010 (95% ci: 97–701) (chase 2011). sex and age structure sex ratios may not make much sense especially for polygynous ungulates such as red lechwe, where only a few reproductive males can mate and fertilise many females (holand et al. 2006; milner et al. 2007; milner-gulland et al. 2003). however, sex ratios are nonetheless critical as the number of males may reach a threshold below which conception time would be delayed and calving rates reduced (holand et al. 2006; milner et al. 2007). in our study, there was no significant difference in the overall male-to-female ratio between habitats. age ratios also did not differ between sites. the short duration of our study and the limited distribution of the transects in the study sites gave us only a snapshot of the numbers, sex and age structure of red lechwe in these ecologically threatened wetlands, and future surveys that aim to have detailed knowledge of the red lechwe should be long term and also cover the area between selinda and kwando on the western side of the linyanti swamps. conclusion the numbers of red lechwe in the chobe floodplains varied widely between seasons, with higher numbers observed in the wet season than in the dry season. in the dry season, red lechwe were mostly restricted to fewer locations within the floodplains. in the linyanti swamps, the numbers did not fluctuate widely over space and time. the findings of this study suggest that the linyanti swamps and the chobe floodplains still have the potential as habitats for red lechwe, and conservation efforts should be directed at minimising factors that limit forage and surface water availability in the two areas (e.g. in bell et al. 1973). the seasonal variations in the red lechwe population in the chobe floodplains may be because of disturbance from tourism activities, competition with other herbivores, displacement by predators or variability in forage availability because of seasonal rainfall and flood pulse. these possible disturbances to red lechwe need to be investigated further as they have important implications for the conservation of this globally near-threatened antelope. conservation implications a better understanding of the seasonal and spatial distribution of red lechwe in the chobe floodplains is critical to mapping and conserving key resource areas that supplement and complement each other to maintain a viable red lechwe population, and which can subsequently boost tourism on botswana side. the understanding will further guide the nature of conservation projects that may be initiated by local communities living within the kavango-zambezi transfrontier conservation area. it is, therefore, imperative that longitudinal studies on population dynamics of red lechwe should be done. acknowledgements this study was made possible by grants and permits from the university of botswana and the ministry of environment natural resource conservation and tourism. the authors acknowledge the valuable comments, which they received from dr richard w.s. fynn and anonymous reviewers. the office of research and development (ord) at the university of botswana provided funding for fieldwork and the ministry of environment natural resource conservation and tourism provided a scholarship to p.g. for mphil studies. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions p.g. conceived and designed the study, performed fieldwork, analysed the data and wrote the manuscript. g.s.m. supervised the work from conception to manuscript writing. references bell, r.h.v., grimsdell, j.j.r., van lavieren, l.p. & sayer, j.a., 1973, ‘census of the kafue lechwe by aerial stratified sampling’, east african wildlife journal 11, 55–74. https://doi.org/10.1111/j.1365-2028.1973.tb00073.x chase, m., 2011, a dry season fixed-wing aerial survey of elephants and wildlife in northern botswana. september-november 2010, a survey was conducted jointly by elephants without borders, forest conservation botswana and the zoological society of san diego, kasane, botswana, p. 82. dipotso, f.m. & skarpe, c., 2006, ‘population status and distribution of puku in a changing riverfront habitat in northern botswana’, south african journal of wildlife research 36, 89–97. dobrouruka, l.j., 1980, ‘the lechwe waterbuck kobus leche, its current status and preliminary studbook’, international zoo yearbook 20, 298–301. https://doi.org/10.1111/j.1748-1090.1980.tb00999.x dugan, p., 1992, wetlands in danger, mitchell beazley london ltd., london, uk. dwnp, 2012, aerial census of animals in botswana: dry season 2012, department of wildlife and national parks, ministry of environment, wildlife and tourism, gaborone, botswana. hachileka, e., 2003, ‘sustainability of wildlife utilisation in the chobe district, botswana’, south african geographical journal 85, 50–57. https://doi.org/10.1080/03736245.2003.9713784 halley, d.j. & mari, m., 2004, ‘dry season social affiliation of african buffalo bulls at the chobe riverfront, botswana’, south african journal of wildlife research 34, 105–111. holand, ø., weladji, r.b., røed, k.h., gjøstein, h., kumpula, j., gaillard, j.m. et al., 2006, ‘male age structure influences females’ mass change during rut in a polygynous ungulate, the reindeer (rangifer tarandus)’, behavioral ecology and sociobiology 59, 682–688. https://doi.org/10.1007/s00265-005-0097-5 hughes, r.s. & hughes, j.s., 1992, a directory of african wetlands, iucn/unep/wwf/world conservation monitoring center, samara publishing ltd., cardigan, uk. iucn, 2015, the iucn red list of threatened species. 2015–14, iucn, cambridge, uk. lin, y.t.k. & batzli, g.o., 2001, ‘the influence of habitat quality on dispersal, demography and population dynamics of voles’, ecological monographs 71, 245–275. https://doi.org/10.1890/0012-9615(2001)071[0245:tiohqo]2.0.co;2 mason, d.r., 1990, ‘monitoring of sex and age ratios in ungulate populations of the kruger national park by ground survey’, koedoe 33, 19–28. https://doi.org/10.4102/koedoe.v33i1.449 mcintyre, c., 2014, botswana safari guide, okavango delta, chobe, northern kalahari, 4th edn., bradt travel guides ltd., the globe pequot press, inc., ct. milner, j.m., nilsen, e.b. & andreassen, h.p., 2007, ‘demographic side effects of selective hunting in ungulates and carnivores’, conservation biology 21, 36–47. https://doi.org/10.1111/j.1523-1739.2006.00591.x milner-gulland, e.j., bukreeva, o.m., coulson, t., lushchekina, a.a., kholodova, m.v., bekenov, a.b. et al., 2003, ‘conservation, reproductive collapse in saiga antelope harems’, nature 422, 135–135. https://doi.org/10.1038/422135a o’hara, r.b. & kotze, d.j. 2010. ‘do not log-transform count data’, methods in ecology and evolution 1, 118–122. https://doi.org/10.1111/j.2041-210x.2010.00021.x o’shaughnessy, r., 2010, ‘a comparative diet and habitat selection of puku (kobus vardonii) and red lechwe (kobus leche) on the chobe river floodplain, botswana’, msc dissertation, university of witwatersrand, johannesburg, south africa. spinage, c.a., williamson, d.t. & williamson, j.e., 1989, ‘botswana antelopes’, global survey and regional action plans 2, 41–49. thirgood, s.j., nefdt, r.j., jeffery, r.c.v. & kamweneshe, b., 1994, ‘population trends and current status of black lechwe (kobus: bovidae) in zambia’, african journal of ecology 32, 1–8. https://doi.org/10.1111/j.1365-2028.1994.tb00549.x thomas, l., buckland, s.t., rexstad, e.a., laake, j.l., strindberg, s., hedley, s.l. et al., 2010, ‘distance software, design and analysis of distance sampling surveys for estimating population size’, journal of applied ecology 47, 5–14. https://doi.org/10.1111/j.1365-2664.2009.01737.x von richter, w., 1974, ‘survey of the adequacy of existing conserved areas in relation to wild animal species’, koedoe 17, 39–69. https://doi.org/10.4102/koedoe.v17i1.902 williams, m., 1993, wetlands, a threatened landscape, blackwell publishers, oxford, uk. williamson, d.t., 1981, ‘the status of red lechwe in the linyanti swamp’, botswana notes and records 13, 101–105. williamson, d.t. 1990, ‘habitat selection by red lechwe (kobus leche leche gray 1850)’, african journal of ecology 28, 89–101. https://doi.org/10.1111/j.1365-2028.1990.tb01141.x abstract introduction research method and design results and discussion conclusion acknowledgements references appendix 1 about the author(s) anna s. dippenaar-schoeman biosystematics arachnology, arc – plant protection research institute, south africa department of zoology, university of venda, south africa charles r. haddad department of zoology & entomology, university of the free state, south africa robin lyle biosystematics arachnology, arc – plant protection research institute, south africa leon n. lotz department of arachnology, national museum bloemfontein, south africa stefan h. foord department of zoology, university of venda, south africa rudy jocque royal museum for central africa, tervuren, belgium peter webb south african national survey of arachnida, pretoria, south africa citation dippenaar-schoeman, a.s., haddad, c.r., lyle, r., lotz, l.n., foord, s.h. & jocque, r. et al., 2018, ‘south african national survey of arachnida: a checklist of the spiders (arachnida, araneae) of the tswalu kalahari reserve in the northern cape province, south africa’, koedoe 60(1), a1486. https://doi.org/10.4102/koedoe.v60i1.1486 checklist south african national survey of arachnida: a checklist of the spiders (arachnida, araneae) of the tswalu kalahari reserve in the northern cape province, south africa anna s. dippenaar-schoeman, charles r. haddad, robin lyle, leon n. lotz, stefan h. foord, rudy jocque, peter webb received: 08 aug. 2017; accepted: 09 apr. 2018; published: 09 july 2018 copyright: © 2018. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract one of the aims of south african national survey of arachnida (sansa) is to survey protected areas to obtain species-specific information and compile inventories to determine species distribution patterns and evaluate their conservation status for red data assessments. the aim of this study, the first in a series of surveys of the diamond route reserves, was to compile the first checklist of the spider species in the northern cape at the tswalu kalahari reserve. spiders were collected during three survey periods (2005−2013) using different collecting methods to sample both the ground and field layers. in total, 32 families represented by 108 genera and 136 species have been collected so far. the most species-rich families are the salticidae (20 spp.) and thomisidae (18 spp.), followed by the gnaphosidae and araneidae (11 spp. each), while nine families are represented by singletons. the free-living wandering spiders represent 97 spp., while 39 spp. are web-builders. information on spider guilds, endemicity value and conservation status are provided. the tswalu kalahari reserve protects approximately 6.1% of the total south african spider fauna, while 24.3% of the species found in the reserve are south african endemics, of which 5.9% are northern cape endemics. approximately 6.0% of the species sampled are possibly new to science or represent new records for south africa. conservation implications: the tswalu kalahari reserve falls within the savanna biome in the northern cape province. only one spider species was previously known from the reserve; a further 135 spp. are reported for the first time, with 5.9% of the species being northern cape endemics and 24.3% south african endemics. approximately 6.0% of the species may be new to science or represent new records for south africa. introduction the south african national survey of arachnida (sansa) was initiated in 1997, with the main aim of making inventories of the arachnid fauna of south africa (dippenaar-schoeman & haddad 2006; dippenaar-schoeman et al. 2015). sansa has several focus areas, such as arachnid diversity in floral biomes, agroecosystems and protected areas. species distribution data are an essential information resource needed for the conservation assessments used to compile a red data list of the arachnida of south africa (lyle & dippenaar-schoeman 2015). surveys are needed to obtain species-specific information, and yield new, rare and/or endemic species and resources for these existing protected areas. the publication of these species distribution records formed the basis of the first spider atlas and national species list (dippenaar-schoeman et al. 2010; dippenaar-schoeman 2013). this study presents the results of sansa sampling in the tswalu kalahari reserve (tkr), falling within the arid parts of the savanna biome (foord, dippenaar-schoeman & haddad 2011a). the reserve is an e. oppenheimer & son property situated in the northern cape (lyle & dippenaar-schoeman 2013). this is the first survey of the arachnid fauna of protected areas in the northern cape province and the first spider checklist for the tkr. information on spider guilds, their habitat preference, web types, and endemicity index and conservation status are provided. checklists for several of the protected areas in south africa have been published but none for the northern cape (mcgeoch et al. 2011; dippenaar-schoeman et al. 2015). research method and design study area and period tswalu kalahari reserve (27°13’30’’s, 22°28’40’’e; 930 m a.s.l.) is the largest (> 100 000 hectares) privately owned wildlife reserve in south africa. it lies in the northern cape province, at the foot of the korannaberg mountains (figure 1). kuruman is the closest large town, some 140 km east of tswalu. figure 1: (a) map of south africa, showing the locality of the tswalu kalahari reserve in northern cape province; and (b) details of the vegetation units in the reserve. tswalu kalahari reserve includes vegetation described by low and rebelo (1998) as shrubby kalahari dune bushveld, kalahari plains bushveld and kalahari mountain bushveld areas of the savanna biome (figure 2a–d). acocks (1988) described the area as kalahari thornveld. the reserve is characterised in certain areas by scattered shrubs and well-developed grass layers, in other areas by a well-developed tree layer and moderately developed grass and shrub layers, and by a poorly developed tree layer and moderately developed grass layers on the mountains and hills (van rooyen 1999). some dominant plant species include the trees vachellia erioloba, boscia albitrunca and terminalia sericea. the four main soil types in the tkr are poorly structured red soils with a high base status; well-drained red, sandy soils with a high base status; red and yellow, well-drained sandy soils with a high base status; and rocky areas with little or no soil (van rooyen 1999). figure 2: habitat types in the tswalu kalahari reserve: (a) well-developed grass layer near the foothills of the korannaberg mountains; (b) kalahari thornveld; (c) sand dune with poorly developed tree layer and moderately developed grass; and (d) one of the south african national survey of arachnida sampling sites. the climate of tkr is highly variable and falls in the summer rainfall area of southern africa (low & rebelo 1998), with a relatively high rainfall occurring from october to april but with a distinct peak in march. the mean annual rainfall is 253.3 mm. the dry season occurs from may to september, with an average of less than 10.0 mm during this period. the peak dry season occurs from june to august, with little or no rainfall. sampling methods and identification material from three surveys (table 1) was used to compile the first checklist of the spiders of tkr (appendix 1). during the first visit to the reserve, spiders were collected ad hoc in all five habitats in the reserve (figure 1) using a variety of methods, and no set protocol was followed. the second and third surveys were carried out using the standardised protocol devised for sansa and described in detail by haddad & dippenaar-schoeman (2015). it can be briefly summarised as follows: four representative habitats in a selected degree-square grid were selected by the field work manager, in this case the third author, and sampled by a team of four collectors. during the second and third surveys, sampling was carried out in grass layer around hills forming part of the korannaberg–langeberg mountain bushveld, olifantshoek plains thornveld, gordonia plains shrubland and kathu bushveld (figure 1). table 1: details of spider surveys undertaken at the tswalu kalahari reserve during three field trips, including some of the authors of this article. in each of these habitats sampled using the sansa protocol, 500 beat samples were taken from woody vegetation using a beating sheet and beating stick; 500 sweep samples were taken from grasses and herbaceous vegetation using a sweep net; 50 pitfall traps (buckets with diameter of 135 mm) were set out 2 m apart and kept open for 3–4 days; ten leaf litter samples were taken and sifted over a white sheet using a steel sieve with a mesh spacing of 9 mm. further, in each habitat, all four team members conducted 2 h of hand collecting during the day from beneath logs, rocks and bark and from vegetation. night collecting (2 h per person) was done in all four habitats, as opposed to the single habitat required by the sansa protocol. winkler traps were used to extract leaf litter samples taken in a single habitat (olifantshoek plains thornveld) during the second survey only; this method yielded poor results and was not used during the third survey. all of the material sampled for each of the above methods was preserved in 70% ethanol, except for pitfall traps, in which propylene glycol was used as a preservative. once the pitfalls were removed from the soil, the material was sieved, and the arachnids removed and preserved in 70% ethanol. species determinations were performed by several of the authors. voucher specimens are deposited in the national collection of arachnida housed at the arc-plant protection research, pretoria (nca), and at the national museum in bloemfontein (nmba). only the generic names were included in the checklist when immature specimens were sampled and in those cases where the family lacks taxonomic resources to make species level identifications possible. functional groups spiders often live in distinct microhabitats with limitations imposed by contrasting biotic and abiotic factors. species can be categorised into particular functional groups or guilds, based on our knowledge of their habitat and microhabitat preferences, as well as their diets and hunting strategies (foord et al. 2011a). this provides valuable ecological information that helps in better understanding the utilisation of habitat structures by different taxa. in general, spiders can be divided into species that are largely or entirely reliant on silk to construct webs to capture prey (web-builders, wb) and those that actively search for prey or ambush prey from burrows or on vegetation (wanderers, w). each of these two major guilds is divided into several subcategories based on the substrates they utilise or the web types that they construct (table 2). table 2: detailed descriptions of foraging guilds that were assigned to spider species sampled in the tswalu kalahari reserve. endemicity value the conservation status of species is important, and as part of the first atlas of south african spiders (dippenaar-schoeman et al. 2010), an endemicity index was provided for each species (table 3, appendix 1) based on its current distribution. seven endemicity categories were considered: 6 = endemic, known only from type locality or one locality only; 5 = known from one province only, wider than type locality; 4 = known from two adjoining provinces only; 3 = south africa, known from more than two provinces or two provinces not adjoining; 2 = southern africa (south of zambezi and kunene rivers); 1 = afrotropical region; 0 = africa and wider. table 3: level of endemicity of the 136 spider species sampled at the tswalu kalahari reserve. regarding conservation status, species that were only recorded from immatures or that represent new taxa were not evaluated, and are considered to be data deficient for taxonomic reasons (ddt). species with a broad distribution (categories 0–2) were considered to be of least concern (lc); those of categories 3 and 4 were considered to be south african endemics (sae); and those of category 5 were considered to be northern cape endemics (nce). no reserve endemics (re, category 6) have been recorded from tkr yet. photography as part of sansa, a photographic virtual museum was developed to access photographs of arachnid species (dippenaar-schoeman, lyle & van den berg 2012; dippenaar-schoeman et al. 2015). spiders sampled during the last surveys at tkr were photographed by the last author. a photo gallery of the spiders will be made available on the sansa website. images can also be viewed at http://www.arc.agric.za:8080/default. ethical considerations permission to collect arachnids in the northern cape province was obtained from the northern cape department of environment and nature conservation. results and discussion spider biodiversity and endemicity thirty-two spider families represented by 108 genera and 136 spp. were collected from tkr between 2006 and 2013 over a total of 16 sampling days (appendix 1, table 4). except for one species, tusitala barbata peckham & peckham, 1902 (salticidae), the rest of the species are reported from the reserve for the first time (azarkina & foord 2015). although the northern cape is south africa’s largest province, covering 29.7% of the land area, only 1990 records sampled from 124 sites in the northern cape are accessioned in the sansa database, represented by 490 spp. from 49 families (dippenaar-schoeman et al. 2015). table 4: spider diversity of tswalu kalahari reserve, with total number of families, genera and species sampled. the northern cape province has been less intensively sampled than the other provinces. except for the field guide on the spiders of the kalahari (dippenaar-schoeman & van den berg 2010), no surveys from protected areas in the province have been published. several surveys are underway in reserves (benfontein, rooipoort and oryx nature reserves) and in the augrabies, richtersveld and namaqua national parks (lyle & dippenaar-schoeman 2013; dippenaar-schoeman 2014a). the only published results are surveys in pistachio orchards in the arid nama karoo near prieska (haddad & dippenaar-schoeman 2005, 2006; haddad, dippenaar-schoeman & pekár 2005; haddad, louw & dippenaar-schoeman 2004; haddad, louw & pekár 2008), where a total of 143 spp. from 31 families were collected (foord et al. 2011a). in a second study, lyons (2009) conducted a broad-scale survey of arthropods in restored alluvial diamond mining sites in the succulent karoo of the northern cape, in which 21 spider families and 51 spp. were sampled. based on these results and information from the sansa database, the number of species sampled in reserves and parks in the northern cape is much lower (80–140 spp.) compared to limpopo reserves, which average 228 spp. per reserve, ranging between 175 and 286 spp. (foord et al. in prep.). of the 136 spp. sampled, ten spp. (7.4%) were ddt and could not be identified to species level, of which four spp. were immature and six spp. are possibly new to science (appendix 1, table 3). however, these putative new species are representatives of species-rich families, and only after revisionary studies would it be possible to tell whether they are indeed new to science. no species sampled from tkr thus far can be considered re. only the south african endemic species falling into categories 3–6 (33 spp., 24.3%) need to be evaluated using the iucn criteria. the majority of the species sampled (93 spp.) can be listed as lc, having a distribution throughout southern africa or wider (table 3). seven northern cape endemic species are protected in the tkr: ancylotrypa pusilla purcell, 1903 (cyrtaucheniidae) (figure 3a); dresserus laticeps purcell, 1904 (eresidae) (figure 3b); allocosa aurichelis roewer, 1959 (lycosidae) (figure 3c); aelurillus cristatopalpus simon, 1902 (salticidae); evarcha brinki haddad & wesołowska, 2011 (salticidae); ariadna jubata purcell, 1904 (segestriidae) (figure 3d) and histagonia deserticola simon, 1895 (theridiidae). figure 3: spiders of the tswalu kalahari reserve: (a) ancylotrypa pusilla (cyrtaucheniidae); (b) dresserus laticeps (eresidae); (c) allocosa aurichelis (lycosidae); (d) ariadna jubata (segestriidae); (e) nest of immature thyene imperialis (salticidae); (f) harpactirella lapidaria (theraphosidae); (g) hogna transvaalica (lycosidae); (h) ammoxenus coccineus (ammoxenidae); (i) loxosceles simillima (sicariidae); (j) sicarius testaceus (sicariidae); (k) peucetia viridis (oxyopidae); (l) hersilia sericea (hersiliidae); (m) isoxya mossamedensis (araneidae); (n) nephila senegalensis (araneidae); and (o) latrodectus geometricus (theridiidae). during this study, ibala okorosave fitzpatrick, 2009 (gnaphosidae) was recorded from south africa for the first time, and the first adult specimens of the monotypic genus mallinus simon, 1893 (zodariidae) were also sampled. currently, 2240 spider species are known from south africa (dippenaar-schoeman 2017), and thus, 6.1% of south african species are protected in this reserve. family diversity results from the savanna biome indicate that four spider families consistently dominate assemblages in terms of species richness (foord, dippenaar-schoeman & haddad 2011b; dippenaar-schoeman, foord & haddad 2013): araneidae, gnaphosidae, salticidae and thomisidae. in this study, the salticidae (20 spp.), thomisidae (18 spp.), gnaphosidae (11 spp.) and araneidae (11 spp.) were the most species-rich families (table 4), consistent with patterns in the savanna biome. nine families are represented by singletons. salticidae: the salticidae are free-living spiders found on vegetation and the soil surface. they build small silk nests attached to various substrates, in which they moult, oviposit and sometimes mate, or which they occupy during periods of inactivity (dippenaar-schoeman & van den berg 2010; dippenaar-schoeman 2014b). during the last survey of this study, a small round densely woven silk retreat attached to grass (figure 3e) was sampled in the tkr, housing an immature thyene imperialis (rossi 1846). one species has been identified as belonging to a new genus (galina azarkina, pers. comm.) and one was immature. the other 17 are new records for the tkr, five spp. are sae, two spp. are nce, while 11 spp. are more widely distributed throughout africa (appendix 1). thomisidae: crab spiders are free-living spiders commonly found on grass, shrubs, flowers and trees, and only few species were sampled from the soil surface (dippenaar-schoeman & van den berg 2010; dippenaar-schoeman 2014b). thomisids are easily dispersed by wind and most species have a wide distribution. in the tkr, 13 genera represented by 18 spp. were sampled. of these, only four spp. are known sae, while the rest (14 spp.) are widely distributed throughout africa (appendix 1). gnaphosidae: the gnaphosids are free-living spiders commonly found on the ground and low vegetation (dippenaar-schoeman & van den berg 2010; dippenaar-schoeman 2014b). one species could not be determined, five of the 11 spp. are sae, and the rest have a wide distribution. one species, aneplasa nigra tucker, 1923, has a restricted distribution and is known from the northern and western cape provinces only (appendix 1). araneidae: the araneidae are web-builders and produce typical orb-webs (owb) and modified orb-webs (mowb) (dippenaar-schoeman & van den berg 2010; dippenaar-schoeman 2014b). all the members of the family (11 spp.) recorded here have a wide african distribution. functional groups for this study, two main guilds were recognised, namely wandering spiders (w) (97 spp.) and web-builders (wb) (39 spp.), with further subdivisions based on microhabitat and general behaviour, as observed during surveys (appendix 1). wanderers: a total of 97 spp. (71.3%) are wandering spiders, with some species living on vegetation (39 spp.) and others on the ground surface (55 spp.), with an additional three species occurring regularly in both strata. the majority of ground-dwellers are free-living soil dwellers (58 spp., 42.6%), while six spp. (4.4%) live in burrows. the salticidae (15 spp.), gnaphosidae (11 spp.) and lycosidae (six spp.) are the most species rich families of ground-dwellers (appendix 1). most of the burrow-dwellers belong to the suborder mygalomorphae and include the trapdoor spider species ancylotrypa pusilla (figure 3a), a bag-nest migid, moggridgea peringueyi simon, 1903, and three theraphosid baboon spider species (figure 3f). one species of wolf spider, possibly hogna transvaalica (simon, 1898) (lycosidae) (figure 3g), also constructs burrows. these spiders use their bright red cheliceral setae to scare off predators (webb 2013). a species of the termite-eating spider, ammoxenus coccineus simon, 1893 (ammoxenidae), was sampled from loose sand (figure 3h). ammoxenids are specialist termite-feeders (petráková et al. 2015) and use the strong setae on their chelicerae to dive into the sand (dippenaar-schoeman, de jager & van den berg 1996a; dippenaar-schoeman, de jager & van den berg 1996b). two species of medical importance were sampled at tkr, the violin spider loxosceles simillima lawrence, 1927 (sicariidae) (figure 3i) and the six-eyed sand spider sicarius testaceus purcell, 1908 (sicariidae) (figure 3j). the plant wanderers sampled from the grass and tree layer are represented by 42 spp. (30.9%). the thomisidae (16 spp.), salticidae (eight spp.) and oxyopidae (six spp.) were the most diverse plant-dwellers found on grasses, shrubs and trees. three salticid species occur both on the ground and on vegetation (appendix 1). some interesting results have already been published regarding the presence of peucetia viridis (blackwall, 1858) (figure 3k) of the family oxyopidae, which was sampled from the unpalatable kalahari sour grass (bushman’s grass), schmidtia kalahariensis. this annual grass is only available for a short period after good rains. it has an unpleasant smell and is covered with glands that produce an acidic substance. during the survey in 2008, this was the dominant grass present and it was intensively swept, but only this one species was recorded from the grass (dippenaar-schoeman 2005; vasconcellos-neto et al. 2007). several species were sampled from trees, including the long-spinnered bark spiders, hersilia sericea pocock, 1898 (hersiliidae) (figure 3l), and the community nest spiders, stegodyphus dumicola pocock, 1898 (eresidae). web-dwellers: the web-dwellers are represented by 39 spp. (28.7%), with the largest number making owb or mowb (14 spp., 10.3%), followed by gumfoot-webs (eight spp., 5.9%), retreat-webs (eight spp., 5.9%), funnel-webs (five spp., 3.7%), sheet-webs (three spp., 2.2%) and space-webs (one sp., 0.7%). the physical structure of the habitat plays a role in the composition of the web-dwelling fauna, as it not only provides the necessary support for anchoring webs but also increases the availability of retreat space and modification of the microclimate, which could have an effect on spiders, as well as their prey. most of the owb recorded belong to the araneidae (11 spp.) (figure 3m and n), which construct large orb-webs between trees and shrubs. some of these species are diurnal and they are found in their webs during the day. some orb-web builders are associated with grasslands (araneus, larinia, nemoscolus and neoscona) and are mostly nocturnal, making their orb-webs at night and resting in retreats, usually constructed in grass inflorescences, during the day. one species is a mowb, the tropical tent-web spider (cyrtophora citricola [forsskål, 1775]). several gumfoot-web spiders of the theridiidae (eight spp.) were sampled, including two button spiders that are of medical importance, latrodectus geometricus c.l. koch, 1841 (figure 3o) and l. renivulvatus dahl, 1902. conclusion as signatories to the convention on biodiversity, south africa has an obligation to develop a strategic plan for the conservation and sustainable utilisation of its fauna and flora. preliminary investigations into the biodiversity of the south african arachnida highlighted the obstacles caused by a lack of baseline biodiversity and ecological information for many of the arachnid orders (dippenaar-schoeman 2002). with this in mind, each biodiversity survey contributes to improving our knowledge of the geographical distribution and biology of south african spider species. this survey forms part of the sansa for the savanna biome, as well as the northern cape province, and as such represents new provincial records for 102 species. although this article probably represents only a portion of the spider fauna present, we hope that this information will stimulate further interest and research. established reserves, such as tkr, can make a substantial contribution towards invertebrate conservation. however, the contribution of existing reserves can only be highlighted through studies such as this. acknowledgements the authors would like to thank the agricultural research council (arc) and the south african national biodiversity institute’s (sanbi) threatened species programme for funding the south african national survey of arachnida (sansa) phase 2; duncan macfadyen of e. oppenheimer & son trust for providing permission to sample in tswalu kalahari reserve and the officials of tswalu for their friendliness and assistance; the staff of the arachnology section of the biosystematics programme, arc – plant protection research, notably connie anderson, sma mathebula and petro marais, for their assistance with processing the material collected; elisabeth tybaert (wife of rudy jocqué), petro marais and michael stiller (arc) for assisting during the fieldwork; and galina azarkina for assistance with the salticidae identifications. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions all the authors are team members of sansa and contributed towards planning this national survey. they participated in field work, identifications of specimens and curation of material. a.s.d-s. and r.l. were involved in surveys, identifications and preparation of the manuscript; c.r.h. assisted with editing the manuscript; l.n.l. and r.j. assisted with field surveys; p.w. participated in field surveys and photographed all the specimens funding information the first, second and fifth authors acknowledge financial support from the national research foundation of south africa. references acocks, j.p.h., 1988, ‘veld types of south africa’, 3rd edn., memoirs of the botanical survey of southern africa 57, 1–146. azarkina, g.n. & foord, s.h., 2015, ‘a review of three tusitala (araneae: salticidae) species from southern africa, with a new synonymy and description of a new species from botswana’, african invertebrates 56, 285–307. https://doi.org/10.5733/afin.056.0204 dippenaar-schoeman, a.s., 2002, ‘status of south african arachnida fauna’, proceedings of the symposium on the status of south african species organized by the endangered wildlife trust (ewt) of south africa, rosebank, september 4–7, 2001, pp. 70–81. dippenaar-schoeman, a.s., 2005, ‘interesting behaviour of a green lynx spider collected at tswalu game reserve (araneae: oxyopidae: peucetia)’, spider club news 20, 8–9. dippenaar-schoeman, a.s., 2013, ‘first national species lists of south african arachnids available’, plant protection news 95, 6. dippenaar-schoeman, a.s., 2014a, ‘what is new in sansa – and plans for 2014’, sansa news 19, 3–6. dippenaar-schoeman, a.s., 2014b, field guide to the spiders of south africa, lapa publishers, pretoria. dippenaar-schoeman, a.s., 2017, ‘feedback on the spider red listing project (srlp)’, sansa news 29, 3. dippenaar-schoeman, a.s., de jager, m. & van den berg, a., 1996a, ‘behaviour and biology of two species of termite-eating spiders, ammoxenus amphalodes and a. daedalus (araneae: ammoxenidae), in south africa’, african plant protection 2, 15–17. dippenaar-schoeman, a.s., de jager, m. & van den berg, a., 1996b, ‘ammoxenus species (araneae: ammoxenidae) – specialist predators of harvester termites in south africa’, african plant protection 2, 103–109. dippenaar-schoemanl, a.s., foord, s.h. & haddad, c.r., 2013, spiders of the savanna biome, university of venda, thohoyandou & agricultural research council, pretoria. dippenaar-schoeman, a.s. & haddad, c.r., 2006, ‘what is the south african national survey of arachnida (sansa) all about?’, sansa news 1, 1–3. dippenaar-schoeman, a.s., haddad, c.r., foord, s.h., lyle, r., helberg, l. & mathebula, s., 2010, first atlas of the spiders of south africa (arachnida: araneae), arc – plant protection research institute, pretoria. dippenaar-schoeman, a.s., haddad, c.r., foord, s.h., lyle, r., lotz, l.n. & marais, p., 2015, ‘south african national survey of arachnida (sansa): review of current knowledge, constraints and future needs for documenting spider diversity (arachnida: araneae)’, transactions of the royal society of south africa 70, 245–277. https://doi.org//10.1080/0035919x.2015.1088486 dippenaar-schoeman, a.s., lyle, r. & van den berg, a.m., 2012, ‘bioinformatics on the spiders of south africa’, serket 13, 121–127. dippenaar-schoeman, a.s. & van den berg, a.m., 2010, spiders of the kalahari, plant protection handbook no. 17, agricultural research council, pretoria. foord, s.h., dippenaar-schoeman, a.s. & haddad, c.r., 2011a, ‘south african spider diversity: african perspectives on the conservation of a mega-diverse group’, in o. grillo & g. venora (eds.), changing diversity in changing environment, pp. 163–182, in tech publishing, rijeka. foord, s.h., dippenaar-schoeman, a.s. & haddad, c.r., 2011b, ‘the faunistic diversity of spiders (arachnida, araneae) of the savanna biome in south africa’, transactions of the royal society of south africa 66, 170–201. https://doi.org/10.1080/0035919x.2011.639406 haddad, c.r. & dippenaar-schoeman, a.s., 2005, ‘epigeic spiders (arachnida: araneae) in nama karoo grassland in the northern cape province’, navorsinge van die nasionale museum, bloemfontein 21, 1–10. haddad, c.r. & dippenaar-schoeman, a.s., 2006, ‘epigeic spiders (araneae) in pistachio orchards in south africa’, african plant protection 12, 12–22. haddad, c.r. & dippenaar-schoeman, a.s., 2015, ‘diversity of non-acarine arachnids of the ophathe game reserve, south africa: testing a rapid sampling protocol’, koedoe 57, 1255. https://doi.org/10.4102/koedoe.v57i1.1255 haddad, c.r., dippenaar-schoeman, a.s. & pekár, s., 2005, ‘arboreal spiders (arachnida: araneae) in pistachio orchards in south africa’, african plant protection 11, 32–41. haddad, c.r., louw, s.vdm. & dippenaar-schoeman, a.s., 2004, ‘spiders (araneae) in ground covers of pistachio orchards in south africa’, african plant protection 10, 97–107. haddad, c.r., louw, s.vdm. & pekár, s., 2008, ‘commercial pistachio orchards maintain lower density and diversity of spiders (araneae): a study from south africa’, african plant protection 14, 24–36. low, a.b. & rebelo, a.g., 1998, vegetation of south africa, lesotho and swaziland, 2nd edn., department of environmental affairs and tourism, government printers, pretoria. lyle, r. & dippenaar-schoeman, a.s., 2013, ‘sampling in the diamond route reserves’, sansa newsletter 18, 10. lyle, r. & dippenaar-schoeman, a.s., 2015, ‘red listing of south african spiders’, sansa newsletter 23, 1. lyons, c.-l., 2009, ‘evaluating restoration success of alluvial diamond mined sites in south africa using invertebrate community indicators’, unpublished msc thesis, university of cape town. mcgeoch, m.a., sithole, h., samways, m.j., simaika, j.p., pryke, j.s., picker, m. et al., 2011, ‘conservation and monitoring of invertebrates in terrestrial protected areas’, koedoe 53, 1000. https://doi.org/10.4102/koedoe.v53i2.1000 petráková, l., líznarová, e., pekár, s., haddad, c.r., sentenská, l. & symondson, w.o.c., 2015, ‘discovery of a monophagous true predator, a specialist termite-eating spider (araneae: ammoxenidae)’, scientific reports 5, 14013. https://doi.org/0.1038/srep14013 van rooyen, n., 1999, the vegetation types and veld condition of tswalu private desert reserve, unpublished report to the management of tswalu kalahari reserve. vasconcellos-neto, j., romero, g.o., santos, a.j. & dippenaar-schoeman, a.s., 2007, ‘association of spiders of the genus peucetia (oxyopidae) with plants bearing glandular hairs’, biotropica 39, 221–226. https://doi.org/10.1111/j.1744-7429.2006.00250.x webb, p., 2013, ‘defense mechanism in burrow-dwelling wolf spiders’, sansa news 18, 4. appendix 1 table 1-a1: checklist of the spiders of tswalu kalahari reserve with guild, endemicity (en) and conservation status (cs). abstract introduction research methods and design results discussion conclusion acknowledgements references appendix 1 appendix 2 about the author(s) wataru tokura percy fitzpatrick institute of african ornithology, university of cape town, south africa department of biological sciences, university of cape town, south africa sam l. jack department of biological sciences, university of cape town, south africa tania anderson private, johannesburg, south africa michael t. hoffman department of biological sciences, university of cape town, south africa citation tokura, w., jack, s.l., anderson, t. & hoffman, m.t., 2018, ‘long-term variability in vegetation productivity in relation to rainfall, herbivory and fire in tswalu kalahari reserve’, koedoe 60(1), a1473. https://doi.org/10.4102/koedoe.v60i1.1473 note: this article is partly based on the thesis by wataru tokura, ‘understanding changes in plant productivity using evi satellite data in tswalu kalahari reserve’, available at https://open.uct.ac.za/handle/11427/20933 original research long-term variability in vegetation productivity in relation to rainfall, herbivory and fire in tswalu kalahari reserve wataru tokura, sam l. jack, tania anderson, michael t. hoffman received: 03 may 2017; accepted: 20 may 2018; published: 31 july 2018 copyright: © 2018. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract exploring the long-term influence of climate and land use on vegetation change allows for a more robust understanding of how vegetation is likely to respond in the future. to inform management, this study investigated the relationship between vegetation productivity trends and potential drivers of change in the 110 000 ha of the tswalu kalahari reserve between 2000 and 2015, using the moderate resolution imaging spectroradiometer enhanced vegetation index (evi, mod13q1). spatio-temporal variability of the evi was mapped and then related to the historical records of precipitation, animal numbers and fire occurrences. long-term trends in productivity were analysed by residual trend analysis (restrend). significantly different evi profiles were found between vegetation types, and this was related to the structure and function of the vegetation, as well as the effects of soil reflectance. the evi time-series signalled spatial and temporal heterogeneity in plant productivity, which was strongly correlated with rainfall, although fire and especially herbivory had noteworthy localised effects on productivity. the restrend identified a significant positive trend in plant productivity in shrub-dominated vegetation types, providing evidence for the ongoing thickening of woody species. significant negative trends in productivity were associated with artificial water points and more heavily stocked areas, leading to degradation. conservation implications: the southern kalahari has a highly variable rainfall regime, which is tied to a dynamic vegetation response. this variability should be taken into account when making management decisions. field-based monitoring together with adaptive management approaches are needed in the face of an uncertain future in which significant warming is expected. introduction vegetation dynamics in arid environments are controlled to a large degree by abiotic factors such as temperature and moisture availability (reynolds et al. 2004; weltzin et al. 2003). changes in moisture supply to vegetation, either directly via rainfall or indirectly in terms of the effects of rising temperatures on evaporation and transpiration rates, are therefore likely to have a significant effect on arid-adapted vegetation (collins et al. 2014; weltzin et al. 2003). consequently, there is an expectation that anthropogenic climate change will have a disproportionate impact on arid systems. indeed, recent declines in the equator-ward populations of certain arid region species, and a poleward shift in species’ ranges as a result of physiological stress and/or changing inter-specific interactions between species, have already been documented (cahill et al. 2012; chen et al. 2011; hickling et al. 2006). because of slow vegetation growth rates and episodic recruitment in arid environments, long-term monitoring studies have been crucial in revealing directional changes in these systems (guo 2004; lindenmayer et al. 2012; rahlao et al. 2008). although it can be difficult to discern the relative influence of different drivers on vegetation dynamics (weltzin et al. 2003), drawing on long-term and large-scale ecological data sets can provide a more accurate representation and robust understanding of complex systems and enable more evidence-based conservation management (lindenmayer et al. 2012). earth observation data (eod) derived from satellites have been used in a number of studies to monitor environmental change (nagendra et al. 2013). with its broad spatial coverage, growing historical archive, temporal consistency and cost efficiency, eod can circumvent the logistical challenges of field-based monitoring techniques (mishra et al. 2015b; nagendra et al. 2013). however, in the sparsely vegetated drylands context, the application of eod for vegetation studies has certain challenges, such as considerable background noise in the normalised difference vegetation index (ndvi) values (huete et al. 2002; kong et al. 2015; palmer & van rooyen 1998; van rooyen 2000), as well as in accounting for the overriding influence of rainfall over other factors (wessels et al. 2007). nevertheless, recent vegetation monitoring studies in the semi-arid kalahari in southern africa have successfully employed time-series ndvi or improved vegetation indices such as the enhanced vegetation index (evi), with careful interpretation aided by an understanding of the ecology and vegetation dynamics of the region (e.g. colditz et al. 2007; mishra et al. 2015a). several techniques, such as rain use efficiency and residual trend analysis (restrend) (wessels et al. 2007, 2012), have also been developed to isolate the effects of rainfall in order to discern the influence of other factors on dry land vegetation. in relation to other drylands of the world, the southern kalahari, located in the central-western region of southern africa, has experienced some of the strongest warming in the historical record (kruger & sekele 2013). moreover, modelled projections are for particularly severe climate change in this region in the 21st century, with reductions in moisture availability and higher evaporative loss as a result of marked temperature increases (dai 2013; engelbrecht et al. 2015; shongwe et al. 2009). several studies have reported on how historical warming may already be contributing to the reduced fitness of certain bird species (e.g. cunningham et al. 2013; du plessis et al. 2012). however, the impact of historical and projected climate change on vegetation change in the southern kalahari has received less attention, probably because of the difficulty of separating the influence of multiple interacting drivers of change that operate at different temporal and spatial scales. changes in the climate are happening against the backdrop of a shift in land use from extensive livestock production to private game reserves for tourism and hunting, as the latter becomes increasingly profitable (cloete et al. 2007; thomas & paul 1991). however, little is known about the long-term consequences of such a shift on vegetation. for example, changes in vegetation composition, structure and productivity on livestock farms in the southern kalahari have traditionally been ascribed to overstocking (rutherford & powrie 2010; skarpe 1990) and the presence of boreholes (palmer & van rooyen 1998; van rooyen & van rooyen 1998). the impact on the environment of free-roaming wild game, often kept at high densities and in the absence of predator species, has not been adequately assessed. both livestock production and game farming are heavily reliant on rainfall to ensure good forage. therefore, an increase in aridity is likely to affect the stocking rates of both industries and erode the profitability of these enterprises. this is a cause of concern because rangelands in the kalahari are known to be sensitive to overgrazing, with irreversible shifts from palatable perennial grasses to unpalatable woody shrubs, or to grasslands with a dominance of annual species (rutherford & powrie 2010; skarpe 1990; van rooyen 2000). additionally, an increase in atmospheric co2 concentration is also likely to facilitate the thickening of woody plants, which might lead to a shift in the suitability of vegetation for grazers to a suitability for browsers, as has been observed elsewhere in southern africa (bond & midgley 2012; o’connor, puttick & hoffman 2014). given the sensitivity of the southern kalahari to projected climate change and the complex ramifications of a shift in vegetation composition and structure on the natural environment and tourism industry, a robust understanding of the strengths of the relationships between arid-adapted vegetation and likely drivers of change, as well as rates of change, should be a priority. this study investigated the relationship between vegetation productivity trends and drivers of change over a 15-year period (2000–2015) in a 110 000 ha mega-reserve in the southern kalahari. this study was conducted in order to determine the dominant drivers of vegetation productivity, and thereby better understand vegetation responses to current and future climate change. earth observation data were used to (1) map plant communities; (2) analyse patterns of plant productivity in space and time; (3) relate changes to historical records for precipitation, animal numbers and fire; and (4) assess where unidirectional changes in productivity might represent the thickening of woody species or degradation. finally, we reflect on the implications of the findings of this study for natural resource management in similarly arid, summer rainfall environments. research methods and design study site tswalu kalahari reserve (tkr, 27.2031 s 22.4673 e, 1020 km2) is situated in the northern cape province of south africa (figure 1). the climate is typically hot and arid with highly variable rainfall occurring mainly during summer (december–march). the mean annual rainfall and standard deviation for the past 25 years is 361.4 mm ± 169.2 mm (south african weather service 2015). the landscape is characterised by sandy plains and parallel sand dunes, while the quartzitic korannaberg mountains extend from north to south through the eastern half of the reserve (davis et al. 2010). five different vegetation units are mapped by mucina and rutherford (2006) for the tkr, including koranna-langeberg mountain bushveld, gordonia duneveld, gordonia plains shrubveld, olifantshoek plains thornveld and kathu bushveld. figure 1: (a) location of tswalu kalahari reserve and major management units, and a comparison of the vegetation maps of the tswalu kalahari reserve adapted from (b) mucina & rutherford (2006) and (c) a landsat-based vegetation map prepared during the course of this study. prior to 1995 the tkr was divided into more than 40 livestock farms, but was converted into a nature reserve by removing farm infrastructure (e.g. fences and livestock handling pens), closing several artificial water points and restocking with wildlife species (davis et al. 2010). the tkr has undergone continual expansion in the last 20 years and the current size of the reserve exceeds 110 000 ha. currently, 75 mammal species are present, most of which are grazing herbivores (e.g. gemsbok [oryx gazelle], springbok [antidorcas marsupialis] and blue wildebeest [connochaetes taurinus]), with a lower number of browser species (e.g. greater kudu [tragelaphus strepsiceros], giraffe [giraffa camelopardalis] and black rhinoceros [diceros bicornis]), omnivores and predators. the reserve is divided into three fenced areas, each with its own management plan (figure 1). these are (1) the ‘predator camp’ in the north-east where lions [panthera leo] are present, (2) the roan [hippotragus equinus] and sable [hippotragus niger] breeding camp in the north-west corner of the reserve, fenced off into ten sections and (3) the remainder of the reserve which supports the majority of the herbivores. data preprocessing vegetation types to derive a vegetation map of the tkr, the cloud-free, geometrically corrected, multispectral landsat 8 operational land imager (oli) data (captured on 02 april 2014, scene id lc81740792014092lgn00), band b2–b7 (blue, green, red, near infrared, swir1 and swir2) were used to perform supervised classification. the bands of images covered the entire reserve and were taken near the peak of the growth season in an above-average rainfall year. the obtained image was converted from a digital number to a top of atmosphere radiance and then corrected for the sun’s angle during preprocessing before the analysis, using geosud toa reflectance plugin (ver 1.0) in qgis (ver 2.10 pisa, qgis development team). preprocessed landsat oli data were used for the supervised classification with a random forest algorithm. a comprehensive set of photographs taken in may 2015 of the main vegetation units in the tkr was used to produce a training data set. this was augmented by high-resolution satellite images from google earth (ver 7.1.5.1557, google inc.) in combination with vegetation classes of the national vegetation map produced by mucina and rutherford (2006). based on this approach, a total of 53 sampling areas were generated and 200 pixels were randomly selected per vegetation class. the 200 pixels were then split into 100 training pixels and 100 validation pixels. subsequently, a supervised classification was performed using the random forest algorithm package ‘rasclass’ (ver 0.2.2) in r (ver 3.1.0, r core team 2013). the routines of post-processing were performed using qgis (ver 2.10) and included filtering isolated pixels or noise. accuracy was assessed by comparing the validation pixels against those in the produced map. the final product comprising a landsat-based vegetation map was amalgamated into a single image and used in further analyses. vegetation productivity satellite images, which encompassed the reserve for the period february 2000 to november 2015, were obtained. the evi product at 250 m resolution derived from the moderate resolution imaging spectroradiometer (modis, mod13q1 ver 005) was used as a proxy for plant productivity. this approach has been used previously for the monitoring of plant productivity in the kalahari (colditz et al. 2007; hüttich et al. 2009). the modis evi product has several advantages such as a reduction in atmospheric noise and a reduction in the variation in canopy background signals (huete et al. 2002), as well as a decoupling of the influence of the variation in soil brightness (solano et al. 2010). the seasonality parameters of the evi were extracted from time-series data using the software package timesat (jönsson & eklundh 2004). this package can quantify noise-corrupted remote sensing time-series data by filtering each pixel for noise and identifying (1) phenological measurements such as the beginning and end of the plant growth season and (2) plant productivity, represented by the small integrated value of evi (siv of evi) (jönsson & eklundh 2004; wessels et al. 2011). the adaptive savitzky–golay filter, with a window width of four data points, was applied to smooth the data. the season per year was set at one because vegetation in the kalahari has one growth season per year. the start and end of the growth seasons were defined as a 20% increase in the seasonal amplitude, measured from the left and right minimum levels to the maximum of the seasonal curve. these values were determined not only by visually inspecting the fitted curve on the timesat graphical user interface but also by referring to an earlier study from the region (wessels et al. 2011). to illustrate spatio-temporal patterns of vegetation productivity, the siv of evi, quantified by timesat, was mapped onto each growth season. the siv of evi provides a good estimate of the production of the seasonally dominant vegetation type (fensholt et al. 2013; jönsson & eklundh 2004) for each pixel. additionally, smoothed time-series data of evi were extracted to understand the temporal variation in plant productivity. rainfall daily rainfall data collected from 30 rain gauges located within the tkr for the period 2001–2014 were used in the analysis. in addition, monthly rainfall records from five neighbouring rainfall stations (van zylsrus, kathu, severn, wildebeesduin and upington) were obtained from the south african weather service (2015). a spatially continuous rainfall surface was generated by interpolating the rainfall records from the 35 stations distributed throughout and around the tkr. firstly, the original data from rain gauges and weather stations were formatted by removing incomplete and suspect records. next, total rainfall for the growth season, which starts at the beginning of october and ends at the end of september in the savanna biome of south africa (wessels et al. 2011), was calculated as the annual rainfall for each rainfall station. to produce annual rainfall surfaces, ordinary kriging was performed because it is commonly used and appears to be preferable (goovaerts 2000; ly, charles & degré 2013). a spherical model was selected to fit a semi-variogram, while other parameters were set at default values. the annual rainfall surfaces were produced using the geostatistical analyst tool in arcmap (ver 10.0, esri) for the 2001–2002 to 2013–2014 growth seasons at 250 m spatial resolution, which matched precisely to the modis raster image. herbivores changes in herbivore pressure were estimated using large animal units (laus), which is a standard metric for calculating commercial stocking densities (van rooyen 2010). the laus were determined from annual aerial count data provided by the tkr for the period 2005–2016 for the predator camp and the remainder of the tkr (excluding the roan and sable breeding camp). fire the modis burned-area product (mcd45a1), which denotes monthly fire occurrence at 500 m resolution, was used to examine the effect of fire on plant productivity in the tkr. these data, from the beginning of the 2000–2001 growth season to the end of the 2014–2015 growth season, were downloaded. all the satellite data were obtained for the tkr from the national aeronautics and space administration’s earth observing system clearing house, reverb (http://reverb.echo.nasa.gov/reverb/). data analysis relationship between vegetation productivity and potential drivers the influence of the potential drivers (rainfall and herbivores) on vegetation productivity in the different vegetation types was analysed as a response variable in a linear mixed-effects model using the siv of evi as the explanatory variable. the interpolated annual rainfall surface and laus were used for this analysis to represent the effects of rainfall and herbivores. vegetation types were assigned by upscaling the 30 m resolution landsat-based vegetation map to a 250 m grid, which matched precisely to the modis data. fire was not included in this model because of a limited number of fire occurrences and its contrasting effect on the siv of evi. as a result of the data availability, only the data between 2005–2006 and 2013–2014 in the predator camp and the remainder of the tkr (excluding the roan and sable breeding camp) were pooled into one data set. however, to avoid potential disproportional bias, the data from the 2006–2007 growth season were excluded as the tkr experienced a severe drought with little rainfall over this period. the growth season and id of pixels were added as random terms to account for pseudoreplication. statistical analyses were done using the lmer function in ‘lme4’ and ‘lmetest’ package in r (ver 3.1.0, r core team 2013). the effect of fire was visually assessed by comparing maps of the siv of evi and fire occurrence. degradation trend analysis (residual trend analysis) residual trend (restrend) was used to analyse each growth season using each growth season (october–september) as a time step for the duration of 13 growth seasons (2001–2002 to 2014–2015). restrend removes the effect of rainfall from the long-term trend in productivity, and in so doing is able to highlight areas where land degradation has occurred (wessels et al. 2007). this analysis followed the method proposed by wessels et al. (2007), except that rainfall was not log transformed because plant productivity was unlikely to level off in the water-limited environment of the kalahari. firstly, the regression analysis of the siv of evi (response variable) and annual rainfall (explanatory variable) was performed for each pixel; then the coefficient of determination (r2) and p-values were mapped. following this, trends in the residuals, expressed as the difference between observed evi and predicted evi by rainfall, were regressed through the growth seasons (wessels et al. 2007, 2012). the long-term trend represented by the slope of the linear regression of the trends in the residuals was mapped to determine the distribution pattern of degraded areas, defined as areas experiencing the degeneration of structure or function. both regression analyses were performed per pixel, using a linear model function ‘lm’ in r (ver 3.1.0, r core team 2013). data from the 2006–2007 growth season (october–september), when the tkr experienced a severe drought, were excluded from this analysis to avoid a potential disproportional bias affecting the underlying regression model. to further assess the progress of degradation, standardised restrend values were computed over time in several areas in the tkr where significant decreasing or increasing trends were detected. results vegetation types five vegetation types were classified for the tkr, with the landsat-based map producing greater spatial detail than that achieved by mucina and rutherford (2006) (figure 1). field observations found a general correspondence between topography, plant community structure and the landsat-based vegetation types. for example, vegetation classified as koranna-langeberg mountain bushveld vegetation was composed predominantly of a mixture of trees and shrubs, gordonia duneveld was dominated by grasses, gordonia plains shrubveld by dwarf shrubs and olifantshoek plains thornveld by short (i.e. < 3 m high) woody shrubs such as senegalia mellifera. in the landsat-based vegetation map, there were a few exceptions such as around old artificial water points, which were often misclassified as gordonia plains shrubland. certain parts in the west of the tkr were also incorrectly designated as being part of the koranna-langeberg mountain bushveld. boundaries of the landsat-based vegetation types were also sometimes misclassified as a result of a similarity in vegetation structure and a mixture of species at the margins. this was particularly evident for gordonia duneveld and gordonia plains shrubveld, and kathu bushveld and olifantshoek plains thornveld, respectively. notwithstanding the above discrepancies, overall classification accuracy was 86.2%, while the kappa statistic was 0.83 (see appendix 1). spatio-temporal patterns and variability of vegetation productivity the overall geographical trend of the mean of the siv of evi was lower in the west and higher in the east, often delineated by the north–south axis of the korannaberg mountains (figure 2). differences in the spatial patterning of evi values were assumed to relate to differences in physical vegetation attributes such as dominant growth forms, species composition and soil types. extremely low evi values corresponded to bare dune crests in the west and provincial roads in the east of the tkr, whereas abrupt changes in the siv of evi were observed at vegetation type boundaries (figure 2). figure 2: a composite image of the annual small integrated value of the enhanced vegetation index in the tswalu kalahari reserve for the 2000–2001 to 2014–2015 growth season, which starts at the beginning of october and ends at the end of september. red colours indicate a lower enhanced vegetation index value corresponding to lower plant productivity. (a) 2000–2001, (b) 2001–2002, (c) 2002–2003, (d) 2003–2004, (e) 2004–2005, (f) 2005–2006, (g) 2006–2007, (h) 2007–2008, (i) 2008–2009, (j) 2009–2010, (k) 2010–2011, (l) 2011–2012, (m) 2012–2013, (n) 2013–2014 and (o) 2014–2015. figure 2 (continues...): a composite image of the annual small integrated value of the enhanced vegetation index in the tswalu kalahari reserve for the 2000–2001 to 2014–2015 growth season, which starts at the beginning of october and ends at the end of september. red colours indicate a lower enhanced vegetation index value corresponding to lower plant productivity. (a) 2000–2001, (b) 2001–2002, (c) 2002–2003, (d) 2003–2004, (e) 2004–2005, (f) 2005–2006, (g) 2006–2007, (h) 2007–2008, (i) 2008–2009, (j) 2009–2010, (k) 2010–2011, (l) 2011–2012, (m) 2012–2013, (n) 2013–2014 and (o) 2014–2015. the time-series analysis of evi values demonstrated a seasonal cycle in plant productivity, with higher values in the summer and lower values in the winter. on average, the beginning of the growth season started in the period from december to january and ended in august to september (figure 3). the time-series of evi for the past 16 years for the tkr illustrated high inter-annual variability in plant productivity (figures 2 and 3). low evi values were evident especially towards the east of the tkr, during the 2002–2003, 2003–2004, 2006–2007 and 2012–2013 growth seasons, which corresponded to periods of low annual rainfall (figure 4). in the 2006–2007 growth season, when the annual rainfall was lowest for the study period, timesat failed to quantify the productivity metrics for some pixels because of a low and indistinct peak in evi values. figure 3: average of smoothed time-series of the enhanced vegetation index using an adaptive savitzky–golay filter of timesat for the different vegetation types in the tswalu kalahari reserve. figure 4: a composite of interpolated annual rainfall surface contour maps computed by ordinary kriging for the growth seasons (october to september) in the tswalu kalahari reserve. black dots in the 2001–2002 map represent the locations of the rainfall stations. (a) 2001–2002, (b) 2002–2003, (c) 2003–2004, (d) 2004–2005, (e) 2005–2006, (f) 2006–2007, (g) 2007–2008, (h) 2008–2009, (i) 2009–2010, (j) 2010–2011, (k) 2011–2012, (l) 2012–2013 and (m) 2013–2014. figure 4 (continues...): a composite of interpolated annual rainfall surface contour maps computed by ordinary kriging for the growth seasons (october to september) in the tswalu kalahari reserve. black dots in the 2001–2002 map represent the locations of the rainfall stations. (a) 2001–2002, (b) 2002–2003, (c) 2003–2004, (d) 2004–2005, (e) 2005–2006, (f) 2006–2007, (g) 2007–2008, (h) 2008–2009, (i) 2009–2010, (j) 2010–2011, (k) 2011–2012, (l) 2012–2013 and (m) 2013–2014. relationship between vegetation productivity and potential drivers rainfall has been highly variable over time and space in the tkr over the past 15 years. however, a general spatial gradient was observed, with mean annual rainfall being higher in the east and lower in the west (figure 4). as a result of the orographic effects of altitude, the korannaberg mountains probably receive higher rainfall than other areas in the tkr, although we could not confirm this because of a lack of rain gauges on the slopes and peaks of these mountains. standard error of the rainfall surface within most of the areas in the tkr was less than 60 mm. the values varied depending on the growth season and location, and higher standard errors were estimated in the south-eastern corner of the reserve where the rainfall stations were relatively scarce (see appendix 2). in general, the number of herbivores within the main section of the tkr increased between 2005 and 2013 and thereafter remained relatively stable, although there was a slight decline in 2016 (figure 5). the recorded decline between 2008 and 2010 was because of the capture and removal of extra-limital species by management and an unexplained decline in springbok numbers over this period. herbivore numbers within the predator camp have remained stable throughout the recorded period. figure 5: large animal units for two management areas within the tswalu kalahari reserve for the period 2005–2016. large animal units were estimated from annual aerial count data and large animal units values for the herbivores, after van rooyen (2010). three fire events affecting the reserve were evident from the modis burned-area product data. however, these fires were restricted to the korannaberg mountains in the south-eastern corner of the reserve, and most of the reserve had not burned during the study period (figure 6). the impact of these fires on plant productivity was captured by the siv of evi, although the response was case specific. for example, the area south of the staff village, which was exposed to fire in january 2013, had a very low siv of evi value in 2012–2013, while the area that was burned in august to september 2012 had an even more muted response (figure 2). conversely, the area that burned outside the tkr in 2010 showed an increase in the siv of evi in the same season. figure 6: the frequency of fire from 2000–2001 to 2014–2015 growth seasons as determined from the moderate resolution imaging spectroradiometer burned-area products (mcd45a1) in the tswalu kalahari reserve. the data show that in the past 15 years the burned areas have been confined to the mountains in the south-eastern corner of the reserve. the dates on the map indicate the year and month in which the fires occurred. results of the analysis using a linear mixed-effects model indicated that there is a significant positive effect of rainfall on the siv of evi (p < 0.001) (table 1). the model also suggested different levels of effects from vegetation types (p < 0.001) and a relatively minor but significantly negative influence of laus (p < 0.001). table 1: results of a linear mixed-effects model that explores the effect of rainfall, large animal units and vegetation type (explanatory variable) on vegetation productivity represented by the small integrated value of enhanced vegetation index (responsive variable) in the tswalu kalahari reserve. the year and id of the pixel are added as random terms in the model. degradation trend analysis (residual trend analysis) regression analysis between the siv of evi and annual rainfall indicated a significant positive correlation for most of the tkr (p < 0.05), except in the roan and sable breeding camp, as well as for parts of the kathu bushveld and olifantshoek plains thornveld vegetation types (figure 7). the trend in plant productivity computed by the restrend detected a significant positive trend in plant productivity in the east and south-west of the tkr, while a negative trend was detected in some locations in the centre and west of the reserve (figure 8). the standardised restrend values from these areas demonstrated a relatively consistent and directional change over time (figure 9). this suggested that the observed change was initiated before the 2000–2001 growth season and that the driver of change has remained the same. most of the area that showed an increasing trend overlapped with shrub-dominated vegetation, especially the olifantshoek plains thornveld in the east and gordonia plains shrubveld in the south-west. conversely, declines in productivity were observed for small areas of gordonia plains shrubveld in the central and western area. figure 7: map of the correlation of determination (r2) of the regression analysis of the annual rainfall for the growth seasons (explanatory variable) and small integrated value of the enhanced vegetation index (responsive variable) between the 2001–2002 and 2013–2014 growth seasons, excluding the drought season of 2006–2007. figure 8: map of the slope of the residual-year regression analysis of the residual trend analysis. the areas where a significant trend was not found are displayed in grey. the residual trend analysis showed a significant positive long-term trend in vegetation productivity in the eastern and south-western parts of the tswalu kalahari reserve. figure 9: examples of significant trends in standardised residual trend analysis values over time from several areas with significant decreasing trends (graphs a and b) or increasing trends (graphs c–g) in the tswalu kalahari reserve. values are shown as mean (± standard deviation) for each area. (a) east of rogella pan, (b) kalkpan, (c) south-west, (d) tarkuni, (e) south-east, (f) dedeben and (g) langkloof/bruwer. (h) the small map indicates the general area covered by each graph. figure 9 (continues...): examples of significant trends in standardised residual trend analysis values over time from several areas with significant decreasing trends (graphs a and b) or increasing trends (graphs c–g) in the tswalu kalahari reserve; (h) the small map indicates the general area covered by each graph. values are shown as mean (± standard deviation) for each area. discussion spatio-temporal patterns and variability of vegetation productivity variations in evi values illustrated the spatio-temporal patterns of vegetation productivity through variable changes in vegetation cover. these included significant increases in annual grass cover in the wet season or conversely, dieback in the dry season (van rooyen et al. 1984). the evi values typically increased with plant greening in late spring and summer, decreased in the late autumn and then remained low during the dry winter months. this cycle matched the findings of previous remote sensing studies in the kalahari (hüttich et al. 2009; jolly & running 2004; mishra et al. 2015a; wessels et al. 2011), as well as field observations (sekhwela & yates 2007), and confirmed the utility of evi as a proxy for seasonal vegetation productivity cycles in this semi-arid region. it also highlighted the potential utility of this approach to aid wide-scale and real-time decision-making in terms of, for example, the determination of optimal stocking rates at a specific time, given the spatial assessment of rainfall in the preceding season. relationship between vegetation productivity and potential drivers not surprisingly, rainfall was found to be the most important factor in determining the spatial success or failure of interand intra-seasonal, as well as inter-annual, productivity (masunga, moe & pelekekae 2013; van rooyen et al. 1990; van rooyen & van rooyen 1998). the effect of other factors such as herbivory was secondary to the influence of rainfall. different vegetation types had significantly different evi phenological profiles, which suggests that plant structural and functional traits have an important influence on plant productivity. for example, woody plants, which increase in abundance from west to east in the tkr, generally demonstrated higher and more persistent greenness (mishra et al. 2015a, 2015b) compared to the grass and low shrub-dominated vegetation of gordonia duneveld and gordonia plains shrubveld. this increased productivity is mediated to a large degree by increasing rainfall and deeper rooting depths typically found in vegetation associated with koranna-langeberg mountain bushveld, olifantshoek plains thornveld and kathu bushveld. the ubiquity of thorny species and the density of the vegetation also contributed to a reduction in browser pressure and therefore the persistence of greenness recorded in these vegetation types. notwithstanding the overwhelming influence of rainfall, the results suggested that herbivores negatively influence vegetation productivity. a decoupling between growth season rainfall and plant productivity was apparent, especially within the ten roan and sable breeding camps, which cover an area of 1250 ha in the north-western corner of the tkr. this area has a different management strategy to the rest of the reserve. it is characterised by high densities of economically valuable grazing herbivores and the provision of supplemental feed and water. the patterns observed within these camps point to a possible threshold stocking rate beyond which herbivory and associated impacts such as trampling and very low vegetation cover in general (which were clearly observable in the field) weakened the correlation between rainfall and evi, and resulted in herbivore-driven degradation. fire has only occurred in limited areas and times in the tkr, and its effect on vegetation productivity has been localised. the observed response of fire in evi was case specific. the siv of evi has decreased nearly to zero after the fire in the tkr in the 2012–2013 growth season, while the increase in the siv of evi, as seen in the area burned outside the tkr in 2010, may have been caused by the post-fire recovery processes. from these observations, it is assumed that the intensity and date of the fire influence several key characteristics of plant productivity in the growth season immediately after the fire. degradation and management implications residual trends provided a useful analytical approach in the southern kalahari where the strong influence of erratic rainfall on annual productivity tends to overwhelm the contribution of other factors. results of the restrend analysis suggested that the thickening of woody species has occurred in the eastern and south-western parts of the tkr. although the process of the thickening of woody species has not been monitored, the direction and slope of trends implies that this process has progressed constantly over the course of the study period. interestingly, certain lowland areas located within the predator camp in the eastern part of the tkr showed a poor correlation between rainfall and evi and might also have experienced a thickening of woody plants, as the poor correlation indicates potential effects on vegetation productivity from factors other than rainfall. we speculate that a combination of lower herbivore stocking rates in the predator camp as a result of the presence of lions allows other factors such as low fire frequency, the densification of thorny vegetation and possibly the effects of high atmospheric co2 concentrations (which favour woody species) to promote more rapid woody thickening in these areas (o’connor et al. 2014). these factors act together to establish a positive feedback loop, which promotes further densification of woody species with a concomitant decrease in more flammable grass cover and the eventual ‘switching off’ of fire as a means of ‘resetting’ the system. earlier studies have noted that the thickening of woody species can have multiple possible drivers (see o’connor et al. [2014] for an overview). establishing the relative contribution of each of the potential drivers is often difficult because they interact and vary in space and time. however, irrespective of the cause for the shift from grassto shrub-dominated vegetation, this change has important implications for several aspects of reserve management, including the ratio of grazers to browsers and associated stocking rates and the frequency of controlled burns. it is therefore likely that a more active management regime will be required in order to maintain the degree of openness in encroached landscapes and the associated ecology in terms of, for example, grazer–browser ratios, as well as the game-viewing experience available to visitors to the reserve. in the drier west of the tkr, restrend identified a significant decreasing trend in vegetation productivity in some areas of gordonia plains shrubveld and on dune slopes in gordonia duneveld. standardised restrend values exhibited a continuously declining trend, which suggested that a persistent driver might be influencing this pattern. because these areas have not been burned since 2000 and restrend analysis is able to exclude the effect of rainfall on vegetation productivity, the decline in productivity has likely been caused by the impact of herbivores. from our field observations, localised heavy grazing appears to have reduced vegetation cover in these areas, especially at sites close to artificial water points. another possibility is that the difference in underlying soil types might influence the palatability and thus the grazing pressure from herbivores. however, validation or simulation of the data to test its robustness is recommended because restrend is statistically underpowered when shorter periods of time are analysed (e.g. less than 16 growth seasons) and is strongly influenced by the timing of degradation (wessels et al. 2012). nevertheless, it would be prudent for management to adopt a more conservative herbivore stocking rate or allow predator control of herbivore numbers in affected areas. this suggestion is underscored by unfavourable climate projections (dai 2013; engelbrecht et al. 2015; shongwe et al. 2009), which are likely to negatively impact vegetation, and the known sensitivity of kalahari rangelands to overgrazing (jeltsch et al. 1997; rutherford & powrie 2010; skarpe 1990; van rooyen 2000). these threats combine to pose a growing risk to natural resource managers in the southern kalahari. given the current uncertainty in especially rainfall projections and inherent fine-scale spatial heterogeneity in vegetation, long-term field-based monitoring is also recommended in order to establish a more detailed understanding of the rate and magnitude of change within different vegetation types and across the rainfall gradient. conclusion variability in the spatial and temporal patterns of vegetation productivity, as well as long-term changes in this measure, were outlined for a large reserve in the southern kalahari using modis evi. the findings of this study confirmed known vegetation dynamics in the region, such as high spatial heterogeneity, seasonal change and extreme inter-annual variability of plant productivity shaped largely by erratic rainfall. through restrend it was possible to isolate the effects of rainfall and provide evidence of potential overgrazing and the thickening of woody species in certain areas of the tkr. the thickening of woody species is likely to intensify and spread in future because of the effects of co2 fertilisation, access by trees and shrubs to deeper and more consistent water supplies under more arid conditions and the establishment of feedback loops which could switch off grass-mediated fires. additionally, degradation trends in the west hint at possibly high and unsustainable stocking rates, which, if left unchecked, may erode the carrying capacity in this area of the tkr and take many years to recover. these findings emphasise the need for a proactive and anticipatory management style, informed by an extensive climate, vegetation and animal monitoring programme. this combination is necessary in order to respond timeously and effectively to both current and future risks facing the southern kalahari environment. acknowledgements we thank julian smit, christiaan j. harmse, claire davis and feroza morris for their valuable technical advice and bonginkosi prince ngomane, armin du plessis and kelsey green for fieldwork assistance. konrad wessels and anthony r. palmer provided comments to improve the manuscript. parts of the rainfall record used in the analysis were provided by the south african weather service. we would also like to thank dylan smith and the tswalu kalahari reserve for providing data, and duncan macfadyen and e oppenheimer & son for enabling this research. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions w.t. and s.l.j. conducted the fieldwork. t.a. and m.t.h. conceptualised and designed the overall research. w.t. performed the data analysis and led the writing. all authors participated in the planning of the research, the interpretation of the findings and the critical revision of the manuscript. references bond, w.j. & midgley, g.f., 2012, ‘carbon dioxide and the uneasy interactions of trees and savannah grasses’, philosophical transactions of the royal society b, 367(1588), 601–612. https://doi.org/10.1098/rstb.2011.0182 cahill, a.e., aiello-lammens, m.e., fisher-reid, m.c., hua, x., karanewsky, c.j., ryu, h.y. et al., 2012, ‘how does climate change cause extinction?’, proceedings of the royal society b, 280(1750), 20121890. https://doi.org/10.1098/rspb.2012.1890 chen, i., hill, j.k., ohlemüller, r., roy, d.b. & thomas, c.d., 2011, ‘rapid range shifts of species associated with high levels of climate warming’, science 333(6045), 1024–1026. https://doi.org/10.1126/science.1206432 cloete, p.c., taljaard, p.r. & grové, b., 2007, ‘a comparative economic case study of switching from cattle farming to game ranching in the northern cape province’, south african journal of wildlife research 37(1), 71–78. https://doi.org/10.3957/0379-4369-37.1.71 colditz, r.r., gessner, u., conrad, c., van zyl, d., malherbe, j., newby, t. et al., 2007, ‘dynamics of modis time series for ecological applications in southern africa’, proceedings of the 2007 international workshop on the analysis of multi-temporal remote sensing images, pp. 1–6, leuven, belgium, 18–20th july. https://doi.org/10.1109/multitemp.2007.4293069 collins, s.l., belnap, j., grimm, n.b., rudgers, j.a., dahm, c.n., d’odorico, p. et al., 2014, ‘a multiscale, hierarchical model of pulse dynamics in arid-land ecosystems’, annual review of ecology, evolution, and systematics 45, 397–419. https://doi.org/10.1146/annurev-ecolsys-120213-091650 cunningham, s.j., martin, r.o., hojem, c.l. & hockey, p.a.r., 2013, ‘temperatures in excess of critical thresholds threaten nestling growth and survival in a rapidly-warming arid savanna: a study of common fiscals’, plos one 8(9), e74613. https://doi.org/10.1371/journal.pone.0074613 dai, a.g., 2013, ‘increasing drought under global warming in observations and models’, nature climate change 3, 52–58. https://doi.org/10.1038/nclimate1633 davis, a.l.v., scholtz, c.h., kryger, u., deschodt, c.m. & strümpher, w.p., 2010, ‘dung beetle assemblage structure in tswalu kalahari reserve: responses to a mosaic of landscape types, vegetation communities, and dung types’, environmental entomology 39(3), 811–820. https://doi.org/10.1603/en09256 du plessis, k.l., martin, r.o., hockey, p.a.r., cunningham, s.j. & ridley, a.r., 2012, ‘the costs of keeping cool in a warming world: implications of high temperatures for foraging, thermoregulation and body condition of an arid-zone bird’, global change biology 18, 3063–3070. https://doi.org/10.1111/j.1365-2486.2012.02778.x engelbrecht, f., adegoke, j., boape, m., naidoo, m., garland, r., thatcher, m. et al., 2015, ‘projections of rapidly rising surface temperatures over africa under low mitigation’, environmental research letters 10, 085004. https://doi.org/10.1088/1748-9326/10/8/085004 fensholt, r., rasmussen, k., kaspersen, p., huber, s., horion, s. & swinnen, e., 2013, ‘assessing land degradation/recovery in the african sahel from long-term earth observation based primary productivity and precipitation relationships’, remote sensing 5(2), 664–686. https://doi.org/10.3390/rs5020664 goovaerts, p., 2000, ‘geostatistical approaches for incorporating elevation into the spatial interpolation of rainfall’, journal of hydrology 228(1–2), 113–129. https://doi.org/10.1016/s0022-1694(00)00144-x guo, q., 2004, ‘slow recovery in desert perennial vegetation following prolonged human disturbance’, journal of vegetation science 15(6), 757–762. https://doi.org/10.1658/1100-9233(2004)015[0757:sridpv]2.0.co;2 hickling, r., roy, d.b., hill, j.k., fox, r. & thomas, c.d., 2006, ‘the distributions of a wide range of taxonomic groups are expanding polewards’, global change biology 12, 450–455. https://doi.org/10.1111/j.1365-2486.2006.01116.x huete, a.r., didan, k., miura, t., rodriguez, e.p., gao, x. & ferreira, l.g., 2002, ‘overview of the radiometric and biophysical performance of the modis vegetation indices’, remote sensing of environment 83, 195–213. https://doi.org/10.1016/s0034-4257(02)00096-2 hüttich, c., gessner, u., herold, m., strohbach, b.j., schmidt, m., keil, m. et al., 2009, ‘on the suitability of modis time series metrics to map vegetation types in dry savanna ecosystems: a case study in the kalahari of ne namibia’, remote sensing 1(4), 620–643. https://doi.org/10.3390/rs1040620 jeltsch, f., milton, s.j., dean, w.r.j. & van rooyen, n., 1997, ‘simulated pattern formation around artificial waterholes in the semi-arid kalahari’, journal of vegetation science 8, 177–188. https://doi.org/188.10.2307/3237346 jolly, w.m. & running, s.w., 2004, ‘effects of precipitation and soil water potential on drought deciduous phenology in the kalahari’, global change biology 10, 303–308. https://doi.org/10.1046/j.1365-2486.2003.00701.x jönsson, p. & eklundh, l., 2004, ‘timesat – a program for analyzing time-series of satellite sensor data’, computers and geosciences 30(8), 833–845. https://doi.org/10.1016/j.cageo.2004.05.006 kong, t.m., marsh, s.e., van rooyen, a.f., kellner, k. & orr, b.j., 2015, ‘assessing rangeland condition in the kalahari duneveld through local ecological knowledge of livestock farmers and remotely sensed data’, journal of arid environments, 113, 77–86. https://doi.org/10.1016/j.jaridenv.2014.10.003 kruger, a.c. & sekele, s.s., 2013, ‘trends in extreme temperature indices in south africa: 1962–2009’, international journal of climatology 33(3), 661–676. https://doi.org/10.1002/joc.3455 lindenmayer, d.b., likens, g.e., andersen, a., bowman, d., bull, c.m., burns, e. et al., 2012, ‘value of long-term ecological studies’, austral ecology 37(7), 745–757. https://doi.org/10.1111/j.1442-9993.2011.02351.x ly, s., charles, c. & degré, a., 2013, ‘different methods for spatial interpolation of rainfall data for operational hydrology and hydrological modeling at watershed scale: a review’, biotechnology, agronomy, society and environment 17(2), 392–406, viewed 21 december 2015, from http://hdl.handle.net/2268/136084 masunga, g.s., moe, s.r. & pelekekae, b., 2013, ‘fire and grazing change herbaceous species composition and reduce beta diversity in the kalahari sand system’, ecosystems 16(2), 252–268. https://doi.org/10.1007/s10021-012-9611-6 mishra, n.b., crews, k.a., miller, j.a. & meyer, t., 2015a, ‘mapping vegetation morphology types in southern africa savanna using modis time-series metrics: a case study of central kalahari, botswana’, land 4(1), 197–215. https://doi.org/10.3390/land4010197 mishra, n.b., crews, k.a., neeti, n., meyer, t. & young, k.r., 2015b, ‘modis derived vegetation greenness trends in african savanna: deconstructing and localizing the role of changing moisture availability, fire regime and anthropogenic impact’, remote sensing of environment 169, 192–204. https://doi.org/10.1016/j.rse.2015.08.008 mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, south african national biodiversity institute, pretoria. nagendra, h., lucas, r., honrado, j.p., jongman, r.h.g., tarantino, c., adamo, m. et al., 2013, ‘remote sensing for conservation monitoring: assessing protected areas, habitat extent, habitat condition, species diversity, and threats’, ecological indicators 33, 45–59. https://doi.org/10.1016/j.ecolind.2012.09.014 o’connor, t.g., puttick, j.r. & hoffman, m.t., 2014, ‘bush encroachment in southern africa: changes and causes’, african journal of range and forage science 31(2), 67–88. https://doi.org/10.2989/10220119.2014.939996 palmer, a.r. & van rooyen, a.f., 1998, ‘detecting vegetation change in the southern kalahari using landsat tm data’, journal of arid environments 39(2), 143–153. https://doi.org/10.1006/jare.1998.0399 rahlao, s.j., hoffman, m.t., todd, s.w. & mcgrath, k., 2008, ‘long-term vegetation change in the succulent karoo, south africa following 67 years of rest from grazing’, journal of arid environments 72(5), 808–819. https://doi.org/10.1016/j.jaridenv.2007.08.003 r core team, 2013, r: a language and environment for statistical computing, r foundation for statistical computing, vienna. reynolds, j.f., kemp, p.k., ogle, k. & fernández, r.j., 2004, ‘modifying the “pulse–reserve” paradigm for deserts of north america: precipitation pulses, soil water, and plant responses’, oecologia 141(2), 194–210. https://doi.org/10.1007/s00442-004-1524-4 rutherford, m.c. & powrie, l.w., 2010, ‘severely degraded dunes of the southern kalahari: local extinction, persistence and natural re-establishment of plants’, african journal of ecology 48(4), 930–938. https://doi.org/10.1111/j.1365-2028.2009.01194.x sekhwela, m.b.m. & yates, d.j., 2007, ‘a phenological study of dominant acacia tree species in areas with different rainfall regimes in the kalahari of botswana’, journal of arid environments 70(1), 1–17. https://doi.org/10.1016/j.jaridenv.2006.12.006 shongwe, m.e., van oldenborgh, g.j., van den hurk, b.j.j.m., de boer, b., coelho, c.a.s. & van aalst, m.k., 2009, ‘projected changes in mean and extreme precipitation in africa under global warming. part i: southern africa’, journal of climate 22(13), 3819–3837. https://doi.org/10.1175/2009jcli2317.1 skarpe, c., 1990, ‘shrub layer dynamics under different herbivore densities in an arid savanna, botswana’, journal of applied ecology 27(3), 873–885. https://doi.org/10.2307/2404383 solano, r., didan, k., jacobson, a. & huete, a., 2010, modis vegetation index user’s guide (mod13 series), viewed 09 november 2015, from http://vip.arizona.edu/documents/modis/modis_vi_usersguide_01_2012 south african weather service, 2015, monthly rainfall data (1990–2015), pretoria. thomas, d.s.g. & paul, a.s., 1991, the kalahari environment, cambridge university press, cambridge. van rooyen, a.f., 2000, ‘rangeland degradation in the southern kalahari’, phd thesis, department of range and forage science, university of natal, pietersmaritzburg. van rooyen, n., 2010, ‘veld management in the african savannas under current climatic conditions: ecological and economic wildlife stocking rates’, in j. du p. bothma & j.g. du toit (eds.), game ranch management, 5th edn., pp. 801–815, van schaik publishers, pretoria. van rooyen, n., bezuidenhout, d., theron, g.k. & bothma, j.d.p., 1990, ‘monitoring of the vegetation around artificial watering points (windmills) in the kalahari gemsbok national park’, koedoe 33(1), 63–88. https://doi.org/10.1006/jare.1994.1037 van rooyen, n., van rensburg, d.j., theron, g.k. & bothma, j.d.p., 1984, ‘a preliminary report on the dynamics of the vegetation of the kalahari gemsbok national park’, koedoe (supplement), 83–102. https://doi.org/10.4102/koedoe.v27i2.570 van rooyen, n. & van rooyen, m.w., 1998, ‘vegetation of the south-western arid kalahari: an overview’, transactions of the royal society of south africa 53(2), 113–140. https://doi.org/10.1080/00359199809520381 weltzin, j.f., loik, m.e., schwinning, s., williams, d.g., fay, p.a., haddad, b.m. et al., 2003, ‘assessing the response of terrestrial ecosystems to potential changes in precipitation’, bioscience 53(10), 941–952. https://doi.org/10.1641/0006-3568(2003)053[0941:atrote]2.0.co;2 wessels, k.j., prince, s.d., malherbe, j., small, j., frost, p.e. & van zyl, d., 2007, ‘can human-induced land degradation be distinguished from the effects of rainfall variability? a case study in south africa’, journal of arid environments 68(2), 271–297. https://doi.org/10.1016/j.jaridenv.2006.05.015 wessels, k.j., steenkamp, k., von maltitz, g. & archibald, s., 2011, ‘remotely sensed vegetation phenology for describing and predicting the biomes of south africa’, applied vegetation science 14(1), 49–66. https://doi.org/10.1111/j.1654-109x.2010.01100.x wessels, k.j., van den bergh, f. & scholes, r.j., 2012, ‘limits to detectability of land degradation by trend analysis of vegetation index data’, remote sensing of environment 125, 10–22. https://doi.org/10.1016/j.rse.2012.06.022 appendix 1 table 1-a1: accuracy assessment of the landsat-based vegetation map for the tswalu kalahari reserve using a confusion matrix. appendix 2 table 1-a2: summary of the cross validation results from the ordinary kriging performed to interpolate rainfall in the tswalu kalahari reserve. figure 1-a2: maps of prediction standard error of rainfall surface (figure 4) in the tswalu kalahari reserve. (a) 2001–2002, (b) 2002–2003, (c) 2003–2004, (d) 2004–2005, (e) 2005–2006, (f) 2006–2007, (g) 2007–2008, (h) 2008–2009, (i) 2009–2010, (j) 2010–2011, (k) 2011–2012, (l) 2012–2013 and (m) 2013–2014. figure 1-a2 (continues...): maps of prediction standard error of rainfall surface (figure 4) in the tswalu kalahari reserve. (a) 2001–2002, (b) 2002–2003, (c) 2003–2004, (d) 2004–2005, (e) 2005–2006, (f) 2006–2007, (g) 2007–2008, (h) 2008–2009, (i) 2009–2010, (j) 2010–2011, (k) 2011–2012, (l) 2012–2013 and (m) 2013–2014. article information authors: susan j. taylor1 doreen atkinson1 affiliations: 1centre for development support, university of the free state, south africa correspondence to: susan taylor postal address: po box 339, internal 100, bloemfontein 9300, south africa dates: received: 15 feb. 2011 accepted: 02 dec. 2011 published: 21 may 2012 how to cite this article: taylor, s.j. & atkinson, d., 2012, ‘delivering community benefits acts as insurance for the survival of small protected areas such as the abe bailey nature reserve, south africa’, koedoe 54(1), art. #1043, 9 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1043 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. delivering community benefits acts as insurance for the survival of small protected areas such as the abe bailey nature reserve, south africa in this original research... open access • abstract • introduction    • protected areas and sharing benefits    • buffer zones and land use changes outside protected areas    • protected areas and human development    • the vulnerability of small protected areas • methods    • procedure    • study area       • abe bailey nature reserve       • profile of khutsong township • results    • community needs relating to the abe bailey nature reserve       • the ‘care for the aged’ home-based care group       • ‘millennium women’, a women’s group involved in home-based care       • khutsong youth group       • bambanani group of traditional healers of khutsong       • new mandela village informal settlement residents    • protected area management needs    • lack of neighbour relations policy    • final project selection • discussion    • protected area buffer zones in south africa    • what protected areas can mean in a ‘township’ context    • the survival of small protected areas • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ the abe bailey nature reserve (abnr) in the gauteng province of south africa is largely unknown and offers little to attract visitors. the biological integrity of the abnr is challenged by the urban poverty in khutsong, the reserve’s immediate neighbour. relations between khutsong and the nature reserve had been hostile for decades as a result of the ‘fortress’ style of conservation protection used for the abnr. however, this situation provided the gauteng directorate of nature conservation with an opportunity to experiment with identifying and transferring benefits to the community, as well as establishing an effective buffer zone between the nature reserve and the informal settlements of khutsong. following an initial rapid rural appraisal and ongoing liaison through specifically appointed project managers, an outreach programme containing two natural resource-based projects was developed. as a result, better relations were established between the abnr and its neighbouring community for the first time since the nature reserve was established in 1977. this acted as ‘insurance’ during violent public protests and vandalism in the khutsong border demarcation dispute (2005–2007), but may not be enough to secure the nature reserve into the future. conservation implications: small protected areas may not be effective in ensuring their biological integrity in the long term, but working cooperatively with existing and future neighbours is an essential strategy to optimise conservation activities in small reserves such as the abnr. introduction top ↑ protected areas and sharing benefits while the primary role of protected areas is the conservation of species diversity, biodiversity conservation, along with sustainable resource management, can and must result in material benefits to neighbouring communities. the united nations convention on biological diversity (secretariat of the convention on biological diversity 2008) states that protected areas are important tools for the conservation of biological and often supply important ecosystem goods and services and must form the cornerstone of sustainable development strategies; yet, they are often under-resourced to play these roles. additionally, individual protected areas are becoming increasingly isolated and ever more in contact with growing human populations, who, at the same time, are struggling to find land and resources on which to survive, specifically in developing countries (straede & treue 2006). conservation and poverty reduction need to be tackled together for best results and, as such, achieving local cooperation and support without jeopardising conservation goals has become a top priority for most parks around the world (adams et al. 2004; wittmayer & büscher 2010). this predicts an ongoing battle between the need to keep protected areas intact and the needs and impact of the people outside these areas. to ensure that protected areas achieve local cooperation and survive into the future, a deep understanding of the socio-economic dynamics that determine the current and future use of land resources in and around the protected area is essential (defries et al. 2007). community-based conservation and development has become the prevailing paradigm of conservation organisations and development donors over the last 20 years (browder 2002; gjertsen 2005). however, there is considerable complexity in ensuring that this happens in practice. considerable management, social, financial and ecological skills, as well as the latest conceptual thinking by park management, needs to be brought to bear on attaining community support. more often than not, successfully managing large parks through trying to reconcile the aspirations of the local people has been difficult to achieve (mbile et al. 2006) and the many failures since the 1980s have resulted in a proliferation of consultative processes to engage with communities living outside or within protected areas. the large, donor-funded community-based integrated conservation and development projects, which have formed a well-known approach to dealing with both conservation and poverty through the attempt to reconcile park management with local needs and aspirations, have drawn criticism from conservation biologists for failing to ensure adequate protection of biodiversity (browder 2002). also, other approaches, such incentive-based programmes, which aim to foster local stewardship by delivering benefits tied directly to local biodiversity conservation by local people, have been tried with mixed results (spiteri & nepal 2008). these have resulted in recent experimental approaches that try to incorporate elements of adaptive management and institutional development, as well as form new partnership models with stakeholders and create the vertical integration of site-level work with policy initiatives (wells & mcshane 2004). baker, milner-gulland and leader-williams (2011) point out that conflicts between protected-area managers and local people are common, but the drivers of such conflicts are rarely analysed. the lack of case studies and data limit opportunities to identify strategies that reduce conflict and the magnitude of resulting threats to conservation; this lack of evidence and analysis may set up some community programmes for failure. reid et al. (2004) point out a growing trend towards protectionism, possibly because of the many failures and ongoing challenges in making the relationship between people and biodiversity conservation work. as an alternative to both community biodiversity partnership models and protectionism, sodhi et al. (2010) warn that ‘local people must be provided with alternative sustenance opportunities and basic education in addition to environmental outreach to reduce their reliance on protected forests and to enhance conservation support’. in south african, nature conservation has moved away from a strictly protectionist approach to the recognition that biodiversity must benefit local people (botha, witkowski & cock 2006; mulder, caro & msago 2007; wittmayer & büscher 2010; wynberg 2002). the principle that biodiversity and protected areas must benefit people is contained in a ‘white paper on the conservation and sustainable use of south africa’s biological diversity’ (department of environmental affairs and tourism 1997) and in the national environmental management: protected areas act 2003 (act no. 57 of 2003) (republic of south africa 2003), which states that protected areas must be able to deliver some needed benefit to neighbours and that mechanisms to ensure this must be outlined in that protected area’s management plan. in south africa, larger conservation projects, such as the maloti-drakensberg transfrontier conservation and development project, are established with the aim of playing a large role in local development (wittmayer & büscher 2010). also, south african national parks (sanparks) plays a large developmental role in the areas surrounding the national parks, through, for example, contractual parks (reid et al. 2004). the challenge is for the smaller protected areas in south africa (i.e. municipal and provincial reserves), which have fewer financial and skills resources, to engage with this complexity. buffer zones and land use changes outside protected areas in general, natural ecosystems are under enormous pressure from the growing demands of human development (guo, zhang & li 2010; iwamura et al. 2010). although protected areas now cover over 12% of the terrestrial surface of the earth, many fail to protect species and ecological processes as originally envisioned. results of recent studies suggest that a critical reason for this failure is an ever-increasing contrast between the protected area and the surrounding matrix of often highly altered land outside the protected area (seiferling et al. 2011).to mitigate against the impact of human development, international best practice for protected areas includes creating ‘buffer zones’ where the interface between park and people is managed. these can include areas designated for conservation, community forestry, limited resource extraction or agriculture (bajimaya 2006; lynagh & urich 2002; sanparks n.d.). buffer zones are intended as areas where communities can engage with natural resource use and generate revenue from biodiversity rather than having access to the protected area itself (bajimaya 2006). whilst the intention of buffer areas is to resolve park–people conflicts over resource use, straede and treue (2006) show that, in nepal, there is a growing gap between local people’s need for natural resources and their rights to satisfy them on a legal basis in the buffer areas, meaning that the situation is unsustainable in the longer term, and this is the situation for many protected areas. other studies show that, inevitably, the buffer zone degrades from overuse and the local communities begin to make forays into the protected area itself to satisfy their needs (lynagh & urich 2002). constant monitoring and adaptive management is needed to ensure a ‘win–win’ situation between protected area and neighbours and that buffer strategies may have to change over the years to ensure sustainability (lynagh & urich 2002). ma et al. (2009) show that even in a biosphere situation meant to accommodate human activity, the declared buffer zone becomes over-utilised with serious consequences for the biodiversity within the ‘core’ region of the biosphere reserve. integrating biodiversity conservation and the development of local communities is therefore a major challenge (ma et al. 2009). protected areas and human development in some rural areas, there are new concerns that the protected areas themselves may cause the human population density to increase artificially on the outside of parks because of the opportunities and benefits protected areas offer in a world of decreasing resource availability (wittemyer et al. 2008). countering this finding, joppa, loarie and pimm (2009) showed that when population growth does occur near protected areas, it results from a general expansion from nearby population centres. working with the kafue national park in zambia, joppa et al. (2009) found that, over time, distant rural population centres grew outwards in all directions, threatening kafue national park. the national park was ‘simply in the way’ of human population expansion. inspection of many other parks shows this to be a common trend (joppa et al. 2009). this is the situation in gauteng, where urban development is beginning to engulf small protected areas such as the abe bailey nature reserve (abnr). in other developed countries, gimmi et al. (2011) also found that human density tended to increase outside protected areas through urban development and increased residential market values of land near ‘nature’. the availability of land to declare an appropriate buffer zone for a protected area is a critical constraint because of the frequent lack of available land surrounding the protected area. in the urban or peri-urban environment, buffer zones are especially difficult to create because land use is usually already designated, or land is highly priced. the vulnerability of small protected the impact of human development on the outside of large reserves is mitigated to some extent by the size of the reserve, but for small reserves, the impact can mean that, over time, the biodiversity within the reserve begins to deteriorate, following the dominant land-use change pattern into which they are located (maiorano, falcucci & boitani 2008). as such, maiorano et al. (2008) suggest that small parks will not be viable in the long term, especially if they are allowed to become ‘islands’ in a human-dominated landscape. identification of appropriate and effective management opportunities for a small nature reserve depends on defining the biodiversity attributes of concern; identifying landscape connections to delineate strong ecological interactions between the protected area and its surrounding landscape and identifying socio-economic dynamics that determine current and future use of land resources in and around the protected area. the challenge is to identify management opportunities that result in the maintenance of ecological function in the wider landscape without imposing undue restrictions on human land use (defries et al. 2007). the abnr is a small (4197 ha) nature reserve located within a human-dominated landscape (urban development, agriculture and mining activities) and this paper reviews some of its struggles to remain as a functioning entity on the edge of urban development, conserve biodiversity and deliver benefits to the neighbouring community. its biodiversity attributes include a sample of carletonville dolomite grassland (listed as vulnerable), a genetically pure herd of black wildebeest, connochaetes gnou (dna tested), a population of the white-tailed rat, mystromys albicaudatus, a wetland, an extensive cave system which is part of the dolomitic geology of the area, as well as 262 bird species. no planning work has been undertaken to understand how this nature reserve contributes to ecosystem or species conservation in the wider landscape. outside the reserve, at the abnr–khutsong interface, the land use has already been designated as urban, with bigger agricultural and agri-business expansion also planned for the area. the merofong city integrated development plan (idp) (merofong municipality 2011) makes scant mention of the nature reserve, merely citing that it is an ‘impediment to the northwards expansion of urban khutsong and wielverdiend’. the notion of a protected area playing a role in determining surrounding land use, as urged by defries et al. (2007), would be very difficult to achieve at the abnr without considerably more resources. methods top ↑ procedure a qualitative approach was used to analyse, post hoc, the history and outcomes of community outreach programme initiated in 2000. this paper is compiled from the personal experiences and notes of the primary author, who was one of the project initiators involved at a strategic level at the gauteng department of agriculture conservation and environment (gdace) from 1999 to 2006, as well as from two formal interviews held with the officer-in-charge at the abnr in 2010 and 2011, and from interviews with two of the former project managers of this project, the late madire malepe (pers. comm., 21 july 2008) and dr thato shale (pers. comm., 30 october 2010). departmental records of the rapid rural appraisal (rra) held between residents from khutsong and the gdace in 2000 were used to review the findings of this initial community consultation. the rra sessions probed general social and economic needs of the community, as well as how the community related to the abnr, and sought to identify any ecological, spiritual or other needs that the nature reserve could provide, which would improve the working relationship between the community and the reserve. in preparation, the governmental group from the gdace was given a one-day training session on how to conduct an rra (most of the departmental participants in the project were nature conservation officials with no formal training in social science methods), held by the late james mascarenhas, an expert in participatory research practice (mascarenhas 1991). although notes were taken at the time, no detailed data were recorded at the rra sessions. the rra techniques used with the khutsong groups included mapping, matrix ranking, seasonality diagrams, venn diagrams and timelines (j. mascarenhas, pers. comm., 24 october 2000). an initial liaison with merofong municipal councillors in 2000 guided the selection of community groups for the rra. the five groups interviewed were a home-based care group called ‘millennium women’, a home-based care group called ‘home for the aged’, a youth group, the bambanani group of traditional healers, and a group of residents of the new mandela village informal settlement adjacent to the nature reserve. study area abe bailey nature reserve the abnr falls within the merafong city municipality and is a provincial nature reserve in gauteng province, located 7 km north-west of carltonville, between 26°16′ s and 26°20′ s and 27°17′ e and 27°22′ e (eber 1998). sir abe bailey, a wealthy ‘randlord’ from johannesburg’s early years, originally owned the land and used it as a hunting preserve around the turn of the 20th century (eber 1998). in 1977, the far west rand dolomitic association (fwrda), who still own much of the land in the far west rand, made two farms available to the transvaal branch of the wildlife society of southern africa for the creation of a nature reserve, and a five-year lease was signed (eber 1998). in 1980, the fwrda offered further land adjoining the reserve to the wildlife society. the wildlife society was unable to afford such a venture and so offered the reserve to the then transvaal provincial administration. transvaal nature conservation assumed control of the reserve on the 01 april 1982. on 11 may 1988, a 50-year lease was signed between the transvaal provincial administration and fwrda for the abnr. this lease was transferred subsequently to the gauteng provincial government on 10 may 1994 (eber 1998). whilst the abnr was proclaimed in 1977 and was intended to be a multi-racial recreational facility, this never came to fruition and the reserve was severely underutilised by all race groups during the apartheid years (eber 1998). there were many management factors which alienated the residents of khutsong from the nature reserve over the years. primarily, the reserve had a history of ‘brutal enforcement of its policy of fences, arrests and fines’ (eber 1998). prior to the democratic elections in 1994, the rangers reportedly patrolled the perimeter with semi-automatic firearms. in 1992, 251 dogs were shot on the reserve, escalating conflict with the community. other ongoing community-related problems on the reserve included arson, poaching of wildlife, removal of fences and the use of the nature reserve as a short cut to a main road leading into carletonville (eber 1998). by the late 1990s, the abnr was in a dilapidated condition, had never reached its potential as a recreational resource for black people in the area and was facing closure (eber 1998). the boundary fence between new mandela village and the abnr had been erected and removed many times and, by 1999, shacks, rubbish dumps, soccer fields and stands of maize could be found within the nature reserve, adding to the picture of dereliction. thus, in 1997, nature conservation officials formally petitioned the gdace head of department for permission to set up a task team to investigate options for the abnr to try and resolve the ongoing conflict with khutsong (eber 1998). officials explained the need to investigate the potential of the reserve as an asset to the people of khutsong. finally, in 2000, this request was taken up with an rra as the first formal post-1994 engagement between the gdace and the khutsong residents. the nature reserve presently covers an area of ± 9780 ha, after the addition of land donated by the fwrda and the local mining houses (merafong municipality 2011). profile of khutsong township khutsong is a crowded urban settlement that borders on the abnr. khutsong was established in 1958 as an apartheid township to relocate people living in slums that had developed on white-owned farms, as well as to provide housing for mine labour (botha 2003). the township is composed of many formal and informal sections. one of the poorest of these is the new mandela village, an informal settlement located on unsafe land (i.e. land with a prevalence of developing sinkholes) without basic services, and which is the immediate neighbour of the abnr. the western border of the nature reserve is also the gauteng–north west provincial border and the border of merofong municipality with north west province. the presence of unstable dolomite geology limits development in the area.after the first national democratic elections in 1994, khutsong was designated as part of the merofong municipality in gauteng. service delivery was reported to be notably poor, experiencing little or no change since khutsong was neglected under apartheid (botha 2003). there was mounting public frustration and violence within khutsong in the post-1994 period in relation to service delivery (botes et al. 2007). in 2003, a social scan undertaken by botha (2003) estimated that unemployment in merofong municipality ranged from 25% to close to 60% in different parts of khutsong. poverty was very prevalent and numerous households were heavily dependent on child and/or disability grants and/or pensions. young men lacked jobs and future prospects, even though many had completed formal schooling (botha 2003). nevertheless, khutsong was a diverse and active place and botha (2003) identified dozens of community groups in khutsong, including women’s groups, hiv and/or aids home-based care groups, faith groups, youth groups, trade unions, men’s stokvel [savings] groups, food garden groups, sports and soccer clubs, arts and culture groups, health groups and business groups. from 2005 to 2007, local merofong municipality residents, particularly those living in khutsong, engaged in a dispute with the national government over their unwillingness to be incorporated into north west province. residents feared the new demarcation would make them worse off in terms of service delivery. the impact of this unrest was manifested through the destruction of public and private property in khutsong, costing an estimated r70 million in damage by april 2006 (botes et al. 2007; mavungu 2011). the violence ended in 2009 when a government decision was gazetted that khutsong would be returned to gauteng province. results top ↑ community needs relating to the abe bailey nature reserve the rra held in 2000 with the selected community groups at khutsong revealed a plethora of needs, concerns and opportunities for action, even though many of these could not be taken further by gdace’s nature conservation or agricultural directorates as they fell outside the departmental mandate. in some cases, the issues were referred to relevant government departments. the rra also noted specifically how the community related to the abnr. key rra findings from each of the five groups relating to the abnr are summarised in the subsections below. the ‘care for the aged’ home-based care group the older persons looked after by the care for the aged group had little knowledge of the abnr and its biodiversity objectives, even though they had lived in the area for decades. what was important for this group was land for gardening and a ‘working centre’ where vegetable gardening could be undertaken in a safe environment. there was also the suggestion that able-bodied elderly people could be employed by the reserve, rather than only young people being selected for work such as burning fire breaks. ‘millennium women’, a women’s group involved in home-based care the members of this group were involved in caring for hiv and/or aids patients and the aged, but stated that little support comes from government and that, generally, the health services in the areas are poor. there were many orphans in the area who had suffered the death of parents caused by hiv-related and/or aids-related diseases. the group also assists the elderly to claim their social security grants. the women’s group had a very good understanding of the abnr and had several suggestions on how it could become relevant to the khutsong people. they did not express any direct ‘needs’ for themselves. the women stated that the abnr should be utilised for the following:• agriculture (food security projects and community cattle grazing) • environmental education • medicinal plants propagation (in a nursery on the reserve). the millennium women’s group also requested access to the nature reserve to collect ’greens‘ (morogo), which refers to a group of at least three different dark-green leafy vegetables found throughout southern africa, harvested for human consumption and which represent a seasonal-free food source. khutsong youth group this was a group of young people who met regularly at the one of the small clinics in the informal settlement. the youth group included 16 young men aged between the ages of 16 and 30 years. the youth group was aware of the abnr and that it promoted nature conservation. its members were also aware that the reserve sometimes offered temporary work opportunities and that this was a benefit. they also acknowledged that the nature reserve provided ‘free medicinal plants’. negative issues from the youth included the statement that the rangers ‘beat us up instead of taking us to the police station if we are caught on the nature reserve and they think we are stealing animals’ (khutsong youth group member, 25 years). the youth did not suggest any ‘needs’ for themselves, but suggested that there is a need to renovate the education centre at the abnr and run an awareness programme about nature conservation. the youth group mentioned that the reserve is not accessible to local people because they do not know anything about it and suggested information about the nature reserve should be presented in local schools. bambanani group of traditional healers of khutsong the government consultation only reached one traditional healer group who called themselves the bambanani group, although continued liaison with khutsong residents revealed that there were at least six traditional healing organisations in khutsong (botha 2003). it was known by the gdace officials that the bambanani group acted as gatekeepers and inhibited other traditional healers or healer groups from participating in activities organised by the nature reserve. this was difficult to overcome with limited resources and, as such, the department resigned itself to dealing with the bambanani group. however, the dealings with this group have proved to be productive over the years and it has been recognised that the aim of the project was not to address all concerns and help all groupings within khutsong.fourteen traditional healers (twelve women of various ages and two men of retirement age) from the bambanani group attended the initial rra workshop in 2000. the traditional healers in this group were all very poor and stated that their patients were even poorer and very often could not pay for treatment. most of the plants that they used in their practices and which they considered important medicinally do not come from the abnr, but are bought from suppliers from kwazulu-natal. the plants the traditional healers harvested in the reserve, or from the surrounding farm lands, are relatively insignificant to them, so access for the sustainable harvesting of these plants in the abnr was not a big need for the healers. what was important for the bambanani group was access to land to grow the specific plants they needed. the traditional healer group requested that provincial government allow them to plant orchards of medicinal tree species and wetlands plants for their use within the boundaries of the abnr. they gave the gdace a confidential list of their required species. new mandela village informal settlement residents this group (about 40 men and women) informed the gdace officials that the municipality did not provide them with basic municipal services and that they lived in a situation of dire need. the lack of jobs was a serious problem. of the 46 households represented at the rra session, only seven people had formal incomes. informal sources of income for new mandela village residents include selling paraffin and vegetables, cooking and selling fresh maize cobs, fixing radios, sewing, informal maize farming, working in carletonville as gardeners and domestic workers, and wood carving for the tourist trade. people in the area also did not have access to land for household cultivation purposes and, as a result, food security was a key issue. it was enlightening to understand how new mandela village residents perceived the ‘good things’ about the abnr. they stated that they received free meat, free firewood, free morogo, free living space (land on which to locate their shacks) and free fencing and poles from the nature reserve. in terms of the nature reserve management and its requirement to protect government assets such as game animals and infrastructure, these ‘benefits’ would all be illegally acquired and essentially constituted theft and poaching. the negative statements about the abnr included: not enough jobs on offer, that they were chased away by the rangers, that the rangers shot their dogs and that there was no communication between reserve staff and new mandela village residents. the new mandela village group also stated that they never see any wild animals so are puzzled about the purpose of the nature reserve; they see it as land that is not being used. new mandela village residents had many suggestions about how the abnr could help them meet their needs. these included suggestions that the reserve create a children’s garden and find other ways to contribute to local children’s education and aspirations. also, new mandela village residents wanted to see reserve land be released for food cultivation. people in the area also wanted to use the abnr for cattle grazing. protected area management needs the gdace officials from the directorate of nature conservation hoped that the rra consultation would enable them to do something ‘good’ for the community and also achieve a ‘win–win’ situation for the nature reserve. they hoped that residents of khutsong would come to understand why the nature reserve land had been set aside for biodiversity conservation and that this understanding would reduce conflicts between the management of the protected area and the community. although not part of the rra, it was recognised that a buffer zone was needed to mitigate the impact of the khutsong settlement on the nature reserve. this need was articulated and negotiated over time by the first project manager, the late mr madire malepe, with representatives of the merofong municipality and with affected residents of khutsong, including those who participated in the original rra. in creating the buffer zone, consideration was given to the fact that some of the shacks of khutsong were already next to the boundary of the abnr and because these shacks could not be moved back from the ‘fence’ to create the buffer, the buffer area was set inside the nature reserve, using nature reserve land. lack of neighbour relations policy before the interaction with khutsong residents, the gdace had a generic ‘neighbour relations’ policy, essentially about ‘nurturing the community’. once the khutsong rra was performed in 2000, a more realistic picture of community engagement emerged (although this did not constitute a replacement policy). whilst post-rra relations were very cordial, it was realised by the gdace officials that a relationship with khutsong residents would need to be ‘toughed out’ over time, whilst engaging in a spirit of openness, and that staff skilled in community liaison would need to be recruited and deployed. over time, skilled staff were recruited to manage the two outreach projects and build the relationship with the community. in response to the findings of the rra in 2000, and in realising that the abnr is very small, the management plan committee took the decision that no direct use of the nature reserve by the neighbouring khutsong community would be considered, although there was no formal departmental policy on transferring benefits to communities. this stance was also informed by a medicinal plant study dzerefos and witkowski (2001), which recommended that no harvesting rights be bestowed on local traditional healers and medicinal plant traders. regular discussions about project challenges were held both at departmental level (in the form of gdace quarterly review meetings) and at the level of the abnr management plan committee. final project selection the one-day rra consultation with five khutsong community groups in january 2000 enabled the gdace’s directorate of nature conservation to plan a programme of two outreach projects linked to the nature reserve, whilst rejecting other suggestions from the community. findings of the rra indicated that most of the problems the residents of khutsong and new mandela village faced were related to poverty and extreme deprivation, lack of work opportunities and poor municipal service delivery and had few links to the mandate of the abnr or the core business of the gdace (i.e. biodiversity conservation, food security and agriculture). this both limited and influenced the final project selection. in the final selection of projects for the outreach programme, some ideas and suggestions (both from the community and from conservation officials) had to be rejected as being outside the mandate of the gdace, difficult to manage, or out of alignment with biodiversity conservation principles. for example, cattle grazing by the local community was considered initially as a potential ‘benefit’ but, in the end, was rejected by the province because of the difficulties in managing the equitable allocation of this benefit. also, the community stated that they wanted to be allowed to hunt with dogs on the nature reserve, but this was deemed an unacceptable practice by the management plan committee. the community also wanted to receive meat from game culls on the reserve, as well animal parts to be used for ‘muthi’ [witchcraft purposes]. these latter requests were not practical, as most of the game surplus was sold live to game capture organisations. the request by the bambanani group for a medicinal tree plantation was also eventually refused by the gdace ecologists on the grounds that these species were foreign to the typical grassland habitat of the abnr and could become invasive over time.during 2000 and 2001, a 50-metre wide strip along the western edges of the nature reserve was set aside to be developed as an agricultural project. the 50 m was easy to designate because of a gravel road which ran along the edge of the nature reserve, 50 m from where the fence line was originally located. this strip ran for approximately 6 km and provided 147 ha of land, or 3.5% of the nature reserve land area, and was formalised in 2002. officials also felt that the words ‘buffer zone’ were unpalatable in that it implied the intention to keep the community at bay; as such, the title ‘green zone’ was used for official communications. the need for land for food gardens had been underestimated by the gdace before the rra consultation and it was agreed by provincial government that the green zone land could be released for food growing. a large portion of this was divided into 196 plots (indicating 196 lease holders) in the initial allocation. a plot size of 50 m × 50 m (about 0.5 ha) was recommended by the gdace directorate of agriculture as a viable plot size and this was also negotiated with the participants in various start-up meetings. participants wanted small plots that were easy to manage. allotment applicants were screened through an interview process according to their willingness to use the land according to the lease conditions, as well as whether they had prior experience in agriculture or vegetable growing. a fenced plot (about 0.5 ha) of land within the nature reserve was also allocated to the bambanani group of traditional healers (14 recipients) in 2000 for medicinal plant cultivation. the group grew wild ginger (siphonochilus aethiopicus) and african potato (hypoxis hemerocallidea). a tap provided municipal water for the plants. the bambanani group also received training from the resident abnr horticulturalist on cultivation methods. a study was later commissioned by the gdace to guide the development of an outreach medicinal plant nursery and overcome predicted failures (botha 2003). the study showed how funding and support has to be provided for a considerable length of time before the projects are handed over to be run independently by the community (botha 2003). although the abnr community projects were financed by provincial government, and an associated medicinal plant rescue scheme and infrastructure (greenhouses) was funded for three years (2006–2008) through the gtz (deutsche gesellschaft für technische zusammenareit, now called giz or deutsche gesellschaft für internationale zusammenareit), the projects ultimately failed as a result of unexpected political reasons. owing to the khutsong border demarcation protests, the gauteng provincial government adopted a holding position and only funded core salaries between 2003 and 2011. the gtz funding ended in 2008 and was not renewed because of the border demarcation issue and, since then, the greenhouse has stood empty (gdard staff, pers. comm., 25 march 2011). as a side note, the gdace became the gauteng department of agriculture and rural development (gdard) in 2009. vegetables continue to be cultivated on some of the ‘green zone’ plots (abnr officer-in-charge, pers. comm., 25 march 2011). discussion top ↑ protected area buffer zones in south africa in south africa, the national environmental management act: protected areas act 2003 (republic of south africa 2003) makes no provision for the declaration of buffer zones and thus any buffer zones have no legal status unless the land is purchased. sanparks (n.d.) have developed their own buffer zone policy to influence burgeoning development outside the national parks, particularly the kruger national park.gauteng province, south africa’s most urbanised province, also has no over-arching buffer zone policy for its six protected areas, although urban development is encroaching rapidly on all these reserves. however, there are internal policies and ‘guidelines’ that deal with the types of development allowed around the protected areas. there is also no general neighbour relations policy to guide how benefits from protected areas are to be transferred (gdard staff, pers. comm., 25 march 2011). what protected areas can mean in a ‘township’ context since its proclamation as a nature reserve in 1977, the abnr it has not been used by black khutsong and carletonville residents as a recreational area, even in the post-1994 era and despite the very over-crowded conditions in khutsong. neither before nor after 1994 have the residents of khutsong ever tried to claim the abnr through land invasions. whilst some living in new mandela village had erected their shacks inside the reserve, they had never gone further than a perimeter gravel road (50 m from the boundary fence). in fact, it would appear that most of the township seems indifferent to the nature reserve. yet, the nature reserve has meaning for some groups. the traditional healers (bambanani group) saw it as an area where the growing of medicinal plants could take place. the youth saw possibilities for hunting with dogs. some people with an agricultural background (many of the eventual allotment holders) saw an opportunity to gain a small plot of land for cultivation. the home-based care group, the new mandela village group and the khutsong youth group, saw the potential educational value of the nature reserve. the value of the nature reserve to khutsong residents is the sum of these perceptions and uses. yet, the environmental education facility identified by khutsong groups that were consulted has never materialised, despite the new facilities built on the abnr (gdard staff, pers. comm., 25 march 2011). this has been largely attributed to the impact of the border demarcation protests. the abe bailey outreach programme based on the rra in 2000 also did not yield the large number of sustainable food gardens and medicinal plant gardens that were originally envisaged by both the khutsong groups and the nature reserve management, and would have needed substantially more inputs from government, donors and the private sector to create sustainable benefits. water and infrastructure for irrigation were the main constraints to the green zone agricultural activities, which could have been rectified with water storage tanks and pumping water from the nearby wetland to a high point and then using flood irrigation to water plots, but this would have required investment in a pump and cement ditch with sluice gates. at that stage, new mandela residents used a ‘pay-as-you-go’ meter system to buy water for their households – a system that was regarded as ‘very expensive’ for householders. yet, one lease holder produced ‘enough spinach for sale’ throughout the year by using household waste (grey) water, which he purchased from nearby shack owners. whilst sanparks has a detailed buffer zone policy, smaller protected areas such as the abnr have to develop their own policy and, with minimal resources, implement this in an era with a strong development focus. whilst there is still no formal buffer zone policy for the abnr, or any overt patrolling of the border zone, it is apparent that the buffer zone land itself is somehow respected by the khutsong residents (particularly the people of new mandela village) as a boundary, although trespass and arson continue. since the gdace–khutsong engagement during 2000 and 2008, no new shacks have been erected in the buffer zone area, nor has there been encroachment of any kind beyond the buffer zone area into the nature reserve (google earth 2012). the only regulatory mechanism used to enforce the green zone was the one-year lease system and leases signed by allotment recipients in 2001. the lease arrangements have all now expired because of the border demarcation protests (abnr officer-in-charge, pers. comm., 25 september 2010). one of the important findings of this exercise was that the development of a community outreach programme does not depend solely on what either party (nature reserve management or community) needs, wants or demands, but depends, instead, on the sustained allocation of provincial resources (staff hours, capital expenditure, government budgets) or donor project funding to maintain relationships and a programme of work. the survival of small protected areas maiorano et al. (2008) have warned that small protected areas in a human-dominated landscape will, in time, lose all the characteristics for which they have been established. to deal with this threat, conservation planners need to devote much more attention to the non-protected matrix in which their small protected areas exist and in which they must survive (maiorano et al. 2008). in the case of the abnr, this would involve paying greater attention to changing land use within the wider area, as well as engaging with stakeholders other than khutsong residents (i.e. municipal planners, farmers and agri-businesses). the merafong idp states that merafong city is responsible for protecting environmentally sensitive areas within the municipal area from exploitation and damage resulting from direct or indirect urban development (merafong municipality 2011). the idp also notes that the abnr is one of several tourism areas and facilities located within the municipal area and acknowledges that important challenges facing the reserve are changes in land use causing a loss and fragmentation of natural habitat in surrounding areas, windborne and waterborne pollution, as well as socio-economic factors such as poverty that lead to the over-exploitation of natural resources (merafong municipality 2011). the most surprising benefit of the abe bailey outreach programme was to the nature reserve staff, infrastructure and other assets. during the khutsong provincial border demarcation riots of 2005–2007, the newly built nature reserve infrastructure valued at r7 million was spared vandalism and staff were not threatened. it is the view of the staff at the abnr (abnr officer-in-charge, pers. comm., 24 april 2011) and also the opinion of the late mr madire malepe (2008) that this protection was largely the result of an improvement in relations between the abnr and its neighbours since 2000. conclusion top ↑ in south africa, the national environmental management act: protected areas act 2003 (republic of south africa 2003) requires government-funded protected areas to provide ‘benefits’ to neighbouring communities and contribute to the development needs of poor communities on the outside of the reserve. in building relationships between small protected areas and neighbouring communities, there are many research and/or consultative approaches, such as the rra method, which can help uncover issues and needs on both sides and identify relevant projects and interventions. whether taking the form of outreach or community-based natural resource management, these programmes can take a long time to implement, mature and deliver benefits. financial support, whether from government or donors, can falter along the way. unexpected impacts, for example, from urban political unrest, can stall projects. yet attending to (selected) development issues on the outside of protected areas through community liaison and joint projects may be a sound way to secure a protected area into a more violent and unstable urban future. surprisingly, just the act of engaging with the community in a fair and transparent way yielded positive results for the abe bailey programme, as the actual outreach projects were only marginally successful. whilst incurring costs in terms of staff hours, building better relationships acted as a form of insurance against the threats that township protests can bring, as seen by the way that the abnr did not suffer the kind of vandalism that other government properties suffered in khutsong during 2003–2008. acknowledgements top ↑ we wish to acknowledge the staff of the gdard (formerly known as gdace), including those in the directorate of conservation and the directorate of agriculture, who worked on both projects from 2000 to 2007. we also wish to acknowledge the contribution of the community groups within khutsong. competing interests the authors declare that they have no financial or personal relationship(s) which may have inappropriately influenced them in writing this article. authors’ contributions s.t. (university of the free state) conducted the interviews with gdard officials, reviewed the written material and records from the original project and wrote the manuscript. d.a. (university of the free state) as the second author was responsible for conceptual contributions and suggestions. references top ↑ adams, w.m., aveling, r., brockington, d., dickson, b., elliott, j., hutton j. et al., 2004, ‘biodiversity conservation and the eradication of poverty’, science 306(5699), 1146–1149. http://dx.doi.org/10.1126/science.1097920, pmid:15539593 baker, j., milner-gulland, e. & leader-williams, n., 2011, ‘park gazettement and integrated conservation and development as factors in community conflict at bwindi impenetrable forest, uganda’, conservation biology 26(1), 160–170. http://dx.doi.org/10.1111/j.1523-1739.2011.01777.x, pmid:22044616 bajimaya, s., 2006, ‘nepal’s buffer zone programme: a showcase of a participatory approach to protected area management’, in p. gyamtsho, b.k. singh & g. rasul (eds.), capitalisation and sharing of experiences on the interaction between forest policies and land use patterns in asia: linking people with resources, vol. 2, technical papers, p. 125–141, international centre for integrated mountain development, kathmandu. botes, l., lenka, m., marais, l., matebesi, z. & sigenu, k., 2007, the cauldron of local protests: reasons, impacts and lessons learned, centre for development support, university of the free state, bloemfontein. botha, j., 2003, ‘a preliminary study towards the setting up on an operational medicinal plant nursery at abe bailey nature reserve’, draft report to the gauteng department of agriculture, conservation and environment, johannesburg. botha, j., witkowski, e.t.f. & cock, j., 2006, ‘the south african experience of conservation and social forestry outreach nurseries’, journal of environmental management 38, 733–749. http://dx.doi.org/10.1007/s00267-005-0184-4, pmid:16508799 botha, j., witkowski, e.t.f. & cock, j., 2007, ‘is something better than nothing’? the impacts of ten outreach nurseries on south african community stakeholders’, forests, trees and livelihoods 17, 199–221. browder, j.o., 2002, ‘conservation and development projects in the brazilian amazon: lessons from the community initiative program in rondônia’, journal of environmental management 29(6), 750–762. http://dx.doi.org/10.1007/s00267-001-2613-3, pmid:11992169 chambers, r., 1994, ‘the origins and practice of participatory rural appraisal’, world development 22(7), 953–969. http://dx.doi.org/10.1016/0305-750x(94)90141-4 defries, r., hansen, a., turner, b.l., reid, r. & liu, j., 2007, ‘land use change around protected areas: management to balance human needs and ecological function’, ecological applications 17, 1031–1038. http://dx.doi.org/10.1890/05-1111, pmid:17555216 department of environmental affairs and tourism, 1997, ‘white paper on the conservation and sustainable use of south africa’s biological diversity’, general notice 1095 of 1997, deat, pretoria, viewed 28 march 2012, from http://www.environment.gov.za/polleg/whitepapers/biodiversity/contents.htm dzerefos, c.m. & witkowski, e.t.f., 2001, ‘density and potential utilization of medicinal grassland plants from abe bailey nature reserve, south africa’, biodiversity and conservation 10, 1–22. eber, s., 1998, ‘establishment of task team for approval by the head of department’, internal departmental memo within the department of agriculture, conservation and environment, johannesburg. gimmi, u., schmidt, s.l., hawbaker, t.j., alcántara c., gafvert u. & radeloff v.c., 2011, ‘increasing development in the surroundings of us national park service holdings jeopardizes park effectiveness’, journal of environmental management 92(2011), 229–239. gjertsen, h., 2005, ‘can habitat protection lead to improvements in human well-being? evidence from marine protected areas in the philippines’, world development 33(2), 199–217. google earth, computer software, viewed 28 march 2012, from http://www.maplandia.com/south-africa/guateng/oberholzer/khutsong/ guo z., zhang l. & li y., 2010, ‘increased dependence of humans on ecosystem services and biodiversity’, plos one 5(10), e13113, viewed 10 november 2011, from http://dx.doi.org/10.1371/journal.pone.0013113 iwamura, t., wilson, k.a., venter, o. & possingham, h.p., 2010, ‘a climatic stability approach to prioritizing global conservation investments’, plos one 5(11), e15103, viewed 10 november 2011, from http://dx.doi.org/10.1371/journal.pone.0015103 joppa, l.n., loarie, s.r. & pimm, s.l., 2009, ‘on population growth near protected areas’, plos one 4(1), e4279, viewed 10 november 2011, from http://dx.doi.org/10.1371/journal.pone.0004279, pmid:19169358 kintz, d.b., young, k.r. & crews-meyer, k.a., 2002, ‘implications of land use/land cover change in the buffer zone of a national park in the tropical andes’, journal of environmental management 28(2), 238–252. lynagh, f.m. & urich, p.b., 2002, ‘a critical review of buffer zone theory and practice: a philippine case study’, society and natural resources 25, 129–145. http://dx.doi.org/10.1080/089419202753403319 ma, z., li, b., lil, w., han, n., chen, j. & watkinson, a., 2009, ‘conflicts between biodiversity conservation and development in a biosphere reserve’, journal of applied ecology 46, 527–535. http://dx.doi.org/10.1111/j.1365-2664.2008.01528.x maiorano, l., falcucci, a. & boitani, l., 2008, ‘size-dependent resistance of protected areas to land-use change’, proceedings of the royal society b: biological sciences 275(1640), 1297–305. http://dx.doi.org/10.1098/rspb.2007.1756, pmid:18319213 malepe, m., 2008, e-mail, 18 july, email address unknown. mascarenhas, j., 1991, ‘participatory rural appraisal and participatory learning methods: recent experiences from myrada and south india’, rra notes 1991 13, 26–32, viewed 28 march 2012, from http://pubs.iied.org/pdfs/g01399.pdf mavungu, e.m., 2011, ‘explaining boundary disputes in post-apartheid south africa: bushbuckridge, matatiele and khutsong’, phd thesis, department of political studies, university of the witwatersrand, viewed 28 march 2012, from http://wiredspace.wits.ac.za/bitstream/handle/10539/10945/phd%20thesis%20mazembo_final%20version.pdf?sequence=2 mbile, p., vabi, m., meboka, m., okon, d., arrey-mbo, j., nkongho, f. et al., 2006, ‘linking management and livelihood in environmental conservation: case of the korup national park, cameroon’, journal of environmental management 76(1), 1–13. http://dx.doi.org/10.1016/j.jenvman.2005.01.009, pmid:15854732 merafong municipality, 2011, the merafong city integrated development plan 2011/2012. annexure j8: growth and development strategy – section 4, viewed 28 march 2012, from http://www.merafong.gov.za/datafiles/file/idp%202011%2012/annexure%20j%208%20-growth%20&%20development%20strategy.pdf mulder, m.b., caro, t. & msago, o.a., 2007, ‘the role of research in evaluating conservation strategies in tanzania: the case of the katavi-rukwa ecosystem’, conservation biology 21(3), 647–658. http://dx.doi.org/10.1111/j.1523-1739.2007.00703.x, pmid:17531043 reid, h., fig, d., magome, h. & leader-williams, h., 2004, ‘co-management of contractual national parks in south africa: lessons from australia’, conservation and society 2(2), 377– 409. cronje&reilley.qxd natural mortality amoung four common ungulate species on letaba ranch, limpopo province, south africa h.p. cronje, b.k. reilly and i.d. macfadyen cronje, h.p., b.k. reilly and i.d. macfadyen. 2002. natural mortality amoung four common ungulate species on letaba ranch, limpopo province, south africa. koedoe 45(1): 79–86. pretoria. issn 0075-6458. five years of mortality data of impala aepyceros melampus, blue wildebeest connochaetes taurinus, buffalo syncerus caffer and kudu tragelaphus strepsiceros have been used to describe the minimum mortality profiles of the respective species in an open savanna system with the full compliment of predators. predation is the principle cause of natural mortalities on the study site, letaba ranch, which is adjacent to the kruger national park. the principle cause of impala mortality are leopards panthera pardus and wild dogs lycaon pictus. lion panthera leo were major contributors to the mortality of wildebeest and buffalo. anthrax bacillus anthracis was the main cause of kudu mortality. key words: predation, predators, impala, blue wildebeest, buffalo, kudu, letaba ranch. h.p. cronje and b.k. reilly, department of nature conservation, technikon pretoria,private bag x680, pretoria. 0001; i.d. macfadyen, 214 leonie street, doringkloof, pretoria 0001. issn 0075-6458 79 koedoe 45/1 (2002) introduction predation constitutes an important feature of the biotic environments of wild ungulates (hirst 1969) and the effect of this on small private nature reserves can lead to financial implications. the increased perception of the benefits of tourism especially if the major predators are accommodated, has led to several private undertakings applying for permits to accommodate lion in particular. the availability of mortality data, expressed as a financial benefit or loss relative to tourism benefits, is an important factor in determining the sustainability of undertakings. several predator-prey studies have been done to describe the impact of predators on wild ungulate species (mitchell et al. 1965; hirst 1969; pienaar 1969; mills & skenk 1992), but they were one-off studies for short periods of time. the effective management of any wildlife undertaking is dependent on the constant availability of information. mortality registers updated on a weekly and annual basis is a valuable source of information particularly during periods in which game censuses are absent. this data is a means to monitor annual mortalities thus effecting the adjustments of ungulate trends over time. the mortality profiles of impala, blue wildebeest, buffalo and kudu are described. data was collected and collated from field ranger observations for the period january 1990 to december 1994. study area the letaba ranch nature reserve is situated at 23º39's and 31º03'e and 100 km east of tzaneen in the limpopo province, south africa. the kruger national park borders the eastern boundary of letaba ranch. the 35 000 ha reserve is now totally enclosed by an electrified fence which was not the case at the time of the study. the topography of the reserve is predominantly gentle undulating landscapes at altitudes between 340–400 m a.s.l. with the cronje&reilley.qxd 2005/12/09 11:29 page 79 occasional hill reaching 450 m above sea level. letaba ranch is located in a summer rainfall region characterised by hot summers and temperate winters, with the highest and lowest daily mean maximum and minimum temperatures of 31.7 ºc in the summer and 10.1 ºc in the winter. the mean annual rainfall is 487 mm. gertenbach (1987), in his study of the vegetation of the kruger national park, distinguishes two landscapes also common to letaba ranch, namely colophospermum mopane veld on sandy soils extending over the southern two-thirds of the reserve and the letaba river rugged veld in the northern third of the reserve. the larger herbivore component common on letaba ranch are impala, blue wildebeest, buffalo, bushbuck, elephant loxodonta africana, zebra equus burchelli, giraffe giraffa camelopardalis, kudu, sable hippotragus niger, waterbuck kobus ellipsiprymnus, warthog phacochoerus africanus and white rhinoceros ceratotherium simum. lion, leopard panthera pardus, spotted hyaena crocuta crocuta and black-backed jackal canis mesomelas are the most numerous ungulate predators. cheetah acinionyx jubatus, wild dog and side-striped jackal canis adustus are present, but not common. in terms of resource utilisation, letaba ranch functioned as a trophy hunting reserve until 1994 (macfadyen pers. comm.). the rugged and rough terrain, poor roads and scrub mopane all contributed to this utilisation option which had financial benefits higher than that of other eco-tourism ventures (macfadyen pers. comm.). methods to collect a large data set on the game mortalities occurring on letaba ranch, the rangers who patrolled the reserve on a daily basis were tasked with noting mortality data. hirst (1969), pienaar (1969) and mills (1984) also used this data collection technique. to standardise the classification of age classes by the rangers, examples of the various ages of the ungulates under investigation were demonstrated to the rangers. the age classes of kill data recorded were adult male (adm), adult female (adf), sub-adult male (sam), sub-adult female (saf), and juvenile (j). the cause of death was determined by the evidence of characteristic predator kill signs (that is the method by which the animals were killed and eaten), dominant tracks at the site of the kill or the visual confirmation of the predator at a carcass. in the event that the cause of death could not be determined, cause of death was recorded as unknown. the miscellaneous (m) category encompassed infrequently recorded mortalities due to crocodile and caracal. natural mortalities refer to mortalities that were caused by predators or due to intraspecific interactions of a particular species. although malnutrition is the direct cause of mortality during drought periods, the category drought was chosen to represent malnutrition. the main source of bias in the technique used to collect mortality data is fortuitous observations (mitchell et al. 1965; hirst 1969; pienaar 1969; mills 1984) in that for the small prey species and younger age classes, the detection of the carcass was negligible due to the rapid rate at which the carcass was devoured. thus the data is biased to the larger ungulate and age classes. data collection is also biased towards the dry season due to the increased visibility and detection of carcasses during this period. with cognisance of these limitations, it is believed that the data presented provides an estimate of the mortalities that can be expected on a reserve accommodating the full spectrum of predators. the agents of mortality, relative to each ungulate species, were collectively summed with the impact of each agent of mortality expressed as a proportion of the total cause of mortality. results and discussion of the four ungulate species, predation on impala was mostly by leopard and wild dog. lion were responsible for the greater part of deaths among blue wildebeest, buffalo and kudu (table 1). over the spectrum of natural mortalities recorded, in descending order of contribution, lion had the greatest impact (27.5 %) followed by leopard (21.3 %), anthrax (20.5 %) and wild dog with 11.8 % (table 2). koedoe 45/1 (2002) 80 issn 0075-6458 cronje&reilley.qxd 2005/12/09 11:29 page 80 issn 0075-6458 81 koedoe 45/1 (2002) ta bl e 1 n at ur al m or ta lit y to ta ls o f f ou r un gu la te s pe ci es a nd th e pr op or tio na l m or ta lit y of e ac h ag en t o f m or ta lit y on l et ab a r an ch , l im po po p ro vi nc e sp ec ie s n a nt hr ax c he et ah fi gh tin g h ya en a ja ck al l io n l eo pa rd w ild d og d ro ug ht m is ce lla ne ou s u nk ow n % % % % % % % % % % % a ep yc er os m el am pu s 49 1 2, 2 4, 7 1 8, 3 1 7, 3 41 22 6, 7 1, 2 5, 9 c on no ch ae te s ta ur in us 10 1 1 4 73 ,3 11 2 1 2 5, 9 sy nc er us c af fe r 16 2 14 ,2 2, 5 71 ,6 6, 8 5 tr ag el ap hu s s tr ep si ce ro s 25 0 69 1, 2 0, 8 1, 2 20 1, 6 3, 2 0, 4 2, 8 ta bl e 2 c um ul at iv e pr op or tio na l m or ta lit y of e ac h ag en t o f m or ta lit y re la tiv e to th e to ta l nu m be r of m or ta lit ie s re co rd ed , l et ab a r an ch , l im po po p ro vi nc e n a nt hr ax c he et ah fi gh tin g h ya en a ja ck al l io n l eo pa rd w ild d og d ro ug ht m is ce lla ne ou s u nk ow n % % % % % % % % % % % 10 04 20 ,5 2, 7 0, 7 5, 2 0, 5 27 ,5 21 ,3 11 ,8 4, 5 0, 9 4, 5 cronje&reilley.qxd 2005/12/09 11:29 page 81 impala impala were vulnerable to all six predators (table 1). although all predators prey on impala to a greater or lesser extent (skinner & smithers 1990), in this study leopard had the greatest impact (41 %). hirst (1969), pienaar (1969) and le roux & skinner (1989) found that leopard are the main predators of impala in the timbavati (20.7 %), kruger national park (34.3 %) and londolozi game reserve (51 %), respectively. wild dog had the second highest impact (22 %) in contrast to the 9.5 % reported by pienaar (1969) for the kruger national park. this may be attributed to the fact that letaba ranch is smaller in size and fully fenced, causing predation in the herbivore component to be higher. hyaena (8.3 %) and lion (7.3 %) had a smaller impact on impala. cheetah (4.7 %) had the lowest impact of all the predators. cheetah were also reported to be of less importance in the kruger national park, where they contributed a 5.8 % mortality of all carcasses recorded (pienaar 1969). however, in areas where cheetah are in high density and where inter-specific competition is minimal, mortality of impala by cheetah would be expected to be higher. cheetah are regularly subjected to inter-specific competition (pienaar 1969; skinner & smithers 1990). miscellaneous mortalities amounted to 1.2 %, occasioned by five crocodile kills and one caracal kill. anthrax was a minor factor, resulting in mean of 2.2 % over the study period, with 9.32 % deaths in 1990, the year of the outbreak. the percentage mortality of impala per predator within each age class is shown in table 3. adult female impala had the highest mortality relative to other sex and age classes by lion (47.2 %), leopard (39.7 %) and wild dog (49.1 %). for hyaena, adult male and adult female age classes had the same mortality rates. cheetah seemed to concentrate on the male component with a higher mortality rate (43.5 %) within the sub-adult male age class. cheetah generally prey on stragglers on the fringes of the main impala herds (skinner & smithers 1990) which are often of the male gender. jackals caused the highest mortality (80 %) of juvenile impala. newborn lambs of the smaller antelope species are prone to the attention of jackals during the lambing season (pienaar 1969). blue wildebeest blue wildebeest were preyed on by five different predators. the main predators were lion (73.3 %) followed by leopard (11 %) and hyaena (4 %) as indicated in table 1. pienaar (1969), hirst (1969) and mills & skenk (1992) also found that lion were the major cause of wildebeest mortality. the data in table 4 shows a high kill ratio for the adult component, with a slightly higher percentage of females (45.9 %) in relation to the male mortality of 40.5 %, resulting from lion. leopard prey mostly on young wildebeest (bourliére 1963), which is supported koedoe 45/1 (2002) 82 issn 0075-6458 table 3 proportional predation mortality of impala aepyceros melampus relative to age and sex class, letaba ranch, limpopo province age-sex class n cheetah(%) hyaena(%) jackal(%) lion(%) leopard(%) wilddog(%) adult male 143 26,1 41,5 30,6 38,2 30,6 adult female 171 21,7 41.5 47,2 39,7 49,1 sub-adult male 58 43,5 9,8 20,0 13,9 12,1 13,0 sub-adult female 33 8,7 7,3 8,3 9,0 6,5 juvenile 7 80 1,0 0,9 cronje&reilley.qxd 2005/12/09 11:29 page 82 by the results of this study (table 4). juveniles had the highest mortality rate (63.6 %), followed by the sub-adult males (27.3 %) and with the lowest, the sub-adult females (9.5 %). kills by wild dog, hyaena and cheetah were low at 2 %, 4 % and 1 % respectively. buffalo as with blue wildebeest, lion predation is also the principle cause of buffalo mortality (71,6 %), which is borne out by the findings of pienaar (1969) and mitchell et al. (1965) in the kruger and kafue national parks respectively. according to bourliere (1963) adult buffalo regularly fall prey to lion. the adult female and sub-adult male buffalo age classes had the highest representation in lion kills viz. 35.4 % and 25.4 %, respectively (table 5). hyaena had a minimal (2.5 %) effect on the mortality of buffalo (table 1). adult buffalo may be too large a prey to be captured by hyaena, especially if hunted singly (cooper 1990). hyaena mostly select medium-sized prey but being opportunistic, would take almost anything they can overpower (cooper 1990). this is supported by the data in table 5 where it is noted that hyaena tend to concentrate on the younger buffalo component where the opportunity arises. over the study period, anthrax was responsible for 14.2 % of all deaths. during 1990, out of a total of 31 mortalities recorded, anthrax was responsible for 23 (74.2 %) of those mortalities. buffalo are infected with anthrax by grazing on vegetation onto which blowflies have regurgitated after feeding on a carcass contaminated with anthrax as well as being infected with anthrax at waterholes (bothma 1996). the principle source of anthrax infection in buffalo in the kruger national park is waterholes (de vos pers. comm.) and is also assumed to be the case on the letaba ranch game reserve. in 5 % of the sample, the cause of death was issn 0075-6458 83 koedoe 45/1 (2002) table 4 proportional predation mortality of blue wildebeest connocheatus taurinus relative to each age and sex class, letaba ranch, limpopo province age-sex class n cheetah(%) hyaena(%) jackal(%) lion(%) leopard(%) wild dog(%) adult male 32 50,0 40,5 adult female 36 50,0 45,9 sub-adult male 4 0 1,4 27,3 sub-adult female 8 9,5 9,1 juvenile 12 100 2,7 63,6 100 table 5 proportional predation mortality of buffalo syncerus caffer relative to each age and sex class, letaba ranch, limpopo province age-sex class n cheetah(%) hyaena(%) jackal(%) lion(%) leopard(%) wilddog(%) adult male 16 13,8 adult female 41 35,3 sub-adult male 30 25,9 sub-adult female 14 12,1 juvenile 19 100 12,9 cronje&reilley.qxd 2005/12/09 11:29 page 83 unknown. two calving-related deaths (1.2 %) were recorded. kudu kudu were subject to high (20 %) lioninduced mortality as corroborated by hirst (1969), with wild dog second highest (3.2 %) (table 1). in the northern and central districts of the kruger national park, pienaar (1969) found that, amongst others, kudu was one of the species heavily preyed upon by wild dog. during the study, both male and female adult animals had equal mortalities (table 6). leopard, hyaena and cheetah were the lowest contributors to kudu mortality. leopard and cheetah were responsible for the mortality within younger age groups (sub-adult and juvenile) as was also noted by pienaar (1969) in the kruger national park. hyaena preyed on both adult and sub-adult females (table 6). anthrax caused the greatest mortality (69 %) of kudu during the study period. in 1990, of 204 deaths recorded, 171 (84 %) were anthrax related. of all ungulates, kudu are particularly susceptible to anthrax (dekker pers. comm.). kudu are infected in a similar fashion to buffalo (pienaar 1961; braack & de vos 1990). one kudu was taken by a crocodile (0.7 %). annual and seasonal mortality variation the mean annual natural mortalities for impala, blue wildebeest, buffalo and kudu are given in table 8. of particular note, are those of buffalo (31 ± 37.37) and kudu (50 ± 75.38). the cause of the wide data dispersion for kudu was due to the anthrax outbreak during 1990. that of the buffalo is attributed to excessively high lion predation during 1992. the factors contributing to this high mortality were the stress of buffalo after a capture operation, which had an adverse affect on the herd structure, as well as the occurrence of a drought during 1992 (macfadyen pers. comm.). re-computation of the dispersion of the data in the absence of the outliers (trimmed mean) show the mortalities for buffalo and kudu at the 95% confidence koedoe 45/1 (2002) 84 issn 0075-6458 table 6 proportional predation mortality of kudu tragelaphus strepsiceros relative to each age and sex class letaba ranch, limpopo province age-sex class n cheetah(%) hyaena(%) jackal(%) lion(%) leopard(%) wild dog(%) adult male 22 40,0 25,0 adult female 24 66,7 40,0 25,0 sub-adult male 6 12,0 sub-adult female 10 33,3 33,3 8,0 75,0 12,5 juvenile 6 66,7 25,0 37,5 table 7 the seasonal distribution of natural mortalities on letaba ranch, limpopo province species n wet season dry season jan mar/oct dec (%) mar sept (%) aepyceros melampus 491 37,5 62,5 connocheatus taurinus 101 47,5 52,5 syncerus caffer 162 42 58 tragelaphus strepsiceros 250 11,6 88,4 cronje&reilley.qxd 2005/12/09 11:29 page 84 limit to be 15 (± 9.92) and 15.8 (± 8.6) respectively. the majority of all deaths recorded occurred during the dry season for all four ungulates (table 7). conclusion the juvenile components of all four species are underrepresented in the data in this study, attributed to the rapid rate at which these animals are disposed by predators (mitchell et al.1965; hirst 1969). migratory animals have a lower mortality in contrast to sedentary animals in national parks that are large enough to allow for the natural migrations of gregarious ungulates (mills & skenk 1992). from this can be concluded that migratory species in restricted areas will be prone to greater mortality, as is the case of letaba ranch. in restricted areas where predators (especially lion), are accommodated and the smaller the area, the greater the number of mortalities that can be expected. anthrax has the potential to adversely effect ungulate species to a greater or lesser extent and must be taken into account. the capture and removal of game is a management action commonly used to regulate ungulate populations and is a prime income generator. on areas accommodating predators, particularly lions, cognizance must be taken of the potential post capture increase in predation which may lead to further losses. the data in this study represents the minimum number of mortalities for impala, blue wildebeest, buffalo and kudu. it is impossible to record the exact number of mortalities occurring in areas where predators are active due to differing hunting techniques, rates at which prey is consumed, remains left and failure of managers to detect carcasses particularly during the wet season when visibility is impaired. mortality data should be used in the development of population models that incorporate game count data to derive more accurate data on population trends. annual game censuses are becoming more irregular due to financial constraints. a population model based on population trends (from long-term species monitoring) and incorporating annual mortality data (natural and unnatural) providing estimates of populations during intervening periods, should be a research priority. models of this nature would ensure continuous data regarding ungulate trends. count data combined with population dynamics data provide better decision support than count driven trends alone. predator prey models also require accurate data on mortalities, particularly where predators are to be reintroduced to conservation areas in the sub-region. very often wildlife managers look to expensive and sophisticated procedures to gather data on game populations. this analysis from daily summaries in a managers diary, from good ground coverage by field rangers, show that “primitive” methods are invaluable in gaining insight into the status of species in protected areas. in the light of issn 0075-6458 85 koedoe 45/1 (2002) table 8 annual variation in the natural mortalities of four ungulate species on letaba ranch, limpopo province species n mean anuual mortality aepyceros melampus 491 98,2 (± 44,12) connocheatus taurinus 101 20,2 (± 5,2) syncerus caffer 162 31 (± 37,37) tragelaphus strepsiceros 250 50,2 (± 75,38) cronje&reilley.qxd 2005/12/09 11:29 page 85 increasing costs of monitoring, declining budgets and expertise in conservation agencies, such inexpensive opportunistic data gathering must be considered in wildlife management undertakings in southern africa. the effectiveness of this approach is, however, dependent on the number of field staff employed for this purpose relative to the surface area that will require regular surveillance. ascertaining the competence of field staff in accurately recording mortality and training them where necessary is of paramount importance for the collection of mortality data. acknowledgements the first author would like to thank ionka du preez for her contiuous support and assitance throughout the preparation of this manuscript. gratitiude is also expressed to the referees that provided valuble input and added value to the paper. references bourliére, f. 1963. specific feeding habits of african carnivores. african wildlife 17(1): 21–27. braak, l.e.o. & v. de vos. 1990. feeding habits and flight range of blow-flies (chrysomyia spp) in relation to anthrax transmission in the kruger national park, south africa. onderstepoort journal of veterinary research 57: 141–142. choquette, l.p.e. & e. broughton. 1981. anthrax pp. 289–295. in: davies, j.w., l.h. karstad & d.o. trainer (ed). infectious diseases of wild mammals. ames, iowa: iowa state university press. cooper, m.s. 1990. the hunting behaviour of spotted hyaenas (crocuta crocuta) in a region containing both sedentary and migratory populations of herbivores. african journal of ecology 28: 131–141. gertenbach, w.p.d. 1987. 'n ekologiese studie van die suidelikste mopanieveld in die nationale krugerwildtuin. d.sc. verhandeling, universiteit van pretoria, pretoria. hirst, s.m. 1969. populations in a transvaal lowveld nature reserve. zoologica africana 4(2): 199–230. le roux, p.g. & j.d. skinner. 1989. a note on the ecology of the leopard (panthera pardus linnaeus) in the londolozi game reserve, south africa. african journal of ecology 27: 167–171. mills, m.g.l. 1984. prey selection and feeding habits of the large carnivores in the southern kalahari. koedoe (suppl.): 281–296. mitchell, b.l., j.b. shenton & j.c.m. uys. 1965. predation on large mammals in the kafue national park, zambia. zoologica africana 1: 297–318. mills, m.g.l. & t.m. skenk. 1992. predator-prey relationships: the impact of lion predation on wildebeest and zebra populations. journal of animal ecology 61: 693–702. pienaar, u. de v. 1961. the outbreak of anthrax amongst game animals in the kruger national park. koedoe 4: 4–17. pienaar, u. de v. 1969. predator-prey relationships amongst the larger mammals of the kruger national park. koedoe 12: 108–187. skinner, j.d. & r.h.n. smithers. 1990. the mammals of the southern african subregion. pretoria: university of pretoria. koedoe 45/1 (2002) 86 issn 0075-6458 cronje&reilley.qxd 2005/12/09 11:29 page 86 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <feff00560065007200770065006e00640065006e0020005300690065002000640069006500730065002000450069006e007300740065006c006c0075006e00670065006e0020007a0075006d002000450072007300740065006c006c0065006e00200076006f006e0020005000440046002d0044006f006b0075006d0065006e00740065006e0020006d00690074002000650069006e006500720020006800f60068006500720065006e002000420069006c0064006100750066006c00f600730075006e0067002c00200075006d002000650069006e00650020007100750061006c00690074006100740069007600200068006f006300680077006500720074006900670065002000410075007300670061006200650020006600fc0072002000640069006500200044007200750063006b0076006f0072007300740075006600650020007a0075002000650072007a00690065006c0065006e002e00200044006900650020005000440046002d0044006f006b0075006d0065006e007400650020006b00f6006e006e0065006e0020006d006900740020004100630072006f0062006100740020006f0064006500720020006d00690074002000640065006d002000520065006100640065007200200035002e003000200075006e00640020006800f600680065007200200067006500f600660066006e00650074002000770065007200640065006e002e00200042006500690020006400690065007300650072002000450069006e007300740065006c006c0075006e00670020006900730074002000650069006e00650020005300630068007200690066007400650069006e00620065007400740075006e00670020006500720066006f0072006400650072006c006900630068002e> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice article information authors: tineke kraaij1,2 richard m. cowling2 brian w. van wilgen3,4 affiliations: 1south african national parks, garden route scientific services, south africa2department of botany, nelson mandela metropolitan university, south africa 3csir natural resources and the environment, stellenbosch, south africa 4centre for invasion biology, department of botany and zoology, stellenbosch university, south africa correspondence to: tineke kraaij postal address: po box 176, sedgefield 6573, south africa dates: received: 25 july 2012 accepted: 10 dec. 2012 published: 15 mar. 2013 republished: 02 apr. 2013 note: this article was republished with the correct affiliation for brian w. van wilgen. how to cite this article: kraaij, t., cowling, r.m. & van wilgen, b.w., 2013, ‘fire regimes in eastern coastal fynbos: imperatives and thresholds in managing for diversity’, koedoe 55(1), art. #1104, 9 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1104 copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. fire regimes in eastern coastal fynbos: imperatives and thresholds in managing for diversity in this review article... open access • abstract • introduction    • study area • fynbos fire regimes across an east-west transect • managing fire for biodiversity conservation: ecological thresholds    • fire return interval    • fire season • fire management approach: operational considerations    • fire risk and safety (with emphasis on protecting neighbouring plantations)    • fire and management of invasive alien plants • future research and monitoring needs • concluding recommendations • acknowledgements    • competing interests    • author' contributions • references abstract top ↑ until recently, fire ecology was poorly understood in the eastern coastal region of the cape floral kingdom (cfk), south africa. rainfall in the area is aseasonal and temperatures are milder than in the winter-rainfall and drier inland parts of the cfk, with implications for the management of fire regimes. we synthesised the findings of a research programme focused on informing ecologically sound management of fire in eastern coastal fynbos shrublands and explored potential east–west trends at the scales of study area and cfk in terms of fire return interval (fri) and fire season. fris (8–26 years; 1980–2010) were comparable to those elsewhere in the cfk and appeared to be shorter in the eastern tsitsikamma than in the western outeniqua halves of the study area. proteaceae juvenile periods (4–9 years) and post-fire recruitment success suggested that for biodiversity conservation purposes, fris should be ≥ 9 years in eastern coastal fynbos. collectively, findings on the seasonality of actual fires and the seasonality of fire danger weather, lightning and post-fire proteoid recruitment suggested that fires in eastern coastal fynbos are not limited to any particular season. we articulated these findings into ecological thresholds pertaining to the different elements of the fire regime in eastern coastal fynbos, to guide adaptive management of fire in the garden route national park and elsewhere in the region.conservation implications: wildfires are likely to remain dominant in eastern coastal fynbos, whilst large-scale implementation of prescribed burning is unattainable. fires occurring in any season are not a reason for concern, although other constraints remain: the need for sufficient fire intensity, safety requirements, and integration of fire and invasive alien plant management. introduction top ↑ fire is instrumental in maintaining the species-rich and endemic-rich fynbos shrublands (myers et al. 2000) of the cape floral kingdom (cfk) of south africa (kruger & bigalke 1984). fire is the dominant natural disturbance in fynbos and managing fynbos equates to managing fire. the fire ecology of fynbos has been well researched in the western, winter-rainfall part of the cfk (kruger & bigalke 1984; van wilgen & viviers 1985) and in the drier inland mountains (bond, vlok & viviers 1984; midgley 1989; seydack, bekker & marshall 2007) but was, until recently, poorly understood in the eastern coastal part of the cfk. the climate of the latter is milder with reduced seasonal extremes in temperature, whilst rainfall occurs year-round. as shown by heelemann et al. (2008) in the inland fynbos of the eastern cfk, overstorey proteoids (non-sprouting, slow-maturing, serotinous proteaceae) show weak seasonal effects on post-fire recruitment, a consequence of a non-seasonal rainfall and fire regime. we expected similar patterns in our region, as well as faster plant growth and maturation rates of proteoids, a consequence of the region’s relatively benign climate. these phenomena will have important implications for the management of fire regimes.the garden route national park (grnp; c. 130 000 ha) was recently established in the eastern coastal part of the cfk in a landscape mosaic of indigenous forests, fire-prone fynbos shrublands and fire-sensitive plantations of invasive alien trees. the park faces considerable ecological and operational challenges pertaining to the management of fire and invasive alien plants (kraaij, cowling & van wilgen 2011). a recent, multifaceted research programme was undertaken to inform ecologically sound management of fire in eastern coastal fynbos and entailed, (1) an assessment of the context within which fire management was practiced during the past century (kraaij et al. 2011), (2) characterisation of the recent fire history and fire regime (1900–2010) in the area (kraaij, baard, cowling, van wilgen & das 2012), (3) characterisation of the seasonality of fire weather and lightning (kraaij, cowling & van wilgen 2012), (4) estimation of minimum fire return intervals (fris) from juvenile periods and post-fire recruitment success of proteoids (kraaij et al. 2013) and (5) determination of the ecologically appropriate fire season from post-fire recruitment seasonality of proteoids (kraaij 2012). here we synthesise the findings of this research in terms of potential east–west trends within eastern coastal fynbos and at the scale of the entire cfk. furthermore, we articulate the findings into ecological thresholds (biggs & rogers 2003; van wilgen et al. 2011) pertaining to the different elements of the fire regime in eastern coastal fynbos, to guide adaptive management of fire in the park and elsewhere in the region. study area the study area comprises the coastal slopes of the outeniqua mountains (east of the touw river) and tsitsikamma mountains (33°48′s 22°35′e – 34°01′s 24°16′e), of which a large portion occurs within the grnp (kraaij et al. 2011). owing to maritime influence, the climate of the area is relatively equable (schulze 1997). rainfall occurs throughout the year, with winter months being the driest. mean annual rainfall increases eastwards, from 820 mm to 1078 mm in the outeniqua and tsitsikamma mountains, respectively. the proportion of summer rain also increases eastwards (schulze 1997). the fire-prone and fire-dependent vegetation of the study area largely comprises south outeniqua sandstone fynbos and tsitsikamma sandstone fynbos in the outeniqua and tsitsikamma mountains, respectively (rebelo et al. 2006). these are tall, medium-dense proteoid shrublands, with an ericoid-leaved shrub understorey (dominated by ericaceae) and a prominent restioid (restionaceae) component. more detailed accounts of the climate, vegetation and fire management history of the study area can be found elsewhere (geldenhuys 1994; kraaij et al. 2011, 2013; kraaij, cowling & van wilgen 2012). fynbos fire regimes across an east–west transect top ↑ kraaij et al. (2011) established that historically, plantation protection enjoyed priority over fynbos conservation, influenced by the large extent of plantations in the study area. fynbos close to plantations has probably been compromised by short-rotation (4–8 years at times) and low-intensity burning in the past, as well as by invasion by alien trees. in terms of area burnt (1900–2010), natural (lightning-ignited) fires dominated the fire regime (59%), particularly in the tsitsikamma (83%), whereas prescribed burning was relatively unimportant (kraaij, baard, cowling, van wilgen & das 2012). fris since 1980 appeared to be shorter in the east of the study area (tsitsikamma region, median fri = 8.3 years, when only complete fire intervals were considered), where rainfall and plant growth rates are higher, than in the west (outeniqua region, median fri = 13.2 years) (kraaij, baard, cowling, van wilgen & das 2012). however, fris for the study area as a whole (8–26 years, considering complete and open-ended fire intervals during the period 1980–2010) were broadly comparable to those of other regions in the cfk (15–55 years – seydack et al. 2007; 10–21 years – van wilgen et al. 2010), suggesting that variation is related to local or regional moisture regimes (kruger & bigalke 1984; seydack et al. 2007) rather than an east–west gradient of increasing fris across the cfk. this is consistent with the lack of a gradient in mean annual rainfall from east to west within the cfk (deacon, jury & ellis 1992). proteaceae juvenile periods (4–9 years) and post-fire recruitment success (following fires in 7-year-old vegetation) suggested that for biodiversity conservation purposes, fris should be no less than 9 years in moist, productive fynbos (kraaij et al. 2013). minimum fris to ensure pre-fire maturation and thus successful post-fire recruitment of proteoids in the study area (kraaij et al. 2013) were similar to those from other parts of the cfk (van wilgen et al. 2011).seasonality in fire occurrence was less pronounced in the east than in the west of the study area, with fires less concentrated in summer in the tsitsikamma (13% of area burnt) than in the outeniqua region (31%) (kraaij, baard, cowling, van wilgen & das 2012). in the study area as a whole, fire occurrence was less seasonal (c. 40% of the total area burnt did so during summer–autumn) than in western (c. 75%) and inland (c. 70%) (seydack et al. 2007; van wilgen et al. 2010) parts of the cfk, consistent with climatic gradients of increasing winter rainfall and summer drought from east to west and increasing summer evapotranspiration from the coast to the interior. the trend of increasing seasonality in fire occurrence and rainfall from east to west was not reflected in the seasonality of lightning occurrence at either the scale of study area or cfk (kraaij, cowling & van wilgen. 2012). the east–west trend of increasing seasonality in fire occurrence was, however, reflected in the biotic post-fire recruitment response. fire season had little consistent effect on post-fire recruitment of proteoids in the study area (kraaij 2012), in contrast to the strong seasonal response in western (midgley 1989; van wilgen & viviers 1985) and inland (bond 1984; bond et al. 1984) parts of the cfk, where summer and autumn fires result in optimal recruitment. collectively, findings in the study area on the seasonality of actual fires (kraaij, baard, cowling, van wilgen & das 2012) and the seasonality of fire danger weather, lightning (kraaij, cowling & van wilgen 2012), as well as post-fire proteoid recruitment (kraaij 2012) suggest that fires in eastern coastal fynbos are not limited to any particular season, and therefore management is not constrained, although the ecological requirements for adequate fire intensity (van wilgen et al. 2011) remain. significant increases in the total area burnt annually since 1980 (kraaij, baard, cowling, van wilgen & das 2012) that were correlated with significant long-term increases in weather conditions conducive to the spread of fires (kraaij, cowling & van wilgen 2012), suggest fire regime changes related to global change. increases in fire frequency or extent (forsyth & van wilgen 2008) and in the severity of fire danger weather conditions (wilson et al. 2010) have similarly been observed elsewhere in the cfk, although evidence is not unequivocal across all meteorological variables (e.g. relative humidity, wind run) affecting fire danger weather (hoffman et al. 2011; kraaij, cowling & van wilgen 2012). managing fire for biodiversity conservation: ecological thresholds top ↑ fire management practices aimed at biodiversity conservation within protected areas are often in conflict with hazard reduction requirements (kraaij et al. 2011) and, where such conflict of interest exists, managers need to know the ecological thresholds within which fire may be managed. adaptive management is based on formulating targets (or thresholds of potential concern) (biggs & rogers 2003) describing acceptable variations in the elements (frequency, season, intensity, size) of fire regimes, with monitoring regularly assessing ‘whether the fire regimes that arise from various forms of management remain within the specified ranges’ (van wilgen et al. 2011). we have set provisional thresholds for eastern coastal fynbos (table 1) and particularly in terms of two elements of the fire regime, namely fire frequency (the lower threshold for fri) (kraaij et al. 2013) and fire season (kraaij 2012). table 1: proposed thresholds of potential concern relating to fire management of fynbos (c. 105 000 ha) in the garden route national park. thresholds pertaining to fire patterns are informed by monitoring against thresholds related to ecosystem responses to fires. the adoption of thresholds in terms of elements of the fire regime may, at first glance, appear to imply that the fire patterns themselves are the desired outcome. however, fire patterns will ultimately manifest themselves in vegetation structure, function and composition, which are assessed in terms of thresholds related to the responses of ecosystems to fire (van wilgen et al. 2011). we distinguished between thresholds pertaining to fire patterns (termed ‘operational thresholds’ by van wilgen et al. 2011) and those pertaining to ecosystem responses to fires (termed ‘ecosystem thresholds’ by van wilgen et al. 2011) (table 1), the ultimate goal being to make the links between fire patterns and ecological outcomes. fire return interval estimates of optimal fire frequencies are often based on the relationship between plant age and the rate of seed accumulation of the slowest-maturing species. in fynbos and in australian shrublands, juvenile periods of proteoids are used as indicators of extreme lower limits for fris, whereas their longevity indicates extreme upper limits (bond 1980; gill & mccarthy 1998). the rule of thumb generally used in fynbos to establish a lower threshold for fri posits that half of the individuals in a population of the slowest-maturing proteoid species should have flowered in at least three successive seasons before the area may be burnt (kruger & lamb 1978; kruger 1982; van wilgen et al. 2011) (table 1). recruitment success of proteoids after fires at different intervals may also indicate the ecological suitability of fris. application of the mentioned rule of thumb to juvenile periods observed in the study area implied a minimum fri of 9 years for moist, productive tsitsikamma fynbos, whereas post-fire recruitment of proteoids was always successful following fris of 7 years (kraaij et al. 2013). considerable variation in juvenile periods, flowering status at given plant ages and post-fire recruitment success at given fris, within and between species and habitats (kraaij et al. 2013), suggests that management should not attempt to rigidly and consistently pursue a fixed or narrow range of fris. besides it being practically unattainable (van wilgen et al. 2010), variability in fris in nature is inevitable and, where fire management is aimed at conserving biodiversity, variation within established limits is desirable (gill & mccarthy 1998; thuiller et al. 2007). variation in fri (or fire season or fire intensity) will induce variation in the density of overstorey proteoids, which, in turn, is associated with maintenance of diversity in understorey species (cowling & gxaba 1990; vlok & yeaton 1999). some of the variation in flowering status appeared to be related to local moisture regimes with fris being inversely related to rainfall or moisture availability. due cognisance should therefore be taken of site-specific requirements, with drier habitats (northerly or westerly aspects or shallow soils) generally needing longer intervals between fires. range-restricted species or habitat specialists may furthermore have very specific fri requirements (kruger & bigalke 1984). slow-growing, high-altitude species, for example protea grandiceps (near threatened) (raimondo et al. 2009), may be slow to mature (> 10 years) (j.h.j. vlok pers. comm., 04 february 2012) and thus require longer fris. although their primary aim was to establish a lower threshold for fri, the results of kraaij et al. (2013) also showed that the upper threshold for fri is likely to be in excess of 38 years. they concurred with van wilgen et al. (2011) that the occurrence of very old vegetation and thus the setting of an upper threshold for fri is not a key concern in the ecological management of fire in montane proteoid fynbos, both because old vegetation is typically limited in extent (bond 1980; kraaij, baard, cowling, van wilgen & das 2012) and because proteoid recruitment does not appear to be negatively affected by relatively long inter-fire periods (kraaij et al. 2013). an assessment of the historical (1980–2010) occurrence of fire in the study area found that the proposed fri thresholds were exceeded over a greater proportion of the area burnt in the tsitsikamma than in the outeniqua region (kraaij, baard, cowling, van wilgen & das 2012). since 1980, 11% of the fynbos of the grnp burnt at least once at post-fire ages of < 7 years. in these areas, proteoids have likely been locally eliminated, as post-fire recruitment success of this guild has been near zero following a 5-year fri (kraaij et al. 2013). measuring against a minimum fri threshold of 9 years resulted in much greater area exceeding this threshold, with 27% of the study area (31% of tsitsikamma and 15% of outeniqua) having burnt at least once at post-fire ages of < 9 years. in these areas, seed banks of proteoids have likely been insufficient at the time of fire to have enabled optimal post-fire recruitment. the extent to which proteoid populations may have been compromised in these areas depends on the particular species present, local moisture regimes and plant maturation rates (kraaij et al. 2013). identification of site and vegetation characteristics and weather conditions before and during short interval fires could provide valuable insight into whether an increase in the occurrence of such burns may be expected. short interval fires, which alter vegetation structure (from woody to herbaceous) (kruger 1984; lloret, pausas & vilà 2003) and thus fuel dynamics, may set up negative feedback loops whereby short fris persist in the landscape (richardson & van wilgen 1992). similar effects on fire regimes have occurred after invasion by alien grasses of californian chaparral (keeley & brennan 2012). in large parts of the study area, graminoid sprouters dominate, whilst proteoids are sparse or absent (heelemann et al. 2010), notably in areas near plantations of alien pine trees (t. kraaij, pers. obs.). this likely resulted from frequent and low intensity burning in the past, aimed at protecting timber plantations from fire (kraaij et al. 2011). facilitation of fris that would ensure the long-term persistence of proteoids in the grnp is thus a priority for fynbos conservation management, particularly where this guild has persisted in the landscape. it is therefore important to map the distribution of species that are vulnerable to too frequent fires and to incorporate this information into the fire management plan for the area. thresholds may also be set for post-fire age class distributions of the vegetation (table 1). relatively equal age class distributions are desirable as these would provide variation in vegetation age and structure, and thus a variety of habitat types for a diversity of species, in addition to spreading out the work load over time for prescribed burning and invasive plant control (van wilgen et al. 2011). the distribution of post-fire vegetation age classes in the grnp (measured in 2011) was highly skewed towards the younger age classes (kraaij, baard, cowling, van wilgen & das 2012), thereby far exceeding proposed thresholds (table 1) and upsetting invasive alien plant clearing schedules. to correct this situation, increased attempts at protection of grnp fynbos from extensive burning would be desirable for the next few year. fire season season of fire has marked effects on floristic composition in fire-prone mediterranean-climate shrublands (bond 1984; enright & lamont 1989; midgley 1989). in these winter-rainfall systems, summer–autumn fires lead to optimal recruitment of proteoids, which are key to the conservation of floral diversity overall. kraaij (2012) established that seasonal patterns in post-fire recruitment of proteoids are weak in eastern coastal fynbos (good recruitment may occur after disturbance in any season), in line with the findings that fire danger weather (kraaij, cowling & van wilgen. 2012) and fire occurrence (kraaij, baard, cowling, van wilgen & das 2012) in this area are also largely aseasonal. this implies a lack of any seasonal constraint on burning, which has encouraging management implications. managers will not have to allocate large amounts of resources to fight wildfires that are burning in what are considered ecologically undesirable seasons based on patterns from the western and inland fynbos regions; they may also conduct back burns during any season. instead of the thresholds applicable to fire seasonality in western and inland fynbos (i.e. that > 80% of area should burn during summer–autumn and < 20% during winter–spring) (van wilgen et al. 2011), variability in fire season is considered desirable in eastern coastal fynbos. the thresholds we propose for the study area accordingly relate to a reasonably even distribution of fires amongst seasons (i.e. the proportion of the area burnt in any particular season should be > 10% and < 40%) (table 1).in aseasonal environments such as the study area and southeastern australia, there is large variation between years in the seasonality of rainfall and an unpredictable response of proteoid recruitment to season of fire. the effect of fire season on proteoids may, in the long term, be no more important than that of the timing of fire in relation to occurrence of wet and dry cycles (bradstock & bedward 1992). particularly in these unpredictable, aseasonal environments, variation in fire regimes (including fire season) is necessary to maintain plant diversity in space and time (gill & mccarthy 1998; thuiller et al. 2007). the range of acceptable variation in fire seasonality that we propose is accordingly wide. although removal of the seasonal constraint should expand the window of opportunity and thus improve the chances of implementing prescribed burns, the ecological need for adequate fire intensity remains (table 1) (van wilgen et al. 2011). fire intensity needs to be sufficiently high to stimulate seed release and germination in serotinous (midgley & viviers 1990) and large or hard-coated, soil-stored seeds (bond et al. 1990; jefferey, holmes & rebelo 1988; knox & clarke 2006), but not excessive such that species with small seeds and shallow seed banks are killed (bond, honig & maze 1999). for instance, a few severely fragmented subpopulations (mostly with < 50 individuals each) of the large-seeded, myrmecochorous mimetes splendidus (endangered) (raimondo et al. 2009) occur within the study area and at least one subpopulation (at buffelsnek) is known to have been lost as a result of low intensity fires (j.h.j. vlok pers. comm., 04 february 2012). furthermore, ecological requirements of fire intensity are often in conflict with safety considerations (morrison, buckney & bewick 1996; van wilgen et al. 2012), presenting considerable management challenges. an assessment of the historical occurrence of fire in the study area found that fire season thresholds were not exceeded when considering the period from 1900–2010 as a whole (kraaij, baard, cowling, van wilgen & das 2012). the proportional areas burnt were fairly evenly distributed amongst seasons (winter 23%, spring 35%, summer 19%, autumn 22%). neither were the thresholds exceeded when assessed over the past 15 years (1996–2010) (cf. van wilgen et al. 2011), when the seasonal distribution of fires was also relatively even (winter 20%, spring 35%, summer 19%, autumn 30%). despite the occurrence of several very large (10 000 ha) fires since 1990, fire seasonality remained within the proposed thresholds, suggesting that inappropriate season of fire generally does not jeopardise ecologically sound management of fynbos in the area. fire management approach: operational considerations top ↑ approaches to fire management in fynbos are mainly distinguished by the extent of intervention pursued, ranging from no manipulation (‘natural fire zone management’) through selective manipulation (‘adaptive interference management’) to (attempted) complete control by means of block burning (seydack 1992; van wilgen et al. 1994). the perspective of this study is that of protected area management, with specific reference to the grnp. the key fire management issues presently faced by the grnp are, (1) to reconcile ecological and fire hazard reduction requirements (cf. gill & mccarthy 1998; morrison et al. 1996; van wilgen et al. 2012), especially in the context of neighbouring commercial plantations and (2) to align invasive alien plant clearing initiatives with fire management strategies to optimise biodiversity conservation and ecosystem service delivery (kraaij et al. 2011). here we explore the options for addressing these two main issues within the established ecological constraints. fire risk and safety (with emphasis on protecting neighbouring plantations) plantation managers are placing pressure on grnp managers to adopt a policy of low-intensity burns at short rotations (4–8 years) in fynbos adjacent to plantations to minimise wildfire hazards and damage (kraaij et al. 2011). substantial legal claims for damages led to a fire management agreement between the main plantation holding company, cape pine (formerly mountain to ocean forestry) and the grnp for the tsitsikamma region (vermeulen et al. 2009). accordingly, fuel reduction burning is to be undertaken in designated fynbos blocks (45 ha – 1178 ha in size) located to the north of plantations where indigenous forest does not form a natural buffer between fynbos and plantations (kraaij et al. 2011). the blocks are strategically placed in relation to fire corridors associated with wildfires that burn under hot, dry, bergwind conditions (geldenhuys 1994) and form two parallel fire belts to be burnt on an 8-year rotation, such that the vegetation in one of the belts would always be ≤ 4 years of post-fire age. the block burn system is to be further supported by strategically placed fire breaks, as well as controlled fuel reduction burning within plantations, particularly along their boundaries (de ronde et al. 1990).however, implementation of this system to date has failed for a number of reasons. the rugged terrain, lack of access roads and natural firebreaks, and dense infestations of invasive alien plants (originating from the plantations), make block burning prohibitively expensive and risky. to date, ecological restrictions on fire season (inherited from winter-rainfall fynbos) combined with fire danger weather restrictions reduced the window of opportunity for prescribed burning to only a few days per year (van wilgen & richardson 1985). furthermore, fire legislation – the national veld and forest fire act (act no. 101 of 1998) (republic of south africa 1998) – acts as a strong deterrent to taking risks with prescribed burning, despite it recognising ‘both the ecological role of fire for maintaining healthy ecosystems and the need to reduce the risks posed by fires’ (van wilgen et al. 2012). even if it were possible to implement prescribed burning according to plan (which may be somewhat more plausible with the relaxation of seasonal constraints) (kraaij 2012), land managers need to realise that fynbos and plantations cannot be fire-proofed (cf. agee 2002). assessments of historical fire regimes in the study area (kraaij, baard, cowling, van wilgen & das 2012) and elsewhere (keeley, fotheringham & morais 1999; moritz 2003; van wilgen et al. 2010) show that rigid control over fire regimes is largely unattainable. globally, low levels of fuel-age dependency in fire-prone shrublands and forests imply that fuel reduction burning may not necessarily prevent the occurrence of runaway wildfires (bessie & johnson 1995; keeley et al. 1999; moritz 2003; van wilgen et al. 2010). the most effective strategy for facilitating fire safety where necessary is to focus effort on strategic locations, bearing in mind that fuel breaks are largely effective at facilitating fire management by providing access for fire fighting activities (syphard, keeley & brennan 2011). given the mentioned physical, ecological, economical and political constraints, wide-scale implementation of prescribed burning is thus likely to remain unattainable (keeley et al. 1999; kraaij et al. 2011; van wilgen 2009) and ineffective at preventing destructive wildfires (keeley & zedler 2009). instead of extensive block burning of fynbos in the grnp, we recommend a system of largely natural fires and minimum interference, but allowing pro-active fire management measures where or when required (‘adaptive interference management’) (seydack 1992). the main focus should be on establishing a network of fire breaks where fynbos abuts plantations or other fire-sensitive land uses. in line with the stipulations of the national veld and forest fire act (act no. 101 of 1998) (republic of south africa 1998), the location, specifications (clearing method, width and maintenance schedule) and maintenance responsibilities pertaining to fire breaks should be formally agreed upon by neighbouring land managing agencies within the institutional framework of the regional (southern cape) fire protection association. in principle, these fire breaks should provide access and relatively safe conditions for fire fighting along boundaries in areas of high fire hazard. fire breaks have to be practically aligned with topographical features and need to avoid sensitive habitats, such as mountain ridges, rather than strictly follow property boundaries. the location and specifications of fire breaks (or other pro-active fire management interventions) will be dynamic in space and time, tracking changes in the vegetation age mosaic and fire risk distribution. regular assessments may be required to identify: • areas where fires should not burn. these include immature fynbos (< 9 years post-fire) (table 1), areas of disturbed or transformed forest where forest recovery should be facilitated (watson & cameron 2001) and where damage to assets or runaway fires may be anticipated. such areas should receive priority for intervention during wildfires. • areas where fires may be allowed to burn unhindered. these would mainly be inaccessible areas where no immediate threat to assets exists. • areas where fires should preferably burn within the next 5 years. these include senescent fynbos (> 40 years old) (table 1) that requires a burn for rejuvenation, areas where excessive fuel build-up poses a fire hazard to human lives or assets, and disturbed areas (primarily decommissioned plantations) (kraaij et al. 2011) where fire may facilitate the recovery of fynbos (holmes & richardson 1999). in these areas prescribed burning should be considered. adoption of the outlined approach should afford sufficient flexibility to integrate fire management with the dynamic plantation decommissioning and rehabilitation process (kraaij et al. 2011) and with invasive alien plant management (roura-pascual et al. 2009; van wilgen 2009). other aspects to fire management are preparedness and response or suppression capacity (van wilgen et al. 2012). in areas where fires occur seasonally, (high) fire danger season(s) are typically recognised and fire-fighting capacity augmented during this time. fire managers conventionally regard the fire danger season in the study area to be during autumn–winter (april–october) when rainfall is relatively low and bergwinds often prevail (geldenhuys 1994; kraaij et al. 2011; le roux 1969). an analysis of the seasonality of fire-prone weather conditions confirmed that fire danger weather peaks in winter (kraaij, cowling & van wilgen 2012); although, average conditions are mild year-round compared to those characteristic of fire danger seasons elsewhere in the cfk (van wilgen 1984). furthermore, the historical occurrence of both lightning-ignited and human-ignited fires in the study area does not attest to a winter fire regime (kraaij, baard, cowling, van wilgen & das 2012) and large fires may occur at any time of the year, even under low or moderate fire danger conditions (kraaij, cowling & van wilgen 2012). lightning, as a source of ignition, also occurs throughout the year (kraaij, cowling & van wilgen 2012). collectively, these findings suggest that there is no clear fire danger season in the study area that would demand increased preparedness or fire fighting capacity compared to other times of the year. fire and management of invasive alien plants the first state plantations in the study area were established in 1883 (kraaij et al. 2011) and the two most widely planted species (pinus pinaster and pinus radiata) (van wilgen & richardson 2012) have become the most problematic invaders in their genus (classed as ‘transformers’) richardson et al. 2000) in the study area and elsewhere in the fynbos (richardson & higgins 1998). key to the invasive success of serotinous pines in fynbos is their response to fire, which is similar to that of proteoids (showing marked spatial and temporal post-fire fluctuations in population sizes). however, pines outperform proteoids in two key facets of demography, that is, seed dispersal and fire-resilience (richardson & higgins 1998). seeds and pollen of pines are exceptionally well dispersed by wind, whilst isolated pioneers can give rise to colonies by selfing (richardson & higgins 1998). furthermore, pines have superior fire-resilience with comparatively short juvenile periods (c. 5 and 6 years in p. radiata and p. pinaster, respectively) (richardson, cowling & le maitre 1990) and better adult fire-survival ability than most proteoids. once pines are established, the fynbos may be transformed into a pine forest over two to three fire cycles (richardson & brown 1986). because the life cycles of serotinous pines (and hakeas) are closely linked to fire, integration of fire and alien plant management strategies is essential (roura-pascual et al. 2009).the conventional methods of managing invasive pines are slashing or hand-pulling of immature saplings, and ring-barking or felling of large mature trees, allowing time for seeds to be released and subsequent burning of new recruits prior to their reaching maturity (holmes & marais 2000; holmes et al. 2000). equally, where prescribed burning of fynbos is intended, invader plants require pre-fire treatment. in some cases, options exist for managing invasions by manipulating disturbance regimes like fire cycles (e.g. esler et al. 2010 for hakea; richardson & higgins 1998 for pinus). unfortunately, potential use of short-rotation fires to control pines is precluded by the fact that juvenile periods of native proteoids in the study area (4–9 years) (kraaij et al. 2013) are mostly longer than those of pines (see above). however, in fynbos devoid of proteoids (naturally, or as a result of inappropriate past management of fire) and mature pines, application of a single fri of < 5 years may, in some instances, provide an inexpensive means of substantially reducing dense infestations of young pine recruits. otherwise, it would generally be in the interest of invasive plant control to prolong fris in order to curb the rate of spread of fire-propagated species such as pines and hakeas (richardson & van wilgen 1992). future research and monitoring needs top ↑ recent research reported here improved knowledge of the historical fire regime in the study area and how it has changed during the past century. accurate spatial recording of future fires is important to supply baseline data to inform the design of natural experiments, fire management decisions and planning (driscoll et al. 2010) and to recognise potential long-term changes in fire regimes (kraaij, baard, cowling, van wilgen & das 2012). ongoing monitoring and regular evaluation of fire regime outcomes in terms of agreed thresholds are essential components of the adaptive management cycle, to enable learning and corrective action where necessary. in cases where thresholds are exceeded, consideration needs to be given to management interventions that will drive the system to be within thresholds, or, alternatively, thresholds may be recalibrated where appropriate. results from the monitoring of ecosystem responses to fires (ecosystem thresholds in table 1) should inform the calibration of fire pattern thresholds. the thresholds we proposed for minimum fri and fire season are first approximations within which fire managers may attempt to resolve the conflicting demands of fire hazard reduction and biodiversity conservation in eastern coastal fynbos. further research on maturation rates of slow-maturing reseeding plant species and success of post-fire vegetation recovery (morrison et al. 1995) in diverse habitats would be required to refine these thresholds. substantial variation, both in flowering status and post-fire recruitment, as well as disparity amongst estimates of minimum fri based on these measures (kraaij et al. 2013), emphasise the need to empirically verify kruger’s (1982) rule of thumb conventionally used to establish the lower threshold for fri. verification may be conducted by relating pre-fire flowering status of proteoid populations to their post-fire recruitment response at corresponding sites (as for a single site by kraaij et al. 2013) across various species and habitat types. the mapping of unburnt areas within the outer perimeter of fire boundaries could enable assessments of the extent to which these areas act as refugia and sources of seed for recruitment into the burnt landscape. further research on plant responses to fire intensity and articulation of the findings into ecological thresholds for fire intensity would also be informative in fynbos generally and, especially, in environments where weather conditions are variable within seasons and the fire season not clearly delineated. making the link between past fire regimes, as opposed to single fire events, and their ecological outcomes also remains a major challenge. however, the nature of post-fire fynbos assemblages does not solely depend on the fire survival attributes of constituent species and greater understanding is required of the effects of dynamic and competitive interactions between overstorey and understorey species (cowling & gxaba 1990; vlok & yeaton 1999). the practicalities of complying with fire legislation in the face of conflicting land management objectives (fire risk vs biodiversity conservation) need to be considered. a legal review should clarify the particular obligations pertaining to the conservation and commercial forestry sectors with respect to fire and invader plant legislation. this should improve compliance and cooperation, whilst averting the need for legal action between roleplayers (kraaij et al. 2011). concluding recommendations top ↑ recent research aimed to establish what constitutes an acceptable fire regime in eastern coastal fynbos and we accordingly determined ecological thresholds for fire season and return interval. whilst prescribed burns do not have to be constrained by season in the study area, other constraints remain, most significant of which are safety requirements and the abundance of fire-adapted invasive alien trees. because large-scale implementation of prescribed burning is unattainable (keeley et al. 1999; kraaij et al. 2011; van wilgen 2009) and mostly ineffective at ensuring fire safety (keeley & zedler 2009) fire managers need to accept that wildfires will remain dominant. fire management should be adaptive in nature and focused on, (1) managing wildfires to stay within the proposed thresholds, (2) monitoring to assess actual fire regimes against proposed thresholds or to refine thresholds where appropriate, (3) protecting boundaries between fynbos and fire-sensitive land uses (such as plantations) through collaborative agreements with neighbours and (4) integration of fire and invasive alien plant management, using scarce resources to treat invasive plants and focus prescribed burns in priority areas. acknowledgements top ↑ we thank jan vlok for conceptual contributions to this study and south african national parks for exposure to the fire management challenges experienced in the grnp. the comments of two reviewers resulted in improvements to this manuscript. south african national parks, nelson mandela metropolitan university, council for scientific and industrial research (csir) and south african environmental observation network funded this study. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions t.k. (garden route scientific services) largely wrote the manuscript and r.m.c. (nelson mandela metropolitan university) and b.w.v.w. (csir natural resources and the environment) made conceptual contributions. references top ↑ agee, j.k., 2002, ‘the fallacy of passive management. managing for firesafe forest reserves’, conservation in practice 3(1), viewed 13 october 2010, from http://www.conbio.org/cip/redesign/article31fpm.cfm, http://dx.doi.org/10.1111/j.1526-4629.2002.tb00023.xbessie, w.c. & johnson, e.a., 1995, ‘the relative importance of fuels and weather on fire behavior in subalpine forests’, ecology 76(3), 747–762. http://dx.doi.org/10.2307/1939341 biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59–80, island press, new york. bond, w., 1980, ‘fire and senescent fynbos in the swartberg, southern cape’, south african forestry journal 114, 68–71. http://dx.doi.org/10.1080/00382167.1980.9629837 bond, w.j., 1984, ‘fire survival of cape proteaceae – influence of fire season and seed predators’, vegetatio 56, 65–74. bond, w.j., honig, m. & maze, k.e., 1999, ‘seed size and seedling emergence: an allometric relationship and some ecological implications’, oecologia 120, 132–136. http://dx.doi.org/10.1007/s004420050841 bond, w.j., le roux, d. & erntzen, r., 1990, ‘fire intensity and regeneration of myrmecochorous proteaceae’, south african journal of botany 56, 326–330. bond, w.j., vlok, j. & viviers, m., 1984, ‘variation in seedling recruitment of cape proteaceae after fire’, journal of ecology 72, 209–221. http://dx.doi.org/10.2307/2260014 bradstock, r.a. & bedward, m., 1992, ‘simulation of the effect of season of fire on post-fire seedling emergence of two banksia species based on long-term rainfall records’, australian journal of botany 40(1), 75–88. http://dx.doi.org/10.1071/bt9920075 cowling, r.m. & gxaba, t., 1990, ‘effects of a fynbos overstorey shrub on understorey community structure: implications for the maintenance of community-wide species richness’, south african journal of ecology 1, 1–7. deacon, h.j., jury, m.r. & ellis, f., 1992, ‘selective regime and time’, in r.m. cowling (ed.), the ecology of fynbos. nutrients, fire and diversity, pp. 6–22, oxford university press, oxford. de ronde, n., goldammer, j.g., wade, d.d. & soares, r.v., 1990, ‘prescribed fire in industrial pine plantations’, in j.g. goldammer (ed.), fire in the tropical biota. ecosystem processes and global challenges, ecological studies 84, pp. 216–272, springer-verlag, berlin. driscoll, d.a., lindenmayer, d.b., bennett, a.f., bode, m., bradstock, r.a., cary, g.j. et al., 2010, ‘fire management for biodiversity conservation: key research questions and our capacity to answer them’, biological conservation 143, 1928–1939. http://dx.doi.org/10.1016/j.biocon.2010.05.026 enright, n.j. & lamont, b.b., 1989, ‘seed banks, fire season, safe sites and seedling recruitment in five co-occurring banksia species’, journal of ecology 77, 1111–1122. http://dx.doi.org/10.2307/2260826 esler, k.j., van wilgen, b.w., te roller, k.s., wood, a.r. & van der merwe, j.h., 2010, ‘a landscape-scale assessment of the long-term integrated control of an invasive shrub in south africa’, biological invasions 12, 211–218. http://dx.doi.org/10.1007/s10530-009-9443-2 forsyth, g.g. & van wilgen, b.w., 2008, ‘the recent fire history of the table mountain national park, and implications for fire management’, koedoe 50(1), 3–9. http://dx.doi.org/10.4102/koedoe.v50i1.134 geldenhuys, c.j., 1994, ‘bergwind fires and the location pattern of forest patches in the southern cape landscape, south africa’, journal of biogeography 21, 59–62. http://dx.doi.org/10.2307/2845603 gill, a.m. & mccarthy, m.a., 1998, ‘intervals between prescribed fires in australia: what intrinsic variation should apply?’, biological conservation 85, 161–169. http://dx.doi.org/10.1016/s0006-3207(97)00121-3 heelemann, s., procheş, ş., porembski, s. & cowling, r.m., 2010, ‘impact of graminoid cover on postfire growth of nonsprouting protea seedlings in the eastern fynbos biome of south africa’, african journal of ecology 59, 51–55. heelemann, s., procheş, ş., rebelo, a.g., van wilgen, b.w., porembski, s. & cowling, r.m., 2008, ‘fire season effects on the recruitment of non-sprouting serotinous proteaceae in the eastern (bimodal rainfall) fynbos biome, south africa’, austral ecology 33(2), 119–127. http://dx.doi.org/10.1111/j.1442-9993.2007.01797.x hoffman, m.t., cramer, m.d., gillson, l. & wallace, m., 2011, ‘pan evaporation and wind run decline in the cape floristic region of south africa (1974–2005): implications for vegetation responses to climate change’, climatic change 109(3–4), 437–452. http://dx.doi.org/10.1007/s10584-011-0030-z holmes, p.m. & marais, c., 2000, ‘impacts of alien plant clearance on vegetation in the mountain catchments of the western cape’, southern african forestry journal 189, 113–117. http://dx.doi.org/10.1080/10295925.2000.9631286 holmes, p.m. & richardson, d.m., 1999, ‘protocols for restoration based on recruitment dynamics, community structure and ecosystem function: perspectives from south african fynbos’, restoration ecology 7, 215–230. http://dx.doi.org/10.1046/j.1526-100x.1999.72015.x holmes, p.m., richardson, d.m., van wilgen, b.w. & gelderblom, c., 2000, ‘the recovery of south african fynbos vegetation following alien woody plant clearing and fire: implications for restoration’, austral ecology 25, 631–639. http://dx.doi.org/10.1111/j.1442-9993.2000.tb00069.x jefferey, d.j., holmes, p.m. & rebelo, a.g., 1988, ‘effects of dry heat on seed germination in selected indigenous and alien legume species in south africa’, south african journal of botany 54, 28–34. keeley, j.e. & brennan, t.j., 2012, ‘fire-driven alien invasion in a fire-adapted ecosystem’, oecologia 169(4), 1043–1052. http://dx.doi.org/10.1007/s00442-012-2253-8, pmid:22286083 keeley, j.e. & zedler, p.h., 2009, ‘large, high-intensity fire events in southern california shrublands: debunking the fine-grain age-patch model’, ecological applications 19, 69–94. http://dx.doi.org/10.1890/08-0281.1 keeley, j.e., fotheringham, c.j. & morais, m., 1999, ‘reexamining fire suppression impacts on brushland fire regimes’, science 284(5421), 1829–1822. http://dx.doi.org/10.1126/science.284.5421.1829 knox, k.j.e. & clarke, p.j., 2006, ‘fire season and intensity affect shrub recruitment in temperate sclerophyllous woodlands’, oecologia 149, 730–739. http://dx.doi.org/10.1007/s00442-006-0480-6, pmid:16847616 kraaij, t., 2012, fire regimes in eastern coastal fynbos: drivers, ecology and management, phd thesis, department of botany, nelson mandela metropolitan university, port elizabeth. kraaij, t., cowling, r.m. & van wilgen, b.w., 2011, ‘past approaches and future challenges to the management of fire and invasive alien plants in the new garden route national park’, south african journal of science 107(9/10), art. #633, 11 pages. http://dx.doi.org/10.4102/sajs.v107i9/10.633 kraaij, t., cowling, r.m. & van wilgen, b.w., 2012, ‘lightning and fire weather in eastern coastal fynbos shrublands: seasonality and long-term trends’, international journal of wildland fire. http://dx.doi.org/10.1071/wf11167 kraaij, t., cowling, r.m., van wilgen, b.w. & schutte-vlok, a-l., 2013, ‘proteaceae juvenile periods and post-fire recruitment as indicators of minimum fire return interval in eastern coastal fynbos’, applied vegetation science 16, 84–94. http://dx.doi.org/10.1111/j.1654-109x.2012.01209.x kraaij, t., baard, j.a., cowling, r.m., van wilgen, b.w. & das, s., 2012, ‘historical fire regimes in a poorly-understood, fire-prone ecosystem: eastern coastal fynbos’, international journal of wildland fire. http://dx.doi.org/10.1071/wf11163 kruger, f.j., 1982, ‘prescribing fire frequencies in cape fynbos in relation to plant demography’, paper presented at the symposium on the dynamics and management of mediterranean-type ecosystems, 22–26 june 1981, san diego, california. general technical report psw-58, pp. 118–122, pacific southwest forest and range experiment station, forest service and department of agriculture, berkeley. kruger, f.j., 1984, ‘effects of fire on vegetation structure and dynamics’, in p. de v. booysen & n.m. tainton (eds.), ecological effects of fire in south african ecosystems, ecological studies 48, pp. 219–243, springer-verlag, berlin. http://dx.doi.org/10.1007/978-3-642-69805-7_10 kruger, f.j. & bigalke, r.c., 1984, ‘fire in fynbos’, in p. de v. booysen & n.m. tainton (eds.), ecological effects of fire in south african ecosystems, ecological studies 48, pp. 67–114, springer-verlag, berlin. http://dx.doi.org/10.1007/978-3-642-69805-7_5 kruger, f.j. & lamb, a.j., 1978, conservation of the kogelberg state forest. preliminary assessment of the effects of management from 1967 to 1978, jonkershoek forestry research centre, report 79-02, stellenbosch. le roux, p.j., 1969, ‘brandbestryding in suid-kaapland met spesiale verwysing na chemiese metodes van beheer [fire fighting in the southern cape with special reference to chemical methods of control]’, msc thesis, department of forestry, university of stellenbosch. lloret, f., pausas, g. & vilà, m., 2003, ‘responses of mediterranean plant species to different fire frequencies in garraf natural park (catalonia, spain): field observations and modelling predictions’, plant ecology 167, 223–235. http://dx.doi.org/10.1023/a:1023911031155 midgley, j.j., 1989, ‘season of burn of serotinous fynbos proteaceae: a critical review and further data’, south african journal of botany 55, 165–170. midgley, j.j. & viviers, m., 1990, ‘the germination of seeds from heated serotinous cones of eight shrubland species’, south african forestry journal 155, 5–9. http://dx.doi.org/10.1080/00382167.1990.9629064 moritz, m.a., 2003, ‘spatiotermporal analysis of controls on shrubland fire regimes: age dependency and fire hazard’, ecology 84(2), 351–361. http://dx.doi.org/10.1890/0012-9658(2003)084[0351:saocos]2.0.co;2 morrison, d.a., buckney, r.t. & bewick, b.j., 1996, ‘conservation conflicts over burning bush in south-eastern australia’, biological conservation 76, 167–175. http://dx.doi.org/10.1016/0006-3207(95)00098-4 morrison, d.a., cary, g.j., pengelly, s.m., ross, d.g., mullins, b.j., thomas, c.r. et al., 1995, ‘effects of fire frequency on plant-species composition of sandstone communities in the sydney region: inter-fire interval and time-since-fire’, australian journal of ecology 20, 239–147. http://dx.doi.org/10.1111/j.1442-9993.1995.tb00535.x myers, n., mittermeier, r.a., mittermeier, c.g., da fonseca, g.a.b. & kent, j., 2000, ‘biodiversity hotspots for conservation priorities’, nature 403, 853–858. http://dx.doi.org/10.1038/35002501, pmid:10706275 noble, i.r., gill, a.m. & bary, g.a.v., 1980, ‘mcarthur’s fire danger meters expressed as equations’, austral ecology 5, 201–203. raimondo, d., von staden, l., foden, w., victor, j.e., helme, n.a., turner, r.c. et al. (eds.), 2009, ‘red list of south african plants’, strelitzia 25, south african national biodiversity institute, pretoria. rebelo, a.g., boucher, c., helme, n., mucina, l. & rutherford, m.c., 2006, ‘fynbos biome’, in l. mucina & m.c. rutherford (eds.), the vegetation of south africa, lesotho and swaziland, strelitzia 19, pp. 158–159, south african national biodiversity institute, pretoria. republic of south africa, 1998, national veld and forest fire (act no. 101 of 1998), government gazette 401(19515), government printers, cape town. richardson, d.m. & brown, p.j., 1986, ‘invasion of mesic mountain fynbos by pinus radiata’, south african journal of botany 52, 529–535. richardson, d.m. & higgins, s.i., 1998, ‘pines as invaders in the southern hemisphere’, in d.m. richardson (ed.), ecology and biogeography of pinus, pp. 450–473, cambridge university press, cambridge. richardson, d.m. & van wilgen, b.w., 1992, ‘ecosystem, community and species response to fire in mountain fynbos: conclusions from the swartboskloof experiment’, in b.w. van wilgen, d.m. richardson, f.j. kruger & h.j. van hensbergen (eds.), fire in south african mountain fynbos: species, community and ecosystem response in swartboskloof, ecological studies 93, pp. 273–284, springer-verlag, berlin. richardson, d.m., cowling, r.m. & le maitre, d.c., 1990, ‘assessing the risk of invasive success in pinus and banksia in south african mountain fynbos’, journal of vegetation science 1, 629–642. http://dx.doi.org/10.2307/3235569 richardson, d.m., pysek, p., rejmanek, m., barbour, m.g., panetta, d.f. & west, c.j., 2000, ‘naturalization and invasion of alien plants – concepts and definitions’, diversity & distributions 6, 93–107. http://dx.doi.org/10.1046/j.1472-4642.2000.00083.x roura-pascual, n., richardson, d.m., krug, r.m., brown, a., chapman, r.a., forsyth, g.g. et al., 2009, ‘ecology and management of alien plant invasions in south african fynbos: accommodating key complexities in objective decision making’, biological conservation 142(8), 1595–1604. http://dx.doi.org/10.1016/j.biocon.2009.02.029 schulze, r.e., 1997, ‘climate’, in r.m. cowling, d.m. richardson & s.m. pierce (eds.), vegetation of southern africa, pp. 21–42, cambridge university press, cambridge. seydack, a.h.w., 1992, ‘fire management options in fynbos mountain catchment areas’, south african forestry journal 161, 53–58. http://dx.doi.org/10.1080/00382167.1992.9630425 seydack, a.h.w., bekker, s.j. & marshall, a.h., 2007, ‘shrubland fire regime scenarios in the swartberg mountain range, south africa: implications for fire management’, international journal of wildland fire 16, 81–95. http://dx.doi.org/10.1071/wf06015 syphard, a.d., keeley, j.e. & brennan, t.j., 2011, ‘comparing the role of fuel breaks across southern california national forests’, forest ecology and management 261, 2038–2048. http://dx.doi.org/10.1016/j.foreco.2011.02.030 thuiller, w., slingsby, j.a., privett, s.d.j. & cowling, r.m., 2007, ‘stochastic species turnover and stable coexistence in a species-rich, fire-prone plant community’, plos one 2(9), e938. http://dx.doi.org/10.1371/journal.pone.0000938 van wilgen, b.w., 1984, ‘fire climates in the southern and western cape province and their potential use in fire control management’, south african journal of science 80, 358–362. van wilgen, b.w., 2009, ‘the evolution of fire and invasive alien plant management practices in fynbos’, south african journal of science 105, 335–342. van wilgen, b.w. & richardson, d.m., 1985, ‘factors influencing burning by prescription in mountain fynbos catchment areas’, south african forestry journal 134, 22–32. http://dx.doi.org/10.1080/00382167.1985.9629590 van wilgen, b.w. & richardson, d.m., 2012, ‘three centuries of managing introduced conifers in south africa: benefits, impacts, changing perceptions and conflict resolution’, journal of environmental management 106, 56–68. http://dx.doi.org/10.1016/j.jenvman.2012.03.052, pmid:22562012 van wilgen, b.w. & viviers, m., 1985, ‘the effect of season of fire on serotinous proteaceae in the western cape and the implications for fynbos management’, south african forestry journal 133, 47–53. http://dx.doi.org/10.1080/00382167.1985.9629574 van wilgen, b.w., forsyth, g.g. & prins, p., 2012, ‘the management of fire-adapted ecosystems in an urban setting: the case of table mountain national park, south africa’, ecology and society 17(1), 8. http://dx.doi.org/10.5751/es-04526-170108 van wilgen, b.w., richardson, d.m. & seydack, a.h.w., 1994, ‘managing fynbos for biodiversity: constraints and options in a fire-prone environment’, south african journal of science 90, 322–329. van wilgen, b.w., govender, n., forsyth, g.g. & kraaij, t., 2011, ‘towards adaptive fire management for biodiversity conservation: experience in south african national parks’, koedoe 53(2), art. #982, 9 pages, http://dx.doi.org/10.4102/koedoe.v53i2.982 van wilgen, b.w., forsyth, g.g., de klerk, h., das, s., khuluse, s. & schmitz, p., 2010, ‘fire management in mediterranean-climate shrublands: a case study from the cape fynbos, south africa’, journal of applied ecology 47(3), 631–638. http://dx.doi.org/10.1111/j.1365-2664.2010.01800 vermeulen, w.j., van der merwe, c.j., seydack, a.h.w., baard, j., cunningham, h., kraaij, t. et al., 2009, fire management system for the tsitsikamma mountain catchment area, south african national parks, knysna. vlok, j.h.j. & yeaton, r.i., 1999, ‘the effect of overstorey proteas on plant species richness in south african mountain fynbos’, diversity & distributions 5, 213–222. http://dx.doi.org/10.1046/j.1472-4642.1999.00055.x watson, l.h. & cameron, m.j., 2001, ‘the influence of fire on a southern cape mountain forest’, south african forestry journal 191, 39–42. http://dx.doi.org/10.1080/20702620.2001.10434149 wilson, a.m., latimer, a.m., silander, j.a. jnr, gelfand, a.e. & de klerk, h., 2010, ‘a hierarchical bayesian model of wildfire in a mediterranean biodiversity hotspot: implications of weather variability and global circulation’, ecological modelling 221, 106–112. http://dx.doi.org/10.1016/j.ecolmodel.2009.09.016 article information authors: gordon c. o’brien1,2 nico j. smit1 victor wepener1 affiliations: 1unit for environmental sciences and management, north-west university, south africa2department of zoology, university of johannesburg, south africa correspondence to: gordon o’brien postal address: private bag x6001, potchefstroom 2520, south africa dates: received: 02 nov. 2012 accepted: 09 sept. 2013 published: 25 feb. 2014 how to cite this article: o’brien, g.c., smit, n.j. & wepener, v., 2014, ‘conservation of fishes in the elands river, mpumalanga, south africa: past, present and future’, koedoe 56(1), art. #1118, 8 pages. http://dx.doi.org/10.4102/ koedoe.v56i1.1118 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. conservation of fishes in the elands river, mpumalanga, south africa: past, present and future in this original research... open access • abstract • introduction • research method and design    • setting    • procedure • results • ethical considerations • discussion    • conservation requirements    • recommendations • conclusion • acknowledgements    • competing interests    • authors’ contributions • references • footnotes abstract top ↑ in an isolated reach, between two large natural waterfalls in the elands river in mpumalanga, populations of a critically endangered kneria sp., the endangered chiloglanis bifurcus and a genetically unique population of labeobarbus polylepis occur. the aim of this article was to evaluate past efforts to conserve these fishes, describe the current status and propose future conservation and management actions. the population status assessments were based on a series of fish community composition and population structure evaluations from surveys undertaken at 22 sites during seven surveys from 2002 to 2006. although water-use activities have continued to increase in the area, impacts have been offset by conservation efforts initiated almost 30 years ago. the existing c. bifurcus population appears to be stable, which is reflected in the downgrading of the conservation status of the species from critically endangered to endangered. the abundance of the kneriid population appears to be increasing and spreading to other tributaries in the study area. the abundance of l. polylepis appears to be increasing but has still not reached historical levels.conservation implications: continued conservation efforts are required to protect these fishes. this case study presented a rare example of how the impacts associated with the use of aquatic resources in south africa can successfully be offset by conservation efforts. introduction top ↑ the elands river is a biologically diverse tributary of the crocodile river in south africa which is being used moderately and has been maintained in a suitable integrity state (o’brien 2012). a large waterfall (> 10 m) isolates the fish of the elands river and some tributaries from the crocodile river catchment (gaigher 1969). within this isolated portion of the catchment, populations of a critically endangered kneria sp., the endangered incomati rock catlet (chiloglanis bifurcus jubb & le roux, 1969) and a genetically unique population of the bushveld smallscale yellowfish (labeobarbus polylepis boulenger, 1907) occur (figure 1) (engelbrecht & bills 2007a; engelbrecht & roux 2002; o’brien 2009, 2012). following an accidental spill of paper mill effluent into the elands river at ngodwana in 1989 (kleynhans et al. 1992), the conservation and management of the river received considerable attention (ferreira, wepener & van vuren 2008, 2009; kleynhans et al. 1992). this included the establishment of a sanctuary in the ngodwana river for c. bifurcus, introduction of a population of kneria sp. from the upper crocodile river and the establishment of the elands river yellowfish conservation area (eryca) for the l. polylepis population. additional conservation efforts include the recommendations that the elands river be declared a fish sanctuary within the national freshwater priority area (nfepa) (driver et al. 2011). the aim of this article is to evaluate the past conservation efforts, describe the current population status and provide management recommendations for the fishes that are threatened and/or unique to the elands river and its tributaries. figure 1: the elands river and associated rivers within the inkomati river catchment, southern africa, where the populations of fishes that were evaluated in the study occur. research method and design top ↑ setting the study area included the 62 km reach of the elands river and its tributaries between the waterval-boven and de villiers waterfalls and a portion of the crocodile river upstream and downstream of the elands river confluence, including the houtbosloop river (figure 1). numerous natural barriers, including the waterval-boven waterfall downstream of waterval-boven and de villiers waterfall upstream of the crocodile river confluence on the elands river, exist in the study area. other artificial barriers include impoundments on the ngodwana and swartkoppies rivers and a gauging weir on the elands river (figure 1). these barriers restrict the movement of fishes in the study area. numerous activities alter the water quality, quantity and fish habitat in the area, including the sappi ngodwana pulp and paper mill, forestry and other agriculture plantations, as well as the urban areas of machadodorp, waterval-boven and ngodwana (ferreira et al. 2008, 2009; hill 2005; o’brien 2012). other impacts associated with the occurrence of extra-limital and alien fishes also exist in the study area.nine representative river reaches were identified in the study area where surveys were carried out to evaluate the status of fish populations (figure 1). the criteria considered for reach selection included nfepa fish sanctuary boundaries, the location of natural and artificial barriers, eco-region boundaries and the spatial extent of important land-use activities. reach 1 (r1) included the elands river from the waterval-boven waterfall and the confluence of the ngodwana river. the elands river in this reach is a fast-flowing, rejuvenated mountain stream with a well-defined incised channel. sites e1–e5 were positioned in this reach. reach 2 (r2) included the elands river between the confluences of the ngodwana and lupulele rivers below the sappi pulp and paper-making activities. the elands river in this reach is dominated by pools and intermittent riffle and rapid complexes. two sampling sites (e6 and e7) were positioned within r2. reach 3 (r3) extended from the lupulele river on the elands river to the de villiers waterfall. the elands river in r3 is similar to r2 and contains two sampling sites (e8 and e9). reach 4 (r4) included a uniform reach of the crocodile river upstream of the confluence with the elands river where two sites (c1 and c2) were located. the habitat type of the crocodile river in r4 is similar to that which occurs in r2 and r3 in the elands river. poorly managed flow releases from the man-made kwena lake are known to impact on this portion of the crocodile river (o’brien 2012). reach 5 (r5) included a portion of the lower elands, upper crocodile and lower houtbosloop rivers. two sampling sites (c3 and h4) were located in r5. reach 6 (r6) included the upper houtbosloop river above an artificial barrier. this part of the houtbosloop river can be described as a small, fast-flowing mountain stream that is dominated by riffles and rapids. three sampling sites (h1–h3) were positioned within r6. reach 7 (r7) included the lupulele river which is a small, slow-flowing clear mountain stream that is dominated by many shallow pools (± 1 m deep). one sampling site (l1) was positioned in the lower reaches of the lupulele river. reach 8 (r8) included the minimally used upper ngodwana river above the man-made ngodwana lake. the habitat of the upper ngodwana river in r8 is comparable to the upper elands (r1) and houtbosloop rivers (r6). one sampling site (n1) was located within r8. reach 9 (r9) included the swartkoppies river, which is a small mountain stream that has been segmented by artificial barriers to establish pools for a local alien fisheries activity. two sampling sites were located in r9, one upstream (s1) and one downstream (s2) of the first artificial barrier on the swartkoppies river. procedure seven fish surveys were undertaken from march 2002 to august 2006 during high summer and low winter flow periods at 22 sites (figure 1) (o’brien 2009, 2012). in addition, data obtained from a single survey carried out to a shallow (< 1 m) tributary of the elands river in r1 was included in the study (figure 1; palmer pers. comm., 06 august 2012)1. electrofishing was used to collect fish in all shallow (< 1 m) habitats and gill nets were used to collect fishes in deep (> 1 m) habitats. the collected fishes were identified, measured (fork length) and released in close proximity to the collection location. the number of fishes collected per minute of electrofishing and gill net deployments were recorded independently as the catch per unit effort (cpue). gill net mesh sizes of 57 mm and 73 mm in 25 m segments were selectively used to target adult l. polylepis individuals in pools. population structure assessments of the c. bifurcus and l. polylepis were used to evaluate the status of these populations. the proportion of juveniles (young of each year surveyed), immature sub-adults and mature adult individuals collected within each reach of the study area were used to describe the population structures. population structures were based on length distributions of fishes in relation to attainable lengths (skelton 2001). temporal comparisons of cpue results from kleynhans et al. (1992) and james (1992) were compared to cpues obtained in the study (table 1 and figure 2). trends in the cpue of c. bifurcus and l. polylepis populations from the study area between 1978–1991 (james 1992; kleynhans et al. 1992) and 2002–2012 (o’brien 2009, 2012) were evaluated. insufficient kneria sp. samples were collected for inclusion in the population assessment; however, comparisons between historical and recent collections allowed for a descriptive assessment of the integrity of the population. table 1: catch per unit effort of the juveniles, sub-adult and adult chiloglanis bifurcus and labeobarbus polylepis collected using electrofishing and gill net sampling techniques during fish community structure assessments of the study from 2002 to 2006. figure 2: population structure of chiloglanis bifurcus and labeobarbus polylepis from each site considered in the study. charts present percentage proportions of juveniles, sub-adults and adults. results top ↑ sixty-nine c. bifurcus were collected using electrofishing during 18 surveys from seven of the nine reaches sampled during the study (table 1 and figure 2). the cpue rates for c. bifurcus ranged between 0.04 and 1.73 individuals per minute when collected. the population structures of this species varied noticeably (figure 2). high proportions of juvenile, sub-adult and adult c. bifurcus individuals were collected at r1 in the elands river upstream of ngodwana, downstream of ngodwana in the elands river at r2, in the crocodile river at r5 and in the upper ngodwana river at r8 (figure 2). only adult c. bifurcus were collected in the elands river below the lupulele river confluence in r3, in the upper houtbosloop river at r6 and in the swartkoppies river at r9. a noticeable decrease in the cpue of c. bifurcus in the elands and crocodile rivers from r1 and r2 to r3 and r5 were observed (table 1). from october 1989, the cpue of c. bifurcus populations in four of the six reaches sampled reduced noticeably (figure 3). figure 3: temporal trends in the catch per unit effort (number of fish per minute) of chiloglanis bifurcus collected in the elands river between 1978 and 1989 (mean of available data) and between october 1989 and 2006. arrow depicts the occurrence of the 1989 black liquor spill into the elands river. the cpue of the population in the upper elands and houtbosloop rivers remained relatively stable for at least another year until march 1990. during june and september 1990, the lowest abundance of c. bifurcus (cpue < 0.20) was observed. from october 1990 to september 1991, the cpues of c. bifurcus populations in many reaches fluctuated considerably and then returned to low levels in october 1991. from march 2002 to august 2006, apart from a cpue of 1.73 in r1 during march 2002, the cpue of c. bifurcus populations did not exceed 0.33 individuals per minute (table 1 and figure 3). one hundred and twenty-three l. polylepis individuals were collected on 13 occasions in the study area during the electrofishing and gill net efforts from eight sites representing five reaches of the study area (table 1). the cpue rates for l. polylepis ranged between 0.00 and 0.04 when collected, to 1.20 fish per minute using electrofishing techniques and 0.00–3.00 fish per minute using gill netting techniques (table 1). shoaling behaviour of l. polylepis was observed where numerous (5–45 individuals) were collected together. the local l. polylepis population preferred deep pools during winter where they were actively targeted using gill nets (o’brien 2009). these deep pool areas were uncommon in the upper reaches of the elands river, the swartkoppies, lupulele and ngodwana rivers but common in the middle and lower elands river (r2 and r3). in r1 alone, all size categories of l. polylepis were collected. adult l. polylepis dominated the upper portion of the elands river in the study area and the lower swartkoppies river sites, (r1, r2 and r9). juvenile l. polylepis were common in the upper reaches of the ngodwana (r8) and swartkoppies (r9) rivers and in the crocodile river in the vicinity of the confluence of the houtbosloop river at r5. although l. polylepis were collected between 1978 and october 1991, no cpue data are available for this period. from 2002, large abundances l. polylepis were initially collected from r1 and r5 in the elands river, and from r8 in the upper ngodwana river (figure 4). the observed cpues varied considerably and were influenced by the shoaling behaviour of l. polylepis. these findings were substantiated in 2004 following the discovery of large shoals of adult l. polylepis in r1 of the elands river in february 2004 (eryca 2004). in addition, during the spring of 2004, large numbers of l. polylepis were observed migrating upstream in the elands river, approximately 5 km below the waterval-boven waterfall (eryca 2004). in 2007, approximately 50 kneria sp. were collected in the unnamed tributary of the elands river (r1) using electrofishing techniques. although no cpue data are available, these findings show that the kneria sp. population relocated into r1 remains in relatively large abundances (palmer pers. comm., 06 august 2012). figure 4: temporal trends in the catch per unit effort (number of fish per minute) of labeobarbus polylepis collected in the elands river between 2002 and 2006. ethical considerations top ↑ the fisheries biology research presented in this paper was carried out under the authorisation of mpumalanga parks and tourism permits issues on the 09 march 2005 (mpb. 5131/2), 31 january 2006 (mpb. 5161), 24 january 2007 (mpb. 5184) and 02 june 2008 (mpb. 5222). discussion top ↑ it is well known that the fish communities of the elands river vary spatially and temporally throughout the study area (gaigher 1969; james 1992; kleynhans et al. 1992; o’brien 2012). apart from the effects of natural barriers, artificial physical and chemical barriers have been identified as major threats that affect fish community structures in the study area (o’brien 2012). other impacts attributed to shifts in fish community structures included habitat availability and water quality and quantity alterations of natural and anthropogenic origin (o’brien 2012). abundant, well-structured populations of c. bifurcus and l. polylepis that contained high proportions of juvenile, sub-adult and adult individuals occurred in the elands river, swartkoppies and ngodwana rivers prior to the pollution events in the mid-to-late 1980s (gaigher 1969; kleynhans et al. 1992). these pollution events included a series of small to moderate (< 5000 l) chemical spills into the elands river in february 1985, july 1986, june 1989 and culminated in the large effluent spill in september 1989 (kleynhans et al. 1992). these events caused the c. bifurcus and l. polylepis population structures to change considerably, particularly downstream of the source of the spills, in the vicinity of ngodwana and the lower elands river into the upper crocodile river (kleynhans et al. 1992). although by 1992 some recovery had occurred, the abundance of individuals has not recovered to pre-1980 levels (james 1992; kleynhans et al. 1992). until the recent discovery of a population of c. bifurcus in swaziland (monadjem et al. 2003), the known distribution of c. bifurcus was restricted to the upper crocodile river and the elands river and its major tributaries (skelton 2001). following an evaluation of the effects of the chemical spill into the elands river on 23 september 1989, conservation efforts for these populations were initiated (kleynhans et al. 1992). james (1992) proposed that the species be allocated critically endangered status because of the noticeable reduction in distribution and abundance of populations in the elands and upper crocodile rivers. in 1996, c. bifurcus was formally assigned critically endangered status by the international union for conservation of nature (iucn) (engelbrecht & bills 2007a). the vulnerability criteria included the restricted distribution of the remaining populations in the inkomati river catchment (figure 1) and intolerance to water quality and flow alterations (engelbrecht & bills 2007a; james 1992). in addition, james (1992) recommended that a sanctuary be established in the upper ngodwana river for the local elands river population and in the houtbosloop river for the crocodile river population. in 1992, a small population of c. bifurcus individuals (n = ± 100) were relocated into the ngodwana river for this purpose. apart from the lupulele river (r7) and upper crocodile river (r4) reaches, the c. bifurcus populations still extend throughout the study area and now include the upper ngodwana river (r8). the most abundant populations occur in the upper elands (r1), ngodwana (r8) and crocodile rivers (r5). from the findings, we can surmise that although the local c. bifurcus population has been affected by the spill in 1989, other events or activities may have affected local populations. some of the reductions in the cpue of c. bifurcus populations were observed from 1990 and occurred in areas that were not affected by the spill. from august 2002 to 2006, 17 years after the spill, the cpue scores of all c. bifurcus populations had still not recovered to levels observed prior to 1989 (figure 3). in 2007, however, after the discovery of a population of c. bifurcus in swaziland (monadjem et al. 2003), the conservation status of c. bifurcus was amended to endangered (engelbrecht & bills 2007a; scott et al. 2006; skelton 1993, 2001). the abundance of the local l. polylepis population also decreased substantially in 1989 following the chemical spill into the elands river (kleynhans et al. 1992). despite the occurrence of known unique morphological features of the local population from 1969 (gaigher 1969), relocations of l. polylepis individuals from the komati river to the elands river occurred between 1993 and 1996 in an attempt to increase the abundances of the local population (engelbrecht pers. comm., 03 june 2006)2. although actual numbers of individuals released were not recorded, it is estimated that between 35 and 50 were released into the elands river, upstream of ngodwana at hemlock, and into the man-made ngodwana lake. these conservation efforts formed part of a conservation plan for the l. polylepis population in the elands river developed by roux (2008). in addition, a captive breeding programme was initiated in an attempt to breed and restock l. polylepis into the elands river (eryca 2004). in 2000, a oncorhynchus mykiss (walbaum 1792) breeding facility (trout farm) at arlie estates (located in the upper reaches of the study area, r1) was stocked with l. polylepis individuals from the komati river for the breeding programme. the culturing efforts were not successful and no l. polylepis from arlie estates were released into the elands river. in 2004, the eryca strategy was developed to conserve the fishes of the elands river using l. polylepis as a flagship species (eryca 2004). then, in 2004, mulder et al. discovered that consistent genetic variation occurred between isolated l. polylepis populations. based on these findings and recommendations made by the eryca strategy, a morphological and genetic diversity and ecological evaluation of the local l. polylepis population was undertaken (o’brien 2009). findings of the study confirmed that the elands river l. polylepis population is genetically and morphologically unique, with unique biological aspects (o’brien 2009). in 1983, approximately 200 individuals of an undescribed, critically endangered kneria sp. population from the alexander spruit were relocated into a tributary of the elands river (figure 1) (engelbrecht & bills 2007b; kleynhans pers. comm., 11 november 2012)3. the distribution range of these knerians is restricted to the upper crocodile river above the confluence with the elands river in mpumalanga (figure 1). the population of the kneria sp. in the alexander spruit was threatened by habitat destruction associated with the construction of the man-made kwena lake (figure 1). other threats include population fragmentation, predation by alien fishes (e.g. o. mykiss) and water quality and habitat altering water resource use activities (engelbrecht & bills 2007b). some fragmented populations have become extinct as a result of these threats (engelbrecht & bills 2007a). the remaining natural area of habitation by the kneria sp. is less than 10 km². the tributary of the elands river where the knerians were relocated to was considered to be suitable for the establishment of a sanctuary for this critically endangered species (engelbrecht & bills 2007b). although no kneria sp. individuals were collected in the elands river in close proximity to the relocation point between 2002 and 2006, the collection of numerous individuals (n = ± 50) in a small tributary of the elands river in 2007 other than the release site, indicates that the relocated population still exists 29 years after the relocation and may now occur in at least two tributaries of the elands river. conservation requirements the conservation efforts in the study area have had varying degrees of success. in particular, the continued survival of the kneria sp. population is encouraging. the endangered c. bifurcus population from the study area still exists in low but stable abundances when compared with historical cpues (figure 3). the conservation efforts to establish a population of c. bifurcus in the ngodwana river have been successful and results showed that this population is stable and increasing. the only population of c. bifurcus that appears to be decreasing is the population in the houtbosloop river, which has continuously reduced from 1992. the local l. polylepis population in the elands river appears to be stable and increasing in abundance. the continued management and conservation of the elands river and its tributaries is of great importance for the continued conservation efforts of these fishes. in accordance, the elands river catchment has been selected as a fish sanctuary within the nfepa programme (nel et al. 2011). the large number and diversity of water resource use activities in the study are known to pose a threat to the viability of the populations of local fishes (o’brien 2012). these stressors include water quality and quantity impacts and threats from alien and invasive fishes and habitat alterations (o’brien 2012). water quality alteration threats include point source pollution events such as the chronic accidental spill and the continuous diffuse release of effluent from the pulp and paper mill, various agriculture activities in the area and partially treated effluent from the machadodorp, waterval-boven and ngodwana wastewater treatment works (ferreira et al. 2009; o’brien 2012). water quantity impacts include the alteration of volume, timing and duration of flows in the elands river by the pulp and paper-making activities at ngodwana, which includes the management of the man-made ngodwana lake (ferreira et al. 2009; o’brien 2012). the man-made ngodwana lake is a 10 m3× 106 m3 water storage facility that was constructed in the early 1980s on the lower ngodwana river, a tributary of the elands river. shortly after construction, a recreational sport angling activity was established by local community members who established extra-limital populations of the mozambique tilapia (oreochromis mossambicus), redbreast tilapia (tilapia rendalli), sharptooth catfish (clarias gariepinus), sidespot barb (barbus neefi), the silver robber (micralestes acutidens) and bushveld smallscale yellowfish (l. polylepis) and alien species including common carp (cyprinus carpio), largemouth bass (micropterus salmoides) and rainbow trout (o. mykiss) in the lake (eryca 2004; schroeder pers. comm., 16 february 2005)4. many of these alien fishes have established populations in the elands river and some of its tributaries (o’brien 2012). oncorhynchus mykiss individuals are still being stocked into the elands river upstream of the swartkoppies river confluence and into the swartkoppies river to maintain the fly-fishing activities in these areas (eryca 2004). recommendations although the abundance of the c. bifurcus population in the ngodwana river is increasing, the abundance of the individuals in the houtbosloop river has decreased. in the elands river, although the c. bifurcus population is lower than historical levels, the abundances do not appear to be decreasing. continued monitoring and protection measures are required. efforts similar to those established in the ngodwana and elands rivers should be developed for the houtbosloop river and other populations in the inkomati catchment. this should include an assessment of the genetic diversity of isolated c. bifurcus populations. the possibility that the critically endangered kneria sp. has established itself in the other tributaries of the elands river is encouraging, but needs validation. the status of the original remaining populations in the upper crocodile river is largely unknown and should also be considered. the critically endangered conservation status of this species is likely to remain for the foreseeable future. although the abundance of the unique l. polylepis population in the elands river appears to be increasing, numerous threats to the sustainability of the population still exist. the continued conservation of the local l. polylepis population should be encouraged through awareness and the implementation of conservation plans. financial resources availability continues to be a limitation for these conservation endeavours. the development of a sustainable catch and release angling industry for the local l. polylepis population should be considered. similar industries have been established in other parts of south africa, such as the vaal river where the local yellowfish-dependent angling industry is valued at over r133 million per annum (brand et al. 2009). wepener and chapman (2012) argue that, for the protection of biodiversity, there should be a shift away from conserving ecosystem health towards ecosystem services. therefore the potential to establish an economic value associated with the conservation of fishes could contribute greatly to the sustainability of conservation efforts. conclusion top ↑ in the almost 30 years following the initiation of conservation efforts for fishes in the elands river, the threats to local populations have continued to increase. the impacts of the threats have, however, been offset by the conservation efforts established in the area. although utilised, the elands river and its associated tributaries are considered to be in a healthy state (o’brien 2012). although recoveries from the 1989 events appear to be ongoing 24 years later, the populations of fishes appear to be growing. this case study presents a rare example of how the impacts associated with the use of aquatic resources in south africa can successfully be offset by conservation efforts. in addition, it is only because of the availability of long-term monitoring data that the status of the populations can be evaluated. these actions have all been established and/or carried out within the context of existing management and conservation endeavours with the willingness of the local stakeholders to maintain biodiversity and keep ecosystem service use sustainable. acknowledgements top ↑ the financial contributions made to this study by sappi (pty) ltd, the department of trade and industry (thrip programme) and the water research commission are acknowledged. we further would like to thank mr n. james, dr c.j. kleynhans, dr j.s. engelbrecht, mr f. roux and dr w. vlok for specialist advice and the bsc honours and msc students from the centre of aquatic research at the university of johannesburg who carried out their studies on the system. finally, we would like to thank participants of the eryca workshop and members of the elands river conservancy who facilitated this work, in particular the late mr george harvermaul, garth johnson and dave hempson from the eryca. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions g.c.o. (north-west university) was responsible for the study conceptualisation, data collection and iteration, analyses, report writing and editing, whilst n.j.s. (north-west university) contributed to the study conceptualisation, data iteration, report writing and editing and v.w. (north-west university) contributed to the study conceptualisation, report writing and editing. references top ↑ brand, m., maina, j., mander, m. & o’brien, g., 2009, ‘characterisation of the social and economic value of the use and associated conservation of the yellowfishes in the vaal river’, water research commission report kv226/09, water research commission, pretoria.driver, a., nel, j.l., snaddon, k., murray, k., roux, d.j., hill, l. et al., 2011, ‘technical report for the freshwater ecosystem priority areas project’, water research commission report 1801/2/11, pp. 16–22, water research commission, pretoria. elands river yellowfish conservation area, 2004, ‘elands river yellowfish conservation area: conservation plan’, outcomes of the stakeholders workshop for the establishment of the elands river yellowfish conservation area strategy, hosted by the elands river conservancy, ntsinini, 28 august. engelbrecht, j. & bills, r., 2007a, ‘chiloglanis bifurcus’, in iucn red list of threatened species, version 2012.1, viewed 03 september 2012, from http://www.iucnredlist.org engelbrecht, j. & bills, r., 2007b, ‘kneria sp. nov. south africa’, in iucn red list of threatened species, version 2012.1, viewed 03 september 2012, from http://www.iucnredlist.org engelbrecht, j.s. & roux, f., 2002, ‘fish’, in a.j. emery, m. lötter & s.d. williamson (eds.), ‘determining the conservation value of land in mpumalanga’, pp. 137–146, unpublished report prepared for the department of water affairs and forestry and department for international development, by the mpululanga parks board, lydenburg. ferreira, m., wepener, v. & van vuren, j.h.j., 2008, ‘die invloed van papierpulpmeule-aktiwiteite op die visgemeenskapstruktuur van die elandsrivier, mpumalanga’, suid-afrikaanse tydskrif vir natuurwetenskap en tegnologie 27, 83–94. ferreira, m., wepener, v. & van vuren, j.h.j., 2009, ‘spatial and temporal variation in the macroinvertebrate community structure of the lower elands river, mpumalanga, south africa’, african journal of aquatic science 34, 231–238. http://dx.doi.org/10.2989/ajas.2009.34.3.4.980 gaigher, i.g., 1969, ‘aspekte met betrekking tot die ekologie, geographie en taksonomie van varswatervisse in die limpopo en incomatiriviersisteem’, phd thesis, department of zoology, rand afrikaans university. hill, l., 2005, ‘elands catchment comprehensive reserve determination study, mpumalanga province, ecological classification and ecological water requirements (quantity) workshop report’, contract report for sappi-ngodwana env-p-c 2004-019, pp. 1–98, submitted to the department water affairs and forestry, by the division of water environment and forestry technology, csir, pretoria. james, n.p.e., 1992, ‘the conservation status of the incomati rock catlet chiloglanis bifurcus, in the elands river, eastern transvaal, and research into artificial propagation’, south african institute for aquatic biodiversity investigational report 40, saiab, grahamstown. kleynhans, c.j., schulz, g.w., engelbrecht, j.s. & roussaeu, f.j., 1992, ‘the impact of a paper mill effluent spill on the fish populations of the elands and crocodile rivers (inkomati system, transvaal)’, water sa 18, 73–80. monadjem, a., boycott, r., parker, v. & culverwell, j., 2003, ‘threatened vertebrates of swaziland’, in swaziland red data book: fishes, amphibians, reptiles, birds and mammals, pp. 66–72, ministry of tourism, environment and communication, mbabane. mulder, p.f.s., engelbrecht, d., engelbrecht, j.s. & roux, f., 2004, ‘biochemical genetic variation between four populations of labeobarbus polylepis from three river systems in south africa’, african journal of aquatic science 29, 97–102. http://dx.doi.org/10.2989/16085910409503797 nel, j.l., murray, k.m., maherry, a., petersen, c.p., roux, d.j., driver, a.l. et al., 2011, ‘technical report for the national freshwater ecosystem priority areas project’, water research commission technical report 1801/2/11, pp. 16–17, water research commission, pretoria. o’brien, g.c., 2009, ‘aspects of the ecology and population management of the bushveld smallscale yellowfish (labeobarbus polylepis)’, water research commission report kv225/09, pp. 21–52, water research commission, pretoria. o’brien, g.c., 2012, ‘regional scale risk assessment methodology using the relative risk model as a management tool for aquatic ecosystems in south africa’, phd thesis, department of zoology, university of johannesburg. roux, f., 2008, ‘status of the bushveld smallscale yellowfish’, in n.d. impson, i.r. bills & l. wolhuter (eds.), technical report on the state of yellowfishes in south africa in 2007 kv212/08, pp. 65–72, water research commission, pretoria. scott, l.e.p., skelton, p.h., booth, a.j., verheust, l., harris, r. & dooley, j., 2006, ‘atlas of southern african freshwater fishes’, monograph 2, smithiana, south african institute for aquatic biodiversity, grahamstown. skelton, p.h., 1993, a complete guide to the freshwater fishes of southern africa, 1st edn., southern publishers (pty.) ltd., halfway house. pmcid:pmc1491714 skelton, p.h., 2001, a complete guide to the freshwater fishes of southern africa, 2nd edn., southern publishers (pty.) ltd., halfway house. wepener, v. & chapman, p.m., 2012 ‘south african ecotoxicology – present status and future prognosis’, african journal of aquatic science 37(3), 229–234. http://dx.doi.org/10.2989/16085914.2012.717051 footnotes top ↑ 1. dr r. palmer is the director of nepid consulting, an environmental and water resource management consultancy operating out of white river. 2. dr johan engelbracht is a former fisheries scientist of the mpumalanga parks and tourism, based in lydenburg, mpumalanga.3. dr c.j. kleynhans is the chief specialist scientist at resource quality services, department of water affairs, roodplaat, and a former ecologist of the transvaal chief directorate of nature and environmental conservation, karenpark. 4. mr e.w. schroeder is the former chairman of the ngodwana boating and angling club, ngodwana. reviewer acknowledgementpage 1 of 1 http://www.koedoe.co.za open access the editorial team of koedoe recognises the value and importance of the peer reviewer in the overall publication process – not only in shaping the individual manuscript, but also in shaping the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely, productive peer review process. we would like to take this opportunity to thank all reviewers who participated in shaping this volume of koedoe. we appreciate the time taken to perform your review(s) successfully. in an effort to facilitate the selection of appropriate peer reviewers for koedoe, we ask that you take a moment to update your electronic portfolio on https://koedoe.co.za for our files, allowing us better access to your areas of interest and expertise, in order to match reviewers with submitted manuscripts. if you would like to become a reviewer, please visit the journal website and register as a reviewer. to access your details on the website, you will need to follow these steps: 1. log into the online journal at https://koedoe.co.za 2. in your ‘user home’ [https://koedoe.co.za/ index.php/koedoe/user] select ‘edit my profile’ under the heading ‘my account’ and insert all relevant details, bio statement and reviewing interest(s). 3. it is good practice as a reviewer to update your personal details regularly to ensure contact with you throughout your professional term as reviewer to koedoe. please do not hesitate to contact us if you require assistance in performing this task. publisher: publishing@aosis.co.za tel: +27 21 975 2602 tel: 086 1000 381 alan whitfield andrew davies andrew t. hudak andy blackmore arne b.r. witt benjamin d. dovie bernard w.t. coetzee carly r. cowell chenay simms cloe guerbois colin everson david le maitre dave f. joubert edmund february edson gandiwa ernest seamark felix patton gareth hempson haemish melville hannes botha helga van der merwe hendrik sithole hindrik bouwman inam yekwayo jackie abell jesse m. kalwij kelly retief klaus kellner lesley henderson louise swemmer maddy case mario melleti mark alderweireldt mark brigham matt w. hayward matthew child mduduzi ndlovu michael h. knight michael j. somers michael t. hoffman michelle greve michelle j. tedder nicky knox nicole bezemer norbert cordeiro pedro m. cardoso peter a. novellie petri c. viljoen ray jansen robert m.r. barclay roland hunter ryan blanchard ryan phillips samuel williams sandra macfadyen sascha buchholz sonja matthee steve johnson sue snyman sue taylor theo h.c. mostert tim haas tony swemmer trevor d. harrison upenyu dzingirai vivienne uys wayne twine wendy annecke xander combrink zoliswa n. nhleko 00 acknowledgement to reviewers https://koedoe.co.za https://koedoe.co.za https://koedoe.co.za/index.php/koedoe/user https://koedoe.co.za/index.php/koedoe/user mailto:publishing@aosis.co.za filelist convert a pdf file! koedoe 19: 63-66 (1976) a note on mountain reedbuck redunca fulvorufula fulvorufula afzelius in the kruger national park l. r. irby caesar kleberg research programme in wildlife ecology department oj wildlife and fisheries sciences texas a & m university college station texas 7784) abstract this survey was conducted to obtain information on the mountain reedbuck population in the kruger national park for comparison with the population in the loskop dam nature reserve, transvaal. during 93 hours of search effort, 36 mountain reedbuck sightings were recorded. all of the sightings were in the malelane hills on slopes above the kwa machiyaliwane drainage. introduction mountain reedbuck in the kruger national park (knp) are found only in the malelane hills and along the lebombo mountain. the population is estimated to number 240 (pienaar 1969). the status of this species in the park is considered precarious by many people, but too little is known of its population structure and distribution to determine whether the population is declining or merely occupying a limited area of suitable habitat (smuts, pers. comm.). during february and september 1970, 19 days were spent in the park surveying the mountain reedbuck population in the malelane hills. the object of the survey was to obtain information for comparison with the mountain reedbuck population in the loskop dam nature reserve located near groblersdal, transvaal. this paper is a condensation of observations made during the survey. methods mountain reedbuck were located in four ways: 1. scanning hillsides with 7 x 35 binoculars and a 60 x spotting scope; 2. waiting by watering sites at dawn and sunset; 3. walking ridgelines accompanied by a game scout; 4. driving firebreak roads. results only six individual mountain reedbuck were sighted during 52,5 63 hours of observation in february (table 1). all sightings were in the hills along the kwa machiyaliwane drainage even though a comparable amount of time was spent in the vicinity of khandizwe hill and along the emcwebeni drainage. table 1 relative intensity of sampling and success using four techniques for sighting mountain reedbuck in the kruger national park february september observation animals observation animals time sighted time sighted technique (h) (h) spotting scope and binoculars 18,6 0 2,1 3 driving firebreaks 15,0 4 3,9 0 waterhole counts 3,0 0 4,3 0 foot transects 15,9 3 30,0 27 efforts in september were more successful (table 1). of 30 animals sighted, at least 25 were not resightings. group size ranged from one to six. table 2 gives group size, composition, and general habitat information for the september sightings. all of the animals were located on hills above the kwa machiyaliwane drainage in combretum woodland. most of the groups were seen on west-facing slopes on a burned area east of the kwa machiyaliwane streambed. none were more than 5 km from the matjulwana windpump. discussion the low number of sightings recorded in february was probably due to visibility restrictions imposed by the fully-leafed vegetation. several of the sightings in september would not have been made if the trees were not bare. the presence of juveniles in september (table 2) indicates that mountain reedbuck are reproducing in the park and that lambs are surviving until the end of the dry season. although the small sample size precludes any conclusions on the population trends of this antelope in the park, the lamb to ewe ratio in the knp is greater than the lamb to ewe ratio of the relatively stable loskop population for the same period (table 3). 64 table 2 group size, composition, and site characteristics rif mountain reedbuck sighted in the kruger national park during september 1970 habitat group composition characteristics time 1515 0840 1012 1655 0705 0810 0830 0855 0710 group size male female imm. (>1 a) (>1 a) «1 a) 1 3 2 6 1 (whistle only) 3 5 4 1 2 2 2 2 1 2 1 table 3 unaged 4 1 1 1 slope h t. on recent face slope burn w 3/4 yes sw 3/4 yes s 3/4 yes w 1/4 yes w 3/4 yes ? 1/4 yes sw 2/3 yes nw 4/5 yes e 3/4 no comparison rif mountain reedbuck sightings in the loskop dam nature reserve during august-september 1970 with sightings in the kruger national park during september 1970* lamb: ewe n males females lambs unaged ratio (> 1 a) ( > 1 a) ( < 1 a) (lambs per 100 females) loskop 50 14 25 6 5 24 kruger 26 3 10 5 8 50 * groups sighted in intensive study areas at loskop have been excluded. the 24 km 2 section of the kwa machiyaliwane drainage surveyed yielded a minimum count of 25 mountain reedbuck. exclusion of 3 km 2 of flats associated with the main streamcourse gave a minimum density of 1,2 mountain reedbuck per km 2 of slope habitat. a similar treatment of sightings at loskop during june-august 1973 (irby 1973), gave a minimum count of 205 individuals on 49 km 2 for a minimum density of 4,2 per km 2 of slope habitat. while the two counts are not directly 65 comparable, since the search intensity was greater at loskop, the data indicate that the kwa machiyaliwane sub-population is less dense than that of loskop. the absence of sightings around khandizwe hill and along the emcwebeni drainage may indicate a decline in the population, and a subsequent decrease in the range occupied, or only insufficient effort expended to locate individuals in a very sparse population. new growth on burned grass clumps in the kwa machiyaliwane area could have concentrated mountain reedbuck in that region. a mass migration from the emcwebeni drainage to this burn seems unlikely since comparable burns, with no mountain reedbuck, were observed in the emcwebeni drainage, and mountain reedbuck are territorial. acknowledgements i wish to thank dr. u. de v. pienaar and the national parks board of trustees for allowing me to work in the kruger national park and the staff of the park for the guidance and assistance provided during my stay. dr. s. l. beasom deserves thanks for reviewing this manuscript. funding for this study was provided by the mammal research institute, university of pretoria; the caesar kleberg research program in wildlife ecology, texas a & m university; and the national science foundation, washington, d.c. material presented in this paper will be used for partial fulfillment of the requirements for a ph.d. degree from texas a & m university. this manuscript is texas agricultural experiment station technical article no. ta 12059. references irby, l. r. 1973. mountain reedbuck in the transvaal. unpublished annual report of the caesar kleberg research program in wildlife ecology. 108-115. pienaar, u. de v. 1969. predator-prey relationships amongst the larger mammals of the kruger national park. koedoe 12: lo8-176. 66 page 1 page 2 page 3 page 4 abstract introduction methods and materials results discussion conclusion acknowledgements references about the author(s) kelly derham department of biology, western kentucky university, united states michelle d. henley applied behavioural ecology and ecosystem research unit, university of south africa, south africa elephants alive, transboundary research programme, south africa bruce a. schulte department of biology, western kentucky university, united states citation derham, k., henley, m.d. & schulte, b.a., 2016, ‘wire netting reduces african elephant (loxodonta africana) impact to selected trees in south africa’, koedoe 58(1), a1327. http://dx.doi.org/10.4102/koedoe.v58i1.1327 note: this article is partially based on kelly derham’s thesis of the degree of master of science at the department of biology, western kentucky university, kentucky, received may 2014, available here: http://digitalcommons.wku.edu/cgi/viewcontent.cgi?amp=&article=2358&context=theses original research wire netting reduces african elephant (loxodonta africana) impact to selected trees in south africa kelly derham, michelle d. henley, bruce a. schulte received: 05 june 2015; accepted: 24 feb. 2016; published: 30 june 2016 copyright: © 2016. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract african elephants (loxodonta africana) are ecosystem engineers in that they substantially alter the environment through their unique foraging and feeding habits. at high densities, elephants potentially have negative impacts on the environment, specifically for large trees. because of this, recent increases of elephants in the associated private nature reserves (apnr) on the western boundary of the kruger national park (knp), south africa, have caused concern regarding the survival of several tree species. our objective was to assess the effectiveness of wrapping protective wire netting around the trunk of the tree for preventing and reducing bark stripping, branch breaking, and felling by elephants. we assessed 2668 trees – 1352 sclerocarya birrea (marula), 857 acacia nigrescens (knobthorn), and 459 lannea schweinfurthii (false marula) – for elephant impact in the apnr, 1387 (52%) of which had previously been wrapped in protective wire netting (789, 548 and 50, respectively). wire netting was effective in reducing the severity of bark stripping and the relative proportion of trees that were bark stripped. in addition, wire netting had an effect on the level of impact, with a higher relative frequency of wire-net-protected trees found in lower impact categories compared with unprotected trees. since tree mortality has been attributed to high levels of elephant impact, the use of wire netting could serve to maintain individual trees or populations particularly vulnerable to elephant impact in areas with locally high densities of elephants. conservation implications: since wire netting is a relatively low cost and ecologically unobtrusive strategy, it could be used to reduce elephant impact in problem areas. this method focuses on protecting trees rather than some other strategies such as environmental manipulation, translocation, contraceptives, and culling that instead focus on reducing elephant numbers. introduction elephants are allogenic ecosystem engineers (jones, lawton & shachak 1994) in that they substantially modify the environment through their unique foraging and feeding habits (laws 1970). in areas where elephants are confined by fences and human settlements, their population density can increase locally, leading to extensive modification of habitat that can potentially have negative consequences on ecosystem processes and other organisms (dublin & hoare 2004). for example, vogel et al. (2014) proposed that elephants could affect vulture and raptor nesting sites by allowing opportunities for insect and fungal attack after bark stripping, and therefore reducing tree survival. guldemond and van aarde (2008) list a number of studies where high densities of elephants were found to negatively impact woody vegetation, but caution that the findings are site-specific and dependent upon environmental conditions. within the kruger national park (knp), managing elephants in relation to vegetation changes has been a longstanding topic of concern (ferreira et al. 2011; gaylard & ferreira 2011; young, ferreira & van aarde 2009). large trees are of particular concern, because elephants are one of only a few biotic forces that can directly and rapidly modify this key feature of the savanna landscape (asner et al. 2015; laws 1970; morrison, holdo & anderson 2015). large trees play an important role in the biogeochemical cycles of the savanna as well as indirectly affect the distribution of numerous sympatric species that use the trees for refuge, shade, nesting, food, and other activities (bernhard-reversat 1982; nasseri, mcbrayer & schulte 2010; vogel et al. 2014). elephants can affect trees in a variety of ways, including bark stripping, breaking branches, breaking the main stem, or uprooting the entire tree (boundja & midgley 2010; henley 2013). bark stripping and branch breaking expose trees to insect attack and greater damage from fire, both of which may result in tree mortality (helm et al. 2011; vogel et al. 2014). trees are particularly vulnerable to a type of bark stripping known as ringbarking, where the bark is removed around the entire circumference of the tree (helm et al. 2011; ihwagi et al. 2009). tree species differ in their vulnerability to impact because elephants selectively browse for particular species or because of certain characteristics that could predispose a tree to extirpation (boundja & midgley 2010; o’connor, goodman & clegg 2007). boundja and midgley (2010), for example, found that elephants preferred larger trees and therefore targeted larger, less abundant species in hluhluwe-imfolozi park, south africa. elephants are also known to favour trees with certain chemical composition, choosing to browse trees with high levels of leaf nutrients such as calcium, magnesium, potassium, and protein (holdo 2003). although extensive branch breaking and felling can alter the form of a tree, species that readily coppice after such events can survive and continue to grow if their roots remain intact (eckhardt, van wilgen & biggs 2000; henley 2013; ihwagi et al. 2009). however, the continuing loss of habitat for elephants because of expanding human populations and/or fencing compresses their range and confines them to certain areas for unnaturally long periods of time. this often results in continuous and persistent impact that alone or in combination with other factors such as fire or insect attack leads to increased risk of tree mortality (boundja & midgley 2010; mapaure & moe 2009; van aarde, jackson & ferreira 2006). many conservation management plans focus on reducing elephant numbers in order to reduce impact, often assuming a direct linear relationship between the two (van aarde et al. 2006). however, some research has suggested this is not the case and that other factors may be involved (young et al. 2009). environmental manipulation, translocation, contraceptives, and culling have all been suggested and used at various times in south africa’s history to reduce elephant densities (du toit, rogers & biggs 2003; van schalkwyk 2008). alternatively, management could focus on protecting the trees themselves. wrapping wire netting around the trunk of a tree is one such technique that was first employed in kenya (gordon 2003). if wire netting decreases elephant impact in other areas and over the long term, it could serve as a cost-effective and ecologically valuable solution to continuous elephant activity in confined areas. the aim would not be to protect all trees of a certain species and thereby potentially increase the impact on other, unprotected species. instead, the technique can be used to protect individual trees or areas of specific species to ensure there will always be enough mature specimens to populate the surrounding landscape with their seeds. the objective of this study was to assess the effectiveness of wire netting in reducing elephant impact to three relatively abundant, iconic, larger (> 5 m) tree species, namely sclerocarya birrea, acacia nigrescens, and lannea schweinfurthii. wire netting is designed to reduce bark stripping, but we examined all forms of impact by elephants to determine if there was a relationship between wire netting and elephant impact in general. methods and materials study site this study was conducted in the associated private nature reserves (apnr), an area of over 1800 km2 adjacent to knp, south africa (figure 1). together, the apnr and knp encompass an area of more than 20 000 km2 of conserved land and sustain numerous valuable animal and plant species (gertenbach 1983; joubert 1996). the apnr includes balule, klaserie, timbavati, and umbabat private nature reserves, each of which is composed of many private properties that have adopted the overall management plan of the apnr. elephant numbers in the apnr have increased over a 10-year period. in 2002, the population was estimated at 952 individuals. in 2012, 1540 elephants were estimated, indicating an increase in density from 0.52 elephants per km2 to 0.84 elephants per km2 over 10 years (peel 2014). a smaller and slower increase in elephant densities occurred in neighbouring knp (0.40 elephants per km2 to 0.70 elephants per km2) from 1904 to 2002 (whyte, van aarde & pimm 2003). figure 1: map of south africa showing the associated private nature reserves with the individual properties where surveys were conducted. overall, the apnr is characterised as a typical savanna ecosystem with a continuous grass understory and isolated trees (scholes & archer 1997). however, the vegetation within the apnr varies regionally. the eastern areas have dense colophospermum mopane (mopane) woodland as well as isolated a. nigrescens and s. birrea. in other areas combretum apiculatum (red bush willow), l. schweinfurthii and terminalia sericea (silver cluster leaf) occur regularly (de villiers 1994). mean annual rainfall varies around 600 mm and temperatures average 22 °c throughout the year (greyling 2004; venter & gertenbach 1986). tree selection and netting procedure properties in the apnr were selected based on the owner’s interest in participating in the study. in 2004, at the onset of the study, 63 sclerocarya birrea trees were tagged. by 2008, the study was expanded to include six properties, two more species, and 2915 tagged trees, of which 53% had been wrapped in wire netting (figure 1). these trees were resurveyed in 2012 and data collected in that year were used for analysis. trees were selected by species and size as determined by both the height and girdle width of the tree. sclerocarya birrea, a. nigrescens, and l. schweinfurthii trees of greater than 5 m in height were tagged and approximately half were wrapped with wire netting. in addition, trees greater than 2 m in height with similar diameter at breast height (dbh) were included if their height had been previously modified by elephant impact. sclerocarya birrea and a. nigrescens were chosen because residents of the apnr expressed concern about these species in surveys conducted since 2003 in the apnr (henley 2013). in addition, l. schweinfurthii trees are readily accepted as a food source in the dry months by elephants in the area (greyling 2004). wire netting was wrapped around the trunk of the tree (figure 2) about 50 cm off the ground to a height of about 230 cm and secured with fencing staples. on average 1.25 m of wire were used per tree. this amount of wire ensured that most of the area in which elephants primarily remove bark was covered (stokke & du toit 2000). average tusk entry height for trees in this study in 2004 was 120.4 cm ± 35.6 cm (n = 63). prior to using wire, the tree was inspected for any holes left by squirrels, nesting birds, or other animals, and if necessary holes were cut in the wire for these species. wire netting was sometimes applied to trees that were already bark stripped by elephants and new impact was recorded based on the extent of wire-net penetration to access the bark beneath in comparison to previous survey data. figure 2: mesh wire netting (13 mm) around a sclerocarya birrea used to reduce elephant bark stripping. assessment of elephant impact a total of 95 of the trees surveyed in 2008 were located again in 2012. of the 2772 relocated, 57 were dead with the cause of death not directly ascribed to elephants as only remains of the trees were found, and elephant impact from previous years of data collection was not severe enough to have caused death. death in these cases was described as either because of fire where fire scaring was clearly visible or unknown causes. the recent mortality of some trees could be ascribed to wind toppling following storms with strong winds in the study area. only those trees whose dbh could be recorded accurately (2668) were used for analysis (table 1). table 1: the number of trees of each species with average diameter at breast height and cost of wire in the associated private nature reserves, south africa. we used the following separate measures of elephant impact in the field: (1) bark stripping (bs), (2) branch breaking to access smaller plant parts (bba), (3) main stem breaking (ms), and (4) uprooting (ur), as defined by greyling (2004). during analysis, a fifth category, felling (fell), was created by combining the categories ms and ur, which individually had small sample sizes. for each measure of elephant impact, a score between 1 and 10 was given based on its severity as adapted from anderson and walker (1974). in instances where multiple impact types occurred, each event was recorded separately. when two or more instances of the same type of impact were present, the most severe impact was recorded. impact from other animals such as black rhinoceros (diceros bicornis), white rhinoceros (ceratotherium simum), or cape buffalo (syncerus caffer) was differentiated based on the height and type of bark stripping or branch breaking and recorded separately from elephant impact. such impact was rare (26 of the 1281 unprotected, tagged trees [2.0%] had been bark stripped by rhinoceros or cape buffalo when sharpening their horns) and therefore damage from other animals was excluded from analysis. dbh was measured and recorded (in centimetres) for each tree. tree height was also measured in the field but was highly correlated with dbh (r = 0.64, p < 0.001). therefore, dbh was used as a proxy for overall tree size in analyses. for bark stripping and branch breaking, impact scores included 1 (0%), 2 (< 1%), 3 (1% – 5%), 4 (6% – 10%), 5 (11% – 25%), 6 (26% – 50%), 7 (51% – 75%), 8 (76% – 90%), 9 (91% – 99%) and 10 (100%). bark stripping percentage was determined by the proportion of the circumference of the tree that had been bark stripped, which is a method commonly used in other studies also considering this impact type (anderson & walker 1974; helm et al. 2011). for branch breaking, the score was determined based on the percentage of all branches that had been broken by elephants. scores were determined for ms and ur based on whether the tree survived the impact and if any new growth was present. a score of 1 was given to trees with no impact and a score of 10 given to trees with heavy elephant impact, resulting in the death of the tree (henley 2013). scores of 6, 7, 8, and 9 were given in situations where trees had their main stem snapped or had been uprooted to a varying degree by elephants but had not died. these scores were dependent on the amount of the upper stem still attached to the lower stem for main stem breakage and the angle of the uprooted tree for the uprooting category. the variable amounts of regrowth left after the impact incident also contributed to the overall assessment of the score. a further description of each score is provided in greyling (2004) and derham (2012). statistical analysis to determine whether wrapping the tree in wire netting affected the likelihood that a tree would be bark stripped or incur any other type of impact by elephants, log linear analyses were performed. because species are known to vary in their vulnerability to elephant impact, species was used as another factor in the log linear analysis. since size (dbh) was significantly different between wire-net protected and unprotected trees (t-test, t = -17.9, df = 2667, p < 0.001) and among species (anova, f = 23.8, df = 2, 2667, p < 0.01), dbh was used as a covariate to account for the influence size might have on elephant impact. the mean dbh of wire-net-protected trees was 44.0 cm ± 13.5 cm, n = 1387 while the mean dbh of unprotected trees was 33.8 cm ± 14.1 cm, n = 1281. acacia nigrescens were generally the largest (mean dbh = 44.9 cm ± 15.2 cm, n = 857), compared with sclerocarya birrea (37.9 cm ± 13.4 cm, n = 1352) and l. schweinfurthii (32.3 cm ± 14.0 cm, n = 459). any significant results from the log linear analysis were investigated further. to assess differences in size between impacted and intact trees, welch’s t-tests to address unequal variance were performed. g-tests of independence were used to further investigate any differences by species (sokal & rohlf 2012). to assess the effectiveness of wire netting in reducing the severity of elephant bark stripping, a g-test was performed with wire netting (2 levels) and score of bark stripping (10 levels). a g-test also was performed to analyse whether the distribution of overall elephant impact across classes differed between wire-net-protected and unprotected trees. for this analysis the highest impact score was recorded for each tree regardless of whether it came from bark stripping, branch breaking, or felling. for example, if a tree had bark stripping impact of class 3 and uprooting impact of class 8, the overall impact for the tree was recorded as class 8 for this analysis. all statistical tests were performed using r statistical software with a type i error rate of 0.05 (r core team 2014). all assumptions were addressed, and a bonferroni’s correction was used when necessary. results elephant impact was common in this study with 2381 out of 2668 (89%) trees being impacted by elephants in some way or the other (table 2). a total of 74 unprotected trees (5.8%) and 213 protected trees (15%) did not have any elephant impact (table 2). in total, 25% of unprotected trees were bark stripped by elephants, while only 1.7% of wire-net-protected trees were bark stripped by elephants (table 2). dbh was not significant in determining the likelihood of bark stripping (p = 0.08); however, wire netting (p < 0.001) and species (p = 0.03) were significant (table 3). acacia nigrescens had significantly more bark stripping than l. schweinfurthii (g = 36, df = 1, p < 0.001), which had significantly more bark stripping than s. birrea (g = 7.2, df = 1, p < 0.01; figure 3a). there was no significant interaction between species and whether a tree was protected by wire netting, meaning that regardless of species differences, wire netting was effective at reducing the number of trees that were bark stripped (p = 0.75; table 3). table 2: the number and percentage of total trees in each category of elephant impact in the associated private nature reserves, south africa. table 3: results of log linear analysis for the presence of bark stripping, branch breaking to access smaller plant parts, and felling (ms + ur) by elephants in the associated private nature reserves, south africa. figure 3: relative frequency of bark stripping (a) and branch breaking (b) caused by elephant for sclerocarya birrea, acacia nigrescens, and lannea schweinfurthii in the associated private nature reserves, south africa. in addition, the severity of bark stripping was reduced on trees protected by wire netting (g = 55, df = 8, p < 0.001) (figure 4). the number of trees in all impact score categories was reduced with wire netting, and no tree wrapped with wire experienced impact in the highest two categories, 9 and 10 (figure 4). only three trees (0.2%) protected with wire netting were bark stripped more than 50% of the circumference of their trunk, compared with 85 (6.0%) without wire netting. in addition, no tree with wire netting was ring barked (i.e. class 10; 100% bark stripped), compared with 23 unprotected trees. figure 4: the relative frequency of trees in each bark stripping class for wire-net protected and unprotected trees in the associated private nature reserves, south africa. the presence or absence of wire netting was not a significant factor for explaining the occurrence of elephant impact other than bark stripping (branch breaking p = 0.49; felling p = 0.51; table 3; figure 3b). branch breaking was recorded for 69% of unprotected trees and 64% of protected trees (table 2). dbh had a significant effect on the occurrence of branch breaking (p = 0.001; table 3), with smaller trees more likely to have their branches broken by elephants (t = 3.80, df = 1788, p < 0.001). in addition, species was an important factor influencing branch breaking (p < 0.001; table 3). acacia nigrescens had significantly fewer branches broken compared with s. birrea (g = 346, df = 1, p < 0.001) and l. schweinfurthii (g = 208, df = 1, p < 0.001) (figure 3b). there was no significant difference in branch breaking between s. birrea and l. schweinfurthii. dbh was a significant factor in influencing whether a tree had been felled (p < 0.001; table 3). felled trees were significantly smaller in dbh than trees that remained standing (t = 11.3, df = 954, p < 0.01). although 15% of trees without wire netting were felled and only 7% with wire netting suffered from this impact, once dbh was accounted for, species and treatment were not significant factors (table 3). although wire netting was not successful at significantly reducing the proportion of trees that had their branches broken or were felled by elephants, it did have a significant influence on the distribution of the level of impact (g = 73.8, df = 9, p < 0.001; figure 5). trees that were protected with wire netting were more likely to experience lower levels of elephant impact than unprotected trees. sixty two per cent of the time the highest impact recorded was branch breaking. therefore, it appears that wire netting reduced the degree of branch breaking in trees that were impacted in this way by elephants. figure 5: the relative frequency of trees in each impact class where the highest level of impact by elephants was recorded within the associated private nature reserves, south africa. discussion wire netting was effective at reducing the relative frequency and severity of bark stripping by elephants in this study. since high levels of bark stripping are known to affect tree survival (helm et al. 2011; ihwagi et al. 2009), wire netting could prevent mortality by decreasing both the number of trees that are bark stripped by elephants and the occurrence of severe impact. although wire netting did not significantly decrease the frequency of branch breaking or felling, it did reduce the occurrence of high levels of all types of impact. trees that are likely to experience severe elephant impact, specifically ringbarking, main stem breaking, or uprooting are susceptible to mortality (o’connor et al. 2007) and therefore wire netting could help these trees survive in areas with high elephant densities. along with the absence or presence of wire netting, tree species and size were important factors in explaining damage for some measures of impact. dbh did not significantly affect the frequency of bark stripping; however, this could be because most of the tagged trees were of a similar size. if we included a wider range of tree sizes in the study, then it is possible that some sizes of trees would benefit more from wire netting. for example, smallie and o’connor (2000) found that trees taller than 4 m were selected by elephants for bark stripping. further research would be needed to see if smaller trees would also benefit from wire netting and how the wire netting of such trees could accommodate girth expansion over time. acacia nigrescens had high levels of bark stripping suggesting it might be preferred by elephants for this type of impact. physical properties of the bark itself may account for the differences in frequencies of bark stripping between the species (o’connor et al. 2007). in addition, elephants are known to bark strip larger trees and this result might also be because a. nigrescens were significantly larger than s. birrea and l. schweinfurthii in this study (boundja & midgley 2010; smallie & o’connor 2000). since wire netting was effective regardless of species, it could benefit all three of the species assessed in this study. however, it might be particularly useful in reducing elephant bark stripping in preferred species such as a. nigrescens. in contrast to other methods of reducing elephant impact such as culling and altering water-surface availability that focus on reducing elephant numbers or altering spatial distribution patterns, wire netting instead attempts to reduce impact by protecting the trees themselves. some research has used elephant exclosures to study the relative impact of elephants on trees (western & maitumo 2004). however, as a management tool these exclosures also exclude other large herbivores such as giraffe from accessing the trees (goheen & palmer 2010; western & maitumo 2004). in contrast, wire netting maintains the functionality of the trees for other organisms in the ecosystem by allowing them to access the trees for refuge, shade, food, and other activities. wire netting is a relatively low maintenance strategy of reducing elephant impact. in the current system, the wire netting did not need to be replaced from the time it was applied in 2004 until the last survey in 2012 and very little maintenance was required. depending on the scale at which trees are wire-net protected, it could be cost-effective. electric fencing has been used to keep elephants out of specific areas but it is more expensive and can be problematic to maintain (kiokoe et al. 2008). in addition, wire netting wrapped around the trunk of a tree is certainly less visible to tourists than electric fencing. conclusion this study aimed to assess the effectiveness of wire netting in reducing elephant impact on trees in the apnr. we found that wire netting reduced the proportion and severity of elephant impact and contributed to the survival of trees. as elephant densities in the apnr and other areas continue to increase, there will need to be strategies to maintain these iconic and valuable large tree species. in such areas, the use of wire netting could serve to alleviate elephant impact. acknowledgements we thank western kentucky university’s graduate school, office of sponsored programs, and department of biology for funding. we also thank the united states fish and wildlife service for covering aspects of the running expenses of the tree-monitoring programme as well as adam baugh, katherine bunney, amy clark, adam edge, dan hibbet, josh hibbet, hannah malin, francois van der merwe and ellary williams for assisting in the field. we are grateful to the property owners in the apnr. cornelius van der waal is thanked for coordinating the wire netting of the trees on sumatra. the van tienhoven foundation for international nature protection financially supported the wire netting of the trees on this property. elephants alive (formerly known as save the elephants − south africa), transfrontier africa, and tanda tula safari camp are thanked for logistical support. thanks to michael collyer for statistical consultation and r scripts. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions k.d. performed most of the fieldwork and was responsible for data analysis and write up. m.d.h. was responsible for experimental and project design and all field surveys prior to 2012. m.d.h. and b.a.s. made conceptual and editorial contributions. references anderson, g.d. & walker, b.h., 1974, ‘vegetation composition and elephant damage in sengwa wildlife research area’, journal of the south african wildlife management association 4, 1–14. asner, g.p., vaughn, n., smit, i.p.j. & levick, s., 2015, ‘ecosystem-scale effects of megafauna in african savannas’, ecography 38, 1–13. bernhard-reversat, f., 1982, ‘biogeochemical cycles of nitrogen in a semi-arid savanna’, oikos 38, 321–332. http://dx.doi.org/10.2307/3544672 boundja, r.p. & midgley, j.j., 2010, ‘patterns of elephant impact on woody species in the hluhluwe-imfolozi park, kwazulu-natal, south africa’, african journal of ecology 48(1), 206–214. http://dx.doi.org/10.1111/j.1365-2028.2009.01104.x derham, k.d., 2012, ‘wire netting reduces african elephant (loxodonta africana) impact to selected large trees in south africa’, ms thesis, western kentucky university, bowling green, ky. de villiers, p.a., 1994, ‘aspect of the behavior and ecology of elephant (loxodonta africana) in the eastern transvaal lowveld with special reference to environmental interaction’, phd thesis, university of the orange free state, bloemfontein. dublin, h.t. & hoare, r.e., 2004, ‘searching for solutions: the evolution of an integrated approach to understanding and mitigating human-elephant conflict in africa’, human dimensions wildlife 9, 271–278. http://dx.doi.org/10.1080/10871200490505701 du toit, j.t., rogers, k.h. & biggs h.c., 2003, ‘the kruger experience: ecology and management of savanna heterogeneity’, island press, washington, dc. eckhardt, h.c., van wilgen, b.w. & biggs, h.c., 2000, ‘trends in woody vegetation cover in the kruger national park, south africa, between 1940 and 1998’, african journal of ecology 38(2), 108–115. http://dx.doi.org/10.1046/j.1365-2028.2000.00217.x ferreira, s.m., deacon, a., sithole, h., bezuidenhout, h., daemane, m. & herbst, m., 2011, ‘from numbers to ecosystems and biodiversity – a mechanism approach to monitoring’, koedoe 53, 193–204. gadd, m.e., 2002, ‘the impact of elephants on the marula tree sclerocarya birrea’, african journal of ecology 40(4), 328–336. gaylard, a. & ferreira, s.m., 2011, ‘modification and maturation of sanpark’s adaptive planning process – making critical linkages between conservation objectives and actions’, koedoe 53, 58–65. gertenbach, w.p.d., 1983, ‘landscapes of the kruger national park’, koedoe 26, 9–121. http://dx.doi.org/10.4102/koedoe.v26i1.591 goheen, j.r. & palmer, t.m., 2010, ‘defensive plant-ants stabilize megaherbivore-driven landscape change in an african savanna’, current biology 20, 1768–1772. http://dx.doi.org/10.1016/j.cub.2010.08.015 gordon, c.h., 2003, ‘the impact of elephants on the riverine woody vegetation of samburu national reserve’, unpublished report for save the elephants, kenya. greyling, m.d., 2004, ‘sex and age related distinctions in the feeding ecology of the african elephant, loxodonta africana’, phd thesis, university of the witwatersrand, johannesburg. guldemond, r. & van aarde, r.j., 2008, ‘a meta-analysis of the impact of african elephants on savanna vegetation’, journal of wildlife management 72(4), 892–899. http://dx.doi.org/10.2193/2007-072 helm, c., wilson, g., midgley, j., kruger, l. & witkowski, e.t.f., 2011, ‘investigating the vulnerability of an african savanna tree (sclerocarya birrea ssp. caffra) to fire and herbivory’, austral ecology 36(8), 964–973. http://dx.doi.org/10.1111/j.1442-9993.2010.02232.x henley, m.d., 2013, save the elephants vegetation report, unpublished report, save the elephants. holdo, r.m., 2003, ‘woody plant damage by african elephants in relation to leaf nutrients in western zimbabwe’, journal of tropical ecology 19(2), 189–196. http://dx.doi.org/10.1017/s0266467403003213 ihwagi, f.w., vollrath, f., chira, r.m., doulas-hamilton, i. & kironchi, g., 2009, ‘the impact of elephants, loxodonta africana, on woody vegetation through selective debarking in samburu and buffalo springs national reserves, kenya’, african journal of ecology 48(1), 87–95. http://dx.doi.org/10.1111/j.1365-2028.2009.01089.x jones, c.g., lawton, j.h. & shachak, m., 1994, ‘organisms as ecosystem engineers’, oikos 69(3), 373–386. http://dx.doi.org/10.2307/3545850 joubert, s.c.j., 1996, ‘master plan for the management of the associated private nature reserves’, unpublished report. kiokoe, j., muruthi, p., omondi, p. & chiyo, p.i., 2008, ‘the performance of electric fences as elephant barriers in amboseli, kenya’, south african journal of wildlife research 38(1), 52–58. http://dx.doi.org/10.3957/0379-4369-38.1.52 laws, r.m., 1970, ‘elephants as agents of habitat and landscape change in east africa’, oikos 21, 1–15. http://dx.doi.org/10.2307/3543832 mapaure, i. & moe, s.r., 2009, ‘changes in the structure and composition of miombo woodlands mediated by elephants (loxodonta africana) and fire over a 26-year period in north-western zimbabwe’, african journal of ecology 47(2), 175–183. http://dx.doi.org/10.1111/j.1365-2028.2008.00952.x morrison, t., holdo, r.m. & anderson, t.m., 2015, ‘elephant damage, not fire or rainfall, explains mortality of overstory trees in the serengeti’, journal of ecology 104(2), 408–418. http://dx.doi.org/10.1111/1365–2745.12517 nasseri, n.a., mcbrayer, l.d. & schulte, b.a., 2010, ‘the impact of tree modification by african elephant (loxodonta africana) on herpetofaunal species richness in northern tanzania’, african journal of ecology 49(2), 133–140. http://dx.doi.org/10.1111/j.1365-2028.2010.01238.x o’connor, t.g., goodman, p.s. & clegg, g., 2007, ‘a functional hypothesis of the threat of local extirpation of woody plant species by elephant in africa’, biological conservation 136(3), 329–345. http://dx.doi.org/10.1016/j.biocon.2006.12.014 peel, m., 2014, ‘ecological monitoring: association of private nature reserves (timbavati, umbabat, klaserie, and balule)’, unpublished report. r core team, 2014, r: a language and environment for statistical computing, r foundation for statistical computing, vienna, austria, viewed 14 february 2014, from http://www.r-project.org/ scholes, r.j. & archer, s.r., 1997, ‘tree-grass interactions in savanna’, annual review of ecology and systematics 28, 517–544. smallie, j.j. & o’connor, t.g., 2000, ‘elephant utilization of colophospermum mopane: possible benefits of hedging’, african journal of ecology 38(4), 352–359. http://dx.doi.org/10.1046/j.1365-2028.2000.00258.x march 10th, 2014 sokal, r.r. & rohlf, f.j., 2012, biometry: the principles and practice of statistics in biological research, 4th edn., w.h. freeman and co., new york. stokke, s. & du toit, j.t., 2000, ‘sex and size related differences in the dry season feeding patterns of elephants in chobe national park, botswana’, ecography 23, 70–80. http://dx.doi.org/10.1111/j.1600-0587.2000.tb00262.x van aarde, r.j., jackson, t.p. & ferreira, s.m., 2006, ‘conservation sciences and elephant management in southern africa’, south african journal of science 102, 385–388. van schalkwyk, m., 2008, ‘national norms and standards for the management of elephants in south africa’, department of environmental affairs and tourism, government gazette no. 30833, 29 february 2008, 251, pp. 3–39, pretoria, south africa. venter, f.j. & gertenbach, w.p.d., 1986, ‘a cursory review of the climate and vegetation of the kruger national park’, koedoe 29, 125–138. http://dx.doi.org/10.4102/koedoe.v29i1.526 vogel, s.m., henley, m.d., rode, s.c., van de vijyer, d., meares, k.f., simmons, g. & de boer, w.f., 2014, ‘elephant (loxodonta africana) impact on trees used by nesting vultures and raptors in south africa’, african journal of ecology 52(4), 458–465. http://dx.doi.org/10.1111/aje.12140 western, d. & maitumo, d., 2004, ‘woodland loss and restoration in a savanna park: a 20-year experiment’, african journal of ecology 42(2), 111–121. http://dx.doi.org/10.1111/j.1365-2028.2004.00506.x whyte, i.j., van aarde, t. & pimm, s., 2003, ‘kruger national park’s elephant population: its size and consequences for ecosystem heterogeneity’, in j.t. du toit, h. biggs & k. rogers (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 332–348, island press, washington, dc. young, k.d., ferreira, s.m. & van aarde, r.j., 2009, ‘the influence of increasing population size and vegetation productivity on elephant distribution in kruger national park’, austral ecology 34, 329–342. http://dx.doi.org/10.1111/j.1442-9993.2009.01934.x http://www.koedoe.co.za open access koedoe african protected area conservation and science issn: (online) 2071-0771, (print) 0075-6458 page 1 of 1 corrigendum read online: scan this qr code with your smart phone or mobile device to read online. authors: valli-laurente fraser-celin1 alice j. hovorka1,2 mark hovorka3 glyn maude4,5 affiliations: 1department of geography, university of guelph, canada 2department of geography and planning, queen’s university, canada 3private, ottawa ontario, canada 4kalahari research and conservation group, maun, botswana 5conservation and research, denver zoo, united states corresponding author: valli-laurente fraser-celin, vfraserc@uoguelph.ca dates: published: 30 nov. 2017 how to cite this article: fraser-celin, v-l., hovorka, a.j., hovorka, m. & maude, g., 2017, ‘corrigendum: farmer–african wild dog (lycaon pictus) relations in the eastern kalahari region of botswana’, koedoe 59(2), a1508. https://doi. org/10.4102/koedoe. v59i2.1508 copyright: © 2017. the authors. licensee: aosis. this work is licensed under the creative commons attribution license. in the version of this article initially published, mark hovorka’s last name was misspelled as ‘hovork’. the error has been corrected in the pdf version of the article. corrigendum: farmer–african wild dog (lycaon pictus) relations in the eastern kalahari region of botswana read online: scan this qr code with your smart phone or mobile device to read online. note: doi of original article: https://doi.org/10.4102/koedoe.v59i2.1366 http://www.koedoe.co.za http://orcid.org/0000-0003-2399-3415 mailto:vfraserc@uoguelph.ca https://doi.org/10.4102/koedoe.v59i2.1508 https://doi.org/10.4102/koedoe.v59i2.1508 https://doi.org/10.4102/koedoe.v59i2.1508 http://crossmark.crossref.org/dialog/?doi=10.4102/koedoe.v59i2.1508=pdf&date_stamp=2017-11-30 https://doi.org/10.4102/koedoe.v59i2.1366 abstract introduction the biodiversity case against implementation of the eradication proposal the biodiversity of the north-eastern kwazulu-natal is more susceptible to the sequential aerosol technique than that of the okavango delta conclusion acknowledgements references about the author(s) adrian j. armstrong scientific services, ezemvelo kzn wildlife, south africa andy blackmore scientific services, ezemvelo kzn wildlife, south africa citation armstrong, a.j. & blackmore, a., 2017, ‘tsetse flies should remain in protected areas in kwazulu-natal’, koedoe 59(1), a1432. https://doi.org/10.4102/koedoe.v59i1.1432 review article tsetse flies should remain in protected areas in kwazulu-natal adrian j. armstrong, andy blackmore received: 08 aug. 2016; accepted: 20 oct. 2016; published: 30 jan. 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the proposal to eradicate tsetse flies from south africa, including its protected areas, via the sequential aerosol technique combined with the sterile insect technique to reduce trypanosomiasis in cattle did not present an appropriate analysis of the impacts that implementation of the proposal would have on biodiversity. not only would the implementation of the proposal be contrary to south african laws protecting and conserving biodiversity, but it would also have negative consequences for the conservation of biodiversity. some of the negative consequences are reviewed, including extirpations and negative impacts on ecological and ecosystem processes and services. alternative strategies to control trypanosomiasis in cattle effectively in a more environment-friendly manner are presently available and others will almost certainly become available in the not-too-distant future. conservation implications: environmental protection, promotion of conservation and sustainable use of the environment are all deeply seated in south africa’s law. rural livestock husbandry considerations and biodiversity conservation are not mutually exclusive and the importance of one cannot supersede the other. the eradication proposal is seen to be environmentally damaging and therefore it is concluded that the purpose of this proposed eradication exercise is unconstitutional, contrary to various multilateral agreements south africa has entered into and contrary to good environmental governance. introduction tsetse flies are the most important vectors for african animal trypanosomiasis (aat) in cattle. although indigenous wild mammals have resistance to trypanosomes, susceptible cattle develop clinical trypanosomiasis. the eradication of tsetse flies from kwazulu-natal (and therefore south africa) has been proposed by several authors (international atomic energy agency [iaea] 2006; kappmeier green, potgieter & vreysen 2007). the iaea has a technical cooperation project, ‘supporting a feasibility study to eradicate tsetse from southern mozambique, south africa and swaziland’, under the joint food and agriculture organization (fao)/iaea programme, ‘nuclear techniques in food and agriculture’, as part of its peaceful use of nuclear technology theme (iaea n.d.). this eradication proposal (ep) recommends using the sequential aerosol technique (sat) over north-eastern kwazulu-natal (nekzn; figure 1), a region with a number of very important protected areas, including a world heritage site (whs) and five ramsar sites. sat involves blanket spraying an insecticide from the air several times to suppress tsetse fly populations. deltamethrin, a synthetic pyrethroid, at a low concentration of active ingredient, has been proposed as the insecticide of choice (iaea 2006). the sat would be followed by the sterile insect technique (sit) in which millions of sterilised male tsetse flies are released to eradicate the flies through suppression of reproduction. figure 1: protected areas, including a world heritage site and five ramsar sites, and priority areas for protection that would be blanket aerially sprayed with deltamethrin and where millions of sterilised male tsetse flies would be released. although generally believed to have low environmental build-up and greater specificity to invertebrates, deltamethrin is pernicious in aquatic systems. it has been recorded in fish liver, water, and sediment samples in south africa, and it has been found in trace amounts in the ndumo game reserve and tembe elephant park (ansara-ross et al. 2012), both of which are in the proposed spray region. further contamination by deltamethrin of protected areas that are held in public trust for the conservation of biodiversity should not be permitted. deltamethrin poses a significant risk to non-target species, including aquatic insects and macrocrustaceans (ansara-ross et al. 2012). implementation of the ep would have detrimental consequences that are not restricted to tsetse flies and trypanosomes, but will include several non-target species of conservation concern and others of great biodiversity and ecosystem service value. area-wide eradication of tsetse flies in kwazulu-natal and the adjacent part of mozambique and swaziland is still promoted in the scientific literature (e.g. de beer, venter & vreysen 2015). the study by perkins and ramberg (2004) on the impacts of deltamethrin applied via the sat in 2001 and 2002 to eradicate tsetse flies from the okavango delta (od) and the recovery study in 2003 have been used to justify why the ep would be acceptable from an environmental viewpoint. little critique of, or research on, the negative impacts that the eradication course of action would have on biodiversity has been published. hence, this review gives reasons why the ep should not be carried out from a biodiversity conservation viewpoint, and indeed may not be carried out in protected areas. the biodiversity case against implementation of the eradication proposal various south african legislations promulgated to conserve biodiversity would be contravened the consideration and implementation of a strategy to eradicate tsetse flies within a region of south africa, and importantly in existing protected areas, is seen to contravene south africa’s constitutional and environmental laws and some african and global multilateral agreements this country has entered into. the environmental right in the bill of rights in the constitution of south africa affords a right to present and future generations to have, inter alia, protection and conservation of the environment which is devoid of ecological degradation, and that all use of the environment must be ecologically sustainable within a context of justifiable economic and social development. therefore, the ep presents a significant risk of contravening the supreme law of the country. derived from the environmental right, the national environmental management act (no. 107 of 1998) (nema) provides for a set of environmental principles that: serve as guidelines by reference to which any organ of state must exercise any function when taking any decision in terms of this act or any statutory provision concerning the protection of the environment. (republic of south africa government 1998, s. 2, ss. 1) the application of these principles is binding on all organs of state. within these, nema brings into south african law the public trust doctrine which is defined as: the environment is held in public trust for the people, the beneficial use of environmental resources must serve the public interest and the environment must be protected as the people’s common heritage. (republic of south africa government 1998, s. 2, ss. 4o) the remainder of the environmental principles provide the guidance and the tools necessary to ensure that the trust entity (the environment) is not depleted or degraded (blackmore 2015a). the application of the public trust doctrine is further reinforced by section 3 of the national environmental management: biodiversity act (no. 10 of 2004) (nemba) and the national environmental management: protected areas act (no. 57 of 2003) (nempaa) in which both biodiversity and protected areas are held, by the state, in trust for the people of south africa, thereby clearly bringing biodiversity, and in this case the tsetse flies, into the public trust entity (the environment). thus, the state has fiduciary duty to ensure, through all of its organs of state, the protection of the tsetse flies as species within south africa as well as a component of biodiversity occurring within the country’s protected areas (blackmore 2014). the ep is in conflict with the 2003 african convention on the conservation of nature and natural resources (the maputo convention) that requires state parties to undertake measures, inter alia, to avoid or eliminate risks to biodiversity that manifest at species and habitat levels (article x). furthermore, the parties to the convention are to ensure that they, inter alia: to the maximum extent possible, take all necessary measures to ensure that development activities and projects are based on sound environmental policies and do not have adverse effects on natural resources and the environment in general. (article xiv, s. 2.a; iucn 2004) the 1999 southern african development community (sadc) protocol on wildlife conservation and law enforcement requires each state party to apply their fiduciary duty to ensure that its wildlife resources are conserved and used sustainably (southern african development community 1999, article 3, s. 1.). the protocol also requires sadc countries to refrain from causing ‘damage to the wildlife resources of other states or in areas beyond the limits of national jurisdiction’ (southern african development community 1999, article 3, s. 1.). the request or expectation of south africa to eradicate tsetse flies in this country, by neighbouring countries, would conflict with this provision (blackmore 2015b). the ep is also in conflict with the convention on biological diversity and the ramsar and world heritage conventions, to which south africa is a signatory. given that the south african distribution of tsetse flies includes protected areas that are ramsar and whs sites (figure 1), the provisions of these conventions also apply. the global recognition of these sites is founded mainly on the species and habitat assemblages that occur within these protected areas. south africa has an obligation to protect the outstanding universal values of the isimangaliso wetland park whs: it fulfilled, inter alia, the criterion of containing ‘the most important and significant natural habitats for in-situ conservation of biological diversity’ (united nations educational, scientific and cultural organisation 2015, s. ii.d, ss. 77[x]); the prerequisite condition of integrity included the condition that ‘the properties should contain habitats for maintaining the most diverse fauna and flora characteristic of the bio-geographic province and ecosystems under consideration’ (united nations educational, scientific and cultural organisation 2015, s. ii.e, ss. 95). it is thus common cause that the whs would include a complete assemblage of co-evolved animals, including indigenous mammals, tsetse flies and trypanosomes. by entering into these multilateral agreements, and thereafter having applied for both listing and the inscription of these protected areas, the state has fundamentally bound itself to conserving and protecting all indigenous species and habitats that comprise the biodiversity of these sites (section 231 of the constitution). furthermore, these multilateral agreements have also been brought into south africa’s jurisprudence by way of the nema (section 2[4][n]) and the nemba (section 5), and within an array of explicit provisions in the nempaa. the nempaa further requires the state to ensure that the management of a protected area is commensurate with the purpose it was declared as such (blackmore 2015a, 2015b). finally, by way of a schedule to the world heritage convention act (no. 49 of 1999), the convention is explicitly brought into south african law. the entire convention and all provisions of this convention are, therefore, binding on all organs of state. even outside protected areas, the state has a duty to conserve indigenous biodiversity. tsetse flies and trypanosomes are part of biodiversity tsetse flies and trypanosomes are ancient and remarkable organisms, being components of indigenous biodiversity, and have intrinsic existence value. a maximum of about nine final instar larvae are born live per female and so tsetse flies are ‘k-selected’ (nagel 1995), rather like humans and elephants. they are the only insects that live entirely on blood, and they have become model organisms in biology (snyder & rio 2013). trypanosomes are remarkable for their ability to live in two hosts, and to evade the mammalian immune system, mediated through changes in their surface glycoproteins (matthews 2005). the trypanosome has become a model organism in the study of cell biology and genetics. ecosystem health and functioning would be compromised indigenous parasites and their vectors are important components of biodiversity. tsetse flies are eaten by many animals (nagel 1995), and so if they are eradicated, the food-web structure will be altered, which would result in a less stable and resilient animal community. the elimination of a parasite from an ecosystem can strongly affect the interactions between a diverse range of species in the community, both hosts and non-hosts, and hence affect biodiversity (lebarbenchon et al. 2007). parasites such as trypanosomes are an integral part, and are vital to the maintenance, of biodiversity. parasitic species increase the species richness of an area dramatically, and parasites alter the outcomes of competition between, and therefore the relative abundance of, other animal species. parasites increase the number of linkages in food webs and therefore the connectivity and cohesion of the food webs, as well as increase the average number of links per species and the lengths of food chains. these factors increase the stability of food webs and therefore of biotic communities. animals at higher trophic levels are less vulnerable to predators but more vulnerable to parasites. all of these factors cause an increase in ecosystem stability (lafferty, dobson & kuris 2006). parasites can divert host energy towards investment in immune function, reduce reproductive success and increase susceptibility to predators or to stress. many parasites have significant effects on the survival, behaviour, growth and competitive ability of their animal hosts, and therefore play a role in natural population regulation and have an important influence on the flow of energy between various trophic levels. parasites influence speciation through changes in animal life histories and gene flow and by influencing the ways in which species interact. the end result is that parasites increase the health of ecosystems (hudson, dobson & lafferty 2006). the following local example is illustrative of a role of tsetse flies and trypanosomes in ecosystems. r.h.t.p. harris discovered in zululand in the 1920s that shape and shading were important in the attraction of tsetse flies to their hosts. a unicoloured mammal of horizontal, cylindrical shape with its undersurface in shadow attracts tsetse flies; the horizontal cylindrical body profile of a zebra is ‘broken up’ by vertical black and white contrasting stripes, rendering them unattractive to biting flies, and they are seldom bitten. therefore, the characteristic striping of zebras evolved to avoid the attention of biting flies, especially tsetse flies (caro et al. 2014; egri et al. 2012; waage 1981). classic equine symptoms of trypanosomiasis are known in zebras, and the disease may come at a greater fitness cost than for wildebeest (caro et al. 2014; ford 1971; glover 1965; neitz 1931; rurangirwa et al. 1986). therefore, the striped coats of zebras appear to have an adaptive function to reduce the risk of parasitism by trypanosomes. direct pathological effects of trypanosomes on populations of wild mammals will most likely occur when animals are stressed (ford 1971; glover 1965; molyneux 1982), such as in times of drought and poor food availability or high competition with other species (cf. melton 1987). in a prolonged drought situation, reduction in population numbers may be crucial to avoid severe habitat degradation (walker et al. 1987). trypanosomes may play a role in reducing populations of certain species more quickly under these circumstances. non-target organisms would be detrimentally impacted the od study documented that the sat had immediate, significant, measurable impacts on non-target organisms, some of which were long lasting (perkins & ramberg 2004; ramberg et al. 2006). abundance of aquatic invertebrates and terrestrial invertebrates was reduced by up to 50% and 70%, respectively, and the losses of higher aquatic invertebrate taxa and of species in the canopies of various tree species was up to 30% and 40%, respectively, after spraying. there was a significant effect of deltamethrin spray on the abundance and community composition of non-target invertebrate organisms (perkins & ramberg 2004). although recovery of assemblages to near those present pre-spray was documented in the recovery study of 2003, some of the spray-affected families of aquatic organisms remained at reduced levels and 10% of the identified species may have been lost because of spraying. in the terrestrial invertebrate recovery study, the long-term fate of 18 rare species flagged as missing could not be determined. however, only four tree species out of the approximately 1300 vascular plant species and subspecies in the od were sampled and five major invertebrate groups analysed (perkins & ramberg 2004; ramberg et al. 2006). given that many invertebrates are host specific, even at lower taxonomic levels, or are microhabitat specific, the long-term impact of the sat on the total terrestrial invertebrate assemblage of the od could not be determined from such a small sample. little follow-up study was done in the od to determine whether any ‘lost’ species may have been extirpated from the sprayed area. for example, various odonata were severely affected by the aerial spraying and the disappearance of 22 species of dragonflies and damselflies from the delta was plausibly the result of the aerial spraying (ramberg et al. 2006). kipping (2010) recorded some of the ‘lost’ species again in the od, as would be expected for non-endemic mobile species, but some species historically recorded in the od were not recorded after the spraying. kurugundla, kgori and moleele (2012) summarised the impacts of the sat using deltamethrin that was carried out in the kwando–linyanti region in 2006, which included severe detrimental impacts on orthopterans. the biodiversity of the north-eastern kwazulu-natal is more susceptible to the sequential aerosol technique than that of the okavango delta although the proposed number of insecticide sprays is fewer than in the od because of the proposal to use the sit in conjunction with insecticide sprays, severe negative impacts can still be expected if the ep were to be implemented. the od is part of a vast, relatively uniform savanna, and generally the species there are widely distributed. no species were known to be endemic to the od (cowling & hilton-taylor 1997; ramberg et al. 2006). the re-appearance of hardy and mobile species after the spraying could be reasonably expected. in contrast, maputaland is hemmed in by the sea on one side and by mountains on another, and consists of a diverse mosaic of habitats. the maputaland centre of endemism is part of the maputaland–pondoland–albany global biodiversity hotspot (cowling & hilton-taylor 1997; steenkamp et al. 2004). the negative impacts of the sat would likely be greater in maputaland than in the od owing to greater habitat heterogeneity, environmental stability, numbers of localised endemics and perhaps more specific relationships between certain invertebrates and particular host plant species or localised vegetation types. for example, moth species assemblages differ between ndumo game reserve, tembe elephant park, manguzi forest reserve and kosi bay which are all in relatively close proximity (figure 1; staude 2016; see staude 1999 for other comparisons). seventy-three per cent of the total 322 species recorded in these protected areas have only been found in one reserve. conversely, in the northern kalahari, there is very little change in looper moth composition between shakawe and maun, which are about 250 km apart (staude 2016). re-establishment of populations of species that were vulnerable to deltamethrin would depend on nearby sources of immigrants and the lack of barriers (e.g. longley et al. 1997). these conditions would not be fulfilled for maputaland endemics and likely not for species with patchy distributions in a transformed and fragmented landscape. extirpation of non-target organisms in north-eastern kwazulu-natal: extermination of tsetse flies can have repercussions for other elements of biodiversity. predators of tsetse flies may be forced to feed on other prey (nagel 1995) and may be reduced in number. parasites (including trypanosomes) and parasitoids of tsetse flies may be concomitantly exterminated or reduced in number. parasitoids and predators are important for the control of tsetse and other fly populations (glasgow 1963). the mourning bee fly exhyalanthrax lugens, which parasitises tsetse fly pupae, was eradicated from areas of nekzn that were aerially sprayed with ddt even before the pale-footed tsetse fly glossina pallidipes itself was eradicated (fiedler, du toit & kluge 1954; fiedler & kluge 1954). this bee fly did not return to the sprayed areas, nor was it and another parasite of tsetse fly pupae, the abrupt bee fly exhyalanthrax abruptus, recorded at the umkhuze game reserve in an extended survey carried out between 2002 and 2005 (greathead et al. 2006). this suggests that the spraying had long-term impacts on some non-target species and that the control of other tsetse flies may have been affected, perhaps allowing populations of other tsetse fly species to expand (cf. esterhuizen et al. 2005). parasitoids endemic to maputaland (figure 2) will be susceptible to reduction in numbers and eventual elimination, depending on their specificity to the tsetse fly hosts and sensitivity to the sat. ecological processes such as the population control of other flies may therefore be affected. figure 2: two maputaland-endemic bee flies, (a) the mozambique bee fly exoprosopa mozambica (female) and (b) the dull-white-banded bee fly exoprosopa albata (males). additionally, some lepidopteran species disappeared from the sprayed area to the south and east of the lebombo mountains. for example, rhodometra satura (figure 3) has not been seen again in the sprayed areas despite the presence of appropriate habitat, host plants and populations to the north. chionopora tarachodes (figure 3) is one of several other examples of maputaland-endemic looper moths that may be vulnerable to sat (staude 2016). figure 3: (a) saturated vestal moth rhodometra satura and (b) chionopora tarachodes. endemic invertebrates on the red list: some rare, non-endemic invertebrate species were present in the od before the commencement of the sat in 2001, but no threatened endemics (ramberg et al. 2006). in contrast, the critically endangered zulu ambush katydid peringueyella zulu, the endangered zululand black millipede doratogonus zuluensis, the endangered orange wisp damselfly agriocnemis ruberrima, the vulnerable umsingazi sprite damselfly pseudagrion coeleste umsingaziense (figure 4) and the vulnerable blue river crab potamonautes lividus are endemic or near-endemic to nekzn and the wider maputaland region (iucn 2016; samways 2006). most, if not all, of these species would be severely affected were the ep implemented. some rare butterfly species, including the white spotted sapphire iolaus lulua (figure 4), pennington’s white mimic ornipholidotos peucetia penningtoni and zulu buff teriomima zuluana, that complete their whole life-cycles on plants, are also endemic to the same region (eds. mecenero et al. 2013). deltamethrin is known to cause mortality of butterfly larvae at very low doses, including relatively long-term delayed mortality of larvae and pupae through sublethal effects, as well as loss in fitness in survivors (çigli & jepson 1995). other non-endemic red list species occur in nekzn. the threat status of most invertebrate groups in south africa is unknown, but some species have their entire life-cycles in the canopy of trees or on bushes in the region, and these may be susceptible to local extinction if the ep were to be implemented. figure 4: (a) the vulnerable maputaland-endemic umsingazi sprite damselfly pseudagrion coeleste umsingaziense and (b) the rare maputaland-endemic white spotted sapphire iolaus lulua. some examples of other susceptible invertebrates: dragonflies and damselflies, beetles, grasshoppers and crickets, and spiders were some groups negatively affected by the sat in the od (kurugundla et al. 2012; perkins & ramberg 2004; ramberg et al. 2006). owing to the diversity of habitats, nekzn is an important area for odonata (50% of the species in south africa are known to occur there) which contribute to various ecosystem services (hart et al. 2014). implementation of the proposed sat would have negative repercussions for odonata. amongst the beetle families in nekzn, leaf beetles (family chrysomelidae) may be severely affected by the sat. many species in the subfamilies chrysomelinae and cassidinae have traits that make them very vulnerable to heavy mortality or extinction through aerial application of insecticides, including traits such as host specificity (often at plant species or genus level), living exposed on plants for most or all life-cycle stages and winglessness (chaboo 2007; grobbelaar 2016; heron & borowiec 1997; figure 5). not only do leaf beetles contribute to ecosystem functioning through their herbivory, but many good flyers are also pollinators. orthoptera are important herbivores in the savannas (scholes & walker 1993), and at least seven genera of grasshoppers and three genera of crickets have flightless species endemic or near-endemic to maputaland. spiders are very important predators of other invertebrates, and some spiders that live above-ground, including on tree trunks and on grass, are known to be sensitive and even very sensitive to deltamethrin (nagel 1995). maputaland has a high diversity of spiders (e.g. haddad & russell-smith 2009). declines in populations of at least some species of spider, if not extinction, would occur if the ep were to be implemented, and non-lethal effects would affect invertebrate community structure and ecological processes to some degree. figure 5: (a) a pitted flightless leaf beetle iscadida alveolata (chrysomelinae) and (b) the black-punctured tortoise beetle aspidimorpha nigropunctata (cassidinae). the negative effects of factors such as climate change on the persistence of elements of biodiversity may either be added to or multiplied with the negative effects of other factors, such as habitat loss (jewitt et al. 2015a), increased pollution, increased use of pesticides and drought, to cause a greater negative effect than if just one factor was operating on biodiversity. the impact of insecticide spray, acting additively or synergistically with other factors, may push some species towards extinction or lead to local extirpation through decrease in population size, including through various genetic effects in small and isolated populations (e.g. pekin 2013). conservation is required in various parts of the range of a species, particularly for endemic species that occur at low densities and in restricted habitats. negative ecosystem effects are likely negative ecosystem effects in the aquatic environment the use of deltamethrin in the od affected the full spectrum of aquatic habitats and animals, from water surface predators to sediment-dwelling mayflies, with negative impacts to a wide range of non-target aquatic organisms, representing many of the functional feeding groups of aquatic macroinvertebrates (perkins & ramberg 2004). community recovery was affected, with some changes to community structure and function. a good example of impacts from the od study concerns freshwater shrimps caridina species that were severely negatively affected by the deltamethrin applications and showed poor recovery. freshwater shrimps (figure 6) are important in various freshwater lakes in nekzn: they are the dominant zoobenthic species of the littoral zone of lake sibaya in terms of biomass, and occur in nearly every marginal habitat, and are abundant in various habitats at lake mgobezeleni (bruton 1980; hart 1979, 1980). these detritivores will be important in the process of the mineralisation of detritus. lake sibaya and many of the wetlands in nekzn are closed systems, being rain fed and with drainage internal to the region, so the shrimps will not be able to re-colonise them from elsewhere should the ep be implemented. figure 6: freshwater shrimp caridina species. in lake sibaya and in pans on the pongolo river floodplain, caridina was found to be a main food in the diet of the tank goby glossogobius giurus, manyspined climbing perch ctenopoma multispine and smaller sharp-tooth catfish clarias gariepinus, and together with aquatic insects form part of the diet of other fish such as young tigerfish hydrocynus vittatus, silver catfish schilbe intermedius, snake catfish clarias theodore, imberi alestes imberi, southern redbreast tilapia tilapia rendalli and adult mozambique tilapia oreochromis mossambicus (bruton 1979; bruton & kok 1980; kok 1980). deltamethrin is known to be toxic to even hardy fish, and can cause mortality of, or non-lethal effects in, fish (e.g. huang et al. 2014). cascading negative ecosystem effects might occur in nekzn should the ep be implemented. impact on livelihoods of poor rural people: fish are important in the diets of poor rural people in maputaland, and the fish species mentioned in the preceding paragraph are important for the local fishermen (bruton & kok 1980; coetzee et al. 2015; kok 1980). the livelihoods of these poor rural people in the spray area would likely be negatively impacted were the ep to be implemented, through impact on the prey base of fish species that are important in their diets. actions that reduce the provision of ecosystem services must be properly weighed up against the benefits of the actions. negative ecosystem effects in the terrestrial environment invertebrates play very important roles in the functioning of ecosystems. implementation of the ep will have impacts on ecological processes and ecosystem services, as illustrated by the selected examples below. nutrient (carbon, nitrogen and phosphorus) cycling from leaves to the soil: herbivorous invertebrate communities play very important roles in the recycling of carbon, nitrogen and phosphorus from vegetation to the soil, making these nutrients more readily available to plants and other micro-organisms than would otherwise happen (metcalfe et al. 2013). the detrimental impacts that the ep would have on insect herbivore communities could negatively affect ecosystem carbon sequestration and soil fertility, particularly in sub-tropical forests and grasslands, at least in the short term. scholes and walker (1993) reported that leaf-feeding grasshoppers and lepidopteran larvae removed about 5% of the foliar primary production in an infertile, sub-tropical, broad-leafed savanna, and a greater proportion of foliar primary production in fertile sub-tropical, fine-leafed savanna. this biomass is returned to the soil as faeces, bodies and cut pieces of leaves that are more easily processed by detritivores and bacteria than abscised leaves. the plant nutrients are thereby more efficiently recycled to the soil to assist plant growth. pollination of flowering plants: reduction in pollination efficiency may occur if non-target pollinators (such as bees, bee flies, beetles and tabanid flies) are reduced in number or extirpated. pollination limitation may lead to a reduction in or absence of recruitment of certain plant species through poor or absent seed production (potts et al. 2010), a reduction in the preferred food supply of certain herbivores, and so on. some wild pollinators, particularly bees, have been declining globally, in part owing to the lethal and sublethal effects of insecticides, which may act synergistically with other drivers such as habitat loss and disease (potts et al. 2010). an estimated 94% of plant species in the latitudinal band 29° – 0° are biotically pollinated, mainly by insects (ollerton, winfree & tarrant 2011), and the impact on the pollination of some of the plant species of maputaland could be severe if the ep were to be implemented. vamosi et al. (2006) show that animal-pollinated plants in plant hotspots have reduced seed-sets owing to greater pollen-limitation than elsewhere, and human-mediated impacts on pollinators may eventually lead to the extinction of plant species. bees may be killed or disabled by deltamethrin, depending on the genus or species of bee (desneux, decourtye & delpuech 2007; scott-dupree, conroy & harris 2009). other pollinating insect groups have not been studied the way bees have been, but the same general impacts are likely. toxicity of a particular insecticide to different pollinators can vary markedly. however, scott-dupree et al. (2009) found that deltamethrin is highly toxic to alfalfa leaf-cutting bees megachile rotundata, an important indigenous north american pollinator of native plants and cultivated crops. at least 20 species of megachile bees occur in nekzn (eardley 2013 and references therein) and deltamethrin may be highly toxic to them. parasitism and predation: parasitoid and predator insects are important for controlling insect populations. deltamethrin is known to have lethal and a wide variety of sublethal effects on insect parasitoids and predators (e.g. desneux et al. 2007; longley et al. 1997). a small change in the reproductive potential of parasitoids can disrupt biological control and make it less successful. deltamethrin can act synergistically with another pesticide in terms of detrimental effects (desneux et al. 2007). ecosystem services relied on by poor rural people: implementation of the ep would affect ecosystem services needed by, or important to, poor rural people. poor rural communities in the proposed spray region are reliant on indigenous plants to maintain health and treat disease, and use indigenous plants and other vegetables and fruits as food (cunningham 1988a, 1988b; ellis, myers & ricketts 2015). many of these are insect-pollinated and a variety of wild insect pollinators are vital for good fruit production in many pollinator-dependent vegetable and fruit crops (garibaldi et al. 2011, 2013). managed honeybee pollination cannot replace wild insect pollination for many of these crops. predatory insects and insect parasitoids that assist with the control of pests of crops (e.g. samways 1988), including those of poor rural people, would be adversely affected by broad-scale sat using deltamethrin, with likely detrimental impacts on crop production, food security and traditional medicine resources. crop production may be affected where poor rural people rely on ecosystem services rather than fertilisers for soil fertility. many rural people in nekzn rely directly or indirectly on activities associated with wildlife for income (aylward & lutz 2003). protected areas are reservoirs of natural resources, including invertebrates that mediate ecosystem processes and services. biological control: alien plant biological control projects in nekzn might be negatively affected were the ep implemented. one of the probable reasons why nekzn does not have great problems with feral and alien invasive mammals is that these mammals are susceptible to diseases such as trypanosomiasis. if trypanosomes, and in future other disease-causing parasites and their vectors, are deliberately exterminated in nekzn, feral and alien invasive mammal problems would likely increase, with corresponding increased impacts on indigenous species. various domesticated animals that have the ability to become feral are susceptible to trypanosomiasis (nwoha 2013), and when controlling factors for those species are reduced or eliminated, the feral species are released from population control and can cause changes in community structure and function, including extinctions of indigenous species (hollings et al. 2016). controlling feral and alien mammals would be costly. stable host-parasite relationships: eradication of tsetse flies and certain trypanosome species would disrupt the stable host–parasite relationship between the trypanosomes and the native mammalian wildlife. should these tsetse flies and trypanosomes return to the region after a certain period, severe clinical trypanosomiasis would be likely in wild mammals. immunity against local trypanosomes but not against foreign trypanosomes is a known phenomenon (ford 1971). south african rhinos that were translocated to areas where different trypanosomes were encountered, and rhinos brought into maputaland from areas that do not have trypanosomes, suffered from trypanosomiasis. some died and others had to be treated. also, if tsetse fly parasites have been eliminated, tsetse fly populations could build up rapidly again. the reasoning that tsetse flies would not come back to the region if the whole population was eradicated can also be used to support the claim that the eradication would then be unnecessary as neither would the population expand northwards. the eradication of trypanosomes without proper cattle population controls would likely exacerbate the destruction of biodiversity forty-four vegetation types occur in the approximate area of occurrence of tsetse flies in kwazulu-natal. ten of these vegetation types are classed as critically endangered, eight as endangered, five as vulnerable and 21 as least threatened. targets for protection have been reached in only 14 of the 44 vegetation types; two of the critically endangered vegetation types are not protected. existing protected areas fall short of conserving a representative sample of the diversity of the region, and accordingly emphasise the importance of areas outside protected areas for meeting south africa’s and mozambique’s and swaziland’s conservation commitments (smith et al. 2008). the rate of land transformation is high at an average of 1.2% of kwazulu-natal per annum between 1994 and 2011 (jewitt et al. 2015b). therefore, reaching the conservation targets in the future is becoming less likely. risks associated with tsetse fly eradication include increasing cattle and other livestock numbers, leading to a reduction of vegetation cover, increasing runoff and erosion, increased pressure on conserved areas, increased transmission of livestock diseases to wild mammals and vice versa, and reduction in biodiversity and increase in alien plants through overgrazing. contrary to the ep (iaea 2006), the argument for not eradicating tsetse flies has in fact become stronger in this respect. environment-friendly methods and options for the control of trypanosomiasis in cattle alternative options currently exist to reduce the incidence and prevalence of trypanosomiasis in cattle without causing extinction of indigenous species, and other options will be available in the future. normal veterinary measures that include trypanosomiasis surveillance, dipping of cattle using appropriate dips or else the use of pour-ons and treatment of infected cattle with trypanocides have proved effective in the past (kappmeier, nevill & bagnall 1998). trypanosomiasis is not the only, nor the most important, disease in cattle in nekzn. tick-borne diseases also have to be controlled using the same methods. decentralised control of animal husbandry at the farmer level using environment-friendly methods could be the ultimate goal. rather than area-wide eradication, creation of local grazing areas free of tsetse flies would assist the farmer and would be more environment-friendly. for many centuries, communities practicing agriculture have been able to reside successfully in areas where tsetse flies existed via local adaptations in the ways the communities formed and lived (e.g. anderson et al. 2015; ford 1971). colonialism and the impacts of its practices and pastoralist migrations often had adverse impacts on these ways of life (e.g. anderson et al. 2015; ford 1971). the consequences of the gradual encroachment of people and cattle into an area where tsetse flies occur on the epidemiology of bovine trypanosomiasis are well known (molyneux 1982; van den bossche 2001). hunting of indigenous mammals and continuous clearing of vegetation eventually removed the hosts and breeding habitat of the tsetse flies, which then disappeared from the area. this enabled farming with crops and cattle to occur even within the tsetse belts (ford 1971; kent 1926; van den bossche 2001). trypanosomiasis in cattle is likely to be more severe where the cattle are close to the boundaries of protected areas or other areas of suitable tsetse fly habitat with wild mammal hosts (de beer et al. 2016; van den bossche 2001). glossina austeni has a short flight distance and is predominantly restricted to forest in kwazulu-natal (esterhuizen et al. 2005). keeping cattle away from forest or bush would greatly reduce the incidence of trypanosomiasis in cattle. this could be achieved by grazing cattle on fenced pastures and by keeping cattle out of protected area buffers or away from protected area boundaries. trypanosomiasis in cattle is more severe where poor nutrition or other stress factors are present (ford 1971; holmes 2013). improvement in the immune system and general health of cattle will occur with improvement in the condition of grazing lands and maintenance of these in good condition through rehabilitation of grazing lands, adherence to appropriate stocking and land resting rates, and implementation of an integrated land-use plan with designated grazing areas (e.g. pastures) where tsetse flies cannot breed. traps and targets are effective for tsetse flies in kwazulu-natal (esterhuizen et al. 2006; kappmeier & nevill 1999), and so could be used to good effect on cattle grazing lands that are within the normal flight range of tsetse flies, especially glossina brevipalpis. certain species of trypanosomes such as trypanosoma vivax are transmitted via the proboscis of blood-feeding flies, including tsetse flies, horse flies (family tabanidae) and stable flies (genus stomoxys). so the eradication of tsetse flies alone will not prevent trypanosomiasis in cattle (ahmed et al. 2016). replacing susceptible cattle by cattle with resistance to trypanosomes in a recapitalisation programme would be fruitful (department of agriculture 1950; holmes 2013). maintenance of heterozygosity by rotation of breeding bulls should be practiced to decrease susceptibility of calves to transmittable diseases such as trypanosomiasis (murray et al. 2013). maintenance of physiological adaptation of the cattle to trypanosomes, as occurred in nguni cattle (department of agriculture 1950), would be required. stress may cause cattle to lose resistance, so it may be necessary to use other means than cattle for ploughing. tractors are alternatives to cattle for draught power and are preferred in terms of modernisation of agriculture. in addition, modern agricultural thinking emphasises the value of no-till farming which has grown remarkably in kwazulu-natal because of its benefits for the farmer and carbon storage. cattle coat colour selection could be used to good effect. white horse models are very unattractive to tabanid flies (egri et al. 2012), and blue and black are most attractive to tsetse flies in nekzn (kappmeier & nevill 1999). if the majority of nguni cattle near the boundaries of protected areas had light coats, the incidence of trypanosomiasis may be reduced further. white-coloured cattle were kept by the king of the mtetwa clan, king dingiswayo, in the imfolozi game reserve area before it was proclaimed a reserve (foster 1955). perhaps this was a local adaptation to the presence of tsetse flies and horse flies, the white coats of the cattle not being attractive to them. maputaland is not south africa’s major beef-producing region, and nature tourism plays a major economic and development role in nekzn (eds. aylward & lutz 2003). the dependence on cattle as a monetary system could be reduced through diversifying income streams, including small business development around wildlife or the green economy. environmental costs (many of which are not financial) have not been quantified or incorporated into a cost–benefit analysis in terms of the ep. further, opportunity costs, depending on what are the pressing social needs, and costs and benefits of alternative management strategies would also have to be included. a significant contribution to livestock productivity could be achieved by investing in the control of invasive alien plants that are invading and reducing the grazing capacity in nekzn, such as famine weed parthenium hysterophorus. with the sequencing of the trypanosome genome complete, and genetic research on trypanosomes proceeding well, there is a good chance that biochemical and molecular genetic techniques can be used to prevent or reduce trypanosomiasis in cattle in the future (e.g. caljon, de vooght & van den abbeele 2013). the development of vaccines could be one outcome. owing to the great strides that are being made in understanding the biology of trypanosomes, long-lasting environment-friendly solutions for the prevention of trypanosomiasis in cattle are likely. conclusion the proposed mass killing of invertebrates, including parasites, as mooted in the tsetse fly ep, would be illegal and detrimental to the biodiversity of south africa, as was the destruction of ‘game’ in the middle of the 20th century. eradication of tsetse flies from protected areas and the use of aerial spraying of insecticides over vast regions is no longer an option in a country that prides itself on its conservation principals with innovative and advanced environmental protection laws. a total ban on insecticide use and the sit is not what is being proposed here, but rather conservative and environmentally judicial use where needed at local grazing land scales, if appropriate. a variety of techniques are now available to combat trypanosomiasis in cattle, and in future others should become available. the more environment-friendly ones should be prioritised for use at the grazing land scale. acknowledgements we thank ezemvelo kzn wildlife for assisting with the research and publication of this work. the ideas, arguments and opinions expressed in this manuscript are that of the authors and do not necessarily represent those of ezemvelo kzn wildlife. we thank the following people for their valuable contributions to this article: beth grobbelaar for information relating to leaf beetles and for comments on the manuscript, camille lebarbenchon for information on the role of parasites in ecosystems, hermann staude for information on and photographs of looper moths in north-eastern kwazulu-natal, nick rivers-moore for assistance with aquatic aspects of the article, roger porter for information on world heritage site matters, and michael samways and steve woodhall for permitting the use of their photographs. we also thank our colleagues debbie jewitt, dave cooper, ian rushworth and john craigie for information on vegetation types, trypanosomes in non-resistant rhinos, alien plant invasion and for providing a photograph, respectively. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions a.j.a. was the project leader and conceptualised, designed and wrote the review. a.b. contributed the legal section. references ahmed, s.k., rahman, a.h., hassan, m.a., salih, e.m., paone, m. & cecchi, g., 2016, ‘an atlas of tsetse and bovine trypanosomosis in sudan’, parasites & vectors 9, 194–201. http://dx.doi.org/10.1186/s13071-016-1485-6 anderson, n.e., mubanga, j., machila, n., atkinson, p.m., dzingirai, v. & welburn, s.c., 2015, ‘sleeping sickness and its relationship with development and biodiversity conservation in the luangwa valley, zambia’, parasites & vectors 8, 224. http://dx.doi.org/10.1186/s13071-015-0827-0 ansara-ross, t.m., wepener, v., van den brink, p.j. & ross, m.j., 2012, ‘pesticides in south african fresh waters’, african journal of aquatic science 37(1), 1–16. http://dx.doi.org/10.2989/16085914.2012.666336 aylward, b. & lutz, e. (eds.), 2003, nature tourism, conservation, and development in kwazulu-natal, south africa, the world bank, washington, dc. blackmore, a., 2014, ‘the interplay between the public trust doctrine and biodiversity and cultural resource legislation in south africa: the case of the shembe church worship site in tembe elephant park in kwazulu-natal’, law, environment and development journal 10(1), 1–15. blackmore, a., 2015a, ‘the relationship between the nema and the public trust doctrine: the importance of the nema principles in safeguarding south africa’s biodiversity’, south african journal of environmental law and policy 20(2), 89–118. blackmore, a., 2015b, ‘legal and public trust considerations for the ndumo game reserve and south africa-mozambique border, following the migration of the usuthu river’, journal of southern african public law 30(2), 347–379. bruton, m.n., 1979, ‘the fishes of lake sibaya’, in b.r. allanson (ed.), lake sibaya, pp. 162–245, dr w. junk bv publishers, the hague. (monographiae biologicae 36). bruton, m.n., 1980, ‘an outline of the ecology of the mgobezeleni lake system at sodwana, with emphasis on the mangrove community’, in m.n. bruton & k.h. cooper (eds.), studies on the ecology of maputaland, pp. 408–426, rhodes university, grahamstown. bruton, m.n. & kok, h.m., 1980, ‘the freshwater fishes of maputaland’, in m.n. bruton & k.h. cooper (eds.), studies on the ecology of maputaland, pp. 210–244, rhodes university, grahamstown. caljon, g., de vooght, l. & van den abbeele, j., 2013, ‘options for the delivery of anti-pathogen molecules in arthropod vectors’, journal of invertebrate pathology 112(suppl. 1), 75–82. http://dx.doi.org/10.1016/j.jip.2012.07.013 caro, t., izzo, a., reiner, r.c., walker, h. & stankowich, t., 2014, ‘the function of zebra stripes’, nature communications 5, 3535. http://dx.doi.org/10.1038/ncomms4535 chaboo, c., 2007, ‘biology and phylogeny of cassidinae gyllenhal (tortoise and leaf-mining beetles) (coleoptera: chrysomelidae)’, bulletin of the american museum of natural history 305, 1–250. http://dx.doi.org/10.1206/0003-0090(2007)305[1:bapotc]2.0.co;2 çigli, t. & jepson, p.c., 1995, ‘the risks posed by deltamethrin drift to hedgerow butterflies’, environmental pollution 87, 1–9. coetzee, h.c., nell, w., van eeden, e.s. & de crom, e.p., 2015, ‘artisanal fisheries in the ndumo area of the lower phongolo river floodplain, south africa’, koedoe 57(1), art. #1248, 1–6. http://dx.doi.org/10.4102/koedoe.v57i1.1248 cowling, r.m. & hilton-taylor, c., 1997, ‘phytogeography, flora and endemism’, in r.m. cowling, d.m. richardson & s.m. pierce (eds.), vegetation of southern africa, pp. 43–61, cambridge university press, cambridge. cunningham, a.b., 1988a, ‘an investigation of the herbal medicine trade in natal/kwazulu’, investigational report no. 29, institute of natural resources, university of natal, pietermaritzburg. cunningham, a.b., 1988b, ‘collection of wild plant foods in tembe thonga society: a guide to iron age gathering activities?’, annals of the natal museum 29(2), 433–446. de beer, c.j., venter, g.j., kappmeier green, k., esterhuizen, j., de klerk, d.g., ntshangase, j. et al., 2016, ‘an update of the tsetse fly (diptera: glossinidae) distribution and african animal trypanosomosis prevalence in north-eastern kwazulu-natal, south africa’, onderstepoort journal of veterinary research 83(1), a1172. http://dx.doi.org/10.4102/ojvr.v83i1.1172 de beer, c.j., venter, g.j. & vreysen, m.j.b., 2015, ‘determination of the optimal mating age of colonized glossina brevipalpis and glossina austeni using walk-in field cages in south africa’, parasites & vectors 8, 467–476. http://dx.doi.org/10.1186/s13071-015-1073-1 department of agriculture, 1950, ‘nguni-cattle’, report on indigenous cattle in south africa, agricultural research institute series no. 22, bulletin no. 311, the government printer, pretoria. desneux, n., decourtye, a. & delpuech, j.-m., 2007, ‘the sublethal effects of pesticides on beneficial arthropods’, annual review of entomology 52, 81–106. http://dx.doi.org/10.1146/annurev.ento.52.110405.091440 eardley, c., 2013, ‘a taxonomic revision of the southern african leaf-cutter bees, megachile latreille sensu stricto and heriadopsis cockerell (hymenoptera: apoidea: megachilidae)’, zootaxa 3601(1), 1–133. http://dx.doi.org/10.11646/zootaxa.3601.1.1 egri, á., blahó, m., kriska, g., farkas, r., gyurkovszky, m., åkesson, s. et al., 2012, ‘polarotactic tabanids find striped patterns with brightness and/or polarization modulation least attractive: an advantage of zebra stripes’, the journal of experimental biology 215, 736–745. http://dx.doi.org/10.1242/jeb.065540 ellis, a.m., myers, s.s. & ricketts, t.h., 2015, ‘do pollinators contribute to nutritional health?’, plos one 10(1), e114805. http://dx.doi.org/10.1371/journal.pone.0114805 esterhuizen, j., kappmeier green, k., marcotty, t. & van den bossche, p., 2005, ‘abundance and distribution of the tsetse flies, glossina austeni and g. brevipalpis, in different habitats in south africa’, medical and veterinary entomology 19, 367–371. http://dx.doi.org/10.1111/j.1365-2915.2005.00582.x esterhuizen, j., kappmeier green, k., nevill, e.m. & van den bossche, p., 2006, ‘selective use of odour-baited, insecticide-treated targets to control tsetse flies glossina austeni and g. brevipalpis in south africa’, medical and veterinary entomology 20, 464–469. http://dx.doi.org/10.1111/j.1365-2915.2006.00650.x fiedler, o.g.h., du toit, r. & kluge, e.b., 1954, ‘the influence of the tsetse fly eradication campaign on the breeding activity of glossinae and their parasites in zululand’, onderstepoort journal of veterinary research 26, 389–397. fiedler, o.g.h. & kluge, e.b., 1954, ‘the parasites of tsetse flies in zululand with special reference to the influence of the hosts upon them’, onderstepoort journal of veterinary research 26, 399–404. ford, j., 1971, the role of the trypanosomiases in african ecology. a study of the tsetse fly problem, clarendon press, oxford. foster, w.e., 1955, history of the umfolozi game reserve, unpublished report, natal parks, game and fish preservation board, pietermaritzburg. garibaldi, l.a., aizen, m.a., klein, a.m., cunningham, s.a. & harder, l.d., 2011, ‘global growth and stability of agricultural yield decrease with pollinator dependence’, pnas 108, 5909–5914. http://dx.doi.org/10.1073/pnas.1012431108 garibaldi, l.a., steffan-dewenter, i., winfree, r., aizen, m.a., bommarco, r., cunningham, s.a. et al., 2013, ‘wild pollinators enhance fruit set of crops regardless of honey bee abundance’, science 339, 1608–1611. http://dx.doi.org/10.1126/science.1230200 glasgow, j.p., 1963, the distribution and abundance of tsetse, pergamon press, oxford. glover, p.e., 1965, review and recent knowledge on the relationship between the tsetse fly and its vertebrate hosts, fauna protection society, london. greathead, d., lovell, s., barraclough, d., slotow, r., hamer, m. & herbert, d., 2006, ‘an ecological and conservation assessment of the fauna of bombyliidae (diptera) occurring in the mkuze, phinda and false bay reserves, kwazulu-natal, south africa’, african invertebrates 47, 185–206. grobbelaar, e., 2016, biology of chrysomelidae: cassidinae, chrysomelinae and galerucinae, unpublished report to ezemvelo kzn wildlife, pietermaritzburg. haddad, c.r. & russell-smith, a., 2009, ‘spider family composition and species richness patterns in two savannah habitats along the eastern coastal plain of africa’, african journal of ecology 48, 418–427. http://dx.doi.org/10.1111/j.1365-2028.2009.01128.x hart, l.a., bowker, m.b., tarboton, w. & downs, c.t., 2014, ‘species composition, distribution and habitat types of odonata in the isimangaliso wetland park, kwazulu-natal, south africa and the associated conservation implications’, plos one 9(3), e92588. http://dx.doi.org/10.1371/journal.pone.0092588 hart, r.c., 1979, ‘the invertebrate communities: zooplankton, zoobenthos and littoral fauna’, in b.r. allanson (ed.), lake sibaya, pp. 108–161, dr w. junk bv publishers, the hague. (monographiae biologicae). hart, r.c., 1980, ‘the aquatic invertebrates of lake sibaya’, in m.n. bruton & k.h. cooper (eds.), studies on the ecology of maputaland, pp. 114–122, rhodes university, grahamstown. heron, h. & borowiec, l., 1997, ‘host plants and feeding patterns of some south african tortoise beetles (coleoptera: chrysomelidae: cassidoid hispinae)’, genus 8, 625–658. hollings, t., jones, m., mooney, n. & mccallum, h., 2016, ‘disease-induced decline of an apex predator drives invasive dominated states and threatens biodiversity’, ecology 97(2), 394–405. holmes, p., 2013, ‘tsetse-transmitted trypanosomes – their biology, disease impact and control’, journal of invertebrate pathology, 112(suppl. 1), 11–14. http://dx.doi.org/10.1016/j.jip.2012.07.014 huang, y., zhang, j., han, x. & huang, t., 2014, ‘the use of zebrafish (danio rerio) behavioral responses in identifying sublethal exposures to deltamethrin’, international journal of environmental research and public health 11, 3650–3660. http://dx.doi.org/10.3390/ijerph110403650 hudson, p.j., dobson, a.p. & lafferty, k.d., 2006, ‘is a healthy ecosystem one that is rich in parasites?’, trends in ecology and evolution 21, 381–385. http://dx.doi.org/10.1016/j.tree.2006.04.007 international atomic energy agency (iaea), 2006, feasibility study – the creation of a tsetse-free zone in the republic of south africa and in the southern part of the republic of mozambique, international atomic energy association, vienna. international atomic energy agency (iaea), n.d., supporting a feasibility study to eradicate tsetse from southern mozambique, south africa and swaziland, viewed 17 october 2016, from https://www.iaea.org/projects/tc/raf5069 iucn, 2004, an introduction to the african convention on the conservation of nature and natural resources, iucn, gland, switzerland and cambridge, uk, viewed n.d., from https://portals.iucn.org/library/sites/library/files/documents/eplp-056.pdf iucn, 2016, the iucn red list of threatened species. version 2016-1, viewed 04 august 2016, from http://www.iucnredlist.org jewitt, d., erasmus, b.f.n., goodman, p.s., o’connor, t.g., hargrove, w.w., maddalena, d.m. et al., 2015a, ‘climate-induced change of environmentally defined floristic domains: a conservation based vulnerability framework’, applied geography 63, 33–42. http://dx.doi.org/10.1016/j.apgeog.2015.06.004 jewitt, d., goodman, p.s., erasmus, b.f.n., o’connor, t.g. & witkowski, e.t.f., 2015b, ‘systematic land-cover change in kwazulu-natal, south africa: implications for biodiversity’, south african journal of science 111(9/10), art. #2015-0019, 1–9. http://dx.doi.org/10.17159/sajs.2015/20150019 kappmeier, k. & nevill, e.m., 1999, ‘evaluation of coloured targets for the attraction of glossina brevipalpis and glossina austeni (diptera: glossinidae) in south africa’, onderstepoort journal of veterinary research 66, 291–305. kappmeier, k., nevill, e.m. & bagnall, r.j., 1998, ‘review of tsetse flies and trypanosomosis in south africa’, onderstepoort journal of veterinary research 65, 195–203. kappmeier green, k., potgieter, f.t. & vreysen, m.j.b., 2007, ‘a strategy for an area-wide control campaign with an sit component to establish a tsetse-(glossina austeni and glossina brevipalpis) free south africa’, in m.j.b. vreysen, a.s. robinson & j. hendrichs (eds.), area-wide control of insect pests, pp. 309–323, iaea, vienna. kent, c.c., 1926, ‘field notes on the mfolozi game reserve’, south african journal of natural history 6, 38–48. kipping, j., 2010, ‘the dragonflies and damselflies of botswana – an annotated checklist with notes on distribution, phenology, habitats and red list status of the species (insecta: odonata)’, mauritiana (altenburg) 21, 126–204. kok, h.m., 1980, ‘ecological studies of some important fish species of the pongolo floodplain, kwazulu, south africa’, phd thesis, university of kwazulu-natal, pietermaritzburg. kurugundla, c.n., kgori, p.m. & moleele, n., 2012, ‘management of tsetse fly using insecticides in northern botswana’, in f. perveen (ed.), insecticides – pest engineering, pp. 449–476, intech, rijeka, viewed 12 august 2016, from http://www.intechopen.com/books/insecticides-pest-engineering/management-of-tsetse-fly-using-insecticides-in-northern-botswana lafferty, k.d., dobson, a.p. & kuris, a.m., 2006, ‘parasites dominate food web links’, pnas 103, 11211–11216. http://dx.doi.org/10.1073/pnas.0604755103 lebarbenchon, c., poulin, r., gauthier-clerc, m. & thomas, f. 2007, ‘parasitological consequences of overcrowding in protected areas’, ecohealth 3, 303–307. http://dx.doi.org/10.1007/s10393-006-0067-z longley, m., jepson, p.c., izquierdo, j. & sotherton, n., 1997, ‘temporal and spatial changes in aphid and parasitoid populations following applications of deltamethrin in winter wheat’, entomologia experimentalis et applicata 83, 41–52. http://dx.doi.org/10.1046/j.1570-7458.1997.00155.x matthews, k.r., 2005, ‘the developmental cell biology of trypanosoma brucei’, journal of cell science 118, 283–290. http://dx.doi.org/10.1242/jcs.01649 mecenero, s., ball, j.b., edge, d.a., hamer, m.l., henning, g.a., kruger, m. et al., (eds.), 2013, conservation assessment of butterflies of south africa, lesotho and swaziland: red list and atlas, saftronics (pty) ltd., johannesburg & animal demography unit, cape town. melton, d.a., 1987, ‘waterbuck (kobus ellipsipyrmnus) population dynamics: the testing of a hypothesis’, african journal of ecology 25, 133–145. http://dx.doi.org/10.1111/j.1365-2028.1987.tb01100.x metcalfe, d.b., asner, g.p., martin, r.e., silva espejo, j.e., huaraca huasco, w., farfán amézquita, f.f. et al., 2013, ‘herbivory makes major contributions to ecosystem carbon and nutrient cycling in tropical forests’, ecology letters 17(3), 324–332. http://dx.doi.org/10.1111/ele.12233 molyneux, d.h., 1982, ‘trypanosomes, trypanosomiasis, and tsetse control: impact on wildlife and its conservation’, in a. marcia edwards & u. mcdonnell (eds.), animal disease in relation to animal conservation: symposia of the zoological society of london, vol. 50, pp. 29–55, academic press, london. murray, g.g.r., woolhouse, m.e.j., tapio, m., mbole-kariuki, m.n., thumbi, s.m., jennings, a.e. et al., 2013, ‘genetic susceptibility to infectious disease in east african shorthorn zebu: a genome-wide analysis of the effect of heterozygosity and exotic introgression’, bmc evolutionary biology 13, 246. http://dx.doi.org/10.1186/1471-2148-13-246 nagel, p., 1995, environmental monitoring handbook for tsetse control operations, the scientific environmental monitoring group, margraf verlag, weikersheim. neitz, w.o., 1931, blood parasites of game in zululand, preliminary report, 17th report of the director of veterinary services and animal industry, union of south africa, pretoria. nwoha, r.i.o., 2013, ‘a review on trypanosomosis in dogs and cats’, african journal of biotechnology 12(46), 6432–6442. http://dx.doi.org/10.5897/ajb2013.12093 ollerton, j., winfree, r. & tarrant, s., 2011, ‘how many flowering plants are pollinated by animals?’, oikos 120, 321–326. http://dx.doi.org/10.1111/j.1600-0706.2010.18644.x pekin, b.k., 2013, ‘effect of widespread agricultural chemical use on butterfly diversity across turkish provinces’, conservation biology 27(6), 1439–1448. http://dx.doi.org/10.1111/cobi.12103 perkins, j.s. & ramberg, l. (eds.), 2004, ‘environmental recovery monitoring of tsetse fly spraying impacts in the okavango delta – 2003 final report may 2004’, the harry oppenheimer okavango research centre, university of botswana, maun. potts, s.g., biesmeijer, j.c., kremen, c., neumann, p., schweiger, o. & kunin, w.e., 2010, ‘global pollinator declines: trends, impacts and drivers’, trends in ecology and evolution 25(6), 345–353. http://dx.doi.org/10.1016/j.tree.2010.01.007 ramberg, l., hancock, p., lindholm, m., meyer, t., ringrose, s., sliva, j. et al., 2006, ‘species diversity of the okavango delta, botswana’, aquatic sciences 68, 310–337. http://dx.doi.org/10.1007/s00027-006-0857-y republic of south africa government, 1998, national environmental management act 107, republic of south africa government, pretoria, viewed n.d., from https://www.environment.gov.za/sites/default/files/legislations/nema_amendment_act107.pdf rurangirwa, f.r., musoke, a.j., nantulya, v.m., nkonge, c., njuguna, l., mushi, e.z. et al., 1986, ‘immune effector mechanisms involved in the control of parasitaemia in trypanosoma brucei-infected wildebeest (connochaetes taurinus)’, immunology 58, 231–237. samways, m.j., 1988, ‘a pictorial model of the impact of natural enemies on the population growth rate of the scale insect aonidiella aurantii’, south african journal of science 84, 270–272. samways, m.j., 2006, ‘national red list of south african odonata’, odonatologica 35, 341–368. scholes, r.j. & walker, b.h., 1993, an african savanna: synthesis of the nylsvley study, cambridge university press, cambridge. scott-dupree, c.d., conroy, l. & harris, c.r., 2009, ‘impact of currently used or potentially useful insecticides for canola agroecosystems on bombus impatiens (hymenoptera: apidae), megachile rotundata (hymentoptera: megachilidae), and osmia lignaria (hymenoptera: megachilidae)’, journal of economic entomology 102(1), 177–182. http://dx.doi.org/10.1603/029.102.0125 smith, b. & nhancale, b., 2010, ‘appendix 1: a conservation assessment for the lubombo transfrontier conservation area’, in b. smith (ed.), final report: tfca data collation and data gap analysis, pp. 1–23, report to tfca – cu conservation planning systems (cps) for the chimanimani, great limpopo and lubombo tfcas, durrell institute of conservation and ecology, university of kent. smith, r.j., easton, j., nhancale, b.a., armstrong, a.j., culverwell, j., dlamini, s.d. et al., 2008, ‘designing a transfrontier conservation landscape for the maputaland centre of endemism using biodiversity, economic and threat data’, biological conservation 141, 2127–2138. http://dx.doi.org/10.1016/j.biocon.2008.06.010 snyder, a.k. & rio, r.v.m., 2013, ‘interwoven biology of the tsetse holobiont’, journal of bacteriology 195(19), 4322–4330. http://dx.doi.org/10.1128/jb.00487-13 southern african development community, 1999, protocol on wildlife conservation and law enforcement, southern african development community, maputo, viewed n.d., from http://www.sadc.int/documents-publications/show/813 staude, h., 2016, an analysis of the moth species composition of the northern kwazulu-natal reserves: ndumo game reserve, tembe elephant park, manguzi forest reserve and kosi bay reserve, indicating a high degree of local endemism, unpublished report to ezemvelo kzn wildlife, pietermaritzburg. staude, h.s., 1999, ‘an illustrated report of 510 geometrid moth taxa (lepidoptera: geometridae) recorded from 28 protected areas from the northern and eastern parts of south africa’, metamorphosis 10, 97–148. steenkamp, y., van wyk, b., victor, j., hoare, d., smith, g., dold, t. et al., 2004, ‘maputaland–pondoland–albany’, in r.a. mittermeier, p.r. gil, m. hoffmann, j. pilgrim, t. brooks, c.g. mittermeier et al. (eds.), hotspots revisited: earth’s biologically richest and most endangered ecoregions, pp. 219–228, cemex, mexico city. united nations educational, scientific and cultural organisation, 2015, operational guidelines for the implementation of the world heritage convention, intergovernmental committee for the protection of the world cultural and natural heritage, united nations educational, scientific and cultural organisation world heritage centre, paris, viewed n.d., from http://whc.unesco.org/en/guidelines/ vamosi, j.c., knight, t.m., steets, j.a., mazer, s.j., burd, m. & ashman, t.-l., 2006, ‘pollination decays in biodiversity hotspots’, pnas 103(4), 956–961. http://dx.doi.org/10.1073/pnas.0507165103 van den bossche, p., 2001, ‘some general aspects of the distribution and epidemiology of bovine trypanosomosis in southern africa’, international journal for parasitology 31, 592–598. http://dx.doi.org/10.1016/s0020-7519(01)00146-1 waage, j.k., 1981, ‘how the zebra got its stripes – biting flies as selective agents in the evolution of zebra coloration’, journal of the entomological society of southern africa 44, 351–358. walker, b.h., emslie, r.h., owen-smith, r.n. & scholes, r.j., 1987, ‘to cull or not to cull: lessons from a southern african drought’, journal of applied ecology 24, 381–401. http://dx.doi.org/10.2307/2403882 article information authors: margaretha (gretel) w. van rooyen1 ronel henstock1 noel van rooyen1 helga van der merwe1 affiliations: 1department of plant science, university of pretoria, south africa correspondence to: gretel van rooyen email: gretel.vanrooyen@up.ac.za postal address: department of plant science, university of pretoria, pretoria 0002, south africa keywords abandoned croplands; annual species; disturbance; fallow lands; succession; vegetation recovery dates: received: 18 may 2010 accepted: 04 aug. 2010 published: 22 nov. 2010 how to cite this article: van rooyen, m.w., henstock, r., van rooyen, n. & van der merwe, h., 2010, ‘plant diversity and flowering displays on old fields in the arid namaqua national park, south africa’, koedoe 52(1), art. #1004, 7 pages. doi: 10.4102/koedoe.v52i1.1004 copyright notice: © 2010. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0771 (online) plant diversity and flowering displays on old fields in the arid namaqua national park, south africa in this original research... open access • abstract • introduction • methods    • study area    • field surveys and statistical analysis • results    • classification of plant communities       • 1. ursinia cakilefolia old field vegetation       • 2. felicia australis old field vegetation       • 3. salsola kali old field vegetation       • 4. oncosiphon grandiflorum old field vegetation    • diversity • discussion • conclusion    • research significance • acknowledgements • references abstract (back to top) the main aim of this study was to classify the vegetation on the abandoned fields in the namaqua national park and to investigate whether the time since abandonment has impacted on the diversity and mass flowering displays. sixty-two old field sample sites were surveyed using the braun-blanquet method. floristic data were analysed to produce a hierarchical classification, while a principal coordinates analysis was used to establish whether the ordination supported the classification. species richness (total number of taxa), as well as the shannon-wiener diversity index, was calculated per vegetation unit for the different life forms. four old field communities were identified, each containing a unique complement of species, which differed in their potential to produce a mass flowering display. on the skilpad section of the park, ursinia cakilefolia dominated the spectacular mass floral displays for which the park has become well known. old fields in other parts of the park did not produce the same spectacular displays. although total and perennial species richness did not differ significantly with time since abandonment, the richness of annual species, in particular of the showy annual species, decreased with time since abandonment. the relative contribution of the perennial species to vegetation cover increased with time since abandonment, whereas the relative contribution made by showy annual species to vegetation cover declined with time since abandonment. although species composition and the potential to produce mass flowering displays of the four communities differed, diversity parameters were similar. however, to maintain a flower display for tourists on the skilpad section, a degree of disturbance is essential. conservation implication: although deliberate disturbance is not desirable in a national park, we find that some disturbance is essential to maintain a spectacular flowering display. however, these intentionally disturbed old fields cover only a small proportion of the total area of the park. all other old fields in the park should not be disturbed, and should be allowed to recover naturally. introduction (back to top) the namaqua national park is situated within the succulent karoo biome – a biome recognised by the international union for conservation of nature (iucn) as one of the global hotspots of diversity (critical ecosystem partnership fund 2003; mittermeier et al. 2000; myers et al. 2000), and one of only two hotspots that are entirely arid (conservation international, undated). the biome is unique because almost half of its plant species are endemic, while the high diversity of leaf succulents and bulbous species is unrivalled amongst similar deserts (cowling et al. 1999; mucina et al. 2006). the park forms an integral part of the conservation network in namaqualand to conserve the rich biodiversity of the region. namaqualand has become synonymous with mass displays of wild flowers (van rooyen 1999). these mass displays of annuals are often a sign of land degradation and are especially prominent on croplands and other disturbed, trampled or overgrazed sites (van rooyen 2002). flowering displays on cropland are often at their best in the fields lying fallow between cultivation (van rooyen et al. 1996). whereas natural rangeland in namaqualand contains a diverse array of showy annual species, the mass effect on disturbed sites is created by the dominance of a few pioneer species and, accordingly, mass displays are believed to be associated with a loss of plant diversity (van rooyen 2002). the namaqua national park developed as an extension to the skilpad wildflower reserve. the 930-ha farm, skilpad, was originally selected by wwf-sa (at that stage the south african nature foundation) as a wildflower reserve because the farm was renowned for its springtime display on the fallow lands (van rooyen et al. 1996). this floral spectacle draws thousands of tourists annually and is a valuable source of income to the region, and especially to the park (james et al. 2007). the expansion of the park, from its small 930 ha beginning to its current size of 140 035 ha (south african national parks 2010), has led to the inclusion of many more vegetation types present in the region, and it now includes old fields situated in many of these different vegetation types. the objectives of the current study were, (1) to classify the old field vegetation, (2) to compare the communities in terms of their potential for producing mass flowering displays to draw tourists, (3) to compare diversity and life form composition across different communities and (4) to investigate the effect of time since abandonment on the diversity and life form composition of the old fields in the skilpad section of the park. methods (back to top) study area geologically, the namaqua national park is located on granites and gneisses of the bushmanland terrane of the namaqua-natal metamorphic province (cornell et al. 2006). the soils in the region are generally shallow and show minimal development, yet the diversity and spatial variability of the soils’s physical and chemical properties are exceptionally high, and could be fundamental in governing biodiversity and ecosystem function (francis et al. 2007; petersen 2008). although a diverse array of land types occurs in the park, croplands are found primarily on the ag, fc and ib land types (agis 2007). land type ag is characterised by red-yellow apedal, freely drained soils with a high base status, and is less than 300 mm deep; land type ib comprises miscellaneous land classes and rocky areas with miscellaneous soils; and land type fb comprises glenrosa and/or mispah soil forms, with lime rare or absent in upland soils, but generally present in the low-lying soils (du plessis 1987). croplands in the park are found primarily towards the east of the park on the high-lying escarpment areas (figure 1). in the mountainous namaqualand landscape suitable sites for crop cultivation are limited and farmers have selected sites with a level terrain or gentle slopes, little rock cover and deep soils. this particular habitat has therefore been largely transformed by agriculture throughout the region. the climate of the region is characterised by relatively reliable, albeit low (50 mm – 250 mm per annum), winter rainfall (> 60% winter precipitation) arriving between may and september (desmet 2007). the mean annual rainfall measured over 15 years in the skilpad section is 340 mm, whereas the mean annual rainfall at soebatsfontein located on the coastal plain below the escarpment is 140 mm per annum (south african national parks 2010). croplands are thus predominantly found on the escarpment in the eastern sections of the park (figure 1), where the rainfall is higher. the presence of the cold atlantic ocean in the west moderates temperatures throughout namaqualand and provides alternative sources of moisture in the form of coastal fog and heavy dew, experienced in the winter months (desmet 2007). figure 1: map indicating the location of the namaqua national park (nnp, in green) and the old homesteads in the vicinity of which various old fields were sampled field surveys and statistical analysis the braun-blanquet method of vegetation sampling (werger 1974) was used to conduct a floristic analysis of the old fields of the namaqua national park. forty-three old fields were sampled in 2006 and an additional 19 were surveyed in 2007. plot sizes of 20 m x 10 m were used to quantify the vegetation on the old fields. global positioning system (gps) coordinates were taken at each site and various environmental data such as aspect, slope, drainage, erosion, soil texture and rock cover were recorded.a cover-abundance value, according to the braun-blanquet scale (van der maarel 2007; werger 1974), was noted for each species present in a plot. an analysis of the floristic data was conducted using the turboveg and megatab computer packages (hennekens & schamineé 2001): turboveg software was used to capture the vegetation data and a twinspan (two way species indicator analysis; hill 1979) was run in megatab as a first step in the classification of the vegetation data. the resulting table of sample plots against species was further refined using braun-blanquet tabulation procedures (werger 1974) to produce a hierarchical classification. to visualise the relationship between the communities the floristic data were ordinated using principal coordinates analysis (pcoa) (mccune & grace 2002) in the syn-tax computer program (podani 2001). cover-abundance values were converted to percentages and the percentage values standardised using a natural logarithmic (loge) standardisation. the bray-curtis distance measure was applied for the ordination. species richness (total number of taxa), as well as the shannon-wiener index of diversity per community or subcommunity, was calculated as the mean for all sample sites for that vegetation unit. both species richness values and the shannon-wiener diversity index were also calculated separately for the perennial, annual and geophyte life forms. the annual species richness was further subdivided into those species that have the potential for producing a flower display (referred to as showy species) and those species that do not contribute to a flower display (non-showy species). diversity data were analysed using a one-way analysis of variance (anova) in the statistica computer program (statistica version 8, statsoft, tulsa). the tukey post hoc test was used to determine significant differences (p ≤ 0.05) between communities or subcommunities. diversity data were first compared across all vegetation units and thereafter the subcommunities within community 1 were compared to one another. nomenclature follows that of ‘plants of southern africa (version 2.5: ’an online checklist’ http://posa.sanbi.org). non-flowering geophytes that could not be identified were excluded from the analysis. these geophytes were infrequently encountered and did not contribute to the delineation of communities. figure 2a, 2b & 2c: principal coordinates analysis on (a) all 62 sites, (b) 31 sites of community 1 and (c) 31 sites of communities 2, 3 and 4, in the namaqua national park results (back to top) classification of plant communities four communities were identified and two of the communities were further divided into subcommunities. the communities and subcommunities are as follows (table 1 [see online appendix] and figure 1):1. ursinia cakilefolia old field vegetation 1.1 leysera gnaphalodes-ursinia cakilefolia old field vegetation 1.2 heliophila variabilis-ursinia cakilefolia old field vegetation 1.3 felicia bergeriana-ursinia cakilefolia old field vegetation 1.4 stipagrostis zeyheri-ursinia cakilefolia old field vegetation 2. felicia australis old field vegetation 2.1 grielum humifusum-felicia australis old field vegetation 2.2 galenia africana-felicia australis old field vegetation 3. salsola kali old field vegetation 4. oncosiphon grandiflorum old field vegetation overall, the classification of the floristic data provided in table 1 was supported by the ordination (figure 2a, 2b and 2c). the pcoa of the floristic data of all 62 relevés clearly indicated the separation between community 1 and communities 2 to 4 (figure 2a). two separate ordinations were then run, the first on the floristic data of community 1 and a second on the floristic data of communities 2, 3 and 4. in the ordination of community 1 the subcommunities were arranged along the first axis from right (subcommunity 1.1) to left (subcommunity 1.4) and this gradient was related to the time since abandonment (figure 2b). the ordination of communities 2, 3 and 4 produced three separate clusters (figure 2c). the current study focused on a vegetation ‘state’ in disturbed habitats, predominantly composed of annual plant species, and thus the vegetation descriptions were based strongly on annual species. because of the dynamic nature of this disturbed vegetation, some of the subcommunities are arranged along a continuum (figure 2b) and some of the vegetation units identified here may even be transient. 1. ursinia cakilefolia old field vegetation this plant community was located on the farm skilpad in the namaqualand klipkoppe vegetation type (skn1; mucina et al. 2006) at altitudes from 620 m to 755 m above sea level. most of this community was found on land type ag94, although land type ib235 also occurred. community 1 was generally found on gentle or moderate slopes, on well-drained sandy loams or loamy sands with no rock cover.the vegetation of this community was characterised by species group g with, generally, a high cover of ursinia cakilefolia. other common species included gazania leiopoda and felicia bergeriana (species group g), heliophila variabilis, leysera gnaphalodes, conicosia elongata and oxalis obtusa (species group ac). this community was divided into four subcommunities. 1.1 leysera gnaphalodes-ursinia cakilefolia old field vegetation: subcommunity 1.1 was found along the circular tourist route on the skilpad section of the namaqua national park and represented those fields in the park that are regularly disturbed (tilled), at approximately four-year intervals, to produce mass flowering displays. the subcommunity occurred exclusively on land type ag94 and the gentle slopes and plains, with no rock cover, comprised well-drained loamy sand and sandy loam soils. altitudes ranged from 686 m to 749 m above sea level. senecio arenarius and helichrysum cf. hebelepis (species group a) differentiated this subcommunity. lotononis leptoloba (species group d), ursinia cakilefolia (species group g), heliophila variabilis and leysera gnaphalodes (species group ac) were present with high cover values. pentaschistis tomentella (species group j), arctotheca calendula (species group v), lapeirousia silenoides (species group y), lessertia diffusa, conicosia elongata, wahlenbergia annularis and dischisma spicatum (species group ac) were always present in subcommunity 1.1. 1.2 heliophila variabilis-ursinia cakilefolia old field vegetation: subcommunity 1.2 was found towards the eastern part of the skilpad section. this subcommunity occurred on land type ag94, on old fields that had been actively cultivated until the skilpad wildflower reserve was established, but have not been disturbed since then. in general, the old fields in subcommunity 1.2 were 12–15 years old. the loamy sands were welldrained and the fields occurred on level to gentle slopes at altitudes ranging from 620 m to 680 m above sea level. no rock cover was present, with the exception of one site with a rock cover of 5%. the species of species group b, such as gymnodiscus linearifolia and lotononis pentaphylla, characterised this subcommunity. ursinia cakilefolia, gazania leiopoda (species group g), pentaschistis tomentella (species group j), heliophila variabilis and leysera gnaphalodes (species group ac) often had high cover values. species with a high constancy were manulea altissima, othonna pavonia (species group c), arctotheca calendula (species group v), limeum africanum, lapeirousia silenoides (species group y), dimorphotheca sinuata and conicosia elongata (species group ac). 1.3 felicia bergeriana-ursinia cakilefolia old field vegetation: subcommunity 1.3 included old fields in the park (eastern part of the skilpad section) as well as in the adjacent farmland at altitudes ranging from 624 m to 680 m above sea level. field age of the sites in the park ranged from approximately 20–60 years. in contrast, the fields on the adjacent farmland were estimated to be approximately 10–20 years old at the time of the study. the gentle to moderately sloping old fields, on land type ag94, were characterised by well-drained loamy sands with a low (< 1%) or zero rock cover. with the exception of the two sites abandoned approximately 60 years ago, all fields were still clearly distinguishable from the surrounding vegetation. this subcommunity had no diagnostic species group and was transitional between subcommunities 1.2 and 1.4. abundant species included ursinia cakilefolia, gazania leiopoda, felicia bergeriana (species group g), trichogyne glomerata (species group i), pentaschistis tomentella, cotula barbata (species group j), and heliophila variabilis (species group ac). species with a high constancy were nylandtia spinosa (species group g), trachyandra falcata, lapeirousia silenoides (species group y), leysera gnaphalodes, conicosia elongata, oxalis obtusa and ehrharta calycina (species group ac). 1.4 stipagrostis zeyheri-ursinia cakilefolia old field vegetation: subcommunity 1.4 represented those fields that had been cultivated more than 50–80 years ago. most of the sites were surveyed on the adjacent farmland. land type ib235 dominated this subcommunity, which was found at altitudes ranging from 660 m to 755 m above sea level. the well-drained loamy sands occurred on level to gentle slopes and usually surface rocks were absent. the vegetation at these sites could hardly be distinguished from the natural vegetation and most sites were pointed out by the farmer. species group e, with species such as searsia horrida, ballota africana and aspalathus cf. pulicifolia, characterised this subcommunity. dorotheanthus bellidiformis (species group f), dodonaea viscosa and nylandtia spinosa (species group g), microloma sagittatum (species group i), as well as heliophila variabilis and oxalis obtusa (species group ac) were generally present. large perennial shrub species such as searsia horrida, euclea tomentosa (species group e), wiborgia mucronata (species group f), nylandtia spinosa and dodonaea viscosa (species group g) and a high cover of stipagrostis zeyheri (species group ab) distinguished this subcommunity from the others. 2. felicia australis old field vegetation this community was located in the vicinity of the farms keerkom and canariesfontein and the old homestead of bitterputs. it occurred predominantly on land type fb154 and in a few cases on land type ib235. the sandy loam soils were usually moderately to welldrained but on occasion waterlogged or poorly drained. rock cover varied between zero and < 10% on the gentle to moderate slopes of these old fields. altitudes ranged from 473 m to 705 m above sea level. the natural vegetation was located within the namaqualand klipkoppe (skn1) vegetation type (mucina et al. 2006). community 2 was characterised by the presence and high cover of felicia australis (species group q) together with galenia africana (species group ac). other conspicuous species were hermannia amoena (species group u), calobota sericea (species group v), zaluzianskya benthamiana (species group y), heliophila variabilis, helichrysum tinctum, wahlenbergia annularis and dischisma spicatum (species group ac). communities 2 and 3 showed strong floristic affinities as indicated by the presence of species groups t and u. 2.1 grielum humifusum-felicia australis old field vegetation: subcommunity 2.1 was found on the farm canariesfontein at altitudes ranging from 483 m to 705 m above sea level. the slopes were gentle to moderate and usually covered with moderately to well-drained, sandy loam soils, and no to little (2% – < 10%) rock cover. this subcommunity did not have a strong species group differentiating it from other communities. commonly occurring species shared with community 1 were trichogyne glomerata (species group i) and pentaschistis tomentella (species group j). dominant species occurring in subcommunity 2.1 included felicia australis (species group q), hermannia amoena (species group u), calobota sericea (species group v), gorteria diffusa and hermannia trifurca (species group ab), galenia africana, leysera gnaphalodes, dimorphotheca sinuata, grielum humifusum and helichrysum tinctum (species group ac). 2.2 galenia africana-felicia australis old field vegetation: subcommunity 2.2 was located on the farm keerkom and in the vicinity of the old bitterputs and rondeklip homesteads. it was found on both land type fb154 and ib235 and the sandy loam soils ranged from waterlogged to well-drained. surface rocks were generally absent, but at two sites a low rock cover (3% – 5%) was found. the altitudes ranged from 473 m to 686 m above sea level. felicia australis (species group q), drosanthemum hispidum (species group t), and galenia africana (species group ac) dominated subcommunity 2.2. hermannia amoena (species group u), senecio cardaminifolius (species group v), zaluzianskya benthamiana (species group y), heliophila variabilis and wahlenbergia annularis (species group ac) had a high constancy in this subcommunity. two variants could be distinguished within this subcommunity: the first occurring on keerkom and characterised by species group l, and the second mostly at bitterputs and rondeklip and characterised by species group o. 3. salsola kali old field vegetation community 3 occurred in the vicinity of platklip and windkraal at altitudes ranging from 477 m to 622 m above sea level. the old fields of this community were predominantly found within land type ib124 on level areas with no rock cover. the soils were usually well-drained loamy sands. the natural vegetation fell within the namaqualand klipkoppe (skn1) vegetation type (mucina et al. 2006). salsola kali and helichrysum oxybelium (species group s) differentiated this community. furthermore, the strong presence of drosanthemum hispidum (species group t), atriplex lindleyi, galenia fruticosa (species group w), zaluzianskya benthamiana, aizoon canariense (species group y), calobota halenbergensis (species group aa) and galenia africana (species group ac) also distinguished this group. community 3 was closely related to community 2 (species groups t and u) as well as community 4 (species group aa), with species groups w and x indicating the affinities between all three communities. 4. oncosiphon grandiflorum old field vegetation located on melkboom, this community was the only community that did not occur in the namaqualand klipkoppe (skn1) vegetation type of mucina et al. (2006), but was located in the namaqualand heuweltjieveld (skn4). with the exception of one plot, these old fields were found on land type ag96, on gentle to moderate slopes, with a rock cover of less than 2%. a low altitude (264 m – 335 m above sea level) and well-drained sandy loams distinguished this community. species group z, with species such as oncosiphon grandiflorum and arctotis fastuosa, differentiated this community. lotononis parviflora, karroochloa schismoides (species group aa) and heliophila variabilis, leysera gnaphalodes, galenia africana, dimorphotheca sinuata, grielum humifusum and helichrysum tinctum (species group ac) were common species that occurred across most, if not all, the relevés representing community 4. diversity a comparison between the four communities revealed no significant differences in the species richness or shannon-wiener diversity index for all the species (p > 0.05), perennial species (p > 0.05) or annual species (p > 0.05). when only community 1 was considered there were, however, significant differences in the diversity parameters between the subcommunities. annual species richness was significantly lower in subcommunity 1.4 than in the other subcommunities (table 2a). these differences were primarily due to differences in the showy component of the annual species, with no differences observed between species richness of the non-showy annual species (table 2a). the shannon-wiener diversity index confirmed the lower annual diversity in subcommunity 1.4 than in subcommunity 1.1 (see table 3 superscripts for significance). the shannon-wiener index also indicated the opposite trend for the perennial species diversity, with subcommunity 1.4 having a significantly higher perennial diversity than subcommunity 1.1 (table 3). most of the differences in species richness among the subcommunities of community 1 were reflected in the contributions made to the vegetation cover by the different life forms (table 2b). in figure 3a and figure 3b the relative contributions of the species per life form to the total number of species (figure 3a) and the relative contribution of vegetation cover per life form to the total vegetation cover (figure 3b) are illustrated. these figures clearly show the increased contribution of perennial species to species richness and cover from subcommunity 1.1 to 1.4, and a concomitant decrease in the contribution by annual species to richness and cover from subcommunity 1.1 to 1.4. the changes in relative contribution to richness and cover were evident for the showy component of the annual species, but not for the non-showy component. figure 3a & 3b: relative contribution of perennial species, annual species (subdivided into showy and non-showy annual species) and geophyte species to (a) the total number of species, and (b) the vegetation cover of community 1, in the namaqua national park table 2a & 2b: 2a: species richness per community or subcommunity for various life forms 2b: vegetation cover per community or subcommunity for various life forms table 3: the shannon-wiener diversity index for the old field communities/subcommunities in the namaqua national park discussion (back to top) a clear distinction could be made between community 1 and the other three communities (figure 2a). community 1 was located on the farm skilpad (in and adjacent to the park) at a fairly high altitude (620 m – 710 m), with subcommunities 1.1–1.3 on land type ag94 and subcommunity 1.4 on land type ib235. the subcommunities of community 1 were associated with time since abandonment, with subcommunity 1.1 being the youngest and subcommunity 1.4 the oldest (figure 2b). the sample sites of subcommunity 1.1 were fairly closely clustered (figure 2b), but as time progressed, the floristic composition of the fields became increasingly diverse indicating that there were divergent pathways along which succession on these old fields proceeded. subcommunity 1.4 represented the oldest fields, however, these sites were also on a different land type to the other subcommunities, and this would have influenced the species assemblage patterns on the old fields in this subcommunity. although all the sample sites within community 1 have a similar ploughing history, divergence among the older abandoned fields could have been the result of different farming practices on the old fields, such as grazing history and intensity (bonet 2004, cadenasso et al. 2002), number of times ploughed, the various crops planted and the last crop planted (bonet 2004; myster & pickett 1990), or the rainfall amount and temporal distribution in the year of abandonment (myster & pickett 1990; pickett et al. 2008). it is notable that old fields surveyed in the adjacent farmland were classified into the same subcommunities of community 1 as those in the park. however, most of the fields surveyed in the adjacent farmland had been abandoned for several decades and belonged to subcommunities 1.3 and 1.4. therefore, for the first years after abandonment both the fields inside and outside the park would have been subjected to livestock grazing. it is only the more recently abandoned fields in the park that were subjected to a conservation land-use with a low grazing intensity after abandonment. these old fields were classified as subcommunity 1.2. a comparison of the subcommunities of different ages in community 1 clearly indicated that secondary succession on these old fields proceeded very slowly. such a slow rate of vegetation change is common in arid environmental conditions (dean & milton 1995; otto et al. 2006). the vegetation on many old fields abandoned more than 50 years ago was still clearly visually distinguishable from the surrounding vegetation, although the vegetation on several sites in subcommunity 1.4 that had not been cultivated for more than 80 years could hardly be distinguished from the surrounding vegetation. the remaining three communities (2, 3 and 4) were clearly separated from each other and from community 1 (figure 2c). they were found in different vegetation types, on different land types and at different altitudes. community 2 was characterised predominantly by land type fb154 and occurred at intermediate altitudes, whereas community 3 occurred predominantly on land type ib124, also at intermediate altitudes. community 4 occurred in heuweltjieveld on land type ag96 and at the lowest altitudes of all sites surveyed. because rainfall is strongly influenced by the altitudinal gradient in this region, some of the differences between the communities could be ascribed to differences in available moisture. comparisons of the diversity among the four communities revealed no significant differences. nevertheless, there were large differences in their potential to produce floral displays. community 1 (subcommunities 1.1 through to 1.3 in particular) was the only community with the mass floral displays of ursinia cakilefolia for which the namaqua national park has become world-renowned. the relative contribution of the perennial, annual (showy and non-showy) and geophyte life forms within community 2 and community 4 were similar. however, subcommunity 2.1 had a higher cover of showy annual plant species than subcommunity 2.2 and community 4. the most prominent annual species in subcommunity 2.1 were felicia australis, dimorphotheca sinuata and grielum humifusum, whereas subcommunity 2.2 was dominated only by the annual felicia australis, and community 4 had high cover values for oncosiphon grandiflorum, arctotis fastuosa and grielum humifusum. overall, community 3 offered the least to tourists by way of mass flowering displays. this community had a high cover of perennial species, largely dominated by galenia africana, calobota halenbergensis and galenia fruticosa, and most of the annuals occurring in the community were inconspicuous. the rocky soils (land type ib124) in this community could have contributed to the low cover of annual species. many different relationships between species richness and age since abandonment have been reported. theoretically, an asymptotic relationship is predicted under environmentally limiting conditions (peet 1992), although bonet (2004) found a parabolic relationship in an arid region in southern spain, whereas otto et al. (2006) reported a linear increase under arid conditions. a parabolic relationship is believed to be typical of the vegetation in the mediterranean basin (bonet 2004; debussche et al. 1996,) and could therefore have been expected in namaqualand with its mediterranean climate, with winter rainfall. however, total and perennial species richness did not differ significantly among the subcommunities of community 1, which reflect a temporal gradient from less than 5 years to about 80 years since cultivation. the relative richness of annual species, and in particular of the showy annual species, showed a decrease with time since abandonment (figure 3a). within community 1 it was clear that while the relative contribution of the perennial species to vegetation cover increased with time since abandonment, the relative contribution made by showy annual species to vegetation cover declined with age since abandonment (figure 3b). these results support the general contention that spectacular mass floral displays rely on regular disturbance (van rooyen 2002), such as experienced in subcommunity 1.1. whether the mass floral displays are associated with a loss in plant diversity depends on how diversity is measured. subcommunity 1.1, with exceptional mass displays, did not have a lower total species richness or richness of perennial species than the other subcommunities of community 1, although the shannon-wiener diversity index indicated a lower diversity in the perennial component in subcommunity 1.1. in contrast, both species richness as well as the shannon-wiener index indicated a higher diversity for the annual component in subcommunity 1.1 than in subcommunity 1.4. understanding the process of natural vegetation change and diversity relationships of old fields in the namaqua national park is essential to evaluate the conservation outcomes of the park. however, the current study is not only relevant for the park. worldwide, there is an increasing trend of land abandonment as a result of environmental and socio-economic changes (cramer & hobbs 2007; cramer et al. 2007). a similar trend can be seen in namaqualand (hoffman & rohde 2007). consequently, many abandoned croplands lie scattered throughout the namaqualand landscape. the insight gained into vegetation dynamics of old fields in the namaqua national park could provide a framework (bartha et al. 2003; pickett et al. 2008) for the investigation of restoration opportunities of old fields in the larger namaqualand region. conlusion (back to top) the aim of this study was to classify the plant communities on the abandoned fields across different vegetation types in the namaqua national park and to investigate the impact of time since abandonment on plant diversity and mass flowering displays. the knowledge gained in this study provides insight that could aid future planning and effective management of the many old fields scattered over this extensive conservation area. old fields located on the skilpad section belong to a single community and produce the characteristic mass floral displays of ursinia cakilefolia for which the namaqua national park has become world-renowned. old fields in other parts of the park belong to other communities, and will not produce the same spectacular ursinia cakilefolia dominated displays as the old fields on skilpad but should contribute to the appeal of the park by increasing the diversity of the springtime displays of wild flowers. on the skilpad section the best flowering displays were associated with the old fields that were tilled approximately every 4 years (subcommunity 1.1), although a variety of showy annual species were still commonly found on fields abandoned about 20 years ago. to maintain a flower display for tourists this type of management intervention appears to be essential. apart from the old fields along the circular tourist route on the skilpad section, all other old fields in the park should not be disturbed, but should be allowed to recover naturally. research significance understanding the process of natural vegetation change and diversity relationships of old fields in the namaqua national park is essential to evaluate the conservation outcomes of the park. the insight gained into vegetation dynamics of old fields in the park could, furthermore, provide a framework for the investigation of restoration opportunities of old fields in the larger namaqualand region. acknowledgements (back to top) the authors gratefully acknowledge the support received from the german federal ministry of education and research (bmbf) through the biota south project, and the national research foundation under grant number 61277. sanparks and in particular, dr h. bezuidenhout and dr h. hendricks are thanked for permission to conduct research on the namaqua national park. we also thank various students from the department of plant science, university of pretoria, and annelise le roux (succulent karoo knowledge centre) for field assistance. editor: llewellyn c. foxcroft references (back to top) agis, 2007, agricultural geo-referenced information system, viewed 10 april 2010, from www.agis.agric.za bartha, s., meiners, s.t., pickett, s.t.a. & cadenasso, m.l., 2003, ‘plant colonization window in a mesic old field succession’, applied vegetation science 6, 205−212. bonet, a., 2004, ‘secondary succession of semi-arid mediterranean old-fields in south-eastern spain: insights for conservation and restoration of degraded lands’, journal of arid environments 56, 213−233. cadenasso, m., pickett, s.t.a. & morin, p.j., 2002, ‘experimental test of the role of mammalian herbivores on old field succession: community structure and seedling survival’, journal of the torrey botanical society 129, 228−237. conservation international, n.d., biodiversity hotspots, viewed 10 april 2010, from http://www.biodiversityhotspots.org cornell, d.h., thomas, r.j., moen, h.f.g., reid, d.l., moore j.m. & gibson, r.l., 2006, ‘the namaqua-natal province’, in m.r. johnson, c.r. anhaeusser & r.j. thomas (eds.), the geology of south africa, pp. 325−379, council for geoscience, pretoria. cowling, r.m., esler, k.j. & rundel, p.w., 1999, ‘namaqualand, south africa – an overview of a unique winter-rainfall desert ecosystem’, plant ecology 142, 3–21. cramer, v.a. & hobbs, r.j. (eds.), 2007, old fields. dynamics and restoration of abandoned farmland, island press, washington. cramer, v.a., hobbs, r.j. & standish, r.j., 2007, ‘what’s new about old fields? land abandonment and ecosystem assembly’, trends in ecology and evolution 23, 104−112. critical ecosystem partnership fund, 2003, ecosystem profile: the succulent karoo hotspot, namibia and south africa, critical ecosystem partnership fund report. dean, w.r. & milton, s.j., 1995, ‘plant and invertebrate assemblages on old fields in the arid southern karoo, south africa’, african journal of ecology 33, 1−13. debussche, m., josé, e., lepart, j., houssard, c., & lavorel, s., 1996, ‘changes in mediterranean plant succession: old-fields revisited’, journal of vegetation science 7, 519−526. desmet, p.g., 2007, ‘namaqualand – a brief overview of the physical and floristic environment’, journal of arid environments 70, 570−587. du plessis, h.m., 1987, ‘land types of the maps 2816 alexander bay, 2818 warmbad, 2916 springbok, 2918 pofadder, 3017 garies, 3018 loeriesfontein’, memoirs on the agricultural natural resources of south africa 9, 1–538. francis, m.l., fey, m.v., prinsloo, h.p., ellis, f., mills, a.j. & medinski, t.v., 2007, soils of namaqualand: compensations for aridity, journal of arid environments 70, 588−603. hennekens, s.m. & schaminée, j.h.j., 2001, ‘turboveg, a comprehensive data base management system for vegetation data’, journal of vegetation science 12, 589−591. hill, m.o., 1979, twinspan – a fortran program for arranging multivariate data in an ordered two-way table by classification of the individuals and attributes, ecology & systematics, cornell university ithaca, new york. hoffman, m.t. & rohde, r.f., 2007, ‘from pastoralism to tourism. the historical impact of changing land use practices in namaqualand’, journal of arid environments 70, 641−658. james, i., munro, a., hoffman, t., o’farrell, p.j. & smart, r. 2007. the economic value of flower tourism at the namaqua national park, south africa, south african journal of economic and management sciences 10, 442−456. mccune, b. & grace, j.b., 2002, analysis of ecological communities, mjm software design, gleneden beach, oregon. mittermeier r.a., myers, n., robles gil, p. & mittermeier, c.g., 2000, hotspots: earth’s biologically richest and most endangered terrestrial ecoregions, cemex, mexico. mucina, l., jürgens, n., le roux, a., rutherford, m.c., schmiedel, u., esler, k.j. et al., 2006, ‘succulent karoo biome’, in l. mucina & m.c. rutherford (eds.), the vegetation of south africa, lesotho and swaziland, strelitzia 19, pp. 220−299, south african national biodiversity institute, pretoria. myers, n., mittermeier, r.a., mittermeier, c.g., de fonseca, g.a.b. & kent, j., 2000, ‘biodiversity hotspots for conservation priorities’, nature 403, 853−858. myster, r.w. & pickett, s.t.a., 1990, ‘initial conditions, history and successional pathways in ten contrasting old fields’, american midland naturalist 124, 231−238. otto, r., krüsi, b.o., burga, c.a., fernández-palacios, j.m., 2006, ‘old-field succession along a precipitation gradient in the semi-arid coastal region of tenerife’, journal of arid environments 65, 156−178. peet, r.k., 1992, ‘community structure and persistence’, in d.c. glenn-lewin, r.k. peet & t.t. veblen (eds.) plant succession: theory and prediction, pp. 103−151, chapman & hall, new york. petersen, a., 2008, ‘pedodiversity of southern african drylands’, phd thesis, institute of soil science, university of hamburg. pickett, s.t.a., cadenasso, m.l. & meiners, s.j., 2008, ‘ever since clements: from succession to vegetation dynamics and understanding to intervention’, applied vegetation science 12, 9−21. podani, j., 2001, syn-tax 2000 computer programs for data analysis in ecology and systematic, scientia publishing, budapest. south african national parks, 2010, ‘namaqua national park: park management plan’, february 2010 (draft), sanparks, pretoria. van der maarel, e., 2007, ‘transformation of cover-abundance values for appropriate numerical treatment – alternatives to the proposals by podani’, journal of vegetation science 18, 767−770. van rooyen, m.w., theron g.k. & van rooyen, n., 1996, ‘skilpad wildflower reserve: a flower-lover’s paradise’, veld and flora 4, 40−42. van rooyen, m.w., 1999. functional aspects of short-lived plants, in w.r. dean & s.j. milton (eds.), the karoo: ecological patterns and processes, pp. 107−122, cambridge university press, cambridge. van rooyen, m.w., 2002, ‘management of the old field vegetation in the namaqua national park, south africa: conflicting demands of conservation and tourism’, geographical journal 168, 211−223. werger, m.j.a., 1974, ‘on concepts and techniques applied in the zürich-montpellier method of vegetation survey’, bothalia 11, 309−323. book review biography of marjorie courtenay-latimer – book review book title: curator and crusader: the life and work of marjorie courtenay-latimer author: mike bruton isbn: isbn 9780639932699 paperback publisher: footprint press, southern africa, 2019, r280.00* *book price at time of review review title: biography of marjorie courtenay-latimer – book review reviewer: peter a. novellie1 affiliation: 1sustainability research unit, nelson mandela university, george, south africa corresponding author: peter novellie, novellie@netactive.co.za how to cite this book review: novellie, p.a., 2020, ‘biography of marjorie courtenay-latimer – book review’, koedoe 62(1), a1607. https://doi.org/10.4102/koedoe.v62i1.1607 copyright notice: © 2020. the authors. licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. this biography tells the enchanting story of how a determined little girl, described by her father as a ‘quaint little serious minded child’, grew up to become freeman dame dr marjorie eileen dorothy courtenay-latimer dsc, fsama, mcc, dmss – this despite her never having received a tertiary education, probably not even a matriculation certificate. based on thorough background research from marjorie’s birth in 1907 to the end of her life in 2004, mike bruton builds a complete picture of how this incredible fairy tale unfolded. it is the story of how a precocious childhood passion for collecting items of natural or cultural historical interest laid the foundation for a career that made a lasting contribution to museology in south africa and attracted an impressive variety of accolades. while still a teenager, marge’s extremely diverse field knowledge of birds, plants and cultural artefacts drew the attention, and mentorship, of a few prominent local biologists and educators and led to her appointment, at the age of only 24 years, as curator of the fledgling east london museum, ahead of many older and better qualified applicants for the position. marge is best known for her role in the discovery, and presentation to the world of science, of a specimen of a living coelacanth. the extraordinary impact of this discovery is intriguingly described. marge’s persistence in ensuring that the coelacanth came to the attention of ichthyologist j.l.b. smith, despite a series of obstacles, showed her clear appreciation of the significance of the discovery, even though she had little knowledge of fishes at the time. her tenacity graphically illustrates louis pasteur’s famous quote: in the fields of observation chance favours only the prepared mind. bruton presents evidence that marge developed a mind prepared for discovery very early in life. on 24 february 1918, marge’s 11th birthday, her father wrote, ‘she is naturally gifted with a flare of finding things and knowing by instinct what is wanted or knowing where to look for what she wants’ (introductory quotation to chapter 3). the discovery of the coelacanth in late 1938 brought world renown to both marge and her museum. however, it would do her a great injustice to see her only in this context. as bruton shows, marge had already made impressive progress with the development of the east london museum before the discovery of the coelacanth, and was to make a wide diversity of important further contributions up to and beyond her retirement at the age of 66. apart from her contributions to science, marge was a natural story teller and inspirational as an educator and conservationist, which enabled her to make her museum a ‘thriving hub of community activity’. readers with an interest in sanparks and its history will appreciate the story of the tsitsikamma conference (pp. 208, 212) convened by the national parks board (as sanparks was formerly called) at the tsitsikamma forest inn in 1964. the aim of the conference was to encourage research in the tsitsikamma coastal national park (tcnp), an early example of research-friendly policies that have long been followed by sanparks. researchers were offered free accommodation, collecting permits, travel allowance and even a daily allowance of r2.00! in this aim, the conference was highly successful; marge and several others made significant contributions. an important sequel was marge’s first scientific book on the flowering plants of the tcnp and tsitsikamma forest (1967), which included extensive comments on traditional medicinal uses of plants, enriched by her vast experience in the field. another sequel was marge’s 115 page manuscript on the birds of the park and forest, written in the style of a field guide. unfortunately – for unclear reasons (bruton suggests that it may have been because of over-sensitivity on marge’s part to criticisms of the manuscript) – it remains unpublished in the archives of the east london museum. bruton raises the question of whether this work may yet be published. one feels it really should be. given her long life filled with achievements and awards, a biographer could easily present a haloed image of ‘marvellous marge’. however, making good use of an abundance of letters and reports written by marge herself, and by people who knew her well, bruton portrays a complex personality: a strong-willed, energetic and engaging people’s person, enjoying the limelight but also shy and sometimes self-doubting, prone to minor deceptions that made her or her museum look good, and overly sensitive to criticism. bruton presents marge’s contributions in relation to a broader perspective of museums and museology in south africa, drawing attention to the long history of museums – going back to the engagement of andrew smith in 1825 to establish a museum in cape town – and their value as sources of information collected over years by numerous dedicated scholars. the postscript to the biography describes the strong current state of the east london museum, as well as the diverse wealth of museums in south africa generally. this will be both reassuring and enlightening to those (including myself) who believed our natural history museums to be neglected and declining. the story of marge’s life and work is well supported by an abundance of background material, which in no way slows down the engaging narrative. i found it gripping and difficult to put down. this book is a must-read for anyone with an interest in museology. however, because of marge’s eclectic interests and her enchanting, fairy tale career, it will be enjoyed by a wide diversity of readers. van staden.qxd major plant communities of the marakele national park p.j. van staden and g.j. bredenkamp van staden, p.j. & g.j. bredenkamp. 2005. major plant communities of the marakele national park. koedoe 48(2): 59-70. pretoria. issn 0075-6458. to manage and conserve any national park efficiently, a profound knowledge of the ecology is a prerequisite, and to achieve that an inventory of the biotic and abiotic components must be undertaken. as a contribution to such a program this information was collected for marakele national park. the study area covers 290.51 km² in the southwestern part of the limpopo province. the underlying parent rock of the study area is sandstone, shale and mudstone with several diabase dykes. the soils range from shallow to deep sandy soils on sandstone and clayey soils on diabase and mudstone. the rainfall varies from 556 mm to 630 mm per annum, mainly during the summer months. the study area experiences warm summers with temperatures of up to 32 ºc and cool, dry winters with frost in the low-lying areas. the vegetation of the study area was classified in a hierarchical, plant sociological system by using twinspan and the braunblanquet technique. the floristic data from 130 relevés were classified to identify five major plant communities, namely one forest community, three savanna/grassland communities and one wetland community. these plant communities were ecologically interpreted by habitat.the phytosociological table was condensed to a synoptic table to describe the major plant communities. key words: braun-blanquet, classification, major plant communities, phytosociology, synoptic table. pj van staden, centre for wildlife management, university of pretoria, 0001 pretoria; gj bredenkamp, dept of botany, university of pretoria, 0001 pretoria (george.bredenkamp@.up.ac.za) issn 0075-6458 59 koedoe 48/2 (2005) introduction the primary objective when proclaiming a national park is to conserve parts of pristine natural ecosystems for future generations (gertenbach 1987). marakele national park (mnp) covers an area of 290.51 km² in the southwestern part of the limpopo province and is managed as a national park since 1988, but was officially proclaimed a national park on 11 february 1994. in order to manage and conserve any conservation area, a profound knowledge of the ecology is a prerequisite, and to achieve that prerequisite, an inventory of the biotic and the abiotic components of that national park must be undertaken (edwards 1972). the “natural systems” as they occur today cannot be viewed and conserved as “natural” any more, because of the influence of man. thus, management recommendations can only be made on the basis of interpreted ecological knowledge to restore the balance of the original natural system. the influence of management recommendations on the system must also be regularly monitored to determine if the aims that were set have been achieved satisfactorily. efficient monitoring systems also depend upon an inventory of the biotic and abiotic components (gertenbach 1987). the primary aim of this study was to classify and describe the vegetation of the marakele national park in the waterberg of the limpopo province (van staden 2003). the purpose of the classification can be described as to form an inventory as a basis of management. study area the study area covers 29051 ha in the southwestern part of the limpopo province of south africa, between 27°30'e–27°45'e and 24°15'–24°30's. the position of the park in relation to towns is shown in fig. 1 (van staden 2003). the underlying parent rock of the study area (sacs 1980) is sandstone of the matlabas subgroup, aasvoëlkop formation in the south-western and southern parts; with shale and mudstone of the matlabas subgroup, aasvoëlkop formation, groothoek mudstone member; a conglomerate outcrop of the matlabas subgroup, aasvoëlkop formation in the west; and with the biggest part of the study area consisting of sandstone of the kransberg subgroup, sandriviersberg formation. the soils that have developed on the parent materials range from shallow to deep sandy soils on sandstone and clayey soils on diabase and mudstone (van staden 2003). the rainfall varies from 556–630 mm per annum and occurs mainly during the summer months. the study area experiences warm, wet summers with average daily temperatures of 32 ºc and cool, dry winters with frost in the low-lying areas (van staden 2003). mnp is situated mainly in the waterberg moist mountain bushveld (low & rebelo 1996) in the savanna biome (rutherford & westfall 1994). the vegetation of the study area includes acocks’ (1988) sour bushveld (veld type 20), mixed bushveld (veld type 18), sourish mixed bushveld (veld type 19) and north-eastern mountain sourveld (veld type 8). this sour bushveld is listed by edwards (1972) as one of 52 of south koedoe 48/2 (2005) 60 issn 0075-6458 fig. 1. a map indicating the location of the study area in relation to towns. african veld types that is extremely lacking in conservation. the sour bushveld covers 18306 km2, occurring in mountainous areas in the previous transvaal province (coetzee 1975; coetzee et al. 1981). previous plant ecological studies in the sour bushveld (acocks 1988) include those by van vuuren & van der schijff (1970), coetzee (1975); coetzee et al. (1981), westfall (1981) and westfall et al. 1985. other plant ecological work of interest in related vegetation types include those of theron (1973) who described the vegetation of the loskopdam nature reserve; van der meulen (1979) who described the vegetation of the bushveld south of the waterberg; van rooyen (1983) who described the vegetation of roodeplaat dam nature reserve; and brown (1997) who described the vegetation of borakalalo game reserve. methods analysis methods applied are described in detail by van staden (2003) and are only summarised here. stereo aerial photographs on a scale of 1:50 000 (task 874 of 1984) were used to delineate homogeneous units on the basis of physiography and physiognomy (bredenkamp & theron 1978; westfall 1981; gertenbach 1987). a total of 130 sample plots were surveyed throughout the study area. the sample plot location was determined by means of stratified-random sampling (westfall 1981). the number of sample plots for each delineated physiographic-physiognomic unit was determined according to the size of each delineated unit. termitaria and riparian vegetation was not included in the placing of the sample plots, due to their limited size. additional sample plots were identified for the termitaria and riparian vegetation and sampling was done in these vegetation types. a sample plot size of 10 m x 20 m was fixed and used throughout the study area. this size is considered adequate for surveys in savanna vegetation by coetzee (1975), coetzee et al. (1976), westfall (1981), van rooyen (1983) and gertenbach (1987). at each sample plot a list of all the species present, was compiled. a cover-abundance value was given to each species according to the braun-blanquet cover-abundance scale, as given by mueller-dombois & ellenberg (1974) and werger (1974), and adapted by barkman et al. (1964). the following habitat information was recorded at each sample plot—symbols are used in the synoptic table: land type was read from the land type map (land type survey staff 1988); soil forms were classified in accordance with macvicar et al. (1977); the altitude of each sample plot was recorded using an altimeter and is given in metres; the slope of the terrain of each sample plot was measured in degrees, using an optical clinometer. the following classification of slope units (westfall 1981) were used in this study: symbol description class l level 0.00° -3.49° g gentle 3.50° 17.62° m moderate 17.63°36.39° s steep 36.40° the aspect of the terrain where each sample plot is situated was determined using a compass. aspect is given in the eight compass directions, namely: n north; s south; ne northeast; sw southwest; e east; w west; se southeast; nw northwest. the surface rock cover in each sample plot was estimated as a percentage stones (> 20 mm diameter), boulders and rocky outcrops. the following five classes were used, based on its potential influence on mechanical use (ploughing) (van der meulen 1979; westfall 1981): symbol class description o < 1 % no limitation on mechanical utilisation l 1–4 % low limitation on mechanical utilisation m 5–34 % moderate limitation on mechanical utilisation h 35–84 % high limitation on mechanical utilisation v 85–100 % no mechanical utilisation possible synthesis the classification is based on the braun-blanquet method of vegetation classification, discussed in detail by westhoff & van der maarel (1980), mueller-dombois & ellenberg (1974) and werger (1974). data were captured on the mainframe computer of the university of pretoria, in the bbnew software package. it was then exported to be used in the software package bbpc (bezuidenhout et al. issn 0075-6458 61 koedoe 48/2 (2005) koedoe 48/2 (2005) 62 issn 0075-6458 table 1 synoptic table of the major plant communities of marakele national park for symbols see text, constancy values = percentage community # 1 2 3 4 5 land type ib/ad ad ib/fa fa ib/fa altitude (x 100 = m) 11-17 12-15 15-20 13-16 13-14 slope g,l g,l m m l aspect all n s,e n n,e surface rock cover o,h o,m h h o species group a pappea capensis 59 mimusops zeyheri 59 olea europaea subsp. africana 55 cussonia paniculata 52 33 diospyros whyteana 52 euphorbia ingens 52 rhus leptodictya 45 cryptolepis transvaalensis 45 podocarpus latifolius 38 maytenus undata 38 zanthoxylum capense 34 myrsine africana 31 canthium gilfillanii 31 calpurnia aurea 31 olea capensis 31 ficus sur 31 grewia occidentalis 31 species group b acacia caffra 65 elionurus muticus 61 faurea saligna 57 aristida congesta 57 solanum incanum 48 dichrostachys cinerea 43 pterocarpus rotundifolius 43 39 eragrostis capensis 39 sida dregei 39 bewsia biflora 39 pogonarthria squarrosa 39 eragrostis lehmanniana 39 perotis patens 35 vernonia oligocephala 39 species group c eragrostis curvula 31 65 dombeya rotundifolia 52 65 33 euclea crispa 41 43 grewia flavescens 34 39 berchemia zeyheri 48 39 ziziphus mucronata 38 35 species group d panicum natalense 75 anthospermum hispidula 64 urelyterum agropyroides 57 thesium utile 52 rhynchosia monophylla 50 acalypha angustata 50 protea caffra 48 aeschynomene rehmannii 45 indigofera hedyantha 43 monocymbium ceresiiforme 43 38 tristachya rehmannii 41 cheilanthus hirta 39 dicoma anomala 39 vernonia galpinii 39 xerophyta retinervus 36 chaetacanthus costatus 36 senecio venosus 36 rhynchosia nitens 34 indigofera mollicoma 34 pentanisia angustifolia 32 parinari capensis 32 species group e eragrostis racemosa 70 82 trachypogon spicatus 30 80 bulbostylis burchellii 30 73 themeda triandra 78 59 gnidia capitata 43 34 species group f setaria lindenbergiana 100 rhoicissus revoilii 89 hypoestes forskaolii 78 pseudolachnostylis maprouneifolia 78 strychnos pungens 72 diplorhynchus condylocarpon 67 ochna pulchra 67 elephantorrhiza burkei 61 littonia modesta 61 stylochiton natalense 56 maytenus tenuispina 56 lantana rugosa 50 tephrosia rhodesica 44 cryptolepis oblongifolia 44 tapiphyllum parvifolium 44 kalanchoe paniculata 39 apodytes dimidiata 39 aristida scabrivalvus 33 turraea obtusifolia 33 ancylbotrys capensis 33 talinum caffrum 33 species group g englerophytum magalismontanum 43 100 aristida transvaalensis 50 61 rhynchosia totta 52 56 loudetia simplex 86 50 tephrosia longipes 68 50 stachys natalensis 36 50 sphenostylis angustifolia 45 33 rhus dentata 45 33 species group h burkea africana 43 94 combretum molle 30 94 lannea discolor 43 83 ozoroa paniculosa 30 50 table 1 (continued) community # 1 2 3 4 5 land type ib/ad ad ib/fa fa ib/fa altitude (x 100 = m) 11-17 12-15 15-20 13-16 13-14 slope g,l g,l m m l aspect all n s,e n n,e surface rock cover o,h o,m h h o issn 0075-6458 63 koedoe 48/2 (2005) phyllanthus parvulus 35 78 raphionachne galpinii 43 56 setaria sphacelata 78 44 heteropogon contortus 87 44 vitex rehmannii 39 61 species group i andropogon schirensis 43 80 61 commelina africana 39 64 100 diheteropogon amplectans 30 73 67 schizachyrium sanguineum 30 50 33 fadogia homblei 39 57 50 melinis repens 78 36 72 brachiaria serrata 70 39 44 species group j vangueria infausta 34 74 55 94 asparagus transvaalensis 38 39 43 56 pellaea calomelanos 31 30 50 89 cheilanthus viridis 38 56 species group k xyris capensis 88 andropogon huilensis 88 miscanthus junceus 63 fuirena pubescens 56 monopsis decipiens 50 helichrysum aureonitens 50 ischaemum fasciculatum 44 aristida junciformis 44 ascolepis capensis 44 sebaea leiostyla 44 hypericum lalandii 44 syzygium cordatum 38 verbena bonariensis 38 cyperus thorncroftii 38 cliffortia linaerifolia 31 arundinella nepalensis 31 drosera madagascariensis 31 results the vegetation composition of the study area is summarised in a synoptic table (table 1). all species with a constancy value of more than 30 % in any community were included in the synoptic table. this table therefore shows the most frequently found species of each major community and summarises the relationship between the five major plant communities recognised. classification the following major communities were identified: a. forest communities: 1. olea europaea subsp. africana diospyros whyteana major community b. sour bushveld and north eastern mountain sourveld communities: 2. acacia caffra-heteropogon contortus major community 3. protea caffra-loudetia simplex major community 4. burkea africana-setaria lindenbergiana major community c. wetland communities: 5. andropogon huilensis-xyris capensis major community. description of the major communities a forest communities 1. olea europaea subsp. africana-diospyros whyteana major community this major community occurs as forests in the kloofs, as dense bush clumps on south and east facing slopes and as bush clumps on termitaria. the kloofs are the least exposed of the geomorphology classes found in the study area, with water in the spruits. the species composition of the olea europaea subsp. africana-diospyros whyteana major community is given in table 1. these forests and dense bush clumps have table 1 (continued) community # 1 2 3 4 5 land type ib/ad ad ib/fa fa ib/fa altitude (x 100 = m) 11-17 12-15 15-20 13-16 13-14 slope g,l g,l m m l aspect all n s,e n n,e surface rock cover o,h o,m h h o 1996), after a twinspan analysis (hill 1979), the output of the resulting classification was imported into a spreadsheet, for refinement by braun-blanquet procedures (behr & bredenkamp 1988; bredenkamp & brown 2003). the final classification of the relevés was then interpreted for identification of major communities, where-after a synoptic table was compiled by calculating the percentage constancy of each species in each of the major communities identified. the result is given in table 1. the following diagnostic species, all being trees or shrubs (species group a, table 1): pappea capensis, mimusops zeyheri, olea europaea subsp. africana, cussonia paniculata, diospyros whyteana, euphorbia ingens, rhus leptodictya, cryptolepis transvaalensis, podocarpus latifolius, maytenus undata, zanthoxylum capense, myrsine africana, canthium gilfillanii, calpurnia aurea, ficus sur, olea capensis and grewia occidentalis. although various different plant communities can be recognised within these forests and bush clumps (van staden 2003), species such as podocarpus latifolius, mimusops zeyheri, diospyros whyteana, canthium gilfillanii and olea capensis are prominent throughout the range of this vegetation. grass and forb species are scanty under the dense forest vegetation, only eragrostis curvula, asparagus transvaalensis, pellaea calomelanos and cheilanthes viridis occurred with a constancy of more than 30 %. the olea europaea subsp. africana-diospyros whyteana major community is floristically related to the sour bushveld communities (communities 2, 3 and 4) within the park (species group j), but shows particular affinity to the acacia caffra-heteropogon contortus major community through species group c. coetzee (1975) described a related community from the rustenburg nature reserve as hypoestes verticillaris-mimusops zeyheri forests, and westfall (1981) described a similar community from the farm groothoek in the waterberg as kloof forest communities on moderately deep soils in moist, sheltered habitats. du preez et al. (1991) classified this type of forest as typical afromontane forests, where these forests occur in specific niches in deep valleys, protected gorges, crevices and ravines along the eastern and western slopes of the drakensberg mountain range. coetzee et al. (1976) described termitaria bush clump communities from the nylsvley nature reserve, which have diagnostic species similar to the bush clump communities on termitaria in the marakele national park. b sour bushveld and north eastern mountain sourveld communities 2. acacia caffra-heteropogon contortus major community this major community represents a sour bushveld (acocks 1988) with open to dense savanna typically found on the slopes of the waterberg, and more open woodland or grassland on the summits. this open to closed woodland vegetation is found on gradual to moderately steep slopes (mainly footslopes) of outcrops, hills, ridges and mountains, which are widely distributed over marakele national park. it is restricted to shallow and rocky soils with a relatively high nutritional status (coetzee 1975), derived from diabase and also from quartzite (bezuidenhout et al. 1994). many of the sample plots were located on diabase, which forms the substrate of this major community. on the lower, less rocky slopes with deeper soils the acacia caffra-dominated vegetation often grades into communities of the plains forming a mixed thornveld, as also described from the rustenburg nature reserve (coetzee 1975). on the other hand, the most mesic acacia caffra-dominated vegetation shows some affinity to higher altitude protea caffra-dominated vegetation that normally occurs at cooler sites (species group e, table 1), at altitudes above the acacia caffra-dominated vegetation, as also reported by coetzee (1975). the soils are mainly shallow, of the mispah or glenrosa form, though deeper soils of the hutton or clovelly form are encountered at the foot of the mountain slopes (macvicar et al. 1977). the soil depth varies from 100 mm to more than 1000 mm (land type survey staff 1988). the species composition of the acacia caffra-heteropogon contortus major community is given in table 1. this major community is differentiated from the other major communities in the area by the following koedoe 48/2 (2005) 64 issn 0075-6458 diagnostic plant species (species group b, table 1): acacia caffra, faurea saligna, dichrostachys cinerea, pterocarpus rotundifolius, elionurus muticus, aristida congesta, eragrostis capensis, bewsia biflora, perotis patens, eragrostis lehmanniana, pogonarthria squarrosa, solanum incanum, sida dregei and vernonia oligocephala. acacia caffra is mostly a dominant woody species, with other prominent species such as faurea saligna, burkea africana, dichrostachys cinerea, pterocarpus rotundifolius, vitex rehmannii, vangueria infausta, ozoroa paniculosa, lannea discolor and combretum molle also prominently present in certain plant communities. quite often the woody species are grouped in bush clumps, resulting in a savanna with scattered individual trees and bush clumps. the prominent grass species include trachypogon spicatus, themeda triandra, setaria sphacelata, schizachyrium sanguineum, melinis repens, heteropogon contortus, eragrostis racemosa, diheteropogon amplectens, brachiaria serrata, andropogon schirensis and the diagnostic elionurus muticus and eragrostis lehmanniana. the most frequently found forbs include pellaea calomelanos, gnidia capitata, fadogia homblei, bulbostylis burchellii, asparagus transvaalensis, solanum incanum and vernonia oligocephala. the sour bushveld (acocks 1988) communities that occur in the park (communities 2, 3 and 4) are all floristically related, indicated by species group i, but also through species groups e, g and h. descriptions of acacia caffra-dominated vegetation are given by bezuidenhout et al. (1988) from the vredefort dome area, bezuidenhout & bredenkamp (1991) from the north-west province grasslands and grobler et al. (2002) from the johannesburgpretoria area. particularly good examples (eustachys mutica-acacia caffra woodlands) are provided by coetzee (1975) from the rustenburg nature reserve, bezuidenhout et al. (1994) from the slopes of the hills and ridges in the north-west province, and coetzee et al. (1994, 1995) from the magaliesberg in the pretoria area. coetzee (1974) described acacia caffra savannas on diabase and in sheltered valleys from the jack scott nature reserve. westfall (1981) described a similar community as woodland on moderately deep soils in moderately exposed habitats, from the farm groothoek in the waterberg. 3. protea caffra-loudetia simplex major community the protea caffra-loudetia simplex major community is representative of acocks’ (1988) sour bushveld on moderately deep to deep soils in moderately exposed habitats. this mountain bushveld is found on gradual to steep rocky hills and ridges in the marakele national park. this vegetation type is prominent on the higher altitude slopes and crests. the soils are mainly very shallow, of the mispah and glenrosa forms. the soil depth varies from 10 mm to 500 mm (land type survey staff 1988). the species composition of the protea caffra-loudetia simplex major community is given in table 1. this major community is differentiated by the following diagnostic plant species (species group d, table 1): protea caffra, urelyterum agropyroides, tristachya rehmannii, panicum natalense, monocymbium ceresiiforme, xerophyta retinervus, vernonia galpinii, thesium utile, senecio venosus, rhynchosia nitens, rhynchosia monophylla, pentanisia angustifolia, parinari capensis, indigofera mollicoma, indigofera hedyantha, dicoma anomala, cheilanthus hirta, chaetacanthus costatus, anthospermum hispidula, aeschynomene rehmannii and acalypha angustata. this major community is dominated by the tree protea caffra. other prominent woody species include vangueria infausta, rhus dentata and englerophytum magalismontanum. the most prominent grasses that are often found in this major community are trachyissn 0075-6458 65 koedoe 48/2 (2005) pogon spicatus, themeda triandra, schizachyrium sanguineum, melinis repens, loudetia simplex, eragrostis racemosa, diheteropogon amplectens, brachiaria serrata, aristida transvaalensis and andropogon schirensis. many forb species are present in this vegetation; the most frequently encountered include tephrosia longipes, stachys natalensis, sphenostylis angustifolia, rhynchosia totta, pellaea calomelanos, gnidia capitata, fadogia homblei, commelina africana, bulbostylis burchellii and asparagus transvaalensis. many authors described protea caffra-dominated vegetation, mainly from sour bushveld and bankenveld, including coetzee (1974, 1975) from the magaliesberg area, bredenkamp & theron (1978) from the suikerbosrand, behr & bredenkamp (1988), from the witwatersrand, bezuidenhout et al. (1994) from the gatsrand area in north-west province, coetzee et al. (1995), bredenkamp & brown (1998a; 1998b) from the natural areas of the western metropolitan local council and grobler et al. (2002) within various natural open spaces in gauteng. this emphasises the floristic relationships between bankenveld, sour bushveld and the drakensberg area (bredenkamp & brown 2003). 4. burkea africana-setaria lindenbergiana major community this major community represents sour bushveld (acocks 1988). it occurs on gentle to moderately steep slopes of rocky sandstone hills, where soils are litholitic and large rock boulders cover 53 % of the soil surface. aspects are mostly northerly, and these slopes are warm and dry. the soils are mainly of the mispah and glenrosa forms. the soil varies from 10 mm to more than 500 mm in depth (land type survey staff 1988). this vegetation type is also found at the northernmost areas of the bankenveld in the pretoria and rustenburg areas and is found at high altitudes on warm northern and northeastern aspects with gradual to very steep slopes (bredenkamp & brown 2003). the shallow soils are covered with large boulders and smaller rocks that in some cases provide moister microhabitats than the surrounding areas. the species composition of the burkea africana-setaria lindenbergiana major community is given in table 1. this major community is differentiated by the following diagnostic plant species (species group d, table 1): tapiphyllum parvifolium, strychnos pungens, rhoicissus revoilii, pseudolachnostylis maprouneifolia, ochna pulchra, maytenus tenuispina, elephantorrhiza burkei, diplorhynchus condylocarpon, cryptolepis oblongifolia, apodytes dimidiata, acylobotrys capensis, setaria lindenbergiana, aristida scabrivalvus, turraea obtusifolia, tephrosia rhodesica, talinum caffrum, stylochiton natalense, littonia modesta, lantana rugosa, kalanchoe paniculata and hypoestes forskaolii. apart from the above prominent diagnostic species, other prominent woody species include vitex rehmannii, vangueria infausta, rhus dentata, ozoroa paniculosa, lannea discolor, englerophytum magalismontanum, combretum molle and burkea africana. grass species that are prominent in this community include setaria sphacelata, schizachyrium sanguineum, melinis repens, loudetia simplex, heteropogon contortus, diheteropogon amplectens, brachiaria serrata, aristida transvaalensis and andropogon schirensis. forbs include tephrosia longipes, stachys natalensis, sphenostylis angustifolia, rhynchosia totta, pellaea calomelanos, fadogia homblei, cheilanthus viridis and asparagus transvaalensis. coetzee (1975) described a similar community as the barleria bremekampiidiplorhynchus tree savanna from rustenburg nature reserve, and westfall (1981) described a similar community as a woodland, representative of acocks’ (1988) sour koedoe 48/2 (2005) 66 issn 0075-6458 bushveld, on moderately deep to deep soils in moderately exposed habitats from the waterberg. further examples of this vegetation were described from the scarps and slopes of the jack scott (coetzee 1974), from the warm temperate mountain bushveld within the pretoria-witbank-heidelberg areas by (coetzee et al. 1993), from the melville koppies nature reserve (ellery 1994), from the natural areas of the western local metropolitan council of gauteng by bredenkamp & brown (1998a) and within various natural open spaces in gauteng by grobler et al. (2002). 5. andropogon huilensis-xyris capensis major community major many streams arise in seepage areas, for instance on mountain slopes within marakele national park. these are seasonally or perennially waterlogged sponges, with vegetation dominated by sedges and other hygrophilous angiosperms and mosses (noble & hemens 1978). this major wetland community occurs along streams and tributaries of the matlabas, mamba and sterkstroom rivers and shallow submerged marshy areas or sponges. the soils are wet, mainly of the avalon, hutton, katspruit, oakleaf and westleigh forms, derived from sandstone of the sandriviersberg formation (de vries 1968, 1969). the soil depth varies between 300–1 200 mm (land type survey staff 1988). the species composition of this wetland major community is given in table 1. this major community is differentiated by the following diagnostic plant species (species group k): syzygium cordatum, cliffortia linaerifolia, andropogon huilensis, aristida junciformis, arundinella nepalensis, ischaemum fasciculatum, miscanthus junceus, ascolepis capensis, cyperus thorncroftii, drosera madagascariensis, fuirena pubescens, helichrysum aureonitens, hypericum lalandii, monopsis decipiens, sebaea leiostyla, verbena bonariensis and xyris capensis. coetzee (1975) described similar communities from the rustenberg nature reserve as the aristida junciformis-arundinella nepalensis grassland and the pteridium aquilinum-phragmites mauritianus reedswamp. discussion the vegetation of the study area was classified in a hierarchical, plant sociological system by using the braunblanquet technique. the floristic data from 130 relevés were classified to identify five major plant communities. these plant communities were ecologically interpreted by habitats. these major plant communities should form the basis for a management plan, and should also be the basis for a more detailed classification of the vegetation of this national park. according to white (1978) the majority of the tree species in south african afromontane forests are widespread. amongst them are, inter alia, podocarpus latifolius, olea capensis and myrsine africana that occur as diagnostic species in the olea europaea subsp. africana-diospyros whyteana major community within the study area. these forests on the waterberg have a clear floristic affinity to the afromontane forests found in the drakensberg (du preez et al. 1991), indicating a much more widespread distribution of forests at previous times, with the waterberg forests being a western relict (bredenkamp et al. 2002). the above mentioned assemblage of species could almost be used to define the afromontane region as a whole. not one species occurs throughout, but most species of the assemblage are present on virtually every “island” of afromontane forest in the region. it is interesting to note that the vegetation on the termitaria have species similar to the forest communities, and are therefore included into this major community. the greater water holding capacity of the soils worked over by the termites renders these habitats suitable for forest development (coetzee et al. 1976). issn 0075-6458 67 koedoe 48/2 (2005) the acacia caffra-heteropogon contortus major community on footslopes, protea caffra-loudetia simplex major community on cooler southerly facing slopes and the burkea africana-setaria lindenbergiana major community on the warmer northerly facing slopes have a clear floristic relationship with similar plant communities in the bankenveld. the relationships between sour bushveld, sourish mixed bushveld and bankenveld woodland communities (acocks 1988) are clearly indicated by bredenkamp & brown (2003) and can further be demonstrated from studies by van vuuren & van der schijff (1970), du plessis (1972), theron (1973), coetzee (1975), van der meulen (1979), westfall (1981) and coetzee (1993). furthermore, coetzee (1993), o’connor & bredenkamp (1997) and bredenkamp & brown (2003) indicated that the bankenveld vegetation also shows an affinity to the drakensberg flora. this relationship may also be seen in the presence of the afromontane forests in the drakensberg, bankenveld and waterberg. in the cooler, southern parts of the bankenveld, acacia caffra-dominated vegetation is usually found on lower and warmer northfacing slopes, e.g., in the vredefort dome area, though in the warmer northern parts e.g. the marakele national park, it may be found at higher altitudes, on crests or southfacing slopes (coetzee 1975; bezuidenhout et al. 1994; coetzee et al. 1995). due to the complex mosaic distribution pattern of the sub-communities that are included under the major communities, a vegetation map is more appropriate when the details of the sub-communities are published. references acocks, j.p.h. 1988. veld types of south africa. memoirs of the botanical survey of south africa 57: 1-146. barkman, j.j., j. moravec & s. rauchert. 1964. kritische bemerkungen und vorschläge zur quantitativen vegetationanalyse. acta botanica neerlandia 13: 394-449. behr, c.m. & g.j. bredenkamp. 1988. a phytosociological classification of the witwatersrand national botanical garden. south african journal of botany 54: 525-533. bezuidenhout, h. 1992. verslag oor die abiotiese komponent van die kransberg nasionale park. unpublished internal report. bezuidenhout, h., h.c. biggs & g.j. bredenkamp. 1996. a process supported by the utility bbpc for analysing braun-blanquet data on a personal computer. koedoe 39(1): 107-112. bezuidenhout, h., & g.j. bredenkamp. 1991. the vegetation of the bc land type in the western transvaal grassland, south africa. phytocoenologia 19: 497-518. bezuidenhout, h., g.j. bredenkamp & j.h. elsenbroek. 1988. die plantegroei van die alkaligraniet en aangrensende kwartsiet in die vredefortkoepel noordwes van parys. suidafrikaanse tydskrif vir natuurwetenskap en tegnologie 7: 4-9. bezuidenhout, h., g.j. bredenkamp & g.k. theron.1994. syntaxonomy of the vegetation of the fb land type in the western transvaal grassland, south africa. south african journal of botany 60: 72-80. bredenkamp, g.j. & l.r. brown. 1998a. a vegetation assessment of the open spaces in the western metropolitan local council area. pretoria: ekotrust cc. bredenkamp, g.j. & l.r. brown.1998b. a vegetation assessment of open spaces in the northern areas of the northern metropolitan local council. pretoria: ekotrust cc. bredenkamp, g.j. & l.r. brown. 2003. a reappraisal of acocks’ bankenveld: origin and diversity of vegetation types. south african journal of botany 69(1): 7-26. bredenkamp, g.j., f. spada & e. kazmierczak. 2002. on the origin of northern and southern hemisphere grasslands. plant ecology 163(2): 209-229. bredenkamp, g.j. & g.k. theron. 1978. a synecological account of the suikerbosrand nature reserve. 1. the phytosociology of the witwatersrand geological system. bothalia 12: 513-529. brown, l.r. 1997. a plant ecological study and wildlife management plan of the borakalalo nature reserve, north-west province. ph.d. thesis. university of pretoria, pretoria. coetzee, b.j. 1974. a phytosociological classification of the vegetation of the jack scott nature reserve. bothalia 11: 329 347. coetzee, b.j. 1975. a phytosociological classification of the rustenburg nature reserve. bothalia 11: 561-580. coetzee, b.j. & m.j.a. werger. 1975. on association—analysis and the classification of plant communities. vegetatio 30: 201-206. koedoe 48/2 (2005) 68 issn 0075-6458 coetzee, b.j., f. van der meulen, s. zwanziger, p. gonsalves & p.j. weisser. 1976. a phytosociological classification of the nylsvley nature reserve. bothalia 12: 137-160. coetzee, b.j., p. van wyk, w.p.d. gertenbach, a. hall-martin & s.c.j. joubert. 1981. ‘n plantekologiese verkenning van die waterberggebied in die noord-transvaalse bosveld. koedoe 24:1-23. coetzee, j.p. 1993. phytosociology of the ba and ib land types in the pretoria-witbank-heidelberg area. msc thesis, university of pretoria, pretoria. coetzee, j.p., g.j. bredenkamp & n. van rooyen. 1993. the sub-humid warm temperate mountain bushveld plant communities of the pretoriawitbank-heidelberg area. south african journal of botany 59(6): 623-632. coetzee, j.p., g.j. bredenkamp & n. van rooyen. 1994. an overview of the physical environment and vegetation units of the ba and ib land types of the pretoria-witbank-heidelberg area. south african journal of botany 60(1): 49-61. coetzee, j.p., g.j. bredenkamp & n. van rooyen. 1995. plant communities of the sub-humid cool temperate mountain bushveld in the pretoria witbank heidelberg area. south african journal of botany 61: 114-122. de vries, w.c.p. 1968-69. stratigraphy of the waterberg system in the southern waterberg area, northwestern transvaal. annals of the geological survey in south africa 43-56. du plessis, c.j. 1972. ‘n floristies-ekologiese studie van die plaas doornkop in die distrik middelburg, transvaal. msc thesis, university of pretoria, pretoria, du preez, p.j., g.j. bredenkamp & h.j.t. venter. 1991. the syntaxonomy and synecology of the forests in the eastern orange free state, south africa. i. the podocarpetalia latifolii. south african journal of botany 57: 198-206. edwards, d. 1972. botanical survey and agriculture. proceedings of the grassland society of south africa 7: 15-19. ellery, w.n. 1994. the vegetation ecology of melville koppies nature reserve and louw geldenhuys view site: proposals for their management. johannesburg: botanical society of south africa. gertenbach, w.p.d. 1987. ‘n ekologiese studie van die suidelikste mopanieveld in die nasionale krugerwildtuin. d.sc proefskrif (ongepubliseer). universiteit van pretoria. grobler, c.h., g.j. bredenkam. & l.r.brown. 2002. natural woodland vegetation and plant species richness of the urban open spaces in gauteng, south africa. koedoe 45: 19-34. hill, m.o. 1979. twinspan a fortran program for arranging multivariate data in an ordered two way table by classification of individuals and attributes. cornell university, ithaca, new york. land type survey staff. 1988. land types of the maps 2426 thabazimbi, 2428 nylstroom. memoirs of the agricultural natural resoures of south africa 10: 1-431. low, a.b. & a.g. rebelo. 1996. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. macvicar, c.n., r.f. loxton, j.j.n. lambrechts, j. le roux, j.m. de villiers, e. verster, f.r. merry-weather, t.h. van rooyen & h.j. von m. harmse. 1977. grondklassifikasie, 'n binomiese sisteem vir suid afrika. pretoria: departement lanbou tegniese dienste. mueller-dombois, d. & h. ellenberg. 1974. aims and methods of vegetation ecology. new york: wiley. noble, r.g. & j. hemens. 1978. inland water ecosystems in south africa a review of research needs. south african natural science progress report 34: 1-150. o’connor, t.g.. & g.j. bredenkamp. 1997. grassland. pp. 215-257. in: cowling r.m., d.m. richardson & s.m. pierce (eds.). vegetation of southern africa. cambridge: cambridge university press. rutherford, m.c. & r.h. westfall. 1994. biomes of southern africa: an objective categorization. memoirs of the botanical survey of south africa 63: 1-94 south african commission for stratigraphy (sacs). 1980. statigraphy of south africa. part 1 (comp. l.e. kent). lithostratigraphy of the republic of south africa, south west africa/ namibië and the republics of bophuthatwana, transkei, and venda. pretoria: government printer. (handbook for geological survey in south africa 8.) theron, g.k. 1973. ‘n ekologiese studie van die plantegroei van die loskopdam natuurreservaat ongepubl. d.sc thesis, university of pretoria, pretoria. van der meulen, f. 1979. plant sociology of the western transvaal bushveld. south africa: syntaxonomic and synecological study. vaduz: cramer. (dissertationes botanica 49.) van rooyen, n. 1983. die plantegroei van die roodeplaatdam natuurreservaat ii. die plantgemeenskappe. south african journal of botany 2: 115 -125. van staden, p.j. 2002. an ecological study of the plant communities of marakele national park. m.sc. thesis, university of pretoria, pretoria. van vuuren, d.r.j. & h.p. van der schijff. 1970. ‘n vergelykende ekologiese studie van die plantegroei van ‘n noordelike en suidelike kloof issn 0075-6458 69 koedoe 48/2 (2005) van die magaliesberg. tydskrif vir natuurwetenskappe 10: 16-75. werger, m.j.a. 1974. on concepts and techniques applied in the zürich-montpellier method of vegetation survey. bothalia 11(3): 309-323. westfall, r.h. 1981. the plant ecology of the farm groothoek, thabazimbi district. m.sc. thesis, university of pretoria, pretoria. westfall, r.h., n. van rooyen & g.k. theron. 1985. the plant ecology of the farm groothoek, thabazimbi district. 2. classification. bothalia 15: 655-688. westhoff, v. & e. van der maarel. 1980. the braun-blanquet approach. pp. 287-399. in: r.h. whittaker (ed.). classification of plant communities. the hague: junk. white, f. 1978. the afromontane region. pp. 465513. in: werger, m.j.a. (ed.). biogeography and ecology of southern africa. the hague: junk. koedoe 48/2 (2005) 70 issn 0075-6458 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <feff00550073006100720065002000710075006500730074006500200069006d0070006f007300740061007a0069006f006e00690020007000650072002000630072006500610072006500200064006f00630075006d0065006e00740069002000500044004600200063006f006e00200075006e00610020007200690073006f006c0075007a0069006f006e00650020006d0061006700670069006f00720065002000700065007200200075006e00610020007100750061006c0069007400e00020006400690020007000720065007300740061006d007000610020006d00690067006c0069006f00720065002e0020004900200064006f00630075006d0065006e00740069002000500044004600200070006f00730073006f006e006f0020006500730073006500720065002000610070006500720074006900200063006f006e0020004100630072006f00620061007400200065002000520065006100640065007200200035002e003000200065002000760065007200730069006f006e006900200073007500630063006500730073006900760065002e002000510075006500730074006500200069006d0070006f007300740061007a0069006f006e006900200072006900630068006900650064006f006e006f0020006c002700750073006f00200064006900200066006f006e007400200069006e0063006f00720070006f0072006100740069002e> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice book review review of national park science: a century of research in south africa by jane carruthers book title: national park science: a century of research in south africa author: jane carruthers isbn: 978-1-107-19144-0 (hardback); 978-1-316-64202-3 (paperback) publisher: cambridge university press, 2017, r1330.00* *book price at time of review review title: review of national park science: a century of research in south africa by jane carruthers reviewer: peter a. novellie1 affiliation: 1conservation services division, sanparks, south africa corresponding author and email: peter novellie, novellie@netactive.co.za how to cite this book review: novellie, p.a., 2018, ‘review of national park science: a century of research in south africa by jane carruthers’, koedoe 60(1), a1550. https://doi.org/10.4102/koedoe.v60i1.1550 copyright: © 2018. the authors. licensee: aosis. this this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. from the perspective of a long-serving scientist – initially of the national parks board, later sanparks – jane carruthers’ book took me on an enchanting and insightful journey. i could readily recognise – and at least partly recollect – the three periods into which she divides the historical narrative, each named in accordance with the prevailing nature conservation paradigm: protecting, preserving and propagating (1900–1960); measuring, monitoring and manipulating (1960–1990); and integration, innovation and internationalisation (1990–2010). the analysis is wide-ranging and thoroughly grounded in numerous sources. particularly, the first and second of the three periods are enriched by many years of research on environmental history and conservation, including a focus on the social and political history of the kruger national park, as well as other protected areas. good scholarship stimulates new ideas, introduces fresh perspectives and alters mindsets. carruthers has achieved this in ways that raise important questions for further analysis. as i travelled through the narrative, with its intriguing reminders of paradigms, institutions and personalities of distant eras, i was struck by my own career-long disregard for the past. i found i had a partial and imperfect knowledge of the contributions of older scientific colleagues whose period of service overlapped mine, and knew little or nothing of those, notably p.j. barnard and t.g. nel, who had departed before my time. during my early years in the organisation, i was dismissive of the work of predecessors of long before. this perspective, as carruthers explains in her preface, is typical of scientists who are trained to feel that there is no point in reading anything but the latest paper. scientists assume that the present includes all pasts, and tend to be impatient with historical context, whereas environmental historians must read all papers to unpack intellectual journeys in light of their political and social contexts (carruthers 2017:xxiv). by the end of the book, i was convinced that an understanding of past journeys, and their contexts, is as important for conservation scientists as it is for historians. like all organisations, sanparks and its predecessor, the national parks board, developed organisational pride – undoubtedly a healthy phenomenon, but which can decline into unwarranted self-congratulation, based on historical half-truths, even outright myths. one long-held myth – that kruger was south africa’s first national park – carruthers exposes as false. the 1960s–1990s were characterised by an element of arrogance and isolationism, convictions that ‘we know best’, and resistance to external influences save those of a select group of trusted collaborators. carruthers indicates how the attitude of this time was not only self-delusional (science in the national parks board was actually behind the times) but also hampered scientific progress. for example, the organisation was extraordinarily slow to embrace the concept of biodiversity. the analysis of publications in the journal koedoe showed that the word ‘biodiversity’ appeared only in 1999 ‘as a keyword for a contribution relating to national parks – almost 20 years after it had entered the lexicon of conservation biology’ (carruthers 2017:209). scientific stagnation was swept away during the period of integration, innovation and internationalisation that followed. political and social transformation in south africa and the world promoted receptivity to new ideas and stimulated the growth of science in national parks. thus, a significant achievement of the book is its analysis of the complex transitions between the three periods, the external developments that influenced and facilitated the transitions, and the processes whereby old ways of thinking were replaced by new ideas. such transitions are in the nature of conservation organisations and sanparks will undoubtedly undergo others in future. a body of theories is developing on how the governance of socio-ecological systems can be transformed from inflexible ‘command and control’ approaches to adaptive governance regimes (e.g. olsson et al. 2006). it could be insightful to examine carruthers’ account of the transition to integration, innovation and internationalisation in light of this theory. importantly, carruthers draws attention to several long-standing problem areas that have remained intransigent, at variance with the flow of transition in national park science. these remain evident to this day. there is no obvious explanation for the shortcomings, and one feels that they merit further investigation. one of the long-standing problem areas is the neglect of the humanities and social sciences in general. the absence of staff members qualified in these fields meant that studies have often been undertaken by untrained people (carruthers 2017:417). richard bell, in his 1995 report on the state of scientific services in the kruger national park, recommended the employment of social scientists. the recommendation went unheeded for many years, and carruthers indicates that the dearth of publications in the humanities, in both kruger and the other national parks, was evident throughout the period covered by her book. a possibly related problem area is the ineffectiveness and limited impact of the social ecology unit of sanparks (carruthers 2017:343), all the more surprising in view of the urgency of reaching out to the historically marginalised communities surrounding national parks. the neglect of social sciences relative to biophysical sciences is not limited to national parks in south africa; it has been reported elsewhere in the world (bennett et al. 2016; correia et al. 2016; moreno et al. 2014). another persistent shortcoming is that of ‘silos’, a lack of collaboration between the national parks board/sanparks and other south african conservation agencies or institutions. as indicated above, isolationism was a particular feature of the 1960s–1990s, but carruthers notes that the lack of collaboration with provincial conservation agencies persisted even into the period of integration, innovation and internationalisation (carruthers 2017): fewer silos and more synergy and collaboration would lead to further integration, innovation and internationalisation to the benefit of conservation science in south africa, and particularly to good and meaningful research in the smaller national parks and protected areas. (p. 464) during my final years with sanparks, the department of environmental affairs repeatedly encouraged collaboration and assistance for particularly certain provincial conservation authorities. it seems worth reflecting why this has been an enduring problem area. a peculiar phenomenon uncovered by carruthers is the apparent discomfort in the national parks board/sanparks with the idea of scientific advisory boards. there was a succession of efforts to provide expert scientific oversight for national parks and other protected areas, starting as early as the 1940s and the 1950s, continuing during the 1980s with the national committee for nature conservation (nacor) and finally, in the mid-1990s, an offer of financial support from the mellon foundation for a guiding body of international scientists. all these efforts came to nothing. carruthers highlights another persistent feature of south africa’s system of protected areas – its misalignment with the international union for conservation of nature (iucn) system of classifying protected areas. harold eidsvik’s 1996 report on national parks pointed out that some did not qualify as such in terms of the criteria of the day. the protected areas act of 2003 ignored the iucn classification, and sanparks continues to manage parks such as bontebok, which do not qualify. a serious problem area is the long-standing discrepancy in research capacity and output between kruger and the other national parks. this is apparent from the numbers of publications from each national park from 2012 to 2016 (carruthers 2017:447–448). an aspect that remains to be clarified is the extent to which the achievement of the conservation objectives set in park management plans may have been compromised in the poorly researched parks. carruthers draws attention to the potential for collaboration and partnerships with other conservation agencies and universities. however, collaboration by itself does not provide a ready solution to the imbalance between parks because the research community is strongly biased towards the larger and older national parks (van wilgen et al. 2016). this is not a uniquely south african phenomenon; correia et al. (2016) show that the larger, older and more protected reserves in ecuador and peru were more likely to have high scientific production. it appears that the potential of a park to deliver science outputs will be influenced by inherent, ‘given’ features – such as size and age – that are beyond the control of management (smit et al. 2017) and do not bear any relationship with the need for research as a guide to realise conservation objectives. smit et al. (2017) suggest that this bias can be mitigated by various measures designed to attract research collaboration. an influence that looms large in my recollection of 32 years with national parks, but which seems understated in the book, is the provision in both the national parks act of 1976 and the protected areas act of 2003 for the organisation to retain self-generated income. its influence was not only a feature of the era of integration, innovation and internationalisation (carruthers 2017:368–369); it was apparent from the 1980s. it empowered the national parks board/sanparks and gave it financial flexibility, but i have the feeling that it may have contributed, directly or indirectly, to some of the problem areas outlined above. it engendered an organisational pride in partial independence of the national fiscus, understandably more among staff members responsible for income generation than among scientists. it introduced a degree of cynicism, often diplomatically hidden but nevertheless prevalent, regarding expenditure on enterprises that do not generate income. this attitude probably constrained significant investments in new fields, however deserving, such as staff qualified in the social sciences, outreach to communities through social ecology, or on collaboration and support for needy provincial conservation authorities. arguably perhaps, it may also account for the apparent discomfort with scientific advisory boards, whose recommendations would often beat variance with the main financial drivers of the organisation. i remember being sent to attend national committee for nature conservation (nacor) meetings because it was politic for the national parks board to show face at that forum, knowing well that my organisation had its own agenda for park expansion and had no intention of following nacor priorities. this agenda prioritised park expansion to enhance ‘big five’ ecotourism prospects, so as to extend the kruger model of income generation to other parks. this is entirely understandable in view of the provisions of the national parks act, which gave the parks board stronger potential to fund land acquisition than nacor could provide. dependence on self-generated income also means that ‘money source’ parks (notably kruger) subsidise other parks, the ‘money sinks’. some of the provincial conservation agencies came to adopt financing models similar to that of sanparks. this makes it very difficult to align south african protected areas with the iucn classification and explains why bontebok national park has remained in the sanparks estate. financial viability requires a balance between money sources and money sinks, and no one is keen to take on the sinks. measures to attract research collaboration, such as providing and servicing research accommodation, are undoubtedly easier to establish in ‘money source’ parks, and it will require innovative approaches to attract collaborators to the money sinks. clearly sanparks’ financial resources allow only limited improvements to science capacity, and initiatives are needed to access additional funding sources. carruthers’ account of the role of the mellon foundation, and of other donors, provides a valuable record of how helpful such sources can be in strengthening science capacity. it is important to understand how sanparks came to exploit windows of opportunity in accessing such funding sources. a vital link in this process does not feature in the book, namely the initiative of middle level scientists who engaged with key individuals in funding organisations to identify opportunities and who convinced their superiors in sanparks of the potential of these opportunities. a few individual scientists in sanparks have been influential in brokering access to important sources of financing. these initiatives are undocumented, least of all in the job descriptions of the scientists involved, and so are easily overlooked. carruthers’ history of science in national parks not only recognises significant and valuable achievements but is most valuable for a critical and sometimes unflattering analysis and for highlighting key questions for further investigation. in recent years, sanparks scientists have entered a new phase, embracing new theoretical principles to manage complex, multi-actor socio-ecological systems. these include broadening participation and collaboration to harness diversity, social learning as concerted action, adaptive governance as a prerequisite for adaptive management and the value of transdisciplinarity. carruthers’ emphasis on collaborative research, and on understanding past intellectual journeys and their contexts, resonates well with these new principles. her work is an essential reading for sanparks scientists embarking on the new phase of science. references bennett, n.j., roth, r., klain, s.c., chan, k., christie, p., clark, d.a. et al., 2016, ‘conservation social science: understanding and integrating human dimensions to improve conservation’, biological conservation 205, 93–108. https://doi.org/​10.1016/j.biocon.2016.10.006 carruthers, j., 2017, national park science: a century of research in south africa, cambridge university press, cambridge, united kingdom. correia, r.a., malhado, a.c.m., lins, l., gamarra, n.c., bonfim, w.a.g., valencia aguilar, a. et al., 2016, ‘the scientific value of amazonian protected areas’, biodiversity and conservation 25(8), 1503–1513. https://doi.org/10.1007/s10531-016-1122-x moreno, j., palomo, i., escalera, j., martín-lópez, b. & montes, c., 2014, ‘incorporating ecosystem services into ecosystem-based management to deal with complexity: a participative mental model approach’, landscape ecology 29, 1407–1421. https://doi.org/10.1007/s10980-014-0053-8 olsson, p., gunderson, l.h., carpenter, s.r., ryan, p., lebel, l., folke, c. et al., 2006, ‘shooting the rapids: navigating transitions to adaptive governance of social-ecological systems’, ecology and society 11(1), 18, viewed 12 july 2018, from http://www.ecologyandsociety.org/vol11/iss1/art18/ smit, i.p.j., roux, d.j., swemmer, l.k., boshoff, n. & novellie, p., 2017, ‘protected areas as outdoor classrooms and global laboratories: intellectual ecosystem services flowing to-and-from a national park’, ecosystem services 28, 238–250. http://doi.org/10.1016/j.ecoser.2017.05.003 van wilgen, b.w., boshoff, n., smit, i.p.j., solano-fernandez, s. & van der walt, l., 2016, ‘a bibliometric analysis to illustrate the role of an embedded research capability in south african national parks’, scientometrics 107, 185–212. https://doi.org/10.1007/s11192-016-1879-4 article information authors: rina c.c. grant1 mike j.s. peel2 hugo bezuidenhout3 affiliations: 1scientific services [kruger national park], south african national parks, south africa 2animal production institute, agricultural research council, south africa 3conservation services, south african national parks, south africa correspondence to: rina grant email: rina.grant@sanparks.org postal address: private bag x402, skukuza 1350, south africa dates: received: 04 june 2010 accepted: 10 mar. 2011 published: 13 may 2011 how to cite this article: grant, r.c.c., peel, m.j.s. & bezuidenhout, h., 2011, ‘evaluating herbivore management outcomes and associated vegetation impacts’, koedoe 53(2), art. #1008, 15 pages. doi:10.4102/koedoe.v53i2.1008 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) evaluating herbivore management outcomes and associated vegetation impacts in this essay... open access • abstract • introduction    • outcomes of plant–herbivore interaction for biodiversity • the adaptive management approach    • defining the desired state or outcomes of herbivore management       • economic objectives      • biodiversity objectives      • herbivory objectives    • indicators of not achieving herbivore objectives      • vegetation changes   • herbivore numbers and distribution   • thresholds of potential concern   • monitoring change in vegetation and herbivores    • evaluation of thresholds of potential concern      • vegetation       • herbivore threshold of potential concerns    • changes in herbivore population • alternative approaches to monitoring?    • site selection    • alternative thresholds of potential concern    • animal condition and nutritional status    • energy flows and sustainability    • economic return • discussion • conclusion • references abstract (back to top) african savannas are characterised by temporal and spatial fluxes that are linked to fluxes in herbivore populations and vegetation structure and composition. we need to be concerned about these fluxes only when management actions cause the system to shift towards a less desired state. large herbivores are a key attribute of african savannas and are important for tourism and biodiversity. large protected areas such as the kruger national park (knp) manage for high biodiversity as the desired state, whilst private protected areas, such as those adjacent to the knp, generally manage for high income. biodiversity, sustainability and economic indicators are thus required to flag thresholds of potential concern (tpcs) that may result in a particular set of objectives not being achieved. in large conservation areas such as the knp, vegetation changes that result from herbivore impact, or lack thereof, affect biodiversity and tpcs are used to indicate unacceptable change leading to a possible loss of biodiversity; in private protected areas the loss of large herbivores is seen as an important indicator of economic loss. therefore, the first-level indicators aim to evaluate the forage available to sustain grazers without deleteriously affecting the vegetation composition, structure and basal cover. various approaches to monitoring for these indicators were considered and the importance of the selection of sites that are representative of the intensity of herbivore use is emphasised. the most crucial step in the adaptive management process is the feedback of information to inform management decisions and enable learning. feedback loops tend to be more efficient where the organisation’s vision is focused on, for example, economic gain, than in larger protected areas, such as the knp, where the vision to conserve biodiversity is broader and more complex. conservation implications: in rangeland, optimising herbivore numbers to achieve the management objectives without causing unacceptable or irreversible change in the vegetation is challenging. this manuscript explores different avenues to evaluate herbivore impact and the outcomes of management approaches that may affect vegetation. introduction (back to top) large mammals are a key attribute of african savannas and many national parks and private protected areas have been developed with the aim of protecting and benefitting from these large mammals. these aims are reflected in the mission statement of the south african national parks (sanparks), namely ‘to develop and manage a system of national parks that is representative of the biodiversity,landscapes, and associated heritage assets of south africa, for the sustainable use and benefit of all’ (south african national parks 2008:4). the herbivore management policy of sanparks states that herbivores are managed primarily as ecosystem drivers and is directed by how much change (through herbivore impacts) is acceptable within the desired state (sanparks 2006). it thus focuses on the importance of spatial heterogeneity and temporal fluxes, and the role of landscape patchiness and disturbance in promoting ecosystem resilience and biodiversity (see grant et al. 2011; walker & goodman 1983). large private protected areas adjacent to the kruger national park (knp) have embraced this philosophy since the removal of the fence between them, but the fact that most function at different spatial scales (peel, biggs & zacharias 1999) influences their specific objectives. private protected areas also have a more explicit objective of being economically sustainable; for example, the primary objective statement of the associated private nature reserves is ‘to provide for ecologically and aesthetically sustainable (nonconsumptive and consumptive) use of the area for its owners, based on wildlife-focused recreation, tourism and hunting’. (peel, stalmans & anderson 2009). scale also influences the management of smaller national parks, which have similar biodiversity objectives to the larger parks, but because animal movement is restricted and areas tend to be less heterogeneous these areas present a unique set of management challenges. we aim to record the lessons learnt during the adaptive management process, including the development of thresholds of concern, for change in vegetation and herbivore numbers in sanparks and private protected areas. the application of these thresholds is discussed using data collected in the knp, the associated private nature reserves, sabi sand wildtuin, some smaller national parks and privately owned protected areas. outcomes of plant–herbivore interaction for biodiversity biodiversity as referred to in this section is as defined by noss (1990). herbivores act not only as important drivers of the savanna ecosystem but also as responders to changes in the ecosystem. the main concerns of plant–herbivore interactions therefore relate to loss of vegetation heterogeneity (and herbivore composition and numbers) as described by grant et al. (2011). these concerns are related to the outcomes of the ecological processes and their effect on biodiversity as a reflection of the predefined desired state (biggs & rogers 2003). this management philosophy allows significantly more flux in the system than the more precautionary historical approach where the best available information was used to determine the number of animals that could be supported by the available forage (coe, cumming & phillipson 1976; fritz & duncan 1994; mentis & duke 1976). the precautionary approach focused on animal numbers rather than impact. this approach may, however, still be more useful in smaller, more homogenous private protected areas and conservation areas until our understanding of how to measure herbivore impact and its outcomes has improved. however, managers of conservation areas and the advising scientists in smaller conservation areas have to take into account that the ecosystem outcomes cannot be predicted by the impact of animal numbers alone (peel, kruger & zacharias 2005). the main factors that interact to deliver various potential outcomes are driven by the values of the community towards a specific desired outcome or state such as a representative herbivore population. the values may include economic goals, goodwill of adjacent stakeholders and/or biodiversity (structure, function and composition at the various scales as defined by noss [1990]). these values interact with the abiotic template to render a specific community of plants and animals over different temporal and spatial scales (figure 1). figure 1: a framework illustrating the top-down influences of the herbivore objective as an outcome of the desired state. the herbivore population is influenced by bottom-up processes that effect these populations at different spatial and temporal scales. this determines the options that management has to affect the shorter-term herbivore population outcomes (adapted from peel, m.j.s., 2005, ‘towards a predictive understanding of savanna vegetation dynamics in the eastern lowveld of south africa: with implications for effective management’, phd thesis, university of kwazulu-natal). most national parks define their desired state with regard to biodiversity as represented by the plant and animal species and the ecosystem processes that used to occur in the region, but this objective is influenced by the economic expectations of the stakeholders. the diversity that can be supported by a protected area is determined by the geomorphological template and climate, which determines the large-scale processes and patterns over time. at the landscape scale and over the medium term, rainfall and soil properties (texture, soil depth, nutrient concentration and moisture) are the most important drivers in african savannas (peel et al. 2005; venter, scholes & eckhardt 2003). this forms the template that determines the moisture and nutrients available to plants, which, in turn, influence the habitats utilised by herbivore populations. herbivores respond indirectly to these drivers and in african savannas 87% of the variance in large ungulate biomass is explained by soil properties (fritz & duncan 1994). management of herbivores focuses on shorter-term outcomes and tend to be at smaller spatial scales. various management factors, such as fences, water distribution and fire, determine these spatial scales. most stakeholders require protected areas to be fenced to limit transgression by damage-causing animals and to retain ownership of ‘their’ animals. these fences determine the scale of the area to be managed as well as the type and size of herbivores that can be supported by the area (du toit & owen-smith 1989; peel et al. 1999; peel et al. 2005). in areas constrained by fences, animals usually have to be supplied with additional water, which not only ensures animal health but also provides tourists with a better wildlife experience by concentrating animals in an area. fire is also an important driver of vegetation because fire intensity and frequency affect vegetation structure (van wilgen, govender & biggs 2007; van wilgen, govender & macfadyan 2008). furthermore, the interaction between fire and herbivory has a significant effect on woody structure (figure 2), with fire, elephant and bark-feeding vertebrates all contributing significantly to the mortality of large trees (bond & van wilgen 1996). figure 2: an illustration of the interaction between herbivores and the ecological influences of soil moisture and nutrient distribution at the medium or landscape scale. at smaller spatial and temporal scales, soil moisture and nutrients interact with other factors such as water distribution and predators to affect herbivore population and distribution. artificial water is mostly provided to enable animals to forage at a similar intensity over the entire landscape. depending on the density of animals present, such wide water distribution may lead to homogenisation of vegetation composition and structure (gaylard, owen-smith & redfern 2003; owen-smith 1996; walker et al. 1987). the provision of artificial water also influences herbivore composition in that it favours species that are attracted to these water sources, such as zebra (equus burchelli) and blue wildebeest (connochaetes taurinus). other species tend to prefer rivers and drainage lines, such as elephant (loxodonta africana) and impala (aepyceros melampus), whilst the distribution of browsers is not significantly influenced by the presence of water (smit, grant & devereux 2007). in smaller, fenced areas in particular, where activities such as tourism, game sales and hunting are an important source of income, water distribution policies generally result in increased water distribution, higher densities of water-dependent herbivores and often, in extreme cases, overutilisation of resources over a wide area (see figure 2). selective herbivory is also a major driver of vegetation structure and composition, creating diversity and heterogeneity at different scales by their preference for certain forage types and patches (adler, raff & lauenroth 2001; gordon, hester & festa-bianchet 2004). different types of herbivore can be grouped together: bulk grazers such as elephant can change the vegetation significantly from a closed savanna with tall trees to a more open savanna with smaller coppicing trees, whilst white rhino (ceratotherium simum) can change areas dominated by tuft grasses to ones dominated by lawn grasses (figure 2). these changes are detrimental to the selective grazers such as sable (hippotragus niger), which prefer stands of lightly used tall grasslands for hiding their young and as preferred forage (collinson & goodman 1982). foraging by bulk grazers also favour herbivores that prefer high-quality grazing, such as blue wildebeest, which prefer open patches of short grass. the browsing herbivores, such as kudu (tragelaphus strepsiceros), that preferentially utilise woody vegetation may be affected by the other herbivores if the available browse is decreased. small-scale heterogeneity is created and enhanced by selective grazing on patches with short, actively growing grasses such as lawn grasses (fuhlendorf & engle 2001; mcnaughton 1979; stock, bond & van de vijver 2009), areas of high forage quality such as sodic patches and termite mounds (grant & scholes 2006), or areas with high biomass or quantity of forage (turner et al. 1997), as illustrated in figure 2. larger-scale heterogeneity may be created by herbivores preferentially utilising areas around natural water sources in the absence of additional artificial sources (fuhlendorf & engle 2001). similarly, fires that burn over large areas may limit the area covered by lawn grasses by drawing animals away from these areas and thus allowing the areas to change back into areas dominated by tuft grasses (archibald 2008). however, fires that cover smaller areas can increase the heterogeneity of herbaceous vegetation created by grazers (archibald et al. 2005; staver et al. 2009) by increasing the difference in biomass between heavily grazed lawn grass with burnt patches and ungrazed, unburnt patches (fuhlendorf & engle 2001). although the expectation is that protected areas have to be large to maintain high biodiversity, evidence suggests that this type of habitat heterogeneity can counterbalance the effect of small property size (cromsigt, prins & olff 2009). the adaptive management approach (back to top) defining the desired state or outcomes of herbivore management economic objectives to be able to manage effectively, the objectives of the protected area must be clearly stated. for national parks the preservation of representative biodiversity is the highest priority, as stated in the sanparks vision. for private protected areas economic return through tourism is often the highest priority, whilst for smaller national parks, biodiversity may be less important than the preservation of a particular community or species. the ideal for both types of reserve would be high species diversity and high income. this is not always achievable, for either private protected areas or national parks, as illustrated in figure 3. in this example, protected areas were scored on the basis of their potential to accrue income through ecotourism (occupancy rates), game sales and hunting as a function of animal diversity. animal diversity values were derived from the annual game counts and subsequent calculations of stocking density based on the vegetation condition in the form of grass species composition and standing crop, specifically (peel et al. 2005). the allocated indices for private protected areas and national parks are illustrated in table 1 and table 2, respectively. figure 3: indices of potential income based on (a) product versus vegetation condition parameters and herbivore biodiversity for the private protected areas and (b) on bed nights sold per area (km2) per year versus biodiversity parameters (as shown in table 2). number next to point indicates number of properties or parks represented with this index where more than one falls into the specific category. table 1: indices for biodiversity and income derived for the private protected areas (also see figure 3a). table 2: indices used to plot income against biodiversity for all parks managed by sanparks. according to the classification of the 24 private protected areas evaluated, 52% were classified as having low vegetation potential and low income, 4% as having high vegetation potential and low income, 30.4% as having low vegetation potential and high income and only 14% in the most desired quadrant of having high vegetation potential and a high income bracket (figure 3a). in 20 national parks managed by sanparks (before the establishment of the garden route national park, where biodiversity is the main priority, 30% had low diversity and low income, 5% had high diversity and low income, 45% had low diversity and high income and 20% achieved the desired combination of high diversity and high income (figure 3b). although these results are preliminary, they illustrate how important the top-down drivers, dictated by the value systems (shown in figure 1), are in herbivore management. biodiversity objectives the biodiversity aspect of the objective of sanparks is to maintain biodiversity in all its natural facets and fluxes. heterogeneity is assumed to be the ultimate source of biodiversity and is also the key link between structure and function (grant et al. 2011; pickett, cadenasso & benning 2003). heterogeneity has therefore been used to predict (murwira 2003; august 1983) and understand (cromsigt & olff 2006) animal distribution. to achieve this ecosystem objective it is necessary to develop an integrated understanding of ecosystem diversity and dynamics and, where necessary, intervene with appropriate strategies, in order to conserve and restore heterogeneity, terrestrial biodiversity and natural processes such as herbivory. herbivory objectives with recognition of the importance of spatial and temporal variation to maintain ecosystem processes, herbivores are managed primarily as ecosystem drivers and management is considered only where the impact is considered excessive. the approach to strategic adaptive management (roux & foxcroft 2011) is used to learn about herbivore–vegetation dynamics. this approach strives to allow herbivory to bring about sufficient ecosystem change or flux whilst providing a safety net to avoid unacceptable or irreversible change to the vegetation and associated faunal communities of the ecosystem. larger protected areas are usually better able to represent a wider range of different states than smaller areas; however, some degree of change is recognised as desirable even in smaller parks. indicators of not achieving herbivore objectives vegetation changes one of the most important indicators of excessive herbivore impact is vegetation homogenisation due to overutilisation, rather than creating the heterogeneity as is expected of herbivory in natural systems (adler et al. 2001; asner et al. 2009; also see grant et al. [2011] for a discussion on evaluating heterogeneity as an indicator of the desired state). changes in the structure of vegetation (especially woody plant species) in savannas may have a significant effect on other dependent species by, for example, altering nesting and perching sites for birds, reptiles and bats (cumming et al. 1997), forage for browsers (skarpe et al. 2004; woolnough & du toit 2001) and habitat for different herbaceous species (scholes & archer 1997). furthermore, large trees (more than 5 m tall) have been shown to play an important role in ecosystem functions such as hydraulic lift (ludwig 2001), erosion control on river margins (rogers & o’keeffe 2003) and nutrient cycling (treydte et al. 2007). mixed feeders, especially elephants, are known to be an important driver of shifts in vegetation structure and composition as they forage selectively at specific height classes (guldemond & van aarde 2007; ihwagi et al. 2010; kerley & landman 2006; shannon et al. 2008). changes in vegetation structure can also be caused by giraffe (bond & loffell 2001) and porcupine (yeaton 1988), as well as by nonherbivore factors such as high winds, disease, floods (rogers & o’keeffe 2003) or drought (viljoen 1995; shackleton 1998). herbivore impact has classically been measured according to vegetation outcomes, which are mainly aimed at avoiding degradation (scholes 2009). degradation can be described as the reduction in capacity of the vegetation or soil to support life through the damage of physical, chemical or biological properties, thereby contributing to an unsustainable ecological system (vlek, le & tamene 2008). overgrazing (which is a function of time and not necessarily a function of animal number) is a concern because it can lead to a degradation of the herbaceous layer. this can be defined as the inability of forage species to maintain themselves over time due to an excess of herbivory and related processes (mysterud 2006). this is a result of a weakening of the grass layer, resulting in reduced root growth and eventually reduced rainfall infiltration (snyman 2009), increased run-off leading to soil erosion and a loss of soil nutrients (mcginty, smeins & merril 1979; van de koppel et al. 2002). the major indicator of such degradation is a loss of the generally palatable and productive perennial grasses, which further results in a decrease in herbaceous basal cover. this is further reflected as a long-term reduction in both above-ground (diaz, noy-meir & cabido 2001), and below-ground (snyman 2005) net primary production. another indicator of excessive herbivore impact is an increase in tree density as a result of their deeper rooting systems, which allow them to access deeper water resources and thus facilitate a cohort of trees to emerge when the basal cover is decreased and grass competition is removed. increases in tree density are exacerbated by a reduced frequency of high-intensity fires as a result of the weakened grass layer, which allows fire-sensitive woody saplings to establish (scholes 2003). conversely, high fire frequencies may result in the proliferation of unpalatable, fire-tolerant grasses such as bothriochloa radicans. certain woody species are also known to increase in response to high levels of herbivory independent of the change in fire history, especially in more arid areas (scholes & archer 1997; ward 2005). herbivore numbers and distribution herbivore numbers are seldom a concern in large parks where flux is required to maintain system heterogeneity; however, herbivore numbers are often used as an indicator in smaller parks and protected areas to avoid possible undesirable change (peel et al. 1999). the only instance where larger parks would be concerned about herbivore numbers is in the case of a decline of a specific group of herbivores. the decline in the selective grazers in the knp is such an example, because the decline indicates unacceptable ecosystem changes (grant et al. 2002; ogutu & owen-smith 2003; owen-smith 1985). an alternative approach in smaller parks and private protected areas is to evaluate animal numbers against the forage available to optimise for economic purposes within a sustainability paradigm. a change in herbivore distribution is an alternative indicator of herbivory, for example the change in elephant distribution following the closure of waterpoints in the knp (smit & ferreira 2010). herbivore impact may be expected to be evident first in favoured habitats, such as those with high nutritional resources like sodic sites and termite mounds (botkin, mellilo & wu 1978; grant & scholes 2006; naiman et al. 2003; wentzel, botma & van rooyen 1991). the absence of unacceptable impact in such areas would be expected to indicate that the less favoured areas would not be undesirably impacted. thresholds of potential concern an important aspect of adaptive management is setting levels of concern that would indicate that the objectives of the conservation area may not be achieved (rogers 2003). thresholds of potential concern (tpcs) can be seen as hypotheses of performance measures that predict management outcomes. scale is explicitly considered in the setting of each tpc, which is, in turn, determined by the scale at which the specific element of function, structure or composition is relevant in the ecosystem. ideally, all biophysical tpcs should relate either to the imminent danger of loss of a species or to a real transition beyond the desired set of varying conditions (as expressed by the tpcs). these changes usually happen gradually (e.g. the conversion of grassland to woody thickets), but sudden changes have been recorded in other ecosystems such as lakes (biggs, carpenter & brock 2009). thus, it is difficult to determine whether a point constitutes a real concern that should lead to management actions and whether the tpc should rather be adjusted as indicated by improved understanding. when a tpc indicates that the management did not have the predicted outcome, the exceedence is discussed with scientists and managers who then decide on appropriate management adjustments. the evaluation of the outcome of the management action is an integral aspect of this process and informs the amendment of the tpc and potential management actions. each tpc requires a threshold or benchmark against which to measure change and to guide the setting of a tpc. such information is not available for most of the indicators and, therefore, the oldest reliable historical data are used as an initial benchmark to be adjusted as knowledge of the system improves. a rate-based approach to setting and evaluating tpcs is probably the best solution at this stage (scholes & kruger 2011). the temporal framework used for tpcs attempts to ensure that future generations still have options of different outcomes. the tpcs were originally developed to try to detect homogenisation or unacceptable change in: • structural diversity in the woody and herbaceous component • patchiness in the woody and herbaceous component • dominant and subdominant tree species • basal herbaceous cover • herbivore species composition • herbivore distribution • landscape function (nutrient cycling, infiltration and soil stability, as described by ludwig et al. [2004]). rare species were evaluated separately and the suggested tpcs and monitoring are discussed by mcgeoch et al. (2011) table 3 summarises the specific concerns raised by park managers in science–management forums. the tpcs listed are all preliminary and will be re-evaluated as more knowledge is gained, if necessary. table 3: examples of management concerns and related thresholds of potential concern in national parks in south africa. monitoring change in vegetation and herbivores effective monitoring is imperative in the adaptive management process. according to duncan and wintle (2008) a monitoring programme aimed at determining whether the management objectives are achieved should: • use clearly defined performance measures (e.g. population size of forest dependent species) • adequately reflect the precision (confidence) requirements implied by the management objectives • be performed at a spatial extent and resolution congruent with the management objectives • avoid covering too many aspects • target indicators for which risk assessments can be undertaken. historically, monitoring in the national parks and protected areas has focused mainly on the herbaceous vegetation and herbivore numbers. in the knp, vegetation monitoring was primarily aimed at informing fire management (trollope 1990a), whilst in the private protected areas monitoring was aimed at determining the available forage. this information was used to determine the number of animals that could be supported by the forage resources without deterioration of the vegetation due to herbivory. this applies in particular to grazers, as grass appears to be more limiting than browse in most instances (peel et al. 1999). in the knp an extensive vegetation monitoring programme was initiated in 1989 with 533 sites across the entire park (approximately 2 million hectares), with the primary intention of guiding fire regimens and, secondly, to assess vegetation change in response to herbivory (trollope 1990b). this programme was expanded to include the woody component in 2002 (zambatis 2002). herbaceous species surveys were performed annually by rangers whilst the woody component was monitored every 3 years. additionally, fixed-point photography sites and aerial photography have been used as vegetation monitoring tools (eckhardt, van wilgen & biggs 2000). detailed vegetation maps have been developed for all the national parks and the private protected areas adjacent to the knp (bezuidenhout 1996, 2004; bezuidenhout & brown 2008; gertenbach 1983; gotze et al. 2008; peel, kruger & macfadyen 2007; peel et al. 2005; rubin 1998; rubin & palmer 1996; siebert & eckhardt 2008; van rooyen et al. 2008; van staden & bredenkamp 2006; venter 1986; venter et al. 2003). these studies are valuable to define monitoring sites and can contribute valuable baseline data to detect future trends. herbivore exclosures, erected in several parks to monitor vegetation change due to herbivory, are also valuable reference sites (asner et al. 2009; blose 2007; landman & kerley 2009; siebert & eckhardt 2008) and have played an important role in understanding elephant impacts in addo elephant national park (lombard et al. 2001). in the knp, large mammals have been counted annually from 1977 to 1994 using a fixed-wing aircraft (redfern et al. 2002). since 1998, the census has been carried out along fixed transects, thus allowing for the precision of the census to be determined (bothma et al. 1990; peel & bothma 1995). in most of the other national parks and private protected areas large mammals are counted annually. the arc-api savanna ecosystem dynamics project was set up in 1989 and covers an area of some 450 000 ha of the eastern lowveld, between the sabie and letaba rivers on private protected areas adjacent to the knp. it includes 700 vegetation sampling sites, which are monitored annually with the aim of detecting vegetation change over time. the database includes environmental (e.g. rainfall) and management (e.g. animal numbers) data as well as a faecal analysis component as an adjunct to the vegetation monitoring. an aerial animal count component includes animal type, number and distribution, with data going back to 1992. the field work of this project is carried out by trained technicians. evaluation of thresholds of potential concern vegetation in the knp one of the main concerns around vegetation change has been the loss of large trees (coetzee et al. 1979; eckhardt et al. 2000; grant et al. 2007). it is acknowledged that a rate-based tpc is the ideal approach to evaluate rate of loss of large trees and data are presently being collected to develop such a tpc. seedling survival and recruitment to the taller tree class will be included in such a tpc and are being addressed by research projects. at this stage the best available data to evaluate the tpc are those of the veld condition assessment woody survey. the data, collected in 2006, were used to determine whether the large tree threshold had been exceeded. vegetation survey data of 1957 (when the experimental burn plots were created as described by van wilgen et al. 2011) were used as the benchmark. only landscapes with burn plots (van wilgen et al. 2007) could be evaluated. the pretoriuskop landscape, situated in a high-rainfall area showed an increase in tall trees, but the decline was much more than the maximum allowed for the low impact zones in the three other major landscapes (table 4). table 4: selected landscapes indicating the threshold of potential concern for loss of large trees (more than 5 metres), the benchmark and the subsequent survey data. to inform appropriate management action the mechanism driving the loss of large trees must be determined. the distribution of elephant suggests that most impact is at a small scale and concentrated along the main perennial and seasonal rivers (smit & ferreira 2010). the monitoring was thus adjusted to include trees along these rivers in the knp (druce et al. 2008; hartmann 2009). in an attempt to mitigate these trends, management actions included an attempt to decrease the frequency and intensity of fires in the areas of concern as well as to investigate methods of decreasing elephant density and utilisation in these areas, as guided by the elephant management plan (sanparks 2010). data collected annually during surveys on private protected areas adjacent to the knp illustrate the use of rate of change as a tpc (figure 4). the proportion of trees in different height classes impacted by elephant is illustrated in figure 5 to evaluate the possible use of impact tpcs. the latter is monitored to determine whether the tree layer is tending towards domination by a single height class (homogeneity), which is problematic with regard to both the biodiversity and sustainability objectives. figure 4: the annual percentage change in trees taller than 5 metres (a) compared to an increasing trend in elephant density in one of the private protected areas and (b) compared to a variable trend in elephant density in a private protected area. figure 5: the proportion of trees of different height classes utilised by elephant. more detailed information from private protected areas can assist in evaluating elephant impact as a mechanism for change in woody composition and structure. an exploratory linear regression (figure 4a) indicates that the density of trees taller than 5 m in protected area 1 has declined markedly since 1992 corresponding to an increase in elephant densities. the decline was slow initially (r2 = 0.54 until 2000), but then became more rapid (r2 = 0.76). the overall regression coefficient was r2 = 0.67. this is similar to the trend recorded in the knp (see earlier) and indicates that elephant utilisation is an important mechanism for the loss of large trees. for protected area 2 (figure 4b) the increase in elephant numbers was slower and decreased when the elephant moved out of the area recently (r2 = 0.54). the impact is influenced by rainfall as indicated by the decline in taller tree densities in dry years (1989–1990, 1993–1994 and 1998–1999). there is some indication of an increase in the density of trees taller than 5 m in protected area 2, which may be related to a recent efflux of elephant from the area. together this may indicate that taller trees may be recruited if the elephant density is reduced. as shown in figure 5, the selection for height classes demonstrate a clear selection for trees between 2 m and 5 m tall, and those taller than 5 m. these results show that the loss of large trees can be effectively monitored using existing monitoring techniques and that tpc exceedences can be associated with elephant impact. change in species and community composition was examined as an indication of elephant impact in the marakele national park (mnp). during 1996, 15 monitoring plots were subjectively placed in the major vegetation communities in the mnp. at that stage the park was dominated by a mountainous area with a very small area of footslopes and plains and the recommendation was to introduce 15 elephants. in 1996, however, 39 elephants were re-introduced. the point-centred quadrate method of coetzee and gertenbach (1977) was adapted to monitor the impact of elephants on the vegetation using randomly distributed plots in four of the major plant communities identified as possible suitable elephant habitat in the mnp (van staden & bredenkamp 2006). monitoring was initiated in march 1996 before the elephants were introduced and repeated during the next 2 years, with a subsequent follow-up in october 2008. a higher diversity of plant species was recorded in the different plant communities owing to the opening of gaps in the plant communities, but there was a negative impact on the microhabitats of the thicket and forest, which exposed rare and endangered species. herbivore threshold of potential concerns several herbivore tpcs have been developed for the knp. however, the only tpcs that were addressed were the declines in and retraction of the distribution of the lower density antelope and the increase in elephant numbers. the tpc developed for detecting unacceptable decreasing trends in the low density antelope resulted in the creation of an additional breeding enclosure (grant & van der walt 2000). this tpc was later supplemented with nutritional status tpc, meant to flag unacceptable low nitrogen and phosphorus concentrations in the faeces of roan antelope (hippotragus equinus). these tpcs were replaced by the tpcs for species of concern as addressed by rebelo et al. (2011). the effect of the increase in elephant is measured using the impact tpcs as discussed above. changes in herbivore population in smaller national parks and private reserves, there are often concerns about declining herbivore populations. based on the data collected from the private protected areas, figure 6 shows a monotonic decline of blue wildebeest, which exceeds the guideline tpc that was set at three monotonic drops of 10% (also see table 3). figure 6: decline in the blue wildebeest population in a private protected area between 1997 and 2009 (figures indicate percentage decline per year). alternative approaches to monitoring? (back to top) site selection acknowledgement of the importance of evaluating a loss of heterogeneity at a large scale as an indication of system change, allows changes at smaller scales to be addressed using a step-wise approach (figure 7). figure 7: a framework for structuring a monitoring programme for detecting vegetation change and feedback mechanisms; thresholds set at 60% are illustrative only. in the model shown in figure 7, monitoring is focused on areas or habitats where a particular concern is most likely to be exhibited by, for example, nutrient hotspots for grazers or large trees for elephants. unacceptable change in a predetermined sample of selected sensitive and key habitats should then trigger a more detailed and expanded monitoring programme in the area of concern to confirm the tpc exceedence. the thresholds set at 60% are only illustrative and should be determined by experts if sufficient data are not available to determine the tpc. such a tpc should reflect not only unacceptable changes due to excessive impact but also a lack of impact that would lead to a loss of, for example, high-nutrient areas such as grazing lawns, which would decrease nutrient availability and habitats used to avoid predation. these areas could be selected for focused monitoring as most grazer populations rely on relatively small areas of key resources (grant & scholes 2003; hanley 1982; mcnaughton et al. 1989; stock et al. 2009). examples of such nutrient hotspots for grazers are sodic sites, termite mounds, calcrete hard pans and grazing lawns (grant & scholes 2006; mutanga et al. 2004; naiman et al. 2003; schmitz 2008; stock et al. 2009; traill 2004; treydte et al. 2008), whilst browsers tend to concentrate in riverine areas and within specific vegetation types. elephants also prefer trees of a certain height class (shannon et al. 2008) and more focused monitoring of this class may detect unacceptable trends in the loss of large trees. alternative thresholds of potential concern we contend that there will be almost no risk of unacceptable herbivore impact as long as these intensely utilised patches are not overutilised (gordon et al. 2004; van de koppel et al. 2002). these areas could therefore be used as indicators of herbivore impact (muller 2005). if plants in these areas are grazed too frequently, composition may change with a concomitant reduction in grass tuft size and basal cover, resulting in an increase in the area of bare soil (senft et al. 1987). this is more likely to happen in arid and semiarid environments (mysterud 2006) and a bare ground index may be a useful tpc for intensely utilised grazing areas in areas with an annual rainfall of less than 650 mm (sankaran, ratnam & hanan 2004). in more mesic systems (rainfall > 650 mm), overgrazing may lead to replacement of palatable species by unpalatable species, and the abundance of the latter as well as the ratios of annual to perennial grasses may therefore be more useful tpcs. species composition and biomass may also be an indicator of a decline in grazer numbers, which could result in a change in the herbaceous structure, with lawn grass areas giving way to tuft grass areas (archibald 2008). animal condition and nutritional status in smaller conservation areas there is often a concern about the loss of herbivores due to a lack of forage resources. forage resources could thus reflect a deleterious change in vegetation and should be considered when management decisions are taken. a decline in the condition of herbivores due to declines in the amount of preferred forage and foraging areas (cromsigt et al. 2009) may thus be an indication of insufficient resources. this is reflected as a decline in phosphorus and nitrogen concentrations in faeces (dörgeloh, van hoven & rethman 1998; grant et al. 2000). nitrogen and phosphorus concentrations in dry faeces below the threshold values of 1.2% and 0.25%, respectively, would constitute a tpc exceedence (grant et al. 2000; wrench, meissner & grant 1997). such a tpc may also be useful for management of rare species, which require more intensive management, and if there are concerns about the loss of such species. for selected species, a tpc that indicates a change in distribution may be valuable. an example is the change in elephant distribution away from rivers in the knp (young, ferreira & van aarde 2009b), which is probably associated with an increase in elephant density. at larger scales a change in the system may be reflected by the existing range contraction tpc, as was exhibited by the roan antelope in the knp (harrington et al. 1999). energy flows and sustainability in smaller parks and private protected areas managers prefer to use a precautionary approach to avoid overutilisation and degradation based on carrying capacity. coe et al. (1976) use energy flows based on rainfall and vegetation biomass as a tpc, rather than population numbers according to a predetermined carrying capacity. another approach to this is to use requirements for endosomatic energy (food and work) as a tpc (peel 2005). in this case the average energy demands of the different species are based on an estimate of the activity patterns, which affect the feeding requirements of the various species. available energy is determined by subtracting the energy fraction that is reduced by internal overheads (e.g. consumption used to maintain the population) and external overheads (e.g. predation that reduces the population) from the useful energy flows available from the forage (peel 2005). environmental loading (the biophysical cost of the diet) is used as an indicator of the forage energy that is available to the animals. it is calculated using the metabolisable energy of the forage (10.5 mj/kg dry matter). estimates vary from 22% to 49% in the broad-leaved savannas to between 15% and 80% in fine-leaved savannas (in highly nutritious systems). in this case, the tpc would be a conservative estimate of the forage energy available that would avoid the loss of grazing animals and a concomitant degradation of the grazing resource. this more precautionary approach may be suitable in smaller national parks and wildlife reserves where fluctuations have to be limited because of the scale and limited habitat heterogeneity of the area. figure 8 illustrates the process taken from a private protected area adjacent to the knp where the resource is not limiting for the period in question. figure 8: the capacity of the grazing resource, presented as the amount of grass required versus that available to provide sufficient energy to a game population of known number. economic return managers of conservation areas tend to avoid issues surrounding economics, but economic return is an integral part of herbivore management. this suggests that a tpc is required that would define the desired position of the enterprise relating to biodiversity and economy (figure 3). discussion (back to top) rangelands have long been managed using animal numbers as an indicator of degradation related to excess herbivory (fynn & o’connor 2000; o’reagain & turner 1992; scarnecchia 1988). using vegetation change as an indicator of the impact of herbivory is a well-explored field and has also been used to determine the impact of wild herbivores (trollope 1990a; peel et al. 2005). however, by acknowledging the importance of flux and heterogeneity in complex systems, the management of large conservation areas such as the knp and the kgalagadi national park have moved to using animal impact and other indicators (e.g. a measure of heterogeneity) to inform management rather than animal numbers (grant et al. 2011). private conservation areas achieve their desired state by ensuring sufficient income from their enterprises by avoiding loss of animals due to a lack of forage resources; in other words a sustainability paradigm. the concern about herbivore management is thus focused on impacts and responses rather than numbers. to measure impacts reliably we need to have some predictive understanding of the response of different herbivores to changes in climate or soil (template) and the related vegetation as well as the response of the vegetation to herbivory. predators, disease and water provision all impact on the way that herbivores utilise the landscape and these factors need to be considered if we are to predict and understand herbivore impacts on their environment (young, ferreira & van aarde 2009a). the tpcs developed for the knp attempted to address the outcomes of this complex system according to vegetation composition and structure, and herbivore populations and distribution. we have evaluated some of the tpcs to determine how useful they would be in practice and although some need to be adjusted, they have proved useful for flagging concerns. the importance of understanding the mechanisms that lead to the tpc exceedence was emphasised and should be considered when designing monitoring programmes. the suggested use of energy flows to determine appropriate animal numbers in smaller protected areas shows great promise. using this method, animal type and number can be manipulated in response to vegetation biomass and habitat change. this approach requires sophisticated data and knowledge, but will allow more accurate estimations of the number of animals that can be sustained by the system. using such an approach will allow more natural flux than the fixed ‘carrying capacity’ approach that has been used in the past. to be able to reflect changes that may be of concern, monitoring must address specific questions for each component at each relevant scale. this is a daunting task for conservation agencies with limited resources and a much more focused approach is thus recommended. to prioritise monitoring efforts it is necessary to identify the main drivers of vegetation change as well as the types of change that may be expected. table 5 is an attempt to list the most important drivers in the savanna parks and the vegetation changes that could be expected as an outcome of the effects of each of these drivers. table 5: summary of the drivers that may be responsible for exceedences of thresholds of potential concern. a tick indicates that the driver may influence this vegetation response and should therefore be associated with monitoring for the specific driver. where resources for monitoring are limited, we suggest that the most sensitive indicators of the expected change be the focus of the monitoring programme. the selected sites should be those that are most likely to be vulnerable to change. once a concerning change at the selected sites is detected, surveys should be expanded to more sites to confirm the trend. knowledge of the mechanisms that lead to these concerning changes is important in informing possible management action. exclosures that are managed differently from the surrounding areas could act as reference points in large protected areas. this more focused approach needs to be tested and evaluated to enable further learning to promote efficient monitoring and management. data collected on the private protected areas adjacent to the knp, where economic goals are paramount, are analysed as soon as surveys are completed, so that timely management can be implemented (peel et al. 1999). in the knp, where biodiversity is the main objective, analysis of previous surveys proved to be very complex. one of the major hurdles in evaluating tpc exceedences in the knp was an under-estimation of the data processing requirements for the evaluation. the problem is currently being addressed through the development of an automated analysis procedure (kruger et al. 2008), which should improve the rate and interpretation of monitoring results significantly. a critical consideration in ensuring the success of the use of tpcs in flagging management concerns is the implementation of the feedback process. each tpc has to be evaluated according to a defined time schedule and the outcome of the evaluation needs to be reported at a predetermined forum where decisions about the management response to tpc exceedences can be made. in the past, this process has proved successful in private protected areas where an agreement between a client and a service provider exists and where regular feedback (at least annually) forms part of a contractual agreement. the whole tpc process depends on this crucial last step; if the feedback mechanisms fail, the tpc process fails. conclusion (back to top) vast resources may be required to determine unacceptable (with regard to the defined desired state) impact on vegetation due to herbivores or other drivers. a framework was developed to guide resource use by identifying areas of concern, which would allow focused monitoring efforts. in cases where knowledge of the system is particularly limited, reference sites that represent the most important ecological features can be considered in addition to the focused monitoring sites to gain more knowledge of the system’s responses. whatever monitoring approach is used, it is essential to analyse and interpret findings promptly to ensure that the appropriate feedback completes the adaptive management loop in time to enhance the necessary management actions. the combination of a focused and a prompt response makes the monitoring initiative both feasible and useful, and hence likely to be used. references (back to top) adler, p.b., raff, d.a. & lauenroth, w.k., 2001, ‘the effect of grazing on the spatial heterogeneity of vegetation’, oecologia 128, 456−479. archibald, s., 2008, ‘african grazing lawns: how fire, rainfall, and grazer numbers interact to affect grass community states‘, journal of wildlife management 72, 429−501. archibald, s., bond, w.j., stock, w.d. & fairbanks, d.h.k., 2005, ‘shaping the landscape: fire–grazer interactions in an african savanna’, ecological applications 15, 96−109. asner, g.p., levick, s., kennedy-bowdoin, t., knapp, a.k., emerson, r., jacobson, j., et al., 2009, ‘large-scale impacts of herbivores on the structural diversity of african savannas’, proceedings of the national academy of sciences of the united states of america 106(12), 4947−4952. august, p.v., 1983, ‘the role of habitat complexity and heterogeneity in structuring tropical mammal communities’, ecology 64, 1495−1507. bezuidenhout, h. & brown, l.r., 2008, ‘vegetation description of the doornhoek section of the mountain zebra national park (mznp), south africa’, koedoe 50, 82−92. bezuidenhout, h., 1996, ‘the major vegetation communities of the augrabies falls national park, northern cape. 1. the southern section’, koedoe 39, 7−24. bezuidenhout, h., 2004, ‘report on the impact of elephants on the vegetation of the zwarthoek section, marakele national park’, unpublished report, arid ecosystems research unit, conservation services, kimberley. biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring and management in practise,’ in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59−80, island press, washington dc. biggs, r., carpenter, s. & brock, w.a., 2009, ‘turning back from the brink: detecting an impending regime shift in time to avert it’, proceedings of the national academy of sciences of the united states of america 106, 826−831. blose, a., 2007, ‘influence of large mammals on woody vegetation heterogeneity along the sabie and letaba rivers in kruger national park’, bsc(hons) dissertation, school of animal, plant and environmental sciences, university of the witwatersrand. bond, w.j. & loffell, d., 2001, ‘introduction of giraffe changes acacia distribution in a south african savanna’, african journal of ecology 39, 286−294. bond, w.j. & van wilgen, b.w., 1996, fire and plants, chapman & hall, london. bothma, j. du. p., peel, m.j.s., pettit, s. & grossman, d., 1990, ‘evaluating the accuracy of some commonly used game-counting methods’, south african journal of wildlife management 20(1), 26−32. botkin, d.b., melillo, j.m. & wu, l.s.y., 1978, ‘how ecosystem processes are linked to large mammal population dynamics,’ in c.w. fowler & t.d. smith (eds.), dynamics of large mammal populations, pp. 373−414, john wiley & sons. coe, m.j., cumming, d.h. & phillipson, j., 1976, ‘biomass and production of large african herbivores in relation to rainfall and primary production’, oecologia 22, 341−354. coetzee, b.j. & gertenbach, w.p.d., 1977, ‘technique for describing woody vegetation composition and structure in inventory type classification, ordination and animal habitat surveys’, koedoe 20(1), 67−75. coetzee, b.j., engelbrecht, a.h., joubert, s.c.j. & retief, p.f., 1979, ‘elephant impact on sclerocarya caffra trees in acacia nigrescens tropical plains thornveld of the kruger national park’, koedoe 22, 39−60. collinson, r.f.h. & goodman, p.s., 1982, ‘an assessment of range condition and large herbivore carrying capacity of the pilanesberg game reserve, with guidelines and recommendations for management’, inkwe 1, 1−54. cromsigt, j.p.g.m. & olff, h., 2006, ‘resource partitioning among savanna grazers mediated by local heterogeneity: an experimental approach’, ecology 87(6), 1532−1541. cromsigt, j.p.g.m., prins, h.h.t. & olff, h., 2009, ‘habitat heterogeneity as a driver of ungulate diversity and distribution patterns: interaction of body mass and digestive strategy’, diversity and distributions 15, 513−515. cumming, d.h.m., fenton, m.b., rautenbach, i.l., taylor, r.d., cumming, g.s., cumming, m.s., et al., 1997, ‘elephants, woodlands and biodiversity in southern africa’, south african journal of science 93, 231−236. diaz, s., noy-meir, i. & cabido, m., 2001, ‘can grazing response of herbaceous plants be predicted from simple vegetative traits?’, journal of applied ecology 38, 497−508. dörgeloh, w.g., van hoven, w. & rethman, n.f.g., 1998, ‘faecal analysis as an indicator of the nutritional status of the diet of roan antelope in south africa’, south african journal of wildlife research 28, 16−23. druce, d.j., shannon, g., page, b., grant, r. & slotow, r., 2008, ’ecological thresholds in the savanna landscape: developing a protocol for monitoring the change in composition and utilisation of large trees’, plos one 3(12): e3979. doi:10.1371 du toit, j.t. & owen-smith, n., 1989, ‘body size, population metabolism and habitat specialization among large african herbivores’, the american naturalist 133(5), 736−740. duncan, d.h. & wintle, b.a., 2008, ‘towards adaptive management of native vegetation in regional landscapes’, in landscape analysis and visualisation. lecture notes in geoinformation and cartography, part 2, pp. 159−182, springer, berlin. eckhardt, h.c., van wilgen, b.w. & biggs, h.c., 2000, ‘trends in woody vegetation cover in the kruger national park, south africa, between 1940 and 1998.’, african journal of ecology 38, 108−115. fritz, h. & duncan, p., 1994, ‘on the carrying capacity for large ungulates of african savanna ecosystems’, proceedings of the royal society of london b 256, 77−82. fuhlendorf, s.d. & engle, d.m., 2001, ‘restoring heterogeneity on rangelands: ecosystem management based on evolutionary grazing patterns’, bioscience 51, 625−632. fynn, r.w.s. & o’connor, t.g., 2000, ‘effect of stocking rate and rainfall on rangeland dynamics and cattle performance in a semi-arid savanna, south africa’, journal of applied ecology 37, 491−507. gaylard, a., owen-smith, n. & redfern, j.v., 2003, ‘surface water availability: implication for heterogeneity and ecosystem processes,’ in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 171−188, island press, washington dc. gertenbach, w., 1983, ‘landscapes of the kruger national park’, koedoe 26, 9−121. gordon, i.j., hester, a.j. & festa-bianchet, m., 2004, ‘the management of wild large herbivores to meet economic, conservation and environmental objectives’, journal of applied ecology 41, 1021−1031. gotze, a.r., cilliers, s.s., bezuidenhout, h. & kellner, k., 2008, ‘analysis of the vegetation of the sandstone ridges (ib land type) of the north-eastern parts of the mapungubwe national park, limpopo province, south africa’, koedoe 50, 72−81. grant, c.c. & scholes, 2006, ‘the importance of nutrient hot-spots in the conservation and management of large wild mammalian herbivores in semi-arid savannas’, biological conservation 130, 426−437. grant, c.c. & scholes, m.c., 2003, ‘determining the scales at which nutrients act as drivers of herbivore distribution in the kruger national park’, african journal of range and forage science 20, 74. grant, c.c. & van der walt, j.l., 2000, ‘towards an adaptive management approach for the conservation of rare antelope in the kruger national park: outcome of a workshop held in may 2000’, koedoe 43, 103−112. grant, c.c., balfour, d., hofmeyr, m. & kruger, l., 2007, ‘a framework for understanding elephant effects on woodland’, in j.j. bester, a.h.w. seydack, t. vorster, i.j. van der merwe & s. dzivhani (eds.), multiple use management of natural forests and woodlands: policy refinement and progress. natural forests and savanna woodlands symposium iv, port elizabeth, south africa, may 15–18, 2006, pp. 163−174. grant, c.c., davidson, t., funston, p.j. & pienaar, d.j., 2002, ‘challenges faced in the conservation of rare antelope: a case study on the northern basalt plains of the kruger national park’, koedoe 45, 45−66. grant, c.c., peel, m.j.s., zambatis, n. & van ryssen, j.b.j., 2000, ‘nitrogen and phosphorus concentration in faeces: an indicator of range quality as a practical adjunct to existing evaluation methods’, african journal of range and forage science 17, 81−92. grant, c.c., smit, i.p.j., novellie, p., kruger, l., vickers, k. & gaylard, a., in review, ‘heterogeneity: what is sufficient and how can it be measured?’, koedoe. guldemond, r. & van aarde, r., 2007, ‘the impact of elephants on plants and their community variables in south africa’s maputaland’, african journal of ecology, 45, 327−335. hanley, t.a., 1982, ‘the nutritional basis for food selection by ungulates’, journal of range management 35, 146−151. harrington, r., biggs, h.c., owen-smith, n. & viljoen, p.c., 1999, ‘the roan antelope decline in northern kruger national park: spatial analysis of herbivore and vegetation changes using gis.’, south african journal of wildlife research 96, 69−78. hartmann, i., 2009, ‘can elephant impact influence the diversity of riparian tree species?’, diploma thesis, department of vegetation ecology, technische universitaet muenchen freising. ihwagi, f.w., vollrath, f., chira, r.m., douglas-hamilton, i. & kironchi, g., 2010, ‘the impact of elephants, loxodonta africana, on woody vegetation through selective debarking in samburu and buffalo springs national reserves, kenya’, african journal of ecology 48, 87−95. kerley, g.i.h. & landman, m., 2006, ‘the impact of elephant on biodiversity in the eastern cape subtropical thickets’, south african journal of science 102, 1−8. kruger, j.m., jones, m., schildauer, m. & stevens, n., 2008, ‘new technologies for automating the management and analysis of conservation data’, ecological circuits 1, 5−29. landman, m. & kerley, g.i.h., 2009, ‘elephant impacts and potential indicators of elephant-induced change to vegetation composition and structure in addo elephant national park: baseline assessment of key-plant species in colchester, kuzuko contractual park and the darlington monitoring exclosures’, report by the centre for african conservation ecology, nelson mandela metropolitan university, port elizabeth. lombard, a.t., johnson, c.f., cowling, r.m. & pressey, r.l., 2001, ‘protecting plants from elephants: botanical reserve scenarios within the addo elephant national park, south africa’, biological conservation 102, 191−203. ludwig, f., 2001, ‘tree–grass interactions on an east african savanna: the effects of competition, facilitation and hydraulic lift’, phd thesis published in tropical resource management papers, no 39, wageningen university and research centre, wageningen. ludwig, j.f., tongway, d., bastin, g.n. & james, c.d., 2004, ‘monitoring ecological indicators of rangeland functional integrity and their relation to biodiversity at local to regional scales’, austral ecology 29, 108−120. mcgeoch, m.a., sithole, h., samways, m.j., simaika, j.p., pryke, j.s., picker, m., et al., 2011, ‘monitoring terrestrial and freshwater invertebrates in protected areas’, koedoe 53(2), art. #1000, 13 pages. doi:10.4102/koedoe.v53i2.1000 mcginty, w.a., smeins, f.e. & merril, l.b., 1979, ‘influence of soil, vegetation, and grazing management on infiltration rate and sediment production of edwards plateau rangeland’, journal of range management 32, 33−37. mcnaughton, s.j., 1979, ‘grazing as an optimization process: grass–ungulate relationships in the serengeti’, the american naturalist 113, 601−703. mcnaughton, s.j., oesterheld, m., frank, d.a. & williams, k.j., 1989, ‘ecosystem-level pattern of primary productivity and herbivory in terestrial habitats’, nature 341, 142−144. doi:10.1038/341142a0, pmid:2779651 mentis, m.t. & duke, r.r., 1976, ‘carrying capacities of natural veld in natal for large wild herbivores’, south african journal of wildlife research 6, 65−74. mills, m.g.l., biggs, h.c. & whyte, i.j., 1995, ‘the relationship between rainfall, lion predation and population trends in african herbivores’, wildlife research 22, 75−88. doi:10.1071/wr9950075 muller, f., 2005, ‘ecosystem indicators for the integrated management of landscape health and integrity’, in s.e. jorgensen, r. costanza & f.l. xu (eds.), handbook of ecological indicators for assessment of ecosystem health, pp. 277−303, taylor & francis, london. doi:10.1201/9780203490181.ch12 murwira, a., 2003, ‘scale matters! a new approach to quantify spatial heterogeneity for predicting the distribution of wildlife’, phd thesis, wageningen university. mutanga, o., prins, h.h.t., skidmore, a.k., van wieren, s., huizing, h., grant, c.c.r., et al., 2004, ‘explaining grass-nutrient patterns in a savanna rangeland of southern africa’, journal of biogeography 31, 819−829. doi:10.1111/j.1365-2699.2004.01072.x mysterud, a., 2006, ‘the concept of overgrazing and its role in management of large herbivores’, wildlife biology 12, 129−141. doi:10.2981/0909-6396(2006)12[129:tcooai]2.0.co;2 naiman, r.j., braack, l.e.o., grant, c.c., kemp, a.c., du toit, j.t. & venter, f.j., 2003, ‘interactions between species and ecosystem characteristics,’ in j.t. du toit, k.h. rogers & h.c. biggs (eds.),the kruger experience: ecology and management of savanna heterogeneity, pp. 221−241, island press, washington dc. noss, r.f., 1990, ‘indicators for monitoring biodiversity: a hierarchical approach’, conservation biology 44, 355–364. ogutu, j. & owen-smith, n., 2003, ‘enso, rainfall and temperature influences on extreme population declines among african savanna ungulates’, ecology letters 6, 412−419. doi:10.1046/j.1461-0248.2003.00447.x o’reagain, p.j. & turner, j.r., 1992, ‘an evaluation of the empirical basis for grazing management recommendations for rangeland in southern africa’, journal of the grassland society of southern africa 9(1), 38−49. owen-smith, n., 1985, ‘the ecology of browsing by african wild ungulates,’ in j.c. tothill & j.j. mott (eds.), international savanna symposium: ecology and management of the world’s savannas, brisbane, australia, may 28–31, 1984, pp. 345−349. owen-smith, n., 1996, ‘ecological guidelines for waterpoints in extensive protected areas’, south african journal of wildlife research 26, 107−112. peel, m., stalmans, m. & anderson, j., 2009, ‘management plan: associated private nature reserves’, unpublished land owner document. peel, m.j.s., 2005, ‘towards a predictive understanding of savanna vegetation dynamics in the eastern lowveld of south africa: with implications for effective management’, phd thesis, university of kwazulu-natal. peel, m.j.s. & bothma, j. du p., 1995, ‘comparison of the accuracy of four methods commonly used to count impala’, south african journal of wildlife research 25(22), 41−43. peel, m.j.s., biggs, h.c. & zacharias, p.j.k., 1999, ‘the evolving use of indices currently based on animal number and type in semi-arid heterogeneous landscapes and complex systems’, african journal of range and forage science 15, 117−127. doi:10.1080/10220119.1998.9647953 peel, m.j.s., kruger, j.m. & macfadyen, s., 2007, ‘woody vegetation changes of a mosaic of protected areas adjacent to the kruger national park, south africa’, journal of vegetation science 18, 807−814. doi:10.1658/1100-9233(2007)18[807:wvoamo]2.0.co;2, doi:10.1111/j.1654-1103.2007.tb02597.x peel, m.j.s., kruger, j.m. & zacharias, p.j.k., 2005, ‘environmental and management determinants of vegetation state on the eastern lowveld of south africa’, african journal of ecology 43, 352−361. doi:10.1111/j.1365-2028.2005.00590.x pickett, s.t.a., cadenasso, m.l. & benning, t.l., 2003, ‘biotic and abiotic variability as key determinants of savanna heterogeneity at multiple spatiotemporal scales,’ in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 22−40, island press, washington dc. rebelo, t.g., freitag-ronaldson, s., cheney, c. & mcgeoch, m., 2011, ‘prioritising species of special concern in the table mountain national park for monitoring: the challenge of a species-rich, threatened ecosystem’, koedoe 53(2), art. #1019, 14 pages. doi:10.4102/koedoe.v53i2.1019 redfern, j.v., viljoen, p.c., kruger, j.m. & getz, w.m., 2002, ‘biases in estimating population size from an aerial census: a case study in the kruger national park, south africa’, south african journal of science 98, 455−461. rogers, k.h., 2003, ‘adopting a heterogeneity paradigm: implications for management of protecteed areas,’ in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 41−58, island press, washington dc. rogers, k.h. & o’keeffe, j., 2003, ‘river heterogeneity: ecosystem structure, function and management’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 189−218, island press, washington dc. roux, d. & foxcroft, l.c., 2011, ‘the development and application of strategic adaptive management within south african national parks’ areas’, koedoe 53(2), art. #1049, 5 pages. doi:10.4102/koedoe.v53i2.1049 rubin, f., 1998, ‘the physical environment and major plant communities of the tankwakaroo national park’, koedoe 41, 61−94. rubin, f. & palmer, a.r., 1996, ‘the physical environment and major plant communities of the karoo national park, south africa’, koedoe 39, 25−52. sankaran, m., ratnam, j. & hanan, n., 2004, ‘tree–grass coexistence in savannas revisited: insights from an examination of assumptions and mechanisms invoked in existing models’, ecology letters 7, 480−490. doi:10.1111/j.1461-0248.2004.00596.x sanparks. see south african national parks. scarnecchia, d.l., 1988, ‘grazing, stocking, and production efficiencies in grazing research’, journal of range management 41, 279−281. doi:10.2307/3899378 schmitz, o.j., 2008, ‘herbivory from individuals to ecosystems’, annual review of ecology, evolution and systematics 39, 133−152. doi:10.1146/annurev.ecolsys.39.110707.173418 scholes, r.j., 2003, ‘convex relationships in ecosystems containing mixtures of trees and grass’, environmental and resource economics 26, 559−574. doi:10.1023/b:eare.0000007349.67564.b3 scholes, r.j., 2009, ‘syndromes of dryland degradation in southern africa’, african journal of range and forage science 26(3), 113−125. doi:10.2989/ajrf.2009.26.3.2.947 scholes, r.j. & archer, s.r., 1997, ‘tree-grass interactions in savannas’, annual review of ecology and systematics 28, 517−544. doi:10.1146/annurev.ecolsys.28.1.517 scholes, r.j. & kruger, j.m., 2011, ‘a framework for deriving and triggering thresholds for management intervention in uncertain, varying and time-lagged systems’, koedoe 53(2), art. #987, 8 pages. doi:10.4102/koedoe.v53i2.987 senft, r.l., coughenour, m.b., baily, d.w., rittenhouse, l.r., sala, o.e. & swift, d.m., 1987, ‘large herbivore foraging and ecological hierarchies’, bioscience 37, 789−799. doi:10.2307/1310545 shackleton, c., 1998, ‘the prediction of woody plant productivity in the savanna biome, south africa’, phd thesis, university of witwatersrand. shannon, g., druce, d.j., page, b., eckhardt, h.c., grant, r. & slotow, r., 2008, ‘the utilization of large savanna trees by elephant in southern kruger national park’, journal of tropical ecology 24, 281−289. doi:10.1017/s0266467408004951 siebert, f. & eckhardt, h.c., 2008, ‘the vegetation and floristics of the nkhuhlu exclosures, kruger national park’, koedoe 50, 126−144. skarpe, c., aarrestad, p.a., andreassen, h.p., dhillion, s.s., dimakatso, t., du toit, j.t., et al., 2004, ‘the return of the giants: ecological effects of an increasing elephant population’, ambio 33, 276−282. doi:10.1639/0044-7447(2004)033[0276:trotge]2.0.co;2, doi:10.1579/0044-7447-33.6.276, pmid:15387059 smit, i.p.j. & ferreira, s.m., 2010, ‘management intervention affects river-bound spatial dynamics of elephants’, biological conservation 143, 2172–2181. smit, i.p.j., grant, c.c. & devereux, b.j., 2007, ‘do artificial waterholes influence the way herbivores use the landscape? herbivore distribution patterns around rivers and artificial surface water sources in a large african savanna park’, biological conservation 136, 85−99. doi:10.1016/j.biocon.2006.11.009 snyman, h.a., 2005, ‘rangeland degradation in a semi-arid south africa – 1: influence of seasonal root distribution, root/shoot ratios and water-use efficiency’, journal of arid environments 60, 457−481. doi:10.1016/j.jaridenv.2004.06.006 snyman, h.a., 2009, ‘root studies on grass species in a semi-arid south africa along a degradation gradient’, agriculture, ecosystems and environment 130, 100−108. doi:10.1016/j.agee.2008.12.003 south african national parks, 2006, coordinated policy framework governing park management plans (draft 3), south african national parks, pretoria. south african national parks, 2008, park management plan: kruger national park (draft), south african national parks, pretoria. south african national parks, 2010, elephant management plan: kruger national park (draft), south african national parks, pretoria. staver, c., bond, w.j., stock, w.d., van rensburg, s.j. & waldram, m., 2009, ‘browsing and fire interact to suppress tree density in an african savanna’, ecological applications 19, 1909−1919. doi:10.1890/08-1907.1, pmid:19831079 stock, w.d., bond, w.j. & van de vijver, c.a.d.m., 2009, ‘herbivore and nutrient control of lawn and bunch grass distributions in a southern african savanna’, plant ecology 206(1), 15−27. doi:10.1007/s11258-009-9621-4 traill, l.w., 2004, ‘seasonal utilization of habitat by large herbivores in semi-arid zimbabwe’, south african journal of wildlife research 34, 13−24. treydte, a.c., heitkonig, i.m.a., prins, h.h.t. & ludwig, f., 2007, ‘trees improve grass quality for herbivores in african savannas’, perspectives in plant ecology, evolution and systematics 8, 197−205. doi:10.1016/j.ppees.2007.03.001 treydte, a.c., van beeck, l., ludwig, f. & heitkonig, i.m.a., 2008, ‘improved quality of beneath-canopy grass in south african savannas: local and seasonal variation’, journal of vegetation science 19, 663−670. doi:10.3170/2008-8-18435 trollope, w.s.w., 1990a, ‘development of a technique for assessing veld condition in the kruger national park’, journal of the grassland society southern africa 7, 46−51. trollope, w.s.w., 1990b, ‘veld management with specific reference to game ranching in the grassland and savanna areas of south africa’, koedoe 33, 77−86. turner, m.g., pearson, s.m., romme, w.h. & wallace, l.l., 1997, ‘landscape heterogeneity and ungulate dynamics: what spatial scales are important?,’ in j.a. bisonette (ed.), wildlife and landscape ecology: effects of pattern and scale, pp. 331−348, springer verlag, new york. van de koppel, j., rietkerk, m., van langevelde, f., kumar, l., klausmeier, c.a., fryxell, j.m., et al., 2002, ‘spatial heterogeneity and irreversible vegetation change in semiarid grazing systems’, the american naturalist 159, 209−218. doi:10.1086/324791, pmid:18707414 van rooyen, m.w., van rooyen, n., bothma, j. du p. & van den berg, h.m., 2008, ‘landscapes in the kalahari gemsbok national park, south africa’, koedoe 50, 99−112. van staden, p.j. & bredenkamp, g.j.l., 2006, ‘a floristic analysis of forest and thicket vegetation of the marakele national park’, koedoe 49, 15−31. van wilgen, b.w., govender, n., forsyth, g.g. & kraaij, t., 2011, ‘towards adaptive fire management for biodiversity conservation: experience in south african national parks’, koedoe 53(2), art. #982, 9 pages. doi:10.4102/koedoe.v53i2.982 van wilgen, b.w., govender, n. & biggs, h.c., 2007, ‘the contribution of fire research to fire management: a critical review of a long-term experiment in the kruger national park, south africa’, international journal of wildland fire 16, 519−530. doi:10.1071/wf06115 van wilgen, b.w., govender, n. & macfadyen, s., 2008, ‘an assessment of the implementation and outcomes of recent changes to fire management in the kruger national park’, koedoe 50(1), 22−31. venter, f.j., 1986, ‘soil patterns associated with the major geological units of the kruger national park’, koedoe 29, 125−138. venter, f.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation pattern,’ in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83−129, island press, washington dc. viljoen, a.j., 1995, ‘the influence of the 1991/1992 drought on the woody vegetation of the kruger national park’, koedoe 38, 85−97. vlek, p.l.g., le, q.b. & tamene, l., 2008, land decline in land-rich africa: a creeping disaster in the making, cgiar science council secretariat, rome. walker, b.h. & goodman, p.s., 1983, ‘some implications of ecosystem properties for wildlife management,’ in n. owen-smith (ed.), management of large mammals in african conservation areas, pp. 79−80, haum, pretoria. walker, b.h., emslie, r.h., owen-smith, n. & scholes, r.j., 1987, ‘to cull or not to cull: lessons from a southern african drought’, journal of applied ecology 24, 381−401. doi:10.2307/2403882 ward, d., 2005, ‘do we understand the causes of bush encroachment in african savannas?’, african journal of range and forage science 22, 101−105. doi:10.2989/10220110509485867 wentzel, j.j., bothma, j. du p. & van rooyen, n., 1991, ‘characteristics of the herbaceous layer in preferred grazing areas of six herbivore species in the south-eastern kruger national park’, koedoe 34(1), 51–59. woolnough, a.p. & du toit, j.t., 2001, ‘vertical zonation of browse quality in tree canopies exposed to a size-structured guild of african browsing ungulates’, oecologia 129, 585−590. wrench, j.m., meissner, h.h. & grant, c.c., 1997, ‘assessing diet quality of african ungulates from faecal analysis: the effect of forage quality, intake and herbivore species’, koedoe 40(1), 125−136. yeaton, r.i., 1988, ‘porcupines, fire and the dynamics of the tree layer of the burkea africana savanna’, journal of ecology 76, 1017−1029. doi:10.2307/2260630 young, k.d., ferreira, s.m. & van aarde, r., 2009a, ‘elephant spatial use in wet and dry savannas of southern africa’, journal of zoology 278, 1−17. doi:10.1111/j.1469-7998.1984.tb06041.x, doi:10.1111/j.1469-7998.2009.00568.x young, k.d., ferreira, s.m. & van aarde, r., 2009b, ‘the influence of increasing population size and vegetation productivity on elephant distribution in the kruger national park’, austral ecology 34, 329−342. doi:10.1111/j.1442-9993.2009.01934.x zambatis, n., 2002, ‘revised procedures for veld condition assessment in the kruger national park’, internal report 1/2002, scientific services, south african national parks, skukuza. article information authors: stephen d. holness1,2 harry c. biggs3 affiliations: 1south african national parks, port elizabeth, south africa 2department of zoology, nelson mandela metropolitan university, south africa 3south african national parks, skukuza, south africa correspondence to: stephen holness email: sholness@nmmu.ac.za postal address: po box 76693, port elizabeth 6031, south africa dates: received: 07 oct. 2010 accepted: 02 feb. 2011 published: 13 may 2011 how to cite this article: holness, s.d. & biggs, h.c., 2011, ‘systematic conservation planning and adaptive management’, koedoe 53(2), art. #1029, 9 pages. doi:10.4102/koedoe.v53i2.1029 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) systematic conservation planning and adaptive management in this essay... open access • abstract • introduction    • why an adaptive approach is required in conservation planning    • similarities in philosophy       • similarity in structural and functional components       • adaptive implementation of systematic conservation planning in a messy world       • maintaining and promoting adaptive thinking in systematic conservation planning • conclusion • references • footnote abstract (back to top) this article argues that systematic conservation planning (scp) is an intrinsic part of the adaptive management approach within sanparks and should not be seen as a separate or different initiative. scp operates within a complex environment that requires a deliberately adaptive approach. the similarities in philosophy, structure and functional elements of the planning process and approach between adaptive management and scp, as applied within sanparks, are highlighted. the article distils requirements for ensuring that scp remains strategically adaptive in its approach. conservation implication: a deliberately adaptive approach to scp improves its effectiveness in guiding the implementation of conservation actions and is a requirement for effective conservation planning in a complex environment. introduction (back to top) in the last decade, conservation planning in south africa (e.g. cadman et al. 2010; cowling et al. 2003a; cowling et al. 2003b; driver et al. 2005) has been strongly influenced by the principles of systematic conservation planning (scp) (margules & pressey 2000). south african national parks (sanparks) is an important participant in this planning, which takes place at nested spatial scales in the country. it involves their entrusted conservation estate, the surrounding seascapes, landscapes and riverscapes in which the parks are embedded, and the planned future expansion of existing and new protected areas. specific theme areas where scp principles underlie sanparks’ spatial planning processes include the zoning, land use and infrastructure planning within reserves, the spatial planning for the buffer areas and areas of influence around parks, and the planning for identifying reserve expansion priorities for all existing national parks and evaluating potential areas for new national parks. scp principles and processes have been applied in a range of environments, including terrestrial, freshwater and marine areas, and underpin the response strategies of conservation agencies to key threats such as climate change and habitat fragmentation around reserves. concurrent with the scp process (which has largely focused on spatial planning issues), sanparks has actively developed a strategic adaptive management (sam) programme, which is the conceptual basis that underpins biodiversity management within reserves (rogers 2005; south african national parks 2008). adaptive management is an attempted response to the uncertainties of the operating context in an environment of complexity (levin 1999) and sam (rogers 2005), with its flexible goal-based view, should be well suited for planning in a complex situation. the importance of an adaptive approach to scp has been highlighted, at least in theory, by conservation planners (bottrill & pressey in prep.; fischer et al. 2009; grantham et al. 2010; groves 2003; knight, cowling & campbell 2006; margules & pressey 2000; pressey & bottrill 2009; salafsky et al. 2002). despite a monitoring and evaluation stage being one of the original six scp steps laid out by the influential paper of margules and pressey (2000), which also highlights the need for an iterative and adaptive approach, there appears to be little evidence that scp is actually practiced in an adaptive manner. a recent literature search1 failed to identify a single documented example of a functioning scp system that fully integrates adaptive management principles. whether the lack of examples of functioning adaptive scp systems is the result of poor documentation or an indication of scp not being generally adaptive in practice, is uncertain. in either case, it is valuable to highlight the sanparks approach to scp where it forms an intrinsic part of adaptive management. this article argues that scp is, in fact, being practiced in an increasingly adaptive manner within sanparks and that scp and sam should not be seen as being separate and unrelated processes. we firstly outline the rationale for why adaptive planning approaches are necessary within scp and then discuss similarities in the underlying philosophies of scp and sam as used by sanparks, point out similarities in structure and components between the planning processes employed, examine existing or potential parallels in application, and conclude by discussing how we may ensure that scp remains appropriately adaptive. emphasis is given to scp-related examples (the ones normally in deficit when it comes to demonstrating adaptiveness) and diagrams, but with parallel detail for sam to support the intention of the article. the article is designed to document the lessons learnt from the sanparks experience in practically applying scp in an adaptive manner within the context of a broader sam system. where possible, specific projects and outputs are referenced, but most remain unpublished or in inaccessible, grey literature. why an adaptive approach is required in conservation planning the physical, social, political, technological and economic environments in which scp takes place are changing – and, in south africa, often rapidly. process relationships in such socio-ecological systems (berkes, colding & folke 2003) are often poorly understood, unpredictable and evolving themselves. in the business domain, snowden and boone (2007) have emphasised the difference between complicated and complex systems. the former refers to systems with a large number of parts but which display predictable behaviour (e.g. an aircraft or a circuit board), whilst the latter refers to a system with any number of parts, but for which dynamic relationships and interactions dominate in a way that can produce surprises (e.g. ecosystems and most socio-ecological systems). ‘complexity’ has a variety of related technical definitions and leading complexity thinker, paul cilliers (2000), finds it useful rather to summarise the general characteristics of a complex system for user disciplines: he emphasises influential nonlinear interactions between the multiple elements of a system(which is invariably open), organised as direct and indirect feedbacks. because of the importance of these interactions, system behaviour as a whole cannot be predicted from inspection of the components alone. history of antecedent configurations is also very important in determining future trajectories. furthermore, these systems can reorganise their internal structure without the intervention of an external agent; that is, they tend to be adaptive. in the biological and ecological sciences, thinking towards complexity has received growing attention. levin (1999) used the diagnostic that systems with diversity, interactions (especially local interactions) and any selection process (such as natural selection) – precisely the kind of systems widely encountered in these fields – would show complex behaviour. resilience thinkers (e.g. gunderson & holling 2001), along with those in fields that focus on sustainability and transdisciplinarity, emphasised adaptive and often heuristic responses as appropriate for learning to cope effectively in managing complex systems, which are often depicted as socio-ecological systems. at the level of the philosophy of science, ulanowicz (2009) argues that ecology can add to (and reorganise) earlier newtonian and darwinian views with new and useful generalities. this results in society dealing more humbly, and ultimately more effectively, with many types of irreducible uncertainty in (in his cases, biophysical) systems, which nevertheless show propensities of pattern that can indeed be usefully understood. against this background it is not surprising that adaptive strategies, difficult though they are to operationalise, are being promoted widely in these fields today. if one specifically examines the environment within which scp operates and plans, there is much evidence that this environment should be seen as complex: • both biodiversity itself and the associated human interactions are changing rapidly (pressey et al. 2007). • there are many steps, often with only partly predictable outcomes, involved in the generation and ultimate implementation of scp. • scp takes place across multiple agencies (e.g. national and provincial conservation agencies, water affairs, agriculture, marine management agencies). • scp needs to engage diverse stakeholder groups (ranging from traditional communities living largely subsistence-based lifestyles on park boundaries to organised industry such as commercial agriculture or fisheries). • scp deals with multiuse landscape mosaics and river catchment networks. • scp deals with interactions between government-driven processes on the one hand (e.g. land reform) and free-market entities on the other (particularly individual land owners). • in the context of a conservation organisation, recommendations that stem from scp processes are implemented through a range of mechanisms, including land purchase, voluntary land stewardship programmes and proactive land use planning (e.g. through guiding regional zoning processes), as well as being used to respond reactively to development proposals (e.g. through environmental impact assessments). • experience indicates that there are often unexpected setbacks (e.g. large reductions in the budget available for land acquisition or changes in institutional capacity or willingness to expand) and surprising windfalls (e.g. sudden availability of high-priority conservation sites due to the death of a land owner or the auction of an insolvent estate). • there is often imperfect knowledge of many aspects(e.g. willingness of land users to be involved in a stewardship programme or to sell their land for incorporation into a reserve) and of the viability of different land uses, let alone imperfect knowledge of the multiple perceptions and mental models held by different people in this regard. if political fickleness, evolution of human values and fluctuating levels of capacity are added as additional attributes, it seems clear that the system is complex. yet, in many cases, we come from a history of viewing such systems simplistically, for example by assuming that there is a single, stable planning approach that will result in an optimal park blueprint for a region (as would be applicable in a ‘complicated’ world). as corollaries, we have often assumed that if we only had more data, we would eventually understand the whole system adequately and that a discrete handover from ’planners’ to ’implementers’ would be possible. in reality, if we accept and internalise the complexity described above along with its consequences, we need to alter our approach. on the other hand, however, we cannot allow ourselves to become paralysed by this complexity and should thus seek the requisite simplicities that allow for action and further learning (stirzaker et al. 2010). the use of scp in sanparks has led to some such adaptation and learning already, and more will follow. such learning provides the basis for the subsequent analysis in this article. whilst broad-level plans may be useful to give society a rough idea of where to conserve land or how to manage it, intermediate and finer-scale planning appear to require more adaptive capacity and response to allow successful navigation (berkes et al. 2003) of the complexity so that both conservation and broader sustainable land use goals can be reached. having discussed why an adaptive approach should be considered appropriate for scp in our (and likely most) circumstances, we now examine more specifically possible underlying and operating similarities between sam and scp, with reference to several scp examples. multiple examples of sam are described in this volume (including management of river systems, fire and alien invasive species), with an introductory editorial (roux & foxcroft 2011), sketching the context. similarities in philosophy sam and scp share some fundamentally similar intentions and much underlying philosophy. although the similarities mentioned below were presented in the original scp texts (e.g. margules & pressey 2000), certain of these issues have been made more explicit in recent renditions (bottrill & pressey in prep.; pressey & bottrill 2009), such as feedback loops being incorporated into key diagrams. scp and sam both take a wide view of the stakeholder base and strive to be inclusive. bottrill and pressey (in prep.) emphasise the importance of the early identification and involvement of stakeholders in the scp process. these stakeholders can include other conservation agencies, governments, ngos, experts and communities. within sanparks, a recent example is the planning of the marine protected area associated with addo elephant national park (on the south-eastern coast of south africa), which included stakeholders such as commercial fisheries, recreational fishermen, local property owners’ associations and subsistence fishers. stakeholder participation has included targeted meetings with specific sectors as well as public participation. another example is a feasibility study led by sanparks for a new protected area in the southern drakensberg, which has had direct representation of local authorities, agriculture, conservation organisations and government bodies such as the department of environmental affairs and the south african national biodiversity institute. whilst stakeholder mapping is less explicitly described in the headline procedures of sam, stakeholder participation is intrinsic to sam processes such as development and implementation of national park management plans. the consistent success of this sam-based initiative relies on the identification and involvement of a broad cross section of stakeholders, with particular emphasis on local government structures, interest groups, local communities and park forums. the latter acts as a particularly useful vehicle for the public to contribute to park decision making (a. symonds; pers. comm.). key benefits of the strong emphasis on stakeholder involvement include the development of a ‘shared rationality’ (rogers 2005) amongst stakeholders and increased trust and credibility amongst key partners. for example, the feasibility study in the southern drakensberg included both national and provincial conservation agencies, which have in the past been drawn into unproductive disputes over the implementation of projects that had originated separately (m. knight; pers. comm.). other benefits include: • access to implementation opportunities that would otherwise not be available (e.g. alignment with payment for ecosystem service projects or land reform initiatives) • improved understanding of the requirements and concerns of stakeholders (e.g. the marine planning process around addo elephant national park now includes the identification of areas which various fishing industries are most concerned about for the impact of the closure of certain areas) • support from a range of organisations that are responsible for implementing conservation actions (e.g. catchment management agencies, local authorities, environmental impact assessment implementing authorities). both scp and sam processes recognise the importance of societal and organisational values and attempt to elicit these values and incorporate them in the planning process. pressey and bottrill (2009) identify the establishment of clearly defined goals as a key component of a number of divergent conservation planning processes. these broad goals are driven by societal and organisational values. for example, in an scp process for a reserve, these goals may influence the relative value of biodiversity, heritage and tourism features, or the extent to which a reserve would be planned to minimise impact on economic productivity (e.g. displacement of viable resource extraction industries or land uses) or social systems (e.g. displacement of farm workers). as part of its overall sam thrust, sanparks developed a clear set of conservation values (sanparks 2008); the adaptive planning procedures in sam (rogers 2005) highlight these and other wider societal values as mandatory and key inputs in the formulation of, for example, park objectives. both scp and sam thus strive to balance a range of societal values in reaching decisions and clearly recognise that in most cases the appropriate management responses are driven not by biodiversity considerations only. both scp and sam strive for integration across sectors (e.g. conservation, agriculture, forestry, mining and water management) and see biodiversity as only one important attribute within a wider view of societal interests, usually within multiuse landscapes. it is clearly recognised that collaboration between a range of sectors are needed for successful implementation to result in the desired outcome (e.g. a resilient system within which a reserve is nested in a matrix of compatible land uses). for example, in scp for reserve expansion, the desired outcome of ensuring that freshwater systems which flow into a reserve remain in or are restored to an acceptable desired state, depends entirely on upstream users, including forestry and local authorities. this parallels what is strived for in other sam-based initiatives in sanparks, such as ongoing river management in park plans. both scp and sam accept that recognition of complexity is paramount to success. the sam process is explicitly designed to accommodate and operate in a complex environment. although some (older) scp projects were undertaken to produce once-off blueprints for reserve expansion, sanparks soon recognised that planning products had to remain ’live’ to ensure their adapting and response to complex and changing environments. for example, for addo elephant national park, a systematic reserve expansion study was commissioned from outside service providers. although a good quality product was delivered (council for scientific and industrial research 2002), it soon became clear that a single static plan for the park was not particularly useful for park expansion implementation. the base data from this project were then incorporated into an internal decision support system housed within the newly established sanparks park planning department. this allowed for response to changes in the implementation environment (e.g. specific properties may have been successfully acquired or negotiations may have fallen through for others), improvements in data (e.g. revision and refinement of land use, land transformation and degradation data), refinements of methodology (e.g. revision of targets or reprioritisation of properties based on emergent considerations) or review of value systems. a key difference may be that sam has broad overall applicability to a range of planning, management and decision making situations, whilst scp is a more focused spatial planning tool for a particular task. in this sense we could argue that scp may philosophically be seen as nested inside sam. we thus promote the idea that scp should generally fall inside the broader sam philosophy. this article discusses the extent to which this does, and should, take place, by considering similarities not only in philosophy but also in structure and process, and then reviewing whether scp implementation proceeds in a way that can be considered sam compliant in reality. similarity in structural and functional components sam, as practiced in sanparks, has a number of clearly articulated stages and processes, of which the adaptive planning process serves as a usual start (figure 1). although scp can vary widely with regard to its objectives, process, data and approach, some themes, stages and processes are common to most of its planning strategies. the iucn initiative on conservation planning convened by the species survival commission and world commission on protected areas identifies 11 major stages within an scp process (bottrill & pressey in prep.; pressey & bottrill 2009), as shown in figure 2. here we explore the similarities between sam and the scp process with regard to their structural and functional components and the overlap of various components, with specific reference to how the scp process is applied in sanparks. figure 1: flowchart of the stages of the adaptive planning component of a strategic adaptive management process (adapted from rogers, k.h., 2005, biodiversity custodianship in sanparks: a protected area management planning framework, report to south african national parks, pretoria). the initial stages of a sam process are aimed at defining the decision making environment within which adaptive management is to take place (figure 1). specific components of this process include defining a vision and explicitly identifying the operating principles and context within which management takes place. these steps implicitly require intensive internal and external stakeholder involvement, which, in practice, enhances the credibility of the process and ensures that it meets legislative requirements and is sensitive to stakeholder needs. the process of defining a vision involves a broad philosophical statement of intent that is clearly linked to the values of the organisation, whilst the operating principles refine these more specifically. the exploration of context examines the range of circumstances that affect the decision making process, including local conditions and surroundings. this leads into a series of steps where the vital attributes (or key features) of the system are identified, along with the main influencing factors and possible threats. very similar stages exist in scp and sam, both with regard to how the process is generally conceptualised (figure 2) and how it is applied within sanparks in practice. stage 1 of the scp process involves scoping and costing the planning process, which defines the way the project will be implemented. key issues include determining a planning domain and identifying the scope and scale at which planning will take place. although this is sometimes, wrongly, seen as a trivial stage, careful consideration of these issues can result in significant implementation benefits. for example, what would have been systematic assessments for buffer regions around agulhas and bontebok national parks were expanded into a systematic assessment of biodiversity priorities for the whole overberg district. this resulted in a critical biodiversity map for the district, which embeds the local park concerns into regional requirements and accesses a far broader range of implementation agents (e.g. the provincial conservation agency and local authorities) and additional resources that support more detailed planning than would otherwise have been possible. figure 2: conceptual framework for systematic conservation planning developed by the iucn initiative on conservation planning (diagram adapted from pressey, r.l. & bottrill, m., 2009, ‘approaches to landscapeand seascape-scale conservation planning: convergence, contrasts and challenges’, oryx 43(4), 464–475. doi:10.1017/s0030605309990500). stage 2 of the scp process involves identifying and involving stakeholders. as discussed earlier, these stakeholders can include other conservation agencies, ngos, local government, experts and local communities. this process is important for developing trust amongst key partners, accessing a range of implementation opportunities, developing an understanding of stakeholders’ requirements and concerns, and obtaining the support of implementing organisations (pressey & bottrill 2009). stage 3 of the scp process provides the context for the subsequent conservation planning steps. the social, economic and political setting within which conservation planning takes place is examined and direct threats (e.g. landcover change or overgrazing in particular areas) as well as indirect threats to natural features (e.g. policies that result in environmental degradation) intended for conservation are examined. thus, scp displays clear parallels with sam in the explicit consideration of the social, economic and political context within which it takes place, the deliberate involvement of stakeholders and the examination of the key features of a region and possible threats. the context examination phases of both sam and scp are followed by a process to identify objectives or goals. in scp, conservation goals are often initially defined as broad qualitative statements that link the underlying values on which conservation efforts are based to more specific quantitative biodiversity targets (pressey & bottrill 2009). these goals may include ensuring that a reserve includes a representative sample of all the habitats and species in a region, or that the ecological processes on which the reserve depends are sufficiently safeguarded (e.g. ensuring that freshwater inputs from catchment areas are of sufficient quality and quantity or that ecological corridors with neighbouring reserves are not excessively impacted). later in the scp process (step 7, after intermediate steps for gathering spatially explicit data on biodiversity and socio-economic features and threats), these broad goals are refined to more specific conservation objectives. the use of explicit and generally quantitative targets forces planners to be clear about the required outcomes (analogous to the objectives and thresholds parts of a ‘desired state’ in sam), and provides measures against which progress towards targets can be measured. these objectives reduce the potential for ad hoc decisions and form the basis for long-term monitoring and evaluation. in addition to these quantitative goals, qualitative goals are also often set. these may relate to improving the connectivity between reserves or their shape and configuration. this process strongly resembles the sam process of defining an objectives hierarchy, which starts off with fairly broad objective statements at a high level, and refines these to very specific, and often quantified, low-level objectives (roux & foxcroft 2011). at the end of this process in both sam and scp, clear, generally quantifiable objectives exist against which both the current state and the success or failure of management actions can be evaluated. in scp these objectives are commonly articulated as targets or thresholds (e.g. at least x hectares of this habitat kept in a natural or near-natural state, or incorporated into a reserve), whilst sam (for biodiversity objectives at least) tends to define the acceptable end state in terms of thresholds of potential concern (roux & foxcroft 2011), such as a water quality measure that needs to be kept within a specific range. although not identical, these concepts clearly have a very similar basis. in both sam and scp, there is concern not just when a threshold or target is exceeded, but also when the trajectory of that variable suggests that a threshold may be exceeded in the future. scp and sam then both have an operationalisation or implementation phase. in scp, stages 8–10 involve the evaluation of the current situation (e.g. a reserve network) against the objectives (e.g. quantitative targets for protection of a range of vegetation types or habitat requirements for a particular range of species), the identification of additional priority areas (e.g. for reserve expansion or for coverage by stricter land use controls), and then the implementation of these place-specific actions (e.g. through purchasing a property for inclusion into a reserve or the contractual inclusion of that land). although this is diagrammatically represented as a mostly linear system, it includes a number of iterations and feedback cycles in practice. for example, the successful purchase of a property could result in the relative value of the remaining properties needing to be re-evaluated. alternatively, properties may not be available owing to changes in willingness of the land owner to sell or socio-economic developments. once established, it would be necessary to evaluate whether the expanded reserve network did in fact maintain the biodiversity features it was designed to protect in the required state. alternatively, where the scp process was aimed at land use planning and controls, the area would need to have been evaluated regularly to ensure that the identified priority areas in a region were in fact appropriately protected and had not been lost to development. in either case, the results of this monitoring and evaluation would need to have been fed back into an ongoing planning and implementation process to ensure that the necessary adjustments were made to the identified priority areas, the implementation mechanisms used and the ongoing planning process itself. in sam, these stages involve the identification of appropriate management actions (e.g. a changed burning regime), implementing these actions and then evaluating the biodiversity outcome against the applicable thresholds of potential concern (roux & foxcroft 2011). although the scp diagram (figure 2) is summarised as a broadly linear sequence, these steps are, in reality, iterative, overlapping and interlinked. further, although adaptive cycles and feedbacks are identified at specific places in the planning process in the diagram, these in fact take place so frequently in the process that the diagram suggests a more linear process than exists in practice, at least where scp is embedded with conservation organisations. it would be helpful for our use if such iterations were made even more explicit in these diagrams. overall, there are thus strong structural and functional similarities between scp and sam, namely: • an emphasis on clearly defining the context within which they operate • explicit consideration of a range of issues such as social and economic factors (as well as biodiversity) • dependence on stakeholder involvement and full consideration of stakeholder issues • initial broad higher-level goals which are then refined to more explicit lower-level targets • explicit monitoring and evaluation stages with feedbacks and learning (in sam, and increasingly also in scp, ongoing learning is regarded as imperative, regardless of success or failure). with all these adaptive and potentially adaptive processes in scp outlined above, it would be instructive to consider the extent to which, as a whole, scp actually functions adaptively. adaptive implementation of systematic conservation planning in a messy world intuitively, it might seem most logical to conceptualise the scp process as a series of planning steps that starts with broad-scale planning and eventually cascades down to detailed fine-scale projects (figure 3). the plans would be spatially nested within one another and specific local projects (e.g. planning for infrastructure within a reserve) would reflect both the values and broad spatial priorities identified by national and regional planning. figure 3: theoretical progression of nested systematic conservation planning processes. however, in our experience, this system does not (and usually cannot) materialise for a number of reasons. firstly, knowledge is incomplete and generally insufficient for useful top-down planning. secondly, immediate requirements (e.g. a specific land parcel for reserve expansion becomes available and rapid answers are required as to whether it is strategically valuable), implementation opportunities (e.g. a short window of opportunity may be available to include biodiversity into local the spatial development framework to guide land use planning decisions for a five-year period) and short-term implementation actions (e.g. land use and zoning planning for short-duration poverty relief projects) all hinder the hypothesised ’ideal’ implementation in figure 3. thirdly, external drivers of project implementation processes (e.g. new requirements and deadlines brought about by government policy or legislative change) or requirements linked to individual projects or initiatives (e.g. a landscape initiative may require a fine-scale assessment of biodiversity priorities for a particular area to support a range of conservation implementation mechanisms, and this may be required before broad national identification of priorities has been undertaken) may require specific planning projects at particular scales for a region to be done as a priority. fourthly, the planning effort in a region is not limited to a particular organisation and the needs and resources of a number of organisations need to be accommodated and utilised. the specific planning work may also be undertaken within different organisations, with varying time availability. fifthly, limited resources preclude fully comprehensive top-down planning. finally, as an important overarching comment, the overall environment being dealt with is complex and neat, linear planning processes generally do not respond ideally to this environment at the range of operative scales. in practice, within sanparks (and probably more widely in south africa) the conservation planning process more closely resembles figure 4. figure 4: example of an actual planning process. within a region the conservation planning would be undertaken by a range of organisations (e.g. academic institutions, national institutes or consulting agencies). early projects for expansion of specific reserves are not necessarily undertaken at a broad national scale; in fact, much of the initial work tends to be undertaken for biomes or focuses at a very local scale. for example, cape for the fynbos (cowling et al. 2003a) and step for the thicket (cowling et al. 2003b) were done before any national integration had occurred. later projects, which theoretically should predate the specific expansion plans for a particular reserve, are often completed only after a range of specific reserve expansion plans have been drawn up, as, for example, in the case of the national protected areas expansion strategy (government of south africa 2010). this approach may well result in a mess of overlapping and conflicting blueprints, with no spatial or strategic cohesion. however, in practice, the approach outlined here can produce a highly responsive system that adapts quickly to implementation requirements, learns from the successes and failures of earlier processes, and incorporates both the values derived from broader integrative processes and the specific conservation priorities identified by local-scale planning. in our view, the reason for the success of this system is that the deliberate adaptive approach taken within sanparks and other organisations allows the process to tend towards robustness rather than fragmentation. there are a number of key elements that contribute to the development of a functional adaptive system. firstly, planning projects do not aim to be perfect. there is a strong prototyping approach applied in sanparks, which aims to deliver sufficiently good answers (the so-called ‘80/20 principle’, meaning that 20% effort produces 80% of the result) for specific programmes and projects. the approach is to support implementation by rapidly integrating the best reasonably available data (rather than striving for a perfect product) into efficient decision support systems. secondly, there are strong adaptive feedbacks between the various components of the system. each project contributes knowledge or techniques or helps refine values. thirdly, there is a fairly small community of scp practitioners who generally know one another, are aware of new methods developed by the others, share data, and interact reasonably frequently (national processes such as an annual biodiversity planning forum organised by the south african national biodiversity institute have significantly contributed to the maintenance of dialogue between practitioners). fourthly, limited budgets and resources have forced projects to utilise the best data and outputs from previous projects, rather than having the scope to go out and develop a whole new suite of products. lastly, the projects have all been implemented in the same conceptual environment and have strong unifying concepts, namely systematic target-driven or threshold-driven conservation planning that strives for pattern and process goals to meet in a way that is least conflicting with other land use. given that we have examined similarities between scp and sam, and how adaptively scp has been able to perform in the ’messy’ real world, we now try to extract a few essential principles that we believe will assist us in maintaining and promoting appropriate adaptiveness in scp in the future. maintaining and promoting adaptive thinking in systematic conservation planning whilst there may be occasional situations where nonresponsive behaviour is desired in scp (e.g. where an organisation may need a stable blueprint to implement a specific action for a particular period, such as in applying land use controls), this article concentrates on how appropriate levels of adaptive capacity and action can be achieved. five factors are synthesised below as key strategies. development of a prototyping approach: always insisting on comprehensiveness and full accuracy is unlikely to facilitate implementation. for example, if prototypes are not available, important windows of opportunity for delivering a product in time to influence a broader development plan may be missed. in the garden route (figure 4), rapid systematic plans were developed as precursors to a robust fine-scale conservation plan (holness, bradshaw & brown 2010; vromans et al. 2010) to ensure that the major priorities were embedded into regional land use plans that were up for review, as well as environmental management frameworks that were being developed. this does not mean that serial prototyping activities should not be constructed to achieve comprehensive goals with products of increasing quality (e.g. for the garden route a formal published product was eventually produced). we recognise that there are ultimately trade-offs between flexibility (such as capacity for frequent updates and maintenance of ‘living products’) and stability, and believe it important for the sanparks scp initiatives to try to stay as near to the flexible edge as possible without losing stability. encouraging ongoing and explicit learning: if management decisions are not seen as learning opportunities and results of recommendations and decisions are not being assessed and reflected upon in an effective way, the central engine for adaptation starts turning more slowly. properly functioning feedbacks are essential to such learning. one way of enhancing learning opportunities is by explicitly setting up contrasting management options. development of sufficient adaptive capacity for action and learning: this includes having a critical mass of skills available, in people who are or are becoming philosophically geared towards such adaptation. this does not include only experts, but also asks for increasing recognition from at least key stakeholders. without this capacity, it is likely that successful adaptation will eventually grind to a halt. we recognise that lack of capacity is often given as a reason for inaction. we consider the better attitude to inculcate to be that if limited resources are anticipated for the foreseeable future, maximal adaptive capacity within the limits of the system’s means should carefully be planned for. maintenance of appreciation for complexity and appropriate attitudes: if both of these are not actively engendered, many individuals may ’roll back’: owing to setbacks in dealing with complexity, they become disheartened and go back to historic patterns of behaviour, even where these have been shown unlikely to be effective in dynamic natural resource or biodiversity settings. a common consequence of internalisation of complexity is that people develop a more modest disposition – and such people seldom tout a single option as ‘the answer’ under all circumstances. shared understanding between groups: if knowledge and derived wisdom do not spread amongst academics, planners, implementers and managers, overall goal achievement inevitably drops. barriers between academics and practitioners, and between agency scientists and managers, are common. perhaps because of recent radical social changes, which generally acted to level the playing field between groups, south africa is often thought to be in a better position in this regard than many other countries, but we do need to take heed of any impending rifts. to ensure alignment with implementation possibilities, a robust and vigorous arena of practitioner-led innovation is required; yet, ongoing academic scrutiny is necessary to ensure conceptual soundness. this underlines the necessity for effective sharing between, as an important and relevant example, these two groups. we believe that applying these five guidelines will go a long way towards maintaining adequate adaptive capacity in scp in the future. conclusion (back to top) this article has highlighted the general similarities between scp and sam as applied within sanparks. we consider it artificial to separate these two processes; in practice, they are two parts of the same system. sam has broader overall applicability to a range of planning, management and decision making situations, whilst scp is a more focused spatial planning tool for a fairly particular task (i.e. the spatial prioritisation of conservation actions). it is our view that scp should philosophically be seen as nested inside sam and that there are significant benefits to approaching scp as an integral part of a broader adaptive management process, rather than trying to add adaptive components to scp. this article has narrated sanparks experiences with scp over the last decade, specifically reviewing the extent to which these can be considered adaptive in the more formal sense defined by the sam processes used in the organisation. comparisons between scp and sam showed pervasive underlying similarities, at least based on the sanparks case history reported here. considerable progress in making scp more adaptive, particularly given the ‘messy’ context in which opportunities present themselves for the productive use of scp, was described. principles were laid out to maintain and enhance this ability to deal with complexity. it is hoped that both sanparks staff and collaborating partners will regard scp as a venture fully inside the set of adaptive principles. references (back to top) berkes, f., colding, j. & folke, c., 2003, navigating social-ecological systems: building resilience for complexity and change, cambridge university press, cambridge. bottrill, m. & pressey, r.l., (in prep.), ‘designs for nature: regional conservation planning, implementation and management’, iucn world commission on protected areas (best practice protected areas guidelines series). cadman, m., petersen, c., driver, a., sekhran, n., maze, k. & munzhedzi, s., 2010, biodiversity for development: south africa’s landscape approach to conserving biodiversity and promoting ecosystem resilience, south african national biodiversity institute, pretoria. cilliers, p., 2000, ‘what can we learn from a theory of complexity?’, emergence 2(1), 23–33. doi:10.1207/s15327000em0201_03 council for scientific and industrial research, 2002, final report: conservation planning framework for the greater addo elephant national park, council for scientific and industrial research, stellenbosch. cowling, r.m., pressey, r.l., rouget, m. & lombard, a.t., 2003a, ‘a conservation plan for a global biodiversity hotspot: the cape floristic region, south africa’, biological conservation 112, 191–216. doi:10.1016/s0006-3207(02)00425-1 cowling, r.m., lombard, a.t., rouget, m., kerley, g.i.h., wolf, t., sims-castley, r., et al., 2003b, ‘a conservation assessment for the subtropical thicket biome’, terrestrial ecology research unit, port elizabeth (report no. 42). driver, a., maze, k., rouget, m., lombard, a.t., nel, j., turpie, j.k., et al., 2005, national spatial biodiversity assessment 2004: priorities for biodiversity conservation in south africa, strelitzia 17, south african national biodiversity institute, pretoria. fischer, j., peterson, g.d., gardner, t.a., gordon, l.j., fazey, i., elmqvist, t., et al., 2009, ‘integrating resilience thinking and optimisation for conservation’, trends in ecology and evolution 24(10), 549–554. doi:10.1016/j.tree.2009.03.020 government of south africa, 2010, national protected areas expansion strategy,government printer, pretoria. grantham, h.s., bode, m., mcdonald-madden, e., game, e.t., knight a.t. & possingham, h.p., 2010, ‘effective conservation planning requires learning and adaptation’, frontiers in ecology and the environment 8, 431–437. doi:10.1890/080151 groves, c.r., 2003, drafting a conservation blueprint: a practitioner’s guide to planning for biodiversity, island press, washington dc. gunderson, l. & holling, c., 2001, panarchy: understanding transformations in human and natural systems, island press, washington dc. holness, s.d, bradshaw, p. & brown, a., 2010, critical biodiversity areas of the garden route, garden route initiative, sanparks, knysna. knight, a.t., cowling, r.m. & campbell, b.m., 2006, ‘an operational model for implementing conservation action’, conservation biology 20(2), 408–419. doi:10.1111/j.1523-1739.2006.00305.x, pmid:16903102 levin, s., 1999, fragile dominion: complexity and the commons, helix books, boston. margules, c.r. & pressey, r.l., 2000, ‘systematic conservation planning’, nature 405, 243−253. doi:10.1038/35012251, pmid:10821285 pressey, r.l. & bottrill, m., 2009, ‘approaches to landscapeand seascape-scale conservation planning: convergence, contrasts and challenges’, oryx 43(4), 464–475. doi:10.1017/s0030605309990500 pressey, r.l, cabeza, m., watts, m.e., cowling, r.m. & wilson, k.a., 2007, ‘conservation planning in a changing world’, trends in ecology and evolution 22(11), 583–592. doi:10.1016/j.tree.2007.10.001 rogers, k.h., 2005, biodiversity custodianship in sanparks: a protected area management planning framework, report to south african national parks, pretoria. roux, d.j. & foxcroft, l.c., 2011, ‘the development and application of strategic adaptive management within south african national parks’, koedoe 53(2), art. #1049, 5 pages. doi:10.4102/koedoe.v53i2.1049 salafsky, n., margoluis, r., redford, k.h. & robinson, j.g., 2002, ‘improving the practice of conservation: a conceptual framework and research agenda for conservation science’, conservation biology 16(6), 1469–1479. doi:10.1046/j.1523-1739.2002.01232.x sanparks. see south african national parks. south african national parks, 2008, ‘policy context: sanparks’ mandate and values’, in south african national parks, a framework for developing and implementing management plans for south african national parks, viewed 15 march 2011, from http://www.sanparks.org/docs/conservation/cpfjanuary2010.pdf snowden, d.j. & boone, m., 2007, ‘a leader’s framework for decision making’, harvard business review november, 69−76. stirzaker, r., biggs, h., roux, d. & cilliers, p., 2010, ‘requisite simplicities to help negotiate complex problems’, ambio 39(8), 600–607. doi:10.1007/s13280-010-0075-7, pmid:21141779 ulanowicz, r., 2009, the third window: natural life beyond newton and darwin, templeton foundation press, west conshohocken. vromans, d.c., maree, k.s., holness, s.d., job, n. & brown, a.e., 2010, the garden route biodiversity sector plan for the george, knysna and bitou municipalities: supporting land-use planning and decision-making in critical biodiversity areas and ecological support areas for sustainable development, garden route initiative, south african national parks, knysna. footnote (back to top) 1. conducted january 2011 on google scholar and science direct using the keywords ‘systematic conservation planning’ and ‘adaptive management’, and examining all papers which had references to both concepts. abstract introduction methods results and discussion conclusion acknowledgements references about the author(s) marietjie landman department of zoology, nelson mandela university, south africa kate kloppers department of zoology, nelson mandela university, south africa graham i.h. kerley department of zoology, nelson mandela university, south africa citation landman, m., kloppers, k. & kerley, g.i.h., 2018, ‘settling the browser–grazer debate for african buffalo in grass-limited eastern cape thicket, south africa’, koedoe 60(1), a1465. https://doi.org/10.4102/koedoe.v60i1.1465 original research settling the browser–grazer debate for african buffalo in grass-limited eastern cape thicket, south africa marietjie landman, kate kloppers, graham i.h. kerley received: 27 feb. 2017; accepted: 20 nov. 2017; published: 22 feb. 2018 copyright: © 2018. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract despite extensive evidence that african buffalo syncerus caffer are grazers, de graaff et al. using rumen content analysis of animals that had starved to death proposed that buffalo in grass-limited eastern cape thicket should be considered browsers. although these anomalous findings were initially accepted, but later challenged, the browse-dominated diet continues to be used as a foundation for hypotheses on the diet of healthy animals. consequently, the debate around buffalo as browsers or grazers in thicket has not yet been settled. we describe the diet of buffalo in the addo elephant national park and include data from other published work from the region to test the importance of grass in buffalo diet. we show that the diet is dominated by grasses, even in grass-limited thicket, and that browse species are seldom dominant foods. thus, there is no empirical evidence to corroborate the notion that buffalo switch their diet to browse when grass availability is low. in an attempt to advance our understanding of buffalo foraging in thicket, we reiterate that de graaff’s work is not a valid measure of buffalo diet in succulent thicket and that additional testing of the browser–grazer hypothesis is not needed. conservation implications: our results confirm that buffalo are grazers, rather than browsers, in grass-limited eastern cape thicket. thus, additional testing of the browser–grazer hypothesis for buffalo in the region is not needed. introduction weak attempts to apply the scientific method may paradoxically improve our understanding if these encourage further robust scientific engagement that recognises the pitfalls of earlier attempts (e.g. hayward et al. 2015; stephens et al. 2015). here we use an example of a diet study of african buffalo syncerus caffer that had starved to death (de graaff, schulz & van der walt 1973), and whose findings were anomalous and initially accepted, only to be challenged later (landman & kerley 2001; novellie, hall-martin & joubert 1991). with this we aim to resolve the debate around buffalo as browsers or grazers in grass-limited eastern cape thicket, south africa. based on their digestive anatomy, hofmann (1989) classified large ruminant herbivores along a continuum from browsers through intermediate feeders to grazers. following this classification, buffalo are specialist grazers, adapted to forage on grasses (as opposed to browse for browsers, or a mixture of grass and browse for intermediate feeders) that are rich in plant cell walls (or fibre). although both the nutritional and physiological bases of hofmann’s classification have been questioned (e.g. gordon & illius 1994; robbins, spalinger & van hoven 1995), most empirical evidence of the food of buffalo corresponds neatly with the grazing-type digestive system. thus, across their range in eastern and southern africa, buffalo diet is dominated by grasses (e.g. cerling, harris & passey 2003; codron et al. 2007; jarman 1971; lamprey 1963; perrin & brereton-stiles 1999; prins 1996; sinclair 1977). however, in the succulent thickets of the eastern cape, de graaff et al. (1973), using rumen content analysis of buffalo that had died in a drought, showed an overabundance of browse in buffalo diet. they presumed that this was a response to low grass availability at the time and proposed that these buffalo should be considered browsers rather than grazers. this led to a number of studies on the food of buffalo in the region (landman & kerley 2001; tshabalala, dube & lent 2009; watermeyer, carroll & parker 2015), each aimed at testing the extent to which buffalo switch their diet between browse and grass. importantly, landman and kerley (2001) concluded that de graaff’s methods were inappropriate for testing diet because their study animals died during a pronounced drought and probably as a consequence of excessive browse to which they were not adapted (novellie et al. 1991). despite this, de graaff’s browse-dominated diet continues to be used as a foundation for hypotheses on the diet of healthy animals (tshabalala et al. 2009; watermeyer et al. 2015). as a consequence, the debate around the feeding strategy of buffalo in thicket has not yet been settled. here we describe the diet of buffalo in the grass-limited succulent thickets of the addo elephant national park. the specific question addressed was how important is grass in buffalo diet in thicket, taking into account other published work from the study site and region (de graaff et al. 1973; landman & kerley 2001; tshabalala et al. 2009; watermeyer et al. 2015). to end off, we consider how our findings influence research perspectives on buffalo resource use in succulent thicket specifically. methods study site we conducted the study in the main camp section (then covering 120 km2) of the addo elephant national park, south africa. during the study, the area supported roughly 300 buffalo and a diverse grazing herbivore community of plains zebra equus quagga, common warthog phacochoerus africanus, eland tragelaphus oryx and red hartebeest alcelaphus buselaphus, and numerous browsers. the region is semi-arid with 260 mm – 530 mm rainfall annually (333 mm over the study period), peaking in late spring (november) and early autumn (march) and experiences frequent droughts. vegetation comprises mostly succulent thicket types (70%), which are evergreen, 2 m – 4 m high and dominated by the tree-succulent portulacaria afra (mucina & rutherford 2006). these thickets are characterised by a high diversity of growth forms: drought-resistant succulents (e.g. p. afra), low trees (e.g. euclea undulata, schotia afra and sideroxylon inerme) and spinescent woody shrubs (e.g. azima tetracantha, capparis sepiaria, carissa spp., gymnosporia spp. and searsia spp.) contribute the bulk of plant biomass, whereas the understory hosts dwarf succulents, forbs and geophytes. although grass availability is generally low and unreliable between years (stuart-hill & aucamp 1993), some species (e.g. cynodon dactylon, digitaria eriantha, eragrostis spp., panicum spp. and sporobolus spp.) may be seasonally abundant in secondary grasslands (covering roughly 15% of the study site) and where intensive utilisation by elephants has removed the canopy shrubs (landman et al. 2014). diet composition we determined the diet of buffalo by identifying plant epidermal fragments in faeces (sparks & malechek 1968). reference slides of the epidermal tissues of potential food items at the site were available from landman and kerley (2001) and landman, schoeman and kerley (2008). holechek, vavra and pieper (1982) described the accuracies and biases of the technique. between january and september 2007, we collected 15 fresh faecal samples in each of the four austral seasons, for a total of 60 samples. seasons were identified based on patterns of temperature, rainfall and frost. faeces were collected opportunistically from apparently healthy buffalo in family groups located throughout the study site, oven-dried and prepared following landman and kerley (2001) and landman et al. (2008). we identified 100 epidermal fragments to species-level per faecal sample and treated each sample as an independent observation. the diet is described as the frequency of occurrence of all the recorded plant species. data analyses to assess the adequacy of our sample size, we generated a mean randomised (50 iterations) accumulation curve of plant species recorded per faecal sample. however, because this curve did not reach a clear asymptote, we estimated total dietary richness with a non-parametric species richness estimator (foggo et al. 2003). differences between observed and expected counts provide an estimate of the variation in dietary information at the upper limit of sampling effort. we generally combined the seasonal data and described the diet in terms of principal diet items and by grouping all plant species into broad growth form categories (i.e. grasses, woody shrubs, succulents, forbs and climbers). principal diet items are those foods consumed in the greatest quantities and were identified as the plant species that contributed > 3% of the diet (landman & kerley 2001; tshabalala et al. 2009). anova procedures (tukeys’ test) were used to test for differences in growth form categories and the number of plant species recorded in the diet across seasons. data were inspected for deviations from normality and homogeneity of variances prior to analyses. to develop broader insight into the range of buffalo diet in succulent thicket, we draw data from two previous studies at the study site (de graaff et al. 1973; landman & kerley 2001) and two others in the region (great fish river nature reserve – tshabalala et al. 2009; kwandwe private game reserve – watermeyer et al. 2015). because de graaff’s study animals died during a drought, we use their data only to support the presence of plant species already recorded. results and discussion this study identified 37 plant species in the diet of buffalo in 60 faecal samples (table 1). these species accounted for roughly 79% of the estimated richness at the upper limit of sampling effort, suggesting that our sample size was adequate to describe the diet. although the observed richness is comparable to that recorded previously (e.g. landman & kerley 2001; tshabalala et al. 2009; venter & watson 2008), 17 species were new to the diet in thicket and the study site specifically (table 1). importantly, > 80% of the browse species recorded in the rumen samples of de graaff et al. (1973) were not recorded in the diet anywhere else in the region. by combining our data with those from others in the region (landman & kerley 2001; tshabalala et al. 2009), we show that buffalo have a relatively broad diet – 44 plant species – exceeding that of other grass-eating ruminants in thicket (e.g. eland, red hartebeest – kerley & landman 2006). thus, buffalo foraging may rely on more plant species than previously thought, and we contribute towards understanding its role in succulent thicket and the addo elephant national park specifically. table 1: plant species identified in the diet of buffalo in the succulent thickets of the eastern cape, south africa. based on their characteristic grazing-type digestive system (hofmann 1989), grasses contributed the bulk of the diet (mean = 75.6%, s.d. = 12.7%) in our study. the remaining browse comprised woody (mean = 23.6%, s.d. = 12.2%) and succulent (mean = 0.8%, s.d. = 2.2%) shrubs and climbers (mean = 0.1%, s.d. = 0.4%), with few browse species contributing sufficiently to the diet to be considered dominant (i.e. > 3%). instead, principal diet items were characterised by grasses across study populations (table 1). our results are within the range of mean grass consumption by buffalo in the region (71.9% – landman & kerley 2001; ± 75% – tshabalala et al. 2009; 93.1% – watermeyer et al. 2015) and support all previous work on their food resources (e.g. cerling et al. 2003; codron et al. 2007; jarman 1971; lamprey 1963; perrin & brereton-stiles 1999; prins 1996; sinclair 1977), including that from other grass-limited habitats (venter & watson 2008) considered marginal to buffalo (i.e. nama-karoo, boshoff, landman & kerley 2016). seasonal variations in diet are well described, particularly as buffalo increase their utilisation of browse towards the dry season when grass availability and quality decline (e.g. prins 1996; sinclair 1977; tshabalala et al. 2009; watermeyer et al. 2015). although we found no seasonal differences in the contribution of browse to the diet in our study (range across seasons: 19.7% – 30.0%; f3,56 = 2.25, p = 0.09), we detected a gradual increase in diet breadth (from 19 spp. to 35 spp.) towards the dry season that coincided with an increase in the utilisation of browse species (from 26% [spring] to 54% [summer] of plant species recorded; f3,56 = 4.22, p = 0.009). these findings follow the predictions of optimality theory which predicts that a loss of preferred foods (such as grasses for buffalo – prins 1996; sinclair 1977) would cause an increase in the utilisation of less nutritious, non-preferred foods (owen-smith & novellie 1982). thus, despite the fact that we had no information on food availability and could not test food preferences per se, we presume that the broader diet and increased utilisation of browse species during the dry season reflects changing diet with changing grass availability and quality. from our results, it is clear that buffalo diet is dominated by grasses, even in grass-limited succulent thicket (landman & kerley 2001; tshabalala et al. 2009; watermeyer et al. 2015). although the diet might include browse in varying quantity between seasons, depending on the availability and quality of grass, browse species are seldom dominant foods and many are probably incidentally utilised as parts of larger mouthfuls. thus, with the exception of the findings of de graaff et al. (1973), there is no empirical evidence to support the notion that buffalo switch their diet to browse when grass availability is low (cerling et al. 2003; codron et al. 2007; jarman 1971; lamprey 1963; perrin & brereton-stiles 1999; prins 1996; sinclair 1977; venter & watson 2008). instead, de graaff’s work generated confusion in our understanding of buffalo foraging in thicket through poor experimental design. even though such weaknesses can undermine science and management (e.g. hayward et al. 2015; stephens et al. 2015), this might still improve our understanding if it encourages additional robust interrogation. for buffalo in thicket, additional testing confirmed what was already known about their diet, which probably limited our ability to develop broader insights because our approaches were motivated by a single hypothesis. thus, in an attempt to advance our understanding of buffalo foraging in thicket, we reiterate the conclusions of landman and kerley (2001) that de graaff’s work is not a valid measure of buffalo diet in succulent thicket and that additional testing of the browser–grazer hypothesis is not needed. conclusion our study questioned the importance of grass in buffalo diet in grass-limited eastern cape thicket. we show that the diet is dominated by grasses, even in grass-limited thicket, and that browse species are seldom dominant foods. thus, there is no empirical evidence to corroborate the notion that buffalo switch their diet to browse when grass availability is low. with this we argue that additional testing of the browser–grazer hypothesis for buffalo in the region is not needed. acknowledgements the authors thank south african national parks for providing permission to conduct this study. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions m.l., k.k. and g.i.h.k. designed the study, analysed the data and wrote the article. references boshoff, a., landman, m. & kerley, g.i.h., 2016, ‘filling the gaps on the maps: historical distribution patterns of some of the larger mammals in part of southern africa’, transactions of the royal society of south africa 71, 23–87. https://doi.org/10.1080/0035919x.2015.1084066 cerling, t., harris, j. & passey, b., 2003, ‘diets of east african bovidae based on stable isotope analysis’, journal of mammalogy 84, 456–470. https://doi.org/10.1644/1545-1542(2003)084%3c0456:doeabb%3e2.0.co;2 codron, d., codron, j., lee-thorp, j.a., sponheimer, m., de ruiter, d., sealy, j. et al., 2007, ‘diets of savanna ungulates from stable carbon isotope composition of faeces’, journal of zoology, london 273, 21–29. https://doi.org/10.1111/j.1469-7998.2007.00292.x de graaff, g., schulz, k.c.a. & van der walt, p.t., 1973, ‘notes on rumen contents of cape buffalo syncerus caffer in the addo elephant national park’, koedoe 16, 45–58. https://doi.org/10.4102/koedoe.v16i1.884 foggo, a., attrill, m.j., frost, m.t. & rowden, a.a., 2003, ‘estimating marine species richness: an evaluation of six extrapolative techniques’, marine ecology progress series 248, 15–26. https://doi.org/10.3354/meps248015 gordon, i.j. & illius, a.w., 1994, ‘the functional significance of the browser-grazer dichotomy in african ruminants’, oecologia 98, 167–175. https://doi.org/10.1007/bf00341469 hayward, m.h., boitani, l., burrows, n.d., funston, p.j., ullas karath, k., mackenzie, d.i. et al., 2015, ‘ecologists need robust survey designs, sampling and analytical methods’, journal of applied ecology 52, 286–290. https://doi.org/10.1111/1365-2664.12408 hofmann, r.r., 1989, ‘evolutionary steps of ecophysiological adaptation and diversification of ruminants: a comparative view of their digestive system’, oecologia 78, 443–457. https://doi.org/10.1007/bf00378733 holechek, j.l., vavra, m. & pieper, r.d., 1982, ‘botanical composition determination of range herbivore diets: a review’, journal of range management 35, 309–315. https://doi.org/10.2307/3898308 jarman, p.j., 1971, ‘diets of large mammals in the woodlands around lake kariba, rhodesia’, oecologia 8, 157–178. https://doi.org/10.1007/bf00345811 kerley, g.i.h. & landman, m., 2006, ‘the impacts of elephants on biodiversity in the eastern cape subtropical thickets’, south african journal of science 102, 395–402. lamprey, h.f., 1963, ‘ecological separation of the large mammal species in the tarangire game reserve, tanganyika’, east africa wildlife journal 1, 63–92. https://doi.org/10.1111/j.1365-2028.1963.tb00179.x landman, m. & kerley, g.i.h., 2001, ‘dietary shifts: do grazers become browsers in the thicket biome?’, koedoe 44, 31–36. https://doi.org/10.4102/koedoe.v44i1.183 landman, m., kerley, g.i.h. & schoeman, d.s., 2008, ‘relevance of elephant herbivory as a threat to important plants in the addo elephant national park, south africa’, journal of zoology, london 274, 51–58. landman, m., schoeman, d.s., hall-martin, a.j. & kerley, g.i.h., 2014, ‘long-term monitoring reveals differing impacts of elephant on elements of a canopy shrub community’, ecological applications 24, 2002–2012. https://doi.org/10.1890/14-0080.1 mucina, l. & rutherford, m.c., 2006, ‘the vegetation of south africa, lesotho and swaziland’, strelitzia 19, 1–807. novellie, p., hall-martin, a.j. & joubert, d., 1991, ‘the problem of maintaining large herbivores in small conservation areas: deterioration of the grassveld in the addo elephant national park’, koedoe 34, 41–50. https://doi.org/10.4102/koedoe.v34i1.413 owen-smith, n. & novellie, p., 1982, ‘what should a clever ungulate eat?’, american naturalist 119, 151–178. https://doi.org/10.1086/283902 perrin, m.r. & brereton-stiles, r., 1999, ‘habitat use and feeding behavior of the buffalo and the white rhinoceros in the hluhluwe-umfolozi game reserve’, south african journal of wildlife research 29, 72–80. prins, h.h.t., 1996, ecology and behavior of the african buffalo, chapman & hall, london. robbins, c.t., spalinger, d.e. & van hoven, w., 1995, ‘adaptation of ruminants to browse and grass diets: are anatomical-based browser-grazer interpretations valid?’, oecologia 103, 208–213. https://doi.org/10.1007/bf00329082 sinclair, a., 1977, the african buffalo: a study of resource limitation of populations, university of chicago press, chicago, il. sparks, d.r. & malechek, j.c., 1968, ‘estimating percentage dry weight in diets using a microscopic technique’, journal of range management 21, 264–265. https://doi.org/10.2307/3895829 stephens, p.a., pettorelli, n., barlow, j., whittingham, m.j. & cadotte, m.w., 2015, ‘management by proxy? the use of indices in applied ecology’, journal of applied ecology 52, 1–6. https://doi.org/10.1111/1365-2664.12383 stuart-hill, g.c. & aucamp, a.j., 1993, ‘carrying capacity of the succulent valley bushveld of the eastern cape’, african journal of range and forage science 10, 1–10. https://doi.org/10.1080/10220119.1993.9638314 tshabalala, t., dube, s. & lent, p.c., 2009, ‘seasonal variation in forages utilized by the african buffalo (syncerus caffer) in the succulent thicket of south africa’, african journal of ecology 48, 438–445. https://doi.org/10.1111/j.1365-2028.2009.01132.x venter, j.a. & watson, l.h., 2008, ‘feeding and habitat use of buffalo (syncerus caffer caffer) in the nama-karoo, south africa’, south african journal of wildlife research 38, 42–51. https://doi.org/10.3957/0379-4369-38.1.42 watermeyer, j.p., carroll, s.l. & parker, d.m., 2015, ‘seasonal consumption of browse by the african buffalo (syncerus caffer) in thicket biome of south africa’, african journal of ecology 53, 599–601. https://doi.org/10.1111/aje.12214 abstract introduction methods results and discussion conclusion and recommendations acknowledgements references footnotes about the author(s) sam m. ferreira scientific services, south african national parks, skukuza, south africa cathy greaver scientific services, south african national parks, skukuza, south africa chenay simms scientific services, south african national parks, skukuza, south africa citation ferreira, s.m., greaver, c. & simms, c., 2017, ‘elephant population growth in kruger national park, south africa, under a landscape management approach’, koedoe 59(1), a1427. https://doi.org/10.4102/koedoe.v59i1.1427 short communication elephant population growth in kruger national park, south africa, under a landscape management approach sam m. ferreira, cathy greaver, chenay simms received: 27 july 2016; accepted: 12 june 2017; published: 25 aug. 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract south african national parks (sanparks) manage landscapes rather than numbers of elephants (loxodonta africana) to mitigate the effects that elephants may have on biodiversity, tourism and stakeholder conservation values associated with protected areas. this management philosophy imposes spatial variability of critical resources on elephants. restoration of such ecological processes through less intensive management predicts a reduction in population growth rates from the eras of intensive management. we collated aerial survey data since 1995 and conducted an aerial total count using a helicopter observation platform during 2015. a minimum of 17 086 elephants were resident in the kruger national park (knp) in 2015, growing at 4.2% per annum over the last generation of elephants (i.e. 12 years), compared to 6.5% annual population growth noted during the intensive management era ending in 1994. this may come from responses of elephants to density and environmental factors manifested through reduced birth rates and increased mortality rates. authorities should continue to evaluate the demographic responses of elephants to landscape scale interventions directed at restoring the limitation of spatial variance in resource distribution on elephant spatiotemporal dynamics and the consequences that may have for other conservation values. conservation implications: conservation managers should continue with surveying elephants in a way that allows the extraction of key variables. such variables should focus on measures that reflect on how theory predicts elephants should respond to management interventions. introduction african savanna elephants (loxodonta africana) significantly influence several conservation management objectives that authorities seek to achieve in protected areas. managers of protected areas typically seek to conserve biodiversity – elephants act as ecological engineers that may alter habitats and various aspects of biological diversity (valeix et al. 2011). protected areas also serve as focal places where elephant presence forms a key component of tourist attractions and experiences (kerley, geach & vial 2003). for people living next to parks, elephants leaving protected areas threaten livelihoods and lives (osborn & parker 2003). south african national parks (sanparks) seek to manage these effects that elephants have on the ability of managers to achieve conservation management objectives. the effects that elephants have on biodiversity, tourist experiences and rural people’s livelihoods may associate primarily with their location rather than abundance or density (van aarde, jackson & ferreira 2006). in small parks, however, density of elephants may aggravate the effects they have on biodiversity. the way elephants use landscapes depends on the distribution of water (chamaillé-jammes, valeix & fritz 2007; loarie, van aarde & pimm 2009), food (codron et al. 2006) and shade (kinahan, pimm & van aarde 2007) – resources that elephants select at different intensities during different seasons (young, ferreira & van aarde 2009a). in addition, elephants avoid people by choice (douglas-hamilton, krink & vollrath 2005; graham et al. 2009). within this context sanparks embraced a landscape approach to elephant management and seek to restore the landscape variability and limitations on elephant spatial and temporal dynamics (harris et al. 2008). within kruger national park (knp), this translated to stopping the control of elephant numbers through culling (van aarde, whyte & pimm 1999), the closure of boreholes and destruction of several earthen dams (gaylard, owen-smith & redfern 2003), the removal of fences between kruger national park and adjacent conservation-friendly land-uses (venter, naiman, biggs & pienaar 2008), and the use of a fire management policy focusing on patch burns (van wilgen et al. 2004). the creation of spatial and temporal heterogeneity in elephant resources predicts responses in how elephants use landscapes. elephants use landscapes more variably (smit & ferreira 2010) following the restoration of various ecological processes, where previous management philosophies and approaches fostered homogeneity (venter et al. 2008). variability in landscape use also predicts population numerical responses – when daily distances travelled by cows increase, weaned calves have higher mortality rates (young & van aarde 2010). the closure of boreholes increased distances and configuration (gaylard 2015) required for travel between water and good quality food. in addition, when sanparks stopped culling, elephant densities increased (van aarde, whyte & pimm 1999). higher densities associated with a less clumped distribution of elephants across the kruger landscape (young, ferreira & van aarde 2009b). these mechanisms predict decreased population growth rates with increasing density and decreasing water provisioning. sanparks seek to evaluate the consequences of reduced management intensity directed at mitigating the effects of elephants on various conservation management objectives. monitoring focuses on spatial and temporal responses of elephants. as part of this monitoring, sanparks survey elephants to derive estimates of population variables. here we report on the knp elephant survey of 2015, trends extracted from recent surveys (sanparks, unpublished data1) and check population growth rates post 1994 against those at the end of the intensive elephant management era in the mid-1990s (whyte 2001). methods the knp, 19 485 km² in size, spans the low-lying savannas of the eastern parts of the limpopo and mpumalanga provinces, south africa. mozambique abuts the park in the east, while zimbabwe forms the boundary to the north. the park has a mean annual rainfall that varies from 750 mm in the south to 450 mm in the north, with ± 80% of rain falling during summer (gertenbach 1980). granite and gneiss soils dominate knp’s western half and nutrient-rich basalt soils dominate the eastern half, with a narrow band of karoo sediment occurring at the junction of the granite and basalt (schutte 1986). the vegetation on the southern basalts is largely wooded savanna, with sclerocarya caffra and vachellia nigrescens dominating the tree canopy. mixed combretum spp. and vachellia spp. dominate the southern granites. in the north colophospermum mopane dominates both the basalt and granite substrates (gertenbach 1983). this produces a diverse range of landscapes resulting in spatial and temporal heterogeneity of resources that elephants use. water is an essential resource provided through several perennial (i.e. crocodile, sabie, olifants and letaba) and large seasonal rivers (i.e. biyamiti, nwaswitshaka, nwatswitsontso, timbavati, tsendze, shingwedzi, levhuvhu and limpopo). the provision of additional water, through drilling of boreholes and construction of several dams that started in 1933, disrupted spatial and temporal heterogeneity of water availability and quality. knp had 365 boreholes (gaylard, owen-smith & redfern 2003) and 109 pipeline troughs, weirs, concrete and earthen dams by 1995. since then, the closure of boreholes, breaching of dams and removal of structures resulted in 41 boreholes and 31 pipeline troughs, weirs, concrete and earthen dams remaining functional during our study.2 we applied a total count approach to survey elephants living in knp during 2015, using the same technique as used by sanparks since 1967 (whyte 2001). the approach divides the park into 21 blocks and uses a helicopter-based observation platform to search each block for elephants. flight paths, flown at 90–100 knots at a height of 91 m above ground level, focused on 2nd order river catchments approximately 500 m – 1000 m parallel to the second order stream. the distance between flight paths vary depending on vegetation cover. we used two helicopters to survey blocks during august 2015 over a 21-day period. each helicopter, surveying a separate block, was crewed by a pilot, two observers and a person recording data. the survey team noted the geo-referenced positions of elephant groups encountered, recorded the number of elephants in the group, identified the group as a bull or mixed herd and counted calves less than 1 year old (see trimble et al. 2011) for mixed herds. from these surveys, we summed all observations to find the total number of elephants observed. we collated the total number of elephants observed in surveys since 1995. these values are estimates of minimum numbers of elephants known to be alive and do not account for availability, detection probability, observer bias or statistical sampling error (caughley 1974). we fitted population models that included exponential: ricker: and heterogeneity: models where nt is minimum number of elephants alive at time t, r is exponential growth rate, rm is maximum growth rate, k is an estimate of equilibrium population size, r is mean annual rainfall and α is a constant. we estimated r, rm, k and α using maximum likelihood approaches (edwards 1972; microsoft excel macro provided by hood 2005). we used akaike’s information criteria (aicc) to select the most appropriate model (johnson & omland 2004). when differences between aicc values were less than 2, we used model averaging (claeskens & hjort 2008) to obtain a population growth estimate and associated confidence intervals. the recording of calves allowed us to derive a recruitment rate as the fraction of elephants less than 1 year old. such a recruitment rate is the consequence of calves born and their subsequent survival during the first year. to estimate overall elephant survival (s) we calculated an age distribution parameter a, the rate at which age-specific frequencies decay with age in stable age distributions from: where nx is the frequency of individual x years of age and w is elephant longevity (ferreira & van aarde 2008). we set w at 55 years (lee et al. 2016). we estimated a using maximum likelihood approaches (edwards 1972; microsoft excel macro provided by hood 2005). this allowed us to calculate survival from a = λs (eberhardt 1988) where λ is finite population growth (λ = 1+r). to minimise the consequences of variation in age distributions, we recalculated population growth using only estimates spanning an elephant generation defined as the age at which member of a given cohort is expected to reproduce (charlesworth 1994), that is, 12 years (robson 2015) from 2004 until 2015 and used this as input to calculate survival rate. we could then estimate birth rate by dividing recruitment rate with the average estimated survival rate across all age classes. sanparks’ intensive management philosophy ceased in 1994 that is, culling stopped, water provisioning decreased, some fences were removed and patch burns were implemented since then. we thus extracted population growth rates as well as estimates of survival and recruitment rates prior to 1994 (whyte 2001) to check against recent estimates of these variables. note that mean annual rainfall prior to 1994 was 510.1 mm (se = 38.2, n = 18) similar to that after culling stopped (565.1, se = 44.5, n = 20) and from 2004 to 2015 (529.4, se = 34.0, n = 11). we thus did not consider rainfall driven influences in our comparisons. results and discussion we encountered elephants throughout knp (figure 1). two of the large perennial rivers dissecting the park – the olifants and sabie rivers served as boundaries for regional estimates. we provide estimates for areas north of the olifants river, between the olifants and sabie rivers and south of the sabie river separately. observers encountered 7107 and 1163 elephants in mixed (n = 527) and bull herds (n = 546), respectively, north of the olifants river at a density of 0.84 n.km-2. between the sabie and olifants rivers observers noted 3526 and 462 elephants living in mixed (n = 289) and bull herds (n = 229), respectively, at a density of 0.81 n.km-2. highest density of 1.15 n.km-2 was south of the sabie river and came from observers recording 4232 and 596 elephants in mixed (n = 306) and bull herds (n = 302), respectively. these regional counts resulted in a minimum of 17 086 elephants living in knp during 2015. figure 1: the distribution of elephants recorded during the survey of the kruger national park in 2015. models describing the trends in minimum elephants known to be alive after culling stopped were equivalent (r2 values ranged from 0.96 to 0.98) (table 1). model averaging (figure 2) thus recorded annual exponential growth rate since 1995 at 4.9% (2.1% – 7.6%) per annum, similar to that reported previously (young et al. 2009b). the point estimates of 6.5% reported prior to 1995 (whyte 2001), the period of intensive management, is located at the 86.7th percentile of growth since 1995. population growth since 2003, an elephant generation, was 4.2% (1.1% – 7.3%) with 6.5% located at the 92.7th percentile of growth since 2004. these results may reflect a likely reduction in elephant population growth rate over time and as densities increased supporting the prediction that population growth rates should be declining. note that poaching had little effect on the elephants living in knp with between 0 and 22 elephants poached during any year (sanparks, unpublished data3). it is uncertain, however, how poaching of elephants could change in future and what the impact would be on the population in knp. figure 2: minimum number of elephants living in the kruger national park since 1995 after intensive management of elephants stopped in 1994. we also illustrate the average model (line) and its 95% confidence intervals (thin lines) since then. table 1: population model comparisons made for minimum number of elephants living in the kruger national park since 1995. we acknowledge that movements of elephants to areas abutting knp can also account for reduction in population growth rates that we noted. for this reason, we also estimated vital rates given that previous studies implied variance in birth rates (robson 2015) and declines in survival rates as mechanisms associated with increase landscape limitations on elephant resources (young et al. 2009b; young & van aarde 2010). the estimated population growth since 2003 and an estimated age distribution parameter over that same period (a = 0.94 ± 0.02, n = 10) allowed us to estimate survival during the first year at 0.982 (0.941–0.999), while other age classes had a survival rate of 0.983 (0.954–0.999). this estimate ignores age-specific survival rates for elephants older than 1 year. the overall recruitment rate (5.67%, 5.23% – 6.02%) predicted an annual birth rate of 5.78% (5.33% – 6.41%). note that during the period of intensive management estimates of annual birth rates (expressed as potential rates of increase – i.e. rate of increase in the absence of mortality) ranged from 7.8% to 16.8% (whyte 2001). demographic responses of elephants to restoration of ecological processes may thus provide some potential mechanisms explaining how elephant growth rates declined since 1995. recent analyses, however, highlight that variability reflecting on ecological productivity during the conception and gestation phases of elephant reproduction drives variability in elephant birth rates (robson 2015). the estimated survival rate translates to an overall mortality rate of 1.8% (1.72% – 1.88%) higher than the point estimated of 1.5% reported for the period of intensive management (whyte 2001). we are mindful of confounding influences of different methods, but our results could support the predictions derived from mechanism of how changes in resource distribution and density influence survival schedules of elephants (young & van aarde 2010). we acknowledge though that sanparks need estimates of age-specific survival to evaluate the various predictions made on how daily distances travelled as a result of variability in resources and elephant density influence survival of specifically weaned calves (young & van aarde 2010). conclusion and recommendations the temporal responses of elephants following the implementation of a philosophy of managing landscapes rather than managing elephants directly (van aarde, jackson & ferreira 2006) support the predictions that elephant population growth rate slows when elephant resources are heterogeneously distributed across the landscape. a minimum of 17 086 elephants were present in knp during 2015 with their temporal dynamics reflecting declining growth rates coming from interactions between decreased birth and increased death rates. sanparks should continue monitoring elephants and check temporal responses complimented by demographic studies in concert with spatial responses derived through distribution and satellite studies. the association between elephant responses to management and the significance of elephants for biodiversity, tourism and stakeholder values associated with protected areas (but see guldemond & van aarde 2008) remains a key requirement. acknowledgements we are grateful to the following observers: sandra visagie, marius snyders, andrew desmet, joe nkuna, richard sowry, rob thompson, pauli viljoen, jacques venter, adolf manganyi, steven oosthuizen, scott ronaldson, ben wigley, corli wigley, izak smit, rudi lorist and the pilots grant knight, charles thompson, jaco mol and brad grafton. we thank dr mike chase, director of elephants without borders, who provided part of the funding through the paul alan foundation. the remaining funding was provided from sanparks baseline funding. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions c.g. coordinated the aerial survey but all authors participated in the aerial survey. s.m.f. performed the statistical analysis. c.s. performed the arcgis analyses. s.m.f. led the writing while c.g. and c.s. assisted with writing and editing. references caughley, g., 1974, ‘bias in aerial survey’, the journal of wildlife management 38(4), 921–933. https://doi.org/10.2307/3800067 chamaille-jammes, s., valeix, m. & fritz, h., 2007, ‘managing heterogeneity in elephant distribution: interactions between elephant population density and surface-water availability’, journal of applied ecology 44(3), 625–633. https://doi.org/10.1111/j.1365-2664.2007.01300.x charlesworth, b., 1994, evolution in age-structured populations, pp. 28–30, university of cambridge press, cambridge. claeskens, g. & hjort, n.l., 2008, model selection and model averaging, vol. 330, cambridge university press, cambridge. codron, j., lee-thorp, j.a., sponheimer, m., codron, d., grant, r.c. & de ruiter, d.j., 2006, ‘elephant (loxodonta africana) diets in kruger national park, south africa: spatial and landscape differences’, journal of mammalogy 87(1), 27–34. https://doi.org/10.1644/05-mamm-a-017r1.1 douglas-hamilton, i., krink, t. & vollrath, f., 2005, ‘movements and corridors of african elephants in relation to protected areas’, naturwissenschaften 92(4), 158–163. https://doi.org/10.1007/s00114-004-0606-9 eberhardt, l.l., 1988, ‘using age structure data from changing populations’, journal of applied ecology 25, 373–378. https://doi.org/10.2307/2403829 edwards, a.w.f., 1972, likelihood, cambridge university press, cambridge. ferreira, s.m. & van aarde, r.j., 2008, ‘a rapid method to estimate population variables for african elephants’, the journal of wildlife management 72(3), 822–829. https://doi.org/10.2193/2007-151 gaylard, a., 2015, ‘adopting a heterogeneity paradigm for understanding and managing elephants for biodiversity: a case study in riparian woodlands in kruger national park’, phd dissertation, university of the witwatersrand, johannesburg, south africa. gaylard, a., owen-smith, n. & redfern, j., 2003, ‘surface water availability: implications for heterogeneity and ecosystem processes’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 171–188, island press, washington, dc. gertenbach, w.d., 1980, ‘rainfall patterns in the kruger national park’, koedoe 23(1), 35–43. https://doi.org/10.4102/koedoe.v23i1.634 gertenbach, w.d., 1983, ‘landscapes of the kruger national park’, koedoe 26(1), 9–121. https://doi.org/10.4102/koedoe.v26i1.591 graham, m.d., douglas-hamilton, i., adams, w.m. & lee, p.c., 2009, ‘the movement of african elephants in a human-dominated land-use mosaic’, animal conservation 12(5), 445–455. https://doi.org/10.1111/j.1469-1795.2009.00272.x guldemond, r. & van aarde, r., 2008, ‘a meta-analysis of the impact of african elephants on savanna vegetation’, the journal of wildlife management 72(4), 892–899. https://doi.org/10.2193/2007-072 harris, g.m., russell, g.j., van aarde, r.j. & pimm, s.l., 2008, ‘rules of habitat use by elephants loxodonta africana in southern africa: insights for regional management’, oryx 42(1), 66–75. https://doi.org/10.1017/s0030605308000483 hood, g.m., 2005, poptools version 2.6.6, viewed 01 june 2017, from http://poptools.org/ johnson, j.b. & omland, k.s., 2004, ‘model selection in ecology and evolution’, trends in ecology and evolution 19(2), 101–108. https://doi.org/10.1016/j.tree.2003.10.013 kerley, g.i., geach, b.g. & vial, c., 2003, ‘jumbos or bust: do tourists’ perceptions lead to an under-appreciation of biodiversity?’, south african journal of wildlife research 33(1), 13–21. kinahan, a.a., pimm, s.l. & van aarde, r.j., 2007, ‘ambient temperature as a determinant of landscape use in the savanna elephant, loxodonta africana’, journal of thermal biology 32(1), 47–58. https://doi.org/10.1016/j.jtherbio.2006.09.002 lee, p.c., fishlock, v., webber, c.e. & moss, c.j., 2016, ‘the reproductive advantages of a long life: longevity and senescence in wild female african elephants’, behavioral ecology and sociobiology 70, 337–345. https://doi.org/10.1007/s00265-015-2051-5 loarie, s.r., van aarde, r.j. & pimm, s.l., 2009, ‘fences and artificial water affect african savannah elephant movement patterns’, biological conservation 142(12), 3086–3098. https://doi.org/10.1016/j.biocon.2009.08.008 osborn, f.v. & parker, g.e., 2003, ‘towards an integrated approach for reducing the conflict between elephants and people: a review of current research’, oryx 37(1), 80–84. https://doi.org/10.1017/s0030605303000152 robson, a.s., 2015, ‘demographic responses to changes in conservation management: a case study on elephants in the kruger national park’, msc-thesis, university of pretoria, pretoria, south africa. schutte, i.c., 1986, ‘the general geology of the kruger national park’, koedoe 29(1), 13–37. https://doi.org/10.4102/koedoe.v29i1.517 smit, i.p. & ferreira, s.m., 2010, ‘management intervention affects river-bound spatial dynamics of elephants’, biological conservation 143(9), 2172–2181. https://doi.org/10.1016/j.biocon.2010.06.001 trimble, m.j., van aarde, r.j., ferreira, s.m., nørgaard, c.f., fourie, j., lee, p.c. & moss, c.j., 2011, ‘age determination by back length for african savanna elephants: extending age assessment techniques for aerial-based surveys’, plos one 6(10), e26614. https://doi.org/10.1371/journal.pone.0026614 valeix, m., fritz, h., sabatier, r., murindagomo, f., cumming, d. & duncan p., 2011, ‘elephant-induced structural changes in the vegetation and habitat selection by large herbivores in an african savanna’, biological conservation 144(2), 902–912. https://doi.org/10.1016/j.biocon.2010.10.029 van aarde, r.j., jackson, t.p. & ferreira, s.m., 2006, ‘conservation science and elephant management in southern africa’, south african journal of science 102(9/10), 385. van aarde, r.j., whyte, i. & pimm, s., 1999, ‘culling and the dynamics of the kruger national park african elephant population’, animal conservation 2(4), 287–294. https://doi.org/10.1111/j.1469-1795.1999.tb00075.x van wilgen, b.w., govender, n., biggs, h.c., ntsala, d., & funda, x.n., 2004, ‘response of savanna fire regimes to changing fire-management policies in a large african national park’, conservation biology 18(6), 1533–1540. https://doi.org/10.1111/j.1523-1739.2004.00362.x venter, f.j., naiman, r.j., biggs, h.c. & pienaar, d.j., 2008, ‘the evolution of conservation management philosophy: science, environmental change and social adjustments in kruger national park’, ecosystems 11(2), 173–192. https://doi.org/10.1007/s10021-007-9116-x whyte, i.j., 2001, ‘the conservation management of the kruger national park elephant population’, phd thesis, university of pretoria, pretoria, south africa. young, k.d., ferreira, s.m. & van aarde, r.j., 2009a, ‘elephant spatial use in wet and dry savannas of southern africa’, journal of zoology 278(3), 189–205. https://doi.org/10.1111/j.1469-7998.2009.00568.x young, k.d., ferreira, s.m. & van aarde, r.j., 2009b, ‘the influence of increasing population size and vegetation productivity on elephant distribution in the kruger national park’, austral ecology 34(3), 329–342. https://doi.org/10.1111/j.1442-9993.2009.01934.x young, k.d. & van aarde, r.j., 2010, ‘density as an explanatory variable of movements and calf survival in savanna elephants across southern africa’, journal of animal ecology 79(3), 662–673. https://doi.org/10.1111/j.1365-2656.2010.01667.x footnotes 1. sanparks data depository, judith botha, judith.botha@sanparks.org 2. sanparks kruger national park conservation management, stephen midzi, stephen.midzi@sanparks.org 3. ken maggs, ken.maggs@sanparks.org haacke.qxd variation in population size of bouton’s snake-eyed skink (reptilia: scincidae) at black rock in northern kwazulu-natal, south africa w.d. haacke haacke, w.d. 2002. variation in population size of bouton’s snake-eyed skink (reptilia: scincidae) at black rock in northern kwazulu-natal, south africa. koedoe 45(1): 93–100. pretoria. issn 0075-6458. bouton’s snake-eyed skink cryptoblepharus boutonii sp. occurs in scattered island or coastal populations in the indian ocean. the most southern known population occurs on black rock on the northern coast of kwazulu-natal. this tiny population was monitored over a period of 14 summer seasons and a final check was made 10 seasons later. this population consisted of an average of about 58 individuals, but has fluctuated by more than 100 %, suggesting that its existence is very tenuous. the re-check during october 2001 produced very positive figures, indicating that this population, observed over 23 years, is doing very well, is maintaining its numbers within acceptable parameters and is in a good position to survive without special precautions. key words: bouton’s snake-eyed skink, cryptoblepharus boutoni, black rock. w.d. haacke, nfi, transvaal museum, p.o. box 413, pretoria, 0001, south africa issn 0075-6458 93 koedoe 45/1 (2002) introduction bouton’s snake-eyed skink is a small, slender skink (fig. 1), of which the individuals of the population under discussion reach a headbody length of just under 50 mm and a tail which may be nearly 20 mm longer. its preferred habitat is the intertidal zone of rocky coastlines and its distribution appears to have been achieved by passive rafting (wilson & knowles 1988; spawls et al. 2002). only a single population of this snake-eyed skink, presently still referred to as cryptoblepharus boutonii africanus (sternfeld), is known to occur within the limits of the republic of south africa (haacke 1977; mclachlan 1978; bruton & haacke 1980; haacke 1988; branch 1988; branch 1998). the locality, black rock (27°08's, 32°50'e; 2732bc), is a coastal cliff on the maputaland coast about 35 km south of the mozambique border in northern kwazulu-natal (fig. 2.). the next and nearest apparently conspecific population is known from about 550 km further north near inhambane on the mozambique coast. mertens (1931, 1933, 1934, 1963) viewed what is now the crytoblepharus complex (fuhn 1970) as 36 subspecies of ablepharus boutonii (desjardin) (type locality: roche noir [= black rock], mauritius) the lizard species with the widest distribution in the world, which extended from the african east coast, across various islands of the indian ocean, australia and islands of the pacific as far as the easter islands. in contrast, a reassessment by storr (1976) suggested a much more complicated species complex and, by raising the status of many taxa to full species, he fragmented this range considerably. the most important study concerning indian ocean populations is that of brygoo (1986). welch (1982), without giving any reasons, raised all the taxa in the indian natural history note haacke.qxd 2005/12/09 11:30 page 93 ocean and on the african east coast to full species status, with c. africanus occurring from coastal somalia to tanzania. brygoo (op. cit.) could not agree without a full assessment of all the populations and retained the subspecies c. boutonii africanus for the coastal populations from mogadishu in somalia to the cabaceira peninsula in mozambique. the populations further south were not known to either of the latter authors. the presence of this most southern population on the african continent was first brought to the attention of science in 1964 by dr ortwin bourquin, then employed as a temporary student helper of the marine turtle surkoedoe 45/1 (2002) 94 issn 0075-6458 fig. 1. cryptoblepharus boutonii in the barnacle zone at black rock. fig. 2. black rock from the air, showing the bay and the sandy channel separating it from the dune forest. vey, who collected some specimens and had them identified by dr. d.g. broadley, of the then umtali museum in rhodesia (= mutare, zimbabwe). after a number of specimens were collected and the basic situation was assessed, it was apparent that a minute population was involved. this was considered a serious conservation problem. accordingly, it was agreed upon by interested individuals in the natal parks board and the transvaal museum not to deplete this small, isolated population any further by unreasonable collecting. it was also decided to monitor this population by an annual census over a number of years. a start was made during february 1978 and, although it was impossible for the author to visit black rock annually, it was visited 11 times until march 1992. in october 2001, a follow-up visit was made to assess the current status after a period of nearly ten summer cycles. habitat on the generally sandy maputaland coastline occasional rocky outcrops, consisting of consolidated, fossilised, ancient dune deposits (dr. a. kaiser pers. comm.), occur. it is probably also of pleistocene origin, with physical, faunal and botanical aspects very similar to those described for cabo inhaca on inhaca island (macnae & kalk 1958). black rock is one of these, forming a prominent cliff and the tip of a short peninsula at the southern end of a shallow bay with a wide sweeping sandy beach (fig.2). the beach is edged by typical low vegetation, which quickly haacke.qxd 2005/12/09 11:30 page 94 changes into wind-trimmed dense coastal forest covering the dune ridge parallel to the coast. this beach vegetation reaches the top edge of the rocks. a broad sandy gully, probably formed by some exceptional storm tides in the past and maintained by previous vehicular traffic and prevailing winds, between the base of the high dune and the cliff, left an elevated vegetation island on the high ground of the peninsula behind the cliff. about a hundred metres south of the above ‘island’, higher up on the beach and only rarely reached by exceptionally high tides, lies a similar, but much smaller, island. no snake-eyed skinks have ever been noticed there. the rocky cliff rests on a broad shelf which becomes exposed during low tides. on the inside of the bay the rocky habitat starts with the presence of loose lying rocks which, according to season and past weather conditions, may be covered in sand or more or less exposed. towards the ne the loose rocks are backed by a continuous rock bank, increasing in thickness and overlain by vegetated white sand. from the most southern point of the bay these rocks lie in a north-eastern direction for about 30 m, before the cliff swings northwards. from this point the cliff consists of jagged, porous, exposed rock without sand cover or vegetation. this bare 25 m point of the peninsula rises to about 4 m above the base shelf, which is about a metre below mean high water mark and the oyster zone. a mushroom-like, free-standing rock forms the most northern point. from there the eastern, seaward edge of black rock runs in a generally southern direction for about 50 m and then swings into a southwesterly direction, terminating in beach sand about 200 m away. the more or less vertical cliff face is often deeply undercut and in places reaches 8–10 m in height. the bottom shelf is only negotiable during low and preferably spring low tide. this is essential as the lizards descend from their roosting sites into the intertidal zone to feed, which is the optimum exposure for counting and thus establishing the population size. as mentioned before, the rock surface is extremely leached and weathered resulting in a viciously rough surface which provides ample retreat for the tiny skinks. at several points on this rock outcrop groups of pecuissn 0075-6458 95 koedoe 45/1 (2002) fig. 3. working sketch of black rock with arbitrary names often indicating concentration areas of lizards. haacke.qxd 2005/12/09 11:31 page 95 liar, circular, vertical, slightly conical holes occur, with an upper diameter of up to about one metre. they are the result of chemical leaching and subsequent erosion. many are partially filled with sand and in some this filling appears to have consolidated and even fossilized. a considerable number extend 3–5 m down and some have actually broken through into the undercut in the base of the cliff face. similar situations along the mozambique coast result in well-developed ‘blow holes’. in the situation at black rock, these shafts provide additional shelter and vertical migration routes from the high tide and night retreats at the top of the cliff to the low tide feeding grounds in the oyster-barnacle zone. koedoe 45/1 (2002) 96 issn 0075-6458 fig. 4. black rock bay at low tide, showing ‘hole rock’ and adjacent terrain. fig. 5. the northern 'tip' of black rock showing the extremely eroded and jagged rock type. the extreme porosity of the northwest-facing rocks facing the bay and the northern tip of the peninsula appears to be favoured by the lizards, as several concentration points exist. the greater part of the main seawardfacing high cliffs, although roughly eroded, is less porous and therefore appears less suited as retreats since fewer skinks were observed there. in a number of places accumulations of large boulders lying against the base of the cliff face provide protected feeding areas and consequently have resident groups of skinks. the most important sites were named and indicated on the sketch plan (fig. 3) on which sightings were recorded. starting in the bay amongst the smaller boulders partially covered in sand, the significant groups of rocks or sites are: a) ‘hole rock’, a small cluster of very leached rocks lying on sand leading up to a slightly isolated, but dominant boulder about 2 m high with a big, cloverleaf shaped diagonal hole through it, which is much favoured by the skinks (fig.4.). b) ‘bay block’, a large block which broke off the main cliff and settled next to it and a few smaller ones which form a bridge for the lizards. c) ‘the tip’, the most northern tip of the peninsula, with lots of cavities, recesses and rock pools, used by ‘rock-hopper’ fish (blenniidae) (day 1968), providing safe feeding areas for lizards (fig. 5). southwards, the first important site is: d) ‘tumbled rocks’, where a group of large rocks provide access to the barnacleoyster zone with safe retreats and a cliff face top with night retreats. these boulders lie in front of e) ‘the hole’, where the roof of an undercut collapsed, provides access to the very jagged cliff-top. ‘the hole’ is surrounded by vertical leach holes. f) ‘big block’. a big section of the cliff face broke off and settled on the bottom shelf with a slight tilt. the sheltered, haacke.qxd 2005/12/09 11:31 page 96 south is the small island, described above. black rock appears to act as an interference to the sea current, carrying debris from the north, as on several occasions huge logs were dumped on the beach. some were of a type, with nooks, crannies and borer holes, which one could imagine to have acted as potential rafts for the little skinks. census methods at high tide only the top of the cliff is accessible to humans. at low tide, preferably spring low, once the base shelf becomes exposed, one can walk along the bottom edge of the cliff in the barnacle-oyster zone, which, on sunny days is the main feeding area for the lizards. returning along the top edge of the cliff, those lizards which had remained there, may be counted. initially repeated rounds were made from very early in the morning until late afternoon, but it soon became apparent that the prime feeding time shifts with the tides and time of the year and on a sunny day or with only a hazy sun may be from about 08:40 to 11:30. weather and tide conditions, combined with wave action and human activity, can influence the results significantly. as a consequence it was found that a single thorough count per day during midmorning, repeated over four or five days around spring tide, reduces the effect of poor weather conditions or human presence and allows the figures obtained by two or more people working as a team to be acceptable. issn 0075-6458 97 koedoe 45/1 (2002) narrow gap between the cliff face and the ‘big block’ attracts lizards. g) ‘the crack’. the cliff face developed a massive crack, where another huge block will part from the rock face to settle next to it. it is partly opened and is used as a safe vertical passage or for basking in the top section. h) ‘bone point’ is the most easterly projection, marked by the exposure of sub-fossil buffalo teeth (id. mrs e. voigt pers. comm.) and some bone splinters. this point has no particular attraction for the lizards, but from here the direction of the cliff swings to the south-west, often facing rough seas and strong wind, making it less attractive to the lizards. i) ‘big cave’. the roof of a large undercut, caused by wave erosion, collapsed as it was weakened by deep, vertical, leaching holes. the remaining leaching holes surrounding the collapsed ceiling are used by the skinks as sheltered living areas. these lizards will not normally descend here into the oyster-barnacle zone as the descent is either exposed or overhanging and in the dark. j) ‘hell’s gate’. of no consequence to the lizards, but it is a gully which observers have to cross by climbing over. it may suddenly fill up by wave action and thereby endanger the safety of the team (fig. 6.). it can only be crossed safely during low, preferably spring low tides. k) ‘raised platform’. a beautiful raised open air aquarium, which indicates the northern beginning of the l) ‘high wall’. a near vertical, very jaggedly eroded cliff face, mostly avoided by the lizards, varying in height from 8–10 m and extending for 40–50 m as far as the k) ‘southern rocks’, which consist of an accumulation of large boulders extending due east for about 15 m. this landmark is much favoured by the skinks. after the last landmark the rock face becomes lower and lower, only occasionally visited by individual lizards, and eventually ends in beach sand. about 100 m further fig. 6. the lower shelf of black rock at low tide, showing the oyster-barnacle zone near the entrance of the 'big cave' on the right, two observers crossing the 'hell's gate' channel, with the 'raised platform' in the back. haacke.qxd 2005/12/09 11:31 page 97 by repeating these census rounds for a few days in succession, it became obvious where concentrations of lizards occur and that individuals stay more or less within a limited home range. this was confirmed when individuals were marked with a red paint spot on the back and individuals with broken tails, as well as juveniles, were recorded. dispersal was quite slow and in principle individuals remained more or less in the same area where first found during a visit. it was also noted that according to circumstances not all lizards are active simultaneously. after each round the ‘total number counted’ was established by adding up all the lizards recorded during one round and the ‘counted maximum’ was established at the end of a visit by picking the highest count of all the results. an ‘estimated maximum’ was also established at the end of each visit to black rock (table 1.). out of the set of four or five counts made during one visit the highest number of lizards per concentration point was selected and added. this figure was accepted as representing the actual total population as closely as it could be established.accordingly indications of population fluctuations can be established. the means of the ‘counted maxima’ and the ‘estimated maxima’ were calculated to indicate the difference between these two parameters. results due to the rugged terrain at black rock it is impossible to do exact counts. by repeating counts during the optimum activity period and by adding up maximum counts at various concentration points including individual vagrants, an estimated population size present during a visit can be established. the range of the ‘counted maximum’ has varied between 28 (feb. 1978) and 83 (mar. 1989), for which seasons the ‘estimated maxima’ were 47 and 106 (dec. 1980 = 112). this suggests a fluctuation in excess of 100 % (range = 47–112) over 15 seasons, with a mean ‘counted maximum’ of 58 (= 57.55) and mean ‘estimated maximum’ of 80 (=79.8). a follow-up check during the first week in october 2001, nearly ten seasons later, provided the very satisfactory results, viz., highest ‘counted maximum’ = 99 and the highest ‘estimated maximum’ = 113, suggesting a generally very positive situation (table 1.). koedoe 45/1 (2002) 98 issn 0075-6458 table 1 population variation of cryptoblepharus boutonii at black rock, kwazulu/natal season counted estimated maximum maximum feb. 1978 28 47 dec. 1978 44 60 apr. 1980 50 63 dec. 1980 75 112 dec. 1981 60 7 9 dec. 1984 41 65 nov. 1986 60 85 mar. 1989 83 106 feb. 1990 66 87 apr. 1991 63 94 feb. 1992 63 81 range 28-83 47-112 n = 11 x = 57. x = 79.8 check oct. 2001 99 113 various questions arise when considering the survival potential of such a small, isolated population of a lizard which reproduces by laying a single, soft-shelled egg (rarely 2) at a time (fricke 1970; own records). biological observations have been made (haacke 1977) during these visits and will be reported on in greater detail elsewhere. in short, these skinks occupy a niche which is ideally suited for their small, thin bodies. their colour matches the dark background and the availability of retreats protects against potential predators and inclement weather. the intertidal zone is without reptile competition. larger lizards, such as mabuya depressa, which may threaten mainly juveniles, occur in the vegetated zone adjacent to the bare rocks and will restrict possible expansion. a single juvenile white-throated monitor (varanus albigularis) was observed during one visit basking in front of a hole on the cliff face above ‘tumbled rocks’. this lizard could be a serious threat, but its presence was apparently only of a temporary nature. on ‘tumbled rocks’, we found a single shed skin and a complete young specimen of a short-snouted haacke.qxd 2005/12/09 11:31 page 98 sand snake psammophis brevirostris which appeared to have been killed and dropped by a bird. hardly any sea-birds were ever noticed in the area. predation by shorebirds in the pacific has been reported (clapp 1967), but has not been confirmed here and it is unlikely, as the local terrain is unsuited for plovers. being tiny and non-spectacular, these lizards are safe from collectors and the pet trade and unless some mischievous persons destroy specimens, people do not appear to be a threat. conclusion under the present conditions, the tiny population of cryptoblepharus boutonii cf. africanus on black rock maintained a population mean of 80 individuals over 15 seasons and consisted of a considerably higher figure when checked ten seasons later. from a conservation point of view this is a commendable situation, and if the conditions controlling access and utilisation can be maintained, no additional measures seem to be necessary to maintain this quaint population of tiny skinks. acknowledgements i am indebted to the various authorities who were responsible for issuing permits to conduct these surveys over the years. these were the director-general for co-operation and development, the regional forester at mbazwana, the director of the bureau of natural resources, kwazulu administration and lately mr. t. ferguson, ranger in charge, department of environment and tourism, kwazulu-natal. in the area concerned, field staff and rangers of the various authorities over many years, too many people to be mentioned individually, gave advice and rendered assistance. a number of people assisted in the counting procedure and without their assistance it would have been a futile exercise. mrs lomi brown, collection manager, herpetology, at that time and my children karl and ingrid were part of the team for a number of years. mrs elizabeth voigt, then paleontologist of the transvaal museum and later on two occasions hans and monika kubierske, assisted with great enthusiasm. on the final visit the team consisted of dr niels jacobsen, dr annémarie van der walt and mr mirko barts. many thanks to all of them. references bruton, m.n. & w.d. haacke. 1980. the reptiles of maputaland. pp. 251–287. in: bruton, m.n. & k.h. cooper (eds.). studies on the ecology of maputaland. cape town: rhodes university and the natal branch of the wildlife society of southern africa. brygoo, e.r. 1986. systématique des lézards de scincidés de la région malgache xviii. les cryptoblepharus. bulletin museum naturel paris, 4e sér., 8. 1986, section a, no 3: 643–690. clapp, r.b. & g.m. tilger. 1967. predation on snake-eyed skink (ablepharus boutonii) by two pacific shorebirds. herpetologica 23(1): 75. day, j.h. 1968. a guide to marine life on south african shores. cape town: a.a. balkema. fricke, h.w. 1970. die ökologische spezialisierung der eidechse cryptoblepharus boutonii* cognatus (boettger) auf das leben in der gezeitenzone (reptilia, skinkidae). oecologia (berl.) 5: 380–391. fuhn, i. e., 1970. contribution à la systématique des lygosominae africain (reptilia, scincidae). i. les espèces attribuées au genre ablepharus. revue roumaine de biologie (zoologie) 15 (6): 379–393. haacke, w.d. 1977. it floated its way across the world—and any scientist with sense leaves it alone! african wildlife 31(1): 30–31. haacke, w.d. 1988. bouton’s skink. pp. 75–76. in: branch w.r. (ed.). south african red data book —reptiles and amphibians. pretoria: council for scientific and industrial research, national scientific programmes unit. (south african national scientific programmes report; no. 151). macnae, w. & m. kalk. 1958. a natural history of inhaca island, mozambique. johannesburg: witwatersrand university press. mclachlan, g.r. 1978. south african red data book —reptiles and amphibians. pretoria: council for scientific and industrial research, national scientific programmes unit. (south african national scientific programmes report; no. 23). mertens, r. 1931. ablepharus boutonii (desjardin) und seine geographische variation. zoologische jahrbücher 61(1 & 2): 63–210. mertens, r. 1933. weitere mitteilungen über die rassen von ablepharus boutonii. zoologischer anzeiger 105: 92–96. mertens, r. 1958. neue eidechsen aus australien. senckenbergiana biologica 39 (1/2): 51–56 issn 0075-6458 99 koedoe 45/1 (2002) haacke.qxd 2005/12/09 11:31 page 99 mertens, r. 1963. weitere mitteilungen über die rassen von ablepharus boutonii, iii. zoologischer anzeiger 173 (2):99–110. spawls, s., k. howell, r. drewes, j. ashe. 2002. a field guide to the reptiles of east africa. london: a p natural world. welch, k.r.g. 1982. herpetology of africa: a checklist and bibliography of the orders amphisbaenia, sauria, and serpentes. malabar, florida: krieger. wilson, k. w. & d. g. knowles 1988. australia’s reptiles. australia: collins. koedoe 45/1 (2002) 100 issn 0075-6458 haacke.qxd 2005/12/09 11:31 page 100 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice about the author(s) leslie r. brown applied behavioural ecology & ecosystem research unit, university of south africa, florida, south africa andri c. van aardt department of plant sciences, faculty of natural & agricultural sciences, university of the free state, bloemfontein, south africa beanelri b. janecke department of animal, wildlife & grassland sciences, faculty of natural & agricultural sciences, university of the free state, bloemfontein, south africa citation brown, l.r., van aardt, a.c., janecke, b.b. 2020, ‘a tribute to pieter johannes (johann) du preez’, koedoe 62(2), a1640. https://doi.org/10.4102/koedoe.v62i2.1640 in memorial: pieter johannes (johann) du preez note: special issue: connections between abiotic and biotic components of a granite catena ecosystem in kruger national park, sub-edited by beanelri janecke and johan van tol. tribute a tribute to pieter johannes (johann) du preez leslie r. brown, andri c. van aardt, beanelri b. janecke copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. pieter johannes du preez passed away on the evening of 29 december 2019 in hermanus after a short fight against cancer. johann as he was known to everyone, was well educated in ecology and had a passion for nature. his knowledge of the environment and willingness to share that with colleagues, students and the general public is how he will be remembered. johann’s passing came as a huge shock to all with whom he has worked and who has known him. he has left a huge gap within the scientific world, especially within the field of vegetation science where he was well known and regarded as one of the top plant ecologists in the country. he will be remembered for his endless passion to study nature and obtain more knowledge on the functioning of ecosystems. his love for nature and conservation could be felt in his presence. he was modest, humble, understanding, supportive and always willing to walk that extra mile for the people that crossed his path. as an academic he influenced the lives of several students who became successful vegetation scientists and researchers under his supervision. he also contributed to the literature in various fields of ecology mostly specialising in the mapping of vegetation and investigating various ecological community compositions. many of the valuable plant samples that he collected are housed at the geo potts herbarium at the university of the free state (ufs). he is well-published and has contributed numerous publications, book chapters, conference presentations and technical reports on the vegetation of southern africa. he produced a detailed vegetation map of the free state province and was co-author of the widely acclaimed ecology book life and the environment: an african perspective for which he received the golden merit award from the south african academy for science and arts for his contribution to science. johann collaborated on the vegetation surveys of the stevenson hamilton research supersite near skukuza in the kruger national park with colleagues from the university of the free state. here he contributed towards our understanding of the interaction between vegetation and the environmental factors that influenced the various plant communities in the area. he was one of the main researchers of this multidisciplinary project and contributed to much of the research published in this special issue. at the time of his passing, johann was an active research fellow at the department of plant sciences, ufs, where he was still involved in supervising students and playing his part in research with colleagues from other south african universities. a day in the field with johann was equivalent to a long time in the classroom. he was cited by many people for his many achievements. johann played a quiet, yet profound role in the advancement of vegetation science in southern africa and we will remember him for his huge contribution not only in terms of scientific knowledge, but his energetic and positive approach to life. this is a tribute to honour prof. johann du preez, and to show our respect, gratitude and admiration when we remember the huge contribution that he has made to the research published in this special issue. – l. brown, a. van aardt, b. janecke and the research team (a special thanks to prof. leslie brown from unisa for his contribution to this tribute). introduction materials and methods results discussion conclusion acknowledgements references about the author(s) marieka gryzenhout department of genetics, faculty of natural and agricultural science, university of the free state, bloemfontein, south africa marcele vermeulen department of microbial biochemical and food biotechnology, faculty of natural and agricultural science, university of the free state, bloemfontein, south africa gilmore pambuka department of genetics, faculty of natural and agricultural science, university of the free state, bloemfontein, south africa riana jacobs department of mycology unit, plant health and protection, agricultural research council, pretoria, south africa citation gryzenhout, m., vermeulen, m., pambuka, g. & jacobs, r., 2020, ‘first report of various fusarium species from the stevenson-hamilton supersite granite catena system in the kruger national park, south africa’, koedoe 62(2), a1599. https://doi.org/10.4102/koedoe.v62i2.1599 note: special issue: connections between abiotic and biotic components of a granite catena ecosystem in kruger national park, sub-edited by beanelri janecke and johan van tol. short communication first report of various fusarium species from the stevenson-hamilton supersite granite catena system in the kruger national park, south africa marieka gryzenhout, marcele vermeulen, gilmore pambuka, riana jacobs received: 28 sept. 2019; accepted: 19 apr. 2020; published: 29 oct. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. keywords: fusarium; neocosmospora; kruger national park; topsoil; rhizosphere; catena. introduction the kruger national park (knp) covers the north-eastern part of southern africa (carruthers 2017) and is also linked with the gonarezhou national park (zimbabwe) and the limpopo national park (mozambique) as the great limpopo transfrontier park. the knp is part of the kruger to canyons biosphere area designated by the united nations educational, scientific and cultural organization (unesco) as an international man and biosphere reserve (the ‘biosphere’) (http://www.unesco.org/new/en/natural-sciences/environment/ecological-sciences/biosphere-reserves/africa/south-africa/kruger-to-canyons/). the stevenson-hamilton supersite, where this study was conducted, is part of four research ‘supersites’ in the knp, with each representing distinct geological, climatic and linked biodiversity patterns (smit et al. 2013). the foundational biological information regarding soil biota in south africa was recently assessed, and it included soil fungi (janion-scheepers et al. 2016). these authors reported that despite south africa being only 0.8% of the earth’s terrestrial area, it contains nearly 1.8% of the world’s described soil species. areas such as the nama-karoo, northern cape and eastern cape are undersampled for most taxa as well as natural soils in biodiversity hotspots. similarly, the knp with its diverse ecosystems is not well explored. the fungal genus fusarium has a cosmopolitan distribution and includes a vast number of species. these species are commonly recovered from a variety of substrates including soil, air, water and decaying plant materials (leslie & summerell 2006). they have diverse ecosystem functions in soils and are also able to colonise living tissues of plants and animals, including humans, acting as endophytes (microbial organisms existing inside plant tissues), secondary invaders or becoming devastating plant pathogens (nelson, dignani & anaissie 1994). in addition to their ability to colonise a multiplicity of habitats, fusarium species are present in almost any ecosystem in the world (leslie & summerell 2006). a number of genera representing previously known fusarium species were established based on deoxyribonucleic acid (dna) sequence data (lombard et al. 2015). for instance, the fusarium solani species complex (fssc) was proposed to be the genus neocosmospora (lombard et al. 2015). however, because of the close association with the name fusarium and the fact that these names serve a large community of end-users, that is, plant pathologists, quarantine officers, veterinarians and medical practitioners, a different system was proposed where the name fusarium was kept, for instance, for the fssc (geiser et al. 2013). the resulting confusion is evident as a number of new species in the complex kept the name of fusarium, for example, f. euwallaceae (freeman et al. 2014), which is the pathogenic fungus associated with the devastating polyphagous shothole borer. these taxa are usually included in fusarium research. a multidisciplinary study was conducted in the knp to study the structure and biodiversity, and the various possible biotic and abiotic interactions of a catena or hill slope ecosystem on the stevenson-hamilton supersite (25°06’28.6s, 31°34’41.9e and 25°06’25.7s, 31°34’33.7e). the aim of this study was to establish a baseline on the species diversity of fusarium occurring at the particular supersite in order to possibly use species in fusarium and closely related genera, which include specialised and generalist species, as a possible focus group to study interactions with the various biotic and abiotic factors in the catena system. previous studies have already described five new species with representative isolates from the knp. these included f. nygamai (burgess & trimboli 1986) and f. fredkrugeri (sandoval-denis et al. 2018) in the ffsc, f. polyphialidicum (marasas et al. 1986) in the fusarium concolor species complex (fcosc), f. convolutans in the fusarium buharicum species complex (fbsc) and f. transvaalense in the fusarium sambucinum species complex (fsasc) from soils (sandoval-denis et al. 2018). jacobs-venter et al. (2018a) separated f. polyphialidicum strains into three species, namely f. concolor, f. babinda and f. austroafricanum, and confirmed that f. polyphialidicum is synonymous with f. concolor. f. fredkrugeri, f. convolutans and f. transvaalense originated from the stevenson-hamilton supersite. in this study, soil and rhizosphere samples from various plants in the stevenson-hamilton supersite, which has a distinct geology and hydrology, were obtained. isolations from these samples revealed a large collection of fusarium isolates. the aim of this study was to identify these fusaria. as information on microbes, including fungi, is scarce for the knp, and basically non-existent for the supersite, the study will contribute pioneering and invaluable biodiversity data on these ill-studied organisms that will be informative and useful for the management and conservation of the knp. materials and methods sampling the study was conducted in the southern granites ‘supersite’ catena close to the stevenson-hamilton memorial (smit et al. 2013). four random soil samples to a depth of 5 cm were taken for each of the components of the catena system in a transect of more or less 500 m following theron, van aardt and du preez (2020). furthermore, roots of pogonarthria squarrosa (sickle grass, poaceae, poales), sporobolus nitens (curly leaved drop seed grass, poaceae) and schkuhria pinnata (dwarf marigold, asteraceae, asterales), which included some of the dominant plants in the catena (theron et al. 2020), were collected. topsoil was deliberately included with the assumption that the soils will contain soil-associated fusaria as well as spores that were aerially distributed from plants in the area. the soils were transported cold to the laboratory, where soil dilution series were made on rose bengal–glycerine–urea medium (leslie & summerell 2006) and 20% potato dextrose agar (pda; biolab, merck, south africa). roots were suspended in sterile water and shaken, and the soil solution was then used in a dilution series. colonies resembling the cultural morphotypes of fusarium species were purified from the primary plates by making single spore cultures from colonies on sna medium (leslie & summerell 2011). cultures were deposited in the national collection of fungi (prem), biosystematics division, agricultural research council (arc), pretoria, south africa (table 1). table 1: list of fusarium isolates identified in this study that originated from the stevenson-hamilton catena in kruger national park, south africa. table 1 (continues...): list of fusarium isolates identified in this study that originated from the stevenson-hamilton catena in kruger national park, south africa. deoxyribonucleic acid sequence-based characterisation inqaba biotec (pretoria, south africa) extracted dna from the scraped mycelium of 1-week-old cultures grown on pda, and the translation elongation factor 1α gene region (tef1α) was amplified and sequenced using primers ef1 and ef2 (o’donnell et al. 2008). sequences obtained were viewed and edited with geneious 7.1.9 (biomatter, auckland). sequences were grouped into fusarium species complexes using a skeleton sequence data set representing all species complexes in fusarium and genera previously named as fusarium, as well as species grouping outside species complexes (data not shown). after the appropriate complex or closest related species has been identified, sequences were included in separate dna data sets representing all known and vouchered sequences of the particular group or complex. all alignments were performed in mafft 7.0 (http://mafft.cbrc.jp/alignment/software/) with the l-ins-i option selected (katoh et al. 2005). the alignments were corrected manually where needed. representative sequences with a high identity to the ffsc were aligned with all currently recognised species and phylogenetic lineages in the ffsc (edwards et al. 2016; geiser et al. 2013; herron et al. 2015; moroti et al. 2016). similarly representative sequences with a high identity to the fusarium chlamydosporum species complex (fcsc) (lombard et al. 2019a; o’donnell et al. 2009b) and fusarium oxysporum species complex (fosc) (laurence et al. 2014; lombard et al. 2019b; o’donnell et al. 2009a) were aligned in data sets linked to the listed references. sequences of the fssc (o’donnell et al. 2008) that are now known as neocosmospora (lombard et al. 2015) were also used. maximum likelihood analyses were conducted in mega v. 7 using the models assigned to each data set and with a 1000 bootstrap replicates to determine the support of the branches. ethical considerations ethical approval for the multidisciplinary project as a whole was obtained from the interfaculty animal ethics committee at the university of the free state (ufs-aed2019/0121). sanparks permit numbers for collection of soil for lab analyses and vegetation for identification purposes are sk069, sk2095 and sk054. results deoxyribonucleic acid sequence-based characterisation isolates (109) characterised in this study from the catena system represented four species complexes, namely ffsc, fcsc, fssc and fosc, and originated from the rhizospheres of all three plants and the topsoil (table 1). each of these complexes includes a diversity of species. in the ffsc, isolate ppri 20296 was identified as f. proliferatum (bootstrap support 98%), and isolates ppri 20281 and ppri 201306 were identified as f. nygamai (bootstrap support 97%) (figure 1). isolates ppri 20610, ppri 19535 and ppri 19537 were grouped in the clade of n. vasinfecta (figure 2) in the fssc (bootstrap support of 97%) and grouped into two haplotype groups. isolates from the knp that were grouped in the fcsc (table 1) constituted a very large number of isolates that did not group with any of the previously known lineages or newly described species (figure 3). between-isolate variation seven haplotypes was seen that could be indicative of more possible cryptic species or significant population structure. based on the tef sequence data alone, all of the novel species described (lombard et al. 2019a) in the fosc could not be resolved but isolates (table 1) formed four haplotypes that grouped together with f. callistephi and f. fabacearum, and isolates from australia (figure 4). figure 1: unrooted maximum likelihood phylogram of all currently sequenced fusarium species in the fusarium fujikuroi species complex based on translation elongation factor 1-a gene sequences. bootstrap support values higher than 80% are shown on the branches. the evolutionary model applied to the analysis is indicated on the figure. isolates from this study are indicated with ppri numbers. figure 2: unrooted maximum likelihood phylogram of all currently sequenced fusarium species in the fusarium solani species complex (also referred to as neocosmospora with some species named as neocosmospora) based on translation elongation factor 1-a gene sequences. bootstrap support values higher than 80% are shown on the branches. the evolutionary model applied to the analysis is indicated on the figure. isolates from this study are indicated with ppri numbers. figure 3: unrooted maximum likelihood phylogram of all currently sequenced fusarium species in the fusarium chlamydosporum species complex based on translation elongation factor 1-a gene sequences. bootstrap support values higher than 80% are shown on the branches. the evolutionary model applied to the analysis is indicated on the figure. isolates from this study are indicated with ppri numbers. figure 4: unrooted maximum likelihood phylogram of all currently sequenced fusarium species in the fusarium oxysporum species complex based on translation elongation factor 1-a gene sequences. bootstrap support values higher than 80% are shown on the branches. the evolutionary model applied to the analysis is indicated on the figure. isolates from this study are indicated with ppri numbers. discussion this study represents the first report of f. proliferatum (ffsc), n. vasinfectum that was previously known as f. cosmosporiellum in the fssc (geiser et al. 2013) and f. oxysporum sensu lato (fosc) from soils in the stevenson-hamilton granite supersite in the knp. possible new species in the fcsc were also detected. the presence of f. nygamai (ffsc) was confirmed. together with other species previously described from the knp (f. fredkrugeri, f. convalutum, f. transvaalense and f. concolor as f. polyphialidicum), there are thus at least nine fusarium species present in the knp. fusarium nygamai, f. proliferatum, f. oxysporum sensu lato, f. chlamydosporum, n. vasinfectum and f. concolor are species that have a world-wide occurrence, including south africa (jacobs et al. 2018a; leslie & summerell 2006). they are associated with various plant hosts as well as soils and can also produce mycotoxins in food commodities or be associated with diseases of animals or humans (leslie & summerell 2006). the new species f. fredkrugeri, f. convalutum and f. transvaalense have only recently been described and have most likely not yet been discovered elsewhere. because these species are generalists that can be isolated from various substrates and plant hosts, these species most likely are not suitable to represent a target group to study unique species associations within a catena system. the majority of strains (90) obtained from the knp sample sites belonged to f. chlamydosporum species complex. a four-locus typing scheme (o’donnell et al. 2009b) revealed mlst and species within the species complex, and lombard et al. (2019a) recently published the description of numerous new species in the complex. isolates obtained from this study appear to represent new species. as before, a number of new species from the knp are yet to be described. what is notable is that several undescribed fusarium species have been discovered in the knp. previously, f. nygamai, f. polyphialidicum, f. fredkrugeri, f. convalutum and f. transvaalense were described from the knp, while possible new species have also been characterised in this study. since their description, f. nygamai and f. concolor (also as the synonym f. polyphialidicum) have been discovered from across the world (leslie & summerell 2006), suggesting a wide substrate, host and geographical range despite being first described from a national conservation park in south africa. the number of undescribed species of fusarium in the knp is not surprising because the biodiversity of fusarium and closely allied genera that were previously called fusarium is largely untouched. this is especially so in pristine natural areas (jacobs et al. 2018b), because most fusarium research in south africa is focused on agricultural problems or animal and human health issues caused by fusarium species. conclusion the knp plays an important role in not only protecting the native ecosystems present in that area and the animals and plants they contain but also protecting fusarium species that occur in south africa, of which some are new to science. the ecological roles of these species in numerous ecosystems are, however, still unknown, and further studies on their impact on ecosystem services and function must be pursued. such studies are important because gryzenhout et al. (2020) showed through environmental sequencing that fusarium species are one of the dominant groups found within the soil-plant root zones of plants occurring in the stevenson-hamilton granite supersite. further sequencing of additional genes, as what has been done in this study, will provide a better estimation on species level of the species that could be involved. acknowledgements the authors thank the university of the free state strategic research fund for providing funding for this research, sanparks scientific services for their assistance during field sampling, dr beanelri janecke for her leadership in the project and the rest of the research team for their insights. the authors are grateful to mrs grace kwanda (arc, pretoria) for her patience and assistance during the submission of the fungal cultures to the national collection of fungi. mr e. theron and profs. johan du preez and piet le roux (ufs) are thanked for the provision of soil and plant samples. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions the authors directly participated in the study design, execution and interpretation of the research. all authors contributed equally to this research work. funding information this study was funded by the university of the free state under the ‘multi-disciplinary program’. data availability data are available from the corresponding author on request. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliate agency of the authors. references burgess, l.w. & trimboli, d., 1986, ‘characterization and distribution of fusarium nygamai, sp. nov.’, mycologia 78(2), 223–229. https://doi.org/10.1080/00275514.1986.12025233 carruthers, j., 2017, national park science: a century of research in south africa, cambridge university press, cambridge. edwards, j., auer, d., de alwis, sk., summerell, b., aoki, t., proctor, r.h. et al., 2016, ‘fusarium agapanthi sp. nov, a novel bikaverin and fusarubin-producing leaf and stem spot pathogen of agapanthus praecox (african lily) from australia and italy’, mycologia 15(2), 333. https://doi.org/10.3852/15-333 freeman, s., sharon, m., maymon, m., mendel, z., protasov, a., aoki, t. et al., 2014, ‘fusarium euwallaceae sp. nov. – a symbiotic fungus of euwallacea sp., an invasive ambrosia beetle in israel and california’, mycologia 105(6), 1595–1606. https://doi.org/10.3852/13-066 geiser, d.m., aoki, t., bacon, c.e., baker, s.e., bhattacharyya, m.k., brandt, m.e. et al., 2013, ‘one fungus, one name: defining the genus fusarium in a scientifically robust way that preserves longstanding use’, phytopathology 103(5), 400–408. https://doi.org/10.1094/phyto-07-12-0150-le gryzenhout, m., cason, e.d., vermeulen, m., kloppers, g.a.e., bailey, b. & ghosh, s., 2020, ‘fungal community structure variability between the root rhizosphere and endosphere in a granite catena system in the kruger national park, south africa’, koedoe 62(2), a1597. https://doi.org/10.4102/koedoe.v62i2.1597 herron, d.a., wingfield, m.j., wingfield, b.d., rodas, c.a., marincowitz, s. & steenkamp, e.t., 2015, ‘novel taxa in the fusarium fujikuroi species complex from pinus spp.’, studies in mycology 80, 131–150. https://doi.org/10.1016/j.simyco.2014.12.001 jacobs, a., laraba, i., geiser, d.m., busman, m., vaughan, m.m. & proctor, r.h., 2018a, ‘molecular systematics of two sister clades, the fusarium concolor and f. babinda species complexes, and the discovery of a novel microcycle macroconidium–producing species from south africa’, mycologia 110(6), 1189–1204. https://doi.org/10.1080/00275514.2018.1526619 jacobs, a., mojela, l., summerell, b. & venter, e., 2018b, ‘characterisation of members of the fusarium incarnatum-equiseti species complex from undisturbed soils in south africa’, antonie van leeuwenhoek 111, 1999–2008. https://doi.org/10.1007/s10482-018-1093-x janion-scheepers, c., measey, j., braschler, b., chown, s.l., coetzee, l., colville, j.f. et al., 2016, ‘soil biota in a megadiverse country: current knowledge and future research directions in south africa’, pedobiologia 59(3), 129–174. https://doi.org/10.1016/j.pedobi.2016.03.004 katoh, k., kuma, k., toh, h. & miyata, t., 2005, ‘mafft version 5: improvement in accuracy of multiple sequence alignment’, nucleic acids research 33(2), 511–518. https://doi.org/10.1093/nar/gki198 laurence, m.h., summerell, b.a., burgess, l.w. & liew, e.c.y, 2014, ‘genealogical concordance phylogenetic species recognition in the fusarium oxysporum species complex’, fungal biology 118(4), 374–384. https://doi.org/10.1016/j.funbio.2014.02.002 leslie, j.f. & summerell, b.a., 2006, the fusarium laboratory manual, blackwell publications, ames. lombard, l., van der merwe, n.a. & groenewald, j.z., 2015, ‘generic concepts in nectriaceae’, studies in mycology 80, 189–245. https://doi.org/10.1016/j.simyco.2014.12.002 lombard, l.a., van doorn, r. & crous, p.w., 2019a, ‘neotypification of fusarium chlamydosporum – a reappraisal of a clinically important species complex’, fungal systematics and evolution 4(1), 183–204. https://doi.org/10.3114/fuse.2019.04.10 lombard, l., sandoval-denis, m., lamprecht, s.c. & crous, p.w., 2019b, ‘epitypification of fusarium oxysporum – clearing the taxonomic chaos’, persoonia 43, 1–47. https://doi.org/10.3767/persoonia.2019.43.01 marasas, w.f.o., nelson, p.e., tousson, t.a. & van wyk, p.w., 1986, ‘fusarium polyphialidicum, a new species from south africa’, mycologia 78(4), 678–682. https://doi.org/10.1080/00275514.1986.12025306 moroti, r.v., gheorghita, v., al-hatmi, a.m.s., de hoog, g.s., meis, j.f. & netea, m.g., 2016, ‘fusarium ramigenum, a novel human opportunist in a patient with common variable immunodeficiency and cellular immune defects: case report’, bmc infectious diseases 16, 79. https://doi.org/10.1186/s12879-016-1382-9 nelson, p.e., dignani, m.c. & anaissie, e.j., 1994, ‘taxonomy, biology, and clinical aspects of fusarium species’, clinical microbiology reviews 7(4), 479–504. https://doi.org/10.1128/cmr.7.4.479-504.1994 o’donnell, k., gueidan, c., sink, s. & johnston, p.s., 2009a, ‘a two-locus dna sequence database for typing plant and human pathogens within the fusarium oxysporum species complex’, fungal genetics and biology 46(12), 936–948. https://doi.org/10.1016/j.fgb.2009.08.006 o’donnell, k., sutton, d.a., rinaldi, m.g., gueidan, c., crous, p.w. & geiser, d.m., 2009b, ‘novel multilocus sequence typing scheme reveals high genetic diversity of human pathogenic members of the fusarium incarnatum-f. equiseti and f. chlamydosporum species complexes within the united states’, journal of clinical microbiology 47(12), 3851–3861. https://doi.org/10.1128/jcm.01616-09 o’donnell, k., sutton, d.a., fothergill, a., mccarthy, d., rinaldi, m.g., brandt, m.e. et al., 2008, ‘molecular phylogenetic diversity, multilocus haplotype nomenclature, and in vitro antifungal resistance within the fusarium solani species complex’, journal of clinical microbiology 46(8), 2477–2490. https://doi.org/10.1128/jcm.02371-07 sandoval-denis, m., swart, w.j. & crous, p.w., 2018, ‘new fusarium species from the kruger national park, south africa’, mycokeys 34, 63–92. https://doi.org/10.3897/mycokeys.34.25974 smit, i.p.j., riddell, e.s., cullum, c. & petersen, r., 2013, ‘kruger national park research supersites: establishing long-term research sites for cross-disciplinary, multiscaled learning’, koedoe 55(1), 1–7. https://doi.org/10.4102/koedoe.v55i1.1107 theron, e.j., van aardt, a.c. & du preez, p.j., 2020, ‘vegetation distribution along a granite catena, southern kruger national park, south africa’, koedoe 62(2), a1588. https://doi.org/10.4102/koedoe.v62i2.1588 vol. 50 no. 1 pp. 93 98koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research the alien invasive land snail theba pisana in the west coast national park: is there cause for concern? abstract the distribution, abundance, size distribution and diurnal activity patterns of invasive land snails, theba pisana, in the west coast national park (wcnp), south africa, were investigated. the park was divided into 1 km2 grids, within each of which five 1 m2 quadrat counts of live snails were recorded. of 106 grids sampled, 19% contained live snails. the average density of snails was 4.04 m-2 ± 24.9, significantly lower than in disturbed habitats adjacent to the park (57 m-2 ± 96.25), but very high densities were recorded at two sites. snails were most abundant along roadsides and densities decreased dramatically with distance from roads. t. pisana in the wcnp appear to have an annual lifecycle, breeding in autumn to winter and growing to adult size of about 14 mm diameter by the end of the following summer. snails were observed on a wide variety of endemic and introduced plant species and appeared to have a catholic diet. they are active mostly at night and especially during periods of high humidity, irrespective of temperature. given the very high densities that t. pisana can attain at some sites, plus their apparently catholic feeding habits, their potential impact on the vegetation of the park is cause for concern and should be further investigated. control of the main colonies should also be considered. keywords: theba pisana, land snail, invasive species, west coast national park, impact of alien species invasion many human activities, including agriculture, recreation, trade and transport, facilitate the movement of species across natural biogeographic barriers; allowing species that have been separated for millions of years over evolutionary time to interact with one another over ecological time. only a small fraction of such non-indigenous species introduced to an area becomes established and only some of these become invasive. nonetheless, they often have far-reaching impacts, many of which are poorly understood (mooney & cleland 2001) and irreversible (herbert & sirgel 2001). invasive species can also drive local native species to extinction via competitive exclusion (e.g. lach et al. 2002), niche displacement, or hybridisation with related native species (mooney & cleland 2001). therefore, besides their economic ramifications, alien invasions may result in extensive changes in the structure, composition and global distribution of the biota of sites of introduction, leading ultimately to the homogenisation of the world’s fauna and flora (olden et al. 2004) and the loss of biodiversity. the cape floristic region (cfr), the smallest of the six recognised floral kingdoms of the world, is an area of extraordinarily high diversity and endemism (cowling et al. 1996), and is home to more than 9 000 vascular plant species, of which 69% are endemic (goldblatt & manning 2002). much of this diversity is associated with the fynbos biome (turpie et al. 2003), a mediterranean-type, fire-prone shrubland (turpie et al. 2003; van wilgen et al. 2001). the economic worth of fynbos biodiversity, based on harvests of fynbos products (e.g. wildflowers) and eco-tourism, is estimated to be in the region of r77 million a year (turpie et al. 2003). thus, it is clear that the cfr has both economic and intrinsic biological value as a biodiversity hot-spot. while the impacts of alien plants (e.g. alien pines from australia) (richardson 1999) and insects (e.g. the argentine ant, linepithema humile, and spotted stem borer, chilo partellus partellus) (lach et al. 2002) on the fynbos biome are well known, the potential impacts of alien terrestrial snails are poorly understood. theba pisana (müller, 1774), a land snail native to the mediterranean region (baker 1986), was introduced into south africa prior to 1881 (durr 1946), yet has received little attention from researchers, despite the fact that the extent of its invasion in the fynbos biome has been known for some time (macdonald & jarman 1984). t. pisana is a well-known agricultural pest in many parts of the world. in australia, it feeds on a range of agricultural plants (baker 1986, 1989) and uses the stalks of cereals as aestivating sites, in turn clogging machinery and fouling produce during harvests (baker & vogelzang 1988). t. pisana is also a significant pest of citrus, vines, legume crops and cereals in south africa (durr 1946, joubert & walters 1951) and according to quick (1952) has been responsible for the extermination of native snail species as a result of competition for available food. currently, t. pisana is widely distributed in the semi-arid coastal fynbos region of the south western cape (mcquaid et al. 1979), and has been recorded as far east as port elizabeth (pers. obs.), although it has not been successful in establishing viable populations in the eastern cape beyond east london, as this region experiences a significantly different climatic regime to that of the western and southern cape (herbert & kilburn 2004). in their report on the status and potential impacts of alien invasive organisms in the fynbos biome, macdonald and jarman (1984) predicted that t. pisana would have very little impact on ecosystem processes, such as nutrient cycling, energy flow and sediment dynamics or on the germination and succession processes of the plant community itself. however, the snail could seriously impact directly on fynbos plants, by feeding on them, and also displace native herbivores by competing with them for resources, and these effects have not been studied. thus, the collection of reliable data on t. pisana is urgently lizelle j. odendaal tanya m. haupt charles l. griffiths zoology department and centre for invasion biology university of cape town south africa correspondence to: lizelle j. odendaal e-mail: lizelle.odendaal@uct.ac.za postal address: zoology department and centre for invasion biology, university of cape town, rondebosch 7700 93 2008 alien invasive land-snail original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 93 98 required to assess and mitigate the impacts of the snail, and to better conserve the rich floral diversity of the fynbos. this study aims to determine the distribution, abundance and size distribution of t. pisana in the west coast national park (wcnp), a reserve largely dedicated to the conservation of coastal fynbos. we have also made preliminary observations on the daily activity pattern and feeding ecology of t. pisana within this conservation area. results of this study can be used to provide a preliminary assessment of the possible impacts of this invasion within the wcnp. methods study area sampling was conducted between march 2006 and january 2007 in the wcnp (35 000 ha) (17o57’e; 33o10’s). the area is characterised by a mediterranean-type climate with cool, wet winters and hot dry summers (cowling et al. 1996; goldblatt & manning 2002), has an average rainfall of 265 mm and is dominated by strandveld vegetation (west coast national park 2006). data collection to establish the density and distribution of t. pisana within the reserve, the entire study area was divided into 1 km2 grids. the mean density of snails in each grid was calculated from five 1 m2 quadrat counts of live snails recorded at 50 m intervals along a transect line placed within each grid. since t. pisana have mostly been observed close to roads in the wcnp (c.l. griffiths pers. obs.), the first quadrat in each line was taken at a random point (determined by vehicle odometer reading) along the roadside, if a road passed through the grid. if no road crossed the grid the transect line was positioned at random. the densities of t. pisana in 11 random areas in and around langebaan town were also collected in the same manner and compared with the density of snails found within the park. where snails were observed feeding, or grazing damage was visible around snail colonies, the species of plant consumed was noted. to confirm whether snail densities decreased with distance from the road, at three sites of high snail density five 1 m2 quadrat counts were taken along the road verge and five additional samples at successive 10 m distances perpendicular 94 figure 1 live specimen of theba pisana in the wcnp figure 2 map of the wcnp showing the density (m-2) and distribution of theba pisana in august 2006 as well as the position of sampled sites outside of the park original research odendaal, haupt & griffiths koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 93 98 to the road. the average densities of snails, and the position of sampling sites outside the reserve were mapped using arcview gis version 3.3 (esri 2002). population structure was determined by measuring the maximum diameter of at least 150 snails (to the nearest 0.1 mm) at the same site within the park during march, august and october 2006 and january 2007, using dial callipers. to determine the diurnal activity rhythm of t. pisana, samples of 70 to 100 snails, together with the vegetation on which they occurred, were collected within the park and placed in open 20l buckets. after 24 hours of acclimatisation, the percentage of snails observed aestivating and active were recorded every two hours over 24 hours. the temperature (oc) and humidity (%) were also recorded every two hours using a temperaturehygrometer reader. because the study was done over a short period of time, observations of the feeding duration under various weather conditions were not possible. for example, very humid conditions in which these snails are reported to be highly active were not experienced (durr 1946). an artificial humid climate was therefore created by placing a plastic bag pierced with holes (to ensure sufficient ventilation) over one of the buckets, after which the contents were moistened with water using an atomiser every four hours. the experiment was repeated under both dry and humid conditions. statistical analyses statistical analyses were performed using statistica version 7.0 (statsoft inc. 2004). to determine if there were significant differences between the average density of snails within the park and outside the park, nonparametric mannwhitney u-tests were used. results theba pisana was the only alien snail species recorded in the surveys. it can be recognised by its shell colour (figure 1), which ranges from white to yellow-brown with light brown spiral markings (myburgh & rust 1986). the density and distribution of theba pisana within the wcnp is illustrated in figure 2. a total of 106 grids were sampled. two grids had dense populations of t. pisana averaging between 30 and 55 m-2 (figure 2). however, one of these contained mostly very young snails (< 5 mm in diameter), most of which were found in the quadrat closest to the road. the other, at the bird hide, was the most heavily infested area within the park, and quadrat taken along the road contained 300 to 700 snails m-2. the colony extended for approximately 200 m on either side of an old road, and road-side plants in this area were so coated with snails that they appeared white (figure 3). eighteen per cent of the remaining grids contained live snails, all at densities less than 10 snails m-2 (figure 2). interestingly, in many areas where no live t. pisana were found, empty shells were common, attesting to the presence of t. pisana in a much larger proportion of the park in the past. the average density (mean ± sd) of snails outside the park (57 ± 96.25) (table 1) was significantly higher than that within the park (4.04 ± 24.91) (mann-whitney; u = 274, p < 0.05). the densities of snails decreased rapidly with distance from the road (figure 4), the most dramatic case being at site two, where the density decreased from 436 snails m-2 to 5.6 snails m-2 within 20 m and to zero within 50 m of the road side. 95 figure 3 dense population of theba pisana at the bird hide in the wcnp figure 4 average snail densities (m-2) of theba pisana (mean ± sd) along 50 m transects extending perpendicular to the road at four sites within the wcnp (site 1: □, site 2: ∆, site 3: ◊, site 4: ○) site number average snail density (m-2 ) 1 43.8 ± 53.6 2 3 ± 6.7 3 54.2 ± 67.8 4 0 5 15.6 ± 14.5 6 12.8 ± 6.2 7 0 8 0 9 2.2 ± 2.3 10 3.8 ± 4.1 11 0 overall mean ± sd 57 ± 96.25 table 1 mean snail density (m-2 ± sd) at 11 sites adjacent to the wcnp 0 10 20 30 40 50 distance from road (m ) -100 0 100 200 300 400 500 600 700 800 a ve ra ge d en si ty m -2 alien invasive land-snail original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 93 98 table 2 lists species of plants that t. pisana was found aestivating on. damage to these plants on which snails occurred abundantly was quite extensive (pers. obs.). the size distributions of t. pisana during march, august and october 2006 and january 2007 are illustrated in figure 5. in march, most snails were between 13 to 15 mm in diameter. in contrast, the majority of snails in august were much smaller (2 to 7 mm) with very few snails exceeding 10 mm surviving. two months later, the juvenile cohort had reached a maximum diameter of 11 mm (figure 5) and by january 2007 an average diameter of 14 to 15 mm. no snails were active during the day and late evening periods during a sunny day (figure 6a) and only minor activity was noted < 10% in the early morning (0:00 – 06:00), when the humidity was highest (85 to 96%) but the temperature low (4 to 8oc). on a humid day (figure 6b) the snails still remained inactive during the day, but showed high rates of activity during the warm night, peaking shortly after nightfall and before dawn, when over 90% of individuals were active. discussion the primary objective of this study was to map the distribution and density of the alien invasive land snail, theba pisana, in the wcnp, with the aim of evaluating its threat to the vegetation of the area. currently, t. pisana is restricted to only 19% of the study area (figure 2) predominately in areas adjacent to roads and where human activity is highest. however, the presence of large numbers of empty shells in more than 50% of the grid cells sampled indicates that its distribution has been more extensive in the recent past. while the average temperatures within the langebaan region rarely exceed 27 ºc, the average rainfall in the wcnp during the previous two summers has been considerably less than normal (table 3). we conclude that the recent low average rainfall of the region has probably resulted in high mortality rates of snails and that the distribution and densities measured here therefore probably represent minimal 96 figure 5 size frequency distributions of theba pisana in the wcnp in march, august and october 2006 and january 2007 outside scientific name common name family atriplex sp. salt bush chenopodiaceae echium sp. boraginaceae lactuca sp. wild lettuce asteraceae lavatera sp. tree mallow malvaceae euphoria helioscopia umbrella milkweed euphorbiaceae inside euphorbia burmannii steenbokbos euphorbiaceae lycium ferocissimum african boxthorn solanaceae nylandtia spinosa tortoiseberry polygalaceae othonna arborescens bobbejaankool asteraceae othonna cylindrica dikblaar bobbejaankool asteraceae putterlickia sp. celastraceae ruschia sp. mesembryanthemaceae zygophyllum spinosum smalblaar spekbos zygophyllaceae table 2 vegetation species on which theba pisana was found and possibly predated on, both within and adjacent to the wcnp (all species inside the park are indigenous) n o v e m b e r d e c e m b e r ja n u a r y fe b r u a r y m a r c h 1961–1990 temperature (°c) 25 26 28 28 27 precipitation (mm) 12 10 8 4 11 2004–2005 temperature (°c) 24.7 26.6 27.9 27.2 26.5 precipitation (mm) 1.6 1 11.4 0.6 2.8 2005–2006 temperature (°c) 23.9 25 26.8 27.4 25.6 precipitation (mm) 5.2 0 0.2 3.6 2.6 table 3 monthly average temperature (°c) and precipitation (mm) during the hottest months of summer at langebaan over a 30-year period (sa weather service, 26 october 2006), compared with that of the previous two summers at the time of the study 0 10 20 30 40 50 60 70 80 90 100 march (n = 593) 0 5 10 15 20 25 30 35 40 45 august (n = 217) 0 5 10 15 20 25 30 35 40 october (n = 150) 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 0 10 20 30 40 50 t ot al n um be r of s na ils january (n = 150) j snail size (mm) original research odendaal, haupt & griffiths koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 93 98 figures, considerably lower than may occur during wetter and more favourable conditions. snail populations occur mainly along roads and around existing or abandoned human establishments. at some of these sites densities were extremely high, to the point that snails coated the stems of the vegetation (figure 3). at such densities it is inevitable that these invasive snails will have significant impacts on the flora, although the nature of such impacts will depend on their feeding rates and dietary selectivity – aspects of their biology that remain unknown at this time. however, several of the plant species on which t. pisana were commonly found is indigenous and showed extensive damage, indicating that t. pisana directly impacts the indigenous fynbos flora (table 3). the presence of such a high biomass of snails is also likely to affect the entire food chain, since they are by far the dominant animal species at these high density sites. more importantly, high density sites did not appear to have a distinct flora, or indeed any other unique characteristics, suggesting that the high snail densities there simply represent a more established population, and that, in time, other areas of the park are likely to become equally seriously infested. no, or very few, snails were active during the day. at night a few snails became active when the weather was cold (figure 6a), but on a warm humid night almost all the snails were active, particularly shortly after dusk and in the hours before dawn (figure 6b). mcquaid et al. (1979) found that t. pisana aroused from aestivation when induced by a combination of low temperature and high humidity. our preliminary observations agree that activity is promoted by high humidity, but suggest that snails were also active when this was combined with relatively high temperature. mcneely (2001) argues that many invasive species do well in areas that have been disturbed by human activities. thus, the low density inside the park could also be due to the relatively undisturbed environment of the wcnp. this is supported by the fact that the average density of t. pisana in the severely disturbed areas outside of the park (langebaan town and adjacent road verges) was more than 14 times higher than that inside the park (table 1). furthermore, t. pisana colonies were mainly found in disturbed areas of the park e.g. adjacent to roads and human establishments. figure 4, showing the decline in snail densities as one moves away from the road at four different sites within the park, further supports this. in march (the beginning of autumn) the population comprised of mostly adult snails with a mean diameter of 14 mm. by august, most of these adults had disappeared and were replaced by juveniles with the diameter averaging 6 mm. the size of snails increased rapidly during spring and by october, when a unimodal cohort of snails with a diameter of 11 mm dominated the population (figure 5), this increasing to 14 mm by midsummer (january). durr (1946) stated that oviposition in t. pisana occurs from march to may and that they have an annual life cycle, and our results support these conclusions. theba pisana in britain and france have an annual life cycle and breed in summer and autumn, while in the mediterranean, they have a biennial life cycle with breeding in autumn and winter (cowie 1985). in conclusion, it is clear that t. pisana are well adapted to conditions in the western cape and can reach very high densities within the wcnp, although our observations of large numbers of dead shells suggest that the population has recently undergone a significant mortality event associated with unusually dry conditions. at peak densities it appears certain that this snail will have profound impacts on the growth and survivorship of favoured food species, although these effects have not yet been quantified. the plant species listed in table 3 could be used as a starting point for investigating the food preferences of t. pisana. to date, the mean density within the park is much lower than the density of more disturbed areas outside the park, suggesting that the populations in the park have not yet reached carrying capacity. more research regarding various life history characters of t. pisana is required to better understand and identify the impacts of these snails on biotic and abiotic ecosystem processes of the fynbos biome. these include studies that would investigate dispersal rates (using mark-recapture methods), as well as competitive interactions between these snails and other fauna. for example, in their study on the unique use of empty shells of indigenous land snails (several species of trigonephrus and tropidophora ligata) as nest sites by several previously undescribed species of silk-spinning wasps belonging to the genus quartinia, gess and gess (2007) found that the relatively small shells of t. pisana were unsatisfactory nesting sites for these wasps. less than 10% of t. pisana shells were used as nests compared to an average use of 53% of shells from indigenous snails. also, in areas where t. pisana was present, they were much more abundant than indigenous snails. since fynbos is not only globally important as a biodiversity hot-spot biome, but also locally important to the economic wellbeing of the western cape, the potential impacts of t. pisana on indigenous flora and fauna is indeed cause for concern. acknowledgements thank you to elizabeth kelly, melinda griffiths, taryn duthie, monique boucher and bradley kruger for their assistance in the field, the management of the west coast national park for permission to conduct this study and p. nel for the use of data. thanks are also extended to d. herbert and the staff of the gis lab (university of cape town). this study was funded by grants to l.j. odendaal and t.m. haupt from the nrf/dst centre of excellence for invasion biology. 97 figure 6 percentage of active individuals in response to temperature (°c) and humidity (%) during (a) a dry sunny and (b) a humid day a) b) 0 10 20 30 40 50 60 70 80 90 100 8:00 10:00 12:00 14:00 16:00 18:00 20:00 22:00 0:00 2:00 4:00 6:00 8:00 time of day a ct iv ity & h um id ity % 0 5 10 15 20 25 30 t em pe ra tu re ( °c ) temperature(°c) 0 20 40 60 80 100 120 08:00 10:00 12:00 14:00 16:00 18:00 20:00 22:00 00:00 02:00 04:00 06:00 08:00 time of day a ct iv ity & h um id ity (% ) 0 5 10 15 20 25 30 35 te m pe ra tu re (° c ) alien invasive land-snail original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 93 98 macdonald, i.a.w. & jarman, m.l. 1984. invasive alien organisms in the terrestrial ecosystems of the fynbos biome, south africa. south african national scientific programmes report, no. 85. mcneely, j. 2001. invasive species: a costly catastrophe for native biodiversity. land use and water resources research, 1(2): 1–10. mcquaid, c.d., branch, g.m. & frost, p.g. 1979. aestivation behaviour and thermal relations of the pulmonate theba pisana in a semi-arid environment. journal of thermal biology, 4: 47–56. mooney, h.a. & cleland, e.e. 2001. the evolutionary impact of invasive species. colloquium, 98(10): 5446–5451. myburgh, a.c. & rust, d.j. 1986. micellaneous pests. in: myburgh, a.c. (ed.). crop pests in southern africa: deciduous fruit, grapes and berries. pretoria: plant protection research institute bulletin 407, pp. 83–85. olden, j.d., poff, n.l., douglas, m.r., douglas, m.e. & fausch, k.d. 2004. ecological and evolutionary consequences of biotic homogenization. trends in ecology and evolution, 19: 18–24. quick, h.e. 1952. emigrant british snails. proceeding of the malacological society of london, 29: 181–189. richardson, d.m. 1999. commercial forestry and agroforestry as sources of invasive alien trees and shrubs. in: o. t. sandlund, p. j. schei & a. viken (eds.). invasive species and biodiversity management. dordrecht: kluwer academic publishers, pp. 237–258. south african weather service. climate data for langebaanweg. available at: http://www.weathersa.co.za/climat/ climstats/langebaanstats.jsp [accessed 26 october 2006]. statistica for windows. 2004. release 7.0. tulsa, usa: statsoft inc. turpie, j.k., heydenrych, b.j. & lamberth, s.j. 2003. economic value of terrestrial and marine biodiversity in the cape floristic region: implications for defining effective and socially optimal conservation strategies. biological conservation, 112(1/2): 233–257. van wilgen. b.w., richardson, d.m., le maitre, d.c. marais, c. & magadlela, d. 2001. the economic consequences of alien plant invasions: examples of impacts and approaches to sustainable management in south africa. environment, development and sustainability, 3(2): 145–168. west coast national park. park management plan october 2006. available at: http://celtis.sanparks.org/conservation/ park_man/westcoast.pdf [accessed 20 december 2006]. references baker, g.h. 1986. the biology and control of white snails (mollusca: helicidae), introduced pests in australia. csiro australia, division of entomology, technical paper, 25: 1–31. baker, g.h. 1989. damage, population dynamics, movement and control of pest helicid snails in southern australia. in: hendersen, i. (ed.). slugs and snails in world agriculture, volume 41. thornton heath, british crop protection council monograph, pp. 175–185 baker, g.h. & vogelzang, b.k. 1988. like-history, population dynamics and polymorphism of theba pisana (mollusca: helicidae) in australia. journal of applied ecology, 25: 867– 887. cowie, r.h. 1985. microhabitat choice and high temperature tolerance in the land snail theba pisana (mollusca: gastropoda) biological conservation, 78: 207–214. cowling, r.m., macdonald, i.a. & simmons, m.t. 1996. the cape peninsula, south africa: physiographical, biological and historical background to an extraordinary hot-spot of biodiversity. biodiversity and conservation, 5(5): 527–550. durr, h.j.r. 1946. a contribution to the morphology and bionomics of theba pisana (muller) (gastropoda: helicidae). south african department of agriculture. stellenbosch-elsenburg science series, 47: 1–34. gess, s. & gess, f. 2007. snail invasion: the potential impact of the invasive mediterranean snail, theba pisana on our coastal dune vegetation. veld and flora, 93: 216–218. goldblatt, p. & manning, j.c. 2002. plant diversity of the cape region of south africa. annals of the missouri botanical garden, 89: 281–302. herbert, d.g. & kilburn, d. 2004. field guide to the land snails and slugs of eastern south africa. pietermaritzburg: natal museum. herbert, d.g. & sirgel, w.f. 2001. the recent introduction of two potentially pestiferous alien snails into south africa and the outcomes of different pest management practices: an eradication and a colonization. south african journal of science, 97: 301–304. joubert, c.j. & walters, s.s. 1951. the control of snails and slugs. farming south africa, 26: 379–380. lach, l., picker, m.d., colville, j.f., allsopp, m.h. & griffiths, c.l. 2002. alien invertebrate animals in south africa. in: pimentel, d. (ed.). biological invasions: economic and environmental costs of alien plant, animal and microbe species. new york: crc press, pp. 267–282. 98 abstract introduction methods results ethical considerations discussion acknowledgements references about the author(s) keoikantse sianga okavango research institute, university of botswana, botswana richard w.s. fynn okavango research institute, university of botswana, botswana mpaphi c. bonyongo okavango research institute, university of botswana, botswana citation sianga, k., fynn, r.w.s. & bonyongo, m.c., 2017, ‘seasonal habitat selection by african buffalo syncerus caffer in the savuti–mababe–linyanti ecosystem of northern botswana’, koedoe 59(2), a1382. https://doi.org/10.4102/koedoe.v59i2.1382 original research seasonal habitat selection by african buffalo syncerus caffer in the savuti–mababe–linyanti ecosystem of northern botswana keoikantse sianga, richard w.s. fynn, mpaphi c. bonyongo received: 26 jan. 2016; accepted: 24 nov. 2016; published: 23 may 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract this study aimed to establish seasonal movement and habitat selection patterns of african buffalo syncerus caffer in relation to a detailed habitat map and according to seasonal changes in forage quality and quantity in the savuti–mababe–linyanti ecosystem (botswana). two buffalo were collared in november 2011 and another in october 2012. all three buffalo had greater activities in the mopane–sandveld woodland mosaic during the wet season, which provided high-quality leafy grasses and ephemeral water for drinking, but moved to permanent water and reliable forage of various wetlands (swamps and floodplains) and riverine woodlands during the dry season. wetlands had higher grass greenness, height and biomass than woodlands during the dry season. buffalo had similar wet season concentration areas in the 2011–2012 and 2012–2013 wet seasons and similar dry season concentration areas over the 2012 and 2013 dry seasons. however, their dry season location of collaring in 2011 differed dramatically from their 2012 and 2013 dry season concentration areas, possibly because of the exceptionally high flood levels in 2011, which reduced accessibility to their usual dry season concentration areas. the study demonstrates that extremely large and heterogeneous landscapes are needed to conserve buffalo in sandy, dystrophic ecosystems with variable rainfall. conservation implications: this study emphasises the importance of large spatial scale available for movement, which enables adaptation to changing conditions between years and seasons. introduction african buffalo (syncerus caffer) have a large distributional range across the savannas of africa. their habitat selection and foraging ecology have been relatively well studied (landman & kerley 2001; macandza, owen-smith & cross 2004; sinclair 1979; taylor 1985). because of their large body size and large groups, buffalo are able to fend off predators (sinclair, mduma & brashares 2003), thereby enabling them to forage in wooded vegetation with relatively low visibility and high predation risk. the large absolute food demands of buffalo, which is a function of their large body size (illius & gordon 1987; wilmshurst, fryxell & bergman 2000), combined with their inability to efficiently crop short grass (use of tongue to increase bite size cannot work on short grass) constrain them to foraging in vegetation with sufficient height and biomass of forage (codron et al. 2008; illius & gordon 1987). buffalo generally avoid heavily grazed regions of short grassland (bhola et al. 2012; jarman & sinclair 1979; traill & bigalke 2007), preferring woodlands dominated by tufted and leafy perennial grasses during the wet season and often relying on more productive riverine vegetation during the dry season (bell 1970; bennitt, bonyongo & harris 2014; fynn, chase & roder 2014; macandza, owen-smith & cain 2012; sinclair 1979). while drylands often support less productive but higher quality forage than wetlands over the wet season (fynn et al. 2014; taylor 1985), their moisture-limited position in the landscape results in forage drying out and declining in forage quality and quantity over the dry season (ellis & swift 1988; owen-smith 2008; taylor 1985). thus, wetlands, where permanent water is easily accessible and perennial grasses continue to produce green forage over the dry season, would be expected to be favoured by many herbivores at this time with drylands being favoured over the wet season (bell 1970; fynn et al. 2014; maddock 1979; taylor 1985). in contrast to more mobile species such as zebra (equus burchellii) and wildebeest (connochaetes taurinus), which often migrate between distinct wetand dry season ranges (bartlam-brooks, bonyongo & harris 2011; maddock 1979), most buffalo studies have found strong overlap between wetand dry season ranges (macandza et al. 2012; ryan 2006; ryan, knechtel & getz 2006; venter & watson 2008). many studies, however, were conducted in small reserves (landman & kerley 2001; ryan et al. 2006; tshabalala, dube & lent 2009) where opportunity for large seasonal movements are limited or in larger reserves where functional heterogeneity of resources may be well developed at the landscape (catena) scale (bell 1970; macandza et al. 2012; perrin & brereton-stiles 1999; sinclair 1979), thereby reducing the need for large seasonal movements (hopcraft, olff & sinclair 2010). however, in large, relatively unfragmented ecosystems and where landscape scale functional heterogeneity may be poorly developed relative to regional-scale heterogeneity (fynn et al. 2014), larger buffalo movements may be expected (naidoo et al. 2012; skarpe et al. 2004). the savuti–mababe–linyanti ecosystem (smle) is part of the > 80 000 km2 northern conservation area of botswana, one of the largest relatively unfragmented wildlife regions in africa (fynn & bonyongo 2011). despite several studies on buffalo in the smle (fynn et al. 2014; patterson 1972), detailed seasonal movement and habitat selection patterns of buffalo in the region have not been fully identified and established, especially seeing that no detailed habitat map has been available until this year (2016). considering their non-specialised mouth anatomy and their large body size–mediated demands for absolute food intake, we expected buffalo in the smle to favour vegetation where tufted, leafy grasses provided sufficient height and biomass of grass to enable food intake requirements to be satisfied (illius & gordon 1987; wilmshurst et al. 2000) (hypothesis 1 – h1). the strongly developed regional-scale distribution (rather than landscape scale) of functional heterogeneity of wetand dry season vegetation types in the smle (fynn et al. 2014) is likely to promote extension of buffalo home ranges beyond the landscape scale (hopcraft et al. 2010). consequently, we expected seasonal movement patterns of buffalo in the smle to match the scale at which functional seasonal vegetation types are distributed in the ecosystem, which is strongly regional (fynn et al. 2014) (hypothesis 2 – h2). the objectives of this study were to (1) to examine seasonal movements and habitat selection of buffalo in the smle of northern botswana and (2) to link seasonal movements to the quality and quantity (grass greenness, height and biomass) of vegetation in favoured seasonal regions of the landscape. methods study area this study was conducted in the smle (northern botswana, figure 1). climate in the study area is described as semi-arid with mean annual rainfall ranging from around 500 mm in the okavango region (western boundary of the smle) to over 600 mm in the chobe enclave (eastern boundary of the smle), most of which is received between november and april (botswana meteorological services). seasons may be functionally separated into a wet season (december–april), a cool early dry season (may–august) and a hot late dry season (september–november), where daily maximum temperatures are between 35 °c and 40 °c (figure 2, adapted from fynn et al. 2014). the smle is defined by a variety of vegetation types from swamps, floodplains and riverine in wetter areas to vast dryland woodland systems away from permanent water comprising a mosaic of mopane woodland on alluvial soils and sandveld woodland on kalahari sands (figure 1; sianga & fynn 2017; wolski & murray-hudson 2006). in addition, very heavy clay soils formed under sedimentation in a paleolake system known as the mababe depression (teter 2007) give rise to open grassland and sparse savanna in the eastern section of the smle (figure 1). another key feature are the dambo grasslands of the chobe enclave, which are seasonally flooded shallow, linear depressions (flooded by runoff from upslope regions) supporting tall grassland. figure 1: savuti–mababe–linyanti ecosystem vegetation. figure 2: rainfall and temperature patterns between 2010 and 2013. (a) 2010/2011, (b) 2011/2012 and (c) 2012/2013 rainfall season. buffalo habitat use single cows in three buffalo herds were fitted with satellite collars (africa wildlife tracking, pretoria, south africa). buffalo 1 (bh1) and buffalo 2 (bh2) were collared in the savuti channel floodplains and near the savuti marsh, respectively, at the end of the late dry season in november 2011. buffalo 3 (bh3) was collared in the savuti marsh in the late dry season (october 2012). anaesthetic m3080 xylazine was used to immobilise cows during capturing and later reversed using naltrexone after fitting collars. collars were programmed to take a global positioning system (gps) position of the animal every 4 hours. every fix obtained by the satellite collars was downloaded and plotted in arcgis 10.1 (esri 2010) for various analyses. for analyses relating to seasonal differences in movement patterns and habitat selection, we defined six seasonal periods to which each gps fix could be allocated. the seasonal periods were early wet season (mid-november to end of december), mid wet season (january and february), late wet season (march and april), early dry season (may and june), mid dry season (july and august) and late dry season (september to mid-november). for visual presentation of seasonal movements and locations for each buffalo, we plotted the home ranges of each buffalo (colour coded for each of the six seasonal periods) in a local convex hull (a-locoh, 95% isopleths) (getz et al. 2007) in r (rcore-team 2013) and later displayed as a shapefile on a habitat map of the region. the habitat map was developed from a detailed vegetation classification and mapping study of the smle funded by southern african science service centre for climate change and adaptive land management (sasscal); a detailed account of the vegetation communities and habitat map can be seen in an accompanying paper in this special issue on botswana (sianga & fynn 2017). locoh has been found to be an appropriate tool in gps studies (getz et al. 2007). vegetation sampling gps data from satellite collars were used to locate sites where buffalo have been during the wet and dry seasons. sampling of buffalo’s wet season vegetation types was conducted in the 2012–2013 wet season, while their dry season vegetation types were sampled in the 2012 dry season. a total of 124 samples (grass greenness, height and biomass) were collected from sites where the buffalo have been. seventy-five samples (20 and 55) were collected for bh1 during the early and late dry season of 2012, while 20 samples were collected for bh2 in the late dry season of 2012. at each site, five 0.25 m2 quadrats were set-up at the gps position obtained from the collar. the first quadrat was set at the 0-m position on the hand-held gps (garmin gps map 62s) and the other four quadrats 5 m each side of the first (central) quadrat. greenness was estimated visually as the percentage of green tissue of grasses and sedges rooted within the quadrat. grass height was measured by lowering a brown paper sample bag at the centre of each quadrat, and the height above the soil surface was measured. all grasses rooted within the quadrats were clipped at ground surface level and air-dried during the field exercise. the air-dried grasses were oven-dried at 60 °c for 48 hours and weighed for biomass at the okavango research institute laboratory (maun, botswana). in addition, to determine the vegetation type, we noted grasses and trees common within approximately a 10-m radius around the central quadrat. statistical analyses home ranges and habitat selection for analysis of habitat selection, we determined seasonal minimum convex polygons (mcps) in arcgis 10.1 (esri 2010) and local convex hulls in r (getz et al. 2007; rcore-team 2013) using seasonal location data of each collared buffalo. seasonal habitat selection indices of each buffalo were determined by dividing the proportion of use of each habitat by the proportion of availability of each habitat following jacobs index (j.i) = (r ‑ p)/(r + p – 2rp), where r is the proportion of habitat used, and p is the proportion of habitat available (jacobs 1974). j.i ranges between –1 (selected against) and +1 (selected for). to consider the effects of scale on resource availability (gustine et al. 2006), we calculated habitat selection at second and third order. second-order selection (johnson 1980) was determined by comparing the availability of various habitat types in the individual buffalo mcps against availability in the overall population mcp (thomas & taylor 1990), while third-order selection (johnson 1980) was determined by comparing the availability of various habitat types in the individual buffalo local convex hulls to availability in the respective individual buffalo mcps (thomas & taylor 1990). owing to having only three collared individual buffalo, there was insufficient replication to effectively statistically test habitat selection. thus, the j.i merely provides a guide to readers on potential habitat selection. to determine home-range overlaps between seasons, we used intersect tool (universal transverse mercator, esri 2010) to estimate area percentage overlaps, where % overlaps = area overlap of two seasons/(area of season 1 + season 2) × 100. to examine how buffalo moved in relation to distance from perennial water sources differed seasonally, we determined the distance to the nearest perennial water source of every gps location of all buffalo throughout the study period using near tool in arcgis 10.1 (universal transverse mercator, esri 2010). vegetation data data of grass greenness, height and biomass were subjected to shapiro–wilk test (test of normality) and levene statistic (test of homogeneity of variance) in r version 2.15.2 (rcore-team 2013). data sets that failed assumptions of normality or homogeneity of variance were natural log transformed, and failure to meet these assumptions after transformation led to the use of non-parametric tests (kruskal–wallis test). multiple comparison tests (kruskalmc’s function) in ‘pgirmess’ package in r (rcore-team 2013) was used to analyse data sets. results all three buffalo had much greater activities in sandveld and mopane woodlands during the wet season and various wetland systems during the dry season (figure 3). bh1 and bh2 had greater separation of their wet and dry season concentration areas than bh3 (quantified by percentage overlap of their wet and dry season 95% local convex hull isopleths – table 1; also see figure 4 vs. figure 5). bh1 and bh2 demonstrated distinct wet and dry season concentration areas, returning to these respective seasonal locations in successive years (table 1; figure 6). figure 3: percentage of various vegetation types observed in the follow-ups of gps collar locations during the wet season and late dry season. figure 4: movement patterns of buffalo 1 (bh1) and buffalo 2 (bh2) over the 2011/2012 annual cycle, in relation to vegetation. figure 5: movement patterns of buffalo 3 (bh3) over the 2012/2013 annual cycle, in relation to vegetation. figure 6: movement patterns of buffalo 1 (bh1) and 2 (bh2) in the mid wet and mid dry season of 2012 and 2013, in relation to vegetation. table 1: percentage overlaps of home ranges by buffalo in different seasons over the 2012 and 2013. for second-order habitat selection, the buffalo selected mopane and sandveld woodlands over the wet season, whereas riverine and wetland habitats were selected over the dry season (table 2); for third-order habitat selection, the buffalo selected mopane woodlands over the wet season and acacia grasslands, whereas riverine and wetland habitats were selected over the dry season (table 3). for more details of the plant species composition and classification of these plant communities and for details of the habitat map development, see a parallel paper in this special issue (sianga & fynn 2017). over both dry seasons, bh1 concentration areas were in wetland and adjacent woodlands where it focused on the linyanti swamp in increasing proportions (and regularly across the international border with namibia) as the dry season progressed (figure 4). the concentrations of bh2 were in the selinda spillway and kwando river wetland (floodplains) and adjacent woodlands in the 2012 and 2013 dry seasons (figures 4 and 6). interestingly, bh2 concentrated its activities in the tsum tsum wetlands (floodplains) of the okavango delta in the late wet season of april 2013 (which it did not do in 2012) before moving to the selinda spillway later in the dry season (figure 6). bh3 had the wetland habitats of the savuti channel and savuti marsh at the core of its range, which it focused on during the dry season, and utilised adjacent woodlands on the peripheries of the home range during the wet season (figure 5). table 2: second-order habitat selection (annual minimum convex polygons vs. wet or dry seasons minimum convex polygons) by three buffalo in the savuti–mababe–linyanti ecosystem (northern botswana). table 3: third-order habitat selection (annual minimum convex polygons vs. wet or dry season local convex hull) by three buffalo in the savuti–mababe–linyanti ecosystem (northern botswana). during the early and late dry season of 2012, wetland habitats (floodplain not sampled in the early dry season) had higher grass greenness, height and biomass than mopane, sandveld and riverine habitats, respectively (p < 0.05; figure 7), while mopane, sandveld and riverine habitats were not significantly different in grass greenness, height and biomass, respectively (p > 0.05; figure 7). figure 7: (a) grass greenness, (b) height and (c) biomass of mopane woodland, sandveld, riverine and wetland used by buffalo herds during the early and late dry season. ethical considerations a veterinarian registered with the government of botswana conducted all darting operations in principle with the research permit (ewt 8/36/4 xvii [31]) and the supplementary collaring permit (ewt 8/36/4 xvii [41]) issued by the ministry of environment, wildlife and tourism and the department of wildlife and national parks (gaborone, botswana). darting operations or removing collars from cows were done from a vehicle and a helicopter, respectively. collars were removed successfully after two successive years. discussion buffalo shifted their seasonal concentration areas between wetlands and woodlands (figures 3 and 4) as an adaptive strategy to seasonal forage dynamics and water availability over the annual cycle, with bh1 and bh2 having a migratory strategy with little overlap between wet and dry season concentration areas. however, the distance moved from perennial water sources over the wet season varied significantly across the buffalo, with bh3 being more sedentary. bennitt et al. (2014) and taolo (2003) reported similar results, noting that buffalo selected contrasting seasonal vegetation types, using woodlands far from permanent water during the rainy season and seasonally flooded vegetation close to permanent water during the early and late flood seasons. as with our findings, several other studies in the region have also found a mix of strategies (migratory vs. sedentary) between buffalo (bennitt et al. 2014; bennitt, bonyongo & harris 2015; naidoo et al. 2012, 2014). cornélis et al. (2011) also noted similar findings where buffalo were observed to range between woodlands and riverine habitats during the wet and dry season, respectively. a key factor that may influence buffalo concentration patterns during the wet season appears to be the availability of higher quality, soft-leaved grasses (such as digitaria eriantha) in back country woodlands than in wetlands (floodplains), which in contrast support tougher-leaved grasses adapted to shading and litter accumulation under the more productive conditions of wetlands (fynn & bonyongo 2011), and where greater biomass dilutes nutrient concentrations (jarrell & beverly 1981), while greater silica contents of wetland versus dryland grasses further reduce digestibility (mosimane 2015). one of the most favoured grasses by buffalo, d. eriantha, peaks in abundance beyond 10 km from the permanent water of the okavango delta and linyanti swamps, whereas shorter grazing lawn–forming grasses such as urochloa trichopus peaked in abundance with 5 km of these extensive wetland systems (sianga et al. 2017). the medium to tall, leafy high-quality d. eriantha would be expected to be optimal in height and digestibility for maximising protein and energy intake rates in buffalo, which explains why buffalo focused their wet season foraging beyond 10 km from wetland systems (figure 4). by contrast, wildebeest and impala, which favour shorter grasses, focused within 5 km of these wetland systems (fynn et al. 2014). this suggests that short and tall grass grazers in the smle have distinct spatial niche separation during the wet season driven by distance to wetland systems. similar observations have been made in the maasai-mara, where buffalo favour the taller grass areas within the reserve and wildebeest and gazelles the shorter grass areas outside the reserve (bhola et al. 2012). thus, a high conception rate of buffalo during the wet season is probably because of the availability of high-quality green forage of optimal height and biomass (ryan, knechtel & getz 2007). thus, concentration during the wet season by the three buffalo in vegetation types (sandveld and mopane) where tufted, soft-leaved grasses are more abundant supports our first hypothesis (h1). drying up of ephemeral pans over the dry season forced buffalo to move away from favoured wet season ranges towards permanent water sources (figures 3 and 4), which included floodplains and swamps (wetlands) and adjacent woodlands within the study area. it is this forced movement away from favoured wet season ranges towards permanent water that likely contributes to reduced grazing pressure in favoured wet season ranges and, therefore, higher abundances of favoured grasses such as d. eriantha far from permanent water. this indicates that artificial water provision in back country woodlands far from permanent water may be an unwise management option that could reduce niche diversity in the system leading to declines in taller grass grazers such as buffalo, as well as roan and sable antelope. these floodplain and swamp (wetlands) systems provided buffalo not only with reliable drinking water during the dry season but also with forage significantly greener than that found in woodlands, as observed in other studies in the region (bennitt et al. 2014; fynn et al. 2014; taolo 2003). more productive wetland areas are likely to provide a critical reserve and critical buffer resources (owen-smith 2002) for the late dry season, or elsewhere referred to as key resources (illius & o’connor 2000), especially during drought years when more preferred resources have been depleted (owen-smith 2002). the ability of wetlands to provide taller green forage with higher energy and protein levels relative to drylands during the late dry season has been shown to greatly elevate buffalo population productivity (taylor 1985). the taller grass of wetlands is likely to be especially critical for buffalo, which are highly vulnerable to competition for forage (bhola et al. 2012), owing to their tongue sweep foraging strategy, which cannot deal with short grass. thus, our findings of increasing use of swamp, marsh and floodplains (wetlands) as the dry season progressed by our three buffalo together with similar observations in other studies (prins & beekman 1989; taylor 1985; tinley 1977; vesey-fitzgerald 1960; western 1973), demonstrate the importance of wetland systems as dry season key-resource habitat for buffalo, as is the case for cattle and other wild herbivores across africa (fynn et al. 2015). our collared buffalo migrated at an intermediate scale from wetlands to woodlands rather than from wetlands to the high-quality mababe depression grasslands, favoured by a migratory zebra population over the wet season (sianga 2014) and, thus, our second hypothesis (h2) that buffalo seasonal movements would match the regional-scale distribution of functional seasonal vegetation types was not supported. though this finding does not support our h2, it does not rule out the possibility of buffalo migrating into the depression during the wet season because not all buffalo herds in the ecosystem were fitted with collars to determine their movements. also, aerial surveys conducted by fynn et al. (2014) did not observe any buffalo in the mababe depression at that time, suggesting that buffalo probably do not favour the mababe depression during the wet season, unlike the zebra population (sianga 2014). however, the buffalo appear to show adaptive variation in seasonal presence to changing environmental conditions in different years. for instance, in 2011, bh1 and bh2 avoided floodplains and swamps (wetlands) in their usual dry season concentration areas of the linyanti swamps and selinda spillway (2012 and 2013) because of the exceptionally high floods of 2011, being forced to use the more elevated floodplains of the savuti channel. this shift in location of different dry season concentration areas emphasises the importance of having a large spatial scale available for movement (fynn et al. 2014), which enables adaptation to changing conditions between years. finally, the finding of cross-border movements by bh1, together with those of naidoo et al. (2014) and patterson (1972), demonstrates the importance of transfrontier conservation areas such as this one (kavango zambezi transfrontier conservation area, kaza), which transcends the borders of angola, namibia, botswana, zimbabwe and zambia allowing adaptive foraging to a variety of different functional seasonal resources. acknowledgements we thank the german ministry of education and research (bmbf) who funded this work through ‘the future okavango project’ (buffalo study) and through southern african science service centre for climate change and adaptive land management (sasscal) (vegetation study and buffalo habitat selection analyses). dr andy lyons is acknowledged for his guidance with tlocoh. dr rob jackson is appreciated for buffalo darting, collar fitting and removal. finally, we thank the ministry of environment, wildlife and tourism and the department of wildlife and national parks (botswana) for permitting us to conduct research in northern botswana. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions all authors of this research article have directly participated in the planning, execution and analysis of this study. references bartlam-brooks, h., bonyongo, m. & harris, s., 2011, ‘will reconnecting ecosystems allow long-distance mammal migrations to resume? a case study of a zebra equus burchelli migration in botswana’, oryx 45, 210–216. https://doi.org/10.1017/s0030605310000414 bell, r., 1970, ‘the use of the herb layer by grazing ungulates in the serengeti’, in a. watson (ed.), animal populations in relation to their food resources, pp. 111–124, blackwell, oxford. bennitt, e., bonyongo, m.c. & harris, s., 2014, ‘habitat selection by african buffalo (syncerus caffer) in response to landscape-level fluctuations in water availability on two temporal scales’, plos one 9, e101346. https://doi.org/10.1371/journal.pone.0101346 bennitt, e., bonyongo, m.c. & harris, s., 2015, ‘behaviour-related scalar habitat use by cape buffalo (syncerus caffer)’, plos one 10, e0145145. https://doi.org/10.1371/journal.pone.0145145 bhola, n., ogutu, j.o., piepho, h.p., said, m.y., reid, r.s., hobbs, n.t. et al., 2012, ‘comparative changes in density and demography of large herbivores in the masai mara reserve and its surrounding human-dominated pastoral ranches in kenya’, biodiversity & conservation 21, 1509–1530. https://doi.org/10.1007/s10531-012-0261-y codron, d., brink, j.s., rossouw, l., clauss, m., codron, j., lee-thorp, j.a. et al., 2008, ‘functional differentiation of african grazing ruminants: an example of specialized adaptations to very small changes in diet’, biological journal of the linnean society 94, 755–764. https://doi.org/10.1111/j.1095-8312.2008.01028.x cornélis, d., benhamou, s., janeau, g., morellet, n., ouedraogo, m. & visscher, m.n., 2011, ‘spatiotemporal dynamics of forage and water resources shape space use of west african savanna buffaloes’, journal of mammalogy 92(6), 1287–1297. https://doi.org/10.1644/10-mamm-a-397.1 ellis, j.e. & swift, d.m., 1988, ‘stability of african pastoral ecosystems: alternate paradigms and implications for development’, journal of range management archives 41, 450–459. https://doi.org/10.2307/3899515 esri, 2010, arcgis desktop, environmental systems research institute, redlands, ca. fynn, r.w.s. & bonyongo, m.c., 2011, ‘functional conservation areas and the future of africa’s wildlife’, african journal of ecology 49, 175–188. https://doi.org/10.1111/j.1365-2028.2010.01245.x fynn, r.w.s., chase, m. & roder, a., 2014, ‘functional habitat heterogeneity and large herbivore seasonal habitat selection in northern botswana’, south african journal of wildlife research 44, 1–15. https://doi.org/10.3957/056.044.0103 fynn, r.w.s., murray-hudson, m., dhliwayo, m. & scholte, p., 2015, ‘african wetlands and their seasonal use by wild and domestic herbivores’, wetlands ecology and management 23, 559–581. https://doi.org/10.1007/s11273-015-9430-6 getz, w.m., fortmann-roe, s., cross, p.c., lyons, a.j., ryan, s.j. & wilmers, c.c., 2007, ‘locoh: nonparametric kernel methods for constructing home ranges and utilization distributions’, plos one 2, e207. https://doi.org/10.1371/journal.pone.0000207 gustine, d.d., parker, k.l., lay, r.j., gillingham, m.p. & heard, d.c., 2006, ‘interpreting resource selection at different scales for woodland caribou in winter’, journal of wildlife management 70, 1601–1614. https://doi.org/10.2193/0022-541x(2006)70[1601:irsads]2.0.co;2 hopcraft, j.g.c., olff, h. & sinclair, a., 2010, ‘herbivores, resources and risks: alternating regulation along primary environmental gradients in savannas’, trends in ecology & evolution 25, 119–128. https://doi.org/10.1016/j.tree.2009.08.001 illius, a.w. & gordon, i.j., 1987, ‘the allometry of food intake in grazing ruminants’, the journal of animal ecology 56, 989–999. https://doi.org/10.2307/4961 illius, a.w. & o’connor, t.g., 2000, ‘resource heterogeneity and ungulate population dynamics’, oikos 89, 283–294. https://doi.org/10.1034/j.1600-0706.2000.890209.x jacobs, j., 1974, ‘quantitative measurement of food selection’, oecologia 14, 413–417. https://doi.org/10.1007/bf00384581 jarman, p.j. & sinclair, a.r.e., 1979, ‘feeding strategy and the pattern of resource partitioning in african ungulates’, in a.r.e. sinclair & m. norton-griffiths (eds.), serengeti: dynamics of an ecosystem, pp. 130–163, university of chicago press, chicago, il. jarrell, w. & beverly, r., 1981, ‘the dilution effect in plant nutrition studies’, advances in agronomy 34, 197–224. https://doi.org/10.1016/s0065-2113(08)60887-1 johnson, d.h, 1980, ‘the comparison of usage and availability measurements for evaluating resource preference’, ecology 61, 65–71. https://doi.org/10.2307/1937156 landman, m. & kerley, g., 2001, ‘dietary shifts: do grazers become browsers in the thicket biome?’, koedoe 44, 31–36. https://doi.org/10.4102/koedoe.v44i1.183 macandza, v.a., owen-smith, n. & cain, j.w., 2012, ‘habitat and resource partitioning between abundant and relatively rare grazing ungulates’, journal of zoology 287, 175–185. https://doi.org/10.1111/j.1469-7998.2012.00900.x macandza, v.a., owen-smith, n. & cross, p.c, 2004, ‘forage selection by african buffalo in the late dry season in two landscapes’, south african journal of wildlife research 34, 113–121. maddock, l., 1979, ‘the migration and grazing succession’, in a.r.e. sinclair & m. norton-griffiths (eds.), serengeti: dynamics of an ecosystem, pp. 104–129, university of chicago press, chicago, il. mosimane, k., 2015, ‘a first quantification of biogenic silica pools in an evaporation-dominated wetland system: the okavango delta’, mphil thesis, university of botswana. naidoo, r., du preez, p., stuart-hill, g., beytell, p. & taylor, r., 2014, ‘long-range migrations and dispersals of african buffalo (syncerus caffer) in the kavango–zambezi transfrontier conservation area’, african journal of ecology 52, 581–584. https://doi.org/10.1111/aje.12163 naidoo, r., du preez, p., stuart-hill, g., jago, m. & wegmann, m., 2012, ‘home on the range: factors explaining partial migration of african buffalo in a tropical environment’, plos one 7, e36527. https://doi.org/10.1371/journal.pone.0036527 owen-smith, n., 2002, adaptive herbivore ecology: from resources to populations in variable environments, cambridge university press, cambridge. owen-smith, n., 2008, ‘changing vulnerability to predation related to season and sex in an african ungulate assemblage’, oikos 117, 602–610. https://doi.org/10.1111/j.0030-1299.2008.16309.x patterson, l., 1972, studies on the population biology of the african buffalo syncerus caffer in botswana, department of wildlife and national parks, gaborone, botswana. perrin, m. & brereton-stiles, r., 1999, ‘habitat use and feeding behaviour of the buffalo and the white rhinoceros in the hluhluwe-umfolozi game reserve’, south african journal of wildlife research 29, 72–80. prins, h.h.t. & beekman, j.h., 1989, ‘a balanced diet as a goal for grazing: the food of the manyara buffalo’, african journal of ecology 27, 241–259. https://doi.org/10.1111/j.1365-2028.1989.tb01017.x rcore-team, 2013, r: a language and environment for statistical computing, r foundation for statistical computing, vienna, austria, viewed 11 april 2015, from http://www.r-project.org ryan, s.j., 2006, ‘spatial ecology of african buffalo and their resources in a savanna ecosystem’, phd thesis, university of california, usa. ryan, s.j., knechtel, c.u. & getz, w.m., 2006, ‘range and habitat selection of african buffalo in south africa’, journal of wildlife management 70, 764–776. https://doi.org/10.2193/0022-541x(2006)70[764:rahsoa]2.0.co;2 ryan, s.j., knechtel, c.u. & getz, w.m., 2007, ‘ecological cues, gestation length, and birth timing in african buffalo (syncerus caffer)’, behavioral ecology 18, 635–644. https://doi.org/10.1093/beheco/arm028 sianga, k., 2014, ‘habitat use by buffalo and zebra in relation to spatial and temporal variability of resources in the savuti-mababe-linyanti ecosystem of northern botswana’, mphil thesis, university of botswana, botswana. sianga, k. & fynn, r., 2017, ‘the vegetation and wildlife habitats of the savuti-mababe-linyanti ecosystem, northern botswana’, koedoe 59(2), a1406. https://doi.org/10.4102/koedoe.v59i2.1406 sianga, k., van telgen, m., vrooman, j., fynn, r.w.s. & van langevelde, f., 2017, ‘spatial refuges buffer landscapes against homogenisation and degradation by large herbivore populations and facilitate vegetation heterogeneity’, koedoe 59(2), a1434. https://doi.org/10.4102/koedoe.v59i2.1434 sinclair, a.r.e., 1979, ‘the eruption of the ruminants’, in a.r.e. sinclair & m. norton-griffiths (eds.), serengeti – dynamics of an ecosystem, pp. 82–103. chicago university press, chicago, il. sinclair, a.r.e., mduma, s. & brashares, j.s., 2003, ‘patterns of predation in a diverse predator–prey system’, nature 425, 288–290. https://doi.org/10.1038/nature01934 skarpe, c., aarrestad, p.a., andreassen, h.p., dhillion, s.s., dimakatso, t., du toit, j.t. et al., 2004, ‘the return of the giants: ecological effects of an increasing elephant population’, ambio: a journal of the human environment 33, 276–282. taolo, c., 2003, ‘population ecology, seasonal movement and habitat use of the african buffalo (syncerus caffer) in chobe national park, botswana’, phd thesis, norwegian university of science and technology, trondheim, norway. taylor, r.d. 1985, ‘the response of buffalo syncerus caffer to the kariba lakeshore grasslands (panicum repens) in matusadona national park’, phd thesis, university of zimbabwe, zimbabwe. teter, k.l., 2007, ‘paleoenvironmental reconstruction of paleolake mababe, northwestern botswana from sediment chemistry and biological productivity data’, msc thesis, oklahoma state university, stillwater, ok. thomas, d.l. & taylor, e.j., 1990, ‘study designs and tests for comparing resource use and availability’, the journal of wildlife management 54, 322–330. https://doi.org/10.2307/3809050 tinley, k.l., 1977, ‘framework of the gorongosa ecosystem’, phd dissertation, university of pretoria, south africa. traill, l.w. & bigalke, r.c., 2007, ‘a presence-only habitat suitability model for large grazing african ungulates and its utility for wildlife management’, african journal of ecology 45, 347–354. https://doi.org/10.1111/j.1365-2028.2006.00717.x tshabalala, t., dube, s. & lent, p.c., 2009, ‘seasonal variation in forages utilized by the african buffalo (syncerus caffer) in the succulent thicket of south africa’, african journal of ecology 48, 438–445. https://doi.org/10.1111/j.1365-2028.2009.01132.x venter, j.a. & watson, l.h., 2008, ‘feeding and habitat use of buffalo (syncerus caffer) in the nama-karoo, south africa’, south african journal of wildlife research 38, 42–51. https://doi.org/10.3957/0379-4369-38.1.42 vesey-fitzgerald, d.f., 1960, ‘grazing succession among east african game animals’, journal of mammalogy 41, 161–172. https://doi.org/10.2307/1376351 western, d., 1973, ‘the structure, dynamics and changes of the amboseli ecosystem’, phd dissertation, university of nairobi, kenya. wilmshurst, j.f., fryxell, j.m. & bergman, c.m., 2000, ‘the allometry of patch selection in ruminants’, proceedings of the royal society of london. series b: biological sciences 267, 345–349. https://doi.org/10.1098/rspb.2000.1007 wolski, p. & murray-hudson, m., 2006, ‘flooding dynamics in a large low-gradient alluvial fan, the okavango delta, botswana, from analysis and interpretation of a 30-year hydrometric record’, hydrology and earth system sciences discussions 2, 1865–1892. https://doi.org/10.5194/hessd-2-1865-2005 koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research vol. 50 no. 1 pp. 54 60 applying economic guidelines for responsible tourism in a world heritage site abstract this article proposes an integrated sustainable tourism development strategy for the vredefort dome as a world heritage site. in particular, it analyses the integration of the south african guidelines for responsible tourism applied to a local community. all tourism product owners in the dome area formed part of this research that was conducted by means of a survey. the strategy that was formulated for the area aims to ensure that future generations will be able to utilise the resources in the same manner as the present generation. it is indicated that this intra-generational equity will only be possible through an integrated approach. the value of this research is its contribution to responsible tourism and, concurrently, its emphasis on the fact that tourism destinations in south africa need to be developed in an integrated and sustainable manner. in order for the vredefort dome to be a world heritage site, and to exist as a protected area, it is imperative that the tourism product owners in the area realise their roles as custodians of this precious site. this proposition leads to the main aim of this study, namely to develop an integrated tourism development strategy for the vredefort dome as a world heritage site. the results of the empirical study indicated that tourism stakeholders are prepared to improve the present situation in the vredefort dome and are willing to adjust their business practices in the future. such changes in practice involve some strategic issues, and the economic, social and environmental strategies will give direction to this world heritage site to become more sustainable. keywords: tourism, sustainable guidelines, integrated development, world heritage site, vredefort dome willem j. l. coetzee mathilda van niekerk department of tourism management tshwane university of technology south africa melville saayman institute for tourism and leisure studies north-west university (potchefstroom campus) south africa correspondence to: willem j. l. coetzee e-mail: coetzeewjl@tut.ac.za postal address: private bag x680, pretoria, 0001, south africa 54 this study focuses on the tourism stakeholders and their economic responsibility regarding sustainable tourism in the rural area of the vredefort dome. the south african guidelines for responsible tourism (social, environmental and economic) were used in the research although only the economic guidelines for responsible tourism will be reflected in this article. these guidelines were used to evaluate the present state of affairs and the stakeholders then had to predict the future importance of each guideline. based on the effect sizes, the most important aspects (according to the stakeholders) are that they will maintain and encourage social and cultural diversity as well as assess the economic impacts, as a prerequisite to developing tourism in the area. rural communities such as the vredefort dome (see figure 1) usually have few resources that can be developed, and because of this many communities are currently examining nontraditional means for economic diversification. in the vredefort dome it is, however, not a lack of natural resources that poses a problem – because the tourism potential of this area is well documented (iied, 2000; contour, 2002). according to reimold and coney (2001) more than seven hundred publications have appeared with some reference to the majestic vaal river and the scenic vredefort hills. it is for this reason that local policy-makers in rural communities such as this one realise the importance of tourism. although they have little or no experience regarding the development of tourism, many have nonetheless already grasped at opportunities for tourism development. however, do communities consider and monitor the effects of such development? a segment of the community in the vredefort dome has realised the importance of tourism, but it can be argued that there is a lack of integrated planning in this regard. according to briedenhann and wickens (2004), the lack of a strategic tourism plan in order to develop tourism in a sustainable manner, and also to alleviate poverty, is a general point of concern. the aim of this article is to report on current tourism practices in the vredefort dome area and to predict future practices in terms of the following: economic impacts as a prerequisite to developing tourism in the area willingness to maximise local economic benefits by increasing linkages and reducing leakages ensuring that the community is involved in, and benefits from, tourism responsible marketing and product development in the vfd equitable business opportunities in the vfd literature review authors such as inskeep (1991) have provided examples of how unfettered tourism growth can lead to detrimental impacts on the socio-cultural values of local residents, the economic diversity/development, and the environment. these impacts and their consequences have been discussed by a number of • • • • • 2008 economic guidelines for responsible tourism original research http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 54 60 55 authors as relevant in terms of economic growth, job creation, income for local governments (see also dwyer et al. 2003:431) and catalysts for other sectors such as agriculture, forestry, and fishing (curtin 2003:173; finkler & higham 2004) and the manufacturing industries, as well as infrastructure development for the local community (han et al. 2003:153). early research from other disciplines such as anthropology (smith 1977), geography and sociology also plays educational roles in the fields of different ideologies, culture, national heritage, the environment, political differences and cross-cultural attitudes towards an integrated development process. careful assessment and implementation of policy tools to minimise negative impacts on social attributes constitutes an important component of integrative tourism planning in rural settings. inskeep (1991) and many others call for integrated tourism planning within a broader regional context. the intent of integrative tourism planning is to provide for tourism development strategies in areas such as the dome that will be effective in enhancing the social, economic and environmental assets in the region and that simultaneously act to minimise the potential liabilities. by its nature, tourism brings outsiders into a local community. as with other development strategies, this global phenomenon is having a dramatic impact at the local level: some traditional cultures and economies are being modified to the extent that they are effectively destroyed (routledge 2002). furthermore, outsiders may eventually decide to move to the local community that once served as their original tourist destination. the literature in this regard has identified unique processes and methods that are specific to tourism planning (oliver & jenkins 2003:293). tourism development planning in rural areas has given rise to a growing body of academic literature (macdonald & jolliffe 2003:307). key unique attributes of more integrative tourism planning approaches include incorporating a broad mix of contemporary issues, initiatives, stakeholders, and objectively based data into the regional planning process. to conclude, this issue calls for a careful approach to tourism development that is integrated with overall regional goals. this approach is founded on murphey’s (1985:176) work on tourism development planning and calls those interested in sustaining rural communities to attention: “economic problems … have led many communities to consider embracing this growth industry [tourism] of the post-industrial era.” an integrated sustainable tourism strategy will take into account the social, economic and environmental issues as set out in agenda 21 – the foundation document of sustainable development. oliver and jenkins (2003:293) define integrated tourism as that which is explicitly linked to the localities in which it takes place and, in practical terms, has clear connections with local resources, activities, products, production service industries, and a participatory local community. as murdoch (1993:225) stated fifteen years ago, the implications of breaking down disciplinary boundaries are far-reaching but necessary to any meaningful research on sustainability. research method the population was identified after discussions with the tourism information centre in parys (the closest town to the dome) as well as the information centre in nearby potchefstroom. a list was compiled of all 43 tourism product owners in the vredefort dome area. ten stakeholders (n = 10) were identified as stakeholders that provide accommodation only. the majority (n = 21) of the respondents belong to the sector that provides accommodation and also act as operators in tourism activities such as rafting, hiking and team building. the hospitality sector comprised only five stakeholders (n = 5) and seven (n = 7) of the identified stakeholders are shop owners who deal mainly with tourists, for example antique, souvenir, and arts and handcraft shops. the forty-three questionnaires were hand-delivered in the area with a covering letter explaining the purpose of the study. a follow-up reminder was faxed after seven days to remind the respondents that the questionnaires would be collected the figure 1 aerial view of the vredefort dome – south africa source: http://www.unb.ca/passc/impactdatabase/images/vredefort.html a frican p rotected a rea c onservation and s cience original research coetzee, van niekerk & saayman vol. 50 no. 1 pp. 54 60koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za following day (eight days after delivery). during the collection phase a follow-up reminder with a copy of the questionnaire was left with each non-respondent to be collected after seven days. twenty-eight questionnaires were collected, representing an overall response rate of 60%. two of the questionnaires were discarded because respondents filled in all the “yes” answers in the yes/no category and all the 3’s on the likert scale. both of these respondents were shop owners and it was clear that they could not agree on all the environmental issues as indicated. finally, twenty-six questionnaires were used in the statistical analysis (n = 26). based on the south african guidelines for responsible tourism, a survey questionnaire was developed to examine respondents’ perceptions regarding the importance of sustainable tourism development and their vision for the future of the vredefort dome. in the guidelines for responsible tourism the criteria are divided into three sections: economic aspects, social aspects, and environmental aspects. the scale that was used entailed requesting two answers from respondents, namely: a yes/no answer (yes = in their organisation this guideline is already in practice; no = in their organisation they do not practice the guideline). a likert three-poi nt scale: (1 = not important, 2 = important and 3 = very important) to evaluate the view of product owners on how important they feel each aspect will be for the future. as the guidelines for responsible tourism have already been grouped into 12 subdivisions, it was not possible to perform a factor analysis of data – and furthermore, apart from that, the population that was selected for this study was not substantial enough for such an exercise. for the purpose of this study, therefore, the following formula was used to measure the differences for dependent data (steyn 2000): diff diff x d s = where diffx is the mean and diffs the standard deviation of each construct under each of the 12 subsections. the statistical analysis was performed by the statistical consultation services of north-west university by making use of the sas-programme (sas institute inc. 2001). results of the frequency analysis regarding economic aspects the results of frequency analysis regarding economic issues will be addressed under the following five methodological subsections: assessing economic impacts as a prerequisite to developing tourism in the vfd maximising local economic benefits – increasing linkages and reducing leakages ensuring that the community is involved in, and benefits from, tourism marketing and product development in the vfd equitable business opportunities in the vfd the discussion of each of the above five subsections is structured in a table that indicates the present situation (if tourism stakeholders apply the guidelines at present) as well as a future perspective (how important the guidelines will be in the future). both the present and the future perspectives are ranked according to importance (1 = the most important) in order to provide a clear indication of how priorities will change from the present towards a strategic future perspective. • • • • • • • assessing economic impacts as a prerequisite to developing tourism in the vfd from the literature it is clear that economic impacts are important in order to maximise the positive impacts of tourism and to minimise the negative ones. from table 1 it is clear that product owners recognise that tourism can create revenue from cultural heritage, traditional ways of life, wildlife and habitats (1.1.3) and they rated this aspect as the most important at present. to exercise a preference for business that directly benefits the local community (1.1.8) was rated second, which means that most of the product owners support local businesses. an interesting fact is that the product owners identified the historically disadvantaged as an emerging domestic tourism market (1.1.2). currently the product owners do not regard it as important to consider the adverse effects of tourism such as local land price inflation, and/or loss of access to resources (1.1.7), but this guideline is ranked as no.1 for the future. it is clear that the product owners regard this issue as imperative and therefore rated the guideline as the most important. losses of access to and land price inflation are well documented in the literature, but rural communities are not always aware of the negative impacts; they are rather blinded by the possible economic benefits. this issue needs to be addressed if the vfd would like to leave something of the present for future generations, otherwise the future generations of this area will not be able to enjoy the current way of living. it is a fact that if tourism development is not properly planned and developed it can lead to an increase in land prices and the area runs the risk of losing its unique character. inskeep (1991:372) states that: “... if residents cannot conveniently use their own facilities, they will become irritated and resentful of tourism”. the following aspects were also rated as important for the future in the vfd: the product owners would encourage business relationships between foreign entrepreneurs and local and emerging entrepreneurs (1.1.4). the product owners also realise that the historically disadvantaged as an emerging domestic tourism market (1.1.2) is an important aspect regarding future success. in 2002, the department of environmental affairs and tourism (deat) embarked on the first national domestic tourism survey to identify existing and potential domestic markets. to exercise a preference for business that directly benefits the local community (1.1.8) will also help to reduce leakage out of the community and will help to spread the economic benefits in the vfd. one point of concern that arises from table 1 is the fact that the product owners do not regard it as important at present or in the future to plan initiatives and investments to contribute to the broader local economic development strategy of the dome (1.1.6). after the survey, the economic development forum of the area 56 present assessing economic impacts as a pre-requisite future 0.851276 cronbach-alpha coefficient 0.585067 r % r % 4 54.69 1.1.1 extend the tourism season by developing new products 7 19.91 3 59.93 1.1.2 identify the historically disadvantaged as an emerging domestic market 3 35.47 1 82.47 1.1.3 recognise that tourism can create revenue from cultural heritage 6 22.40 5 50.06 1.1.4 encourage business relationships between foreign entrepreneurs 2 41.14 6 49.51 1.1.5 maintain and encourage economic diversity 5 28.23 7 47.20 1.1.6 plan initiatives and investments to contribute to local economy 9 9.02 8 45.82 1.1.7 consider adverse effects of tourism such as land price inflation 1 45.62 2 76.18 1.1.8 exercise a preference for businesses that benefit the community 4 30.32 6 49.51 1.1.9 conduct market feasibility before raising expectations 8 17.59 table 1 assessment of economic impacts as a prerequisite when developing tourism economic guidelines for responsible tourism original research http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 54 60 artists’ community has established an art route, and product owners that elect not to sell local art and handcraft at their establishments can assist by distributing maps of the route. in future, this guideline will help the vfd community to tap into new tourism markets, other than adventure tourism: namely those of art lovers and international craft buyers. maximising local economic benefits by increasing linkages and reducing leakages is a complicated issue. for the present and future generations it is essential for product owners to establish an amalgamated strategy without compromising the economic well-being of present establishments. from table 2 it is encouraging to see where the future increases will occur – in community enhancement and locally produced crafts and curios. ensuring that the community is involved in, and benefits from, tourism community involvement is an essential element of a successful strategy for sustainable tourism development. the product owners are willing to encourage visitors to spend more money in the local economy, bringing business to the local community (1.3.8) and to encourage tour operators to be more innovative in their itineraries, by, for example, including shebeens, local museums and art galleries (1.3.9). it is clear from the above that the product owners are prepared to encourage tourists to spend more money and to assist tour operators in programme planning. tour operators are not always aware of secondary attractions in an area and it is hence the responsibility of established product owners to spread the economic benefits. the vfd has established an economic development forum, which is an excellent way to start, so that businesses can become aware of each other and network in the business environment. the following are ideal guidelines, but an elected person in each organisation in the vfd needs to consider marketing, training and managerial support for promising tourism projects (1.3.6) and the product owners will hopefully consider using local entrepreneurs in developing community initiatives (1.3.10). some product owners are willing to consider establishing targets to monitor progress in achieving sustainable tourism objectives (1.3.12). the definitions of sustainable development are clear on the fact that it is a futureorientated concept and therefore it is essential that targets or indicators are needed to reach these objectives (bell & morse 1999). transparency is also important when reporting community 57 employed an economic development officer to facilitate the economic development idea that might lead to an enhanced economic strategy. maximising local economic benefits – increasing linkages and reducing leakages two important aspects of economic enhancement programming are firstly, to increase the linkages between tourism and other economic elements, for example agriculture, and secondly, to reduce leakage of money generated in the area. leakage is the purchasing power that is spent on imports to an area, resulting in a transfer of income out of the local economy (cook et al. 2002:299). the longer money circulates in the area, the more economic benefits will be generated. the vfd is suitable for becoming a self-sufficient area, and in this way jobs can be generated, which will subsequently lead to economic growth. in order to achieve a reliable measuring instrument, the following two guidelines were left out of the frequency analysis: (i) to buy locally-made goods and use locally-provided services from locally-owned businesses (1.2.3) and (ii) to cooperate with other formal sector businesses to maximise benefits for local community enterprises (1.2.5). the reason for the inconsistent rating of these two guidelines might be that the guidelines incorporate more than one issue per guideline or that the formulation of these guidelines is not acceptable for statistical analyses. for any future research it would be interesting to see how these two guidelines are rated by respondents. it is noteworthy that product owners would like to encourage all establishments to upgrade their standards of service and in that way maximise their revenue earning potential (1.2.1). other communities with similar ideas introduced a local grading system that forced the product owners to register with the local tourism information office, and in return the information office would have an idea of the services offered by different establishments. an essential and highly rated guideline is that the product owners must help the local community to develop its products so that the products can be more easily used by others (1.2.4). product development and distribution will lead to higher amounts spent by both tourists and by product owners in the area. this guideline was rated very highly for the future and it is encouraging to see that product owners seem willing to assist the local community with product development. give customers the opportunity to purchase locally produced crafts and curios (1.2.6) is an excellent way of generating jobs for the local artists and craftspeople. at present, the product owners feel that they are not doing enough to achieve this guideline, but it is encouraging to see how highly they rate the increase for the future. the vfd is well known for the number of craft shops and the density of artistic activity in the area. the vfd present maximising local economic benefits future 0.569278 cronbach-alpha coefficient 0.593118 r % r % 2 55.48 1.2.1 encourage all establishments to upgrade their standards 3 23.59 3 27.05 1.2.2 encourage the informal sector to become part of the formal sector 4 22.43 n/a 1.2.3 buy locally-made goods and use locally-made goods n/a 1 59.72 1.2.4 help the local community to develop its products 2 52.95 n/a 1.2.5 cooperate with other formal businesses to maximise benefits n/a 4 6.63 1.2.6 give customers the opportunity to purchase locally 1 54.44 table 2 maximising local economic benefits present community involvement in, and benefit from, tourism future 0.846828 cronbach-alpha coefficient 0.848138 r % r % 6 52.8 1.3.1 enable the historically disadvantaged to engage in tourism 11 11.35 2 68.2 1.3.2 work closely with local communities to develop new products 8 43.91 10 36.4 1.3.3 develop business partnerships in which the community has a significant stake 4 59.63 3 59.9 1.3.4 identify projects that can support the poor 2 82.63 5 57.2 1.3.5 assist in the development of local entrepreneurs 2 82.63 7 45.6 1.3.6 consider marketing, training and managerial support 10 16.52 9 37.5 1.3.7 foster the development of community-based tourism products 1 82.76 4 58.2 1.3.8 encourage visitors to spend more money in the local economy 9 24.59 8 44.9 1.3.9 encourage tour operators to be more innovative 6 51.34 4 58.2 1.3.10 consider using local entrepreneurs in community initiatives 5 57.10 1 70.3 1.3.11 be transparent when reporting community benefits 7 47.96 11 29.4 1.3.12 consider establishing targets to monitor progress 3 67.97 table 3 community involvement in, and benefit from, tourism a frican p rotected a rea c onservation and s cience original research coetzee, van niekerk & saayman vol. 50 no. 1 pp. 54 60koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za58 benefits (1.3.11) in order to distribute the benefits and minimise the effects where, otherwise, one establishment dominates the tourism development process and in that way keeps all the potential benefits to itself. for the future it is encouraging to see the commitment from product owners to identify projects that the enterprise can support that will benefit the poor (1.3.4). poverty alleviation is a global problem, and it is particularly in rural areas like the vfd that tourism can contribute to reducing this. assisting the development of local entrepreneurs with visitor feedback on their products (1.3.5) will allow the entrepreneurs to adapt their products to the market needs and in this way the product owners will foster the development of community-based tourism products by providing marketing and mentoring support (1.3.7) to entrepreneurs in the community. the community-based approach between entrepreneur and product owner should foster a close relationship that should lead to the development of new products that provide complementary products for tourism enterprises (1.3.2). especially the historically disadvantaged community will receive the opportunity to engage in the tourism sector (1.3.1). developing business partnerships in which the community has a significant stake (1.3.3) will also contribute towards economic enhancement of the broader community. marketing and product development providing information about local services and attractions provided in the dome, and encouraging tourists to use them (1.4.1) was rated as important to the product owners. the high rating that this aspect received is an indication that the product owners are committed to marketing the area to visitors. to consider cooperative advertising, marketing and the promotion of new and emerging products and attractions (1.4.2) was rated low, and for a community this small it might be a good idea to look into this opportunity. as the vfd is regarded by many as an adventure destination, it is not surprising to see the high rating that the product owners will foster the development of access opportunities for all visitors and potential visitors, regardless of physical or mental conditions (1.4.5). two factors that were not highly rated for the future (table 4) are to consider developing and marketing fairly traded tourism products (1.4.4) and to ensure that the visual way in which the product is presented includes local cultural elements (1.4.3). reasons for this might be that the product owners are already practising these two guidelines and one can thus expect them to rate these guidelines as low for the future. however, to rate fair trade tourism negatively might be regarded as a warning sign for the product owners. from table 4 it is clear that curios constitute an important aspect for the vfd, but the community should be made aware that they should guard against unfair curio practices such as child labour and environmental degradation that might occur from “unfair” products. from table 4 it is clear that the vfd needs an aggressive marketing strategy in which product owners cooperate with each other as well as with the local community. this will provide access to smaller niche markets such as the physically challenged tourists who are not able to participate in adventure activities. fair trade tourism products are part and parcel of ecotourism and are also closely related to sustainable development, and product owners cannot disregard this in the future. equitable business opportunities transparency and equity are central to sustainable development. as florini (1999) puts it: “... transparency is always closely connected to accountability, the purpose of calls for transparency is to permit citizens, markets or governments to hold others accountable for their polities and performances. thus, transparency can be defined as the release of information by institutions that is relevant to evaluating those institutions”. in table 5 equity and transparency are addressed and the results of the survey are presented in terms of how the product owners perceive these issues. to set targets for increasing the proportion of the enterprise wage bill going to community within 20 km of the enterprise (1.5.4) was most highly rated at present and also for the future (table 5). from this it is clear that product owners are currently employing locals where possible, but also regard it as an aspect in which the situation can be improved. employment is a global problem, especially in rural communities where opportunities are scarce. however, with the commitment of product owners, as shown in table 5, the problem might be reduced for future generations in the vfd. from table 5 it is clear that to develop transparent systems of sharing the benefits of tourism through equitable contracts. (e.g. tendering processes) (1.5.2) is rated relatively highly at present, and it is clear that the product owners feel that their current practices are effective. they also rated this guideline as important for the future, which indicates that there might be an increase in systems that will enhance the sharing of tourism benefits. one such system might be to advertise on the local website (http:// www.vredefortdome.co.za) any goods or services needed, and in that way local entrepreneurs might be able to render such services. an important consideration arises from the following, as entrepreneurs can be assisted to learn from their mistakes: when entering into agreements with the local community or emerging entrepreneurs ensure that the risk is equitably shared (1.5.3). if the product owners are willing to share the risk there might be an increase in entrepreneurial activities in the vfd area, and that will lead to the improvement of the community on many levels. product owners were aware of the issue of abusing their market power and imposing unfair commissions or pushing down prices inequitably (1.5.1), and in table 5 the product owners also indicated that they will do more in the future to minimise market domination. the guideline that states to go beyond the bare minimum wage rate and invest in local staff – quality is dependent upon well-motivated staff (1.5.6) was left out of the statistical analysis because of the inconsistency when product present marketing and product development future 0.640074 cronbach-alpha coefficient 0.498744 r % r % 3 35.88 1.4.1 provide information about local services and attractions 3 27.44 5 22.43 1.4.2 consider cooperative advertising, marketing and promotion 1 59.60 2 49.32 1.4.3 ensure that product presentation includes local cultural elements 4 8.41 1 71.66 1.4.4 consider developing and marketing fairly traded products 5 -0.52 4 22.76 1.4.5 foster the development of access opportunities for all visitors 2 51.12 table 4 marketing and product development present equitable business future 0.546882 cronbach-alpha coefficient 0.800850 r % r % 5 11.84 1.5.1 beware of abusing market power 2 65.38 2 43.03 1.5.2 develop transparent systems of sharing the benefits 3 52.03 3 38.48 1.5.3 ensure that the risk is equitably shared 5 39.13 1 44.66 1.5.4 set targets for wage bill to community within 20 km of the enterprise 1 88.07 4 19.83 1.5.5 develop a community labour agreement with targets 4 51.01 table 5 equitable business opportunities economic guidelines for responsible tourism original research http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 54 60 owners evaluated the guideline. the reason might be that the guideline addresses more than one issue, namely minimum wages and well-motivated quality staff. for future research, this guideline needs some contemplation in order to address the reliability quandary. from the results it is clear that the environmental aspects are regarded as the most important. product owners are committed to looking after the natural diversity in the area and to practise conservation strategies where possible. from an economic point of view it is clear that product owners are not always aware of the linkages between tourism and other economic activities in the area. for future benefits it is important that the product owners should practise activities that will reduce leakage out of the vfd and increase linkages with other economic partners. the social aspects were not rated as highly as one may have hoped for, but the future perspective looks prosperous. tourism in the vfd is in a relatively healthy state at present and product owners are committed to increasing their positive performances in the future. in the following section the measurement instrument that was used will be tested for reliability by making use of cronbachs alpha coefficients. use of cronbachs alpha reliability coefficient for subsections the cronbachs alpha reliability coefficient (cronbach 1951) was calculated to determine whether the questions in each subsection constituted reliable measuring instruments for that particular subsection. from the literature it is clear that instruments with a cronbachs alpha reliability coefficient of larger that 0.8 are very reliable, but that any instrument with a coefficient larger than 0.5 constitutes an acceptable measurement instrument (anastasi 1988:817). all except one of the subsections (marketing and product development) are reliable measuring statements. the future perspective regarding the marketing and product development aspects are the only subsections that are not reliable and it will be interesting to observe and compare results in this regard in the future. after the frequency analysis it was established that the measurement instrument is reliable, but the question remains: do the results have any significance to the product owners in the vfd? in order to measure the relevance of the results in such a small population it is necessary to measure practically significant differences between present and future opinions. the reason for this calculation is that the population that was used in the survey is too small to generalise the results to all tourism product owners in south africa. practically significant difference between the present and future opinions in order to determine if there is a difference between the product owners’ opinion regarding the importance of each guideline at present and for the future, the data for future importance was converted. opinions of “important” or “very important” were recoded as 1, while “not important” was recoded as 0. the mean difference between these recoded future and present opinions was determined. effect sizes were calculated to determine if there is a practically significant difference between the present and future opinions (see steyn 2000:2). effect sizes for dependent data (steyn 2000) for each subsection were calculated by making use of the following formula: diff diff x d s = 59 where diffx is the mean difference between future and present opinions and diffs the standard deviation of the difference between future and present opinions. cohen (1988) gives the following guidelines for the interpretation of the effect size in the current case: (a) small effect: d = 0.2, (b) medium effect: d = 0.5, (c) large effect: d = 0.8. we consider data with d ≥ 0.8 as practically significant, since it is the result of a difference having a large effect. these results are only practically significant for the population that participated in this survey. table 6 presents the practically significant difference between the present and future opinions of product owners in the vfd. from table 6 it is apparent that all the social aspects are significant. the present situation compared with the increase in the future is of importance to the product owners. some of the economic aspects are also practically significant, but none of the environmental aspects are significant. the reason for these environmental results is that the vfd is already developed and marketed as a nature-based tourism destination. product owners in the area are already environmentally sensitive and the difference between the present and the future is not viewed as significant to this community. thus, it is clear that if the vfd would like to be more sustainable, product owners should improve their social and economic aspects and continue with their environmental practices as at present. conclusions this paper presents the frequency analysis of the guidelines tested on the vfd. the analysis was based on the present situation, regarding how product owners currently conduct their activities, as well as a future perspective. the future perspective indicated how product owners perceive an increase or decrease of each guideline, not the importance of each guideline, but rather how they perceive the increase or decrease of the specific guideline to be. after the frequency analysis was tabulated and discussed in the text, the reliability of the measurement instrument was tested. the results of cronbachs alpha coefficients indicated that the instrument is reliable for the situation where it was utilised. the last part of the paper evaluated the practical significance of the results. in other words, the question as to whether the results mean anything for a community this small was addressed. because of the size of the population it would have been senseless to use means as a measurement criterion and therefore the results were evaluated by using a special formula suited for this particular situation. based on the results presented it is possible to formulate a strategy for sustainable tourism development in the vredefort dome area as a world heritage site. for sustainable tourism development in the vredefort dome the following strategies are proposed on how tourism stakeholders can assist to achieve sustainability: economic aspects diffx diffs effect size 1 assessing economic impacts as a prerequisite to developing tourism in vfd 0.333 0.327 1.018 2 maximising local economic benefits 0.271 0.410 0.661 3 ensuring that community is involved in and benefits from tourism 0.347 0.454 0.764 4 marketing and product development 0.233 0.262 0.889 5 equitable business 0.417 0.429 0.972 table 6 practically significant difference between the present and future opinions a frican p rotected a rea c onservation and s cience original research coetzee, van niekerk & saayman vol. 50 no. 1 pp. 54 60koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za60 follow an integrated approach to the planning and management of land resources. this implies that all stakeholders should work together and that the development of the area should take into consideration the economic, social and environmental impacts. in doing so, poverty alleviation can also be addressed in a sustainable manner. stakeholders in the area include government, communities, product owners and investors. protect and promote human health in the vfd. promote education and training in the vfd. promote sustainable human settlement development in the vfd area. promote sustainable agriculture and rural development. establish a system that guarantees supply that is in line with fair trade in tourism. reduce leakages and other purchasing practices. the results indicated that tourism stakeholders are willing to improve the present situation in the vredefort dome and to practise their businesses in a different way in the future. these changed practices entail that some strategic issues will need to be addressed, and the strategies mentioned above will give direction to this world heritage site to become more sustainable. references anastasi, a. 1988. cronbachs alpha psychological testing (6th ed.). new york: macmillan. bell, s. & morse, s. 1999. sustainability indicators: measuring the immeasurable. london: earthscan. briedenhann, j. & wickens, e. 2004. tourism routes as a tool for the economic development of rural areas – vibrant hope or impossible dream? tourism management, 25(1): 71–79, february. cohen, j. 1988. statistical power analysis for the behavioural sciences (2nd ed.). hillsdale: erlbaum. contour project managers. 2002. strategic development and management plan for the vredefort dome. johannesburg: grant thornton kessel feinstein. cook, r.a., yale, l.j. & marqua, j.j. 2002. tourism – the business of travel (2nd ed.). london: prentice hall. cronbach, j.l. 1951. coefficient alpha and internal structure of tests. psychometrika, 16: 297–334. curtin, s. 2003. whale-watching in kaikoura: sustainable destination development? journal of ecotourism, 2(3): 173– 195, december. deat – see south africa dwyer, l., forsyth, p., spurr, r. & vanho, t. 2003. tourism’s contribution to a state economy: a multi-regional general equilibrium analysis. tourism economics, 9(4): 431–448, december. • • • • • • • finkler, w. & higham, j. 2004. the human dimensions of whale watching: an analysis based on viewing platforms. human dimensions of wildlife, 9(2): 103–117, january. florini, a.m. 1999. does the invisible hand need a transparent glove? the politics of transparency. paper prepared for the annual world bank conference on development economics, washington, dc, 28–30 april 1999. available: http://www. worldbank.org/research/abcde.papers.html [accessed: 5 september 2002]. han, q., dellaert, b.g.c., van raaij, f. & timmermans, h.j.p. 2003. supporting tourist activity planning decisions from an urban tourism management perspective. tourism analysis, 8(2): 153–157. iied see international institute for environment and development. inskeep, e. 1991. tourism planning: an integrated and sustainable development approach. new york: van nostrand reinhold. international institute for environment and development. 2000. rural planning in south africa: a case study. london: iied. macdonald, r. & jolliffe, l. 2003. cultural rural tourism evidence from canada. annals of tourism research, 30(2): 307–322, april. murdoch, j. 1993. sustainable rural development: towards a research agenda. geoforum, 24(3): 25–241. murphey, p.e. 1985. tourism: a community approach. new york: routledge. oliver, t. & jenkins, t. 2003. sustaining rural landscapes: the role of integrated tourism. landscape research, 28(3): 293– 307, july. reimold, w.u. & coney, l. 2001. the vredefort dome impact structure and directly related subjects: an updated biography. johannesburg: economic geology research institute, university of the witwatersrand. routledge, p. 2002. resisting and reshaping the modern: social movements and the development process. in johnston, r.j., taylor, p. j., watts, m.j. & blackwell, o., (eds.). geographies of global change. remapping the world in the late twentieth century. london: blackwell publishing. p. 544. sas institute inc., 2001. the sas system for windows release 8.02 ts level 02m0 copyright© 1999–2001 by sas institute inc., cary: usa. smith, v.l. 1977. hosts and guests: the anthropology of tourism. philadelphia: university of pennsylvania press. south africa. 1996. department of environmental affairs and tourism. the development of tourism in south africa. pretoria: government printer. south africa. 2002. department of environmental affairs and tourism. guidelines for responsible tourism. pretoria: government printer. steyn, h.s. (jr.). 2000. practical significance of the difference in means. journal of industrial psychology, 26(3): 1–3. abstract introduction site description and methodology results and discussions conclusion acknowledgements references about the author(s) johan van tol department of soil, crop and climate sciences, faculty of science and agriculture, university of the free state, bloemfontein, south africa stefan julich institute of soil science and site ecology, technical university of dresden, dresden, germany darren bouwer department of soil, crop and climate sciences, faculty of science and agriculture, university of the free state, bloemfontein, south africa edward s. riddell department of conservation management, south african national parks, skukuza, south africa centre for water resources research, university of kwazulu-natal, pietermaritzburg, south africa citation van tol, j., julich, s., bouwer, d. & riddell, e.s., 2020, ‘hydrological response in a savanna hillslope under different rainfall regimes’, koedoe 62(2), a1602. https://doi.org/10.4102/koedoe.v62i2.1602 note: special issue: connections between abiotic and biotic components of a granite catena ecosystem in kruger national park, sub-edited by beanelri janecke and johan van tol. original research hydrological response in a savanna hillslope under different rainfall regimes johan van tol, stefan julich, darren bouwer, edward s. riddell received: 11 oct. 2019; accepted: 22 apr. 2020; published: 29 oct. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract soil water is a link between precipitation and the functioning of ecological systems. it is therefore critical to understand exactly how soil water regimes are affected by changes in precipitation. this is especially true for the variable water regimes of savanna ecosystems. therefore, understanding the effects of precipitation on soil water was the central goal of this article. the hydropedological behaviour of a catena in the stevenson hamilton research supersite of the kruger national park was configured as a conceptual model of catchment modelling framework, a toolbox of various model structures and processes. the model was parameterised using measured hydraulic properties of the soils, and calibrated and validated using measured soil matric potentials and derived actual evapotranspiration (aet) data. the model was then used to simulate hydrological response under five different rainfall scenarios, ranging from 30% drier than the normal rainfall to 30% wetter than the normal rainfall. the scenarios also included rainfall years with fewer but larger rain events, that is, more intense rainfall events. in general, the model performed well with pearson’s correlation coefficient (r) values ranging between 0.66 and 0.87 and between 0.58 and 0.69 for correlations with daily soil matric potential and daily aet, respectively. scenario analysis indicates non-linearity in the response of hydrological processes to changes in precipitation. this is especially evident in a seven-fold increase in the duration of saturation at the seepage surface associated with a 30% increase in rainfall. in general, the impact of drying conditions (30% below average rain) has a greater influence on soil water contents, overland flow and percolation from the riparian zone to bedrock than a 30% increase in rainfall would have on the same process. conservation implications: this article presents realistic predictions of the potential impact of changes in precipitation on hydrological processes in an important area of the kruger national park. these predictions would enable decision-makers to be prepared for the anticipated changes in near-surface hydrological processes associated with climate changes. keywords: hydropedology; hydrological modelling; soil water; catchment modelling framework; kruger national park. introduction savannas are currently experiencing extensive population and agricultural water resource pressure globally (rockström et al. 2014). because we are now moving into an era of adaptive management of ecosystems where information on state change thresholds is a prerequisite for their sustained management, it is necessary to invest in understanding the biophysical processes that maintain these systems. in savanna landscapes, soil water is the direct link between precipitation and ecological patterns (weltzin et al. 2003). the co-domination by trees and grasses is bound by soil water availability, a key factor in establishing form and function. therefore, spatial soil moisture dynamics is a crucial link in the equilibrium between climate, soil and vegetation in these systems (rodriguez-iturbe et al. 1999). the resulting hydrological processes contribute to the biophysical template of these semi-arid systems, controlling the distribution of water and other resources along a continuum within the landscape, which often comprises compound effects of non-linear relationships and threshold-triggered responses. given this complexity, novel interdisciplinary approaches should be sought to understand hydrological processes in such heterogeneous landscapes (troch et al. 2008). moreover, inter-disciplinarity is increasingly valuable for successful landscape management, given the emphasis on hydrological connectivity at landscape scale (michaelides & chappell 2009). the general framework that process hydrologists have used is to garner knowledge of water flow pathways and residence times, as this is essential for predicting the response of a catchment to rainfall (uhlenbrook, wenninger & lorentz 2005). it is suggested that hydropedology is a vital interdisciplinary science for contributions to the earth’s critical zone (lin 2009). whilst the fields of pedology and hydrology have long been mutually exclusive, there is a need for the soil scientist to benefit from flow theory when transcribing qualitative descriptions into quantitative expressions, and vice versa, for the hydrologist to develop representative pedotransfer functions in hydrological modelling (bouma 2006). for instance, ticehurst et al. (2007) demonstrated that marrying these two disciplines, most notably using soil colour and presence of redox concretions, was effective in indicating depths of saturation and lateral flow occurring on hillslopes in new south wales, australia. testing their conceptual flow path model against hillslope hydrometric observations yielded a satisfactory agreement, but they warned that further catchment information should be sought to reduce model uncertainty. meanwhile, sivapalan (2003) asked why watershed hydrological responses were seemingly simple and hillslope processes comparatively complex, suggesting that by aggregating hillslope complexities into dominant processes, one could parameterise the hillslope as the basic unit within a catchment model. ridolfi et al. (2003) highlighted these hillslope complexities, including inter alia the spatial heterogeneity of soil properties; climatic variability (although generally uniform at the hillslope scale, may trigger mechanisms within the hillslope to alter its spatial dynamics); lateral redistribution of water along a hillslope because of the formation of a saturated zone within the soil; lateral sub-surface flow in the unsaturated zone; the longitudinal form and profile of a hillslope; and boundary conditions at the valley bottom of the hillslope. it has been advocated that the use of perceptual hydrological models, whilst being largely qualitative conceptualisations, offers good potential based on process understanding of key zones or ‘reservoirs’, and that these ‘soft’ data could be married to ‘hard’ hydrological observations (streamflow and soil water content) to facilitate the reduction in parameter uncertainty (bouwer et al. 2015; lorentz et al. 2003; seibert & mcdonnell 2002). arid and semiarid regions of the world are highly dependent on the availability of water, affecting recruitment, growth and reproduction, nutrient cycling and net ecosystem productivity. quantifying the potential consequences of changing precipitation patterns is important in these regions. it is expected that varying regional precipitation regimes will have important ramifications for the distribution, structure, composition and diversity of plant, animal and microbe populations and communities (easterling et al. 2000; houghton et al. 2001; weltzin & mcpherson 2003). although some effects have been estimated, for instance, increasing precipitation variability – but not necessarily the total amount of precipitation – may reduce grassland productivity and livestock-carrying capacity, exacerbate overgrazing and increase rangeland susceptibility to invasive weed species (knapp et al. 2002). given the above rationale, the following three primary objectives are addressed in this article: (1) to configure and parameterise a hillslope-scale hydrological model using soil and environmental data by bouwer et al. (2020); (2) to calibrate and/or validate the model using hydrometrical measurements presented in riddell et al. (2020); and (3) to use the calibrated model to simulate hillslope hydropedological processes under different rainfall scenarios. site description and methodology site characteristics and soil information a granitic catena (hillslope) of a third-order stream in the southern granite supersite (smit et al. 2013) in the kruger national park (knp), south africa, was the focus area in this article. the mean annual precipitation is 560 mm/a (smit et al. 2013), and the geological formation is granite and gneiss of the nelspruit suite (venter 1990). vegetation on the crest to upper-midslope positions is dominated by woody vegetation such as combretum apiculatum and combretum zeyheri. a prominent seep line exists on the transition between the midst to footslope, which is marked by terminalia sericea. below the seep line, there is a long sodic site frequented by euclea divinorium. along the footslope, fine-leaved woody species such as vachellia nilotica dominates (smit et al. 2013). bouwer et al. (2020) conducted a hydropedological survey where 49 soil profiles were classified and interpreted in terms of their hydropedological behaviour (figure 1). this conceptual model of hillslope hydropedological behaviour served as a basis for both the model set-up and identification of dominant soils and horizons for undisturbed sampling for soil physical measurements. watermark sensors were installed in representative soil horizons (see riddell et al. 2020). figure 1: hydropedological behaviour inferred from soil distribution pattern, different landscape units and location of watermark sensors on the studied hillslope. four modal profiles were identified based on terrain position and soil type. these modal profiles were sampled at 100-mm depth intervals for particle size distribution using the pipette method and chemical analysis. soil hydraulic properties were determined on five of the dominant horizons (orthic a, yellow-brown apedal b, prismacutanic b, melanic a and pedocutanic b). the saturated hydraulic conductivity of each diagnostic horizon in the modal profiles was measured in duplicate using a modified bouwer and rice (1976) double-ring falling-head method: where ks is the saturated hydraulic conductivity (mm hr−1), l is the thickness of the horizon, t is the time until constant infiltration and h0 and h1 are the head of water above the surface before and after time t, respectively. water retention characteristics were measured for each horizon on undisturbed cores. the samples were de-aired at –70 kpa at room temperature for 24 h. water was then allowed to gradually fill the cores. the saturated samples were mounted on a hanging water column set-up in accordance with the procedure of dirksen (1999). the suction level (h) was set at 0, 38, 50, 100, 200, 400, 600, 800 mm to capture important lower suctions of the retention curves. the gravimetric water content was determined at each level. the samples were then oven-dried at 105°c to determine the bulk density to calculate the volumetric water content (θv). the measured bulk density, water retention measurements, together with the particle size distribution, was used to derive van genuchten parameters in the retention curve (retc) program (van genuchten, leij & yates 1991). these parameters were used as inputs in the hydrological model. model configuration and parameterisation the catchment modelling framework (cmf; kraft et al. 2011) was used to construct the framework for the hydrological modelling of this hillslope. the cmf is essentially a toolbox to configure a wide range of different model structures based on the finite volume approach. water fluxes through the landscape are represented as a network of storages and boundary conditions in cmf. flux-governing equations could be assigned to link the storage units with the next one. these equations can either be fairly simple, for instance, linear storage or tipping bucket approaches, or more complex, for instance, solving of kinematic wave or richard’s equation. the compounds of the model are assembled using the scripting language python. here we used the open-source platform, spyder version 3.3.3, as the environment for scientific programming in python (python software foundation 2019). we divided the 480-m long slope into 24 cells, each covering 2 000 m2. these cells were 100-m wide and stretched 20 m downslope. the elevation profile was determined with a differential global positioning system (gps; riddell et al. 2020). the hydraulic parameters of soils (table 1), and their spatial distribution presented by bouwer et al. (2020: figure 1), guided the spatial distribution of the hydraulic parameters (figure 2). the vertical discretisation within each cell was 10 cm and the richard’s equation was used to estimate vertical flow through profiles. overland flow was calculated using the kinematic wave approach. water ponding on the surface flows to the next downslope cell, from where it is allowed to re-infiltrate the soil. the thickness of horizons, as observed in the soil survey, was included in the set-up. below the soils, a 2 m thick ‘bedrock’ layer was included in the model set-up. the hydraulic properties below each of the cells were adjusted as part of the calibration, but in general it reflected the conceptual understanding of ‘recharge’ of groundwater in the crest and riparian zone, and limited deep drainage in the midslope positions. below the bedrock layer, free drainage into a groundwater store was allowed. figure 2: hillslope configuration and hydrological parameters/processes employed in catchment modelling framework for the simulations. table 1: summary of climatic parameters used in simulations; total rainfall and monthly averages of temperature, humidity, wind and radiation. an atmospheric boundary condition was applied to the surface of each cell. measured climatic parameters (riddell et al. 2014), including rainfall, minimum and maximum temperatures, relative humidity, solar radiation and wind speed, were included in the simulation on a daily time-scale. a summary of these parameters is presented in table 1. potential evapotranspiration (et0) was calculated from the climatic parameters using the penman–monteith equation, and the shuttleworth–wallace approach was used to distinguish between actual evaporation and transpiration (shuttleworth & wallace 1985). the simulation period commenced on 01 may 2011 and ended on 13 january 2013. evaluation of the model outputs commenced on 03 november 2011 and 10 november 2011 for evaporation and soil water contents, respectively, because of the availability of reliable data. this allowed the model to ‘settle’ for approximately six months prior to evaluation. evaluation and calibration of the model soil matric pressures were measured at different horizons of the four sites on the hillslope (figure 1) and converted to daily average values. details on the instrumentation and calibration of these measurements are discussed in more detail in riddell et al. (2014) and riddell et al. (2020). actual evapotranspiration (aet) was obtained from van eekelen et al. (2015) using the surface energy balance algorithm for land (sebal) model (bastiaanssen et al. 1998). the weekly measurements were then converted into daily values for different landscape positions using the approach reported in riddell et al. (2020). measured daily soil matric pressure and daily sebal-derived aet values were used in a manual calibration process. in this process, parameters for which no measured values were available have been varied stepwise in a defined range (table 2). after each simulation, the pearson’s correlation coefficient (r) values and root mean square errors (rmse) between measured and simulated values were calculated. the calibrated parameters comprise the conductivity and porosity of the bedrock layers as well as rooting depth and leaf area index (lai) of the respective vegetation types. for example, if the simulated water contents of sub-soils accumulated more than what was measured, the conductivity and porosity of the bedrock was increased. the aim of the calibration was not simply to improve model outputs but to maintain the measured parameters and make the unmeasured parameters (i.e. bedrock hydraulic parameters, rooting depth and lai) more realistic and congruent with existing data for the region and reduce parameter uncertainty. table 2: calibration ranges and final values for selected model parameters. precipitation scenario analysis the calibrated model was then used to simulate hydrological processes under five different precipitation scenarios. an average rainfall year (583 mm), represented by the 2009–2010 rain season (i.e. august to july), was identified from the riddell et al. (2014) database. this rainfall year represented the ‘average’ scenario, or the baseline. the rainfall was then manually changed to reflect a ‘dry year’ by reducing the measured daily rainfall by 30%. similarly, a ‘wet year’ was created by increasing the measured rainfall by 30%. ‘dry extreme’ and ‘wet extreme’ scenarios were created by lumping the rainfall of consecutive rainfall days (i.e. >1 mm day−1) and using this value as the daily rainfall on the first day of the rainfall sequence whilst deleting rainfall on the other days. this was carried out to reflect an increase in rainfall intensity and also longer dry periods. these adjustments were applied to both ‘dry’ and ‘wet’ years to create ‘dry extreme’ and ‘wet extreme’ scenarios. a summary of this climatic data is presented in table 3. table 3: average monthly values for precipitation scenarios. the soils were initially dry for the model runs, with a saturated zone 10 m below the lowest node of the bedrock; then, different yearly climate inputs were repeated twice. the first year was to allow a settling period for the model and the second year for evaluation purposes. the simulation results presented are centred on the key hydrological processes as presented in figure 1, namely, storage of water in the crest zone, seepage above the sodic site, overland flow in the midslope, and recharge and storage in the riparian zone. ethical considerations the full ethical clearance statement was obtained from the interfaculty animal ethics committee of the university of the free state – clearance number ufs-aed2019/0121. results and discussions hydraulic parameters of the different soils the crest is dominated by freely drained clovelly (cv) soils consisting of an orthic a (ot) horizon on a yellow-brown apedal (ye) b horizon. these soils are marked by high hydraulic conductivity (> 50 mm h−1; table 4). in the upper-midslope, an albic horizon (gs) was formed, predominantly because of lateral movement on the soil–bedrock interface. the difference in conductivity between the albic horizon and impermeable bedrock (table 4) promotes the generation of lateral flow and the formation of albic properties (van tol, hensley & le roux 2013). table 4: van genuchten parameters of dominant soil horizons. the conductivity of both the top soil and sub-soils of the sodic site (see the orthic horizon above prismacutanic [pr] b horizon – table 4) restricts vertical infiltration, especially during high-intensive rain events, and promotes the generation of overland flow. the well-developed structure of the melanic a horizon (ml) of the bonheim soils in the footslope is likely to allow infiltration and storage of water (figure 1) with the pedocutanic (vp) also allowing drainage towards the bedrock. in the valley bottom, the dundee (du) soils have the same hydropedological behaviour as the clovelly soil in the crest (bouwer et al. 2020). the ot and the stratified alluvium (al) horizons of the dundee form were consequently assigned similar hydraulic properties as that of ot and ye of the clovelly soil, respectively (table 4). infiltration and vertical drainage are the dominant hydrological processes in the valley bottom. model validation pearson’s correlation coefficients between measured and simulated soil matric potentials were acceptably high (when compared to similar studies in the same area – see van tol et al. 2015) for most of the soil horizons (table 5), especially in the crest and the upper-midslope. this suggests that there is a good (positive) correlation between measured and simulated changes in the soil water content. pearson’s correlation coefficient values obtained in a previous modelling study for the same area for the riparian soil ranged between 0.42 and 0.75 (van tol et al. 2015), rendering >0.7 r-values of this exercise acceptable. the lower correlation coefficients in the sterkspruit soil of the midslope could be attributed to macropore flow dominant between the structure units of the prismacutanic b horizon. this bypass flow is not simulated well with the model. the rmse values between measured and simulated matric, however, are also high, suggesting that the absolute measured matric potentials are not predicted very well. because the measurements of soil matric potential are also an estimation, albeit a calibrated estimation, there is a degree of error in these measurements. future work should focus on direct measurements of soil water content (gravimetric measurements) to validate modelling and to confirm the accuracy of, or recalibrate, the watermark sensors. table 5: correlations between measured and simulated matric potentials (kpa) of different soil horizons along the catena. the sebal-derived aet and simulated aet values correspond relatively well for all landscape units during the first part of the simulations (figure 3) (note that the cumulative graphs in figure 3 are not of consecutive days, as there were gaps in the sebal-derived aet values. the corresponding simulated values were also omitted from the statistics). a relative under-simulation of sebal-derived aet compared to simulated aet is observed towards the end of the simulation period. the differences could be explained in the dynamic nature of vegetation, which flourishes after wet periods. this is not simulated properly by the model, hence the underestimation. in theory, it implies that parameters such as lai and root depth should also be dynamic and change with the changes of actual vegetation characteristics during a simulation. for this modelling exercise, the relatively high r and low rmse values were deemed acceptable for establishing the impacts of changing precipitation patterns in the next section. figure 3: cumulative surface energy balance algorithm for land-derived actual evapotranspiration and cumulative-simulated actual evapotranspiration for: (a) the crest, (b) the footslope and (c) the valley bottom of the studied hillslope together with (d) corresponding correlation coefficients and root mean square error of daily actual evapotranspiration (mm day−1). precipitation scenarios a summary of the impacts of the different precipitation scenarios is presented in table 6. it is important to note that there is non-linearity between different processes and properties. for example, a 30% decrease in rainfall will result in an average decrease of around 12% in the water content of the a horizon of the soils of the crest, but a 30% increase in precipitation will only yield an average increase of around 5% in the water content of the same horizon. this non-linearity is more pronounced in complex processes such as overland flow. here, a 30% decrease in rainfall will result in a 67% decrease in overland flow volume, whereas a 30% increase will only cause a 17% increase in the volume of overland flow during a rainfall year (table 6). table 6: summary of impacts of different precipitation scenarios on key hydrological properties and processes. values are followed by percentage differences from ‘average’ scenario. soil water contents of selected soil horizons figure 4 presents exceedance curves of soil water contents of various soil horizons under different precipitation scenarios. sub-soil horizons of both the clovelly and dundee soils have higher water contents, for longer, than surface horizons. this is presumably because of the impact of evaporation on surface horizons. in general, the water content is considerably lower for the dry and dry extreme scenarios when compared with average or wet scenarios. the duration curves of the ‘average’ scenario are more closely associated with wet scenarios than dry ones. this could imply that a drying climate will have a greater impact on soil water contents than a wetting climate (see also differences in the average water content presented in table 6). in the simulations, soil water content affects evapotranspiration; therefore, higher soil water content has a higher potential to produce evapotranspiration. for this reason, the impact of an increase in rainfall patterns on soil water contents is not as clear as the impact of a decrease in rainfall. figure 4: duration of exceedance curves of soil water contents of (a) orthic a horizon and (b) yellow-brown apedal b horizon of the clovelly soil on the crest position; and (c) orthic a horizon and (d) stratified alluvium b horizon of the dundee soil in the valley bottom. there is very little difference between the ‘dry’ and ‘dry extreme’, and ‘wet’ and ‘wet extreme’ scenarios in all the simulations. this is likely because of relatively high hydraulic conductivity of all horizons in the clovelly and dundee soils (table 4). an increase in rainfall intensity will therefore have little impact on infiltration in the soils when simulated with a daily time-step. simulations indicate that the dundee soil will be wetter than the clovelly soil (figure 4). this despite higher simulated evapotranspiration from the dundee soil (figure 3) and similar hydraulic properties of the two soils (table 4). overland flow from the midslope to the dundee soil contributes to higher water contents in this soil. seepage line functioning under the ‘dry’ and ‘dry extreme’ scenarios, a positive matric pressure (i.e. seepage) was simulated for one day only: a decrease of 85% compared to the ‘average’ (table 6). the ‘wet’ and ‘wet extreme’ scenarios could result in fivefold and sevenfold increase, respectively, in the duration of saturation at seepage line. the lateral flux volume in the profile under the ‘dry’ and ‘dry extreme’ simulations will decrease by 45% and 31%, respectively, whilst the ‘wet’ year will increase lateral flux by 66% and a ‘wet extreme’ scenario by 122%. lower fluxes associated with a drying climate could result in the accumulation of salts at the seepage line and reduce the spreading of salts over the sodic site. a wetting climate, on the other hand, could result in excessive leaching of salts and nutrients from the slope, and hinder the normal functioning of the sodic site. overland flow in midslope the functioning of the seepage line impacts the soils downslope. owing to drier soils and lower than average rain events, overland flow is likely to decrease between 65% and 67% for ‘dry extreme’ and ‘dry’ scenarios, respectively. it appears that under dry conditions, the soil water content has a greater influence on overland flow generation than the nature of precipitation events (i.e. only a difference of 2%). this assumption is, however, only applicable to the daily time-step, which was used in these simulations; using a finer time-step (e.g. hourly), the rainfall intensity might be higher than the infiltration rates of the soils and could produce significant overland flow. under wet conditions, the rainfall intensity has a much stronger influence (18% increase from ‘wet’ to ‘wet extreme’ scenario). under ‘wet’ and ‘wet extreme’ scenarios, the seepage line produces more return flow (66% and 122%, respectively), and the surface horizons of the soils lower in the hillslope transect are expected to be close to saturation. large rain events are then more likely to generate significant overland flow when compared to the drier antecedent soil water conditions expected under the dry climate. percolation from riparian zone soils the assumption is that large rain events are more likely to generate significant groundwater recharge than a number of smaller events (riddell et al. 2020). our simulations however indicate that the highest volume of percolation from the dundee soil of the riparian zone will occur during the ‘wet’ scenario (33% increase) (note that percolation in this sense does not necessarily imply recharge into the groundwater but merely exfiltration into the bedrock layer below the soil). the ‘wet extreme’ scenario only resulted in an increase of 17% (table 6). a possible reason for this is that the simulations based on matrix flow do not take macropore flow into account; the latter is often dominant during extreme events. this is also evident in the percolation duration curves (figure 5). percolation from the ‘dry’ and ‘dry extreme’ scenarios will decrease by 49% and 53%, respectively, which will impact groundwater levels negatively. figure 5: duration of exceedance curves of percolation (mm day−1) from the lowest node of the dundee soil in the riparian zone under different precipitation scenarios. conclusion in this article, we applied pedology and soil physics to quantify the hydrological response of a hillslope under different rainfall scenarios. the quantification of hydrological processes based on this hydropedological approach is important for an improved understanding of the functioning of the hydrological cycle and contribution towards better resource management in the savanna ecosystems. the cmf model was relatively accurate in predicting soil matric potentials and aet. the model structure could, however, be improved by including a dual porosity module as well as time-dependent vegetation parameters (e.g. lai and rooting depth). the findings presented suggest that hillslope processes in savanna ecosystems are especially vulnerable to a drying climate. it also highlights non-linearity in the response of hillslope hydrological processes to changes in rainfall. this non-linearity is caused not only by non-linearity in the soil-water retention function of soils but also by the connectivity of landscape. for example, the seepage line would remain saturated seven times longer, with a 30% increase in rainfall because of increased lateral flow from upslope positions. similarly, drainage from the riparian zone decreases by more than 50%, with a 30% decrease in rainfall because of lower overland flow volumes received from upslope. it is important that resource managers and policymakers in the savanna ecosystems understand and quantify this non-linearity and the influence of hydrological connectivity at different scales. higher or lower rainfall cannot be simplistically translated as higher or lower recharge, streamflow, runoff, soil water storage and seepage. future work should investigate whether these hillslope-scale processes could be extrapolated to catchment and landscape scale. different processes (e.g. groundwater contributions and return flow from streams) are associated with higher-order streams, which influence the sensitivity of hydrological processes to changing rainfall regimes. it would also be important to establish the impact of changes in hydrological processes on other ecosystem components through interdisciplinary research. acknowledgements competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions j.v.t. was responsible for modelling and compiling the first draft of the manuscript. d.b. and e.s.r. conducted the fieldwork and improved the first draft. s.j. assisted with the modelling and the final write-up of the manuscript. funding information this work was partially funded by the university of the free state under the ‘multi-disciplinary program’ and the water research commission (wrc) of south africa under project k5/2051. data availability data are available and may be provided, on request, by the corresponding author. data from all research done within kruger national park is placed within the sanparks repository (not for free, open access). disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references bastiaanssen, w.g.m., menentia, m., feddesb, r.a. & holtslagc, a.a.m., 1998, ‘a remote sensing surface energy balance algorithm for land (sebal) 1. formulation’, journal of hydrology 198–212, 212–213. https://doi.org/10.1016/s0022-1694(98)00253-4 bouma, j., 2006, ‘hydropedology as powerful tool for environmental policy research’, geoderma 131, 275–286. https://doi.org/10.1016/j.geoderma.2005.03.009 bouwer, h. & rice, r.c., 1976, ‘a slug test for determining hydraulic conductivity of unconfined aquifers with completely or partially penetrating wells’, water resource research 12(3), 423–428. https://doi.org/10.1029/wr012i003p00423 bouwer, d., le roux, p.a., van tol, j.j. & van huyssteen, c.w., 2015, ‘using ancient and recent soil properties to design a conceptual hydrological response model’, geoderma 241, 1–11. https://doi.org/10.1016/j.geoderma.2014.10.002 bouwer, d., le roux, p.a.l. & van tol, j., 2020, ‘identification of hydropedological flow paths in stevenson–hamilton catena from soil morphological, chemical and hydraulic properties’, koedoe 62(2), a1584. https://doi.org/10.4102/koedoe.v62i2.1584 dirksen, c., 1999, soil physics measurements, catena verlag, reiskirchen. easterling, d.r., meehl, g.a., parmesan, c., changnon, s.a., karl, t.r. & mearns, l.o., 2000, ‘climate extremes: observations, modeling, and impacts’, science 289(5487), 2068–2075. https://doi.org/10.1126/science.289.5487.2068 houghton, j.t., ding, y., griggs, d.j., noguer, m., van der linden, p.j., dai, x. et al., 2001, climate change 2001: the scientific basis, cambridge university press, cambridge. knapp, a.k., fay, p.a., blair, j.m., collins, s.l., smith, m.d., carlisle, j.d. et al., 2002, ‘rainfall variability, carbon cycling, and plant species diversity in a mesic grassland’, science 298(5601), 2202–2205. https://doi.org/10.1126/science.1076347 kraft, p., vaché, k.b., frede, h.g. & breuer, l., 2011, ‘cmf: a hydrological programming language extension for integrated catchment models’, environmental modelling & software 26(6), 828–830. https://doi.org/10.1016/j.envsoft.2010.12.009 lin, h.s., 2009, ‘earth’s critical zone and hydropedology: concepts, characteristics, and advances’, hydrology and earth systems science discussions 6, 3417–3481. https://doi.org/10.5194/hessd-6-3417-2009 lorentz, s., thornton-dibb, s., pretorius, j.j. & goba, p., 2003, ‘hydrological systems modelling research programme: hydrological processes, phase ii: quantification of hillslope, riparian and wetland processes’, report no. k5/1061, water research commission, pretoria. michaelides, k. & chappell, a., 2009, ‘connectivity as a concept for characterising hydrological behaviour’, hydrological processes 23(3), 517–522. https://doi.org/10.1002/hyp.7214 riddell, e.s., nel, j., fundisi, d., jumbi, f., van niekerk, a. & lorentz, s., 2014, ‘ephemeral hydrological processes in savannas’, report no. tt 619/14, water research commission, pretoria. riddell, e.s., nel, j., van tol, j., fundisi, d., jumbi, f., van niekerk, a. et al., 2020, ‘groundwater–surface water interactions in an ephemeral savanna catchment, kruger national park’, koedoe 62(2), a1583. https://doi.org/10.4102/koedoe.v62i2.1583 ridolfi, l., d’odorico, p., porporato, a. & rodriguez-iturbe, i., 2003, ‘stochastic soil moisture dynamics along a hillslope’, journal of hydrology 272(1–4), 264–275. https://doi.org/10.1016/s0022-1694(02)00270-6 rockström, j., falkenmark, m., folke, c., lannerstad, m. & barron, j., 2014, water resilience for human prosperity, cambridge university press, cambridge. rodriguez-iturbe, i., d’odorico, p., porporato, a. & ridolfi, l., 1999, ‘tree-grass coexistence in savannas: the role of spatial dynamics and climate fluctuations’, geophysical research letters 26(2), 247–250. https://doi.org/10.1029/1998gl900296 seibert, j. & mcdonnell, j.j., 2002, ‘on the dialog between experimentalist and modeler in catchment hydrology: use of soft data for multicriteria model calibration’, water resources research 38(11), 1241–1255. https://doi.org/10.1029/2001wr000978 sivapalan, m., 2003, ‘process complexity at the hillslope scale, process simplicity at the watershed scale: is there a connection?’, hydrological processes 17(5), 1037–1041. https://doi.org/10.1002/hyp.5109 shuttleworth, w.j. & wallace, j.s., 1985, ‘evaporation from sparse crops – an energy combination theory’, quarterly journal of the royal meteorological society 111(469), 839–855. https://doi.org/10.1002/qj.49711146910 smit, i.p.j., riddell, e.s., cullum, c. & petersen, r., 2013, ‘kruger national park research supersites: establishing long-term research sites for cross-disciplinary, multiscaled learning’, koedoe 55(1), art. #1107, 7 pages. https://doi.org/10.4102/koedoe.v55i1.1107 spyder version 3.3.3., 2019, computer software, python software foundation, viewed 21 june 2019, from https://www.spyder-ide.org/. ticehurst, j.l., cresswell, h.p., mckenzie, n.j. & glover, m.r., 2007, ‘interpreting soil and topographic properties to conceptualise hillslope hydrology’, geoderma 137(3–4), 279–292. https://doi.org/10.1016/j.geoderma.2006.06.016 troch, a.p., carillo, g.a., hiedbuchel, i., rajgopal, s., switanek, m., volkmann, t.h.m. et al., 2008, ‘dealing with landscape heterogeneity in watershed hydrology: a review of recent progress toward new hydrological theory’, geography compass 3(1), 375–392. uhlenbrook, s., wenninger j. & lorentz s., 2005, ‘what happens after the catchment caught the storm? hydrological processes at the small, semi-arid weatherley catchment, south africa’, advances in geosciences 2, 237–241. https://doi.org/10.5194/adgeo-2-237-2005 van eekelen, m.w., bastiaanssen, w.g.m., jarmain, c., jackson, b., ferreira, f., van der zaag, p. et al., 2015, ‘a novel approach to estimate direct and indirect water withdrawals from satellite measurements: a case study from the incomati basin’, agriculture, ecosystems & environment 200, 126–142. https://doi.org/10.1016/j.agee.2014.10.023 van genuchten, m., th., leij, f.j. & yates, s.r., 1991, the retc code for quantifying the hydraulic functions of unsaturated soils, version 1.0, computer software, us salinity laborotory, california, ca. van tol, j.j., hensley, m. & le roux, p.a.l., 2013, ‘pedological criteria for estimating the importance of subsurface lateral flow in e-horizons of south african soils’, water sa 39(1), 47–56. https://doi.org/10.4314/wsa.v39i1.7 van tol, j.j., van zijl, g.m., riddell, e.s. & fundisi, d., 2015, ‘application of hydropedological insights in hydrological modelling of the stevenson hamilton research supersite, kruger national park, south africa’, watersa 41(4), 525–533. https://doi.org/10.4314/wsa.v41i4.12 venter, f.j., 1990, a classification for land management planning in the kruger national park, phd thesis, department of geography, university of south africa, pretoria. weltzin, j.f., loik, m.e., schwinning, s., williams, d.g., fay, p.a., haddad, b.m. et al., 2003, ‘assessing the response of terrestrial ecosystems to potential changes in precipitation’, bioscience 53(10), 941–952. https://doi.org/10.1641/0006-3568(2003)053[0941:atrote]2.0.co;2 weltzin, j.f. & mcpherson, g.r., 2003, changing precipitation regimes and terrestrial ecosystems: a north american perspective, university of arizona press, tuscon, az. abstract introduction methods results discussion conclusion acknowledgements references about the author(s) jody m. barends department of biodiversity and conservation biology, university of the western cape, cape town, south africa darren w. pietersen department of zoology and entomology, university of pretoria, pretoria, south africa guinevere zambatis scientific services, kruger national park, skukuza, south africa donovan r.c. tye organization for tropical studies, kruger national park, skukuza, south africa bryan maritz department of biodiversity and conservation biology, university of the western cape, cape town, south africa citation barends, j.m., pietersen, d.w., zambatis, g., tye, d.r.c. & maritz, b., 2020, ‘sampling bias in reptile occurrence data for the kruger national park’, koedoe 62(1), a1579. https://doi.org/10.4102/koedoe.v62i1.1579 original research sampling bias in reptile occurrence data for the kruger national park jody m. barends, darren w. pietersen, guinevere zambatis, donovan r.c. tye, bryan maritz received: 25 july 2019; accepted: 24 feb. 2020; published: 11 may 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract to effectively conserve and manage species, it is important to (1) understand how they are spatially distributed across the globe at both broad and fine spatial resolutions and (2) elucidate the determinants of these distributions. however, information pertaining to the distributions of many species remains poor as occurrence data are often scarce or collected with varying motivations, making the resulting patterns susceptible to sampling bias. exacerbating an already limited quantity of occurrence data with an assortment of biases hinders their effectiveness for research, thus making it important to identify and understand the biases present within species occurrence data sets. we quantitatively assessed occurrence records of 126 reptile species occurring in the kruger national park (knp), south africa, to quantify the severity of sampling bias within this data set. we collated a data set of 7118 occurrence records from museum, literature and citizen science sources and analysed these at a biologically relevant spatial resolution of 1 km × 1 km. as a result of logistical challenges associated with sampling in knp, approximately 92% of knp is data deficient for reptile occurrences at the 1 km × 1 km resolution. additionally, the spatial coverage of available occurrences varied at species and family levels, and the majority of occurrence records were strongly associated with publicly accessible human infrastructure. furthermore, we found that sampled areas within knp were not necessarily ecologically representative of knp as a whole, suggesting that areas of unique environmental space remain to be sampled. our findings highlight the need for substantially greater sampling effort for reptiles across knp and emphasise the need to carefully consider the sampling biases within existing data should these be used for conservation management decision-making. modelling species distributions could potentially serve as a short-term solution, but a concomitant increase in surveys across the park is needed. conservation implications: the sampling biases present within knp reptile occurrence data inhibit the inference of fine-scale species distributions within and across the park, which limits the usage of these data towards meaningfully informing conservation management decisions as applicable to reptile species in knp. keywords: conservation management; kruger national park; occurrence data; reptile fauna; sampling bias. introduction effective conservation and management of organisms require an understanding of how species are spatially distributed at both broad and fine spatial resolutions, and ideally also the underlying determinants of their distribution patterns (hurlbert & jetz 2007; kery 2011). however, species geographic data that may help inform conservation management decisions are often limited and biased in their collection strategies (franklin 2010). for example, although museum databases often include occurrence data of collected specimens, the principal purpose of most museum collections is to act as reference catalogues for species identification rather than for species distribution mapping (newbold 2010). it is important to note that although several museum specimens are collected directly as a result of systematic sampling, many specimens are collected opportunistically (kadmon, farber & danin 2004; pyke & ehrlich 2010). as a result, collection effort and spatial coverage within museum data naturally vary depending on the interests of the collection. despite this, a recently increased urgency in the need for species distribution information has placed a greater emphasis on the use of museum databases for amassing species occurrence records (syfert, smith & coomes 2013). in recent years, the capture of museum data within electronic databases, the establishment and continued activities of atlasing projects, and the growth of citizen science projects have provided a wealth of species occurrence data that are accessible online (newbold 2010). these data are undoubtedly valuable but are subject to a multitude of biases, errors and uncertainties that need to be considered should these data be used for environmental research. generally, species occurrence records are susceptible to geospatial or taxonomic sampling biases and on their own do not explain the full extents of species distributions (bird et al. 2014; botts, erasmus & alexander 2011; reddy & davalos 2003). for example, museum data are often biased towards heavily sampled areas (newbold 2010), atlas data tend to be vulnerable to omission errors (botts et al. 2011) and citizen science records are often imprecise (geldmann et al. 2016; mcgrath et al. 2015). for rare and understudied species, bias in occurrence data sets exacerbates an already severe issue of misinformation and overall data deficiency (reddy & davalos 2003). with recorded occurrences of these animals already limited, the presence of an assortment of sampling biases within databases further restricts our understanding of these species’ distributions and curtails our ability to manage them effectively. for cryptic species such as some species of reptiles (bates et al. 2014; mcgrath et al. 2015), there is often a distinct lack of high-quality records of these animals’ occurrences within their natural environments (böhm et al. 2013; tolley et al. 2016), even within areas specifically designated for conservation (ferreira et al. 2011; venter et al. 2008; zielinski 2001). in south africa, the kruger national park (knp) is home to approximately 126 reptile species (bates et al. 2014; branch 1998; pienaar 1978). the presence of reptiles promotes ecological diversity within knp, and more broadly southern africa, as many reptile species are likely to have important ecological roles or carry out ecologically beneficial functions within a variety of habitats and ecosystems (trimble & aarde 2014). overall, reptiles comprise approximately 14% of vertebrate species within knp (parr, woinarski & pienaar 2009), and the conservation of these animals is essential for maintaining diversity within this important protected area (gascon et al. 2015; parr et al. 2009; venter et al. 2008). the knp biological reference collection houses thousands of preserved specimens across a wide variety of taxa and includes hundreds of individual reptiles collected within the park over the past 80 years. the collection also includes an extensive electronic database that catalogues each specimen along with its respective biological and locality information where available. this collection places knp among the best sampled protected areas in south africa (and probably in africa) for reptiles (bates et al. 2014). however, the very nature of such reference collections is that sampling intensity and objectives vary over time, with earlier sampling efforts focused primarily on compiling inventory lists and collecting reference material. as such, the knp biological reference collection database for reptiles was never intended as a systematic survey across all habitats and reflecting all patterns of occurrence within the park. in recent years, however, the need for spatially explicit species occurrence data sets to inform modern conservation tools requires that the data from biological reference collections should be co-opted into conservation analyses. accordingly, there is a need to critically evaluate such existing data sets to understand any inherent patterns of bias they possess. in this study, we (1) collate and synthesise available occurrence data for reptile species in knp from reference collections, museum databases and literature sources; (2) assess patterns of geographic and taxonomic biases within this data set; and (3) evaluate whether areas of spatial bias are environmentally representative of knp as a whole, including under-sampled regions. methods reptile occurrence data we collated reptile locality and occurrence data from literature sources, museum and reference collection databases, a virtual museum platform, citizen science sightings from social media platforms and field data gathered under various teaching, monitoring and inventorying exercises by the organization for tropical studies (table 1). additionally, two of the authors (j.m.b. and b.m.) provided 151 novel records from personal observations in knp (listed as ‘this study’). in total, we collated 14 533 records, but after georeferencing these to match locality descriptions and removing duplicates across sources, we had a final data set of 7118 records representing 126 reptile species occurring in knp. this data set is available upon request from sanparks scientific services. table 1: sources of reptile occurrence records used in this study. coverage biases we summarised reptile species occurrence data to identify geographic and taxonomic biases in coverage across knp. by carrying out regression analyses, we tested if reptile families were evenly represented across knp by comparing the relationship between the number of occurrences for each reptile family to the extent of the geographical areas (in km2) surrounding those occurrences of each reptile family (i.e. the area of the minimum convex polygon enclosing all occurrences). next, we assessed if residual values of each reptile family fell within the range of mean ± standard deviation of all residual values to identify which families were spatially overor under-represented (i.e. above or below the range) across knp in our sample data set. to evaluate the proportion of knp for which reptile occurrence data exist and quantify the extent of unsampled areas, we divided knp into equal-sized grid cells at 1 km × 1 km, 2 km × 2 km, 4 km × 4 km and 9 km × 9 km (pentad scale) resolutions, respectively. these resolutions allowed us to identify patterns of geographic sampling bias across a range of biologically appropriate spatial resolutions. however, the 1 km × 1 km resolution was preferred for most analyses. this resolution subjectively offered the best trade-off between the spatial error associated with historical records of occurrence data (newbold 2010) and the relatively small spatial scale at which many reptiles utilise landscapes (fischer, lindenmayer & cowling 2004; price, kutt & mcalpine 2010). we plotted reptile occurrences across the grid cells of each spatial resolution by using quantum geographic information system (qgis) version 3.4 (qgis development team 2018) and counted the number of occurrences per grid cell. by carrying out regression analyses, we also tested whether a relationship exists between the numbers of reptile occurrences recorded within each grid cell and the proximity of those grid cells to the nearest publicly accessible infrastructure areas of knp (defined here as all camps, gates, picnic sites and public roads) at the finest spatial resolution (1 km × 1 km). are sampled areas representative of the kruger national park as a whole? we downloaded environmental and infrastructural data layers at a spatial resolution of 1 km × 1 km to represent the overall environmental space of knp. these included 20 bioclimatic layers representing current climate (1970–2000) and elevation from the worldclim database (http://www.worldclim.org), soil type classifications of south africa from the international soil resource and information centre (isric; https://www.isric.org), vegetation type classifications of south africa from the south african national biodiversity institute (sanbi; www.bgis.sanbi.org) and infrastructural layers for publicly accessible camps, gates, picnic sites and roads within knp from south african national parks (sanparks; http://dataknp.sanparks.org/sanparks). we also generated ‘slope’, ‘aspect’ and ‘distance to water bodies’ layers for knp by using arcgis version 10.4 (esri 2016), resulting in a total of 27 representative layers for the environmental space of knp. to reduce the effects of spatial autocorrelation between layers, we performed a principal component analysis by using r verswion 3.5.3 (r core team 2018) to summarise the layers into 27 new, uncorrelated principal component layers. we retained the first six principal component layers as representatives of the overall environmental variability of knp as they cumulatively represented 85% of all modelled variation, which we selected as an effective stopping point as per jackson (1993). we tested if sampled areas within knp (i.e. grid cells containing at least one reptile occurrence) were environmentally representative of knp as a whole. to do this we separated each of our six principal component layers into new layers representing (1) sampled areas and (2) unsampled areas and compared the kernel density estimates of each pair per component via six separate two-sample kolmogorov–smirnov tests. ethical considerations this article followed all ethical standards for a research without direct contact with human or animal subjects. results summary of occurrence data our database contained 7118 reptile occurrence records, unevenly distributed across 60 lizard species, 59 snake species, 6 testudine species and 1 crocodylian species (table 2). as such, the majority of occurrences were of squamates (lizards: 48% of all records; snakes: 41% of all records), with the less speciose testudine and crocodilian groups having less representation (8% and 3% of all records, respectively). this was not the case at the species level where the nile crocodile (210 records) and the leopard tortoise (232 records) ranked only below the rainbow rock skink (242 records) for species with the highest numbers of occurrence records in our data set. the number of occurrence records per reptile family was positively related to the number of species per said family (linear regression analysis: f1, 17 = 28.45, p < 0.01, r2 = 0.63). the uneven distributions of records across reptile families were likely present as a product of collection bias and the specific combination of species occurring within knp rather than being solely because of collection bias on its own. table 2: summary of records of reptile occurrences within the kruger national park. coverage biases representation based on spatial coverage was unevenly distributed across reptile families in knp. we found a significant positive relationship between the cumulative number of records and the cumulative extents of the areas encompassing records of each reptile species per reptile family (linear regression analysis: f1, 17 = 76.60, p < 0.01, r2 = 0.81). we identified reptile families that appeared to be significantly under-represented (lacertidae, leptotyphlopidae and typhlopidae) and those that were over-represented (crocodylidae and scincidae) geographically across knp (figure 1), providing evidence of taxonomic sampling bias at the family level within our data set. figure 1: linear regression residuals demonstrating coverage biases across reptile families within the kruger national park. families with positive residuals occur more frequently than expected based on family-level species richness, although families with negative residuals are considered under-sampled. at the biologically appropriate spatial resolution of 1 km × 1 km, we found that only 1751 of 21 761 grid cells (8%) contained any reptile occurrence records at all (figure 2). moreover, 52% of these grid cells contained only a single record (911 grid cells; figure 3). we found that as the numbers of records per grid cell increased, the numbers of grid cells containing records decreased (regression analysis: f1, 114 = 9.34, p < 0.01, r2 = 0.08). this pattern held true at resolutions of 2 km × 2 km and 4 km × 4 km, respectively, but was not present at 9 km × 9 km (figure 3), demonstrating that geographic sampling bias is strongest at fine resolutions but weakens as resolution becomes coarser. we also found a significant relationship between the number of records present within each grid cell and its proximity to publicly accessible human infrastructure within the park (regression analyses: public roads ‒f1, 1749 = 6.75, p < 0.01, r2 = 0.06; camp sites and picnic spots ‒f1, 1749 = 9.01, p < 0.01, r2 = 0.10; figure 4). as the distance to infrastructure increased, the frequency of recorded reptile occurrences per grid cell significantly decreased, providing evidence of sampling bias towards publicly accessible areas. figure 2: spread of reptile occurrence records across the kruger national park at a 1 km × 1 km spatial resolution. figure 3: frequency distributions of the numbers of kruger national park reptile occurrence records per grid cell at 1 km × 1 km, 2 km × 2 km, 4 km × 4 km and 9 km × 9 km resolutions. figure 4: distances of grid cells (1 km × 1 km) containing records to nearest publicly accessible infrastructure (camp sites, picnic spots and public roads) within the kruger national park. environmental representation of sampled areas at the spatial resolution of 1 km × 1 km, sampled areas of knp were not representative of the full range of environmental space of knp as a whole (figure 5). the results of six separate kolmogorov–smirnov tests showed that there were significant differences in environmental variability between sampled and unsampled areas across each of the six principal components representing the overall environmental space of knp (d = 0.06–0.27, p < 0.01 in all cases). grid cells containing records of reptile occurrences were thus not statistically representative of the overall ecological variability of knp. figure 5: overlap in environmental variation between sampled and unsampled grid cells within the kruger national park at the 1 km × 1 km resolution for (a) principal component 1, (b) principal component 2, (c) principal component 3, (d) principal component 4, (e) principal component 5, and (f) principal component 6. bars beneath kernel density plots represent the range of environmental variation between sampled and unsampled grid cells. discussion at a fine spatial resolution ecologically relevant to reptiles, occurrence data for reptile species in knp are geographically and taxonomically biased. as a consequence of overall data deficiency, representation within our reptile occurrences data set varied, with highly detectable reptile species and families having had significantly more records than those with comparatively lower detectability. moreover, the majority of reptile occurrence data were associated with human infrastructure. approximately 68% of all records occurred in close proximity (< 2 km) to publicly accessible human infrastructure areas in knp. unsurprisingly, grid cells associated with major tourist camps and surrounding areas were considerably better sampled than the remainder of the park. importantly, sampled areas were not representative of the complete range of environmental variability across knp. this suggests that regions of the park that comprised unique environmental space are not represented in the current data set. spatial sampling biases associated with human infrastructure are common in biological sampling data sets (newbold 2010). most notably, presence-only data sets derived from atlas projects, citizen science data and museum records are typically susceptible to geographic bias in collection effort (botts et al. 2011; geldmann et al. 2016; reddy & davalos 2003; zielinski 2001;). geographic bias in collection effort is often present within these data sets as a result of sampling being inhibited in certain areas but facilitated in others (botts et al. 2011; pyke & ehrlich 2010). for example, some areas may be difficult to sample because of extreme weather conditions, rough terrain, the presence of dangerous animals, distance from roads or restricted access (bird et al. 2014; freitag et al. 1998). conversely, other areas facilitate more complete sampling by providing ease of access and associated increased visitation. in this context, sampling intensity within certain areas is likely to be dramatically lower in comparison with that of less restrictive areas that offer greater accessibility. this is certainly the case in knp where publicly accessible infrastructural areas have increased human presence and accessibility from staff and visitors alike in comparison with the remainder of the park. consequently, our data set seemingly represents areas of high sampling intensity rather than true biological patterns. areas of high sampling intensity seldom represent the full range of environmental space and ecological factors associated with determining species distributions (tolley et al. 2016). several studies have found substantial differences in climate between welland under-sampled areas (see botts et al. 2011; kadmon et al. 2004; martinez & wool 2006; reddy & davalos 2003; stockwell & peters 1999), with many of these highlighting the significance of climatic biases towards assessments of the true biological distributions of species. botts et al. (2011) found that sparse sampling effort in areas away from human infrastructure resulted in incomplete representations of amphibian distributions across south africa. here, where we have encountered similar geographical sampling biases with our reptile data set to those encountered by botts et al. (2011), we similarly conclude that our data set is unlikely to reflect the complete range of real-world distributions of reptile species across knp. the biased nature of our data set has important implications for sanpark’s management of reptiles in knp. despite being perhaps the most comprehensive collation of reptile occurrence data for knp to date, the use of this data set for informing robust conservation management decisions would need to be considered with caution. because of the large variation in geographical sampling intensity within our data set and the associated biases within the underlying occurrence data, it would be inappropriate to use this data set in its current form within the context of spatial planning for species conservation management. because a large proportion of our data was not collected explicitly for the purposes of mapping species distributions, the biological patterns as presented within our data set may confound comparisons among species, or comparisons of a particular species’ abundance in different habitat types, across environmental gradients, or across time series (bird et al. 2014; fischer et al. 2004). however, should the geographic biases be minimised or reversed without concurrently increasing other forms of bias (botts et al. 2011), this data set could become an important resource for knp conservation management. minimising the biases within our data set could be achieved by targeted sampling of data-deficient areas in knp. although the majority of 1 km × 1 km grid cells in knp could benefit from supplemental sampling, priority should be given to grid cells that contain no data and are distant from publicly accessible areas that are steadily subject to human visitation. in particular, we recommend that the mopane woodlands-dominated north-western region of the park (i.e. areas demarked as ecozones p and p1 as per sanparks 2016) should be targeted for additional sampling. the majority of this region is poorly sampled, with most of its grid cells containing no reptile records. moreover, this region has few public roads and is largely lacking in human visitation. here, we recommend that an approach emulating the karoo biogaps project led by sanbi (main et al. 2019) should be implemented, in which specific grid cells are selected as sites in which to extensively collect occurrence records on the basis of a statistical sampling design. this method of grid cell selection would involve the use of a statistical algorithm (such as latin hypercube sampling) that seeks to maximise coverage across the region although minimising the total number of grid cells to be sampled. traversal to grid cells targeted for sampling could be facilitated through the use of management roads unavailable to the general public. together with public tourist roads, this offers the widest range of vehicle access across knp. although these roads do not cover the full extent of knp, their usage can alleviate some of the challenges associated with inaccessibility for many grid cells and offers a feasible option towards future sampling campaigns. systematically sampling for reptile occurrences in targeted and supplementary areas within knp will substantially improve upon the overall coverage and comprehensiveness of available data. it is important to note however that the challenges associated with reptile sampling, such as low detectability (mcgrath et al. 2015), may result in underestimations of species richness and occurrences at specific sites. false absences as a result of underestimations can falsely inform on species’ performance within monitoring frameworks, including those relating to thresholds of potential concern, and may result in incorrect assignments of conservation priority (botts et al. 2011; ferreira et al. 2011). compiling complete inventories for targeted grid cells will thus be vitally important, but this may require several sampling trips to ensure that comprehensive species lists are compiled. such an approach would be unavoidably costly, time-consuming and could delay investigations of the statuses of reptile species within the park. in the meantime, other options are available to fill gaps in sampling within knp reptile data. over the last two decades, an increasing number of studies have used species distribution models (sdms) to extrapolate spatially explicit predictions of the distributional ranges of species (bird et al. 2014; stockwell & peters 1999). species distribution models predict environmental suitability for species, which can be used to infer species’ presence or absence within a given area (guisan et al. 2013; hurlbert & jetz 2007; kery 2011). these types of models are typically referred to as ecological niche models as the predictions produced are based on statistical relationships between species occurrences and environmental descriptor variables (guisan et al. 2013; kadmon et al. 2004). importantly, studies have found that sdms based on biased data with limited occurrences can produce strong models with accurate predictions (e.g. pearson et al. 2007; syfert et al. 2013), if the underlying biases are corrected for during model production and high-quality predictor variables are available. studies that aim to identify sources of data bias, particularly within presence-only data sets, offer invaluable insights into bias correction within the context of sdms (syfert et al. 2013). by understanding the sources of bias, it may be possible to correct historical and current population distributions modelled through sdm frameworks using mathematically inferred or experimentally determined bias correction factors. a good example of this is the use of visibility bias correction factors on aerial census data to improve the accuracy of geographical distributions and population size estimates of large herbivores in knp (see redfern et al. 2002). potential sdm frameworks for knp reptile species should seek to correct for the proximity of reptile occurrences to publicly accessible areas within the park. overcoming the challenges associated with this bias may require innovative solutions; however, if implemented correctly, such a framework could offer a feasible approach towards obtaining meaningful reptile distribution information for use within conservation management in knp. conclusion we sought to collate occurrence records for knp reptile species and provide a quantitative assessment of the sampling biases within these data. we have shown that at biologically relevant resolutions, knp is largely data deficient for reptile occurrences, with existing data being geographically biased towards publicly accessible areas. we further show that sampled areas were not environmentally representative of knp as a whole and from this, we conclude that our data set does not provide a true reflection of real-world reptile species distributions across knp. because the majority of the data within our database were not explicitly collected with species mapping in mind, additional sampling is needed to reverse the biases present. we recommend that future sampling efforts should target historically poorly sampled regions in the park that are distant from publicly accessible locations. finally, we suggest that in the meantime sdms may offer a more feasible approach for use within conservation management decision-making relating to reptile species within knp. acknowledgements the authors thank the following institutions for providing reptile occurrence data: the animal demography unit (university of cape town), the ditsong national museum of natural history, organization for tropical studies (ots) and sanparks. we acknowledge and thank all of the various collectors for their valuable contributions towards expanding the the knp reference collection over the last 80 years. they further thank ots, including bernard coetzee and laurence kruger for providing logistical support and assistance in the field. they also thank robin maritz for input towards the discussion. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions b.m. conceptualised this study and obtained data provided by d.w.p., d.r.c.t. and g.z. j.m.b. collated the data and performed all analyses. j.m.b. and b.m. co-wrote the manuscript, with additional assistance from d.r.c.t. funding information the authors acknowledge the ada and bertie levenstein foundation, merseta and the national research foundation (nrf grant no. 99186) for providing funding that facilitated the completion of this study. data availability statement reptile occurence data used in this study are available from south africa national parks. disclaimer all the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references bates, m.f., branch, w.r., bauer, a.m., burger, m., marias, j., alexander, g.j. et al., 2014, suricata 1: atlas and red list of the reptiles of south africa, lesotho, and swaziland, south africa national biodiversity institute, pretoria. bird, t.j., bates, a.e., lefcheck, j.s., hill, n.a., thomson, r.j., edgar, g.j. et al., 2014, ‘statistical solutions for error and bias in global citizen science datasets’, biological conservation 173(1), 144–154. https://doi.org/10.1016/j.biocon.2013.07.037 böhm, m., collen, b., baillie, j.e., bowles, p., chanson, j., cox, n. et al., 2013, ‘the conservation status of the world’s reptiles’, biological conservation 157(1), 372–385. https://doi.org/10.1016/j.biocon.2012.07.015 botts, e.a., erasmus, b.f.n. & alexander, g.j., 2011, ‘geographic sampling bias in the south african frog atlas project: implications for conservation planning’, biodiversity conservation 20(1), 119–139. https://doi.org/10.1007/s10531-010-9950-6 branch, w.r., 1998, field guide to snakes and other reptiles of southern africa, struik, cape town. esri, 2016, arcgis desktop 10.4, environmental systems research institute, redlands, ca, viewed 28 february 2018, from https://www.arcgis.com/index.html. ferreira, s., deacon, a., sithole, h., bezuidenhout, h., daemane, m. & herbst, m., 2011, ‘from numbers to ecosystems and biodiversity: a mechanistic approach to monitoring’, koedoe 53(2), 1–12. https://doi.org/10.4102/koedoe.v53i2.998 fischer, j., lindenmayer, d.b. & cowling, a., 2004, ‘the challenge of managing multiple species at multiple scales: reptiles in an australia grazing landscape’, journal of applied ecology 41(1), 32–44. https://doi.org/10.1111/j.1365-2664.2004.00869.x franklin, j., 2010, mapping species distributions: spatial inference and prediction, cambridge university press, cambridge. freitag, s., hobson, c., biggs, h.c. & van jaarsveld, a.s., 1998, ‘testing for potential survey bias: the effect of roads, urban areas and nature reserves on a southern african mammal data set’, animal conservation forum 1(2), 119–127. https://doi.org/10.1111/j.1469-1795.1998.tb00019.x gascon, c., brooks, t.m., contreras-macbeath, t., heard, n., konstant, w., lamoreux, j., et al., 2015, ‘the importance and benefits of species’, current biology 25(10), r431–r438. https://doi.org/10.1016/j.cub.2015.03.041 geldmann, j., heilmann-clausen, j., holm, t.e., levinsky, i., markussen, b., olsen, k. et al., 2016, ‘what determines spatial bias in citizen science? exploring four recording schemes with different proficiency requirements’, diversity and distributions 22(11), 1139–1149. https://doi.org/10.1111/ddi.12477 guisan, a., tingley, r., baumgartner, j.b., naujokaitis-lewis, i., sutcliffer, p.r., tulloch, a.i.t. et al., 2013, ‘predicting species distributions for conservation decisions’, ecology letters 16(12), 1424–1435. https://doi.org/10.1111/ele.12189 hurlbert, a.h. & jetz, w., 2007, ‘species richness, hotspots, and the scale dependence of range maps in ecology and conservation’, proceedings of the national academy of sciences 104(33): 13384–13389. https://doi.org/10.1073/pnas.0704469104 jackson, d.a., 1993, ‘stopping rules in principal components analysis: a comparison of heuristical and statistical approaches’, ecology 74(8), 2204–2214. https://doi.org/10.2307/1939574 jacobsen, n.g.h., 1989, ‘a herpetological survey of the transvaal’, phd thesis, dept. of biological sciences, university of natal. kadmon, r., farber, o. & danin, a., 2004, ‘effects of roadside bias on the accuracy of predictive maps produced by bioclimatic models’, ecological applications 14(2), 40–413. https://doi.org/10.1890/02-5364 kery, m., 2011, ‘towards the modelling of true species distributions’, journal of biogeography 38(4), 617–618. https://doi.org/10.1111/j.1365-2699.2011.02487.x main, d., tensen, l., gihring, k., bronner, g., aboul-hassan, n., blanckenberg, m. et al., 2019, ‘unravelling the taxonomy and distribution of two problematic small mammal genera in the karoo biome’, african zoology 54(3), 125–135. https://doi.org/10.1080/15627020.2019.1628661 martinez, j.j.l. & wool, d., 2006, ‘sampling bias in roadsides: the case of galling aphids on pistacia trees’, biodiversity & conservation 15(7), 2109–2121. https://doi.org/10.1007/s10531-004-6685-2 mcgrath, t., guillera-arroita, g., lahoz-monfort, j.j., osborne, w., hunter, d. & sarre, s.d., 2015, ‘accounting for detectability when surveying for rare or declining reptiles: turning rocks to find the grassland earless dragon in australia’, biological conservation 182(1), 53–62. https://doi.org/10.1016/j.biocon.2014.11.028 newbold, t., 2010, ‘applications and limitations of museum data for conservation and ecology, with particular attention to species distribution models’, progress in physical geography 34(1), 3–22. https://doi.org/10.1177/0309133309355630 parr, c.l., woinarski, j.c.z. & pienaar, d.j., 2009, ‘cornerstones of biodiversity conservation? comparing the management effectiveness of kruger and kakadu national parks, two key savanna reserves’, biodiversity conservation 18(1), 3643–3662. https://doi.org/10.1007/s10531-009-9669-4 pearson, r.g., raxworthy, c.j., nakamura, m. & peterson, a.t., 2007, ‘predicting species distributions from small numbers of occurrence records: a test case using cryptic geckos in madagascar’, journal of biogeography 34(1), 102–117. https://doi.org/10.1111/j.1365-2699.2006.01594.x pienaar, u.d.v., 1978, the reptile fauna of the kruger national park, national parks board of south africa, pretoria. price, b., kutt, a.s. & mcalpine, c.a., 2010, ‘the importance of fine-scale savanna heterogeneity for reptiles and small mammals’, biological conservation 143(1), 2504–2513. https://doi.org/10.1016/j.biocon.2010.06.017 pyke, g.h. & ehrlich, p.r., 2010, ‘biological collections and ecological/environmental research: a review, some observations and a look to the future’, biological reviews 85(2), 247–266. https://doi.org/10.1111/j.1469-185x.2009.00098.x qgis development team, 2018, qgis geographic information system version 3.4. open source geospatial foundation project, viewed 28 february 2018, from https://www.qgis.org/en/site/index.html. r core team, 2018, r: a language and environment for statistical computing, r foundation for statistical computing, viewed 05 march 2018, from https://www.r-project.org. reddy, s. & davalos, l., 2003, ‘geographic sampling bias and its implications for conservation priorities in africa’, journal of biogeography 30(11), 1719–1727. https://doi.org/10.1046/j.1365-2699.2003.00946.x redfern, j.v., viljoen, p.c., kruger, j.m. & getz, w.m., 2002, ‘biases in estimating population size from an aerial census: a case study in the kruger national park, south africa: starfield festschrift’, south african journal of science 98(9), 455–461. sanparks, 2016, kruger national park official guide, jacana media, melville. stockwell, d. & peters, d., 1999, ‘the garp modelling system: problems and solutions to automated spatial prediction’, international journal of geographical information science 13(2), 143–158. https://doi.org/10.1080/136588199241391 syfert, m.m., smith, m.j. & coomes, d.a., 2013, ‘the effects of sampling bias and model complexity on the predictive performance of maxent species distribution models’, plos one 8(2), e55158. https://doi.org/10.1371/journal.pone.0055158 tolley, k.a., alexander, g.j., branch, w.r., bowles, p. & maritz, b., 2016, ‘conservation status and threats for african reptiles’, biological conservation 204(part a), 63–71. https://doi.org/10.1016/j.biocon.2016.04.006 trimble, m.j. & aarde, r.j., 2014, ‘amphibian and reptile communities and functional groups over a land use gradient in a coastal tropical forest landscape of high richness and endemicity’, animal conservation 17(5), 441–53. https://doi.org/10.1111/acv.12111 venter, f.j., naiman, r.j., biggs, h.c. & pienaar, d.j., 2008, ‘the evolution of conservation management philosophy: science, environmental change and social adjustments in kruger national park’, ecosystems 11(2), 173–192. https://doi.org/10.1007/s10021-007-9116-x zielinski, p., 2001, ‘dealing with uneven recording effort in regional atlas projects on the distribution of amphibians and reptiles’, acta zoologica cracoviensia 44(2), 85–92. introduction the wildlife industry in south africa climate change and range shifts potential options to mitigating the impact of climate change conclusion acknowledgements references about the author(s) haemish i. melville department of environmental sciences, college of agriculture and environmental sciences, university of south africa, florida, south africa robyn s. hetem school of animal, plant and environmental sciences, faculty of science, university of the witwatersrand, johannesburg, south africa w. maartin strauss department of environmental sciences, college of agriculture and environmental sciences, university of south africa, florida, south africa citation melville, h.i., hetem, r.s. & strauss, w.m., 2021, ‘is climate change a concern for the ownership of game within fenced wildlife areas?’, koedoe 63(1), a1673. https://doi.org/10.4102/koedoe.v63i1.1673 scientific letter is climate change a concern for the ownership of game within fenced wildlife areas? haemish i. melville, robyn s. hetem, w. maartin strauss received: 09 feb. 2021; accepted: 08 june 2021; published: 27 july 2021 copyright: © 2021. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. introduction the game theft act 105 of 1991 (hereafter the game theft act) stipulates that ownership of wild animals on a property is vested in the landowner, provided that the landowner has taken steps to prevent the wild animals from escaping. furthermore, should such wild animals escape, the landowner retains ownership and has the right to recapture these animals, even if the animals escaped onto land owned by someone else (blackmore 2020). in a recent article entitled ‘climate change and the ownership of game: a concern for fenced wildlife areas’, published in koedoe, blackmore (2020) suggested that south african legislation relating to game ownership (specifically the clauses within the game theft act that relate to the retention of ownership rights in relation to escaped animals) may require revision under future climate change scenarios. blackmore (2020) argued that in response to changing climates, wild animals will increasingly escape from fenced wildlife areas (e.g. reserves and game ranches) to seek more favourable habitats, which may result in loss of ownership and reduced economic integrity of the south african wildlife industry. we argue that wildlife enterprises with a commercial focus, which potentially face the greatest risk of reduced economic integrity should their wild animals escape, potentially have access to mitigation options such as the provisioning of food and water resources to retain high value species. conversely, wildlife enterprises with a conservation focus may benefit from increased connectivity of wildlife areas, which may result in species diversification that would attract more game viewers. whilst we agree that the game theft act may need revision, we argue that the specific local circumstances, that we outline below, should be taken into account and that additional research and monitoring would be required to inform any proposed revisions. for the purposes of this narrative, we will confine our frame of reference to larger species of wild animals that may best be described as charismatic megafauna. these include large (≥ 10 kg) mammals, both carnivores and herbivores, that are typically the focus of south african commercial wildlife enterprises. wildlife enterprises refer to all wildlife related businesses that involve the containment of wild animals and include national parks as well as small private game reserves. in south africa, ecosystem processes are often rigorously managed on both commercial and conservation based wildlife enterprises, for example, animal populations may be subject to control and/or vegetation may be manipulated through controlled burns. managing animal numbers and the availability of resources (natural or provisioned water and forage) may buffer the adverse effects of increased temperatures and alleviate the pressure for animals to track their moving bioclimatic envelope with changing climates. however, animals might escape from a wildlife enterprise for a variety of reasons, not necessarily linked to climate change. should new legislation be considered in relation to wildlife ownership in the face of climate change, we would need a better understanding of the drivers underlying animal escapes from wildlife enterprises. we suggest that even if the lower level of evidence, fraction of attributable risk (pfrommer et al. 2019), were the qualification for the contention that animals were lost or escaped because of climate change, it would be difficult, if not impossible, to prove. the wildlife industry in south africa wildlife management is the management of wild animals and their habitats with specific economic, ecological and conservation goals in mind. the wildlife industry in south africa is unique insofar as it is one of the few countries where wildlife ownership is vested in a landowner, provided the landowner has taken the necessary precautions to establish ownership rights by enclosing the periphery of their property (blackmore 2020). most wildlife enterprises, whether national, provincial or private, are enclosed by some form of game fence or barrier (natural or manmade) (hayward & somers 2012). therefore, in the south african context, all wildlife enterprises may be viewed as islands of habitat within a broader land use matrix. the wildlife industry in south africa comprises a variety of animal management approaches (modified from taylor, lindsey & davies-mostert 2016), which are not necessarily mutually exclusive. these are summarised as: ecotourism enterprises intensive wildlife production enterprises that include both captive breeding of high value and common species and the intensive breeding of colour morphs of various game species trophy hunting enterprises focus on the production of high value individual animals of a variety of species which are offered as trophies (outstanding exemplar individuals) for organised hunting expeditions biltong (dried cured meat) hunting and venison production enterprises typically produce regional species for the local recreational hunting fraternity and a growing venison market. generally, these management approaches can be divided into those with a conservation focus (e.g. ecotourism) or those with a commercial focus (e.g. intensive wildlife production and hunting) (taylor et al. 2016). however, it is not unusual for a wildlife enterprise with a commercial focus to incorporate more than one of the above management approaches to generate income. the profitability of intensively managed production systems hinges on the ability of landowners to exert ownership rights over wild animals (koelble 2011). consequently, the game theft act is intended to ‘protect the interests of the commercial industry’ (blackmore 2020). the fundamentally different objectives of the various wildlife enterprises may require unique approaches in terms of both management and legislation. a recent amendment to the animal improvement act includes 24 indigenous mammals as landrace breeds (somers et al. 2020). arguably, some commercial wildlife enterprises should be reclassified as agricultural because conserving biodiversity is not the priority. the motivation of conserving profitability rather than biodiversity is evident. few wildlife enterprises manage their land with the objective of conserving habitat for common species (wade 2020). we emphasise that, from an ecological perspective, all species are valuable and contribute functionally to the ecosystems in which they occur. therefore, incentivising biodiversity conservation on commercial wildlife enterprises may be worth formalising. even if few wildlife enterprises prioritise conservation, one cannot overlook the potential conservation value of large tracts of land dedicated to wildlife production. for example, there may be substantial benefits to species, and natural guilds or assemblages resulting from land under wildlife production. however, management practices purely in pursuit of commercial gain may result in long-term, potentially permanent, alteration of ecosystem function (holmes et al. 2020). despite a large portion of the south african landscape (16.8%) being under wildlife production (taylor et al. 2016), much of the land is not in a pristine ecological state and has recently been converted from stock farming. furthermore, it typically comprises relatively small (median size: 2100 ha), fenced parcels (taylor et al. 2016) situated in a diverse matrix of land uses, including mining, subsistence and intensive stock and crop production, forestry, rural habitation and urban development (spierenberg & brooks 2014). hence, the expansion of existing wildlife enterprises and the development of wildlife corridors to facilitate dispersal may not be practicable on a large scale and there may not be much opportunity for individual animals to move. climate change and range shifts climate change undoubtedly influences many of the species to which blackmore (2020) referred. in open systems, mobile animals shift their distribution both polewards latitudinally and upwards altitudinally to track suitable bioclimatic envelopes (parmesan & yohe 2003). however, the geographic pattern of these range shifts may vary and are unlikely to be consistent between species that differ in their thermal tolerance and capacity to persist, which may in turn modify ecosystem structure and function (walther 2010). indeed, these indirect effects of climate change may be more pervasive than the direct effects thereof. in short, we need more information on acclimatisation capacity on a variety of species before we can accurately predict how ecosystems may change. the charismatic megafauna on which the wildlife industry depends may be particularly vulnerable to future climate change because of their long generation times and limited capacity to adapt genetically (hetem et al. 2014). hence, the future profitability of the game ranching sector is under threat, but it is unclear how a change in the game theft act could ameliorate such challenges. within the southern african context, the projected increases in temperature may be exacerbated by a reduction in water availability as a result of increased frequency and severity of droughts, reduced precipitation, increased evapotranspiration and increased water usage as the human population grows (engelbrecht et al. 2015). many animals depend on water to dissipate excess heat loads. reliance on water will likely increase as ambient temperatures increase, consequently the management and availability of free water will influence where, and in what densities, large mammals are able to persist (fuller et al. 2021). with costs of water provisioning likely to increase, water provisioning may not be a sustainable mitigation option and the threshold at which costs outweigh benefits are likely to differ depending on the management model of the wildlife enterprise in question. for conservation enterprises, water management should aim to maintain landscape heterogeneity without disadvantaging water-independent species (gaylard, owen-smith, & redfern 2003) or reducing resilience of water-dependent species (selebatso et al. 2018). wildlife enterprises that focus on the production of high value game species may be prepared to invest heavily in resource provisioning negating the need for animals to shift their ranges (mwakiwa et al. 2013). potential options to mitigating the impact of climate change where expansion of wildlife enterprises is feasible, the likely buy-in from landowners may depend on whether the landowners maintain a conservation or commercial focus. whilst the removal of fences to create corridors or expand wildlife areas may reduce ‘economic integrity’ of the wildlife production industry that relies on ownership of wildlife for profit (blackmore 2020), larger wildlife areas that expand suitable habitat for the ‘big five’ and other wide-ranging species (e.g. african wild dogs, lycaon pictus) may enhance tourism attractiveness and improve profitability of ecotourism. indeed, experienced wildlife viewers and local tourists show an increased interest in returning to protected areas that offer opportunities to view rarer species, such as wild dogs (lindsey et al. 2007). whilst we support the suggestion that wildlife enterprises expand their properties to try to accommodate the likely movement of animals because of climate change, we appreciate that it may not always be practical given the amount of additional land likely required to try and ameliorate shifts in the current spatial distribution of megafauna. a more feasible approach, which has already been used to great effect, would be the amalgamation of contiguous wildlife enterprises to allow larger patches of continuous habitat for wild animals. the development of templates for the successful negotiation of ownership rights to overcome challenges associated with varying economic models of component wildlife production strategies will have a direct bearing on the feasibility of this approach. an example of an approach that might inform the development relationships between alternative wildlife management strategies, to the benefit of conservation, is that between the associated private nature reserves (apnr) and the kruger national park. contrary to the assertion by blackmore (2020), the use of corridors need not automatically result in wild animals reverting to their res nulius status. a preventative measure would be for the corridors themselves to be fenced, which would maintain the ‘sufficiently enclosed’ status of wildlife as is required by the game theft act. whilst the successful establishment and maintenance of corridors between wildlife areas may be challenging, where practical, corridors in combination with the amalgamation of individual wildlife enterprises may still be the most feasible way to allow for the movement of wild animals in search of resources or more suitable climates. conclusion whilst both the economic and ecological underpinnings of wildlife management have advanced substantially within the past four decades, legislation that pertains to the ownership and management of wild animals has largely remained unchanged. as our social, economic and ecological contexts change, it will be necessary for legislation to remain relevant. we support the notion that legislation in relation to the management of wildlife in south africa, such as the game theft act, should be subject to review. however, this should not simply be justified in the light of a perceived response to climate change but should take the local circumstances and practicalities into consideration. acknowledgements competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions haemish melville was responsible for the initiation and conceptualisation of the manuscript and compiling the first draft document. dr hetem and dr strauss, being experts in the field of mammalian response to climate change, advised and added to the narrative in relation to the specifics that relate to climate change. they also contributed substantively to editing the initial document for submission and contributed by assisting in addressing the reviewers’ comments. ethical considerations this article followed all ethical standards for research without direct contact with human or animal subjects. funding information this research received no specific grant from any funding agency in the public, commercial or not-for-profit sectors. data availability data sharing is not applicable to this article as no new data were used in this scientific letter. disclaimer the views expressed in this scientific letter are those of the contributing authors and are not necessarily a reflection of any official policy or the stance of the agencies or institutions to which the authors are affiliated. references blackmore, a.c., 2020, ‘climate change and the ownership of game: a concern for fenced wildlife areas’, koedoe 62(1), a1594. https://doi.org/10.4102/koedoe.v62i1.1594 engelbrecht, f., adegoke, j., bopape, m.-j., naidoo, m., garland, r., thatcher, m. et al., 2015, ‘projections of rapidly rising surface temperatures over africa under low mitigation’, environmental research letters 10(8), 085004. https://doi.org/10.1088/1748-9326/10/8/085004 fuller, a., mitchell, d., maloney, s.k., hetem, r.s., fonseca, v.f.c., meyer, l.c.r. et al., 2021, ‘how dryland mammals will respond to climate change: the effects of body size, heat load and a lack of food and water’, journal of experimental biology 224(1), jeb238113. https://doi.org/10.1242/jeb.238113 gaylard, a., owen-smith, n. & redfern, j.v., 2003, ‘surface water availability: implications for heterogeneity and ecosystem processes’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeniety, pp. 171–188, island press, washington, dc. hayward, m.w. & somers, m.j., 2012, ‘an introduction to fencing for conservation’, in m.j. somers & m.w. hayward (eds.), fencing for conservation, pp. 1–6, springer, new york, ny. hetem, r.s., fuller, a., maloney, s.k. & mitchell, d., 2014, ‘responses of large mammals to climate change’, temperature 1(2), 115–127. https://doi.org/10.4161/temp.29651 holmes, p.m., esler, k.j., gaertner, m., geerts, s., hall, s.a., nsikani, m.m. et al., 2020, ‘chapter 23: biological invasions and ecological restoration in south africa’, in: b. van wilgen, j. measey, d. richardson, j. wilson & t. zengeya (eds.), biological invasions in south africa. invading nature – springer series in invasion ecology, vol 14, pp. 665–700, springer, cham. koelble, t.a., 2011, ‘ecology, economy and empowerment: ecotourism and the game lodge industry in south africa’, business and politics 13(1), 1–24. https://doi.org/10.2202/1469-3569.1333 lindsey, p.a., alexander, r., mills, m.g.l., romañach, s. & woodroffe, r., 2007, ‘wildlife viewing preferences of visitors to protected areas in south africa: implications for the role of ecotourism in conservation’, journal of ecotourism 6(1), 19–33. https://doi.org/10.2167/joe133.0 mwakiwa, e., de boer, w.f., hearne, j.w., slotow, r., van langeveld, f., peel, m. et al., 2013, ‘optimization of wildlife management in a large game reserve through waterpoints manipulation: a bio-economic analysis’, journal of environmental management 114(2013), 352–361. https://doi.org/10.1016/j.jenvman.2012.10.029 parmesan, c. & yohe, g., 2003, ‘a globally coherent fingerprint of climate change impacts across natural systems’, nature 421(6918), 37–42. https://doi.org/10.1038/nature01286 pfrommer, t., goeschl, t., proelss, a., carrier, m., lenhard, j., martin, h. et al., 2019, ‘establishing causation in climate litigation: admissibility and reliability’, climatic change 152, 67–84. https://doi.org/10.1007/s10584-018-2362-4 selebatso, m., bennitt, e., maude, g. & fynn, r.w., 2018, ‘water provision alters wildebeest adaptive habitat selection and resilience in the central kalahari’, african journal of ecology 56(2), 225–234. https://doi.org/10.1111/aje.12439 somers, m.j., walters, m., measey, j., strauss, w.m., turner, a.a., venter, j.a. et al., 2020, ‘the implications of the reclassification of south african wildlife species as farm animals’, south african journal of science 116(1–2), 1–2. https://doi.org/10.17159/sajs.2020/7724 spierenberg, m. & brooks, s., 2014, ‘private game farming and its social consequences in post-apartheid south africa: contestations over wildlife, property, and agrarian futures’, journal of contemporary african studies 32(2), 151–172. https://doi.org/10.1080/09637494.2014.937164 taylor, w.a., lindsey, p.a. & davies-mostert, h., 2016, an assessment of the economic, social and commercial value of the wildlife ranching industry and its potential to support the green economy in south africa, endangered wildlife trust, johannesburg. wade, j., 2020, ‘application of harm reduction thinking to the conservation of uncharismatic species’, in c. stephen (ed.), animals, health and society, pp. 271–278, crc press, london. walther, g.r., 2010, ‘community and ecosystem responses to recent climate change’, philosophical transactions of the royal society b: biological sciences 365(1549), 2019–2024. https://doi.org/10.1098/rstb.2010.0021 http://www.koedoe.co.za open access page 1 of 1 reviewer acknowledgement acknowledgement to reviewers in an effort to facilitate the selection of appropriate peer reviewers for koedoe, we ask that you take a moment to update your electronic portfolio on https://koedoe.co.za for our files, allowing us better access to your areas of interest and expertise, in order to match reviewers with submitted manuscripts. if you would like to become a reviewer, please visit the journal website and register as a reviewer. to access your details on the website, you will need to follow these steps: 1. log into the online journal at https://koedoe. co.za 2. in your ‘user home’ [https://koedoe.co.za/ index.php/koedoe/user] select ‘edit my profile’ under the heading ‘my account’ and insert all relevant details, bio statement and reviewing interest(s). 3. it is good practice as a reviewer to update your personal details regularly to ensure contact with you throughout your professional term as reviewer to koedoe. please do not hesitate to contact us if you require assistance in performing this task. publisher: publishing@aosis.co.za tel: +27 21 975 2602 tel: 086 1000 381 the editorial team of koedoe recognises the value and importance of the peer reviewer in the overall publication process – not only in shaping the individual manuscript, but also in shaping the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely, productive peer review process. we would like to take this opportunity to thank all reviewers who participated in shaping this volume of koedoe. we appreciate the time taken to perform your review(s) successfully. alicia t.r. acosta aniruddha belsare antoinette swart corli wigley-coetsee danny govender dave a. balfour eduardo m. arraut eric mashimbye hugo bezuidenhout jaco barendse jesse m. kalwij jessica cockburn joana bezerra louise swemmer mariette van der walt mariette marais michael d. panagos ndukuyakhe ndlovu nkabeng mzileni nonhlanhla vilakazi peter a. novellie peter chatanga sam m. ferreira sellina e. nkosi sue taylor thabiso mokotjomela theuns c. de klerk wayne twine werner nell yolandi ernst yusuf a. ahmed http://www.koedoe.co.za� https://koedoe.co.za� https://koedoe.co.za� https://koedoe.co.za� https://koedoe.co.za/index.php/koedoe/user https://koedoe.co.za/index.php/koedoe/user mailto:publishing@aosis.co.za abstract introduction materials and methods results and discussion conclusion acknowledgements references footnotes about the author(s) babatope e. akinyemi agricultural economics and extension department, faculty of science and agriculture, university of fort hare, alice, south africa abbyssinia mushunje agricultural economics and extension department, faculty of science and agriculture, university of fort hare, alice, south africa citation akinyemi, b.e. & mushunje, a., 2020, ‘community-based ecotourism project in communities adjacent to the addo elephant national park: will households pay for it?’, koedoe 62(1), a1571. https://doi.org/10.4102/koedoe.v62i1.1571 original research community-based ecotourism project in communities adjacent to the addo elephant national park: will households pay for it? babatope e. akinyemi, abbyssinia mushunje received: 27 mar. 2019; accepted: 13 mar. 2020; published: 22 july 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract management of protected areas (pas) that includes previously excluded households is gaining approval from both households living in communities adjacent to pas and pa managers globally. this article examined households willingness to pay (wtp) for community-based ecotourism (cbe) project in adjacent communities to the addo elephant national park (aenp). the study adopted two-stage sampling procedure to interview 198 respondents using double-bounded contingent valuation (cv) questionnaire administered through face-to-face interviews in five communities adjacent to the aenp. quantitative data analysis was performed with descriptive statistics, pairwise correlation and seemingly unrelated bivariate probit. findings reveal a strong association between respondent’s support for cbe implementation and wtp. the null hypothesis that wtp is independent of the first bid price is strongly rejected. it was observed that firstand second-bid prices, and seven other variables determined wtp. lastly, households are willing to pay between r121.00 and r125.00 monthly for a period of 3 years to support development of cbe initiative in their communities. conservation implications: this study established that households in communities adjacent to the aenp are in support of the establishment of cbe projects as long as they will be beneficial to them and their communities. this support is demonstrated by households’ willingness to pay for the implementation of community-based projects. hence, we concluded that the implementation of a cbe project can offer a market for craft-making, village accommodation and village tours that can showcase adjacent communities to visitors and thus assist the park management in strengthening the existing relationship between the park and communities. keywords: addo elephant; bivariate probit; contingent valuation; community-based ecotourism; protected areas. introduction south africa’s approach to the conservation of nature was a colonialist method, which was built on the idea that households living in communities adjacent to the protected areas (pas) should be excluded to ensure optimum protection of plants, animals and the environment (paterson 2007). however, this exclusionary approach to conservation is gradually giving way to a more inclusive approach to conservation, whereby communities surrounding pas actively participate in both conservation and the sustainable use of the resources in pas, which is a hallmark of the modern conservation model (algotsson 2006; balint 2007; fabricius, koch & magome 2001; pelser, redelinghuys & velelo 2013). the inclusion of local communities in conservation is important when viewed from the perspective of the effective environmental governance (eg), whereas, their exclusion often makes enforcement of conservation policies extremely difficult (aswani & weiant 2004). to formally integrate households with conservation efforts in south africa, the south african national parks (sanparks), which is the largest conservation and ecotourism provider within the country, established a social ecology unit (sanparks 2011). this unit served as the precursor of the households and conservation directorate in 2003 aimed at bringing about a households–parks interface by inculcating a culture of stewardship of the environment and raising awareness about conservation issues amongst local households (sanparks 2011). the legal framework of the households and parks support the preservation of the integrity of the ecosystems, whilst simultaneously acknowledging the vital role played by humans in the conservation of nature. moreover, a draft policy on buffer zones for national parks was published in 20101 to encourage sustainable development compatible with conservation values of the surrounding national park, and to help adjacent and affected communities obtain suitable and sustainable benefits. to fully harness these benefits, namely, promotion of conservation economy, ecotourism and its supporting structure, services and sustainability through a planned harvesting of biological resources have been suggested.2 ecotourism concept ecotourism is a component of sustainable tourism that has been receiving growing attention for over three decades because of the environmental benefits that are intertwined with the well-being of local households. although there is no universally accepted definition of ecotourism, the first definition given by ceballos-lascurain in 1987 defines ecotourism as ‘travelling to relatively undisturbed or uncontaminated natural areas with the specific objective of studying, admiring and enjoying the scenery and its wild plants and animals’. the international ecotourism society (ties) (1990) further defined ecotourism as a ‘… responsible travel to natural areas which conserves the environment and sustains the well-being of local households’. ecotourism has become one of the world’s major economic sectors with the capability to play a significant role in sustainable development in areas where nature attracts tourists (fennel 2002; powell & ham 2008). the business model of sanparks is anchored on three core pillars, namely: conservation, nature-based tourism and community, which are perfectly aligned with the core values of ecotourism. in line with these core values, buckley (2001) recommended that ecotourism product owners like sanparks should have a ‘best practice model’ that can be used as a benchmark to evaluate environmental performance and help in planning new ecotourism products responsibly. it is very important that sanparks implement an ecotourism model to successfully develop and manage its ecotourism products. rest assured that sanparks management operations are well aligned with the core values of ecotourism principles, it has been established that no concrete model of ecotourism is currently in place to clearly show how ecotourism should be developed and managed in south africa (de witt, van der merwe & saayman 2011). willingness to pay concept the concept of willingness to pay (wtp) or reservation price can be defined as the maximum price a given consumer accepts to pay for a product or service (marine 2009). this concept is often used to identify the number of individuals that are willing to pay a given price for a particular product or service. ability to measure consumer wtp makes it possible to calculate the demand curve according to price and to determine the price that offers the optimum feasible margin. if prices can be tailored, knowledge of wtp could enable optimisation of both sales volumes and margins. knowing the factors that drive wtp allows it to be raised and offers the opportunity of increasing sales volumes for a given price or, when possible, to customise prices. some studies have adopted the concept of wtp to examine households’ readiness to contribute or pay for ecotourism in communities adjacent to pas in developing countries. for instance, ezebilo, mattsson and afolami (2010) estimated communities’ willingness to contribute for an ecotourism improvement project and its determinants in the okwangwo division of the cross river national park, nigeria. the study revealed that households were willing to contribute an average of about 1% of their mean annual income and that willingness to contribute was influenced by income, distance of respondents’ residence to the park, post-high school education, occupation and membership of an environmental conservation group. similarly, kimengsi et al. (2019) examined the extent to which respondents’ assets drive ecotourism choices in cameroon. the study revealed a high preference for the production and sale of arts and craft items and the promotion of cultural heritage sites as key ecotourism choices. moreover, women were found to participate more in conservation education, as opposed to culture-related activities such as arts and crafts. also, access to education and training was observed to be inversely related to the promotion of cultural festivals in the study (kimengsi et al. 2019). hence, this study elicited households’ wtp for community-based ecotourism (cbe) initiative in the five villages bordering the addo elephant national park (aenp). the study adopted the double-bounded contingent valuation (dbcv) method to elicit information on households’ wtp for cbe in their villages. the ‘materials and methods’ and the ‘results and discussion’ sections present detailed descriptions of the methodology used, and the results and discussion of the study, respectively. materials and methods description of the study area this study was conducted in five communities bordering the aenp in the eastern cape province of south africa. the park straddles the following local and district authority boundaries: sundays river local municipality; ndlambe local municipality; ikwezi local municipality; blue crane local municipality; nelson mandela metropolitan; and cacadu district municipality. eight adjacent communities that share proximities to the park include nomathamsanqa, valencia, paterson, enon, bersheba, joe slovo, moses mabida and colchester. historical records showed that khoekhoen (khoi khoi) households arrived and settled in the area now designated as aenp between 2000 and 1500 years ago, and many of them died during an outbreak of smallpox that hit the area in the early 1700s (south africa nature reserves & national parks 2016). almost around the same period, the nomadic xhosa tribes also arrived in the region, and many of them settled down in the northern-most part of wit river. several clashes ensued between the already settled khoi khoi settlers and the newly arrived nomadic xhosa (south africa nature reserves & national parks 2016). over a century ago, the area was also a focal point of bitter conflicts between pastoralists of european and african origin. during the anglo-boer war, a boer commando, led by general jan smuts, entered the zuurberg mountains in september 1901 (reitz 1929). the following day, the commando being severely pressed for food, cooked and ate the fruit of a cycad encephalartos longifolius. about 80 men were poisoned, and 20 were very ill, some becoming unconscious, but fortunately all survived. the sick men made a dramatic escape from the advancing british forces (ferreira 1980; reitz 1929). whilst the war lasted, hunters poached elephants and other game animals in the area unabatedly and eventually by the early 1900s, there were only a few small populations of elephants left (ferreira 1980). a ruling passed in 1919 by major pj pretorious ordering that all of the remaining elephants in the area were to be exterminated saw 114 of these animals shot in just over a year. by 1931, only 11 elephants remained, and a national park was proclaimed in an effort to protect these wild animals (south africa nature reserves & national parks 2016). since the proclamation of the national park, efforts have been made to minimise poaching and other human–wildlife conflicts in and around the pas. one such effort is the mayibuye ndlovu (meaning ‘let the elephant return’ in xhosa) community forum, which was established in 1993 to resolve conflicts between the park and the adjacent nomathamsaqa community (kate 2006). in recent times, due recognition has been accorded in the form of concessions and ecotourism development opportunities in and around the park, along with associated opportunities to supply goods and services to these businesses (kate 2006). this has resulted in the expansion of the community forum to become more representative of all the local communities and the formation of the mayibuye ndlovu development trust (mndt). this trust is represented by the park, the local government, the sunday river valley tourism forum and the eight surrounding local communities (sanparks 2005). conceptual framework and empirical strategy the empirical analysis of a respondent’s wtp for cbe development in this study is based on the bivariate probit model. the bivariate normal density function is appealing to statisticians because it allows non-zero correlation, whilst the logistic distribution does not. in addition, constraining the parameters of the bivariate probit model yields other models such as the interval data model and the random effect probit model (cameron & quiggin 1994). econometrically, modelling the data generated by the dbcv relies on the formulation given by: where wtpij represents the jth respondent’s wtp and i = 1, 2, denoting the first and the second questions as the means for the first and second responses. as before, setting u1 = x’ijb allows the means to be dependent upon the characteristics of the respondent. following haab and mcconnell (2002), the jth contribution to the likelihood function is given as: where yy = 1 for a ‘yes-yes’ answer, 0 otherwise, ny = 1 for a ‘no-yes’ answer, 0 otherwise and so on. this is the bivariate discrete choice model. assuming normally distributed error terms with mean zero and respective variances σ1 and σ2, then wtp1j and wtp2j have a bivariate normal distribution with means µ1 and µ2, variances σ1 and σ2 and correlation coefficient ρ. as a result, the jth contribution to the bivariate probit in equation 3 is the likelihood function for the bivariate probit: where φε1ε2 is the standardised bivariate normal distribution function with zero means; y1j =1 if the response to the first question is yes and 0 otherwise y2j = 1 if the response to the second question is yes and 0 otherwise d1j = 2y1j −1 and d2j = 2y2j −1 ρ = correlation coefficient σ = standard deviation. although the double-bounded question format may be potentially biased, its use may be justified because it leads to lower mean squares error (alberini 1995). it may also yield more conservative wtp estimates (banzhaf et al. 2004) by narrowing down the confidence interval around wtp measures. the mean and median wtp as given by huang and smith (1998) was estimated using equation 4: where is the coefficient on the bid amount. as shown in cameron and james (1987), the coefficient on the bid amount is a point estimate of . the dispersion parameter or standard deviation of wtp is thus estimated at: presented in table 1 are the variables included in the bivariate probit model. it shows the variables, types of measures and their descriptions. two categories of variables dealing with respondents’ socio-economic characteristics and their knowledge of deforestation in the aenp, tourism and ecotourism are also presented. table 1: a priori expectations of the determinants of willingness to pay for community-based ecotourism. description of data the contingent valuation survey was implemented amongst respondents selected from five villages (enon, bersheba, moses mabida, nomathamsanqa and valencia) adjacent to the aenp. the park ranks as the third-largest national park after the kruger national park and the kgalagadi transformation park in south africa. it is unique for being the most biologically diverse park in africa (aenp 2015). in its present state, it represents five of south africa’s seven biomes, namely, the nama karoo, fynbos, forest, thicket, grassland and the azonal wetland (only lacking are the succulent karoo and savannah) (vlok, euston-brown & cowling 2003). the park is also the only park in south africa with the big seven because of its coastal area. the study employed a two-stage sampling technique to identify 198 households as presented in table 2. in the first stage, the names of eight villages adjacent to the aenp were retrieved from sanparks with which authors signed a contractual agreement. from these eight villages, five villages adjacent to the pa were selected using simple random sampling, which is a probability sampling method. the second stage involved non-probability convenience sampling of households that are willing to participate in the study from the selected villages. the choice of the convenience sampling method was informed by non-availability of sampling frame from which random sample can be drawn for the study. five enumerators from the five selected communities were recruited through the assistance of mndt, and the enumerators were trained by the researcher on how to administer the questionnaire to the respondents in their local language to ensure a clear understanding of the study. the data were collected in march 2016. table 2: sampling procedure for households. the respondents were assured of their confidentiality, and their privacy was respected. enumerators informed the respondents of the aim of the study and secured their consents before commencing the interviews. in all, a total of 210 copies of questionnaires were administered to households in communities adjacent to the pas. of all the copies of the questionnaire administered, 198 representing a 94% response rate, contained meaningful information for analysis. this sample size aligned with the findings of heechan (2006), which confirmed that the statistical gain of the bivariate probit model is negligible for sample sizes ranging from 200 to 1000. moreover, similar studies (ezebilo et al. 2010; kimengsi et al. 2019) implemented in nigeria and cameroon used sample sizes ranging from 150 to 200 respondents. this study was ethically approved by the university of fort hare, with the ethic clearance certificate number (rec-270710-028-ra level 01). the researcher assured the study respondents that the findings of the study will be shared with them through mndt, and the research findings were shared with mndt at the completion of the study in september 2016. description of the hypothetical market before posing the dbcv wtp questions to the respondents, description of the contingent good (cbe project) was established. it was explained to respondents that ‘visitors to the aenp may be interested in participating in cbe projects’. these projects include: participating in craft-making market, using village accommodation and engaging in village tours that will take visitors to historic sites within their villages and also expose visitors to their cultural values. this project will provide alternative sources of livelihood, which may reduce exploitation of natural resources, thereby minimising the rate of deforestation. the respondents were informed that the establishment of cbe in their villages would require each household to contribute a monthly payment for the funding of the project for the next 3 years. this is important because evidence reveals that individuals respond differently when responding to hypothetical questions than when having to make real payment (list & gallet 2001). because consumers usually dislike increase in prices (jeanty 2006), payment methods like an increase in federal or provincial taxes and prices were not considered. the elicitation method used in the households’ survey is the double-bounded dichotomous choice (db dc) format. the db dc format questions ask the respondent whether they would vote ‘yes’ on a policy that would cost her household an initial amount in south african rands. if the respondent says ‘yes’ (respectively, no), another wtp question was asked using a higher (respectively, lower) price. a common problem found in this survey design is starting point bias, as recognised by herriges and shrogren (1996). it has to do with the design of the bid vector and occurs when the initial bid introduced in the survey influences the respondent’s level of wtp. according to mitchell and carson (1989), anchoring effect and yes-saying bias are two major sources of starting point bias. to deal with starting point bias in this study, contingent valuation method (cvm) questionnaire was designed with an optimal number and the distribution of bids to offer improved efficiency (alberini 1995). the questionnaire has three sections. the first section deals with the respondent’s knowledge of deforestation and information relating to tourism and ecotourism activities in the aenp. the second section contains the valuation scenario, which attempts to provide as much information as possible about the hypothetical market. guidelines suggested by carson (2000) and the noaa panel (arrow et al. 1993) for valid contingent valuation analysis were followed as much as possible. the sets of bids used in the scenario were: 25, 12, 50; 40, 20, 80; 60, 30, 120 and 100, 45, 200, where the first element of each set represents the first bid. the second element corresponds to the lower bid if the respondent answers ‘no’ to the first bid and the third element corresponds to the higher bid if the response to the first bid is a ‘yes’. all bids are in south african rands. to establish the institutional setting in which the cbe project was to be implemented, a scenario was presented to respondents: [w]hereby the south african national parks (sanparks) was considering a community-based ecotourism project that will provide village accommodation, craft market and village tour that will expose visitors to the local culture of the households. however, consistent with the previous work by loureiro gracia and nayga (2006), the respondents were not explicitly told whether findings from this study would affect these considerations. providing this information to the respondents could have affected their decisions, given the context in which the project is to be implemented. the last part of the questionnaire deals with the socio-economic characteristics of the respondents. ethical consideration ethical approval to conduct the study was obtained from the university of fort hare (urec). (reference number mus181saki01) results and discussion the demographic characteristics of the respondents that participated in the study are presented in table 3. on an average, respondents were willing to give r57.00 and r80.00 for the first and second bids presented to them. the mean age of the participants in the study is 40 years. this mean age is 11 years lower than the 51 years reported for the entire province in 2014 general households survey (statistics south africa 2014). this suggests that these respondents are in their economically active years. majority of the participants are males, represented by 52% of the study population. an average household head in the study area attended school for 9 years, whilst 35% are married. the dependency ratio of the households defined by the ratio of non-working household members to the working household members indicates that 78% have a dependency ratio that is greater than 1. seventy-five per cent of the respondents are of xhosa descent, which is similar to the census 2011 figure. it was reported that xhosa constituted 77% of the population in the study area. table 3: descriptive statistics of independent variables. furthermore, the mean distance of household’s residence to the aenp is 17.57 km. the fact that respondents live a mere 17 km away from the pa suggests that they may be familiar enough with the pa in order to value it. studies have also shown that the proximity to a resource, the more the wtp for it (biadgilign et al. 2015). a monthly income of r2265.15 was reported as the mean monthly income in the study. this mean monthly income is comparatively lower than the average annual income of r6400.00 for the eastern cape province, as reported in statistics south africa, 2012. however, we believe that this figure is commensurate with the level of education reported by the respondents in the study area. regarding the six variables measuring respondents’ knowledge of deforestation, deforestation to the aenp, concerns about deforestation, contribution to environmental cause, awareness of ecotourism and involvement in tourism-related business, 43%, 54%, 82%, 15%, 63% and 5%, respectively, were affirmative to these questions. when asked about the usefulness of the information provided in the questionnaire for the respondents to make decisions, 68% affirmed that the questionnaire was useful. to establish zero wtp amongst the study participants, they were first asked whether or not they would favour cbe if funding was made available from either government or non-governmental organisations (ngos). as indicated in table 4, about 84% of the respondents indicated that they would favour the cbe project if money is made available by the government and/or ngos. amongst the respondents that support the project, about 73% were affirmative in their response to the first bid. of the 15.6% that were against the cbe project, all responded negatively to the first wtp question. correlation test results presented in table 4 reject the null hypothesis that whether respondents would favour cbe development is independent of their responses to the first wtp question at 1% level of significance. it invariably implies that household’s response to the first wtp question is strongly associated with either they are in favour or against the implementation of the cbe project. table 4: association between community-based ecotourism support and the first bid payment. distribution of respondents’ responses to the first bid and the results testing the null hypothesis that the responses are independent of the first -bid amount in rand is presented in table 5. the null hypothesis is strongly rejected at 1% level of significance. this means that respondents take into account the bid amounts when stating their wtp. the first three rows in table 5 describe each bid level. the first row corresponds to the frequency of ‘no’ and ‘yes’ answers to the specified bid amount in rand and to the total number of the respondents who were exposed to that bid amount. the second row represents the percentage of ‘no’ and ‘yes’ responses amongst those who were offered the bid amount. the third row indicates the percentage of respondents who indicated ‘no’ and ‘yes’ for that particular bid across all respondents that were offered the bid. table 5: response distribution for the first bids. for bid r25.00, the first row indicates that 44 respondents were exposed to this bid, amongst which 17 rejected the bid price (i.e. no) and 27 accepted the bid price (i.e. yes), whereas the second row implies that 17 represent 38.64% and 27 represent 61.36%. the third row suggests that 22.22% of the respondents were offered the first bid of r25.00 and that amongst all respondents, 22.67% said ‘no’ and 21.95% said ‘yes’ to the first bid of r25.00. theoretically, the third row indicates that the bids satisfy the monotonicity assumption, which implies that the percentage of the respondents who indicated ‘no’ should monotonically increase as the bid amount increases. as indicated in table 5, both ‘no’ and ‘yes’ responses satisfy monotonicity, as suggested by upward and downward sloping hicksian demand function. the graphical representation of the monotonicity assumption is shown in figure 1. it is anticipated that the likelihood of respondents saying ‘yes’ to the lowest bid should be close to 1, and the likelihood of saying ‘yes’ to the highest bid should be close to 0. in this study, 61% of the respondents indicated ‘yes’ to r25.00 the lowest first bid, and 48% indicated ‘no’ to the highest first bid of r100.00. figure 1: percentage of ‘no’ and ‘yes’ to the first bid amounts. the contingency table, which compares the first and second responses of the bids, is presented in table 6. it shows that amongst those who said ‘no’ to the first bid, 69.3% said ‘no’ and 30.4% said ‘yes’ to the second bid. amongst those who answered affirmatively to the first bids, 19.5% said ‘no’ and 80.4% said ‘yes’ to the second bid. counterintuitively, ‘dominant effect’ phenomenon, whereby respondents who initially said ‘yes’ persisted in saying yes and those who initially said ‘no’ are inclined to persist in saying ‘no’ was not satisfied for both ‘no’ and ‘yes’ responses in this study (cameron & quiggin 1994). the test of the null hypothesis that the first and second responses are independent was implemented, and the results indicated that the null hypothesis should be rejected. this implies that the responses to the first and the second questions are correlated. table 6: test of independence between the first and the second responses. the results of the seemingly unrelated bivariate probit model showing the summary statistics of coefficients, standard errors and levels of significance for first and second responses are presented in table 7. when implementing the model, we assumed that the same exogenous variables appear in both equations because there are no requirements that different variables should appear in the two equations (green 2018). the conditional tetrachoric correlation (r) of the error terms (green 2018) capturing the correlation between the binary variables (responses to first and second bids) of 11.70 indicates that there is a correlation between the binary variables. of all the 17 variables included in the bivariate probit model, the first bid: residents’ distance to the aenp, xhosa, deforestation concerns, ecotourism awareness and instrument usefulness are significant in the first equation. whilst, that of second bid: being married, log of income, deforestation knowledge, deforestation concerns and instrument usefulness are significant in the second equation. table 7: seemingly unrelated bivariate probit regression results. respondents’ wtp for cbe in communities adjacent to the aenp is strongly related to both first and second bid amounts. it also follows a priori negative sign as anticipated (du preez, tessendorf & hosking 2010; jeanty 2006). the negative signs indicate that as the bid amount offered to the household increases, their wtp for cbe decreases. the finding follows the popular axiom of demand theory, which states that the higher the price, the lower the quantity demanded. although being married is not significant in the first equation, it significantly influences respondents’ response to the follow-up wtp question. the positive coefficient of the married variable implies that respondents that are married are most likely to be willing to pay for the project than their counterparts who are single, divorced or widowed. because marriage often comes with responsibility in africa, married respondents may see this project as a way of contributing to society. in addition, the log of monthly household’s income is significant with a positive sign as intuitively anticipated. this obviously means that respondents with a higher income are likely to pay for cbe than respondents with a lower income. this finding corroborated past studies that show that households’ wtp is often contingent on the level of income of the households (bonan, lemay-boucher & tenikue 2013; hadker et al. 1997). the variable distance is another important variable that may greatly influence wtp. it is strongly significant (1%) with a negative sign as expected in the first equation but insignificant with a priori negative coefficient sign in the second equation. this result is particularly true of the study area, as previous studies revealed inter-community conflict relating to the location of developmental programmes in the past (timmermans & naicker 2002). the xhosa variable is significant at 10% with a positive sign coefficient estimate. this suggests that respondents who are of xhosa descent are more likely to answer ‘yes’ to the wtp question than other tribes living in the study area. this finding may not be unconnected to the sense of ownership shared by the xhosa on their rights to the natural resource stemming from the age-long history of the households (timmermans & naicker 2002). also, the traditional cultural circumcision of young male xhosa in the forest may be connected to their wtp for the cbe, which will help in the conservation of forest resources around this pa. similarly, it was observed that being deforestation concerned significantly influenced respondents’ wtp for cbe when presented with both bid amounts, albeit at 5% for the first equation and 10% for the second equation. this finding aligns with jeanty’s (2006) study that individuals concerned about deforestation tend to express positive wtp. however, findings of the study contradict akinyemi and mushunje (2017) which found that respondents are concerned about deforestation are less willing to pay for cbe in eastern cape wild coast, south africa. finally, it was observed that respondents’ awareness of ecotourism at 10% significantly affects respondents’ wtp for the first bid. meanwhile, the coefficient on instrument usefulness variable is positive and strongly significant at 1% for both equations. as a result, respondents that find information in the questionnaire useful tend to be more willing to pay for the project than their counterparts who found it less informative. both findings were intuitively anticipated a priori because it is only right to expect that the more the level of awareness of ecotourism and its benefits households have, the more their wtp. similarly, if respondents clearly understand the hypothetical market described in the questionnaire and its environmental market value, they are probably more likely to be willing to pay than those who do not understand. the distribution of the average wtp estimates, the lower and upper bound parameters from the dbcv, achieved significance level, confidence intervals using krinsky and robb procedures are presented in table 8. the confidence intervals reported in this study were estimated with 50 000 draws as recommended by krinsky and robb (1986) and park, loomis and creel (1991). in computing mean/median wtp, mean income from survey data is used. in the first instance, it was observed that the estimated mean/median wtp increased from r121.78 to r125.84 when the second question was introduced. this is a counterintuitive result of the double bounded modelling. the variation in the binary responses conveys information about the error in each equation because the bid amounts vary across individuals. the results of this study imply that the actual ‘willingness to pay’ of the respondents in communities adjacent to the pas in the study area may be lower than the amount estimated by the dbcv. therefore, further study may be required to empirically clarify this. table 8: mean/median willingness to pay the amount. conclusion the aim of this study was to measure household’s wtp for the development of cbe in communities adjacent to the aenp, south africa. in addition, we measured the association between households’ support for cbe implementation and their wtp the first bid price. lastly, we determined the variables that influenced households’ wtp for cbe development amongst households that participated in the study. findings from the study established a strong association between households’ support for cbe implementation and wtp the first bid price (25, 40, 60, 100) offered to the households in the dbcv questionnaire implemented in the study. the null hypothesis that households’ support for the implementation of the cbe development is independent of the first bid price is strongly rejected. this result established that households that support the development of cbe project will probably pay the first bid price when asked to do so. the estimate of the wtp from the bivariate probit model shows that households are willing to pay between r121 and r125 every month for a period of 3 years to support the establishment of the cbe project in their communities, thus confirming the findings of ezebilo et al. (2010) and kimengsi et al. (2019) that the community will contribute or pay for the ecotourism project if they stand to benefit from the project. in conclusion, this study established that households in communities adjacent to the aenp are in support of the establishment of cbe project as long as it will benefit their communities in general and individuals in particular. this support is demonstrated by households’ willingness to contribute money for the implementation of a community-based project in their communities. based on this finding, the study concluded that the implementation of cbe project that offers craft-making market, village accommodation and village tour can showcase adjacent communities to visitors and thus assist the park management to strengthen the existing relationship between the park and the communities. acknowledgements the authors thank the managements of the south african national parks (sanparks) and the addo elephant national park for the provision of accommodation and survey logistics during the tourists’ survey in 2016 and for assisting with the recruitment of enumerators for the household survey. the authors convey their special thanks to marna herbst for the invaluable contribution to the tourists’ questionnaire guide and for facilitating authors’ contractual agreement with sanparks. competing interests the authors have declared that no competing interest exists. authors’ contributions b.e.a. designed this study under the supervision of a.m. the study survey, data analysis and the writing of the first draft of the manuscript were performed by b.e.a. a.m. improved substantially on the subsequent drafts that culminated in the final draft submitted to koedoe. funding information this study was self-funded. data availability statement data sharing is not applicable to this article as no new data were created or analysed in this study. disclaimer all the views and opinions expressed in this article are that of the authors and do not represent that of the university of fort hare, the south african national parks and the addo elephant national park. the authors are responsible for errors and omissions in the submitted content, if any. references addo elephant national park, 2015, park management plan, south african national parks, pretoria. akinyemi, b.e. & mushunje, a., 2017, ‘willingness to pay for wild coast nature reserves conservation through community-based ecotourism projects’, international journal of applied business and economic research 15(16), 57–73. alberini, a., 1995, ‘optimal designs for discrete choice contingent valuation surveys: single-bound, double-bound, and bivariate models’, journal of environmental economics and management 28(3), 287–306. https://doi.org/10.1006/jeem.1995.1019 algotsson, e., 2006, ‘wildlife conservation through households-centred approaches to natural resource management programmes and the control of wildlife exploitation’, local environment 11(1), 79–93. https://doi.org/10.1080/13549830500396230 arrow, k., solow, r., portney, p., leamer, e., radner, r. & schuman, h., 1993, ‘report of the noaa panel on contingent valuation’, federal register 58(10), 4601–4614. aswani, s. & weiant, p., 2004, ‘scientific evaluation in women’s participatory management: monitoring marine invertebrate refugia in the solomon islands’, human organization 63(3), 301–319. https://doi.org/10.17730/humo.63.3.r7kgd4thktmyf7k1 balint, p.j., 2007, ‘a proposed general model for southern african community-based wildlife management’, human dimensions of wildlife 12(3), 169–179. https://doi.org/10.1080/10871200701322829 banzhaf, s., dallas, b., evans, d. & krupnick, a., 2004, valuation of natural resource improvements in the adronacks, resource for the future, washington, dc. biadgilign, s., reda, a.a. & kedir, h., 2015, determinants of willingness to pay for the retreatment of insecticide treated mosquito nets in rural area of eastern ethiopia’, international journal for equity in health 14, a99. https://doi.org/10.1186/s12939-015-0249-9 bonan, j., lemay-boucher, p. & tenikue, m., 2013, ‘households’s willingness to pay for health microinsurance and its impact on actual take-up results from a field experiment in senegal’, the journal of development studies 50(10), 1445–1462. https://doi.org/10.1080/00220388.2014.940909 buckley, r., 2001, ‘major issues in tourism ecolabelling’, in x. font & r.c. buckley (eds.), tourism ecolabelling: certification and promotion of sustainable management, pp. 19–26, cabi, new york, ny. cameron, t. & james, m.d., 1987, ‘efficient estimation methods for “close-ended” contingent valuation surveys’, the review of economics and statistics 69(2), 269–276. https://doi.org/10.2307/1927234 cameron, t. & quiggin, j., 1994, ‘estimation using contingent valuation data from a “dichotomous choice with follow-up” questionnaire’, journal of environmental economics and management 27(3), 218–234. https://doi.org/10.1006/jeem.1994.1035 carson, r.t., 2000, ‘contingent valuation: a user guide’, environmental science and technology 34(8), 1413–1418. https://doi.org/10.1021/es990728j ceballos-lascurain, h., 1987, tourism, ecotourism and protected areas, international union for conservation of nature (iucn) (world conservation union), gland. de witt, l., van der merwe, p. & saayman, m., 2011, ‘an ecotourism model for south african national parks’, in book of proceedings, vol. ii: international conference on tourism and management studies, university of the algarve, algarve, 26–29 october 2011. du preez, m., tessendorf, s. & hosking, s., 2010, ‘application of the contingent valuation method to estimate the willingness to pay for restoring indigenous vegetation in underberg, kwazulu-natal, south africa’, south african journal of economic and management sciences 13(2), 135–157. https://doi.org/10.4102/sajems.v13i2.42 ezebilo, e.e., mattsson, l. & afolami, c.a., 2010, ‘economic value of ecotourism to local communities in the nigerian rainforest zone’, journal of sustainable development 3(1), 51–60. https://doi.org/10.5539/jsd.v3n1p51 fabricius, c., koch, e. & magome, h., 2001, community wildlife management in southern africa: challenging the assumptions of eden, russell press, nottingham. fennel, d.a., 2002, ecotourism programme planning, cabi, new york, ny. ferreira, o., 1980, memoirs of general ben bouwer, human sciences research council (hsrc), pretoria. greene, w.h., 2018, econometric analysis, 8th edn., pearson, new york, ny. haab, t.c. & mcconnell, k.e., 2002, ‘referendum models and economic values: theoretical, intuitive, and practical bounds on willingness to pay’, land economics 74(1998), 216–229. https://doi.org/10.2307/3147052 hadker, n., sharma, s., david, a. & muraleedharan, t., 1997, ‘willingness to pay for borivli national park: evidence from a contingent valuation’, ecological economics 21(2), 105–122. heechan, k., 2006, ‘essays on methodologies in contingent valuation and the sustainable management of common pool resources’, phd dissertation, the ohio state university, columbus, oh. herriges, j.a. & shogren, j.f., 1996, ‘starting point bias in dichotomous choice valuation with follow-up questioning’, journal of environmental economics and management 30(1), january, 112. https://doi.org/10.1006/jeem.1996.0008 huang, j. & smith, v.k., 1998, ‘monte carlo benchmarks for discrete valuation methods’, land economics 74(2), 186–202. https://doi.org/10.2307/3147050 jeanty, w.p., 2006, ‘two essays on environmental and food security’, phd dissertation, graduate school of ohio state university, s.l. kate, h., 2006, partnerships to manage conservation areas through tourism: some best practice models between government, indigenous communities and the private sector in canada and south africa, government of western australia, department of conservation and land management, perth. kimengsi, j.n., kechia, m.a., azibo, b.r., pretzsch, j. & kwei, j., 2019, ‘households’ assets dynamics and ecotourism choies in the western highlands of cameroon’, sustainability 11(7), 1844. https://doi.org/10.3390/su11071844 krinsky, i. & robb, a.l., 1986, ‘on approximating the statistical properties of elasticities’, the review economics and statistics 68(4), 715–719. https://doi.org/10.2307/1924536 list, j.a. & gallet, c., 2001, ‘what experimental protocol influence disparities between actual and hypothetical stated values?’, environmental and resource economics 20, 241–254. https://doi.org/10.1023/a:1012791822804 loureiro, m., gracia, a. & nayga, r.m., 2006, ‘do consumers value nutritional labels?’, european journal of agricultural economics 33(2), 249–268. https://doi.org/10.1093/erae/jbl005 marine le, g., 2009, ‘definition, measurement and determinants of the consumer’s willingness to pay: a critical synthesis and directions for further research’, recherche et applications en marketing 24(2), 91–113. https://doi.org/10.1177/205157070902400205 mitchell, r. & carson, r., 1989, using surveys to value public goods. the contingent valuation method, resources for the future, washington, dc. park, t., loomis, j. & creel, m., 1991, ‘confidence intervals for evaluating benefits from dichotomous choice contingent valuation studies’, land economics 67(1), 64–73. https://doi.org/10.2307/3146486 paterson, a.r., 2007, ‘wandering about south africa’s new protected areas regime’, south african public laws journal 22, 1–3. pelser, a., redelinghuys, n. & velelo, n., 2013, ‘protected areas as vehicles in population development: lessons from rural south africa’, environment, development and sustainability 15(1), 1205–1226. https://doi.org/10.1007/s10668-013-9434-4 powell, r.b. & ham, s.h., 2008, ‘can ecotourism interpretation really lead to pro-conservation knowledge, attitudes and behaviour? evidence from the galapagos islands’, journal of sustainable tourism 16(4), 467–489. https://doi.org/10.1080/09669580802154223 reitz, d., 1929, commando, faber & faber, london. south africa nature reserves & national parks, 2016, march 08, the bloody history of addo elephant national park, siyabonga africa, viewed 08 march 2019, from https://www.nature-reserve.co.za/history-addo-elephant-natio. south african national parks (sanparks), 2005, ‘addo elephant national park official guide’, the tourism blueprint, south african national parks, pretoria. south african national parks (sanparks), 2011, people and conservation: a brief history, viewed 08 march 2019, from http://www.sanparks.org/people/about/history.php. statistics south africa, 2014, general household survey, statistical release p0318 statistics south africa, pretoria. the international ecotourism society (ties), 2015, what is ecotourism?, the international ecotourism society, washington, dc, viewed 25 january 2019, from http://www.ecotourism.org/what-is-ecotourism. timmermans, h. & naicker, k., 2002, ‘the land’, in r. palmer, h. timmermans & d. fay (eds.), from conflict to negotiation: natural-based development on south africa’s wild coast, chap. 1, pp. 2–14, human science research council, pretoria. vlok, j.h., euston-brown, d. & cowling, r., 2003, ‘acocks’ valley bushveld 50 years on: new perspective on the delimitation, characterisation and origin of the thicket vegetation’, south african journal of botany 69(1), 27–51. https://doi.org/10.1016/s0254-6299(15)30358-6 footnotes 1. draft policy on buffer zones for national parks. 2. draft policy on buffer zones for national parks 48. introduction materials and methods results and discussion acknowledgements references about the author(s) albert chakona national research foundation – south african institute for aquatic biodiversity, grahamstown, south africa department of ichthyology and fisheries science, rhodes university, grahamstown, south africa gavin gouws national research foundation – south african institute for aquatic biodiversity, grahamstown, south africa department of ichthyology and fisheries science, rhodes university, grahamstown, south africa wilbert t. kadye department of ichthyology and fisheries science, rhodes university, grahamstown, south africa pule p. mpopetsi department of ichthyology and fisheries science, rhodes university, grahamstown, south africa paul h. skelton national research foundation – south african institute for aquatic biodiversity, grahamstown, south africa citation chakona, a., gouws, g., kadye, w.t., mpopetsi, p.p. & skelton, p.h., 2020, ‘probing hidden diversity to enhance conservation of the endangered narrow-range endemic eastern cape rocky, sandelia bainsii (castelnau 1861)’, koedoe 62(1), a1627. https://doi.org/10.4102/koedoe.v62i1.1627 short communication probing hidden diversity to enhance conservation of the endangered narrow-range endemic eastern cape rocky, sandelia bainsii (castelnau 1861) albert chakona, gavin gouws, wilbert t. kadye, pule p. mpopetsi, paul h. skelton received: 03 mar. 2020; accepted: 22 july 2020; published: 29 sept. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. introduction accurate delimitation of species boundaries is a fundamental requirement for formulating environmental policies and spatial conservation planning to prevent loss of biodiversity (e.g. nel et al. 2011). however, because morphological differences may remain undetected as a result of their subtlelty and the experience or expertise of the observer, there are several cases where two or more morphologically similar species have been mistakenly classified into a single taxon, thus compromising conservation of rare, cryptic and narrow-range species (bickford et al. 2007). the use of molecular data has resulted in the discovery of new species and several historically isolated lineages within many groups of freshwater fishes that were previously considered to be single wide-ranging species. this is particularly true for the cape fold freshwater ecoregion (cfe) in south africa and the eastern zimbabwe highlands freshwater ecoregion (ezh), where new species, unique lineages and taxonomic conflicts have been discovered in various species groups (e.g. bronaugh, swartz & sidlauskas 2020; chakona et al. 2018a; chakona, swartz & gouws 2013; swartz, skelton & bloomer 2009; wishart et al. 2006). such findings have stimulated renewed interest in the systematics and taxonomic revisions of freshwater fishes in southern africa (e.g. chakona & skelton 2017; chakona & swartz 2013; chakona, swartz & skelton 2014; maake, gon & swartz 2014). this information has also been critical in guiding accurate international union for conservation of nature (iucn) redlist assessments for freshwater fishes in south africa (chakona et al. in prep). in the present study, mitochondrial 16s ribosomal ribonucleic acid (rrna) sequences were used to explore patterns of spatial genetic structuring in the eastern cape rocky, sandelia bainsii, with the aim of illuminating the implications of incomplete systematic knowledge on the conservation of aquatic biodiversity. the eastern cape rocky attains the largest size of all anabantid species, reaching a standard length of about 260 mm (skelton 2001). this species is endemic to south africa, where it has a restricted distribution, occurring in short sections of the kowie, great fish, keiskamma, igoda, buffalo and nahoon river systems in the eastern cape province (skelton 2001; figure 1). the species is listed by the iucn as endangered, and the persistence of remnant populations is uncertain as a result of deterioration in water and habitat quality, the spread of non-native piscivores, habitat fragmentation and hydrological modifications (chakona, sifundza & kadye 2018b; figure 1). many of the known remnant populations of this species are highly fragmented, and there are concerns that some of these populations may not be viable in the long term (chakona et al. 2018a). figure 1: historical distribution of the eastern cape rocky, sandelia bainsii, based on records from the national fish collection at the nrf-saiab (black circles), the localities where tissues samples were collected from the kowie (red triangle), great fish (green triangles), keiskamma (orange triangles), igoda (turquoise triangle) and buffalo (navy blue triangles) river systems. the yellow points represent localities where sandelia bainsii was not recorded from recent surveys (2009–2014). a previous genetic study based on mitochondrial cytochrome b data provided the first evidence for the existence of historically isolated lineages within the eastern cape rocky (roos 2005). this raised concerns that these lineages could be facing a higher risk of extinction as they may potentially have narrower geographic ranges than currently recognised for this species. however, as a result of the small sample sizes and large geographic sampling gaps, this study could not provide a clear determination of the number of lineages and their distribution ranges. the present study builds on the findings of roos (2005), by collecting samples from all known extant populations of the eastern cape rocky to (1) determine the number of unique lineages within this species and (2) provide more accurate distribution ranges of the lineages. we discuss critical future research directions and conservation options to ensure the long-term persistence of the lineages identified within the eastern cape rocky. materials and methods sample collection the present study used data that were collected between 2010 and 2017. the samples were collected from 11 localities in the kowie, great fish, keiskamma, igoda and nahoon river systems, representing all the known remnant populations of s. bainsii (table 1; figure 1). sampling was done using fyke nets, seine nets and electric fishing. because s. bainsii is a threatened species, only fin clips were collected for the present study, and all sampled fish were returned to their habitat alive. table 1: sequence numbers and locality details for the lineages identified within sandelia bainsii. deoxyribonucleic acid extraction, amplification and sequencing laboratory work and sequencing were conducted at the aquatic genomic research platform, nrf-south african institute for aquatic biodiversity (saiab). deoxyribonucleic acid (dna) was extracted from 21 fin clip samples using the salting-out method (sunnucks & hales 1996). a fragment of the mitochondrial 16s rrna gene was amplified using the primer pair 16sar and 16sbr (palumbi, 1996) and sequenced using the forward primer (16sar). each pcr mixture (25 µl) contained 1 × buffer, 2.5 mm mgcl2, 0.8 mm deoxyribonucleotide triphosphates (dntps), 0.2 µm of each primer and 0.5 u taq polymerase and dna template (final concentration, 4.3 ng/µl – 9.5 ng/µl). the reaction volume was adjusted with dh2o to a final volume of 25 µl, depending on the volume of the dna template used. the amplification profile was initial denaturing at 95 °c for 3 min, followed by 35 cycles of denaturing at 95 °c for 50 s, annealing at 50 °c for 30 s and extension at 72 °c for 50 s. the reaction was then completed by a final extension at 72 °c for 10 min. the products were purified using an exosap method (thermofisher scientific). the purified products were then cycle sequenced using abi (applied biosystems, austin, texas) big dye version 3.1 cycle sequencing. sequencing was done using an abi hitachi 3500 genetic analyser at the nrf-saiab. data analysis the sequences were assembled and manually edited to equal lengths using seqman version 7.2.1 (dna star lasergene segman pro) and then aligned in clustal x2 (larkin et al. 2007). jmodeltest (darriba et al. 2012) was used to select the best-fit model of nucleotide evolution based on the akaike information criterion (aic) as implemented on the cipres science gateway (miller, pfeiffer & schwartz 2010). a maximum likelihood (ml) tree was generated using raxml version 8.2.6 (stamatakis 2014), performed on the cipres science gateway (miller et al. 2010). sandelia capensis, ctenopoma kingsleyae, ctenopoma patherici and ctenopoma occelatum were used as out-groups. to further assess the genealogical relationships between sequences among the populations of s. bainsii, a haplotype network was generated using the statistical parsimony method implemented in the program tcs (clement posada & crandall 2000). we used paup* (swofford 2002) to estimate the model-corrected genetic distances using the three parameter model (tpm2uf+i) (kimura, 1981) model of nucleotide evolution. results and discussion the edited alignment of 21 mitochondrial dna (mtdna) 16s sequences was 439 bp in length with 13 polymorphic sites that defined four unique haplotypes (figure 2). the tpm2uf + i (kimura 1981) was selected as the best model of sequence evolution. the ml tree and the templeton, crandall and sing (tcs) haplotype network revealed strong geographic structuring within s. bainsii, with the sequences split into three distinct lineages (figure 2). these lineages are herein referred to as sandelia sp. ‘bainsii kowie’, which is confined to the kowie and great fish river systems, sandelia sp. ‘bainsii keiskamma’, which is confined to the keiskamma river system and sandelia sp. ‘bainsii buffalo’, which occurs in the buffalo and igoda river systems (figure 2). these findings are consistent with those of roos (2005), who identified two lineages corresponding to sandelia sp. ‘bainsii kowie’ and sandelia sp. ‘bainsii buffalo’. however, roos’ (2005) study did not include samples from the keiskamma river system. all samples from the kowie and great fish river systems comprised a single haplotype (figure 2), suggesting that this lineage could have experienced recent range expansion, possibly because of human-mediated translocation, palaeodrainage connections, river captures or intermittent connection of low-drainage divides as inferred for other stream fishes in the genera galaxias, pseudobarbus and sandelia in the cfe (bronaugh et al. 2020; chakona et al. 2013; swartz et al., 2009). similarly, the sharing of a haplotype between the buffalo and igoda river systems also suggests recent connectivity or range expansion between these systems. the existence of these geographically isolated lineages clearly indicates the need for studies that, for example, use multiple genetic markers to determine the patterns of gene flow and investigate the ecology and population dynamics for each lineage. this would provide a better understanding of the mechanisms that have shaped the evolutionary history of s. bainsii sensu lato, as well as inform revision of the freshwater fish sanctuaries in south africa (nel et al. 2011). figure 2: maximum likelihood phylogeny (a) and tcs haplotype network (b) showing the existence of three lineages (sandelia sp. ‘bainsii kowie’, sandelia sp. ‘bainsii keiskamma’ and sandelia sp. ‘bainsii buffalo’) within the eastern cape rocky, sandelia bainsii. the river systems where each of these lineages were recorded are indicated. the low haplotype diversity observed in the present study suggests that population decline (cambray 1996) could have reduced genetic diversity within the eastern cape rocky. advances in molecular techniques that allow extraction and sequencing of dna from formalin-preserved specimens would provide an opportunity to leverage sequence data from old historical specimens to investigate changes in genetic diversity over time and the possibility of uncovering species or lineages that are now extinct in the wild (ruane & austin 2017). the existence of comprehensive samples of s. bainsii within the nrf-saiab national fish collection facility offers a unique opportunity for future studies to investigate the changes in the genetic diversity of the eastern cape rocky over time and identify possible causes for those changes. the range of genetic divergence values between sandelia sp. ‘bainsii kowie’ and sandelia sp. ‘bainsii buffalo’ (2.42% – 2.73%) is consistent with that found for interspecific mtdna sequence divergences in a number of fish genera (ward 2009) and was also comparable to interspecific differences among some species in the genus ctenopoma (see table 2), which is the sister genus to sandelia (rüber, britz & zardoya 2006). our results highlight the need for further studies that integrate genetic (both mitochondrial and nuclear sequences), morphological, osteological, ecological and other biological characteristics (e.g. puillandre et al. 2012) to determine the taxonomic status of the lineages identified within s. bainsii. a recent study by bronaugh et al. (2019) also revealed the existence of at least three deeply divergent lineages and substantial intralineage genetic and geographic structuring within s. capensis, indicating that the current taxonomy obscures the diversity of these anabantid fishes that are endemic to the cfe of south africa. table 2: mitochondrial 16s ribosomal ribonucleic acid sequence divergence estimates (%) among the three lineages identified within the eastern cape rocky, sandelia sp. ‘bainsii kowie’, sandelia sp. ‘bainsii keiskamma’ and sandelia sp. ‘bainsii buffalo’ sandelia bainsii was described by castelnau (1861) based on samples that were collected from the kowie river system. in the same year, günther (1861) described another species, ctenopoma microlepidotum, with the type locality details of the holotype vaguely presented as ‘freshwaters of the cape of good hope, south africa’. this species was subsequently synonymised with s. bainsii (skelton 2018). morphological examination of the types of s. bainsii and c. microlepidotum is currently ongoing to determine whether they are conspecific and to evaluate whether sandelia sp. ‘bainsii kowie’, sandelia sp. ‘bainsii keiskamma’ and sandelia sp. ‘bainsii buffalo’ constitute distinct taxonomic entities. the discovery of three genetically distinct and historically isolated lineages within the eastern cape rocky, which is already listed as endangered by the iucn (chakona et al. 2018b), supports the growing need to expedite discovery and documentation of biodiversity because the existence of hidden diversity in many taxa hampers conservation efforts (e.g. bickford et al. 2007). as recommended by moritz (1994), to preserve the evolutionary processes that shaped the genetic patterns within s. bainsii, the three lineages need to be managed separately. this requires close collaboration between researchers, conservationists and local stakeholders (landowners) to develop a sustainable partnership programme that will promote improved catchment management practices for sustainable ecological functioning of the rivers and to protect critical habitats for these lineages. previous and ongoing field surveys have reported significant declines in the historical distribution range of s. bainsii, with the population in the kowie river system feared to be nearing extinction, as only two specimens were recorded from comprehensive surveys that were conducted between 2010 and 2017 (figure 1). the eastern cape rocky appears to be highly sensitive to poor water quality, as it has not been recorded from heavily polluted sections of the kowie and bloukranz rivers where it was historically abundant. all remnant populations of s. bainsii are highly fragmented, because they are isolated by instream impoundments and invasion of the mainstem sections of the rivers by non-native fishes, which may prevent connectivity and hamper gene flow. field observations suggest that most of these remnant populations are likely to be represented by small population sizes, as they were only recorded in short stretches of the streams where they were found. empirical data show that population fragmentation leads to rapid deterioration in genetic diversity in small populations because of genetic drift and inbreeding (bessert & ortí 2008; pavlova et al. 2017). as genetic diversity underpins the ability of populations to persist and adapt to environmental changes (hughes, schmidt & finn 2009), there are concerns that the small isolated populations of sandelia sp. ‘bainsii kowie’, sandelia sp. ‘bainsii keiskamma’ and sandelia sp. ‘bainsii buffalo’ could be at high risk of extinction as a result of the possibility of inbreeding depression, loss of fitness and reduced adaptive potential. future studies should focus on assessing the implications of population fragmentation on genetic diversity and the evolutionary potential of each of these newly identified lineages. in addition, studies should determine effective conservation strategies, such as securing remnant populations to prevent invasion by non-native fishes, rehabilitation of degraded habitats and genetic rescue through assisted gene flow (see pavlova et al. 2017), where such measures are deemed necessary interventions. acknowledgements the authors hereby acknowledge the use of the equipment provided by the nrf-saiab molecular genetic laboratory and the nrf-saiab collection management centre. greg brett from east london museum, leah sloman, apelele zonda and bosupeng motshegoa are thanked for their assistance with fieldwork; leah sloman for generating the sequences. competing interests the authors have declared that no competing interests exist. authors’ contributions this work formed part of p.p.m.’s hons project; a.c. secured research funding; a.c. and p.h.s. conceived the research; a.c., p.p.m. and w.t.k. conducted the fieldwork; a.c. and g.g. performed the genetic analyses. all authors contributed equally to writing and revising the manuscript. ethical consideration this research was carried out following the evaluation and approval of the sampling protocols by the national research foundation – south african institute for aquatic biodiversity animal ethics committee (reference 2014/03). permits to carry out this research were obtained from the department of economic development, environmental affairs and tourism (eastern cape province) (permit numbers cro 88/15cr and cro 44/18cr). funding information this work was supported by the national research foundation (nrf) of south africa under the foundational biodiversity information programme (fbip) through biodiversity surveys in priority inland areas (ibip) grant (grant no. ibip-bs13100251309) through a joint initiative with the department of science of technology, the nrf and the south african national biodiversity institute. data availability statement the sequences generated from this study were submitted to genbank. disclaimer the authors acknowledge that the opinions, findings and conclusions or recommendations expressed in this publication generated by nrf-supported research belong to the authors and that the nrf accepts no liability whatsoever in this regard. references bessert, m.l. & ortí, g., 2008, ‘genetic effects of habitat fragmentation on blue sucker populations in the upper missouri river (cycleptus elongatus lesueur, 1918)’, conservation genetics 9, 821–832. https://doi.org/10.1007/s10592-007-9401-4 bickford, d., lohman, d.j., navjot, s.s., ng, p.k.l., meier, r., winker, k. et al., 2007, ‘cryptic species as a window on diversity and conservation’, trends in ecology and evolution 22(3), 148–155. https://doi.org/10.1016/j.tree.2006.11.004 bronaugh, w.m., swartz, e.r. & sidlauskas, b.l., 2020, ‘between an ocean and a high place: coastal drainage isolation generates endemic cryptic species in the cape kurper sandelia capensis (anabantiformes: anabantidae), cape region, south africa’, journal of fish biology 96(5), 1087–1099. https://doi.org/10.1111/jfb.14182 cambray, j.a., 1996, ‘threatened fishes of the world: sandelia bainsii castelnau, 1861 (anabantidae)’, environmental biology of fishes 1(3), 159–168. chakona, a. & skelton, p.h., 2017, ‘a review of the pseudobarbus afer (peters 1864) species complex (teleostei, cyprinidae) in the eastern cape fold ecoregion of south africa’, zookeys 657, 109–140. https://doi.org/10.3897/zookeys.657.11076 chakona, a. & swartz, e.r., 2013, ‘a new redfin species, pseudobarbus skeltoni (cyprinidae, teleostei), from the cape floristic region, south africa’, zootaxa 3686(5), 565–577. https://doi.org/10.11646/zootaxa.3686.5.5 chakona, a., kadye, w.t., bere, t., mazungula, d.n. & vreven, e., 2018a, ‘evidence of hidden diversity and taxonomic conflicts in five stream fishes from the eastern zimbabwe highlands freshwater ecoregion’, zookeys 768, 69–95. https://doi.org/10.3897/zookeys.768.21944 chakona, a., sifundza, d. & kadye, w.t., 2018b, ‘sandelia bainsii, the iucn red list of threatened species 2018, e.t19889a99447325’, https://doi.org/10.2305/iucn.uk.2018-1.rlts.t19889a99447325.en chakona, a., swartz, e.r. & gouws, g., 2013, ‘evolutionary drivers of diversification and distribution of a southern temperate stream fish assemblage: testing the role of historical isolation and spatial range expansion’, plos one 8(8), e70953. https://doi.org/10.1371/journal.pone.0070953 chakona, a., swartz, e.r. & skelton, p.h., 2014, ‘a new species of redfin (teleostei, cyprinidae, pseudobarbus) from the verlorenvlei river system, south africa’, zookeys 453, 121–137. https://doi.org/10.3897/zookeys.453.8072 clement, m., posada, d. & crandall, k.a., 2000, ‘tcs: a computer program to estimate gene genealogies’, molecular ecology 9(10), 1657–1659. https://doi.org/10.1046/j.1365-294x.2000.01020.x darriba, d., taboada, g.l., doallo, r., posada, d., 2012, ‘jmodeltest 2: more models, new heuristics and parallel computing’, nature methods 9(8), 772. https://doi.org/10.1038/nmeth.2109 gunther, a. c., 1861, catalogue of acanthopterygian fishes in the british museum 3., addenda et corrigenda, trustees of the british museum, london. hughes, j.m., schmidt, d.j. & finn, d.s., 2009, ‘genes in streams: using dna to understand the movement of freshwater fauna and their riverine habitat’, bioscience 59(7), 573–583. https://doi.org/10.1525/bio.2009.59.7.8 kimura, m., 1981, ‘estimation of evolutionary distances between homologous nucleotide sequences’, proceedings of the national academy of sciences usa 78(1), 454–458. https://doi.org/10.1073/pnas.78.1.454 larkin, m.a., blackshields, g., brown, n.p., chenna, r., mcgettigan, p.a., mcwilliam, h. et al., 2007, ‘clustal w and clustal x version 2.0’, bioinformatics 23(21), 2947–2948. https://doi.org/10.1093/bioinformatics/btm404 maake, p.a., gon, o. & swartz, e.r., 2014, ‘descriptions of three new species of marcusenius gill, 1862 (teleostei: mormyridae) from south africa and mozambique’, zootaxa 3780(3), 455–480. https://doi.org/10.11646/zootaxa.3780.3.2 miller, m.a., pfeiffer, w. & schwartz, t., 2010, ‘creating the cipres science gateway for inference of large phylogenetic trees’, in proceedings of the gateway computing environments workshop (gce), new orleans, la, pp. 1–8. moritz, c., 1994, ‘defining ‘evolutionary significant units’ for conservation’, trends in ecology & evolution 9(10), 373–375. https://doi.org/10.1016/0169-5347(94)90057-4 nel, j.l., driver, a., strydom, w.f., maherry, a., petersen, c., hill, l. et al., 2011, ‘atlas of freshwater ecosystem priority areas in south africa: maps to support sustainable development of water resources’, wrc report no. tt 500/11, water research commission, gezina, south africa palumbi, s.r., 1996, ‘nucleic acids ii: the polymerase chain reaction’, in d.m. hillis, c. moritz & b.k. mable (eds.), molecular systematics, pp. 205–247, sinauer associates, inc, sunderland, massachusetts. pavlova, a., beheregaray, l.b., coleman, r., gilligan, d., harrisson, k.a., ingram, b.a. et al., 2017, ‘severe consequences of habitat fragmentation on genetic diversity of an endangered australian freshwater fish: a call for assisted gene flow’, evolutionary applications 10(6), 531–550. https://doi.org/10.1111/eva.12484 puillandre, n., modica, m.v., zhang, y., sirovich, l., boisselier, m.c., cruaud, c. et al., 2012, ‘large-scale species delimitation method for hyperdiverse groups’, molecular ecology 21(11), 2671–2691. https://doi.org/10.1111/j.1365-294x.2012.05559.x roos, h., 2005, ‘genetic diversity in the anabantids sandelia capensis and sandelia bainsii: a phylogeography and phylogenetic investigation’, msc thesis, university of pretoria, south africa. ruane, s. & austin, c.c., 2017, ‘phylogenomics using formalin-fixed and 100+ year-old intractable natural history specimens’, molecular ecology resources 17(5), 1003–1008. https://doi.org/10.1111/1755-0998.12655 rüber, l., britz, r. & zardoya, r., 2006, ‘molecular phylogenetics and evolutionary diversification of labyrinth fishes (perciformes: anabantoidei)’, systematic biology 55(3), 374–397. https://doi.org/10.1080/10635150500541664 skelton, p.h., 2001, a complete guide to freshwater fishes of southern africa, southern book publishers pty ltd., jlb smith institute of ichthyology, grahamstown. skelton, p.h., 2018, ‘sir andrew smith – his contributions and connections to freshwater ichthyology in southern africa’, transactions of the royal society of south africa, 73(1), 42–55. https://doi.org/10.1080/0035919x.2017.1383322 stamatakis, a., 2014, ‘raxml version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies’ bioinformatics 30(9), 1312–1313. https://doi.org/10.1093/bioinformatics/btu033 sunnucks, p. & hales, d.f., 1996, ‘numerous transposed sequences of mitochondrial cytochrome oxidase i-ii in aphids of the genus sitobion (hemiptera: aphididae)’, molecular biology and evolution 13(3), 510–524. https://doi.org/10.1093/oxfordjournals.molbev.a025612 swartz, e.r., skelton, p.h. & bloomer, p., 2009, ‘phylogeny and biogeography of the genus pseudobarbus (cyprinidae): shedding light on the drainage history of rivers associated with the cape floristic region’, molecular phylogenetics and evolution 51(1), 75–84. https://doi.org/10.1016/j.ympev.2008.10.017 swofford, d.l., 2002, paup*, phylogenetic analysis using parsimony (*and other methods), version 4, sinauer associates, sunderland, ma. ward, r.d., 2009, ‘dna barcode divergence among species and genera of birds and fishes’, molecular ecology resources 9(4), 1077–1085. https://doi.org/10.1111/j.1755-0998.2009.02541.x wishart, m., hughes, j., stewart, b. & impson, d., 2006, ‘extreme levels of intra-specific divergence among cape peninsula populations of the cape galaxias, galaxias zebratus castelnau 1861, reveals a possible species complex’, african journal of aquatic science 31(1), 99–106. https://doi.org/10.2989/16085910609503876 article information authors: mohamed elhag1 ahmad k. hegazy2 abdulrahman a. alatar3 mohamed faisal3 magdi el-bana4 jarbou a. bahrawi1 amal a.m. al-ghamdi5 affiliations: 1department of hydrology and water resources management, king abdulaziz university, saudi arabia 2department of botany and microbiology, cairo university, egypt 3department of botany and microbiology, king saud university, saudi arabia 4department of botany, port said university, egypt 5department of botany, environment program, king abdul aziz university, saudi arabia correspondence to: mohamed elhag email: melhag@kau.edu.sa postal address: po box 80208, jeddah 21589, saudi arabia dates: received: 27 apr. 2015 accepted: 10 sept. 2015 published: 30 nov. 2015 how to cite this article: elhag, m., hegazy, a.k., alatar, a.a., faisal, m., el-bana, m., bahrawi, j.a. et al., 2015, ‘population demography and global sensitivity analysis of avicennia marina on the eastern and western coasts of saudi arabia’, koedoe 57(1), art. #1317, 9 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1317 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. population demography and global sensitivity analysis of avicennia marina on the eastern and western coasts of saudi arabia in this original research... open access • abstract • introduction • materials and methods    • study area    • plant populations    • basics of variance-based sensitivity analysis    • the concept of global sensitivity analysis    • global sensitivity analysis procedures • results and discussion • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ mangrove ecosystems are one of the habitats that host high environmental diversity at the level of physical, geomorphological and biological features in arid regions. in saudi arabia, mangrove ecosystems are heavily threatened by both natural hazards and human pressure. the total estimated area of mangroves in safwa al khurais, saudi arabia, is approximately 20 000 ha in extent and comprises two species: avicennia marina and rhizophora mucronata. they supply detritus to the marine food web and play a significant role in the conservation of biological diversity. the main objective of this study was to analyse the demographic population sensitivity of a. marina in two representative sites on the red sea and the eastern coast of saudi arabia. a sensitivity analysis was used to assess the contributions of the inputs to the total uncertainty in the analysis outcomes. demographic features affecting mangroves in saudi arabia are complex and include various aspects. the phenological phase, tree size, density, cover, number of dead trees and pneumatophore characteristics were considered in this study. a comparative analysis of gaussian process emulators for performing a global sensitivity analysis was used to conduct a variance-based sensitivity analysis to identify which uncertain inputs drive the output uncertainty. the results showed that the interconnections between different demographic features were predictable, but that the extent of the sensitivity was uncertain. findings from the current study are anticipated to contribute significantly towards an inclusive mangrove demographic features assessment, and towards the subsequent conservation of mangroves in saudi arabia. conservation implications: the findings of the current research are discussed in light of the application of sensitivity analyses outputs in the conservation and management of mangrove ecosystems at a national level introduction top ↑ mangroves in saudi arabia are found in the form of fragmented populations or thin stands in many tidal areas of the red sea and the arabian gulf coasts. they consist mainly of two species: avicennia marina (forsk.) vierh. (locally known as qurum), which is the common species; and rhizophora mucronata lam. (locally known as qundel), which is limited to a few localities. the northerly limit of distribution of a. marina in the red sea is debah, saudi arabia, and extends south to the border with yemen (khan, kumar & muqtadir 2010; kumar, khan & muqtadir 2010). in the arabian gulf, mangroves are found in very small patches in the north, on jazeerat qurma (translated as mangrove island) and in tarut bay, where they form well-developed stands consisting of a. marina. although the mangroves of saudi arabia are not as luxurious as those on other tropical shores, they play similarly significant ecological and environmental roles. these locations are nurseries for numerous profitable fish species and they shelter coral reefs by retaining sediment loads from the periodic influx of rain water. because they grow in very different environments, mangroves are very sensitive to over-exploitation. the degradation of mangrove ecosystems has occurred in various parts of the region, due to excessive browsing by camels, overcutting, impeding rainwater from draining through valleys, pollution, and coastal construction (mandura 1997; mandura & khafaji 1993). human activities such as coastal urbanisation and related wastewater manage ment problems, industrialisation and related emissions, chemical pollutants, fishing activities and aquaculture development, tourism and the consequent increase of pressures on coastal resources are some of the main stresses introduced by humankind on coastal ecosystems (hegazy 1998; kumar et al. 2010). global sensitivity analysis (gsa) methods assign the output inconsistency to the inconsistency of the input parameters when they fluctuate over their entire uncertainty dimension (petropoulos et al. 2009). the sensitivity of input parameters is fundamentally based on the generation of samples dispersed across the parameter dimension of interest. extensive scholarly work of the obtainable gsa approaches and their applications is reported, for example, saltelli, chan and scott (2000), saltelli (2002), and saltelli et al. (2004). global sensitivity analysis is an influential method due to its capability to assimilate the influence of the input parameters over their whole range of inconsistency (saltelli, tarantola & chan 1999). global sensitivity analysis techniques are able to deliver quantitative assessments not only of the most sensitive model inputs, but also of the interactions of the model input parameters (schwieger 2004). this yields measureable evidence of the degree of difficulty of the model input–output interactions (petropoulos et al. 2009). the aim of this study is to analyse the uncertainty of the survival of a. marina based on the demographic features of five populations, located in the red sea and the arabian gulf of saudi arabia. this is useful as a guiding tool for the conservation and sustainable use of mangrove ecosystems in the region. materials and methods top ↑ study area five sites were identified for this study (figure 1). two sites are located on the red sea coast: al mudhaylif (19°30′76.1″ n, 40°56′45.6″ e) is located north of al qunfudhah city, and usfan (21°59′18.1″ n, 39°00′42.0″ e) is located north of jiddah city. the remaining three sites are located in the arabian gulf: sehat (26°30′12.8″ n, 50°02′35.0″ e) and safwa al khurais (26°39′43.8″ n, 49°59′15.8″ e) are located in tarut bay, and qurum island (27°07′56.4″ n, 49°29′13.0″ e) is located north of al jubail city. figure 1: the location of the five study sites in the red sea (usfan and al mudhaylif) and the arabian gulf (qurum island, sehat and safwa al khurais). plant populations measurements of the population characteristics of a. marina were recorded in the five study sites. plant density was estimated as the number of individuals per 100 m2 for seedlings and adult individuals. the number of dead or partially dead trees was counted. plant cover was measured by the line intercept method. the density of respiratory roots (pneumatophores) was estimated by counting the number of roots per m2. basics of variance-based sensitivity analysis sensitivity analysis approaches are categorised according to the outcome of the related sensitivity procedures into local or global methods, and methods that depend on or are independent of the model characteristics (schwieger 2004). the concept of global sensitivity analysis consistent with saltelli et al. (2000), gsa is the study of the relationship between the input and output of a model. in essence, gsa deals with the variation that corresponds to the uncertainties of the input magnitudes. moreover, input parameters are introduced to the uncertainties of the model parameters and to the overall model structure. the discrepancy of the input parameters results in discrepancies of the output magnitudes. the interconnections between speckled input and output are measured by various sensitivity measures that are the basis for model validation and optimisation (schwieger 2004). the broad practice of sensitivity analysis is shown in figure 2. global sensitivity analysis emphasises the variance-based techniques to quantitatively estimate the global variances of the outputs and to investigate the independent sensitivity of the input measures. figure 2: the general procedure for a sensitivity analysis. global sensitivity analysis procedures based on the monte carlo methods, sensitivity analysis methods include regression and correlation analyses as well as the analysis of the rank of the transformed data. the general procedure to estimate global sensitivity measures is based on the following equations: where: for non-correlated input additive models: according to schwieger (2004), this translates to an easy numerical interpretation of the sensitivity indices, because each si assures a direct measure for the portion of xi on the output variance . for non-additive models, the interactions amongst the input quantities within the model need to be considered. non-additive models require a comprehensive disintegration of the function y into summons of increasing order: the terms of higher order are estimated by taking other fixed input quantities into consideration: the estimation of higher order terms leads to the estimation of the total effects sti with respect to an input quantity xi, to be computed as follows: the corresponding total effect is computed as: consistently, a judgement between si and sti leads to a conclusion concerning the additivity of models with non-correlated inputs: results and discussion top ↑ the results from the sensitivity analysis focus specifically on the decomposition of variance (%) of the mean total variance in emulator output, where input parameters have been assumed to be non-correlated, normally distributed and varying within their whole range. a sensitivity analysis of mangrove demographic features was carried out on five different mangrove stands: two stands located on the west coast and three located on the east coast. both of the western mangrove stands showed a lower sensitivity to the demographic feature number of respiratory roots, where this was observed to be 0.05 and 7.5, respectively (see table 1). conversely, the stands located on the east coast showed a different sensitivity to the demographic features. in sehat stand, the number of dead trees was the least sensitive mangrove demographic feature. the safwa al khurais stand demonstrated that the average height of respiratory roots was the least sensitive, whilst on qurum island the number of respiratory roots was the least sensitive feature. the conclusion drawn from these findings implies interactions or dependency on the rest of the demographic features (hegazy 1998; holvoet et al. 2005). table 1: population characteristics and sensitivity analysis in the five study sites. histogram chart representations of the total effect of the sensitivity analysis demonstrated interand intra comparability differences, shown collaterally in figure 3. in the western region of the study area, the mangrove stand in usfan is considered to be more stable compared to the al mudhaylif region, where none of the demographic features count for over 33% individually (the average height of respiratory roots). in the eastern part of the study area, the mangrove stand on qurum island is considered to be the most sensitive stand. all of the demographic features were very sensitive and showed a higher uncertainty total variance, except for the average height of respiratory roots. this could be explained by the extensive anthropogenic activities taking places in and around the stand (kumar et al. 2010). figure 3: the sensitivity analysis total effect (%). in the usfan stand, the results indicated that the most sensitive demographic feature was the average height of respiratory roots, which accounted for 33%. the average height of respiratory roots of mangroves alone accounts for one third of the total effect of the sensitivity analysis. the higher total effect of the average height of respiratory roots explains the lower stability of such demographic features (elhag 2014; petropoulos et al. 2009). the total variance of the sensitivity analysis of mangrove seedlings was calculated to be 0.9%. such a minor value indicated a robust stability of the mangrove demographic feature in the study area. the next stand on the western side of saudi arabia, located at al mudhaylif, showed a relative stability compared to the stand located in usfan (according to the demographic features used in the current study). both the number of respiratory roots and the number of dead trees (8% for both) were considered to be the least sensitive demographic features according to petropoulos et al. (2009) and elhag (2014). on the eastern side of saudi arabia, three different stands of mangroves were considered for sensitivity analysis. sehatstand is a moderately stable stand, and both the number of dead trees and average height of respiratory roots accounted for 3% of the total variance. the low total variance percentages represent more stable demographic features, which correspond to environmental variability (holvoet et al. 2005). within the same geographical region, on the eastern side of the current study, both mangrove stands of safwa al khurais and qurum island were found to be extremely sensitive to environmental impacts. the number of seedlings (38%) and tree density of mangroves (39%) were found to be the most unstable demographic features in safwa al khurais and on qurum island, respectively. the average height and number of respiratory roots were the most stable demographic features in safwa al khurais and on qurum island, respectively. the uncertainty analysis of mangrove demographic features of the usfan area is shown in figure 4. there were three mangrove demographic features in usfan. density, tree cover and crown diameter showed a higher precision of uncertainty towards higher corresponding demographic values. figure 4: usfan demographic features sensitivity analysis. (a) density (100 m2); (b) cover (%); (c) tree height (m); (d) crown diameter (m); (e) number of dead trees (100 m2); (f) number of seedlings (100 m2); (g) number of respiratory roots (100 m2) and (h) average height of respiratory roots (m). the values of mangrove demographic features were proportionally related to the corresponding uncertainty as the main effect, except for the number of seedlings, which was inversely proportionate to its uncertainty value. the number of respiratory roots of mangroves in the usfan area displayed a steady behaviour of uncertainty levels, with different values of the total number of respiratory roots. figure 5 shows that the interactions of the mangrove demographic features of the al mudhaylif area were not strong enough to suggest a variance dependency and moderate interactions uncertainty (elhag 2014; petropoulos et al. 2009). the mangroves in al mudhaylif expressed more uncertainty levels with different demographic features. uncertainty levels of mangrove demographic features in al mudhaylif were higher when compared to usfan. higher precisions of uncertainty were expressed in six different demographic features, either in a higher or lower corresponding demographic feature value. all of the demographic features were proportionally related to its corresponding uncertainty as the main effect. moreover, the number of dead mangroves was inversely proportionate to its uncertainty value (elhag 2014). figure 5: al mudhaylif demographic features sensitivity analysis: (a) density (100 m2); (b) cover (%); (c) tree height (m); (d) crown diameter (m); (e) number of dead trees (100 m2); (f) number of seedlings (100 m2); (g) number of respiratory roots (100 m2) and (h) average height of respiratory roots (m). a gsa was also applied to the eastern mangrove stands. sehat, safwa al khurais and qurum island showed that sehatis was the most stable mangrove stand in the area of investigation (east and west). the mangrove stand at sehatstand showed good resilience to environmental impacts (figure 6). the tree cover of mangroves and height of mangroves in safwa al khurais showed sensitive behaviour to environmental impacts (figure 7), due its higher contributions to the two demographic features of the total variance of the gsa. mangrove trees confined to qurum island were considered to be the most degraded mangrove stand because all of the demographic features implemented in the gsa (except the average height of respiratory roots) were remarkably unstable to the surrounding environmental impacts (figure 8). figure 6: sehat demographic features sensitivity analysis: (a) density (100 m2); (b) cover (%); (c) tree height (m); (d) crown diameter (m); (e) number of dead trees (100 m2); (f) number of seedlings (100 m2); (g) number of respiratory roots (100 m2) and (h) average height of respiratory roots (m). figure 7: safwa al khurais demographic features sensitivity analysis: (a) density (100 m2); (b) cover (%); (c) tree height (m); (d) crown diameter (m); (e) number of dead trees (100 m2); (f) number of seedlings (100 m2); (g) number of respiratory roots (100 m2) and (h) average height of respiratory roots (m). figure 8: qurum island demographic features sensitivity analysis (a) density (100 m2); (b) cover (%); (c) tree height (m); (d) crown diameter (m); (e) number of dead trees (100 m2); (f) number of seedlings (100 m2); (g) number of respiratory roots (100 m2) and (h) average height of respiratory roots (m). conclusion top ↑ the gsa of the mangroves were measured against eight different demographic features: density, cover, tree height, crown diameter, number of dead trees, number of seedlings, number of respiratory roots and average height of respiratory roots. the gsa delivered quantitative and model-independent sensitivity measures to each of the input factors, and to the input factors collectively, in response to the simulated outputs under consideration. the results of this study show the model concept to be sufficiently sensitive to represent the behaviour of the natural systems. the sensitivity analysis confirms that demographic features were alternately sensitive to different locations of mangrove stands and that input parameters related directly to the estimated variables derived from the uncertainty analysis. the purpose of implementing a gsa approach was to recognise variance associated with different input measures. the gsa model is independent of the features of the investigated model. conducted findings shall mainly be used for better model performance, for example, comparing different demographic features from different locations. immediate remediation and restoration techniques need to be applied urgently in order to conserve the mangrove stands, especially in the eastern section of the study area. acknowledgements top ↑ this work is funded by the project number 11-env1756-02, npst-kacst, saudi arabia, entitled ‘conservation and utilisation of mangrove ecosystems in saudi arabia: from knowledge to development’. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions a.k.h. (king saud university) proposed the project idea, participated in all the field work, data analysis and writing of the manuscript. m.e. (king abdulaziz university) and j.a.b. (king abdulaziz university) were responsible for the data analysis, and wrote most of the manuscript. m.e.b. (port said university) participated in the field work and data collection. a.a-g. (king abdulaziz university) made conceptual contributions in the writing of the manuscript. a.a.a. (king saud university) and m.f. (king saud university) performed some of the experiments. references top ↑ elhag, m., 2014, ‘sensitivity analysis assessment of remotely based vegetation indices to improve water resources management’, environment, development and sustainability 16(6), 1209–1222. http://dx.doi.org/10.1007/s10668-014-9522-0 hegazy, a.k., 1998, ‘perspectives on phenology, survival and productivity of avicennia marina in the qatari coasts, arabian gulf’, journal of arid environments 40, 417–429. holvoet, k., van griensven, a., seuntjens, p. & vanrolleghem, p.a., 2005, ‘sensitivity analysis for hydrology and pesticide supply towards the river in swat’, physics and chemistry of the earth, parts a/b/c 30, 518–526. http://dx.doi.org/10.1016/j.pce.2005.07.006 khan, m.a., kumar, a. & muqtadir, a., 2010, ‘distribution of mangroves along the red sea coast of the arabian peninsula: part 2. the southern coast of western saudi arabia’, earth science india 3(3), 154–162. kumar, a., khan, m.a. & muqtadir, a., 2010, ‘distribution of mangroves along the red sea coast of the arabian peninsula: part 1. the northern coast of western saudi arabia’, earth science india 3(1), 28–42. mandura, a.s., 1997, ‘a mangrove stand under sewage pollution stress: red sea’, mangroves and salt marshes 1(4), 255–262. http://dx.doi.org/10.1023/a:1009927605517 mandura, a.s. & khafaji, a.k., 1993, ‘human impact on mangrove of khor farasan island, southern red sea coast of saudi arabia’, in h. leith & h. al masoom (eds.), towards the rational use of high salinity tolerant plants, vol. 1, pp. 353–261, kluwer academic publishers, dordrecht. petropoulos, g., wooster, m.j., carlson, t.n., kennedy, m.c. & scholze, m., 2009, ‘a global bayesian sensitivity analysis of the 1d simsphere soil–vegetation– atmospheric transfer (svat) model using gaussian model emulation’, ecological modelling 220(19), 2427–2440. http://dx.doi.org/10.1016/j.ecolmodel.2009.06.006 saltelli, a., 2002, ‘sensitivity analysis for importance assessment’, risk analysis 22(3), 549–590. http://dx.doi.org/10.1111/0272-4332.00040 saltelli, a., chan, k. & scott, e.m., 2000, ‘sensitivity analysis’, in wiley series in probability and statistics, p. 467, wiley, new york. saltelli, a., tarantola, s., campologno, f. & ratto, m., 2004, sensitivity analysis in practice: a guide to assessing scientific models, p. 217, john wiley & sons, ltd., chichester. saltelli, a., tarantola, s. & chan, k.p.s., 1999, ‘a quantitative model-independent method for global sensitivity analysis of model output’, technometrics 41(1), 39–56. http://dx.doi.org/10.1080/00401706.1999.10485594 schwieger, v., 2004, ‘variance-based sensitivity analysis for model evaluation in engineering surveys’, ingeo 2004 and fig regional central and eastern european conference on engineering surveying, bratislava, slovakia, november 11–13, 2004. abstract acknowledgements references about the author(s) charles w. helm african centre for coastal palaeoscience, nelson mandela university, port elizabeth, south africa hayley c. cawthra african centre for coastal palaeoscience, nelson mandela university, port elizabeth, south africa richard m. cowling african centre for coastal palaeoscience, nelson mandela university, port elizabeth, south africa jan c. de vynck african centre for coastal palaeoscience, nelson mandela university, port elizabeth, south africa martin g. lockley african centre for coastal palaeoscience, nelson mandela university, port elizabeth, south africa curtis w. marean african centre for coastal palaeoscience, nelson mandela university, port elizabeth, south africa mark g. dixon african centre for coastal palaeoscience, nelson mandela university, port elizabeth, south africa carina j.z. helm african centre for coastal palaeoscience, nelson mandela university, port elizabeth, south africa willo stear african centre for coastal palaeoscience, nelson mandela university, port elizabeth, south africa guy h.h. thesen african centre for coastal palaeoscience, nelson mandela university, port elizabeth, south africa jan a. venter african centre for coastal palaeoscience, nelson mandela university, port elizabeth, south africa citation helm, c.w., cawthra, h.c., cowling, r.m., de vynck, j.c., lockley, m.g., marean, c.w. et al., 2021, ‘protecting and preserving south african aeolianite surfaces from graffiti’, koedoe 63(1), a1656. https://doi.org/10.4102/koedoe.v63i1.1656 short communication protecting and preserving south african aeolianite surfaces from graffiti charles w. helm, hayley c. cawthra, richard m. cowling, jan c. de vynck, martin g. lockley, curtis w. marean, mark g. dixon, carina j.z. helm, willo stear, guy h.h. thesen, jan a. venter received: 17 sept. 2020; accepted: 01 feb. 2021; published: 11 mar. 2021 copyright: © 2021. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract pleistocene aeolianite surfaces on the south african coastline, which occur in national parks, other protected areas, as well as unprotected areas, are of profound scientific, cultural, palaeoenvironmental and heritage importance. a threat is posed to these surfaces by the increasing presence of graffiti, which may deface or destroy fossil tracksites and other evidence of events that transpired on these surfaces when they were composed of unconsolidated sand tens of thousands of years ago. increased awareness of the importance of this heritage resource is desirable, along with the development of strategies to prevent further damage. keywords: protected areas; management; pleistocene; conservation; graffiti. aeolianite surfaces on the south african coastline are of profound scientific, cultural, palaeoenvironmental and heritage importance, and a threat is posed to these surfaces by graffiti artists. the south african aeolianites are not only an outstanding, global example of this rock type, but are also sensitive palaeoenvironmental indicators. they happen to occur in an area of critical importance for palaeoanthropology and, hence, in the study of human origins. research on the cape south coast, for example, has documented the oldest known working of ochre and adornment through its use (marean et al. 2007:905–908); ornamentation in the form of perforated nassarius krausssianus shells (d’errico et al. 2005:3–24); bone awl and bone tool manufacturing (henshilwood et al. 2001:632–678); development of microlithic technology (brown et al. 2012:590–593); heat treatment of stone tools (brown et al. 2009:859–862); early art, or the use of abstract symbols, as engravings in ochre (e.g. henshilwood et al. 2002:1278–80), a drawing on a silcrete flake (henshilwood et al. 2018:115–118), and engravings on ostrich eggshells (henshilwood et al. 2014:284–303); the first use of marine or intertidal resources by humans, and the first systematic exploitation of these resources (marean et al. 2007:905–908; marean 2010:425–443). in combination, these innovations strongly indicate the emergence of cognitive complexity in our species (wadley 2015:155–226). the coastal aeolianites are cemented pleistocene dune surfaces, and the cemented foreshore deposits represent pleistocene beach surfaces. they crop out intermittently along the south african coast, and occur in, for example, west coast national park, garden route national park, addo elephant national park, adjacent protected areas such as goukamma nature reserve, and within numerous unprotected areas. whilst the majority have been dated to ~90 000–130 000 years (e.g. bateman et al. 2011:63–81; cawthra et al. 2018:156–172; roberts et al. 2008:261–279), the age of these rocks extends from ~35 000 years at robberg (carr et al. 2019:1–15) to ~400 000 years at dana bay (roberts et al. 2012:226–237), and as much as 720 000 years in a cave at pinnacle point (pickering et al. 2013:39–52). since 2007, we have, through the cape south coast ichnology project, documented more than 250 fossil vertebrate tracksites in these rock surfaces (helm et al. 2020a:1–15). thus, we know that they have the capacity to faithfully record, sometimes in exquisite detail, the events that transpired on them when they were composed of unconsolidated sand tens of thousands of years ago. some tracks represent the traces of extinct members of the pleistocene megafauna (e.g., the long-horned buffalo, giant cape zebra and cape subspecies of the african lion). others provide evidence, which is not available through the traditional body fossil record, of the presence of creatures that no longer inhabit the region, such as hatchling sea turtles (lockley et al. 2019:626–640), crocodiles, giraffe and large avian forms, with resulting palaeoenvironmental inferences (helm et al. 2020a:1–15). these tracksites would have been situated at the margin of the vast, now-submerged, palaeo-agulhas plain, harbouring an extinct ecosystem that was of pivotal significance for early modern human beings (marean, cowling & franklin 2020:1–17). in addition, these exceptional rocks provide evidence of the activity of our ancestors: to date, six hominin tracksites have been identified (helm et al. 2018a:1–13, 2020b:1–13; roberts 2008:190–207), providing the oldest known ichnological evidence of homo sapiens, thereby complementing the documented middle stone age archaeological record from the cape coast. sometimes, erosive forces acting on tracksites lead to the development of remarkably beautiful sculpted forms. aeolianite surfaces on the southern african coastline, therefore, are of profound scientific, cultural, palaeoenvironmental, heritage and aesthetic importance. unfortunately, their relatively soft, friable nature makes them suitable canvases for graffiti artists. a hammer and chisel can easily deface or completely destroy these surfaces, and their scientific, heritage and aesthetic attributes. repeat visits indicate that this is an active problem, and that the prevalence of graffiti is increasing. possibly, the presence of graffiti acts as an incitement to other potential graffiti artists. a notable example is from kraalbaai in west coast national park, where roberts (2008:190–207) identified hominin tracks now famously known as ‘eve’s footprints’, dated to ~117 000 years. this surface was found in an area with multiple examples of graffiti (figure 1a). a graffiti artist had reached the surface containing the hominin tracks first; through good fortune, the graffiti narrowly missed defacing the tracks. nonetheless, a decision was taken that airlifting the track-bearing aeolianite slab to the iziko museum in cape town constituted the most secure means of preservation. figure 1: (a) dense concentrations of graffiti on aeolianites at kraalbaai, west coast national park; (b) graffiti on a cave floor surface at brenton-on-sea, rendering what may have been hominin tracks (two of which are indicated with arrows) undecipherable; (c) graffiti on an aeolianite surface in goukamma nature reserve, metres from an important hominin tracksite; (d) graffiti on a loose aeolianite slab exhibiting well-preserved invertebrate burrow traces; (e) graffiti crossing a lagomorph trackway in garden route national park; (f) bovid tracks (indicated by white arrows) exposed on a cemented foreshore deposit surface at great brak river following the failure of a superficial graffiti layer – vestigial areas of graffiti are evident on the overlying layer. the outcome was less fortunate at brenton-on-sea, an area that now also boasts at least one hominin tracksite (helm et al. 2018b:1–13). in a small cave, the sloping floor is adorned with multiple examples of graffiti, as a result of which the underlying trackways are deformed beyond recognition and interpretation. tragically, there were indications, noted by our research team, that these may also have been hominin tracks (figure 1b). at another site, in goukamma nature reserve (helm et al. 2020b:1–13), graffiti (figure 1c) has been observed within a few metres of a hominin tracksite. graffiti in the shape of letters and hearts may be readily identifiable as such, but abstract or older examples have the potential to create identification challenges in the interpretation of such sites, which we regard as an unwelcome intrusion. we have observed that tracksites seem to encourage the creative urge in the graffiti artist, as such sites appear to harbour a disproportionate amount of recent etchings, either in direct association with the tracks or on adjacent surfaces. a surface at brenton-on sea that contains well-preserved late pleistocene invertebrate burrow traces also contains deeply etched graffiti (figure 1d). furthermore, a surface on a fallen slab in garden route national park is unusual, in that it not only contains etched graffiti, but also the resulting grooves have been daubed with an erosion-resistant black substance, making the graffiti visible from a considerable distance (figure 1e). unfortunately, the graffiti transects the only example we have observed thus far of a late pleistocene lagomorph (hare) trackway. another area that we have examined in the goukamma nature reserve (helm et al. 2018b:89–101), which comprises a dense concentration of fossil tracksites, including the longest long-horned buffalo trackway thus far identified, also contains its share of graffiti. areas with a high public recreational presence not surprisingly also exhibit a larger amount of graffiti. however, an example from the rocks near the mouth of the great brak river illustrates a rare positive effect of graffiti: a large fallen block, from a layer dated to ~121 000 years (cawthra et al. 2018:156–172), contained a substantial flat surface, which attracted graffiti artists, and this came to be known as ‘graffiti rock’. then around 2010, the surface layer failed and much of it broke off, probably as a result of the pounding by hammers and chisels. this 4-cm-thick layer tumbled into the river, leaving just a small corner of graffiti as a reminder, and exposing a number of bovid trackways on the underlying surface, which were identified by our team (figure 1f). these trackways provided the first unequivocal evidence of pleistocene vertebrates leaving their tracks on beaches in southern africa (helm et al. 2021:59–74). however, this result is an exception to a trend of increasing destruction and vandalism. there are sites in the western cape, such as the heerenlogement (near graafwater: https://sahris.sahra.org.za/node/15040) and the herrieklip (in meiringspoort: https://sahris.sahra.org.za/sites/920680010), where examples of lettering that was carved into rock surfaces, as much as three centuries ago, have been declared provincial historic sites. while such sites may be of historical importance, we are not aware of such significance for any of the graffiti found on aeolianite surfaces. it appears that the erodible nature of these surfaces prevents graffiti (and tracksites) from enduring for more than a few generations. we also noted that while some forms of graffiti can be erased (e.g., paint on walls in public spaces), defacement of aeolianite surfaces is essentially irreversible. we noted a regrettable irony with regard to the hominin tracksites. the surfaces that recorded the footprints and activities of our own species became buried and, over time, became cemented, and then became re-exposed through forces of erosion. it is incredibly fortuitous that such re-exposed surfaces occur in southern africa, where our distant ancestors ‘found their feet’, and that they are available to us for interpretation. yet, some modern-day descendants of those ancestral pioneers carve new forms into those surfaces, sometimes before any scientific appraisal can occur. what can be performed to stop this phenomenon? while regulations and punitive fines may have their place, prevention through education and awareness may be more suitable. examples of graffiti on trees and in association with rock art can be used to inform such discussion. for example, in cases involving etching into ancient trees, signage can alert visitors to the dangers of such behaviour, and can direct would-be graffiti artists to nearby alternative stumps on which they can ply their craft. over time, the importance of not defacing rock art sites has been appreciated, although the only certain means of protection in some cases has been to construct physical barriers and gates, despite the associated aesthetic drawbacks, for example, at schaapplaats in free state province (ouzman 2000:8–10). unfortunately, access to areas of the aeolianite-containing coastline is harder to prevent. signage at access points to areas of coastline could include catch phrases such as ‘please don’t destroy in five minutes what took 120 000 years to form’. while we are uncertain as to their potential effectiveness, display boards explaining what aeolianites are and emphasising their geological and ichnological importance could be erected at such access points – much like signs that provide information on archaeological sites and coastal fauna and flora. because many aeolianites occur in south africa’s national parks and other protected areas, we suggest that authorities such as sanparks and capenature are well placed to lead such initiatives, ideally in collaboration with municipalities that have jurisdiction over unprotected areas. regardless of how this issue is eventually approached, we believe that the initial step consists simply of promoting an awareness of the importance of protecting aeolianites amongst preservation-minded people. aeolianites are remarkable rock types, and their surfaces may be worthy of protection in their own right. however, we make the case that their heritage and scientific value strengthen the argument for their protection. it is our view that this value is often underappreciated, and the scientific importance of an aeolianite surface may not readily be apparent to a non-ichnologist. furthermore, whilst aeolianites worldwide are worthy of protection, the information that can be gleaned from them regarding hominin origins places the south african examples in a unique context and makes them of critical international importance. acknowledgements the support of sanparks and cape nature in providing permission for the authors to study pleistocene palaeosurfaces in national parks and nature reserves is appreciated. the authors wish to acknowledge the two anonymous peer reviewers who provided insightful contributions, and the input of the editor. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this letter. authors’ contributions c.w.h., h.c.c., r.m.c., j.c.d.v., m.g.l., c.w.m., m.g.d., c.t.z.h., w.s., g.h.h.y. and j.a.v contributed to and agreed upon the contents of the scientific letter. ethical consideration this article followed all ethical standards for a research without direct contact with human or animal subjects. funding information the authors received no financial support for the research, authorship and/or publication of this letter. data availability the authors confirm that the data supporting the findings of this study are available within the letter. disclaimer the views expressed in the submitted letter are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the author. references bateman, m.d., carr, a.s., dunajko, a.c., holmes, p.j., roberts, d.l., mclaren, s.j. et al., 2011, ‘the evolution of coastal barrier systems: a case study of the middle-late pleistocene wilderness barriers, south africa’, quaternary science reviews 30(1–2), 63–81. https://doi.org/10.1016/j.quascirev.2010.10.003 brown, k.s., marean, c.w., herries, a.i.r., jacobs, z., tribolo, c., braun, d. et al., 2009, ‘fire as an engineering tool of early modern humans’, science 325(5982), 859–862. https://doi.org/10.1126/science.1175028 brown, k.s., marean, c.w., jacobs, z., schoville, b.j., oestmo, s., fisher, e.c. et al., 2012, ‘an early and enduring advanced technology originating 71 000 years ago in south africa’, nature 491(1), 590–593. https://doi.org/10.1038/nature11660 carr, a.s., bateman, m.d., cawthra, h.c. & sealy, j., 2019, ‘first evidence for on-shore marine isotope stage 3 aeolianite formation on the southern cape coastline of south africa’, marine geology 407(1), 1–15. https://doi.org/10.1016/j.margeo.2018.10.003 cawthra, h.c., jacobs, z., compton, j.s., fisher, e.c., karkanas, p. & marean, c.w., 2018, ‘depositional and sea-level history from mis 6 (termination ii) to mis 3 on the southern continental shelf of south africa’, quaternary science reviews 181(1), 156–172. https://doi.org/10.1016/j.quascirev.2017.12.002 d’errico, f., henshilwood, c., vanhaeren, m. & van niekerk, k., 2005, ‘nassarius kraussianus shell beads from blombos cave: evidence for symbolic behaviour in the middle stone age’, journal of human evolution 48(1), 3–24. https://doi.org/10.1016/j.jhevol.2004.09.002 helm, c.w., cawthra, h.c., cowling, r.m., de vynck, j.c., lockley, m.g., marean, c.w. et al., 2020a, ‘pleistocene vertebrate tracksites on the cape south coast of south africa and their potential palaeoecological implications’, quaternary science reviews 235(1), article id 105857. https://doi.org/10.1016/j.quascirev.2019.07.039 helm, c.w., cawthra, h.c., de vynck, j.c., dixon, m. & stear, w., 2021, ‘elephant tracks – a biogenic cause of potholes in pleistocene south african coastal rocks’, journal of coastal research 37(1), 59–74. https://doi.org/10.2112/jcoastres-d-20-00064.1 helm, c.w., lockley, m.g., cawthra, h.c., de vynck, j.c., dixon, m.g., helm, c.j.z. et al., 2020b, ‘newly identified hominin trackways from the cape south coast of south africa’, south african journal of science 116(9/10), art. #8156, 13 pages. https://doi.org/10.17159/sajs.2020/8156 helm, c.w., mccrea, r.t., cawthra, h.c., cowling, r.m., lockley, m.g., marean, c.w. et al., 2018a, ‘a new pleistocene hominin tracksite from the cape south coast, south africa’, scientific reports 8(1), 3772. https://doi.org/10.1038/s41598-018-22059-5 helm, c.w., mccrea, r.t., cawthra, h.c., thesen, g.h.h. & mwankunda, j.m., 2018b, ‘late pleistocene trace fossils in the goukamma nature reserve, cape south coast, south africa’, palaeontologia africana 52(1), 89–101. henshilwood, c., d’errico, f., marean, c., milo, r. & yates, r., 2001, ‘an early bone tool industry from the middle stone age at blombos cave, south africa: implications for the origins of modern human behaviour, symbolism and language’, journal of human evolution 41(6), 632–678. https://doi.org/10.1006/jhev.2001.0515 henshilwood, c.s., d’errico, f., van niekerk, k.l., dayet, l., queffelec, a. & pollarolo, l., 2018, ‘an abstract drawing from the 73,000-year-old levels at blombos cave, south africa’, nature 562(1), 115–118. https://doi.org/10.1038/s41586-018-0514-3 henshilwood, c.s., d’errico, f., yates, r., jacobs, z., tribolo, c., duller, g.a.t. et al., 2002, ‘emergence of modern human behavior: middle stone age engravings from south africa’, science 295(5588), 1278–1280. https://doi.org/10.1126/science.1067575 henshilwood, c.s., van niekerk, k.l., wurz, s., delagnes, a., armitage, s.j., rifkin, r.f. et al., 2014, ‘klipdrift shelter, southern cape, south africa: preliminary report on the howiesons poort layers’, journal of archaeological science 45(1), 284–303. https://doi.org/10.1016/j.jas.2014.01.033 lockley, m.g., cawthra, h.c., de vynck, j.c., helm, c.w., mccrea, r.t. & nel, r., 2019, ‘new fossil sea turtle trackway morphotypes from the pleistocene of south africa highlight role of ichnology in turtle palaeobiology’, quaternary research 92(3), 626–640. https://doi.org/10.1017/qua.2019.40 marean, c.w., 2010, ‘pinnacle point cave 13b (western cape province, south africa) in context: the cape floral kingdom, shellfish, and modern human origins’, journal of human evolution 59(3–4), 425–443. https://doi.org/10.1016/j.jhevol.2010.07.011 marean, c.w., bar-matthews, m., bernatchez, j., fisher, e., goldberg, p., herries, a.i.r. et al., 2007, ‘early human use of marine resources and pigment in south africa during the middle pleistocene’, nature 449(7164), 905–908. https://doi.org/10.1038/nature06204 marean, c.w., cowling, r.c. & franklin, j., 2020, ‘the palaeo-agulhas plain: temporal and spatial variation in an extraordinary extinct ecosystem of the pleistocene of the cape floristic region’, quaternary science reviews 235(1), article id 106161. https://doi.org/10.1016/j.quascirev.2019.106161 ouzman, s., 2000, ‘public rock art sites of the free state: schaapplaats’, culna 55(1), 8–10. pickering, r., jacobs, z., herries, a.i.r., karkanas, p., bar-matthews, m., woodhead, j.d. et al., 2013, ‘paleoanthropologically significant south african sea caves dated to 1.1–1.0 million years using a combination of u–pb, tt-osl and palaeomagnetism’, quaternary science reviews 6(1), 39–52. https://doi.org/10.1016/j.quascirev.2012.12.016 roberts, d.l., 2008, ‘last interglacial hominid and associated vertebrate fossil trackways in coastal eolianites, south africa’, ichnos 15(3), 190–207. https://doi.org/10.1080/10420940802470482 roberts, d.l., bateman, m.d., murray-wallace, c.v., carr, a.s. & holmes, p.j., 2008, ‘last interglacial fossil elephant trackways dated by osl/aar in coastal aeolianites, still bay, south africa’, palaeogeography, palaeoclimatology, palaeoecology 257(3), 261–279. https://doi.org/10.1016/j.palaeo.2007.08.005 roberts, d.l., karkanas, p., jacobs, z., marean, c.w. & roberts, r.g., 2012, ‘melting ice sheets 400,000 yr ago raised sea level by 13 m: past analogue for future trends’, earth and planetary science letters 357–358(1), 226–237. https://doi.org/10.1016/j.epsl.2012.09.006 wadley, l., 2015, ‘those marvellous millennia: the middle stone age of southernmost africa’, azania 50(2), 155–226. https://doi.org/10.1080/0067270x.2015.1039236 abstract introduction study area methods results plant community descriptions discussion conclusion acknowledgements references about the author(s) nqobile s. zungu department of botany, university of zululand, south africa theo h.c. mostert department of botany, university of zululand, south africa rachel e. mostert department of biology, felixton college, south africa citation zungu, n.s., mostert, t.h.c. & mostert, r.e., 2018, ‘plant communities of the umlalazi nature reserve and their contribution to conservation in kwazulu-natal’, koedoe 60(1), a1449. https://doi.org/10.4102/koedoe.v60i1.1449 note: additional supporting information may be found in the online version of this article as online appendix 1: https://doi.org/10.4102/koedoe.v60i1.1449-1 and online appendix 2: https://doi.org/10.4102/koedoe.v60i1.1449-2 original research plant communities of the umlalazi nature reserve and their contribution to conservation in kwazulu-natal nqobile s. zungu, theo h.c. mostert, rachel e. mostert received: 18 nov. 2017; accepted: 06 feb. 2018; published: 28 may 2018 copyright: © 2018. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract vegetation research is an important tool for the simplified and effective identification, management and conservation of the very complex ecosystems underlying them. plant community descriptions offer scientists a summary and surrogate of all the biotic and abiotic factors shaping and driving ecosystems. the aim of this study was to identify, describe and map the plant communities within the umlalazi nature reserve. a total of 149 vegetation plots were sampled using the braun-blanquet technique. thirteen plant communities were identified using a combination of numeric classification (modified two-way-indicator species analysis) and ordination (non-metric multidimensional scaling). these communities were described in terms of their structure, floristic composition and distribution. an indirect gradient analysis of the ordination results was conducted to investigate the relationship between plant communities and their potentially important underlying environmental drivers. based on the results, the floristic conservation importance of each plant community was discussed to provide some means to evaluate the relative contribution of the reserve to regional ecosystem conservation targets. conservation implications: the umlalazi nature reserve represents numerous ecosystems that are disappearing from a rapidly transforming landscape outside of formally protected areas in zululand. the descriptions of the plant communities of these relatively pristine ecosystems provide conservation authorities with inventories and benchmarks with which the ecological health of similar ecosystems in the region can be measured. introduction vegetation research is an important tool for the simplified and effective identification, management and conservation of the very complex underlying ecosystems. plant community descriptions offer scientists a summary and surrogate for the bewildering combinations of biotic and abiotic factors shaping and driving ecosystems. in response to a recent request by ezemvelo kwazulu-natal wildlife (ekznw), this study of the plant communities of the umlalazi nature reserve (umnr) was conducted as a first attempt to provide such a vegetation description. in order to establish the uniqueness and conservation importance of the umnr’s plant communities, comparisons were made with existing vegetation descriptions from the literature. without scientifically based ecological information and interpretation of vegetation data, conservation authorities will not be able to either report meaningfully on the state of the vegetation under their management, or set and adapt conservation priorities as required by the national environmental management biodiversity act (act 10 of 2004) (von maltitz, van wyk & everard 1996:188–195). previous vegetation studies in the umnr were conducted at a relatively coarse scale, aimed at providing very general trends within the major vegetation types (nevill & nevill 1995:51–58; todd 1994; weisser 1978a:95–97). vegetation maps that were compiled in the past were based on dominant species and vegetation structure classes (todd 1994; weisser 1978b). to date, no detailed plant community descriptions based on total floristic composition have been conducted for the umnr. the only detailed description of these ecosystems comes from the neighbouring pennington park by moll (1972:615–626). detailed vegetation descriptions for other ecosystems within the indian ocean coastal belt (iocb) biome have been published for tshanini game reserve (gaugris et al. 2004:9–29), tembe elephant park (matthews et al. 2001:573–594), lake eteza nature reserve (neumann, scott, bousman & van as 2010:39–53) and sileza nature reserve (matthews et al. 1999:151–167). other detailed plant ecology studies have focussed on foredune advancement, dune vegetation changes and changes in grassland communities (weisser 1978a:95–97; weisser, garland & drews 1982:127–130; weisser & muller 1983:661–667). study area the umnr (28°56′s, 31°46′e) is situated directly south of mtunzini, within the northern half of kwazulu-natal, in the southern section of zululand, south africa (todd 1994; traynor et al. 2010). this coastal reserve (1469 ha) forms part of the maputaland-pondoland-albany biodiversity hotspot and the maputaland centre of floristic endemism (van wyk & smith 2001). the centre contains approximately 2500 infraspecific taxa, of which 9.2% are regarded as near-endemics. the climate of the study area is subtropical with hot, humid summers and mild winters with no frost (nevill & nevill 1995:51–58; todd 1994). the mean annual rainfall fluctuates between 819 mm and 1272 mm, with approximately two-thirds falling in mid-summer and the remaining one-third falling in early winter (mucina & rutherford 2006; tyson & preston-whyte 2000). the ages of the dune fields of the maputaland coastal plain range from the early pleistocene (± 5 ma) to 500 years ago. the dunes of the maputaland coastal plain are among the most recent geological formations that are found in southern africa (gaugris et al. 2004:9–29), with the umnr beachfront classified as prograding (actively expanding). the soil of the umnr consists of fine-grained marine sands, typically yellowish or grey apedal soil with early horizon development. the a-horizon is usually thin and enriched with organic matter. subsoil horizons often show sparse ferruginous mottles. the resulting soil types are classified according to the south african system as mainly cover sands, with some fernwood soil forms and some very limited champagne soil formations along the waterlogged bottomlands (fey 2010:32–35; matthews et al. 2001:573–594). this combination of a water-rich environment and highly permeable sandy substrates is the main driver of the vegetation patterns of this biome. the region forms part of the iocb biome, which is a complex mosaic of zonal, intrazonal and azonal vegetation types (mucina & rutherford 2006). the umnr and its immediate surroundings contain the following major vegetation types: foz 7 northern coastal forest, foa 2 swamp forest, foa 3 mangrove forest, aze subtropical estuarine salt marshes, azs 3 subtropical dune thicket, azd 4 subtropical seashore vegetation and azf 6 subtropical freshwater wetlands (mucina & rutherford 2006). cultivation, afforestation, mining, urban sprawl and invasive alien plants are major threats to the remaining untransformed patches of iocb vegetation (ekznw 2009). methods the study area was stratified into homogenous vegetation units using texture and colour classes on aerial imagery (google earth 7.1.2.2041 2016). after a reconnaissance of the area to ensure the homogeneity of vegetation units, 149 plots were randomly selected within each of the different homogenous units on the imagery. in the field, however, each sampling plot was critically evaluated according to the first rule of the zurich–montpellier sampling method (the placement of the sampling plot should be within a homogeneous vegetation patch that is representative of the perceived plant community). if the sampling plot did not fall within a homogeneous representative vegetation stand, it was moved to the nearest locality that fulfils this criterion. accessibility to the plot was also taken into consideration. the braun-blanquet sampling method (werger & coetzee 1978) was specifically chosen for its international recognition as the most appropriate technique for the description of vegetation when based on total floristic composition (brown et al. 2013). by using an internationally accepted standard, the data and vegetation description will be compatible and comparable with data from other regions and landscapes. such comparability and compatibility is crucial for the regional and international coordination of vegetation and ecosystem conservation. based on the recommendations by brown et al. (2013), plot sizes varied according to the vegetation types sampled (grasslands 9 m2, salt marshes 25 m2, wetland vegetation 25 m2, foredune communities 49 m2, swamp communities 49 m2 and forest communities 1000 m2) and were marked out in the field to ensure consistency. in all sample plots, each plant species was recorded and the cover–abundance value of each species determined using the modified braun-blanquet cover–abundance scale: r (very rare, with a negligible cover), + (present but not abundant, with a cover value of < 1% of the quadrat), 1 (numerous but covering less than 1% of the sample area), 2a (covering 5% – 12% of the sample area), 2b (covering 13% – 25% of the sample area), 3 (covering 25% – 50% of the sample area), 4 (covering 50% – 75% of the sample area), 5 (covering 75% – 100% of the sample area) (brown et al. 2013). the names of the plant species follow the latest taxonomy, as provided by the south african biodiversity institute online from the plants of south africa (posa), which was last accessed at http://www.ville-ge.ch/musinfo/bd/cjb/africa/ on 15 june 2017. fieldwork was conducted during the peak growing season (october 2014 – february 2015). the vegetation structure was recorded and described based on the height and canopy cover of the different strata (tall trees, short trees, shrubs, lianas and herbaceous strata) within each plant community and classified in accordance with edwards’ structural classification (edwards 1983:705–712). the phytosociological data were captured in turboveg (hennekens & schaminee 2001:589–591), where relevés were created and exported as a cornell condensed format file into juice 7.0.84 (tichy 2002) for analysis. the modified two-way-indicator species analysis (twinspan) algorithm as contained within juice (tichy 2002) was used to classify the floristic data. the classification of different plant communities was based on total floristic composition, with pseudo-species cut levels set at 0, 1, 5, 12, 25, 50 and 75. before the classification was performed, all relevé cluster sizes were standardised in order to remove phi coefficient dependence on cluster size. the fidelity of species to the various plant communities was calculated using the phi coefficient. fisher’s exact test was used to calculate the statistical significance (p > 0.001) of the fidelity values calculated for each species. the diagnostic values of species were based on their fidelity to each community, with phi coefficient threshold values set at > 33% (chytrý et al. 2002:79–90), while dominant species were identified based on their relative cover–abundance within each of the resulting plant communities. an ordination of the floristic data was performed using a non-metric multidimensional scaling (nmds) within pc-ord (mccune & mefford 1999) in order to visualise dissimilarities between samples in a two-dimensional scatter plot. environmental data gathered during the vegetation surveys were mostly qualitative descriptions of the physical parameters associated with the various sample sites. at each sample site, attention was focussed on those environmental variables that were deemed more influential with regard to driving vegetation composition and structure. a conscious decision was therefore made to gather detailed environmental data specific and relevant to each sample site, and not a standardised matrix of generalised environmental parameter data. for this reason, it was decided that an indirect gradient analysis would be attempted, based on the detailed sample site descriptions, instead of a direct gradient analysis based on generalised and often irrelevant environmental parameters. all the observations were made based on field testing, with no laboratory testing conducted. the following scales were used to ensure robust and reliable categories for the environmental variables recorded: percentage of soil clay content (sausage texture method): < 10, 10–14, 15–19, 20–34, 35–55, > 55 water drainage from the soil column based on soil texture: fast, medium, slow, stagnant water drainage from the landscape based on runoff potential: fast, medium, slow, stagnant soil moisture: saturated, moist, dry soil salt content (electrical conductivity): high (oceanic saltwater contamination), low (no oceanic saltwater contamination) organic content of soil (visual recognition of fibre particles): high (> 10% of horizon volume), some (< 10%), none anthropogenic disturbance levels: high, moderate, low, none wind exposure: high, medium, low sunlight penetration to lower vegetation strata: full shade (> 80%), partial shade (40% – 79%), highly exposed (< 40%). plant community names were assigned following the guidelines suggested by brown et al. (2013), namely, a diagnostic species followed by a dominant species followed by a structural or landscape description. the mapping of the plant communities was performed using google earth 7.1.2.2041 (unprojected) and the final output was compiled in quantum gis 2.14.1. ethical considerations a plant collection permit (op 4829/2014) for identification and herbarium preparation was obtained from ezemvelo kwazulu-natal wildlife (ekznw) for use in the umlalazi nature reserve. results classification the results from the modified twinspan classification and the nmds ordination are presented in a dendrogram (figure 1), an ordination scatter plot (figure 2), a synoptic table (online appendix 1) and a full phytosociological table (online appendix 2). the environmental parameters associated with the segregation of the various relevé clusters are presented as part of the classification dendrogram (figure 1). the results from the indirect gradient analysis superimposed on the nmds ordination scatter plot are presented in figure 2. based on the combined results from the modified twinspan classification and the nmds ordination of the umnr vegetation, 13 different plant communities were identified: salicornia meyeriana–avicennia marina salt marsh community bruguiera gymnorrhiza–avicennia marina mangrove forest phragmites australis–juncus kraussii saline wetland scaevola plumieri–gazania rigens foredune community typha capensis–cyperus dives wetland community digitaria eriantha–dactyloctenium australe secondary coastal grasslands stenotaphrum secundatum–phragmites australis temporary wetlands passerina rigida–carpobrotus dimidiatus dune scrub community adenopodia spicata–vachellia robusta riverine woodland community albizia adianthifolia–trichilia emetica disturbed coastal dune forest tricalysia sonderiana–apodytes dimidiata dune forest margin gymnosporia arenicola–protorhus longifolia coastal dune forest carissa bispinosa–mimusops caffra climax coastal dune forest. figure 1: dendrogram illustrating the modified twinspan classification of the vegetation of the umlalazi nature reserve into 13 plant communities. figure 2: plant community clusters of the umlalazi nature reserve as produced by a non-metric multidimensional scaling ordination. representative photographs of the various plant communities are presented in figure 3. figure 3: representative photographs of the umlalazi nature reserve plant communities: (a) salicornia meyeriana–avicennia marina salt marsh community, (b) bruguiera gymnorrhiza–avicennia marina mangrove forest, (c) phragmites australis–juncus kraussii saline wetland, (d) scaevola plumieri–gazania rigens foredune community, (e) typha capensis–cyperus dives wetland community, (f) digitaria eriantha–dactyloctenium australe secondary coastal grasslands, (g) stenotaphrum secundatum–phragmities australis freshwater wetland, (h) passerina rigida–carpobrotus dimidiatus dune scrub community, (i) adenopodia spicata–vachellia robusta riverine woodland community, (j) albizia adianthifolia–chromolaena odorata disturbed coastal dune forest, (k) tricalysia sonderiana–apodytes dimidiata dune forest margin, (l) gymnosporia arenicola–protorhus longifolia coastal dune forest, (m and n) carissa bispinosa–mimusops caffra climax coastal dune forest. figure 3 (continues ...): representative photographs of the umlalazi nature reserve plant communities: (a) salicornia meyeriana–avicennia marina salt marsh community, (b) bruguiera gymnorrhiza–avicennia marina mangrove forest, (c) phragmites australis–juncus kraussii saline wetland, (d) scaevola plumieri–gazania rigens foredune community, (e) typha capensis–cyperus dives wetland community, (f) digitaria eriantha–dactyloctenium australe secondary coastal grasslands, (g) stenotaphrum secundatum–phragmities australis freshwater wetland, (h) passerina rigida–carpobrotus dimidiatus dune scrub community, (i) adenopodia spicata–vachellia robusta riverine woodland community, (j) albizia adianthifolia–chromolaena odorata disturbed coastal dune forest, (k) tricalysia sonderiana–apodytes dimidiata dune forest margin, (l) gymnosporia arenicola–protorhus longifolia coastal dune forest, (m and n) carissa bispinosa–mimusops caffra climax coastal dune forest. figure 3 (continues ...): representative photographs of the umlalazi nature reserve plant communities: (a) salicornia meyeriana–avicennia marina salt marsh community, (b) bruguiera gymnorrhiza–avicennia marina mangrove forest, (c) phragmites australis–juncus kraussii saline wetland, (d) scaevola plumieri–gazania rigens foredune community, (e) typha capensis–cyperus dives wetland community, (f) digitaria eriantha–dactyloctenium australe secondary coastal grasslands, (g) stenotaphrum secundatum–phragmities australis freshwater wetland, (h) passerina rigida–carpobrotus dimidiatus dune scrub community, (i) adenopodia spicata–vachellia robusta riverine woodland community, (j) albizia adianthifolia–chromolaena odorata disturbed coastal dune forest, (k) tricalysia sonderiana–apodytes dimidiata dune forest margin, (l) gymnosporia arenicola–protorhus longifolia coastal dune forest, (m and n) carissa bispinosa–mimusops caffra climax coastal dune forest. ordination the nmds ordination (149 relevés) resulted in a scatter diagram with nine distinct clusters when viewed as a two-dimensional plot along axes 1 and 2 (figure 2). the indirect gradient analysis superimposed onto the scatter diagram revealed the following potential environmental gradients between plant communities, based on the relative distances and sequences among clusters: soil moisture availability, water drainage capability of soils within different parts of the landscape, clay and salt content of soils, effects of fire and grazing, effects of organic content in the upper soil profile and effects of salt-clipping by oceanic salt spray driven by onshore winds. species richness the mean species richness per relevé within plant communities is indicated in figure 4. plant communities 1–5 show very low levels of species richness and are associated with wetlands and newly formed dunes. these communities can generally be regarded as very harsh to plant life, with only a few specialist species adapted to survive in them. these environmental conditions include soils that are flooded for prolonged periods of time, tidal folding regimes, hyper-saline soils, mobile structureless soils and salt-laden oceanic wind. plant communities 6, 8, 9, 10, 12 and 13 show relatively high species richness. these plant communities are associated with the more stable soils and more mesic conditions of the sheltered dune forests and grasslands. communities 9, 10, 12 and 13 revealed the highest species richness in the umnr. figure 4: species richness for the 13 plant communities of the umlalazi nature reserve. plant community descriptions plant community 1: salicornia meyeriana–avicennia marina salt marsh community the salicornia meyeriana–avicennia marina salt marsh community is located within certain estuarine sections of the umlalazi river floodplains (figure 5). these sections of the floodplains do not drain freely after the regular seasonal flooding events, and act as natural evaporation pans. the combination of saltwater exposure from tidal movements and the evaporation of water from the temporary pans lead to the hyper-accumulation of salt in the soil of this plant community. these alluvial soils generally contain large percentages of silt, with very little organic matter. the community has a very low vegetation cover and a simple vegetation structure. its structure varies from open (10%) to patches of closed (75%), low herbaceous vegetation, mostly clumped into colonies of s. meyeriana. figure 5: vegetation map of the umlalazi nature reserve. the diagnostic species for this plant community, the succulent species s. meyeriana, is listed in species group (sg a) of the synoptic table (online appendix 1). the dominant species for this community are displayed in the full phytosociological table (online appendix 2) and include the low-growing halophytic succulent s. meyeriana (sg a) and the tree species a. marina (sg b). however, a. marina has a low cover–abundance value within this plant community. based on the vegetation structure and composition, this plant community is regarded as a part of the major vegetation type aze 3 subtropical estuarine salt marshes, as described by mucina and rutherford (2006). although some authors have described similar subtropical salt marsh vegetation (colloty, adams & bate 2002; lubke 1997; venter 1972), no equivalent to that of the umnr was observed in the literature. despite the very low species richness recorded for this plant community (2 ± 0.5 s.d.), its conservation value is considered to be high, based on its unique floristic composition and the unique and dynamic nature of this ecosystem. it acts as a refuge for benthic organisms during low tides and as rich feeding grounds for many vertebrate species during unusually high tides (bromberg-gedan, silliman & bertness 2009). plant community 2: bruguiera gymnorrhiza–avicennia marina mangrove forest the bruguiera gymnorrhiza–avicennia marina mangrove forest community is restricted to some of the very narrow intertidal zones of the umlalazi river in the study area (figure 5). the soils of this community are poorly drained, saline, anoxic and fine-grained. these siltand clay-rich soils are mostly waterlogged. this community is typically species-poor, dominated by one or two species, and with a dense vegetation cover. structurally, this community varies from medium to tall, closed mangrove forest (5 m – 8 m). the diagnostic tree species for this community are b. gymnorrhiza and a. marina (sg b, online appendix 1). the most dominant tree species in this community are presented in the full phytosociological table (sg b, online appendix 2) and include the tree a. marina, with only a few other visually prominent species such as the perennial sedge juncus kraussii. based on the vegetation structure and composition, this plant community is regarded as a part of the major vegetation type foa 3 mangrove forest, as described by mucina and rutherford (2006). the mangrove community of the umnr is similar to those described by moll and werger (1978), steinke (1995) and colloty et al. (2002). the very extreme and constantly changing environmental conditions of these intertidal ecosystems make them very challenging for terrestrial plant life. only those plant species that are specifically adapted to deal with both flooded saline conditions, when the tide is in; and extreme desiccation, when the tide is out, are able to colonise these ecosystems. without competition from less-adapted plant species, such plant communities become dominated by a handful of specialists, such as mangrove tree species. the conservation value of such plant communities therefore does not lie in its plant diversity, but in its ecosystem functionality. plant community 3: phragmites australis–juncus kraussii saline wetland the phragmites australis–juncus kraussii saline wetland plant community occurs within the floodplains of the umlalazi river in places where saltwater contamination occurs during super-tidal events. it is periodically flushed by freshwater flooding events from the river (figure 5). within the floodplains, this community forms a mosaic distribution pattern with the salicornia meyeriana–avicennia marina salt marsh community. soils are typically saline, with high silt, clay and organic components. water drainage is slow and even stagnant in some cases, leading to the accumulation of organic material in these anaerobic conditions. the vegetation is structurally characterised by a medium to tall, closed reed and sedgeland (1.5 m – 3.5 m). the vegetation of this very dynamic ecosystem is regularly harvested of the sedge species j. kraussii and the reed species p. australis for the weaving and building industry. the only diagnostic species for this community is the sedge species j. kraussii (sg c, online appendix 1). the dominant species (online appendix 2) include the reed species p. australis (sg h), the sedge species j. kraussii (sg c), the forb species ipomoea cairica (sg d) and the grass species stenotaphrum secundatum (sg g). this plant community, with its mosaic of reed and sedge-dominant patches, is the result of fluctuating salinity at both spatial and temporal scales. wherever and whenever salinity rises, the reed species are negatively affected, and the more salt-tolerant j. kraussii outcompetes the reeds. when salt is flushed from the system, the reeds gain the competitive advantage and outcompete j. kraussii. the relatively high cover–abundance values recorded for the grass species s. secundatum are regarded as an artefact of the drought conditions that prevailed at the time when surveys were conducted. under more normal flood conditions, this grass species would be drowned, only occupying the better drained fringes of these wetlands. based on the vegetation structure and composition, this plant community is regarded as a part of the major vegetation type aze subtropical estuarine salt marshes, as described by mucina and rutherford (2006). juncus kraussii-dominated saline sedgelands occur widely, spread along the estuaries of the east coast of south africa (colloty et al. 2002; taylor, adams & haldorsen 2006). just like in the umnr, these communities are dynamic in their response to fluctuations in salinity. plant community 4: scaevola plumieri–gazania rigens foredune community the scaevola plumieri–gazania rigens foredune plant community occupies the seaward side of the first dune above the high-water mark along the beach (figure 5). at this point in the landscape the sand is still very mobile and contains virtually no organic matter as yet. water drainage is fast and salinity is relatively high because of oceanic salt spray. the vegetation structure is relatively simple, with mainly one layer of herbaceous plants with an average height of 0.4 m. as a result, the vegetation structure for this community can be described as patches of low, closed herblands, with large stretches of open, uncolonised mobile sand between the vegetation patches. the diagnostic species for this plant community include the succulent shrublet s. plumieri (sg d online appendix 1), the perennial trailing herb ipomoea pes-caprae (sg d) and the creeping perennial herb g. rigens (sg d). the most dominant species in this community (online appendix 2) is the creeping perennial herb g. rigens (sg d). other visually prominent species include the forbs i. pes-caprae (sg d) and s. plumieri (sg d), the succulent forb carpobrotus dimidiatus (sg i) and the shrubs rhynchosia nitens (sg i) and chrysanthemoides monilifera (sg i). although the recorded species richness is relatively low, this plant community plays a very important ecological role in stabilising the prograding (expanding) beaches of zululand. this can be seen as the first in a long series of successional stages from dune colonisation to the final climax state of mature coastal dune forest. based on the vegetation structure and composition, this plant community is regarded as a part of the major vegetation type azd 4 subtropical seashore vegetation, as described by mucina and rutherford (2006). plant communities and vegetation types similar to the one described for the umnr occur along large sections of the zululand coast and have been described by lubke et al. (1997). plant community 5: typha capensis–cyperus dives wetland community the typha capensis–cyperus dives wetland community occurs in interdune depressions where groundwater breaks the surface to form perennial wetlands of various sizes (figure 5). the sandy soils are waterlogged and contain high percentages of clay particles (30%) and very high percentages of organic matter (> 10%). no distinction was made between groundwater and surface water sources within these highly permeable sandy substrates. the structure of this community can be described as a tall, closed reedland (2 m – 4 m), with a cover value of > 85% at the height of the growing season. the diagnostic species for this plant community (sg e, online appendix 1) include the perennial sedge c. dives, the forb persicaria serrulata, the bulrush t. capensis, the fern cyclosorus interruptus and the palm phoenix reclinata. the aforementioned species were, incidentally, also recorded as the most dominant in terms of cover–abundance values (sg e, online appendix 2). such patterns of low species richness coupled with an equally low evenness of dominant species are very typical of wetland ecosystems where only specially adapted species can compete for the available resources. as with many south african wetlands, the conservation value of the associated plant community does not lie in its floristic diversity or uniqueness, but rather in its ecosystem functionality. within the southern sections of the umnr, this plant community has been disturbed and degraded by illegal grazing practised by neighbouring rural communities. based on the vegetation structure and composition, this plant community is regarded as a part of the major vegetation type azf 6 subtropical freshwater wetlands, as described by mucina and rutherford (2006). very few vegetation descriptions that are floristically similar to the typha capensis–cyperus dives wetland community occurring within the umnr could be found in the literature (venter 1972). despite its relatively common occurrence throughout the coastal dune systems of zululand, it is very common for azonal vegetation types such as this to be overlooked and neglected during conventional mapping exercises. plant community 6: digitaria eriantha–dactyloctenium australe secondary coastal grasslands the digitaria eriantha–dactyloctenium australe secondary coastal grassland community includes one section along the southern section of the umnr and one section of the newly acquired land along the north-eastern border of the umnr (figure 5). the southern section was mainly forest and woodlands before being transformed into grassland by human activities. it is characterised by frequent disturbances in the form of fire and cattle grazing induced by the neighbouring human population along the southern border of the reserve. soils are sandy with low levels of organic content. this fire-suppressed, subclimax plant community has a short, closed vegetation structure. the northern section of secondary grassland is a remnant of grassland transformed by agricultural activities. the diagnostic species for this plant community (sg f, online appendix 1) include the grass species d. eriantha, d. australe, sporobolus africanus, imperata cylindrica and stiburus alopecuroides, the sedges kyllinga alata and cyperus species and the forbs helichrysum ruderale, rhynchosia caribaea, wahlenbergia benghalensis and manulea parviflora. the dominant species (online appendix 2) in this community include the grass species d. australe, s. africanus, i. cylindrica and d. eriantha (sg f). some woody species that occur within this community include the low shrub eugenia capensis (sg n), the straggling shrub searsia nebulosa (sg h) and the dwarf shrublet chironia baccifera (sg h). in its current state as a fire-suppressed, subclimax plant community, the conservation value of the southern secondary grassland, based on floristic composition, is regarded as relatively low. although ekznw strives to conserve as much tropical grasslands as possible, this secondary grassland should not be calculated as part of the proposed target set for grassland conservation. from a plant conservation perspective, it is recommended that this plant community be allowed to return to its climax state of coastal dune forest. the northern grassland, however, with its very different origins, is regarded as potentially very important for conservation, based on its potential recovery towards a tall and dense wet grassland. its conservation value will predominantly lie in creating habitat for fauna species of potential concern. based on the vegetation structure and composition, this northern grassland is regarded as a part of the very diverse major vegetation type cb 1 maputaland coastal belt, as described by mucina and rutherford (2006). although no equivalent for this umnr plant community was found in the literature, mucina and rutherford (2006) specifically mentioned the abundance of these secondary coastal grasslands. however, all of them are floristically and structurally very different because of their varied origins and the varied disturbance regimes that created them. most of them are species-poor and occur as early successional stages. plant community 7: stenotaphrum secundatum–phragmites australis temporary wetlands the stenotaphrum secundatum–phragmites australis temporary wetlands community occurs along depressions within the umlalazi river floodplain where enough surface water accumulates to form temporary wetlands (figure 5). the alluvial soil contains enough clay to form a water-impermeable layer that prevents water from draining away. this ecosystem is driven by surface water from seasonal flooding events and rain water. it also occurs, to a limited extent, along small interdune sections where fluctuating groundwater levels create temporary wetlands. depending on the duration and seasonality of each wetland, varying amounts of organic matter accumulate within them. the vegetation structure can be described as tall, closed reedland (2 m – 4 m), with a cover value of > 85%. however, there is a cyclical alteration of dominance between the grass and the reed component of this wetland community. during high rainfall, soils are waterlogged, promoting reed growth and effectively drowning stoloniferous grass species. during low rainfall, soils tend to be better aerated, promoting the grass component. at the time of the field surveys, zululand experienced severe drought conditions, with the lowest average annual rainfall in recorded history for the umlalazi region (sa weather service 2017). the diagnostic species include the grasses s. secundatum and paspalum dilatatum, herbs such as ipomoea cairica, hibiscus trionum and cissampelos hirta and the sedge cyperus eragrostis (sg g, online appendix 1). the most dominant species (online appendix 2) for this community is the sedge p. australis (sg h). other dominant species include the grass s. secundatum (sg g) and the herbs i. cairica (sg g), h. trionum (sg g) and asystasia gangetica (sg u). based on the vegetation structure and composition, this plant community is regarded as a part of the major vegetation type azf 6 subtropical freshwater wetlands, as described by mucina and rutherford (2006). venter (1972), and begg and carser (1988) described similar plant communities and indicated their widespread occurrence within the zululand region. plant community 8: passerina rigida–carpobrotus dimidiatus dune scrub community the passerina rigida–carpobrotus dimidiatus dune scrub community is found along the second and third dunes from the beach (figure 5). this community is established on young dunes, which have slightly higher organic matter content than beach sand with no vegetation. the vegetation structure can generally be described as low (1 m) coastal scrub and thickets, with an open to closed canopy structure (30% – 90%). the trees and shrubs that grow in this community are dwarfed, with a compact canopy flattened by wind-shearing and salt-clipping on the windward sides of dunes. the more protected leeward slopes of the older dunes, however, are characterised by a significantly taller thicket vegetation (3 m – 4 m). these tall thickets along the leeward sides are regarded as the early stages of forest formation. the diagnostic species (sg i, online appendix 1) with the highest fidelity for this community is the shrub p. rigida, which usually grows up to 1.5 m in height. other diagnostic species in this community include the shrubs carissa bispinosa, c. monilifera, osyris compressa, brachylaena discolor, tephrosia purpurea, dichrostachys cinerea and searsia nebulosa; the grass stipagrostis zeyheri; the sedges kyllinga species and cyperus species; the perennial trailing succulent c. dimidiatus; the forbs senecio species and rhynchosia nitens; the creeper abrus precatorius; and the woody climber rhoicissus digitata. the dominant species (online appendix 2) in this community include c. dimidiatus (sg i), the grass s. zeyheri (sg i), the sedges kyllinga species (sg i) and cyperus species (sg i), the herbs senecio species (sg i) and r. nitens (sg i) and the shrubs eugenia capensis (sg n), c. bispinosa (sg q), c. monilifera (sg i), b. discolor (sg i), t. purpurea (sg i), d. cinerea (sg i), s. nebulosa (sg i) and chironia baccifera (sg p). other visually prominent trees in this community include mimusops caffra (sg q), apodytes dimidiata (sg l), kraussia floribunda (sg r) and allophylus natalensis (sg u). based on the vegetation structure and composition, this plant community is regarded as a part of the major vegetation type azs 3 subtropical dune thicket, as described by mucina and rutherford (2006). weisser (1978b), weisser et al. (1982:127–130) and donnelly and pammenter (1983:705–712) also described this widespread plant community at numerous other zululand locations. plant community 9: adenopodia spicata–vachellia robusta riverine woodland community the adenopodia spicata–vachellia robusta riverine woodland community occurs along the better-draining sections of umlalazi river floodplain (figure 5). the structure comprises a tall, dense tree layer and a well-developed dense shrub layer. it is associated with deep, fine-textured soils of recent alluvial deposits that are subject to frequent seasonal flooding. the soils drain freely and do not stay waterlogged for extended periods of time. deep-rooted trees along the riverbank have year-round access to water from this perennial river. the diagnostic species for this plant community (sg j, online appendix 1) include the shrubs a. spicata, tricalysia lanceolata, hibiscus tiliaceus, canthium inerme, pavetta lanceolata, scutia myrtina, tecoma capensis and clausena anisata. the diagnostic herbs include scadoxus membranaceus, chenopodium ambrosioides, oxalis droseroides, nidorella undulata and scadoxus puniceus. the diagnostic trees include v. robusta and tarenna pavettoides. the most dominant species for this community (online appendix 2) include the tree species v. robusta (sg j); the shrubs a. spicata, h. tiliaceus, c. inerme, p. lanceolata, s. myrtina, t. capensis and c. anisata; the herb asystasia gangetica; and the geophytes s. membranaceus and s. puniceus (sg j). because of the very dynamic nature of flooding events along this section of the umlalazi river, the floodplain is highly heterogeneous at both a special and a temporal level. this prohibits the formation of typical lowveld riverine forest (foa 1) (mucina & rutherford 2006). however, we argue that the adenopodia spicata–vachellia robusta riverine woodland community is essentially an early successional stage within the major regional vegetation type. similar plant communities were described by whateley and porter (1983:745–758) further inland along the nyalazi, hluluwe and umfolozi rivers. plant community 10: albizia adianthifolia–trichilia emetica disturbed coastal dune forest the albizia adianthifolia–trichilia emetica disturbed coastal dune forest community has weakly developed sandy–loamy soils with a high organic content in the upper layers. the average total cover for this community is 80% and bare patches make up 20% – 25%. structurally, this community qualifies as a forest. however, because of varying degrees of disturbance to the different vegetation strata, these forests have lost their structural diversity and complexity. disturbances recorded throughout this community include fire; wind; the presence of charcoal pits, animal traps, footpaths and roads; subtropical storms; the harvesting of timber, firewood and medicinal plants; alien plant invasions; and the slumping of unstable dunes and substrates. footpaths were particularly widespread throughout the study area, leading to soil erosion. the diagnostic species for this community include the trees t. emetica, a. adianthifolia, euclea natalensis, erythrina lysistemon, apodytes dimidiata, clerodendrum glabrum, cussonia zuluensis, trimeria grandifolia, deinbollia oblongifolia and ekebergia capensis; the invasive alien shrubs chromolaena odorata and lantana camara; the native shrubs rhynchosia totta and searsia nebulosa; the forb bidens pilosa; and the grass species digitaria longiflora (sg k, online appendix 1). the most dominant species (online appendix 2) include the forb b. pilosa; the trees t. emetica (sg k), e. lysistemon (sg k), a. dimidiata (sg l), c. glabrum (sg k), c. zuluensis (sg k), e. capensis (sg k), t. grandifolia (sg k) and d. oblongifolia (sg k); and the shrubs c. odorata (sg k), l. camara (sg k), r. totta (sg k), s. nebulosa (sg i) and e. natalensis (sg m). based on the vegetation structure and composition, this plant community is regarded as a middle-to-late successional stage of coastal dune forest within the major vegetation type foz 7 northern coastal forest, as described by mucina and rutherford (2006). very few descriptions were found in the literature of the middle-to-late successional stages of forest communities before they reach their climax states (grainger, van aarde & wassenaar 2011; von maltitz, van wyk & everard 1996:188–195). however, most of the coastal dune forests within zululand that occur outside of formally protected areas are in this anthropogenic subclimax state (berliner 2005). plant community 11: tricalysia sonderiana–apodytes dimidiata dune forest margin the tricalysia sonderiana–apodytes dimidiata dune forest margin community occurs along the forest edges. it is demarcated as a narrow band along the forest (figure 5). it has deep apedal sandy–loamy soils with a high accumulation of organic matter within the upper 150 mm of the soil column. structurally, this community can be described as a medium-to-tall dune forest. the diagnostic species for this plant community (sg l, online appendix 1) include the small tree species t. sonderiana and the tree species a. dimidiata. the most dominant species for this community (online appendix 2) includes the tree a. dimidiata (sg l), which grows best in these well-drained, organic-rich soils of the forest margin. other dominant species include psydrax obovata (sg r), the shrubs euclea natalensis (sg m), kraussia floribunda (sg r) and ekebergia capensis (sg n) and the fern microsorum scolopendrium (sg s). this plant community contains surprisingly few plant species (8.3 ± 0.8 s.d.) when compared to the climax dune forest community (19.2 ± 4.9 s.d.). generally, forest ecotones are relatively species rich, containing species from both the forest and the neighbouring plant community. in retrospect, the anomaly recorded here may very well have been an artefact of incorrect sample site selection. it is recommended that the forest ecotones of the umnr be mapped and managed as an integral part of the dune forests. based on the vegetation structure and composition, this plant community is regarded as a part of the major vegetation type foz 7 northern coastal forest, as described by mucina and rutherford (2006). no formal plant community descriptions of coastal dune forest margins were found in the literature. plant community 12: gymnosporia arenicola–protorhus longifolia coastal dune forest the gymnosporia arenicola–protorhus longifolia coastal dune forest community was recorded between the tricalysia sonderana–apodytes dimidiata dune forest margin and the carissa bispinosa–mimusops caffra climax coastal dune forest; as a result, it displays strong floristic affinities towards these plant communities (figure 5). the soils underlying this community are deep apedal sands, ranging from mediumto course-grained, with a high organic component within the upper 150 mm of the soil column. structurally, this community is classified as a tall dune forest. the forest canopy is less dense than the climax dune forests because of the absence of very old and large trees with wide canopies. this results in more light reaching the forest floor, leading to a relatively well-developed shrub layer. the diagnostic species for this plant community include the tree species p. longifolia and tricalysia capensis, the straggling shrub g. arenicola and the straggling climber smilax anceps (sg o, online appendix 1). the dominant species for this community are displayed in the full phytosociological table (online appendix 2) and include psydrax obovata (sg r) and p. longifolia (sg o). other dominant species include the tree species sideroxylon inerme (sg t) and garcinia gerrardii (sg t); the climbers smilex anceps (sg o) and rhoicissus tomentosa (sg q); the shrubs k. floribunda (sg r), g. arenicola, peddia africana (sg r), putterlickia verrucosa (sg q) and c. baccifera (sg q); the grass panicum coloratum (sg o); and the geophyte dietes species (sg m). based on the lack of older and larger trees normally found in a climax dune forest, this plant community may very well be a younger successional stage of dune forest. some preliminary investigations showed that there may very well be a correlation between fire scars on old aerial photographs and the distribution of the gymnosporia arenicola–protorhus longifolia coastal dune forest community. however, the verification of this pattern has not been attempted in this study. this plant community is therefore regarded as a very late successional stage of dune forest within the major vegetation type foz 7 northern coastal forest, as described by mucina and rutherford (2006). very few descriptions were found in the literature of the very late successional stages of forest communities (venter 1972; von maltitz et al. 1996:188–195). some authors (venter 1972; von maltitz et al. 1996:188–195) view these very late successional stages of forest communities simply as variants of climax states. plant community 13: carissa bispinosa–mimusops caffra climax coastal dune forest the carissa bispinosa–mimusops caffra climax coastal dune forest community is associated with the oldest and more protected dune forests, extending up to the northern edge of the umnr (figure 5). the soil underlying this community is deep sandy soil with a very high organic component within the upper 150 mm. structurally, it can be described as a tall, closed forest. the diagnostic species for this plant community (sg q, online appendix 1) include the tree species with the highest fidelity values: m. caffra, dovyalis longispina, vepris lanceolata and cussonia spicata. the diagnostic shrubs and small trees include c. bispinosa, putterlickia verrucosa, brachylaena discolor and bersama lucens. the diagnostic woody climbers include monanthotaxis caffra, rhoicissus rhomboidea, dalbergia armata, rhoicissus tomentosa and dalbergia obovata. the diagnostic species include the grass oplismenus hirtellus, the geophyte dietes species as well as the orchids cyrtorchis praetermissa, polystachya sandersonii and ansellia africana. the most dominant species for this community (online appendix 2) include the tree species m. caffra and shrub c. bispinosa (sg q). other dominant species include the trees psydrax obovata, b. lucens, kraussia floribunda (sg r), ficus natalensis (sg u), sideroxylon inerme (sg t), psydrax obovata (sg t) and d. longispina (sg q); the shrubs and small trees peddia africana (sg t), b. discolor (sg q), grewia occidentalis (sg u), p. verrucosa (sg q), allophylus natalensis (sg u) and carissa macrocarpa (sg q); the woody climbers r. rhomboidea (sg u), asparagus falcutus (sg u) and r. tomentosa (sg q); the grasses o. hirtellus (sg q) and panicum coloratum (sg u); the geophyte dietes species (sg q); and the ferns microsorum scolopendrium (sg s) and microsorum punctatum (sg t). as was expected, the recorded species richness was relatively high (19.2 ± 4.9 s.d.). this community also contains numerous protected plant species, including m. caffra, s. inerme, c. praetermissa, p. sandersonii, a. africana and the extremely rare and endangered saprophytic orchid didymoplexis verrucosa (recorded just outside the umnr). the conservation value of these forests is regarded as very high, based on their floristic composition and ecosystem functionality. they also contribute greatly to the stability of the dune fields of southern zululand. based on the vegetation structure and composition, this plant community is regarded as a part of the major vegetation type foz 7 northern coastal forest, as described by mucina and rutherford (2006). although none of the forest communities described in the literature were based on total floristic composition, similar forest types were recorded by venter (1972), weisser (1978b) and macdevette et al. (1989). discussion the ordination clusters closely resemble the classification groupings of plant communities with relatively unique species assemblages. for heterogeneous vegetation types, such as forests, the ordination patterns were less distinct, reflecting the floristic overlap between such plant communities. when viewing the complex of the forest communities along ordination axes 1 and 3, slightly more defined clustering of the various communities occurred. the patterns of species richness recorded within the various plant communities are generally what would be expected. ecosystems that tend to fluctuate between extremes, such as intertidal zones, show relatively low species richness and are dominated by highly adapted stress-tolerant species such as mangrove species. stable, mesic ecosystems tend towards high species richness levels dominated by competitive species. the ecotonal forest edge community, which was predicted to show the highest species richness, showed surprisingly few plant species. however, the suspicion is that the anomaly recorded here may very well have been an artefact of suboptimal sample site selection. the indirect gradient analysis from the ordination scatter plot provided some valuable insights into the general trends in plant community affinities to a variety of environmental variables. based on our deductions, specific environmental drivers of vegetation structure and plant community composition within the umnr include the soil moisture availability, water drainage capability of soils within different parts of the landscape, clay and salt content of soils, the effects of fire and grazing, the effects of organic content in the upper soil profile and the effects of salt-clipping by oceanic salt spray driven by onshore winds. the secondary grasslands within the southern sections of the umnr show very low species richness, and no regional endemic species were recorded. the conversion of forest to secondary grasslands is therefore considered to contribute very little in terms of tropical grassland conservation. these secondary grasslands do not represent any stage or form of the pristine tropical grasslands observed in other nearby reserves under formal protection. however, it must be added that the grasslands and wetlands within the recently acquired northern sections of the umnr are essentially very different from the secondary grasslands along the southern sections. the northern grasslands are recovering remnants of grasslands after the impact of agricultural activities. these tall, northern grasslands provide very valuable habitat for threatened bird species such as the eurasian bittern (botaurus stellaris), african marsh harrier (circus ranivorus), corncrake (crex crex), swamp nightjar (caprimulgus natalensis) and grass owl (tyto capensis). their conservation status, based on the habitat provided to a wide range of fauna species, is therefore regarded as relatively high. the climax coastal dune forest sections within the umnr are species rich, with an even distribution of dominance among the more visually prominent species. despite the many laws and regulations protecting coastal forests, very few outside of formally protected areas are still in an ecologically healthy condition. in light of the ongoing decimation of forests outside of protected areas within southern zululand, the dune forests of the umnr are regarded as forests of very high conservation value. the riverine woodlands and wetlands within the umlalazi river floodplain are relatively species poor and no endemic plant species were recorded there. although these plant communities are not regarded as of very high conservation value based on their floristic composition, they are regarded as very important for ecosystem functioning and stability. however, the recorded levels of invasion by alien plant species within this heterogeneous mosaic of plant communities are of great concern. the saline and intertidal ecosystems within the umnr are not species rich, nor do they contain many endemic plant species. however, based on the ecological importance of mangroves and salt flats along tidal rivers, these ecosystems with their plant communities are critically endangered and under very strict formal protection. it is for this reason that the two mangrove tree species recorded in the umnr, bruguiera gymnorrhiza and avicennia marina, are protected by law. from a sustainable use perspective, the biannual harvesting of materials from the phragmites australis–juncus kraussii saline wetland offers an important example of how nature reserves can contribute to the controlled use of natural resources. the relatively unique prograding (extending) dune system along the umnr provides a classical example of primary succession, the autogenic changes that drive coastal dune succession and the natural rates at which serial stages replace one another. conclusion the described plant communities of the umnr can be seen as surrogates for the ecosystems underlying them. these plant communities should form the basis for conservation and management planning within the umnr. they should also be used as benchmarks and reference examples of undisturbed primary vegetation as well as successional stages in plant community development in order to measure the ecological integrity of similar systems within the maputaland region. based on the floristic similarities between the plant communities of clusters of certain vegetation types, such as forest communities, these vegetation units can be managed and conserved as an integral unit. however, the monitoring of vegetation changes should be conducted on the basis of the individual plant communities. while the management of nature reserves can be done at a relatively coarse scale, monitoring should always be done at the finest relevant scale possible. acknowledgements competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions n.s.z. was the main data collector as part of an msc research project. he contributed to data analysis, interpretations and preparation of the original manuscript. t.h.c.m. was the co-worker and supervisor and contributed to data collection, analysis, interpretations and preparation of the original manuscript and its revision. r.e.m. contributed to literature reviews and comparisons, data analysis, and the editing and preparation of the extensively revised manuscript. references begg, g. & carser, a., 1988, the wetlands of natal (part 2). the distribution, extent and status of wetlands in the mfolozi catchment, natal town and regional planning commission, pietermaritzburg. berlilner, d., 2005, systematic conservation planning for the forest biome of south africa, water and forestry support programme and the department of water affairs and forestry, pretoria. bromberg-gedan, k., silliman, b.r. & bertness, m.d., 2009, ‘centuries of human-driven change in salt marsh ecosystems’, annual review of marine science 1, 117–141. https://doi.org/10.1146/annurev.marine.010908.163930 brown, l.r., du preez, p.j., bezuidenhout, h., bredenkamp, g.j., mostert, t.h.c. & collins, n.b., 2013, ‘guidelines for phytosociological classifications and descriptions of vegetation in southern africa’, koedoe 55(1), art. #1103, 1–10. https://doi.org/10.4102/koedoe.v55i1.1103 chytrý, m., tichý, l., holt, j. & botta-dukát, z., 2002, ‘determination of diagnostic species with statistical fidelity measures’, journal of vegetation science 13, 79–90. https://doi.org/10.1111/j.1654-1103.2002.tb02025.x colloty, b.m., adams, j.b. & bate, g.c., 2002, ‘classification of estuaries in the ciskei and transkei regions based on physical and botanical characteristics’, south african journal of botany 68, 312–321. https://doi.org/10.1016/s0254-6299(15)30392-6 donnelly, f.a. & pammenter, n.w., 1983, ‘vegetation zonation on a natal coastal sand-dune system in relation to salt spray and soil salinity’, south african journal of botany 2, 46–51. https://doi.org/10.1016/s0022-4618(16)30144-9 edwards, d., 1983, ‘a broad-scale structural classification of vegetation for practical purposes’, bothalia 14, 705–712. https://doi.org/10.4102/abc.v14i3/4.1231 ekznw, 2009, amatikulu nature reserve: integrated management plan: 2009–2013, version 1.0, ezemvelo kzn wildlife, pietermaritzburg, p. 82, and 7 maps. fey, m.v., 2010, ‘a short guide to the soils of south africa, their distribution and correlation with world reference base soil groups’, in 19th world congress of soil science, soil solutions for a changing world, brisbane, 1st–6th august, pp. 32–35. gaugris, j.y., matthews, w.s., van rooyen, m.w. & bothma, j.d.p., 2004, ‘the vegetation of tshanini game reserve and a comparison with equivalent units in the tembe elephant park in maputaland, south africa’, koedoe 47(1), 9–29. https://doi.org/10.4102/koedoe.v47i1.67 google earth 7.1.2.2041, 2016, umlalazi nature reserve. 28°57′ 29.53″ s,31°46′ 09.75″ e, elevation 55 m, viewed 22 march 2016, from http://www.google.com/earth/index.html grainger, m.j., van aarde, r.j. & wassenaar, t.d., 2011, ‘landscape composition influences the restoration of subtropical coastal dune forest’, restoration ecology 19, 111–120. https://doi.org/10.1111/j.1526-100x.2009.00630.x hennekens, s.m. & schaminée, j.h.j., 2001, ‘turboveg, a comprehensive data base management system for vegetation data’, journal of vegetation science 12, 589–591. https://doi.org/10.2307/3237010 lubke, r.a., 1997, ‘vegetation and flora of the kosi bay coastal forest reserve in maputaland, northern kwazulu-natal, south africa’, msc thesis, university of pretoria. lubke, r.a., avis, a.m., steinke, t.d. & boucher, c., 1997, ‘coastal vegetation’, in r.m. cowling & s.m. pierce (eds.), vegetation of southern africa, pp. 300–321, cambridge university press, cambridge. macdevette, d.r., macdevette, d.k., gordon, i.g. & bartholomew, r.l.c., 1989, ‘the floristics of the natal indigenous forests’, in i.g. gordon (ed.), natal indigenous forests. a preliminary collection of reports on indigenous forest in natal, pp. 1–20, natal parks board, pietermaritzburg. matthews, w.s., van wyk, a.e. & van rooyen, n., 1999, ‘vegetation of the sileza nature reserve and neighbouring areas, south africa, and its importance in conserving the woody grasslands of the maputaland centre of endemism’, bothalia 29(1), 151–167. https://doi.org/10.4102/abc.v29i1.586 matthews, w.s., van wyk, a.e., van rooyen, n. & botha, g.a., 2001, ‘vegetation of the tembe elephant park, maputaland, south africa’, south african journal of botany 67, 573–594. mccune, b. & mefford, m.j., 1999, pc-ord for windows. multivariate analysis of ecological data, version 4.10, mjm software, gleneden beach, or. moll, e.j., 1972, ‘a preliminary account of the dune communities at pennington park, mtunzini, natal’, bothalia 10(4), 615–626. https://doi.org/10.4102/abc.v10i4.1571 moll, e.j. & werger, m.j.a., 1978, ‘mangrove communities’, in m.j.a. werger & a.c. van bruggen (eds.), biogeography and ecology of southern africa, pp. 1231–1238, dr w. junk, the hague. mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, strelitzia 19, south african biodiversity institute, pretoria. neumann, f.h., scott, l., bousman c.b. & van as, l., 2010, ‘a holocene sequence of vegetation change at lake eteza, coastal kwazulu-natal, south africa’, review of palaeobotany and palynology 162, 39–53. https://doi.org/10.1016/j.revpalbo.2010.05.001 nevill, h. & nevill, e.m., 1995, ‘a survey of the culicoides (diptera: ceratopogonidae) of the umlalazi nature reserve in zululand, south africa, with notes on two species biting man’, onderstepoort journal of veterinary research 62, 51–58. south african weather service, 2017, viewed n.d., from http://www.weathersa.co.za/compliments-complaints/climate-data-requests steinke, t.d., 1995, ‘a general review of the mangroves of south africa’, in g.i. cowan (ed.), wetlands of south africa, pp. 53–74, department of environmental affairs and tourism, pretoria. taylor, r.h., adams, j.b. & haldorsen, s., 2006, ‘primary habitats in st lucia estuarine system, south africa, and their response to mouth management’, african journal of aquatic science 31, 31–41. https://doi.org/10.2989/16085910609503869 tichy, l., 2002, ‘juice, software for vegetation classification’, journal of vegetation science 13, 451–453. https://doi.org/10.1658/1100-9233(2002)013 todd, c.b., 1994, ‘a comparison of the reproductive strategies of key species of a prograding dune system in the mlalazi nature reserve, natal’, msc thesis, rhodes university. tyson, p.d. & preston-whyte, r.a., 2000, the weather and climate of southern africa, 2nd edn., oxford university press, cape town. van wyk, a.e. & smith, g.f., 2001, regions of floristic endemism in southern africa, umdaus press, pretoria. venter, h.j.t., 1972, ‘die plantekologie van richardsbaai, natal’, dsc thesis, university of pretoria. von maltitz, g.p., van wyk, g.f. & everard, d.a., 1996, ‘successional pathways in disturbed coastal dune forest on the coastal dunes in north-east kwazulu-natal, south africa’, south african journal of botany 62(4), 188–195. https://doi.org/10.1016/s0254-6299(15)30633-5 weisser, p.j., 1978a, ‘changes in area of grasslands on the dunes between richards bay and the mfolozi river, 1937 to 1974’, proceedings of the annual congresses of the grassland society of southern africa 13(1), 95–97. https://doi.org/10.1080/00725560.1978.9648841 weisser, p.j., 1978b, ‘conservation priorities in the dune area between richards bay and mfolozi mouth based on a vegetation survey’, natal town and regional planning report 38, 1–64. weisser, p.j., garland, i.f. & drews, b.k., 1982, ‘dune advancement 1937–1977 at the mlalazi nature reserve, mtunzini, natal, south africa, and a preliminary vegetation-succession chronology’, bothalia 14(1), 127–130. https://doi.org/10.4102/abc.v14i1.1152 weisser, p.j. & muller, r., 1983, ‘dune vegetation dynamics from 1937 to 1976 in the mlalazi-richards bay area of natal, south africa’, bothalia 14(3), 661–667. https://doi.org/10.4102/abc.v14i3/4.1225 werger, m.j.a. & coetzee, b.j., 1978, ‘the sudano–zambesian region’, in m.j.a. werger (ed.), biogeography and ecology in southern africa, pp. 231–299, junk, the hague. introduction methods results discussion acknowledgements references about the author(s) clement j. arendse rondevlei scientific services, garden route national park, south african national parks (sanparks), sedgefield, south africa ian a. russell rondevlei scientific services, garden route national park, south african national parks (sanparks), sedgefield, south africa citation arendse, c.j. & russell, i.a., 2020, ‘morphometric regressions for the endangered knysna seahorse, hippocampus capensis, in the swartvlei estuary from mass stranding events’, koedoe 62(1), a1632. https://doi.org/10.4102/koedoe.v62i1.1632 short communication morphometric regressions for the endangered knysna seahorse, hippocampus capensis, in the swartvlei estuary from mass stranding events clement j. arendse, ian a. russell received: 08 apr. 2020; accepted: 22 july 2020; published: 28 sept. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. introduction the knysna seahorse, hippocampus capensis, is listed as endangered on the iucn red list of threatened species, with the only known, apparently isolated, populations occurring in the knysna, swartvlei and keurbooms estuaries along the south coast of south africa. the swartvlei estuary population is the most genetically distinct (mkare et al. 2017). despite all three known populations occurring within protected areas and there being no commercial or subsistence use of the species, population sizes are still thought to be declining because of habitat degradation across much of its range (pollom 2017). environmental management authorities have outlined strategies for the conservation of h. capensis, including listing it as a species of special concern in the estuarine lower level plan of the garden route national park (south african national parks 2020) and setting management objectives for the keurbooms population in the keurbooms–bitou estuarine management plan (western cape government 2018). assessments of population size and habitat use of h. capensis have focussed mainly on the relatively larger and more stable population in knysna estuary (bell et al. 2003; claassens 2016; claassens & hodgson 2018; teske et al. 2007). the lesser studied swartvlei estuary population is subject to large fluctuations in abundance, with past densities ranging from 0.04 individuals per m2 to 0.26 individuals per m2 (lockyear et al. 2006). the swartvlei estuary is the only intermittently open host estuary, with elevated water levels and prolonged flooding of land adjacent to the shores common during closed periods. periodic mass strandings and associated die-offs of h. capensis have been recorded in swartvlei estuary when the estuary is breached (russell 1994; skelton 1987), mainly by artificial means to prevent inundation of properties on the estuarine floodplain. the breaching protocol and criteria are contained with the garden route national park management plan (sanparks 2020). south african national parks (sanparks) as the management authority of swartvlei estuary has since 1991 intermittently collected dead h. capensis from the banks of the swartvlei estuary after breaching events. this now regular sampling has evolved into a sanparks co-ordinated citizen science programme. the only known morphometric descriptions of h. capensis come from small samples of flood-stranded individuals from the knysna estuary (n = 57) and swartvlei estuary (n = 39) (toefy 2000), despite the usefulness of such data in allowing comparison between studies and the conversion of data where different measures of length are used (lourie 2003). this article aims to use the seahorse morphometric data collected following two mass stranding events (> 500 individuals) in 1991 and 2017 to provide more robust estimates of the relationship between key morphometric characteristics of h. capensis. in addition, data from mass strandings are compared to those collected from seine netting of live individuals in 2019 to determine the feasibility of using these periodic events as a means of assessing population attributes. methods sample sites hippocampus capensis were sampled from the swartvlei estuary (34° 01’ 48.58” s, 22° 47’ 50.66” e), situated along the south coast of south africa. collection protocol dead h. capensis were collected after mass stranding events in february 1991 and november 2017. specimens collected in 1991 were preserved in 10% neutral freshwater-buffered formalin immediately after collection (russell 1994), whereas in 2017 dead individuals were frozen immediately after collection. in 2017, samples were collected within 1 day of stranded individuals first being noticed, and for 1 week after estuary breaching. sampling effort was spread throughout the estuary, although repeat sampling, where sites were searched daily over the sampling period, was undertaken mostly in areas with previous high abundance (lockyear et al. 2006). approximately 1000 individuals were recorded, of which half (52%) were dead. all live individuals were returned to the water. retained dead individuals showed various levels of desiccation, likely a result of differences in exposure time because of the intermittent stranding of h. capensis over several days following breaching. samples were pooled for all sites, and all samples were preserved without manipulating (i.e. straightening) body parts. live h. capensis were collected in march 2019 using a 10 m × 1.5 m seine net with a stretched mesh size of 8 mm. three effective hauls were undertaken in ≤ 1.3 m water depth in areas of previous high h. capensis density, with each haul being approximately 3 m wide, ranging from 12 m to 21 m long, and spaced at least 20 m apart. data were pooled for all replicates. live samples were compared to preserved samples to determine if size classes and sex ratios differed between sampling techniques. morphometrics preserved samples were photographed and measured using imagej software. where necessary, individuals were flattened before being photographed to minimise bias in measurements because of body arching (discussed in sotola et al. 2019). all individuals were sexed and then measured using the protocol set by lourie (2003). head length (hdl), curved trunk length (trl), curved tail length (tal) and height (ht) were measured, with standard length (sl) calculated as the sum of the first three measurements. straight-line sl (slstraight) and straight-line ht (htstraight) were also determined for a subsample of individuals whose heads were positioned at a right angle to their trunk. the trunk measurements for the straight-line sizes were measured as straight-line distances from the respective anterior starting points ending at the last trunk ring. regression equations of size measurements of h. capensis included measurement of htstraight and ht because htstraight is often the size measurement used when sampling live specimens (foster & vincent 2004; lourie 2003), likely to avoid handling stress, and ht may be useful if the snout of a dead seahorse is damaged. live individuals collected in 2019 were sexed and htstraight was measured before the animals were released at the location of capture. the effect of preservation techniques on morphometric relationships was investigated by comparing the regression equations between sl and hdl, trl, tal and htstraight of formalin preserved (collected 1991) and frozen (collected 2017) specimens. preservation of samples in a fixative can cause shrinkage, and different preservation techniques may result in different shrinkage rates of seahorse body parts (nadeau, curtis & lourie 2009). also, further shrinkage is known to occur in fish when multiple preservation techniques, in this case freezing or preservation in formalin after desiccation, are applied (sotola et al. 2019). statistical analysis sex ratio bias within the 1991, 2017 and 2019 datasets was tested using an exact binomial test (crawley 2007; zar 2010). sexes were analysed separately because of known sexual dimorphism (bell et al. 2003). normality of sl (between preservation techniques) and htstraight (between alive and preserved specimens) measurements for years and sexes were tested using a shapiro–wilk test. homogeneity of variances was tested using a fligner–killeen test before differences in means were tested using either a one-tailed welch’s two-sample t-test or a one-tailed wilcoxon rank-sum test. differences in mean htstraight between years were tested using a kruskal–wallis rank-sum test. regression equations of sl and htstraight and their component measurements for years and sexes were produced separately using ordinary least squares regressions and compared to determine if sex and preservation techniques influenced correlation between variables. regression equations of sl and htstraight, separated by year and sex, were tested using an analysis of covariance (ancova) to determine if regression equations obtained differed significantly (crawley 2007; zar 2010). data analyses were undertaken using the programming language r (r core team 2019). results sex ratios sex ratios were biased towards females in the 1991 sample (1.025:1; n = 170) and males in the 2017 and 2019 samples (0.974:1; n = 304 and 0.814:1; n = 78, respectively), although none differed significantly from the predicted 1:1 ratio (p > 0.05). size class distribution mean length of h. capensis collected in 2017 (62 mm – 100 mm sl; n = 274) was significantly larger than the mean length of those collected in 1991 (33 mm – 100 mm sl; n = 161) when sexes were grouped (w = 7800; p < 0.05) as well as for females (w = 1650; p < 0.05) and males (w = 2227; p < 0.05) separately. the majority of individuals collected in 2017 were above the size of sexual maturity (≥ 65 mm sl [whitfield 1995]), whereas most individuals collected in 1991 were likely sexually immature (figure 1a). mean sl differed significantly between sexes in both 1991 (w = 2255; p < 0.05) and 2017 (t = −2.989; p < 0.05), with the mean size of males being larger than that of females in both years. figure 1: box and whisker plots of hippocampus capensis separated by sex showing the minimum, first quartile, median, third quartile and maximum sizes of (a) sl for preserved (1991 and 2017) samples, and (b) htstraight for preserved (1991 and 2017) and live samples (2019). measurements of htstraight from live h. capensis collected in 2019 were normally distributed for both females (w = 0.946; p > 0.05) and males (w = 0.984; p > 0.05). mean htstraight did not differ significantly between sexes (t = −0.432; p > 0.05). the 2019 size range (38 mm – 71 mm htstraight; n = 78) was similar to those collected in both 1991 (35 mm – 76 mm htstraight; n = 53) and 2017 (54 mm – 77 mm htstraight; n = 97), though mean sizes differed significantly for both females (chi-squared = 51.875; p < 0.05) and males between years (chi-squared = 52.689; p < 0.05) (figure 1b). morphometric regressions the fit of regression models between sl and hdl, trl and tal was better for the 1991 samples compared to those collected in 2017, as evidenced by the high r2 values in the former (tables 1 and 2). test statistics for the regressions between sl and hdl, trl, tal and htstraight are presented in table 3. hdl, trl and tal were significantly correlated to sl for both sexes when years were analysed separately, whilst sex effected all regressions except hdl in 1991. interaction did not significantly effect the fit of the models in either year (p > 0.05) and the slope of the models did not differ significantly between sexes (p > 0.05), with the exception of hdl in 2017. when differences between years were calculated separately for sexes, interaction effects were evident for the female datasets for trl and tal and male datasets for hdl and trl (p > 0.05) but not the female dataset for hdl or the male dataset for tal. year significantly effected all regressions except the male dataset for trl. slopes differed between years for all variables for both sexes. preservation technique, therefore, influenced trl for both sexes, whereas hdl only differed between males and tal between females. when years were analysed individually, htstraight significantly effected sl in 1991 and 2017 (p < 0.05), but sex effected the regressions only in 2017 (p < 0.05). no interaction effects were evident in either year (p > 0.05). when sexes were analysed separately, htstraight and year significantly influenced the regressions for both sexes, with no interaction effects evident for females or males (p > 0.05). the linear regression for males collected in 1991 (table 1) was used to calculate htstraight at sexual maturity (56 mm) (figure 1b). table 1: correlation between selected morphometric characteristics of hippocampus capensis collected in 1991. table 2: correlation between selected morphometric characteristics of hippocampus capensis collected in 2017. table 3: test statistics of regression models of standard length with head length, curved trunk length and curved tail length, showing interaction between the measured variables and sexes and years, respectively. discussion the 1991 and 2017 datasets provided reliable correlations between the morphometric variables measured for the different preservation techniques. both preservation techniques used in this study are known to result in significant shrinkage of fish bodies (berbel-filho, jacobina & martinez 2013; buchheister & wilson 2005; nadeau et al. 2009) with the preservation of samples in formalin and ethanol resulting in ongoing shrinkage of body parts (martinez, berbel-filho & jacobina 2013; sotola et al. 2019). to compound the complexity of using preserved specimens in morphometric investigations, shrinkage in fish may not be uniform across body parts and differences can even be observed at an individual level (sotola et al. 2019). this effect was noted in this study with the regressions of sl with hdl, trl and tal of h. capensis collected in 2017, showing greater variation than those collected in 1991. although the individuals collected in 2017 were frozen before processing, they showed differing levels of desiccation when collected. this, coupled with an extended, staggered collection period for the stranded seahorses, and the relatively shorter period between preservation and processing of samples compared to the 1991 dataset, could have contributed to the larger variation in the body length ratios. the greater correlation between morphometric measurements of sl and hdl, trl and tal in 1991 was therefore likely because of more even shrinkage resulting from prolonged exposure of the samples to the fixative. this result, together with the differences in slopes between the 2 years, suggests that different regression equations are needed for each preservation type and that comparisons between samples preserved in different fixatives may require correction factors to be calculated by measuring samples before and at several periods after preservation. in addition, using regression equations obtained from preserved specimens to describe morphometric relationships of live specimens is not recommended. the mean length (htstraight) of h. capensis differed significantly between 1991, 2017 and 2019 for both sexes. differences could relate to the timing of the breaching events, with both the 1991 and 2019 samples collected towards the end of the breeding period for h. capensis (austral summer – whitfield [1995]), whereas the 2017 samples were collected at the start of the breeding period, potentially explaining the lower proportion of smaller individuals in that year. the duration of preservation of specimens could also have contributed to differences in mean length, with specimens collected in 1991 being subjected to longer exposure to a fixative likely having shrunk to a greater degree. although differences in the size data existed, the slopes of the sl versus htstraight regressions for 1991 and 2017 did not differ significantly from one another, indicating that the relationship between sl and htstraight remains relatively constant regardless of preservation technique used. this was, however, not the case for the relationship between sl and hdl, trl and tal, where preservation technique, as well as sex, influenced the slopes of the regressions. as various length measurements are often used for measuring seahorses, standardisation of seahorse measurements is difficult (foster & vincent 2004). the morphometric regressions presented in tables 1 and 2 can assist in comparing the results of previous studies where different measurements and preservation methods were used. equal sex ratios observed in 1991 and 2017 were similar to those found in in situ studies of natural populations (lockyear et al. 2006) and in 2019, suggesting that stranding affected both sexes equally. the use of stranding data for h. capensis in the swartvlei estuary could be a novel way of post hoc determination of population attributes, as data collected from these events are comparable to those collected from the study of in situ populations. the effects of preservation on morphometric variables also need additional consideration, and shrinkage correction equations should be developed for different preservation methods and durations. acknowledgements the authors would like to thank the citizen scientists, sanparks staff and honorary rangers for their assistance with data collection. they would also like to thank dr mohlamatsane mokhatla for his help with the statistical programme r. competing interests the authors have declared that no competing interest exists. authors’ contributions c.j.a. and i.a.r. conceptualised the study, conducted fieldwork and contributed to writing the article. c.j.a. performed all laboratory and statistical analysis. ethical consideration this study fully meets the ethical standards for handling of hippocampus capensis as set out by the sanparks animal use and care committee (004/17). funding information this research received no specific grant from any funding agency in the public, commercial or not-for-profit sectors. data availability statement data can be requested via the sanparks data repository (http://dataknp.sanparks.org/sanparks/). data use limitations may apply. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references bell, e.m., lockyear, j.f., mcpherson, j.m., marsden, a.d. & vincent, a.c.j., 2003, ‘first field studies of an endangered south african seahorse, hippocampus capensis’, environmental biology of fishes 67(1), 35–46. https://doi.org/10.1023/a:1024440717162 berbel-filho, w.m., jacobina, u.p. & martinez, p.a., 2013, ‘preservation effects in geometric morphometric approaches: freezing and alcohol in a freshwater fish’, ichthyological research 60(3), 268–271. https://doi.org/10.1007/s10228-013-0339-x buchheister, a. & wilson, m.t., 2005, ‘shrinkage correction and length conversion equations for theragra chalcogramma, mallotus villosus and thaleichthys pacificus’, journal of fish biology 67(2), 541–548. https://doi.org/10.1111/j.0022-1112.2005.00741.x claassens, l., 2016, ‘an artificial water body provides habitat for an endangered estuarine seahorse species’, estuarine, coastal and shelf science 180, 1–10. https://doi.org/10.1016/j.ecss.2016.06.011 claassens, l. & hodgson, a.n., 2018, ‘gaining insights into in situ behaviour of an endangered seahorse using action cameras’, journal of zoology 304(2), 98–108. https://doi.org/10.1111/jzo.12509 crawley, m.j., 2007, the r book, john wiley & sons, ltd, the atrium, southern gate, chichester, west sussex, england. foster, s.j. & vincent, a.c.j., 2004, ‘life history and ecology of seahorses: implications for conservation and management’, journal of fish biology 65(1), 1–61. https://doi.org/10.1111/j.0022-1112.2004.00429.x lockyear, j.f., hecht, t., kaiser, h. & teske, p.r., 2006, ‘the distribution and abundance of the endangered knysna seahorse hippocampus capensis (pisces: syngnathidae) in south african estuaries’, african journal of aquatic science 31(2), 275–283. https://doi.org/10.2989/16085910609503897 lourie, s., 2003, measuring seahorses, project seahorse technical report no. 4, version 1.0, p. 15, project seahorse, fisheries centre, university of british columbia. martinez, p.a., berbel-filho, w.m. & jacobina, u.p., 2013, ‘is formalin fixation and ethanol preservation able to influence in geometric morphometric analysis? fishes as a case study’, zoomorphology 132(1), 87–93. https://doi.org/10.1007/s00435-012-0176-x mkare, t.k., jansen van vuuren, b. & teske, p.r., 2017, ‘conservation implications of significant population differentiation in an endangered estuarine seahorse’, biodiversity and conservation 26(6), 1275–1293. https://doi.org/10.1007/s10531-017-1300-5 nadeau, j.l., curtis, j.m.r. & lourie, s.a., 2009, ‘preservation causes shrinkage in seahorses: implications for biological studies and for managing sustainable trade with minimum size’, aquatic conservation: marine and freshwater ecosystems 19(4), 428–438. https://doi.org/10.1002/aqc.1002 pollom, r.a., 2017, ‘hippocampus capensis’, the iucn red list of threatened species 2017, e.t10056a54903534. r core team, 2019, r: a language and environment for statistical computing, r foundation for statistical computing, vienna, viewed 07 october 2019, from https://www.r-project.org/. russell, i.a., 1994, ‘mass mortality of marine and estuarine fish in the swartvlei and wilderness lake systems, southern cape’, southern african journal of aquatic sciences 20(1–2), 93–96. https://doi.org/10.1080/10183469.1994.9631353 skelton, p.h., 1987, south african red data book – fishes, pretoria, south africa. sotola, v.a., craig, c.a., pfaff, p.j., maikoetter, j.d., martin, n.h. & bonner, t.h., 2019, ‘effect of preservation on fish morphology over time: implications for morphological studies’, plos one 14(3), e0213915. https://doi.org/10.1371/journal.pone.0213915 south african national parks, 2020, garden route national park management plan, p. 222, groenkloof, south africa. teske, p.r., lockyear, j.f., hecht, t. & kaiser, h., 2007, ‘does the endangered knysna seahorse, hippocampus capensis, have a preference for aquatic vegetation type, cover or height?’, african zoology 42(1), 23–30. https://doi.org/10.3377/1562-7020(2007)42[23:dteksh]2.0.co;2 toefy, z., 2000, ‘a preliminary examination of the morphological and genetic structure within populations of the knysna seahorse hippocampus capensis (pisces: syngnathidae) from two south african estuaries’, m.sc dissertation, university of the western cape, bellville, south africa, p. 83. western cape government, 2018, keurbooms-bitou estuarine management plan, cape town, south africa, p. 139. whitfield, a.k., 1995, ‘threatened fishes of the world: hippocampus capensis boulenger, 1900’, environmental biology of fishes 44(2), 362. https://doi.org/10.1007/bf00008251 zar, j.h., 2010, biostatistical analysis, 5th edn., prentice hall, inc, upper saddle river, new jersey. abstract introduction research method and design confounding effects results discussion conclusion acknowledgements references about the author(s) hloniphani moyo school of animal, plant and environmental sciences, university of the witwatersrand, south africa mary c. scholes school of animal, plant and environmental sciences, university of the witwatersrand, south africa wayne twine school of animal, plant and environmental sciences, university of the witwatersrand, south africa citation moyo, h., scholes, m.c. & twine, w., 2016, ‘effects of water and nutrient addition on the coppice growth response of cut terminalia sericea’, koedoe 58(1), a1371. http://dx.doi.org/10.4102/koedoe.v58i1.1371 original research effects of water and nutrient addition on the coppice growth response of cut terminalia sericea hloniphani moyo, mary c. scholes, wayne twine received: 25 nov. 2015; accepted: 30 mar. 2016; published: 22 july 2016 copyright: © 2016. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the ability of a woody plant to coppice and remain vigorous largely depends on the severity of disturbances, resource availability and the mobilisation of stored reserves. there is limited information about the role played by resource limitation on the recovery of cut trees. this study investigated the effects of water and nutrient supplementation on coppice growth responses of resprouting cut trees in a semi-arid savannah in south africa. cut trees were exposed to different levels of water and nutrient (nitrogen and phosphorus) supplementation over a period of 2 years in a factorial experimental design. we hypothesised that adding water and nutrients would result in an increased coppice growth response and replenishment of stored structural reserves. adding water and nutrients significantly increased shoot diameter, shoot length and resprouting ratio for the initial 12 months after cutting but not stored structural reserves. such a response pattern suggests that the initial growth of resprouting shoots may be strongly resource-limited, while resources are concentrated on supporting fewer resprouting shoots compared to a higher number. conservation implications: if practicing rotational tree harvesting, trees resprouting in resource-poor locations need a longer resting period to recover stored reserves and to also recover lost height after cutting. introduction plants are subjected to various forms of disturbances and stresses that affect their growth, production and survival. when recovering lost biomass after a disturbance, resource availability and exposure to further disturbances limit the full recovery of plants growing in a disturbed ecosystem (hoffmann, schroeder & jackson 2002; holdo, holt & fryxell 2009). thus, there is a need to understand the importance of tree resource use as well as the factors that affect tree recovery as influenced by climatic factors. an understanding of how disturbances and resource availability interact will have further management implications for the long-term productivity and sustainability of woody vegetation (neke 2004; neke, owen-smith & witkowski 2006; pote et al. 2006). as a result of exposure to disturbances, coppicing (resprouting) may be central to the survival and resilience of trees in disturbance-prone systems (avohou et al. 2011). coppicing can be defined as the production of vegetative shoots at the base of the stem or sprouts arising from a stump, resulting in the emergence of new shoots from the stump or roots (hardesty 1984; laureysens et al. 2003; van wyk & van wyk 1997). following a disturbance event, an increase in nutrient and water availability stimulates growth rates and allocation to storage in resprouting organs (cruz et al. 2002; sankaran, ratnam & hanan 2008). this is supported by the study of cruz et al. (2002), who suggest that resprout growth is limited by the low amount of moisture during the early stages of regeneration. in other instances, an increase in water and nutrient availability after a disturbance may increase woody plant vegetation, leading to bush encroachment and therefore reducing biodiversity and the overall productivity of the ecosystem (ward 2005). for example, water availability was shown to be positively correlated with regeneration in acacia tortilis (forssk) in tunisia (noumi et al. 2010) and in populus tristis fisch and populus balsamifera in the usa (dickmann, nguyen & pregitzer 1996). nutrients play a key role in replenishing depleted stored nutrient reserves of plants after a disturbance such as tree cutting (bowen & pate 1992) and their interaction with moisture affects coppice regrowth (castell & terradas 1994). because nitrogen (n) in the remaining plant parts is used for reconstructing new tissue, adding more n increases root n storage for later use after a disturbance (el omari et al. 2003; kabeya & sakai 2005). more n also increases concentrations of photosynthetic enzymes, which in turn cause a higher rate of photosynthesis in leaves (chapin iii 1991). in fertilised resprouting erica australis l. com., trees grew faster and produced higher biomass than un-fertilised trees 2 years after fertiliser application in spain (cruz et al. 2002). nutrient addition also increased both diameter and height of betula pendula roth. and willow (salix spp.) stands in finland (hytonen & kaunisto 1999). it has been suggested that stored carbohydrate reserves and surviving meristems are the most important resources controlling resprouting after disturbances, with differences in soil nutrient levels also affecting stored carbohydrate reserves (bellingham & sparrow 2000; bond & midgley 2001; kabeya & sakai 2005). for example, carbohydrate reserve concentration in resprouting quercus crispula blume was positively correlated to resprouting shoot length and leaf number (kabeya et al. 2003), while starch concentrations in roots of coppicing salix viminalis l., were higher in trees that received higher levels of n compared to plants that received lower levels (fircks & sennerby-forsse 1998). surprisingly, little is known about the regeneration strategies of woody species in tropical and subtropical savannahs (neke 2004; pote et al. 2006). this is because most studies have focused on ecosystems that differ markedly from african woodlands, such as temperate ecosystems (wilson 2002). such ecosystems differ in terms of the seasonality and distribution of rainfall, tree population structure, moisture levels and tree densities. most studies on tree responses have also strongly focused on the effects of varying resource levels on seedling growth and survival (cheng & fuchigami 2002; druege et al. 2000; kraaij & ward 2006; scogings & mopipi 2008; wendler & millard 1996; wilson & witkowski 1998), rather than mature and established trees. knowing such information will provide clues for understanding mechanisms underlying a tree’s regrowth strategies after a disturbance. also, such information would assist in evaluating how changes in resource availability may impact a community exposed to different disturbance types and frequencies. this study was carried out to investigate how the addition of water and nutrients interact to influence tree regrowth (in terms of shoot production, shoot length and diameter, and resprout ratio) as well as the total non-structural carbohydrate (tnc) storage capacity after cutting. the following question was asked: how do varying levels of water and nutrient supplementation influence the coppice response and stem tnc reserves of cut trees? it was hypothesised that supplementing resprouting trees with water and nutrients would increase coppice response and would also result in higher tnc levels in supplemented trees compared with unsupplemented trees. research method and design the study site and the experimental design were described previously (moyo, scholes & twine 2015b); a brief description of both the site and the design follows. setting the experiment was conducted at the wits rural facility (wrf), a 350-ha research station in the central savannah low-veld region of limpopo province, south africa (24° 30′ s; 31°06′ e). the study site is semi-arid, with a mean annual rainfall of ~650 mm, concentrated in the summer season (between october and april) (figure 1b) (kaschula, twine & scholes 2005a; neke et al. 2006; shackleton 1997). the study spanned a 2-year period, starting at the end of the dry season of 2010 (september 30), during the period when trees begin leaf-flush, and ending in september 2012 (moyo, scholes & twine 2015a; moyo et al. 2015b). figure 1: (a) long-term mean monthly rainfall distribution (1992–2010) for the wits rural facility and (b) cumulative monthly rainfall data for the duration of the study period (september 2010 – september 2012). january 2012 received exceptionally high amounts of rainfall compared to other january figures for the past 18 years. design a 3 × 3 factorial experiment (nine plots in each site) replicated in three sites, was established in september 2010 to determine the effects of water and nutrient additions on the coppice response of terminalia sericea. all three sites were on the slope crests. therefore, they had shallow, coarse-textured and dystrophic soils (neke 2004; shackleton 1997; shackleton 1999) (moyo et al. 2015). plots within a site were separated by a distance of approximately 15 m, with a density of approximately 1400 trees per ha. ten trees were selected per plot (with a distance of about 2 m separating each tree from the adjacent one) with the number totalling 270 for the experiment. while root systems of t. sericea extend horizontally, this distance was considered adequate to avoid uptake of water and nutrients by the neighbouring trees. trees were cut at a standard height of approximately 25 cm from the ground, because research has shown that cutting height influences coppice response (ibrahima et al. 2007; kaschula, twine & scholes 2005b; khan & tripathi 1986; shackleton 1997) (moyo et al. 2015b). a 30-cm radius was marked around each tree, and grass was cleared to reduce competition for water and nutrients. long-term average monthly (18 years) rainfall data (figure 1) from the wrf records were used to derive monthly water treatments. water addition treatments were as follows (moyo et al. 2015b): no water addition (w0) – no water additions throughout the study. low (w+) – trees were supplemented with an amount of water 0.5 times the long-term mean rainfall for that month. high (w++) – trees were supplemented with the long-term mean rainfall for that month (moyo et al. 2015b). this monthly amount of rainfall was then divided by four to obtain weekly amounts for supplementing trees for a period of about 18 months beginning in september 2010. water was applied next to the base of each stump using a watering can (moyo et al. 2015b). for the experimental plots that had nutrient additions, nitrogen (n) and phosphorus (p) were supplied using a commercial fertiliser in the form of ammonium phosphate, with a total of 80 kg n ha−1 for the high–fertiliser addition treatment and 40 kg n ha−1 for the low–fertiliser addition treatment. considering that the amount of fertiliser was added once at the beginning of each of the two growing seasons during the study (therefore, twice during the study – in october 2010 and october 2011), this amount (160 kg n ha−1 in total was added) was regarded as a high rate of n application in several studies (le roux & mentis 1986; mbatha & ward 2010; tilman 1987) (moyo et al. 2015b). fertiliser was added after the first rains fell in october 2010 and also in october 2011. fertiliser application was done in split applications compared to a single application to avoid, over the course of the wet season, possible leaching of fertiliser through rainfall and added water if applied all at once. a commercial dry fertiliser blended as 4:3:4:1 (n, p, k, and zn) and mixed according to 120 g/kg n and 90 g/kg p was purchased. fertiliser additions, calculated according to n and p on the commercial fertiliser, were as follows (figure 2): no fertiliser addition (n0) – no fertiliser additions throughout the study. low (n+) – 0.13 g of n and 0.1 g of p per tree. high (n++) – 0.27 g of n and 0.2 g of p per tree (moyo et al. 2015). figure 2: factorial experiment designed to investigate the influence of water and nutrient additions on the coppice response of harvested trees. the dry fertiliser was applied by coring a hole next to the base of the stump with a diameter of 3 cm and a depth of about 10 cm and depositing the fertiliser into the hole. the assumption was that all the fertiliser added using this method was utilised exclusively by the fine tree roots with loss, if any, of approximately the same for the treatments that received fertiliser addition (moyo et al. 2015b). a combination of the treatments, for example, w++n0 would mean the treatment is high water addition (w++) and no nutrient addition (n0). morphological measurements from each resprouting stump, the following variables were measured per stump on a monthly basis: (1) total number of shoots, (2) resprout shoot diameter (measured above the resprouting zone) and (3) resprout shoot length. for (2) and (3), the leader shoot showing the greatest length was selected and marked using strings. shoot production was calculated as the number of shoots produced per unit area of stump basal area. the tree’s original stump diameter was measured at the beginning of the experiment and used together with resprout shoot diameter to calculate the resprout shoot diameter as a ratio of the original tree’s stump diameter. this was termed the resprout ratio. total non-structural carbohydrate sampling and determination to obtain an indication of tnc dynamics in t. sericea and also to understand how adding water and nutrients affected tnc levels, all trees exposed to treatments as illustrated in figure 2 were sampled for stem tnc levels twice during the course of the study: after a year of treatment application (30 september 2011) after 18 months of treatment application, that is, at the end of the wet season (30 april 2012). three trees per treatment were sampled using an increment borer (4.3 mm diameter) to extract two 3-cm-long wood cores from the stumps. before storing the cores in airtight plastic bottles, the outer tree bark was immediately removed from cores after sampling. core samples were then bottled and placed inside a freezer. drying of samples was done at 65 °c for 2 days (assumed dry) and samples were ground using a coffee grinder before finer grinding through a 40-µm mesh wiley mill screen, at the university of florida, usa (moyo, scholes & twine 2015a). a composite sample for each treatment was obtained after grinding. the tnc was determined on a dry weight basis using the anthrone method described by edwards, downie & clingeleffer (2011). the tnc values reported here are the sum of the soluble (glucose and sucrose) and insoluble (starch) fractions (moyo et al. 2015a). for comparison, cores from three non-harvested and unsupplemented trees, about 20 m from the experimental sites, with the same diameter (measured 25 cm above ground) were sampled for tnc analysis. confounding effects because of the presence of antelopes at the study site, the possibility of browsing after tree cutting was very high. the proportion of shoots that were browsed was estimated by counting the total number of shoots resprouting per stump, at the time of recording morphological changes, together with the total number of shoots browsed. the browse proportion was then calculated as the number of browsed shoots relative to the total number of shoots resprouting per stump, and expressed as a percentage. the browse proportion was compared across all the treatments for the duration of the study. analyses the data were partitioned into two 12-month time intervals to test for the effects of water and nutrient additions on cut trees using linear mixed models implemented in statistical analysis software (sas) with proc generalised linear model. the september 2010 to september 2011 interval represented the end of the initial 12 months while the september 2011 to september 2012 interval represented the second 12-month period. prior to analysis, the total number of shoots, shoot diameter and length data were log-transformed because data were not normally distributed. for the number of shoots, a generalised linear mixed model was fitted using an underlying poisson distribution with a logit link, which included the fixed effects of treatment. in addition, random effects in terms of site and tree within site were included in the model. all models were fitted using the procedure glimmix as implemented in sas v. 9.2 (sas institute, cary, nc, usa) (moyo et al. 2015a). significance of fixed effect terms were evaluated with an approximated f-test, with mean comparisons between treatment regimens obtained using least square means with dunnett’s t3. the simple means comparison (unprotected) was used to obtain the level of significance for the total number of shoots. the effects of water and nutrient addition on coppice response (shoot diameter and length) and tncs were analysed using analysis of variance, and dunnett’s t3 test was used for mean comparison at p < 0.05 (moyo et al. 2015a). results effects of water and nutrient additions on coppice response twelve months after trees were cut (september 2011), in the no–water addition treatment (w0), shoot production was significantly higher when water was interacting with high nutrient addition (w0n++) than the other nutrient addition treatments. shoot production was lowest when water was interacting with high nutrient addition, in the low–water addition treatment (w+n++) (figure 3a). there were no significant differences in shoot production in the w++ treatment interacting with nutrient treatments in september 2011. the second year after cutting (september 2012), there were no significant differences in shoot production in all water treatment levels interacting with nutrient addition treatments. between the sampling dates, shoot production reduced markedly across all treatments (figure 3a). figure 3: effects of water and nutrient addition on (a and b) shoot production (calculated as number of shoots per unit area of original stump circumference cm2), (c and d) shoot diameter, (e and f) shoot length and (g and h) resprout ratio (calculated as the ratio of the tree’s original stump diameter to that of the leader shoot resprout diameter) between september 2011 and september 2012. a year after trees were cut and under w0, w0n0 recorded the lowest shoot diameter compared to w0n+ and w0n++ (figure 3b). for the w+ treatment, shoot diameter was significantly higher under the w+n++, compared with w+n0 and w+n+. for the w++ treatment, significantly higher shoot diameter was only noted when water was interacting with lower nutrient addition (w++n+) compared to w++n0, and not when compared with w++n++. the second year after cutting, there were no significant differences in shoot diameter in the w0 and w++ treatments interacting with nutrient addition treatments, while in the w+ treatment, w+n+ and w+n++ recorded significantly higher shoot diameter than w+n0 (figure 3b). a year after cutting, under w0, shoot length was highest under no nutrient addition (w0n0); in both the w+ and w++ treatments, shoot length was lowest at the high–nutrient addition level (w+n++ and w++n++). the second year after cutting and under w0, shoot length was highest under high nutrient addition (w0n++), while under w+ and w++, shoot length was lowest under no nutrient additions (w+n0 and w++n0) (figure 3c). a year after cutting, resprout ratio was significantly lowest for w0n0 and highest under w0n++ and w++n++ for the respective water treatments. for the w0 treatment in september 2012, resprout ratio was lowest in w0n+, while highest for the w++n++ under the w++ (figure 3d). interactive effects of water and nutrient additions on coppice response a year after cutting, shoot production was highest when water and nutrient additions were low (w+n+), with no significant differences observed 2 years after the trees were cut (figure 4a). in september 2011, shoot diameter was significantly highest under the w+n++, with w+n0 recording the lowest shoot diameter in september 2012 (figure 4b). in september 2011, shoot length was significantly higher under the w++n++, while in september 2012, w++n++ and w++n+ recorded the highest shoot length (figure 4c). the resprout ratio was significantly highest for the w++n++ for both the sampling dates, that is, september 2011 and 2012 (figure 4d). figure 4: effects of water and nutrient interaction on (a) shoot production (calculated as the number of shoots per unit area of original stump circumference cm2), (b) shoot diameter, (c) shoot length and (d) resprout ratio (calculated as the ratio of the tree’s original stump diameter to that of the leader shoot resprout diameter) between september 2011 and september 2012. effect of water and nutrient additions on stem total non-structural carbohydrate levels supplementing cut trees with water and nutrients had no significant effect on stem tnc concentration for both sampling dates (september 2011 and april 2012) (table 1). uncut trees recorded significantly higher tnc levels compared to unsupplemented trees (w0n0) in september 2011. table 1: differences in total non-structural carbohydrate (mg g−1 dw) levels between september 2011 and april 2012 as a function of water and nutrient additions. confounding effects: browsing browsing (percent of shoots browsed per stump) on resprouting trees was widespread in both growing seasons during the study, although it was lower in the second compared to the first. there were no significant differences (f = 1.57, p = 0.167) in browsing proportion across all treatments during the study. discussion water and nutrient addition had initial significant effects on shoot production, while, as hypothesised, the addition of water and nutrients resulted in significant increases in resprout shoot diameter and shoot length. contrary to the hypothesis, adding water and nutrients did not result in significant increases in stem tnc levels at both sampling dates, september 2011 and april 2012. effects of water and nutrients on coppice response of cut trees high water and nutrient addition had a significant effect on shoot production after 12 (but not after 24) months. the differences in tree responses can possibly be attributed to the higher amount of rainfall received in the 2012 rainfall season compared to the 2011 season, contributing to treatment effects being non-measurable in 2012. alternatively, this pattern could also be interpreted as generally the speeding up of the resprouting process because resprouting trees start off with many small shoots and, over time, tend to invest in fewer and larger shoots. therefore, the effect of water and nutrients could have been to speed up the rate at which this shift in allocation happens. similar nutrient addition effects have been reported when there was an increase in the above-ground shoot production in pinus taeda l., pinus elliottii and liquidambar styraciflua l. (albaugh et al. 2004; cobb et al. 2008), in populus deltoids bartr. and planatus occidentalis l. (coyle & coleman 2005), and shoot biomass and growth in quercus macrocarpa michx. and northern pin oak quercus ellipsoidalis e. j. hill (davis et al. 1999). shoot length and diameter for trees supplemented with water and nutrients were significantly higher in the w++n++ treatment between september 2011 and september 2012. under high soil moisture conditions, there is a general increase in sprouting vigour leading to the transfer and supply of growth promoters such as auxins, cytokinins and gibberellins (ferm & kauppi 1990). this is not the case when there is very low soil moisture (liu & dickmann 1996). consequently, adding water for this study may have led to an overall increase in cell division and resprouting shoots exhibit stronger apical dominance, resulting in higher average shoot diameters and lengths compared to w0n0 trees. it has been proposed that fertiliser addition increases foliar n concentrations as well as chlorophyll levels, leading to an increase in photosynthetic efficiency (scott et al. 2004). in this study, nutrient addition possibly resulted in increased leaf area (ewers et al. 1999), thus increasing the surface area for elevated photosynthetic activity. this may have contributed to an increase in shoot elongation and shoot thickness. results from this study are similar to those obtained by osman and abohassan (2010), when shoot production and resprout shoot length increased after the addition of n and addition of p to rhizophora mangle l. (lovelock et al. 2004). in addition, current findings are also supported by results from the addition of n to thinned p. taeda, which increased resprout diameter and length (sayer et al. 2004). in line with this study’s findings, shoot diameter and height were higher in pinus ponderosa dougl. ex p. laws. and c. laws, which received n addition compared to trees which received no additions (tingey et al. 1996), with both height and diameter growth in fertilised stands doubling in comparison to the non-fertilised stands in norway spruce (picea abies [l.] karst.) (bergh et al. 1999). an overall reduction in the number of shoots for all treatments at the beginning of the wet season in september 2012 implies that there was self-thinning. between september 2011 and september 2012, there was over 50% reduction in the number of shoots in all treatments. while self-thinning could be viewed as a way of reducing high respiration demands associated with high leaf material, this growth pattern was observed even on supplemented trees. findings from this study could possibly mean that resource availability does not necessarily control self-thinning, that is, self-thinning may, in fact, be controlled within the trees and not by environmental factors. effect of water and nutrients on stem total non-structural carbohydrate levels adding water and nutrients had no significant effect on stem tnc levels at both time points. this was surprising given that an increase in soil moisture and nutrient levels through resource additions on the same trees extended leaf survival early into the dry season (moyo et al. 2015), thereby possibly extending the period of photosynthesis for trees supplemented. such an extension of the period of photosynthesis maintained greater leaf area, possibly leading to higher carbon assimilation in supplemented trees. this was in comparison to unsupplemented trees, which had leaf survival that only lasted to the end of the growing season, thereby photosynthesising for a shorter period. given this, the expectation was that these phenological shifts in the dry season (september 2011) extended the photosynthetically active period so that supplemented trees rely less on stored reserves for regrowth compared to unsupplemented trees. because tnc concentrations in tree tissues are considered a measure of carbon storage or a demand for growth (druege et al. 2000; kabeya & sakai 2005; sakai & sakai 1998), the lack of significant effects the addition of water and nutrients may indicate that the carbon demand in resprouting trees, even for supplemented trees which had a longer period to photosynthesise, was exceeding the supply from photosynthesis alone, as stored tnc were mobilised for growth. resprouting trees appear to continuously draw from stored reserves. findings from this study are different to work on q. crispula blume, where root tnc levels were higher after resource addition (kabeya & sakai 2005). similar to findings from this study, other studies also found that n additions had no effect on carbohydrate reserves in beech (fugus sylvatica l.) (pahlsson 1992), while it had little effect on starch levels for chrysanthemum cuttings (druege et al. 2000). in protected areas such as game reserves and conservation areas, a fire can open up patches of land (ward 2005), exposing seeds and seedlings to resources such as light and nutrients. such disturbances, coupled with the fluctuation in rainfall distribution in semi-arid areas (scholes & archer 1997), create bush encroachment patches absent when no fire events or disturbances occur (ward 2005). in agreeing with this study’s findings, this is true when considering the quick recovery of shoot length and diameter in t. sericea after a cutting event. this quick recovery may eventually result in this species encroaching large areas of land, thus reducing the biodiversity and species distribution of the herbaceous vegetation. browsing proportion was the same across all treatments, implying that while browsing may have influenced shoot length and, possibly, the number of shoots produced as well as total biomass produced, the influence was evenly spread across all the treatments. therefore, browsing did not unduly influence the relative differences in the variables measured between treatments. water and nutrients were supplemented through addition immediately adjacent to the tree stump. water addition may have facilitated nutrient uptake, and we assumed that this type of water and nutrient addition enhanced the tree’s ability of fine roots to absorb most of the resources added. the mean annual rainfall in the study area (650 mm per annum) lies between the upper and lower limits of the study species range. considering that t. sericea adapts well to droughts (griffiths 1959), the rainfall in the area is intermediate when compared to the lower limit of the species range. for a tree species adapted to growing in a resource-poor environment, this may mean that the weaker effects of water addition in the second season may have been a result of the study species’ allocation of resources towards storage, instead of growth, as suggested by witkowski, mitchell and stock (1990). such allocation may help explain the low effectiveness of high water addition on coppice regrowth during the second season of growth. conclusion this study demonstrated that resprouting is water and nutrient limited in cut trees and that resource addition does not replenish the stem carbon storage. the results suggest that water is limiting for resprout shoot diameter and length, but not for shoot production and stem storage carbohydrates. results from this study thus support the hypothesis that supplementing cut trees with water and nutrients increases coppice growth, but this effect holds primarily only for the first growing season after cutting, with the effect declining over time. results from a follow-up study also suggest that even though a tree supplemented with water and nutrients may end up photosynthesising for a longer period compared to an unsupplemented tree, this does not necessarily change the tree’s reliance on stored carbon reserves for regrowth early in the growing season. total number of shoots produced displays clear self-thinning and it is interesting to note that water and nutrient additions do not impact this self-thinning. this suggests that self-thinning may be light related and is not positively related with resource availability. future research could focus on how adding water and nutrients changes the balance of tnc in roots and stems, and not only the stem tnc as covered in this current study. cutting of trees at shorter intervals, for example, logging and short coppice rotation can be done in environments that have higher water and nutrient levels, because a quicker recovery of lost biomass is expected in resource-rich environments compared to resource-poor environments (moyo et al. 2015). in an ecosystem where competition for food is high and resources for plant growth are limited, new leaves can be a major source of food at a time when most herbivores struggle to fulfil their nutritional needs, especially at the end of the dry season. therefore, this would ensure that herbivores are assured of a food source, because cattle have been shown to browse more than graze during the dry season (katjiua & ward 2006). in order to reduce the possibility of this study species encroaching large areas, disturbances such as hot fires, which burn and open up large patches, should be controlled. this is because hot fires open up large patches, leading to vigorous resprouting of this species, which may threaten biodiversity in the long run. acknowledgements this research was funded by the mellon foundation and the centre for tree health and biotechnology (fabi, university of pretoria). the assistance of julia reiskind, ed witkowski, sally archibald, ricardo holdo, michelle mack, thabo sibuyi and frank nyathi is appreciated. the wrf is appreciated for making the sites available. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions h.m. was responsible for designing the study, field work and data collection including the analysis and writing up of the paper. m.c.s. guided the first author and read drafts of the paper, while w.t. also read and corrected drafts of the paper as well as formulated the study design. references albaugh, t.j., lee allen, a., dougherty, p.m. & johnesen, k.h., 2004, ‘long term growth responses of loblolly pine to optimal nutrient and water resource availability’, forest ecology and management 192, 3–19. http://dx.doi.org/10.1016/j.foreco.2004.01.002 avohou, t.h., houehounha, r., glele-kakai, r., assogbadjo, a.e. & sinsin, b., 2011, ‘firewood yield and profitability of a traditional daniellia oliveri short-rotation coppice on fallow lands in benin’, biomass and bioenergy 35, 562–571. http://dx.doi.org/10.1016/j.biombioe.2010.10.030 bellingham, p.j. & sparrow, a.d., 2000, ‘resprouting as a life history strategy in woody plant communities’, oikos 89, 409–416. http://dx.doi.org/10.1034/j.1600-0706.2000.890224.x bergh, j., lindera, s., lundmark, t. & elfving, b., 1999, ‘the effect of water and nutrient availability on the productivity of norway spruce in northern and southern sweden’, forest ecology and management 119, 51–62. bond, w.j. & midgley, j.j., 2001, ‘ecology of sprouting in woody plants: the persistence niche’, trends in ecology & evolution 16(1), 45–51. http://dx.doi.org/10.1016/s0169-5347(00)02033-4 bowen, b.j. & pate, j.s., 1992, ‘the significance of root starch in post-fire shoot recovery of the resprouter stirlingia latifoliai r. br. (proteaceae)’, annals of botany 72, 7–16. http://dx.doi.org/10.1006/anbo.1993.1075 castell, c. & terradas, j., 1994, ‘effects of water and nutrient availability on water relations, gas exchange and growth rate of mature plants and resprouts of arbutus unedo l.’, annals of botany 73, 595–602. http://dx.doi.org/10.1006/anbo.1994.1074 chapin iii, f.s., 1991, ‘integrated responses of plants to stress’, bioscience 41(1), 29–36. http://dx.doi.org/10.2307/1311538 cheng, l. & fuchigami, l.h., 2002, ‘growth of young apple trees in relation to reserve nitrogen and carbohydrates’, tree physiology 22, 1297–1303. http://dx.doi.org/10.1093/treephys/22.18.1297 cobb, w.r., will, r.e., daniels, r.f. & jacobson, m.a., 2008, ‘aboveground biomass and nitrogen in four short-rotation woody crop species growing with different water and nutrient availabilities’, forest ecology and management 255, 4032–4039. http://dx.doi.org/10.1016/j.foreco.2008.03.045 coyle, d.r. & coleman, m.d., 2005, ‘forest production responses to irrigation and fertilization are not explained by shifts in allocation’, forest ecology and management 208, 137–152. http://dx.doi.org/10.1016/j.foreco.2004.11.022 cruz, a., perez, b., quintana, j.r. & moreno, j.m., 2002, ‘resprouting in the mediterranean-type shrub erica australis afffected by soil resource availability’, journal of vegetation science 13, 641–650. http://dx.doi.org/10.1658/1100-9233(2002)013[0641:ritmse]2.0.co;2 davis, m., wrage, k.j., reich, p.b., tjoelker, m.g., schaeffer, t. & muermann, c., 1999, ‘survival, growth, and photosynthesis of tree seedlings competing with herbaceous vegetation along a water-light-nitrogen gradient’, plant ecology 145, 341–350. http://dx.doi.org/10.1023/a:1009802211896 dickmann, d.i., nguyen, p.v. & pregitzer, k.s., 1996, ‘effects of irrigation and coppicing on above-ground growth, physiology, and fine-root dynamics of two field-grown hybrid poplar clones’, forest ecology and management 80, 163–174. http://dx.doi.org/10.1016/0378-1127(95)03611-3 druege, u., zerche, s., kadner, r. & ernst, m., 2000, ‘relation between nitrogen status, carbohydrate distribution and subsequent rooting of chrysanthemum cuttings as affected by pre-harvest nitrogen supply and cold-storage’, annals of botany 85, 687–701. http://dx.doi.org/10.1006/anbo.2000.1132 edwards, e.j., downie, a.f. & clingeleffer, p.r., 2011, ‘a simple microplate assay to quantify non-structural carbohydrates of grapevine tissues’, american journal of ecology and viticulture 62(1), 133–137. http://dx.doi.org/10.5344/ajev.2010.10051 el omari, b., aranda, x., verdaguer, d., pascual, g. & fleck, i., 2003, ‘resource remobilization in quercus ilex l. resprouts’, plant and soil 252, 349–357. http://dx.doi.org/10.1023/a:1024792206369 ewers, b.e., oren, r., albaugh, t.j. & dougherty, p.m., 1999, ‘carry-over effects of water and nutrient supply on water use of pinus taeda’, ecological applications 9(2), 513–525. http://dx.doi.org/10.1890/1051-0761(1999)009[0513:coeowa]2.0.co;2 ferm, a. & kauppi, a., 1990, ‘coppicing as a means for increasing hardwood biomass production’, biomass 22(1–4), 107–121. http://dx.doi.org/10.1016/0144-4565(90)90010-h fircks, y.v. & sennerby-forsse, l., 1998, ‘seasonal fluctuations of starch in root and stem tissues of coppiced salix viminalis plants grown under two nitrogen regimes’, tree physiology 18, 243–249. http://dx.doi.org/10.1093/treephys/18.4.243 griffiths, m.e., 1959, ‘a revision of the african species of terminalia’, journal of the linnean society of london botany 25, 364. http://dx.doi.org/10.1111/j.1095-8339.1959.tb00042.x hardesty, l.h., 1984, ‘the challenge of integrated brush management in semi-arid tropics’, rangelands 6(6), 249–253. hoffmann, w.a., schroeder, w. & jackson, r.b., 2002, ‘positive feedbacks of fire, climate, and vegetation and the conversion of tropical savanna’, geophysics research letters 29(22), 2052–2056. http://dx.doi.org/10.1029/2002gl015424 holdo, r.m., holt, r.d. & fryxell, j.m., 2009, ‘grazers, browsers, and fire influence the extent and spatial pattern of tree cover in the serengeti’, ecological applications 19(1), 95–109. http://dx.doi.org/10.1890/07-1954.1 hytonen, j. & kaunisto, s., 1999, ‘effect of fertilization on the biomass production of coppiced mixed birch and willow stands on a cut-away peatland’, biomass and bioenergy 17, 455–469. http://dx.doi.org/10.1016/s0961-9534(99)00061-6 ibrahima, a., mapongmetsem, p.m., bouitang, d. & hassana, b., 2007, ‘regeneration of some fuelwood tree species of humid savanna of adamawa, cameroon: effects of season and cutting height’, ghana journal of science 47, 45–57. kabeya, d. & sakai, s., 2005, ‘the relative importance of carbohydrate and nitrogen for the resprouting ability of quercus crispula seedlings’, annals of botany 96, 479–488. http://dx.doi.org/10.1093/aob/mci200 kabeya, d., sakai, a., matsui, k. & sakai, s., 2003, ‘resprouting ability of quercus crispula seedlings depends on the vegetation cover of their microhabitats’, journal of plant research 116, 207–216. http://dx.doi.org/10.1007/s10265-003-0089-3 kaschula, s., twine, w. & scholes, m., 2005a, ‘coppice harvesting of fuelwood species on a south african common: utilizing scientific and indigenous knowledge in community based natural resource management’, human ecology 33, 387–418. http://dx.doi.org/10.1007/s10745-005-4144-7 kaschula, s.a., twine, w.c. & scholes, m.c., 2005b, ‘the effect of catena position and stump characteristics on the coppice response of three savannah fuelwood species’, environmental conservation 32(1), 76–84. katjiua, m.l.j. & ward, d., 2006, ‘cattle diet selection during the hot-dry season in a semi-arid region of namibia’, african journal of range & forage science 23, 59–67. khan, m.l. & tripathi, r.s., 1986, ‘tree regeneration in a disturbed sub-tropical wet hill forest of north-east india: effect of stump diameter and height on sprouting of four tree species’, forest ecology and management 17, 199–209. http://dx.doi.org/10.1016/0378-1127(86)90112-x kraaij, t. & ward, d., 2006, ‘effects of rain, nitrogen, fire and grazing on tree recruitment and early survival in bush-encroached savanna, south africa’, plant ecology 186(2), 235–246. http://dx.doi.org/10.1007/s11258-006-9125-4 laureysens, i., deraedt, w., indeherberge, t. & ceulemans, r., 2003, ‘population dynamics in a 6-year old coppice culture of poplar. i. clonal differences in stool mortality, shoot dynamics and shoot diameter distribution in relation to biomass production’, biomass and bioenergy 24, 81–95. http://dx.doi.org/10.1016/s0961-9534(02)00105-8 le roux, n.p. & mentis, m.t., 1986, ‘veld compositional response to fertilization in the tall grassveld of natal’, south african journal of plant and soil 3, 1–10. http://dx.doi.org/10.1080/02571862.1986.10634177 liu, z. & dickmann, d.i., 1996, ‘effects of water and nitrogen interaction on net photosynthesis, stomatal conductance, and water-use efficiency in two hybrid poplar clones’, physilogia plantarium 97, 507–512. http://dx.doi.org/10.1111/j.1399-3054.1996.tb00510.x lovelock, c.e., feller, i.c., mckee, k.l., engelbrecht, b.m.j. & ball, m.c., 2004, ‘the effect of nutrient enrichment on growth, photosynthesis and hydraulic conductance of dwarf mangroves in panama’, functional ecology 18, 25–33. http://dx.doi.org/10.1046/j.0269-8463.2004.00805.x mbatha, k.r. & ward, d., 2010, ‘the effects of grazing, fire, nitrogen and water availability on nutritional quality of grass in semi-arid savanna, south africa’, journal of arid environments 74, 1294–1301. http://dx.doi.org/10.1016/j.jaridenv.2010.06.004 moyo, h.p.m., scholes, m.c. & twine, w.c., 2015a, ‘the effects of repeated cutting on coppice response of terminalia sericea’, trees 29(1), 161–169. moyo, h.p.m., scholes, m.c. & twine, w.c., 2015b, ‘the effects of water and nutrient addition on the pehnological response of cut terminalia sericea’, south african journal of botany 96, 85–90. http://dx.doi.org/10.1016/j.sajb.2014.10.009 neke, k.s., 2004, the regeneration ecology of savanna woodlands in relation to human utilisation, university of the witwatersrand, johannesburg. neke, k.s., owen-smith, n. & witkowski, e.t.f., 2006, ‘comparative resprouting response of savanna woody plant species following harvesting: the value of persistence’, forest ecology and management 232(1–3), 114–123. http://dx.doi.org/10.1016/j.foreco.2006.05.051 noumi, z., touzard, b., michalet, r. & chaieb, m., 2010, ‘the effects of browsing on the structure of acacia tortilis (forssk.) hayne ssp. raddiana (savi) brenan along a gradient of water availability in arid zones of tunisia’, journal of arid environments 74(6), 625–631. http://dx.doi.org/10.1016/j.jaridenv.2009.11.007 osman, h.e. & abohassan, a.a., 2010, ‘effect of npk fertilization on growth and dry matter accumulation in mangrove [avicennia marina (forssk) vierh] grown in western saudi arabia’, environmental & arid land agricultural science 21(2), 57–70. http://dx.doi.org/10.4197/met.21-2.5 pahlsson, a.b., 1992, ‘influence of nitrogen fertilization on minerals, carbohydrates, amino acids and phenolic compounds in beech (fugus sylvatica l.) leaves’, tree physiology 10, 93–100. http://dx.doi.org/10.1093/treephys/10.1.93 pote, j., shackleton, c., cocks, m. & lubke, r., 2006, ‘fuelwood harvesting and selection in valley thicket, south africa’, journal of arid environments 67(2), 270–287. http://dx.doi.org/10.1016/j.jaridenv.2006.02.011 rooke, t. & bergstrom, r., 2007, ‘growth, chemical responses and herbivory after simulated leaf browsing in combretum apiculatum’, plant ecology 189, 201–212. http://dx.doi.org/10.1007/s11258-006-9177-5 sakai, a. & sakai, s., 1998, ‘a test for the resource remobilization hypothesis: tree sprouting using carbohydrates from above-ground parts’, annals of botany 82, 213–216. http://dx.doi.org/10.1006/anbo.1998.0672 sankaran, m., ratnam, j. & hanan, n., 2008, ‘woody cover in african savannas: the role of resources, fire and herbivory’, global ecology and biogeography 17(2), 236–245. http://dx.doi.org/10.1111/j.1466-8238.2007.00360.x sayer, s.m.a., goelz, j.c.g., chambers, j.l., tang, z., dean, t.j., haywood, j.d., et al., 2004, ‘long-term trends in loblolly pine productivity and stand characteristics in response to thinning and fertilization in the west gulf region’, forest ecology and management 192, 71–96. http://dx.doi.org/10.1016/j.foreco.2004.01.006 scholes r.j. & archer, s.r., 1997, ‘tree-grass interactions in savannas’, annual review ecological systems 28, 517–544. scogings, p.f. & mopipi, k., 2008, ‘effects of water, grass and n on responses of acacia karroo seedlings to early wet season simulated browsing: leaf n, fibre and tannin concentrations’, journal of arid environments 72(9), 1666–1674. http://dx.doi.org/10.1016/j.jaridenv.2008.03.008 scott, d.a., burger, j.a., kaczmarek, d.j. & kane, m.b., 2004, ‘growth and nutrition response of young sweetgum plantations to repeated nitrogen fertilization on two site types’, biomass and bioenergy 27, 313–325. http://dx.doi.org/10.1016/j.biombioe.2004.02.003 shackleton, c.m., 1997, the prediction of woody plant productivity in the savanna biome, south africa, university of the witwatersrand, johannesburg. shackleton, c.m., 1999, ‘rainfall and topo-edaphic influences on woody community phenology in south african savannas’, global ecology and biogeography 8, 125–136. http://dx.doi.org/10.1046/j.1365-2699.1999.00128.x tilman, d., 1987, ‘secondary succession and the pattern of plant dominance along experimental nitrogen gradients’, ecological monographs 57(3), 189–214. http://dx.doi.org/10.2307/2937080 tingey, d.t., johnson, m.g., phillips, d.l., johnson, d.w. & ball, t.j., 1996, ‘effects of elevated co2 and nitrogen on the synchrony of shoot and root growth in ponderosa pine’, tree physiology 16, 905–914. http://dx.doi.org/10.1093/treephys/16.11-12.905 van wyk, b. & van wyk, p., 1997, field guide to trees of southern africa, struik publishers, cape town. ward, d., 2005, ‘do we understand the causes of bush encroachment in african savannas?’, african journal of range & forage science 22(2), 101–105. http://dx.doi.org/10.2989/10220110509485867 wendler, r. & millard, p., 1996, ‘impacts of water and nitrogen supplies on the physiology, leaf demography and nitrogen dynamics of betula pendula’, tree physiology 16, 153–159. http://dx.doi.org/10.1093/treephys/16.1-2.153 wilson, b., 2002, population dynamics and the regeneration ecology of the african savanna tree burkea africana, university of the witwatersrand, johannesburg. wilson, t.b. & witkowski, e.t.f., 1998, ‘water requirements for germination and early seedling establishment in four african savanna woody plant species’, journal of arid environments 38, 541–550. http://dx.doi.org/10.1006/jare.1998.0362 witkowski, e.t.f., mitchell, d.t. & stock, w.d., 1990, ‘response of a cape fynbos ecosystem to nutrient additions: shoot growth and nutrient contents of a proteoid (leucospermum parile) and ericoid (phylica cephalantha) evergreen shrub’, acta ecologica 11(3), 311–326. abstract introduction methods results discussion conclusion acknowledgements references about the author(s) beanelri b. janecke department of animal, wildlife and grassland sciences, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa jeremy g. bolton bushcam consulting, stonehurst mountain estate, lakeside, cape town, south africa citation janecke, b.b. & bolton, j.g., 2020, ‘variation in mammal diversity and habitat affect heterogeneity and processes of a granite catena’, koedoe 62(2), a1592. https://doi.org/10.4102/koedoe.v62i2.1592 note: special issue: connections between abiotic and biotic components of a granite catena ecosystem in kruger national park, sub-edited by beanelri janecke and johan van tol. original research variation in mammal diversity and habitat affect heterogeneity and processes of a granite catena beanelri b. janecke, jeremy g. bolton received: 13 sept. 2019; accepted: 19 apr. 2020; published: 29 oct. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract a higher variety of habitats normally result in higher diversity of species. the granite catenas near skukuza, kruger national park (knp), consist of different soil types along the hillslope, creating different habitats. objectives were to determine the mammal species present on a catena and surrounding areas; to indicate their main period of activity; and to indicate human visibility in each catenal zone to explain landscape of fear principles. camera trap surveys were conducted for short periods and repeated over three years. in total, 31 mammal species were observed on the catena, and its nearest waterholes. small to mega-sized mammals were present, but some species were only observed during one survey period. small changes were noticed in activity periods between survey periods, probably due to the drought. a severe drought changed vegetation structure and visibility, but the study area appeared to act as a drought forage refuge. the lowest visibility was found at the sodic patch upper-midslope ecotone, and shrub veld. this can possibly explain the lower number of mammal observations in these areas. different habitats and habitat features were described which can affect the presence of mammals, i.e. the mud wallows that were created and maintained by the mammals. future studies can focus on the impact of seasonal changes in mammal presence and on mammal diversity during a normal rainfall year. conservation implications: to understand the mechanisms of herbivores as ecosystem drivers, aspects such as vegetation, soil and mammals should be combined. better understanding of mammals, their habitats and associated processes can lead to better conservation actions. keywords: animal presence; ecosystem functioning; herbivores and predators; landscape of fear; stevenson-hamilton supersite. introduction acceptable habitat conditions are one of the most important requirements for successful maintenance of animal populations. resources in the habitat are exploited differently in order for animals to fulfil their requirements for survival, reproduction and growth (owen-smith 2002). hall, krausman and morrison (1997) define habitat as follows: [t]he resources and conditions present in an area that produce occupancy – including survival and reproduction – by a given organism. habitat is organism-specific; it relates the presence of a species, population or individual (animal or plant) to an area’s physical and biological characteristics. habitat implies more than vegetation or vegetation structure; it is the sum of the specific resources that are needed by organisms. (p. 175) habitat is thus any area where an animal has the necessary resources that allow it to survive, including migration paths, dispersal corridors and areas occupied during breeding times (hall et al. 1997; krausman & morrison 2016). habitat selection refers to a process where an animal selects a specific set of environmental factors that can provide all the essential resources it needs to survive and reproduce (bonyongo 2005; johnson 1980; krausman & morrison 2016). the basic habitat requirements of animals in general are food, water, cover (i.e. thermal cover, shade, shelter, escape cover and cover for new-borns) and space to perform normal daily activities (i.e. resting, feeding, rumination, reproduction, socialising, avoiding competition, etc.) (hansen et al. 2009; janecke 2011). other features that are included when an animal selects a habitat are the vegetation structure of the area, certain geomorphological features, topography, seasonal availability of resources, distance from resources (such as water and grazing), presence of predators and the landscape of fear principle, to name but a few. the specific animal species, their social structure, age, sex, physiological condition (gestation and lactation) and behavioural aspects play a role in the needs of the animal when selecting a habitat (bonyongo 2005; grant et al. 2011; joubert 2016; owen-smith 2002). the presence or absence of an animal species is determined by how well the species’ own special, or general habitat requirements and basic needs are met in that specific area, amongst other reasons, and probably also what its niche role is in maintaining ecosystem processes of the area. a variety of available habitats in an area will usually result in a variety of animals in that area (joubert 2016). it is essential for long-term management and conservation of species to relate the distribution patterns of animals to certain characteristics of their chosen habitat (ben-shahar & skinner 1988; pieterse 2018). this study forms part of a larger multidisciplinary study on the stevenson-hamilton southern granite supersite of the kruger national park (knp), with the main aim of determining specific abiotic processes and some biotic diversity that may play a role in the functioning of the catena ecosystem. weil and brady (2016) define a catena as a soil sequence (soilscape) where each soil type occurs on the same parent material, but in different arrays from the crest down to the footslope. this hillslope facilitates the transfer of solutes, colloids or other particles from upslope areas downhill along an increasing environmental gradient, resulting in a variation of soil properties and associated vegetation in different catenal zones (khomo et al. 2011). the focus of this current study was on whether specific mammal species use certain catenal zones more frequently than others; the specific objectives of this study were to: determine the mammal species present in the study area, at the nearest waterholes and granite outcrops with a snapshot approach indicate the main period of activity of these mammals during three different survey periods establish human visibility (based on density of vegetation) in each catenal zone in order to explain the landscape of fear principle usually connected to the absence or presence of mammals in the area. methods study area the supersite concept was established formally in 2013 to try and focus research effort geographically in knp, and to allow data integration over long periods and across different research themes. one of the four research supersites is located in a specific area on the wetter, southern granite landscape in the stevenson-hamilton area (smit et al. 2013). this study was conducted on a hillslope in the southern granite supersite, located between 25°06’28.6 s, 31°34’41.9 e and 25°06’25.7 s, 31°34’33.7 e, approximately 10 km from skukuza, knp. the study area falls in the renosterkoppies land type, described as an ecotonal area between land types that are associated with the sabie river catchment (smit et al. 2013). the average herbivore biomass on the supersite is 2.1 kg grazers/ha, 3.0 kg browsers/ha and 9.9 kg mixed feeders/ha (smit et al. 2013). this study focused on one catena or hillslope from the crest to third-order watercourse or drainage line. the vegetation in this area is described as moderately dense bush savanna on midslopes and as shrub savanna with a dense riverine forest at the footslope and valley bottom (smit et al. 2013). as part of the effort to focus different research fields together on the supersites, the following research was conducted on one specific study area: bouwer, le roux and van tol (2020) described the soil types and properties, theron, van aardt and du preez (2020) did a detailed vegetation classification and janecke (2020) provided a description of the vegetation structure. four catenal zones were identified along the slope based on their position on the catena and associated vegetation, namely, crest and upper-midslope, lower midslope or sodic patch, footslope shrub veld and a riparian area around the dry drainage line. a seepline defines the transition between the upper and lower-midslope. to give an indication of the size of each zone, a linear measurement was done from crest to drainage line (but that does not indicate the surface area of each zone): upper-midslope is 140 m in width, lower midslope is 220 m, footslope is 70 m (on its shortest end) and the riparian area is roughly 50 m but more difficult to indicate because of the winding of the drainage line. the study area was approximately 1.5 km in length. a visual illustration of the catena studied is provided by janecke et al. (2020). camera trapping the location of each camera trap was determined after a visual survey of the study area in order to represent the four identified catenal zones. the three nearest permanent waterholes (de la porte, kwaggaspan and renosterkoppiesdam) and two rocky outcrops with granite boulders close to the study area, as well as two large mud wallows inside the study area (sodic patch and riparian zone), were also surveyed through cameras. in total, 30 camera traps were deployed during each of the three survey periods, which lasted for approximately a fortnight each during september 2015, march 2016 and march–april 2017. of the 30 cameras used, 23 were bushnell models, 4 were cuddeback, 2 were scoutguard and 1 was little acorn model. cameras were selected with similar attributes in terms of trigger response time (between 0.25 s and 0.9 s) and field of view ranges, and all utilised infrared flashes. a camera trap positioned to monitor a game trail was required to have a faster trigger response time than one monitoring a wide clearing. the same locations were used in all three surveys unless the camera-holding tree had been damaged or vegetation conditions dictated that a better (nearby) location was essential to obtain good results. the majority of cameras were set up to monitor mammals passing at close range, while four were set up to monitor on a time-lapse basis from a distance as follows: passing mammals (26 cameras). cameras were positioned between 50 cm and 170 cm height to give the best uninterrupted view of the area around a well-used game trail, termite mound or an open clearing where there was evidence of the presence of mammals. where possible, cameras were orientated in a southerly direction to optimise the quality of images obtained (sunlight). cameras were installed as low as possible, provided that the camera’s field of view was not obstructed by vegetation, to also capture small mammals. cameras were programmed to take two photographs per trigger event, with a rest interval of 5 s between image pairs. time lapse (four cameras). these cameras were installed between 1.5 m and 3 m height so as to give a clear view of the entire waterhole or sodic patch and were programmed to take one image every 5 min. this was deemed an appropriate interval to ensure that any mammal drinking at the waterhole or crossing the sodic patch would likely be recorded. in addition to the camera taking time lapse images, it was also programmed to record the movement of mammals passing the camera at close range. visibility during the vegetation sampling of the study area in 2015, a visibility index was determined. the distance of 20 m was used after testing different possible distances – in many of the plant communities, the vegetation was too dense to see much further than 20 m or 30 m. a 100-m measuring tape was used as a transect line in the different catenal zones. transects were placed on a horizontal plane in each zone (in other words, not running from top to bottom down the slope) at approximately 20 m intervals but excluding the transition areas (janecke 2020). a 1 m2 cloth with a checkerboard pattern of 10 cm blocks and dowel sticks on the edges was used. the method was slightly adjusted from that of bissett and bernard (2007). the field assistant held this apparatus at ground level, 20 m from the observer and at 90°, 180° and 270° angles, respectively, from the observer standing on the transect line. the observer then counted the number of blocks visible at an observing height of 70 cm and 170 cm from the ground level to imitate different sized animals. these observations were repeated at 20 m intervals along the 100 m transect line (0 m, 20 m, 40 m, 60 m, 80 m and 100 m) for each catenal zone. the average in the bar graph illustrating the results was calculated from all the values on the 100 m line (20 m intervals) for each angle (90°, 180° and 270°) observed, respectively. data and statistical analyses camera trapping images were analysed as follows. for the purpose of this study, an independent trigger event is defined as an image or a series of images that record the passing of an individual or group of individuals at a specific time. for example, an event could include the movement of a herd of impala, or a single hyena, passing the camera. the number of events thus represents the number of observations of an animal species. no specific interval was used to determine the independence of consecutive events. however, every effort was made to determine whether the individual or individuals were different from those in preceding sequences of images to prevent double-counting. if an animal species could not be identified, for example, because of badly blurred photos usually taken at night, it was indicated as unknown. from these, the following data per independent trigger event were extracted: date and time, animal species, number of individuals in the view of the camera and the primary activity of the animals. data from the camera traps were grouped according to the catenal zone that the camera was located in, with all three waterholes combined together as one zone. there were five cameras per catenal zone, five at waterholes and surrounds (the two larger waterholes had two cameras each pointing in different directions), and five at special locations (granite outcrops and mud wallows). the number of trigger events (or observations of each species) was totalled for each mammal species per catenal zone and graphically presented as a grayscale gradient of increasing number of events for each survey period (year) of research. a pareto graph was used to indicate the total number of events of that species in the study area in descending order of frequency and its percentage of the total as a cumulative line. only mammal species were indicated in the results, but all information on the two rhinoceros species was excluded (for security reasons). the minimum to maximum range in general group size of individuals visible in the cameras’ view and the median value of typical group sizes observed were tabulated per survey period. the excel median function was used to determine a median value of those records of group sizes greater than 1. this was verified by a visual observation of the data to ensure that the median value looked reasonable. the sum total of all mammals observed per species in each survey period was also indicated in a table (this is the total number of all individuals captured on the cameras and not the number of times/events that they were observed). other information calculated from photographs included the percentage of the total number of events that a species was observed on the cameras to be active (grouped into day, night and civil twilight) and species that utilised the catena was also indicated. utilisation of the catena was described as mammals captured on the cameras doing one or more of these activities: feeding: obvious feeding activity, or a species moving slowly across the camera’s field of view in a feeding position, for example, head down for grazers and head near vegetation for browsers. socialising: chasing each other, making body contact, locking horns. resting: lying down for long periods, usually at night for diurnal animals. wallowing: lying in the mud holes on the catena and a mammal next to the wallow that showed mud on its body were included. drinking: some animals were drinking from the mud wallows, but mostly mammals standing at the edge of a waterhole were presumed to be drinking. if mammals were just in the vicinity, although it can be assumed that they might have been drinking, these mammals are not listed in table 3 (but are indicated in figure 1). figure 1: number of events (observations) of each mammal species through camera trapping in the southern granite supersite of kruger national park. cameras were set up to represent each catenal zone from crest to drainage line, while waterholes and granite hills in the immediate vicinity of the catena were also covered. results are indicated in class ranges in each catenal zone during the different survey periods. species are grouped in the figure based on their feeding habits: other, herbivores, carnivores and the last seven species only occurred in surrounding areas but not on the catena itself. the normal distribution of data was determined by the shapiro–wilk test. as it was not normally distributed, the kruskal–wallis test (https://www.socscistatistics.com/) for non-parametric data was used to determine significant differences in number of animal species observations per catenal zone between the three different survey periods. a 5% level of significance was used. variability in sample means of the number of blocks visible on the checkerboard was measured through standard errors. error bars were included for standard error of the mean (sem) in the visibility graph and were calculated by excel using the following formula: sem = standard deviation/square root of total number of samples. ethical considerations ethical approval for the multidisciplinary project as a whole, with specific mention to the camera trapping, was obtained from the interfaculty animal ethics committee at the university of the free state (ufs-aed2019/0121). results a snapshot approach was followed to give an idea of the mammal species that frequent the study area – in other words, the cameras were only left in the veld for short periods during each survey period. a total of 31 mammal species were observed during the study (table 1). tortoises, terrapins, (large) millipedes and several bird species (including ground birds and smaller birds) were also observed on the cameras but are not included in the species list which focuses only on mammals. all data on the black and white rhinoceroses have been removed from the figures because of the sensitive nature of this data (i.e. poaching concerns). although exclusion of these two threatened species affects the results, we decided to remove such data as a precaution. table 1: common and scientific names of all mammals observed in the study area by means of camera trapping. figure 1 indicates the events (or in other words, the number of independent observations) of each mammal species observed during the three survey periods, divided into catenal zones (determined from its position on the catena and associated vegetation). herbivores (16 species), carnivores (11 species), insectivores (2 species: aardvark and civet) and primates (2 species) were recorded. of these, seven species are small mammals, such as the mongoose species, genet, scrub hare and tree squirrel. figure 2 presents the total number of events that each species was observed and its percentage of the total number of events (including all species). this graph clearly shows the species that were observed more frequently than others, in contrast to species that were recorded the least or were completely absent during all the survey periods. the order of species also changes through the survey periods (figure 2). the more common species during all three survey periods were impala (603 events in total), elephant (289), spotted hyena (74), kudu (76), giraffe (72), buffalo (54), zebra (51), warthog (48) and grey duiker (43). blue wildebeest (28 events in total), lion and steenbok (approximately 20 events each) were also observed more than the remainder of the species. figure 2: pareto chart indicating the decreasing frequency of occurrence (number of observation events) of mammal species during each survey period ([a] 2015, [b] 2016 and [c] 2017) on the left axis (bars) and the cumulative percentage of the total number of occurrences on the right axis (line). the general group sizes of each species observed are shown in table 2. it is noted that some individuals may have passed outside the view of the cameras; thus, the totals are just an indication and not the true possible numbers of herds or groups in the area. the sum total of all animals of each species that were observed during a survey period (table 2) clearly shows the following: the rise and fall of total elephant numbers observed during the three surveys, the increasing total number of impalas observed and the declining total number of kudu recorded over the three periods, to mention but a few. these totals are not herd sizes but all the individuals in total observed during the survey period, which can be the same individuals seen repeatedly. if these totals are compared to herd sizes and the number of events (table 2), it may be interpreted that individuals of these specific species are probably resident in the study area and will therefore utilise the area more than other species just passing through the area and are not observed so frequently. table 2: different numbers of mammals observed during each survey period and the percentage of events that they were active at a specific period during a 24-h day. group size range (minimum and maximum numbers recorded during the events) and the median value of group sizes (excluding solitary individuals from calculation), as well as the sum totals of all individuals of each mammal species observed, are summarised. if no active period is indicated, the animal was not observed in that year’s camera trapping survey. the total number of mammal species observed during each survey period (24, 25 and 24 species, respectively) seem very similar, but they include some species that were only observed during one survey period but were absent from other periods (figure 1). this was confirmed by significant statistical differences between the number of observations of each animal species in a catenal zone between the three different survey periods (n = 122 per survey period; h = 7.295; p = 0.026 < 0.05). the total number of mammal species in each catenal zone was as follows: crest and upper-midslope (19); sodic patch on lower midslope (17); shrub veld on footslope (14); riparian area close to the drainage line (15); permanent waterholes (22); rocky hills (16); and mud wallows, temporarily filled with water (13). many of these mammal species were present in more than one catenal zone. the highest species diversity was found on the upper-midslope and sodic patch, and as was expected, at the waterholes (figure 1). the period of a 24-h day when the mammals were observed on the cameras is indicated in table 2. the obvious assumption was made that animals engaging in activities such as feeding, resting, wallowing, drinking and socialising were spending more time in the study area than animals just passing the cameras. these results may differ seasonally, but there is insufficient data from this study to confirm that, and it is possible that more mammal species may utilise the area than that found from the short duration of camera surveys (table 3). the duiker was included, as they are territorial and not water-dependent animals (meaning that their presence indicates utilisation of the area and not just moving through the area to perhaps reach waterholes), and the civet was included because of the presence of active civetories (i.e. locations of frequent civet defecation) on the catena. table 3: these mammals were observed on the cameras to utilise the study area and surrounds in the day, night or civil twilight (twi’t) during each survey period by feeding, resting, socialising, wallowing or drinking. it was uncertain if hippopotamus and aardvark utilised the area. many animals just seemed to be passing through the granite catena, not utilising the area, and therefore their activity is not indicated in the table. figure 3 indicates the human visibility (average of six distances in one direction) in the different directions (or angles from the transect line) in each catenal zone surveyed. because of the vegetation structure, the sodic patch had the highest average visibility, while the transition zone between the upper-midslope and sodic patch had the lowest average visibility, followed by the shrub veld on the footslope. there were differences observed in figure 3 between the 70 cm and 170 cm observation heights, especially in the riparian zone (t7). tall, dense grasses, clumps of shrubs, large trees and the presence of the drainage line (with its relatively higher banks than the surrounding area) contributed to lowered visibility in this zone. figure 3: average number of blocks visible on a 1 m2 checkerboard in each transect at three angles to the transect line (90°, 180° and 270°). standard error bars are also indicated. observations were performed at 20 m intervals with a line distance of 100 m and the average number of blocks visible per line is indicated. two observation heights were used – the observer sitting on the ground (eye level at 70 cm) and the observer standing (eye level at 170 cm). during the initial vegetation sampling as part of the large multidisciplinary project, the veld was still in relatively good condition and the vegetation was dense with long grass, especially in the shrub veld and the riparian areas. this changed during 2016, with the extreme drought, not only for the study area, but many other areas in the southern and central knp were also similarly affected (malherbe et al. 2020). the visibility changed to an estimated 80% – 100% in different zones of the study area (pers. obs. october 2016) – the grasses that were present were only small tufts grazed down to stubble height. the trees were mostly bare or sparsely leafed and many trees died or were uprooted and damaged by elephants. after the drought in 2016, many of the tall grasses died and were lying flat on the ground (covering the soil – see janecke 2020). all of this greatly increased the visibility in that area during the last two years of camera data collection. discussion mammals are usually present in a habitat if their needs and requirements are met or if they move between preferred habitats. with regards to the usual terrestrial mammal habitat requirements, the assumption can be made that the basic needs are most probably accounted for in the study area and surrounds, based on the presence of the listed mammals (figure 2) as indicated by the definition of habitat (hall et al. 1997; krausman & morrison 2016). there are various reasons why mammal species will be present in or absent from a specific area or catenal zone (see introduction) and some of these reasons will be focussed on in this section. food, water, cover (shade, shelter, etc.), space and certain geomorphological characteristics (soil types, topography, geological formations, etc.) are considered to be the basic habitat requirements of most mammals. the mediumto large-sized individual trees (with canopy cover ranging from 10.5% to 33.6%) present on the upper-midslope and in the lower-lying riparian area (janecke 2020), together with scattered clumps of shrubs and trees, should provide food for herbivores and sufficient shade and shelter from the extreme temperatures associated with savannas. the characteristic granite boulders on small hills or outcrops (geomorphology) in the surrounding area had their own mammal species associated with that habitat which mostly differed from that of the catena (figure 1). the nearest drinking water is available to animals at the sabie river and some of its smaller tributaries, and at three nearby permanent waterholes located in different directions from the study area. the rationale with placing camera traps at the three permanent waterholes was that mammals that were not observed in the study area but were caught at the waterholes are actually present in the local area and might also use the study area from time to time as habitat. waterbuck, side-striped jackal and banded mongoose were only found at the waterholes and not on the catena or granite hills (figure 1). there were three large mud wallows in the study area that might have been created by mammals looking for water. it might fill up with rain water, surface run-off, and there is a possibility that water seepage from underground may also play a role (depending on the position of the mud wallow on the catena). many animals were also observed to drink at these temporary holes when water was present. water sources are generally crucial components in habitat selection by many water-dependent herbivores, and also for predators like lions. habitat selection of most herbivores is also influenced by distance to water; hence, predators will have a greater chance of encountering prey in the areas closer to water (davidson et al. 2012; gaylard et al. 2003). the temporary mud wallows contained rain water during the 2015 and 2016 survey periods, but during 2017 no water was present. this might have affected the presence of species on the catena and the number of individuals present during the 2017 observation period. the group numbers of elephant, buffalo, warthog and blue wildebeest were lower in this period (table 2), possibly as a result of the absence of water in the temporary holes, and also possibly because of obvious food shortages caused by the drought (2015–2017). mongoose was not observed during 2017, while other smaller-sized mammals like scrub hare, vervet monkey, duiker and steenbok had the lowest number of events (observations). this can be because of various reasons, such as food limitations, the absence of cover (higher visibility to predators) and high temperatures, to name but a few (see seydack et al. 2012). the number of observations of kudu and giraffe did not change drastically over the survey periods, although it was only a few individuals observed at a time (table 2). during the extreme el niño event of 2015–2017, the study area appeared to be a refuge island of green vegetation surrounded by the desert-like areas around skukuza and lower sabie where mainly bare soil, leafless trees and dead trees were present during the drought (janecke et al. 2020; van aardt et al. 2020). the long-term (76 years) average annual rainfall total for skukuza is 550.4 mm. during the drought, the cumulative total of rainfall measured at the end of the climatic year (june 2015–july 2016) for the skukuza area was 194 mm (skukuza scientific services weather data). this is lower than the lowest annual rainfall on record (251 mm during 1990–1991 – zambatis & biggs 1995) for that area. staver, wigley-coetsee and botha (2019) concluded that the droughts of 1991–1992 and 2015–2017 were exceptionally severe and that it probably exceeded the tolerance of the grasses. the impact of browsing or grazing across a landscape is differentially affected by the dispersion of herbivores as an agent of heterogeneity, while further controls include the social structure of these animals and their density. dispersion (and thus the presence or absence in the study area) may also be controlled by the degree of the drought (pickett et al. 2003). there was still grass available on the catena (although it was grazed to stubble height on most of the sodic patch, but not on the upper-hillslope) and the trees on the catena were alive and full of leaves (pers. obs. april 2016 and visible on the cameras). this can provide an explanation for the presence of the herbivores in the study area at the peak of the drought (figure 1), where they appeared to be mostly absent from surrounding areas closer to skukuza. hippos were captured on the cameras at the de la porte and renosterkoppies waterholes and in the dry drainage line of the study area during 2016, but not during other survey periods (figure 1). as expected, they seemed to have moved further away from the river in search of food (and possibly other large waterbodies for cooling their body temperature) during the drought period (table 3). elephants were observed during the day in 87% of events in 2015, 66% in 2016 and 52% in 2017 (table 2); giraffe were observed during the day for 73%, 69% and 53% of each of the three survey periods; kudu for 85%, 100% and 55%; and zebra for 86%, 44% and 56% of the survey periods. these changes in their active period might be a way to deal with intensifying heat during the day as the drought progressed, by being more active in cooler periods of the night, or it might be that as more rains were experienced from 2017, the food resources were less scarce and animals could spend less time on feeding and moving in the hot period of the day; however, more long-term research is needed to confirm any of these statements for the catena. white rhino and hippo grazing, together with continuous grazing from other species, and the trampling effect on the soil are important contributors to maintaining the short grass state on grazing lawns. active dung middens also contribute to the nutrient value of a local area. high-quality forage that is seasonally available on sodic sites may have important consequences for population dynamics and the behaviour of grazers (grant & scholes 2006; jacobs & naiman 2008; khomo & rogers 2005). du toit (2003) stated that grazers concentrate on zones that shift up and down a catenary drainage gradient on a seasonal basis, and they move progressively down the hillslope in the dry season as green grass declines, while they switch back to nutritional swards on the uplands as rains commence. the same pattern is described for browsers, usually resulting in the higher herbivory in riparian areas during the dry season (du toit 2003). in the current study, the riparian areas had lower average visibility, and these areas were not frequented by mammals more than other catenal zones at the end of the growing season when the surveys were conducted. visibility might affect the presence of herbivores in some of the catenal zones, or parts thereof, because of predation risk and the landscape of fear principles. the presence of carnivores is also affected by visibility in the area because smaller carnivores, as possible prey for larger carnivores, might also experience a landscape of fear in certain areas. terrain heterogeneity differs over landscapes and a predator is not adapted to hunt skilfully in all types of landscapes. a system can be conceived where the lethality of a predator and predation risk will vary with spatial changes in different habitats. this is described as the landscape of fear: ‘a three dimensional landscape whose peaks and valleys are defined by the level of predation risk related to changes in habitat as they affect the lethality of the predator’ (laundre, hernandez & ripple 2010:2). the landscape of fear thus reflects the level of fear of predation that a prey will experience in various parts of the area it uses as habitat (laundre et al. 2010). specific areas of the riparian zone proved to have the lowest overall visibility of all the catenal zones, especially for smaller-sized mammals (everything smaller than, and including, a dwarf antelope), because of tall grasses and dense shrubs. although camera traps were specifically placed on footpaths and open areas where more mammal activity was expected, the lower recorded visibility score (figure 3) might explain why relatively few observations of mammals were made in the shrub veld and riparian zones (figure 1). the percentage of the total number of events (observations) in each catenal zone was recorded as follows: 13.3% on the crest and upper-midslope, 19.1% on the sodic patch, 9.0% in the shrub veld on the footslope, 10.7% in the riparian area, 10.5% at mud wallows and the remainder at waterholes (25.2%) and surrounding areas (12.2%). however, these percentages are slightly biased because of different capturing techniques used by the cameras and because of impala that ‘camped out’ in vicinity of some cameras while feeding and ruminating – increasing the number of events on a single day but spaced out with longer time periods in between (see the definition of ‘event’ under ‘methods’ section). lions and leopards are stalk and ambush predators and are expected to be more successful where they can use dense vegetation as concealment for hunting. lions in hwange national park, zimbabwe, preferred to be located in denser vegetation, next to more open, bushed grassland areas, from where they can observe prey inside the bushed grasslands (davidson et al. 2012). trees and bushes are also used by predators when they approach prey, which to a certain extent might counteract a tree’s attractiveness to herbivores for feeding and/or resting. furthermore, areas with high densities in trees, especially trees with low branching heights, are known to be less attractive because of impeded predator visibility (riginos & grace 2008; treydte, riginos & jeltsch 2010). in the study area, leopards were observed on the upper-midslope and in the riparian area, while lions were observed on the upper-midslope, sodic patch and riparian area of the catena (figure 1). they may have similar tendencies in using the denser areas around the sodic patch to observe potential prey, as recorded in the literature, but this assumption has not been researched in the current study and these predators were also observed in other catenal zones. the vegetation structure and density, together with other factors such as the presence of termite mounds and the higher banks of the drainage line, made visibility poor in certain zones on the catena. this can create a top-down regulation by predators (or their possible presence anticipated by prey) on herbivore numbers and species in these areas, based on landscape of fear principles (see laundre et al. 2010), especially in riparian areas known to usually have higher grazing value (du toit 2003; naiman & rogers 1997). lower observations than expected were made of herbivores in the riparian zone (figure 1) and only the more open areas were used by animals that were present. bottom-up controls (vegetation structure, cover and food availability) can also play an equally important role in the presence of mammals. there may be feedbacks present between mammals and vegetation heterogeneity, for example, herbivores selecting open areas in order to reduce exposure to predation risk, which results in them maintaining these areas as open through their grazing and trampling effects (i.e. a positive feedback loop). factors that play an important role in controlling species richness on sodic areas specifically are biomass consumption, trampling or hoof action and other activities. to quantify these relationships will enhance the understanding of the mechanistics of herbivores as ecosystem drivers, and also of riparian areas at the bottom of a catena as refuges for biodiversity and as nutrient sinks (jacobs & naiman 2008). localised patches known to have higher nutrient value (sub-canopy habitats, termite mounds and sodic patch), as well as permanent (waterholes) and ephemeral (mud wallows) water sources, were used by herbivores and can also explain their presence in the study area. this article aimed to provide a basic list of mammals present on the catena that might have a role in the functioning and interactions of this heterogeneous ecosystem. a general description was also provided of habitat and habitat features that determine the presence of mammals on the catena. there is scope for future studies on the southern granite supersite to expand on this knowledge and information. conclusion there is nothing obvious that can prohibit movement of animals through the study area, and considering the park has a large surface area catering for various species, the possibility is high that some mammal species, especially the smaller mammals, were not listed. some species might be present for short periods, or during specific seasons, increasing the heterogeneity of the larger supersite even more than what was found. furthermore, camera traps do not monitor the entire site, which will further contribute towards under-reporting or entirely missing lower density and transient species. this study at least provided a basic list of 31 mammal species present in the southern granite supersite, and some measure of relative abundance. the more common species were found in almost all catenal zones; thus, the question about certain mammal species that might use specific zones more often could not be answered with this study. distances between the different catenal zones were relatively short, which made it more difficult to differentiate between utilisation of zones separately. large and small predators, as well as herbivores of all sizes, were present in the study area, while poor visibility in some areas might explain the absence or lower presence of mammals based on the landscape of fear principles and/or poor detectability. small changes were also observed in activity periods of specific species between different years of surveys, probably in reaction to the drought. differences in species presence between seasons might be because of animals’ migration, or localised movement patterns. the extreme drought has changed the vegetation structure and availability of food, which had a pronounced effect on the presence and movement of animals (e.g. hippo appearing at the site during the drought as it grazed further from perennial rivers), but this aspect was not studied specifically. animals move to areas where their needs are best met, some will move locally in search of food or water, but others might be bound to strict territorial boundaries that restrict their movement to local areas. future studies can focus on longer periods of data collection and exploring the possible differences in the presence of mammals between seasons. it would also be interesting to study any possible changes in the presence of mammal species in a normal rainfall year after the veld recovered from the impact of the severe 2015–2016 drought. acknowledgements the following people and institutions are acknowledged: the ufs strategic research fund, as well as the nrf thuthuka grant for funding the research; martin tinneveld for assistance with the checkerboard; dr izak smit from sanparks for his time, guidance and valuable comments on this manuscript; the staff of sanparks scientific services for administrative arrangements; the game guards who kept us safe during field work; dr tascha vos for her time, patience and effort with figure and table formatting; and erneli steyn for assistance with data processing and literature searches. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions both authors were co-responsible for experimental layout, camera trapping, data gathering, data processing and interpretation. b.b.j. was responsible for writing the majority of the article and creating the figures. j.g.b. processed the camera trap data and compiled the tables. funding information the university of the free state (ufs) strategic research fund largely funded the multidisciplinary project as a whole, including this part of the study; and the national research foundation thuthuka grant also partially funded this research. data availability study data are available and may be provided, on request, by the corresponding author. data from all research done within kruger national park is placed within the sanparks repository (not for free, open access). disclaimer the views and opinions expressed in this article are the authors’ own and do not necessarily reflect the official policy or position of the institution or funder. references ben-shahar, r. & skinner, j.d., 1988, ‘habitat preferences of african ungulates derived by uniand multivariate analyses’, ecology 69(5), 1479–1485. https://doi.org/10.2307/1941645 bissett, c. & bernard, r.t.f., 2007, ‘habitat selection and feeding ecology of the cheetah (acinonyx jubatus) in thicket vegetation: is the cheetah a savanna specialist?’, journal of zoology 271(3), 310–317. https://doi.org/10.1111/j.1469-7998.2006.00217.x bonyongo, c.m., 2005, ‘habitat utilization by impala (aepyceros melampus) in the okavango delta’, botswana notes and records 37, 227–235. bouwer, d., le roux, p.a.l. & van tol, j., 2020, ‘identification of hydropedological flowpaths in stevenson-hamilton catena from soil morphological, chemical and hydraulic properties’, koedoe 62(2), a1584. https://doi.org/10.4102/koedoe.v62i2.1584 davidson, z., valeix, m., loveridge, a.j., hunt, j.e., johnson, p.j., madzikanda, h., et al., 2012, ‘environmental determinants of habitat and kill site selection in a large carnivore: scale matters’, journal of mammalogy 93(3), 677–685. https://doi.org/10.1644/10-mamm-a-424.1 du toit, j.t., 2003, ‘large herbivores and savanna heterogeneity’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 292–309, island press, washington, wa. gaylard, a., owen-smith, n. & redfern, j., 2003, ‘surface water availability: implications for heterogeneity and ecosystem processes’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 171–188, island press, washington, wa. grant, c.c. & scholes, m.c., 2006, ‘the importance of nutrient hot-spots in the conservation and management of large wild mammalian herbivores in semi-arid savannas’, biological conservation 130(3), 426–437. https://doi.org/10.1016/j.biocon.2006.01.004 grant, r.c.c., peel, m.j.s., bezuidenhout, h. & grant, r., 2011, ‘evaluating herbivore management outcomes and associated vegetation impacts’, koedoe 53(2), 116–130. https://doi.org/10.4102/koedoe.v53i2.1008 hall, l.s., krausman, p.r. & morrison, m.l., 1997, ‘the habitat concept and a plea for standard terminology’, wildlife society bulletin 25(1), 173–182. hansen, b.b., herfindal, i., aanes, r., sæther, b.e. & henriksen, s., 2009, ‘functional response in habitat selection and the tradeoffs between foraging niche components in a large herbivore’, oikos 118(6), 859–872. https://doi.org/10.1111/j.1600-0706.2009.17098.x jacobs, s.m. & naiman, r.j., 2008, ‘large african herbivores decrease herbaceous plant biomass while increasing plant species richness in a semi-arid savanna toposequence’, journal of arid environments 72(6), 891–903. https://doi.org/10.1016/j.jaridenv.2007.11.015 janecke, b.b., 2020, ‘vegetation structure and spatial heterogeneity in the granite supersite, kruger national park’, koedoe 62(2), a1591. https://doi.org/10.4102/koedoe.v62i2.1591 janecke, b.b., 2011, ‘a plant based study of the feeding ecology of introduced herbivore game species in the central free state’, phd thesis, department of animal, wildlife and grassland sciences, university of the free state, bloemfontein. janecke, b.b., van tol, j., smit, i.p.j., van aardt, a.c., riddell, e.s., seaman, m.t. et al., 2020, ‘biotic and abiotic connections on a granitic catena: framework for multidisciplinary research’, koedoe 62(2), a1600. https://doi.org/10.4102/koedoe.v62i2.1600 johnson, d.h., 1980, ‘the comparison of usage and availability measurements for evaluating resource preference’, ecology 61(1), 65–71. https://doi.org/10.2307/1937156 joubert, s.c.j., 2016, ‘animal behaviour’, in j.d.p. bothma & j.t. du toit (eds.), game ranch management, pp. 385–392, van schaik publishers, pretoria. khomo, l., hartshorn, a.s., rogers, k.h. & chadwick, o.a., 2011, ‘impact of rainfall and topography on the distribution of clays and major cations in granitic catenas of southern africa’, catena 87(1), 119–128. https://doi.org/10.1016/j.catena.2011.05.017 khomo, l.m. & rogers, k.h., 2005, ‘proposed mechanism for the origin of sodic patches in kruger national park, south africa’, african journal of ecology 43(1), 29–34. https://doi.org/10.1111/j.1365-2028.2004.00532.x krausman, p.r. & morrison, m.l., 2016, ‘another plea for standard terminology: editor’s message’, the journal of wildlife management 80(7), 1143–1144. https://doi.org/10.1002/jwmg.21121 laundre, j.w., hernandez, l. & ripple, w.j., 2010, ‘the landscape of fear: ecological implications of being afraid’, the open ecology journal 3(3), 1–7. https://doi.org/10.2174/1874213001003030001 malherbe, j., smit, i., wessels, k. & beukes, p., 2020, ‘recent droughts in the kruger national park as reflected in the extreme climate index’, african journal of range and forage science, 37(1), 1–17. naiman, r.j. & rogers, k.h., 1997, ‘large animals and system-level characteristics in river corridors’, bioscience 47(8), 521–529. https://doi.org/10.2307/1313120 owen-smith, r.n., 2002, adaptive herbivore ecology from resources to populations in variable environments, cambridge university press, cambridge. pickett, s.t.a., cadenasso, m.l. & benning, t.l., 2003, ‘biotic and abiotic variability as key determinants of savanna heterogeneity at multiple spatiotemporal scales’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 22–40, island press, washington, wa. pieterse, m., 2018, ‘the ecology of impala (aepyceros melampus) in a dystrophic system: a case study from welgevonden game reserve, limpopo province, south africa’, msc dissertation, school of natural resource management, nelson mandela metropolitan university, george. riginos, c. & grace, j.b., 2008, ‘savanna tree density, herbivores, and the herbaceous community: bottom-up vs top-down effects’, ecology 89(8), 2228–2238. https://doi.org/10.1890/07-1250.1 seydack, a.h., grant, c.c., smit, i.p., vermeulen, w.j., baard, j. & zambatis, n., 2012, ‘climate and vegetation in a semi-arid savanna: development of a climate–vegetation response model linking plant metabolic performance to climate and the effects on forage availability for large herbivores’, koedoe 54(1), art. #1046, 12 pages. http://dx.doi.org/10.4102/koedoe.v54i1.1046 smit, i.p.j., riddell, e.s., cullum, c. & petersen, r., 2013, ‘kruger national park research supersites: establishing long-term research sites for cross-disciplinary, multiscaled learning’, koedoe 55(1), art. #1107, 7 pages. http://dx.doi.org/10.4102/koedoe.v55i1.1107 staver, a.c., wigley-coetsee, c. & botha, j., 2019, ‘grazer movements exacerbate grass declines during drought in an african savanna’, journal of ecology 107(3), 1482–1491. https://doi.org/10.1111/1365-2745.13106 theron, e.j., van aardt, a.c. & du preez, p.j., 2020, ‘vegetation distribution along a granite catena, southern kruger national park’, koedoe 62(2), a1588. https://doi.org/10.4102/koedoe.v62i2.1588 treydte, a.c., riginos, c. & jeltsch, f., 2010, ‘enhanced use of beneath-canopy vegetation by grazing ungulates in african savannahs’, journal of arid environments 74(12), 1597–1603. https://doi.org/10.1016/j.jaridenv.2010.07.003 van aardt, a.c., codron, d., theron, e. & du preez, p.j., 2020, ‘plant community structure and vegetation changes after drought on a granite catena in the kruger national park, south africa’, koedoe 62(2), a1585. https://doi.org/10.4102/koedoe.v62i2.1585 weil, r.r. & brady, n.c., 2016, the nature and properties of soils, 15th edn., pearson education limited, harlow. zambatis, n. & biggs, h.c., 1995, ‘rainfall and temperatures during the 1991/92 drought in the kruger national park’, koedoe 38(1), 1–16. https://doi.org/10.4102/koedoe.v38i1.301 abstract introduction study area methods results and discussion conclusion acknowledgements references about the author(s) andri c. van aardt department of plant sciences, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa daryl codron department of zoology and entomology, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa ettienne j. theron department of plant sciences, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa pieter j. du preez † department of plant sciences, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa citation van aardt, a.c., codron, d., theron, e.j. & du preez, p.j., 2020, ‘plant community structure and possible vegetation changes after drought on a granite catena in the kruger national park, south africa’, koedoe 62(2), a1585. https://doi.org/10.4102/koedoe.v62i2.1585 note: additional supporting information may be found in the online version of this article. online appendix 1; online appendix 2; and online appendix 3. note: special issue: connections between abiotic and biotic components of a granite catena ecosystem in kruger national park, sub-edited by beanelri janecke and johan van tol. †, 1960-2019. review article plant community structure and possible vegetation changes after drought on a granite catena in the kruger national park, south africa andri c. van aardt, daryl codron, ettienne j. theron, pieter j. du preez received: 04 sept. 2019; accepted: 14 apr. 2020; published: 29 oct. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract a preliminary study investigated the associations between vegetation communities along catenary soil gradients in 2015. the severe drought of 2016 in south africa presented the opportunity to study post-drought savanna vegetation changes. this hillslope transect was surveyed for five successive seasons. the braun-blanquet method was used, and the data were analysed by means of the twinspan algorithm, which resulted in the classification of different communities on the crest, sodic site and riparian area. change in herbaceous and grassy vegetation composition and diversity in the transect is compared between rainfall years, wet and dry seasons, and three different zones (crest, sodic site and riparian areas). spatial and temporal autocorrelation of the woody component shifted the focus to variance within the graminoid and herbaceous layers. clear vegetation changes were observed on the crest and the sodic sites, whereas changes in the riparian area were less obvious. in all three habitats, species richness decreased after the drought and did not reach pre-drought levels even after two years. however, plant species diversity was maintained as climax species were replaced by pioneer and sub-climax species. these changes in community structure, which had reverted to systems dominated by climax species by the end of the sampling period, might be an indication of the savanna ecosystem’s resilience to drought conditions. conservation implications: although clear vegetation changes were observed in the five successive seasons after the drought, this study showed that the savanna ecosystem is relatively resistant to drought and that human intervention is not needed. keywords: drought; vegetation classification; savanna; diversity; catena. introduction the earth’s environment is dominated by three great natural components, namely, climate, vegetation and soil. climate is considered the most important factor influencing the distribution and composition of vegetation on a micro and sub-continental scale (campbell et al. 2008; furley 2010; scholes 1997; schulze 1997). vegetation development is controlled largely by light, temperature and moisture (bond, midgley & woorward 2003; schulze 1997). topography and the chemical and physical compositions of the soil also influence vegetation and, in conjunction with climate, are responsible for the intricate interactions that govern the worldwide distribution of vegetation (campbell et al. 2008; furley 2010; scholes 1997). understanding how these interactions regulate the ecology of plant communities is critical for characterising the impacts of global change on biodiversity at local and regional scales. the savanna biome is unique because it consists of both woody vegetation and a grass layer. climate and other regulating factors likely affect these two components differently, resulting in spatio-temporal heterogeneity of tree:grass compositions. severe droughts, for example, may remove trees, leading to negative effects on woody plant diversity (swemmer 2016; walker et al. 1987; zambatis & biggs 1995). by reducing tree densities, droughts in savanna provide opportunities for drought-adapted flora to thrive, for instance, by promoting seedling recruitment of fast-growing, palatable shrub species and the re-establishment of a grassy layer (swemmer et al. 2018; vetter 2009). in this way, drought can help maintain the balance between trees and grasses (swemmer 2016). grasses, on the other hand, can take decades to recover their productive potential or might recover comfortably before the next drought (swemmer et al. 2018). the herbaceous layer thus also regularly experiences negative responses to drought (zambatis & biggs 1995); however, abbas, bond and midgley (2019) indicated that grasses can resprout vigorously after the onset of rainfall events. in fact, this layer usually responds to droughts and other climate changes first, primarily because of the shallow depth of root penetration. upper soil layers are more susceptible to desiccation than the deeper strata penetrated by many woody plants. furthermore, the extensive root structures of trees increase their access to subterranean reserves of ground water. shorter term responses of grassy and herbaceous vegetation were highlighted by buitenwerf, swemmer and peel (2011), who showed that dynamics of this savanna component are mainly controlled by interannual changes in rainfall. the response of the grass layer to climate is of importance for conservation planning and application, because it is an important food source for grazer populations (staver, wigley-coetzee & botha 2018). the savanna regions of south africa are considered semi-arid, receiving rainfall mostly during the summer months between october and april (walker et al. 1987). fluctuations in annual rainfall, including droughts, are a regular and recurrent feature of the climate (rouault & richard 2003). in more than half of the 80 summer rainfall districts identified by rouault and richard (2003), droughts were recorded during 1926, 1933, 1945, 1949, 1952 1970, 1983 and 1992 (fauchereau et al. 2003; rouault & richard 2003; gommes & petrassi 1996). rouault and richard (2003) and staver et al. (2018) indicated that the 1982–1983 drought was the worst drought recorded since 1922; however, swemmer (2016) indicated that the drought of 2015–2016 was the worst drought that the lowveld experienced in the past 33 years. in the savanna areas of kwazulu-natal, this drought was shown to be the worst in 50 years by abbas et al. (2019). research by hu and fedorov (2019) indicated that the drought of 2015–2016 was worse than the droughts of 1982 and 1997. these studies show that, since the 1960s, drought is more often associated with el niño events; notably, however, annual rainfall during wet years has also increased since the 1970s. south african savannas experienced drought conditions during the rainfall seasons of 2014–2015 and 2015–2016. in the kruger national park (knp), and the surrounding areas of the lowveld, below average rainfall occurred at annual (255 mm) and monthly scales (swemmer 2016). this resulted in devastating effects on vegetation, animal and human welfare in certain areas. these years were also marked by unusually high temperatures, resulting in higher evaporation rates, further reducing water availability (swemmer 2016). the severity of these conditions provided us with the opportunity to study their effects on short-term responses of vegetation, specifically on the grassy and herbaceous component. we conducted a study of seasonal and annual plant community dynamics along a granitic catenal gradient. this catena forms part of a research supersite, where long-term research is needed to establish baselines for monitoring and understanding ecological change (smit et al. 2013). we describe taxonomic community changes, as well as testing for shifts in diversity, over two wet and two dry seasons through the drought period and compare these with pre-drought conditions (april 2015) described elsewhere (theron, van aardt & du preez 2020). we focused only on the herbaceous and grassy components of the vegetation because we were interested in resolving short-term responses in savanna plant resilience to drought. study area the study site is in the southern parts of knp south of skukuza (see study area figure in theron et al. 2020) at 25.111ºs and 31.579ºe. kruger national park falls within the arid ‘bsh’ (hot semi-arid climate) climate type according to the köppen–geiger classification system (venter, scholes & eckhardt 2003). ‘bsh’ is one of the four climate types within this category. the main features of ‘bsh’ climate are distinct seasonal rainfall and temperature variations. mean annual precipitation in knp is generally in the range of 650 mm annually (smit et al. 2013). on a local scale, map of the granite lowveld varies between 450 and 900 mm along the eastern plains and the western escarpment, respectively (eds. mucina & rutherford 2006). however, the average annual total rainfall as recorded at the skukuza meteorological station is 553 mm (zambatis 2006). the mean annual temperature in the vicinity of the study area varies between 21ºc and 22ºc (khomo et al. 2011; scholes, bond & eckhardt 2003). this area experiences an insignificant seasonal and diurnal temperature variation with extreme periods of inundation and aridity (kruger, makamo & shongwe 2002). the study site is underlain by the nelspruit suite geological formation and consists of granite and gneiss mostly occurring in the eastern parts of knp (alard 2009; smit et al. 2013; van zijl & le roux 2014). granite gneiss is widespread in the eastern regions of knp and results in shallow, nutrient-poor soils that vary from grey to red to brown in colour (venter 1990). descriptions of the different soil forms found along the catena at the site were provided in figure 2 within the article by theron et al. (2020). the vegetation type at the study site is mostly granite lowveld (svi3), characterised by a ground layer of tall grasses with intermittent trees and other woody species (eds. mucina & rutherford 2006). methods data collection the same hillslope transect was surveyed for five seasons; the first survey was conducted prior to the onset of severe drought conditions (theron et al. 2020) during december 2015 and april 2016 (figure 1). the second and fourth surveys represent the start of the rainy summer season, while the third and fifth surveys reflect the end thereof (figure 1). relevés of 10 m2 were aligned along a 500 m transect. cover abundance was recorded per species according to the modified braun-blanquet scale (kent 2012; kent & coker 1992; van der maarel & franklin 2013; theron et al. 2020). figure 1: survey events timeline: first season represented by april 2015; december 2015 and april 2016 no sampling because of lack of vegetation; second season represented by december 2016; third season represented by april 2017; fourth season represented by december 2017; and fifth season represented by april 2018. classification, richness and diversity analysis the analysis done by theron et al. (2020) indicated that the catenal vegetation communities can be divided into crest, sodic site and riparian areas. each of these habitat types contains different plant communities that are bound by different soil forms. thus, the analysis of data for the seasons after the drought (december 2016–april 2018) was guided by these differentiations. each topographical unit was thus analysed separately to look at the vegetation composition or change over the period of december 2016–april 2018. during this study, december samples were regarded as wet seasons, and april samples were regarded as dry seasons, irrespective of the delayed effect, because most summer rainfall usually occurred during december. classification vegcap (unpublished database tool designed by n. collins) was used to capture vegetation data into a macro-enabled excel spreadsheet. from there, the data were imported into juice© (tichý & holt 2006) where a modified twinspan classification (roleček et al. 2009) analysis was carried out. parameters for this analysis included the following: pseudo-species cut level (5); analysis was constrained to a minimum group size of 3–54 clusters; and division reached an endpoint if dissimilarity went lower than 0.3 based on average sorensen dissimilarity. the resulting clusters were then arranged within both juice© and excel to form the final vegetation communities. although all the species were recorded during the field surveys, woody species were removed from the data in order to look at the change in graminoids and herbaceous species after the drought. this follows, for example, rouault and richard (2003), who indicated that trees and other vegetation with extensive root structures have access to subterranean reserves of groundwater and will thus not be immediately affected by the drought. the naming of communities and sub-communities was carried out according to the guidelines presented in brown et al. (2013). in order to obtain diagnostic, constant and dominant species, we made use of the analysis of columns of a synoptic table in juice. the frequency thresholds were set at 75, 60 and 50 for the respective diagnostic, constant and dominant species. an asterisk indicates alien invasive species. diversity and richness in addition to descriptions of community composition and how this changed over time, we evaluated changes in diversity and compared these across time for each of the three communities. we compared changes in species richness as well as changes in alpha-diversity. we used the chao estimator as an indicator of species richness, as this index accounts for the occurrences of singletons and doubletons, and the shannon index was used to quantify alpha-diversity. for each sample (i.e. per season and per habitat), ordinal abundance data as scored by the braun-blanquet system were converted to abundance cover data, rounded to integer values, following van der maarel (2007): r = 1; + = 2; 1 = 3; 2a = 8; 2b = 18; 3 = 38; 4 = 63; 5 = 88. diversity estimates were computed using the inext package (hsieh, ma & chao 2016) for r (r core team 2015). the inext function was used for extrapolation and prediction of diversity indices based on rarefaction procedures, with the expected means and standard errors extrapolated from the asymptotes of the fitted accumulation curves (see figure 2). in all cases, accumulation curves approached or reached an asymptote, and observed data represented between 80 and 100% of extrapolated estimates (in the case of species richness), and between 94% and 100% of extrapolated estimates (for shannon diversity), depending on the sample. thus, sampling effort is considered sufficient for reliable estimations of diversity in these communities. figure 2: rarefied species accumulation curves for herbaceous vegetation from the three habitats (cr = crest, s = sodic, r = riparian) sampled at five time intervals from a single catenal transect (10 m2 neighbouring relevés forming 500 m long belt transect) in the southern knp. error bars depict 95% confidence intervals of the richness estimates. ethical considerations ethical approval was obtained from the interfaculty animal ethics committee of the university of the free state (ufs-aed2019/0121). results and discussion classification different plant communities were classified for each topographical unit as defined by theron et al. (2020). in this article, the data for 2015 were not included in the classification in order to prevent a repetition of information. crest communities (december 2016–april 2018) these communities located on the crest zone and upslope beyond the sodic site occur on the clovelly, pinedene, fernwood, estcourt, mispha and sterkspruit soil forms (theron et al. 2020). the soil depth varies from 533 to 620 mm deep, with an average phh₂o of 5.95–6.08. soil texture is mostly loamy sand to coarse loamy sand (theron et al. 2020). vegetation classification resulted in three communities and two sub-communities that perfectly align with the different sampling seasons, showing a clear change in vegetation composition since the onset of the rainy season in december 2016 (online appendix 1). although there are only three communities, the sub-communities of community 3 distinguish between season 4 (s4) (december 2017) and season 5 (s5) (april 2018), although their composition was very similar. the vegetation of the crest communities can be compared to community 3 (vachellia excuvialis–pogonarthria squarrosa) from theron et al. (2020). as indicated in the ‘materials and methods’ section, data on the woody species were removed as it obscured the focus of this study: heliotropium ciliatum–cleome monophylla community zornia glochidiata–crotalaria sphaerocarpa subsp. sphaerocarpa community aristida congesta subsp. barbicollis–bulbostylis *barbata community 3.1. aristida congesta subsp. barbicollis–bulbostylis *barbata–melhania acuminata sub-community 3.2. aristida congesta subsp. barbicollis–bulbostylis *barbata–schmidtia pappophoroides sub-community crest community descriptions: 1. heliotropium ciliatum – cleome monophylla community diagnostic species: cleome monophylla 84.3, heliotropium ciliatum 93.4 constant species: bare soil 77, bulbostylis hispidula 77, chlorophytum recurvifolium 62, cleome monophylla 92, dipcadi papillatum 62, heliotropium ciliatum 100, kyllinga alba 85, phyllanthus maderaspatensis 62, tragus berteronianus 77, urochloa mosambicensis 85 dominant species: none this community mostly represents vegetation sampled during the december 2016 (s2) season. species from species group a (online appendix 1) define this community. these species are mostly absent or occur with very low cover-abundance values in the other communities. from a growth-form perspective, it is notable that this community contains the most geophytic plants. there is also a strong presence of species from species group b and species group i and the ‘pseudo-species’ indicated as bare soil (species group j): 2. zornia glochidiata–crotalaria sphaerocarpa subsp. sphaerocarpa community diagnostic species: crotalaria sphaerocarpa s. sphaerocarpa 82.3, zornia glochidiata 84.8 constant species: aristida congesta s. congesta 79, crotalaria sphaerocarpa s. sphaerocarpa 74, eragrostis superba 63, pogonarthria squarrosa 95, setaria sphacelata v. sericea 63, tricholaena monachne 68, vernonia fastigiata 79, zornia glochidiata 89 dominant species: none vegetation found in this community represents the sampling during april 2017, which is mostly dominated by species from species group c (online appendix 1). again, the species found here do not occur in other communities. notable is the high cover abundance of species found in this community when compared to that of community 1. furthermore, species from species group b are shared between community 1 and community 2; however, aristida congesta subsp. congesta occur with much higher cover abundance in community 2 than in community 3. a possible explanation for that might be the increase in rainfall after the severe drought experienced in 2015–2016. from species group h, it is also clear that the grasses eragrostis cylindriflora, aristida adscensionis and melinis repens start to establish with average cover-abundance values: 3. aristida congesta subsp. barbicollis–bulbostylis *barbata community diagnostic species: aristida congesta s. barbicollis 77.1, bulbostylis barbata 79.4 constant species: aristida congesta s. barbicollis 75, bare soil 81, bulbostylis barbata 78, schmidtia pappophoroides 66 dominant species: none this community represents sampling seasons 4 and 5 (december 2017 and april 2018), which is more or less one year after rainfall occurred that terminated the 2015–2016 drought. species from species group f distinguishes this community from the other communities. plants from species group h also started to occur in more relevés during these seasons, which might indicate that the veld was starting to improve after the drought conditions: 3.1. aristida congesta subsp. barbicollis–bulbostylis *barbata–melhania acuminata sub-community diagnostic species: melhania acuminata 79.4, panicum coloratum 78.5 constant species: aristida congesta s. barbicollis 94, bare soil 100, bulbostylis barbata 81, eragrostis cylindriflora 69, hibiscus micranthus v. micranthus 62, melhania acuminata 88, panicum coloratum 75, panicum maximum 69, perotis patens 88, pogonarthria squarrosa 94, tricholaena monachne 75 dominant species: none sub-community 3.1 mostly represents vegetation sampled during april 2018 (s5). this sub-community is distinguished by the presence of species from species group e, which are either absent from other communities or occur with very low cover-abundance values. when looking at species group d, it is clear that the graminoids (pogonarthria squarrosa, tricholaena monachne, eragrostis superba and digitaria eriantha) mostly occur during the december sampling seasons (s3 and s5). although these grasses do occur in other communities, it is with very low cover abundance: 3.2. aristida congesta subsp. barbicollis–bulbostylis *barbata–schmidtia pappophoroides sub-community diagnostic species: none constant species: aristida adscensionis 62, bare soil 62, bulbostylis barbata 75, schmidtia pappophoroides 75, urochloa panicoides 62 dominant species: none this sub-community represents crest vegetation during april 2017 (s4). species group g distinguishes this sub-community from the aristida congesta subsp. barbicollis–bulbostylis *barbata–melhania acuminate sub-community 3.1. furthermore, the absence of species from species group e is also very prominent in this sub-community. however, from species group f it is clear that the cover-abundance values of aristida congesta subsp. barbicollis and panicum maximum decreased from sub-community 3.1 to 3.2. a possible explanation might be that during december 2017 (s4; sub-community 3.2), the species only started to establish at the site and favourable environmental conditions such as an increase in rainfall allowed the improvement of cover in april 2018 (s5; sub-community 3.1). the above community descriptions cannot be directly compared to what was found in 2015 (theron et al. 2020) because of the removal of the woody species which then dominated the community. there are, however, species such as aristida congesta, tricholaena monachne, melhania acuminata, panicum maximum and perotis patens that occur on the site during most of the sampling seasons. it is, nevertheless, clear that the grass pogonarthria squarrosa (species group d) only started to reappear in the vegetation in growing season 3 (april 2017), then diminished and reappeared again in growing season 5 (april 2018). this might indicate that this grass is also restricted to certain sampling seasons and does not occur on the crest sites throughout the year. van oudtshoorn (2018) indicated that p. squarrosa is a weak perennial tufted grass that can grow for two to five seasons. diminishing of this grass during season 4 (december 2017) might therefore still be due to the effects of the drought; indicating that the drought still affected vegetation composition one year after the onset of the rainy season. sodic site communities (december 2016–april 2018) the communities occur between the crest and the riparian area on the mid-slope of the hill, and are also sodic sites. soils are mostly of the sterkspruit form; however, there were also instances of mispah soil forms present. the depth varies between 180 mm and 500 mm with an average phh2o of 6.20–6.43. soil texture is coarse sandy loam. the vegetation classification resulted in two communities and four sub-communities (online appendix 2). in terms of vegetation composition, these communities can be compared to the dactyloctenium aegyptium–sporobolus nitens (community 4) of theron et al. (2020): tribulus terrestris–portulaca *oleracea community tribulus terrestris–portulaca *oleracea–urochloa panicoides sub-community tribulus terrestris–portulaca *oleracea–heliotropium ciliatum sub-community chloris virgata–eragrostis cylindriflora community chloris virgata–eragrostis cylindriflora–sporobolus nitens sub-community chloris virgata–eragrostis cylindriflora–chloris gayana sub-community sodic site community descriptions: 1. tribulus terrestris–portulaca *oleracea community diagnostic species: portulaca oleracea 78.5, tribulus terrestris 89.8 constant species: bare soil 96, cynodon dactylon 64, portulaca *oleracea 86, schkuhria pinnata 100, tribulus terrestris 89, urochloa mosambicensis 68 dominant species: none this community is defined by species from species group c, which occur here and are absent from other communities or occur with low cover-abundance values. cynodon dactylon is known as a pioneer grass (van oudtshoorn 2018) and tribilus terrestris is known to occur in disturbed areas (van wyk & malan 1998). portulaca *oleracea is a creeping succulent that grows vigorously under warm conditions covering the soil surface (bromilow 2018). this community was mostly restricted to april 2017 (s2) and april 2018 (s4). thus, again as seen in the crest communities, there are certain species that show preferences for certain sampling seasons: 1.1. tribulus terrestris–portulaca *oleracea–urochloa panicoides sub-community diagnostic species: urochloa panicoides 91.7 constant species: alternanthera pungens 62, bare soil 94, portulaca oleracea 81, schkuhria pinnata 100, sporobolus nitens 69, tribulus terrestris 81, urochloa panicoides 88 dominant species: none the vegetation found in this sub-community mostly represents species from growing season 4 with a single occurrence of season 2. species from species group a (online appendix 2) define this sub-community. these species are completely absent or occur with very low cover-abundance values in other communities and sub-community on the sodic site. urochloa panicoides, which defines this sub-community, is known as a pioneer annual tufted grass and will thus only be present for one season (van oudtshoorn 2018). in this subcommunity, this grass co-occurs with sporobolus nitens, which defined the communities found in 2015 before the drought: 1.2. tribulus terrestris–portulaca *oleracea–heliotropium ciliatum sub-community diagnostic species: none constant species: bare soil 100, cynodon dactylon 75, gomphrena celosioides 67, heliotropium ciliatum 92, ledebouria luteola 67, portulaca oleracea 92, schkuhria pinnata 100, tribulus terrestris 100, urochloa mosambicensis 92 dominant species: none this sub-community is mostly represented by growing season 2 (december 2016) at the onset of the rainy season after the severe drought. furthermore, this sub-community is defined by the presence of species from species group b, which include two geophytic species. there is also a complete absence of the species from species group a in this sub-community. very notable in this sub-community is the almost complete absence of sporobolus nitens (species group g) and dactyloctenium aegyptium (species group f), which completely dominated the vegetation during 2015 (growing season 1) (theron et al. 2020): 2. chloris virgata–eragrostis cylindriflora community diagnostic species: chloris virgata 83.5 constant species: alternanthera pungens 78, bare soil 85, chloris virgata 100, dactyloctenium aegyptium 67, eragrostis cylindriflora 74, schkuhria pinnata 96, sporobolus nitens 93, urochloa mosambicensis 70 dominant species: bare soil 4 this community is defined by the presence of species from species group f. although some of the species that occur in this species group were also present in community 1, they occur with much higher cover-abundance values in community 2: 2.1. chloris virgata–eragrostis cylindriflora–sporobolus nitens sub-community diagnostic species: none constant species: alternanthera pungens 92, bare soil 100, chloris virgata 100, dactyloctenium aegyptium 62, gomphrena celosioides 92, schkuhria pinnata 92, sporobolus nitens 100, urochloa mosambicensis 92 dominant species: none vegetation in this sub-community is mostly from growing season 5 (april 2018) with a single occurrence of vegetation from growing season 3 (april 2017). although s. nitens is the diagnostic species for this sub-community, the presence of species from species group d defines this sub-community. these species are completely absent from sub-community 2.2. season 5 marks the return of s. nitens (with high cover abundance) and dactyloctenium aegyptium (with low cover-abundance and only in some relevés) which dominated the communities found on the sodic site by theron et al. (2020) in 2015: 2.2. chloris virgata–eragrostis cylindriflora–chloris gayana sub-community diagnostic species: none constant species: alternanthera pungens 64, bare soil 71, chloris gayana 71, chloris virgata 100, dactyloctenium aegyptium 71, eragrostis cylindriflora 93, schkuhria pinnata 100, sporobolus nitens 86 dominant species: bare soil 7 sub-community 2.2 is defined by the presence of perennial grasses from species group e, which are absent from sub-community 2.1. although having low cover abundances and not occurring in all relevés, this is the only season in which these grass species were found. all three of these grass species (chloris gayana, eragrostis gummiflua and aristida stipitata) are regarded by van oudtshoorn (2018) as sub-climax species, which might indicate that after the third season, the sodic site started to recover from the severe drought of 2015–2016. species such as schkuhria pinnata, urochloa mosambicensis and chloris virgata occurred on the site through most of the sampling seasons since 2015. sporobolus nitens that formed part of the diagnostic species that defined the sodic site communities in 2015 (theron et al. 2020) only started to appear in april 2017 (s3) with low cover-abundance values. the high cover-abundance values of this diagnostic species only started returning in december 2017 (s4) and increased in april 2018 (s5). from online appendix 2, it is clear that certain species on the sodic site are restricted to certain sampling seasons such as april or december. however, it is also clear that the vegetation composition on the sodic site changed from december 2016 until april 2018. the mentioned changes can possibly be assigned to the recovery of the site after the drought of 2015–2016. riparian area communities (december 2016–april 2018) the communities occur between the sodic site on the lower midslope of the hill and the drainage line. soil forms found in this area include dundee, mispah, bonheim and sterkspruit. the depth of these soils varies from 100 mm to 600 mm with an average phh20 of between 6.21–6.73. soil texture also varies from sandy loam to loamy to sandy clay loam. in contrast to the other terrain units depicted along the catena, the riparian area’s classification did not result in communities that could depict the different seasons of sampling. the vegetation classification resulted in five communities (online appendix 3). the vegetation of the riparian communities can be compared to communities 1 (panicum maximum–pupalia lappacea) and 2 (themeda triandra–flueggea virosa) from theron et al.’s (2020) 2015 study: eragrostis cylindriflora–urochloa mosambicensis community themeda triandra–panicum maximum community eragrostis superba–bothriochloa insculpta community eragrostis rigidior–urochloa mosambicensis community bothriochloa radicans–eragrostis superba community riparian area community descriptions: 1. eragrostis cylindriflora–urochloa mosambicensis community diagnostic species: none constant species: bare soil 67, eragrostis cylindriflora 83, panicum maximum 75, urochloa mosambicensis 83 dominant species: none eragrostis cylindriflora (species group g) and urochloa mosambicensis (species group h) define this community. species from species group a are mostly present in community 1 and absent or occur with low cover-abundance value in other communities in the riparian areas. this community represents sampling seasons 2, 4 and 5. it is notable that none of the relevés done during season 2 (just at the onset of the rainy season) is present in this community. community 1 also share a lot of species from species group b with community 2: 2. themeda triandra–panicum maximum community diagnostic species: none constant species: cymbopogon caesius 68, panicum maximum 95, themeda triandra 95, urochloa mosambicensis 74 dominant species: none community 2 is defined by the presence of species from species group c, which are mostly restricted to this community although they occur with low cover-abundance values. notable in this community is the strong presence of themeda triandra (species group d) and panicum maximum (species group h), which were also present as diagnostic species defining the riparian areas in theron et al. (2020). it seems as if themeda triandra is mostly limited to this community with high cover-abundance values. however, panicum maximum occurs throughout all the communities present in the riparian area throughout all the sampling seasons. this community is also mostly represented by sampling seasons 3 and 5 with some instances of sampling season 4: 3. eragrostis superba–bothriochloa insculpta community diagnostic species: none constant species: bare soil 100, bothriochloa insculpta 67, eragrostis superba 100, panicum maximum 67, urochloa mosambicensis 67 dominant species: none community 3 is the community with the lowest number of species in all the communities found in the riparian area, and there are no species that clearly distinguish this community from all the other communities in the riparian area. the cover abundance of species in this community is also low, and species do not occur in all the relevés found in this community. it is only the grass eragrostis superba (species group h), known to grow in disturbed areas (van oudtshoorn 2018), that occurs in all three relevés that make up the community. vegetation in this community mostly represents sampling seasons 2 and 5. the reason for the low number of species might be that the vegetation still needed to recover after the drought. 4. eragrostis rigidior–urochloa mosambicensis community diagnostic species: none constant species: bare soil 100, eragrostis rigidior 77, eragrostis superba 62, urochloa mosambicensis 92 dominant species: none vegetation in this community is dominated by species from species group e, which are mostly absent from the other communities in the riparian area. furthermore, urochloa mosambicensis (species group h) also occurs more frequently and with a higher cover abundance in this community. according to van oudtshoorn (2018), u. mosambicensis grows in disturbed or overgrazed and trampled areas. the high occurrence of this species in the riparian area might indicate that animals were seeking shade in order to evade the heat of the day during the drought (2015–2016). he further also indicated that eragrostis rigidior is known to occur in disturbed soil. it is also important to note that most of the relevés present in this community represent sampling season 2, which was just after the 2015–2016 drought: 5. bothriochloa radicans–eragrostis superba community diagnostic species: none constant species: bare soil 62, bothriochloa radicans 77, dicoma tomentosa 62, eragrostis superba 69 dominant species: bare soil 8 this is the only community that is solely represented by vegetation sampled during sampling season 4. the vegetation is mostly dominated by the presence of species from species group f, which is mostly absent or occurs with low cover-abundance values in other communities of the riparian area. the grasses bothriochloa radicans and eragrostis trichophora are known to occur in areas with additional moisture or where water collects (van oudtshoorn 2018). a possible explanation for this might be that after rains, water can remain close to the surface in the vicinity of the riparian area, which contributes to the additional moisture that is favourable for these grasses. although there is no distinction to be made between the sampling seasons in the riparian area of the study site, there are differences in the vegetation composition over the study period. when comparing the vegetation of the riparian area with communities 1 and 2 (theron et al. 2020), it is clear that panicum maximum, urochloa mosambicenis and themeda triandra remained an important part of the vegetation composition over all the different sampling seasons. richness and diversity of plant communities from figure 3a, it seems as if the species richness decreased at all the sites during the drought and subsequently increased more-or-less progressively through time as the communities recovered from the drought between 2015 and the onset of the current sampling period. however, preversus post-drought richness estimates are only significantly different for the sodic and riparian habitats (non-overlapping 95% confidence intervals between groups); variance in estimates for the crest communities is high and overlaps with the pre-drought estimate. interestingly, however, the recovery in species richness in sodic and riparian habitats appeared to slow or even reverse by the end of the study period (april 2018), although this could be because the final sample was taken in the dry season. overall, species richness in crest habitats was greater than in both sodic and riparian habitats. figure 3: bar chart representing the species richness (chao estimate) and diversity (shannon index) for the different sampling seasons at the different topographical units. the height of each column represents the mean, and the error bar represents the upper portion of the 95% confidence interval. figure 3b represents the changes that took place in diversity over the different sampling seasons. in contrast to richness, species diversity did not differ between preand post-drought periods. however, a more cyclic seasonal shift is apparent, in that diversity was often highest in the wet seasons (december samples), compared with both dry season samples (april). the sodic and riparian habitats are an exception to this trend, because diversity in these areas was low in december 2016, perhaps because of a lag in recovery from the drought. as with species richness, diversity was also consistently greater in the crest, compared with the other two habitats. while these indices of diversity provide some indication about changes in the studied communities, their overall function might be better represented in terms of changes in plant functional groups. indeed, in all three habitats, the proportional representation of plant functional groups differed between 2015 and 2016, with climax and subclimax species being replaced by pioneers, perennials, annuals and – in some cases, especially in the sodic habitat – bare soil (figure 4). by the end of the sampling period, however, the frequency distribution of functional groups at each habitat was qualitatively similar to pre-drought conditions. figure 4: graphs showing the different percentage covers of different growth forms ([a] crest, [b], sodic site and [c] riparian) during the different sampling seasons. general discussion with this study, we aimed to determine how savanna plant communities along a catenal gradient changed over time following a severe drought. the catenal gradient studied could be divided into three plant communities – crest and midslope with the highest diversity; sodic site, and riparian areas. the crest and sodic sites further showed a definite change in species composition among the different sampling seasons. there was also an association between april sampling seasons for the crest as well as associations between the december and april sampling sites for the sodic site. vegetation in the riparian section of the study revealed no clear distinction between different sampling seasons or any correlation between april and december. in a study by scholes (1985), he investigated the drought of 1981–1983 and found that the grasses were more adversely affected by the drought than the trees. although we excluded data for woody plants from this study, it is clear that vegetation changes took place in the ground layer (graminoids, forbs, herbs and geophytes), especially in the crest and sodic site communities (see janecke 2020). previous studies have indicated that the physical and chemical properties of soils would affect grass mortality rates during drought conditions (khomo & rogers 2005; khomo et al. 2011; scholes 1985). specifically referring to the characteristics of the study site and its catenary properties, it is expected that grasses inhabiting the sandy crest and valley bottoms would have a higher mortality rate than those inhabiting the clay-rich sodic sites and downslopes. the physical properties of sandy soils would compound the effects of droughts because they retain less water than do clay soils, and also through exasperating water infiltration and percolation of any available surface water. the effect of soil properties was shown to also affect this catena complex (theron et al. 2020). this is also comparable to this study because most of the grass species dominating the climax community (sampling s1; 2015) returned to the vegetation composition of communities during sampling season 3 (april 2017). we furthermore found that richness and diversity declined and that recovery was not complete two years after the drought, especially in the sodic and riparian habitats, which have maintained a low level of species richness throughout the sampling period. these shifts coincided with changes in functional group representation following the drought. conclusion definite changes in plant community composition were seen in the crest, midslope and sodic sites during the different sampling seasons. shifts were also seen in terms of species composition at certain times of the year. this was not always clear in terms of richness and diversity of plant species. we would, however, be cautious to extrapolate these findings to all vegetation successions along a catena. in the riparian area, no distinctions were clear between the different sampling seasons and no cyclic correspondence was observed between april and december. this phenomenon might be ascribed to water movement through the process of hydraulic lift from deeper soil layers which lessen the impact of drought on the vegetation. we recommend that future studies following droughts should be done over more sampling seasons than reported here to better relate seasons to plant assemblages. lastly, the recovery of the plant growth forms from 2015 to 2018 might be an indication of the resilience of the savanna ecosystem, in spite of the recovery not being complete. acknowledgements the authors thank the south african national parks for providing them with access to the research sites within kruger national park. a special thanks goes to the field rangers who accompanied them during the surveys. the authors also thank louis scott and leslie brown for suggestions on the writing of the manuscript. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions e.j.t. and a.c.v.a. (partially) were responsible for the fieldwork and data collection during field surveys. a.c.v.a. and p.j.d.p. contributed towards the analysis and interpretation of the plant communities. d.c. contributed towards the analysis and interpretation of the statistical elements of the article. all authors contributed to the writing of the manuscript. funding information the authors are grateful to the university of the free state (ufs) strategic research fund for partially funding this multidisciplinary research. data availability study data are available and may be provided, on request, by the corresponding author. data from all research done within kruger national park is placed within the sanparks repository (not for free, open access). disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references abbas, h.a., bond, w.j. & midgley, j.j., 2019, ‘the worst drought in 50 years in a south african savannah: limited impact on vegetation’, african journal of ecology 57(4), 1–10. https://doi.org/10.1111/aje.12640 alard, g.f., 2009, ‘a comparison of grass production and utilization in sodic and crest patches on a semi-arid granitic savanna catena in the southern kruger national park, south africa’, msc thesis, faculty of science, university of the witwatersrand. bond, w.j., midgley, g.f. & woorward, f.i., 2003, ‘what controls south african vegetation – climate or fire?’ south african journal of botany 69(1), 79–91. https://doi.org/10.1016/s0254-6299(15)30362-8 bromilow, c., 2018, problem plants and alien weeds of southern africa, 4th edn., briza publications, pretoria. brown, l.r., du preez, p.j., bezuidenhoudt, h., bredenkamp, g.j, mostert, t.h.c. & collins, n.b., 2013, ‘guidelines for phytosociological classifications and descriptions of vegetation in southern africa’, koedoe 55(1), art. #1103, 10. https://doi.org/10.4102/koedoe.v55i1.1103 buitenwerf, r., swemmer, a.m. & peel, m.j.s., 2011, ‘long-term dynamics of herbaceous vegetation structure and composition in two african savanna reserves’, journal of applied ecology 48(1), 238–246. https://doi.org/10.1111/j.1365-2664.2010.01895.x campbell, n.a., reece, j.b, urry, l.a., cain, m.l., wasserman, s.a., minorsky, p.v., 2008, biology, 8th edn., pearson benjamin cummings, san francisco, ca. fauchereau, n., trzaska, s., rouault, m. & richard, y., 2003, ‘rainfall variability and changes in southern africa during the 20th century in the global warming context’, natural hazards 29(2), 139–154. https://doi.org/10.1023/a:1023630924100 furley, p., 2010, ‘tropical savannas: biomass, plant ecology, and the role of fire and soil on vegetation’, progress in physical geography 34(4), 563–585. https://doi.org/10.1177/0309133310364934 gommes, r. & petrassi, f., 1996, rainfall variability and drought in sub-saharan africa, food and agriculture organization of the united nations, rome. hsieh, t.c., ma, k.h. & chao, a., 2016, ‘inext: an r package for rarefaction and extrapolation of species diversity (hill numbers)’, methods in ecology and evolution 7(12), 1451–1456. https://doi.org/10.1111/2041-210x.12613 hu, s. & fedorov, a.v., 2019, ‘the extreme el niño of 2015–2016: the role of westerly and easterly wind bursts, and preconditioning by the failed 2014 event’, climate dynamics 48(1), 1–19. https://doi.org/10.1007/s00382-017-3531-2 janecke, b.b., 2020, ‘vegetation structure and spatial heterogeneity in the granite supersite, kruger national park’, koedoe 62(2), a1591. https://doi.org/10.4102/koedoe.v62i2.1591 kent, m., 2012, vegetation description and data analysis: a practical approach, 2nd edn., wiley-blackwell publishers, west sussex. kent, m. & coker, c., 1992, vegetation description and analysis: a practical approach, bellhaven press, west sussex. khomo, l.m., hartshorn, a.s., rogers, k.h. & chadwick, o.a., 2011, ‘impact of rainfall and topography on the distribution of clays and major cations in granitic catenas of southern africa’, catena 87(1), 119–128. https://doi.org/10.1016/j.catena.2011.05.017 khomo, l.m. & rogers, k.h., 2005, ‘proposed mechanism for the origin of sodic patches in kruger national park, south africa’, african journal of ecology 43(1), 29–34. https://doi.org/10.1111/j.1365-2028.2004.00532.x kruger, a.c., makamo, l.b. & shongwe, s., 2002, ‘an analysis of skukuza climate data’, koedoe 45(1), 1–7. https://doi.org/10.4102/koedoe.v45i1.16 mucina, l. & rutherford, m.c. (eds.), 2006, the vegetation of south africa, lesotho and swaziland, strelizia 19, south african national biodiversity institute, pretoria. r core team, 2015, computer software, r: a language and environment for statistical computing, r foundation for statistical computing, vienna. roleček, j., lubomír, t., david, z., & milan, c., 2009, ‘modified twinspan classification in which the hierarchy respects cluster heterogeneity’, journal of vegetation science 20(4), 596–602. https://doi.org/10.1111/j.1654-1103.2009.01062.x rouault, m. & richard, y., 2003, ‘intensity and spatial extension of drought in south africa at different time scales’, water sa 29(4), 489–500. https://doi.org/10.4314/wsa.v29i4.5057 scholes, r.j., 1985, ‘drought related grass, tree and herbivore mortality in a southern african savanna’, in j.c. tothill & j.j. mott (eds.), ecology and management of the world’s savannas, pp. 350–353, australian academy of science, washington, dc. scholes, r.j., 1997, ‘savanna’, in r.m. cowling, d.m. richardson & s.m. pierce (eds.), vegetation of southern africa, pp. 258–277, cambridge university press, cambridge. scholes, r.j., bond, w.j. & eckhardt, h.c., 2003, ‘vegetation dynamics in the kruger ecosystem’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 242–262, island press, london. schulze, r.e., 1997, ‘climate’, in r.m. cowling, d.m. richardson & s.m. pierce (eds.), vegetation of southern africa, pp. 21–42, cambridge university press, cambridge. smit, i.p.j., riddell, e.s., cullum, c. & petersen, r., 2013, ‘kruger national park research supersites: establishing long-term research site for cross-disciplinary, multiscaled learning’, koedoe 55(1), art. #1107, 7, https://doi.org/10.4102/koedoe.v55i1.1107 staver, a.c., wigley-coetsee, c. & botha, j., 2018, ‘grazer movements exacerbate grass declines during drought in an african savanna’, journal of ecology 107(1), 1482–1491. swemmer, a.m., bond, w.j., donaldson, j., hempson, g.p., malherbe, j. & smit, i.p.j., 2018, ‘the ecology of drought – a workshop report’, south african journal of science 114(9–10) art.#5098, 3. https://doi.org/10.17159/sajs.2018/5098 swemmer, t., 2016, the lowveld’s worst drought in 33 years? understanding the long-term impacts, viewed 13 april 2017, from http://www.saeon.ac.za/newsletter/archives/2016february/doc02. theron, e.j., van aardt, a.c. & du preez, p.j., 2020, ‘vegetation distribution along a granite catena, southern kruger national park, south africa’, koedoe 62(2), a1588. https://doi.org/10.4102/koedoe.v62i2.1588 tichý, l. & holt, j., 2006, juice: a program for management, analysis and classification of ecological data, vegetation science group, brno. van der maarel, e., 2007, ‘transformation of cover-abundance values for appropriate numerical treatment – alternatives to the proposals by podani’, journal of vegetation science 18(5), 767–770. https://doi.org/10.1111/j.1654-1103.2007.tb02592.x van der maarel, e. & franklin, j., 2013, vegetation ecology, 2nd edn., john wiley & sons, chichester. van oudtshoorn, f., 2018, guide to grasses of southern africa, briza publications, pretoria. van wyk, b. & malan, s., 1998, field guide to the wild flowers of the highveld, struik nature, cape town. van zijl, g. & le roux, p., 2014, ‘creating a conceptual hydrological soil response map for the stevenson hamilton research supersite, kruger national park, south africa’, water sa 40(2), 331–336. venter, f.j., 1990, ‘a classification of land for management planning in the kruger national park’, phd thesis, university of south africa, pretoria. venter, f.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation pattern’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83–129, island press, london. vetter, s., 2009, ‘drought, change and resilience in south africa’s arid and semi-arid rangelands’, south african journal of science 105(1–2), 29–33. https://doi.org/10.4102/sajs.v105i1/2.35 walker, b.h., emslie, r.h., owen-smith, r.n. & scholes, r.j., 1987, ‘to cull or not to cull: lessons from a southern african drought’, journal of applied ecology 24(2), 381–401. https://doi.org/10.2307/2403882 zambatis, n., 2006, average monthly and seasonal temperatures (°c) of the kruger national park, scientific services, kruger national park, cape town. zambatis, n. & biggs, h.c., 1995, ‘rainfall and temperatures during the 1991/92 drought in the kruger national park’, koedoe 38(1), 1–16. https://doi.org/10.4102/koedoe.v38i1.301 abstract background scope and context monitoring system implementation and adaptive management conclusion acknowledgements references about the author(s) wessel vermeulen garden route scientific services, conservation services, south african national parks, knysna, south africa nicola van wilgen cape research centre, conservation services, south african national parks, steenberg, south africa centre for invasion biology, stellenbosch university, south africa kyle smith garden route scientific services, conservation services, south african national parks, sedgefield, south africa mbulelo dopolo cape research centre, conservation services, south african national parks, steenberg, south africa louise swemmer savanna and arid research unit, conservation services, south african national parks, phalaborwa, south africa wendy annecke cape research centre, conservation services, south african national parks, steenberg, south africa hugo bezuidenhout savanna and arid research unit, conservation services, south african national parks, kimberley, south africa applied behavioural ecology and ecosystem research unit, university of south africa, south africa graham durrheim garden route scientific services, conservation services, south african national parks, knysna, south africa nick hanekom garden route scientific services, conservation services, south african national parks, sedgefield, south africa howard hendricks conservation services, south african national parks, pretoria, south africa melodie mcgeoch cape research centre, conservation services, south african national parks, steenberg, south africa ntombizodwa ngubeni garden route scientific services, conservation services, south african national parks, knysna, south africa alexis symonds conservation services, south african national parks, pretoria, south africa citation vermeulen, w., van wilgen, n., smith, k., dopolo, m., swemmer, l., annecke, w. et al., 2019, ‘monitoring consumptive resource use in south african national parks’, koedoe 61(1), a1516. https://doi.org/10.4102/koedoe.v61i1.1516 original research monitoring consumptive resource use in south african national parks wessel vermeulen, nicola van wilgen, kyle smith, mbulelo dopolo, louise swemmer, wendy annecke, hugo bezuidenhout, graham durrheim, nick hanekom, howard hendricks, melodie mcgeoch, ntombizodwa ngubeni, alexis symonds received: 17 jan. 2018; accepted: 14 sept. 2018; published: 30 jan. 2019 copyright: © 2019. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract monitoring is an essential component of measuring the performance of protected areas. this requirement led to the development of a biodiversity monitoring system for south african national parks (sanparks). the system comprises of ten major programmes, each focusing on a core area of conservation biodiversity monitoring, with resource use being one of the focal areas. with the growing appreciation of the importance of natural resources for the socio-economic well-being of communities and other stakeholders, sustainable resource use is an important component of the management of natural areas and national parks. to gauge sustainability, a sound monitoring and research programme that fits within the context of the sanparks’ adaptive management approach towards social-ecological system management is required. the purpose of this article was to define the context and scope in which consumptive resource use takes place within sanparks and to outline the criteria necessary for developing a sound monitoring programme to assess the sustainability of such use. the monitoring programme is structured in view of the fact that sustainable resource use is achievable only where all dimensions of sustainability (social, economic and ecological) are considered simultaneously. in terms of the social and economic dimensions of sustainability, the programme provides for assessing stakeholder needs, trends in resource use and the social and economic impacts of resource use. monitoring that relates to the ecological dimension of sustainability of biological resource use deals with the rate of turnover and population dynamics of target species, as well as harvest impact. in terms of abiotic (non-renewable) resources, monitoring deals with sound management practices to minimise impact on the environment, and to optimise benefits through responsible use. conservation implications: the resource use monitoring programme is intended to ensure that monitoring relating to the harvesting of natural resources from national parks is scientifically sound and conducted in a structured way, towards meeting the objective of sustainable use and compliance with national legislation. the article illustrates how sanparks meets its obligation to monitor biodiversity conservation while at the same time meeting the needs for the consumptive use of resources. background the primary mandate of protected areas is the conservation of biodiversity both for its intrinsic value and for the conservation-related benefits for people. although protected areas provide a wide range of opportunities from which many stakeholders benefit, they are under increasing threat from a range of external and internal factors. as a result, protected areas are under pressure to do more in terms of their ecological, social and economic contributions than ever before (ervin et al. 2010). monitoring is an essential component of both measuring the performance of protected areas and ensuring their sustainability. the requirement for biodiversity monitoring in national parks is specified in national legislation and international policy, as well as by south african national parks’ (sanparks) own adaptive management philosophy. these guiding frameworks and principles have led to the development of a biodiversity monitoring system (bms) for sanparks (mcgeoch et al. 2011). this system comprises ten major programmes, each focusing on a core area of conservation biodiversity monitoring. resource use has been identified as one of these areas, together with, for example, freshwater and estuarine systems, habitat representation and rehabilitation, species of special concern, invasive species, disease and climate. with an increasing population and high unemployment levels in south africa (stats sa 2017), the demand for access to natural resources (subsistence, recreational and commercial purposes) within protected areas is growing (van wilgen & mcgeoch 2014). with the greater appreciation of the importance of natural resources for the socio-economic well-being of people (clarke & grundy 2004; shackleton, shanley & ndoye 2007; twine 2011), sustainable resource use is an important aspect of the management of natural areas and increasingly also national parks. as such, the importance of resource use in national parks is embedded in the sanparks’ mission (sanparks 2016a): to develop, expand, manage and promote a system of sustainable national parks that represents biodiversity and heritage assets, through innovation and best practice for the just and equitable benefit of current and future generations. (p. 10) while access to resources has the potential to provide varied benefits to people, overharvesting and/or the illegal or unauthorised harvesting of resources has been identified as an international threat to biodiversity and ecosystem services and is a driver of global change, particularly in the marine environment (butchart et al. 2010; millennium ecosystem assessment 2005; pauly 2008; van wilgen & herbst 2017). in south africa, uncontrolled and illegal harvesting from both terrestrial and aquatic ecosystems in many areas and the lack of harvesting systems that ensure the sustainable use of resources are of growing concern (ouédraogo 2001; ticktin 2004; van wilgen & herbst 2017). overharvesting and unsustainable use not only challenge meeting biodiversity management objectives but also threaten sustained social and economic benefits undermining system sustainability. paradoxically, a lack of access to harvest resources (under-harvesting) may similarly impact system sustainability. the challenge for protected areas is therefore to put resource harvesting systems in place that can facilitate equitable, legal and controlled access to resources inside national parks that enhance social relevance (through impacting positively human well-being), and in so doing build a vested interest in conservation while maintaining ecological integrity and economic viability (swemmer, mmethi & twine 2017). this article provides a brief overview of the resource use monitoring programme adopted by sanparks (vermeulen et al. 2011) and describes how a sound, holistic resource use monitoring and research programme can contribute towards the sustainability of protected areas through more effective social, economic and ecological system management. as the monitoring framework is based on an established, generic process for harvest system development, and consolidates the ecological, social and economic aspects of sustainability, it can be replicated by other conservation and management agencies in south africa and further afield. although the context in which it is applied may differ, the system is flexible enough to accommodate different challenges and focus areas. its wider application would contribute to consistency in resource use monitoring and reporting. two case studies are presented to illustrate challenges and success in implementing the programme in sanparks. scope and context local context natural resources play a significant role in the well-being of millions of south africans. despite widespread electrification, low household income levels result in a significant number of people in the country still relying on firewood as a primary fuel source or using a combination of fuel sources (department of energy 2013; uhunamure, nethengwe & musyok 2017). timber, thatching grass and other resources are widely used for building material, while many people rely on natural pastures for livestock fodder. millions of people use a wide variety of plants and animal products as medicines (mander et al. 2007; van wyk, oudtshoorn & gericke 2009) and natural foods (plants, animals and fish) are harvested on a wide scale around the country (paumgarten, locatelli & witkowski 2018; shackleton & shackleton 2004). south african national parks manages 19 national parks in south africa, ranging in size from approximately 50 km2 (bontebok national park) to roughly 20 000 km2 (kruger national park) and covering nearly 40 000 km2. four parks also have marine-protected areas (mpas) adjoining protected land, and collectively, the parks span eight of the nine biomes present in south africa. historically access to parks and their resources has been confined to exclusive groups. some open access parks, such as agulhas national park and garden route national park (grnp), have experienced more unregulated extractive use than others, such as kruger national park (knp), that have hard boundaries. as a result of this inequality in access, many people are resentful about the lack of resources and access to them that they experience. parks are embedded within a significant array of diverse social, political, historical and economic contexts, some with a history of forced removals of local residents during the establishment of parks and others with a history of restricted access policies that did not allow black visitors until south africa became a democracy in 1994. as such sanparks are fraught with contrasts both in terms of constituency and visitation (biggs et al. 2014) as well as stark contrasts in land use on either side of park boundaries. such contrasts pose a number of biological and social threats to biodiversity, providing important context for implementing locally appropriate resource use projects and programmes. threats include the introduction of alien species (spear et al. 2013), pollution of freshwater entering parks (roux & nel 2013) and spread of agriculturally relevant diseases or genes between plants and animals inside and outside of parks (see, e.g., sanparks [2016b] for threats to the cape mountain zebra equus zebra zebra in this regard). in some cases, increasing contrasts in natural resource abundance and diversity inside and outside parks reinforces the historical alienation that many people living adjacent to certain national parks feel at not being able to access resources within protected areas. many of these threats are further exacerbated by climate change impacts (van wilgen et al. 2016), while also impacting the livelihoods of people living outside of national parks. as such, with biodiversity threats increasing outside of parks, the resources within parks are increasingly important not only for conservation but also as resources for livelihoods and to meet social and cultural needs. while the formal unemployment rate is reported at 26.7%, non-participation in the labour force means that of 38 million people of working age in south africa, only 43.5% are employed (stats sa 2018). this extremely high level of unemployment increases reliance on natural resources, which can serve as an entry point into the informal economy in terms of the traditional medicine trade (petersen et al. 2014). in general, people or communities living near more isolated or rural parks tend to have limited employment opportunities compared to urban parks. table mountain national park (tmnp), for example, is situated within the city of cape town, where 3.7 million people (growing at ~2%; 2011 census data) live on the park periphery. this calls for a relatively strict control and regulation of resource harvesting from the park. the tmnp-mpa is located adjacent to communities for whom fishing forms part of their livelihoods and who are in competition with more expansive commercial operators for the limited resources. resentment over the allocation of fishing licenses and quotas (not administered by sanparks) contributes to illegal resource harvesting, including rock lobster and abalone. overall, good progress has been made with the co-management of resources within parks (e.g. the harvesting of mopane worms, thatching grass and sour figs in some parks) (scheepers, swemmer & vermeulen 2011; swemmer et al. 2015a). however, the same cannot be said of co-management of parks, with the exception of the richtersveld national park that is situated in a very remote and sparsely populated part of the country, typified by desert landscapes with very few permanent residents. this is also the only park that is jointly managed by sanparks and local communities in its entirety, with co-owners having considerable authority and responsibility. livestock graze freely in the park, providing an important contribution to local livelihoods through resource access. the makuleke concession agreement in the knp also claims some successful components of co-management, while the khomani san co-management agreement in the kgalagadi transfrontier park has yielded few benefits for the majority of the san. the knp has a particularly complex history with a legacy of forced removals of residents to outside of the park during its establishment as well as policies of restricted access to certain cultural groups. furthermore, a large portion of the western boundary of the park is bordered by former homelands (bantustans) where thousands of people were forcibly moved to live from other areas in the country. as such, not only does the knp have approximately 2 million people living within the eight municipalities that border the western boundary of the park, but the 1074 km of perimeter is adjoined by segments of industry, urban settlements, and private, communal and public nature reserves, as well as state-owned communal land. unemployment rates in all eight adjacent municipalities are higher than the national average (swemmer & mmethi 2017). as such, many people living next to the park rely on natural resources (shackleton 2000). this offers significant opportunity for positive well-being and constituency building as a result of controlled resource use from the park. the vast area covered by the various parks, the wide variety of biodiversity within them and the complexity of the range of stakeholders involved mean that there are a large number of resources within parks that are sought for a variety of purposes (van wilgen et al. 2013). defining resource use resource use is either consumptive or non-consumptive, while indirect use-values (e.g. ecosystem services such as soil and water conservation, genetic resources and landscape aesthetics) are also recognised (lawes, obiri & eeley 2004). non-consumptive use refers to the passive or intangible value of resources and includes activities such as the tourism experience (often commercial ventures) and cultural practices of a non-extractive nature. consumptive use implies the removal or withdrawal of all or part of the natural resource from its origin. natural resources are both biological and renewable (i.e. able to be replenished in the course of natural events within the limits of human time) or abiotic and non-renewable (i.e. cannot be produced, grown or generated on a scale that can sustain their consumption rate) (symonds 2010). the extractive use of biological resources is broad in scope and includes parts of individual plants and animals such as leaves and bark, or by-products from animal activities (e.g. honey), as well as entire plants or animals and their different life-history stages, such as seeds, flowers and eggs (lawes et al. 2004), each of which has a unique social, economic and ecological impact. consumptive use may either be for commercial gain (e.g. game sales, to support the tourism industry and fishing) or for domestic (often traditional) purposes, such as subsistence and recreational use (e.g. medicinal plant harvesting and recreational fishing), with sanparks and/or external stakeholders as the beneficiaries. consumptive use could also include the removal of plants and animals (or products thereof) as by-products of management actions or interventions (e.g. game stock reductions and plants and cultural artefacts collected for research purposes). the monitoring programme outlined in this article deals specifically with the consumptive use of biotic (living, e.g., thatching grass and fish) and abiotic (non-living, e.g., rocks and minerals) resources from terrestrial and aquatic ecosystems as the broader biodiversity monitoring framework was established within the context of direct threats to biological resources. the importance of non-extractive use of resources is acknowledged, but is beyond the scope of the programme and this article. legislative and policy context having policies, laws and institutions in place at all levels of governance with effective linkages between them is one of the addis ababa principles for the sustainable use of biodiversity (cbd 2004). relevant south african legislation, as well as international agreements and conventions relating to resource use, is provided in vermeulen et al. (2011). the south african national environmental management: protected areas act (act no. 57 of 2003) states that one of the purposes of protected areas is to provide for the sustainable use of natural and biological resources. various components of the marine living resources act (act no. 18 of 1998) and the international convention on biological diversity are also relevant. implementation of and compliance with legislation dealing with resource use is directed by internal sanparks policies, in particular, the sanparks resource use policy (sanparks 2010). some key principles that relate to this policy, and which potentially impact on monitoring, include the following: the consumptive use of renewable resources is allowed, provided that the use is sustainable and does not threaten the resource, nor any other resource, ecosystem functioning or element of biodiversity dependent on it. a complex system view is adopted and therefore a certain amount of change to ecosystems (including that brought about by resource use) is acceptable, provided that the change remains within the limits of natural variation exhibited by the ecosystem. south african national parks follows an approach of adaptive management based on (1) a combination of science and traditional or local knowledge, (2) iterative, timely and transparent feedback derived from adequately monitoring the resource and (3) adjusting the management of the resource based on feedback from monitoring. south african national parks supports interdisciplinary research (including the development of internal expertise and harvesting systems) into the ecological, social and economic opportunities and constraints of resource use. the effective implementation of the policy requires that resource use takes place according to principles of sustainability, and without compromising the integrity of ecosystems and other services that it sustains. monitoring system sustainability is of overriding importance in the context of sound conservation management. as such, the development of a robust, goal-orientated monitoring programme must consider the interdependent social, economic and ecological dimensions of sustainability, within a political or policy framework (cunningham 2001). the sanparks resource use monitoring programme is therefore structured in view of the fact that sustainable resource use is achievable only where all dimensions of sustainability are considered simultaneously and where investment in assessment and monitoring of these dimensions, including the trade-offs within and between them (swemmer et al. 2015a), is in place to ensure that sustainability objectives are achieved. ecological sustainability, as defined in the national environmental management: biodiversity act (act no. 10 of 2004), involves the use of a biological resource in a way and at a rate that does not lead to its long-term decline, does not disrupt the ecological integrity of the ecosystem in which it occurs and ensures its continued use to meet the needs and aspirations of present and future generations. social sustainability refers to social justice and equity and stresses community participation, paying particular attention to the most vulnerable people in society (dea 2002). economic sustainability refers to economic viability and integrity and focuses on economic growth that is viable and fair, and which occurs at a rate that does not exceed the ability of natural and social systems to support this growth (dea 2002). the sanparks resource use monitoring programme, described in detail by vermeulen et al. (2011), follows the generic process for harvest system development (figure 1) and is summarised in table 1. two case studies provide examples and challenges in programme implementation (see section ‘implementation and adaptive management’). the first case study deals with medicinal plants in the grnp and the second case study covers recreational and subsistence line fisheries. figure 1: flow diagram indicating the generic process for the development of harvest systems and management prescriptions for sustainable resource use. table 1: components of the south african national parks resource use monitoring programme for the three dimensions of sustainability. monitoring the social and economic dimension of sustainability the overall objective of this component of the programme is to (1) assess the extent to which sanparks meets legislative and internal objectives pertaining to the promotion of access to social and economic benefits from resource use and (2) to monitor the extent to which having access to resources impacts the perception of stakeholders regarding the value of conservation and protected areas. to this end, the programme entails assessing the resource use needs of key stakeholders, the status and trends of resource use in protected areas managed by sanparks, and the social and economic impact thereof (see table 1). also the development of alternative resources is of relevance not only to address needs where demand exceeds supply, but also to build more robust systems and reduce vulnerability. local stakeholder resource uses and needs baseline information on the history of the park and the people living in proximity to park, as well as local stakeholder needs for and uses of various natural resources, provides useful context with which conservation organisations can frame specific resource use projects and programmes. stakeholder needs and uses are dynamic; hence, ongoing engagement is required to maintain the relevance of resource use programmes in the context of changing needs. data to be collected would be context-specific depending on the type of resource, but some useful variables may include species (fauna and flora) used, parts harvested, the motivation for harvest, type of use (domestic or commercial), quantity used, product quality requirements, number of people to be involved in the harvesting, level of dependence on the resource and acceptability of alternative resources should demand exceed supply (see case study 1 in terms of medicinal plant harvesting and surveys with traditional healers). broad-scale status and trends in resource use broad-scale status of and trends in resource use are useful for assessing the degree to which strategic objectives for resource use are being met, as well as for long-term monitoring of resource use dynamics to inform an adaptive approach to management, specifically in relation to threat identification. the known status of resource use in sanparks is provided by van wilgen et al. (2013) and forms the baseline for monitoring long-term trends in resource use (in terms of species and products harvested, volumes harvested, user groups, etc.) as does the annual sanparks resource use report (see, e.g. symonds 2014 and 2018 for the reports for 2013/2014 and 2017/2018, respectively). broad-scale trend indicators to monitor the extent of resource use and the benefits accrued include quality and quantity of products harvested, satisfaction of participants, income generated and number of stakeholders directly involved. setting easy-to-measure species or group-specific indicators of quality is an important step when new projects are set up. for the forest fern rumohra adiantiformis, for example, it involved recording the number and length of ferns fronds with < 10% blemish, harvested during successive harvest cycles (vermeulen 2009). the programme also includes indices for illegal, unreported and uncontrolled harvesting (see case study 1). local-scale social and economic impact of resource use monitoring the simultaneous social and economic impact of resource use enables identification of costs and benefits at multiple stakeholder levels, as well as between stakeholders and the natural environment, in support of a net benefit outcome at most levels (swemmer & taljaard 2011; swemmer et al. 2015a). examples of social impacts include developing or maintaining social cohesion and continuity as well as changes that occur in people’s way of life (i.e. how they live, work, play and interact with one another on a daily basis), their culture (i.e. shared beliefs, customs and values) and their community (i.e. its cohesion, stability, character, services and facilities) (lahiri-dutt, nair & dowling 2008). not only does the sanparks resource use programme aim to promote access to resources but it also aims to build support for conservation through the creation of both positive relationships and vested interest in national parks (swemmer et al. 2017). monitoring the economic impact of resource use on the user at various scales (e.g. individual, household and broader scales) is equally important. identifying indicators for social and economic objectives is not easy because in most cases the outcomes are both quantitative and qualitative. detailed monitoring of the social and economic aspects is not always needed on an annual basis. numerous generic well-being indicators exist, but indicators are context-specific in that they must be seen as legitimate by all parties involved. this is best achieved through co-defining programme objectives and indicators through multiple stakeholder lenses together with resource harvesters (swemmer et al. 2015a). a lack of cohesion and formal representation of user groups, for example, limited inclusion of resource users in developing indicators within the grnp line fisheries (case study 2) and will require novel methods to ensure inclusivity in the future. in contrast, the rastafarian community was directly involved in the monitoring of bulbine latifolia, a plant used for medicinal purposes (case study 1) within the park. monitoring of the indicators is determined at a project level based on local context and should be based on needs, resources and expertise available. the use of community-based monitors has proven successful in natural resource-based programmes (swemmer et al. 2015b). for example, community monitors employed through south africa’s expanded public works’ environmental monitor programme patrol certain boundary fences of the knp, including areas where there have been historical incidents of illegal harvesting of high-value medicinal plants. the community monitors both come from and work in the areas adjacent to the park. despite advantages of using community monitors (local expertise, connections with community and buy-in to the project) funding for salaries can be a challenge (swemmer et al. 2015b). monitoring the ecological dimension of sustainability (biological resources) the overall objective of this component is to ensure the ecological sustainability of resource use by providing a scientific basis for harvest system development and refinement, and implementing relevant monitoring of key aspects of target species for continual assessment of and feedback to adaptive management. of specific relevance here are baseline data to support sustainable yield determination and harvest system development, including (1) species distribution and populations status (locally and nationally) and (2) rate of recruitment or replacement (i.e. rate of turnover) of the target species or product. harvest impact on the resource must be assessed to allow for harvest system refinement (table 1). although monitoring is an integral component of specific, formal resource use projects and concessions cognisance must be given to the potential impacts of illegal and unauthorised use of resources. where necessary, monitoring of relevant population dynamics should be compatible with and be able to feed into national monitoring programmes. this could be particularly important within the aquatic environment where fish stocks are assessed, and species-specific regulations are generated on a national basis (case study 2). rate of turnover and population dynamics of target species or products quantitative, field-based resource inventories or other relevant methods (e.g. life tables for animal populations, biomass estimates for fish and size–class distributions for timber – see table 1) can provide baseline data on how much of a resource targeted for use is available. data on the rate of production, which depend on the demography of the target species or the parts harvested, are crucial in determining the sustainable off-take of the available resource. for example, in the case of b. latifolia harvested for medicinal purposes in the grnp, the size–class distribution of the population and monitoring data on the growth rate of the species were used to formulate harvest prescriptions in terms of harvestable size and harvest rotation, reflecting the sustainable off-take, while rate of bark regrowth was used to assess options for sustainable medicinal bark harvesting (vermeulen 2009, case study 1). population transition and other mathematical models could also be used to determine the potential yield or to assess harvest impact (ngubeni 2015; pfab & scholes 2004) for harvest prescription development and refinement. where necessary (e.g. large mammals), ecological complexity should be incorporated into the decision-making process and population management should seek heterogeneity favouring biodiversity and ecosystem resilience (ferreira & hofmeyr 2011). indicators of turnover would depend on the species or products harvested, for example, diameter or length growth for whole plants, recruitment and regeneration (including coppice growth), fruit or leaf or bark production for specific plant parts and catch per unit effort for fish. harvest impact the impact of harvesting according to specific harvest intensities on the target species, as well as on other components of the ecosystem, is an important component of monitoring. two approaches are followed here: monitoring trends in quality and quantity of the product harvested from the same harvest area and the health of the population after harvesting. data collection depends on the specific species and product harvested and could include the number or volumes harvested (quantity) and size (or any other measures that define the quality, e.g., condition factor) of the individuals or products harvested. monitoring population health entails assessing the quantity and quality of the remaining product in the harvest area, as well as the population status as a whole. for example, in the grnp, monitoring included assessing tree response to bark stripping and susceptibility to insect and fungal damage following bark stripping (case study 1). in some instances, population parameter monitoring should be implemented within areas where resource use occurs, and also in ‘no-take’ areas to provide an indication of population trends independent of anthropogenic influence. for example, target populations of various fish species are known to show natural fluctuations in population abundance (götz et al. 2008), while a natural decline in r. adiantiformis fern populations has also been recorded (vermeulen 2009), which may need to be taken into consideration. monitoring the impact of harvesting of abiotic resources as abiotic resources are considered to be non-renewable, sustainable use (e.g. the harvesting of rock, gravel and sand from quarries) in this context refers to sound harvest and management practices to minimise the impact on the environment, and to ensure the responsible use of the resource in terms of its availability to optimise social and economic benefits. this includes assessing the needs and trends in the harvesting of non-renewable resources from national parks in terms of products, volumes and frequencies, as well as whether sound management practices and measures are in place to minimise environmental impact. in line with policy directives (sanparks 2010), the total available original quantity of the resource would need to be determined and its use should be curtailed such that only a minor portion of this original quantity is used and no ecosystem process or biodiversity is threatened by the use. implementation and adaptive management in terms of ecological sustainability, the programme has been implemented at different levels in different parks for specific resource use projects, while a baseline has been established to monitor trends in resource use and benefits accrued (swemmer & mmethi 2016; van wilgen et al. 2013). lessons learnt are that the precautionary principle should apply with the formulation of harvest prescriptions when limited monitoring and research data are available and that roles and responsibilities in project implementation are clearly defined, including that of external beneficiaries. also, new projects may fail if the necessary financial and human resources are not available to support the development of harvest prescriptions and associated social, economic and ecological monitoring. depending on the nature of the resource use project, different role-players in sanparks are responsible for the successful development and implementation of the programme. these include scientists (working in the biology and social science fields), park management staff and resource users. in line with sanparks strategic adaptive management approach (biggs & rogers 2003), management intervention could be required in terms of both the socio-economic (resource use trends, benefits accrued and social impact) and ecological dimensions (e.g. harvest impact) of sustainability, based on monitoring results. adaptive management lies on the concept of thresholds of potential concern (tpcs), which defines acceptable upper and lower levels of change (biggs & rogers 2003). such tpcs could also be developed for resource use to close the adaptive loop, and be linked to park management objectives. science–management interface meetings are the ideal platform for the development and refinement of such tpcs. this approach allows for the setting of conservative harvest levels where data deficiencies exist, to be refined as monitoring results become available. examples of applying adaptive management in resource use in sanparks are provided in the two case studies, but also include the harvesting of seven-weeks fern (r. adiantiformis) in the grnp, medicinal bark from the pepper-bark tree (warburgia salutaris) in the knp and sour fig (carpobrotus acinaciformis) in agulhas national park (scheepers et al. 2011). applying a holistic, sustainability-based focus on the application of strategic adaptive management of resource use (social, ecological and economic) has the potential to reduce the risk of resource use project failure. ensuring the maintenance of ecological integrity (through ecological harvest system development), promoting economic viability (by identifying, monitoring and managing the cost–benefit trade-offs within and between stakeholder groups) and establishing social relevance (by facilitating access to resources using governance processes and promoting shared decision-making) are key requirements. also, as lessons are learnt with the implementation of the monitoring programme, the programme itself could be adapted and refined to ensure that it remains appropriate and practical to adequately serve its purpose within a specific context. case study 1: demand and access for medicinal plant harvesting in the garden route national park context the overharvesting of resources has been identified as an international threat to biodiversity and ecosystem services and as such a driver of global change (van wilgen & herbst 2017). this is no different in sanparks where the demand for medicinal plants is increasing (van wilgen et al. 2013), particularly so in the grnp where demand for forest and fynbos medicinal plants species is exacerbated by the nature of the open access park, changes in stakeholder dynamics and commercialisation (vermeulen 2009). assessing demand and monitoring harvesting table 2 summarises garden route projects in terms of monitoring objectives, methods, frequency of monitoring and important indicators. a baseline inventory of medicinal plant harvesting was provided by van wilgen et al. (2013), while local traditional healers were surveyed to determine their requirements for plant species in the region (ngubeni 2015). medicinal tree bark was identified as a high priority, considering the demand for bark for private and commercial use, and the destructive nature of strip harvesting. tree response to bark stripping and the rate of bark regrowth were assessed (vermeulen 2009). monitoring of b. latifolia, also a species in high demand and subjected to overharvesting, was aimed at assessing the rate of growth and population turnover to inform harvest prescriptions for sustainable use (vermeulen 2009). this monitoring was conducted in consultation with the rastafarian community as an important user group. a project was initiated to monitor the increasing unauthorised use of medicinal and other plants (vermeulen et al. 2015). table 2: monitoring projects and surveys relating to the sustainable harvesting of medicinal plant products from the garden route national park. summary of results a wide range of medicinal plant species and products are in demand (ngubeni 2015; vermeulen 2018). users seem to be receptive to exploring alternatives to assist sustainability (ngubeni 2015). harvest prescriptions have been formulated for high-demand species such as b. latifolia, which is slow growing with a low rate of turnover of populations in the wild (vermeulen 2009). however, a sustainable supply of b. latifolia can be provided by growing this in gardens and engagements are underway with stakeholders to facilitate this. for medicinal tree bark, harvest prescriptions have been developed for ocotea bullata, but for most other species, little scope exists for sustainable strip harvesting because of poor bark regrowth, and susceptibility to fungal and insect damage (vermeulen 2009; vermeulen, geldenhuys & esler 2012). harvesting bark as by-product from timber harvesting or full-tree harvesting under sustainable timber harvesting systems (see seydack et al. 1995) would be more viable options than strip harvesting. management actions and challenges managing unauthorised resource use in a fragmented and open access park, such as the grnp, requires pro-active engagement with stakeholders together with law enforcement and vigilant monitoring of population health. improved understanding of stakeholder dynamics and needs enable research and monitoring to focus on priority species and products, and assess to what extent needs could realistically be addressed. achieving inclusive engagement with all plant harvesters is key but difficult because many are not members of a user group. park management needs to drive a more structured effort to identify stakeholders because the demand for medicinal plant products is expected to increase. case study 2: monitoring the recreational and subsistence line fisheries occurring within the garden route national park context recreational fishing managed nationally under the marine living resources act (1998) has been primarily top-down and subsistence (or small-scale) fisheries, in particular, were historically marginalised and neglected within policies and management systems (sowman 2006). recreational and subsistence fishing occurs within the grnp, but little information about participants or trends in fishing effort and catches was available to management. monitoring related to recreational and subsistence fishing to evaluate the sustainability of marine and estuarine fishing, a monitoring programme was implemented in 2008. two estuaries and two coastal sections were selected as monitoring sites through a prioritisation exercise. objectives, key questions and hypotheses were co-developed by scientists and park management. sustainability indicators along with trends, performance criteria and tpcs were set within the ecological and social domains. indicators were standard between estuarine systems and coastal areas but reference points were area specific. for example, in estuarine systems, targeted catch per unit effort (cpue) for cape stumpnose is an indicator but the tpc differs between systems. feedback and challenges the programme provides spatially explicit information including catch rates, proportion of different user groups, cpue, retention rates, proportion of undersized fish, number of patrols and annual estimates (e.g. total fishing effort and total catch). results are fed into the adaptive management loop and used to guide management interventions and identify further research questions (figure 2). for example, monitoring highlighted a large proportion of undersized fish being retained, and as a result, research is being conducted regarding the drivers of angler behaviour and non-compliance, while management increased its emphasis on law enforcement. similarly, poor angler awareness and knowledge of fishery regulations resulted in management interventions, including the development of illustrative and locally relevant fishery signboards, pocket field guides and brochures. the influence of these interventions is currently being re-assessed on a 5-year basis. figure 2: example of how the monitoring framework has been adopted in the garden route national park recreational and subsistence line fisheries. text in red shows how areas of concern (identified through monitoring key indicators) were addressed in management actions. the impact of these actions is then re-assessed through the monitoring programme. currently, the monitoring programme is in partial compliance with the desired resource use monitoring system. the impact on the targeted populations and biodiversity is less well covered and, in particular, social and economic indicators need to be developed in collaboration with the local anglers. future research and monitoring should strengthen the interdisciplinary nature of the work, with the aim of gaining greater understanding on how recreational and subsistence fisheries respond to socio-ecological change. of particular importance is further engagement with local fishermen and enabling their input into the monitoring framework and future refinements. this is a challenge as most anglers do not belong to a club and, in particular, subsistence fishermen are not well organised or represented. conclusion the monitoring programme needs to be supported by applied research on selected target species and products to ensure sustainable use and optimise benefits. the resources required to develop harvest systems for sustainable use and to conduct initial and ongoing monitoring to support this are often underestimated (cunningham 2001; vermeulen 2009). costing for a specific project would be possible, but a detailed costing of the programme would be difficult as this would be influenced by, for example, diversity of species and products harvested, number of users and stakeholder dynamics, further impacted on by the precision and accuracy of inventories and monitoring required, and the spatial and temporal scale at which monitoring needs to be conducted (cunningham 2001). constraints on the availability of resources (financial and human resources) could potentially hamper the successful implementation of the programme. in many cases (as described in the different examples and the two case studies), components of the programme have already been implemented, integrated with park management activities and as part of the established sanparks research and monitoring programme (sanparks 2014). however, prioritising monitoring and resource use projects is essential. these are influenced by sociopolitical or institutional factors (including political and societal values and regulatory frameworks for resource use), economic factors (such as the economic status of local communities and the demand for and value of the resource), as well as user group and local community factors (e.g. the nature of the demand and history of use, dependence on the resource and the availability of alternatives). also, the ecological factors such as the status and availability of a resource and the characteristics of the target species are of importance (sanparks 2011). considering the long-term nature of the monitoring programme, the development of in-house skills and expertise is essential, while partnering with research institutions such as universities, is also important. in terms of commercial resource use projects, the costs of monitoring and research should ideally be covered by the commercial venture. acknowledgements the authors would like to thank sanparks for providing financial assistance for the project. they also thank two anonymous reviewers for their comments and input that helped improve the manuscript. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions w.v., together with n.v.w., k.s., m.d., l.s. and w.a., conceptualised the article. w.v. drafted the manuscript with input from n.v.w., k.s., m.d., l.s. and w.a. further comments and editorial input for the article were provided by h.b., g.d., n.h., h.h., m.m., n.n. and a.s. all authors formed part of a working group that developed the monitoring system. references biggs, d., swemmer, l., phillips, g., stevens, j., freitag, s. & grant, r., 2014, ‘the development of a tourism research framework by south african national parks to inform management’, koedoe 56(2), art. #1164, 1–9. https://doi.org/10.4102/koedoe.v56i2.1164 biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring, and management in practice’, in j.t du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59–80, island press, washington, dc. butchart, s.h.m., walpole, m., collen, b., van strien, a., scharlemann, j.p.w., almond, r.e.a. et al., 2010, ‘global biodiversity: indicators of recent declines’, science 328, 1164–1168. https://doi.org/10.1126/science.1187512 cbd, 2004, addis ababa principles and guidelines for the sustainable use of biodiversity, secretariat of the convention of biological diversity, montreal. clarke, j. & grundy, l.m., 2004, ‘the socio-economics of forest and woodland resource use: a hidden economy’, in m.j. lawes, h.a.c. eeley, c.m. shackleton & b.g.s. geach (eds.), indigenous forests and woodlands in south africa. policy, people and practice, pp. 167–193, university of kwazulu-natal press, pietermaritzburg. cunningham, a.b., 2001, applied ethnobotany. people, wild plant use and conservation, people and plants conservation manuals, earthscan publications, london. department of energy, 2013, a survey of energy-related behaviour and perceptions in south africa. the residential sector, department of energy, pretoria. department of environmental affairs and tourism (now department of environmental affairs), 2002, national framework document: strengthening sustainability in the integrated development planning process, deat, pretoria. ervin, j., sekhran, n., dinu, a., gidda, s., vergeichik, m. & mee, j., 2010, protected areas for the 21st century: lessons from undp/gef’s portfolio, united nations development programme and montreal: convention on biological diversity, new york. ferreira, s.m. & hofmeyr, m., 2011, reconciling conservation objectives through wildlife sales as an ecosystem service, unpublished report, south african national parks, skukuza. götz, a., kerwath, s.e., attwood, c.g. & sauer, w.h.h., 2008, ‘effects of fishing on population structure and life history of roman chrysoblephus laticeps (sparidae)’, marine ecology progress series 362, 245–259. https://doi.org/10.3354/meps07410 lahiri-dutt, k., nair, a. & dowling, s., 2008, social impact assessment: a manual for mining projects, resource management in asia pacific programme, australian national university, canberra. lawes, m.j., obiri, j.a.f. & eeley, h.a.c., 2004, ‘the uses and value of indigenous forest resources in south africa’, in m.j. lawes, h.a.c. eeley, c.m. shackleton & b.g.s. geach (eds.), indigenous forests and woodlands in south africa. policy, people and practice, pp. 227–273, university of kwazulu-natal press, pietermaritzburg. mander, m., ntuli, l., diederichs, n. & mavundla, k., 2007, economics of the traditional medicine trade in south africa, report for ezemvelo kzn wildlife, futureworks!, durban. mcgeoch, m.a., dopolo, m., novellie, p., hendriks, h., freitag, s., ferreira, s. et al., 2011, ‘a strategic framework for biodiversity monitoring in south african national parks’, koedoe 53(2), art. #991, 1–10. https://doi.org/10.4102/koedoe.v53i2.991 millennium ecosystem assessment, 2005, ecosystems and human well-being: synthesis, island press, washington, dc. ngubeni, n., 2015, ‘bark re-growth and wood decay in response to bark stripping for medicinal use’, msc thesis, department of conservation ecology and entomology, stellenbosch university, stellenbosch. ouédraogo, a.-s., 2001, ‘conservation, management, and use of forest genetic resources’, in o. eyog-matig, b. kigomo & j.-e. boffa (eds.), recent research and development in forest genetic resources, proceedings of the training workshop on the conservation and sustainable use of forest genetic resources in eastern and southern africa, december 6–11, 1999, pp. 1–14, international plant genetic resources institute, nairobi. pauly, d., 2008, ‘global fisheries: a brief review’, journal of biological research-thessaloniki 9, 3–9. paumgarten, f., locatelli, b. & witkowski, e.t.f., 2018, ‘wild foods: safety net or poverty trap? a south african case study’, human ecology 46, 183–195. https://doi.org/10.1007/s10745-018-9984-z peters, c.m., 1996, the ecology and management of non-timber forest resources, world bank technical paper no. 322, the world bank, washington, dc. petersen, l.m., charman, a.j.e., moll, e.j., collins, r.j. & hockings, m.t., 2014, ‘“bush doctors and wild medicine”: the scale of trade in cape town’s informal economy of wild-harvested medicine and traditional healing’, society & natural resources 27, 315–336. https://doi.org/10.1080/08941920.2013.861558 pfab, m.f. & scholes, m.a., 2004, ‘is the collection of aloe peglerae from the wild sustainable? an evaluation using stochastic population modeling’, biological conservation 118, 695–701. https://doi.org/10.1016/j.biocon.2003.10.018 roux, d.j. & nel, j.l., 2013, ‘freshwater conservation planning in south africa: milestones to date and catalysts for implementation’, water sa 39, 151–164. https://doi.org/10.4314/wsa.v39i1.15 scheepers, k., swemmer, l. & vermeulen, w.j., 2011, ‘applying adaptive management in resource use in south african national parks: a case study approach’, koedoe 53(2), art. #999, 1–14. https://doi.org/10.4102/koedoe.v53i2.999 seydack, a.h.w., vermeulen, w.j., heyns, h.e., durrheim, g.p., vermeulen, c., willems, d. et al., 1995, ‘an unconventional approach to timber yield regulation for multi-aged, multispecies forests. ii. application to a south african forest’, forest ecology and management 77, 155–168. https://doi.org/10.1016/0378-1127(95)03578-x shackleton, c. & shackleton, s., 2004, ‘the importance of non-timber forest products in rural livelihood security and as safety nets: a review of evidence from south africa’, south african journal of science 100, 658–664. shackleton, c.m., 2000, ‘comparison of plant diversity in protected and communal lands in the bushbuckridge lowveld savanna, south africa’, biological conservation 94, 273–285. https://doi.org/10.1016/s0006-3207(00)00001-x shackleton, s., shanley, p. & ndoye, o., 2007, ‘invisible but viable: recognising local markets for nontimber forest products’, international forestry review 9(3), 697–712. https://doi.org/10.1505/ifor.9.3.697 south african national parks (sanparks), 2010, sanparks resource use policy, ref. 17/p – csd/pol/resource use/03-10/vs 1, sanparks, pretoria. south african national parks (sanparks), 2011, notes on work session to explore and compile criteria for prioritizing particular resource uses in parks, sanparks resource use working group workshop ii, 7–8 september, cape research centre, steenberg. south african national parks (sanparks), 2014, sanparks research report 2013, sanparks, pretoria. south african national parks (sanparks), 2016a, south african national parks strategic plan 2016/17–2019/20, sanparks, pretoria. south african national parks (sanparks), 2016b, mountain zebra national park. park management plan for the period 2016–2026, sanparks, pretoria. sowman, m., 2006, ‘subsistence and small-scale fisheries in south africa: a ten-year review’, marine policy 30, 60–73. https://doi.org/10.1016/j.marpol.2005.06.014 spear, d., foxcroft, l.c., bezuidenhout, h. & mcgeoch, m.a., 2013, ‘human population density explains alien species richness in protected areas’, biological conservation 159, 137–147. https://doi.org/10.1016/j.biocon.2012.11.022 stats sa, 2017, mid-year population estimates 2017, statistical release p0302, statistics south africa, pretoria. stats sa, 2018, youth unemployment still high in q1: 2018, statistics south africa, viewed 14 august 2018, from http://www.statssa.gov.za. swemmer, l.k., grant, r., annecke, w. & freitag-ronaldson, s., 2015a, ‘toward more effective benefit sharing in south african national parks’, society & natural resources 28(1), 4–20. https://doi.org/10.1080/08941920.2014.945055 swemmer, l.k. & mmethi, a.h., 2016, biodiversity for society – a reflection on the diversity of direct, local impacts (benefits and costs) of the kruger national park, kruger national park, south african national parks, pretoria. swemmer, l.k., mmethi, h. & twine, w., 2017, ‘tracing the cost-benefit pathway of protected areas, a case study of the kruger national park, south africa’, ecosystem services 28, 162–172. https://doi.org/10.1016/j.ecoser.2017.09.002 swemmer, l.k. & taljaard, s., 2011, ‘sanparks, people and adaptive management: understanding a diverse field of practice during changing times’, koedoe 53(2), art. #1017, 1–7. https://doi.org/10.4102/koedoe.v53i2.1017 swemmer, l.k., uys, m.t., marshall, t., malepe, m., morale, k. & buthelezi, i., 2015b, the social and economic impact of the kruger to canyon biosphere environmental monitor programme – impact on the environmental monitors, sanparks scientific report 05/2015. symonds, a., 2010, towards a common understanding of resource use in sanparks, internal memorandum, south african national parks, pretoria. symonds, a., 2014, report on the sustainable use of biological resources. 1 april 2013 – 31 march 2014. annual return to the minister of the department of environment affairs in compliance with section 7 of the nem:paa regulations for the proper administration of special nature reserves, national parks and world heritage sites, south african national parks, pretoria. symonds, a., 2018, report on the sustainable use of biological resources. 1 april 2017 – 31 march 2018. annual return to the minister of the department of environment affairs in compliance with section 7 of the nem:paa regulations for the proper administration of special nature reserves, national parks and world heritage sites, south african national parks, pretoria. ticktin, t., 2004, ‘the ecological implications of harvesting non-timber forest products’, journal of applied ecology 41, 11–21. https://doi.org/10.1111/j.1365-2664.2004.00859.x twine, w., 2011, ‘drivers of natural resource use by rural households in the central lowveld’, in h.l. zietsman, j. pauw, a. van jaarsveld & a. saidi (eds.), observations on environmental change in south africa: causes, consequences and responses, pp. 17–19, sun press, pretoria. uhunamure, s.e., nethengwe, n.s. & musyok, a., 2017, ‘driving forces for fuelwood use in households in the thulamela municipality, south africa’, journal of energy in southern africa 28, 25–34. https://doi.org/10.17159/2413-3051/2017/v28i1a1635 van wilgen, n.j., dopolo, m., symonds, a., vermeulen, w., bester, e., smith, k. et al., 2013, ‘an inventory of natural resources harvested from national parks in south africa’, koedoe 55(1), art. #1096, 1–5. https://doi.org/10.4102/koedoe.v55i1.1096 van wilgen, n.j., goodall, v., holness, s., chown, s.l. & mcgeoch, m.a., 2016, ‘rising temperatures and changing rainfall patterns in south africa’s national parks’, international journal of climatology 36, 706–721. https://doi.org/10.1002/joc.4377 van wilgen, n.j. & herbst, m. (eds.), 2017, taking stock of parks in a changing world: the sanparks global environmental change assessment, south african national parks, cape town. van wilgen, n.j. & mcgeoch, m.a., 2014, ‘balancing effective conservation with sustainable resource use in protected areas: precluded by knowledge gaps’, environmental conservation 42(3), 246–255. https://doi.org/10.1017/s0376892914000320 van wyk, b.e., van oudtshoorn, b. & gericke, n., 2009, medicinal plants of south africa, briza, pretoria. vermeulen, w., dopolo, m., bezuidenhout, h., durrheim, g., hanekom, n., hendricks, h. et al., 2011, south african national parks biodiversity monitoring programme: resource use (consumptive), scientific report 08/2011, sanparks, knysna. vermeulen, w.j., 2009, ‘the sustainable harvesting of non-timber forest products from natural forests in the southern cape, south africa: development of harvest systems and management prescriptions’, phd dissertation, department of conservation ecology and entomology, stellenbosch university, stellenbosch. vermeulen, w.j., 2018, incidences of illegal harvesting of terrestrial plants from the garden route national park, garden route scientific services, knysna, south african national parks. unpublished report. vermeulen, w.j., baard, j.a., durrheim, g.p., kraaij, t., ngubeni, n. & van der vyver, l., 2015. monitoring of environmental indicators in the garden route national park: terrestrial ecosystems, scientific services, south african national parks, knysna. vermeulen, w.j., geldenhuys, c.j. & esler, k.j., 2012, ‘response of ocotea bullata, curtisia dentata and rapanea melanophloeos to medicinal bark stripping in the southern cape, south africa: implications for sustainable use’, southern forests 74(3), 183–193. https://doi.org/10.2989/20702620.2012.717384 koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research major vegetation types of the soutpansberg conservancy and the blouberg nature reserve, south africa abstract the major megetation types (mvt) and plant communities of the soutpansberg centre of endemism are described in detail, with special reference to the soutpansberg conservancy and the blouberg nature reserve. phytosociological data from 442 sample plots were ordinated using a detrended correspondence analysis (decorana) and classified using two-way indicator species analysis (twinspan). the resulting classification was further refined with table-sorting procedures based on the braun–blanquet floristic–sociological approach of vegetation classification using megatab. eight mvt’s were identified and described as eragrostis lehmanniana var. lehmanniana–sclerocarya birrea subsp. caffra blouberg northern plains bushveld, euclea divinorum–acacia tortilis blouberg southern plains bushveld, englerophytum magalismontanum–combretum molle blouberg mountain bushveld, adansonia digitata–acacia nigrescens soutpansberg arid northern bushveld, catha edulis–flueggia virosa soutpansberg moist mountain thickets, diplorhynchus condylocarpon–burkea africana soutpansberg leached sandveld, rhus rigida var. rigida–rhus magalismontanum subsp. coddii soutpansberg mistbelt vegetation and xymalos monospora–rhus chirendensis soutpansberg forest vegetation. keywords: phytosociology, vegetation ecology, grassland, savanna, thicket the soutpansberg conservancy (sc) and the blouberg nature reserve (bnr) reveal extremely rich diversities of plant communities relative to the sizes of these conservation areas (van wyk & smith 2001). although van rooyen and bredenkamp (1996) recognised this diversity and unique composition of plant communities within the soutpansberg–blouberg complex, the lack of detailed research in the region forced them to merge the area’s vegetation under the broad term of soutpansberg arid mountain bushveld. acocks (1953) recognised four different veld types for the greater surrounding region and described them as arid sweet bushveld, mixed bushveld, sourish mixed bushveld and sour bushveld. most of these veld types were described as heterogeneous (acocks 1953), comprising many sub-communities with different agricultural and production potentials. in addition to the savanna vegetation of the area, geldenhuys and murray (1993) and lubke and mckenzie (1996) described and mapped the patches of afromontane forest associated with the region. van wyk and smith (2001) only briefly mention the occurrence of “fynbos-type” vegetation along the summit of the mountain. they also refer to dense, almost monospecific stands of lebombo ironwood (androstachys johnsonii) on the arid northern slopes of the mountain. due to major gaps in the available vegetation data, no attempt has yet been made to synthesise, classify and to describe the plant communities of this region. the study area location the bnr (7 000 ha) and sc (100 000 ha) are located in the arid northern regions of the limpopo province of south africa (fig. 1). for the purpose of this study the sc is defined as a section of the soutpansberg mountain range contained by four provincial and national roads. the eastern boundary (e29º55’) is the n1 from makhado in the south to wyllies poort in the north. the northern boundary (s22º52’) is the r523 road from wyllies poort in the east to kalkheuwel in the west. the western boundary (e29º15‘) is the r521 road from kalkheuwel in the north to vivo in the south. the southern boundary (s23º12’) is the r522 road from vivo in the west to makhado in the east. from east to west it spans approximately 70 km and from north to south approximately 25 km at its widest. the bnr lies approximately 40 km to the west, situated along the eastern half of the blouberg mountain. topography the sc and bnr are part of the blouberg–soutpansberg mountain range, with ene–wsw orientation. although the blouberg and soutpansberg belong to the same geological formation they are referred to as separate entities. successive faulting along the tshamuvhudzi, kranspoort, nakab and zoutpan strike-faults, followed by the northwards tilting of the theo h.c. mostert george j. bredenkamp hannes l. klopper cornie verwey 1african vegetation and plant diversity research centre department of botany university of pretoria south africa rachel e. mostert directorate nature conservation gauteng department of agriculture conservation and environment south africa norbert hahn1 correspondence to: theo mostert e-mail: tmostert@tuks.co.za postal address: african vegetation and plant diversity research centre, department of botany, university of pretoria, pretoria, 0002 32 vol. 50 no. 1 pp. 32 48 2008 major vegetation types original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe area, created these quartzite mountains within the surrounding limpopo plain. this gave the blouberg–soutpansberg mountain range a wedge-shaped appearance with steep southern slopes and moderate northern slopes. the ridges are highest at the western extremity of this range, which gradually descend until they finally plunge beneath the karoo supergroup along the northern reaches of the lebombo mountains near the northeastern border of the limpopo province. the sc’s altitude ranges from 750 m above sea level at waterpoort to 1 748 m at lejuma. the bnr’s altitude ranges from 850 m above sea level in the east to 1 400 m in the west. the highest peak of the blouberg lies further to the west, and reaches 2 051 m above sea level (bumby 2000). the surrounding plains are approximately 850 m above sea level. geology and soils the geology of the sc and bnr is dominated by pink, erosionresistant quartzite, and sandstone, with minor pebble washes of the wyllies poort geological formation of the soutpansberg group. other less prominent rock types include shale, conglomerate, basalt and diabase intrusions. the rocks of the study area do not contain large amounts of minerals that are of economic value. soils derived from quartzite and sandstone are generally shallow, gravelly, skeletal and well drained, with low nutrient content and acidic characteristics. soils derived from the basalt and diabase dykes are fine-textured, clayey, well weathered, and generally deep. these poorly drained soils are prone to erosion along the higher rainfall southern slopes. soils derived from the aeolian kalahari sands are fine-grained deep sands. large areas along the northern slope contain no soil, and comprise only the exposed underlying mother material. peat soils occur along the cooler high lying wetlands of the sc. the deeper soils within the mistbelt act as sponge areas, which slowly release water to feed mountain streams over extended periods. climate the sc and bnr fall within the summer rainfall zone of southern africa. a climate diagram (fig. 2) depicts typical rainfall patterns and temperature gradients within the study area. due to the east-west orientation of the soutpansberg it experiences orographic rainfall. this phenomenon is due to moisture-laden air from the indian ocean, driven by the prevailing south-easterly winds into the southern scarp of the soutpansberg. the north-south orientated wolkberg– drakensberg escarpment further blocks the westerly movement of the atmospheric moisture, forcing it into the wedge created by the two mountain systems in the vicinity of entabeni. large amounts of rain are discharged onto these southern slopes of the soutpansberg and eastern slopes of the wolkberg– drakensberg escarpment. entabeni receives an annual rainfall of 1 874 mm. orographic mist along this southern slope may increase annual precipitation to 3 233 mm (hahn 2002; olivier & rautenbach 2002). this creates a rain-shadow effect along the western slopes of the wolkberg–drakensberg escarpment and the northern slopes of the soutpansberg. with the sc and bnr located northwest of the soutpansberg–wolkberg– drakensberg escarpment junction, a double rain-shadow effect is experienced along the northern slopes of the study area. waterpoort, located north of the soutpansberg, receives only 367 mm rain annually. due to the extreme topographic diversity and altitude changes over short distances within the study area, the climate (especially rainfall and mist precipitation) varies dramatically. the amount of orographic rain associated with the southern ridges varies considerably in accordance to the changing landscape. the venturi effect caused by certain narrow gorges when mist is forced through them by orographic, anabatic and catabatic winds can lead to abnormally high localised rainfall (hahn 2002; matthews 1991). the areas just below the escarpment crest, where atmospheric moisture can be trapped most effectively against the south-facing escarpment, generally yield the highest precipitation (matthews 1991). 33 figure 1 the location of the blouberg nature reserve and western soutpansberg conservancy within south africa. a weather station name j average daily temp fluctuation b altitude k average monthly temp c number of years recorded [temp – rainfall] l average monthly precipitation d average annual temp (ºc) m dry season e average annual rainfall (mm) n wet season f average daily min (coldest month) o rainfall axis g lowest temp recorded p temperature axis h average daily max (warmest month) q time axis (months) i highest temp recorded legend figure 2 climate diagram for the alldays/waterpoort area sa nd r iv e r mu tam ba riv erho ut r ive r li mpopo ri ver nz hel ele ri ver w estern soutpansberg cons ervancy blouberg mountain range blou berg na ture reserve soutpansberg mountain range bra k r ive r 0 50 100 150 200 kilometers n study area w o lk b e rg d ra k e n s b e rg e s c a rp m e n t gauteng limpopo mpumalanga south africa northern c ape w estern cape eastern cape free state kwazulu-natal north-west lesotho n vol. 50 no. 1 pp. 32 48 original research mostert, bredenkamp, klopper, verwey, mostert & hahn koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za the diversity of rainfall in the study area is reflected in the long-term average rainfall recorded for three farms, namely ventersdorp, hanglip and schyffontein with annual rainfall figures of 585 mm, 774 mm and 835 mm respectively (south african weather bureau 2004). apart from the spatial variation in rainfall, the area reveals a high temporal variation in recorded rainfall (geldenhuys & murray 1993). mean annual rainfall for makhado fluctuated between 571 mm for the period 1965–1971 and 1027 mm for the period 1979–1988 (south african weather bureau 2004). in addition to these rainfall figures, the amount of precipitation as a result of mist can be substantial (schutte 1971). in the higher lying areas of the kwazulu-natal drakensberg, for example, the orographic fog contribution at 1 800 m altitude is an additional 403 mm per annum, which amounts to one third of the mean annual precipitation (matthews 1991). as with many mountainous areas, the daily weather of the higher altitude crests and summits of the soutpansberg is very unpredictable; it fluctuates between extremes within a matter of hours. the higher lying crests and ridges within the mistbelt are exposed to strong winds. during the summer months these winds carry moisture in from the indian ocean, creating a seasonal mistbelt, which gives rise to an abundance of rock and bark lichens and bryophytes. the combination of frequent orographic rain and mist during the summer months leaves the available soil drenched, and sometimes flooded for extended periods. during the extended dry season the prevailing winds are dry, causing dehydration and desiccation of the soils and vegetation (hahn 2002). these extreme and fluctuating environmental conditions have led to specialisation among the plants and may explain the relatively high level of endemisity within this vegetation type (hahn 2002). soutpansberg centre of biological diversity the soutpansberg–blouberg region has been recognised as a centre of endemism by van wyk and smith (2001). however, little ecological knowledge of the area exists (anderson 2001, berger et al. 2003). some floristic surveys conducted by hahn (1994, 1996 1997, 1999, 2002), stirton (1982), obermeyer et al. (1937) and van wyk (1984, 1996) indicated that the soutpansberg centre of endemism is exceptionally diverse and species-rich for its size (van wyk & smith 2001). the conservation value of this centre lies in its unique ability to house a wide variety of floristic elements from the surrounding floristic regions (hahn 2002). the region is an outstanding centre of plant diversity, with approximately 2 500–3 000 recorded vascular plant taxa (hahn 1997). according to van wyk and smith (2001) 41% of genera and 68% of families occurring in southern africa are represented within the soutpansberg centre of endemism. altogether 595 specific and infra-specific trees and shrubs are known from the soutpansberg, amounting to one third of all the known tree species in the entire southern africa region (hahn 1994, 1997, 2003). this constitutes one of the highest tree counts in southern africa (hahn 1997). the kruger national park, which covers an area of two million hectares, contains approximately 380 tree species (van wyk 1994), whereas 321 tree species have been recoded by hahn (2002) in an area of only 2 000 hectares within the sc. trees and shrubs accounts for approximately 24% of the vascular plants of the soutpansberg and play an important role in the species composition, vegetation structure and relative dominance within the different plant communities. more than 500 bird species have been recorded throughout the soutpansberg mountain range, amounting to approximately 56% of the recorded species for the entire southern africa (harrison et al. 1997; hockey et al. 2005). the soutpansberg and its surroundings have some unique reptile habitats, and seven endemic species are found there (branch 1988). a total of 46 spider families, 110 genera and 130 species have been recorded in the sc on the farm lejuma (< 50 km2), which represent 70% of the families, 26% of the genera and 5% of the species recorded for south africa (foord et al. 2002, 2003). the high biological diversity of the soutpansberg and blouberg can possibly be attributed to the fact that the mountain range acts as a refuge in times of environmental flux (hahn 2003). in the light of the high diversity recorded for the blouberg– soutpansberg expanse, it is proposed that the region be given the status of the soutpansberg centre of biological diversity (scbd) in addition to its recognised status as the soutpansberg centre of plant endemism. there are currently efforts underway to create a biosphere reserve in this area, which will include the sc and bnr (hahn in prep.). aims of the study this study is a first attempt at understanding the complex ecological patterns and processes observed within the scbd. it is concerned with the phytosociology and synecology of the sc and bnr. the aim is to identify the different plant communities and to investigate the interrelationships between plant communities and their physical and biological environments. in an attempt to create a holistic image and to explain the macro-ecology of the region, disciplines such as climatology, geology, pedology, physical geography, history and anthropology are drawn upon and integrated. this study provides a first approximation of the vegetation and proposes nine mvt’s for the study area. it aims to define and describe the characteristics of these mvt’s within the context of the sc and bnr. this will assist scientists, conservationists and land-use planners when future projects are conducted within the surrounding areas. these plant communities from the sc and bnr will serve as reference sites with which to compare proposed development sites from the surrounding unprotected areas. environmentally sound development is the ethical responsibility to base decisions regarding resourse utilisation on all available information in order to make holistical and long term sense (siebert 2001). this can be achieved if basic information, such as this account, is actively drawn upon during the planning phases of development and the management of natural resources. methods aerial photographs (scale 1:50 000) were used to stratify the study area into physiographic–physiognomic units. a total of 466 sample plots were randomly placed within each of these stratified units. the sample plot size was set at 400 m2 in accordance with vegetation studies elsewhere in the semiarid environments of southern africa (siebert 2001). the cover-abundance value for every species recorded within each sample plot was assessed according to the braun-blanquet cover-abundance scale (mueller-dombois & ellenberg 1974). the taxon names conform to those of germishuizen and meyer (2003). environmental data include soil type, aspect, slope, surface rock cover and disturbance to the soil and vegetation. the data set, containing 846 specific and infraspecific taxa, was entered into a vegetation database created in turboveg (hennekens & schamineé 2001). a first approximation of the main communities was arrived at by applying the twinspan classification program (hill 1979a) to the floristic data, and subsequent refinement of the classification was achieved by applying braun-blanquet procedures (bredenkamp et al. 1989; fuls et al. 1993; kent & coker 1996; siebert et al. 2003; van staden & bredenkamp 2006). a synoptic table was constructed to represent the major groups defined by the twinspan classification (table 1). refinement of the synoptic table was done with braun-blanquet procedures (bredenkamp et al. 1989; fuls et al. 1993; van staden 34 vol. 50 no. 1 pp. 32 48 major vegetation types original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe & bredenkamp 2006). the synoptic table comprises species in each of the identified mvt’s on constancy values of 20% ordinal scale (i–v). only species with a minimum constancy value of 20% (ii), in any of the given mvt’s were included in the table. all the excluded taxa will be included into detailed phytosociological tables of subsequent papers that will focus on the individual major vegetation types of the sc and bnr. the ordination algorithm detrended correspondence analysis (decorana) (hill 1979b) was applied, using the computer software package pc-ord (mccune & mefford 1999), to determine gradients in vegetation and the relationship between these plant communities and the physical environment. results are depicted on scatter diagrams. results eight mvt’s were identified for the sc and the bnr. these represent the forest biome, grassland biome, savanna biome and some azonal plant communities. a dendrogram (fig. 3) of the hierarchical classification was produced with the software package twinspan (hill 1979a). the first division separated the xymalos monospora–rhus chirendensis soutpansberg forest vegetation from the savanna and grassland vegetation types. the second division separated the vegetation of the sc from that of the bnr. a third division split off the rhus rigida var. rigida–rhus magalismontanum subsp. coddii soutpansberg mistbelt vegetation from the sc–cluster, and split off the blouberg plains bushveld from the blouberg mountain bushveld. further divisions separated the blouberg northern plains bushveld from the blouberg southern plains bushveld, while the vegetation of the sc was divided into adansonia digitata–acacia nigrescens soutpansberg arid northern bushveld, catha edulis–flueggia virosa soutpansberg moist mountain thickets and diplorhynchus condylocarpon– burkea africana soutpansberg leached sandveld. the dataset was classified into different plant communities based on diagnostic species. strong emphasis was placed on long-lived perennial species for the purpose of community description and syntaxonomy. these species were specifically chosen to ensure relatively long-term predictability regarding effective plant community identification by future fieldworkers and managers. due to the fleeting existence and unpredictable appearance of annual and weak perennial species within communities of arid and semi-arid ecosystems it was decided to treat such species as the more temporary and fluctuating component within the vegetation of these event-driven systems (westoby et al. 1989). the floristic composition of the eight major vegetation types is given in a synoptic table (table 1). these mvt’s are discussed below. description of the major vegetation types 1. eragrostis lehmanniana var. lehmanniana–sclerocarya birrea subsp. caffra blouberg northern plains bushveld major vegetation type the eragrostis lehmanniana var. lehmanniana–sclerocarya birrea subsp. caffra blouberg northern plains bushveld mvt occurs on the northern foot slopes and plains of the blouberg nature reserve. the terrain is generally flat, with a maximum incline of three degrees. this mvt is associated with the hutton soil form (macvicar et al. 1991) of the ae land type derived from alluvium on sandstone of the wyllies poort geological formation (botha 2004a; patterson & ross 2004a). the diagnostic species for this group are presented in species group a (table 1). this group includes the woody species spirostachys africana. diagnostic grass species include eragrostis lehmanniana var. lehmanniana, tragus berteronianus, dactyloctenium aegyptium and pogonarthria squarrosa. this diagnostic group contains numerous herbaceous species such as phyllanthus burchellii, indigofera species, limeum fenestratum, erythrophleum africanum, bulbostylis hispidula subsp. pyriformis, ruellia species, indigofera rhytidocarpa, chamaecrista absus, hermannia grisea, tragia minor, corchorus species, talinum crispatulum, monechma divaricatum, zornia species, blepharis subvolubilis, limeum viscosum, leucas sexdentata, secamone parvifolia, chamaecrista biensis and asparagus exuvialis forma exuvialis. dominant woody species of this mvt include spirostachys africana (species group a), commiphora species, grewia flavescens var. flavescens (species group f), boscia albitrunca var. albitrunca, cissus cornifolia, combretum apiculatum subsp. apiculatum, combretum mossambicense, commiphora mollis, grewia bicolor (species group h), acacia nigrescens, dichrostachys cinerea subsp. africana, ehretia rigida, markhamia zanzibarica and sclerocarya birrea subsp. caffra (species group l). dominant grass species include eragrostis lehmanniana var. lehmanniana (species group a), eragrostis rigidior, urochloa mosambicensis (species group l), digitaria eriantha, panicum maximum and aristida stipitata subsp. graciliflora (species group p). prominent herbaceous species include phyllanthus burchellii (species group a), acanthospermum species, arctotis species, aristida congesta subsp. congesta, evolvulus alsinoides, hibiscus calyphyllus, hibiscus praeteritus, sida ovata (species group c), cyperus angolensis (species group f), solanum panduriforme (species group l) and waltheria indica (species group p). the vegetation of the eragrostis lehmanniana var. lehmanniana– sclerocarya birrea subsp. caffra blouberg northern plains bushveld mvt shows some floristic affinities with the acacia mellifera– eragrostis lehmanniana vegetation class, informally described by smit (2000) as part of the eastern kalahari thornveld. shared prominent species within the blouberg northern plains bushveld and the acacia mellifera–eragrostis lehmanniana 35 1 eragrostis lehmanniana–sclerocarya birrea blouberg northern plains bushveld 2 euclea divinorum–acacia tortilis blouberg southern plains bushveld 3 englerophytum magalismontanum–combretum molle blouberg mountain bushveld 4 adansonia digitata–acacia nigrescens soutpansberg arid northern bushveld 5 catha edulis–flueggia virosa soutpansberg moist mountain thickets 6 diplorhynchus condylocarpon–burkea africana soutpansberg leached sandveld 7 rhus rigida–rhus magalismontanum soutpansberg mistbelt vegetation 8 xymalos monospora–rhus chirendensis soutpansberg forest vegetation figure 3 a dendrogram showing the hierarchical divisions created by the twinspan (two way indicator species analysis) computer software package (hill 1979a) legend vol. 50 no. 1 pp. 32 48 original research mostert, bredenkamp, klopper, verwey, mostert & hahn koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za36 vegetation type 1 2 3 4 5 6 7 8 number of relevés 98 84 16 65 20 50 33 70 species group a diagnostic spp for the eragrostis lehmanniana var. lehmanniana– sclerocarya birrea subsp. caffra blouberg northern plains bushveld eragrostis lehmanniana var. lehmanniana v i i i phyllanthus burchellii iv indigofera sp. iii i i leucas sexdentataw iii limeum fenestratum iii erythrophleum africanum iii bulbostylis hispidula subsp. pyriformis iii i i i ruellia sp. ii indigofera rhytidocarpa ii chamaecrista absus ii hermannia grisea ii tragus berteronianus ii tragia minor ii corchorus sp. ii i talinum crispatulum ii monechma divaricatum ii dactyloctenium aegyptium ii zornia sp. ii pogonarthria squarrosa ii blepharis subvolubilis ii i limeum viscosum ii secamone parvifolia ii i spirostachys africana ii chamaecrista biensis ii asparagus exuvialis fo. exuvialis ii species group b diagnostic spp for the euclea divinorum–acacia tortilis blouberg southern plains bushveld lepidagathis scabra i v rhinacanthus xerophilus iii melhania prostrata iii i i rhus leptodictya i ii i i enteropogon macrostachyus ii combretum imberbe ii i schotia brachypetala ii i i aristida congesta subsp. barbicollis ii i i corbichonia decumbens ii species group c hibiscus praeteritus v iv acanthospermum sp. v ii evolvulus alsinoides iv ii melhania forbesii iv ii aristida congesta subsp. congesta iii ii i hibiscus calyphyllus iii v sida ovata iii ii i blepharis integrifolia iii ii i table 1 synoptic table of the major vegetation types of the soutpansberg conservancy and the blouberg nature reserve vegetation type 1 2 3 4 5 6 7 8 number of relevés 98 84 16 65 20 50 33 70 lantana rugosa ii iv i acacia tortilis w. heteracantha ii iii i i aristida adscensionis ii iii arctotis sp. ii ii acalypha indica ii ii philenoptera violacea ii ii i ocimum gratissimum subsp. gratissimum ii ii species group d diagnostic spp for the englerophytum magalismontanum–combretum molle blouberg mountain bushveld loudetia filifolia iv rhynchosia vendae i iii trichoneura grandiglumis iii aristida sp. i iii combretum zeyheri i ii elephantorrhiza sp. ii dalechampia sp. ii tricliceras schinzii ii species group e cheilanthes involuta ii iii tephrosia purpurea iii iv species group f cyperus angolensis iii iii iii commiphora sp. iv ii iii grewia flavescens var. flavescens iv iv ii eragrostis biflora ii ii ii species group g diagnostic spp for the adansonia digitata–acacia nigrescens soutpansberg arid northern tribulus terrestris iii grewia hexamita iii commiphora glandulosa iii blepharis diversispina iii adansonia digitata ii grewia flava i i ii grewia subspathulata ii cordia monoica ii i kirkia acuminata i i ii maerua parvifolia ii terminalia prunioides i ii maerua edulis ii i commiphora tenuipetiolata ii tephrosia macropoda ii solanum lichtensteinii ii i cassia abbreviata i ii cleome angustifolia subsp. petersiana ii ochna inermis ii i lannea schweinfurthii i ii ledebouria apertiflora ii table 1 (cont...) vol. 50 no. 1 pp. 32 48 major vegetation types original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe 37 vegetation type 1 2 3 4 5 6 7 8 number of relevés 98 84 16 65 20 50 33 70 commiphora viminea ii sterculia rogersii i ii grewia villosa ii chamaecrista mimosoides ii i i boscia foetida subsp. rehmanniana ii sansevieria aethiopica ii heliotropium steudneri i i ii i commiphora africana var. africana ii species group h commiphora mollis iv ii iv combretum apiculatum subsp. apiculatum iii iii i iii i boscia albitrunca var. albitrunca iii ii ii grewia bicolor var. bicolor iii iii ii i i cissus cornifolia ii ii ii ii combretum mossambicense iii ii pristimera longipitiolata ii i ii i species group i diagnostic spp for the catha edulis–flueggia virosa soutpansberg moist mountain thickets grewia occidentalis var. occidentalis iv dovyalis zeyheri iv acalypha glabrata i iv dombeya rotundifolia var. rotundifolia i iv i catha edulis iv rhus pentheri iii i carissa edulis iii rhoicissus tridentata subsp. tridentata iii i senna petersiana i i iii diospyros lycioides i iii i berchemia zeyheri iii dovyalis caffra ii brachiaria deflexa ii capparis tomentosa ii euphorbia ingens i ii acacia ataxacantha i ii i euclea undulata i ii pavetta schumanniana ii i acacia rehmanniana ii commelina benghalensis ii i gymnosporia senegalensis ii acokanthera oppositifolia ii mystroxylon aethiopicum subsp. schlechteri i ii rhus pyroides ii bridelia mollis i i ii clerodendrum glabrum var. glabrum ii ekebergia capensis ii allophylus africanus var. africanus ii tarchonanthus camphoratus ii maerua caffra ii coddia rudis ii i table 1 (cont...) vegetation type 1 2 3 4 5 6 7 8 number of relevés 98 84 16 65 20 50 33 70 lippia javanica ii i jasminum multipartitum ii i i combretum hereroense i i i ii eragrostis superba ii i panicum deustum i i ii acacia caffra ii setaria megaphylla ii christella guenziana ii euclea crispa subsp. crispa ii cyperus albostriatus ii i olea europaea subsp. africana ii bridelia micrantha ii buddleja saligna ii solanum tettense var. renschii i ii cussonia natalensis ii ficus sycomorus subsp. sycomorus ii dicliptera heterostegia i ii scolopia zeyheri ii canthium inerme ii buddleja salviifolia ii podocarpus falcatus ii pyrenacantha grandiflorus ii aloe greatheadii var. greatheadii ii i canthium mundianum ii i bothriochloa insculpta ii ficus sur ii i sansevieria hyacinthoides i ii cyperus sphaerospermus ii i bulbostylis burchellii ii i syzygium cordatum ii aloe marlothii subsp. marlothii ii barleria gueinzii ii pavetta eylesii ii capparis fascicularis var. fascicularis ii acacia gerrardii var. gerrardii ii justicia flava i i ii species group j plectroniella armata ii iii i hibiscus meyeri iii ii peltophorum africanum i ii iii i ximenia caffra var. caffra ii ii i gossypium herbaceum subsp. africanum ii ii species group k euclea divinorum v ii ii flueggia virosa i iii ii iv ximenia americana var. microphylla ii ii ii heteropogon contortus i ii ii ii i themeda triandra ii ii i iii i acacia karroo ii v table 1 (cont...) vol. 50 no. 1 pp. 32 48 original research mostert, bredenkamp, klopper, verwey, mostert & hahn koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za38 vegetation type 1 2 3 4 5 6 7 8 number of relevés 98 84 16 65 20 50 33 70 gymnosporia buxifolia i ii i iii pappea capensis ii i i ii acacia nilotica ii i ii species group l dichrostachys cinerea subsp. africana iv iv iv iii solanum panduriforme iii iv ii i ii acacia nigrescens ii iv iv ii i urochloa mosambicensis iii ii ii sclerocarya birrea subsp. caffra iv ii ii ii eragrostis rigidior iii iii i ii grewia monticola iii iv iii ii i ehretia rigida ii iv ii iii markhamia zanzibarica iv i i ii species group m diagnostic spp for the diplorhynchus condylocarpon–burkea africana soutpansberg leached sandveld centropodia glauca iii elephantorrhiza burkei iii diplorhynchus condylocarpon iii ochna pulchra iii i hexalobus monopetalus var. monopetalus i ii grewia retinervis ii ipomoea albivenia ii strychnos pungens ii eragrostis pallens ii ozoroa paniculosa var. salicina i i ii schizachyrium jeffreysii ii selaginella dregei i ii i euphorbia aeruginosa ii aloe angelica ii pterocarpus angolensis ii garcinia livingstonei ii eragrostis gummiflua ii i euphorbia zoutpansbergensis ii commiphora marlothii i ii ficus abutilifolia ii euphorbia cooperi ii cineraria parvifolia ii artabotrys brachypetalus ii portulaca kermesina ii i aristida canescens subsp. ramosa ii tephrosia longipes ii aristida diffusa subsp. burkei ii indigofera cryptantha var. cryptantha ii ficus tettensis ii adenia spinosa ii orthosiphon labiatus i ii i isoglossa hypoestiflora ii anacampseros subnuda ii i table 1 (cont...) vegetation type 1 2 3 4 5 6 7 8 number of relevés 98 84 16 65 20 50 33 70 asparagus laricinus ii loudetia flavida ii androstachys johnsonii ii dicoma montana ii species group n tricalysia junodii var. kirkii ii ii euclea natalensis i ii ii ii species group o pseudolachnostylis maprouneifolia i iv i iv burkea africana iii iv species group p panicum maximum v v iv iii iv ii digitaria eriantha iv i iv i i ii i ziziphus mucronata ii iii i ii v ii strychnos madagascariensis ii i iv ii iii enneapogon cenchroides iii iii ii ii iii i waltheria indica iv ii ii ii ii schmidtia pappophoroides iii i ii iii ii terminalia sericea ii i ii ii iii aristida stipitata subsp. graciliflora iii ii ii stipagrostis uniplumis var. uniplumis ii i ii ii species group q diagnostic spp for the rhus rigida var. rigida–rhus magalismontanum subsp. coddii soutpansberg mistbelt vegetation rhus rigida var. rigida iv melinis nerviglumis iii helichrysum kraussii iii brachiaria serrata iii cryptolepis cryptolepioides i iii parinari capensis subsp. capensis i iii fadogia homblei iii coleochloa setifera iii setaria sphacelata var. torta iii rhynchosia monophylla i ii olea capensis subsp. enervis ii senecio barbertonicus ii syzygium legatii ii aloe arborescens ii rotheca myricoides ii euclea linearis ii crassula swaziensis i ii khadia borealis ii trachypogon spicatus ii rhus tumulicola var. meeuseana ii commelina erecta i i ii olinia rochetiana ii cyperus obtusiflorus var. obtusiflorus ii combretum moggii ii vernonia natalensis ii table 1 (cont...) vol. 50 no. 1 pp. 32 48 major vegetation types original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe 39 vegetation type 1 2 3 4 5 6 7 8 number of relevés 98 84 16 65 20 50 33 70 dicoma anomala i ii vangueria soutpansbergensis i ii schistostephium crataegifolium ii coptosperma supra-axillare ii plectranthus neochilus ii gnidia cuneata ii elionurus muticus ii protea caffra subsp. caffra ii elephantorrhiza elephantina ii tetradenia riparia ii wahlenbergia undulata ii hypoxis argentea var. argentea ii eulophia ensata ii pteridium aquilinum ii kalanchoe sexangularis var. sexangularis ii sarcostemma viminale i i ii aristea woodii ii apodytes dimidiata subsp. dimidiata ii anthospermum welwitschii ii vernonia oligocephala ii protea roupelliae subsp. roupelliae ii pentanisia prunelloides subsp. prunelloides ii lopholaena coriifolia ii senecio oxyriifolius ii coptosperma rhodesiacum ii viscum rotundifolium ii ekebergia pterophylla ii myrsine africana ii ipomoea oblongata ii helichrysum cerastioides ii bulbostylis contexta ii hypoxis hemerocallidea ii asparagus falcatus i ii species group r loudetia simplex iii iv xerophyta retinervis iii ii corchorus kirkii i iii ii commelina africana var. lancispatha i ii ii landolphia kirkii ii ii combretum vendae ii ii vangueria parvifolia i ii ii myrothamnus flabellifolius ii ii species group s zanthoxylum capense iv ii dioscorea sylvatica ii ii heteropyxis natalensis ii ii species group t pellaea calomelanos var. calomelanos ii ii iii ii table 1 (cont...) vegetation type 1 2 3 4 5 6 7 8 number of relevés 98 84 16 65 20 50 33 70 species group u combretum molle v iv ii iii vitex rehmannii i iv i iii ii englerophytum magalismontanum iv i iii vangueria infausta subsp. infausta i i iii ii ii ii rhus magalismontanum subsp. coddii ii iv hyperacanthus amoenus ii ii ii ii mimusops zeyheri ii ii iii species group v diagnostic spp for the xymalos monospora–rhus chirendensis soutpansberg forest vegetation xymalos monospora iv zanthoxylum davyi ii celtis africanaw i ii nuxia floribunda i ii rhoicissus tomentosa i ii kiggelaria africana ii vepris lanceolata i ii rapanea melanophloeos i ii rothmannia capensis i ii brachylaena discolor i ii ficus craterostoma ii combretum kraussii ii trichilia dregeana ii trimeria grandifolia ii drypetes gerrardii ii oxyanthus speciosus subsp. gerrardii ii species group w diospyros whyteana ii ii species group x maytenus undata iii iii ii rhus chirindensis ii iv cussonia spicata ii i ii species group y maesa lanceolata ii ii vegetation class include the grasses eragrostis lehmanniana, pogonarthria squarrosa and schmidtia pappophoroides. a shared prominent woody species is boscia albitrunca. however, these five species all show wide distribution ranges and exhibit wide ecological tolerance and adaptation within the sandveld areas of southern africa, and are therefore not exclusive to the blouberg arid mountain bushveld. this major vegetation type shares some of the drought-resistant woody species with the adansonia digitata–acacia nigrescens soutpansberg arid northern bushveld major vegetation type, such as commiphora mollis, combretum apiculatum, boscia albitrunca var. albitrunca, grewia bicolor, combretum mossambicense, commiphora africana, dichrostachys cinerea subsp. africana and acacia nigrescens. these species are also commonly found in the adansonia mixed thornveld (14e) (acocks 1953), the adansonia digitata–colophospermum mopane rugged veld (gertenbach 1983) and the commiphora–terminalia prunioides community (louw 1970). table 1 (cont...) vol. 50 no. 1 pp. 32 48 original research mostert, bredenkamp, klopper, verwey, mostert & hahn koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za40 2. euclea divinorum–acacia tortilis blouberg southern plains bushveld major vegetation type the euclea divinorum–acacia tortilis blouberg southern plains bushveld mvt is restricted to the plains and foot slopes south of the blouberg within the bnr. the terrain is generally flat, with a slope that varies between one to five degrees. it is predominantly associated with the hutton soil form (macvicar et al. 1991), derived from alluvium on sandstone of the ae land type from the wyllies poort geological formation (botha 2004b; patterson & ross 2004a). other soil forms associated with this major vegetation type are of less importance. the oakleaf and valsrivier soil forms (macvicar et al. 1991) are associated with alluvium, sand and calcrete of the quaternary deposits of the ia land type (botha 2004b). the glenrosa and mispah soil forms (macvicar et al. 1991) of the fc land type (botha 2004b) are associated with basalt of the letaba geological formation in the lebombo group–karoo sequence, as well as aphibolite and metapelite of the malala drift group. the mispah, avalon and hutton soil forms (macvicar et al. 1991) of the bc land type (botha 2004b) are associated with leucocratic migmatite and gneiss, grey and pink hornblende–biotite gneiss, grey biotite gneiss, minor muscovite–bearing granite, pegmatite and gneiss of the hout river gneis geological formation, as well as metapelite of the bandelierkop complex. the diagnostic species for this group are presented in species group b (table 1). the diagnostic woody species characterising the communities of this mvt are combretum imberbe, rhus leptodictya and schotia brachypetala. diagnostic grass species include enteropogon macrostachyus and aristida congesta subsp. barbicollis. diagnostic herbaceous species within this group are lepidagathis scabra, rhinacanthus xerophilus, melhania prostrata and corbichonia decumbens. dominant woody species of this mvt include acacia tortilis subsp. heteracantha, lantana rugosa, rhus leptodictya (species group c), grewia flavescens var. flavescens (species group f), combretum apiculatum subsp. apiculatum, grewia bicolor var. bicolor (species group h), acacia nilotica, euclea divinorum (species group k), acacia nigrescens, dichrostachys cinerea subsp. africana, ehretia rigida, eragrostis rigidior (species group l) and ziziphus mucronata (species group p). dominant grass species include enteropogon macrostachyus (species group b), aristida congesta subsp. congesta, aristida adscensionis (species group c), aristida species (species group d), eragrostis rigidior, urochloa mosambicensis (species group l), enneapogon cenchroides and panicum maximum (species group p). prominent forbs include hibiscus praeteritus, sida ovata, blepharis integrifolia (species group c), tephrosia purpurea (species group e), cyperus angolensis (species group f) and solanum panduriforme (species group l). the euclea divinorum–acacia tortilis blouberg southern plains bushveld major vegetation type is regarded as a variant of the rhus leptodictya–acacia tortilis bushveld of the acacietalia rehmannianae–tortilis of the acacienea nilotico–tortilis of the panico maximi–acacietea tortilis, described by winterbach (1998) and winterbach et al. (2000) for the north-western savanna of south africa. acocks (1953) described similar vegetation as the knoppiesdoring veld (13b) of the other turf thornveld (13). it also compares floristically with the acacia tortilis–panicum maximum–ziziphus mucronata major plant community of the waterberg (henning 2002). these communities are generally not geographically restricted to a certain part of south africa, but occur in a patchy distribution where conditions are favourable. important taxa binding these communities together are acacia tortilis subsp. heteracantha rhus leptodictya, grewia species, acacia nilotica, euclea divinorum, acacia nigrescens, dichrostachys cinerea subsp. africana, ehretia rigida, eragrostis rigidior, ziziphus mucronata, aristida species eragrostis rigidior, urochloa mosambicensis and panicum maximum. 3. englerophytum magalismontanum–combretum molle blouberg mountain bushveld major vegetation type the englerophytum magalismontanum–combretum molle blouberg mountain bushveld major vegetation type can be described as the mountain bushveld component of the bnr. this mvt is restricted to the higher lying mountainous terrain of the bnr, ranging from 975–1 465 m above sea level. west of the bnr the blouberg rises to 2 051 m above sea level, where more temperate vegetation types occur (van jaarsveld & hardy 1991). average slopes range from moderate to very steep, with northern, southern and eastern aspects. soils are generally shallow or skeletal (< 100 mm), and are associated with mispah and glenrosa soil forms (macvicar et al. 1991) of the fa land type (botha 2004a; patterson & ross 2004a). the underlying geology is dominated by pink quartzite and minor conglomerates of the wyllies poort geological formation of the soutpansberg group. the diagnostic species for this group are presented in species group d (table 1). diagnostic woody species include combretum zeyheri and an elephantorrhiza species. the diagnostic grass species include loudetia filifolia, trichoneura grandiglumis and aristida species. diagnostic herbaceous species include rhynchosia vendae, a dalechampia species and tricliceras schinzii. dominant woody species of the englerophytum magalismontanum–combretum molle blouberg mountain bushveld include combretum zeyheri, elephantorrhiza species (species group d), commiphora species (species group f), burkea africana, pseudolachnostylis maprouneifolia (species group o), strychnos madagascariensis (species group p), combretum molle, englerophytum magalismontanum, hyperacanthus amoenus, mimusops zeyheri, rhus magalismontanum subsp. coddii, vangueria infausta subsp. infausta and vitex rehmannii (species group u). dominant grass species include aristida species, loudetia filifolia, trichoneura grandiglumis (species group d), digitaria eriantha, enneapogon cenchroides, panicum maximum and schmidtia pappophoroides (species group p). prominent herbaceous species include rhynchosia vendae (species group d), cheilanthes involuta, tephrosia purpurea (species group e), cyperus angolensis (species group f) and waltheria indica (species group p). the englerophytum magalismontanum–combretum molle blouberg mountain bushveld major vegetation type shares floristic elements with the englerophyto magalismontani–acacietea caffrae of the waterberg and magaliesberg, as described by winterbach et al. (2000). it shares many of the relatively drought tolerant species associated with the rhus rigida var. rigida–rhus magalismontanum subsp. coddii soutpansberg cool mistbelt major vegetation type. it does however lack some of the more mesic species recorded within the mistbelt of the sc. the central variation of the bankenveld (acocks 1953) and the rocky highveld grassland of the grassland biome (bredenkamp & van rooyen 1996) share limited floristic and structural elements with the rhynchosia vendae–englerophytum magalismontanum blouberg moist mountain bushveld major vegetation type. although the vegetation structure of the diplorhynchus condylocarpon–englerophytum magalismontanum rocky slope community of the waterberg biosphere (henning 2002) is very similar to that of the blouberg moist mountain bushveld, the floristic composition differs considerably. 4. adansonia digitata–acacia nigrescens soutpansberg arid northern bushveld major vegetation type the adansonia digitata–acacia nigrescens soutpansberg arid northern bushveld major vegetation type is confined to the rain-shadow northern ridges of the sc. due to the wide range of vol. 50 no. 1 pp. 32 48 major vegetation types original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe 41 topographic and edaphic conditions found within this mvt, it comprises a complex of very diverse plant communities. despite the diversity of these plant communities, most are adapted to prolonged water-stress conditions and unpredictable rainfall events. these drought events are likely to be the main driving factors behind the vegetation structure and species composition of the various plant communities within this major vegetation type. this soutpansberg arid northern bushveld mvt is associated with the clovelly soil form (macvicar et al. 1991) of the ae, ag, ia, ib, and fa land types derived from sandstone, quartzite and conglomerate of the wyllies poort geological formation, basalt from the musekwa geological formation, as well as from narrow diabase intrusions or dykes within the wyllies poort geological formation (botha 2004a; patterson & ross 2004a). the ia land type in particular is associated with the aeolian sands (kalahari sand) covering the musekwa sediments at the foot of the mountain. the vegetation structure can generally be described as open woodland (edwards 1983), with a very sparse field layer. this is especially true during dry cycles, when only the most hardy perennial grass and herbaceous species survive. the woody layer ranges from open along the deeper well-drained sandy soils of the northern plains, to sparse along the shallower clayey foot slopes and around the saltpan. the diagnostic species for this group are presented in species group g (table 1). diagnostic woody species characterising the communities of this mvt along the northern sandy plains include adansonia digitata, boscia foetida subsp. rehmanniana, commiphora glandulosa, commiphora tenuipetiolata, cordia monoica, blepharis diversispina, grewia flava, grewia subspathulata, grewia villosa and grewia hexamita. diagnostic woody species along the basaltic-clay foot slopes are kirkia acuminata, maerua parvifolia, maerua edulis, terminalia prunioides, cassia abbreviata, sterculia rogersii and commiphora viminea. although tribulus terrestris was recorded as a relatively strong diagnostic herbaceous species for the group, due to its status as a pioneer annual herbaceous species that dominates disturbed patches, it is not recommended as a reliable indicator species for the soutpansberg arid northern bushveld. instead, sansevieria aethiopica and ledebouria apertiflora should be seen as more reliable perennial herbaceous indicator species for this broad mvt. prominent woody species of this mvt include grewia hexamita, commiphora glandulosa, blepharis diversispina, adansonia digitata (species group g), commiphora mollis, combretum apiculatum (species group h), hibiscus meyeri (species group j), acacia nigrescens, dichrostachys cinerea and grewia monticola (species group l). although none of the grass species could be labelled as prominent during the time of the surveys, the most dominant grass species included panicum maximum and schmidtia pappophoroides (species group p). forbs recorded from the poorly developed field layer are tephrosia macropoda, solanum lichtensteinii, ledebouria apertiflora, chamaecrista mimosoides, heliotropium steudneri (species group g), hibiscus meyeri, gossypium herbaceum subsp. africanum (species group j), waltheria indica (species group p) and pellaea calomelanos var. calomelanos (species group t). the adansonia digitata–acacia nigrescens soutpansberg arid northern bushveld major vegetation type has been described by acocks (1953) as the adansonia–mixed thornveld variant (14e) of the arid sweet bushveld (14). due to the scale at which these veld types were mapped, acocks’ (1953) description includes more variation along the lower lying microphyllous plains between the blouberg and soutpansberg mountain ranges and less along the foot slopes of these mountains. the adansonia–mixed thornveld variant (14e) described by acocks (1953) is therefore very similar to the associations along the northern sandy plains described here under the soutpansberg arid northern bushveld major vegetation type. 5. catha edulis–flueggia virosa soutpansberg moist mountain thickets major vegetation type the catha edulis–flueggia virosa moist mountain thickets mvt is a mixture of plant communities situated at different altitudes. they are, however, all associated with soils of a high clay content and relatively moist conditions. even during dry cycles, the moisture-laden air from the south-east ensures at least some orographic rain and moisture during the summer months. these plant communities are primarily confined to the steep southern slopes where igneous rock in the form of basalt and diabase settled after flowing through the cracks and tears in the upper sedimentary rock layers. however, it also contains plant communities associated with the illuvial clays found in some of the valleys cutting through the mountain ridges. the moist mountain thickets are associated with the shortlands soil sorm (macvicar et al. 1991) derived from basalt and tuff associated with the fa land type of the sibasa geological formation, as well as from narrow diabase intrusions or dykes associated with the ib land type of the wyllies poort geological formation (botha 2004b; patterson & ross 2004b). the vegetation structure can be described as low, closed thickets (edwards 1983), with no definite separation between the tree and shrub layers. the woody layer contains a mixture of trees and shrubs, ranging from 1.5–4 m in height. a very high percentage canopy cover (> 80%) blocks sunlight from reaching the understory and prevents the establishment of a dense field layer. this major vegetation type is characterised by a high diversity of trees and shrubs. the diagnostic species are presented in species group i (table 1) and include woody species from relatively moist thickets and wet riverine thickets such as catha edulis, grewia occidentalis, dovyalis zeyheri, acalypha glabrata, dombeya rotundifolia, rhus pentheri, carissa edulis, rhoicissus tridentata subsp. tridentata, senna petersiana, diospyros lycioides, berchemia zeyheri, dovyalis caffra, capparis tomentosa, euphorbia ingens, acacia ataxacantha, euclea undulata, pavetta schumanniana, acacia rehmanniana, gymnosporia senegalensis, acokanthera oppositifolia, mystroxylon aethiopicum subsp. schlechteri, rhus pyroides, bridelia mollis, clerodendrum glabrum var. glabrum, ekebergia capensis, allophylus africanus var. africanus, tarchonanthus camphoratus, maerua caffra, coddia rudis, lippia javanica, jasminum multipartitum, combretum hereroense, acacia caffra, euclea crispa subsp. crispa, olea europaea subsp. africana, bridelia micrantha, buddleja saligna, cussonia natalensis, ficus sycomorus subsp. sycomorus, scolopia zeyheri, canthium inerme, buddleja salviifolia, podocarpus falcatus, pyrenacantha grandiflorus, canthium mundianum, ficus sur, syzygium cordatum, pavetta eylesii, capparis fascicularis var. fascicularis and acacia gerrardii var. gerrardii. the recorded diagnostic succulent species listed in species group i are generally widespread throughout south africa and include aloe greatheadii var. greatheadii, sansevieria hyacinthoides and aloe marlothii subsp. marlothii. diagnostic grass species recorded from the poorly developed field layer are brachiaria deflexa, eragrostis superba, panicum deustum, setaria megaphylla and bothriochloa insculpta. diagnostic herbaceous species include commelina benghalensis, christella guenziana, cyperus albostriatus, solanum tettense var. renschii, dicliptera heterostegia, cyperus sphaerospermus, bulbostylis burchellii, barleria gueinzii and justicia flava. other prominent woody species include plectroniella armata, peltophorum africanum (species group j), flueggia virosa subsp. virosa, acacia karroo, gymnosporia buxifolia (species group k), ehretia rigida (species group l), ziziphus mucronata (species group p), heteropyxis natalensis, zanthoxylum capense (species vol. 50 no. 1 pp. 32 48 original research mostert, bredenkamp, klopper, verwey, mostert & hahn koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za group s), combretum molle (species group u) and maytenus undata (species group x). only a few shade-tolerant grass and herbaceous species are prominent within this mvt such as brachiaria deflexa, panicum deustum, setaria megaphylla (species group i) and panicum maximum (species group p). the shaded conditions also favour fern species such as christella gueinziana (species group i) and pellaea calomelanos var. calomelanos (species group t). acocks (1953) considered this major vegetation type one of the many variants of the “sourish mixed bushveld” and the “north-eastern mountain sourveld”. due to its association with the narrow intrusive basalt, tuff and diabase dykes, it is often difficult to map this major vegetation type when dealing with coarse or small-scale vegetation studies. the southernmost ridge of the mountain with its large southern basalt slope is one of the few extensive and mappable areas with this major vegetation type. the strong mixture of temperate and sub-tropical plant species reflects the gradient of plant communities up the southern slopes of the sc. while those communities along the warm and relatively humid foot slopes are very tropical, the plant communities of clayey depressions along the cooler higher lying areas tend to contain many temperate species. the lower lying sub-tropical plant communities seem to be unique in their species composition and structure. the higher lying temperate communities compare with the hillside scrub of the transitional cymbopogon–themeda veld (49) of the pure grassveld types described by acocks (1953). they share species such as acacia karroo, acacia caffra, grewia occidentalis, ehretia rigida, euclea crispa subsp. crispa, olea europaea subsp. africana, buddleja saligna, buddleja salviifolia, rhus pyroides, tarchonanthus camphoratus, diospyros lycioides, ziziphus mucronata, dombeya rotundifolia and cussonia species. these temperate clay communities of the sc also share some floristic links with the riverine woodland zizipho mucronatae–acacietum karroo described by brown (1997). according to du preez and bredenkamp (1991), bezuidenhout et al. (1994) and winterbach (1998), the acacia karroo-dominated vegetation of the southern african grassveld and savanna should be classified as a separate syntaxonomical class. the high-lying communities associated with fine textured clayey soils along the southern slopes of the sc may be classified as part of this proposed class. 6. diplorhynchus condylocarpon–burkea africana soutpansberg leached sandveld major vegetation type the diplorhynchus condylocarpon–burkea africana leached sandveld mvt is confined to the warmer northern slopes of the mountain, as well as some of the more arid southern slopes along the northernmost ridges of the mountain range, which falls within the rain shadow zone of the mountain. it is associated with the mispah and hutton soil forms (macvicar et al. 1991) derived from sandstone, quartzite and conglomerate associated with the ae, fa, and ib land types of the wyllies poort geological formation (botha 2004a; patterson & ross 2004a). one plant community in particular is associated with deep regic sands of the namib soil form, which is of an aeolian origin from the kalahari (brandl 2002). it also includes the vegetation associated with the almost bare sheets of exposed rock against the warm northern slopes. the plant communities of this mvt occur on both very shallow and very deep sands of the relatively dry landscapes of the sc. the shallow soils are situated on steep rocky inclines, while the deep sands are associated with relatively high-lying flat plateaus. the combination of the underlying nutrient poor quartzite parent material and the eluviation of silt and clay particles from these well drained sandy soils have left this system extremely nutrient poor. with the exception of the plant communities associated with the rock sheets, the leached sandveld communities are relatively poor in plant species diversity. the diagnostic species for this group are presented in species group m (table 1). the leached sandveld mvt is characterised by diagnostic woody species such as elephantorrhiza burkei, diplorhynchus condylocarpon, ochna pulchra, grewia retinervis and strychnos pungens. diagnostic grass species include centropodia glauca, eragrostis pallens, schizachyrium jeffreysii, eragrostis gummiflua, aristida canescens subsp. ramosa, aristida diffusa subsp. burkei and loudetia flavida. the herbaceous layer is sparse and species poor and contains some weak indicator species such as selaginella dregei, cineraria parvifolia and tephrosia longipes. prominent woody species include pseudolachnostylis maprouneifolia, burkea africana (species group o), strychnos madagascariensis, terminalia sericea (species group p), xerophyta retinervis (species group r) and vitex rehmannii (species group u). locally dominant grass species include centropodia glauca, eragrostis pallens, schizachyrium jeffreysii, eragrostis gummiflua, aristida canescens subsp. ramose, aristida diffusa, loudetia flavida (species group m), stipagrostis uniplumis var. uniplumis, enneapogon cenchroides, schmidtia pappophoroides, aristida stipitata subsp. graciliflora (species group p) and loudetia simplex (species group r). some of the locally prominent herbaceous species include selaginella dregei, portulaca kermesina, tephrosia longipes, indigofera cryptantha var. cryptantha, orthosiphon labiatus, isoglossa hypoestiflora, anacampseros subnuda subsp. subnuda (species group m), waltheria indica (species group p), corchorus kirkii, commelina africana subsp. lancispatha, myrothamnus flabellifolius (species group r) and pellaea calomelanos var. calomelanos (species group t). deep soil depth shallow low soil moisture high low surface rock cover high low altitude high flat slope steep arid plant–available moisture moist fine sands soil texture coarse a eragrostis lehmanniana–sclerocarya birrea blouberg northern plains bushveld b euclea divinorum–acacia tortilis blouberg southern plains bushveld c englerophytum magalismontanum–combretum molle blouberg mountain bushveld legend figure 4 relative positions of relevés along the second and third axes of the decorana ordination (hill 1979b) of the vegetation data from the blouberg nature reserve. cooler, moist high-lying slopes and crests c b a arid northern plains cooler southern plains and high-lying steep slopes and crests 42 vol. 50 no. 1 pp. 32 48 major vegetation types original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe type, diagnostic species comprise of a mixture of inconspicuous perennials, and do not include any dominant or abundant species. diagnostic species for this group are presented in species group q (table 1). the most important diagnostic woody species include rhus rigida var. rigida, helichrysum kraussii, cryptolepis cryptolepioides and parinari capensis subsp. capensis. other, less constant woody indicator species of this mvt include olea capensis subsp. enervis, syzygium legatii, aloe arborescens, rotheca myricoides, euclea linearis, rhus tumulicola var. meeuseana, olinia rochetiana, combretum moggii, vangueria soutpansbergensis, coptosperma supra–axillare, protea caffra subsp. caffra, elephantorrhiza elephantina, tetradenia riparia, apodytes dimidiata subsp. dimidiata, protea roupelliae subsp. roupelliae, lopholaena coriifolia, coptosperma rhodesiacum, ekebergia pterophylla and myrsine africana. diagnostic grass species include melinis nerviglumis, brachiaria serrata, setaria sphacelata var. torta, trachypogon spicatus and elionurus muticus. diagnostic herbaceous species recorded are fadogia homblei, coleochloa setifera, rhynchosia monophylla, senecio barbertonicus, crassula swaziensis, khadia borealis, vernonia natalensis, dicoma anomala, plectranthus neochilus, gnidia cuneata, eulophia ensata, pteridium aquilinum, kalanchoe sexangularis, aristea woodii, anthospermum welwitschii, vernonia oligocephala, pentanisia prunelloides subsp. prunelloides, senecio oxyriifolius, ipomoea oblongata, helichrysum cerastioides, bulbostylis contexta, hypoxis hemerocallidea and asparagus falcatus. prominent species within this collection of diverse communities include the woody species rhus rigida var. rigida, helichrysum kraussii, cryptolepis cryptolepioides, parinari capensis subsp. capensis (species group q), combretum molle, englerophytum magalismontanum, rhus magalismontanum subsp. coddii, mimusops zeyheri (species group u) and maytenus undata (species group x). locally prominent grass species include melinis nerviglumis, brachiaria serrata, setaria sphacelata var. torta, trachypogon spicatus, elionurus muticus (species group q) and loudetia simplex (species group r). due to the topographic heterogeneity of this mvt, and the diversity of plant communities it contains, prominent herbaceous species are only acocks (1953) described the various plant communities of the diplorhynchus condylocarpon–burkea africana soutpansberg leached sandveld vegetation as variations of the mixed terminalia–dichapetalum veld (18b) of the mixed bushveld (18). the burkea africana–setaria lindenbergiana major community described by van staden (2002) and van staden and bredenkamp (2005), the barleria bremekampii–diplorhynchus tree savanna (coetzee et al. 1976), as well as the burkea africana– setaria sphacelata undulating plains, foot slopes, terraces and plateaus community and the terminalia sericea–eragrostis pallens deep sandy lowlands community described by henning (2002) from the waterberg area share many diagnostic species with the diplorhynchus condylocarpon–burkea africana soutpansberg leached sandveld major vegetation type of the sc. the coarse vegetation described by van den berg (1993) of the sour and mixed bushveld emphasises the heterogeneity of these veld types. the soutpansberg leached sandveld mvt shares numerous diagnostic species with the terminalio sericeae–combretetea apiculati described by winterbach (1998) and winterbach et al. (2000). more specifically, this major vegetation type shares numerous diagnostic species with the burkea africana–perotis patens woodland alliance described by van der meulen (1979). due to the nutrient poor nature of this major vegetation type, grazing and browsing fodder production is generally low. most of the landscape is homogeneous, resulting in low species richness. however, the more rugged landscapes with their higher diversity of micro-habitats are relatively rich in species. 7. rhus rigida var. rigida–rhus magalismontanum subsp. coddii soutpansberg cool mistbelt major vegetation type the rhus rigida var. rigida–rhus magalismontanum subsp. coddii mistbelt vegetation of the sc is situated higher than 1200 m above sea level and is confined to the mistbelt region. it is associated with glenrosa and mispah soil forms (macvicar et al. 1991) derived from sandstone, quartzite and conglomerate associated with the fa and ib land types of the wyllies poort geological formation (botha 2004b; patterson & ross 2004b). the champagne soil form (macvicar et al. 1991) was recorded along some of the localized high-lying wetlands and peatlands. this mvt is associated with the rugged landscapes of the upper plateaus and crests of the sc. the soils derived from the underlying sandstone and quartzite can be described as extremely shallow, coarse sands. the depth of the soil and the extent of rock cover determine the vegetation structure and species composition within this mvt. during the rainfall season these plant communities are covered in mist on an almost daily basis, contributing towards the abundance of rock and bark lichens and bryophytes. the combination of frequent orographic rain and mist during the summer months leaves the available pockets of soil among the rock sheets drenched and sometimes flooded for extended periods. the deeper soil pockets and the half-weathered matrix of saprolite within the mistbelt can be regarded as the sponge areas, which slowly release water to feed mountain streams over extended periods. during the dry season the shallow sandy soils rapidly dry out. the local and surrounding topography of the landscape plays a major role in the rate of desiccation or water retention of a specific patch of soil. extended flooding is often a product of water seepage into an area from the surrounding catchments and sponge areas. in cases where water floods the soil for most of the year, wetlands and peatlands have formed. however, these high-lying wetlands and their surrounding catchments are prone to severe periodic droughts. these fluctuating moisture conditions favour those plant species with strategies such as the ephemerals, xerophytes and succulents. structurally, the plant communities of the mistbelt mvt are extremely diverse. it includes peatlands, low open grasslands and islands of short thickets or bush clumps (edwards 1983). due to this structural diversity within this major vegetation high soil moisture low high low clay content high a adansonia digitata–acacia nigrescens soutpansberg arid northern bushveld b catha edulis–flueggia virosa soutpansberg moist mountain thickets c diplorhynchus condylocarpon–burkea africana soutpansberg leached sandveld d rhus rigida var. rigida–rhus magalismontanum subsp. coddii soutpansberg mistbelt vegetation e xymalos monospora–rhus chirendensis soutpansberg forest vegetation legend figure 5 relative positions of relevés along the first and second axis of the decorana ordination (hill 1979b) of the vegetation data from the soutpansberg conservancy. high-altitude moist leached soils low altitude arid nutrient-rich soils d wetlands b c d a arid savanna and woodlands e b moist grasslands wet forests 43vol. 50 no. 1 pp. 32 48 original research mostert, bredenkamp, klopper, verwey, mostert & hahn koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za locally prominent and may include fadogia homblei, coleochloa setifera, rhynchosia monophylla, senecio barbertonicus, crassula swaziensis, khadia borealis, vernonia natalensis, dicoma anomala, plectranthus neochilus, gnidia cuneata, wahlenbergia undulata, hypoxis argentea var. argentea, pteridium aquilinum, kalanchoe sexangularis, vernonia oligocephala, pentanisia prunelloides subsp. prunelloides, hypoxis hemerocallidea (species group q) and pellaea calomelanos var. calomelanos (species group t). the rhus rigida var. rigida–rhus magalismontanum subsp. coddii mistbelt vegetation type of the sc is a mosaic of closed bush clumps from the englerophyto magalismontani–acacietea caffrae in the savannas (winterbach et al. 2000), the patches of high-lying, low open grasslands from the loudetio simplicis–alloteropsidetea semialatae in the grasslands (matthews et al. 1994), the loudetia simplex–aristida aequiglumis woodlands, shrublands and grasslands (coetzee 1975) and the protea caffra–loudetia simplex major community (van staden 2002). the bush clump communities show strong floristic affinities with the rhus tumulicola–aloe arborescens mist belt bush clumps (matthews 1991; matthews et al. 1991) of the north-eastern mountain sourveld (acocks 1953) and the landolphia capensis–crassula argyrophylla savanna (bredenkamp 1975). although the grassland patches share structural similarities with the diheteropogono amplectentis–proteetum gaguedi described by matthews et al. (1994) and the protea caffra–helichrysum setosum savanna described by bredenkamp (1975), they show stronger floristic affinities with the grassland patches of the structurally different helichrysum kraussii–englerophytum magalismontanum bush clump communities described by matthews (matthews 1991; matthews et al. 1991). 8. xymalos monospora–rhus chirendensis soutpansberg forest major vegetation type the forests of the sc are confined to the southern slopes of the southern most ridges of the mountain. it is associated with the glenrosa, mispah and shortlands soil forms (macvicar et al. 1991) derived from basalt, tuff, sandstone, and conglomerate associated with the fa land type of the sibasa geological formation (botha 2004b; patterson & ross 2004b). the soils are generally rich in organic matter and contain relatively high percentages of clay in areas where basalt and tuff form the underlying geological material. defining the geology and soil formations is often difficult along this rupture section of the mountain where the upper sedimentary plates have torn and became mixed with volcanic material (barker 1979, 1983; bumby 2000). should vegetation be removed, soil erosion becomes a major problem along the steep southern slopes with its relatively shallow soils and high rainfall. this major vegetation type is dependent on the orographic rain driven onto the southern slopes by a south-easterly wind during summer. the evergreen high forests are confined to the mistbelt of the mountain, which reaches down as far as 1380 m above sea level (geldenhuys & murray 1993). deciduous shrub forest forms a fire resistant ecotone of thickets, which extends to below the mistbelt zone of the southern slopes. the diagnostic species for this major vegetation type are presented in species group v (table 1). the diagnostic woody species include xymalos monospora, zanthoxylum davyi, celtis africana, nuxia floribunda, rhoicissus tomentosa, kiggelaria africana, vepris lanceolata, rapanea melanophloeos, rothmannia capensis, brachylaena discolor, ficus craterostoma, combretum kraussii, trichilia dregeana, trimeria grandifolia, drypetes gerrardii and oxyanthus speciosus subsp. gerrardii. other prominent woody species include diospyros whyteana (species group w), maytenus undata, rhus chirindensis, cussonia spicata (species group x) and maesa lanceolata (species group y). a recent classification of forest vegetation data by von maltitz et al. (2003) merged the forest communities of the blouberg, soutpansberg, north eastern escarpment, mariepskop and barberton regions under the name northern mistbelt forest. these afrotemperate forests have been described by numerous authors, and under various different names, such as afromontane forest (cooper 1985, white 1978), temperate, transitional and scrub veld types (acocks 1953), uplands vegetation (edwards 1967; moll 1976), interior forests (macdevette et al. 1989), montane podocarpus forest (cooper 1985; edwards 1967; moll 1976), highland sourveld (acocks 1953), mist belt mixed podocarpus forest (cooper 1985; edwards 1967; moll 1976) and natal mist belt ‘ngongoni veld (45) (acocks 1953). the evergreen xymalos monospora–rhus chirendensis soutpansberg forest vegetation, which forms part of the northern mistbelt forests, share some floristic affinities with the highland sourveld, the dohne sourveld (44b) and the natal mist belt ‘ngongoni veld (45), as well as the coastal tropical forest patches associated with the ’ngongoni veld (5) (acocks 1953). important shared species include the valuable timber species podocarpus falcatus, podocarpus latifolius and ocotea kenyensis. these species have been exploited for financial gain to such an extent that they are no longer dominant indicator species within all the soutpansberg forest associations. they now only occur in relatively high abundances where difficult terrain has inhibited people from accessing these species. this phenomenon of low cover-abundance values of the two podocarpus species within the sc is therefore an artefact of human intervention (mccracken 1986; obiri et al. 2002). the historical prominence of the two podocarpus species therefore justifies the soutpansberg forests’ inclusion into the temperate forest category (acocks 1953). the widdringtonia nodiflora dominated cliff forests along the scarp faces of the southern slopes of the soutpansberg share some weak floristic and structural affinities with the highland sourveld (acocks 1953) and the widdringtonia nodiflora–podocarpus latifolius short forest of the waterberg (van staden & bredenkamp 2006). ordination the rich diversity of plant communities within the sc and bnr is closely associated with the topographic and geologic diversity of these areas. the variation in topography and geology of the landscape contributes to variation in localised climatic conditions, seasonal precipitation, predictability of precipitation, processes and tempos of weathering, degrees of nutrient leaching from the soil, accumulation tempo of organic matter in the soil and plant available moisture in the soil. an ordination of the vegetation data on these two areas revealed distinct groupings of relevés, which in turn represent different major vegetation types and plant communities. in order to display and interpret these vegetation groupings in an ecologically meaningful manner, it was decided to separate the two distinct clusters representing the sc and the bnr. vegetation of the blouberg nature reserve a scatter diagram (fig. 4) displays the distribution of relevés along the second and third ordination axes. the vegetation units are represented as groups and their distribution on the scatter diagram corresponds with certain physical environmental conditions. the lack of any distinct groupings, with some relative distance between groupings, reflects the floristic similarities among the plant communities of the bnr. however, numerous environmental gradients could be correlated with the distribution of the plant communities and individual relevés along the xand y-axes. the second axis (eigen value = 0.5828), orientated along the xaxis, represents gradients in soil depth, soil moisture, surface rock cover, altitude, slope, plant-available moisture and soil texture. those plant communities along the left side of the diagram represent the dry, flat areas with relatively deep fine44 vol. 50 no. 1 pp. 32 48 major vegetation types original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe grained soils. these include the eragrostis lehmanniana var. lehmanniana–sclerocarya birrea subsp. caffra blouberg northern plains bushveld and the euclea divinorum–acacia tortilis blouberg southern plains bushveld. those plant communities along the right side of the diagram represent the wetter steep, rocky slopes with relatively shallow coarse sandy soils. these are mostly represented by the englerophytum magalismontanum– combretum molle blouberg mountain bushveld. the third axis (eigen value = 0.3918), orientated along the yaxis, represents gradients in plant-available moisture, soil depth and slope. those plant communities along the top of the diagram represent the wetter, steep rocky slopes with relatively shallow coarse sandy soils, represented by the englerophytum magalismontanum–combretum molle blouberg mountain bushveld. those plant communities along the bottom of the diagram represent the dry flat areas with relatively deep finegrained soils, represented by the eragrostis lehmanniana var. lehmanniana–sclerocarya birrea subsp. caffra blouberg northern plains bushveld and the euclea divinorum–acacia tortilis blouberg southern plains bushveld. vegetation of the soutpansberg conservancy the scatter diagram (fig. 5) displays the distribution of relevés along the first and second ordination axes. the vegetation units are represented as groups and their distribution on the scatter diagram corresponds with certain physical environmental conditions. the very distinct groupings of relevés within the virtual space of the scatter plot are a good indication of the unique nature of each of the different plant communities it represents. these pronounced ordination clusters reinforce those groupings created by the numerical classification done on the available vegetation data of the sc. numerous environmental gradients could be correlated with the distribution of the plant communities and individual relevés along the xand yaxes. the first axis (eigen value = 0.9501), orientated along the xaxis, represents gradients in plant-available soil moisture and soil clay content. those plant communities along the left side of the diagram represent the arid areas with relatively sandy soils. these include the adansonia digitata–acacia nigrescens soutpansberg arid northern bushveld and diplorhynchus condylocarpon–burkea africana soutpansberg leached sandveld. those plant communities along the right side of the diagram represent the relatively moist areas with soils that have a higher clay content. these include the catha edulis–flueggia virosa soutpansberg moist mountain thickets and the xymalos monospora–rhus chirendensis soutpansberg forest vegetation. the second axis (eigen value = 0.6412), orientated along the y-axis, represents gradients in plant-available moisture, soil mineral content and altitude. those plant communities along the top of the diagram represent the lower lying mineral rich arid areas of the sc. these include the adansonia digitata–acacia nigrescens soutpansberg arid northern bushveld and catha edulis–flueggia virosa soutpansberg moist mountain thickets. those plant communities along the bottom of the diagram represent the moist high altitude major vegetation types on the leached soils of the sc. this includes the. rhus rigida var. rigida– rhus magalismontanum subsp. coddii soutpansberg mistbelt vegetation. discussion the results of both the ordination and the classification indicate a considerable difference between the vegetation types of the blouberg nature reserve and the soutpansberg conservancy (fig. 4 & fig. 5; table 1). species group f is restricted to the bnr, showing the relationship among the bnr plant communities, and indicates that it has unique vegetation. although individual major vegetation types of the sc are exceptionally unique (with many diagnostic species, e.g. species groups g, i, m, q and v), no species groups emphasise the coherence of the vegetation of the sc. on the contrary, species groups h, k, l, o, p, u and y indicate the relationship of sc plant communities with some bnr communities. all these heterogeneous vegetation types and plant communities fall within an estimated area of only 108 000 hectares (1 080 km2). the conservation of such a biological hotspot should be a provincial, national and international priority (unesco 2002; walker 1989; wessels et al. 2003). the soutpansberg conservancy and the blouberg nature reserve represent numerous major vegetation types of the scbd, and form the core of present conservation efforts. these nature reserves should act as benchmark sites for the monitoring of the impact of development on the surrounding unprotected areas. the observed diversity in vegetation patterns is the result of the region’s high spatial variation (topography, geology, pedology, extreme localised climate patterns) as well as the region’s high temporal variability (irregular climatic cycles, periodic stochastic events) (gibson et al. 2004). in addition to the environmental factors influencing the observed vegetation and floristic patterns, certain regions of the study area have been altered through intense anthropogenic activities over extended periods of time. the stochasticity with which humans have impacted on this environment has led to even higher levels of spatial and temporal variation in habitat heterogeneity. numerous stone tools and artefacts indicate that humans have occupied the region on a periodic basis since the early stone age (coles & higgs 1975). more recent times have seen cultures and civilisations such as the khoisan (eloff 1979), the people of mapungubwe (huffman 1996), the vhenda people (nemudzivhadi 1985), and european settlers utilising the region for hunting, livestock farming and the cultivation of crops (voigt & plug 1984). it is important to identify and to understand the major ecological processes driving this particular ecosystem in order to conserve and manage it effectively (wessels et al. 2003). the major vegetation patterns seen among the plant communities of the sc and bnr are largely related to the availability of soil moisture and the rate of environmental desiccation (bond et al. 2003). the underlying geology and soils as well as altitude seem to play only secondary roles in the community composition of these event-driven systems (krebs 2001). soil moisture availability within the study area is governed by mainly four environmental factors: (1) the amount of precipitation of atmospheric moisture, (2) the rate of water loss through evaporation (3) the soil’s ability to capture and keep moisture within reachable depth of the plant roots, and (4) the available soil water capacity (kramer 1969; scott & le maitre 1998). white (1995) defines the available soil water capacity as the amount of water in a soil that is available for plant growth. the upper limit is set by the soil’s field capacity (water-saturated soil), while the lower limit is set by the volumetric water content value at which plants lose turgor and wilt, or the permanent wilting point. the sc and bnr are surrounded by many poverty stricken informal settlements of rural venda. these people rely on the savanna and forest plant communities to supply grazing, fuel wood, timber and agricultural produce. approximately 58% of the province’s land area is used for grazing, and 22% used for agriculture (hoffman & ashwell 2000). the limpopo province contributes considerably to the formal economy through its ecotourism, livestock, mining, timber and export fruit and vegetable industries (adams et al. 2000). sadly though, this culturally, historically and naturally rich and diverse province of south africa is a poverty stricken region (shackleton & shackleton 2000). the province’s rural communities are often 45vol. 50 no. 1 pp. 32 48 original research mostert, bredenkamp, klopper, verwey, mostert & hahn koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za locked in a struggle for survival against the frequent and severe droughts. this has led to a culture where “if it does not pay, it does not stay” (goudie 2000). in addition, the ever-expanding population of south africa is making increasing demands on the natural resources of the limpopo region. this will inevitably lead to the expansion of agriculture and industry into marginal and ecologically sensitive areas. in order for the government to plan development, management and conservation sensibly, we need the necessary ecological knowledge of the area. without this baseline information and insight on the region’s driving ecological processes and patterns, the much needed development of the limpopo’s infrastructure and the utilisation of its natural resources will be unsustainable, with only limited short-term benefits for a few selected individuals. acknowledgements we thank the referees for their constructive comments on the manuscript, the national research foundation for funding this project and all the landowners within the sc and the management of the bnr for access to gather field data. thank you to goro lodge, lajuma mountain retreat, medike mountain reserve, and the soutpansberg herbarium for accommodation during fieldwork. references acocks, j.p.h. 1953. veld types of south africa. memoirs of the botanical survey of south africa, 28: 1–192. (3rd edition with updated names and illustrations published in 1988 as memoirs of the botanical survey of south africa 57: 1–146). adams, m., cousins, b. & manona, s. 2000. land tenure and economic development in rural south africa constraints and opportunities. in: cousins, b. (ed.) at the crossroads: land and agrarian reform in south africa into the 21st century, cape town: university of the western cape, pp. 111-128. anderson, j.m. (ed.). 2001. towards gondwana alive, vol. 1 (assoc. eds. berger, l., de wit, m., fatti, l.p., holm, e., rubidge, b., smith, g., thackeray, f., & van wyk, b.). pretoria: gondwana alive society. barker, o.b. 1979. a contribution to the geology of the soutpansberg group, waterberg supergroup, northern transvaal. msc thesis, johannesburg: university of the witwatersrand. barker, o.b. 1983. a proposed geotectonic model for the soutpansberg group within the limpopo mobile belt, southern africa. in: van biljon, w.j. and j.h. legg (eds.). the limpopo belt. geological society of south africa. volume 8: 181–190. berger, k., crafford, e., gaiger, i., gaiger, m.j., hahn, n., & macdonald, i. 2003. a first synthesis of the environmental, biological and cultural assets of the soutpansberg. makhado: leach printers & signs. bezuidenhout, h., bredenkamp, g.j. & theron, g.k. 1994. phytosociological classes of the western transvaal grassland, south africa. koedoe, 37(1): 1–18. bond, w.j., midgley, g.f. & woodward, f.i. 2003. what controls south african vegetation – climate or fire? south african journal of botany, 69(1): 79–91. botha, m.j. 2004a. soil mineralogy. in: macvicar, c.n. (ed.). land types of the map: 2228 alldays. memoirs on the agricultural natural resources of south africa. department of agriculture and water supply. available at: www.arc. agric. za/institutes/iscw/main/maps/htm [accessed 2004] botha, m.j. 2004b. soil mineralogy. in: macvicar, c.n. (ed.). land types of the map 2328: pietersburg. memoirs on the agricultural natural resources of south africa. department of agriculture and water supply. available: www.arc.agric. za/institutes/iscw/main/maps/htm [accessed 2004]. branch, w.r. 1988. south african red data book – reptiles and amphibians. south african national scientific programmes report 151. pretoria: csir. brandl, g. 2002. the geology of the alldays area. explanation, sheet 2228: alldays. council for geoscience, south africa, pretoria. bredenkamp, g.j. 1975. ’n plantsosiologiese studie van die suikerbosrand-natuurreservaat. msc thesis, pretoria: university of pretoria. bredenkamp, g.j., joubert, a.f. & bezuidenhout, h. 1989. a reconnaissance survey of the vegetation of the plains in the potchefstroom-fochville-parys area. south african journal of botany 55(2): 199–206. bredenkamp, g.j. & van rooyen, n. 1996. 34. rocky highveld grassland. in: low, a.b. and a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. p 39, pretoria: department of environmental affairs and tourism. brown, l.r. 1997. a plant ecological study and wildlife management plan of the borakalalo nature reserve, north west province. phd thesis. pretoria: university of pretoria. bumby, a.j. 2000. the geology of the blouberg formation, waterberg and soutpansberg groups in the area of blouberg mountain, northern province, south africa. phd thesis. pretoria: university of pretoria. coetzee, b.j. 1975. a phytosociological classification of the rustenburg nature reserve. bothalia, 11: 561–580. coetzee, b.j., van der meulen, f., zwanziger, s., gonsalves, p. & weisser, p.j. 1976. a phytosociological classification of the nylsvlei nature reserve. bothalia 12: 137–160. coles, j.m. & higgs, e.s. 1975. the archaeology of early man. great britain: peregrine books. cooper, k.h. 1985. the conservation status of indigenous forests in transvaal, natal and o.f.s., south africa. durban: wildlife society of south africa. du preez, p.j. & bredenkamp, g.j. 1991. vegetation classes of the southern and eastern orange free state (republic of south africa) and the highlands of lesotho. bloemfontein: navorsinge van die nasionale museum, volume 7, pp. 477–526. edwards, d. 1967. a plant ecological survey of the tugela river basin. botanical survey of south africa memoir no. 36. pietermaritzburg: town and regional planning commission. edwards, d. 1983. a broad–scale structural classification of vegetation for practical purposes. bothalia, 14: 705–712. eloff, j.f. 1979. iii die kulture van greefswald – ‘n argeologiese studie van die ystertydperkkulture op die plaas greefswald. pretoria: university of pretoria. foord, s.h., dippenaar–schoeman, a.s. & van der merwe, m 2002. a checklist of the spider fauna of the western soutpansberg, south africa (arachnida: araneae). koedoe, 45(1): 35–43. foord, s.h. & dippenaar-schoeman, a.s. 2003. spiders (araneae). in: berger, k., craffort, j.e., graiger, i., graiger, m.j., hahn, n. & macdonald, i. (eds.). a first synthesis of the environmental, biological and cultural assets of the soutpansberg. makhadu: leach printers & signs, pp. 45–50. fuls, e.r., bredenkamp, g.j., van rooyen, n. & theron, g. k. 1993. the physical environment and major plant communities of the heilbron–lindley–warden–villiers area, northern orange free state. south african journal of botany, 59(4): 345–359. geldenhuys, c.j. & murray, b. 1993. floristic and structural composition of hanglip forest in the soutpansberg, northern transvaal. south african forestry journal, 165: 9–20. germishuizen, g. & meyer, n.l. 2003. plants of southern africa: an annotated checklist. strelitzia, 14: 1–1231. gertenbach, w.p.d. 1983. landscapes of the kruger national park. koedoe, 26: 9–121. gibson, l.a., wilson, b.a. & aberton, j.g. 2004. landscape characteristics associated with species richness and occurrence of small native mammals inhabiting a coastal heathland: a spatial modelling approach. biological conservation, 120(1): 75–89. goudie, a. 2000. the human impact on the natural environment. oxford: blackwell publishers. 46 vol. 50 no. 1 pp. 32 48 major vegetation types original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe hahn, n. 1994. tree list of the soutpansberg. pretoria: fantique publishers. hahn, n. 1996. rubiaceae. a new species of vangueria from the soutpansberg. bothalia, 27(1): 45–48. hahn, n. 1997. rare and endangered flora of tshikondeni. inhouse manuscript of the tshikondeni mining company (pty) ltd. tshikondeni. hahn, n. 1999. rubiaceae. a new species of pavetta from the soutpansberg, south africa. bothalia, 29(1): 107–109. hahn, n. 2002. endemic flora of the soutpansberg. msc thesis, pietermaritzburg: university of natal. hahn, n. 2003. endemic flora. in: berger, k., crafford, e., gaiger, i., gaiger, m.j., hahn, n. & macdonald, i. (eds.). a first synthesis of the environmental, biological and cultural assets of the soutpansberg. makhado, leach printers & signs. hahn, n. endemism and biogeography. phd thesis, in preparation. pretoria: university of pretoria. harrison, j.a., allan, d.g., underhill, l.g., herremans, m., tree, a.j., parker, v. & brown, c.j. 1997. the atlas of southern african birds. avian demography unit, bird life south africa. hennekens, s.m. & schamineé, j.h.j. 2001. turboveg, a comprehensive database management system for vegetation data. journal of vegetation science, 12: 589–591. henning, b.j. 2002 the relevance of ecosystems to ecotourism in the waterberg biosphere reserve. phd thesis, pretoria: university of pretoria. hill, m.o. 1979a. twinspan – a fortran program for arranging multivariate data in an ordered two-way table by classification of individuals and attributes. new york: cornell university. hill, m.o. 1979b. decorana – a fortran program for detrended correspondence analysis and reciprocal averaging. new york: cornell university. hockey, p.a.r., dean, w.r.j. & ryan, p.g. 2005. roberts – birds of southern africa (7th edition). the trustees of the john voelcker bird book fund, cape town. hoffman, t. & ashwell, a. 2000. land degradation: northern province fact sheet. cape town, salty print. huffman, t. n. 1996. archaeological evidence for climatic change during the last 2000 years in southern africa. quaternary international, 33: 55–60. kent, m. & coker, p. 1996. vegetation description and analysis: a practical approach. john wiley & sons ltd., sussex, england. kramer, p.j. 1969. plant and soil water relationships: a modern synthesis. new delhi, united states of america, mcgraw– hill publishing company. krebs, c.j. 2001. ecology: the experimental analysis of distribution and abundance (5th edition). san francisco: benjamin cummings. louw, a.j. 1970. mopane veld noord van die soutpansberg. dsc thesis, pretoria: university of pretoria. lubke, r.a. & mackenzie, b. 1996. afromontane forest. in: low, a.b. and a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism, p. 12. macdevette, d. r. 1993. the woody vegetation of the zululand coastal dunes. in: van der sejde (ed.). south african forestry handbook/suid-afrikaanse bosbouhandboek. southern african institute of forestry, pretoria, pp. 633–637. macdevette, d.r., mac devette, d.k., gordon, i.g. & bartholomew, r.i.c. 1989. floristics of the natal indigenus forests of southern africa. occasional report 45. pretoria: foundation for research and development. macvicar, c.n., bennie, a.t.p. & de villiers, j.m. 1991. soil classification: a taxonomic system for south africa. pretoria: department of agricultural development. matthews, w.s. 1991. phytosociology of the north-eastern mountain sourveld. msc thesis, pretoria: university of pretoria. matthews, w.s., bredenkamp, g.j. & van rooyen, h. 1991. the grassland-associated vegetation of the black reef quartzite and associated large rocky outcrops in the north-eastern mountain sourveld of the transvaal escarpment. south african journal of botany, 57: 143–150. matthews, w.s., bredenkamp, g.j. & van rooyen, n. 1994. the phytosociology and syntaxonomy of relatively low-altitude areas in the north-eastern mountain sourveld, in the eastern transvaal escarpment region. koedoe, 37(2): 73–87. mccracken, d.p. 1986. the indigenous forests of colonial natal and zululand. natalia, 16: 19–38. mccune, b. & mefford, m.j. 1999. pc-ord for windows. multivariate analysis of ecological data, version 4.10. mjm software, gleneden beach, oregon, u.s.a. moll, e.j. 1976. the vegetation of the three rivers region, natal. natal town and regional planning reports 33. pietermaritzburg: town and regional planning commission. mueller-dombois, d. & ellenberg, h. 1974. aims and methods of vegetation ecology. new york: wiley. nemudzivhadi, m. 1985. when and what? an introduction to the evolution of the history of venda. thohoyandou: department of land affairs. obermeyer, a.a., scweickerdt, h.g. & verdoorn, i.c. 1937. an enumeration of the plants collected in the northern transvaal. bothalia, 3(2): 223–258. obiri, j., lawes, m. & mukolwe, m. 2002. the dynamics and sustainable use of high-value tree species of the coastal pondoland forests of the eastern cape province, south africa. forest ecology and management, 199: 131–148. olivier, j. & rautenbach, c.j. 2002. the implementation of fog water collection systems in south africa. atmospheric research, 64(4): 227–239. patterson, d.g. & ross, p.g. 2004a. soil mineralogy. in: macvicar, c.n. (ed.). land types of the map 2228 alldays. memoirs on the agricultural natural resources of south africa. department of agriculture and water supply. available: www.arc.agric.za/institutes/iscw/main/maps/ htm [accessed 2004]. patterson, d.g. & ross, p.g. 2004b. soil mineralogy. in: macvicar, c.n. (ed.). land types of the map 2328 pietersburg. memoirs on the agricultural natural resources of south africa. department of agriculture and water supply. available at: www.arc.agric.za/institutes/iscw/main/maps/htm [accessed 2004]. schutte, j.m. 1971. die onttrekking van water uit die newellaag en lae wolke by mariepskop. s.a. department of water affairs, division hydrological research, technical note 20: 1–21. scott, d.f. & le maitre, d.c. 1998. the interaction between vegetation and groundwater: research priorities for south africa. pretoria, wrc report no. 730/1/98. water research commission. shackleton, c.m. & shackleton, s.e. 2000. direct use values of secondary resources harvested from communal savannas in the bushbuckridge lowveld, south africa. journal of tropical forest products 6: 28–47. siebert, s.j. 2001. vegetation on the ultramafic soils of the sekhukhuneland centre of endemism. phd thesis, pretoria: university of pretoria. siebert, f., bredenkamp, g.j. & siebert, s.j. 2003. a comparison of mopane vegetation in south africa, namibia and zimbabwe. bothalia, 33(1): 121–134. smit, j.h.l. 2000. phytosociology and veld management of the eastern kalahari thornveld. msc thesis, pretoria: university of pretoria. south african weather buro 2004. south african weather service. available: http://www.weathersa.co.za [accessed 2004]. stirton, c.h. 1982. a new species of rhynchosia from venda. bothalia, 14(1): 76–77. unesco. 2002. convention concerning the protection of the world cultural and natural heritages. unesco world heritage centre, france http://www.unesco.org [accessed 2004] 47vol. 50 no. 1 pp. 32 48 original research mostert, bredenkamp, klopper, verwey, mostert & hahn koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za van den berg, h.m. 1993. die identifisering en kartering van veldekotope in die noordwes-transvaalse suuren gemengdebosveld. msc thesis, pretoria: university of pretoria. van der meulen, f. 1979. plant sociology of the western transvaal bushveld, south africa: a syntaxonomic and synegological study. dissertationes botanicae, 49: 1–192. van jaarsveld, e. & hardy, d. 1991. the blouberg botanical expedition, with specific reference to the succulent vegetation. aloe, 28(3): 78–83. van rooyen, n. & bredenkamp, g.j. 1996. 17. sweet bushveld.. in: low, a.b. and a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. p. 21. van staden, p.j. 2002. an ecological study of the plant communities of marakele national park. msc thesis, pretoria: university of pretoria. van staden, p.j. & bredenkamp, g.j. 2005. major plant communities of the marakele national park. koedoe, 48(2): 59–70. van staden, p.j. & bredenkamp, g.j. 2006. a floristic analysis of forest and thicket vegetation of the marakele national park. koedoe, 49(1): 15–31. van wyk, a.e. 1984. a new species of combretum from venda and taxonomic notes on the section angustimarginata (combretaceae). south african journal of botany, 3(2): 125– 135. van wyk, a.e. & smith, g.f. 2001. regions of floristic endemism in southern africa. pretoria: umdaus press. van wyk, p. 1994. field guide to the trees of the kruger national park. cape town: struik publishers. van wyk, a.e. 1996. biodiversity of the maputaland centre. in: van der maesen, l.j.g., x.m. van der burgt and j.m. van medenbach de rooy (eds.). the biodiversity of african plants. wageningen: kluwer academic press. pp. 198–207. voigt, e.a. & plug, i. 1984. happy rest: the earliest iron age fauna from the soutpansberg. south african journal of science, 80: 22–227. von maltitz, g.p., mucina, l., geldenhuys, c., lawes, m., eeley, h., adie, h., vink, d., flemming, g. & voster, t. 2003. classification system for south african indigenous forests: an objective classification for the department of water affairs and forestry. environmentek report env–p–c 2003–017. pretoria: csir. walker, b. 1989. diversity and stability in ecosystem conservation. in: western, d. and m. pearl (eds.). conservation for the twenty-first century. new york: oxford university press. westoby, m., walker, b. & noy-meir, i. 1989. opportunistic management for rangelands not at equilibrium. journal of rangeland management, 42(4): 266–274. wessels, k.j., reyers, b., van jaarsveld, a.s. & rutherford, m.c. 2003. identification of potential conflict areas between land transformation and biodiversity conservation in north-eastern south africa. agriculture, ecosystems and environment, 95: 157–178. white, f. 1978. the afromontane region. in: werger, m.j.a. (ed.). biogeography and ecology of southern africa. the hague, junk, pp. 463–513. white, r.e. 1995. introduction to the principles and practices of soil science (2nd edition). oxford: blackwell sciences. winterbach, r. 1998. a phytosociological synthesis of acacia tortilis communities in the north-western savanna of south africa. msc thesis, pretoria: university of pretoria. winterbach, r., bredenkampl, g.j., deutschlander, m.s. & mucina, l. 2000. preliminary syntaxonomic scheme of vegetation classes for the central bushveld of south africa. in: whyte, p.s., mucina, l., leps, j.s. & van der maarel, e. (eds.). proceedings iavs symposium. upsala: opulus press. pp. 123–127. 48 vol. 50 no. 1 pp. 32 48 abstract introduction why are estuaries important to fishes? current protection status for fishes in south african estuaries fish species examples of why estuarine-protected areas are required some reasons why existing law enforcement measures are not working conclusion acknowledgements references about the author(s) alan k. whitfield south african institute for aquatic biodiversity, makhanda, south africa colin g. attwood department of biological sciences, faculty of science, university of cape town, rondebosch, south africa paul d. cowley south african institute for aquatic biodiversity, makhanda, south africa stephen j. lamberth department of environment, forestry and fisheries, cape town, south africa bruce q. mann oceanographic research institute, durban, south africa citation whitfield, a.k., attwood, c.g., cowley, p.d., lamberth, s.j. & mann, b.q., 2020, ‘no-take estuarine-protected areas: the missing armour for the conservation of fishes’, koedoe 62(1), a1648. https://doi.org/10.4102/koedoe.v62i1.1648 essay no-take estuarine-protected areas: the missing armour for the conservation of fishes alan k. whitfield, colin g. attwood, paul d. cowley, stephen j. lamberth, bruce q. mann received: 10 july 2020; accepted: 27 aug. 2020; published: 05 nov. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the focus of conservation attention over the past few decades has been on marine-protected areas (mpas) providing protection for heavily exploited marine fish species. although some estuaries are included in most large mpas, specific attention on the protection needs of fish species in estuaries has been lacking. furthermore, many of the estuaries located within conservation areas are open to angling activities and this has been exacerbated by the proliferation of illegal gillnet fishing in many systems during recent decades. three fish species, the dusky kob argyrosomus japonicus, white steenbras lithognathus lithognathus and spotted grunter pomadasys commersonnii – are used as examples of estuary-dependent taxa whose populations have been decimated by fishing over-exploitation, as well as habitat degradation caused by various human activities. by having complete protection for vulnerable fish species in certain estuaries, the potential benefits of increased catches for both subsistence and recreational anglers along the entire south african coastline could be substantial. if such protection is not offered to these species, then the downward spiral in fish catches will continue, to the ultimate detriment of both the people who currently use these protein resources for food security and those who are part of the economically important recreational fishing industry. conservation implications: based on collapsing populations of targeted fishery species in estuaries, there is an urgent need to implement no-take estuarine-protected areas in each of the biogeographic regions along the south african coast. keywords: estuarine fishes; threatened species; protected areas; conservation; fisheries management; environmental management. introduction south africa was one of the few countries that took an early initiative in declaring marine-protected areas (mpas) for biodiversity conservation (attwood et al. 1997a; attwood, harris & williams 1997b). support for these protected areas intensified when it became clear that they could assist in the recovery of depleted fishery resources (attwood et al. 1997a; buxton 1993). although representation of biogeographic zones and habitats was recognised in 1995, the declaration of new mpas at the end of that decade did not address the paucity of estuarine fish protection that was highlighted (attwood et al. 1997b). despite the fact that a number of marine fish species use estuaries as facultative or obligate nursery areas, there was little acknowledgement of this link in the proclamation of mpas (whitfield 1997), a situation that has not changed in the intervening decades (whitfield & cowley 2010). more recent studies in south africa have provided further conclusive evidence of the benefits of no-take mpas to harvested fish stocks (kerwath et al. 2013; maggs, mann & cowley 2013; mann et al. 2016), including those found in estuarine-protected areas (attwood et al. 2007; da silva et al. 2013; hedger et al. 2010; padare et al. 2020). south african estuaries are home to 172 fish species (whitfield 2019). of these species, 37 (22%) are marine ‘stragglers’ and make very limited use of estuaries. marine fish species that use estuaries as nurseries and/or foraging areas comprise 73 (42% of the total), but only 36 (21%) of these are closely associated with estuaries (whitfield 2019). fish species that are resident and breed within estuaries comprise 25% (43 species) of all the taxa recorded in these systems. there is a high level of endemism, with 38 fish species (22% of all estuary-associated taxa) occurring only in southern african waters (whitfield 2019). why are estuaries important to fishes? the importance of connectivity between marine and estuarine ecosystems is globally recognised (e.g. able 2005). in recent years, two diadromous fish species have gone extinct (allibone et al. 2010; freyhof & christian 2005), yet no fully marine fish species has gone extinct despite the latter being vastly more numerous and the target of industrial-scale fisheries speaks about the high risks confronted by estuary-dependent fish. our local piscine extinction candidates are either estuarine or freshwater. two species of sawfish pristis spp. have disappeared from south african estuaries (everett et al. 2015), and the estuarine pipefish syngnathus watermeyeri, once thought to be extinct (whitfield & bruton 1996), is not far from that listing (whitfield et al. 2017). estuaries are the conduits between marine and freshwater habitats – critical for the completion of life cycles of many fishes but threatened by land-based pollutants and growing numbers of catchment impoundments. past mpas did not factor in the important role of estuaries in the ecology of catadromous fish species, such as anguillid eels, whose glass eel larvae have to enter catchment rivers through estuaries (bruton, bok & davies 1987). although the eels spend more than a decade of development in the freshwater environment (mcewan & hecht 1984), the ability of the adults to reach spawning grounds in the indian ocean (tsukamoto, aoyama & miller 2002) is dependent on safe passage through healthy and protected estuarine environments. estuaries are possibly the most threatened habitat realm in the country. there are 22 estuarine types represented in south africa and 86% of these are threatened (van niekerk et al. 2019, 2020). about one-third of south africa’s freshwater is abstracted for agricultural, industrial and domestic use before it reaches the sea. this alone has reduced estuarine habitat, but what remains has been degraded further by habitat modification and pollution (van niekerk et al. 2019). the loss of fish nursery habitat, including saltmarsh, mangroves and intertidal foraging area, has effectively reduced a large number of fish populations by suppressing recruitment. over-exploitation poses the single biggest threat to estuarine fish conservation (whitfield & cowley 2010; table 1). although more than 200 species are caught in south africa’s marine linefishery, very few species are targeted in estuarine fisheries, with the dominant species being the spotted grunter pomadasys commersonnii, dusky kob argyrosomus japonicus, leervis lichia amia, white steenbras lithognathus lithognathus and cape stumpnose rhabdosargus holubi (e.g. baird, marais & daniel 1996; cowley et al. 2004). the number of species targeted in subtropical estuaries is slightly higher (james et al. 2001) than those in the more temperate systems. with the exception of cape stumpnose, the stock status of all the highly targeted species is considered to be either over-exploited or collapsed. table 1: estimates of the relative importance of current major fish conservation issues in (1) freshwater, (2) estuarine and (3) marine ecosystems in south africa. a pressing need for protecting estuaries resides with several iconic fish families that contain species targeted by recreational and subsistence fishers. the main use of estuaries by marine fish taxa in south africa is as 0+ juveniles that usually spend between 1 and 3 years in these habitats (wallace & van der elst 1975). species belonging to the families mugilidae, sparidae, haemulidae, sciaenid and carangid are particularly well represented in estuaries on the subcontinent. members of these families are economically important and well represented in estuarine and coastal fisheries. the juveniles of most of these species are largely restricted to estuaries but make occasional forays into the open sea (e.g. bennett et al. 2017; dames et al. 2017; grant et al. 2017). once they approach adulthood, these individuals return to the sea on a more permanent basis, where they are targeted with a variety of fishing methods (e.g. childs et al. 2015; murray et al. 2018). these fish contribute more than 90% of the catch of south africa’s shore-based and nearshore marine fisheries (lamberth & tripe 2003). current protection status for fishes in south african estuaries over 3730 tonne of fish is caught annually in south africa’s estuaries, of which 2200 tonne (60%) is by the illegal gillnet fishery (van niekerk et al. 2019). these estimates are conservative as the combined legal and illicit catch in st lucia alone may be over 300 tonne to as much as 800 tonne per annum (mann 1995; turipe et al. 2014). of over 500 gillnets confiscated from estuaries throughout south africa that had some degree of protection in the form of conservator or fishery patrols, 60% were retrieved before having caught any fish. this implies that even current low levels of protection (and support for conservators) are helping to keep estuarine gillnet catch from surging past 4000 tonne per year. increasing the number of estuarine-protected areas, accompanied by well-trained, motivated staff would most likely reduce gillnet catch to even lower levels. the latest assessment of the coverage of estuarine-protected areas (epas), of which several types exist, shows that 48% of estuarine habitat is protected, but that the vast majority of this protection is either partial or ineffective (box 1; van niekerk et al. 2019). turipe, wilson and van niekerk (2012) modelled that 133 south african estuaries, including those already protected, would be required to meet defined biodiversity targets, including protection of fish. of these, 61 should be fully protected no-take estuaries, whereas 72 require partial protection, including zoned no-take areas. this represents 46% of estuaries and 79% of estuarine area. consequently, all epas should be explicit in the individual estuary management plans required in terms of the integrated coastal management act (act 24 of 2008). box 1: steps required for recovery of over-exploited fish species in south african estuaries. current estuarine ecosystem protection levels are low, both in terms of number of types and in area. overall, 82% (19 out of 22 types) of south africa’s estuarine ecosystem types are under-protected. of estuarine area, less than 2% is well protected, 24% moderately protected, 63% poorly protected and 11% not-protected (van niekerk et al. 2019). seventy-two estuaries are in terrestrial protected areas and flow into mpas, but few have any no-take status. only 25 (8%) of estuaries have no-take zones, representing less than 1% of total estuarine area. this is because of most of these being very small systems that fall entirely within an mpa (e.g. klipdriftsfontein in de hoop) but make a minimal contribution to total estuarine fish biomass or productivity (these very small systems are excluded from figure 1). figure 1: important south african functional estuaries (from a fish diversity and production perspective) that have a formally declared conservation status, even though fish stocks within these areas are often not protected from fishing activities. very small protected estuaries and those larger systems that have only a minor portion of the water area conserved are not shown. the inclusion of estuaries within an mpa, or other types of protected areas, does not imply that fish are protected, as recreational and subsistence fishing in protected estuaries is usually permitted, for example, mbhashe and mbhanyana where angling effort is five times the national average. many estuarine systems that have a national, provincial or municipal conservation status have management authorities that allow fishing, for example, knysna and goukamma. langebaan is a ramsar site and foundation of the west coast national park but supports a legal commercial gillnet fishery as well as commercial, small-scale and recreational line fishing. the combined catch exceeds 250 tonne per annum. our concern is that the word ‘protection’ has been misapplied and that the two most common deficiencies in protected areas are the management of fishing and water quality (van niekerk et al. 2019). important estuaries (from a fish diversity and production perspective) currently situated in the footprint of conservation areas are concentrated in the eastern and south-eastern parts of the country (figure 1). the largest and most important estuary in south africa is the lake st lucia system, which covers almost 50% of the estuarine surface area in the country. although forming a part of the world heritage site and a ramsar site of international importance, the fishes of the st lucia system are not protected from fishing and illegal gill netting is common (mann 1995, 2003). furthermore, this system has been effectively closed to the sea since 2002, thus greatly reducing its nursery function for marine species (vivier, cyrus & jerling 2010). also part of the isimangaliso wetland park is the large kosi estuarine lake system that, in addition to recreational angling, permits the extensive use of modified traditional thonga fish traps and is simultaneously plagued by illegal gill netting (kyle 2013). clearly, the conservation status of fishes within these key estuarine systems needs to be reviewed and elevated. fish species examples of why estuarine-protected areas are required why are effective epas for fishes such an urgent requirement for the sustainability of fish stocks all along the south african coast? just three examples should provide ample evidence to support epa proclamations, not as addendums to mpas, but as fully protected areas in their own right. the first species is the dusky kob a. japonicus, a member of the sciaenidae family, which is the primary target of most recreational and subsistence fishers in the subtropical and warm temperate regions (cowley, childs & bennett 2013; crook & mann 2002; mann, james & beckley 2002; potts et al. 2005). dusky kob is now estimated to have a spawner stock biomass below 3% of their pre-exploitation biomass (winker et al. 2015). the south african dusky kob is now considered to be endemic to the subcontinent and is genetically distinct from a. japonicus in the rest of the indo-pacific region (barnes et al. 2016). an additional threat is the reported hybridisation with the heavily depleted silver kob argyrosomus inodorus (mirimin et al. 2014). the second example is the white steenbras l. lithognathus (sparidae), which is targeted by fishers in warmand cold-temperate regions. this large seabream had a spawner stock biomass < 6% of pristine levels in 1990 (bennett 1993) and ongoing monitoring has suggested further declines since then (mann et al. 2014, http://dehoopfishmonitoring.uct.ac.za). the discovery of a handful of 0+ juveniles in the cunene estuary on the angola-namibia border attests to what is left of a large historical west coast spawning population depleted to near extinction by eight decades of intensive beach-seine netting straddling the late 19th and early 20th centuries. white steenbras bones and otoliths are a dominant fish component of archaeological middens on the west coast (poggenpoel 1996). by inference, it took just a few decades to nearly extirpate a population that had been in existence for more than 100 000 years. the third example relates to a tagging study by padare et al. (2020) on spotted grunter p. commersonnii (330 mm – 650 mm fork length) in the goukou estuary (western cape). the results of this study further confirm the high site and estuarine fidelity by spotted grunter and the potential value of estuarine no-take area closures as a management option. the high site fidelity suggests that the closure of even a section of an estuary has the potential to reduce the vulnerability to capture of a portion of the locally resident population. the vulnerability of this species to fishing pressure within estuaries was highlighted by the loss of 43% of the tagged fish. whilst this study revealed the potential benefit of no-take epas, the possible capture of several tagged fish within the no-take zone suggests that compliance is low and that law enforcement is inadequate at the goukou estuary. some reasons why existing law enforcement measures are not working whilst in theory improved management of the above iconic species could be achieved by catch restrictions, or even a total moratorium on catches of these species, law enforcement, and hence compliance with fishing regulations, is very poor across the country (bova et al. 2017). a recent detailed study conducted on the sundays estuary revealed that more than 90% of the targeting effort was levelled at only two species, dusky kob and spotted grunter (cowley et al. 2013). a large portion of the dusky kob catch comprised fish with lengths below the minimum legal size limit (i.e. 60 cm total length). the high retention rate of undersized fish is a major cause for concern. sixty-three per cent of dusky kob and all white steenbras catches were below the legal size limit of 60 cm, and 30% of spotted grunter were below the 40 cm size limit. size limits are a type of catch restriction that should become more effective as fishing pressure increases and reduces the average length of fish, but it still requires effective enforcement. the other commonly applied catch restriction, namely, bag limits, is more problematic. this type of restriction becomes less effective as fishing pressure increases (attwood & bennett 1995), as the following evidence demonstrates. the daily bag limit (one fish per person per day) of legal-sized dusky kob in the sundays estuary was attained on only 2.6% of angler outings, whilst that for spotted grunter (five fish per person per day) was attained on only 0.1% of outings. no white steenbras larger than the legal size were captured (cowley et al. 2013). the bag limits therefore have little practical value in these instances. a detailed questionnaire survey revealed that most anglers were unaware of the fishing regulations pertaining to their targeted species and that only 71% of interviewees claimed to have a valid fishing permit (cowley et al. 2013). the apparent lack of compliance can largely be ascribed to poor law enforcement. most (59%) of the respondents had never had their catch inspected, whilst 11% had encountered a law enforcement officer on only one previous outing. it is our opinion that epas (in which some or all of the estuary is protected from fishing) would offer far greater protection to estuarine fishery species, as long as their formal declaration includes dedicated management (conservation staff and budgets), as has been accorded to other protected areas. not surprisingly, both dusky kob and white steenbras are now listed as critically endangered on the international union for conservation of nature red list (fennessy 2020; mann et al. 2014). similarly, many estuary-associated marine fish species are on the world wildlife fund southern african sustainable seas initiative red list for the restaurant or retail trade as these may not be commercially sold or bought in south africa. these include leervis, tenpounder, kingfish species, spotted grunter, two stumpnose species, estuarine bream and mangrove snapper. conclusion the current political and economic environment makes it difficult to restrict fishing activities in any estuaries that are currently open to this activity. indeed, the trend in recent years has been to open up parts of mpas to fishing that were previously closed to angling (lombard et al. 2020). however, by having complete protection for vulnerable and over-exploited fish species in certain estuaries, the potential benefits of increased catches for both subsistence and recreational anglers along the entire south african coastline could be substantial. if such protection is not offered to these species, then the downward spiral in fish catches will continue, to the ultimate detriment of both the people who currently use these protein resources for food security and those who are part of the economically important recreational fishing industry. indeed, the long-term survival of certain endemic species in estuaries on the subcontinent could well be threatened if these fish populations drop below critical levels, and the possibility then arises that recovery of the stocks becomes increasingly unlikely. in conclusion, we recognise that the advantage of combining well-managed mpas and epas is the complete protection of at least some vulnerable fish species in a particular area during all stages of their life cycle. examples of this are the stilbaai mpa and greater addo np, which include no-take estuarine areas, and substantial parts of the adjacent coast. protected areas of this type, and community-based conservancies or stewardship initiatives, will greatly assist in achieving the recovery of over-exploited estuarine fish stocks and support the survival of critically endangered species, such as the estuarine pipefish. however, we also see a need to protect catchment connectivity, estuarine habitat and vegetation, which are all vital for young catadromous fish stages and the return migration of adult eels to the sea. to accomplish this, it will be necessary to declare additional epas and to substantially improve the management of existing epas to ensure that we have adequate protection of our essential fish nursery habitats. acknowledgements the authors thank their respective research organisations for financial, laboratory and logistical support to conduct fish research in south african estuaries. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions this article was initiated by a.k.w. and all authors made contributions to the writing of the review. ethical consideration this article followed all ethical standards for a research without direct contact with human or animal subjects. funding information this research received no specific grant from any funding agency in the public, commercial or not-for-profit sectors. data availability statement data sharing is not applicable to this article as no new data were created or analysed in this study. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references able, k.w., 2005, ‘a re-examination of fish estuarine dependence: evidence for connectivity between estuarine and ocean habitats’, estuarine and coastal shelf science 64(1), 5–17. https://doi.org/10.1016/j.ecss.2005.02.002 allibone, r., david, b., hitchmough, r., jellyman, d., ling, n., ravenscroft, p. & waters, j., 2010, ‘conservation status of new zealand freshwater fish, 2009’, new zealand journal of marine and freshwater research 44(4), 271–287. https://doi.org/10.1080/00288330.2010.514346 attwood, c.g. & bennett, b.a., 1995, ‘a procedure for setting daily bag limits on the recreational shore-fishery of the south-western cape, south africa’, south african journal of marine science 15(1), 241–252. https://doi.org/10.2989/025776195784156377 attwood, c.g., cowley, p.d., kerwath, s.e., naesje, t.f., okland, f. & thorstad, e.b., 2007, first tracking of white stumpnose rhabdosargus globiceps (sparidae) in a south african marine protected area’, african journal of marine science 29(1), 147–151. https://doi.org/10.2989/ajms.2007.29.1.15.80 attwood, c.g., harris, j.m. & williams, a.j., 1997b, ‘international experience of marine protected areas and their relevance to south africa’, south african journal of marine science 18(1), 311–332. https://doi.org/10.2989/025776197784161162 attwood, c.g., mann, b.q., beaumont j. & harris, j.m., 1997a, ‘review of the state of marine protected areas in south africa’, south african journal of marine science 18(1), 341–367. https://doi.org/10.2989/025776197784160910 baird, d., marais, j.f.k. & daniel, c., 1996, ‘exploitation and conservation of angling fish in two south african estuaries’, aquatic conservation: marine and freshwater ecosystems 6(4), 319–330. https://doi.org/10.1002/(sici)1099-0755(199612)6:4%3c319::aid-aqc201%3e3.0.co;2-h barnes, t.c., junge, c., myers, s.a., taylor, m.d., rogers, p.j., ferguson, g.j. et al., 2016, ‘population structure in a wide-ranging coastal teleost (a. japonicus, sciaenidae) reflects marine biogeography across southern australia’, marine and freshwater research 67(8), 1103–1113. https://doi.org/10.1071/mf15044 bennett, b.a., 1993, ‘the fishery for white steenbras lithognathus lithognathus off the cape coast, south africa, with some considerations for its management’, south african journal of marine science 13(1), 1–14. https://doi.org/10.2989/025776193784287185 bennett, r.h., cowley, p.d., childs, a.-r., attwood, c.g., swart, l. & næsje, t.f., 2017, ‘movement patterns of an endangered fishery species lithognathus lithognathus (sparidae) and the role of no-take marine protected areas as a management tool’, african journal of marine science 39(4), 475–489. https://doi.org/10.2989/1814232x.2017.1404493 bova, c.s., halse, s.j., aswani, s. & potts, w.m., 2017, ‘assessing a social norms approach for improving recreational fisheries compliance’, fisheries management and ecology 24(2), 117–125. https://doi.org/10.1111/fme.12218 buxton, c.d., 1993, ‘life-history changes in exploited reef fishes on the east coast of south africa’, environmental biology of fishes 36(1), 47–63. https://doi.org/10.1007/bf00005979 bruton, m.n., bok, a.h. & davies, m.t.t., 1987, ‘life history styles of diadromous fishes in inland waters of southern africa’, american fisheries society symposium 1, 104–121. childs, a.-r., cowley, p.d., næsje, t.f. & bennett, r.h., 2015, ‘habitat connectivity and intra-population structure of an estuary-dependent fishery species’, marine ecology progress series 537, 233–245. https://doi.org/10.3354/meps11456 cowley, p.d., childs, a.-r. & bennett, r.h., 2013, ‘the trouble with estuarine fisheries in temperate south africa, illustrated by a case study on the sundays estuary’, african journal of marine science 35(1), 117–128. https://doi.org/10.2989/1814232x.2013.789079 cowley, p.d., wood, a.d., corroyer, b., nsubuya, y. & chalmers, r., 2004, ‘a survey of fishery resource utilization on four eastern cape estuaries (great fish, west kleinemond, east kleinemond and kowie)’, in protocols contributing to the management of estuaries in south africa, with a particular emphasis on the eastern cape province, vol. iii, project c, supplementary report c5, pp. 129–165, water research commission, pretoria. crook, b.j.s. & mann, b.q., 2002, ‘a critique of and recommendations for a subsistence fishery, lake st lucia, south africa’, biodiversity and conservation 11(7), 1223–1235. https://doi.org/10.1023/a:1016074802295 dames, m.h., cowley, p.d., childs, a.-r., bennett, r.h., thorstad, e.b. & næsje, t.f., 2017, ‘estuarine and coastal connectivity of an estuarine-dependent fishery species, pomadasys commersonnii (haemulidae)’, african journal of marine science 39(1), 111–120. https://doi.org/10.2989/1814232x.2017.1305991 da silva, c., kerwath, s.e., attwood, c.g., thorstad, e.b., cowley, p.d., okland, f., wilke, c.g. & naesje, t.f., 2013, ‘quantifying the degree of protection afforded by a no-take marine reserve on an exploited shark’, african journal of marine science 35(1), 57–66. https://doi.org/10.2989/1814232x.2013.769911 everett, b.i., cliff, g., dudley, s.f.j., wintner, s.p. & van der elst, r.p., 2015, ‘do sawfish pristis spp. represent south africa’s first local extirpation of marine elasmobranchs in the modern era?’, african journal of marine science 37(2), 275–284. https://doi.org/10.2989/1814232x.2015.1027269 fennessy, s., 2020, argyrosomus japonicas, ‘the iucn red list of threatened species 2020’, e.t49145403a49234015. https://doi.org/10.2305/iucn.uk.2020-1.rlts.t49145403a49234015.en freyhof, j. & christian, s., 2005, ‘the houting coregonus oxyrinchus (l.) (salmoniformes: coregonidae), a globally extinct species from the north sea basin’, journal of fish biology 67(3), 713–729. https://doi.org/10.1111/j.0022-1112.2005.00771.x grant, g.n., cowley, p.d., bennett, r.h., murray, t.s. & whitfield, a.k., 2017, ‘space use by rhabdosargus holubi in a southern african estuary, with emphasis on fish movements and ecosystem connectivity’, african journal of marine science 39(2), 135–143. https://doi.org/10.2989/1814232x.2017.1327887 hedger, r.d., naesje, t.f., cowley, p.d., thorstad, e.b., attwood, c.g., okland, f. et al., 2010, ‘residency and migratory behaviour by adult pomatomus saltatrix in a south african coastal embayment’, estuarine, coastal and shelf science 89(1), 12–20. https://doi.org/10.1016/j.ecss.2010.04.013 james, n.c., beckley, l.e., mann, b.q. & kyle, r., 2001, ‘the recreational fishery in the kosi estuarine lake system, south africa’, african zoology 36(2), 217–228. https://doi.org/10.1080/15627020.2001.11657140 kerwath, s.e., winker, h., gotz, a. & attwood, c.g., 2013, ‘marine protected area improves yield without disadvantaging fishers’, nature communications 4(2347), 1–6. https://doi.org/10.1038/ncomms3347 kyle, r., 2013, ‘thirty years of monitoring traditional fish trap catches at kosi bay, kwazulu-natal, south africa, and management implications’, african journal of marine science 35(1), 67–78. https://doi.org/10.2989/1814232x.2013.769905 lamberth, s.j. & turpie, j.k., 2003, ‘the role of estuaries in south african fisheries: economic importance and management implications’, south african journal of marine science 25(1), 131–157. https://doi.org/10.2989/18142320309504005 lombard, a.t., durbach, i., harris, j.m., mann-lang, j.b., mann, b.q., branch, g.m. et al., 2020, ‘south africa’s tsitsikamma marine protected area – winners and losers’, in j. humphreys & r.w.e. clark (eds.), marine protected areas: science, policy and management, pp. 237–260, elsevier, amsterdam. maggs, j.q., mann, b.q. & cowley, p.d., 2013, ‘contribution of a large no-take zone to the management of vulnerable reef fishes in the south-west indian ocean’, fisheries research 144, 38–47. https://doi.org/10.1016/j.fishres.2012.10.003 mann, b.q., 1995, ‘quantification of illicit fish harvesting in the lake st lucia game reserve, south africa’, biological conservation 74(2), 107–113. https://doi.org/10.1016/0006-3207(95)00019-z mann, b.q., 2003, ‘the st lucia subsistence gillnet fishery’, in m. hauck & m. sowman (eds.), waves of change: coastal and fisheries co-management in southern africa, pp. 99–122. uct press, cape town. mann, b.q., buxton, c.d., pollard, d., carpenter, k.e. & iwatsuki, y., 2014, ‘lithognathus lithognathus. the iucn red list of threatened species 2014’, e.t12137a505458. https://doi.org/10.2305/iucn.uk.2014-3.rlts.t12137a505458.en mann, b.q., james, n.c. & beckley, l.e., 2002, ‘an assessment of the recreational fishery in the st lucia estuarine system, kwazulu-natal, south africa’, south african journal of marine science 24(1), 263–279. https://doi.org/10.2989/025776102784528330 mann, b.q., winker, h., maggs, j.q. & porter, s., 2016, ‘monitoring the recovery of a previously exploited surf-zone fish community in the st lucia marine reserve using a no-take sanctuary area as a benchmark’, african journal of marine science 38(3), 423–441. https://doi.org/10.2989/1814232x.2016.1224779 mcewan, a. & hecht, t., 1984, ‘age and growth of the longfin eel anguilla mossambica peters, 1852 (pisces, anguillidae) in transkei rivers’, african zoology 19(4), 280–285. https://doi.org/10.1080/02541858.1984.11447894 mirimin, l., kerwath, s.e., macey, b.m., bester-van der merwe, a.e., lamberth, s.j., bloomer, p. et al., 2014, ‘identification of naturally occurring hybrids between two overexploited sciaenid species along the south african coast’, molecular phylogenetics and evolution 76, 30–33. https://doi.org/10.1016/j.ympev.2014.02.010 murray, t., cowley, p., bennett, r. & childs a.-r., 2018, ‘fish on the move: connectivity of an estuary-dependent fishery species evaluated using a large-scale acoustic telemetry array’, canadian journal of fisheries and aquatic sciences 75(11), 2038–2052. https://doi.org/10.1139/cjfas-2017-0361 padare, g., cowley, p.d., vumazonke, l.u., murray, t.s., du plessis, j. & lamberth, s.j., 2020. ‘can partial estuarine area protection reduce the vulnerability to capture of spotted grunter pomadasys commersonnii?’, in t.s. murray, m.i. duncan, a.c. winkler, a.-r childs, b.q. mann & w.m. potts (eds.), linefish resilience in the anthropocene: the proceedings of the 5th southern african marine linefish symposium, 113 pp., rhodes university, makhanda. poggenpoel, c.a., 1996. ‘the exploitation of fish during the holocene in the south-western cape, south africa’, ma thesis, university of cape town, rondebosch. potts, w.m., cowley, p.d., corroyer, b. & næsje, t.f., 2005, ‘trends in fishery resource utilisation on the great fish estuary’, nina report 50, 34. turpie, j.k., feigenbaum, t., hayman, m., hutchings, k., cousins, t., chipeya, t. et al., 2014, ‘analysis of alternatives to determine the most feasible solution to the hydrological issues of the lake st lucia estuarine system, volume iv: socio-economic assessment’, report prepared by anchor environmental consultants for isimangaliso wetland park authority, kwazulu-natal. turpie, j.k., wilson, g. & van niekerk, l., 2012, ‘national biodiversity assessment 2011: national estuary biodiversity plan for south africa’, anchor environmental consultants report no aec2012/01, report produced for the council for scientific and industrial research and the south african national biodiversity institute, cape town. tsukamoto, k., aoyama, j. & miller, m.j., 2002, ‘migration, speciation, and the evolution of diadromy in anguillid eels’, canadian journal of fisheries and aquatic science 59(12), 1989–1998. https://doi.org/10.1139/f02-165 van niekerk, l., adams, j.b., james, n.c., lamberth, s.j., mackay, c.f., turpie, j.k. et al., 2020, ‘an estuary ecosystem classification that encompasses biogeography, estuary size and types in support of estuarine protection, conservation and management’, african journal of aquatic science 45(1–2), 199–216. https://doi.org/10.2989/16085914.2019.1685934 van niekerk, l., adams, j.b., lamberth, s.j., mackay, c.f., taljaard, s., turpie, j.k. et al. (eds.), 2019, ‘south african national biodiversity assessment 2018: technical report. volume 3: estuarine realm’, report no. sanbi/nat/nba2018/2019/vol3/a, south african national biodiversity institute, pretoria. vivier, l., cyrus, d.p. & jerling, h.l., 2010, ‘fish community structure of the st lucia estuarine system under prolonged drought conditions and its potential for recovery after mouth breaching’, estuarine, coastal and shelf science 86, 568–579. https://doi.org/10.1016/j.ecss.2009.11.012 wallace, j.h. & van der elst, r.p., 1975, ‘the estuarine fishes of the east coast of south africa. part 4. occurrence of juveniles in estuaries. part 5. ecology, estuarine dependence and status’, investigational report of the oceanographic research institute 42, 1–63. whitfield, a.k., 1997, ‘fish conservation in south african estuaries’, aquatic conservation: marine and freshwater ecosystems 7(1), 1–11. https://doi.org/10.1002/(sici)1099-0755(199703)7:1%3c1::aid-aqc213%3e3.0.co;2-8 whitfield, a.k., 2019, ‘fishes of southern african estuaries: from species to systems’, smithiana monograph no. 4, p. 495. whitfield, a.k. & bruton, m.n., 1996, ‘extinction of the river pipefish syngnathus watermeyeri in the eastern cape province, south africa’, south african journal of science 92(2), 59–60. whitfield, a.k. & cowley, p.d., 2010, ‘the status of fish conservation in south african estuaries’, journal of fish biology 76(9), 2067–2089. https://doi.org/10.1111/j.1095-8649.2010.02641.x whitfield, a.k., lamberth, s., cowley, p. & mann, b., 2019, ‘fisheries in south african estuaries – are we on the right road?’, the water wheel 18(6), 12–15. whitfield, a.k., mkare t., teske, p.r., james, n.c. & cowley, p.d., 2017, ‘life-histories explain the conservation status of two estuary-associated pipefishes’, biological conservation 212, 256–264. https://doi.org/10.1016/j.biocon.2017.06.024 winker, h., attwood, c. & kerwath, s., 2015. ‘assessment of stock abundance of inshore fish resources included in the “basket-of-species” to be allocated under the small-scale fisheries policy’, unpublished report, department of agriculture, forestry and fisheries, lswg feb 2015-1, p. 69. abstract introduction materials and methods results discussion acknowledgements references about the author(s) isiah nthenga faculty of agrisciences, stellenbosch university, stellenbosch, south africa department of biology, nematology research group, faculty of biological sciences, gent university, gent, belgium rinus knoetze department of conservation ecology and entomology, faculty of agrisciences, stellenbosch university, stellenbosch, south africa plant protection division, agricultural research council (arc), stellenbosch, south africa antoinette p. malan department of conservation ecology and entomology, faculty of agrisciences, stellenbosch university, stellenbosch, south africa citation nthenga, i., knoetze, r. & malan, a.p., 2021, ‘distribution and diversity of entomopathogenic nematodes (steinernematidae and heterorhabditidae) in a south african nature reserve’, koedoe 63(1), a1661. https://doi.org/10.4102/koedoe.v63i1.1661 original research distribution and diversity of entomopathogenic nematodes (steinernematidae and heterorhabditidae) in a south african nature reserve isiah nthenga, rinus knoetze, antoinette p. malan received: 30 oct. 2020; accepted: 18 sept. 2021; published: 22 nov. 2021 copyright: © 2021. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract entomopathogenic nematodes (epns) are microscopic roundworms that are found in soil worldwide. they deliver an important ecosystem service through preventing natural flares in insect reproduction by means of utilising the soil stages of insects as a food source and by acting as natural biocontrol agents. a survey of epns was conducted in the js marais nature reserve, stellenbosch, in the western cape province of south africa. soil samples were baited with the larvae of three susceptible hosts, codling moth (cydia pomonella), wax moth (galleria mellonella) and mealworm (tenebrio molitor) to determine the presence of epn. of the 76 soil samples collected across the reserve, 39 were found to be positive for the presence of epn (51.32%). among the positive samples, 87% contained steinernema isolates, 8% contained heterorhabditis and 5% contained the oscheius sp. morphological and molecular studies were performed to characterise the isolates to species level. the steinernema species were identified as steinernema khoisanae in 34 samples, and as steinernema nguyeni in five samples. the only species of heterorhabditis found was h. safricana, which was identified from three samples. an unknown oscheius sp. was found in two samples. the reserve’s population of s. khoisanae showed interesting inter-individual variation (93%) early in the internal transcribe spacer (its) region, leading to short single-usable sequences, which, in most cases, included only the its1 or its2 region. however, using the d2d3 confirmed their identity as s. khoisanae, with such occurring in all areas and soil types of the reserve. conservation implications: the undisturbed alluvial fynbos and renosterveld of the js marais nature reserve showed high epn abundance and diversity in stark contrast to the agro-ecosystems present in the cape floristic region. this finding, on a micro level, should be conserved for future bioprospecting in the fynbos for epns with potential as biocontrol agents. keywords: distribution; fynbos; js marais nature reserve; natural habitat; natural veld. introduction entomopathogenic nematodes (epns) in the families steinernematidae and heterorhabditidae are of great interest because of their potential as commercial biological control agents against insects. entomopathogenic nematodes are lethal, obligate parasites of soil-dwelling stages of insect pests (burnell & stock 2000). they occur naturally in all soils worldwide, with their control of the soil stages of insects acting as an ecosystem service for keeping insect populations at bay. entomopathogenic nematodes have been successfully used worldwide as biological control agents to suppress over 200 economically important insect pests (grewal, ehlers & shapiro-ilan 2006; kaya & gaugler 1993; shapiro-ilan, gouge & koppenhofer 2002). entomopathogenic nematode surveys during the last decades yielded an increasing number of new species. entomopathogenic nematodes are found globally, except for antarctica, where they have yet to be found (griffin, downes & block 1990; hominick 2002). the study of natural epn occurrence and distribution, as characterised by different ecological habitats in various geographical areas, provides baseline information for incorporating the pathogens in biological control programmes, and for understanding the complexities of their biodiversity. the factors such as geographical location, climatic conditions, soil properties and habitat type may determine the occurrence and distribution of the epns. to be effective as biological control agents, epns need to be adapted to the local environmental conditions of the site of application (bedding 1990). however, with inundative application, with an immediate effect on the target organism, temperature and humidity are the most important environmental factors to take into consideration. in south africa, five major surveys have been conducted to isolate epns for use as biological control agents (hatting & malan 2017; malan & hatting 2015). a survey was conducted between 2004 and 2005, during which 498 soil samples were collected, with only 36 representing 7% of the total number of samples testing positive for epns (malan, nguyen & addison 2006). this survey, aimed at obtaining epns for use as biological control agents against codling moth (cydia pomonella linnaeus), were restricted to the southern part of the western cape province. another survey, aimed at determining the distribution and diversity of epns, was conducted between 2003 and 2005, during which 1508 soil samples were collected. this survey, in which only 79 (5%) of the total samples tested positive for epns (hatting, stock & hazir 2009), was conducted in the western cape, free state, gauteng, mpumalanga and kwazulu-natal provinces. the third survey, aimed at determining the potential of epns for the control of the soil stages of a false codling moth, thaumatotibia leucotreta (meyrick), consisted of 202 soil samples, of which 35 (17%) were found to test positive for the presence of epns (malan, knoetze & moore 2011). three provinces were targeted in the survey, namely the western cape, the eastern cape and mpumalanga provinces. recent surveys were conducted by steyn et al. (2017b) from avocado, litchi and macadamia orchards in the subtropical regions of mpumalanga, limpopo and kwazulu-natal provinces, while abate et al. (2018) surveyed eucalyptus spp., pinus spp. and acacia mearnsii de wild plantations in kwazulu-natal and mpumalanga. both surveys reported new steinernema species, steinernema fabii abate, malan, tiedt, wingfield, slippers & hurley (abate et al. 2016) and steinernema litchi steyn, knoetze, tiedt and malan (steyn et al. 2017a), as well as new reports for south africa of heterorhabditis taysearae shamseldean, el-sooud, abd-elgawad & saleh (steyn et al. 2017b) and heterorhabditis baujardi phan, subbotin, nguyen and moens (abate et al. 2018; steyn et al. 2017b). the above surveys have contributed to the existing knowledge of the geographical distribution and diversity of epns. the natural vegetation type in the western cape consists of fynbos, with such vegetation consisting primarily of natural shrubland, which occurs in a small belt of the western cape, which has a mediterranean climate, characterised by winter rainfall (esler, pierce & de villiers 2014). the fynbos biogeography is known for its exceptional degree of biodiversity and endemism. the diversity of fynbos plants is extremely high, with over 9000 species of plants occurring in the area, of which about 6200 are endemic. the extremely high level of diversity is comparable to that of tropical rainforests or large islands, and it is unique to the relatively dry african continent (esler et al. 2014). it is hypothesised that the diversity of epns in the fynbos will be as diverse, as different vegetation types it is associated with. the objective of the current study was to determine the distribution and composition of indigenous epns in the natural vegetation of the western cape province, in terms of the fynbos found in the js marais nature reserve in the town of stellenbosch. the information obtained should add to the knowledge of the occurrence and distribution of epns in mostly undisturbed soils in south africa. materials and methods study area the js marais nature reserve, which is composed of natural veld and managed areas, covers an area of 23 ha, situated on the eastern side of the stellenbosch university campus, in the town of stellenbosch, western cape. the area was proclaimed as a nature reserve in 2019. the natural veld is found in the a blocks, with the managed areas in blocks b, c and d (figure 1). in the western cape, very little natural vegetation remains between the mountains and the sea, making the conservation of the two veld types very important. the reserve protects the original habitats of the alluvial terraces on which the town of stellenbosch was established. the reserve contains the only remaining of boland granite fynbos and swartland renosterveld (mucina & rutherford 2006). alien vegetation found in the reserve includes pinus radiata d., pinus pinea l. and various eucalyptus species. figure 1: map showing sites where samples were taken, indicating the positive and negative sampling sites and the different species of entomopathogenic nematodes isolated. the climatic data about the reserve, incorporating a long-term summary over a period of 33 years, were obtained from the arc infruitec-nietvoorbij weather station at nietvoorbij farm, being the closest weather station to the nature reserve. the reserve is in a winter rainfall region, with an average rainfall of between 700 mm and 1500 mm. the maximum monthly temperature for a 33-year period was during november 1972, with it being 41.2 °c and with the lowest minimum monthly temperature being 1.1 °c. the very low temperatures occurring in the reserve are frequently accompanied by snowfall on the mountain peaks surrounding stellenbosch. the general surface layer of the js marais nature reserve consists of loose gravel, with a highly variable sand matrix. the soil of the reserve is largely eerste river alluvial gravel, which is derived from table mountain sandstone and some granite. collection of soil samples the reserve is demarcated into 22 block sections (figure 1), with each block being chosen as a sampling site. a total of 76 soil samples were collected in the reserve during august and september of 2012. at each sampling site, in an area of approximately 50 m2, five subsamples, consisting of approximately 200 g of soil, were taken at least 4 m apart, in a cross-sampling design and at a depth of 10–15 cm, using a small hand trowel. the subsamples from each site were combined into a single plastic bag and mixed thoroughly. between samples, the shovel was thoroughly cleaned with water and dried with paper towels, to prevent contamination between the next sampling site. approximately 500 g of the soil from each sample was placed in polyethylene bags to prevent loss of moisture, whereupon the bags were labelled and taken to the laboratory for processing. the following information was recorded: the site location, the date, the global positioning system (gps) reading and the vegetation and soil type. source of insect hosts the greater wax moth, galleria mellonella (linnaeus) (lepidoptera: phylaridae), was cultured in a growth medium and maintained at 25 °c. it is the most frequently used insect for the propagation of epns, because it is easy to maintain and very susceptible to epns. mealworm, tenebrio molitor (linnaeus) (coleoptera: tenebrionidae) larvae were cultured, using bran, with carrots for moisture (van zyl & malan 2015). the culture was maintained at room temperature. codling moth, cydia pomonella (linnaeus) larvae, which were kept at 4 °c, were obtained from the entomon facility at welgevallen experimental farm. isolation of nematodes entomopathogenic nematodes were recovered from the soil samples, using an insect-baiting method described by bedding and akhurst (1975). in the laboratory, soil samples from each site were placed into 500 ml plastic containers. five last instars each of wax moth larvae, mealworm larvae and codling moth larvae were added to each soil sample and stored at room temperature of 20 ± 22 °c for 21 days. the soil traps were checked every 7 days and dead larvae removed from the pots and replaced with healthy ones. the dead insects from each sample were rinsed in water and placed in a 9 cm petri dish lined with moist filter paper for 5–6 days. the larvae showing signs of epn attack were transferred to a modified white trap (kaya & stock 1997; woodring & kaya 1988) to collect the infective juveniles (ijs). to verify the pathogenicity of collected nematodes and to establish new cultures, the emerging nematodes were collected for each sample and used to infect fresh g. mellonella larvae. parasitised g. mellonella were placed on a white trap to collect the ijs, which were then stored in culture flasks at 14°c in water. molecular characterisation the molecular approach in identifying the nematodes was according to nguyen and smart (1996) and nguyen, maruniak and adams (2001). dna was extracted from a single nematode female, using a modification of a method reported by nguyen (2007). the its-rdna regions were amplified using the its primers 18s: 5’-ttgattacgtccctgcccttt-3’ (forward) and 26s: 5’-tttcactcgccgttactaagg-3’ (reverse) as reported by vrain et al. (1992). the forward primer d2f: 5’-ccttagtaacggcgagtgaaa-3’ (nguyen, malan & gozel 2006) and the reverse primer 536: 5’-cagctatcctgaccaaac(stock, pryor & kaya 1999) were used to amplify the d2–d3 expansion segments of 28s rrna. polymerase chain reaction (pcr) products were cleaned up and sequenced by the central analytical facilities (caf) of the stellenbosch university. the sequence was assembled and edited using the clc dna workbench (http://www.clcbio.com). the sequences generated of the its region of all the positive samples were compared with those of the species made available on genbank (ncbi). morphological and morphometric characterisation to determine the genus of the isolated epns, the diagnostic criteria suggested by stock and kaya (1996) were used. microscopic examination of live ijs, males and females was performed to support the molecular identity of the different isolates, using key diagnostic features (nguyen & smart 1996). measurements were made by means of a leica dm2000 compound microscope (leica microsystems, wetzlar, germany), fitted with a digital camera and with software leica application suite v3.5.0. results more than half (39) of the samples collected the nature reserve contained epns. two endemic species of steinernema (29 isolates) were isolated and identified with molecular techniques viz. steinernema khoisanae (24 isolates) and steinernema nguyeni malan, knoetze and tiedt (five isolates). heterorhabditis safricana malan, nguyen, de waal and tiedt (malan et al. 2008), the only heterorhabditis found, was isolated from two samples. an unknown oscheius sp. was also isolated from two soil samples (table 1). table 1: identification of entomopathogenic nematodes (steinernema, heterorhabditis and oscheius) isolated from soil collected at the js marais nature reserve. sequences of the its region of s. khoisanae generated with the two primer pairs, tw81 (f) and ab28 (r) and 18s (f) and 26s (r), did not produce good-quality sequences. only partial sequences of either the its1 with the forward primer, or its2 with the reverse primer, could be submitted to genbank. only in two cases was a full its (its1-2.8s-its2) region for s. khoisanae was obtained. however, sequencing of the d2–d3 regions of the 28s gene large subunit produced good-quality sequencing > 880 base pairs and confirmed their identity (100%) with the isolates of populations of s. khoisanae (table 1) reported from other areas in the western cape province. all species isolated were trapped using mealworm, except at two samples, where s. khoisanae was trapped with wax moth. in 11 samples, both the mealworm and the codling moth trapped both s. khoisanae and s. nguyeni. entomopathogenic nematodes were trapped in 14 samples using wax moth. in two samples, epns were trapped with all three trapping hosts, whereas, in 10 samples, both wax moth and mealworm trapped epns (figure 1). the oscheius sp. was trapped using mealworm (table 1). the soil analysis showed that s. khoisanae were found in all the soil types throughout the nature reserve (table 2). table 2: representative soil samples analysed using a three-fraction soil analysis to indicate percentages differences in the different sampling regions. of all the isolates, the body length of 10 ijs was measured to confirm the molecular identification. the body length of s. khoisanae specimens from the js marais nature reserve was found to be slightly shorter than those of the type specimens 1010 (916–1127) µm versus 1062 (904–1159) µm. in the case of s. nguyeni, the body length of the ij correlated with that of the type specimen. for h. safricana, the ij body length of the type specimen 697 (656–725) µm was found to be slightly longer, versus 600 (550–676) µm of the specimens found in this study. the sequences derived from the oscheius sp. could not be matched to any known sequences in genbank (table 3); thus, possibly, it is a new, undescribed species of the genus. table 3: measurements of the body length of the infective juveniles. discussion the occurrence and diversity of the epns found in the js marais nature reserve were high (51%), with four species being identified, namely s. khoisanae, s. nguyeni, h. safricana and an oscheius sp. the steinernema spp. were more abundant than the others, which is found in 94% of the positive samples. steinernema khoisanae was the most common species, which is found in 86% of the positive samples, with it being distributed all over the nature reserve. of the positive soil samples, three contained s. nguyeni, three h. safricana and two with an unknown oscheius sp. all four species are endemic to south africa. the trend of more steinernematids being recovered than heterorhabditis as stated by hominick (2002) was confirmed in the current survey. in previous surveys undertaken to test for the presence of epns, the recovery rate was, in general, very low. a non-targeted survey by malan et al. (2006) showed a 7% recovery rate, while, in a targeted citrus orchard survey, 17% recovery was obtained for a total of 119 sampling sites. hatting et al. (2009) undertook a comprehensive survey (of 1500 samples), with a very low recovery rate of 5%. most of these samples were taken from agricultural soil, which could be the reason for the low recovery rate obtained. in the current survey, less than four samples were collected per hectare, which is not regarded as intensive, as in orchards, 1 ha is usually divided into four quadrants, from which five subsamples are combined to give one sample, with a total of four samples per hectare (steyn, malan & addison 2020). from the information obtained, it can be extrapolated that the occurrence of epns in the js marais nature reserve was exceptionally high. three susceptible hosts were chosen for the isolation of epns because some species or strains could prove to be host-specific (malan et al. 2011). during the present study, the most effective trapping host was t. molitor, from which all positive samples were trapped, except for two species of s. khoisanae, which were isolated with wax moth larvae. unfortunately, the natural insect host of none of the epn species isolated are known (table 1). some epn species were found to be highly specific, like steinernema scapterisci nguyen and smart, which appears to reproduce only in adult mole crickets (scapteriscus spp.) (nguyen & smart 1990). this indicates that many more epns might have been present in the soil but were not trapped by the three hosts used. steinernema bertusi katumanyane, malan, tiedt, hurley, a newly described species for south africa, was successfully isolated using wax moth and mealworm on two separate occasions and at two separate locations, from the kwazulu-natal and mpumalanga provinces (katumanyane et al. 2020). the high abundance of s. khoisanae in the western cape province could suggest the presence of a wide range of suitable endemic hosts and favourable conditions for the species, which might contribute to its high observed frequency (hatting et al. 2009; malan et al. 2006, 2011). from the soil analysis (table 2), s. khoisanae was noted to be present in all the soil types (lmsa, salm, lm) of the nature reserve. intra-individual variability of the its region of most of the s. khoisanae individuals influenced the ability of standard primer pairs to produce high-quality sequences for this region that could successfully be aligned to identify the species. however, in this study, only two sequences were obtained by means of aligning the forward and the reverse sequences, both using the 28s and 26s primer pairs (table 1). using the d2d3 region of the 28s gene region, high-quality sequences were obtained with both the forward and the reverse primers, thus confirming their identity as s. khoisanae. the its region, which is among the most important molecular markers used in identification, taxonomy and phylogeny of epns, was used with for the species identified from south africa, as well as from other parts of the world (malan & hatting 2015). půža et al. (2015) found that, in steinernema feltiae (filipjev) wouts, mráček, gerdin & bedding, intra-individual variability was found in 65% of specimens, and in almost 90% of the glaseri group, sampled from different localities. however, in the js marais nature reserve, 93% of the individuals in the population showed intra-individual variation early in the process of sequencing the its region, with the generation of short sequences at the 3’ and 5’ ends, which could not be aligned. půža et al. (2015) also suggested the importance of using the d2d3 region to confirm the new species status of an epn isolate. in this case, the d2d3 was used to confirm the species identification of most of the populations found in the same area, which is very rare occurrence. de waal et al. (2010) tested the virulence of six epn species against diapausing codling moth, an insect of agricultural importance. the results showed the high susceptibility of codling moth for s. khoisanae, causing 97% and 88% infection, respectively. during a field trail against the false codling moth in citrus, s. khoisanae did not perform as well as did the other species tested. as the ij of s. khoisanae is relatively large (≈1 cm), it generally cannot be used as a biocontrol against such micro insects as bradysia impatiens and frankliniella occidentalis (pergande) (thysanoptera: thripidae) (dlamini, allsopp & malan 2019; katumanyane, ferreira & malan 2018). this specific population of s. khoisanae from the js marais nature reserve with its different its region should be tested against pest insects to compare it to other s. khoisanae populations with a normal its profile. steinernema nguyeni was first isolated from soil collected from beneath olea europaea subsp. africana l. (miller), an indigenous tree, near clanwilliam in the western cape. the nematode was trapped using both g. mellonella and t. molitor. similarly, it was reported from a survey conducted by de waal, malan and addison (2011), as being isolated from undisturbed soil in the knysna area of the western cape, but not in any other surveys. heterorhabditis safricana (sf281), which was described from a peach orchard in the western cape, has been isolated by trapping, using galleria as host. it seems to be a rare species in south africa, as previous surveys have detected its presence only in a peach orchard (malan et al. 2008). just two isolates were recovered during the current survey. some controversy surrounds the status of the oscheius (= heterorhabditidoides) in its role as an entomopathogen, but the species found in the js marais nature reserve could most probably be a new species. in terms of the recovery rate and frequencies reported by previous surveys, caution must be taken because of the differences in the sampling design, the time of sampling and the insect bait used. the high number of positive samples observed in the current survey may be attributed to the use of three different trapping hosts, and the increased number of sampling sites for the area to compensate for the patchy distribution of epns, as reported by stock et al. (1999). the use of different insect hosts has shown that some isolates prefer certain hosts, which may lead to an underestimation of the natural occurrence of epns. however, the current study confirms that the use of mealworms is very effective for the isolation of species. the present study provides an understanding of the distribution and diversity of the epns found in the js marais nature reserve, which is a natural ecosystem surrounded by urban development, which has remained undisturbed for over a century, in the heart stellenbosch, in the western cape province. the unique intraspecific variation found in the its region of the s. khoisanae populations of the js marais nature reserve should be further investigated, as well as its implications for the use of the its region as a molecular marker. acknowledgements the author wished to thank the flemish inter-university council – university development cooperation (vlir-uos) and the technology and human resources for industry programme (thrip) for funding of the current project. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions i.n. contributed towards the investigation and writing the original draft. r.k. contributed to the formal analysis, writing, reviewing and editing of the manuscript. a.p.m. was responsible for conceptualisation, methodology, fomal analysis, writing, reviewing and editing of the manuscript as well as supervision of the study and funding acquisition. ethical considerations this article followed all ethical standards for research without direct contact with human or animal subjects. funding information the authors would like to thank the flemish inter-university council – university development cooperation (vlir-uos) and the technology and human resources for industry programme (thrip) for funding of the current project. data availability the authors confirm that the data supporting the findings of this study are available within the article. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references abate, b.a., malan, a.p., tiedt, l.r., wingfield, m.j., slippers, b. & hurley, b.p., 2016, ‘steinernema fabii n. sp. (rhabditidae: steinernematidae), a new species of entomopathogenic nematodes from south africa’, nematology 18, 235–255. https://doi.org/10.1163/15685411-00002956 abate, b.a., slippers, b., wingfield, m.j., malan, a.p. & hurley, b.p., 2018, ‘diversity of entomopathogenic nematodes and their symbiotic bacteria in south african plantations and indigenous forests’, nematology 20, 335–317. https://doi.org/10.1163/15685411-00003144 bedding, r.a., 1990, ‘logistics and strategies for introducing entomopathogenic nematode technology in developing countries’, in r. gaugler & h.k. kaya (eds.), entomopathogenic nematodes in biological control, pp. 233–248, crc press, boca raton, fl. bedding, r.a. & akhurst, r.j., 1975, ‘a simple technique for the detection of insect parasitic rhabditid nematodes in soil’, nematologica 21, 109–110. https://doi.org/10.1163/187529275x00419 burnell, a.m. & stock s.p., 2000, ‘heterorhabditis, steinernema and their bacterial symbionts – lethal pathogens of insects’, nematology 2(1), 31–42. https://doi.org/10.1163/156854100508872 de waal, j.y., malan a.p., levings, j. & addison, m.f., 2010, ‘key elements in the successful control of diapausing codling moth, cydia pomonella (l.) (lepidoptera: tortricidae) in fruit bins with a south african isolate of heterorhabditis zealandica (rhabditida: heterorhabditidae)’, biocontrol science and technology 20(5), 489–502. https://doi.org/10.1080/09583151003599708 de waal, j.y., malan, a.p. & addison, m.f., 2011, ‘efficacy of entomopathogenic nematodes (rhabditida: heterorhabditidae and steinernematidae) against codling moth, cydia pomonella (lepidoptera: tortricidae) in temperate regions’, biocontrol science and technology 20(5), 489–502. https://doi.org/10.1080/09583157.2011.607922 dlamini, t.m., allsopp, e. & malan, a.p., 2019, ‘efficacy of entomopathogenic nematodes against western flower thrips, frankliniella occidentalis (thysanoptera: thripidae), under laboratory conditions’, african entomology 27(2), 322–335. https://doi.org/10.4001/003.027.0322 esler, k.j., pierce, s.m. & de villiers, c., 2014, fynbos – ecology and management, briza publications, pretoria. grewal, p.s., ehlers, r. & shapiro-ilan, d.i., 2006, nematodes as biocontrol agents, cabi, wallingford. griffin, c.t., downes, m.j. & block, w., 1990, ‘tests of antarctic soils for insect parasitic nematodes’, antarctic science 2, 221–222. https://doi.org/10.1017/s095410209000030x hatting, j., stock, p.s. & hazir, s., 2009, ‘diversity and distribution of entomopathogenic nematodes (steinernematidae, heterorhabditidae) in south africa’, journal of invertebrate pathology 102, 120–128. https://doi.org/10.1016/j.jip.2009.07.003 hatting, j.l. & malan, a.p., 2017, ‘status of entomopathogenic nematodes in integrated pest management strategies in south africa’, in m.m.m. abd-elgawad, t.h. askary & j. coupland (eds.), biocontrol agents: entomopathogenic and slug parasitic nematodes, pp. 409–428, cabi, wallington. hominick, w.m., 2002, ‘biogeography’, in r. gaugler (ed.), entomopathogenic nematology, pp. 115–143, cabi publishing, wallingford. katumanyane, a., ferreira, t. & malan, a.p., 2018, ‘bradysia species (diptera: sciaridae) as pests of covered crops, with special reference to biological control using entomopathogenic nematodes’, african entomology 26(1), 1–13. https://doi.org/10.4001/003.026.0001 katumanyane, a., malan, a.p., tiedt, l.r & hurley, b.p., 2020, ‘steinernema bertusi n. sp. (rhabditida: steinernematidae), a new entomopathogenic nematode from south africa’. nematology 22, 343–360. https://doi.org/10.1163/15685411-00003309 kaya, h.k. & gaugler, r., 1993, ‘entomopathogenic nematodes’, annual review of entomology 38, 181–206. https://doi.org/10.1146/annurev.en.38.010193.001145 kaya, h.k. & stock, s.p., 1997, ‘techniques in insect nematology’, in l. lacey (ed.), manual of techniques in insect pathology, pp. 281–324, academic press, london. malan, a.p. & hatting, j., 2015, ‘entomopathogenic nematode exploitation: case studies in laboratory and field applications from south africa’, in r. campos-herrera (ed.), sustainability in plant and crop protection: ecology and applied technologies for sustainable plant and crop protection, pp 475–506. cham, springer. malan, a.p., knoetze, r. & moore, s.d., 2011, ‘isolation and identification of entomopathogenic nematodes from citrus orchards and their biocontrol potential against false codling moth’, journal of invertebrate pathology 108, 115–125. https://doi.org/10.1016/j.jip.2011.07.006 malan, a.p., nguyen, k.b. & addison, m.f., 2006, ‘entomopathogenic nematodes (steinernematidae and heterorhabditidae) from the southwestern parts of south africa’, african plant protection 12, 65–69. malan, a.p., nguyen, k.b., de waal j.y. & tiedt, l., 2008, ‘heterorhabditis safricana n. sp. (rhabditida: heterorhabditidae), a new entomopathogenic nematode from south africa’, nematology 10, 381–396. https://doi.org/10.1163/156854108783900258 malan, a.p., tiedt, t.r. & knoetze, r., 2016, ‘a new entomopathogenic nematode, steinernema nguyeni from south africa’, nematology 18, 571–590. https://doi.org/10.1163/15685411-00002978 mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, strelitzia 19, south african national biodiversity institute, pretoria. nguyen, k.b., 2007, ‘methodology, morphology and identification’, in k.b. nguyen & d.j. hunt (eds). entomopathogenic nematodes: systematics, phylogeny and bacterial symbionts, pp. 59–119, leiden, brill. nguyen, k.b., malan, a.p. & gozel, u., 2006, ‘steinernema khoisanae sp. n. (rhabditida: steinernematidae), a new entomopathogenic nematode from south africa’, nematology 8, 157–175. https://doi.org/10.1163/156854106777998728 nguyen, k.b., maruniak j. & adams, b.j., 2001, ‘diagnostic and phylogenetic utility of the rdna internal transcribed spacer sequences of steinernema’, journal of nematology 33(2–3), 73–82. nguyen, k.b. &. smart, g.c., jr., 1990, ‘steinernema scapterisci n. sp. (rhabditida: steinernematidae)’, journal of nematology 22, 187–199. nguyen, k.b. & smart, g.c., jr., 1996, ‘identification of entomopathogenic nematodes in the steinernematidae and heterorhabditidae (nemata: rhabditida)’, journal of nematology 28(3), 286–300. půža, v., chundelová, d., nemut’, j., žurovcová, m. & mráček, z., 2015, ‘intra-individual variability of its regions in entomopathogenic nematodes (steinernematidae: nematoda): implications for their taxonomy’, biocontrol 60, 547–554. https://doi.org/10.1007/s10526-015-9664-5 shapiro-ilan, d.i., gouge, d.h. & koppenhofer, a.m., 2002, ‘factors affecting commercial success: case studies in cotton, turf and citrus’, in r. gaugler (ed.), entomopathogenic nematology, pp. 333–356, cabi, wallingford. steyn, v.m., malan a.p. & addison, a., 2020, ‘efficacy of entomopathogens against thaumatotibia leucotreta (lepidoptera: tortricidae) under laboratory conditions’, entomologia experimentalis et applicata 168, 499–461. https://doi.org/10.1111/eea.13044 steyn, w.p., knoetze, r., tiedt t.r. & malan, a.p., 2017a, ‘steinernema litchi n. sp. (rhabditida: steinernematidae), a new entomopathogenic nematode from south africa’, nematology 19, 1157–1177. https://doi.org/10.1163/15685411-00003116 steyn, w.p., malan, a.p., daneel, m.s. & slabbert, r.m. 2017b, ‘entomopathogenic nematodes from north-eastern south africa and their virulence against the false codling moth, thaumatotibia leucotreta (lepidoptera: tortricidae)’, biocontrol science and technology 27(11), 1265–1278. https://doi.org/10.1080/09583157.2017.1391174 stock, s.p. & kaya, h.k., 1996, ‘a multivariate analysis of morphometric characters of heterorhabditis species (nematoda: heterorhabditidae) and the role of morphometrics in the taxonomy of species of the genus’, journal of parasitology 82, 806–813. stock, s.p., pryor b.m. & kaya, h.k., 1999, ‘distribution of entomopathogenic nematodes (steinernematidae and heterorhabditidae) in natural habitats in california’, biodiversity & conservation 8(4), 535–549. van zyl, c. & malan, a.p., 2015, ‘optimisation of inoculation techniques for in vivo mass culture of entomopathogenic nematodes through nematode and insect host manipulation’, african entomology 24, 405–416. https://doi.org/10.4001/003.022.0221 vrain, t.c., wakarchuck, d.a., lèvesque a.c. & hamilton r.i., 1992, ‘intraspecific rdna restriction fragment length polymorphism in the xiphinema americanum group’, fundamental and applied nematology 15, 563–573. woodring, j.l. & kaya, h.k., 1988, ‘steinernematid and heterorhabditid nematodes: a handbook of techniques’, southern cooperative series bulletin 331, arkansas agricultural experimental station, fayetteville, arkansas, 30 pp. introduction background and context of dwesa-cwebe nature reserve missed opportunities in mitigating people-conservation tension in dwesa-cwebe nature reserve discussion and conclusion acknowledgements references about the author(s) deon de villiers eastern cape department of economic development, environmental affairs and tourism, bisho, south africa thembela kepe department of human geography, university of toronto, toronto, canada department of geography, faculty of science, rhodes university, makhanda, south africa citation de villiers, d. & kepe, t., 2023, ‘missed opportunities in resolving people-conservation tensions in dwesa-cwebe, south africa’, koedoe 65(1), a1750. https://doi.org/10.4102/koedoe.v65i1.1750 short communication missed opportunities in resolving people-conservation tensions in dwesa-cwebe, south africa deon de villiers, thembela kepe received: 25 dec. 2022; accepted: 23 july 2023; published: 22 aug. 2023 copyright: © 2023. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. introduction the importance of biodiversity conservation is arguably undisputed globally, as people generally realise that their survival is intertwined with biodiversity, and that threats to it could diminish their quality of life (sanderson & redford 2003). but tensions between people and conservation management are a worldwide global phenomenon, particularly in the global south, where past injustices saw conservation needs trumping land and natural resource justice for local people (hutton, adams & murombedzi 2005) and where widespread poverty is seen by many as putting pressure on biodiversity (du toit, walker & campbell 2004). given this state of affairs, numerous attempts to mitigate people-conservation tension have been made by various constituencies, including international bodies concerned with environmental health, state agencies tasked with management of protected areas, academics, businesses, organisations and individuals (dressler et al. 2010). kepe (2018) argues that these stakeholders commonly form alliances (ideological or actual) to advance or oppose the mitigation of the people-conservation tension. strategies in mitigating people-conservation conflict have included, but are not limited to, erection and maintenance of fences and/or issuing of fines to trespassers. in this approach, enclosures and punitive measures are implemented as deterrents (hutton et al. 2005). alternative approaches include ‘inclusive conservation’, where local people are afforded a greater stake in managing local natural resources (e.g. community based natural resource management), (dressler et al. 2010), ‘conservation against poverty’, where alternative livelihood means are promoted to meet local livelihood needs, while aiming to reduce pressure on biodiversity (salafsky 2011), and neoliberal conservation, where markets and the private sector are seen to play a role in making conservation commercially viable (igoe & brockington 2007). although individual site-level successes have been documented, in general, these strategies have not succeeded as a universal blueprint for mitigating people-conservation conflict. this article argues that there is a need to acknowledge that the four strategies mentioned here are not a panacea for all contexts, and that there is a need to explore every opportunity, at multiple scales, in a manner that is context-specific. specifically, this article argues that in each case of people-conservation conflict, as much as there is a need to look forward, in terms of plans and resolutions, it is important to look back at important points in history, where opportunities existed for mitigating the tension. an assessment of the history of a site would allow planners to develop strategies that avoid repeating past mistakes, while at the same time opening opportunities to explore how past mistakes are addressed in future plans. using dwesa-cwebe nature reserve located on the wild coast of the eastern cape, a 300 km stretch of coastline, as a case study, this article explores missed opportunities that explain the lingering people-conservation tensions in the area. it is argued that acknowledging these failures, broadly, is a viable pathway to mitigating people-conservation tensions in the area. the information discussed in this article is based on a number of sources. firstly, there was extensive review of literature, including published and unpublished material. this material included books and journal articles, policy and legislation documents, and archival material. secondly, the authors’ long-term experience working on the intersection of natural resources and livelihoods, as well as conservation on the wild coast, including insights from one of the authors’ doctoral study (de villiers 2021), allowed for insights to be drawn from detailed direct observations in the area for a period of over 30 years. background and context of dwesa-cwebe nature reserve although small in size, dwesa-cwebe nature reserve is known in south africa for biodiversity conservation, as well as issues around violent conflict and land justice (timmermans 2004). the 5700 hectares dwesa-cwebe nature reserve is an amalgamation of two smaller terrestrial reserves (dwesa nature reserve and cwebe nature reserve), which were proclaimed in 1975. thesse are adjacent to the 18 000 ha dwesa-cwebe marine protected area (de villiers 2021). dwesa was officially declared as a state forest in 1891 (henkel 1895), arguably making it the first protected area in south africa (feely 2013). by the early 1970s, with the looming independence of the ‘transkei’, an apartheid-created homeland for black south africans, the south african government identified dwesa as the ideal locality for the declaration of a protected area (timmermans 2004). the biological importance of dwesa and the adjacent cwebe indigenous forests had long been recognised and were considered as being among the most valuable and ecologically significant forests in south africa (sim 1906). they are some of the most species-rich non-tropical forests globally (moll 1974), with high levels of endemism (cooper & swart 1992). moll’s (1974) preliminary report on the dwesa forest reserve not only highlighted the scientific value of the forests but also of the grasslands and adjacent marine resources. the establishment of these protected areas resulted in evictions of african people who resided inside these forests, as late as the 1890s (henkel 1895). the act of evicting people from dwesa and cwebe state forests, or preventing them from reoccupying or utilising the resources of the land was sustained by the various successive governments that assumed management authority over the protected areas (ntshona et al. 2010). more recently, dwesa-cwebe nature reserve gained national prominence during the early 1990s as a result of mass protests and reserve invasion by people from the surrounding seven villages, demanding access to grazing, forest resources and marine resources (ntshona et al. 2010). the nationally televised account of local village protesters forcefully removing natural resources from the reserve was a stark reminder about the tensions that exist between people and conservation, something that led to numerous meetings between the authorities and local communities, including their representatives (e.g. non-governmental organisation), to mitigate the tension. these protests coincided with a changed political climate that signalled the end of official apartheid, and the ushering in of democratic governance in south africa. in 1996, utilising the legislated land claims process, members of the seven villages lodged a land claim for the return of their land (fay 2007). even though the land claim was subsequently resolved, declaring the local communities rightful owners of the reserve, use of natural resources within dwesa-cwebe continues to be restricted because of a clause in the land restitution settlement agreement that the reserve will continue to operate as such, and be managed by a state conservation authority, while local people would benefit through ecotourism (ntshona et al. 2010). the dwesa-cwebe land claim and settlement was seen as probably the most high-profile in the country, following the makuleke claim and settlement for sections of kruger national park (de villiers 2021). tensions between conservation authorities and neighbouring communities of dwesa-cwebe nature reserve continue to this day, and occasionally these turn violent. the causes of the tensions include a range of issues, from poor implementation of the land claim settlement agreement, to clashes over access to and use of the locally available natural resources (de villiers 2021). given the aforementioned context, the following section explores how the people-conservation tensions could have been and could still be mitigated in the case of dwesa-cwebe nature reserve. missed opportunities in mitigating people-conservation tension in dwesa-cwebe nature reserve linking dwesa-cwebe nature reserve to local heritage history history of place has been central in the origins of people-conservation tensions, globally (agrawal & redford 2009). arguably, the history of settlement, natural resource use, and subsequent dispossession of local and indigenous people of their land to make way for biodiversity conservation, has been the central cause of tension. however, even with that history of injustice, there are potential opportunities to use other histories of places where the conservation area is, to give a sense of pride and belonging among the local people, and thus reduce potential tensions (de villiers 2021). in dwesa-cwebe nature reserve there is currently one historical linkage that could be made between local people and this protected area. the reserve is situated in the heartland of amaxhosa kingdom. more specifically, dwesa forest has direct linkages with one of the famous amaxhosa kings, sarhili, the son of king hintsa who was assassinated by the british in 1835 (peires 1989). part of king sarhili’s fame among amaxhosa comes from his several conflicts with the british armies, as part of frontier wars. slade (2010) also argues that sarhili was the last independent king of amaxhosa before they were defeated by the british, and thus is commonly seen as a symbol of amaxhosa life that was disrupted by europeans’ arrival. this is why his gravesite at tsholorha is now a tourist site in the area (mbashe local municipality 2022). there are two important links between sarhili and the dwesa-cwebe nature reserve. firstly, in his battles with the british, and on the verge of defeat, king sarhili and his bodyguard hid in the cwebe forests, just east of mbhashe river (peires 1989). secondly, long before the colonial government recognised the need to conserve dwesa, king sarhili is known to have had it set aside as a protected forest presumably for his royal hunting ground (henkel 1889), but cawe (1992) argues that he may have protected the forests because he admired their grandeur. irrespective of the reasons for protecting it, there is clear evidence that king sarhili protected dwesa forest long before the colonial forestry department (henkel 1895). this article argues that this history of protection of dwesa by sarhili, and his use of the cwebe forest as a refuge when his life was in danger, could be promoted to allow local people to be reminded of their history in association with the protected area. just like sarhili’s gravesite has become a tourist attraction, dwesa-cwebe nature reserve could elevate its historical relevance to develop a sense of belonging and stewardship among local people, as well as have a commercial value as a tourist attraction. planners’ recommendations on inclusive conservation during establishment of the reserve studies on people-conservation tensions regarding dwesa-cwebe nature reserve mostly mention local people’s discontent with not being consulted and being restricted on natural resource use in the reserve (ntshona et al. 2010). yet the history of planning of this protected area shows that during the planning stages, these issues were considered, and recommendations were made. firstly, moll’s (1974) preliminary report on the dwesa forest reserve not only highlighted the scientific value of the forests but also acknowledged the historical utilisation of the forests, grasslands and marine life, and recommended that they be proclaimed as a nature reserve to protect them from destruction. while recommending restrictions to harvesting of marine resources, he did not state that utilisation should be outlawed forever, but recommended, ‘… the amount of cropping must be limited to the sustainable yield available from the area’ (moll 1974). his final recommendation was: … [t]o re-emphasize that consultation and planning are the most important features of good long-term development. by having a well-planned, scientifically sound management plan for the future of an area like dwesa, one will ensure that the maximum benefit for all will be obtained. (moll 1974:5) secondly, in fulfilling their brief to develop a management plan for dwesa and cwebe nature reserves, tinley and van riet (1975) considered moll’s report and recognised that local people depended upon certain resources from the indigenous forests. they made specific recommendations about natural resource use by local people and how the reserves should respond to their needs. they recommended that thatch grass collection should be allowed to satisfy local requirements as well as to reduce fire hazard in the forests. they concurred with moll (1974) that rotational harvesting of shellfish should be implemented. tinley was particularly adamant that herbalists should be given full cooperation in the collection of traditional medicines from the forests but overuse should be guarded against. besides the natural resource use, tinley and van riet recommended that local people should receive preferential employment opportunities. they suggested benefit sharing of revenue generated from tourism with neighbouring communities. the recommendations made by moll, tinley and van riet were ignored by the transkei government in favour of a fences and fines conservation model. the post-apartheid era, the land claim and its aftermath since the advent of post-apartheid south africa in 1994, many opportunities opened up for the biodiversity conservation sector to also ride the waves of change and adopt more inclusive approaches. this happened, but for the most part it failed to consider local people’s historical links to a conservation area, including historical loss of land and other natural resource rights (kepe 2018). this is where the greatest opportunity for a reset lies. in dwesa-cwebe nature reserve the nationally televised invasion of the reserve by local people in september 1994, demanding access to resources, as well as the lodging of the land claim for the reserve in 1996, were opportunities to finally put this protected area on a new path (i.e. switch from a regime of exclusion and dispossession to one of inclusion and partnership), given all the publicity and attention. ntshona et al. (2010) argue that despite their successful land claim in 2001, the dwesa-cwebe communities have not been able to fully benefit from it, as a myriad of government policies and legislation have made it almost impossible for them to access certain resources within the reserve. in addition, following the ‘successful’ land claim resolution in 2001, the state, in its multiple forms (departments) has virtually taken a back seat, leaving confusion and conflict between local people and conservation managers to go on unchecked. the absence of a clear and honest implementation of land claim settlement terms is a lost opportunity. across the country, there is a need for intervention that should begin by acknowledging the failed implementation, or absence thereof, of land claim settlement agreements. in agreement with ntshona et al. (2010), local people’s: [r]ights that are enshrined in policy documents and legal agreements should be informed by the capacity of people to access those rights. there have to be accompanying legal provisions and the capacity to enforce them. (p. 360) currently, local people are generally left on their own, and in constant conflict among themselves, and with the state, despite policies, legislation and legal entities (e.g. community property association) that were supposed to mitigate the people-conservation tensions. a structural tension between legal entities such as the communal property associations and local, traditional governance structures are often mired in structural tensions that limit success of mitigating people-conservation tensions (ntshona et al. 2010). lessons from research on dwesa-cwebe and the wild coast at large the national profile of the dwesa-cwebe nature reserve over the last two decades or so, has encouraged a flurry of researchers from within and outside south africa, writing about people-conservation tensions, the land claim, use of natural resources within and outside the reserve, the marine protected area and so forth (e.g. de villiers 2021; fay 2007; ntshona et al. 2010; shackleton et al. 2007; sunde 2014; timmermans 2004). there is also some writing by journalists and consultants. whether explicit or not, collectively, these studies contain recommendations that, if taken seriously, could mitigate aspects of the current people-conservation tensions in dwesa-cwebe nature reserve. translating research findings into policy, or making them accessible to practitioners is challenging, but in the case of dwesa-cwebe nature reserve post-study dissemination workshops, policy briefs, op-eds and media coverage have become accessible to mitigate this past challenge. discussion and conclusion this article argues that people-conservation conflicts are common globally, but that their mitigation is of primary importance. while much has been done to mitigate these tensions, including ‘inclusive conservation’ (e.g. co-management between local people and conservation agencies), community based natural resource management, ‘conservation against poverty’, where alternative livelihood means are promoted to meet local livelihood needs, and neoliberal conservation, where markets and the private sector are seen to play a role in making conservation commercially viable, more is needed. this article argues that in each case of people-conservation conflict, there is a need to look back at important points in history, where opportunities existed that could have forged a better scenario and at least reduced the current tension. the article shows that this assessment of history allows for strategies to avoid past mistakes, while exploring how past opportunities could still be embraced or revived. in the case of dwesa-cwebe nature reserve, the article explores some missed opportunities that can explain some of the lingering people-conservation tensions in the area. firstly these include the opportunity to use the historical ties of the famous amaxhosa king sarhili to dwesa and cwebe forests, including his exclusive protection of dwesa. given that sarhili’s grave is already promoted as a tourist attraction, local people could renew their sense of belonging to the dwesa-cwebe nature reserve, and therefore see it in fresh light, were this similarly promoted the same way as the king’s grave. secondly, advice about community consultation and resource use that were given by the planners of dwesa-cwebe nature reserve during the 1970s should have been heeded. evidence shows that had some of those suggestion not been ignored, some of the tensions would have been much less. thirdly, the ushering in of the post-apartheid era that saw enthusiastic changes in policy and legislation, including in biodiversity conservation, as well as the ‘successful’ land claim for dwesa-cwebe nature reserve by local communities in 2001, was another opportunity that could have been used to reduce the historical tensions in the area. with almost no positive intervention by state departments that were supposed to implement the land claim settlement agreement for the benefit of the local communities, it is easy to see this as another missed opportunity. lastly, dwesa-cwebe nature reserve and the local communities with ties to it, have seen numerous research studies that yielded important recommendations. it is argued that lessons from these studies could shed light on a number of issues that currently contribute to the tension. these include information about natural resource use by local people, which vacillates between sustainable use of some resources, and extra-legal and sometimes destructive means of natural resource use (e.g. poaching using dogs) (de villiers 2021), understandings of the kind of rights that local people seek, and broadly, the political ecology of conservation in the wild coast. acknowledgements the authors are grateful to environmental officers and traditional authorities around dwesa-cwebe reserve. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions d.d.v. conducted the original fieldwork, wrote sections of the article and provided editing of sections of the final draft. t.k. conceptualised the idea of the article, wrote the first draft of the article and edited it for submission to the journal. ethical considerations ethical clearance to conduct this study was obtained from the rhodes university, science faculty ethics committee (no. sci2017/075). funding information this research received financial support from the economic development, environmental affairs and tourism, rhodes university and university of toronto. data availability data sharing is not applicable to this article, as no new data were collected or analysed. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references agrawal, a. & redford, k., 2009, ‘place, conservation, and displacement’, conservation & society 7(1), 56–58. https://doi.org/10.4103/0972-4923.54797 cawe, s., 1992, ‘coastal forests of transkei: their history and conservation value’, veld and flora 78(4), 114–117. cooper, k.h. & swart, w., 1992. transkei forestry survey. wildlife society of southern africa, durban. de villiers, d., 2021, ‘biodiversity conservation of south africa’s wild coast through the years: exploring the tensions between western-style and local traditional conservation practices’, unpublished phd thesis, rhodes university. dressler, w., büscher, b., schoon, m., brockington, d., hayes, t., kull, c. et al., 2010, ‘from hope to crisis and back again? a critical history of the global cbnrm narrative’, environmental conservation 37(1), 5–15. https://doi.org/10.1017/s0376892910000044 du toit, j.t., walker, b.h. & campbell, b.m., 2004, ‘conserving tropical nature: current challenges for ecologists’, trends in ecology and evolution 19(1), 12–17. https://doi.org/10.1016/j.tree.2003.09.018 fay, d., 2007, ‘struggles over resources and community formation at dwesa-cwebe, south africa’, the international journal of biodiversity science and management 3(2), 88–10. https://doi.org/10.1080/17451590709618165 feely, j., 2013, ‘guide to the natural history of the qawukeni coast eastern cape’, unpublished document. nelson mandela metropolitan university, port elizabeth. henkel, c.c., 1889, ‘report of the superintendent of woodland forests for the year 1888: report on crown forests in the transkeian territories of kaffraria proper with rough sketch map’, in cape of good hope annexures 1889/2, pp. 57–65. henkel, c.c., 1895, ‘report of the conservator of forests, transkeian conservancy, for the year 1894’, in cape of good hope annexures to the votes and proceedings of the house of assembly 6, pp. 130–135. hutton, j., adams, w.m. & murombedzi, j., 2005, ‘back to the barriers? changing narratives in biodiversity conservation’, forum for development studies 32(2), 341–370. https://doi.org/10.1080/08039410.2005.9666319 igoe, j. & brockington, d., 2007, ‘neoliberal conservation: a brief introduction’, conservation and society 5(4), 432–449. kepe, t., 2018, ‘meanings, alliances and the state in tensions over land rights and conservation in south africa’, in s. mollett & t. kepe (eds.), land rights, biodiversity and justice: rethinking parks and people, routledge publishers, abingdon, pp. 18–30. mbashe local municipality, 2022, tourism, viewed 26 october 2022, from https://www.mbhashemun.gov.za/tourism-2/. moll, e.j., 1974, a preliminary report on the dwesa forest reserve, transkei, wildlife society and university of cape town, rondebosch. ntshona, z., kraai, m., kepe, t. & saliwa, p., 2010, ‘from land rights to environmental entitlements: community discontent in the “successful” dwesa-cwebe land claim in south africa’, development southern africa 27(3), 353–361. https://doi.org/10.1080/0376835x.2010.498942 peires, j., 1989, the dead will arise: nongqawuse and the great xhosa cattle-killing of 1856–7, raven press, johannesburg. salafsky, n., 2011, ‘integrating development with conservation: a means to a conservation end or a mean end to conservation?’, biological conservation 144(3), 973–978. https://doi.org/10.1016/j.biocon.2010.06.003 sanderson, s.e. & redford, k.h., 2003, ‘contested relationship between biodiversity conservation and poverty alleviation’, oryx 37(4), 389–390. https://doi.org/10.1017/s003060530300070x shackleton, c., timmermans, h., nongwe, n., hamer, n. & palmer, r., 2007, ‘direct-use values of non-timber forest products from two areas on the transkei wild coast’, agrekon 46(1), 113–134. https://doi.org/10.1080/03031853.2007.9523764 sim, t.r., 1906, the forests and forest flora of the colony of the cape of good hope, government of the cape of good hope, aberdeen. slade, v.c., 2010, ‘producing sarhili: the colonial archive and the biographical limits of writing a history of a nineteenth century xhosa king’, unpublished m.a. thesis, university of the western cape. sunde, j., 2014, ‘customary governance and expressions of living customary law at dwesa-cwebe: contributions to small-scale fisheries governance in south africa’, phd thesis, university of cape town. timmermans, h., 2004, ‘rural livelihoods at dwesa/cwebe: poverty, development and natural resource use on the wild coast, south africa’, unpublished ma thesis, rhodes university. tinley, k.l. & van riet, w.f., 1975, planning and management proposals for the dwesa forest reserve, transkei, 88 pp., department of forestry and nature conservation, umtata. 161-12 oct 2009.indd [final version].indd a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe original research a rticle #161 (page number not for citation purposes) 2 spiders and scorpions (arachnida: araneae, scorpiones) of the nylsvley nature reserve, south africa abstract among other activities, the south african national survey of arachnida (sansa) aims to survey the biodiversity of arachnids in protected areas of south africa. the study presented here documents the diversity of spiders and scorpions collected from the nylsvley nature reserve (nnr), south africa over a 30-year period. the spider fauna of nnr contains 175 species (7.5% of the total recorded in south africa), in 131 genera and 37 families. thomisidae is the most diverse spider family in the reserve, with 33 species (18.9% of the total), followed by salticidae, with 20 species (11.4%), and araneidae, with 18 species (10.3%). the majority of species (125) are wandering spiders (71.4%), whereas 50 species (28.6%) build webs. wandering grounddwelling spiders comprise 52 species, whereas 73 wandering species have been collected from the vegetation. a total of 158 species are new records for the reserve and oxyopes tuberculatus lessert, 1915 is newly recorded for south africa. six spider species may be new to science. the scorpion fauna of nnr comprises fi ve species (5% of the total recorded in south africa) in three genera and two families. buthidae are more diverse in the reserve, with four species and two genera represented. the scorpion fauna of the reserve includes two fossorial and three epigeic species, representing fi ve ecomorphotypes: semi-zpsammophilous, pelophilous, lithophilous, corticolous and lapidicolous. five additional scorpion species may be recorded if the reserve is sampled more intensively using appropriate techniques. introduction the south african national survey of arachnida (sansa) was initiated in 1997 to survey the arachnid diversity of south africa (dippenaar-schoeman & craemer 2000). sansa encompasses several projects, including surveys and inventories of arachnids that are protected in parks and reserves; surveys and inventories of arachnid diversity in the fl oral biomes; and checklists of arachnid species for each province in south africa. although arachnids constitute an abundant and diverse group of invertebrates, knowledge of their diversity in south africa remains poor (dippenaar-schoeman 2002). the aim of the study presented here was to survey the spider and scorpion fauna of the nylsvley nature reserve (nnr), situated in the limpopo province of south africa. the reserve lies in the upper reaches of the nyl river on the nyl fl oodplain, the largest inland fl oodplain in south africa, which was recognised as a ramsar site in 1998. the savanna biome, a vegetation type where trees and grass are equally important components, covers approximately half of africa’s land surface and about 35% of south africa. nnr falls entirely within the biome and is one of the most intensively studied savanna sites in the world. more than 100 scientifi c papers and reports, many postgraduate degrees and several books were produced during the south african savanna biome project, conducted at nnr from 1974 to 1990. the aim of this project was to develop the understanding necessary to predict changes in the biome’s stability (scholes & walker, 1993). however, little information is available for the arachnids. the only published data on the spiders of nnr are a survey of spiders from abandoned mammal holes (heidger, 1988) and notes on the behaviour of a trapdoor spider, ancylotrypa brevicornis (hewitt 1919; leroy & leroy 2005). the study presented in this contribution is the eighth survey of the arachnid fauna of the savanna biome in south africa and the fi rst arachnid inventory for the nnr. other surveys of arachnids in the savanna biome were undertaken at roodeplaat dam nature reserve (dippenaar-schoeman, van den berg & van den berg 1989), makalali nature reserve (whitmore, slotow, crouch & dippenaarschoeman 2002; druce, hamer, slotow & prendini, 2004), western soutpansberg (foord, dippenaarschoeman & van der merwe 2002; in press), kruger national park (dippenaar-schoeman & leroy 2003), the inselberg at polokwane (pietersburg) (modiba, dippenaar & dippenaar-schoeman 2005), ndumo game reserve (haddad, dippenaar-schoeman & wesolowska 2006) and polokwane nature reserve (dippenaar, modiba, khoza & dippenaar-schoeman 2008). this contribution does not present the results of a quantitative survey, but instead summarises data from collections of spiders and scorpions made in the nnr over a period of 30 years, including specimens collected during the south african savanna biome project (scholes & walker 1993). although this survey may not refl ect the true diversity and species richness of spiders and scorpions in the reserve, it nonetheless provides an estimate of the species presently protected within its boundaries. study area nylsvley nature reserve (24º39’s 28º42’e) is situated 12 km south of mole mole (nylstroom) in the limpopo province of south africa. the altitude of nnr ranges between 1 080 m and 1 154 m above sea level, with an average altitude of 1 100 m. the reserve, which is located in the upper reaches of the nyl river fl oodplain, comprises 3 970 ha, of which 500 ha (10%) falls within authors: ansie s. dippenaarschoeman1 annette van den berg1 lorenzo prendini2 affi liations: 1arc-plant protection research institute, queenswood, south africa 1department of entomology and zoology, university of pretoria, south africa 2division of invertebrate zoology, american museum of natural history, new york correspondence to: ansie s. dippenaarschoeman e-mail: dippenaara@arc.agric.za postal address: arc-plant protection research institute, private bag x134, queenswood, 0121, south africa keywords: biodiversity; faunistics; savanna biome; south african national survey of arachnida dates: received: 26 june 2008 accepted: 31 aug. 2008 published: 23 jan. 2009 how to cite this article: dippenaar-schoeman a.s., van den berg a. & prendini l., 2009, ‘spiders and scorpions (arachnida: araneae, scorpiones) of the nylsvley nature reserve, south africa’, koedoe 51(1), art. #161, 9 pages. doi: 10.4102/koedoe. v51i1.161 this article is available at: http://www.koedoe.co.za © 2009. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. vol. 51 no. 1 page 1 of 9 koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research dippenaar-schoeman, van den berg & prendini a rt ic le # 16 1 (page number not for citation purposes) the nnr. the nyl river floodplain, the largest inland wetland in south africa (noble & hemens 1978), extends from middelfontein, west of mole mole in the south-west, to moorddrift, near molopane (potgietersrus) in the north-east. nnr is situated in the summer rainfall region, receives rainfall during the hot summer months and experience cool, dry winters. the rainfall is variable: the 69-year mean annual rainfall at nnr is 623 mm, with an annual coefficient of variation of 24%. the mean annual temperature is 19ºc (scholes & walker 1993). the maximum daily temperature ranges from a mean of 29ºc in december/january to 21ºc in june/july, whereas the minimum daily temperature varies from 17ºc in december/january to 4ºc in june/july. the nyl river floodplain comprises short grassy plains, reed beds, stands of long rice grass, open water patches, marshes, acacia (fineleaf) woodland and broadleaf woodland. approximately 600 plant species have been recorded in the nnr, and scholes and walker (1993) have distinguished nine vegetation types: burkea africana savanna; diplorhynchus condylocarpon savanna; combretum savanna; acacia tortilis savanna; old village sites; acacia karroo savanna; floodplain grasslands; grasslands on vertic soils; and seepline grassland. these vegetation types may be further grouped into two broad categories (low & rebelo 1996), namely mixed bushveld and clay thorn bushveld. methods collection techniques during the south african savanna biome project (1974 to 1990), spiders were collected from grass with sweepnets and from trees by beating. this material was donated to the national collection of arachnida, where it was sorted and identified. additional sporadic collecting was undertaken in the nnr from 1985 to 2005. during this period, spiders were collected by hand (ground and plant search, turning rocks, peeling bark, sifting leaf litter) or by using a sweepnet for grass and a beating tray for low shrubs. scorpions were collected by hand (mainly by turning rocks and peeling bark). material examined the spiders examined during the course of the study were identified by the first author and are deposited in the national collection of arachnida (nca) at the arc-plant protection research institute, pretoria. the scorpions were identified by the third author and are deposited in the nca, the american museum of natural history in new york, and the transvaal museum in pretoria. the lack of taxonomic research on certain spider families (e.g. lycosidae and theridiidae) in southern africa prevented the identification of some specimens to species. in some families, only immature specimens were collected and these were impossible to identify to species level. ecology a guild is a group of species that potentially compete for jointly exploited limited resources (uetz, halaj & cady 1999). because most spiders live in a defined environment with limitations imposed by both abiotic and biotic factors, species can be grouped into guilds based on information about their habitat preferences and predation methods. two main guilds of spiders were recognised in the present study, i.e. wandering spiders (w) and web builders (wb), with further subdivisions based on microhabitat and general behaviour, as defined by dippenaarschoeman and leroy (2003). guilds of scorpions were defined 3 and subdivided on the basis of the ecomorphotypes defined by prendini (2001, 2005). results and discussion spiders diversity a total of 175 spider species (7.5% of the total recorded in south africa), representing 132 genera and 37 families (tables 1 and 2) were collected in the nnr. thomisidae was the most diverse spider family, with 33 species (18.9% of the total), followed by salticidae, with 20 species (11.4%), araneidae, with 18 species (10.3%), and corinnidae, gnaphosidae, oxyopidae and theridiidae, with 10 (5.7%) species each. fifteen families were each represented by a single species (table 2) and 159 species are new records for the reserve (table 1). oxyopes tuberculatus lessert, 1915 is newly recorded for south africa. the pholcid, quamtana nylsvley huber, 2003, was originally described from the reserve (huber, 2003). six species of the genera anahita (ctenidae), diores and ranops (zodariidae), hamataliwa (oxyopidae), hypsosinga (araneidae) and theuma (prodidomidae), may be new to science (table 1). guilds although the majority of the spiders collected in the nnr, namely 125 species (71.4%) representing 24 families, are wanderers, 50 species (28.6%) in 13 families build webs. a total of 52 species (30%) in 22 families live on the ground. seven species (4%) in four families of ground-dwelling spiders are fossorial, whereas 45 species (26%) in 14 families are free-living. seventy-three species (42%) in 12 families inhabit vegetation, of which 36 species (20.8%) in 11 families construct webs to catch prey in the grass and herbaceous layer. fossorial species: seven species of the suborder mygalomorphae (baboon and trapdoor spiders) inhabit silk-lined burrows and are presently protected in most provinces of south africa. the baboon spider families barychelidae and theraphosidae are represented by one species, sipalolasma humicola (benoit, 1965), and four species, augacephalus junodi (simon, 1904), brachionopus pretoriae purcell, 1904, ceratogyrus darlingi pocock, 1897 and an unidentified species of harpactirella (gallon 2005) respectively. two trapdoor spider families are known from the nnr: segregara transvaalensis (hewitt, 1913) (idiopidae) and ancylotrypa brevicornis (hewitt, 1919) (cyrtaucheniidae). observations of the burrow-constructing behaviour of a. brevicornis showed that, in addition to constructing a thin wafer-lid trapdoor, these spiders use a hard, spherical plug (or marble) made of soil particles, held together by silk, to close the burrow entrance (leroy & leroy 2005). free-living ground species: of the 45 species in 14 families of free-living ground spiders collected in the nnr, gnaphosidae and corinnidae, each represented by 10 species, are the most abundant, followed by zodariidae, with six species, and lycosidae, with three species. plant-dwelling species: thirty-six species (20.8%) in 10 families are commonly found on the five dominant tree species in the nnr (table 3). eleven species (seven families) occur on burkea africana hock (wild syringa), eight species (five families) on combretum molle (r.br ex g. don) (velvet bush willow), 13 species (six families) on dombeya rotundifolia (hochst) planch. (wild pear), 14 species (six families) on grewia flavescens juss. (raisin bush), and 21 species (five families) on ochna pulchra hook (peeling plane). species that live permanently on tree bark include the long-spinnered bark spiders, hersilia sericea pocock, 1898 and h. setifrons lawrence, 1927 (foord & dippenaarschoeman, 2006). four tmarus spp. (thomisidae) and oxytate argenteooculata (strand, 1886) (thomisidae) are also exclusively arboreal. vol. 51 no. 1 page 2 of 9 spiders and scorpions original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #161 (page number not for citation purposes) 4 table 1 checklist of spider and scorpion species recorded in the nylsvley nature reserve, limpopo province, south africa species guild reference distribution araneae 1. family agelenidae c.l. koch 1837 (funnel-web spiders) benoitia ocellata (pocock 1900) fwb nr 4 2. family ammoxenidae simon 1893 (termite-eating spiders) ammoxenus amphalodes (dippenaar & meyer 1980) gw nr 4 3. family araneidae simon 1895 (orb-web spiders) araneilla sp. 1 owb nr araneus apricus (karsch 1884) owb nr 6 argiope australis (walckenaer 1805) owb nr 6 a. trifasciata (forskål 1775) owb nr 7 caerostris sexcuspidata (fabricius 1793) owb nr 6 cyphalonotus larvatus (simon 1881) owb nr 6 gasteracantha sanguinolenta (c.l. koch 1844) owb nr 6 hypsosinga lithyphantoides (caporiacco 1947) owb nr 6 hypsosinga sp. 2* owb nr isoxya cicatricosa (c.l. koch 1844) owb nr 6 nemoscolus elongatus (lawrence 1947) owb nr 4 neoscona blondeli (simon 1885) owb nr 6 n. moreli (vinson 1863) owb nr 6 n. subfusca (c.l. koch 1837) owb nr 6 n. triangula (keyserling 1864) owb nr 7 poltys furcifer (simon 1881) owb nr 6 pycnacantha tribulus (fabricius 1781) owb nr 6 singa lawrencei (lessert 1930) owb nr 6 4. family barychelidae simon 1889 (lesser baboon spiders) sipalolasma humicola (benoit 1965) bgw nr 5 5. family clubionidae wagner 1887 (sac spiders) clubiona africana (lessert 1921) pw nr 6 c. revillioidi (lessert 19360) pw nr 6 6. family corinnidae karsch 1880 (dark sac spiders) apochinomma formicaeforme (pavesi 1881) gw nr 6 brachyphaea sp. 1 gw nr castianeira fulvipes (simon 1896) gw nr 6 castianeira sp. 1 gw nr copa flavoplumosa (simon 1885) gw nr 6 corinnomma semiglabrum (simon 1896) gw nr 5 lessertina mutica (lawrence 1942) gw nr 4 merenius alberti (lessert 1923) gw nr 5 pronophaea natalica (simon 1897) gw nr 4 trachelas sp. 1 gw nr 7. family ctenidae keyserling 1877 (tropical wolf spiders) anahita sp. 1* gw nr 8. family cyrtaucheniidae simon 1892 (wafer-lid trapdoor spiders) ancylotrypa brevicornis (hewitt 1919) bgw leroy & leroy 2005 3 9. family deinopidae c.l. koch 1850 (net-throwing spiders) menneus camelus (pocock 1902) owb nr 4 10. family eresidae c.l. koch 1851 (velvet spiders) dresserus colsoni (tucker 1920) rwb nr 4 stegodyphus dumicola (pocock 1898) rwb nr 6 11. family gnaphosidae pocock 1898 (ground spiders) aphantaulax inornata (tucker 1923) gw nr 4 asemesthes ceresicola (tucker 1923) gw nr 4 asemesthes sp. 2 gw nr camillina corrugata (purcell 1907) gw nr 4 vol. 51 no. 1 page 3 of 9 original research dippenaar-schoeman, van den berg & prendini koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 16 1 (page number not for citation purposes) species guild reference distribution c. maun (platnick & murphy 1987) gw nr 5 echemus sp. 1 gw nr pterotricha varia (tucker 1923) gw nr 4 scotophaeus sp. 1 gw nr setaphis arcus (tucker 1923) gw nr 4 xerophaeus appendiculatus (purcell 1907) gw nr 4 12. family hersiliidae thorell 1870 (long-spinnered bark spiders) hersilia sericea (pocock 1898) pw foord & dippenaar-schoeman 2006 4 h. setifrons (lawrence 1928) pw foord & dippenaar-schoeman 2006 5 13. family idiopidae simon 1892 (front-eyed trapdoor spiders) segregara transvaalensis (hewitt 1913) bgw nr 4 14. family linyphiidae blackwall 1859 (hammock-web spiders) ceratinopsis sp. 1 swb nr microlinyphia sterilis (pavesi 1883) swb nr 6 tybaertiella krugeri (simon 1894) swb nr 6 15. family lycosidae sundevall 1833 (wolf spiders) lycosa sp. 1 gw nr pardosa crassipalpis (purcell 1903) gw nr 5 proevippa albiventris (simon 1898) gw nr 5 16. family miturgidae simon 1885 (sac spiders) cheiracanthium africanum (lessert 1921) pw nr 6 c. furculatum (karsch 1879) pw lotz 2007 6 17. family nephilidae simon 1894 (golden orb-web spiders) nephila fenestrata (thorell 1859) owb nr 6 n. senegalensis (walckenaer 1842) owb nr 6 18. family oecobiidae blackwall 1862 (ant eaters) oecobius navus (blackwall 1859) rwb nr 7 19. family oxyopidae thorell 1870 (lynx spiders) hamataliwa rostrifrons (lawrence 1928) pw nr 5 hamataliwa sp. 2* pw nr oxyopes affinis (lessert 1915) pw nr 6 o. jacksoni (lessert 1915) pw nr 6 o. russoi (caporiacco 1940) pw nr 6 o. schenkeli (lessert 1927) pw nr 6 o. tuberculatus (lessert 1915) pw nr (new record for south africa) 6 oxyopes sp. 6 pw nr oxyopes sp. 7 pw nr peucetia transvaalica (simon 1896) pw van niekerk & dippenaar-schoeman 1994 6 20. family palpimanidae thorell 1870 (palp-footed spiders) palpimanus transvaalicus (simon 1893) gw nr 4 21. family philodromidae thorell 1870 (small huntsman spiders) hirriusa variegata (simon 1895) gw nr 4 philodromus bigibbus (o.p. cambridge 1876) pw nr 6 p. guineensis (millot 1941) pw nr 6 philodromus sp. 3 pw nr philodromus sp. 4 pw nr thanatus africanus (karsch 1878) pw nr 6 tibellus hollidayi (lawrence 1952) pw nr 6 22. family pholcidae c.l. koch 1851 (daddy long-legs) quamtana nylsvley (huber 2003) spwb huber 2003 (type locality) 3 smeringopus natalensis (lawrence 1947) spwb nr 4 23. family pisauridae simon 1890 (nursery-web spiders) afropisaura rothiformis (strand 1908) pw nr 6 euprosthenops australis (simon 1898) fwb nr 6 table 1 (cont...) 5 vol. 51 no. 1 page 4 of 9 spiders and scorpions original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #161 (page number not for citation purposes) 6 species guild reference distribution euprosthenopsis armata (strand 1913) fwb nr 6 maypacius bilineatus (pavesi 1895) swb nr 6 thalassius spinosissimus (karsch 1879) gw nr 6 24. prododomidae simon 1884 (long-spinnered ground spiders) theuma sp. 1* gw nr 25. family salticidae blackwall 1841 (jumping spiders) afraflacilla sp. 1 pw nr asemonea sp. 1 pw nr baryphas ahenus (simon 1902) pw nr 5 bianor albobimaculatus (lucas 1846) gw nr 7 brancus bevisi (lessert 1925) pw nr 6 festucula lawrencei (lessert 1933) pw nr 6 habrocestum sp. 1 pw nr heliophanus insperatus (wesolowska 1986) gw nr 5 heliophanus sp. 2 pw nr hyllus bevisi (lessert 1925) pw nr 6 h. brevitarsis (simon 1902) pw nr 6 h. treleaveni (peckham & peckham 1902) pw wesolowska & cumming 2004 6 menemerus sp. 1 pw nr mogrus albogularis (simon 1901) pw nr 4 myrmarachne sp. 1 pw nr pachyballus transversus (simon 1900) pw nr 6 pellenes sp. 1 gw nr phlegra sp. 1 gw nr rhene machadoi (berland & millot 1941) pw nr 6 thyene coccineovittata (simon 1885) pw nr 6 26. family scytodidae blackwall 1864 (spitting spiders) scytodes sp. 1 gw nr 27. family segestriidae simon 1893 (tube spiders) ariadna sp. 1 rwb nr 28. family selenopidae simon 1897 (flatties) anyphops sp. 1 gw nr 29. family sparassidae bertkau 1872 (huntsman spiders) olios correvoni (lessert 1921) pw nr 6 palystes superciliosus (l. koch 1875) pw nr 5 pseudomicrommata longipes (bösenberg & lenz 1895) pw nr 6 30. family tetragnathidae menge 1866 (water orb-web spiders) leucauge festiva (blackwall 1866) owb nr 6 tetragnatha demissa (l. koch 1872) owb nr 6 tetragnatha isidis (simon 1880) owb nr 7 31. family theraphosidae thorell 1870 (baboon spiders) augacephalus junodi (simon 1904) bgw gallon 2005 5 brachionopus pretoriae (purcell 1904) bgw nr 4 ceratogyrus darlingi (pocock 1897) bgw gallon 2005 (as bechuanicus) 5 harpactirella sp. 1 bgw nr 32. family theridiidae sundevall 1833 (comb-footed spiders) achaearanea sp. 1 gwb nr argyrodes convivans (lawrence 1937) gwb nr 4 dipoena sp. 1 gwb nr enoplognatha inornata (o.p. cambridge 1904) gwb nr 4 episinus bilineatus (simon 1894) gwb nr 4 euryopis episinoides (walckenaer 1847) gw nr 4 latrodectus geometricus (c.l. koch 1841) gwb nr 7 l. renivulvatus (dahl 1902) gwb nr 6 phoroncidia eburnea (simon 1895) gwb nr 4 table 1 (cont...) vol. 51 no. 1 page 5 of 9 original research dippenaar-schoeman, van den berg & prendini koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 16 1 (page number not for citation purposes) 7 species guild reference distribution steatoda capensis (hann 1990) gwb nr 6 33. family thomisidae sundevall 1833 (crab spiders) camaricus nigrotesselatus (simon 1895) pw nr 6 diaea puncta (karsch 1884) pw nr 6 heriaeus transvaalicus (simon 1895) pw nr 5 hewittia gracilis (lessert 1928) pw nr 6 misumenops rubrodecoratus (millot 1942) pw dippenaar-schoeman 1983 6 monaeses gibbus (dippenaar-schoeman 1984) pw nr 4 m. pustulosus (pavesi 1895) pw nr 6 m. quadrituberculatus (lawrence 1927) pw nr 5 oxytate argenteooculata (strand 1886) pw nr 6 pactactes compactus (simon 1895) pw nr 4 parabomis martini (lessert 1919) pw nr 6 pherecydes lucinae (dippenaar-schoeman 1980) pw dippenaar-schoeman 1980 4 p. zebra (lawrence 1927) pw nr 6 runcinia affinis (simon 1897) pw nr 6 r. erythrina (jézéquel 1964) pw nr 6 r. flavida (simon 1881) pw dippenaar-schoeman 1980 6 r. grammica (l. koch 1937) pw nr 7 smodicinus coroniger (simon 1895) pw nr 6 synema imitator (pavesi 1883) pw nr 6 thomisops bullatus (simon 1895) pw nr 5 t. pupa (karsch 1879) pw nr 6 t. sulcatus (simon 1895) pw nr 6 thomisus blandus (karsch 1880) pw nr 6 t. citrinellus (simon 1875) pw dippenaar-schoeman 1983 6 t. congoensis (comellini 1957) pw dippenaar-schoeman 1983 6 t. kalaharinus (lawrence 1936) pw nr 6 t. scrupeus (simon 1886) pw dippenaar-schoeman 1983 6 t. stenningi (pocock 1900) pw nr 6 tmarus africanus (lessert 1919) pw dippenaar-schoeman 1984 6 t. cameliformis (millot 1942) pw dippenaar-schoeman 1984 6 t. comellinii (garcia-neto 1989) pw nr 6 t. planetarius (simon 1903) pw nr 6 xysticus fagei (lessert 1919) gw nr 6 34. family trochanteriidae karsch 1879 (scorpion spiders) platyoides walteri (karsch 1886) gw nr 6 35. family uloboridae thorell 1869 (hackled orb-web spiders) miagrammopes brevicaudus (o.p. cambridge 1882) owb nr 4 uloborus plumipes (lucas 1846) owb nr 6 uloborus sp. 2 owb nr 36. family zodariidae thorell 1881 (burrowing spiders/ant-eating spiders) caesetius sp. 1 gw nr capheris decorata (simon 1904) gw nr 4 chariobas sp. 1 gw nr diores recurvatus (jocqué 1990) gw nr 4 diores sp. 2* gw nr ranops sp. 1* gw nr 37. family zoropsidae bertkau 1882 (ground spiders) griswoldia sp. rwb nr scorpiones 1. family buthidae c.l. koch 1837 (thick-tail scorpions) parabuthus mossambicensis (peters 1861) fs prendini 2001, 2005 5 uroplectes planimanus (karsch 1879) li prendini 2001, 2005 5 u. triangulifer (thorell 1876) la prendini 2001, 2005 4 table 1 (cont...) vol. 51 no. 1 page 6 of 9 spiders and scorpions original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #161 (page number not for citation purposes) 8 families genera species % of total thomisidae 17 33 18.86 salticidae 17 20 11.43 araneidae 13 18 10.29 corinnidae 10 10 5.71 gnaphosidae 8 10 5.71 oxyopidae 3 10 5.71 theridiidae 9 10 5.71 philodromidae 4 7 4.00 zodariidae 5 6 3.43 pisauridae 5 5 2.86 theraphosidae 4 4 2.29 linyphiidae 3 3 1.71 lycosidae 3 3 1.71 sparassidae 3 3 1.71 uloboridae 2 3 1.71 eresidae 2 2 1.14 hersiliidae 1 2 1.14 miturgidae 1 2 1.14 nephilidae 1 2 1.14 pholcidae 2 2 1.14 tetragnathidae 2 2 1.14 clubionidae 1 2 1.14 agelenidae 1 1 0.57 ammoxenidae 1 1 0.57 barychelidae 1 1 0.57 ctenidae 1 1 0.57 cyrtaucheniidae 1 1 0.57 deinopidae 1 1 0.57 idiopidae 1 1 0.57 oecobiidae 1 1 0.57 palpimanidae 1 1 0.57 prodidomidae 1 1 0.57 scytodidae 1 1 0.57 segestriidae 1 1 0.57 selenopidae 1 1 0.57 trochanteriidae 1 1 0.57 zoropsidae 1 1 0.57 total spiders 132 175 100.00 buthidae 2 4 80.00 scorpionidae 1 1 20.00 total scorpions 3 5 100.00 table 2 spiders and scorpions collected in the nylsvley nature reserve, limpopo province, south africa, indicating the number of species as a percentage of the total collected table 1 (cont...) species guild reference distribution u. vittatus (thorell 1876) c prendini 2001, 2005 5 2. family scorpionidae latreille, 1802 (burrowing scorpions) opistophthalmus glabrifrons (peters 1861) fp prendini 2001, 2005 5 * possible new species guild: bgw = burrow ground dwellers; fwb = funnel-web; gw = ground dwellers; gwb = gumfoot-web; pw = plant dwellers; rwb = retreat webs; swb = sheet-web; fs = fossorial semi-psammophilous; fp = fossorial pelophilous; c = corticolous; li = lithophilous; la = lapidicolous. reference: nr = new fossorial record. distribution: 1 = endemic to reserve; 2 = endemic to the limpopo province; 3 = near endemic to the limpopo province (occurs in two provinces); 4 = endemic to south africa; 5 = endemic to southern africa; 6 = endemic to the afrotropical region; 7 = cosmopolitan, occurs outside the afrotropical region. most of the araneid orb-web species found on vegetation are nocturnal and remove their webs early each morning. six araneid species, araneus apricus (karsch 1884), caerostris sexcuspidata (fabricius 1793), cyphalonotus larvatus (simon 1881), neoscona subfusca (c.l. koch 1837), n. triangula (keyserling 1864) and poltys furcifer simon, 1881, were collected from trees to which they retreat when inactive during the day. the nnr has a rich fauna of graminicolous spider species, most with elongated bodies resembling grass stems. these include the runcinia spp. (thomisidae), tibellus hollidayi lawrence 1952 (philodromidae) and pseudomicrommata longipes (bösenberg & lenz 1895) (sparassidae). other graminicolous spider species belong to the families oxyopidae, salticidae and thomisidae. web-building species: the orb-web spiders, comprising 25 species in the families araneidae (18 species), nephilidae (two), tetragnathidae (three) and uloboridae (three), are the most diverse group recorded. nephilidae and tetragnathidae are often seen in their large webs during the day. species of tetragnathidae (tetragnatha and leucauge) are associated with wetlands and found commonly in the nnr. ten species construct webs on or close to the soil surface. benoitia ocellata (pocock 1900) an agelenid, is often observed in the nnr, especially early in the mornings when dew collects on the sheet part of its web. the very large euprosthenops australis simon, 1898 (pisauridae) is another common spider, which constructs very large funnel webs next to trees, the funnel usually originating from an abandoned termite nest or mammal burrow. the retreat webs of eresidae (dresserus colsoni tucker 1920), oecobiidae, segestriidae and zoropsidae are constructed under rocks and plant debris, whereas the sheet webs of linyphiidae and gumfoot webs of theridiidae are constructed in grass close to the ground. the space webs of pholcidae are constructed mainly in abandoned mammal holes (heidger 1988). stegodyphus dumicola pocock, 1898 is a community web spider that constructs large nest-like retreats containing numerous spiders on vegetation. scorpions diversity five scorpion species (5% of the total recorded in south africa), representing three genera and two families, have been collected in the nnr (table 1). family buthidae is represented by two genera and three species, whereas scorpionidae is represented by a single genus and species. guilds the five scorpion species recorded in the nnr can be grouped into two different guilds: fossorial or burrowing species (40% of the species), and epigean species, which do not construct burrows (60%). each of these guilds can be further subdivided into distinct ecomorphotypes. fossorial species: two of the scorpion species collected in the nnr are adapted for constructing burrows and spend most of their time below the soil surface, i.e. they are fossorial. these species represent two distinct ecomorphotypes, associated with different substrata: semi-psammophilous and pelophilous species (prendini 2001). parabuthus mossambicensis (peters 1861) is a semi-psammophilous species that constructs burrows in semi-consolidated sand to sandy-loam substrata. burrows are constructed in open ground using the thickened metasoma to loosen the soil and the anterior two pairs of legs to scrape and rake soil out of the burrow. vol. 51 no. 1 page 7 of 9 original research dippenaar-schoeman, van den berg & prendini koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 16 1 (page number not for citation purposes) trees (especially acacias), often several metres above the ground. this species exhibits a few ecomorphological adaptations, for example moderate dorsoventral compression, elongation of the metasoma and pedipalps, and well-developed telotarsal ungues. uroplectes triangulifer (thorell 1876) is a lapidicolous species that shelters under loose stones, wood or debris at ground level, and displays few ecomorphological adaptations for this generalist lifestyle. species not recorded: up to five additional scorpion species, representing five genera and two families, may be recorded in the nnr if the scorpion fauna is more intensively sampled using appropriate techniques, e.g. pitfall trapping and night collecting by ultraviolet detection. the nnr falls within the known distributional range of these widespread savanna scorpion species, and suitable habitat occurs within its boundaries, suggesting that at least some of them may be present. the species include two buthids, the semipsammophilous parabuthus transvaalicus purcell, 1899 and lapidicolous uroplectes carinatus (pocock 1890), and three liochelids, the pelophilous cheloctonus jonesii pocock, 1892, lithophilous hadogenes sp., and corticolous opisthacanthus asper (peters 1861). conclusion arachnids form an important component of healthy ecosystems. any approach to conservation must consider the composition of the arachnid fauna. inventories with resulting checklists provide valuable baseline data about the species present in reserves and are the first step towards a better understanding of the fauna. preliminary investigations of the biodiversity of arachnids in south africa have highlighted the lack of baseline data on the ecology and diversity of most arachnid orders (dippenaarschoeman 2002). the survey presented here forms part of the south african national survey of arachnida (sansa) and the data gathered will be used in the savanna biome database. the nnr has a rich fauna of arachnid species (180) and, this species displays several ecomorphological adaptations to increase locomotor and burrowing efficiency in soft substrata, including carinae and spiniform processes on the metasoma and comb-like rows of long macrosetae on the prolateral and retrolateral margins of the tibia and basitarsi of the first and second pairs of legs. opistophthalmus glabrifrons peters, 1861 is a pelophilous species that constructs burrows in hard sandy-loam and clay substrata. burrows are constructed in open ground or under stones using the enlarged chelicerae to loosen the soil, and the anterior two pairs of legs to scrape and rake soil out of the burrow; the metasoma is also used for tail-scraping operations. this species displays several ecomorphological adaptations to increase burrowing efficiency in hard substrata, including short, robust legs and telotarsal ungues; stout, spiniform macrosetae distributed laterally and distally on the basitarsi; and carinae and spiniform processes on the metasoma. epigean species: three (60%) of the scorpion species recorded in the nnr are found at or above the soil surface and are unable to construct burrows, i.e. they are epigean. these species represent three distinct ecomorphotypes associated with different substrata: lithophilous, corticolous and lapidicolous species (prendini 2001). uroplectes planimanus (karsch 1879) is a lithophilous species adapted to life in the narrow cracks and crevices of rocks and under rocks resting on bedrock. this species is characterised by moderate dorsoventral compression; elongation of the metasoma and pedipalps; and stout macrosetae on the telotarsi, operating in conjunction with curved telotarsal ungues to provide grip on rock surfaces. these specialised adaptations facilitate rapid locomotion in any spatial plane of their rock habitats, but hinder locomotion on other substrata. these scorpions are therefore restricted to weathered rock outcrops. uroplectes vittatus (thorell 1876) is an obligate corticolous species, found in holes or under the loose bark of old or dead tree species spider species burkea africana hock araneidae: neoscona triangula hersiliidae: hersilia sericea oxyopidae: oxyopes russoi philodromidae: philodromis bigibbus salticidae: afraflacilla sp.; baryphas ahenus; brancus bevisi; rhene machadoi. sparassidae: olios sp. thomisidae: oxytate argenteooculata; tmarus cameliformis combretum molle (r.br ex g. don) araneidae: araneus apricus miturgidae: cheiracanthium furculatum oxyopidae: oxyopes russoi philodromidae: philodromus guineensis thomisidae: pactactes compactus; tmarus africanus; t. cameliformis; t. comellini dombeya rotundifolia (hochst) planch araneidae: poltys furcifer oxyopidae: hamataliwa rostrifrons philodromidae: philodromus guineensis salticidae: mogrus albogularis theridiidae: dipoena sp. thomisidae: diaea puncta; parabomis martini; misumenops rubrodecoratus; monaeses pustulosus; oxytate argenteooculata; synema sp.; tmarus africanus; t. cameliformis grewia flavescens juss araneidae: cyphalonotus larvatus oxyopidae: hamataliwa rostrifrons; oxyopes russoi; o. schenkeli salticidae: baryphas ahenus; brancus bevisi; festucula lawrencei. theridiidae: episinis sp. thomisidae: oxytate argenteooculata; synema sp.; tmarus africanus; t. cameliformis; t. planetarius uloboridae: miagrammopes brevicaudus ochna pulchra hook araneidae: araneus apricus; cyphalonotus larvatus; neoscona subfusca oxyopidae: hamataliwa rostrifrons; oxyopes schenkeli; peucetia transvaalica philodromidae: philodromus bigibbus salticidae: afraflacilla sp.; baryphas ahenus; brancus bevisi; mogrus albogularis; myrmarachne sp.; rhene machadoi thomisidae: diaea puncta; misumenops rubrodecoratus; oxytate argenteooculata; pactactes compactus; synema sp.; thomisus citrinellus; tmarus africanus; t. cameliformis table 3 spider species collected from five tree species commonly found in the nylsvley nature reserve, limpopo province, south africa 9 vol. 51 no. 1 page 8 of 9 spiders and scorpions original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #161 (page number not for citation purposes) 36 10 gallon, r.c., 2005, ‘a new species of theraphosid spider from southern africa (araneae, theraphosidae, harpactirinae) with distributional notes on other harpactirines’, bulletin of the british arachnological society 13, 179–184. haddad, c.r., dippenaar-schoeman, a.s. & wesolowska, w., 2006, ‘a checklist of the non-acarine arachnids (chelicerata: arachnida) of the ndumo game reserve, maputaland, south africa’, koedoe 49, 1–22. heidger, c., 1988, ‘ecology of spiders inhabiting abandoned mammal burrows in south african savanna’, oecologia 76, 303–306. huber, b.a., 2003, ‘southern african pholcid spiders: revision and cladistic analysis of quamtana gen. nov. and spermophora hentz (araneae: pholcidae), with notes on male-female covariation’, zoological journal of the linnean society 139, 77–527. leroy, a. & leroy, j., 2005, ‘notes on the natural history of a trapdoor spider ancylotrypa simon (araneae, cyrtaucheniidae) that constructs a spherical burrow plug’, journal of arachnology 33, 558–562. lotz, l.n., 2007, the genus cheiracanthium (araneae: miturgidae) in the afrotropical region. 1. revision of known species, navorsinge van die nasionale museum, bloemfontein 23, 1–76. low, a.b. & rebelo, h.g., 1996, vegetation of south africa, lesotho and swaziland, department of environment affairs and tourism, pretoria. modiba, m.a., dippenaar, s.m. & dippenaar-schoeman, a.s., 2005, ‘a checklist of spiders from sovenga hill, an inselberg in the savanna biome, limpopo province, south africa (arachnida: araneae)’, koedoe 48, 109–115. noble, r.g. & hemens, j., 1978, ‘inland water ecosystems in south africa – a review of research needs’, south african national scientific programmes report 34, 1–150. prendini, l., 2001, ‘substratum specialization and speciation in southern african scorpions: the effect hypothesis revisited’, in v. fet & p.a. selden (eds.), scorpions 2001. in memoriam gary a. polis, pp. 113–138. british arachnological society, burnham beeches. prendini, l., 2005, ‘scorpion diversity and distribution in southern africa: pattern and process’, in b.a. huber, b.j. sinclair & k.-h. lampe (eds.), african biodiversity: molecules, organisms, ecosystems: 5th international symposium on tropical biology proceedings, bonn, germany, 2-6 may, 2004, pp. 25– 68. scholes, r.j. & walker, b.h., 1993, an african savanna, synthesis of the nylsvley study, cambridge university press, cambridge. uetz, g.w., halaj, j. & cady, a.b., 1999, ‘guild structure of spiders in major crops’, journal of arachnology 27, 270–280. van niekerk, p. & dippenaar-schoeman, a.s., 1994, ‘a revision of the afrotropical species of peucetia (araneae: oxyopidae)’, entomology memoir 89, 1–50. wesolowska, w. & cumming, m.s., 2004, ‘a redescription and natural history of hyllus treleaveni peckham et peckham, 1902, the largest jumping spider in africa (araneae: salticidae)’, annales zoologici fennici 54, 579–586. whitmore, c., slotow, r., crouch, t.e. & dippenaar-schoeman, a.s., 2002, ‘diversity of spiders (araneae) in a savanna reserve, northern province, south africa’, journal of arachnology 30, 344–356. although this contribution reports on sporadic collecting and probably represents only a subset of the arachnids present, we hope it will stimulate further research on the arachnids of the nnr and savanna biome. acknowledgements we thank gherda ferreira, who collected arachnids during the savanna biome survey, the spider club of southern africa, and the friends of nylsvley for additional collecting. this work forms part of the south african national survey of arachnida (sansa), which is coordinated by the south african national biodiversity institute and the agricultural research council. references dippenaar, s.m., modiba, m.a., khoza, t.t. & dippenaarschoeman, a.s., 2008, ‘a checklist of spiders (arachnida, araneae) of the polokwane nature reserve, limpopo province, south africa’, koedoe 50, 10–17. dippenaar-schoeman, a.s., 1980, ‘the crab-spiders of southern africa (araneae: thomisidae). 1. the genus runcinia simon, 1875’, journal of the entomological society of southern africa 43, 303–326. dippenaar-schoeman, a.s., 1983, ‘the spider genera misumena, misumenops, runcinia and thomisus (araneae: thomisidae) of southern africa’, entomology memoirs of the department of agricultural technical services, republic south africa 55, 1–66. dippenaar-schoeman, a.s., 1984, ‘the crab-spiders of southern africa (araneae: thomisidae). 4. the genus monaeses thorell, 1869’, phytophylactica 16, 101–116. dippenaar schoeman, a.s., 2002, ‘status of south african arachnida fauna’, in endangered wildlife trust (ewt) of south africa symposium on the status of south african species proceedings, rosebank, south africa, september 4–7, 2002, n.p. dippenaar-schoeman, a.s. & craemer, c., 2000, ‘the south african national survey of arachnida’, plant protection news 56, 11–12. dippenaar-schoeman, a.s. &. leroy, a., 2003, ‘a checklist of the spiders of the kruger national park, south africa (arachnida: araneae)’, koedoe 46, 91–100. dippenaar-schoeman, a.s., van den berg, a.m. & van den berg, a., 1989, ‘species composition and relative seasonal abundance of spiders from the field and tree layers of the roodeplaat dam nature reserve’, koedoe 32, 51–60. druce, d., hamer, m., slotow, r. & prendini, l., 2004, ‘checklist of millipedes (diplopoda), centipedes (chilopoda) and scorpions (arachnida: scorpionida) from a savanna ecosystem, limpopo province, south africa’, african invertebrates 45, 315–322. foord, s.h. & dippenaar-schoeman, a.s., 2006, ‘a revision of the afrotropical species of hersilia audouin (araneae: hersiliidae)’, zootaxa 1347, 1–92. foord, s.h., dippenaar-schoeman, a.s. & van der merwe, m., 2002, ‘a checklist of the spider fauna of the western soutpansberg, south africa (arachnida: araneae)’, koedoe 45, 35–43. foord, s.h., mafadza, m.m., dippenaar-schoeman, a.s. & van rensburg, b.j., in press, ‘micro-scale heterogeneity of spiders (arachnida: araneae) in the soutpansberg, south africa: a comparative survey and inventory in representative habitats’, african zoology. vol. 51 no. 1 page 9 of 9 introduction material and methods results discussion acknowledgements references about the author(s) alice bernard rehabs, international research laboratory, french national centre for scientific research (cnrs), university of lyon 1, nelson mandela university, george, south africa biometry and evolutionary biology laboratory (lbbe), french national centre for scientific research (cnrs) (unit 5558), university of lyon 1, villeurbanne, france sustainability research unit, nelson mandela university, george, south africa lizette moolman scientific services, south african national parks, knysna, south africa melanie a. de morney scientific services, south african national parks, knysna, south africa chloé guerbois rehabs, international research laboratory, french national centre for scientific research (cnrs), university of lyon 1, nelson mandela university, george, south africa sustainability research unit, nelson mandela university, george, south africa jan a. venter rehabs, international research laboratory, french national centre for scientific research (cnrs), university of lyon 1, nelson mandela university, george, south africa department of conservation management, faculty of science, nelson mandela university, george, south africa hervé fritz rehabs, international research laboratory, french national centre for scientific research (cnrs), university of lyon 1, nelson mandela university, george, south africa sustainability research unit, nelson mandela university, george, south africa citation bernard, a., moolman, l., de morney, m.a., guerbois, c., venter, j.a. & fritz, h., 2023, ‘height-related detection bias in camera trap surveys: insights for combining data sets’, koedoe 65(1), a1734. https://doi.org/10.4102/koedoe.v65i1.1734 note: additional supporting information may be found in the online version of this article as online appendix 1. short communication height-related detection bias in camera trap surveys: insights for combining data sets alice bernard, lizette moolman, melanie a. de morney, chloé guerbois, jan a. venter, hervé fritz received: 24 aug. 2022; accepted: 17 mar. 2023; published: 30 may 2023 copyright: © 2023. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. introduction camera trap surveys is a non-invasive method for monitoring wildlife (royle 2011). camera traps inevitably record non-target species (burton et al. 2015) and this by-catch is potentially useful (edwards et al. 2011; hofmeester et al. 2020; mazzamuto, lo valvo & anile 2019). the usefulness of by-catch data is determined by the probability of capturing an animal within a given survey design (findlay, briers & white 2020). as described by findlay et al. (2020), an animal has to pass in front of (encounter probability), trigger (trigger probability), be photographed by the camera (registration probability), and the image must be of sufficient quality to be able to identify the animal (capture quality probability). the probability of a camera trap being triggered differs between species and can be influenced by their body mass, size or behaviour, environmental variables (land cover, temperature), as well as camera trap brand, model and height of set-up (apps & mcnutt 2018; hofmeester et al. 2019; kolowski, oley & mcshea 2021; meek et al. 2016a; swann et al. 2004). to ensure the highest registration and capture probabilities for a target species or to maximise the diversity of photographed species, camera placement is crucial (burton et al. 2015). camera height is one of the critical settings, athough few studies have focused on it, whereas it has been shown to affect species detection (anile & devillard 2016; burton et al. 2015; hofmeester et al. 2019; meek et al. 2016a; palencia et al. 2021). camera height settings usually differ between surveys aimed at monitoring human activities and surveys aimed at monitoring biodiversity (burton et al. 2015). little emphasis has been put on using by-catch data from surveys targeting humans for biodiversity monitoring purposes. the effect of camera height on detection probability of mammal species was tested by comparing a standard height for monitoring biodiversity (50 cm, used in the african snapshot safari survey, meek et al. 2016b; pardo et al. 2021) with a height chosen to monitor human-related events (130 cm, used in a human-wildlife interface monitoring project, moolman et al. 2019). it was hypothesised that lower cameras would detect smaller mammals at higher frequencies than the higher cameras, but that no difference would occur between camera heights in detecting larger species. the aim was to assess if data sets from various surveys (humans or mammals as primary survey targets) using different camera heights could be combined. an investigation to test whether one of the height settings could maximise the detectability of both human and other mammal species activity in order to optimise camera trap survey design was also done. material and methods study site ten passive infrared-triggered (bushnell trophy cam hd aggressor no glow) cameras were set up at five different locations in the harkerville section of the garden route national park in south africa (-34.046731, 23.209715; online appendix figure 1-a1). all cameras were deployed in afrotemperate forest, on hiking trails. two cameras were set up at each location, at two different heights, on the same tree, no further than 1 m away from the hiking trails (online appendix figure 1-a1). high cameras were placed at a height of 130 cm (± 5 cm) from the ground level of the trail, which was the set-up for the human-wildlife interface survey, and low cameras at 50 cm (± 3 cm), which is the average shoulder height of the mammal species (mean height = 53.1 cm; height range: 15 cm to 80 cm) in the area. the two cameras were oriented and angled towards the same direction to photograph the same area. they were diagonally facing the trails, so that animals had to walk towards or away from the camera, to have a longer area of detection. the horizontal and vertical shift from the high camera compared to the low camera was measured. five points on the low camera image were identified and the relative position of these on the high camera was compared by measuring the relative deviation on the photograph (online appendix figure 2-a1 [a]). the mean vertical deviation was 5% and horizontal deviation was less than 2% between low and high cameras (online appendix figure 2-a1 [b]). the cameras used black flash (no glow), that is, infrared illumination, that prevents animals from being disturbed and being detected by humans at night. to reduce the risk of theft, cameras were protected by cable locks and metal casings. cameras were active 24 h per day and were set up to take one eight mega-pixels image per trigger, with a delay of 1 s between subsequent triggers. the sensitivity was set on ‘normal’. the flash intensity was set on ‘high’. data were collected from 09 march 2020 to 06 august 2020. humans moved cameras at two sites. data from after they were moved were excluded from the analyses. cameras were visited every 3 months to change batteries and sd cards. the researchers ensured that the time set on the cameras was identical. when servicing the cameras, both cameras took pictures of the researchers walking towards them; hence, the differences observed in the study were not induced by a malfunctioning camera. data processing the images were processed using the camerabase software (tobler 2007). for each image, the species name and the number of individuals were manually tagged, while the software automatically extracted the time and date of each photo. humans, their activities (walking, motorcycling), vehicles and dogs were tagged as such (online appendix table 1-a1). when an image was dark, the brightness was adjusted. when it was not possible to identify a species because a part of it was visible, the photo was tagged as non-identified. in order to test the impact of camera height setting from these two types of surveys on the analysis of a potential combined data set and because there was an average of 9.5 photos for each event, only photos from the same camera and with the same species that had a minimum period of 30-min between image time stamps were retained. this procedure is standard practice to obtain a list of independent capture events for camera trap data analysis, and the authors wanted to position their comparison within commonly used practices. similar events of both camera heights for a given location was merged by date and time (‘merge’ function, ‘base’ r package), in order to produce a comparative list of all the different capture events per camera. when the two cameras were triggered simultaneously and successfully captured the same event, the pair was classified as identical. events that could not be identified on either camera were deleted (hofmeester et al. 2020). for the events that were not similar between the two cameras, the remaining capture events were manually processed by comparing the images of both cameras. when only one of the two cameras was triggered, meaning that one could not find any photo for the other camera at that time, the capture event was classified as not triggered. when the image of one of the two cameras was empty (i.e. only background vegetation is visible but no mammal or human), corresponding to a lack of capture, the photo was coded as empty, and when a camera was triggered but it was not possible to identify any species, for example, because the image was too dark even after adjusting the brightness, corresponding to a poor image quality, it was classified as dark. for each non-identical event (i.e. events classified as not triggered, empty or dark), the species that was detected by the other camera was classified as specified. the information was noted in excel. finally, each capture event was categorised according to whether it was taken during the day or at night based on the sunset or sunrise time (‘sunrise.set’ function, ‘streammetabolism’ r package). data analyses all statistical analyses were performed in r (r core team 2020) and all the graphics were plotted using the ‘ggplot2’ r package (wickham 2016). general detection model to compare the detection similarity of different species and human activities between the two treatments (low camera vs. high camera), a 0 was assigned to events classified as not triggered, empty or dark, and a 1 was assigned to identical events. a generalised linear model was built to perform a logistic regression, that is, with a binomial distribution of the response variable, to explain the detection of the cameras. the ‘mass’ package (venables & ripley 2002) was used and the site was included as a variable to control for deviation between two cameras at the same site. the authors also tested the influence of the period of the day (day vs. night) on the detection probability. for each detected species, the average weight and height were compiled to test how they could affect the variation in detection between camera heights (child et al. 2016; hofmeester et al. 2020) (online appendix table 1-a1). weight and height were log-transformed to conform with parametric modelling constraints. thereafter, a backward stepaic model selection procedure (‘stepaic’ function, ‘mass’ package) was used to search for the best combination of variables to get a simplified model. the predicted detection probabilities were calculated (‘predict’ function, ‘car’ package) and plotted (‘ggpredict’ function, ‘ggeffects’ package, fox & weisberg 2019; lüdecke 2018). species detection and by-catch frequency for each event, a mcnemar test (‘mcnemar.test’ function, ‘stats’ package), which compares the proportions of paired data to highlight species for which there were differences in the number of detections between the two camera heights, was independently performed. the authors plotted the type of non-identical events (not triggered, empty, dark), coloured by species to describe the causes behind discrepancies between paired cameras. results the camera trap survey ran over a period of 150 days. camera pairs were collectively active for a total of 595 trapping days and produced 15 623 photographs, corresponding to 987 captures. most of the photographs were taken during the day (70% for high cameras and 62% for low cameras). low and high respectively accounted for 53% (519) and 47% (468) of the captures. these corresponded to 559 unique independent events recorded during the trapping survey. low and high cameras respectively captured 90% and 81% of the events. the high cameras missed 16% of the events (90) compared to 9% for the low cameras (48). the percentage of empty photos was fairly low for both heights, but was twice as low for the low cameras (0.6% for low cameras and 1.8% for high cameras). the most frequently captured species were bushbuck (n = 146) and humans (n = 147). cape grey mongoose (n = 2), vervet monkey (n = 6) and vehicles (n = 5) were the rarest (online appendix table 1-a1). ten wild mammal species were identified from the survey: (1) chacma baboon (papio ursinus), (2) honey badger (mellivora capensis), (3) bushbuck (tragelaphus sylvaticus), (4) bushpig (potamochoerus larvatus), (5) caracal (caracal caracal), (6) south african large-spotted genet (genetta tigrina), (7) cape leopard (panthera pardus), (8) vervet monkey (chlorocebus pygerythrus), (9) cape grey mongoose (galerella pulverulenta), and (10) cape porcupine (hystrix africaeaustralis). general detection model high cameras detected more humans than low cameras (respectively 146 and 124), whereas high cameras detected less wild mammal species events than low cameras (266 and 330 respectively). the best model includes three significant interactions. low cameras were more efficient in detecting smaller species (weight and height) than high cameras (p: 0.00539; online appendix figure 3-a1 and table 2-a1). additionally, lower cameras were more efficient for detecting species at night (p = 0.00940; online appendix table 2-a1), especially smaller species (p: 0.00798; online appendix figure 3-a1 and table 2-a1). the site was not part of the selected variables, so it was assumed that it had no effect on the difference in detection probability between the two camera heights (online appendix table 2-a1). species detection and by-catch frequency the only species not detected by high cameras, the cape grey mongoose, was detected only twice by low cameras. it was found that high cameras were less effective at capturing porcupine (p: 0.014), large-spotted genet (p: 5.35e-13) and honey badger (p: 0.059), compared to low cameras (online appendix table 3-a1 and figure 1). low cameras detected most of the human-related events (i.e. vehicles, motorcycles and dogs with humans) but had a lower detection success for humans (p: 7.10e-06). figure 1: number of independent detections by low (blue) and high (green) cameras for each species photographed in the harkerville forest, garden route national park, western cape, south africa. furthermore, for some species (bushbuck, human, bushpig, large-spotted genet and honey badger), the total sum of detections (in grey on figure 1) was greater than the maximum number of detections for each camera, proving that both cameras missed some images. the non-identical events were mostly due to cameras not being triggered by a by-catch species, which concerned small mammal species for high cameras (e.g. 63% of genets not detected) and humans for low cameras (56% of the non-detected events; figure 2). figure 2: type and number of non-identical events for (a) high camera and (b) low camera, coloured by species and sorted by empty images, not triggered events, and technical issues. discussion lower placed cameras were more effective for capturing small and nocturnal mammals, while higher cameras did marginally better at detecting humans and large nocturnal species. low cameras detected most of the human-related events successfully, but to a lesser extent than the higher cameras, which may be because humans are bipedal and have a higher centre of gravity than quadrupedal species, with the bulk of their mass being higher (alexander 2004; soni et al. 2020). moreover, the high cameras missed the detection of most large-spotted genets and all cape grey mongooses, which were by-catch species. this might be because species walked under the triggering sensor of the camera or that the latter was too far to detect smaller species, which is consistent with other work showing that missed detections were the result of failed triggers that increased as species size decreased (jacobs & ausband 2018). this supports that a setup height which is not adapted to the target species misses most of its detections, mainly because the cameras are not triggered, and not because of the registration or capture quality probability (findlay et al. 2020). as the cameras from both heights were oriented to photograph the same vertical and horizontal area (online appendix figure 2-a1), the non-identical events were mostly due to the blind spot from the high camera (near the ground at the bottom of the tree) or because the individual moved outside of the range of detection of the motion sensor. empty photos were due to species triggering the camera and moving out of the detection zone too quickly to be photographed (findlay et al. 2020). this study concurs with other studies that found a significant effect of the camera height on species detection (findlay et al. 2020; mcintyre et al. 2020; meek et al. 2016b) and that the probability of detection for cameras positioned at different heights depends on the species size-class (palencia et al. 2021). it confirms the necessity to adapt the height and the angle of the camera according to the targeted species or community of species to optimise detection rate (hofmeester et al. 2020; mcintyre et al. 2020; meek et al. 2016b). the results of this study were obtained in a dense forest, which can limit the potential area of movement for the species and the detection range of the cameras, yet they are similar to results found in open landscapes (mcintyre et al. 2020; palencia et al. 2021). failure to take into consideration the effect of camera height on animal detection will lead to poor-quality data, and will likely bias analysis, interpretation and conservation outcomes. the survey focused on a single camera trap model at five sites, all distributed in the same habitat and at the same distance of a trail, in order to test for the effect of camera height on detection. other survey designs that might impact the detection were not taken into consideration. the results showed that cameras set up higher from the ground are not suitable for studying small mammals (under 30 cm and 10 kg); however, the by-catch data can be used for studying human activities and large mammals (above 50 cm or 20 kg). furthermore, the low cameras appeared to be the most relevant for studying the biodiversity of small to medium-sized mammals (mean height = 53.1 cm, height range = 15 cm to 80 cm). using cameras set at a low height is suitable for the study of human and wildlife coexistence, as they performed similarly to high cameras in recording human-related events. the study expands knowledge on how to incorporate by-catch data into camera trap studies by providing a case study comparing human-focused surveys with mammal-focused surveys in forest habitat. camera trap by-catch should not be carelessly used to address a multitude of research objectives. pilot studies are paramount before merging data sets acquired from surveys with differing objectives and camera trap setups. although imperfect detection needs to be accounted for when combining different studies of camera traps (hofmeester et al. 2019), by-catch data can play a crucial role in providing valuable information on management, conservation and environmental processes (edwards et al. 2018; hofmeester et al. 2020). acknowledgements the authors would like to thank south african national parks for allowing this study in the garden route national park, and the rangers who assisted with the service of the cameras. the authors thank jérémy tornos for his help at various stages of the work as well as justin bellengé for the drawings. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions l.m. and h.f. conceived the original idea. a.b., l.m. and m.a.m. carried out the experiment. a.b. wrote the manuscript with support from all the authors. a.b. performed the analysis. h.f. and c.g. provided critical feedback on the analysis. all authors discussed the results and contributed to the final manuscript. ethical considerations this article followed all ethical standards for research without direct contact with human or animal subjects. funding information this study was funded as part of a grant from the ‘idex de l’université de lyon’ and field support from the french national centre for scientific research (cnrs) institute of ecology and environment (inee) allocated to the international research laboratory rehabs. cameras for this experiment were provided by the scientific services from sanparks, garde route national park. data availability the data that support this study will be available from the corresponding author, a.b., upon reasonable request. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references alexander, r.m., 2004, ‘bipedal animals, and their differences from humans’, journal of anatomy 204(5), 321–330. https://doi.org/10.1111/j.0021-8782.2004.00289.x anile, s. & devillard, s., 2016, ‘study design and body mass influence rais from camera trap studies: evidence from the felidae’, animal conservation 19(1), 35–45. https://doi.org/10.1111/acv.12214 apps, p. & mcnutt, j.w., 2018, ‘are camera traps fit for purpose? a rigorous, reproducible and realistic test of camera trap performance’, african journal of ecology 56(4), 710–720. https://doi.org/10.1111/aje.12573 burton, a.c., neilson, e., moreira, d., ladle, a., steenweg, r., fisher, j.t. et al., 2015, ‘review: wildlife camera trapping: a review and recommendations for linking surveys to ecological processes’, journal of applied ecology 52(3), 675–685. https://doi.org/10.1111/1365-2664.12432 child, f.m., roxburgh, l., do linh san, e., raimondo, d. & davies-mostert, h.t., 2016, the red list of mammals of south africa, swaziland and lesotho, south african national biodiversity institute and endangered wildlife trust, south africa, viewed 14 march 2022, from www.ewt.org.za. edwards, m.a., derocher, a.e., hobson, k.a., branigan, m. & nagy, j.a., 2011, ‘fast carnivores and slow herbivores: differential foraging strategies among grizzly bears in the canadian arctic’, oecologia 165(4), 877–889. https://doi.org/10.1007/s00442-010-1869-9 edwards, s., cooper, s., uiseb, k., hayward, m., wachter, b. & melzheimer, j., 2018, ‘making the most of by-catch data: assessing the feasibility of utilising non-target camera trap data for occupancy modelling of a large felid’, african journal of ecology 56(4), 885–894. https://doi.org/10.1111/aje.12511 findlay, m.a., briers, r.a. & white, p.j.c., 2020, ‘component processes of detection probability in camera-trap studies: understanding the occurrence of false-negatives’, mammal research 65(3), 167–180. https://doi.org/10.1007/s13364-020-00478-y fox, j. & weisberg, s., 2019, an {r} companion to applied regression, 3rd edn., sage, thousand oaks, ca. hofmeester, t.r., cromsigt, j.p.g.m., odden, j., andrén, h., kindberg, j. & linnell, j.d.c., 2019, ‘framing pictures: a conceptual framework to identify and correct for biases in detection probability of camera traps enabling multi-species comparison’, ecology and evolution 9(4), 2320–2336. https://doi.org/10.1002/ece3.4878 hofmeester, t.r., young, s., juthberg, s., singh, n.j., widemo, f., andrén, h. et al., 2020, ‘using by-catch data from wildlife surveys to quantify climatic parameters and timing of phenology for plants and animals using camera traps’, remote sensing in ecology and conservation 6(2), 129–140. https://doi.org/10.1002/rse2.136 jacobs, c.e. & ausband, d.e., 2018, ‘an evaluation of camera trap performance – what are we missing and does deployment height matter?’, remote sensing in ecology and conservation 4(4), 352–360. https://doi.org/10.1002/rse2.81 kolowski, j.m., oley, j. & mcshea, w.j., 2021, ‘high-density camera trap grid reveals lack of consistency in detection and capture rates across space and time’, ecosphere 12(2), e03350. https://doi.org/10.1002/ecs2.3350 lüdecke, d., 2018, ‘ggeffects: tidy data frames of marginal effects from regression models’, journal of open source software 3(26), 772. https://doi.org/10.21105/joss.00772 mazzamuto, m.v., lo valvo, m. & anile, s., 2019, ‘the value of by-catch data: how species-specific surveys can serve non-target species’, european journal of wildlife research 65(5), 68. https://doi.org/10.1007/s10344-019-1310-6 mcintyre, t., majelantle, t.l., slip, d.j. & harcourt, r.g., 2020, ‘quantifying imperfect camera-trap detection probabilities: implications for density modelling’, wildlife research 47(2), 177–185. https://doi.org/10.1071/wr19040 meek, p., ballard, g., fleming, p. & falzon, g., 2016a, ‘are we getting the full picture? animal responses to camera traps and implications for predator studies’, ecology and evolution 6(10), 3216–3225. https://doi.org/10.1002/ece3.2111 meek, p.d., ballard, g.a. & falzon, g., 2016b, ‘the higher you go the less you will know: placing camera traps high to avoid theft will affect detection’, remote sensing in ecology and conservation 2(4), 204–211. https://doi.org/10.1002/rse2.28 moolman, l., de morney, m.a., ferreira, s.m., ganswindt, a., poole, j.h. & kerley, g.i.h., 2019, ‘and then there was one: a camera trap survey of the declining population of african elephants in knysna, south africa’, african journal of wildlife research 49(1), 16–26. https://doi.org/10.3957/056.049.0016 palencia, p., vicente, j., soriguer, r.c. & acevedo, p., 2021, ‘towards a best-practices guide for camera trapping: assessing differences among camera trap models and settings under field conditions’, journal of zoology 316(3), 197–208. https://doi.org/10.1111/jzo.12945 pardo, l.e., bombaci, s.p., huebner, s., somers, m.j., fritz, h., downs, c. et al., 2021, ‘snapshot safari: a large-scale collaborative to monitor africa’s remarkable biodiversity’, south african journal of science 117(1/2). https://doi.org/10.17159/sajs.2021/8134 r core team, 2020, r: a language and environment for statistical computing, r foundation for statistical computing, vienna. royle, j.a., 2011, ‘camera trap in ecology’, in a.f. o’connell, j.d. nichols & k.u. karanth (eds.) camera traps in animal ecology, pp. 163–190, springer, tokyo. soni, a., mishra, s., santra, a., khune, v., pathak, r., dubey, a. et al., 2020, ‘position of centre of gravity in different species: a review’, journal of entomology and zoology studies 8(1), 496–499. swann, d.e., hass, c.c., dalton, d.c. & wolf, s.a., 2004, ‘infrared-triggered cameras for detecting wildlife: an evaluation and review’, wildlife society bulletin 32(2), 357–365. https://doi.org/10.2193/0091-7648(2004)32[357:icfdwa]2.0.co;2 tobler, m.w., 2007, camera base version 1.3, viewed 10 march 2021, from http://www.atrium-biodiversity.org/tools/camerabase/. venables, w.n. & ripley, b., 2002, modern applied statistics with s, 4th edn., springer, new york, ny. wickham, h., 2016, ggplot2: elegant graphics for data analysis, springer-verlag, new york, ny. article information authors: natasha nikodinoska1 llewellyn c. foxcroft2,3 mathieu rouget4 alessandro paletto5 sandra notaro6 affiliations: 1department of environmental sciences, parthenope university of naples, italy2south african national parks, conservation services, skukuza, south africa 3centre for invasion biology, department of botany and zoology, stellenbosch university, south africa 4centre for invasion biology, school of agricultural, earth and environmental sciences, university of kwazulu-natal, south africa 5consiglio per la ricerca e la sperimentazione in agricoltura, forest monitoring and planning research unit (cra-mpf), villazzano, trento, italy 6department of economics, university of trento, italy correspondence to: natasha nikodinoska postal address: centro direzionale, isola c4, 80143 napoli, italy dates: received: 18 nov. 2013 accepted: 27 apr. 2014 published: 02 july 2014 republished: 15 july 2014 how to cite this article: nikodinoska, n., foxcroft, l.c., rouget, m., paletto, a. & notaro, s., 2014, ‘tourists’ perceptions and willingness to pay for the control of opuntia stricta invasion in protected areas: a case study from south africa’, koedoe 56(1), art. #1214, 8 pages. http://dx.doi.org/10.4102/ koedoe.v56i1.1214 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/ koedoe.v56i1.1214-1. this article was republished with the full second affiliation for llewellyn c. foxcroft and full affiliation for mathieu rouget. copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. tourists’ perceptions and willingness to pay for the control of opuntia stricta invasion in protected areas: a case study from south africa in this original research... open access • abstract • introduction • material and methods    • study area    • data collection    • analysis • results    • willingness to pay for the control of opuntia stricta    • perceptions of tourists to invasive alien plant species in south africa • discussion    • willingness to pay for the control of opuntia stricta    • perceptions of tourists to invasive alien plant species in south africa • conclusion • acknowledgments    • competing interests    • authors’ contributions • references • appendix 1 • appendix 2 abstract top ↑ invasive alien plants have a long history of establishment in the national parks of south africa. in particular, opuntia stricta (sour prickly pear) has invaded several protected areas in the country, threatening the biodiversity conservation mandate of these conservation areas. this article focuses on the economic estimation of o. stricta’s negative impacts in protected areas by using contingent valuation surveys conducted amongst a sample of tourists in the pilanesberg national park (north west parks and tourism board, south africa). tourists’ familiarity and awareness of selected invasive alien plants and their willingness to pay for the implementation of a control programme for o. stricta were assessed. the results show that many tourists are familiar with invasive alien plants and their (positive and negative) impacts and, in particular, perceived the presence of o. stricta to be negative, due to the impacts on aesthetics and recreation. socio-demographic characteristics, as well as individual attitudes and biocentric beliefs, have an influence on the willingness to contribute financially to a control programme for o. stricta. the individual willingness to pay assessment found that the majority of respondents (78%) were willing to pay a higher entrance fee (an additional r57.30 or $7.00 per day) for a hypothetical programme to control the invasion of o. stricta in the pilanesberg national park.conservation implications: the willingness of tourists to pay for o. stricta management provides useful insights in the decision-making process of park management. the results are encouraging, since, in general, tourists are aware of the problem and are in support of providing additional economic input for preventing future alien plant invasions. introduction top ↑ biophysical aspects of invasive alien plants (iaps) have been relatively well studied over a long period of time (e.g. simberloff, martin & genovesi 2013; vilà et al. 2011), whilst socio-economic aspects linked to alien plant invasions have been analysed only during the last two decades (born, rauschmayer & bräuer 2005; charles & dukes 2007). this was mainly attributed to (1) the inability of the markets to capture the economic value of the damage caused by iaps on many ecosystem services, (2) limited knowledge and experience with iap impacts on many ecosystems and (3) conflicts of interest often associated with iaps (pejchar & money 2009; van wilgen, khan & marais 2011), which can heavily influence iap management and conservation policy. constructive consideration of these challenges is of crucial importance, especially for protected areas (pas) where management needs to ensure biodiversity conservation in the face of serious challenges (emerton, bishop & thomas 2006), such as human-induced pressures, lack of financial resources and competing conservation projects (e.g. animal poaching).several methods for the economic valuation of iap impacts have been employed in the past (garrods & willis 1999; pejchar & money 2009), namely cost-based (production function, replacement cost and avoided damage cost) and preference-based (travel cost method, contingent valuation and choice experiment) methods. amongst the preference-based techniques, contingent valuation (cv) has commonly been used to assess the public’s perception of the impacts of iaps and their willingness to pay (wtp) for control programmes. cv is a survey-based method where respondents directly state their preferences for quantitative or qualitative changes in ecosystem goods or services (alberini & kahn 2006; mitchell & carson 1989). despite the fact that cv has its own limitations (venkatachalam 2004), it is still viewed as a useful way to provide input into the decision-making process, especially when dealing with effects that are not expressed through market signals (bräuer 2003). cv was employed to assess the economic value of potentially eradicating a single iap, such as eichhornia crassipes (law 2008), acacia saligna (lehrer, becker & kurtiel 2013) or a set of iaps (bardsley & jones 2006; turpie 2004; van wilgen et al. 2001). still, few studies have examined both tourists’ perceptions and their wtp for iap control, especially in pas that are directly dependent on tourism for their income. the aim of this article is to assess tourists’ perceptions and knowledge of selected iaps present in south africa (sa) and, in particular, to investigate tourists’ wtp for the control of opuntia stricta in pas, using combined cv and visual aids surveys. opuntia stricta was chosen because it can have severe impacts on pas, without any important potential benefits for pas, or for local communities in or around conservation areas, as the plant is not used for animal forage or human consumption. other representatives of this group, like opuntia ficus-indica, are important food and fodder resources for humans and animals and play an important role in the livelihood of local communities in africa (larsson 2004). opuntia species were abundant in the pilanesberg national park (pnp) in the period from 1960–1984 and were treated mechanically and with herbicides (carruthers 2011). although it was not specified whether o. stricta was included in the species controlled, the plant has not become established; or if previously locally eradicated, has not become re-established and invaded pnp. however, we use this species as an example of a species (and specifically an opuntia spp.) that could become problematic in pnp, as it has in other pas in sa for example, kruger national park (foxcroft et al. 2007) and mapungubwe national park (sanparks 2012). native to south-eastern usa, eastern mexico and cuba, o. stricta (haw.) haw. was introduced worldwide through natural (wind, bird, animal and floodwater) and anthropogenic pathways (monteiro et al. 2005; vilà et al. 2003). the causes for the man-made pathways include intentional introduction for ornamental purposes and as hedge plants (monteiro et al. 2005). opuntia stricta is present across much of the north-eastern region of sa (henderson 2001; rouget et al. 2004), including several pas (kruger national park, mapungubwe national park, camdeboo national park and karongwe private game reserve, amongst others), indicating that even conservation areas are not immune to its invasion (foxcroft et al. 2007; masubelele, foxcroft & milton 2009). opuntia stricta can reach high densities and cause multiple negative ecological and economic impacts, such as the reduction of food production and the loss of grazing potential (jullien, mcfadyen & cullen 2012; menkins 2010); the restriction of human access that impedes people’s ability to travel and move without hindrance, mainly in rural areas (larsson 2004); the reduction of the aesthetic value of a landscape and recreational activities (parsons & cuthbertson 2001); habitat transformation (walters et al. 2011) and changes in biodiversity (robertson et al. 2011). given the above-mentioned impacts and need for control, we estimated the potential economic value of reducing the negative impacts of o. stricta, as perceived by tourists in pas. tourists in pnp were considered as a sample group of visitors of pas to test their wtp in a hypothetical situation of o. stricta invasion in a protected area. firstly, the knowledge and the perceptions of tourists towards iaps in general, and o. stricta specifically, were investigated. secondly, the tourists’ wtp for the implementation of an o. stricta control programme was determined. studies on the outcomes of the working for water programme have shown that the control of iaps results in increased water runoff, and enhances biodiversity (van wilgen et al. 2011). the associated cost–benefit analysis of this programme highlighted the net economic benefits in the long-term. the hypothesis of this study is that such a control programme produces extended benefits by improving the landscape value in pas, enhancing financial support from tourists. we then discuss the advantages and drawbacks of using cv to elicit tourists’ perceptions and their willingness to contribute to a control programme, as well as the potential value of our preliminary results in reducing the knowledge gap in this field. moreover, we discuss the potential contribution of the cv method in quantifying the negative impacts of an iap that do not hold any market price, which could assist policy makers in placing an economic tolerance threshold on the invasiveness of alien plants. material and methods top ↑ study area the pnp is situated in the north west province of sa, and has a total surface area of 57 200 ha. it is set in the crater of an extinct volcano and appears as a complex series of eroded rings of low mountains and hills that rise approximately 300 m – 600 m above the surrounding land (carruthers 2011).the pnp was proclaimed in 1977, but was not considered a ‘natural’ area because of the various anthropogenic activities by many inhabitants in the previous centuries. nevertheless, it was restored ecologically from previous farmlands and converted into a conservation area (national park) where a variety of indigenous animals were reintroduced (anderson 1986). this area is significant for biodiversity conservation because it is a transition zone between the arid savanna and the moist savanna biome (carruthers 2011). the landscape is highly heterogeneous and has many landscapes and habitat types, providing refuge for many species of plant and animal. it also has aesthetic attributes that attract many domestic and foreign tourists. emerging from its recent complex political and management history, the number of visitors has increased steadily from fewer than 50 000 in 1992 to over 500 000 in the most recent records, which place pnp as the leading tourist attraction in north west province (ndabeni et al. 2007). its close proximity to gauteng (less than 3 hs’ drive) favours the presence of day (short-term) visitors from this province. data collection tourists’ perceptions of o. stricta invasion and their wtp for the implementation of a control programme were assessed using a cv survey (alberini & kahn 2006). we pre-tested the cv survey with students doing their master’s degrees in ecology, and visited pnp a week before the formal survey to ask tourists in pnp for their comments. we discussed respondents’ understanding of iaps and their impacts, and obtained feedback on the clarity and flow of the questionnaire content, provision of sufficient information, and time needed to fill in the questionnaire. modifications were made based on the participants’ comments. once finalised, the questionnaire was presented in person (by n. nikodinoska) to a random sample of 61 visitors in pnp in december 2011 (one out of ten visitors sitting in the picnic site was interviewed). the survey lasted approximately 10 min per person. the semi-structured questionnaire comprised 24 closed-form questions (see online appendix 1), subdivided into four parts, following standard procedures described by arrow et al. (1993).part 1 focused on tourists’ motivations, and total costs to visit the park (including tickets, travel, meals and lodging). in part 2, we evaluated tourists’ perceptions about the current status of selected iaps and their knowledge of the negative and positive impacts that these have on ecosystem services, and function, using a 5-point likert scale (1 = very low impact, 2 = low impact, 3 = medium impact, 4 = high impact, 5 = very high impact). the questions on familiarity and knowledge of the selected iaps were important in understanding the quality of information provided in the wtp section (check questions). in the case of the negative impacts, a high value on the likert scale is associated with high concerns amongst the respondents, whereas a high value given for the positive impacts reflects the perception of the historical or current usefulness of iaps for industries or local livelihoods. positive and negative impacts of iaps were selected based on studies of social perceptions of the impacts of iaps in different areas of sa (e.g. bardsley & jones 2006; joubert & mclahren 2002; tessendorf 2007). six of the main genera and invasive species in sa were included: acacia sp. (wattle), pinus sp. (pine tree), lantana camara (lantana), eucalyptus sp. (gum tree), e. crassipes (water hyacinth) and opuntia spp. (the common name ‘prickly pear’ is generally used for most species of platyopuntoid cacti). the negative impacts included in the list were agricultural weeds, damage to livestock, human health, ecosystem functioning, landscape aesthetics and costs of control methods. positive impacts included the use or role of iaps in the forestry sector, ornamental uses, historical uses and landscape aesthetics. part 3 assessed the visual preferences of the presence of o. stricta, using landscape photographs across a gradient of o. stricta abundance (see online appendix 2), as well as the wtp for the implementation of a control programme of o. stricta invasion. the photographs represented four scenarios of o. stricta infestation: about 10%, 20%, 40% and 60% of the total vegetation cover. a ‘business as usual’ scenario is characterised by a percentage of cover less than 5%. thereafter, we presented a hypothetical scenario where o. stricta had invaded pnp and required a control programme to determine the invasion status, select and implement an appropriate control method, and conduct further monitoring. we used the first and the last photograph to represent two invasion scenarios: (1) less than 10% invasion, where a prompt control programme is necessary (i.e. rapid response) and (2) where a control programme is not in place (more than 50% infestation). an open-ended response format for the wtp question was adopted. the daily entrance fee was used as a means of determining tourists’ wtp for the implementation of the control programme, as respondents had already paid an entry fee for pnp, and this could be increased (or remain the same) depending on their response. respondents were first asked to state whether they were prepared to pay a higher fee if they knew that this contribution would go directly to an o. stricta control programme, and then to state their maximum wtp. whilst the surveys were conducted in december 2011, all values presented here are based on 2012 values. the entrance fee for 2012 was r65 for adults and r20 for pensioners and children. part 4 of the questionnaire requested personal information from the respondent (gender, age, level of education, home language and income). analysis a tobit model was fitted to analyse the data and generate a predictive model of wtp (tobin 1958). it is a censored (or truncated) regression model in which the range of the dependent variable (wtp values) is constrained in one way: above or below. in this particular case the censoring was made from below, with the threshold set to zero, meaning that values below zero were constrained. in cases of relatively large numbers of zero values, tobit regression models using maximum likelihood estimation are preferred to linear models using ordinary least squares, as they predict only positive rational wtp values (maddala 1983; tobin 1958). ordinary least squares models are in these cases downward biased. the tobit model is still widely employed in cv surveys that use open-ended wtp questions (e.g. du preez, tessendorf & hosking 2010; halkos & jones 2012; halstead, lindsay & brown 1991).the selection of the variables (see table 1 for variables and definitions) and their expected relationship with the individual wtp per day was based on behavioural theory review and previous studies on tourists’ perceptions of iaps in sa (de wit 2006; tessendorf 2007). the explanatory variables are of both a qualitative and a quantitative nature. qualitative variables are represented by dummy variables, where a value of 1 indicates the presence of the subject (e.g. previous visits to the park) and 0 the absence of the subject, whilst the quantitative variables are expressed by a continuous variable. the expected relationship between the explanatory variables and the wtp is positive if the monetary value increases in response to higher values of the predictive variable (e.g. respondents with a higher income are expected to state higher wtp values). the expected effect is negative if the amount increases in response to lower values of the predictive variables (e.g. tourists who give lower values for photographs with major infestations are expected to state high wtp amounts). table 1: explanatory variables used in the tobit model and their theoretical (expected) relationship with the individual willingness to pay. results top ↑ willingness to pay for the control of opuntia stricta the results from the cv show that 78% of the respondents were willing to pay additional fees in order to control the spread of o. stricta (table 2). the value of wtp for these respondents ranged from r5 – r150 per day. about one third of the respondents who were willing to pay gave r100 as their wtp amount, 17% indicated r20 and 13% stated r10 as their wtp value. table 2: willingness to pay values (2012 values). of the remaining responses, 13 tourists (22% of the respondents) refused to pay higher fees, and 12 had valid wtp zero values (i.e. tourist responses showed that they did not perceive invasion as a problem, therefore were not willing to pay). the reasons for valid zero wtp were mainly that the respondents perceived o. stricta and iaps in general as a way of enhancing species diversity (three respondents), they were not interested in plants (one respondent), and they have become familiar with seeing them in the natural environment (one respondent). there was one protest response that stated his refusal to pay a higher price as a consequence of already high fees in the park. of the respondents who were willing to pay higher fees, two did not quantify the amount they were willing to pay, and responded with ‘not sure’ and ‘reasonable amount’. one respondent did not provide his wtp. in this survey the ‘i don’t know’-type answers were excluded from the wtp analysis. the tobit model selection was based on the bayesian information criterion value, where a lower value indicates a better fitting model. the variables in the tobit model provide a relatively good explanation of wtp (table 3). the mean daily wtp estimated by the tobit model was found to be r57.30 (± 28.80 standard deviation [s.d.]). the positive value of the coefficient for the variable ‘previous visits in south african parks’ shows that respondents who had previously visited parks are willing to pay a higher entrance fee for an iap control programme. similarly, the coefficient for the variable ‘conservation’ (16.747) indicates that visitors who rated the conservation of the environment and typical landscape in pas highly are willing to pay more. the respondents who perceived the photograph that represents the scenario of higher infestation levels (30% – 50% density) as unpleasant are willing to pay more in order to prevent the possible spread of o. stricta (figure 3, in online appendix 2). foreign respondents have a higher wtp for controlling iaps than domestic respondents (variable ‘nationality’). tourists who come from gauteng are also prepared to pay more than visitors from other regions (variable ‘province’). personal income, not surprisingly, influences the respondents’ wtp; namely, respondents with higher incomes are willing to pay a higher entrance fee (variable ‘income’). the average annual income of the interviewed tourists is relatively high, with 26% earning more than r385 711 and 11% earning less than r55 100 per year (26% did not respond; table 4). the rest of the tourists’ socio-demographics are included in table 5. table 3: tobit model. table 4: categories of respondents’ income per year. table 5: socio-demographics of tourists in the pilanesberg national park. perceptions of tourists to invasive alien plant species in south africa tourists’ knowledge of iaps was relatively high, with 75% of respondents having previously heard of the six invasive species (figure 1). where the respondents were asked to select the iaps (of the given options) they know or have heard about, acacia and pinus (pine) spp. were the best known, followed by eucalyptus sp. and opuntia spp. when asked which species they could recognise when they see them, opuntia spp., together with pinus sp., were listed most frequently. eichhornia crassipes was indicated as a problematic species in sa by 27% of the respondents, followed by acacia sp. (23%) and pinus sp. (22%). opuntia species (generally prickly pear) and acacia sp. (wattle) were indicated as problematic species within pas (figure 1). with the ‘other’ option where respondents could list other species they were familiar with, cereus jamacaru (queen of the night) and chromolaena odorata (chromolaena) were frequently identified as problematic iaps in sa. figure 1: tourists’ perceptions of the negative impacts of invasive alien plants in south africa and in protected areas in general. in assessing the impacts from the list of benefits of iaps in sa (table 6), relatively high positive impact values were attributed to forestry uses (mean of 3.4), and pastoral and ornamental uses (3.03). historical uses as well as landscape aesthetics were comparatively less ‘valued’ and with more variation in the responses. table 6: mean value and standard deviation of positive and negative impacts of invasive alien plants in south africa. all listed negative impacts of the iaps had an average value higher than 3 (out of 5), which means that respondents recognised the serious potential treats that invasions can cause. the highest negative values, 4.39 and 4.26 (out of 5), are related to the potential reduced performance of ecosystem functioning and increased costs for implementation of control programmes, respectively. discussion top ↑ willingness to pay for the control of opuntia stricta this survey design included landscape visual aids to better explain the hypothetical scenario. we believe that the use of photographs in cv surveys helps in the design of valuation scenarios, because landscape changes appear more understandable or visual to respondents (madureira et al. 2011). tourists’ perception of the aesthetic value of o. stricta control is therefore easier to convey using visual aids. a possible drawback of using these photographs could be due to the fourth photograph (see online appendix 2) representing an o. stricta invasion of more than 50%, in the description of the hypothetical scenario, which could have partially influenced some of the wtp positive values. whilst this may not be a common situation, it is not unfeasible. nevertheless, the outcomes of the photograph preference have shown that people attribute negative values to the o. stricta invasion, starting from the second photograph (11% – 30% invasion). moreover, the outcome from the tobit model showed that people who expressed their displeasure for the third photograph (30% – 50% invasion) were willing to pay more for the control programme.one of the advantages of using an open-ended wtp question format is that the use of data sets are smaller than required by, for example, a dichotomous choice, which reduces the time and expense of the survey process (halstead et al. 1991). this approach can, however, provide a lower (conservative) wtp value than other methods. critics of this format assert that it can create a large number of non-responses, protest bids and outliers (carson, flores & meade 2001; hanemann 1994), but is efficient if respondents are familiar with paying for the subject of the survey and does not result in a starting point bias (mitchell & carson 1989). in our case study, the respondents demonstrated their awareness of the problem of iaps and thus we are confident in the validity of our results. the similarity of the tourists’ profile between pnp and other national pas in sa (sanparks 2012; van der merwe & saayman 2008) gives relatively higher validity to some of the socio-demographics features that were used as explanatory variables in the tobit model. in particular, we focused on the variable ‘income’ as an important predictor of the stated wtp. the average income of the respondents is relatively high, with 15% earning around r220 400 and another 15% earning more than r385 711 per year. this could be due to the tourists’ profile that (1) a large percentage comes from gauteng, widely known as the most developed south african province and (2) eco-tourists are often people with a middle-high income (statistics south africa 2013). however, a non-negligible number of respondents (approximately 26%) refused to answer this question, which could have altered this outcome. the individual daily wtp to finance a control programme in pnp is, on average, an additional r57.30, which is a substantial amount when considering the entrance fee for an adult is r65.00. the fact that 78% of the visitors are prepared to support an o. stricta control programme, of which a third are willing to pay r100.00 or more, strongly suggests that visitors to national parks are aware of the threat, and the seriousness, posed by iaps. the individual wtp lies within a range of values obtained for another iaps control programme in sa (du plessis 2003; du preez et al. 2010). a study on the wtp for the removal of alien vegetation and restoration of indigenous vegetation in several sites (including conservation areas) in underberg (kwazulu-natal province, sa), by the working for water programme, derived a wtp value of r26.40 (r36.30 in 2012 values) per project, and r27.34 (r37.30 in 2012 values) per hectare (du preez et al. 2010). du plessis (2003) estimated the individual average wtp for the removal of iaps and biodiversity conservation in several sites in the eastern and western cape provinces (nature reserves included) using a cv survey conducted amongst both residents and tourists. the mean individual wtp was r111.54 (r153.57 in 2012 values). our estimation of the wtp appears to be relatively low, which could be explained by the fact that we investigated tourists’ preferences for a control programme implemented for just one iap species. there is still a possibility that respondents overstated their wtp and that if o. stricta was to be included as only one of many iaps to be controlled, the wtp amount could have been even lower (garrods & willis 1999). another constraint is that due to the limited time available, there were fewer questionnaires than would be ideal; however, with the similarities in visitor profiles to other pas, the spread of questionnaires across the range of tourists is acceptable. perceptions of tourists to invasive alien plant species in south africa the respondents rated the impact to ecosystem function and the expense of control methods as the highest concern, which is similar to the results found in a study in the mediterranean region (bardsley & jones 2006). these were followed by the impacts to landscape aesthetics and by agricultural weeds. however, another study conducted in spain shows that the changes in ecosystem stability is considered the third greatest impact of alien plants, after competition with native species and the effects on fauna (andreu, vilà & hulme 2009). the results show that the impacts of iaps on ecosystems and management control programmes are broadly recognised by the respondents.the benefits of iaps that were ranked highly by the respondents included use of species for forestry (commercial plantation forestry), pastoral uses, food, ornamental uses and historical uses. greater importance placed on iaps for forestry uses is perhaps related to the fact that there are still large areas of commercial plantations in sa (e.g. eucalyptus and pinus spp.). in the context of national parks, on the other hand, trees and woody shrubs (acacia spp., pinus spp. and l. camara) as well as cactaceae (opuntia spp.) were most frequently considered to have negative impacts. conclusion top ↑ reducing the knowledge gap in the public perception of iap impacts by providing quantitative data that could be integrated with other research in pas can contribute to the scientific understanding on the economic valuation of iap control programmes. the outcomes of studies on people’s perceptions of both positive and negative impacts of iaps in sa can play a role in improving the development and implementation of conservation projects, especially in cases where ‘conflict’ alien species are considered. moreover, it can also provide insight into the long-term potential of visitor-funded iap control programmes as a feasible option. the benefits of a control programme could be over-estimated if it has not been established whether the beneficiaries (i.e. tourists and local community) recognise the problem and are willing to contribute to the restoration of the invaded areas. acknowledgments top ↑ dr llewellyn c. foxcroft acknowledges the support from the south african national parks, the centre for invasion biology, stellenbosch university and the national research fund (sa) incentives fund. we also thank marc hockings and the anonymous reviewers for their constructive comments that helped improve this article. competing interests the authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article. authors’ contributions the authors contributed equally to this work. references top ↑ alberini, a. & kahn, j.r. (eds.), 2006, handbook on contingent valuation, edward elgar publishing, london. http://dx.doi.org/10.4337/9781845427917anderson, j.l., 1986, ‘restoring a wilderness: the reintroduction of wildlife to an african national park’, international zoo yearbook 24(1), 192–199. http://dx.doi.org/10.1111/j.1748-1090.1985.tb02538.x andreu, j., vilà, m. & hulme, p.e., 2009, ‘an assessment of stakeholder perceptions and management of noxious alien plants in spain’, environmental management 43, 1244–1255. http://dx.doi.org/10.1007/s00267-009-9280-1 arrow, k., solow, r., portney, p., leamer, e., radner, r. & schuman, h., 1993, ‘contingent valuation methodology report’, report of the noaa panel on contingent valuation. fed. reg. 58, resources for the future, washington, pp. 4602–4614. bardsley, d. & jones, e.g., 2006, ‘stakeholders perception of the impacts of invasive exotic plants species in the mediterranean basin’, geojournal 65, 199–210. http://dx.doi.org/10.1007/s10708-005-2755-6 born, w., rauschmayer, f. & bräuer, i., 2005, ‘economic valuation on biological invasions – a survey’, ecological economics 55(3), 321–336. http://dx.doi.org/10.1016/j.ecolecon.2005.08.014 bräuer, i., 2003, ‘money as an indicator: to make use of economic evaluation for biodiversity conservation’, agriculture, ecosystems and environment 98, 483–491. http://dx.doi.org/10.1016/s0167-8809(03)00107-5 carruthers, j., 2011, ‘pilanesberg national park, north west province, south africa: uniting economic development with ecological design – a history, 1960s to 1984’, koedoe 53(1), 1–10. carson, r.t., flores, n.e. & meade, n.r., 2001, ‘contingent valuation: controversies and evidence’, environmental and resource economics 19, 173–210. http://dx.doi.org/10.1023/a:1011128332243 charles, h. & dukes, j.s., 2007, ‘impacts of invasive species on ecosystem services’, in w. nentwig (ed.), biological invasions, vol. 193, pp. 217–237, springer-verlag berlin heidelberg, new york. de wit, m.p., 2006, ‘the value of biodiversity to the south african economy: a preliminary study’, report prepared for south african national biodiversity institute, project ‘development of the sanbi business case’. du plessis, l.l., 2003, ‘an assessment of selected non-water benefits of the working for water programme in the eastern and southern cape’, master’s thesis, faculty of business and economic sciences, nelson mandela metropolitan university. du preez, m., tessendorf, s. & hosking, s.g., 2010, ‘application of the contingent valuation method to estimate the willingness-to-pay for restoring indigenous vegetation in underberg, kwazulu-natal, south africa’, south african journal of economic and management sciences 13(2), 135–157. emerton, l., bishop, j. & thomas, l., 2006, sustainable financing of protected areas: a global review of challenges and options, iucn, gland, switzerland and cambridge, uk. foxcroft, l.c., rouget, m., richardson, d.m. & macfadyen, s., 2007, ‘reconstructing 50 years of opuntia stricta invasion in the kruger national park, south africa: environmental determinants and propagule pressure’, diversity and distributions 10, 427–437. http://dx.doi.org/10.1111/j.1366-9516.2004.00117.x garrods, g. & willis, k., 1999, economic valuation of the environment: methods and case studies, edward elgar, cheltenham. halkos, g.e. & jones, n., 2012, ‘modeling the effect of social factors on improving biodiversity protection’, ecological economics 78, 90–99. http://dx.doi.org/10.1016/j.ecolecon.2012.04.003 halstead, j.m., lindsay, b.e. & brown, c.m., 1991, ‘use of tobit model in contingent valuation: experimental evidence from the pemigewasset wilderness areas’, journal of environmental management 33, 79–89. http://dx.doi.org/10.1016/s0301-4797(05)80049-0 hanemann, w.m., 1994, ‘valuing the environment through contingent valuation’, journal of economic perspective 8, 19–43. http://dx.doi.org/10.1257/jep.8.4.19 henderson, l., 2001, alien weeds and invasive plants: a complete guide to declared weeds and invaders in south africa, plant protection research institute handbook no. 12., agricultural research council, pretoria. joubert, p. & mclahren, l., 2002, ‘awareness on alien vegetation in kruger national park’, sanparks (unpublished data). jullien, m., mcfadyen, r. & cullen, j., (eds.), 2012, biological control of weeds in australia, csiro publishing, melbourne, pp. 351–364. larsson, p., 2004, ‘introduced opuntia spp. in madagascar, problems and opportunities’, swedish university of agricultural sciences, slu/repro, uppsala. law, m.c., 2008, ‘willingness to pay for the control of water hyacinth in an urban environment of south africa’, master’s thesis, rhodes university. lehrer, d., becker, n. & kurtiel, p., 2013, ‘the value of coastal sand dunes as a measure to plan an optimal policy for invasive plant species: the case of the acacia saligna at the nizzanim lter coastal sand dune nature reserve, israel’, springer series on environmental management, 273–288. maddala, g.s., 1983, limited-dependent and qualitative variables in econometrics, cambridge university press, cambridge. http://dx.doi.org/10.1017/cbo9780511810176 madureira, l., nunes, l.c., borges, j.g. & falcão, a.o., 2011, ‘assessing forest management strategies using a contingent valuation approach and advanced visualisation techniques: a portuguese case study’, journal of forest economics 17, 399–414. http://dx.doi.org/10.1016/j.jfe.2011.04.001 masubelele, m.l., foxcroft, l.c. & milton, s., 2009, ‘alien plant species list and distribution for cambedoo national park, eastern cape province, south africa’, koedoe 51, 1–16. menkins, i., 2010, ‘a reassessment of the type of opuntia stricta (haw.) haw.’, viewed 20 september 2013, from http://opuntiads.com/zpdf/a_reassessment_of_the_type_of_opuntia_stricta.pdf mitchell, r.c. & carson, r.t., 1989, using surveys to value public goods: the contingent valuation method, resources for the future, washington. monteiro, a., cheia, v.m., vasconcelos, t. & moreira, i., 2005, ‘management of the invasive species opuntia stricta in a botanical reserve in portugal’, weed research 45(3), 193–201. http://dx.doi.org/10.1111/j.1365-3180.2005.00453.x ndabeni, c., shroyer, m., boonzaaier, w., mokgoko, g. & mochin, s., 2007, ‘the heritage park model: a partnership approach to park expansion in poor rural areas’, in a. watson, j. sproull & l. dean (eds.), science and stewardship to protect and sustain wilderness values: eight world wilderness congress symposium, proceedings rmrs-p-49, n.p., department of agriculture, forest service, rocky mountain research station, us. parsons, w.t. & cuthbertson, e.g., 2001, noxious weeds of australia, 2nd edn., csiro publishing, victoria, australia. pejchar, l. & money, h., 2009, ‘invasive species, ecosystem services and human well-being’, trends in ecology and evolution 24(9), 497–504. http://dx.doi.org/10.1016/j.tree.2009.03.016 robertson, m.p., harris, k.r., coetzee, j., foxcroft, l.c., dippenaar-schoeman, a.s. & van rensburg, b.j., 2011, ‘assessing local scale impacts of opuntia stricta (cactaceae) invasion on beetle and spider diversity in kruger national park, south africa’, african zoology 46(2), 205–223. http://dx.doi.org/10.3377/004.046.0202 rouget, m., richardson, d.m., nel, j.a., le maitre, d.c. & egoh, b.t., 2004, ‘mapping the potential ranges of major plant invaders in south africa, lesotho and swaziland using climatic suitability’, diversity and distributions 10, 475–484. http://dx.doi.org/10.1111/j.1366-9516.2004.00118.x south african national parks (sanparks), 2012, annual report 2011/2012, viewed n.d., from www.sanparks.org/about/annual south african national parks (sanparks), 2012, mapungubwe national park and world heritage site management plan, viewed n.d., from http://www.sanparks.org.za/conservation/park_man/approved_plans.php statistics south africa, 2013, domestic tourism survey 2012. statistical release, viewed n.d., from http://www.datafirst.uct.ac.za/dataportal/index.php/catalog/440/download/5677 simberloff, d., martin, j.l. & genovesi, p., 2013, ‘impacts of biological invasions: what’s what and the way forward’, trends in ecology and evolution 28, 58–66. http://dx.doi.org/10.1016/j.tree.2012.07.013 tessendorf, s.e., 2007, ‘estimating the willingness-to-pay for restoring indigenous vegetation at selected sites in south africa’, master’s thesis, nelson mandela metropolitan university. tobin, j., 1958, ‘estimation of relationships for limited dependent variables’, econometrica 26(1), 24–36. http://dx.doi.org/10.2307/1907382 turpie, j., 2004, ‘the role of resource economics in the control of invasive alien plants in south africa: working for water’, south african journal of science 100(1 & 2), 87. van der merwe, p. & saymaan, m., 2008, ‘travel motivations of tourists visiting kruger national park’, koedoe 50, 154–159. http://dx.doi.org/10.4102/koedoe.v50i1.140 van wilgen, b.w., khan, a. & marais, c., 2011, ‘changing perspectives on managing biological invasions: insights from south africa and the working for water programme’, in d.m. richardson (ed.), fifty years of invasion biology: the legacy of charles elton, pp. 377–393, blackwell publishing ltd, uk. van wilgen, b.w., richardson, d.m., le maitre, d.c, marais, c. & magadlela, d., 2001, ‘the economic consequences of alien plant invasion: examples of impacts and approaches to sustainable management in south africa’, environment, development and sustainability 3, 145–168. http://dx.doi.org/10.1023/a:1011668417953 venkatachalam, l., 2004, ‘the contingent valuation method: a review’, environmental impact assessment review 24(1), 89–124. http://dx.doi.org/10.1016/s0195-9255(03)00138-0 vilà, m., burriel, j.a., pina, j., chamizo, j., llach, e., porterias, m. et al., 2003, ‘association between opuntia species invasion and changes in land-cover in the mediterranean region’, global change biology 9(8), 1234–1239. http://dx.doi.org/10.1046/j.1365-2486.2003.00652.x vilà, m., espinar, j.l., hejda, m., hulme, p.e., jarošík, v., maron, j.l. et al., 2011, ‘ecological impacts of invasive alien plants: a meta-analysis of their effects on species, communities and ecosystems’, ecological letters 14, 702–708. http://dx.doi.org/10.1111/j.1461-0248.2011.01628.x walters, m., figueiredo, e., crouch, n.r., winter, p.j.d., smith, g.f., zimmermann, h.g. et al., 2011, ‘naturalised and invasive succulents of southern africa’, abc taxa 11, 1–10. appendix 1 top ↑ questionnaire on invasive alien plants in national parksintroduction i’m a student enrolled in a master’s course in environmental and tourism economics at the university of trento, italy. the questionnaire is part of the research for my thesis in a cost–benefit analysis of clearing invasive alien plants in national parks, which i’m currently conducting at the university of pretoria. the completion of this questionnaire will take approximately 10 minutes. if you are unable to provide me with exact details, please answer as precisely and correctly as possible. part 1 1. have you visited any parks in south africa? 2. if yes, list the parks that you’ve visited __________________________ __________________________ 3. what is your main reason for visiting the park? (the answer can be more than one) a. to see animals b. to see plants c. to see endangered wildlife d. to take photographs of wildlife and landscapes e. educational reasons f. other, specify_________________________________ 4. how many days will you spend in this park? 5. could you give an estimate of the total cost (including travel, meals and lodging, visits) for the holiday in this park? 6. did you book any of these services? if yes, please select the appropriate service. otherwise, select ‘no’ and go to question number 8. a. lodge (suite, cottage) b. chalet c. safari tent d. tour and safari e. other services ___________________ f. no 7. if yes, could you please give the amount that you have paid for the service? 8. how important is the conservation of the environment and the typical landscape within protected areas? (1 = not at all, 5 = very important). we would like to remind you that there isn’t either correct or incorrect answer. 9. are you member of an environmental group? (e.g. wwf, greenpeace, etc.) 10. have you heard about invasive alien plants? if no, go directly to part 2 of the questionnaire. 11. below is a list of plants. in the first column please tick the plants that you know or have heard of. in the second column, tick the plants that you can recognise when you see them. in the third column, tick the plants that you think are a problem in south africa. in the fourth column, tick the ones that you think are a problem in national parks. people have different opinions and information about invasive alien plants. please use the scale from 1 to 5 to respond the questions below (1 = not at all important, 5 = very important). we would like to remind you that there isn’t either a correct or incorrect answer. 12. what are the negative impacts of invasive alien plants? please answer all the questions. 13. what are the positive impacts of invasive alien plants? please answer all the questions. part 2 14. the following four photographs indicate the possible state of invasion of prickly pear within a park. express your preferences related to each of these photographs using the scale from -2 to 2 (-2 = not pleasant at all, 0 = indifferent, 2 = very pleasant). 15. please select the entrance fee that you have paid in this park. would you be willing to pay a higher entrance fee in order to contribute to the prevention of the spread of sour prickly pear in the park (through operations such as detection, research, eradication and monitoring)? if yes, how much more would you be willing to pay to avoid opuntia stricta from spreading in the park? ___________________ 16. could you please give reasons why you are willing to pay a higher entrance fee? ___________________ could you please give reasons why you are willing to pay the same entrance fee? _______________________________________________________________________________________ general information 17. gender 18. age ____________ 19. nationality ______ 20. if south african, please answer the following questions: 21. number of people in your family who have paid the entrance fee of r65. 22. number of pensioners and children in your family who have paid the entrance fee. 23. qualification • school • graduate degree • post graduate – honours • doctorate degree 24. please indicate if you are currently employed in one of these sectors. if not, select ‘other’ and specify the sector or indicate your position as student, pensioner or unemployed. • agriculture • forest industry • tourism • conservation • education • other (please specify which sector __________________) • student • pensioner • unemployed 25. income per year (€1 = r11.02; €1= $1.35). choose one of the following categories: appendix 2 top ↑ figure 1: invasion with less than 10% cover. figure 2: invasion with between 11% and 30% cover. figure 3: invasion with between 31% and 50% cover. figure 4: invasion with more than 50% cover. note: this is the online appendix of nikodinoska, n., foxcroft, l.c., rouget, m., paletto, a. & notaro, s., 2014, ‘tourists’ perceptions and willingness to pay for the control of opuntia stricta invasion in protected areas: a case study from south africa’, koedoe 56(1), art. #1214, 8 pages. http://dx.doi.org/10.4102/koedoe.v56i1.1214 abstract introduction materials and methods results and discussion conclusion acknowledgements references appendix 1 about the author(s) daniel m. parker school of biology and environmental sciences, faculty of agriculture and natural sciences, university of mpumalanga, mbombela, south africawildlife and reserve management research group, department of zoology and entomology, rhodes university, makhanda, south africa citation parker, d.m., 2021, ‘mammals in the mountains: an historical review and updated checklist of the mammals of the mountain zebra national park’, koedoe 63(1), a1683. https://doi.org/10.4102/koedoe.v63i1.1683 original research mammals in the mountains: an historical review and updated checklist of the mammals of the mountain zebra national park daniel m. parker received: 30 apr. 2021; accepted: 26 may 2021; published: 12 july 2021 copyright: © 2021. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract based on published and unpublished records, together with original data collected from regular field trips over a 15-year period, 68 mammal species have been reliably recorded from the mountain zebra national park. i assessed the current status of all mammal species, in relation to park expansion and research effort over time (1937–2020). although numerous large and charismatic mammal species have been reintroduced to the park since it was gazetted in 1937, both in an attempt to restore the historical diversity of the region and to attract tourists, research effort in the surveying of the smaller and more cryptic mammal species has been sorely lacking. i recommend that future survey work targets the small, mostly fossorial mammals (i.e. golden and rodent moles, elephant shrews and gerbils) and insectivorous bats. conservation implications: this work provides critical presence data for several mammal species from an important protected area that straddles three biomes in south africa. keywords: species richness; biodiversity; surveys; time-series; south africa; semi-arid. introduction south africa boasts a network of 20 national parks that are situated across a range of vegetation biomes. the primary function of these national parks is to protect the ecological integrity of these various natural ecosystems for current and future generations (bezuidenhout & brown 2008). however, several national parks within this network were originally gazetted in an attempt to preserve a single species (often large mammals) because of pressure from over-hunting or poaching (e.g. addo elephant national park and bontebok national park). the mountain zebra national park (hereafter mznp) is one such ‘single-species’ national park. the park was founded in 1937 to protect the endangered cape mountain zebra (equus zebra zebra). to sustain the viability of the mountain zebra population, the park was extended in 1964 and 1996 by incorporating various farms adjacent to the park (bezuidenhout & brown 2008). the park was expanded again in 2002, but by this stage, the conservation and management focus had shifted more towards the conservation of biodiversity as a whole, rather than just the viability of the mountain zebra population (sanparks 2016). although the mznp has been the focus of much mammalian-related research since it was proclaimed, the park has expanded to 12 times its original size since 1937, and the current checklist of mammals appears to be a transcript of several earlier (flawed) lists. in addition, since the proclamation of the park, several technological advancements (e.g. the introduction of passive infra-red camera traps) are likely to have improved the detectability of some species that other more traditional techniques failed to detect in the past (de bondi et al. 2010). moreover, the nomenclature (and, effectively, the number) of individual mammal species may change over time as new knowledge about the evolutionary history of a species or group of species becomes available and they are either divided or synonymised (taylor et al. 2019). thus, the aims of this study were to provide an historical review of the mammal species of the mznp and to update the mammal checklist for the park. materials and methods study site the mznp (32°18’s, 25°24’e) is a south african national park (sanparks) and initially included only the farm ‘babylons toren’ which was just 1712 ha in size (grobler & hall-martin 1982). in 1964, the park increased to 6536 ha with the acquisition of the farms ‘zebrahoek’, ‘pretoriuskraal’, ‘wildepaardenek’, ‘sneeuberg’ and a portion of ‘doornhoek’ (grobler & hall-martin 1982). the park stayed at this size until 1996 when the farms ‘ingleside’, ‘welgedacht’, ‘de rust’, ‘ebenhaeser’, ‘jurisdam’, ‘zeekoeigat’ and the remaining portion of ‘doornhoek’ were acquired, taking the park to 18 000 ha (craig, hulley & parker 2005). these newly acquired farms only became available for use by the non-volant animals residing in the park in the early 2000s when they were adequately fenced into the original 6536 ha portion. in approximately 2002, the farms ‘toekoms’ (also referred to as ‘sonnenrust’) on the western boundary and portions of farm numbers 595 and 596 (previously owned by wp erasmus) on the southern tip were incorporated into the park, taking it to 21 412 ha (figure 1, sanparks 2016). although the park is now officially 28 386 ha in extent, the additional areas are not located directly adjacent to the existing park and have not yet been gazetted as national park land (sanparks 2016). figure 1: the current (2020) extent of the mountain zebra national park in the eastern cape, south africa. the park is situated in a transition zone between the nama-karoo, grassland and albany thicket biomes and is characterised by a semi-arid climate (mucina et al. 2006). the southern section of the park is mountainous with altitudinal peaks of up to 1960 m, whereas the northern section consists of lower lying areas ranging from 1000 m to 1500 m. historical mammal records and updating of the mammal checklist mammal records for the mznp since its proclamation were collated from (1) relevant published articles, (2) relevant unpublished reports and student theses, (3) unpublished, annual and individual field trip reports submitted to sanparks for the period 2001–2016 under approved projects 2000-11-08rber and bisc864 and (4) direct observations during field trips not captured under (3). for published articles, a google scholar search was conducted using the search term ‘mountain zebra national park’. this search produced 1200 potential articles which i then screened to determine their relevance. if an article’s title alluded to the manuscript including records for mammals, i read the abstract (where applicable) to ascertain if such records were in fact provided. in cases where mammal records were included in the manuscript, these data were extracted (table 1). the articles that included mammal records then had their reference lists perused to determine the relevance of any citation which was not included in the list of articles from the original google scholar search. in most instances, such citations were unpublished internal reports or student theses. wherever possible, i attempted to obtain copies of these reports and or theses. table 1: the mammal species recorded at mountain zebra national park between 1937 and 2020. for unpublished reports, student theses and annual and field trip reports, relevant sections (including appendices) were checked and any reliable mammal data were extracted (table 1). for the unpublished or grey literature, i deemed mammal presence data reliable only when it was supported by firm evidence (i.e. a specimen had been collected, the animal had been reliably observed by the authors or photographic evidence was provided). the extracted data were then organised taxonomically and chronologically to match the four separate periods of park expansion described above (table 1). only in instances where a species had apparently not been recorded during a certain period, but was known to be present in the park, were direct observations used. typically, the inclusion of such direct observations was for the more charismatic and larger species or those that could not be easily confused with any other species for the period 1999–2002. for example, published and unpublished data demonstrated that porcupines (hystrix africaeaustralis) were present in the park from 1937 to 1998 but only recorded again between 2003 and 2020, leaving the period 1999–2002 without a confirmed record. however, field trips to mznp by the rhodes university department of zoology and entomology began in 2001, and porcupines were sighted fairly regularly on night drives in 2001 and 2002. thus, porcupines and nine other species (aardvark orycteropus afer, springhare pedetes capensis, ground squirrel xerus inauris, chacma baboon papio ursinus, vervet monkey chlorocebus pygerythrus, aardwolf proteles cristata, meerkat suricata suricatta, yellow mongoose cynictis penicillata and cape grey mongoose galerella pulverulenta) were recorded as being present between 1999 and 2002. i was confident that these 10 species could not be confused with any other mammal species. to update and revise the checklist of mammals for mznp, i assessed the reliability of the time-series data generated using the methods described above (table 1). i adopted a conservative approach and only included a species on the revised checklist if a specimen had been reliably collected or observed or if there was photographic evidence of its presence in the park. importantly, during the initial 27 year existence of the park (1937–1964), when the park was just 1712 ha in size, the only published account of the mammals that might be present was published by the renowned naturalist jack skead (1958). however, this publication is essentially a ‘best guess’ estimate of the mammals that could be found in the cradock district as a whole and was not specific to the park (skead 1958). whilst skead (1958) does rely on museum specimens to a limited extent, many of the species included in his publication were based on ‘….distributional evidence from contiguous districts’ and, in some cases, the diaries and notes of early european settlers. although such data are valuable from an historical perspective, their reliability can be questionable, especially for non-charismatic and smaller species (bernard & parker 2006). nevertheless, for the sake of completeness, all of skead’s (1958) proposed records (except species which he deemed to be extinct at the time) were initially included in the generation of the time-series data (table 1). however, if these records could not be corroborated by subsequent (reliable) records, they were not included in the revised checklist (appendix 1). results and discussion in its first 27 years of existence, it was estimated that a total of 60 mammalian species could have been present in mznp (table 1). between 1965 and 1998, with the addition of several dedicated mammal surveys, this number increased to 62 (table 1). however, the 1999 to 2002 period saw the number of mammals supposedly present in the park decline to just 35 (table 1). however, this is likely an artefact of a lack of sampling and/or documented fieldwork in the park during this period as the number of mammal species present in the park increased to 63 between 2003 and 2020 (table 1) when dedicated fieldwork was being conducted by the rhodes university department of zoology and entomology. interestingly, despite the park increasing more than 12 times in size between 1965 and 2020, the overall number of mammal species present has stayed remarkably stable (figure 2). however, the actual species composition present has changed substantially (table 1). for example, many of the larger ungulate species were re-introduced and several surveys of the smaller mammals revealed previously unrecorded species (table 1). the advent of passive infra-red camera traps to photograph medium and large species between 2003 and 2020 also likely contributed to the observed changes. figure 2: the four periods of park expansion for the mountain zebra national park, eastern cape, south africa between 1937 and 2020. the total number of mammal species recorded during each period (grey bars) is also shown. the solid line denotes the cumulative increase in park area (hectares) over time. the revised and updated checklist for the park includes 68 species from 25 families representing 13 orders (appendix 1). with 14 species present in the park, the bovidae is the best represented family, followed by the murid rodents (11 species; appendix 1). as a group, the carnivores (hyaenidae, felidae, viverridae, canidae and mustelidae) are also fairly well represented (20 species) in the park (appendix 1). all other families present in the park currently have less than three species represented (appendix 1). many of these remaining families represent the small, and arguably understudied, mammal groups (appendix 1). i consider these 68 species to be the best current estimate of the mammal richness of the park. the species which i rejected from the final checklist are improbable and likely based on outdated distributional and ecological data or were originally mis-identified. i briefly discuss the 18 rejected species below. rejected species bushveld elephant shrew (elephantulus intufi) the inclusion of this species was based on what was believed to be the most plausible distribution at the time (skead 1958). as no additional specimens have been collected, or verifiable sightings made since its initial inclusion, i rejected it from the updated checklist of mznp. round-eared elephant shrew (macroscelides proboscideus) the inclusion of this species was based on what was believed to be the most plausible distribution at the time (skead 1958). as no additional specimens have been collected, or verifiable sightings made since its initial inclusion, i rejected it from the updated checklist of mznp. spectacled dormouse (graphiurus ocularis) like the previous two species, the spectacled dormouse appears to have been included based on what was believed to be the most plausible distribution at the time (skead 1958). although the first published field guide for the park lists the species as being present, it provides no corroborating evidence for its listing (grobler & hall-martin 1982). certainly, none of the more comprehensive small mammal surveys conducted at mznp have recorded the species (de graaff & nel 1970; kok, parker & barker 2012; nel & pretorius 1971). as such, i rejected it from the updated checklist of mznp. brown mouse (rattus norvegicus) the first published field guide for the park lists this alien species as being present, but without any corroborating evidence (grobler & hall-martin 1982). as none of the more comprehensive small mammal surveys conducted at mznp have recorded the species (de graaff & nel 1970; kok et al. 2012; nel & pretorius 1971), i rejected it from the updated checklist of mznp. grant’s rock mouse (michaelamys granti) similarly, grobler and hall-martin (1982) list grant’s rock mouse as being present without any supporting evidence. since none of the more comprehensive small mammal surveys conducted at mznp have recorded the species (de graaff & nel 1970; kok et al. 2012; nel & pretorius 1971), i rejected it from the updated checklist of mznp. fat mouse (steatomys pratensis) the inclusion of this species was based on what was believed to be the most plausible distribution at the time (skead 1958). as no additional specimens have been collected, or verifiable sightings made since its initial inclusion, i rejected it from the updated checklist of mznp. gerbil mouse (malacothrix typica) the inclusion of this species was based on what was believed to be the most plausible distribution at the time (skead 1958). as no additional specimens have been collected, or verifiable sightings made since its initial inclusion, i rejected it from the updated checklist of mznp. reddish-grey musk shrew (crocidura cyanea) the inclusion of this species was based on what was believed to be the most plausible distribution at the time (skead 1958). as no additional specimens have been collected, or verifiable sightings made since its initial inclusion, i rejected it from the updated checklist of mznp. straw-coloured fruit bat (eidolon helvum) the inclusion of this species was based on what was believed to be the most plausible distribution at the time (skead 1958). as no additional specimens have been collected, or verifiable sightings made since its initial inclusion, i rejected it from the updated checklist of mznp. geoffroy’s horseshoe bat (rhinolophus clivosus) this species is listed in the first field guide for the park (grobler & hall-martin 1982). although no corroborating evidence is supplied in the guide, horseshoe bats are relatively easily distinguished from other insectivorous bats based on their characteristic nose-leaf patterns (monadjem et al. 2020). in addition, hans grobler was the resident biologist in the mznp at the time and he could easily have observed horseshoe bats roosting in the cave or culvert. however, despite geoffroy’s horseshoe bat (rhinolophus clivosus) being broadly distributed across the eastern cape (monadjem et al. 2020), without any corroborating morphometric or echolocatory evidence, it could be confused with at least one other horseshoe bat species (monadjem et al. 2020). as such, i rejected it from the updated checklist of mznp. mauritian tomb bat (taphozous mauritianus) the inclusion of this species was based on what was believed to be the most plausible distribution at the time (skead 1958). as no additional specimens have been collected, or verifiable sightings made since its initial inclusion, i rejected it from the updated checklist of mznp. natal long-fingered bat (miniopterus natalensis) the inclusion of this species was based on what was believed to be the most plausible distribution at the time (skead 1958). as no additional specimens have been collected, or verifiable sightings made since its initial inclusion, i rejected it from the updated checklist of mznp. leopard (panthera pardus) skead (1958) was the only author who believed that leopards could be present in the mznp. despite photographic evidence of leopards elsewhere in the karoo, leopards have never been reliably recorded in the mznp where numerous camera trapping surveys have been (and continue to be) conducted. as such, i rejected it from the updated checklist of mznp. white-tailed mongoose (ichneumia albicaudatus) the inclusion of this species was based on what was believed to be the most plausible distribution at the time (skead 1958). as no additional specimens have been collected, or verifiable sightings made since its initial inclusion, i rejected it from the updated checklist of mznp. spotted-neck otter (hydrictis maculicollis) the inclusion of this species was based on what was believed to be the most plausible distribution at the time (skead 1958). as no additional specimens have been collected, or verifiable sightings made since its initial inclusion, i rejected it from the updated checklist of mznp. plains zebra (equus quagga) plains zebra were intentionally introduced into mznp in 1998 (sanparks 2016). as plains zebra are known to hybridise with cape mountain zebras, producing fertile hybrids (kotze et al. 2017), plains zebras resembling the extinct quagga were introduced in an attempt to recreate the quagga through selective breeding. however, in 2014, sanparks discontinued the programme and removed all plains zebra individuals from the park. by 2015, plains zebra were no longer recorded during annual aerial game censuses and are believed to have been eradicated from the park (bissett et al. 2019). as such, i rejected it from the updated checklist of mznp. common reedbuck (redunca arundinum) common reedbuck (redunca arundinum) were also intentionally introduced into mznp in the 1960s (penzhorn 1971). however, grobler and bronkhorst (1981) note that they failed to thrive and by 1981 were no longer present in the park. as such, i rejected it from the updated checklist of mznp. re-introductions, immigrations and introductions at the time of proclamation (1937), the only larger herbivores believed to be present in the park were cape mountain zebra, klipspringer (oreotragus oreotragus), steenbok (raphicerus campestris), common duiker (sylvicapra grimmia), grey rhebok (pelea capreolus), mountain reedbuck (redunca fulvorufula) and rock hyrax (procavia capensis) (grobler & bronkhorst 1981; grobler & hall-martin 1982; penzhorn 1971). the first ungulate re-introduction occurred in the 1940s when a small group of springbok (antidorcas marsupialis) were translocated from a farm near bedford (penzhorn 1971). re-introductions of blesbok (damaliscus pygargus phillipsi), black wildebeest (connochaetes gnou), red hartebeest (alcelaphus buselaphus), gemsbok (oryx gazella) and eland (tragelaphus oryx) followed in the 1960s (grobler & hall-martin 1982; novellie & knight 1994; penzhorn 1971). all of these ungulate re-introductions were successful except for that of the gemsbok. the entire re-introduced gemsbok population was removed in 1981 because ‘they were not doing well’ (grobler & bronkhorst 1981). the reason for their failure to thrive is likely because of the poor habitat available to the animals at the time. till the early 2000s, only the more mountainous, sourveld region of the park was available to the animals, and the overall poor palatability of the grass species in this section, and greater climatic fluctuations in the mountains, could have negatively affected the gemsbok population (grobler & hall-martin 1982). however, with the expansion of the park in the early 2000s, the additional ‘sweeter’ veld became available (brown & bezuidenhout 2005), and a second re-introduction of gemsbok to the park in 2004 was much more successful. gemsbok numbers are now in excess of 200 individuals and they are very seldom sighted in the southern (more mountainous) section of the park (bissett et al. 2019). more recent, large mammal re-introductions include buffaloes (syncerus caffer) in 1997, black rhinoceros (diceros bicornis) in 2002, cheetah (acinonyx jubatus) in 2007, brown hyaena (parahyaena brunnea) in 2008 and lion (panthera leo) in 2013 (bissett et al. 2019; sanparks 2016). all of these species were re-introduced in accordance with the park’s objective of restoring the historical diversity of large mammals of the region (sanparks 2016) and have been successful (bissett et al. 2019). the late 1970s saw the natural immigration of bushpig (potomochoerus larvatus), greater kudu (tragelaphus strepsiceros) and bushbuck (tragelaphus scriptus) into the park (grobler & bronkhorst 1981). the only other non-volant mammal which is believed to have naturally immigrated into the park is the extralimital warthog (phacochoerus africanus) in approximately 2012 (parker, bissett & craig 2013). although thought to be present in the region at the time of establishment of the park (skead 1958), it was not until mid-1979 that two african clawless otters (aonyx capensis) escaped from a temporary cage on the babylons toren section of the park (grobler & bronkhorst 1981). grobler and bronkhorst (1981) speculated that as the pair was a male and a female that perhaps they would establish themselves in the park. however, it was not until one of the first intensive camera trapping surveys of the park that african clawless otters were reliably recorded again (figure 3; comley 2016). figure 3: three african clawless otters (aonyx capensis) photographed on a camera trap during a brown hyaena (parahyaena brunnea) population assessment (comley 2016) in the mountain zebra national park, eastern cape, south africa. permission to use the image granted by jessica comley. taxonomic conundrums and future research priorities the number of species within the genus lepus has been a matter of debate amongst taxonomists for some time (skinner & chimimba 2005). despite some regional differences in skull and ear length, it is now accepted that the only two species to occur in south africa are l. saxatilis and l. capensis (skinner & chimimba 2005). both of these species have been reliably collected as specimens in mznp (grobler & bronkhorst 1981; nel & pretorius 1971). however, all subsequent records have been through direct observations or camera trap photographs. as the most reliable way to distinguish between the two species is by comparison of the incisors (skinner & chimimba 2005), direct observations and camera trap photographs of hares, with potentially variable pelage, should be interpreted with caution. thus, either the collection of museum specimens or the sampling of dna from captured specimens should be a future research priority. cryptic speciation amongst the laminate toothed rats (family muridae: subfamily murinae: tribe otomyini), to which the vlei (otomys irroratus), bush karoo (o. unisulcatus) and sloggett’s ice rat (o. sloggetti) belong, is common, especially in mountainous habitats (taylor et al. 2019). in fact, recent taxonomic work has demonstrated that o. irroratus senso lato can be confused morphologically with at least one other co-occurring species o. karoensis (taylor et al. 2019). as such, i recommend that the specific assignment of the three laminate toothed rats putatively present in mznp be verified through the collection of specimens for morphological and genetic analyses. notwithstanding the taxonomic conundrums that require resolution, it is clear that there has been unequal surveying and documenting of the mammals present within the mznp over time. there has clearly been a bias towards the larger, more charismatic species and less of a focus on the small, more cryptic mammal groups (but see de graaff & nel 1970; kok et al. 2012; nel & pretorius 1971). as such, i believe that a concerted effort should be made to comprehensively sample the mznp for golden moles, elephant shrews, rodent moles, gerbils and bats. whilst the sampling of the fossorial small mammals can be challenging and labour intensive, recent technological advances in acoustic technology make the sampling of echolocating bats (a notoriously difficult mammal group to sample) much more feasible (parker & bernard 2019). conclusion like most of the initial national parks gazetted in south africa, the mznp was initially proclaimed to protect the survival of just one species, the mountain zebra. however, as time has progressed, and conservation priorities have changed (sanparks 2016), the conservation footprint of the mznp has been expanded and the diversity of mammals receiving formal protection has been increased. whilst the updated and revised checklist presented here is the most comprehensive current list available, i am confident that with targeted survey work, additional mammal species will be added. acknowledgements i would like to thank my colleagues, ric bernard, adrian craig and chris brown, for getting much of the more recent mammal survey work off the ground in the late 1990s and early 2000s. thanks are also because of the park managers and section rangers at mznp for their support of the work. thank you to paddy gordon, johan de klerk, phumla mzazi, lesley-ann brown, robyn woods, craig williams, megan taplin and greg bond. additionally, thank you to all the students, colleagues and other ‘hangers on’ who assisted in the field over the years. special mention must be made of charlene bissett for all the glorious hours in the field, the fire-side chats (and arguments) and her critical and constructive eye on the manuscript. finally, i acknowledge rhodes university for its financial assistance. competing interests the author declares that he has no financial or personal relationships that may have inappropriately influenced him in writing this article. author’s contributions d.m.p. contributed solely to this work. ethical considerations this article followed all ethical standards for research without direct contact with human or animal subjects. funding information this research received no specific grant from any funding agency in the public, commercial or not-for-profit sectors. data availability the data that support the findings of this study are available from the corresponding author, d.m.p., upon reasonable request. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the author. references bernard, r.t.f., craig, a.j.f.k. & hulley, p.e., 2005, the effects of past land use on biodiversity, mountain zebra national park, annual progress report to sanparks, department of zoology and entomology, rhodes university, grahamstown. bernard, r.t.f. & parker, d.m., 2006, ‘the use of archaeological and ethnographical information to supplement the historical record of the distribution of large mammalian herbivores’, south african journal of science 102, 117–119. bezuidenhout, h. & brown, l.r., 2008, ‘vegetation description of the doornhoek section of the mountain zebra national park (mznp), south africa’, koedoe 50(1), 32–41. https://doi.org/10.4102/koedoe.v50i1.142 bissett, c., 2012, space use and feeding ecology of large carnivores in small, fenced reserves: case study of cheetahs and brown hyaenas in mountain zebra national park, annual progress report to sanparks, department of zoology and entomology, rhodes university, grahamstown. bissett, c., ferreira, s., ramoelo, a., bezuidenhout, h., januarie, r. & raselabe, m., 2019, mountain zebra national ark herbivore off-take recommendations 2019: an integrated approach combining local knowledge with data derived from animal census, herbivore models and satellite imagery, internal report 30/2019, scientific services, sanparks, pretoria. brown, l.r. & bezuidenhout, h., 2005, ‘a vegetation description and classification of the farms ingleside and welgedacht in the mountain zebra national park, eastern cape’, koedoe 48(2), 23–42. https://doi.org/10.4102/koedoe.v48i2.92 comley, j., 2016, ‘population assessment and feeding ecology of brown hyaenas (hyaena brunnea) in mountain zebra national park, eastern cape, south africa’, msc thesis, department of zoology and entomology, rhodes university, grahamstown. craig, a.j.f.k., hulley, p.e. & parker, d., 2005, ‘a re-assessment of the avifauna of the mountain zebra national park’, koedoe 48(1), 95–113. https://doi.org/10.4102/koedoe.v48i1.169 craig, a.j.f.k., parker, d.m., hulley, p.e. & bernard, r.t.f., 2009, the effects of past land use on biodiversity, mountain zebra national park, annual progress report to sanparks, department of zoology and entomology, rhodes university, grahamstown. craig, a.j.f.k., parker, d.m., hulley, p.e. & bernard, r.t.f., 2010, the effects of past land use on biodiversity, mountain zebra national park, annual progress report to sanparks, department of zoology and entomology, rhodes university, grahamstown. craig, a.j.f.k., parker, d.m., hulley, p.e. & bernard, r.t.f., 2011, the effects of past land use on biodiversity, mountain zebra national park, annual progress report to sanparks, department of zoology and entomology, rhodes university, grahamstown. de bondi, n., white, j.g., stevens, m. & cooke, r., 2010, ‘a comparison of the effectiveness of camera trapping and live-trapping for sampling terrestrial small mammal communities’, wildlife research 37(6), 456–465. https://doi.org/10.1071/wr10046 de graaff, g. & nel, j.a.j., 1970, ‘notes on the smaller mammals of the eastern cape national parks’, koedoe 13(1), 147–149. https://doi.org/10.4102/koedoe.v13i1.736 de graaff, g. & penzhorn, b.l., 1976, ‘the reintroduction of springbok antidorcas marsupialis into south african national parks – a documentation’, koedoe 19(1), 75–82. https://doi.org/10.4102/koedoe.v19i1.1184 du toit, c.f., 1980, ‘the yellow mongoose cynictis penicillata and other small carnivores in the mountain zebra national park’, koedoe 23(1), 179–184. https://doi.org/10.4102/koedoe.v23i1.645 grobler, j.h., 1981, ‘feeding behaviour of the caracal felis caracal shreber 1776 in the mountain zebra national park’, south african journal of zoology 16(4), 259–262. https://doi.org/10.1080/02541858.1981.11447764 grobler, j.h. & bronkhorst, p.j., 1981, ‘additions and amendments to the bird and mammal lists of the mountain zebra national park’, koedoe 24, 199–203. https://doi.org/10.4102/koedoe.v24i1.630 grobler, h. & hall-martin, a., 1982, a guide to the mountain zebra national park, national parks board of trustees, pretoria. horak, i.g. & fourie, l.j., 1986, ‘parasites of domestic and wild animals in south africa. xix. ixodid ticks and fleas on rock dassies (procavia capensis) in the mountain zebra national park’, onderstepoort journal of veterinary research 53(2), 123–126. horak, i.g., fourie, l.j., novellie, p.a. & williams, e.j., 1991, ‘parasites of domestic and wild animals in south africa. xxvi. the mosaic of ixodid tick infestations on birds and mammals in the mountain zebra national park’, onderstepoort journal of veterinary research 58(3), 125–136. jackson, c. & bernard, r.t.f., 2005, ‘effects of supplementary food on the winter inhibition of reproduction in male and female four-striped field mice (rhabdomys pumilio)’, reproduction, fertility and development 17(4), 393–400. https://doi.org/10.1071/rd04134 kok, a.d., parker, d.m. & barker, n.p., 2012, ‘life on high: the diversity of small mammals at high altitude in south africa’, biodiversity and conservation 21(11), 2823–2843. https://doi.org/10.1007/s10531-012-0340-0 kotze, a., zimmerman, d., dalton, d.l., mnisi, c., taplin, m., novellie, p. et al., 2017, ‘hiding in plain sight: evidence of hybridization between cape mountain zebra (equus zebra zebra) and plains zebra (equus quagga burchelli)’, african journal of wildlife research 47(1), 59–64. https://doi.org/10.3957/056.047.0059 monadjem, a., taylor, p.j., cotterill, f.p.d. & schoeman, m.c., 2020, bats of southern and central africa: a biogeographic and taxonomic synthesis, wits university press, johannesburg. moolman, l.c., 1984, ‘n vergelyking van die voedingsgewoontes van die rooikat felis caracal binne en buite die bergkwagga nasionale park’, koedoe 27(1), 121–129. https://doi.org/10.4102/koedoe.v27i1.554 morris, d., 2010, ‘the effects of land use change on the small mammal communities of the mountain zebra national park, eastern cape, south africa’, honours thesis, department of zoology and entomology, rhodes university. mucina, l., rutherford, m.c., palmer, a.r., milton, s.j., scott, l., lloyd, j.w. et al., 2006, ‘nama-karoo biome’, in l. mucina & m.c. rutherford (eds.), the vegetation of south africa, lesotho and swaziland, pp. 325–347, south african national biodiversity institute, pretoria. nel, j.a.j. & pretorius, j.j.l., 1971, ‘a note on the smaller mammals of the mountain zebra national park’, koedoe 14, 99–110. https://doi.org/10.4102/koedoe.v14i1.719 novellie, p.a. & knight, m., 1994, ‘repatriation and translocation of ungulates into south african national parks: an assessment of past attempts’, koedoe 37(1), 115–119. https://doi.org/10.4102/koedoe.v37i1.328 parker, d.m. & bernard, r.t.f., 2019, ‘the use of acoustic detectors for assessing bat species richness and functional activity in a south african national park’, mammalia 83(1), 53–63. https://doi.org/10.1515/mammalia-2017-0055 parker, d.m., bissett, c. & craig, a.j.f.k., 2013, the effects of past land use on biodiversity, mountain zebra national park, mid-year fieldtrip report to sanparks, department of zoology and entomology, rhodes university, grahamstown. parker, d.m., bissett, c. & craig, a.j.f.k., 2014, the effects of past land use on biodiversity, mountain zebra national park, mid-year fieldtrip report to sanparks, department of zoology and entomology, rhodes university, grahamstown. penzhorn, b.l., 1971, ‘a summary of the reintroduction of ungulates into south african national parks (to december 1970)’, koedoe 14(1), 145–159. https://doi.org/10.4102/koedoe.v14i1.725 sanparks, 2016, mountain zebra national park: park management plan for the period 2016–2026, south african national parks, pretoria. skead, c.j., 1958, ‘mammals of the uitenhage and cradock districts c.p. in recent times’, koedoe 1(1), 19–59. https://doi.org/10.4102/koedoe.v1i1.864 skinner, j.d. & chimimba, c.t., 2005, the mammals of the southern african subregion, cambridge university press, cape town. stuart, c.t., 1987, ‘the vlei rat otomys irroratus recorded in the mountain zebra national park’, koedoe 30, 167. https://doi.org/10.4102/koedoe.v30i1.511 taylor, p.j., kearney, t., dalton, d.l., chakona, g., kelly, c.m.r. & barker, n.p., 2019, ‘biomes, geology and past climate drive speciation of laminate-toothed rats on south african mountains (murinae: otomys)’, zoological journal of the linnean society 189(3), 1046–1066. https://doi.org/10.1093/zoolinnean/zlz134 van der walt, p.t., 1980, ‘a phytosociological reconnaissance of the mountain zebra national park’, koedoe 23(1), 1–32. https://doi.org/10.4102/koedoe.v23i1.632 whittington-jones, g.m., bernard, r.t.f. & parker, d.m., 2008, ‘bushclumps as refugia for small mammals in two eastern cape conservation areas’, african zoology 43(2), 273–276. https://doi.org/10.1080/15627020.2008.11657244 appendix 1 table 1-a1: the updated (2020) checklist of 68 mammal species in the mountain zebra national park, eastern cape, south africa. see text for details on the criteria used for inclusion. abstract introduction methods results ethical considerations discussion conclusion acknowledgements references appendix 1 about the author(s) elizabeth j. opperman department of botany and zoology, stellenbosch university, south africa michael i. cherry department of botany and zoology, stellenbosch university, south africa nokwanda p. makunga department of botany and zoology, stellenbosch university, south africa citation opperman, e.j., cherry, m.i. & makunga, n.p., 2018, ‘community harvesting of trees used as dens and for food by the tree hyrax (dendrohyrax arboreus) in the pirie forest, south africa’, koedoe 60(1), a1481. https://doi.org/10.4102/koedoe.v60i1.1481 review article community harvesting of trees used as dens and for food by the tree hyrax (dendrohyrax arboreus) in the pirie forest, south africa elizabeth j. opperman, michael i. cherry, nokwanda p. makunga received: 22 june 2017; accepted: 20 nov. 2017; published: 28 feb. 2018 copyright: © 2018. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract forests in south africa are harvested by local communities for multiple purposes and this affects the animals that inhabit them. the tree hyrax (dendrohyrax arboreus) has a restricted distribution and utilises various tree species as dens and a source of food. in this article, we determined, through a series of interviews in the communities surrounding the pirie forest, which plant species are harvested by local people and whether these overlap with those used by the tree hyrax. in addition, we determined the extent to which tree hyraxes are hunted by these communities. of the trees used by the hyrax as dens in the pirie forest, afrocarpus falcatus, schotia latifolia, andrachne ovalis, teclea natalensis and apodytes dimidiata are important resources for local communities. but as these are harvested at relatively low levels, it is unlikely that current harvesting has a large impact on the tree hyrax. opportunistic hunting occurs, but the hyrax is not targeted by hunters. very limited commercial harvesting of a. falcatus has been taking place in the pirie forest since 1975, but its impact on the hyrax population, although undetermined, is also unlikely to be high. we recommend that the pirie forest tree hyrax population should be monitored by forest management in order to ascertain the impact of both commercial and community harvesting over the past quarter century. conservation implications: tree hyrax populations in the pirie forest should be actively monitored by management on an annual basis. introduction the world’s forests have experienced high levels of destruction (owing to deforestation) and degradation (owing to unsustainable harvesting). in south africa, indigenous forests cover only 0.56% of the land surface (low & rebelo 1996), but these forests have a long history of exploitation. timber harvesting, in particular, increased between 1890 and 1940 but has since declined owing to increased use of alien vegetation for timber used in construction (brown 2003; cunningham & davis 1997; lawes, midgley & chapman 2004). however, many forests are harvested by proximate local communities which are dependent on forests as a source of food, medicine and timber for both household consumption and income; harvesting also forms an integral part of the culture of these communities (dold & cocks 2012). as south africa has very little forest cover, forest-specialised species are restricted to relatively few forests, which in many cases are subject to extensive human exploitation (castley & kerley 1996). in forests exploited by humans, the forest biota is clearly affected. for example, cooper, wannenburgh and cherry (2017) have found that half of south africa’s forest-dependent bird species have declined over the past quarter century, particularly in the eastern cape forests, despite the fact that the forests have not decreased in area. the tree hyrax (dendrohyrax arboreus) is an arboreal, nocturnal and folivorous small mammal with a disjunct distribution across southern and eastern africa (see figure 1 in gaylard et al. 2016). the tree hyrax uses the hollows of trees for their dens and prefers hollow, large, upright and dying trees with many cavity entrances (gaylard & kerley 2001). in south africa, it is confined to relatively few forests in the eastern cape and kwazulu-natal: it is estimated that only 1% of its former range in south africa, lesotho and swaziland is currently occupied (gaylard et al. 2016). interestingly, different tree species are used as dens and for foraging in the coastal forests of alexandria and springmount as opposed to the afromontane mistbelt pirie forests (gaylard & kerley 1997, 2001). although the tree hyrax is listed as of least concern by the international union for conservation of nature (iucn), the listing mentions that as the southern and eastern african forests are under threat, the situation needs to be monitored (butynski, hoeck & de jong 2015). the local red listing is endangered (gaylard et al. 2016). figure 1: map showing the pirie forest and the locations of the villages in which the interviews were conducted. the yellow pins represent the communities in this study. this study focused on community harvesting of trees in the vicinity of the pirie forest (figure 1), which forms part of the amatole mistbelt forests in the eastern cape province of south africa, and the possible threat it poses to the forest’s tree hyrax populations. in the pirie forest, the hyrax utilises only six tree species as dens: searsia chirindensis (red currant), schotia latifolia (forest boerboon), andrachne ovalis (bastard lightning bush), apodytes dimidiata (white pear), afrocarpus falcatus (outeniqua yellowwood) and teclea natalensis (small-fruited teclea) (gaylard & kerley 2001). by contrast, tree hyraxes have been recorded as foraging on the leaves of 36 different plant species in the pirie forest (gaylard & kerley 1997). the pirie forest is one of only two forests nationally (the other is the knysna forest) where commercial harvesting is permitted by the department of agriculture, forestry and fisheries (daff) for two yellowwood species, a. falcatus and podocarpus latifolius (real yellowwood) (mpisekaya & kameni 2007). the former is one of the species in the pirie forest used as a den by the hyrax (gaylard & kerley 2001). although harvesting is permitted only of trees which are dead or dying, are wind fallen or have lost their crowns, this may still have an impact on the hyrax as they depend on dead or dying trees with hollows for their dens (lawes, mealin & piper 2000). commercial wood harvesting in the amatole forests was once again permitted from 1975 onwards, having been terminated in 1940 (mpisekaya & kameni 2007). since 2007, harvesting has been limited to 0.1% of available stems of these species per annum. indiscriminate historical harvesting may explain why hyraxes occur at a low density in the pirie forest, in contrast to the springmount forest, a drier dune forest where it occurs at higher densities (gaylard 1994). but as a. falcatus comprised only 14.3% of the trees used by the hyrax as dens in the pirie forest in 1992, despite it being a dominant species there (gaylard & kerley 2001; greyling & huntley 1984), it is unlikely that current levels of commercial harvesting impact negatively tree hyrax numbers. the pirie forest was chosen as a study site because of its proximity to villages, and the fact that like most forests protected by daff, it is not fenced and so can be easily accessed. in addition to commercial harvesting, local community members are officially allowed to take only a daily head-load (inyanda in isixhosa but popularly referred to as theza) of wood lying on the ground, intended primarily for use as firewood, which should have no impact on the hyrax (j. feely [nelson mandela metropolitan university, port elizabeth] pers. comm., 01 october 2015). despite this, illegal harvesting of bark for spiritual and medicinal purposes, as well as cutting trees for poles for building material, is widespread in eastern cape forests (dold & cocks 2012). the former can result in ring-barking of trees, and cutting of trees for poles can result in canopy reduction, both of which could negatively affect the tree hyrax. in addition, illegal hunting of small game with dogs is also an important pastime among young men and boys (dold & cocks 2012). in this study, we used a series of semi-structured interviews in the communities surrounding the pirie forest in order to determine the plant species harvested by the local communities. in addition, we aimed to gauge the extent of harvesting in the pirie forest and for what purposes harvested material is used. we also aimed to determine the extent to which hunting for tree hyrax takes place in the forest and whether communities living around the pirie forest were aware of the tree hyrax and, if so, knew anything about its behaviour. we predicted that harvested tree species would overlap with those utilised by the hyrax as den and food sources described in gaylard and kerley (2001). methods the study was conducted in seven randomly selected communities in close proximity to the pirie forest in the eastern cape of south africa in july 2015 (figure 1). the pirie forest (32°43’s, 27°16’e) is situated between keiskammahoek and king williams town at an altitude between 540 m and 1300 m, is 3173 ha in extent and has a stand density of 2115.6 stems/ha (von maltitz et al. 2003). the area has a very steep sloping pediment with cliffs as well as several rock faces, and the major river running through it is the buffalo river (greyling & huntley 1984). interviews were conducted in the villages of pirie mission (32°47’33.03”s, 27°14’54.86”e), cwengcwe (32°46’15.91”s, 27°20’38.47”e), mxhalanga (32°44’1.37”s, 27°21’31.35”e), tyusha (32°41’51.01”s, 27°22’16.30”e), new rest (33°57’23.95”s, 24°18’42.69”e), nakani (32°47’45.49”s, 27°13’5.51”e) and gubevu (32°5’6.00”s, 28°23’9.00”e) (google earth 2013). at each village, the village leader or chief was asked for permission to conduct interviews in the community. the leaders, who were informed of the purpose of the survey as well as the rights of each participant, assisted in identifying known harvesters and those who hunt animals within the forest. semi-structured questionnaires (appendix 1) were used to determine which tree species are used, and to what extent and purpose. participants were identified by a snowballing or referral technique in which the person interviewed provided possible future participants, based on existing connections within the community. the purpose of the study and the rights of the participants were explained to each participant in isixhosa before the interview was undertaken. each participant was also asked to sign a consent form. anonymity of participants was guaranteed in terms of reporting results from the study. but as harvesters and hunters may be reluctant to share information openly, our results should be regarded as reporting minimum levels of both activities. one of the interviews was conducted in english, and the rest in isixhosa with the help of a translator. answers were noted down and one of the interviews was recorded after consent from the participant was obtained. a total of 28 people were interviewed, who were asked basic biographical information plus whether they harvest, which species they harvest, how much they harvest and how often they harvest. in addition, they were asked about hunting practices surrounding the hyrax, and general knowledge of the hyrax. the questionnaire consisted of multi-choice and open-ended questions. the sample comprised 20 men and 8 women. the participants interviewed were evenly spread across age classes (table 1); the majority of the participants earned less than r500 per month and had an education lower than grade 7. table 1: the demographics of the participants interviewed at the pirie forest, south africa. a list of all the plants harvested by the participants was made, and the isixhosa vernacular names were translated into scientific names using dold and cocks (1999). the relative frequency of citation (rfc) for each plant species was determined. the rfc can be used to estimate the local importance of a certain plant species based on the number of times it was mentioned and is calculated using the equation: where rfc = relative frequency of citation, fc = number of participants mentioning the use of the species and n = number of participants taking part in the survey (samoisy & mahomoodally 2015). the perceptions of local people, distilled from open-ended interviews, were also documented qualitatively. results of the six tree species used by the hyrax for both dens and forage, the participants reported harvesting of five: s. latifolia, t. natalensis, a. falcatus, and. ovalis and apo. dimidiata. of these, a. falcatus and s. latifolia had the highest rfc values (figure 2). of the respondents, 57% harvested, 22% harvested and hunted, 7% hunted and the remaining 14% did neither but had general knowledge of harvesting practices in the area (figure 3a). of the harvesters, 36% reported doing so only for personal use; the remaining 64% did so both for personal use and selling. regarding frequency of harvesting, only 7% harvested daily, with most participants reporting that the frequency of harvesting depends either on what they need or that they harvest only every few months or on a monthly basis (figure 3b). of the 12 respondents who were able to estimate quantities harvested, 25% reported harvesting less than 1 kg per month; 25% between 1 kg and 50 kg; and just under 17% reported harvesting in each of the categories, namely 50 kg – 100 kg, 100 kg – 200 kg and over 200 kg per month. figure 2: relative frequency of citation of tree species harvested near the pirie forest which are used as dens by the tree hyrax. figure 3: (a) the occupation of the participants, (b) the frequency of harvesting by participants, (c) the relative frequency of citation of the plant parts used by participants and (d) the rfc of the uses of the plants described by harvesters near the pirie forest. the participants mentioned that the bark of the plants was used most (figure 3c), followed by the stems, leaves, branches and lastly the roots and/or tubers. most participants said that the products they harvest are used for medicinal purposes, followed by firewood, building material and customs and traditions (figure 3d), including the passage of boys to manhood and ancestral worship ceremonies. a total of 42 plant species was reported to be harvested. the alien acacia mearnsii (black wattle) and ptaeroxylon obliquum (sneezewood) were mentioned most by participants. other trees mentioned with a high rfc were curtisia dentata (assegai), strychnos henningsii (panda’s walking stick) and olea europaea (wild olive). figure 4 shows the rfc values of the plants used (1) medicinally, (2) spiritually, (3) as building material and (4) as firewood in these separate categories. as expected, a wide range of plants are used for traditional medicines and for all tree species that are harvested from the forest, the bark is utilised; three of these species have been reported as being utilised as dens by the tree hyrax in the pirie forest (gaylard & kerley 2001; figure 4a). a majority (61%) of the respondents indicated that the products harvested were both for personal use and as a source of income, being sold either within their village or at local markets for plant medicines. figure 4: the relative frequency of citation for plants used (a) medicinally, (b) spiritually, (c) as building materials and (d) as firewood by harvesters in the pirie forest. plant species used by the tree hyrax as dens are indicated in grey. of the respondents, 89% knew about the hyrax; 61% had heard the hyrax calling; 54% had seen it; and all eight hunters had hunted it. of the respondents, 46% indicated that the hyrax is hunted for meat; 12.5% of hunters reported hunting the hyrax for medicinal purposes and for entertainment, respectively. those respondents familiar with tree hyraxes indicated that these animals were active at night and were ‘difficult to catch as they live in tall trees’. many respondents started to mimic the call of the tree hyrax during the interview, indicating that the isixhosa word for it (umqha) when pronounced has a similar sound to its call. those that had seen the tree hyrax likened it to imbila (rock dassie) but emphasised that it lives in trees and is shy. one of the respondents, who identified himself as a hunter, remarked that the hyrax is a ‘tame, shy animal that is not intimidating and it does not really inspire one to try and hunt it’ (participant 24, male, matric, unemployed, 18–29). another conversation referred to the isixhosa idiom unyabe njengomqha which is translated as ‘as shy as a tree hyrax’ (participant 19, female, tertiary education, pensioned high school principal, 60–69). a traditional healer indicated that there is a general belief that eating the meat of the tree hyrax will make you tame, introverted and generally docile, but mentioned that it is not common to hunt the hyrax as it is difficult to catch them owing to their nocturnal habits. another man had tried to hunt hyraxes but had never been successful. in general, those familiar with the hyrax also mentioned that it calls in the evenings, before proceeding to imitate the call, and noted that its call is very loud (and also peculiar). to assess the importance of the forest and its likelihood as a resource in future generations, we were interested in knowing at what age villagers started to learn about the forest or use the forest, and whether they perceived the forest as important for future generations. lastly, we wanted to understand whether they saw the forest as a renewable resource. all the participants indicated that their ancestors had utilised the forest and many of them indicated that they would want their descendants to continue the culture of utilising the forest, or at least understand the value of the forests in their lives. one of the respondents indicated that they would teach their children as it will ‘restore and uphold the cultural heritage and traditions of the amaxhosa’ (participant 16, male, grade 7 or below, unemployed, 40–49). it became apparent that linking the forest and its future regeneration is also important as ‘ancestors are linked to nature and the forest is a refuge for their souls’ (participant 4, male, grade 7 or below, pensioner, 50–59). the villagers regarded the forests as being renewable, and that it is renewable as some plants: re-sprout through roots and also some trees regenerate through seeds; and those animals reliant on trees for their habitat and food as a resource would thus be maintained in the future. (participants 8, female, grade 7 or below, unemployed, 50–59) with regard to knowledge transmission pertaining to the hyrax and harvesting of plant material, 79% of the interviewees expressed the importance of future generations carrying on these traditions. they emphasised this as a way of staying connected with their ancestors. one of the respondents said: ‘although it is our heritage, we now live in a modern context and traditions are not always passed on. we want our children to know and appreciate our culture’ (participant 19, female, tertiary education, pensioned high school principal, 60–69). our data illustrated that men mostly received knowledge from their fathers or grandfathers, whilst women mostly received knowledge from their mothers or grandmothers, and thus knowledge was transferred vertically, and mostly gender specifically. some of the participants indicated that the sharing of knowledge with peers is also important. one man indicated that the tree hyrax is not only important medicinally but it is also safe for women to eat it ‘as it does not interfere with their ability to bear children’ (participant 16, male, grade 7 or below, unemployed, 40–49). although they have no substantial uses for the tree hyrax, it was perceived as an important part of the general ecology of the forest. this was further echoed by those without children, who confirmed that being in the forest whether for hunting, harvesting and/or leisure allowed them to commune with nature, rejuvenating their spirits and essentially giving them a ‘new life’. many respondents indicated that they had started to visit the forest and to be taught how to use the forest from a young age (between the ages of 8 and 16). ethical considerations participation in the interviews was voluntary and the ethical guidelines of stellenbosch university were followed. ethical clearance for the interviews was obtained from the departmental ethical clearance committee (desc) at the university of stellenbosch (reference number: desc/opperman/may2015/1). discussion although five (s. latifolia, and. ovalis, apo. dimidiata, a. falcatus, t. natalensis) of the six tree species used by the tree hyrax as dens in the pirie forest (gaylard & kerley 2001) were reported (figure 2) as being harvested (out of a total of 42 reported harvested species), all five had low rfc values. of these, s. latifolia and a. falcatus had the highest rfc values, (0.14 in each case). the other tree species had very low rfc values of 0.04 (figure 2), suggesting that they were being harvested less. searsia chirindensis was the only tree used by the hyrax (gaylard & kerley 2001) which was not reported as being harvested. as the bark of s. latifolia is used medicinally (dold & cocks 2012) and as a. falcatus is in addition commercially logged (mpisekaya & kameni 2007), it may be worth investigating community harvesting levels of these tree species in more detail. of the 36 plant species reported in the diet of the tree hyrax in the pirie forest (gaylard & kerley 1997), only seven were mentioned as being harvested (rfc values in brackets): o. europaea (0.32), hippobromus pauciflorus (false horsewood) (0.11), pleurostylia capensis (coffee pear), behnia reticulata (forest smilax), olea capensis (ironwood) (0.07), apo. dimidiata and ekebergia capensis (cape ash) (0.04). however, as tree hyraxes eat leaves from a wide range of species, harvesting is unlikely to impact their foraging habits heavily. disturbance relating to the harvesting of trees other than those it uses as dens or for foraging could conceivably be detrimental to the tree hyrax, owing to a loss of canopy cover. topp-jørgensen et al. (2008) emphasised how tree hyraxes preferred undisturbed forests with closed canopies, with the number of calling individuals being highest in undisturbed closed canopies. more open canopies thus lead to habitat unsuitability, as the hyrax uses arboreal pathways as a predator escape strategy. with regard to hunting, none of the female participants and only a minority (8) of the 20 male participants reported to hunt or have hunted the hyrax during the course of their lives. hunting of the hyrax that does take place is opportunistic and largely for recreational purposes (the experience of tasting hyrax meat). for subsistence, bush pigs (potamochoerus larvatus) are hunted, but the time investment required to hunt the hyrax is not justified owing to its relatively small size and as it is not used for traditional purposes. in xhosa culture, the tree hyrax is associated with being ‘timid’ and is often used to express perceptions being linked to having a docile personality embodied in the idiom that states ‘unyabe ngathi utye umqha – you are as docile as if you ate a tree hyrax’. this saying may even dissuade many from actively hunting the tree hyrax. a study in the mau forest of kenya on the tree hyrax reported similar low levels of hunting (milner 1994). in the udzungwa mountains in tanzania (where the tree hyrax is found), bush meat comprises little of the meat intake of the locals, but when the local fauna is hunted larger fauna are preferred, provided they are available (nielsen 2006). low income and educational levels explain the reliance of the communities on the forest as a source of additional income and for cultural practices, with them obtaining firewood, building material and medicines from the forest (dold & cocks 2012). most participants indicated that harvesting is both for the generation of income and for personal use. the fact that participants indicated that most of the natural material obtained is for medicinal purposes suggests the importance of traditional medicine in these communities (figure 3d); it serves as the primary source of healthcare in the province (dold & cocks 2012; makunga, philander & smith 2008). participants indicated that they mainly use the bark (figure 3c), which could be problematic as in some trees ring-barking often leads to the death of the tree and thus extensive harvesting of bark may lead to a decrease in tree density (e.g. delvaux, sinsin & van damme 2010). there were few indications of ring-barking in the pirie forest (e.j.o., m.i.c., n.p.m. pers. obs., 13–17 july 2015), but this merits further investigation. some of the respondents indicated that they were aware that forest guards were monitoring the practices of harvesting associated with the pirie forest. the respondents also informed us that forest guards provided education about the forest ecosystem in an attempt to promote sustainable harvesting practices in the pirie forest. regarding the frequency of harvesting, it is evident that villagers go out harvesting depending on when they need the material, with a low frequency of participants saying that they harvest daily (figure 3b). most participants reported harvesting between 1 kg and 25 kg per trip, which amounts to what a person is able to carry either in a rucksack or on their head. some participants remarked that they harvest only as much as they need; others said that they harvested only pieces of bark. this can be in accordance with the theza load allowed in forests, provided that only dead or fallen trees are harvested and no cutting implements are employed. however, three participants declared taking more, with one mentioning 70 kg, one 500 kg and one saying he takes a tractor load per month. the participant who reported 500 kg harvested dead trees (mainly used for kraals) and the one who indicated a tractor load felled the trees and sold them, although this comprised mainly the invasive species, aca. mearnsii. this particular species was often the first species mentioned by respondents and this may be linked to the fact that harvesting alien invasives is not only legal but also actively promoted by nature conservation agencies. the tree listed most was acacia owing to its use as firewood, followed by p. obliquum, which is used medicinally, culturally and as building material. the latter is considered a sacred tree by the amaxhosa owing to it being symbolically linked to the ancestors (dold & cocks 2012) and is currently classified as being of least concern according to the south african red data list (raimondo et al. 2009). other trees mentioned in high frequencies were c. dentata, eucalyptus, o. europaea and s. henningsii. these plants also had the highest rfc values, indicating their importance in the community. of these trees, c. dentata is used as building material as well as for medicinal and spiritual purposes (dold & cocks 2012). for the continued ethnobotanical use of this particular tree, harvesting of c. dentata is of concern, as it has a ‘near threatened’ status according to the red data list (raimondo et al. 2009) and thus the high levels of harvesting of this species are problematic. the extent of harvesting of this tree species needs to be monitored. worthy of mention is the medicinal utilisation of rapanea melanophloeos, hypoxis hemerocallidea (african potato), ilex mitis (cape holly) and cassipurea flanaganii (cape onionwood). these species, although having low rfc values (figure 4), were all recorded in the medicinal plant trade and in addition c. flanaganii is listed as endangered, whilst r. melanophloeos, h. hemerocallidea and i. mitis are all listed as declining (raimondo et al. 2009; williams, victor & crouch 2013). it is therefore important to place restrictions on trade of these species, which could be compensated for by utilising common species with the same active compounds. furthermore, some of the respondents indicated that they are conservative in the quantities that they harvest for their personal subsistence but that gatherers and medicinal plant vendors from other areas or villages distant from pirie harvest illegally at night and often in large quantities for the medicinal trade. villagers were aware of the risk of unsustainable utilisation of forest resources. all the participants had a high appraisal of the socio-psychological experiences linked to activities that take them into the forest, indicating that knowledge and utilisation of natural products collected from the forest is an important aspect of their cultural identity that should be continued by future generations. hunting that occurs is mainly opportunistic and is unlikely to have a significant impact on hyrax populations. the expanse of the forest is perceived to have undergone little change, but changes to the quality of the forest need monitoring as local communities and commercial activities may lead to ecosystem changes. disturbances in the form of harvesting and hunting may have an effect on hyrax populations, as these activities could potentially alter the canopy, leading to a reduction in the capability of the hyrax to avoid natural predators. although commercial harvesting in the forest has remained at a constant level since 2007 (mpisekaya & kameni 2007), the impact of this harvesting on the hyrax population is unknown and should be monitored in future. although labour intensive, this can be accomplished by monitoring den occupation by ascertaining the presence or absence of hyrax faeces in tree hollows, and the estimate of gaylard (1994), using this technique, provides a baseline against which population changes can be measured. conclusion of the tree species used by the hyrax as dens in the pirie forest, a. falcatus, s. latifolia, a. ovalis, t. natalensis and a. dimidiata are important resources for local communities. as these are harvested at relatively low levels, it is unlikely that current harvesting has a large impact on the tree hyrax. we recommend ongoing monitoring of the pirie forest tree hyrax population in order to ascertain the impact of both commercial and community harvesting over the past quarter century. acknowledgements we would like to thank the interviewees from the communities surrounding the pirie forest and the herbal vendors interviewed in king williams town. mr mzonke ‘ronnie’ makhahla and mr khanyisa mzamo are thanked for their assistance and ms bukiwe mzamo is thanked for translating the questionnaire into isixhosa. this study was funded by a bursary to e.j.o. from the sasol inzalo foundation; running expenses were provided by grants (96257) to m.i.c. and to n.p.m. (76555) from the national research foundation of south africa. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions e.j.o. and n.p.m. conducted interviews, whereas e.j.o. analysed the data. all the authors wrote the manuscript. references brown, k., 2003, ‘trees, forests and communities’: some historiographical approaches to environmental history on africa’, area 35(4), 343–356. butynski, t., hoeck, h. & de jong, y.a., 2015, dendrohyrax arboreus. the iucn red list of threatened species. 2015: e.t6409a21282806, intrenational union for the conservation of nature, cambridge, uk. https://doi.org/10.2305/iucn.uk.2015-2.rlts.t6409a21282806.en castley, j.g. & kerley, g.i.h., 1996, ‘the paradox of forest conservation in south africa’, forest ecology and management 85, 35–46. https://doi.org/10.1016/s0378-1127(96)03748-6 cooper, t.j.g., wannenburgh, a.m. & cherry, m.i., 2017, ‘atlas data indicate forest dependent bird species declines in south africa’, bird conservation international 27, 337–354. https://doi.org/10.1017/s095927091600040x cunningham, a.b. & davis, c.w., 1997, ‘human use of plants’, in r.m. cowling, d.m. richardson & s.m. pierce (eds.), vegetation of southern africa, pp. 477–501, cambridge university press, cambridge, uk. delvaux, c., sinsin, b. & van damme, p., 2010, ‘impact of season, stem diameter and intensity of debarking on survival and bark re-growth pattern of medicinal tree species, benin, west africa’, biological conservation 143, 2664–2671. https://doi.org/10.1016/j.biocon.2010.07.009 dold, a.p. & cocks, m.l., 1999, ‘preliminary list of xhosa plant names from eastern cape, south africa’, bothalia 29, 267–292. https://doi.org/10.4102/abc.v29i2.601 dold, a.p. & cocks, m.l., 2012, voices from the forest: celebrating nature and culture in xhosaland, craft print international ltd., auckland park, south africa. gaylard, a., 1994, ‘the tree hyrax (dendrohyrax arboreus) as a rare forest species: habitat and diet’, msc thesis, university of port elizabeth. gaylard, a. & kerley, g.i.h., 1997, ‘diet of tree hyraxes dendrohyrax arboreus (hyracoidea: procaviidae) in the eastern cape, south africa’, journal of mammalogy 78, 213–221. https://doi.org/10.2307/1382654 gaylard, a. & kerley, g.i.h., 2001, ‘habitat assessment for a rare, arboreal forest mammal, the tree hyrax dendrohyrax arboreus’, african journal of ecology 39, 205–212. https://doi.org/10.1046/j.0141-6707.2000.301.x gaylard, a., venter, j., ehlers-smith, y. & child, m.f., 2016, ‘a conservation assessment of dendrohyrax arboreus’, in m.f. child, l. roxburgh, e. do linh san, d. raimondo & h.t. davies-mostert (eds.), the red list of mammals of south africa, swaziland and lesotho, south african national biodiversity institute and endangered wildlife trust, south africa, viewed 22 june 2017, from https://www.ewt.org.za/reddata/pdf/hyracoidea/2016%20mammal%20red%20list_dendrohyrax%20arboreus_en.pdf google earth 7.1.1.1888, 2013, pirie main forest reserve 32°46’59”s, 27°13’0”e elevation: 686 m, viewed 04 march 2015, from https://earth.google.com/web/@-32.84609917,27.47770818,506.94577214a,77550.7433243d,35y,0.00000009h,2.62787946t,-0r/data=cksasrjdciuwedflnjzlmte2zdvkowq3yju6mhg5ztfjmwe0njqzndrmnthmgr5cpsavgkdaiy50bkze3dtakghozxcgumvzdbgbiae greyling, t. & huntley, b.j., 1984, directory of southern african conservation areas, south african national scientific progress report no. 98, council for scientific and industrial research, pretoria, 311 p. lawes, m.j., mealin, p.e. & piper, s.e., 2000, ‘patch occupancy and potential metapopulation dynamics of three forest mammals in fragmented afromontane forests in south africa’, conservation biology 14, 1088–1098. https://doi.org/10.1046/j.1523-1739.2000.99120.x lawes, m.j., midgley, j.j. & chapman, c.a., 2004, ‘south africa’s forests: the ecology and sustainable use of indigenous timber resources’, in m.j. lawes, h.a.c. eeley, c.m. shackleton & b.g.s. geach (eds.), indigenous forests and woodlands in south africa: policy, people and practice, pp. 31–75, university of kwazulu-natal press, scottsville. low, a.b. & rebelo, a.g., 1996, vegetation of south africa, lesotho and swaziland, department of environmental affairs and tourism, pretoria. makunga, n.m., philander, l.e. & smith, m., 2008, ‘current perspectives on an emerging formal natural products sector in south africa’, journal of ethnopharmacology 119, 365–375. https://doi.org/10.1016/j.jep.2008.07.020 milner, j., 1994, ‘relationships between the forest-dwelling people of south-west mau and tree hyrax, dendrohyrax arboreus’, journal of east african natural history 83, 17–29. https://doi.org/10.2982/0012-8317(1994)83[17:rbtfdp]2.0.co;2 mpisekaya, s.r. & kameni, c., 2007, amatole forest yellowwood harvesting levels, department of water affairs and forestry, pretoria. nielsen, m.r., 2006, ‘importance, cause and effect of bushmeat hunting in the udzungwa mountains, tanzania: implications for community based wildlife management’, biological conservation 128, 509–516. https://doi.org/10.1016/j.biocon.2005.10.017 raimondo, d., von staden, l., foden, w., victor, j.e., helme, n.a., turner, r.c. et al. (eds.), 2009, ‘red list of south african plants 2009’, in strelitzia, p. 25, south african national biodiversity institute, pretoria. samoisy, a.k. & mahomoodally, m.f., 2015, ‘ethnopharmacological analysis of medicinal plants used against non-communicable diseases in rodrigues island, indian ocean’, journal of ethnopharmacology 173, 20–38. https://doi.org/10.1016/j.jep.2015.06.036 topp-jørgensen, j.e., marshall, a.r., brink, h. & pederson, h.b., 2008, ‘quantifying the response of tree hyraxes (dendrohyrax validus) to human disturbance in the udzungwa mountains, tanzania’, tropical conservation science 1, 63–74. https://doi.org/10.1177/194008290800100106 von maltitz, g., mucina, l., geldenhuys, c., lawes, m., eeley, h. & adie, h., 2003, classification system for south african indigenous forests: an objective classification for the department of water affairs and forestry, environmental report: 1-284, council for scientific and industrial research, pretoria. williams, v.l., victor, j.e. & crouch, n.r., 2013, ‘red listed medicinal plants of south africa: status, trends and assessment challenges’, south african journal of botany 86, 23–25. https://doi.org/10.1016/j.sajb.2013.01.006 appendix 1 questionnaire gender:        [male] [female] age:          [18–29] [30–39] [40–49] [50–59] [60–69] [70+] education:       [grade 7 or below] [grade 10] [matric] [bachelors] [more] birthplace: race:          [african] [mixed race] [white] [indian or asian] [other] income per month:   [less than r500] [r500–r1000] [r1000–r3000] [r3000–r10 000] [more than r10 000] occupation: do you harvest in the pirie forest?  [yes] [no] which trees do you harvest? what part of the tree do you harvest? how often do you harvest?   [yearly] [twice a year] [every few months] [monthly]                  [twice a month] [weekly] [daily] how much do you harvest at a time? what do you use the products you harvest for? are the products you get from harvesting only for personal use or do you sell them as well?   [sell them]  [personal use]  [both] since what age have you been harvesting in the forest? who taught you to harvest and use the product like that? do you see the forest as a resource that can be renewed?   [yes] [no] what do you know about the tree hyrax present in the forest? do you hear the tree hyrax in the forest?            [yes] [no] have you ever seen the tree hyrax and do you still see them around? do you hunt the tree hyrax or know of someone who does?   [yes] [no] if you answered ‘yes’ to the previous question, what do you and/or they hunt the tree hyrax for? do you know anyone else i can talk to about this? thank you so much for your time and patience. i will come back to you with the results of this survey. abstract introduction research methods and design results discussion conclusion acknowledgements references appendix 1 about the author(s) chad keates department of zoology and entomology, rhodes university, makhanda, south africasouth african institute for aquatic biodiversity (saiab), makhanda, south africa werner conradie port elizabeth museum, humewood, south africadepartment of nature conservation management, natural resource science and management cluster, faculty of science, nelson mandela university, george, south africa tatenda dalu school of biology and environmental sciences, university of mpumalanga, mbombela, south africa farai dondofema department of ecology and resource management, university of venda, thohoyandou, south africa eddie s. riddell kruger national park, south african national parks, skukuza, south africacentre for water resources research, university of kwazulu-natal, pietermaritzburg, south africa ryan j. wasserman department of zoology and entomology, rhodes university, makhanda, south africa south african institute for aquatic biodiversity (saiab), makhanda, south africa citation keates, c., conradie, w., dalu, t., dondofema, f., riddell, e.s. & wasserman, r.j. 2022, ‘phylogenetic placement of the enigmatic floodplain water snake, lycodonomorphus obscuriventris fitzsimons, 1964’, koedoe 64(1), a1698. https://doi.org/10.4102/koedoe.v64i1.1698 original research phylogenetic placement of the enigmatic floodplain water snake, lycodonomorphus obscuriventris fitzsimons, 1964 chad keates, werner conradie, tatenda dalu, farai dondofema, eddie s. riddell, ryan j. wasserman received: 07 oct. 2021; accepted: 08 feb. 2022; published: 28 june 2022 copyright: © 2022. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract lycodonomorphus is a genus of lamprophiid water snake endemic in africa. although widespread, abundant and presumably an important component of many aquatic and semi-aquatic food webs, these snakes are poorly understood taxonomically, particularly from a phylogenetic perspective. with only four of the nine species currently sequenced, this study attempts to improve our understanding of the evolutionary relationships within the genus through the phylogenetic placement of one of the most elusive species, lycodonomorphus obscuriventris. collected in the ramsar declared makuleke wetlands in northern kruger national park (south africa), the sample used in this study not only yielded the first dna sequences for the taxon but also represented the most northerly south african record, bridging the gap between the southern and northern populations. the snake was sequenced for three partial mitochondrial genes (16s, cyt-b, nd4) and one partial nuclear gene (c-mos) and phylogenetically placed, relative to the rest of the genus, using maximum likelihood (ml) and bayesian inference (bi). sequence divergences between sister taxa were also estimated using pairwise distance analysis. the concatenated phylogenetic reconstruction yielded similar topological structuring when compared to phylogenies from past articles, with both the ml and bi algorithms recovering strong support for l. obscuriventris as sister to a clade comprising of l. whytii + l. laevissimus + l. rufulus. the phylogenetic placement, albeit based on a single sample, challenges the original placement (morphological) of l. obscuriventris as sub-specific within l. whytii, suggesting that multiple species concepts should be considered when delineating species within this group. conservation implications: prior to the discovery of the new record, the global distribution of l. obscuriventris was characterised by two disjunct populations. the new record bridges the distribution gap between these two populations, rendering the distribution continuous. this bodes well for the species as there is likely no barrier to gene flow, thereby buffering the species from localised threats given the more expansive distribution. furthermore, given that the specimen was sampled from the kruger national park, the species is likely to be well-protected as much of its distribution within south africa seems to fall within protected areas. keywords: lamprophiidae; molecular systematics; water snake; range expansion; wetlands; african herpetology; kruger national park; southern africa. introduction lycodonomorphus is an endemic genus of medium-sized, semi-aquatic-to-aquatic lamprophiids occurring in south-central africa that are characterised by their small heads, which are virtually indistinguishable from the neck (branch 1998; broadley 1983; broadley & blaylock 2013; broadley, doria and wigge 2003; marais 2004; pietersen, verburgt & davies 2021; spawls et al. 2018; wallach, williams & boundy 2014). all species within the genus are oviparous, harmless to humans and do not possess enlarged fangs or venom glands (branch 1998; broadley 1983; broadley et al. 2003; marais 2004; spawls et al. 2018). whilst mostly nocturnal, several species in the genus forage actively during the day (branch 1998; broadley 1983; kyle, alexander & du preez 2021). little is known about the ecology of these presumably important predators and their trophic role in aquatic habitats is largely underexplored (madsen & osterkamp 1982). several studies (e.g. kyle et al. 2021; madsen & osterkamp 1982; raw 1973; taylor 1970) have, however, reported members of the genus as feeding on fringes of water bodies for tadpoles, frogs, fish and other small vertebrates. some of the more aquatic species have also been observed to ambush fish from amongst submerged rocks within waterbodies (branch 1998). lycodonomorphus has undergone substantial taxonomic restructuring since the last major revision by loveridge (1958), in which he only recognised four species with six subspecies. the genus currently contains nine accepted species (uetz et al. 2022): l. bicolor (günther, 1893), l. inornatus (duméril, bibron & duméril, 1854), l. laevissimus (günther, 1862), l. leleupi (laurent, 1950), l. mlanjensis loveridge, 1953, l. obscuriventris fitzsimons, 1964, l. rufulus (lichtenstein, 1823), l. subtaeniatus laurent, 1954 and l. whytii (boulenger, 1897). raw (1973) further described two subspecies, l. laevissimus natalensis and l. laevissimus fitzsimonsi, which were later synonymised with the nominate form (haagner & branch 1994). although currently recognised as a full species, the species has a complex origin (broadley 1967, 1983, 1995; rasmussen 2004). it was originally described as a subspecies of l. whytii based on the very dark ventrum, compared to the uniform immaculate white in the nominal form (fitzsimons 1964). up until this point, however, l. whytii was considered a subspecies of l. rufulus (loveridge 1958), necessitating the elevation of l. whytii to species level to accommodate the sub-specific recognition of l. w. obscuriventris. when broadley (1967) reviewed newly collected material of l. whytii from central mozambique, he found the ventral colouration to be variable and thus regarded them as the same species, although he did record differences in ventral and subcaudal scale counts. based on this, scale differences and different habitat preferences, broadley (1983) tentatively proceeded and recognised the southern material, including central mozambique material, as l. w. obscuriventris. the discovery of a specimen that conforms to l. w. obscuriventris near the boundary of lengwe game reserve in southern malawi in 1995 raised interest in the taxonomical relationship between the two subspecies, especially because the type locality of the nominal form is from fort hill (= chitipa) in northern malawi (broadley 1995). based on morphological data gathered from the three l. w. whytii (1 malawi, 2 tanzania [all females]) type specimens available at the time, it was noted that l. w. whytii had lower ventral scale counts, higher subcaudal scale counts, it lacked the distinctive labial markings and displayed different habitat preferences (montane streams versus lowland floodplains) when compared to l. w. obscuriventris (see broadley 1995). these results were expanded on with the incorporation of four additional specimens (including a male) of the nominal form by rasmussen (2004). based on these different traits, l. obscuriventris was then elevated to full species. given the lack of phylogenetic work on the taxon, the species is considered related to l. whytii, given its past sub-specific placement and shared morphology (broadley 1967, 1983, 1995; fitzsimons 1964; rasmussen 2004). although relatively widespread (figure 1), with a distribution that stretches from eastern kwazulu-natal province (south africa) into eswatini, kruger national park, zimbabwe and northern mozambique into southern malawi (broadley 1983, 1995; brown & wilkey 2019; kyle et al. 2021), l. obscuriventris remains elusive and poorly understood because of its original description. figure 1: (a) current global distribution of lycodonomorphus obscuriventris with black triangles denoting historical localities and the red triangle denoting the new sample used in this study from the makuleke contractual national park. (b) full body photograph of pem r27786. (c) head shot of pem r27786. aim and objectives whilst the group has received much attention from traditional taxonomists (morphology), genetic work on lycodonomorphus is severely lacking, with no dedicated phylogenetic study of the genus to date. the phylogenetic work that has included lycodonomorphus spp. mainly focussed on the higher-level taxonomy (pyron, burbrink & wiens 2013; vidal et al. 2008; zaher et al. 2019) and other closely related genera within lamprophiidae such as lamprophis, boaedon and file snakes (branch et al. 2019; broadley et al. 2018; ceríaco et al. 2021; greenbaum et al. 2015; hallermann et al. 2020; keates et al. 2019; kelly et al. 2008, 2011). only limited representative sampling has been used for members of lycodonomorphus, meaning only four of the nine species have been sequenced. these include l. rufulus, l. inornatus, l. laevissimus and l. whytii, with the last two species only having one sequence each (https://www.ncbi.nlm.nih.gov/genbank/). as stated in vidal et al. (2008), the uncertainty over species boundaries within the lycodonomorphus whytii-mlanjensis-obscuriventris complex needs to be addressed prior to the onset of systematic reordering in the group to ensure that taxonomical redundancies (e.g. synonyms, homonyms) are not incorporated into the nomenclature. to this end, we endeavour to improve our understanding of the phylogenetics of the group through the phylogenetic placement of l. obscuriventris using a recently acquired genetic sample of the species. research methods and design sample site and data collection in early-april 2021, a multidisciplinary group of biologists embarked on a field survey of the ramsar declared makuleke wetlands, in northern kruger national park, south africa. the kruger national park itself forms a part of the great limpopo transfrontier park shared with zimbabwe and mozambique. the hydro-geomorphic setting of the seasonal pans was within the flood plain of the limpopo and luvuvhu river systems. the trip was focussed on surveying the food web dynamics of the temporary and permanent pans. on the 8th of april 2021, an inactive snake was discovered concealed beneath the bark of a large dead tree trunk lying on a small island in banyini pan. the snake was discovered approximately 2 km from the zimbabwean border on the western limits of the makuleke contractual national park (–22.365750, 31.075306) (figure 1). the specimen was removed from the log and identified as l. obscuriventris based on colouration and more specifically the yellow upper labials (marais 2004). the specimen was humanely euthanised by placing it in a solution of clove oil until it was dead. subsequently, a liver sample was dissected out for genetic analysis, and preserved in 99% ethanol. the specimen was then fixed in 10% formalin for 72 h after which it was transferred into 70% ethanol for long-term storage in the herpetological collection at the port elizabeth museum (pem), south africa. the male specimen was catalogued under the number pem r27786 and measured: 318 mm snout-vent length and 72 mm tail length (the terminal tip was slightly truncated). data analysis dna extraction, amplification and sequencing dna was isolated from the tissue sample with a standard salt extraction method (bruford et al. 1992) using lysis (buffer atl) and elution (buffer ae) buffers. standard polymerase chain reaction (pcr) procedures were utilised to amplify one partial mitochondrial ribosomal gene (ribosomal ribonucleic acid [16s]), two partial mitochondrial genes (cytochrome b [cyt-b] and nadh-dehydrogenase subunit 4 [nd4]) and one partial nuclear gene (oocyte maturation factor [c-mos]) (table 1). the pcr amplification was carried out using the primer pairs listed in table 1. amplification of the selected genes was carried out using 20 ng/μl – 50 ng/μl extracted genomic dna. each amplification was conducted with a pcr mixture to the total volume of 25 μl containing 12.5 μl taq dna polymerase 2x master mix (ampliqon; 3 mm mgcl2, 0.4 mm dntps and ampliqon taq dna polymerase), 2 μl forward primer (10 μm), 2 μl reverse primer (10 μm), 6.5 μl denucleated water and 2 μl genomic dna. the cycling profile for all the genes was as follows: initial denaturing step at 94 °c for 5 min, followed by 35 cycles of 94 °c for 30 s, 50 °c – 56 °c for 45 s and 72 °c for 45 s, with a final extension at 72 °c for 8 min. the prepared pcr products were sent to macrogen corp. (amsterdam, netherlands) for sequencing (after purification) with the forward primers only. table 1: primers and pcr protocols used to generate sequences for the study. the phylogenetic placement of l. obsuriventris was estimated by comparing the genetic information of the newly collected sample with published sequences from all currently sequenced species of lycodonomorphus: four l. rufulus, one l. laevissimus, one l. whytii and seven l. inornatus (appendix 1). in addition to the ingroup taxa, the dataset was supplemented with sequences from closely related genera that were obtained from genbank, to root the tree (appendix 1). phylogenetic analysis the sequence trace files were checked using bioedit sequence alignment editor v.7.2.5 (hall 1999) and aligned with accessioned genbank sequences using mega v.6.0 (tamura et al. 2013) and the clustalw alignment method. prior to further analysis, the hyper-variable region of 16s was removed. four individual alignments were created and used to construct individual gene trees using the maximum likelihood (ml) algorithm (100 bootstrap replicates), and the gtr + g + i nucleotide substitution model. the individual consensus trees were used to determine the congruence of the topologies of the different genetic markers using the congruence index (icong; http://max2.ese.u-psud.fr/icong/index.help.html; vienne, giraud & martin 2007). all gene-tree combinations were found to be congruent and a concatenated dataset of the four genes was created for additional phylogenetic analyses. saturation was tested in dambe v.6.4.67 (xia 2013) using the individual as well as the combined first and second codon positions of each gene. saturation was absent from every marker, necessitating the use of a gene-partitioned dataset for the phylogenetic reconstruction. the optimal partition scheme and best-fitting models of molecular evolution were selected using iq-tree v.2.1.2 (minh et al. 2021) with the following settings: p-partition file (each partition has its own evolution rate), a greedy strategy and the freerate heterogeneity model excluded (only invariable site and gamma rate heterogeneity considered) (chernomor, von haeseler & minh 2016; kalyaanamoorthy et al. 2017). the greedy strategy implemented in modelfinder via iq-tree v.2.1.2 resembles the one used in partitionfinder 2 (lanfear et al. 2016), in the way it starts the full partition model and subsequently merges two genes until the model fit no longer increases (minh et al. 2021). the best-fitting model scheme selected included the following three partitions and models of evolution: tim2 + i + g (16s); tim2 + i +g (cyt-b, nd4); gtr + i + g (c-mos). mrbayes v.3.2.7a (ronquist et al. 2012) was not able to implement tim2, so the next best alternative (gtr) was used in its place. phylogenetic tree and p-distance analysis maximum likelihood (ml) analysis was conducted using iq-tree v.2.1.2 (nguyen et al. 2015). a random starting tree was used, and the ml analysis was assessed using the gene-partitioned scheme mentioned above and 1000 standard nonparametric bootstraps. bayesian inference (bi) analysis (mrbayes v.3.2.7a; ronquist et al. 2012) was implemented on the cipres science gateway xsede online resource (http://www.phylo.org; miller et al. 2010; tamura et al. 2013) using the best-fit nucleotide substitution models and partition scheme listed above. two parallel runs of 20 million generations were performed, with trees being sampled every 1000 generations using beagle (high performance likelihood calculation library). psammophylax rhombeatus was used as an outgroup as only a single sample can be used as an outgroup using mrbayes. the number of generations discarded as burn-in was determined using tracer v.1.6.0. (rambaut & drummond 2007). the effective sample size (ess) was above 200 for all parameters and the runs reached convergence, indicating that a burn-in of 15% was adequate. both trees were viewed in figtree v.1.4.2 (rambaut 2014). pairwise distance analysis was conducted in mega x (kumar et al. 2018) using the cyt-b alignment from the phylogenetic reconstruction. sequences were grouped according to species and pairwise distance analysis was conducted on mega x, using uniform rates, pairwise deletion and 500 bootstrap replicates. ethical considerations all procedures performed in this study followed all international, national and/or institutional guidelines for the care and use of animals. ethical permission was acquired from the university of venda (reference number: ses/18/erm/10/1009) and sample collection permits were acquired from the kruger national park (reference number: skz 132). results both phylognetic algorithms (ml and bl) showed strong support for the monophyly of lycodonomorphus (bootstrap probability [bp] 91%, posterior probability [pp] 1.0) (figure 2). lycodonomorphus obscuriventris was recovered as sister to l. whytii + l. laevissimus + l. rufulus, with strong support from both phylogenetic algorithms (bp 89%, pp 1.0). furthermore, both algorithms recovered a supported sister relationship between l. rufulus and l. whytii + l. laevissimus. both l. rufulus and l. inornatus were characterised by substantial topological sub-structuring with both algorithms recognising two supported clades in both species. using cyt-b, the average pairwise distance separating species within the genus was 15.88 ± 2.04% (mean ± standard deviation), whilst the average pairwise distance separating l. obscuriventris from other members of lycodonomorphus was 15.87% ± 0.90% (table 2). the lowest intrageneric pairwise distance separating l. obscuriventris was observed in l. rufulus (15.23%; table 2). the structured clades observed within l. inornatus and l. rufulus (figure 2) were supported by pairwise distance analysis with intraspecific divergences of 6.77% and 3.89%, respectively. due to the lack of sampling for l. laevissimus, l. whytii and l. obscuriventris, intraspecific divergences could not be assessed for these species. figure 2: maximum likelihood phylogenetic reconstruction with bayesian inference support overlaid. table 2: sequence divergences (uncorrected pairwise distance values) separating the species of lycodonomorphus and lamprophis using cytochrome b (cyt-b). discussion aside from the addition of l. obscuriventris, the phylogenetic reconstruction of lycodonomorphus yielded identical topological structuring when compared to past publications (kelly et al. 2011; vidal et al. 2008; zaher et al. 2019). whilst only a single sample was available, the strong phylogenetic support (ml and bi) coupled with the divergent pairwise distance values separating l. obscuriventris from its congeners ratified the assertions of broadley (1995) that the species is valid. whilst the species was elevated out of the synonymy of l. whytii based on morphological and geographical grounds, the recovery of l. obscuriventris as sister to l. laevissimus + l. rufulus + l. whytii (figure 2) suggests that the original sub-specific assignment of l. obscuriventris was erroneous and the current taxonomy reflects correct nomenclature. the sister relationship between l. laevissimus and l. whytii is interesting to note given the large geographical distance separating the two species. lycodonomorphus laevissimus is restricted to eastern south africa, lesotho and eswatini, whilst l. whytii is found in northern malawi and tanzania (branch 1998; rasmussen 2004; wallach et al. 2014). these findings would suggest that geographical proximity at least in southern africa plays a reduced role in evolutionary relatedness. the topological structuring observed (figure 2) may be better explained by the habitat preferences and associated ecologies of the different species with l. obscuriventris showing an affinity to lowland floodplains and pans, whilst l. laevissimus prefers slow moving streams, l. whytii prefers upland montane streams and l. rufulus being more generalist (branch 1998; rasmussen 2004). strengths and limitations whilst robust sampling was absent for many of the species in this study (only one sample each for l. obscuriventris, l. whytii and l. laevissimus), both l. rufulus and l. inornatus displayed intraspecific substructuring consistent with geographical variability. within l. inornatus, the intraspecies diversity was 6.77% for cyt-b. whilst higher than the other species of lycodonomorphus, it must be noted that seven samples (kelly et al. 2011) were available for this study, five from haenertsberg and two from the eastern cape. the large geographical distance separating the samples may explain the increased intraspecies diversity found with l. inornatus. the addition of new samples, representing the full distribution of the species, would thus likely support the recognition of l. inornatus as a single species, especially because the average interspecific divergence separating species is approximately 16%. implications or recommendations in a recent study, greenbaum et al. (2015) elevated the poorly known lycodonomorphus subtaeniatus upembae (laurent 1954) to full species status and transferred it to the genus boaedon, whilst lycodonomorphus subtaeniatus was retained, pending further phylogenetic evidence. the findings from this article coupled with the small morphological characteristics (mainly the lower number of midbody scale rows, simpler unforked-to-weakly forked hemipenis, no diastema separating maxillary teeth) separating lycodonomorphus from boaedon (kelly et al. 2011) casts doubt on the validity of lycodonomorphus, especially because several members of the genus show a strong terrestrial affinity (as opposed to being aquatic). for this reason, it is recommended that future work on the group endeavours to sequence all the species associated with the genus to determine the most accurate systematic structuring of the group. in addition to being the first sequenced sample of l. obscuriventris, the sample also represents the most westerly located record for the species and the most northerly located record for the kruger national park. the species’ previous most northern record (within the kruger national park) was near the border of eastern mozambique, approximately 30 km north of the letaba river in the central kruger national park (pienaar 1976). the newly collected specimen was found in the western reaches of the makuleke contractual park approximately 150 km north of the previous most northern kruger national park record. conclusion prior to the discovery of the new records, the known distribution of l. obscuriventris was characterised by two disjunct populations, with a southerly population in eastern south africa and eswatini and a northern population in zimbabwe, mozambique and malawi (broadley, 1983, 1995). this sample, thus fills the gap between the two populations meaning that the distribution is likely continuous. additionally, the recovery of the snake in the northern kruger national park bodes well for the conservation of the species as a large portion of its south african distribution seems to fall within this protected area, although this may be a result of sampling bias. given the large proportion of records for this species inside the kruger national park, future effort should be directed to adjacent areas, including the great limpopo transfrontier park regions, to elucidate more clearly the ecology, biology and phylogeographic structuring of the enigmatic and elusive water snake. acknowledgements the authors would like to thank south african national parks, and more specifically, the staff of kruger national park and the makuleke contractual national park for facilitating this research. without their dedicated managers, section rangers and field guides, this project would not have been possible. competing interests the authors would like to declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions c.k., r.j.w., t.d. and w.c. conceived the study. c.k. carried out lab work and genetic analyses. all authors contributed to the interpretation of the results. t.d., f.d. and e.s.r. arranged permitting and all other necessary permissions. t.d., f.d., r.w. and w.c. provided funding for the project. c.k. wrote the first draft of the manuscript. all authors provided a critical feedback and helped shape subsequent drafts of the manuscript. funding information this research was funded by the university of venda (niche grant [ses/18/erm/10]) and university of mpumalanga (institutional research themes support grant). data availability the data that support the findings from this study are openly available on genbank (https://www.ncbi.nlm.nih.gov/genbank/) and can be downloaded using the unique genbank numbers found in appendix 1. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references arevalo, e., davis, s.k. & sites, j.w., 1994, ‘mitochondrial dna sequence divergence and phylogenetic relationships among eight chromosome races of the sceloporus grammicus complex (phrynosomatidae) in central mexico’, systematic biology 43(3), 387–418. https://doi.org/10.1093/sysbio/43.3.387 branch, w.r., 1998, field guide to the snakes and other reptiles of southern africa, struik, cape town. branch, w.r., baptista, n., keates, c. & edwards, s., 2019, ‘rediscovery, taxonomic status, and phylogenetic relationships of two rare and endemic snakes (serpentes: psammophiinae) from the southwestern angolan plateau’, zootaxa 4590(3), 342–366. https://doi.org/10.11646/zootaxa.4590.3.2 broadley, d.f., doria, c.t. & wigge, j., 2003, snakes of zambia: an atlas and field guide, chimaira, frankfurt. broadley, d.g., 1967, ‘a review of the genus lycodonomorphus fitzinger (serpentes: colubridae) in southeastern africa, with a key to the genus’, arnoldia 3(16), 1–9. broadley, d.g., 1983, fitzsimons’ snakes of southern africa, delta books, johannesburg. broadley, d.g., 1995, ‘a small collection of reptiles and amphibians from central and southern africa’, african herp news 24, 16–18. broadley, d.g. & blaylock, r., 2013, the snakes of zimbabwe and botswana, chimaira, frankfurt. broadley, d.g., tolley, k.a., conradie, w., wishart, s., trape, j.-f., burger, m. et al., 2018, ‘a phylogeny and genus-level revision of the african file snakes gonionotophis boulenger (squamata: lamprophiidae)’, african journal of herpetology 67(1), 43–60. https://doi.org/10.1080/21564574.2018.1423578 brown, g. & wilkey, r., 2019, reptiles of malawi: a photographic guide to 145 species, unknown publisher. bruford, m.w., hanotte, m., brookfield, j.f.y. & burke, t., 1992, ‘single locus and multilocus dna fingerprint’, in a.r. hoelzel (ed.), molecular genetic analysis of populations: a practical approach, pp. 225–270, irl press, oxford. ceríaco, l.m., arellano, a.l., jadin, r.c., marques, m.p., parrinha, d. & hallermann, j., 2021, ‘taxonomic revision of the jita snakes (lamprophiidae: boaedon) from são tomé and príncipe (gulf of guinea), with the description of a new species’, african journal of herpetology 70(1), 1–31. https://doi.org/10.1080/21564574.2020.1832152 chernomor, o., von haeseler, a. & minh, b.q., 2016, ‘terrace aware data structure for phylogenomic inference from supermatrices’, systematic biology 65(6), 997–1008. https://doi.org/10.1093/sysbio/syw037 fitzsimons, v.f.m., 1964, ‘a new subspecies of water-snake from kruger national park’, koedoe 7(1), 26–29. https://doi.org/10.4102/koedoe.v7i1.796 greenbaum, e., portillo, f., jackson, k. & kusamba, c., 2015, ‘a phylogeny of central african boaedon (serpentes: lamprophiidae), with the description of a new cryptic species from the albertine rift’, african journal of herpetology 64(1), 18–38. https://doi.org/10.1080/21564574.2014.996189 haagner, g.v. & branch, w.r., 1994, ‘a taxonomic revision of the dusky-bellied water snake, lycodonomorphus laevissimus (serpentes: colubridae)’, journal of african zoology 108(3), 237–250. hall, t.a., 1999, bioedit: ‘a user-friendly biological sequence alignment editor and analysis program for windows 95/98/nt’, nucleic acids symposium series 41, 95–98. hallermann, j., ceríaco, l.m.p., schmitz, a., ernst, r., conradie, w., verburgt, l. et al., 2020, ‘a review of the angolan house snakes, genus boaedon duméril, bibron and duméril (1854) (serpentes: lamprophiidae), with description of three new species in the boaedon fuliginosus (boie, 1827) species complex’, african journal of herpetology 69(1), 1–50. https://doi.org/10.1080/21564574.2020.1777470 kalyaanamoorthy, s., minh, b.q., wong, t.k.f., von haeseler, a. & jermiin, l.s., 2017, ‘modelfinder: fast model selection for accurate phylogenetic estimates’, nature methods 14(6), 587–589. https://doi.org/10.1038/nmeth.4285 keates, c., conradie, w., greenbaum, e. & edwards, s., 2019, ‘a snake in the grass: genetic structuring of the widespread african grass snake (psammophylax fitzinger 1843), with the description of a new genus and a new species’, journal of zoological systematics and evolutionary research 57(4), 1039–1066. https://doi.org/10.1111/jzs.12337 kelly, c.m.r., barker, n.p., villet, m.h., broadley, d.g. & branch, w.r., 2008, ‘the snake family psammophiidae (reptilia: serpentes): phylogenetics and species delimitation in the african sand snakes (psammophis boie, 1825) and allied genera’, molecular phylogenetics and evolution 47(3), 1045–1060. https://doi.org/10.1016/j.ympev.2008.03.025 kelly, c.m., barker, n.p., villet, m.h. & broadley, d.g., 2009, ‘phylogeny, biogeography and classification of the snake superfamily elapoidea: a rapid radiation in the late eocene’, cladistics 25(1), 38–63. https://doi.org/10.1111/j.1096-0031.2008.00237.x kelly, c.m.r., branch, w.r., broadley, d.g., barker, n.p. & villet, m.h., 2011, ‘molecular systematics of the african snake family lamprophiidae fitzinger, 1843 (serpentes: elapoidea), with particular focus on the genera lamprophis fitzinger 1843 and mehelya csiki 1903’, molecular phylogenetics and evolution 58(3), 415–426. https://doi.org/10.1016/j.ympev.2010.11.010 kumar, s., stecher, g., li, m., knyaz, c. & tamura, k., 2018, ‘mega x: molecular evolutionary genetics analysis across computing platforms’, molecular biology and evolution 35(6), 1547–1549. https://doi.org/10.1093/molbev/msy096 kyle, k.j., alexander, g.j. & du preez, l.h., 2021, ‘reproduction, geographic distribution and habitat association of lycodonomorphus obscuriventris (serpentes: lamprophiidae)’, herpetology notes 14, 865–867. lanfear, r., frandsen, p.b., wright, a.m., senfeld, t. & calcott, b., 2016, ‘partitionfinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses’, molecular biology and evolution 34(3), 772–773. https://doi.org/10.1093/molbev/msw260 lawson, r., slowinski, j.b., crother, b.i. & burbrink, f.t., 2005, ‘phylogeny of the colubroidea (serpentes): new evidence from mitochondrial and nuclear genes’, molecular phylogenetics & evolution 37(2), 581–601. https://doi.org/10.1016/j.ympev.2005.07.016 loveridge, a., 1958, ‘revision of five african snake genera’, bulletin of the museum of comparative zoology 119, 1–198. madsen, t. & osterkamp, m., 1982, ‘notes on the biology of the fish-eating snake lycodonomorphus bicolor in lake tanganyika’, journal of herpetology 16(2), 185–188. https://doi.org/10.2307/1563817 marais, j., 2004, a complete guide to the snakes of southern africa, struik nature, cape town. miller, m.a., pfeiffer, w. & schwartz, t., 2010, ‘creating the cipres science gateway for inference of large phylogenetic trees’, in gateway computing environments workshop, new orleans, november, 14, 2010, pp. 1–8. minh, b.q., lanfear, r., trifinopoulos, j., schrempf, d. & schmidt, h.a., 2021, iq-tree version 2.1.2: tutorials and manual phylogenomic software by maximum likelihood, viewed n.d., from http://www.iqtree.org/doc/iqtree-doc.pdf. nguyen, l.t., schmidt, h.a., von haeseler, a. & minh, b.q., 2015, ‘iq-tree: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies’, molecular biology and evolution 32(1), 268–274. https://doi.org/10.1093/molbev/msu300 palumbi, s.r., 1996, ‘the polymerase chain reaction’, in d. hillis, c. moritz & b. mable (eds.), molecular systematics, 2nd edn., pp. 205–247. sinauer associates, sunderland, ma. pienaar, u.v., haacke, w.d., & jacobsen, n.h.g., 1976, the reptiles of the kruger national park, national parks board of south africa, pretoria. pietersen, d., verbught, l., & davies, j., 2021, snakes and other reptiles of zambia and malawi, struik nature, cape town. pyron, r.a., burbrink, f.t. & wiens, j., 2013, ‘a phylogeny and revised classification of squamata, including 4161 species of lizards and snakes’, bmc evolutionary biology 13, 93. https://doi.org/10.1186/1471-2148-13-93 rambaut, a., 2014, figtree, viewed 15 september 2021, from http://tree.bio.ed.ac.uk/software/figtree/. rambaut, a. & drummond, a.j., 2009, tracer, viewed 15 september 2021, from http://beast.bio.ed.ac.uk/tracer. rasmussen, j.b., 2004, ‘a review of whyte’s water-snake, lycodonomorphus whytii (serpentes: colubridae: lamprophiinae)’, african journal of herpetology 53(2), 155–162. https://doi.org/10.1080/21564574.2004.9635508 raw, l.r.g., 1973, ‘a review of the dusky-bellied water snake, lycodonomorphus laevissimus (günther), with description of two new subspecies’, annals of the natal museum 21(3), 713–718. ronquist, f., teslenko, m., van der mark, p., ayres, d.l., darling, a., hohna, s. et al., 2012 ‘mrbayes 3.2: efficient bayesian phylogenetic inference and model choice across a large model space’, software for systematics and evolution 61(3), 539–542. https://doi.org/10.1093/sysbio/sys029 slowinski, j.b. & lawson, r., 2002, ‘snake phylogeny: evidence from nuclear and mitochondrial genes’, molecular phylogenetics and evolution 24, 194–202. https://doi.org/10.1016/s1055-7903(02)00239-7 spawls, s., howell, k., hinkel, h. & menegon, m., 2018, a field guide to east african reptiles, 2nd edn., bloomsbury publicshing plc, london. tamura, k., stecher, g., peterson, d., filipski, a. & kumar, s., 2013, ‘mega6: molecular evolutionary genetics analysis version 6.0’, molecular biology and evolution 30(12), 2725–2729. https://doi.org/10.1093/molbev/mst197 taylor, m., 1970, ‘an observation on the feeding habits of lycodonmorphus rufulus’, african journal of herpetology 6(1), 19–20. https://doi.org/10.1080/04416651.1970.9650767 vidal, n., branch, w.r., pauwels, o.s.g., hedges, s.b., broadley, d.g., wink, m. et al., 2008, ‘dissecting the major african snake radiation: a molecular phylogeny of the lamprophiidae fitzinger (serpentes, caenophidia)’, zootaxa 66(1945), 51–66. https://doi.org/10.11646/zootaxa.1945.1.3 vienne, d.m., giraud, t. & martin, o.c., 2007, ‘a congruence index for testing topological similarity between trees’, bioinformatics 23, 3119–3124. https://doi.org/10.1093/bioinformatics/btm500 wallach, v., williams, k.l. & boundy, j., 2014, snakes of the world: a catalogue of living and extinct species, crc press, boca raton, fl. whiting, a.s., bauer, a.m. & sites, j.w., 2003, ‘phylogenetic relationships and limb loss in sub-saharan african scincine lizards (squamata: scincidae)’, molecular phylogenetics and evolution 29(3), 582–598. https://doi.org/10.1016/s1055-7903(03)00142-8 uetz, p., freed, p., aguilar, r. & hošek, j. (eds.), 2021, the reptile database, viewed 19 january 2021, from http://www.reptile-database.org. xia, x., 2013, ‘dambe5: a comprehensive software package for data analysis in molecular biology and evolution’, molecular biology and evolution 30(7), 1720–1728. https://doi.org/10.1093/molbev/mst064 zaher, h., murphy, r.w., arredondo, j.c., graboski, r., machado-filho, p.r., mahlow, k. et al., 2019, ‘large-scale molecular phylogeny, morphology, divergence-time estimation, and the fossil record of advanced caenophidian snakes (squamata: serpentes)’, plos one 14(5), e0217959. https://doi.org/10.1371/journal.pone.0217959 appendix 1 table 1-a1: list of samples used in the study. filelist convert a pdf file! page 1 page 2 page 3 page 4 a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe original research vol. 50 no. 1 pp. 145 153 springbok behaviour as affected by environmental conditions in the kalahari abstract springbok behavioural ecology in the kalahari was examined with the use of public questionnaires and field forms. springbok favoured grass and forbs overall more than shrubs and trees, but diet selection was influenced by time of day and season. feeding was the most common activity and the frequency of occurrence varied during the day and between seasons. weather and microhabitat conditions were found to have a significant effect on the feeding behaviour. springbok fed in direct sunlight in the mornings and moved into the shade during the afternoon. more time was spent feeding in the shade during the warmer months than during the colder months, especially under northerly to northeasterly wind directions. natural licks were commonly utilised. herd sizes were found to increase during the cold-dry season and decrease during the hot-wet season. springbok and blue wildebeest appeared to avoid competition by niche separation. the study showed that springbok behaviour was significantly affected by environmental conditions. these results imply that changes in climatic conditions, such as those predicted by climate change, or changes in vegetation structure due to degradation, can negatively affect springbok behaviour. keywords: antidorcas marsupialis, climate, feeding ecology, questionnaire, seasonality hein stapelberg margaretha w. van rooyen department of plant science university of pretoria south africa jacobus du p. bothma centre for wildlife management university of pretoria south africa michael j. van der linde hendrik t. groeneveld department of statistics university of pretoria south africa correspondence to: margaretha w. van rooyen e-mail: gretel.vanrooyen@up.ac.za postal address: department of plant science, university of pretoria, pretoria 0002, south africa 145 the kalahari plays host to a range of herbivores of which springbok antidorcas marsupialis (zimmermann, 1780) are characteristic. it has even been suggested that the kalahari springbok be declared a distinct subspecies (bigalke 1970, peters & brink 1992). springbok morphology and anatomy have been studied in detail (skinner & louw 1996), but there are few indepth ecological studies of springbok in the kalahari. studies on the morphology and anatomy of the springbok show that they are well suited to arid areas and are able to cope well with changing environmental pressures (louw & seely, 1982; mills & retief, 1984; spinage & matlhare, 1992; skinner & louw, 1996; skinner & moss, 2004). springbok are selective feeders (bothma et al. 2002) and the quality of their diet is a significant driving force behind their behaviour and movements (mills & retief 1984). they have an exceptionally high annual population growth of 35 to 45% (mills & haagner, 1989; conroy 2005). recent declining springbok populations in south africa have been reported from the kalahari, while other areas such as the karoo have shown a significant increase in recent times (skinner & moss 2004). historical records describe numerous massive migrations (eloff 1961, child & le riche 1969) which are well reviewed by skinner (1993). however, the population numbers of springbok have been declining in the kalahari over the past 50 years, and rapidly so in the past 20 years (skinner, pers. comm.; engelbrecht, pers. comm.; skarpe, 2001; skinner & moss, 2004). studies of sex ratios within springbok populations (bednekoff & ritter 1997) and of morphometrics and reproduction (skinner et al. 1996) do not give any clues to the reason for the decline in numbers, and no significant increase in depredation or predator numbers has been noticed. the focus of this paper is on the activities of springbok on the south african side of the kgalagadi transfrontier park, the kalahari gemsbok national park. the hypothesis that was put forward was that springbok behaviour, in particular feeding and food selection, is significantly affected by environmental and seasonal conditions. to test the hypothesis, data were gathered by means of public questionnaires and researcher observations. study area the kalahari consists of the largest continuous stretch of sand in the world, located more or less diagonally across southern to central africa (van rooyen 2001). the southwestern kalahari is considered a relatively unspoilt, ecologically fragile, arid savanna (van rooyen & van rooyen 1998). the largest conservation unit within the kalahari ecosystem, the kgalagadi transfrontier park, is shared across the borders of south africa and botswana (van rooyen 2001). the present study was carried out on the south african side of the park, called the kalahari gemsbok national park (between 24° 15’ s and 26° 30’ s, and 20° 00’ e and 20° 45’ e), an area that covers some 9 600 km² (jackson 1995). 2008 original research stapelberg, van rooyen, bothma, van der linde & groenewald koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 145 153 33 the southwestern kalahari consists mainly of red sand dunes that are underlain by calcrete. the two main dry riverbeds of the auob and the nossob rivers (jackson 1995; van rooyen & van rooyen, 1998), cut through the dunes and converge just north of the twee rivieren rest camp. among the dunes are some dispersed pans, increasing in abundance from around the centre of the park northwards and eastwards into botswana. the kalahari receives the majority of its rain during summer and has a highly erratic (skarpe, 1986; tyson, 1986) annual rainfall of around 180 mm in the southwest to 400 mm in the northeast. during the study period, the mean rainfall amounted to 213.6 mm over 19 months across all weather stations within the kalahari gemsbok national park (data supplied by the kgalagadi transfrontier park 2004). three ecological seasons occur, namely the hot-wet season (january to april), cold-dry season (may to august) and hot-dry season (september to december) (leistner 1967, mills & retief 1984, van rooyen & van rooyen 1998). the temperature range is immense, with recorded winter lows of around -10°c and summer highs of around 45°c (van rooyen et al. 1990). materials and methods properly designed questionnaires can be used as an effective and affordable tool for gathering large amounts of data within a relatively short period (hague 1993). however, the use of questionnaires is not without flaws and difficulties and those discovered during the course of this study are analysed in stapelberg (2007). constructing meaningful and interpretable questions is a complex process and requires careful word selection to ensure that the questions are asked in such a way that they are fully understood by the respondent. whenever a public questionnaire is used to collect data in conjunction with a researcher, public respondents’ answers should be checked for differences against the researcher’s answers and ground-truthed, where possible. only once these differences are described and understood can the data be used (stapelberg 2007). properly applied statistics are invaluable for filtering such data. a public questionnaire and field data sheet were compiled of observable variables relating to springbok behaviour (appendix a). the questions consisted of factual information (such as date and time), closed format questions (where choices were given) as well as open-ended questions about animal activity. the questionnaire was distributed to staff and visitors of the kalahari gemsbok national park over a period of 13 months. during this time, informative talks were also held at the main rest camp, twee rivieren, explaining the project and the questionnaire. one questionnaire per herd (solitary springbok were also considered as a herd) had to be completed and collection boxes were positioned at the reception office of every camp and emptied periodically. during the same period, the researcher collected data by using a field data sheet that was based on the public questionnaire. the researcher also recorded some additional variables (appendix a). observations were mostly conducted at random within strips (bothma 2002), and were done from a vehicle en route between destinations. most observations were made within the dry riverbeds of the nossob and auob rivers, since this is where the springbok tend to congregate and most of the tourist routes in the park are. visibility varied, but was generally up to around 150 m on either side of the vehicle. binoculars and/or a spotting-scope were used when necessary. data were transformed into numerical codes and these numerical codes were captured into a matrix using microsoft excel. data were classified into ranges or categories wherever it best suited the analytical model or question at hand. all analyses were done using sas software (sas® software version 8.2, sas institute inc., sas campus drive, cary, nc 27512, usa). chisquare tests for independence were conducted to examine the independence of variables. the hypotheses of dependence were expressed by p-values, where p ≤ 0.05 indicated that the variables being tested were independent. generalised linear models were also used to examine the data, incorporating either classed or continuous independent variables. in most cases the independent variable used in the generalised linear models was time. repeated measures analyses of variance were also used in generalised linear models to consider differences between dependent variables at independent variable intervals. logistic procedures were used to calculate odds ratios for binary variables. where necessary, regression analyses were used to clarify trends. where the means for collective data were used, these were calculated as least square means so as to equalise the data for a more accurate mean representation. analyses of variance (anova) were used to distinguish the various attributes of the variables. this paper explores daily and seasonal trends in a selection of springbok activities collected by means of the questionnaire. the analysis of activity data generally involved two steps. first, the activity data were compared with and without the influence of a specified variable to investigate the effect of said variable. thereafter, the means of the activity data were determined according to each class of the specified variable and compared with each other. to investigate the effect of rainfall on herd size, rain was accepted as being present only when a minimum of 10 mm had fallen in a particular month. this was to ensure that a sufficient amount of water had been received to induce vegetation sprouting. the generalised linear model incorporated a lag effect in order to accommodate a response time for vegetation after the occurrence of rain. a total of 17 rainfall stations are located in the kalahari gemsbok national park. a georeferenced grid was created to match specific rainfall stations with herd locations. results and discussion two thousand questionnaires were distributed and 955 (47.7%) were returned. the researcher completed 31.7% and public respondents the balance (68.3%). observations lasted from one to more than ten minutes, most observations not continuing more than three minutes. food item selection food item selection was considered at growth form level. observations were made of springbok feeding either on grass/ forbs, shrubs or trees at different times of the day to determine whether there was a change in food item selection during the course of the day. feeding on grass/forbs, shrubs and trees could not be explained effectively by using time as a continuous independent variable. even when time was reclassified into hourly categories a linear model could not be fitted appropriately for feeding on grass/ forbs (r2 = 0.03; df = 12), shrubs (r2 = 0.02; df = 12) or trees (r2 = 0.01; df = 12). springbok favoured grass/forbs overall more than shrubs and trees (figure 1). the percentage of the herd feeding on grass/forbs reached a maximum in the early morning from > 07:00–08:00 to > 09:00–10:00. this trend is supported by other research (nagy & knight 1994) and is likely related to the increased moisture content of grass during early morning (taylor 1968). tree leaf consumption remained low throughout the day. shrubs were consumed at a lower rate the later in the day it became, possibly due to decreasing moisture content. compared to grass/forb consumption, the relative consumption of shrubs reached a peak at > 11:00–12:00 and again at > 15:00–16:00. diet selection of springbok is affected by season. an investigation 146 springbok behaviour in the kalahari original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 145 15334 into the seasonal changes in nutrient content of springbok food plant species in the kalahari showed that the crude protein and phosphorus levels were especially low during the cold-dry season (stapelberg et al. in press). browse species had a higher crude protein and phosphorus content than grass species. consequently, springbok could consume a higher quality diet if they changed their feeding strategy to include more browse than grasses in their diet during the winter months. springbok were observed to eat rhigozum trichotomum leaves throughout the year, but especially so when the plants were freshly sprouting. this supports observations by bothma (1971), who described a rapid response by springbok to newly emerging green sprouts of r. trichotomum. this species was not specifically selected as food during times when a diversity of fresh plant growth was present, except when in flower. when flowering, springbok were observed to move through r. trichotomum thickets, only feeding on the flowers. it has also been reported that springbok consume plant underground storage organs as a source of moisture (williamson 1987), but that could not be confirmed during the present study. nocturnal feeding no data were collected on nocturnal feeding by public respondents because of tourist travel restrictions, and the data collected by the researcher were mostly qualitative. springbok were observed to feed at night during all three seasons of the year. two peak feeding times were reported during the hot-dry season as opposed to one in the cold-dry and hot-wet seasons. feeding periods were also longer during the two dry seasons than in the wet season. these observations could indirectly be seen as supporting the hypothesis that animals feed on plant material at night because of elevated moisture levels. nocturnal feeding behaviour could be a form of depredation prevention by the herd through having vigilant individuals awake while some continue resting (siegfried, 1980; treves, 2000). licks ‘natural licks’, i.e. areas where animals lick hardened, clayish soil, occur in several places in the kgalagadi transfrontier park. these licks are mostly associated with pans (van rooyen et al. 1994). kalahari springbok make regular use of natural licks, and clear evidence of the presence of clay has been found in the guts of springbok that were shot close to pans in the kalahari in botswana (bergström & skarpe 1999). in the present study, utilisation of such licks was especially prominent around the kannaguass waterhole in the nossob riverbed. springbok utilise licks presumably to supplement their dietary requirements and are reportedly sometimes attracted to artificial waterholes more figure 1 mean percentage of springbok herd feeding on grass/forbs, shrubs and trees at hourly intervals from november 2002 to may 2004 in the kalahari gemsbok national park 0 5 1 0 1 5 2 0 2 5 3 0 3 5 4 0 4 5 5 0 06 :0 0 0 7: 00 >0 7: 00 08 :0 0 >0 8: 00 09 :0 0 >0 9: 00 10 :0 0 >1 0: 00 11 :0 0 >1 1: 00 12 :0 0 >1 2: 00 13 :0 0 >1 3: 00 14 :0 0 >1 4: 00 15 :0 0 >1 5: 00 16 :0 0 >1 6: 00 17 :0 0 >1 7: 00 18 :0 0 >1 8: 00 19 :0 0 time of day p er ce nt ag e of h er d g r a s s & f o r b s s h r u b s tr e e s figure 2 mean percentage of springbok herd involved in different activities, over all time periods, during the study period from november 2002 to may 2004 in the kalahari gemsbok national park. the “other activity” category contains a collection of activities, all of which were recorded infrequently. the value on each bar in the graph indicates the sample size being used to calculate the percentage for that particular activity 9 2 1 9 3 79 3 39 3 6 9 3 1 9 3 89 3 7 9 3 4 9 3 1 9 3 2 9 2 8 9 3 5 9 2 6 0 5 1 0 1 5 2 0 2 5 3 0 3 5 4 0 4 5 fe ed in g in th e su nl ig ht fe ed in g in th e sh ad e st an di ng in th e su nl ig ht st an di ng in th e sh ad e ly in g in th e su nl ig ht ly in g in th e sh ad e sl ee pi ng in th e su nl ig ht sl ee pi ng in th e sh ad e w al kin g in th e su nl ig ht w al kin g in th e sh ad e pl ay in g in th e su nl ig ht pl ay in g in th e sh ad e o th er a ct ivi ty type of activity p er ce nt ag e of h er d in vo lv ed 147 original research stapelberg, van rooyen, bothma, van der linde & groenewald koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 145 153 35 for the licks than the water itself (knight & knight-eloff, 1988; van rooyen et al., 1994). it is unclear whether this was the case in the present study, as springbok were observed to drink water intermittently while also utilising licks. the licks that were observed all occurred within the piosphere zone (jeltsch et al. 1997) around waterholes, and their frequent use could therefore have a degrading effect on the vegetation around waterholes. springbok feeding times although only 21 activities were listed in the questionnaire, up to 31 different activities were specified by respondents. the mean percentage of springbok involved in these activities over all time periods during the whole study period showed that feeding in sunlight occurred most often (38.6%, figure 2). six of all the recorded activities related directly to feeding. neither feeding in the sunlight (p = 0.220; χ² = 0.003; df = 2) nor feeding in the shade (p = 0.150; χ² = 0.004; df = 2) showed any significant changes in frequency of occurrence across different seasons. when these two activities were compared with the rest of the activities by analysis of variance of contrast variables, it was found that feeding in the sunlight occurred significantly more frequently than all other activities (table 1). frequency of feeding in the shade (2.3%) was also significantly different from most other activities, but not from standing in the shade (5.1%; p = 0.265; df = 1), lying down in the shade (3.1%; p = 0.712; df = 1), sleeping in the sunlight (0.3%; p = 0.097; df = 1), walking in the shade (0.3%; p = 0.084; df = 1) and playing in the sunlight (0.7%; p = 0.2745; df = 1) (figure 2; table 1). the lack of a significant difference in the frequency of shaderelated activities (feeding, standing, lying down and walking) indicated that the time spent in the shade was divided equally between these activities. springbok herds were often observed to rest in the shade during the heat of the day. during such resting periods, different animals in the herd would be involved in different activities, such as standing, walking and feeding. the main objective of spending time in the shade other than feeding is assumed to be to rest. daily trends of all activities are presented in figure 3. fewer springbok were observed to be feeding in the sunlight the later in the day it became up to approximately 16:00 (figure 3a). significantly more springbok per herd were feeding in the sunlight at early morning (> 07:00–08:00) than at mid-morning (> 10:00–11:00) (p = 0.0002; df = 1) or mid-afternoon (> 15:00– 16:00) (p < 0.0001; df = 1). in contrast, a significantly larger percentage of the herd was feeding in the shade at midday > 12:00–13:00 than at > 09:00–10:00 (p = 0.032; df = 1). activities that reached their peak frequency in the hour during sunrise included sleeping in direct sunlight (figure 3d) and playing in the shade (figure 3f). feeding and playing in direct sunlight (figures 3a and 3f) reached their peak frequency during early morning, while lying down and playing in the shade (figures 3c and 3f) occurred at their lowest frequency then. lying down in direct sunlight (figure 3c) reached a peak during the late morning, while feeding in the shade (figure 3a) and standing in direct sunlight (figure 3b) had a low frequency of occurrence then. activities that peaked at midday included feeding in the shade (figure 3a) and standing in direct sunlight (figure 3b), while feeding in direct sunlight (figure 3a) occurred at its lowest frequency. during early afternoon standing and lying down in the shade (figures 3b and 3c) and walking in direct sunlight (figure 3e) had a high frequency of occurrence, whereas lying down in direct sunlight (figure 3c) did not occur often then. feeding by season a generalised linear model was used to test the null hypothesis that the frequency of an activity in springbok herds did not differ significantly between the three different seasons. the results showed that none of the feeding-related activities produced either a good linear model fit or a significant dependence when analysed by season (p > 0.130 and r² < 0.005 in all cases). although fewer animals were found to feed in the direct sunlight in the hot-dry season than in the hot-wet season, this difference was not significant (p = 0.087; r² = 0.003). it appeared that more animals fed in the shade during the hotdry season compared with the cold-dry season, but again this difference was not significant (p = 0.075; r² = 0.004) (figure 4). the effect of wind on feeding wind strength analyses relating to wind strength were based on relatively few observations (n = 97). nonetheless, they showed a few notable results. none of the feeding activities seemed to be influenced by wind strength (p > 0.160 in every case), nor did the feeding activities differ significantly from each other with the overall effect of wind strength taken into account (p = 0.352; df = 3). however, standing in the shade was positively linearly related to wind strength (r² = 0.225; p < 0.0001), although the fit was poor. wind direction a larger percentage of the herd fed in the sunlight when there was no wind, a southeasterly, or a random wind than when there was a northerly wind (p = 0.022, p = 0.043 and p = 0.016 table 1 p-values of comparative tests between sets of two variables (activities) to examine independence of the frequency of different activities in springbok herds from november 2002 to may 2004 in the kalahari gemsbok national park. statistical independence is where p ≤ 0.05 fe e d in g in s u n fe e d in g in s h a d e s ta n d in g in s u n s ta n d in g in s h a d e ly in g d o w n in s u n ly in g d o w n in s h a d e s l e e p in g in s u n s l e e p in g in s h a d e w a l k in g in s u n w a l k in g in s h a d e p l a y in g in s u n p l a y in g in s h a d e o t h e r a c t iv it y feeding in sun <0.0001 <0.0001 <0.0001 <0.0001 <0.0001 <0.0001 <0.0001 <0.0001 <0.0001 <0.0001 <0.0001 <0.0001 feeding in shade 0.0051 0.2654 0.0002 0.7123 0.0969 0.0462 0.0280 0.0842 0.2745 0.0493 <0.0001 standing in sun 0.1035 0.2074 0.0173 <0.0001 <0.0001 0.6365 <0.0001 0.0002 <0.0001 0.2406 standing in shade 0.0074 0.4442 0.0150 0.0074 0.2796 0.0141 0.0434 0.0089 0.0005 lying down in sun 0.0005 <0.0001 <0.0001 0.1075 <0.0001 <0.0001 <0.0001 0.6604 lying down in shade 0.0564 0.0275 0.0690 0.0535 0.1618 0.0333 <0.0001 sleeping in sun 0.2869 0.0006 0.9124 0.3252 0.5393 <0.0001 sleeping in shade 0.0003 0.5665 0.0968 0.6795 <0.0001 walking in sun 0.0005 0.0016 0.0003 0.0712 walking in shade 0.3112 0.6864 <0.0001 playing in sun 0.1353 <0.0001 playing in shade <0.0001 other activity 148 springbok behaviour in the kalahari original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 145 15336 figure 3 mean percentage of springbok per herd involved in different activities at hourly intervals during the study period from november 2002 to may 2004 in the kalahari gemsbok national park. (a) feeding in sunlight and shade, (b) standing in sunlight and shade, (c) lying down in sunlight and shade a b c 149 original research stapelberg, van rooyen, bothma, van der linde & groenewald koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 145 153 figure 3 (cont...) mean percentage of springbok per herd involved in different activities at hourly intervals during the study period from november 2002 to may 2004 in the kalahari gemsbok national park: (d) sleeping in sunlight and shade, (e) walking in sunlight and shade, and (f ) playing in sunlight and shade d e f 150 springbok behaviour in the kalahari original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 145 153 respectively). a larger percentage of the herd was observed feeding in the shade in northeasterly wind conditions than in northwesterly conditions or in no wind (p = 0.009 and p = 0.022 respectively). this may suggest that winds from the north to northeast are hotter winds that add to thermoregulation stress in springbok. springbok may subsequently adapt by changing their feeding behaviour to specifically feed in the shade. herd size and rainfall rainfall is a key environmental factor in the southern kalahari (van rooyen et al. 1990) and consequently, the effect of rainfall on herd size was examined. herd sizes varied between seasons and months. significantly larger mean herd sizes were recorded during months of no rain, but when it had rained in the previous month, compared with no rain for two consecutive months (p = 0.047); rain during both consecutive months (p = 0.014; df = 3); or rain during the surveying month, but not the previous month (p = 0.012) (table 2). the largest mean herd sizes were recorded during june 2003 at 102.1 animals (n = 25) and the smallest during november 2002 at 9.1 individuals (n = 57) (table 3). the largest herd recorded during the study period consisted of 476 springbok found in the far northern part of the nossob riverbed, close to kannaguass waterhole, in april 2003. mean herd sizes by season were largest during the cold-dry season (40 springbok, n = 329), intermediate during the hot-wet season (35 springbok, n = 294) and smallest during the hot-dry season (19 springbok, n = 326). these springbok herd sizes coincide roughly with those reported by other researchers in the same area as having a peak from february to may and a trough from october to november (mills & retief 1984, jackson et al. 1993). springbok herd sizes therefore increased during the colddry season and decreased during the hot-dry season. this phenomenon appeared to be linked to specific rainfall patterns rather than to rigid seasonal boundaries. these results also support the results of other studies (bigalke 1972, nagy & knight 1994, bergström & skarpe 1999), and are probably due to the concurrence of lambing with rainfall. bothma and mills (1977), although using total counts instead of herd sizes, reported similar trends. springbok and other animals springbok are known to associate with blue wildebeest connochaetes taurinus and gemsbok oryx gazella (skinner & louw 1996). however, during the present study period, blue wildebeest were either not present or were some distance away from the springbok herds studied in 88.5% of all observations. blue wildebeest were sometimes witnessed to feed in conjunction with springbok, as if moving in a convoy. the distance from the blue wildebeest to the springbok herds was mostly recorded as being at > 50 m away (5.2%), followed by 0–20 m away (4.0%) and then by 21–50 m away (2.3%; p = 0.060; χ² = 7.4; df = 3). gemsbok proximities showed similar trends (p = 0.007; χ² = 9.86; df = 2). moreover, gemsbok were close to springbok more often than blue wildebeest. according to knight (1995), the blue wildebeest grazes at a higher level than the springbok and seems to precede springbok as low-level grazer. qualitative observations confirmed this during the study period. blue wildebeest have wide muzzles that are suited for grazing relatively close to the ground. springbok have narrow muzzles suited for selective feeding (skinner & louw 1996). the results of the present study suggest that springbok and blue wildebeest coexist in the kalahari by avoiding competition through niche separation. they also suggest that although these two animal types occur together, springbok and blue wildebeest more often prefer not to feed in close proximity. a similar situation occurs between plains zebra equus quagga, white-bearded wildebeest connochaetes taurinus subsp. mearnsi and thomson’s gazelle gazella thomsoni in the serengeti in tanzania (sinclair & norton-griffiths 1982). the thomson’s gazelle is morphologically similar to the springbok and is also a selective feeder. however, in their study, sinclair and norton-griffiths (1982) concluded that the white-bearded figure 4 mean percentage of springbok herd feeding in sunlight and shade respectively during the hot-wet, cold-dry and hot-dry seasons from november 2002 to may 2004 in the kalahari gemsbok national park 0 5 1 0 1 5 2 0 2 5 3 0 3 5 4 0 4 5 h o t -w e t s e a s o n c o ld -d ry s e a s o n h o t -d ry s e a s o n season m ea n pe rc en ta ge o f h er ds f e e d in g in s u n lig h t f e e d in g in s h a d e table 2 relationship between herd size of springbok and rainfall in the kalahari gemsbok national park during the study period from november 2002 to may 2004. rainfall in the current and previous months were noted as present (1) or absent (0) respectively. values with the same superscript do not differ significantly (p > 0.05) previous month current month mean herd size 0 0 40.915a 0 1 24.189 a 1 0 71.413b 1 1 15.099 a table 3 mean herd sizes of springbok by month for the period from november 2002 to may 2004 in the kalahari gemsbok national park season month and year mean herd size ± standard deviation number of observations hot-dry november 2002 9.1 ± 20.97 57 december 2002 . . hot-wet january 2003 . . february 2003 . . march 2003 43.7 ± 76.67 73 april 2003 31.1 ± 69.36 209 cold-dry may 2003 53.4 ± 83.43 58 june 2003 102.1 ± 85.47 25 july 2003 56.2 ± 75.07 48 august 2003 23.0 ± 36.45 180 hot-dry september 2003 17.2 ± 25.47 141 october 2003 13.5 ± 21.05 51 november 2003 34.6 ± 59.88 73 december 2003 29.7 ± 13.25 4 hot-wet january 2004 . . february 2004 49.2 ± 76.48 10 march 2004 74.0 ± 36.77 2 april 2004 . . cold-dry may 2004 31.9 ± 55.63 18 total . 40.6 ± 25.26 949 151 original research stapelberg, van rooyen, bothma, van der linde & groenewald koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 145 153 wildebeest were regulating the thomson’s gazelle in the serengeti through competition. competition can therefore not be excluded as a factor regulating the springbok population in the kalahari and more research on this aspect is required. gemsbok are considered roughage feeders and the majority of their diet consists of grass. it is not a strongly territorial animal and is believed to be fairly water-independent (bothma 2002). few similarities between the preferred plant species selected by springbok and gemsbok could be found in the literature (leistner 1959). conclusions the diet of kalahari springbok is highly variable and diet selection is influenced by season and time of day. kalahari springbok make regular use of natural licks, presumably to supplement their dietary requirements. this may have an environmentally degrading effect, especially due to lick locations, but may not be so serious as to require conservation efforts. compared to other activities, feeding was the most important activity of springbok and had the highest frequency of occurrence during all seasons and at all times of the day in the present study. climate and weather conditions were found to have a significant effect on the feeding behaviour, as did shade and direct sunshine. in general, the springbok fed in direct sunlight in the mornings and the frequency of feeding in the shade increased towards midday. they also spent more time feeding in the shade during the hot-dry season than during the cold-dry and hot-wet ones. wind strength did not affect feeding behaviour, although wind direction did. springbok fed in the shade more often under northerly and northeasterly wind directions. herd sizes were significantly related to specific rainfall patterns rather than to rigid seasonal boundaries. the ecological relationship between springbok and blue wildebeest was investigated and evidence was found of competition avoidance through niche separation. the study showed that springbok behaviour was significantly influenced by time of day, season, wind and rainfall and the hypothesis that environmental conditions affect springbok behaviour significantly can confidently be accepted. the presence of large trees that provide shade was important as these seemed to afford some protection against the extreme heat in the middle of the day or against hot winds from the north to northeast. these results imply that changes in climatic conditions, such as those predicted by climate change, or changes in vegetation structure due to degradation, will affect springbok behaviour. the information that was gained from this research project gives an overview of the current springbok ecology in the kalahari; it serves as a valuable benchmark for future studies on springbok behaviour; and provides background knowledge to generate hypotheses on possible reasons for the decline in springbok numbers that can be investigated in future studies. acknowledgements we thank camelbak international for their support and express our gratitude to the kalahari gemsbok national park for approving this project and the university of pretoria and the national research foundation (gun 2053522) for their financial support. references bednekoff, p.a. & ritter, r.c. 1997. adult sex ratio of a wild population of springbok (antidorcas marsupialis) at nxai pan, botswana. african journal of wildlife research, 27: 22–26. bergström, r. & skarpe, c. 1999. the abundance of large wild herbivores in a semi-arid savanna in relation to seasons, pans and livestock. african journal of ecology, 37(1): 12–26. bigalke, r.c. 1970. observations on springbok populations. zoologica africana, 5: 59–70. bigalke, r.c. 1972. observations on the behaviour and feeding habits of springbok, antidorcas marsupialis. zoologica africana, 7: 333–359. bothma, j. du p. 1971. notes on river habitat use by larger ungulates in the kalahari gemsbok national park. koedoe, 14: 33–48. bothma, j. du p. (ed.) 2002. game ranch management, 4th ed. pretoria: van schaik. bothma, j. du p. & mills, m.g.l. 1977. ungulate abundance in the nossob river valley, kalahari desert. proceedings of the xiiith international congress of game biologists, atlanta, georgia, pp. 90–102. bothma, j. du p., van rooyen, n. & du toit, j.g. 2002. antelope and other smaller herbivores. in: bothma, j. du p. (ed.) 2002. game ranch management, 4th ed. pretoria: van schaik, pp. 149–175. child, d. & le riche, j.d. 1969. recent springbok treks (mass movements) in south-western botswana. mammalia, 33: 499–504. conroy, a.m. 2005. the springbok. in: bothma, j. du p. & van rooyen, n. (eds). intensive wildlife production in southern africa. pretoria: van schaik, pp. 214–226. eloff, f.c. 1961. observations on the migrations and habits of the antelope of the kalahari gemsbok park, part 3. koedoe, 4: 18–30. engelbrecht, d. dries@sanparks.org, 2002. stapelberg, h, hein. staplberg@gmail.com, november 2002. hague, p. 1993. questionnaire design. london: kogan page. jackson, t.p. 1995. the role of territoriality in the mating system of the springbok antidorcas marsupialis (zimmermann 1780). phd dissertation, pretoria: university of pretoria. jackson, t.p., skinner, j.d. & richardson, p.r.k. 1993. some costs of maintaining a perennial territory in the springbok, antidorcas marsupialis. african journal of ecology, 31(3): 242– 254. jeltsch, f., milton, s.j., dean, w.r.j. & van rooyen, n. 1997. simulated pattern formation around artificial waterholes in the semi-arid kalahari. journal of vegetation science, 8(2): 177–188. johnson, d.h. 1980. the comparison of usage and availability measurements for evaluating resource preference. ecology, 61(1): 65–71. knight, m.h. 1995. tsamma melons, citrullus lanatus, a supplementary water supply for wildlife in the southern kalahari. african journal of ecology, 33: 71–80. knight, m.h. & knight-eloff, a.k. 1988. the importance of borehole water and lick sites for kalahari south africa ungulates. journal of arid environments, 15: 269–282. leistner, o.a. 1959. notes on the vegetation of the kalahari gemsbok national park with special reference to its influence on the distribution of antelopes. koedoe, 2: 128– 151. leistner, o.a. 1967. the plant ecology of the southern kalahari. memoirs of the botanical survey of south africa, 38: 1–172. louw, g.n. & seely, m.k. 1982. ecology of desert organisms. london: longman. mills, m.g.l. & haagner, c. 1989. guide to the kalahari gemsbok national park. johannesburg: southern book publishers. mills, m.g.l. & retief, p.f. 1984. the response of ungulates to rainfall along the riverbeds of the southern kalahari. koedoe (suppl.), 27: 129–141. nagy, k.a. & knight, m.h. 1994. energy, water, and food use by springbok antelope (antidorcas marsupialis) in the kalahari desert. journal of mammalogy, 75: 860–872. peters, j. & brink, j.s. 1992. comparative postcranial osteomorphology and osteometry of springbok antidorcas marsupialis (zimmermann, 1780) and grey rhebok, pelea capreolus (forster, 1790) (mammalia: bovidae). memoirs of the national museum bloemfontein, 8: 161–207. siegfried, w.r. 1980. vigilance and group size in springbok. madoqua, 12: 151–154. 152 springbok behaviour in the kalahari original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 145 153 sinclair, a.r.e. & norton-griffiths, m. 1982. does competition or facilitation regulate migrant ungulate populations in the serengeti? a test of hypotheses. oceologia, 53(3): 364–369. skarpe, c. 1986. plant community structure in relation to grazing and environmental changes along a north-south transect in the western kalahari. vegetatio, 68(1): 3–18. skarpe, c. 2001. resource use and rangeland products. available at: http://www.maposda.net/global/reports/lesotho/ session2b_botswana_2.pdf [accessed 2006]. skinner, j.d. 1993. springbok (antidorcas marsupialis) treks. transactions of the royal society of south africa, 48: 291–305. skinner, j.d., john.skinner@up.ac.za, 2003. stapelberg, h, hein. stapelberg@gmail.com, february 2003. skinner, j.d. & louw, g.n. 1996. the springbok antidorcas marsupialis (zimmermann, 1780). pretoria: transvaal museum. skinner, j.d. & moss, d.g. 2004. kgalagadi springbok (antidorcas marsupialis): bucking the trend. transactions of the royal society of south africa, 59(2): 119–121. skinner, j.d., van aarde, r.j., knight, m.h. & dott, h.m. 1996. morphometrics and reproduction in a population of springbok antidorcas marsupialis in the semi-arid southern kalahari. african journal of ecology, 34(3): 312–330. spinage, c.a. & matlhare, j.m. 1992. is the kalahari cornucopia fact or fiction? a predictive model. journal of applied ecology, 29(3): 605–610. stapelberg, f.h. 2007. feeding ecology of the kalahari springbok antidorcas marsupialis in the kgalagadi transfrontier park, south africa. msc. dissertation. pretoria: university of pretoria. stapelberg, f.h., van rooyen, m.w. & bothma, j. du p. in press. seasonal nutrient fluctuation of selected plant species in the kalahari. african journal of range and forage science. taylor, c.r. 1968. hygroscopic food: a source of water for desert antelope? nature, 219(4): 181–182. treves, a. 2000. theory and methods in study of vigilance and aggregation. animal behaviour, 60(6): 711–722. tyson, p.d. 1986. climatic change and variability in southern africa. cape town: oxford. van rooyen, n. 2001. flowering plants of the kalahari dunes. pretoria: ekotrust. van rooyen, n., bezuidenhout, d., theron, g.k. & bothma, j. du p. 1990. monitoring of the vegetation around artificial watering points (windmills) in the kalahari gemsbok national park. koedoe, 33(1): 63–88. van rooyen, n., bredenkamp, g.j., theron, g.k., bothma, j. du p. & le riche, e.a.n. 1994. vegetational gradients around artificial watering points in the kalahari gemsbok national park. journal of arid environments, 26(4): 349–361. van rooyen, n. & van rooyen, m.w. 1998. vegetation of the south-western arid kalahari: an overview. transactions of the royal society of south africa, 53(2): 113–140. williamson, d.t. 1987. plant underground storage organs as a source of moisture for kalahari wildlife. african journal of ecology, 25(1): 63–64. appendix a variables, listed in the questionnaire, that were recorded by staff, the researcher and visitors to the kalahari gemsbok national park from november 2002 to may 2004 to study springbok behaviour in the park. note that not all variables are reported on in this publication. date location time when observation starts total number of springbok riverbed habitat (yes or no) plains habitat (yes or no) duneveld (yes or no) 1. 2. 3. 4. 5. 6. 7. pan-type habitat (yes or no) tree-dominated habitat (yes or no) grass-dominated habitat (yes or no) shrub-dominated habitat (yes or no) other (three alternative dominant habitat features recorded) near a camp (yes or no) near a waterhole (yes or no) near a picnic spot (yes or no) other (none specified) total number of blue wildebeest total number of red hartebeest total number of gemsbok total number of other species (13 other animal species recorded) distance between springbok and blue wildebeest (0–20 m, 21–50 m and > 50 m specified) distance between springbok and red hartebeest (0–20 m, 21–50 m and > 50 m specified) distance between springbok and gemsbok (0–20 m, 21–50 m and > 50 m specified) distance between springbok and other animal species (0–20 m, 21–50 m and > 50 m specified) percentage of the springbok herd feeding in sunlight* percentage of the springbok herd feeding in the shade* percentage of the springbok herd standing in sunlight* percentage of the springbok herd standing in shade* percentage of the springbok herd lying down in sunlight* percentage of the springbok herd lying down in shade* percentage of the springbok herd sleeping in sunlight* percentage of the springbok herd sleeping in shade* percentage of the springbok herd walking in sunlight* percentage of the springbok herd walking in shade* percentage of the springbok herd playing in sunlight* percentage of the springbok herd playing in shade* percentage of the springbok herd doing something else* (19 alternative actions recorded) percentage of the springbok herd feeding on grass* percentage of the springbok herd feeding on shrubs* percentage of the springbok herd feeding on trees* percentage of the springbok herd feeding on something else* (22 alternative options recorded) direction in which animals are moving (north, northeast, east, southeast, south, southwest, west and northwest specified) time when observation ends sheet number total number of male springbok in the herdφ total number of female springbok in the herdφ total number of immature springbok in the herdφ springbok are feeding on flowers (yes or no) φ springbok are feeding on fruit (yes or no) φ nearest waterhole (47 waterholes specified) φ habitat type (four options specified) φ additional variables, only recorded by the researcher springbok are feeding on fresh, new plant growth (yes or no) springbok are feeding on old plant growth (yes or no) wind direction (north, northeast, east, southeast, south, southwest, west and northwest specified) wind strength (none, breeze, moderate, strong, extreme) * numbers were converted into percentages of the total herd after collection. φ although variables 45 to 51 did not appear on the questionnaire, they were recorded as additional notes by the researcher, as well as some of the visitors/ staff. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. 51. 1. 2. 3. 4. 153 filelist convert a pdf file! page 1 page 2 http://www.koedoe.co.za open access page 1 of 1 reviewer acknowledgement acknowledgement to reviewers in an effort to facilitate the selection of appropriate peer reviewers for koedoe, we ask that you take a moment to update your electronic portfolio on https://koedoe.co.za for our files, allowing us better access to your areas of interest and expertise, in order to match reviewers with submitted manuscripts. if you would like to become a reviewer, please visit the journal website and register as a reviewer. to access your details on the website, you will need to follow these steps: 1. log into the online journal at https://koedoe. co.za 2. in your ‘user home’ [https://koedoe.co.za/ index.php/koedoe/user] select ‘edit my profile’ under the heading ‘my account’ and insert all relevant details, bio statement and reviewing interest(s). 3. it is good practice as a reviewer to update your personal details regularly to ensure contact with you throughout your professional term as reviewer to koedoe. please do not hesitate to contact us if you require assistance in performing this task. publisher: publishing@aosis.co.za tel: +27 21 975 2602 tel: 086 1000 381 the editorial team of koedoe recognises the value and importance of the peer reviewer in the overall publication process – not only in shaping the individual manuscript, but also in shaping the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely, productive peer review process. we would like to take this opportunity to thank all reviewers who participated in shaping this volume of koedoe. we appreciate the time taken to perform your review(s) successfully. adrian j. armstrong alexander n. songorwa althea l. davies bernard w.t. coetzee bettine jansen van vuuren brian taylor chloe guerbois colin everson david le maitre doug harebottle edmund february elmarie slabbert helen suich hendri coetzee hugo bezuidenhout james pryke jan a. venter jan vlok jesse m. kalwij keith bosak lizette moolman louise swemmer maartin strauss marna herbst paul martin peet van der merwe peter a. novellie priscilla burgoyne robert f. brand roland hunter sam m. ferreira theo h.c. mostert yoshan moodley http://www.koedoe.co.za� https://koedoe.co.za� https://koedoe.co.za� https://koedoe.co.za� https://koedoe.co.za/index.php/koedoe/user https://koedoe.co.za/index.php/koedoe/user mailto:publishing@aosis.co.za filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 abstract introduction methods results and discussion acknowledgements references about the author(s) robin m. petersen scientific services, south african national parks, kruger national park, south africa andries c. hoffman aquatic systems, mpumalanga tourism and parks agency, south africa pieter kotze clean stream biological services, malelane, south africa sean m. marr south african institute for aquatic biodiversity, grahamstown, south africa citation petersen, r.m., hoffman, a.c., kotze, p. & marr, s.m., 2017, ‘first record of the invasive australian redclaw crayfish cherax quadricarinatus (von martens, 1868) in the crocodile river, kruger national park, south africa’, koedoe 59(1), a1435. https://doi.org/10.4102/koedoe.v59i1.1435 short communication first record of the invasive australian redclaw crayfish cherax quadricarinatus (von martens, 1868) in the crocodile river, kruger national park, south africa robin m. petersen, andries c. hoffman, pieter kotze, sean m. marr received: 16 aug. 2016; accepted: 11 nov. 2016; published: 31 mar. 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the redclaw crayfish cherax quadricarinatus (von martens, 1868), a robust freshwater crayfish native to australia and papua new guinea, has now been recorded from the kruger national park (knp). previously absent from the crocodile river, san parks received a report in february 2016 of redclaw crayfish below the van graan dam on the border of the knp. here, we provide evidence of the presence of redclaw crayfish in the crocodile river. a better understanding of the redclaw crayfish distribution, habitat preferences, rate of spread and impacts on the local aquatic ecosystems in the crocodile river is urgently required to develop mitigation strategies that minimise the spread of this invasive crayfish in the knp and the komati catchment. the negative impacts of global crayfish introductions justify efforts to discourage further introductions and prevent their secondary spread. conservation implications: a better understanding of the redclaw crayfish distribution, habitat preferences, rate of spread and impacts on the local aquatic ecosystems in the crocodile river is urgently required to develop mitigation strategies that minimise the spread of this invasive crayfish in the kruger national park and the komati catchment. introduction redclaw crayfish cherax quadricarinatus (von martens, 1868) is a robust freshwater crayfish native to australia and papua new guinea (ruscoe 2002). tolerant of a broad range of environmental conditions (masser & rouse 1997; medley, jones & avault 1994), the species has a simple reproductive cycle and fast growth rates (masser & rouse 1997; ruscoe 2002), making it of specific interest to the commercial aquaculture (masser & rouse 1997; medley et al. 1994; ruscoe 2002; saoud et al. 2013) and ornamental aquarium industries (ahyong & yeo 2007; belle et al. 2011). as a result, redclaw crayfish have been introduced into many regions including latin america, the caribbean islands, china, taiwan, swaziland, zimbabwe, zambia and the united states (harlioğlu & harlioğlu 2006; tyser & douthwaite 2014). concern has been raised that the rapid expansion of redclaw crayfish in the zambezi basin could threaten the okavango delta (nunes et al. 2016). a meta-analysis of global crayfish introductions found an overall negative ecological impact affecting all levels of freshwater food webs including reductions in the abundance and growth of macrophytes, aquatic invertebrates, amphibians and fish (twardochleb, olden & larson 2013). negative impacts on macrophytes, invertebrates and fish were often coupled with positive effects on algal growth. the impact of a crayfish introduction was determined more by extrinsic characteristics of the recipient ecosystem than the species of crayfish introduced (twardochleb et al. 2013). the potential impacts of c. quadricarinatus in south africa include the destruction of aquatic macrophytes, disturbance of gravel beds used by substrate-spawning fish, predation on benthic macro-invertebrate communities and introduction of commensals, for example, parasites (de moor 2002). although the redclaw crayfish has been evaluated as a potential aquaculture species in south africa, the importation of this species has remained prohibited with all permit applications to import the species denied (de moor 2002). however, redclaw crayfish were permitted to be imported into swaziland in the 1990s for an aquaculture venture. the venture failed and a wild population of redclaw crayfish established in the sand river dam in swaziland (de moor 2002), and subsequently in the sand river, a tributary of the komati river, komati catchment by 1999 (de moor 2002). the sand river population have spread into the komati river and its tributaries, including the lomati river (de villiers 2015) but was first recorded in south africa in the komati river in 2002 (de villiers 2015). it was first reported in scientific literature from the outlet of lake nyamiti in ndumo game reserve, usuthu-phongolo/maputo river system in 2012 (du preez & smit 2013) and subsequently from two sites on the komati river in 2015, one just upstream from the confluence with the crocodile river (tavakol et al. in press). the origin of the lake nyamiti population remains unresolved (du preez & smit 2013), as the usuthu-phongolo/maputo river system has no connection to the komati catchment. although previously absent from the crocodile river, san parks received a report in february 2016 of redclaw crayfish below the van graan dam on the southern border of the kruger national park (knp). here, we report a new locality record for redclaw crayfish in the crocodile river. methods intensive sampling was conducted below van graan dam on the crocodile river on the southern boundary of the knp in march 2016 because this was the location where the crayfish had been reported. the sluices of van graan dam were temporarily closed (for roughly 40 minutes) to allow a thorough examination of the site below the dam wall. while the sluices were closed, specimens were collected using an electro-shocking device (samus 4254) and hand netting, and some specimens were caught by hand. the cephalothorax length, weight and sex of crayfish captured were recorded. specimens collected were preserved as vouchers for the south african national parks skukuza biological reference collection. to establish the extent of the invasion, five additional sites, including above the van graan dssam wall, along the crocodile river were sampled over 3 days using five baited traps per site set overnight (figure 1). the traps were constructed from pvc pipes (length = 700 mm, diameter = 60 mm) and chicken mesh wire to create a non-return entrance on each side. commercial dog food pellets were used as bait. the traps were set in deep, slow-flowing habitats typically found above off-take weirs and bridges. intensive sampling by wading at these sites was not possible because of the presence of crocodiles and hippopotamus. figure 1: map showing the location of the sites sampled in this study. results and discussion crayfish (six males and three females) were only caught below van graan dam (s -25.439773°; e 31.635407°), while the sluices were closed, using electro-shocking. no crayfish were caught at the additional sites using the baited traps. the largest of which had 56 mm cephalothorax length and weighed 150 g (figure 2); the smallest measured 39 mm and weighed 52 g (table 1). the crayfish were captured in areas that are normally deep and fast flowing while the sluices are open. this was surprising because crayfish have in past surveys mostly been caught in deep, slow-moving water in the komati river (a. hoffman, pers. obs., n.d.). our sampling in the crocodile river intentionally targeted deep, slow-flowing habitats, but this study indicates that future surveys should target other habitats on the crocodile river. figure 2: dorsal view of a male redclaw crayfish found in the crocodile river, south africa. table 1: record of measurements recorded for redclaw crayfish captured below van graan dam, crocodile river, on the border of the kruger national park. this study confirms the presence of c. quadricarinatus in the crocodile river. it is unclear at present whether redclaw crayfish colonised the site from downstream or were illegally introduced at the site. possession of this species remains illegal in south africa (de moor 2002; republic of south africa 2014) but articles in farming magazines have extolled the economic potential of the species in aquaculture (copeland 1999; van rooyen 2013) without demonstrating a clear understanding of the threat this species poses to aquatic biodiversity (de moor 2002). from the current surveys, redclaw crayfish in the crocodile river appears to be isolated to just below the van graan dam, which may provide credence to suggestions that they have been illegally stocked at this locality. however, we cannot exclude colonisation from the komati river and upstream via the crocodile river because they were recently recorded from the komati river just upstream of its confluence with the crocodile river. the densities are currently low and no evidence of recruitment has been recorded to date, raising questions whether the population is self-sustaining at present. both sexes and a range of size classes were, however, recorded at the site. redclaw crayfish may have naturally colonised the site from the population in the komati river, but in this scenario, one would expect to have evidence of the species at suitable sites between van graan dam and the confluence between the crocodile and komati rivers. further sampling is required to determine the extent of invasion of redclaw crayfish in the crocodile river. freshwater biotic invasions are generally irreversible once the species have become established (cucherousset & olden 2011). a better understanding of the redclaw crayfish distribution, habitat preferences, rate of spread and impacts on the local aquatic ecosystems in the crocodile river is urgently required to develop mitigation strategies that minimise the spread of this invasive crayfish in the knp and the komati catchment. the negative impacts of crayfish introductions justify efforts to discourage further introductions and prevent their secondary spread (de moor 2002; twardochleb et al. 2013). acknowledgements the authors thank employees from sanparks, mr isaac sedibe, our game guard and ms chenay simms for producing the map. the authors also thank mr willie du toit from the crocodile irrigation board, dr t.a. zengeya from south african national biodiversity institute (sanbi) and dr f. roux from mpumalanga tourism and parks agency (mtpa). the comments from the reviewers are also acknowledged. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions r.m.p. was the project leader; a.c.h. and p.k. were responsible for experimental and project design and implementation. s.m.m. co-wrote the manuscript. references ahyong, s.t. & yeo, d.c., 2007, ‘feral populations of the australian red-claw crayfish (cherax quadricarinatus von martens) in water supply catchments of singapore’, biological invasions 9, 943–946. https://doi.org/10.1007/s10530-007-9094-0 belle, c.c., wong, j.q., yeo, d.c., tan, s., tan, h.h., clews, e. et al., 2011, ‘ornamental trade as a pathway for australian redclaw crayfish introduction and establishment’, aquatic biology 12, 69–79. https://doi.org/10.3354/ab00324 copeland, j., 1999, ‘rich pickings from crayfish’, farmer’s weekly, 8 october, 1999, p. 89–91. cucherousset, j. & olden, j.d., 2011, ‘ecological impacts of non-native freshwater fishes’, fisheries 36, 215–230. https://doi.org/10.1080/03632415.2011.574578 de moor, i., 2002, ‘potential impacts of alien freshwater crayfish in south africa’, african journal of aquatic science 27, 125–139. http://dx.doi.org/10.2989/16085914.2002.9626584 de villiers, m., 2015, ‘freshwater crayfish found in the komati river’, lowvelder, 08 july, 2015, viewed n.d., from http://lowvelder.co.za/279853/crayfish-web/ du preez, l. & smit, n., 2013, ‘double blow: alien crayfish infected with invasive temnocephalan in south african waters’, south african journal of science 109, 1–4. https://doi.org/10.1590/sajs.2013/20130109 harlioğlu, m.m. & harlioğlu, a.g., 2006, ‘threat of non-native crayfish introductions into turkey: global lessons’, reviews in fish biology and fisheries 16, 171–181. https://doi.org/10.1007/s11160-006-9010-1 masser, m.p. & rouse, d., 1997, australian red claw crayfish, southern regional aquaculture center, stoneville, srac publication no. 244. medley, p.b., jones, c.m. & avault, j.w., jr., 1994, ‘a global perspective of the culture of australian redclaw crayfish, cherax quadricarinatus: production, economics and marketing’, world aquaculture 25, 4. nunes, a.l., douthwaite, r.j., tyser, b., measey, g.j. & weyl, o.l.f., 2016, ‘invasive crayfish threaten okavango delta’, frontiers in ecology and the environment 14, 237–238. https://doi.org/10.1002/fee.1287 republic of south africa, 2014, government notice no. 37885. vol. 590, regulation gazette no. 10244, government publishing works. ruscoe, i., 2002, fishnote 32: redclaw crayfish aquaculture, darwin fisheries, northern territory government, australia. saoud, i.p., ghanawi, j., thompson, k.r. & webster, c.d., 2013, ‘a review of the culture and diseases of redclaw crayfish cherax quadricarinatus (von martens, 1868)’, journal of the world aquaculture society 44, 1–29. https://doi.org/10.1111/jwas.12011 tavakol, s., luus-powell, w.j., smit, w.j., baker, c., hoffman, a. & halajian, a., in press, ‘first introduction of two species of australian temnocephalans into africa with an alien host: double trouble’, journal of parasitology. https://doi.org/10.1645/15-936 twardochleb, l.a., olden, j.d. & larson, e.r., 2013, ‘a global meta-analysis of the ecological impacts of nonnative crayfish’, freshwater science 32, 1367–1382. https://doi.org/10.1899/12-203.1 tyser, a. & douthwaite, r., 2014, ‘predation on invasive redclaw crayfish cherax quadricarinatus by native fishes in the kafue river, zambia’, african journal of aquatic science 39, 473–477. https://doi.org/10.2989/16085914.2014.980774 van rooyen, l., 2013, ‘feral freshwater crayfish: ideal food’, farmer’s weekly, 16 september, 2013, viewed n.d., from http://www.farmersweekly.co.za/agri-technology/farming-for-tomorrow/feral-freshwater-crayfish-ideal-food/ kyle fish mortality.qxd a selective fish mortality and some notes on rhabdosargus sarba at kosi bay, south africa r. kyle issn 0075-6458 117 koedoe 45/2 (2002) on 6 july 2000, while crossing lake nhlange, the largest of the kosi bay lakes, several dead fish were seen, lying bloated on the surface. on investigation they were found to be large natal stumpnose rhabdosargus sarba. occasionally, during twenty years of residence at the kosi lakes, the author has seen individual natal stumpnoses dead or dying in this lake, particularly in winter, but never so many at one time. on investigation, by questioning other local residents, it was found that the mortality had been taking place for several days and it subsequently extended for at least another 14 days. efforts were made to identify and measure as many dead and dying fish as possible (table 1) but those recovered were a small proportion of the total number of fish which died. the total mortality was probably several thousand fish. the vast majority of dead fish were mature r. sarba (tables 1 &2) and dead fish were not reported in any of the other kosi bay lakes. nearly all the fish were affected by a white fungal growth over much of their bodies, particularly behind the dorsal and anal fins. initially there was deep concern among locally resident anglers that a sickness or poison was killing the fish, as many rely on the fish resource for their livelihood. soon, however, they began collecting dead and dying fish on the lee shore each morning and eating or selling them. no illness or deaths were reported from people eating the fish and so the mortality was later regarded by local residents as fortuitous. results of several days’ collecting and measuring of dead fish show that 85 % of the fish by number, and almost 94 % by mass, were r. sarba (table 1). the next most common species were r. holubi (cape stumpnose), a much smaller member of the genus, then pomadasys commersonni (spotted grunter) followed by a few very small fish and a single acanthopagrus berda (riverbream). many other r. sarba were collected by locals but they reported few other species and natural history note table 1 species composition and total mass estimates of dead fish found in lake nhlange between 6 and 28 july 2000 species number % total mass estimate (kg) % rhabdosargus sarba 215 85 322.5 93.9 rhabdosargus holubi 17 6.7 2.5 0.8 pomadasys commersonni 6 2.4 12 3.5 gilchristella aestuaria 6 2.4 0.1 0.0 caranx spp. 4 1.6 6 1.7 hyporhamphus capensis 3 1.2 0.1 0.0 ambassis spp. 2 0.8 0.1 0.0 acanthopargus berda 1 0.4 0.3 0.1 total 254 100 343.6 100.1 kyle fish mortality.qxd 2005/12/09 10:06 page 117 would probably not have noticed the very small fish. mass fish mortalities have been recorded in several estuaries including kosi bay (blaber & whitfield 1976). although many estuarine fish are resistant to extremes of temperature (kyle 1984) this is often the reason suggested for mortalities. these have been ascribed to particularly high temperatures (kyle 1998) or particularly low temperatures (blaber & whitfield 1976, kyle 1989). they have also been ascribed to chemical pollutants (ramm 1992) as well as a combination of low temperatures and low salinity (blaber 1973a, 1973b) sometimes they appear to have affected most fish present (blaber & whitfield 1976; kyle 1989) while often they only affect certain species or size classes (blaber 1973a, 1973b; kyle 1998). a large amount of information is now available on the fish and the environment of the kosi bay lakes, and it is possible to use this to examine the selectivity of this mortality. the mortality took place in winter and there were no records of, or information on any unusual chemicals in the lake at the time. the catches of the legal gillnet fishery, operating in these margins, should give an koedoe 45/2 (2002) 118 issn 0075-6458 table 2 species composition of gillnet catches in lake nhlange for may and june 2000 and for larger fish found dead during july 2000 species % gillnet % mortality gerres methueni 56.9 0 mugil cephalus 20.1 0 rhabdosargus sarba 9.1 88.9 pomadasys commersonni 6.6 2.4 myxus capensis 2.9 0 acanthopagrus berda 2.3 0.4 oreochromis mossambicus 1.7 0 rhabdosargus holubi 0 6.7 caranx spp. 0 1.6 other species 0.4 0 number of fish examined 145 253 insight into the fish species and size present at the time. although gillnetting is a selective way of catching fish, gillnet catch species composition (table 2) shows that there were large numbers of species other than r. sarba present, which were not recorded in the mortality. the gillnet catch species composition in the two months prior to the mortality (table 2) show that gerres methueni, mugil cephalus, myxus capensis and oreochromis mossambicus, together comprised over 80 % of the catch but were not recorded in the mortality. pomadasys commersonni and acanthopagrus berda appeared to die in numbers lower than expected. only rhabdosargus sarba, r. holubi and caranx species were found in numbers greater than their gillnet catch abundance would have suggested, had all fish been equally affected. the mean total length of gillnetted r. sarba (395 mm) was similar to that of the fish found dead (389 mm). although local residents occasionally report dead and dying fish, when questioned they said that they had never seen so many. temperatures are recorded daily at a jetty in the ne of nhlange lake and these showed levels no lower than usual for this time of year. salinity is also recorded in the lake and over the last few years it had been slowly rising, from about 0.5 parts per thousand to almost three parts per thousand. extremely heavy rainfall in late summer, more than double the average, flushed the system and reduced salinities markedly and rapidly back to about 0.5 parts per thousand in nhlange lake. the salinity in lakes mpugwini and makawulani, which also contained large r. sarba, remained above 3 parts per thousand. data from kosi bay national marine linefish survey voluntary catchcards (pradervand pers. comm.) show an increase in mean r. sarba mass, from 0.7 kg in 1994 to 1.4 kg. in 1999, which was the period of rising salinity. most of the r. sarba found dead were well above the size reported to be found in estuaries by whitfield (1998) who stated that a proportion of the spent individuals return to estuaries but these are confined to fish less than 35 cm (standard kyle fish mortality.qxd 2005/12/09 10:06 page 118 length). whitfield also reported in an earlier publication (blaber & whitfield 1976) that a combination of salinities below 3 parts per thousand and temperatures below 12 ºc were lethal to r. sarba. the lowest daily water temperature recorded in nhlange in july 2000 was 15 ºc. most of whitfield’s work was carried out south of, and thus in cooler areas, than in kosi bay. data reported here suggest that salinities of around 3 parts per thousand and warmer temperatures allowed much larger r. sarba than previously reported to enter or grow up in the kosi bay lakes. when temperatures and salinities fell, the r. sarba probably suffered severe osmoregulatory stress which led to often fatal fungal infections (blaber & whitfield 1976). it appears that, in the preceding years a combination of salinities above 3 parts per thousand and temperatures above 15 ºc allowed r. sarba well above the predicted maximum size, to live in the kosi bay lakes. it seems likely that the larger r. sarba will now disappear from lake nhlange until salinities rise again. other larger marine-origin fish species found in estuaries seem to be much more resistant to falling salinities and temperatures. acknowledgements i thank ezemvelo kwazulu natal wildlife for permission to publish this paper. references blaber, s.j.m. 1973a. temperature and salinity tolerance of juvenile rhabdosargus holubi (steindachner) (teleostei). journal of fish biology. 5 : 593 599. blaber, s.j.m. 1973b.population size and mortality of juveniles of the marine teleost rhabdosargus holubi (pisces : sparidae) in a closed estuary. international journal on life in oceans and coastal waters. 21 (3) : 219 225. blaber, s.j.m. & whitfield, a.k. 1976. large scale mortality of fish at st. lucia. south african journal of science. 72: 218. kyle, r. 1984. tolerance of high temperature by the cichlid oreochromis mossambicus in the kosi system. lammergeyer. 32:50. kyle, r. 1989. a mass mortality of fish at kosi bay. lammergeyer. 40:39. kyle, r. 1998. an interesting, possibly man enhanced, fish mortality at kosi bay. lammergeyer 45:37-38. ramm, a.e. l. 1992. aspects of the biochemistry of sulphur in lake mpugwini, southern africa. estuarine, coastal and shelf science 34: 253 261. r. kyle, p.o. box 43, kwangwanase, 3973, south africa. (ezemvelo kwazulu natal wildlife; rkyle@iafrica.com). issn 0075-6458 119 koedoe 45/2 (2002) kyle fish mortality.qxd 2005/12/09 10:06 page 119 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <feff004200720075006b00200064006900730073006500200069006e006e007300740069006c006c0069006e00670065006e0065002000740069006c002000e50020006f00700070007200650074007400650020005000440046002d0064006f006b0075006d0065006e0074006500720020006d006500640020006800f80079006500720065002000620069006c00640065006f00700070006c00f80073006e0069006e006700200066006f00720020006800f800790020007500740073006b00720069006600740073006b00760061006c00690074006500740020006600f800720020007400720079006b006b002e0020005000440046002d0064006f006b0075006d0065006e0074006500720020006b0061006e002000e50070006e006500730020006d006500640020004100630072006f0062006100740020006f0067002000520065006100640065007200200035002e00300020006f0067002000730065006e006500720065002e00200044006900730073006500200069006e006e007300740069006c006c0069006e00670065006e00650020006b0072006500760065007200200073006b00720069006600740069006e006e00620079006700670069006e0067002e> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice book review eden lost, and regained – book review book title: eden lost, and regained: the story of south africa’s wild life: from decimation to conservation author: a.p. cartwright isbn: 978-0-6398050-1-6. hardcover publisher: footprint press, south africa, 2019, r500.00* *book price at the time of review review title: eden lost, and regained – book review reviewer: peter a. novellie1 affiliation: 1sustainability research unit, nelson mandela university, george, south africa corresponding author: peter novellie, novellie@netactive.co.za how to cite this book review: novellie, p.a., 2020, ‘eden lost, and regained – book review’, koedoe 62(1), a1628. https://doi.org/10.4102/koedoe.v62i1.1628 copyright notice: © 2020. the authors. licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. this book begins by describing the incredible richness of south africa’s wildlife, continues with its progressive decimation by hunters during the 19th century and ends with the remarkable contribution of the james stevenson-hamilton, against all odds, to reversing the trend towards decimation. it constitutes a reprint of a book originally published in the late 1950s, under a pseudonym and a different title, by the journalist and prolific writer a.p. cartwright. in a foreword to the reprint, the author’s son suggests that his father used a pseudonym because the book was written quickly, within approximately 12 weeks, without the diligent research that characterised his later works. the history of south africa’s wildlife has been often told – and much researched – so i confess to being initially sceptical as to the value of a book that is many years out of date and possibly inadequately researched. in fact, i was pleasantly surprised, the book provides an essentially true, lively and highly readable account, based on a well-chosen bibliography. the personal recollections of cartwright’s son yield an enchanting insight into his father’s state of mind when writing the book. cartwright commenced work on it shortly after being dismissed from a stressful, tiring position as editor of the rand daily mail, and was visibly rejuvenated and enthused by the task of writing. cartwright’s creative joy is reflected in the vibrant, engaging style which he maintains throughout the work. the book contains inaccuracies and errors of interpretation, some of which stem from the fact that it is old and pre-dates much relevant research into wildlife conservation and its history in south africa. for example, cartwright suggests that the tusklessness of addo elephants was because of an adaptation to the particular vegetation in the area, whereas more recent research has indicated that it is probably as a result of non-selective genetic changes resulting from the population’s isolation, small size and population bottlenecks during the 1800s and 1920s (whitehouse 2002). following misconceptions prevalent at the time, cartwright gave paul kruger too much credit for the establishment of the sabi game reserve (now part of the kruger national park) towards the end of his period as president of the republic of south africa. carruthers (1995) showed that the main motivator for the sabi game reserve was not kruger, but various other members of the volksraad, notably r.k. loveday. carruthers exposed the politically driven myths surrounding kruger’s supposed role in initiating what later became the kruger national park. she furthermore presented evidence that would undoubtedly have interested and amused cartwright – namely that stevenson-hamilton shrewdly used the political appeal of the mythology, which he called the ‘kruger stunt’, even though he was well aware of the truth (carruthers 1995:61). another minor error is that de kuyper’s expedition into what is now the crocodile bridge area of the kruger national park occurred in 1725 (punt 1958), and not in 1825 (p. 111). inaccuracies such as these are understandable given that cartwright’s aim was to inspire and amuse, rather than to produce a work of complete and detailed scholarship. nevertheless, the book has potential value for scholars. for example, cartwright gave a fascinating account of the life and death of the colourful colonel steinaecker, supported by details from kruger park ranger harry wolhuter, who was a former member of steinaecker’s horse. information from cartwright’s original book is quoted in van vollenhoven’s (2014) report on an archaeological investigation of steinaecker’s horse in the kruger national park. apart from steinaecker, cartwright related an amusing and lively account of the lives and deeds of a selection of 19thand early 20th-century hunters and travellers: william burchell, cornwallis harris, gordon cumming, frederick selous, major pretorius, all of whom contributed significantly to our knowledge of an exciting fauna as it was in the past, and some of whom contributed even more significantly to its decimation. the best part of the book is devoted to its hero, james stevenson-hamilton, self-effacing but determined, shrewdly diplomatic, a hugely persuasive and effective leader, whose work laid firm and enduring foundations for the kruger national park. an energetic, heart-warming story by a gifted storyteller; it will appeal even to those already familiar with the story, even now 60 years after its original publication. references carruthers, j., 1995, the kruger national park: a social and political history, university of natal press, pietermaritzburg, viewed 27 february 2020, from http://www.environmentandsociety.org/node/4665. punt, w.h.j., 1958, ‘die verkenning van die krugerwildtuin deur die hollandse oos-indiese kompanje, 1725’, koedoe 1, 1–18. https://doi.org/10.4102/koedoe.v1i1.863 van vollenhoven, a.c., 2014, a historical archaeological investigation of the ngotso mouth camp of steinaecker’s horse, close to balule, kruger national park, report no. ii, viewed 27 february 2020, from https://sahris.sahra.org.za/heritage-reports/ngotso-mouth-ii. whitehouse, a.m., 2002, ‘tusklessness in the elephant population of the addo elephant national park, south africa’, journal of zoology 257(2), 249–254. https://doi.org/10.1017/s0952836902000845 filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 page 3 page 4 koedoe 61-1_2019_contents.indd http://www.koedoe.co.za open access table of contents i original research estimating conservation effectiveness across protected areas in limpopo province, south africa luna milatović, brandon p. anthony, anthony swemmer koedoe | vol 61, no 1 | a1530 | 19 june 2019 original research changes in bird assemblages because of vegetation homogenisation in communal livestock systems mathew a. harris, michael d. voysey, samantha-leigh jamison, michelle greve koedoe | vol 61, no 1 | a1543 | 17 july 2019 original research a bimodal pollination system enhances reproductive potential of translocated populations of an endangered grassland succulent peter g. jansen, stefan j. siebert, frances siebert, johnnie van den berg, anine jordaan koedoe | vol 61, no 1 | a1562 | 14 august 2019 original research measuring kruger visitors’ place attachment to specific camps anneli douglas, jan-albert wessels, jenny pope, angus morrisonsaunders, mike hughes koedoe | vol 61, no 1 | a1559 | 17 september 2019 original research outbreak of avian botulism and its effect on waterbirds in the wilderness lakes, south africa ian a. russell, rodney m. randall, david zimmerman, danny govender koedoe | vol 61, no 1 | a1553 | 31 october 2019 original research ecology of palustrine wetlands in lesotho: vegetation classification, description and environmental factors peter chatanga, erwin j.j. sieben koedoe | vol 61, no 1 | a1574 | 31 october 2019 checklist a checklist of the termites of kruger national park, south africa brianna m. lind, andrew b. davies koedoe | vol 61, no 1 | a1531 | 09 april 2019 correction corrigendum: good governance and tourism development in protected areas: the case of phong nha-ke bang national park, central vietnam anna hübner, tuan phon ly, trương s.h. châu koedoe | vol 61, no 1 | a1601 | 06 december 2019 reviewer acknowledgement koedoe | vol 61, no 1 | a1611 | 11 december 2019 79 89 107 120 131 144 160 165 166 page i of i table of contents i essay koedoe: african protected area conservation and science – a retrospection: 1958 to 2018 jane carruthers, llewellyn c. foxcroft koedoe | vol 61, no 1 | a1556 | 08 april 2019 short communication reproductive biology of the sausage tree (kigelia africana) in kruger national park, south africa jah namah, jeremy j. midgley, laurence m. kruger koedoe | vol 61, no 1 | a1512 | 29 april 2019 short communication the genetic status of the introduced giraffe population in central south africa marieka e. van niekerk, francois deacon, paul j. grobler koedoe | vol 61, no 1 | a1570 | 16 september 2019 review article the management dilemma: removing elephants to save large trees michelle d. henley, robin m. cook koedoe | vol 61, no 1 | a1564 | 15 august 2019 original research use of a rapid roadside survey to detect potentially invasive plant species along the garden route, south africa johan a. baard, tineke kraaij koedoe | vol 61, no 1 | a1515 | 28 january 2019 original research monitoring consumptive resource use in south african national parks wessel vermeulen, nicola van wilgen, kyle smith, mbulelo dopolo, louise swemmer, wendy annecke, hugo bezuidenhout, graham durrheim, nick hanekom, howard hendricks, melodie mcgeoch, ntombizodwa ngubeni, alexis symonds koedoe | vol 61, no 1 | a1516 | 30 january 2019 original research the feasibility of national parks in south africa endorsing a community development agenda: the case of mokala national park and two neighbouring rural communities hendri coetzee, werner nell koedoe | vol 61, no 1 | a1470 | 28 february 2019 original research vachellia erioloba dynamics over 38 years in the kalahari gemsbok national park, south africa helga van der merwe, noel van rooyen, hugo bezuidenhout, jacobus du p. bothma, margaretha w. van rooyen koedoe | vol 61, no 1 | a1534 | 29 april 2019 1 7 14 21 33 43 54 67 vol 61, no 1 (2019) issn: 0075-6458 (print) | issn: 2071-0771 (online)koedoe overview and literature analysis status and trends in koedoe’s global reach: 1958–2016 acknowledgements references about the author(s) llewellyn c. foxcroft conservation services, south african national parks, skukuza, south africa centre for invasion biology, stellenbosch university, south africa marna herbst conservation services, south african national parks, cape research centre, south africa citation foxcroft, l.c. & herbst, m., 2017, ‘status and trends in the global growth of koedoe between 1958 and 2016’, koedoe 59(1), a1500. https://doi.org/10.4102/koedoe.v59i1.1500 editorial status and trends in the global growth of koedoe between 1958 and 2016 llewellyn c. foxcroft, marna herbst copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. the process of scientific publishing is highly competitive and the prestige associated with journals is a key consideration of where authors choose to publish their research. journals need to provide relevant, high-quality publications that add to the body of literature to provide updated information upon which evidence-based management can be implemented. in addition, scientists want their work to be accessible and with the evolution of web-based publishing, articles that were difficult to access in the past can now be made immediately available worldwide. to remain relevant and expand globally beyond its previous reach, koedoe launched its online platform in 2008. since then it has shown strong annual growth rates, which is illustrated by the number of articles being downloaded more than 265 000 times in 2016 alone. in addition, by 2016, the total number of all koedoe articles downloaded exceeded 1.3 million. the launch of koedoe in 1958 was aimed at providing a platform to capture important baseline information and research specifically on protected areas and national parks in south africa. the focus has expanded to include science across the african continent, addressing important management questions both in the biophysical sciences and more recently socio-ecological and economic studies. to date, about 1250 articles have been published in koedoe. publishing koedoe as a fully open access online journal also brought with it additional opportunities. for example, all articles from the first issue of koedoe in 1958 were scanned and included in the koedoe website, making over 50 years’ worth of scientific knowledge easily available. the importance of making past research available can be illustrated by the example of the focus around savanna fire ecology, a key research theme in the kruger national park. the third article published in koedoe was a report on fire research in the kruger national park (inleidende verslag oor veldbrandnavorsing in die nasionale krugerwildtuin; van der schijff 1958). this information would not have been easily accessible and knowledge may have been lost; however, since 2008 this article has been viewed about 4456 times on the website. the increasing visibility and use of articles positioned koedoe to be able to apply for accreditation with international bodies that use various metrics to benchmark journals. one of the frequently used benchmarks is clavariate analytics journal citation reports (formerly thomson-reuters web of science). koedoe received its first impact factor in 2013 with 1.48, and has since fluctuated between 1.48 and 0.52. this article provides (1) a brief overview and literature analysis of articles published in koedoe since 1958 and (2) an update on the status and trends of readers interacting with the koedoe website and articles being downloaded. most of the results are only for data up to 2016 to include a complete year; however, in some cases, information from 2017 is also given. overview and literature analysis a bibliometric analysis of keywords that have featured in koedoe articles since 1958, summarised by decades, provides some insights into the themes that were researched over time. keywords were taken from the titles of all articles and grouped a priori to show patterns of priorities. trends indicate that articles have consistently shown an increase in the number of downloads, even when accounting for the number of articles published in that decade (figure 1). for example, except for 1958–1969, the highest number of downloads was for articles published between 2000 and 2009; however, the lowest number of articles were published during this period. although the period from 2010 to 2019 is not complete and only 142 articles have been published to date, the number of downloads is already higher than pre-2000 publications. this also illustrates that while koedoe has maintained a constant level of interest, it suggests an increase in readership and the value of articles. figure 1: the number of downloads divided by the number of articles published per decade. values above the columns indicate the numbers of articles published per decade. encouragingly, articles published before 2000 are still consistently downloaded, showing the importance of accessibility to the information that has been published in koedoe in the past (figure 2). the number of articles published per year has fluctuated, often when special issues are published. however, since the late 1980s, there have been, on average, more than 20 articles a year. figure 2: the number of downloads divided by the number of articles published per year. the number of articles downloaded/articles published from about 2013 onwards have not had enough time to give an accurate reflection. an analysis of trends in keywords extracted from the titles of articles published in koedoe provides an indication of some of the priorities over time (figure 3). publication of inventories has remained important throughout the history of koedoe, with species lists being continuously updated as poorly surveyed areas or taxa receive attention. water-related themes became a major focus area in the 1990s, linked in part to the kruger national park rivers research programme (knprrp; mcloughlin et al. 2011) and the ongoing concern around river health. this can be seen with the increasing amount of work around fish monitoring. adaptive management only emerged between 1990 and 2009, partly as an outcome of the knprrp, but expanded to become a major focus area since 2010 (roux & foxcroft 2011). as part of the development of knowledge on wildlife management, animal behavioural studies remained important themes until 2000–2009 era. this evolved more broadly into biodiversity studies from about 2000 and has continued growing as an area of interest. figure 3: bubble plot showing the most frequently listed words in article titles, analysed per decade. the size of the circle indicates the frequency of keywords as an indication of active fields of research. analysing the origin of the research published provides indirect insight into two aspects: (1) for sanparks especially, priority research sites over time and (2) in general, the distribution of scientists across africa who have published in koedoe. articles have originated overwhelmingly from the kruger national park, and until the end of the 1990s also in the kalahari gemsbok national park, although with a declining focus from the latter in recent years (figure 4). in the first 10 years of koedoe, articles largely originated from nine protected areas or national parks. since the start of koedoe, articles have originated from at least 58 different areas, and from 2010 from 30 different protected areas. figure 4: bubble plot showing the most frequently listed national parks or other research sites in article titles, analysed per decade. the size of the circle indicates the frequency of the protected area name as an indirect indication of key focus areas over time. status and trends in koedoe’s global reach: 1958–2016 koedoe continued to show substantial growth in 2016, with the number of articles downloaded from the entire collection (1958–2016) exceeding the 1.3 million milestone, having been downloaded a total of 1 311 840 times (only pdf and html, but excluding epub and xml versions, which totals 3046 417 downloads). this showed a 27.8% increase from the 2015 cumulative total of 945 950 times. the 2015 total was already a 30.1% increase from 660 472 in 2014. there was an increase in the number of papers downloaded annually, from 285 475 downloads in 2015 to 365 890 downloads in 2016 (figure 5). figure 5: the number of article downloads and visitors to the koedoe website. download data in 2010 were only available for part of the year and not recorded between 2007 and 2009. while the number of website visitors has decreased slightly, there were still more than 130 000 unique site visitors and 225 000 visitors, leading to about 800 000 page views. this indicates that while there are fewer site visits, more articles are being downloaded by these visitors. data from crossref (http://www.crossref.org) showed that koedoe received a total of 268 unique citations (articles cited for the first time) in 2016. although the number of citations accumulates with time, the 268 citations in 2016 have already surpassed the 200 citations in 2015. the citations to articles published in 2014 continued to grow steadily to 602. the article that received the most citations (100) in 2016 is ‘landscapes of the kruger national park’ (gertenbach 1983). data from google analytics provide insights into the distribution of countries from which site visits originate. the top 10 countries from which visits to the koedoe website originated accounted for 42 551 of the visits, with south africa representing approximately half of these. as koedoe aims to make science available across the continent, it is encouraging to see that nine african countries have all had more than 100 site visits. these include kenya, also being the fourth highest overall (1626), botswana (746) and zimbabwe (462) (figure 6). the number of visits to the website from the united states, united kingdom, australia, india and germany illustrates the widespread reach of koedoe. figure 6: the number of visitors to the koedoe website per country (google analytics). the darker shade of green shows the highest number of visitors by origin, with the lighter shade indicating the number of site visits from african countries. between 2008 and 2016, 173 articles were published, including a special issue on adaptive management in 2011 (19 articles) and a special issue on tourism in 2014 (10 articles). both of the special issues have received much interest, with the special issue on adaptive management (http://www.koedoe.co.za/index.php/koedoe/issue/view/82) being downloaded a total of about 212 700 times (as at september 2017). the more recent special issue on tourism (http://www.koedoe.co.za/index.php/koedoe/issue/view/86) has been downloaded about 63 800 times (as at september 2017). a third special issue, titled ‘natural resources research: implication for management and conservation’, was published during 2017. koedoe will soon be celebrating its 60th anniversary in the publication of protected area management and conservation science. many milestones have been reached, with numbers of downloads increasing annually, the publication of two highly successful special editions and the journal citation report impact factor. koedoe is growing in the publication of manuscripts that address pressing challenges to conservation across the continent, including adaptive management, water, tourism, climate change and social-ecological systems. koedoe is providing a niche for relevant publications that are widely accessed and highly visible in the management of protected areas and conservation science. acknowledgements we thank south african national parks for ongoing support for koedoe. we thank all editors and reviewers for their valuable time and expertise given to koedoe. we acknowledge the aosis editorial team for their support. sandra macfadyen conducted the analysis and provided figures 3 and 4. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this editorial. authors’ contributions l.c.f. and m.h. developed the concept. l.c.f. led the writing and m.h. gave input on the manuscript. references gertenbach, w.p.d., 1983, ‘landscapes of the kruger national park’, koedoe 26(1), 9–121. https://doi.org/10.4102/koedoe.v26i1.591 mcloughlin, c.a., deacon, a., sithole, h. & gyeduababio, t., 2011, ‘history, rationale, and lessons learned: thresholds of potential concern in kruger national park river adaptive management’, koedoe 53(2), art. #996, 1–27. https://doi.org/10.4102/koedoe.v53i2.996 roux, d.j. & foxcroft, l.c., 2011, ‘the development and application of strategic adaptive management within south african national parks’, koedoe 53(2), art. #1049, 1–5. https://doi.org/10.4102/koedoe.v53i2.1049 van der schijff, h.p., 1958, ‘inleidende verslag oor veldbrandnavorsing in die nasionale krugerwildtuin’, koedoe 1(1), 60–94. https://doi.org/10.4102/koedoe.v1i1.865 http://www.koedoe.co.za open access page 1 of 1 reviewer acknowledgement acknowledgement to reviewers in an effort to facilitate the selection of appropriate peer reviewers for koedoe, we ask that you take a moment to update your electronic portfolio on https://koedoe.co.za for our files, allowing us better access to your areas of interest and expertise, in order to match reviewers with submitted manuscripts. if you would like to become a reviewer, please visit the journal website and register as a reviewer. to access your details on the website, you will need to follow these steps: 1. log into the online journal at https://koedoe. co.za 2. in your ‘user home’ [https://koedoe.co.za/ index.php/koedoe/user] select ‘edit my profile’ under the heading ‘my account’ and insert all relevant details, bio statement and reviewing interest(s). 3. it is good practice as a reviewer to update your personal details regularly to ensure contact with you throughout your professional term as reviewer to koedoe. please do not hesitate to contact us if you require assistance in performing this task. publisher: publishing@aosis.co.za tel: +27 21 975 2602 tel: 086 1000 381 the editorial team of koedoe recognises the value and importance of the peer reviewer in the overall publication process – not only in shaping the individual manuscript, but also in shaping the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely, productive peer review process. we would like to take this opportunity to thank all reviewers who participated in shaping this volume of koedoe. we appreciate the time taken to perform your review(s) successfully. alan k. whitfield alexander christianini althea l. davies andrew taylor beanelri b. janecke benjamin j. wigley betsie milne bram van schoenwinkel brett summerell bronwyn kirby-mccullough caswell munyai christo fabricius corli wigley-coetsee dave balfour david le maitre duan biggs elmarie kotze elmarie slabbert gabriela bucini george branch george bredenkamp gijsbert hoogendoorn graham alexander gregor feig haemish melville helen suich helga van coller hendrik sithole hugo bezuidenhout ian a. russell james pryke jan a. venter jessica cockburn johan van tol joyce jefwa kaera coetzer keith bosak kevin p. kirkman kolobe mmonwa laura becerra laven naidoo leanne k. faulks lesego khomo leslie r. brown liandi slabbert llewellyn c. foxcroft louis du preez manqhai kraai matthew bird matthys a. dippenaar michael bates miranda deutschländer mmoto masubelele mohlamatsane mokhatla nick hanekom nicola j. van wilgen pablo gonzález-moreno peet van der merwe petro marais robert guldemond robert j. scholes rudzani a. makhado sandra macfadyen shaun levick silvia giulio stefan j. siebert sue taylor teresa coutinho theuns de klerk tommie steyn tshoanelo miya wayne twine werner voigt willem froneman http://www.koedoe.co.za� https://koedoe.co.za� https://koedoe.co.za� https://koedoe.co.za� https://koedoe.co.za/index.php/koedoe/user https://koedoe.co.za/index.php/koedoe/user mailto:publishing@aosis.co.za book review review: conservation biology in sub-saharan africa book title: conservation biology in sub-saharan africa authors: john w. wilson & richard b. primack isbn: isbn paperback: 9781783747504 isbn hardback: 9781783747511 isbn digital (pdf): 9781783747528 isbn digital ebook (epub): 9781783747535 isbn digital ebook (mobi): 9781783747542 isbn digital (xml): 9781783747559 publisher: open book publishers (2019) £59.95 (r1399.13) *book price at time of review review title: review: conservation biology in sub-saharan africa reviewer: marna herbst1 affiliation: 1conservation services, cape research centre, south african national parks, cape town, south africa corresponding author: marna herbst, assoc.editor.koedoe@gmail.com how to cite this book review: herbst, m., 2020, ‘review: conservation biology in sub-saharan africa’, koedoe 62(1), a1635. https://doi.org/10.4102/koedoe.v62i1.1635 copyright notice: © 2020. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. there are many well-written textbooks on biodiversity conservation. what makes this textbook so valuable is that it is ‘the very first conservation biology textbook dedicated entirely to an african audience’. conservation biology in africa faces difficult and often unsurmountable challenges; the authors acknowledge these and describe it as a crises discipline! it might come across as overwhelming at first, however, the challenges are real and the authors manage to write with in-depth understanding of these issues, in an african context, with explanations that young and practising african conservation biologists will relate to. case studies and inspirational stories together with ‘an agenda for the future’ give guidance, foster a wider perspective and provide encouragement to readers. the first three chapters present the general introduction of conservation biology in sub-saharan africa, followed by a chapter linking the human component and well-being into the field of conservation biology. in chapter 1, the authors set the foundations for conservation biology, a multidisciplinary research field, where scholars need to reason within sound scientific principles and ethics to understand the broader societal environment and beliefs in which conservation biology functions. the authors reflect on the economic principles as well as the importance of communication to a wide audience. chapter 2 sets the scene for a sub-saharan context. it explains why the region is so diverse in terms of climate, geography and biodiversity. the authors succeed in discussing the historical context of africa, the socio-political relationships and the history of conservation. they are cognisant of the relationship that the people of africa have with nature, the use of resources and the influence of practices of the developed world. i must applaud the authors for opening discussions on the multitude of challenges the region is facing, and how solutions need to be sought that will benefit local communities in addition to conservation efforts to promote wildlife biodiversity. chapter 3 explores the knowledge conservationists and ecologists need to define biodiversity, from species to ecosystems, and the various methodologies used in conservation biology to determine diversity and patterns in biodiversity. in chapter 4, the connection and importance of people and nature, and how people value biodiversity, are described. this expands both the material benefits and regulating services to the non-material values and experiences people have in nature. several examples of ecosystem services are presented in this chapter (i.e. the importance of wetlands in flood control, insects in pollination, biocontrol and the enjoyment of activities like scuba diving and wildlife viewing – all very relevant examples). the authors also touch on the sub-discipline of environmental economics and the contribution of ecosystem services to global economies. after this broad introduction and developing an understanding of the underlining principles of conservation biology, the second part of the book follows with the threats to biodiversity and current scenarios, encouraging open-minded understanding and actions that are needed to address these challenges. chapter 5 starts with the very fitting title ‘scramble for space’, which emphasises the big conundrum of a growing human population as a driver for habitat loss, habitat degradation, fragmentation and the subsequent consequences that these factors have on remaining natural areas. the authors end with a request to readers to think about sustainable development and to reflect on the impact the expanding human population, with excessive consumption of natural resources, has on our lives. the current concerns about environmental change (anthropogenic climate change), and the impact of this on biodiversity, are discussed in chapter 6. drivers of climate change and the importance of understanding and predicting future changes on all ecosystems (terrestrial, freshwater and marine) are debated. the authors point out that climate change interacts with other drivers and that the resulting impacts, therefore, are exaggerated and that not all species will suffer to the same extent – some might even benefit. the important message, that climate change has the means to affect the world and economies on a global scale, is highlighted. the topics for discussion in this chapter are very relevant and encourage readers to think clearly and deeply about these scenarios and mitigation actions. chapter 7 continues with the threats to biodiversity from various forms of pollution, overharvesting, persecutions, invasive species, parasites and diseases. although the authors touch on poaching under the theme of overharvesting, and again later (chapter 12) under law enforcement, i do feel that this was the chapter where it may have been discussed in more depth. it is not just an overharvesting problem at issue, as the challenges involved are much deeper rooted in a social context. a stronger argument, with perhaps more controversial discussion points, could have highlighted this complex issue that is very explicit in african countries, the focus group of readers on conservation biology in sub-saharan africa. the last section of the book focusses on overarching solutions to the current biodiversity crisis. chapter 8 reflects on the meaning of an extinction, what species are at risk, how to classify them (iucn guidelines) and the use of these criteria to guide biologists and ecologists to focus on specific species and habitat-specific interventions and conservation efforts. in chapters 9 and 10, the authors discuss applied conservation biology principles that are exercised in the conservation of species (chapter 9) and ecosystems (chapter 10). the authors explain how ecosystems can be restored through rehabilitation to a specifically identified benchmark state. chapter 11 discusses how species’ extinctions can be prevented, understanding threatened species and populations, the factors influencing them, how to maintain viable populations, and the feasibility and ethics involved when considering breeding or re-introducing species back into natural areas. chapter 12 touches on environmental laws and regulations. this is probably one of the more difficult themes to cover, however, the authors provide a good and thorough overview. the importance of international treaties, which guide national and local laws and regulations, are explained. here the authors refer to poaching in a legislative context, which indeed needs discussing. i do however think it could have been emphasised more strongly in chapter 7. nonetheless, it is clear that the authors have a deep understanding of the challenges concerning the environmental laws and regulations of sub-saharan africa. in chapter 13, the importance of the prioritisation and declaration of protected areas (pas) and the different management options and authorities involved (government, community, private or co-managed) are reviewed. the authors also reflect on the design and effective management of pas. they acknowledge the significance of the interactions between pas and the surrounding local communities, other stakeholders and the benefits of sharing and/or co-management opportunities that are critical to the existence of these areas. in chapter 14, the authors focus their attention on the importance and values of areas and spaces outside pas and their contribution to functioning ecosystems. human welfare and biodiversity are interlinked, and green spaces in cities and environmental-friendly farming practices contribute to economic benefits. the fact that conservationists depend on many of the extractive industries makes it important to explore working together and finding mutual balances for a healthier environment and being mindful of sustainable development. the human–wildlife conflict in africa remains an important issue and it is imperative to find workable solutions (generally very case-specific). this chapter highlights several case studies on integrated conservation, development projects and community-based natural resource management. these examples are valuable for guidance in different scenarios that are playing out across sub-saharan africa. in summary, conservation biology is indeed facing massive challenges and uncertainties. the goals range from describing biodiversity to species and environmental protection and restoration in the face of global threats and economic markets. in the last chapter of conservation biology in sub-saharan africa, the authors set out an ‘agenda for the future’. sustainable development is essential as conservation biologists aim to address future needs without compromising the natural world. finding a balance and making informed decisions about how to address the challenges for the future is the key message in this textbook. being mindful that conservation funding continues to be limited, the authors show some optimism with ways to direct multilateral consortiums and non-governmental conservation organisations to support local conservation projects. building partnerships and forming relationships with local people and conservation professionals remains important. the focus on environmental education, leadership, successful communication, media and the involvement of general citizens remains imperative in order for conservation biologists to succeed. in conservation biology in sub-saharan africa, the authors, with their background and understanding, were able to bring the basic principles of conservation biology together, and lead readers to understand the challenges and threats we face in the conservation of sub-saharan africa today. through multiple case studies and guidance, it expands readers’ perspective through real-life scenarios and challenges. with their writing skills and expertise, the authors have been able to produce an outstanding textbook. i am convinced that this book will be a fundamental resource for future conservation biologists and ecologists. it is published under a creative attribution international license, making this textbook accessible to all, which will be especially valuable in sub-saharan african institutions. references about the author(s) abel ramoelo centre for environmental studies, department of geography, geoinformatics and meteorology, university of pretoria, pretoria, south africa citation ramoelo, a., 2022, ‘a reflection about the recent koedoe publications (volume 64, no 1, 2022)’, koedoe 64(1), a1736. https://doi.org/10.4102/koedoe.v64i1.1736 editorial a reflection about the recent koedoe publications (volume 64, no 1, 2022) abel ramoelo copyright: © 2022. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. welcome to volume 64, 1 (2022) of koedoe: african protected area conservation and science journal! it has been a privilege for me to oversee, edit and manage the collection of such astounding journal outputs. koedoe continues to be a premier open-access journal supporting and publishing research on the conservation of natural and cultural assets in protected areas, especially in africa. it is within the thrust of the journal to acknowledge the human and biophysical dimensions of many conservation issues by combining every relevant natural and social science discipline, especially interdisciplinary investigations and perspectives. as it was established in 1958, koedoe aims to provide a forum that facilitates the integration and sharing of learning about protected areas that have become increasingly important for maintaining habitats, species and ecosystem services and for ensuring socio-environmental resilience at many levels. it is still true that the management of conservation or protected areas cannot be successful without biodiversity monitoring and a holistic understanding of the system. the current issue provides yet another series of research outputs that are critical to inform decision-making processes pertaining to conservation management. a short communication, seven original research articles and two review article were published. the collection of published papers is mostly about the state of the ecosystems for the south african national parks’ conservation assets, with one paper reporting research results from one of the private game reserves. in the quest to understand river systems, evidence in the geomorphic characterization of sabie river through mapping geomorphic patterns and change, sediment dynamics and mobility as well as dating periods of sediment deposition (knight & evans 2022), diversity and distribution of benthic invertebrates, which are crucial for bio-monitoring (majdi et al. 2022), and the enigmatic floodplain water snake’s phylogenetic placement (keates et al. 2022) were presented. these three papers presented results covering broader biodiversity topics, including molecular, species and ecosystem levels for diagnosing the state and condition of the river systems. the authors found residues of tebuthiuron arboricide in the soils of mokala national park, which were postulated to be phytotoxic to the woody and grass species. the latter study provided information that could explain the failure of the natural recruitment of vegetation in sections of the park, where arboricides were applied and degraded woody plants more than a decade ago. looking at ecotourism, the potential for frogging (frog conservation) through ecotourism was explored. last but not the least, sustainability of solid waste management in protected areas was presented, focusing on the challenges and opportunities. it was evident that the implementation of solid waste management has the potential to contribute to pollution prevention, community upliftment and other secondary benefits, which could enhance conservation efforts. finally, the reflections of over 60 years of research on herbivore exclosures in the kruger national park (knp) were reported. herbivore exclosures are global research platforms that enable the training of ecologists, veterinarians and field biologists. they provide an opportunity to understand savanna dynamics that would otherwise have been hard to gain. the authors recommended continued use and support of exclosures as learning tools in complex african savannas. i hope you will enjoy reading this collection of original research, short communication and review articles, and i invite you to consider submitting your scientific papers to the upcoming issue. please distribute this issue within your networks. everyone at koedoe: african protected area conservation and science, including editors, reviewers and publishers, appreciates the support you as readers are giving us and providing an opportunity to offer the premier open access scientific outputs. references keates, c., conradie, w., dalu, t., dondofema, f., riddell, e.s. & wasserman, r.j. 2022, ‘phylogenetic placement of the enigmatic floodplain water snake, lycodonomorphus obscuriventris fitzsimons, 1964’, koedoe 64(1), a1698. https://doi.org/10.4102/koedoe.v64i1.1698 knight, j. & evans, m., 2022, ‘characterising the geomorphic dynamics of river systems: an example of the sabie river, south africa’, koedoe 64(1), a1700. https://doi.org/10.4102/koedoe.v64i1.1700 majdi, n., de necker, l., fourie, h., loggenberg, a., netherlands, e.c., bunte-tschikin, j. et al., 2022, ‘diversity and distribution of benthic invertebrates dwelling rivers of the kruger national park, south africa’, koedoe 64(1), a1702. https://doi.org/10.4102/koedoe.v64i1.1702 abstract introduction research method and design results discussion conclusion acknowledgements references appendix 1 about the author(s) peter g. jansen unit for environmental sciences and management, north-west university, potchefstroom, south africa stefan j. siebert unit for environmental sciences and management, north-west university, potchefstroom, south africa frances siebert unit for environmental sciences and management, north-west university, potchefstroom, south africa johnnie van den berg unit for environmental sciences and management, north-west university, potchefstroom, south africa anine jordaan laboratory of electron microscopy, chemical resource beneficiation, north-west university, potchefstroom, south africa citation jansen, p.g., siebert, s.j., siebert, f., van den berg, j. & jordaan, a., 2019, ‘a bimodal pollination system enhances reproductive potential of translocated populations of an endangered grassland succulent’, koedoe 61(1), a1562. https://doi.org/10.4102/koedoe.v61i1.1562 original research a bimodal pollination system enhances reproductive potential of translocated populations of an endangered grassland succulent peter g. jansen, stefan j. siebert, frances siebert, johnnie van den berg, anine jordaan received: 27 nov. 2018; accepted: 02 apr. 2019; published: 14 aug. 2019 copyright: © 2019. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract a major concern during the translocation of higher plant species is related to habitat suitability and the availability of pollination services. should these not meet the requirements of the plant, then successful reproduction and establishment cannot occur. we studied an endangered succulent, frithia humilis, which had previously been translocated to typical and atypical habitats, to assess the occurrence of potential pollinators at these sites. insects visiting f. humilis flowers and showing signs of pollen were captured, preserved and studied using a scanning electron microscope. pollen of f. humilis was searched for. abundance and diversity patterns of these pollen carriers across edaphic habitats of translocated populations were compared with those in a natural occurring population. pollination success of guilds was compared amongst translocated f. humilis populations by considering the number of seedlings in a new season. across f. humilis populations, hymenopteran species had the largest pollen loads, making this a melittophilous pollination system, typical for the aizoaceae. additionally, anemophilous syndrome was described for translocation sites which provide reserve pollinators. fruit formation was more favourable in suitable edaphic habitat on ecca sandstones. presence of seedlings at both translocation sites was indicative of successful pollination events of the self-incompatible species, but recruitment was proportionally higher in suitable habitat. habitat suitability, in the case of this threatened species, is more of a limiting factor than pollination services after a translocation event. conservation implications: translocations are fast becoming an attractive alternative for developers. this study cautions that the presence of pollinators and successful reproduction in translocated populations are only effective if the populations were translocated to an ideal habitat. keywords: aizoaceae; coal mining; conservation ecology; frithia humilis; mesembryanthemaceae. introduction frithia humilis burgoyne is an endangered, cryptic, dwarf succulent in the aizoaceae (burgoyne & krynauw 2005; burgoyne, smith & du plessis 2000). it is endemic to the rand highveld grassland of gauteng and mpumalanga in south africa, specifically the area between bronkhortspruit, ogies and middelburg. here it is restricted to flat sandstone plates of the dwyka and ecca formations in microhabitats comprising aggregates of weathered rock and organic materials (burgoyne & hoffman 2011). the succulent leaves of the species generally grow from beneath the soil surface (a typical window plant), seldom protruding more than 20 mm above ground level. the leaves are contractile, allowing the plant to retract into the soil where it is protected from desiccation and frost in the winter months (burgoyne et al. 2000). consequently, leaves of this species are only visible during active growth, specifically when flowering during the summer months (figure 1a). figure 1: frithia humilis plants in flower (a) and displaying a fresh (left) and an expired flower (right) on the same plant (b). in 2008, a population of f. humilis was discovered at a coal mine in the north of emalahleni after a mining license had already been granted (harris et al. 2014). in situ conservation was impossible because of the destructive nature of open-cast mining practices. considering the species’ endangered status (burgoyne & krynauw 2005), translocation was regarded as a last resort to save the population (godefroid et al. 2011; gordon 1994). translocation is the process whereby a population of living organisms is deliberately moved from one area to another suitable habitat within its existing distribution range (iucn 2013). consequently, conservation agencies translocated the population to pre-selected receptor sites (burgoyne & hoffmann 2011). a rapid pre-translocation habitat assessment was conducted to assess the suitability of receptor sites in terms of substrate, ecological integrity, genetic impacts and long-term protection (burgoyne & hoffmann 2011). a major critique of such rapid assessments is that in-depth assessments of pollinators are generally lacking (forup et al. 2008), putting the translocated population at risk, especially in the case of a self-incompatible species such as f. humilis (harris, van den berg & siebert 2016). however, it could be argued that by the very nature of a translocation the number of plant individuals should be high enough to attract pollinators from elsewhere in the distribution range and provide for their dietary requirements (menz et al. 2011). the translocation of f. humilis provided an ideal opportunity to test whether the pollination guild of the translocated populations is similar to that of a naturally occurring population. knowledge of this provides a measure to gauge whether a population is only reproducing because of chance pollination, or it persists because of effective pollination (montalvo et al. 1997). a poorly functioning pollination system can have several genetic consequences (armstrong & seddon 2008; montalvo et al. 1997; moritz 1999) and may sooner or later affect population viability, eventually leading to population extinction (kearns & inouye 1997). johnson (2010) has called for pollination research in south africa to improve our understanding of the role of pollination niches for the maintenance of plant species diversity in a rapidly changing environment. this study heeds the call to supplement existing knowledge concerning the pollinators of f. humilis (harris et al. 2016). it highlights the importance of pollination studies in translocation initiatives and provides insight into the status of a biotic factor that could limit reproductive success after a founder population was established in a harsh environment. flowers of frithia humilis the flowers of f. humilis are 15 mm–20 mm in diameter and white with a yellow centre (figure 1a, b). the petals, especially towards the tip, are occasionally tinged pink. flowers are either carried on very short stalks or are entirely stalkless. the pollen is yellow and tricolpate (figure 2). the sepals resemble the leaves and form a short tube (burgoyne et al. 2000). frithia flowers are self-incompatible (burgoyne et al. 2000), a common characteristic of south african aizoaceae (mayer & pufal 2007). after pollination the flowers turn yellow or pink before expiring (figure 1b), which may serve as an indication to pollinators that the flower is pollinated and without reward (burgoyne et al. 2000). figure 2: scanning electron micrographs of frithia humilis pollen grains in its distal polar-equatorial view (a) and polar view (b). based on familial pollination syndromes (hartmann 1991), f. humilis flowers could be melittophilous because they are wide open, petals are shiny, whitish and open diurnally (from mid-morning to mid-afternoon) for several days. pollen is less abundant than in most melittophilous species but easily accessible. further support for this syndrome is the self-incompatible sexual phase, with flowers reported to have a distinct male phase (protandrous) followed by a female phase approximately 4 days later (hammer 1995; hartmann 1991). stigmata are initially shorter than stamens, elongating and emerging only at the beginning of the female phase when the male phase ends. research method and design study area and site selection two sites were chosen as receptor sites based on pre-selected criteria (burgoyne & hoffmann 2011) for plants translocated from a coal mine (figure 3). half of the affected population was translocated to typical ecca group sandstone which corresponded with the geology of the donor site. the other half of the population was translocated to less ideal outcrops of the sedimentary wilge river formation of the waterberg group to test whether other related rock types could serve as an alternative should mining activities warrant future translocations (harris et al. 2014). figure 3: localities of the receptor sites (ecca sandstone at goedvertrouwdt farm and sedimentary rock at eagle’s rock private estate) and control population at ezemvelo nature reserve on ecca and dwyka sandstone in south africa. the donor population at inyanda coal mine is indicated and the shaded grey triangle represents the approximate endemic distribution of frithia humilis. distances between study populations are indicated. insect observations qualitative observations for pollinators were made at the receptor sites of goedvertrouwdt and eagle’s rock, and a large, natural population at ezemvelo nature reserve. as the flowers open only on sunny days, observations were made on non-overcast days of march 2016 and insects that visited the flowers were collected by hand. one observation session (of 3.5 h) was performed per day to coincide with the flowering time of the species for each population, on 3 suitable days (table 1). three 1-m2 patches with most flowers were visually estimated and demarcated at each site for observation. one person per patch observed for pollinators for the entire period of anthesis from 11:30 to 15:00. thus 10.5 h of observation was conducted per population. insects alighting or foraging on flowers were captured using an insect net and those species observed for the first time were euthanised in a killing jar with ethyl acetate. these specimens and a set of duplicates were then stored in individually marked containers for later identification and pollen assessments. repeat visitors of the same species were only counted and released. table 1: overview of time spent and flower visitors recorded during the survey. verification of pollen and insect identification all euthanised specimens were photographed with a nikon az1000 stereomicroscope to assist in identification and scanned for pollen grains. particular attention was paid to the head and legs, where pollen was most likely to be found. those specimens carrying pollen were sputter-coated with a thin gold-palladium layer and inspected under an fei quanta feg 250 scanning electron microscope (sem) and micrographs of relevant pollen loads were taken. duplicate insect specimens were pinned and labelled as prescribed by uys and urban (2006) and submitted to the biosystematics division of the plant protection research institute of the agricultural research council (arc) for identification (appendix 1). identification to species level was hampered by the taxonomic impediment whereby large numbers of species still require classification or have gone undiscovered (eardley, gikungu & schwarz 2009). verification of f. humilis pollen was based on pollen micrographs in burgoyne et al. (2000). plant counts in each of the three subplots per population, all the adult plants and flowers were counted according to the technique described by harris et al. (2014). the fruits were counted per subplot 30 days later. to estimate seed set per capsule in each subplot, 20 capsules were removed from plants bordering on the subplot (after 6 months) and the seeds were counted to determine the mean number of seeds per capsule. the sites were revisited after the first follow-up rains in october 2017 to count the seedlings per subplot. ethical considerations ethical clearance was obtained from the north-west university. this article followed all ethical standards for research without direct contact with human or animal subjects. results pollinator diversity a total of 46 insects were captured whilst alighting on f. humilis flowers. these comprised nine different species of which six were hymenopterans and three were dipterans (table 2). no lepidopterans were captured during this survey although species were observed making darting visits to frithia flowers. harris et al. (2016) did record two butterfly species alighting on frithia flowers but these were void of pollen. table 2: frithia humilis pollen carriers with collection locality and pollen load. at goedvertrouwdt, 14 individuals belonging to six species were recorded, with exoprosopa eluta (loew) (hymenoptera: bombyliidae) being the most frequent visitor (five records) of frithia flowers (table 2). species of lipotriches (gerstaecker) (hymenoptera: bombyliidae) and quartinia (andré) (hymenoptera: vespidae) were recorded the most at eagle’s rock private estate (table 2). at ezemvelo nature reserve, 26 individuals (four species) were captured, with amegilla fallax (smith) (hymenoptera: apidae) and e. eluta accounting for 65% of the visitations (table 2). pollen carriers harris et al. (2016) already provided evidence that the bees amegilla fallax and megachile niveofasciata (friese) (hymenoptera: megachilidae), together with the bee fly e. eluta, carry f. humilis pollen. scanning electron microscope evidence reported here was, therefore, specifically aimed at the five newly captured flower visitors and also on recaptured lipotriches sp. previously identified as a potential pollen carrier by harris et al. (2016). frithia pollen was found in varying amounts on the selected insect specimens and these species are therefore considered pollen carriers (table 2). some pollen grains were visible on notolomatia sp. (figure 4), on and immediately adjacent to the mouth parts. although only three pollen grains were found on the abdomen of paragus sp. (figure 5), a significant clump of frithia pollen was found directly on the anus of the specimen. pollen found on the quartinia sp. (figure 6) was sparsely distributed over its entire body. copious amounts of pollen were found beneath the mouth parts of the ammophila sp. (figure 7). halictidae specimens (seladonea sp. and lipotriches sp.) (figures 8 and 9) were carrying copious amounts of frithia pollen, particularly in their pollen baskets. figure 4: (a) stereomicrograph of notolomatia sp. (bombyliidae). scanning electron micrographs of several frithia humilis pollen grains on the eyes (b) and proboscis (c). figure 5: (a) stereomicrograph of paragus sp. (syrphidae). scanning electron micrographs of frithia humilis pollen grains above the anus (b), on the body (c) and on the anus (d). figure 6: (a, b) stereomicrographs of quartinia sp. (vespidae). scanning electron micrographs of several frithia humilis pollen grains on the body (c) and the head and antenna (d). figure 7: (a, b) stereomicrographs of ammophila sp. (sphecidae). scanning electron micrographs of frithia humilis pollen grains on the underside of the mouth parts (c) and on a maxillary palp (d). figure 8: (a) stereomicrograph of seladonea sp. (halictidae). (b, c) scanning electron microscope micrographs of frithia humilis pollen grains in the pollen baskets. figure 9: (a, b) stereomicrograph of lipotriches sp. (halictidae). (c, d) scanning electron micrographs of frithia humilis pollen grains in the pollen baskets. pollination system the two translocation sites and the control populations were situated in an area of approximately 200 km2. within this area, three bee species (a. fallax, lipotriches sp. and m. niveofasciata) were recorded from at least two of the sites (figure 10) and a bee fly species, e. eluta, was recorded from all three sites (table 2). the 39 flower visits by these four species accounted for 85% of all recorded visitations. this indicates that the pollination system of f. humilis is dominated by generalist bee pollinators that carry medium to high loads of pollen. although the bee fly was a more regular flower visitor (14 visits), its pollen load was low (table 2). reserve pollinators were observed at goedvertrouwdt and eagle rock private estate (figure 10). these pollinators were mostly flies and wasps and generally had medium to low pollen loads. a sweat bee (seladonea sp.) was only recorded from goedvertrouwdt and had a high pollen load. these five reserve pollinators accounted for 15% of flower visitations. figure 10: pollination system of translocated frithia humilis populations (g; e) in relation to a control population (enr). pollination success when mature plants, flowers, fruits and seedlings were compared across sites per 1 m2, ezemvelo nature reserve had significantly higher numbers, as would be expected for a natural population (table 3). however, no significant differences were recorded between the typical and atypical habitats. plants from ezemvelo nature reserve had the highest number of flowers as a percentage of total number of mature plants, with fruit set as a percentage of flowers the highest for the sites at goedvertrouwdt and ezemvelo nature reserve (table 3). as this study was conducted 7 years after translocation, it may be assumed that the seed bank would make a minor contribution to recruitment and that recent seed production is a consequence of recent flowering seasons. based on this premise, seedling emergence at goedvertrouwdt receptor site as a percentage of seed numbers was considerably higher than that of the control population and is indicative of viable seed (table 3). considering the 7-year time span since translocation, the presence of six pollinators (figure 10) and seedlings, it can be deduced that effective pollination is taking place. table 3: mean number of mature plants, flowers, fruit, seed and seedlings per 1 m2 for receptor sites and the control, as well as the flowering, fruiting and seeding percentage in brackets. discussion flower visitation and pollen carriers exoprosopa and notolomatia of the bombyliidae are primarily afrotropical in occurrence (greathead & evenhuis 2001). larvae are either predators or parasitoids, whilst adult flies in most genera are adapted for nectar feeding. female flies also feed on pollen for reproductive purposes and many of them have special adaptations for the collection of pollen (greathead et al. 2006). bombyliidae are most active during sunny middays that coincide with the anthesis of f. humilis. little is known about their role as pollinators; it is generally believed that most of them are generalists (szucsich & krenn 2002) although some specialised relationships are known (johnson & steiner 1997). the small number of pollen grains on the notolomatia specimen could be a consequence of the species or individuals (the inherent interested in pollen), the gender of the specimen (male flies may not be as interested in pollen, if at all) and the fact that the specimen was captured as soon as it landed on the flower, thereby shortening its potential visitation time. larvae of the hover fly genus paragus (diptera: syrphidae) are recognised as aphid predators (hayat & claussen 1997), whilst adult flies are nectar feeders. female flies consume pollen because it is a rich protein source necessary for reproduction (haslett 1989). even if pollen is consumed and digested by all syrphids, it still retains its shape after passing through the gut, making identification possible (holloway 1976). this explains the large cluster of pollen grains from f. humilis and other plant species on the anus of the specimen. the specimen only foraged for a short period before it was captured. although the pollen load was low on the insect itself, the pollen cluster on the anus suggested that it had recently fed on f. humilis pollen. the halictidae, represented by the species lipotriches and seladonea in this study, is the second largest family of bees with over 3500 members. many of them are pollen specialists with distinctive adaptations for collecting nectar and/or pollen from a small number of closely related plant species (danforth et al. 2008). in southern africa, these non-apis bees visit numerous plant families but the most predominant is the aizoaceae, along with asteraceae, fabaceae and zygophyllaceae (gess & gess 2004a). gess and gess (2004a) reported that as many as 13 different plant families may be visited by a single species but visitation records pointed to preferences for certain families. seladonea sp., for example, seemed to prefer asteraceae compared with other families. however, when considering the species richness and diversity of the halictidae, along with their polylectic manner of feeding, it can be concluded with a reasonable degree of certainty that they are important pollinators for many seed-bearing plants (dikmen 2007). all of the collected halictidae specimens displayed a typical pollen collection behaviour, which explains the copious amounts of pollen observed on their bodies and in their pollen baskets. ammophila is a genus of thread-waisted wasps in the sphecidae well known to be parasitic and predatory in nature. preferences for specific flowers are known amongst the genera of sphecidae but such relationships are not as strong as in bees or masarids (bohart & menke 1976). spechids, in general, feed on a variety of food including nectar, honeydew and bodily fluids of their prey. genera with short tongues seek nectar from flowers with short corollas such as the apiaceae, asteraceae, euphorbiaceae and polygonaceae (gess & gess 1991). ammophila has elongated mouthparts that enable it to feed on a greater variety of flowers. flowers known to be visited by ammophila include acanthaceae, apiaceae and lamiaceae (gess & gess 1991; herrera 1989; weaving 1989). the captured ammophila specimen was observed to make precise efforts to probe the nectaries of the f. humilis flower by circumnavigating the flower in such a way to ensure contact between the anthers and the underside of its mouth parts where pollen grains were found. all species of pollen wasps (vespidae), including those belonging to the genus quartinia, are nectar feeders and pollen collectors (gess & gess 2010). of the 40 species recorded to visit flowers, 75% preferred the aizoaceae. quartinia is also generally regarded as an effective pollinator for many aizoaceae species and numerous specialised relationships have been noted (gess & gess 2004b, 2010). during the survey, one of the quartinia specimens continued to forage in a f. humilis flower even after the hand net was placed over it and the flower. even though little pollen was found on the specimen, the fact that all species of quartinia are pollen feeders and collectors indicate that this genus may be an efficient reserve pollinator of f. humilis (gess & gess 2010). pollination system although few specimens (46 individuals) of pollen carriers were captured during this study, our observations provided enough evidence to support the hypothesis presented by hartmann (1991) that f. humilis belongs to the melittophilous syndrome, thus being pollinated mostly by bee and bee-like pollinators. this was especially the case for the natural f. humilis population at ezemvelo nature reserve, where bee pollinators were predominant. at translocated sites, additional observations were made of flies and the ammophila wasps which support the presence of anemophilous syndrome and imply a bimodal pollination system. these ‘reserve’ pollinators (johnson, harris & procheş 2009) associated with the anemophilous syndrome provide plausible alternatives for the translocated populations. however, it must be cautioned that even though specimens might carry f. humilis pollen, it does not conclusively prove that any of these species are pollinators. in addition, the amount of pollen on a pollinator’s body is regarded as an unreliable determinant of efficiency (adler & irwin 2006). identifying a primary pollinator depends on the pollination efficiency of the species, which is defined as the amount of con-specific pollen transferred to a stigma in any one visit (inouye et al. 1994). however, determining pollination efficiency is problematic because pollination is a highly variable ecological interaction and can be influenced by factors such as pollinator abundance across years, flower visit duration and frequency, contact with flower parts as well as pollen removal and deposition (ivey, martinez & wyatt 2003). with this in mind, the bees amegilla fallax, megachile niveofasciata and lipotriches sp. that form part of the melittophilous syndrome are cautiously proposed as primary pollinators of f. humilis because of their higher visitation frequencies and larger pollen loads. the sweat bee, seladonea sp., could also be a primary pollinator because of its higher observed pollen loads, despite only being observed twice. whilst pollen load may not be a direct indication of pollination efficiency, the larger pollen loads of bees compared with other f. humilis visitors were particularly evident on sem micrographs. the number of observations recorded as visitation frequency is also in favour of bees compared with the various other species. one generalist pollinator, e. eluta, was observed at all sites but is likely an inefficient pollinator because of the limited amount of pollen found only on the eyes (harris et al. 2016). further evidence for bee pollination is supported by the pollination system of a closely related species, bergeranthus multiceps (aizoaceae), from spring and autumn rainfall areas of the eastern cape province, south africa. this plant is also self-incompatible and flowers open between 15:30 and 18:30. bees were determined to be the most important pollinators to this species, placing it in the melittophilous syndrome (peter et al. 2004). other potential pollinators included butterflies, bees and hover flies belonging to the families bombyliidae, tachinidae and syrphidae (peter et al. 2004). this study and harris et al. (2016) made similar observations for f. humilis. pollination syndromes may maximise successful pollination by a specific species whilst still reserving the option for chance pollination by a less suitable species. the non-melittophilous frithia pollen carriers are suggested to be reserve pollinators because of their frequent visits and exclusive occurrence in translocated populations. they are primarily wasp and fly species that form part of an anemophilous syndrome. whilst bees may be primary pollinators, the small size and isolation of f. humilis populations supports the idea that f. humilis is a generalist rather than a specialist, and it can rely on reserve pollinators as its flower structure allows pollination by a different syndrome. a bimodal pollination system has also been identified previously for a member of the ruschioideae in the aizoaceae (zietsman 2013). pollination efficacy pollination was equally successful at the natural and translocated goedvertrouwdt populations although flower density differed (harris et al. 2014). this is in accordance with the flower density hypothesis (pufal et al. 2008). flower density is known to have a strong influence on pollinator visits (kunin 1997) but the size of the population had little effect on pollination efficacy. in other words, more flowers do not imply higher proportional frequencies of pollination. seedling recruitment per seed quantity was the highest for goedvertrouwdt, but this could be ascribed to lower plant densities and more suitable niches (eriksson & ehrlen 1992). eagle’s rock has a different geology and as expected did not have ideal conditions for seedling germination and establishment (harris et al. 2014). the presence of seedlings, therefore, is no indication of pollination efficacy but only that successful pollination takes place. this indicates that at least some of the pollen carriers recorded in this study are also pollinators. implications for translocation science this study showed that plant populations translocated to atypical habitats and geology had lower levels of fruit formation despite having higher levels of flowering compared to translocated populations of typical habitat. site selection is important and should be performed carefully to avoid plant stressors and nutrient deficiencies which may affect plant and flower health, and the availability of pollen (mayer 2004). reduced flowering can affect the number of pollinators that may visit a population and might limit outcrossing in small populations resulting in pollen transfer between relatives or not being pollinated at all (kearns & inouye 1997). the sparsely populated translocated sites had higher levels of fruit set and seedling occurrence per individual than the natural population. this shows that it is important to consider transplant density for xenogamous species, as too high densities influence the available pollinators and could result in many unpollinated flowers, which could restrict geneflow in the newly established population. pufal et al. (2008) suggested that decreased density of flowers increases pollination success and fruit set per plant as pollinators are known to be scarce, with a few taxa accounting for most of the flowers at a site (herrera 1989). translocation requires relocating a species to areas within its existing distribution range (iucn 2013). this study has shown that generalist pollinators of f. humilis occur throughout its natural distribution range and are common and mobile enough to locate newly established populations. translocation is therefore a low risk conservation option in terms of generalist plant species. pollination is further enhanced by the bimodal pollination system that is characteristic for some of members of the aizoaceae (zietsman 2013), which makes them resilient to shifts within their distribution range. conclusion nine arthropod species have now been confirmed as f. humilis pollen carriers. hymenopteran species had the largest pollen loads in pollen baskets and on front legs. this positioning of pollen ensures contact with the stigmata of the open flowers of f. humilis and could result in effective pollination. the pollination system is melittophilous as would be expected for the family, with an anemophilous syndrome providing reserve pollinators at translocation sites. the presence of seedlings at both translocation sites was indicative of successful pollination by insects of this self-incompatible species. pollination of flowers was proportionally similar between the varying population densities of natural and translocated populations on ecca sandstone. this indicated that flower density did not influence fruit formation in this species. when the two translocated populations were compared, it was evident that fruit formation was more effective on ecca sandstone. this indicated that reproductive success of this edaphic specialist decreases on non-typical rock habitats. micro-habitat conditions, planting density and locality within the species’ natural distribution range require an in-depth understanding before sites are selected to enhance pollination probability. however, even if predictions based on this are not accurate, it can be assumed that generalist and ‘reserve’ pollinators are widespread and abundant enough to locate translocated populations of f. humilis within the natural distribution range of the species. it can be concluded that pollinators are not limiting factors when this species is translocated. acknowledgements the authors thank dr duncan macfayden (manager: research and conservation at e. oppenheimer & son) for providing access to the frithia population at ezemvelo nature reserve. they are grateful to the landowners for providing access to translocated populations at eagle’s rock private estate and goedvertrouwdt farm and giving permission to collect insect specimens on their properties. mr scott van eeden prepared the locality map and mr koos smit photographed the species in habitat. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions p.g.j. conducted fieldwork and species identification and wrote part of the manuscript. s.j.s. designed the study, conducted initial surveys and wrote part of the manuscript. f.s. contributed to the writing of the manuscript and data analyses. j.v.d.b. performed sorting of arthropod material and coordinated the identification of pollinators. a.j. conducted scanning electron microscopy. funding information financial assistance for this study was provided by the national research foundation (grant uid: 103370). data availability statement data sharing is applicable to this article as new data were created or analysed in this study. disclaimer opinions expressed and conclusions arrived at in the article are those of the authors and are not necessarily to be attributed to the national research foundation (nrf). references adler, l.s. & irwin, r.e., 2006, ‘comparison of pollen transfer dynamics by multiple floral visitors: experiments with pollen and fluorescent dye’, annals of botany 97(1), 141–150. https://doi.org/10.1093/aob/mcj012 armstrong, d.p. & seddon, p.j., 2008, ‘directions in reintroduction biology’, trends in ecology and evolution 23(1), 20–25. https://doi.org/10.1016/j.tree.2007.10.003 bohart, r.m. & menke, a.s., 1976, sphecid wasps of the world: a generic revision, university of california press, berkeley. burgoyne, p.m., smith, g.f. & du plessis, f., 2000, ‘notes on the genus frithia (mesembryanthemaceae) and the description of a new species, f. humilis, in south africa’, bothalia 30(1), 1–7. https://doi.org/10.4102/abc.v30i1.531 burgoyne, p.m. & krynauw, s., 2005, frithia humilis burgoyne, national assessment: red list of south african plants version 2017.1, south african national biodiversity institute, viewed 19 march 2018, from http://redlist.sanbi.org/species.php?species=126-3. burgoyne, p.m. & hoffmann, a., 2011, ‘frithia humilis – notes on the translocation of a red listed succulent’, aloe 48(2), 38–42. danforth, b.n., eardley, c., packer, l., walker, k., pauly, a. & randrianambinintsoa, f.j., 2008, ‘phylogeny of halicitidae with an emphasis on the african halictinae’, apidologie 39(1), 86–101. https://doi.org/10.1051/apido:2008002 dikmen, f., 2007, ‘the role and the importance of the family halictidae (apiformes: apoidea) in pollination of natural and agricultural vegetation’, mellifera 7, 16–19. eardley, c.d., gikungu, m. & schwarz, m.p., 2009, ‘bee conservation in sub-saharan africa and madagascar: diversity, status and threats’, apidologie 40(3), 355–366. https://doi.org/10.1051/apido/2009016 eriksson, o. & ehrlén, j., 1992, ‘seed and microsite limitation of recruitment in plant populations’, oecologia 91(3), 360–364. https://doi.org/10.1007/bf00317624 forup, m.l., henson, k.s., craze, p.g. & memmott, j., 2008, ‘the restoration of ecological interactions: plant–pollinator networks on ancient and restored heathlands’, journal of applied ecology 45(3), 742–752. https://doi.org/10.1111/j.1365-2664.2007.01390.x gess, f.w. & gess, s.k., 1991, ‘a preliminary survey of the aculeate wasps and the bees of the lower reaches of the nossob river valley, kalahari gemsbok national park, south africa’, koedoe 34(2), 77–88. https://doi.org/10.4102/koedoe.v34i2.424 gess, s.k. & gess, f.w., 2004a, ‘a comparative overview of flower visiting by non-apis bees in the semi-arid to arid areas of southern africa’, journal of the kansas entomological society 77, 602–618. https://doi.org/10.2317/e7.1 gess, s.k. & gess, f.w., 2004b, ‘distributions of flower associations of pollen wasps (vespidae: masarinae) in southern africa’, journal of arid environments 57(1), 17–44. https://doi.org/10.1016/s0140-1963(03)00093-4 gess, s.k. & gess, f.w., 2010, pollen wasps and flowers in southern africa, sanbi biodiversity series 18, south african national biodiversity institute, pretoria. godefroid, s., piazza, c., rossi, g., buord, s., stevens, a.d., aguraiuja, r. et al., 2011, ‘how successful are plant species reintroductions?’, biological conservation 144(2), 672–682. https://doi.org/10.1016/j.biocon.2010.10.003 gordon, d., 1994, ‘translocation of species into conservation areas: a key for natural resource managers’, natural areas journal 14(1), 31–37. greathead, d.j. & evenhuis, n.l., 2001, ‘annotated keys to the genera of african bombylioidea (diptera: bombyliidae; mythicomyiidae)’, african invertebrates 42(1), 105–224. greathead, d.j., lovell, s., barraclough, d., slotow, r., hamer, m. & herbert, d., 2006, ‘an ecological and conservation assessment of the fauna of bombyliidae (diptera) occurring in the mkhuze, phinda and false bay reserves, kwazulu-natal, south africa’, african invertebrates 47(1), 185–206. hammer, s., 1995, ‘mastering the art of growing mesembs’, cactus and succulent journal 67, 195–247. harris, e., siebert, s.j., smit, j.h.l. & van den berg, j., 2014, ‘translocation of an endangered succulent plant species from sandstone outcrops earmarked for coal mining’, journal of the southern african institute of mining and metallurgy 114(11), 904–912. harris, e., van den berg, j. & siebert, s.j., 2016, ‘notes on the pollinators of frithia humilis burgoyne, an endangered highveld succulent’, african entomology 24(1), 250–256. https://doi.org/10.4001/003.024.0250 hartmann, h.e.k., 1991, ‘mesembryanthema’, contributions from the bolus herbarium 13, 75–157. haslett, j.r., 1989, ‘adult feeding by holometabolous insects: pollen and nectar as complementary nutrient sources for rhingia campestris (diptera: syrphidae)’, oecologia 81(3), 361–363. https://doi.org/10.1007/bf00377084 hayat, r. & claussen, c., 1997, ‘a new species and new records of the genus paragus latreille, 1804 from turkey (diptera: syrphidae)’, zoology in the middle east 14(1), 99–108. https://doi.org/10.1080/09397140.1997.10637710 herrera, c.m., 1989, ‘pollinator abundance, morphology, and lower visitation rate: analysis of the “quantity” component in a plant–pollinator system’, oecologia 80(2), 241–248. https://doi.org/10.1007/bf00380158 holloway, b.a., 1976, ‘pollen-feeding in hover-flies (diptera: syrphidae)’, new zealand journal of zoology 3(4), 339–350. https://doi.org/10.1080/03014223.1976.9517924 inouye, d.w., gill, d.e., dudash, m.r. & fenster, c.b., 1994, ‘a model and lexicon for pollen fate’, american journal of botany 81(12), 1517–1530. https://doi.org/10.1002/j.1537-2197.1994.tb11462.x international union for conservation of nature (iucn), 2013, guidelines for reintroductions and other conservation translocations, iucn, gland. ivey, c.t., martinez, p. & wyatt, r., 2003, ‘variation in pollinator effectiveness in swamp milkweed, asclepias incarnata (apocynaceae)’, american journal of botany 90(2), 214–225. https://doi.org/10.3732/ajb.90.2.214 johnson, s.d. & steiner, k.e., 1997, ‘long-tongued fly pollination and evolution of floral spur length in the disa draconis complex (orchidaceae)’, evolution 51(1), 45–53. https://doi.org/10.1111/j.1558-5646.1997.tb02387.x johnson, s.d., 2010, ‘the pollination niche and its role in the diversification and maintenance of the southern african flora’, philosophical transactions of the royal society b 365(1539), 499–516. https://doi.org/10.1098/rstb.2009.0243 johnson, s.d., harris, l.f. & procheş, ş., 2009, ‘pollination and breeding systems of selected wildflowers in a southern african grassland community’, south african journal of botany 75(4), 630–645. https://doi.org/10.1016/j.sajb.2009.07.011 kearns, c.a. & inouye, d.w., 1997, ‘pollinators, flowering plants, and conservation biology’, bioscience 47(5), 297–307. https://doi.org/10.2307/1313191 kunin, w.e., 1997, ‘population size and density effects in pollination: pollinator foraging and plant reproductive success in experimental arrays of brassica kaber’, journal of ecology 85(2), 225–234. https://doi.org/10.2307/2960653 mayer, c., 2004, ‘pollination services under different grazing intensities’, international journal of tropical insect science 24(1), 96–103. https://doi.org/10.1079/ijt20047 mayer, c. & pufal, g., 2007, ‘investigation of breeding systems for four aizoaceae species in namaqualand, south africa’, south african journal of botany 73(4), 657–660. https://doi.org/10.1016/j.sajb.2007.06.001 menz, m.h.m., phillips, r.d., winfree, r., kremen, c., aizen, m.a., johnson, s.d. et al., 2011, ‘reconnecting plants and pollinators: challenges in the restoration of pollination mutualisms’, trends in plant science 16(1), 4–12. https://doi.org/10.1016/j.tplants.2010.09.006 montalvo, a.m., williams, s.l., rice, k.j., buchmann, s.l., cory, c., handel, s.n. et al., 1997, ‘restoration biology: a population biology perspective’, restoration ecology 5(4), 277–290. https://doi.org/10.1046/j.1526-100x.1997.00542.x moritz, c., 1999, ‘conservation units and translocations: strategies for conserving evolutionary processes’, hereditas 130(3), 217–228. https://doi.org/10.1111/j.1601-5223.1999.00217.x peter, c.i., dold, a.p., barker, n.p. & ripley, b.s., 2004, ‘pollination biology of bergeranthus multiceps (aizoaceae) with preliminary observations of repeated flower opening and closure’, south african journal of science 100(11–12), 624–629. pufal, g., mayer, c., porembski, s. & jürgens, n., 2008, ‘factors affecting fruit set in aizoaceae species of the succulent karoo’, basic and applied ecology 9(4), 401–409. https://doi.org/10.1016/j.baae.2007.06.001 szucsich, n.u. & krenn, h.w., 2002, ‘flies and concealed nectar sources: morphological innovations in the proboscis of bobyliidae (diptera)’, acta zoologica 83(3), 183–192. https://doi.org/10.1046/j.1463-6395.2002.00111.x weaving, a.j.s., 1989, ‘nesting strategies in some southern african species of ammophila (hymenoptera: sphecidae)’, journal of natural history 23(1), 1–16. https://doi.org/10.1080/00222938900770011 uys, v.m. & urban, r.p., 2006, how to collect and preserve insects and arachnids, 2nd edn., plant protection research institute, queenswood. zietsman, p.c., 2013, ‘reproductive biology of stomatium bolusiae (aizoaceae: ruschioideae)’, bothalia 43(1), 23–27. https://doi.org/10.4102/abc.v43i1.83 appendix 1 table 1-a1: specialist identifications of frithia humilis pollen carriers not considered in harris et al. (2016). abstract introduction research strategy existing schemata results: the people parks win-win framework discussion conclusion acknowledgements references about the author(s) dorothy r. queiros department of applied management, college of economic and management sciences, university of south africa, pretoria, south africa citation queiro, d.r., 2022, ‘people parks win-win framework: integrating components that can influence people-park relationships’, koedoe 64(1), a1723. https://doi.org/10.4102/koedoe.v64i1.1723 original research people parks win-win framework: integrating components that can influence people-park relationships dorothy r. queiros received: 28 mar. 2022; accepted: 29 aug. 2022; published: 26 sept. 2022 copyright: © 2022. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract protected areas are often surrounded by impoverished communities. biodiversity must be conserved while improving community well-being. greater insight is required into what influences pro-conservation attitudes and behaviour in these communities. much appears to rest on the relationships between protected area staff and local communities surrounding the parks, yet there is limited understanding of stakeholders’ perceptions and how to pragmatically achieve win-win solutions. with the current lack of a multidimensional framework to enhance understanding of this complex and dynamic relationships, this research aimed to construct a comprehensive integrated framework representing the components that can influence people-park relationships. the framework was constructed via a threefold approach, namely a broader literature review, a focused study of existing schemata and primary research regarding the attitudes and behaviour of three local communities bordering three different protected areas in south africa. the resultant people parks win-win framework consists of four layers (each with its own components): ‘external context’, ‘stakeholders’, ‘community beneficiation’ and ‘outputs’. its unique arrangement focuses on beneficiation, inclusion of more stakeholders and their characteristics, the centrality of relationships and demonstration of outputs (how preceding layers can culminate in win-wins and how pro-conservation attitudes and behaviour fit into this). a simplified framework is also provided, for stakeholders to superimpose their own characteristics, benefits, influences and beneficiation principles. this research draws on the work of others as well as primary research to produce this multidimensional framework capturing the influences on people-park relationships with a focus on achieving both community well-being and biodiversity conservation. conservation implications: win-wins for community well-being and biodiversity conservation are complex. yet potential exists for tangible and intangible beneficiation, which can foster positive attitudes resulting in pro-conservation behaviour and robust reciprocate relationships between parks and neighbouring communities. to this end, the framework serves as a practical tool for protected area managers and stakeholders involved in the people-park relationships, which can be customised to particular contexts. keywords: benefits; biodiversity conservation; local community well-being; people-park relationships; people parks win-win framework; pro-conservation attitudes. introduction the conservation crisis, together with impoverished communities on the verges of natural protected areas, creates a major challenge. the natural environment globally is losing biodiversity, highlighting the urgency to conserve protected areas in order to halt this loss (abrams et al. 2009; allen et al. 2019). ceballos et al. (2015:1) confidently conclude that there has been an ‘exceptionally rapid loss of biodiversity over the last centuries, indicating that a sixth mass extinction is already under way’, with a devastating loss of ecosystem services looming. the crisis, however, can be lessened through rapid intensified conservation efforts, but this window of opportunity is quickly closing (ceballos et al. 2015). at the same time, the majority of protected areas in africa are surrounded by poor communities (davies et al. 2014; shackleton et al. 2010) and are experiencing increasing illegal harvesting of natural resources (mutanga, muboko & gandiwa 2017). according to recent forecasts, by 2050, africa’s population is expected to double and half of that population will be under 25 years of age (baker 2015; united nations 2021). these youngsters will also carry the majority of the unemployment burden (african development report 2015). in south africa, where this research was conducted, the ‘triple problem’ of poverty, unemployment and inequality is commonplace (world economic forum 2019). the dual crises of poverty and biodiversity loss are therefore two distinct but connected concerns in the 21st century (davies et al. 2014; nyaupane & poudel 2011) – not only in africa, but globally. this linkage has resulted in attempts to decrease poverty through management systems whereby local community livelihoods are incorporated into action plans and locals become involved in and support the conservation of biodiversity (abrams et al. 2009; davies et al. 2014; stoll-kleemann 2005). in south africa, government is also increasingly highlighting the twofold achievement of peoples’ well-being and park conservation and the participation of local communities in this. for example, while comparing the national environmental management: protected areas act 57 of 2003 (south africa 2003) and the national environmental management: biodiversity act 10 of 2004 (south africa 2004) with the more recently released ‘norms and standards for the management of protected areas in south africa’ (south africa 2016), the responsibilities placed on protected areas to improve human well-being are much more distinct. these include community welfare, development, education, communication, use of natural resources, access to the land and long-term economic benefits. with stakeholders within protected areas worldwide, as well as in africa and south africa, working towards improved involvement of neighbouring local communities and with governments keenly advocating this, the integration of these two agendas is becoming increasingly pressing, but proving more challenging and costly than anticipated (adams 2013). yet this integration is essential as the future success of conserving wildlife and their habitats could depend on the attitudes and behaviour of communities living in and nearby protected areas (snyman 2014; tessema et al. 2007) and on the extent to which their lives are improved (diedrich 2007; hackel 1999). however, greater insight into what influences pro-conservation attitudes and behaviour is first required, that is, what causes local people to want to conserve and engage in pro-environmental behaviour? (berkes 2004; coria & calfucura 2012; imran, alam & beaumont 2014; kiss 2004; mbaiwa & stronza 2010; walpole & goodwin 2001). ferse et al. (2010:7) said: ‘a deeper understanding of what drives positive conservation behaviour and what hampers it, is needed’. a great deal appears to rest on the relationships between protected area staff and the local communities surrounding the parks. in south africa, despite growing literature regarding protected area management, there is still limited understanding of the perceptions of both park officials and local communities regarding these inter-relationships (thondhlana & cundill 2017). mccleave, espiner and booth (2006), who conducted a literature review of new zealand case studies, highlight the significance of this issue but add that there is a scarcity of theory on people-park relationships. mccleave et al. (2006) warn that failing to recognise this important relationships can hamper effective management, cause the community to resent conservation and result in reduced social well-being and loss of tourism opportunities. anthony (2021) who considers this relationships in the context of human-wildlife conflict and mutanga et al. (2017) who investigate a range of influences on this relationship, contend that a deeper understanding of what affects people-park relationships is required. stoll-kleemann (2005), reporting on the international governance of biodiversity (gobi) research project, which considered the integration of ecological and socio-economic approaches in improving protected area management, appeals for better relationships between protected areas and surrounding communities. these authors contend that pragmatic integration of the interests of both protected areas and neighbouring communities is an urgent requirement. pragmatic integration, however, requires achievement of the dual goals of biodiversity conservation and improvement in the well-being of communities bordering protected areas (davies et al. 2014; ferse et al. 2010). this is vital to impact positively on the biodiversity and poverty crises. yet, as pointed out by wali et al. (2017), there is much work needed on how to integrate biodiversity conservation with human well-being. in addition, solutions must be realistic, considering the very real constraints experienced by protected areas in south africa, and elsewhere. this research refers to achieving this pragmatic integration as aiming for win-wins between parks and people. furthermore, answers to these problems need to be represented in a format that is useful to practitioners in nature conservation and community development, local communities and academics. however, following a review of existing frameworks (outlined later in this article), it would appear that a comprehensive integrated framework or schema such as this does not exist. allendorf (2010) identifies this research gap: the lack of a common framework limits not only our understanding, but also the development of future research areas and approaches to balancing conservation and sustainable development around protected areas (pas). if we are to understand and address fundamental pa–people issues, we need a descriptive framework that enables the understanding of the relationships that people have with pas and that facilitates comparisons among pas. (p. 417) published in 2018, stone and nyaupane deliver a brief overview of different frameworks and models that link conservation, tourism and the livelihood of local people. they contend that existing frameworks: [a]re based on simple static concepts and fail to capture the complexity and dynamic nature of the relationships … moving forward in addressing this complexity, there is a need to overcome the development and reliance on simplified models of complex systems. (stone & nyaupane 2018:309) wali et al. (2017) also argue for more multidimensional tactics to address community well-being, arguing that economic indices alone are insufficient and exclude other interrelated components that also account for community well-being. the aim of this research was therefore to construct a comprehensive integrated framework representing the components that can influence people-park relationships. this research forms part of a larger study carried out (queiros 2020), wherein this framework was the final output. this article first outlines the research strategy used to construct the framework followed by examination of existing schemata. in the results section, the people parks win-win framework (ppww) is introduced, followed by a discussion on its distinctiveness and the various layers and components. the article is then concluded. research strategy figure 1 depicts the threefold approach used to construct the framework: a broad literature review, a focused study of existing schemata and primary research conducted in south africa regarding the attitudes and behaviour of three local communities bordering three different protected areas. each approach contributed elements to the final framework. existing schemata are discussed in detail as they inform the research gap. the current section briefly covers the design of the literature review followed by the methodology used in the primary research. figure 1: threefold approach to construct the people parks win-win framework. design of the literature review the literature review focused on the influences on pro-conservation attitudes and behaviour. the author sought to conduct a wide-ranging search so as to provide multiple lenses at the start of the research. this aligns with maxwell (2013) who warns researchers to include concepts from beyond the traditional field of study. the language of the sources was restricted to english. various keyword combinations were used in both google scholar and within the e-journals search engines of the university of south africa’s library. the researcher consulted the work of different researchers worldwide in the fields of tourism, the natural environment, development and social studies. a wide range of models and frameworks related to communities or communities and conservation were also consulted. the majority of sources consulted were in the fields of protected areas, biodiversity conservation and local communities. some focused entirely on these domains, while others incorporated different combinations of the following keywords: attitudes, behaviour, benefits, participation, poverty reduction and natural resource management. some of this research was from the perspective of a particular approach, such as sustainable tourism development, ecotourism or one of the numerous different approaches to community-based conservation and/or tourism. the literature review was updated after the collection of primary data, in order to include newer research. the findings from the literature review were clustered under the different foci of the research. a key investigation was the contrasting views on the different linkages between benefits and losses or costs, pro-conservation attitudes and pro-conservation behaviour. this included research where connections were made between benefits or costs and attitudes; research that had very specific links between benefits or costs and behaviour and between attitudes and behaviour; and studies that argue against the presence of links between benefits and pro-conservation attitudes and behaviour. a large section of the literature review documented the different benefits (tangible and intangible) received by local communities bordering protected areas as well as the losses or costs incurred by these communities because of proximity to a protected area (since these can influence attitudes and behaviour). finally, ‘other factors’ that were not benefits but could influence pro-conservation attitudes and behaviour were also considered. some examples include the presence of tourism, local participation, multiple livelihood strategies and building trust. the literature review informed the questions posed to the participants in the primary research. the full literature review as per queiros (2020) is beyond the scope of this article. however, the specific components from literature that were integrated into the framework, which is the focus of this article, are clearly discussed in the tables that form part of the discussion. primary research methodology the primary research was qualitative as the researcher sought to explore the feelings, opinions and perceptions of the participants in their own words (roulston 2014). while much conservation-related research is quantitative, this research was firmly positioned in the interpretivism or constructivism paradigm, which acknowledges the multiple subjective interpretations that individuals will have of their world (creswell 2014). in the context of african society and community, it was vital to work with social systems, where individuals’ lived realities are constructed by both individuals and groups, and discovered via the interactions within these social groups (bann 2001; lune & berg 2017). in this vein, and in order to hear the voices of the participants in ways with which they could identify, multiple methods were used to gather data. these methods were semi-structured individual interviews, focus group interviews, adapted nominal grouping technique and drawings (of the communities living area in contrast to the protected area). atlas.ti, as a data software package for qualitative research, was instrumental in organising and analysing the large body of data. data were collected at three case study sites within south africa between december 2015 and september 2017. each case study site comprises the following: (1) protected area where a tourism venture exists and (2) the local community located closest to the protected area (as these are usually the communities most affected by the protected area). in other words, within each case study, two groups of people (constituencies) formed part of the research. constituency 1 (c1) refers to the local community living closest to the protected area. constituency 2 (c2) denotes the protected area: conservation authorities (e.g. reserve management), conservationists, as well as other stakeholders that are involved in managing the tourism venture. participants were selected using non-probability purposive sampling. for c1, it was important that the group constituted a balance of participants in terms of age, gender and societal position. the three case studies varied from each other in the sense that each protected area had differing management models and ownership structures and were at different stages in the level of improvement in human well-being offered to the neighbouring community. using open-ended questions around several themes, the contrasting cases assisted the researcher in determining how very different contexts influence attitudes and behaviour related to conservation. this enabled the framework to be more comprehensive. the first case study site was dinokeng game reserve and kekana gardens community (13 c1 participants; 4 c2 participants) in the gauteng province. dinokeng is a public–private partnership between provincial government and landowners, with the latter running various tourism establishments varying from budget to luxury. regarding the level of improvement in human well-being for bordering communities, as a fairly new reserve opened in 2011 (dinokeng 2017), reserve management and landowners are still developing different economic and social well-being programmes. mkhambathi nature reserve and khanyayo community (19 c1 participants; 5 c2 participants) in the eastern cape province formed the second case. mkhambathi has been a nature reserve since 1982 (queiros 2000). it is a provincial reserve on land owned by the surrounding local communities, and hence jointly managed by eastern cape parks and tourism agency and a community trust. income from tourism is currently minimal, as only a few mid-range accommodation options still exist, but a private company is developing some luxury accommodation. low tourism numbers affect benefit flow and employment possibilities, but reserve staff have been interacting with the community for several years and run various economic and social upliftment programmes. the third case study was phinda private game reserve and mnqobokasi community (24 c1 participants; 5 c2 participants) in kwazulu-natal. this reserve was established in 1991, is managed by &beyond and offers luxury tourism (&beyond 2018). some land is owned by local communities which rent it to &beyond. unlike the other two case studies, phinda has a dedicated organisation, africa foundation, focusing on community involvement and beneficiation. the relationships between phinda and the mnqobokazi community is well established, with employment opportunities and economic and social upliftment programmes in place. within the conservation and tourism realm, the model followed by phinda is widely perceived as successful. at each site, c2 participants had individual semi-structured interviews. for c1, at each case study site, participants self-divided into two groups. each group had a focus group interview and a drawing session. for the adapted nominal grouping technique, both groups came together. the research was conducted with a translator, in the mother tongues of participants, which at the three case study sites were sepedi, isimpondo and isizulu, respectively. access to the communities was facilitated by the protected area management and enabled via permission from community leadership. at each site, the same questions were asked to participants regarding the following themes: the community’s knowledge and experience of the protected area. the relationships between the community and the protected area. positive and negative changes that the protected area had brought to the community’s way of life. elements that could increase positive attitudes of the community towards the protected area. the community’s responsibilities towards the protected area. benefits and losses incurred by the community because of living nearby a protected area. participants’ ideas of an ideal future in the context of being a community bordering a protected area. for each case study, the data were transcribed and then coded inductively, creating a coding frame for each of the above-mentioned themes. after this, cross-case analysis took place to compare the findings across the three case studies, for each of these themes. using this analysis, meta-themes were identified that captured the most pertinent influences on pro-conservation attitudes and behaviour across the three case studies. it was these meta-themes that helped to construct the framework. as a result of the extent of the analysis (three case studies comprising 1217 coded quotes in total), followed by a cross-case analysis, the presentation and discussion of primary findings is beyond the scope of this article but can be found in queiros (2020). existing schemata in investigating existing schemata, two types were relevant to the people-park relationships – those representing the community and the conservation area and those focusing on the community only. certain components from these existing schemata were then incorporated into the final framework. schemata that include the community and conservation area (c & c) the ‘theory of resource use’ by firey (1960) was arguably the first model on protected area–people relationships and indicates the overlaps between the ecological, economic and social dimensions. following on from that several authors included these components within more detailed models, such as bennett (2016) and ross and wall (1999). abel and blaikie (1986) later developed a very detailed ‘management model for national parks’. it focused on elephants and ivory and different habitats. a few people elements were also included, such as employment opportunities and roads. in 2006, mccleave et al. published the ‘model of the new zealand people-park relationships’ and what affects the relationships. the park and its neighbours are the core of the model, surrounded by three dimensions: lifestyle, recreation and place attachment; tourism; and interactions with the park management agency. this model applies more to developed countries and to parks with a well-established tourism product. it is therefore less applicable to impoverished communities, resource access, meeting basic needs and so forth. of interest to an integrated model, however, is that the ‘stage of tourism development’ is included as a component that plays a part in relationships (and is hence included in the final framework). another approach was that of zube and busch (1990) who proposed four models of ‘park-local population relationships’. each model depicts a differing management approach that can encourage positive people-park relationships. the following six schemas directly focused on both the community and the conservation area, with each one contributing to the final framework. three of the models use a triangular structure. nyaupane and poudel (2011) created the model ‘linkages among biodiversity conservation, livelihood improvement and tourism development’, which illustrates the two-way relationships between these three aspects. their research used appreciative inquiry in three communities surrounding nepal’s chitwan national park. uni-directional arrows between the three aspects highlight different linkages, such as investments due to tourism development result in improved livelihoods. ross and wall’s (1999) ‘ecotourism paradigm’ has a similar structure in the sense that it considers the balance and synergistic relationships between local communities, biodiversity and tourism. however, they also add ‘management’ to the centre of the model because it plays a pivotal role in maintaining balance via effective management. for example, a one-way arrow links management to the local community, with the mention of ‘outreach programmes’ and ‘enforcement of use zones’. later, tsaur, lin and lin (2006) adapted ross and wall’s (1999) framework. they represented ‘tourism’, ‘local community’ and ‘the resource’ as the three corners of a triangle, with bi-directional arrows connecting the corners and representing economic, social and environmental relationships. the linked incentives model of direct linkage as a conservation strategy is the work of salafsky and wollenberg (2000). they had previously proposed three conservation strategies, namely no linkage, indirect linkage and direct linkage. it is the latter, as the ideal scenario, that became this model. using a series of blocks with connecting arrows, it represents different connections between livelihood activities and conservation (labelled as ‘biodiversity’). in this model, to protect the environment, alternative livelihood options that support biodiversity are provided to communities inside the conservation area (who might be an internal threat) and to communities outside the park (as a possible external threat). as a result of the linkages of increased benefits received, capacity building and locals recognising the value of biodiversity conservation, the notion is that local people will act to conserve. mutanga et al. (2015a) designed a framework for assessing the relationships between a protected area and community. they also used an arrangement of blocks (but in columns) with ‘protected area (staff)’ and ‘community’ as the two outer columns linked by the middle column of ‘human–wildlife interface’. their assertion is that conservation is enhanced if these relationships are positive. mutanga et al.’s study is based on a literature review and demonstrates that the attitudes of both parties’ shape relationships. four major factors play a role, namely the history of the protected area in the sense of forced relocation and a fences or fines approach; benefits and costs because of living near to a protected area; socio-demographic factors, for example, level of education and household size and income; and community involvement in conservation initiatives via integrated conservation and development projects. based on research in myanmar and nepal, the ‘framework for the protected area–people relationships’ by allendorf (2010) introduces the idea of layers within a framework. allendorf selected universal terms that can be applied and compared across different cases. the inner layer represents people’s physical relationships with the protected area (how locals interact with and use the area and the impact of the protected area on them in terms of costs). this layer also represents their attitudes towards the protected area (including their perceptions of the area) and concerns whether they like or dislike the protected area. the middle layer relates to people’s perception of other entities, for example, management, non-governmental organisations (ngos) and government (which can perform a direct or indirect role). finally, the outer layer concerns the broader context of the social, political, cultural, historical and economic realms and their linkages to the people-park relationships. lastly, bennett’s (2016) article investigates research regarding perceptions and the contribution thereof to improving adaptive management and evidence-based conservation. he maintains that perceptions are often dismissed in conservation science, but are research-worthy because they provide valuable insights into how to ensure the support of the community and thereby long-term conservation. bennett’s (2016) schema, in the form of a flow chart, therefore depicts the importance of knowing the perceptions regarding different facets (social, ecological, governance and management), which will then generate or undermine support for conservation. these perceptions are influenced by the social context and the individual context. schemata focusing on the community (c) four schemas with a community focus contained valuable elements for inclusion into an integrated framework. these are briefly discussed next. the ‘sustainable livelihoods framework’ was developed by scoones (1998) and is a multifaceted tool for analysing sustainable rural livelihoods. it includes five key indicators, namely context, livelihood resources, institutional processes, livelihood strategies and sustainable outcomes. in different contexts, several livelihood resources are available that combine to form a particular livelihood strategy, which in turn influences sustainable livelihood outcomes. this is affected by different processes and organisational structures. the different options aligned with each of the five key indicators are located in the centre of the framework (krantz 2001; scoones 1998). garrod et al. (2001) in garrod (2003) present the ‘revised model of local participation in planning and managing ecotourism’. it has a different focus – outlining eight consecutive steps that can be followed to foster local participation and good planning and management in ecotourism projects. originally developed for marine ecotourism in the european union’s atlantic region, this model has wider applicability to various ecotourism projects in different locations (garrod 2003). garrod argues that participation should be well integrated into these planning steps and not just be an ‘add-on’. quite different in structure, the stool model was developed by brook et al. (2009) for a project in northern canada. it depicts four critical supports (the legs of the stool) that are necessary in order to foster community-based wildlife health monitoring and research. these supports are collaboration, information and analysis, funding and education. the seat of the stool, being a local champion, is a key contribution of this model – coordinating, ‘holding’ the supports together and keeping participants engaged and informed. finally, based on a literature review of community-based tourism across rich and poor countries, giampiccoli, jugmohan and mtapuri (2015) generated the ‘community-based tourism e model’ (cbt e model). this model contains eight es, which represent the fundamental pillars against which a community-based tourism project can be evaluated. they are endogenous (reliance on natural resources); environment (care thereof as well as physical infrastructure); education (to advance both skills and education); empowerment (economic, psychological, social and political); equity (in terms of distributing benefits and resources); evolving (continuous improvement and adaptation); enduring (long-term sustainability); and entrepreneurship (the support thereof) (giampiccoli et al. 2015). the research gap presented in the introduction revealed that a comprehensive integrated framework that captures the components that could influence people-park relationships does not exist. this section has examined some of the existing schemata. these schemata hold value and components thereof are built into the framework resulting from this research. results: the people parks win-win framework by integrating existing schemata and existing literature with primary research findings, a newly synthesised framework was constructed. this section introduces the ppww – a comprehensive integrated framework representing the components that can influence people-park relationships (figure 2). through the threefold approach, comprehensiveness and generalisability was increased and a tool was created that can be used by relevant stakeholders. following the framework, its various layers, components and linkages are discussed via several tables. its contribution to the field of nature conservation is also discussed. figure 2: people parks win-win framework. discussion the ppww framework provides a detailed overview of the different layers and the components within each layer that can influence people-park relationships. four layers are set out, namely ‘external context’, ‘stakeholders’, ‘community beneficiation’ and ‘outputs’ and each contain their own components. to demonstrate validity and reliability, four tables accompany the discussion to review the origin of each component and layer. the ppww framework can be used by stakeholders involved in the relationships between people and parks, as a practical tool to facilitate win-win situations for both communities and biodiversity conservation. by representing multiple influences on this relationships between people and parks, the ppww can assist in shaping protected area management strategies for varying people-park arrangements, ranging from benefit-sharing only to true co-management. the ppww components neither operate in isolation nor as a complete package. it depends on the context of each protected area and its adjacent or resident community. stakeholders (often from trans-disciplines) should adapt it to their own context, taking cognisance that all these components can impact the relationships between people and parks. stakeholders also need to be aware that these relationships are complex and no single framework will entirely capture all impacting components. the ppww is a significant departure from other schemata for the following reasons: it is specific to the components that emerged in this research via primary and secondary research. its arrangement and the detailed focus on beneficiation (and principles to follow in benefit distribution) is novel. the ppww incorporates multiple stakeholders and indicates that all can be involved in beneficiation with these relationships affecting the achievement of a win-win scenario. as the characteristics of the stakeholders have considerable influence, they are included. it concludes with an ‘outputs’ layer, which suggests that preceding layers and components can culminate in the provision of benefits by stakeholders and represents the results this can have on community members – potentially leading to pro-conservation attitudes and behaviours and ultimately to a successful relationships between people and park. moreover, what takes place in this final layer can feed back into preceding layers, creating a positive cycle. finally, the centrality of relationships in the framework differentiates it from other schemata. while relationships are inherent in some of the linkages represented in the schemata that focus on both the community and the conservation area, only mutanga et al. (2015a) include the word ‘relationships’ within their framework, while ross and wall (1999) used it in the title of their framework. furthermore, the current framework is different in showing that all layers and components influence relationships. the discussion continues throughout table 1, table 2, table 3 and table 4. each layer is numbered and main headings are in bold. grey and white sections differentiate components from each other and linking arrows are indicated in italics. table 1: layer 1 and components used to build the people parks win-win framework. table 2: layer 2 and components used to build the people parks win-win framework. table 2 (continues...): layer 2 and components used to build the people parks win-win framework. table 3: layer 3 and components used to build the people parks win-win framework. table 3 (continues…): layer 3 and components used to build the people parks win-win framework. table 3 (continues…): layer 3 and components used to build the people parks win-win framework. table 4: layer 4 and components used to build the people parks win-win framework. the key for table 1, table 2, table 3 and table 4 and their preamble is as follows: c = existing schemata that focus on the community only. c & c = existing schemata that include community and conservation area. pr = primary research. lr = literature review. table 1 outlines the origin of each component within the ‘external context’ layer. as the outer layer, the ‘external context’ influences all other layers and components within the framework. the idea of layers of influence is borrowed from allendorf’s (2010) ‘framework for the protected-area–people relationships’ (c & c). people parks win-win framework uses its ‘context’ layer as the outer layer, renaming it as ‘external context’. however, while allendorf (2010) includes economic, historical, political, social and cultural within this outer layer, ppww only uses the first three components. it integrates social and cultural within the next layer for ‘stakeholders’. this is because the framework contains a dedicated layer for ‘stakeholders’ and the ‘social’ component is therefore better positioned alongside ‘community’ as a stakeholder. table 2 considers the components within the ‘stakeholders’ layer of ppww. allendorf (2010) (c & c) has a similar layer but labels it as ‘entities’. people parks win-win framework includes ‘community’ and ‘tourism management’ as stakeholders, thus adding to what allendorf included, namely ‘park management’, ‘ngos’ and ‘government’. table 2 also contains a sub-layer, entitled ‘stakeholder characteristics’ (highlighted in brown) – included because the pr reveals that the specific characteristics/circumstances of each stakeholder can influence this relationships. the characteristics of the community influence their perception of benefits (vertical green text in framework), while the characteristics of the other stakeholders will influence the generation of benefits (vertical peach text in framework). table 3 contains the origin, context and discussion around the components in layer 3, namely ‘community beneficiation’. the lr revealed various schools of thought on the linkages between benefits and losses, pro-conservation attitudes and behaviour. several authors establish a link between benefits and attitudes (liu et al. 2014; reimer & walter 2013; scanlon & kull 2009; strickland-munro & moore 2014) while others include lack of benefits or costs as influencing attitudes (black 2015; kideghesho, røskaft & kaltenborn 2007; mehta & heinen 2001). some researchers extend receipt of benefits to changing attitudes and behaviour (biodiversity conservation network 1999; gadd 2005; hulme & murphree 1999; kideghesho et al. 2007; mbaiwa 2005; ogunbode 2013; shibia 2010; tran & walter 2014) and some contend that it is also a lack of benefits or losses that influence attitudes and behaviour (burgoyne & mearns 2017; odindi & ayirebi 2010; thondhlana, cundill & kepe 2016). considered cumulatively, there is strong support in literature for the notion that benefits and losses influence attitudes, which in turn influence behaviour. in this pr, benefits and losses emerged as key in influencing attitudes and some links to behaviour were also found. therefore, based on the importance of benefits, as emerging in the lr and pr, ppww includes a layer dedicated to ‘community beneficiation’. some c & c schemata mention benefits (mutanga et al. [2015a] include ‘benefits and costs’; ross and wall [1999] refer to ‘outreach programmes’; salafsky and wollenberg [2000] mention ‘benefits’); and nyaupane and poudel (2011) list a few examples of benefits (such as jobs, education and infrastructure). however, ppww’s focus on the main benefits is a departure from existing schema. losses or costs incurred by the community are excluded because ppww focuses on requirements for a win-win relationships. it is important to observe that it is the presence of beneficiation and beneficiation principles that lead to the outputs (discussed in table 4). the stakeholder characteristics of the community will influence their perception of benefits (vertical green text in framework) and the characteristics of park management, tourism management, external stakeholders and government will influence the generation of benefits (vertical peach text in framework). the final section of the discussion (in table 4) centres on the ‘outputs’ layer, which suggests the outcomes that could be expected should enough of the components discussed in table 1, table 2 and table 3 be in place. finally, for practitioners who would prefer a simplified framework on which they can superimpose their own characteristics, benefits, influences and beneficiation principles, figure 3 provides the ‘simplified people parks win-win framework’. figure 3: simplified people parks win-win framework. conclusion the biodiversity and poverty crises are realities and impact each other. in the african context, protected area management confronts very real constraints in terms of budgets and a limited number of staff, who need to accomplish a multitude of tasks. communities bordering these parks are often impoverished and may not share the sentiment that these parks should be protected. the conservation of this crucial biodiversity thus depends on good relationships with surrounding communities and finding win-win solutions. yet, there are gaps regarding what influences local communities to have, or not to have, pro-conservation attitudes and behaviour and regarding people-park relationships. furthermore, an extensive framework that captures the multiple influences on this relationships did not exist. the people parks win-win framework was constructed from literature, existing schema and primary research to illustrate the components that can influence the people-park relationships. the framework provides a more complex, multidimensional type of framework. it deviates from others in terms of its structure and arrangement; detailed focus on beneficiation; incorporation of more stakeholders, as well as their characteristics; strong focus on community well-being being equally important to biodiversity conservation; centrality of the people-park relationship; and inclusion of an output layer, which demonstrates how the preceding layers could culminate in a win-win scenario, and also how pro-conservation attitudes and behaviour fit into this. while the primary research was carried out in the south african context, the literature and schemata that contributed to the framework come from various authors worldwide. the resultant framework has global applicability to other settings where people live side by side with parks and can be adapted to suit different contexts. it can be used by stakeholders to: understand the multiple forces at play regarding the people-park relationship; consider the range of benefits and customise these to their own context and particular constraints faced; and consider the other internal influences and beneficiation principles which can maximise a good relationships and contribute to win-wins. this framework could be a practical tool for protected area management, communities and other stakeholders when developing strategies and plans for the future. it may also be of interest to academics – testing, customising or adding to it. rode et al. (2016:46) suggest that ‘“win-win solutions” that can combine conservation and livelihoods benefits may not always be found, but that it is worth looking for them’. win-wins are indeed a complex process involving negotiations between different stakeholders with different agendas and clarity in planning as well as implementation. with commitment from government, conservation agencies, tourism bodies and other stakeholders, as well as informed realistic expectations from local communities and recognition of their role and responsibilities, there is potential for tangible and intangible benefit provision within a healthy context. this could foster positive attitudes that could lead to pro-conservation behaviour and the instilling of robust reciprocate relationships between the park and the people living nearby – where communities benefit from the wildlife on their doorstep and are part of its conservation. the alternative of not investing in this crucial relationships and not ensuring a healthy sustainable flow of benefits (i.e. not looking for ‘win-win solutions’) could result in significant losses on both sides, with the final score being lose–lose. acknowledgements the author would like to acknowledge the communities and protected area staff who participated in the primary research. she would also wish to thank the supervisors of her phd in environmental management, prof. kevin mearns and prof. ciné van zyl for their invaluable guidance. this article stems from the thesis of the author, titled ‘towards pro-conservation attitudes and behaviour by local communities bordering protected areas in south africa’ submitted to the university of south africa in fulfilment of doctoral degree, available here: http://hdl.handle.net/10500/26663 competing interests the author declares that she has no financial or personal relationships that may have inappropriately influenced her in writing this article. author’s contributions d.r.q. is the sole author of this article. ethical considerations ethical clearance was issued by the university of south africa (unisa) (reference number: 2015/caes/016) and permission obtained from the participating reserves. informed consent was obtained from each chief or community leader, protected area manager as well as all participants. funding information funding for this research was provided by the university of south africa through its academic qualification improvement programme. data availability the primary data analysed to construct the framework is available in queiros (2020), but the original transcripts are protected by the ethics rules of the university. disclaimer the views and opinions expressed in this article are those of the author and do not necessarily reflect the official policy or position of any affiliated agency of the author. references &beyond, 2018, luxury experiential travel in africa, asia and south america, viewed 02 august 2018, from https://www.andbeyond.com/. abel, n. & blaikie, p., 1986, ‘elephants, people, parks and development: the case of the luangwa valley, zambia’, environmental management 10(6), 735–751. https://doi.org/10.1007/bf01867727 abrams, r.w., anwana, e.d., ormsby, a., dovie, d., ajagbe, a. & abrams, a., 2009, ‘integrating top-down with bottom-up conservation policy in africa’, conservation biology 23(4), 799–804. https://doi.org/10.1111/j.1523-1739.2009.01285.x adams, w.m., 2013, ‘conservation in the anthropocene: biodiversity, poverty and sustainability’, in m. roe, d. elliott, j. sandbrook & c. walpole (eds.), biodiversity conservation and poverty alleviation: exploring the evidence for a link, pp. 304–315, john wiley & sons, ltd., chichester. african development report, 2015, africa’s youth in the labour market, african development bank group, abidjan. allen, l., holland, k.k., holland, h., tome’, s., nabaala, m., seno, s. et al., 2019, ‘expanding staff voice in protected area management effectiveness assessments within kenya’s maasai mara national reserve’, environmental management 63(1), 46–59. https://doi.org/10.1007/s00267-018-1122-6 allendorf, t., swe, k.k., oo, t., htut, y., aung, m., aung, m. et al., 2006, ‘community attitudes toward three protected areas in upper myanmar (burma)’, environmental conservation 33(4), 344–352. https://doi.org/10.1017/s0376892906003389 allendorf, t.d., 2010, ‘a framework for the park-people relationship: insights from protected areas in nepal and myanmar’, international journal of sustainable development and world ecology 17(5), 417–422. https://doi.org/10.1080/13504501003788180 anthony, b.p., 2021, ‘paying for the past: the importance of fulfilling promises as a key component to resolving human–wildlife conflict’, sustainability 13(7407), 1–20. https://doi.org/10.3390/su13137407 baker, a., 2015, ‘let’s prepare for africa’s population surge now – or face the consequences’, time magazine, 22 june, p. 44. bann, e., 2001, ‘effects of media representations of a cultural ideal of feminine beauty on self body image in college-aged women: an interactive qualitative analysis’, phd thesis, the university of texas at austin, austin, tx. bennett, n.j., 2016, ‘using perceptions as evidence to improve conservation and environmental management’, conservation biology 30(3), 1–11. https://doi.org/10.1111/cobi.12681 berkes, f., 2004, ‘rethinking community-based conservation’, conservation biology 18(3), 621–630. https://doi.org/10.1111/j.1523-1739.2004.00077.x bezerra, j., 2018, ‘land claims in south africa: it’s about the meaning of the land, not just money’, the conversation: academic rigour, journalistic flair, 10 august 2018, n.p. biodiversity conservation network, 1999, evaluating linkages between business, the environment, and local communities: final stories from the field, biodiversity support program, washington, dc. black, r., 2015, social and economic impacts of tourist lodges on local communities: case studies from rwanda and botswana, a report for the african wildlife foundation, institute for land, water and society, albury. brook, r.k., kutz, s.j., veitch, a.m., popko, r.a., elkin, b.t. & guthrie, g., 2009, ‘fostering community-based wildlife health monitoring and research in the canadian north’, ecohealth 6(2), 266–278. https://doi.org/10.1007/s10393-009-0256-7 burgoyne, c. & mearns, k., 2017, ‘managing stakeholder relations, natural resources and tourism: a case study from ololosokwan, tanzania’, tourism and hospitality research 17(1), 68–78. https://doi.org/10.1177/1467358416639088 campbell, l.m., haalboom, b.j. & trow, j., 2007, ‘sustainability of community-based conservation: sea turtle egg harvesting in ostional (costa rica) ten years later’, environmental conservation 34(2), 122–131. ceballos, g., ehrlich, p., barnosky, a.d., garcia, a., pringle, r. & palmer, t.m., 2015, ‘accelerated modern human-induced species losses: entering the sixth mass extinction’, science advances 1(5), 1–6. https://doi.org/10.1126/sciadv.1400253 cetas, e.r. & yasué, m., 2017, ‘a systematic review of motivational values and conservation success in and around protected areas’, conservation biology 31(1), 203–212. https://doi.org/10.1111/cobi.12770 cobbinah, p.b., black, r. & thwaites, r., 2015, ‘biodiversity conservation and livelihoods in rural ghana: impacts and coping strategies’, environmental development 15, 79–93. https://doi.org/10.1016/j.envdev.2015.04.006 collins, s., 2016, ‘no easy road: an evaluation of the makuleke community conservation model’, in african wildlife foundation, nairobi’, african conservancies: towards best practices 1, 74–81. coria, j. & calfucura, e., 2012, ‘ecotourism and the development of indigenous communities: the good, the bad, and the ugly’, ecological economics 73, 47–55. https://doi.org/10.1016/j.ecolecon.2011.10.024 creswell, j.w., 2014, research design: qualitative, quantitative and mixed methods approaches, 4th edn., sage, ca. davies, t.e., fazey, i.r.a., cresswell, w. & pettorelli, n., 2014, ‘missing the trees for the wood: why we are failing to see success in pro-poor conservation’, animal conservation 17(4), 303–312. https://doi.org/10.1111/acv.12094 de los angeles somarriba-chang, m. & gunnarsdotter, y., 2012, ‘local community participation in ecotourism and conservation issues in two nature reserves in nicaragua’, journal of sustainable tourism 20(8), 1–19. https://doi.org/10.1080/09669582.2012.681786 de pourcq, k., thomas, e., arts, b., vranckx, a., léon-sicard, t. & van damme, p., 2015, ‘conflict in protected areas: who says co-management does not work?’, plos one 10(12), 1–15. https://doi.org/10.1371/journal.pone.0144943 diedrich, a., 2007, ‘the impacts of tourism on coral reef conservation awareness and support in coastal communities in belize’, coral reefs 26(4), 985–996. https://doi.org/10.1007/s00338-007-0224-z dinokeng, 2017, the hub of dinokeng game reserve, viewed 10 april 2017, from http://www.gauteng.net/dinokeng/hubs/dinokeng-game-reserve. ferse, s.c.a., máñez costa, m., máñez, k.s., adhuri, d.s. & glaser, m., 2010, ‘allies, not aliens: increasing the role of local communities in marine protected area implementation’, environmental conservation 37(1), 23–34. https://doi.org/10.1017/s0376892910000172 firey, w., 1960, man, mind and land, the free press, glencoe, il. gadd, m.e., 2005, ‘conservation outside of parks: attitudes of local people in laikipia, kenya’, environmental conservation 32(1), 50–63. https://doi.org/10.1017/s0376892905001918 garrod, b., 2003, ‘local participation in the planning and management of ecotourism: a revised model approach’, journal of ecotourism 2(1), 33–53. https://doi.org/10.1080/14724040308668132 ghoddousi, s., pintassilgo, p., mendes, j., ghoddousi, a. & sequeira, b., 2018, ‘tourism and nature conservation: a case study in golestan national park, iran’, tourism management perspectives 26, 20–27. https://doi.org/10.1016/j.tmp.2017.12.006 giampiccoli, a., jugmohan, s. & mtapuri, o., 2015, ‘community-based tourism in rich and poor countries: towards a framework for comparison’, african journal for physical health education, recreation and dance 21(4), 1200–1216. gurung, d.b. & seeland, k., 2011, ‘ecotourism benefits and livelihood improvement for sustainable development in the nature conservation areas of bhutan’, sustainable development 19(5), 348–358. https://doi.org/10.1002/sd.443 hackel, j.d., 1999, ‘community conservation and the future of africa’s wildlife’, conservation biology 13(4), 726–734. https://doi.org/10.1046/j.1523-1739.1999.98210.x harihar, a., veríssimo, d. & macmillan, d.c., 2015, ‘beyond compensation: integrating local communities’ livelihood choices in large carnivore conservation’, global environmental change 33, 122–130. https://doi.org/10.1016/j.gloenvcha.2015.05.004 hulme, d. & murphree, m., 1999, ‘communities, wildlife and the ‘new conservation’ in africa’, journal of international development 11(2), 277–285. https://doi.org/10.1002/(sici)1099-1328(199903/04)11:2<277::aid-jid582>3.0.co;2-t imran, s., alam, k. & beaumont, n., 2014, ‘environmental orientations and environmental behaviour: perceptions of protected area tourism stakeholders’, tourism management 40, 290–299. https://doi.org/10.1016/j.tourman.2013.07.003 infield, m. & namara, a., 2001, ‘community attitudes, and behaviour towards conservation: an assessment of a community conservation programme around lake mburo, national park, uganda’, oryx 35(1), 48–60. https://doi.org/10.1046/j.1365-3008.2001.00151.x kideghesho, j.r., røskaft, e. & kaltenborn, b.p., 2007, ‘factors influencing conservation attitudes of local people in western serengeti, tanzania’, biodiversity and conservation 16(7), 2213–2230. https://doi.org/10.1007/s10531-006-9132-8 kiss, a., 2004, ‘is community-based ecotourism a good use of biodiversity conservation funds?’, trends in ecology and evolution 19(5), 232–237. kohler, f. & brondizio, e.s., 2017, ‘considering the needs of indigenous and local populations in conservation programs’, conservation biology 31(2), 245–251. https://doi.org/10.1111/cobi.12843 krantz, l., 2001, the sustainable livelihood approach to poverty reduction: an introduction, swedish international development cooperation agency, stockholm. krűger, o., 2005, ‘the role of ecotourism in conservation: panacea or pandora’s box?’, biodiversity and conservation 14, 579–600. https://doi.org/10.1007/s10531-004-3917-4 larson, l.r., conway, a.l., krafte, k.e., hernandez, s.m. & carroll, j.p., 2016, ‘community-based conservation as a potential source of conflict around a protected area in sierra leone’, environmental conservation 43(3), 242–252. lee, t.h., 2013, ‘influence analysis of community resident support for sustainable tourism development’, tourism management 34, 37–46. https://doi.org/10.1016/j.tourman.2012.03.007 liu, j., qu, h., huang, d., chen, g., yue, x., zhao, x. et al., 2014, ‘the role of social capital in encouraging residents’ pro-environmental behaviors in community-based ecotourism’, tourism management 41, 190–201. https://doi.org/10.1016/j.tourman.2013.08.016 lucas, k., brooks, m., darnton, a. & jones, j.e., 2008, ‘promoting pro-environmental behaviour: existing evidence and policy implications’, environmental science and policy 11(5), 456–466. https://doi.org/10.1016/j.envsci.2008.03.001 lune, h. & berg, b.l., 2017, qualitative research methods for the social sciences, 9th edn., pearson education limited, harlow. matarrita-cascante, d., brennan, m.a. & luloff, a.e., 2010, ‘community agency and sustainable tourism development: the case of la fortuna, costa rica’, journal of sustainable tourism 18(6),735–756. https://doi.org/10.1080/09669581003653526 maxwell, j.a., 2013, qualitative research design: an interactive approach, 3rd edn., california, sage, ca. mbaiwa, j., 2005, ‘wildlife resource utilization at moremi game reserve and khwai community area in the okavango delta, botswana’, journal of environmental management 77(2), 144–156. https://doi.org/10.1016/j.jenvman.2005.03.007 mbaiwa, j.e. & stronza, a.l., 2010, ‘the effects of tourism development on rural livelihoods in the okavango delta, botswana’, journal of sustainable tourism 18(5), 635–656. https://doi.org/10.1080/09669581003653500 mccleave, j., espiner, s. & booth, k., 2006, ‘the new zealand people-park relationship: an exploratory model’, society and natural resources 19(6), 547–561. mcshane, t.o., hirsch, p.d., trung, t.c., songorwa, a.n., kinzig, a., monteferri, b. et al., 2011, ‘hard choices: making trade-offs between biodiversity conservation and human well-being’, biological conservation 144(3), 966–972. mehta, j.n. & heinen, j.t., 2001, ‘does community-based conservation shape favorable attitudes among locals? an empirical study from nepal’, environmental management 28(2), 165–177. https://doi.org/10.1007/s002670010215 mutanga, c.n., muboko, n. & gandiwa, e., 2017, ‘protected area staff and local community viewpoints: a qualitative assessment of conservation relationships in zimbabwe’, plos one 12(9), 1–21. https://doi.org/10.1371/journal.pone.0184779 mutanga, c.n., vengesayi, s., muboko, n. & gandiwa, e., 2015a, ‘towards harmonious conservation relationships: a framework for understanding protected area staff-local community relationships in developing countries’, journal for nature conservation 25, 8–16. https://doi.org/10.1016/j.jnc.2015.02.006 mutanga, c.n., vengesayi, s., gandiwa, e. & muboko, n., 2015b, ‘community perceptions of wildlife conservation and tourism: a case study of communities adjacent to four protected areas in zimbabwe’, tropical conservation science 8(2), 564–582. niedziałkowski, k., komar, e., pietrzyk-kaszyńska, a., olszańska, a. & grodzińska-jurczak, m., 2018, ‘discourses on public participation in protected areas governance: application of q methodology in poland’, ecological economics 145, 401–409. https://doi.org/10.1016/j.ecolecon.2017.11.018 nyaupane, g.p. & poudel, s., 2011, ‘linkages among biodiversity, livelihood, and tourism’, annals of tourism research 38(4), 1344–1366. https://doi.org/10.1016/j.annals.2011.03.006 odindi, j.o. & ayirebi, g.k., 2010, ‘communities and conservation: in search for a win-win situation in the great fish river reserve’, journal of sustainable development in africa 12(1), 13–26. ogunbode, c.a., 2013, ‘the nep scale: measuring ecological attitudes/worldviews in an african context’, environment, development and sustainability 15(6), 1477–1494. https://doi.org/10.1007/s10668-013-9446-0 pechacek, p., li, g., li, j., wang, w., wu, x. & xu, j., 2013, ‘compensation payments for downsides generated by protected areas’, ambio 42(1), 90–99. https://doi.org/10.1007/s13280-012-0330-1 pfueller, s.l., lee, d. & laing, j., 2011, ‘tourism partnerships in protected areas: exploring contributions to sustainability’, environmental management 48(4), 734–749. https://doi.org/10.1007/s00267-011-9728-y pretty, j. & smith, d., 2004, ‘social capital in biodiversity conservation and management’, conservation biology 18(3), 631–638. https://doi.org/10.1111/j.1523-1739.2004.00126.x queiros, d.r., 2000, ‘implementing the fundamentals of ecotourism: the case of mkambati nature reserve, wild coast, south africa’, unpublished master’s dissertation, university of pretoria. queiros, d.r., 2020, ‘towards pro-conservation attitudes and behaviour by local communities bordering protected areas in south africa’, unpublished phd thesis, university of south africa, pretoria. rahman, m.m., mahmud, m.a.l. & shahidullah, m., 2017, ‘socioeconomics of biodiversity conservation in the protected areas: a case study in bangladesh’, international journal of sustainable development & world ecology 24(1), 65–72. https://doi.org/10.1080/13504509.2016.1169453 reimer, j.k.k. & walter, p., 2013, ‘how do you know it when you see it? community-based ecotourism in the cardamom mountains of southwestern cambodia’, tourism management 34, 122–132. https://doi.org/10.1016/j.tourman.2012.04.002 rode, j., wittmer, h., emerton, l. & schröter-schlaack, c., 2016, ‘ecosystem service opportunities’: a practice-oriented framework for identifying economic instruments to enhance biodiversity and human livelihoods’, journal for nature conservation 33, 35–47. https://doi.org/10.1016/j.jnc.2016.07.001 roe, d. & elliott, j., 2006, ‘pro-poor conservation: the elusive win-win for conservation and poverty reduction?’, policy matters 14, 53–63. ross, s. & wall, g., 1999, ‘ecotourism: towards congruence between theory and practice’, tourism management 20(1), 123–132. roulston, k., 2014, ‘analysing interviews’, in u. flick (ed.), the sage handbook of qualitative data analysis, pp. 297–312, sage, london. sachedina, h. & nelson, f., 2010, ‘protected areas and community incentives in savannah ecosystems: a case study of tanzania’s maasai steppe’, oryx 44(3), 390–398. https://doi.org/10.1017/s0030605310000499 salafsky, n. & wollenberg, e.v.a., 2000, ‘linking livelihoods and conservation: a conceptual framework and scale for assessing the integration of human needs and biodiversity’, world development 28(8), 1421–1438. https://doi.org/10.1016/s0305-750x(00)00031-0 saufi, a., o’brien, d. & wilkins, h., 2014, ‘inhibitors to host community participation in sustainable tourism development in developing countries’, journal of sustainable tourism 22(5), 801–820. https://doi.org/10.1080/09669582.2013.861468 scanlon, l.j. & kull, c.a., 2009, ‘untangling the links between wildlife benefits and community-based conservation at torra conservancy, namibia’, development southern africa 26(1), 75–93. https://doi.org/10.1080/03768350802640107 scoones, i., 1998, sustainable rural livelihoods: a framework for analysis, institute of development studies (ids), brighton. shackleton, c.m., willis, t.j., brown, k. & polunin, n.v.c., 2010, ‘reflecting on the next generation of models for community-based natural resources management’, environmental conservation 37(1), 1–4. https://doi.org/10.1017/s0376892910000366 shibia, m.g., 2010, ‘determinants of attitudes and perceptions on resource use and management of marsabit national reserve, kenya’, journal of human ecology 30(1), 55–62. https://doi.org/10.1080/09709274.2010.11906272 simpson, m.c., 2008, ‘community benefit tourism initiatives: a conceptual oxymoron?’, tourism management 29(1), 1–18. https://doi.org/10.1016/j.tourman.2007.06.005 snyman, s., 2012, ‘the role of tourism employment in poverty reduction and community perceptions of conservation and tourism in southern africa’, journal of sustainable tourism 20(3), 395–416. https://doi.org/10.1080/09669582.2012.657202 snyman, s., 2014, ‘assessment of the main factors impacting community members’ attitudes towards tourism and protected areas in six southern african countries’, koedoe 56(2), 1–12. https://doi.org/10.4102/koedoe.v56i2.1139 sommerville, m., jones, j.p.g., rahajaharison, m. & milner-gulland, e.j., 2010, ‘the role of fairness and benefit distribution in community-based payment for environmental services interventions: a case study from menabe, madagascar’, ecological economics 69(6), 1262–1271. https://doi.org/10.1016/j.ecolecon.2009.11.005 south africa, 2003, national environmental management: protected areas act (act no. 57 of 2003), government notice no. 181, government gazette 26025, february 2003. south africa, 2004, national environmental management: biodiversity act (act no. 10 of 2004), government notice no. 700, government gazette 26436, june 2004. south africa, 2016, national environmental management: protected areas act, 2003 (act no. 57 of 2003): norms and standards for the management of protected areas in south africa, government notice no. 382, government gazette 39878, march 2016. spenceley, a., 2008, ‘local impacts of community-based tourism in southern africa’, in a. spenceley (ed.), responsible tourism: critical issues for conservation and development, pp. 285–304, earthscan, london. spenceley, a., rylance, a., nanabhay, s. & van der watt, h., 2016, operational guidelines for community-based tourism in south africa, department of tourism, pretoria. stem, c.j., lassoie, j.p., lee, d.r., deshler, d.d. & schelhas, j.w., 2003, ‘community participation in ecotourism benefits: the link to conservation practices and perspectives’, society & natural resources 16(5), 387–413. https://doi.org/10.1080/08941920309177 stoll-kleemann, s., 2005, ‘voices for biodiversity management in the 21st century’, environment: science and policy for sustainable development 47(10), 24–36. https://doi.org/10.3200/envt.47.10.24-36 stoll-kleemann, s., de la vega-leinert, a.c. & schultz, l., 2010, ‘the role of community participation in the effectiveness of unesco biosphere reserve management: evidence and reflections from two parallel global surveys’, environmental conservation 37(3), 227–238. https://doi.org/10.1017/s037689291000038x stone, m.t. & nyaupane, g.p., 2018, ‘protected areas, wildlife-based community tourism and community livelihoods dynamics: spiraling up and down of community capitals’, journal of sustainable tourism 26(2), 307–324. https://doi.org/10.1080/09669582.2017.1349774 strickland-munro, j. & moore, s., 2014, ‘exploring the impacts of protected area tourism on local communities using a resilience approach’, koedoe 56(2), 1–10. https://doi.org/10.4102/koedoe.v56i2.1161 stronza, a. & gordillo, j., 2008, ‘community views of ecotourism’, annals of tourism research 35(2), 448–468. https://doi.org/10.1016/j.annals.2008.01.002 swemmer, l., mmethi, h. & twine, w., 2017, ‘tracing the cost/benefit pathway of protected areas: a case study of the kruger national park, south africa’, ecosystem services 28(part b), 62–172. https://doi.org/10.1016/j.ecoser.2017.09.002 tessema, m.e., ashenafi, z.t., lilieholm, r.j. & leader-williams, n.l., 2007, ‘community attitudes towards wildlife conservation in ethiopia’, in proceedings of the 2007 george wright society conference, assessing public attitudes and experiences, st paul, the george wright society, minnesota, april 16–20, 2007, pp. 287–292. thondhlana, g. & cundill, g., 2017, ‘local people and conservation officials’ perceptions on relationships and conflicts in south african protected areas’, international journal of biodiversity science, ecosystem services & management 13(1), 204–215. https://doi.org/10.1080/21513732.2017.1315742 thondhlana, g., cundill, g. & kepe, t., 2016, ‘co-management, land rights, and conflicts around south africa’s silaka nature reserve’, society and natural resources 29(4), 403–417. tolkach, d. & king, b., 2015, ‘strengthening community-based tourism in a new resource-based island nation: why and how?’, tourism management 48, 386–398. https://doi.org/10.1016/j.tourman.2014.12.013 torquebiau, e. & taylor, r.d., 2009, ‘natural resource management by rural citizens in developing countries: innovations still required’, biodiversity and conservation 18(10), 2537–2550. https://doi.org/10.1007/s10531-009-9706-3 tran, l. & walter, p., 2014, ‘ecotourism, gender and development in northern vietnam’, annals of tourism research 44(1), 116–130. https://doi.org/10.1016/j.annals.2013.09.005 tsaur, s.h., lin, y.c. & lin, j.h., 2006, ‘evaluating ecotourism sustainability from the integrated perspective of resource, community and tourism’, tourism management 27(4), 640–653. https://doi.org/10.1016/j.tourman.2005.02.006 united nations, 2021, global population growth and sustainable development (report by united nations department of economic and social affairs, population division), viewed 03 june 2022, from https://www.un.org/development/desa/pd/sites/www.un.org.development.desa.pd/files/undesa_pd_2022_global_population_growth.pdf. wali, a., alvira, d., tallman, p.s., ravikumar, a. & macedo, m.o., 2017, ‘a new approach to conservation: using community empowerment for sustainable well-being’, ecology and society 22(4), 1–14. https://doi.org/10.5751/es-09598-220406 walpole, m.j. & goodwin, h.j., 2001, ‘local attitudes towards conservation and tourism around komodo national park, indonesia’, enviromental conservation 28(2), 160–166. https://doi.org/10.1017/s0376892901000169 waylen, k.a., fischer, a., mcgowan, p.j.k., thirgood, s.j. & milner-gulland, e.j., 2010, ‘effect of local cultural context on the success of community-based conservation interventions’, conservation biology 24(4), 1119–1129. https://doi.org/10.1111/j.1523-1739.2010.01446.x world economic forum, 2019, how can south africa reduce working poverty?, viewed 09 november 2019, from https://www.weforum.org/agenda/2015/08/how-can-south-africa-reduce-working-poverty. zhang, l., luo, z., mallon, d., li, c. & jiang, z., 2017, ‘biodiversity conservation status in china’s growing protected areas’, biological conservation 210(part b), 89–100. https://doi.org/10.1016/j.biocon.2016.05.005 zube, e.h. & busch, m.l., 1990, ‘park-people relationships: an international review’, landscape and urban planning 19(2), 117–131. https://doi.org/10.1016/0169-2046(90)90030-6 abstract introduction materials and methods results discussion conclusion acknowledgements references appendix 1 appendix 2 about the author(s) lizanne basson department of veterinary tropical diseases, university of pretoria, south africa centre for emerging, zoonotic and parasitic diseases, national institute for communicable diseases, south africa ayesha hassim department of veterinary tropical diseases, university of pretoria, south africa at dekker state veterinarians office, kruger national park, south africa allison gilbert centre for emerging, zoonotic and parasitic diseases, national institute for communicable diseases, south africa wits research institute for malaria, university of the witwatersrand, south africa wolfgang beyer department of livestock infectiology and environmental hygiene, university of hohenheim, germany jennifer rossouw centre for emerging, zoonotic and parasitic diseases, national institute for communicable diseases, south africa henriette van heerden department of veterinary tropical diseases, university of pretoria, south africa citation basson, l., hassim, a., dekker, a., gilbert, a., beyer, w., rossouw, j. & van heerden, h., 2018, ‘blowflies as vectors of bacillus anthracis in the kruger national park’, koedoe 60(1), a1468. https://doi.org/10.4102/koedoe.v60i1.1468 short communication blowflies as vectors of bacillus anthracis in the kruger national park lizanne basson, ayesha hassim, at dekker, allison gilbert, wolfgang beyer, jennifer rossouw, henriette van heerden received: 29 mar. 2017; accepted: 09 apr. 2018; published: 26 june 2018 copyright: © 2018. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract anthrax, caused by bacillus anthracis, is endemic in the kruger national park (knp). the epidemiology of b. anthracis is dependent on various factors including vectors. the aims of this study were to examine non-biting blowflies for the presence of b. anthracis externally and internally after feeding on an anthrax-infected carcass and to determine the role of flies in disseminating b. anthracis onto the surrounding vegetation. during an anthrax outbreak in 2014 in the endemic pafuri region, blowflies associated with two 2–3-day-old anthrax-positive carcasses (kudu and impala) as well as surrounding vegetation were collected and investigated for the presence of b. anthracis spores. the non-biting blowflies (n = 57) caught included chrysomya albiceps, ch. marginalis and lucilia spp. bacillus anthracis spores were isolated from 65.5% and 25.0% of blowflies collected from the kudu and impala carcasses, respectively. chrysomya albiceps and ch. marginalis have the potential to disseminate b. anthracis to vegetation from infected carcasses and may play a role in the epidemiology of anthrax in the knp. no b. anthracis spores were initially isolated from leaves of the surrounding vegetation using selective media. however, 170 and 500 spores were subsequently isolated from abutilon angulatum and acacia sp. leaves, respectively, when using sheep blood agar. conservation implications: the results obtained in this study have no direct conservation implications and only assist in the understanding of the spread of the disease. introduction anthrax is a serious zoonotic disease affecting mainly herbivores. it is caused by the soil-borne, gram-positive, spore-forming organism bacillus anthracis. in unfavourable conditions, b. anthracis forms endospores that can remain dormant in the environment for long periods of time surviving outside the host. the spores are ingested by a susceptible host, where germination and multiplication occur in vivo. the host dies and the carcass is opened by scavengers or human, resulting in the vegetative cells being exposed to oxygen and sporulating; remaining in the soil until another host ingests the spores and the cycle is repeated (dragon & rennie 1995). anthrax occurs endemically in the northern kruger national park (knp) and the northern cape province (ghaap plateau area) in south africa (hugh-jones & de vos 2002). the disease cannot spread from one living animal to the next; therefore, the life cycle is dependent on various factors including rainfall, temperature, type of soil, animal densities and the presence of susceptible hosts (dragon & rennie 1995). a number of abiotic and biotic vectors such as insects (biting flies, blowflies, mosquitoes and ticks) (blackburn et al. 2010; fasanella et al. 2010; graham-smith 1913; turell & knudson 1987), scavengers (pienaar 1961, 1967), soil (hugh-jones & blackburn 2009) and water (hugh-jones & de vos 2002; pienaar 1961) have also been implicated in the dissemination of anthrax. the main insect vectors implicated with anthrax in the knp are the blowflies chrysomya albiceps and ch. marginalis (braack 1985) as these species are the first (and most abundant) insects to arrive shortly after death. these two species consume infected bodily fluids and are probably the principal vectors in the dissemination of b. anthracis (braack 1985; braack & retief 1986). chrysomya albiceps and ch. marginalis have been hypothesised to spread the bacteria over short distances despite the fact that blowflies have the ability to travel great distances (35 km – 65 km). they normally rest on nearby vegetation when engorged and deposit most of the contaminated defaecation and/or vomit droplets close to the carcass (braack & de vos 1990; de vos 1990) of vegetation (reviewed by de vos 1994), which pose a risk of infection to susceptible herbivores. in knp, outbreaks were usually associated with the driest periods of the year (winter and early spring) or during climatic dry cycles (pienaar 1967). however, since the 2010 outbreak onwards, outbreaks mainly occurred in wet periods (summer months; e.h. dekker [skukuza state veterinary office] pers. comm., 2014). this is of importance as insect activity and abundance on a carcass is influenced by climate. in the rainy season, the insect abundance (especially the blowflies) will increase, which was speculated by braack (1985) to increase b. anthracis dissemination by blowflies to vegetation during an outbreak. scavengers have also been speculated to disseminate spores at drinking holes after opening and feeding on anthrax-infected carcasses (pienaar 1961). research conducted by turner et al. (2014, 2016) indicated that grazing seems to play a significant role in the dissemination of b. anthracis in etosha national park. similar investigations must be performed in knp to determine the factors influencing the epidemiology of anthrax as well as the amount of spores found on flies and the bacterial load defaecated or regurgitated by flies on vegetation. in this study, we investigated the possible role of blowflies in carrying and/or spreading b. anthracis during an anthrax outbreak in the knp. this aim was investigated by quantifying the presence of b. anthracis on the exterior and interior of blowflies after feeding on an anthrax-contaminated carcass. furthermore, the role of blowflies in the dissemination of b. anthracis onto the surrounding vegetation during the anthrax outbreak in the knp was also investigated. materials and methods blowfly collection and identification in march 2014 – april 2014, blowflies were collected from two confirmed anthrax-positive carcasses (a kudu and an impala) during an anthrax outbreak in the endemic pafuri region of the knp. the adult male kudu carcass was approximately 2–3 days old, lying in the open with clear indications that scavengers (vultures and/or hyenas) had started to feed on it. the adult male impala carcass infected with anthrax was about three days old, lying under a low thorn tree (acacia sp.). the carcass was mostly intact, except for a large opening in the abdomen region. a net trap was used to collect the blowflies on the carcass. trapped flies were collected and placed in liquid nitrogen for 30 seconds and transported at 4 °c to the national institute for communicable diseases (nicd) for further bacteriological analysis. all blowflies were kept, while other flies were discarded. each blowfly was identified and sexed using the identification method described by zumpt (1965). blowflies that were not phenotypically or morphologically distinguishable were identified by sequencing of the ribosomal dna internally transcribed spacer 2 (its 2) region that differentiates calliphoridae as described by koekemoer et al. (2002). isolation of bacillus anthracis from blowflies each blowfly was tested for the presence of b. anthracis vegetative cells and spores on the interior and exterior surfaces. to detect the presence of b. anthracis on exterior surfaces, each blowfly was washed in 1 ml sterile saline to remove any bacteria. the presence of b. anthracis vegetative cells was determined by plating 10-fold dilutions (100–108) from the external wash onto polymyxin–edta–thallium acetate (pet; modified polymyxin-lysozyme-edta-thallium acetate [plet] where lysozyme was omitted) agar plates followed by incubation at 37 °c for 48 hours. the presence of b. anthracis spores was determined by plating 100 µl heat-treated (62.5 °c for 15 min) undiluted external wash on pet agar, followed by incubation at 37 °c for 48 h. after the exterior wash step, each blowfly was disinfected with 1 ml 0.1% (v/v) peracetic acid for 1 h to inactivate any residual bacteria left over on the exterior of the blowfly after the wash step. the 0.1% peracetic acid (v/v) was neutralised by 1 ml 0.8% sodium bicarbonate – sodium chloride (v/v) solution for 1 h. one-hundred microliters of the neutralised external wash was inoculated onto 5% sheep blood agar (sba) and incubated at 37 °c for 24 h to ensure that no bacteria were present on exterior surface of the blowflies. the presence of internal (ingested) b. anthracis was determined by homogenising each blowfly in 1 ml saline. the homogenised blowfly was briefly centrifuged (500×g for 5 s) to separate the blowfly debris from the supernatant. tenfold serial dilutions (100–108) of the supernatant were prepared and plated on pet agar and incubated at 37 °c for 48 h. the presence of b. anthracis spores was determined by plating 100 µl heat-treated undiluted supernatant onto pet followed by incubation at 37 °c for 48 h. isolation of bacillus anthracis from vegetation leaves from shrubs with blowfly defaecation or discard droplets around the infected carcasses were collected and transported at 4 °c to the nicd. each defaecation or discard droplet was removed from the leaves by using a transport swab pre-wetted with saline, followed by the swabs being rinsed in 1 ml sterile water. bacillus anthracis was detected by plating 100 µl undiluted wash before and after heat treatment on pet and sba plates followed by incubation at 37 °c for 48 h. leaves without any defaecation or discard droplets were sent to prof. k. eloff at the university of pretoria for identification and to determine antibacterial activity. the minimum inhibitory concentration (mic in mg/ml) of acetone extracts of the plant was tested against five bacterial organisms (escherichia coli, enterococcus faecalis, pseudomonas aeruginosa, staphylococcus aureus and b. anthracis sterne) using the protocol as previously described by eloff (1998). confirmation of bacillus anthracis bacillus anthracis isolates were identified based on the characteristic colony morphology (ground glass appearance, fairly flat, more tacky and white or grey-white), no haemolytic activity, penicillin sensitivity and gamma phage sensitivity after 24 h incubation at 37 °c on 5% sba as described by world health organization (2008). each confirmed b. anthracis colony of vegetative cell and spores was counted and recorded, followed by the calculation of total colony forming units per fly (cfu/fly). results the two carcasses were confirmed to be anthrax-positive through the presence of bacilli and endospores in a giemsa-stained blood smear as well as through the isolation of b. anthracis through culture. twenty-nine blowflies were collected from the kudu carcass and 28 blowflies from the impala carcass (table 1). chrysomya marginalis, ch. albiceps and lucilia spp. females were collected from the kudu carcass (with a female:male ratio of 3:1), whereas only ch. marginalis males and females were collected from the impala carcass (female:male ratio of 2:5) (table 1). table 1: summary of blowfly species collected from two anthrax carcasses during an anthrax outbreak in march 2014 in the kruger national park. only b. anthracis spores were isolated from the blowflies. a total of 26 (46%; 26/57) blowflies were positive for the presence of b. anthracis spores, with 66% (19/29) of blowflies collected from the kudu carcass and 25% (7/28) of the blowflies collected from the impala carcass (table 2). the ch. marginalis blowflies collected from the kudu carcass had an average internal and external spore count of 128 cfu/fly. the ch. albiceps blowfly collected from the kudu carcass was only positive on the interior with 20 cfu/fly. a low b. anthracis spore count of 10 cfu/fly was isolated from the exterior of the two lucilia spp. (appendix 1). the ch. marginalis blowflies collected from the male impala carcass had an average internal and external spore count of 26 cfu/fly (appendix 2). table 2: blowflies positive for bacillus anthracis spores collected from the confirmed anthrax kudu and impala carcasses during an anthrax outbreak in the kruger national park in march 2014. the shrub where blowflies defaecated or regurgitated after feeding on infected carcasses was identified as abutilon angulatum approximately 2.5 m from the kudu carcass and leaves were collected with blowfly defaecation or regurgitation spots. an average of ten defaecation or regurgitation spots was found per a. angulatum leaf. no b. anthracis was isolated from the a. angulatum shrub leaves using selective medium combined with or without heat treatment, but 170 spores were isolated from one defaecation or regurgitation spot cultured on 5% sba with no heat treatment. from a single thorny stem of an acacia tree, 500 spores per droplet for three out of eight defaecation or regurgitation spots were isolated on 5% sba without heat treatment. because of the absence of spores on non-selective medium, the antimicrobial activity of the a. angulatum shrub was tested and the results indicated that the mic of the a. angulatum acetone extract was most effective against e. coli (0.08 mg/ml) and had a moderate inhibitory effect towards e. faecalis, s. aureus and b. anthracis (sterne) with 0.31 mg/ml. discussion chrysomya albiceps, ch. marginalis and lucilia spp. were collected from two b. anthracis carcasses and their abundance confirms that these carcasses were 2–3 days old as indicated by the investigation of carcass-attending insect communities in the knp of braack (1985). chrysomya marginalis usually arrives within a couple of minutes to hours after death of animals and stays at a carcass for approximately 4 days, whereas ch. albiceps arrives a couple of hours after the death and remains at a carcass for about 3 days in the knp (braack 1985). lucilia spp. usually arrives at the same time as ch. marginalis, but in far less numbers. the major blowfly species collected from anthrax-infected carcasses was ch. marginalis of which 88% was b. anthracis culture-positive. no viable b. anthracis was isolated from any of the vegetation using selective media and could only be isolated using sba. based on the culture results using selective (pet) and enrichment (sba) media, it seems that the selective medium has an inhibitory effect on b. anthracis spore germination from leaves, which was not the case with germination from the blowflies themselves. because of initial negative results of the selective media from the a. angulatum leaves, we tested for antibacterial activity and found a moderate inhibitory effect on the non-virulent sterne strain of b. anthracis by phytocompounds within a. angulatum. only spores, and no vegetative organisms, from b. anthracis were isolated from the interior and exterior of blowflies, which could be attributed to various factors that could have killed the more labile vegetative organisms. these factors include the freezing of the flies in liquid nitrogen, collection time and the period between collection and processing of the flies. graham-smith (1913) fed musca domestica and calliphora erythrocephala on b. anthracis vegetative cells and spores and recovered only b. anthracis spores for up to ten days from fly legs and wings and seven days from both the gut and the crop. vegetative cells were present on legs and wings for up to 24 h, lending further evidence to our findings. in this study, the collected flies were frozen in liquid nitrogen. haines (1938) found that 90% b. mesentericus spores were viable after –70 °c freezing with varying loss of viability of the vegetative cells and young et al. (1968) suggested that vegetative cells of spore-forming bacteria are more susceptible to damage from freeze-thawing than vegetative cells of non-spore-forming bacteria. therefore, it is presumed that the freezing of the flies in liquid nitrogen did not influence the spore count significantly, as we only found spores. the absence of vegetative cells could be because of the use of liquid nitrogen to kill the blowflies. various researchers have determined that flies can potentially transmit b. anthracis to their surroundings and that these flies are carriers of the bacteria for numerous days (blackburn et al. 2010, 2014; hugh-jones & blackburn 2009; von terzi et al. 2014). according to de vos (1990), explosions of blowfly populations preceded the anthrax outbreaks in the knp in 1960 and 1970. blackburn et al. (2010) speculated that blowflies and flesh flies are responsible for the spread of an anthrax outbreak in white-tailed deer in the usa, which was later confirmed by isolating b. anthracis dna from leaves collected 21 days post-outbreak as well as viable b. anthracis from blowfly larvae and one adult that can potentially infect the surrounding vegetation (blackburn et al. 2014). this coincides with the findings of this study where b. anthracis could not be isolated from leaves using selective medium. this could explain why only b. anthracis dna on leaves was detected by blackburn et al. (2014). the suppressive nature of selective media on b. anthracis from environmental samples needs further investigation. two recent studies investigated fly species as potential mechanical or biological vectors of b. anthracis. fasanella et al. (2010) allowed the common housefly (m. domestica) to feed on b. anthracis–infected rabbit carcasses or b. anthracis–contaminated blood and indicated the presence of b. anthracis spores in defaecation or vomit spots as well as hypothesised that anthrax spores are able to germinate and replicate in the gut content of m. domestica. these authors hypothesise that the housefly has the potential to act as a biological vector for b. anthracis. von terzi et al. (2014) demonstrated that b. anthracis vegetative cells are unable to multiply inside the gut of calliphora vicina and cannot survive for longer than a week. these authors’ results show high numbers of viable b. anthracis indicating the role of c. vicina as only mechanical vectors as previously hypothesised (blackburn et al. 2010; hugh-jones & blackburn 2009; hugh-jones & de vos 2002). the ch. albiceps had less spores per fly (20 cfu/fly) compared to ch. marginalis (128 cfu/fly) collected at the kudu carcass. only ch. marginalis was caught at the impala carcass with 25 cfu/fly. the reason for the larger spore count present on flies at the kudu carcass could be attributed to the bacterial load of the host, or more sporulation as the kudu carcass was opened by scavengers compared to the partially opened impala carcass that might have been protected from scavengers by the acacia shrub. the total cfus isolated per fly reflect the potential infectious load of a fly with an average of 87 b. anthracis cfu/fly depending on the blowfly species. it is not known how many spores were defaecated onto the surrounding vegetation by flies, but 170 spores were isolated from a defaecated spot on the a. angulatum leave. our results show that b. anthracis is present in and on blowflies that actively fed on an anthrax carcass and that the faecal or discard droplets left on leaves are infective. this can then lead to the assumption that blowflies can aid in the dissemination of b. anthracis to susceptible host species. as anthrax carcass sites are the primary sources for future infections (turner et al. 2014, 2016), it will depend if the spore doses from surrounding infected vegetation are lethal. considering the 170–500 spores per leaf or twig (that weighed approximately 1 g – 2 g) found in this study, a lethal dose of around 105–107 for gastrointestinal anthrax (turner et al. 2016) can easily be ingested during anthrax outbreaks as impala ingests 0.9 kg – 1.9 kg per day (furstenburg 2002) and kudu 3.7 kg – 5.0 kg per day (http://www.wildliferanching.com/content/kudu-tragelaphus-strepsiceros). however, because of the small sample size (host species and blowflies), further research is necessary to investigate the spore load present in the different host species and cfu on and defaecated by the different blowfly species. despite the unique findings of this study, specifically in the knp, it only involved two carcasses of different species with blowflies being collected at one specific time point. to obtain a clearer view of the infectious load of blowflies feeding on a carcass under different climatic conditions, a temporal study over a longer time period should be conducted. ideally, blowflies should be collected daily from carcasses and the surrounding vegetation until the carcasses are completely decomposed giving insights into the extent of the role that blowflies may play in anthrax transmission. while most studies indicate that blowflies play a role as a vector of b. anthracis, the epidemiological significance of this role is difficult to determine because various factors need to be considered, including the survival of the spores on or in blowflies, the spore load in defaecation or vomit spots of blowflies, and hence, on the vegetation, the latter depending on climatic conditions and antimicrobial activity of the plant(s), and last but not least the effective dose required by susceptible host species. conclusion in conclusion, the results obtained from this study indicate that blowflies feeding on an anthrax-contaminated carcass can contaminate surrounding vegetation and aid in the continuous cycle of b. anthracis. if there are high numbers of blowflies feeding on a carcass, it can lead to high bacillus or spore numbers being deposited on the surrounding vegetation. this can then lead to a new infection of a susceptible host through the ingestion of contaminated vegetation. acknowledgements this research was funded by the national research foundation, deutsche forschungsgemeinschaft and agricultural sector education and training authority (agriseta). the authors would like to acknowledge sanparks for approving this research in the kruger national park and skukuza state veterinary services, department of agriculture, forestry and fisheries for support during collection. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions l.b. did the experiments and wrote the manuscript. knowledge transfer of techniques to l.b. used in this study was done by a.h., a.d. and a.g. a.h. did most of the vegetation experiments. a.g. was responsible for the polymerase chain reaction (pcr) identification of the blowflies. w.b., j.r. and h.v.h. participated in the design of the study and co-supervised the first author. w.b. has been the principal investigator of the dfg project be2157/4-1. h.v.h. and w.b. were provided funding through agriseta and the deutsche forschungsgemeinschaft, respectively. l.b. received a national research foundation (nrf) freestanding innovation scholarship. all authors participated in critical revision of the manuscript. references blackburn, j.k., curtis, a., hadfield, t.l., o’shea, b., mitchell, m.a. & hugh-jones, m., 2010, ‘confirmation of bacillus anthracis from flesh-eating flies collected during a west texas anthrax season’, journal of wildlife diseases 46, 918–922. https://doi.org/10.7589/0090-3558-46.3.918 blackburn, j.k., mullins, j.c., van ert, m., hadfield, t.l., o’shea, b. & hugh-jones, m., 2014, ‘the necrophagous fly anthrax transmission pathway: empirical and genetic evidence from a wildlife epizootic in west texas 2010’, vector-borne and zoonotic diseases 14, 576–583. https://doi.org/10.1089/vbz.2013.1538 braack, l.e.o., 1985, ‘an ecological investigation of the insects associated with exposed carcasses in the northern kruger national park: a study of populations and communities’, phd thesis, department of entomology, university of natal, south africa. braack, l.e.o. & de vos, v., 1990, ‘feeding habits and flight range of blow-flies (chrysomyia spp.) in relation to anthrax transmission in the kruger national park, south africa’, the onderstepoort journal of veterinary research 57, 141–142. braack, l.e.o. & retief, p.f., 1986, ‘dispersal, density and habitat preference of the blow-flies chrysomya albiceps (wd.) and chrysomya marginalis (wd.) (diptera: calliphoridae)’, the onderstepoort journal of veterinary research 53, 13–18. de vos, v., 1990, ‘the ecology of anthrax in the kruger national park, south africa’, salisbury medical bulletin 68s, 19–23. de vos, v., 1994, ‘anthrax’, in j.a.w. coetzer, g.r. thomson & r.c. tustin (eds.), infectious diseases of livestock with special reference to southern africa, pp. 1262–1289, vol. 2, oxford university press, cape town, south africa. dragon, d.c. & rennie, r.p. 1995, ‘the ecology of anthrax spores: tough but not invincible’, the canadian veterinary journal 36, 295–301. eloff, j.n., 1998, ‘a sensitive and quick microplate method to determine the minimal inhibitory concentration of plant extracts for bacteria’, planta medica 64, 711–713. https://doi.org/10.1055/s-2006-957563 fasanella, a., scasciamacchia, s., garofolo, g., giangaspero, a., tarsitano, e. & adone, r., 2010, ‘evaluation of the house fly musca domestica as a mechanical vector for an anthrax’, plos one 5, 1–5. https://doi.org/10.1371/journal.pone.0012219 furstenburg, d., 2002, impala ‘rooibok’ aepyceros melampus, viewed 24 november 2010, from http://gadi.agric.za/articles/furstenburg_d/impala.php graham-smith, g.s., 1913, flies and disease: non-bloodsucking flies, cambridge university press, london. haines, r.b., 1938, ‘the effect of freezing on bacteria’, proceedings of the royal society b: biological sciences 124, 451–463. https://doi.org/10.1098/rspb.1938.0005 hugh-jones, m.e. & blackburn, j.k., 2009, ‘the ecology of bacillus anthracis’, molecular aspects of medicine 30, 356–367. https://doi.org/10.1016/j.mam.2009.08.003 hugh-jones, m.e. & de vos, v., 2002, ‘anthrax and wildlife’, revue scientifique et technique (international office des epizootics) 21, 359–383. https://doi.org/10.20506/rst.21.2.1336 koekemoer, l.l., kamau, l., hunt, r.h. & coetzee, m., 2002, ‘a cocktail polymerase chain reaction assay to identify members of the anopheles funestus (diptera: culicidae) group’, american journal of tropical medicine and hygiene 6, 804–811. https://doi.org/10.4269/ajtmh.2002.66.804 pienaar, u., de v., 1961, ‘a second outbreak of anthrax amongst game animals in the kruger national park 5th june to 11th october 1960’, koedoe 4, 4–14. https://doi.org/10.4102/koedoe.v4i1.824 pienaar, u., de v., 1967, ‘epidemiology of anthrax in wild animals and the control of anthrax epizootics in the kruger national park, south africa’, federation proceedings 26, 1496–1502. turell, m.j. & knudson, g.b., 1987, ‘mechanical transmission of bacillus anthracis by stable flies (stomoxys calcitrans) and mosquitoes (aedes aegypti and aedes taeniorhynchus)’, infection and immunity 55, 1859–1861. turner, w.c., kausrud, k.l., beyer, w., easterday, w.r., barandongo, z.r., blaschke, e. et al., 2016, ‘lethal exposure: an integrated approach to pathogen transmission via environmental reservoirs’, scientific reports 6, 27311. https://doi.org/10.1038/srep27311 turner, w.c., kausrud, k.l., krishnappa, y.s., cromsigt, j.p.g.m., ganz, h. h., mapaure, i. et al., 2014, ‘fatal attraction: vegetation responses to nutrient inputs attract herbivores to infectious anthrax carcass sites’, proceedings of the royal society of london series b: biological sciences 281, 20141785. https://doi.org/10.1098/rspb.2014.1785 von terzi, b., turnbull, p.c.b., bellan, s.e. & beyer, w., 2014, ‘failure of sterneand pasteur-like strains of bacillus anthracis to replicate and survive in the urban bluebottle blow fly calliphora vicina under laboratory conditions’, plos one 9, 1–7. https://doi.org/10.1371/journal.pone.0083860 world health organization, 2008, anthrax in humans and animals, 4th edn., who press, geneva. young, r.s., deal, p.h. & whitfield, o., 1968, ‘the response of spore-forming vs. nonspore-forming bacteria to diurnal freezing and thawing’, origins of life and evolution of biospheres 1, 113–117. https://doi.org/10.1007/bf00924233 zumpt, f., 1965, myiasis in man and animals in the old world, butterworths, london. appendix 1 table 1-a1: presence and number of colony forming units of bacillus anthracis on and/or in blowflies collected from the confirmed anthrax-positive kudu carcass (2–3 day old) during anthrax outbreak in kruger national park in 2014. appendix 2 table 1-a2: presence and number of colony forming units of bacillus anthracis on and/or in blowflies collected from the confirmed anthrax-positive impala carcass (2–3 days old) during anthrax outbreak in kruger national park in 2014. filelist convert a pdf file! page 1 page 2 page 3 page 4 koedoe 63-1_2021_contents.indd http://www.koedoe.co.za open access table of contents original research mammals in the mountains: an historical review and updated checklist of the mammals of the mountain zebra national park daniel m. parker koedoe | vol 63, no 1 | a1683 | 12 july 2021 original research first guidelines and suggested best protocol for surveying african elephants (loxodonta africana) using a drone welsey l. hartmann, vicki fishlock, alison leslie koedoe | vol 63, no 1 | a1687 | 06 september 2021 original research the spatial distribution of the woodland communities and their associated environmental drivers in the golden gate highlands national park, south africa mahlomola e. daemane, abel ramoelo, samuel adelabu koedoe | vol 63, no 1 | a1672 | 23 september 2021 original research distribution and diversity of entomopathogenic nematodes (steinernematidae and heterorhabditidae) in a south african nature reserve isiah nthenga, rinus knoetze, antoinette p. malan koedoe | vol 63, no 1 | a1661 | 22 november 2021 original research a reclassification and description of the waterberg mountain vegetation of the marakele national park, limpopo province, south africa petrus j. van staden, george j. bredenkamp, hugo bezuidenhout, leslie r. brown koedoe | vol 63, no 1 | a1689 | 10 december 2021 original research the first sub-meter resolution digital elevation model of the kruger national park, south africa kai heckel, marcel urban, jean-sébastien bouffard, jussi baade, peter boucher, andrew davies, evan g. hockridge, wolfgang lück, jonas ziemer, izak smit, bernhard jacobs, mark norris-rogers, christiane schmullius koedoe | vol 63, no 1 | a1679 | 17 december 2021 correction erratum: elephant population responses to increased density in kruger national park albertus s. louw, sandra macfadyen, sam ferreira, cang hui koedoe | vol 63, no 1 | a1691 | 13 december 2021 reviewer acknowledgement koedoe | vol 63, no 1 | a1704 | 21 december 2021 76 86 95 104 111 127 140 141 page i of i table of contents book review review of bats of southern and central africa, second edition leigh r. richards koedoe | vol 63, no 1 | a1678 | 30 april 2021 short communication protecting and preserving south african aeolianite surfaces from graffiti charles w. helm, hayley c. cawthra, richard m. cowling, jan c. de vynck, martin g. lockley, curtis w. marean, mark g. dixon, carina j.z. helm, willo stear, guy h.h. thesen, jan a. venter koedoe | vol 63, no 1 | a1656 | 11 march 2021 scientific letter is climate change a concern for the ownership of game within fenced wildlife areas? haemish i. melville, robyn s. hetem, w. maartin strauss koedoe | vol 63, no 1 | a1673 | 27 july 2021 review article a review of fire management practices in african savannaprotected areas willem a. nieman, brian w. van wilgen, alison j. leslie koedoe | vol 63, no 1 | a1655 | 15 february 2021 original research management guidelines for the conservation of heritage resources in wakkerstroom, mpumalanga reitumetse m.h. sethaba, mary scholes koedoe | vol 63, no 1 | a1647 | 20 january 2021 original research a preliminary investigation of exposure to rabies virus in selected wildlife in the kruger national park, south africa leana rossouw, carin boshoff, claude sabeta, johann kotzé koedoe | vol 63, no 1 | a1651 | 12 march 2021 original research adaptive and transformative learning in environmental water management: implementing the crocodile river’s ecological reserve in kruger national park, south africa craig a. mcloughlin, eddie s. riddell, robin m. petersen, jacques venter koedoe | vol 63, no 1 | a1663 | 20 april 2021 original research elephant population responses to increased density in kruger national park albertus s. louw, sandra macfadyen, sam ferreira, cang hui koedoe | vol 63, no 1 | a1660 | 10 june 2021 1 3 8 12 25 36 44 63 vol 63, no 1 (2021) issn: 0075-6458 (print) | issn: 2071-0771 (online)koedoe the contemporary geomorphology of the letaba ri ver in the kr uger national pa rk b.p. moon a nd g. l. h e ritage moon. b.p. and g.l. heritage. 200 1. the contemporary geomorphology of the le taba ri ver in the kruger national park . koedoe 44( 1): 45-55. pretoria. lssn 0075-6458. the letaba rive r drains part of nonhem provi nce in nonh-east south africa. its catchment has been mod ified significantl y by hu man activ ity which has affec ted the flow regime: it experiences only ephemera l flows through the kruger national park to its confl uence wi th the oli fants river. although the lelaba is similar to the other rivers in the kruger national park in thai it displays some bedrock-infl uenced channel features, increased sediment delivery from the degraded catchment upstream has resulted in extensive atl uviation within the channel. sections of channel flowing over bedrock with no sediment covering are rare, and the river com prises a series of channel types: mixed anastomosi ng, all uvial braided, mixed pool-rapid and al luvial single thread. eac h is characterised by a differe nt combi nation of morphological units which relate to the degree of all uviation in the channel. these channel types are described in detai l and infere nces are made concern in g the ir fonna tion and maintenance from fie ld observat ion and measu rement. key words: channel type, channe l change, geomorphology, lelaba river, mot'phological units. b.p. moon. centre/or water in the environment. universityo/ the witwatersrand. i jon smuts avenue. johannesburg. republic 0/ soulh a/rica (i32abm@atlas.wits.ac.tjl); g.l heritage, department 0/ geography, peel building, unil'ersity 0/ salford, manchester m54wt, united kingdom (gl heritage@saljorri.ac.uk). int rod uct io n duri ng the las t ce ntury the leta ba ri ver in the kru ger national park has changed fro m a perenn ial to a n ep heme ral ri ver (steffen robe rtso n & ki rste n i 990a). in c reased abstraction of water fo r development. and the construc ti on of da ms, have led to a modi fication of the flow regime and a s hift in the ba lance of catc hme nt c ont rol va ria bles (heritage et a l. 1997). th e consequ e nt marked geomorph olog ica l c hanges have bee n doc umen te d ( vog t 1992), and a recen tly-comp leted fi ve-year proj ect o n the ri vers of th e kruge r nat io na l park has focused o n an unde rsta nd ing of th ose changes (he ri tage et aj. 1997). increas ing water abstrac tion and an expansion of agricu ltural practices are like ly to result in further flow reduc tio n and increased sedime nt issn 0075-6458 " inputs to the ri ver ( birkhead et al. 1995) leading to furth er mo rph olog ica l c han ge . recently the re has bee n a marked growth in research into th e natu re and d ynamics of south afri can ri ver systems (fo r exam ple, the work of ro wntree & wadeson 1996; wadeson & rowntree 1996; heritage et al. in press; ro untree et al. in press). in th is paper, fo llowi ng an ove rvi ew of the letaba catc hme nt. the contemporary geomorphology of the le taba ri ver is descri bed within the framework of its compone nt c hannel types. four di ffere nt c hanne l types have been recogni sed : mi xed anas to mos ing, all uvial braided. mi xed pool-rapid and all uv ia l s ingle thread, each re flec ting a different combi nation of mo rphological units. the nature of change in the differe nt cha nne l types is discussed in re lat ion to reduced flo ws and in creased a lluvi ation ove r time. kued(n 4411 (200t) fig. i. the letaba. river catchment. nonhern province. south africa. (a) location. (b) precipitation (mmla) and evaporation (mmla). (c) sed iment prod uction (tkm"a). (after stelten robertson & kirsten i99ob). kodof' 4411 (200 1) 46 the le ta ba catchment the letaba catc hmenl covers an area of 13400 km1 in the northern prov in ce of south afri ca. in the lowveld region its major tri butaries are the molotsi and kl ei n-letaba rive rs, and the let aba join s the olifan ts near the mozambique border (fi g. la). the catchment is class ified as se mi-arid. receivin g be tween 500 and 1800 mm of ra infa ll in the moun · tai nous western parts of the catchment , fa llin g to between 450 mm and 700 mm in the east (fi g. ib). evaporation is hi gh, rangi ng from 1400 mm in the west to 1900 mm in the east (fig. ib). precipitation is concentrat· ed in the summer month s whereas evaporat ion potent ial is more even ly di stri buted (s teffen robert son & ki rs ten i 99oa). runoff is now sea· sonal and became ephemera l during the drought of the 1990s when sur· face flow in the lower reac hes occurred onl y in response to severe stonn eve nts in the catchment . virg in mean an nua l run off has been est imated at 556 milli on cubic metres per annu m (m ml/a) (s teffe n robenson & kirsten 1990a) compared with the present mar of 323 mml/a. in the upper catchment . outsi de the park, ex tensive areas of land are used for fo restry and commerc ial agri cul ture. intense subsistence fann ing and irrigation also occ urs along th e ri ver where access to water is good (s teffen robe rtso n & ki rste n i 99oa). forested areas (47600 ha in 1985) require a water vol um e of 64 mm'/a (s teffe n robenson & ki rsten 1990a). irrigati on of fru it, vegetables and grai n cash crops req ui res 220 mm'/a. and domesti c and indu stri al water use amoun ts to approx imatel y 17 mml/a. a furthe r approximately 7 mm1/a is exported to urban areas ou tside the ca tchment. potenti al in crease in demand as a result of increasing agricu llu ral. industrial and rural water tssn 0075-6458 • ---""""'• + .........'-'*"~ d _siibrim . miqd_~ cj ............ _~1:!i:!i ciiannei typ. btuk --.... 1 51101 co w% 189 1011 1213 , b _sllbri"". ubed ""sior'nosing t2'2i -----+ ',---,'~ it : c + • " ......... siieid .... _ miled~ ~ mixed poolirapid cj """"" "'" ---iii: i: ;: ... iii 0123 . s 8 r'il01112131. ' 516171ai9 . , ~ fig. 2: the sequence of c hannel types along the letaba river in the kroger national park (seq uence a-d: upstream 10 downstream, wesi to east through the reserve 10 the conllllence with the olifanls ri ver). usage is like ly to o utstrip water availability within the catchme nt , and conseq ue ntly land degradation and sedime nt produc tion will increase (s teffen, robertson & kirsten 1990b; birkhead et ai. 1995). it is therefore likely that the already degrad ed letaba rive r will ex peri e nce furt he r reductions in water vol ume and an in crease in sed ime nt input. geomorpholog ica l and ecological c hanges will occur as a resu lt . and it is for ih is reason that the nature of geo morphologica l chan ge needs to be understood. sediment produ ction in the letaba catc hment has been estimated from land use, so il type. geomorpho logy, vegetation an d rainfall variab les (s te ffe n ro bertso n & kirsten 1990b). yi e lds ra nge fro m 150 tkm2/a to in excess of 400 tkm1/a (fig. i c) with the hi ghest yields occuning in the de nse ly populated rura l areas in the west of the catc hment . thi s issn 0015-6458 47 material will eve ntu all y be tran sferred from th e drainage bas in into the ri ver (knigh ton 1991). affecting a ll reaches of the letaba river to vary ing degrees. increas ing press ures on the catchme nt as a result o f greate r water abstracti o n and an ex pa nsio n of agric ultural practices are like ly to result in fu rthe r now reduction and inc reased sediment inputs to the ri ver (birkhead et of. 1995 ). geomorp hol ogy the letaba ri ve r in the lowveld cons ists of a mac ro-channel th at is inc ised into the surrounding landscape by so me 510 m o n average and ove r 30 m in the gorge close to th e conflu e nce wi th the o li fa nt s ri ve r ( heritage el 01. 1997). with in the mac roc hannel there are o ne or more sub-channel s that are ac tivated as discharge ri ses follo wk~d~441 t (2001) table i description of the morphological units found on the utan river in the kurger national park morphological unit rapid rime pool braid bar lateral bar point bar bedrock core bar l<ebm bedrock backwater alluvial backwater river cliff apical poo l rip channel boulder bed armoured =, alluvial distributary bedrock distributary terrace descri pt ion steep bedrock sections, hi gh veloc ity concentrated fl ow. accumulation of coarser sediment as a topographic high point as part of a pool rime seque nce. topographic low point characterised by finer sediments, as part of a pool-ri m e sequence. accumu lation of sediment in mid-channe l causing the flow to diverge over a scale \hat approx imates to the channel width. acc um ulation of sedi ment attached to the side of the channel. may occur sequentially down· stream as altern ate bars. accumulation of sediment on the inside of a meander bend. accumulatio n o f finer sediment on top of bedrock in bedrock anastomosing areas. accumulation o f sediment in the lee of flow obstructions. stationary or near stationary bodies o f water in bedroc k, adjace nt to the active c hannel. stationary o r near stationary bodies of water in alluvium, adjacent to the active channel. vertical or near vertical alluvial erosion face . deep section of channel located on the outer bend of a meander, associa ted wi th point bars. hi gh discharge distributary channel on the inside of poi nt and lateral bars. accumulation of locally derived material exceeding 0.25 m. accumulation of coarser sedi ments due to winnowing of finer material. individua l active channel in an alluvial braided or anastomosi ng system. individual active channe l in a bedrock anastomos in g system. relic fl oodplain or valley floor deposits above the present river level. in g rainfall eve nts. th e macro-c hanne l ex tends across th e width of the incised valley and is characterised by discrete sedimentary depos its forming all uvial morphological units many of which are colonised by ripari an vegetation. that on the sabie river (heritage & moon 2(00). sedi ment from the degraded catchment has accu mulated within th e macrochannel creatin g a preponderance of allu vial reac hes. these, often ex tensive, allu vial areas are characterised by sparsel y vegetated sand sheets and associated lateral terrace-like depos its of cohesive all uvium. wet season floods inundate the sand sheets whereas the terrace features are overtopped only by less frequent flows of greater mag nitude . although the river has incised several metre s in to bedrock and allu vial material to fo nnthe macro-c hannel, the influ ence of bedrock on th e channe l floor is considerabl y less than koedoe 4411 (200 1) 48 tssn 0075-6458 table 2 distributiq1i of tht morphological u1iils fou1id 011 tht utaba ril,tr;1i tht knlgtr national park --morphological unit mixed anastomosing -rapid ./ rime pool rime apical rapid braid bar lateral bar point bar bedrock core bar ./ '" "" ./ backwater bedroc k ./ allu vial mixed river cliff apical pool rip channe l bou lder bed """""" ~ ./ distributary bedrock ./ alluvial ./ mixed ./ t,~ bedroc k· influenced reaches are limited in exten t and ex hibit considerable alluv ial cove r in the fonn of un co nsolidaled sands and fine gravel s. in such reaches the channel divides 10 fonn a multipl e di stributary net· work flowin g around bedrock o ut crops exposed in th e bed of the channel. aerial photographs taken in 1989 together with field observations were used to categorise the letaba ri ver in a hi erarchi ca l framewor k. the basic components of the framework are morpholog ica l units. eac h with a di stinctive character and microhabi· issn 0075-6458 ----alluvial mixed pool. alluvial single. braided rapid thread -----------49 ./ ./ ./ ./ ./ ./ ./ ./ ./ = ./ ./ ./ ./ = ./ ./ ./ ./ ./ ./ ./ ./ ./ ./ ./ ./ tat (van niekerk & heritage 1993). morpho-logical un its are seen to associate together to fonn different channel types (heritage et al. in press). there is a functi onal re lati onship belween differem chann el lypes wh ich may be grouped int o reac hes. where one channel type may be influ encing directl y the nature of the channel type upstream or downstream . the interaction of a range of flows on bedrock and all uv ium has ge neraled a varied assemblage of morpho logical un its (tab le i). kfh dfh 4411 (200 1) fig. 3: a mixed anastomosing channel type on the: letaba river in ihe kruge r national park . visual in terpre tation of aerial photogra ph s ( i: 10 000 sca le), ae ri al video footage take n during winter low flow co ndition s and extensive fieldwork prov ided the basis for identify in g characteristic mo rphological asse mblage s or channe l types. four common chan nel types (mi xed anastomos ing, mixed pool-rap id, alluv ial single thread and allu vial braided) are found along the letaba ri ve r (fig. 2). gradations betwee n the four primary channel types indicate that individua l channe l types are pan of a spat ial and tempora l continuum of cha nne l fonn. the channel types ex hibit ed by th e letaba are described below. mixed anastomosillg some sect ions of the river di sp lay a wider macro-c hannel than the average (fig. 3). bot h sedimentary and bedrock features (table 2) are found in the channel which is characterised by a tonuous distri butary netkoedoe4411 (200 1) work that follows weakn esses in th e bedrock and covers the ent ire width of the macrochannel . waler leve ls within the various di stributary chann els are oft en al different elevations as a result of flow being conce ntrated along lines of weakness as defined by the bedroc k template. loca ll y bedroc k out crops within the channe ls act to obstruct the flow effectively acting as local hydrauli c co ntrol s. this results in a di scontinuou s phreati c surface with no sub-s urface water fl ow linkage. the all uvia l deposits covering muc h of the bedrock exist as a thin (less than 1.5 m) ve neer on the interfl uves be twee n the dislributary chan nels. the material is generally unconso lidated or only se mi -consol idated bei ng composed of sa nds and fi ne gravel s with little clay or silt and ih ere are few large bedrock core bar deposits . occasional deposi ts of coarse r grave ls and cobb les exist as armoured areas an d lag deposits in the sect ions of the channel dominated by bedrock. distri bu tary channels tend to be lined with reed and sc rub vegetation. aerial photographic analy sis of the effects of high flow s (w hich inundate the entire macrochan nel ) on mix ed anastomos ing channels has revea led that thei r planfonn is stable in that fl ow returns 10 pre-existing channels as the flood stage d imi ni shes. alii/vial braided short sec ti ons of river are characleri sed by ex tensive san d sheets (table 2) dissected by a multi -c hannel network of alluvial di stributaries that flow over the sandy surface . the lowfl ow d istributary channels split and rejo in arou nd unco nsol idated mid-channel deposil s over di slances that approximate the distri bulary width (fig. 4). flow leve ls in these chan nels are similar, ind icati ng a wellconnecled phreatic surface across the macrochanne l. pl anfor m change is rapid and observat ion of the river afler a flood reveals that thi s is often associated with new channel cutti ng on the falling lim b of the flood hydrograph , except where reed growth has paniall y slabil ised and restricted the lateral issn 0075-6458 fig. 4: }\ bmidcd ch;lii lleltypc on the letaha river in the kn' gcr national p;lr k. migrat io n of di slribu taries. exlens ive sa nd sheel s dom ina te the macro-c han nel: sta lled s and dunes and sca~()nal d istribu lary channel s formed d uring the prev io us flood event complicate iheir morpho logy. fi ne ma leria l doc s not accumu late on the s urface of the s:.i11 d shee ts (fig. 4 ). it is suggested that this is due to the li ne ma te rial. carr ied by flood flow s. re mains in s uspe ns ion because of hig h e ne rgy level s :l nd contin ue dow i1 stream. lower e nergy levels arc expe rienced close to the mac ro-channel margi ns whe re li nes arc de l)qs iled c re at in g s ublinear terraceli ke featu re s composed of clays. s ilts a nd sand y depo s it s. occ as ion:t1 allu vial issn 0075-6-t58 " fig . 5: ;\ mixed ixkji-rdpid dwnncltypc un lhe lc1~ ­ ha ri .'cr in the kn' ger national p;,rk. backwat ers develop (fig. 4) fo llowing now recessio n and i)qo i iso lation. fin er material characterises the bed o f these areas. mixl'{/l'orj/ -mpid bedrock ex i)qsures o n th e macro-c hannel floor influence ihe course and st nl ct ure of ihe lowfl ow channe l and ge nera le a se ries o f bedrock ra pid s and assoc iated upstrea m pool s (fig. 5). o flelll he now netwoi-k for ms a number of sub -pa rallel pools separat ed by bedrock rap ids ( fig. 5). evide nce from s im ilar fealures on the sabi e river (c hes hire 1994) ind ica te ihall ocal geology. wi th alternating weake !" (pools) and more res isl:11l1 litho log ies (rapi ds). influence s the nature of these morpho logical u nils (tab le 2). mixed poo l-rapid c hannel s arc common in th e highe r ene rgy zones along ihe ri"c r. p:lniclilarly ko.·,/"" -1 -1 / 1 ( 2001) fig . 6: ,\ 11 ~liu\'i,ll s ingte thread i:hannc t !ype on ih e lc!:iba river in ihe kru ger nali()n~1 park (w,l\'c trai ns ca n be m:c n in !he lop ri ght uf ihe ,a nd ~hcc!). whe re sedimc nt suppl y is locall y redu ced. for examp lc dow nstrea m of s mall dams. cohes ive sed imen ta ry de pos it s ar..: res tric ted to occ as ion al dissec ted re mn a nt s o f more ex te ns ive tc rrace fe mures o ft e n fo rmin g ex posed 'ri ve r c li ffs' nca r the s ides of the mac ro-c hannel. allu viati oll across mu ch o f th e mac ro-c hannel f0 1"ms cx te nsive but thin sand sheets infillin g lowe r areas of ex pose d bedroc k pa ve mc nt. coarser arm o ured area s occ asionally fo rm rime -lik e fe atures of la g mat e rial in th e ac ti ve c hannel (fi g. 7a). ailil 1"ia/ .~il1gll! ihread therc is ev id e nce o f swi tching bet wee n bra ided. mi xed an asto mosing a nd s ingle thread sys t e l1l ~ in allu vial sec tions of the rive r. in s ingle th read reaches low n ows are conveyed along a s ingle a ll uv ial channel across an exten sive sand s hee t whe re no bedrock feature s arc ev ident (table 2). oft e n suc h channel s arc close 10 the s ide of the mac ro-chan nel. :md alternate fro m s ide to side bearing no rei;ltion to the m:!c ro·c hannel planfonn (fi g . 6). llle re arc s igns of in cipi ent braidin g with the comm o n devel o pm e nt of shallow lo w now rime s composed of s and and fin e gra vel s and m id -c hann e l bars (fig.7b). t he e xten s ive c hann el-s ide rced growth. howeve r. restricts lat e ral e rosio n :lnd stabili ses the planfo rm . th e restric tion o n lateral movement is often compe nsated for by venical eros ioll. and as a res ult the lown o w channe l appears ove r-deep in re lati o n to its width. an exte nsive network of seasonal c hannel s exist s on the surf,lce of th e sand sheet . s imilar to that found in bra id ed sectio ns. their dcvelopmcn t is esse nt ia ll y the same in both c hannel types (fi g. 7b ). t he pres enc e of e xte ns ive coh es ive alluv ial deposi ts as lateral mac ro-c hannel te rra cclike feature s is also related 10 s i milar features co mpo sed of fine r mate ri .. 1 in braided secti o ns. and .. gg rad e follo wing majo r n ood e ve nts. 52 c h a nnel c han ge 0 11 t he lel a ba rive r chan ges 10 th e let ab:. river h:! \'e bee n an alysed us in g ae rial pholograph s of the ri ver in the kru ge r national park fro m the 1960s 10 the 1980s (vogt 1992). th ese includ e pos iti o nal c han ges in sin gle thre ad c hannels. occas io nal ne w c h;mnels be in g cui throu g h ex isting sedim entary de pos it s. and occasio na l ne w in -channel late ral fea ture s whi ch wc re depos ited in th e 1960s and 1980s. limit ed di sscc ti o n o f large po int bars occ urred in 1974 . in multi -thread reac hes. the abando nment of di stributary c hannel s as a res ult of bar coalesce nce occ urred in the 1960s and 1980 ), co ntc mpo raneo us ly w ith th e depositi o n of in -c hann el lat e ral feature s. new channel s were clltthro ugh ex istin g sedim e nt s in th e 1970s. th e ev ide ncc sugge sts pe riods of de pos it io n in the 1960s and 198 0s se parated by an e ros io nal phase in th e 1970s (vog t 1992). tssn 0075·6-' 51; .. . ' . , ~ •• . ' .. fig. 7: (a) ,\rmourcu :orca in a mixed pool-r..pitl channel. nol\ i~ right 10 lefl :ulu ihe ch:umd "idlh is approximately 40m. (b) ~i ng lc thread channel hightigluing :111 cx!.::il,j,·c \;ij\u ,heet. f\. ... -d ·cu\crcd mid-channel b;,r ~nd ct)hc~i\'c iuw ictr.i!.-c deposits <ii the m:trt;in\ of the m:,cro.-challncl. til\' ;icli\'c ('halilid is appm.xirn:ucly 10m ilidc. ev id e nce of short -term c han ge has bee n ob tained from a fe-survey of twenly c ro:.ssections c~tablishcd on the lelah:! i~ivcr in 1993 (heri tage l 'l al. 1997). th e oood of february 1995. whic h geller.lled it i>cak now gauge reading of 655 1111/s oil enge lha rd o;ull. close to lelab:! rest camp in the kruger national park, resulted in rcrna rbbly litt le cha nnel change. low-flow channel:issn 0()75 · (>-i5!! rema in ed slab lc. in so me sec tions there was dissection of sa nd ~ho.!ets a~ a res ult of cha nnel dev el oplllo.!l1\ during the waning-flow of a flood. the sig nili cant c han ges we re in b ulk sediment mass. "tor:.jge o n the sand sheets with deposi ti o n and erosion occurring a lo ng the river. the p.l\lcrn of c hange was not related 10 th e c hann eltypc distribution ( heritage ;'/ 01. 1997). this may reflecl ihe 1ll0\'e rn e llt of sedime nt s lu gs as d une field wavc trains al o ng th e system ( fi g. 6 ). con s id er.lble depos its of fi ne se dime nt we re left o n teit:icc fe'ltures o n the s ides o f the n1ac rochannel . th e available evide nce s uggests, wi th the cxcept ion o f loc .. li \cd e rosio n and some minor c hannel c hange resulling from infre(iuent hi g h. n ow cvenl ~, that the le t:tba ri\'e r is gener-ill y becoming more allu viated ( bi rkhe:ld e l al. 19(5). with decreasi ng now volumes and inc re:ls ing sediment inputs the prog ress io n of e h.mge in the different c han nel types is li kel y to be from mi xed 1>001mpid or mi xed :lnasto mos ing 10 allu vi al s in· glc thread or .. l1ll vial bmidcd ( herit .. gc ('/ lil. 19(9 ). c onclu s ioll despitc th e lc taba ri vcr ha vin g incised into bed roc k in the pa sl. allu vial c hannel t)' pe s domina lc ihe rive r. th e in n uc ncc o f the underlyi ng bedrock has bee n l o~ t du e to the depos it ion o f un co nsolid:ned and con so lidated all uv ial mat e rial. bed roc kinnu e nced ch .. nncl types arc resuic led to shon le ng ths of c ha nnel .. nd arc oft e n as~oc i a t ed wi th ani fieia l stmclures that alter the sedim e nt ba l:mce loca ll y. la nd deg radat io n within the catchmc nt alld assoc i:ltcd red uc lio ns in now reach ing the do wnstream c nd of ihe rive r. ha ve undoubted ly hel ped 10 mai nt a in the ri ver in its present stat e ( birkhe.1l1 ('i (ii. 1(95). rat es of ch:tnnel c h.rn ge arc slo wer .md th e l1ia£ nitud e of geo mo rpholog ical adj us tm e nt is mino r in compa ri~on wi lh syste ms such as the sabie ri ve r (va n nie kerk & i-i e riw ge 199 3: i·lcrita ge ('/ (ii. 1997: heritage & moon 2000). the relati ve slabi lil)' of the le taba is probabl y rcl ;li ed to the declin e in o ve mll n o w e ne rgy con sc(lil e nt upon th e mu ch reduced now freque ncy .md ill :lgnitude as a result of int e ns ive wal c r abstmction wi lhi n the catc hme nt (ste ffen ro bert s & kirste n i 99();i) it is anti cipated that c han ge wilhin ihe syste m will now occ ur o nl y in re latio n to s ignific:l nl e \'c nts th at n ood ih e mac ro-ch.mn e!. the in fe rred con se(llle nce of s uch nood s will be the de posi lion of fine se dime nt at the base o f the macro·c hann e l bank and the rewo rk ing o f un conso lidated s.mds and fine growe l ove r th e surf;lce o f e xt e ns ive sand shee ts. s tabil isati o n o f some :ireas. p.1nic ularly the ba nk s o f ih e lo wnow di stribut a ry c ha nnel s lli ay funh er inhib it chan nel c h:mge, and the inferred con seque nce is the de vel o pme nt o f stable. o\'e r-dl..-cp ch .. nnel ne two rks in la rgescale deposi ts o f unconso lid ated allu vium . obse rvation of the tre nd s on th e lelab:l ri ve r suggesl that de ve lopmc nt of th e te rracelike cohes ive de posi ls <ii the s ides of the macro·chann el is li kel y to co ntinu e b), ven ical acc re tion provi ded nond fl ows o f suffi ci e nt ma gni tude colllinu c to reac h the knl ger nati o na l park . similar nuws :ire rctlu ire<l 10 maintain the sand shee ts, it is infe rred tha t the se remai n free o f fine sedime nt as a re sult o f prolon ged nood recess io ll . !\ rl..'<iuc tio n in the dura ti o n of inte rmediate wanin g nows may lead to th e de pos itio n of fine suspe nded mate ria l across the surface of ihc sand s hee ts in stea d o f this mate ri:.l con tinuin g to be transpon cd do wn strea m. t he lack of pmlorl£ed flo ws cap .. ble of re lilo bili sing fine mat erial. du e 10 inc reas in g wat er usage within the catc hment . will further e xacerb:ne the poss ibilit y of cohes i\'e st.-d iment build· up. ack nowled ge mell ts the a uthor.. gr.lldulj)' aei",o\'. led gc lhe fun d ing proi'idcd fo r ih i~ projcci by ihc walcr re~arch corn· rni ... ~ion o f soul h a rric a. logi'lic;,1 -'111111011 \'.·a~ provided by ihe n:u ioll al i' arks board of soulh africa. p<lrt ic il1 ~r ih~,,' ks 111 u,1 be cxiclldcd iii m r bcl1)., us erl~jill :1nd /i.'tr mitt iun xho~~, for ihei r hc tp in ihc fic ld. 1\·l r g. 1)obn), nski is ih ;111kcd for h i~ c~," (j· gr.'phic ~k i ll s. anorly 11111 11 ~ rerci'ces ~ ,re ilmn kcd fo r thc ma ny enn ~i iljc li \'e comme nt , in ihei r i'cl'iews. rcfercncl..'s 1~ir kht:ad. 1\ .1.. & g.lll o m i\cl 1995. t<."chniquc .. for c\' alualing ri'·c r :tnd i'c-cn ·oi r !>cd irncnl~llion 011 ihe sabic a nll l..ciao:l rh·cr ~ysl cms. l'nlc .. • (fllft's of iht' m,·" " ,/r smllil ,v,;("/i<i nli/;mllll issn 0075·6458 '(h/m/og.\' s.i'iii{i<jsiuiii (e/ummicfllrtlml), s",,· /c",/}cr /995, cre.'''''''s/o ..... , s",,/it ,vri ... /, c!testtllu,. 1'. 1994, geology and geornorl'hol,lsy of ihe sahie ril'er, kruger nalionji pl.rk. ;md it~ el.lcllillenl area. cemre for wjter in the environ· ment, imeml.1 ~p()rt: no, ii9·t univenily of the wilwl'lersrjnd. johannesburg. iiertt,igi ,. g. 1... m . e. charl.ton & s. o'ru;an (ill '''''.1.<). morpho logical ela~~ifiemilln of flllvia! environmenls: an iui'estigalinn nflhe emllinuurn uf channell}'pe,. jullmll/ 'ifc"o/"i"i'. 1·iel!rr,\ge. g.l. & by. ]l.i oon, 20(xl the coillellll'omry geo llhlrphology of ihe sabie ri ver in the krusei' nalional park. koe,i",-, 43 : 39-55 . ii eritage. g.l.. ,\,w. v,is niekerk & b.i'. moon. 1999, cll.~~ifying bedrock influcnced ..:mi-;irill river ~y~l elll~: extending ihe conti rllluui l·oneepl. pp. 53-so. "r: miu.ei!. aj. & a. cu'!',\ (eoj~ ) . 11i';l'iif.i' 'if i:""'io/ "-onll. ch iche~ter: wiley ii ei!itagi:. g.l.. a.w. van nieki:rk. h.i'. /l ioon. l.j. b ro,\ijil urst. k, h , rqgi:i!s & c.s. j a~ils. 11)')7. 71/1' k('(jm(jfijlw/ogic<l/ "'.'i'",'.i'f' /0 dlllllkillg flo\\' rfgilll/',i' (jfl/ii" sabi" <i",i i..<'/ll/m d.".,. ,1'.1',1' 1,'m,i', preloria: watcr resean:h c()l1l1l1i~,ion. (w:lter re search collllllis,ioll report: nn. 3761 11')7 .) ii ci!itage. g, l, a. l. birkilic,\d. l. j. bito,\llnuiist & b. r, i iait."n. 1999. the inl1uence of 110oding on lhe erodabilit)' of cobe,ive m:dimem, allllls lhe s;lbie r iver. south africa. s,'di,ii/'rrwl0,l:y28.131 . 145, k .... iglrron. a.d. 1997. flrll'i,,/ ftmm /i1111,,,,w,'u,',.: /i nfli' lj('f,fpl'clil'l'. london: edward ,\moll!. row","j1(t:e. k. m . &. r. a wa[)"-.'ion. 199(,. tr:iii~' blillg dlanncl geomorphology into i1yttr:urlie h~lbitm : applil';ltion of ihe hydr:lulk hiotope co n· cepl to :lil asse.,sillc/ll nf disch~rge related habiillt dwnses. i' p. 342-351. i,,: lecu:rc, 1\'1.. ii. capita. s, v"uxtl:<-:. a. !j ouditeaul 'r & y. cott:, (cds,), lii:""ydrauli'lfi<' zooo: i'm/'/'/"ifissn 0075 ·(,458 " ill,~s of 11/1' ~/'('ollli it\/fr illl,'m(l/i",,,,/ j."mp"· si"111 ,iii /tydmu/io lii,,1 i",bilo'." quebec. canl.da: ia fir . rou:-.-rltu .. m. w .. k. ii , rogus & g. l h crltagt, (ill 1m', ... ). llllld!ocl' pc 't~le change in ihe -emi· arid s;,bic rilei'. kruger n~tion31 park. in rc~p(m'e 10 11(1)0.1. droughl and fire. south african gcol:l'llfllril',,, jmmw/. sti:ffi: n r oiierts01'o' & klksti: 1'o' . 1990". \1~ii<'1' ll'iliilin' f!{{//mill,~ ,if ilw /..<'w/", hin'r blliill. miu/v of d<'l'e/iiiii/wili {11i/"l1/it'/ 11",1 ""111(1):1'111('11/ of/it<' 'riil,', /l'.fomr/·.' (if.l'dm/ii,~.": fli"" c"i/("mioll). preloria: dejl:,rtmenl of waler affair,. ( dep;lrlrn cnt of waler affair,' report: no. bsooioo 13'jo) s-n.i'i'i;_" roiii.rtso" & k iitstl'. 1990b_ \i~lil'r "'-,""i(fn ' /i/mlllilll: ,if iiil' /..i.'/(lfh' hi"cr ill/s ill. sii/(~\' of dfi·t'/"lljill'ill/hi/,·"lil/l (/11(/ mmlllgfii//'''' of /i,t' ,,·(!/t'r rfs",,,,,,,'i (exccutil'e suiiul);,,),). pretoria: depllrtl11cl11 of w:l1er aff;lirs. (department uf waler affair~ report: no, i' bsoofooioi'jo.) v,\n niu';erk. /\ .w. & g,t. ih]oin~\c,l 1993. c"i!mmplrolrwy 'if ii", slibi,' hi,./"(: """n';",,· mul .-j('~'sific/llion. johai\ne~h lt rg: university nf the witw~len.r;ii'd, (c(1i1re for water in the environmenl reporl: iill , 2/1)3) vovt. i. 19'12 . shurhcrm geomorphologic;" eh~nges in lhe s;lhic and lelab" ri'e", in the kru~er nationll! i'arl , unpubli,llctl ,\ i. sc. lii,~el1~ti()n. dep;1i1mcnl tlf gcogr:l]lhy and enl iroml)erlla\ slodie,. ullin·.." il y tlf the wilwat~r· .'fand. johanne,burg, walllison. r, $; k, rowf',"l'iu]:, 1(1)5. a hierjrchkal geom0'l'llolugil-al ",,,.lei ftlr classification of snuth african river ,y'ic11i'. i'p. 49-68. iii : uvs. m.c . (cd.). ci(i."ii/il-lili"" of rh" 'f,i' iliid '."iil'irml· 111"11/(11 hl'lillh illdklll'jfj. i'rclori,,: w3ter re:-c;lrch connl1i,~iun, (water rc.,e3f('h comrni~~ ion reporl : no. tt6319-l ,) ko .. ,11jl' +111 (200!) abstract introduction study area methods results and discussion acknowledgements references about the author(s) hendrik sithole scientific services, south african national parks, hadison park, kimberley, south africa school of animal, plant and environmental sciences, university of the witwatersrand, johannesburg, south africa nolubabalo tantsi department of agriculture and rural development, provincial government of south africa, johannesburg, south africa hugo bezuidenhout scientific services, south african national parks, hadison park, kimberley, south africa applied behavioural ecology and ecosystem research unit, university of south africa, pretoria, south africa thinandavha c. munyai school of life science, university of kwazulu-natal, durban, south africa lufuno munyai scientific services, south african national parks, hadison park, kimberley, south africa citation sithole, h., tantsi, n., bezuidenhout, h., munyai t.c. & munyai, l., 2020, ‘reconnaissance of epigeal ants at the degraded and control sites of mountain zebra and mokala national parks’, koedoe 62(1), a1542. https://doi.org/10.4102/koedoe.v62i1.1542 short communication reconnaissance of epigeal ants at the degraded and control sites of mountain zebra and mokala national parks hendrik sithole, nolubabalo tantsi, hugo bezuidenhout, thinandavha c. munyai, lufuno munyai received: 03 may 2018; accepted: 08 mar. 2020; published: 25 may 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract south african national parks (sanparks) is implementing rehabilitation projects in parks where acquired lands have degrees of degradation. such parks need to have empirical data to determine the degree of degradation and the success of the projects. we sampled epigeal ants at degraded and control sites in mountain zebra and mokala national parks to acquire the empirical data. with the data, our main aims were to: 1) determine the impact of degradation on the ant community; 2) determine if the rehabilitation period influences the difference between the degraded and control sites; and, 3) identify the degradation that affected the ant community the most. the majority of the degraded sites had lower abundance, species richness, shannon-wiener diversity, commonand exclusive-species than their respective control sites. all the degraded sites had a lower abundance of ants compared to their respective controls. the majority of the degraded sites had lower species richness, higher shannon-weiner diversity index and higher ant species dominance than their respective controls. the rehabilitation periods differently influenced the difference between the degraded and control sites. degraded sites that had undergone longer rehabilitation times had lower dissimilarity to their respective controls, while a degraded site with shorter rehabilitation period was very different to its control. ploughing appears to have longer lasting degradation-impact on ant communities than herbicides applications. a site with ploughing degradation was very dissimilar to its control while sites with chemical degradation had lower disparities to their respective controls. different degradation activities, therefor, affect ant recovery differently. conservation implications: attention should be paid to the type of degradations that are at the lands identified for expanding the conservation areas. some of these degradations seem persistent and slow to rehabilitate. variety of organisms (such as ants in this case) should be included when determining the degradation status of a conservation land. keywords: disparity; indicator species; rehabilitation age; degradation method; conservation; ecology. introduction habitat loss is the decline of ecological (or environmental) area (or community) by any action resulting in unfavourable conditions for species to complete their lifecycles (galvin 2007). habitat loss is one of the main drivers for the decline of especially endemic species in south africa as well as globally (international union for conservation of nature (iucn) 2015). to preserve the indigenous habitats that are within its jurisdiction, the south african national parks (sanparks) is implementing rehabilitation projects where varying degrees and types of habitat degradation occur (south african national parks (sanparks) 2016). a reconnaissance was carried out in two sanparks, mountain zebra (mznp) and mokala (monp), to compare ant diversity and abundance in degraded and undisturbed control areas. over-grazing, ploughing and eradication of alien species lead to degradation in mznp, whilst chemical treatment to control woody plant species was the main factor in monp. we chose to use ants as they often respond temporally and spatially to degradation (gollan et al. 2011). they exhibit relatively low mobility or dispersion within their habitats (barrow & parr 2008), and interact with different organisms at different trophic and symbiotic levels (agosti, schultz & majer 2000; hölldobler & wilson 1990). sampling ants is also relatively cheap (mcgeoch et al. 2011). the main aims for this study are as follows: (1) to determine whether degradation affects the ant communities; this was conducted by comparing ant composition diversity between degraded and control sites of a specific locality. (2) to determine whether rehabilitation periods influence ant communities by comparing the specific biodiversity index disparity between the localities with varying rehabilitation periods. (3) to identify which of these degradation activities impacted most on ant communities by comparing the specific biodiversity index disparity between the localities. study area the mznp (32.20500° s, 25.45000° e) is situated in the eastern cape province of south africa (figure 1). its vegetation consists of a mixture of dwarf shrubs and grasses (bezuidenhout & brown 2008). the area receives late summer rain, with the highest rainfall recorded in march and the lowest in june. average temperature ranges from 0.5 °c to 28.4 °c (bezuidenhout & brown 2008). three localities were selected in this park. the first locality was groenfontein with the degraded site, at 32.215628° s, 25.119439° e, consisting of bare soil surface and debris left after clearance of the alien plant species populus canescens. the control site for this was at groenfontein (32.21563° s, 25.11844° e), which had no history of any degradation. its proximity to the degraded site and similar habitat makes it a valid control site. the second locality was juriesdam. this degraded site, at 32.18846° s, 25.49268° e, suffered from sheep overgrazing with its undisturbed control site situated at 32.19030° s, 25.40922° e on a similar habitat (brown & bezuidenhout 2018; van der walt 1980). the third locality was ubejani. the degraded site, at 32.16859° s, 25.47663° e, was formally a ploughed field, and the undisturbed control site was situated at 32.17062° s, 25.46189° e. the proximity and habitat similarity between these sites make the site a valid control site. figure 1: the map showing the localities where ants were sampled from both degraded and control sites in mountain zebra and mokala national parks. the monp (29.10350° s, 24.39410° e) is located in the northern cape province of south africa. the dominant vegetation types are kimberley thornveld, vaalbos rocky shrubland and northern upper karoo (bezuidenhout et al. 2015a) (figure 1). the area receives late summer rain with the highest rainfall recorded in march and the lowest in june. the temperature ranges from −6.61 °c to 39.95 °c (bezuidenhout et al. 2015a). the two localities that were selected were windpomp and puntberg (figure 1), where bush densification of senegalia mellifera (swarthaak) was treated chemically (bezuidenhout, kraaij & baard 2015b). the degraded site of windpomp was located at 29.11187° s, 24.37244° e and the control site was at 29.11069° s, 24.37556° e. the degraded site at puntberg was situated at 29.13396° s, 24.31461° e with the control site at 29.14880° s, 24.32222° e. methods twenty pitfall traps (100 ml plastic cups), each containing 50 ml of 50% propylene glycol–water solution (adis 1979), were placed at the centre of each site in a grid of 5 × 4 rows (each 10 m apart) for 96 hours. ants were sampled in november 2015 at the mznp sites, and in february 2016 for the monp sites. only female workers were then identified to genus level using the keys of bolton (1994). the morphospecies of the identified specimens from each genus were then given a code based on their external morphological characteristics (viz. body sculptures, body colouring, hair/setae colour, hair/setae shape and petiole shape). all samples were kept at scientific services, kimberley, sanparks, for the future reference. the biodiversity indices used to determine the composition of ant communities from each site were abundance, species richness, dominant species (i.e. the most abundant) and similarity (anderson et al. 2008; dandan & zhiwei 2007; spellerberg & fedor 2003). sampled ant specimens and the morphospecies were counted to get the ant abundance and ant species richness for each site. the total abundance of all ant species for each site and the relative abundance of each ant species were calculated. the relative abundance of all ant species was ranked in decreasing order to get species that contributed ≥ 75% of the total abundance of each site. the shannon–wiener index and simpson’s diversity index were used to calculate the ant diversity per site (spellerberg & fedor 2003). sorenson’s coefficient (dandan & zhiwei 2007) was used to determine ant community similarities between the sites of each locality. the difference between the degraded and control sites of each locality, for each biodiversity index, was compared with its counterparts from other localities. the differences were given a score ranging from 1 (for the least difference) to 5 (for the highest difference) (roux et al. 2002) for the following indices: abundance, species richness, shannon–wiener diversity index and simpson’s diversity index. the scores for the dominant species were given 0 (where at least three of the five most abundant species at the degraded and control sites were of the same species – regardless of the order of their abundance at top five) or 1 (where at least three of the five most abundant species were different between the degraded and the control sites – regardless of the order of their abundance at top five). the scores for sorenson’s coefficient index ranged from 1 (for a locality with the highest sorenson’s coefficient) to 5 (for the lowest sorenson’s coefficient). all the scores for each locality were then summed up and compared with other localities. ethical considerations this article followed all ethical standards for a research without direct contact with human or animal subjects. results and discussion all the degraded sites had lower abundance values than their controls (figure 2), with the largest difference at groenfontein and the least difference at juriesdam (figure 2, table 1). figure 2: diversity indices of ants sampled on degraded and control sites from localities at the mountain zebra national park and mokala national park. table 1: index values from the degraded and control sites of each locality and their disparity rankings relative to other localities. species richness was lower at the degraded sites of groenfontein, ubejani and puntberg relative to their controls (figure 2) but was higher at the degraded sites of juriesdam. the species richness was similar at the degraded and control sites of windpomp. the degraded and control sites of the ubejani locality had the most dissimilar species richness, whilst the degraded and control sites of the windpomp locality were least dissimilar (figure 2, table 1). the shannon–wiener diversity index indicated that ant diversity was higher at the degraded sites of groenfontein, juriesdam, ubejani and windpomp, and lowest at puntberg (figure 2). the largest shannon–wiener diversity index difference was at ubejani and the least difference was at juriesdam (figure 2, table 1). the simpson’s index showed that groenfontein, ubejani and windpomp had higher dominance at the degraded sites, whilst the degraded sites of juriesdam and puntberg had lower dominance in relation to their control sites (figure 2). the ubejani sites had the highest simpson’s index difference, whilst puntberg sites had the least simpson’s index difference (figure 2, table 1). although the degraded and control sites of most localities had similar dominating ant species (except juriesdam and windpomp) (table 1), most of the ant species contributing ≥75% of the total abundance of the locality were less abundant (and in fewer traps) at the degraded sites (table 2). table 2: species list of ants sampled from each locality with their abundance and indications stating their roles in dissimilarity contributions and exclusivity between the degraded and control sites. the low abundance and species richness at the degraded sites suggest that the treatments applied in the habitats were not supportive to ant communities (table 1). ant sampling should be included in the methodology used to assess achievement of the aims of rehabilitation interventions. these data and the sampling methodology can also be used in monitoring projects to assess the temporal trends of ant communities to assess rehabilitation actions. ants are dominant in the terrestrial ecosystems based on their biomass, interactions with other organisms, their roles in the ecosystem and their response to disturbances, as suggested by this study. we suggest that the lower biodiversity indices from the degraded sites (figure 2) are driven by the disturbance (or even the diminution) of microhabitats. these disturbances may have altered the effects of driving factors (such as temperature and humidity) to levels beyond the tolerance of some species (gibb et al. 2015; hölldobler & wilson 1990). these changes may have favoured species that are able to adapt to harsher conditions, for instance, ocymyrmex species (hölldobler & wilson 1990; marsh 1985; sommer et al. 2013). ocymyrmex sp. 1 was more abundant at degraded sites of all the localities, except juriesdam, where it was not recorded (table 2). the abundance of ocymyrmex species, where it occurs, could further be used as a general indicator that determines the progress of rehabilitation actions (table 2). this could be performed by comparing the abundance disparities of ocymyrmex sp. 1 at a degraded site relative to its control, or by temporally comparing its abundance in a specific degraded site to determine the trend followed by abundance. the lower ant abundance at all the degraded sites, and the lower species richness at most of the degraded sites (figure 2), could have affected the important myrmecophile roles at these sites as there could be relatively fewer ants to fulfil their roles in ecosystems in both intraspecific and interspecific symbioses such as competition, mutualism, parasitism and feeding behaviours (hölldobler & wilson 1990). therefore, this could have affected organisms that rely on myrmecophile relationships (positive interactions between ants and other organisms) (hölldobler & wilson 1990). differences between degraded and control sites suggest that time plays an important role in the rehabilitation process. sites that had been recently rehabilitated, such as groenfontein, had the highest overall differences, whilst the ones that had been rehabilitated before, such as windpomp and puntberg, had the lowest overall (table 1) differences. the narrower disparities at puntberg and windpomp could be attributed to improved habitat because of longer recovery time (table 1). this follows data findings of andersen, hoffmann and somes (2003), who also found ants recover better with increasing rehabilitation time. we therefore anticipate that in the future the differences in younger rehabilitated localities would decrease. in addition to the time of rehabilitation, the type of degradation also influences the recovery (vasconcelos 1999). ubejani site has the longest rehabilitation time but had the second highest overall differences between its sites (table 1). we relate this difference to the ploughing that took place at the degraded site. chan (2001) reported that ploughing disturbs the soil profile and its associated organisms. this again could have had an impact on the ant community by affecting their myrmecophily association with the soil, other arthropods and plant species (agosti et al. 2000; hölldobler & wilson 1990). we anticipate there will be a slower narrowing of differences at ubejani relative to other localities because of the type of degradation it experienced. the type of degradation can sometimes have no direct impact on ants, as was seen at puntberg, juriesdam and windpomp which have the least disparities between the sites and controls (see overall score in table 1). the degraded sites at puntberg and windpomp experienced chemical degradations, mostly with tebuthiuron as the active ingredient (bezuidenhout et al. 2015b) to eradicate woody plant species. as this chemical is systemic and absorbed through the roots (blaise & harwood 1991), it may not have been in direct contact with epigeal ants. the chemical may, therefore, had little impact on the ants. we are unsure whether the difference in these localities would remain relatively low because johnsen and morto (1989) reported that the concentration of tebuthiuron and its depth in the soil changes under different rainfall conditions. it may be possible to show this by samplings during different rainfall conditions. because this is only a preliminary study conducted during local drought conditions, we recommend further sampling when the rainfall is back to previous averages. we hope also that additional samplings could verify whether the differences still hold for all different sampled degradations. in conclusion, we suggest that environmental authorities should consider the importance of the degradation types when acquiring lands for expanding conservation areas with consideration for brother organisms. as some of these degradation types seem to be persistent and take longer and more effort to be rehabilitated whilst others need lesser effort. they should also consider including different organisms when determining the degradation condition of the concerned conservation land. including different organisms when assessing such land would inform the management as to how degraded is that land, in general, and therefore make the decision based on the assessment that would include different avenues. acknowledgements sanparks and biodiversity social projects in sanparks provided funding and manpower for this project. the following people contributed immensely towards collecting data for this article: clodene ackerman, pulane daemane, letlhogonolo kotsokoane, abel matsapula, phindile mbele, kelebogile mohudi, spencely motloung, mazwi mthethwa, shadi ndouvhada, lucky ramabulana, bruce taplin and nosidima vumindaba. laboratory space to identify ant specimen was provided by the arid lands node of the south african environmental observation network (saeon) in kimberley. the authors would like to thank the referees who reviewed the article. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions h.s. was the project leader and responsible for sampling design, project implementation, data analysis and article writing. n.t. was responsible for project implementation, data analyses and article writing. h.b. and t.c.m. were responsible for article writing. l.m. was responsible for sampling. funding information sanparks and biodiversity social projects in sanparks provided funding and manpower for this project. data availability statement data sharing is not applicable to this article as no new data were created or analysed in this study. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references adis, j., 1979, ‘problems of interpreting arthropod sampling with pitfall traps’, zoologischer anzeiger 202, 117–184. agosti, d., schultz, t. & majer, j.d. (eds.), 2000, ants, smithsonian institution press, washington, dc. andersen, a.n., hoffmann, b.d. & somes, j., 2003, ‘ants as indicators of minesite restoration: community recovery at one of eight rehabilitation sites in central queensland’, ecological management & restoration 4(s1), s12–s19. https://doi.org/10.1046/j.1442-8903.4.s.2.x anderson, m., gorley, r. & clarke, k.p., 2008, permanova+ for primer: guide to software and statistical methods, primer-e, plymouth. barrow, l. & parr, c.l., 2008, ‘a preliminary investigation of temporal patterns in semi-arid ant communities: variation with habitat type’, austral ecology 33(5), 653–662. https://doi.org/10.1111/j.1442-9993.2008.01832.x bezuidenhout, h. & brown, l.r., 2008, ‘vegetation description of the doornhoek section of the mountain zebra national park (mznp), south africa’, koedoe 50(1), 82–92. https://doi.org/10.4102/koedoe.v50i1.142 bezuidenhout, h., bradshaw, p.l., bradshaw, m. & zietsman, p.c., 2015a, ‘landscape units of mokala national park, northern cape province, south africa’, navorsinge van die nasionale museum bloemfontein 31, 1–27. bezuidenhout, h., kraaij, t. & baard, j., 2015b, ‘persistent effects of chemicals used to control shrub densification in semi-arid savanna’, earth science research 4(1), 31–39. https://doi.org/10.5539/esr.v4n1p31 blaise, c. & harwood, m., 1991, ‘ecotoxicological assessment of tebuthiuron – a substituted urea class herbicide’, revue des sciences de l’eau/journal of water science 4(1), 121–134. https://doi.org/10.7202/705093ar bolton, b., 1994, identification guide to the ant genera of the world, harvard university press, cambridge. brown, l.r. & bezuidenhout, h., 2018, ‘ecosystem description and diversity of the jurisdam-seekoegat sections of the mountain zebra national park, south africa’, south african journal of botany 118, 166–178. https://doi.org/10.1016/j.sajb.2018.07.013 chan, k.y., 2001, ‘an overview of some tillage impacts on earthworm population abundance and diversity – implications for functioning in soils’, soil and tillage research 57(4), 179–191. https://doi.org/10.1016/s0167-1987(00)00173-2 dandan, z. & zhiwei, z., 2007, ‘biodiversity of arbuscular mycorrhizal fungi in the hot-dry valley of the jinsha river, southwest china’, applied soil ecology 37(1), 118–128. https://doi.org/10.1016/j.apsoil.2007.06.003 galvin, k.a., 2007, fragmentation in semi-arid and arid landscapes: consequences for human and natural systems, springer, dordrecht. gibb, h., sanders, n.j., dunn, r.r., watson, s., photakis, m., abril, s. et al., 2015, ‘climate mediates the effects of disturbance on ant assemblage structure’, proceedings of the royal society b 282(1808), 20150418. https://doi.org/10.1098/rspb.2015.0418 gollan, j.r, de bruyn, l.l., reid, n., smith, d. & wilkie, l., 2011, ‘can ants be used as ecological indicators of restoration progress in dynamic environments? a case study in a revegetated riparian zone’, ecological indicator 11(6), 1517–1525. https://doi.org/10.1016/j.ecolind.2009.09.007 hölldobler, b. & wilson, e.o., 1990, the ants, springer-verlag, berlin. international union for conservation of nature (iucn), 2015, the iucn red list of threatened species, version 2015.1, viewed october 2016, from http://www.iucnredlist.org. johnsen, t.n. & morton, h.l., 1989, ‘tebuthiuron persistence and distribution in some semiarid soils’, journal of environmental quality 18(4), 433–438. https://doi.org/10.2134/jeq1989.00472425001800040006x marsh, a.c., 1985, ‘microclimatic factors influencing foraging patterns and success of the thermophilic desert ant’, ocymyrmex barbiger. insectes sociaux 32(3), 286–296. https://doi.org/10.1007/bf02224917 mcgeoch, m.a., sithole, h., samways, m.j., simaika, j.p., pryke, j.s., picker, m. et al., 2011, ‘conservation and monitoring of invertebrates in terrestrial protected areas’, koedoe 53(2), 137–149. https://doi.org/10.4102/koedoe.v53i2.1000 roux, d., de moor, f., cambray, j. & barber-james, h., 2002, ‘use of landscape-level river signatures in conservation planning: a south african case study’, conservation ecology 6(2), 6, viewed october 2016, from http://www.consecol.org/vol6/iss2/art6. sommer, s., weibel, d., blaser, n., furrer, a., wenzler, n.e., rossler, w. et al., 2013, ‘group recruitment in a thermophilic desert ant, ocymyrmex robustior’, journal of comparative physiology a – neuroethology sensory neural and behavioral physiology 199(8), 711–722. https://doi.org/10.1007/s00359-013-0830-x south african national parks (sanparks), 2016, park management plan for the period 2016–2026, viewed october 2016, from https://www.sanparks.org/assets/docs/conservation/park_man/mznp-plan.pdf. spellerberg, i.f. & fedor, p.j., 2003, ‘a tribute to claude shannon (1916–2001) and a plea for more rigorous use of species richness, species diversity and the “shannon–wiener” index’, global ecology and biogeography 12(3), 177–179. https://doi.org/10.1046/j.1466-822x.2003.00015.x van der walt, p.t., 1980, ‘a phytosociological reconnaissance of the mountain zebra national park’, koedoe 23(1), 1–32. https://doi.org/10.4102/koedoe.v23i1.632 vasconcelos, h.l., 1999, ‘effects of forest disturbance on the structure of ground-foraging ant communities in central amazonia’, biodiversity & conservation 8(3), 407–418. https://doi.org/10.1023/a:1008891710230 abstract introduction materials and methods results discussion conclusion acknowledgements references about the author(s) martin k.s. smith rondevlei scientific services, south african national parks, south africa demi rodrigues school of natural resource management, nelson mandela university, south africa bianca currie sustainability research unit, nelson mandela university, south africa citation smith, m.k.s., rodrigues, d. & currie, b., 2018, ‘a survey of the ichthyofauna in the noetsie estuary, western cape province, south africa’, koedoe 60(1), a1513. https://doi.org/10.4102/koedoe.v60i1.1513 original research a survey of the ichthyofauna in the noetsie estuary, western cape province, south africa martin k.s. smith, demi rodrigues, bianca currie received: 18 dec. 2017; accepted: 11 apr. 2018; published: 16 july 2018 copyright: © 2018. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the fish assemblage in the noetsie estuary, a temporarily open and closed estuary on the southern coast of south africa, was sampled using multiple gears. a total of 12 species from 8 families were recorded. collectively, estuarine-dependent marine species dominated seine net catches numerically and in terms of biomass for both sampling seasons. estuarine round herring (gilchristella aestuaria) was numerically the dominant species in late summer, while juvenile mugilidae dominated catches in winter. size class distributions of various fish species indicate that the estuary both serves a nursery function for important euryhaline marine species and supports estuarine resident taxa. application of the estuarine fish community index indicates the ecological condition of the estuary to be ‘good’. this study contributes to the species list for the estuary while also reporting the presence of an alien invasive freshwater species, gambusia affinis. recommendations include the development of a management plan and the formalisation of an estuarine management committee. conservation implications: the noetsie estuary serves a nursery function for important euryhaline marine species, while supporting healthy populations of estuarine resident taxa. the presence of one alien invasive fish species is documented with potential implications for the conservation of biodiversity in the estuary. introduction south africa has approximately 300 estuaries along its coastline (whitfield 2000). collectively, they play an important role in promoting fish species richness in south africa (harrison 2003) and are important nursery areas for several species of marine fishes, many of which are exploited (james & harrison 2008). the ichthyofauna in estuaries along the southern coast of south africa is fairly well known (hall, whitfield & allanson 1987; james & harrison 2008; kok & whitfield 1986; olds et al. 2011; russell 1996; whitfield & kok 1992). in a review, james et al. (2007) showed that the fish fauna in southern coast estuaries are dominated by juvenile estuary-dependent marine species, with strong contributions by mugilidae and sparidae. native estuarine resident species are also abundant in most estuaries in this region (james & harrison 2008), but a number of smaller estuaries, including the noetsie, are data deficient (whitfield 2000). anthropogenic influences on estuarine environments can impact food resources, distribution, breeding, growth and survival of fish assemblages (whitfield & elliott 2002). despite the dynamic nature of fish assemblages within estuaries, fish communities have been used as indicators of estuary health (harrison & whitfield 2004) and can illustrate changes in the condition of estuarine environments (whitfield 1997). the estuarine fish community index (efci) is a multi-metric fish index that integrates structural and functional attributes of estuarine fish communities to provide a robust method for assessing the ecological condition of estuarine systems (harrison & whitfield 2004). understanding and being aware of changes in relative abundance and species composition is important for guiding and evaluating management actions (olds et al. 2016) as they can reflect the state of the estuary and impact of management interventions. here we assess the diversity, abundance and size structure of the fish community within the noetsie estuary and compare the current efci scores with a previous assessment (harrison & whitfield 2006b). materials and methods study site the noetsie estuary, situated just east of knysna, borders the garden route national park (figure 1) and falls within the warm-temperate bioregion (harrison 2003). classified as a temporarily open and closed system, the overall condition is considered excellent (whitfield 2000) and the ecological state is defined as largely natural with few modifications (dwaf 2008). however, based on the fish community, harrison and whitfield (2006b) rated the system as ‘poor’. the noetsie river has a total catchment of 38.8 km2 (nrio 1987) and is one of the few estuaries that receives most of its natural mean annual run-off (bornman & adams 2005). figure 1: map of the noetsie estuary in relation to the garden route national park and placement of sampling positions. physico-chemical properties during each survey, selected physico-chemical parameters including water temperature (°c), ph, salinity (‰) and dissolved oxygen (mg l−1) were measured at six sites situated up the estuary (figure 1). all readings were taken at the surface using a multi-parameter water analyser (ysi 550a dissolved oxygen; ysi model 60 handheld ph and temperature; ysi model 30 handheld salinity, conductivity and temperature instrument). ichthyofauna sampling was conducted in late summer (march) and winter (july) of 2015. fish were sampled in the main channel of the estuary using a 30 m beach seine net (30 m × 2 m × 15 mm multifilament bar mesh in the wings and 5 mm bar mesh in the purse) at five sites spaced longitudinally up the estuary (figure 1). dense stands of common reeds (phragmites australis) in the lower and middle reaches as well as steep sides and dense bush in the upper reaches limited beach seine net sites. one seine net pull was executed at each site on each sampling excursion. at each site, a scoop net (54 cm diameter hoop and 2 mm bar mesh) was also used to sample the shallows along roughly 15 m of shoreline (three samples each of 5 m). a multi-mesh multifilament gillnet (stretched mesh sizes: 35 mm, 45 mm, 57 mm, 73 mm, 93 mm, 118 mm and 150 mm) with each panel being 5 m long was deployed for 2 hours at two sites, one middle and one lower (figure 1). three double-ended fyke nets (10 mm stretched mesh, 5 m leader and 60 cm first hoop diameter) were set within the lower reaches of the estuary (figure 1) parallel to the shoreline. fyke nets were set at sunset in water approximately 1 m deep and retrieved the following day at sunrise. all fish caught were identified to the lowest possible taxonomic level and measured to the nearest millimetre fork length (fl) before being released. mullet (mugilidae) caught below 80 mm were only recorded at family level. data analysis diversity was calculated as the total number of species and the number of species sampled per sampling trip. total species composition, by number and mass, was calculated for each sampling period with the relative biomass contribution of each species calculated using masses derived from length–mass relationships presented in harrison (2001). where appropriate, species length-frequency histograms (20 mm size classes for lichia amia and 10 mm size classes for all other species) were generated for each sampling trip. species recorded were divided into the estuarine association categories described by whitfield (1994): freshwater species, estuarine resident species, estuarine-dependent marine species and marine species. the per cent contribution made by each category to the total ichthyofaunal assemblage of each sampling trip was calculated in terms of number of species, relative abundance and relative mass. estuarine fish community index the efci developed by harrison and whitfield (2004) comprises 14 metrics that represent four broad fish community attributes: species diversity and composition, species abundance, nursery function and trophic integrity (table 1). metric reference conditions applicable to the noetsie estuary were developed from table 5 presented in harrison and whitfield (2006a) and followed procedures set out in harrison and whitfield (2006b). each metric was assessed according to the extent of its deviation from the reference condition with thresholds and scores being based on harrison and whitfield (2006b) (table 1). the species assemblage was compared to the reference assemblage using the bray–curtis similarity measure based on presence or absence. relative (%) contribution of each species for each sampling period and reference assemblage was 4th root transformed prior to running the bray–curtis similarity measure. for both these analyses, mugilidae were included as a ‘species’ because of the large proportion of juvenile mullet that were not identified down to species level. the final efci score was calculated by summing the various scores. the biological condition of the estuary was designated a qualitative rating (very poor to very good) based on ranges described in harrison and whitfield (2006b). the efci was applied to the late summer and winter data sets, and final scores were compared to an assessment by harrison and whitfield (2006b). bray–curtis similarity analyses were performed using the plymouth routines in multivariate ecological research package (primer) (clarke & warwick 1994). table 1: metric scoring criteria for small closed estuaries in the warm-temperate biogeographic region. results physico-chemical surface water temperature ranged between 18.9 °c and 22.6 °c during late summer and 11.3 °c and 13.2 °c in winter with only a slight drop in temperature occurring in the upper reaches for both sampling periods. salinity was low throughout the estuary during late summer decreasing from 1.7‰ near the mouth to 0.2‰ at sites five and six. the estuary mouth was open during the winter sampling period, and a salinity gradient was present with the highest salinity (17.2‰) occurring near the mouth and the lowest (0.4‰) recorded at transect six. surface water ph was relatively constant throughout the estuary during late summer ranging between 7.59 near the mouth and 7.02 at site six. during winter, the ph was generally lower and ranged from 5.95 near the mouth up to 6.59 at transect five. oxygen levels in late summer showed a slight increase from transect one (4.65 mg l−1) to transect five (6.60 mg l−1) but varied between transects in winter with the lowest level (4.66 mg l−1) recorded at transect five and the highest (8.88 mg l−1) recorded in the lower reaches. ichthyofauna a total of 731 fish representing 12 species and 9 families were sampled. of these, one was catadromous (category v), two were estuarine residents (category i), eight were marine migrants (category ii) and one was a freshwater species (category iv) (table 2). table 2: relative abundance and biomass of fishes captured during surveys conducted in the noetsie estuary between late summer and winter 2015. seine net catches were dominated numerically by gilchristella aestuaria and mugilidae (unidentified mullet species) comprising 37.8% and 30.8% of total catch in late summer respectively. however, mugilidae dominated catches (31.6%) during winter followed by g. aestuaria (25.8%). euryhaline marine species (categories iia, iib and iic) contributed 54.3% and 56.1% of total numbers of fish caught during the late summer and winter sampling trips, respectively, and dominated catches in terms of biomass comprising 85.5% and 60.7% of the entire sampled biomass, respectively (table 2). although estuarine resident species (ib) were prominent in terms of numbers together comprising 43.3% and 28.4% of the total number of fish caught during late summer and winter, respectively, they only contributed 14.5% and 3.9% of the total biomass harvested. fyke net catches in late summer were dominated by rhabdosargus holubi (78.2%) both numerically and in terms of biomass (65.4%), while winter catches were dominated by monodactylus falciformis (table 2). one species, anguilla mossambica, sampled in late summer, was not caught in any other gear type. lichia amia was the only species caught within the gill nets (table 2) during late summer, while m. falciformis (69.6%) dominated gill net catches numerically during winter followed by liza richardsonii (17.4%). scoop netting only resulted in one species, gambusia affinis, being caught during both sampling periods with fewer and smaller individuals being caught in winter (table 2). length–frequency distributions the length-frequency distribution for g. aestuaria showed a high proportion of adult fish in the 50 mm – 70 mm (fl) size range during both late summer and winter with very few larger individuals or juveniles present during the late summer period (figure 2). lichia amia were only sampled during late summer with fish ranging in size from 440 mm fl to 515 mm fl. most mugilidae sampled during the two surveys were juveniles ranging in size from 11 mm to 71 mm fl (figure 2). a greater distribution in size frequency for mugillids was noted during late summer. lithognathus lithognathus showed a shift in size class distribution between late summer and winter with larger individuals being sampled in winter (figure 2). both r. holubi and m. falciformis showed a wider size frequency distribution in late summer compared to winter with the larger size classes absent during the second sampling period (figure 2). figure 2: length–frequency distribution of the more abundant species caught within the noetsie estuary. (a) rhabdosargus holubi, (b) monodactylus falciformis, (c) lithognathus lithognathus, (d) mugilidae, (e) lichia aestuaria and (f) gilchristella aestuaria. estuarine fish community index the total efci score for the late summer sampling period was 49, while the winter sampling scored 46. the final index scores of both correspond to a qualitative rating of good (scores fall between 46 and 62) (harrison & whitfield 2006b). discussion although temperature gradients along the length of small temporarily open and closed estuaries do not generally occur during their closed phase (perissinotto et al. 2004), a difference of 3.4 °c was recorded during the late summer survey. rainfall experienced just prior and during the sampling period lead to increased river inflow which is likely to have influenced both water temperatures in the upper sections and the lower salinity levels recorded throughout the estuary during late summer. bornman and adams (2005) indicate that limnetic conditions (0.1 ppt – 0.5 ppt) may prevail in the noetzie estuary through most of a closed phase. although the lowest ph was recorded at transect five, near one of the main tributaries, ph was generally higher during the late summer survey. this is unusual as freshwater, because of the humic acid leached from catchment vegetation (dwaf 1995), is generally more acidic than sea water. dissolved oxygen levels were generally lower than those reported by james and harrison (2008) (6.2 mg l−1 – 6.6 mg l−1). this survey recorded 12 species from 8 families, which is a substantial increase from the previous work by james and harrison (2008) during which 7 species from 4 families were recorded. additional species include l. amia, m. falciformis, a. mossambica and l. macrolepis, which are all species known to occur within southern cape estuarine systems (james & harrison 2008; olds et al. 2011; whitfield 1998). however, two species, mugil cephalus and myxus capensis, sampled by james and harrison (2008) were not sampled during this survey. in addition, one alien invasive species, gambusia affinis, was recorded for the first time within the system. the numerical dominance of an estuarine resident species, in this case g. aestuaria, is not surprising as this group generally comprise over 50% of catches (numbers) in warm-temperate temporarily open and closed estuaries (james et al. 2007). gilchristella aestuaria breed throughout the year completing their entire life cycle within the estuary (whitfield 1998), and although the majority of g. aestuaria sampled were mature individuals; a wide range of size classes were sampled indicating that breeding was occurring within the estuary during the closed phase. somewhat surprisingly, no atherina breviceps were sampled during this or previous surveys (james & harrison 2008). although there is an overlap in diet between the two species (whitfield 1998), g. aesturia has been shown to switch diets and feeding strategies dependent on water clarity and food resources (blaber, cyrus & whitfield 1981) which may provide a competitive edge in certain circumstances. atherina breviceps was among the most abundant species captured in adjacent estuaries both to the west (james & harrison 2008) and to the east of the noetsie (james & harrison 2010); however, although the frequency of occurrence within small closed estuaries in the warm-temperate region is quite high at 60%, the relative abundance is usually low at 4.98% (harrison & whitfield 2006a). determining the reasons for their omission from noetsie estuary requires further work but is probably because of environmental preferences rather than competition. estuarine fish communities, in particular the estuary associated marine component, depend heavily on mouth state, and for temporarily open and closed estuaries, the frequency, duration and timing of opening events are important (james et al. 2007; kok & whitfield 1986). the near-natural mean annual run-off (bornman & adams 2005) controls the mouth dynamics of the system, which allows the migration of fish into and out of the system. the size class frequency distribution of r. holubi, l. lithognathus, l. amia, m. falciformis and mugilidae within the noetsie estuary indicates that the estuary serves as both a viable nursery and a feeding area for juveniles. the absence of myxus capensis and mugil cephalus from the catches was unusual as these two species comprised 51.9% and 41.6% of the catch or 9.7% and 41.4% of the biomass, respectively, in a previous study (james & harrison 2008). identifying muligids to species level would have provided a better understanding of recruitment and usage of the noetsie estuary by this family while providing a more robust data set for the efci calculations. the improvement in the efci scores from 32 (harrison & whitfield 2006b) to 49 and 46 for the late summer and winter surveys, respectively, pushes the ecological condition category up from ‘poor’ to ‘good’. a significant correlation between efci scores and mean efci values for systems sampled multiple times suggests that the efci is reproducible (harrison & whitfield 2006b) despite variability in fish communities. however, the improvement in score for the noetsie is more than likely as a result of a more intensive sampling regime and the use of multiple gear types than an actual directional change in ecological condition. however, when utilising the efci on a single system over time, it would be beneficial to standardise sampling effort, sampling seasons and gear types to minimise potential biases. the introduction of alien species is seen as one of the leading causes of biodiversity loss in aquatic ecosystems (mack et al. 2000). occurring naturally within south-eastern north america (pyke 2008), g. affinis has been successfully introduced to most parts of the world (lloyd 1986). preferring sheltered, shallow and well-vegetated freshwater habitats (arthington & lloyd 1989), g. affinis are highly tolerant to a wide range of physico-chemical conditions (pyke 2008) and can occur in waters with temperatures ranging from 0 °c to 45 °c (cherry et al. 1976), salinities from 0 ‰ to 41 ‰ (hubbs 2000) and dissolved oxygen from 1 mg l−1 to 11 mg l−1 (odum & caldwell 1955). the physico-chemical parameters measured during this and previous surveys of the noetsie estuary (bornman & adams 2005; james & harrison 2008) were all within the g. affinis tolerance range. in freshwater environments, g. affinis has been shown to impact ecosystems through both predation on and competition with native biota (pyke 2008), but little is known about the impacts of g. affinis within estuarine environments. potential impacts could include predation on juvenile fish and, in particular, the eggs of estuarine resident taxa. work by sloterdijk et al. (2015) indicates that in southern cape estuaries, g. affinis populations seem to undergo a boom and bust scenario, with a rapid increase in abundance over spring and summer and a collapse during winter. our results suggest g. affinis are limited in distribution to littoral waters and occur in low numbers in the noetsie estuary. however, a more detailed study with more sample sites over a longer period would be needed to accurately describe the population characteristics of this invasive species. conclusion the noetsie estuary is important for estuarine resident species while serving as a viable nursery area for estuarine-associated marine species. the species list of fishes utilising the estuary has increased with an additional five indigenous species and one alien invasive freshwater species. the state of the estuary mouth is the single most important factor driving the ecology in the noetsie estuary (bornman & adams 2005), and in turn, mouth dynamics are largely determined by river inflow. changes in river flow will influence the relationship between open, semi-closed and closed mouth conditions. a decrease in the open and semi-closed phases would limit larval fish recruitment and adult movement out of the estuary, potentially having a negative influence on the associated fish community which should be avoided. the ecological condition of the noetsie estuary is rated as ‘good’ showing an improvement from a previous assessment of ‘poor’. to maintain the condition (or improve it), we recommend that an estuarine management plan be drafted which includes the monitoring and management of river inflows and the potential control or eradication measures needed for the recorded alien species. the plan should also include a sustainable participatory process for stakeholder involvement. we suggest that an estuarine management forum be established, which includes stakeholders influencing or are influenced by river inflows. future research should include assessing the change in physico-chemical characteristics and estuarine fish community between stable closed and open phases, as well as directed research assessing the distribution, abundance and ecological impacts of g. affinis throughout the catchment of the noetsie estuary. acknowledgements bheki maphanga, pierre mouskie, brian du preez and alison macallister are thanked for assisting with data collection. wendy dewberry assisted with logistics, provided background information on noetsie and helped catalyse this research. aubrey wynne-jones is thanked for covering running costs and providing accommodation. sanparks provided all sampling equipment, and part of the running costs was covered by the rondevlei marine protected area budget. the two anonymous reviewers are thanked for their positive and helpful comments to improve the manuscript. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions m.k.s.s. was responsible for the study conceptualisation, data collection, analyses, manuscript writing and editing. d.r. contributed to the study design, data collection, analyses and manuscript editing; and b.c. contributed to the study design and manuscript editing. references arthington, a.h. & lloyd, l.n., 1989, ‘introduced poeciliidae in australia and new zealand’, in g.k. meffe & f.f. snelson (eds.), evolution and ecology of livebearing fishes (poeciliidae), pp. 33–50, prentice hall, new york. blaber, s.j.m., cyrus, d.p. & whitfield, a.k., 1981, ‘the influence of zooplankton food resources on the morphology of the estuarine clupeid gilchristella aestuaria (gilchrist, 1914)’, environmental biology of fishes 6, 351–355. https://doi.org/10.1007/bf00005764 bornman, t.g. & adams, j.b., 2005, present state of the noetsie estuary, iecm research report no. 45, institute for environmental and coastal management, nelson mandela metropolitan university, nelson mandela bay, 43p. cherry, d.s., guthrie, r.k., rodgers, j.h., cairns, j. & dickson, k.l., 1976, ‘response of mosquito fish (g. affinis) to ash affluent and thermal stress’, transaction of the american fisheries society 105, 686–694. https://doi.org/10.1577/1548-8659(1976)105%3c686:romgat%3e2.0.co;2 clarke, k.r. & warwick, r.m., 1994, change in marine communities: an approach to statistical analysis and interpretation, 1st edn., plymouth marine laboratory, plymouth, uk, 144p. department of water affairs and forestry (dwaf), 1995, south african water quality guidelines for coastal marine waters. volume 1: natural environment, the government printer, pretoria. department of water affairs and forestry (dwaf), 2008, reserve determination studies for selected surface water, groundwater, estuaries and wetlands in the outeniqua (groot brak and other water resources, excluding wetlands) catchment: ecological water requirement study – estuarine rdm report, volume 1.3: assessment report rdm/k40 – k50/00/con/0307, vol. 1, department of water affairs and forestry, pretoria. hall, c.m., whitfield, a.k. & allanson, b.r., 1987, ‘recruitment, diversity and the influences of constrictions on the distribution of fishes in the wilderness lakes system, south africa’, south african journal of zoology 22, 163–168. https://doi.org/10.1080/02541858.1987.11448038 harrison, t.d., 2001, ‘length-weight relationships of fishes from south african estuaries’, journal of applied ichthyology 17, 46–48. https://doi.org/10.1046/j.1439-0426.2001.00277.x harrison, t.d., 2003, ‘biogeography and community structure of fishes in the southern african estuaries’, phd thesis, rhodes university. harrison, t.d. & whitfield, a.k., 2004, ‘a multi-metric fish index to assess the environmental condition of estuaries’, journal of fish biology 65, 683–710. https://doi.org/10.1111/j.0022-1112.2004.00477.x harrison, t.d. & whitfield, a.k., 2006a, ‘estuarine typology and the structuring of fish communities in south africa’, environmental biology of fishes 75, 269–293. https://doi.org/10.1007/s10641-006-0028-y harrison, t.d. & whitfield, a.k., 2006b, ‘application of a multimetric fish index to assess the environmental condition of south african estuaries’, estuaries and coasts 29, 1108–1120. https://doi.org/10.1007/bf02781813 hubbs, c., 2000, ‘survival of gambusia affinis in a hostile environment’, southwestern naturalist 45, 521–522. https://doi.org/10.2307/3672601 james, n.c., cowley, p.d., whitfield, a.k. & lamberth, s.j., 2007, ‘fish communities in temporarily open/closed estuaries from the warmand cool-temperate regions of south africa: a review’, reviews in fish biology and fisheries 17, 565–580. https://doi.org/10.1080/00359190809519216 james, n.c. & harrison, t.d., 2008, ‘a preliminary survey of the estuaries on the south coast of south africa, cape st blaize, mossel bay robberg peninsula, plettenberg bay, with particular reference to the fish fauna’, transactions of the royal society of south africa 63, 111–127. https://doi.org/10.1080/00359190809519216 james, n.c. & harrison, t.d., 2010, ‘a preliminary survey of the estuaries on the southeast coast of south africa, cape st francis – cape padrone, with particular reference to the fish fauna’, transactions of the royal society of south africa 65, 69–84. https://doi.org/10.1080/00359191003652116 kok, h.m. & whitfield, a.k., 1986, ‘the influence of open and closed mouth phases on the marine fish fauna of the swartvlei estuary’, south african journal of zoology 21, 309–315. https://doi.org/10.1080/02541858.1986.11448004 lloyd, l., 1986, ‘an alternative to insect control by “mosquito fish”, gambusia affinis’, arbovirus research in australia 1986, 156–163. mack, r.n., simberloff, d., lonsdale, w.m., evans, h., clout, m. & bazzaz, f.a., 2000, ‘biotic invasions: causes, epidemiology, global consequences, and control’, ecological applications 10, 689–710. https://doi.org/10.1890/1051-0761(2000)010[0689:bicegc]2.0.co;2 national research institute for oceanology (nrio), 1987, basic physical geography/hydro data for ‘estuaries’ of the southern cape (cms 0–50), nrio data report d8706, sediment dynamics division, coastal engineering and hydraulics, nrio, council for scientific and industrial research, stellenbosch, 18 p. odum, h.t. & caldwell, d.k., 1955, ‘fish respiration in the natural gradient of an anaerobic spring in florida’, copeia 1955, 104–106. https://doi.org/10.2307/1439312 olds, a.a., james, n.c., smith, m.k.s. & weyl, o.l.f., 2016, ‘fish communities of the wilderness lakes system in the southern cape, south africa’, koedoe 58(1), a1364. https://doi.org/10.4102/koedoe.v58i1.1364 olds, a.a., smith, m.k.s., weyl, o.l.f. & russell, i.a., 2011, ‘invasive alien freshwater fishes in the wilderness lakes system, a wetland of international importance in the western cape province, south africa’, african zoology 46, 179–184. https://doi.org/10.1080/15627020.2011.11407491 perissinotto, r., blair, a., connell, a., demetriades, n.t., forbes, a.t., harrison, t.d. et al., 2004, ‘literature review: ecology of south african temporarily open/closed estuaries: a review of current knowledge’, in j.b. adams (ed.), contributions to the information requirements for the implementation of resource directed measures for estuaries. volume 2. responses of the biological communities to flow variation and mouth state in two kwazulu-natal temporarily open/closed estuaries, water research commission report no. 1247/2/04, pp. 1–55, water research commission, pretoria. pyke, g., 2008, ‘plague minnow or mosquito fish? a review of the biology and impacts of introduced gambusia species’, annual review of ecology, evolution, and systematics 39, 171–191. https://doi.org/10.1146/annurev.ecolsys.39.110707.173451 russell, i.a., 1996, ‘fish abundance in the wilderness and swartvlei lake systems: changes relative to environmental factors’, south african journal of zoology 31, 1–9. https://doi.org/10.1080/02541858.1996.11448389 sloterdijk, h., james, n.c., smith, m.k.s., ekau, w. & weyl, o.l.f., 2015, ‘population dynamics and biology of an invasive population of mosquito fish gambusia affinis in a temperate estuarine lake system’, african zoology 50(1), 31–40. https://doi.org/10.1080/15627020.2015.1021169 whitfield, a.k., 1994, ‘an estuary-association classification for the fishes of southern africa’, south african journal of science 90, 411–417. whitfield, a.k., 1997, ‘fish conservation in south african estuaries’, aquatic conservation: marine and freshwater ecosystems 7, 1–11. https://doi.org/10.1002/(sici)1099-0755(199703)7:1%3c1::aid-aqc213%3e3.0.co;2-8 whitfield, a.k., 1998, biology and ecology of fishes in southern african estuaries, ichthyological monographs of the j.l.b. smith institute of ichthyology, no 2, grahamstown, p. 223. whitfield, a.k., 2000, available scientific information on individual southern african estuarine systems, water research commission report 577/3/00, water research commission, pretoria, 217p. whitfield, a.k. & elliott, m., 2002, ‘fishes as indicators of environmental and ecological changes within estuaries: a review of progress and some suggestions for the future’, journal of fish biology 61, 229–250. https://doi.org/10.1111/j.1095-8649.2002.tb01773.x whitfield, a.k. & kok, h.m., 1992, ‘recruitment of juvenile marine fishes into permanently open and seasonally open estuarine systems on the southern coast of south africa’, ichthyological bulletin 57, 1–15. filelist convert a pdf file! page 1 reilly.qxd precision of helicopter-based total-area counts of large ungulates in bushveld b.k. reilly reilly, b.k. 2002. precision of helicopter-based total-area counts of large ungulates in bushveld. koedoe 45(2): 77–83. pretoria. issn 0075-6458. precision in helicopter total-area counts in bushveld for a range of common species are evaluated. counts were conducted on properties ranging in size from 250 ha to 10 000 ha as part of experiments on precision and power. counts were conducted in three vegetation types: mopane veld, sourish mixed bushveld and arid bushveld. scant regard is given to precision as a factor in large herbivore monitoring, with more effort often devoted to accuracy. coefficients of variation varied from 3.2 % to 70.9 %. median values are generally acceptable for the establishment of trends for long-term monitoring, but might be inadequate for annual population size monitoring and evaluation. a regression of all species and vegetation types by the standard error showed a significant correlation. similar regressions were also developed for impala, kudu, warthog, and zebra. regressions by vegetation type for mopane veld, sourish mixed bushveld, and northwest arid bushveld were also significant. however, the high correlation for some vegetation types might be an artifact of the small sample sizes. keywords: census, game counting, helicopter counts, impala, kudu, zebra, wildebeest, giraffe, large herbivore monitoring. b.k. reilly, department of nature conservation, technikon pretoria, private bag x680, pretoria, 0001, republic of south africa. issn 0075-6458 77 koedoe 45/2 (2002) introduction precision of game counts is often neglected in the development of aerial large ungulate counting strategies. replicated counts provide opportunities to assess the precision of the total-area helicopter counts for a range of species and conservation areas. the use of aerial surveys as a tool in the monitoring of large ungulates can be traced back to 1935 (cahalane 1938). the use of aircraft, particularly helicopters, has grown consistently since the 1950s and today is almost universally applied in enumerating many ungulate species, for example, white tailed deer, oidocoileus virginianus (beasom et al. 1986), mule deer, oidocoileus hemionus (bartman et al. 1986) and elk, cervus elaphus in michigan (otten et al. 1993). furthermore, total counts have been applied to many species of african ungulates and results have been recorded by several authors. van lavieren & esser (1979) report results of counts of giant eland tragelaphus derbianus, roan antelope hippotragus equinus, defassa waterbuck kobus ellipsiprymnus defassa, bohor reedbuck redunca redunca and bubal hartebeest alcelaphus buselaphus in west africa and several species such as impala aepyceros melampus, topi damaliscus lunatus, sitatunga tragelaphus spekii, eland tragelaphus oryx and waterbuck kobus ellipsiprymnus in rwanda. total counts have been used in east africa (dasmann & mossman 1962; jolly 1969) and in south africa (hirst 1969; goodman 1977; melton 1978; collinson 1985; bothma et al. 1990; eiselen 1994; van hensbergen et al. 1996). total aerial counts have moved beyond application in protected areas and, particularly in the south african context, are now widely applied to monitor population size on commercial game ranches. for smaller propreilly.qxd 2005/12/09 10:11 page 77 erties it provides information in a short time and is less sensitive to habitat diversity and ungulate population sizes than many other sampling methods. in most bushveld applications visibility from the air is also generally better than from the ground. the helicopter surveys have, however, many detractors due often to failure to ensure that underlying assumptions have been met (caughley 1974; melton 1978). the requirement for experienced observers, poor performance caused by observer fatigue, difficulty in standardisation of methodology between (and within) counts, and high costs often mitigate against the technique (caughley 1979; norton-griffiths 1978; seber 1992). consequently, these methods are often applied by inexperienced staff who are unschooled in the underlying principles involved. the non-standardisation of approaches to game counts, in general, lengthens the causal chain of poor precision. yet, it remains the census technique of choice in south africa. this could be due to a misconception that number of animals counted equate to the actual population size, i.e., the technique is accurate. the accuracy of helicopter counts has been questioned (rice & harder 1977; beasom 1979; payne 1981; de young 1985; beasom et al. 1986; seber 1992; bothma et al.1990) and the effect of various factors, for example, height and speed, on accuracy is unclear. other investigations quantify observer bias (pennycuick & western 1972; norton-griffiths 1978; melton 1978; graham & bell 1989) but there are few references on precision and power of these applications. each survey produces a count of the population, which is some fraction, hopefully large, of the actual population. in practice, individual counts differ for a variety of reasons, including systematic and random errors. the extent to which repeated counts differ can be stated in terms of the precision of a count. if precision is high, then repeated counts would all have similar values. when repeated survey counts of a population are conducted, the results will vary because of a variety of factors. beasom (1979) reports variations in replicated helicopter counts in texas, ranging from 0.9 % to 32.3 %, while le resche & rausch (1974) report coefficients of variation ranging from 16 % to 41 % in fixed-wing counts of moose. they also report significantly lower cvs in some seasons (spring, p < 0.05). hirst (1969) recorded variability of 12.8 % to 32.2 % on strip counts of blesbok on rietvlei nature reserve. beasom (1979) indicates a significant negative correlation between cv and mean estimated deer population and area size. rice & harder (1977) comment on the high precision attained using helicopters for mark-recapture counts in ohio and reports a cv of 7 %. mccullough (1994), in documenting herd composition counts of black tailed deer, describes large variances, and consequently questioning the value of the data in decision making. this study analyses replicate count data from several typical bushveld total helicopter surveys in an effort to quantify precision in addition to power analyses results published by reilly(2000) and brockett (in press). study areas three study areas comprised 1) an 8500-ha block located in the loskop dam nature reserve situated in sourish mixed bushveld (acocks veldtype 19), (acocks 1988); 2) a number of conservation areas surrounding atherstone nature reserve (ranging in size from 500 ha to 2500 ha) situated in arid bushveld (acocks veldtype 18); and 3) two areas in mopane veld (acocks veldtype 15)—a 10000-ha block of letaba ranch in the limpopo province, and five conservation areas adjacent to the dongola nature reserve (ranging in size from 250 ha to 1500 ha) in limpopo province, also in mopane veld. all areas fall in the savanna biome. method data collection the method has been described in detail by reilly (2000). a summary of the methodology was as follows: strips (330 m wide) were flown by heading and counter heading, tracked by a navigator using aerial koedoe 45/2 (2002) 78 issn 0075-6458 reilly.qxd 2005/12/09 10:11 page 78 photos, 1:50 000 topocadastral maps and a grid overlay of 1 km². flying at 96 km/h, an attempt was made to cover the total area. height was regulated using a radar altimeter set at 53 m above ground level and counts were accomplished with the rear doors removed. all counts were done between july and september, post leaf drop. replicates were conducted on consecutive days but at the same time of day, and using the same observers. handheld tape recorders were used to record data and were later transcribed to data sheets using the grid numbers of the overlay. all ungulate species seen were counted and, where necessary, herds were flushed to ensure accurate enumeration. during each aerial survey on a property an attempt is made to completely enumerate the population of each species. data analysis precision is most commonly stated in terms of a variance or standard deviation, or may be scaled and presented in the form of a coefficient of variation. in this study, coefficients of variation are used to present information on relative precision since the absolute sizes of populations differ greatly, both between species and between sites. the coefficient of variation (cv) is a scaled measure of precision that provides a useful indication of a technique’s ability to show change over time. precision of the counts is expressed as a coefficient of variation (scaled measure of dispersion that allows for comparison between sites), i.e., where s is the sample standard deviation and x (x bar) the mean of the replicate counts. issn 0075-6458 79 koedoe 45/2 (2002) cv = (100)x s the median value of each set of cvs for each species provides a measure of location of the sample that is insensitive to outliers. bootstrapped standard deviation each survey can be considered as an independent sample from the population of all possible surveys for that site. to estimate the variance of these counts, the standard estimator of variance could be used but this is problematic since we need to know the estimate of variance for the statistical population of enumerable counts rather than sample parameters obtained from the three and four replicates. to overcome this problem the estimate of the variance was obtained by bootstrapping (efron 1974). samples (4000) were generated randomly with replacement from the sample values. from each of these samples, a variance was calculated. the mean of these variances is an estimate of the statistical population variance. other measures of dispersion (standard error) were then based on these estimators. correlations for species and vegetation type a regression analysis for all species and vegetation type (site) of the mean count (population) size by the standard error (se) was also conducted. for the purpose of discussion in the following review of results by species and area-habitat type the cv was used as a measure of precision. results estimates of precision derived from bootstrap replications are presented in table 1 as means, standard errors of the mean and as coefficients of variation. it can be seen that there is considerable variation in the estimates of cv, they range from < 5.3 % to > 70 % for some species. kudu ten estimates of precision were obtained for kudu ranging from 5.3 % to 54.2% with a median of 14.6 % (n = 10). single observations were obtained for lowveld mopane veld and the sourish mixed bushveld (29.6 % and 5.3 % respectively). the results for the northern mopane veld were poor with a high of 70.9 %, while a median for five observations in northwestern arid bushveld had a precision of 12.5 %. zebra six estimates of precision were obtained for zebra with cvs ranging from 4.9 % to 61.9 %. single observations for lowveld mopane veld, sourish mixed bushveld and northwestern arid bushveld were 18.6 %, 7.2 % and 11.3 %, respectively. the median of three observations for northern mopane veld is 9.8 %. reilly.qxd 2005/12/09 10:11 page 79 koedoe 45/2 (2002) 80 issn 0075-6458 table 1 summary precision data for 61 observations during this study study site species count replicates mean bootstrapped se cv 1 2 3 4 sd letaba ranch kudu 25 48 57 35 41.3 12.2 6.1 29.6 mopanieveld zebra 76 60 54 46 59.0 11.0 5.5 18.6 wildebeest 67 62 59 70 64.5 4.4 2.2 6.8 impala 633 727 615 446 605.3 102.4 51.2 16.9 giraffe 25 48 57 35 41.3 12.4 6.2 30.1 loskop dam kudu 263 274 260 237 258.5 13.8 6.9 5.3 mixed bushveld zebra 262 236 243 214 238.8 17.2 8.6 7.2 wildebeest 160 192 200 192 186.0 15.2 7.6 8.2 impala 285 346 291 345 316.8 28.6 14.3 9.0 giraffe 33 14 19 26 23.0 7.2 3.6 31.3 warthog 35 37 46 37 38.8 4.2 2.1 10.8 sable 43 41 45 36 41.3 3.4 1.7 8.2 white rhino 32 33 25 30 30.0 3.2 1.6 10.7 mountain reedbuck 163 149 170 141 155.8 11.4 5.7 7.3 vhembe kudu 3 13 7 7.7 4.2 2.4 54.2 mopanieveld kudu 26 30 50 35.3 10.4 6.0 29.4 kudu 28 26 21 25.0 2.9 1.7 11.8 zebra 13 3 5 7.0 4.3 2.5 61.9 zebra 55 55 61 57.0 2.8 1.6 4.9 zebra 24 25 20 23.0 2.3 1.3 9.8 wildebeest 23 0 21 14.7 10.4 6.0 70.9 wildebeest 94 106 100 100.0 4.8 2.8 4.8 wildebeest 32 76 54 54.0 18.0 10.4 33.4 wildebeest 70 64 69 67.7 2.6 1.5 3.8 impala 120 85 122 109.0 17.1 9.9 15.7 impala 210 160 178 182.7 20.4 11.8 11.2 impala 122 114 137 124.3 9.5 5.5 7.7 impala 175 288 372 278.3 81.9 47.3 29.4 giraffe 22 18 21 20.3 1.7 1.0 8.5 giraffe 4 4 4 4.0 0.0 0.0 0.0 warthog 5 3 2 3.3 1.2 0.7 36.4 warthog 21 26 11 19.3 6.2 3.6 32.3 eland 3 7 3 4.3 1.9 1.1 44.0 eland 5 11 10 8.7 2.6 1.5 30.0 eland 5 3 2 3.3 1.2 0.7 36.4 hartebeest 22 15 26 21.0 6.2 3.6 29.7 gemsbok 26 27 28 27.0 0.9 0.5 3.2 waterbuck 5 14 16 11.7 4.8 2.8 41.6 waterbuck 27 15 15 19.0 5.7 3.3 30.1 waterbuck 12 10 12 11.3 0.9 0.5 7.6 athersone kudu 132 116 108 118.7 9.9 5.7 8.3 nw arid kudu 22 26 33 27.0 4.5 2.6 16.7 bushveld kudu 19 33 24 25.3 5.7 3.3 22.6 kudu 34 33 28 31.7 2.6 1.5 8.2 kudu 9 7 9 8.3 1.0 0.6 12.5 zebra 37 39 30 35.3 4.0 2.3 11.3 wildebeest 219 185 230 211.3 19.1 11.0 9.0 reilly.qxd 2005/12/09 10:11 page 80 blue wildebeest similarly, in the case of blue wildebeest, seven cvs obtained range from 3.8 % to 70.9 % with a median of 8.2 %. the cvs for lowveld mopane veld, sourish mixed bushveld and arid bushveld are 6.8 %, 8.2 % and 9.0 % respectively while a median of 19.1 % was obtained for the northern mopane veld. impala eleven estimates of precision were obtained for impala with cvs ranging from the high of 70.9 % to 7.7 % with a median of 14.3 %. single values for lowveld mopane veld and sourish mixed bushveld are 16.9 % and 9.0 %, respectively. the median cv for northern mopane veld is 9.5 % and that for northwestern arid bushveld 9.7 %. giraffe there were only four estimates of precision obtained for giraffe with cvs of 30.1 %; 31.3 %; 19.0 % and 0 % for lowveld mopane veld, sourish mixed bushveld and the last two for northern mopane veld, respectively. this computes to a median of 24.6 %. all results for other species are presented in table 1. the regression analysis for all species and vegetation type (site) of the mean count (population) size by the standard error (se) shows significant correlation (r² = 0.71; p < 0.074; n = 63). this is an indication that the cv is constant in relation to population size indicating that the technique is robust. similar regressions by species for those species having several observations show a similar result. for impala, r² = 0.67, p < 0.002, n = 10; kudu, r² = 0.39, p < 0.054, n = 9; warthog, r² = 0.64, p < 0.018, n = 7; and zebra, r² = 0.86, p < 0.021, n = 8. in the case of wildebeest however, the result is not significant (r² = 0.17, p < 0.361, n = 6). further regression analysis by vegetation type shows significant coefficients of determination for lowveld mopane veld (letaba ranch) (r² = 0.99, p < 0.001, n = 4); sourish mixed bushveld (loskop dam) (r² = 0.86, p < 0.001, n = 8); northern mopane veld (vhembe) (r² = 0.72, p < 0.01, n = 25) and arid bushveld (atherstone) (r² = 0.81, issn 0075-6458 81 koedoe 45/2 (2002) table 1 (continued) study site species count replicates mean bootstrapped se cv 1 2 3 4 sd impala 477 477 346 433.3 62.0 35.8 14.3 impala 117 78 123 106.0 19.7 11.4 18.6 impala 17 16 0 11.0 7.8 4.5 70.9 impala 141 179 167 162.3 15.8 9.1 9.7 impala 69 82 66 72.3 6.9 4.0 9.6 warthog 87 79 32 66.0 24.9 14.4 37.8 warthog 71 81 49 67.0 13.5 7.8 20.2 warthog 52 29 36 39.0 9.7 5.6 24.9 warthog 54 33 41 42.7 8.7 5.0 20.3 warthog 67 39 27 44.3 16.3 9.4 36.7 hartebeest 7 10 32 16.3 11.1 6.4 67.9 hartebeest 29 22 30 27.0 3.6 2.1 13.5 hartebeest 79 91 71 80.3 8.3 4.8 10.3 waterbuck 18 23 29 23.3 4.5 2.6 19.3 reilly.qxd 2005/12/09 10:11 page 81 koedoe 45/2 (2002) 82 issn 0075-6458 p < 0.01, n = 20). the first mentioned of these could be an artefact of small sample size however. discussion comparing these results with other published results (hirst 1969; le resche & rausch 1974; rice & harder 1977; beasom 1979; mccullough 1994; brockett 2002) suggest that generally these results obtained appear consistent in range with those of other authors and are mainly unpredictable. the role of precision in monitoring is often misunderstood; in essence the precision can be seen as the “best minimum estimate” of ability of a chosen technique or application to show change in a population over time. this relates back to the fact that monitoring techniques show two sources of variation, real change over time and “within technique variation” which is an artifact of all contributory biases from the physical application of the technique. misconstruing the last mentioned as real change is statistically termed a type i error. the cv can be seen as a valuable guide in that a cv value less than the population change (%) to be detected increases the probability of showing real change over time whilst a cv that exceeds percentage change to be detected increases the probability of the change shown having actually not taken place in reality. singlepass aerial surveys or any other monitoring results that are unreplicated or lacking in a strategy to estimate precision cannot fulfill the role of decision support in this regard. in this study the means of the cv indicate that, in most cases, the precision is inadequate for decision making on an annual basis, making the use of single-pass surveys doubtful as a decision support tool in ungulate management. the ranges of results also indicate precision to be highly variable. to improve the situation, extending the time between counts will increase the change to be detected, thence improving the probability of detecting real change rather than artifacts related to external sources of bias. references acocks, j.p.h. 1988. veld types of south africa. memoirs of the botanical survey of south africa 57:1–146. bartmann, r.m.; l.h. carpenter; r.a. garrott & d.c. bowden. 1986. accuracy of helicopter counts of mule deer in pinyon-juniper woodland. wildlife society bulletin 14(4): 356–363. beasom, s. l. 1979. precision in helicopter censusing of white-tailed deer. journal of wildlife management 43(3): 777–780. beasom, s.l.; f.g. leon iii & d.r. synatzske. 1986. accuracy and precision of counting whitetailed deer with helicopters at different sampling intensities. wildlife society bulletin 14(4): 364–368. bothma, j. du p.; m.j.s. peel; s. pettit & d. grossman. 1990. evaluating the accuracy of some commonly used game-counting methods. south african journal of wildlife research 20(1): 26–32. brockett, b.h. 2002. accuracy, precision and bias of helicopter-based counts of black rhinoceros in pilanesburg national park, south africa. south african journal of wildlife research 32(2): 121136. cahalane, v.h. 1938. the annual northern yellowstone elk herd count. transactions of the north american wildlife conference 3: 388–389. caughley, g. 1974. bias in aerial survey. journal of wildlife management 38: 921–933. caughley, g. 1979. analysis of vertebrate populations. new york: wiley. collinson, r.h.f. 1985. selecting wildlife census techniques. monograph 6, institute of natural resources, university of natal, pietermaritzburg. dasmann, r.f. & a.s. mossman. 1962. road strip counts for estimating numbers of african ungulates. journal of wildlife management 26: 101–104. deyoung, c. a. 1985. accuracy of helicopter surveys of deer in south texas. wildlife society bulletin 13 (2): 146–149. efron, b. 1979. bootstrap methods: another look at the jackknife. annals of statistics 7(1): 1–26. eiselen, r. 1994. estimating the trends in herbivore numbers in the southern district of the kruger national park. south african journal of wildlife research 24(4): 95–100. gerrodette, t. 1993. trends: software for a power analysis of linear regression. wildlife society bulletin 21: 515–516. goodman, p.s. 1977. an evaluation of three techniques for estimating animal numbers in a wildlife population. unpub. mimeographed report., natal parks board, pietermaritzburg. graham, a. & r. bell. 1989. investigating observer bias in aerial survey by simultaneous doublereilly.qxd 2005/12/09 10:11 page 82 issn 0075-6458 83 koedoe 45/2 (2002) counts. journal of wildlife management 53(4): 1009–1010. hirst, s.m. 1969. road-strip census techniques for wild ungulates in african woodland. journal of wildlife management 33 (1): 40–48. jolly, g.m. 1969. the treatment of errors in aerial counts of wildlife populations. east african agricultural & forestry journal, special issue, 34: 50–55. leresche, r.e. & r.a. rausch. 1974. accuracy and precision of aerial moose censusing. journal of wildlife management 38: 175–182. mccullough, d.r. 1994. what do herd composition counts tell us? wildlife society bulletin 22(2): 295–300. melton, d.a. 1978. undercounting bias of helicopter census in umfolozi game reserve. lammergeyer 26: 1–6. norton-griffiths, m. 1978. reducing counting bias in aerial census by photography. east african wildlife journal 12: 245–248. otten, m. r.m.; j. b. haufler; s. r. winterstein & l. c. bender. 1993. an aerial censusing procedure for elk in michigan. wildlife society bulletin 21(1): 73–80. payne, n. f. 1981. accuracy of aerial censusing for beaver colonies in newfoundland. journal of wildlife management 45(4): 1014–1016. pennycuick, c.j. & d. western. 1972. an investigation of some sources of bias in aerial transect sampling of large mammal populations. east african wildlife journal 10: 175–191. reilly, b.k. 2000. the statistics of helicopter total counts of large ungulates in sourish mixed bushveld, northwest arid bushveld and mopane veld, republic of south africa. unpublished phd thesis, university of stellenbosch. rice, w.r. & j.d. harder. 1977. application of multiple aerial sampling to mark-recapture census of white-tailed deer. journal of wildlife management 41: 197–206. seber, g.a.f. 1992. a review of estimating animal abundance ii. international statistical review 60(2): 129–166. van hensbergen, h.j., m.p.s. berry & j. juritz. 1996. helicopter based line transect estimates of african herbivore populations. south african journal of wildlife research 26(3): 81–87. van lavieren, l.p. & esser. j.d. 1979. numbers, distribution and habitat preference of large mammals in bouba ndjida national park, cameroon. african journal of ecology 17: 141–153. reilly.qxd 2005/12/09 10:11 page 83 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 filelist convert a pdf file! page 1 page 2 kyle mark recapture.qxd more information from the mark and recapture of fish at kosi bay, south africa r. kyle issn 0075-6458 121 koedoe 45/2 (2002) a considerable amount of work has been carried out on the mark and recapture of fish at kosi bay (kyle 1986). this resulted in the publication of data (kyle & robertson 1997) suggesting fairly low rates of exploitation by man but warning that further information was needed to confirm apparent levels. fish marking has been ongoing (kyle 2000) and results of more recent mark and recaptures are given here. the methodology employed was similar to that described in kyle & robertson (1997) and tagging has continued since the early 1980s at similar levels, apart from a focal study in 1984/1985. fish were marked in the southern lakes of kosi bay and most recoveries were made in the northern channels as the fish migrated out of the lakes towards the ocean to spawn (garratt 1992). to facilitate recoveries, the fish marking was widely discussed with local fishermen. all the fish marked were caught on rod and line. most recoveries are made by trap, rod and line or gillnet subsistence fishermen. the mean time between marking and recovery was 186 days, with a minimum of 98 days between marking and recovery. from august 1988 to july 1999 a total of 73 marked fish were recovered, giving a mean annual recovery rate of 6.6 fish from the lakes and ocean. during the twelve month period august 1999 to july 2000 a total of 17 fish were recovered from local subsistence fishermen, suggesting a higher level of exploitation. analysis of mark and recapture data for the lakes, for the period august 1999 to july 2000 (table 1), shows that 24.3 % of all the fish were recovered within one year of marking and 38.9 % of the acanthopagrus berda were recovered. this compares with 4.7 % from a table 1 data on marking and recapture of fish in the kosi lakes from august 1999 to july 2000 species marked recaptures % acanthopagrus berda 18 7 38.9 lutjanus argentimaculatus 5 0 0 caranx ignobilis 5 1 20 caranx sexfasciatus 4 0 0 caranx papuensis 4 0 0 sphyrnidae 1 1 100 total 37 9 24.3 similar multispecies exercise in 1983/1984 and 5.4 % for a focal study on a. berda in 1984/1985 (kyle & robertson 1997). although little has changed in the technique of fish mark and recapture in the kosi lakes, it is possible that notification of the capture of tagged fish improved, as knowledge of the small financial reward for marked fish spread, resulting in some of the increase in the proportion of fish recovered. over the whole period of marking and recovery, three sphyraena jello gave useful information on growth rates (table 2) as well as suggesting their residence for several years in the lakes. data suggest a remarkably standard growth rate of about 1 mm/day. three caranx ignobilis of between 10–13 kg each, were marked. one was recovered in a fishtrap inside the lakes. another was recaught by a recreational angler in the ocean, about 30 km south of kosi mouth, released again and recovered and released again back in the lakes by another recreational angler. this suggests heavy fishing pressure on these large fish and the marine recovery is the first for a fish marked inside the kosi lakes. the subsequent natural history note kyle mark recapture.qxd 2005/12/09 10:07 page 121 recovery again, inside the lakes, confirms the multiple entry and exit of this species, the usefulness of fish mark and recovery generally, and the suitability of the species for mark and recapture work. one 12 kg c. ignobilis was hooked on light tackle and played gently for about 1.5 h before being carefully tagged, “walked” in the shallows to recover, and released. it was found dead two days later. a second, similar fish was caught the day following the first capture, but it was hooked on heavy tackle, brought in as soon as possible, marked quickly and released immediately. it was not seen again and it is assumed to have survived. this phenomenon has been reported before (bagnall 2000) and it appears that survival of marked fish may be enhanced by rapid capture and release, while gentle tactics result in exhausted fish which may die after release. many recaptured a. berda were recovered very close to the marking sites suggesting a high degree of residency in this species (kyle 2000). although not as many data are available, there is little such evidence for other species marked at kosi bay. the kosi lakes fishery is probably the most important true subsistence fishery in south africa and the recreational fishery is of increasing importance to the local and national economy. to date, it has been thought that fish stocks are adequate for these, plus a limited gillnet fishery, to exist in relative harmony without unduly impacting fish stocks. data presented here, however, suggest that total offtake levels have increased in recent years and would suggest a cautious approach to allowing any significant increase in fishing effort. some recovered marked fish are still, however, probably not recorded and known recovery levels of almost 40 % for some species are cause for alarm about overexploitation. little accurate information is available on illegal fishing, such as unlicensed gillnetting, and no recoveries were recorded recently from this method. as it is known that numbers of a. berda are caught by these fishermen it is likely that true overall fish catch rates are markedly higher than those indicated above. marked fish recovery data thus suggest that if kosi bay fish stocks are to be maintained, illegal fishing at kosi bay should be greatly reduced and no new fishing should be allowed unless there is a marked reduction in some present fishing levels or methods. acknowledgements i thank the ezemvelo kwazulu natal wildlife for permission to publish this paper. references bagnall, r. 2000. giving him the stick. the fishing journal 3(3): 10-11. garratt, p. a. 1992. spawning of riverbream, acanthopagrus berda, in kosi estuary. south african journal of zoology 28(1): 26-31. kyle, r. 1986. aspects of the ecology and exploitation of the fishes of the kosi bay system, kwazulu, south africa. unpublished phd thesis, university of natal, durban. kyle, r. 2000. some interesting information from tagging returns for the riverbream, acanthopagrus berda, in the kosi lakes, kwazulu/natal. koedoe 43(1): 99-100. kyle, r. & w.d. robertson. 1997. preliminary estimates of population size and capture rates of mature acanthopagrus berda in the kosi lakes system, south africa, using mark-recapture methods. south african journal of zoology 32(4): 124-128. r. kyle, p.o. box 43, kwangwanase, 3973, rsa. koedoe 45/2 (2002) 122 issn 0075-6458 table 2 growth information, of total length in millimetres, from three sphyraena jello marked and recovered in the kosi lakes date marked date recovered days free mark l. recovery l. growth 29 december 1999 10 march 2000 120 800 940 140 31 march 1999 9 april 2000 390 550 940 390 11 december 1997 15 april 2000 540 380 920 540 kyle mark recapture.qxd 2005/12/09 10:07 page 122 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <feff004200720075006700200064006900730073006500200069006e0064007300740069006c006c0069006e006700650072002000740069006c0020006100740020006f0070007200650074007400650020005000440046002d0064006f006b0075006d0065006e0074006500720020006d006500640020006800f8006a006500720065002000620069006c006c00650064006f0070006c00f80073006e0069006e0067002000740069006c0020007000720065002d00700072006500730073002d007500640073006b007200690076006e0069006e0067002000690020006800f8006a0020006b00760061006c0069007400650074002e0020005000440046002d0064006f006b0075006d0065006e007400650072006e00650020006b0061006e002000e50062006e006500730020006d006500640020004100630072006f0062006100740020006f0067002000520065006100640065007200200035002e00300020006f00670020006e0079006500720065002e00200044006900730073006500200069006e0064007300740069006c006c0069006e0067006500720020006b007200e600760065007200200069006e0074006500670072006500720069006e006700200061006600200073006b007200690066007400740079007000650072002e> /nld <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> /esp <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> /suo <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> /ita <feff00550073006100720065002000710075006500730074006500200069006d0070006f007300740061007a0069006f006e00690020007000650072002000630072006500610072006500200064006f00630075006d0065006e00740069002000500044004600200063006f006e00200075006e00610020007200690073006f006c0075007a0069006f006e00650020006d0061006700670069006f00720065002000700065007200200075006e00610020007100750061006c0069007400e00020006400690020007000720065007300740061006d007000610020006d00690067006c0069006f00720065002e0020004900200064006f00630075006d0065006e00740069002000500044004600200070006f00730073006f006e006f0020006500730073006500720065002000610070006500720074006900200063006f006e0020004100630072006f00620061007400200065002000520065006100640065007200200035002e003000200065002000760065007200730069006f006e006900200073007500630063006500730073006900760065002e002000510075006500730074006500200069006d0070006f007300740061007a0069006f006e006900200072006900630068006900650064006f006e006f0020006c002700750073006f00200064006900200066006f006e007400200069006e0063006f00720070006f0072006100740069002e> /nor <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> /sve <feff0041006e007600e4006e00640020006400650020006800e4007200200069006e0073007400e4006c006c006e0069006e006700610072006e00610020006e00e40072002000640075002000760069006c006c00200073006b0061007000610020005000440046002d0064006f006b0075006d0065006e00740020006d006500640020006800f6006700720065002000620069006c0064007500700070006c00f60073006e0069006e00670020006600f60072002000700072006500700072006500730073007500740073006b0072006900660074006500720020006100760020006800f600670020006b00760061006c0069007400650074002e0020005000440046002d0064006f006b0075006d0065006e00740065006e0020006b0061006e002000f600700070006e006100730020006d006500640020004100630072006f0062006100740020006f00630068002000520065006100640065007200200035002e003000200065006c006c00650072002000730065006e006100720065002e00200044006500730073006100200069006e0073007400e4006c006c006e0069006e0067006100720020006b007200e400760065007200200069006e006b006c00750064006500720069006e00670020006100760020007400650063006b0065006e0073006e006900740074002e> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice introduction methods results discussion conclusion acknowledgements references about the author(s) gary k. nicolau department of zoology and entomology, faculty of science, rhodes university, makhanda, south africa emily a. jackson department of zoology and entomology, faculty of science, rhodes university, makhanda, south africa ryan j. wasserman department of zoology and entomology, faculty of science, rhodes university, makhanda, south africa south african institute for aquatic biodiversity, makhanda, south africa citation nicolau, g.k., jackson, e.a. & wasserman, r.j., 2023, ‘mass vehicle induced mortalities of giant bullfrogs in nylsvley nature reserve’, koedoe 65(1), a1760. https://doi.org/10.4102/koedoe.v65i1.1760 short communication mass vehicle induced mortalities of giant bullfrogs in nylsvley nature reserve gary k. nicolau, emily a. jackson, ryan j. wasserman received: 25 oct. 2022; accepted: 10 may 2023; published: 08 aug. 2023 copyright: © 2023. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. introduction scientific interest in roadkill and road ecology is on the increase, given the massive role roads play in dividing the earth’s terrestrial surfaces (e.g. collinson et al. 2014; perkins, shilling & collinson 2022; sutherland, dunning & baker 2010). linear structures such as road networks have the capability of substantially compromising terrestrial landscapes and their associated biota (forman & alexander 1998). while the threats imposed by road networks vary considerably (trombulak & frissell 2000), the most commonly observed impact is that of wildlife-vehicle collisions, causing what is commonly referred to as roadkill. in some regions, amphibians have been observed to be the most impacted vertebrates, by road networks (glista, devault & dewoody 2008), yet more work is needed to fully assess impacts on populations and roadkill mitigation (beebee 2013). certain amphibians are at risk of roadkill predominantly because of their life history strategies, where they have large congregations, often moving onto nearby roads during or after breeding seasons, while their slow movement and the lack of detectability by drivers add to their vulnerability (puky 2005; trombulak & frissell 2000). when roads are built through or near wetlands and other water sources used for breeding by amphibians, the threat of amphibian roadkill is intensified. the nylsvley nature reserve (nnr) is a protected area situated in the waterberg district of the limpopo province, south africa. within the centre of the nnr is the nyl river floodplain, a vital inland wetland fed from the waterberg massif. because of its conservation importance, specifically to that of avifauna, the nnr has been declared a ramsar site. as a result of its proximity to the major cities of johannesburg and pretoria, the exceptional biodiversity, and self-driving option for visitors, the reserve is a tourism hotspot, leading to heavy traffic during peak holiday periods. jacobsen (1977) published extensive work on the amphibian diversity of the nnr, whereby a total of 18 species of amphibians were documented within its borders. because of taxonomic changes and increased sampling efforts, approximately 24 species are now expected to occur within nnr. pyxicephalus adspersus (giant bullfrog) (jacobsen 1977) and more recently pyxicephalus edulis (african bullfrog) are both species that have been recorded within the boundaries of the reserve. members of the pyxicephalus genus display mass breeding congregations, typically taking place in temporary pans or wetlands for a few weeks after heavy rainfall in their breeding season. pyxicephalus adspersus and p. edulis are listed under the least concern category of the international union for conservation of nature (iucn) red list (iucn ssc amphibian specialist group 2013, 2016). however, both species have been reported to be experiencing population declines, primarily because of urbanisation causing conversion and degradation of suitable habitats (iucn ssc amphibian specialist group 2013, 2016). roadkill is considered a significant contributor to reductions in pyxicephalus populations; however, no major work has been performed to investigate the true extent of this threat. this study reports on short-term, mass vehicle-induced mortalities of juvenile p. adspersus within the borders of the nnr. the study serves as an ad hoc observation of how even minimal vehicle traffic during spawning events can generate high levels of roadkill. methods on 14 february 2021, during a relatively wet season, ad hoc observations of extensive dead p. adspersus were observed on the main road and on many other roads of the nnr. this prompted the researchers (g.k.n. and e.a.j.) to walk a transect of approximately 2.7 km on the main road (figure 1a), starting at the campsite (24° 38′ 58″ s; 28° 40′ 24″ e) and ending at the turnoff to jacana hide (24° 38′ 34″ s; 28° 41′ 37″ e). during the walked transect, the climatic conditions were hot and humid with no wind and heavy cloud cover. each researcher walked along the main tyre tracks of the road, able to observe each half of the axial transect. the transect of the road was divided into 11 quadrants of c.a. 250 m long and dead pyxicephalus spp. identified and enumerated within each section (live pyxicephalus spp. were roughly estimated based on the size of groups present on the road and not counted because of the high density, of which many retreated into surrounding vegetation when approached). all roadkill consisted of individuals ~500 mm in size, clearly indicating they had recently completed metamorphosis. for identification, approximately every 20th dead on-road individual was closely inspected for the presence or absence of the following: (1) white dot on the tympanum, (2) several prominent, (3) interrupted skin ridges on the back and (4) a pale vertebral stripe, all characters that assist in distinguishing between juvenile p. adspersus and p. edulis. because of the quantity of roadkill along the transect, and that no material was handled, the given method of identifying a subset of individuals was the most practical approach. as all tourist vehicles entering the nrr are required to sign a guest book upon entrance to the protected area, a record of traffic levels was also available. as such, the number of vehicles that entered the provincial park over the weekend was determined from the guest book. the number of times a car drove the single transect could not, however, be determined. yet, all vehicles that enter the park will likely take the road surveyed at least once when entering or exiting the reserve. figure 1: (a) study site: red line – transect walked to identify pyxicephalus spp. roadkill on the main road within the nylsvley nature reserve, limpopo, south africa. white dots – start and end of each section of the transect. green line – proposed alternative route to mitigate excessive roadkill during breeding season. blue – nylsvley floodplain. insert – location of nylsvley nature reserve. (b) dead on road juvenile pyxicephalus spp. on the main tyre tracks in nylsvley nature reserve, limpopo, south africa. (c) a juvenile pyxicephalus spp. scavenging on other juvenile pyxicephalus spp. killed on the road. (d) a depression in the road formed high congregations of juvenile pyxicephalus spp. results the total length of the transect produced 1005 dead juvenile pyxicephalus spp. (table 1). all ~ 50 dead bullfrogs assessed were identified as p. adspersus. most roadkill observed were fresh (figure 1b, c) and likely driven over either on 14 february, with a few expected to have been killed on the 13 february 2021. of the 1005 dead bullfrogs, only 15, based on their dehydrated appearance, were considered to have been killed more than 48 h before the observation and were, thus, excluded from the total observations. table 1: observations of pyxicephalus spp. killed on a transect of the main road within the nylsvley nature reserve between 24° 38′ 58″ s; 28° 40′ 24″ e and 24° 38′ 34″ s; 28° 41′ 37″ e dor = dead on road. a total of 81 vehicles signed in and entered the gate between 13 february 2021 and 14 february 2021. the highest density of roadkill was present in the first 250 m road (1.02 dead on-road individual per meter), located in the flooded grasslands of a drainage line north of the main floodplain (figure 1a). the density of roadkill started to decrease after 1.25 km, once the road bordered the main nylsvley floodplain. estimates of live individuals also began to decline within the latter part of the transect. on the day of the survey and the following day, many active individuals were seen in the early mornings (figure 1d) and late afternoons. while midday had active individuals on the road, intensity of activity declined. when a vehicle approached, juveniles would scatter across to the sides of the road. yet even with the slow speed of a tourist vehicle, many were still unable to escape death. large congregations were often observed, these were often in muddy depressions in the road (figure 1d) or in sections were marshy grassland bordered the transect. discussion despite minimal, slow-moving traffic within a protected area, excessive pyxicephalus roadkill was observed along a single road in the protected area nnr. while all inspected pyxicephalus dead on-road were identified as p. adspersus, the presence of p. edulis among all the mortalities is plausible given that they have been recorded in the nnr. these findings further highlight how road networks that skirt wetlands are likely to increase vehicle-collision risk for certain taxa. these water bodies, embedded within a terrestrial matrix, serve as foraging and breeding patches for various invertebrate and vertebrate species (cuthbert et al. 2022). amphibians, as semi-aquatic organisms that typically require waterbodies to complete their life-cycles, are vulnerable to road mortalities in these areas. those species with explosive breeding strategies, such as pyxicephalus spp., are likely particularly vulnerable given their proclivity for mass aggregation by juveniles upon metamorphosis. while this brief ad hoc study aimed to highlight the levels of bullfrog roadkill in a protected area over a short period, the quantified results only represented a fraction of mortalities that took place over the 2 days within the reserve’s boundaries and the surrounding roads. other roads within the park showed mortalities of similar quantities. in contrast, road networks outside the park appeared to have lower mortality numbers but would have likely had a greater traffic volume. it is possible that this is either because of more suitable breeding habitats within the reserve, these roads are less accessible for the young pyxicephalus or that populations along the roads had already decreased because of roadkill. it is possible that pyxicephalus roadkill numbers were an underestimation, given that an abundance of predators were observed scavenging the dead on road and alive juvenile pyxicephalus spp. these were not only predominantly avifauna but also included other opportunists such as a southern tree agama (jackson & nicolau 2022) and other pyxicephalus individuals (figure 1c). indeed, santos, carvalho and mira (2011) found that amphibian carcases typically do not last long on the road relative to other taxonomic groups, with scavenging contributing to this observation. in several cases during this study, scavenging events by juvenile pyxicephalus on dead on-road pyxicephalus resulted in further deaths, after failing to move for oncoming vehicles. it has been estimated that roadkill can significantly influence certain amphibian populations. hels and buchwald (2001) reported that up to 10% of certain populations of rana temporaria (common frog) and rana arvalis (moor frog) were killed annually through vehicle-induced mortalities. smith and dodd (2003), over a period of 1 year, found that 91% of all mortalities documented on a single road were herpetofauna, of which the majority were amphibians. similar results were found by glista et al. (2008), where 95% of all roadkill were reptiles and amphibians. the findings of the present study suggest that pyxicephalus spp. may be particularly vulnerable to roadkill at certain times of the year. similarly, a study assessing amphibian roadkill in the north of the limpopo province found that p. edulis roadkill densities were among the highest of all species observed (hlatshwayo et al. 2022). areas of concern are likely those where breeding and tadpole metamorphosis takes place. this was presumably the case within the first section of the transect at nnr, where both roadkill and live estimates were elevated. there are many variables that could influence congregations of metamorphs, with the most likely being that this area was the most suitable breeding ground during the season. it is plausible that simple mitigation measures for the species are possible, particularly in protected areas. practical roadkill mitigation measures for amphibians have been through volunteer groups, along with temporary fencing and pitfall traps, allowing amphibians to be captured and moved across the road (beebee 2013). mitigation efforts such as small road tunnels and fences have also proven to be effective in reducing amphibian deaths (beebee 2013), with one study showing a decrease in roadkill between 85% and 100% (helldin & petrovan 2019). similarly, signage has been proven to be successful in limiting roadkill (beebee 2013), through the alerting of drivers to the potential presence of fauna. in the nrr specifically, there is no dangerous megafauna within the reserve, which often hinders the implementation of temporary fencing and pitfall traps (jordaan 2022). a simple culvert and drift fences over the first 250 m of the transect could potentially have mitigated much of the mortalities that occurred over 2 days. similarly, as the public road network within nnr is a looped route, tourists could be restricted to entering and exiting the high-mortality sections during key periods. a short alternative road leading to the jacana hide is present within the reserve (figure 1a). despite still going past the floodplain, this route mitigates driving through the section with the highest mortalities from this study. should the decision of alternative routes be taken by conservation officials, it is essential to determine whether the spawning events take place in the same locations each season. alternative routes would likely be more successful than signs, given that the observed numbers of pyxicephalus were so high that even cautious drivers would not be able to avoid all individuals on the road. conclusion this study further highlights that even in formally protected areas, roadkill can be significant and warrants continuous management consideration. the extent a short road network with minimal traffic had on amphibian mortalities was considerable. this protected area could represent an ideal pilot site to investigate the efficacy of local simple and cost-effective mitigation measures on road networks in the proximity of wetland ecosystems, within the context of amphibian roadkill. as a part of this process, further data and monitoring of pyxicephalus spp. roadkill and population levels within the ramsar-protected area are required. acknowledgements the authors would like to thank the staff at nylsvley for allowing access to the guest book to obtain the number of persons entering the park over the period of time needed. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions g.k.n. and e.a.j. carried out the data collection. g.k.n. wrote the article with input from all authors e.a.j. and r.j.w. all authors g.k.n., e.a.j. and r.j.w. conceptualised the discussed project, discussed the results of the findings and contributed to the final manuscript. g.k.n., e.a.j. and r.j.w. contributed funds to the project. ethical considerations this article followed all ethical standards for research without direct contact with human or animal subjects. funding information this work was supported by the south african institute for aquatic biodiversity (project name: assessing herpetofauna diversity associated with freshwater ecosystems). data availability data sharing is not applicable to this article, as no new data were created or analysed in this study. disclaimer the views expressed in the article are that of the authors and not an official position of the institution or funder. references beebee, t.j.c., 2013, ‘effects of road mortality and mitigation measures on amphibian populations’, conservation biology 27(4), 657–668. https://doi.org/10.1111/cobi.12063 collinson, w.j., parker, d.m., bernard, r.t.f., reilly, b.k. & davies-mostert, h.t., 2014, ‘wildlife road traffic accidents: a standardized protocol for counting flattened fauna’, ecology and evolution 4(15), 3060–3071. https://doi.org/10.1002/ece3.1097 cuthbert, r.n., wasserman, r.j., keates, c. & dalu, t., 2022, ‘food webs’, in t. dalu & r.j. wasserman (eds.), fundamentals of tropical freshwater wetlands, pp. 517–547, academic press, london. forman, r.t.t. & alexander, l.e., 1998, ‘roads and their major ecological effects’, annual review of ecology and systematics 29, 207–231. https://doi.org/10.1146/annurev.ecolsys.29.1.207 glista, d.j., devault, t.l. & dewoody, j.a., 2008, ‘vertebrate road mortality predominantly impacts amphibians’, herpetological conservation and biology 3(1), 77–87. helldin, j.o. & petrovan, s.o., 2019, ‘effectiveness of small road tunnels and fences in reducing amphibian roadkill and barrier effects at retrofitted roads in sweden’, peerj 7, e7518. https://doi.org/10.7717/peerj.7518 hels, t. & buchwald, e., 2001, ‘the effect of road kills on amphibian population’, biological conservation 99, 331–340. https://doi.org/10.1016/s0006-3207(00)00215-9 hlatshwayo, t.i., stam, e.m., collinson-jonker, w.j. & dawood, a., 2022, ‘an inventory of amphibian roadkill in the western soutpansberg, limpopo province, south africa’, african journal of herpetology 72(1), 1–17. https://doi.org/10.1080/21564574.2022.2115154 iucn ssc amphibian specialist group, 2013, pyxicephalus adspersus. the iucn red list of threatened species 2013, viewed 10 november 2022, from https://doi.org/10.2305/iucn.uk.2013-2.rlts.t58535a3070700.en iucn ssc amphibian specialist group, 2016, pyxicephalus edulis. the iucn red list of threatened species 2016, viewed 10 november 2022, from https://doi.org/10.2305/iucn.uk.2016-1.rlts.t76317591a87648183.en jackson, e.a. & nicolau, g.k., 2022, ‘acanthocercus atricolus diet’, african herp news 81, 27–29. jacobsen, n.h.g., 1977, an annotated checklist of the amphibians, reptiles and mammals of the nylsvley nature reserve, cooperative scientific programmes, csir, pretoria. jordaan, p.r., 2022, ‘practical considerations for conducting surveys in the presence of potentially dangerous wildlife’, african herp news 79, 17–31. perkins, s.e., shilling, f. & collinson, w., 2022, ‘anthropause opportunities: experimental perturbation of road traffic and the potential effects on wildlife’, frontiers in ecology and evolution 10, 833129. https://doi.org/10.3389/fevo.2022.833129 puky, m., 2005, amphibian road kills: a global perspective, uc davis: road ecology center, viewed 10 november 2022, from https://escholarship.org/uc/item/7j7546qv. santos, s.m., carvalho, f. & mira, a., 2011, ‘how long do the dead survive on the road? carcass persistence probability and implications for road-kill monitoring surveys’, plos one 6, e25383. https://doi.org/10.1371/journal.pone.0025383 smith, l.l. & dodd, jr. c.k., 2003, ‘wildlife mortality on us highway 441 across paynes prairie, alachua county, florida’, florida scientist 66(2), 128–140. sutherland, r.w., dunning, p.r. & baker, w.m., 2010, ‘amphibian encounter rates on roads with different amounts of traffic and urbanization’, conservation biology 24(6), 1626–1635. https://doi.org/10.1111/j.1523-1739.2010.01570.x trombulak, s.c. & frissell, c.a., 2000, ‘review of ecological effects of roads on terrestrial and aquatic communities’, conservation biology 14(1), 18–30. https://doi.org/10.1046/j.1523-1739.2000.99084.x 157table-20 july 2009[final version].indd a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe original research a rticle #157 1vol. 51 no. 1 page 1 of 3 date tpc level1 tpc detail species of concern management response expected outcome at the time of the tpc notification and current status oct. 1999 multi-level tpcs not in use imminent outside threat to the knp (crocodile river) chromolaena odorata (chromolaena) a special working for water contract was given to have the localised patch of plants eradicated the aim was to eradicate the patch of plants, but it soon became evident that the patch would not be completely removed. c. odorata become part of a long-term, ongoing management programme oct. 1999 multi-level tpcs not in use new record in the knpplants observed at sunset dam and orpen dam azolla filiculoides (red water fern) introduction of biological control agents long-term, ongoing control using biocontrol agents nov. 1999 multi-level tpcs not in use new distribution in the knp and an increase in distribution (luvuvhu and limpopo river systems) oreochromis niloticus (nile tilapia) to support the efforts of the provincial conservation authorities to curb the dispersal of these fish. to monitor the increase in distribution of the fish. to research the genetic mixing of o. niloticus and o. mossambicus (indigenous species) management / control of a fish species is unlikely in most situations, although further spread through anthropogenic means should be prevented. however, information is needed on the potential impacts to the ecosystem and other related species mar. 2000 multi-level tpcs not in use new distribution within the knp (n’waswitshaka firebreak, skukuza) agave sisalana (sisal) detailed survey of the area and chemical control of the plants the aim was to eradicate this patch of plants. while the plants appear to be eradicated, periodic site visits are still carried out mar. 2000 multi-level tpcs not in use increase in distribution in the knp (new records of both species along the skukuza – lower sabie road) senna occidentalis (wild coffee) and senna bicapsularis (rambling cassia) detailed survey of the roadsides and mechanical / chemical control of the plants although the aim was to eradicate the plants this was found to be unfeasible and the sites still require ongoing control may 2000 multi-level tpcs not in use new occurrence of an invasive alien plant species within the boundary of the knp (near skukuza) helianthus annuus (sunflower) and nicandra physaloides (apple of peru) survey of the area for additional plants and physical removal as there were only one plant of each species found, they were removed and it was felt unlikely that either species had established populations in the knp july 2000 multi-level tpcs not in use imminent outside threat to the crocodile river tithonia diversifolia (mexican sunflower) t. diversifolia was added to the list of species targeted by the alien plant control teams ongoing control is required, as it is unlikely that this species will ever be eradicated oct. 2000 multi-level tpcs not in use imminent outside threat (malelane) acridotheres tristis (indian myna) requested provincial conservation authorities to control the myna’s. requested all staff to report observations of myna’s in the knp myna’s are closely associated with human settlements and are likely to increase in numbers in towns bordering the knp. myna’s found in the knp should be destroyed. sept. 2000 multi-level tpcs not in use imminent outside threat to the knp, (hans merensky country club, phalaborwa). chromolaena odorata (chromolaena) control efforts were required to start immediately (mechanical / chemical). ongoing monitoring and follow-up work the aim was to eradicate the patches of plants, but it was unlikely that these plants could be eradicated and would have to form part of ongoing follow-up operations sept. 2000 multi-level tpcs not in use any new occurrence of an invasive alien plant species within the boundary of the knp, (on the tributary of the tshutshi spruit, phalaborwa) chromolaena odorata (chromolaena) control efforts were required to start immediately (mechanical / chemical). ongoing monitoring and follow-up work the aim was to eradicate the patches of plants, but it was unlikely that these plants could be eradicated and would have to form part of ongoing follow-up operations nov. 2000 multi-level tpcs not in use new occurrence in the knp reported from numbi and shitlave, pretoriuskop region varroa (bee) mite ongoing monitoring unknown, potentially large impacts on bee colonies dec. 2000 multi-level tpcs not in use new distribution in knp (talamati and lower sabie) acridotheres tristis (indian myna) requested provincial conservation authorities to control the myna’s. requested all staff to report observations of myna’s in the knp myna’s are closely associated with human settlements and are likely to increase in numbers in towns bordering the knp. myna’s found in the knp should be destroyed feb. 2001 multi-level tpcs not in use increase in distribution of a species in the knp (mlondozi dam, n’waswitsontso river and sabie river) cardiospermum halicacabum (balloon vine) survey of the areas invaded and c. halicacabum to be added to the list of species controlled c. halicacabum to form part of the ongoing maintenance control work in the knp feb. 2001 multi-level tpcs not in use new occurrence of an invasive alien plant species within the boundaries of the knp (two small patches at pafuri and vlakteplaas) arundo donax (giant reed) plants to be manually removed and any coppice to be sprayed with an appropriate herbicide these small patches were to have been eradicated. while it appears that they have been eradicated, periodic site visits are still carried out june 2001 multi-level tpcs not in use new distribution in the knp and an increase in distribution (olifants and crocodile rivers) hypophtalmichthys molitrix (silver carp) to support the efforts of the provincial conservation to curb the dispersal of these fish. to monitor the increase in distribution of the fish. management / control of a fish species is unlikely in most situations, although further spread through anthropogenic means should be prevented. information is however needed on the potential impacts to the ecosystem and other related species oct. 2002 2 first occurrence from a new grid cell and any new grid cell invaded that is not contiguous with grid cells invaded previously numerous patches along the luvuvhu , letaba, klein letaba, olifants, sabie and crocodile rivers chromolaena odorata (chromolaena) continued surveys, monitoring and increased vigilance to determine the extent of the distribution of these plants. immediate control of all plants. a number of regional workshops. an internal scientific report. control and containment of the plants still continues may 2003 2 first ever record from a new grid cell (or new area)gutchwa spruit and nsikazi river eichhornia crassipes (water hyacinth) chemical control of plants upstream of the knp and biological control for plants from the knp boundary downstream. eventual eradication of the plants outside the knp and containment / long-term biological control of the plants inside the knp table 2 list of alien species tpc exceedences to date koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research foxcroft a rt ic le # 15 7 2 vol. 51 no. 1 page 2 of 3 may 2003 2 first ever record from a new grid cell (or new area)roadsides in the crocodile bridge and lower sabie areas parthenium hysterophorus (parthenium) ongoing mapping / surveying. develop research projects on the potential spread and impacts of p. hysterophorus. chemical control. containment of the plants at the current sites. while much research and distribution mapping has taken place, little chemical control has been done may 2003 2 first ever record from a new grid cell (or new area)sabie river, near skukuza colocasia esculenta (elephants ears) the two patches of plants to be removed through manual (digging) means. follow-up by means of herbicides the aim was eradication of these two patches of plants. it appears that the patches have been eradicated, although periodic site visits are still required may 2003 2 first ever record from a new grid cell (or new area)numerous dams and rivers in the knp aplexa marmorata; physa acuta; lymnaea columella (alien snails) follow-up survey in 5 years time and research to be initiated on the potential impacts of the snails control unlikely, but an understanding of the potential impacts is necessary july 2003 3 increase in density at engelhard dam eichhornia crassipes (water hyacinth) biological control of the makhadzi spruit and chemical control on the main body of engelhard dam long-term sustainable control largely through the use of biocontrol. chemical control has continued to be used as an integrated management option on engelhard dam july 2003 1 new occurrence of a species in the knppafuri region australocylindropuntia cylindrical chemical control eradication of this species from the knp. periodic site visits are still required sept. 2003 1 new occurrence of a species in the knpmalelane, crocodile river tarebia granifera (snail) monitoring of spread, research on possible future impacts control unlikely, but an understanding of the potential impacts is necessary oct. 2003 2 increase in distribution in the knp skukuza animal holding bomas acridotheres tristis (indian myna) control if possible and removal of nests if necessary eradication of birds at this site (the bird disappeared shortly after tabling this tpc) oct. 2003 2 increase in distribution – crocodile river. although e. crassipes had previously been known to occur in the crocodile river, it had been absent since the floods of 2000. eichhornia crassipes (water hyacinth) integrated control using biological control at selected sites and aerial application of herbicides as dam walls and weirs long-term integrated control feb. 2004 1 new occurrence of a species in the knpcrocodile bridge harrisia martinii (moon cactus) chemical control and follow-up operations to ensure patch is eradicated. surveys to determine how far the plants have spread. landowner in adjacent property (source) to control the plants as well although the aim was eradication of the patch of plants, ongoing site visits will still be required feb. 2004 2 increase in distribution in the knp crocodile bridge bryophyllum delagoense (mother of millions) chemical control and follow-up operations to ensure patch is eradicated. surveys to determine how far the plants have spread. landowner in adjacent property (source) to control the plants as well although the aim was eradication of the patch of plants, ongoing site visits will still be required feb. 2004 2 increase in distribution in the knp crocodile bridge opuntia stricta (sour prickly pear) chemical control and follow-up operations to ensure patch is eradicated. surveys to determine how far the plants have spread. landowner in adjacent property (source) to control the plants as well although the aim was eradication of the patch of plants, ongoing site visits will still be required mar. 2004 1 & 2 tabling tpc of btb introduction and spread within the knp mycobacterium bovis (btb) ongoing monitoring of spread and prevalence per herd and region; detection of btb in other species; containment at the human / wildlife interface to develop long-term control methods july 2004 2 first ever record from a new area (crocodile river) pistia stratiotes (water lettuce) determine the extent of the invasion, survey for the presence of biocontrol agents and release agents if necessary long-term control using biological control agents july 2004 1 re-invasion of a controlled speciesmtshawu dam. this species was previously completely controlled by the biocontrol agents and absent from the dam as from late 2001 salvinia molesta (kariba weed) re-release of the biological control agents unlikely that eradication will take place. ongoing monitoring and if necessary re-release of the biological control agents, to ensure a long-term control programme dec. 2004 2 first ever record from a new area phabeni, sabie river thelechitonia trilobata (singapore daisy) chemical control ongoing chemical control operations to eradicate these patches from the sabie river, and prevent re-introduction from further upstream in the sabie river. ongoing site visits continue july 2005 2 first ever record from a new grid celltshokwane opuntia imbricata (imbricate cactus) chemical control of the plants and surveys to ensure any plants that have started to spread are also controlled eradication of this patch of plants. although the plants appear to have been eradicated, ongoing site visits will be required oct. 2005 2 first record in new grid cells (patches along the sabie river, just upstream from kruger gate) arundo donax (giant reed) plants to be manually controlled and coppice to be sprayed with an appropriate herbicide this invasion will require ongoing follow-up as part of routine operations, as it is unlikely that these patches will be eradicated date tpc level1 tpc detail species of concern management response expected outcome at the time of the tpc notification and current status online supplementary table 2 (cont...) developing thresholds of potential concern original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #157 3vol. 51 no. 1 page 3 of 3 oct. 2005 1 imminent external threat (a species on the boundary of the knp which might invade within the next 12 months)sabie river, hazyview area acacia decurrens (green wattle) mechanical / chemical control of the trees in hazyview. survey between the plants and the knp boundary although this patch can be eradicated, it is likely that there is a larger source somewhere in the upper catchment and continued vigilance is required sept. 2007 2 first record in new grid cell (small patch of plants near olifants tourist camp) cereus jamacaru (queen of the night) plants to be chemically controlled with an appropriate herbicide this small patch should be eradicated, but will require ongoing site visits sept. 2007 2 first ever records in a new grid cellolifants river opuntia stricta (sour prickly pear) chemical control and surveys to determine whether the plants have already spread eradication of the patches of plants, but longterm monitoring for re-growth and re-invasion from sources in the upper olifants river catchment 1level 1: tpcs that deal with new invasions of a species in the knp. level 2: tpcs that deal with an increase in distribution of a species (or all species combined) in the knp, over a 12 month period. date tpc level1 tpc detail species of concern management response expected outcome at the time of the tpc notification and current status online supplementary table 2 (cont...) abstract introduction research methods and design results discussion conclusions acknowledgements references about the author(s) carl f. reinhardt unit for environmental sciences and management, faculty of natural and agricultural sciences, north-west university, potchefstroom, south africa department of plant and soil sciences, faculty of natural and agricultural sciences, university of pretoria, pretoria, south africa hugo bezuidenhout nature conservation, scientific services, sanparks, kimberley, south africa applied behavioural ecology and ecosystem research unit, department of conservation, university of south africa, johannesburg, south africa judith m. botha scientific services, sanparks, skukuza, south africa citation reinhardt, c.f., bezuidenhout, h. & botha, j.m., 2022, ‘evidence that residues of tebuthiuron arboricide present in soil of mokala national park can be phytotoxic to woody and grass species’, koedoe 64(1), a1658. https://doi.org/10.4102/koedoe.v64i1.1658 original research evidence that residues of tebuthiuron arboricide present in soil of mokala national park can be phytotoxic to woody and grass species carl f. reinhardt, hugo bezuidenhout, judith m. botha received: 04 oct. 2020; accepted: 25 nov. 2021; published: 18 mar. 2022 copyright: © 2022. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract mokala national park (monp) has a history of arboricide use through south african national parks (sanps) having bought commercial game farmland for its establishment in 2007. tebuthiuron arboricide is known to have been applied for controlling bush densification during the period 1996 to 2004. persistent negative impacts on monp vegetation, which are ascribed to the historical arboricide use, have prompted this investigation from 2016 to 2017. bioassay experiments employing as test plants the tree species vachellia erioloba and vachellia tortilis, the shrub species senegalia mellifera and the grass tragus berteronianus were conducted in a glasshouse. growth responses of these species were assessed upon their exposure to a tebuthiuron concentration range that simulated expected levels in monp soil soon and long after application. chemical analysis as well as bioassay with the test species lycopersicon esculentum (tomato) were performed on soil samples collected from three depths (0−30, 30−60 and 60−90 cm) of the soil profile at two sites in monp where tebuthiuron was applied in the past. the three woody test species showed differential, negative growth response to tebuthiuron, and even growth of the grass species (t. berteronianus) was affected at the higher concentrations. evidence provided by the tomato bioassay and analysis performed on soil samples collected in situ points to the putative presence of tebuthiuron, more than a decade after the last use of arboricides for controlling bush densification. conservation implications: if the reported evidence of the presence of phytotoxic residue of tebuthiuron in soil of monp would be substantiated through further research, such findings could at least partly explain the failure of natural recruitment of vegetation in those areas where the woody component was degraded because of arboricide application more than a decade ago. keywords: arboricide; bush densification; ecosystems; environment; herbicide residues; problem plants; soil contamination. introduction it is generally accepted that dense stands of shrubs and trees can suppress herbaceous vegetation, especially grasses (harmse, kellner & dreber 2016; o’connor & crow 1999; smit 2005). the main concern about ‘densification’ of trees, or an undesirably dense woody component of vegetation, is in relation to reduction in grazing capacity and loss of land productivity in commercial farming settings. on game and livestock farms, in addition to its impact on grazing capacity, dense stands of the shrub senegalia mellifera (vahl) seigler & ebinger subsp. detinens (common name: black thorn; ‘swarthaak’) can be the bane of wildlife, livestock and humans alike, because of its ability to physically restrict access and mobility of humans and animals alike (ward 2005). thorns impart formidable defence against most herbivores, and this characteristic is also responsible for the notoriety of s. mellifera as an obstruction or hindrance on farmland and for hunters. of course, what is considered ‘undesirably dense’ or ‘problem plant’ by some (e.g. farmers and hunters) is a valuable food and ideal habitat for the black rhinoceros (buk & knight 2010) and other browsers. an increase in abundance of certain shrubs and trees is commonly experienced when land is dedicated to commercial production of the game and/or livestock (smit 2005) − the phenomenon is referred to as bush ‘densification’ or ‘thickening’ (la grange 2010). the terminology ‘bush encroachment’ is inter alia used by o’connor and crow (1999) to signify ‘invasion’ of grasslands and ‘thickening’ of savanna by indigenous woody species. its prolific growth habit is responsible for the classification of s. mellifera as ‘problem plant’ or ‘bush encroacher’, although it must be emphasised that such terminology with negative connotation is used in relation to human perceptions based on perceived threats to man’s aspirations. most savanna ecosystems in southern africa, and especially semi-arid savanna, such as that which predominates in mokala national park (monp), are water limited and, consequently, increased abundance of woody species typically results in the suppression of herbaceous layer (which include forbs and grass species) (harmse et al. 2016). s. mellifera has a shallow root system with extensive horizontal spread, in addition to a deep taproot, which makes it more competitive for water and nutrients than most other species, and hence explains its tendency to dominate plant communities of semi-arid savanna (adams 1967; donaldson 1967; smit 1999). reduction of grazing capacity is the inescapable consequence of unnatural bush densification that is brought on mainly by mismanagement of land by humans. to rectify both historical and more recent mistakes responsible for disruption of ecological balance in natural vegetation, the human solution is to institute control measures for the thinning or, even eradication, of the woody element. chemical and mechanical methods of control are basically the only options available to deal with bush encroachment and/or densification. mechanical control involves cutting and/or uprooting of plants, which is time-consuming and demands high levels of energy inputs, and effects are of limited duration in the case of cutting. the chemical option, which involves the use of arboricides, is by far the most economical in terms of both time and money. the glaring drawback of arboricides used for the control of woody species is that they lack selectivity in their effects on trees, shrubs and forbs, and they are also not innocuous towards grasses (du toit & sekwadi 2012; wssa 2014). basically, all dicotyledonous species and even grasses under certain conditions, particularly at the juvenile stage, will be susceptible to arboricides that are supposed to target only certain woody species. dosage of the arboricide, level of exposure and susceptibility of species are important determinants of eventual effects. selectivity of arboricides can be improved or directed through the way in which they are applied. application of arboricides by hand or by knap-sack sprayer is far more selective than aerial application by aircraft, but the latter method is far more economical than the former methods, and therefore, aerial application is the most common way in which arboricides are applied for the control of bush densification in south africa. an often overlooked or ignored negative characteristic of arboricides is their relatively long persistence in soil. persistence or half-life of arboricides in soil is typically measured in months and even years (wssa 2014), with dosage and frequency of use often the main determinants of their persistence in the environment. the department of environmental affairs (dea) publishes regulations pertaining to alien and invasive species (ais) in terms of the national environmental management biodiversity act (nemba), which was promulgated on 01 october 2014. regulation 16a of nemba deals with ‘indicators of bush encroachment’, that is indigenous species that are not a problem per se but are considered symptomatic (indicative) of poor land management. the nemba regulation 16a lists 62 indigenous tree and 16 shrub species as ‘indicators of bush encroachment’ (bromilow 2018). s. mellifera appears in the nemba regulation 16a list, and its control with specific herbicides (arboricides) is authorised in the following provinces: gauteng, limpopo, mpumalanga, northern cape, and north-west (bromilow 2018). arboricides that are currently registered in south africa for the control of s. mellifera are marketed under various product names, but all contain at least one of the following active ingredients: bromacil and tebuthiuron (croplife south africa 2016). it is on record that at least one tebuthiuron-containing product (molopo 200 gg) has apparently been used during the period 1996 to 2004 for the control of woody species (particularly s. mellifera) on a commercial game farm that was acquired by south african national parks (sanparks) to proclaim monp in 2007 (bezuidenhout, kraaij & baard 2015b). tebuthiuron is an urea-type, non-selective herbicide that controls broadleaf (including woody) plants at relatively low application rates and controls nearly all species at high rates. tebuthiuron (e.g. molopo 500 sc) is used for ‘bush control’ where woody species dominate. prior to 2003, that game farm was dedicated to livestock production. mokala national park is located (29° 10’ 20.7’’ s, 24° 21’ 00.5’’ e) in the northern cape province, about 80 km from the city of kimberley. at the time of it being a farming enterprise, the landowners had used chemical (arboricide) and mechanical methods for the control and, even eradication, of prolific shrubs (main target species s. mellifera) in certain areas on the farm during the period 1996 to 2004. the method of application of molopo 200 gg product varied among areas on the farm – plant-specific application was done at a dosage of 6 kg/ha while application by aeroplane was done at a dosage of 3 kg/ha. an unknown number of treatments were done on different occasions during the period 1996 to 2004, mostly in the spring (bezuidenhout et al. 2015b). bromacil and tebuthiuron arboricides have inherently long persistence in soil, and their half-life in soil is measured in months when applied once at recommended dosages, but when overdosing occurs or when used at high frequency and/or multiple applications, residues can remain biologically active in soil for several years (cox 1998; du toit & sekwadi 2012; moore, van niekerk & knight 1985; van eck & van lill 2008; wssa 2014). in the case of tebuthiuron, it is reported that phytotoxic residue was present in soil for at least 8 years following application of this arboricide for the control of the encroacher shrub seriphium plumosum (du toit & sekwadi 2012). residual effects of tebuthiuron are reported to be long-lasting; in areas that receive 1000 mm to 1500 mm of annual rainfall, the half-life of tebuthiuron is 12–15 months (wssa 2014). in areas with lower rainfall, the persistence can be longer. microbial action is primarily responsible for the degradation of tebuthiuron. a common characteristic of arboricides is their high molecular stability in the environment, and therefore, phytotoxic activity remains intact, that is persist, for months or even years, especially under dry soil conditions, near-neutral and alkaline soil ph, and low soil microbial activity (wssa 2014). those soil conditions are typical of semi-arid savanna ecosystems such as monp. rainfall in monp occurs mainly in the summer months, but it is erratic and drought a common occurrence. annual rainfall ranges from 304 mm to 622 mm (mean 415 mm) per annum. geographical features of monp include rocky hills, sandy plains with underlying calcrete, plains of exposed calcrete, degraded old lands and drainage lines (bezuidenhout et al. 2015b). the present study aimed to determine whether tebuthiuron arboricide residues could be responsible for the persistent impacts on vegetation, which were first reported by bezuidenhout et al. (2015b), in those areas of monp treated with tebuthiuron arboricide more than a decade ago. specific objectives were to (1) follow the bioassay approach for the detection of phytotoxic residues of tebuthiuron in different depths of the hutton soil form profile (soil classification working group 1991), (2) assess the response and relative susceptibility of two trees (protected vachellia erioloba and vachellia tortilis) and one shrub (s. mellifera) species to tebuthiuron and (3) determine how a grass species responds to tebuthiuron. research methods and design general mokala national park is situated about 70 km southwest of the city of kimberley in the northern cape province monp gps: 29° 15’ 5.19” s 24° 14’ 12.16” e to 28° 59’ 56.90” s 24° 23’ 8.29” e. the puntberg (pb) soil collection site is in the western section adjacent to puntberg (gps: 29° 7’ 41.93” s 24° 18’ 38.00” e). the soil is deep (> 2.0 m), well-drained, aeolian sand (hutton soil form) that was deposited on a hard calcrete layer which forms a water basin (srk 2009) for v. erioloba (camel thorn tree) as well as other associated plant species to grow in, the same habitat description applies for the windpomp (wp) site (gps: 29° 5’ 44.38” s 24° 22’ 23.65” e), which is situated on the northern side of puntberg (figure 1). figure 1: landscape units of mokala national park with locations of puntberg and windpomp soil sampling sites indicated experiments with potted soil collected from two locations (puntberg (pb) and windpomp (wp) in monp (refer map in figure 1) were conducted in a glasshouse at hatfield experimental farm, university. soil samples were collected in 2014 from the 0 cm – 30 cm, 30 cm – 60 cm and 60 cm – 90 cm depths in the soil profile at locations pb and wp and subjected to bioassay with various test species for determination of the presence of biologically active (phytotoxic) residues of tebuthiuron. the pb and wp locations were chosen for investigation because both locations occur in those areas of monp where it is known that tebuthiuron and other control methods were applied as described by bezuidenhout et al. (2015b). bezuidenhout et al. (2015a) describe both pb and wp habitat as deep (> 1.2 m) well-drained red sandy (less than 6% clay content in a-horizon and less than 15% in b-horizon) soil. it is restricted to the undulating plains, which are overlain by aeolian sand covering the dwyka formation. surface limestone occurs sporadically in this woodland and shrubland landscape units. no rocks or stones were noted on the soil surface. the dominant soil form was hutton, while clovelly and mispah soil forms were also found (soil classification working group 2015). at the pb site, the landscape is closely associated with the undulating plains of v. erioloba-v. tortilis open woodland. the wp site is on the interface between the two landscape units, namely the flat plains landscape unit of s. mellifera-v. erioloba open to closed woodland, while the other landscape unit is associated with slightly undulating footslopes/midslopes of s. mellifera–v. tortilis open shrubland (figure 1) (bezuidenhout et al. 2015b). selected soil properties of samples collected at pb are presented in table 1. soil analysis was done by intertek agricultural services, johannesburg, south africa. samples from wp site have similar properties as that of pb site, and therefore, only samples from the latter site were analysed for certain characteristics (personal communication: dr hugo bezuidenhout, sanparks, kimberley). table 1: selected properties of the hutton soil form samples collected from three depths (cm) of the soil profile at the puntberg location in mokala national park. (analyses were performed by intertek agricultural services). treatments (see next section for detail) in the bioassay experiments were replicated three times and pots were arranged in a completely randomised design. all statistical analyses were performed in r v.2.8.1. analyses of variance (anova) were used to detect significant differences between and within treatment groups. for chemical detection of tebuthiuron in all three depths of the soil profile that were sampled in 2014 at both locations, sub-samples were sent to the laboratory of rps mountainheath ltd, united kingdom, for analysis using chromatographic techniques (smith, shuman & lokey 1984; wells, michael & neary 1984). bioassay experiments tree and shrub test species the tree species v. tortilis, v. erioloba and the shrub species s. mellifera were grown from seed collected in monp. seedlings were transplanted into soil treated with tebuthiuron to achieve a concentration range that simulates extremely low to low residual concentrations, which conceivably, might be present in monp soil for several months, and even years, after application of recommended dosages of this tebuthiuron arboricide. soil used in this experiment had to be free of any arboricide residues, therefore collection was done from the 0 cm – 30 cm soil depth of a control area of the pb site where tebuthiuron application had never been done in the past (bezuidenhout et al. 2015b). because arboricides do not affect only woody species, it was decided to monitor the presence of any other plant species that would spontaneously establish on soil in this experiment. all other species that emerged and grew spontaneously with the three transplanted woody species would be exposed to the same range of tebuthiuron concentrations serving as treatments in this experiment. tebuthiuron was sourced from villa crop protection (pty) ltd in the form of the product tebuthiuron 500 sc. selection of the tebuthiuron dosage range used in this experiment was based on the following measurements, assumptions and calculations: specific weight of mokala soil = 1.5 g/cm3, therefore, 1 ha soil 10 cm deep (109 cm3) weighs 1.5 × 109 g, therefore, 1 m2 area soil 10 cm deep weighs 1.5 × 102 kg. molopo 500 sc label recommendation (500 g active ingredient l-1): stock solution: 1.5 l product in 8.5 l water (= 750 g in 10 l stock sol; = 75 mg/ml). apply 8 ml (4 × 2-ml dosage stock sol) at base of 2-m tall multi-stemmed tree, therefore, 8 × 75 mg = 600 mg tebuthiuron, therefore, assume 600 mg tebuthiuron is applied per 1 m2 soil surface area, and it is uniformly distributed 10 cm deep, therefore, 600 mg tebuthiuron per 1.5 × 102 kg soil, therefore, 1 kg soil contains 4 mg tebuthiuron [= 4 mg/kg (ppm) tebuthiuron in 1 m2 area, 10 cm deep]; [= 0.4 mg/kg tebuthiuron in 1 m2 area, 100 cm deep]. based on the above assumptions and calculations, tebuthiuron treatments applied on 06 february 2017 comprised the following dosage range: 0 mg/kg (untreated control), 0.005, 0.01, 0.05, 0.1, 0.5 mg/kg. treatment was done by applying predetermined volumes of 10 000-times diluted stock solution, which were prepared from the commercial product, in 80 ml total volume (made up with pure water) per pot containing 800 g soil. seed of v. erioloba, v. tortilis and s. mellifera were collected in monp on 08/09 january 2017. for promotion of germination, seeds of v. erioloba and v. tortilis was scarified on sanding paper prior to soaking in warm water overnight – temperature of water was 70 °c at immersion of seeds, with no further heating of water. seeds of s. mellifera was simply immersed overnight in water at room temperature (25°c – 28°c). following soaking in water, seeds were placed on, and covered with, moist paper towels to allow germination at room temperature. after 7 days, germination had occurred to the extent that it was possible to select germinated seeds from which the radicle extended at least 5 mm. seedlings were transplanted to pots containing 800 g monp soil sampled near the pb locality in an area that do not have a history of tebuthiuron application on 20 january 2017. water was added to soil in pots to attain field capacity level at the onset of the experiment. following treatment with tebuthiuron, total mass of each experimental unit (pot with soil) comprising pot, soil, seedlings and water was measured by weighing, and total mass recorded for each pot. subsequently, pots were weighed twice a week to replenish as accurately as was practically possible water lost through evaporation and transpiration. each week a total volume of 100 ml of a commercial complete nutrient solution (multifeed) was applied to each pot to prevent the risk of nutrient deficiencies confounding the results. the experiment was terminated when herbicide effects were judged to have reached maximum level. visual estimate of % damage of total leaves affected (expressed as percentage damage) caused by tebuthiuron was made with 2-week (14-day) intervals, which commenced 14 days after treatment (dat). the final measurement (visual assessment of damage expressed as percentage of the control) was done 56 days after herbicide application, and the experiment was terminated at 60 dat. statistical analysis employed the r v.2.8.1 programme. for the tree and shrub species, the amount of damage was compared across locations, species and soil depth at different treatment levels using the multifactor anova approach. common carrot-seed grass (tragus berteronianus) a single plant species was observed to establish spontaneously at all treatments in the experiment discussed above, namely the pioneer annual grass tragus berteronianus. its presence was monitored at different treatments and growth compared between tebuthiuron treatments and the control treatment (zero tebuthiuron). arboricide effect on this grass species was measured through visual assessment of growth reduction at treatments relative to growth at the control treatment. statistical analysis of data was done with the r v.2.8.1 programme. damage over time was compared between the different treatments with a simple anova. tomato (lycopersicon esculentum mill.) test species seed of cultivar heinz 1370 was sown on soil collected from three soil depths (0 m – 30 m, 30 m – 60 m and 60 cm – 90 cm) at both pb and wp locations where tebuthiuron had been applied in the past (bezuidenhout et al. 2015b). soil samples serving as untreated controls were extracted from the three soil depths by means of a soil auger at both sites in those areas where tebuthiuron was not applied at any stage in the past (control sites). six seeds were sown in each pot, and following emergence, seedlings were thinned to three per pot. seedling height and aboveground fresh biomass were measured at termination of the experiment, that is 30 days after sowing of seed. water and nutrient solution were applied in the same manner and frequency as that described above for the ‘tree and shrub test species’ experiment. data were expressed as mean plant height and biomass per pot prior to statistical analysis performed with r v.2.8.1. height and biomass were compared between locations and depths using a multifactor anova. chemical analysis for detection of arboricide residues in soil samples soil analysis was conducted by rps mountainheath ltd (http://www.rpsgroup.com), united kingdom, and was focused on tebuthiuron in all soil samples. analyses results are presented in table 2 in the following section. table 2: tebuthiuron presence in soil samples from different depths at puntberg and windpomp sites in mokala national park. (minimum detection level was set at 0.1 mg/kg). results bioassay experiments tree and shrub test species seedling growth response, which was expressed as estimated damage percentage based on visual comparison of tebuthiuron-treated seedlings of v. erioloba, v. tortilis and s. mellifera with their respective controls (untreated seedlings), appear in figure 2. averaged across tebuthiuron concentration, damage was already visible at 14 dat in the form of symptoms, which manifest as chlorosis on leaves, that are typical of a photosynthesis-inhibiting arboricide (tebuthiuron in this case). the severity of damage symptoms, that is, the intensity of chlorosis increased with tebuthiuron concentration and time. no damage symptoms caused by tebuthiuron were observed on any of the seedling types at 0.005 and 0.01 mg/kg concentrations for the duration of the experiment (56 dat). figure 2: growth reduction (% damage) calculated across a range of tebuthiuron concentrations that were established in soil collected from site puntberg in an area which did not have a history of tebuthiuron application. observations for which data are presented here were made 14 days after treatment (14 dat –‘day a’), 28 dat (‘day b’) and 56 dat (‘day c’) (erioloba – vachellia erioloba, mellifera – senegalia mellifera, tortilis – vachellia tortilis). over the 14 dat to 56 dat period s. mellifera and v. erioloba showed a significant increase in damage beyond 28 dat. v. tortilis showed a significant increase in damage earlier on between days 14 and 28. all three species showed a similar tendency for increased damage with time (figure 2). in all three species, leaf senescence and shedding of leaves followed the manifestation of chlorosis on leaves, but the former symptoms (leaf senescence) manifested only at the relatively high tebuthiuron concentrations of 0.1 mg/kg and 0.5 mg/kg. of the three species, across tebuthiuron concentration and time, v. tortilis tended to be the most susceptible to this arboricide, with v. erioloba and s. mellifera showing lower and similar tolerance to tebuthiuron. common carrot-seed grass (tragus berteronianus) the effect of tebuthiuron arboricide on the growth of this grass species could be assessed because of the presence of seeds of t. berteronianus occurring naturally in the soil collected for bioassay from monp. tebuthiuron effects, which were expressed as damage percentage, were measured by visual comparison of the grass’ growth and development at tebuthiuron treatments and the untreated control (figure 3). damage symptoms that were visible as leaf chlorosis and failure of seedling emergence were observed at tebuthiuron concentrations of 0.05 mg/kg, 0.1 mg/kg and 0.5 mg/kg, and symptoms increased significantly with an increase in concentration across this higher end of the concentration range. relatively low percentage damage was already observed at 14 dat and damage increased significantly over time. failure of seedling emergence was the common observation at the maximum concentration of the arboricide. figure 3: growth reduction (% damage) caused to the grass species tragus berteronianus by different tebuthiuron concentrations that were established in untreated (control) soil collected at site puntberg. observations for which data are presented here were made 14 days after treatment (14 dat – ‘day a’), 28 dat (‘day b’) and 56 dat (‘day c’). tomato (lycopersicon esculentum l.) test species in this experiment, the soil in bioassay was not treated with tebuthiuron or any other arboricide and was sourced from three depths (0 cm − 30 cm; 30 cm − 60 cm and 60 cm − 90 cm) in the soil profile of the two sites pb and wp that have a history of arboricide use, that is, tebuthiuron according to available information (bezuidenhout et al. 2015b). a third site near the aforementioned pb site does not have a history of arboricide application and consequently served as ‘control’, that is, soil not contaminated with arboricide residues. growth of tomato seedlings at different treatments was measured by harvesting and weighing of seedlings when the experiment was terminated. seedling weight in gram per pot (treatment combination) appears in figure 4. figure 4: growth response (seedling weight) of tomato cultivar heinz 1378 on soil collected from three depths (‘a’ = 0 cm – 30 cm; ‘b’ = 30 cm – 60 cm); ‘c’ = 60 cm – 90 cm) of the soil profile at three locations (puntberg-west; windpomp; untreated control). significant reduction in tomato weight measured for soil collected from both the 0 cm − 30 cm and 60 cm − 90 cm depths in the soil profile at both the pb and wp sites (figure 4) is indicative of the presence of a growth-inhibiting factor, such as putative phytotoxic residue(s) of an arboricide(s). tomato growth on soil from the 30 cm − 60 cm depth of both those sites was not significantly different from growth attained on control soil and therefore is indicative of lack of a growth-inhibiting factor(s) or to the presence of factors in too low amount to cause growth reduction in the test species. none of the soil characteristics presented in table 1 can explain the significant tomato growth reductions relative to the control treatments, which occurred on soil from specific depths, at both pb and wp sites with a history of arboricide application. chemical analysis for the presence of arboricide residues in soil samples the presence of tebuthiuron arboricide was revealed by chemical analysis in all three soil depths (0−30, 30−60 and 60−90 cm) at both sites (table 2). in all samples, the amount of tebuthiuron was found to be lower than the minimum detection level (mdl), which means it was detected but could not be accurately quantified. the mdl applied by the laboratory for these analyses, that is, 0.1 mg/kg, which is the same concentration as one of the tebuthiuron treatments that caused phytotoxic plant responses (damage symptoms) on test plants in the bioassays with tree, shrub and the grass species. discussion commercial products of arboricides such as molopo 200 gg, in which the active ingredient is tebuthiuron, represent one method (chemical) used for the control of woody species on the commercial livestock and game farm that was acquired by sanparks to establish monp in 2007. bezuidenhout et al. (2015b) document the history of arboricide applications and other vegetation management practices and contend that impacts of these treatments on vegetation in monp have persisted for more than a decade. according to bezuidenhout et al. (2015b), the arboricide tebuthiuron was applied in granular formulation (molopo 200 gg product) on said commercial farm in the period 1996 to 2004 for the control of the woody plant species s. mellifera and v. tortilis. because tebuthiuron is nonselective in terms of effect on all dicotyledonous species, more herbaceous and woody species would succumb to this arboricide than just the main target species, s. mellifera. in the present study, all three the woody species assessed, viz. v. tortilis, v. erioloba and s. mellifera, were susceptible (showed damage symptoms) to effects of tebuthiuron at 0.1 mg/kg and 0.5 mg/kg concentrations when exposed at the seedling stage. these concentrations could conceivably be relevant to tebuthiuron levels present in soil following application of recommended dosages of a product such as molopo 200 gg for ‘bush control’ or ‘controlling encroacher species’, that is, for killing mature woody plants. it should be considered that under natural conditions, over a prolonged period of months and even years, relatively low, non-phytotoxic arboricide concentrations in soil can be taken up by plants and progressively accumulate to phytotoxic amounts in the plant system and eventually cause damage to plants long after initial arboricide application. as an example of herbaceous species’ response to tebuthiuron, the tomato test species used in this study proved to be an effective indicator of the presence of phytotoxic residues of tebuthiuron in monp soil (figure 4). the bioassay technique that we used is generally accepted to be effective for determining the presence of plant-damaging (phytotoxic) residues of herbicides, including arboricides, in soil. the presence of bioactive residues of tebuthiuron in the 0 cm − 90 cm soil profile of the two sites monitored in monp was confirmed by chemical analysis (table 2). arboricides used for the control of woody species are generally considered to be selective towards grass species, that is, grass species are expected to be impervious to arboricides but then only under specific circumstances (du toit & sekwadi 2012). at dosage rates significantly higher than field-use recommended rates, arboricides can damage all types of vegetation, especially herbaceous and woody species, and even grasses can be damaged or killed when exposed to excessive amounts of arboricides (cobb & reade 2010). we observed that the grass species t. berteronianus could be severely affected by tebuthiuron depending on its concentration in soil of monp (figure 3), even to the extent that the emergence of seedlings is prevented. this points to the possibility that recruitment of grasses in general can be curtailed and even prevented when seeds are exposed to relatively high concentrations of an arboricide such as tebuthiuron. du toit and sekwadi (2012) report that tebuthiuron remains biologically active for at least 8 years after application in semi-arid grassland (cymbopogon-themeda veld type). they used the bioassay technique with oats (avena sativa) and cabbage (brassica oleracea) as indicator species for the presence of tebuthiuron in soil to explain the development of bare patches in the grass layer where this arboricide was applied for the control of the encroacher shrub seriphium plumosum. bauer (1978) found that tebuthiuron in either liquid or granular formulation prevented ryegrass (lolium perenne) establishment when applied at the 3.4 kg/ha dosage rate 261 days before planting of this grass pasture. picloram arboricide applied after the emergence of bermuda grass (cynodon dactylon) temporarily reduced the yield of this grass, and yield recovery only occurred the following season (bauer, bovey & holt 1977). generally, the grass component of areas that previously was dominated by dense stands of s. mellifera is reported to show good long-term recovery after the dominant woody species are killed by arboricides (moore et al. 1985), but there is a scant in-depth research on how different grass species respond to specific arboricides in semi-arid environments. just as woody species differ in their susceptibility to arboricides, grasses are likely to respond differently as well. bezuidenhout et al. (2015b) advise consideration of active rehabilitation measures if the natural recovery in the arboricide-treated areas of monp proves to be inadequate to ensure the conservation of ecological pattern and process. they contend that phytotoxic residues of tebuthiuron arboricide present in soil could be impacting vegetation recruitment even 10 years after this arboricide was applied. key edaphic factors that prolong the period for which arboricides can remain biologically active (phytotoxic) in soil are the following: low rainfall, near-neutral soil ph, low microbial activity, low clay content in soil profile and low organic carbon content of soil (wssa 2014). dosage and frequency of use remain key drivers of the persistence of arboricides in soil. the aforementioned long periods of impact following their use are likely to occur only if excessive dosages and/or high frequency of use were the historical practice. in the case of monp, it appears that the records of historical arboricide application and current contamination levels are incomplete, and only through further investigation can the real situation be revealed. we contend that only once such information is available can vegetation rehabilitation efforts have a high probability of being successful. conclusions although conclusive proof is lacking at this stage, both the bioassay approach and chemical analysis in the present investigation revealed the presence of phytotoxic residue of the arboricide tebuthiuron in soil samples collected from sites in monp with a history of application of this herbicide for controlling bush densification. effects of tebuthiuron observed on seedlings of v. erioloba, v. tortilis and s. mellifera suggest that recovery of the woody species following tebuthiuron application, especially at relatively high dosage rates and high frequency, for the control of mature plants could impede or even prevent the establishment of seedlings or saplings of those species for a significant period post-use. the establishment of grass species could also be impacted by tebuthiuron, as shown by the response of t. berteronianus in our bioassay experiment. soil and environmental conditions in monp favour the long persistence (half-life) of arboricides such as tebuthiuron and probably explain the apparent excessively long persistence of tebuthiuron in this setting. many key drivers of the long persistence of arboricides in soil are characteristic of semi-arid savanna areas of southern africa where arboricide use for controlling bush densification is often common practice, and unfortunately, excessive in terms of various arboricide combinations, high dosage rates and high frequency of application. for any rehabilitation effort of affected areas in monp to have a reasonable chance of success, further investigation on the nature and extent of soil contamination with arboricide residues is imperative. acknowledgements competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions c.r. and h.b. were responsible for conceptualisation. h.b. was the project leader and contributed to the sampling of soil and seeds in mokala national park. c.r. was the main writer and was responsible for the experiments. c.r. and h.b. organised the resources. j.b. conducted statistical analyses and produced graphs based on the data that was collected by c.r. and contributed towards describing the methods. all authors contributed to the final drafting of the manuscript. ethical considerations this article followed all ethical standards for research without direct contact with human or animal subjects. funding information sanparks and mokala national park management provided technical assistance for the field work, and provided accommodation for the researchers. the university of pretoria provided support in the form of glasshouse facilities, and villa crop protection (pty) ltd provided partial financial support. data availability data sharing is not applicable to this article as no new data were created or analysed in this study. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references adams, m.e., 1967, ‘a study of the ecology of acacia mellifera, a. seyal and balanites aegyptica in relation to land-clearing’, journal of applied ecology 4(1), 221–237. https://doi.org/10.2307/2401420 bauer, j.r., 1978, ‘effects of picloram and tebuthiuron on establishment on ryegrass winter pasture’, journal of range management 31(6), 450–455. https://doi.org/10.2307/3897205 bauer, j.r., bovey, r.w. & holt, f.c., 1977, ‘effect of herbicides on production and protein level in pasture grasses’, agronomy journal 69(5), 846–851. https://doi.org/10.2134/agronj1977.00021962006900050030x bezuidenhout, h., bradshaw, p.l., bradshaw, m. & zietsman, p.c., 2015a, ‘landscape units of mokala national park, northern cape province, south africa’, navorsinge van die nasionale museum bloemfontein 31(1), 1–27. bezuidenhout, h., kraaij, t. & baard, j., 2015b, ‘persistent effects of chemicals used to control shrub densification in semi-arid savanna’, earth science research 4(1), 31–39. https://doi.org/10.5539/esr.v4n1p31 bromilow, c., 2018, problem plants and invasive weeds of southern africa, briza publications, pretoria. buk, k.g. & knight, m.h., 2010, ‘seasonal diet preferences of black rhinoceros in three arid south african national parks’, african journal of ecology 48(4), 1064–1075. https://doi.org/10.1111/j.1365-2028.2010.01213.x cobb, a.h. & reade, j.p.h., 2010, herbicides and plant physiology, 2nd edn., wiley-blackwell, uk. cox, c., 1998, ‘picloram, herbicide fact sheet’, journal of pesticide reform 18(1), 13–20. croplife south africa, 2016, problem plant control compendium, avcasa, halfway house. donaldson, c.h., 1967, bush encroachment with special reference to the black thorn problem of the molopo area, department of agricultural technical services, armoedsvlakte research station, vryburg. du toit, j.c.o. & sekwadi, k.p., 2012, ‘tebuthiuron residues remain active in soil for at least eight years in a semi-arid grassland, south africa’, african journal of range & forage science 29(2), 85–90. https://doi.org/10.2989/10220119.2012.705325 harmse, c.j., kellner, k. & dreber, n., 2016, ‘restoring productive rangelands: a comparative assessment of selective and non-selective chemical bush control in a semi-arid kalahari savanna’, journal of arid environments 35, 39–49. https://doi.org/10.1016/j.jaridenv.2016.08.009 la grange, m., 2010, ‘vegetation classification of the proposed heritage park, north-west province, south africa’, msc thesis (environmental sciences), north-west university, potchefstroom. moore, a., van niekerk, j.p. & knight, i.w., 1985, ‘the effect of tebuthiuron on the vegetation of the thorn bushveld of the northern cape – a preliminary report’, journal of the grassland society of south africa 2(4), 7–10. https://doi.org/10.1080/02566702.1985.9648012 o’connor, t.g. & crow, v.r.t., 1999, ‘rate and pattern of bush encroachment in eastern cape savanna and grassland’, african journal of range and forage science 16(1), 26–31. https://doi.org/10.2989/10220119909485715 smit, g.n., 1999, guide to the acacias of south africa, briza publications, pretoria. smit, g.n., 2005, ‘tree thinning as an option to increase herbaceous yield of an encroached semi-arid savanna in south africa’, bmc ecology 5(4), 15. https://doi.org/10.1186/1472-6785-5-4 smith, a.e., shuman, l.m. & lokey, n., 1984, ‘high-pressure liquid chromatographic analysis of tebuthiuron in soil’, journal of agricultural food and chemistry 34, 416–418. https://doi.org/10.1021/jf00122a060 soil classification working group, 1991, ‘soil classification: a taxonomic system for south africa’, memoirs on the agricultural natural resources of south africa 15, 1–262. srk, 2009, ‘consulting engineers and scientists: mokala yield testing. internal report, park manager: mokala national park, south african national park. van eck, j.a.j. & van lill, c., 2008, ‘chemical bush control on rhigozum trichotomum and acacia nebrownii in southern namibia using different arboricides and control methods’, agricola 15, 51–55. ward, d., 2005, ‘do we understand the causes of bush encroachment in african savannas?’, african journal of range & forage science 22(2), 101–105. https://doi.org/10.2989/10220110509485867 weed science society of america (wssa), 2014, herbicide handbook, 10th edn., wssa, lawrence, ks. wells, m.j.m., michael, j.l. & neary, d.g., 1984, ‘determination of picloram in soil and water by reversed-phase liquid chromatography’, archives of environmental contamination and toxicology 13, 231–235. https://doi.org/10.1007/bf01055881 abstract introduction research method and design results discussion conclusion acknowledgements references appendix 1 about the author(s) charles r. haddad department of zoology & entomology, university of the free state, south africa vivian p. butler department of animal, wildlife and grassland sciences, university of the free state, south africa citation haddad, c.r. & butler, v.p., 2018, ‘ground-dwelling spider assemblages in contrasting habitats in the central south african grassland biome’, koedoe 60(1), a1482. https://doi.org/10.4102/koedoe.v60i1.1482 original research ground-dwelling spider assemblages in contrasting habitats in the central south african grassland biome charles r. haddad, vivian p. butler received: 28 july 2017; accepted: 09 apr. 2018; published: 31 may 2018 copyright: © 2018. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract background: ground-dwelling spider assemblages in shrublands and cultivated pastures in the south african grassland biome have never been comprehensively studied. objectives: epigeic spiders were collected in eight different habitats in the amanzi private game reserve in the free state to determine assemblages of different vegetation types. methods: three of the sampled habitats were contrasting low-lying shrublands; three were contrasting hill aspects (northern slope, southern slope and plateau) in the buddleja saligna–searsia burchellii–olea europaea africana subcommunity; one habitat was cultivated digitaria eriantha pastures, and the last habitat was an area in and around a freshwater dam. spiders were sampled by pitfall trapping in early spring (sept. 2012), mid-summer (jan. 2013), mid-autumn (apr. 2013) and mid-winter (july 2013). results: a total of 2982 adult spiders were collected, representing 129 species and 33 families. ammoxenidae was the most abundant family (40.85%), followed by gnaphosidae (21.26%), zodariidae (10.80%) and salticidae (10.26%). gnaphosidae was the most species-rich family (24.81%), followed by salticidae (13.18%), lycosidae (11.63%) and zodariidae (6.20%). spider activity densities and species richness did not differ significantly between habitats, although significant seasonal fluctuations were detected. the three hill aspects and cultivated d. eriantha pastures had the most distinct assemblages, while those of the three low-lying shrublands and freshwater dam showed considerable overlap. conclusions: our results indicate that the aspect of hills has a significant effect in shaping spider assemblages, while the vegetation composition of shrublands is not strongly influential. the unique spider assemblages of cultivated d. eriantha pastures can be attributed to the absence of woody plants. conservation implications: this was the first study to investigate ground-dwelling spider assemblages in shrublands and cultivated pastures in the south african grassland biome. our study confirms that hill aspects, shrublands and pastures harbour very different spider faunas. when identifying land for potential expansion or establishment of protected areas, conservation planners should ensure that the greatest diversity of vegetation units are included to optimise the conservation of biodiversity. introduction in africa, the grassland biome is largely limited to the central plateau of south africa, lesotho and parts of swaziland (mucina & rutherford 2006). it is characterised by extremely high plant biodiversity, second only to that of the fynbos biome (low & rebelo 1996). grasslands can be defined as a single-layered herbaceous plant community, with a few woody plants, which are usually restricted to specific habitats, including drainage lines and rocky hilltops (carbutt et al. 2011). the grassland biome is one of the most transformed biomes in south africa and is under continuous threat from cultivation, overgrazing and urban expansion (bredenkamp & van rooyen 1996). only an estimated 2.04% to 2.80% of this biome is formally conserved (carbutt et al. 2011; o’connor & kuyler 2005), and therefore, effective management and conservation of private land is critical to protect its highly endemic fauna and flora (wessels et al. 2003). although nearly 910 point localities have been sampled for spiders in south african grasslands, only 27 of these have more than 100 specimen records (foord, dippenaar-schoeman & haddad 2011). only as recently as three decades ago were the first ecological studies on spiders undertaken in the biome, focusing on the biodiversity of ground-dwelling (haddad et al. 2015; jansen et al. 2013; lotz, seaman & kok 1991; van den berg & dippenaar-schoeman 1991), plant-dwelling (dippenaar-schoeman, hamer & haddad 2011; fourie et al. 2013; haddad 2005; neethling & haddad 2013), litter-dwelling (butler & haddad 2011) and termitophilous assemblages (haddad & dippenaar-schoeman 2002, 2006). different species are adapted to particular microhabitats within grasslands, either living on the ground, on grasses or on foliage of woody plants, with few species abundant in multiple strata (haddad et al. 2013). consequently, there is considerable scope for research on spider biodiversity, ecology and biology in this unique biome. only three of the aforementioned studies have investigated finer scale differences in spider assemblages associated with different plant species, notably litter-dwellers (butler & haddad 2011) and foliage-dwellers (fourie et al. 2013; neethling & haddad 2013). however, assemblage structure in contrasting vegetation communities has so far only been investigated for grass-dwellers in structurally variable grasslands (fourie et al. 2013). all the pitfall trapping surveys listed above were conducted in open grasslands with sparse or absent woody vegetation, and spider assemblages in shrublands and woodlands remain largely unknown in this biome (butler & haddad 2011). the aims of this study were (1) to sample ground-dwelling spider assemblages in different plant communities (predominantly shrublands) to determine possible habitat associations of spider species; (2) assess how habitats affected the activity density and species richness of spiders; (3) determine seasonality of ground-dwelling spider assemblages, and (4) determine whether indicator species could be identified for any of the sampled habitats. further, this study aims to add to the current knowledge base of ground-dwelling spider biodiversity in south africa, for which relatively little information is currently available (dippenaar-schoeman et al. 2015; janion-scheepers et al. 2016). research method and design study area amanzi private game reserve (apgr) is located about 80 km north-east of bloemfontein in the central free state (figure 1a) and falls within the summer rainfall region of central south africa, with an average of approximately 475 mm of rainfall received annually (butler 2017). the area experiences hot summers, with day temperatures sometimes between 35 °c and 40 °c (averaging above 30 °c), and cold winters, with night temperatures frequently below freezing and day temperatures usually ranging between 15 °c and 20 °c (butler 2017). figure 1: (a) location of the amanzi private game reserve, brandfort district, south africa (star), north-east of bloemfontein (circle) and (b) vegetation map of the western section of the amanzi private game reserve. the study area is located in the grassland biome (rutherford & westfall 1994), with the vegetation of the surrounding area being described as dry sandy highveld grassland (bredenkamp & van rooyen 1996) or vaal-vet sandy grassland (mucina & rutherford 2006). the vegetation in apgr is, however, more representative of winburg grassy shrubland, which occurs in a series of larger patches from trompsburg through bloemfontein and winburg to ventersburg (mucina & rutherford 2006). the landscape of this vegetation type consists of isolated hills, slopes and escarpments, creating habitats ranging from open grassland to shrubland (mucina & rutherford 2006). a comprehensive vegetation survey was conducted in the western section of apgr, and details of the community composition in this part of the reserve are provided in butler (2017). based on the results of the vegetation survey, spiders were collected in eight different habitats representing four plant communities (table 1; figure 1b). in the buddelja saligna–searsia burchellii community, two different subcommunities were sampled, one of which included three habitats associated with the northern and southern slopes and plateau of a hill (buddleja saligna–searsia burchellii–olea europaea africana subcommunity), and the second including two habitats, one dominated by searsia burchellii and the other by tarchonanthus camphoratus (buddleja saligna–searsia burchellii–vachellia karroo subcommunity). the remaining three habitats sampled were the themeda triandra–digitaria eriantha community, which was dominated in the woody layer by vachellia karroo; cultivated d. eriantha pastures (digitaria eriantha–cynodon dactylon community); and an area in and around a freshwater dam (persicaria lapathifolia–panicum coloratum community). the latter two communities had little or no plants contributing to the woody layer (table 1). table 1: detailed description of plant communities in each of the habitats sampled in the amanzi private game reserve from september 2012 to august 2013. spider sampling two sites were sampled in each habitat, with at least 100 m separating them to avoid pseudoreplication. five pitfall traps were placed 5 m apart in a straight line at each site. a soil auger was used for drilling holes, and plastic buckets 10 cm in diameter were used as pitfall traps. ethylene glycol (100 ml) was added to each pitfall trap to preserve terrestrial arthropods for later identification. collected material was removed and the pitfall traps refilled at the end of each month, from the start of september 2012 (early spring) to the end of august 2013 (end of winter). the sampled material was sorted in the laboratory and all arachnids were extracted from the samples and preserved in 70% ethanol. following identification and tallying of adult spiders, material was deposited in the national collection of arachnida at the arc–plant protection research in pretoria, south africa. although spiders were sampled for a full year, several sampling months represented incomplete sampling efforts as a result of flooding of the pitfall traps at some sites following heavy rains, or damage caused to pitfall traps by large herbivorous mammals or vervet monkeys. to ensure that sampling effort was equal between habitats, we opted to provide data for 1 month in each season for which all sites had a complete sampling effort: early spring (september 2012), mid-summer (january 2013), mid-autumn (april 2013) and mid-winter (july 2013). only adult spiders were included in the analysis. statistical analysis we calculated the estimated species richness for each habitat using the equation: where the number of species represented by a single individual (i.e. singletons) and two individuals only (i.e. doubletons) are represented by f1 and f2, respectively (chao 1984). chao1 is an estimator calculated using the available abundance data and is a function of the ratio between the singletons and doubletons in the data. with an increase in the number of samples, an accumulation curve reaches an asymptote when all species in the community are represented by at least two individuals. we calculated sampling completeness as the ratio of the observed species richness (sobs) and the chao1-estimated species richness (schao1) (e.g. cardoso et al. 2008; scharff et al. 2003; sørensen, coddington & scharff 2002). chao and jost (2012) proposed the use of coverage-based rarefaction and extrapolation to assess community richness and sampling effort. they defined sample coverage as the proportion of the total number of individuals in a community that belong to the species represented in the sample. subtracting the sample coverage from unity gives the proportion of the community belonging to as yet unsampled species, which they referred to as the ‘coverage deficit’. this can be inferred as the likelihood that a new, previously unsampled species will be found if the sample was increased by one individual (chao & jost 2012). coverage for each habitat and for the total spider assemblage was calculated using the following equation: where n represents the number of individuals in the sample and f1 and f2 represent the number of singleton and doubleton species, respectively. chao and lee (1992) proposed that an estimated coverage value should be at least 50%, that is, 0.5. inventory completeness was analysed in paleontological statistics (past) version 2.07 using the sample rarefaction function, which implements the ‘mao tau’ analytical procedure, with standard errors indicated as 95% confidence intervals on the resulting graphs. we produced curves for each of the habitats, as well as the for the whole spider assemblage. using past versions 2.07 and 3.06 (hammer, harper & ryan 2001), we calculated whether the abundance and species richness of spiders differed between the eight habitats and also whether these parameters varied seasonally, using linear multivariate regression. in each analysis, habitats and samples were used as independent variables. because count data follows a poisson distribution, values were log-transformed prior to analysis to approach a normal distribution. we then determined whether the two sites from each habitat sampled similar spider assemblages by performing a two-dimensional non-metric multidimensional scaling (nmds), based on a bray–curtis similarity matrix. ideally, the stress value of an nmds should be lower than 0.2; otherwise, the resulting diagram needs to be interpreted with caution (clarke 1993). further, we performed a cluster analysis using the cluster function and unweighted pair-group average algorithm (clarke & warwick 2001). in both analyses, the pooled data from each of the 16 sampling sites were used. further, a permutational multivariate analysis of variance (permanova) was performed to test for differences in assemblages between the eight habitats, using the bray–curtis distance measure and 10 000 permutations. these analyses were carried out in past version 2.07. lastly, we attempted to identify indicator spider species, which are considered to be characteristic of a particular habitat. indicator values were obtained by multiplying a species’ relative abundance in a particular habitat, expressed as a percentage of its total abundance, with its relative frequency of occurrence in that particular habitat, that is, proportion of samples in which a species was collected (dufrene & legendre 1997). thus, a species’ specificity (narrow association with a particular habitat) and fidelity (frequency of occurrence in that habitat) is expressed as a percentage that can be compared with other species in the sampled habitats (dufrene & legendre 1997). a high indicator value illustrates a high affiliation of a species to a particular habitat, with a suitable benchmark of 70% being suggested (e.g. haddad et al. 2010; mcgeoch, van rensburg & botes 2002; van rensburg et al. 1999). ethical consideration permission to collect arachnids in the free state province was obtained from the free state department of economic development, tourism and environmental affairs. results a total of 2982 adult spiders were collected, representing 129 species and 33 families (table 2; appendix 1). ammoxenidae was the dominant family (n = 1218, 40.85%), followed by gnaphosidae (n = 634, 21.26%), zodariidae (n = 322, 10.80%) and salticidae (n = 306, 10.26%). ammoxenus amphalodes dippenaar & meyer, 1980 strongly dominated the fauna overall (n = 1218, 40.85%), largely because of its extremely high activity densities in the cultivated d. eriantha pastures, where it represented 66.15% of the fauna. ranops sp. (7.38%) was the second most abundant species. other common species include proevippa sp. 1 (4.33%), phlegra karoo wesołowska, 2006 (3.66%), zelotes sclateri tucker, 1923 (3.66%) and drassodes sp. 2 (3.29%). table 2: family activity density and species richness of ground-dwelling spiders sampled by pitfall trapping from eight habitats in the amanzi private game reserve. gnaphosidae was the most species-rich family (32 spp., 24.81%), followed by salticidae (17 spp., 13.18%), lycosidae (15 spp., 11.63%) and zodariidae (8 spp., 6.20%). total species richness was quite similar between habitats, ranging between 34 and 48 species. however, chao1-estimated species richness varied considerably, between 58 and 106 species per habitat, with the total ground-dwelling assemblage in the sampled habitats estimated at 167 species (table 3). although coverage values were above 0.85 for all the habitats, sample completion was much more variable (0.41–0.76), with 0.77 for the total assemblage (table 3). this pattern was confirmed by the sample rarefaction curve for the whole spider assemblage (figure 2a), which approached an asymptote but did not level out, indicating that the majority of the ground-dwelling species in the sampled habitats had been collected. however, none of the habitats’ rarefaction curves approached an asymptote (figures 2b–i), indicating that a considerable portion of the fauna of each was still unsampled. because open grasslands were not sampled in this study, it could be expected that the total species richness at apgr may further exceed this projected value of 167 species. figure 2: sample rarefaction curves (mao tau) with 95% confidence intervals for ground-dwelling spiders sampled in the amanzi private game reserve from september 2012 to august 2013: (a) all habitats combined; (b) searsia burchellii closed evergreen shrubland; (c) cultivated digitaria eriantha pastures; (d) freshwater dam; (e) olea europaea closed evergreen shrubland on northern slopes of hill; (f) o. europaea closed evergreen shrubland on plateau of hill; (g) o. europaea closed evergreen shrubland on southern slope of hill; (h) tarchonanthus camphoratus closed evergreen shrubland; and (i) vachellia karroo open deciduous shrubland. table 3: summary of actual and estimated species richness of ground-dwelling spiders sampled by pitfall trapping from eight habitats in the amanzi private game reserve. nearly half of all the spiders (n = 1480, 49.63%) were collected from the two cultivated d. eriantha pasture sites; indeed, spider numbers in this habitat were exactly five times higher than the habitat with the second highest activity densities, searsia burchellii closed evergreen shrubland (n = 296). however, linear multivariate regression showed no significant differences between habitats in log-transformed spider activity densities (f2,61 = 1.125, p = 0.3313) or species richness (f2,61 = 0.3105, p = 0.7343). linear multivariate regression showed that there was significant seasonality across all sites in log-transformed spider activity densities (f2,61 = 8.388, p = 0.0006), with summer and spring activity densities clearly much higher than those of the colder seasons (figure 3a). species richness showed a similar pattern (f2,61 = 13.72, p < 0.0001), although spring and summer species richness was very similar (figure 3b), although markedly higher than autumn and winter species richness. figure 3: seasonal variation in mean ± sd of (a) activity densities and (b) species richness of ground-dwelling spiders across 16 sites (eight habitats) sampled in the amanzi private game reserve. spider assemblages showed some interesting patterns. not surprisingly, the assemblages of the cultivated d. eriantha pasture sites were the most distinct and showed the greatest similarity to one another compared to all the other habitat site-pairs. among the remaining seven habitats, only the three habitats associated with the hill had their paired sites grouping together (figures 4a, b). there was considerable overlap in the assemblages of the lower lying shrubland types and the dam, with most site-pairs not grouping close together (figure 4b). these results were supported by the permanova analysis, which showed highly significant differences between the spider assemblages in the sampled habitats (pseudo-f = 2.614, p < 0.0001). particularly, the pair-wise post hoc comparisons showed significant differences in assemblages between the cultivated pastures (p < 0.0007), plateau (p < 0.0128) and southern slope (p < 0.0105) and all the other habitats (table 4). for the other habitats, several other paired habitats were also significantly different from each other, although the assemblages of the freshwater dam and searsia burchellii closed evergreen shrubland had the fewest significant paired values, confirming their assemblage overlap with other habitats (table 4). figure 4: (a) non-metric multidimensional scaling plot and (b) cluster analysis dendrogram using unweighted pair-group average algorithm of ground-dwelling spider assemblages sampled from eight habitats in the amanzi private game reserve. analyses are based on a bray–curtis dissimilarity matrix using untransformed data. habitat abbreviations: b, searsia burchellii closed evergreen shrubland; c, cultivated digitaria eriantha pastures; d, freshwater dam; n, olea europaea closed evergreen shrubland on northern slopes of hill; p, o. europaea closed evergreen shrubland on plateau of hill; s, o. europaea closed evergreen shrubland on southern slope of hill; t, tarchonanthus camphoratus closed evergreen shrubland; v, vachellia karroo open deciduous shrubland. table 4: post hoc p-values calculated from permutational multivariate analysis of variance analysis of spider assemblages associated with eight habitats in the amanzi private game reserve. of the eight habitats sampled, indicator species with a percentage value > 70% were only found in one of the habitats, with a. amphalodes (80.38%) and ranops sp. (77.95%) both being indicator species for the cultivated d. eriantha pastures. of the remaining species, only three had indicator values > 50.00%: setaphis browni (tucker, 1923) (62.50%) in the northern slope sites, diores poweri tucker, 1920 (66.18%) in the plateau sites and proevippa sp. 1 (58.72%) for the southern slopes. discussion previous studies on ground-dwelling spiders in the grassland biome have focused on open grasslands, while the shrubland faunas have received little attention (butler & haddad 2011). in this study, the first focused on shrubland habitats and cultivated pastures in the grasslands of central south africa, the hill-associated habitats and cultivated d. eriantha pastures had the most distinct assemblages, while those of the lower lying shrublands and freshwater dam showed considerable overlap and often lacked clear distinction. the sample completion values varied between 0.41 and 0.76 for each habitat, which suggests that further sampling is necessary for a better representation of the species richness of those habitats with sample completion < 0.5. however, had the data from the other months sampled, but not included in this analysis, been incorporated into this study, then it is quite likely that this threshold value would have been exceeded in all the habitats. our results indicate that the aspect of hillside habitats has a strong influence in shaping assemblages. this could be because of the northern slopes getting more exposure to direct sunlight compared to the southern slopes. southern slopes have denser vegetation cover in the woody layer, with especially olea europaea africana providing a lot more shade. assemblages of the plateau are also very unique, which can possibly be attributed to the differences in the herbaceous layer and slope of the habitat. we would propose conducting a large-scale survey of various hills in central south africa to determine whether each aspect of these hills (including east and west that were not sampled here) contains a distinct assemblage and whether this pattern varies geographically. the assemblages of cultivated d. eriantha pastures were very unique, largely because of the absence of woody plants. although studies in open grasslands in south africa have shown very contrasting patterns of family dominance (haddad et al. 2015; jansen et al. 2013; lotz et al. 1991; van den berg & dippenaar-schoeman 1991), the dominance of ammoxenidae in the cultivated d. eriantha pastures studied here is quite extreme (66.15%). this could be attributed to the high abundance of hodotermes mossambicus (hagen, 1853) termites in the pastures, which are the sole prey of a. amphalodes (petráková et al. 2015). similarly, the high activity of ranops sp. in this habitat (13.24%) can be attributed to high activity densities of their mimetic model and possible prey ant, anoplolepis custodiens (f. smith, 1858) (haddad 2012). interestingly, these were the only two spider species with indicator values above 70.0%, suggesting that the other species sampled had more general habitat preferences, were more strongly seasonal in occurrence or were too scarce to serve as meaningful indicators of particular habitats. the ten species of mygalomorph trapdoor spiders collected in four months’ sampling (~9600 pitfall trap-days) is quite remarkable. this is higher than the eight species collected during nearly 22 000 pitfall trap-days’ sampling in open grasslands in the erfenis dam nature reserve (haddad et al. 2015) and the five species collected in open grassland during 36 500 trap-days’ sampling in the free state national botanical gardens (j.a. neethling & c.r. haddad [university of the free state] unpubl., august 2010 to may 2011), both within a radius of 70 km from amanzi. this is probably because of the more structurally and topographically variable habitats sampled in the current study compared to open grasslands sampled in the latter two studies. the inclusion of open grassland habitats in the current study would likely have increased the trapdoor spider diversity at this site. for example, calommata meridionalis fourie, haddad and jocqué, 2011 (atypidae) was not collected in this study but has been recorded from the other two sites (fourie et al. 2011; haddad et al. 2015). in addition, an unidentified harpactira sp. (theraphosidae) has also been collected from burrows and/or at night at all three localities, but has yet to be sampled using pitfall trapping. two faunistic records are of particular interest: the tree-trapdoor genus moggridgea (migidae) is recorded from the free state province for the first time (dippenaar-schoeman 2002; dippenaar-schoeman et al. 2010; griswold 1987), with five male spiders sampled in very contrasting habitats (appendix 1). this suggests that this species may occur in shrublands throughout central south africa. this study also yielded the first records of opiliones from the central free state (assamiidae: polycoryphus asper loman, 1902); all previous records of harvestmen in the province are from the eastern or southern fringes (lotz 2002). previously, this species was only known from the kogelbeen caves in northern cape, port elizabeth in the eastern cape and on the namibia-angola border, and is thus widespread although scarce (lotz 2009). although only two specimens were collected in pitfall traps from the southern slope sites in this study, more than 20 additional specimens were collected from beneath large rocks on various hills at amanzi, all on the southern slopes. further studies are needed elsewhere in central south africa to clarify the distribution and microhabitat preferences of these two arachnids. conclusion this is the first study to investigate spider assemblages in shrubland, hill and pasture habitats in the grassland biome of the free state, south africa. our results indicate that activity densities of spiders are lower in shrubland habitats than cultivated pastures and open grasslands previously sampled in central south africa. shrublands accommodate very different assemblages to pastures and grasslands, and therefore, conservation efforts for arachnids will benefit considerably from sampling a broader habitat diversity to identify potential indicator species and species of potential conservation importance. acknowledgements kobie fourie is thanked for permission to conduct this study at amanzi private game reserve and for providing accommodation during the field work to the second author. ansie dippenaar-schoeman (arc – plant protection research institute, pretoria) and leon lotz (national museum, bloemfontein) are thanked for identifications of some of the species. the free state department of economic development, tourism and environmental affairs is thanked for collecting permits that made the study possible. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions c.r.h. performed initial sorting of arthropod material, identified the arachnid material, performed the statistical analysis and wrote part of the manuscript. v.p.b. performed the vegetation analysis, conducted the field work, assisted with initial sorting of arthropod material and wrote part of the manuscript. funding information this study was funded through a grant to the senior author from the national research foundation of south africa in the thuthuka programme (#69017). references bredenkamp, g.j. & van rooyen, n., 1996, ‘dry sandy highveld grassland’, in a.b. low & a.g. rebelo (eds.), vegetation of south africa, lesotho and swaziland, p. 41, department of environmental affairs and tourism, pretoria. butler, v.p., 2017, ‘feeding ecology of the greater kudu (tragelaphus strepsiceros) in the central free state’, unpublished m.sc dissertation, university of the free state, bloemfontein. butler, v.p. & haddad, c.r., 2011, ‘spider assemblages associated with leaf litter of three tree species in central south africa (arachnida: araneae)’, african journal of ecology 49, 301–310. https://doi.org/10.1111/j.1365-2028.2011.01265.x carbutt, c., tau, m., stephens, a. & escott, b., 2011, ‘the conservation status of temperate grasslands in southern africa’, grassroots 11, 17–23. cardoso, p., scharff, n., gaspar, c.s., henriques, s.s., carvalho, r., castro, p.h. et al., 2008, ‘rapid biodiversity assessment of spiders (araneae) using semi-quantitative sampling: a case study in a mediterranean forest’, insect conservation and diversity 1, 71–84. https://doi.org/10.1111/j.1752-4598.2007.00008.x chao, a., 1984, ‘nonparametric estimation of the number of classes in a population’, scandinavian journal of statistics 11, 265–270. chao, a. & jost, l., 2012, ‘coverage-based rarefaction and extrapolation: standardizing samples by completeness rather than size’, ecology 93, 2533–2547. https://doi.org/10.1890/11-1952.1 chao, a. & lee, s.m., 1992, ‘estimating the number of classes via sample coverage’, journal of the american statistical association 87, 210–217. https://doi.org/10.1080/01621459.1992.10475194 clarke, k.r., 1993, ‘non-parametric multivariate analyses of changes in community structure’, australian journal of ecology 18, 117–143. https://doi.org/10.1111/j.1442-9993.1993.tb00438.x clarke, k.r. & warwick, r.m., 2001, change in marine communities: an approach to statistical analysis and interpretation, 2nd edn., primer-e, plymouth. dippenaar-schoeman, a.s., 2002, baboon and trapdoor spiders of southern africa: an identification manual, agricultural research council, pretoria. dippenaar-schoeman, a.s., haddad, c.r., foord, s.h., lyle, r., helberg, l., mathebula, s. et al., 2010, first atlas of the spiders of south africa (arachnida: araneae), arc – plant protection research institute, pretoria, pp. 1158. dippenaar-schoeman, a.s., haddad, c.r., foord, s.h., lyle, r., lotz, l.n. & marais, p., 2015, ‘south african national survey of arachnida (sansa): review of current knowledge, constraints and future needs for documenting spider diversity (arachnida: araneae)’, transactions of the royal society of south africa 70, 245–275. https://doi.org/10.1080/0035919x.2015.1088486 dippenaar-schoeman, a.s., hamer, m. & haddad, c.r., 2011, ‘spiders (arachnida: araneae) of the vegetation layer of the mkambati nature reserve, eastern cape, south africa’, koedoe 53(1), 1058. https://doi.org/10.4102/koedoe.v53i1.1058 dufrene, m. & legendre, p., 1997, ‘species assemblages and indicator species: the need for a flexible asymmetrical approach’, ecological monographs 67, 345–366. https://doi.org/10.2307/2963459 foord, s.h., dippenaar-schoeman, a.s. & haddad, c.r., 2011, ‘south african spider diversity: african perspectives on the conservation of a mega-diverse group’, in o. grillo & g. venora (eds.), changing diversity in changing environment, pp. 163–182, intech publishing, rijeka. fourie, r., haddad, c.r., dippenaar-schoeman, a.s. & grobler, a., 2013, ‘ecology of the plant-dwelling spiders (arachnida: araneae) of the erfenis dam nature reserve, south africa’, koedoe 55(1), 1113. https://doi.org/10.4102/koedoe.v55i1.1113 fourie, r., haddad, c.r. & jocqué, r., 2011, ‘a revision of the purse-web spider genus calommata lucas, 1837 (araneae: atypidae) in the afrotropical region’, zookeys 95, 1–28. https://doi.org/10.3897/zookeys.95.745 griswold, c.e., 1987, ‘the african members of the trap-door spider family migidae (araneae: mygalomorphae) 1: the genus moggridgea o. p.-cambridge, 1875’, annals of the natal museum 28, 1–118. haddad, c.r., 2005, ‘ecology of spiders (arachnida: araneae) inhabiting themeda triandra forskål grassland in semi-arid south africa’, navorsinge van die nasionale museum, bloemfontein 21, 25–36. haddad, c.r., 2012, ‘advances in the systematics and ecology of african corinnidae spiders (arachnida: araneae), with emphasis on the castianeirinae’, unpublished phd thesis, university of the free state, bloemfontein. haddad, c.r. & dippenaar-schoeman, a.s., 2002, ‘the influence of mound structure on the diversity of spiders (araneae) inhabiting the abandoned mounds of the snouted harvester termite trinervitermes trinervoides (sjöstedt)’, journal of arachnology 30, 403–408. https://doi.org/10.1636/0161-8202(2002)030[0403:tiomso]2.0.co;2 haddad, c.r. & dippenaar-schoeman, a.s., 2006, ‘spiders (arachnida: araneae) inhabiting abandoned mounds of the snouted harvester termite trinervitermes trinervoides (sjöstedt) (isoptera: termitidae: nasutitermitinae) in the free state, with notes on their biology’, navorsinge van die nasionale museum, bloemfontein 22, 1–15. haddad, c.r., dippenaar-schoeman, a.s., foord, s.h., lotz, l.n. & lyle, r., 2013, ‘the faunistic diversity of spiders (arachnida: araneae) of the grassland biome in south africa’, transactions of the royal society of south africa 68, 97–122. https://doi.-org/10.1080/0035919x.2013.773267 haddad, c.r., foord, s.h., fourie, r. & dippenaar-schoeman, a.s., 2015, ‘effects of a fast-burning spring fire on the ground-dwelling spider assemblages (arachnida: araneae) in a central south african grassland habitat’, african zoology 50, 281–292. https://doi.org/10.1080/15627020.2015.1088400 haddad, c.r., honiball, a.s., dippenaar-schoeman, a.s., slotow, r. & van rensburg, b.j., 2010, ‘spiders (arachnida: araneae) as indicators of elephant-induced habitat changes in the maputaland centre of endemism, south africa’, african journal of ecology 48, 446–460. https://doi.org/10.1111/j.1365-2028.2009.01133.x hammer, ø., harper, d.a.t. & ryan, p.d., 2001, ‘past – palaeontological statistics’, palaeontologia electronica 4, 1–33. janion-scheepers, c., measey, j., braschler, b., chown, s.l., coetzee, l., colville, j. et al., 2016, ‘soil biota in a megadiverse country: current knowledge and future research directions in south africa’, pedobiologia 59, 129–174. https://doi.org/10.1016/j.pedobi.2016.03.004 jansen, r., makaka, l., little, i.t. & dippenaar-schoeman, a.s., 2013, ‘response of ground-dwelling spider assemblages (arachnida, araneae) to montane grassland management practices in south africa’, insect conservation and diversity 6, 572–589. https://doi.org/10.1111/icad.12013 lotz, l.n., 2002, ‘the opiliones (arachnida) of the free state province, south africa’, navorsinge van die nasionale museum, bloemfontein 18, 161–188. lotz, l.n., 2009, ‘harvestmen (arachnida: opiliones) in southern africa – an annotated catalogue with notes on distribution’, navorsinge van die nasionale museum, bloemfontein 25, 1–46. lotz, l.n., seaman, m.t. & kok, d.j., 1991, ‘surface-active spiders (araneae) of a site in semi-arid central south africa’, navorsinge van die nasionale museum, bloemfontein 7, 530–540. low, a.b. & rebelo, a.g., 1996, vegetation of south africa, lesotho and swaziland, department of environmental affairs and tourism, pretoria. mcgeoch, m.a., van rensburg, b.j. & botes, a., 2002, ‘the verification and application of bioindicators: a case study of dung beetles in a savanna ecosystem’, journal of applied ecology 39, 661–672. https://doi.org/10.1046/j.1365-2664.2002.00743.x mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, strelitzia 19, south african national biodiversity institute, pretoria. neethling, j.a. & haddad, c.r., 2013, ‘arboreal spider assemblages associated with four tree species in the grassland biome of central south africa (arachnida: araneae)’, transactions of the royal society of south africa 68, 123–131. https://doi.org/10.1080/0035919x.2013.806374 o’connor, t.g. & kuyler, p., 2005, national grasslands initiative: identification of compatible land uses for maintaining compatible biodiversity integrity, unpublished report, south african national biodiversity institute, pretoria. petráková, l., líznarová, e., pekár, s., haddad, c.r., sentenská, l. & symondson, w.o.c., 2015, ‘discovery of a monophagous true predator, a specialist termite-eating spider (araneae: ammoxenidae)’, scientific reports 5, 14013. https://doi.org/10.1038/srep14013 rutherford, m.c. & westfall, r.h., 1994, ‘biomes of southern africa: an objective characterization’, memoirs of the botanical survey of south africa 6, 1–94. scharff, n., coddington, j.a., griswold, c.e., hormiga, g. & bjorn, p., 2003, ‘when to quit? estimating spider species richness in a northern european deciduous forest’, journal of arachnology 31, 246–273. https://doi.org/10.1636/0161-8202(2003)031[0246:wtqess]2.0.co;2 sørensen, l.i., coddington, j.a. & scharff, n.j., 2002, ‘inventorying and estimating subcanopy spider diversity using semiquantitative sampling methods in an afromontane forest’, environmental entomology 31, 319–330. https://doi.org/10.1603/0046-225x-31.2.319 van den berg, a. & dippenaar-schoeman, a.s., 1991, ‘ground-living spiders from an area where the harvester termite hodotermes mossambicus occurs in south africa’, phytophylactica 23, 247–253. van rensburg, b.j., mcgeoch, m.a., chown, s.l. & van jaarsveld, a.s., 1999, ‘conservation of heterogeneity among dung beetles in the maputaland centre of endemism, south africa’, biological conservation 88, 145–153. https://doi.org/10.1016/s0006-3207(98)00109-8 wessels, k.j., reyers, b., van jaarsveld, a.s. & rutherford, m.c., 2003, ‘identification of potential conflict areas between land transformation and biodiversity conservation in north-eastern south africa’, agriculture, ecosystems and environment 95, 157–178. https://doi.org/10.1016/s0167-8809(02)00102-0 appendix 1 table 1-a1: ground-dwelling spiders collected by pitfall trapping from eight habitats in the amanzi private game reserve. table 1-a1 (continues...): ground-dwelling spiders collected by pitfall trapping from eight habitats in the amanzi private game reserve. table 1-a1 (continues...): ground-dwelling spiders collected by pitfall trapping from eight habitats in the amanzi private game reserve. abstract introduction materials and methods results discussion conclusion recommendations acknowledgements references about the author(s) thobile b. dlamini department of nature conservation, tshwane university of technology, pretoria, south africa south african environmental observation network (saeon), ndlovu node, phalaborwa, south africa brian k. reilly department of nature conservation, tshwane university of technology, pretoria, south africa dave i. thompson south african environmental observation network (saeon), ndlovu node, phalaborwa, south africa school of geography, archaeology and environmental studies, university of the witwatersrand, johannesburg, south africa deron e. burkepile department of ecology, evolution, and marine biology, university of california, santa barbara, united states judith m. botha scientific services, south african national parks, skukuza, south africa anthony g. rebelo south african national biodiversity institute, brummeria, pretoria, south africa citation dlamini, t.b., reilly, b.k, thompson, d.i, burkepile, d.e., botha, j.m. & rebelo, a.g., 2020, ‘differential herbivore occupancy of fire-manipulated savannas in the satara region of the kruger national park, south africa’, koedoe 62(1), a1603. https://doi.org/10.4102/koedoe.v62i1.1603 original research differential herbivore occupancy of fire-manipulated savannas in the satara region of the kruger national park, south africa thobile b. dlamini, brian k. reilly, dave i. thompson, deron e. burkepile, judith m. botha, anthony g. rebelo received: 14 oct. 2019; accepted: 20 aug. 2020; published: 26 oct. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the kruger national park’s (knp) long-running experimental burn plots (ebps) have a history of research projects, which improve the understanding of fire in savanna ecosystems. using data from knp’s aerial censuses (2005–2016) and in situ dung count data (2008–2017), this study assessed (1) herbivore densities on the satara, n’wanetsi and marheya ebps, on annual, triennial and no-burn treatments and across pre-, during and post-drought climate conditions; (2) herbivore densities of these ebps relative to their non-manipulated surroundings and (3) the extent to which distance to water and rainfall influence ungulate densities. the results revealed that herbivore mean density differed significantly between the three ebps of satara and across their fire treatments. n’wanetsi showed the highest density (0.30 animals/ha), whilst the lowest was found at marheya (0.12 animals/ha). overall, pre-drought density was higher on the annual plots (0.56 animals/ha), whilst higher post-drought density was evidenced on the triennial plots (0.80 animals/ha). on average, there were significantly higher herbivore densities on the ebps (2.54 animals/ha) compared to the surrounding matrix at the larger scales of the satara management section (0.15 animals/ha) and the central knp (0.18 animals/ha). a positive correlation between herbivore mean density estimate and distance to water was shown. however, grazer mean density across fire treatments was strongly correlated to rainfall. conservation implications: given the variation in fire regimes and their application, and the non-uniform and elevated herbivore densities of the ebps, inferences from the ebps cannot be made to the larger knp. the trials should rather be viewed as an isolated, fire herbivory experiment. it is also recommended to align the experiment with south african national parks’ mandate by including biodiversity parameters like small mammals and insects in the monitoring of the plots. keywords: conservation; protected areas; ecosystem management; ecology; fire; herbivore densities; ebps. introduction fire has long been considered an essential determinant of vegetation structure in savanna ecosystems (sankaran & hanan 2008:17) and is frequently utilised as a management tool by conservationists. changes in fire management in southern africa over the past century reflect an increasing level of ecological consideration (van wilgen 2009:105), partially through the insight of manipulations such as the experimental burn plots (ebps) in the kruger national park (knp), south africa. this fire experiment began in 1954 (biggs et al. 2003:46) and has facilitated research projects on fire ecology in savanna ecosystems (govender, trollope & van wilgen 2006:43). fire extent, frequency and burn shape influence wildlife distributions across landscapes (cromsigt, archibald & owen-smith 2017:47; roberts 2008:1). moreover, spatially patchy fire drives the spatial and temporal distribution of grazers, in particular, to congregate on immediate post-fire patches (mcgranahan & kirkman 2013:176). burning affects forage quality and quantity as well as animal behaviour and diet (roberts 2008:1). fire frequency further influences ungulate distribution through altered predation risk (burkepile et al. 2013:139). at a landscape scale, fire reduces plant biomass (hanan et al. 2008:171), resulting in temporary displacement of herbivores. fire affects nutrient cycling (van der vijver, proot & prins 1999:173) and soil organic nitrogen and carbon (holdo et al. 2009:115), which influence ecosystem productivity. it also modifies plant species composition (anderson et al. 2007:343) and diversity (belsky 1992:187). on the other hand, herbivores also drive changes in vegetation cover (holdo et al. 2009:7). thus, understanding fire’s influence for the management of grazers is essential (eby et al. 2014:1196). post-fire stimulation of plant nutrients is thought to benefit grazing mammals (eby et al. 2014:1198) as plant re-growth is higher in ca, p, mg and k and protein (van der vijver et al. 1999:173). eby et al. (2014:1196) argued that herbivores are attracted to burned patches due to forage quality, rather than a decrease in predation risk. woolfolk et al. (1975:190) and anderson et al. (2007:343) reported a greater body mass gain in herbivores on post-burn areas. fire may affect ungulate density and, together with annual rainfall and distance to water drives herbivore densities and distribution in savannas. artificial surface water provision is widely practised in south african conservation. access to water constrains herbivore movement and alters their distribution, particularly during drier periods (bleich, marshal & andrew 2010:638; deleeuw et al. 2001:297; smith et al. 2007:85). reduced rainfall in drier habitats has an impact on the life history characteristics, physiology and composition of plants and animals (chesson et al. 2004:236). rainfall not only provides water for herbivores but also drives forage quality and quantity (marshal, krausman & bleich 2005:360). this study aimed to (1) quantify herbivore (buffalo, impala, kudu, giraffe, wildebeest and zebra) densities on the knp satara ebps relative to surrounding control areas (central knp and satara management section); (2) relate fire frequencies to herbivore densities and (3) track species density changes across the ebps over time and across rainfall (pre-, during and post-drought) to relate herbivore densities to mean distance to water and rainfall (mm). materials and methods study area the knp (1 948 528 ha) is located in the low-lying savannas of the eastern limpopo and mpumalanga provinces of south africa. mozambique lies to the east and zimbabwe to the north (van wilgen et al. 2000:167). the study was conducted on the controlled ebps of the satara region in the knp. the vegetation community of the area is described as a sclerocarya birrea–vachellia (previously acacia) nigrescens savanna (gertenbach 1983:11). common woody plants include v. nigrescens, s. birrea and pterocarpus rotundifolius (shackleton & scholes 2000:76). the shrub layer is dominated by dichrostachys cinerea, flueggea virosa and grewia sp. the area is underlain by basalts with eutrophic fertile clay soils (gertenbach 1983:11). mean annual rainfall is 548 mm (shackleton & scholes 2000:76), with most of the precipitation falling between november and february (strydom et al. 2013:19). the region experiences variable annual rainfall, with a recent drought < 30% – 40% of long-term average knp rainfall experienced between 2015 and 2016 (smith & bond 2020:120). methodology the study site comprised three burn strings, namely, satara, marheya and n’wanetsi, each comprising 14 plots, each with different fire treatments (biggs et al. 2003:46). the treatments consist of five biennially burned plots, five triennially burned plots and single annual, quadrennial, sexennial and no-burn plots. each string included one additional fire exclusion plot. within the five biennial and triennial treatments, the season of burn varied, and a plot was burnt in one of each of february, april, august, october or december fire seasons. annual plots were burnt in august each year, and the quadrennial and sexennial burned plots were burnt in october. this study focussed on herbivore densities at the august annual and triennial burn treatments in each string, as well as the control plots. treatment plots were 6 ha – 12 ha in size and arranged in strips parallel to the long axis of each string (shackleton & scholes 2000:76). within each string, treatments were randomly assigned to plots within strips (called burn blocks) (shackleton & scholes 2000:76). the control plots were considerably larger and varied between 500 000 ha and 100 000 ha. the density of buffalo, giraffe, impala, kudu, zebra and wildebeest in the control plots, for the period 2005–2016, was determined using the knp’s aerial census data. these data were collected using the protocol detailed in kruger et al. (2008:371). the census data were clipped, using qgis 2.2 into different strata of decreasing spatial scale, that is, central knp (marula north region: ±494 594 ha) and the satara management section (±91 404 ha) prior to density estimation. species density was estimated using the programme distance 6 per determined scale. relative herbivore use was derived from data generated during the 2008–2013 and 2016–2017 dung surveys on the different fire treatments, as detailed by burkepile et al. (2013:136). in brief, data were collected from a series of 50 m × 4 m transects established parallel to the short axis of each burn plot, with seven (2008–2013) and five (2016–2017) transects per plot. herbivore dung piles encountered on transects in january, march and may of each survey year from 2008 to 2013 and monthly in 2016–2017 were identified to the species level. once counted, dung was removed from each transect to avoid recounting during subsequent surveys. the mean number of dung piles per species was calculated for each transect in each burn plot and averaged across surveyed periods. dung density estimates per species were converted to species density estimates following the method put forth by bowland and perrin (1994:505) and smith (2012): where, for each species, d = density (number/ha), n = dung piles (number), t = interval between transect clearance and assessment (days), dr = defecation rate (number of dung piles deposited per day) and a constant 0.02. dung piles deposited per day were determined as the number of dung piles per species divided by the total number of transects multiplied by time. dung decay rate was not included in species density estimates as the time interval between visits (< 3 month) was too short for dung to have decayed in semi-arid savanna (plumptre 2000:356). for the determination of mean distance to water sources, google earth pro was used to measure the distance from the furthest corner of each plot on individual burn strings to active and accessible water sources, both perennial and non-perennial (in 2016–2017), including rivers and artificial water points. mean distance was calculated by summing the distances to water per burn string, divided by the total number of active water sources. the maximum and minimum distances were also indicated. annual rainfall (2005–2017) data for the satara section were used, as there were no data for individual burn strings. these data were provided by the knp. statistical analyses data were transformed (fowler, cohen & javis 2009:179) using square root transformation as the variance of the data was approximately equal to the mean, indicating a poisson distribution. each observation (x) was replaced by and zero counts were replaced by + 0.05. all analyses for dung counts were performed using programme r 3.2.2. distance 6 was used to derive density estimates from aerial counts. a single-factor analysis of variance (anova) was utilised to determine significant differences between estimates. the tukey honest significant differences (tukey hsd) post hoc test was applied for multiple comparisons of means to highlight differences in density estimates across fire treatments and burn strings. pearson’s correlation coefficient was used to quantify the relationship amongst density estimates, annual rainfall and mean distance to water. ethical consideration the authors declare that ethical clearance was not required for this study. results fire treatment mean herbivore density estimates for all years and strings combined were highest on the triennial burn plots (0.24 animals/ha) and lowest on the no-burn plots (0.12 animals/ha) (figure 1a). analysis of variance and a post hoc hsd test across different rainfall conditions (pre-, during and post-drought) revealed a significant difference (p = 0.002) across fire treatments before the drought (2008–2013). pre-drought herbivore mean density on the annual burn plots (0.56 animals/ha) was significantly higher than the triennial (0.33 animals/ha) (p = 0.006) and no-burn (0.32 animals/ha) (p = 0.007) plots (figure 1b). during the drought (2016), herbivore density estimates on the triennial burn plots were significantly (p = 0.01) higher (0.25 animals/ha) than the annual burn plot’s estimates (0.06 animals/ha) (figure 1b). post-drought (2017), the density estimates from the triennial burn plots were significantly (p = 0.003) higher (0.80 animals/ha) than the no-burn plots (0.18 animals/ha) (figure 1b). however, the annual (0.41 animals/ha) and no-burn plots were not significantly different (p = 0.09) in density estimates after the drought. figure 1: pooled herbivore density estimates (a) for three fire treatments (annual burn, triennial burn and no-burn plots) on the satara experimental burn plots. (b) herbivores mean density estimate shown across rainfall conditions (before, during and after drought) and fire treatments (95% confidence interval shown). burn strings an anova and a post hoc (tukey hsd) test revealed significant differences (p = 0.001) in herbivore mean density estimates between individual burn strings. the n’wanetsi burn string (figure 2a) had significantly (p = 0.001) higher pooled (all years combined) density estimates (0.30 animals/ha), whilst the marheya string showed the lowest density at 0.12 animals/ha (p = 0.03). within burn string, a significant (p = 0.01) difference was shown for n’wanetsi between fire treatments (figure 2b). the triennial burn plots had a significantly higher (0.19 animals/ha) estimate, with the lowest shown for the no-burn (0.04 animals/ha) (p = 0.02) treatment plot (figure 2b). similarly, the satara burn string revealed a significant difference (p = 0.02) in herbivore density estimates across fire treatments (figure 2b). herbivore mean density estimates were significantly higher (p = 0.01) on the annual burn plots (0.11 animals/ha) and lowest on the triennial burn (0.06 animals/ha) plots. the no-burn plot revealed a density estimate of 0.10 animals/ha, similar to that of the annual burn plot. the marheya string also revealed significant differences (p = 0.0003) across fire treatments (figure 2b), with density estimates being significantly higher (p = 0.001) on the annual burn plot (0.06 animals/ha) and lowest on the triennially burned plot (0.01 animals/ha). figure 2: (a) overall herbivore density estimates, on the experimental burn strings (n’wanetsi, satara and marheya) of satara, kruger national park, and (b) herbivore density estimates across fire treatments per burn string (95% confidence interval shown). experimental burn plots versus controls herbivore mean density between fire treatments and the adjacent non-manipulated control areas suggests that, overall, the burn strings are preferred by herbivores compared to the control areas. the mean density estimate was seven times higher (2.54 animals/ha) on the burn strings than the non-manipulated control areas (0.33 animals/ha; p = 001) (figure 3a). furthermore, the mean density estimate from the burn strings was 14 times higher than central knp (0.18 animals/ha) and 16 times higher than the satara management section (0.15 animals/ha; p = 001) (figure 3b). figure 3: overall herbivore density estimates compared (a) between experimental burn plot strings and non-manipulated control areas, and (b) amongst burn strings, central kruger national park and the satara management section (95% confidence interval shown). distance to water and rainfall on average, the burn strings were at a mean distance of 3.47 km from active water sources in 2016–2017. pearson’s correlation coefficient revealed a weak correlation (r = 0.57) between the mean distance to water and water abundance (r = 0.50) with the estimated herbivore density across the burn strings. marheya was shown with the lowest herbivore density estimate despite having the most abundant (8), and second closest active water source. a minimum of 0.30 km and a maximum of 6.78 km to water were indicated, with a mean distance of 2.60 km. the n’wanetsi burn string had the highest herbivore density estimate, but the fewest (2) and the farthest active water sources (range, 5.00 km–6.42 km; mean, 5.71 km). the satara burn string had an intermediate herbivore density estimate and an intermediate number of active water sources (5), but the shortest mean distance to water (1.99 km; range, 0.60 km–4.96 km). therefore, the abundance of water sources and their distance to the burn strings showed a petite effect on herbivore density. rainfall showed a significant impact on herbivore densities across fire treatment plots and per feeding guild (figure 4). a strong correlation (r = 0.96) (figure 4a) between herbivore densities on the no-burn and annual burn (r = 0.71) plots and annual rainfall was evident. disaggregation by feeding guild across fire treatments revealed rainfall and the density of grazers to be strongly correlated (r = 0.91) (figure 4b) on annual burn plots. similarly, on the no-burn plots, a strong correlation (r = 0.85) (figure 4d) was shown between the density of grazers and rainfall. a proportional strong correlation (r = 0.84) (figure 4c) was shown between rainfall and the density of grazers and browsers on the triennial burn plots. browser estimates also revealed a positive, but weak, correlation on the annual burn plots (r = 53). conversely, mixed feeders showed a correlation (r = 65) with rainfall on the triennial burn plots and on the no-burn plots (r = 55). this illustrates the significance of annual rainfall as a driver of herbivore densities in fire-prone savannas. figure 4: pearson’s correlation showing the relationship between rainfall (r) and overall herbivore density estimate (mf, mixed feeders; br, browsers and gr, grazers), across all fire treatments (a), and per individual fire treatment, (b) annual (an), (c) triennial (te) and (d) no burn (nb), plots of satara knp. discussion fire showed a vast influence on herbivore densities across the ebps. the higher collective density of herbivores on the ebps compared to the non-fire manipulated areas of the larger central knp and satara section is because of variances in fire regimes. a decrease in abundance of herbs and small shrubs was found by morrison et al. (1995:240) on infrequently burned landscapes such as the non-fire manipulated parts of knp. fire regimes influence changes in habitat structure, plant species composition and palatability (lyon et al. 2000:23). a study by morrison et al. (1995:144) suggests that animals are attracted to an area because of forage quality rather than the fear of predation. therefore, the ebps may be dominated by palatable plant species to attract higher herbivore densities compared to their non-fire manipulated surroundings. the significant differences on herbivore densities across individual burn strings (n’wanetsi, satara and marheya) and their fire treatment plots suggest differences in the temporal nature of each. selective herbivore use of the ebps may be because of the differences in vegetation quality and quantity caused by fire frequency and intensity and rainfall (shackleton & scholes 2000:81). therefore, vegetation palatability across these burn strings may differ, causing herbivores to congregate and forage selectively. selective foraging of herbivores influences the distribution and biomass of plants, in turn affecting fire regime attributes (smith et al. 2013:72). the annual burn plots of the satara ebps have been documented with low grass species richness than the triennial burn plots because of fire frequency and intensity (shackleton & scholes 2000:79). therefore, the continued burning of the annual plots, whilst exposed to higher densities, may have decreased grass species richness even further in the past decade. this causes a shift from annual to triennial plots on overall herbivore densities to occur. an island study conducted by feeley and terborgh (2005:116) suggests that higher herbivore densities in annually burned areas cause a shift in the plant communities towards unpalatable plant species over time. additionally, annually burned plots exposed to grazing because of high herbivore densities reduce the abundance of palatable plant species relative to unpalatable ones (jourbert et al. 2014:126). therefore, plots become dominated by fire-tolerant plant species with reduced composition (boakye et al. 2014:603), in turn affecting herbivore densities. however, burning at a 3-year interval supports both fire-tolerant and fire-intolerant species with high species composition (boakye et al. 2014:603), further facilitating higher herbivore densities and composition. research also outlines that fire frequency and grazing intensity have a detrimental effect on plant species diversity, composition and their nutritional levels (jourbert et al. 2014:126). palatable grass species like panicum maximum were reported to increase in infrequently burned areas like triennial burn plots (smith et al. 2013:72), whereas urochloa mosambicensis was reported to increase in abundance in intermediate and greater fire intensity (smith et al. 2013:80). therefore, higher herbivore densities shifting to triennially burned plots suggest that herbivores follow quality forage and diverse plant species that the annual plot may have lacked. moreover, a 3-year interval burning results in high-vegetation biomass, improving the proportion of palatable species (jourbert et al. 2014:128) compared to the annual burn frequency. however, an increase in the relative abundance of unpalatable species because of annual burning results in a decline in quality of leaf litter, causing a decline in nutrient revenues and primary production rates (pastor & naiman 1992:690). this motivates selective foraging of herbivores, whilst directly affecting herbivore populations across treatment plots. although the loss of grass biomass with frequent burning is attributed to reduced plant diversity (auld & denham 2006:15; snyman 1998:645), herbivore densities are also affected. water abundance and mean distance to water showed a slight effect on the herbivore densities on the ebps. however, this does not exclude abundance of water as the cause of high herbivore congregations on the ebps but may indicate that it is not the only factor. factors like rainfall over the medium term drive forage quality and quantity in african savanna ecosystems (peel & bothma 1995:41; venter, scholes & eckhardt 2003:5). herbivores respond indirectly to this driver, with 87% of the variance in large ungulate biomass being explained by rainfall and fire (fritz & duncan 1994:77). the strong correlation between rainfall and herbivore densities on the ebps coincides with fritz and duncan (1994:77) that rainfall is significantly important to herbivore densities in fire-prone semi-arid savanna ecosystems. rainfall forms the archetype that determines the moisture and nutrients available to plants, therefore, influencing the way habitats are selectively utilised by herbivore populations. hence, impacts on the ebp trial cannot be attributed to fire alone but to a combination of fire, past herbivory and rainfall. in events like ‘drought’, where rainfall is scarce, changes in different functional plant responses are experienced (silver et al. 2013:17), indirectly causing changes in herbivore population densities and distributions. ecosystems like semi-arid savannas with intermittent low annual rainfall seasonally modify co2 and n cycles, resulting in poor water and mineral uptake, hindering growth and causing reduced litter decomposition, alternately altering above ground net primary production (silver et al. 2013:17), in turn influencing changes in herbivore densities before and after a drought period. conclusion the ebps are preferred grazing patches for buffalo, giraffe, impala, kudu, zebra and wildebeest, compared to the non-fire manipulated surrounding areas. our findings suggest that the differences in the burn strings of satara may not only be because of fire but also because of higher herbivore densities. higher herbivore densities impose increased grazing pressure and this questions the level of impact caused by the fire and grazing on the ebps or the impact of both events in succession (shackleton & scholes 2000:80). moreover, the vast differences between the ebps, satara section and the central knp in herbivore densities validate that they are different in the nature of fire regimes and applications, and thus cannot be used to make informed decisions on the broader central knp or satara management section. this motivates management to reconsider the ebps as a fire–herbivory experiment, especially when the density estimate is more than seven times higher in places. although fire proved to have an influence on the herbivore densities on the ebps, factors like rainfall may have contributed to some extent, as the correlation between herbivore densities and rainfall was stronger. this further accentuates that fire and rainfall are critical drivers of herbivore densities on the ebps and fire-prone semi-arid savanna ecosystems. nonetheless, results from this study may be influenced by the 2015–2016 drought period, as this event can affect an ecosystem to some degree. recommendations careful interpretation of results is needed to unbundle the effects of fire, herbivory and rainfall, all of which may have a greater impact on the ebps. the annual treatments have not burned effectively since 1996 compared to the triennials because of inadequate fuel load to sustain a fire. in some cases, the herb layer is too wet and green to facilitate and enable burning. this disrupts the burning schedule for the fire treatment. in addition, burning the annuals in august no longer seems effective, suggesting a revised burning season or a renaming of the treatment as near-annual burns rather than annual burns. the ebps being focal points for research in knp, long term rainfall data for individual burn strings should be available to prevent researchers from relying on regional or section data. putting up rain gauges on the ebps for long-term data collection is advisable, as regional rainfall has proven to have a greater effect on herbivore densities within the ebps. acknowledgements competing interests the authors have declared that no competing interests exist. authors’ contributions all authors contributed equally to this work. funding information this study was funded by the south african national biodiversity institute (sanbi) and the department of nature conservation at tshwane university of technology (tut). the financial assistance of the national research foundation (nrf) and the south african environmental observation network (saeon) towards this research is also acknowledged. data availability statement data of the study belongs to the south african environmental observation network and therefore access can only be permitted by this organisation. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references anderson, t.m., ritchie, m.e., mayemba, e., eby, s., grace, j b. & mcnaughton, s.j., 2007, ‘forage nutritive quality in the serengeti ecosystem: the roles of fire and herbivory’, the american naturalist 170(3), 343–357. https://doi.org/10.1086/520120 auld, t.d. & denham, a.j., 2006, ‘how much seed remain in the soil after a fire?’, plant ecology 187(1), 15–24. https://doi.org/10.1007/s11258-006-9129-0 belsky, j.a., 1992, ‘effects of grazing, competition, disturbance and fire on species composition and diversity in grassland communities’, vegetation science 3(2), 187–200. https://doi.org/10.2307/3235679 biggs, h.c. & rogers, k.r., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59–80, island press, washington, dc. bleich, v.c., marshal, j.p. & andrew, n.g., 2010, ‘habitat use by a desert ungulate: predicting effects of water availability on mountain sheep’, journal of arid environment 74(6), 638–645. https://doi.org/10.1016/j.jaridenv.2009.10.019 boakye, m.k., little, i.t., panagos, m.d. & jansen, r., 2014, ‘effects of burning and grazing on plant species percentage cover and habitat condition in the highland grassland of mpumalanga province, south africa’, journal of animal and plant science 23(2), 603–610. bowland, a.e. & perrin, m.r., 1994, ‘density estimate methods for blue duikers (philantomba monticola) and red duiker (cephalophus natalensis) in natal, south africa’, journal of african zoology 108, 505–519. burkepile, d.e., burns, c.j., tambling, e.a., buis, g.m., govender, n., nelson, v. et al., 2013, ‘habitat selection by large herbivores in a southern african savanna: the relative roles of bottom-up and top-down forces’, ecosphere 4(11), 139. https://doi.org/10.1890/es13-00078.1 chesson, p., gebauer, r.l., schwinning, s., huntly, n., wiegand, k., ernest, m.s. & weltzin, j.f., 2004, ‘resource pulses, species interactions, and diversity maintenance in arid and semi-arid environments’, oecologia 141(2), 236–253. https://doi.org/10.1007/s00442-004-1551-1 cromsigt, j.p.g.m., archibald, s. & owen-smith, r.n., 2017, conserving africa’s mega-diversity in the anthropocene: the hluhluwe imfolozi park story, cambridge university press. london. deleeuw, j., maweru, m.n., okello, o.o., maloba, m., nguru, p. & said, m.y., 2001, ‘distribution and diversity of wildlife in northern kenya in relation to livestock and permanent water points’, bioogical conservation 100(3), 297–306. https://doi.org/10.1016/s0006-3207(01)00034-9 eby, s.l., anderson, t.m., mayembe, e.p. & ritchie, m.e., 2014, ‘the effect of fire on habitat selection of mammalian herbivores: the role of body size and vegetation characteristics’, journal of animal ecology 83(5), 1196–1205. https://doi.org/10.1111/1365-2656.12221 feeley, k. & terborgh, j., 2005, ‘the effect of herbivore density on soil nutrients and tree growth in tropical forest fragments’, ecology 86(1), 116–124. https://doi.org/10.1890/03-0657 fowler, j., cohen, l. & javis, p., 2009, practical statistics for field biology, 2nd edn., wiley john and sons, west sussex. fritz, h. & duncan, p., 1994, ‘on the carrying capacity for large ungulates of african savanna ecosystems’, proceedings of the royal society of london 256(1345), 77–82. https://doi.org/10.1098/rspb.1994.0052 gertenbach, w.p.d., 1983, ‘landscapes of kruger national park’, koedoe 26, 9–121. https://doi.org/10.4102/koedoe.v26i1.591 govender, n., trollope, w.s. & van wilgen, b.w., 2006, ‘the effect of fire season, fire frequency, rainfall and management on fire intensity in savanna’, journal of applied ecology 43, 748–758. https://doi.org/10.1111/j.1365-2664.2006.01184.x hanan, n.p., sea, w.b., dangelmayr, g. & govender, n., 2008, ‘do fires in savannas consume woody biomass? a comment on approaches to modelling savanna dynamics’, the american naturalist 171(6), 851–856. https://doi.org/10.1086/587527 holdo, r.m., sinclair, a.r.e., dobson, a.p., metzger, k.l., bolker, b.m. & ritchie, m.e., 2009, ‘a disease-mediated trophic cascade in the serengeti and its implications for ecosystem’, c.plos biology 7(9), e1000210. https://doi.org/10.1371/journal.pbio.1000210 jourbert, l., pryke, j.s. & samways, m.j., 2014, ‘annual burning drives plant community in remnant grassland ecological networks in an afforested landscape’, south african journal of botany 92, 126–133. https://doi.org/10.1016/j.sajb.2014.02.009 kruger, j.m., reilly, b.k. & whyte, i.-j., 2008, ‘application of distance sampling to etimate population densities of large herbivores in kruger national park’, wildlife research 35, 371–376. lyon, l.j., huff, m.h., hoper, r.g., telfer, e.s., schreiner, d.s. & smith, j.k., 2000, ‘wildland fire in ecosystems’, wildland fire in ecosystems: effects of fire on fauna. usda forest service research; pap. se -234, pp. 23, fort collins, colorado. marshal, j.p., krausman, p.r. & bleich, v.c., 2005, ‘rainfall, temperature and forage dynamics affect nutritional quality of desrt mule deer forage’, rangeland ecology and management 58(4), 360–365. https://doi.org/10.2111/1551-5028(2005)058[0360:rtafda]2.0.co;2 mcgranahan, d.a. & kirkman, k.p., 2013, ‘multifunctional rangeland in southern africa: managing for production, conservation and resilience with fire and grazing’, land 2(2), 176–193. https://doi.org/10.3390/land2020176 morrison, d.a., cary, g.t., pengelly, s.m., ross, d.g., mullins, b.j., thomas, c.r. et al., 1995, ‘effects of fire frequency on plant species composition of sandstone communities in the sydney region: inter-fire interval and time-since-fire’, australia journal of ecology 20(2), 239–247. https://doi.org/10.1111/j.1442-9993.1995.tb00535.x pastor, j. & naiman, r.j., 1992, ‘selective foraging and ecosystem processes in boreal forests’, the american naturalist 139(4), 690–705. https://doi.org/10.1086/285353 peel, m.j.s. & bothma, j.d.p., 1995, ‘comparison of accuracy of four methods commonly used to count impala’, south african journal of wildlife research 25(2), 41–43. plumptre, a.j., 2000, ‘monitoring mammal populations with line transect technique in african forests’, journal of applied ecology 37(2), 356–368. https://doi.org/10.1046/j.1365-2664.2000.00499.x roberts, l.s., 2008, ‘special issue: fire and wildlife interaction’, fire ecology 4(2), 1–2. https://doi.org/10.4996/fireecology.0402001 sankaran, m.r. & hanan, n.p., 2008, ‘woody cover in african savannas: the role of resources, fire and herbivory’, global ecology & biogeography 17(2), 236–245. https://doi.org/10.1111/j.1466-8238.2007.00360.x shackleton, c.m. & scholes, r.j., 2000, ‘impact of fire frequency on woody community structure and soil nutrients in the kruger national park’, koedoe 43(1), 75–81. https://doi.org/10.4102/koedoe.v43i1.210 silver, e.c., albuquerque, m.b., azevedo neto, a.d. & silver junior, c.d., 2013, ‘drought and its consequences to plants from individual to ecosystem’, in s. akinci (ed.), responses of organisms to water stress, pp. 17–47, in tech, croatia. smith, d., 2012, inventory and monitoring toolbox: animal pests: faecal pellet counts, department of conservation, te papa atawhai. smith, i.p.j. & bond, w.j., 2020, ‘observations on the natural history of a savanna drought’, african journal of range & forage science 37(1), 119–136. https://doi.org/10.2989/10220119.2020.1723695 smith, i.p.j., grant, c.c. & devereux, b.j., 2007, ‘do artificial water holes influence the way herbivores use the landscape? herbivore distributions patterns around rivers and artificial surface water sources in a large african savanna park’, biological conservation 136(1), 85–99. https://doi.org/10.1016/j.biocon.2006.11.009 smith, m.d., van wilgen, b.w., burns, c.e., govender, n., potgieter, a.l.f., andelman, s. et al., 2013, ‘long-term effects of fire frequency and season on herbaceous vegetation in savannas of the kruger national park, south africa’, journal of plant ecology 6(1), 71–83. https://doi.org/10.1093/jpe/rts014 snyman, h.a., 1998, ‘dynamics and sustainable utilization of the rangeland ecosystem in arid and semi-arid climates of southern africa’, journal of arid environment 39(4), 645–666. https://doi.org/10.1006/jare.1998.0387 strydom, t., riddell, e., lorentz, s. & le roux, p., 2013, ‘the effects of longterm fire frequencies on soil hydralic properties in semi-arid savannas in kruger national park’, msc thesis, center for water resource research, school of agriculture, earth and environmental sciences, pietermaritzburg. van der vijver, c.a.d.m., proot, p. & prins, h.h.t., 1999, ‘causes of increased nutrient concetrations in post-fire regrowth in an east afrcan savanna’, plant and soil 214(1), 173–185. https://doi.org/10.1023/a:1004753406424 van wilgen, b.w., 2009, ‘the evlolution of fire management in savanna protected areas in south africa’, south african journal of science 105(9/10). https://doi.org/10.4102/sajs.v105i9/10.107 van wilgen, b.w., biggs, h.c., o’regan, h. & mare, n., 2000, ‘a fire history of the savanna ecosystems in the kruger national park, between 1941 and 1996’, south african journal of science 96(4), 167–178. venter, f.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation pattern’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83–129, island press, washington, dc. woolfolk, j.s., smith, e.f., schalles, r.r., brent, b.e., harbes, l.h. & owensby, c.e., 1975, ‘effects of nitrogen fertilization and late-spring burning on bluestem range on diet and performance of steers’, journal of rangeland management 28(3), 190–193. https://doi.org/10.2307/3897523 http://www.koedoe.co.za open access page 1 of 1 reviewer acknowledgement acknowledgement to reviewers in an effort to facilitate the selection of appropriate peer reviewers for koedoe, we ask that you take a moment to update your electronic portfolio on https://koedoe.co.za for our files, allowing us better access to your areas of interest and expertise, in order to match reviewers with submitted manuscripts. if you would like to become a reviewer, please visit the journal website and register as a user. in order to be considered, please email submissions@koedoe. co.za indicating your intention to register as a reviewer for the journal. to access your details on the website, you will need to follow these steps: 1. log into the online journal at https://koedoe. co.za 2. in your ‘user home’ [https://koedoe.co.za/ index.php/koedoe/user] select ‘edit my profile’ under the heading ‘my account’ and insert all relevant details, bio statement and reviewing interest(s). 3. it is good practice as a reviewer to update your personal details regularly to ensure contact with you throughout your professional term as reviewer to koedoe. please do not hesitate to contact us if you require assistance in performing this task. publisher: publishing@aosis.co.za tel: +27 21 975 2602 the editorial team of koedoe recognises the value and importance of the peer reviewer in the overall publication process – not only in shaping the individual manuscript, but also in shaping the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely, productive peer review process. we would like to take this opportunity to thank and recognise the following reviewers for their precious time and dedication, regardless of whether the papers they reviewed were finally published. we apologise for any names that have been inadvertently left out. these individuals provided their services to the journal as a reviewer from 01 october 2021 to 30 september 2022. ana novoa andrinajoro r. andrinajoro babatope e. akinyemi conrad a. matthee corli wigley-coetsee david h.m. cumming dian spear eddie riddell graham durrheim isaac mapaure jasper knight jessica comley keenan stears krystal tolley liaan minnie marna herbst megan loftie-eaton michael f. bates nick rivers-moore oupa e. malahlela peet van der merwe peter j. apps sandy steenhuisen shané gertse tara massad tatenda dulu tercia strydom theo h. mostert tshepiso mangani http://www.koedoe.co.za� https://koedoe.co.za� mailto:submissions@koedoe.co.za mailto:submissions@koedoe.co.za https://koedoe.co.za� https://koedoe.co.za� https://koedoe.co.za/index.php/koedoe/user https://koedoe.co.za/index.php/koedoe/user mailto:publishing@aosis.co.za edge.qxd some ecological factors influencing the breeding success of the brenton blue butterfly, orachrysops niobe (trimen) (lepidoptera: lycaenidae) d.a. edge edge, d.a. 2002. some ecological factors influencing the breeding success of the brenton blue butterfly, orachrysops niobe (trimen) (lepidoptera: lycaenidae). koedoe 45(2): 19–34. pretoria. issn 0075-6458. the brenton blue butterfly, orachrysops niobe (trimen, 1862) (lepidoptera: lycaenidae), is endemic to the southern cape and is currently listed as endangered. this study looks at some of the key ecological factors influencing the breeding success of the species—host plant abundance and condition, nectar sources, climate/ microclimate, and vegetation management techniques. the adult butterfly population was monitored over an entire breeding season; host plants were identified and individually monitored; and egg counts were done. this enabled the effects of a number of different management techniques to be evaluated (burning, cutting, physical removal of invasive elements, and combinations thereof). a fivefold increase in the population of o. niobe was observed over the breeding season. this increase was positively correlated to a similar increase in host plant abundance in the areas where cutting and physical removal of invasive elements was practiced. burning, by contrast, appeared to have a negative impact on host plant and butterfly abundance over the same period. impacts of other factors such as climate, nectar sources and the natural strength of the second brood are discussed. a hypothesis, of megaherbivore activity as the principal historical disturbance mechanism promoting locally favourable conditions for o. niobe to establish and maintain colonies, is proposed. recommendations for reserve management and future research are made. key words: butterfly ecology, endangered species, nectar, population dynamics, population counts, reserve management. d.a. edge, school of environmental sciences and development, potchefstroom university, p o box 2586, knysna, 6570, western cape province, south africa. issn 0075-6458 19 koedoe 45/2 (2002) introduction the brenton blue butterfly orachrysops niobe (trimen, 1862) (lepidoptera: lycaenidae) is one of south africa’s most threatened butterfly species, being endemic to the southern cape and currently red-listed as endangered (henning & henning 1995). it is now only known to occur at one locality —at brenton-on-sea near knysna. it previously occurred in nature’s valley 50 km to the east but has become extinct there as a result of anthropogenic activities (property development, habitat fragmentation, prevention of fires, and exclusion of larger herbivores). the colony at brenton-on-sea has been subject to similar pressures but has been given a reprieve from a similar fate following a protracted and highly publicised campaign to save the species (steenkamp & stein 1999). this has resulted in the procurement of the land on which the colony occurs by central government and the impending proclamation of a reserve which covers 1.47 ha. research has been conducted on the life history (edge & pringle 1996; williams 1996); butterfly population size and distribution (britton & silberbauer 1997); the ant populations found at the site (robertson 1997, 1998, 2000); and on the floristics and plant communities (lubke et al. 1996). case histories from other endangered butterfly species (mostly from the northern hemiedge.qxd 2005/12/09 10:02 page 19 sphere) have shown that it is essential to understand the ecology of each endangered butterfly species in detail if conservation efforts are to succeed (new 1997). the aim of the current research is to determine some of the critical ecological factors influencing the breeding success of o. niobe. since edge & pringle (1996) demonstrated by rearing adult o. niobe from ova that the larvae are essentially phytophagous the abundance and condition of the sole larval host plant indigofera erecta thunberg is one of the factors examined in detail. several vegetation management techniques to promote proliferation of the host plant have been proposed by other workers (lubke et al. 1996; schutte-vlok 2001), and their efficacy needs to be tested. other factors dealt with in the present study are nectar sources and preferences; the influence of climate; and the population dynamics of o. niobe. study site the study site is the brenton blue butterfly reserve, situated in brenton-on-sea near knysna (34º04'20''s; 23º02'00''e) in the western cape province of south africa (fig. 1). this figure is to scale and depicts areas that were treated as well as the network of paths that were cut. the reserve has a total area of 14 673 m². koedoe 45/2 (2002) 20 issn 0075-6458 fig. 1. brenton blue butterfly reserve, near knysna. edge.qxd 2005/12/09 10:02 page 20 the reserve is bounded to the north by a tarred road, w.k. grobler drive. to the north of this road is a public open space, uitzicht 216 portion 81, which is owned by the district municipality. to the south, the reserve boundary is fynbosoord, a tarred access road to the residential properties which lie south of fynbosoord and at the south-eastern corner of the reserve. the reserve is bounded on the west by residential properties and stands, and mountain rose drive. the eastern boundary abuts partly on residential property and partly onto public open space that does not presently form part of the reserve. the study site is on a well-drained south (seaward) facing slope of average inclination of 1 in 4. it is at an altitude of 95 m to 120 m and is 450 m from the sea. the geology, climate and floristics of the site were described by lubke et al. (1996). the vegetation can be characterised as asteraceous coastal fynbos with patches of candlewood-dominated thickets [pterocelastrus tricuspidatus (lamont) sonder]. materials and methods habitat modification habitat modifications (treatments) were carried out in the areas indicated on the site plan (fig. 1) and summarised in table 1. the procedure for the burn on 20 september 2000 was described by hiseman (2000). the “skoffeling” treatment applied to area b involved cutting down of the bracken fern (pteridium aquilinium (linnaeus) kuhn), and turning over the surface layer down to a depth of 0.2 m so that the roots of the bracken were pulled out. the intention was to simulate the activities of bush pigs, which it had been hypothesised by schutte-vlok (2001) played an important role in the past in controlling the bracken fern. the network of access paths (area d) were cut during the period from 4–25 july 2001, using shears at a level of approximately 10 cm above the ground, to a width of 1–1.5 m. the paths wind to avoid larger bushes and trees and any patches of the host plant i. erecta, and have been subsequently kept open by pulling out any new bracken fern plants. the blocks contained within the paths (area e) have not been disturbed, but difficulty of access has precluded any systematic host plant observations in this study. area f consists of a steep embankment approximately 1.5 m high running east to west which resulted from excavations for a sewerage line in the early 1990s. indigofera erecta monitoring immediately after the fire careful watch was kept for any i. erecta seedlings. as soon as they appeared they were marked and identified with 1-m sticks and numbered tags. thereafter, during host plant surveys, any new plant which had sprouted was similarly identified. these plants were given a letter following the number to show that they were not amongst the original cohort. when the paths were cut all existing i. erecta plants exposed were also marked, as were the new ones that germinated and/or sprouted later on. no distinction was made as to whether the new plant had originated from seed or had resprouted from an old root stock. issn 0075-6458 21 koedoe 45/2 (2002) table 1 summary of treatment areas treatment area (m²) description of treatment a 1400 burnt on 20 september 2000. natural succession thereafter. b 1000 burnt on 20 september 2000. “skoffeled” prior to fire, natural succession thereafter. c 300 burnt on 20 september 2000. bracken fern controlled by manually pulling new plants out for 6 months thereafter. d 1500 network of access paths cut through the unburnt portion of the reserve, and kept open thereafter. e 10450 undisturbed, unburnt blocks inside paths f 120 steep 1.5 m high embankment created by excavation for sewerage line in early 1990s edge.qxd 2005/12/09 10:02 page 21 surveys of all the identified i. erecta plants were conducted at regular intervals (19 march, 9 may, 18 july, 25 july, 10 october, 10–25 november 2001 and 28 january–1 march 2002). during the march and may surveys the health of each plant was recorded. this was a subjective measure made by eye on a scale from 1 to 10, and took into account the overall size of the plant and its stolons, withering of leaves and stems, and whether the plant appeared to be spreading or shrinking. from july the size of each plant was recorded by counting the number of stolons and the number of inflorescences. a stolon, for the purpose of this study, is defined as an aerial shoot of the plant (horizontal or vertical) which originates near the root stock. stolons vary in length from 150 mm to 500 mm. the inflorescences of i. erecta are vertical racemes that carry the flowers on sturdy peduncles. figure 2 is a diagrammatic sketch of a typical plant (the detailed morphology of the root system is not yet known). adult butterfly population counts in 2000 and 2001 experiments by the author with mark, release, recapture (mrr) techniques had shown that the trauma to the butterflies caused by this method posed an unacceptable risk to the small and fragile population. consequently a transect method was devised, using the network of paths, to yield a comparative count of butterflies which could be used to assess population trends. this method required similar weather conditions (sunny, warm, little wind) and was carried out at the same time each day (11:30–12:30). the route through the network was the same for each count, at a fairly constant pace of 20 m/min., and took about one hour, allowing some time for stops to record significant observations such as nectar plant visitation, oviposition or mating. during the population counts all adults seen were recorded with the place, gender and activity. multiple sightings of a particular insect inevitably took place but each sighting was recorded. the observations for each day were recorded on a data sheet that has a schematic layout of the paths in the reserve incorporated and shows the traverse route (fig. 3). a second counting method was used which is referred to as the “fixed point” method. the observer was positioned at point a2 (which is on the normal male patrolling route) and recorded all o. niobe that came within 10 metres of this point during a 30 minute period (at the same time each day). the fixed point method was “calibrated” on several separate occasions by capturing all the butterflies present and retaining them in a gauze netting cage. this enabled a relationship to be derived between the “fixed point” count and the total population of butterflies patrolling (referred to as the repetition factor for this method). repetition factor = total count made / actual butterflies present koedoe 45/2 (2002) 22 issn 0075-6458 fig. 2. typical growth form of indigofera erecta (diagrammatic). edge.qxd 2005/12/09 10:02 page 22 the repetition factor for the transect method could only be estimated (as between 1.25 and 2.0). the butterflies on the paths were much less mobile than the patrolling males mostly seen from the fixed point and the observer moved quickly along each path, reducing the chance of repeat observations of the same insect. observations made during the transects that fell within the “fixed point” area (defined as the three paths observable from the fixed point a2) were treated as if they were made during a fixed point count for the purpose of applying a repetition factor and calculating the actual butterflies present. adult butterfly behaviour during the adult butterfly counts and the plant surveys, all noteworthy adult butterfly behaviour was recorded. categories were defined as fluttering (typical slow female flight within 15 cm of the ground or vegetation); patrolling (more vigorous male flight up to a metre from the ground or over vegetation); mate search and acquisition (males only); nectar plant visitation (plant species and butterfly gender recorded); host plant search; oviposition (plant number recorded); copulation (time and location recorded); basking, and resting. issn 0075-6458 23 koedoe 45/2 (2002) early stages (eggs, larvae) all the identified host plants were searched thoroughly during the period from 10–25 november 2001 and again from 28 january–1 march 2002 for eggs and larvae. the search method involved examining both sides of each stolon. in order to standardise the search about one minute was taken to search each stolon. when eggs or larvae were found the host plant number was recorded with the count and any unusual features (e.g., eggs were usually laid on the lower surface of a leaf—so if they were in another position this was recorded). eggs were recorded as unhatched (pale blue, entire) or hatched (white ring of eggshell remnant). climate and microclimate rainfall records from the previous ten years were obtained from a weather station located at 140 watsonia road, brenton-on-sea. this station is situated at 90 m altitude, 400 m from the sea, and 500 m away from point a1 in a westerly direction and is also sited on a south-facing slope. an additional weather station was established at the site (position near a1 as shown on fig. 1) and daily recordings commenced in june 2001. data recorded was mm rainfall using a conical rain gauge and the maximum and minimum temperatures using a thermometer. the only microclimate factor recorded during the present study was a subjective estimate by eye of the fig. 3. data sheet incorporates a schematic layout of the paths in the reserve and shows the traverse route. edge.qxd 2005/12/09 10:02 page 23 koedoe 45/2 (2002) 24 issn 0075-6458 table 2 plants and stolons of indigofera erecta counted in each area between march 2001 and february 2002 treatment area original a plants per 100 m² stolons per 100 m² plants/ oct nov feb oct nov feb 100 m² 2001 2001 2002 2001 2001 2002 fire/ bracken control (c) 16.3 16.7 19.0 14.0 32 82 54 fire/ no control (a) 1.7 1.9 2.0 1.9 10 26 13 embankment (f) 10.0 13.3 15.0 14.2 62 128 92 paths in unburnt area (d) 7.0 8.5 12.7 14.3 24 109 133 totals 5.7 6.6 8.9 9.0 20 72 73 a start date for areas (c), (a) and (f) was 19 march 2001, and for area (d) was 25 july 2001. table 3 shade factors, population and growth records of i. erecta in each treatment area treatment area shade original new deaths percentage increase in % plants plants stolons counted a fire/ bracken control (c) 22.4 49 17 24 68 fire/ no bracken control (a) 47.6 24 5 3 24 embankment (f) 81.2 12 9 4 49 paths in unburnt area (d) 38.3 105 121 12 463 totals/averages 38.6 190 152 43 265 a defined as: (final stolon count initial stolon count) / (initial stolon count) x 100 table 4 orachrysops niobe total adult population counts between october 2001 and march 2002, using two methods and standardised as counts per hour first brood second brood date total count /h date total count/h transects fixed point transects fixed point 29.10.01 3.81 12.0 14.01.02 1.60 08.11.01 4.86 13.3 28.01.02 19.09 42.9 09.11.01 4.55 30.01.02 15.86 63.2 10.11.01 4.07 12.0 04.02.02 18.21 35.0 11.11.01 5.41 17.1 08.02.02 11.79 54.0 12.11.01 4.20 27.0 13.02.02 19.29 87.0 13.11.01 3.85 10.5 17.02.02 13.58 39.0 16.11.01 4.76 11.4 26.02.02 4.14 18.0 18.11.01 3.53 6.0 08.03.02 2.67 averages 4.34 13.7 averages 11.8 42.9 edge.qxd 2005/12/09 10:02 page 24 maximum and minimum temperatures using a thermometer. the only microclimate factor recorded during the present study was a subjective estimate by eye of the extent of shade at each host plant site, ranging from 0 % (no shade for whole day) to 100 % (full shade for whole day). results indigofera erecta monitoring the number of living i. erecta plants located and the number of stolons counted in each treatment area at different dates is given in table 2, and a summary of the i. erecta microclimate and population dynamics data gathered appears in table 3. adult butterfly population counts a summary of the data obtained using the two comparative methods (transect and fixed point) is presented in table 4. each day’s count using the two methods has been standardised to a count per hour so that the figures for each day are directly comparable. no butterflies were observed between 21 november 2001 and 10 january 2002. table 5 presents the total counts from the two methods taken over six days, during the peak emergence of each brood with favourable weather conditions. these results are another assessment of the relative strength of the two broods. table 6 presents the results of calculations made from the observations to determine the sex ratio of the adults (assuming a repetition issn 0075-6458 25 koedoe 45/2 (2002) table 5 orachrysops niobe adult population counts for six days at the peak of the november 2001 and february 2002 broods using two counting methods period of surveys entire reserve transects fixed point a2 (11:30-12:30) (20 minutes per day) males females totals males females totals november 2001 (1st brood) 12 17 29 25 2 27 jan/feb 2002 (2nd brood) 39 54 93 99 14 113 ratios (2nd:1st brood) 3.25 3.18 3.21 3.96 7.0 4.19 table 6 calculation of o. niobe sex ratios during the november 2001 and february 2002 broods (male and female) november 2001 brood february 2002 brood method/area transect fixed total transect fixed total time (mins) 740 276 1016 450 195 645 male count 20.5 44.5 65 22.8 135.2 158 female count 25.6 7.4 33 63.3 16.7 80 repetition/ hr 1.25 6.56 1.25 6.56 actual male/h 1.330 1.475 2.805 2.432 6.341 8.773 actual female/h 1.661 0.245 1.906 6.752 0.783 7.535 male:female ratio 0.80 6.02 1.47 0.36 8.10 1.16 edge.qxd 2005/12/09 10:02 page 25 koedoe 45/2 (2002) 26 issn 0075-6458 fig. 4. o. niobe comparative population counts using the fixed point method. fig. 5. o. niobe comparative population counts using the transect method. table 7 orachrysops niobe egg counts in each treatment area during november 2001 and january/february 2002 (h = hatched eggs and u = unhatched eggs) treatment areas period of surveys increase 10 nov. 2001 25 nov. 2001 28 jan. 2002 27 feb. 2002 factor 1st–2nd h u totals h u totals brood fire/ bracken control (c) 2 0 2 15 1 16 8.00 fire/ no bracken control (a) 2 7 9 10 11 21 2.33 embankment (f) 3 1 4 11 2 13 3.25 paths in unburnt area (d) 106 109 215 440 606 1046 4.87 totals 113 117 230 476 620 1096 4.76 factor of 1.25 for the transect method). the repetition factor for the fixed point method was determined to be 6.6. the overall male to female ratio in the colony falls within the range from 1.2 to 1.5. if a repetition factor for the transect method of 2.0 is used the male to female ratio falls within the range 1.6 to 1.8. figures 4 and 5 present graphically the population fluctuations over a 21 weeks period from 17 october 2001 to 12 march 2002, using the two methods. oviposition the i. erecta plants preferred for oviposition were mostly healthy and vigorous, on well drained sloping ground and in semi shade with bare ground in the immediate vicinity of the plant. the eggs were usually laid on the underside of leaves on a horizontal stolon close to the ground. the eggs were therefore well concealed and in a cool, damp microclimate. only occasionally (3.2 % of observations) were eggs laid on the upper surface of the leaves, on stems (as opposed to leaves) or on vertical stolons (as opposed to horizontal stolons). the egg counts (h = hatched and u = unhatched) for each brood are presented in table 7. the data from tables 7 & 2 can be used to calculate egg densities (per plant and per stolon) for each treatment area as shown in table 8. edge.qxd 2005/12/09 10:02 page 26 issn 0075-6458 27 koedoe 45/2 (2002) table 8 orachrysops niobe average egg densities per plant and per stolon in each treatment area during november 2001 and january/ february 2002 areas period of surveys 10 nov. 2001–25 nov. 2001 28 jan. 2002 –27 feb. 2002 per 100 plants per 100 stolons per 100 plants per 100 stolons fire/ bracken control (c) 0.035 0.008 0.38 0.10 fire/ no bracken control (a) 0.32 0.02 0.81 0.12 embankment (f) 0.22 0.026 0.77 0.12 paths (d) 1.13 0.13 4.89 0.53 totals 0.78 0.096 3.65 0.45 table 9 nectar sources and visitations by o niobe adults plant species family number of months areas flower symmetry b visitations a colour commelina africana l. commelinaceae 2f february d2 yellow z hypoxis villosa l. hypoxidaceae 1f november c1 yellow a felicia echinata (thunb.) asteraceae 1m february c1 lilac & a yellow lobelia tomentosa l. f. campanulaceae 1m;3f january a1;c2 february d1 blue z erica speciosa andrews ericaceae 2f january d2 red a indigofera erecta thunb. fabaceae 1f november d1 pink z indigofera verrucosa fabaceae 2m;3f january eckl.& zeyh. february c5 pink z rhynchosia caribea (jacq.) fabaceae 1m february c1 yellow z rhynchosia chrysoscias fabaceae 6m;6f november c8 benth. ex harv. february d4 yellow z geranium incanum burm. geraniaceae 3m;2f november c4 a1 lilac a sebaea aurea (l.f.) roem. & schult. gentianaceae 1m november a1 yellow a polygala fruticosa p.j.bergius polygalaceae 1f february d1 purple z selago glomerata thunb. selaginaceae 8m;12f january c15 a february 3 d2 pale blue a totals 10 families 23m a6 c37 33f d13 a m = male and f = female b a = actinomorphic and z = zygomorphic nectar plant visitation a summary of the observations made of o. niobe adults visiting nectar plants appears in table 9. thirteen angiosperm species were recorded from 10 different families, with selago glomerata thunberg being the most frequently selected nectar source (35.7 % of observations). actinomorphic (radially symmetrical) flowers were narrowly preferred over zygomorphic (bilaterally symmetrical) flowers (53.6 %) and lilac or pale blue flowedge.qxd 2005/12/09 10:02 page 27 ers were chosen over other colours (57.1 %). the majority of the nectar plant visitation observations were made in the burnt areas (76.8 %), where the preferred nectar plants (which were mostly forbs) were more abundant. climate the weather data in table 10 was obtained from the 140 watsonia road weather station (rainfall); the weather station at the reserve (rainfall and temperatures); and the knysna records of s.a. weather services (saws). the watsonia road and reserve stations agreed for rainfall records over fourteen months to within 1.5 %. there is also good agreement between the watsonia road station and saws over the longer term. overall comparison of the burnt areas versus the paths drawing the data together from the various sources in this study the summary compiled in table 11 was produced. discussion reproduction and growth of indigofera erecta the genus indigofera is large, with 78 species occurring in the cape floristic region (cfr) and 720–730 worldwide (goldblatt & manning 2000). the taxonomy is still being actively researched and goldblatt & manning identify 19 undescribed species in the cfr (24 % of the total). a variety of reproductive strategies are evident in the genus, koedoe 45/2 (2002) 28 issn 0075-6458 table 10 weather records for the knysna area month 140 watsonia road brenton blue knysna s.a. rainfall (mm) a butterfly reserve b weather service c max. min. mean rainfall average average rainfall average average mean max. min mean max. min mm °c ° c mm °c ° c january 82 21 52.4 53 28.6 14.5 56.8 26.2 17.1 february 85 12 47.3 16 28.4 14.9 47.9 27.2 16.7 march 107 6 57.0 14 26.8 15.5 59.7 27.1 17.0 april 122 0 60.8 37 24.8 13.2 65.7 20.1 10.1 may 128 6 51.3 62 20.4 10.7 64.7 21.4 11.4 june 74 8 34.3 44 17.4 8.4 64.0 19.5 8.3 july 117 18 61.0 62 17.5 9.0 60.9 19.0 8.7 august 200 0 85.2 103 20.5 9.9 84.6 20.1 10.0 september 216 35 61.7 54 19.9 10.9 64.7 20.9 11.9 october 192 25 87.4 42 22.8 13.5 74.2 22.9 14.7 november 122 0 87.3 85 24.9 13.8 59.0 23.3 15.9 december 144 12 62.5 53 27.8 15.1 50.8 24.9 17.1 annual 1078 564 756.3 625 753.1 a from january 1992 to august 2002 b from july 2001 to august 2002 c from january 1951 to august 2002 edge.qxd 2005/12/09 10:02 page 28 issn 0075-6458 29 koedoe 45/2 (2002) table 11 summary of data comparing the burnt areas with the paths description of data burnt areas paths a, b &c (area d) total area (m²) 2700 1500 final i. erecta stolons count 341 1988 final i. erecta stolons per 100 m² 13 130 i. erecta stolon increase (%) 41.5 463 o. niobe eggs laid per 100 m² (november) 0.4 14.3 o. niobe eggs laid per 100 m² (february) 1.3 70 o. niobe eggs laid per plant (november) 0.129 1.13 o. niobe eggs laid per plant (february) 0.544 4.89 % of total o. niobe eggs laid (november) 6.5 93.5 % of total o. niobe eggs laid (february) 4.6 95.4 % of male o. niobe sightings (november) 69.0 31.0 % of female o. niobe sightings (november) 22.2 77.8 % of male o. niobe sightings (february) 31.1 68.9 % of female o. niobe sightings (february) 10.6 89.4 adult o. niobe nectar visitations 43 13 average shading factor (%) 43 38 with six species being described as resprouting and six as reseeding shrubs or shrublets. the biology and phenology of i. erecta have not yet been studied as far as can be ascertained. two other indigofera species have been identified at the site (i. verrucosa and i. glaucescens ecklon & zeyher). schutte-vlok (2001) hypothesised that the germination of i. erecta was principally fire induced, and used this as one of the main arguments for management of the reserve by regular controlled burns. jeffrey et al. (1997) found that not all legume species tested germinated in response to dry heat treatment and hypothesised that species found in dune or forest communities, which were less fire prone, required other stimuli to break seed dormancy. cocks & stock (1997), in a study of germination stimulation of 16 fabaceae species, found that response to fire varied and seemed to be correlated with certain seed characteristics. observations and experimentation on the host plant of orachrysops ariadne (butler, 1898) which is indigofera woodii h.bolus var. laxa — indicated that whilst fire and heat seemed to play a role in seed germination, seedlings emerged in unburned areas as well as burnt areas. (lu & samways 2002b). the evidence accumulated in the present study indicates that the germination of i. erecta is stimulated not only by fire as expected (table 3—areas a and c—“new plant” column) but also by surface disturbance and by removal of competing plants to admit more light (access paths—area d). effects similar to this could have been caused naturally in the past by megaherbivores creating paths for access and passage, and by their browsing and grazing on the plants along the paths. elephants may have occasionally visited the area in the past and their partiality to many typical fynbos plants as well as p. tricuspidatus, and the rhizomes of p. aquilinium has been recorded recently in fynbos areas adjacent to the knysna forest (milewski 2002). it is interesting that these two species have a tendency to proliferate at the study site in the absence of any control agent(s). at the study site the predominant growth form of i. erecta is depicted diagrammatically in fig. 2. most of the vegetative mass is contained in the horizontal stolons, which are edge.qxd 2005/12/09 10:02 page 29 koedoe 45/2 (2002) 30 issn 0075-6458 up to 500mm long and spread along the ground in a horizontal or downward direction, depending on the slope of the ground. vertical stolons are much less frequent and appear to be a response to there being greater ground cover by competing plants in the vicinity. since the stems of i. erecta are herbaceous the height of the vertical stolons is limited to 100mm by their mechanical strength unless support is gained from neighbouring plants, in which case a height of a metre or more can be attained. leaves are sparser on these vertical stolons compared to the horizontal stolons, which are supported by the ground and therefore able to bear many more leaves. the growth form described above is markedly different from the host plant of o. ariadne which is an upright plant growing to over a metre and which has woody stems (lu & samways 2001). this type of growth form can compete successfully with taller plants and does not require bare ground to grow vigorously which appears to be the case with i. erecta. the influence of growth form can be seen by comparing the response of i. erecta to fire as opposed to cutting and clearing around the plant. the i. erecta plants that sprouted after the fire did well at first, while the ground was bare and there was not much shade. however they were out-competed later as the ground became covered with grass and sedge species and the taller more vigorous species (p. aquilinium, helichrysum spp, rhyncosia spp, etc) shaded out the smaller i. erecta plants. consequently growth of both horizontal and vertical stolons was inhibited and not only did a high proportion of i. erecta plants die during the observation period (refer to table 3), but also the overall increase in the number of stolons was low. by contrast the i. erecta plants which were present on the paths, together with those that sprouted subsequently, had a much lower mortality and a far greater rate of increase in the number of stolons (refer to tables 2 & 3). this shows clearly that growth of i. erecta was encouraged to a far greater extent by the activity (disturbance) of cutting the access paths rather than by the fire. it is noteworthy that parts of the reserve area were regularly disturbed in the 1980s and early 1990s by the property owner who used to cut down the taller plants every few years to gain access for surveying. escom also used to keep the strip of ground along their power servitude just below the present w. k. grobler drive open for maintenance access. these sort of activities may have been critical sustaining factors for the butterfly at the site over the last two decades. orachrysops niobe adult population fluctuations the author has been keeping records of adult population counts dating back to 1993. in that year, using a method similar to but not as rigorous as in the present study, a maximum count of 10/h was obtained for the november brood using a fixed point observation method. in november 2000 the maximum count was 16/h also using a fixed point method, and in november 2001 the maximum count was 15/h. it can therefore be stated with reasonable confidence that the november population of adult o. niobe has not shown any significant decline over the eight year period, and if anything seems to have increased. however, in previous years, whilst the author has sparse records for december, january and february, a major second brood such as emerged in february 2002 has never been detected, with at most a hand full of individuals being seen. the three to four fold increase in the adult numbers between november 2001and february 2002 (figs. 4 & 5), and the nearly five fold increase in the egg counts between the two broods, show that o. niobe is strongly bivoltine if favourable conditions exist, with early and late summer emergences. this contrasts greatly with o. ariadne which is univoltine and only emerges in autumn (april/may). (lu & samways 2001) edge.qxd 2005/12/09 10:02 page 30 there are undoubtedly many other factors which control the population of o. niobe, such as predation and parasitism in the larval stage; the influence of the larval-ant relationship and the availability of sufficient ant colonies in proximity to the host plant concentrations. these factors are beyond the scope of the present study but will be the focus of future investigations. oviposition preferences of o. niobe all members of the genus orachrysops so far studied in any detail appear to specialise on a single larval host plant in the genus indigofera (refer to table 12). in the case of the four colonies of o. ariadne studied by lu & samways (2001, 2002a) they found that an abundance of the host plant was an essential pre-condition for a colony to become established and to be maintained. the abundance of i. erecta at the brenton blue butterfly reserve is greater than at any other place on the brenton peninsular, and indeed greater than all other sites investigated for the presence of o. niobe over a number of years. (ball 1997) an earlier study of oviposition sites found that the host plant and egg densities varied across the reserve (britton & silberbauer 1997), but were generally positively correlated. a similarly patchy distribution of host plant has been found in the present study, with the females concentrating their oviposition activities in areas containing the most host plant. in the present study, both in the november and february broods the majority of the eggs were laid on the host plants growing on the paths, and the density of eggs per plant and per stolon were also much higher on these host plants. the preference of the females of o. niobe for oviposition on host plants on or near the paths as opposed to plants in the burnt areas could have the following possible causes: the very high density of host plant in some areas of the paths attracted the females by sight and smell. predominance on the paths of horizontal stolons close to the ground which are preferred by the females. accessibility of the host plants on the paths no need to flutter between other plants. health and vigour of the host plants on the paths (see table 3). more favourable microclimate for larval survival may exist along the paths (sheltered, cool and humid). presence of an (as yet unknown) host ant in these areas. it was concluded from a study of the habitat of the endangered lycaenid aloeides dentatis dentatis (swierstra, 1909) that this butterfly had a preference for a disturbed community in an early stage of succession, which seemed to encourage colonisation by the host ant. (deutschländer & bredenkamp 1999). several observations were made during the present study of oviposition by another polyissn 0075-6458 31 koedoe 45/2 (2002) table 12 larval host plants of orachrysops species species host plant reference o. lacrimosa (bethune-baker, 1923) indigofera species unknown clark & dickson 1971 o. mijburghi henning & henning, 1994 indigofera species unknown williams 1996 o. niobe (trimen) i. erecta thunberg edge & pringle 1996 o. brinkmani heath, 1997 i. declinata e.meyer heath 1997 o. ariadne (butler) i. woodii var. laxa h. bolus lu & samways 2001 o. subravus henning & henning, 1994 i. woodii var. woodii h. bolus lu & samways 2001 edge.qxd 2005/12/09 10:02 page 31 ommatine lycaenid lampides boeticus (linnaeus, 1767) on the reserve and in all cases this was on the host plant rhynchosia chrysoscias bentham ex harvey, particularly in the burnt area where this climbing shrublet thrives. the niche differentiation between these two closely related butterfly species is clear and reduces potentially harmful larval competition. nectar sources nectar sources were hypothesised to be a limiting factor for maintaining colonies of o. ariadne in natal where only eight nectar plants were identified, with an overwhelming preference for two of them. (lu & samways 2001). the wider variety of flowers used by o. niobe (13 species) and their relative abundance at the study site leads one to surmise that this is not currently a limiting factor for the population. furthermore the wide choice of flowers of different symmetry, morphology, size and colour indicates that o. niobe is a generalist which does not fulfil any pollination function and is thus a nectar thief. the dominance of forbs and thus nectar plant visitations in the burnt area is probably because the woodier shrubs had not yet gained ascendancy in the succession process. influence of climate and microclimate from table 10 it can be seen that rainfall was well below the average for previous years during the period from october 2001 to march 2002, which is the emergence time for adult o. niobe. the increase in butterfly population that has been recorded in this study took place despite the adverse dry and hot conditions. it should be noted that moisture appears to be a critical factor in all orachrysops species studied so far (pringle 1997; lu & samways 2001) from a microclimate point of view, the shade factor and the angle of slope are two of the most important factors. it can be seen from table 3 that the butterfly larval host plant does better in intermediate shade (approx. 40 %) situations. generally the area where the paths were cut had a steeper slope than the burnt area and this made for a cooler microclimate as well as good drainage. overall comparison of burning versus cutting the summary in table 11 clearly demonstrates that the cut access paths created a superior habitat for both i. erecta to grow and for o. niobe to breed. the burnt area did however provide more male o. niobe activity, principally because it included the current patrolling path of the males. another important attraction and contribution of the burnt area was as a nectar source because of the prevalence of nectar rich forbs in the early successional stages after a fire. this may have some significance in the overall ecology of the butterfly and needs to be taken into account. conclusions the germination of the host plant indigofera erecta is not just fire induced. there was a doubling of the number of host plants on the access paths (area d), where no fire had taken place, during the observation period. indigofera erecta plants in the burnt areas a and c, and on the embankment (area f) had a higher mortality rate than those on the paths. the growth rate of the plants on the paths measured by increase in the number of stolons was an order of magnitude higher than in the burnt area. orachrysops niobe females had an overwhelming preference to lay eggs on the i erecta plants on or near the access paths and almost completely avoided the plants on the burnt area. there was a significant increase in the population of the butterfly between the first brood and the second brood—three to four times as many adults, and nearly five times as many eggs laid. orachrysops niobe uses a wide range of nectar sources, most of which are abundant at the reserve and not likely to be a limiting koedoe 45/2 (2002) 32 issn 0075-6458 edge.qxd 2005/12/09 10:02 page 32 factor in controlling the population, although the higher abundance of nectar sources in the burnt area may have some significance. the rainfall during this season has been significantly below the historical average so this factor could not have caused the population increase. the evidence suggests rather that both the proliferation of the host plant on the access paths and the increase in the butterfly population came about as a result of the access path cutting and maintenance activities. the areas that were burnt have produced relatively small numbers of i erecta plants on which hardly any eggs were laid. under natural conditions the type of disturbance caused by the cutting of paths is similar to that created by large and small herbivores grazing. burning is likely to cause a short-term negative impact on butterfly populations since it not only does not create conditions suitable for breeding, but the danger exists of larvae perishing in any fire occurring during the months from october to april. recommendations burning should only be carried out at the reserve in areas where the host plant is absent. the network of paths should be maintained and kept open through the next breeding season in order to continue monitoring of the host plant and to re-assess the butterfly population in november 2002. the morphology, reproductive biology (specifically the germination stimuli) and phenology of indigofera erecta should be studied in detail. the existence of a larval-ant association needs to be confirmed and investigated to find out if this could be a regulating factor on the populations of the butterfly. acknowledgements my thanks are due to cape nature conservation for permission to work within the brenton blue butterfly reserve, and to rhett hiseman in particular for loan of a gps instrument belonging to cape nature conservation. professor brian allanson is also thanked for his advice on methods to be used and for the use of laboratory equipment belonging to the knysna basin project. i am also grateful to renier terblanche, dr hamish robertson and ernest pringle for their guidance on methods, data analysis, review of drafts and general discussion and encouragement. wiehan properties’ role in making available their records from a weather station at brentonon-sea and brian young’s assistance in gathering and compiling of data from the weather station at the brenton blue butterfly reserve are also gratefully acknowledged. references ball, j.b. 1997. some notes on the brenton blue butterfly, orachrysops niobe (trimen) (lepidoptera, lycaenidae). unpubl. research report for the endangered wildlife trust of southern africa, johannesburg attachment 1: 22–27. cocks, m.p. & w.d stock. 1997. heat stimulated germination in relation to seed characteristics in fynbos legumes of the western cape province, south africa. south african journal of botany 63(3): 129–132. britton, d.r. & l.x. silberbauer. 1997. the lifehistory, ecology and conservation of the brenton blue butterfly, orachrysops niobe (trimen) (lycaenidae) at brenton-on-sea. unpubl. research report for the endangered wildlife trust of southern africa, johannesburg. clark, g.c. & c.g.c. dickson. 1971. life histories of the south african lycaenid butterflies. cape town: purnell. deutschländer, m.s. & g.j. bredenkamp. 1999. importance of vegetation analysis in the conservation management of the endangered butterfly aloeides dentatis dentatis (swierstra) (lepidoptera, lycaenidae). koedoe 42(2): 1–12. edge, d.a. & e.l. pringle. 1996. notes on the natural history of the brenton blue orachrysops niobe (trimen) (lepidoptera: lycaenidae). metamorphosis 7(3): 109–20. goldblatt, p. & j. manning. 2000. cape plants: a conspectus of the cape flora of south africa. pp. 486–492. in: strelitzia 9. cape town: national botanical institute of s a & missouri botanical garden. heath, a. 1997. description of a new species of orachrysops vári from the western cape province of south africa (lepidoptera: lycaenidae). metamorphosis 8(1): 26–32. issn 0075-6458 33 koedoe 45/2 (2002) edge.qxd 2005/12/09 10:02 page 33 henning, g.a. & s.f. henning. 1995. updating the status of south african red data butterfly species. metamorphosis 6(2): 96– 98. hiseman, r.e. 2000. brenton blue butterfly reserve management report. internal report for cape nature conservation (unpublished, dated 30 november 2000). jeffrey, d.j., p.m. holmes & a.g. rebelo. 1988. effects of dry heat on seed germination in selected indigenous and alien legume species in south africa. south african journal of botany 54(1): 28–34. lu, s-s. & m.j. samways. 2001. life history of the threatened karkloof blue butterfly, orachrysops ariadne (lepidoptera: lycaenidae). african entomology 9(2): 137–151. lu, s-s. & m.j. samways. 2002a. behavioural ecology of the karkloof blue butterfly, orachrysops ariadne (lepidoptera: lycaenidae) relevant to its conservation. african entomology 10(1): 137–147. lu, s-s. & m.j. samways. 2002b. conservation management recommendations for the karkloof blue butterfly, orachrysops ariadne (lepidoptera: lycaenidae). african entomology 10(1): 149–159. lubke, r.a., d.b. hoare & j.e. victor. 1996. the vegetation and floristics of the habitat of the brenton blue butterfly. unpubl. research report for the endangered wildlife trust of southern africa, johannesburg. milewski, a.v. 2002. elephants and fynbos. veld & flora march 2002: 28. new, t.r. 1997. butterfly conservation. melbourne: oxford university press. pringle, e.l. 1997. butterfly breeding areas—a microclimate perspective. metamophosis 8(2): 82–84. robertson, h.g. 1997. report on the ants at brenton-on-sea and their possible interactions with the brenton blue butterfly, orachrysops niobe. unpubl. research report for the endangered wildlife trust of southern africa, johannesburg. robertson, h.g. 1998. report on field trip to brenton-on-sea from 15–26 november 1998. internal report for life sciences division, south african museum, cape town (unpublished). robertson, h.g. 2000. report on field trip to brenton-on-sea from 6-10 november 2000. internal report for life sciences division, south african museum, cape town. schutte-vlok, a.l. 2001. environmental management plan for the brenton blue butterfly reserve. internal report for cape nature conservation (unpublished). steenkamp, c. & r. stein. 1999. the brenton blue saga—a case study of south african biodiversity conservation. johannesburg: endangered wildlife trust/nedbank/danish co-operation for environment and development. trimen, r. 1862–1864. on some new species of south african butterflies. transactions of the entomological society, london, 3rd series vol i: 279–291. williams, m.c. 1996. report on research findings concerning the life history and ecology of the brenton blue (orachrysops niobe). metamorphosis 7(1): 3–7. koedoe 45/2 (2002) 34 issn 0075-6458 edge.qxd 2005/12/09 10:02 page 34 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice abstract introduction methods results discussion management implications acknowledgements references about the author(s) ian a. russell conservation services, south african national parks, sedgefield, south africa rodney m. randall private, sedgefield, south africa david zimmerman conservation services, south african national parks, kimberley, south africa danny govender conservation services, south african national parks, skukuza, south africa citation russell, i.a., randall, r.m., zimmerman, d. & govender, d., 2019, ‘outbreak of avian botulism and its effect on waterbirds in the wilderness lakes, south africa’, koedoe 61(1), a1553. https://doi.org/10.4102/koedoe.v61i1.1553 original research outbreak of avian botulism and its effect on waterbirds in the wilderness lakes, south africa ian a. russell, rodney m. randall, david zimmerman, danny govender received: 16 aug. 2018; accepted: 05 sept. 2019; published: 31 oct. 2019 copyright: © 2019. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract avian botulism has been reported in many areas worldwide, particularly in north america, where at times it has resulted in die-offs of hundreds of thousands of waterbirds. by contrast, there are few reported cases in wild waterbird communities in south africa. mass die-offs of waterbirds in the wilderness lakes, south africa, were first observed in january 2015, raising concerns about the effect of disease on the status of waterbird communities in this ramsar site. surveys of bird carcasses were undertaken between 2015 and 2017. an estimated 1115 individuals from 23 species were affected, with sick individuals displaying symptoms of avian type c, c and d or e botulism. the highest mortality was amongst red-knobbed coot (fulica cristata), cape shoveler (anas smithii) and yellow-billed duck (anas undulata), respectively comprising 60%, 18% and 9% of affected individuals. cape shoveler was found to be particularly susceptible to the disease. deaths occurred mainly during summer and autumn (november–april). no mortalities were recorded amongst 44 waterbird species, of which common moorhen (gallinula choropus), african purple swamphen (porphyrio madagascariensis) and reed cormorant (phalacrocorax africanus) are widespread and abundant. the reasons for the outbreaks remain elusive, as environmental changes characteristic of a botulism outbreak, notably a decline in water level and dissolved oxygen, did not differ substantially from earlier years when no disease outbreaks were observed. the role of fish, and in particular the recently introduced common carp (cyprinus carpio) in the progression of the disease, remains speculative, although worthy of investigation. the removal of carcasses for disease control should be continued. conservation implications: the long-term effect of repeated outbreaks of avian botulism on the abundance of susceptible waterbird species in the wilderness lakes is of concern. the previous regular high abundance of duck species, particularly yellow-billed duck and cape shoveler, was the initial reason for the wilderness lakes being declared a ramsar site. the effect of ongoing high disease-related mortalities may, in part, prevent these wetlands from continuing to regularly support globally significant populations of some waterbird species. the containment of the disease must be attempted by the regular collection and removal of dead birds and fish during the outbreak periods of november–june, and carcasses should be disposed of off-site. further testing of affected individuals should be undertaken and the monitoring of environmental variables and affected individuals continued to improve the understanding of the drivers and progression of the disease. keywords: botulism; disease; wetlands; waterbirds; ramsar; wilderness lakes. introduction avian botulism is a non-infectious bird disease resulting from the ingestion of toxins produced by the gram-positive bacterium clostridium botulinum (rocke & friend 1999). seven different types of botulism neurotoxins occur, labelled a to g (collins & east 1998), as well as mosaics that have characteristics of two toxins (rocke & friend 1999). avian botulism outbreaks are typically caused by type c (wobeser 1997), mosaic c/d (anza et al. 2014) or type e (getchell & bowser 2006; rocke & friend 1999). fish die-offs are typically caused by type e (cromie et al. 2012). clostridium botulinum is reportedly widespread in wetland sediments where outbreaks are common, and typically requires a source of protein, warm temperatures and low oxygen conditions to activate and produce toxin (espelund & klaveness 2014). the salinity and ph of sediments and water may also affect the probability of botulism outbreaks in aquatic ecosystems (rocke & friend 1999). the disease in birds is characterised by a flaccid paralysis of the muscles and is invariably fatal (rocke & friend 1999). death typically results from respiratory failure or drowning. avian botulism has been reported from several territories worldwide (rocke 2006), including countries in both north america and south america (rocke & friend 1999; wobeser 1997), europe (włodarczyk et al. 2014; woudstra et al. 2012), africa (blaker 1967), australia (grubb 1964) and new zealand (martinovich et al. 1972). botulism is one of the most common causes of death in waterfowl in north america (smith 1976; wobeser 1997) where it has resulted in die-offs of hundreds of thousands of waterfowl in single events (rocke & bollinger 2007). by contrast, there are only three published accounts comprising one peer-reviewed article (blaker 1967) and two semi-popular articles (ramollo 2016; van heerden 1972) of separate, smaller outbreaks in south africa, suggesting that the disease is far less prevalent in southern africa than in north america and possibly europe. in this study, we document the repeated outbreak of disease affecting waterbirds and fish in the wilderness lakes, and discuss its relevance to the conservation of these aquatic ecosystems. methods study area the wilderness lakes system (33°59′ to 34°02′ s and 22°35′ to 22°46′ e) is located on the southern coastline of south africa, and includes the touw estuary and three interconnected estuarine lakes (rondevlei, langvlei and eilandvlei) (figure 1). the touw estuary is a temporarily open-closed system, with closure of the estuary mouth dictated mostly by marine condition and river flow (whitfield, allanson & heinecken 1983). under natural conditions, the breaching of the estuary would occur when the water level exceeds the height of the sandbar at the estuary mouth, which at times can attain 3.5 m above mean sea level (amsl). natural breaching, however, seldom occurs with the estuary being artificially breached when water levels are between 2.1 m amsl and 2.4 m amsl, in order to prevent the flooding of residential and other developed areas on the estuary floodplain (south african national parks 2012). fijen and kapp (1995) estimated that under pristine conditions the estuary would have been open approximately 40% of the time. open phases have, however, been reduced to an average of 30% (1991–2017) (sanparks unpublished data) as a result of both artificial breaching and freshwater deprivation. estuary breaching and closure results in substantial variability in water levels and physico-chemical properties in the lakes. figure 1: map of study area showing relative position of estuarine lakes within the wilderness lakes complex. rainfall occurs year-round, with a mean of between 900 mm/year and 1000 mm/year in the upper catchments (adamson 1975). one perennial and two intermittently flowing rivers provide surface freshwater flow to the lake system. the lower catchment of the larger (catchment ~89.1 km2) perennial touw river and the majority of the catchments of the smaller, intermittently flowing duiwe river (~32.5 km2) and langvlei spruit (~17.8 km2) support intensive, mostly livestock-based agricultural practices (filmalter & o’keeffe 1997). river water has a high humic matter content, is low in electrolytes (robarts & allanson 1977) and is acidic (median ph in touw river = 5.1, duiwe river = 6.7 in 1991–2014 [sanparks unpublished data]). submerged aquatic plants are widespread and abundant in the shallower (< 3 m) littoral areas of the lakes. dominant taxa are fennel-leaved pondweed (potamogeton pectinatus), water hornwart (ceratophyllum demersum), stoneworts (charophyta) and filamentous algae (howard-williams & liptrot 1980; weisser & howard-williams 1982; whitfield et al. 1983). stands of emergent aquatic plants occur on the lake margins and floodplains, with abundant species including common reed (phragmites australis), clubrush (schoenoplectus scirpoides) and bullrush (typha capensis). the fish community consists of estuarine, euryhaline marine and freshwater alien species (hall, whitfield & allanson 1987; olds et al. 2011; russell 1996; whitfield 1984). the lakes, and in particular langvlei and rondevlei, support abundant and diverse waterbird communities (boshoff, palmer & piper 1991a, 1991b, 1991c; russell, randall & hanekom 2014). the wilderness lakes system is located in the garden route national park. the lakes and their interleading channels were designated as a wetland of international importance, or ramsar site, in 1991 (ramsar convention bureau 2019). abundance of living and dead waterbirds surveys of the abundance of all waterbirds were undertaken biannually during summer (january–february) and winter (july–august) from 2005 to 2017. surveys were undertaken from a boat over two days by four observers using binoculars. the boat route used enabled the observation of all open water areas and exposed sandbanks and mudflats. the first bird carcasses were observed and reported by members of the public in january 2015 in front of a bird hide on the northern shoreline of rondevlei. shortly thereafter, during regular boat-based surveys on the lakes by south african national parks (sanparks) personnel to assess aspects of water quality and waterbird abundance, the scale and extent of the outbreak became more apparent, with the observation of multiple bird and fish carcasses on eilandvlei, langvlei and rondevlei. to monitor the carcass occurrence, regular boat-based surveys were undertaken of all the waterbodies between january 2015 and may 2017, following routes that enabled observation of all the open water areas. twenty surveys were conducted on eilandvlei over a period of 256 days, 37 on langvlei over 462 days and 56 on rondevlei over 528 days. these equate to surveys being conducted, on average, once every 13, 12 and 9 days respectively, although, during the height of the outbreak, surveys were conducted at shorter intervals. intermittent surveys were undertaken on the touw estuary and adjacent swartvlei system. all observed and accessible bird and fish carcasses were collected and, where possible, identified. live birds (approximately 30) with paralysis and those that were unable to walk or fly were captured and identified. those in an advanced state of paralysis and in the process of drowning when captured were euthanised. all collected dead and sick birds, and all dead fish, were removed from the lakes. all carcasses not used for laboratory analyses were disposed of by burying at a site removed from the lakes in an attempt to try and contain the disease. disease identification in waterbirds state veterinarians were consulted to provide guidance on the probable cause of the bird deaths. three dead and one paralysed yellow-billed duck (anas undulata) were collected in early march 2015. a full post-mortem was conducted, including a histopathological examination of the lung, liver, intestine, pancreas, proventriculus and brain tissues. organ swabs were tested by means of polymerase chain reaction (pcr) for avian influenza and newcastle disease. bacterial cultures were undertaken of organ fluids. four frozen whole carcasses of affected and euthanased waterbirds comprising two cape shoveler (anas smithii), one red-knobbed coot (fulica cristata) and one blacksmith lapwing (vanellus armatus), collected in march 2015, and two water samples from rondevlei and langvlei were submitted to the agricultural research council-onderstepoort veterinary institute (pretoria) by sanparks. mouse bioassays for the toxin of c. botulinum were conducted on samples of both water and waterbird intestinal contents. post-mortem and histopathological examinations of heart, small intestine, gizzard, pancreas, aorta, liver, trachea, proventriculus, kidneys, lungs, cerebrum, cerebellum and testes were conducted. bacterial cultures were undertaken of pooled liver swabs. tissue samples of major organs and muscles of three disease-affected and euthanased waterbirds, namely, one each of yellow-billed duck, great crested grebe (podiceps cristatus) and southern pochard (netta erythrophthalma), and one recently dead adult fish, a white steenbras (lithognathus lithognathus), were harvested by sanparks veterinary wildlife services on 14 december 2015. all tissue samples were preserved in 10% formalin and submitted to the wildlife pathology research of the national zoological gardens (pretoria) for histopathological examination. presence of toxic algae and diatoms water samples were collected in polypropylene bottles from nine localities (2 × rondevlei, 2 × rondevlei-langvlei channel, 1 × langvlei, 1 × langvlei-eilandvlei channel, 2 × eilandvlei, 1 × serpentine channel) on 20 february 2015, during the height of the first disease outbreak. sample preparations and species identifications were undertaken by the stellenbosch office of the council for scientific and industrial research (csir). all water column algae were identified using a compound microscope (carl zeis, germany) at 1250 × magnification and keys of truter (1987), wehr and sheath (2003), van vuuren et al. (2006) and taylor, harding and archibald (2007). diatom samples were heated in a sulphuric acid and potassium dichromate solution to clear them of organic matter and thereafter rinsed, diluted and mounted in a pleurax medium for microscopic examination. samples were sedimented in an algae chamber and evaluated using the strip-count method (american public health association, american water works association & water pollution control federation 1992). environmental parameters prior to and during the disease outbreak water level data for the lakes were extracted from the unpublished databases of the south african department of water and sanitation. temperature (°c), salinity, dissolved oxygen (mg/l) and ph of lake waters were measured quarterly by sanparks personnel at 30 cm depth at five localities in each lake using ysi model 30 s-c-t, model 550a o2 and model 60 ph meters. the standing biomass of submerged plants was determined during may and june from 1992 to 2017. in rondevlei and langvlei, assessments between 2000 and 2004 were undertaken biennially. four littoral transects extending from the inner edge of the emergent macrophyte zone to the 2 m depth contour were randomly positioned around each lake each year. the above-ground portions of aquatic plants were collected at five 0.0625 m2 sample points along each transect with a submerged macrophyte sampler (howard-williams & longman 1976). living plant tissue was oven-dried at 55 °c for approximately 1 week and thereafter weighed on an electronic balance (sartorius) to the nearest gram. ethical considerations this article followed all ethical standards for research without direct contact with human or animal subjects. results waterbirds affected a total of 1115 dead or disease-affected waterbirds were recorded on rondevlei, langvlei and eilandvlei between january 2015 and may 2017. this constituted 20% of the number of individuals that, on average, occur on these waterbodies. twenty-three species from 13 families were affected (table 1), with the highest mortality being amongst red-knobbed coot (670 individuals), cape shoveler (204) and yellow-billed duck (100), respectively comprising 60%, 18% and 9% of affected individuals. table 1: population statistics and number of mortalities of waterbird species affected by botulism. there is a strong similarity for most species between the percentage of individuals affected per species and the percentage representation in the waterbird community (table 1). this is particularly apparent in two of the three most affected species, namely red-knobbed coot and yellow-billed duck, where the percentages of community representation and disease-affected individuals are identical. by contrast, cape shoveler appeared to be particularly susceptible to the disease, with the percentage of individuals affected (18%) far exceeding community representation (4%). the number of dead and dying cape shoveler (204 individuals), a regularly occurring species on the lakes, is a substantial portion (78%) of the average population (261 individuals). although the number of dead greater flamingo (phoenicopterus ruber) (12 individuals) exceed the average population size (six individuals), this does not necessarily suggest above average susceptibility to the disease, as this species is an intermittent, mostly winter migrant to the wilderness lakes, and abundances can vary substantially from year to year. greater flamingo were a typical abundant in 2016, with arrivals in late summer, and 453 individuals were counted in the winter 2016 surveys. relatively abundant species that were affected by the disease, but where the percentage of affected individuals was notably lower than community representation, included both little grebe (tachybaptus ruficollis) and great crested grebe (table 1). the highest number of waterbird deaths over the 3-year study period occurred on rondevlei (570), followed by langvlei (473) and eilandvlei (72) (table 2). no affected waterbirds were observed on either the touw estuary or the adjacent swartvlei system. the highest mortality occurred on langvlei in 2015 and on rondevlei in 2016. mortalities were recorded only on langvlei in 2017 (table 2). table 2: the number of disease-affected waterbirds located in rondevlei, langvlei and eilandvlei in the three different botulism outbreak periods encompassing january–june 2015, november 2015 – august 2016 and january–may 2017. there was temporal variability in the disease outbreak, with the majority (95%) of waterbird deaths occurring in the summer and autumn months of november–april, with no deaths recorded in the late winter months of september and october (figure 2a). figure 2: number of (a) waterbird and (b) fish mortalities recorded in the wilderness lakes per month between january 2015 and july 2017. no mortalities were recorded amongst 44 additional waterbird species (table 3), of which common moorhen (gallinula choropus), african purple swamphen (porphyrio madagascariensis) and reed cormorant (phalacrocorax africanus) are widespread and abundant. table 3: population statistics of waterbird species not affected by botulism. for waterbird communities surveyed biannually in rondevlei, langvlei and eilandvlei over 10 years (2005–2014) prior to the initial outbreak of botulism. symptoms seen in clinically affected birds most affected birds were found in areas of dense flooded plants, or on the shoreline, and were frequently grouped in one or more localities on a waterbody. birds that were on the shoreline, or propped up by emergent plants, were generally hunched over and unable to stand. all affected individuals were either reluctant or unable to fly, and when attempting to do so were only able to propel themselves for very short distances across the water surface or mudflats with their wings. most displayed paralysis of the neck and were unable to hold up their head. this was particularly prevalent in the longer-necked species such as ducks and cormorants, many of which, when located, were struggling to hold their head above water. when on land, none were able to stand, and when attempting to walk exhibited a floppy stumbling gate for very short distances. most were unable to walk, and either sat hunched over or lay laterally recumbent with their legs and necks stretched out. in extreme cases, individuals seemed completely paralysed and unresponsive. most had their eyes either partially or fully closed. fishes affected a total of 124 dead fish from eight species were observed in the wilderness lakes over the same period when the waterbird deaths occurred. all dead fish, with the exception of one cape stumpnose (rhabdosargus holubi) and one oval moonie (monodactylus falciformis), were large (> 30 cm total length). common carp (cyprinus carpio) was the most abundant species of dead fish in both eilandvlei (48%) and langvlei (84%), as well as collectively across all waterbodies (72%) (table 4). observed fish mortality was highest in langvlei (72%) and comparatively uncommon in rondevlei (5%) (table 4). temporal variability in fish deaths coincided with that of waterbirds, with dead individuals observed from november to may (figure 2b). table 4: the number of dead fish located in rondevlei, langvlei and eilandvlei in the three different botulism outbreak periods encompassing january–june 2015, november 2015 – august 2016 and january–may 2017. post-mortal findings laboratory assessments undertaken by state veterinarians on four yellow-billed duck indicated minimal to severe autolysis. all of the birds’ gastrointestinal tracts were empty. pcr tests were negative for both newcastle disease and avian influenza. no significant bacteria isolates were cultured from body fluids. a histological assessment of lung, liver, intestine, pancreas and brain tissue did not indicate any diagnostic significant microscopic lesions. oedema intermingled with large numbers of bacteria and non-cellular debris within the parabronchi, most probably from aspirated water, was indicative of drowning. similarly, a post-mortem and histopathological examination of carcasses submitted to the arc-onderstepoort veterinary institute and wildlife pathology research revealed no specific lesions that could be associated with mass waterbird deaths. water and intestinal samples were negative on the mouse bioassay for c. botulinum toxins. these samples were also negative on anaerobic culture for clostridium species or any other significant bacteria species. based on the negative histopathology and laboratory results and clinical symptoms of affected birds, a presumptive diagnosis of avian botulism was made. the single fish examined histologically, a white steenbras, only revealed epithelial hyperplasia of the gills, indicating subacute bronchitis, possibly because of altered environmental conditions, with death likely as a result of respiratory failure. environmental conditions during disease outbreak all of the waterbodies are interconnected; as a result, variability in their water levels is similar. within the lake system, this variability is depicted by data from rondevlei (figure 3). breaching of the touw estuary, both natural and artificial, and resultant reductions in water levels throughout the lakes system is a common occurrence (figure 3). post-breaching reduction in water level occurred just prior to the first disease outbreak (07 december 2014). the following three mechanical breachings during the disease period had a relatively minor effect on water levels. subsequent natural breaching on 30 august 2015 and mechanical breaching on 03 september 2016 resulted in larger and longer duration reductions in water levels. post-breaching reductions in water levels just prior to and during the outbreak period are, however, not atypical for these waterbodies with, as an illustration, 19 breachings in the prior 13 years resulting in either comparable or larger and longer duration reductions in lake water levels (figure 3). figure 3: water level relative to sea level in rondevlei from 2002 to 2017. arrows indicate times of breaching of the touw estuary mouth (black = artificial breaching, red = natural breaching). shaded area indicates the period when botulism-related deaths of waterbirds were recorded in the wilderness lakes. seasonal variations in water temperature were similar in all waterbodies (figure 4a), with the highest temperature occurring in january (summer) and the lowest in july (mid-winter). variability and extremes in water temperature during the disease outbreak did not notably differ from that recorded in earlier years (figure 4a). figure 4: time series plots of (a) mean water temperature, (b) salinity, (c) dissolved oxygen and (d) ph collected in january, april, july and october of each year from 2004 to 2017 in rondevlei (blue), langvlei (red) and eilandvlei (green). horizontal dotted lines indicate environmental limits conducive for botulism occurrence for temperature (> 15 °c), salinity (< 5) and ph (7.5 < ph < 9.0) (rocke & friend 1999). the shaded area indicates the period when botulism-related deaths of waterbirds were recorded in the wilderness lakes. salinity in the wilderness lakes fluctuated temporally depending on the timing and volume of inflows of fresh river water, saline marine waters and brackish groundwater, the state of the estuary mouth, the movement of water between lakes, and the timing and rate of water loss through evaporation, evapotranspiration and groundwater movement. all of the lakes became progressively less saline until early 2008 after which an extended low rainfall period resulted in increases in salinity, which stabilised mostly within the range of 5–10 between 2010 and 2017 (figure 4b). rondevlei is typically more saline than both langvlei and eilandvlei. variability in salinity did not differ substantially in the period of disease outbreak (2015–2017) from the prior 5 years, and from 2010 onwards never fell below 5, which rocke and friend (1999) describe as more conducive for avian botulism outbreaks. dissolved oxygen in the lakes is naturally variable, with higher oxygen levels frequently occurring in the cooler winter months, and relatively lower dissolved oxygen levels occurring during the warmer summer months. this distinctive variability also occurred during the period of disease outbreak, and with both upper and lower dissolved oxygen levels not being atypical, or exceeding those recorded in the 11 years prior to the disease outbreak (figure 4c). the ph within the lakes mostly fluctuated within the range of 7–9 (figure 4d), and was rarely lower than 6.5, below which botulism outbreaks do not occur (rocke & friend 1999), except for short periods (< 3 months) in eilandvlei following high freshwater inflows. variability in ph mostly within the range conducive for botulism outbreaks did not differ between the period of disease outbreak and earlier years (figure 4d). little to no algae were detected in water samples collected during the height of the first disease outbreak. the dominant diatom was pleurosigma salinarum, an indicator of brackish water. toxic cyanobacteria detected were oscillatoria tenue and oscillatoria princeps. these were only observed in samples from the rondevlei-langvlei channel, and abundances were too low to cause toxicity in the water (p. oberholster pers. comm., 09 november 2015). temporal variation in the biomass of submerged aquatic plants was substantial in all three estuarine lakes (figure 5a–c). declines in the standing biomass of aquatic plants, and consequently potential increases in decaying organic plant matter, occurred prior to the initial 2015 avian disease outbreak in both rondevlei (figure 5a), and to a lesser extent in langvlei (figure 5b). no comparable substantial declines were recorded in eilandvlei (figure 5c). the variability in submerged aquatic plant biomass recorded in rondevlei and langvlei since 2015 was, however, not atypical, with periodic substantial declines having occurred in all waterbodies in the years prior to the avian disease outbreak, notably in 2000–2003 in eilandvlei and in 2008–2010 in all three lakes. figure 5: the biomass of submerged plants determined annually in the period of maximum biomass (may–june) in (a) rondevlei, (b) langvlei and (c) eilandvlei from 1991 to 2017. discussion a diagnosis of avian botulism is frequently based on a combination of symptoms displayed by sick birds, and the absence of specific post-mortal findings (rocke & bollinger 2007). patterns of mortality and behaviour of affected, and in extremis, waterbirds on the wilderness lakes were classic for avian botulism, as described by rocke and friend (1999) and rocke and bollinger (2007), which along with the absence of any obvious lesions in dead birds, and the absence of toxic algae, indicate that a diagnosis of avian botulism is reasonable. the absence of a positive mouse bioassay does not invalidate this diagnosis, as although it is still considered the most sensitive test for all c. botulinum types (rocke & bollinger 2007), false negatives may occur (rocke et al. 2004; thomas 1991). the number of observed dead and sick waterbirds is substantial, comprising approximately 20% of the average number of all susceptible waterbird species on the affected lakes. this is quite probably an underestimation, as assessments of total waterbird mortality based only on carcass retrieval can result in the underestimation of actual mortality by between three (cliplef & wobeser 1993) and 10 times (stutzenbaker et al. 1986 cited in rocke & bollinger 2007). the effect of the disease on dabbling ducks in general is of concern, and particularly cape shoveler, a near-endemic species to southern africa (hockey, dean & ryan 2005), the recorded deaths of which constitute approximately 75% of their average population on the affected lakes. one obvious question relevant to the management of the wilderness lakes is why has there been an outbreak of botulism here and now? clostridium botulinum spores occur naturally in most wetland sediments where they can persist for decades and can be found in the body tissues of most wetland biota, including healthy birds (rocke & friend 1999). the mechanisms triggering spore germination and bacterium growth leading to botulism outbreaks remain poorly understood (espelund & klaveness 2014), although several predisposing environmental factors have been proposed. the prevalence of higher environmental botulism has been linked to falling water levels, higher summer surface-water temperatures, decaying organic material, water ph of between 7.5 and 9.0, water salinity of below 5, and low dissolved oxygen levels (perez-fuentetaja et al. 2006; rocke, euliss & samuel 1999; rocke & samuel 1999). therefore, many of the environmental conditions prevalent at the time of the outbreak were not conducive to botulism outbreaks, and the lakes were not undergoing any atypical fluctuations compared to historical data. water level fluctuations, temperature, salinity, dissolved oxygen and level of decaying senescent aquatic plants were all very consistent with years when there were no disease outbreaks. however, the one environmental factor that does differ from the years prior to the botulism outbreaks is the introduction and proliferation of alien common carp (olds et al. 2011), the most abundant fish species recorded in botulism concurrent mortalities in the wilderness lakes. common carp, a primarily freshwater species, was first observed in eilandvlei in 2005 (personal observation made by i.a. russell), when water salinity was relatively low, although the exact date and place of introduction is unknown. they now occur in all lakes in the wilderness system. the role that common carp play in triggering botulism outbreaks by scavenging in the benthic layer where c. botulinum may have proliferated in vegetation and insects, mobilising organic-rich, potentially anoxic sediments during feeding (skelton 2001) or perpetuating outbreaks with the carcasses of large individuals providing a decaying organic source, remains speculative, although worthy of investigation. all birds, except vultures (cohen et al. 1969), are thought to be susceptible to ingested avian botulism neurotoxin, although interspecific differences in susceptibility may occur (bollinger 2011), with diet and feeding patterns playing a role in their exposure to the toxin. in the wilderness lakes, the cape shoveler, a dabbling duck (brickell 1988), was found to be highly susceptible, whereas many species, particularly most piscivorous species and waders were unaffected. this is consistent with the findings of rocke (2006) that dabbling ducks are amongst the waterbirds most susceptible to type c/d botulism, with diving ducks seemingly less affected. the particularly high mortality of cape shoveler also mirrors the observation that mallards and shovelers are the species at greatest risk from type c (rocke & bollinger 2007). a high mortality amongst red-knobbed coot in the wilderness lakes is similar to the findings of anza et al. (2016) who found coot to be highly vulnerable to type c/d botulism, with the source of the toxins thought to be invertebrates on plants and carcasses. both similarities and differences occur in the species affected in this and earlier reported botulism outbreaks amongst waterbirds in south africa. whereas van heerden (1972) similarly reported mostly duck species to be affected in a botulism outbreak in a pan in central south africa (welkom, south africa), along with red-knobbed coot, grey-headed gull (larus cirrocephalus), greater flamingo, blacksmith lapwing and black-winged stilt (himantopus himantopus), observed mortalities differed from that in the wilderness lakes in that both common moorhen and waders in the family scolopacidae were also affected. similarly, blaker (1967) also recorded the mortality of several small waders at the strandfontein sewage works (cape town, south africa). the botulism outbreak recorded by blaker (1967), similar to that in the wilderness lakes, also affected several duck species, although it differed in that while the cape shoveler was highly susceptible in the wilderness lakes, only one individual (1% of the population) of this species was reported by blaker (1967), with cape teal (anas capensis), another dabbling duck (brickell 1988), being the most affected species. the susceptibility of greater flamingo also appears to differ between outbreaks, with a high mortality reported on kamfers dam (kimberley, south africa) (ramollo 2016), whereas few deaths of this species occurred on wilderness lakes, despite being abundant in the summer of 2016 (± 450 individuals). similarly, no deaths of greater flamingo were recorded by van heerden (1972) despite being common during that botulism outbreak. greater flamingo in europe are considered by anza et al. (2016) to be less susceptible to type c/d botulism. differences in the species affected between outbreaks suggest variances in the mode of disease transmission. the transfer of toxins to waterbirds is thought to occur mainly through zooplankton or invertebrate food items that have assimilated toxins (rocke & friend 1999). fish may also act as vectors for botulinum type e toxin, with affected fish being consumed by piscivorous or scavenging birds (getchell & bowser 2006), or their carcasses providing a protein source for bacterial growth and consequently toxin production (eklund et al. 1984), thereby affecting birds via the carcass-maggot cycle. the presence of dead fish during botulism outbreaks in the wilderness lakes suggests that botulinum type e toxin may be implicated. however, rather unusually, most piscivorous birds, including the abundant reed cormorant, african darter (anhinga rufa) and black-necked grebe (podiceps nigricollis), as well as herons, terns and most kingfishers, were unaffected in this instance. more remains to be learned about the origin, strain, drivers and transmission of the disease in the wilderness lakes, and the cause and ecology of the disease remains speculative. management implications the long-term effect of repeated outbreaks of avian botulism on the abundance of susceptible waterbird species in the wilderness lakes is of concern. the previous regular high abundance of duck species, particularly yellow-billed duck and cape shoveler, was the initial reason for the wilderness lakes being declared a ramsar site. the effect of ongoing high disease-related mortalities may, in part, prevent these wetlands from continuing to regularly support globally significant populations of some waterbird species. the containment of the disease must be attempted by the regular collection and removal of dead birds and fish during the outbreak periods of november–june, and the disposal of carcasses off-site. further testing of affected individuals should be undertaken and monitoring of environmental variables and affected individuals continued, to improve the understanding of the drivers and progression of the disease. serotyping c. botulinum in the soil of various important wetlands in southern africa will assist in detecting hotspots for potential outbreaks. the regular monitoring of environmental conditions will increase predictive capability, informing both proactive management of these important wetland and lake environments, to prevent outbreaks and to allow for early detection if an outbreak occurs, and therefore enable rapid response to minimise environmental contamination. acknowledgements the authors wish to thank paul oberholster for identification of algae species, emily mitchell for assistance with the histopathology and members of the garden route national park ranger team, notably davie antas, nelson tyhali, petrus mitchell, donavan mckeith, charles april and jonathan britton, for undertaking the unpopular task of carcass location and disposal. the authors also wish to thank two anonymous reviewers for their helpful comments on an earlier draft of the article. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions r.m.r. initiated and oversaw surveys of dead birds in 2015. he also arranged a visit by state veterinarians. i.a.r. participated in dead bird surveys from 2015 to 2017, undertook water quality and aquatic plant surveys and maintained databases on water heights and estuary breaching. he undertook the data analysis and wrote the article. d.g. arranged for botulism testing. d.z. collected tissue samples and arranged for some histopathological examinations. r.m.r., d.g. and d.z. proofread and contributed to the article. funding information the south african national parks (sanparks) funded this research. the state veterinarian office of the western cape government undertook infield inspections, post-mortem and histopathological examinations, and provided advice and diagnoses. the wildlife pathology research of the national zoological gardens undertook post-mortem and histopathological examinations. the agricultural research council-onderstepoort veterinary institute undertook mouse bioassays, bacterial cultures and histopathological examinations. the stellenbosch office of the council for scientific and industrial research (csir) identified algae species. data availability statement data are available from the first author upon request. disclaimer the views and opinions expressed in this article are those of the authors and not an official position of sanparks or any institute or organisation that undertook tests in support of this study. references adamson, p.t., 1975, extension of monthly runoff records in the catchments of the wit els, diep and karatara rivers – wilderness – cape province, department of water affairs, pretoria. american public health association, american water works association & water pollution control federation, 1992, standard methods for the examination of water and wastewater, 18th edn., american public health association, american water works association & water pollution control federation, washington, dc. anza, i., skarin, h., vidal, d., lindberg, a., båverud, v. & mateo, r., 2014, ‘the same clade of clostridium botulinum strains is causing avian botulism in southern and northern europe’, anaerobe 26, 20–23. https://doi.org/10.1016/j.anaerobe.2014.01.002 anza, i., vidal, d., feliu, j., crespo, e. & mateoa, r., 2016, ‘differences in the vulnerability of waterbird species to botulism outbreaks in mediterranean wetlands: an assessment of ecological and physiological factors’, applied and environmental microbiology 48(10), 3092–3099. https://doi.org/10.1128/aem.00119-16 blaker, d., 1967, ‘an outbreak of botulinus poisoning among waterbirds’, ostrich 38(2), 144–147. https://doi.org/10.1080/00306525.1967.11783617 bollinger, t.k., evelsizer, d.d., dufour, k.w., soos, c., clark, r.g., wobeser, g. et al., 2011, ecology and management of avian botulism on the canadian prairies. prairie habitat, joint venture science committee. boshoff, a.f., palmer, n.g. & piper, s.e., 1991a, ‘spatial and temporal abundance patterns of waterbirds in the southern cape province. part 1: diving and surface predators’, ostrich 62(3–4), 156–177. boshoff, a.f., palmer, n.g. & piper, s.e., 1991b, ‘spatial and temporal abundance patterns of waterbirds in the southern cape province. part 2: waterfowl’, ostrich 62(3–4), 178–196. boshoff, a.f., palmer, n.g. & piper, s.e., 1991c, ‘spatial and temporal abundance patterns of waterbirds in the southern cape province. part 3: wading birds’, ostrich 62(3–4), 197–214. brickell, n., 1988, ducks, geese and swans of africa and its outlying islands, frandsen publishers, fourways. cliplef, d.j. & wobeser, g., 1993, ‘observations on waterfowl carcasses during a botulism epizootic’, journal of wildlife diseases 29(1), 8–14. https://doi.org/10.7589/0090-3558-29.1.8 cohen, g.m., pates, a.l., easton, d.m. & peterson, m.g., 1969, ‘vulture and rooster resistance to botulinus toxin’, american zoologist 9, 584. collins, m.d. & east, a.k., 1998, ‘phylogeny and taxonomy of the food-borne pathogen clostridium botulinum and its neurotoxins’, journal of applied microbiology 84, 5–17. https://doi.org/10.1046/j.1365-2672.1997.00313.x cromie, r.l., lee, r., delahay, r.j., newth, j.l., o’brien, m.f., fairlamb, h.a. et al., 2012, ramsar wetland disease manual: guidelines for assessment, monitoring and management of animal disease in wetlands, ramsar technical report no. 7, ramsar convention secretariat, gland. eklund, m.w., poysky, f.t., petersen, m.e., peck, l.w. & brunson, w.d., 1984, ‘type e botulism in salmonids and conditions contributing to outbreaks’, aquaculture 41(4), 293–309. https://doi.org/10.1016/0044-8486(84)90198-4 espelund, m. & klaveness, d., 2014, ‘botulism outbreaks in natural environments – an update’, frontiers in microbiology 5, 1–7. https://doi.org/10.3389/fmicb.2014.00287 fijen, a.p.m. & kapp, j.f., 1995, wilderness, swartvlei and groenvlei lakes catchment, water management strategy. proposed water management strategy, objectives and goals, department of water affairs and forestry report no. wq k100/00/1395 – z/z/z, pretoria. filmalter, e. & o’keeffe, j.h., 1997, effects of land-use changes on the rivers of the wilderness lakes, report to department of environment affairs, institute for water research, rhodes university, grahamstown. getchell, r.g. & bowser, p.r., 2006, ‘ecology of type e botulism within dreissenid mussel beds’, aquatic invaders 17(2), 1–8. grubb, w.b., 1964, ‘avian botulism in western australia’, the australian journal of experimental biology and medical science 42(1), 17–26. https://doi.org/10.1038/icb.1964.2 hall, c.m., whitfield, a.k. & allanson, b.r., 1987, ‘recruitment, diversity and the influence of constrictions on the distribution of fishes in the wilderness lakes system, south africa’, south african journal of zoology 22(2), 163–169. https://doi.org/10.1080/02541858.1987.11448038 hockey, p.a.r., dean, w.r.j. & ryan, p.g. (eds.), 2005, roberts birds of southern africa, 7th edn., the trustees of the john voelcker bird book fund, cape town. howard-williams, c. & liptrot, m.r.m., 1980, ‘submerged macrophyte communities in a brackish south african estuarine-lake system’, aquatic botany 9, 101–116. https://doi.org/10.1016/0304-3770(80)90012-1 howard-williams, c. & longman, m., 1976, ‘a quantitative sampler for submerged aquatic macrophytes’, journal of the limnological society of southern africa 2(1), 31–33. https://doi.org/10.1080/03779688.1976.9632918 martinovich, d., carter, m.e., woodhouse, d.a. & mccausland, i.p., 1972, ‘an outbreak of botulism in wild waterfowl in new zealand’, new zealand veterinary journal 20(5), 61–65. https://doi.org/10.1080/00480169.1972.34012 olds, a.a., smith, k.s., weyl, o.l.f. & russell, i.a., 2011, ‘occurrence of alien invasive freshwater fishes in the wilderness lakes system, a wetland of international importance, western cape, south africa’, african zoology 46(1), 179–184. https://doi.org/10.1080/15627020.2011.11407491 perez-fuentetaja, a., clapsad, m.d., einhouse, d., bowser, p.r., getchell, r.g. & lee, w.t., 2006, ‘influence of limnological conditions on clostridium botulinum type e presence in eastern lake erie sediments (great lakes, usa)’, hydrobiologia 563(1), 189–200. https://doi.org/10.1007/s10750-005-0011-1 ramollo, p., 2016, ‘jewel of the northern cape is fading due to pollution’, the water wheel jan/feb 2016, 36–38. ramsar convention bureau, 2019, the list of wetlands of international importance, ramsar convention bureau, gland, switzerland. robarts, r.d. & allanson, b.r., 1977, ‘meromixis in the lake-like upper reaches of a south african estuary’, archiv für hydrobiologie 80(4), 531–540. rocke, t.e., 2006, ‘the global importance of avian botulism’, in g.c. boere, c.a. galbraith & d.a. stroud (eds.), waterbirds around the world, pp. 422–426, the stationery office, edinburgh. rocke, t.e. & bollinger, t.k., 2007, ‘avian botulism’, in n.j. thomas, d.b. hunter & c.t. atkinson (eds.), infectious diseases of wild birds, pp. 377–416, blackwell publishing, ames, ia. rocke, t.e. & friend, m., 1999, ‘avian botulism’, in m. friend & j.d. franson (eds.), field manual of wildlife disease: general procedures and diseases of birds, pp. 271–281, usgs national wildlife health center, washington, dc. rocke, t.e. & samuel, m.d., 1999, ‘water and sediment characteristics associated with avian botulism outbreaks in wetlands’, journal of wildlife management 63(4), 1249–1260. https://doi.org/10.2307/3802842 rocke, t.e., euliss, n.h. & samuel, m.d., 1999, ‘environmental characteristics associated with the occurrence of avian botulism in wetlands of a northern california refuge’, journal of wildlife management 63(1), 358–368. https://doi.org/10.2307/3802520 rocke, t.e., nol, p., pelliza, c. & sturm, k., 2004, ‘type c botulism in pelicans and other fish-eating birds at the salton sea’, studies in avian biology 27, 36–140. russell, i.a., 1996, ‘fish abundance in the wilderness and swartvlei lake systems: changes relative to environmental factors’, south african journal of zoology 31(1), 1–9. https://doi.org/10.1080/02541858.1996.11448389 russell, i.a., randall, r.m. & hanekom, n., 2014, ‘spatial and temporal patterns of waterbird assemblages in the wilderness lakes complex, south africa’, waterbirds 37(1), 1–18. https://doi.org/10.1675/063.037.0104 skelton, p.h., 2001, a complete guide to the freshwater fishes of southern africa, southern book publishers, halfway house. smith, g.r., 1976, ‘botulism in waterfowl’, wildfowl 27, 129. south african national parks, 2012, garden route national park: park management plan appendices, pp. 135–150, south african national parks, pretoria. stutzenbaker, c.d., brown, k. & lobpries, d., 1986, ‘special report: an assessment of the accuracy of documenting waterfowl die-offs in a texas coastal marsh’, in j.s. feierabend & a.b. russel (eds.), lead poisoning in wild waterfowl, pp. 88–95, national wildlife federation, washington, dc. taylor, j.c., harding, w.r. & archibald, c.g.m., 2007, an illustrated guide to some common diatom species from south africa, wrc report no. tt 282/07, water research commission, pretoria. thomas, r.j., 1991, ‘detection of clostridium botulinum type c and d toxin by elisa’, australian veterinary journal 68(3), 111–113. https://doi.org/10.1111/j.1751-0813.1991.tb00769.x truter, e., 1987, an aid to the identification of the dominant and commonly occurring genera of algae observed in some south african impoundments, technical report no. tr 135, south african department of water affairs, pretoria. van heerden, j., 1972, ‘botulisme breek uit in welkom’, bokmakierie 24(3), 60–61. van vuuren, s., taylor, j.c., gerber, a. & van ginkel, c., 2006, easy identification of the most common freshwater algae, north-west university and department of water affairs and forestry, pretoria. wehr, j.d. & sheath, r.g., 2003, freshwater algae of north america: ecology and classification, academic press, san diego, ca. weisser, p.j. & howard-williams, c., 1982, ‘the vegetation of the wilderness lakes system and the macrophyte encroachment problem’, bontebok 2, 19–40. whitfield, a.k., 1984, ‘the effects of prolonged aquatic macrophyte senescence on the biology of the dominant fish species in a southern african coastal lake’, estuarine, coastal and shelf science 18(3), 315–329. https://doi.org/10.1016/0272-7714(84)90074-x whitfield, a.k., 2019, fishes of southern african estuaries: from species to systems, smithiana monograph no. 4, south african institute for aquatic biodiversity, grahamstown. whitfield, a.k., allanson, b.r. & heinecken, t.j.e., 1983, estuaries of the cape, swartvlei (cms 11), report no. 22, council for industrial and scientific research, stellenbosch. włodarczyk, r., minias, p., kukier, e., grenda, t., śmietanka, k. & janiszewski, t., 2014, ‘the first case of a major avian type c botulism outbreak in poland’, avian diseases 58(3), 488–490. https://doi.org/10.1637/10669-091913-case.1 wobeser, g.a., 1997, diseases of wild waterfowl, plenum press, new york. woudstra, c., skarin, h., anniballi, f., fenicia, l., bano, l., drigo, i. et al., 2012, ‘neurotoxin gene profiling of clostridium botulinum types c and d native to different countries within europe’, applied and environmental microbiology 78(9), 3120–3127. https://doi.org/10.1128/aem.07568-11 abstract introduction methodology results discussion conclusion acknowledgements references appendix 1 about the author(s) luna milatović department of environmental sciences and policy, central european university, budapest, hungary brandon p. anthony department of environmental sciences and policy, central european university, budapest, hungary anthony swemmer south african environmental observation network (saeon), phalaborwa, south africa citation milatović, l., anthony, b.p. & swemmer, a., 2019, ‘estimating conservation effectiveness across protected areas in limpopo province, south africa’, koedoe 61(1), a1530. https://doi.org/10.4102/koedoe.v61i1.1530 original research estimating conservation effectiveness across protected areas in limpopo province, south africa luna milatović, brandon p. anthony, anthony swemmer received: 13 mar. 2018; accepted: 14 feb. 2019; published: 19 june 2019 copyright: © 2019. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract protected areas (pas) are considered the cornerstone of biodiversity conservation, and their establishment is often used to demonstrate the success of conservation policies. however, despite the increase in number and extent of pas, global trends of biodiversity loss remain largely unchanged. it has become apparent that simply establishing a pa is not enough, and effective management is equally important for achieving conservation success. the importance of management monitoring and evaluation systems has been recognised by numerous institutions, triggering the development of various conservation performance assessment frameworks and tools, each varying in their accuracy, scale and application. modified threat reduction assessment (mtra) was selected to evaluate the management effectiveness of the letaba ranch nature reserve (lrnr), which serves as a buffer area of the kruger national park (knp), south africa, and two knp sections bordering it (mahlangeni and phalaborwa). the aim of this research was to determine the nature of threats to biodiversity and their changes between 2013 and 2017 in all three sites. the research revealed 13 threats and determined negative mtra indices of −33.6% in the lrnr and −14.7% in the phalaborwa section, but a 13.2% reduction in threats in the mahlangeni section. combined with the results of geospatial visualisation of changes in threats, we suggest that the lrnr is not sufficiently meeting the objective of supporting the ecological integrity of the knp, but further exposes it to threats, and that mtras can be useful for measuring and comparing threats across pa boundaries. conservation implications: south africa has initiated a strategy on buffer zones which seeks to buffer threats to national parks. evaluating how well buffer reserves function in this capacity is crucial for understanding management effectiveness and resource allocation. utilising a combination of mtras and geospatial tools can assist managers in understanding threats and their mitigation across pas. keywords: protected areas; conservation success; management effectiveness; threat reduction assessment; biodiversity; buffer zone. introduction as we enter what many scientists call the earth’s six mass extinction (mccallum 2015; pimm et al. 2014; thomas et al. 2004), the importance of biodiversity conservation is more apparent than ever. declines in biodiversity are associated with changes that are reducing or homogenising biological diversity at many levels, from genes to habitats and ecosystems (gaston & spicer 2004). probably the most efficient and widespread tool developed in response to the biodiversity crisis is the establishment of protected areas (pas). today, there are 202 467 designated terrestrial and inland water pas, covering 14.7% of the land surface (unep-wcmc & iucn 2016). however, despite the increased number and extent of the protected area (pa) portfolio over recent decades (chape et al. 2005), biodiversity loss continues (bertzky et al. 2012) and, in some cases, even within pas (craigie et al. 2010; françoso et al. 2015; laurance et al. 2012). this has led to the growing recognition that effective management of pas is at least as important as their size, number and physical characteristics (iucn-wcpa 2009; leverington et al. 2010). there is therefore a need for management effectiveness evaluation (mee) which can assist pa management staff to assess their current management strategies, and identify shortcomings and successes, so they can modify interventions and allocate resources more efficiently (salafsky & margoluis 1999). monitoring is a crucial component of any evaluation needed for detecting changes and tracking progress towards management objectives (tucker 2005). protected area management effectiveness (pame) schemes were developed as a way of examining pa performance using management indicators (hockings & phillips 1999) and gained momentum by the obligations for convention on biological diversity signatory countries to achieve management effectiveness assessments for 60% of their pas by 2015 (cbd 2010). although there are many approaches and tools used to evaluate management effectiveness, no single pame evaluation system is suitable for all pas, and the aims and circumstances under which pame is conducted are very diverse. in fact, there are more than 50 different mee methodologies (leverington et al. 2010), most of which are modified standard approaches adapted to specific local conditions (hockings 2003). threat reduction assessment (tra), developed by salafsky and margoluis (1999), measures management effectiveness indirectly by concentrating on changes in direct biodiversity threats within a pa. threat reduction assessment is particularly useful where little or no baseline data exist on biodiversity, and when comparing this across sites, it yields a more focused and detailed approach on direct threats to biodiversity and conservation than other widely used mee tools. it is used to measure outcomes and identify effective management strategies, while overcoming the limitations of using biological indicators (salafsky & margoluis 1999). the tool was later modified (modified threat reduction assessment [mtra]) to assess changes in threats more accurately by allowing for the inclusion of worsening or emerging threats – a recognised deficiency in the original tool (anthony 2008). republic of south africa the republic of south africa (rsa) covers a total land surface area of 1 221 037 km2 (un 2017) and is listed by conservation international as one of the 17 megadiverse countries of the world, because of its highly diverse ecosystems, species richness and endemism (driver et al. 2011). however, its biodiversity is under increasing pressure because of inter alia loss and degradation of natural habitats, overexploitation of natural resources (including poaching) and invasive alien species (ias) (driver et al. 2011). the republic of south africa has a number of national policies and legislative frameworks addressing conservation and management of natural resources and biodiversity. the national environmental management: protected areas act of 2004 (nem: paa) is the main legislative document concerning the protection of the rsa’s biodiversity and conservation, while the national biodiversity framework (deat 2009) serves to align conservation efforts across institutions. another important aspect of pas in rsa is the recently legally recognised concept of buffer zones around pas, outlined in the biodiversity policy and strategy for south africa: strategy on buffer zones for national parks (notice 106 of 2012). globally, the concept originated by a growing realisation that pas cannot be managed as static, isolated islands, but are often subsets of larger ecosystems (hansen et al. 2011), and that the long-term conservation success of pas can be enhanced only when socio-ecological relationships with the surrounding landscape are considered (bengtsson et al. 2003; palomo et al. 2014). this includes a heightened appreciation of effective reserve sizes, ecological flows in and out of pas, crucial habitats as well as negative edge effects from pa peripheries (hansen & defries 2007). buffer zones gained momentum as a conservation tool in the 1970s through unesco’s man and biosphere programme (unesco 1974), which introduced the zoning hierarchy of buffer areas: the first layer of protection around the core area is the buffer zone which is surrounded by a transition area, where a greater level of land use is allowed. the principal idea is to safeguard areas that have biodiversity conservation as their primary objective (core areas) from harmful external impacts, while allowing a certain level of sustainable human activities that are consistent with the protection of the core area. this is particularly relevant for rsa’s pas, many of which are located in densely populated and impoverished parts of the country. the strategy on buffer zones for national parks, adopted in 2012, outlines a clear mission ‘to best protect the integrity of national parks, their purpose and values while enabling sustainable benefits to those persons and communities living next to the national parks’. therefore, the strategy presents not only an invaluable contribution to the protection of pas but also a tool for increased inclusion of the local community. there are a number of different forms of buffer zones, all with somewhat different objectives, including the so-called buffer parks around the kruger national park (knp). this mosaic of buffer areas around the knp, and how they contribute to ecosystem resilience, can only be better understood when both governance structures and broader socio-ecological processes are considered together in monitoring and management (cumming and allen 2017; cumming et al. 2015). although previous studies have explored large-scale, satellite-based land use transformations just south of our study area (coetzer et al. 2013), and threat mitigation by the knp and limpopo province to the north (anthony 2008), our study is the first to compare finer scale threats across adjacent pas with differing governance structures in the region. methodology study area kruger national park the kruger national park (knp) covers approximately 20 000 km2 in the north-eastern part of rsa and attracts over 1 million visitors annually. the kruger national park’s biodiversity is significant because of its size, distinctive landscapes and geological diversity (sanparks 2018). one of the key features of the knp is its protection of 150 mammal species (sanparks 2018), including many of the earth’s remaining megafauna populations (ripple et al. 2016). the park is divided into 22 sections, each managed under a common knp management plan. owing to their location, the phalaborwa and mahlangeni sections were selected for this study (figure 1). the mahlangeni section covers an area of approximately 1160 km2 of mopane-dominated woodlands on granite (gertenbach 1983), whereas the phalaborwa section is slightly smaller, covering 1035 km2 of similar vegetation (gertenbach 1983). one of the park’s entry gates (phalaborwa) is located in this section. figure 1: location of study areas. letaba ranch nature reserve the letaba ranch nature reserve (lrnr) is a provincial nature reserve located in the ba-phalaborwa and greater giyani municipalities of the mopani district in the limpopo province, covering an area of 420 km2. it is positioned along the western boundary of the knp, north of the phalaborwa gate, and serves as a buffer between the park and settlements and agricultural areas to the west. fences between the lrnr and the knp were dismantled in 2006 (swemmer & mmethi 2016), allowing the free movement of fauna between the pas. under the most recent management plan (ledet 2013), a number of management issues have been highlighted inter alia susceptibility to drought, unsustainable hunting and invasive species control. the mthimkhulu nature reserve (mnr), a community-owned nature reserve of 63.49 km2 bordering the lrnr to the north, is managed by the same provincial authority (ledet 2013) under the same management plan and is thus treated as part of the lrnr here. modified threat reduction assessment tool to evaluate management effectiveness in the selected sites, it was important to utilise a suitable pame tool. modified threat reduction assessment was chosen as it allows for comparison between sites, does not require previously collected baseline data, is relatively easy to apply, facilitates management monitoring without highly trained staff and is cost-effective (anthony 2008; salafsky & margoluis 1999). previously conducted mtras have identified group discussions as the most effective way of applying the tool, with a workshop style being particularly fitting (anthony 2008; matar & anthony 2010). the kruger national park section rangers and the lrnr managers were invited to participate and were provided with information on criteria for workshop participant selection, that is, experience working in the area during the entire assessment period; intimate knowledge of the site; and familiarity with concepts such as biodiversity, habitat conditions and ecosystem services. three workshops were conducted: two in the mahlangeni and phalaborwa sections of the knp, and one in the lrnr. each workshop commenced with a presentation introducing the methodology, key concepts and definitions, as well as a step-by-step explanation of the workshop. the mtra assessed the area of 5 km from the border inside the mahlangeni and phalaborwa sections of the knp (assumed by workshop participants to adequately account for border effects) and the whole of the lrnr. the assessment period was selected to be from 2013, the year the lrnr adopted its 5-year strategic plan (ledet 2013), until present (may 2017). the same timeframe was used for the knp to allow for comparison. utilising the iucn standard lexicon of threats (salafsky et al. 2008), participants were instructed to identify and list direct threats present at the beginning of the assessment period in their area and any emerging threats over the assessment period. after developing the final list, the top ten threats were chosen and categorised according to the iucn lexicon. participants were asked to describe the specifics of the identified threats in their area to minimise the possible loss of information resulting from threat standardisation (anthony 2008) and consulting the lexicon if required. participants were then asked to agree on detailed definitions of the identified threats and determine what a 100% reduction of that threat would be. the moderator defined a 100% threat reduction ‘as complete eradication of a given threat’, following anthony (2008), who notes that any other definition may cause ambiguity. however, if participants recognised that total threat elimination is not feasible, a different definition of 100% reduction was made. the participants were then asked to rank each threat in regard to its area, intensity and urgency. the scoring scale was defined depending on the number of identified threats, with number 1 being the minimum score without the possibility of equal scoring. participants were advised in advance to consult any reports to minimise subjectivity and increase validity of the method. criteria scores were summed to calculate a ‘total ranking’ for each individual threat. participants were then asked to review the rankings and decide if this is the true representation of the threats in their area, allowing them to modify their rankings and increase the legitimacy of the results. participants were then given time to independently estimate (as a percentage) to what degree the threat had increased or decreased over the assessment period. after group discussion, they corporately decided on the final percentage for each threat (anthony 2008). if the threat had been mitigated, they gave a positive score, with the top score being +100% if the threat had been eliminated. if a threat had worsened since the start date, the score was negative and had no lower bound. if a threat was absent at the assessment start date, but emerged since then, that threat was given a score of −100%. dividing the sum of the raw scores for each threat by the total possible rankings of all the threats and multiplying by 100 computed the threat reduction index: the second part of the workshop focused on the spatial demarcation of threats. geospatial data collection was carried out following anderson (2012). management teams were given two blank site maps and instructed to collectively draw the approximate location of threats identified during the mtra. after discussing each threat and deciding on its approximate location, threats at the beginning (2013) and end (2017) of the assessment period were drawn on separate maps to avoid overlapping and confusion during data analysis. to orientate themselves and substantiate the method, participants consulted their daily ranger patrol maps. maps produced during the mtra workshops were digitised and analysed in arcgis 10.2. each threat was constructed as a separate layer and then merged to produce final maps. in total, two maps displaying the spatial arrangement of threats to biodiversity were produced across all sites, that is, the start and end of the assessment period. all information was collated by the moderator and projected on the wall during the workshop to increase workshop coherence. at the end of each workshop, there was an informal discussion concerning the nature of the threats identified, management interventions and how successful or unsuccessful management teams had been in their mitigation. ethical considerations this research was approved by, and adhered to, the central european university, research ethics policy and guidelines. results a total of 28 participants contributed to the mtra workshops (8, 12 and 8 in mahlangeni, phalaborwa and the lrnr, respectively). the majority of participants were field rangers; however, section rangers from mahlangeni and the lrnr also participated. in total, 13 threats were identified across the three sites (table 1), with ‘hunting and collection of terrestrial animals’ and ‘invasive alien species’ present across all study sites, ‘fire and fire suppression’ and ‘utility and service lines’ were restricted to the knp only, and six threats isolated to one site only. table 1: identified threats to biodiversity and percent mitigation from 2013 to 2017 in the letaba ranch nature reserve, and mahlangeni and phalaborwa sections of the kruger national park. letaba ranch nature reserve the reserve management team identified 10 major threats to the reserve’s biodiversity (table 1-a1). the mtra index of −33.6% indicated that total threats had worsened since 2013. this was largely because of two newly emerged threats, ‘(sand) mining’ and ‘drought’, which were given a score of −100% accordingly (figure 2b). the threat of ‘problematic native species’ had doubled since 2013, which contributed heavily to the negative overall index. despite a reduction of 5%, (mostly illegal) ‘hunting’ was recognised as the top threat, followed by ‘drought’ and ‘fishing’, the latter remaining unchanged according to the management team. the threats of ‘invasive alien species’ and ‘wood harvesting’ had both halved, while pollution from ‘agricultural effluents’ from surrounding crop land and ‘garbage and solid waste’ had worsened. rangers also recognised ‘dams’ and ‘illegal fishing’ in the reserve as threats, but identified no change during the assessment period. figure 2: spatial demarcation of threats to biodiversity in assessed areas in (a) 2013 and (b) 2017. mahlangeni there was a moderate improvement of 13.2% in the mtra index for mahlangeni, as a result of the reduction of three of seven identified threats (table 2-a1; figure 2b). despite considerable reduction (+49%), ‘hunting’ still emerged as the top threat, followed by ‘fishing’ that had worsened (−50%), and ‘invasive alien species’ that had been significantly mitigated (+70%). ‘mining’ and ‘garbage and solid waste’ were both identified as worsening threats, originating from outside the park. lastly, ‘utility and service lines’ within the knp were not perceived as a serious problem, and the impact of their threat remained constant. phalaborwa management staff identified five threats (table 3-a1; figure 2a). ‘hunting’ and ‘fire’ had declined; however ‘hunting’ remained the top threat in 2017 in terms of the area affected, intensity and urgency. ‘air pollution’ from a nearby mine remained the same, as did the threat of ‘utility and service lines’. on the contrary, the problem of ‘invasive alien species’ had worsened and was given a score of −200%, which was primarily responsible for the negative overall mtra index of −14.67%. discussion knowledge gained (threats and drivers) a number of threats that were identified in more than one site (table 1) and their spatial organisation, as delineated from the threat maps, provide insights into their source(s) and reasons for change. although our study identified a total of 13 threats, we focus on five categorical threats common across at least two sites, which we believe are functionally linked. hunting and collection of terrestrial animals was recognised in all three sites as the top threat. in the lrnr, the management team expressed concern about poaching, commercial hunting activities and hunting quotas given to neighbouring communities. poaching was identified as the dominant threat in the reserve, and it was alleged that this illegal activity is usually carried out by people living adjacent to the reserve, who hunt for bushmeat as a source of food or as a source of income. most of the subsistence poaching is carried out using snares, targeting impala (aepyceros melampus), kudu (tragelaphus strepsiceros) and buffalo (syncerus caffer). a select group of poachers target elephants (loxodonta africana) and rhinos (ceratotherium simum simum) which move freely into the reserve from the knp. apart from poaching, hunting quotas given to the communities present a threat as they are purportedly not scientific estimates of sustainable levels of offtakes, an issue previously identified as a challenge in the lrnr strategic plan (ledet 2013). irregularities concerning issuing of these quotas and hunting rights have been investigated by the letaba herald (2015), a local newspaper and ems foundation (2016), which published its findings in a report prepared for cites cop 2017. both sources list individuals being involved with reselling of hunting rights in a network underpinned by corruption in the reserve. no official statements regarding the issue were made by the limpopo department of economic development, environment and tourism (ledet), and these claims are based on grey information. however, it is important to keep this in mind to understand the context in which the reserve is being managed. commercial hunting in the reserve, which was previously common, ceased in 2015, but is likely to resume according to workshop participants. the light reduction in the threat of poaching was ascribed to the increased frequency of patrols by the rangers. in mahlangeni, poaching for bushmeat, ivory, rhino horns and vulture poisoning was identified, as well as illegal harvesting of mopani caterpillars (imbrasia belina). rangers from the phalaborwa section identified rhino poaching as the dominant threat, while there was a concern that poaching for ivory would likely become the prevailing threat in the future. in addition to anti-poaching measures, the reduction of the threat in both sections can be explained by the fact that there are far fewer rhinos in the area than in 2013. according to both management teams, the number of poachers recorded in the area decreased in accordance with a decline in rhino numbers. the threat maps illustrate a potential pattern that could help understand poachers’ entrance points into the knp. in mahlangeni, areas in which poaching incidents were recorded are prevalent in the southern portion of the assessed area, which borders the northern half of the lrnr (figure 2a, b). similarly, the phalaborwa management team identified a high-intensity poaching zone in the far north of the section, which borders the southern half of the lrnr (figure 2a, b). the letaba ranch nature reserve therefore appears to function as a gateway for poachers into the knp, rather than a buffer that restricts the movement of poachers. this reflects the widely held perception of poor management practices and law enforcement within the lrnr. invasive alien species was also recognised in all three sites as one of the top four threats, with the situation improving in the lrnr and mahlangeni since 2013 but deteriorating in phalaborwa. in mahlangeni, invasive alien plants occur primarily along the letaba river which flows between the two sections. periodic flooding of this river results in frequent colonisation of exposed river banks for a variety of alien plant species. while the phalaborwa section also borders this river, rangers were more concerned with indian myna (acridotheres tristis) and large patches of alien vegetation that have emerged in terrestrial areas since 2013, contributing to the doubling of this threat for this section. rangers ascribed the −200% worsening of ias to staff shortages in the section, claiming that there are not enough people to regularly clear alien plants. in the lrnr, the extent and intensity of ias invasion are not as well researched as in the knp, but there are several known patches along the groot letaba river (figure 2a, b). in mahlangeni, rangers attributed the 70% reduction in the ias threat to successful control completed in the same manner as in the lrnr, on an ad hoc basis (ledet 2013). however, the threat map for mahlangeni did not reflect this, with only a minimal decrease in area in 2017 compared to 2013 but with continued urgency and intensity of the problem (figure 2b). the geospatial data for fishing indicated that illegal fishing in mahlangeni only takes place on the western side of the section, again suggesting that the lrnr acts as a conduit for illegal fishermen into the knp. the letaba ranch nature reserve rangers indicated no specific areas where fishing was prevalent and spatially defined the threat as present along the entire length of the two largest rivers flowing through the reserve. the discrepancy, between the mtra results and extent of fishing, indicated on the threat maps implies that rangers either overestimated the reduction in this threat or have a poorer ability of spatially describing the threat. garbage and solid waste were recognised as a threat in both the lrnr and the mahlangeni section of the knp. it was identified as a threat to wildlife, which may get entangled in waste or die from consuming it. in the lrnr, garbage is being dumped across the fence into the lrnr (figure 2a, b). the western part of the reserve is primarily (but not only) being affected, probably because the fence is more easily accessible from the road that runs along the border. rangers try to collect as much garbage as they can during patrols, but this is not a priority. in mahlangeni, solid waste is brought into the section by the klein letaba river from outside the park. during the dry season, when the management staff start eradicating ias along the river, they also collect deposited garbage. again, direct threats that originate from outside our study sites should be addressed by relevant authorities both within and outside the pas. finally, in all three sites, rangers identified fire as crucial to the health of savanna ecosystems, as long as it is managed and controlled. however, (illegal) runaway fires were identified as a threat in both sections of the knp. possible causes of these uncontrolled fires could be fires lit in the rural and urban areas, as well as along the roads to the west of the study area, which can then spread to the east because of westerly winds in the spring, or fires lit by poachers. additional management challenges the large increases for most threats found in the study suggest that management of both the buffer reserve and the studied sections of the knp should be improved. discussions with rangers and managers revealed that governance is a key problem in the lrnr, particularly in the northern mnr part. according to the strategic plan (ledet 2013), that portion of the reserve should be managed jointly with the mthimkhulu community and a co-management agreement was being drafted in 2013. however, during the mtra workshop, it was clear that the ledet management staff from the lrnr have a strong opinion that only they manage the entire area and made no mention of a co-management agreement or any sort of cooperation between the ledet and the community. interviews conducted with the community chief and mthimkhulu reserve manager suggest that the community is not benefitting from the reserve; hunting quotas are currently non-existent and they do not receive any other sort of compensation. the community also employs its own guards who supervise the north entrance to the reserve and conduct bike patrols. the fact that these stakeholders have a very different perception of the lrnr management constrains cohesion among managing institutions, increasing vulnerability to threats (as evidenced by negative mtra index scores), and threatens the long-term resilience of both the lrnr and proximate sections of the knp (cumming et al. 2015). poor governance seems to be the key reason why the lrnr does not function as an effective buffer, which has been demonstrated to inhibit conservation efforts both locally (anthony, scott & antypas 2010) and in a wider variety of contexts (lockwood 2010). in our case, questions concerning good governance principles of legitimacy, inclusiveness and connectivity emerged which necessitate further investigation now that their impacts on managing threats to biodiversity have been elucidated. it is clear that a more co-ordinated effort by diverse management institutions is needed in this buffer area which cannot be sidestepped if threat mitigation is to improve (palomo et al. 2014). this is of particular importance if the knp and the ledet would like to enhance adaptive management as a key component of their overall management strategies (gaylard & ferreira 2011). advantages and limitations of the modified threat reduction assessment our study confirmed some advantages of the tool, mentioned in earlier studies where tra was applied (anthony 2008): it is not time-consuming (no workshop lasted longer than 3 h), does not require previously collected baseline data and enables all workshop participants’ opinions to be taken into account, as the workshop moderator can facilitate the discussion and prevent individuals from dominating the discussion. in particular, the workshop setting allows for a more diversified group of participants, where rangers with different levels of experience and knowledge can contribute equally. in case of the mtra, the added advantage of incorporating worsening and emerging threats into the assessment increases the accuracy and representativeness of the results, a point workshop participants also noted. we suggest that, where appropriate, the mtra should be routinely utilised by pa management teams as a complementary mee tool, integrating the geospatial exercise that can assist management teams in synthesising and communicating threat dynamics. there are some disadvantages with the mtra method, particularly subjectivity in measurements (anthony 2008; margoluis & salafsky 2001). for instance, the discrepancy between the mtra results and the extent of fishing indicated on the threat maps implies that rangers either overestimated the reduction in this threat or have a poorer ability of spatially describing the threat. however, the high number of participants, although at times making the workshop execution more challenging, increased the objectivity and the validity of the results. potential bias further originates from the susceptibility to memory loss (papworth et al. 2009). sending information about workshops beforehand, encouraging participants to bring supporting documentation about threats and choosing a shorter assessment period (5 years) were all strategies employed to minimise this bias (as suggested in matar & anthony 2010). furthermore, when conducting tras, there is a risk of management staff overstating the threat reduction percentage in order to present their pa management as successful (margoluis & salafsky 2001), especially if being compared to other pas. nonetheless, this problem was not evident in either of the pas, as workshop participants were willing to assign high negative scores to certain threats, without considering the final tra index, and were not informed about the intention to conduct workshops in other pas. finally, the mtra tool does not explicitly reveal reasons for threat dynamics (e.g. a decrease in poaching because of decline in rhino numbers) and should be followed by a thorough discussion with management teams. capturing these nuances and the multifaceted legal and illegal drivers of these threats under the larger threat category remains an important requirement in the mtra methodology (anthony 2008) and allows management teams to devise appropriate interventions to address these threats. spatially defining threats can be an extremely useful tool to incorporate into the mtra as it allows enhanced analysis of the sources of identified threats in what cumming et al. (2015) term the ‘proximate institutional context’ which, in our case, may extend beyond the lrnr and include other buffer areas of the knp, and beyond. however, a caveat here is related to capacity. in some cases, rangers faced difficulties with spatially describing the threat suggesting either the need for more frequent training workshops or further restricting mtra workshop participants to those competent in map reading. based on our experience, in addition to the strategies addressed in matar and anthony (2010), we recommend the following: when administering the tool, it is vital to ensure all participants’ focus and engagement during the workshop. in our case, some individuals dominated the discussion and restricted full participation of other attendees. depending on context, it is important to ensure a suitable number of workshop participants (6–8) and plan extra time for workshops, allowing for ample explanations, protracted discussions and/or reaching consensus. during the workshops, to enhance data validity, it is crucial to ensure that any supporting reference material is requested beforehand, accessible and consulted. in cases where multi-level governance structures are in place across pas, a combined follow-up workshop may be useful to coordinate findings from individual sites, which can contribute to more strategic and collaborative threat mitigation efforts. conclusion through the administration of a mtra tool, a number of threats to biodiversity were explicated in the lrnr and two neighbouring sections of the knp, mahlangeni and phalaborwa. although some of these threats were reduced, the overall mtra indices of −33.6% in the lrnr and −14.67% in phalaborwa indicate that total threats have worsened since the beginning of the assessment period in these two sites. in mahlangeni, there was a reduction in threats of 13.2%. five of seven threats, spanning more than one assessment site and believed to be functionally related between the knp and the lrnr, were discussed in this article. threat mapping elucidated the probable facilitation of these threats between sites, for example, areas in which poaching incidents were recorded in the knp are proximally prevalent in the portion of the assessed area which borders the lrnr and are likely facilitated by road networks. our key finding that the lrnr is currently not serving as an effective buffer to the knp, manifested in a wide variety of threats and facilitating access to the knp, may, in large part, be because of poor governance. further research is required to delineate the drivers and nuances behind this finding, as well as explore other factors which emerged in our study and which may contribute to its relatively poor performance including lack of funding, socio-political factions and confusion over governance responsibilities (cumming et al. 2015). acknowledgements the authors would like to thank all workshop participants from the letaba ranch nature reserve and the kruger national park management teams. they also thank the journal editor and two anonymous reviewers for comments on an earlier draft of this article. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions l.m. and b.p.a. designed the research. l.m. carried out the fieldwork and l.m., b.p.a. and a.s. analysed the data and wrote the article. funding information this research was funded by the central european university, master research support program. references anderson, b., 2012, ‘assessing biodiversity threat mitigation efficacy: a case study of kakum conservation area (2004 to 2012)’, master of science thesis, central european university, budapest. anthony, b.p., 2008, ‘use of modified threat reduction assessments to estimate success of conservation measures within and adjacent to kruger national park, south africa’, conservation biology 22(6), 1497–1505. https://doi.org/10.1111/j.1523-1739.2008.01030.x anthony, b.p., scott, p. & antypas, a., 2010, ‘sitting on the fence? policies and practices in managing human-wildlife conflict in limpopo province, south africa’, conservation & society 8(3), 225–240. https://doi.org/10.4103/0972-4923.73812 bengtsson, j., angelstam, p., elmqvist, t., emanuelsson, u., folke, c., ihse, m. et al., 2003, ‘reserves, resilience and dynamic landscapes’, ambio 32(6), 389–396. https://doi.org/10.1579/0044-7447-32.6.389 bertzky, b., corrigan, c., kemsey, j., kenney, s., ravilious, c., besancon, c. et al., 2012, protected planet report 2012: tracking progress towards global targets for protected areas, international union for the conservation of nature (iucn) and united nations environment programme (unep)-world conservation monitoring centre (wcmc), gland and cambridge. chape, s., harrison, j., spalding, m. & lysenko, i., 2005, ‘measuring the extent and effectiveness of protected areas as an indicator for meeting global biodiversity targets’, philosophical transactions of the royal society a 360, 443–455. https://doi.org/10.1098/rstb.2004.1592 coetzer, k.l., erasmus, b.f.n., witkowski, e.t.f. & reyers, b., 2013, ‘the race for space: tracking land-cover transformation in a socio-ecological landscape, south africa’, environmental management 52(3), 595–611. https://doi.org/10.1007/s00267-013-0094-9 convention on biological diversity (cbd), 2010, conference of the parties (cop) 10, decision x/31. protected areas section 19(a), viewed 20 june 2017, from http://www.cbd.int/decision/cop/default.shtml?id=12297. craigie, i.d., baillie, j.e.m., balmford, a., carbone, c., collen, b., green, r.e. et al., 2010, ‘large mammal population declines in africa’s protected areas’, biological conservation 143, 2221–2228. https://doi.org/10.1016/j.biocon.2010.06.007 cumming, g.s. & allen, c.r., 2017, ‘protected areas as social-ecological systems: perspectives from resilience and social-ecological systems theory’, ecological applications 27(6), 1709–1717. https://doi.org/10.1002/eap.1584 cumming, g.s., allen, c.r., ban, n.c., biggs, d., biggs, h.c., cumming, d.h. et al., 2015, ‘understanding protected area resilience: a multi-scale, social-ecological approach’, ecological applications 25(2), 299–319. https://doi.org/10.1890/13-2113.1 department of environmental affairs and tourism (deat), 2009, national biodiversity framework, department of environmental affairs and tourism, pretoria. driver, a., sink, k.j., nel, j.n., holness, s., van niekerk, l., daniels, f. et al., 2011, national biodiversity assessment 2011: an assessment of south africa’s biodiversity and ecosystems. synthesis report, south african national biodiversity institute and department of environmental affairs, pretoria. ems foundation, 2016, enough is enough: how south africa’s policies are killing africa’s wild animals. a document prepared for cites cop 2017, ems foundation, honeydew. françoso, r.d., brandão, r., nogueira, c.c., salmona, y.b., machado, r.b. & colli, g.r., 2015, ‘habitat loss and the effectiveness of protected areas in the cerrado biodiversity hotspot’, natureza & conservação 13(1), 35–40. https://doi.org/10.1016/j.ncon.2015.04.001 gaston, k.j. & spicer, j.i., 2004, biodiversity: an introduction, 2nd edn., blackwell, oxford, uk. gaylard, a. & ferreira, s., 2011, ‘advances and challenges in the implementation of strategic adaptive management beyond the kruger national park – making linkages between science and biodiversity management’, koedoe 53(2), art. #1005, 1–8. https://doi.org/10.4102/koedoe.v53i2.1005 gertenbach, w.p.d., 1983, ‘landscapes of the kruger national park’, koedoe 26(1), 9–121. https://doi.org/10.4102/koedoe.v26i1.591 hansen, a.j., davis, c.r., piekielek, n., gross, j., theobald, d.m., goetz, s. et al., 2011, ‘delineating the ecosystems containing protected areas for monitoring and management’, bioscience 61(5), 363–373. https://doi.org/10.1525/bio.2011.61.5.5 hansen, a.j. & defries, r., 2007, ‘ecological mechanisms linking protected areas to surrounding lands’, ecological applications 17(4), 974–988. https://doi.org/10.1890/05-1098 hockings, m., 2003, ‘systems for assessing the effectiveness of management in protected areas’, bioscience 53, 823–832. https://doi.org/10.1641/0006-3568(2003)053[0823:sfateo]2.0.co;2 hockings, m. & phillips, a., 1999, ‘how well are we doing? some thoughts on the effectiveness of protected areas’, parks 9(2), 5–14. international union for the conservation of nature and natural resources (iucn) – world commission on protected areas (wcpa), 2009, wcpa strategic plan 2005–2012, iucn, gland. laurance, w.f., carolina useche, d., rendeiro, j., kalka, m., bradshaw, c.j.a., sloan, s.p. et al., 2012, ‘averting biodiversity collapse in tropical forest protected areas’, nature 489(7415), 290–294. https://doi.org/10.1038/nature11318 letaba herald, 2015, ‘letaba ranch: a nest of corruption’, letaba herald, viewed 19 july 2018, from http://letabaherald.co.za/15823/letaba-ranch-a-nest-of-corruption/. leverington, f., costa, k.l., courrau, j., pavese, h., nolte, c., marr, m. et al., 2010, management effectiveness evaluation in protected areas – a global study, 2nd edn., the university of queensland, brisbane. limpopo department of economic development, environment and tourism (ledet), 2013, five-year strategic plan for the letaba ranch nature reserve, limpopo province, south africa, limpopo department of economic development, environment and tourism, polokwane. lockwood, m., 2010, ‘good governance for terrestrial protected areas: a framework, principles and performance outcomes’, journal of environmental management 91, 754–766. https://doi.org/10.1016/j.jenvman.2009.10.005 margoluis, r. & salafsky, n., 2001, is out project succeeding? a guide to threat reduction assessment for conservation, biodiversity support program, washington d.c. matar, d. & anthony, b.p., 2010, ‘application of modified threat reduction assessments in lebanon’, conservation biology 24(5), 1174–1181. https://doi.org/10.1111/j.1523-1739.2010.01575.x mccallum, m.l., 2015, ‘vertebrate biodiversity losses point to a sixth mass extinction’, biodiversity and conservation 24(10), 2.497–2519. https://doi.org/10.1007/s10531-015-0940-6 palomo, i., montes, c., martín-lópez, b., gonzález, j.a., garcía-llorente, m., alcorlo, p. et al., 2014, ‘incorporating the social–ecological approach in protected areas in the anthropocene’, bioscience 64(3), 181–191. https://doi.org/10.1093/biosci/bit033 papworth, s.k., rist, j., coad, l. & milner-gulland, e.j., 2009, ‘evidence for shifting baseline syndrome in conservation’, conservation letters 2, 93–100. https://doi.org/10.1111/j.1755-263x.2009.00049.x pimm, s.l., jenkins, c.n., abell, r., brooks, t.m., gittleman, j.l., joppa, l.n. et al., 2014, ‘the biodiversity of species and their rates of extinction, distribution, and protection’, science 344(6187), 1246752. https://doi.org/10.1126/science.1246752 ripple, w.j., chapron, g., lópez-bao, j.v., durant, s.m., macdonald, d.w., lindsey, p.a. et al., 2016, ‘saving the world’s terrestrial megafauna’, bioscience 66, 807–812. https://doi.org/10.1093/biosci/biw092 salafsky, n. & margoluis, r., 1999, ‘threat reduction assessment: a practical and cost-effective approach to evaluating conservation and development projects’, conservation biology 13, 830–841. https://doi.org/10.1046/j.1523-1739.1999.98183.x salafsky, n., salzer, d., stattersfield, a.j., hilton-taylor, c., neugarten, r., butchart, s.h.m. et al., 2008, ‘a standard lexicon for biodiversity conservation: unified classifications of threats and actions’, conservation biology 22(4), 897–911. https://doi.org/10.1111/j.1523-1739.2008.00937.x south african national parks (sanparks), 2018, draft kruger national park management plan 2018–2028, south african national parks, skukuza. swemmer, l.k. & mmethi, a.h., 2016, biodiversity for society – a reflection on the diversity of direct, local impacts (benefits and costs) of the kruger national park, south african national parks, muckleneuk, pretoria. thomas, j.a., telfer, m.g., roy, d.b., preston, c.d., greenwood, j.j.d., asher, j. et al., 2004, ‘comparative losses of british butterflies, birds, and plants and the global extinction crisis’, science 303(5665), 1879–1881. https://doi.org/10.1126/science.1095046 tucker, g., 2005, a review of biodiversity conservation performance measures, earthwatch institute, oxford. united nations (un), 2017, un data: a world of information, viewed 03 july 2017, from http://data.un.org/countryprofile.aspx?crname=south%20africa. united nations educational, scientific and cultural organization (unesco), 1974, task force on criteria and guidelines for the choice and establishment of biosphere reserves, mab report series 22, unesco, bonn. united nations environment programme’s world conservation monitoring centre (unep-wcmc) & international union for conservation of nature (iucn), 2016, protected planet report 2016, unep-wcmc and iucn, cambridge and gland. appendix 1 table 1-a1: modified threat reduction assessment index for the letaba ranch nature reserve. table 2-a1: threat reduction assessment index for mahlangeni section of the kruger national park. table 3-a1: threat reduction assessment index for phalaborwa section of the kruger national park. abstract introduction materials and methods results discussion conclusion acknowledgements references about the author(s) michelle l. hamer zoological systematics & research collections, south african national biodiversity institute, south africa school of life sciences, university of kwazulu-natal, south africa rob slotow school of life sciences, university of kwazulu-natal, south africa school of genetics, evolution and environment, university college, united kingdom citation hamer, m.l. & slotow, r., 2017, ‘a conservation assessment of the terrestrial invertebrate fauna of mkambati nature reserve in the pondoland centre of endemism’, koedoe 59(1), a1428. https://doi.org/10.4102/koedoe.v59i1.1428 note: additional supporting information may be found in the online version of this article as online appendix 1: https://doi.org/10.4102/koedoe.v59i1.1428-1 checklist a conservation assessment of the terrestrial invertebrate fauna of mkambati nature reserve in the pondoland centre of endemism michelle l. hamer, rob slotow received: 29 july 2016; accepted: 02 feb. 2017; published: 04 may 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract mkambati nature reserve (nr) falls within the pondoland centre of endemism, which is part of the maputaland-pondoland-albany global biodiversity hotspot. the biodiversity status of this area is based largely on its flora, and the invertebrates are poorly known. the area is under threat from various proposed developments. we surveyed 14 orders in three invertebrate phyla at 26 sites with two main objectives: (1) to assess the fauna in terms of conservation value, and, (2) to identify habitats and sites of conservation concern. from the survey, 3231 samples were sent for identification and 425 species were identified. a minimum of 18 new species were confirmed. mkambati nr shows exceptional diversity for molluscs (gastropoda, 51 species), bees (apoidea, 48 species) and true bugs (heteroptera, 65 species). at least 43 species collected from the reserve are south african endemics, 31 have a restricted distribution within south africa and 18 are only known from the reserve itself. conservation implications: the authors provide the first assessment of the invertebrate fauna of the mkambati nr, which indicates that it is a rich and important fauna. the results highlight the need to consider invertebrates in other biodiversity assessments in the pondoland region. in terms of habitats, for both forest and grassland there was a large difference in the invertebrate communities at different sites, even over relatively short distances in grassland; shared habitat attributes clustered sites with more similar communities, for example, rocky ledges or the sea shore. all forest patches are a priority for protection. introduction biodiversity inventories are essential for planning, managing and monitoring activities, including in protected areas. surveys and inventories are considered by conservationists as being good investments (balmford & gaston 1999; rohr, mahan & kim 2006). they are essential to determining the level and patterns of diversity within an area (deharveng et al. 2015), including the identities and number of taxa, their distribution and the diversity of communities. invertebrates are a major component of biodiversity. they are important in all ecosystems in terms of species richness (an estimated 95% of all species are invertebrates [myers et al. 2000]), abundance and biomass, and they play vital roles in processes such as pollination, soil formation and fertility, plant productivity, organic decomposition and the regulation of populations of other organisms. invertebrates are also part of nearly every food chain, and they are a food source for many vertebrates (kellert 1993). losey and vaughan (2006) estimated that the annual financial benefit of insects through just four ecosystem services (pollination, dung burial, control of native herbivores and ecotourism) in the united states alone added up to $57 billion per annum. however, the positive role of invertebrates in ecosystems was not considered in the millennium assessment or the intergovernmental panel on climate change (prather et al. 2013), and they are generally not represented in inventories nor are they considered in local-scale conservation or management activities (mcgeoch et al. 2011). in addition to their functional roles, many invertebrate taxa have small distribution ranges, that is, they are narrow endemics (harvey 2002), and they are therefore of conservation importance. invertebrates are also recognised as being important indicators of environmental condition or change (hodkinson & jackson 2005; mcgeoch et al. 2011; paoletti, thomson & hoffmann 2007). there are a number of reasons for the lack of inclusion of invertebrates in conservation and management activities. cardoso et al. (2011) listed seven main constraints for invertebrate conservation, including a lack of awareness of the ecological role of invertebrates among the public and decision-makers, lack of basic scientific knowledge, including changes in abundance in space and time, way of life and sensitivity of species to habitat change, a large number of undescribed species and poor knowledge about the distribution of known species. the authors suggested mainly high-level and longer term solutions, but they did include provision of funds for inventories, and the use of repeatable sampling methods for surveys in their suggestions. the maputaland-pondoland-albany (mpa) is one of 34 recognised global biodiversity hotspots (mittermeier et al. 2004) and one of three that are represented in south africa. the pondoland region of the eastern cape coastal area is recognised as a centre of high plant diversity and endemism (van wyk & smith 2001) with over 200 endemic plant species. an analysis of the vertebrate fauna of the broader mpa has been carried out (perera, ratnayake-perera & proches 2011) and identified two endemic reptile species in the pondoland coastal forests. a herpetological survey of the protected areas of the wild coast part of the eastern cape, which includes pondoland, resulted in a list of 22 amphibian and 37 reptile species (venter & conradie 2015). the invertebrate fauna of pondoland are very poorly studied. existing information comprises one survey of the herbivorous insect species associated with 12 tree species in a forest patch (moran et al. 1994) and one publication on the spider fauna of mkambati nature reserve (nr) which resulted from the study presented here (dippenaar-schoeman, hamer & haddad 2011). there are published records of individual species or taxa that are in taxonomic revisions or descriptions, and there are specimens in collections, but these data are scattered through museums or in various, often difficult to access, publications. no multi-taxa, quantified survey of invertebrates has previously been carried out in pondoland. quantified surveys are critical for comparison of areas and for providing future baseline data for monitoring (lovell et al. 2010; slotow & hamer 2000). mkambati nr falls within the pondoland area and was selected for the survey of invertebrates because of proposals for expanding tourism development in the reserve, the proposed construction of a major highway adjacent to the reserve, the threat of dune mining in the region, and because of management concerns about frequent fires in some grassland areas. this study provides the first multi-taxa assessment of invertebrates of the reserve, but it will also have relevance for and application to conservation in the surrounding pondoland region. the two objectives of the study were (1) to assess the fauna in terms of diversity, level of knowledge (new species) and conservation value (endemic species and threatened species) and (2) to identify sites and habitats of high value for invertebrate conservation. we recognise that the extent of the survey may be limited in terms of sampling completeness especially considering likely temporal changes in invertebrate communities (janzen 1973; lovell et al. 2010), but the resources (capacity, time and funding) to process and identify the number of samples that would be required to achieve sampling completeness are extensive (deharveng et al. 2015; lovell et al. 2010). the results presented here should be seen in the context of a snapshot sample of the invertebrates of a threatened and neglected part of a global biodiversity hotspot and centre of endemism, and also as a data set that can contribute to decision-making for the reserve and for future monitoring programmes. materials and methods study site mkambati nr (7720 ha) is situated on the north eastern cape coast between the mtentu and msikaba rivers (31°13’ – 31°20’s/29°55’ – 30°04’e) (figure 1), and it comprises mainly open grasslands dominated by fire-prone pondoland-ugu sandstone coastal sourveld (cb 4 in mucina et al. 2006), with patches of scarp (foz5) and swamp forest (foa 2) (mucina & geldenhuys 2006; mucina et al. 2006). the climate is subtropical, with summer rainfall that averages over 1000 mm per annum. the geology is predominantly pondoland or natal sandstone, of the msikaba formation. details of geology and climate for the reserve are provided by shackleton et al. (1991). figure 1: (a) south africa showing position of mkambati nature reserve and map of mkambati nr showing rivers and boundaries and (b) map showing vegetation types and location of the forest and grassland sampling sites. mkambati was a leper colony from 1922 to 1976 (kepe 2004) with restricted access by local communities who had been removed from the area. in 1976, the leper colony was closed down, and in 1977, part of the land was declared a state protected area. since the early 1990s, access by the communities to the resources within the reserve has been permitted (kepe 2004), and after a protracted process, a land claim by the original inhabitants of the area was approved, which restored ownership to the community. there are, however, several restrictions associated with the title deeds, including that the mkambati nr must remain a protected area in perpetuity and that it may not be used for residential purposes (kepe 2008). the reserve is currently managed by eastern cape parks and tourism. sampling sites we surveyed mkambati nr between 24 january 2008 and 03 february 2008. the survey team comprised two invertebrate researchers and five research assistants, as well as seven volunteers recruited through the earthwatch institute, and a scientist from eastern cape parks and tourism. we sampled a total of 26 sites (details provided in figure 2) that included six forest patches, representing scarp and swamp forest, and 20 grassland sites, covering sites from the coast at 16 m a.s.l., those near wetlands, on rocky hills or slopes, and sites inland at 311 m a.s.l. (figures 1 and 3). additional photographs of selected sites were presented in dippenaar et al. (2011). figure 2: localities sampled at mkambati nature reserve: 24 january 2008 to 3 february 2008, with number of species and endemics sampled at each site. figure 3: selected sampling sites at mkambati nature reserve. (a) site 6, (b) site 16, (c) site f11, (d) site f15, (e) site 20 and (f) site 24. target taxa not all invertebrates were targeted for inclusion in the survey because of the enormous diversity and abundance of invertebrates, and the implications of this for capacity, time and funds required for processing and identifying material to species level. we selected invertebrates for which expertise to identify material exists, and that would represent a range of functional roles (predators, herbivores and detritivores), mobilities, body sizes and life histories. in forests, the focus was on ground-dwelling invertebrates such as gastropoda, chilopoda, diplopoda, opilionida and oligochaeta because narrow endemism of these taxa was expected to be higher in fragmented habitats (harvey 2002). other taxa (araneae, odonata, heteroptera, carabidae, cicindellidae, cetoniinae, scarabaeinae, neuroptera, lepidoptera [butterflies only], asilidae, platystomatidae, syrphidae and apoideae) were sampled in both forest and grassland habitats. sampling methods a variety of sampling methods were used because these target different microhabitats and taxa. the methods are effective and efficient and can be implemented by non-specialists, and they do not have a major impact on the environment (deharveng et al. 2015; lovell et al. 2010). grassland sampling twenty grassland sites, each covering approximately 1 ha and at least 200 m apart, were sampled. sampling at each site included setting a malaise trap at the centre of the site and five each of yellow, blue and white pan traps. these were filled with soapy water and placed on the ground 10 m apart. the traps were collected after 5 days, of which at least three were sunny. six sweep samples, each comprising 40 sweeps with a net along a 50-m strip, with each sweep approximately 1 m apart, were taken from selected sites (figure 2). the sweep samples were collected at least 50 m away from the colour pan and malaise traps. active searching for invertebrates on 40 flowers and with a net to sample flying insects for a total of 3 h per site was carried out at all 20 grassland sites (figure 2). the flower sampling attempted to cover all the obvious plant species at the site and to spread the samples across the entire site area. the active searching was only performed in the morning between 09:00 and 12:00 on days when there was sunshine and minimum wind. forest sampling six forests, ranging in area from 30 ha to 640 ha and classified as scarp (either along a river or on a sand dune) or swamp forest, were sampled. we sampled two to four (depending on forest size) 20 m × 20 m plots for 1 h each, with three people searching for 20 min in each plot, and thoroughly searched a 10 m × 2 m quadrat divided into five contiguous 2 m × 2 m blocks, with one person searching each block. four leaf litter samples, each comprising a 2-l bag of litter and the surface layer of soil, were collected from each forest to sample micromolluscs. ten trees were each beaten 10 times by hitting a branch with a large, robust truncheon to sample selected invertebrate taxa. a cloth or beating tray was placed under the branch being beaten, and target invertebrates were collected from the cloth or tray using an aspirator or vial. six fruit-baited butterfly traps were hung from trees in each of the forests and left for 2–5 days to collect butterflies, flies and fruit chafer beetles that were attracted by the rotting fruit bait. the traps could not be left for the same period of time in all of the forests because a limited number of traps were available for the study. this may have impacted the results. additional specimens were collected opportunistically outside the sampling sites. these 11 species are included in the checklist (online appendix 1), and in the species and specimen totals, but were excluded from the analyses. sample processing and identification for active sampling methods, where accurate identification in the field was possible, a maximum of five individuals were collected, and additional specimens were recorded and released. this was especially important in the case of long-lived taxa such as millipedes and large molluscs. for collected material, each sample, or specimen where these were separate, was labelled with a unique numeric code in the field. in the laboratory, mixed samples such as malaise traps, pan traps and sweep samples were sorted to separate out each of the target taxa. each target taxon was then separated into morphospecies, which were sent to experts for identification. where identification lower than family was not possible, these records were removed from the data. in cases where the genus could be identified and species could be separated by the taxonomist but not named (morphospecies), we included these records in the data set. specimens from the survey have been deposited into the agricultural research council (arc) national collections, the kwazulu-natal museum and the ditsong national museum of natural history in pretoria. for micromolluscs, the leaf litter samples were air dried for approximately 1 week and sifted through a series of four sieves of decreasing mesh sizes. the four fragments from each sample were then searched using a dissecting microscope, and micromolluscs were removed by hand. data analysis sampling completeness was determined using estimates (version 9.1.0), copyright r. k. colwell: http://purl.oclc.org/estimates, and has been plotted separately for forests and grassland. level of endemism could only be determined where comprehensive distribution data sets are available (e.g. the south african national survey of arachnida or arc spider database), or where there are checklists (e.g. hamer [1999] for millipedes), conservation assessments such as that for butterflies (mecenero et al. 2013), or field guides that give species’ distributions (tarboton & tarboton [2002, 2005] for odonata, herbert & kilburn [2004] for terrestrial molluscs). the following categories of endemism were slightly modified from those described by hamer and slotow (2002): (1) national endemic = confined to south africa, (2) regional endemic = confined to an area where the two furthest locality points are between 60 km and 330 km apart, (3) local endemic – multiple localities known = more than one locality known and confined to an area where the two furthest points are less than 60 km apart and (4) known only from mkambati nr. we assessed differences in communities across sites and related these to the vegetation characteristics of the site. we analysed the grassland and forest sites separately, and to assess species assemblages sampled at each site, we constructed bray–curtis similarity matrices (presence–absence), followed by analysis of similarity (anosim) and non-metric multi-dimensional scaling (mds) plots using primer version 5.2.9 (clarke & warwick 2001). the bray–curtis calculated on presence–absence data in primer provides the same outcome as jaccard and sørensen, which are the two commonly used measures of similarity for this type of data (clarke & gorley 2001). from anosim, if r > 0.75, groups are well separated; if r > 0.5, groups are over-lapping but clearly different; if r < 0.25, groups are barely separable (clarke & gorley 2001). only those grassland sites for which the full suite of sampling methods had been used (figure 2) were included in these analyses, that is, effort was equal across sites. results assessment of diversity or species richness a total of 2531 samples were sent for expert identification. of these, 71 could not be identified to any useful level for various reasons. for some taxa, notably the millipedes (diplopoda) and earthworms (oligochaeta), a large proportion of the specimens were juveniles that could not be identified even to genus level. for other taxa such as the true bugs (heteroptera), a large number of specimens could not be identified to species level because the expertise or taxonomic information does not currently exist. a total of 425 species were identified from the target taxa during the survey, with 211 from grassland, 155 from forest, 46 that were sampled in both biomes and 11 species without habitat details from opportunistic collecting. sampling was not complete (figure 4). the chao 2 values calculated were 270.11 ± 35.06 sd species for forests and 373.64 ± 39.8 sd species for grassland, which suggests that approximately 74% of forest and 69% of grassland species present were actually sampled. one exotic earthworm was recorded from the forest biome. an annotated checklist of taxa sampled is presented as online appendix 1. this list excludes the spiders, which were published separately in dippenaar et al. (2011). figure 4: species accumulation curves for (a) forest and (b) grassland sites at mkambati nature reserve, calculated based on samples and using s(est), the expected number of species in pooled samples given the reference sample (189 = 163 species for forest, and 291 = 228 species for grassland), and showing the upper and lower 95% confidence limits (dashed lines). the number of new species identified is a minimum of 18, with the possibility of this number being as high as 40 species (table 1). the uncertainty in the figure is based on the need for dna analysis in some cases, or the need for taxonomic revision or more specialist expertise to verify suspected new species. table 1: summary of number of specimens collected, identified, number of species and number of new species collected for each of the target taxa at mkambati nature reserve. mkambati nr shows exceptional richness for some taxa, such as the molluscs (gastropoda) (51 species), bees (apoidea) (48 species) and true bugs (hemiptera) (65 species). the bee and true bug richness is likely to be higher if sampling was repeated at different times of the year, but the molluscs do not show much temporal turnover (uys, hamer & slotow 2010), and the richness value for molluscs is unlikely to increase much with additional sampling. the total number of endemics, at 104 species (25% of the total number of species identified) (table 2) that are endemic to south africa or a smaller area within the country, is likely to be an underestimate because for a large proportion of species the level of endemism could not be assessed because of a lack of accessible distribution data. forty-three species (10% of the total) are restricted to the pondoland area and 18 species (4% of the total) are only known from mkambati nr. the molluscs have a remarkably high level of endemism, with 43 species (68%) qualifying as endemic within south africa, and 47% of these being restricted to the pondoland region. table 2: number of species in different categories of endemism for invertebrate taxa surveyed at mkambati nature reserve. few invertebrate taxa have been formally assessed according to the international union for conservation of nature (iucn) red list criteria (iucn 2001). the red list currently includes south african butterflies and odonata, but none of the species sampled at mkambati nr are considered as threatened. one mollusc species on the current red list occurs in mkambati nr but was not collected during this survey. natalina beyrichi is categorised as vulnerable because it has only been collected from dwessa and mkambati nrs. the exact locality within mkambati nr for this species is not known. habitats and sites of special concern species presence or absence data for different grassland habitats showed that different habitats were all more than 40% different (figure 5a). however, the two sites that were closest to the seashore had similar invertebrate communities (40% similar), which were distinct from other sites, whereas the rocky ledges also grouped (± 50% similar), as did the thick, unburnt grassland sites (figure 5a and b). distance may also be influencing the community structure because sites 16–19 were similar (50%) (figure 5a and b) and were also close in terms of distance between them (figure 1). these sites were also known to burn at least annually as a result of fires spreading from the adjacent communal areas so fire frequency may also influence invertebrate diversity. sites 17 and 18 also had the highest species richness (54 and 55 species) (figure 2). site 24 had thick, unburnt grassland with few flowers, and sloped down to a forest patch, and this site was distinct from the other thick, unburned sites (sites 2, 7, 8) that could be explained by its higher altitude (figure 2). figure 5: community relationships among grassland sites at mkambati based on those sites at which all sampling methods were used. (a) similarity analysis and (b) multi-dimensional scaling analysis. species presence or absence data for the different forest types sampled showed that none of the forests were more than 50% similar if data for all target taxa were used. however, the swamp forest and the swamp or scarp forest had some similarities (40% similar), as did the two scarp forests associated with rivers (50% similar) (figure 6a). in the mds plot, the two swamp forests (sites f11 and f15) are separated from the other forests (figure 6b). forest type appeared to be a stronger influence on community structure than distance, with the closest forests not showing the most similar invertebrate community structure. figure 6: community relationships among forest sites at mkambati for all taxa. (a) similarity analysis and (b) multi-dimensional scaling analysis. the forest molluscs were explored further because of the large diversity and the low mobility of this taxon and because it is likely that a high proportion of species present were sampled. analyses for the molluscs showed similar patterns to those for all taxa, with low levels of similarity in community structure across sites (figure 7a and b). one swamp forest (f15) was most different from the scarp forests (< 40% similarity), but this is related to the low number of species (8). the scarp forest on the dunes (f12) was also distinct (± 45% similarity), and this was related to the high number of species (28), including seven that were unique to this forest. figure 7: community relationships among forest sites at mkambati for mollusca, gastropoda. (a) similarity analysis and (b) multi-dimensional scaling analysis. discussion information about species’ distributions is critical for spatial planning and protected area expansion (polak et al. 2015), and an understanding of the habitat associations and local distribution patterns of species is essential for local-scale planning and monitoring (corona et al. 2011). this preliminary survey provides new data, which can contribute to understanding the conservation value of mkambati nr, for planning, development and monitoring within the reserve. it must be recognised, however, that the survey represents a limited data set in terms of temporal variation and sampling effort, and the estimated species richness is higher than that sampled. increasing the number of samples collected at each site, as well as the number of sites sampled (ashcroft et al. 2010), and resampling to capture temporal change would increase the number of species. increasing sampling effort in invertebrate surveys increases the time and cost of processing and identifying the material (ward & larivière 2004), and these limitations have to be weighed up against the urgent need for data for areas such as those in global biodiversity hotspots. this survey did highlight the high richness, poor knowledge and uniqueness of the invertebrate fauna, and it provides a start in identifying some trends relating to habitats. the mpa hotspot and the pondoland centre of endemism are recognised as having high plant species richness and levels of endemism. although it is difficult to compare the invertebrate richness of mkambati nr with that for other localities because of differences in extent of the area surveyed, sampling methods and intensity and taxa included, two other multi-taxa invertebrate surveys within the mpa hotspot do provide at least an indication of the relative richness of the mkambati nr fauna. a 3-year survey of mkhuze and phinda game reserves in maputaland, about 410 km2 in extent, covered 77 sites, most of which were sampled more than once in different seasons or years. a total of 49 961 specimens were identified as 716 invertebrate species (lovell et al. 2010). a survey of 28 forests and 55 grassland sites along the length of the 14 740 km2 maloti-drakensberg mountains produced 10 168 specimens, which yielded 832 species (hamer & slotow 2007). in comparison with these extensive surveys, the 425 species identified from the relatively small 77 km2 area of mkambati nr suggest exceptional richness. at the level of individual taxa, the number of species sampled at mkambati nr was also high relative to the maloti-drakensberg mountains for some taxa, despite the smaller area and limited sampling effort in the former area. for example, 66 bee species were sampled in the maloti-drakensberg survey, 51 species at mkuze-phinda and 48 species at mkambati nr; 48 molluscs were sampled in the maloti-drakensberg, 50 in mkhuze-phinda and 51 species in mkambati nr. for the other taxa, the richness was not notably high relative to other areas. for example, the spider richness at mkambati nr was 132 species (dippenaar-schoeman et al. 2011), which is moderate relative to surveys of other protected areas. haddad and dippenaar-schoeman (2009) provided a range of between 76 and 431 species for south african conserved areas for which published surveys exist but none of the reserves they provided data for were in the forest or grassland biome. millipede and earthworm richness were low relative to mkhuze-phinda and the maloti-drakensberg mountains, but for both taxa a large number of juvenile specimens were collected and these cannot be identified to species level. the high overall species richness in mkambati nr is likely to be related to the moderate, stable coastal temperatures relative to those in the high altitude drakensberg mountains. the biogeography of the fauna needs to be analysed, especially in the context of the flora, which has both palaeoendemics and neoendemics with strong drakensberg and fynbos associations (mucina et al. 2006; van wyk & smith 2001), but this is beyond the scope of this article. in addition to species richness, the level of endemism in mkambati nr is high, with approximately 25% of species (104) sampled endemic to south africa. at least 18 of these species are known only from mkambati nr, and another 12 are local endemics (7% of the total). these percentages are likely to be underestimates because where there was uncertainty about the distribution or identity of a species, this was not included in the count of endemics. comparison of the values with those for other areas in south africa is difficult because few published studies have carried out this type of assessment and calculation. the cape peninsula, a locality known to have habitats associated with high levels of endemism, was found to have 32% of invertebrates endemic to that area (picker & samways 1996). the percentage of endemics varied according to taxon with opilionida having the highest level at 67% endemic species and the butterflies having the lowest level at 5%. at mkambati nr, the molluscs had the highest level of endemism, with 15 species (29%) that are local endemics or only known from mkambati nr. across all taxa, most of the local endemics and those species known only from mkambati nr are in forests. all forests in mkambati have high conservation value, because all of them, except site f11, produced local endemics or species only known from mkambati nr that were unique to that forest. even site f11 had five species that are regional or local endemics, although these were not unique to the forest, and 18 species that were not collected in any of the other forests, although these species are not narrow endemics. van wyk and smith (2001) suggested that swamp forests are small, rare and are impoverished in terms of species and endemics but this does not seem to reflect the situation for invertebrates at mkambati nr. the mkambati nr forests illustrate that small forest patches have value for invertebrate conservation; they do not just hold subsets of the species that are present in large forests in the area. for the grassland biome, rocky ledges should be protected because of their rich and distinct invertebrate fauna, and rocky outcrops were also identified by perera et al. (2011) as being important for vertebrate endemics. the seashore sites were recognised as being floristically unique by shackleton et al. (1991), and they are also distinct in terms of their invertebrate community. these habitats are threatened because they cover a limited area and are prime sites for the development of tourist accommodation and facilities. within the grassland, distance appears to have some effect on community structure, which means that sites prioritised for conservation should cover both altitudinal and latitudinal gradients. the three richest grassland sites were those that had burnt in spring or early summer and that are known to burn at least annually. it is, however, not possible to draw any conclusions about fire impacts on invertebrates or make any recommendations on fire management based on the limited data. the large number of gaps in knowledge and the large number of new species collected highlight the need to carry out additional research in the pondoland region. additional surveys at different times of the year are important, not only for sampling a different suite of insects, but also for collecting adult males of millipedes and mature specimens of earthworms and spiders. there are many forest patches that remain to be sampled, and given the high levels of uniqueness of the forests that were surveyed, this is likely to increase the species list for the region and the number of new species. the data collected through this survey, but also through future work, will be useful for planning sustainable development within mkambati nr, and the trends identified here (high richness and endemism, large differences in communities across forest types and the differences in community even in seemingly similar or homogenous habitats) should also be considered in biodiversity assessments outside the reserve. we therefore recommend (1) additional surveys, (2) that invertebrate conservation be considered in management and development plans for mkambati nr and (3) that the results from this study should be considered in the pondoland area until more robust data for those area have been collected. conclusion the main objectives of this study were to assess the terrestrial invertebrate fauna in terms of conservation value and to identify habitats and sites of conservation concern in mkambati nr which falls within the pondoland centre of endemism. a range of sampling methods were used to collect various taxa across 26 sites covering different forest types and grassland habitats. although far from complete, this survey indicates high species richness especially for bees, true bugs and terrestrial snails, and high levels of endemism. all forests within mkambati nr have conservation value because of their uniqueness and high number of endemic species. rocky ridges and the seashore habitats in the grassland had the highest richness and uniqueness, and distance may also influence the similarity of the invertebrate communities in grassland. the results indicate the need for further invertebrate surveys both in mkambati nr and more widely in pondoland, and highlight the need to include invertebrates in planning and management of the reserve. acknowledgements the authors are very grateful to the following experts for identifying the material collected: oligochaeta: dr danuta plisko (kwazulu-natal museum), molluscs: prof. dai herbert (kwazulu-natal museum), araneae: prof. ansie dippenaar-schoeman (arc), opilionida: mr. leon lotz (national museum), scorpionida: dr ian engelbrecht (private), neuroptera: dr mervyn mansell (private), odonata: dr tanza crouch (private), butterflies: mr kevin cockburn (lepidopterist society of south africa), cetoninae: prof. renzo perissonotto (nelson mandela metropolitan university), cicindellinae and carabidae: dr peter schule (private), scarabaeinae: dr adrian davis (university of pretoria), asilidae: dr jason londt (kwazulu-natal museum), syrphidae and platystomatidae: dr andrew whittington (private), heteroptera: dr dawid jacobs (university of pretoria), cicadidae: prof. martin villet (rhodes university), and apoidea: dr connal eardley (arc). diplopoda and chilopoda were identified by the first author. the authors thank eastern cape parks board for providing funding and logistical support to carry out this survey. jan venter, dave balfour and lwazi khuzwayo, in particular are thanked for their assistance. the authors thank the earthwatch volunteers who assisted in the field, as well as university of kwazulu-natal (ukzn) technicians warren matten, lynne stone and jessica cockburn. ukzn provided access to a vehicle, and laboratory facilities. two anonymous experts are thanked for their useful comments on the manuscript. earthwatch institute provided funding through the support of british american tobacco. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions m.l.h. led the fieldwork, sorting and processing of samples and wrote the manuscript; r.s. assisted with the design of the study and the field sampling, carried out the community analyses and provided input on the manuscript. references ashcroft, m.b., gollan, j.r., faith, d.p., carter, g.a., lassau, s.a., ginn, s.g. et al., 2010, ‘using generalised dissimilarity models and many small samples to improve the efficiency of regional and landscape scale invertebrate sampling’, ecological informatics 5(2), 124–132. https://doi.org/10.1016/j.ecoinf.2009.12.002 balmford, a. & gaston, k., 1999, ‘why biodiversity surveys are good value’, nature 398, 204–205. https://doi.org/10.1038/18339 cardoso, p., erwin, t.l., borges, p.a. & new, t.r., 2011, ‘the seven impediments in invertebrate conservation and how to overcome them’, biological conservation 144(11), 2647–2655. https://doi.org/10.1016/j.biocon.2011.07.024 clarke, k.r. & gorley, r.n., 2001, primer v5, primer-e ltd, plymouth, uk. clarke, k.r. & warwick, r.m., 2001, primer v5: user manual/tutorial, primer-e ltd, plymouth, uk. corona, p., chirici, g., mcroberts, r.e., winter, s. & barbati, a., 2011, ‘contribution of large-scale forest inventories to biodiversity assessment and monitoring’, forest ecology and management 262(11), 2061–2069. https://doi.org/10.1016/j.foreco.2011.08.044 deharveng, l., bedos, a., daugeron, c., villemant, c. & judson, m.l., 2015, ‘organization, usefulness and limitations of an atbi (all taxa biodiversity inventory): the inventory of terrestrial invertebrates in the mercantour national park’, zoosystema 37(1), 9–30. https://doi.org/10.5252/z2015n1a1 dippenaar-schoeman, a.s., hamer, m. & haddad, c.r., 2011, ‘spiders (arachnida: araneae) of the vegetation layer of the mkambati nature reserve, eastern cape, south africa’, koedoe 53(1), art. #1058, 1–10. https://doi.org/10.4102/koedoe.v53i1.1058 haddad, c.r. & dippenaar-schoeman, a.s., 2009, ‘a checklist of the non-acarine arachnids (chelicerata: arachnida) of the de hoop nature reserve, western cape province, south africa’, koedoe 51, 1–9. hamer, m.l., 1999, ‘an annotated and illustrated key to the southern african genera of spirostreptida (diplopoda)’, annals of the natal museum 40, 1–22. hamer, m.l. & slotow, r., 2002, ‘conservation application of existing data for south african millipedes’, african entomology 10(1), 29–42. hamer, m.l. & slotow, r., 2007, ‘an assessment and ecological interpretation of the terrestrial invertebrate fauna of the south african portion of the maloti-drakensberg bioregion’, unpublished report to the maloti drakensberg transfrontier project. harvey, m.s., 2002, ‘short-range endemism amongst the australian fauna: some examples from non-marine environments’, invertebrate systematics 16(4), 555–570. https://doi.org/10.1071/is02009 herbert, d. & kilburn, r., 2004, field guide to the land snails and slugs of eastern south africa, natal museum, pietermaritzburg. hodkinson, i.d. & jackson, j.k., 2005, ‘terrestrial and aquatic invertebrates as bioindicators for environmental monitoring, with particular reference to mountain ecosystems’, environmental management 35(5), 649–666. https://doi.org/10.1007/s00267-004-0211-x iucn, 2001, 2001 categories and criteria, iucn, gland, switzerland, viewed 17 july 2016, from http://www.iucnredlist.org/info/categories_criteria2001 janzen, d.h., 1973, ‘sweep samples of tropical foliage insects: effects of seasons, vegetation types, elevation, time of day, and insularity’, ecology 54(3), 687–708. https://doi.org/10.2307/1935359 kellert, s.r., 1993, ‘values and perceptions of invertebrates’, conservation biology 7(4), 845–855. https://doi.org/10.1046/j.1523-1739.1993.740845.x kepe, t., 2004, ‘land restitution and biodiversity conservation in south africa: the case of mkambati, eastern cape province’, canadian journal of african studies/la revue canadienne des études africaines 38(3), 688–704. https://doi.org/10.2307/4107262 kepe, t., 2008, ‘land claims and co-management of protected areas in south africa: exploring the challenges’, environmental management 41(3), 311–321. https://doi.org/10.1007/s00267-007-9034-x losey, j.e. & vaughan, m., 2006, ‘the economic value of the ecosystem services provided by insects’, bioscience 56(4), 311–323. https://doi.org/10.1641/0006-3568(2006)56[311:tevoes]2.0.co;2 lovell, s.j., hamer, m.l., slotow, r.h. & herbert, d., 2010, ‘assessment of sampling approaches for a multi-taxa invertebrate survey in a south african savanna-mosaic ecosystem’, austral ecology 35, 357–370. https://doi.org/10.1111/j.1442-9993.2009.02052.x mcgeoch, m.a., sithole, h., samways, m.j., simaika, j.p., pryke, j.s., picker, m. et al., 2011, ‘conservation and monitoring of invertebrates in terrestrial protected areas’, koedoe 53(2), art. #1000, 1–13. https://doi.org/10.4102/koedoe.v53i2.1000 mecenero, s., ball, j.b., edge, d.a., hamer, m.l., henning, g.a., kruger, m. et al. (eds.), 2013, conservation assessment of butterflies of south africa, lesotho and swaziland: red list and atlas, saftronics (pty) ltd, johannesburg & the animal demography unit, cape town. mittermeier, r.a., robles-gil, p., hoffmann, m., pilgrim, j.d., brooks, t.b., mittermeier, c.g. et al. (eds.), 2004, hotspots revisited: earth’s biologically richest and most endangered terrestrial ecoregions, cemex, mexico, mi. moran, v.c., hoffmann, j.h., impson, f.a.c. & jenkins, j.f.g., 1994, ‘herbivorous insect species in the tree canopy of a relict south african forest’, ecological entomology 19(2), 147–154. https://doi.org/10.1111/j.1365-2311.1994.tb00404.x mucina, l. & geldenhuys, c.j., 2006, ‘afrotemperate, subtropical and azonal forests’, in l. mucina & m.c. rutherford (eds.), the vegetation of south africa, lesotho and swaziland, pp. 584–614, south african national biodiversity institute, pretoria. mucina, l., scott-shaw, c.r., rutherford, m.c., camp, k.g.t., matthews, w.s., powrie, l.w. et al., 2006, ‘indian ocean coastal belt’, in l. mucina & m.c. rutherford (eds.), the vegetation of south africa, lesotho and swaziland, pp. 568–583, south african national biodiversity institute, pretoria. myers, n.a., mittermeier r.a., mittermeier, c.g., da fonseca, g.a.b. & kent, j., 2000, ‘biodiversity hotspots for conservation priorities’, nature 403, 853–858. https://doi.org/10.1038/35002501 paoletti, m.g., thomson, l.j. & hoffmann, a.a., 2007, ‘using invertebrate bioindicators to assess agricultural sustainability in australia: proposals and current practices’, animal production science 47(4), 379–383. https://doi.org/10.1071/ea05288 perera, s.j., ratnayake-perera, d. & proches, s., 2011, ‘vertebrate distributions indicate a greater maputaland-pondoland-albany region of endemism’, south african journal of science 107(7–8), 52–66. https://doi.org/10.4102/sajs.v107i7/8.462 picker, m.d. & samways, m.j., 1996, ‘faunal diversity and endemicity of the cape peninsula, south africa – a first assessment’, biodiversity & conservation 5(5), 591–606. https://doi.org/10.1007/bf00137611 polak, t., watson, j.e.m., fuller, r.a., joseph, l.n., martin, t.g., possingham, h.p. et al., 2015, ‘efficient expansion of global protected areas requires simultaneous planning for species and ecosystems’, royal society open science 2, 150107. https://doi.org/10.1098/rsos.150107 prather, c.m., pelini, s.l., laws, a., rivest, e., woltz, m., bloch, c.p. et al., 2013, ‘invertebrates, ecosystem services and climate change’, biological reviews 88(2), 327–348. https://doi.org/10.1111/brv.12002 rohr, j.r., mahan, c.g. & kim, k.c., 2006, ‘developing a monitoring program for invertebrates: guidelines and a case study’, conservation biology 21, 422–433. https://doi.org/10.1111/j.1523-1739.2006.00578.x shackleton, c.m., granger, j.e., mckenzie, b. & mentis, m.t., 1991, ‘multivariate analysis of coastal grasslands at mkambati game reserve, north-eastern pondoland, transkei’, bothalia 21(1), 91–107. https://doi.org/10.4102/abc.v21i1.869 slotow, r. & hamer, m., 2000, ‘biodiversity research in south africa: comments on current trends and methods’, south african journal of science 96(5), 222–225. tarboton, w. & tarboton, m., 2002, a fieldguide to the dragonflies of south africa, tarboton & tarboton, modimolle. tarboton, w. & tarboton, m., 2005, a fieldguide to the damselflies of south africa, tarboton & tarboton, modimolle. uys, c., hamer, m. & slotow, r., 2010, ‘step process for selecting and testing surrogates and indicators of afrotemperate forest invertebrate diversity’, plos one 5(2), e9100. https://doi.org/10.1371/journal.pone.0009100 van wyk, a.e. & smith, g.f., 2001, regions of floristic endemism in southern africa. a review with emphasis on succulents, umdaus press, pretoria. venter, j.a. & conradie, w., 2015, ‘a checklist of the reptiles and amphibians found in protected areas along the south african wild coast, with notes on conservation implications’, koedoe 57(1), art. #1247, 1–25. https://doi.org/10.4102/koedoe.v57i1.1247 ward, d.f. & larivière, m.c., 2004, ‘terrestrial invertebrate surveys and rapid biodiversity assessment in new zealand: lessons from australia’, new zealand journal of ecology 28, 151–159. filelist convert a pdf file! page 1 page 2 page 3 page 4 abstract introduction research outputs from kruger national park research supersites (up to august 2019) effectiveness of kruger national park research supersites in achieving original objectives discussion, recommendations and conclusions acknowledgements references footnotes about the author(s) izak p.j. smit scientific services, south african national parks, skukuza, south africa centre for african ecology, school of animal, plant and environmental sciences, university of the witwatersrand, johannesburg, south africa citation smit, i.p.j., 2020, ‘integrating multi-scaled and multidisciplinary studies: a critical reflection on the kruger national park research supersites’, koedoe 62(2), a1586. https://doi.org/10.4102/koedoe.v62i2.1586 note: additional supporting information may be found in the online version of this article. online appendix 1; online appendix 2; and online appendix 3. note: special issue: connections between abiotic and biotic components of a granite catena ecosystem in kruger national park, sub-edited by beanelri janecke and johan van tol. essay integrating multi-scaled and multidisciplinary studies: a critical reflection on the kruger national park research supersites izak p.j. smit received: 12 sept. 2019; accepted: 08 jan. 2020; published: 29 oct. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the kruger national park (knp) research supersites were designed to encourage place-based research in order to geographically focus research activities on known and well described study sites as opposed to ad hoc site selection practiced previously. this was done by (i) delineating sites using a clear rationale, (ii) providing basic meta-data for these sites, and (iii) actively encouraging scientists to conduct research on these sites and share data freely. the underlying concept was that geographically focused research would facilitate data and knowledge exchanges and lead to long-term, multi-scaled and cross-disciplinary studies at these data-rich sites, facilitating an integrated and collectively developed understanding that would be hard to achieve otherwise. this essay acts as a short-term reflection on the knp supersites and an introductory text for the special issue focusing on the outcomes from a multi-disciplinary study conducted on the southern granitic supersite. it starts off by briefly introducing the supersite concept, followed by a reflection on the achievements and challenges towards achieving the main objectives of the supersites. in addition, and as part of the “data-begets-data” philosophy underlying the supersites (i.e positive feedback of place-based data attracting further research and hence collection of further data), updated lists of references and available datasets are provided. conservation implications: this paper highlights the successes and challenges of geographically focusing research in the knp to the research supersites in order to facilitate integrative and multi-scaled learning in savanna systems. it also provides updated lists of references and available datasets to further stimulate research at these sites. keywords: long-term ecological research sites (lters); monitoring; multidisciplinary studies; integrative studies; research supersites. introduction over the years, the kruger national park (knp) has established a reputation as an ‘outdoor laboratory’, and has registered more than 700 research projects between 2005 and 2016, and published 556 papers between 2003 and 2013, both by in-house scientists and external collaborators from around the world (smit et al. 2017; van wilgen et al. 2016). as a result, the park has become one of the most studied savanna conservation areas in africa (smit et al. 2017). many of these studies have focussed on specific experimental sites where underlying drivers are manipulated, for example, herbivore exclosures (e.g. asner et al. 2009) or plots where fire regimes have been manipulated (e.g. higgins et al. 2007). however, for many projects that aim to study or monitor patterns and processes emerging under non-manipulated conditions, sites were selected in a haphazard and uncoordinated manner and, as a consequence, because of underlying heterogeneity, it was often hard to integrate data sets and knowledge across these disparate sites. in an attempt to geographically focus research effort and allow data integration over time and across themes, the ‘knp research supersites’ was conceptualised. it was envisaged that through establishing these areas, some geographic focussing of research would be achieved with these sites increasingly acting as data-rich, long-term sites for monitoring and research. in many respects, the knp supersites have objectives similar to long-term ecological research sites (lters) (e.g. gosz, waide & magnuson 2010; mirtl et al. 2018). the four knp research supersites that cover the rainfall gradient (lower rainfall northern knp and higher rainfall southern knp) and geological contrast (basalt in east and granite in west) were formally introduced to the scientific community by smit et al. (2013) who described the rationale, selection criteria and location of the sites, and introduced existing data sets describing environmental variables for each of these sites. the supersites’ conceptualisation and delineation process started earlier (around 2011). six years after formally introducing the supersites and coinciding with this special issue focussing on a multidisciplinary project on the southern granitic supersite (stevenson-hamilton supersite), it is an opportune time to reflect on the successes of the knp research supersites, the shortcomings and possible ways to further increase the value of these sites going forward. it is hoped that this reflection will also be useful for other situations where long-term research sites are being established. this essay will also act as a valuable reference document by providing consolidated and updated lists of research outputs and data sets (online appendix 1 – online appendix 3). research outputs from kruger national park research supersites (up to august 2019) since establishment, four honours, four master’s and two phds have been completed on the supersites (online appendix 1). in addition, 24 peer reviewed articles (11 as part of this special issue, excluding this essay); two scientific reports; one book chapter; and one peer-reviewed conference proceeding have been published based on work fully or mostly conducted on the supersites (online appendix 1). considering the relatively short existence of the supersites, the research outputs compare favourably to the number of research outputs associated with the knp’s well-established experimental burn plots (ebps) that have been in existence since the 1950s (by june 2020, there were 79 papers published on the ebps tercia strydom [south african national parks] pers. comm., 19 june 2020). the skukuza flux tower is another example of place-based research in the knp, linked to specialised equipment, which has attracted research attention and has been used as a study site for a range of studies (e.g. majozi et al. 2017). it is also expected that as the supersites become better known and more data were accumulated, research at these sites will be further stimulated. it is anticipated that this special issue, which reports on integrative learning happening at the southern granitic supersite across a range of disciplines, will also advance the profile and increase the understanding of these sites, and will act as a further catalyst for stimulating research interest. online appendix 2 provides a list of a diverse range of papers and dissertations/theses known to the author that cite the supersites or make a reference to the knp supersites concept, reflecting also on the broader impact the knp supersites had in recent years. effectiveness of kruger national park research supersites in achieving original objectives the following sections highlight six of the original objectives of the knp research supersites and reflect on the effectiveness of the sites in achieving these objectives. objective 1: attracting and geographically focussing research to established and well-described study sites based on a subjective assessment, it seems as if the knp research supersites have, at least partially, managed to become geographical focal areas for research within the knp. this geographical focus of studies makes it easier to integrate data sets or infer conditions between studies. the initial smit et al. (2013) paper, which introduced the supersites concept, delineated the sites and provided a backbone of metadata that has proven valuable as a reference document for studies conducted on the supersites ever since. most of the studies that have been conducted on the supersites refer to and cite this paper, as it provides a finer-scale description of the supersites than most other study site references that are usually available only at the regional or park-wide scale. a comment that was raised by critics during the initial phases of delineation of the supersites was that the ‘ideal’ location of the supersites would be highly dependent on the specific objectives of each study, and that it would be hard to select sites that would ‘fit-all’. although this was acknowledged, the idea was never to optimise the supersites for a specific application but rather to delineate sites based on ‘generic’ principles and to make them potentially useful for a wide range of studies. as such, the final sites were selected based on a delineation of nested firstto third-order catchments entirely embedded within a single geology, across the rainfall gradient (north and south of park) and contrasting the geological divide (basalt and granite) in order to represent the dominant abiotic drivers in a nested hierarchy. secondly, these sites had to adhere to some logistical criteria as well (e.g. close to research accommodation, accessible by all-weather roads from multiple directions and outside wilderness zones in order to allow instrumentation). as such, it is encouraging to note that the final supersite locations have proven suitable for a suite of themes, including studies on different taxonomic groups (vegetation, microbes, aquatic invertebrates, small mammals, bats, birds, and large mammals), abiotic patterns and processes (geology, soils, topography and hydrology) and development and use of technology (remote sensing) (see focal themes in online appendix 1). objective 2: promoting integrated understanding across multiple disciplines and scales although there was some cross reference and data sharing within and between hydrological and geological/soil studies, it was apparent that many studies did not integrate with other studies nor were they following a multidisciplinary approach. this is probably to be expected as studies considering very dissimilar taxa or processes and at very different scales would not have obvious and direct linkages or these linkages may not be fully appreciated. also, science is often still approached using traditional discipline and specialist focal areas, with cross-disciplinary studies still in the minority. the multidisciplinary project conducted by the university of the free state, and on which this special issue focusses, is an example where some level of coordination and integration has happened during the initial project design phase, continued through coordinated and spatiotemporally aligned field campaigns, and ultimately resulted in integrative dissemination (this special issue). furthermore, some papers in this special issue attempt to make explicit the links between how the abiotic patterns (soils, geology and topography) influence the abiotic processes (hydrology, soil chemistry), ultimately giving rise to the biotic communities responding to the resulting heterogeneity (vegetation, large mammals, aquatic invertebrates and microbes). this project involved eight principal researchers representing six different departments (soil sciences, groundwater studies, plant sciences, wildlife sciences, environmental sciences and microbiology), doing collective fieldwork campaigns and sharing ideas and data across disciplinary boundaries. janecke et al. (2020) (co-authored by all principle investigators from diverse disciplines) makes a concerted effort to not only summarise but to some degree also integrate between the different studies conducted as part of this project, providing a conceptual framework for biotic and abiotic interactions and feedbacks. objective 3: promoting free sharing of data the supersites concept aims to leverage the ‘data-begets-data’ principle, and for this to function optimally, free sharing and easy accessibility of data are critical. this has been partially successful with the supersites, with data and metadata being archived on a centrally managed south african national parks data repository.1 however, various challenges were also experienced in this regard. some researchers were not allowed or prepared to share data because of a range of reasons (e.g. restrictions linked to funding bodies). in other cases, researchers failed to respond to requests to share data, and knp project coordinators were not effective in following up to ensure all data were appropriately archived. another challenge is that in some cases data got shared and centrally archived, but search terms were ineffective for the data to be associated with the supersites, reducing the likelihood of other researchers working on the supersites being aware of the existence of the data. online appendix 3 provides a list of data sets and key reports currently available for the supersites and also indicates for which supersites the respective data sets are available. objective 4: comparing abiotic contrasts it was initially hoped that studies would collect and compare data across all four supersites in order to better understand the role of rainfall gradients and disparate geologies. this has proven problematic from the start – most of the studies focus only on one of the two southern supersites. this special issue is a point in case, focussing only on one of the four supersites. this is inter alia because of the fact that the southern supersites are in close proximity to skukuza (the main research hub of knp and also the closest supersite to reach from most universities, with the northern supersites adding another day of travelling), and these sites have higher rainfall, deeper soils, more heterogeneity and higher levels of biodiversity. this is further compounded by the financial implications of repeating fieldwork at all supersites. with the exception of remote sensing studies and one geology study, all of the studies in online appendix 1 were conducted on the southern supersites. as such, the northern sites (which represent about half of the knp abiotic template) have not received any notable research attention. it is anticipated that the knowledge and data disparities between the southern and northern supersites will continue to increase. although not a problem per se, it is foreseen that this gap will increasingly widen and the value of the northern supersites may prove to be very limited. research coordinators within knp could play a role in promoting these sites if and where appropriate and where logistical constraints allow. objective 5: understanding long-term ecological dynamics another objective of the supersites was that it would become important long-term monitoring sites, similar to lters. too little time has passed after the initiation of the sites to really assess whether the sites are contributing towards this objective. however, colleagues from the skukuza science leadership initiative (ssli) in partnership with florida university, usa, are already exploring some short-term vegetation and biodiversity trends and dynamics on the southern basalt supersite (2013–2019) (unpublished presentations). van aardt et al. (2020) and janecke et al. (2020) also provide insights into short-term drought dynamics. in addition, various remote sensing projects have also explored long-term woody cover patterns on the supersites (although the historical data sets were collected independently of the establishment of the supersites). objective 6: training sites for remote sensing products although various remote sensing projects have been conducted on the supersites using a range of sensors (aerial light detection and ranging [lidar], aerial photography and optical satellite sensors [i.e. satellite pour l’observation de la terre {spot} 6]), the supersites have not been used as training or validation sites for remote sensing products. this is likely because of relatively little vegetation-related fieldwork and at scales inappropriate for linking to the remote sensing products. it is anticipated that wall-to-wall aerial lidar coverage across the supersites would significantly increase the value of these sites as training sites for development of global remote sensing products for savannas (multiple such requests have been received). unfortunately, it has not been possible to acquire such data sets yet, but potential opportunities are being explored. discussion, recommendations and conclusions the knp supersite concept has gained traction, with research on a range of topics conducted on these sites since their establishment. many of the studies in online appendix 1 could have been conducted at a number of locations within the park, yet the conceptualisation and delineation of the supersites seemed to have been instrumental in geographically focussing research. in the absence of supersites, these studies would most likely have happened at various unrelated sites across the park. furthermore, the conceptual appeal and the rationale behind the supersites may even have attracted research projects to the park that may otherwise not have happened. it is believed that some of the reasons for research projects focussing on the supersites may have been (1) the clear ecological rationale behind the site selection, (2) the logistical benefits of working at these sites (i.e. easy access to research camps and other research support services), (3) the availability of good basic metadata and descriptions of these sites and (4) other research projects also being conducted at these sites, allowing opportunities for data sharing and collaboration. going forward, a concerted effort should be made to ensure that data management is improved to ensure that all supersite data are systematically collated, archived and easily accessible. even if data archiving infrastructure is in place, follow-up and coordination is important to ensure that datasets get collated and centrally archived in a database with appropriate search terms. it is proposed that this database be continually updated in order to (1) increase awareness of the available data sets, (2) attract further research to the sites, (3) facilitate data sharing and integration, and ultimately (4) increase understanding of the role of top-down and bottom-up processes in savanna ecosystems. some limited measuring equipment (e.g. soil moisture meters) and small-scale manipulations (herbivore exclosures) have been added in recent years at some of the sites, which may further increase the value and research uptake. installation of further long-term equipment (e.g. weather stations) and wall-to-wall coverage of valuable datasets (e.g. high-resolution airborne lidar and derived products like digital terrain model [dtm] and digital surface model [dsm]) should be promoted, as it will act as additional catalysts for further studies on the supersites. in addition, the sites may gain further traction if they become more formally part of the research infrastructure networks and increase their involvement with international collaborators and lter initiatives. dedicated research budget, solicited research and well-funded projects on these sites can also contribute towards the objectives set out in this essay. where the knp research supersites provide a good ‘fit’ for the purposes of studies, they should be actively promoted as potential field sites. as per their original conceptualisation, it is believed that the longer these sites are in existence and the more they are studied (and with associated data sets becoming freely available), the more valuable they will become and the more research attention they will attract, contributing towards answering questions that would not be possible with individual projects, which are typically funded only for threeto five-year cycles. this special issue, together with the earlier work of riddell et al. (2014), and other studies listed in online appendix 1 are a testament to how the knp supersites are contributing towards improved understanding of bottom-up and top-down drivers of and responders to savanna heterogeneity. it is hoped that over time, and as more studies are conducted on these sites, more integration would emerge between studies that can benefit from multidisciplinary approaches. acknowledgements the author thanks edward s. riddell, beanelri janecke and johan van tol for comments on an earlier draft. competing interests the author declares that he has no financial or personal relationships that may have inappropriately influenced him in writing this essay. author’s contribution i.p.j.s. is the sole author of this research article. ethical considerations this research followed all ethical standards for research without direct contact with human or animal subjects. funding information this research received no specific grant from any funding agency in the public, commercial or not-for-profit sectors. data availability data sharing is not applicable to this essay as no new data were created or analysed. disclaimer the views and opinions expressed in this essay are those of the author and do not necessarily reflect the official policy or position of any affiliated agency of the author. references asner, g.p., levick, s.r., kennedy-bowdoin, t., knapp, d.e., emerson, r., jacobson, j. et al., 2009, ‘large-scale impacts of herbivores on the structural diversity of african savannas’, proceedings of the national academy of sciences 106(12), 4947–4952. https://doi.org/10.1073/pnas.0810637106 gosz, j.r., waide, r.b. & magnuson, j.j., 2010, ‘twenty-eight years of the us-lter program: experience, results, and research questions’, in f. müller, c. baessler, h. schubert & s. klotz (eds.), long-term ecological research, pp. 59–74, springer, dordrecht. higgins, s.i., bond, w.j., february, e.c., bronn, a., euston-brown, d.i., enslin, b. et al., 2007, ‘effects of four decades of fire manipulation on woody vegetation structure in savanna’, ecology 88(5), 1119–1125. https://doi.org/10.1890/06-1664 janecke, b.b., 2020, ‘vegetation structure and spatial heterogeneity in the granite supersite, kruger national park’, koedoe 62(2), a1591. https://doi.org/10.4102/koedoe.v62i2.1591 janecke, b., van tol, j., smit, i.p.j., van aardt, a.c., riddell, e.s., seaman, m.t. et al., 2020, ‘biotic and abiotic connections on a granitic catena: framework for multidisciplinary research’, koedoe 62(2), a1600. https://doi.org/10.4102/koedoe.v62i2.1600 majozi, n.p., mannaerts, c.m., ramoelo, a., mathieu, r.s., nickless, a. & verhoef, w., 2017, ‘analysing surface energy balance closure and partitioning over a semi-arid savanna fluxnet site in skukuza, kruger national park, south africa’, hydrology and earth system sciences 21(7), 3401–3415. https://doi.org/10.5194/hess-21-3401-2017 mirtl, m., borer, e.t., djukic, i., forsius, m., haubold, h., hugo, w. et al., 2018, ‘genesis, goals and achievements of long-term ecological research at the global scale: a critical review of ilter and future directions’, science of the total environment 626, 1439–1462. https://doi.org/10.1016/j.scitotenv.2017.12.001 riddell, e.s., nel, j., fundisi, d., jumbi, f., van niekerk, a. & lorentz, s.a., 2014, ephemeral hydrological processes in savannas, water research commission report, gezina, pretoria. smit, i.p.j., riddell, e.s., cullum, c. & petersen, r., 2013, ‘kruger national park research supersites: establishing long-term research sites for cross-disciplinary, multiscaled learning’, koedoe 55(1), 01–07. https://doi.org/10.4102/koedoe.v55i1.1107 smit, i.p.j., roux, d.j., swemmer, l.k., boshoff, n. & novellie, p., 2017, ‘protected areas as outdoor classrooms and global laboratories: intellectual ecosystem services flowing to-and-from a national park’, ecosystem services 28, 238–250. https://doi.org/10.1016/j.ecoser.2017.05.003 van aardt, a.c., codron, d., theron, e.j. & du preez, p.j., 2020, ‘plant community structure and possible vegetation changes after drought on a granite catena in the kruger national park, south africa’, koedoe 62(2), a1585. https://doi.org/10.4102/koedoe.v62i2.1585 van wilgen, b.w., boshoff, n., smit, i.p.j., solano-fernandez, s. & van der walt, l., 2016, ‘a bibliometric analysis to illustrate the role of an embedded research capability in south african national parks’, scientometrics 107(1), 185–212. https://doi.org/10.1007/s11192-016-1879-4 footnotes 1 see https://dataknp.sanparks.org/sanparks/ (using ‘supersites’ as the search keyword). filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 koedoe 60-1_2018_contents.indd http://www.koedoe.co.za open access table of contents i original research plant communities of the umlalazi nature reserve and their contribution to conservation in kwazulu-natal nqobile s. zungu, theo h.c. mostert, rachel e. mostert koedoe | vol 60, no 1 | a1449 | 28 may 2018 original research ground-dwelling spider assemblages in contrasting habitats in the central south african grassland biome charles r. haddad, vivian p. butler koedoe | vol 60, no 1 | a1482 | 31 may 2018 original research movement patterns and home range size of tigerfish (hydrocynus vittatus) in the incomati river system, south africa francois roux, gert steyn, clinton hay, ina wagenaar koedoe | vol 60, no 1 | a1397 | 27 june 2018 original research a survey of the ichthyofauna in the noetsie estuary, western cape province, south africa martin k.s. smith, demi rodrigues, bianca currie koedoe | vol 60, no 1 | a1513 | 16 july 2018 original research long-term variability in vegetation productivity in relation to rainfall, herbivory and fire in tswalu kalahari reserve wataru tokura, sam l. jack, tania anderson, michael t. hoffman koedoe | vol 60, no 1 | a1473 | 31 july 2018 original research the 2013–2014 vegetation structure map of hwange national park, zimbabwe, produced using free satellite images and software eduardo m. arraut, andrew j. loveridge, simon chamaillé-jammes, hugo valls-fox, david w. macdonald koedoe | vol 60, no 1 | a1497 | 27 september 2018 original research seasonal variation in species richness and abundance of waterbirds in mole national park, ghana: implication for conservation and ecotourism timothy k. aikins, francis gbogbo, erasmus h. owusu koedoe | vol 60, no 1 | a1466 | 02 october 2018 reviewer acknowledgement koedoe | vol 60, no 1 | a1561 | 27 november 2018 59 73 85 98 105 123 133 142 page i of i table of contents i book review review of national park science: a century of research in south africa by jane carruthers peter a. novellie koedoe | vol 60, no 1 | a1550 | 06 september 2018 short communication blowflies as vectors of bacillus anthracis in the kruger national park lizanne basson, ayesha hassim, at dekker, allison gilbert, wolfgang beyer, jennifer rossouw, henriette van heerden koedoe | vol 60, no 1 | a1468 | 26 june 2018 short communication challenges and opportunities for monitoring wild nile crocodiles with scute mark-recapture photography bernard w.t. coetzee, sam m. ferreira, kristine maciejewski koedoe | vol 60, no 1 | a1505 | 19 july 2018 short communication occupancy and habitat use by six species of forest ungulates on tiwai island, sierra leone kathryn r. mccollum, emily belinfonte, april l. conway, john p. carroll koedoe | vol 60, no 1 | a1484 | 29 august 2018 checklist south african national survey of arachnida: a checklist of the spiders (arachnida, araneae) of the tswalu kalahari reserve in the northern cape province, south africa anna s. dippenaar-schoeman, charles r. haddad, robin lyle, leon n. lotz, stefan h. foord, rudy jocque, peter webb koedoe | vol 60, no 1 | a1486 | 09 july 2018 review article community harvesting of trees used as dens and for food by the tree hyrax (dendrohyrax arboreus) in the pirie forest, south africa elizabeth j. opperman, michael i. cherry, nokwanda p. makunga koedoe | vol 60, no 1 | a1481 | 28 february 2018 original research settling the browser–grazer debate for african buffalo in grass-limited eastern cape thicket, south africa marietjie landman, kate kloppers, graham i.h. kerley koedoe | vol 60, no 1 | a1465 | 22 february 2018 original research the operational competitiveness of public protected areas managed by ezemvelo kzn wildlife karabo d. motau, edilegnaw wale koedoe | vol 60, no 1 | a1452 | 27 march 2018 1 4 10 15 20 31 40 44 vol 60, no 1 (2018) issn: 0075-6458 (print) | issn: 2071-0771 (online)koedoe abstract introduction description of the study area methods and material results discussion conclusion acknowledgements references about the author(s) manish ramjeawon south african environmental observation network, pietermaritzburg, south africa centre for water resources, college of agriculture, engineering and science, university of kwazulu-natal, pietermaritzburg, south africa molla demlie department of geology, college of agriculture, engineering and science, university of kwazulu-natal, pietermaritzburg, south africa michele l. toucher south african environmental observation network, pietermaritzburg, south africa centre for water resources, college of agriculture, engineering and science, university of kwazulu-natal, pietermaritzburg, south africa susan janse van rensburg south african environmental observation network, pietermaritzburg, south africa citation ramjeawon, m., demlie, m., toucher, m.l. & janse van rensburg, s., 2020, ‘analysis of three decades of land cover changes in the maputaland coastal plain, south africa’, koedoe 62(1), a1642. https://doi.org/10.4102/koedoe.v62i1.1642 original research analysis of three decades of land cover changes in the maputaland coastal plain, south africa manish ramjeawon, molla demlie, michele l. toucher, susan janse van rensburg received: 08 june 2020; accepted: 18 aug. 2020; published: 09 nov. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the northern half of the maputaland coastal plain (mcp) of south africa has undergone significant change in land-use over the last five decades, including afforestation, agriculture and rural settlements. to understand the extent of land-use changes that took place in the northern half of the mcp from 1986 to 2019 and its efficacy, various landsat satellite images that are freely available were processed, analysed and interpreted. the cloud-based google earth engine (gee) platform was used to determine the land-use changes. the random forest classification algorithm available within gee was used to classify the landsat 5 thematic mapper, landsat 7 enhanced thematic mapper plus and landsat 8 operational land imager images for 1986, 1991, 1996, 2001, 2006, 2011, 2016 and 2019. the development of plantations was tracked, which indicated that forestry increased by more than 100% between 1986 and 2019. over the same period, surface water bodies and wetlands decreased by 36.1% and 49.1%, respectively. in addition to forestry, climate had a major impact on water resources in the mcp. given that the mcp is a predominately groundwater-driven system, the impact of increased plantations on groundwater is an area that requires more investigation. this will improve the understanding of water resources in the area. conservation implications: the management, protection and conservation of water resources within protected areas are entwined with land-use decisions and planning outside of its boundaries. the rapid change in land-use experienced outside of protected areas and its impact on water resources disregard boundaries and may transgress on protected areas. keywords: google earth engine; landsat imageries; land-use change; remote sensing; north-eastern south africa; water resources; random forest classification; land cover. introduction changes in land use and land cover over time and space indicate socioeconomic change. population growth and consequent demand for economic development have increased the conversion of natural vegetation. the kwazulu-natal province has experienced an accumulated transformation of 45.6% of natural vegetation between 1994 and 2008 (jewitt 2012:12). one area that has been affected by land cover transformation is the maputaland coastal plain (mcp) located in the north-east of the kwazulu-natal province of south africa, which is rich in biodiversity and other natural resources (orimoloye et al. 2019:3). the mcp has several wetlands, lakes and game reserves, which play an integral role in maintaining a healthy ecosystem. the largest freshwater lake in south africa, lake sibayi, and the isimangaliso wetland park, which is a renowned ramsar and world heritage site, are located within the mcp (weitz & demlie 2014:1). however, recent studies indicated that surface waters and wetland systems in the mcp are under severe threat because of plantations and water extraction (everson et al. 2019:1; orimoloye et al. 2019:9; smithers et al. 2017:490). a large part of the lake sibayi catchment area is covered by plantations (everson et al. 2019:34; weitz & demlie 2014:1). mbazwana and manzegwena, the two largest plantations, cover approximately 8500 and 15 000 ha, respectively, and were established in the 1960s by the kwazulu department of forestry (sa forestry 2012:1). pine plantations are deep-rooted, evergreen, non-native tree species that are known to extract soil water from below the root zone, limiting aquifer recharge and resulting in a lowered groundwater table (kienzle & schulze 1992:1). as a result of this, the nati onal water act (act no. 36 of 1998) of south africa considers plantations as a streamflow reduction activity, and as such, plantations require a water use license from the government. in coastal areas such as the mcp, favourable solar radiation and air temperatures facilitate year-round growth and provide plantations the opportunity to access the shallow aquifers, resulting in higher water consumption than the surrounding natural vegetation (everson et al. 2019:34). this has had a significant impact on surface water and wetland systems in the mcp as many of the surface water systems and wetlands in the mcp are predominately groundwater driven (kelbe & germishuyse 2010:124). studies as early as the 1990s highlighted the negative impact that plantations had on the hydrology and ecology of the mcp. a study using aerial photography conducted by rawlins (1991:30) indicated that the size and number of wetlands reduced as a result of the introduction of plantations in the area. wetlands were reported to exist during the early growth stages of the plantations; however, these dried up as the plantations matured and were only visible for a short period after heavy rainfall (rawlins 1991:30). smithers et al. (2017:490) indicated that 35% of the total decrease in the level of lake sibayi was a result of forest plantations surrounding the lake. orimoloye et al. (2019:1) opined that a 38% decrease in wetlands in the isimangaliso wetland park between 1987 and 2017 was because of the impact of plantations. everson et al. (2019:5–8) investigated the water use of plantations and agroforestry systems in the mcp and deduced that eucalyptus trees use more water in a day when compared with the indigenous vegetation. modelling results from the study indicated that the removal of forest plantations from the vasi area of the mcp will result in a 2-m increase in the groundwater table. furthermore, a 40% increase in water yield was predicted if the plantations were to be removed from the immediate vicinity of wetlands and replaced by agroforestry. despite these limited and specific spatiotemporal studies, the long-term impacts of land-use change, including plantations on water resources and wetland systems in the mcp, are not yet well understood and documented. although land cover information such as the ezemvelo kwazulu-natal wildlife (ekznw) maps is available, they do not extend back beyond 2005. thus, there is a need for regularly updated and long-term land-use data to improve the understanding of changes in the land surface and the impact on water resources including wetlands. recently, the google earth engine (gee) platform has been increasingly used in conjunction with earth observation data in change detection studies (google earth engine 2018). as the gee platform is cloud based, it increases the efficiency of producing long-term records of data. a 40-year collection of landsat earth observation data is available within the gee platform, providing an opportunity to produce land-use data over a longer time frame. previous studies are limited in their temporal and spatial investigation of afforestation in the mcp. the availability of a long-term data set of afforestation for the mcp provides an opportunity to expand on the outcomes of short-term studies to gain a better understanding of the hydrology over time. this study aims to track the development of plantations over a 30-year period and its potential impacts on surface water and wetlands systems in the mcp using supervised classification. description of the study area location and climate the mcp is located in the north-eastern coastal plain of the kwazulu-natal province, south africa (figure 1). the coastal plain extends from mlalazi in the south to mozambique in the north. the coastal plain is bounded by the lebombo mountains in the west and the indian ocean in the east. the present study focuses on the region stretching from the northern perimeter of the mkhuze swamp in the south to the south africa–mozambique national boundary in the north (figure 1). the mcp falls under the usutu to mhlatuze water management area and has a population of approximately 573 353 people (statistics sa 2011). it is described as one of the most underdeveloped areas in south africa as unemployment, access to piped water, lack of electricity and low education levels are prevalent in the region (statistics sa 2011). figure 1: location map of the study area in north-eastern south africa. the climate of the study area is subtropical and receives summer rainfall, with a 30-year mean annual precipitation of approximately 580 mm (figure 4) (saws 2018). the majority of the rainfall occurs during the summer months, between october and march, and is highly spatially and temporally variable. the rainfall decreases from the coast inland towards the lebombo mountains. mean daily temperatures vary from 21.0 °c to 32.1 °c in summer and 10.9 °c to 26.4 °c in winter (saws 2018). the mean potential evaporation rate for the study area is approximately 1458 mm year-1 (dws 2018). topography, drainage and land cover the topography of mcp is generally flat. the highest elevation in the area is approximately 700 m above mean sea level and occurs along the lebombo mountain range in the western boundary of the study area. from here, the elevation drops to sea level at the coast. the surface water bodies (lakes, rivers and wetlands) are predominately groundwater driven (kelbe & germishuyse 2010:124). the main rivers that drain the mcp are the umfolozi, mhlathuze, pongola and mkuze rivers (kelbe & germishuyse 2010:124). the umfolozi river that once formed part of the st. lucia wetland has since separated from the wetland. the mhlathuze river flows into the richards bay harbour, and the pongola river flows into mozambique towards the maputo bay. the mkuze river, along with the nylazi, hluhluwe and mzinene rivers, flow into lake st. lucia (kelbe & germishuyse 2010:124). lake sibayi and the kosi bay lake systems are the two main lakes in mcp (figure 1). the study area forms part of the maputaland-pondoland-albany hotspot. indian ocean coastal belt (iocb), savanna and forest are the three biomes covering the study area. the iocb runs along the coast and up to 20 km inland. it consists of the maputaland coastal belt and maputaland wooded grassland (mucina & rutherford 2006:573). the central and western portions consist of vegetation forming the savanna biome. this includes the maputaland pallid sandy bushveld, muzi palm veld and wooded grassland, tembe sandy bushveld, makatini clay thicket, western maputaland sandy bushveld and southern lebombo bushveld, which cover the entire lebombo mountain range (mucina & rutherford 2006:573). the sand forest, the lowveld riverine forest and the northern coastal forest are distributed throughout the study area (mucina & rutherford 2006:571). adjacent to st. lucia, the wetland type vegetation is evident through the azonal vegetation identified in sanbi (2018). large areas of the iocb and savanna biomes have been replaced by plantations and commercial agriculture, with homesteads distributed throughout the biome regions. methods and material earth observation data and data processing the multi-year land-use maps reported in this study were produced using javascript in gee application programming interface. google earth engine (gee) is a cloud-based platform for earth science and data analyses (google earth engine 2018). google earth engine (gee) contains a multi-petabyte catalogue of over 40 years of satellite imagery and geospatial data sets with massive computational abilities available for users to perform studies relating to change detection, trend mapping and quantify changes on the earth’s surface (google earth engine 2018). landsat 5 thematic mapper (tm), landsat 7 enhanced thematic mapper plus (etm+) and landsat 8 operational land imager (oli) surface reflectance data were used. as the operational period for each satellite is different, a combination of three satellites was used. landsat 5 tm was launched in march 1984 and decommissioned in june 2013. landsat 7 etm+ and landsat 8 oli were launched in april 1999 and february 2013, respectively, and are still operational (usgs 2018). all have the same 30-m spatial resolution. the flow diagram (figure 2) describes the process undertaken in gee to classify the landsat images. the landsat collection was filtered to acquire one cloud-free image every 5 years from 1986 to the most recent 2019 cloud-free images. moreover, a cloud-free landsat image for 2018 was obtained and used to validate against the 2018 south african national land-cover (sanlc) map. all but one of the cloud-free images were available during the winter months. the aim of this study was to determine long-term trends in land use and land cover, not the seasonal variability. thus, although it is acknowledged that in the wetland, lake levels will be lower in the dry, winter season, the consistency of using only winter images will allow the long-term changes to be identified. the images used in this study were for 04 november 1986, 23 july 1991, 18 june 1996, 16 june 2001, 14 june 2006, 23 august 2011, 12 august 2016, 15 june 2018 and 04 july 2019. the availability of images was limited because of cloud cover. figure 2: flow chart summarising the methods followed. supervised classification supervised classification is based on the premise that prior knowledge of the land classes in the study area is known (richards & jia 2006:78). the users determine the land cover types to be investigated, and representative pixels for each type are identified. this is known as training, and the information is obtained either from field surveys, aerial photographs, maps, satellite imagery or a combination of these (richards & jia 2006:80). once the pixels are trained, a classifier is used to classify the rest of the image (richards & jia 2006:298). several classifiers are available, including the maximum likelihood, minimum likelihood, support vector machine and random forest. training data training data were obtained by visual inspection of terrametrics 15-m spatial resolution satellite imagery, the ekznw land cover map and field data. ezemvelo kwazulu-natal wildlife land cover map is a free data set available on request for 2005, 2008, 2011 and 2017. the ekznw data sets were selected as they are well-recognised published maps for kwazulu-natal that have relatively high overall accuracies (thompson 2018:21). the ekznw land cover maps were produced using sentinel 2 satellite imagery based on a 20-m spatial resolution (thompson 2018:5). in addition to the use of the ekznw maps, a field survey was conducted between 24 june 2019 and 28 june 2019, where a total of 65 in situ land-use data points were collected. the location of the data points was generated using a random point generator tool. the data points were collected around the study site and consist of 18 grasslands, 16 plantations, 14 indigenous vegetation, 7 wetlands, 5 urban, 4 bare ground and 1 agriculture data point. these data were used to assist in identifying land uses in the study area for training purposes. overall, 35 water body, 25 urban, 25 bare soil, 25 agriculture, 35 plantations, 50 grasslands, 50 natural vegetation, 30 wetlands and 30 degraded land reference polygons, accounting for over 70 000 pixels, formed the reference data set, which was used to identify landsat pixels that made up one of the nine land-use classes. the nine land-use classes were chosen as these were identified as the dominant land covers, which could be clearly distinguished for the study area. the nine land-use classes are water bodies, urban, bare ground, agriculture, plantations, grasslands, indigenous vegetation, wetlands and degraded land. bare ground is described as a sand cover found typically on the coastline, whereas degraded land consisted of partial vegetation cover. from the reference data set, 70% of the sample data were randomly selected for training, and the remaining 30% was used to validate the data set. the training data was used for the supervised classification algorithm, and the validation data were used in the accuracy assessment of the classified land-use maps. image classification the random forest classification algorithm has become a popular model for land-use and land cover classification as it has produced more accurate results in comparison to other classification algorithms (li et al. 2014:21; reynolds et al. 2016:2). the random forest classification is easier to operate, more efficient and robust to outliers and can handle a large amount of training data (rodriguez-galiano & chica-rivas 2012:13). the random forest classifier has been used in several studies to map forests (mellor et al. 2012; senf, hostert & van der linden 2012), wetlands (berhane et al. 2018; chignell et al. 2018), agriculture (lebourgeois et al. 2017; ok, akar & gungor 2012) and land cover (jin et al. 2018; na et al. 2010). the random forest classifier consists of many decision trees that operate as an ensemble and requires two parameters to be identified to make accurate class predictions, namely, the number of predictor variables and the number of classification trees (yiu 2019:1; zurqania et al. 2018:178). consequently, the random forest classification algorithm was adopted and used in gee for this study. accuracy assessment accuracy assessment techniques are used to determine how well an image has been classified (nasa 2018:2). the validation data set was used to produce an error matrix for each of the classified images. the error matrix was used to determine the overall accuracy, kappa coefficient, user’s accuracy and producer’s accuracy using equations 1, 2, 3 and 4, respectively. the user’s accuracy (error of commission) occurs when a pixel is incorrectly included in a category being evaluated, whereas the producer’s accuracy (error of omission) occurs when a pixel is left out of the category being evaluated (nasa 2018:17). the kappa coefficient is a measure of inter-rater reliability and is used to determine the agreement between two data sets. the user’s and producer’s accuracies were generated for all nine classes for each image: where pagree is the agreement observed and pchance is the agreement expected by chance. in addition to the error matrix, a land-use map was produced for 2018 and compared with the sanlc (2018) map. the sanlc map was chosen as it is a well-recognised published map for south africa with an overall accuracy of 90.1%. classes from the sanlc (2018) map had to be merged to form classes that were comparable to the nine classes defined for this study. open water bodies, such as rivers, estuaries, lakes, dams and natural pans, were merged to form the ‘water’ land cover. residential areas including informal settlements, scattered and dense villages, commercial, industrial and roads were merged to form the ‘urban’ land cover. natural rock, dry pans, coastal dunes and dry riverbeds were merged to form ‘bare ground’. cultivated commercial permanent orchards, cultivated commercial sugarcane (pivot and non-pivot), emerging farmer sugarcane non-pivot, commercial annual crops (irrigated and drylands) and subsistence farms were merged to form the ‘agriculture’ land cover. contiguous and dense plantation forests, open and sparse plantation forests and temporary unplanted (clear-felled) plantation forests were merged to form ‘plantations’. natural grasslands were merged to form ‘grasslands’. contiguous (indigenous) forests, contiguous low forests and thicket, dense forests and woodland, open woodland and low shrubland were merged to form ‘indigenous vegetation’. herbaceous wetlands and mangrove wetlands were merged to form ‘wetlands’. ethical consideration the authors confirm that ethical clearance was not required for the study. results the results obtained from the accuracy assessment, the comparison between the 2018 classified landsat image and the sanlc (2018) map as well as the spatiotemporal changes for plantations, water bodies and wetlands from 1986 to 2019 are presented in this section. accuracy assessment the overall accuracies achieved are all greater than 92% with the highest accuracy of 96.3% for the 1996 image (table 1). the highest and lowest kappa coefficients are 0.95 (1996) and 0.89 (2011) (table 1). the 2011 image was the only image based on the landsat 7 etm+ satellite data and has the lowest overall accuracies and kappa coefficient. this could be a result of scan lines present on the landsat 7 etm+ images. table 1: the user’s accuracy, producer’s accuracy, overall accuracies and kappa coefficients for the classified landsat images. the user’s accuracy and producer’s accuracy range from 60.2% to 99.9% and from 64.8% to 100%, respectively (table 1). water bodies proved to be the most accurately predicted land-use class across all images with user’s and producer’s accuracies ranging between 99.7% and 99.9% and 99.7% and 100%, respectively. the least accurately predicted class was the urban areas with a maximum user and producer accuracy of 72.3% and 76.5%, respectively. a majority of the settlements in the study area are located within grasslands; as a result, some settlement areas, which should have been classed as urban land use, are classed as grasslands in the majority of the images classified. the plantation land-use class produced user’s and producer’s accuracies ranging from 71.6% to 92% and 72.0% to 91%, respectively. validation of the 2018 landsat classified map the 2018 landsat classified land-use map was compared with the 2018 sanlc map (2018) as part of the validation study. the water bodies, wetlands and plantations land-use classes were compared. the regression indicates a good correlation between the two data sets with a coefficient of determination (r2) of 0.92. water bodies have the least difference in area between the two maps followed by wetlands and plantations. water bodies covered a total area of 10 303 ha according to the sanlc (2018) map, whereas the landsat classified map had a total area of 11 260 ha (table 2). the landsat classified map overestimated water bodies, wetlands and plantations by approximately 9.3%, 4.4% and 29.3%, respectively. these small discrepancies may be attributed to the fact that the sanlc (2018) map was produced at a 20-m spatial resolution compared with the landsat image, which has a 30-m spatial resolution. table 2: area comparison between south african national land-cover map (2018) land cover and the landsat 2018 classified map. land-use changes in the maputaland coastal plain from 1986 to 2019 plantations, water bodies and wetland land-use classes from 1986 to 2019 were selected and assessed. most of the plantations are located along the coastline, just north and south of lake sibayi (figure 3). plantations increased at an average rate of 235 ha year-1 between 1986 and 1991, which increased to approximately 2372 ha year-1 by 2019 (table 3). this indicates that the rate of change increased 10-fold between 1986 and 2019. the plantation coverage increased by about 22 549 ha between 1986 and 2019. initially, the plantation was concentrated around lake sibayi. between 1986 and 2001, a 5936-ha increase in plantations was observed. this expansion of plantation occurred in the region north of lake sibayi with smaller plantations developing west of lake kosi bay. in 2006, there was a decrease in forestry, possibly because of felling. following this, a 35.4% (11 193 ha) increase in plantations occurred between 2006 and 2019. the classified images between 2006 and 2019 (figure 3) indicate that the expansion of commercial plantations was concentrated in the area west of kosi bay and to a smaller extent west of lake sibayi. figure 3: spatial distribution of forests, water bodies and wetlands for the maputaland coastal plain between 1986 and 2019. figure 3 (continues...): spatial distribution of forests, water bodies and wetlands for the maputaland coastal plain between 1986 and 2019. table 3: net area change (ha year−1) between measurement periods for plantations, surface water bodies and wetlands. surface water bodies and wetlands have experienced a decline since 1986. the largest rate of change was experienced during the 2001–2006 period for surface water bodies with an average decrease of 444 ha year-1. during the same period, wetlands experienced the largest rate of decline of 1383 ha year-1 (table 3). open water bodies decreased by 17.5% between 1986 and 1996 (figure 4). however, water bodies increased by 0.5% between 1996 and 2001. from 2001 to 2019, a more significant decrease by 22.9% was observed. an overall decrease of 36.1% occurred between 1986 and 2019. the largest rate of change was experienced during the 2001–2006 period with an average decrease of 444 ha year-1 (table 3). the extent of lake sibayi and water bodies located in the middle of the study area visibly declined in the images from 1986 to 2019 (figure 3). between 2001 and 2006, the southern basin of lake sibayi became noticeably smaller (figure 3). by 2016, the southern portion of the lake was disconnected from the main lake and continued to decrease, as can be seen in the 2019 image (figure 3). the water bodies located near the middle of the study area have also decreased between 1986 and 2019 and, in some instances, disappeared. the water bodies in the kosi bay area do not seem to have been affected that much. within the conservation area, water bodies decreased by 17.5% between 1986 and 2019. figure 4: change in area covered by plantations, water bodies and wetlands, as well as annual precipitation for the study area, for the period from 1986 to 2019. the distribution of wetlands located in the study area is shown in the classified images of figure 3, where the most extensive one is located in the southern section of the study area. the extent of wetlands is visibly smaller in the 1986 image as a result of below-average rainfall received in 1985 (dws 2018). between 1986 and 1991, a 16.7% increase in wetlands occurred (figure 4), particularly along and around water bodies in the northern region of the study area and in the south-west section of the study area (figure 3). the extent of wetlands remained relatively constant with a 705.7-ha difference between 1996 and 2001. the extent began to decrease significantly from 2001 onwards (figure 4). between 2001 and 2019, wetlands decreased by 47.9%. the highest rate of wetland decline was experienced during the 2001 and 2006 period at a rate of 1383 ha year-1 (table 3). a decrease in the extent of the isimangaliso wetland in the south and wetlands located on the northern boundary of the study area is noticeable in the classified maps. generally, wetlands experienced an overall decrease of 15 862.6 ha (49.1%) during the study period. wetlands within the conservation area increased between 1986 and 1996, and after which, their extent decreased by an overall 18%. urban areas increased by 36.5% between 1986 and 1996 (figure 4). this was followed by a decrease of 4.8% between 1996 and 2001. overall, the urban land class increased by 38% between 1986 and 2019. discussion image processing, classification and validation the random forest classification algorithm was used within the gee platform to produce land-use maps for the northern mcp for the years 1986, 1991, 1996, 2001, 2006, 2016 and 2019 to assess long-term trends and changes. the results of the validation study indicate a good overall goodness of fit (r2 = 0.92). the overall accuracy ranged between 92.3% and 96.3%. the 2011 image had the lowest overall accuracy (92.3%) because of the gap-filled scan lines present in the landsat 7 etm+ image, which may have affected the accuracy of the classification. the landsat 7 etm+ 2011 image was used as there was no cloud-free image available from landsat 5. the overall accuracy decreased from 1986 to 2019. this could be zattributed to the change in land cover over time. as land cover heterogeneity increases, the probability of misclassifying pixels increases, which influences the overall accuracy (smith et al. 2002:69). the error matrix for all eight images indicated that urban areas were the least accurate land cover class predicted. the reason for the poor estimation of the urban land cover class could be attributed to the 30-m spatial resolution of the landsat satellite data, which failed to capture the homesteads in the mcp, as many of them are smaller than the satellite resolution. furthermore, the yards are not tarred or paved and are covered by grass or shrub vegetation and small subsistence vegetable plots, which may have been incorrectly mapped as grassland land cover. improved classification of urban areas can be achieved using a finer resolution satellite such as sentinel (10 m resolution); however, imagery is not available for long-term studies. the lower accuracies achieved for the grasslands, plantation and agricultural classes may be attributed to the random forest classification algorithm. jin et al. (2018:17) identified normalized difference vegetation index (ndvi) as the most important variable that influenced the random forest classifier. the limited availability of water, particularly in recent years, may have influenced the ndvi of grasslands, plantations and rainfed agriculture lands during the winter season and hence affected the results achieved. the 30-m spatial resolution appeared to be suitable for water bodies and wetlands as the overall accuracies for these classes were relatively high. land-use/land cover change the landsat classified maps from 1986 to 2019 indicated an overall increase in size and number of plantations, wherein it more than doubled during the last three decades. the manzengwenya and mbazwana plantations have expanded but most notable was the development of new plantations near the kosi bay area and west of lake sibayi. these findings were in line with earlier reports by weitz and demlie (2014:1) and everson et al. (2019:69), who expressed concern that plantations in the mcp were increasing. the new plantation developments are visibly smaller than the manzengwenya and mbazwana plantations and are likely related to small-scale woodlots grown by subsistence farmers. sa forestry (2012:1) reported that mondi, a timber production company, has been assisting the local communities that are located from sodwana bay to the mozambique border in establishing small-scale commercial plantations as means of livelihoods. another timber production company, sappi, has established a programme that provided small-scale growers with a minimum of 5 ha (sa forestry 2012:1). one such site is established near manguzi, near lake kosi bay (sappi 2020:1). many studies highlighted the high water use of plantations compared with the natural vegetation it replaces (dye 1996:33; dye & versfeld 2007:122; everson et al. 2019:6). the evergreen canopies, high leaf area index and deep roots coupled with favourable climatic conditions are the drivers behind the high water use of plantations (dye & versfeld 2007:122; everson et al. 2019:138). the deep root systems of plantations allow the trees to access groundwater, which is not accessible by shallow-rooted grasslands and poses a threat to groundwater systems. water use of pinus plantations from three summer-rainfall catchments in south africa, namely, cathedral peak, mokobulaan a and mokobulaan b, ranged between 750 and 900 mm year-1 (dye 1996:32). recently, everson et al. (2019:5) estimated the average daily water use of individual trees in the mcp. small eucalyptus trees had an average daily water use of 19.2 l day-1, whereas the average daily water use of mature trees was 8.0 and 11.41 l day-1 for pinus and eucalyptus, respectively. in nebraska sand hill, united states, adane et al. (2018:180) reported that annual average groundwater recharge rates decreased from 9.65 to 0.07 cm year-1 when natural grasslands were replaced with pinus plantations. ndlovu and demlie (2018:1) reported that groundwater level around the tembe and mbazwana areas in mcp decreased by 0.7 m and 2.7 m, respectively. the landsat classified maps revealed a decrease in water bodies and wetlands in the study area during the same period. the main drivers for the reduction in the extent of open water bodies and wetland systems are the expansion of plantations and the change in climate. the results of this study are consistent with the findings of rawlins (1991:30) and smithers et al. (2017:490), where plantations are reported to have induced a reduction in the extent of surface water and wetland systems. since the early 1990s, studies have indicated that plantations have reduced the size and number of wetlands in the mcp (rawlins 1991:30). rawlins (1991:30), everson et al. (2019:24) and orimoloye et al. (2019:1) noted the reduction in wetlands linked to plantations, with orimoloye et al. (2019:1) finding a 38% decrease in wetlands in the isimangaliso wetland park between 1987 and 2017. moreover, grundling, van den beg & price (2013:135) reported that wetlands in the mcp decreased as a result of land-use change and drought. the present study found an overall decrease of 49.1% in wetlands that is far more than that reported by orimoloye et al. (2019), which is related to the difference in the size of the study area investigated and the length of time considered. smithers et al. (2017:490) indicated that plantations were the cause of a 35% drop in the water level of lake sibayi and that removal of the forest from the catchment area would increase water yield substantially. elsewhere in the world, d’amato et al. (2017:270) stated that plantations in southern china had negative impacts on water quantity and suggested the removal of plantations to improve water availability. immediate increase in water yield following the removal of plantations has been reported in many countries, including south-eastern australia (bren & hopmans 2007:1), india (sikka et al. 2003:1) and several andean countries (bonnesoeur et al. 2019:575). similarly, woodward et al. (2014:1) reported that the removal of forests has increased the water available to wetlands and, in some instances, created new wetlands, particularly in australia and new zealand. climate has a major impact on water resources in the mcp. pitman and hutchison (1975) highlighted that lake sibayi was highly sensitive to changes in local climate. in addition to forestry, the decrease in the extent of water bodies reported in this study for the period between 1991 and 1996 may be attributed to a severe drought that occurred between 1992 and 1995 (kelbe & germishuyse 2010:183; ndlovu & demlie 2020:8). rainfall data from the makatini research centre (figure 4) recorded an average annual precipitation of 471 mm between 1992 and 1994. this was lower than the 30-year average annual precipitation of 580 mm between 1988 and 2018. the increase in water bodies found between 1996 and 2001 is attributed to the higher rainfall the area received during the same period, where surface water bodies increased in area by approximately 12 ha year-1. however, the following decade recorded lower-than-average rainfall, which was a major contributor to the shrinkage of wetlands and lake level reduction reported for the study area (nsubuga, mearns & adeola 2018:1; smithers et al. 2017:490). climatic patterns investigated over the study period found that below-average rainfall has been experienced in the study area since 2001 (blamey et al. 2018:6). nsubuga et al. (2018) analysed rainfall data from eight stations in the mcp and reported a strong correlation (0.88) between precipitation and surface water levels. the study reported in this article shows that, the extent of surface water bodies decreased by about 444 ha year-1 between 2001 and 2006; this decrease was consistent with that reported in smithers et al. (2017:490) and nsubuga et al. (2018:1). implications on conservation areas this study showed that expansion of plantations and decrease in rainfall in the mcp have resulted in a decrease in the extent of surface water bodies and wetlands. the expansion of plantations is concentrated around the south–north axis of the recharge area of the isimangaliso wetland park. consequently, the extent of water bodies and wetlands in the conservation areas of the mcp decreased by 17% and 24%, respectively. because of the interconnected nature of the hydrologic system of the mcp, any stress on the groundwater systems will transmit automatically into surface water bodies and wetlands, including the conservation areas. this suggests that land-use planning and decisions outside of conservation areas and their impact on water resources disregard boundaries and may transgress on conservation areas. conclusion the north-eastern mcp has undergone significant change in land use over the last five decades, including afforestation, agriculture and rural settlements. to understand and map these land-use changes, the cloud-based gee platform was used to process and classify satellite data for the period from 1986 to 2019. the random forest classifier was used to classify a total of eight images for the study period. the overall accuracies of the classification achieved range from a minimum of 92.3% (2011) to a maximum of 96.3% (1996). the spatial expansion of plantations is clearly indicated, and an inverse relationship with surface water bodies and wetlands is noticed over the study period. the development of plantations was tracked between 1986 and 2019, which indicated that during the last decade, the rate of development has increased. the landsat classified images indicated that the expansion of plantation was concentrated in the area west of lake kosi bay and lake sibayi. over the same period, surface water bodies and wetlands decreased by 36.1% and 49.1%, respectively. as a result of the shrinkage in the areal extent of the lakes, the southern basin of lake sibayi was disconnected from the main basin of the lake in 2016, and the lake area continued to decrease. water bodies located around the middle of the study area have also been negatively affected, and some of them have disappeared in the 2019 classified map. similarly, the extent of wetlands decreased significantly, and a 47.9% reduction in area was noted between 2001 and 2019. the decrease in the extent of surface water systems, including lakes and wetlands, is not only attributed to land-use changes such as the expansion of plantation but also to climate changes in the form of below-average rainfall. this study confirms that the gee platform is an ideal tool for processing earth observation data at large spatial and long-time scales as it contains a multi-petabyte catalogue of satellite imagery and geospatial data sets with massive computational abilities, which allow users to efficiently determine long-term land-use changes for large areas. the results of the multi-temporal land-use and land cover information reported in this article will undoubtedly inform policyand decision-making related to the impacts of land-use change, including critical evaluation of water use license issues related to various scales of plantations and undertaking consistent compliance monitoring. acknowledgements the south african weather service is acknowledged for the meteorological data; dr. gokool of the university of kwazulu-natal, for assisting in the use of google earth engine; ezemvelo kwazulu-natal wildlife, for providing the land-use maps, and the south african national biodiversity institute, for providing the south african national land-cover map. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions m.r., m.d., m.l.t. and s.j.v.r. contributed to the design and implementation of the research, analysis of the results and writing of the manuscript. funding information the financial assistance of the expanded freshwater and terrestrial environmental observation network, hosted by the national research foundation’s south african environmental observation network business unit, and the south african department of science and innovation towards this research is acknowledged. data availability statement the data that support the findings of this study are available from the corresponding author (m.r.) upon reasonable request. disclaimer opinions expressed and conclusions arrived at in this article are those of the authors and are not necessarily to be attributed to the national research foundation or the south african environmental observation network. references adane, z.a., nasta, p., zlotnik, v. & wedin, d., 2018, ‘impact of grassland conversion to forest on groundwater recharge in the nebraska sand hills’, journal of hydrology: regional studies 15, 171–183. https://doi.org/10.1016/j.ejrh.2018.01.001 berhane, t.m., lane, c.r., wu, q., autrey, b.c., anenkhonov o.a., chepinoga v.v. & liu, h., 2018, ‘decision-tree, rule-based, and random forest classification of high-resolution multispectral imagery for wetland mapping and inventory’, remote sensing 10(580), 1–26. https://doi.org/10.3390/rs10040580 blamey, r.c., kolusu, s.r., mahlalela, p., todd, m.c. & reason, c.j.c., 2018, ‘the role of regional circulation features in regulating el niño climate impacts over southern africa: a comparison of the 2015/2016 drought with previous events’, international journal of climatology 38(11), 4276–4295. https://doi.org/10.1002/joc.5668 bonnesoeur, v., locatelli, b., guariguata, m.r., ochoa-tocachi, b.f., vanacker, v., mao, z., stokes, a. & mathez-stiefel, s., 2019, ‘impacts of forests and forestation on hydrological services in the andes: a systematic review’, forest ecology and management 433, 569–584. https://doi.org/10.1016/j.foreco.2018.11.033 bren, l. & hopmans, p., 2007, ‘paired catchments observations on the water yield of mature eucalypt and immature radiata pine plantations in victoria, australia’, journal of hydrology 336(3), 416–429. https://doi.org/10.1016/j.jhydrol.2007.01.018 chignell, s.m., luizza, w.m., skach, s., young, n.e. & evangelista, p.h., 2018, ‘an integrative modeling approach to mapping wetlands and riparian areas in a heterogeneous rocky mountain watershed’, remote sensing in ecology and conservation 4(2), 150–165. https://doi.org/10.1002/rse2.63 d’amato, d., rekola, m., wan, m. & cai, d., 2017, ‘effects of industrial plantations on ecosystem services and livelihoods: perspectives of rural communities in china’, land use policy 63, 266–278. https://doi.org/10.1016/j.landusepol.2017.01.044 department of water and sanitation (dws), 2018, daily evaporation data. dye, p., 1996, ‘climate, forest and streamflow relationships in south african afforested catchments’, commonwealth forestry association 75(1), 31–38. dye, p. & versfeld, d., 2007, ‘managing the hydrological impacts of south african plantation forests: an overview’, forest ecology and management 251(1–2), 121–128. https://doi.org/10.1016/j.foreco.2007.06.013 everson, c.s., scott-shaw, b.c., kelbe, b.e., starke, a., pearton, t., geldenhuys, c., vather, t. & maguire m., 2019, ‘quantifying the water-use of dominant land uses in the maputaland coastal plain’, in water research commission report no. tt 781/18. pretoria, south africa. google earth engine, 2018, meet earth engine, viewed 01 november 2019, from https://earthengine.google.com/. grundling, a.t., van den berg, e.c. & price, j.s., 2013, ‘assessing the distribution of wetlands over wet and dry periods and land-use change on the maputaland coastal plain, north eastern kwazulu-natal, south africa’, south african journal of geomatics 2(2), 1–20. jewitt, d., 2012, ‘land cover change in kwazulu-natal’, environment science and policy for sustainable development 10, 12–13. jin, y., liu, x., chen, y. & lang, x., 2018, ‘land-cover mapping using random forest classification and incorporating ndvi time-series and texture: a case study of central shandong’, international journal of remote sensing 39(23), 1–12. https://doi.org/10.1080/01431161.2018.1490976 kelbe, b. & germishuyse, t., 2010, ‘groundwater/surface water relationships with specific reference to maputaland’, in water research commission report no. 1168/1/10. pretoria, south africa. kienzle, s.w. & schulze, r.e., 1992, ‘a simulation model to assess the effect of afforestation on ground-water resources in deep sandy soils’, water sa 18(4), 265–272. lebourgeois, v., dupuy, s., vintrou, e. & ameline, m., 2017, ‘a combined random forest and obia classification scheme for mapping smallholder agriculture at different nomenclature levels using multisource data (simulated sentinel-2 time series, vhrs and dem)’, remote sensing 9(259), 1–21. https://doi.org/10.3390/rs9030259 li, c., wang, j., wang, l., hu, l. & gong, p., 2014, ‘comparison of classification algorithms and training sample sizes in urban land classification with landsat thematic mapper imagery’, remote sensing 6(2), 964–983. https://doi.org/10.3390/rs6020964 mellor, a., haywood, a., jones, a. & wilkes, p., 2012, ‘forest classification using random forests with multisource remote sensing and ancillary gis data’, in 16th australasian remote sensing and photogrammetry conference proceedings, january 2012, pp. 40–44, melbourne. mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, strelitzia 19, south african national biodiversity institute, pretoria. na, x., zhang, s., li, x., yu, h. & liu, c., 2010, ‘improved land cover mapping using random forests combined with landsat thematic mapper imagery and ancillary geographic data’, photogrammetric engineering & remote sensing 76(7), 833–84. https://doi.org/10.14358/pers.76.7.833 nasa, 2018, ‘advanced webinar: accuracy assessment of a landcover classification’, in remote sensing training program, february 13–20, 2018, pp. 1–20. national water act (nwa), 1998, act no. 36 of 1998, government printer, pretoria. ndlovu, m.s. & demlie, m., 2018, ‘statistical analysis of groundwater level variability across kwazulu-natal province, south africa’, journal of environmental earth science 77(739), 1–15. https://doi.org/10.1007/s12665-018-7929-x ndlovu, m.s. & demlie, m., 2020, ‘assessment of meteorological drought and wet conditions using two drought indices across kwazulu-natal province, south africa’, atmosphere 11(623), 1–20. https://doi.org/10.3390/atmos11060623 nsubuga, f.n.w., mearns, k.f. & adeola, a.m., 2017, ‘lake sibayi variations in response to climate variability in northern kwazulu-natal, south africa’, theoretical and applied climatology 137, 1233–1245. https://doi.org/10.1007/s00704-018-2640-0 ok, a., akar, o. & gungor, o., 2012, ‘evaluation of random forest method for agricultural crop classification’, european journal of remote sensing 45(3), 421–432. https://doi.org/10.5721/eujrs20124535 orimoloye, i.r., mazinyo, s.p., kalumba, a.m., nel, w., adigun, a.i. & ololade, o.o., 2019, ‘wetland shift monitoring using remote sensing and gis techniques: landscape dynamics and its implications on isimangaliso wetland park, south africa’, earth science informatics 12, 553–563. https://doi.org/10.1007/s12145-019-00400-4 pitman, w.v. & hutchison, i.p.g., 1975, ‘a preliminary hydrological study of lake sibaya’, in hydrological research unit, report 4/75, pp. 35. university of the witwatersrand, johannesburg. rawlins, b.k., 1991, ‘a geohydrological assessment of the behaviour and response of the zululand coastal plain to both environmental influences and human activity’, unpublished msc, university of zululand, richards bay. reynolds, j., wesson, k., desbiez, a.l.j., ochoa-quintero, j.m. & leimgruber, p., 2016, ‘using remote sensing and random forest to assess the conservation status of critical cerrado habitats in mato grosso do sul, brazil’, land 5(12), 1–12. https://doi.org/10.3390/land5020012 richards, j.a. & jia, x., 2006, remote sensing digital image analysis: an introduction, springer, heidelberg. rodriguez-galiano, v.f. & chica-rivas m., 2012, ‘evaluation of different machine learning methods for land cover mapping of a mediterranean area using multi-seasonal landsat images and digital terrain models’, international journal of digital earth 7(6), 492–509. https://doi.org/10.1080/17538947.2012.748848 sa forestry, 2012, sa’s biggest land reform forestry project, viewed 29 january 2020, from http://saforestryonline.co.za/articles/land_and_community/sas_biggest_land_reform_forestry_project/. sappi, 2020, sappi khulisa, viewed 09 august 2020, from https://www.sappi.com/sappi-khulisa. senf, c., hostert, p. & van der linden, s., 2012, ‘using modis time series and random forest classification for mapping land use in south-east asia’, in international geoscience and remote sensing symposium (igarss), munich. sikka, a.k., samra, j.s., sharda, v.n., samraj, p. & lakshmanan, v., 2003, ‘low flow and high flow responses to converting natural grassland into blue gum (eucalyptus globules) in nilgiri watersheds of south india’, journal of hydrology 270(1–2), 12–26. https://doi.org/10.1016/s0022-1694(02)00172-5 smith, j.h., wickham, j.d., stehman, s.v. & yang, l., 2002, ‘impacts of patch size and land cover heterogeneity on thematic image classification accuracy’, photogrammetric engineering and remote sensing 68(1), 65–70. smithers, j., gray, r., johnson, s. & still, d., 2017, ‘modelling and water yield assessment of lake sibhayi’, water sa 43(3), 480–491. https://doi.org/10.4314/wsa.v43i3.13 south african national biodiversity institute (sanbi), 2018, vegetation map of south africa, lesotho and swaziland, viewed 01 april 2020, from http://bgis.sanbi.org/spatialdataset/detail/669. south african weather service (saws), 2018, climate date, viewed 12 february 2019, from http://www.weathersa.co.za/climate. statistics south africa, 2011, department of statistics, viewed 30 july 2019, from http://www.statssa.gov.za/?page_id=993&id=the-msunduzi-municipality. thompson, m., 2018, ekznw 27/2017: updating the existing kzn provincial land cover map 2011 to 2017 conditions. united states geological survey (usgs), 2018, what is the landsat satellite program and why is it important? viewed 20 june 2019, from https://www.usgs.gov/faqs/what-landsat-satellite-program-and-why-it-important?qt-news_science_products=0#qt-news_science_products. weitz, j. & demlie, m., 2014, ‘conceptual modeling of groundwater-surface water interactions in the lake sibayi catchment, eastern south africa’, journal of earth science 99(2), 613–624. https://doi.org/10.1016/j.jafrearsci.2013.11.018 woodward, c., shumeister, j., larsen, j., jacobsen, g.e. & zawadzki, a., 2014, ‘the hydrological legacy of deforestation on global wetlands’, science 346(621), 844–847. https://doi.org/10.1126/science.1260510 yiu, t., 2019, understanding random forest, viewed 31 january 2020, from https://towardsdatascience.com/understandingrandom-forest-58381e0602d2. zurqania, h.a., posta, c.j., mikhailovaa, e.a., schlautmanb, m.a. & sharpc, j.l., 2018, ‘geospatial analysis of land use change in the savannah river basin using google earth engine’, international journal of applied earth observation geoinformation 69, 175–185. https://doi.org/10.1016/j.jag.2017.12.006 kopij.qxd the birds of sehlabathebe national park, lesotho g. kopij kopij, g. 2002. the birds of sehlabathebe national park, lesotho. koedoe 45(1): 65–78. pretoria. issn 0075-6458. a total of 117 bird species has been recorded in sehlabathebe national park, south-east lesotho, consisting of 29 vagrants, 18 visitors and 70 breeding and probable breeding residents. for each species status was determined and abundance roughly estimated. quantitative studies on breeding bird communities were carried out by means of the line transect method on four transects with the total length ca 30 km in the park and on two transects with the total length of ca 20 km outside the park. in the park, dominant species were represented by the stonechat saxicola torquata, ayres’ cisticola cisticola ayresii, yellow-rumped widow euplectes capensis and wailing cisticola cisticola lais. outside the park dominants were represented by cape weaver ploceus capensis, cape sparrow passer melanurus, cape canary serinus canicollis, common quail coturnix coturnix, stonechat, cape butning emberiza capensis and drakensberg siskin pseudochloroptila symonsi. characteristic, high-altitude species in the park included drakensberg siskin, mountain pipit anthus hoeschi, orange-breasted rockjumper chaetops auriantius, banded martin riparia cincta and sentinel rock thrush monticola explorator. species such as the laughing dove streptopelia senegalensis, sicklewinged chat cercomela sinuata, mountain chat oenanthe monticola, thick-billed lark galerida magnirostris, red-winged starling onychognathus morio, alpine swift apus melba, cape sparrow, grey-headed sparrow passer diffusus, red bishop euplectes orix and golden bishop euplectes afer were absent or occurred in very low densities in the park, although they are widespread and common in the maluti/drakensberg grasslands (including areas neighbouring to the park). the lack of trees and shrubs for nesting, the lack of cultivated fields as feeding places and competition with related species both for food and nesting sites, may partly play a role in this regard. key words: sehlabathebe national park, lesotho, birds, quantitative studies, conservation. g. kopij, department of biology, national university of lesotho, p. o. roma 180, lesotho. issn 0075-6458 65 koedoe 45/1 (2002) introduction sehlabathebe national park (snp) was proclaimed on 27 february 1970 and to date it is the only national park in lesotho (pomela et al. 2000). plant diversity of snp has been studied by kopij (2001b), and animal diversity by lynch & watson (1990) and kopij (2001b). the first ornithological observations in snp were made during the years 1975–1982 by j. belcomb, c. brown, r. howard, m. howard and l. passineau. from 1987 to 1990, a total of 16 visits to record birds of snp were conducted. each visit lasted 1–4 days (11 visits in summer (november-february) and five in winter (april-september)). the following persons participated in this project: l. p. abbott, e. bolding, j. cambridge, k. e. deas, s. fairman, j. m. gosnell, r. guy, s. haines, j. jones, p. osborne, d. p. smits, b. tigar, s. van der veur, j. p. watson, h. wilson, a. wolfromm and h. wrayner. these observers used the standard southern african bird atlas project cards for reporting their records and also provided details of unusual records. to date, results of these observations are available mainly in the form of unpublished reports (howard 1977; howard & howard 1977; belcomb 1980). although a booklet on birds of the park was compiled by passineau (1977), this is only a popular account of more common species, together with a simple, not kopij.qxd 2005/12/09 11:32 page 65 annotated and not critically evaluated checklist of all species. the present study aims to determine and report on the status (based on my own and all previous records) and abundance (based on my own studies) of bird species in snp and the surrounding area. to date, quantitative studies of bird communities in lesotho have only been conducted in the lowlands: in urbanised areas of maseru (kopij 2000) and in rural areas and in farmlands in the roma valley (kopij 2001a). study area located in the qacha’s nek district of southeastern lesotho, snp (7239 ha) was established in 1970. ranging in elevation from 2200 m to 2600 m, it falls entirely within the themeda-festuca alpine grassland. snp comprises short grasses (69 %), escarpments and colluvial lands (13 %), wetlands (11 %), water-courses (4 %) and roads and other man-modified areas (3 %). except for a few small isolated willow salix subserrata clumps, no other woody vegetation occurs in the park. water in the form of streams, rivers and pools is abundant. mean annual rainfall is 769 mm, with most rains (75 %) in november-march. mean annual maximum and minimum temperatures are 19.6 ºc and 2.2 ºc respectively (kopij 2001b). a map of snp was published by lynch & watson (1990). methods quantitative studies quantitative studies were carried out from 12 to 15 january 2001. the american version of the line transect method (bibby et al. 1992) was employed to assess density and dominance of breeding species. the observer walked slowly, counting his paces and stopping often to observe and record birds. only the birds in front and on either side of the observer were recorded—those behind were not. where the transect ended, birds beyond the end point were not counted. distant birds only detectable using binocular were not searched for. a pair, not an individual, was the censused unit. a singing or otherwise observed male, in a habitat suitable for breeding, was interpreted as one pair. if a male was not observed, a female, a group of fledglings or an occupied nest was also interpreted as one pair (koskimies & vaisanen 1991; bibby et al. 1992). density estimates are based on morning counts only, as birds were markedly less active in the afternoon. for calculation of relative abundance (dominance), morning and afternoon counts were pooled. for comparison of the breeding bird communities in the park with that in man-modified neighbouring areas, two transects were also undertaken outside the park. morning counts covered four transects: along the road from the research station to the lodge (9 km); grassland with numerous stream valleys at the foot of thaba ntšo; surveyed from 6:00 to 11:00 on 12 january 2001; from the research station along the north-west border of the park through koti-se-seholo tsoelikane to tsoelikane falls (7 km); mainly short grassland with a few stream valleys and sandstone formations; surveyed from 6:00 to 10:00 on 13 january 2001; along the tsoelikane river from letamong village to the waterfall (6 km); extensive marshland with fairly tall and dense grass and sedges; surveyed from 10:00 to 12:30 on 13 january 2001; from the research station through the north-western corner of the park, herdboy and central valleys to the waterfall (9 km); grassland with sandstone formations and stream valleys; surveyed from 6:00 to 11:00on 15 january. afternoon counts were conducted along three transects: from the lodge to the research station; short grass with numerous stream valleys, ca 0.5 km to 2 km south of the main road; surveyed from 12:00 to 16:00 on 12 january 2001; from the waterfall along romoluma and moseha to the research station; short grass with numerous sandstone formations; surveyed from 13:00 to 16:00 on 13 january 2001; from the waterfall to the lodge along tsoelikane valley (marshland dominated by merxmuelleria spp. with seven rock pools vegetated with typha capensis, aponogeton ranunculiflorus and other water plants); from the lodge to the research station through short grass with stream valleys south of the main road; surveyed from 12:00 to 16:00 on 15 january 2001. outside the park counts were conducted on two transects: along the road between the main gate to the park and the crossroad near the roman catholic church in sehlabathebe (ca 10 km); pastures, two villages, arable fields of wheat, sorghum and peas; surveyed from 9:45 to 12:15 on koedoe 45/1 (2002) 66 issn 0075-6458 kopij.qxd 2005/12/09 11:32 page 66 14 january 2001; along the leqooa valley upstream from the bridge between ha mavuka and ha semenyane to the confluence of leqooa river and thamathu river near ha soloja; salix trees grow along the river banks; the valley is partly cultivated (mainly with wheat) and marshland is utilised as pasture for cattle and horses; surveyed from 12:45 to 15:45 on 15 january 2001. dominant species are those represented by more than 5.0 %, and subdominant species by 2–4.9 % of all breeding pairs recorded. annotated checklist the checklist is based on results of all unpublished records by howard (1977), howard & howard (1977), passineau (1977) and balcomb (1980) and those of g. kopij (this study) the following criteria were used to determine the status and abundance of each species: status: vagrant—single records or once in a five-year period; visitor – not resident, and nonbreeding, but recorded regularly in each year, often resident in neighbouring areas; resident – direct breeding evidence (nest with eggs or chicks, egg shells); probable resident – indirect breeding evidence (occupied territory, simultaneously singing males in optimal habitats, breeding display etc.); palearctic migrant – breeds in the palearctic region during the northern summer, wintering in snp during the austral summer; intra-african migrant – breeds in snp, but winters in tropical africa; altitudinal migrant – breeding in snp during the austral summer, but wintering outside the park, at lower altitudes; nomad – appears erratically, at no particular season, sometimes in large numbers that may breed. abundance: very common—more than 200 breeding pairs (b.p.) in the entire park; common: 70–200 b.p.; fairy common – 20–69 b.p.; uncommon: 6–19 b.p.; rare – <5 breeding pairs. abundance has been roughly estimated through extrapolations from transect counts and from observations. results of quantitative studies among 1120 potential breeding pairs represented by 51 species recorded during 12 to 15 january 2001 in sehlabathebe national park, four species were classified as dominant: stonechat, ayres’ cisticola, yellowissn 0075-6458 67 koedoe 45/1 (2002) table 1 relative abundance (dominance) of birds in afroalpine grassland in sehlabathebe national park in january 2001. dominant species are given in bold case. data from all transects is pooled species pairs dominance (%) stonechat 127 11.3 ayres’ cisticola 125 11.2 yellow-rumped widow 95 8.5 wailing cisticola 77 6.9 drakensberg siskin 54 4.8 mountain pipit 54 4.8 long-tailed widow 50 4.5 levaillant’s cisticola 46 4.1 orange-throated longclaw 44 3.9 common quail 36 3.2 black swift 34 3.0 cape bunting 33 2.9 sentinel rock thrush 32 2.9 cape wagtail 31 2.8 cape canary 29 2.6 orange-breasted rockjumper 26 2.3 ground woodpecker 24 2.1 greater striped swallow 24 2.1 quail finch 22 2.0 cape weaver 21 1.9 banded martin 19 1.7 malachite sunbird 13 1.2 jackal buzzard 13 1.2 cape robin 9 0.8 white-breasted swallow 8 0.7 rock pigeon 6 0.5 red-collared widow 5 0.4 grey-wing francolin 5 0.4 horus swift 5 0.4 yellow-breasted pipit 5 0.4 rock kestrel 5 0.4 black crow 5 0.4 rock martin 4 0.4 black duck 3 0.3 hadeda ibis 3 0.3 yellow-billed duck 3 0.3 red-winged starling 3 0.3 cape sparrow 3 0.3 pied starling 3 0.3 common waxbill 3 0.3 cape turtle dove 2 0.2 bearded vulture 1 0.1 african marsh harrier 1 0.1 black harrier 1 0.1 lanner falcon 1 0.1 pied kingfisher 1 0.1 cape eagle owl 1 0.1 rock pipit 1 0.1 spotted prinia 1 0.1 grey-headed sparrow 1 0.1 yellow canary 1 0.1 mountain chat 1 0.1 total 1120 100 kopij.qxd 2005/12/09 11:32 page 67 koedoe 45/1 (2002) 68 issn 0075-6458 table 2 breeding bird community of afro-alpine grassland in sehlabathebe national park in january 2001. dominant species are given in bold case species morning transects 1 2 3 4 pairs p./10 km dom.(%) ayres’ cisticola 6 22 19 39 86 26.9 11.7 stonechat 19 18 23 20 80 25.0 10.9 long-tailed widow 4 13 14 12 43 13.4 5.8 yellow-rumped widow 17 15 24 15 71 22.2 9.6 wailing cisticola 17 11 4 10 42 13.1 5.7 levaillant’s cisticola 2 3 24 8 37 11.6 5.0 drakensberg siskin 13 7 2 8 30 9.4 4.1 mountain pipit 4 15 10 29 9.1 3.9 orange-throated longclaw 5 5 5 12 27 8.4 3.7 common quail 5 11 10 26 8.1 3.5 black swift 4 5 16 25 7.8 3.4 cape bunting 8 9 2 5 24 7.5 3.3 cape wagtail 3 5 12 1 21 6.6 2.9 sentinel rock thrush 3 6 1 7 17 5.3 2.3 banded martin 1 4 10 2 17 5.3 2.3 cape weaver 7 8 1 1 17 5.3 2.3 ground woodpecker 4 5 2 4 15 4.7 2.0 cape canary 12 3 15 4.7 2.0 greater striped swallow 3 2 3 3 11 3.4 1.5 orange-breasted rockjumper 2 1 2 6 11 3.4 1.5 quail finch 1 4 6 11 3.4 1.5 jackal buzzard 1 4 3 1 9 2.8 1.2 white-breasted swallow 4 1 2 7 2.2 1.0 cape robin 3 2 5 1.6 0.7 malachite sunbird 3 1 1 5 1.6 0.7 red-collared widow 3 1 1 5 1.6 0.7 grey-wing francolin 1 3 4 1.3 0.5 white-rumped swift 4 4 1.3 0.5 yellow-breasted pipit 1 1 1 1 4 1.3 0.5 black duck 2 1 3 0.9 0.4 rock pigeon 1 2 3 0.9 0.4 rock martin 1 1 1 3 0.9 0.4 hadeda ibis 1 1 2 0.6 0.3 yellow-billed duck 1 1 2 0.6 0.3 rock kestrel 1 1 2 0.6 0.3 black crow 1 1 2 0.6 0.3 red-winged starling 1 1 2 0.6 0.3 cape sparrow 1 1 2 0.6 0.3 bearded vulture 1 1 0.3 0.1 african marsh harrier 1 1 0.3 0.1 black harrier 1 1 0.3 0.1 lanner falcon 1 1 0.3 0.1 cape turtle dove 1 1 0.3 0.1 pied kingfisher 1 1 0.3 0.1 cape eagle owl 1 1 0.3 0.1 rock pipit 1 1 0.3 0.1 spotted prinia 1 1 0.3 0.1 pied starling 1 1 0.3 0.1 common waxbill 1 1 0.3 0.1 grey-headed sparrow 1 1 0.3 0.1 yellow canary 1 1 0.3 0.1 total 736 kopij.qxd 2005/12/09 11:32 page 68 rumped widow and wailing cisticola (table 1). together they comprised 37.9 % of all breeding pairs recorded. subdominants, represented by 15 species (table 1), constituted another 48 %. drakensberg siskin, mountain pipit, orange-breasted rockjumper, banded martin and sentinel rock thrush can be classified as characteristic species. six species (ayres’ cisticola, stonechat, long-tailed widow, yellow-rumped widow, wailing cisticola and levaillant’s cisticola) exceeded the density of 10 pairs/100 ha, 23 species nested in a density ranging from 1 to 10 pairs/10 km. the remaining species nested in a density lower than 1 pair/10 km (table 2). issn 0075-6458 69 koedoe 45/1 (2002) table 3 breeding bird communities in farmland bordering sehlabathebe national park in january 2001. dominant species are given in bold case species transect 1 transect 2 pairs dom. (%) pairs dom. (%) cape weaver 2 1.6 25 14.0 cape sparrow 10 7.9 13 7.3 cape canary 8 6.3 13 7.3 common quail 16 12.6 4 2.2 stonechat 7 5.5 13 7.3 cape bunting 10 7.9 8 4.5 drakensberg siskin 7 5.5 9 5.1 cape wagtail 4 3.1 12 6.7 mountain pipit 12 9.4 1 0.6 ayres’ cisticola 6 4.7 7 3.9 yellow-rumped widow 4 3.1 8 4.5 common waxbill 4 3.1 5 2.8 long-tailed widow 5 3.9 3 1.7 banded martin 8 4.5 black-throated martin 7 3.9 red-collared widow 2 1.6 4 2.2 greater striped swallow 6 3.4 wailing cisticola 4 3.1 1 0.6 sentinel rock thrush 3 2.4 1 0.6 pied starling 4 3.1 white-breasted swallow 4 2.2 malachite sunbird 1 0.8 3 1.7 sentinel rock thrush 3 2.4 1 0.6 pin-tailed whydah 2 1.6 1 0.6 cape robin 1 0.8 2 1.1 rock martin 3 1.7 black duck 2 1.1 jackal buzzard 1 0.8 1 0.6 horus swift 1 0.8 1 0.6 rock pigeon 2 1.1 quail finch 2 1.6 house sparrow 2 1.6 grey-headed sparrow 2 1.6 black swift 2 1.1 red bishop 2 1.1 black crow 1 0.8 hadeda ibis 1 0.8 hamerkop 1 0.6 rock kestrel 1 0.6 cape turtle dove 1 0.8 ground woodpecker 1 0.6 thick-billed lark 1 0.8 mountain chat 1 0.6 red-winged starling 1 0.6 levaillant’s cisticola 1 0.6 total 127 100.0 178 100.2 kopij.qxd 2005/12/09 11:32 page 69 koedoe 45/1 (2002) 70 issn 0075-6458 dabchick grey heron black-headed heron hamerkop black stork white stork sacred ibis bald ibis hadeda ibis spur-winged goose egyptian goose black duck yellow-billed duck secretarybird cape vulture lanner falcon rock kestrel yellow-billed kite black eagle bearded vulture jackal buzzard steppe buzzard african marsh harrier black harrier gymnogene grey-winged francolin quail red-knobbed coot crowned crane wattled crane blue crane ethiopian snipe greenshank wood sandpiper rock pigeon cape turtle dove barn owl spotted eagle owl black swift white-rumped swift alpine swift giant kingfisher ground woodpecker table 4 temporal distribution of birds in sehlabathebe national park species j f m a m j j a s o n d kopij.qxd 2005/12/09 11:32 page 70 issn 0075-6458 71 koedoe 45/1 (2002) european swallow white-throated swallow greater striped swallow rock martin african sand martin banded martin black crow white-necked raven rock jumper cape rock thrush sentinel rock thrush mountain chat stonechat cape robin ayres’ cisticola wailing cisticola levaillant’s cisticola spotted prinia spotted flycatcher cape wagtail mountain pipit yellow-breasted pipit orange-thr. longclaw red-winged starling pied starling malachite sunbird house sparrow cape sparrow cape weaver red bishop cape widow red-collared widow long-tailed widow quail finch pin-tailed whydah drakensberg siskin cape canary yellow canary cape bunting species j f m a m j j a s o n d table 4 (continued) kopij.qxd 2005/12/09 11:32 page 71 in areas bordering snp, the dominant species were quite different. in farmland the group was composed of the common quail, mountain pipit, cape sparrow, cape bunting, cape canary, stonechat and drakensberg siskin, while in tsoelikane valley the dominants were the cape weaver, cape sparrow, cape canary, stonechat, cape wagtail and drakensberg siskin (table 3). also, composition of the group of subdominant species outside the park (table 3) differed markedly from that inside the park (table 2). annotated checklist nomenclature, systematics and species numbers follow maclean (1993). 6. podiceps cristatus great crested grebe vagrant. recorded in january 1980 (j. balcomb). this is the only lesotho record of the species during the past 50 years. 8. tachybaptus ruficollis dabchick vagrant. 55. phalacrocorax carbo white-breasted cormorant vagrant. recorded in january 1988 (s. fairman). 62. ardea cinerea grey heron visitor. probably resident in neighbouring areas. on 23 december 1976, one aberrant individual with red bill and legs was observed (r. howard). 63. ardea melanocephala black-headed heron visitor. resident in neighbouring areas. 71. bubulcus ibis cattle egret vagrant. 76. nycticorax nycticorax black-crowned night heron vagrant. 81. scopus umbretta hamerkop visitor. resident in neighbouring areas. 83. ciconia ciconia white stork palearctic migrant (table 4). most birds forage in the tsoelikane valley below the waterfall, e.g. from 12 to 15 january 2001, a flock of 35 birds were observed there, and the whole population in the park was estimated at 40–50 pairs. 84. ciconia nigra black stork visitor. resident in neighbouring areas. 91. threskiornis aethiopicus sacred ibis vagrant. 92. geronticus calvus bald ibis visitor. resident in neighbouring areas. 94. bostrychia hagedash hadeda ibis visitor. breeding in neighbouring areas. 95. platalea alba african spoonbill vagrant. 102. alopochen aegyptiacus egyptian goose visitor. probably resident in neighbouring areas. 104. anas undulata yellow-billed duck probable resident on the tsoelikane river; uncommon. 105. anas sparsa african black duck resident on the tsoelikane river; uncommon. 108. anas erythrorhyncha red-billed teal vagrant. recorded in january 1980 (j. balcomb), 01 october 1989 (j. tigar & p. osborne). 116. plectropterus gambensis spur-winged goose visitor. probably resident in neighbouring areas. 118. sagittarius serpentarius secretarybird visitor. probably resident in neighbouring areas. 119. gypaetus barbatus bearded vulture koedoe 45/1 (2002) 72 issn 0075-6458 kopij.qxd 2005/12/09 11:32 page 72 resident at thaba ntšo, where one pair nests. 122. gyps coprotheres cape vulture visitor. resident in neighbouring areas. until late 1980’s there was a breeding colony of 100–200 individuals in snp (barnes 1998). 126. milvus migrans yellow-billed kite vagrant. recorded on 01 september 1985 (h. wilson). 127. elanus caeruleus black-shouldered kite vagrant. recorded in february 1990 (r. guy) and november 1990 (j. jones). 131. aquila verreauxii black eagle vagrant. recorded in january 1977 (r. howard). during the years 1986–89 a pair was recorded; probably bred at thaba ntšo (osborne & tigar 1990). 149. buteo buteo steppe buzzard palearctic migrant; uncommon. 152 buteo rufofuscus jackal buzzard uncommon resident. 162. melierax canorus pale chanting goshawk vagrant. single birds were recorded in april 1976 and december 1979 (j. belcomb). 165. circus ranivorus african marsh harrier probable altitudinal migrant (table 4). during the years 1975–80, only one record (28 october 1976; j. balcomb). on 15 january 2001 an adult with two juveniles was observed 3–4 km south of the lodge (pers. obs.). 168. circus maurus black harrier probable altitudinal migrant (table 4). single pairs probably nested, at least during 1978/79 (j. balcomb), 1980/81 (c. brown), 1989/90 (r. guy) and 2000/01 breeding seasons (pers. obs.). 169. polyboroides typus gymnogene vagrant. 172. falco biarmicus lanner falcon rare resident. 181. falco tinnunculus rock kestrel uncommon resident. 190. francolinus africanus grey-wing francolin fairly common resident in grassy areas. 200. coturnix coturnix common quail common altitudinal migrant (tables 4 & 5). 203. numida meleagris helmeted guineafowl vagrant. 207. bugeranus carunculatus wattled crane probably 1–2 breeding pairs. a pair, 5 january 1977 (r. howard). january, april, december 1976; march 1980 (j. balcomb); 3–4 january 1976, december 1982 (d. cochlan). no records since 1983. snp is the only locality in lesotho where the species has been recorded. 208. anthropoides paradiseus blue crane probably 1–2 breeding pairs. a pair, 28 december 1976 (r. howard). september, 1975, october, december 1976 (j. belcomb), 3 january 1976 (d. cochlan). not recorded after 1976. issn 0075-6458 73 koedoe 45/1 (2002) table 5 habitat selection by common quails (number of breeding pairs) habitat snp neighbouring area grasses 14 5 sedges 5 grasses + sedges 4 marsh 3 wheat 3 peas 2 kopij.qxd 2005/12/09 11:32 page 73 209. balearica regulorum crowned crane vagrant. probably single pairs bred irregularly in summer 1975/76 (j. balcomb). snp is the only locality in lesotho where the species has been recorded. 211. crex crex corncrake vagrant. recorded on 3 january 1976 (bonde 1990). 228. fulica cristata red-knobbed coot visitor. 231. neotis denhami stanley’s bustard vagrant. recorded in february and march 1979 (j. balcomb). 264. actitis hypoleucos common sandpiper palearctic migrant (table 4). 266. tringa glareola wood sandpiper vagrant. two birds were recorded on 6 january 1977. 269. tringa stagnatilis marsh sandpiper vagrant. recorded in november 1990 (s. fairman, j. cambridge). 270. tringa nebularia greenshank palearctic migrant (table 4). 274. calidris minuta little stint palearctic migrant (table 4). recorded on 8 january 1977. 284. philomachus pugnax ruff palearctic migrant (table 4). recorded on 29 december 1976 (d. cochlan). 286. gallinago nigripennis ethiopian snipe probable altitudinal migrant (table 4). recorded in march 1977, january, february, june 1980 (j. balcomb), november 1990. 297. burhinus capensis spotted dikkop vagrant. recorded in 5 january 1977. 349. columba guinea rock pigeon fairly common resident, nesting on sandstone cliffs. 354. streptopelia capicola cape turtle dove rare resident in larger willow clumps. 382. clamator jacobinus jacobin cuckoo vagrant. recorded in november 1978 (j. belcomb). 392. tyto alba barn owl visitor. resident in sehlabathebe village (osborne & tigar 1990). 400. bubo capensis cape eagle owl probable resident, 1–2 pairs on thaba ntšo. 401. bubo africanus spotted eagle owl probable resident, 1–2 pairs on thaba ntšo. 412. apus barbatus black swift common intra-african migrant. nesting colonies were localised mainly around thaba ntšo and on sandstone cliffs around the waterfall. 415. apus caffer white-rumped swift fairly common intra-african migrant (table 4). 416. apus horus horus swift uncommon intra-african migrant (table 4), nesting along sandstone cliffs. 418. apus melba alpine swift visitor. intra-african migrant to neighbouring areas (table 4). 428. ceryle rudis pied kingfisher probable resident on the tsoelikane river below the waterfall. not recorded since 1976 (r. howard). 429. megaceryle maxima giant kingfisher probable resident on the tsoelikane river. 431. alcedo cristata malachite kingfisher probable resident on the tsoelikane river. 480. geocolaptes olivaceus ground woodpecker koedoe 45/1 (2002) 74 issn 0075-6458 kopij.qxd 2005/12/09 11:32 page 74 common resident. breeds in the banks of rivers and streams and in erosion gulleys. 500. mirafra curvirostris long-billed lark probable resident; rare. 507. calandrella cinerea red-capped lark probably rare nomadic. 512. galerida magnirostris thick-billed lark rare resident. 518. hirundo rustica european swallow palearctic migrant (table 4). 520. hirundo albigularis white-throated swallow intra-african migrant (table 4); uncommon. 526. hirundo cucullata greater striped swallow intra-african migrant (table 4). present along sandstone cliffs and on thaba ntšo. 529. hirundo fuligula rock martin fairly common altitudinal migrant (table 4); nesting on sandstone cliffs. 530. delichon urbica house martin palearctic migrant (table 4). recorded in february 1990 (r. guy). 533. riparia paludicola brown-throated martin fairly common resident, nesting mainly along tsoelikane river. 534. riparia cincta banded martin fairly common intra-african migrant, nesting mainly along tsoelikane river. 547. corvus capensis black crow resident; 2–5 pairs on larger sandstone cliffs and at thaba ntšo. 548. corvus albus pied crow resident; 1–2 pairs at thaba ntšo. 550. corvus albicollis white-necked raven probably resident. one probable breeding pair at thaba ntšo. 551. parus afer southern grey tit vagrant. recorded on december 1988 (s. fairman). 581. monticola rupestris cape rock thrush rare resident; nesting along sandstone cliffs. 582. monticola explorator sentinel rock thrush common resident in short grasses with rocky outcrops or along rocky cliffs. 586. oenanthe monticola mountain chat probable rare resident. 589. cercomela familiaris familiar chat probable rare resident. 591. cercomela sinuata sickle-winged chat visitor. breeding in neighbouring areas. 596. saxicola torquata stonechat most birds are altitudinal migrants, some are resident. very common in grassland, marshes and along streams and valleys during the breeding season. breeds mainly in december/january, e.g. on 12–15 january 2001, most pairs were seen with 2–3 juveniles. 601. cossypha caffra cape robin fairly common resident in willow clumps. 612. chaetops aurantius orange-breasted rockjumper fairly common resident; nesting along sandstone formations. 667. cisticola ayresii ayres’ cisticola very common altitudinal migrant; breeding mainly in short grass. 670. cisticola lais wailing cisticola very common resident, mainly in shorter grasses on sloping hillsides. issn 0075-6458 75 koedoe 45/1 (2002) kopij.qxd 2005/12/09 11:32 page 75 677. cisticola tinniens levaillant’s cisticola very common resident in marshes and longer grasses, especially along water courses. 686. prinia hypoxantha spotted prinia rare resident on lower slopes of thaba ntšo. 654. muscicapa striata spotted flycatcher palearctic migrant. recorded on december 1977 and january 1980 (j. belcomb). 698. sigelus silens fiscal flycatcher vagrant. 713. motacilla capensis cape wagtail common resident along water courses, at rock pools and dams. 701. anthus hoeschi mountain pipit very common intra-african migrant, nesting mainly in short grass. 721. anthus crenatus rock pipit probable resident on lower rocky slopes of thaba ntšo. 725. hemimacronyx chloris yellow-breasted pipit uncommon altitudinal migrant (table 4); breeding in short grass. 727. macronyx capensis orange-throated longclaw common resident in marshes and in grass, especially along streams and river valleys. 732. lanius collaris fiscal shrike visitor. resident in neighbouring areas. 759. spreo bicolor pied starling uncommon resident on lower slopes of thaba ntšo. 769. onychognathus morio red-winged starling altitudinal migrant (table 4); nesting along sandstone cliffs; uncommon. december-april. 775. nectarinia famosa malachite sunbird fairly common altitudinal migrant (table 4), usually close to sandstone formations and kniphofia and aloe communities. 801. passer domesticus house sparrow visitor, resident at sehlabathebe village. 803. passer melanurus cape sparrow uncommon resident. nesting recorded in old cape weaver nests in willows. 804. passer diffusus grey-headed sparrow rare resident recorded in willow clumps. 813. ploceus capensis cape weaver common altitudinal migrant (table 4). nests were located exclusively in willows, both in snp and in the neighbouring area. outside the park, on the banks of the tsoelikane river, most nests were located 1–2 m above the water level. the number of nests per group ranged from 3 to 20. 824. euplectes orix red bishop probable altitudinal migrant (table 4); rare. breeding was recorded in old cape weaver nests in willows on banks of the tsoelikane river outside snp. this nest site is unusual for this species. 826. euplectes afer golden bishop probably altitudinal migrant (table 4); rare. 827. euplectes capensis yellow-rumped widow very common altitudinal migrant (table 4); breeding in marshes and in grass along water-courses and at dams. 831. euplectes ardens red-collared widow fairly common altitudinal migrant (table 4); breeding usually in marshes. 832. euplectes progne long-tailed widow koedoe 45/1 (2002) 76 issn 0075-6458 kopij.qxd 2005/12/09 11:32 page 76 very common resident in marshes and in grass along water-courses and at dams. 846. estrilda astrild common waxbill fairly common resident, mainly along the tsoelikane river. 852. ortygospiza atricollis quail finch fairly common altitudinal migrant (table 4); breeding in grass, usually in close proximity of pools and dams. 860. vidua macroura pin-tailed whydah uncommon altitudinal migrant (table 4). 872. serinus canicollis cape canary common altitudinal migrant (table 4); breeding in grassy areas. 875. pseudochloroptila symonsi drakensberg siskin very common resident; breeding in grassy areas and marshes. feeds mainly on seeds of helichrysum spp. and polygonum spp. a nest with one egg (20.5 x 16.5 mm) was found on 13 january 2001, located in short grass mixed with numerous helichrysum flowers. 878. serinus flaviventris yellow canary probable altitudinal migrant (table 4); rare. 885. emberiza capensis cape bunting very common resident, usually along sandstone formations. 886. emberiza tahapisi rock bunting vagrant. recorded on 1 september 1985 (h. wilson), 13 march 1987 (c. brown). the following species need confirmation before their inclusion into the list: little swift apus affinis, whinchat saxicola rubetra white-winged widow euplectes albonotatus and bully canary serinus sulphuratus. all these species were listed by passineau (1977), but without corroborating evidence. they are regarded as very rare in lesotho highlands (osborne & tigar 1990, bonde 1993) and can be confused with congeneric species. the records may be considered, therefore, to derive from misidentification. conclusions to date 117 bird species have been recorded in the snp, 29 of them are vagrants, 18 are visitors, and 70 are breeding or probable breeding residents. dominant species are represented by stonechat, ayres’ cisticola, yellow-rumped widow and wailing cisticola. there are 15 subdominant species. characteristic species include drakensberg siskin, mountain pipit, orange-breasted rockjumper, banded martin and sentinel rock thrush. it is important to note the absence or very low densities of the following species in snp, which are widespread and common in the maloti/drakensberg grasslands (including areas neighbouring to snp): laughing dove streptopelia senegalensis, sicklewinged chat, mountain chat, thick-billed lark, red-winged starling, alpine swift, cape sparrow, grey-headed sparrow, red bishop and golden bishop (cf. osborne & tigar 1990, little & bainbridge 1992, bonde 1993). the lack of trees and shrubs for nesting, the lack of cultivated fields as feeding places and competition with related species both for food and nesting sites, may partly play a role in this regard. acknowledgements this study was sponsored by the rcc committee of the national university of lesotho. dr d. n. johnson, prof. o. b. kok and mr r. j. nuttall are thanked for reviewing the paper. my thanks are also due to prof d. ambrose for indicating some unpublished reports. i also thank the administration of the snp for providing me accommodation during this study. issn 0075-6458 77 koedoe 45/1 (2002) kopij.qxd 2005/12/09 11:32 page 77 references balcomb, j. 1980. birds sighted in sehlabathebe national park, 1975–1980. unpublished internal report. barnes, k. n. 1988. important bird areas of lesotho. pp. 281–294. in: barnes, k. n. (ed.). the important bird areas of southern africa. johannesburg: birdlife south africa. bibby, c.l., n.d. burgess & d.a.hill. 1992. bird census techniques. london: academic press. bonde, k. 1993. birds of lesotho. a guide to distribution past and present. pietermaritzburg: university of natal press. brown, c.j. & p.r. barnes. 1984. birds of the natal alpine belt. lammergeyer 33: 1–13. howard, r. 1977. lesotho bird list. unpublished internal report. howard, r. & m. howard. 1977. report on ornithological work, [sehlabathebe national park], january 2 to 9, 1977. unpublished report. koskimies, p. & r.a. väisänen. 1991. monitoring bird populations. helsinki: zoological museum, finnish museum of natural history. kopij, g. 2000. birds of maseru. nul journal of research 8: 104–151. kopij, g. 2001a. birds of roma valley, lesotho. roma: national university of lesotho. kopij, g. 2001b. sehlabathebe national park. roma: national university of lesotho. little, r.m. & w.r. bainbridge. 1992. birds of the natal drakensberg parks. pietermaritzburg: wildlife society. lynch, c.d. & j.p. watson. 1990. the mammals of sehlabathebe national park, lesotho. navorsinge van die nasionale museum, bloemfontein 6: 523–554. osborne, p.e. & b.j. tigar. 1990. the status and distribution of birds in lesotho. unpublished report. passineau, l. 1977. birds of sehlabathebe national park. maseru: lesotho national parks. pomela, e.m., c. mokuku, s. ‘mabataung, r. phororo, m. mokoae, m. majoro, d. may, d. ambrose & s. talukdar. 2000. biological diversity in lesotho: a country study. maseru: lesotho government, national environmental secretariat. koedoe 45/1 (2002) 78 issn 0075-6458 kopij.qxd 2005/12/09 11:32 page 78 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <feff00560065007200770065006e00640065006e0020005300690065002000640069006500730065002000450069006e007300740065006c006c0075006e00670065006e0020007a0075006d002000450072007300740065006c006c0065006e00200076006f006e0020005000440046002d0044006f006b0075006d0065006e00740065006e0020006d00690074002000650069006e006500720020006800f60068006500720065006e002000420069006c0064006100750066006c00f600730075006e0067002c00200075006d002000650069006e00650020007100750061006c00690074006100740069007600200068006f006300680077006500720074006900670065002000410075007300670061006200650020006600fc0072002000640069006500200044007200750063006b0076006f0072007300740075006600650020007a0075002000650072007a00690065006c0065006e002e00200044006900650020005000440046002d0044006f006b0075006d0065006e007400650020006b00f6006e006e0065006e0020006d006900740020004100630072006f0062006100740020006f0064006500720020006d00690074002000640065006d002000520065006100640065007200200035002e003000200075006e00640020006800f600680065007200200067006500f600660066006e00650074002000770065007200640065006e002e00200042006500690020006400690065007300650072002000450069006e007300740065006c006c0075006e00670020006900730074002000650069006e00650020005300630068007200690066007400650069006e00620065007400740075006e00670020006500720066006f0072006400650072006c006900630068002e> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <feff004e00e4006900640065006e002000610073006500740075007300740065006e0020006100760075006c006c006100200076006f0069006400610061006e0020006c0075006f006400610020005000440046002d0061007300690061006b00690072006a006f006a0061002c0020006a006f006900640065006e002000740075006c006f0073007400750073006c00610061007400750020006f006e0020006b006f0072006b006500610020006a00610020006b007500760061006e0020007400610072006b006b007500750073002000730075007500720069002e0020005000440046002d0061007300690061006b00690072006a0061007400200076006f0069006400610061006e0020006100760061007400610020004100630072006f006200610074002d0020006a0061002000520065006100640065007200200035002e00300020002d006f0068006a0065006c006d0061006c006c0061002000740061006900200075007500640065006d006d0061006c006c0061002000760065007200730069006f006c006c0061002e0020004e00e4006d00e4002000610073006500740075006b0073006500740020006500640065006c006c00790074007400e4007600e4007400200066006f006e0074007400690065006e002000750070006f00740075007300740061002e> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice acknowledgements references about the author(s) beanelri b. janecke department of animal, wildlife & grassland sciences, faculty of natural & agricultural sciences, university of the free state, bloemfontein, south africa johan van tol department of soil, crop and climate sciences, faculty of natural & agricultural sciences, university of the free state, bloemfontein, south africa citation janecke, b.b. & van tol, j., 2020, ‘connections between abiotic and biotic components of a granite catena ecosystem in kruger national park’, koedoe 62(2), a1638. https://doi.org/10.4102/koedoe.v62i2.1638 note: special issue: connections between abiotic and biotic components of a granite catena ecosystem in kruger national park, sub-edited by beanelri janecke and johan van tol. editorial connections between abiotic and biotic components of a granite catena ecosystem in kruger national park beanelri b. janecke, johan van tol received: 15 june 2020; accepted: 17 june 2020; published: 29 oct. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. this special issue is a compilation of multidisciplinary articles on a granitic ecosystem in the stevenson-hamilton supersite of kruger national park. the majority of research was conducted on a single catena during the same period (2015–2018), to establish connections between abiotic and biotic components. a framework for the multidisciplinary approach is presented and followed by a series of articles focussing on soils, hydrology, microorganisms in rhizospheres, vegetation and mammals. the catena concept refers to the co-evolution and distribution pattern of different soils along a hillslope, from the crest to the valley bottom. these differences in soils are largely driven by hydrological processes in savanna landscapes. the combination of soils and soil water regimes results in different vegetation patterns along the hillslope gradient, which affects other biotic components such as microorganisms and mammals. the catena in this study is located in a third-order catchment of the southern granite supersite, near skukuza in the kruger national park (knp). the supersite concept was introduced by smit et al. (2013) to encourage multidisciplinary research within a specific focus area of the park. the special issue is the product of a dedicated effort to bring eight principle, multidisciplinary research fields together to explore connections between ecosystem components within the same local study area. the underlying hypothesis was that heterogeneity of abiotic and biotic components at catena scale can be better understood and, ultimately, better managed through a multidisciplinary approach. this research has been conducted on a small scale to provide not only a basis for understanding, but also a framework for other similar studies. during the course of this project, two team members, dr fred kruger and prof. johann du preez, who were giants in ecological research, passed away. the tributes in this issue acknowledge their immense contribution not only to the project but also to natural science in general – our friends, we miss you dearly! two other colleagues stepped up to help with publishing the results of this project, to whom we are eternally grateful. the project commenced in 2015 and covered soil types and properties, hydropedological flowpaths, dry mud in depressions (small aquatic organisms), microorganisms and fungi in the rhizosphere of plants, vegetation, mammals, and the ecosystem. during 2015–2016, knp was badly affected by a severe drought. this impacted collection of certain samples that are dependent on sufficient rainfall to wet the soils, fill temporary pools, or lead to flow in the seep lines on the catena. samples were taken of dry mud in depressions to search for the presence of small aquatic organisms or their eggs. unfortunately, the drought hampered this research and no substantial results were obtained. vegetation structure, composition and mammal presence were still recorded during 2016 and 2017. the drought was partially broken during 2017. this special issue consists of a series of 11 articles and one short communication. it starts with an article, which summarises the main findings of the other articles and highlights connections between the abiotic and biotic trophic levels and components that formed part of this study (janecke et al. 2020). the southern granite supersite and research that was performed up to this point on the supersite is described in detail by smit (2020). he also provided an integration of multidisciplinary research as an introduction to this project. the next article describes the morphological and chemical properties of soil types along the catena (bouwer, le roux & van tol 2020). it further provides a conceptualisation of dominant hydrological processes based on the hydropedological interpretation of the spatial distribution of soils. the article by riddell et al. (2020) builds on this topic and quantified specific hydrological processes and properties (e.g. evapotranspiration, soil water contents and groundwater recharge). van tol et al. (2020) then used the soil and hydrological measurements to predict how changes in rainfall regimes will impact hydrological processes on this catena. the next article deals with the microbiome of the root zone (rhizosphere) of three plant species growing in different positions on the catena (vermeulen, cason & swart 2020). the variability of the fungal community structure in the rhizosphere follows on this topic (gryzenhout et al. 2020a). a short communication wraps up this topic and provides a first report of the presence of various fusarium species on the catena (gryzenhout et al. 2020b). soil, among other environmental factors, determines the type of vegetation that can grow in specific areas. different soil types were linked to plant communities and sub-communities distributed along the catena, from the top to the bottom in the third order drainage line (theron, van aardt & du preez 2020). the vegetation structure was also found to differ between the hillslope zones (janecke 2020), and this led to a variety of habitat types for animals. the impact of the drought on vegetation and the recovery of plants on the catena over a period of 3–4 years were covered in an article by van aardt et al. (2020). the variety of habitat types created by different plant communities in the catenal zones were also linked to mammal presence on the catena and at the waterholes in the vicinity (janecke & bolton 2020). this special issue provides a basis for possible future studies that can build on the foundation laid by this project. it also supports the concept encouraged by knp to focus multidisciplinary research on areas such as the supersites. systems are unstable, complex and open, with random reactions and different processes that continuously need further research and investigation. we trust that this research will spark the interest of other researchers and emphasise the importance of multidisciplinary studies to provide links and further connections, and in the process fill any gaps in knowledge that might still exist of the various components of an ecosystem. acknowledgements the research presented in this special issue was financially supported by several institutions; each is acknowledged in the specific publications. as guest editors, we would, however, like to acknowledge the university of the free state central research fund and sanparks for their financial contributions towards publishing the issue, as well as for their vision in supporting the multidisciplinary project which forms the core of articles in this special issue. further, we would like to thank all the reviewers who contributed to the quality of the manuscripts, and our research team (the eight principle researchers, 12 authors from ufs and two from sanparks in associated collaboration with authors from 14 other institutions) for all their hard work, long hours and dedication to this project. lastly, this issue would not have been possible without the assistance of the friendly staff of sanparks scientific services and aosis publishers who were always ready to assist. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this editorial. references bouwer, d., le roux, p.a.l. & van tol, j., 2020, ‘identification of hydropedological flowpaths in stevenson–hamilton catena from soil morphological, chemical and hydraulic properties’, koedoe 62(2), a1584. https://doi.org/10.4102/koedoe.v62i2.1584 gryzenhout, m., cason, e.d., vermeulen, m., kloppers, g.a.e., bailey, b. & ghosh, s., 2020a, ‘fungal community structure variability between the root rhizosphere and endosphere in a granite catena system in kruger national park, south africa’, koedoe 62(2), a1597. https://doi.org/10.4102/koedoe.v62i2.1597 gryzenhout, m., vermeulen, m., pambuka, g. & jacobs, r., 2020b, ‘first report of various fusarium species from the stevenson-hamilton supersite granite catena system in the kruger national park, south africa’, koedoe 62(2), a1599. https://doi.org/10.4102/koedoe.v62i2.1599 janecke, b.b., 2020, ‘vegetation structure and spatial heterogeneity in the granite supersite, kruger national park’, koedoe 62(2), a1591. https://doi.org/10.4102/koedoe.v62i2.1591 janecke, b.b. & bolton, j.g., 2020, ‘variation in mammal diversity and habitat affect heterogeneity and processes of a granite catena’, koedoe 62(2), a1592. https://doi.org/10.4102/koedoe.v62i2.1592 janecke, b.b., van tol, j., smit, i.p.j., van aardt, a.c., riddell, e.s., seaman, m.t. et al., 2020, ‘biotic and abiotic connections on a granitic catena: framework for multidisciplinary research’, koedoe 62(2), a1600. https://doi.org/10.4102/koedoe.v62i2.1600 riddell, e.s., nel, j., van tol, j., fundisi, d., jumbi, f., van niekerk, a. et al., 2020, ‘groundwater–surface water interactions in an ephemeral savanna catchment, kruger national park’, koedoe 62(2), a1583. https://doi.org/10.4102/koedoe.v62i2.1583 smit, i.p.j., riddell, e.s., cullum, c. & petersen, r., 2013, ‘kruger national park research supersites: establishing long-term research sites for cross-disciplinary, multiscaled learning’, koedoe 55(1), art. #1107, 7 pages. http://dx.doi.org/10.4102/koedoe.v55i1.1107 smit, i.p.j., 2020, ‘integrating multi-scaled and multidisciplinary studies: a critical reflection on the kruger national park research supersites’, koedoe 62(2), a1586. https://doi.org/10.4102/koedoe.v62i2.1586 theron, e.j., van aardt, a.c. & du preez, p.j., 2020, ‘vegetation distribution along a granite catena, southern kruger national park, south africa’, koedoe 62(2), a1588. https://doi.org/10.4102/koedoe.v62i2.1588 van aardt, a.c., codron, d., theron, e.j. & du preez, p.j., 2020, ‘plant community structure and possible vegetation changes after drought on a granite catena in the kruger national park, south africa’, koedoe 62(2), a1585. https://doi.org/10.4102/koedoe.v62i2.1585. van tol, j., julich, s., bouwer, d. & riddell, e.s., 2020, ‘hydrological response in a savanna hillslope under different rainfall regimes’, koedoe 62(2), a1602. https://doi.org/10.4102/koedoe.v62i2.1602 vermeulen, m., cason, e.d. & swart, w.j., 2020, ‘the rhizobiome of herbaceous plants in clovelly and sterkspruit soils of the stevenson-hamilton supersite’, koedoe 62(2), a1596. https://doi.org/10.4102/koedoe.v62i2.1596 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 filelist convert a pdf file! page 1 page 2 page 3 kruger andries final.qxd an analysis of skukuza climate data a.c. kruger, l.b. makamo and s. shongwe kruger, a.c., l.b. makamo and s. shongwe. 2002. an analysis of skukuza climate data. koedoe 45(1): 1–7. pretoria. issn 0075-6458. data from the climate station at skukuza, which has been open from 1912 to the present, are analysed. this exercise was done to provide the south african regional science initiative (safari-2000) experimental program with long-term climate statistics and trends. climate parameters analysed are rainfall, temperature, humidity and sunshine. in the case of rainfall and temperature, the data was first tested for homogeneity and in only three out of 36 monthly cases, the data proved not to be homogeneous. no monthly rainfall trends proved to be significant (at the 5 % level), with five months indicating slightly negative trends and seven slightly positive. only the monthly maximum temperature series for june proved to be non-homogeneous. the june maximum temperature trend and the february, march, may, july and december minimum temperature trends were significantly positive. the annual time series for minimum temperature were also significantly positive. the monthly results were reiterated by the seasonal results, with the winter maximum temperature trend and the autumn and summer minimum temperature trends significantly positive. ten months showed negative temperature diurnal range trends with only march being significant. all long-term statistics of rainfall, temperature, humidity and sunshine were found to be typical of a savanna type climate in the southern hemisphere, although average monthly sunshine hours were somewhat less than the norm, due to frequent influx of moist air from the mozambique channel. key words: skukuza, safari-2000, climate, trends. a.c. kruger, directorate: climate systems, south african weather service, private bag x097, pretoria, 0001 south africa (andries@weathersa.co.za); l.b. makamo and s. shongwe, directorate: climate systems, south african weather service, private bag x097, pretoria, 0001 south africa. issn 0075-6458 1 koedoe 45/1 (2002) introduction the south african regional science initiative (safari-2000) experimental program is developed for southern africa to explore, study and address linkages between landatmosphere processes and the relationship of biogenic, pyrogenic or anthropogenic emissions and the consequences of their deposition to the functioning of biogeophysical and biogeochemical systems (safari-2000 home page). one of the safari-2000 sites is about 13 km wsw of skukuza in the mpumalanga province. many scientists are involved in the project or will use data forthcoming from it. however, climate data from the site itself only has a measuring period of about two years, from 2000 to 2001. the purpose of this paper is to provide an analysis of the data of the south african weather service climate station at skukuza, entailing the calculations of long-term averages and trends, for the purpose of background data to interpret measurements at the flux measurement site. general climate characteristics of the area the climate station of skukuza is situated at 24º59's, 31º36'e at a height of 263 m above sea level. the area the station is situated in is generally known as the lowveld and is kruger andries final.qxd 2005/12/09 11:32 page 1 directly east of the eastern escarpment. according to the new climate regions developed by the south african weather service, loosely based on the vegetation regions of low & rebelo (1996) and the köppen climate classification, skukuza is situated in the lowveld bushveld region, which receives moderate summer precipitation (500–700 mm p.a.) with maximum rainfall in january. warm to hot temperatures are usually experienced and no frost occurs. the humidity is usually fairly high and this makes summer days uncomfortable. sunshine duration during summer is below average for this typical savanna climate, due to the influx of moist air from the coast and exacerbated by the proximity of the southern african escarpment close by towards the west. this type of climate lends itself to game, cattle and goat farming, subtropical fruit, vegetables and sugarcane through irrigation, and ecotourism. analysis of rainfall to obtain a general idea of the rainfall climate of skukuza, trends and long-term statistics are calculated. but before any analyses can be done, the data first have to be checked for overall quality, as inhomogeneities can influence the results significantly, especially in the case of trend calculations. homogeneity of rainfall data to test for the homogeneity of skukuza rainfall data, the non-parametric run test described by thom (1966) was applied to monthly time series for january to december. this test is based on the number of runs (or groups of consecutive data points) above or below the median value of the time series. the results are shown in table 1. in the second column the number of runs are shown which should ideally be about half of the total years used in the calculation (in this case about 80 years of data), to accept the time series as homogeneous. the third column shows the results when compared to the koedoe 45/1 (2002) 2 issn 0075-6458 table 1 results of run test to determine homogeneity of monthly rainfall time series month number of runs result jan 49 homogeneous feb 38 homogeneous mar 34 non-homogeneous apr 37 homogeneous may 34 non-homogeneous jun 44 homogeneous jul 39 homogeneous aug 37 homogeneous sep 36 homogeneous oct 43 homogeneous nov 38 homogeneous dec 38 homogeneous lower and upper 10 % significance limits. two of the months, march and may, showed possible inhomogeneities in their series, with much fewer than the ideal number of runs. this might indicate a possible trend or slippage in the mean. this is however not evident from the trend calculations, of which the results are shown in the following section. thus, another possible explanation for the inhomogeneity results is that there might be relatively large clusters of consecutive months with rainfall above or below the median value, but not distributed in a way to cause a trend in the data. for march the largest clusters or runs with four or more consecutive values above or below the median are from 1912 to 1919 (below the median), 1929 to 1934 (above the median), 1943 to 1946 and 1962 to 1966 (below the median), 1967 to 1972 (above the median), and 1973 to 1976 and 1982 to 1986 (below the median). for may the runs are 1934 to 1937 (above the median), 1943 to 1946 (below the median), 1948 to 1951 (above the median), 1962 to 1965 and 1967 to 1970 (below the median), and 1979 to 1983 and 1990 to 1993 (above the median). except for 1943 to 1946 and 1962 to 1965, none of the above periods are the same for both march and may, thus mostly excluding the probability of artificial biases introduced to the climate series, causing recorded values to be too high or low. also, no mention is made in the metadata of kruger andries final.qxd 2005/12/09 11:32 page 2 the station of changes in exposure during, just before or just after the mentioned periods, which lets one come to the conclusion that the above clusters are most possibly natural phenomena, a true reflection of the rainfall received at the station. no attempt was thus made to adjust data in the series before further analysis. trends in rainfall table 2 shows the analysis of linear trends in monthly rainfall time series for skukuza for the period 1912 to 2001. the month with the highest trend is december with a very small positive value of 0.41 mm p.a. none of the trends calculated were significant at the 5% significance level. trends were also calculated for seasonal rainfall, where summer is defined as the months from december to february, autumn from march to may, winter from june to august and spring from september to november. the results are shown in table 3. again, very small trends were detected, the highest being 1.4 mm p.a. for summer while none of the trends were significant. these results indicate that the long-term rainfall average for skukuza has remained fairly constant over the past 90 years. issn 0075-6458 3 koedoe 45/1 (2002) long-term rainfall statistics figure 1 and table 4 show long-term rainfall statistics for the period 1912 to 2001. similar statistics is supplied by scholes et al. (2001), but only for a shorter period, ending in 1999. the analyses reveal a typical summer rainfall climate in the southern hemisphere. from the absolute maximum and minimum monthly rainfall it is clear that the rainfall is highly variable, making the area prone to frequent droughts and floods. temperature the same approach was followed with temperature analyses as in the case of rainfall. firstly the data was checked for homogeneity, and after that long-term trends and statistics were calculated. homogeneity of temperature data as with rainfall, the run test described by thom (1966) was used to test monthly time series for homogeneity. the total number of years of data is 42 years (1960 to 2001). table 2 monthly rainfall trends (mm p.a.) for the period 1912 to 2001 month jan feb mar apr may jun jul aug sep oct nov dec trend –0.25 0.25 –0.13 0.13 –0.02 –0.03 –0.03 0.08 0.07 0.18 0.08 0.41 table 3 seasonal rainfall trends (mm p.a.) for the period 1912 to 2001 season autumn winter spring summer trend –0.07 0.07 1.1 1.4 fig. 1. average monthly rainfall (mm) for the period 1912 to 2001. kruger andries final.qxd 2005/12/09 11:32 page 3 koedoe 45/1 (2002) 4 issn 0075-6458 table 4 rainfall statistics (mm) for the period 1912 to 2001 statistic jan feb mar apr may jun jul aug sep oct nov dec absolute monthly maximum 835 531 260 222 83 110 132 182 193 231 434 331 absolute monthly minimum 8 0 4 0 0 0 0 0 0 0 0 1 maximum daily rainfall 103.5 119.5 86.5 71.5 56.5 40.1 75.2 40.0 85.6 44.4 67.3 99.5 average amount of rainy days (>= 0.1 mm) 9.8 9.6 8.8 5.7 3.0 2.1 1.8 2.3 3.3 7.0 10.6 9.8 table 5 results of run test to determine homogeneity of maximum and minimum temperature time series month maximum maximum minimum minimum temperature temperature temperature temperature runs result runs result jan 23 homogeneous 17 homogeneous feb 19 homogeneous 16 homogeneous mar 21 homogeneous 24 homogeneous apr 24 homogeneous 20 homogeneous may 23 homogeneous 24 homogeneous jun 26 non-homogeneous 19 homogeneous jul 19 homogeneous 18 homogeneous aug 23 homogeneous 17 homogeneous sep 22 homogeneous 22 homogeneous oct 25 homogeneous 19 homogeneous nov 23 homogeneous 17 homogeneous dec 23 homogeneous 25 homogeneous according to the test, the ideal number of runs should be approximately 21 for a homogeneous data set. the results for the maximum and minimum temperatures are shown in table 5. the third and last columns show the results of the test when compared to the lower and upper 10 % significance limits. only the maximum temperature series for july showed possible inhomogeneities in its data series, indicating a high number of short runs in the series. this may in turn indicate a climatic vaccilation of the mean, or relatively short cycles in the series, although this was not verified by graphical analysis. trends in maximum and minimum temperature table 6 shows the trends of monthly average maximum and minimum temperatures. for the maximum temperature, only july showed kruger andries final.qxd 2005/12/09 11:32 page 4 a significant trend at the 5 % level. for the minimum temperature, five of the months showed significantly positive trends, as well as the annual average. these results are consistent with global trends where the minimum temperatures show higher positive trends than the maximum temperature (easterling et al. 2000), causing the diurnal range in temperature to become progressively smaller. table 7 shows the seasonal trends for maximum and minimum temperatures. the maximum temperature trend in winter, as well as the minimum temperature trend in autumn and summer, showed significantly positive trends. issn 0075-6458 5 koedoe 45/1 (2002) table 6 monthly and average trend of maximum and minimum temperatures (°c p.a.) for the period 1960 to 2001 (* indicates significance at the 5% level) month maximum minimum temperature temperature jan –0.020 0.007 feb –0.005 0.029* mar –0.006 0.035* apr 0.026 0.021 may 0.010 0.034* jun 0.037* 0.011 jul 0.008 0.041* aug 0.009 0.015 sep –0.002 0.026 oct –0.009 0.018 nov 0.015 0.018 dec –0.001 0.019* average 0.005 0.024* table 7 seasonal trends of maximum, minimum and average temperatures (°c p.a.) for the period 1960 to 2001 (* indicates significance at the 5 % level) season maximum minimum average temperature temperature temperature autumn 0.010 0.030* 0.020 winter 0.018* 0.029 0.024* spring 0.000 0.020 0.010 summer –0.009 0.018* 0.013 table 8 trends of monthly diurnal temperature range (°c p.a.) for the period 1960 to 2001 (* indicates significance at the 5 % level) month jan feb mar apr may jun jul aug sep oct nov dec trend -0.027 -0.034 -0.041* 0.005 -0.025 0.027 -0.033 -0.007 -0.028 -0.027 -0.004 -0.020 trends in diurnal range of temperature table 8 shows the monthly trends in diurnal range of temperature. as can be expected, corresponding with the relative trends in maximum and minimum temperature, the trend is negative for most months, although only the trend for march is significant. for the seasonal trends of diurnal range, as shown in table 9, all the results were negative. autumn shows the highest negative trend of –0.023 ºc per year. long-term temperature statistics figure 2 and table 10 show temperature statistics for the total period of record (1960 to 2001). similar statistics is supplied by scholes et al. (2001), but for the period until 1999. summer temperatures are hot, while during the winter it is mostly warm and pleasant. minimum temperatures rarely drop below freezing point during winter, but even at this time of the year maximum temperatures can reach levels in the mid-thirties. kruger andries final.qxd 2005/12/09 11:32 page 5 humidity because of frequent influxes of moist air from the east, the humidity should be relatively high. this is reflected in table 11. the average monthly humidity at 14:00 sast, koedoe 45/1 (2002) 6 issn 0075-6458 table 9 trends of seasonal diurnal temperature range (°c p.a.) for the period 1960 to 2001 season autumn winter spring summer trend –0.023 –0.010 –0.021 –0.017 table 10 long-term temperature statistics (°c) for the period 1960 to 2001 month jan feb mar apr may jun jul aug sep oct nov dec absolute minimum 11.3 10.0 7.6 5.6 0.8 –4.4 –3.8 –4.2 1.3 6.4 9.9 2.0 absolute maximum 43.0 45.6 42.0 41.3 39.2 35.3 36.4 38.0 42.6 43.6 42.8 42.4 table 11 relative humidity (%) at 14:00 sast for the period 1978 to 2001 month jan feb mar apr may jun jul aug sep oct nov dec average 53 54 53 48 43 38 39 38 41 45 52 51 lowest monthly 39 37 40 35 30 29 28 30 33 36 39 36 highest monthly 66 77 67 57 64 57 58 60 57 56 84 62 table 12 monthly sunshine data (hours) for the period 1960 to 2001 month jan feb mar apr may jun jul aug sep oct nov dec total average 7.3 7.3 7.2 7.5 8.0 8.1 8.1 8.1 7.5 6.7 6.2 7.0 lowest average 4.8 5.2 4.4 5.4 6.4 6.7 6.4 5.8 5.5 5.0 4.2 4.8 highest average 9.1 9.6 9.3 9.4 9.4 9.4 9.6 9.5 9.3 8.5 7.8 10.8 which is usually the driest time of the day, ranges from a maximum of 54 % for february to a minimum of 38 % for june and august. sunshine average sunshine data for 1960 to 2001 is shown in table 12. one can see that, although daylight hours in summer is much longer than in winter, the sunshine hours show the opposite, indicating cloudier weather during summer. this region actually receives markedly less sunshine than the kruger andries final.qxd 2005/12/09 11:32 page 6 norm for savanna regions, the reason being the frequent influx of moist air from the east, with its accompanying fog and cloud. discussion and conclusions the motivation for this paper is to supply researchers, and other interested parties involved in the safari-2000 project, sufficient long-term background data to serve as a baseline for comparison to future observations at the flux measurement site close by. meta data and homogeneity tests suggest that the data is generally of a high quality, suitable for trend analyses. the average rainfall (monthly and seasonal) stayed fairly constant during the previous century. howissn 0075-6458 7 koedoe 45/1 (2002) fig. 2. average monthly maximum and minimum temperature (°c) for the period 1960 to 2001. ever, there has been a marked increase in minimum temperature since the 1960s, especially during the summer and autumn months. this result corresponds to the average global trend (easterling et al. 2000), and can also be considered a true reflection of temperature trends in the area, since no meaningful urbanisation or increased pollution has taken place since the starting point of the time series. long-term rainfall and temperature statistics indicate a highly variable climate, typical of the african savanna. references easterling d.r., t.r. karl, k.p. gallo, d.a. robinson, k.e. trenberth & a. dai. 2000. observed climate variability and change of relevance to the biosphere. journal of geophysical research 105: 20101–20114. low, a.b. & a.g. rebelo (eds.). 1996. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. scholes, r.j., n. gureja, m. giannecchinni, d. dovie, b. wilson, n. davidson, k. piggott, c. mcloughlin, k. van der velde, a. freeman, s. bradley, r. smart & s. ndala. 2001. the environment and vegetation of the flux measurement site near skukuza, kruger national park. koedoe 44(1): 73–83. safari-2000 www home page. 2002. virginia state university. virginia. u.s.a. (http:// safari.gecp.virginia.edu/) thom, h.c.s. 1966. some methods of climatological analysis. geneva, switzerland: world meteorological organization. (wmo technical note; no. 81.) kruger andries final.qxd 2005/12/09 11:32 page 7 koedoe 45/1 (2002) 8 issn 0075-6458 kruger andries final.qxd 2005/12/09 11:32 page 8 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <feff004200720075006700200064006900730073006500200069006e0064007300740069006c006c0069006e006700650072002000740069006c0020006100740020006f0070007200650074007400650020005000440046002d0064006f006b0075006d0065006e0074006500720020006d006500640020006800f8006a006500720065002000620069006c006c00650064006f0070006c00f80073006e0069006e0067002000740069006c0020007000720065002d00700072006500730073002d007500640073006b007200690076006e0069006e0067002000690020006800f8006a0020006b00760061006c0069007400650074002e0020005000440046002d0064006f006b0075006d0065006e007400650072006e00650020006b0061006e002000e50062006e006500730020006d006500640020004100630072006f0062006100740020006f0067002000520065006100640065007200200035002e00300020006f00670020006e0079006500720065002e00200044006900730073006500200069006e0064007300740069006c006c0069006e0067006500720020006b007200e600760065007200200069006e0074006500670072006500720069006e006700200061006600200073006b007200690066007400740079007000650072002e> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 reviewer acknowledgement http://www.koedoe.co.za open access koedoe recognises and acknowledges the value and importance of reviewers in the publication process – not only in shaping the individual manuscript, but also in upholding the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely, productive peer review process. we would like to take this opportunity to thank all reviewers who participated in shaping this volume of koedoe: page 1 of 1 in an effort to facilitate the selection of appropriate peer reviewers for koedoe, we ask that you take a moment to update your electronic portfolio at http://www.koedoe. co.za for our records. this will allow us better access to your areas of interest and expertise, in order to match reviewers with submitted manuscripts. if you would like to become a reviewer, please visit the journal website and register as a reviewer. to access your details on the website, you will need to follow these steps: 1. log into the online journal at http://www. koedoe.co.za 2. in your ‘user home’ [http://www.koedoe. co.za/index.php/koedoe/ user] select ‘edit my profile’ under the heading ‘my account’ and insert all relevant details, bio statement and reviewing interest. 3. it is good practice as a reviewer to update your personal details regularly, to ensure contact with you throughout your professional term as reviewer to koedoe. please do not hesitate to contact us if you require assistance in performing this task. publisher: publishing@aosis.co.za tel: +27 21 975 2602 fax: +27 21 975 4635 adrian j. armstrong adrian m. shrader alta de vos amanda bastos ana l. nunes andrew drake andrew r. deacon angela gaylard anna songhurst arthur perroten benjamin j. wigley brian jones brian reilly carrie roever christiaan w. winterbach christina skarpe coral birss danny govender dave balfour david le maitre doug harebottle ed witkowski eddie riddell edson gandiwa elizabeth berkeley elizabeth freeman elmer topp-jorgensen emily bennitt felix patton florian weise frances siebert francois deacon gareth hempson glyn vale gregory rasmussen gretel van rooyen guillaume latombe guy castley haemish melville harriet bartlam-brooks helga van der merwe hendrik sithole hugo bezuidenhout ian a. russell ingrid wiesel ioannis tsiripidis james pryke jan a. venter jeanetta selier jeremy perkens jose a. corronca keith leggett kerry slater kristine maciejewski leanne faulks leo braack leon n. lotz lesley henderson lochran traill mamokete dingaan mariska te beest mark keith markus gusset mduduzi ndlovu michael j. somers neville d. impson paul cross peter goodman rickert van der westhuizen rowshyra castañeda sadie ryan sam williams samantha williams samuel osinubi sean m. marr sharon thompson simon chamaille sindiso chamane solomon mombeshora stefan schindler stefan h. foord sue snyman susan cunningham tess cole tony booth trevor booth acknowledgement to reviewers http://www.koedoe.co.za http://www.koedoe.co.za http://www.koedoe.co.za http://www. koedoe.co.za http://www. koedoe.co.za http://www.koedoe. co.za/index.php/koedoe/ user http://www.koedoe. co.za/index.php/koedoe/ user http://www.koedoe. co.za/index.php/koedoe/ user mailto:publishing@aosis.co.za about the author(s) andy blackmore ezemvelo kzn wildlife, pietermaritzburg, south africa school of law, university of kwazulu-natal, pietermaritzburg, south africa citation blackmore, a., 2020, ‘erratum: climate change and the ownership of game: a concern for fenced wildlife areas’, koedoe 62(1), a1659. https://doi.org/10.4102/koedoe.v62i1.1659 note: doi of original article: https://doi.org/10.4102/koedoe.v62i1.1594 correction erratum: climate change and the ownership of game: a concern for fenced wildlife areas andy blackmore published: 11 dec. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. in the version of this article initially published, blackmore, a., 2020, ‘climate change and the ownership of game: a concern for fenced wildlife areas’, koedoe 62(1), a1594. https://doi.org/10.4102/koedoe.v62i1.1594, the author’s second affiliation was omitted in the ‘authors’ and ‘affiliations’ sections. the author affiliations are now corrected as: 1ezemvelo kzn wildlife, pietermaritzburg, south africa 2school of law, university of kwazulu-natal, pietermaritzburg, south africa this correction does not alter the study’s findings of significance or overall interpretation of the study’s results. the publisher apologises for any inconvenience caused. abstract introduction discussion conclusion acknowledgements references about the author(s) andy blackmore integrated environmental management and protected area planning, scientific services, ezemvelo kzn wildlife, durban, south africa citation blackmore, a., 2020, ‘climate change and the ownership of game: a concern for fenced wildlife areas’, koedoe 62(1), a1594. https://doi.org/10.4102/koedoe.v62i1.1594 original research climate change and the ownership of game: a concern for fenced wildlife areas andy blackmore received: 16 sept. 2019; accepted: 20 mar. 2020; published: 22 june 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract wildlife and particularly economically valuable game are likely to be displaced as a result of the habitat change. this displacement is expressed, inter alia, in the emigration of game to a more suitable habitat. the impacts of climate change, therefore, may have significant consequences on the economic well-being of wildlife areas, which are derived from, amongst others, sale of excess animals, hunting and tourism. this article investigates whether the south african statute and common law provide sufficient protection to landowners, from a game ownership perspective, as the impacts of climate change become evident. it was discovered that the complexity derived from the relationship between landownership, legislation and common law (1) required wildlife areas to be isolated fenced areas, (2) may lead to loss of ownership of game which escapes as a consequence of climate change and (3) provided for the possible loss of ownership of all game occurring in the wildlife area and those emigrating when all or part of an encircling boundary fence is removed to establish a wildlife or climate change corridor. it is further recommended that the game theft act 105 of 1991 requires substantial amendment to enable owners of wildlife areas to retain ownership of game that escapes or emigrates in response to climate change. finally, it is recommended that landowners acquire and include into their fenced wildlife areas an additional area as an interim measure to mitigate the impacts of climate change, until such time that the desired legislative change is implemented. conservation implications: climate change has serious implications for continued ownership of escaped wildlife as well as for the implementation of adaptive strategies to mitigate the impacts of a changing climate on fenced wildlife areas. the south african law needs to be revised to protect wildlife owners as the impacts of climate change become evident. keywords: climate change; common law; conservation; game fences; game farms; game theft act 105 of 1991; game; ownership; protected areas; rural communities; wildlife. introduction climate change is increasingly being cited as one of the major challenges facing the conservation of biodiversity (bellard et al. 2012), and with this realisation a greater emphasis is being placed on protected areas and other conserved natural areas (i.e. game farms and ranches) as refugia as well as ‘island habitats’ for wildlife. although this argument may hold in open systems within which protected areas occur and where, at least, wildlife may freely move beyond the boundary of the conserved area, it may not hold for areas that are fenced, as is predominantly the case in south africa, in a manner that limits the movement of species responding to climate change. in such circumstances, the recommendation is to physically relocate climate change-displaced species to areas of more suitable habitat, or establish biodiversity or wildlife corridors that are abundant in the scientific literature (davidson et al. 2012; di minin et al. 2013; groves et al. 2012; hartter, goldman & southworth 2011; lister et al. 2015; madden 2008; mcdowell 2013; minteer & collins 2010; pittiglio et al. 2014; sax, smith & thompson 2009; schwartz et al. 2012; songhurst et al. 2016; syombua 2013; treves 2009; wilke & rannow 2014). although this argument is sound from many perspectives, there are a number of social, economic, conservation, legal and practical challenges that may need to be overcome before a species can be either extracted and re-introduced elsewhere, or stable and effective corridors can be established. as the impacts of climate change manifest, conserved natural areas are likely to experience changes in rainfall (viz. water availability), temperature and vegetation type and abundance. wildlife will most likely respond to these environmental changes by either a reduction in density or by moving out of the conserved area to seek a more suitable habitat or prey, or a combination of both (corlett & westcott 2013; davidson et al. 2012; hannah et al. 2005; monzón, moyer-horner & palamar 2011). thus, from a wildlife perspective, particularly in countries that have inherited the roman common law such as south africa (pienaar 2012; van der merwe & rabie 1974), one of the biggest challenges to be overcome before implementing the recommendation either to relocate animals or allow wildlife to move freely out and away from the conserved area is one of ‘ownership’. this has received little emphasis in terms of management of conserved areas in response to climate change. this article, from a south african perspective, therefore, focusses on the legal challenges that are likely to arise from game (a subset of wildlife) escaping from a fenced wildlife area or moving out of such area, by way of corridors, to a more suitable habitat, in response to altered habitats as a result of climate change. discussion ownership of game ownership of game in the roman common law, as with many legal matters, may be complex. in brief, however, game in south africa are considered ‘res nullius’, meaning that they are unowned or ‘property of no one’ (blackmore 2017; muir 2016; richter v. du plooy 1921; selier et al. 2016; van der merwe 2002). ownership, however, can be achieved by a person taking possession of an animal, that is, to be in purposeful physical control of it. the possession of game is commonly achieved, according to south african common law, by a person through hunting, seizing, capturing or uniquely marking (e.g. branding) an animal with a predetermined intention to take possession of it. thus, wildlife roaming on a game farm or ranch or in a protected area (a ‘wildlife area’) would not necessarily be owned by the landowner. as such, a person who has hunted an animal, even without permission, could not be charged with theft (i.e. poaching), but may be liable for trespassing (eastern cape parks and tourism agency v. medbury (pty) ltd 2018; richter v. du plooy 1921; van der merwe 2002). irrespective of the legality of the hunt, the hunter retains legal ownership of the animal in that the hunter has purposefully taken physical control of it (eastern cape parks and tourism agency v. medbury (pty) ltd 2018; richter v. du plooy 1921). thus, for a landowner to possess and own the wildlife on his or her property, the landowner must take physical control of each animal. once achieved, if any other person takes an animal, it would be considered theft (magudu game company (pty) ltd v. mathenjwa no and others 2008; van der merwe 2002). as noted by muir (2016), it is undesirable, impractical and illogical for an owner of a wildlife area to take physical control of each individual animal on the property. for this reason, the game theft act 105 of 1991 was promulgated (couzens et al. 2019; muir 2016; republic of south africa 1991; rumsey 2009; van der merwe 2002). this statute grants the landowner ownership of certain game on his or her property that has been adequately enclosed with a game-proof fence. should such circumstance arise, the landowner would be reasonably entitled to retrieve the game that are known to have escaped into neighbouring properties (the ‘escape rule’) (eastern cape parks and tourism agency v. medbury (pty) ltd 2018; freedman 2019). furthermore, the landowner would not lose ownership of an animal to another person who without permission gains physical control of it (i.e. through capture or hunting) within the fenced property. game ownership in terms of the game theft act is, however, not all encompassing in that its provisions are limited to its purpose (i.e. ‘to regulate the ownership of game in certain instances’ – preamble to the act) and ultimately the purpose for which the act was promulgated (to protect the interests of the commercial industry from the theft of game) (freedman 2019; muir 2016). it is, therefore, unlikely that the writers of the act took into consideration the common law on the natural ‘re-wilding’ of wildlife, a principle applied to game that escape from control as a result of natural processes. acknowledging that certainty may only be achieved in the courts, the application of this principle, within the context of climate change, may, therefore, lead to the loss in ownership of animals by the landowner. thus, without going into detailed argument, for the purposes of this article, it is cautiously assumed that the game theft act does not modify or set aside the common law principle of re-wilding. the legal status of wildlife, therefore, as described briefly above, has significant consequences for the ownership of game and, in particular, those species that may be displaced by the impacts of climate change. game and wildlife industry since the 1990s, south africa has witnessed an accelerated increase in the numbers and extent of wildlife ranches and private protected areas as landowners have responded to the increased demand for and the concomitant increased economic value of game. this demand was facilitated, if not created, predominantly by the security of ownership of game conferred by the game theft act (carruthers 2008; cousins, sadler & evans 2010; eastern cape parks and tourism agency v. medbury (pty) ltd 2018; kamuti 2014; muir 2016; vos et al. 2019). following the democratisation of south africa in 1994 and the subsequent restitution of land to communities and people dispossessed of land during the apartheid era, there has been a marked increase in the number of independent rural community protected areas and rural communities acquiring ownership of part or all of existing protected areas (koelble 2011). in addition, a growing interest in the biodiversity-tourism economy has also facilitated a number of rural communities contributing land to and expanding existing state and private protected areas (koelble 2011). the economic value, via ownership secured by the provisions of the game theft act, enables these emergent wildlife stakeholders to benefit from the wildlife economy and associated tourism industry (koelble 2011). two aspects that are common to private, state and communal wildlife areas are that they have well-defined boundaries and are fenced to (1) prevent the game from escaping and (2) confer statutory ownership of wild animals on the property, irrespective of whether they are naturally occurring or introduced. the complexity derived from the relationship between landownership (i.e. the ownership of distinct parcels of land forming constituting a ranch or wildlife management area), the game theft act and the common law pertaining to game has facilitated (if not caused to) the network of wildlife areas in south africa being isolated from each other and managed independently of neighbouring areas. the corollary of this is that wildlife areas have few functional linkages and migratory corridors for large wildlife species to, inter alia, respond to the impacts of climate change (cushman et al. 2018; newmark 2008). game ownership and climate change the impact of climate change on enclosed wildlife areas is likely to become a significant consideration with the passage of time (barbier et al. 2011; hulme 2005; mawdsley, o’malley & ojima 2009; selier et al. 2016). these impacts include, amongst others, a significant change in vegetation assemblages and the abundance of species (carter et al. 2014; davidson et al. 2012), displacement of habitats and species (agrawal & redford 2009; batllori et al. 2017; carter et al. 2014; monzón et al. 2011) and an increased risk of human–wildlife conflict as potential damage-causing animals emigrate from wildlife areas into neighbouring areas as a consequence of reduction in either the quantity of quality habitat or prey species, or both (johnson, karanth & weinthal 2018; lamichhane et al. 2018; nyhus 2016; selier et al. 2016). although the impacts of climate change may be mitigated and ameliorated through adaptive management of the wildlife area (e.g. changes in burning regimes and addition of fodder), a threshold exists beyond which the costs and practicality of such actions outweigh the benefits gained (koprowski & krausman 2019). at this point, be it a limit of acceptable change or a cautious or risk-averse threshold of potential concern (blackmore 2015), a decision needs to be taken to either allow, inter alia, local populations to substantially reduce in numbers and condition, go extinct or to actively facilitate emigration of affected wildlife to a more suitable habitat. notwithstanding these consequences that climate change will have on the integrity and tourism attractiveness of wildlife areas, it has serious implications for game ownership. these implications are discussed below. common law re-wilding notwithstanding the principal purpose of the game theft act to provide certainty of ownership of game that have been adequately enclosed, this certainty may not be absolute or perpetual (muir 2016). for instance, the act may not necessarily modify or override the common law principle of re-wilding. such circumstances occur when one or more animals escape from the wildlife area as a result of natural processes, that is, when the ecological carrying capacity is exceeded. south african common law views these animals as reverting to their wild [ferae naturae] non-captive state. such escaped animals would be considered ‘res nullius’ and, as such, can be legally seized (i.e. hunted or captured) and hence owned by another person (freedman 2019; guyon 2018; magudu game company (pty) ltd v. mathenjwa no and others 2008). it is therefore conceivable that wildlife, particularly those species that are considered ‘game’ in terms of the game theft act, that are being displaced by the impacts of climate change, and escape from the adequately enclosed wildlife area into adjacent areas, are likely to be considered to be ‘re-wilding themselves’ or regaining their natural freedom (eastern cape parks and tourism agency v. medbury (pty) ltd 2018; freedman 2019). the potential consequence of climate change in this regard, together with a laissez-faire approach to this cause of displacement of wildlife, therefore, may lead to a potential loss of valuable animals (without compensation) from the wildlife area. although the concept of re-wilding has not been prominent in legal arguments pertaining to game ownership disputes, its consideration in the medbury case (and others) confirms, unless successfully challenged, the existence of this principle in south african common law. it is therefore surmised that re-wilding is likely to become a key basis for legal argument challenging the applicability of the escape rule, as the impacts of climate change become evident at a wildlife area level. with the progression of time, a continual reliance on the escape clause (as provided by the game theft act) may, therefore, be an ill-advised strategy to be followed by game owners in that they risk losing ownership of game that escape from their fenced wildlife areas. it is thus recommended that wildlife areas should be progressively expanded, at least, as an interim measure to decrease the risk of climate change-induced escape of game and to counter arguments of re-wilding should they arise. climate change corridors establishing wildlife corridors that enable wildlife to emigrate to a more suitable habitat has been advocated as a viable adaption strategy to mitigate the impacts of climate change (caro et al. 2009; landguth et al. 2012; lister et al. 2015; mawdsley et al. 2009; nuñez et al. 2013; olson et al. 2009). in order for this strategy to be implemented for fenced wildlife areas, one or more corridors may need to be established through predominantly rural communities that are likely to have either a limited history of co-existence with wildlife or one of conflicts caused by damage-causing animals (guerbois, chapanda & fritz 2012; pinter-wollman 2012). notwithstanding the social and economic challenges that need to be overcome to establish and maintain such corridors, several legal considerations predominate in this. of these, the risk associated with the loss in ownership of wildlife is paramount. in order for a fenced wildlife area to be incorporated into a corridor network, from a game perspective, the fence separating the wildlife area and the corridor would need to be removed. for south african wildlife areas, such action would result in its ‘sufficiently enclosed’ status falling away in terms of the game theft act. as such, the landowner would immediately lose ownership of the game in that the legal status of the game would have, on removing the interleading fence between the wildlife area and an unfenced corridor, revert to its res nullius status. the removal of all or part of a restraining fence to enable game and other wildlife to migrate in response to the impacts of climate change therefore stands as a significant threat to, at least, the economic integrity and tourism attractiveness of any wildlife area. conclusion facilitating the movement of wildlife to a more suitable habitat via corridors has been advocated in the literature as one of the key long-term management measures that can be considered to mitigate the impacts of climate change on wildlife. although this option is plausible from a conservation and academic perspective, it may have significant negative implications from a continued ownership of, and with that the legal protection afforded to, wildlife within fenced south african wildlife areas (e.g. game reserves, game ranches and private and communal protected areas). this article highlights the potential legal constraints facing landowners who intend to apply this mitigation measure. the game theft act 105 of 1991 overrides the res nullius common law status of game by conferring ownership on the landowner if the game in question have been adequately enclosed by a game fence. thus, in order for the wildlife areas mentioned above (i.e. those areas that have been fenced in a manner that fulfils the requirements of the game theft act) to enable game to move to a more suitable habitat, boundary fences would need to be removed or altered in a manner that would be permeable to affected wildlife. should the climate change corridor not be fenced in compliance with the act, the legal status of the game (those within the wildlife area and corridor) will instantly revert to its common law res nullius status, that is, owned by no one and available to be claimed by any person through purposefully taking physical possession of them. it is thus concluded that the game theft act should be adjusted by an amendment legislation, to keep pace with the challenges facing acquisition and loss of ownership of wildlife in a changing environment caused by climate change. a similar conclusion is advocated by freedman (2019) in his observation that the game theft act may not necessarily provide equivalent protection to state-protected areas as that afforded to private and communal wildlife areas. furthermore, as an interim measure, and until legislative amendments are implemented, the impacts of climate change may be decreased by a variety of management interventions, for example, a progressive expansion of the wildlife area as a means to include a suitable habitat. this action is likely to lessen the pressures inflicted by game that have been displaced by climate change on the area’s fenced boundaries. it would also strengthen legal arguments against the application of the common law principle of re-wilding to and the resultant res nullius status of game that have escaped from the wildlife area. this may require redefining the ‘ownership’ in a manner that allows for continued ownership of wildlife that may be displaced by the impacts of climate change, and an unambiguous separation of continued ownership of an ‘escaped animal’ and the loss of ownership through natural re-wilding. it is finally concluded that the practicality of recommendations eliminating out of biodiversity, and in particular wildlife research, cannot be assured without consideration of the legal context of the recommendations. acknowledgements the supportive environment of ezemvelo kzn wildlife is correct in as this is the wording of the registered trade name. wildlife and the university of kwazulu-natal are acknowledged. magda goosen of ezemvelo kzn wildlife is thanked for her comments on an earlier draft of this article. competing interests the author declares no conflict of interests with regard to the writing of this article. author contributions the author conceptualised the study, performed the principal analysis and wrote the manuscript. ethical consideration this article followed all ethical standards for a research without direct contact with human or animal subjects. funding information this research received no specific grant from any funding agency in the public, commercial or not-for-profit sector. data availability statement data sharing is not applicable to this article as no new data were created or analysed in this study. disclaimer the views and opinions expressed in this article are those of the author and do not necessarily reflect the official policy or position of any affiliated agency of the author. references agrawal, a. & redford, k., 2009, ‘conservation and displacement: an overview’, conservation and society 7(1), 1–10. barbier, e.b., hacker, s.d., kennedy, c., koch, e.w., stier, a.c. & silliman, b.r., 2011, ‘the value of estuarine and coastal ecosystem services’, ecological monographs 81(2), 169–193. https://doi.org/10.1890/10-1510.1 batllori, e., parisien, m.-a., parks, s.a., moritz, m.a. & miller, c., 2017, ‘potential relocation of climatic environments suggests high rates of climate displacement within the north american protection network’, global change biology 23(8), 3219–3230. https://doi.org/10.1111/gcb.13663 bellard, c., bertelsmeier, c., leadley, p., thuiller, w. & courchamp, f., 2012, ‘impacts of climate change on the future of biodiversity’, ecology letters 15(4), 365–377. https://doi.org/10.1111/j.1461-0248.2011.01736.x blackmore, a., 2015, ‘the relationship between the nema and the public trust doctrine: the importance of the nema principles in safeguarding south africa’s biodiversity’, south african journal of environmental law and policy 20(2), 89–118. blackmore, a.c., 2017, ‘public trust doctrine, research and responsible wildlife management in south africa’, bothalia 47(1), 1–9. https://doi.org/10.4102/abc.v47i1.2217 caro, t., jones, t. & davenport, t.r.b., 2009, ‘realities of documenting wildlife corridors in tropical countries’, biological conservation 142(11), 2807–2811. https://doi.org/10.1016/j.biocon.2009.06.011 carruthers, j., 2008, ‘wilding the farm or farming the wild? the evolution of scientific game ranching in south africa from the 1960s to the present’, transactions of the royal society of south africa 63(2), 160–181. https://doi.org/10.1080/00359190809519220 carter, r.w., walsh, s.j., jacobson, c. & miller, m.l., 2014, ‘in the george wright forum’, george wright society 31(3), 245–255. corlett, r.t. & westcott, d.a., 2013, ‘will plant movements keep up with climate change?’, trends in ecology & evolution 28(8), 482–488. https://doi.org/10.1016/j.tree.2013.04.003 couzens, e., 2019, international encyclopaedia of law: environmental law south africa, 2nd edn., p. 600, kluwer law international, the netherlands, isbn: 9789065449450. couzens, e. international encyclopaedia of law: environmental law south africa (2nd ed, 2019) kluwer law international, the netherlands. 600p isbn: 9789065449450 cushman, s.a., elliot, n.b., bauer, d., kesch, k., bahaa-el-din, l., bothwell, h. et al., 2018, ‘prioritizing core areas, corridors and conflict hotspots for lion conservation in southern africa’, plos one 13(7), e0196213. https://doi.org/10.1371/journal.pone.0196213 davidson, a.d., boyer, a.g., kim, h., pompa-mansilla, s., hamilton, m.j., costa, d.p. et al., 2012, ‘drivers and hotspots of extinction risk in marine mammals’, proceedings of the national academy of sciences 109(9), 3395–3400. https://doi.org/10.1073/pnas.1121469109 di minin, e., hunter, l.t.b., balme, g.a., smith, r.j., goodman, p.s. & slotow, r., 2013, ‘creating larger and better connected protected areas enhances the persistence of big game species in the maputaland-pondoland-albany biodiversity hotspot’, plos one 8(8), e71788. https://doi.org/10.1371/journal.pone.0071788 eastern cape parks and tourism agency v medbury (pty) ltd, (2018), 206, zasca34. freedman, w., 2019, ‘a critical analysis of the escape rule – eastern cape parks and tourism agency v medbury (pty) ltd t/a crown river safari 2018 4 sa 206 (sca)’, journal of south african law 2019(2), 374–382. groves, c.r., game, e.t., anderson, m.g., cross, m., enquist, c., ferdaña, z. et al. (2012). ‘incorporating climate change into systematic conservation planning’, biodiversity and conservation 21, 1651–1671. https://doi.org/10.1007/s10531-012-0269-3 guerbois, c., chapanda, e. & fritz, h., 2012, ‘combining multi-scale socio-ecological approaches to understand the susceptibility of subsistence farmers to elephant crop raiding on the edge of a protected area’, journal of applied ecology 49(5), 1149–1158. https://doi.org/10.1111/j.1365-2664.2012.02192.x guyon, f., 2018, ‘le massif des vosges face à un cas de “ré-ensauvagement” par le cerf: analyse de l’influence des facteurs sociaux (professionnels et culturels) sur les représentations et qualifications attribuées au cerf, à la forêt et aux activités des hommes’, vertigo-la revue électronique en sciences de l’environnement 18(3), 1–29. https://doi.org/10.4000/vertigo.23363 hannah, l., midgley, g., hughes, g. & bomhard, b., 2005, ‘the view from the cape: extinction risk, protected areas, and climate change’, bioscience 55(3), 231–242. https://doi.org/10.1641/0006-3568(2005)055[0231:tvftce]2.0.co;2 hartter, j., goldman, a. & southworth, j., 2011, ‘responses by households to resource scarcity and human–wildlife conflict: issues of fortress conservation and the surrounding agricultural landscape’, journal for nature conservation 19(2), 79–86. https://doi.org/10.1016/j.jnc.2010.06.005 hulme, p.e., 2005, ‘adapting to climate change: is there scope for ecological management in the face of a global threat?’, journal of applied ecology 42(5), 784–794. https://doi.org/10.1111/j.1365-2664.2005.01082.x johnson, m.f., karanth, k.k. & weinthal, e., 2018, ‘compensation as a policy for mitigating human-wildlife conflict around four protected areas in rajasthan, india’, conservation and society 16(3), 305–319. https://doi.org/10.4103/cs.cs_17_1 kamuti, t., 2014, ‘the fractured state in the governance of private game farming: the case of kwazulu-natal province, south africa’, journal of contemporary african studies 32(2), 190–206. https://doi.org/10.1080/02589001.2014.936678 koelble, t.a., 2011, ‘ecology, economy and empowerment: eco-tourism and the game lodge industry in south africa’, business and politics 13(1), 1–24. https://doi.org/10.2202/1469-3569.1333 koprowski, j.l. & krausman, p.r., 2019, international wildlife management: conservation challenges in a changing world, jhu press, baltimore. lamichhane, b.r., persoon, g.a., leirs, h., poudel, s., subedi, n., pokheral, c.p. et al., 2018, ‘spatio-temporal patterns of attacks on human and economic losses from wildlife in chitwan national park, nepal’, plos one 13(4), e0195373. https://doi.org/10.1371/journal.pone.0195373 landguth, e.l., hand, b.k., glassy, j., cushman, s.a. & sawaya, m.a., 2012, ‘unicor: a species connectivity and corridor network simulator’, ecography 35(1), 9–14. https://doi.org/10.1111/j.1600-0587.2011.07149.x lister, n.-m., brocki, m. & ament, r., 2015, ‘integrated adaptive design for wildlife movement under climate change’, frontiers in ecology and the environment 13(9), 493–502. https://doi.org/10.1890/150080 madden, f.m., 2008, ‘the growing conflict between humans and wildlife: law and policy as contributing and mitigating factors’, journal of international wildlife law & policy 11(2–3), 189–206. https://doi.org/10.1080/13880290802470281 magudu game company (pty) ltd v mathenjwa n.o. (2008) 2 all sa 338 (n) mawdsley, j.r., o’malley, r. & ojima, d.s., 2009, ‘a review of climate-change adaptation strategies for wildlife management and biodiversity conservation’, conservation biology 23(5), 1080–1089. https://doi.org/10.1111/j.1523-1739.2009.01264.x mcdowell, c., 2013, ‘climate-change adaptation and mitigation: implications for land acquisition and population relocation’, development policy review 31(6), 677–695. https://doi.org/10.1111/dpr.12030 minteer, b.a. & collins, j.p., 2010, ‘move it or lose it? the ecological ethics of relocating species under climate change’, ecological applications 20(7), 1801–1804. https://doi.org/10.1890/10-0318.1 monzón, j., moyer-horner, l. & palamar, m.b., 2011, ‘climate change and species range dynamics in protected areas’, bioscience 61(10), 752–761. https://doi.org/10.1525/bio.2011.61.10.5 muir, a., 2016, ‘offences, game and property – some unresolved issues surrounding ownership of wild animals in south africa’, stellenbosch law review 27(1), 136–160. newmark, w.d., 2008, ‘isolation of african protected areas’, frontiers in ecology and the environment 6(6), 321–328. https://doi.org/10.1890/070003 nuñez, t.a., lawler, j.j., mcrae, b.h., pierce, d.j., krosby, m.b., kavanagh, d.m. et al., 2013, ‘connectivity planning to address climate change: climate change and connectivity’, conservation biology 27(2), 407–416. https://doi.org/10.1111/cobi.12014 nyhus, p.j., 2016, ‘human–wildlife conflict and coexistence’, annual review of environment and resources 41, 143–171. https://doi.org/10.1146/annurev-environ-110615-085634 olson, d., o’connell, m., fang, y.-c., burger, j. & rayburn, r., 2009, ‘managing for climate change within protected area landscapes’, natural areas journal 29(4), 394–399. https://doi.org/10.3375/043.029.0406 pienaar, g., 2012, ‘the methodology used to interpret customary land tenure’, potchefstroom electronic law journal 15(3), 153–183. https://doi.org/10.4314/pelj.v15i3.7 pinter-wollman, n., 2012, ‘human–elephant conflict in africa: the legal and political viability of translocations, wildlife corridors, and transfrontier parks for large mammal conservation’, journal of international wildlife law & policy 15(2), 152–166. https://doi.org/10.1080/13880292.2012.678793 pittiglio, c., skidmore, a.k., van gils, h.a.m.j., mccall, m.k. & prins, h.h.t., 2014, ‘smallholder farms as stepping stone corridors for crop-raiding elephant in northern tanzania: integration of bayesian expert system and network simulator’, ambio 43, 149–161. https://doi.org/10.1007/s13280-013-0437-z richter v du plooy (1921), opt 117. rumsey, a.b., 2009, terrestrial wild animals, 2nd edn., h. strydom & n. king (eds.), juta, cape town. sax, d.f., smith, k.f. & thompson, a.r., 2009, ‘managed relocation: a nuanced evaluation is needed’, trends in ecology & evolution 24, 472–473. https://doi.org/10.1016/j.tree.2009.05.004 schwartz, m.w., hellmann, j.j., mclachlan, j.m., sax, d.f., borevitz, j.o., brennan, j. et al., 2012, ‘managed relocation: integrating the scientific, regulatory, and ethical challenges’, bioscience 62(8), 732–743. https://doi.org/10.1525/bio.2012.62.8.6 selier, s.j., slotow, r., blackmore, a. & trouwborst, a., 2016, ‘the legal challenges of transboundary wildlife management at the population level: the case of a trilateral elephant population in southern africa’, journal of international wildlife law & policy 19(2), 101–135. https://doi.org/10.1080/13880292.2016.1167460 songhurst, a., mcculloch, g. & coulson, t., 2016, ‘finding pathways to human–elephant coexistence: a risky business’, oryx 50(4), 713–720. https://doi.org/10.1017/s0030605315000344 syombua, j., 2013, ‘land use and land cover changes and their implications for human-wildlife conflicts in the semi-arid rangelands of southern kenya’, journal of geography and regional planning 6(5), 193–199. https://doi.org/10.5897/jgrp2013.0365 treves, a., 2009, ‘the human dimensions of conflicts with wildlife around protected areas’, in m.j. manfredo, j.j. vaske, p. brown, d.j. decker & e.a. duke (eds.), wildlife and society: the science of human dimensions, pp. 214–228, island press, washington, dc. van der merwe, c.g., 2002, ‘things’, in w.a. joubert (ed.), the law of south africa (lawsa), butterworths, durban. van der merwe, c.g. & rabie, m.a., 1974, ‘eiendom van wildre diere’, tydskrif vir hedendaagse romeins-hollandse reg 37, pp. 38–48. vos, a., clements, h.s., biggs, d. & cumming, g.s., 2019, ‘the dynamics of proclaimed privately protected areas in south africa over 83 years’, conservation letters 12(6), e12644. https://doi.org/10.1111/conl.12644 wilke, c. & rannow, s., 2014, ‘a methodical framework for climate change-adapted management in protected areas’, in s. rannow & m. neubert (eds.), managing protected areas in central and eastern europe under climate change, pp. 159–172, springer netherlands, dordrecht. abstract introduction methods results discussion conclusion acknowledgements references about the author(s) kudakwashe musengi school of animal, plant and environmental science, university of the witwatersrand, south africa sally archibald school of animal, plant and environmental science, university of the witwatersrand, south africa citation musengi, k. & archibald, s., 2017, ‘demographics of eucalyptus grandis and implications for invasion’, koedoe 59(1), a1437. https://doi.org/10.4102/koedoe.v59i1.1437 original research demographics of eucalyptus grandis and implications for invasion kudakwashe musengi, sally archibald received: 30 aug. 2016; accepted: 21 feb. 2017; published: 30 mar. 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract alien invasive species can have negative impacts on the functioning of ecosystems. plantation species such as pines have become serious invaders in many parts of the world, but eucalypts have not been nearly as successful invaders. this is surprising considering that in their native habitat they dominate almost all vegetation types. available theory on the qualities that characterise invasive species was used to assess the invasive potential of eucalyptus grandis – a common plantation species globally. to determine rates of establishment of e. grandis outside plantations, we compared population demographics and reproductive traits at two locations in mpumalanga, south africa: one at higher elevation with more frost. eucalyptus grandis has a short generation time. we found no evidence that establishment of e. grandis was limiting its spread into native grassland vegetation, but it does appear that recruitment is limited by frost and fire over much of its range in mpumalanga. populations at both study locations displayed characteristics of good recruitment. size class distributions showed definite bottlenecks to recruitment which were more severe when exposed to frost at higher elevations. generally, the rate of spread is low suggesting that the populations are on the establishing populations’ invasion stage. this research gives no indication that there are any factors that would prevent eucalyptus from becoming invasive in the future, and the projected increase in winter temperatures should be a cause for concern as frost is currently probably slowing recruitment of e. grandis across much of its planted range. conservation implications: eucalyptus plantations occur within indigenous grasslands that are of high conservation value. frost and fire can slow recruitment where they occur, but there are no obvious factors that would prevent e. grandis from becoming invasive in the future, and monitoring of its rates of spread is recommended. introduction the deliberate or unintentional introduction of new species by humans is a leading cause of the global biodiversity crisis (wilcove et al. 1998). invasive species are defined as naturalised alien plants outside their native range as a result of relocation and often have the ability to produce very large numbers of offspring at considerable distances from parent plants (richardson et al. 2000). the most aggressive invaders can spread far from parent plants and cover large areas. up until the last few hundred years, geographical barriers to immigration and emigration of species by mountains and oceans, led ecosystems to evolve in relative isolation. increased rates of human movement around the world saw the intentional introduction of alien species (wilcove et al. 1998). today’s spread and global trade of species, which are backbones of horticulture, agriculture and forestry, act as constant sources of alien invasive species. fixed transport infrastructures can also provide platforms for biotic interchange, for example, the panama and suez canals (wilson et al. 2009). when looking at aspects of invasions, it is useful to conceptualise processes that limit or facilitate invasions as an invader negotiating a series of barriers (blackburn et al. 2011; kruger, richardson & van wilgen 1986; richardson et al. 2000). there are many factors that affect the chances of a species moving from one stage of invasion to another. the invasion process has a series of stages, and in each stage, a species or population needs to overcome a barrier to proceed to the next stage (blackburn et al. 2011). failure by a species to overcome the barriers to proceed to the next stage reduces the chances of the species becoming invasive. one of the most fascinating questions in ecology that has been the focus of increased research efforts in the last 50 years is what controls invasiveness of alien species (richardson & pysek 2008). potential determinants of invasiveness that are frequently studied include introduction history and pattern, species traits, and ecological and evolutionary processes. the invasion probability of a species is usually high if it was introduced a long time ago (long residence time) as it may take time for introduced species to build up to the point that they start spreading (richardson et al. 2000). additionally, if a species is introduced in multiple locations, it can also dramatically affect the rate at which it spreads (richardson & pysek 2008). the number of individuals introduced is the most prevalent correlate of establishment success in non-native populations. studies have shown that the number of individuals introduced can lead to establishment success (blackburn et al. 2013; lockwood, cassey & blackburn 2009). species traits normally associated with invasiveness are short generation times, production of copious amounts of viable seeds and dispersal ability of the seeds (rejmánek 2000). the first two traits affect the rate at which new individuals can be produced, and the third trait affects the ease with which individuals spread into new environments. for example, eucalypts produce seeds that just fall from the tree and is only carried to very short distances by the wind. the very small seed sizes mean that fire is often required to clear competing vegetation and allow eucalypt seedlings to establish. both biotic (competition or predation) and abiotic factors can control the ease with which a species spreads into new environments. invasives are often released from natural predators or pests when introduced to new environments (kruger et al. 1986; richardson et al. 2000), but the extent to which the abiotic environment limits alien spread appears to depend on particular circumstances; therefore many invasive species have a larger climatic niche than in their natural habitat. this enemy release hypothesis is also a good example for arguable results of empirical tests, for example, maron and vilà (as cited in jeschke & heger 2014) found that exotic plants can attract native herbivores which negatively affected the survival of the alien plants. invasive species are a good ecological test of our understanding of vegetation distribution, and the relative roles of ecological versus evolutionary processes in determining a species range. size (or age) class distributions give information on the proportion of a population that is at different stages of growth and can give insights into regeneration levels and recruitment bottlenecks of a population. in terms of invasive species, they could give insights into which stage of invasion a population is at, and what is preventing them from invading. an inverse j-shaped curve – with more juveniles than adults and a smoothly declining function – is expected in a recruiting population (lehmann, prior & bowman 2009; moritz 2003), because it allows for mortality and competitive effects on the probability of progressing to the next size class. demographic bottlenecks are usually deduced when there is a very steep slope between classes, indicating that something is preventing transition from one class to the next. these demographic bottlenecks can also result in bimodal or unimodal size class distributions among plant populations (lehmann et al. 2009) when the bottleneck is released. a u-shaped (bimodal) curve has got slightly more juveniles than adults indicating some form of recruitment but there is a sharp decline in the frequency of the middle-aged class. this could indicate ‘cohort’ recruitment – where the probability of moving between size classes depends on particular environmental conditions that happen seldom. a unimodal curve shows the absence of juveniles indicating that there is no recruitment at all. this could be as a result of something preventing seedling establishment. the natural occurrence of eucalyptus grandis w. hill ex maiden is in australia from newcastle, new south wales northwards to bundaberg, queensland on lower slopes of fertile valleys and is usually found in open woodlands or grasslands fringing rainforests (crisp, cook & steane 2012). the altitude is mainly from sea level to 600 m, but up to 1100 m in the tropical north (nsw department of primary industry 2010). the mean maximum temperature of the hottest month across its distribution ranges from 24 °c to 30 °c, and the mean minimum temperature of the coldest month ranges around 7.3 °c – 19.4 °c. rose gum is well adapted to fire – it resprouts vigorously and maintains a seed bank in its canopy for recruiting after fire (booth 2012; williams & woinarski 1997). in queensland (northern australia) where e. grandis is native, fire is a recurring element where most of the landscape is burnt every year (booth 2012). eucalyptus grandis was brought to south africa in 1890, and by 1973, it represented about 75% of all eucalyptus species planted in south africa (daff 2000). south africa’s eucalyptus plantations cover a broad range of environments, particularly in the subtropical and the humid warmer temperate regions (otim 2008). eucalyptus grandis plantations cover a much wider range of climates in south africa than it does in australia. the plantation areas in south africa fall into high rainfall zones. the fast growth rate of e. grandis has resulted in e. grandis being the most extensively planted hardwood in these regions. however, the increased demand for hardwoods has resulted in the expansion of hardwoods into sites that are colder and drier, or warmer and drier, than the conditions e. grandis evolved in australia. we studied the invasion potential of e. grandis in south africa. eucalyptus grandis has not been nearly as successful in invading alien environments as other widely planted trees such as pines (forsyth et al. 2004; higgins & richardson 1998) and legumes. the reasons why eucalypts are not more invasive are not well understood but it is agreed that the limited dispersal of the seeds, high mortality of the seeds and lack of compatible mycorrhizal fungi make the eucalypts less invasive (lake & fisherman 2004). although most eucalypts are not currently highly problematic invaders in south africa, they are still listed as potential invaders in many national and international lists (dept. of agriculture, regulation no. 15, 2001; richardson, williams & hobbs 1994). eucalyptus grandis is a successful invader in the tzaneen area and in kwazulu-natal along rivers (versfeld, le maitre & chapman 1998). four major frost events in the forestry areas of south africa have occurred in the past 30 years, on average a frequency of one event every 7.5 years (sappi 2006). cold-damaged plantations are more vulnerable to fires and are prone to consequential damage through pest or disease attacks (swain & gardner 2003). if the frost is severe, the affected leaves wither and the shoots may die back. due to its sensitivity to cold, mean annual temperatures (mats) that are below 18 °c and altitudes above 1050 m are not suited for planting e. grandis, according to swain and gardner (2003). cold-tolerant species such as eucalyptus nitens or hybrids of eucalyptus dunnii and e. grandis are sometimes used at sites above 1200 m which are prone to frost and frequent snowfalls. the frequent frosts in the highveld of south africa might therefore be a barrier to the spread of e. grandis because seedlings and saplings are ‘top killed’ by frosts. most of this frost damage occurs in winter in the form of tip scorching or total scorching, depending on the severity and frequency of the frost (otim 2008). scorch forms on the leaves as an irregular browning, yellowing or bronzing of tissues along the margins and tips of leaves (integrated pest management 1997). in some ways, a frost event can be similar to a fire in its impacts on woody plants (whitecross et al. 2012). both fire and frost can damage above-ground biomass of small trees (‘top-kill’) and therefore act to prevent trees from growing to adults. although frost is rare in the australian range of e. grandis, fire is common, and the species resprouts vigorously and is able to survive top-kill. it is therefore not clear what effect frost (and fire) would have on the recruitment and spread of this alien species. this study used the theory on the qualities that characterise invasive species to assess the invasion potential of e. grandis with the help of a field study. the study had the following objectives: (1) to test whether e. grandis is expected to be a serious invader based on its reproductive traits and (2) to assess the impact of frost on the rate of spread of e. grandis into neighbouring vegetation from plantations. methods study area a field study was used to determine whether there is any indication that e. grandis is in fact invading from plantations in mpumalanga (figure 1). to test specifically for the role of frost in preventing e. grandis invasion, we chose sites that were similar in terms of their fire regimes, but were exposed to very different frost frequencies. five sites were selected near graskop at high elevation (1600 m a.s.l.) where they are exposed to frost (24° 55” s, 30° 46” e) with mean minimum temperature of 4.8 °c and the other five at a low elevation area (953 m a.s.l.) with infrequent frost near white river (25° 09” s, 31° 02” e) with mean minimum temperature of 7 °c. sites at graskop experience ‘heavy’ frost in more than 80% of the years, whereas in white river severe frost occurs less than 20% of the time (schulze & maharaj 2007). ‘heavy’ frost is when the temperature of ≤ 0 °c is recorded on a stevenson screen assuming that the temperature on the ground would be even lower. figure 1: site map showing sampling locations. the plantations have trees that can be as tall as 60 m and are being managed for timber. fire return periods at these sites were similar – on average once every 3–4 years (archibald et al. 2010). plantations are usually protected from wild fires by annual firebreaks to protect the plantations from grassland fires, and during this process, saplings are usually cleared. however, many plantations are no longer actively managed in this way, which will probably increase invasion probability. this neglected management provided an opportunity to assess natural rates of spread of eucalypts into surrounding natural grassland vegetation. we studied populations that were close to the plantation plots where no firebreaks had been made before. we identified sites that had not been actively managed for at least 8 years. we sampled five different populations at each location to account for variability related to topographic position and plantation age. the veld surrounding the plantation plots consisted of grass with a mixture of old and young pine trees – pinus patula. we did not measure the diameters and heights of the pine trees. the sampling was done over a period of 17 days, the first field work was carried out from 07 to 14 june 2013, and the second one was carried out from 19 to 29 november 2013. transect layout belt transects (5 m by 50 m) were laid out from the edge of the plantation into the natural veld in each of the naturalised populations that are next to the plantations. we ensured that we sampled pure e. grandis stands – not hybrid stands – as hybrids of e. grandis and e. dunnii are often used in colder parts of the country (k. padayachee [university of the witwatersrand], pers. comm., 2013). on each transect, the height, distance along the transect (figure 2) and stem diameter at 30 cm above the ground of all the e. grandis individuals encountered within 2.5 m either side of the transect, were measured and recorded. tree callipers were used to measure the stem diameters, and a 2.5-m pole was used to visually help estimate the tree heights. for trees that have multiple stems, the diameter for all the living stems was noted, and the average basal area was calculated and converted back to average stem diameter: [basal area = sum (pi × r2)] ÷ (total number of stems). figure 2: heights of the invasive eucalypts along the 50-m transects. (a) graskop and (b) white river. demographic study we assessed a minimum of 50 trees on each site. the locations’ starting and ending points of all transects, as well as the location of all e. grandis individuals, were recorded in the field on a handheld gps. the distances of the e. grandis individuals along the transects were also noted, which allowed us to calculate the population density at the site [number of trees/(transect length × transect width)]. if plants had resprouted, this was noted, as well as any signs of pests or pathogens. the presence of flowers or fruits on each tree was also recorded, so as to have an idea of the size when e. grandis starts to become reproductively mature (i.e. generation time). size structure studies in tree communities usually measure dbh (diameter at breast height), basal diameter (baker et al. 2005; coomes & allen 2007; wang, hao & zhang 2009) and tree height (helm & witkowski 2012; lehmann 2009). these are usually strongly correlated with each other, but in a situation where there is a great deal of top-kill they could sometimes vary. we assessed size structure both in terms of basal diameter (diameter at 30 cm) and tree height. the number of saplings and adult individuals was determined. saplings were defined as those individuals less than 3 m in height – which is generally considered the cut-off height for when saplings have escaped the fire trap (bond & midgley 1995). germination rates study a germination study was undertaken to assess the vigour of e. grandis seeds. the aim was to test how quickly eucalyptus seeds germinate and what proportion of their seeds is viable. seeds were harvested from mature fruits of a random selection of five trees in graskop and white river to be germinated in the lab. the seeds from different trees at each site were mixed together, and 20 seeds were placed on a petri dish (5 replicates for each location). two 90-mm sheets of filter papers were placed in the lid of the 90-mm petri dishes; the seeds were placed in the lid and moistened with distilled water. the base of the petri dish was used to seal, and the seeds were incubated at 25 °c. the number of seedlings germinating was counted at 24-h intervals and continued until there was no change in germination percentage for 10 days. a seed was considered germinated if it had radicle longer than 1 mm. the proportion of viable seeds (pv), germination index (gi, zanjan & asli 2012) and the time taken for 50% of the seeds to have germinated (t50) germination were computed by using the following formulas: where n is the final number of germinated seeds and s is the total number of seeds placed in each dish. where ti is the number of days after planting, ni is the number of seeds that germinated on day 1, and s is the total number of seeds placed in each dish (zanjan & asli 2012). where n is the final number of germinated seeds, ni and nj are the cumulative number of seeds germinated by adjacent counts at times ti and tj when ni < 0.5 n and nj > 0.5 n. the gi is a measure of viability and seed vigour that incorporates both rate and amount of germination (seednet 2014). seed lots with greater germination indices and which take a short time to germinate are considered to be more vigorous. statistical analysis a multiple logistic regression analysis was performed where flowering state (yes or no) was predicted by the two explanatory variables (height and location). we also looked for interactions between height and location, to see whether size of first flowering was influenced by the environmental conditions of the site. the bayesian information criterion (bic) was used to compare the models. for each site, we determined the height when 80% of the trees were reproductively mature, and using literature on growth rates of e. grandis at the two locations (daff 2000), we converted this height to an estimated generation time. statistical modelling was carried out using the software r (version 3.0.2). the data were not normally distributed so we used the non-parametric mann–whitney u test to analyse the difference in diameters, number of resprouting trees, number of trees affected by pests and densities between the locations, graskop and white river. comparison between the size class distributions of the e. grandis populations in graskop and white river was made using the kolmogorov–smirnov test. to give a quantitative basis for the comparison of population size structures, the weibull function was fitted to diameter frequency distribution at each site using the fbasics package in r. this weibull distribution has been frequently used to characterise tree size distributions (baker et al. 2005) because it can have a variety of different shapes (defined by the shape parameter) and can indicate the degree to which a distribution fits an inverse-j shape. the bayesian framework was used to fit the weibull model to diameter data from each site. the parameterisation, which was used for the probability density of the weibull distribution, is as follows: where diameter d > 0, b > 0 and c ≥ 0. the scale parameter (b) gives an indication of the characteristic diameter or height size of the population (it represents the 63.2 quantile). the shape parameter (c) defines the shape of the frequency distribution. when the value of c < 1, the function is falling sharply (monotonic function) showing that there are fewer juveniles than adults in the population; when near or equal to 1 (c = 1), it is a negative exponential distribution representing a healthy recruiting population with more juveniles than adults (everard 1995; moritz 2003). when c > 1, the function is unimodal and it shows lack of recruitment. when 1 < c < 3.6, the distribution will be having a positive skew (baker et al. 2005). the germination data were submitted to mann–whitney u test using r statistical package to determine if any of the measured parameters differed statistically between the graskop and white river populations. results generation time there was high variability in flowering height in the populations – some very small individuals (< 2 m) did have flowers or fruit, but other individuals greater than 20 m showed no signs of being reproductively active. for every one metre increase in height, the odds of having flowered increased by 1.18 [exp (0.16934)] overall. of the three logistic models considered, the one with the highest explanatory power included an interaction between height and location (i.e. the effect of height on flowering varied between graskop and white river). this model had a bic of 515 (table 1) which was an improvement on a model with height alone (bic 594). table 1: comparison three logistic models. location independent of height was not significant and had no effect on the explanatory power. therefore, although the average number of flowering trees in each location is similar, the height at which they start flowering at each site varies significantly (figure 3). figure 3: height at first flowering of graskop sites (black circles–dotted black line) and white river sites (dark grey solid circles–solid dark grey line). the graskop data have been shifted up slightly (using the r function ‘jitter’) to make the comparison clear. it appears that reproductive maturity occurs more quickly at white river than graskop (figure 3) with individuals ± 6 m tall having a 60% probability of being reproductively mature, whereas at graskop it took until ± 15 m for 70% of the individuals to be reproductively mature. on the contrary, at graskop a significant proportion (± 20%) of the very small trees (< 3 m) were already flowering. demographics trees in white river were taller than those found in graskop. the tall trees were found closer to the edge of the plantations at both locations, distance less than 10 m along the transects. these tall trees produced seeds thus spreading the eucalypts along the transects. the population densities in the sampling sites (from the edge of the plantation into the natural veld) ranged from 1000 stems/ha to 3378 stems/ha. there was no significant difference between the population densities in graskop and white river (mann–whitney u test: p = 0.64). resprouting was significantly different between sites (mann–whitney u test: p = 0.02) – with resprouts representing 46% of the population in graskop and only 11.6% in white river. for all locations, resprouting increased with altitude (r2 = 0.70, p = 0.02447; figure 4). figure 4: (a) boxplots showing the percentage of resprouts across the sites in graskop and white river. (b) relationship between altitude and percentage of resprouts. graskop and white river populations both had large numbers of individuals in the smallest stem diameter classes which follow the reverse-j curve, characteristic of good regeneration (figure 5). however, in graskop, there was a sharp step between the smallest and next smallest size class (67% of all individuals were < 5 cm) suggesting some sort of bottleneck. in white river, the distribution was smoother. graskop also had fewer old trees (diameter > 20 cm) in the ‘adults’ class compared to white river. figure 5: diameter size class distributions of all the populations in (a) graskop and (b) white river. using height instead of diameter to define the size class distributions gave similar patterns (figure 6). the step drop in individuals from the smallest size classes at graskop is even more apparent when considering height – with most of the individuals being less than 3 m tall. figure 6: height size class distributions of all the populations in (a) graskop and (b) white river. there was a strong relationship between height and diameter, as would be expected (figure 7). however, at graskop, there were some very small trees that still had significant diameters. these trees could be the result of several resprouting events after disturbance – where the tips and shoots of the sapling were damaged but the stem survived and continued to increase in size. figure 7: relationship between height and diameter in graskop (grey circles) and white river (solid black circles). although there is obviously strong recruitment at both locations, there was considerable variability in size class distributions between sites (figure 8) with both locations having some sites with strong reverse-j distributions, and others with unior bi-modal distributions. this indicates that various site-specific factors might also control the spread rates. four of the sites in graskop exhibited a negative exponential distribution (shape parameter not significantly different from 1) indicating healthy populations with good recruitment at those sites, whereas at site 2 the function was unimodal (c = 1.2) showing little recruitment as evidenced by few juveniles. figure 8: density plot showing the variability in diameter size class distributions between sites: (a) graskop sites and (b) white river sites (five sites for each location). two of the sites in white river had a negative exponential distribution (healthy and stable populations), whereas all the other sites had unimodal functions: site 1 (1.5, absence of juveniles), site 3 (1.1, little recruitment) and site 4 (2.1, no recruitment at all). there were no monotonic functions exhibited in white river. the (b) values confirm what was found previously, that is, trees in white river are larger on average than in graskop. shape parameter (c) values of the weibull distribution decreased with increasing altitude, indicating that the proportion of juveniles: adults were generally higher at the high-altitude sites. germination rates of all the e. grandis seeds harvested, 97% were viable and there was no significant difference between the proportion of viable seeds between the sites, graskop and white river. there was also no significant difference between the germination indices (figures 8b and 9a) in graskop and white river population (mann–whitney u test: p = 0.29). figure 9: (a) boxplots showing germination indices of graskop and white river seeds. (b) histogram showing the range of germination indices recorded in the 10 replicate samples (both white river and graskop are included here as there was no difference between sites). the time taken for 50% of the seeds to germinate ranged from 1.83 to 4 days (figure 10), but again there was no significant difference between the germination rates between the two sites (mann–whitney u test: p = 0.141; table 2). figure 10: histogram of the time taken to reach 50% germination. table 2: summary of the descriptive statistics of eucalyptus seeds germination. discussion in terms of reproductive traits assessed, e. grandis has a potential to be a serious invader. its seeds are viable, with short germination times, and no dormancy. the germination parameters of e. grandis are similar to those of other known invasive species such mikania micrantha which has a mean germination index of 2.5 (yang et al. 2005). at both sites, a significant proportion of the population was reproductively active by the time it was 5 m, and given that e. grandis can grow several meters a year (oballa et al. 2010), this is in line with the 2to 3-year generation times reported in the literature (nsw department of primary industry 2010). importantly, these generation times are in the same range of other known invasive species which are causing a lot of problems, for example, pinus patula can be reproductively mature in 2 years (barrett & mullin 1968), eucalyptus camaldulensis in 3 years (cab international 2000) and acacia mearnsii in 20 months (fao 2014). the demographics show that frost and other unknown demographic factors are slowing the rates at which e. grandis individuals mature. the high-altitude graskop site had a clear reverse-j-shaped curve, with a sharp drop in frequency from the 2 cm to 6 cm diameter size classes (the 2 m – 5 m height size classes) (see figures 4 and 5): evidence of a frost or fire trap which is supported by the substantial number (> 30%) of resprouting individuals in these populations. these results suggest that frost might be hindering the rate of spread. interestingly, we also showed that most of these resprouting individuals in graskop seem to delay reproduction for longer than individuals at white river – so not only is the rate of transition to adult size classes slower in graskop, but the generation time is longer and the total amount of seed produced probably is lower than in the white river populations. whether these factors might act to prevent invasion in the long term in high-altitude sites in south africa is not clear. considering the high number of juvenile individuals in both populations and lack of evidence of mortality in the resprouting individuals at graskop, these results suggest that the e. grandis populations are still at the establishing population stage of invasiveness (williamson 2006). however, unlike other invasive species, which often have a dense seedling layer of newly recruiting individuals (richardson et al. 1994), we recorded very few seedlings at our sites. thus, there could be a demographic bottleneck at the seedling establishment phase that was not considered in this study. conclusion about 7% of south africa’s water resources are being used by invasive alien trees and eucalypts are well known for using a lot of water (le maitre, versfeld & chapman 2000). also, there is a shift in recent years from growing eucalyptus as part of large commercial industries to having emerging farmers and subsistence farmers growing individual stands of eucalyptus on their properties as a means to earn extra income (sappi 2006). this means that e. grandis will now be able to spread more widely throughout the country and that its rates of spread into native vegetation will probably be less strictly controlled. given that one of the strongest predictors of invasion is the number of introduction events and the spatial distribution of these introduction events, this is likely to increase the probability that e. grandis could become invasive. in this study, we found evidence that frost and some other factors (table 3) limit the rate of spread of e. grandis into native vegetation. other factors like the limited seed dispersal of eucalypts might limit their spread. the seeds are dispersed by wind over quite short distances, and they have no adaptations for dispersal (wings or fleshy tissues). the lack of compatible ectomycorrhizal fungi can also be a factor limiting the spread. table 3: summary of results in relation to demographic stage – and which factors might be important but were not considered in this study. there have been suggestions that lack of ectomycorrhizal fungi may limit the spontaneous establishment of seedlings away from plantations (rejmánek & richardson 2011). however, in this study, we found absolutely no indication that there was a fundamental constraint on its ability to spread into native vegetation. for this reason, the new dispersed forestry industry that is developing around e. grandis should be a cause for concern. similarly, if the incidence of frost decreases with the projected increase in winter temperatures (engelbrecht, mcgregor & engelbrecht 2009), then more parts of the country will become suitable for rapid expansion of eucalyptus. this article demonstrates that looking at the demographics of alien populations can give clues about (1) the rates at which they might be spreading and (2) the factors preventing them from spreading. analysis of the population structure is the first step to gather information about the factors that might be affecting populations of alien species. a similar approach might well be used for other species of concern such as eucalyptus camaldulensis dehnh. however, this study also, demonstrates that the factors controlling transition between all demographic stages need to be considered to conclusively assess invasive potential. nonetheless, we show here that the chances of natural regeneration and spread of e. grandis have proven to be very high if the demographic bottlenecks are suppressed. protected areas such as the wolkberg centre of biodiversity which has red-listed plant species are already under threat and this can be worsened by the spreading of e. grandis once the demographic bottlenecks are suppressed. acknowledgements competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions s.a. was the project leader and did the experimental and project design. k.m. was also responsible for the project design and collected the data in the field. references archibald, s., scholes, r.j., roy, d.p., roberts, g. & boschett, l., 2010, ‘southern african fire regimes as revealed by remote sensing’, international journal of wildland fire 19, 861–878. https://doi.org/10.1071/wf10008 baker, p.j., bunyavejchewin, s., oliver, c.d. & ashton, p.s., 2005, ‘disturbance history and historical stand dynamics of a seasonal tropical forest in western thailand’, ecological monographs 75, 317–343. https://doi.org/10.1890/04-0488 barrett, r.l. & mullin, l.j., 1968, ‘a review of introductions of forest trees in rhodesia’, the rhodesia bulletin of forestry research, 1. blackburn, t.m., pysek, p., bacher, s., carlton, j.t., duncan, r.p., jarosik, v. et al., 2011, ‘a proposed unified framework for biological invasions’, trends in ecology and evolution 26, 333–339. https://doi.org/10.1016/j.tree.2011.03.023 blackburn, t.m., prowse, t.a., lockwood, j.l. & casey, p., 2013, ‘propagule pressure as a driver of establishment success in deliberately introduced exotic species: fact or artefact?’, biological invasions 15, 1459–1469. https://doi.org/10.1007/s10530-013-0451-x bond, w.j. & midgley, j.j., 1995, ‘kill thy neighbour: an individualistic argument for the evolution of flammability’, oikos 73, 79–85. https://doi.org/10.2307/3545728 booth, t., 2012, ‘eucalypts and their potential for invasiveness particularly in frost-prone regions’, international journal of forestry research 2, article id 837165. https://doi.org/10.1155/2012/837165 cab international, 2000, eucalyptu camaldulensis: forestry compendium global module, cab international, wallingford, uk. coomes, d.a. & allen, r.b., 2007, ‘mortality and tree-size distributions in natural mixed-age 430 forests’, journal of ecology 95, 27–40. https://doi.org/10.1111/j.1365-2745.2006.01179.x crisp, m., cook, l. & steane, d., 2012, ‘radiation of the australian flora’, philosophical transactions of the royal society of london. series b, biological sciences 359, 1551–1571. https://doi.org/10.1098/rstb.2004.1528 department of agriculture, conservation of agricultural resources act, 2001, (act number 43 of 1983), viewed n.d., from https://www.sanbi.org/sites/default/files/documents/documents/conservationofagriculturalresourcesact.pdf department of agriculture, forestry and fisheries (daff), 2000, viewed 11 october 2013, from http://forestry.daff.gov.za engelbrecht, f.a., mcgregor, j.l. & engelbrecht, c.j., 2009, ‘dynamics of the conformal-cubic atmospheric model projected climate-change signal over southern africa’, international journal of climatology 29, 1013–1033. https://doi.org/10.1002/joc.1742 everard, d.a., midgley, j.j. & van wyk, g.f., 1995, ‘dynamics of some forests in kwazulu-natal, south africa, based on ordinations and size-class distributions’, south african journal of botany 61(6), 283–292. https://doi.org/10.1016/s0254-6299(15)30548-2 fao, 2014, case studies of two invasive alien tree species in southern africa, viewed 08 march 2014, from http://www.fao.org/docrep/005/ac846e/ac846e09.htm#topofpage forsyth, g.g., richardson, d.m., brown, p.j. & van wilgen, b.w., 2004, ‘a rapid assessment of the invasive status of eucalyptus species in two south african provinces’, south african journal of science 100, 75–77. helm, c.v. & witkowski, e.t.f., 2012, ‘characterising wide spatial variation in population size structure of a keystone african savanna tree’, forest ecology and management, 263, 175–188. higgins, s.i. & richardson, d.m., 1998, ‘pine invasions in the southern hemisphere: modelling interactions between organism, environment and disturbance’, plant ecology 135(1), 79–93. https://doi.org/10.1023/a:1009760512895 integrated pest management, 1997, leaf scorch of woody plants, viewed 02 december 2013, from http://ipm.illinois.edu/diseases/series600/rpd620/ jeschke, j.m. & heger, t., 2014, ‘the enemy release hypothesis as a hierarchy of hypotheses’, oikos 123, 741–750. https://doi.org/10.1111/j.1600-0706.2013.01263.x kruger, f.j., richardson, d.m. & van wilgen, b.w., 1986, ‘processes of invasion by alien plants’, in i.a.w. macdonald, f.j. kruger & a.a. ferrar (eds.), the ecology and management of biological invasions in southern africa, pp. 145–155, oxford university press, cape town. lake, j.c. & leishman, m.r., 2004, ‘invasion success of exotic plants in natural ecosystems: the role of disturbance, plant attributes and freedom from herbivores’, biological conservation 117, 215–226. https://doi.org/10.1016/s0006-3207(03)00294-5 lehmann, c.r.e., prior, l.d. & bowman, d.m.j.s., 2009, ‘fire controls population structure in four dominant tree species in a tropical savanna’, oecologia 161(3), 505–515, viewed 10 march 2014, from http://www.jstor.org/stable/40310158?origin=jstor-pdf le maitre, d.c., versfeld, d.b. & chapman, r.a., 2000, ‘the impact of invading alien plants on surface water resources in south africa: a preliminary assessment’, water sa 26(3), 397–408. lockwood, j.l., cassey, p. & blackburn, t.m., 2009, ‘the more you introduce the more you get: the role of colonization pressure and propagule pressure in invasion ecology’, diversity and distributions 15, 904–910. https://doi.org/10.1111/j.1472-4642.2009.00594.x moritz, a.m., 2003, ‘spatiotemporal analysis of controls on shrubland fire regimes: age dependency and fire hazard’, ecological society of america 84, 351–361. https://doi.org/10.1890/0012-9658(2003)084[0351:saocos]2.0.co;2 nsw department of primary industry, 2010, eucalyptus, viewed 04 november 2013, from http://www.dpi.nsw.gov.au/primefacts oballa, p.o., konuche, p.k.a., muchiri, m.n. & kigomo, b.n., 2010, facts on growing and use of eucalyptus in kenya, kenya forestry research institute, nairobi, viewed 02 november 2013, from http://www.kefri.org/eucalyptus_facts.pd otim, c.k., 2008, ‘investigation of growth potential of alternative eucalyptus species for mid and high altitude sites in the summer rainfall region in south africa’, msc thesis, school of biochemistry, genetics, microbiology and plant pathology, university of kwazulu-natal. rejmánek, m., 2000, ‘invasive plants: approaches and predictions’, austral ecology 25, 497–506. https://doi.org/10.1046/j.1442-9993.2000.01080.x rejmánek, m. & richardson, d.m., 2011, ‘eucalypts’, in d. simberloff & m. rejmanek (eds.), encyclopaedia of biological invasions, pp. 203–208, university of california press, berkeley, ca. richardson, d.m. & pyek, p., 2008, ‘fifty years of invasion ecology: the legacy of charles elton’, diversity and distributions 14, 161–168. richardson, d.m., pyek, p., rejmánek, m., barbour, m.g., panetta, f.d. & west, c.j., 2000, ‘naturalization and invasion of alien plants: concepts and definitions’, diversity and distributions 6, 93–107. https://doi.org/10.1046/j.1472-4642.2000.00083.x richardson, d.m., williams, p.a. & hobbs, r.j., 1994, ‘pine invasions in the southern hemisphere: determinants of spread and invadability’, journal of biogeography 21, 511–527. https://doi.org/10.2307/2845655 sappi, 2006, tree farming guidelines for private growerspart 2 silviculture, viewed 30 september 2013, from http://www.sappi.com/regions/sa/sappisouthernafrica/sappi%20forests/tree%20farming%20guidelines/part%202_silviculture_chapter%201_species%20selection.pdf schulze, r.e. & maharaj, m., 2007, frequency of frost occurrences. south african atlas of climatology and agro hydrology, water research commission, pretoria, rsa, wrc report 1489/1/06, section 9.3. seednet, 2014, handbook of seed testing, viewed 19 april 2014, from http://seednet.gov.in/material/handbook_of_seed_testing/chapter%2015.pdf swain, t.l. & gardner, r.a.w., 2003, a summary of current knowledge of cold tolerant eucalypt species (cte’s) grown in south africa, icfr bulletin 03/2003, institute for commercial forestry research, pietermaritzburg. versfeld, d.b., le maitre, d.c. & chapman, r.a., 1998, alien invading plants and water resources in south africa: a preliminary assessment, wrc report no tt 99/98, water research commission, pretoria. wang, x., hao, z. & zhang, j., 2009, ‘tree size distributions in an old-growth temperate forest’, oikos 118, 25–36. https://doi.org/10.1111/j.0030-1299.2008.16598.x whitecross, m.a., archibald, s. & witkowski, e.f.t., 2012, ‘do freeze events create a demographic bottleneck for colophospermum mopane?’, south african journal of botany 83, 9–18. wilcove, d.s., rothstein, d., dubow, j., phillips, a. & losos, e., 1998, ‘quantifying threats to imperilled species in the united states’, bioscience 48, 607–615. https://doi.org/10.2307/1313420 williams, j. & woinarski, j.c.z., 1997, eucalypt ecology: individuals to ecosystems, cambridge university press, united kingdom. williamson, m., 2006, ‘explaining and predicting the success of invading species at different stages of invasion’, biological invasions 8, 1561–1568. wilson, j.r.u., dormontt, e.e., prentis, p.j., lowe, a.j. & richardson, d.m., 2009, ‘something in the way you move: dispersal pathways affect invasion success’, trends in ecology and evolution 24, 136–144. https://doi.org/10.1016/j.tree.2008.10.007 wits university department of geography. mpumalanga region [map], 2013, scale 1:60,000, st. catharines, wits university department of geography, on. yang, q., wan-hui, y., xiong, d., hong-ling, c., yun, z. & kai-yan, x., 2005, ‘seed germination eco-physiology of mikania micrantha, h.b.k’, botanical bulletin academia sinica 46, 293–299. zanjan, g.m. & asli, d.e., 2012, ‘seed germination and early seeding growth of wheat genotypes affected by different seed pyridoxine-priming duration’, annals of biological research 3(12), 5687–5691. power.qxd prey selection of lions panthera leo in a small, enclosed reserve r.j. power power, r.j. 2002. prey selection of lions panthera leo in a small, enclosed reserve. koedoe 45(2): 67–75. pretoria. issn 0075-6458. annual trends in numbers of ungulate species on a 15 km² reserve from 1993 to 1998, were evaluated in the context of lion panthera leo reintroduction during 1996, and subsequent predation by them. the ungulate prey base was enumerated annually by aerial counts and a road count that took place during 1998. the lion prey record was obtained from direct observations of a radio-located pride of eight lions and daily reserve management records. all ungulate species that underwent precipituous declines were also the most important prey to lions, comprising over 80 % of their prey, and they were preyed upon according to their availability. lion predation was causal for the declines in wildebeest connochaetes taurinus, blesbok damaliscus pygargus phillipsi and warthog phacochoerus africana, while the decline in kudu tragelaphus strepsiceros was only partly ascribed to lions, as other non-lion related mortality sources were identified. the only ungulate species to increase subsequent to lion reintroduction was the impala aepyceros melampus, which was furthermore under-selected by lions. the uncontrolled population growth of impala could have elicited ecological degradation, and it was advised to either not stock impala, or otherwise control their numbers if lions are unable to do so. lion hunting success and kill rate, were 21 % (n = 63) and 1 kill/4.4 days, respectively. three bushpigs potamochoerus larvatus were killed but not utilised, and this finding is corroborated by an intensive study in kwazulu-natal, and this aversion is discussed. predators can cause unprecedented declines of their prey where the prey are confined to small reserves that have no refuge from predation. on an annual basis, prey may need to be augmented to sustain predators on small reserves. key words: predation, predator-prey relationships, small population management, small reserves, wildlife management. r.j. power, centre for wildlife management, university of pretoria, pretoria, 0002 republic of south africa. issn 0075-6458 67 koedoe 45/2 (2002) introduction predators are unable to regulate their prey where their prey populations undergo seasonal migrations of varying degrees (sinclair et al. 1985; mills & retief 1984; mills & shenk 1992). however, where prey populations are resident, the predator can regulate and even limit its prey populations (hirst 1969; smuts 1978; fryxell et al. 1988; caughley & sinclair 1994; sinclair 1995; harrington et al. 1999; peel & montagu 1999). where the prey are at very low densities, the predator can even eliminate the prey (fryxell et al. 1988). analogous to this, enclosed reserves become islands in which the prey are compelled to be resident, and depending on the number of predators, the prey can be regulated in the same way. on a managed reserve of this nature, with lions panthera leo, the impact that predation can have on the prey base is an important consideration (mills 1991; van schalkwyk 1994; viljoen 1996; van dyk 1997; hunter 1998). to address this, one requires annual prey numbers at least, and a record of what the lions prey upon (mills 1991; van schalkwyk 1994; hunter 1998). however, to accurately and unequivocally demonstrate that a predator has had an impact on the prey; one requires additional information on the power.qxd 2005/12/09 10:10 page 67 demography of the prey populations; other non-predation related mortality sources in the dynamics of the prey populations; and how the predator selects for species, gender, age or condition (mills 1991; hunter 1998). this paper aimed to investigate lion prey selection in southern africa’s smallest reserve containing free-ranging lions. this reserve was the madjuma lion reserve (mlr), that was a part of the greater mabula game reserve. the surface area size was 15 km², which was even smaller than the smallest extensively managed reserves (< 1 000 km²) that were under review by van schalkwyk (1994). on fairly small reserves (40–140 km²), predators like lion and cheetah acinonyx jubatus, have caused drastic declines in their preferred prey (hunter 1998; peel & montagu 1999). hunter (1998) pointed out that it remains to be seen whether predators can cause extinctions in their prey in a small reserve. contrary to the difficulties of studying a typical african multi-predator system (smuts 1978; mills 1991; bothma 1997), this reserve with one notable predator, was expected to yield a less complex understanding of a predator-prey relationship. various aspects of predation such as hunting success, kill rates and prey species selection were examined in the light of prey abundance, and inference was to be made on the impact of this predation on prey populations. in 1998, the mlr pride consisted of eight lions, which were borne from a reintroduced stock of three lionesses and an adult male that came from the pilanesberg national park (pnp). the lionesses were born in pnp, of which these lions originally hailed from etosha national park (enp), and the extent to which they behave like them (stander 1992a; 1992b; stander & albon 1993) would be investigated in conjunction with the main objectives. one of the females already had a litter of four cubs born at the beginning of june 1997. the sex ratio was 3:1, males to females. in may 1998, the roster was brought to 10 when two cubs were born. study area madjuma lion reserve is situated in the limpopo province, south africa, between latitudes 24º40'–24º44's, and longitudes 27º57'–27º59'e. the underlying geology is comprised mainly of granitic rock types, as well as sandstone and metamorphic rock, that are, in turn, overlain by six soil types (bredenkamp & van rooyen 1990). the annual rainfall varies between 300 mm and 900 mm, with a mean of 602 mm per annum (south african weather bureau: rooiberg station). the vegetation type of the region is mixed bushveld (van rooyen & bredenkamp 1996), which is dominated by combretum apiculatum–woodland and smaller patches of old lands grassland undergoing succession with cynodon dactylon and cenchrus ciliaris (bredenkamp & van rooyen 1990). the only other large carnivore present is the brown hyaena hyaena brunnea (power 1998), which is not known to be an active predator, as it is mostly a scavenger (mills 1990). the reserve contains 10 ungulate species, of which nine can be regarded as prey, and serve that purpose to varying degrees. the only function of the mlr is tourism, where rangers escort guest clientele from the mabula game lodge to see the lions. methods the study period during 1998 was 95 days in total (power 1998), and that from which data was available from in 1997 was 285 days (after jakoby 1997). during 1998, the study period spanned intermittent intervals of two weeks between march and october. during 1997, data was made available outside the researcher’s study period too, hence the longer effective study period. radio-located lions within the pride were followed in a landrover and directly observed in the same way as other workers had done (schaller 1972; bryden 1978; elliot & cowan 1978; mcbride 1984; van orsdol 1982; 1984; packer et al. 1990; scheel & packer 1991; stander 1992a; b; stander & albon 1993; mills & shenk 1992; scheel 1993; viljoen 1993; mills 1996; hunter 1998; funston et al. 2001). four lions were fitted with 148 mhz radio-collars, with a 0.25 wave antennae (mk6, telonics, arizona). the radio signals were received by a portable receiver (telonics, arizona) koedoe 45/2 (2002) 68 issn 0075-6458 power.qxd 2005/12/09 10:10 page 68 connected to a hand-held two-element yagi antennae. the radio-collared individuals were members of the four different subgroups of the pride that frequently split apart from one another (power 1998). observations were undertaken daily in a non-random fashion, which is perceived as biased (mills 1996), but regarded as unimportant, as the lions and their kills could be readily located and it was assumed that all prey killed during the study period could be tallied. nocturnal observations ranged between short-duration observations up to six hours, and long-duration observations which were at least 12 hours, and were dependent on the likelihood that lions would kill (cf. mills 1996). the adult females were the focal animals as they were regarded as the hunting lions (van orsdol 1982; mills & shenk 1992), and all their activities were observed. a coleman spotlight (300 000 cp) was used, with an attached red-filter (stander 1992a; viljoen 1993) to observe the lions at night, but was switched off when the lions hunted (mills & shenk 1992). observations were made with a pair of leica 10 x 42 ba binoculars, at distances up to 400 m. all kills were recorded, including the species, sex and age where possible. it was not always possible to gather the last two characteristics under direct observations, and where that was the case, this information was obtained from returning to the carcass the following day. known ageing and sexing criteria were used (mcbride 1984; viljoen 1993; bothma 1996) to assign three age-classes: adults, subadults and juveniles. the reserve management provided information on kills made from their daily monitoring. in some cases the age and sex were not recorded and hence they were assigned as unknown. data collected from the reserve management in the previissn 0075-6458 69 koedoe 45/2 (2002) ous year and that collected by researchers in 1997 (jakoby 1997) and 1998 (power 1998) was used, and re-analysed for this paper. lion hunting success was determined as the number of kills made per species, out of all hunting attempts for all prey species grouped (van orsdol 1984; stander & albon 1993; funston et al. 2001). a hunt was defined according to schaller (1972), and the outcome of a hunt ended in prey being killed or otherwise fleeing upon detecting hunting lions. when examining prey species selection, overt selection was inferred when killed proportions of a certain species exceeded the proportions with which they occurred in, in the study area (karanth & sunquist 1995). the road-count numbers of the five most common ungulates, and the prey numbers killed by lions during 1998 (power 1998) were used fig. 1. line graphs showing long-term trends in the ungulate and lion numbers of mlr since 1993. lion reintroduction took place in 1996. the blesbok damaliscus pygargus phillipsi numbers were supplemented during 1996. to calculate the expected and observed number of prey killed respectively. the log-likelihood ratio or g-test statistic (zar 1986) was used to detect prey species selection. where a difference was found, bonferroni simultaneous confidence intervals were proceeded to, as in other use-availability data (neu et al. 1974; byers et al. 1986). the kill rate by hunting lionesses (van orsdol 1982; mills & shenk 1992) over the specified study period was determined as the total number of kills/study period (days). this was done for each species and extrapolated over a one year period. results based on the aerial census figures, the trend was for most ungulate prey populations to grow prior to lion reintroduction, and then decline subsequent to lion reintroduction. the impala aepyceros melampus numbers however increased, while the warthog phacochoerus africana numbers remained relatively stable with minor fluctuations (fig. 1). over the period (1997–1998), 86 lion kills were recorded (table 1), 87 % were compower.qxd 2005/12/09 10:10 page 69 koedoe 45/2 (2002) 70 issn 0075-6458 table 1 summary of all lion kills (n = 86) made from march 1997 to october 1998 which include gender and relative age characteristics of prey prey species gender totals males females unknown adult subadult juvenile ad. subad. juv. ad. subad. juv. wildebeest 13 2 12 4 31 blesbok 5 6 2 1 14 kudu 8 3 11 warthog 4 9 1 4 18 impala 1 1 1 3 bushpig 3 3 genet 1 1 porcupine 1 1 aardvark 1 1 2 ostrich 2 2 table 2 summary of total mortality percentages of five ungulates on mlr, inclusive of all lion kills and other mortality sources and the annual kill rate prey species % lion kills in % lion kills in other non-lion related per annum 1998 (n = 30) 1997 (n = 55) mortality (1997), kill rate as % of total mortality wildebeest 40.0 36.4 16.7 (n = 24) 30.7 blesbok 10.0 18.2 9.1 (n = 11) 13.5 kudu 16.7 10.9 72.7 (n = 22) 10.6 warthog 20.0 20.0 0 (n = 11) 13.6 impala 0.0 5.4 0 (n = 3) 2.8 prised of four ungulate species, and 93 % were prey between 50 and 300 kg. wildebeest connochaetes taurinus comprised 37 %, warthog 21 %, kudu tragelaphus strepsiceros 13 %, and blesbok 16 %. a small spotted genet genetta genetta and three bushpigs potamochoerus larvatus were not utilised. the one bushpig was though partially eaten, but the lions regurgitated the meat subsequently. anomalous prey like porcupine hystryx africaeaustralis, aardvark orycteropus afer and ostrich struthio camelus were also recorded. male individuals comprised 55 % (n = 66) of kills where gender was known, while adults comprised 81 % of all prey, while younger individuals (subadults & juveniles) made up the remainder (table 1). there were a total of 63 hunts observed, of which 13 were successful. therefore hunting success was 21 % for all prey species. the kill rate was one kill every 4.4 days, implying that 83 kills were made annually, regardless of whether they were utilised or not. the annual off-take by lions for the prey animals was calculated (table 2). other non-lion related mortality (table 2) sources included fence electrification, old age, and various unidentified sources, which could have included food shortages. excluding the small recruitment of the year for each species (madjuma census figures 1997), the calculatpopo uma ures ht © ve etc. nk it mabend. djuma game power.qxd 2005/12/09 10:10 page 70 ed per annum kill rate does conform to the same number at which ungulate populations declined by (table 2) suggesting that predation over-rode the net growth rate there was an overall significant degree of prey species selection by the mlr lions from the road count data (g = 47, df = 4, p < 0.001). owing to this, it was decided to proceed to construct bonferroni confidence intervals and further analyse this claim. it was found that the mlr lions appeared to prey on four ungulate prey species according to their numbers in the populations, and impala were significantly under-utilised (table 3). discussion as predicted by predation theory (caughley & sinclair 1994), small resident prey populations, when subject to predation, without refuge from predators, or where migration is inhibited (fencing), can undergo significant declines (fryxell et al. 1988; mills 1991; sinclair 1995; hunter 1998; peel & montagu 1999). at mlr, the four ungulate species that underwent dramatic declines, were also the most important prey to lions, comprising over 80 % of lion kills. it was imminent that some ungulate populations could have become extinct within a year (fig. 1), which remains to be seen on a small reserve, as hunter (1998) pointed out. in identifying the cause of prey declines, it is imperative to also account for other sources of mortality, other than predation (mills 1991; hunter 1998), and accordingly, all non-lion related mortality was recorded for the mlr (table 2). for wildebeest and blesbok, nonlion related mortality accounted for a small fraction (< 17 %) of total mortality, and in warthogs no other source of mortality could be identified other than lion predation (table 2). however, in the case of kudu, nonlion related mortality was much greater (> 70 %) than that contributed by predation per se. it is thus argued that lion predation was the main cause of population declines in wildebeest, blesbok and warthog, while predation might have had a secondary role in hastening the decline in the kudu population. in kudu, mortality is often associated with dry season food shortages (owen-smith 1984) and the consequences of dimorphism in the long-term, particularly in adult males, which may be further exacerbated by predation (owen-smith 1993). lion predation was aimed at the adult segment of ungulate populations, moreover juvenile predation, which is most certainly because the 1998 study period (power 1998), fell outside the main ungulate seasonal birth peak. alternatively, the study methodology could have been the reason. prey like warthogs and juvenile ungulates are rapidly consumed (8 min for juvenile warthog: power 1998) and a study of this nature might have overlooked the fraction of smaller prey (mills 1996). elsewhere, higher juvenile predation by lions was reported, which was preissn 0075-6458 71 koedoe 45/2 (2002) table 3 prey species selection by lions in the mlr based on the 1998 road count and kill record, using bonferroni simultaneous confidence intervals. po = observed use proportion, pio = expected use proportion, n = kill sample, total z = 2.34, selection status: -ve = avoidance, +ve = overt selection, 0 = selection in accordance with numbers prey species po pio n bonferroni intervals selection (1998 road counts) status wildebeest 0.460 0.44 12 -0.06 < pio < 0.98 0 blesbok 0.115 0.104 3 -0.22 < pio < 0.45 0 kudu 0.192 0.061 5 -0.02 < pio < 0.37 0 warthog 0.231 0.100 6 -0.21 < pio < 0.67 0 impala 0.000 0.288 0 -0.2 < pio < 0.20 -ve power.qxd 2005/12/09 10:10 page 71 sumably because of the continuous presence of observers with study lions (mcbride 1984; viljoen 1993; stander 1992a). notwithstanding, if adult mortality was important at mlr, it could have been a contributor to the observed population declines, as lion predation was simulated to have the greatest impact in the knp when the adult segment of the zebra (mills & shenk 1992) and roan (harrington et al. 1999) populations were preyed upon. the mlr lions favoured wildebeest, warthog and kudu as prey, like they do in savannas elsewhere (mills 1991), that was in accordance with their numbers (table 3). this would however be the first published record of blesbok as lion prey, as lions have been eliminated from the blesbok’s distribution range, though this species is non-native to the limpopo province bushveld (skinner & smithers 1990). the absence of zebra predation was a reflection of their numbers, and it is suggested that they should be stocked on account of them being an important prey species (pienaar 1969; whateley & brooks 1985; mills 1991; mills & shenk 1992; mills & shenk 1992; scheel 1993; stander & albon 1993; viljoen 1993; funston et al. 2001). the fact that bushpig were not utilised, and when so, regurgitation took place can be corroborated with the findings of hunter (1998) at phinda. he reported 10 out of 13 bushpig kills that were abandoned and only partially fed upon (hunter 1998). boma-contained lions have also been known to show disdain to eating bushpig meat (van dyk pers. comm.). two reasons are put forward why this could be so. firstly, bushpig might have glandular secretions that may impart some distastefulness to the flesh, which would be to the disdain of the eater and explain why lions regurgitate this once eating. secondly, lions could be construed as ‘finicky’ carnivores that feed only on true herbivores on the appropriate secondary trophic level. the bushpig is more omnivorous than the warthog (skinner & smithers 1990), and it could be likened to another carnivore, which is infrequently preyed upon (cf. leopards). however, nutritionallystressed lions, or any other predator for that matter, would not have an aversion to feeding on non-herbivore prey (griffiths 1975; eloff 1984; mills 1984; skinner & smithers 1990). this phenomenon has largely been overlooked as bushpig are either rare or absent from many of the sites where lions have been intensively studied. only a fullyfledged research study on the myochemistry of bushpig meat could properly elucidate this answer. the mlr hunting success is intermediary between that reported for enp (15 %: stander & albon 1993) and other areas such as the r’wenzori national park (28.9 %: van orsdol 1984). hunting success was highest on dark moonless nights (power 1998), which is due to the lessened ability of prey in detecting hunting lions (van orsdol 1984; stander & albon 1993; funston et al. 2001). contrary to enp lions (stander & albon 1993), mlr lions had exercised some diurnal hunting with fairly high success (power 1998), which is perhaps because of the closed nature of vegetation (funston et al. 2001), and the availability of diurnal warthog, which are a main contributor to diurnal hunting success in lions (van orsdol 1984). the extent to which the mlr lions hunted with a coordinated strategy of enp lions (stander 1992b), could not be properly ascertained owing to the conditions of the vegetation and darkness. for those that were witnessed (n = 6) during daylight, the largest lioness consistently occupied a central position in all hunting, whilst the two smaller lionesses were wings that encircled the prey as described by stander (1992b). a kill rate of seven wildebeest per killing lioness per annum for knp (mills & shenk 1992; funston 1999) is lower than the 10.2 wildebeest per lioness per year for mlr, which is perhaps because of the limitations in prey switching on mlr when compared to the more diverse knp prey base. mlr lions exhibited no significant selection for any species, except for impala. non-selective predation patterns occur where large prey are scarce (karanth & sunquist 1995), and where prey choice is seemingly limiting (griffiths 1975). at phinda, wildebeest were koedoe 45/2 (2002) 72 issn 0075-6458 power.qxd 2005/12/09 10:10 page 72 preyed upon at three times their availability, warthogs twice, and kudu at their availability, while impala were under-selected (hunter 1998) like the mlr. it appears that impala are more frequently preyed upon in larger reserves (bryden 1978; whateley & brooks 1985; mills & biggs 1993; funston et al. 2001), and the fact that they were significantly under-represented in the diet of mlr lions, might be ascribed to the extreme alertness and superior vigilance behaviour of this species (see mooring 1999). at phinda, impala underwent a 200 % increase in vigilance (measured as the proportion of individuals being vigilant) following felid reintroduction (hunter & skinner 1997), which suggests that the slightly higher predation upon them during 1997 (jakoby 1997), was because impala were more naïve to the presence of lion. with a constrained and below average pride range size (ca. 15 km²), it is contended that the encounter rate with impala would be much higher than it would be in a larger reserve. in larger reserves, then, larger lion prides frequently range out of the ranges of impala, and the impala thus have a reprieve in awareness for lions. with the higher predator-prey encounter rates experienced on smaller reserves, it could be the primer to ‘fine-tune’ their senses to a familiar predator. it is hypothesised then that impala will feature less in the diet of lions when their pride ranges are small, whether this is because of confinement or ecological reasons. since impala were infrequently preyed upon they would not be serving their purpose as prey. despite this, what is more at stake, though, is that an ever increasing abundance of impala could severely impact upon the vegetation and other herbivore habitats. impala are highlyselective mixed feeders (fairall & klein 1984; du toit 1988; skinner & smithers 1990), that have the ability to denude the amount and change the composition of the herbaceous layer to the detriment of other herbivores (wentzel et al. 1991), and reduce predator stalk cover (power 1998), which is an important variable influencing hunting success (kruuk 1986; sunquist & sunquist 1989; van orsdol 1984; funston et al. 2001). following this, for ecological reasons, managers should closely monitor impala numbers, and intervene to control impala numbers if lions are unable to do so. impala appear to successfully utilise a habitat that can be construed as ‘enemy-free space’ (jeffries & lawton 1984; begon et al. 1996), which would not be favoured by potentially competing herbivores (wentzel et al. 1991), or otherwise not conducive to the hunting antics of lions, owing to the reduction in stalk cover (van orsdol 1984), which is typical of impala habitat (wentzel et al. 1991). prey like wildebeest, kudu, warthog, zebra and blesbok would serve effective buffer prey species, if there is a desire to safeguard rare or expensive ungulate populations. wildebeest are especially a favoured prey species, and lions are known to have a great impact on their populations in both large and small reserves (mills & shenk 1992; mills & biggs 1993; hunter 1998; peel & montagu 1999). on small reserves (< 2 000 ha) managers might be compelled to restock reserves with prey or otherwise increase area size through the conservancy approach, and if that is not an option, to not liberally stock small reserves with lions. large protected areas that create prey confinement by water provisioning can be prone to prey declines by predators (kruuk 1986; young 1992; hunter 1998; harrington et al. 1999), in the same way that small enclosed reserves would. in managing an assemblage of predators on a reserve, it can be expected that similar prey declines of the same species will occur, and the real challenge is to accommodate this. research would need to be aimed at sustainability of reserves for predators, but in the meantime the suggested practice is to have to re-supplement prey populations on small reserves. acknowledgements i would like to thank the management of the mabula game reserve, in particular mr. danie de bruyn, riaan van zyl and chris van der valk; all rangers from the mabula game lodge for information dissemination; mr. ben orban of the university of preissn 0075-6458 73 koedoe 45/2 (2002) power.qxd 2005/12/09 10:10 page 73 toria; various members of the african lion working group (alwg) for allowing me to present this information at their last meeting, and for encouragement to present it in this paper. references begon, m., j.l. harper & c.r. townsend. 1996. ecology. oxford: blackwell science. bothma, j. du p. 1996. game ranch management. pretoria: van schaik. bothma, j. du p. 1997. larger carnivores on game ranches. pp. 58 81. in: van heerden, j (ed.). proceedings of a symposium on lions and leopards as game ranch animals. pretoria: wildlife group of the south african veterinary association, onderstepoort. bredenkamp, g. j. & n. van rooyen. 1990. the vegetation and wildlife management of the mabula game reserve. pretoria: ekotrust. bryden, b.r. 1978. the biology of the african lion (panthera leo linnaeus, 1758) in the kruger national park. msc thesis, university of pretoria, pretoria. byers, c.r., r.k. steinhorst & p.r. krausman. 1984. clarification of a technique for analysis of utilisation-availability data. journal of wildlife management 48: 1050–1053. caughley, g. & a.r.e. sinclair. 1994. wildlife ecology and management. cambridge: blackwell science. du toit, j.t. 1988. patterns of resource use within the browsing ruminant guild in the central kruger national park. phd dissertation, university of witwatersrand, johannesburg. elliot, j.p. & i.m. cowan. 1978. territoriality, density and prey of lions in ngorongoro crater, tanzania. canadian journal of zoology 56: 1726–1734. eloff, f.c. 1984. food ecology of the kalahari lion panthera leo vernayi. koedoe (supplement) 27: 249–258. fairall, n. & d.r. klein. 1984. protein intake and water turnover: a comparison of two equivalently sized african antelope, the blesbok (damaliscus dorcas) and the impala (aepyceros melampus). canadian journal of animal science 64: 212–214. fryxell, j.m., j. greever & a.r.e. sinclair. 1988. why are migratory ungulates so abundant? american naturalist 131: 781–798. funston, p.j., m.g.l. mills & h.c. biggs. 2001. factors affecting the hunting success of male and female lions in the kruger national park. journal of zoology, london 253: 419–431. griffiths, d. 1975. prey availability and the food of predators. ecology 56: 1209–1214. harrington, r., n. owen-smith, p.c. viljoen, h.c. biggs, d.r. mason & p.c. funston. 1999. establishing the causes of the roan antelope decline in the kruger national park, south africa. biological conservation 90: 69–78. hirst, s.m.1969. predation as a regulating factor of wild ungulate populations in a transvaal lowveld nature reserve. zoologica africana 4(2): 199–230. hunter, l.t.b. 1998. the behavioural ecology of reintroduced lions and cheetahs in the phinda resource reserve, kwazulu-natal, south africa. phd thesis, university of pretoria, pretoria. hunter, l.t.b & j.d. skinner. 1997. vigilance behaviour in african ungulates: the role of predation pressure. behaviour 135: 195–211. jakoby, m. 1997. the nutritional requirements of the predators in the mabula game reserve and the sustainability of the lions in the madjuma lion reserve. honours project (wildlife management), university of pretoria. pretoria. jeffries, m. j & j.h. lawton. 1984. enemy-free space and the structure of ecological communities. biological journal of the linnaen society 23: 269–286. karanth, k.u. & m.e. sunquist. 1995. prey selection by tiger, leopard and dhole in tropical forests. journal of animal ecology 64: 439–450. kruuk, h. 1986. interactions between felidae and their prey species: a review. pp. 353–374. in: miller, s.d. & d. everett (eds.). cats of the world: biology, conservation and management. kingsville: caesar kleberg wildlife research institute. madjuma lion reserve. 1997. madjuma census figures. mabula game reserve mcbride, c.j. 1984. age and size categories of lions prey in the chobe national park, botswana. botswana notes and records 16: 139–143. mills, m.g.l. 1984. prey selection and feeding habits of the larger carnivores in the southern kalahari. koedoe (supplement) 27: 281–294. mills, m.g.l. 1990. kalahari hyaenas: comparative behavioural ecology of two species. london: unwin. mills, m.g.l. 1991. conservation management of large carnivores in africa. koedoe 34: 81–90. mills, m.g.l. 1996. methodological advances in capture, census, and food-habits studies of large african carnivores. pp. 223–242. in: gittleman, j.l (ed.). carnivore behaviour, ecology and evolution. london: comstock. mills, m.g.l & h.c. biggs. 1993. prey apportionment and related ecological relationships between large carnivores in the kruger national park. symposia of the zoological society of london 65: 253–266. koedoe 45/2 (2002) 74 issn 0075-6458 power.qxd 2005/12/09 10:10 page 74 mills, m.g.l. & p.f. retief. 1984. the response of ungulates to rainfall along the riverbeds of the southern kalahari. koedoe (supplement) 27: 129–141. mills, m.g.l & t.m. shenk. 1992. predator-prey relationships: the impact of lion predation on wildebeest and zebra populations. journal of animal ecology 61: 693–702. mooring, m. 1999. impala: the living fossil. africa, environment & wildlife 7(5): 53–61. neu, c.w., c.r. byers & j.m. peek. 1974. a technique for analysis of utilisation-availability data. journal of wildlife management 38: 541–545. owen-smith, n. 1984. demography of the greater kudu, tragelaphus strepsiceros in relation to rainfall. acta zoologica fennica 172: 197–199. owen-smith, n. 1993. comparative mortality rates of male and female kudus: the costs of sexual size dimorphism. journal of animal ecology 62: 428–440. packer, c., d. scheel & a. e. pusey. 1990. why lions form groups: food is not enough. american naturalist 136: 1–19. peel, m.j.s. & g.p. montagu.1999. modelling predator-prey interactions on a northern province game ranch. south african journal of wildlife research 29(2): 31–34. pienaar, u. de v. p. 1969. predator prey relationships amongst the large mammals of the kruger national park. koedoe 12: 108–177. power, r.j. 1998. the impact of lion panthera leo predation and the sustainability of the madjuma lion reserve, mabula game reserve, northern province. bsc (hons) (wildlife management) project, university of pretoria, pretoria. schaller, g.b. 1972. the serengeti lion: a study of predatorprey relations. chicago: university of chicago. scheel, d. 1993. profitability, encounter rates, and prey choice of african lions. behavioural ecology 4: 90–97. scheel, d. 1993. & c. packer. 1991. group hunting behaviour of lions: a search for cooperation. animal behaviour 41: 697–709. sinclair, a.r.e. 1995. population consequences of predation-sensitive foraging: the serengeti example. ecology 76: 882–891. sinclair, a.r.e., h.t. dublin & m. borner. 1985. population regulation of serengeti wildebeest: a test of food hypothesis. oecologia 65: 266–268. skinner, j.d & r.h.n. smithers. 1990. the mammals of the southern african subregion. pretoria: university of pretoria. smuts, g.l. 1978. interrelations between predators, prey and their environment. bio science 28: 316–320. sunquist, m. e. & f.c. sunquist. 1989. ecological constraints on predation by large felids. pp. 283–301. in: gittleman, j.l (ed). carnivore behaviour, ecology and evolution. london: comstock. stander, p.e. 1992a. foraging dynamics of lions in a semi-arid environment. canadian journal of zoology 70: 8–21. stander, p.e. 1992b. cooperative hunting in lions: the role of the individual. behavioural sociobiology 29: 445–454. stander, p.e & s.d. albon. 1993. hunting success of lions in a semi-arid environment. symposium of the zoological society of london 65: 127–142. van dyk, g. 1997. reintroduction techniques for lion panthera leo. pp. 82–91. in: van heerden. j. (ed.). proceedings of a symposium on lions and leopards as game ranch animals. pretoria: wildlife group of the south african veterinary association, onderstepoort. van orsdol, k.g. 1982. ranges and food habits of lions in rwenzori national park, uganda. symposia of the zoological society of london 49: 325–340. van orsdol, k.g. 1984. foraging behaviour and hunting success of lions in queen elizabeth national park, uganda. african journal of ecology 22: 79–99. van rooyen, n. & g. bredenkamp. 1996. mixed bushveld. pp 26. in: (eds.). low, a.b & a.g rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. van schalkwyk, a.g. 1994. riglyne vir die bestuur van leeus panthera leo in ingeperkte gebiede in suid-afrika. msc thesis (wildlife management), university of pretoria, pretoria. viljoen, p.c. 1993. the effects of changes in prey availability on lions predation in a natural ecosystem in northern botswana. symposium of the zoological society of london 65: 193–213. viljoen, p.c. 1996. lions. pp. 467–471. in: bothma, j. du p (ed.). game ranch management. pretoria: van schaik. whateley, a. & p.m. brooks.1985. the carnivores of the hluhluwe and umfolozi game reserves: 1973–1983. lammergeyer 35: 1–27. wentzel, j.j., j du p. bothma & n. van rooyen. 1991. characteristics of the herbaceous layer in preferred grazing areas of six herbivore species in the south-eastern kruger national park. koedoe 34(1): 57-58. young, e. 1992. game farming and wildlife management. nylstroom: eddie young publishers, zar, j.h.1986. biostatistical analysis. englewood cliffs: prentice-hall. issn 0075-6458 75 koedoe 45/2 (2002) power.qxd 2005/12/09 10:10 page 75 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <feff004200720075006700200064006900730073006500200069006e0064007300740069006c006c0069006e006700650072002000740069006c0020006100740020006f0070007200650074007400650020005000440046002d0064006f006b0075006d0065006e0074006500720020006d006500640020006800f8006a006500720065002000620069006c006c00650064006f0070006c00f80073006e0069006e0067002000740069006c0020007000720065002d00700072006500730073002d007500640073006b007200690076006e0069006e0067002000690020006800f8006a0020006b00760061006c0069007400650074002e0020005000440046002d0064006f006b0075006d0065006e007400650072006e00650020006b0061006e002000e50062006e006500730020006d006500640020004100630072006f0062006100740020006f0067002000520065006100640065007200200035002e00300020006f00670020006e0079006500720065002e00200044006900730073006500200069006e0064007300740069006c006c0069006e0067006500720020006b007200e600760065007200200069006e0074006500670072006500720069006e006700200061006600200073006b007200690066007400740079007000650072002e> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice abstract introduction methods results discussion acknowledgements references appendix 1 about the author(s) welsey l. hartmann department of conservation ecology and entomology, faculty of agrisciences, stellenbosch university, stellenbosch, south africa vicki fishlock amboseli trust for elephants, nairobi, kenyacentre for ecology and conservation, university of exeter, cornwall, united kingdom alison leslie department of conservation ecology and entomology, faculty of agrisciences, stellenbosch university, stellenbosch, south africa citation hartmann, w.l., fishlock, v. & leslie, a., 2021, ‘first guidelines and suggested best protocol for surveying african elephants (loxodonta africana) using a drone’, koedoe 63(1), a1687. https://doi.org/10.4102/koedoe.v63i1.1687 original research first guidelines and suggested best protocol for surveying african elephants (loxodonta africana) using a drone welsey l. hartmann, vicki fishlock, alison leslie received: 06 may 2021; accepted: 28 june 2021; published: 06 sept. 2021 copyright: © 2021. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract unmanned aerial vehicles, commonly known as drones, are increasingly used in ecological management, conservation and research. numerous reviews on drones tout almost unlimited potential within the wildlife sciences as they open up inaccessible habitats to observation. however, the influence of drones on the animals themselves is far less understood, and impact studies to construct protocols for best practices are urgently needed to minimise the potential for stress on target species. the impact of a quadcopter drone’s approach speed, angle of approach and initial starting altitude was tested on the behavioural responses of african elephants (loxodonta africana), along with sustained speed and flight pattern. seventy-nine approach flights and 70 presence flights were conducted. the speed and angle of approach significantly impacted the success of a flight, but neither speed nor flight pattern had any measurable impact on elephants’ behaviour during sustained flights. it is recommended that drones be launched at a distance of 100 m from an elephant or a herd of elephants, ascending to a height of 50 m by using an approach speed of 2m/s and an approach angle of 45 ° or less to successfully contact elephant targets. conservation implications: this study aimed to provide a significant step towards the ethical use of drones in wildlife research. further research is required to investigate the impacts of drones on other taxa. physiological responses to drones, for example, would determine if physiological stress responses unlinked to behavioural indicators are of concern in elephants. keywords: african elephant; aerial monitoring; drones; ethology; elephant behaviour; technology; unmanned aerial vehicles. introduction wildlife science usually focusses on the study, monitoring and management of animals and their habitats (chabot & bird 2015). although these goals may be relatively simple, achieving them can be extremely challenging, particularly as resources are often limited and target species can be elusive, wide-ranging, sensitive to anthropogenic disturbances and/or dangerous to approach (chabot & bird 2015). additionally, many target animals occupy habitats that are extensive, remote and often impossible to access at ground-level. new technologies have greatly aided accessing these difficult subjects in their challenging habitats. examples include motion-triggered camera traps (o’connell, nichols & karanth 2011), aircraft (fleming & tracey 2008), remote sensing satellites (kerr & ostrovsky 2003), radar (larkin 2005), thermal cameras (o’neil et al. 2005), projectile-based animal-capturing devices and chemical immobilisation agents (roffe, sweeney & aune 2005; schemnitz 2005) and a vast array of electronic tracking devices and accompanying software (thomas, holland & minot 2011). one technology that is rapidly gaining popularity are the aerial units known variously as unmanned aircraft systems (uas), unmanned aerial vehicles (uav), remotely piloted aircraft systems or (mostly popularly) drones. the popularity of drones amongst wildlife biologists, ecologists and conservationists is clear from the many review articles investigating the applications and proliferation of drone use in remote sensing, natural resource sciences and ecology (allan et al. 2015; anderson & gaston 2013; christie et al. 2016; colomina & molina 2014; jones, pearlstine & percival 2006; koh & wich 2012; pajares 2015; shahbazi, theau & menard 2014; watts, ambrosia & hinkley 2012; whitehead & hugenholtz 2014; whitehead et al. 2014). chabot and bird (2015) conducted an extensive review of drone use in wildlife management in which they highlighted optical surveying and observation of animals, uses of drones in autonomous wildlife telemetry tracking, habitat research and monitoring and a review of the broader potential for uavs. although the capabilities and potential practical uses of drones in the field of wildlife and conservation biology has been investigated thoroughly, their effects on the animals themselves have not been widely addressed (leslie 2018). this study assessed the possible effects of drones on african elephants (loxodonta africana). previous drone research on elephants has investigated the use of drones in mitigating human–elephant conflict (hahn et al. 2016) and for population surveys (vermeulen et al. 2013), but no work has examined the impact of drones on elephants or suggested any ethical recommendations for drone use. because elephants exhibit high sensitivity to anthropogenic impacts and fit into many criteria for which drones might be considered as suitable monitoring tools (wide-ranging, sometimes in inaccessible habitats or perhaps dangerous to approach), there is a clear need to examine the potential impact of drones on elephants. the protocols developed in this study can be used to minimise the stress when utilising drones for elephant research. our specific objectives were to determine the following: what are the ideal approach speed and angles for drones to observe elephants without disturbance? what is the ideal sustained flight protocol for drones to observe elephants without disturbance? do different types or sizes of elephant groups (breeding herds, lone bulls or bachelor herds) respond differently to the presence of a drone? do elephant populations with greater exposure to previous, potentially negative drone or other aircraft encounters differ in their response to drone presence? we hypothesised the following points: (1) slower speeds and a less steep angle of approach would minimise the disturbance to elephants as these patterns were successful with other taxa (vas et al. 2015); (2) slower speeds and steady altitude would result in greater success rates for the sustained flight, that is keeping the drone with in close proximity to the elephants and collecting data; (3) herds containing dependent offspring would be more sensitive to the approach and presence of a drone; and (4) because of the higher levels of poaching, and subsequent helicopter and drone activity experienced by liwonde national park (lnp), we hypothesise that the elephants in the lnp would be more sensitive to the approach and presence of a drone (i.e. the approach or presence of a drone will be more likely to elicit higher response levels [see table 2]). table 1: the variables used throughout the approach and presence portions of data collection with the corresponding ‘shortcuts’ and abbreviations. table 2: the various possible elephant responses recorded during both the approach and presence data collection. methods study area majete wildlife reserve majete wildlife reserve (mwr) is located at the southern tip of the great rift valley in the lower shire valley region of southern malawi (15.9364 °s, 34.6414 °e) (figure 1). the reserve is 700 km2 in size with two perennial rivers, the mkulumadzi and the shire. the altitude within the reserve varies greatly, with the western region containing steeply undulating hills disrupted by river valleys with the terrain flattening towards the shire river in the east. two well-defined seasons occur in majete: the wet season (december to may) and the dry season (june to november). annual precipitation varies according to topography from 680 to 1000 mm annually (wienand 2013). whilst surface water availability is dependent on seasonal rainfall, there are 10 artificial borehole-fed waterholes, as well as several naturally occurring perennial springs. mwr is not only dominated by multi-altitude woodlands, but also contains open savanna grassland areas (forrer 2017). figure 1: africa is shown with malawi’s location highlighted. a map of malawi is also displayed in the centre, in which the locations of majete wildlife reserve and liwonde national park can be seen. on the left, majete wildlife reserve is displayed with perennial and seasonal rivers indicated, and on the right liwonde national park is shown with its perennial and seasonal rivers. in 1955, mwr was gazetted as a protected area, but poaching and poor management led to most of its large game being decimated by 2003 (forrer 2017). as a result, a public private partnership (ppp) agreement was made between african parks, majete (pty) ltd. and the malawian department of national parks and wildlife (dnpw) to initiate one of africa’s greatest reintroduction programmes. over 2550 individual animals, comprising 14 different species, were reintroduced into mwr. elephant, black rhino (diceros bicornis), buffalo (syncerus caffer), sable (hippotragus niger), hartebeest (alcelaphus buselaphus), several other antelope species and many predators were all included in the reintroduction programme. the reintroduction was considered successful as all wildlife populations are continuously increasing, including the elephant population which was steadily impacting the vegetation in mwr. the combination of increased rates of vegetation damage, because of the rapidly expanding elephant population within mwr, and the desire to reintroduce elephants to nkhotakota wildlife reserve led to a decision to conduct a major translocation operation. during the months of june and july of 2017, 154 elephants were translocated out of mwr. liwonde national park liwonde national park is located in the upper shire valley, which forms part of the great east african rift valley in southern malawi (14.8441 °s, 35.3466 °e) (figure 1). the park is 548 km2 in size and is generally flat except for three separate groups of hills. the dominant vegetation type in the park is colophospermum mopane woodland, which occupies approximately 70% of the total area of the park (morris 2006). liwonde’s dry season lasts from april to october and the rainy season from november to march. annual rainfall ranges from 700 mm to 1400 mm. there are four artificial water points, which provide perennial water, located throughout the park (morris 2006). historically a sport hunting ground for european planters and administrators from 1920 to 1969 (morris 2006; taylor 2002), lnp went through several status evolutions before being gazetted as a national park in 1973, and formally opened to the public for wildlife viewing in 1978 (morris 2006). in partnership with malawi’s dnpw, the african parks network assumed management of lnp in 2015. the new partnership saw significant investment in law enforcement and the construction of a new perimeter fence (sievert, reid & botha 2018) reduced elephant and rhino poaching. additionally, 1329 animals were translocated for restocking of other malawian reserves, and the new partnership and protection resulted in the return of five, vulture species, the supplementation of the remnant black rhino population and the reintroduction of lion (panthera leo) and cheetah (acinonyx jubatus) (sievert & reid 2018; sievert et al. 2018). unmanned aerial vehicle (drone) this study used a uav quadcopter, the mavic pro platinum (dji, shenzhen, china). this specific model was chosen for its compact design, ability to launch and land in most places and noise reduction blades (4 db quieter than the traditional dji mavic pro), which at the time of the study, september 2018, made it the quietest commercial drone available. the drone is equipped with a gps and internal measurement unit, which both aid in determining the position and height of the drone. the drone is remote controlled with a maximum flight range of 7 km. more information regarding the drone’s specifications can be found in appendix 1, table 1-a1. data collection data collection in mwr occurred between september 2018 and april 2019, and in lnp between april and may 2019. the same methodology was applied within both reserves. as elephants range widely and unpredictably, the sampling scheme was opportunistic in nature. when an elephant or herd was encountered, primarily by vehicle, the elephant or herd was given time to ‘settle’ (i.e. display no outward signs of agitation or aggression) before the drone was launched and flown towards them. if the elephant or herd did not settle within 60 min, the drone was not launched. the dji flight software displays the appropriate metadata (altitude, distance, speed, etc.) and was captured by screen-recording the playback device (apple iphone 7+, apple inc., usa). the location of launch and landing, the ambient temperature, the wind strength and the wind direction (towards launch point, away from launch point, across launch point) were all recorded. wind strength was determined by using the beaufort scale (appendix 1, table 2-a1). flights were not conducted if the wind speed was greater than a ‘strong breeze’ (beaufort number 6, 10.8–13.8 m/s) as this exceeds the drone’s wind tolerance capabilities. the drone was launched at a minimum distance of 100 m from the elephants to ensure a safe distance should the launch induce an aggressive response. drone approach the approach methodology largely followed that conducted by vas et al. (2015). from the launch site, the drone ascended vertically to either a height of 35 m, 50 m or 100 m and then approached the elephants. each flight had a pre-determined approach pattern, selected at random from 18 possible combinations that varied speed and angle of approach (table 1). during each approach, the live on-screen video was recorded and analysed post-flight for elephant responses, which were scored by using a standardised scoring system adapted from langbauer et al. (1991), poole (1999), o’connell-rodwell et al. (2006) and soltis et al. (2014), with a five-point scale: no response; vigilance; agitated; flight; or aggressive (table 2). for each approach, the drone height, distance from the elephants and elephants’ responses were recorded at 1-min intervals. approaches were classed as successful if the drone was able to reach a distance of 30 m or closer to the elephants without inducing a type 3, 4 or 5 response from at least one adult individual. thirty metres was determined to be close enough to enable high-resolution observations with this drone model. flights were terminated if the elephants displayed either a flight (4) or an aggressive (5) response at any point during the flight. drone presence if an approach did not elicit a type 4 or 5 response, the drone was kept in contact with the elephants and a sustained flight was initiated, varying drone height and speed (table 1) according to a randomly selected flight pattern. drone height and distance and elephant response were scored at 1-min intervals up to a maximum flight duration of 25 min (because of the battery life of the drone). flights were terminated if the elephants displayed either a flight (4) or an aggressive (5) response at any point during the flight (approach or presence). statistical analysis data were captured by using ms excel and analysed by using statistica 13 (tibco software inc. (2017); http://statistica.io., dell). relationships between continuous variables and a nominal predictor variable were examined by using one-way anova or non-parametrically by using mann–whitney or kruskal–wallis tests. nominal variable relationships were investigated with contingency tables and likelihood ratio or pearson’s chi-square tests. generalised linear models (glz) were used to investigate the influence of environmental and flight variables (appendix 1, table 3-a1) on the success of flight approach or presence. model fits were reported via multiple measures. ethical considerations this study was cleared ethically by the university of stellenbosch’s research ethics committee: animal care and use (protocol number: acu-2019-7822). results contrary to expectations, there was no population level effect on drone flight success (χ2, [df = 1] = 0.53, p = 0.46555) and the subsequent analyses dropped population as a variable. instead, response as a function of previous drone experience was examined within the study as a shorter-term measure of potential disturbance. drone approach seventy-nine approach flights were conducted, and lone males were the most frequently contacted group type (table 3). group type did not affect the likelihood of an approach being successful (glz, estimate = 0.44879, p = 0.324), but slower approach speeds increased the likelihood of a successful flight (glz [akaike information criterion {aicc} = 64.98, bic = 83.70, nagelkerke r2 = 0.75], estimate = 0.9085, p = 0.004; figure 2). approach angles of 45° also increased approach success (glz [aicc = 64.98, bic = 83.70, nagelkerke r2 = 0.75], estimate = 0.0472, p = 0.01456; figure 2). starting altitude had no significant effect on the success of an approach (glz [aicc = 64.98, bic = 83.70, nagelkerke r2 = 0.75], estimate = 0.0055, p = 0.718), nor did environmental factors (glz [aicc = 64.98, bic = 83.70, nagelkerke r2 = 0.75], temperature: estimate = -0.06374, p = 0.638; season: estimate = 0.40891, p = 0.310; weather: estimate = -0.35869, p = 0.622; wind: estimate = 1.4039460, p = 0.536). figure 2: the percentage of successful approaches, regardless of elephant category, when broken down by approach speed (2 m/s, 4 m/s or 6 m/s), angle of approach (45 ° or 90 °) and starting altitude (30 m, 50 m or 100 m). table 3: the breakdown in the effort for approach flights across majete wildlife reserve and liwonde national park. seventy presence flights were conducted with a 60% average success rate (table 4). the average presence flight length was 15 min; however, the flight length of presence flights with a preceding successful approach differed significantly from those with unsuccessful preceding approach flights (kruskal–wallis chi-squared = 42.369, p < 0.0001, figure 3). successful approaches allowed a presence flight to be around the target elephants for an average of seven and a half minutes more than presence flights with an unsuccessful approach. figure 3: a significant difference was detected between the two groups (whether an elephant or group had a successful or unsuccessful approach flight) with significance levels measured at p < 0.05. table 4: the breakdown in effort for presence flights. once again, group type did not affect the likelihood of a presence flight being successful (glz [aicc = 52.20, bic = 67.83, nagelkerke r2 = 0.79], estimate = 0.68829, p = 0.422). although some trends could be seen when broken down into flight variables (figure 4), no presence flight variable (speed and flight pattern [fixed vs varied]) was found to significantly influence the success of a sustained presence flight (glz [aicc = 49.84, bic = 66.63, nagelkerke r2 = 0.78], speed: estimate = -0.01996, p = 0.956; flight pattern: estimate = 0.54753, p = 0.259). likewise, no environmental factors were found to influence a sustained presence flight’s success significantly (glz [aicc = 49.84, bic = 66.63, nagelkerke r2 = 0.78], temperature: estimate = -0.07591, p = 0.658; season: estimate = -0.34112, p = 0.136; weather: estimate = 0.16054, p = 0.212; wind: estimate = 0.46393, p = 0.290). notably, only the success of the preceding approach was found to influence the success of the following presence flight (glz [aicc = 52.20, bic = 67.83, nagelkerke r2 = 0.79], estimate = 2.39497, p < 0.0001). figure 4: the percentage of successful presence flights, regardless of elephant category, when broken down by flight speed (2, 4 or 6 m/s) and flight pattern (fixed or varied). discussion the timely and repeatable manner in which drones deliver high-definition (hd) picture and video footage of animals in often hard-to-reach places is a tool that every conservationist undoubtedly desires. a limited number of previous studies have indicated the usefulness of drones for collecting observational data (ivoševi et al. 2015), and have quantified responses, for example with seals (pomeroy & connor 2015), birds (vas et al. 2015) and bears (ditmer et al. 2015). in this study, we quantified how elephants responded to various drone approach patterns and sustained drone flights. although only a 54% and 60% success rate were achieved for the approach and presence flights, respectively, key insights into which factors influence elephants’ behavioural responses were obtained. environmental variables such as ambient temperature, weather, wind speed and season did not influence success rates of approach flights. it is likely that lone males were the most frequently contacted group type because the 2016–2017 translocation removed only female breeding groups, skewing the population structure towards males, and possibly resulting in female herds to be more cautious around vehicles. however, contrary to predictions, the category of elephant group targeted for a flight also had no effect on approach flight success, so females with calves and infants were not more sensitive to the approach of a drone, nor did starting altitude affect approach success, even though it would allow more time for elephants to assess whether the drone was a threat or not. perhaps most surprisingly, there was no difference between populations, even though lnp elephants had high exposure to drones and helicopters and would be expected to show lower responses in general. this result might be interpreted with caution because of the low sample size in lnp. approach speed and angle significantly affected the success of an approach flight. the speed aspect makes logical sense, as the drone is quieter at slower speeds and may allow elephants additional time to identify (or attempt to determine) whether the drone is a threat or not. the reason a 45 ° angle of approach was preferable to a 90 ° of approach may very well follow a similar logic. in the field, when elephants elicited an agitated response, they were often observed to turn towards the direction in which the drone was approaching. with a 45 ° angle of approach, elephants were observed to tilt their heads upwards and appeared to be looking directly at the drone, once again, determining whether it was a ‘real’ threat or not. however, with a 90 ° angle of approach, elephants were unable to look directly above them and thus the noise from an unknown source descending upon them intuitively seems to explain their discomfort with this approach angle. as for flight variables, very surprisingly neither pattern (a fixed or varied flight height) nor speed had a significant effect on success rates. it was assumed that slower speeds would yield greater rates of success as would a fixed flight height. a fixed flight height was assumed to be preferable to elephants as the pitch of the drone does not change much if kept at a constant height. this is shown in figure 4, as across the three speeds, a fixed flight pattern consistently yielded a higher percentage of successful sustained flight. likewise, slower speeds resulted in higher percentages of successful sustained presence flights. however, for both variables there was no statistical difference. the only variable that was found to have a significant effect on the success of a sustained presence flight was whether the preceding approach flight had been successful or not. it was found that if an approach was successful, a sustained presence would almost always be too. it is therefore recommended that the approach protocols are always strictly adhered to, so as to prevent the initial disturbance and subsequent sustained disturbance of the elephants being droned. if the elephants are disturbed upon approach, it is unlikely they will return to an undisturbed state throughout the rest of the droning session. should this occur, it is recommended that the droning session be cancelled for those elephants and only attempted again at a later stage. as only just over a week was spent in lnp, a limited amount of data was collected. not all approach or presence flight patterns were conducted, and thus a full pair-wise comparison was not possible with all the factors in one model. however, from the data collected, no clear trends were apparent in either the approach flights or presence flights. this indicates that there appears to be a critical amount or number of flights required for patterns to emerge. somewhere between the number of flights conducted within mwr (63 approaches and 56 presences), where clear trends and significant differences were observed, and lnp (16 approaches and 14 presences) is the minimum amount or number of flights required for statistical validity. unlike in previous studies, where the animals were easy to locate and almost always readily accessible (e.g. vas et al. 2015), droning sessions took place opportunistically when elephants were found in the field. despite a lengthy field data collection period, a relatively low number of observational flights (both approaches and presences) were conducted. additionally, not all patterns were flown the same number of times, with some never flown at all (the case for the lnp population). this may account for some of the variables not being significant in influencing elephant behavioural responses to the drone. this study has clearly demonstrated that the speed and angle at which elephants are approached by a drone play a critical role in the elephants’ ability to tolerate drones. because presence flights depended only on the approach success, early exposure to drones determines the ability to stay with target subjects. drone pilots, regardless of purpose (i.e. scientific or cinematic), need to be made aware of this and understand that speed and angle of approach will affect elephant responses, but once the approaches are unsuccessful, mitigating the effects in order to remain with target elephants becomes difficult. drone pilots should always approach elephants with extreme care as reckless flying and repeated negative experiences are likely to limit drone success for elephants and may contribute to population stress. furthermore, because drones are more likely to be used on stressed populations, pilots need to exert caution. further highlighting the importance of carefully approaching elephants aerially is the difference between the average length of time of sustained flights, for flights that had a successful versus unsuccessful approach, as seen in the time differences in the results. aggressive or reckless approaches not only create negative associations between the target elephant population and drones, but they also ultimately inhibit long periods of observations and compromise the data and/or footage captured. a major assumption of this study, as well as two previous studies (pomeroy & connor 2015; vas et al. 2015), is the use of observable behaviour as a proxy for the animals’ response to drones. whilst this may serve as an acceptable model for the time being, greater efforts should be made to additionally measure the physiological response of the animals. ditmer et al. (2015) found that although bears showed little to no outward behavioural response to drones, their heart rates increased significantly during drone flights. for elephants, the use of faecal samples to measure cortisol levels, to measure stress levels of elephants (foley, papageorge & wasser 2002; ganswindt et al. 2010; viljoen et al. 2008), would provide information as to whether the outward behavioural response elicited by a drone is a good indicator of its physiological response as well. this study is expected to act as a platform from which future research can be built upon. it aimed to outline a methodology of how to quantify an animal’s response to the approach and presence of a drone. although the sfocus of this study was elephants, the methodology employed can be easily manipulated to a wide variety of other wildlife species. if we wish to utilise new technology to aid in the efforts of conservation, at the very least we should understand how these technologies affect those species that we are trying to conserve. acknowledgements the authors would like to thank the earthwatch institute for funding the animals of malawi project and for funding this study. they also thank the management team of majete wildlife reserve (mwr) for their support, and mwr’s various staff members, without whom data collection would not have been possible. thanks go to african parks for allowing the study to take place in mwr. additionally, thanks go to dji and the d1store, australia, and ilenna copley for the sponsorship and acquisition of the dji mavic pro platinum and accompanying spare parts used throughout this study. finally, the authors thank sally reece, fleur visser and rhiannon jones for their assistance in the field, and professor daan nel for help with statistics. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions w.l.h. conceived the original idea, carried out the experiment and wrote the manuscript with support from a.l. and v.f. a.l. supervised the project. funding information this work and the animals of malawi research programme was supported by the earthwatch institute (a grant to ajl) and the university of stellenbosch, south africa. data availability the data that support the findings of this study are available from the corresponding author, w.l.h., upon reasonable request. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references allan, b.m., ierodiaconou, d., nimmo, d.g., herbert, m. & ritchie, e.g., 2015, ‘free as a drone: ecologists can add uavs to their toolbox’, frontiers in ecology and the environment 13(7), 354–355. https://doi.org/10.1890/15.wb.015 anderson, k. & gaston, k.j., 2013, ‘lightweight unmanned aerial vehicles will revolutionize spatial ecology’, frontiers in ecology and the environment 11(3), 138–146. https://doi.org/10.1890/120150 chabot, d. & bird, d., 2015, ‘wildlife research and management methods in the 21st century: where do unmanned aircraft fit in?’, journal of unmanned vehicle systems 3(4), 137–155. https://doi.org/10.1139/juvs-2015-0021 christie, k.s., gilbert, s.l., brown, c.l., hatfield, m. & hanson, l., 2016, ‘unmanned aircraft systems in wildlife research: current and future applications of a transformative technology’, frontiers in ecology and the environment 14(5), 241–251. https://doi.org/10.1002/fee.1281 colomina, i. & molina, p., 2014, ‘unmanned aerial systems for photogrammetry and remote sensing: a review’, isprs journal of photogrammetry and remote sensing 92, 79–97. https://doi.org/10.1016/j.isprsjprs.2014.02.013 ditmer, m.a., vincent, j.b., werden, l.k., tanner, j.c., laske, t.g., iaizzo, p.a. et al., 2015, ‘bears show a physiological but limited behavioral response to unmanned aerial vehicles’, current biology 25(17), 2278–2283. https://doi.org/10.1016/j.cub.2015.07.024 fleming, p.j.s. & tracey, j.p., 2008, ‘aerial surveys of wildlife: theory and applications’, wildlife research 35(4), 258–267. https://doi.org/10.1071/wr07081 foley, c.a.h., papageorge, s. & wasser, s.k., 2002, ‘non-invasive stress and reproductive measures of social and ecological pressures in free-ranging african elephants’, conservation biology 15(4), 1134–1142. https://doi.org/10.1046/j.1523-1739.2001.0150041134.x forrer, f.a., 2017, ‘the population status, habitat use and seasonal diet of african elephant (loxodonta africana) in majete wildlife reserve, malawi’, master’s thesis, stellenbosch university, stellenbosch. ganswindt, a., munscher, s., henley, m., palme, r., thompson, p. & bertschinger, h., 2010, ‘concentrations of faecal glucocorticoid metabolites in physically injured free-ranging african elephants loxodonta africana’, wildlife biology 16(3), 323–332. https://doi.org/10.2981/09-081 hahn, n., mwakatobe, a., konuche, j., de souza, n., keyyu, j., goss, m. et al., 2016, ‘unmanned aerial vehicles mitigate human-elephant conflict on the boarders of tanzanian parks: a case study’, oryx 51(3), 513–516. https://doi.org/10.1017/s0030605316000946 ivoševi, b., han, y., cho, y. & kwon, o., 2015, ‘the use of conservation drones in ecology and wildlife research’, journal of ecology and environment 38(1), 113–118. https://doi.org/10.5141/ecoenv.2015.012 jones, g.p., pearlstine, l.g. & percival, h.f., 2006, ‘an assessment of small unmanned aerial vehicles for wildlife research’, wildlife society bulletin 34(3), 750–758. https://doi.org/10.2193/0091-7648(2006)34[750:aaosua]2.0.co;2 kerr, j.t. & ostrovsky, m., 2003, ‘from space to species: ecological applications for remote sensing’, trends in ecology and evolution 18(6), 299–305. https://doi.org/10.1016/s0169-5347(03)00071-5 koh, l.p. & wich, s.a., 2012, ‘dawn of drone ecology: low-cost autonomous aerial vehicles for conservation’, tropical conservation science 5(2), 121–132. https://doi.org/10.1177/194008291200500202 langbauer, j.r., payne, w.r., charif, r.a., rapaport, l. & osborn, f., 1991, ‘african elephants respond to distant playbacks of low-frequency conspecific calls’, journal of experimental biology 157(1), 35–46. https://doi.org/10.1242/jeb.157.1.35 larkin, r.p., 2005, ‘radar techniques for wildlife biology’, in c.e. braun (eds.), techniques for wildlife investigations and management, 6th edn., pp. 448–464, the wildlife society, bethesda, md. leslie a.j., 2018, ‘earthwatch annual field report’, animals of malawi [unpublished]. morris, b., 2006, the history and conservation of mammals in malawi, monograph no. 21, malawi publishers zomba, kachere series, 2006. o’connell, a.f., nichols, j.d. & karanth, k.u., 2011, camera traps in animal ecology, springer, new york, ny. o’connell-rodwell, c.e., wood, j.d., rodwell, t.c., puria, s., partan, s.r. & keefe, r., 2006, ‘wild elephant (loxodonta africana) breeding herds respond to artificially transmitted seismic stimuli’, behavioural ecology and sociobiology 59, 842–850. https://doi.org/10.1007/s00265-005-0136-2 o’neil, t.a., bettinger, p., marcot, b.g., luscombe, b.w., koeln, g.t., bruner, h.j. et al., 2005, ‘application of spatial technologies in wildlife biology’, in c.e. braun (eds.), techniques for wildlife investigations and management, 6th edn., pp. 418–447, the wildlife society, bethesda, md. pajares, g., 2015, ‘overview and current status of remote sensing applications based on unmanned aerial vehicles (uavs)’, photogrammetric engineering and remote sensing 81(4), 281–329. https://doi.org/10.14358/pers.81.4.281 pomeroy, p. & connor, l.o., 2015, ‘assessing use of and reaction to unmanned aerial systems in gray and harbor seals during breeding and molt in the uk’, journal of unmanned vehicle systems 3(3), 102–113. https://doi.org/10.1139/juvs-2015-0013 poole, j.h., 1999, ‘signals and assessment in african elephants: evidence from playback experiments’, animal behaviour 58, 185–193. https://doi.org/10.1006/anbe.1999.1117 roffe, t.j., sweeney, s.j. & aune, k.e., 2005, ‘chemical immobilization of north american mammals’, in c.e. braun (ed.), techniques for wildlife investigations and management, 6th edn., pp. 286–302, the wildlife society, bethesda, md. schemnitz, s.d., 2005, ‘capturing and handling wild animals’, in c.e. braun (ed.), techniques for wildlife investigations and management, 6th edn., pp. 239–285, the wildlife society, bethesda, md. sievert, o. & reid, c., 2018, ‘liwonde national park aerial survey’, african parks, liwonde. sievert, o., reid, c. & botha, a.j., 2018, ‘the re-emergence of african white-backed vultures (gyps africanus), white-headed vultures (trigonoceps occipitalis) and lappet-faced vultures (torgos tracheliotos) in liwonde national park, malawi’, vulture news 74(1), 12–19. https://doi.org/10.4314/vulnew.v74i1.2 shahbazi, m., theau, j. & menard, p., 2014, ‘recent applications of unmanned aerial imagery in natural resource management’, giscience & remote sensing 51(4), 339–365. https://doi.org/10.1080/15481603.2014.926650 soltis, j., king, l.e., douglas-hamilton, i., vollrath, f. & savage, a., 2014, ‘african elephant alarm calls distinguish between threats from humans and bees’, plos one 9(2), e89403. https://doi.org/10.1371/journal.pone.0089403 taylor, p., 2002, ‘good news for liwonde national park, malawi’, oryx 36(4), 323–324. thomas, b., holland, j.d. & minot, e.o., 2011, ‘wildlife tracking technology options and cost considerations’, wildlife research 38(8), 653–663. https://doi.org/10.1071/wr10211 vas, e., lescroël, a., duriez, o., boguszewski, g. & gremillet, d., 2015, ‘approaching birds with drones: first experiments and ethical guidelines’, biology letters 11(2), 20140754. https://doi.org/10.1098/rsbl.2014.0754 vermeulen, c., lejeune, p., lisein, j., sawadogo, p. & bouche, p., 2013, ‘unmanned aerial survey of elephants’, plos one 8(2), e54700. https://doi.org/10.1371/journal.pone.0054700 viljoen, j.j., ganswindt, a., palme, r., reynecke, h.c., du toit, j.t. & langbauer, jr, w.r., 2008, ‘measurement of concentrations of faecal glucocorticoid metabolites in free-ranging african elephants within the kruger national park’, koedoe 50(1), 18–21. https://doi.org/10.4102/koedoe.v50i1.129 watts, a.c, ambrosia, v.g. & hinkley, e.a., 2012, ‘unmanned aircraft systems in remote sensing and scientific research: classification and considerations of use’, remote sensing 4(6), 1671–1692. https://doi.org/10.3390/rs4061671 whitehead, k. & hugenholtz, c.h., 2014, ‘remote sensing of the environment with small unmanned aircraft systems (uass), part 1: a review of progress and challenges’, journal of unmanned vehicles and systems 2(3), 69–85. https://doi.org/10.1139/juvs-2014-0006 whitehead, k., hugenholtz, c.h., myshak, s., brown, o., leclair, a., tamminga, a. et al., 2014, ‘remote sensing of the environment with small unmanned aircraft systems (uass), part 2: scientific and commercial applications’, journal of unmanned vehicle systems 2(3), 86–102. https://doi.org/10.1139/juvs-2014-0007 wienand, j.j., 2013, ‘woody vegetation change and elephant water point use in majete wildlife reserve: implications for water management strategies’, master’s thesis, stellenbosch university, stellenbosch. appendix 1 table 1-a1: aircraft specifications taken from the website of dji (2018). table 2-a1: the beaufort scale was used to determine the strength of the wind (wind speed) during drone flights. table 3-a1: input variables used to investigate possible factors influencing drone approach and presence flights around elephants. abstract introduction data and methods methodology results discussion acknowledgements references about the author(s) marcel urban department for earth observation, friedrich-schiller-university jena, jena, germany kai heckel department for earth observation, friedrich-schiller-university jena, jena, germany christian berger department for earth observation, friedrich-schiller-university jena, jena, germany patrick schratz department of statistics, computational statistics group, ludwig-maximilian university of munich, münchen, germany department of geography, giscience group, friedrich-schiller-university jena, jena, germany izak p.j. smit scientific services, south african national parks, skukuza, south africa centre for african ecology, school of animal, plant and environmental sciences, university of the witwatersrand, johannesburg, south africa tercia strydom scientific services, south african national parks, skukuza, south africa jussi baade department for physical geography, friedrich-schiller-university jena, jena, germany christiane schmullius department for earth observation, friedrich-schiller-university jena, jena, germany citation urban, m., heckel, k., berger, c., schratz, p., smit, i.p.j., strydom, t. et al., 2020, ‘woody cover mapping in the savanna ecosystem of the kruger national park using sentinel-1 c-band time series data’, koedoe 62(1), a1621. https://doi.org/10.4102/koedoe.v62i1.1621 short communication woody cover mapping in the savanna ecosystem of the kruger national park using sentinel-1 c-band time series data marcel urban, kai heckel, christian berger, patrick schratz, izak p.j. smit, tercia strydom, jussi baade, christiane schmullius received: 13 jan. 2020; accepted: 02 june 2020; published: 04 aug. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the savanna ecosystems in south africa, which are predominantly characterised by woody vegetation (e.g. shrubs and trees) and grasslands with annual phenological cycles, are shaped by ecosystem processes such as droughts, fires and herbivory interacting with management actions. therefore, monitoring of the intraand inter-annual vegetation structure dynamics is one of the essential components for the management of complex savanna ecosystems such as the kruger national park (knp). to map the woody cover in the knp, data from european space agency’s (esa) copernicus sentinel-1 radar satellite (c-band vertical–vertical [vv]/vertical–horizontal [vh]) for the years 2016 and 2017, at 10 m spatial resolution and repeated acquisitions every 12 days, were utilised. a high-resolution light detection and ranging (lidar) data set was reclassified to produce woody cover percentages and consequently used for calibration and validation. woody cover estimation for different spatial resolutions was carried out by fitting a random forest (rf) model. model accuracy was assessed via spatial cross-validation and revealed an overall root mean squared error (rmse) of 22.8% for the product with a spatial resolution of 10 m and improved with spatial averaging to 15.8% for 30 m, 14.8% for 50 m and 13.4% for 100 m. in addition, the product was validated against a second lidar data set, confirming the results of the spatial cross-validation of the model. the methodology of this study is designed for savanna vegetation structure mapping based on height estimates by using open-source software and open-access data, to allow for a continuation of woody cover classification and change monitoring in these types of ecosystems. conservation implications: information about the state and changes in woody cover are important for park management and conservation efforts. both increasing (e.g. because of atmospheric carbon fertilisation) and decreasing (e.g. because of elephant impact) woody cover patterns will have cascading effects on other ecosystem processes such as fire and herbivory. keywords: woody cover; earth observation; lidar; radar; machine learning. introduction savanna ecosystems are dominated by different densities of grasses and woody plants with inter-annual changes because of dry and wet seasons. they cover half of the african continent and around 20% of the global land surface and are of great significance for ecology (e.g. living environment) and economy (e.g. fuelwood, timber) (kutsch et al. 2008; main et al. 2016; scholes & walker 1993). the savanna ecosystems of south africa are shaped by disturbance processes such as droughts, fire and herbivory (druce et al. 2008; scholes & archer 1997; stevens et al. 2016), as well as anthropogenic impacts like climate change (e.g. increase in atmospheric co2) or management actions. it is therefore not surprising that these savannas have been undergoing various changes during the last decades (buitenwerf et al. 2012; skowno et al. 2016). information about changes in woody cover and above-ground biomass (agb) in national parks (e.g. kruger national park [knp]) is important for park management and conservation efforts, as changes in woody cover are likely to have effects on other ecosystem patterns and processes. for example, an increase in woody cover will lead to a reduction in grass and herbaceous biomass (berger et al. 2019), which will have cascading effects on herbivores (i.e. favouring browsers to grazers [smit & prins 2015]) and fire regimes (i.e. reducing fire frequency [smit et al. 2012]). furthermore, woody cover maps are useful for a range of other park management applications. for example, the knp has used earlier versions of woody cover maps for adjusting rhino census estimates based on visibility (ferreira et al. 2015) and understanding how woody encroachment may impact the large mammal communities of the park (smit & prins 2015). also, although remote sensing monitoring of woody cover will not be able to replace extensive field monitoring of woody vegetation composition (kiker et al. 2014), it does provide a cost-effective and wall-to-wall approach for monitoring fractional cover, which is considered as a satellite remote sensing essential biodiversity variable (pettorelli et al. 2016). the recent launch of new satellite missions, such as the copernicus sentinel fleet of the european space agency (esa), has led to a tremendous increase in the availability of freely available earth observation (eo) data that can be utilised for woody cover-monitoring applications. sentinel-1a/b, two c-band synthetic aperture radar (sar) satellites, have a revisit time of up to 12 days in south africa and acquire images with a spatial resolution of 10 m (attema et al. 2009). estimation of woody cover by using eo data from different sources across different wavelengths was investigated in various studies (bucini et al. 2010; higginbottom et al. 2018; main et al. 2016; urbazaev et al. 2015). however, the reproducibility of these applications might be limited because of the utilisation of eo data, which are not systematically acquired or cost intensive (e.g. japan aerospace exploration agency advanced land observing satellite [jaxa’s alos], aerial images and so on). the goal of this article was to introduce a workflow to derive woody cover information from freely available sentinel-1a/b time series and light detection and ranging (lidar) data (smit et al. 2016). this workflow was applied to produce a wall-to-wall woody cover map in different spatial resolutions for the knp for 2016 and 2017. the estimation of woody cover by using c-band data has shown great potential in the south african lowveld region (where the knp is located) because of deeper penetration of the radar signals into the open savanna vegetation (mathieu et al. 2013). the workflow is publicly available to allow the continuation of woody cover monitoring, transferability to other savanna regions and future woody cover change mapping. data and methods sentinel-1 data the wall-to-wall woody cover estimation for the knp was carried out by using dense time series information from sentinel-1a/b c-band data between january 2016 and april 2017. the sentinel-1 data were retrieved via the copernicus open access hub (https://scihub.copernicus.eu/ – copernicus sentinel data [2016, 2017]). the data sets were acquired in ground range detected (grd) format from relative orbit 145, which covers the entire knp within one overpass. the temporal resolution of the time series is 12 days and its spatial resolution is 10 m. sentinel-1 sar backscatter in vv (vertical–vertical/co-polarisation) and vh (vertical–horizontal/cross-polarisation) were used as inputs for the woody cover prediction model. the sentinel-1 data were pre-processed by using gamma routines (wegmüller et al. 2016) implemented in pyrosar, a free python framework for large-scale sar satellite data processing (truckenbrodt et al. 2019), which provides a user-friendly solution for the pre-processing of sar satellite data from recent and historical eo missions. pyrosar also offers the possibility to use open-source software packages (e.g. esa’s sentinel application platform [snap]), resulting in comparable backscatter information in comparison with gamma. the pre-processing steps for the sentinel-1 data include (1) radiometric calibration to convert from digital values to radar backscatter; (2) orthorectification, utilising height information from the freely available shuttle radar topographic mission (srtm) (united states geological survey [usgs] 2017) with a spatial resolution of 30 m and the precise orbit state vectors (precise orbit ephemerides [poe]); and (3) terrain flattening utilising srtm to correct for radiometric differences caused by local incidence angles (small 2011). light detection and ranging data a lidar data set with a spatial resolution of 2 m representing vegetation heights was reclassified to woody cover estimates. these were used as input for the random forest (rf) modelling. the data set is available for downloading through the carnegie airborne observatory (cao) maps server (smit et al. 2016). the lidar data were acquired by the cao (asner et al. 2007) at the end of wet season in april/may 2010, 2012 and 2014, respectively, and covered a rectangular footprint of approximately 52 km² (2 km × 26 km) (smit et al. 2016). for this study, we used the data from 2014, to reduce the time gap between the training data and sentinel-1 time series and thus diminish the influence of land cover changes. it needs to be mentioned that abrupt disturbances (e.g. fire, herbivory and drought) during that time are influencing the woody cover estimation. an independent validation was carried out using a lidar data set collected in may 2012, which covers the areas (500 m of each riverfront) along the sabie, olifants and letaba river in the knp (milan, heritage & tooth 2018). the conversion from lidar vegetation heights to woody cover percentages was carried out for both data sets identically, which is dealt with in the next section. methodology preparation of training and prediction data as a first step, both lidar canopy height model (chm) was converted to percentage woody cover. we assumed that each pixel with a height value below 0.5 m represents ground or grassland pixels, whereas all other pixels represent woody cover pixels (smit et al. 2016; urban et al. 2018). the resulting binary mask (0 = non-woody, 1 = woody) was then used to calculate the woody cover percentage for a regular grid with a cell size congruent to the spatial resolution of the sentinel-1 (10 m) (sum of the woody pixel divided by the amount of lidar pixel fitting in one sentinel-1 cell and multiplied by 100). the individual sentinel-1 images were co-registered, mosaicked for the area of the knp and combined into a layer stack for each polarisation (vv and vh), respectively. both sentinel-1 stacks (predictor variables) were then fused with the lidar woody cover (response variable) and utilised as input data for the training of the rf model (figure 1). figure 1: the methodological framework for deriving woody cover information from sentinel-1 time series by using light detection and ranging information (scripts 1–4 are freely available via zenodo.org). random forest model the woody cover estimation was performed utilising the decision tree classifier rf (breiman 2001) through an implementation of the statistical software r (machine learning in r [mlr] package [bischl et al. 2016]). the modelling workflow included (1) tuning, (2) spatial cross-validation, (3) training and (4) prediction (figure 1). during model fitting, the independent decision trees are controlled by two main parameters: mtry, which describes the number of available predictor variables to split each node, and ntrees, the number of regression trees (breiman 2001). these so-called ‘hyperparameters’ were optimised (tuning) before model fitting, as they were found to be the most sensitive variables with respect to accuracy and computation time, to identify the best parameter set for the spatial cross-validation and training (heckel et al. 2020; huang & boutros 2016). the tuning was accomplished for mtry [1, 2] and ntrees [10, 50, 100, 300, 700] with five repetitions and internal spatial cross-validation for each parameter combination. with an increasing mtry value, the runtime of the model rises likewise because of the large number of observations. therefore, we conducted a grid search approach by using a priori knowledge of the hyperparameter search space utilising only a limited number of mtry values. the tuning revealed that the optimal parameter set, resulting in the highest accuracies and best model computation duration ratio, is mtry = 1 and ntrees = 300. according to this, spatial cross-validation (brenning 2012) and training of the model were performed utilising these parameters. we used a spatial cross-validation with five folds and 100 repetitions to ensure the derivation of stable results, for example, consistent accuracies. the trained model was utilised for the prediction on the sentinel-1 time series for the entire knp to produce the first wall-to-wall woody cover map for the year 2016/2017 at spatial resolutions of 10 m, 30 m, 50 m and 100 m. the r scripts as well as example data and the final woody cover product are freely available through zenodo.org for future usage. ethical consideration i confirm that ethical clearance was not needed/required for the study. results the woody cover map, derived from sentinel-1 time series data acquired between 2016 and 2017, is shown in figure 2. the estimations for woody cover range from 0% to 100%. the highest woody cover is reached in the northern part and the mopane-dominated landscapes on granite in the mid-west of the knp as well as the mixed thorn and woodland areas in the southern part (30% – 50% and for some regions even higher than 50% on average). lowest woody cover values were found in the open grasslands on basalt in the eastern part (below 20% on average). these broad-scale spatial patterns indicate that woody cover in knp is significantly controlled by the underlying geology and the north-to-south rainfall gradient (venter et al. 2003). finer-scale contrasts in woody cover are also clearly discernable on the woody cover map. for example, the contrast between the lower woody cover gabbro areas within the higher woody cover granitic areas and the more open sodic areas as opposed to higher woody cover riparian zones are clearly visible. figure 2: wall-to-wall woody cover map of the kruger national park with a spatial resolution of 10 m (right). the footprints of the light detection and ranging data used for training (red, smit et al. 2016) and independent validation (orange, milan et al. 2018) are shown in the woody cover map. comparison of the woody cover information to high-resolution imagery from google earth for three selected regions (a–c) as well as a scatterplot visualising the independent validation of the woody cover map with a woody cover estimate based on light detection and ranging data from milan et al. (2018) (lower left). the rf approach revealed that the sentinel-1 time series data from the dry season of 2016 were the most important predictors for deriving woody cover information. furthermore, the cross-polarised scenes between july and september 2016 have been identified to have the highest ranking in the variable importance, followed by the co-polarised scenes of the same time span. this implies that sentinel-1 predictors representing the dry season are key variables for retrieving woody cover in heterogeneous savanna ecosystems. this is most certainly caused by a lower impact of surface moisture on the c-band backscatter as well as a better separation between woody plants and grasses during the dry season (bucini et al. 2010; heckel et al. 2020; urbazaev et al. 2015). the spatial cross-validation, which was performed on the training data, resulted in a root mean squared error (rmse) of 22.8% for the product with a spatial resolution of 10 m, and improved with spatial averaging to 15.8% for 30 m, 14.8% for 50 m and 13.4% for 100 m as has been found in other radar studies such as santoro et al. (2013). the spatial cross-validation revealed coefficient of determination values of r2 = 0.3 (10 m), r2 = 0.5 (30 m), r2 = 0.6 (50 m) and r2 = 0.6 (100 m), respectively. these error estimates are slightly lower in comparison with the accuracy of other woody cover products with spatial resolution ranging between 50 m and 120 m (bucini et al. 2010; higginbottom et al. 2018; main et al. 2016; urbazaev et al. 2015), with rmse of around 10%. in comparison with this study, these products are lacking a spatially weighted cross-validation (except higginbottom et al. 2018), resulting in modelling accuracies that do not take into account effects of spatial autocorrelation (brenning 2012). an additional independent validation was carried out by using woody cover estimates based on the lidar data from milan et al. (2018), confirming the results of the spatial cross-validation of the model (figure 2 – scatterplot in the lower left for the spatial resolution of 10 m). furthermore, the following rmse and r2 are found for the other spatial resolutions: 30 m (rmse = 19%, r2 = 0.5), 50 m (rmse = 18%, r2 = 0.6) and 100 m (rmse = 16%, r2 = 0.6). discussion the presented workflow was applied to produce a wall-to-wall woody cover map in different spatial resolutions (e.g. 10 m, 30 m, 50 m and 100 m) for the knp for 2016 and 2017, which are freely available via a data repository on zenodo.org. the user can choose between different spatial resolutions based on their needs. the woody cover maps provide insightful information for sustainable park management and future planning and conservation-related activities. the overestimation of woody cover below 20% is a typical phenomenon in radar retrievals (mathieu et al. 2013; santoro et al. 2011) caused by surface contributions to the signal, such as roughness. underestimation for woody cover above 60% has been observed likewise in other studies (bouvet et al. 2018) and is explained by saturation of the c-band backscatter in dense canopies. the presented workflow shows the capability of sentinel-1 radar time series to retrieve fine-scale woody cover estimates in savanna ecosystems at regional scale. we found similar error estimates when comparing the accuracies to other woody cover products (e.g. bucini et al. 2010; higginbottom et al. 2018; main et al. 2016; urbazaev et al. 2015) at comparable spatial resolutions. this article describes the approach to derive woody cover from sentinel-1 time series based on open-source software and open-access data to foster the reproducibility of the framework for individual research interests as well as to facilitate the potential of transferring the presented methodology to other savanna ecosystems. it is worth mentioning that radar data are sensitive to environmental conditions, e.g., surface roughness and surface moisture, which need to be considered in future applications utilising this approach. moreover, the definition of a height threshold for woody cover estimation, to exclude the non-woody vegetation component, might require an adjustment in other regions. we are currently focussing on comparing woody cover maps of different years to build up a woody cover change-monitoring system, where the mentioned environmental factors are analysed. in the near future, such a monitoring system will likely benefit from open access lidar data from the global ecosystem dynamics investigation (gedi) (coyle et al. 2015) on board of the international space station (iss) and holds the potential to become an essential component of park management in the knp and other national parks in south africa. acknowledgements competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions the processing, analysis, interpretation of the used data and the preparation of the manuscript were completed by the first author m.u. k.h., c.b. and p.s. supported the analysis and interpretation of the data as well as proofreading of the manuscript. i.s. and t.s. supported during writing of the manuscript and proofreading. j.b. and c.s. supported the interpretation of the results and revised the manuscript. funding information the study was funded by the project south african land degradation monitor (saldi) sponsored by the german federal ministry of education and research (bmbf) in the framework of the science partnerships for the assessment of complex earth system processes (spaces ii) under the grant 01ll1701a. it was also partly funded by the european union (eu) horizon 2020 research and innovation program under the grant agreement no. 640176 (baci – towards a biosphere atmosphere change index). co-authors received funding from the ars africae and emsafrica research projects, which are part of the spaces and spaces ii initiatives and funded by the bmbf (grant numbers 01ll1303d and 01ll1801d) and the project munich center for machine learning (mcml) from the bmbf (grant number 01is18036a). substantial scientific and logistical support in the knp was received through our two sanparks projects ars africae eo (earth observation) (ref. no. schcc1235) and co-ld-ems (ref. no. baaj1519). the carnegie airborne observatory (cao) lidar data collection from smit et al. (2016) was funded by the andrew mellon foundation. the cao was supported by the avatar alliance foundation, john d. and catherine t. macarthur foundation, andrew mellon foundation, mary anne nyburg baker and g. leonard baker jr., and william r. hearst iii. the lidar data for independent validation of the woody cover map were provided by g. heritage (university of salford, uk) and d. milan (university of hull, school of environmental sciences, uk). data availability statement the programming code (statistical software r), sentinel-1 and lidar training data as well as data sets for model prediction of three selected regions in the knp are freely available via the following link: https://doi.org/10.5281/zenodo.3728186. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references asner, g.p., knapp, d.e., kennedy-bowdoin, t., jones, m.o., martin, r.e., boardman, j.w. et al., 2007, ‘carnegie airborne observatory: in-flight fusion of hyperspectral imaging and waveform light detection and ranging for three-dimensional studies of ecosystems’, journal of applied remote sensing 1(1), 013536. https://doi.org/10.1117/1.2794018 attema, e., davidson, m., snoeij, p., rommen, b. & floury, n., 2009, ‘sentinel-1 mission overview’, in international geoscience and remote sensing symposium (igarss), cape town, south africa, july 12–17, 2009, vols i–36, 1–4. berger, c., bieri, m., bradshaw, k., brümmer, c., clemen, t., hickler, t. et al., 2019, ‘linking scales and disciplines: an interdisciplinary cross-scale approach to supporting climate-relevant ecosystem management’, climatic change 156(1–2), 139–150. https://doi.org/10.1007/s10584-019-02544-0 bischl, b., lang, m., kotthoff, l., schiffner, j., richter, j., studerus, e. et al., 2016, ‘mlr: machine learning in r’, journal of machine learning research 17(170), 1–5. bouvet, a., mermoz, s., toan, t.l., villard, l., mathieu, r., naidoo, l. et al., 2018, ‘an above-ground biomass map of african savannahs and woodlands at 25 m resolution derived from alos palsar’, remote sensing of environment 206, 156–173. https://doi.org/10.1007/s10584-019-02544-0 breiman, l., 2001, ‘random forests’, machine learning 45(1), 5–32. https://doi.org/10.1023/a:1010933404324 brenning, a., 2012, ‘spatial cross-validation and bootstrap for the assessment of prediction rules in remote sensing: the r package sperrorest’, in international geoscience and remote sensing symposium (igarss), munich, germany, july 22–27, 2012, pp. 5372–5375. bucini, g., hanan, n., boone, r., smit, i., saatchi, s., lefsky, m. et al., 2010, ‘woody fractional cover in kruger national park, south africa: remote sensing–based maps and ecological insights’, in m.j. hill & n.p. hanan (eds.), ecosystem function in savannas, pp. 219–237, crc press, boca raton, fl. buitenwerf, r., bond, w.j., stevens, n. & trollope, w.s.w., 2012, ‘increased tree densities in south african savannas: savannas: > 50 years of data suggests co2 as a driver’, global change biology 18(2), 675–684. https://doi.org/10.1111/j.1365-2486.2011.02561.x coyle, d.b., stysley, p.r., poulios, d., clarke, g.b. & kay, r.b., 2015, ‘laser transmitter development for nasa’s global ecosystem dynamics investigation (gedi) lidar’, society of photo-optical instrumentation engineers (spie) 9612, 12th 13th august 2015, lidar remote sensing for environmental monitoring xv, 961208, pp. 1–7, san diego, ca. druce, d.j., shannon, g., page, b.r., grant, r. & slotow, r., 2008, ‘ecological thresholds in the savanna landscape: developing a protocol for monitoring the change in composition and utilisation of large trees’, plos one 3(12), e3979. https://doi.org/10.1371/journal.pone.0003979 ferreira, s.m., greaver, c., knight, g.a., knight, m.h., smit, i.p.j. & pienaar, d., 2015, ‘disruption of rhino demography by poachers may lead to population declines in kruger national park, south africa’, b.l. allen (ed.), plos one 10(6), e0127783. https://doi.org/10.1371/journal.pone.0127783 heckel, k., urban, m., schratz, p., mahecha, m.d. & schmullius, c., 2020, ‘predicting forest cover in distinct ecosystems: the potential of multi-source sentinel-1 and -2 data fusion’, remote sensing 12(2), 302. https://doi.org/10.3390/rs12020302 higginbottom, t.p., symeonakis, e., meyer, h. & van der linden, s., 2018, ‘mapping fractional woody cover in semi-arid savannahs using multi-seasonal composites from landsat data’, isprs journal of photogrammetry and remote sensing 139, 88–102. https://doi.org/10.1016/j.isprsjprs.2018.02.010 huang, b.f.f. & boutros, p.c., 2016, ‘the parameter sensitivity of random forests’, bmc bioinformatics 17(1), 331. https://doi.org/10.1186/s12859-016-1228-x kiker, g.a., scholtz, r., smit, i.p.j. & venter, f.j., 2014, ‘exploring an extensive dataset to establish woody vegetation cover and composition in kruger national park for the late 1980s’, koedoe 56(1), 1–10. https://doi.org/10.4102/koedoe.v56i1.1200-1 kutsch, w., hanan, n., scholes, b., mchugh, i., kubheka, w., eckhardt, h. et al., 2008, ‘response of carbon fluxes to water relations in a savanna ecosystem in south africa’, biogeosciences 5, 1797–1808. https://doi.org/10.5194/bg-5-1797-2008 main, r., mathieu, r., kleynhans, w., wessels, k., naidoo, l. & asner, g.p., 2016, ‘hyper-temporal c-band sar for baseline woody structural assessments in deciduous savannas’, remote sensing 8(8), 1–19. https://doi.org/10.3390/rs8080661 mathieu, r., naidoo, l., cho, m.a., leblon, b., main, r., wessels, k. et al., 2013, ‘toward structural assessment of semi-arid african savannahs and woodlands: the potential of multitemporal polarimetric radarsat-2 fine beam images’, remote sensing of environment 138, 215–231. https://doi.org/10.1016/j.rse.2013.07.011 milan, d., heritage, g. & tooth, s., 2018, dataset record: kruger national park rivers lidar data (may 2012), centre for environmental data analysis, didcot. pettorelli, n., wegmann, m., skidmore, a., mücher, s., dawson, t.p., fernandez, m. et al., 2016, ‘framing the concept of satellite remote sensing essential biodiversity variables: challenges and future directions’, remote sensing in ecology and conservation 2(3), 122–131. https://doi.org/10.1002/rse2.15 santoro, m., beer, c., cartus, o., schmullius, c., shvidenko, a., mccallum, i. et al., 2011, ‘retrieval of growing stock volume in boreal forest using hyper-temporal series of envisat asar scansar backscatter measurements’, remote sensing of environment 115(2), 490–507. https://doi.org/10.1016/j.rse.2010.09.018 santoro, m., cartus, o., fransson, j.e.s., shvidenko, a., mccallum, i., hall, r.j. et al., 2013, ‘estimates of forest growing stock volume for sweden, central siberia, and québec using envisat advanced synthetic aperture radar backscatter data’, remote sensing 5(9), 4503–4532. https://doi.org/10.3390/rs5094503 scholes, r.j. & archer, s.r., 1997, ‘tree-grass interactions in savannas’, annual review of ecology and systematics 28(1), 517–544. https://doi.org/10.1146/annurev.ecolsys.28.1.517 scholes, r.j. & walker, b.h., 1993, an african savanna: synthesis of the nylsvley study, cambridge university press, cambridge. skowno, a.l., thompson, m.w., hiestermann, j., ripley, b., west, a.g. & bond, w.j., 2016, ‘woodland expansion in south african grassy biomes based on satellite observations (1990–2013): general patterns and potential drivers’, global change biology 23(6), 2358–2369. https://doi.org/10.1111/gcb.13529 small, d., 2011, ‘flattening gamma: radiometric terrain correction for sar imagery’, ieee transactions on geoscience and remote sensing 49(8), 3081–3093. https://doi.org/10.1109/tgrs.2011.2120616 smit, i.p.j., asner, g.p., govender, n., vaughn, n.r. & van wilgen, b.w., 2016, ‘an examination of the potential efficacy of high-intensity fires for reversing woody encroachment in savannas’, journal of applied ecology 53(5), 1623–1633. https://doi.org/10.1111/1365-2664.12738 smit, i.p.j. & prins, h.h.t., 2015, ‘predicting the effects of woody encroachment on mammal communities, grazing biomass and fire frequency in african savannas’, m.s. crowther (ed.), plos one 10(9), e0137857. https://doi.org/10.1371/journal.pone.0137857 smit, i.p.j., smit, c.f., govender, n., van der linde, m. & macfadyen, s., 2012, ‘rainfall, geology and landscape position generate large-scale spatiotemporal fire pattern heterogeneity in an african savanna’, ecography 36(4), 447–459. https://doi.org/10.1111/j.1600-0587.2012.07555.x stevens, n., erasmus, b.f.n., archibald, s. & bond, w.j., 2016, ‘woody encroachment over 70 years in south african savannahs: overgrazing, global change or extinction aftershock?’, philosophical transactions of the royal society b: biological sciences 371(1703), 20150437. https://doi.org/10.1098/rstb.2015.0437 truckenbrodt, j., cremer, f., baris, i. & eberle, j., 2019, ‘pyrosar: a framework for large-scale sar satellite data processing’, in proc. of the 2019 conference on big data from space (bids’19), munich, germany, february 19–21, 2019, pp. 197–200. urban, m., berger, c., mudau, t.e., heckel, k., truckenbrodt, j., odipo, v.o. et al., 2018, ‘surface moisture and vegetation cover analysis for drought monitoring in the southern kruger national park using sentinel-1, sentinel-2, and landsat-8’, remote sensing 10(9), 1482. https://doi.org/10.3390/rs10091482 urbazaev, m., thiel, c., mathieu, r., naidoo, l., levick, s.r., smit, i.p.j. et al., 2015, ‘assessment of the mapping of fractional woody cover in southern african savannas using multi-temporal and polarimetric alos palsar l-band images’, remote sensing of environment 166, 138–153. https://doi.org/10.1016/j.rse.2015.06.013 usgs (united states geological survey), 2017, shuttle radar topography mission (srtm) 1 arc-second global – version 3. usgs, sioux falls. venter, f.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation pattern’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience. ecology and management of savanna heterogeneity, pp. 83–129, island press, washington, dc. wegmüller, u., werner, c., strozzi, t., wiesmann, a., frey, o. & santoro, m., 2016, ‘sentinel-1 support in the gamma software’, procedia computer science 100, 1305–1312. https://doi.org/10.1016/j.procs.2016.09.246 koedoe 64-1_2022_contents.indd http://www.koedoe.co.za open access table of contents original research diversity and distribution of benthic invertebrates dwelling rivers of the kruger national park, south africa nabil majdi, lizaan de necker, hendrika fourie, ané loggenberg, edward c. netherlands, jasmine bunte-tschikin, walter traunspurger, gerhard c. du preez koedoe | vol 64, no 1 | a1702 | 27 june 2022 original research phylogenetic placement of the enigmatic floodplain water snake, lycodonomorphus obscuriventris fitzsimons, 1964 chad keates, werner conradie, tatenda dalu, farai dondofema, eddie s. riddell, ryan j. wasserman koedoe | vol 64, no 1 | a1698 | 28 june 2022 original research the potential of frogging as an ecotourism product for south african national parks zoëgné luyt, peet van der merwe koedoe | vol 64, no 1 | a1725 | 21 july 2022 original research the pongola bush: tree diversity assessment in a kwazulu-natal forest patch timothy f. hall, mary c. scholes, robert j. scholes koedoe | vol 64, no 1 | a1724 | 20 september 2022 original research people parks win-win framework: integrating components that can influence people-park relationships dorothy r. queiros koedoe | vol 64, no 1 | a1723 | 26 september 2022 reviewer acknowledgement koedoe | vol 64, no 1 | a1739 | 19 december 2022 50 68 77 88 96 112 page i of i table of contents editorial a reflection about the recent koedoe publications (volume 64, no 1, 2022) abel ramoelo koedoe | vol 64, no 1 | a1736 | 19 december 2022 short communication characterising the geomorphic dynamics of river systems: an example of the sabie river, south africa jasper knight, mary evans koedoe | vol 64, no 1 | a1700 | 29 march 2022 review article reflecting on research produced after more than 60 years of exclosures in the kruger national park corli wigley-coetsee, tercia strydom, danny govender, dave i. thompson, navashni govender, judith botha, chenay simms, adolf manganyi, laurence kruger, jacques venter, cathy greaver, izak p. smit koedoe | vol 64, no 1 | a1674 | 28 february 2022 review article the ephemeral pans of gras-holpan: mokala national park, northern cape, south africa nkabeng t. mzileni, hendrik sithole, hugo bezuidenhout, roxanne erusan, rodney makwakwa koedoe | vol 64, no 1 | a1709 | 09 december 2022 original research evidence that residues of tebuthiuron arboricide present in soil of mokala national park can be phytotoxic to woody and grass species carl f. reinhardt, hugo bezuidenhout, judith m. botha koedoe | vol 64, no 1 | a1658 | 18 march 2022 original research challenges and opportunities for sustainable solid waste management in private nature reserves: the case of sabi sand wildtuin, south africa claudine roos, reece c. alberts, francois p. retief, dirk p. cilliers, william hodgson, iain olivier koedoe | vol 64, no 1 | a1710 | 26 may 2022 1 3 9 23 32 41 vol 64, no 1 (2022) issn: 0075-6458 (print) | issn: 2071-0771 (online)koedoe filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 castley.qxd searching for common ground, a scientific approach to subjective environmental impact assessments: an example from the kgalagadi transfrontier park j.g. castley, h. bezuidenhout and m.h. knight castley, j.g., h. bezuidenhout and m.h. knight. 2003. searching for common ground, a scientific approach to subjective environmental impact assessments: an example from the kgalagadi transfrontier park. koedoe 46(1): 107–114. pretoria. issn 0075-6458. in order to break away from the frequent subjective nature of assessing potential environmental impacts and alternative scenarios when undertaking new developments a structured scientific approach was adopted in the present analysis. an assessment of the proposed upgrading of a 31-km stretch of road between leeudril and kij-kij waterholes on the dry nossob river in the kgalagadi transfrontier park (ktp) was undertaken. relocation of the existing road was proposed as an alternative to upgrading the road as the potential and future impacts on the riverbed ecosystem could be reduced. a systematic assessment of biophysical variables within the riverbed habitat was made. a multivariate clustering analysis revealed a high level of congruency with a purely subjective assessment that potentially supports an “apples and oranges” comparison of such approaches in environmental assessments. furthermore, the present and potential environmental impacts of each of the various upgrading options were compared. significant environmental impacts were envisaged when considering surface and ground water hydrology, flora and the sensitive nature of the landscape. given the requirements of the park in terms of providing a tourism product the preferred action in terms of upgrading the road would require a compromise between ecological and aesthetic requirements in order to provide reasonable tourism opportunities. the scientific approach adopted in the current analysis appears to offer a more defendable result upon which management decisions can be based compared to purely subjective assessments but is not without limitations. key words: environmental impact assessments, kalahari, dry riverbed, road. j.g. castley and m.h. knight, conservation services, south african national parks, p.o. box 20419, humewood, 6013 republic of south africa (gcastley@upe.ac.za; mknight@upe.ac.za); h. bezuidenhout, conservation services, south african national parks, p.o. box 110040, hadison park, 8306 republic of south africa(hugob@parkssa.co.za). issn 0075-6458 107 koedoe 46/1 (2003) introduction roads affect ecosystems, biological communities and species in numerous and different ways, determined largely by their location, density, and distribution across the landscape (hourdequin 2000). roads generally have negative effects on the biotic integrity in both terrestrial and aquatic ecosystems, and these effects can be classified under seven categories (trombulak & frissell 2000): increased mortality from road construction; increased mortality from vehicle collisions; news and views castley.qxd 2005/12/09 10:23 page 107 modification of animal behaviour; alteration of the physical environment; alteration of the chemical environment; spread of exotic species; and increased alteration and use of the habitats by humans. in the kgalagadi transfrontier park (ktp) where most of the tourist road network is constructed in the dry riverbeds of the nossob and auob, the impact of the roads are likely to be limited to a number of the above categories, particularly the physical and chemical properties of the surrounding ecosystems, and to a lesser extent on the mortality of animals (although plant mortality has yet to be quantified) and the modification of animal behaviour (but again where the road is in a vicinity of breeding sites this may create a problem for breeding birds). the precise ecological effects of roads on biota may be uncertain but the selective removal, relocation and rehabilitation of roads may provide certain ecological benefits, although mitigatory measures to address these impacts may be costly, vague and have a negative impact themselves (trombulak & frissell 2000). roads are known to alter at least eight physical characteristics of the environment, such as: soil density, temperature, soil water content, light penetration, dust production, surface water flow, run-off pattern and sedimentation. dust is a major factor to consider as this can settle on plants and block photosynthesis, respiration and transpiration in addition to causing physical injuries of plants (farmer 1993), also acting as a possible deterrent to grazing. roads on floodplains can redirect water, sediments and nutrient dispersal between streams and wetlands and their riparian ecosystems to the detriment of water quality and ecosystem health (trombulak & frissel 2000). the dry riverbed systems of the ktp may be no different to these systems. the development of road networks within protected areas to enhance the tourism potential or act as management tools (such as firebreaks) is a generally accepted practise (pienaar 1968). however, such projects also have dramatic impacts on biodiversity conservation (miller et al. 1995). fortunately, the widespread adoption of environmental impact assessments (eias) has allowed such developments to be assessed in terms of their possible impacts on the environment and the need to assess alternatives (deat 1998). however, few of the experts building or ‘restoring’ roads are trained to recognise and address the full spectrum of ecological issues associated with road construction (trombulak & frissell 2000). the existing 490 km of tourist roads on the south african side of the ktp (formerly the kalahari gemsbok national park) are largely restricted to the dry nossob and auob riverbeds, owing to the flat consolidated surfaces, and lack of permanent water (van wyk & le riche 1984; mills & retief 1984), combined with the difficulty of constructing roads in the unconsolidated sand dunes. furthermore, the higher nutrient status of the riverbed habitat makes it the single most important habitat in the park supporting a large proportion of the indigenous ungulate populations (mills & retief 1984; knight et al. 1987). in addition, the characteristically large trees in the riverbed provide important nesting sites for most of the large raptors species (herholdt 1995). the riverbeds, therefore, offer the best wildlife viewing potential in the park, and with the riverine habitat comprising about 4 % of the 36 000 km² large ktp, it is undesirable that a relatively large proportion of it is taken up by road surface. along the entire length of the auob river and the nossob river north of melkvlei waterhole the road has been largely moved out of the riverbed onto the calcrete footslope. however, along the 45 km section along the nossob river between twee rivieren and melkvlei, the road has remained in the riverbed owing to high dunes on the western bank and calcrete cliffs on the botswanan eastern bank. this particular section, and the focus of the study, remains a management problem as the prekoedoe 46/1 (2003) 108 issn 0075-6458 castley.qxd 2005/12/09 10:23 page 108 sent ungravelled road has been lowered below the riverbed surface through continual maintenance grading. this has made it particularly prone to flooding, making it inaccessible to tourist traffic for extended periods during the summer months. in the case of the twee rivieren to melkvlei road, a number of options exist with regards to altering the road configuration within the riverbed, such as: 1) upgrading the present road within the riverbed, 2) moving the road onto the western or eastern river margins, 3) moving the road onto the eastern calcrete terrace, 4) or a combination of the above alternatives. the option of re-routing the road onto the western bank was disregarded at the outset as this particular section of road is characterised by high sand dunes orientated at right angles to the riverbed along its entire length. the eastern alternatives are more suitable owing to the level nature of the calcrete terrace but one complication is the fact that the nossob river forms an international boundary between south africa and botswana. furthermore, the selection of options is also affected by the desire to provide varied tourist experiences. the aim of this paper is to assess the most likely options proposed for improving the road alignment through the consideration of a combination of expected physical impacts and diverse tourism experiences, through the interpretation of two different approaches, that of a normal subjective assessment and an objective assessment in the analysis of environmental impacts while also providing a platform for future decision making supported by defendable scientific results. description of the habitat the nossob river south of kij-kij is characterised by a relatively narrow (100–500 m across) valley, with the riverbed margins dominated by high dunes on the western south african side and calcrete terraces on the eastern botswanan side. towards the north the nossob changes to a wide, shallow, sandy riverbed and is more than 1 km wide in places (bothma & de graaff 1973). the silt and clayey soils (oakleaf and valsrivier soil forms) of the riverbeds give way to a combination of more rocky (coega and prieska soil forms) and shallow sandy soils (clovelly soil form) along the riverbed margins (van rooyen 1984; soil classification working group 1991). the calcrete banks and ridges of the kalahari group (malherbe 1984), are usually steep (10°–15°) with vertical cliffs a few metres high in the upper zones (parris 1984). however, areas comprising red, brown, yellow, cream and green sandstones, grits as well as lenses of conglomerates are also present (malherbe 1984). the red sandy soils associated with the large dunefield are notably infertile while the pans and dry riverbeds are more fertile owning to the higher concentrations of soluble salts and exchangeable cations such as sodium, calcium, magnesium and potassium (leistner 1959). vegetation along the riverbed margins comprises scattered acacia erioloba trees with rhigozum trichotomum, acacia hebeclada and galenia africana shrubs and the dominant grasses schmidtia kalahariensis and panicum coloratum. the plant communities in the section of particular interest in the lower nossob riverbed have been described as rhigozum trichotomum galenia africana (2b) riverine vegetation, while the adjacent dune habitat is described as an acacia erioloba boscia albitrunca (8b) tree savanna (lubbinge 1999). methods the assessment of environmental impacts is often a subjective exercise given the diverse backgrounds of individual reviewers. in an attempt to determine whether a more rigorous approach to such assessments could be adopted a single survey of the lower nossob river was undertaken by the scientific and technical services staff of the south african national parks (sanparks) in december 1998. twenty-four sites distributed evenly along the 31-km stretch of road were surveyed. two methods were adopted in the current analysis. a purely subjective assessment, based on whether it would be desirable to have the road within the riverbed at any particular issn 0075-6458 109 koedoe 46/1 (2003) castley.qxd 2005/12/09 10:23 page 109 site, was made using aesthetic criteria such as the scenic quality, width of the riverbed, density of trees, position of the road in the riverbed, the accessibility to the riverbed margin and perceived environmental impact. in comparison, a more rigorous assessment of each site was made by measuring a series of criteria (table 1) that were assigned a priori values considered desirable with respect to removing the road from the riverbed. criteria were either ranked or assigned an absolute value and then compared to that required to remove the road using multivariate statistical analysis. a bray-curtis agglomerative hierarchical clustering procedure that determined the similarity among sites in relation to the desirable state was undertaken to provide supporting evidence for management decisions. criteria that determined whether the road should be removed from the riverbed included: 1) the existing road was in the centre of the riverbed; 2) the riverbed was narrow (<50 m); 3) the slope and height of the riverbed margin habitat would allow a road to be taken out; 4) the straight line visual distance of the existing road was greater than 500 m in either direction; 5) the existing road was below the current level of the riverbed; 6) high side-walls were present; 7) there were a large number of a. erioloba trees within the riverbed system; and 8) there were a number of alternative options and the potential to rehabilitate the riverbed was high. results and discussion comparison between three options in jointly assessing the potential impacts of the three options (in riverbed, river margins or on calcrete) open to providing a road up the nossob river, the environmental screening revealed that from an ecological perspective the best option would be to remove the road from the riverbed. although re-routing koedoe 46/1 (2003) 110 issn 0075-6458 table 1 criteria used in a quantitative assessment of sites along a 31-km stretch of the nossob riverbed, kgalagadi transfrontier park. ranks for certain criteria are in bold. the 'desired' column lists either ranks or values classified as meeting the requirements to remove the road from the riverbed criteria parameters format desired 1. position of road in riverbed botswana s. africa centre out ranked 3 1 2 3 4 2. width of riverbed (m) narrow intermediate wide absolute 50 50-80 80-120 > 120 3. slope of calcrete/dune margins flat shallow gentle steep ranked 1 1 2 3 4 4. height (m) of calcrete/dune margins absolute 2 5. straight line length (m) of road n/s north south absolute > 500 6. height of road above riverbed below level above absolute 0.3 7. height of road sidewalls none absolute 0.8 8. large trees and shade provision none few intermed many ranked 4 1 2 3 4 9. alternative road option possible no slight possible good ranked 4 1 2 3 4 10. potential for riverbed rehabilitation poor fair good ranked 3 1 2 3 castley.qxd 2005/12/09 10:23 page 110 the road onto the edge of the riverbank alleviated some of the problems, the best option would be to re-route the road onto the calcrete terraces along the eastern bank of the riverbed within botswana (table 2). many of the impacts were not considered to be significant in the long-term but three were identified where the potential impacts could be significant and included the impact on surface and ground water, and flora as summarised below: i) surface water raising the road surface above the adjacent riverbed surface could have a significant negative impact on the surface water flow owing to water collecting next to the road (currently water collects within the road and behind the road sidewalls). this could also possibly influence the sub-surface water flow and consequently vegetation growth and animal use of the habitat. in this regard the technical services thought the raising of the road would have no ecological impact with damming alleviated through the construction of series of shallow drifts. ii) ground water the compaction of the new road by heavy machinery as well as using gravel (a relatively foreign material in the predominantly clay based riverbed) for the upgrading process as opposed to alternative materials deposited on the road side-walls from grading activity was expected to have a negative issn 0075-6458 111 koedoe 46/1 (2003) table 2 potential impacts of road alignment on three alternatives. ‘-ve’, ‘+ve’refer to negative and positive impacts respectively while, st, lt refer to impactgs in the short and long term respectively. the significance of the impact is depicted by the box shading with low significance (light shading) progressing to high significance (dark shading) impacts riverbed river edge calcrete geology -ve -ve -ve soil -ve -ve +ve topography -ve -ve air quality -ve (st) +ve (lt) noise -ve (st) -ve -ve (st) surface water -ve -ve +ve (lt) ground water -ve (lt) +ve +ve (lt) land use -ve (lt) -ve +ve (lt) sensitive landscape -ve -ve +ve (lt) visual aspects -ve +ve archaeological sites -ve ? cultural interest -ve ? socio-economics ? ? i&ap’s -ve +ve (lt) fauna -ve -ve -ve (st) +ve (lt) flora -ve -ve -ve (st) +ve (lt) threatened species ? ? ? wilderness experience -ve -ve +ve (lt) game migration wildlife management -ve ecosystem functioning -ve (lt) ? -ve +ve tourism -ve (st) +ve (lt) +ve +ve (lt) castley.qxd 2005/12/09 10:23 page 111 impact on the subsurface ground water movement. although calcrete from the adjacent ridges and river banks is periodically assimilated within the riverbeds the use of gravel may affect the drainage patterns in the riverbed and consequently vegetation growth. the duration of these effects is also unknown although it could be reduced with time as sediments stabilise. the gravel requirements would also need to be met using local material and extensive burrow pits in the adjacent calcrete outcrops would have to be excavated. iii) flora the significant impact on the flora of the riverbed is largely influenced by the effects on the surface and ground water components as discussed above. should damming of the water continue irrespective of whether the road is elevated above the existing riverbed level then it is likely that there will continue to be clumping of vegetation alongside the road margins. however, should the water koedoe 46/1 (2003) 112 issn 0075-6458 fig. 1: bray-curtis clustering dendrogram identifying the associations between the 24 sites and a hypothetical control. flow be more evenly spread across the riverbed away from the road then there is a possibility that it may encourage vegetation growth within the riverbed. impacts on ecological processes were considered signifitable 3 comparison of 24 sites along the nossob river bed based on a subjective and objective analysis of aesthetic and bio-physical criteria site subjective objective definite removal indifferent leave road 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 castley.qxd 2005/12/09 10:23 page 112 cant although uncertain as to what the expected outcome may be, given the degree to which water was dispersed and allowed to penetrate into the subsoil. whatever the case, with the riverine vegetation considered the single most important habitat in the park, the ramifications would be more than likely long-term and negative. moving the road onto the river edge or bank was expected to only have a significant negative affect on the geology owing to the need to cross drainage lines, while having relatively minor negative impacts on most of the other bio-physical aspects, but a minor positive affect on tourism, aesthetic appeal and ground water movement. the calcrete terrace option offered mainly positive impacts on the soils, ground water, sensitive landscape, and tourism, with minor, short-term negative impacts on noise, fauna and flora. the noise element was considered to be a factor given the greater noise produced by driving on the gravelled surface. issn 0075-6458 113 koedoe 46/1 (2003) fig. 2: map depicting zones within the river bed which were identified based on the objective classification of bio-physical data. castley.qxd 2005/12/09 10:23 page 113 assessing re-routing options: “apples and oranges” although the option of removing the road out of the riverbed was considered the most desirable alternative, the need for a compromise between the perceived negative ecological impacts of maintaining the road in the river and the need for providing a variety of tourism viewing and experience options existed. in this context the selection of definite removal sites had to be identified. three broad groupings were identified within the cluster analysis approach for dealing with the road (fig. 1). eight sites (33 %) were associated with a 68 % similarity with the desirable state, and thus represented those positions along the riverbed where it was most desirable to remove the road. four sites (17 %) were identified as being acceptable to retain the road at its present alignment within the riverbed, while the remaining sites were considered to be acceptable either within the riverbed or moved out. the subjective assessment of whether or not the road should be removed from the riverbed produced a similar picture to the clustering analysis (table 3). twelve sites (50 %) were identified as candidate areas to re-route the road of which there was 70 % congruence with the quantitative analysis. however, the subjective approach identified a further four sites (numbers: 5, 6, 13, & 23) as not being ideal to keep the road within the riverbed while a further five sites (numbers: 1, 2, 12, 19, & 22) were identified as being acceptable to keep the road in the riverbed. the higher percentage contributions to either the retention or removal of the road may reflect the subjective nature of the assessment where one would be inclined to group sites within specific “yes / no” groups as opposed to an undecided grouping. the conformity of the two approaches supports the notion of using the statistical approach in selecting pertinent alternatives when one has to produce a defendable product against criticism of subjective bias. conclusions although the removal of the road entirely from the sensitive riverine habitat was considered the most ecologically responsible action there is a need to compromise the ecological and aesthetic requirements of providing reasonable tourism opportunities within the park. it is recommended that the road be re-routed in and out of the riverbed, avoiding sensitive sites as indicated in fig. 2. the use of an alternative quantitative approach in selecting these sensitive sites proved successful, making its recommendations more defendable than biased subjective alternatives. acknowledgements the management staff of the kgalagadi transfrontier park are thanked for their cooperation during the assessment. andre du plessis of south african national park’s technical department is also thanked for his input. valuable comments were received from two anonymous reviewers. references bothma, j. du p. & g. de graaff. 1973. a habitat map of the kalahari gemsbok national park. koedoe 16: 181-188. deat 1998. eia regulations. implementation of sections 21,22 & 26 of the environmental conservation act. pretoria: department of environmental affairs and tourism. farmer, a.m. 1993. the effects of dust on vegetation—a review. environmental pollution 79: 63–75. herholdt, j.j. 1995. the breeding status (19881994) and management of raptors in the kalahari gemsbok national park. unpublished m.tech thesis. pretoria technikon. hourdequin, m. 2000. introduction: ecological effects of roads. conservation biology 14: 16–17. knight, m.h., a.k. knight-eloff, & j.j. bornman. 1987. the importance of borehole water and lick sites to kalahari ungulates. journal of arid environments 15: 269–281. leistner, o.a. 1959. notes on the vegetation of the kalahari gemsbok national park with special reference to its influence on the distribution of antelopes. koedoe 2: 128–151. koedoe 46/1 (2003) 114 issn 0075-6458 castley.qxd 2005/12/09 10:23 page 114 lubbinge, j.w. 1999. a phytosociological analysis of the southern kalahari dunes. unpubl. m.sc. thesis, university of pretoria. malherbe, s.j. 1984. the geology of the kalahari gemsbok national park, 1931–1981. koedoe supplement: 33–44. miller, k., m.h. allegretti, n. johnson, & b. jonsson. 1995. measures for conservation of biodiversity and sustainable use of its components. pp. 915–1062. in: heywood v.h. & r.t. watson (eds.). global biodiversity assessment. cambridge: unep, cambridge university press mills, m.g.l. & p.f. retief. 1984. the response of ungulates to rainfall along the riverbeds of the southern kalahari, 1972–1982. koedoe supplement, 129–142. parris, r. 1984. pans, rivers and artificial waterholes in the protected areas of the south-western kalahari. koedoe supplement, 63–82. pienaar u. de v. 1968. the ecological significance of roads in a national park. koedoe 11:169–174. trombulak, s.c. & c.a. frissell. 2000. review of ecological effects of roads on terrestrial and aquatic communities. conservation biology 14: 18–30. van rooyen, t.h. 1984. the soils of the kalahari gemsbok national park. koedoe supplement 4562. van wyk p. & e.a.n. le riche. 1984. the kalahari gemsbok national park, 1931–1981. koedoe supplement: 21–33. issn 0075-6458 115 koedoe 46/1 (2003) castley.qxd 2005/12/09 10:23 page 115 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice abstract introduction ezemvelo kzn wildlife and public protected areas study area the data empirical model: the operational competitiveness rating analysis procedure the results and discussion conclusion and management implications acknowledgements references about the author(s) karabo d. motau discipline of agricultural economics, university of kwazulu-natal, south africa edilegnaw wale discipline of agricultural economics, university of kwazulu-natal, south africa citation motau, k.d. & wale, e., 2018, ‘the operational competitiveness of public protected areas managed by ezemvelo kzn wildlife’, koedoe 60(1), a1452. https://doi.org/10.4102/koedoe.v60i1.1452 original research the operational competitiveness of public protected areas managed by ezemvelo kzn wildlife karabo d. motau, edilegnaw wale received: 04 dec. 2016; accepted: 21 nov. 2017; published: 27 mar. 2018 copyright: © 2018. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the purpose of this study was to measure the operational competitiveness of public protected areas (ppas) in the kwazulu-natal province, south africa. financial data for ecotourism operations in ppas were collected from ezemvelo kzn wildlife (ekznw) for 2007–2013, to construct an operational competitiveness profile for each ppa by using a non-parametric method called operational competitiveness rating analysis (ocra). the results show that permanent staff, utilities, maintenance and repairs, and cost of sales were cost items with the highest average share of total costs, whereas accommodation, admissions, sales and tours, and rides and hikes received higher average shares of total revenues for most ppas. the identification of the most important cost and revenue items was followed by the computation of resource consumption and revenue generation inefficiency ratings from 2007 to 2013, with the results showing that resource competitiveness had more impact on operational competitiveness relative to revenue competitiveness. this suggests that ppas under ekznw can improve operational competitiveness by reducing costs. ecotourism is an economic incentive used in several countries to encourage biodiversity conservation. because of declining public funding, conservation agencies such as ekznw in south africa should find new sources of funding or find cost-effective ways of managing ecotourism operations. conservation implications: this information will provide insights into the quality of operational efficiencies of ecotourism activities at ekznw-controlled ppas and motivate management to adopt cost-cutting and revenue-increasing strategies to improve operational competitiveness. introduction the ownership of wildlife largely remains under state control in several countries, managed by conservation agencies (muir-leresche & nelson 2000). however, despite government protection of wildlife and biodiversity, a record number of species are classified as threatened, endangered or vulnerable by the international union for conservation of nature (iucn), because of pressures emanating from economic activities such as agriculture, mining, timber production and poaching (damania & hatch 2005). conservationists recognise that to prevent further deterioration, the preservation of many species rests on establishing their economic value and providing incentives for sustainable use (baker 1997; lindsey, romanach & davies-mostert 2009). ecotourism as an economic incentive has become a tool for biodiversity conservation in public protected areas (ppas) of many developing countries (lindsey et al. 2005). this is based on the principle that nature or biodiversity must pay for itself by generating economic benefits (kiss 2004). ecotourism supports biodiversity conservation and at the same time promotes sustainable local development (ross & wall 1999). in south africa, ecotourism is part of the sustainable development agenda, and it is viewed as an instrument of empowerment for underprivileged communities as it provides employment for rural communities (holden 2013). for instance, several protected areas, both public and private, have promoted joint economic initiatives whereby specific services and functions are outsourced to local communities (honey 2003; mahony & van zyl 2002; myburgh & saayman 1999). however, the significance of ecotourism is underrated, mainly because of a lack of information on the financial and economic performance of the ecotourism operations of ppas (barnes & de jager 1996; child et al. 2012; eagles 2003; musengezi 2010; porter, ferrer & aylward 2003). this could in turn lead to an under-representation of the significance of ecotourism within fiscal sectors of government. this, according to james, gaston and balmford (2001), could lead to a lack of understanding on the expenditure of biodiversity conservation and create a perception that conservation initiatives are unfeasible. as eagles (2003) states, this could result in cutbacks in funding for conservation agencies. given that funding for ppas has been inadequate and declining over the years (emerton, bishop & thomas 2006), research on the financial and economic performance of ecotourism operations of ppas will justify an investment in ppas. the shortfall in annual spending for ppas in developing countries is estimated to be in the range of $1 billion and $1.7 billion, which could be influenced by distinctive management objectives and activities (bruner, gullison & balmford 2004). moreover, easily accessible ppas face an increased threat of degradation and thus would require more investment. because of their respective unique desirable features, some ppas will generate more revenue than others, making cross-subsidisation inevitable. as ppas get larger, budgets have remained static, and therefore management costs per hectare in such ppas would normally be higher, especially in easily accessible ppas located close to disadvantaged communities (bruner et al. 2004; emerton et al. 2006). in south africa, funding for ezemvelo kzn wildlife (ekznw), kwazulu-natal’s wildlife agency, has been reduced by provincial government (khumalo & molla 2012). reduced funding coupled with low admission fees is creating financial constraints and undermine the capacity of the agency. eventually, this will compromise biodiversity conservation and local development. therefore, for conservation managers to find cost-effective ways of managing ecotourism operations, they will need empirical information on detailed evaluations of income and expenditure patterns of ecotourism operations. measuring the financial performance of ecotourism operations in ppas will enable ekznw to control and improve operational practices in the respective ppas. there are several operational performance measurement methods that have been employed in the literature to measure efficiency, prioritise conservation and justify investment in ppas in various contexts. the most important of these include appraisal methods such as cost-effective analysis (e.g. laycock et al. 2009; moran, pearce & wendelaar 1997), benefit–cost ratios (e.g. dixon & sherman 1991), cost–benefit analysis (e.g. dixon & sherman 1990) and so on. the various methods of comparing the costs and benefits of protected areas are summarised in dixon and sherman (1991). data envelopment analysis (dea) is a non-parametric method used to measure the relative efficiency of decision-making units (dmus) (charnes, cooper & rhodes 1978; speelman et al. 2008). it has often been employed to assess the relative efficiency of dmus of protected areas and to indicate how it could be improved by providing a set of guidelines (bosetti & locatelli 2006). moreover, another method that imposes internal benchmarks to measure operational performance is total factor productivity (tfp), which is used to measure the changes in aggregate output per unit of aggregate input (thirtle & bottomley 1992). nevertheless, these ratios and methods have various features that make it challenging to aggregate them to provide an understanding of the overall operational performance (parkan 1996). only time series data, unit prices and quantities can be used in the model to obtain performance measurements with tfp (parkan 1996). operational performance can be used for comparative analyses, as explained by ghalayini and noble (1996). according to parkan (1996), examining the operational performance of a firm overtime introduces managers to aspects of comparison between operations across time and thus competition. therefore, in each ppa, ecotourism operations that consume fewer resources and generate higher revenues in a specific year would have performed better or more competitively relative to other years. this means that operational performance measures the relative competitiveness of ecotourism operations over a period (parkan 1996). therefore, his study aims to examine the operational competitiveness of ecotourism operations in each ppa managed by ekznw. this is performed using a non-parametric method called operational competitiveness rating analysis (ocra). this method was selected because, according to the authors’ knowledge, it is the first of its kind in south africa to use the ocra procedure to measure and assess the operational competitiveness of ppas and it has an advantage over other methods. the advantage of the ocra model is its capacity to show the period during which ekznw’s overall operational performance has been inadequate or more than adequate compared to other methods as well as the sources of those shortcomings and strengths (parkan 1996). this is possible because of the way operational competitiveness ratings (revenue generation and resource consumption inefficiency ratings) are computed and its ability to incorporate management’s perceptions of the relative importance of the cost and revenue categories. the article begins with a brief account of the competitiveness of ppas, followed by the study area and data description. then the ocra method, the empirical model, is explained. thereafter, the operational competitiveness profiles of each ppa are presented. this is followed by the presentation of the results and discussion on the operational competitiveness of ecotourism operations. finally, conclusions and strategic implications are drawn. ezemvelo kzn wildlife and public protected areas the majority of ppas are managed by central governing bodies or conservation agencies (eagles 2002; porter et al. 2003). these conservation agencies collect revenues from ppas and allocate operating budgets (eagles 2002). however, budget allocations are not closely linked with ecotourism use levels (eagles 2002) and the environmental value of the area, which reduces the incentives of ppas to manage their operations sustainably and profitably. this is true of ekznw because part of its budget comes from government and the the rest from own revenue generated. according to ekznw management authorities, the current budget of ekznw is around r890 million, of which 25% is generated from its own operations and the rest is a government subsidy. ekznw retains all its revenues and submits a budget request to the kzn department of economic development, tourism and environmental affairs. the department often allocates its financial resources on a 3-year cycle to different sectors (including ekznw) in the province, based on the availability of its financial resources and priorities. furthermore, when there is a budget deficit, ekznw will negotiate with the department for more funding, seek other funding sources or attempt to increase its internal sources of revenue. however, if these options cannot make up the budget shortfall, ekznw will typically reduce its budget for lower priority areas. still, increasing budget cuts from government have affected the capacity of ekznw to cover its costs. ekznw manages several protected areas of which some have ecotourism features that generate sufficient income whilst others have biodiversity value that lack income-generating features. therefore, income from profitable protected areas with ecotourism features is used to cross-subsidise those with pure public good features in terms of biodiversity conservation. this has made it necessary for ekznw to prioritise projects in its ppas according to their ability to generate revenue. there have been frequent demands by the south african government for ekznw to design and implement a strategy that is aligned with current trends on sustainable funding for protected area management, in which there is a need to balance between biodiversity conservation objectives and revenue generation (dube 2011). the reasons for this request range from poor corporate governance, recurrent financial mismanagement and pressing socio-economic development needs (dube 2011; ridl 2012). ekznw is considering new business models aimed at achieving business efficiency by optimising the use of financial resources and increasing its resource base (ekznw 2009). furthermore, ekznw plans to focus on more effective marketing strategies to increase revenues (ekznw 2009). according to dube (2011), for ekznw to reduce its dependence on government finance, it needs to focus on three areas, namely, payment for ecosystems services, public–private partnerships and co-management with the private sector and communities. moreover, several ppas in kzn and most of south africa typically generate insufficient revenues to finance operations and cover costs, and as such, most are managed at a loss (dube 2011; myburgh & saayman 1999). according to officials at ekznw, ppas are not mandated to aim for profit even though it is desirable to at least cover their full costs. typical revenue sources for most ppas, including the ekznw ppas, include accommodation, wildlife product sales, admission fees, rentals and concessions, and wildlife viewing (eagles 2002; parker & khare 2005; porter et al. 2003). according to dixon and sherman (1991), there are three types of costs in maintaining protected areas: direct costs (recurrent costs of maintaining and managing a protected area), indirect costs (damages caused by wildlife) and opportunity costs (foregone losses resulting from protecting such areas). the benefits and costs of ecotourism interact in complex ways, but it is imperative that ppas maximise their benefits whilst minimising costs (eagles et al. 2002). several ppas in kzn and most of south africa generate insufficient revenues to finance operations and cover their running costs (dube 2011; myburgh & saayman 1999). there have been studies conducted on ppas in kzn that have touched on the theme of performance or competitiveness in ecotourism (e.g. flanagan 2014; porter et al. 2003). internationally, for instance, a report by bovarnick et al. (2010) was compiled for the united nations development programme (undp) to analyse the financial status and sustainability of protected areas in several latin american and caribbean countries. this report found that protected areas in these regions are underfunded and that funding needs are likely to increase considering the implications of climate change. moreover, rylance (2017) conducted a study to assess the revenue generation of tourism in 93 mozambican protected areas. the findings of the study were that the total annual revenue generation from protected areas was $24m from tourism-related activities, contributing around 10% to the tourism sector. accordingly, identifying and reporting all possible revenue flows will assist in justifying public support for protected areas. however, the literature on this subject in south africa is very limited, particularly on operations cost estimation and assessment. therefore, to contribute to this body of knowledge, this article will examine the allocation of resources or funds by ekznw to ecotourism operations in ppas and the financial performance or competitiveness of these operations relative to each other over time. study area the kzn province covers an area of 92 285 km2, and is situated on the eastern coast of south africa in a biologically rich transition zone between tropical biota in the north and subtropical biota in the south (eeley, lawes & piper 1999; goodman 2003). the abundance of kzn’s biodiversity comes from its altitudinal gradient and its varied geology, topography and climate (eeley et al. 1999; goodman 2003). ekznw is a government agency under the kzn department of agriculture, environmental affairs and rural development and manages biodiversity conservation in the kzn province (goodman 2003). ekznw was created in 1997 via a merger between the natal parks board and the kwazulu department of nature conservation (goodman 2003). it manages 110 ppas that cover over 675 000 hectares (aylward & lutz 2003). moreover, it receives financial support from government through the kzn provincial treasury (dube 2011; goodman 2003). furthermore, the agency receives additional funding from non-governmental organisations (ngos) and philanthropic organisations (dube 2011). protected areas considered in the study are shown in figure 1. figure 1: ezemvelo kzn wildlife public protected areas. the data to evaluate the trends in the competitiveness of commercial operations in ppas, financial data were collected from ekznw for 2007–2013. originally, the ppas required were to be selected using the stratified random sampling strategy across all ekznw administrative regions (ukhahlamba, zululand and coastal regions), where between 35 and 50 ppas mainly focused on ecotourism operations would have been selected. however, ekznw was only able to provide 32 randomly selected ppas from the 110 ppas, some with a stronger conservation focus than a commercial operation, citing organisational privacy concerns. ekznw provided ppas selected randomly across ukhahlamba, zululand and coastal regions with 14, 7 and 11 ppas, respectively. therefore, there is a reduced possibility of sample selection bias in the study. the financial data provided consisted of annual cost and revenue values of commercial operations. cost and revenue values for each protected area were in nominal terms. hence, the south african consumer price index was used to deflate the cost and revenue values for 2007–2013, taking 2005 as the base year. cost and revenue items for each protected area were disaggregated and measured separately. in this study, cost categories are described as resources consumed and revenue categories as revenues generated. thus, 11 cost categories and seven revenue categories were analysed per year, 2007–2013 (table 1). table 1: categories for resource consumption and revenue generated for commercial operations in each public protected area. empirical model: the operational competitiveness rating analysis procedure operational competitiveness rating analysis is a relative performance method used to measure the performance of operating entities called production units (pus). pus are purposeful entities that convert resources into goods and services (parkan 1991). the ocra procedure involves simple ratio-type, non-iterative computations that measure the pus relative to operational competitiveness (parkan et al. 1997). it is suitable for time series data: financial value, on quantities and unit prices, and has been used in different industries to measure operational competitiveness (parkan 1996, 1999, 2003; parkan et al. 1997; parkan & wu 1999). following parkan (1996), the model can further be described as follows. in the model, an operating entity is represented by a pu in each year. a comparison is conducted of operational performance of k pus that consume resources in c categories and generate revenues in the r categories. to compute cost and revenue inefficiency ratings, the prices and quantities of inputs and outputs could be used to obtain information about k pus’ relative input or output efficiency. nonetheless, resource cost and revenue values can be used to obtain relative cost and revenue inefficiency ratings, respectively (see parkan 1996 for details on derivation). in this study, because data on only cost and revenue values were used, the kth pu’s cost and revenue vectors can be represented as vectors , respectively, with as the cost incurred for the ith resource and as the revenue generated from the jth output. cost and revenue at puk are denoted as and , where ckm = costkm = 1ukm is the cost of the mth resource category, m = 1,……, m, and rkh = revenuekh = 1vkh is the revenue generated from the hth category of outputs, h = 1, …, h, at puk, k = 1,…., k. according to parkan and wu (1999), the relative importance of a pu’s performance in a cost or revenue category is dependent on that category’s impact on the overall performance of the pu. this relative importance is reflected by calibration constants, denoted as akm and bkh for resource consumption and revenue generation, respectively, for puk. the calibration constants can be computed by: the first equation defines am as the average cost share of the mth cost category, and the second equation describes bh as the average revenue share of the hth revenue category (for details, see parkan 1996). the resource consumption inefficiency model is meant to determine whether input quantities would give information about a pu’s relative input inefficiency. the kth pu’s resource inefficiency rating, ckm, is computed with respect to the mth input category and is expressed as: where the is the lowest cost incurred by a pu amid k pus with respect to the mth cost category. then, the sum is linearly scaled by: which is the kth pu’s relative resource inefficiency rating. the least inefficient pu will receive an inefficiency rating of zero. for details of the derivation, see parkan (1999). the same approach can be adopted to determine revenue generation inefficiency ratings for related pus. the first step is to compute the relative inefficiency rating of the kth pu in relation to the hth revenue category by: where is the highest realised revenue in relation to the hth revenue category. the scaled sum was calculated (see parkan 1999) and the equation is: which is the kth pu’s relative revenue generation inefficiency rating. the assessment is such that the least inefficient rating has a lower value relative to other pus in the hth revenue category. the computation of the kth pu’s combined inefficiency rating, ek, as the scaled sum of its combined resource consumption and revenue generation inefficiency ratings is as follows: according to parkan (1996), the rating ek measures the overall performance of kth pu relative to other pus. the best performing pu or least inefficient rating receives a rating of zero and larger ek values reflect inferior performance or higher inefficiency. for details on the derivation, see parkan (1999). the results and discussion combined inefficiency ratings for public protected areas the results in table 2 show that the combined inefficiency ratings for kamberg were at their lowest in 2007, improved in 2008 and then increased in 2009. from 2010 to 2013, they fluctuated several times, improving again in 2011. the results suggest that disparities in operational competitiveness can be accounted for by the high share of total revenues and costs that largely influenced the performance of revenue and resource competitiveness, which in turn affected operational competitiveness. a closer examination reveals that the fluctuations of combined inefficiency ratings coincide with observed fluctuations in resource consumption inefficiency ratings, implying that the performance of resource competitiveness had more influence on the operational competitiveness of kamberg. table 2: combined inefficiency ratings (ek) computed for each public protected area. in midmar, efficiency improvements coincided with resource competitiveness improvements in 2008 and 2010 (tables 2 and 3), whereas with revenue competitiveness it was from 2010 to 2012. it seems that midmar was able to improve operational competitiveness when the revenue competitiveness was low and resource competitiveness was high in 2008 and vice versa in 2012. for ntshondwe, the combined inefficiency ratings were closely linked to the performance of resource consumption inefficiency ratings, implying that these had a greater impact on combined inefficiency ratings and thus operational competitiveness. at mpila, meanwhile, the years that registered improvements in resource consumption inefficiency ratings, 2007–2009, coincided with the years that registered improvements in the combined inefficiency ratings. furthermore, it was only in the year 2011 that mpila registered the lowest relative combined inefficiency rating (table 2), the same year resource consumption and revenue generation inefficiency ratings performed better. first, this implies that resource competitiveness had more influence on operational competitiveness because it performed better than revenue competitiveness in some years. second, the results also imply that a combination of better performing revenue and resource competitiveness had a greater effect in improving operational competitiveness. the results seem to support the findings by shieh (2012) that better cost efficiency leads to improved financial performance and by tsaur (2001) that both low inefficiencies in revenue and costs can enhance operational performance. at the phongolo controlled hunting area (pcha), revenue competitiveness influenced operational competitiveness more than resource competitiveness because revenue generation inefficiency ratings were more than 10 times the size of resource consumption inefficiency ratings (table 3). this suggests that, in this study, high inefficiencies in revenue generation have resulted in poor operational competitiveness. table 3: resource consumption inefficiency ratings (cu) for each public protected area. for rugged glen stables (rgs), table 3 shows that resource consumption inefficiency ratings were higher than revenue generation inefficiency ratings between 2007 and 2010. these coincide with the years that operational competitiveness was worse, whereas revenue generation inefficiency ratings were lower in the same period. moreover, between 2011 and 2012, resource competitiveness improved whilst revenue competitiveness declined, again coinciding with better operational competitiveness in this period. therefore, in this study, it suggests that resource competitiveness had a greater effect than revenue competitiveness on operational competitiveness. resource consumption and revenue generation inefficiency ratings for public protected areas resource consumption and revenue inefficiency ratings reflect the resource and revenue competitiveness of each pu. examples of resource consumption and revenue generation inefficiency ratings corresponding to each ppa are illustrated graphically to obtain a better sense of relative competitiveness of resources consumed and revenues generated by each ppa between 2007 and 2013 (figure 2). figure 2: graphical illustration of resource consumption and revenue generation inefficiency ratings for each public protected area (examples). (a) kamberg, (b) midmar, (c) ntshondwe, (d) rugged glen stables, (e) phongolo controlled hunting area and (f) mpila. according to the results of table 4 and figure 2, for kamberg in 2007, the most competitive year was 2007. revenue became increasingly less competitive from 2008 to 2013. mostly, the average shares of total revenues come from the r1, r2 and r3 categories. this suggests that from 2008 to 2013, revenue generators did not produce significant returns to offset the increasing inefficiency. there were sharp declines and increases in resource competitiveness between 2008 and 2011, and then a gradual decrease in competitiveness between 2012 and 2013. from table 5, staff costs take the largest share of costs incurred by kamberg, followed by utilities. this result suggests that increased cost control in staff and utility costs could improve kamberg’s resource competitiveness profile. table 4: revenue generation inefficiency ratings (rv) computed for each public protected area. table 5: resource consumption (input) calibration constants (am) for each public protected area the results presented in table 3 (also see figure 2) show that midmar’s resource competitiveness declined from 2007 to 2008. however, the inefficiency ratings increased in 2009 and declined in 2010. there was a gradual increase in the value of the ratings from 2011 to 2013. this suggests that categories with a higher average share of total costs (c1, c3, c5 and c9) had minimal impact on the improvement of resource competitiveness because of a lack of cost control, and improvements in this area will occur only if midmar adopts cost-cutting strategies, without compromising on the quality of its services. revenue generation inefficiency ratings increased slightly from 2007 to 2009, then declined dramatically from 2010 to 2012 and increased again in 2013. the revenue generation inefficiency ratings were still unacceptably high. the main revenue generators came from the r1 and r2 categories, and these did little to improve revenue competitiveness. this has the important implication that increasing the revenue share of accommodation and entrance fees can impact revenue competitiveness positively. according to dube (2011), ekznw offers outdated products that few clients can relate to and is stuck in traditional conservation practices that prevent it from taking advantage of new markets. it must modernise its services and products, maintaining natural capital and traditions of heritage value. revenue generation inefficiency ratings in ntshondwe gradually increased from 2007 to 2013, suggesting that revenue competitiveness worsened (figure 2). although a large share of revenue was derived from accommodation and sales, this share was insufficient to improve revenue competitiveness. moreover, ntshondwe offers trails, rides and tours, but the share of revenue from these activities was insufficient to make meaningful improvements on revenue competitiveness. noticeable inefficiencies for resource consumption were in 2009 and 2011. an improvement in the revenue and resource competitiveness profile of ntshondwe will occur if management diversifies ecotourism activities and reduces costs. cost-effective marketing strategies can be adopted to increase demand, which can lead to lower inefficiency (barros 2005). for mpila, the best performing revenue generation inefficiency ratings lie in the r2 and r7 categories, implying that management can redirect efforts to improve revenue competitiveness by focusing on these categories. as for resource consumption inefficiency ratings, there was a notable decline from 2007 to 2009 (table 3 and figure 2), and a slight increase in 2010, followed by a dramatic decrease in resource consumption inefficiency ratings in the remaining years. this improvement stems from a decrease in costs in the c1, c2, c5, c9 and c11 categories in that period. furthermore, the results indicate that if management shifted resources away from the c5, c9 and c11 categories, this could improve resource competitiveness. in pcha, revenue generation inefficiency ratings increased from 2007 to 2008, suggesting that revenue competitiveness declined in this period. the least inefficient year was 2009, and then revenue generation inefficiency ratings increased in 2010, and then decreased in 2012, increasing again in 2013 (figure 2). in 2008, resource competitiveness was stable, but resource consumption inefficiency ratings increased in 2009 and 2010. a slight upward trend continued from 2011 to 2013, which may have been because of weak cost control. the c1, c5 and c7 categories had more influence on resource competitiveness because of higher average shares of total costs (table 5). the main revenue generators in pcha were hunting and accommodation. the result is not surprising because many studies (absa 2003; baker 1997; damm 2005; mossman & mossman 1976; van der merwe, saayman & krugell 2004) have shown that hunting is the most lucrative ecotourism enterprise and as such, should have a greater effect on increasing revenue competitiveness. in rgs, revenue generation inefficiency ratings increased from 2007 to 2008 and then remained steady until 2013 (figure 2). the r1, r4 and r7 categories had larger shares of total revenue. on the contrary, resource competitiveness worsened between 2011 and 2013. activities such as services and utilities took a considerable share of costs, next to personnel-related expenses. resource consumption calibration constants for public protected areas the evaluation of resource consumption calibration constants indicates, for instance, that in kamberg, for category c1 (basic salary, unemployment insurance fund [uif], housing subsidy, pension, medical aid and service bonus) had the highest average share of total costs at 4.8%. moreover, category c2 (overtime, subsistence, danger, night shift and standby allowance) had the least average share of total costs at 0.18%. this suggests that kamberg allocates more funds to paying permanent staff members. the category with the highest average share of total costs for midmar was the c1 category at 3.5%, followed by the c3 (short-term contractual work) category because its average share of total cost was 0.64%. the use of temporary staff or contractual workers has several benefits such as a reduction in recruitment costs and employee costs (david, brendan & mike 2006). therefore, midmar should use contractual workers for short-term tasks and assignments to control staffing costs. the c9 (gas, water and electricity) and c5 (roads, tools and equipment, furniture and fittings, buildings and structures) categories each had an average share of total costs of 0.55% and 0.53%, respectively. high utility costs can reduce the revenue maximisation of any organisation significantly (hassanien & dale 2013). midmar has campsites with electric plug points and chalets fitted with refrigerators, electric stoves and cable tv (dstv), and with running cold and hot water (ekznw 2014). furthermore, midmar hosts sporting and music events during weekends and peak periods and has multiple recreational activities which include boating, swimming, water skiing, picnicking and fishing (ekznw 2014). according to flanagan (2014), the provision of electricity or plug-in points at ekznw has a positive effect on the desirability of a ppa site as a tourist destination. thus, ekznw could introduce cheaper and greener alternatives such as solar panels to reduce the cost of utilities. moreover, the c6 (security expenses, security bank expenses and fire extinguisher services) category had an average share of total costs of 0.41%. spending on security should be informed by assessed risks and threats. considering that crime in south africa, especially in kwazulu-natal, has been increasing consistently in all crime categories (saps 2016) and that the dam is adjacent to mpophomeni township, midmar management is advised to increase funds allocated for security to ensure the safety of tourists and integrity of its conservation area. townships are places of socio-economic hardship and this often compels some people to turn to criminal activities. for ntshondwe, the c1 category had the highest calibration constant and thus the highest average share of total costs at 4.1%. this result is consistent with several studies that have shown that salaries and wages are the largest costs for tourism operations (e.g. bovarnick et al. 2010; eagles 2002). labour costs should be assessed against the marginal value product of each labour category, and it will remain essential to align the qualities of labour with their respective costs and revenue contributions. for instance, management can review salaries and wages, reduce weekday hours, remunerate employees based on their level of expertise or contribution to the organisation, reward exceptional performance and eliminate redundant positions that add no significant value to ntshondwe and ekznw. the c11 (cost of sales fuel, curios and other) category had the second largest average share of total costs at 0.9%. cost of sales refers to the purchase price of goods sold at ntshondwe such as fuel and curios. because this cost is high, ekznw can revise its business model around, for instance, outsourcing fuel sales to fuel companies or curio sales to local craft shops. the c9, c7 and c5 categories stand at 0.53%, 0.52% and 0.48%, respectively. one way in which management can reduce the high cost of utilities (c9) is by ‘going green’ (pfister & tierney 2008). this approach provides ntshondwe and other ppas with the opportunity to improve long-term profitability by lowering expenses and enshrining environmental stewardship, adding value to the ekznw brand image of ecological responsibility. ntshondwe and most ppas can install solar panels in their accommodation facilities and in the camping area where electric plug points are available. the upfront cost of implementing these eco-friendly alternatives is high, but in most cases the returns on investment in the long term are excellent, largely because of recurrent cost savings (rahman, reynolds & svaren 2012). moreover, management should also review its maintenance strategies. according to plantweb (2003), the most basic and common maintenance strategy is the reactive and needs-based maintenance (‘fix it when it breaks’ strategy), but repair costs for infrastructure and equipment are higher than in most strategies. therefore, management at ntshondwe could utilise preventive, predictive and proactive maintenance strategies interchangeably, depending on the scale of the maintenance required. in mpila, the relative importance of the c11 category comes second to the c1 category. still, the c10, c5 and c9 categories had an average share of total cost of 0.7%, 0.5% and 0.47%, respectively. it appears that mpila incurred higher costs from its bar and restaurant sales. mpila can reduce the relatively high average share of total costs in the c11 category by offering concessions to private enterprises which are motivated by profits and may have the management skills and operating procedures required to run such operations efficiently (guasch 2004). the c10 and c5 categories had higher shares; however, these costs are to some extent necessary because it is another way mpila can offer quality ecotourism products to retain its customers. according to beerli and martin (2004), cited in mmopelwa, kgathi and molefhe (2007), the image of a tourist destination is essential in influencing the satisfaction of visitors. moreover, tourists are willing to pay higher fees if services are improved and funds are invested in environmental conservation (mmopelwa et al. 2007), especially wealthy consumers. therefore, management needs to ensure that adequate funds are available to conduct maintenance operations on the infrastructure and necessary equipment. utilities were mostly driven up by high electricity accounts. the best alternative for mpila to reduce utility costs in the long run is to invest in sustainable and energy efficient technologies such as solar power. the results presented in table 5 show that at pcha, the c3, c10 and c7 categories had average shares of total costs at 0.8%, 0.76% and 0.42%, respectively. much of the costs are centred on contractual work which at pcha includes live animal capture using helicopters and bomas, an expensive task routinely conducted by professionals (hudson, drew & baskin 1989). moreover, hudson et al. (1989) state that fencing is a major development cost with different animals requiring fences with different heights, numbers of strands and types of poles. furthermore, table 5 shows that rgs, sodwana bay resort and kosi bay had average shares of total costs of 6.5%, 2.44% and 5.2%, respectively, in the c1 category. the high costs in the c1 category seems to be a common phenomenon across ekznw ppas. in rgs, the c6 and c9 categories had the second and third highest average cost shares of 0.95% and 0.34%, respectively. the high cost of stable management services in the c6 category, relative to other categories, is mainly because of costly horse and stable management activities (knight 2010). revenue generation calibration constants for public protected areas according to table 6, kamberg’s main revenue generators between 2007 and 2013 were ‘accommodation’ (r1-chalets, rustic cottages and rondavels), ‘admissions’ (r2-entrance fees) and ‘sales’ (r5-sales revenue from operating retail stores). the revenue calibration constant or average share of total revenue for the ‘trails, tours and rides’ (r7) was low at 0.016%. in addition, giants castle, midmar and ntshondwe also derived a large share of their total revenues from the r1 category with average revenue shares of 5.5%, 5.1% and 4.2%, respectively. furthermore, the r5 category had the second highest relative importance to giants castle and ntshondwe with average shares of total revenues of 1.3% and 1.2%, respectively. meanwhile, midmar’s second highest revenue generator came from the r2 category, and this category had 1.2% of average shares of total revenues. in midmar, the main activities were boating, yachting and fishing (ekznw 2014). therefore, it was rather surprising to find that the r3 category had the second lowest average share of total revenue at 0.047%. giants castle and ntshondwe received significant revenues from the r7 category with average shares of total revenue at 0.34% and 0.47%, respectively. table 6: revenue generation calibration constants (bh) for each public protected area. kamberg offers tours to san rock art, and is regarded as the highlight of visiting the area (ekznw 2014). however, the results above show that this activity generated very low revenue despite it being one of the main attraction at kamberg. similarly, the r7 category in giants castle and ntshondwe generated lower revenues relative to other revenue-generating categories. these results are consistent with the findings of flanagan (2014), who suggested that cultural or tour areas did not have any significant impact on revenues received. moreover, in midmar, strict regulations around freshwater recreational use could be causing the low revenues because boat skippers are required to adhere to strict zoning areas (ekznw 2014). furthermore, the results suggest that accommodation is the main revenue generator for kamberg, giants castle, midmar and ntshondwe. flanagan (2014) also found that increases in occupation rates in ekznw accommodation have significant impacts on revenue, especially when associated with popular tourist activities. the results further indicate that mpila derived a considerable proportion of its average shares of total revenues from the r1 category at 4.7%. this result is not surprising as accommodation has been found to be one of the main revenue generators for ppas (eagles 2002; flanagan 2014; porter et al. 2003). accommodation at mpila consists of 20 chalets and 15 tented camps. according to flanagan (2014), chalets on average experience higher visitor occupation than other accommodation types offered by ekznw. therefore, ppas with fewer chalets (like mpila and hilltop) should have been generating less revenue. furthermore, mpila, found in the hluhluwe-imfolozi park, with an array of wildlife including the ‘big five’, derives low revenue from the r7 category with an average share of total revenue of 0.43%, which rather is surprising. this result, however, is consistent with flanagan (2014), who has shown that the ‘big five’ animals had a statistically insignificant effect on revenues in ppas. it could be that instead of paying for guided tours, visitors prefer to pay entrance fees to such areas and drive themselves to view wildlife. this also explains the 1.7% average share of total revenue in the r2 category, higher than the r7 category. pcha had two categories (r1 and r3) generating revenues. the r3 category for pcha had the largest average share of total revenues at 11.7%, making this category the best revenue generator in the commercial operations of ekznw across all categories considered. permits and licences in pcha are usually granted for hunting and fishing in the phongola river and the phongolopoort dam (ekznw 2014). this indicates that hunting and fishing contribute significantly to revenues in pcha. the result is consistent with many studies in the past (e.g. absa 2003; damm 2005; van der merwe et al. 2004), which show that hunting is a highly profitable operation in the nature-based tourism industry. thus, ekznw management could be forced to enact stricter permit and licencing regulations to control the quantity of wildlife being harvested. in rgs, the category with the highest average share of total revenues was r7 at 4.9%. according to ekznw (2014), rgs mainly conducts horse riding operations, the main revenue generator. this operation mainly attracts europeans ollenburg (2005). considering the growing african middle class in south africa (van loggerenberg & herbst 2010), rgs needs to tap into this lucrative market to grow its revenue. according to cini and saayman (2014), younger people at national parks have been shown to have a higher propensity to spend money at national parks. therefore, ekznw can also market to these younger groups. conclusion and management implications the aim of this study was to measure the operational competitiveness of ecotourism operations of ppas in ekznw. this was performed through constructing an operational competitiveness profile for each ppa, using the ocra procedure. financial data for ecotourism operations in ppas were collected from ekznw for 2007–2013. the study showed that hunting generated more revenue than most of ekznw’s ecotourism operations and had the greatest effect on operational competitiveness. this implies that for management in ekznw to improve its revenue competitiveness, it should implement strategies to scale up hunting-operations. however, it will only be possible to implement this strategy if ekznw increases the number of hunting-sanctioned protected areas because currently only five have that status. this result has other implications as well, one of which is creating perverse incentives that could encourage ekznw to move away from ‘deep ecotourism’, which is in line with conservation objectives, to ‘shallow ecotourism’, which is driven by local community development and beneficiation other than just conservation without the human dimension. ecotourism activities such as trails, rides and tours generally had the lowest average share of total revenue. the implication is that management either needs to make guided tours more attractive or increase park fees to generate more income. the implementation of the latter will depend on the demand elasticities of park visits at ekznw. the results show that ppas in ekznw place a higher importance on salaries and wages, maintenance and repairs, and utilities. therefore, improved cost control is required in these resource consumption activities that adversely affect operational competitiveness. for instance, management needs to explore different methods of minimising labour costs by reducing the number of staff or the level of compensation. moreover, considering that biodiversity management is a labour-intensive activity and relies heavily on human capital, ekznw can ensure that personnel are remunerated in accordance with skill sets that add value to ppas. however, minimising the labour force might reduce incentives for staff to work effectively and lower morale, which in turn might reduce the quality of services provided and thus the capacity of ppas to generate revenue. ekznw should revise its utility expenses (especially electricity) and find alternative sustainable energy sources to reduce not only the utility bill but also its environmental footprint. the capital outlay for such a project in the short term is expected to be high. thus, it is recommended for ekznw to pursue this incrementally. also, the agency should continue allocating funds for maintenance and repairs to improve tourist flows in the long term. it is also recommended that ekznw implement policies and strategies to increase its share of revenue from the main revenue-generating operations (accommodation, permits, licences and admissions) to improve revenue competitiveness. moreover, ekznw needs to appeal to new markets based on demographic changes (the african middle class) as a strategy to increase revenues. by constantly reviewing its policies and strategies, ekznw will be better positioned to take advantage of such changes. furthermore, this result implies that by upgrading accommodation facilities and its product offering, ekznw could provide better services which would increase its revenue and ultimately improve its operational competitiveness. however, this will require major investment from the government, which will prove difficult given the current economic climate and will also depend on whether the supply of facilities in a ppa will match visitor demand. nonetheless, ekznw can initiate partnerships with the private sector and environmental ngos. again, caution should be taken in increasing tourist flows to these areas as this could reduce the quality, and aesthetic and amenity value of the environment. to create lower inefficiencies in overall operations, ppas under ekznw should employ various strategies to increase revenues or reduce costs. each ppa allocates resources and generates revenues differently. therefore, flexibility in the centralised resource allocation system is required so that ppas that perform better and manage funds properly can retain some fraction of their earnings to reward staff and pay for maintenance, repairs, utilities and new projects. however, this will only be possible if ekznw raises the standard of its facilities in some of its (ecotourism-based) ppas to appeal more to a broader market. ppas with high costs, low revenues and poor operational competitiveness but high ecological or environmental value should be monitored closely, with stricter control by ekznw so that they can develop strategies to diversify revenues and improve cost management. in other words, ekznw should generate better incentives for ecotourism in ppas to increase competitiveness and for nature to contribute to its conservation. revenue and resource competitiveness varied erratically throughout the study period, with minor improvements in some years and a worsening competitiveness in most of the years. therefore, ekznw needs to ensure that both cost reduction and high revenue generation are a priority at any one time to improve operational competitiveness. however, the combination of these strategies will vary among ppas in ekznw and will depend on the nature and degree of operations in the ppa concerned. moreover, increasing revenues through increasing service charges will require information on the price elasticity of demand for the services of ppas under investigation. this information will assist in devising a fee system that could be adopted to maximise revenues and improve the economic efficiency of ppas. for instance, rural communities that surround these ppas generally have a higher price elasticity of demand than wealthier international visitors, making them more sensitive to fee increases. therefore, ekznw can adopt a multi-layered fee system (price discrimination), drawing lessons from other countries so as to make their services sensitive to income inequalities among local visitors whilst generating adequate revenues from international visitors. this is certainly an area of study that needs further investigation. acknowledgements the authors wish to thank mr rahman devduth, the natural resources trade coordinator, and mr ebrahim mulla, the finance manager, both at ezemvelo kzn wildlife, for providing the data and additional information. insightful comments from two anonymous reviewers of the journal are duly acknowledged. any remaining errors or omissions are the responsibilities of the authors. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions k.d.m. was the main author responsible for data collection and analysis. e.w. was responsible for supervising the study, contributing to the write-up in terms of regular inputs and comments for improvement at various stages of the study, identifying relevant literature, reviewing the article, revising the article following reviewers’ comments and responding to reviewers’ comments. references absa, 2003, game ranch profitability in south africa, absa group economic research, rivonia, south africa. aylward, b.a. & lutz, e., 2003, nature tourism, conservation, and development in kwazulu-natal, south africa, world bank, washington, dc. baker, j.e., 1997, ‘trophy hunting as a sustainable use of wildlife resources in southern and eastern africa’, journal of sustainable tourism 5, 306–321. https://doi.org/10.1080/09669589708667294 barnes, j.i. & de jager, j., 1996, ‘economic and financial incentives for wildlife use on private land in namibia and the implications for policy’, south african journal of wildlife research 26, 37–46. barros, c.p., 2005, ‘measuring efficiency in the hotel sector’, annals of tourism research 32, 456–477. https://doi.org/10.1016/j.annals.2004.07.011 beerli, a. & martin, j.d., 2004, ‘tourists’ characteristics and the perceived image of tourist destinations: a quantitative analysis – a case study of lanzarote, spain’, tourism management 25, 623–636. https://doi.org/10.1016/j.tourman.2003.06.004 bosetti, v. & locatelli, g., 2006, ‘a data envelopment analysis approach to the assessment of natural parks’ economic efficiency and sustainability: the case of italian national parks’, sustainable development 14(4), 277–286. https://doi.org/10.1002/sd.288 bovarnick, a., fernández-baca, j., galindo, j. & negret, h., 2010, financial sustainability of protected areas in latin america and the caribbean: investment policy guidance, united nations development programme and the nature conservancy. bruner, a.g., gullison, r.e. & balmford, a., 2004, ‘financial costs and shortfalls of managing and expanding protected area systems in developing countries’, bioscience 54, 12. https://doi.org/10.1641/0006-3568(2004)054%5b1119:fcasom%5d2.0.co;2 charnes, a.c., cooper, w.w. & rhodes, e., 1978, ‘measuring the efficiency of decision making units’, european journal of operational research 2, 429–444. https://doi.org/10.1016/0377-2217(78)90138-8 child, b.a., musengezi, j., parent, g.d. & child, g.f., 2012, ‘the economics and institutional economics of wildlife on private land in africa’, pastoralism 2, 1–32. https://doi.org/10.1186/2041-7136-2-18 cini, f. & saayman, m., 2014, ‘which age group spends the most in a national park?’, koedoe 56(2), art. #1158, 8 pages. https://doi.org/10.4102/koedoe.v56i2.1158 damania, r. & hatch, j., 2005, ‘protecting eden: markets or government?’, ecological economics 53, 339–351. https://doi.org/10.1016/j.ecolecon.2004.09.015 damm, g., 2005, ‘hunting in south africa: facts, risks and opportunities’, african indaba 3, 1–23. david, b., brendan, b. & mike, m., 2006, ‘the changing world of the temporary worker: the potential hr impact of legislation’, personnel review 35, 191–206. https://doi.org/10.1108/00483480610645821 dixon, j.a. & sherman, p.b., 1990, economics of protected areas: a new look at benefits and costs, island press, washington, dc. dixon, j.a. & sherman, p.b., 1991, ‘economics of protected areas’, ambio 20, 68–74. dube, t.b., 2011, ‘an analysis of alternative funding strategies for protected area management: a case study of ezemvelo kzn wildlife’, master of development finance thesis, stellenbosch university. eagles, p.f., 2002, ‘trends in park tourism: economics, finance and management’, journal of sustainable tourism 10, 132–153. https://doi.org/10.1080/09669580208667158 eagles, p.f., 2003, ‘international trends in park tourism: the emerging role of finance’, the george wright forum 20(1), 25–57. eagles, p.f., mccool, s.f., haynes, c.d. & phillips, a., 2002, sustainable tourism in protected areas: guidelines for planning and management, vol. 8, iucn, gland, switzerland. eeley, h.a., lawes, m.j. & piper, s.e., 1999, ‘the influence of climate change on the distribution of indigenous forest in kwazulu-natal, south africa’, journal of biogeography 26, 595–617. https://doi.org/10.1046/j.1365-2699.1999.00307.x ekznw, 2009, ezemvelo kwazulu-natal wildlife five year strategic and performance plan for 2009 and 2014, viewed 05 november 2013, from http://www.kznwildlife.com ekznw, 2014, ezemvelo kzn widlife profile, viewed 18 december 2014, from http://www.kznwildlife.com emerton, l., bishop, j. & thomas, l., 2006, sustainable financing of protected areas: a global review of challenges and options, iucn, gland, switzerland. flanagan, o.j., 2014, ‘spatial prioritisation of conservation areas on the fringes of kwazulu-natal protected areas: application of the characteristics framework using tourism competitiveness’, master of science in agriculture (agribusiness) thesis, university of kwazulu-natal. ghalayini, a.m. & noble, j.s., 1996, ‘the changing basis of performance measurement’, international journal of operations and production management 16, 63–80. https://doi.org/10.1108/01443579610125787 goodman, p.s., 2003, ‘assessing management effectiveness and setting priorities in protected areas in kwazulu-natal’, bioscience 53, 843–850. https://doi.org/10.1641/0006-3568(2003)053%5b0843:ameasp%5d2.0.co;2 guasch, l., 2004, ‘granting and renegotiating infrastructure concessions: doing it right’, the world bank, washington, dc. harris, p.a., 1999, ‘review of equine feeding and stable management practices in the uk concentrating on the last decade of the 20th century’, equine veterinary journal 31, 46–54. https://doi.org/10.1111/j.2042-3306.1999.tb05156.x hassanien, a. & dale, c. (eds.), 2013, ‘tourism, hospitality and events facilities’, in facilities management and development for tourism, hospitality and events, pp. 20–36, cabi, edinburgh. holden, a., 2013, tourism, poverty and development, routledge, london. honey, m., 2003, ‘protecting eden: setting green standards for the tourism industry’, environment: science and policy for sustainable development 45(6), 8. https://doi.org/10.1080/00139157.2003.10544694 hudson, r.j., drew, k. & baskin, l.m., 1989, wildlife production systems: economic utilisation of wild ungulates, cup, cambridge, uk. archive. james, a., gaston, k.j. & balmford, a., 2001, ‘can we afford to conserve biodiversity?’, bioscience 51, 1. https://doi.org/10.1641/0006-3568(2001)051%5b0043:cwatcb%5d2.0.co;2 khumalo, s. & molla, y., 2012, ‘funding shock for kzn wildlife’, the mercury newspaper, 28 september, viewed n.d., from https://www.iol.co.za/news/politics/funding-shock-for-kzn-wildlife-1392062 kiss, a., 2004, ‘is community-based ecotourism a good use of biodiversity conservation funds?’, trends in ecology & evolution 19, 232–237. https://doi.org/10.1016/j.tree.2004.03.010 knight, r., 2010, ‘horse business management: managing a successful yard’, australian veterinary journal 88, 509–509. https://doi.org/10.1111/j.1751-0813.2010.00656.x laycock, h., moran, d., smart, j., raffaelli, d. & white, p., 2009, ‘evaluating the cost-effectiveness of conservation: the uk biodiversity action plan’, biological conservation 142, 3120–3127. https://doi.org/10.1016/j.biocon.2009.08.010 lindsey, p.a., alexander, r.r., du toit, j.t. & mills, m., 2005, ‘the potential contribution of ecotourism to african wild dog (lycaon pictus) conservation in south africa’, biological conservation 123, 339–348. https://doi.org/10.1016/j.biocon.2004.12.002 lindsey, p.a., romanach, s. & davies-mostert, h., 2009, ‘the importance of conservancies for enhancing the value of game ranch land for large mammal conservation in southern africa’, journal of zoology 277, 99–105. https://doi.org/10.1111/j.1469-7998.2008.00529.x mahony, k. & van zyl, j., 2002, ‘the impacts of tourism investment on rural communities: three case studies in south africa’, development southern africa 19, 83–103. https://doi.org/10.1080/03768350220123891 mmopelwa, g., kgathi, d. & molefhe, l., 2007, ‘tourists’ perceptions and their willingness to pay for park fees: a case study of self-drive tourists and clients for mobile tour operators in moremi game reserve, botswana’, tourism management 28, 1044–1056. https://doi.org/10.1016/j.tourman.2006.08.014 moran, d., pearce, d. & wendelaar, a., 1997, ‘investing in biodiversity: an economic perspective on global priority setting’, biodiversity and conservation 6, 1219–1243. https://doi.org/10.1023/a:1018360924079 mossman, s.l. & mossman, a.s., 1976, wildlife utilization and game ranching, report on the recent progress in this field in southern africa, iucn occasional paper 17, 97, international union for the conservation of nature, world wildlife fund and humbolt state university, gland, switzerland. muir-leresche, k. & nelson, r., 2000, private property rights to wildlife: the southern african experiment, international centre for economic research, washington, dc. musengezi, j., 2010, ‘wildlife utilization on private land: understanding the economics of game ranching in south africa’, doctor of philosophy, university of florida. myburgh, e. & saayman, m., 1999, ecotourism in action: practical guidelines and principles, 1st edn., leisure consultants and publications, potchefstroom. ollenburg, c., 2005, ‘worldwide structure of the equestrian tourism sector’, journal of ecotourism 4, 47–55. https://doi.org/10.1080/14724040508668437 parkan, c., 1991, ‘the calculation of operational performance ratings’, international journal of production economics 24, 165–173. https://doi.org/10.1016/0925-5273(91)90163-n parkan, c., 1996, ‘measuring the performance of hotel operations’, socio-economic planning sciences 30, 257–292. https://doi.org/10.1016/s0038-0121(96)00013-4 parkan, c., 1999, ‘performance measurement in government services’, managing service quality 9, 121–135. https://doi.org/10.1108/09604529910257911 parkan, c., 2003, ‘measuring the effect of a new point of sale system on the performance of drugstore operations’, computers & operations research 30, 729–744. https://doi.org/10.1016/s0305-0548(02)00047-3 parkan, c., lam, k. & hang, g., 1997, ‘operational competitiveness analysis on software development’, journal of the operational research society 48, 892–905. https://doi.org/10.1057/palgrave.jors.2600446 parkan, c. & wu, m.l., 1999, ‘measurement of the performance of an investment bank using the operational competitiveness rating procedure’, omega 27, 201–217. https://doi.org/10.1016/s0305-0483(98)00041-3 parker, s. & khare, a., 2005, ‘understanding success factors for ensuring sustainability in ecotourism development in southern africa’, journal of ecotourism 4, 32–46. https://doi.org/10.1080/14724040508668436 pfister, r.e. & tierney, p.t., 2008, recreation, event, and tourism businesses: start-up and sustainable operations, human kinetics, il. plantweb, 2003, white paper: reducing operations and maintenance costs, viewed 13 january 2015, from http://www.plantweb.com porter, s., ferrer, s. & aylward, b., 2003, ‘the profitability of nature tourism in zululand: a survey of private reserves and public protected areas’, in b. aylward & e. lutz (eds.), nature tourism, conservation, and development in kwazulu-natal, south africa, pp. 287–324, the world bank, washington, dc. rahman, i., reynolds, d. & svaren, s., 2012, ‘how “green” are north american hotels? an exploration of low cost adoption practices’, international journal of hospitality management 31, 720–727. https://doi.org/10.1016/j.ijhm.2011.09.008 ridl, j., 2012, ‘ezemvelo kzn wildlife on its knees’, the witness newspaper, 04 october, viewed n.d., from https://www.news24.com/archives/witness/ezemvelo-kzn-wildlife-on-its-knees-20150430 ross, s. & wall, g., 1999, ‘ecotourism: towards congruence between theory and practice’, tourism management 20, 123–132. https://doi.org/10.1016/s0261-5177(98)00098-3 rylance, a., 2017, ‘estimating tourism’s contribution to conservation area financing in mozambique’, tourism and hospitality research 17, 24–33. https://doi.org/10.1177/1467358415613119 shieh, h.s., 2012, ‘does cost efficiency lead to better financial performance? a study on taiwan international tourist hotels’, the journal of hospitality financial management 20, 17–30. https://doi.org/10.1080/10913211.2012.10721889 south african police services (saps), 2016, annual crime report 2015/2016: addendum to the saps annual report, report no. 218/2016, viewed 28 september 2017, from https://www.saps.gov.za/services/c_thumbnail.php?id=282 speelman, s., d’haese, m., buysse, j. & d’haese, l., 2008, ‘a measure for the efficiency of water use and its determinants, a case study of small-scale irrigation schemes in north-west province, south africa’, agricultural systems 98, 31–39. https://doi.org/10.1016/j.agsy.2008.03.006 thirtle, c. & bottomley, p., 1992, ‘total factor productivity in uk agriculture, 1967–90’, journal of agricultural economics 43, 381–400. https://doi.org/10.1111/j.1477-9552.1992.tb00233.x tsaur, s.h., 2001, ‘the operating efficiency of international tourist hotels in taiwan’, asia pacific journal of tourism research 6, 73–81. https://doi.org/10.1080/10941660108722090 van der merwe, p., saayman, m. & krugell, w., 2004, ‘factors that determine the price of game’, koedoe african protected area conservation and science 47(2), 105–113. https://doi.org/10.4102/koedoe.v47i2.86 van loggerenberg, m. & herbst, f., 2010, ‘brand-building: a telling story’, usb leaders’ lab 4(2), 13–17. http://www.usb.ac.za/thoughtprint/leaders%20lab%20pdfs/brand-building%20-%20a%20telling%20story.pdf abstract introduction methods results discussion acknowledgements references about the author(s) bernard w.t. coetzee global change institute, university of the witwatersrand, south africa scientific services, organisation for tropical studies, south africa sam m. ferreira scientific services, sanparks, skukuza, south africa kristine maciejewski centre for complex systems in transition, stellenbosch university, south africa citation coetzee, b.w.t., ferreira, s.m. & maciejewski, k., 2018, ‘challenges and opportunities for monitoring wild nile crocodiles with scute mark-recapture photography’, koedoe 60(1), a1505. https://doi.org/10.4102/koedoe.v60i1.1505 short communication challenges and opportunities for monitoring wild nile crocodiles with scute mark-recapture photography bernard w.t. coetzee, sam m. ferreira, kristine maciejewski received: 07 nov. 2017; accepted: 10 may 2018; published: 19 july 2018 copyright: © 2018. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the global conservation status of nile crocodiles (crocodylus niloticus) was last assessed in 1996. the species presents particular difficulty in monitoring because it can be cryptic, require expertise to handle, and caudal tail tags and transmitters are often lost. some studies advocate mark-recapture techniques based on photograph identification of the unique scute markings of crocodile tails as a non-invasive means of monitoring their populations. researchers developed this method with crocodiles in captivity. in this study, we test the technique under field conditions by monitoring crocodiles from 2015 to 2017 in the sunset dam in the kruger national park. using a cormack-jolly-seber open population model, we found that the dam may host 15–30 individuals, but that there is a high turnover of individuals and much uncertainty in model outputs. the dam’s population thus has high rates of immigration and emigration. the method proved challenging under field conditions, as there was bias in identifying scute markings consistently. the efficient use of the method requires an exceptional quality of photographic equipment. animal crypsis, however, remains an issue. in this study, we discuss how to improve the mark-recapture photography methodology, especially to adapt the technique for citizen science initiatives. conservation implications: using scute mark-recapture photography presents challenges under field conditions. these challenges require innovative, practical and analytical solutions to successfully use the technique before monitoring programmes, aimed at ensuring the persistence of crocodiles in the wild, can be implemented. introduction the nile crocodile, crocodylus niloticus, is an apex predator and keystone species across africa (ashton 2010). it serves as an indicator species and populations are proxies for aquatic ecosystem health (ashton 2010). the current global conservation status of the nile crocodile, as assessed by the international union for conservation of nature (iucn), is least concern, but the species was previously listed as vulnerable from 1982 to 1990 (iucn 2017) and is listed as vulnerable under the south african red list assessment (marais 2014). even so, recent declines in crocodile populations recorded across several rivers and lakes, with particularly marked declines in the olifants gorge within the kruger national park (ferreira & pienaar 2011), raise conservation concerns. consequently, there is a need to ascertain the conservation status of nile crocodile populations. this relies on efficient monitoring techniques. nile crocodiles present particular challenges for monitoring, especially over large spatial areas. aerial surveys are useful for counting crocodiles, but consistently yield lower estimates than spotlight surveys at night (combrink et al. 2011; ferreira & pienaar 2011), especially for smaller size classes. crocodiles can be cryptic, which reduces their detectability (thomas et al. 2010). capture mark-recapture studies rely on individually marked crocodiles using scute-removal techniques or the attachment of numbered plastic tags on their tails, or vhf (very high frequency) transmitters (bourquin 2007; leslie 1997). the capturing of crocodiles, however, can be challenging, as the signal of vhf transmitters is often difficult to detect owing to thick vegetation; it is also time-consuming, expensive and dependent on individual behaviour that may vary (bourquin & leslie 2011). these may also bias the re-catchability of certain sexes, sizes or age classes (bayliss et al. 1987). this is not only as a result of improper signal detection from vhf transmitters but also because of the plastic caudal tags, often cattle ear tags, which fail when they fall off, reducing the robustness of mark-recapture estimates (bourquin 2007; swanepoel 1999). in addition, methods employed to monitor key species may have negative impacts on the welfare of individuals as well as species persistence. the fitment of devices on birds, for instance, has significant effects on energy expenditure, nesting likelihood (barron et al. 2010) and population dynamics (saraux et al. 2011). ethical considerations should be a key element of biological monitoring designs (putman 1995), which require the development of less invasive monitoring techniques. researchers have advocated a scute identification technique to better monitor crocodile populations (bouwman & cronje 2016; swanepoel 1996). it is less invasive, less time-consuming and potentially more accurate. the scute markings on the tail provide a unique identification for each crocodile, allowing comparisons between individuals in a population. this method does not require the capture of an animal, but observers need to positively identify an individual visually or through the use of photography. using mark-recapture photography, the method may ‘capture’ a portion of a population with photographs, and identify and catalogue unique markings. then the proportion of ‘marked’ individuals recaptured in subsequent photographic sampling events allows the estimation of population sizes (bouwman & cronje 2016). this approach has potential use for population monitoring, research and application for citizen science initiatives aimed at monitoring individuals in the wild (bouwman & cronje 2016). the development of the mark-recapture photography techniques, however, focused on captive crocodiles (bouwman & cronje 2016; swanepoel 1996). identifying animals in captivity enables researchers to handle specimens, which allows for clearly identified scute markings, removing any issues with regard to individual detectability. here, we develop and test a monitoring protocol by assessing the suitability of mark-recapture photography for crocodiles under typical field conditions. our first aim is to use the technique to calculate the population size and monitor crocodiles at a single dam in the kruger national park. our second aim is to report on the challenges and opportunities with this approach. this allows us to better guide the use of such techniques when operationalised under field conditions. to test the potential role of citizen scientists in such initiatives, we conducted our study at a popular tourist site, at a dam frequented by crocodiles. our study thus also replicated techniques with equipment that we considered tourists might typically employ and in conditions they might face. methods study site and methods this study took place over two years at sunset dam (25°06.972′ s 031°54.729′ e) near the lower sabie rest camp in the kruger national park. sunset dam is 200 m from the sabie river. tourists have regular sightings of crocodiles at this dam. our sampling focused on four events of three days each during april 2015, february 2016, september 2016 and february 2017. sampling at the study site was only conducted when heavy wind, rain or overcast conditions were absent. each sampling event consisted of counting crocodiles from a game drive vehicle parked at sunset dam. observers counted crocodiles every 30 min, starting at 09:00 and ending at 12:00, noting all visible crocodiles on land and in the water at the time. when visible, observers also photographed the tails of all crocodiles for later individual identification using the scute marking technique (details follow). to emulate the equipment that tourists might use, we used a combination of photography techniques including a dslr (75–300 mm lens dslr canon els rebel t3i camera), a canon h50 powershot with 50 × digital zoom and ‘digiscoping’ by taking photographs through a nikon ed field spotting-scope, mounted on a tripod inside the vehicle. during the first sampling event (april 2015), observers also measured the distance to all crocodiles, noted from the observation point with a laser range finder (foresty 550 6 × 21). the distinct markings on scutes running laterally down the tail allows the individual identification of nile crocodiles (bouwman & cronje 2016; swanepoel 1996). we applied swanepoel’s (1996) method, although subsequent work suggests an alternative 11-scute system (see bouwman & cronje 2016). we, however, generated a unique nine-scute code by recording the markings of the last anterior, unfused scute and the subsequent scutes moving towards the head (figure 1). the code was generated by recording the number of black marks on each scute and the relative position of the scute on which the mark(s) were located. if there were multiple black marks per tail segment, we repeated the corresponding scute number to the number of those marks. adding parentheses to numbers represented single marks that occasionally extend across different scutes. square brackets signified marks that are of a lighter colouration. figure 1: conceptual diagram demonstrating the scute encoding procedure followed, based on photographs of crocodile tail scutes. we created a crocodile photographic database containing either the left or the right scute markings identified during the study. each record contains a unique id; an exemplar photograph, whether it was the left or right tail observed; and a unique scute marking code, whether it was present or absent during subsequent sampling events. at least four observers captured data from photographs, assigned scute codes by consensus and drew conceptual diagrams of the tails (figure 1b). four observers in 2017 set out to test the level of agreement rates between different observers and independently captured data for each newly photographed individual. we considered ‘disagreement’ between assigned scute codes to be when three or more scute numbers differed from at least two observers among the codes assigned to each crocodile. data analysis firstly, we provide summary statistics of the overall crocodile photographic database created during the study period. then, because it was particularly important to survey crocodiles when they were basking out of the water, and available for photographic mark-recapture, we estimated the change in crocodile abundance over time during the half-hourly sampling periods recorded on both land and in the water. to estimate the population size in order to determine changes at sunset dam in our dataset, we applied mark-recapture analysis. for this purpose, we considered individuals with unique left and unique right scute codes as ‘marked’. subsequent photographs and scute codes matched to previously coded individuals were ‘recaptured’. we implemented a cormack-jolly-seber (cjs) open population model to calculate the estimated population size (pledger, pollock & norris 2003). model implementation in ‘r’ (2017) used the package ‘mra’ (amstrup, mcdonald & manly 2005) to calculate a population size estimate and standard error. since crocodiles have different leftand right-hand scute marks (bouwman & cronje 2016; swanepoel 1996), we ran a ‘left’ model (left side of tail photographs only) and a ‘right’ model (right side of tail photographs only). results a total of 113 left and right scute markings made up our sample during 2015–2017. after a total of 36 h of observation, only three individuals had both leftand right-hand sides photographed with certainty (based on visually tracking the individual for changing orientation). our data comprised a total of 56 left tail and 57 right tail scute markings. the number of crocodiles on land decreased during the morning counts (figure 2; pearson’s r = −0.94; p < 0.01; n = 7), while those counted in water did not (figure 2; pearson’s r = 0.65; p = 0.117; n = 7). standard deviation bars, however, overlapped half-hourly, indicating no appreciable peak in numbers of crocodiles basking on land (figure 2). when comparing individual scute markings, the four observers collecting data during february 2017 disagreed on 8 out of 14 photographs they collected in total, a 57% disagreement rate. photographed crocodiles were on average more than 120 m away (mean = 124 m; sd = 37.5 m; n = 53). figure 2: individual counts at half-hourly intervals of four sampling events, during 12 days, of crocodiles seen on land and in the water in and around sunset dam at lower sabie, kruger national park. overall, there was a low recapture rate. we recaptured only 13 scute markings and no unique scute markings were captured in all four sampling events. despite the ratio of left to right photographed tails being similar (56 left vs. 57 right), the mark-recapture analysis indicated different population size estimates during sampling events, as the analysis is dependent on the accumulating recapture rate. in 2017, which indicates the longest time interval of mark-recapture data, population sizes were 11 (± 5.0 se) and 18 (± 8.1 se) for the left and right tails, respectively (figure 3). these modelled population size estimates overlapped with the data obtained from morning counts (see figure 2). model fit can be considered robust (left model: aic = 54.17; log likelihood = −24.08; deviance = 48.17; n = 4 and right model: aic = 41.61; log likelihood = −16.80; deviance = 33.61; n = 4). figure 3: modelled crocodile abundance across three repeat sampling events at sunset dam, lower sabie, kruger national park, using a cormack-jolly-seber mark-recapture analysis. see text for model coefficients. discussion the crocodile population at the study site is dynamic. low rates of individual recaptures suggest high rates of movement of crocodiles to and from sunset dam during the study period. crocodiles most likely move into and from the nearby sabie river. despite this finding, the population model demonstrated a relatively consistent number of crocodiles across years. we infer that 15–30 crocodiles regularly use the dam. the low rate of recaptures emphasise that the use of the dam by crocodiles is in flux. the results suggest high levels of emigration and immigration, consistent with observations that crocodiles can move long distances over land. nile crocodiles in the kruger national park moved up to 36 km along the olifants river between south africa and mozambique (swanepoel 1999). fergusson (2010) recorded crocodile movements of up to 90 km. crocodile movement may be affected by changes in water levels. in the dry season when water levels receded, crocodiles moved from the ume river towards the kariba dam (games 1990). the same was found in the amazon basin with melanosuchus niger and caiman crocodylus retreating from the flooded forests in the wet season to the lakes in the dry season (ron,vallejo & asanza 1998). the error rates in our modelled population estimates are high, as they cannot account for crocodile tails being obscured from view in the water (which may underestimate numbers), and cannot account for double counting either sides of the same individual (which would overestimate numbers). the development of statistical techniques that accommodate single-side marking and detectability biases can improve the use of unique tail markings to estimate nile crocodile population sizes. the low recapture rate and difficulties with scute identification using the photographic technique will further increase error in the model. the present study tested and demonstrated the feasibility of a photographic mark-recapture technique for monitoring nile crocodiles. however, the method proved challenging under field conditions and for citizen science application. the use of this approach requires addressing three major challenges. firstly, successful identification relied heavily on the quality of the photographs. only photographs of exceptional quality were suitable. slight decreases in image resolution or poor lighting rendered photographs unsuitable. in addition, most images taken in the study were at distances greater than 100 m. our experience suggests that an slr camera lens of 500 mm or greater could produce consistent and reproducible crocodile images. secondly, the identification of scute markings was subjective for most photographs, regardless of their quality. the disagreement rate between observers of more than 50% implies observer difficulty and potentially bias in identifying scute markings from photographs. we attempted to overcome this by determining scute markings by consensus. although statistical techniques can address observer bias in aerial surveys, for instance (lubow & ransom 2016), such bias is problematic to resolve analytically in our case. we propose the use of paper cards (figure 1b) to help facilitate identification between observers. multiple observers classify the scutes independently and then reach the final codes by consensus. the approach thus requires training observers for consistency. we acknowledge that the use of automated picture identification software (e.g. kuhl & burghardt 2013) and machine learning (e.g. michalski, carbonell & mitchell 2013) could greatly enhance the consistent identification of scute markings. the third challenge is species crypsis, which alters the detectability of crocodile tails and so reduces the ability to monitor populations effectively (thomas et al. 2010). observers can only photograph basking crocodiles. even if away from the water, debris and mud or the orientation of the individual may obscure tail scutes. orientation is particularly problematic as scutes differ between the left and right side of the tail, and our results demonstrate that it may alter population estimates. nile crocodiles may also spend more time out of the water during winter, and therefore sampling may better detect the species during june–july. while the scute marking method would be appropriate for captured animals, using photography presents challenges under field conditions. these challenges require innovative, practical and analytical solutions to successfully use a photographic mark-recapture method based on scute markings that can inform interventions aimed at ensuring the persistence of crocodiles in the wild. acknowledgements the authors thank the following observers: tavis dalton, sam kubica, olivia vennaro, claire weston, brian brooks, lauren chang, catherine craighill, tatiana henry, katie diggs, brianna mathias, annie stevens, jana woerner, joey binder, max israelit, drew perlmutter and henry stevens. funding was provided by the organisation for tropical studies, south africa. h. coetzee kindly redrew figure 1. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions k.m. and s.m.f. conceptualised the study. k.m. and b.w.t.c. conducted and coordinated all the fieldwork. b.w.t.c. performed the analysis and wrote the first draft. all the authors contributed to subsequent drafts of this article. references amstrup, s.c., mcdonald, t.l. & manly, b.f.j., 2005, handbook of capture-recapture analysis, princeton university press, princeton, nj. ashton, p.j., 2010, the demise of the nile crocodile (crocodylus niloticus) as a keystone species for aquatic ecosystem conservation in south africa: the case of the olifants river, csir report, csir, pretoria. barron, d.g., brawn, j.d. & weatherhead, p.j., 2010, ‘meta-analysis of transmitter effects on avian behaviour and ecology’, methods in ecology evolution 1, 180–187. https://doi.org/10.1111/j.2041-210x.2010.00013.x bayliss, p., 1987, ‘survey methods and monitoring within crocodile management programs’, in g.j.w. webb, s.c. manolis & p.j. whitehead (eds.), wildlife management: crocodiles and alligators, pp. 157–176, surrey beatty and sons, sydney. bourquin, s.l., 2007, ‘the population ecology of the nile crocodile (crocodylus niloticus) in the panhandle region of the okavango delta, botswana’, phd thesis, stellenbosch university, stellenbosch. bourquin, s.l. & leslie, a.j., 2011, ‘estimating demographics of the nile crocodile (crocodylus niloticus laurenti) in the panhandle region of the okavango delta, botswana’, african journal of ecology 50, 1–8. https://doi.org/10.1111/j.1365-2028.2011.01285.x bouwman, h. & cronje, e., 2016, ‘an 11-digit identification system for individual nile crocodiles using natural markings’, koedoe 58, 1–6. https://doi.org/10.4102/koedoe.v58i1.1351 combrink, x., korrûbel, j.l., kyle, r., taylor, r. & ross, p., 2011, ‘evidence of a declining nile crocodile (crocodylus niloticus) population at lake sibaya, south africa’, south african journal of wildlife research 41, 145–157. https://doi.org/10.3957/056.041.0201 fergusson, r., 2010, wildlife survey phase 2 and management of human–wildlife conflicts in mozambique, survey of crocodile populations in mozambique, vol. 2 part 5, agreco g.e.i.e., brussels, belgium. ferreira, s.m. & pienaar, d., 2011, ‘degradation of the crocodile population in the olifants river gorge of kruger national park, south africa’, aquatic conservation 21, 155–164. https://doi.org/10.1002/aqc.1175 games, i., 1990, ‘the feeding ecology of two nile crocodile populations in the zambezi valley’, phd thesis, university of zimbabwe, harare. iucn, 2017, iucn red list: crocodylus niloticus, viewed 23 october 2017, from http://www.iucnredlist.org/details/46590/0 kuhl, h.s. & burghardt, t., 2013, ‘animal biometrics: quantifying and detecting phenotypic appearance’, trends in ecology and evolution 28, 432–441. https://doi.org/10.1016/j.tree.2013.02.013 leslie, a.j., 1997, ‘the ecology and physiology of the nile crocodile, crocodylus niloticus’, phd thesis, drexel university philadelphia, pa. lubow, b.c. & ransom, j.i., 2016, ‘practical bias correction in aerial surveys of large mammals: validation of hybrid double-observer with sightability method against known abundance of feral horse (equus caballus) populations’, plos one 11, e0154902. https://doi.org/10.1371/journal.pone.0154902 marais, j., 2014, ‘crocodylus niloticus (laurenti, 1768)’, in m.f. bates, w.r. branch, a.m. bauer, m. burger, j. marais, g.j. alexander et al., (eds.), atlas and red list of the reptiles of south africa, lesotho and swaziland, suricata 1, south african national biodiversity institute, pretoria. michalski, r.s., carbonell, j.g. & mitchell, t.m., 2013, machine learning: an artificial intelligence approach, springer science & business media, palo alto, ca. pledger, s., pollock, k.h. & norris, j.l., 2003, ‘open capture-recapture models with heterogeneity: i. cormack-jolly-seber model’, biometrics 59, 786–794. https://doi.org/10.1111/j.0006-341x.2003.00092.x putman, r.j., 1995, ‘ethical considerations and animal welfare in ecological field studies’, biodiversity and conservation 4, 903–915. https://doi.org/10.1007/bf00056197 ron, s.r., vallejo, a. & asanza, e., 1998, ‘human influence on the wariness of melanosuchus niger and caiman crocodilus in cuyabeno, ecuador’, journal of herpetology 32, 320. https://doi.org/10.2307/1565444 saraux, c., le bohec, c., durant, j.m., viblanc, v.a., gauthier-clerc, m., beaune, d. et al., 2011, ‘reliability of flipper-banded penguins as indicators of climate change’, nature 469, 203–206. https://doi.org/10.1038/nature09630 swanepoel, d.j.g., 1996, ‘identification of the nile crocodile crocodylus niloticus by the use of natural tail marks’, koedoe 39, 113–115. https://doi.org/10.4102/koedoe.v39i1.287 swanepoel, d.j.g., 1999, ‘movements, nesting and the effects of pollution on the nile crocodile crocodylus niloticus in the olifants river, kruger national park’, msc thesis, university of natal, pietermaritzburg. thomas, l., buckland, s.t., rexstad, e.a., laake, j.l., strindberg, s., hedley, s.l. et al., 2010, ‘distance software: design and analysis of distance sampling surveys for estimating population size’, journal of applied ecology 47, 5–14. https://doi.org/10.1111/j.1365-2664.2009.01737.x biggs.qxd experimental burn plot trial in the kruger national park: history, experimental design and suggestions for data analysis r. biggs, h.c. biggs, t.t. dunne, n. govender and a.l.f. potgieter biggs, r., h.c. biggs, t.t. dunne, n. govender and a.l.f. potgieter. 2003. experimental burn plot trial in the kruger national park: history, experimental design and suggestions for data analysis. koedoe 46(1): 1-15. pretoria. issn 0075-6458. the experimental burn plot (ebp) trial initiated in 1954 is one of few ongoing long-term fire ecology research projects in africa. the trial aims to assess the impacts of different fire regimes in the kruger national park. recent studies on the ebps have raised questions as to the experimental design of the trial, and the appropriate model specification when analysing data. archival documentation reveals that the original design was modified on several occasions, related to changes in the park's fire policy. these modifications include the addition of extra plots, subdivision of plots and changes in treatments over time, and have resulted in a design which is only partially randomised. the representativity of the trial plots has been questioned on account of their relatively small size, the concentration of herbivores on especially the frequently burnt plots, and soil variation between plots. it is suggested that these factors be included as covariates in explanatory models or that certain plots be excluded from data analysis based on results of independent studies of these factors. suggestions are provided for the specification of the experimental design when analysing data using analysis of variance. it is concluded that there is no practical alternative to treating the trial as a fully randomised complete block design. key words: burn plots, fire, policy, experimental design, season, frequency. r. biggs, csir environmentek, p.o. box 395, pretoria, 0001, republic of south africa; t.t. dunne, statistical sciences, university of cape town, rhodes gift 7707, republic of south africa; h.c. biggs, n. govender and a.l.f. potgieter, south african national parks, private bag x 402, skukuza, 1350, republic of south africa. issn 0075-6458 1 koedoe 46/1 (2003) introduction the experimental burn plot trial initiated in 1954 in the kruger national park is one of few long-term fire ecology research projects in africa. although attracting much energy and attention during its first decade, scientific interest in the trial declined considerably between the mid-1960s and the late 1990s. this was largely due to a perception that the trial was irrevocably confounded by 'unnaturally' high levels of herbivory (trollope et al. 1998). following a management review in 1997, the possibility of closure of the trial prompted a surge of scientific activity on the plots. the ensuing analysis of data has raised questions about the experimental design of the trial and how the design should be specified in analysis of variance (anova). in the hope of uncovering the original documentation describing the design of the trial, a detailed search of archival documentation was undertaken. this paper outlines the history of the ebp trial and describes the current experimental design. suggestions for the appropriate specification of explanatory models in anova are provided. the paper also discusses confounding factors in the trial, such as herbivory and soil variation, and suggests approaches for taking these into account. history of the ebp trial the trial was started in 1954 by the newly formed biological section of the kruger national park in response to a lack of information on the role of burning in the preserbiggs.qxd 2005/12/09 10:21 page 1 vation of fauna and flora (national parks board 1953). the objectives of the trial were to assess the influence of fire on what were perceived as the four most important veld types (landscapes) and their associated biotic populations in the kruger national park (national parks board 1954). it was envisaged that vegetation surveys would be conducted annually, while animal life would be observed from 'time to time'. the intention was that the investigation would, in due course, be expanded to other important vegetation landscapes (nel 1953). prior to the commencement of the trial, veld burning had become a highly contentious management practice in south africa, and a negative attitude towards controlled burning had developed (trollope et al. 1998). the effects of the fire suppression policy applied in the kruger national park between 1948 and 1956 (van wilgen et al. 2000), led to concerns about bush encroachment, deterioration of grazing and fuel accumulation (trollope et al. 1998). in 1953, approximately one quarter of the kruger national park burnt in uncontrolled wild fires (van der schijff 1958; brynard 1971), raising significant concern about the impacts of fire on the wild plants and animals the park was mandated to protect and preserve. while the need for research on fire impacts had been a primary reason for the appointment of the first biologist in 1951, the fires of 1953 provided the impetus for drawing up the park's first concrete fire research proposal (nel 1953). the original proposal was based on recommendations for a fire research programme in the kruger national park, emanating from a special visit in 1950 by dr. turpin, dr. rowland and mr. bonsma of the department of agriculture. the proposal was that the trial consist of the following six fully randomised treatments in each of the four major veld types: a fire exclosure, an annual and triennial burn in winter, an annual and triennial burn in spring after the first rains, and a triennial burn in late summer (nel 1953). however, the actual plots laid out between 1954 and 1957 consisted of the following seven treatments: a fire exclosure, an annual burn in august and biennial burns in august, october, december, february and april (van der schijff 1958). each treatment was replicated four times within each landscape, with replicates (strings of treatment plots) spaced across the landscape to maximise their representativity of the landscape (van der schijff 1958). the reason for the departure from the original six, carefully considered treatments was not recorded. during 1955, the board approved the expansion of the experiment to include annual and triennial treatments in each of the five existing treatment seasons (i.e., february, april, august, october and december) (national parks board 1955a). the triennial treatments were added between 1956 and 1958 by demarcating five additional plots at one end of each of the existing replicates. where this was not feasible, the additional plots were laid out parallel to the existing replicates (national parks board 1956; van der schijff 1958). the reason why the additional annual treatments were never laid out is not mentioned in the archival material, but probably relates to the physical limitations of burning annually in seasons other than winter. in addition, the fire policy was changed in 1957 to a formal system of burning once every three years in spring (van wilgen et al. 2000), so that the effects of triennial burns would have been of greater interest than those of annual burns. the meaningfulness of certain treatments was questioned early on in the trial (national parks board 1962). according to this report, an investigation into the background of the trial clearly showed that a number of treatments were included to illustrate or prove the damaging impacts of natural and artificially ignited fires, rather than to meet the original objectives. the report concludes that this value-loaded agenda resulted in the inclusion of certain unnecessary treatments and the exclusion of some crucial treatments, such as treatments with a lower burning frequency or a treatment directly after the first spring rains which was not coupled to a specific calendar month (national parks board 1962). koedoe 46/1 (2003) 2 issn 0075-6458 biggs.qxd 2005/12/09 10:21 page 2 following detailed vegetation surveys conducted during the early 1970s, it was concluded that the february treatments did not differ significantly from the fire exclosure (national parks board 1976). a decision was taken to divide the february plots in half and change the treatments on the resulting four subplots to fourand six-year burns, both before and after the first rains of the season (national parks board 1976). this decision accompanied an amendment of the park's fire policy in 1975, which allowed for longer periods between fires in drier areas (van wilgen et al. 2000). the decision to abandon the february treatments was, however, recalled soon after on the basis that midsummer burns were practised as part of the general fire policy (national parks board 1976) and should therefore be retained in the trial (national parks board 1981). where plots had already been subdivided, one half retained the original treatment (february biennial or triennial) and the other half was intended to receive respectively a fouror six-year treatment before the spring rains. these fourand six-year treatments were soon modified to being applied after the first spring rains because it was felt that, since the general fire policy was to burn after the spring rains, treatments with this timing would yield more useful information. issn 0075-6458 3 koedoe 46/1 (2003) fig. 1: the trial consists of four replicates (strings of plots) in each of four major vegetation landscapes in the park. each replicate consists of 12 (sourveld and combretum replicates) or 14 (knobthorn/marula and mopani replicates) plots, each of which receives a unique fire treatment (see table 2). various soil, vegetation and animal surveys were carried out during the first 30 years of the trial (national parks board 1984). botanical surveys were conducted on a regular basis from the inception of the trial until the 1980s and narrative-style summaries of results generally recorded in the annual reports of the biological section. these reports may provide a qualitative means of investigating the rate of vegetation change in the trial. survey methods were of particular concern (national parks board 1960; national parks board 1962) and were addressed by a number of studies (brynard et al. 1960; von broembsen 1964; national parks board 1974). several studies investigated treatment-induced soil changes on the plots (national parks board 1971; national parks board 1978; webber 1979). further details of studies conducted on the ebps, including the detailed half-hectare vegetation surveys conducted during the early 1970s and the 'wrap-up' surveys conducted in the late 1990s, are given by trollope et al. (1998) and govender et al. (2003). details of more recent studies can be found biggs.qxd 2005/12/09 10:21 page 3 in the annual project reports compiled by the scientific services division of the park. current experimental design in statistical terms, the ebp trial is a fractional factorial, partly randomised, complete block design. the experimental design terminology used in this paper is defined in table 1. the trial is referred to as a fractional factorial design as all treatment frequencies do not occur in combination with all treatment seasons (table 2). it is a complete block design as all treatments are applied once within each replicate. the trial is only partly randomised as described below. the trial currently consists of 16 replicates (strings of plots), made up of four replicates in each of the four major gertenbach (1983) landscapes: lowveld sour bushveld of pretoriuskop (sandy granitic soils), combretum collinum/combretum zeyheri woodland (sandy granitic soils), sclerocarya birrea/ acacia nigrescens savanna (clay basaltic soils), and colophospermum mopane shrubveld on basalt (clay basaltic soils) (fig. 1). koedoe 46/1 (2003) 4 issn 0075-6458 fig. 2: each replicate consists of two randomised sections of plots. the first section consists of the fire exclosure, august annual and the five biennial treatments. the second, adjoining section consists of the five triennial treatments. the fourand six-year treatments (only on the mopani and knobthorn/marula replicates) were created by dividing the february biennial and triennial treatments respectively in half. each replicate consists of 12 full plots, each of which receives a different fire treatment. in the knobthorn/marula and mopani landscapes, october four and six year treatments were introduced in the late 1970s by respectively dividing the february biennial and triennial treatments in half; these landscapes therefore have 14 plots per replicate (table 2). the plots within each replicate are not fully randomised, but consist of two adjacent randomised sections (fig. 2). each full plot is approximately 7 ha in size (370 m x 180 m) and protected by a double firebreak. only four treatment randomisations were constructed and each randomisation applied once in each landscape. a replicate in, for example, the lowveld sour bushveld of pretoriuskop, will have a different plot layout to all other replicates in the sourveld, but an identical layout to one replicate in each of the other three landscapes. it appears that the four treatment randomisations were not randomly allocated to the replicates in a particular landscape, but were assigned starting at either the most southerly or most northerly replicate. table 1 experimental design terminology as used in this paper term definition treatment a combination of fire season and frequency (e.g. annual burn in august) applied to a specific plot. plot the smallest experimental unit, approximately 7 ha in size. plots are grouped together in replicates (fig. 2). replicate a string of 12 or 14 plots. there are four replicates in each land scape (fig. 1). trial the full set of 16 replicates, or 208 plots, spaced across four land scapes. biggs.qxd 2005/12/09 10:21 page 4 confounding factors confounding refers to a situation where the effects of experimental treatments cannot be separated from other factors that might be causing the observed differences. the ebp trial was designed with the aim of testing the effects of different seasons and frequencies of burning. these effects have been partly obscured by factors not fully taken into account by the experimental design, most importantly, herbivory, artificial water provision and soil variation. herbivory and artificial water provision it was realised from the outset that the trial would be characterised by strong fire-herbivory interactions. herbivore numbers typically increase for a period following fire (scholes & walker 1993). the small size of the plots (approximately 7 ha) compared to the scale of fires in the kruger national park (average fire size approximately 2500 ha and median fire size approximately 1000 ha, van wilgen et al. 2000) result in 'unnaturally high' concentrations of herbivores on the plots (van der schijff 1958; gertenbach & potgieter 1975; van wyk 1975). this problem is mentioned in almost every annual report and highlights the concern it raised, especially with regard to frequently burnt plots. the implication that the plots are therefore not representative of processes in the wider landscape is one of the main reasons for a widely-held perception (trollope et al. 1998) that the trial is of little relevance. the large plot size (in experimental terms) (nel 1953) and the intended practice of burning veld in the immediate vicinity of the plots when applying treatments (van der schijff 1958) were measures aimed at reducing the impacts of herbivory. post-fire herbivory has been exacerbated by the introduction of artificial, permanent waterholes into the areas surrounding certain replicates (gertenbach & potgieter 1975). recently gorman (in prep.) compared herbivore numbers on the plots to those in the adjacent non-experimental areas. initial results indicate that there are significantly more herbivores on the plots than in the adjacent areas, as well as significant variations in herbivore numbers between different treatments. in response to the herbivory problem, herbivore exclosures were erected during the 1970s in the three southern landscapes in which the trial was being conducted to determine the effect of fire in the absence of herbivory (national parks board 1972-1974; national parks board 1988-1990). each exclosure contained a number of treatment plots that were significantly smaller than the issn 0075-6458 5 koedoe 46/1 (2003) table 2 current fire treatments applied in each replicate of the ebp trial. treatments were started between 1954 and 1958. the fourand six-year treatments were started between 1976 and 1979 season frequency february april august october december (years) (late summer) (autumn) (late winter) (after first (mid-summer) spring rains) 1 x 2 x x x x x 3 x x x x x 4 xa 6 xa >45 fire exclosure aonly on the knobthorn/marula plots (landscape 17) and mopani plots (landscape 23) biggs.qxd 2005/12/09 10:21 page 5 koedoe 46/1 (2003) 6 issn 0075-6458 table 3 suggestions for model specification when analysing ebp data using analysis of variance (anova). these model elements apply more broadly to the specification of generalised linear models, such as log-linear and logistic models. the suggestions below relate to the analysis of data from a single landscape. when analysing data from multiple landscapes, model specification becomes more complex (see text) issue blocking pseudoreplication estimation of fire season and frequency effects problem in heterogeneous environments, background variation may obscure treatment effects. such variation can be reduced by grouping experimental units (plots) into 'blocks' with similar background conditions. repeated measurements on individual plots are incorrectly treated as independent replicates in the model. a very common mistake in ecological field experiments (hurlbert 1984; crawley 1993). as a result of the season and frequency factors not being fully crossed or complete factorials (i.e. not all fire frequencies are applied across all seasons, see table 2), the separate effects of season and frequency of burn can only be examined for biennial and triennial treatments. for the other treatments (august annual, october four and six yearly treatments), only the combined influence of season and frequency can be investigated. suggested approach replicates should be specified as blocks in the model to remove the between-replicate variation from the overall variation in the data, thereby allowing more powerful tests of treatment effects. replicates should be specified as a random factor, as they represent a subset of the landscape about which conclusions are being drawn. 1) calculate the mean value of observations on each plot, and use the single mean value for each plot as input into the model. this, however, excludes the possibility of testing for interactions between factors. if interactions are likely to be important, calculate two mean values for each plot by randomly dividing the observations on each plot into two halves. 2) individual observations (instead of plot means) can be included in the analysis by specifying ‘plot’as an additional blocking factor nested within ‘replicate’. this does not change the analysis results. the simpler approach of using mean values as input is therefore recommended. where all treatments are included in a study, and it is of interest to separate the effects of season and frequency, a possible approach is to run two separate analyses (see enslin et al. 2000): 1) an analysis which includes all treatments without separating the effect of season of burn from that of frequency (i.e. one explanatory variable 'treatment' with 12 or 14 levels), and; 2) a second analysis which includes only the biennial and triennial treatments, with season and frequency specified as independent explanatory variables (season with five levels, and frequency with two levels). results should be adjusted for multiple testing using the bonferroni procedure or other suitable multiple comparisons adjustment (quinn & keough 2002). treatment should be specified as a fixed factor in the model. biggs.qxd 2005/12/09 10:21 page 6 ebps (exclosure plot size 0.04 ha; total exclosure size 2 ha). no results were compiled from these herbivore exclosures, and treatments in the last of these camps, the makhohlola exclosure, were stopped during the early 1990s. soil variation soil variation between replicates in the same landscape, as well as within individual replicates, also raises the issue of representativity of the trial (national parks board 1971; van wyk 1975; trollope et al. 1998). webber (1979) concluded that the plots in his study area (the combretum woodland) were not fully representative of the area and that the original siting of the plots should have been preceded by a soil survey. replicates in the granites are mainly located on the relatively homogenous watersheds (crests), which comprise the majority of the area in these landscapes. this implies that the effects of fire on the low-lying parts of the drainage basins cannot be inferred from the trial. the geomorphic and soil representativity of the plots has been investigated by venter (in prep.), who rated each plot for representativity of its own replicate, as well as of the landscape in which it lies. each replicate was also rated as a whole for representativity of the landscape in which it is located. the conclusion was that the majority of plots are fair to good representatives of the geomorphic and soil characteristics of their landscapes, but that there are a few outliers. treatment compliance more than 80 % of the total number of scheduled burns that should have taken place to date have been applied. appendix 1 tabulates the treatment history in each of the four landscapes in which the trial is being conducted. the main reason for deviation from the planned schedule was that certain plots could not be burnt in specific years because the vegetation cover was either too sparse issn 0075-6458 7 koedoe 46/1 (2003) problem 'unnaturally high' herbivory on certain plots, particularly frequently burnt plots and plots in the vicinity of artificial waterpoints. certain plots are located on soils that are not representative of the general landscape. current differences between plots may reflect not only the effect of differing treatments, but may also be partly related to variation between plots prior to the start of the trial. suggested approach 1) include 'distance to nearest permanent water' as a covariate (see enslin et al. 2000). 2) use the herbivore density data of gorman (in prep.) as a covariate in the model. use the findings of venter (in prep.) as a basis for excluding certain plots or replicates as unrepresentative of the landscape in which they are located. this could also be done using multivariate analysis. 1) include the 1954 baseline vegetation data as a covariate in analyses. 2) define the response variable as the change between 1954 and present (if baseline data were collected for the variable of interest) including a baseline covariate may partly correct for soil variation e.g. local depressions along a replicate string. table 3 (continued) issue herbivory soil variation pre-treatment variation between plots biggs.qxd 2005/12/09 10:21 page 7 (due to drought and/or herbivory impact) or conditions were too moist for burning. a number of accidental burns have occurred, either from fires outside the plots jumping the firebreaks or due to fires jumping between plots during treatment applications. many of the accidental burns occurred during the first few years of the trial before the firebreaks were widened (national parks board 1956), and led to the swapping around of several treatment plots during the early years of the trial (national parks board 1955a). other deviations from the original plot layout resulted from plots being damaged by roadworks during the initial demarcation (national parks board 1955b). a number of accidental burns have recently occurred on the fire exclosure plots (napi, satara, mbyameti and n'shawu replicates) due to the accumulation of biomass resulting from several years of good rain (appendix 1). suggestions for sampling and data analysis the design of the ebp trial poses a number of challenges for sampling and analysing data from the trial. while the nature of these problems will vary depending on the specific study, several issues are likely to crop up in almost all studies. sampling the sequence of rainfall-fire interaction on different plots is likely to have an effect on many parameters of interest (mentis & bailey 1990; oelofse et al. 1999), and raises the question of comparable periods of observation. rather than sampling across different treatments at the same point in time, it may be more appropriate to sample certain responses at a set period after a burning treatment has been applied. however, as treatments are applied in different years, such sampling would need to account for interannual variation in factors such as rainfall. by sampling in this manner, it may be possible to conduct specific analyses to determine the generality of post-burn phenomena such as herbivory across treatments. data analysis data from the ebp trial typically consist of observations (e.g. tree height, termite counts) collected from a selected number of treatments in a specific landscape. usually multiple observations are made on each plot, and all observations are made at a single time period (i.e. the aim is to detect differences between treatments at a specific point in time, rather than to observe changes on a specific treatment over a period of time). such data are typically analysed using analysis of variance (anova). it is important that the design be correctly specified in the anova, in order to ensure that the correct error term is used in hypothesis testing. table 3 provides suggestions for model specification where data have been collected from several replicates in a single landscape. herbivory and soil differences between plots can be accounted for to some extent by making use of results from independent studies of these factors. where data have been collected from multiple landscapes, 'landscape' should be specified as an additional explanatory factor, and 'replicate' should be specified as nested within 'landscape'. the lack of full randomisation within replicates (each replicate consists of two separate randomised sections of plots (fig. 2)), could be accounted for by regarding the trial as having a split-plot design. 'section' would be specified as nested within 'replicate', and 'plot' as nested within 'section'. in this interpretation, frequency (biennial or triennial) is applied per section, and season of burn applied to the plots within each section. specifying the analysis in this way is, however, likely to be unnecessarily complicated, and would be largely theoretical as treatment frequency is not in reality applied separately from treatment season. where multiple observations on specific plots are conducted over a period of time, and this generates time series data, observations cannot simply be regarded as indepenkoedoe 46/1 (2003) 8 issn 0075-6458 biggs.qxd 2005/12/09 10:21 page 8 dent. in some cases, a repeated measures form of anova could be used, with time specified as either an explanatory or blocking factor. in other cases, specific time series analysis techniques may be necessary. three sets of detailed vegetation surveys are available for quantitatively investigating the rate and direction of vegetation change: i) the baseline surveys conducted prior to the start of the trial, ii) the surveys conducted in the early 1970s, and iii) the 'wrap-up' surveys conducted in the late 1990s. the survey method used in the 1970s survey, however, differs substantially from the method used in the baseline and wrap-up surveys and limits the use of this data. inference the small size of the plots in comparison to the typical area burnt by a fire in the park (van wilgen et al. 2000), implies that inference about scale-dependent processes should be approached with caution. the lack of randomisation in the experiment has resulted in a number of treatments nearly always being adjacent, which may have implications for parameters that are sensitive to edge effects. for example, all triennial treatments are located next to at least one other triennial treatment, 75 % of exclosure plots lie adjacent to a december biennial treatment, and all october four-year treatments lie next to a february two-year treatment. edge effects may be important especially on the half-size plots (february twoand three-year and october fourand six-year treatments on the basalts). the responses on the half-size october fourand six-year treatments should be interpreted as the composite effect of 25 years of a february twoand three-year treatment, followed respectively by 20 years of an october fourand six-year treatment. conclusions the ebp trial is one of few ongoing longterm fire ecology projects in africa and provides a basis for valuable research. the trial has several inferential complications. statistically, the most serious is the lack of full randomisation; other complicating factors are herbivory and soil variability between plots. while the lack of randomisation should be kept in mind, there seems no practical alternative to treating the trial as a fully randomised block design. as argued by trollope et al. (1998) the trial should be interpreted as a fire-herbivory interaction study. the results of venter (in prep.) can be used to investigate soil-vegetation relationships under different treatment regimes, while the data of gorman (in prep.) can be used to test for the additional effects of herbivory. the scale of the plots, and hence their representativity of especially scale-dependent processes, is an issue as plot size fires are infrequent in the park and account for only a small proportion of the total area burnt (van wilgen et al. 2000). if the trial is to continue in the long-term, it may be worth considering several adjustments. for example, the closure of waterholes within a certain radius of the ebp replicates would remove some of the variation in herbivory. the idea of erecting herbivore exclosures within specific plots, or of entirely fencing certain plots, could be reexplored. if particular treatments are to be suspended or changed (e.g. as proposed by trollope et al. 1998), and the separate effects of season and frequency of burn are of interest, it is essential that these factors be fully crossed (i.e. a minimum of four treatments are needed, consisting a combination of two seasons by two frequencies). the timing of burning should be related to specific conditions (e.g. after the first spring rains), rather than to specific calendar months, and named accordingly. any modifications should, however, be thoroughly and carefully considered to avoid loss of the long-term value of the trial. acknowledgements beukes enslin explicitly pointed out to the current community of researchers that the trial is not fully randomised. the two reviewers are thanked for their insightful contributions. sandra macfadyen and issn 0075-6458 9 koedoe 46/1 (2003) biggs.qxd 2005/12/09 10:21 page 9 don ntsala are thanked for their assistance in the preparation of the map. references brynard, a.m., r.l. davidson, p. gillard, g. lecatsas & j. leigh. . 1960. calibration of the belt transect method in combretum woodland in the knp. annexure 'c', annual report of the biological department, april 1959 march 1960. report, skukuza, south african national parks. brynard, a.m. 1971. controlled burning in the kruger national park: history and development. proceedings of the annual tall timbers fire ecology conference. tallahasee, tall timbers research station 11: 219-232. crawley, m.j. 1993. glim for ecologists. london: blackwell. enslin, b.w., a.l.f. potgieter, h.c. biggs & r. biggs. 2000. long term effects of fire frequency and season on the woody vegetation dynamics of the sclerocarya birrea/acacia nigrescens savanna of the kruger national park. koedoe 43(1): 27-37. gertenbach, w.p.d. & a.l.f. potgieter. 1975. veldbrandnavorsing in die knoppiesdoringmaroelaveld van die sentrale distrik, nasionale krugerwildtuin. unpublished memorandum, skukuza, south african national parks. gertenbach, w.p.d. 1983. landscapes of the kruger national park. koedoe 26: 9-121. govender, n., a.l.f. potgieter & h.c. biggs. (in press). scientific value of research conducted on the experimental burn plots in the kruger national park. proceedings of the international rangelands conference. 26 july 1 august 2003, durban, south africa. hurlbert, s.h. 1984. pseudoreplication and the design of ecological field experiments. ecological monographs 54(2): 187-211. mentis, m.t. & a.w. bailey. 1990. changing perceptions of fire management in savanna parks. journal of the grassland society southern africa 7(2): 81-85. national parks board. 1953. jaarverslag van bioloog 1951. report, skukuza, south african national parks. national parks board. 1954. jaarverslag van bioloog 1953. item 37, appendix "n" of the agenda of the board meeting held on 25/26 june 1954. report, skukuza, south african national parks. national parks board. 1955b. jaarverslag van bioloog 1954. item 4, appendix "c" of the agenda of a board meeting held on 21/22 november 1955. report, skukuza, south african national parks. national parks board. 1955a. veldbrandproewe. letter from the director of national parks to the warden of the kruger national park informing him of decisions regarding the experimental burning trial taken at a scientific steering committee meeting. reference k.1/14/2(37). typescript, skukuza, south african national parks. national parks board.1956. jaarverslag: biologiese afdeling 1955. item 4, appendix "c" of the board meeting held on 22/23 june 1956. report, skukuza, south african national parks. national parks board. 1960. jaarverslag: biologiese afdeling april 1959 maart 1960. report, skukuza, south african national parks. national parks board.1962. jaarverslag: departement natuurbewaring april 1961 maart 1962. report, skukuza, south african national parks. national parks board.1971. afdeling natuurbewaring jaarverslag 1970 1971. report, skukuza, south african national parks. national parks board.1972. afdeling natuurbewaring jaarverslag 1971 1972. report, skukuza, south african national parks. national parks board. 1973. afdeing natuurbewaring jaarverslag 1972-1973. report, skukuza, south african national parks. national parks board.1 974. afdeling natuurbewaring jaarverslag 1973-1974. report, skukuza, south african national parks. national parks board. 1976. jaarverslag: departement natuurbewaring apr 1975 mar 1976. report, skukuza, south african national parks. national parks board. 1978. department of nature conservation knp: six-monthly report april september 1978. report, skukuza, south african national parks. national parks board. 1981. jaarverslag: departement navorsing en inligting april 1980 maart 1981. report, skukuza, south african national parks. national parks board. 1984. nbp research report 1983 1984. report, skukuza, south african national parks. national parks board. 1988. jaarverslag: afdeling navorsing 1987 1988. report, skukuza, south african national parks. national parks board. 1989. jaarverslag: afdeling navorsing 1988 1989. report, skukuza, south african national parks. national parks board. 1990. jaarverslag: departement natuurbewaring april 1989 maart 1990. report, skukuza, south african national parks. nel, t.g. 1953. verslag oor veldbrand. typescript, skukuza, south african national parks. koedoe 46/1 (2003) 10 issn 0075-6458 biggs.qxd 2005/12/09 10:21 page 10 oelofse, j., b.h. brockett, h.c. biggs & c. ebersohn. 1999. the effect of drought and post-fire grazing on the herbaceous layer of shrubmopane veld on basalt in the kruger national park, south africa. in: eldridge, d. & d. freudenberger (eds.). people and rangelands building the future. proceedings of the vith international rangeland congress, 19-23 july, townsville, australia. quinn, g.p. & m.j. keough. 2002. experimental design and data analysis for biologists. cambridge: cambridge university press. scholes, r.j. & b.h. walker. 1993. an african savanna. synthesis of the nylsvley study. cambridge: cambridge university press. trollope, w.s.w., a.l.f. potgieter, h.c. biggs & l.a. trollope. 1998. report on the experimental burning plots (ebp) trial in the major vegetation types of the kruger national park. unpublished report no. 9/98, skukuza, south african national parks. van der schijff, h.p. 1958. inleidende verslag oor veldbrandnavorsing in die nasionale krugerwildtuin. koedoe 1: 60-93. van wilgen, b.w., h.c. biggs, s.p. o'regan & n. mare. 2000. a fire history of the savanna ecosystems in the kruger national park, south africa between 1941 and 1996. south african journal of science 96: 167-178. van wyk, p. 1975. memorandum: veldbrand in die krugerwildtuin. 14pp. typescript, skukuza, south african national parks. von broembsen, h.h. 1964. report on two visits to the kruger national park. annexure 'b' of the annual report: nature conservation knp nov 1963 oct 1964. report, skukuza, south african national parks. webber, c.n. 1979. the effects of fire on soil-plant ecological relationships in the southern part of the kruger national park: a study in soil geography. m.sc thesis, university of pretoria, pretoria. issn 0075-6458 11 koedoe 46/1 (2003) appendix 1 treatment history and compliance in the ebp trial. for each treatment, the tables reflect the replicates burnt according to schedule in a particular year. where deviations from the schedule occurred, the reasons and specific plots involved are indicated. where treatment reports are missing or incomplete, and probable treatment compliance could be inferred with certainty from the rainfall conditions in the particular year and expert knowledge of the plots in question, the inferred treatments are indicated. for example, treatment reports are consistently missing for all the major drought years (1964-65, 1983, 1992-94), and it was assumed that no treatments were applied in these years. biggs.qxd 2005/12/09 10:21 page 11 koedoe 46/1 (2003) 12 issn 0075-6458 appendix 1 (continued) sourveld (pretoriuskop) landscape (f) fayi, (k) kambeni, (n) numbi, (s) shambeni shaded cells indicate planned treatment schedule. brackets indicate starting year of a modified treatment schedule. (√) plots on all four replicates burnt according to schedule. where treatments could not be applied on certain replicates due to sparse vegetation or overly moist conditions, the specific plots not burnt in the particular year are indicated (remaining plots burnt according to schedule) (#) accidental burn on indicated plot (?) reports missing. biggs.qxd 2005/12/09 10:21 page 12 issn 0075-6458 13 koedoe 46/1 (2003) appendix 1 (continued) combretum (skukuza) landscape (m) mbyameti, (n) napi, (s) skukuza, (w) n'waswitshaka shaded cells indicate planned treatment schedule. brackets indicate starting year of a modified treatment schedule. (√) plots on all four replicates burnt according to schedule. where treatments could not be applied on certain replicates due to sparse vegetation or overly moist conditions, the specific plots not burnt in the particular year are indicated (remaining plots burnt according to schedule) (#) accidental burn on indicated plot (?) reports missing. biggs.qxd 2005/12/09 10:21 page 13 koedoe 46/1 (2003) 14 issn 0075-6458 appendix 1 (continued) knobthorn / marula (satara) landscape (l) lindanda, (m) marheya, (n) n'wanetsi, (s) satara shaded cells indicate planned treatment schedule. brackets indicate starting year of a modified treatment schedule. (√) plots on all four replicates in the landscape burnt according to schedule. where treatments could not be applied to certain replicates due to sparse vegetation or overly moist conditions, the specific plots not burnt in the particular year are indicated (remaining plots burnt according to schedule) (#) accidental burn on indicated plot (?) reports missing biggs.qxd 2005/12/09 10:22 page 14 issn 0075-6458 15 koedoe 46/1 (2003) appendix 1 (continued) mopani landscape (d) dzombo, (m) mooiplaas, (n) n'shawu, (t) tsendi shaded cells indicate planned treatment schedule. brackets indicate starting year of a modified treatment schedule. (√) plots on all four replicates in the landscape burnt according to schedule. where treatments could not be applied to certain replicates due to sparse vegetation or overly moist conditions, the specific plots not burnt in the particular year are indicated (remaining plots burnt according to schedule) (#) accidental burn on indicated plot (?) reports missing biggs.qxd 2005/12/09 10:22 page 15 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <feff00560065007200770065006e00640065006e0020005300690065002000640069006500730065002000450069006e007300740065006c006c0075006e00670065006e0020007a0075006d002000450072007300740065006c006c0065006e00200076006f006e0020005000440046002d0044006f006b0075006d0065006e00740065006e0020006d00690074002000650069006e006500720020006800f60068006500720065006e002000420069006c0064006100750066006c00f600730075006e0067002c00200075006d002000650069006e00650020007100750061006c00690074006100740069007600200068006f006300680077006500720074006900670065002000410075007300670061006200650020006600fc0072002000640069006500200044007200750063006b0076006f0072007300740075006600650020007a0075002000650072007a00690065006c0065006e002e00200044006900650020005000440046002d0044006f006b0075006d0065006e007400650020006b00f6006e006e0065006e0020006d006900740020004100630072006f0062006100740020006f0064006500720020006d00690074002000640065006d002000520065006100640065007200200035002e003000200075006e00640020006800f600680065007200200067006500f600660066006e00650074002000770065007200640065006e002e00200042006500690020006400690065007300650072002000450069006e007300740065006c006c0075006e00670020006900730074002000650069006e00650020005300630068007200690066007400650069006e00620065007400740075006e00670020006500720066006f0072006400650072006c006900630068002e> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice abstract introduction research method and design results discussion conclusion acknowledgements references appendix 1 about the author(s) christiaan w. winterbach tau consultants (pty) ltd, maun, botswana centre for wildlife management, university of pretoria, south africa glyn maude kalahari research and conservation, maun, botswana gosiame neo-mahupeleng botswana university of agriculture & natural resources, gaborone, botswana rebecca klein cheetah conservation botswana, gaborone, botswana lorraine boast cheetah conservation botswana, gaborone, botswana lindsey n. rich department of fish and wildlife conservation, virginia tech, united states michael j. somers centre for wildlife management, university of pretoria, south africa centre for invasion biology, university of pretoria, south africa citation winterbach, c.w., maude, g., neo-mahupeleng, g., klein, r., boast, l., rich, l.n. et al., 2017, ‘conservation implications of brown hyaena (parahyaena brunnea) population densities and distribution across landscapes in botswana’, koedoe 59(2), a1441. https://doi.org/10.4102/koedoe.v59i2.1441 original research conservation implications of brown hyaena (parahyaena brunnea) population densities and distribution across landscapes in botswana christiaan w. winterbach, glyn maude, gosiame neo-mahupeleng, rebecca klein, lorraine boast, lindsey n. rich, michael j. somers received: 13 sept. 2016; accepted: 05 dec. 2016; published: 23 may 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the brown hyaena (parahyaena brunnea) is endemic to southern africa. the largest population of this near-threatened species occurs in botswana, but limited data were available to assess distribution and density. our objectives were to use a stratified approach to collate available data and to collect more data to assess brown hyaena distribution and density across land uses in botswana. we conducted surveys using track counts, camera traps and questionnaires and collated our results and available data to estimate the brown hyaena population based on the stratification of botswana for large carnivores. brown hyaenas occur over 533 050 km² (92%) of botswana. our density estimates ranged from 0 brown hyaenas/100 km² in strata of northern botswana to 2.94 (2.16–3.71) brown hyaenas/100 km² in the southern stratum of the central kalahari game reserve. we made assumptions regarding densities in strata that lacked data, using the best references available. we estimated the brown hyaena population in botswana as 4642 (3133–5993) animals, with 6.8% of the population in the northern conservation zone, 73.1% in the southern conservation zone, 2.0% in the smaller conservation zones and 18.1% in the agricultural zones. the similar densities of brown hyaenas in the central kalahari game reserve and the ghanzi farms highlight the potential of agricultural areas in botswana to conserve this species. the conservation of brown hyaenas in the agricultural landscape of botswana is critical for the long-term conservation of the species; these areas provide important links between populations in south africa, namibia and zimbabwe. conservation implications: botswana contains the core of the brown hyaena population in southern africa, and conflict mitigation on agricultural land is crucial to maintaining connectivity among the range countries. introduction the brown hyaena (parahyaena brunnea) has an estimated population size of < 10 000 mature individuals and is therefore listed as near threatened on the international union for conservation of nature (iucn) red list (wiesel 2015). the species is endemic to southern africa with range countries including botswana, namibia, south africa, angola and zimbabwe (mills & hofer 1998; wiesel 2015). the largest population of brown hyaena is found in botswana (wiesel 2015), a country that hosts one of the most diverse carnivore assemblages in africa. the large carnivore guild includes lion (panthera leo) (iucn/ssc 2006), cheetah (acinonyx jubatus) and wild dog (lycaon pictus) (iucn/ssc 2007) and significant populations of spotted hyaena (crocuta crocuta) and leopard (panthera pardus) (jacobson et al. 2016). the brown hyaena occurs over most of botswana, except the okavango delta and sections in the north (wiesel 2015). the range includes a diversity of human land uses such as conservation areas (keeping 2014; maude & mills 2005), commercial farms (boast & houser 2012; kent & hill 2013) and communal land used for subsistence livestock farming (muir 2009; schiess-meier et al. 2007). the large carnivore guild in botswana comprises a strong, interspecific dominance hierarchy including subordinate competitors (cheetahs, african wild dogs and brown hyaenas), dominant competitors (lions and spotted hyaenas) and leopards (i.e. the large carnivore least affected by interspecific competition) (marker & dickman 2005; mills 2015). the interactions among carnivore guild members as well as their interactions with prey species are important components of biodiversity (dalerum et al. 2008; mills 2005). conservation of the intact carnivore guild has a higher priority than the conservation of individual species (woodroffe & ginsberg 2005). to conserve the intact guild, a mosaic of high and low densities of the dominant competitors is required to provide refuge areas for the subordinate competitors (winterbach et al. 2013). botswana has two large conservation zones with the potential to conserve the intact carnivore guild (winterbach, winterbach & somers 2014). the southern conservation zone is characterised by a mosaic of low and medium densities of wild prey for lions and spotted hyaenas (winterbach et al. 2014), and both carnivores occur at low to medium densities (funston et al. 2001; maude & selebatso 2012; mills 2015; mudongo & dipotso 2010). the northern conservation zone has a mosaic of high, medium and low prey densities for large carnivores, with the highest prey densities in and around the okavango delta and along the kwando/linyanti/chobe river system (winterbach et al. 2014). high densities of lions and spotted hyaenas have been recorded in the okavango delta and low densities in the dry parts of the northern conservation zone (cozzi et al. 2013; winterbach & maude 2015; winterbach & winterbach 2003). in addition to these conservation zones, the long-term conservation of brown hyaenas depends on the agricultural areas in botswana (boast & houser 2012; kent & hill 2013). these areas do not have the potential to conserve the intact carnivore guild (winterbach et al. 2014). they do, however, provide an opportunity to conserve the less dominant species including brown hyaenas, cheetahs, leopards and wild dogs (boast & houser 2012; iucn/ssc 2007; kent & hill 2013; klein 2007; maude & mills 2005; winterbach et al. 2015). conservation of carnivores in the agricultural areas requires mitigation measures (winterbach et al. 2013). although brown hyaenas are mainly scavengers (maude & mills 2005; mills 1990, 2015) and a low level of conflict is expected with livestock owners, human persecution is a significant threat to the species in the range countries (mills & hofer 1998; wiesel 2015). in the north west province of south africa, 40% of livestock owners regard the brown hyaena as a problem animal (thorn 2008), and in namibia, 72% of livestock owners believe that brown hyaenas were responsible for livestock depredations (lindsey et al. 2013). weise et al. (2015) confirmed conflict with brown hyaenas in namibia, and according to farmers, conflict happens especially during peak calving or lambing seasons. although faecal analysis and inspection of food remains at den sites showed cattle were a significant food source in the farming areas of the gauteng and limpopo, discussions with farmers indicated that the killing of cattle by brown hyaena was probably rare and that removing the individual responsible solved the problem (skinner 1976). although translocation can solve the conflict, one should consider brown hyaena sociality as part of the decision process to translocate individuals (weise et al. 2015). some cattle farmers in botswana believe that brown hyaenas kill new born calves when the calves are left hidden while the mothers go foraging (m. bing, pers. comm., n.d.). maude and mills (2005) reported that only 25% of cattle post owners around makgadikgadi pans national park, botswana, believed that brown hyaenas might cause livestock losses. they also found no loss of livestock because of brown hyaena predation around makgadikgadi pans national park in a 5-year period, but did show that carcasses of livestock were an important food source to sustain brown hyaena populations in agricultural areas. furthermore, brown hyaenas were a minority of the problem animal incidents reported in the kweneng district, a livestock area south of the central kalahari game reserve in botswana (schiess-meier et al. 2007). boast (2014) recorded low levels of conflict with brown hyaenas and that they are the most tolerated large carnivore in botswana. the wide distribution of brown hyaenas on both conservation and agricultural land requires density estimates for all ecosystems and land uses in botswana. data to assess the large carnivore populations in botswana are available in peer-reviewed articles and internal and unpublished reports. we reviewed articles and reports to identify some of the areas that were data deficient and conducted large carnivore surveys at multiple sites in botswana. here we present our data on brown hyaenas, collated with other published studies and unpublished or internal reports from the research community in botswana to provide a countrywide population estimate of brown hyaenas in botswana. we discuss the implications of the results for carnivore conservation. research method and design study area the republic of botswana is a landlocked country of 582 000 km2, sharing international boundaries with namibia, south africa, zimbabwe and zambia. altitude ranges from 515 m a.s.l. to 1491 m a.s.l. most of botswana is arid to semi-arid, with the kalahari ecosystem occupying approximately 82% of the country. rainfall is erratic and the mean rainfall ranges from 250 mm per year in the south-west to over 650 mm in the north-east (department of surveys and mapping 2001). over 90% of rainfall occurs in the summer months between november and april. apart from the okavango delta and the perennial kwando/linyanti/chobe river system, the only other surface water occurs in rivers and pans during the rainy season (department of surveys and mapping 2001). the mean minimum temperatures range from 5 °c in july to 19 °c in january with the mean maximum daily temperatures ranging from 22 °c in july to 33 °c in january (department of surveys and mapping 2001). vegetation over most of the country is shrub and tree savannah, which is classified as sandveld (department of surveys and mapping 2001). the hardveld vegetation types are associated with hills and rocky outcrops in the eastern part of botswana. the northern conservation zone comprises the wetland of the okavango delta, sandveld, mopane colophospermum mopane dominated vegetation types and limited miombo woodland in the north-east. the okavango delta consists of a mosaic of islands, waterways and seasonal floodplains (department of surveys and mapping 2001). seventeen percent of botswana is fully protected as designated national parks and game reserves, and a further 21% is partially protected as designated wildlife management areas (wmas). only 5% of the country is designated as urban. the balance of 57% comprises uncultivated rangeland consisting of approximately 5% freehold land, 25% state land and 70% tribal/communal grazing land (department of surveys and mapping 2001). commercial livestock production occurs on freehold, state and tribal lands. winterbach et al. (2014) identified four conservation zones and four agricultural zones in botswana based on primary land use. the conservation zones consist of the large northern conservation zone and southern conservation zone, and the two smaller conservation zones of xaixai and tuli. the northern conservation zone includes chobe national park, moremi game reserve, nxai pan national park, makgadikgadi pans national park and the associated wmas. the central kalahari game reserve, kgalagadi transfrontier park and associated wmas form the southern conservation zone (figure 1). the agricultural zones are ngami, central, ghanzi and kgalagadi (figure 1). figure 1: primary land use in botswana, including the conservation zones (national parks, game reserves and wildlife management areas) and agricultural zones (commercial farms and communal land with cattle posts). insert shows botswana in relation to southern africa. livestock (mainly cattle) rearing forms 70% – 80% of the contribution of the agricultural sector to the gross domestic product (botswana ministry of agriculture 2011). the national herd was 2.6 million cattle, 1.8 million goats and 300 000 sheep in 2012. traditional cattle posts on communal grazing land are the most common livestock production system (botswana ministry of agriculture 2011). there were 109 ranches producing game or cattle and game in 2013; these cover approximately 11 500 km2 (boast 2014). approximately half of the population of 2 million people live in rural villages and small settlements and are thus partially or fully dependent on livestock for their livelihoods (central statistics office 2014). procedure we conducted track counts following the methodology of stander (1998) and funston et al. (2010). existing roads and 4 × 4 trails were used for transects, covering parts of the northern conservation zone and the central kalahari game reserve in the southern conservation zone (figure 2). two trackers identified the tracks from a vehicle driven at slow speeds not exceeding 15 km/h. all fresh tracks (< 24 h old) of large carnivores were identified and recorded with the date, gps location, species and number of individual animals. data recording excluded roads disturbed by vehicles or rain in the previous 24 h. each individual should only be recorded once per day on the survey (stander 1998). multiple track incidences (observations of tracks) from the same species on the same transect were reviewed with the trackers to determine whether the tracks belonged to the same or new individuals. figure 2: stratification of botswana and locations of track surveys and camera trap survey (detail on inset) conducted between 2005 and 2015 to estimate brown hyaena (parahyaena brunnea) densities. we deployed panthera v4 incandescent-flash and bushnell trophycam infra-red camera traps at 221 locations across a 550 km2 study area in the northern conservation zone and ngami agricultural zone between february and july 2015. we used 5 km2 grid cells to guide the placement of cameras and ensure systematic coverage of the entire study area. we deployed two camera stations within each grid cell, one on the road closest to the predetermined centre point of each grid cell and the second on the road closest to a predetermined random point within each grid cell. we placed all cameras on sand roads to increase our probability of photographing carnivores given that large predators and carrion feeders often use lightly travelled roads as movement corridors (forman & alexander 1998). if cameras had been placed randomly or on the extensive network of game trails, we believe our detection rates would have been prohibitively low. each camera station included two opposing cameras mounted on trees, offset by 0.5 m – 1 m. if there were no trees available, we mounted cameras on metal fence posts hammered into the ground. we secured cameras at knee height and positioned cameras to photograph flanks of passing animals. we programmed cameras to take three photos at each trigger event in the daytime with a delay of 30 s between trigger events. at night-time, the infra-red cameras took three photos when triggered but the incandescent-flash cameras could only take one photo every 15 s because of the flash having to recharge. for each station, we combined information from the two opposing cameras using the time or date stamps on the photographs. we used a rotational system for camera deployment. we divided our study area into five sub-areas of approximately 110 km2 each and sequentially sampled each area for 30 nights. we deployed an average of 44 camera stations (i.e. 88 cameras) within each sub-area. we checked cameras every 5–10 days to download photos, replace batteries and ensure cameras were still operational. distribution records of brown hyaenas consisted of track observations we recorded during our surveys, supplemented with brown hyaena observation records from publications, reports and verifiable observations between january 2005 and april 2016. brown hyaena status, collected as part of a larger questionnaire survey conducted during 2012 and 2013, was recorded as present (visual sightings or tracks seen at least quarterly), transient (visual sightings or tracks seen less frequently than quarterly) or absent (never seen brown hyaena or its tracks). klein (2013) surveyed cattle posts and commercial farms in the kalahari region of botswana, and boast (2014) targeted primarily game ranchers and commercial livestock farmers in the game ranching regions of the central, ghanzi, ngamiland and north east regional districts. data analysis transects were pooled per stratum for analysis, following the landscape stratification of botswana for large carnivores (figure 2) from winterbach et al. (2015). the ratio of stratum size and transect length was used to calculate penetration (km²/km of transect) as an index of sampling effort (funston et al. 2010). we calculated track density (number of individual tracks/100 km sampled) per transect, treating each repetition an individual record. following funston et al. (2010), we calculated the mean and standard deviation of the distance (km) between track incidences for each stratum sampled. funston et al. (2010) recommended that the coefficient of variance (cv) of distance between track incidences (standard deviation × 100/mean km per track incidence) should be less than 20% to ensure appropriate precision of the density estimate. this generally occurs after 19–30 track incidences (funston et al. 2010). we provide penetration, distance between track incidences with the cv and number of track incidences as indicators of the quality of density estimates in each stratum. we estimated brown hyaena densities from track densities using the extended carnivore model for sandy substrates formula ‘animal density = track density/3.26’ from winterbach et al. (2016). we included density data from other studies for the national population estimate (table 1-a1). we sourced articles and reports from the research community in botswana. we recalculated the density estimates from other track surveys using the extended carnivore model for sandy substrates (winterbach et al. 2016) for consistency of density calculation methods across studies. the population estimates were calculated per stratum from the density estimates, size of the stratum and the proportion of that stratum included in the brown hyaena range (stratum population = density × stratum size/100 km2). no density estimates were available for some strata. where we deemed strata to be similar (general habitat and general land use) to areas with reference data, we assumed similar population densities. in all other cases for strata lacking suitable reference data, we followed the same approach as the namibia large carnivore atlas (hanssen & stander 2004; stein et al. 2012) using standardised density categories of low, medium or high: assumed low density: 0.085 brown hyaenas/100 km² (range 0.07–0.10). assumed medium density: 0.125 brown hyaenas/100 km² (range 0.10–0.15). assumed high density: 0.575 brown hyaenas/100 km² (range 0.15–1.0). because of the limited number of brown hyaenas recorded during the camera trap survey we could not calculate density. the camera trap records were used to indicate the presence and absence as part of mapping distribution. locations from our surveys and questionnaires were plotted with other reliable location records collected from publications, reports and personal communications. we used a chi-square test with bonferroni simultaneous confidence intervals (byers, steinhorst & krausman 1984) to compare frequencies that respondents reported brown hyaenas as present or transient or absent on commercial livestock farms and cattle posts compared to game farms, whose frequencies were defined as the expected values. results density estimates we present data from track surveys we conducted between 2005 and 2007 (chobe district and okavango delta) and between 2011 and 2013 (ng43, makgadikgadi pans national park and the central kalahari game reserve) (table 1). the lack of roads and 4 × 4 trails limited access to large parts of the study area and resulted in a low sampling effort in some strata as measured by the penetration index (table 1). penetrations ranged from 8.5 km²/km to 118.5 km²/km of transect. the surveys conducted during 2005 and 2006 totalled 3000 km of north-eastern botswana (figure 2). we completed 844 km in ng43 (wmas near the okavango delta) during 2011, partially covering stratum 1.3.2 (figure 2). as part of long-term monitoring, we repeated surveys in ng43 during 2012 and 2013 (table 1). during 2012, we completed the following transects: a total of 482 km in makgadikgadi pans national park (stratum 1.5.3), 1120 km in the northern central kalahari game reserve (stratum 2.1.1), 1022 km in southern and eastern central kalahari game reserve (strata 2.1.2 and 2.1.3) and 397 km in khutse game reserve (stratum 2.1.4). table 1: summary of track surveys completed between 2005 and 2013 to estimate brown hyaena (parahyaena brunnea) densities in study areas across botswana. the track incidences recorded in makgadikgadi pans national park and the central kalahari game reserve ranged from 17 to 87 per stratum with mean distance between track incidences of 11.76 km – 22.47 km (table 2). the cv was between 102% and 143%, exceeding the 20% guideline (funston et al. 2010). although we completed 5550 km of track surveys north of makgadikgadi pans national park, we only recorded tracks of brown hyaenas on five occasions. we could not calculate cv for the track frequency in this area because of the small sample size. the density estimates were between 0.00 animals/100 km² and 0.14 animals/100 km². the southern central kalahari game reserve had the highest density of brown hyaenas (2.94 animals/100 km²), followed by 2.55 animals/100 km² in makgadikgadi pans national park (table 2). table 2: results of track surveys completed between 2005 and 2013 to estimate brown hyaena (parahyaena brunnea) densities in study areas across botswana. the camera trap survey yielded only five records of brown hyaenas from 11 618 trap nights. these data were not sufficient to estimate density, but provided confirmation of the edge of the brown hyaena range in part of northern botswana. distribution brown hyaenas occur over 533 050 km² (92%) of botswana (figure 3). previous studies recorded the presence of brown hyaenas in the kgalagadi transfrontier park and surrounding wmas (funston et al. 2001; keeping 2014; mills 1990, mudongo & dipotso 2010) and ghanzi (boast & houser 2012; kent 2011; kent & hill 2013). these studies and the location data from our track surveys, camera trap survey, questionnaire survey and additional observation records are shown in figure 3. only a few records of brown hyaenas were recorded north of makgadikgadi pans national park despite 5550 km of track surveys completed. the concessionaire reported that they only saw brown hyaena tracks in the southern half of ng43 during 15 years of operation (1999–2013) (j. van rensburg, kgori safaris, pers. comm., n.d.). the results of the camera trap survey support that this was the northern edge of the brown hyaena range as brown hyaena were only recorded at the southern stations of the camera survey. figure 3: brown hyaena (parahyaena brunnea) distribution range in botswana based on occurrences recorded between 2005 and 2016. eighty-two of 418 respondents did not record brown hyaenas as present, transient or absent and were excluded from further analysis. the questionnaire survey provided presence records of brown hyaenas in all four agricultural zones (figure 3). respondents recorded brown hyaenas as present (91.6%) and absent (8.4%) on game ranches (n = 107). presence of brown hyaena was 85.5% on commercial farms (n = 55) and 53.4% on cattle posts (n = 174). the observed frequency of presence and absence or transience differed (χ² = 331.25, df = 3, p < 0.001). brown hyaenas were reported absent or transient more frequently and present less frequently on cattle posts than on game ranches ( p = 0.001). it did not differ significantly between commercial livestock farms and game ranches (table 3). table 3: bonferroni simultaneous confidence intervals for the presence and absence ortransience of brown hyaena (parahyaena brunnea) based on questionnaires (2008–2009 and 2011–2012) completed by farmers on game ranches (n = 107), commercial livestock farms (n = 55) and traditional cattle posts (n = 174) in the agricultural zones of botswana. population estimates our data and data from other sources that we used for the population estimates are summarised in table 1-a1. large parts of the northern conservation zone are outside brown hyaena range or are part of the northern limit of their range. we calculated three density estimates north of makgadikgadi pans national park, all between 0.04 animals/100 km² and 0.14 animals/100 km². the five track records obtained along the 5550 km track survey were too low for robust density estimates and may indicate a patchy distribution towards the northern limit of the brown hyaena range. we therefore assumed (table 1-a1) that brown hyaenas occur at extremely low densities north of makgadikgadi pans national park (range 0.0 animals/100 km² – 0.1 animals/100 km² and mean 0.05 animals/100 km²). recorded densities (table 1-a1) varied between 1.13 animals/100 km² and 3.90 animals/100 km² in the southern conservation zone (funston et al. 2001; keeping 2014, maude & selebatso 2012, mills 1990; mudongo & dipotso 2010). brown hyaena densities were estimated between 1.89 animals/100 km² and 3.10 animals/100 km² in the ghanzi farms (boast & houser 2012; kent 2011; kent & hill 2013). no density data were available for the central agricultural zone (north-east botswana) and kgalagadi agricultural zone (south-east botswana). these two zones border on the north west province in south africa where thorn et al. (2011) estimated overall density of brown hyaenas in agricultural land as 0.15/100 km² ± se 0.08. based on this, we assumed a medium density with 0.125 brown hyaenas/100 km², ranging from 0.1 animals/100 km² to 0.15 animals/100 km² in these two agricultural zones (table 1-a1). for substrata 6.5.0 (ngamiland) and 7.2.0 (ghanzi), we assumed high density (0.575 brown hyaenas/100 km² [range 0.15–1.00]) based on the high densities recorded in neighbouring areas (table 1-a1). the low-density assumption (0.085 brown hyaenas/100 km² [range 0.07–0.10]) was applied to the substrata of ngami agricultural zone, as this is part of the northern edge of brown hyaena distribution (table 4). table 4: population estimates per stratum of brown hyaenas (parahyaena brunnea) in botswana based on surveys conducted between 2005 and 2015. we estimated the brown hyaena population of botswana as 4642 animals (3133–5993) (table 4). seventy-three percent of the brown hyaena population (approximately 3393 animals) occurred within the southern conservation zone (table 4) and 18.1% in the agricultural zones (843 brown hyaenas). the remaining 6.8% occurred in the northern conservation zone and 2% in conservation zones of xaixai and tuli. the estimated and assumed densities are shown in figure 4. figure 4: estimated and assumed densities per stratum of brown hyaenas (parahyaena brunnea) in botswana based on surveys conducted between 2005 and 2015. discussion our study combined information from track surveys, camera trap surveys, questionnaires and previous studies to estimate local densities and distribution of brown hyaenas across botswana. information on brown hyaena populations is vital to making informed conservation decisions and to mitigating population declines, particularly as the species is listed as near threatened (wiesel 2015). because of the large scale of this study, however, our data had several limitations. the data collection span over a decade of research, and there is a risk that carnivore densities may have changed because of ecological factors or changes in land use as demonstrated in zimbabwe (williams et al. 2016). no large-scale changes in land use occurred in botswana during the past 15 years. the brown hyaena density estimates in kgalagadi transfrontier park area of mills (1990), funston et al. (2001) and keeping (2014) were very similar despite covering more than 20 years. the long-term study of maude (2010) found no population declines in the makgadikgadi region. despite this long timespan, our data sets are the best indication of the national population of brown hyaena in botswana. the recommended sampling effort of 19–30 recorded track incidences per sampling unit should result in a cv below 20% for mean distance between track incidences (funston et al. 2010; kent 2011). despite us recording track incidences of 37, 53 and 87 in three strata of the central kalahari game reserve, the cvs of these strata still exceeded 100%. this contradicts the results from localised surveys in the ghanzi farms area (boast & houser 2012; kent 2011) that achieved cvs of less than 20%. these studies recorded densities varying between 1.89 brown hyaenas/100 km² and 3.10 brown hyaenas/100 km² on small units within the stratum 7.1.0 ghanzi farms. similarly, brown hyaena densities varied among habitats within stratum 2.2.1, the kgalagadi transfrontier park (1.13 brown hyaenas/ 100 km² – 2.17 brown hyaenas/100 km²) (funston et al. 2001) and among locations in the surrounding wmas (2.36 brown hyaenas/100 km² – 3.90 brown hyaenas/100 km²) (mudongo & dipotso 2010). the above mentioned studies show heterogeneity in brown hyaena densities across landscapes in conservation and agricultural zones of botswana on a finer resolution than the stratification we used. therefore, we should expect heterogeneity in brown hyaena densities in the large survey strata that result in high cvs. although our current estimate of 4642 (3133–5993) brown hyaenas is higher than the previous estimates of 3900 (3500–4500) (mills & hofer 1998) and 2636 (1990–3282) (winterbach & winterbach 2003), it probably does not reflect a population increase. rather, it indicates that the population was previously underestimated because of a lack of comprehensive data. although there is evidence that brown hyaenas are persecuted in botswana (maude & mills 2005), we found no indication of a population decline. this is supported by a long-term study on brown hyaenas in the makgadikgadi region of botswana by maude (2010). he concluded that brown hyaenas are sufficiently adaptable to live alongside people in agricultural areas and are thus not vulnerable to significant population declines under current land uses (maude 2010). the similar brown hyaena densities recorded in ghanzi and the southern conservation zone (table 1-a1) highlight the importance of agricultural zones in botswana for brown hyaena conservation (boast & houser 2012; kent & hill 2013). furthermore, maude and mills (2005) showed that brown hyaenas occurred at higher densities in the communal land in the agricultural zone around makgadikgadi nxai pan national park, where home range sizes are smaller and clan numbers are higher than inside the national park. in contrast to these findings, thorn et al. (2011) estimated a much lower density of brown hyaenas in agricultural land (0.15/100 km² ± se 0.08) than in pilanesberg national park, north west province of south africa. less antagonism, lower human density, a lack of large-scale crop production and differences in livestock management practices may contribute to a more hospitable environment for brown hyaenas in ghanzi than the north west province (kent & hill 2013). the findings of maude and mills (2005), schiess-meier et al. (2007) and kent and hill (2013) indicated that we can potentially have relatively high brown hyaena densities in the ngami, central and kgalagadi agricultural zones. we lacked density data in these areas that include large parts of communal land with cattle posts. brown hyaena densities in botswana, apart from the north, varied between 1.2 animals/100 km² and 3.9 animals/100 km² (table 1-a1). however, our questionnaire surveys showed that brown hyaenas were more likely to be reported as transient or absent on the communal land with cattle posts than on commercial and game farms. this can be the result of observer bias, with the respondents either overor under-reporting the presence of brown hyaena. the alternative is that brown hyaena did occur less on communal land. our current data are not suitable to clarify this. the brown hyaena densities recorded elsewhere in botswana might not be representative of these agricultural zones, and therefore, we opted for the conservative density assumption of 0.15 animals/100 km² based on the results of thorn et al. (2011). the communal farmlands with cattle posts in these three agricultural zones represent a major gap in our knowledge of brown hyaenas in botswana. brown hyaenas can thrive in agricultural areas of botswana (boast & houser 2012; kent & hill 2013; maude & mills 2005). therefore, we recommend that future brown hyaena surveys focus particularly on the ngami, central and kgalagadi agricultural zones, which may be strongholds for the species. a significant proportion (18.1%) of the brown hyaena population in botswana occurred in agricultural areas of botswana. for example, our estimates showed more brown hyaenas in the ghanzi farms (stratum 7.1.0) than all of the northern conservation zone. the agricultural zones are not only key areas for the conservation of brown hyaenas in botswana but also essential to maintain links with populations in namibia and south africa. the stronghold for brown hyaenas in the northern conservation zone is the makgadikgadi pans. this area and the strata in the northern conservation zone without perennial water have low prey densities and should be refuge areas for subordinate carnivores (winterbach et al. 2014). however, we recorded brown hyaena infrequently in the strata north of makgadikgadi, despite low densities of lion and spotted hyaena (unpublished data from our surveys). annual rainfall increases from 400 mm in makgadikgadi to 700 mm in the north-east of the northern conservation zone (department of surveys and mapping 2001). this may be a factor that limits brown hyaena in the northern conservation zone directly or indirectly, because brown hyaena occurs in areas with rainfall up to approximately 700 mm (wiesel 2015). also, the species does not occur in the okavango delta, the area around moremi game reserve, an area that supports high densities of lions and spotted hyaenas (cozzi et al. 2013; winterbach & winterbach 2003). mills and mills (1982) and mills (2015) found that spotted hyaenas out-compete brown hyaenas and this interspecific competition may explain why the okavango delta is not part of the brown hyaena’s range (mills & hofer 1998; wiesel 2015). although it is not clear from our distribution and density data whether brown hyaenas are resident in localised areas or merely transient through the northern part of their range in botswana, the northern conservation zone provides the link between the botswana and zimbabwe populations. botswana supports the highest number of brown hyaenas of all the range countries (wiesel 2015), and a significant proportion of this national population occurs on agricultural land. the focus of carnivore conservation in botswana should be on maintaining the intact large carnivore guild in the conservation zones, complimented by a species conservation approach in the agricultural zones. we recommend that future surveys in botswana specifically include density estimates for brown hyaenas on communal farmland in the ngami, central and kgalagadi agricultural zones. less livestock owners in botswana regard brown hyaenas as a risk to livestock than their counterparts in south africa and namibia (lindsey et al. 2013; maude & mills 2005; thorn 2008), which may be related to farming practices, that can increase the risk of brown hyaenas killing new born and weak calves (m. bing, pers. comm., n.d.). regional and individual differences in behaviour of brown hyaenas may contribute to the lower perceived threat in parts of botswana. although brown hyaenas may on rare occasions cause livestock losses (skinner 1976; weise et al. 2015), they are not a significant problem animal species. persecution of brown hyaenas in the livestock areas of botswana may impact negatively on the long-term conservation of the species across its distribution range in southern africa. the resident brown hyaena population in the livestock areas of botswana is not only a significantly large population, but is also critical to maintain links among the conservation zones in botswana and the neighbouring range countries namibia, south africa and zimbabwe. the information provided here should be incorporated into conservation strategies for large carnivores in botswana. conclusion we found no indication that the estimated population of 4642 (3133–5993) brown hyaenas in botswana declined. brown hyaena densities were heterogeneous across landscapes in conservation and agricultural zones on a finer resolution than the stratification we used. we lack data to assess the population in large parts of the agricultural areas and recommend that future brown hyaena surveys focus particularly on the ngami, central and kgalagadi agricultural zones, which may be strongholds for the species. the agricultural areas in botswana are important to maintain population links between namibia, south africa and zimbabwe. acknowledgements south africa’s national research foundation provided financial assistance for c.w.w. (grant no: 95399). kgori safaris (pty) ltd funded data collection in ng43. the african wildlife foundation’s large carnivore research project funded the surveys in the chobe district. the authors thank panthera for providing the panthera v4 camera traps used in this study, ron crous for allowing them to survey his property, and the botswana predator conservation trust for providing support for the camera trap survey. kanabo conservation link and denver zoo helped to fund the central kalahari game reserve survey. cheetah conservation botswana, leopard ecology and conservation and anton van schalkwyk participated in the central kalahari game reserve survey. the authors would like to acknowledge the expert skills of all the trackers involved in the spoor surveys, without whom data collection for this study would not have been possible. opinions expressed and conclusions arrived at in the article are those of the authors and are not necessarily to be attributed to the funders. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions c.w.w. was the project leader. c.w.w., g.n-m. and g.m. designed and conducted the track surveys. l.n.r. designed and conducted the camera trap survey. r.k. and l.b. provided the brown hyaena results from their questionnaire surveys. c.w.w. performed the data analysis. m.j.s. made conceptual contributions. all authors contributed to the writing and editing of the manuscript. references boast, l.k., 2014, ‘exploring the causes of and mitigation options for human-predator conflict on game ranches in botswana: how is coexistence possible?’, phd, university of cape town. boast, l.k. & houser, a., 2012, ‘density of large predators on commercial farmland in ghanzi, botswana’, south african journal of wildlife research 42, 118–143. https://doi.org/10.3957/056.042.0202 botswana ministry of agriculture, 2011, botswana agricultural marketing strategy (2011–2016), government of botswana, gaborone, p. 39, veiwed n.d., from http://www.moa.gov.bw./downloads_agricultural_marketing_strategy.pdf byers, r., steinhorst, r.k. & krausman, p.r., 1984, ‘clarification of a technique for analysis of utilization-availability data’, journal of wildlife management 48, 1050–1053. https://doi.org/10.2307/3801467 central statistics office, 2014, population and housing census 2011 analytical report, viewed 20 march 2014, from http://www.cso.gov.bw/images/analytical_report.pdf cozzi, g., broekhuis, f., mcnutt, j.w. & schmid, b., 2013, ‘density and habitat use of lions and spotted hyenas in northern botswana and the influence of survey and ecological variables on call-in survey estimation’, biodiversity and conservation 22, 2937–2956. https://doi.org/10.1007/s10531-013-0564-7 dalerum, f., somers, m.j., kunkel, k.e. & cameron, e.z., 2008, ‘the potential for large carnivores to act as biodiversity surrogates in southern africa’, biodiversity conservation 17, 2939–2949. https://doi.org/10.1007/s10531-008-9406-4 department of surveys and mapping, 2001, botswana national atlas, arkpressen, västerås, sweden. forman, r.t.t. & alexander, l.e., 1998, ‘roads and their major ecological effects’, annual reveiwof ecology and systematics 29, 207–231. https://doi.org/10.1146/annurev.ecolsys.29.1.207 funston, p.j., frank, l., stephens, t., davidson, z., loveridge, a., macdonald, d.m. et al., 2010, ‘substrate and species constraints on the use of track incidences to estimate african large carnivore abundance’, journal of zoology 281, 56–65. https://doi.org/10.1111/j.1469-7998.2009.00682.x funston, p.j., herrmann, e., babupi, p., kruiper, a., kruiper, h., jaggers, h. et al., 2001, ‘spoor frequency estimates as a method of determining lion and other large mammal densities in the kgalagadi transfrontier park’, in p.j. funston (ed.), kalahari transfrontier lion project, pp. 36–52, endangered wildlife trust, johannesburg. hanssen, l. & stander, p., 2004, namibia large carnivore atlas, predator conservation trust, namibia. iucn/ssc, 2006, regional conservation strategy for the lion panthera leo in eastern and southern africa, iucn, gland, switzerland. iucn/ssc, 2007, regional conservation strategy for the cheetah and african wild dog in southern africa, iucn species survival commission, gland, switzerland. jacobson a.p., gerngross p., lemeris jr j.r., schoonover r.f., anco c., breitenmoser-würsten c., et al., 2016, ‘leopard (panthera pardus) status, distribution, and the research efforts across its range’, peerj 4, e1974. https://doi.org/10.7717/peerj.1974 keeping, d., 2014, ‘rapid assessment of wildlife abundance: estimating animal density with track counts using body mass-day range scaling rules’, animal conservation 17(5), 486–497. https://doi.org/10.1111/acv.12113 kent, v., 2011, ‘the status and conservation potential of carnivores in semi-arid rangelands, botswana’, phd, durham university. kent, v.t. & hill, r.a., 2013, ‘the importance of farmland for the conservation of the brown hyaena parahyaena brunnea’, oryx 47, 431–440. https://doi.org/10.1017/s0030605312001007 klein, r., 2007, ‘status report for cheetah in botswana’, in c. breitenmoser, u. breitenmoser & s. durant (eds.), status and conservation needs of cheetahs in southern africa, pp. 14–21, iucn/cat specialist group, gland, switzerland. klein, r., 2013, ‘an assessment of human carnivore conflict in the kalahari region of botswana’, msc, rhodes university. lindsey, p.a., havemann, c.p., lines, r., palazy, l., price, a.e., retief, t.a. et al., 2013, ‘determinants of persistence and tolerance of carnivores on namibian ranches: implications for conservation on southern african private lands’, plos one 8, e52458. https://doi.org/10.1371/journal.pone.0052458 marker, l.l. & dickman, a.j., 2005, ‘factors affecting leopard (panthera pardus) spatial ecology, with particular reference to namibian farmlands’, south african journal of wildlife research 35, 105–115. maude, g., 2010, ‘the spatial ecology and foraging behaviour of the brown hyaena (hyaena brunnea)’, phd thesis, university of bristol. maude, g. & mills, m.g.l., 2005, ‘the comparative feeding ecology of the brown hyaena in a cattle area and a national park in botswana’, south african journal of wildlife research 35, 201–214. maude, g. & selebatso, m., 2012, first summary of large carnivore estimates obtained from the track survey conducted in the ckgr and khutse game reserves in march 2012. unpublished report. mills, m.g.l., 1990, kalahari hyaenas. comparative behavioural ecology of two species, unwin hyman, london. mills, m.g.l., 2005, ‘large carnivores and biodiversity in african savanna ecosystems’, in j.c. ray, k.h. redford, r.s. steneck & j. berger (eds.), large carnivores and the conservation of biodiversity, pp. 208–229, island press, washington, dc. mills, m.g.l., 2015, ‘living near the edge: a review of the ecological relationships between large carnivores in the arid kalahari’, african journal of wildlife research 45(2), 127–137. https://doi.org/10.3957/056.045.0127 mills, m.g.l. & hofer, h., 1998, hyaenas: status survey and conservation action plan, iucn/ssc hyaena specialist group, gland, switzerland. mills, m.g.l. & mills, m.e.j., 1982, ‘factors affecting the movement patters of brown hyaenas, hyaena brunnea, in the southern kalahari’, south african journal of wildlife research 12, 111–117. mudongo, e. & dipotso, f.m., 2010, predator population estimates in kgalagadi wildlife management areas, through track counts and call up, department of wildlife and national parks, gaborone, botswana. muir, m.j., 2009, ‘predator conservation and conflict in the rangelands of western botswana: evaluating the strategies and conditions that mitigate livestock loss’, phd, university of california. schiess-meier, m., ramsauer, s., gabanapelo, t. & könig, b., 2007, ‘livestock predation – insights from problem animal control registers in botswana’, journal of wildlife management 71, 1267–1274. https://doi.org/10.2193/2006-177 skinner, j.d., 1976, ‘ecology of the brown hyaena hyaena brunnea in the transvaal with a distribution map for southern africa’, south african journal of science 72, 262. stander, p.e., 1998, ‘track counts as indices of large carnivore populations: the relationship between track frequency, sampling effort and true density’, journal of applied ecology 35, 378–385. https://doi.org/10.1046/j.1365-2664.1998.00313.x stein, a.b., aschenborn, o., kastern, m., andreas, m. & thompson, s., 2012, namibia large carnivore atlas, ministry of environment and tourism, windhoek, namibia. thorn, m., 2008, preliminary results from a questionnaire survey regarding predators in the commercial landscapes of the north west province of south africa, university of brighton, brighton, england. thorn, m., green, m., bateman, p.w., waite, s. & scott, d.m., 2011, ‘brown hyaenas on roads: estimating carnivore occupancy and abundance using spatially auto-correlated sign survey replicates’, biological conservation 144, 1799–1807. https://doi.org/10.1016/j.biocon.2011.03.009 weise, f.j., wiesel, i., lemeris, j., jr. & janse van vuuren, r., 2015, ‘evaluation of a conflict-related brown hyaena translocation in central namibia’, african journal of wildlife research 45, 178–186. https://doi.org/10.3957/056.045.0178 wiesel, i., 2015, parahyaena brunnea. the iucn red list of threatened species 2015: e.t10276a82344448, viewed 07 january 2016, from http://www.iucnredlist.org williams, s.t., williams, k.s., joubert, c.j. & hill, r.a., 2016, ‘the impact of land reform on the status of large carnivores in zimbabwe’, peerj 4, e1537. https://doi.org/10.7717/peerj.1537 winterbach, c.w., ferreira, s.m., funston, p.j. & somers, m.j., 2016, ‘simplified large african carnivore density estimators from track indices’, peerj 4, e2662. https://doi.org/10.7717/peerj.2662 winterbach, c.w. & maude, g., 2015, ‘summary of lion population estimates and trends in botswana’, unpublished report, maun, botswana, p. 18. winterbach, c.w. & winterbach, h.e.k., 2003, draft predator management strategy, department of wildlife and national parks, gaborone, botswana. winterbach, h.e.k., winterbach, c.w., boast, l.k., klein, r. & somers, m.j., 2015, ‘relative availability of natural prey versus livestock predicts landscape suitability for cheetahs acinonyx jubatus in botswana’, peerj 3, e1033. https://doi.org/10.7717/peerj.1033 winterbach, h.e.k., winterbach, c.w. & somers, m.j., 2014, ‘landscape suitability in botswana for the conservation of its six large african carnivores’, plos one 9, e100202. https://doi.org/10.1371/journal.pone.0100202 winterbach, h.e.k., winterbach, c.w., somers, m.j. & hayward, m.w., 2013, ‘key factors and related principles in the conservation of large african carnivores’, mammal review 43, 89–110. https://doi.org/10.1111/j.1365-2907.2011.00209.x woodroffe, r. & ginsberg, j.r., 2005, ‘king of the beasts? evidence for guild redundancy among large mammalian carnivores’, in j.c. ray, k.h., redford, r.s. steneck and j. berger (eds.), large carnivores and the conservation of biodiversity, pp. 154–175, island press, washington, dc. appendix 1 table 1-a1: summary of reference data to estimate density of brown hyaenas (parahyaena brunnea) in botswana. table 1-a1:(continues…) summary of reference data to estimate density of brown hyaenas (parahyaena brunnea) in botswana. table 1-a1:(continues…) summary of reference data to estimate density of brown hyaenas (parahyaena brunnea) in botswana. filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 acknowledgements references about the author(s) anthony j. mills department of soil science, faculty of agrisciences, stellenbosch university, stellenbosch, south africa tercia strydom scientific services, south african national parks, skukuza, south africa jessica l. allen oneworld sustainable investments, cape town, south africa julia baum african wildlife economy institute, stellenbosch university, stellenbosch, south africa citation mills, a.j., strydom, t., allen, j.l. & baum, j., 2023, ‘potential geochemical constraints on tree seedlings in northern kruger national park grasslands’, koedoe 65(1), a1773. https://doi.org/10.4102/koedoe.v65i1.1773 short communication potential geochemical constraints on tree seedlings in northern kruger national park grasslands anthony j. mills, tercia strydom, jessica l. allen, julia baum received: 19 apr. 2023; accepted: 27 june 2023; published: 26 july 2023 copyright: © 2023. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. in the northern kruger national park (knp), south africa, there are extensive areas of treeless grassland interspersed within treed savannas (figure 1) (pienaar 1963). to the authors’ knowledge, there have been no investigations into the factors preventing trees from establishing in these grasslands. figure 1: study site location and habitats within the kruger national park (knp), south africa, showing (a) the distribution of treeless grasslands in the northern kruger national park, photographs of (b) grassland and (c) adjacent treed savanna and (d) the location of sampling plots (~100 m2) in grassland (squares) and savanna (triangles). there are many potential factors that could lead to poor recruitment of tree seedlings in these grasslands relative to the savanna matrix. such factors, including for example competition from grass and herbivory pressure (morrison et al. 2019; porensky & veblen 2012; riginos 2009; van auken & bush 1988), warrant further investigation. soil nutrients are likely to interact with herbivory and grass competition in the tree constraint given that sub-optimal availability of any particular nutrient will adversely affect a recently germinated tree seedling’s rate of growth as well as recovery from herbivory (cohn, van auken & bush 1989; kambatuku, cramer & ward 2011; vadigi & ward 2012). consequently, the soil surface of the grasslands and adjacent savannas was sampled as a first step in assessing the potential influence of nutrition on grass competition as well as tree seedling growth immediately after germination. forty composite soil samples (0 cm–2 cm) were taken from an extensive treeless grassland and adjacent savanna north of letaba rest camp in the knp, south africa with permission from the south african national parks. the dominant tree in the savanna is colophospermum mopane. the underlying geology of both vegetation types is letaba basalt. the study did not include excavation of soil pits and consequently the researchers did not classify the soils. however, a study by venter (1990) near the study site found that the two most common soil forms were oakleaf and valsrivier. the oakleaf soil form is a cumulic soil characterised by young, unconsolidated sediment, either colluvial, alluvial or aeolian in origin (fey 2010) with a horizons ~10 cm–20 cm deep (scwg 2018). the valsrivier soil form is a clay-enriched duplex soil (fey 2010) with a horizons ~15 cm–25 cm deep (scwg 2018). each composite sample comprised 8–10 sub-samples taken from 100 cm2 within a plot of 100 m2. the samples in savannas were taken outside of tree canopies. the influence of land use on the geochemistry of the 0 cm–2 cm surface layer of the soils sampled is likely to be insignificant because no agriculture has taken place in the area for at least one century. soil samples were air-dried and sieved to < 2 mm. ph was determined using a 1:2.5 soil:1m kcl solution. inductively coupled plasma mass spectrometry was used to analyse: (1) potassium; calcium; magnesium and sodium (1% citric acid extract) (division of chemical services 1956); (2) phosphorous (1% citric acid extract) (du plessis & burger 1965); (3) sulfur (calcium phosphate extract) (beaton, burns & platou 1968); (4) boron (hot water extract) (bingham 1982); and (5) manganese, copper, zinc (0.02m di-ammonium ethylenediaminetetraacetic acid extract) (beyers & coetzer 1971; trierweiler & lindsay 1969). organic carbon was determined using the walkley-black method (nelson & sommers 1982; walkley 1935). the results showed that relative to the savanna the soil surface in the grassland was enriched in phosphorous, magnesium and zinc, impoverished in boron and more alkaline (figure 2). the only nutrient that emerged from this study as potentially constraining tree seedlings shortly after germination in the grassland was boron, with a mean concentration of 0.4 mg kg−1. assessing whether a concentration of 0.4 mg b kg−1, extracted using hot water from a specific soil, will impede growth in a particular plant species is complex, as multiple soil properties such as ph, clay mineralogy, soil water content and phosphate concentration influence boron uptake by roots (bell 1997; shorrocks 1997). notwithstanding this complexity, on finding such concentrations of b and assuming that the goal (at any given site, regardless of soil properties, climate, or plant species involved) was to optimise plant growth, it would be prudent, based on the existing literature (see e.g., bell 1997; tariq & mott 2007), to experimentally determine whether boron is limiting plant growth. particularly relevant studies in this context include those by li et al. (1978), rashid, rafique and bughio (1994), rafique et al. (2002) and yadav et al. (2017), which found that concentrations of 0.4, 0.5, 0.6 and 0.5 mg b kg−1, respectively, restricted the growth of the plants under investigation. figure 2: the soil chemistry of composite pedoderm samples from grassland (n = 20) and adjacent treed savanna (n = 20) at the study site in the northern kruger national park, south africa. it is noteworthy that boron requirements of dicotyledonous plants, such as the tree seedlings at the study site, are likely to be considerably greater than those of the grass species (bell 1997; tariq & mott 2007). scarcity of boron would consequently be expected to constrain the tree seedlings more than the competing grasses. it is also noteworthy that within fertilisation experiments in the brazilian cerrado, plant growth was constrained by deficiencies of boron more than deficiencies of any other nutrient (lannes et al. 2020), and that a pot experiment in south africa, using a wide range of soils from natural ecosystems, found that boron was the nutrient most strongly correlated with growth of indigenous tree seedlings (wakeling, cramer & bond 2010). pot experiments with a wide range of local grass and tree species are likely to be of value for further investigation into the hypothesis that a relative scarcity of boron in the northern knp grasslands enhances the competitive strength of grasses more than recently germinated tree seedlings. given that nutrient deficiencies in plants are usually identified most effectively using foliar nutrient analyses, the authors suggest that such pot experiments should include nutrient analyses of foliage as well as soil. acknowledgements the authors would like to thank sanparks for permission to take soil samples in the kruger national park, the western cape department of agriculture laboratory at elsenburg for soil chemical analysis, antoni milewski for discussions on concepts presented, section ranger karien keet and game guard desmond siza for support in the field, adelé cormac, selina mochrie, zurelda m. le roux and ruan de wet for technical support, and nicholas salonen and ruan van mazijk for preparation of the manuscript for submission. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions a.j.m. conceived and carried out the study and acquired funding. j.b. and t.s. provided administrative assistance and t.s. facilitated access to the field site where j.b. and a.j.m. conducted field work. j.b. and j.l.a. conducted a literature review for this study. with a.j.m., j.l.a. conducted the statistical analyses. j.l.a. prepared the figures. a.j.m. wrote the manuscript, with all authors approving its contents. ethical considerations ethical clearance to conduct this study in the kruger national park was obtained from south african national parks (permit reference number mila1364). funding information the authors would like to gratefully acknowledge the south african department of environmental affairs, natural resources management programme and the national research foundation (grant number fa2005040700027) for funding this research. data availability the raw data from this study are available at the stellenbosch university institutional research data repository ‘sunscholardata’: https://doi.org/10.25413/sun.23659920. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references beaton, j.d., burns, g.r. & platou, j., 1968, determination of sulphur in soils and plant material, sulphur institute, washington, dc. bell, rw., 1997, ‘diagnosis and prediction of boron deficiency for plant production’, plant and soil 193(1), 149–168. https://doi.org/10.1023/a:1004268110139 beyers, c. p. de l. & coetzer, f.j., 1971, ‘effect of concentration, ph and time on the properties of di-ammonium edta as a multiple soil extractant’, agrochemophysica 3(4), 49–53. bingham, f.t., 1982, ‘boron’, in al (ed.), methods of soil analysis: part 2 chemical and microbiological properties, pp. 431–47, 9, american society of agronomy, soil science society of america, madison, wi. cohn, e.j., van auken, o.w. & bush, j.k., 1989, ‘competitive interactions between cynodon dactylon and acacia smallii seedlings at different nutrient levels’, american midland naturalist 121(2), 265–272. https://doi.org/10.2307/2426030 division of chemical services, 1956, analytical methods, pretoria, south africa. du plessis, r.f. & burger, r.d.t., 1965, ‘a comparison of chemical extraction methods for the evaluation of phosphate availability of top soils’, south african journal of agricultural science 8(4), 1113–1122. fey, m.v., 2010, ‘introduction to the soil groups and the naming of forms and families’, in m.v. fey (ed.), ch. 1 in soils of south africa, p. 11, cambridge university press, cambridge. kambatuku, j.r., cramer, m.d. & ward, d., 2011, ‘savanna tree–grass competition is modified by substrate type and herbivory’, journal of vegetation science 22(2), 225–237. https://doi.org/10.1111/j.1654-1103.2010.01239.x lannes, l.s., venterink, h.o., leite, m.r., silva, j.n. & oberhofer, m., 2020, ‘boron application increases growth of brazilian cerrado grasses’, ecology and evolution 10(13), 6364–6372. https://doi.org/10.1002/ece3.6367 li, b.h., li, w.h., kui, m.c., chao, w.s., jern, h.p., li, c.r. et al., 1978, ‘studies on the cause of sterility of wheat’, journal of the northeastern agriculture college 3(1), 19. morrison, t.a., holdo, r.m., rugemalila, d.m., nzunda, m. & anderson, t.m., 2019, ‘grass competition overwhelms effects of herbivores and precipitation on early tree establishment in serengeti’, journal of ecology 107(1), 216–228. https://doi.org/10.1111/1365-2745.13010 nelson, d.w. & sommers, l.e., 1982, ‘total carbon, organic carbon and organic matter’, in: a.l. page, r.h. miller & d.r. keeney (eds.), methods of soil analysis: chemical and microbiological properties, pp. 539–579, american society of agronomy, soil science society of america, madison, wi. pienaar, u.d.v., 1963, ‘the large mammals of the kruger national park – their distribution and present-day status’, koedoe 6(1), 1–37. https://doi.org/10.4102/koedoe.v6i1.810 porensky, l.m. & veblen, k.e., 2012, ‘grasses and browsers reinforce landscape heterogeneity by excluding trees from ecosystem hotspots’, oecologia 168(3), 749–759. https://doi.org/10.1007/s00442-011-2123-9 rafique, e., rashid, a., bhatti, a.u., rasool, g. & bughio, n., 2002, ‘boron deficiency in cotton grown on calcareous soils of pakistan i. distribution of b availability and comparison of soil testing methods’, in h.e. goldblach, p.h. brown, b. rerkasem, m. thellier, m.a. wimmer & r.w. bell (eds.), boron in plant and animal nutrition, pp. 349–358, kluwer academic/plenum publishers, new york, ny. rashid, a., rafique, e. & bughio, n., 1994, ‘diagnosing boron deficiency in rapeseed and mustard by plant analysis and soil testing’, communications in soil science and plant analysis 25(17&18), 2883–2897. https://doi.org/10.1080/00103629409369232 riginos, c., 2009, ‘grass competition suppresses savanna tree growth across multiple demographic stages’, ecology 90(2), 335–340. https://doi.org/10.1890/08-0462.1 shorrocks, v.m., 1997, ‘the occurrence and correction of boron deficiency’, plant and soil 193(1), 121–148. https://doi.org/10.1023/a:1004216126069 soil classification working group (scwg), 2018, soil classification: a natural and anthropogenic system for south africa, arc-institute for soil, climate and water, pretoria. tariq, m. & mott, c.j.b., 2007, ‘the significance of boron in plant nutrition and environment – a review’, journal of agronomy 6(1), 1–10. https://doi.org/10.3923/ja.2007.1.10 trierweiler, j.f. & lindsay, w.l., 1969, ‘edta-ammonium carbonate soil test for zinc’, soil science society of america journal 33(1), 49–54. https://doi.org/10.2136/sssaj1969.03615995003300010017x vadigi, s. & ward, d., 2012, ‘fire and nutrient gradient effects on the sapling ecology of four acacia species in the presence of grass competition’, plant ecology 213(11), 1793–1802. https://doi.org/10.1007/s11258-012-0134-1 van auken, o.w. & bush, j.k., 1988, ‘competition between schizachyrium scoparium and prosopis glandulosa’, american journal of botany 75(6 part 1), 782–789. https://doi.org/10.1002/j.1537-2197.1988.tb13501.x venter, f.j., 1990, ‘letaba land system’, in a classification of land for management planning in the kruger national park, pp. 277–279. phd thesis. university of south africa. wakeling, j.l., cramer, m.d. & bond, w.j., 2010, ‘is the lack of leguminous savanna trees in grasslands of south africa related to nutritional constraints?’, plant and soil 336, 173–182. https://doi.org/10.1007/s11104-010-0457-4 walkley, a., 1935, ‘an examination of methods for determining organic carbon and nitrogen in soils1. (with one text-figure.)’, the journal of agricultural science 25(4), 598–609. https://doi.org/10.1017/s0021859600019687 yadav, s.n., singh, s.k., kumar, o., latare, a.m. & kumar, m., 2017, ‘evaluation of a suitable extractant and determination of critical limit of boron in soil and mustard (brassica juncea l.) plant by various extractants in inceptisols of varanasi’, communications in soil science and plant analysis 48(10), 1145–1153. https://doi.org/10.1080/00103624.2017.1323103 filelist convert a pdf file! page 1 page 2 page 3 page 4 abstract introduction methods results discussion acknowledgements references appendix 1 about the author(s) timothy f. hall school of animal, plant and environmental sciences, faculty of science, university of the witwatersrand, johannesburg, south africa mary c. scholes school of animal, plant and environmental sciences, faculty of science, university of the witwatersrand, johannesburg, south africa robert j. scholes global change institute, university of the witwatersrand, johannesburg, south africa citation hall, t.f., scholes, m.c. & scholes, r.j., 2022, ‘the pongola bush: tree diversity assessment in a kwazulu-natal forest patch’, koedoe 64(1), a1724. https://doi.org/10.4102/koedoe.v64i1.1724 original research the pongola bush: tree diversity assessment in a kwazulu-natal forest patch timothy f. hall, mary c. scholes, robert j. scholes received: 30 mar. 2022; accepted: 22 aug. 2022; published: 20 sept. 2022 copyright: © 2022. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the pongola bush nature reserve lies in a narrow band along the escarpment between mpumalanga and kwazulu-natal (kzn). although referred to as ‘bush’, this vegetation type may be considered to fit into the intersection of two types of scarp forest: the northern afrotemperate forest and the southern mistbelt forest. the area was heavily logged in the early 1900s with the need for timber for the emerging gold mining industry in barberton. the forest fragment is now protected but lacks formal tree diversity assessments, and this study sets out to establish its community composition and richness using a range of standard techniques. distributed throughout the forest, 127 circular plots (each 80 m2) were surveyed. in these, 1152 stems were measured for species, height, and diameter at breast height (dbh). four species contributed roughly 70% of the 70 mg ha−1 above ground woody biomass. stem size class distributions showed a high recruitment rate of small stemmed individuals and few large individuals. this pattern is consistent with the disturbance history of the site, with limited recruitment of certain species, mainly limited to early successional species. the pongola bush forest is particularly diverse in terms of tree species: 41 species were recorded, and it is estimated from the species area curve that there may be 70 tree species present. conservation implications: this survey is the first formal tree diversity survey conducted on the pongola bush which may assist future research and conservation strategies. keywords: afromontane; forest ecology; fragmentation; tree ecology; above ground biomass. introduction roughly half of the world species are found in forests and they continue to be exploited for their socio-economic value, including the need for industrial timber, medicinal products, and use of productive agricultural land (grainger 2013). this exploitation often leads to the transformation of systems, negatively influencing diversity and ecosystem functionality (falk, palmer & zedler 2013; hooper et al. 2005; johnson et al. 1996). subject to the extent of transformation, ecosystems can regenerate after disturbance events (vieira & scariot 2006). regeneration is dependent on a variety of factors including seed dispersal and seedling establishment, structure and richness of existing species, and resprouting ability (vieira & scariot 2006). identifying natural ecosystem restoration processes in forests is challenging and requires systematic assessment of structural composition (chazdon et al. 2016). forests are scattered along eastern and southern margins of south africa, from the soutpansberg in the north and maputaland in the east to the cape peninsula in the west (mucina & rutherford 2006a). with their patchy distribution within more open systems, limited to fire protected refugia and steep south facing slopes, indigenous forests account for less than 0.5% of the land cover in south africa (berliner 2009; fao 2010; geldenhuys 1994; huntley 1984; rutherford, mucina & powrie 2006). mucina and rutherford (2006a), using a floristic biogeographic classification of the indigenous forests of south africa, recognised 26 forest types, grouped into eight zonal groups and one azonal group. afrotemperate forests stretch from the cape peninsula via the southern cape coast, the eastern cape and kwazulu-natal (kzn) midlands and drakensberg to the north-eastern escarpment and soutpansberg mountains. each of these forest types can have both high and scrub forest depending on climatic conditions and landscape. afromontane forests are generally found on steep south-facing slopes with lowland forests on the coast having a lower canopy. the pongola bush is placed in the northern afrotemperate zone by mucina and rutherford (2006b), but lötter, mucina and witkowski (2014) provided an updated classification of the indigenous forests of the mpumalanga province and recommended that the wakkerstroom midlands forest subtype be embedded within the northern highveld forest type, and not the low escarpment mistbelt forest type as is currently recognised in the national forest classification. it is unclear whether the pongola bush was included in the study by lötter et al. (2014). the pongola bush is believed to be a remnant of ancient forests (perhaps from the tertiary), as evidenced by the presence of onychophora (velvet worms or peripatus), for instance, that flourished in the area (ruhberg & hamer 2005). it does not have a formal inventory of tree species or community structure. this pilot study determined the tree diversity and community composition which would provide a foundation for a more extensive study on changes in the floristic-structural composition along an altitudinal gradient. the diversity and composition was achieved through the documentation of species and species metrics to quantify the richness and relative abundances as well as the total above ground biomass of species within the forest. this study also investigated whether there would be a high dominance of early successional species. methods study site the pongola bush nature reserve covering 8.82 km2 (882 ha) is managed by ezemvelo wildlife (kwazulu-natal nature conservation board). it is situated immediately south of the border of mpumalanga and kzn (figure 1; 27.32349s, 30.47642n). the pongola bush is the largest remaining forest fragment in the area, which historically provided timber for the development of both the barberton and witwatersrand goldfields. sawpits and wagon roads are present in the upper reaches of the forest, suggesting an extensive logging history. much of the remaining extent of the pongola bush is situated within the reserve boundaries: an additional 3 km2 lies outside the borders of the reserve. the forest extends less than 1 km wide in a north-south band, and several kilometres in an east-west direction. the forest lies along the southern edge of an escarpment cliff, extending down a steep slope to grassland on a gentle gradient, marking the southern forest edge. the vegetation is tall (> 10 m) and has a closed, interlocking canopy with multiple layers. the climate in the area is temperate, with an average annual temperature of 14.1 °c, and is characterised by wet summers (november–february) and dry winters (may–august) with an average annual precipitation of 790 mm. figure 1: the sampled locations in the pongola bush, situated on the border of mpumalanga and kwazulu-natal. species data collection transects were walked from the southern edge of the pongola bush patch towards the northern cliff base. approximately every 50 m along each transect that was on average 600 m long (450 m – 800 m), a circular plot with a 5 m radius was surveyed. a total of 127 circular plots were sampled through the forest. all tree stems greater than 20 cm in circumference (diameter ≥ 6.37 cm) at breast height (1.3 m) where the centroid of the stem cross-section is within 5 m of the plot centre were identified, and the height of the top of the crown of that individual was estimated using a ranging rod and clinometer with a precision of about 1 m. statistical analyses species biomass values were estimated using the above ground biomass (agb) model of chave et al. (2014): where ρ is wood density in mg m−3, d is stem diameter at breast height (dbh) in m and h is height in m. wood densities of the species encountered were taken from the african wood density database (carsan et al. 2012), shown in appendix 1 (table 1-a1). species were ranked based on their total above ground biomass (in mg), and statistical significance for biomass measures among and between species was determined using an anova and a post-hoc tukey test at a 95% confidence interval (α = 0.05). stem density (stems ha−1) was determined for all species using count data within the known plot area. stem size-class distributions (in terms of dbh) were determined for the entire sampled population as well as the four most abundant species, plus an economically valuable species (afrocarpus falcatus thunb.). species area relationship analyses were conducted using the ‘sars’ (species-area relationships) software developed by matthews et al. (2019). the species-area model (the increase in species richness recorded as the sampled area increases) was computed using eight random bootstrapped samples from the plot data, which were then averaged. biodiversity and community structure estimates were conducted using paleontological statistical software package (past) 1.81 (hammer, harper & ryan 2001). diversity indices were calculated in two ways: firstly, for number of individuals, which is most appropriate for population dynamics interpretations, and secondly, based on contribution to biomass, which is more important for functionality (mason et al. 2013; schleuter et al. 2010). three diversity indices were calculated: dominance (1-simpson), simpson 1-d and shannon (entropy). community composition analyses using ordination techniques were conducted using a presence-absence matrix to run a non-metric multi-dimensional scaling (nmds). the similarity index input for the nmds was the bray-curtis index, as it is the most valuable metric for showing the compositional dissimilarity of sites using abundance data (gauch 1973). ethical considerations this article followed all ethical standards for research without direct contact with human or animal subjects. results a total of 1152 stems with a circumference greater than 20 cm were sampled, in an aggregated area comprising one ha, from a forest with a total area of about 882 ha. the sampled stems represented 41 tree or tall shrub species (table 1). table 1: the total estimated biomass, stem density and total basal area for the sampled species across 1 ha. the equivalent agb, estimated by allometric equations, is 70 mg ha−1 ± 21 mg ha−1. four canopy-forming species contributed 70% of the agb, namely searsia chirindensis (baker f.), combretum kraussii (hochst.), kiggelaria africana (l.) and celtis africana (burm.f.) of which s. chirindensis contributed to the greatest agb, at roughly 18 mg ha−1 (figure 2). eleven of the species contributed 95% of the agb with over 30 other species accounting for the remaining 5% of the pongola bush tree agb (figure 2). figure 2: total above ground biomass (agb) (mg species−1) of selected species within the pongola bush. there seems to be no clear dominance in terms of stem density within the pongola bush (figure 3). the four species which constituted the greatest biomass are not the four species with the greatest stem densities, but they do appear in the quartile of species with the greatest stem densities. gymnosporia mossambicensis (koltzsch), afrocarpus falcatus, cussonia spicata (thunb.) and diospyros whyteana (hiern) contributed the largest number of stems per unit ground area (figure 3). gymnosporia mossambicensis has the greatest stem density yet it does not fall into the list of species which contribute to the greatest agb, because it is a multi-stemmed tall shrub, rather than a canopy-forming tree. figure 3: stem densities (stems ha−1) of selected species within the pongola bush. the stem size distributions (in dbh) of all individuals of all species, and the stem size distributions for species with the highest biomass and a previously economically important species (a. falcatus) are shown in figure 4. stem diameters, both in aggregate and for some individual species populations, follow a classical ‘reverse j’ distribution – a more-or-less smooth exponential decrease in numbers with increasing stem diameter. this indicates steady recruitment following a period of past disturbance, and a constant stem mortality rate. the populations of s. chirindensis, c. kraussii and k. africana all show a hump-shaped distribution, with few small (< 20 cm) and very large (> 80 cm) stemmed individuals. the bulk of the population have medium stem diameters (20 cm – 60 cm), indicating an episode of strong recruitment a century ago, followed by reduced recruitment. this is contrasted by the size class distribution of a. falcatus and c. africana, which both have populations dominated by smaller stemmed individuals and fewer larger individuals, suggesting that the demographic is in a recruitment and regeneration phase, following exploitation in the past. for all five of the chosen species, there are individuals within each population which have very large dbhs compared to the rest of the population, suggesting these are the remnants of large individuals that were established before the timber exploitation era, and were missed in harvesting operations. figure 4: stem size class distribution in diameter at breast height of five common species as well as the total sampled species of the pongola bush. (a) total sampled species; (b) combretum kraussii; (c) afrocarpus falcatus; (d) kiggelaria africana; (e) celtis africana; and (f) searsia chirindensis. the significant relationship between species richness and area, expressed as a log-shaped curve, shows the asymptotic species-area boundary of this curve has not been reached at 1 ha (10 000 m2), the upper limit of the aggregate sample (figure 5; r2 = 0.84 , p < 0.01). at this point, the graph still has an upward trend. extrapolating the relationship to the full extent of the pongola bush (~882 ha) suggests that there may be 70 woody plant species in the forest as a whole, thus about 30 undiscovered by this survey. figure 5: the multi modelled average species area relationship of the pongola bush both in terms of species richness against (a) area and (b) the log-log relationship. diversity indices, calculated on two fundamentals, are shown in table 2. the dominance index ranges from 0 to 1 with 1 being total dominance by an individual species, in this case dominance was lower for species diversity (0.065) compared to functional diversity (0.1492), which is supported by the biomass dominance by a small number of species shown in figure 3, and little dominance in figure 4, in terms of the gradual decrease of stem densities with no clear dominance. the simpsons index for individual counts (0.94) shows a high diversity and evenness (the scale is 0–1, with 1 high richness and evenness). when expressed as functional diversity, there is still a high degree of richness and evenness (0.85). the upper limit of the shannon index is unbounded, and shows the species diversity (2.94) to be higher than that of the functional diversity (2.25). table 2: the functional and species diversity based on three indices. one invasive species was identified in this study, solanum mauritianum scoop., but few individuals existed and were confined to areas where the canopy had been disturbed and establishment could take place, suggesting that the potential of rapid colonisation is unlikely given the interlocking canopy of the pongola bush. upon calculating community composition using multi-dimensional scaling based on the bray-curtis similarity coefficient, there appeared to be no clear tree compositional changes along the forest gradient. in the field there were apparent compositional changes of canopy floor species (forbs) and species which fell outside of the threshold stem diameter; thus these were not explicitly addressed in this study. this means that there are no predictable communities of certain species based on depth into the forest (or equivalently, altitude, because the slope is always upward with distance into the forest), and that the spread of species is relatively even throughout the forest. discussion the biomass is higher than those of fragmented dryer afromontane forests found in kenya and other regions (~50 mg ha−1), but lower than tropical african forests (150 mg ha−1 – 300 mg ha−1) in the albertine rift region (adhikari et al. 2017; baccini et al. 2008; glenday 2008; imani et al. 2017). the pongola bush is similar in biomass compared to african scarp subtypes at roughly 70 mg ha−1 (glenday 2008), but, as expected, only roughly a third of the biomass of pristine mature afrotemperate forests (adie, rushworth & lawes 2013). biomass in the pongola bush is not spread evenly among species but rather is dominated by four species; this dominance mirrors some of the other afrotemperate studies (adie et al. 2013). species that are commonly found in northern afrotemperate forests include many that were found in this study including, celtis, afrocarpus, halleria and scolopia; those missing were olinia, pittosporum and rothmania. when extrapolated, the species area relationship of roughly 70 woody species in the forest patch as a whole is consistent with other afromontane forest patches of similar size (50–110 species) (gemeda, lemenih & gole 2018; giliba et al. 2011; mensah et al. 2020). these northern afrotemperate forest patches are generally dominated by late successional, canopy forming podocarpus but due to relatively recent disturbance events common genera such as combretum and celtis have taken their place (asrat et al. 2020) resulting in different types of forest. diversity indices coincide with forests of similar composition in areas around africa, particularly afromontane forests. in terms of dominance, afromontane forests in tanzania have a range of dominance (0.043–0.135) for differing forest subtypes and fragments (giliba et al. 2011). in the cases where there was higher dominance, these were due to disturbances experienced by the forest subtypes (mafupa 2006). species richness is comparatively higher per ha in the pongola bush compared to patches seen both in specific studies in ethiopia and tanzania (gemeda et al. 2018; giliba et al. 2011), yet lower than southern african forests of like composition (adie et al. 2013). comparisons with other south african forests have also been made (geldenhuys 1992; mucina & rutherford 2006). shannon indices were generally lower in ethiopian afromontane forests (2.67–2.87). this may be due to various factors including disturbance and limited rainfall (500 mm per annum) compared to the pongola bush (gemeda et al. 2018; mekonnen 2006). geldenhuys (1992) stated that the number of tree species in southern africa is lower than that in the tropics. it was unfortunate that information on other plant forms and species, for example, ferns was not collected as this would have provided an additional element to this biodiversity study. there are cases, in other scarp forests, of unsustainable harvesting to the point of local extinction of medicinal species such as ocotea bullata. this species was not encountered in this survey, suggesting it may too have extirpated from this particular forest but has been known to occur in the forest in the past. medicinal plant exploitation may reduce the species diversity of the forest which would be of concern if there was extensive extraction of all medicinal species. regeneration potential using stand density in the pongola bush forest fragment is similar to some other studies in southern african afromontane patches, but these studies show highly variable results based on a range of biotic and abiotic factors. the pongola bush shows a high recruitment of small stems to a decrease in larger stems in a ‘reverse j’ distribution, as demonstrated in other studies (adie et al. 2013; asrat et al. 2020; eriksson, teketay & granström 2003; mensah et al. 2020). smaller-stemmed plants such as s. mossambicensis and many smaller individuals of potentially large trees, such as a. falcatus dominate the forest. populations of c. kraussii have few small stems which contrasts with previous stem diameter studies of forests in the region (mensah et al. 2020). the limited small stem recruits could potentially affect the future composition of the forest as c. kraussii is a significant contributor to agb and stem density. further studies on this species in this location should be included in an integrated monitoring plan. this trend of small stemmed recruits is also occurring for s. chirindensis and k. africana. combretum erythrophyllum has been recorded in northern afrotemperate forests but the occurrence of c. kraussi and k. africana are interesting. given that these species form a significant part of the forest composition there is likely to be a large shift in dominant species many decades into the future. species such as a. falcatus and c. africana show the classical ‘j’ shaped curve which shows that there is a stable regeneration potential for the future of their populations (botzat, fischer & farwig 2015). these stem diameter differences between species may be due to previous exploitation which has put the forest in earlier stages of succession as seen with other south african forests as well as south american forests of similar composition (adie et al. 2013; echeverría et al. 2007). emergence of earlier successional features is correspondingly correlated to a decreased patch size or fragmentation (echeverría et al. 2007), but this forest extent is regulated by various external factors such as historic grassland fire regimes and specific abiotic factors; hence, fragmentation or decreased patch size may not have induced the early successional traits. despite there being limited current forest fragmentation of this patch due to its conservation, other patches of like composition may be under threat from fragmentation, particularly from fire, and this patch may have been part of a much larger constellation of related forests in the distant past (e.g. in the pliocene, when the climate was wetter). in conclusion, the pongola bush is a highly diverse, relatively dense, and stable habitat at present due to current conservation efforts. at the time of the study, four species dominated in terms of biomass, but this was only a preliminary study. a much more detailed study over time is needed in order to contribute to the management and conservation of these types of forests. there remains an information gap on non-tree biodiversity elements within the forest, such as forest floor species, forbs and animal diversity. having a comprehensive study on the full diversity of the forest will boost the knowledge of the forest and may aid in future conservation. additionally, the dynamic between forest patches in the region and dispersal mechanisms of both seeds and pollen as genetic material between patches could be better understood. this forest is an understudied remnant of perhaps an early evolutionary stage in the northern afrotemperate forest system and its conservation should be encouraged to ensure further studies on the flora and fauna. acknowledgements the authors thank rickert van der westhuizen and amon ndlovu from ezemvelo wildlife (kzn wildlife) for organising travel and research permits, and facilitating accommodation at the pongola bush. the authors are also grateful for the field assistance provided by bianca ferreira and zoe khumalo. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions r.j.s. conceived and designed the project, m.c.s. supervised the project and t.f.h. carried out the data collection, analysis and writing of the manuscript in consultation with r.j.s. and m.c.s. funding information the authors thank the national research foundation (sarchi: 101057) for providing the funding that made this project possible. data availability data supporting the findings of this study are available from the corresponding author, t.f.h., upon reasonable request. disclaimer the views expressed in the submitted article are those of the authors and not an official position of the affiliated institution or those of the funders, or publisher. references adhikari, h., heiskanen, j., siljander, m., maeda, e., heikinheimo, v. & pellikka, p.k.e., 2017, ‘determinants of aboveground biomass across an afromontane landscape mosaic in kenya’, remote sensing 9(8), 827. https://doi.org/10.3390/rs9080827 adie, h., rushworth, i. & lawes, m.j., 2013, ‘pervasive, long-lasting impact of historical logging on composition, diversity and above ground carbon stocks in afrotemperate forest’, forest ecology and management 310, 887–895. https://doi.org/10.1016/j.foreco.2013.09.037 asrat, z., eid, t., gobakken, t. & negash, m., 2020, ‘aboveground tree biomass prediction options for the dry afromontane forests in south-central ethiopia’, forest ecology and management 473, 118335. https://doi.org/10.1016/j.foreco.2020.118335 baccini, a., laporte, n., goetz, s.j., sun, m. & dong, h., 2008, ‘a first map of tropical africa’s above-ground biomass derived from satellite imagery’, environmental research letters 3(4), 045011. https://doi.org/10.1088/1748-9326/3/4/045011 berliner, d., 2009, ‘systematic conservation planning and south africa’s forest biome: an assessment of the conservation status of south africa’s forest and recommendations for their conservation’, phd thesis, university of cape town. botzat, a., fischer, l. & farwig, n., 2015, ‘regeneration potential in south african forest fragments: extinction debt paid off or hampered by contemporary matrix modification?’, plant ecology 216(4), 535–551. https://doi.org/10.1007/s11258-015-0457-9 carsan, s., orwa, c., harwood, c., kindt, r., stroebel, a., neufeldt, h. et al., 2012, african wood density database, world agroforestry centre, nairobi. chave, j., réjou-méchain, m., búrquez, a., chidumayo, e., colgan, m.s., delitti, w.b.c. et al., 2014, ‘improved allometric models to estimate the aboveground biomass of tropical trees’, global change biology 20(10), 3177–3190. https://doi.org/10.1111/gcb.12629 chazdon, r.l., brancalion, p.h.s., laestadius, l., bennett-curry, a., buckingham, k., kumar, c. et al., 2016, ‘when is a forest a forest? forest concepts and definitions in the era of forest and landscape restoration’, ambio 45(5), 538–550. https://doi.org/10.1007/s13280-016-0772-y echeverría, c., newton, a.c., lara, a., benayas, j.m.r. & coomes, d.a., 2007, ‘impacts of forest fragmentation on species composition and forest structure in the temperate landscape of southern chile’, global ecology and biogeography 16(4), 426–439. https://doi.org/10.1111/j.1466-8238.2007.00311.x eriksson, i., teketay, d. & granström, a., 2003, ‘response of plant communities to fire in an acacia woodland and a dry afromontane forest, southern ethiopia’, forest ecology and management 177(1), 39–50. https://doi.org/10.1016/s0378-1127(02)00325-0 falk, d.a., palmer, m.a. & zedler, j.b., 2013, foundations of restoration ecology, island press, washington, dc. fao, 2010, global forest resources assessment 2010 – food and agriculture organization of the united nations, viewed 20 april 2020, from http://www.fao.org/3/a-i1757e.pdf. gauch, h.g., 1973, ‘a quantitative evaluation of the bray-curtis ordination’, ecology 54(4), 829–836. https://doi.org/10.2307/1935677 geldenhuys, c.j., 1992, ‘richness, composition and relationships of the floras of selected forests in southern africa’, bothalia 22(2), 205–233. https://doi.org/10.4102/abc.v22i2.847 geldenhuys, c.j., 1994, ‘bergwind fires and the location pattern of forest patches in the southern cape landscape, south africa’, journal of biogeography 21(1), 49–62. https://doi.org/10.2307/2845603 gemeda, w., lemenih, m. & gole, t., 2018, ‘analysis of plant species diversity and forest structure in arero dry afromontane forest of borena zone, south ethiopia’, tropical plant research 5(2), 129–140. https://doi.org/10.22271/tpr.2018.v5.i2.018 giliba, r.a., boon, e.k., kayombo, c.j., musamba, e.b., kashindye, a.m. & shayo, p.f., 2011, ‘species composition, richness and diversity in miombo woodland of bereku forest reserve, tanzania’, journal of biodiversity 2(1), 1–7. https://doi.org/10.1080/09766901.2011.11884724 glenday, j., 2008, ‘carbon storage and emissions offset potential in an african dry forest, the arabuko-sokoke forest, kenya’, environmental monitoring and assessment 142(1), 85–95. https://doi.org/10.1007/s10661-007-9910-0 grainger, a., 2013, controlling tropical deforestation, routledge, abingdon, oxfordshire. hammer, o., harper, d.a.p. & ryan, p.d., 2001, ‘paleontological statistical software package (past) for education and data analysis’, palaeontologia electronica 4(1), 1–9. hooper, d.u., chapin, f.s., ewel, j.j., hector, a., inchausti, p., lavorel, s. et al., 2005, ‘effects of biodiversity on ecosystem functioning: a consensus of current knowledge’, ecological monographs 75(1), 3–35. https://doi.org/10.1890/04-0922 huntley, b.j., 1984, ‘characteristics of south african biomes’, in p.v. de booysen & n.m. tainton (eds.), ecological effects of fire in south african ecosystems, pp. 1–17, springer, colfax county, new mexico. imani, g., boyemba, f., lewis, s., nabahungu, n.l., calders, k., zapfack, l. et al., 2017, ‘height-diameter allometry and above ground biomass in tropical montane forests: insights from the albertine rift in africa’, plos one 12(6), e0179653. https://doi.org/10.1371/journal.pone.0179653 johnson, k.h., vogt, k.a., clark, h.j., schmitz, o.j. & vogt, d.j., 1996, ‘biodiversity and the productivity and stability of ecosystems’, trends in ecology & evolution 11(9), 372–377. https://doi.org/10.1016/0169-5347(96)10040-9 lötter, m.c., mucina, l. & witkowski, e.t.f., 2014, ‘classification of the indigenous forests of mpumalanga province, south africa’, south african journal of botany 90, 37–51. https://doi.org/10.1016/j.sajb.2013.09.010 mafupa, c., 2006, ‘impact of human disturbances in miombo woodlands of igombe river forest reserve, nzega district, tanzania’, msc thesis unpublished, sokoine university of agriculture, morogoro. mason, n.w.h., de bello, f., mouillot, d., pavoine, s. & dray, s., 2013, ‘a guide for using functional diversity indices to reveal changes in assembly processes along ecological gradients’, journal of vegetation science 24(5), 794–806. https://doi.org/10.1111/jvs.12013 matthews, t.j., triantis, k.a., whittaker, r.j. & guilhaumon, f., 2019, ‘sars: an r package for fitting, evaluating and comparing species – area relationship models’, ecography 42(8), 1446–1455. https://doi.org/10.1111/ecog.04271 mekonnen, d., 2006, ‘woody species composition of dilfaqar regional park and its socioeconimic importance’, masters thesis, addis ababa university. mensah, s., egeru, a., assogbadjo, a.e. & glèlè kakaï, r., 2020, ‘vegetation structure, dominance patterns and height growth in an afromontane forest, southern africa’, journal of forestry research 31(2), 453–462. https://doi.org/10.1007/s11676-018-0801-8 mucina, l. & rutherford, m.c., 2006a, vegetation atlas of south africa, lesotho and swaziland, south african botanical institute, pretoria. mucina, l. & rutherford, m.c. (eds.), 2006b, the vegetation of south africa, lesotho and swaziland, strelitzia 19. south african national biodiversity institute, pretoria. ruhberg, h. & hamer, m., 2005, ‘a new species of opisthopatus purcell, 1899 (onychophora: peripatopsidae) from kwazulu-natal, south africa’, zootaxa 1039(1), 27. https://doi.org/10.11646/zootaxa.1039.1.3 rutherford, m.c., mucina, l. & powrie, l.w., 2006, ‘biomes and bioregions of southern africa’, in l. mucina & m.c. rutherford (eds.), the vegetation of south africa, lesotho and swaziland, vol. 19, pp. 30–51, south african national biodiversity institute, pretoria. schleuter, d., daufresne, m., massol, f. & argillier, c., 2010, ‘a user’s guide to functional diversity indices’, ecological monographs 80(3), 469–484. https://doi.org/10.1890/08-2225.1 vieira, d.l.m. & scariot, a., 2006, ‘principles of natural regeneration of tropical dry forests for restoration’, restoration ecology 14(1), 11–20. https://doi.org/10.1111/j.1526-100x.2006.00100.x appendix 1 table 1-a1: wood density (g cm−3) for the sampled species. about the author(s) wijnand j. swart department of plant sciences, faculty of natural & agricultural sciences, university of the free state, bloemfontein, south africa maitland t. seaman centre for environmental management, faculty of natural & agricultural sciences, university of the free state, bloemfontein, south africa pieter a.l. le roux digital soils africa, bloemfontein, south africa institute for groundwater studies, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa beanelri b. janecke department of animal, wildlife & grassland sciences, faculty of natural & agricultural sciences, university of the free state, bloemfontein, south africa citation swart, w.j., seaman, m.t., le roux, p.a.l., janecke, b.b. 2020, ‘a tribute to frederick (fred) j. kruger’, koedoe 62(2), a1639. https://doi.org/10.4102/koedoe.v62i2.1639 in memorial: frederick (fred) j. kruger note: special issue: connections between abiotic and biotic components of a granite catena ecosystem in kruger national park, sub-edited by beanelri janecke and johan van tol. tribute a tribute to frederick (fred) j. kruger wijnand j. swart, maitland t. seaman, pieter a.l. le roux, beanelri b. janecke copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. frederick j. kruger, passed away in his sleep in may 2017 in stellenbosch after a short illness. fred, as he was known to everyone, was highly educated in forest science and ecology and a pioneer of forest hydrology, fynbos ecology and invasive species. fred was a member of the royal society, and a former director of the south african forestry research institute, and of the csir’s division of forest science and technology. his passing left an indelible mark on forestry and environmental science in south africa. fred was an intellectual (although he refused to be called an academic) and uncompromising in his pursuit of answers. he was supportive, understated but provocative, with dignity and integrity and always a twinkle in his eyes. in recent years, he was a major contributor to scholarly literature about the environmental history of south africa. he often collaborated in ecological research and was actively involved from 2015 to 2017 in this multidisciplinary research project near skukuza in the kruger national park (published in this special issue) together with colleagues from the university of the free state. the main concept of the study was the ‘brain child’ of fred and piet le roux. fred stimulated the team’s thoughts on the interaction between the biosphere and lithosphere and he encouraged us to always focus on the bigger ecological picture. his active involvement in this project led to the discovery of a new species of fungus, which was subsequently described and named after him (fusarium fredkrugeri sandoval-denis, crous & w.j. swart, sp. nov.). his extensive knowledge and experience will be sorely missed by all those people he interacted with over many years. at the time of his passing, he was an active research fellow of the centre for environmental management at the university of the free state, and was investigating and documenting studies on the role of water in rivers in the catchments of the kruger national park. this is a tribute to honour dr fred kruger, and to show our respect, gratitude and admiration when we remember the huge contribution that he has made to the research published in this special issue. – w. swart, m. seaman, p. le roux, b. janecke and the research team abstract introduction literature review materials and methods results discussion conclusion acknowledgements references about the author(s) kevin m. coldrey environmental policy research unit, department of economics, university of cape town, cape town, south africa jane k. turpie environmental policy research unit, department of economics, university of cape town, cape town, south africa citation coldrey, k.m. & turpie, j.k., 2020, ‘potential impacts of changing climate on nature-based tourism: a case study of south africa’s national parks’, koedoe 62(1), a1629. https://doi.org/10.4102/koedoe.v62i1.1629 original research potential impacts of changing climate on nature-based tourism: a case study of south africa’s national parks kevin m. coldrey, jane k. turpie received: 09 mar. 2020; accepted: 07 sept. 2020; published: 29 oct. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract climate change could potential have significant impacts on the global tourism industry through changes in accessibility, comfort levels, and geographic or seasonal changes to attractions. global and regional studies based on climate indices suggest that there will be global shifts in tourism, with many of the currently warmer areas potentially experiencing decreases in tourism. these include much of the global south, where nature-based tourism is both an important contributor to economic output and believed to be vulnerable to climate change. protected areas form the core of nature-based tourism in many developing countries and are also directly or indirectly dependent on this tourism for their funding. however, comparatively little empirical work has been done on protected area tourism, particularly in developing countries. we modelled the relationship between occupancy and mean monthly temperature at each of south africa’s 19 national parks and estimated the potential change in demand as a result of climate change. we found significant quadratic or linear relationships between occupancy and temperature for all parks. a single network-wide model could not capture all the location-specific temperature and tourism demand dynamics occurring at individual parks. the temperature models explained over 50% of the variance in historical occupancy for 17 parks, suggesting that climate change may have a significant impact on the timing and magnitude of demand across the network. based on projected future temperatures, visitation rates across all parks could decline by nearly 4% by 2050, with most of the decline in kruger national park, which has the lion’s share of visitors. conservation implications: the results are of concern given that tourism proceeds generated by kruger national park are used to subsidise the operating budgets of the entire park system, which would put the capacity for conservation activities at risk, suggesting that measures to alleviate climate discomfort to tourists should be explored by management. keyword: climate change; tourism; protected areas; statistical analysis; national parks. introduction climate change has the potential to have significant impacts on the global tourism industry through changes in accessibility and comfort levels, as well as geographic or seasonal changes to attractions (rosselló-nadal 2014; scott, gössling & hall 2012). global and regional studies based on climate indices and data suggest that there will be global shifts in tourism, with an improvement in conditions in many temperate areas and a deterioration in conditions in many subtropical and tropical areas (bigano, hamilton & tol 2007; rosselló & santana 2012; tol & walsh 2012). indications are that tourism may decline in much of the global south. this is of particular concern, not only because tourism is a significant source of foreign exchange earnings in developing countries (scott et al. 2012), but also because of the global importance of biodiversity conservation in these countries and the direct and indirect reliance of conservation on nature-based tourism. however, comparatively little empirical work has been done on nature-based or protected area tourism, particularly in developing countries. protected areas form the core of both biodiversity conservation and nature-based tourism throughout africa. in south africa, the national parks are semi-autonomous and derive much of their operating budget from tourism revenues. in addition to the threats of climate change on park biodiversity and infrastructure, climate change may also have an impact on tourism numbers and revenues as a result of extreme events such as drought and changes in expected daily maximum temperatures. indeed, nature-based tourism has been identified as being particularly vulnerable to climate change, and this has been verified in several recent studies undertaken in the region (dube & nhamo 2018, 2019, 2020; dube et al. 2018; macfadyen et al. 2019; mushawemhuka, rogerson & saarinen 2018; sutherland, ndlovu & pérez-rodríguez 2018). this study extends the existing literature by undertaking an empirical analysis of the sensitivity of park tourism to temperature, based on historical data. south africa has a large nature-based tourism industry, a well-managed national park system and reliable historic tourism data. furthermore, substantial and varying climate change is predicted for south africa (davis-reddy & vincent 2017), which has the potential to alter demand for this industry (pandy & rogerson 2018; rogerson 2016). a study by van wilgen et al. (2016) on the change in climate already experienced in south africa’s national parks over the past few decades indicates that significant temperature increases have already been experienced in most parks. the observed temperature changes over the last 20–50 years have, in several instances, already reached those predicted for near future scenarios. these changes in temperature may have already influenced the timing of park visitation in the hotter, non-coastal areas of the country, where peaks have shifted to the cooler winter months. this study uses south africa’s national parks as a case study to explore the influence of climate on tourism demand by using empirical models, so that the potential unmitigated impact of climate change on tourism demand can be estimated. following fisichelli et al.’s (2015) study of parks in the united states of america, we analysed historical monthly visitation data for parks in relation to mean monthly air temperature data obtained from the climatic research unit (cru). our study shows that impacts are positive for some more temperate parks, but that overall visitation rates are likely to decrease in the absence of mitigation. literature review tourism is a highly climate-dependent industry (amelung, nicholls & viner 2007; hamilton, maddison & tol 2005), which can either act to compel tourists to leave their own countries in search of more favourable locations, or it can compel tourists to seek out locations with climates that lend themselves to specific tourism attractions, such as snowfall for winter sports tourism or temperate weather for coastal and nature-based tourism (rosselló-nadal 2014; scott et al. 2012). apart from providing the means for tourists to partake in specific activities, a pleasant, comfortable climate has the ability to draw tourists as much as other more catalogued destination attributes (de freitas 2003; kozak 2002). in fact, many regions rely on their climate as the natural resource that draws tourists (scott et al. 2006). it is important to note that tourists’ choices are different from those of recreationists, in that the latter can react to immediate weather, whereas tourism decisions are planned more in advance (amelung et al. 2007). thus, tourism will be a function of expected conditions for a locality in any given month. furthermore, climate is one of the factors affecting the seasonality of tourism, along with other natural and institutional factors such as the timing of holidays. thus, climate change is likely to impact the seasonality of tourism as well as overall tourism arrivals (koenig & bischoff 2004). although earlier models of tourism demand seldom considered climate (pike 2002), the impact of climate on tourism was first studied in the 1980s and became a dominant theme in the tourism literature by the 2000s (hamilton & tol 2007; scott et al. 2012). studies on the potential impacts of climate change on tourism date back to the 1990s, when abegg and koenig (1997) predicted a reduction in the reliability of snow for swiss ski areas. studies have used a range of qualitative and statistical approaches to consider the potential impacts of changes in climate and/or its impacts on physical conditions for tourism (moore 2010; rosselló-nadal 2014). irrespective of the approach, studies have consistently found that there is a quadratic (inverted u) relationship between tourism and climate (hamilton et al. 2005; rosselló-nadal 2014). in other words, preferences tend to be for moderate or ‘comfortable’ conditions (lise & tol 2002; mieczkowski 1985), and climate change could positively or negatively impact on tourism demand. many studies have made use of a composite index of climate suitability, notably mieczkowski’s (1985) tourism climate index (tci), which is a weighted index composed of seven normalised variables. this index has been used to assess potential changes in tourism demand in various parts of the world (e.g. amelung & viner 2006; amelung et al. 2007; perch-nielsen, amelung & knutti 2010; roshan, yousefi & fitchett 2015; scott & mcboyle 2001; scott et al. 2016). for example, by using the tci, amelung et al. (2007) explored the potential impacts of climate change on tourism at a global scale. they calculated the tci for each grid cell across the land surface of the earth for different decades in the past and future and classified ‘good’ months as those with a tci greater than 70. they found that although some locations are likely to experience substantial increases in attractiveness, others may become significantly less appealing. indeed, these studies have consistently shown that temperate regions tend to become more attractive, whereas tropical and subtropical areas tend to become less attractive. however, this approach is criticised because ideal conditions might vary across destinations (moore 2010; scott, gossling & freitas 2007), as well as for tourists from different origins (lin & matzarakis 2008). these rather qualitative approaches, such as those predicting negative impacts of heat waves in mediterranean countries, also came under criticism for being unable to define what conditions would be too severe for tourists (scott et al. 2012). predicting future changes is arguably best dealt with through statistical and econometric approaches that use historical arrivals data. such approaches have used time series, cross-sectional and panel data analyses (maddison 2001; moore 2010; rosselló-nadal 2014). although time series and cross-sectional approaches are vulnerable to model specification and other biases, panel data approaches have the advantage of controlling for fixed or random effects. moore (2010) incorporated the tci as a variable in a dynamic panel approach with monthly observations over a number of years, to predict the effects of climate change for caribbean destinations. their rigorous analysis suggested reductions in tourism arrivals by 1% – 5% under a 2071–2100 climate. nevertheless, use of the tci is still questionable as it has not been tested empirically (scott et al. 2012), and use of a weighted index still implies a constant weighting of temperature variables within the index across all tourist types and destinations. avoiding this problem, globaland regional-scale tourism impacts have also been modelled by using simple climate variables such as temperature and/or precipitation (hamilton & tol 2007; rosselló & santana 2012; tol & walsh 2012). for example, using pooled ordinary least squares (ols) regression and controlling for a range of factors that attract or deter tourists, such as coastline, poverty and heritage, tol and walsh (2012) found that tourists were attracted by intermediate precipitation and temperatures, with those from warmer countries being more sensitive to climate. few, if any, statistical or econometric studies have used the range of climate variables incorporated into mieczkowski’s (1985) tci. this may be partly because of the difficulty of obtaining suitable data to properly populate the tci, as was found in south africa (fitchett, robinson & hoogendoorn 2017). studies that have focussed on the global tourism industry at the macro-country level have indicated that climate change could lead to a gradual shift of tourists towards higher latitudes and that climate change would lead to tourists from temperate nations altering their travel patterns to take advantage of new climatic opportunities closer to home. this trend is likely to be exacerbated by the rising aversion to long-haul flights owing to tourists’ considerations of their carbon footprints. this shift in tourism destinations globally would be at the expense of tropical and sub-tropical destinations (bigano, hamilton & tol 2007; hamilton et al. 2005). however, such analyses may overlook the niche climates within countries that attract tourists. with climate change predicted to vary spatially across countries (chown 2010; collier, conway & venables 2008), changes in climate at a more micro-scale are likely to alter the distribution or seasonality of tourism demand within countries. although many of the well-studied northern hemisphere destinations expect to see moderate changes or increases because of warmer temperatures, destinations that are already relatively warm and have relatively good tci indices, such as south africa (fitchett et al. 2017), may be expected to experience decreased demand because of increased heat discomfort as well as other climate change effects such as increased drought, habitat and wildlife changes and poverty. impacts on tourism are also likely to have a disproportionately higher significance in the global south (rogerson 2016). indeed, given the severity of projected climate change in southern africa, this region is considered one of the most vulnerable in the world (hoogendoorn & rogerson 2015; pandy 2017; rogerson 2016). in terms of the risks to the tourism industry, nature-based tourism is considered particularly vulnerable (pandy & rogerson 2018; preston-whyte & watson 2005; steyn & spencer 2012). apart from the expected changes in comfort levels for visitors, nature-based tourism destinations are vulnerable to changes in the species and ecosystems that attract those visitors (belle et al. 2016; dunlop & brown 2008; lemieux et al. 2010; van der merwe & saayman 2008). changing demand for nature-based tourism will add to the range of threats that face protected areas under climate change. protected area systems are often directly dependent on revenues from tourism or on government support, which is partly leveraged because of their contribution to national tourism. as for tourism in general, the impacts of climate change on demand for park visitation could be very different in different parts of the world. in one of the first studies focussing on climate change impacts on protected area tourism, jones and scott (2006) used ols regressions between visitation and climate variables (temperature and precipitation) to predict an increase in visitation to canadian protected areas under future climate conditions. fisichelli et al. (2015) modelled historical monthly visitation data for parks in the united states of america in relation to mean monthly air temperature by using generalised linear models. their models (for the park system as a whole as well as for individual parks) also predicted an increase in future visitation as a result of climate change, particularly in the low season. in their study, the response variable was the proportion of the park’s annual visits occurring in a month (to standardise across parks with different overall rates of visitation), and the explanatory variables were mean monthly temperature and park name. in southern africa, on the other hand, climate change is a serious concern for park tourism (dube & nhamo 2020; macfadyen et al. 2019; sutherland et al. 2018). tourists to south africa are potentially sensitive to high temperatures (giddy, fitchett & hoogendoorn 2017), and in kruger national park the annual average maximum temperature has increased from around 29 °c to over 31 °c since the 1970s (dube & nhamo 2020). a series of mixed-methods studies involving semi-structured interviews have shown that rising temperatures and drought have already impacted on tourism experiences in flagship parks of the region such as kruger national park, victoria falls, the okavango delta and the hwange national park (dube & nhamo 2018, 2019; dube et al. 2018; mushawemhuka et al. 2018), as well as in private nature reserves (smith & fitchett 2020). these conclusions are supported by empirical evidence of the negative impact of drought on numbers of visitors in kruger national park (mathivha, tshipala & nkuna 2017). however, to date, no empirical studies have been carried out on the impacts of temperature on park tourism in southern africa. materials and methods study area south africa has 19 national parks, which are managed by south african national parks (sanparks), a state-owned parastatal. these parks constitute 52% of terrestrial protected area in the country and are located throughout south africa (figure 1). each park has its own unique climate, topography and water resources, resulting in a diversity of vegetation and species conserved at the national level, and a unique tourist offering at most parks. because of the large spatial diversity between parks, the projected change in climate for individual parks varies greatly, with some parks expected to experience far larger changes in temperature and precipitation than others (table 1). figure 1: map showing the national parks of south africa. table 1: projected percentage change in mean annual temperature and total annual precipitation for each park from historic average (1960–1990) to mid-century (2040–2060), by using coupled model inter-comparison project data. climate data following fisichelli et al. (2015), historical monthly occupancy data were matched with gridded historical monthly mean temperature (°c) data obtained from the cru high-resolution time series version 4.03 (harris et al. 2020). climatic research unit data are derived by the interpolation of monthly climate anomalies from extensive networks of weather station observations and are globally available at 0.5 decimal degrees (~50 km2) for each month and year of occupancy data (n = 81). cru data were deemed preferable to south african weather service (saws) data for this study as they are publicly available and free to download, cru incorporates quality-controlled data from individual stations (including saws data) to produce a uniform grid of data points spanning the entire earth and saws data is not available for all of south africa’s national parks and there are considerable gaps in the data and differences in the type of data available at the park level (see van wilgen et al. 2016). although a concerted effort is being made globally to curb greenhouse gas emissions, up to at least 2014, emissions have been tracking just above the representative concentration pathway (rcp) 8.5 scenario, considered the worst-case scenario by the intergovernmental panel on climate change (ipcc) (sanford et al. 2014). owing to the consensus that the climate change we experience over the next few decades will primarily be caused by past emissions (glick, stein & edelson 2011), the rcp 8.5 scenario was used in this study. furthermore, because near-term projections of climate change scenarios tend to have a higher degree of certainty, a relatively short time horizon of mid-century (2040–2060) was used. apart from the uncertainty inherent in longer time frames, the choice of mid-century limits the magnitude of divergence between the different rcps. statistical analysis monthly occupancy data, spanning the period april 2012 to december 2016, for both accommodation units and campsites were used as the dependent variable in this study (n = 57), a proxy for tourism demand. occupancy levels were chosen instead of total visitor numbers (which includes day visitors) because occupancy is driven largely by tourist perceptions of mean monthly weather for holiday months, and day visitors may time their visits to parks to avoid the hottest or coldest times of the day. although tourism demand is influenced by a number of factors (biggs et al. 2014), this article investigates the influence of climate, which was not considered by their study. relationships between mean monthly occupancy rates, mean monthly temperature and rainfall were investigated at park level and overall. on this basis the more detailed statistical analysis omitted rainfall. this is consistent with the findings of mathivha et al. (2017) that there was no statistically significant relationship between tourist arrivals and rainfall. therefore, mean monthly temperature and holiday months were used as explanatory variables, as well as the interaction between the two. in support of the use of temperature, giddy et al. (2017) found that american tourists most often cited very hot temperatures as negative experiences with weather during their stay in south africa. the holiday months included the peak tourism months of april, july, september and december, which cover the main school and university holidays. the majority of domestic tourists only have these small windows in which to plan their holiday activities (amelung et al. 2007) and climate change may make it unbearable to visit certain parks during these months. the remaining 24 months (january 2017 to december 2018) of occupancy data was held back for out-of-sample model evaluation. park-specific models were developed to assess the relationship between historical occupancy and climate by using r statistical software (r core team 2014). for each park, ols regression was used, and three models of increasing complexity were tested to quantify the relationship between occupancy levels and temperature and holiday months: a null model (intercept only); the first-order (linear) polynomial function of temperature and holiday months; and the second-order (quadratic) polynomial function of temperature and holiday months. bayesian information criterion (bic) was used to identify the best model for each park, and the model selections were intuitively assessed to make sure that the signs of the variables were consistent with expectations. the best-fit models were then used to estimate the change in annual occupancy between the baseline period (2012–2016) and the future period (2040–2060). the turning point of the quadratic best-fit functions was calculated by using the formula b1/-2b2. park-specific model performance was assessed by using root mean square error (rmse), an indicator of prediction accuracy. rmse for each park was calculated for both the estimation period (in-sample; n = 57) and the validation period (out-of-sample; n = 24) by using the following formula: where and yt are the predicted monthly occupancy and observed monthly occupancy rates, respectively. fixed effects and random effects panel regression analysis was also performed to determine if a single model could be fitted to the entire park system. three separate models were tested: the quadratic function of temperature and holiday months on all 19 parks; the same quadratic function but on only the parks that exhibited a quadratic best-fit model; and the linear function of temperature and holiday months on only the parks that exhibited a linear best-fit model. lagrange multiplier and f tests were performed on the three models to determine if there were significant differences between the random effects model and the ols model and the fixed effects model and the ols model, respectively. finally, a hausman test was performed to determine whether the fixed effects or random effects model should be considered. all models assumed that visitor numbers were minimally constrained by bed capacity, and that installation of visitor facilities responded to demand consistently across all parks. ethical consideration the authors declare that ethical clearance was not required for the study. results influence of temperature on tourist numbers in nearly all cases, mean monthly occupancy rates were strongly related to mean monthly temperatures (figure 2). depending on the range of temperatures experienced at each park, the relationships were either positive, negative or an inverse u. however, the relationship to temperature was not consistent across parks. occupancy rates were also correlated with rainfall, but the r2 value was also lower than for the relationship with temperature in all but four cases. rainfall was strongly correlated with temperature in all but two parks, either positively in the case of the summer rainfall areas (13 parks) or negatively in the winter rainfall area parks (agulhas, bontebok, namaqua, table mountain, tankwa karoo and west coast national parks). inclusion of mean monthly rainfall generally made little difference to model fit, and frequently did not have the expected negative sign on the coefficient. therefore, rainfall was not considered to be an important driver of occupancy rates. figure 2: relationship between mean monthly temperature (°c) and mean monthly percentage occupancy rate for each national park and overall. parks are presented in alphabetical order. the results of the statistical analysis in combination with other control variables indicated that for 13 of the parks, occupancy and temperature exhibit a significant quadratic relationship, although for six of the parks, occupancy and temperature exhibit a significant linear relationship. two of the parks (augrabies falls and mokala national parks) had an r2 value of less than 0.5 and for augrabies falls national park, which has one of the highest year-round average temperatures, the temperature variables are insignificant (table 2). the temperature models explained more than 50% of the variance in historical occupancy for the remaining 17 parks, with 14 parks having more than 60% of the variance in historical occupancy explained by the models and seven of those parks had more than 70% of the variance in historical occupancy explained by the models (table 2). table 2: results of the statistical analysis, indicating the best-fit model for each park. for the parks exhibiting a quadratic function, the temperature turning point is also shown. significance: < 0.001 ***, < 0.01 **, < 0.05 * for the parks exhibiting a quadratic temperature function, the turning point above which occupancy levels decreased ranged from 14.2 °c for golden gate national park to 25.7 °c for bontebok national park (table 2). the turning points for these parks were not significantly correlated with the mean annual temperatures of the parks. however, with the exception of bontebok national park, the turning points were significantly correlated with mean occupancy level (n = 12, r2 = 0.57, p < 0.01) and maximum occupancy level (r2 = 0.62, p < 0.01). note that mean occupancy level was a logarithmic function of maximum occupancy (y = 36.318ln(x)-66.194, r² = 0.87), suggesting that parks with the highest occupancy levels were experiencing some level of capacity constraints. the results of the fixed effects and random effects panel regression analyses of the overall dataset were all highly significant, but explained less of the variance, suggesting that a single model could not capture all the location-specific temperature and tourism demand dynamics (table 3). for each of the three models tested, both the fixed effects and random effects models were superior to the ols model (table 3). in addition, the hausman test indicated that the random effects models were better than the fixed effects models. for the two quadratic panel regression models (model 1 and model 2), the turning points (19.9 °c and 18.5 °c, respectively) were close to the mean turning point of the 13 individual park models of 18.8 °c (table 3), as well as the overall mean average temperature of the parks of 18.4 °c. table 3: results of the fixed effects and random effects panel regressions and the results of the lagrange multiplier test for balanced panels, the f test for individual effects and the hausman test. how accurate were the models in predicting occupancy rates the rmse values for the estimation period were all below 10.0, except for west coast national park (10.8), which indicates that the model predictions of monthly occupancy rates were out by less than 10% on average (table 4). the rmse values for the validation period improved for five of the parks but worsened marginally for the remaining 14 parks, which is to be expected. these results indicate that the models have a less than 10% error margin for 15 parks, with the remaining four parks having an error margin marginally higher than 10% (table 4). table 4: root mean square error for both the estimation and validation periods, and the predicted change in annual units occupied for each park based on the individual best-fit models. how will climate change impact on occupancy based on the empirical models the best-fit models were used to predict the change in annual units occupied for all parks except for augrabies falls national park, for which temperature variables were insignificant (table 4). the results indicate that only six of the parks assessed are expected to experience a decline in annual occupancy, despite 13 parks exhibiting a quadratic best-fit model. the predicted impact of a warmer climate on the total park system is a decline of nearly 4%, largely owing to the predicted decline in annual occupancy for kruger national park (6.4%), which accounts for the lion’s share of the national park systems’ overnight visitors. the other parks that contributed to the total park system decline were kgalagadi national park (the south african component of the kgalagadi trans-frontier park; 2.4% decline), mapungubwe national park (9.7%), marakele national park (8.7%), mountain zebra national park (0.7%) and tankwa karoo national park (6.6%; table 4). discussion if high mean monthly temperatures were a significant deterrent to tourists, then it would be expected that, if all other variables are adequately controlled for, there should be a relatively consistent marginal effect of temperature, and a general threshold mean monthly temperature above which visitation levels are negatively affected. the fixed effects panel data regression, which controlled for time invariant differences between the parks (i.e. a whole suite of park characteristics such as locational and wildlife attributes), confirmed that temperature does have a significant effect on overall occupancy rates. however, this model suggests a turning point temperature of around 18.8 °c. interestingly, this threshold was close to the overall mean average temperature of the parks of 18.4 °c. this suggests that, after controlling for school holiday periods, visitors prefer to visit parks in reasonably cool–mild temperatures. the slightly lower threshold temperature obtained in the panel data analyses warranted closer inspection of the relationships in individual parks. the results of the empirical analysis suggest that tourists to south africa’s national parks are sensitive to temperature. for 13 parks (72%), the best-fit models conform to the expectation of an inverted u-shape relationship between the expected mean monthly temperature and visitation (rosselló-nadal 2014), indicating that above a certain mean monthly temperature threshold, tourism demand decreases. for the five coastal parks, which are in year-round (garden route national park) and winter-rainfall areas (agulhas, namaqua, table mountain and west coast national parks), and where mean monthly temperatures are moderated by cool onshore winds, the relationship between temperature and occupancy is positive and linear for the range of mean monthly temperatures experienced up to now. these are not wildlife parks and tend to be more popular in summer when visitors can enjoy beach activities. the linearity of the relationship suggests that mean monthly temperatures have not yet reached the levels above which tourists would be deterred. with the exception of namaqua national park, these parks are also all located in close proximity to large urban areas and receive a larger proportion of day visitors than overnight visitors owing to the extensive alternative accommodation available in close proximity. this study has not captured the impact of climate change on day visitors as park accommodation occupancy levels were used. day visitors would be more likely to have more flexibility in timing their visits to the parks for the most comfortable parts of the day and might therefore exhibit a different relationship. the park-specific model predictions of monthly occupancy rates were out by less than 10% on average during the estimation period for all but west coast national park. for five parks (augrabies falls, bontebok, kruger, mokala and tankwa karoo national parks) rmse values improved in the validation period compared with the estimation period, with the remaining parks having marginally worse rmse values during the validation period, an expected result given that the models were generated by minimising the sum of the squares in the difference between the observed and predicted values, by using the estimation period data. in some cases, and west coast national park specifically, the rmse values were worse during the validation period because overnight capacity had been increased during the period, resulting in observed occupancy rates in the validation period being lower than predicted for a number of months as demand lagged in its reaction to a change in supply. for the parks for which temperature and occupancy exhibit a quadratic relationship, it was possible to calculate the turning points, or the temperature thresholds, above which occupancy rates started to decline. these turning points varied substantially among the individual parks, and many occurred at relatively mild temperatures. it should be noted that these effectively reflect average monthly temperatures, rather than daily temperatures. thus, a mean monthly temperature of 25 °c could be interpreted as having a relatively high probability of encountering daily temperatures over 30 °c. in cases such as golden gate national park, which experiences relatively low year-round temperatures and where occupancy was mostly positively related to temperature, the modelled threshold of 14 °c seemed too low to be a plausible threshold. indeed, the direct relationship to temperature was fairly weak in the case of this park, and little seasonality in visitation rates. given that visitors tend to avoid the coldest months because of the extremely low overnight temperatures, an increase in temperature might start to deter visitors in the hotter months but might also increase occupancy in the lower months. in the case of the hot arid parks (richtersveld, tankwa karoo and augrabies national parks) which experience very high temperatures and where occupancy was mostly negatively related to temperature, the data and thresholds (15.8 °c and 16.5 °c for the first two, and undefined for the third) reflected that the parks are fullest in their cooler but not coldest months. overall increases in mean monthly temperatures would be expected to lead to overall decreases in visitor bookings but could be averted by increasing the available accommodation so that more people can visit in the coolest months. the other arid zone park, kgalagadi national park, experiences slightly more moderate conditions and had a higher turning point of 18.2 °c. this park is most popular in spring and autumn to avoid the extreme cold nights and extreme hot days of winter and summer, respectively. for the parks in the summer rainfall areas of the savanna biome (the ‘bushveld’), temperatures can reach uncomfortable levels in the summer months, which are also the wet months. wildlife viewing also tends to be better in the dry winter months when there is less foliage and grass, and when animals tend to congregate at watering points. therefore, it is unsurprising that there is a steeply negative association between occupancy and temperature in kruger and mapungubwe national parks. as with the desert parks, higher overall temperatures would be expected to shift demand to the cooler months, leading to reduced numbers unless park capacity is increased. in karoo national park, where visibility is less of a factor, there is less of an association with temperature, apart from a steep drop off in occupancy in the hottest months. in bontebok national park and the eastern cape parks (addo, camdeboo and mountain zebra national parks), which experience relatively moderate temperatures and relatively little seasonality in their fynbos, karoo and thicket vegetation cover, visitors tend to increase in warmer months, apart from the very hottest months (figure 2). increasing temperatures would be expected to shift demand to cooler months, and this could be better accommodated than in parks with an inverse relationship with temperature. thus, the impact of a degree rise in temperature would be expected to be lower for these parks than for the bushveld and arid zone parks. there was also an interesting positive relationship between the turning point threshold and overall occupancy level of parks. bearing in mind that occupancy levels are not a measure of park popularity per se (which is reflected in bed numbers), higher occupancy levels would be expected to reflect more consistent rates of occupancy throughout the year (such as for karoo national park), or periods of the year with very high occupancy (such as the flagship kruger and addo elephant national parks). these parks have more to lose with an increase in temperature by virtue of their already high turning points, which more closely align with comfort levels and the fact that they are likely to be close to their environmental capacity in terms of accommodating visitors. the results of the empirical analysis are important for exploring and estimating the impact that climate change may have on tourism demand for individual parks and at the park system level, especially if tourism is a major financial contributor to the park system’s operational budget as is the case in south africa (novellie 2018). applying climate change predictions for the period 2040–2060 to these models suggested that the majority of parks in south africa (61%) will see an increase in occupancy. this is in line with the explanations above and with other international studies (e.g. fisichelli et al. 2015; jones & scott 2006). this is largely attributed to these parks moving up the occupancy–temperature curve for most months of the year and these increases in occupancy outweighing the decreases experienced during the hotter months of the year. it must be noted that the estimated change is indicative only as, although the extent that future predicted mean monthly temperatures exceed the range of modelled data is marginal for most of the parks, tourism demand may respond differently to these unobserved novel climates at specific locations. for instance, tipping points in temperature may be reached that have not been observed at a specific location in the past, resulting in changes to tourism demand that cannot be modelled. this could well be the case for the coastal parks that currently exhibit a linear relationship between occupancy and temperature but which, under higher predicted future temperatures, may reach a tipping point above which occupancy begins to fall. although the impact on the entire park system is negative (~4% decline by 2050), if the impact on tourism demand for kruger national park was excluded from the analysis, the result would be an overall increase for the park system in the order of ~2%. the projected impact on kruger national park is of special concern because the revenue generated by the park helps subsidise the rest of the park system, and a decline in tourism revenue would put the capacity for conservation activities at risk (novellie 2018). given the high density of beds in this park, there is little option for increasing capacity to accommodate more tourists in the cooler months without leading to congestion and ecological impacts. understanding the potential impact of climate change on tourism demand is important for park planning and management, as it helps to focus adaptation considerations. for example, if a protected area manager were to see that his/her protected area is vulnerable to a decrease in tourism demand because climate conditions may make it unbearable to visit the park during certain periods of the year, then this should inspire adaptation considerations around how to make it more bearable for tourists, such as installing air conditioning in the accommodation units or installing a swimming pool so tourists can cool off during the hotter times of the day. dube and nhamo (2019) found that tourism operators in kruger national park are already considering and implementing strategies to deal with the observed changes in tourist preferences and responses to unbearable temperatures, including green building design to ensure guest comfort. alternatively, considerations around increasing tourism capacity to take advantage of tourists’ climate preferences should be explored, with consideration given to congestion and ecological constraints. these results also have important bearings on sanparks’ marketing strategy with regard to timing and targeting of directed marketing campaigns to increase occupancy and target groups who are perhaps less sensitive to expected temperatures. implementing adaptation strategies will place a further burden on constrained national park budgets in south africa. although useful for identifying potential climate change impacts on tourism demand owing to tourists’ climate preferences, the empirical models are unable to capture the climate change impact on destination attractions that influence the destination decision of tourists, such as the change in snowfall, hydrology, bleaching of reefs, species and vegetation changes, etc. furthermore, the empirical models do not consider higher levels of domestic tourism in key origin countries owing to more favourable conditions closer to home (rosselló & santana 2012; tol & walsh 2012) or the rising aversion to long-haul flights. for these reasons, tourism operators should consider both a change in tourism comfort levels and the potential climate impacts on attractions when estimating how climate change may influence tourism demand. we suggest that further research that extends the analysis to a larger number of parks and broader climatic area, as well as to include visitor surveys, would be useful to further explore the role of variables such as rainfall and accommodation characteristics, as well as to validate the findings on visitor motivations. conclusion tourists to south africa’s national parks are sensitive to temperature, but only six of the parks assessed are predicted to experience a decline in occupancy by 2050, as movements up the quadratic temperature curves during the coldest months of the year outweigh movements down the temperature curves during the hottest months of the year. despite this, the predicted unmitigated impact of climate change on total tourism demand across the national park network is for a 4% decline, largely owing to the roughly 6.5% decline for kruger national park outweighing the predicted changes for the other 18 parks. these results are of concern given that tourism proceeds generated by kruger national park are used to subsidise the operating budgets of the entire park system, which would put the capacity for conservation activities at risk, suggesting that measures to alleviate climate discomfort to tourists should be explored by management. acknowledgements we are grateful to south african national parks for provision of tourism statistics, and to two anonymous referees for their inputs on an earlier draft. competing interests the authors have declared that no competing interests exist. authors’ contributions all authors contributed equally to this work. funding information this research received no specific grant from any funding agency in the public, commercial or not-for-profit sectors. data availability statement data sharing is not applicable to this article as no new data were created or analysed in this study. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references abegg, b. & koenig, u., 1997, ‘impacts of climate change on winter tourism in the swiss alps’, journal of sustainable tourism 5(1), 46–58. https://doi.org/10.1080/09669589708667275 amelung, b. & viner, d., 2006, ‘mediterranean tourism: exploring the future with the tourism climate index’, journal of sustainainable tourism 14(4), 349–366. https://doi.org/10.2167/jost549.0 amelung, b., nicholls, s. & viner, d., 2007, ‘implications of global climate change for tourism flows and seasonality’, journal of travel research 45(3), 285–296. https://doi.org/10.1177/0047287506295937 belle, e.m.s., burgess, n.d., misrachi, m., arnell, a., masumbuko, b., somda, j. et al., 2016, climate change impacts on biodiversity and protected areas in west africa, summary of the main outputs of the parcc project, protected areas resilient to climate change in west africa, p. 62, unep-wcmc, cambridge. bigano, a., hamilton, j. & tol, r., 2007, ‘the impact of climate change on domestic and international tourism: a simulation study’, integrated assessment journal 7(1), 25–49. https://doi.org/10.2139/ssrn.907454 biggs, d., swemmer, l., phillips, g., stevens, j., freitag, s. & grant, r., 2014, ‘the development of a tourism research framework by south african national parks to inform management’, koedoe 56(2), 1–9. https://doi.org/10.4102/koedoe.v56i2.1164 chown, s.l., 2010, ‘temporal biodiversity change in transformed landscapes: a southern african perspective’, philosophical transactions of the royal society b biological sciences 365(1558), 3729–3742. https://doi.org/10.1098/rstb.2010.0274 collier, p., conway, g. & venables, t., 2008, ‘climate change and africa’, oxford review of economic policy 24(2), 337–353. https://doi.org/10.1093/oxrep/grn019 davis-reddy, cl. & vincent, k., 2017, climate risk and vulnerability: a handbook for southern africa 2nd edn., pp. 202, csir, pretoria. de freitas, cr., 2003, ‘tourism climatology: evaluating environmental information for decision making and business planning in the recreation and tourism sector’, international journal of biometeorology 48, 45–54. https://doi.org/10.1007/s00484-003-0177-z dube, k. & nhamo, g., 2018, ‘climate variability change and potential impacts on tourism: evidence from the zambian side of the victoria falls’, environmental science & policy 84, 113–123. https://doi.org/10.1016/j.envsci.2018.03.009 dube, k. & nhamo, g., 2019, ‘climate change and potential impacts on tourism: evidence from the zimbabwean side of the victoria falls’, environmental develpoment and sustainability 21(4), 2025–2041. https://doi.org/10.1007/s10668-018-0118-y dube, k. & nhamo, g., 2020, ‘evidence and impact of climate change on south african national parks. potential implications for tourism in the kruger national park’, environmental development, 33, 100485. https://doi.org/10.1016/j.envdev.2019.100485 dube, k., mearns, k., mini, s. & chapungu, l., 2018, ‘tourists’ knowledge and perceptions on the impact of climate change on tourism in okavango delta, botswana’, african journal of hospitality, tourism and leisure 7(4), 1–18. dunlop, m. & brown, p., 2008, implications of climate change for australia’s national reserve system: a preliminary assessment, report to the department of climate change, and the department of the environment, water, heritage and the arts by csiro sustainable ecosystems, p. 155, csiro australia, canberra. fisichelli, n.a., schuurman, g.w., monahan, w.b. & ziesler, p.s., 2015, ‘protected area tourism in a changing climate: will visitation at us national parks warm up or overheat?’, plos one 10(6), e0128226. https://doi.org/10.1371/journal.pone.0128226 fitchett, j.m., robinson, d. & hoogendoorn, g., 2017, ‘climate suitability for tourism in south africa’, journal of sustainable tourism 25(6), 851–867. https://doi.org/10.1080/09669582.2016.1251933 giddy, j.k., fitchett, j.m. & hoogendoorn, g., 2017, ‘insight into american tourists’ experiences with weather in south africa’, bulletin of geography, socio-economic series 38(38), 57–72. https://doi.org/10.1515/bog-2017-0034 glick, p. stein, b.a. & edelson, n.a., 2011, scanning the conservation horizon: a guide to climate change vulnerability assessment, p. 176, national wildlife federation, washington, dc. hamilton, j.m., maddison, d.j. & tol, r.s., 2005, ‘effects of climate change on international tourism’, climate research 29(3), 245–254. https://doi.org/10.3354/cr029245 harris, i., osborn, t.j., jones, p. & lister, d., 2020, ‘version 4 of the cru ts monthly high-resolution gridded multivariate climate dataset’, scientific data 7(1), 1–18. https://doi.org/10.1038/s41597-020-0453-3 hoogendoorn, g. & rogerson, c.m., 2015, ‘tourism geography in the global south: new south african perspectives’, south african geographical journal 97(2), 101–110. https://doi.org/10.1080/03736245.2015.1034574 jones, b. & scott, d.j., 2006, ‘climate change, seasonality and visitation to canada’s national parks’, journal of recreation administration 24(2), 42–62. koenig, n. & bischoff, e.e., 2004, ‘seasonality research: the state of the art’, working paper 2004/3, european business management school, university of wales, swansea, viewed 21 august 2005, from http://www.swan.ac.uk/ebms/research/working%20papers/ebms%202004%203.pdf. kozak, m., 2002, ‘comparative analysis of tourist motivations by nationality and destinations’, tourism management 23(3), 207–20. https://doi.org/10.1016/s0261-5177(01)00090-5 lemieux, c.j., beechey, t.j., scott, d.j. & gray, p.a., 2010, protected areas and climate change in canada: challenges and opportunities for adaptation, canadian council on ecological areas (ccea) occasional paper no. 19, pp. xii + 170, ccea secretariat, ottawa, ontario. lin, t.p. & matzarakis, a., 2008, ‘tourism climate and thermal comfort in sun moon lake, taiwan’, international journal of biometeorology 52(4), 281–290. https://doi.org/10.1007/s00484-007-0122-7 lise, w. & tol, r.s.j., 2002, ‘impact of climate on tourist demand’, climatic change 55(4), 429–449. https://doi.org/10.1023/a:1020728021446 macfadyen, s., hui, c., verburg, p.h. & van teeffelen, a.j., 2019, ‘spatiotemporal distribution dynamics of elephants in response to density, rainfall, rivers and fire in kruger national park, south africa’, diversity and distribution 6(25), 880–894. https://doi.org/10.1111/ddi.12907 maddison, d., 2001, ‘in search of warmer climates? the impact of climate change on flows of british tourist’, climatic change 49, 193e208. https://doi.org/10.1023/a:1010742511380 markham, a., osipova, e., lafrenz samuels, k. & caldas, a., 2016, world heritage and tourism in a changing climate, united nations environment programme, nairobi, and united nations educational, scientific and cultural organization, paris, pp. 18–28. mathivha, f.i., tshipala, n.n. & nkuna, z., 2017, ‘the relationship between drought and tourist arrivals: a case study of kruger national park, south africa’, jàmbá: journal of disaster risk studies 9(1), 1–8. https://doi.org/10.4102/jamba.v9i1.471 mieczkowski, z., 1985, ‘the tourism climatic index: a method of evaluating world climates for tourism’, canadian geographer/le géographe canadien 29(3), 220–233. https://doi.org/10.1111/j.1541-0064.1985.tb00365.x moore, w.r., 2010, ‘the impact of climate change on caribbean tourism demand’, current issues in tourism 13(5), 495–505. https://doi.org/10.1080/13683500903576045 mushawemhuka, w., rogerson, j.m. & saarinen, j., 2018, ‘nature-based tourism operators’ perceptions and adaptation to climate change in hwange national park, zimbabwe’, bulletin of geography socio-economic series 42(42), 15–127. https://doi.org/10.2478/bog-2018-0034 novellie, p.a., 2018, ‘review of national park science: a century of research in south africa by jane carruthers’, koedoe 60(1), a1550. https://doi.org/10.4102/koedoe.v60i1.1550 pandy, w.r., 2017, ‘tourism enterprises and climate change: some research imperatives’, african journal of hospitality, tourism and leisure 6(4), 1–18. pandy, w.r. & rogerson, c.m., 2018, ‘tourism and climate change: stakeholder perceptions of at risk tourism segments in south africa’, euro economica 37(2), 104–118. perch-nielsen, s.l., amelung, b. & knutti, r., 2010, ‘future climate resources for tourism in europe based on the daily tourism climatic index’, climatic change 103(3–4), 363–381. https://doi.org/10.1007/s10584-009-9772-2 perry, a., 2006, ‘will predicted climate change compromise the sustainability of mediterranean tourism?’, journal of sustainainable tourism 14(4), 367–375. https://doi.org/10.2167/jost545.0 pike, s., 2002, ‘destination image analysis – a review of 142 papers from 1973 to 2000’, tourism management 23(5), 541–549. https://doi.org/10.1016/s0261-5177(02)00005-5 preston-whyte, r.a. & watson, h.k., 2005, ‘nature tourism and climatic change in southern africa’, in c.m. hall & j. higham (eds.), tourism, recreation and climate change, pp. 130–142, channelview, bristol. r core team, 2014, r: a language and environment for statistical computing, r foundation for statistical computing, r package version 1.1–7, viewed from http://cran.r-project.org/package=lme4. rogerson, c.m., 2016, ‘climate change, tourism and local economic development in south africa’, local economy 31(1–2), 322–331. https://doi.org/10.1177/0269094215624354 roshan, g., yousefi, r. & fitchett, j.m., 2015, ‘long-term trends in tourism climate index scores for 40 stations across iran: the role of climate change and influence on tourism sustainability’, international journal of biometeorology 60, 33–52. https://doi.org/10.1007/s00484-015-1003-0 rosselló, j. & santana, m., 2012, climate change and global international tourism: an evaluation for different scenarios, working paper: departament d’economia aplicada, universitat de les illes balears, palma, p. 16. rosselló-nadal, j., 2014, ‘how to evaluate the effects of climate change on tourism’, tourism management 42, 334–340. https://doi.org/10.1016/j.tourman.2013.11.006 sanford, t., frumhoff, p.c., luers, a. & gulledge, j., 2014, ‘the climate policy narrative for a dangerously warming world’, nature climate change 4(3), 164–166. https://doi.org/10.1038/nclimate2148 scott, d. & mcboyle, g., 2001, ‘using a ‘tourism climate index’ to examine the implications of climate change for climate as a tourism resource’, first international workshop on climate, tourism and recreation, pp. 69–88, international society of biometeorology, porto carras. scott, d., gossling, s. & freitas, c.r., 2007, ‘climate preferences for tourism: an exploratory tri-nation comparison’, in a. matzarakis, c.r. de freitas & d. scott (eds.), developments in tourism climatology, pp. 18–23, commission on climate, tourism and recreation, freiburg. scott, d., gössling, s. & hall, c.m., 2012, ‘international tourism and climate change’, wiley interdisciplinary reviews: climate change 3(3), 213–232. https://doi.org/10.1002/wcc.165 scott, d., mcboyle, g., minogue, a. & mills, b., 2006, ‘climate change and the sustainability of ski-based tourism in eastern north america: a reassessment’, journal of sustainable tourism 14(4), 376–398. https://doi.org/10.2167/jost550.0 scott, d., rutty, m., amelung, b. & tang, m., 2016, ‘an inter-comparison of the holiday climate index (hci) and the tourism climate index (tci) in europe’, atmosphere 7(6), 80. https://doi.org/10.3390/atmos7060080 smith, t. & fitchett, j.m., 2020, ‘drought challenges for nature tourism in the sabi sands game reserve in the eastern region of south africa’, african journal of range & forage science 37(1), 107–117. https://doi.org/10.2989/10220119.2019.1700162 steyn, j.n. & spencer, j.p., 2012, ‘climate change and tourism: implications for south africa’, african journal for physical, health education, recreation and dance 18(1), 1–19. sutherland, k., ndlovu, m. & pérez-rodríguez, a., 2018, ‘use of artificial waterholes by animals in the southern region of the kruger national park, south africa’, south african journal of wildlife research 48(2), 023003. https://doi.org/10.3957/056.048.023003 tol, r.s.j. & walsh, s., 2012, ‘climate change and tourism in the arctic circle’, working paper series 5212, department of economics, university of sussex business school, falmer, viewed 10 july 2019, from https://ideas.repec.org/p/sus/susewp/5212.html. van der merwe, p. & saayman, m., 2008, ‘travel motivations of tourists visiting kruger national park’, koedoe 50(1), 54–159. https://doi.org/10.4102/koedoe.v50i1.140 van wilgen, n.j., goodall, v., holness, s., chown, s.l. & mcgeoch, m.a., 2016, ‘rising temperatures and changing rainfall patterns in south africa’s national parks’, international journal of climatology, 36(2), 706–721. https://doi.org/10.1002/joc.4377 filelist convert a pdf file! page 1 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 abstract introduction methods results discussion acknowledgements references about the author(s) kathryn r. mccollum school of natural resources, university of nebraska-lincoln, united states emily belinfonte warnell school of forestry and natural resources, university of georgia, united states april l. conway warnell school of forestry and natural resources, university of georgia, united states john p. carroll school of natural resources, university of nebraska-lincoln, united states citation mccollum, k.r., belinfonte, e., conway, a.l. & carroll, j.p., 2018, ‘occupancy and habitat use by six species of forest ungulates on tiwai island, sierra leone’, koedoe 60(1), a1484. https://doi.org/10.4102/koedoe.v60i1.1484 short communication occupancy and habitat use by six species of forest ungulates on tiwai island, sierra leone kathryn r. mccollum, emily belinfonte, april l. conway, john p. carroll received: 04 aug. 2017; accepted: 12 july 2018; published: 29 aug. 2018 copyright: © 2018. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract forest ungulates in west africa are common bushmeat species and are subject to habitat degradation through deforestation. based on historical data, there are possibly 12 species of forest bovidae and tragulidae found in eastern sierra leone. we used camera trapping to assess occupancy by forest ungulates on and around a small protected area, tiwai island, sierra leone. we then assessed habitat over two field seasons during 2008–2011 for those species where we had sufficient numbers of detections. we detected 6 of 12 potential species and obtained enough data to further assess the habitat of two species. species detected included the black duiker (cephalophus niger), bongo (tragelaphus eurycerus), bushbuck (tragelaphus scriptus), maxwell’s duiker (philantomba maxwellii), water chevrotain (hyemoschus aquaticus) and yellow-backed duiker (cephalophus silvicultor). among detected species, the bongo is considered near threatened. several of the species not detected might be extirpated from the region, but for several species we found no records of them in the area. for the two species with sufficient detections for analysis, we found that maxwell’s duikers were common throughout woody and swamp habitat and yellow-backed duikers preferred old growth forests with open understory. despite widespread deforestation in sierra leone, a recent civil war and continued bushmeat trade, it appears that small wildlife refuges such as tiwai island continue to provide sanctuary for many of the forest ungulates of the region. conservation implications: the guinea rainforest ecosystem of west africa has undergone significant human impact and deforestation, negatively impacting all aspects of the biodiversity of the region. in addition, a long-standing civil war in sierra leone further exacerbated conservation concerns of many wildlife species. there are some recognised reserves in sierra leone, but small reserves managed by local people and conservation organisations have a role to play. our work on tiwai island, along the moa river in sierra leone, demonstrated that a significant proportion of the forest dwelling ungulate biodiversity of the region has been maintained in a small reserve despite isolation and effects of the war. our work also suggests that tiwai island continues to have significant ecological value for ungulate conservation in the region and should be considered a model for establishment of other small reserves to help maintain the region’s biodiversity. introduction the upper guinea forests of western africa contain some of the most diverse animal species on earth and are global hotspots for conservation (klop, lindsell & siaka 2008). several species of ungulates are endemic or near endemic to this area. the westernmost block of the upper guinea forest stretches into sierra leone, and although there are some protected areas, such as gola forest national park, forest ungulates are experiencing decreasing population trends within the area (international union for conservation of nature [iucn] 2015). few studies have been conducted on duikers and other forest ungulates of western africa, although they are a major food source for villagers in many rural forested areas (anadu, elamah & oates 1988; newing 2001). several species may inhabit the same habitat with little interspecies competition by isolating themselves temporally, spatially and by body size (newing 2001). there are 12 forest ungulate species that could possibly be found in the upper guinea forest of sierra leone: maxwell’s duiker, black duiker, water chevrotain, yellow-backed duiker, bushbuck, bongo, zebra duiker (cephalophus zebra), jentink’s duiker (cephalophus jentinki), red-flanked duiker (cephalophus rufilatus), ogilby’s (brooke’s) duiker (cephalophus ogilbyi), african forest buffalo (syncerus caffer nanus) and bay duiker (cephalophus dorsalis). even though all 12 species are found within forested habitats, some are much more adaptable to human encroachment, hunting and deforestation. in this research, we will collectively refer to them as ‘forest ungulates’. duikers and other forest ungulates are commonly utilised for bushmeat in rural areas throughout guinea rainforest, bringing food and an economic benefit to people of all social statuses (anadu et al. 1988; newing 2001; ripple et al. 2016; wilkie & carpenter 1999). domestic livestock have difficulty surviving in the humid, dense rainforests of sierra leone and guinea, and in some areas the taste of bushmeat is preferred to that of domestic bovids (anadu et al. 1988). maxwell’s duikers are a particularly common species for the bushmeat trade in the past few decades and are regarded favourably by consumers (anadu et al. 1988). however, it is unclear how development and alteration of the landscape affects many ungulate species commonly used for bushmeat. the purpose of this study was to determine occupancy of the tiwai island region by forest ungulates. to focus on bovid-related species, the red river hog (potamochoerus porcus), although observed, were excluded from this study (mccollum et al. 2016). methods study area our study was conducted on and around the tiwai island wildlife sanctuary in southeastern sierra leone (figure 1). tiwai has an area of 12 km2 and is located on the moa river, including two islands, upper tiwai and lower tiwai (conway 2013; oates et al. 1990). the climate is wet and warm, typical of western guinean rain forest, with an average annual rainfall of 3000 mm. a distinguishable dry season takes place between december and march in which the temperature averages 30 °c. the island has tropical vegetation consisting of old forests, regenerating secondary forests and swamps (fimbel 1994; oates et al. 1990). the island is managed as an ecotourism site with a non-government organisation with proceeds contributed back to surrounding host villages. this encourages them to cooperate with conservation priorities of the area rather than clearing it for farming or exploiting other resources (conway 2013). figure 1: tiwai island wildlife sanctuary, lower tiwai, and the surrounding small islands in eastern sierra leone, surveyed using camera traps during 03 november 2008–22 february 2009 and 10 may 2010–23 july 2011. camera traps and occupancy analysis camera traps were employed to detect duikers and forest ungulates because of their efficiency in collecting data on elusive or shy species (swann et al. 2004). to select locations at which cameras were installed, the island was mapped out in fifteen 1 km2 grids (figure 1). within each grid section, a moultrie game spy camera was placed at a random location and set up to take two photographs (with a 10 second delay) upon trigger by movement. the random locations were selected using a geographic information system (arcgis) and cameras were attached to trees facing down trails at or near the selected point. there were two field seasons: 03 november 2008–22 february 2009 and 10 may 2010–23 july 2011. the first season and six of the periods in the second season were conducted on upper tiwai, while two of the periods in the second season were conducted on lower tiwai. for field season 1, the cameras were set up for 14 days before relocation to another location. for field season 2, the cameras were employed for 21 days before relocation. species of duikers and ungulates were identified from the images captured by cameras, and each species was recorded as detection (1) or non-detection (0). data were combined into two 7-day periods for season 1, and three 7-day periods for season 2, in which one or more detections within the 7 days warranted presence in the period. on the lower island, there were delays retrieving the cameras, which continued to collect data for 10–12 days after the end of the third period. we extended the detection time frame to three periods in season 2 to potentially capture more rare events. although possibly making the models more robust, we controlled for possible detection bias by analysing the data with season as a covariate. for species with sufficient detection, data were modelled using program presence for analysis, and the success of the habitat parameters was compared. five habitat covariates were used in modelling occupancy: the understory density (cover), understory type (understory), canopy type (canopy), whether the camera was on a human, animal or no trail (trails) and the standardised distance to the moa river (distmoa). covariates had multiple parameters: cover was classified as open, semi-dense or dense; understory was woody, swamp, riparian or bush fallow; canopy was young secondary, mature, swamp, riparian or bush fallow; and trails were human trail, animal trail or no trail. we measured the strength of our models using the akaike’s information criterion (aic), which represents the goodness-of-fit of the models. the models were compared using the change in the aic value (δ aic); the best-fitting model had a δ aic of 0.00, with the value increasing for less-fitting models. all models with a δ aic less than 4.00 were considered a good fit. we then calculated the occupancy probability and 95% confidence interval of each parameter. results we detected 6 of the 12 potential ungulate species on tiwai island wildlife sanctuary and vicinity over two seasons (838 and 2474 camera-trap days, for seasons 1 and 2, respectively) (table 1). a total of 280 ungulate photographs were recorded at 100 of 180 sample locations on upper tiwai, lower tiwai and the smaller surrounding islands. of the five habitat covariates, there were insufficient detections of any species in bush fallow habitat; therefore, that habitat was omitted from further analysis. table 1: potential and observed species of bovidae and tragulidae found on and in the vicinity of tiwai island, sierra leone, using camera traps during 2008–2011. maxwell’s duikers were the most commonly trapped with 175 photographs, or 5 detections per 100 trap nights. yellow-backed duikers were recorded 55 times or 2 detections per 100 trap nights. water chevrotains had 40 total detections or 1 detection per 100 trap nights. there were eight photographs of bushbucks, 0.24 detections per 100 trap nights. bongo and black duiker were each detected once during the study, less than 0.03 detections per 100 trap nights (table 1). because of the insufficient detections, water chevrotain, bushbuck, black duiker and bongo were excluded from habitat analysis. maxwell’s duiker in season 1, naïve occupancy estimate (ψ) was 0.30. occupancy of maxwell’s duiker was influenced by both the distance to the moa river and understory type (table 2). in particular, the distance from the moa river, strongly affected occupancy of maxwell’s duikers, β = 7.87. maxwell’s duikers appeared to prefer swamp and riparian understory, with ψ = 0.83 (±0.50 standard error [se]) (table 3). occupancy was also high in woody habitat, with ψ = 0.77 (±0.35 se). table 2: occupancy analysis of maxwell’s and yellow-backed duikers determined from camera trapping data on tiwai island and vicinity, sierra leone, during 03 november 2008–22 february 2009 and 10 may 2010–23 july 2011. table 3: duiker occupancy of specific tiwai island habitat parameters for best-fitting models (δ aic < 4) used in analysis. in season 2, the naïve ψ was 0.50. occupancy of maxwell’s duikers was associated with canopy cover and understory type (table 2). swamp cover and young secondary growth were similarly preferred, with swamp having ψ = 0.75 (±0.28 se) and young secondary growth having ψ = 0.71 (±0.09 se). mature canopy was also occupied often, with ψ = 0.65 (±0.11 se). there were too few detections of maxwell’s duiker in riverine habitat for analysis. as observed in season 1, maxwell’s duikers preferred swamp habitat, with ψ = 0.83 (±0.25 se). occupancy of woody habitat was common, with ψ = 0.69 (±0.08 se). riparian habitat was not commonly occupied, with ψ = 0.28 (±0.17 se) (table 3). yellow-backed duiker we did not have enough detections of yellow-backed duikers in any habitat during season 1 to run working models. however, in season 2, we obtained a naïve occupancy rate of 0.26. the best-fitting model of season 2 was understory type, although the only significant parameter was woody habitat with ψ = 0.70 (±0.21 se) (table 2). occupancy of riverine or swamp habitat by yellow-backed duiker was infrequent with too few detections to assess. occupancy was also influenced by the distance to the moa river, β = 0.56. yellow-backed duikers greatly preferred open cover, with ψ = 0.92 (±0.34 se) (table 3). semi-dense cover occupancy was also common and had ψ = 0.51 (±0.24 se). dense understory cover was not preferred, with ψ = 0.30 (±0.23 se). discussion forest ungulates can be secretive and solitary animals which make them difficult to detect and estimate abundance. camera trapping provides access to these species and can assist in cost-effective conservation efforts in regions where dramatic changes in land use and other human activity might be rapidly impacting species. species detection our detection of half of the potential species that might be found in this region suggests that despite significant negative impacts of a long civil war, chronic deforestation in the region and continued bushmeat trade, at least some species are apparently quite resilient. those species not detected might simply have been undetected, are extirpated or are marginal in distribution for the region or habitat. based on known habitat, some species, such as forest buffalo; and zebra, bay, ogilby’s and jentinks’ duikers may have historically occurred in this area but were not detected during our study. red-flanked duikers are more of a savanna edge species, but were included because they are found to the north of our study area, and might actually respond to deforestation in the region. in addition, we had very few detections of some important species, such as bongo and black duiker, which may have persisted at low numbers following the civil war or are recolonising the region from protected areas associated with the gola forest and further from liberia. water chevrotains had one of the largest numbers of detections, suggesting that they are commonly found in this protected area. they are not as commonly found in bushmeat markets as some of the duikers, but if larger species continue to be overharvested, hunters may turn to the smaller chevrotains (anadu et al. 1988). we had very low detections of bushbucks, suggesting that they are not commonly found in the study area, although they are common in other parts of their range. bushbucks are commonly hunted for bushmeat in sierra leone, and the species could be more threatened in western africa than in the rest of its range, where they are often not hunted owing to superstitious beliefs (anadu et al. 1988). we had two detections of black duiker. very little is known about populations and activity of this secretive species, as they are often mistaken for other species (klop et al. 2008). oates et al. (1990) noted the absence of bongo on tiwai island prior to the civil war; however, we detected only a single young male in our study. this observation highlights the value of camera trapping as we had no personal observation of bongo in 4 years of research on the island. habitat use our camera trapping survey demonstrated that maxwell’s and yellow-backed duikers occupy a variety of habitats. riverine, woody and swamp habitats were similarly utilised during season 1, whereas in season 2 swamp was occupied most often, followed by woody habitat. riverine habitat was occupied infrequently in season 2. this suggests that while they do use all three habitats, they may use them for different needs. detections of maxwell’s duikers also changed dramatically during our camera trapping periods, suggesting that longer-term studies may be needed to study them fully. yellow-backed duikers did not occupy the islands as extensively as maxwell’s duikers, and appeared more specific in their habitat preferences. they were detected in open woody, understory or old forests. decreases in populations of yellow-backed duikers throughout this area could occur as human impacts on land use continue to meet increasing demands for food. detection rates of yellow-backed duikers varied throughout the second season, suggesting that they utilise both tiwai island and the mainland, and perhaps often travel between the two. long-term studies may be needed for this species to fully understand their habitat usage and range. as human activity continues to change natural landscapes, more species are losing suitable habitat and facing increased hunting pressure, resulting in decreasing populations. bushmeat hunting is unlikely to diminish in the future unless other sustainable food sources are established. illegal poaching makes it difficult to enforce hunting regulations, especially as the majority of poaching occurs in less accessible areas. in some areas, farming of wild meat species, such as the cane rat, has become an alternative to unregulated snaring and shooting. however, this management practice poses concerns for solitary species that do not reproduce often. cryptic species are difficult to study and provide a particular challenge when trying to manage for conservation. acknowledgements we would like to thank the communities of tiwai island for their support, as well as the tiwai island administrative committee and the ministry of agriculture forestry and food security for granting us permission to work on tiwai island. finally, we would like to acknowledge the hard work of our local field assistants, kenewa koroma, bockary koroma, alusine koroma and mohamed lahai. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions k.r.m. analysed data and wrote the manuscript. e.b. summarised photos and wrote the first draft of the manuscript. a.l.c. designed the study and carried out field research. j.p.c. supervised the research, analysed data and wrote the manuscript. funding information funding for the main project was provided by conservation international, tulsa zoo conservation grant, minnesota zoo ulysses s. seal conservation grant, oregon zoo future for wildlife grants program, columbus zoo conservation fund, riverbanks zoo conservation support fund, okc zoo conservation action now, idea wild, the young explorer’s fund and the iucn hippopotamus specialist group. additional funding was provided by the warnell school of forestry and natural resources, university of georgia and the school of natural resources, university of nebraska. references anadu, p., elamah, p. & oates, j.f., 1988, ‘the bushmeat trade in southwestern nigeria: a case study’, human ecology 16, 199–208. https://doi.org/10.1007/bf00888092 conway, a.l., 2013, ‘conservation of the pygmy hippopotamus in sierra leone’, phd dissertation, university of georgia, athens, ga. fimbel, c., 1994, ‘the relative use of abandoned farm clearings and old forest habitats by primates and a forest antelope at tiwai, sierra leone, west africa’, biological conservation 70, 277–286. https://doi.org/10.1016/0006-3207(94)90173-2 international union for conservation of nature (iucn), 2015, the iucn red list of threatened species. version 2015–4, viewed 07 march 2016, from http://www.iucnredlist.org klop, e., lindsell, j. & siaka, a., 2008, biodiversity of gola forest, sierra leone, royal society for the protection of birds, conservation society of sierra leone, and government of sierra leone, sandy, uk, freetown, sierra leone. mccollum, k.r., conway, a.l., lee, m.b. & carroll, j.p., 2016, ‘occupancy and demographics of red river hog potamochoerus porcus on tiwai island, sierra leone’, african journal of ecology 55, 47–55. https://doi.org/10.1111/aje.12314 newing, h., 2001, ‘bushmeat hunting and management: implications of duiker ecology and interspecific competition’, biodiversity conservation 10, 99–118. https://doi.org/10.1023/a:1016671524034 oates, j.f., whitesides, g.h., davies, a.g., waterman, p.g., green, s.m., dasilva, g.l. et al., 1990, ‘determinants of variation in tropical forest primate biomass: new evidence from west africa’, ecology 71, 328–343. https://doi.org/10.2307/1940272 ripple, w.j., abernathy, k., betts, m.g., chapron, g., dirzo, r., galetti, m. et al., 2016, ‘bushmeat hunting and extinction risk to the world’s mammals’, royal society open science 3(10), 160498. https://doi.org/10.1098/rsos.160498 swann, d.e., hass, c.c., dalton, d.c. & wolf, s.a., 2004, ‘infrared-triggered cameras for detecting wildlife: an evaluation and review’, wildlife society bulletin 32, 357–365. https://doi.org/10.2193/0091-7648(2004)32[357:icfdwa]2.0.co;2 wilkie, d.s. & carpenter, j.f., 1999, ‘bushmeat hunting in the congo basin: an assessment of impacts and options for mitigation’, biodiversity conservation 8, 927–955. https://doi.org/10.1023/a:1008877309871 acknowledgements references about the author(s) llewellyn c. foxcroft scientific services, south african national parks, skukuza, south africa centre for invasion biology, department of botany and zoology, stellenbosch university, cape town, south africa citation foxcroft, l.c., 2020, ‘koedoe: changing of the guard’, koedoe 62(1), a1645. https://doi.org/10.4102/koedoe.v62i1.1645 editorial koedoe: changing of the guard llewellyn c. foxcroft copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. in a recent essay marking the 60th anniversary of koedoe, carruthers and foxcroft (2019) reflected on the history of the journal. when koedoe was first instituted by south african national parks (sanparks) in 1958, research in sanparks was still in its infancy and the journal was considered a novel and bold approach to integrating protected area management with research in africa. since then, koedoe has enjoyed sustained support, and the readership has grown exponentially. koedoe has published over 1,250 articles, from a wide range of fields (figure 1, reprinted from foxcroft & herbst 2017), covering more than 60 protected areas across south africa and africa (figure 2, reprinted from foxcroft & herbst 2017). over the last decade it became increasingly important to expand the scope of koedoe beyond the biophysical to include research areas that were less commonly published in koedoe but crucial to conservation. these included subjects like tourism, adaptive management and social-ecological sciences. koedoe is well placed to play an important role in publishing relevant and robust science for the benefit of conservation broadly. figure 1: bubble plot showing the most frequently listed words in article titles, analysed per decade. the size of the circle indicates the frequency of keywords as an indication of active fields of research. figure 2: bubble plot showing the most frequently listed national parks or other research sites in article titles, analysed per decade. the size of the circle indicates the frequency of the protected area name as an indirect indication of key focus areas over time. it was within this setting that in 2007 i took up position as the seventh editor of koedoe, setting out to continue where my predecessors left off and raise koedoe into the foreground of 21st-century protected area research. my first priority was to increase access to koedoe via an online publishing platform, opting for open access since much of koedoe’s readership, across academia, conservation agencies and other research facilities, does not have access to expensive journal subscriptions. in addition to academic use, open access ensures conservation practitioners can apply credible science to practical challenges being faced. this increased visibility of the journal from the traditional print run of just under 1,000 copies to, for example, a total of about 196 000 articles being downloaded from april 2019 to march 2020. there are, of course, challenges with open-access publishing, but we are constantly looking at ways to address these. secondly, i aimed to ‘get koedoe rated’ with a clarivate journal citation reports impact factor. although it took a few years to be realised, i am pleased to say that koedoe has had an impact factor rating since 2013, and i am confident that it will increase with time. in the highly competitive world of science and scholarly publishing, journals need to evolve and bring in new ideas and perspectives. with time, editors and scientists also reach a stage where their focus needs to shift to take on new challenges. with this in mind, after much thought and consultation, koedoe is in the process of embracing a new era of editorial leadership. after 13 years as editor-in-chief, i have decided to hand over the reins to dr abel ramoelo. dr ramoelo is a regional ecologist in sanparks with a background in remote sensing, who brings with him experience as associate editor from various international remote sensing journals. dr ramoelo will continue to be supported by dr marna herbst, who has been associate editor since 2016. i will take up a position on the koedoe editorial advisory board and will play an active advisory role where necessary, especially during the transition period as dr ramoelo assumes management of koedoe. it has been a privilege and an exciting challenge to have played a part in maintaining koedoe as a knowledge source for future generations. i wish dr ramoelo the very best as he takes over and leads koedoe onward, knowing he has a great support team surrounding him. acknowledgements there are too many people to whom i owe a debt of gratitude to be listed here, but i greatly appreciate the support by everyone involved. i do wish to acknowledge south african national parks (sanparks), and especially thank peter novellie, stefanie freitag-ronaldson and danie pienaar, for providing the opportunity to lead koedoe. the editorial advisory board has provided me with invaluable advice, which was greatly appreciated. thanks to aosis (pty) ltd for their role in the evolution of koedoe and especially to those with whom i have worked closely while managing koedoe. thanks also to marna herbst for the substantial support over the last 4 years. references carruthers, j. & foxcroft, l.c., 2019, ‘koedoe: african protected area conservation and science – a retrospection: 1958 to 2018’, koedoe 61(1), a1556. https://doi.org/10.4102/koedoe.v61i1.1556 foxcroft, l.c. & herbst, m., 2017, ‘status and trends in the global growth of koedoe between 1958 and 2016’, koedoe 59(1), a1500. https://doi.org/10.4102/koedoe.v59i1.1500 abstract introduction methods and materials results discussion conclusion acknowledgements references about the author(s) brianna m. lind plant and environmental sciences department, new mexico state university, new mexico, united states andrew b. davies department of global ecology, carnegie institution for science, california, united states citation lind, b.m. & davies, a.b., 2019, ‘a checklist of the termites of kruger national park, south africa’, koedoe 61(1), a1531. https://doi.org/10.4102/koedoe.v61i1.1531 checklist a checklist of the termites of kruger national park, south africa brianna m. lind, andrew b. davies received: 14 mar. 2018; accepted: 12 nov. 2018; published: 09 apr. 2019 copyright: © 2019. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the protection of biodiversity is critical to ecosystem function and is a primary management goal for conservation areas globally. maintaining a current inventory of known diversity is a central component of achieving this goal and serves as an essential starting point for future research endeavours. since the first published survey of termites in south africa’s kruger national park (knp) over 55 years ago, our understanding of termite diversity has expanded sufficiently to merit an update and formal checklist. here we revise the inventory of termite diversity in knp and summarise the taxonomic and functional diversity of termites in the park. a thorough review of recent termite research in knp added 6 new genera and 13 species to what was found in coaton’s original survey, with one genus, anenteotermes, recorded for the first time in southern africa. based on the updated species checklist, the majority of genera in the park belong to feeding group ii (39%) and the termitidae family (75%). conservation implications: in savannas, termites play crucial roles in nutrient cycling, water redistribution and plant dynamics. systematically cataloguing termite diversity and assemblage composition in the park provides an essential baseline for scientific research, aids biodiversity conservation efforts and encourages scientists and managers to consider termites in ecosystem functioning and management. having more detailed descriptions of genera, species and feeding groups allows for more tangible, ecologically relevant attributions of termite influence, facilitates enhanced inquiry and allows for more realistic quantification of termite roles in key ecosystem processes. keywords: termite; kruger national park; feeding groups; diversity; savanna. introduction a principal goal of most conservation areas is to conserve biodiversity in all its facets. to achieve this goal, scientists must first catalogue the known diversity of organisms to serve as a baseline for future conservation and research endeavours. this is particularly pertinent for groups that perform important ecological functions, such as termites. termites are a prominent source of habitat heterogeneity in african savannas and have a substantial impact on ecosystem functioning (jouquet et al. 2011; sileshi et al. 2010). termites are the dominant invertebrate decomposers in tropical forests and savannas (collins 1981; schuurman 2005), and, through their foraging activities, alter soil processes, redistribute nutrients and modify soil hydrology (jouquet et al. 2011). these ecosystem alterations in turn affect vegetation distributions and assemblage composition (davies, baldeck & asner 2016a; joseph et al. 2014; sileshi et al. 2010), resulting in a patchy landscape that alters both mammalian and insect herbivore foraging behaviour (davies et al. 2016b, 2016c; grant & scholes 2006; levick et al. 2010a). considering its latitude, southern africa has a surprisingly high diversity of termites, with over 50 genera present (eggleton 2000; uys 2002). the termite fauna of the region is generally considered to be well documented as a result of the national survey of isoptera conducted by w.g.h. coaton and co-workers between 1956 and 1979. during this survey, almost every quarter-degree grid cell of south africa, swaziland, zimbabwe and namibia was sampled for termites. much of the information gained from this survey appeared in a series of publications in cimbebasia between 1971 and 1980 (uys 2002), and collected specimens are currently located in the south african national collection of insects (uys 2002). kruger national park (knp), south africa, was also included in these surveys, with a checklist of the park’s termite fauna published in 1962 (coaton 1962). however, since this time, a considerable amount of termite research has been conducted in knp, warranting an update of the checklist in order to have a comprehensive and current inventory of termite diversity in the park. in his survey, coaton collected 22 termite genera from four families within knp (coaton 1962). while the last few decades have seen a large increase in the number of studies investigating termite ecology in knp, none have focused exclusively on termite taxonomy or sought to update the checklist, despite new genus and species records. here we synthesise the termite occurrence records from published studies in knp and compile an updated list of termite richness and functional diversity to facilitate better understanding of knp’s biodiversity, which will aid both management and future research endeavours. by characterising the park’s termite assemblages, a fuller understanding of the fundamental roles termites play in critical ecosystem processes within kruger’s savannas can also be realised. methods and materials kruger national park is located in low-lying savanna in the northeast corner of south africa between the foot slopes of the drakensberg escarpment and mozambican coastal plains (venter, scholes & eckhardt et al. 2003). the region experiences high temperatures in the summer, mild winters and the majority of rainfall between october and april (ranging from ~300 mm/year in the north to ~700 mm/year in the south) (venter et al. 2003). the park has two main underlying geologies, basalt in the east and granite in the west (venter et al. 2003). along with varying rainfall, these geologies contribute to a diverse range of soil types and influence vegetation patterns (venter et al. 2003). tree canopy cover ranges from 5% to 60%, and 75% of the park is covered by broad-leaved savanna with the remaining 25% composed of fine-leaved savanna (venter et al. 2003). we conducted a comprehensive literature review of research involving termites in knp by searching web of science and google scholar and compiled a list of termite genera and species sampled from each study. termites were listed by family, subfamily, genus and species and categorised into feeding groups (fg) following donovan, eggleton and bignell (2001) to characterise the assemblage. feeding groups consist of four categories (i–iv) based on the level of decay of food items, which ranged from live plants to decayed organic material. donovan et al. (2001) defines fgs to include a non-termitid group, fg-i, that feeds on dead wood and grass and three termitid groups that eat dead wood, litter and micro-epiphytes, fg-ii, humus, fg-iii, and soil minerals, fg-iv. results our literature review of 20 termite-related studies conducted in knp (see table 1) revealed that since coaton’s 1962 survey, six previously unlisted termite genera and 13 species have been sampled in the park (see table 2). in addition, one sampled genus, anenteotermes, had not been previously recorded in southern africa. three studies in particular contributed to checklist updates: meyer et al. (1999); davies et al. (2012); and davies et al. (2014a). while termites from all southern african families are present in the park, the majority (75%) belong to the termitidae family (see figure 1a). in terms of fgs, fg-ii makes up 40% of termite genera in the park, fg-i and fg-iii each make up 25% and fg-iv, which is the least represented, makes up 10% (see figure 1b). figure 1: summary of the families (a) and feeding groups (b) of termites sampled in kruger national park, south africa. table 1: history of published, termite-focused research conducted in the kruger national park, south africa. table 2: checklist of termites known to occur in the kruger national park, south africa. discussion termite ecology in knp, and in savannas globally, has advanced substantively since coaton’s 1962 survey. our literature review revealed 6 previously unrecorded genera and 13 previously unrecorded species in knp. termite diversity in knp appears to mirror the generalised diversity pattern of termites in southern africa, with relatively few genera in the hodotermitidae, kalotermitidae and rhinotermitidae families, with the majority of termite diversity being found within the termitidae family (uys 2002). as knowledge about how termites affect the structure and function of savanna ecosystems continues to accumulate, it will be increasingly important to understand how different species and genera interact with and affect the environment. maintaining an accurate record of the termites that occur in knp is a critical first step to achieving this understanding and provides an essential baseline for ongoing research. there were two major research papers that contributed most significantly to updating the inventory of knp’s taxonomic diversity: meyer et al. (1999), and davies et al. (2012). meyer et al. (1999) made substantial contributions to advancing our understanding of termite diversity in knp through research that involved mapping the distribution and density of termite mounds in northern kruger (meyer et al. 1999, 2000a). meyer et al. (1999) documented macrotermes natalensis as the most abundant mound-building termite species in this region, which led to several follow-up studies on the species, including an investigation of intracolonial demography (meyer et al. 2000), biomass (meyer et al. 2001) and food consumption (meyer et al. 2003). while meyer’s research into m. natalensis provides an excellent baseline for the ecological influence of this termite species in northern knp, future research should strive to account for the full breath of foraging power (i.e. litter removal) that all fg-ii termites effect in knp. when fg-ii termites are evaluated collectively, we may find that litter removal by termites competes with burning and herbivory and could in turn subsequently influence fire regimes and patterns. while meyer et al. (1999) increased our knowledge of mound-building termites, davies et al. (2012) employed a broad range of sampling strategies that captured the full range of termite families in the park. davies et al. (2012) sampled termites across varying burning regimes across knp and found that termites are broadly resilient to fire. they found that fg-ii termites are dominant across knp’s rainfall gradient with the caveat that the proportion of other fgs changed with rainfall, with the greatest fg diversity in the wettest region, pretoriuskop. this overall finding is similar to our synthesis, where we also found a dominance of fg-ii termites in knp. davies et al. (2012) also documented a first record of the soil-feeding genus anenteotermes (sands 1998) in knp and in southern africa. although this genus is known to range from semi-arid savanna to rainforests (sands 1998), davies et al. (2012) were the first to document the genus in southern africa. anenteotermes has previously been recorded from as far south as malawi (donovan et al. 2002) and is also well represented in central (e.g. deblauwe et al. 2008; eggleton et al. 1996) and east (e.g. wanyonyi et al. 1984) africa. although we have added numerous additions to coaton’s original 1962 checklist, many of the specimens he sampled still await identification. as such, our ‘new’ findings compiled from recent research may be a consequence of a more comprehensive identification process rather than of sampling in a new environment. this highlights the lack of taxonomic resolution of termites (especially non-mound-building, soil-feeding termites) in africa. indeed, almost a third of termite genera in southern africa are in need of taxonomic revision (uys 2002). because listing and categorising taxa is such an important component of conserving biodiversity and understanding ecosystems, a priority of future termite research should be to resolve these taxonomic difficulties (wilson 2017). deo-xyribonucleic acid metabarcoding, in conjunction with taxonomic and natural history data components, may facilitate enhanced taxonomic clarity and enable more species-level identifications. conclusion collectively, termites as an insect group play considerably important roles in the functioning of dryland and subtropical environments. maintaining a termite checklist in knp allows us to begin the important work of attributing what we understand as ‘termite influence’ to specific termite species and genera. it also provides an invaluable baseline for future research in the park and facilitates the implementation of biodiversity conservation, a key park management goal. the ecological significance and conservation value of insects in savannas has traditionally received little attention relative to vertebrates (braack & kryger 2003). knowledge of termite diversity and assemblage composition will hopefully help bridge this gap and facilitate the consideration of this important insect group in ecosystem functioning and management planning. moving forward, assessing termite diversity in ecologically relevant fgs, that have targeted and discernible ecosystem impacts, will advance research endeavours and allow for more realistic quantification of their roles in key processes such as vegetation dynamics, nutrient cycling, litter removal and fuel-load management. acknowledgements the authors would like to thank dr niall hanan for insightful comments on an earlier version of the manuscript. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions b.m.l. and a.b.d. designed the study, analysed the data and wrote the article. references braack, l.e.o., 1995, ‘seasonal activity of savanna termites during and after severe drought’, koedoe 38, 73–82. https://doi.org/10.4102/koedoe.v38i1.307 braack, l.e.o. & kryger, p., 2003, ‘insects and savanna heterogeneity’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83–129, island press, washington, dc. coaton, w.g.h., 1962, ‘survey of the termites of the kruger national park’, koedoe 5, 144–168. https://doi.org/10.4102/koedoe.v5i1.842 coetsee, c., jacobs, s. & govender, n., 2012, ‘an overview of nitrogen cycling in a semiarid savanna: some implications for management and conservation in a large african park’, environmental management 49, 387–402. https://doi.org/10.1007/s00267-011-9779-0 collins, n.m., 1981, ‘populations, age structure, and survivorship of colonies of macrotermes bellicosus (isoptera: macrotermitinae)’, journal of animal ecology 50, 293–311. https://doi.org/10.2307/4046 davies, a.b., baldeck, c.a. & asner, g.p., 2016a, ‘termite mounds alter the spatial distribution of african savanna tree species’, journal of biogeography 43, 301–313. https://doi.org/10.1111/jbi.12633 davies, a.b., eggleton, p., van rensburg, b.j. & parr, c.l., 2012, ‘the pyrodiversity–biodiversity hypothesis: a test with savanna termite assemblages’, journal of applied ecology 49, 422–430. https://doi.org/10.1111/j.1365-2664.2012.02107.x davies, a.b., eggleton, p., van rensburg, b.j. & parr, c.l., 2013, ‘assessing the relative efficiency of termite sampling methods along a rainfall gradient in african savannas’, biotropica 45, 474–479. https://doi.org/10.1111/btp.12030 davies, a.b., eggleton, p., van rensburg, b.j. & parr, c.l., 2015, ‘seasonal activity patterns of african savanna termites vary across a rainfall gradient’, insectes sociaux 62, 157–165. https://doi.org/10.1007/s00040-014-0386-y davies, a.b., levick, s.r., asner, g.p., robertson, m.p., van rensburg, b.j. & parr, c.l., 2014a, ‘spatial variability and abiotic determinants of termite mounds throughout a savanna catchment’, ecography 37, 852–862. https://doi.org/10.1111/ecog.00532 davies, a.b., levick, s.r., robertson, m.p., van rensburg, b.j., asner, g.p. & parr, c.l., 2016b, ‘termite mounds differ in their importance for herbivores across savanna types, seasons and spatial scales’, oikos 125, 726–734. https://doi.org/10.1111/oik.02742 davies, a.b., robertson, m.p., levick, s.r., asner, g.p., van rensburg, b.j. & parr, c.l., 2014, ‘variable effects of termite mounds on african savanna grass communities across a rainfall gradient’, journal of vegetation science 25, 1405–1416. https://doi.org/10.1111/jvs.12200 davies, a.b., van rensburg, b.j., robertson, m.p., levick, s.r., asner, g.p. & parr, c.l., 2016c, ‘seasonal variation in the relative dominance of herbivore guilds in an african savanna’, ecology 97, 1618–1624. https://doi.org/10.1890/15-1905.1 deblauwe, i., dibog, l., missoup, a.d., dupain, j., van elsacker, l., dekoninck, w. et al., 2008, ‘spatial scales affecting termite diversity in tropical lowland rainforest: a case study in southeast cameroon’, african journal of ecology 46, 5–18. https://doi.org/10.1111/j.1365-2028.2007.00790.x donovan, s.e., eggleton, p. & bignell, d.e., 2001, ‘gut content analysis and a new feeding group classification of termites’, ecological entomology 26, 356–366. https://doi.org/10.1046/j.1365-2311.2001.00342.x donovan, s.e., eggleton, p. & martin, a., 2002, ‘species composition of termites of the nyika plateau forests, northern malawi, over an altitudinal gradient’, african journal of ecology 40, 379–385. https://doi.org/10.1046/j.1365-2028.2002.00397.x eggleton, p., 2000, ‘global patterns of termite diversity’, in t. abe & m. higashi (eds.), termites: evolution, sociality, symbioses, ecology, pp. 25–51, springer, netherlands. eggleton, p., bignell, d.e., sands, w.a., mawdsley, n.a., lawton, j.h., wood, t.g. et al., 1996, ‘the diversity, abundance and biomass of termites under differing levels of disturbance in the mbalmayo forest reserve, southern cameroon’, philosophical transactions of the royal society of london b 351, 51–68. https://doi.org/10.1098/rstb.1996.0004 grant, c.c. & scholes, m.c., 2006, ‘the importance of nutrient hot-spots in the conservation and management of large wild mammalian herbivores in semi-arid savannas’, biological conservation 130, 426–437. https://doi.org/10.1016/j.biocon.2006.01.004 joseph, g.s., seymour, c.l., coetzee, b.w.t., ndlovu, m., de la torre, a., suttle, r. et al., 2016, ‘microclimates mitigate against hot temperatures in dryland ecosystems: termite mounds as an example’, ecosphere 7, 1–10. https://doi.org/10.1002/ecs2.1509 joseph, g.s., seymour, c.l., cumming, g.s., cumming, d.h. & mahlangu, z., 2014, ‘termite mounds increase functional diversity of woody plants in african savannas’, ecosystems 17, 808–819. https://doi.org/10.1007/s10021-014-9761-9 jouquet, p., traoré, s., choosai, c., hartmann, c. & bignell, d., 2011, ‘influence of termites on ecosystem functioning. ecosystem services provided by termites’, european journal of soil biology 47, 215–222. https://doi.org/10.1016/j.ejsobi.2011.05.005 levick, s.r., asner, g.p., chadwick, o.a., khomo, l.m., rogers, k.h., hartshorn, a.s. et al., 2010b, ‘regional insight into savanna hydrogeomorphology from termite mounds’, nature communications 1, 1–7. https://doi.org/10.1038/ncomms1066 levick, s.r., asner, g.p., kennedy-bowdoin, t. & knapp, d.e., 2010a, ‘the spatial extent of termite influences on herbivore browsing in an african savanna’, biological conservation 143, 2462–2467. https://doi.org/10.1016/j.biocon.2010.06.012 meyer, v.w., braack, l.e.o. & biggs, h.c., 2000, ‘distribution and density of cubitermes wasmann (isoptera: termitidae) mounds in the northern kruger national park’, koedoe 43, 57–65. https://doi.org/10.4102/koedoe.v43i1.208 meyer, v.w., braack, l.e.o., biggs, h.c. & ebersohn, c., 1999, ‘distribution and density of termite mounds in the northern kruger national park, with specific reference to those constructed by macrotermes holmgren (isoptera: termitidae)’, african entomology 7, 123–130. meyer, v.w., crewe, r.m. & braack, l.e.o., 2003, ‘estimates of food consumption by the fungus-growing termite macrotermes natalensis in a south african savanna-woodland: news and views’, south african journal of science 99, 207–208. meyer, v.w., crewe, r.m., braack, l.e.o., groeneveld, h.t. & van der linde, m.j., 2000, ‘intracolonial demography of the mound-building termite macrotermes natalensis (haviland)(isoptera, termitidae) in the northern kruger national park, south africa’, insectes sociaux 47, 390–397. meyer, v.w., crewe, r.m., braack, l.e.o., groeneveld, h.t. & van der linde, m.j., 2001, ‘biomass of macrotermes natalensis in the northern kruger national park, south africathe effects of land characteristics’, sociobiology 38, 431–448. sands, w.a., 1998, the identification of worker castes of termite genera from soils of africa and the middle east, cab international, wallingford, uk. schuurman, g., 2005, ‘decomposition rates and termite assemblage composition in semiarid africa’, ecology 86, 1236–1249. https://doi.org/10.1890/03-0570 sileshi, g.w., arshad, m.a., konaté, s. & nkunika, p.o., 2010, ‘termite induced heterogeneity in african savanna vegetation: mechanisms and patterns’, journal of vegetation science 21, 923–937. https://doi.org/10.1111/j.1654-1103.2010.01197.x sponheimer, m., lee-thorp, j., de ruiter, d., codron, d., codron, j., baugh, a.t. et al., 2005, ‘hominins, sedges, and termites: new carbon isotope data from the sterkfontein valley and kruger national park’, journal of human evolution 48, 301–312. https://doi.org/10.1016/j.jhevol.2004.11.008 uys, v.m., 2002, a guide to the termite genera of southern africa, plant protection research institute handbook no. 15, agricultural research council, pretoria. venter, f.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation pattern’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83–129, island press, washington, dc. wanyonyi, k., darlington, j.p.e.c. & bagine, r.k.n.b., 1984, ‘checklist of the species of termite (isoptera) recorded from east africa’, journal of the east africa natural history society and national museum 181, 1–12. wilson, e.o., 2017, ‘biodiversity research requires more boots on the ground’, nature ecology & evolution 1, 1590. https://doi.org/10.1038/s41559-017-0360-y abstract introduction study area methods results discussion conclusion acknowledgements references about the author(s) bayanda sonamzi centre for functional biodiversity, school of life sciences, university of kwazulu-natal, pietermaritzburg, south africa school of biology and environmental sciences, mpumalanga university, nelspruit, south africa matthew burnett centre for functional biodiversity, school of life sciences, university of kwazulu-natal, pietermaritzburg, south africa robin petersen scientific services, kruger national park, skukuza, south africa gordon o’brien centre for functional biodiversity, school of life sciences, university of kwazulu-natal, pietermaritzburg, south africa school of biology and environmental sciences, mpumalanga university, nelspruit, south africa colleen t. downs centre for functional biodiversity, school of life sciences, university of kwazulu-natal, pietermaritzburg, south africa citation sonamzi, b., burnett, m., petersen, r., o’brien, g. & downs, c.t., 2020, ‘assessing the effect of tagging and the vulnerability to predation in tigerfish (hydrocynus vittatus, castelnau 1861) in a water-stressed system using telemetry methods’, koedoe 62(1), a1649. https://doi.org/10.4102/koedoe.v62i1.1649 original research assessing the effect of tagging and the vulnerability to predation in tigerfish (hydrocynus vittatus, castelnau 1861) in a water-stressed system using telemetry methods bayanda sonamzi, matthew burnett, robin petersen, gordon o’brien, colleen t. downs received: 13 july 2020; accepted: 07 oct. 2020; published: 17 dec. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract increase in anthropogenic pressures on freshwater ecosystems, globally and locally in south africa, has made it difficult to meet environmental flow requirements and maintain these systems. the letaba-olifants catchment is one such example, where the upstream water demands place pressure on the river downstream as it flows through the kruger national park. we used the activity rates of hydrocynus vittatus as a line of evidence to assess (1) the effects of telemeter tagging on their activity rates in three potential post-tagging recovery periods and (2) their vulnerability to predation and the causality related to the environmental stressors placed on the river catchment in the olifants river gorge, kruger national park, south africa. we determined h. vittatus activity rates as locomotive movement using radio telemetry methods linked remotely to an online data management system. we telemeter tagged nine individuals from 08 may to 28 june 2018. however, only seven fish were successfully tracked for the duration of our study, and two individuals moved out of range of the remote network shortly after release and could not be located. the tagged h. vittatus individuals were found to have reduced activity at least within the first 7 days after tagging compared with the time after that. the results showed that three individuals were preyed on by predators after the tagging procedure recovery period. this coincided with abnormal low flow conditions, where the letaba river ran dry. african fish eagles haliaeestus vocifer were the only confirmed predator, whilst predation by other species was also evident. conservation implications: monitoring of h. vittatus using telemetry is a viable tool to use when assessing environmental stressors in remote locations. the abnormally low water levels in the olifants and letaba rivers (cessation of flow in the letaba river) during our study may have compromised the predator avoidance strategies of tagged h. vittatus and may further be affecting the viability of the h. vittatus populations in kruger national park. the synergistic effects of natural and anthropogenic stressors impact negatively h. vittatus populations and potentially other aquatic biotas. keywords: fish behaviour; conservation; ecological indicators; environmental stressors; e-flows; fish recovery. introduction fish are well established around the world as indicators of ecological integrity (harrison & whitfield 2006; mims & olden 2013; sosa-lópez et al. 2005). similar to other aquatic organisms, fish species face several challenges from the environment they occur in and are subject to a plethora of stressors, both natural and anthropogenic (du plessis 2019; o’brien et al. 2018). the ever-increasing anthropogenic pressures have generally led to a steady decline in aquatic organisms, including fish species (dudgeon 2014; rodell et al. 2018). fish kills and population declines are often reported worldwide as a result of deteriorating aquatic ecosystems (ferreira & pienaar 2011; thronson & quigg 2008). this deterioration has been attributed to several factors, including river fragmentation, flow modification, pollution, the introduction of alien species and global climate change (de moor 1996; ellender & weyl 2014; o’brien et al. 2019). fish make use of the available physical habitat as refugia, for feeding and spawning, and as nurseries for their offspring (godin 1997; o’brien et al. 2013). maintaining these environments is essential, but they are often inaccessible and difficult to sample, making observations hard to determine (hussey et al. 2015; lennox et al. 2017). field-based studies are becoming increasingly important because of their holistic approach to measure animal behaviour and movements attributed to the environment they inhabit (cooke & shcreer 2003; cooke et al. 2017). techniques such as capture-mark-re-capture (cmr) and environmental monitoring generally lack the representation of aquatic organisms’ ecological interactions and fine-scale movements within their environments (cooke et al. 2017). increasingly telemetry techniques are being used to determine various stressors of aquatic organisms in the wild and their interactions with other organisms (burnett et al. 2020; cooke et al. 2004; thorstad et al. 2013). telemetry techniques are employed to monitor fish within their natural environment and can provide the required information needed to understand fish behaviour and fine-scale movements (burnett et al. 2020; wisniewska et al. 2016). consequently, this technique has become the preferred method to study fish behaviour worldwide (cooke et al. 2013; hussey et al. 2015; lennox et al. 2017). recent developments in telemetry studies have presented opportunities for better management actions to safeguard the sustainability of natural resources (abecasis et al. 2018; block et al. 2016; cooke et al. 2017). fish telemetry studies involve the attachment of electronic tags, and it has been shown that these affect the initial behaviour of tagged fish following tagging procedures (økland et al. 2005). this has revealed that the predation of tagged fish is more likely to occur directly after the tagging procedure (beland, kocik & sheehan 2001; halfyard et al. 2017; schultz et al. 2017; thorstad et al. 2003). in southern africa, there have been reports of african fish-eagles, haliaeetus vocifer, preying on telemeter tagged fish shortly after release (burnett et al. 2020; thorstad et al. 2003). furthermore, the movement of telemeter tagged individuals does not always reflect normal behaviour until several days after tagging procedure, possibly the reason for the predation of individuals shortly after release (thorstad et al. 2003). the predation effect on adult fish is generally unknown and can be exacerbated when using telemetry methods (bolland et al. 2019; moxham et al. 2019; thorstad et al. 2003). despite the above, the effects of telemeter tagging of fish are regarded as relatively minimal compared with the amount of information that is obtained through telemetry studies (jepsen et al. 2002, 2004). the effect of predation, and other mortality accounts, after recovery from the telemeter tagging procedure, has largely been neglected, with mortality increasingly becoming an important consideration in aquatic telemetry studies (klinard & mately 2020). these effects have been known primarily during the recovery period post-telemeter tagging procedure; however, predation thereafter could be higher than previously considered because of natural causes and environmental stressors (jacobs 2017; thorstad et al. 2003). anthropogenic changes to the environment take precedence in threatening local populations of fish within south africa and elsewhere (dudgeon et al. 2006; rodell et al. 2018; woodborne et al. 2012). in recent years, there were increased studies within the kruger national park, particularly in the olifants river, to understand the biological and ecological effects of anthropogenic stressors on various fish species and the nile crocodile crocodylus niloticus (ashton 2010; bouwman et al. 2014; de villiers & mkwelo 2009; ferreira & pienaar 2011; smit et al. 2013). moreover, telemetry has successfully been conducted on tigerfish hydrocynus vittatus within southern africa to determine their biology and ecology (jacobs et al. 2019; o’brien et al. 2012; roux et al. 2018). these studies noted some interference of aerial predation on h. vittatus; however, none have specifically addressed the predation issue on h. vittatus post-telemeter tagging nor during recovery. the tigerfish, hydrocynus vittatus, is a relatively large, migratory characin that is known to make extensive use of river reaches (jacobs et al. 2019; roux et al. 2018). their higher trophic level and sensitivity to water temperature and oxygen levels make them good ecological indicators of environmental change in the riverine ecosystems they occur (roux et al. 2018; smit et al. 2013). they are listed as protected in the south african national environmental management of biodiversity act for critically endangered, endangered, vulnerable and protected species (deat 2004). these characteristics make h. vittatus an ideal candidate for telemetry studies (burnett et al. 2020; jacobs et al. 2019; jepsen et al. 2015). in the present study, we used the activity rates of h. vittatus as a line of evidence to assess (1) the effects of telemeter tagging on their activity rates in three potential post-tagging recovery periods and (2) their vulnerability to predation and the causality related to environmental stressors placed on the river catchment in the kruger national park, south africa. we predicted that (1) h. vittatus are affected by the tagging procedure for only a few days and (2) the predation pressure on h. vittatus populations here would be higher than previously perceived. we then highlight the potential causalities of the increase in predation of h. vittatus based on environmental variables measured in the study area and similar environmental impacts elsewhere. the knowledge of natural stressors on h. vittatus populations can contribute to sound decision-making for their conservation and management and their river ecosystems. understanding the impact of predation of fish species may explain the effect these predators have on the taxa and the aquatic environment. study area the letaba river, which originates in the limpopo province of south africa, joins the larger olifants river in the kruger national park and then flows into mozambique (figure 1). our study was conducted around the confluence of the two rivers. the olifants river flows through the gauteng, mpumalanga and limpopo provinces. it is socio-economically important and subjected to anthropogenic factors (e.g. intensive mining, industrialisation, urbanisation and agriculture) in most of its secondary catchments before it enters the kruger national park (de villiers & mkwelo 2009; gerber et al. 2016; rogers & o’keeffe 2003). as a result, a wide range of environmental stressors impact the water quality and quantity entering the kruger national park, ultimately affecting the aquatic biota here (du preez & steyn 1992; gerber et al. 2016; roux et al. 2008). the letaba river has anthropogenic impacts similar to the olifants river that affect the water quality and quantity downstream (rogers & o’keeffe 2003; woodborne et al. 2012). figure 1: the kruger national park within south africa and the positioning of the remote network consisting of base and relay stations. the hydrological gauging weirs at balule and engelhardt on the olifants and letaba rivers, respectively, are shown. methods to track tagged h. vittatus individuals remotely and in real time, we established a remote network of receivers (base [bs] and relay [rs] stations) in the olifants and letaba rivers (figure 1) as per the fish tracking (fishtrac) methodology described by burnett et al. (2020). capturing and tagging we captured nine adult h. vittatus using various techniques and methods that included electro-fishing, angling with artificial lures and the use of seine nets. the specimens were collected from 08 may to 28 june 2018 (table 1). after capture, body weight (where possible) and body length of the individuals were measured and recorded. in cases where this was not possible because of field technicalities, such as electronic scale failure, the body mass of the individuals was estimated using the standard length based on known measured body mass to length ratios, hence the use of an estimate (table 1). according to jepsen et al. (2004), tags weighing less than 2% of the fish body mass were used to ensure that the tag mass did not compromise the mobility of the subjects. suitable individuals were anaesthetised in a water-filled container with a 0.4 millitres (ml)/l concentration of clove oil. we observed for signs of anaesthesia before we commenced with the tagging as described by burnett et al. (2020). these signs included, but were not limited to, reduced fin activities and involuntary upside-down swimming motions that were shown by the individuals (o’brien et al. 2013). after that, we surgically attached wireless wildlife external radio tags (ww series iv) using sterilised hollow surgical needles to thread the tag attachment wire through the muscular tissue below the dorsal fin before removing them and securing the tag (burnett et al. 2018; jacobs, o’brien & smit 2016). an antibiotic (1.0 ml/kilograms [kg] of terramycin® containing oxytetracycline) was administered into the surrounding muscular tissue, followed by the application of wound care gel (aqua vet, veterinary hospital, lydenburg, south africa) to minimise infection and assist with wound recovery (burnett et al. 2020). after the tagging procedure was complete, we released the tagged individuals back into the river at their site of capture after signs of recovery were observed as detailed by burnett et al. (2020). the tagging process from capture to tag attachment took ~< 10 min per fish and at least 30 min was allocated for recovery from anaesthesia before releasing them into the river (at capture site). table 1: summary data for hydrocynus vittatus individuals that were telemeter tagged in the olifants gorge, kruger national park, in the present study. data acquisition and analyses the telemeter tags used house omnidirectional tilt vibration sensors that are programmed to record motion as integer counts per unit time (burnett et al. 2020). motion is an inherent attribute of activity, and as such, these integer counts can be used as measures for locomotive movement recorded as activity rates (burnett et al. 2020; horn et al. 2011). based on this concept, one of the assumptions in this study was that if a tag detects no motion and this persists, the tagged individual is assumed to be dead or the tag has been expelled. we set telemeter tags to record data hourly and transmit the stored data when in range of a station (< 500 metres [m]) to a central data management system (dms). each data point from the telemetry tag recorded the activity rate, water temperature (°c), time, the station that detected the tag and the signal strength. the data were downloaded from the dms and then processed and analysed in ms-excel© and r programming (rstudio team 2020). the data were analysed descriptively using non-parametric tests. we compared our three hypothesised recovery periods using the kruskal–wallis test for significance (mckight & najab 2010). we used principal components analysis (pca) to describe activity groupings immediately after the fish were released accounting for the consecutive days post-tagging procedure (bengraïne & marhaba 2003; nasir et al. 2011; o’brien et al. 2009; van den brink, van den brink & ter braak 2003). the pca allows for the analyses of large, complex data sets and accounts for individual variability in a linear response model relating activity rates against environmental variables and time periods (nasir et al. 2011; skalski, richins & townsend 2018). when activity rates monitored recorded zero, we tracked the respective telemetered tags on foot using a ww mobile receiver connected to an omnidirectional yagi antenna (ww, potchefstroom, south africa) to determine their location and status. we retrieved the telemeter tag where possible. we carried this out in the study area using triangulation, that is, gain reduction between the telemeter tag in consideration and the mobile receiver as detected through the directional antenna (burnett et al. 2020). flow data we acquired the hydrological data from the south african department of water and sanitation (dws) website (http://www.dwa.gov.za/hydrology/verified/hymain.aspx) for the gauging stations at balule weir (17.8 kilometres [km]; dws weir no. b7h026) on the olifants river and engelhardt weir (27 km; dws weir no. b8h018) on the letaba river. ethical consideration all ethical standards were followed for the research and ethical clearance was obtained for the surgical tagging procedure of fish species from the relevant authority before the commencement of the study (university of kwazulu-natal animal ethics permit no. arec/070/017d). permission to sample and conduct the study in the olifants river system was obtained from sanparks. results we captured and tagged a total of nine adult h. vittatus individuals, with a mean (± standard deviation [sd]) standard length of 452 mm ± 35 mm. of these, we successfully tracked and monitored seven (table 1). two tags (hvit 6 and hvit 9) went out of range immediately after release and did not contribute to the data collected (table 1). telemeter tagged h. vittatus were monitored for a minimum of 22 days, with the longest tracked for 156 days (mean ± sd: 66 ± 52 days). the variability in tracked days was because of the mortality of tagged individuals or fish moving out of the network range. the sample size for field studies met the criteria recommended by sequeira et al. (2019), stating that individuals of ≤ 10 should be used for studies that are still at an exploration phase, such as the present study. a typical activity pattern, as displayed by an h. vittatus individual (figure 2), showed diurnal behaviour, with regular higher activity rates during the day when temperatures were warmer. this individual was tagged on 10 may and is indicative of the behaviour after recovery. the individual showed daily diel activity rates as they fluctuated throughout the study showing periods of lower activity rates interspersed with periods of higher activity rates, still maintaining a diurnal diel behavioural pattern. figure 2: the daily behaviour of hydrocynus vittatus as represented by the activity rates of individual hvit3 (table 1) showed typical fluctuations of activity rates, highlighting the diel behavioural pattern between night (non-shaded) and day (shaded). recovery period we compared each of the hypothesised recovery periods (3, 7 and 14 days) with the remainder of the data set, excluding abrupt activity events before the cessation of activity rates where applicable. when the first potential recovery period was examined, there was a significant difference in the median of the activity rates between the data of the first 3 days and the rest of the data set (figure 3). the activity rates of the initial first 3 days were lower than the activity rates after that (table 2). except for hvit5 and hvit7, this difference was statistically significant (p < 0.05; table 2). similarly, for the 7and 14-days recovery periods assessed, all fish activity rates were shown to be significantly different (p < 0.05; table 2), showing a lower activity rate in comparison with the stipulated recovery period (figure 3). the activity rates for all telemeter tagged individuals combined showed an exponential increase in activity rates until the 14-day period (figure 4); after this period, the activity rates fluctuated, indicating normalisation of activity rates for tagged individuals. the fluctuation in activity after 14 days decreased to a level similar to the day 7 post-tagging procedure (figure 4a). this indicated that the effect of the tagging procedure was negligible after 7 days. this was consistent with the pca analyses that showed a change in behaviour post-tagging, highlighting alternative variables affecting the activity rates of fish (figures 4 and 5). the analyses showed that 71% of the individuals (n = 5) exhibited significantly low activity rates during the first 24 h. all individuals showed significantly low activity during the 7and 14-days monitoring periods. figure 3: box and whisker plots comparing the logarithmic activity rates of hydrocynus vittatus individuals during the three potential recovery periods. the activity rates amongst the recovery periods were significantly different from the activity rates following each period (p < 0.05). note: median is denoted by the thick middle line of the box. the upper and lower limits of the box represent the first and third quartiles. figure 4: the logarithmic activity rates post-tagging of all telemetered hydrocynus vittatus individuals in the present study where (a) shows the linear regression relationship and (b) the logarithmic regression relationship. in (a), the blue dotted lines represent the recovery periods tested (3 days, 1 week and 2 weeks after tagging), the red dashed lines highlight when the death of a fish was detected, and the solid red lines indicate when the signal was lost. figure 5: principal components analysis plot explaining 57.9% of variance between the activity rates after the tagging procedure and the time of day across the entire data set of all telemetered hydrocynus vittatus individuals in the present study. the principal components analysis used the activity rates of fish, the time of day (pc1) and the day after the tagging procedure (pc2) to visualise the relationship between these three variables. table 2: a summary of the difference between activity rates between the three hypothesised recovery periods and the rest of the activity rates for the rest of the data set in the present study. predation events on 03 july 2019, tagged individual hvit2 recorded a spike in activity and shortly after that the activity rapidly decreased to zero, indicating cessation of activity and possible mortality (figure 6a). after this event, the telemeter tag also recorded temperatures no longer associated with the aquatic environment. we manually tracked the telemeter tag and found it ~300 m from the riverbank under a feeding roost of an h. vocifer. scattered fish bones surrounded the telemeter tag (figure 7a; personal observation). outputs from individual hvit5 on 04 june were similar to hvit2 and demonstrated changes in temperature and activity rates no longer associated with the aquatic environment and the tagged individual, respectively (figure 6b). manual tracking found the telemeter tag to be ~1 km from the river. figure 6: hydrocynus vittatus activity rates with the change in environmental variables in the present study where (a), (b) and (c) show individuals’ associated tag activity rates (dashed green) shown alongside the tag (grey) and water (black) temperatures (°c), whilst (d) shows verified daily average flows from the department of water and sanitation of south africa at the points (grey bars) at which the predation events took place. figure 7: these photographs depict (a) the recovered telemetry tag (tag32: hvit2) as found by researchers (circled) and (b) no flow on the letaba river downstream of engelhardt weir during the telemeter tag recovery survey. individual hvit8 was monitored for approximately a month until two spikes in the activity rates that happened on 20 july 2018 were recorded (figure 6c). the initial spike brought the telemeter tag out of the aquatic environment for 24 h, and the second spike moved the telemetered tag out of range of the network. these spikes, similar to individuals hvit 2 and hvit 5, indicated some form of a predatory event resulting in mortality of the tagged individual (figure 6c). this predation event took place at night, without removing the fish from the vicinity of the riverbed, with the second activity spike moving the telemetered tag back into the water below detectable limits of the network, given adequate network coverage in the area. the river flow dynamics of the olifants and letaba rivers during the time of tracking and monitoring of h. vittatus are shown in figure 6d. the water discharge in the olifants river was ~14 m3s−1 but decreased steadily from may through to september whilst the letaba river had substantially low flows (~1.4 m3s−1) and eventually ceased to flow, leaving behind scattered pools (figures 6d and 7b). the water discharge for the olifants river at balule weir ranged between 14.10 m3s−1 and 0.78 m3s−1, whilst the letaba river at engelhardt gauging weir received between 1.20 m3s−1 and 0.30 m3s−1 during the study period, drying up entirely in places downstream of the study, as observed during manual monitoring surveys (figures 6d and 7b). discussion in our study, we successfully showed the effects of telemeter tagging on the post-tagging activity rates of h. vittatus. typically, the impact of the tagging procedure on tagged fish species in their natural environment is difficult to measure because of the limited in situ measurements and the inability to provide a control experiment (frank et al. 2009, jacobs et al. 2019). despite this, we showed that the fine-scale variability in the movement was evident between different days in the study. this indicated that the response of h. vittatus was unique per day, and effects from the tagging procedure were no longer reflective from as early as 3 days post-tagging procedure, as shown in our linear model and the pca analyses. this supports jepsen et al.’s (2004) finding that the minimal disturbance in fish behaviour is seen during recovery, and the tagging procedure may only affect behaviour within 7 days post-tagging. knowing that the tagging procedure is regarded as a stressor on the behaviour, the fish will navigate its recovery whilst maintaining daily circadian rhythms whilst facing associated environmental conditions. although none of the individuals were predated with 14 days after the tagging procedure, this period of recovery suggests that telemeter tagged h. vittatus may be susceptible to predation as in other studies (thorstad et al. 2003). the minimum length of the recovery period for fish can be as short as 8 h to a day depending on the species, tagging methodology and the fish species’ environment (jepsen et al. 2002; metcalfe et al. 2016). økland et al. (2005) suggested 7 days, specifically for h. vittatus. our study suggests that recovery can extend up to 14 days, however, but can be seen from as early as 3 days post-tagging procedure. on the contrary, the diel behaviour of tigerfish can return to normalcy within the 24-h period and may be crucial to the recovery of the fish. more recently, jacobs et al. (2016) used upstream and downstream movement of h. vittatus to assess the immediate and long-term effects of telemeter tags and found that the movement of the individuals was predominantly downstream (60%). this was associated with reduced spatial movement within the first 3 days of tagging. as such, tagging procedures have immediate effects (< 3 days after tagging) and minimal in the long term (> 7 days) (jacobs et al. 2016; jepsen et al. 2004). in our study, 2 weeks was shown to fall within the periods of an exponential increase in activity rates; however, fluctuations after this period fell as low as activity rates seen after 7 days. paukert et al. (2001) found similar findings and concluded that activity patterns of individuals could be abnormal to up to 2 weeks following release after tagging. although these studies (jacobs et al. 2016; økland et al. 2005; paukert et al. 2001) used manual monitoring techniques, in our study, the activity rates showed that daily circadian rhythms were reflective of normal behaviour from within 3 days post-tagging. the fine-scale observations using activity rates in our study were conclusive. the activity rates of h. vittatus were not only reflective of their full recovery within 2 weeks of tagging but could also be indicative of their need to maintain biological processes (feeding and resting) as seen in the diel behavioural activity patterns. this supports the use of h. vittatus as suitable candidates for telemetry studies (økland et al. 2005) and can be used to understand the response of the fish to environmental variables (burnett et al. 2020). the present study also used the activity rates of h. vittatus as a line of evidence to assess their vulnerability to predation and the causality related to flows and predatory events in inland aquatic ecosystems. of the seven h. vittatus that were successfully monitored, three were removed by predators after the tagging procedure recovery periods tested. through manual tracking and recovery of radio tags, the african fish eagle (h. vocifer) appeared to be a key predator of adult h. vittatus. this was supported by the activity rates for individuals hvit2 and hvit5 and their change in environments from the freshwater to terrestrial environments. during the tag recovery exercise, hvit2 and hvit5 tags were found below a feeding roost and surrounded by fish scales and bones, confirming that h. vocifer were the likely predators of h. vittatus. real-time data showed that these events occurred during the day when h. vocifer and other piscivorous birds such as p. haliaetus were active, of which both are known to be diurnal species, the latter being migratory (østnes et al. 2019; stewart et al. 1997). notwithstanding h. vocifer being a common natural predator of h. vittatus, otters (lutrinae family) and nile crocodiles cannot be ruled out as potential predators as they are known to feed on fish (games & moreau 1997; rowe-rowe 1977). as shown by hvit8, a predation event took place at night, resulting in the tag being removed from the water and later returned to below detectable limits of the telemeter tag, given adequate network coverage in the area. the occurrence of this event at night suggests that a mesocarnivore like the cape clawless otter (aonyx capensis) were responsible as they are nocturnal predators, known to be common in the olifants and letaba rivers (pienaar 1964; personal observation). the high presence of piscivorous predators in our study area suggests the high impact of these predators on fish communities. our study suggests that the impact of predation is considerably higher than what was initially thought and should be considered as a factor in disturbances to fish communities. similar studies have shown this using an avian approach in estuaries (cowley, terörde & whitfield 2017), yet little is known about riverine ecosystems where predators are linearly spaced. this study suggests that predation on fish species is not solely avian related and that mammalian and reptilian predation could also be significant and should be considered especially for adult fish. we acknowledge that one of the limitations in our study was the relatively small sample size, which was partly because of the cost of telemeter tags and accessibility to the field sites that limited the ability to increase the sample size. as such, larger sample sizes are recommended in further studies to improve confidence in using activity rates as a line of evidence and understanding the results shown here. in the present study, we were also limited to the use of adult fish because of the mass of the telemeter tag (20 grams [g]); thus, only individuals above 1000 g could be tagged as determined by the 2% tag to body weight ratio (jepsen et al. 2002). despite these restrictions, our findings provide inferred population responses, strengthening the cause and effect data associated with h. vittatus. considering this, the results of our study are useful in the management of water resources, especially for downstream users such as the kruger national park, and can lay a foundation for future use of this technique. the low flow observed in both rivers during the survey is also a matter of concern. one of the immediate results of these low flows, potentially resulting from water abstraction, dams and drought, is the formation of isolated pools with little to no flows (pearce 2018). stears and mccauley (2018) reported that during these low flow and dry periods, the lack of sufficient oxygenated water and removal of nutrients could cause oxygen levels to decrease and nutrients to reach toxic levels. although not measured in the current study, stears and mccauley (2018) reported that in hypoxic conditions oxygen-dependent fish species such as yellowfishes (labeobarbus spp.) tend to swim close to the water surface where oxygen levels are comparatively higher, making them vulnerable to predators. hydrocynus vittatus is known to use the entire water column to adapt to changes in food resources and water quality (o’brien et al. 2014; smit et al. 2013; steyn et al. 1996). these adaptations to altering environmental conditions could increase their exposure to predators as they swim closer to the surface in hypoxic conditions (stears & mccauley 2018). other opportunistic predators such as baboons papio ursinus have been observed predating on stressed fish in these circumstances (stears & mccauley 2018). hydrocynus vittatus are oxygen-dependent species (vignaud et al. 2002) and are likely to be under more pressure during periods of low flow when the river becomes fragmented into isolated pools than other species. they are also known to generally stay within their home ranges (jacobs et al. 2019; økland et al. 2005), with their movement often restricted by in-stream barriers (roux et al. 2018). as such, they are more likely to be affected by low oxygen levels when trapped in isolated pools. the reduction in flows and in-stream barriers (dams and natural waterfalls) on both the olifants and letaba rivers resulted in relatively few opportunities for h. vittatus to move to reaches where flows were favourable. as observed in this study, this traps them in isolated pools until conditions improve or mortality ensures. this was supported by the remote network established for this study, not detecting reach scale movement of tagged fish. the reduction in water flows from both river systems coincided with the predation events, showing that flow stressors increase the risk to predation and the ecological degradation of the ecosystem for h. vittatus. two main in-stream barriers (engelhard and mingerhout weirs) upstream on the letaba river form artificial barriers obstructing already low flow conditions and have no economic or biodiversity value for the lower letaba river system (o’brien et al. 2019). flows beyond these gauging weirs typically go unmonitored, and our study detected no flow in the letaba river whilst engelhard dam recorded minimal flows at < 0.314 m3s−1. the activity rates and predation events coincided with this reduction in the flow and susceptibility of h. vittatus to hypoxic conditions (smit et al. 2013). this highlights the importance of maintaining environmental flows into the olifants gorge. the discrepancy in the flows between engelhard weir and the olifants gorge suggests that this weir cannot meet environmental flows downstream and questions the ability of the current infrastructure to adequately monitor and meet the environmental flow requirements for the olifants and letaba rivers. with this, the engelhardt wier makes for a good case to be removed as it jeopardises the biodiversity in the letaba river as with other such in-stream barriers (o’brien et al. 2019). all-inclusive real-time and remote monitoring techniques have been developed to minimise the loss in river connectivity and monitor rivers, and these can continue to be implemented in the olifants-letaba catchment (burnett et al. 2020; pollard et al. 2012). conclusion telemetry methods should, where possible, have larger sample sizes and make use of smaller and lighter tags (< 20 g air mass) with proportionally less body to tag mass ratios for adult fish. these recommendations will improve the confidence in results and the recovery process as well as increase the representable size classes in telemetry studies of this nature. this will further ensure the inclusion of smaller fish species as well as earlier life stages in telemetry studies. our study also demonstrated that external telemeter tags affected the activity patterns of individuals in the short term (< 3 days) but minimal in the long term. despite the stress caused by tagging, inadequate flows can contribute to the vulnerability of h. vittatus to predation, especially when environmental flows are not adequately maintained. the reduction in flows may play a role in increasing the risk of predation of h. vittatus and their overall condition in the freshwater environment. similarly, additional environmental stressors may be compounded by reduced flows, and if not addressed, it would further jeopardise the population of h. vittatus in the kruger national park. the excessive use of water resources by users upstream of the kruger national park (dabrowski, oberholster & dabrowski 2014; de villiers & mkwelo 2009; fouché 2008) is typically the main drivers of reduced flows in the downstream reaches of the letaba and olifants rivers. although compounded by other environmental stressors, altered flows expose h. vittatus to predation, increasing the likelihood of such events during the low flow season when exacerbated by abstraction upstream. in our study, the extremely low flows observed in the letaba river were further exacerbated by the presence of unused dams in the kruger national park, mainly engelhardt wier, that limited the release of water downstream. its removal should be considered to contribute to maintaining low flows and the biodiversity in the letaba river to ensure that environmental flow conditions for kruger national park and into mozambique are met. setting environmental flows and implementing them could be the solution to the conservation and sustainability of h. vittatus in the kruger national park. suitable allocations of environmental flows to the letaba and olifants rivers will have a direct influence on this species by reducing environmental stress, avoiding predators and improving their general fitness. the fine-scale detection of activity rates recorded using the telemeter tagged fish could further contribute to an understanding of the response of fish to environmental conditions enabling long-term monitoring of anthropogenic stressors placed on these and other river systems. the value of telemetry studies such as ours is that they can significantly assist in understanding the behavioural responses of fish species to environmental stressors, such as flows, and also other water quality parameters. monitoring the behavioural responses in real time and remotely can further assist the implementation of environmental flows in ensuring optimal functioning of the ecosystems and increasing the output of ecosystem services provided by freshwater ecosystems. the conservation of h. vittatus is essential in south africa as they serve as keystone species and meeting their requirements will assist in the conservation of other aquatic species maintaining the integrity of those freshwater ecosystems. acknowledgements the authors are grateful to the south african national parks (south africa) staff members, especially game guards, for their contribution to the success of the project. they thank the reviewers for their constructive that improved this manuscript. competing interests the authors have declared that no competing interest exists. authors’ contributions b.s., m.b., r.p., g.o.b. and c.t.d. conceptualised the study. g.o.b. and c.t.d. funded the study. b.s., m.b., r.p. and g.o.b. collected the data. b.s. and m.b. did the data analyses. b.s. and m.b. wrote the draft manuscript. r.p., g.o.b. and c.t.d. made contributions to this. funding information the would like to thank the water accelerator programme (wader) (south africa) and water research commission (south africa) for funding the fishtrac programme to monitor the water quality and flow variability, and fish response to these changes, in real time remotely in the olifants river, kruger national park. they would also like to acknowledge the national research foundation (south africa) and the associated glowsens project for providing operational support for this project. a further thanks to the university of mpumalanga (south africa) and the university of kwazulu-natal (south africa) for funding this project. the authors also thank idea-wild (the united states of america) for equipment used in this study. data availability statement the data are available on request from the corresponding author and are stored at the university of kwazulu-natal. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references abecasis, d., steckenreuter, a., reubens, j., aarestrup, k., alós, j., badalamenti, f. et al., 2018, ‘a review of acoustic telemetry in europe and the need for a regional aquatic telemetry network’, animal biotelemetry 6, 1–7. https://doi.org/10.1186/s40317-018-0156-0 ashton, p.j., 2010, ‘the demise of the nile crocodile (crocodylus niloticus) as a keystone species for aquatic ecosystem conservation in south africa: the case of the olifants river’, aquatic conservation: marine and freshwater ecosystems 20(5), 489–493. https://doi.org/10.1002/aqc.1132 beland, k.f., kocik, j.f. & sheehan, t.f., 2001, ‘striped bass predation upon atlantic salmon smolts in maine’, northeastern naturalist 8(3), 267–275. https://doi.org/10.2307/3858483 bengraı̈ne, k. & marhaba, t.f., 2003, ‘using principal component analysis to monitor spatial and temporal changes in water quality’, journal of hazardous materials 100(1–3), 179–195. https://doi.org/10.1016/s0304-3894(03)00104-3 block, b.a., holbrook, c.m., simmons, s.e., holland, k.n., ault, j.s., costa, d.p. et al., 2016, ‘toward a national animal telemetry network for aquatic observations in the united states’, animal biotelemetry 4, 6. https://doi.org/10.1186/s40317-015-0092-1 bolland, j.d., nunn, a.d., angelopoulos, n.v., dodd, j.r., davies, p., roberts, c.g. et al., 2019, ‘refinement of acoustic-tagging protocol for twaite shad alosa fallax (lacépède), a species sensitive to handling and sedation’, fisheries research 212, 183–187. https://doi.org/10.1016/j.fishres.2018.12.006 bouwman, h., booyens, p., govender, d., pienaar, d. & polder, a., 2014, ‘chlorinated, brominated, and fluorinated organic pollutants in nile crocodile eggs from the kruger national park, south africa’, ecotoxicology and environmental safety 104, 393–402. https://doi.org/10.1016/j.ecoenv.2013.12.005 burnett, m.j., o’brien, g.c., jacobs, f.j., botha, f., jewitt, g. & downs, c.t., 2020, ‘southern african inland fish tracking programme (fishtrac): an evaluation of the approach for monitoring ecological consequences of multiple water resource stressors, remotely and in real time’, ecological indicators 111, 106001. https://doi.org/10.1016/j.ecolind.2019.106001 burnett, m.j., o’brien, g.c., wepener, v. & pienaar, d., 2018, ‘the spatial ecology of adult labeobarbus marequensis and their response to flow and habitat variability in the crocodile river, kruger national park’, african journal of aquatic science 43(4), 375–384. https://doi.org/10.2989/16085914.2018.1517077 cooke, s.j., hinch, s.g., wikelski, m., andrews, r.d., kuchel, l.j., wolcott, t.g. et al., 2004, ‘biotelemetry: a mechanistic approach to ecology’, trends in ecology & evolution 19(6), 334–343. https://doi.org/10.1016/j.tree.2004.04.003 cooke, s.j., midwood, j.d., thiem, j.d., klimley, p., lucas, m.c., thorstad, e.b. et al., 2013, ‘tracking animals in freshwater with electronic tags: past, present and future’, animal biotelemetry 1, 5. https://doi.org/10.1186/2050-3385-1-5 cooke, s.j., nguyen, v.m., kessel, s.t., hussey, n.e., young, n. & ford, a.t., 2017, ‘troubling issues at the frontier of animal tracking for conservation and management’, conservation biology 31(5), 1205–1207. https://doi.org/10.1111/cobi.12895 cooke, s.j. & schreer, j.f., 2003, ‘environmental monitoring using physiological telemetry – a case study examining common carp responses to thermal pollution in a coal-fired generating station effluent’, water, air, and soil pollution 142, 113–136. https://doi.org/10.1023/a:1022082003932 core, r., 2015, r: a language and environment for statistical computing, r foundation for statistical computing, vienna, austria. cowley, p.d., terörde, a.i. & whitfield, a.k., 2017, ‘birds as major predators of fishes in the east kleinemonde estuary’, african zoology 52, 147–154. https://doi.org/10.1080/15627020.2017.1361865 dabrowski, j., oberholster, p.j. & dabrowski, j.m., 2014, ‘water quality of flag boshielo dam, olifants river, south africa: historical trends and the impact of drought’, water sa 40(2), 345–358. https://doi.org/10.4314/wsa.v40i2.17 de moor, i.j., 1996, ‘case studies of the invasion by four alien fish species (cyprinus carpio, micropterus salmoides, oreochromis macrochir and o. mossambicus) of freshwater ecosystems in southern africa’, transactions of the royal society of south africa 51(1), 233–255. https://doi.org/10.1080/00359199609520609 department of environmental affairs and tourism, 2007, national environmental management: biodiversity act 2004 (act 10 of 2004): amendment of critically endangered, endangered, vulnerable and protected species list, no. r. 1187, department of environmental affairs, pretoria. de villiers, s. & mkwelo, s.t., 2009, ‘has monitoring failed the olifants river, mpumalanga?’, water sa 35(5), 671–676. https://doi.org/10.4314/wsa.v35i5.49193 du plessis, a., 2019, ‘evaluation of southern and south africa’s freshwater resources’, in a. du plessis (ed.), water as an inescapable risk, pp. 147–172, springer, cham. du preez, h.h. & steyn, g.j., 1992, ‘a preliminary investigation of the concentration of selected metals in the tissues and organs of the tigerfish (hydrocynus vittatus) from the olifants river, kruger national park, south africa’, water sa 18(2), 131–136. dudgeon, d., 2014, ‘threats to freshwater biodiversity in a changing world’, in b. freedman (ed.), global environmental change: handbook of global environmental pollution, vol. 1, pp. 243–253, springer, dordrecht. dudgeon, d., arthington, a.h., gessner, m.o., kawabata, z.i., knowler, d.j., lévêque, c. et al., 2006, ‘freshwater biodiversity: importance, threats, status and conservation challenges’, biological reviews 81(2), 163–182. https://doi.org/10.1017/s1464793105006950 ellender, b.r. & weyl, o.l., 2014, ‘a review of current knowledge, risk and ecological impacts associated with non-native freshwater fish introductions in south africa’, aquatic invasions 9(2), 117–132. https://doi.org/10.3391/ai.2014.9.2.01 ferreira, s.m. & pienaar, d., 2011, ‘degradation of the crocodile population in the olifants river gorge of kruger national park, south africa’, aquatic conservation: marine and freshwater ecosystems 21(2), 155–164. https://doi.org/10.1002/aqc.1175 fouché, p.s.o., 2008, ‘status of the lowveld largescale yellowfish labeobarbus marequensis (a. smith, 1841)’, in n.d. impson, i.r. bills & l. wolhuter (eds.), technical report on the state of yellowfishes in south africa, wrc report no. kv 212/08, water research commission, pretoria. frank, h.j., mather, m.e., smith, j.m., muth, r.m., finn, j.t. & mccormick, s.d., 2009, ‘what is “fallback”? metrics needed to assess telemetry tag effects on anadromous fish behavior’, hydrobiologia 635, 237–249. https://doi.org/10.1007/s10750-009-9917-3 games, i. & moreau, j., 1997, ‘the feeding ecology of two nile crocodile populations in the zambezi valley’, in j. moreau (ed.), advances in the ecology of lake kariba, pp. 183–195, uz publications, harare. gerber, r., smit, n.j., van vuuren, j.h. & wepener, v., 2016, ‘metal concentrations in hydrocynus vittatus (castelnau 1861) populations from a premier conservation area: relationships with environmental concentrations’, ecotoxicology and environmental safety 129, 91–102. https://doi.org/10.1016/j.ecoenv.2016.03.009 godin, j.g., 1997, behavioural ecology of teleost fishes, oxford university press, oxford. halfyard, e.a., webber, d., del papa, j., leadley, t., kessel, s.t., colborne, s.f. et al., 2017, ‘evaluation of an acoustic telemetry transmitter designed to identify predation events’, methods in ecology and evolution 8(9), 1063–1071. https://doi.org/10.1111/2041-210x.12726 harrison, t.d. & whitfield, a.k., 2006, ‘application of a multimetric fish index to assess the environmental condition of south african estuaries’, estuaries and coasts 29, 1108–1120. https://doi.org/10.1007/bf02781813 horn, j.a., mateus-pinilla, n., warner, r.e. & heske, e.j., 2011, ‘home range, habitat use, and activity patterns of free-roaming domestic cats’, journal of wildlife management 75(5), 1177–1185. https://doi.org/10.1002/jwmg.145 hussey, n.e., kessel, s.t., aarestrup, k., cooke, s.j., cowley, p.d., fisk, a.t. et al., 2015, ‘aquatic animal telemetry: a panoramic window into the underwater world’, science 348(6240), 1–10. https://doi.org/10.1126/science.1255642 jacobs, f., o’brien, g. & smit, n., 2016, ‘diel movement of smallmouth yellowfish labeobarbus aeneus in the vaal river, south africa’, african journal of aquatic science 41(1), 73–76. https://doi.org/10.2989/16085914.2015.1136804 jacobs, f.j., 2017, the conservation ecology of the african tigerfish hydrocynus vittatus in the kavango river, namibia, phd thesis, university of kwazulu-natal, pietermaritzburg. jacobs, f.j., naesje, t.f., ulvan, e.m., weyl, o.l.f., tiyeho, d., hay, c.j. et al., 2019, ‘implications of the movement behaviour of african tigerfish hydrocynus vittatus for the design of freshwater protected areas’, journal of fish biology 1–9. https://doi.org/10.1111/jfb.14158 jepsen, n., koed, a., thorstad, e.b. & baras, e., 2002, ‘surgical implantation of telemetry transmitters in fish: how much have we learned?’, hydrobiologia 483, 239–248. https://doi.org/10.1023/a:1021356302311 jepsen, n., schreck, c., clements, s. & thorstad, e.b., 2004, ‘a brief discussion on the 2% tag/body weight rule of thumb’, in m.t. spedicato, g. lembo & g. marmulla (eds.), aquatic telemetry: advances and applications. proceedings of the fifth conference on fish telemetry held in europe, ustica, italy, 9–13 june 2003, pp. 255–259, food and agriculture organization of the united nations (fao)-coispa tecnologia & ricerca, rome. jepsen, n., thorstad, e.b., havn, t. & lucas, m.c., 2015, ‘the use of external electronic tags on fish: an evaluation of tag retention and tagging effects’, animal biotelemetry 3, 49. https://doi.org/10.1186/s40317-015-0086-z. klinard, n.v. & matley, j.k., 2020, ‘living until proven dead: addressing mortality in acoustic telemetry research’, reviews in fish biology and fisheries 22, 1–5. https://doi.org/10.1007/s11160-020-09613-z lennox, r.j., aarestrup, k., cooke, s.j., cowley, p.d., deng, z.d., fisk, a.t. et al., 2017, ‘envisioning the future of aquatic animal tracking: technology, science, and application’, bioscience 67(10), 884–896. https://doi.org/10.1093/biosci/bix098 mckight, p.e. & najab, j., 2010, ‘kruskal-wallis test’, in i.b. weiner & w.e. craighead (eds.), the corsini encyclopedia of psychology. https://doi.org/10.1002/9780470479216.corpsy0491 metcalfe, j.d., wright, s., tudorache, c. & wilson, r.p., 2016, ‘recent advances in telemetry for estimating the energy metabolism of wild fishes’, journal of fish biology 88(1), 284–297. https://doi.org/10.1111/jfb.12804 mims, m.c. & olden, j.d., 2013, ‘fish assemblages respond to altered flow regimes via ecological filtering of life history strategies’, freshwater biology 58(1), 50–62. https://doi.org/10.1111/fwb.12037 moxham, e.j., cowley, p.d., bennett, r.h. & von brandis, r.g., 2019, ‘movement and predation: a catch-and-release study on the acoustic tracking of bonefish in the indian ocean’, environmental biology of fishes 102, 365–381. https://doi.org/10.1007/s10641-019-00850-1 nasir, m.f.m., samsudin, m.s., mohamad, i., awaluddin, m.r.a., mansor, m.a., juahir, h. et al., 2011, ‘river water quality modeling using combined principle component analysis (pca) and multiple linear regressions (mlr): a case study at klang river, malaysia’, world applied sciences journal 14, 73–82. o’brien, g.c., bulfin, j.b., husted, a. & smit, n.j., 2012, ‘comparative behavioural assessment of an established and a new tigerfish hydrocynus vittatus population in two man-made lakes in the limpopo river catchment, southern africa’, african journal of aquatic science 37(3), 253–263. https://doi.org/10.2989/16085914.2012.723196 o’brien, g.c., dickens, c., hines, e., wepener, v., stassen, r., quayle, l. et al., 2018, ‘a regional-scale ecological risk framework for environmental flow evaluations’, hydrology and earth system sciences 22(2), 957–975. https://doi.org/10.5194/hess-22-957-2018 o’brien, g.c., jacobs, f., cronje, l., wepener, v. & smit, n.j., 2013, ‘habitat preferences and movement of adult yellowfishes in the vaal river, south africa’, south african journal of science 109(7/8), 1–8. https://doi.org/10.1590/sajs.2013/20120095 o’brien, g.c., jacobs, f., evans, s.w. & smit, n.j., 2014, ‘first observation of african tigerfish hydrocynus vittatus predating on barn swallows hirundo rustica in flight’, journal of fish biology 84(1), 263–266. https://doi.org/10.1111/jfb.12278 o’brien, g.c., ross, m., hanzen, c., dlamini, v., petersen, r., diedericks, g.j. et al., 2019, ‘river connectivity and fish migration considerations in the management of multiple stressors in south africa’, marine and freshwater research 70(9), 1254–1264. https://doi.org/10.1071/mf19183 o’brien, g.c., swemmer, r. & wepener, v., 2009, ‘ecological integrity assessment of the fish assemblages of the matigulu/nyoni and umvoti estuaries, kwazulu-natal, south africa’, african journal of aquatic science 34(3), 293–302. https://doi.org/10.2989/ajas.2009.34.3.11.987 økland, f., thorstad, e.b., hay, c.j., næsje, t.f. & chanda, b., 2005, ‘patterns of movement and habitat use by tigerfish (hydrocynus vittatus) in the upper zambezi river (namibia)’, ecology of freshwater fish 14(1), 79–86. https://doi.org/10.1111/j.1600-0633.2004.00080.x østnes, j.e., kroglund, r.t., kleven, o. & nygård, t., 2019, ‘migratory patterns of ospreys (pandion haliaetus) from central norway’, ornis fennica 96(3), 101–111. paukert, c.p., chvala, p.j., heikes, b.l. & brown, m.l., 2001, ‘effects of implanted transmitter size and surgery on survival, growth, and wound healing of bluegill’, transactions of the american fisheries society 130(5), 975–980. https://doi.org/10.1577/1548-8659(2001)130%3c0975:eoitsa%3e2.0.co;2 pearce, f., 2018, when the rivers run dry, water-the defining crisis of the twenty-first century, beacon press, boston, ma. pienaar, u.d.v., 1964, ‘the small mammals of the kruger national park-a systematic list and zoogeography’, koedoe 7(1), 1–25. https://doi.org/10.4102/koedoe.v7i1.795 pollard, s., mallory, s., riddell, e. & sawunyama, t., 2012, towards improving the assessment and implementation of the reserve: real-time assessment and implementation of the ecological reserve, report, water research commission, pretoria. rodell, m., famiglietti, j.s., wiese, d.n., reager, j.t., beaudoing, h.k., landerer, f.w. et al., 2018, ‘emerging trends in global freshwater availability’, nature 557, 651–659. https://doi.org/10.1038/s41586-018-0123-1 rogers, k.h. & o’keeffe, j., 2003, ‘river heterogeneity: ecosystem structure, function and management’, in a.r. sinclair & b. walker (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 189–218, island press, washington, dc. roux, d.j., nel, j.l., ashton, p.j., deacon, a.r., de moor, f.c., hardwick, d. et al., 2008, ‘designing protected areas to conserve riverine biodiversity: lessons from a hypothetical redesign of the kruger national park’, biological conservation 141(1), 100–117. https://doi.org/10.1016/j.biocon.2007.09.002 roux, f., steyn, g., hay, c. & wagenaar, i., 2018, ‘movement patterns and home range size of tigerfish (hydrocynus vittatus) in the incomati river system, south africa’, koedoe 60(1), 1–3. https://doi.org/10.4102/koedoe.v60i1.1397 rowe-rowe, d.t., 1977, ‘food ecology of otters in natal, south africa’, oikos 28(2/3), 210–219. https://doi.org/10.2307/3543973 schultz, a.a., afentoulis, v.b., yip, c.j. & johnson, m.n., 2017, ‘efficacy of an acoustic tag with predation detection technology’, north american journal of fisheries management 37(3), 574–581. https://doi.org/10.1080/02755947.2017.1290720 sequeira, a.m.m., heupel, m.r., lea, m.a., eguíluz, v.m., duarte, c.m., meekan, m.g. et al., 2019, ‘the importance of sample size in marine megafauna tagging studies’, ecological applications 29(6), 1344–1360. https://doi.org/10.1002/eap.1947 skalski, j.r., richins, s.m. & townsend, r.l., 2018, ‘a statistical test and sample size recommendations for comparing community composition following pca’, plos one 13(10), 1–19. https://doi.org/10.1371/journal.pone.0206033 smit, n.j., wepener, v., vlok, w., wagenaar, g.m. & van vuuren, j.h.j., 2013, conservation of tigerfish, hydrocynus vittatus, in the kruger national park with the emphasis on establishing the suitability of the water quantity and quality requirements for the olifants and luvuvhu rivers, wrc report no. 1922/1/12, water research commision, pretoria. sosa-lópez, a., mouillot, d., do chi, t. & ramos-miranda, j., 2005, ‘ecological indicators based on fish biomass distribution along trophic levels: an application to the terminos coastal lagoon, mexico’, ices journal of marine science 62(3), 453–458. https://doi.org/10.1016/j.icesjms.2004.12.004 stears, k. & mccauley, d.j., 2018, ‘hippopotamus dung inputs accelerate fish predation by terrestrial consumers’, african journal of ecology 56(4), 1034–1038. https://doi.org/10.1111/aje.12543 stewart, k., matthiesen, d., leblanc, l. & west, j., 1997, ‘prey diversity and selectivity of the african fish eagle: data from a roost in northern kenya’, african journal of ecology 35(2), 133–145. https://doi.org/10.1111/j.1365-2028.1997.066-89066.x steyn, g.j., gagiano, c.l., deacon, a.r. & du preez, h.h., 1996, ‘notes on the induced reproduction and development of the tigerfish, hydrocynus vittatus (characidae), embryos and larvae’, environmental biology of fishes 47, 387–398. https://doi.org/10.1007/bf00005052 thorstad, e.b., hay, c.j., næsje, t.f., chanda, b. & økland, f., 2003, ‘space use and habitat utilisation of tigerfish and the two cichlid species nembwe and threespot tilapia in the upper zambezi river’, in implications for fisheries management, nina project report no. 24, pp. 1–22, norwegian institute for nature research, trondheim, norway. thorstad, e.b., rikardsen, a.h., alp, a. & økland, f., 2013, ‘the use of electronic tags in fish research – an overview of fish telemetry methods’, turkish journal of fisheries and aquatic sciences 13, 881–896. https://doi.org/10.4194/1303-2712-v13_5_13 thronson, a. & quigg, a., 2008, ‘fifty-five years of fish kills in coastal texas’, estuaries and coasts 31, 802–813. https://doi.org/10.1007/s12237-008-9056-5 van den brink, p.j., van den brink, n.w. & ter braak, c.j., 2003, ‘multivariate analysis of ecotoxicological data using ordination: demonstrations of utility on the basis of various examples’, australasian journal of ecotoxicology 9(2), 141–156. vignaud, p., duringer, p., mackaye, h.t., likius, a., blondel, c., boisserie, j.r. et al., 2002, ‘geology and palaeontology of the upper miocene toros-menalla hominid locality, chad’, nature 418, 152–155. https://doi.org/10.1038/nature00880 wisniewska, d.m., johnson, m., teilmann, j., rojano-doñate, l., shearer, j., sveegaard, s. et al., 2016, ‘ultra-high foraging rates of harbor porpoises make them vulnerable to anthropogenic disturbance’, current biology 26(11), 1441–1446. https://doi.org/10.1016/j.cub.2016.03.069 woodborne, s., huchzermeyer, k.d.a., govender, d., pienaar, d.j., hall, g., myburgh, j.g. et al., 2012, ‘ecosystem change and the olifants river crocodile mass mortality events’, ecosphere 3(10), 1–17. https://doi.org/10.1890/es12-00170.1 kyle nyamithi.qxd interesting new record and further notes on the occurrence of marine fish in nyamithi pan, ndumo game reserve, south africa r. kyle issn 0075-6458 123 koedoe 45/2 (2002) nyamithi pan, situated in ndumo game reserve, south africa, is a floodplain pan near the confluence of the usuthu and pongolo rivers. it lies approximately 75 km from the indian ocean. the floodplain and its fish have been extensively surveyed (coke & pott 1970; kok 1980; merron et al 1993, 1994, 1994a, 1994b, 1994c, 1994d; pooley 1975) and there are many records of the occurrence of marine fish in this and other pans of the pongolo and usuthu rivers. these are, however, usually isolated instances of individual fish being caught and attracting attention. many of the records (table 1) are old and the pongolo and usuthu river systems have been markedly modified in the last few decades through the construction of dams, particularly the pongolopoort dam, agriculture and water abstraction. fishing, mainly gillnetting, has intensified greatly and several exotic fish, particularly cyprinus carpio (carp) have become well established in many pongolo floodplain pans (pers. obs.) in the last few years. more recent, although erratic, fish monitoring in nyamithi pan by ekzn wildlife staff has yielded some interesting records (table 2) confirming the continued immigration of some species into the system and also added a new species record for the pan and river systems. more formal, though intensive, fish sampling (merron et al. 1993, 1994a, 1994b, 1994c) failed to show the presence of any marine fish in nyamithi. in 1999, tourists reported a dead shark in nyamithi pan and on investigation it was found to be a carcharhinus leucas of almost two metres. shark fin sightings had been reported for several months but the reason for its death was not clear. there are often well over one hundred crocodylus niloticus (nile crocodile) in the pan but the dead shark was mostly intact and it was reported that an adult hippo appeared to be guarding the cartable 1 historical records of the occurrence of marine fish in the pongolo and usuthu river systems species name place no. reference pristis microdon sawfish pongolo river 1(?) campbell 1969 carcharhinus leucas zambezi shark confluence of 1(?) crass 1964 pongolo/usuthu megalops cyprinoides oxeye tarpon pongolo & usuthu several crass 1964 rivers and pans coke & pott 1970 merron et al. 1994a mugil cephalus bull mullet nyamithi pan two pooley 1975 acanthopagrus berda river bream pongolo river and pans several pooley 1975 bruton & cooper 1980 microphis fluviatilis pipefish usuthu river ? merron et al. 1993 natural history notes kyle nyamithi.qxd 2005/12/09 10:08 page 123 cass and was seen to drive away any crocodiles which approached the dead fish. one megalops cyprinoides was caught in 1996 while, early in 2002, nine more were caught and many more were seen breaking the surface in the manner typical of the species (skelton 1993). in 1998, one acanthopagrus berda, of 420 mm (fork length), was caught, tagged and released and another, smaller, a. berda was caught in 2000 (tables 1 & 2). in early 2002, a total of nine elops machnata, of fork lengths 650–800 mm, were caught and released. several more were seen and the fish were in good condition. these fish represent the first records of this species in these river systems. this species has often been recorded in estuaries, but a survey of the literature ( bruton & kok 1980; skelton 1993; van der elst 1998; whitfield 1998) yielded no mention of e. machnata appreciably upstream of estuaries, despite their ability to survive in fresh water. all the recent nyamithi records were made during relatively low water levels, several months after any flooding episodes. it seems likely that the fish enter the pan during high flood levels and then stay there for extended periods. the fish were released alive and so not examined internally for gonad condition. as the m. cyprinoides and e. machnata recorded recently were in good condition and the species were still apparently abundant several months after they had immigrated, it appears that they compete well with the hydrocynus vittatus (tigerfish) which were abundant in the pan at the time. they also appear to survive well in spite of high densities of large c. niloticus. no c. carpio have yet been recorded in nyamithi, though they are abundant in namanini pan, just outside the reserve. it is possible that they fall easy prey to c. niloticus. the most recent records are the first evidence of large scale sustained immigration of m. cyprinoides and e. machnata far up rivers. using nyamithi as an indicator of the system’s fish species composition, it appears that despite their modifications, the immigration of marine species to the usuthu and pongolo river systems has not stopped and may have increased in some instances. acknowledgements i thank ezemvelo kwazulu natal wildlife for permission to publish this paper. references bruton, m.n. & h.m. kok. 1980. the freshwater fishes of maputaland. pp. 210-244. in: bruton, m.n. & k.h. cooper (eds.). studies on the ecology of maputaland. grahamstown: rhodes university. campbell, g.c. 1969. a review of scientific investigations in the tongaland area of northern natal. transactions of the royal society of south africa 38(4): 305-316. coke, m.m. & mcc. pott. 1970. the pongolo floodplain pans. pietermaritzburg: natal parks board. crass, r.s. 1964. freshwater fishes of natal. pietermaritzburg: shuter & shooter. kok, h.m. 1980. ecological studies on some important fish species of the pongolo floodplain, kwazulu, south africa. phd. thesis, university of natal. merron, g.s., s.k. weldrick & m.n. bruton. 1993. a fisheries survey of the phongolo floodkoedoe 45/2 (2002) 124 issn 0075-6458 table 2 recent records of marine fish in nyamithi pan, ndumo game reserve species name number year carcharhinus leucas zambezi shark 1 1999 megalops cyprinoides oxeye tarpon 1, 9 1996, 2002 acanthopagrus berda river bream 1, 1 1998 & 2000 elops machnata springer 9 2002 kyle nyamithi.qxd 2005/12/09 10:08 page 124 plain, zululand, south africa—december 1993. j.l.b. institute of ichthyology, grahamstown. (investigational report; no 45.) merron, g.s., s.k.weldrick & h. kaiser. 1994. the second fisheries survey of the phongolo floodplain, zululand, south africa—february 1994. j.l.b. institute of ichthyology, grahamstown. (investigational report; no 46.) merron, g.s., s.k.weldrick, h. kaiser & m.n. bruton. 1994a. the third fisheries survey of the phongolo floodplain, zululand, south africa— april 1994. institute of ichthyology, grahamstown. (investigational report; no 47.) merron, g.s., s.k.weldrick, h. kaiser & m.n. bruton. 1994b. the fourth fisheries survey of the phongolo floodplain, zululand, south africa—june 1994. institute of ichthyology, grahamstown. (investigational report; no 48.) merron, g.s., s.k.weldrick, h. kaiser & m.n. bruton. 1994c. the fifth fisheries survey of the phongolo floodplain, zululand, south africa—september 1994. institute of ichthyology, grahamstown. (investigational report; no 49.) merron, g.s., s.k.weldrick, h. kaiser & m.n. bruton. 1994d. the sixth fisheries survey of the phongolo floodplain, zululand, south africanovember 1994. institute of ichthyology, grahamstown. (investigational report; no 50.) pooley, a.c. 1975. new fish records for ndumo game reserve. lammergeyer 22: 50-51. skelton, p.h. 1993. a complete guide to the freshwater fishes of southern africa. halfway house: southern book publishers. van der elst. r.p. 1988. a guide to the common sea fishes of southern africa. cape town: struik. whitfield, a.k. 1998. biology and ecology of fishes in southern african estuaries. ichthyological monographs of the j.l.b. smith institute of ichthyology 2: 1-223. r. kyle, p.o. box 43, kwangwanase, 3973, south africa. (ezemvelo kwazulu natal wildlife; rkyle@iafrica.com). issn 0075-6458 125 koedoe 45/2 (2002) kyle nyamithi.qxd 2005/12/09 10:08 page 125 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <feff005500740069006c0069007a006500200065007300740061007300200063006f006e00660069006700750072006100e700f5006500730020007000610072006100200063007200690061007200200064006f00630075006d0065006e0074006f0073002000500044004600200063006f006d00200075006d00610020007200650073006f006c007500e700e3006f00200064006500200069006d006100670065006d0020007300750070006500720069006f0072002000700061007200610020006f006200740065007200200075006d00610020007100750061006c0069006400610064006500200064006500200069006d0070007200650073007300e3006f0020006d0065006c0068006f0072002e0020004f007300200064006f00630075006d0065006e0074006f0073002000500044004600200070006f00640065006d0020007300650072002000610062006500720074006f007300200063006f006d0020006f0020004100630072006f006200610074002c002000520065006100640065007200200035002e00300020006500200070006f00730074006500720069006f0072002e00200045007300740061007300200063006f006e00660069006700750072006100e700f50065007300200072006500710075006500720065006d00200069006e0063006f00720070006f0072006100e700e3006f00200064006500200066006f006e00740065002e> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 page 3 de kock.qxd freshwater mollusc diversity in the kruger national park: a comparison between a period of prolonged drought and a period of exceptionally high rainfall k.n. de kock, c.t. wolmarans and l.h. du preez de kock, k.n., c.t. wolmarans and l.h. du preez. 2002. freshwater mollusc diversity in the kruger national park: a comparison between a period of prolonged drought and a period of exceptionally high rainfall. koedoe 45(2): 1-11. pretoria. issn 0075-6458. most of the previous records of the freshwater molluscs from the kruger national park date back to 1966 and earlier. on account of several droughts between 1966 and 1995, a survey was done in 1995 to evaluate the effect of these droughts on the mollusc population. as a result of extensive rainfall between 1995 and 2000 another survey was conducted to establish the effect of a period of exceptionally high rainfall on the species’ diversity of the mollusc population. with the exception of three habitats, an increase in mollusc diversity was recorded for all the other habitats previously surveyed. one of the invader freshwater snail species, aplexa marmorata, collected in only one habitat in 1995, was found in seven habitats located mainly in the south during the 2001 survey. another interesting finding was that juvenile specimens of lanistes ovum, of which large specimens were present prior to 1966 but none found in 1995, were present in the sirheni dam in 2001. from this study the positive effect of the high rainfall on the species’ diversity is highly evident. key words: mollusca, freshwater molluscs, invader species, rainfall, kruger national park, south africa. k.n. de kock, c.t. wolmarans and l.h. du preez, school of environmental sciences and development (zoology), potchefstroom university for christian higher education, private bag x6001, potchefstroom 2520, republic of south africa. issn 0075-6458 1 koedoe 45/2 (2002) introduction the importance of regular, normal rainfall to maintain the bio-diversity in conservation areas in south africa cannot be over-emphasized, especially for those aquatic organisms which lack the ability to migrate to other habitats during periods of serious drought. this inability to migrate leads to the situation where re-population of isolated habitats, such as dams and pans, is largely dependent on time and passive dispersal of biota. certain freshwater invertebrates, however, have the ability to aestivate, a strategy enabling them to survive prolonged periods of drought. apart from these freshwater snail species, among others, bulinus reticulatus mandahl-barth, b. forskalii (ehrenberg, 1831), b. tropicus (krauss, 1848) and the b. africanus group of species (brown 1994), it is mostly the eggs of some freshwater crustaceans such as triops, eocyzicus and branchipodopsis which can survive periods of drought (davies & day 1997). with regard to the freshwater molluscs, it is known that the largest bio-diversity is to be found in the tropical and sub-tropical areas of the world. in south africa it is mainly the limpopo, mpumalanga and kwazulu-natal provinces that represent the tropical and subtropical regions with the kruger national park situated in both the first two mentioned provinces. a large variety of freshwater molluscs collected up to 1966 were reported by oberholzer & van eeden (1967) for the kruger national park. de kock.qxd 2005/12/09 10:01 page 1 koedoe 45/2 (2002) 2 issn 0075-6458 the distribution of freshwater molluscs is not a static process and is mainly influenced by the availability of suitable water habitats. the most recent account of the diversity of freshwater molluscs in the park, prior to the 2001 survey, dates back to 1995 and is a report by de kock & wolmarans (1998) on a survey done after the drought of 1991-1992. between 1966 and 1995, the park was struck by several droughts, namely those in 1982-1983 and 1986-1987—the former also being the severest—prior to the third drought of 1991-1992. according to zambatis & biggs (1995) several of the rainfall monitoring stations recorded the lowest rainfall in history. between 1995 and 2001, however, average to above normal high rainfall figures were recorded for most of the rainfall monitoring stations. this situation prompted the idea to do a follow-up of the 1995 survey to evaluate the effect of the availability of more permanent water habitats on the freshwater mollusc diversity, and to compare results with the 1995 survey. methods and materials during the present investigation we concentrated mainly on water bodies that had been surveyed for snails during the 1995 collection. we also concentrated on several other water bodies that were dry during the 1995 survey, or seemed suitable for supporting molluscs but of which there were no previous records in the national freshwater snail collection (nfsc). a total of 43 habitats were surveyed during 2001. in each survey, all the habitat types were screened for molluscs by means of aluminium mollusc scoops. the collected molluscs were identified by the senior author who had mainly been responsible for the identification of the molluscs in the nfsc since 1973. the molluscs were subsequently incorporated into the national freshwater snail collection. habitats without surface water were screened for mollusc shells. rainfall records, for the periods 1982-1983, 1985-1987, 1991-1994 and 19952001, were obtained from the kruger national park for the monitoring stations nearest to the habitats surveyed. fig. 1. water bodies (•) sampled during the 2001 survey. rainfall monitoring stations (♦). de kock.qxd 2005/12/09 10:01 page 2 results water bodies sampled during the 2001 survey are depicted in fig. 1. the results of this investigation are presented in tables 1, 2 & 3 and figs. 2–5. a comparison of the species diversity recorded during the 1995 survey with that of the present survey (table 1) clearly shows a substantial increase in the diversity in most of the dams and rivers since 1995. a decrease in the species diversity during the 2001 survey is only evident for the sabie river, the nwaswitsontso river and the orpen dam (which forms part of the nwaswitsontso river drainage system). during 1995, molluscs were found in only eight of the 24 sites surveyed during both years, while mollucs were present in all of these habitats in 2001. taking into account the number of habitats sampled during both surveys, it is evident that b. forskalii was present in 16 of the 24 habitats surveyed in 2001, while it was recovered from only four of the habitats during the 1995 survey (table 1). furthermore, this snail was recorded from nine of the 18 habitats surveyed for the first time in 2001. with regard to its distribution in 1995, it is clear that it was only found in the southern part of the area, while it was more or less evenly distributed throughout the whole area in 2001 (fig. 2). in contrast to this, physa acuta draparnaud, 1805, which is well known for its invading potential, in spite of exceptionally high mean rainfall for the last five years (table 4), was present in only two of the habitats, while it was found only once during the 1995 survey (fig. 3). aplexa marmorata (guilding, 1828) which was present only in the sabie river during the 1995 survey was collected in seven habitats located mainly in the south during the 2001 survey (fig. 4). lymnaea columella say, 1817 another invader species, was recorded only three times before 1966 in the kruger national park and only once during the 1995 survey (fig. 5). in contrast to this, it was collected in eight habitats during the 2001 survey. issn 0075-6458 3 koedoe 45/2 (2002) fig. 2. distribution of bulinus forscali in 1995 (o) and 2001 (•). rainfall monitoring stations (♦). de kock.qxd 2005/12/09 10:01 page 3 koedoe 45/2 (2002) 4 issn 0075-6458 table 1 habitats of the kruger national park surveyed in both 1995 and 2001 rivers matjulu spruit mbyamiti river mlambane spruit nwanetsi river nwaswitsontso river sabie river timbavati river engelhard dam grid reference 25° 21' 26"s; 31°31'11"e 25°13'27"s; 31°34'38"e 25°21'26"s; 31°31'11"e 24°25"12"s; 31°47'01"e 24°37'10"s; 31°47'28"e 25°07'06"s; 31°55'00"e 24°13'50"s; 31°38'03"e 23°50'28"s; 31°38'07"e 1995 no molluscs bulinus globosus spathopsis wahlbergi no molluscs bulinus globosus bulinus natalensis bulinus forskalii biomphalaria pfeifferi lymnaea natalensis burnupia sp melanoides tuberculata cleopatra bulimoides spathopsis wahlbergi spathopsis petersi corbicula fluminalis eupera ferruginea cleopatra ferruginea spathopsis petersi bulinus globosus biomphalaria pfeifferi lymnaea natalensis lymnaea columella physa acuta aplexa marmorata melanoides tuberculata unio caffer eupera ferruginea oxyloma patentissima no molluscs no molluscs 2001 bulinus globosus bulinus forskalii biomphalaria pfeifferi lymnaea natalensis lymnaea columella aplexa marmorata burnupia caffra melanoides tuberculata bulinus globosus lymnaea columella aplexa marmorata bulinus forskalii biomphalaria pfeifferi aplexa marmorata bulinus tropicus bulinus forskalii biomphalaria pfeifferi ceratophallus natalensis burnupia caffra melanoides tuberculata spathopsis petersi corbicula fluminalis oxyloma striata bulinus forskalii physa acuta aplexa marmorata bulinus africanus lymnaea natalensis burnupia caffra bulinus globosus bulinus tropicus bulinus forskalii biomphalaria pfeifferi physa acuta melanoides tuberculata corbicula fluminalis pisidium pirothi bulinus tropicus de kock.qxd 2005/12/09 10:01 page 4 issn 0075-6458 5 koedoe 45/2 (2002) table 1 (continued) rivers gezantombi dam gudzani dam klopperfontein dam krapkuil dam kumana dam manzimahle dam mazithi dam nkaya pan nsemani dam nwanetsi water hole olifantdrinkgat orpen dam pioniers dam grid reference 25°20'05"s; 31°52'35"e 24°22'05"s; 31°55'33"e 22°37'42"s; 31°10'20"e 23°14'43"s; 31°18'07"e 24°37'11"s; 31°47'40"e 24°53'51"s; 31°43'55"e 24°43'20"s; 31°50'34"e 24°31'54"s; 31°45'25"e 24°23'32"s; 31°43'05"e 24°28'22"s; 31°58'51"e 24°53'56"s; 31°44'54"e 24°47'31"s; 31°53'51"e 23°31'29"s; 31°23'56"e 1995 no molluscs bulinus globosus bulinus forskalii biomphalaria pfeifferi lymnaea natalensis spathopsis petersi unio caffer no molluscs no molluscs no molluscs no molluscs no molluscs no molluscs no molluscs bulinus globosus bulinus natalensis bulinus forskalii biomphalaria pfeifferi lymnaea natalensis burnupia sp. melanoides tuberculata cleopatra bulimoides spathopsis wahlbergi spathopsis petersi corbicula fluminalis eupera ferruginea no molluscs bulinus globosus bulinus tropicus bulinus forskalii biomphalaria pfeifferi lymnaea natalensis ceratophallus natalensis melanoides tuberculata spathopsis wahlbergi spathopsis petersi bulinus globosus biomphalaria pfeifferi lymnaea natalensis 2001 bulinus forskalii lymnaea. columella aplexa marmorata burnupia caffra bulinus globosus bulinus tropicus biomphalaria pfeifferi lymnaea natalensis ceratophallus natalensis melanoides tuberculata spathopsis petersi corbicula fluminalis bulinus tropicus bulinus forskalii burnupia caffra bulinus forskalii bulinus forskalii bulinus forskalii bulinus forskalii bulinus forskalii burnupia caffra bulinus globosus bulinus tropicus bulinus forskalii biomphalaria pfeifferi ceratophallus natalensis burnupia caffra spathopsis wahlbergi spathopsis petersi corbicula fluminalis bulinus forskalii lymnaea columella ceratophallus natalensis bulinus globosus bulinus forskalii biomphalaria pfeifferi de kock.qxd 2005/12/09 10:01 page 5 koedoe 45/2 (2002) 6 issn 0075-6458 table 1 (continued) rivers shitlhave dam silolweni dam vervoer dam grid reference 25°08'38"s; 31°19'31"e 24°29'21"s; 31°50'11"e 25°07'07"s; 31°29'29"e 1995 melanoides tuberculata spathopsis petersi corbicula fluminalis no molluscs no molluscs no molluscs 2001 lymnaea natalensis melanoides tuberculata biomphalaria pfeifferi (shell) bulinus forskalii burnupia caffra lymnaea columella aplexa marmorata discussion from the results of this investigation it is clear that the high rainfall average recorded for 1995-2001 period had a significant influence on the number of habitats from which molluscs were recorded as compared to the 1995 survey. for the majority of the habitats the mean rainfall for the 2001 period was nearly double the figures recorded for the 1995 period. the decrease in snail bio-diversity in the sabie and nwaswitsiontso rivers and orpen dam (which forms part of the nwaswitsontso river drainage system, as previously mentioned) could probably be ascribed to heavy floods in these habitats prior to the present survey. in the same localities investigated, b. forskalii was found only four times in 1995 as compared to 16 times in 2001 and was, furthermore, present in nine of the 18 habitats surveyed for the first time in 2001. this snail is able to exploit a wide variety of natural and artificial habitats, including the margins of lakes and permanent swamps, but is most abundant in small water bodies and was also collected in alluvial pools associated with rice cultivation in gambia (brown 1994). the fact that this snail was found in many more habitats during the 2001 survey may be ascribed to its ability to aestivate through long periods of drought. however, the possibility that some of these habitats were populated by this snail species for the first time due to the high rainfall of the 19952001 period cannot be ruled out. aplexa marmorata, a confirmed invader species collected only once during the 1995 survey, was present in seven of the localities in 2001, which was investigated during both surveys. although it was collected several times during the past 15 years in kwazulunatal, it was recorded only once from the limpopo province (dana 1999) at the time of the present investigation. the fact that this species was found in a number of habitats during the 2001 survey seems to suggest that it could be in the process of becoming well established in the southern parts of the kruger national park. whether it has already invaded habitats in the adjacent swaziland needs further investigation. although this species is usually reported from water bodies modified by man (dana 1999), it was present in several rivers and dams in the kruger national park in contrast to this, p. acuta, which has a worldwide distribution and which is established in many african rivers and lakes from south africa to morocco, and is considered the second most successful invader snail species in south africa (de kock et al. 1989), was recovered from only two habitats during the 2001 survey as compared to the one in 1995. its distribution pattern in the de kock.qxd 2005/12/09 10:01 page 6 issn 0075-6458 7 koedoe 45/2 (2002) table 2 molluscs found in waterbodies of the kruger national park surveyed for the first time in 2001 waterbodies grid reference 2001 grootvlei dam 23°20'41"s; 31°30'56"e spathopsis sp. jones se dam 24°51'23"s; 31°45'13"e bulinus forskalii ceratophallus natalensis burnupia caffra kanniedood dam 23°08'40"s; 31°27'48"e bulinus tropicus bulinus forskalii lymnaea natalensis letaba river 23°45'28"s; 31°22'20"e bulinus forskalii physa acuta melanoides tuberculata mestel dam 25°06'54"s; 31°12'50"e bulinus globosus lymnaea columella aplexa marmorata mpondo dam 25°12'12"s; 31°43'08"e bulinus forskalii lymnaea natalensis lymnaea columella spathopsis petersi mtombeni spruit 24°15'40"s; 31°43'40"e bulinus forskalii nshawu dam 23°31'40"s; 31°29'09"e ceratophallus natalensis olifantsdrinkbad 23°19'45"s; 31°19'44"e bulinus forskalii piet grobler dam 24°13'54"s; 31°38'03"e bulinus forskalii rabelais dam 24°27'30"s; 31°29'58"e bulinus tropicus bulinus forskalii sable dam 23°56'12"s; 31°14'04"e bulinus tropicus bulinus forskalii shisa spruit 22°57'32"s; 31°15'01"e cleopatra ferruginea melanoides tuberculata spathopsis petersi corbicula fluminalis sirheni dam 22°56'57"s; 31°14'02"e bulinus globosus biomphalaria pfeifferi lymnaea natalensis lymnaea columella lanistes ovum sunset dam 25°06'57"s; 31°54'41"e aplexa marmorata table 3 waterbodies of the kruger national park surveyed for the first time in 2001 and in which no molluscs were found waterbodies grid reference luvuvhu river 22°26'38"s; 31°13'02"e ngotso dam 24°10'31"s; 31°43'09"e nhlanganzwane dam 25°14'40"s; 31°58'30"e de kock.qxd 2005/12/09 10:01 page 7 koedoe 45/2 (2002) 8 issn 0075-6458 rsa shows concentrations of collections around major urban areas (brackenbury & appleton 1993). it is now found in various artificial and natural habitat types and taking into account that this species is not highly specific with regard to its habitat preferences it is quite surprising that it has a relatively limited distribution in the kruger national park. however, it occurs commonly in polluted water and has therefore been nicknamed the sewage snail and seems to have difficulties in establishing itself far from human works or habitats (brown 1978; brackenbury & appleton 1993). this could be one of the reasons for its limited occurrence in the kruger national park, especially with regard to the dams and rivers, which originate in or near the park. although the drought of 1995 may also serve as an explanation for this situation, it must be kept in mind that it had been recovered only three times from the kruger national park prior to 1966. lymnaea columella, which is considered the most successful invader snail species in south africa (brown 1967) was recovered from five habitats during the 2001 survey and from only one during the 1995 survey. scrutiny of its distribution in the whole of the rsa (de kock et al. 1989) shows clearly why it has earned the title as the most successful invader snail species, because it has successfully established itself in vast areas of the country since its introduction in the early 1940s. before 1966 it was found in 132 loci, and in 225 additional loci from 1966 up to the present (de kock et al. 1989). this species was first reported from the kruger national park in 1967 from three localities by oberholzer & van eeden (1967) and was collected in only three additional localities during the 2001 survey. in contrast to this, a. marmorata was collected in six additional localities since the 1995 survey. this increase in collection sites over a period of only six years seems to suggest that a. marmorata could become just as successful as l. columella in establishing itself in this country. fig. 3. distribution of physa acuta in 1995 (o) and 2001 (•). rainfall monitoring stations (♦). de kock.qxd 2005/12/09 10:01 page 8 issn 0075-6458 9 koedoe 45/2 (2002) fig. 5. distribution of lymnea columella in 1995 (o) and 2001 (•). rainfall monitoring stations (♦). fig. 4. distribution of aplexa marmorata in 1995 (o) and 2001 (•). rainfall monitoring stations (♦). de kock.qxd 2005/12/09 10:01 page 9 koedoe 45/2 (2002) 10 issn 0075-6458 table 4 rainfall records of the kruger national park at the different monitoring stations nearest to collection sites over different time periods rainfall monitoring station berg-en-dal crocodile bridge houtboschrand king fisher spruit letaba lower sabie mooiplaas nwanetsi pafuri phalaborwa pretoriuskop punda maria satara shingwedzi skukuza tshokwane collection site matjulu spruit mlambane spruit mpanamana dam gezantombi dam nhlanganzwanedam mtombeni spruit ngotso dam timbavati river rabelais dam engelhard dam letaba river olifants river lower sabie rest camp mpondo dam sabie river grootvlei dam nshawu dam olifantdrinkbad pioniers dam tsende drif gudzani dam nwanetsi river nwanetsi water hole luvuvhu river sable dam mbyamiti river mestel dam shitlhave dam klopperfontein dam nkaya pan nsemani dam piet grobler dam kanniedood dam krapkuil dam sirheni dam sisha spruit manzimahle dam olifantdrinkgat skukuza fish pond sunset dam vervoer dam jones se dam kumana dam leeupan mazithi dam nwaswitsontso river orpen dam silolweni dam average rainfall of all records available (mm) a 547 386 523 435 577 449 515 332 523 709 528 526 462 533 533 average rainfall for 1882 1983 (mm) a 380 304 265 228 493 196 308 98 234 417 142 326 248 272 295 average rainfall for 1985 1987 (mm) a 521 355 545 335 536 447 437 422 484 428 520 428 356 547 547 average rainfall for 1991 1994 (mm) a 506 279 440 281 441 281 376 297 412 507 327 507 334 411 411 average rainfall for 1995 2001 (mm) 832 814 560 673 418 787 663 653 652 673 989 722 595 704 674 761 de kock.qxd 2005/12/09 10:01 page 10 with regard to lanistes ovum peters in troschel, 1845 it needs to be mentioned that large specimens were collected before 1966, while none were found during the 1995 survey. however, juvenile specimens were found in the sirheni dam during the 2001 survey which might indicate that this species could be in the process of recovering from the previous droughts. more or less the same conclusions could be drawn for the bivalves on record for the park in general and the spathopsis spp. in particular. conclusions from the results of this investigation it is clear that there was a significant increase in the number of localities which yielded molluscs during the present study as compared to the 1995 survey. this could mainly be ascribed to the abnormal high rainfall during 1995-2001 period. with regard to the invader species a. marmorata, a follow-up survey in about five years’ time is recommended to establish whether the northern parts of the kruger national park are also going to be invaded by this species. acknowledgements the authors wish to thank the following institution and persons for assistance: south african national parks for permission to conduct the research; dr andrew deacon for coordinating this project on behalf of the kruger national park and mr nick zambatis for kindly making the necessary rainfall data available. references brackenbury, t.d. & c.c. appleton. 1993. recolonization of the umsindusi river, natal, south africa, by the invasive gastropod, physa acuta (basommatophora, physidae). journal of medical & applied malacology 5:39-44. brown, d.s. 1978. 35. freshwater molluscs. pp. 1155-1180. in: werger, m.j.a. (ed.). biogeography and ecology of southern africa 1. the hague: junk. brown, d.s. 1994. freshwater molluscs of africa and their medical importance. london: taylor & francis. dana, p. 1999. aplexa marmorata (guilding, 1828) (basommatophora: physidae) — an invasive freshwater snail in south africa. m.sc thesis, university of natal, durban. davies, b. & j. day. 1989. vanishing waters. cape town: university of cape town press. de kock, k.n., p.h. joubert. & s.j. pretorius. 1989. geographical distribution and habitat preferences of the invader freshwater snail species lymnaea columella (mollusca: gastropoda) in south africa. onderstepoort journal of veterinary research 56: 271-275. de kock, k.n. & c.t. wolmarans. 1989. a reevaluation of the occurrence of freshwater molluscs in the kruger national park. koedoe 41:1-8. oberholzer, g. & j.a. van eeden. 1967. the freshwater molluscs of the kruger national park. koedoe 10: 1-42. zambatis, n. & h.c. biggs. rainfall and temperatures during the 1991/92 drought in the kruger national park. koedoe 38: 1-16. issn 0075-6458 11 koedoe 45/2 (2002) de kock.qxd 2005/12/09 10:01 page 11 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <feff004e00e4006900640065006e002000610073006500740075007300740065006e0020006100760075006c006c006100200076006f0069006400610061006e0020006c0075006f006400610020005000440046002d0061007300690061006b00690072006a006f006a0061002c0020006a006f006900640065006e002000740075006c006f0073007400750073006c00610061007400750020006f006e0020006b006f0072006b006500610020006a00610020006b007500760061006e0020007400610072006b006b007500750073002000730075007500720069002e0020005000440046002d0061007300690061006b00690072006a0061007400200076006f0069006400610061006e0020006100760061007400610020004100630072006f006200610074002d0020006a0061002000520065006100640065007200200035002e00300020002d006f0068006a0065006c006d0061006c006c0061002000740061006900200075007500640065006d006d0061006c006c0061002000760065007200730069006f006c006c0061002e0020004e00e4006d00e4002000610073006500740075006b0073006500740020006500640065006c006c00790074007400e4007600e4007400200066006f006e0074007400690065006e002000750070006f00740075007300740061002e> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice abstract introduction conclusion acknowledgements references about the author(s) jane carruthers department of history, university of south africa, pretoria, south africa centre for invasion biology, stellenbosch university, stellenbosch, south africa llewellyn c. foxcroft centre for invasion biology, stellenbosch university, stellenbosch, south africa conservation services, south african national parks, skukuza, south africa citation carruthers, j. & foxcroft, l.c., 2019, ‘koedoe: african protected area conservation and science – a retrospection: 1958 to 2018’, koedoe 61(1), a1556. https://doi.org/10.4102/koedoe.v61i1.1556 essay koedoe: african protected area conservation and science – a retrospection: 1958 to 2018 jane carruthers, llewellyn c. foxcroft received: 19 sept. 2018; accepted: 19 nov. 2018; published: 08 apr. 2019 copyright: © 2019. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract in 2018, koedoe celebrated an unbroken publication record of 60 years. from uncertain beginnings in 1958, it is now a mature and important internationally recognised scientific journal focussing on conservation and science in national parks in south africa and beyond into the african continent. after an overview of the emergence of national parks in the 20th century, this retrospective essay reflects on koedoe’s long and significant contribution to the field of national parks research and management. we identify and make easily available some of the seminal and influential articles that have appeared in the journal over this long time span. principally relating to matters in national parks in south africa, these articles (some coming from special issues of koedoe) have been chosen for their variety as well as for the broader perspectives they open into the longer trajectory of national park conservation and management. articles illustrate the evolution of paradigms from protectionist and species centric, to ecosystem focus, to complex socio-ecological systems and adaptive management. conservationists, scientists and managers alike will benefit from an understanding of the transformations in their field over six decades together with appreciating the importance and usefulness of unpacking the intellectual journey of national park science in order to contextualise and enrich – even encourage and direct – present and future research. conservation implications: the articles included in this essay produced important information that informed and guided later research. introducing these contributions to a fresh audience we hope will also tempt readers to consult other back issues of this journal, which will benefit conservation by providing an understanding of the long-term transformations in the field. keywords: biodiversity conservation; conservation journals; conservation science; ecology; environmental history; environmental science; south african national parks. introduction an anniversary is conventionally an occasion on which to celebrate, but it is also a time for reflection and for taking stock. the 60th anniversary of the publication record of the journal koedoe: african protected area conservation and science is no exception. after six decades, koedoe is a mature and important journal. for this commemorative retrospective issue we have decided to republish a number of papers on seminal work from the distant and more recent past that give a small taste of the contribution that koedoe has made to the literature over 60 years. we also hope that this will tempt readers to turn to other articles in order to follow long-term threads in their own disciplines – as well as others – so as to appreciate how patterns have emerged and paradigms and priorities have changed. all too often science in the past is forgotten, or ignored, while some of the key work that has laid the foundation for future scientific endeavours lies in the rich knowledge accumulated over many decades. the 1950s were important in the history of national parks worldwide (carruthers 2017), and the remarkable longevity of koedoe is a tribute to the enthusiasm of biologists during that decade in south africa. in 1958, it was a novelty – indeed a very bold step – to publish any research conducted in a national parks organisation, and it is greatly to the credit of south african national parks (sanparks) that support for koedoe has never wavered. national parks have a pedigree dating back to the 19th and early 20th centuries. however, it was only after world war ii that, together with a rash of other institutions designed to herald a new era, the international union for the protection of nature (iupn) was founded at a meeting on 05 october 1948 at fontainebleau, france, under the auspices of unesco (iupn 1948). no delegate from southern africa was present at that event, but before long there was strong engagement from the subcontinent. throughout the 1950s and, indeed, beyond and into the present, south africans – some affiliated to the national parks board (npb), renamed sanparks in 1996 – played a role in shaping many of the international union for the conservation of nature and natural resources’ (iucn) initiatives and policies as they carved a reputation for national park science at home. in 1958 at the 6th general assembly in athens, the international union for the conservation of nature and natural resources (iucn, to which the iupn changed its name in 1956; iucn 1956) established a provisional commission on national parks (resolution 247), planned regular world congresses and decided to compile a list of the world’s national parks. interestingly, however, there was then no mention of defining a national park or any other protected area, although this, together with other bureaucratic steps, was to come in time. a number of south africans at the athens meeting participated in a variety of committees; among them were rocco knobel, director of the npb, who was elected to the executive board, and his colleague, r.j. labuschagne. representatives of the transvaal and natal conservation authorities were also active. the minutes record discussion on the importance of scientific, especially ecological, research in matters relating to conservation (iucn 1960). within africa, in 1953, the bukavu conference of the iupn had prioritised the protection of the fauna and flora of the continent. importantly too, glimpses of the emerging disciplines of conservation science and, indeed the explosions of the environmental era, were increasingly visible in the 1950s. this was evident in south africa, for example, with the introduction of various environmental or nature and wildlife conservation-oriented courses at universities, the establishment of the percy fitzpatrick institute of african ornithology (1959) and the zoological society of southern africa (1958). south african national parks were in the vanguard of these developments with the establishment of the biology section of the kruger national park (knp) in 1951. after a bruising internal commission of enquiry in 1952, and the consequent professionalisation of the npb, stable management was critical to future scientific endeavour (carruthers 2017). it was into this wider scientific and organisational environment that the first issue of koedoe appeared as an annual journal. koedoe, which is the afrikaans name for tragelaphus strepsiceros, ‘kudu’ in english, had become the emblem of the npb in 1955. there was an english subtitle – the journal was bilingual – journal for scientific research in the national parks of the union of south africa (in afrikaans, tydskrif vir wetenskaplike navorsing in die nasionale parke van die unie van suid-afrika). it is a matter of regret that there was no initial foreword or editor’s message to explain exactly what koedoe aimed to accomplish, and who the intended readership might be. national parks board liaison officer n.j. van der merwe noted in 1962 that koedoe was useful for exchanges and that the library in skukuza had, by this means, received 92 scientific journals, 28 semi-scientific journals and 27 annual reports. this meant significant scientific material was available for research staff to use, thanks to the regular publication of koedoe. a network had been established. the first issue of koedoe was impressive, with articles about an expedition to the lowveld from mozambique in 1725, a list of mammals of the mountain zebra national park, a mosquito survey and fire research in the knp, hyena observations and seven articles on the animals of the kalahari gemsbok national park (kgnp). five contributions were in afrikaans and eight in english. only two, however, were by research staff in the employ of the npb: a.m. brynard (on conditions in the kgnp) and h.p. van der schijff (fire research in the knp). at the time, there were five national parks under the banner of the npb: kruger, bontebok, addo elephant, kalahari gemsbok and mountain zebra, and the staff was very small; since then, the number of parks has grown to 19 and staff numbers have burgeoned. with the passing years koedoe has become a highly respected, peer reviewed, open access academic journal of a kind that could never have been foreseen in the 1950s (foxcroft 2008; foxcroft & herbst 2017). the credibility of koedoe was further strengthened in 2013 when koedoe received its first impact factor of 1.48, fluctuating to the current rating of 0.9 (clavariate analytics journal citation reports). moreover, although koedoe has always welcomed contributions beyond national parks, its current subtitle african protected area conservation and science reflects its formally changed focus to cover the entire continent and all kinds of protected areas, not only national parks and not only about south africa. not surprisingly, the content of koedoe has varied enormously in scope and quality since 1958: there have been substantial articles and extensive lists but also, often, short pieces of a page or two about an unusual sighting or a new record. naturally, as a largely in-house journal until recently, the interest, importance, topicality or institutional relevance of an article has sometimes exceeded the value of the research it communicated. in addition, although research articles were mostly reviewed, professional, rigorous, peer review was not the requisite that it is now. nonetheless, one must emphasise the novelty and importance of such an interdisciplinary research journal, as well as its pioneering role in providing a platform for disseminating the knowledge generated in or about national parks in our country. as a record of a long period of locality-based research, koedoe is an incomparably rich resource, and it records the names of the most eminent scientists in south african national park science, both within sanparks and outside it. the short selection of just more than a dozen past articles included in this celebratory issue is, naturally, subjective, perhaps idiosyncratic. it was no easy task to decide what to include from the more than 1250 articles – for koedoe is brim-full of interesting contributions, all of which have a point to make. however, we have focused on some of those that, in our view, capture changes in the emphasis, or type, of scientific research conducted in south africa’s national parks and the philosophy that underpinned research and management (see figure 1). we have also sought out different disciplines and tried to convey a little of the variety of national parks that have been highlighted in koedoe since 1958. figure 1: a sample of articles discussed in this article and journal editors from koedoe’s 60-year history. in addition, we want to flag special issues of koedoe, the contents of which examine a topic or national park in detail and that are certainly worth revisiting, even re-enacting. the earliest was in 1977, ‘proceedings of the symposium, “the state of nature conservation in southern africa”’, held to commemorate the 50th anniversary of the south african national parks act of 1926. the meeting was a large affair held in the knp with about 200 delegates, of whom some 18 were black africans, and there was one lone woman. many parts of southern africa had a presence, including swaziland, rhodesia, lesotho and botswana, as well as qwaqwa, lebowa, ‘vendaland’, ‘zululand’, ciskei and other ‘self-governing’ states in south africa. the proceedings encompassed wilderness, education, private nature reserves, the status of nature conservation in various states and regions, tourism, marine parks, and botanical and zoological gardens. the 1977 symposium was path-breaking in including landscape planning as part of the general brief for protected areas, as was a call from the director of the newly established department of planning and the environment, who urged national park authorities to work with all the other state departments so that nature conservation would become part of a coordinated national system of land-use planning. of the many contributions to the 1977 symposium, one of the few to deal explicitly with research was u. de v. (tol) pienaar’s (1977) ‘research objectives in south african national parks’. for this reason, and as pienaar was one of the foremost scientists and managers of the npb, becoming warden of the kpn and later chief director of the npb, it has a place in our selection. pienaar (1977) ventured that: … the primary objectives of any research program in our national parks should therefore be a study of ecosystems and their interrelationships, with the objective of supplying the managerial basis for maintaining optimum conditions within the ecosystems, without endangering the interdependency of the components or the quality of the environment. (p. 39) the occasion recorded in koedoe 1977 served as a mirror of the southern african nature conservation community at that time, and perhaps an analogous meeting for our era should be put on the agenda, given the immensity of the environmental, scientific and political changes since the 1970s and the challenges that confront the conservation science community in the 21st century. (pienaar, u.d.v., 1977, ‘research objectives in south african national parks’, in proceedings of a symposium on the state of nature conservation in southern africa, kruger national park, 1976, g. de graaff & p.t. van der walt (eds.), supplement, koedoe 20(2), 38–48. https://doi.org/10.4102/koedoe.v20i2.1266) a decade later, in 1986, another article in koedoe also dealt with the wider principles of national park science. the author was salomon joubert, who was later to become warden of the knp. in ‘management and research in relation to ecosystems of the kruger national park’, joubert (1986) maintained the knp’s commitment to preserving and perpetuating ecosystems. this was to be done because these ‘living organisms’ (biodiversity only entered the lexicon of koedoe in 1995) counteracted the depletion of natural resources and provided recreational facilities for ‘modern day technological man’. management options were to be guided by the ‘minimum intervention’ principle, although other than emphasising monitoring, precisely how and when such intervention might be scientifically determined was left unstated. (joubert, s.c.j., 1986, ‘management and research in relation to ecosystems of the kruger national park’, koedoe 29, 157–163. https://doi.org/10.4102/koedoe.v29i1.528) the increasing complexity of national park science was aired in an important special issue in 2011. by then the time had arrived – somewhat haltingly – for an entirely new approach to science and management in south africa’s national parks. as the old guard left with changes to the country in 1994, and many staff and board retirements coincided, a new generation of scientists and managers proved ready to strike out in a new direction. that change was made especially noticeable through the adoption of strategic adaptive management (sam), in place of the command and control paradigm that had been introduced in the 1960s. terminology began to change from ‘national parks’ and ‘ecosystems’ to be replaced by ‘protected areas’ and ‘complex social ecological systems’. the 2011 special issue of koedoe gave considerable attention to dissecting sam, and we reproduce here the introduction to the volume by dirk roux and llewellyn foxcroft (2011). (roux, d.j. & foxcroft, l.c., 2011, ‘the development and application of strategic adaptive management within south african national parks’, koedoe 53(2), 5 pages. https://doi.org/10.4102/koedoe.v53i2.1049) two other koedoe special issues are also worth noting. in 1983 a symposium was held in pretoria on the kalahari ecosystem, and the voluminous proceedings were published the following year. the profile of the kgnp has faded in recent years, but it was once extremely high in npb research circles, and the idea of an ecosystem-wide research think tank is certainly a worthwhile project even now. the large 1983 meeting was a tribute to dr f.c. (fritz) eloff, chairman of the npb since 1979 and soon to retire as professor of zoology at the university of pretoria. eloff had taken a close interest in the lions of the kgnp and became a consultant for the cat specialist group as well as for the survival service commission (later renamed the ‘species survival commission’) of the iucn. the contents of this special issue were wide-ranging and included articles on geology and soil; water resources and vegetation; fire; mammal, bird and invertebrate studies; as well as a list of relevant publications on the kgnp. as anthony hall-martin (1984) pointed out in his conclusion, the symposium and the publication provided a welcome synthesis of the state of knowledge about the kgnp, while also identifying important gaps. (hall-martin, a.j., 1984, ‘symposium on the kalahari ecosystem: summary and conclusions’, in ‘proceedings of a symposium on the kalahari ecosystem 1983’, eds g. de graaff and d.j. van rensburg, supplement, koedoe, 327–333. https://doi.org/10.4102/koedoe.v27i2.589) the second special issue that opens a window into national park science is more recent – 2014. edited by stephen mccool and anna spenceley (2014), two members of the iucn’s tourism and protected areas specialist group, world commission on protected areas, this highlighted the importance of tourism, as a research science, to aid funding and capacity-building. it further emphasised the importance of national parks to local communities and local and international visitors. the introductory article, ‘tourism and protected areas: a growing nexus of challenge and opportunity’, is reproduced here. one article in the issue is of particular importance to south africa, and it is pleasing to see the wide range of international case studies that are presented. (mccool, s. and spenceley, a., 2014, ‘tourism and protected areas: a growing nexus of challenge and opportunity’, in mccool, s. & spenceley, a., eds, ‘tourism and protected areas: a growing nexus of challenge and opportunity’, koedoe 56(2), art. #1221, 2 pages. https://dx.doi.org/10.4102/koedoe.v56i2.1221) south african national parks were once well known (and vilified) internationally for policies that manipulated the environment, provided water, fresh seasonal grazing thanks to a regulated fire regime and that aimed to keep large mammal populations at predetermined annual levels. koedoe appeared in the era when drugs and darting of wildlife became possible, and this soon became both refined and routine, enabling selected wild animals to be translocated and thus saved from extinction, to repopulate areas where they once occurred and to be culled. between 1960 and 1990, koedoe published nearly 20 articles on marking and darting, explaining the drugs applicable to certain species and their advantages and disadvantages. in preparation for the culling programme, pienaar, van wyk and fairall (1966) published ‘an aerial census of elephant and buffalo in the kruger national park and the implication thereof on intended management schemes’ in koedoe. ending elephant culling was controversial and difficult as the koedoe article by whyte et al. (1999) explains. (pienaar, u. de v., van wyk, p. & fairall, n., 1966, ‘an aerial census of elephant and buffalo in the kruger national park and the implication thereof on intended management schemes’, koedoe 9, 40–107. https://doi.org/10.4102/koedoe.v9i1.781, and whyte, i.j., biggs, h.c., gaylard, a. & braack, l.e.o., 1999, ‘a new policy for the management of the kruger national park elephant population’, koedoe 42(1), 111–32. https://doi.org/10.4102/koedoe.v42i1.228) the small group of scientists employed in the npb, together with outside experts, generally invited, were diligent in publishing their work in koedoe. it is not surprising that for at least the first 30 years, most articles dealt with individual species, of which mammal species were the most numerous by far. other taxa were not, however, totally neglected: invertebrates, birds and fishes were included by way of checklists, surveys and species descriptions. given the international focus on endangered animals, this listing of species and, where possible, recording their abundance or scarcity coincided with the initial publication of the iucn red data lists that began in 1964. a creative research article that brought together the elements of an ecosystem in a national park was that by peter novellie (1987). it has been influential, often cited, and is important for its relevance to an endangered species that npb struggled for decades to preserve. (novellie, p., 1987, ‘interrelationships between fire, grazing and grass cover at the bontebok national park’, koedoe 30, 1–17. https://doi.org/10.4102/koedoe.v30i1.498) veld-burning (now called ‘fire ecology’) was given attention in the very first issue of koedoe in 1958. in an extended article, van der schijff (1958) explained why regulated burning was necessary, using four veld types for his experiment. it is fascinating to compare his article with that 40 years later by van wilgen, biggs and potgieter (1998). (van der schijff, h.p., 1958, ‘inleidende verslag oor veldbrandnarvorsing in die nasionale krugerwildtuin’, koedoe 1, 60–94. https://doi.org/10.4102/koedoe.v1i1.865 and van wilgen, b.w., biggs h.c. & potgieter, a.l.f., 1998, ‘fire management and research in the kruger national park, with suggestions on the detection of thresholds of potential concern’, koedoe 41, 69–87. https://doi.org/10.4102/koedoe.v41i1.248) invasion ecology (or invasion biology) is a science that has become a priority within sanparks since 1958. despite managing for a ‘pristine’ ecosystem, alien flora quickly invaded the knp and other national parks, and a list was published in koedoe in 1988. at that time, there were 156 alien plant species recorded in the knp, of which 113 were considered ‘invasive’. the authors, ian macdonald and willem gertenbach (macdonald and gertenbach 1988), warned that a coordinated regional campaign to control them would be required. the invasion of alien biota, however, seems unstoppable and it is salutary to compare the 1988 article with that by foxcroft (2009) in koedoe. (macdonald, i.a.w. & gertenbach, w.p.d., 1988, ‘a list of alien plants in the kruger national park’, koedoe 31, 137–150. https://doi.org/10.4102/koedoe.v31i1.491, and foxcroft, l.c., 2009, ‘developing thresholds of potential concern for invasive alien species: hypotheses and concepts”, koedoe 51(1), art. #157, 6 pages. https://doi.org/10.4102/koedoe.v51i1.157) the expansion of the national park system under sanparks over the past two decades has provided opportunities for a wider variety of scientific research for inclusion in koedoe, although some of the newer parks have yet to find a research advocate and provide more than checklists of biota. the knp continues to dominate koedoe and indeed the research agenda of sanparks. however, as the garden route national park took shape – including the lake systems – the protection and study of these water bodies became critically important. ian russell’s (1999) article entitled ‘changes in the water quality of the wilderness and swartvlei lake systems, south africa’ is well worth reading, as is his (2002) article entitled ‘freshwater fishes of tsitsikamma national park’. (russell, i.a., 2002, ‘freshwater fishes of tsitsikamma national park’, koedoe 45(2), 13–17. https://doi.org/10.4102/koedoe.v45i2.34, and russell, i.a., 1999, ‘changes in the water quality of the wilderness and swartvlei lake systems, south africa’, koedoe 42(1), 57–72. https://doi.org/10.4102/koedoe.v42i1.222) the very first article in the 1958 line-up in koedoe was by willem h.j. punt (1958), a local antiquarian with a flair for colonialist micro-history, particularly the minutiae of how explorers opened up the southern african interior to ‘white civilisation’. the object of punt’s attention was a group from the netherlands that briefly penetrated the lowveld from lourenço marques (maputo) in mozambique, thereby, in his view, initiating a european presence of long standing in the vicinity of the knp. naturally, this kind of historical research that valorised white conservation efforts chimed with the socio-political era of apartheid in the 1950s. not until the 1990s was another historical perspective introduced to koedoe with jane carruthers’s (1993) article on the contributory role of africans in national park developments. regrettably, the number of papers in koedoe on historical and archaeological topics relating to sanparks has diminished over the years, although these are critically important scientific disciplines in protected areas and humanise the landscape. (punt, w.h.j., 1958, ‘die verkenning van die krugerwildtuin deur die hollandse oos-indiese kompanje, 1725‘, koedoe 1, 1–18. https://doi.org/10.4102/koedoe.v1i1.863, and carruthers, j., 1993, ‘“police boys” and poachers: africans wildlife protection and national parks, the transvaal 1902–1950’, koedoe 36(2), 11–22. https://doi.org/10.4102/koedoe.v36i2.371) conclusion while a large portion of articles published in koedoe have, understandably, emanated from kruger, here we have included the kalahari gemsbok, bontebok and tsitsikamma (now garden route) national parks to provide other perspectives. the kalahari gemsbok had a multifaceted and productive research programme, while work at bontebok illustrates early grappling with an ecosystems approach – fire, grazing and grass interactions. the special issues of koedoe were influential in that they brought together a wide range of thinking and research on specific and very relevant topics, thus providing accessible information that might, more often, only be found in a book. the 1977 symposium on the state of conservation resulted in a large workshop of practitioners who debated the merits and flaws of the science at the time and suggested possibilities for the future. over the years, tourism has been the financial bedrock of national parks and the science around that discipline is an expanding and important field. of equal importance is the need for humanity to interact with nature in an era when urbanisation is burgeoning, and this is provided by visitor access. the special issue on adaptive management is vitally important because it has been promoted in recent years as a favourable method of managing adaptive, complex social-ecological systems. sanparks has been hailed as a forerunner in the continued development and implementation of adaptive management and articles from the special issue have been downloaded ~225 000 times (as of march 2018). we hope that the articles we have highlighted will inspire readers to consult the older issues of the journal and that charting the changing contours of national park science will be informative and instructive, as well as illuminate how fresh and complex ecological fields, such as fire ecology and invasion biology, have emerged as large areas of research. no doubt, the learning process will continue. the ease of open access through the koedoe website, and the publication of a number of indexes and statistical material, will make further search easy and rewarding. change is a certainty, and we can be sure that the scientific record of today that is considered to be unassailable will also alter with time. acknowledgements the authors thank the editor and anonymous reviewers for their comments. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions j.c. identified the articles reviewed and wrote the first draft of the manuscript. l.c.f. collaborated on identifying the articles reviewed and wrote the manuscript. funding information koedoe is supported by south african national parks. references carruthers, j., 1993, ‘“police boys” and poachers: africans wildlife protection and national parks, the transvaal 1902–1950’, koedoe 36(2), 11–22. https://doi.org/10.4102/koedoe.v36i2.371 carruthers, j., 2017, national park science: a century of research in south africa, cambridge university press, cambridge, united kingdom. foxcroft, l.c. & herbst, m., 2017, ‘status and trends in the global growth of koedoe between 1958 and 2016’, koedoe 59(1), 5 pages. https://doi.org/10.4102/koedoe.v59i1.1500 foxcroft, l.c., 2008, ‘fifty years of koedoe: current status and future directions’, koedoe 50(1), art. #124, 2 pages. https://doi.org/10.4102/koedoe.v50i1.124 foxcroft, l.c., 2009, ‘developing thresholds of potential concern for invasive alien species: hypotheses and concepts”, koedoe 51(1), art. #157, 6 pages. https://doi.org/10.4102/koedoe.v51i1.157 hall-martin, a.j., 1984, ‘symposium on the kalahari ecosystem: summary and conclusions’, in proceedings of a symposium on the kalahari ecosystem 1983, g. de graaff & d.j. van rensburg (eds.), supplement, koedoe 27(2), 327–333. https://doi.org/10.4102/koedoe.v27i2.589 iucn (international union for the conservation of nature and natural resources), 1960, proceedings of the 6th general assembly, athens, 11–19 september 1958, (brussels, iucn, 1960), viewed 30 january 2019, from https://portals.iucn.org/library/efiles/documents/ga-6th-005.pdf. iupn (international union for the protection of nature), 1948, international union for the protection of nature established at fountainebleau, 05 october 1948, viewed 30 january 2019, from https://portals.iucn.org/library/efiles/documents/1948-001.pdf joubert, s.c.j., 1986, ‘management and research in relation to ecosystems of the kruger national park’, koedoe 29, 157–163. https://doi.org/10.4102/koedoe.v29i1.528 macdonald, i.a.w. & gertenbach, w.p.d., 1988, ‘a list of alien plants in the kruger national park’, koedoe 31, 137–150. https://doi.org/10.4102/koedoe.v31i1.491 mccool, s. & spenceley, a., 2014, ‘tourism and protected areas: a growing nexus of challenge and opportunity’, in s. mccool & a. spenceley (eds.), tourism and protected areas: a growing nexus of challenge and opportunity, koedoe 56(2), art. #1221, 2 pages. https://doi.org/10.4102/koedoe.v56i2.1221 novellie, p., 1987, ‘interrelationships between fire, grazing and grass cover at the bontebok national park’, koedoe 30, 1–17. https://doi.org/10.4102/koedoe.v30i1.498 pienaar, u.d.v., 1977, ‘research objectives in south african national parks’, in proceedings of a symposium on the state of nature conservation in southern africa, kruger national park, 1976, g. de graaff & p.t. van der walt (eds.), supplement, koedoe 20(2), 38–48. https://doi.org/10.4102/koedoe.v20i2.1266 pienaar, u.d.v., van wyk, p. & fairall, n., 1966, ‘an aerial census of elephant and buffalo in the kruger national park and the implication thereof on intended management schemes’, koedoe 9, 40–107. https://doi.org/10.4102/koedoe.v9i1.781 punt, w.h.j., 1958, ‘die verkenning van die krugerwildtuin deur die hollandse oos-indiese kompanje, 1725‘, koedoe 1, 1–18. https://doi.org/10.4102/koedoe.v1i1.863 roux, d.j. & foxcroft, l.c., 2011, ‘the development and application of strategic adaptive management within south african national parks’, koedoe 53(2), 5 pages. https://doi.org/10.4102/koedoe.v53i2.1049 russell, i.a., 2002, ‘freshwater fishes of tsitsikamma national park’, koedoe 45(2), 13–17. https://doi.org/10.4102/koedoe.v45i2.34 russell, i.a., 1999, ‘changes in the water quality of the wilderness and swartvlei lake systems, south africa’, koedoe 42(1), 57–72. https://doi.org/10.4102/koedoe.v42i1.222 van der schijff, h.p., 1958, ‘inleidende verslag oor veldbrandnarvorsing in die nasionale krugerwildtuin’, koedoe 1, 60–94. https://doi.org/10.4102/koedoe.v1i1.865 van wilgen, b.w., biggs, h.c. & potgieter, a.l.f., 1998, ‘fire management and research in the kruger national park, with suggestions on the detection of thresholds of potential concern’, koedoe 41, 69–87. https://doi.org/10.4102/koedoe.v41i1.248 whyte, i.j., biggs, h.c., gaylard, a. & braack, l.e.o., 1999, ‘a new policy for the management of the kruger national park elephant population’, koedoe 42(1), 111–32. https://doi.org/10.4102/koedoe.v42i1.228 abstract introduction literature background research methods and design results discussion conclusion acknowledgements references about the author(s) petrus van der merwe school of tourism management, tourism research in economics, environs and society, faculty of economic and management sciences, north-west university, potchefstroom, south africa melville saayman school of tourism management, tourism research in economics, environs and society, faculty of economic and management sciences, north-west university, potchefstroom, south africa elricke botha department of applied management, college of economic and management sciences, university of south africa, pretoria, south africa citation van der merwe, p., saayman, m. & botha, e., 2020, ‘interpretation needs and preferences of visitors to kgalagadi transfrontier park’, koedoe 62(1), a1613. https://doi.org/10.4102/koedoe.v62i1.1613 original research interpretation needs and preferences of visitors to kgalagadi transfrontier park petrus van der merwe, melville saayman, elricke botha received: 05 dec. 2019; accepted: 17 june 2020; published: 11 aug. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract south african national parks is one of south africa’s main drawcards for ecotourists. one of these parks is the kgalagadi transfrontier park (ktp), which is also a world heritage site (whs). because tourism is key to national parks and whss, visitor management strategies such as interpretation are vital for these destinations to provide tourists with better experiences and knowledge. the aim of this research was therefore to determine the interpretation needs and preferences of visitors to ktp. the factor analyses revealed that tourists felt that park guidelines would add the most to a quality experience. tourists preferred interpretation topics on nature and heritage and would like to receive in-depth information on these topics. interpretation should also preferably be offered through audio and visual media. these findings impact the development of relevant interpretation services for the park, which will contribute towards visitors’ experiences. the research is original as it examined visitors’ interpretation needs and preferences on topics, the extent of information tourists would like to receive and the type of media through which information should be presented. conservation implications: designing relevant interpretation services based on tourists’ needs and preferences will contribute towards satisfactory experiences. in turn, this may lead to an increase in profits, which can support conservation practices. effective interpretation messages can also foster additional support towards conservation efforts. keywords: ecotourism; wildlife tourism; national parks; education; interpretation; world heritage site. introduction the main objective of most national parks in the world, and in south africa specifically, is to conserve the environment or biodiversity (south african government 1976), but this has proven to be quite a difficult task to perform because of declining public funding (eagles 2014). south african national parks (sanparks; the managing body of south african national parks) is therefore primarily a self-funding entity, as ecotourism activities generate 80% of the total income of national parks (sanparks 2018). ecotourism thus contributes to sustaining the conservation objectives of the national parks (sanparks 2018). considering that there has been a decline in public funding and that ecotourism activities generate such a large percentage of sanparks’ income (80%), its focus concerning visitor management is to manage visitor experience through soft visitor management (e.g. influencing enjoyment, attitudes and values through displays, signage or the website, to name but a few) and hard visitor management (e.g. regulating human impact through physical facilities such as bird hides or boardwalks, to name but a few) (sanparks 2018). the manager of visitor management and interpretation of sanparks’ tourism division stated that interpretation has been identified by sanparks as a means of not only adding to the funding required for conservation, but in itself also contributing towards conservation and the tourism experience (moore 2017; personal interview 17 january 2017). the visitor manager further indicated that the requirement is to deliver well-planned and carefully designed interpretation services to help visitors understand what happens in national parks and also educate them on conservation, culture and ecotourism. the park selected as a case study for this research was kgalagadi transfrontier park (hereafter ktp), as it had been identified by sanparks’ management as a priority park for developing interpretation services (moore 2017). kgalagadi transfrontier park has a unique cultural heritage and rich history, but also varied biomes and plentiful fauna and flora (south africa 2019), which draw visitors to the park. visitors want information about these aspects, but the park has limited facilities and services to satisfy their needs. at present only three tourist camps, twee rivieren, nossob and mata mata, have information and/or interpretation centres; however, they are very basic with nothing but a few displays and posters (moore 2017). moreover, the park was added to the united nations educational, scientific and cultural organisation (unesco) world heritage site (whs) list in 2017 (unesco 2019a), which inevitably will increase tourist interest in the park. in addition, the protected areas act (57 of 2003) indicates that protected areas (south africa 2003) such as ktp are mandated to offer visitor facilities and that such facilities should be established in response to market demands. it is against this backdrop that the park has considered refining their interpretation services based on visitors’ needs and preferences, hence this research. determining visitors’ needs and preferences for interpretation services will assist sanparks to develop relevant interpretation services for the park that will contribute towards the visitor experience. literature background educational experiences within ecotourism it is well known that education and learning form part of an ecotourism product. this is evident in both the range of definitions of ecotourism as well as research conducted on the topic. ceballos-lascurain (1987), the first author to do so, defined ecotourism as travelling to relatively undisturbed or uncontaminated natural areas for studying, admiring and enjoying the scenery, its wild plants and animals, or any past and existing cultural manifestations found in the area. fennell (2003), on the other hand, defines ecotourism as a sustainable form of natural resource-based tourism that focuses primarily on experiencing and learning about nature and should contribute to the conservation of such areas. geldenhuys (2009) concurs with fennell’s (2003) definition and explains that ecotourism occurs in natural settings that are ecologically sustainable and enables tourists to interpret and learn about the environment they are visiting (i.e. environmental education). the notion of learning in ecotourism is also evident in the range of research studies conducted on ecotourists. even though ecotourists are not homogenous, research has shown that most ecotourists prefer information-rich experiences. sheena, mariapan and aziz (2015) found that ‘hard’ ecotourists rated ‘tendency to learn’ significantly higher than other ecotourists, whilst ‘structured’ ecotourists (falling between ‘hard’ and ‘soft’ ecotourists) also indicated a strong preference for the learning component. the quality of a tour guide (verbal transfer of information), the quality of information and digital forms of information were found to be key factors in selecting ecotourism products in the study conducted by cheung and jim (2013). these authors also researched the difference between nature tourists and general tourists in a follow-up study and found that nature tourists had higher expectations of the guide, the information provided and the ecological importance of the destination than general tourists did (cheung & jim 2014). chen and jim (2012) researched the potential value of ecotourism development (i.e. country parks) in hong kong and found that the motivation ‘to experience and learn more about nature’ ranked as the most important factor for ecotourism development, followed by ‘rest and relax in a pleasant setting’. in south africa, ecotourists to the kruger national park expect interpretation services in the park because all three factors identified (i.e. primary, secondary and knowledgeable staff) were revealed to be very important to the quality of visitors’ experience (botha, saayman & kruger 2016). primary interpretation refers to immediately recognised interpretation (e.g. visitor centre), whereas secondary interpretation refers to interpretation that is auxiliary to the broader activity (e.g. commentary on a game drive) or indistinct (e.g. web-based marketing of the game drive). these factors were labelled according to stewart et al.’s (1998) classification (discussed later in the study). interpretation carmody and prideaux (2011) state that interpretation is similar to communication. the reason is that interpretation incorporates all the various ways in which attraction or destination managers (in this case, ktp) seek to communicate with their visitors (moscardo & ballantyne 2008) (see figure 1). interpretation, therefore, also includes a context (i.e. the national park’s wildlife and heritage); a sender (e.g. the national park’s managers through interpretation centres); a message (e.g. information regarding conservation and heritage), which can be verbal or non-verbal; a medium (e.g. a podcast, brochure, game drive or exhibit); a receiver (e.g. the ecotourist); and feedback (e.g. visitor enjoyment) (schiffman & kanuk 2007). figure 1: components of interpretation. what makes interpretation different from other forms of information transfer is that interpretation aims to translate the technical language and content of natural science and related fields into simple ideas and illustrations that any person can understand (ham 1992). against this background, various authors have researched this concept and concluded that successful interpretation should be enjoyable, personally relevant and organised, make use of themes and a variety of presentations or mediums, provide time for reflection (allowing visitors to think deeply and draw their own conclusions), involve all the senses, communicate the significance of the place visited and allow personal control (ballantyne, packer & falk 2011; ham 1992; ham, housego & weiler 2005; tilden 1977). tilden (1977), considered to be the father of interpretation, explains interpretation as an educational activity that aims to reveal meanings and relationships through first-hand experiences, the use of original objects and illustrative media, rather than simply to communicate factual information. from a national park’s perspective, interpretation is a means to manage visitors’ educational expectations, because visitors to national parks are well educated and expect information-rich experiences (botha et al. 2016; jurdana 2009). interpretation services, therefore, add value to the tourist’s experience (ballantyne, packer & sutherland 2011; ham & weiler 2006) and lead to a range of other benefits such as increased satisfaction, loyalty, purchasing and revenue, as well as visitors spending more time in the national park, encouraging other visitors to visit the park and providing positive word-of-mouth referrals for the park (de rojas & camarero 2008; lee 2009; zeppel & muloin 2008). added to the various benefits associated with interpretation is also the potential to improve conservation initiatives. the information communicated to tourists broadens their knowledge about the place they are visiting by focusing the message on the significance of their experiences and deepens their understanding of conservation (reisinger & steiner 2006). this understanding creates respect and concern for species (ballantyne et al. 2011) and leads to support for nature conservation work (zeppel & muloin 2008). as stated by baba dioum, ‘in the end, we will conserve only what we love, we will love only what we understand and we will understand only what we are taught’ (dioum 1968). interpretation can be delivered in a variety of ways. tilden (1977) explains that interpretation can either be attended or unattended. attended interpretation refers to services delivered in person, such as discussions or talks, whereas unattended interpretation (i.e. without personal contact) refers to services delivered through signs, labels, self-guided trails (for hikes or vehicles), exhibits, publications and visitor centres. ward and wilkinson (2006) concur with this classification and refer to interpretation as personal or non-personal. other authors have more elaborate classifications. kuo (2002) based his on orams’ (1996) ecotourism management strategies: hard interpretation includes physical (e.g. pathways or bird hides), regulatory (e.g. rules and regulations) and economic (e.g. fines and fees) management strategies, whereas soft interpretation refers to educational (e.g. behavioural information and education) management strategies (kuo 2002; orams 1996). further to this, stewart et al. (1998) refer to primary, secondary and tertiary interpretation. primary interpretation overlaps with kuo’s (2002) soft interpretation or orams’ (1996) educational management strategy, where the interpretation is conspicuous, noticeable and immediately recognised as interpretation. secondary interpretation ‎is auxiliary to a wider activity and forms an integral feature of the activity. tertiary interpretation is hidden, obscure and indistinct as an interpretive activity and is not always considered to be interpretation. both secondary and tertiary interpretation can therefore be considered to overlap with kuo’s (2002) hard interpretation and orams’ (1996) physical, regulatory and economic management strategies. as visitor services, such as interpretation, should be established in response to market demands (haely, van riper & boyd 2016; south africa 2003), it is imperative to have a closer look at the market’s (i.e. the receiver’s) demands in terms of the components of interpretation to effectively design such services and facilities (i.e. product development). the next sections will therefore present a closer look at the context, message and medium of interpretation for which tourists may have specific needs and preferences. the context of interpretation keyton and beck (2010) explain that the context in communication, and therefore interpretation, is multidimensional because it is composed of all internal and external influences that affect the communication process. destinations typically cannot control, or may find it difficult to control, the internal influences (i.e. within the visitor) that can affect the communication process, but they are in a position to control the external influences. bidder, kibat and fatt (2016) found that some of the top motivations to visit the kinabalu park were destination specific, such as ‘to climb mount kinabalu’, ‘because the park was one of the must-see attractions in sabah’ and ‘to experience the indigenous cultural value of mount kinabalu’. cheung and jim (2014) found that ‘ecological value’ and ‘good destination management’ were the two most important factors in destination choice in hong kong’s conservation areas. the factor ‘ecological importance of destination’ was also positively correlated with willingness to pay (cheung & jim 2013). sheena et al. (2015) concluded that high-interaction programmes involving nature in kinabalu park should be offered because the park enhances a profound spiritual connection. from an interpretation perspective, ecotourism managers should identify the unique destination characteristics related to physical, social, historical, psychological and cultural factors to guide the interpretation topics that could be of interest to tourists. ballantyne, hughes and bond (2016) determined the preferred topics among canterbury cathedral whs owners or managers and reduced the 68 topics to 21. different from this study, however, is that their research was not within a natural destination context such as ktp and was based on the preferences of owners or managers and not tourists. as moscardo (2014) explains, those responsible for interpretation are also the ones in power, and they decide which aspects will be presented to tourists. it is therefore crucial to determine tourists’ preferences in terms of interpretation topics. one such study that did research from a tourist perspective, but within a marine context in new zealand, was that by lück (2015). although tourists were generally satisfied with the whale and dolphin tours, they also wanted to learn more about wildlife and the sea in general (lück 2015), thereby amplifying the fact that tourists might have different preferences than managers or owners. as mentioned by the visitor manager of sanparks, in the case of ktp the context that needs interpretation is the culture of the area (ǂkhomani and the related san people) and the rich fauna and flora of the semi-desert habitat (moore 2017). (more detail on the unique context of this park can be found under the ‘research methods and design’ section.) although this is the case, it would be worthwhile to determine the preferences of tourists in this regard. the message of interpretation as previously explained, ecotourists have a high tendency to learn (sheena et al. 2015) and consequently have higher expectations of interpretation (cheung & jim 2014). careful consideration should therefore be given to the extent of information on the topics selected (see the ‘context of interpretation’ discussed previously) and how the topics are presented in interpretation. whilst topics and themes are often used interchangeably, ham (1992) explains that topics are the subject matter, whilst themes refer to the message about the topic. for example, one of the topics in an interpretation centre might be ‘water’. the theme, on the other hand, places specific emphasis on the message to ‘conserve water’ by providing relevant information to the tourists that is enjoyable. the method in which this message is delivered not only highlights the importance of water but also illustrates how the tourist can contribute to the conservation thereof. it is against this background that the theme refers to the messages of interpretation. according to ham (1992), themes should have the qualities of a story: a beginning and end, moral and little meanings (analogies and concepts) that add to the big meaning. langer (1989), a theorist on mindfulness–mindlessness, explains that environments characterised by novelty, surprise, complexity and ambiguity could induce mindful states (i.e. readiness for cognitive processing). interpretation should also be relevant (ham 1992) by including practical examples of how visitors could contribute towards conservation (ballantyne et al. 2011; marschall, granquist & burns 2017); interpretation should provide personal connection (tilden 1977) and allow visitors to reflect on personal meaning (kolb 1984). interpretation messages that focus on emotions have a greater effect on conservation intentions (jacobs & harms 2014). ballantyne et al. (2011) explain that interpretation should let visitors use their imagination, provide information about the dangers faced by animals as a result of human actions, provide examples of how visitors’ everyday behaviours can have either a positive or negative impact on the animals being observed and supply practical yet achievable examples of what visitors can do to contribute to animal and environmental welfare. ecotourism destinations should therefore not only select their topics carefully, but also consider the theme or message to portray the topic wisely. further advice from ballantyne, packer and hughes (2009) is that ecotourism destinations should make sure that these themes or messages are delivered in a consistent manner, because inconsistencies can undermine the credibility of the interpretive message. to the researchers’ knowledge, little research has been conducted to determine tourists’ preferences in terms of the extent of information on topics for interpretation services. however, marschall et al. (2017) found that teleological signs (i.e. information with explanations) were more effective than ontological signs (i.e. information without explanations) in modifying visitors’ behaviour. this may indicate that visitors rather prefer more information than less information on topics (i.e. the extent of their interest). however, some topics might be of more interest than others and may have different preferences in terms of the extent of information visitors prefer. the medium of interpretation the medium of communication is defined as the physical space, means of sending information or the route the message travels (schindler, ruoppolo & barillari 2010). the medium in interpretation can therefore be seen as the vehicle through which the themes and messages are delivered to tourists (ham et al. 2005). ballantyne et al. (2011) suggest that interpretation design should incorporate multiple senses (sound, smell, textures and visually appealing aspects) and use interpretive commentaries and signage to reinforce a sense of wonder, awe, excitement and privilege (i.e. non-verbal communication); time and space should be created for visitors to reflect on the experience and meaning (i.e. organised interpretation); staff or volunteer guides should be available to answer visitors’ questions (i.e. verbal communication); resources should be provided to visitors to access after the visit to follow up on specific interests (i.e. off-site interpretation). bidder et al. (2016), on the other hand, explain that information should be presented in various forms (print, display and audiovisual shows), and souvenirs should be sold as part of the interpretation experience (i.e. post-experience). interestingly, haely et al. (2016) found that low-intensity technologically driven interpretation displays were preferred over high-intensity displays at the cliffs of moher in ireland’s (i.e. natural area) visitor centre. marschall et al. (2017) explain that illustrations in interpretation signs are necessary, not only to attract visitors but also to enable children to understand the interpretation. the positioning of signs is important to attract visitors, and the height at which these signs are positioned should be suitable for children to admire (marschall et al. 2017). other authors studied the effectiveness of certain mediums. cheung and jim (2013) found that respondents participating in a study in natural environments in hong kong preferred digital over printed interpretation. henker and brown (2011) found that online interpretive programmes (i.e. podcasts) were virtually as effective a means as traditional ranger programmes to foster visitor connections. kang and gretzel (2012) concurred with the findings of henker and brown (2011) and advised that podcast tours developed by multiple narrators using a conversational narration style would probably increase the effectiveness of the visitor’s experience. mitsche et al. (2008) explored e-services (e.g. interactive maps, 3-d applications, virtual tours, online exhibitions, interactive resources, games, online collections and online shops) for interpretation purposes and found them to be effective; they also identified a need to cater for different target groups. as this study aims to determine the interpretation needs and preferences of visitors to ktp, the objective is to provide answers to the following questions to develop relevant interpretation services: what are the interpretation services needed by tourists visiting the national park? what are the topics or types of information visitors are interested in? do visitors prefer specific mediums through which the topics or information should be presented? answering these questions will assist the park in developing successful interpretation services and facilities according to the needs and preferences of tourists and will provide researchers with greater insight into visitors’ needs and preferences concerning interpretation. research methods and design study site the ktp is one of the 19 national parks managed by sanparks, and it is unique in that it is a transfrontier park extending into namibia and botswana. transfrontier parks or transfrontier conservation areas (tfca) are defined as ‘relatively large areas, straddling frontiers between two or more countries and covering large-scale natural systems encompassing one or more protected areas’ (south africa 2004). kgalagadi transfrontier park was the first chapter in tfcas in southern africa and was opened on 12 may 2000 (south africa 2004). as indicated previously, the park was also included on the unesco whs list in 2017, based on the evidence of human occupation from the stone age. it is associated with the ǂkhomani san people, which is a unique african cultural group (unesco 2019a). among the 19 national parks, ktp and mapungubwe national park are the only two national parks inscribed as whss (sanparks 2018). these two national parks feature among other well-known south african unesco cultural sites, such as fossil hominid sites, richtersveld cultural and botanical landscape and robben island (unesco 2019b). study design a quantitative study was conducted by means of a structured questionnaire to determine tourists’ interpretation needs and preferences in ktp. study population and sampling strategy a non-probability sampling method, convenience sampling, was used to conduct the survey. convenience sampling means that the researchers include participants who were the most conveniently available (quinlan et al. 2019). krejcie and morgan (1970) state that when the population is over 1 000 000, the minimum sample size should be 384. the level of precision is 5% to compensate for questionnaires with sampling errors (e.g. questionnaires that are not returned or are incomplete). the research population for this research consisted of overnight visitors to ktp for the period february 2016 to january 2017. the number (n) of visitors was 1983, and the questionnaire was sent to their email addresses by sanparks with a letter explaining the importance and aim of the research. the completed questionnaires received back from respondents totalled 562. questionnaire development the questionnaire consisted of six sections comprising openand closed-ended questions as well as likert and semantic differential scale-type questions. this research formed part of a bigger project, and only selected sections of the questionnaire were used for this article. section a consisted of questions relating to the respondents’ demographic and biographical profiles. section d determined tourists’ agreement with the inclusion of interpretation topics and/or information in a visitor centre (1 = strongly disagree to 5 = strongly agree) and prompted the respondents to indicate to what extent they would like information on the topics and/or information listed in the questions (1 = to a very little extent to 5 = to a very great extent). the variables (i.e. topics) in this section were based on park-specific characteristics such as the fauna and flora of the park (sanparks 2020a), the transfrontier aspects (south africa 2019), and the world heritage listing (unesco 2019a). section e contained questions about the importance of interpretation services for a quality experience in the park as a whole, as well as the importance of specific media in an interpretation visitor centre. both questions had to be answered according to a scale of 1 (not at all important) to 5 (extremely important) and were based on the works of ballantyne et al. (2011), botha et al. (2016), bidder et al. (2016), cheung and jim (2013), henker and brown (2011), jacobs and harms (2014), kang and gretzel (2012), kolb (1984), kuo (2002), langer (1989), marschall et al. (2017), mitsche et al. (2008), orams (1994), stewart et al. (1998) and tilden (1977). data analysis the data obtained from the survey were captured in microsoft excel and subsequently statistically analysed using statistical package for social sciences version 16 (field 2013). the data analysis consisted of three phases. in the first phase, data analysis of the descriptive results (e.g. age, gender, home language and place of residence) revealed the profile (demographics) of the respondents. in the second phase, a principal component analysis with an oblimin rotation and kaiser normalisation was performed on the 14 interpretation service preference items, 18 preferred topics and/or information items, 18 items on the extent of information on the topics and/or information, and 18 preferred medium items in order to improve the interpretability of each factor structure. a kaiser–meyer–olkin (kmo) value > 0.7 is deemed acceptable (kaiser 1974). as a result, all factors with eigenvalues larger than one were used, and all items with a factor loading > 0.3 were considered as contributing towards a factor (field 2013). items with a factor loading > 0.4 that cross-loaded onto two factors were categorised in the factor where interpretability was best. all factors with a cronbach’s alpha above 0.6 and inter-item correlations between 0.15 and 0.55 (cohen 1988) were considered acceptable as reliability indicators. factor scores were calculated as the average of all items contributing to that particular factor to interpret them on the applicable five-point likert scale. ethical considerations the research went through a rigorous ethics application process (the university of south africa, clearance number 2016_cems_esttl_007). although the research involved human participants, the research was deemed ‘low risk’. the questions in the questionnaire did not include any personal identifiers or sensitive information, and even if the informed consent was obtained before a respondent completed a questionnaire, the respondent was entitled to withdraw from the study at any point in time. results this section consists of two parts: first, the profile of the respondents, and second, the factor analysis-based preferences for interpretation services, topics and/or information, the extent of information and interpretation mediums. profile of respondents the profile of respondents to the ktp questionnaire can be summarised as male, with an average age of 59, mainly afrikaans or english speaking, from the western cape and gauteng provinces, with a diploma or degree, staying on average 6.4 nights and having visited the park on average seven times. the sociodemographics of the research subjects (profile), namely average age, language spoken, place of residence and level of education, correspond with the results of previous research into the profile (sociodemographics) of visitors to mapungubwe national park conducted by hermann et al. (2015), to kruger national park conducted by saayman and van der merwe (2017), van der merwe, saayman and pienaar (2009) and van der merwe and saayman (2008) and to all south african national parks conducted by de witt, van der merwe and saayman (2015) and du plessis, van der merwe and saayman (2013). the fact that the profile resulting from this study coincides with previous research adds to the validity of the profile of visitors to natural areas in south africa (table 1). table 1: park respondents’ profile. interpretation services the aim of the questions in figure 2 was to determine the interpretation services wanted by tourists to ktp. respondents indicated that they needed park guidelines (factor 1), activity involvement (factor 2) and information structures (factor 3) for a quality experience in the park. all three factors had a cronbach’s alpha loading of above 0.7 with an inter-item correlation above 0.4. a factor loading of 0.30 is considered significant, whilst a factor loading of 0.50 is considered highly significant (field 2013). figure 2: preferred interpretation services. the highest rated factor was factor 1, park guidelines, with a mean value of 4.30. this factor consisted of constructs such as park rules and regulations clearly communicated through signs, brochures and so on (see figure 2). the second most important factor was factor 3, information structures, with a mean value of 4.15. factor 2, activity involvement, received the lowest mean value of the three factors at 3.36. these factors concur with orams’ (1996) and kuo’s (2002) classification of hard interpretation through regulatory practices such as rules and regulations (factor 1) and physical structures such as bird hides (factor 3) and soft interpretation through education practices such as game drives with a field guide (factor 2). information and topics in figure 3, respondents were asked to rate two aspects, namely what information should be presented at interpretation centres (i.e. topics) and the extent to which they would like to receive information on specific topics in interpretation. two factors were identified for both cases: nature and heritage. both factors’ cronbach’s alphas were above 0.8, with inter-item correlations above 0.4. respondents rated nature as the most important factor for both information that should be presented at interpretation centres (mean value 4.23) and the specific topic they would like to receive the most information on (mean value 3.99). therefore, education (interpretation) on nature is key for the interpretation centre in ktp. this research corresponds with the definitions of geldenhuys (2009) and fennell (2003), which highlight that tourism in natural settings must include interpretation and learning about the environment (environmental education). heritage was also rated as significant, with mean values of 3.97 and 3.53. it is therefore also important to present information about the heritage of ktp in its interpretation centres. figure 3: desired topics and extent of information in the interpretation centre. interpretation media media interpretation needed in the park was measured and is presented in figure 4. three factors were identified. factor 1 was named ‘audio and visual’ because the two highest-ranking constructs were audio and video and because the rest of the constructs related to them. factor 2 was named ‘creative text’, as the construct that measured the highest referred to easy, readable text, and the rest of the constructs related to how text is presented. factor 3 was named ‘associations’, as the highest rating construct was souvenirs related to the interpretation, because tourists buy souvenirs to remind or associate themselves with tourism destinations they have visited. all three factors’ cronbach’s alphas were above 0.6, with inter-item correlations above 0.4. the factor with the highest mean value (3.50) was factor 1, audio and visual, followed by factor 2, creative text, with a mean value of 3.49; the last was factor 3, associations, with a mean value of 2.30. figure 4: important media needed for interpretation in the visitor centre at kgalagadi transfrontier park (interpretation centre). discussion the research set out to determine the interpretation needs and preferences of visitors to ktp, and three questions were asked to achieve the aim of the study: what interpretation services do tourists visiting ktp need? what topics or types of information are visitors interested in? do visitors prefer that the topics or information be presented through specific mediums? interpretation services needed by visitors the results revealed that the interpretation services needed are in the form of park guidelines, activity involvement and information structures. park guidelines refer to rules and regulations, clear directions in the park and park maps. this makes perfect sense, as tourists spend most of their time within the park partaking in game drives and less time at the rest camps (maciejewski & kerley 2014). park guidelines are viewed as an inconspicuous form of interpretation, but as kuo (2002) explains, they create an enjoyable visit where visitors can appreciate the value of the site. if all these aspects are well presented, they will influence learning and the overall experience of visitors in the park and therefore contribute to better interpretation within the park. the second aspect, activity involvement, refers to cuisine experiences, events and local culture and therefore indicates that visitors to ktp require interpretation services that focus on the heritage and culture within the park. this was confirmed by mearns and botha (2017), who explain that heritage and culture form part of natural area interpretation. as indicated in the literature, this park has plenty to offer regarding culture and heritage, as it is a whs. currently, as stated by moore (2017), the park lacks interpretation in this regard, thereby confirming what park management expected. information structures are the third interpretation service identified by visitors as important for a quality experience in the park. this includes structures such as bird hides, lookout points, information boards and tree identification. the park does possess a variety of lookout points, picnic areas and hides (sanparks 2020b). management can review these sites and identify ways in which different topics could perhaps be linked through additional approaches. as one would not want to distract from the aesthetics of the natural area, or provide too much or irrelevant information, a possible solution could be to add quick response (qr) scan codes, for example at tree identification nameplates, where tourists can find additional information on the tree. interpretation topics the research revealed that the main interpretation topics visitors are interested in are about nature and heritage and that the extent of information they prefer is quite high. similar to other national parks, ktp was established to protect the natural environment, and it is not surprising that visitors would like to receive information about nature during their stay. however, the park was only recently (i.e. 2017) inscribed as a whs because of the unique cultural landscape, and it was interesting to see the strong preference for heritage information that is currently absent from the park (moore 2017). this is an aspect that requires immediate attention. specific mediums through which the topics or information should be presented the research identified three mediums through which visitors would like to receive interpretation in an interpretation centre, namely audio and visual, creative text and associations. as stated by the visitor manager of sanparks, currently the park lacks such a centre (moore 2017), and identifying the mediums through which the previous identified topics (nature and heritage) should be displayed will help with the design of such a centre. when considering the preferred mediums, it is evident that the visitors are in need of technological forms of interpretation (audio and visual), which are currently not on offer in the limited interpretation offerings (mearns & botha 2017; moore 2017). it is also evident from the results that careful consideration should be given to the delivery of information. as ham (1992) explains, interpretation should be enjoyable, personally relevant and organised, and it is therefore not surprising that visitors prefer information that is creative, fun (i.e. game-like), non-technical and concise. considering that visitors tend to spend more time within the park than at rest camps, it could be worthwhile to incorporate interpretation through mobile devices that tourists can access during their self-drives. an interactive map could highlight points of interest related to either nature or heritage, where on-site interpretation (at the points of interest) could be useful for additional information (such as through the use of qr codes as explained earlier). conclusion the aim of this research was to determine the interpretation needs and preferences of visitors to ktp, and to address this aim, three questions were raised: what are the interpretation service preferences of tourists visiting this national park? what are the topics or types of information visitors are interested in? do visitors prefer specific mediums through which the topics and information should be presented? the answers to these questions revealed several practical implications for the design of interpretation services in ktp based on tourists’ needs and preferences rather than park management’s needs and preferences. customising interpretation services to the tourists’ needs and preferences may add to the visitors’ enjoyment, create loyalty, extend stays, and increase expenditure and revenue for parks (mearns & botha 2017). interpretation is therefore no longer seen as a ‘luxury’ service but is rather viewed worldwide as an essential management function of national parks, and this is also the case in ktp. the research makes the following contributions to the field of interpretation in ecotourism and natural area tourism. first, this study was the first of its kind conducted in ktp regarding interpretation. second, the study revealed the preferred interpretation services, topics, extent of information and media for interpretation, which will assist management with interpretation development and management. as this research focused on the general needs and preferences of tourists for interpretation services, context (i.e. topics) and media, it would be worthwhile to research effective approaches to deliver specific themes (i.e. messages about the topics). it would also be valuable to determine whether there are specific thresholds of information consumption (extent of information) per specific medium of interpretation to determine the most suitable medium for each topic. acknowledgements the authors acknowledge south african national parks for the opportunity to conduct this research and for its support. competing interests the authors declare that they have no conflicts of interest. authors’ contributions all authors contributed to the study conception and design. material preparation and data collection were performed by p.v.d.m., m.s. and e.b. the literature was written by e.b.; the methodology, results and findings were written by p.v.d.m. all authors had the opportunity to comment on one another’s versions, and all read and approved the final manuscript. funding information this research was funded by south african national parks. data availability statement the raw data of this study will not be made available to external third parties, as this was not applied for during the ethics clearance process, and consequently, the respondents have not provided consent to do so. disclaimer the views expressed in this article are those of the authors and are not an official position of the institution or funder. references ballantyne, r., hughes, k. & bond, n., 2016, ‘using a delphi approach to identify managers’ preferences for visitor interpretation at canterbury cathedral world heritage site’, tourism management 54(june), 72–80. https://doi.org/10.1016/j.tourman.2015.10.014 ballantyne, r., packer, j. & falk, j., 2011, ‘visitors’ learning for environmental sustainability: testing shortand long-term impacts of wildlife tourism experiences using structural equation modelling’, tourism management 32(6), 1243–1252. https://doi.org/10.1016/j.tourman.2010.11.003 ballantyne, r., packer, j. & hughes, k., 2009, ‘tourists’ support for conservation messages and sustainable management practices in wildlife tourism experiences’ tourism management 30(5), 658–664. https://doi.org/10.1016/j.tourman.2008.11.003 ballantyne, r., packer, j. & sutherland, l.a., 2011, ‘visitors’ memories of wildlife tourism: implications for the design of powerful interpretive experiences’, tourism management 32(4), 770–779. https://doi.org/10.1016/j.tourman.2010.06.012 botha, e., saayman, m. & kruger, m., 2016, ‘expectations versus experience – the kruger national park’s interpretation services from a regional approach’, journal of ecotourism 15(2), 158–183. https://doi.org/10.1080/14724049.2016.1178753 bidder, c., kibat, s.a. & fatt, b.s., 2016, ‘cultural interpretation toward sustainability: a case of mount kinabalu’, procedia: social and behavioural sciences 224(15), 632–639. https://doi.org/10.1016/j.sbspro.2016.05.454 carmody, j. & prideaux, b., 2011, ‘enhancing the role of host communities in the management of protected areas through effective two-way communications: a case study’, asia pacific journal of tourism research 16(1), 89–104. https://doi.org/10.1080/10941665.2011.539393 ceballos-lascurain, h., 1987, ‘the future of ecotourism’, mexico journal (january), 13–14. chen, w.y. & jim, c.y., 2012, ‘contingent valuation of ecotourism development in country parks in the urban shadow’, international journal of sustainable development & world ecology 19(1), 44–53. https://doi.org/10.1080/13504509.2011.588727 cheung, l.t.o. & jim, c.y., 2013, ‘ecotourism service preference and management in hong kong’, international journal of sustainable development & world ecology 21(2), 149–159. https://doi.org/10.1080/13504509.2013.859183 cheung, l.t.o. & jim, c.y., 2014, ‘expectations and willingness-to-pay for ecotourism service in hong kong’s conservation areas’, international journal of sustainable development & world ecology 20(2), 182–194. https://doi.org/10.1080/13504509.2013.775192 cohen, j.w., 1988, statistical power analysis for the behavioural sciences, 2nd edn., l. erlbaum associates, hillsdale, nj. de rojas, c. & camarero, c., 2008, ‘visitors’ experience, mood and satisfaction in a heritage context: evidence from an interpretation centre’, tourism management 29(3), 525–537. https://doi.org/10.1016/j.tourman.2007.06.004 de witt, l., van der merwe, p. & saayman, m., 2015, ‘key environmental management factors in protected areas: an eco-tourist perspective’, african journal of hospitality, tourism and leisure 4(2), 1–12. dioum, b., 1968, paper presented at the general assembly of the international union for the conservation of nature and natural resources, new delhi: seattle public library archive. du plessis, l., van der merwe, p. & saayman, m., 2013, ‘tourist perception on whether south african national parks are environmentally friendly’, acta academica 44(3), 52–76. eagles, p.f.j., 2014, ‘research priorities in park tourism’, journal of sustainable tourism 22(4), 528–549. https://doi.org/10.1080/09669582.2013.785554 fennell, d., 2003, ecotourism: an introduction, 4th edn., sage, london. field, a., 2013, discovering statistics using spss, 2nd edn., sage, new york, ny. geldenhuys, s., 2009, ‘ecotourism criteria and context’, in m. saayman (ed.), ecotourism: getting back to basics, pp. 1–24, institute for tourism and leisure studies, potchefstroom. haely, n., van riper, c.j. & boyd, s.w., 2016, ‘low versus high intensity approaches to interpretive tourism planning: the case of the cliffs of moher, ireland’, tourism management 52(february), 574–583. https://doi.org/10.1016/j.tourman.2015.08.009 ham, s., 1992, environmental interpretation: a practical guide for people with big ideas and small budgets, north american press, golden, co. ham, s., housego, a. & weiler, b., 2005, tasmanian thematic interpretation planning manual, tourism tasmania, hobart. ham, s.h. & weiler, b., 2006, development of a research-based tool for evaluating interpretation, crc for sustainable tourism, gold coast. henker, k.b. & brown, g., 2011, ‘as good as the real thing? a comparative study of interpretive podcasts and traditional ranger talks’, journal of interpretation research 16(1), 7–23. hermann, u.p., van der merwe, p., coetzee, w.j.l. & saayman, m., 2015, ‘a visitor perspective of conservation management at a south african national park and world heritage site’, african journal of hospitality tourism and leisure 4(1), 1–11. jacobs, m.h. & harms, m., 2014, ‘influence of interpretation on conservation intentions of whale tourists’, tourism management 42(june), 123–131. https://doi.org/10.1016/j.tourman.2013.11.009 jurdana, d.s., 2009, ‘specific knowledge for managing ecotourism destinations’, tourism hospitality management 15(2), 267–278. kaiser, h., 1974, ‘an index of factorial simplicity’, psychometrika 39(1), 31–36. https://doi.org/10.1007/bf02291575 kang, m. & gretzel, u., 2012, ‘effects of podcast tours on tourist experiences in a national park’, tourism management 33(2), 440–455. https://doi.org/10.1016/j.tourman.2011.05.005 keyton, j. & beck, s.t. 2010, ‘perspective: examining communication as macrocognition in sts’, human factors 52(2), 335–339. https://doi.org/10.1177/0018720810371338 kolb, d.a., 1984, experiential learning: experience as the source of learning and development, prentice-hall, englewood cliffs, nj. krejcie, r.v. & morgan, d.w. 1970, ‘determining sample size for research activities’, educational and psychological measurement 30(3), 607–610. https://doi.org/10.1177/001316447003000308 kuo, i., 2002, ‘the effectiveness of environmental interpretation at resource-sensitive tourism destinations’, international journal of tourism research 4(2), 87–101. https://doi.org/10.1002/jtr.362 langer, e.j., 1989, ‘minding matters: the consequences of mindlessness-mindfulness’, advances in experimental social psychology 22, 137–173. https://doi.org/10.1016/s0065-2601(08)60307-x lee, t.h., 2009, ‘a structural model for examining how destination image and interpretation services affect future visitation behaviour: a case study of taiwan’s taomi eco-village’, journal of sustainable tourism 17(6), 727–745. https://doi.org/10.1080/09669580902999204 lück, m., 2015, ‘education on marine mammal tours – but what do tourists want to learn?’, ocean & coastal management 103(january), 25–33. https://doi.org/10.1016/j.ocecoaman.2014.11.002 maciejewski, k. & kerley, g.i.h., 2014, ‘understanding tourists’ preference for mammal species in private protected areas: is there a case for extralimital species for ecotourism?’, plos one 9(2), 1–8, viewed 30 april 2020, from https://journals.plos.org/plosone/article/file?id=10.1371/journal.pone.0088192&type=printable. marschall, s., granquist, s.m. & burns, g.l., 2017, ‘interpretation in wildlife tourism: assessing the effectiveness of signage on visitor behaviour at a seal watching site in iceland’, journal of outdoor recreation and tourism 17(march), 11–19. https://doi.org/10.1016/j.jort.2016.11.001 mearns, k. & botha, e., 2017, ‘the roles of interpretation in the management of national parks in south africa’, in m.z. suratman (ed.), management and conservation, intechopen, viewed 30 april 2020, from https://www.intechopen.com/books/national-parks-management-and-conservation/the-roles-of-interpretation-in-the-management-of-national-parks-in-south-africa. mitsche, n., reino, s., knox, d. & bauernfeind, u., 2008, ‘enhancing cultural tourism e-services through heritage interpretation’, in p. o’connor, w. höpken & u. gretzel (eds.), information and communication technologies in tourism 2008: proceedings of the international conference in innsbruck, austria 2008, pp. 419–429, wien, springer, new york, ny. moore, k., 2017, interpretation needs for kgalagadi transfrontier park [personal interview], 17 jan 2017, kgalagadi transfrontier park. moscardo, g., 2014, ‘interpretation and tourism: holy grail or emperor’s robes?’, international journal of culture, tourism and hospitality research 8(4), 462–476. https://doi.org/10.1108/ijcthr-08-2014-0071 moscardo, g. & ballantyne, r., 2008, ‘interpretation and attractions’, in a. fyall, b. garrod, a. leask & s. wanhill (eds.), managing visitor attractions, 2nd edn., pp. 237–252, elsevier, oxford. orams, m., 1994, ‘creating effective interpretation for managing interaction between tourists and wildlife’, australian journal of environmental education 10(1), 21–34. https://doi.org/10.1017/s0814062600003062 orams, m., 1996, ‘a conceptual model of tourist-wildlife interaction: the case for education as a management strategy’, the australian geographer 27(1), 39–51. https://doi.org/10.1080/00049189608703156 quinlan, c., babin, b., carr, j., griffin, m. & zikmund, w., 2019, business research methods, 2nd edn., cencage, united kingdom. reisinger, y. & steiner, c., 2006, ‘reconceptualising interpretation: the role of tour guides in authentic tourism’, current issues in tourism 9(6), 481–498. https://doi.org/10.2167/cit280.0 saayman, m. & van der merwe, p., 2017, ‘identifying ecotourist accommodation needs and preferences’, the south african journal of business management 48(1), 67–75. https://doi.org/10.4102/sajbm.v48i1.21 sanparks (south african national parks), 2018, annual report 2017/2018, viewed 28 april 2020, from https://www.sanparks.org/assets/docs/general/annual-report-2018.pdf. sanparks (south african national parks), 2020a, kgalagadi transfrontier park: natural & cultural history, viewed 28 april 2020, from https://www.sanparks.org/parks/kgalagadi/tourism/history.php. sanparks (south african national parks), 2020b, kgalagadi transfrontier park, viewed 30 april 2020, from https://www.sanparks.org/parks/kgalagadi/. schiffman, l.g. & kanuk, l.l., 2007, consumer behaviour, 8th edn., prentice hall, upper saddle river, nj. schindler, a., ruoppolo, g. & barillari, u., 2010, ‘communication and its disorders: definition and taxonomy from a phoniatric perspective’, audiological medicine 8(4), 163–170. https://doi.org/10.3109/1651386x.2010.530023 sheena, b., mariapan, m. & aziz, a., 2015, ‘characteristics of malaysian ecotourist segments in kinabalu park, sabah’, tourism geographies 17(1), 1–18. https://doi.org/10.1080/14616688.2013.865069 south africa, 2003, protected areas act 57 south africa (2003), author, cape town, south africa. south africa (department of environmental affairs), 2019, transfrontier conservation areas, viewed 28 april 2020, from https://www.environment.gov.za/projectsprogrammes/transfrontier_conservation_areas#kgalakgadi. south africa (international relations and cooperation), 2004, transfrontier conservation areas (tfcas), viewed 15 july 2019, from http://www.dirco.gov.za/foreign/multilateral/inter/tfcas.htm. south african government, 1976, national parks act 57 south africa (1976), author, cape town, south africa. stewart, e.j., hayward, b.m., devlin, p.j. & kirby, v.g., 1998, ‘the ‘place’ of interpretation: a new approach to the evaluation of interpretation’, tourism management 19(3), 257–266. https://doi.org/10.1016/s0261-5177(98)00015-6 tilden, f., 1977, interpreting our heritage, 3rd edn., university of north carolina press, chapel hill, nc. united nations educational, scientific and cultural organisation (unesco), 2019a, ǂkhomani cultural landscape, viewed 15 july 2019, from http://whc.unesco.org/en/list/1545. united nations educational, scientific and cultural organisation (unesco), 2019b, state parties: south africa, viewed 15 july 2019, from http://whc.unesco.org/en/statesparties/za/. van der merwe, p. & saayman, m., 2008, ‘travel motivation of visitors to the kruger national park’, koedoe 50(1), 154–159. https://doi.org/10.4102/koedoe.v50i1.140 van der merwe, p., saayman, m. & pienaar, j., 2009, ‘expenditure-based segmentation of tourists to the kruger national park’, acta academica 41(3), 107–127. ward, c.w. & wilkinson, a.e., 2006, conducting meaningful interpretation: a field guide for success, fulcrum publishing, golden, co. zeppel, h. & muloin, s., 2008, ‘conservation benefits of interpretation on marine wildlife tours’, human dimensions of wildlife 13(4), 280–294. https://doi.org/10.1080/10871200802187105 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 filelist convert a pdf file! page 1 page 2 page 3 page 4 abstract introduction materials and methods results discussion conclusion acknowledgements references about the author(s) zishan ebrahim cape research centre, south african national parks, tokai, cape town, south africa centre for invasion biology, stellenbosch university, stellenbosch, south africa atherton de villiers scientific services, capenature, jonkershoek, stellenbosch, south africa john measey centre for invasion biology, stellenbosch university, stellenbosch, south africa citation ebrahim, z., de villiers, a. & measey, j., 2020, ‘assessing water conditions for heleophryne rosei tadpoles and the conservation relevance’, koedoe 62(1), a1581. https://doi.org/10.4102/koedoe.v62i1.1581 note: additional supporting information may be found in the online version of this article as online appendix 1 short communication assessing water conditions for heleophryne rosei tadpoles and the conservation relevance zishan ebrahim, atherton de villiers, john measey received: 06 aug. 2019; accepted: 18 mar. 2020; published: 11 aug. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the table mountain ghost frog (heleophryne rosei) is endemic to the table mountain massif and is critically endangered. other than clear, clean perennial stream flow, the optimal aquatic conditions required by their larvae are unknown. dissolved oxygen, temperature, ph, electro-conductivity, aspect and permanence of flow are the independent variables measured seasonally at two sampling altitudes at 12 rivers of the massif. using a logistic regression model we found that a permanence of water flow and lower water temperature were significant predictors of tadpole presence. streams with mean summer temperature above 17.2 °c, at 300 m – 400 m above sea level, do not have tadpoles. summer and autumn abstraction should be avoided, while a summer water temperature above an average of 17.2 °c is a threshold of potential concern for management authorities responsible for biodiversity conservation, threat mitigation efforts, and bulk-water supply and abstraction. conservation implications: the environmental water reserve has not been determined for streams of table mountain. the requirements of the critically endangered table mountain ghost frog (heleophryne rosei) can be adopted as the minimum conditions to support this species and associated communities. perennial flow, an average january water temperature of 17.2 °c or lower. keywords: amphibian; environmental water reserve; management; protected areas; temperature; water flow. introduction southern africa has a rich assemblage of anuran amphibians (south africa has 12 families). of these, one entire family is endemic to the southern african region (poynton 1964): the ghost frog family, heleophrynidae. these torrent-adapted species live in and around fast-flowing montane streams. adults are cryptic, hiding in crevices and emerging at night, while tadpoles are relatively easy to find in streams and easily identified by their large, specially adapted oral suckers used for grazing on algae-covered rocks (boycott 2004). the two genera of the ghost frog family contain seven species, one in the genus hadromophryne (natal cascade frog) and six in the genus heleophryne (ghost frogs); the latter is confined to the cape fold mountains (channing, boycott & van hensbergen 1988) across south-west south africa’s winter rainfall region (colville et al. 2014). the table mountain ghost frog, heleophryne rosei, is confined to the table mountain massif (boycott & de villiers 1986). the extreme endemism of h. rosei (range < 10 km2), and a set of ongoing threats including invasive vegetation and habitat degradation (boycott & de villiers 1986; ed. measey 2011), have meant that it has long been considered critically endangered (south african frog re-assessment group [sa-frog]; iucn 2017). ironically, it occurs only within areas protected by the south african national parks (sanparks) and the south african national biodiversity institute (sanbi). of the 12 streams draining off table mountain, only seven (counting upper disa and lower disa as two streams) are currently inhabited by tadpoles, but the species’ past distribution has been larger (boycott & de villiers 1986). table mountain has undergone many transformations, with the plateau holding several large dams; it was used as an alien timber plantation and subsequently cleared. the slopes are currently being cleared piecemeal of invasive acacias, pines and eucalypts (amongst others) (cheney et al. 2019). changes in flow regime and abstraction in the lower reaches of the mountain’s rivers may have led to the loss of h. rosei tadpole habitat, but we are still largely ignorant of the physiological requirements of all ghost frog species. the variables ph, salinity and water temperature are known to influence stream-breeding anurans with respect to metabolism, growth rates, emergence, fecundity and ultimately survival (rivers-moore & karssing 2014). tadpole health may reflect total population health, as well as ecosystem resilience (mcdiarmid & altig 1999). it is assumed that the larval stage of h. rosei lasts more than 12 months. this is based on congeneric assumptions, their observed occurrence only in sections of streams that are perennial (boycott & de villiers 1986), and because two size cohorts are observed after the breeding and spawning season (ze, adv & jm pers. obs.). the aim of this study is to determine which of these water variables influence the presence or absence of h. rosei tadpoles. in addition, we discuss known threats to this species in relation to the findings of the study in order to infer conservation implications and management directions. materials and methods site description the table mountain massif is a sandstone ridge with granite outcrops, constituting the northern section of south africa’s table mountain national park. twelve streams flowing off the mountain were chosen for this study (figure 1). afromontane forests are the natural riverine vegetation with adjacent sandstone and granite fynbos (mucina & rutherford 2006). the fynbos is fire-driven (van wilgen et al. 2010). forests exclude small fires while anthropogenically altered landscapes alter fire return intervals (slingsby et al. 2020). capenature, the nature conservation authority for the western cape province, monitors h. rosei tadpole populations within the table mountain national park and kirstenbosch national botanical garden estate at the end of the southern summer (ed. measey 2011; measey et al. 2019), before the wet season. this allows for the greatest tadpole visibility during the lowest flow. this monitoring is through an annual timed (30 min) count on five streams at altitudes of about 300 m – 400 m, and two streams at about 800 m. the former, lower-altitudinal monitoring provides the presence/absence response variable for this study. sanparks, the conservation authority for national parks in south africa, measures (in situ, seasonally) the water chemistry of selected streams within table mountain national park in order to compare conditions where tadpoles are present/absent, controlled for altitudes of 300 m and 400 m above sea level. figure 1: findings of this study are extrapolated for each stream between the lower limit of the species and the massif’s plateau. water temperature and water flow results are mapped as ‘dry’ to represent ‘drought’ and ‘warm’ to represent ‘temperature extremes’, interpolated for 400 m and above (to the plateau at 700 m), and for 300 m and below (to 200 m). green areas are favourable for tadpoles as water conditions are both below a summer mean temperature of 17.2 °c and water flows during all four seasons. the red bands indicate where water conditions are either above a summer mean temperature of 17.2 °c or water stops flowing during at least one season, or both, i.e. unfavourable for tadpoles. the yellow star represents adult cave habitats (gow 1963; poynton 1964). five dams on table mountain act as water reservoirs, (1) woodhead, (2) hely-hutchinson, (3) de villiers, (4) victoria, and (5) alexandra. while two reservoirs, (6) molteno and (7) kirstenbosch, are situated away from rivers. contour-lines and river-lines are sourced from the national geo-spatial institute of the department of land affairs and rural development. catchments were calculated by the council for scientific and industrial research (csir). water chemistry monitoring habitat suitability for aquatic larvae is entirely dependent on water conditions. the variables measured in this study are a global standard for monitoring, and prescribed by sanparks’ biodiversity monitoring system: the freshwater and estuarine monitoring programme (russell, roux & randall 2012). a hand-held, multi-probe, water chemistry instrument (models ph100, do200, ec300; ysi, yellow springs instrument, yellow spring, united states) was used. monitoring began in the summer of 2014 (inclusive of the el niño event), ending in autumn 2016. measuring these variables in situ is cost-effective and convenient. measurement was taken in the middle of a stream, mid-profile, after at least 10 min of stabilisation. the explanatory variables assessed in this study are: aspect, electro-conductivity (salinity), dissolved oxygen, ph, water temperature, and permanence of water flow. although this is not a comprehensive water analysis, these integrated metrics provide an overall indication of habitat health. statistical analyses a stepwise linear regression was used to remove variables that do not explain observed presence/absence. linear models are constructed using the retained explanatory variables. the binomial response variable (presence or absence) was tested for, while abundances are displayed in relation to the ecological boundary for the h. rosei tadpoles (table 1). the log-likelihood of 15 models was derived, and akaike’s information criterion calculated (akaike 1973). all analyses were conducted in r (r core team 2019). table 1: four variables were considered for linear models. results tadpoles are found in water with an annual mean temperature from 13.4 °c to 14.1 °c (10 consecutive seasons), rising to a summer mean temperature of 16.2 °c – 17.2 °c (mean of three consecutive summer seasons). three variables best predict the presence of h. rosei tadpoles below the table mountain plateau: the three interacting effects of a stream’s permanence of flow, mean water temperature, and electro-conductivity (table 1). the two best models have a difference in akaike’s information criterion of less than two, thus the model with the lowest number of parameters is preferred (anderson & burnham 2002). the lower the water temperature, the more likely it is that tadpoles are present. permanent water flow is essential for tadpoles. the discerning effect of electro-conductivity is least significant, yet positive, indicating that higher conductivity is preferred, but lower conductivity still tolerated, by tadpoles. with this regression model, over 60% of the variance in the dataset is explained by these three variables (r2 = 0.602). only permanence of flow and mean temperature are significant contributors to this and to the state of the dependent variable (p < 0.001). four of the five extant streams had constantly flowing surface water at both measured altitudes during all four seasons (table 2, where n = 10). nursery ravine, much like cecelia stream, had no surface flow for large sections below the 400 m contour during spring and summer. table 2: the mean summer and mean annual temperatures (2014–2016), indicating permanence of flow (n = 3 and n = 10 respectively). discussion results suggest that permanence of water flow, and low water temperature, are the best habitat predictors for the presence of h. rosei tadpoles; more so than electro-conductivity, aspect, and the other variables measured. the data (for summer) suggest that an upper bound of a putative temperature envelope for h. rosei tadpoles is 17.2 °c. the lower end of the temperature envelope cannot be measured in situ as it is not likely to be reached naturally. permanence of water flow in table mountain rivers water yield and water temperature are known to be influenced by land-use practices (bosch & hewlett 1982) that affect adjacent riparian vegetation cover and fluvial connectivity, and they are also affected by water abstraction and water storage (baxter 1977; cooper et al. 2013; davies & day 1998). both no-flow of water and water of too high a temperature are indicative of habitat-disconnect. nursery ravine, cecelia ravine and fernwood gully each have only one of two altitudes with favourable conditions for tadpoles. of these, nursery ravine is the only stream that supports tadpoles in the 21st century. much like cecelia ravine, adverse conditions are most likely due to lack of water flow rather than higher temperatures. dams and water management dams transform and flood lotic stream habitats, and divide continuous rivers. regulation may allow for cooler summer water temperatures below the dam wall compared to if the dam were absent (baxter 1977), while supplementing summer flow when water is released unseasonably, but this may cause other problems if there are regular releases of high water volumes over a short time. the waters of disa stream flow permanently even though water is diverted from above the study sites westward through the twelve apostles tunnel. water that would have naturally flowed east, to reach cecelia (and spilhaus, rooikat) ravine, is dammed on the plateau (victoria and alexandra dams) and directed south-westwards via the de villiers dam. the alexandra dam’s excess waters could best be utilised for a restorative ecological experiment: returning water volumes to the cecelia/rooikat streams (see the bypass between dam no. 5 and dam no. 3 in figure 1). the kirstenbosch reservoir stores water abstracted from the weirs on window gorge and nursery ravine, which siphon off much of the summer flow, such that there is reduced tadpole habitat below both weirs (adv & ze pers. obs.). water abstractions should be limited to below the habitat of this species. pine plantations the table mountain plateau was initially felled of plantations in the 1970s and 1980s, in a phased harvest (van wilgen 1996). table mountain ghost frogs still breed there today, even though pines are still present at low densities. however, it is unknown whether tadpole numbers have improved since then, as capenature’s systematic h. rosei tadpole monitoring programme only started in 2003. table mountain national park is removing pine plantations from the tokai and cecelia forests in a piecemeal fashion. an increase in water yield would be a positive sign of habitat rehabilitation (bosch & hewlett 1982). fire, erosion, and siltation threat fires do not occur frequently through h. rosei stream habitats, even though frequent fire is listed as a potential threat to h. rosei (south african frog re-assessment group [sa-frog], iucn 2017). fire is naturally excluded by afromontane forest vegetation, but frequent fires do occur at the urban edge of table mountain national park. much of the fynbos covered back (south) table and table mountain plateau has not burned for over six decades (pooley 2014). table mountain national park attempts to mimic natural fire cycles through ecologically prescribed piecemeal burning of senescent vegetation (slingsby et al. 2020). siltation has been regarded as a threat to frogs of the family heleophrynidae (de villiers 2004; minter 2004). one stream with a seemingly consistent anthropogenic sand-input is skeleton gorge. this is because of a dune that formed from weathered sandstone off the bottom of the hely-hutchinson reservoir. dune stabilisation is the only viable mitigation. erosion and siltation monitoring should include fixed-point photography to determine rates of change and whether improvements are being made. temperature re-colonisation of h. rosei tadpoles in the streams of cecelia ravine would represent an indication for successful rehabilitation of that system, but this has not yet occurred. the average summer temperature for the cecelia/spilhaus stream is 17.0 °c; however, the summer of 2015 had an average of 17.4 °c. presumably this stream was cooler in the past if greater volumes of water flowed through it then. factors influencing water temperature are flow rate and the presence of dams. both tributaries of the disa river, disa stream (in disa gorge) and original disa stream, flow from dams. the precautionary approach should apply in this regard: release of cooler (hypolimnion) waters is preferred, rather than the over-flow of warmer surface waters (from the epilimnion). the mean water temperature of mountain streams is expected to increase, largely due to increased air temperature attributed to climate change (isaak et al. 2012). riparian vegetation has a cooling effect on the water in the adjoining stream (davies & day 1998). the table mountain ghost frog is associated with afromontane forests. these forests have shrunk from their pre-colonial extents (sim 1907), but have rebounded on the southern slopes of table mountain (poulsen & hoffman 2015). the latter authors suggest that in the past the indigenous forest canopy was significantly greater than its current extent, and the rebound is limited to the southern slopes of table mountain. campbell and moll (1977) show that the platteklip and kasteelspoort (valley of the red gods) streams flowed under the cover of indigenous trees (circa 1600s) for much of their length. mitigation of high temperatures could include the arrest of energy (heat) inputs through the restoration of indigenous forest canopies for cooling and shade. conclusion the extent of expansion of the table mountain ghost frog’s reproductive metapopulations may be predicted based on suitable tadpole habitat. the environmental reserve (of water) is a legal obligation, but it has not been determined for rivers in the study area. the flow requirements of a critically endangered freshwater species may be useful as a proxy for the environmental reserve. environmental water (flow) requirements may not be met (for h. rosei tadpoles) in cecelia ravine, possibly due to pine plantations and a water bypass between the alexandra and devilliers dams (figure 1). nursery ravine is the extant stream in which the observed presence of h. rosei tadpoles is lowest (ze & adv pers. obs.). cecelia and streams with abstractive weirs should be prioritised for intervention. recommendations in addition to existing measures (e.g. alien clearing), two actions to mitigate temperature extremes and droughts could be considered: (1) streams can be shaded by rehabilitating natural vegetation (through a properly managed burn cycle, not manual planting), and (2) the environmental water reserve should be determined, met and/or mitigated, by supplementing water inputs (to the upper catchment) where possible, and by limiting water abstractions to an altitude below that used by h. rosei tadpoles. there is no forum currently to debate solutions. the city of cape town, capenature, kirstenbosch botanical gardens, and sanparks (and perhaps the national and provincial authorities that issue water-use licenses) need to be open to options and innovations regarding water resilience, research and monitoring. a cape peninsula freshwater conservation forum would be helpful to co-ordinate conservation and freshwater management, not only for human needs but for freshwater biodiversity and ecosystem functionality as well. capenature, south african national biodiversity institute and table mountain national park should co-ordinate non-invasive tadpole surveys and water chemistry monitoring more than once a year; perhaps on consecutive days, and weeks apart. the mimicry of natural fire-return cycles in an appropriate, controlled manor is needed to rehabilitate the upper catchment area of the table mountain plateau. supplementation of water as a mitigation measure is conceivable for cecelia ravine if some waters of the victoria and alexandra dams are not diverted to the de villiers dam, but rather left on their original eastward course towards cecelia and rooikat ravines. the cecelia/rooikat streams, nursery stream, and the front and back table (north and south plateau) are the sources of four of cape town’s rivers. these four streams represent four approaches to water course rehabilitation (in addition to alien clearing): reservoir-overflow diversion for environmental water requirements (cecelia/rooikat is the source of the zand river catchment), mitigation of water-abstraction from small catchments (nursery is the source of the liesbeek river catchment), indigenous shading-plant rehabilitation (platteklip is the source of the pre-colonial – now subterranean – camissa river), and ecological burns (the back table is the source of the disa river). acknowledgements competing interests the authors declare that they have no competing interests. authors’ contributions z.e. and j.m. conceptualised the article and attended to data analysis. z.e. and a.d.v. were responsible for data collection. z.e., a.d.v. and j.m. wrote the article. funding information funding for this article were received from a.w. mellon foundation, department of science and innovation-national research fund (dsi-nrf), centre of excellence for invasion biology at stellenbosch university, south african national parks and capenature. data availability statement the data is available in the sanparks data repository, cc by 3.0. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors references akaike, h., 1973, ‘maximum likelihood identification of gaussian autoregressive moving average models’, biometrika 60(2), 255–265. https://doi.org/10.1093/biomet/60.2.255 anderson, d.r. & burnham, k.p., 2002, ‘avoiding pitfalls when using information-theoretic methods’, the journal of wildlife management 66(3), 912–918. https://doi.org/10.2307/3803155 baxter, r.m., 1977, ‘environmental effects of dams and impoundments’, annual review of ecology and systematics 8, 255–283. https://doi.org/10.1146/annurev.es.08.110177.001351 bosch, j.m. & hewlett, j.d., 1982, ‘a review of catchment experiments to determine the effect of vegetation changes on water yield and evapotranspiration’, journal of hydrology 55(1–4), 3–23. https://doi.org/10.1016/0022-1694(82)90117-2 boycott, r.c., 2004, ‘genus heleophryne sclater, 1898 (family heleophrynidae)’, in minter, l.r., burger, m., harrison, j.a., braack, h.h., bishop, p.j. & knoepfer, d., atlas and red data book of frogs of south africa, pp. 95–97. si/mab series no.9, smithsonian institution press, washington dc boycott, r.c. & de villiers, a.l., 1986, ‘the status of heleophryne rosei hewitt (anura: leptodactylidae) on table mountain and recommendations for its conservation’, south african journal of wildlife research 16(4), 129–134. campbell, b.m. & moll, e.j., 1977, ‘the forest communities of table mountain, south africa’, vegetatio 34(2), 105–115. https://doi.org/10.1007/bf00054478 channing, a., boycott, r. & van hensbergen, h.j., 1988, ‘morphological variation of heleophryne tadpoles from the cape province, south africa (anura: heleophrynidae)’, journal of zoology 215(2), 205–216. https://doi.org/10.1111/j.1469-7998.1988.tb04895.x cheney, c., esler, k. j., foxcroft, l. c., & van wilgen, n. j. 2019, ‘scenarios for the management of invasive acacia species in a protected area: implications of clearing efficacy’, journal of environmental management 238, 274–282. https://doi.org/10.1016/j.jenvman.2019.02.112 colville, j.f., potts, a.j., bradshaw, p.l., measey, g.j., snijman, d., picker, m.d. et al., 2014, ‘floristic and faunal cape biochoria: do they exist?’, in n. allsopp, j.f. colville, g.a. verboom (eds.), fynbos: ecology, evolution and conservation of a megadiverse region, pp. 73–93, oxford university press, usa. cooper, s. d., lake, p. s., sabater, s., melack, j. m., & sabo, j. l., 2013, ‘the effects of land use changes on streams and rivers in mediterranean climates’, hydrobiologia 719(1), 383–425. https://doi.org/10.1007/s10750-012-1333-4 davies, b. & day, j., 1998, vanishing waters, university of cape town press, cape town. de villiers, a., 2004, ‘species account: heleophryne rosei (hewitt, 1925)’, in minter, l.r., burger, m., harrison, j.a., braack, h.h., bishop, p.j. & knoepfer, d., atlas and red data book of the frogs of south africa, lesotho and swaziland, pp. 108–111. si/mab series no.9, smithsonian institution press, washington dc. gow, c.e., 1963, ‘notes on the habitat of heleophyrne rosei’, african wildlife 17, 113–116. isaak, d. j., wollrab, s., horan, d., & chandler, g., 2012, ‘climate change effects on stream and river temperatures across the northwest us from 1980–2009 and implications for salmonid fishes’, climatic change 113(2), 499–524. https://doi.org/10.1007/s10584-011-0326-z iucn ssc amphibian specialist group, south african frog re-assessment group (sa-frog), 2017. heleophryne rosei. the iucn red list of threatened species 2017: e.t9773a13015448, veiwed 14 october 2017 from https://doi.org/10.2305/iucn.uk.2017-2.rlts.t9773a77164671.en. mcdiarmid, r.w. & altig, r., 1999, tadpoles: the biology of anuran larvae, university of chicago press. chicago. measey, g.j. (ed.), 2011, ensuring a future for south africa’s frogs: a strategy for conservation research. sanbi biodiversity series 19. south african national biodiversity institute, pretoria. measey, j., tarrant, j., rebelo, a., turner, a., du preez, l., mokhatla, m. et al., 2019, ‘has strategic planning made a difference to amphibian conservation research in south africa?’, bothalia 49(1), 1–13. https://doi.org/10.4102/abc.v49i1.2428 minter, l.r., burger m., harrison j.a., braack h.h., bishop p.j. & kloepfer d. (eds.), 2004, atlas and red data book of the frogs of south africa, lesotho, and swaziland. si/mab series no.9, smithsonian institution press, washington dc. mucina, l. & rutherford, m.c. (ed.), 2006, the vegetation of south africa, lesotho and swaziland. strelitzia 19. south african national biodiversity institute, pretoria. pooley, s., 2014, burning table mountain: an environmental history of fire on the cape peninsula. palgrave macmillan, london. poulsen, z.c. & hoffman, m.t., 2015, ‘changes in the distribution of indigenous forest in table mountain national park during the 20th century’, south african journal of botany 101, 49–56. https://doi.org/10.1016/j.sajb.2015.05.002 poynton, j.c., 1964, amphibia of southern africa; a faunal study. annals of the natal museum 17. natal museum, pietermaritzburg. r core team, 2019, r: a language and environment for statistical computing, r foundation for statistical computing, vienna. rivers-moore, n.a. & karssing, r.j., 2014, ‘water temperature affects life-cycle duration of tadpoles of natal cascade frog’. african journal of aquatic science, 39(2), 223–227. russell, i., roux, d. & randall, r., 2012, ‘biodiversity monitoring programme: freshwater and estuarine ecosystems’, south african national parks, skukuza. sim, t.r., 1907, forests and forest flora of the colony of the cape of good hope. taylor & henderson, aberdeen. slingsby, j. a., moncrieff, g. r., rogers, a. j., & february, e. c., 2020, ‘altered ignition catchments threaten a hyperdiverse fire-dependent ecosystem’, global change biology, 26(2), 616–628. van wilgen, b.w., 1996, ‘management of the natural ecosystems of the cape peninsula: current status and future prospects’, biodiversity & conservation 5(5), 671–684. https://doi.org/10.1007/bf00137614 van wilgen, b.w., forsyth, g. g., de klerk, h., das, s., khuluse, s., & schmitz, p., 2010, ‘fire management in mediterranean-climate shrublands: a case study from the cape fynbos, south africa’, journal of applied ecology 47(3), 631–638. https://doi.org/10.1111/j.1365-2664.2010.01800.x filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 abstract introduction research method and design results discussion conclusion acknowledgements references footnote about the author(s) hendri coetzee institutional sustainability and community impact office and unit for environmental sciences and management, north-west university, south africa werner nell optentia research programme, north-west university, south africa citation coetzee, h. & nell, w., 2019, ‘the feasibility of national parks in south africa endorsing a community development agenda: the case of mokala national park and two neighbouring rural communities’, koedoe 61(1), a1470. https://doi.org/10.4102/koedoe.v61i1.1470 original research the feasibility of national parks in south africa endorsing a community development agenda: the case of mokala national park and two neighbouring rural communities hendri coetzee, werner nell received: 04 apr. 2017; accepted: 19 sept. 2018; published: 28 feb. 2019 copyright: © 2019. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract this article explores the feasibility of south african national parks (sanparks) endorsing a community development agenda, using mokala national park (mnp) and two neighbouring rural communities as case study. a three-phase sequential exploratory, mixed-methods approach was followed: an initial exploratory qualitative phase aimed at identifying the development needs of the two communities; a quantitative phase aimed at verifying and quantifying the identified needs; and a final qualitative phase (with a minor quantitative component) to determine what parks can reasonably achieve in terms of community development based on their available resources, capacity and expertise. qualitative data were collected via semi-structured interviews (phase 1: n = 22; phase 3: n = 6), which were thematically analysed. quantitative data were collected via a structured questionnaire (phase 2: n = 484; phase 3: n = 6) and analysed using spss 23. findings revealed that the communities’ most significant needs centred on employment opportunities; improved healthcare, service delivery and waste management; and education. community members also expressed the need for improved community policing, safety and security; social services; agricultural support and training; general skills development and training; local leadership; recreational facilities; local economic development and conservation initiatives. results from the third phase of the study suggest that parks such as mnp can realistically only address some of the identified community needs significantly; primarily job creation (via temporary employment), skills development, local economic development, support of local conservation (especially via environmental education) and, to a lesser extent, agricultural support and training and permanent job creation. conservation implications: the findings could be of practical use to sanparks to steer its community development initiatives towards attaining a more optimal balance between actual community needs and what the organisation can realistically offer, thus rendering sanparks’ efforts more efficient and effective in supporting the establishment of equitable and sustainable rural communities. introduction national parks can potentially play an important role in rural community development. this sentiment is shared by researchers from all over the world, including africa (newmark & hough 2000; petursson & vedeld 2017), the americas (machlis & field 2000; nelson & serafin 2013), europe (barrow 2015; lundmark, fredman & sandell 2010), asia (kusters et al. 2006; tisdell 1999) and australia (stolton & dunley 2015). this notion is also supported by a recent report by the global environmental fund’s scientific and technical advisory panel (pullin 2014), which outlines the impacts that protected areas (e.g. national parks) can have on rural communities, especially with regard to environmental capital (e.g. resource use and access to land), economic and social capital, health and inequality. the idea of national parks adopting a developmental agenda stems from a paradigm shift away from traditional forms of conservation (preservation) to more contemporary forms that encompass both conservation and development objectives (anthony 2007). this led to what is widely known today as community-based conservation (cbc) (berkes 2007). the underlying principle of this combined approach is that poor rural communities should benefit directly from conservation (cock & fig 2000). several variations of the approach developed over the years, including (1) that communities should develop their own traditional or cultural land into conservation areas (iucn 2010; stevens 2014); (2) that communities should support protected areas and, in return, that they ought to benefit from job creation, skills development and other types of development in their communities (bennet 2014; pelser, redelinghuis & velelo 2013); (3) that communities should be allowed to harvest grass, wood, medicinal plants and other natural resources from parks (cock & fig 2000; sanparks 2016); and (4) the use of ‘trade-offs’, where certain areas (and the biodiversity contained therein) are protected whilst other parts of the park are made available for development (rights & resource initiative 2015). mokala national park (mnp), which serves as a case study in this article, is a good example of a trade-off approach being applied within a south african context, as it is basically a ‘trade-off park’1 in the place of the former vaalbos national park that was handed back to a local community as part of a land restitution case (sanparks 2017). elements of these variations to cbc can also be found in the latest version of sanparks’ 5-year strategic management plan (2015/2016–2019/2020) and, more specifically, in its strategic outcome-orientated goal 3, in terms of which sanparks’ goal is (sanparks 2016): [t]o foster an efficient, effective and development oriented public service and an empowered, fair and inclusive citizenship that will enable the creation of decent employment through inclusive economic growth, vibrant, equitable and sustainable rural communities. (p. 28) as outlined in the document, this plan is underpinned by the current national government’s ‘five key pillars’, which include job creation, rural development, education and health, as well as a reduction in the levels of crime in society. sanparks adopted a vision ‘connecting to society’ (sanparks 2012), created a social ecology unit (swemmer & taljaard 2011), expanded the scope of its people and conservation programme and added several programmes aimed at promoting access and benefit-sharing, socio-economic development and improved living conditions for local communities adjacent to national parks. this encompasses the implementation of national government’s expanded public works programmes (epwp) (e.g. working for water, land, wetlands, the coast, fire, etc.), value-adding industries, wildlife economy, the development of small, medium and micro-sized enterprises (smmes) and a number of social investment programmes at almost all of the national parks (sanparks 2012). it is not difficult to justify national parks in south africa endorsing a development agenda. from a conservation perspective, an interdependence between biodiversity and sustainable rural development is needed (faasen 2006), and much has been written about the threat of unsustainable land-use practices as well as the impact that resource-poor, hunger-stricken rural communities may have on biodiversity and wildlife (vira & kontoleon 2010). from a development perspective, this approach also makes sense as parks are scattered throughout south africa, often in deep rural areas where government cannot always address all the needs in these communities, which are often resource-poor and desperate for help and support from others (flora, flora & gasteyer 2015; mohan 2009). the benefits of a cbc approach, as well as the costs thereof and the challenges it poses, have been widely reported (botha, witkofski & cock 2007; murphee & hulme 2001; western, wright & strum 2013). most authors seem to concur that cbc, at best, has only achieved mixed results (berkes 2004; conley & moote 2003; mansuri & rao 2004). many reasons for this have been given (cox, arnold & villamayor-tomas 2010), which centre mainly around the fact that community development objectives and conservation objectives are often incompatible and that the benefits derived from community-conservation partnerships are often one-sided, primarily benefitting a conservation rather than a community development agenda (emerton 2001). one possible strategy to investigate and address these concerns further is to assess the actual needs of communities adjacent to national parks empirically (wells & mcshane 2004) in light of existing literature as well as on the basis of a qualitative exploration pertaining to national parks’ current and proposed community development strategies as well as their pool of available resources, skills et cetera. this would enable the feasibility of various development initiatives to be assessed in the light of empirical evidence about what specific communities identify as their particular needs, which, in turn, will allow for distinctions to be made in terms of the viability of specific strategies as outlined in existing sanparks documentation. furthermore, this approach could potentially address the one-sidedness of the relationship between parks and communities by ensuring that the development side of cbc initiatives articulates self-identified community needs. although sanparks has already adopted a developmental approach, given limitations such as restrictions on specific development-related skills and expertise (berkes 2007; biggs et al. 2014; pelser et al. 2013) that fall outside the typical skills required for park personnel, capacity and resources, the extent to which national parks can play a substantive role in community development needs to be investigated. to address these questions, a sequential exploratory, mixed-methods study was conducted in two rural communities (ritchie and ratanang) near mnp in order to explore what the typical developmental needs within communities situated near a national park are. this was followed by an additional qualitative study (with a minor quantitative component) aimed at exploring mnp’s available capacity, skills and expertise and current cbc initiatives. following this, the identified community needs were evaluated in the light of a theoretical and empirical investigation of sanparks’ cbc initiatives as well as the organisation’s capacity, resources and expertise with the intention to assess the feasibility of such needs being addressed by parks such as mnp. research method and design study area mokala national park is situated in a semi-arid part of central south africa near kimberley in the northern cape province (figure 1). ritchie, with a total estimated population size of 3504 (stats-sa 2011), is situated on the northern cape side of the border near modderrivier, an area that falls within the frances baard district municipality and sol plaatjies local municipality, whilst ratanang, with a total estimated population size of 4213 (stats-sa 2011), borders jacobsdal, which is located on the free state side of the border and falls within the xhariep district and letsemeng local municipality (see figure 1). the largest portion of the population in both areas classify themselves as mixed race and speak mostly afrikaans (stats-sa 2011). the two communities were selected as they are the communities closest to mokala. figure 1: map of the study area. design a mixed-methods approach, based on a pragmatic paradigm (creswell & plano clark 2007; johnson, onwuegbuze & turner 2007), was followed in the present study, which was executed in three sequential phases: an initial exploratory qualitative phase aimed at identifying the development needs in the two communities; a quantitative phase, where a structured survey was developed and administered to verify and quantify the needs that were identified during the first phase; and a final qualitative phase (with a minor quantitative component) to determine what parks can reasonably achieve based on their available resources, capacity and expertise. this type of approach is widely accepted as one of the most useful approaches to research, because it provides multiple perspectives on a given topic (creswell 2003) and also affords researchers the opportunity to further explore and explain quantitative results, which, according to tashakkori and teddlie (2003), will add richness, depth and greater credibility to any study. participants during the course of the study, three different groups of participants were recruited. a similar procedure was followed during the first (n = 22) and third (n = 6) qualitative phases of the study, where participants were purposively selected (creswell 2013) based on their knowledge and experience relevant to the research aims. this approach was supplemented with a snowball sampling strategy, where initially selected participants were requested to refer the researchers to other potential participants who met the sampling criteria (creswell 2013). the purposive sampling criteria that were set for the first phase required that participants had to be active participants in, or residents of, one of the two communities and had to occupy a central or leadership role in the community. the community leaders were identified via snowball sampling. the initial sample was comprised of two staff members from mnp, two staff members from sanparks, two ward councillors, a school principal, two local policemen, two nurses heading up community clinics and the two social workers appointed to work in these communities. an additional number of nine community members representing different stakeholder groups (e.g. youth, elderly, women, etc.) were recruited by means of typical instance sampling (tracy 2013). four hundred and eighty-four participants participated in the second, quantitative phase of the study. of these, 300 resided in ritchie (the larger of the two communities) and 184 in ratanang. a systematic sampling frame (creswell 2003) was used in order to ensure that all areas of each community were surveyed. using this approach, every nth household (determined by the total number of participants required for the study in relation to the number of households in each community) was surveyed in each community. the mean age of the participants was 35.97 years (sd = 14.18), with ages ranging from 18 to 86 years. other relevant demographic information pertaining to the sample is presented in table 1. table 1: characteristics of the participants who took part in the quantitative phase of the study (n = 483). during the third phase, six participants were purposively selected (tracy 2013) on the basis of having first-hand experience in what mnp or sanparks can realistically achieve in the development space. the participants consisted of a very experienced park manager who managed several national parks in south africa, a people and conservation officer who had also worked in multiple parks, the head of the park interface programme, a section ranger and two scientists from sanparks’ regional office in kimberley who worked in all the parks in the arid node. procedure and ethics the first phase of the study took place in early 2016 after ethics clearance was obtained from the north-west university’s human research ethics committee (nwu-00342-15-s1). once entry into the community had been negotiated, semi-structured interviews were conducted with key participants by the authors and, based on the findings, a structured questionnaire was developed that was subsequently administered to residents of the two communities during the second phase of the study, which took place 1 year later. eighteen fieldworkers (8 from ritchie and 10 from ratanang) were recruited from the respective communities and subsequently trained to administer the survey. fieldworkers were compensated financially for their services. informed consent was obtained from all participants. data were collected over a period of 1 week. privacy and confidentiality (which are also covered in the informed consent form) were protected by conducting the interviews in a place where the participants felt comfortable (e.g. their homestead) and by making sure that no one other than the researcher had access to the participant’s identifiable characteristics linked to specific results. furthermore, fieldworkers were required to sign confidentiality agreements. completed questionnaires were inspected on receipt to ensure completeness and correctness and were subsequently taken to the north-west university’s statistical consultation services for data capturing and analysis. during the third qualitative phase of the study, an interview guide and a structured questionnaire were developed based on the results of phases 1 and 2 and were subsequently utilised to gather data from six purposively selected senior members of mnp and sanparks. in total, the project involved 4 full weeks of fieldwork, which spanned a period of 3 years. data gathering methods during the first (qualitative) phase of the study, data were gathered by means of semi-structured interviews with purposively selected members of the two communities. an interview guide (tracy 2013) was used to facilitate the interviews. in particular, all participants were asked what they believed to be the most pressing needs in their respective communities. during the second (quantitative) phase of the study, a structured survey was developed on the basis of the themes that were identified from the first set of semi-structured interviews as outlined previously (see table 2 for the complete list of items). this strategy was followed in order to ensure that the scale was contextually sensitive and that the items on the scale tapped into actual community needs, as well as to enable the qualitatively identified needs to be empirically quantified and verified. specifically, participants were requested to rate each of the listed needs on a five-point likert scale ranging from 1 (‘no need’) to 5 (‘a very big need’). in addition, the survey was structured to gather biographical data pertaining to participants’ gender, age, race and employment status. table 2: community needs. during the third qualitative phase of the study, data were gathered by means of an interview schedule and structured questionnaire, which were developed on the basis of the findings that emerged from phases 1 and 2 of the study. participants were requested to rate a list of community needs on a five-point likert scale (ranging from 1 = ‘not at all’ to 5 = ‘to a great extent’) based on their professional view of the extent to which parks like mnp could potentially address each specific need. participants were then requested to qualitatively elaborate on the reason for their ratings via semi-structured interviews or questionnaires. data analysis qualitative data derived from phases 1 and 3 of the study were analysed by means of thematic content analysis (creswell 2003; tracy 2013). the transcribed data were first read multiple times to ensure immersion in the data, after which data were inductively coded by assigning a brief descriptive label to each segment of text. based on conceptual similarities, codes were grouped together into categories and overarching themes. these themes formed the basis for the development of the structured survey, which was used to gather data during the second round of the study. quantitative data were statistically analysed using spss 23 (field 2005). descriptive statistics (means and measures of central tendency) were calculated for all items. results phase 1: identified community needs the framework developed by sirgy et al. (2009) was used to cluster the 17 themes and subthemes that were identified via the thematic analysis of the semi-structured interviews (phase 1) under the four categories: business services, government services and non-profit-related services, as well as community conditions. table 2 summarises the main findings that emerged from the analysis and illustrates each with a relevant excerpt from the interviews. phase 2: empirical verification and quantification of community needs on the basis of the qualitative themes outlined in the previous section, a structured questionnaire was developed and subsequently administered during the second phase of the study. the survey results (see table 3) confirmed that all the needs that were identified during the qualitative phase of the study are actual needs in the communities, whilst it also enabled the empirical quantification of the relative strength and prevalence of these needs. table 3: needs of total group and two communities. according to the participants from both communities, the need for job opportunities was the most pressing (mean = 4.16, sd = 1.22). this was confirmed by the fact that 53.3% of adult participants surveyed were unemployed (see table 1). the second biggest need was for improved healthcare services (mean = 4.1, sd = 1.14). this was followed by a number of government-related services such as the need for improved infrastructure (mean = 4.09; sd = 1.14), improved waste management (mean = 4.08; sd = 1.13), improved municipal service delivery (mean = 4.07; sd = 1.11), improved safety and security (mean = 4.03; sd = 1.30), improved community policing (mean = 4.00; sd = 1.24), skills development and training (mean = 3.96; sd = 1.27) and improved social services (mean = 3.96; sd = 1.11). residents of both communities also felt that there was a significant need for an improvement in public education in their communities, both in terms of the quality and quantity of available educational institutions (mean = 3.95, sd = 1.09). the survey results also indicate that residents of ritchie and ratanang experienced the need for enhanced conservation of their local environments (mean = 3.91, sd = 1.2). a marked need for recreational and sporting facilities is evident not only from the survey results (mean = 3.9, sd = 1.2) but was particularly emphasised during the semi-structured interviews that were conducted with community members. other needs such as improved local leadership, agricultural support and training and the need for local economic development also achieved an above-average score. phase 3: extent to which mokala national park and south african national parks can potentially address identified community development needs during the third phase of the study, semi-structured qualitative interviews as well as structured and unstructured questionnaires were used to gather data from purposively selected staff members from mnp and sanparks as well as other experts in relation to the extent to which parks such as mnp can potentially address the development needs that were identified by the communities. whilst the sample was small, given the expertise of the participants and the need to identify the relative feasibility of each need being addressed, it was deemed to be of value to quantitatively assess all participants’ views on the feasibility of each need being addressed (which was done on a five-point scale ranging from 1 = ‘not at all’ to 5 = ‘to a great extent’). this rating was used as the basis for a qualitative exploration of why the participant assigned the given score. participants’ responses to the survey are reflected in table 4. following on table 4, the needs – in order of their ranked importance and based on the themes derived from the qualitative data – will be discussed in greater detail. table 4: extent to which mokala national park and south african national parks could potentially address development needs in communities. more temporary job and employment opportunities according to the participants, the biggest direct impact that mnp and sanparks can potentially have is via temporary job or employment opportunities (mean = 4.00). as sketched by one of the participants: ‘mokala currently employs nine contractors and each contractor employs 10 or 11 people. this means that nearly 100 contract workers are currently working on one of the expanded public works programme (epwp)/biodiversity social project (bsp) activities in the park, for example infrastructure projects, working for ecosystems, working for water, environmental monitors, et cetera.’ (participant 22, male, park manager) according to the same participant, these temporary job opportunities are creating employment in all national parks in south africa. the participant also commented that these projects have a lot of potential to address needs related to employment, because the projects’ lifecycle typically extends over a period of 2 to 3 years and are often repeated, based on the availability of funding and the need for such services. the participants also indicated that another opportunity for job creation in parks is to offer concessions to private investors, for example, by outsourcing a park’s restaurants, shops, et cetera. according to one of the participants, some of the larger parks such as the kruger national park (knp) are already engaging in this practice, resulting in an increase in the availability of jobs for people from local communities. however, as pointed out by the same participant, this is currently not a feasible strategy at mnp because the park is very remote, its restaurant is not showing much of a profit and accommodation for staff is limited. local conservation initiatives according to expert participants, the second biggest potential impact that mnp and sanparks could have is in the local conservation space, especially via environmental education (ee). not only is this in line with sanparks’ primary mandate, but sanparks already has dedicated staff who can attend to this (its people and conservation staff) and has a whole array of educational material on hand. as one participant pointed out, ‘sanparks also has a fully developed curriculum for environmental education and it has specific programmes aimed at stimulating local conservation initiatives such as its kids in park programme’ (participant 23, female, people and conservation officer). according to the same participant, part of mnp’s efforts to stimulate local conservation initiatives includes taking children from local communities to visit the park for up to 2 to 3 days to learn and to be exposed to natural environments and wildlife. this programme, according to another participant (participant 25, male, head of the park interface programme), has been rolled out at all the local schools in the area. skills development and training the third biggest potential impact that mnp and sanparks can have is via skills development and training. this is largely because of the government-funded bsp/epwp programmes. according to the participants, training is specifically budgeted for in each of these projects because it is mandatory to train community members temporarily employed through bsp/epwp in a variety of fields like health, safety and first aid. furthermore, given that many of these projects are often infrastructure related, for example, building staff accommodation at and erecting a new entrance gate for mnp’s lilydale camp, participants have the opportunity to acquire practical skills pertaining to the construction industry, which they can subsequently apply in their own communities in support of development (e.g. building various structures). according to the participants, contractor development also formed an important part of skills development (business management, health and safety, etc.) in the past, the idea being that sanparks should help to empower contractors so that they can secure other job opportunities (contracts) once their contracts with sanparks have expired. local economic development according to one of the participants, mnp spends a portion of its budget in the local economies of jacobsdal and modderrivier, but the majority of its budget is spent in kimberley: ‘we do buy local as far as possible. however, some of the foods are not locally available. we therefore have to get it from kimberley’ (participant 22, male, park manager). however, according to the same participant, mokala, like all other parks, spends a large portion of its budget on human resources. according to him, at least some of the staff reside in local communities, implying that they also spend a portion of their income at local shops in towns near parks. as such, parks likely have a significant indirect impact on local economic development by means of the disposable income their employees spend in these communities. another potential option, according to the same participant, is to focus on supply-chain development. he explained that mnp currently purchases all the meat it uses in its restaurant from a local butcher near the park. most of the dry food is, however, bought in kimberley. according to him, much more can be done to identify, source and register local suppliers. in fact, mnp recently undertook an exercise to meet and register local service providers. a potential obstacle that was identified in this process is that suppliers have to be registered on a government database. however, this process is reported to be fairly rapid and inexpensive and, as such, need not pose an undue obstacle to local community members registering their businesses as suppliers. in this regard, national parks like mnp can and have played a supportive role by assisting new service providers with the registration process. agricultural support and training according to the participants, mnp and some of the other national parks are in the process of providing agricultural support and training to local communities. this is currently happening in the salt lake community near mokala. other parks, such as the karoo national park, also focused on this in the past with the help of external service providers. in this regard, the participants felt that the people and conservation officers can make a huge contribution and that basic skills can be transferred to local communities near parks, which, in turn, can contribute to food security in these communities. another participant also indicated that yet another way in which sanparks currently supports agriculture is through its wildlife economy programme, an initiative whereby sanparks gives animals to upcoming farmers on loan to build up their own breeding stock for commercial purposes. mnp is also taking part in this initiative. more permanent job and employment opportunities the participants indicated that mnp and sanparks’ capacity to provide permanent job opportunities is limited (mean = 2.33). this state of affairs was confirmed in the present study where only two individuals from the ritchie community were reported to be employed by mnp and only one resident was employed by sanparks’ regional office in kimberley. this, according to the participants, can be ascribed to the fact that mokala is still a developing park and small compared to other parks: ‘there are some expectations, but mokala is a small park compared to kruger and can only contribute a small fraction’ (participant 23, female, people and conservation officer). according to another participant, the creation of job opportunities in parks will also depend on the staff turnover at the parks (via natural attrition, resignations and retirements). this, according to him, is relatively low at present, thus limiting the capacity of parks like mnp to address this need in a feasible manner. another limiting factor that has an influence on the creation of job opportunities, according to the participants, is that some of the higher (more specialised) post levels require some formal education (graduate or postgraduate qualification), which ordinary community members from local communities often do not have. one of the participants also indicated that in her experience, local community members are not aware of the employment opportunities that exist in parks. as a result, mnp is in the process of implementing the yes programme, which is a programme by the national government that is aimed at creating awareness among the youth of the type of job opportunities that exist in parks and within other government institutions. improved quality and quantity of education the participants agreed that they, as a park and as part of sanparks, will not be able to improve the quality of education offered by local primary and secondary schools, nor the quantity of those schools in the area. they did, however, concede that sanparks can support the curriculum by providing context and practical exposure to school children, especially via their ee programmes and initiatives. recreation facilities and activities according to one of the participants, sanparks can play a supporting role in the development of municipalities’ integrated development plans, especially when it comes to providing inputs regarding the development of urban parks and open spaces. a second option, according to the participants, is for communities to make use of the free-entrance week that is normally offered by all national parks in south africa in the course of september. yet another alternative mentioned by participant 22 is to make use of the honorary ranger’s project that, among others, involves hosting an annual mountain bike race in mnp. this initiative, according to him, has a lot of potential to involve developmental groups from the local community. improved waste management although mnp, and parks in general, cannot take the responsibility for waste management outside the park or in local communities, the participants indicated that improved waste management is a need that they share with local communities. one of the participants also indicated that mnp is in the process of developing a recycling plant at the park. according to him, this can potentially provide opportunities for local communities. however, given that there are no waste depots or landfill sites near the park, all waste will have to be transported to the town of kimberley over a distance of 90 km, which is likely to affect the feasibility of this strategy adversely from a community development perspective. access to tertiary education according to the participants, sanparks at present only offers bursaries to its own staff and their children, which implies that even though some of these employees do reside in local communities, the capacity to address this need is limited. however, sanparks also has a junior scientist programme that provides opportunities for upcoming scientists to obtain a master’s degree or phd. according to one of the participants, the programme started in knp and will be expanded to other parks as well. in addition, sanparks is providing in-service training to graduates with a diploma or degree in conservation, and some of the beneficiaries have even been absorbed as section or field rangers or biotechnicians. improved infrastructure the participants indicated that it would be difficult (if not impossible) for them as a park to address the larger infrastructural needs of their surrounding communities. however, according to them, sanparks does have a socio-economic development department that assists with some of the infrastructure needs in communities (e.g. at schools and with regards to libraries). these interventions are funded by a percentage of the conservation fees that are paid by tourists. improved leadership here, too, the participants agreed that sanparks cannot really make a significant direct contribution. however, according to one of the participants, sanparks is making an indirect contribution through its kids in parks programme, because this programme is aimed at instilling leadership ethics at an early age, especially with regards to the disciplines of conservation and biodiversity. improved municipal service delivery the participants indicated that they, as a park and sanparks in general, will not be able to address needs related to service delivery in local communities outside the park. the participants were fairly unanimous that this is the responsibility of local municipalities. improved community safety, security and policing participants felt that the need for improved safety and security and better policing in communities cannot be feasibly addressed by sanparks to any significant extent. nevertheless, given that parks do indeed take measures to ensure security both inside the park and along their boundaries, it stands to reason that their immediate neighbours must be benefitting from these measures to some degree. establishing more schools the participants unanimously felt that mnp and sanparks will not be able to make a significant contribution towards the actual building of more schools. improved availability of and/or access to healthcare services (e.g. clinics) participants indicated that the need for improved availability of and/or access to healthcare is a need they share with local communities. according to them, some of their staff members have to travel all the way to kimberley to get treatment and/or chronic medication. because the park is quite a distance from kimberley, this involves hours of travel and adversely affects productivity. negotiations with the department of health to build a clinic nearby or in the park are currently underway. nevertheless, participants indicated that sanparks’ direct potential contribution to addressing this need is minimal. improved social services (e.g. social workers) all the participants agreed that addressing the need for improved social services falls completely outside their mandate and capacity. discussion this study investigated the feasibility of national parks in south africa endorsing a community development agenda, using mnp and two neighbouring communities as case study. a sequential mixed-methods approach was followed, comprising three phases that spanned a 3-year period. in relation to the first two phases of the study, which aimed to investigate community needs both qualitatively and quantitatively, findings revealed that the communities’ most significant needs centre around employment opportunities; improved healthcare, service delivery and waste management; an improvement in the quality of and access to primary and secondary education opportunities (i.e. schools); and more opportunities for local residents to afford and access tertiary education opportunities. community members also expressed the need for improved community policing, safety and security; social services and agricultural support and training as well as general skills development and training, local leadership, recreational facilities, local economic development and conservation initiatives. as will be outlined in the section to follow, the results from the third phase of the study suggest that mnp (and sanparks in general) could realistically only address some of the identified community needs to a significant degree. the need for employment opportunities as well as the need for skills development and training ranked tops among those identified by the communities targeted in this study. given that studies by tepela and omara-ojungu (2012) in communities near knp and by pelser et al. (2013) near the golden gate highlands national park revealed similar results, it should come as no surprise that addressing the high unemployment rate in south africa – identified as one of the ‘five key pillars’ of national government – has been adopted as a key element in sanparks’ vision. in reaction to national government’s call to address the unemployment rate, mnp/sanparks indicated that their biggest potential contribution would be in the form of temporary job creation, mainly via the epwp/bsp programmes. in 2015–2016, three epwp/bsps were in operation at mokala (spies 2015). since then, this number has increased to nine – which shows that there is still potential for growth. findings derived from the semi-structured interviews in this study confirmed that members of the ritchie community did indeed benefit from the epwp/bsp initiative and that over 100 local community members are currently benefitting from these programmes initiated by mnp, but the findings also revealed that several challenges are being experienced in as far as job allocation and the timely settlement of payments are concerned. from the park’s perspective, spies (2015) found the initiatives to be fairly successful, despite some challenges. according to sanpark’s 2015–2016 annual report, epwp/bsp programmes are on offer in all 19 south african national parks, and 23  298 local community members are being employed on a semi-permanent basis (given that projects typically span 2 to 3 years and are often renewed). these projects simultaneously also address the community need for skills development and training, as all temporary employees are being trained in a whole array of skills ranging from first aid to construction, some of which might increase their future employability and/or their ability to support local economic or infrastructure development in their own communities by, for example, making use of the skills they have acquired in the construction industry to erect structures in their own communities. the creation of permanent jobs, though, seems to be a totally different matter that depends on the size of a particular park, its potential for expansion and the turnover of existing staff members. however, given that sanparks employs only around 4000 people countrywide (sanparks 2016) and that funds allocated to national parks are limited (sanparks 2016), the number of people that can be employed by the organisation on a permanent basis is clearly limited. two potential avenues for job creation that have probably hitherto been underutilised are to make private concessions available to private investors and to create more opportunities for self-employment. in case of the former, it is likely that private investors will have the capital to grow sanparks’ facilities such as restaurants and accommodation and, in the process, to create more job opportunities. given that mnp and other parks are already engaging in the practice of sourcing supplies locally, another feasible strategy that could be investigated would be to establish and develop vetted local supply chains with communities adjoining national parks. given that they are mostly rural (i.e. agricultural), residents could be afforded the opportunity to provide parks with an array of fresh vegetables, fruit and meat or an assortment of arts and crafts. such a practice is also likely to make a meaningful contribution to local economic development. the capacity to address the need for local conservation initiatives was ranked second highest. given that environmental conservation is one of its core mandates, national parks can definitely play a significant role in this regard. not only would such initiatives fall within sanparks’ expertise, but the organisation also has the capacity to address these needs via its people and conservation division (a programme by sanparks that is used to drive the dual agenda of conservation and development) and the various extension programmes it has on offer. furthermore, sanparks can also play a role in the development of new conservation areas in traditional and cultural land via collaborative engagement with local communities. a very pertinent need that was identified in the communities was for recreational facilities and activities. whilst participants felt that direct involvement in the establishment of recreational facilities generally falls outside the scope of sanparks’ mandate, the results of this study do suggest a couple of strategies that could be assessed for feasibility. these include sponsoring recreational facilities (e.g. participants in both communities expressed the need for a skateboard park), sponsoring local residents’ participation in the annual park mountain bike race or promoting the park itself as recreational facility, perhaps via sponsoring occasional transport and entry into the park as is currently the case with the free-entrance week that usually occurs in september of each year. the latter strategy would represent a synergistic approach as it combines the park’s core asset with a community need, which, as repeatedly revealed in the qualitative interviews, is likely to make a significant positive difference in the lives of residents of impoverished communities. the findings suggest that mnp and other parks can play a significant role in agricultural support and training. in this regard, parks can partner with external service providers and provide wildlife to emerging farmers. agricultural support and development is an international priority (linked to sustainable development goals) simply because it has a bearing on food security (fao 2018). it also forms an integral part of south africa’s national development plan, given that it is part of government’s efforts to stimulate integrated and inclusive rural economies in south africa (the conversation 2017). a shared need that was identified by the communities and mnp was for improved waste management. whilst direct involvement in refuse removal falls outside the auspices of sanparks’ community development mandate and cbc initiatives, in this case, it could potentially play a role in terms of promoting environmental awareness and education. this would also be in the interest of national parks, given that refuse and pollution generated by local communities adjacent to national parks have the potential to impact the park and visitors’ perceptions and experiences of the park adversely. in partnership with local authorities, the barriers that prevent people from keeping their communities clean can be identified and addressed. for example, macallister (2015) found that there were instances where managing farm waste as a health problem served as a means to help address this issue in communities and to prevent concomitant health issues. although mnp indicated that it cannot directly contribute to infrastructure development in local communities (e.g. roads, buildings, sanitation, etc.), it can play a role through its socio-economic development initiatives by supporting local supply-chain development and providing skills (such as construction skills) and opportunities to local community members. the beneficiaries of these initiatives can then, in turn, use the skills they have acquired to benefit their communities. a huge need that was identified by the communities was to improve education. whilst the findings clearly reveal that sanparks can only play a limited role in the context of formal education and cannot do much to increase the quantity and quality of schools in the region, it can nevertheless play a meaningful educational role by providing new learning opportunities and contexts for children. a number of parks already have some very good educational programmes and projects in place. included among these are the heritage project imbewu (developed to promote the transference of traditional cultural knowledge from older to younger people), the junior rangers programme (aimed at promoting civil service and volunteerism in parks), the kids in parks programme (aimed at exposing youngsters to parks and pristine natural environments) and the koedoe green schools programme (an ee programme focusing on, among others, climate change) (sanparks 2017). the potential value of projects such as these were underscored in the present study, where several participants reported that their children showed a marked change in their attitudes and enthusiasm towards the natural environment after a visit to mnp. findings from the interviews revealed that whilst children would often kill animals such as snakes on sight, those who experienced a guided visit to mnp not only ceased this practice but actively encouraged others in the community to refrain from doing so. nevertheless, whilst valuable in itself, exposure to parks and ee will likely not promote the educational agenda necessary for people to matriculate with flying colours or to find gainful employment. this notion is supported by van der berg et al. (2011), who found that the level of education in rural communities bordering national parks is typically insufficient for learners to further their tertiary education at a university or college. limited access to tertiary educational institutions (which was also found in the survey to be a significant community concern and need) and lack of funding, as well as lack of awareness of funding opportunities, conspire to exacerbate this problem. even though both government and universities are currently doing a lot to assist communities in this regard (dhet 2017), national parks could also potentially play a significant role by, for example, offering bursaries to communities for those who wish to study science or conservation. in the long run, this will enhance the probability that residents originating from these communities may well end up finding permanent employment within a park structure. finally, the results indicate that there is very little that mnp and sanparks can feasibly do to directly address needs related to healthcare services (even though they share the need), community safety and security, community policing, social services (e.g. social workers), local leadership and the establishment and erection of more schools, as these needs generally fall outside the auspices and capacity of national parks in south africa. elsewhere, naughton-treves, buck holland and brando (2005) came to the same conclusion, whilst andam et al. (2010) also questioned the actual impact national parks could have on poverty. in light of the above, it seems necessary to rethink and possibly reconceptualise the developmental agenda that national parks are expected to drive so that these aims are more in line with (1) what parks can reasonably achieve based on resources, capacity and expertise and (2) what communities surrounding national parks need according to their own articulation. failure to do so could invite the risk of creating unrealistic expectations among communities, resulting in an increased likelihood of general resentment and negative attitudes towards conservation among communities when such expectations are not fulfilled (anthony 2007). furthermore, though well-intentioned, community development initiatives that are not aligned with community needs or the capacity, resources and expertise of national parks are likely to either result in costly failures or might fail to truly have a meaningful impact on communities. it is hoped that the findings of this study will be of use in informing sanparks’ community development initiatives and ensuring that those initiatives are optimally aligned with actual community needs. conclusion using a sequential mixed-methods design, this study set out to investigate the feasibility of national parks in south africa endorsing a community development agenda based on an exploration of community needs using mnp and two neighbouring rural communities as a case study. based on the results, the feasibility of national parks substantively addressing all community needs is questioned as many of these needs fall outside the scope, capacity, skills and resource capability of the organisation. nevertheless, as was revealed in this study, national parks have the ability to directly or indirectly support several pressing community needs by creating jobs (especially by means of temporary projects and, to a far lesser extent, by means of direct employment) and by enabling those who partake in such employment opportunities to be trained and to develop specific skills. furthermore, initiatives aimed at promoting awareness and education around waste management and local environmental conservation, supporting local economic development, sponsoring recreational facilities and/or access to parks as well as offering agricultural support and training have been identified as means whereby mokala can respond to the needs experienced by its surrounding communities. such initiatives would not only articulate with genuine community needs but could be feasibly pursued based on sanparks’ expertise, resources and core mandate and thus hold the promise of truly achieving a positive impact on community development. acknowledgements competing interests the authors declare that they have no financial or personal relationship that may have inappropriately influenced them in writing this article. authors’ contributions both authors contributed equally to conceptualising the study as well as towards collecting and analysing the data and writing the manuscript. funding information this study was funded through a strategic research grant of the unit of environmental sciences and management, faculty of natural and agricultural sciences, north-west university. references andam, k.s., ferraro, p.j., sims, k.r.e., healy, a. & holland, m.b., 2010, ‘protected areas reduced poverty in costa rica and thailand’, proceedings of the national academy of sciences 107(22), 9996–10001. https://doi.org/10.1073/pnas.0914177107 anthony, b., 2007, ‘the dual nature of parks: attitudes of neighbouring communities towards kruger national park, south africa’, environmental conservation 34(3), 236–245. https://doi.org/10.1017/s0376892907004018 barrow, g.c., 2015, ‘the socio-economic benefits of new national parks designations in scotland’, association for the protection of rural scotland 1, 2–13. bennet, n.j., 2014, ‘why local people do not support conservation: community perceptions of marine protected area livelihood impacts, governance and management in thailand’, marine policy 44, 107–116. https://doi.org/10.1016/j.marpol.2013.08.017 berkes, f., 2004, ‘rethinking community-based conservation’, conservation biology 18(3), 621–630. https://doi.org/10.1111/j.1523-1739.2004.00077.x berkes, f., 2007, ‘community-based conservation in a globalized world’, pnas 104(39), viewed 10 january 2017, from http://www.pnas.org/content/104/39/15188.full.pdf biggs, d., swemmer, l., phillips, g., stevens, j., freitag, s. & grant, r., 2014, ‘the development of a tourism research framework by south african national parks to inform management’, koedoe 56(2), art. #1164, 9 pages. botha, j., witkofski, e.t.f. & cock, j., 2007, ‘is “something better than nothing”? the impacts of ten outreach nurseries on south african community stakeholders’, forests, trees and livelihoods 17, 199–221. https://doi.org/10.1080/14728028.2007.9752597 cock, j. & fig, d., 2000, ‘from colonial to community based conservation: environmental justice and the national parks of south africa. society in transition 31(1), 22–35. https://doi.org/10.1080/21528586.2000.10419008 conley, a. & moote, m.a., 2003, ‘evaluating collaborative natural resource management’, society & natural resources 16(5), 371–386. https://doi.org/10.1080/08941920309181 cox, m., arnold, g. & villamayor-tomas, s., 2010, ‘a review of design principles for community-based natural resource management’, ecology and society 15(4), 38. https://doi.org/10.5751/es-03704-150438 creswell, j.w., 2003, research design: qualitative, quantitative, and mixed methods approaches, sage, thousand oaks, ca. creswell, j.w., 2013, qualitative inquiry and research design choosing among five approaches, sage, thousand oaks, ca. creswell, j.w. & plano clark, v.l., 2007, designing and conducting mixed methods research, sage, thousand oaks, ca. department of higher education and training, 2017, official website, viewed 10 january 2017, from http://www.dhet.gov.za/ emerton, l., 2001, ‘the nature of benefits and the benefits of nature: why wildlife conservation has not economically benefited communities in africa’, in d. hulme & m. murphree (eds.), african wildlife and livelihoods: the promise and performance of community conservation, pp. 208–226, david philip publ, cape town, south africa. faasen, h., 2006, ‘synergies between biodiversity conservation and sustainable rural development of adjacent communities: a case study of the tsitsikamma national park’, phd thesis, university of stellenbosch, faculty of agriculture and forestry sciences. field, a., 2005, discovering statistics using spss, sage, london. flora, c.b., flora, j.l. & gasteyer, s.p., 2015, rural communities: legacy and changes, westview press, boulder, co. food and agriculture organization of the united nations, 2018, viewed 10 august 2018, from http://www.fao.org/sustainable-development-goals/en/ iucn, 2010, policy matters 17, viewed 10 january 2017, from https://www.iucn.org/downloads/policy_matters_17___pg_173_204.pdf johnson, r.b., onwuegbuze, a.j. & turner, l.a., 2007, ‘toward a definition of mixed methods research’, journal of mixed methods research 1(2), 112–133. https://doi.org/10.1177/1558689806298224 kusters, k., achdiawan, r., belcher, b. & perez, m., 2006, ‘balancing development and conservation? an assessment of livelihood and environmental outcomes of nontimber forest product trade in asia, africa, and latin america’, ecology and society 11(2), 20. https://doi.org/10.5751/es-01796-110220 lundmark, j.l., fredman, p. & sandell, k., 2010, ‘national parks and protected areas and the role for employment in tourism and forest sectors: a swedish case’, ecology and society 15(1), 19. https://doi.org/10.5751/es-03175-150119 macallister, j., 2015, factors influencing solid-waste management in the developing world, all graduate plan b and other reports. paper 528, utah state university, digitalcommons@usu, utah. machlis, g.e. & field, d.r., 2000, national parks and rural development: practice and policy in the united states, island press, washington, dc. mansuri, g. & rao, v., 2004, ‘community-based and -driven development: a critical review’, world bank research observer 19(1), 1–39. https://doi.org/10.1093/wbro/lkh012 mohan, g., 2009, ‘the significance of rural areas in south africa for tourism development through community participation with special reference to umgababa, a rural area located in the province of kwazulu-natal’, university of south africa, pretoria, viewed 16 january 2017, from http://hdl.handle.net/10500/2159 murphee, m.w. & hulme, d., 2001, african wildlife and livelihoods: the promise and performance of community conservation, james currey ltd, oxford. naughton-treves, l., holland, m. & brando, k., 2005, ‘the role of protected areas in conserving biodiversity and sustaining local livelihoods’, annual review of environment and resources 30, 219–252. https://doi.org/10.1146/annurev.energy.30.050504.164507 nelson, j.g. & serafin, r., 2013, national parks and protected areas: keystones to conservation and sustainable development, nato asi book series 40, springer-verlag, berlin heidelberg. newmark, w.d. & hough, j.l., 2000, ‘conserving wildlife in africa: integrated conservation and development projects and beyond’, bioscience 50(7), 585–592. https://doi.org/10.1641/0006-3568(2000)050[0585:cwiaic]2.0.co;2 pelser, a., redelinghuis, n. & velelo, n., 2013, protected areas as vehicles in population development: lessons from rural south africa, viewed 10 january 2017, from http://conferences.ufs.ac.za/dl/userfiles/documents/00001/654_eng.pdf petursson, j.g. & vedeld, p., 2017, ‘rhetoric and reality in protected area governance: institutional change under different conservation discourses in mount elgon national park, uganda’, ecological economics 131, 166–177. https://doi.org/10.1016/j.ecolecon.2016.08.028 pullin, a.s, 2014, assessing the effects of terrestrial protected areas on human well-being: astap advisory document, global environmental facility, washington, dc. rights and resource initiative, 2015, protected areas and the land rights of indigenous peoples and local communities: current issues and future agenda, viewed 10 january 2017, from file:///c:/users/12894451/desktop/rrireport_protectedareas-and-landrights_web.pdf sanparks, 2012, sanparks annual performance plan, viewed 10 january 2017, from https://www.sanparks.org/assets/docs/about/annual_performance_plan_2012013.pdf sanparks, 2016, sanparks annual report 2015/16, viewed 10 january 2017, from https://www.sanparks.org/assets/docs/general/annual-report-2016.pdf sanparks, 2017, official website of south african national parks, viewed 10 january 2017, from https://www.sanparks.org/about/connecting_to_society/ sirgy, m.j., widgery, r.n., lee, d. & yu, g.b., 2009, ‘developing a measure of community well-being based on perceptions of impact in various life domains’, social indicators research 96, 295–311. https://doi.org/10.1007/s11205-009-9479-9 spies, m.o., 2015, ‘an exploration of whether the people and parks programme contributes to capacity building and awareness in the ritchie community near mokala national park’, unpublished honours dissertation, north-west university, potchefstroom campus. stats-sa, 2011, official website of statistics south africa, viewed 10 january 2017, from http://www.statssa.gov.za/?page_id=3839 stevens, s., 2014, indigenous peoples, national parks, and protected areas: a new paradigm linking conservation, culture, and rights, university of arizona press, tucson, az. stolton, s. & dunley, n., 2015, values and benefits of protected areas, anu press, canberra, australia. swemmer, l.k. & taljaard, s., 2011, ‘sanparks, people and adaptive management: understanding a diverse field of practice during changing times’, koedoe 53(2), 1–7. https://doi.org/10.4102/koedoe.v53i2.1017 tashakkori, a. & teddlie, c., 2003, handbook of mixed methods in social and behavioral research, sage, thousand oaks, ca. tepela, b.n. & omara-ojungu, p.h., 2012, ‘towards bridging the gap between wildlife conservation and rural development in post-apartheid south africa: the case of the makuleke community and the kruger national park’, south african geography journal 81(3), 1999. the conversation, 2017, how land reform and rural development can help reduce poverty in south africa, viewed 10 august 2018, from http://theconversation.com/how-landreform-and-rural-development-can-help-reduce-poverty-in-south-africa84146 tisdell, c., 1999, biodiversity, conservation and sustainable development: principles and practices with asian examples, cheltenham, edward elgar. tracy, s.j., 2013, qualitative research methods: reflecting evidence, crafting analysis, communicating impact, wiley-blackwell, london. van der berg, s., taylor, s., gustafsson, m., spaull, n. & armstrong, p., 2011, improving education quality in south africa, report for the national planning commission, department of economics, university of stellenbosch. vira, b. & kontoleon, a., 2010, ‘dependence of the poor on biodiversity: which poor, what biodiversity?’, cprc international conference 2010, manchester, september 8–10, 2010. wells, m.p. & mcshane, t.o., 2004, ‘integrating protected area management with local needs and aspirations’, ambio 33, 513–519. https://doi.org/10.1579/0044-7447-33.8.513 western, d., wright, r.m. & strum, s.c., 2013, perspectives in community based conservation, island press, washington, dc. footnote 1. a ‘trade-off park’ is where one park (normally an existing park) is de-proclaimed so that it can be used for reasons other than conservation (normally development), and in its place another park is proclaimed. abstract introduction methods review findings implications and recommendations for establishing exclosures conclusion acknowledgements references about the author(s) corli wigley-coetsee savanna and grassland node, scientific services, kruger national park, skukuza, south africa school of natural resource management, george campus, nelson mandela university, george, south africa tercia strydom savanna and grassland node, scientific services, kruger national park, skukuza, south africa department of soil, crop and climate sciences, university of the free state, bloemfontein, south africa danny govender savanna and grassland node, scientific services, kruger national park, skukuza, south africa department of paraclinical sciences, university of pretoria, pretoria, south africa dave i. thompson saeon ndlovu node, scientific services, kruger national park, phalaborwa, south africa school of geography, archaeology and environmental studies, university of the witwatersrand, johannesburg, south africa navashni govender school of natural resource management, george campus, nelson mandela university, george, south africa conservation management, kruger national park, skukuza, south africa judith botha savanna and grassland node, scientific services, kruger national park, skukuza, south africa chenay simms savanna and grassland node, scientific services, kruger national park, skukuza, south africa adolf manganyi savanna and grassland node, scientific services, kruger national park, skukuza, south africa laurence kruger organization for tropical studies, faculty of science, university of cape town, cape town, south africa jacques venter savanna and grassland node, scientific services, kruger national park, skukuza, south africa cathy greaver savanna and grassland node, scientific services, kruger national park, skukuza, south africa izak p. smit savanna and grassland node, scientific services, kruger national park, skukuza, south africa centre for african ecology, school of animal, plant and environmental sciences, university of the witwatersrand, johannesburg, south africa citation wigley-coetsee, c., strydom, t., govender, d., thompson, d.i., govender, n., botha, j. et al., 2022, ‘reflecting on research produced after more than 60 years of exclosures in the kruger national park’, koedoe 64(1), a1674. https://doi.org/10.4102/koedoe.v64i1.1674 note: additional supporting information may be found in the online version of this article as online appendix 1 and online appendix 2. review article reflecting on research produced after more than 60 years of exclosures in the kruger national park corli wigley-coetsee, tercia strydom, danny govender, dave i. thompson, navashni govender, judith botha, chenay simms, adolf manganyi, laurence kruger, jacques venter, cathy greaver, izak p. smit received: 19 feb. 2021; accepted: 01 dec. 2021; published: 28 feb. 2022 copyright: © 2022. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract herbivores are a main driver of ecosystem patterns and processes in semi-arid savannas, with their effects clearly observed when they are excluded from landscapes. starting in the 1960s, various herbivore exclosures have been erected in the kruger national park (knp), for research and management purposes. these exclosures vary from very small (1 m2) to relatively large (almost 900 ha), from short-term (single growing season) to long-term (e.g. some of the exclosures were erected more than 60 years ago), and are located on different geologies and across a rainfall gradient. we provide a summary of the history and specifications of various exclosures. this is followed by a systematic overview of mostly peer-reviewed literature resulting from using knp exclosures as research sites. these 75 articles cover research on soils, vegetation dynamics, herbivore exclusion on other faunal groups and disease. we provide general patterns and mechanisms in a synthesis section, and end with recommendations to increase research outputs and productivity for future exclosure experiments. conservation implications: herbivore exclosures in the knp have become global research platforms, that have helped in the training of ecologists, veterinarians and field biologists, and have provided valuable insights into savanna dynamics that would otherwise have been hard to gain. in an age of dwindling conservation funding, we make the case for the value added by exclosures and make recommendations for their continued use as learning tools in complex african savannas. keywords: disease; fire; herbaceous layer; herbivory; plant productivity; savanna; species diversity. introduction the earliest research exclosures were erected in the kruger national park (knp) in the late 1960s and early 1970s (e.g. makhohlola exclosure) with the intention ‘to provide areas where the grazing effects of animals could be excluded’ (joubert 1986). this was done partly to complement the experimental burn plots (ebps), a fire-manipulation experiment initiated in the 1950s which focused on studying the effects of different fire seasons and frequencies on vegetation, but where herbivores had free access (see biggs et al. 2003). however, some of these fenced-off areas (e.g. hlangwini and n’waswitshumbe exclosures) were also used to support a range of operational management needs, including the breeding of locally rare antelope species (e.g. roan, hippotragus equinus equinus and tsessebe, damaliscus lunatus lunatus) and as holding areas for reintroduced white (ceratotherium simum simum) and black (diceros bicornis) rhinoceros (e.g. hlangwini and black rhino exclosures). after the detection of widespread tuberculosis in buffalo (syncerus caffer) in southern knp, a buffalo breeding enclosure was set up at satara in 2000 (i.e. satara [buffalo] exclosure). then, in 2002, herbivore exclosures were erected along the letaba and sabie rivers after extensive floods removed most of the riparian vegetation (i.e. letaba and nkhuhlu exclosures) (o’keefe & alard 2002). these exclosures include both full exclusion (i.e. where all but the smallest mammals are excluded) and partial exclusion (excluding only elephant [loxodonta africana] and, because of their height, giraffe [giraffa camelopardalis giraffa]), and each divided into burned and unburned sections. the primary purpose here was to understand the effects of herbivore size and the interaction with fire, on vegetation dynamics. a sub-objective was to document the recovery of riparian vegetation after the floods in 2000 (o’keefe & alard 2002). during the early 2000s, several external research programmes which included exclosures in their experiments, were established in knp. the university of cape town (uct) erected two electrified exclosures, one on the october triennial fire treatment of the shabeni ebp string at pretoriuskop and another in the north-west corner of the satara buffalo exclosure (hereafter uct satara exclosure). these exclosures formed part of the broader tree grass programme, with the main objective to investigate hypotheses predicting tree-grass coexistence in savannas (february & higgins 2010). in addition, several international universities (california santa barbara, colorado state, florida international, new mexico, north carolina greensboro, wyoming, and yale) and local institutions (university of kwazulu-natal and south african environmental observation network [saeon]; hereafter the usa-sa consortium) have, since 2005, used multiple small exclosures near satara (including on the ebps and in the satara (buffalo) exclosure) to study the interactions between landscape position, fire, rainfall and herbivore size on herbaceous dynamics (buis et al. 2009; koerner et al. 2014). full exclosures exclude all herbivores larger than a hare (buis et al. 2009) and partial exclosures only allowed access to herbivores ≤ 0.85 m at shoulder height (e.g. impala [aepyceros melampus] and steenbok [raphicerus campestris]) (burkepile et al. 2016). in 2018, researchers from several institutions (universities of florida and eswatini being prominent), erected an exclosure experiment (called browse, including two full and three partial exclosures) south of lower sabie tourist camp, close to the location of the makhohlola exclosure. this experiment is not reviewed here as the exclosures are relatively new. apart from these, a host of other external research groups have used small exclosures (i.e. 1 m2), short-term (one growing season) and long-term, to answer various research questions. although exclosures can be valuable research sites, they are expensive research infrastructure to install and maintain, especially in areas with high densities of elephants (grant et al. 2008; kioko et al. 2008). although many of the exclosures were initially erected using external funds (e.g. andrew. w. mellon foundation and the national science foundation [usa]), south african national parks (sanparks) assumed responsibility for their maintenance and repair, with the assistance of saeon in some cases. sanparks, like many other conservation agencies with largely self-generated funding through tourism and with a decreasing government grant, has limited funds to acquire and maintain research infrastructure. additionally, sanparks is registered as a non-profit parastatal agency, which means operational funds are not guaranteed and cannot be carried over from year to year, thus making long-term budgeting difficult. many of the exclosures (e.g. uct shabeni, makhohlola, black rhino exclosure, capricorn; see online appendix 1 or figure 1 for current status) have seen a steady deterioration of fences or have been decommissioned because of the rising costs of maintaining what is now an ageing infrastructure, in an environment with an increasing population of elephants (smit et al. 2020). the more recent financial impacts of the global covid-19 (coronavirus disease 2019) pandemic on tourism specifically, has further exacerbated the situation (smith et al. 2021), with wide-scale cost curtailment across sanparks. figure 1: map of the kruger national park showing the locations and status of past and present exclosures (map produced in december 2021). here, we provide a review of exclosure-based research in knp. additionally, we were interested to evaluate the realised impact of these exclosures from the perspective of research output. research published in peer-reviewed literature was reviewed under five different groupings, based on the history and objectives of the respective exclosures; that is, (1) the exclosures with the longest history (hlangwini, n’waswitshumbe and makhohlola), (2) the nkhuhlu and letaba exclosures, (3) disease work in the satara (buffalo) exclosure, (4) the tree grass programme (uct shabeni and uct satara), and (5) a grouping capturing all the smaller exclosure work around satara. online appendix 1 provides information about the exclosures, including age, size, experimental design and current status. furthermore, online appendix 2 provides details of the sources used during this review, including the exclosures used and the main variables measured. considering all these fenced areas as ‘herbivore exclosures’, even when some of them have enclosed selected herbivores for certain periods, stems from the fact that (1) elephant, an important landscape engineer in the park, was fenced out of structures, (2) herbivore densities in the exclosures were generally lower than areas outside the exclosures, (3) a few of them have been used both as enclosures and exclosures, and (4) fire has been excluded or carefully managed in many of the enclosures and exclosures. methods this review examines the utility of small to large (1 m2 – 870 ha) and temporary (one growing season) to long-term (≥ 15 years) herbivore exclosures in knp as research infrastructure. to gather all relevant literature, we conducted a google scholar database search using keywords including various combinations of ‘exclosure or enclosure #name’, ‘enclosure knp’ or ‘exclosure knp’, as well as the names of the sections of knp in which the exclosures are located. there are various spellings of the same exclosure name in the literature and all the spelling variations were included in the search. a single standardised spelling for each is used here. we focussed on published work; however, where work was not published, we referred to the corresponding thesis, if available. work done within exclosures was included regardless of whether the objectives were related directly to herbivory or not. research on the breeding of rare antelope was not included, as this information is poorly documented in the formal literature. much of the history and lessons learned are captured in unofficial reports, meeting minutes, correspondence and ranger diaries, but not formally curated (i.e. not listed in the sanparks literature bibliography). although this history and the associated lessons learned are important and worthy of publication, it requires a dedicated study beyond the scope of the current review. review findings a total of 75 peer-reviewed published works were retrieved during the search which ended on 31 august 2021. hlangwini, n’waswitshumbe and makhohlola exclosures eighteen published studies include the hlangwini (figure 2), makhohlola (figure 3) and n’waswitshumbe (figures 4 and 5) exclosures. these exclosures were reviewed together as they are often used as replicates in the same studies, and share more or less the same long-term history of low to no herbivore presence. although some soil research has taken place (e.g. craine, morrow & fierer 2007; craine, morrow & stock 2008; holdo & mack 2014; wigley et al. 2014), the exclosures have mainly been used to compare the effects of herbivore presence or absence on woody vegetation. complementary work from elsewhere in knp that used small temporary exclosures to investigate bottlenecks in tree recruitment is included here (midgley et al. 2020; namah, midgley & kruger 2019). in addition, some studies have investigated the consequences of excluding herbivores for other faunal groups (macfadyen et al. 2012; mccleery et al. 2018) and they, too, are included here. figure 2: a fence-line contrast of the southern edge of the hlangwini exclosure showing the relatively similar woody densities inside (right) and outside (left) the fence. figure 3: a tall woodland establishes over time (right side) when herbivores are excluded on the southern basalts in kruger national park, here shown in the makhohlola exclosure. figure 4: an aerial view of the eastern boundary fence of the n’waswitshumbe exclosure, showing higher woody density where herbivores are excluded. figure 5: the lower (eastern) corner of the n’waswitshumbe exclosure in (a) 1986 and (b) 2015 showing the thickening and expansion of woody vegetation inside of the exclosure compared to the persistent open landscape outside the exclosure. herbivore effects on woody cover airborne lidar (light detection and ranging) technology has shown that herbivores increase tree-fall rates, decreases woody cover, and alter 3-d structure of woody vegetation and canopy height (asner et al. 2009; asner & levick 2012; colgan et al. 2012; levick et al. 2009, 2010) (figures 3–5). however, the magnitude of the effects is dependent on landscape position and soils (asner et al. 2009; colgan et al. 2012). this is apparent in figure 2, where the tree cover is quite similar inside and outside of the hlangwini exclosure, situated on nutrient-poor sandy soils, compared to the very pronounced differences visible at makhohlola and n’waswitshumbe (figures 3–5), where the soils are much more fertile. herbivores impact the abundance of certain species. for instance, dalbergia melanoxylon, combretum imberbe, c. hereroense and lannea schweinfurthii were rarely found outside the exclosures. alternatively, dichrostachys cinerea and albizia harveyii are either more common outside of exclosures or increases faster outside of exclosures over time (levick & rogers 2008; wigley et al. 2014). work from elsewhere in knp using small 1 m2 exclosures showed that the recruitment bottleneck for some tree species such as kigelia africana and senegalia (acacia) nigrescens occurs via predation on seedlings (namah et al. 2019; voysey et al. 2021). even when trees succeed through the recruitment bottleneck, persistent and chronic pruning of plants by elephants and other herbivores can result in mass sterilisation and a change in the adult population demography. from work that included the n’waswitshumbe exclosure, it was shown that most african palms (hyphaene petersiana) in knp are prevented from reaching sexual maturity and thus the ability to form seeds, because of chronic elephant herbivory (midgley et al. 2020). herbivore effects on other faunal groups these exclosures have hardly been used to study the impacts of herbivores on other faunal groups; only four publications were found that addressed it. one study at n’waswitshumbe exclosure found no differences in small mammal species composition in and out of the exclosure (macfadyen et al. 2012). small mammal densities were more affected by time since the last veldfire where a drastic reduction in densities were observed post-fire. alternatively, at the makhohlola exclosure, which until 2016 supported a far greater biomass of large trees and shrubs than the surrounds, the diversity of small mammals and birds were far greater than the surrounding control plots (mccleery et al. 2018). this work, which included many additional sites, led to the conclusion that the greater density of woody plants when protected from herbivores, results in higher habitat heterogeneity, which supports greater species and functional guild diversity. the mass sterilisation of h. petersiana, mentioned before, may have cascading effects on biodiversity because many species feed on the fruits and flowers (e.g. vervet monkeys, porcupines, squirrels and wasps) and birds would nest within tall individuals (e.g. woodpeckers) (midgley et al. 2020). we suggest that these cascading effects on biodiversity may be true for any loss of plant species or reduction in vegetation structural diversity with herbivory, although the removal of herbivores may also result in the loss of dispersal opportunities for some species (midgley, gallaher & kruger 2012). nkhuhlu and letaba exclosures the nkhuhlu exclosures (full exclosure shown in figures 6, 7; partial in figure 8) underpin 26 publications; while the letaba exclosures were used in 9 articles. the nkhuhlu exclosures are situated about 25 km away from skukuza, which is knp’s administrative headquarters and includes a large research camp as well as research support facilities, which likely contributes to its popularity as a research site. the letaba exclosures on the other hand, are quite remote with an access road that is often inaccessible after rains. the main body of work that has taken place in the nkhuhlu exclosures addresses the effects of herbivores, and drought on the woody and herbaceous layers. figure 6: fence-line contrast of a sodic site between full exclosure (left) and area open to herbivores (right) at the nkhuhlu exclosures. figure 7: aerial view of the nkhuhlu exclosure showing the same sodic site as in figure 6 in the foreground with the open-access control (left) and the full exclosure (right). figure 8: aerial view of the nkhuhlu exclosures comparing the crest without elephants and giraffe (partial exclosure, left) and the control open to all herbivores (right). soil and plant nutrients thorough soil classification was done when the exclosures were erected (paterson & steenekamp 2003), but limited follow-up soil work has taken place. jacobs et al. (2007) investigated nitrogen (n) fixation in philenoptera violacea in the nkhuhlu exclosure. they found that p. violacea fixes n and this ability varies with the soil conditions and life history stage; that is, leaves of saplings and juveniles fix higher levels of atmospheric n than mature plants. although these studies utilised the exclosures as study sites, they did not compare n and phosphorus (p) limitation with and without herbivores or n fixation with and without herbivores. as part of their study on arthropod diversity in the two exclosures, thoresen et al. (2021) measured a range of soil fertility indicators (including the macro-elements, ph and soil moisture), and found that herbivore exclusion increased both soil nutrients and moisture. however, herbivores had little effect on soil bacterial and fungal communities (vermeire et al. 2021). herbivore effects on the woody layer the earliest vegetation work from the two exclosures described and mapped their plant communities (siebert & eckhardt 2008; siebert, siebert & eckhardt 2010). the woody cover generally increases with herbivore exclusion (figures 6–8) (asner & levick 2012; asner et al. 2009). these increases are accompanied by increases in woody litter and dead or senescing material, which asner et al. (2009) suggested could result in more intense fires, where areas are protected from herbivores. using lidar-derived metrics for aboveground carbon (c) pools, davies and asner (2019) showed that aboveground c increased over a 6-year period, regardless of herbivore presence, but gains were highest where herbivores were completely excluded. the woody species composition was slow to react to the exclusion of herbivores at nkhuhlu, with no change detected after 5 years (scogings et al. 2012). however, some changes in species abundance were seen after 13 years (combrink 2016). similar to what was observed at the n’waswitshumbe and makhohlola exclosures, combrink (2016) and scogings (2011) also found that d. cinerea’s abundance increased and growth rates were faster when not protected from herbivores. although herbivores may negatively impact woody vegetation survival, severe droughts can be more acute in the short-term. during the 2015–2016 drought, case et al. (2019) investigated the role of herbivores on tree mortality at nkhuhlu, and concluded that drought impacts outweighed herbivore effects on tree mortality during below-average rainfall periods. mortality, however, was most pronounced in the partial exclosure where only elephants and giraffes were excluded, than in the full exclosure. this was in contrast to the average rainfall years where mortality probability was highest in the area open to herbivores, more so than in the two fenced areas (scogings 2014). it was suggested that this may reflect a combination of meso-herbivore displacement (e.g. impala preferring the partial exclosure to avoid elephants) and herbivore diet-switching; for example work elsewhere showed that mixed feeding impalas utilised a higher proportion of trees during drought than during average rainfall years (abraham, hempson & staver 2019). work from knp exclosures, including nkhuhlu, found little consistent responses of c-based secondary metabolites (cbsms) to herbivore exclusion (scogings, hjältén & skarpe 2011, 2013; scogings et al. 2014, 2015; wigley et al. 2014). some species’ cbsms were the highest (e.g. c. apiculatum) with herbivore absence, but responses were species-dependent (scogings et al. 2013, 2014). for trees in general, the time since first rainfall and season seem to be more reliable predictors of nutrient and tannin concentrations, than the presence of mammal herbivores (scogings, hjältén, et al. 2013; scogings et al. 2013, 2014). in contrast to cbsms, the presence of herbivores has more consistent effects on structural defences. intense browsing results in increased shoot growth and branching, and increased spine density and length (engdahl 2009; scogings et al. 2013; wigley et al. 2014). herbivore effects on the herbaceous layer the utilisation of sodic patches is very high when accessible to herbivores at the nkhuhlu exclosure (figure 6), resulting in higher n and sodium (na) and higher n cycling (alard 2009). compared to sodic patches, crest patches are relatively water and n limited environments that result in less nutritious grass forage, which is generally less utilised. nutrient-rich, sodic sites seem to be resilient in the face of high utilisation, with exclusion of herbivores decreasing herbaceous species richness as well as functional diversity (jacobs & naiman 2008; van coller & siebert 2015, 2020; van coller, siebert & siebert 2013). work from nkhuhlu exclosure has highlighted the important contribution that forbs make to herbaceous species richness and diversity (e.g. ~80% of herbaceous species richness) (siebert & scogings 2015; van coller & siebert 2020), as well as functional richness with many palatable forbs being heavily utilised by herbivores (siebert & scogings 2015). herbivore effects on other faunal groups generally, mammalian herbivores had negative effects on the total invertebrate abundance at the two exclosures (jonsson et al. 2010; thoresen et al. 2021), but their presence also resulted in novel species being present (haba, herbert & yang 2019). depending on landscape position, some groups’ abundance (e.g. spiders in riparian areas) increased the most when meso-herbivores were present (i.e. partial exclosure). while arthropod abundance decreased with herbivores presence, its diversity increased (thoresen et al. 2021). these patterns were related to the indirect effects of herbivores through altering interactions between vegetation, fire, microclimate, and soil nutrients. at nkhuhlu exclosure, haba et al. (2019) found an interaction between herbivore presence, s. nigrescens, ants and scale insects. ants were present with and without herbivores, but tachardina africana, a scale insect, was only found on s. nigrescens trees where herbivore was present. disease work in the satara buffalo exclosure specific-pathogen-free buffalo breeding project the satara (buffalo) exclosure was erected to breed specific-pathogen-free buffalo that could be translocated out of the foot and mouth disease (fmd) endemic area in knp, and into the fmd-free zone of south africa. this involved setting up founder herds of tuberculosis (btb)-negative and brucellosis-negative animals caught in northern knp and transferring them to the satara (buffalo) exclosure, to produce calves that would go through a series of quarantine procedures after weaning. the project produced over 400 btb-, brucellosis-, fmdand corridor disease-free calves that were used to restock addo elephant and mokala national parks in south africa and gorongosa national park in mozambique (j. malan [sanparks], pers. comm., 17 september 2021). oregon state university disease ecology programme from 2015 to 2017, the remaining animals in the satara exclosure were used in an oregon state university (osu) disease ecology research programme that produced 8 publications and 10 postgraduate qualifications. the unique, longitudinal nature of the osu study allowed for several diseases to be tracked over time, in individual animals and through the population, revealing temporal patterns not discoverable through typical cross-sectional studies. the satara exclosure enabled this research, since studying a high number of buffalo, subject to environmental and seasonal disease dynamics, at bi-monthly intervals over 3 years, would otherwise not have been possible. the study was unique in that it used a large social animal in a natural setting; most other epidemiology studies of this kind are laboratory-based. the work helped establish that buffalo could serve as an ideal wild model system, linking individual immunology, environmental change (linked to metabiome health), and parasite transmission dynamics (combrink et al. 2020; couch et al. 2017, 2021; ezenwa et al. 2019; glidden et al. 2017, 2020; jolles & ezenwa 2015; sisson et al. 2017). tree grass programme the uct tree grass programme was established in knp in 2005 and operated for 10 years. the two exclosures (uct shabeni and uct satara) were part of a broader programme which delivered more than 30 publications and had > 20 postgraduate students involved (e. february [uct], pers. comm., 03 september 2021). the main objective of the programme was to investigate tree-grass coexistence in terms of two dominant hypotheses, namely, resource-based and disturbance-based. much of the research that took place in the exclosures addressed the issues around allocation of resources between trees and grasses (february & higgins 2010; february et al. 2013; higgins, keretetse & february 2015; february, pausch & higgins 2020). the results show a high degree of overlap of tree and grass roots, which gives rise to high potential competition (february et al. 2013). trees had increased growth rates when grasses were removed, but not when they received extra water. however, grass biomass was not influenced by the presence of trees, but was significantly and positively influenced by total precipitation increases, thus suggesting that increasing rainfall facilitates grass competition with trees. later work by other researchers found that although grasses, saplings and trees used water from the shallower depths when water is not limiting, saplings and trees shifted water uptake to deeper depths when resources are scarce (kulmatiski & beard 2013a). also, small experimental increases in rainfall intensity, without changing the total amount, benefits trees as water is pushed deeper into the soil (kulmatiski & beard 2013). if precipitation intensity increases in the future, deep soil water will increase, which may benefit shrubs more than grasses, with associated increases in woody encroachment as a result (berry & kulmatiski 2017; kulmatiski & beard 2013b). smaller-scale exclosures around satara during 2005–2006, a usa-sa consortium of researchers initiated a programme with the main objective to broadly test whether general rules can be formulated in terms of drivers, responses and functioning, across grassy ecosystems in north america and south africa (knapp et al. 2004, 2006). the group established 38.5 m2 exclosures on (1) the annual, triennial and unburned blocks of the satara, n’wanetsi and marheya ebp strings (figure 9), (2) annual and no-burn blocks inside the satara (buffalo) exclosure, and (3) at two sites off the ebps at midand foot-slope catenal positions. in addition, from 2006 to 2008, 1 m2 exclosures were erected on fire treatments to quantify aboveground net primary productivity (anpp) (figure 9). the focus of the work evolved from comparing systems across continents to focusing on the single and combined effects of fire, soil fertility (landscape position) and herbivory in african savannas. this research infrastructure has produced 25 publications and supported 17 graduate students and 8 postdoctoral researchers. figure 9: full (left) and partial (right) 7 m2 exclosures used by the usa-sa consortium on the satara and n’wanetsi experimental burn plots and affiliated high-low productivity sites. a smaller 1 m2 exclosure for quantifying anpp is visible in the middle. additionally, in 2013, a research group from the universities of the witwatersrand (wits) and liverpool (hereafter the fire-grazer research programme) established an experiment north of the n’wanetsi ebp string to test the influence of fire season and fire size on grazers, and the effect of post-burn herbivory on the grass layer (donaldson et al. 2018). the programme has produced nine publications, and supported seven postgraduate students and two postdoctoral researchers (s. archibald [wits], pers. comm., 21 september 2021). for the experiment, 1 m2 exclosures were erected on (1) four replicates each of three fire treatments (unburned, early-burns and late-burns), two fire sizes (0.25 and 5 ha), and three 25 ha late-burn plots, and (2) lawn areas where the grass was generally below 5 cm tall, tussock (bunch) grass areas where the grass was taller than 30 cm and fires were excluded for 3 years, and communities burned at the beginning of the experiment; that is, the end of dry season in 2015. the research of both the usa-sa consortium (now in its 16th year of herbivore exclusion) and the fire-grazer research programme (8th year of fire manipulation in 2021), is ongoing. grazer and fire effects on productivity the initial research investigated the drivers of annual herbaceous anpp, using the exclosures on the ebps and in the satara (buffalo) exclosure. grass and forb anpp were unaffected by fire regime or whether recent grazing was by a single species (i.e. buffalo) or a mixed suite of species (buis et al. 2009). furthermore, knapp et al. (2012) tested whether overcompensation (i.e. higher anpp when grazed) took place and showed that regardless of whether areas were burned or not, and regardless of whether herbivory took place by a single species or a diverse suite of species, the total anpp was most often fully compensated for but not overcompensated. undercompensated grass anpp during drier years may be off-set by increased forb anpp. the research which took place during drought (either 2015–2016 or experimentally induced) supported emerging patterns elsewhere that show that the herbaceous layer of grasslands and savannas have low resistance but high resilience to drought (koerner & collins 2014; donaldson et al. 2019; wilcox et al. 2020).working in full exclosures on the three different fire regimes of the ebps immediately after the 2015–2016 drought, wilcox et al. (2020) found that the total anpp was very resilient to drought and normalised 1 year post-drought when herbivores were excluded. however, responses are affected by the interaction between herbivory and fire. in the absence of herbivores, tussock grasses are generally able to withstand severe drought and maintain cover, and when burned, there is a significant increase in productivity when rainfall returns (donaldson et al. 2019). unburned, non-grazed tussock grasses respond poorly to rain. with intense herbivory, however, burned tussock grasses had significant mortality during the drought. a low severity 3-year drought was experimentally imposed by koerner and collins (2014) at plots in an annually burned block of the satara (buffalo) exclosure, where herbivory was also excluded for the duration of the experiment, as well as in the uct satara exclosure, where herbivory and fire had been excluded since 2002. twice a year herbivory was simulated by clipping grasses, but not forbs,. with infrequent burning and a history of grazing exclusion (uct satara), grass cover and total anpp were decreased by clipping (i.e. grazing) and drought, as well as by a clipping and drought interaction. with a history of disturbance (i.e. annual burning and low intensity grazing by a single herbivore), there was no effect of clipping or drought on grass cover or total anpp. in summation, the productivity of the herbaceous layer is resilient in the face of drought, however, the grazing and burning history as well as current precipitation of the area govern responses. grazer and fire effects on species richness similar to the nkhuhlu exclosures, the exclusion of herbivores in general decreases herbaceous plant diversity (burkepile et al. 2017; burns, collins & smith 2009). eby et al. (2014), working inside the satara (buffalo) exclosure, compared plots that excluded buffalo for 7 years with buffalo present, in areas that burned annually and where fire was excluded for 10 years. the removal of buffalo resulted in decreased herbaceous richness and diversity when areas were burned annually, but not when areas were not burned. the increased dominance of one of themeda triandra, panicum coloratum or digitaria eriantha, in annually burned areas with herbivore exclusion, resulted in decreases in richness and diversity (figure 10). figure 10: removing mammalian herbivory drives a change in species composition and the accumulation of biomass in the herbaceous layer inside exclosures. although species richness and diversity decreased with herbivore removal during some years on the infrequently burned and unburned areas, there was no definitive change in species or functional community structure after 7 years of herbivore removal at any fire frequency (forrestel, donoghue & smith 2015; koerner et al. 2014). a follow-up study compared species richness and dominance when different sized herbivores were excluded with either full or partial exclosures (figure 11) (burkepile et al. 2016). on more frequent burns, exclusion of only larger herbivores (e.g. elephants, zebra [equus quagga burchellii], wildebeest [connochaetus taurinus]) increased plant abundance and total cover but not plant diversity, while also excluding smaller species (e.g. impala, warthog [phacochoerus africanus], steenbok) resulted in modest and transient declines in plant diversity. on unburned areas, herbivore exclusion led to increased dominance and shading, which resulted in declines in plant richness and diversity. figure 11: where smaller-bodied herbivores, primarily the numerically dominant impala, are excluded (a, full exclosure), there is substantial increase in herbaceous biomass compared to where they have access to grazing (b, partial exclosure). this is apparent even in early season growth. furthermore, the effect of herbivore exclusion on species richness was shown to depend on site productivity (catenal position) and herbivore size (burkepile et al. 2017). the exclusion of all herbivores generally increased plant species richness (i.e. forbs and grasses combined) at low productivity, but decreased richness at high productivity (figure 12). the exclusion effects of only smaller herbivores showed declines in species richness across the productivity gradient. grass species richness declined regardless of the type of herbivore exclusion or level of productivity, while forb richness depended on productivity and herbivore size, with the largest decreases as a result of the exclusion of smaller herbivores at high productivity. figure 12: exclosures (left) clearly show the impact of herbivory on this productive, heavily-grazed foot-slope even in the first year after establishment. herbivore habitat preferences both the usa-sa consortium and the fire-grazer research programme showed that grazers respond positively to areas burned recently, and also prefer more frequently burned areas in general (burkepile et al. 2013; donaldson et al. 2018). as a result, fires can be used to create intensely used patches (donaldson et al. 2018). fires less than 25 ha attracted sufficient grazers to shorten grass height, but only for one growing season. the repetition of late burns (i.e. september/october) for an additional 2 years resulted in the active selection of these areas by wildebeest and impala, and subsequently, grazing pressure was high enough to initiate positive feedbacks and maintain lawns that excluded repeat fires. when grazing was excluded, grasses quickly recovered to pre-fire levels by the end of the season. herbivore effects on grass nutrition the results from research that included the satara (buffalo) exclosure as a single grazer site with less intense utilisation, found that grasses are higher in n and p in areas with a combination of wildlife species present, as opposed to areas with only a single species present (baumgartner et al. 2015; treydte et al. 2013). these results reflect in part a switch in the dominant grass species when a diverse suite of wildlife species is present. urochloa mosambicensis, for instance, which was common in the mixed wildlife sites, is higher in n and p when compared to d. eriantha, which was common in the single species sites. however, the same grass species were also more nutrient rich in the mixed wildlife sites, as opposed to the single herbivore species site, which may reflect a positive feedback between being frequently grazed and increased nutrient content. synthesis within the context of a large protected area such as knp, the herbivore exclusion sites enable intensive study of one of the main drivers in disturbance-driven savanna systems, that is herbivory by mammalian herbivores. they allow the partitioning of effects of those disturbances that managers can partly control (e.g. fire regimes, herbivore densities and disease management) from those that cannot be manipulated easily (e.g. droughts, floods and atmospheric co2 concentrations). the understanding and knowledge that result from this research inform management decisions and policies, allowing sanparks to learn and adapt in an ever-changing environment. realising the value of exclosures as research infrastructure in knp has been varied. the largest exclosures, which also have the longest histories, have been relatively underutilised as research sites (e.g. hlangwini and n’waswitshumbe). this may be partly because of the lack of replicated treatments and the problem of pseudoreplication (holdo & mack 2014), the relatively remote location of these exclosures, and the lack of dedicated research funding within sanparks. the more successful exclosures, from a research output perspective, are those that have been erected through fully-funded research programmes that include dedicated personnel and bursaries for students, with specific objectives in mind, and where logistics such as distance to laboratories and research accommodation have been taken into account. the two areas of savanna ecology in which a significant body of work has been produced, across multiple exclosures in knp, are the woody layer and herbaceous layer dynamics. most of the woody work was done in the nkhuhlu exclosure, with additional work from the hlangwini, n’waswitshumbe and makhohlola exclosures. multiple studies from the nkhuhlu exclosure and the satara-based programmes focused on the interactions between herbivores and fire on the herbaceous layer, with some additional work addressing potential competition between trees and grasses. woody layer it is clear that herbivory decreases woody cover and simplifies vegetation structure (table 1, figures 3–8). the effects on woody species richness is less clear; unique species occur with or without herbivory, but the most pronounced effects of herbivores are changing abundances, rather than creating unique woody communities. for instance, certain species thrive when faced with disturbance by herbivores (and fire) and decrease when protected from herbivory. the knp subscribes to the assumption that increased heterogeneity (e.g. a range of woody cover states ranging from open to dense areas) will increase resilience and biodiversity (see rogers 2003; smit & prins 2015). more open savanna is often more productive in terms of herbaceous productivity and the herbivore assemblage may change along a woody cover gradient (smit & prins 2015). woodlands of intermediate densities harbour high faunal biodiversity, but this may be reversed at high levels of woody densities (mccleery et al. 2018; stanton et al. 2018). the implication is that a range of herbivore densities should ensure a range of woody densities, which in turn will maximise biodiversity. table 1: woody layer responses to herbivore exclusion in the kruger national park. table 2: herbaceous layer responses to herbivore exclusion in the kruger national park. herbaceous layer herbaceous productivity is mostly maintained with high herbivory and frequent fire, and is only impacted during extreme droughts, but recovers rapidly post-drought (buis et al. 2009; knapp et al. 2012). species diversity also depends on the presence of herbivory and often decreases when herbivores are excluded (burkepile et al. 2016; eby et al. 2014; van coller et al. 2013, 2018). there seems to be an intermediate level of herbaceous biomass that ensures highest herbaceous species richness and diversity, and which is maintained by herbivore presence (van coller & siebert 2015). however, grazing and fire have very important interactive effects on herbaceous dynamics; herbaceous richness is the highest in the absence of fire and declined more with herbivore exclusion than when burned (smith et al. 2016a). the initial dominance of b. radicans compared to more palatable grasses often governs responses to herbivore removal (eby et al. 2014; forrestel et al. 2015). low levels of herbivory (such as in satara (buffalo) exclosure) or herbivory exclusion (such as uct satara) give rise to herbaceous layers dominated by tall, palatable grasses (figure 10). high levels of herbivory and fire on the other hand give rise to the dominance of bothriochloa radicans. when palatable grasses are dominant, the removal of herbivory results in palatable grasses being stimulated when burned, and the increase in the dominance of these in turn result in lower species richness (eby et al. 2014). on the ebps however, b. radicans is dominant in the more frequently burned plots. as b. radicans is mostly avoided by herbivores, herbivore exclusion results in little overall change in species richness because b. radicans remains the dominant species. even though it is difficult to decrease the dominance of b. radicans through shorter-term fire or herbivore exclusion, b. radicans was greatly reduced during the recent drought, with drought-recovery seeing an increase in palatable grasses (swemmer et al. 2018; wigley-coetsee & staver 2020). apart from an interaction between fire regime, grazer exclusion and herbaceous dynamics, effects of herbivore exclusion depend on the types and sizes of animals and their interactions with different fire regimes (burkepile et al. 2016), herbivore densities (burkepile et al. 2017; treydte et al. 2013), and site productivity (burkepile et al. 2017). another important consideration is the balance of grazer, mixed feeder and browsing herbivores (burkepile et al. 2017; burns et al. 2009; treydte et al. 2013). in systems where the removal of herbivores results in pronounced directional loss of herbaceous species richness over time, the herbivores often belong to one type, for example only grazers (eby et al. 2014; smith et al. 2016b). the removal of smaller herbivores, which include the mixed-feeding impala, has the most conspicuous impacts; for example lowered species richness and diversity, increased plant dominance and increased plant litter (burkepile et al. 2016, 2017). this may be a result of both their numerical dominance in knp as well as their considerable utilisation of the forb layer. a review of herbivore exclusion effects on the herbaceous as well as woody layers across african savannas (which included data from the knp exclosures) also showed that herbivore impacts are dominated by small-bodied mixed feeders, which was mostly because of their high population densities (staver et al. 2021). lastly, little work has compared mammalian herbivory with invertebrate herbivory impacts. the only study on this topic compared termite herbivory when mammals were excluded, and found that although both types of herbivory were more intense on termite mounds than off, termite herbivory removed more biomass than mammal herbivores during the wet season (davies et al. 2016). given the experimental design and obvious effects of herbivory on savanna systems, exclosures have provided excellent learning opportunities for students across the field of education. we have highlighted the number of graduate students who were involved in exclosure research in knp. yet, these exclosures have also played a major role in undergraduate training. for example, the organization of tropical studies (ots) has been working in knp since 2004 and has conducted student projects in multiple exclosures across the park. to date about 35 undergraduate student research projects from ots have used sanparks exclosures (unpublished data; ots) to understand the effect of mega-herbivores on woody plant demography, changes in community composition of woody plants and grasses, the consequences of mutualism breakdown for plant demography, the effect of invertebrate versus mammalian herbivory on termite mounds, phenotypic variation in physical defences of woody plants and the influence of vegetation structure on invertebrate (butterflies and dung beetles) and small vertebrate communities. many of these smaller research projects served as pilot projects for graduate studies and some of these have been published (e.g. haba et al. 2019; midgley et al. 2020; namah et al. 2019). multiple local and foreign universities, including uct, wits, north west university, bayreuth (germany), university of florida (usa), and western sydney university (australia), currently utilise exclosures as part of their field-based experiential training. importantly, the exclosures create opportunities that would not otherwise be available. for instance, studying disease ecology and eco-immunology in large ungulates in a natural system with a diversity of large dangerous animals is both expensive and logistically challenging. the satara (buffalo) exclosure created a tractable and unique, longitudinal study system which has allowed training in a fairly safe environment. implications and recommendations for establishing exclosures the lessons learned from setting up and managing exclosures, and increasing the research uptake thereof, include the following: give consideration to logistics such as ease of access, including distance from research camps and presence of all-weather access roads in selecting exclosure sites. ensure a budget and maintenance plan that stretches beyond the establishment of the exclosure, and which is linked to the time frame for which the exclosure is needed (e.g. 20 years). build collaborations and joint ownership of exclosures with land managers during the planning phase. purposefully design the exclosure experiment with research questions in mind, for example variable fire treatments, across catenal positions, allowing selective exclusion of different-size herbivores. adapt the size of the exclosures to fit the research questions asked; for example the usa-sa consortium exclosures have been very productive in terms of research outputs. although these exclosures cover a relatively small area, they are numerous and large enough to capture dynamic herbaceous layer changes. employing ‘clever’ design may save much money in the long-term; for instance, electrified 2or 3-strand wire exclosures with electrified wires positioned horizontally up to 1 m, such as those used at mpala in kenya and grumeti in tanzania (a. uys [maritaba], pers. comm. 20 september 2021), are cost effective and extremely efficient as a barrier against megafauna movement. full exclosures inside partial exclosures allow for some level of protection of the full exclosure against elephant damage and can decrease overall costs of maintenance. canvas the broader research community to solicit baseline information which will add value to the exclosure and stimulate further research (e.g. floristic composition, pre-treatment conditions, aerial photographic and lidar surveys, etc.). make baseline information available as a ‘starter pack’ for researchers and students – this can include information pertaining to the year of establishment, size, spatially-explicit treatment history (e.g. fire treatments), long-term average and yearly rainfall, estimates of herbivore density outside exclosures, and so on. collaboration beyond the initial group and objectives have made exclosures such as the usa-sa consortium exclosures very productive and maintains high energy and interest regarding the exclosures. prepare a closure or hand-over plan for the exclosure, which may include a review of research (i.e. knowledge generated, publications, students graduated, etc.), and a long-term maintenance plan including costing. do not assume that the land manager or owner will be in the position to maintain ageing infrastructure beyond the lifespan of the programme. conclusion fence-line contrasts between exclosures and the surrounding landscape are very effective in demonstrating the significant effects and interactions of herbivores in savanna ecosystems to both managers and the public alike, and for stimulating new research ideas for scientists. in addition, exclosures are excellent outdoor classrooms and laboratories for training the next generation of ecologists. with a renewed interest in the implications of the mass extinction of mammal herbivores (e.g. rowan & faith 2019) and ‘rewilding’ as an ecological restorative mechanism in response, exclosures are one of the critical tools to explore these concepts (see e.g. two recent special issues on mega-herbivore extinction and rewilding by malhi et al. 2016; smith et al. 2016a). the loss of exclosure infrastructure in knp represents a missed opportunity, in that it may form a global research platform similar to the ebps. this fire experiment has been running uninterrupted for seven decades and have become one of the primary sites for studying fire effects globally (e.g. pellegrini et al. 2021). these types of long-term research infrastructure and associated data that outlive the typical 3–5 year funding and project cycles are some of the main reasons researchers choose to conduct research in knp. as a result, knp is the premier savanna research destination in africa (smit et al. 2017). lastly, some of the long-term exclosure work from knp reminds us not only how resilient savannas are in the face of disturbance, but also how they are shaped by disturbances, realising outcomes which are difficult to achieve by management alone. acknowledgements the authors would like to thank all the south african national parks (sanparks) staff including section rangers, research assistants and technicians who have worked so hard for many years to service, maintain, and survey exclosures in the knp. the authors also wish to thank dr. anthony swemmer and south african environment observation network (saeon) for all the support, both logistically and financially, in maintaining and surveying some of the exclosures, especially the letaba exclosure. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions c.w-c., t.s., d.g., d.i.t., l.k. and i.p.s. contributed to the text; c.w-c., t.s., c.g., n.g., d.g. and i.p.s. conceptualised the work; j.b., a.m., l.k., and j.v. provided stats on exclosure maintenance; c.w-c. and t.s. analysed literature and data and c.s. provided the map. ethical considerations this article followed all ethical standards for research without direct contact with human or animal subjects. funding information this research received no specific grant from any funding agency in the public, commercial or not-for-profit sectors. data availability all data, in this case works of literature reviewed have been summarised in online appendix 2. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references abraham, j.o., hempson, g.p. & staver, a.c., 2019, ‘drought-response strategies of savanna herbivores’, ecology and evolution 9(12), 7074–7056. https://doi.org/10.1002/ece3.5270 alard, g.f., 2009, ‘a study of selected grass ecophysiological characteristics as well as a comparison of the quality and quantity of grass in sodic and crest patches within a semi-arid granitic savanna catena in the southern knp, south africa’, phd thesis, university of the witwatersrand, johannesburg. asner, g.p. & levick, s.r., 2012, ‘landscape-scale effects of herbivores on treefall in african savannas’, ecology letters 15(11), 1211–1217. https://doi.org/10.1111/j.1461-0248.2012.01842.x asner, g.p., levick, s.r., kennedy-bowdoin, t., knapp, d.e., emerson, r., jacobson, j. et al., 2009, ‘large-scale impacts of herbivores on the structural diversity of african savannas’, proceedings of the national academy of sciences 106(12), 4947–4952. https://doi.org/10.1073/pnas.0810637106 baumgartner, s.a., treydte, a.c., grant, c.c. & van rooyen, j., 2015, ‘can diverse herbivore communities increase landscape heterogeneity? comparing wild and domestic herbivore assemblages in a south african savanna’, perspectives in plant ecology, evolution and systematics 17(1), 34–43. https://doi.org/10.1016/j.ppees.2014.11.002 berry, r.s. & kulmatiski, a., 2017, ‘a savanna response to precipitation intensity’, plos one 12(4), e0175402. https://doi.org/10.1371/journal.pone.0175402 biggs, r., biggs, h.c., dunne, t.t., govender, n. & potgieter, a.l.f., 2003, ‘experimental burn plot trial in the kruger national park: history, experimental design and suggestions for data analysis’, koedoe 46(1), 1–15. https://doi.org/10.4102/koedoe.v46i1.35 buis, g.m., blair, j.m., burkepile, d.e., burns, c.e., chamberlain, a.j., chapman, p.l. et al., 2009, ‘controls of aboveground net primary production in mesic savanna grasslands: an inter-hemispheric comparison’, ecosystems 12(6), 982–995. https://doi.org/10.1007/s10021-009-9273-1 burkepile, d.e., burns, c.e., tambling, c.j., amendola, e., buis, g.m., govender, n. et al., 2013, ‘habitat selection by large herbivores in a southern african savanna: the relative roles of bottom-up and top-down forces’, ecosphere 4(11), 1–19. https://doi.org/10.1890/es13-00078.1 burkepile, d.e., fynn, r.w., thompson, d.i., lemoine, n.p., koerner, s.e., eby, s. et al., 2017, ‘herbivore size matters for productivity–richness relationships in african savannas’, journal of ecology 105(3), 674–686. https://doi.org/10.1111/1365-2745.12714 burkepile, d.e., thompson, d.i., fynn, r.w., koerner, s.e., eby, s., govender, n. et al., 2016, ‘fire frequency drives habitat selection by a diverse herbivore guild impacting top–down control of plant communities in an african savanna’, oikos 125(11), 1636–1646. https://doi.org/10.1111/oik.02987 burns, c.e., collins, s.l. & smith, m.d., 2009, ‘plant community response to loss of large herbivores: comparing consequences in a south african and a north american grassland’, biodiversity and conservation 18(9), 2327–2342. https://doi.org/10.1007/s10531-009-9590-x case, m.f., wigley-coetsee, c., nzima, n., scogings, p.f. & staver, a.c., 2019, ‘severe drought limits trees in a semi-arid savanna’, ecology 100(11), e02842. https://doi.org/10.1002/ecy.2842 colgan, m.s., asner, g.p., levick, s.r., martin, r.e. & chadwick, o.a., 2012, ‘topo-edaphic controls over woody plant biomass in south african savannas’, biogeosciences 9(5), 1809–1821. https://doi.org/10.5194/bg-9-1809-2012 combrink, l., 2016, ‘savanna woody regeneration in response to different treatments of herbivory and fire’, phd thesis, north-west university (south africa), potchefstroom campus. combrink, l., glidden, c.k., beechler, b.r., charleston, b., koehler, a.v., sisson, d. et al., 2020, ‘age of first infection across a range of parasite taxa in a wild mammalian population’, biology letters 16(2), 20190811. https://doi.org/10.1098/rsbl.2019.0811 couch, c.e., movius, m.a., jolles, a.e., gorman, m.e., rigas, j.d. & beechler, b.r., 2017, ‘serum biochemistry panels in african buffalo: defining reference intervals and assessing variability across season, age and sex’, plos one 12(5), e0176830. https://doi.org/10.1371/journal.pone.0176830 couch, c.e., stagaman, k., spaan, r.s., combrink, h.j., sharpton, t.j., beechler, b.r. et al., 2021, ‘diet and gut microbiome enterotype are associated at the population level in african buffalo’, nature communications 12(1), 1–11. https://doi.org/10.1038/s41467-021-22510-8 craine, j.m., morrow, c. & fierer, n., 2007, ‘microbial nitrogen limitation increases decomposition’, ecology 88(8), 2105–2113. https://doi.org/10.1890/06-1847.1 craine, j.m., morrow, c. & stock, w.d., 2008, ‘nutrient concentration ratios and co-limitation in south african grasslands’, new phytologist 179(3), 829–836. https://doi.org/10.1111/j.1469-8137.2008.02513.x davies, a.b. & asner, g.p., 2019, ‘elephants limit aboveground carbon gains in african savannas’, global change biology 25(4), 1368–1382. https://doi.org/10.1111/gcb.14585 davies, a.b., van rensburg, b.j., robertson, m.p., levick, s.r., asner, g.p. & parr, c.l., 2016, ‘seasonal variation in the relative dominance of herbivore guilds in an african savanna’, ecology 97(6), 1618–1624. https://doi.org/10.1111/gcb.14585 donaldson, j.e., archibald, s., govender, n., pollard, d., luhdo, z. & parr, c.l., 2018, ‘ecological engineering through fire-herbivory feedbacks drives the formation of savanna grazing lawns’, journal of applied ecology 55(1), 225–235. https://doi.org/10.1111/1365-2664.12956 donaldson, j.e., parr, c.l., mangena, e.h. & archibald, s., 2019, ‘droughts decouple african savanna grazers from their preferred forage with consequences for grassland productivity’, ecosystems, 23, 689–701. https://doi.org/10.1007/s10021-019-00438-x eby, s., burkepile, d.e., fynn, r.w., burns, c.e., govender, n., hagenah, n. et al., 2014, ‘loss of a large grazer impacts savanna grassland plant communities similarly in north america and south africa’, oecologia 175(1), 293–303. https://doi.org/10.1007/s00442-014-2895-9 engdahl, f., 2009, ‘herbivory on woody plants and induced responses in two similar species of acacia in the kruger national park, south africa’, phd thesis, wildlife, fish, and environmental studies, swedish university of agricultural sciences. ezenwa, v.o., jolles, a.e., beechler, b.r., budischak, s.a. & gorsich, e.e., 2019, ‘the causes and consequences of parasite interactions: african buffalo as a case study’, in k. wilson, a. fenton & d. tompkins (eds.), wildlife disease ecology: linking theory to data and application, pp. 129–160, cambridge university press, cambridge. february, e., pausch, j. & higgins, s.i., 2020, ‘major contribution of grass roots to soil carbon pools and co 2 fluxes in a mesic savanna’, plant and soil 454, 207–215. https://doi.org/10.1007/s11104-020-04649-3 february, e.c. & higgins, s.i., 2010, ‘the distribution of tree and grass roots in savannas in relation to soil nitrogen and water’, south african journal of botany 76(3), 517–523. https://doi.org/10.1016/j.sajb.2010.04.001 february, e.c., higgins, s.i., bond, w.j. & swemmer, l., 2013, ‘influence of competition and rainfall manipulation on the growth responses of savanna trees and grasses’, ecology 94(5), 1155–1164. https://doi.org/10.1890/12-0540.1 forrestel, e.j., donoghue, m.j. & smith, m.d., 2015, ‘functional differences between dominant grasses drive divergent responses to large herbivore loss in mesic savanna grasslands of north america and south africa’, journal of ecology 103(3), 714–724. https://doi.org/10.1111/1365-2745.12376 glidden, c., beechler, b., buss, p.e., charleston, b., de klerk-lorist, l.-m., maree, f.f. et al., 2017, ‘detection of pathogen exposure in african buffalo using non-specific markers of inflammation’, frontiers in immunology 8, 1944. https://doi.org/10.3389/fimmu.2017.01944 glidden, c.k., koehler, a.v., hall, r.s., saeed, m.a., coppo, m., beechler, b.r. et al., 2020, ‘elucidating cryptic dynamics of theileria communities in african buffalo using a high-throughput sequencing informatics approach’, ecology and evolution 10(1), 70–80. https://doi.org/10.1002/ece3.5758 grant, c.c., bengis, r., balfour, d., peel, m.,davies-mostert, w., killian, h.,little, r., smit, i., garai, m. & henley, m., 2008, ‘controlling the distribution of elephants’, in r.j scholes & k.g mennell (eds.), elephant management: a scientific assessment for south africa, pp. 329–369, wits university press, johannesburg. haba, y., herbert, r. & yang, l., 2019, ‘an undescribed relationship between the knobthorn tree (senegalia nigrescens), ants, and a lac scale insect in south africa’, african journal of ecology 57(4), 595–598. https://doi.org/10.1111/aje.12627 higgins, s.i., keretetse, m. & february, e.c., 2015, ‘feedback of trees on nitrogen mineralization to restrict the advance of trees in c4 savannahs’, biology letters 11(8), 20150572. https://doi.org/10.1098/rsbl.2015.0572 holdo, r.m. & mack, m.c., 2014, ‘functional attributes of savanna soils: contrasting effects of tree canopies and herbivores on bulk density, nutrients and moisture dynamics’, journal of ecology 102(5), 1171–1182. https://doi.org/10.1111/1365-2745.12290 jacobs, s.m. & naiman, r.j., 2008, ‘large african herbivores decrease herbaceous plant biomass while increasing plant species richness in a semi-arid savanna toposequence’, journal of arid environments 72(6), 891–903. https://doi.org/10.1016/j.jaridenv.2007.11.015 jacobs, s.m., pettit, n.e. & naiman, r.j., 2007, ‘nitrogen fixation by the savanna tree philenoptera violacea (klotzsch) schrire (apple leaf) of different ages in a semi-arid riparian landscape’, south african journal of botany 73(1), 163–167. https://doi.org/10.1016/j.sajb.2006.09.001 jolles, a.e. & ezenwa, v.o., 2015, ‘ungulates as model systems for the study of disease processes in natural populations’, journal of mammalogy 96(1), 4–15. https://doi.org/10.1093/jmammal/gyu007 jonsson, m., bell, d., hjältén, j., rooke, t. & scogings, p.f., 2010, ‘do mammalian herbivores influence invertebrate communities via changes in the vegetation? results from a preliminary survey in kruger national park, south africa’, african journal of range & forage science 27(1), 39–44. https://doi.org/10.2989/10220111003703468 joubert, s.c.j., 1986, ‘masterplan for the management of kruger national park’, vol 1–6. unpublished document, skukuza library archives, kruger national park, south africa kioko, j., muruthi, p., omondi, p. & chiyo, p.i., 2008, ‘the performance of electric fences as elephant barriers in amboseli, kenya’, south african journal of wildlife research 38(1), 52–58. https://doi.org/10.3957/0379-4369-38.1.52 knapp, a.k., burns, c.e., fynn, r.w., kirkman, k.p., morris, c.d. & smith, m.d., 2006, ‘convergence and contingency in production–precipitation relationships in north american and south african c 4 grasslands’, oecologia 149(3), 456–464. https://doi.org/10.1007/s00442-006-0468-2 knapp, a.k., hoover, d.l., blair, j.m., buis, g., burkepile, d.e., chamberlain, a. et al., 2012, ‘a test of two mechanisms proposed to optimize grassland aboveground primary productivity in response to grazing’, journal of plant ecology 5(4), 357–365. https://doi.org/10.1093/jpe/rts020 knapp, a.k., smith, m.d., collins, s.l., zambatis, n., peel, m., emery, s. et al., 2004, ‘generality in ecology: testing north american grassland rules in south african savannas’, frontiers in ecology and the environment 2(9), 483–491. https://doi.org/10.1890/1540-9295(2004)002[0483:gietna]2.0.co;2 koerner, s.e., burkepile, d.e., fynn, r.w., burns, c.e., eby, s., govender, n. et al., 2014, ‘plant community response to loss of large herbivores differs between north american and south african savanna grasslands’, ecology 95(4), 808–816. https://doi.org/10.1890/13-1828.1 koerner, s.e. & collins, s.l., 2014, ‘interactive effects of grazing, drought, and fire on grassland plant communities in north america and south africa’, ecology 95(1), 98–109. https://doi.org/10.1890/13-0526.1 kulmatiski, a. & beard, k.h., 2013a, ‘root niche partitioning among grasses, saplings, and trees measured using a tracer technique’, oecologia 171(1), 25–37. https://doi.org/10.1007/s00442-012-2390-0 kulmatiski, a. & beard, k.h., 2013b, ‘woody plant encroachment facilitated by increased precipitation intensity’, nature climate change 3(9), 833. https://doi.org/10.1038/nclimate1904 levick, s. & rogers, k., 2008, ‘patch and species specific responses of savanna woody vegetation to browser exclusion’, biological conservation 141(2), 489–498. https://doi.org/10.1016/j.biocon.2007.11.001 levick, s.r., asner, g.p., kennedy-bowdoin, t. & knapp, d.e., 2009, ‘the relative influence of fire and herbivory on savanna three-dimensional vegetation structure’, biological conservation 142(8), 1693–1700. https://doi.org/10.1016/j.biocon.2009.03.004 levick, s.r., asner, g.p., kennedy-bowdoin, t. & knapp, d.e., 2009, ‘the spatial extent of termite influences on herbivore browsing in an african savanna’, biological conservation 143(11), 2462–2467. https://doi.org/10.1016/j.biocon.2010.06.012 macfadyen, d.n., avenant, n.l., van der merwe, m. & bredenkamp, g.j., 2012, ‘the influence of fire on rodent abundance at the n’washitshumbe enclosure site, kruger national park, south africa’, african zoology 47(1), 138–146. https://doi.org/10.3377/004.047.0116 malhi, y., doughty, c.e., galetti, m., smith, f.a., svenning, j.-c. & terborgh, j.w., 2016, ‘megafauna and ecosystem function from the pleistocene to the anthropocene’, proceedings of the national academy of sciences 113(4), 838–846. https://doi.org/10.1073/pnas.1502540113 mccleery, r., monadjem, a., baiser, b., fletcher jr, r., vickers, k. & kruger, l., 2018, ‘animal diversity declines with broad-scale homogenization of canopy cover in african savannas’, biological conservation 226, 54–62. https://doi.org/10.1016/j.biocon.2018.07.020 midgley, j.j., coetzee, b.w., tye, d. & kruger, l.m., 2020, ‘mass sterilization of a common palm species by elephants in kruger national park, south africa’, scientific reports 10(1), 1–5. https://doi.org/10.1038/s41598-020-68679-8 midgley, j.j., gallaher, k. & kruger, l.m., 2012, ‘the role of the elephant (loxodonta africana) and the tree squirrel (paraxerus cepapi) in marula (sclerocarya birrea) seed predation, dispersal and germination’, journal of tropical ecology 28(2), 227–231. https://doi.org/10.1017/s0266467411000654 namah, j., midgley, j.j. & kruger, l.m., 2019, ‘reproductive biology of the sausage tree (kigelia africana) in kruger national park, south africa’, koedoe 61(1), 1–7. https://doi.org/10.4102/koedoe.v61i1.1512 o’keefe, t. & alard, g.f., 2002, ‘effects of herbivores and fire on riparian and upland savanna ecosystems: field operations manual for herbivore & fire exclosures on the sabie and letaba rivers in the kruger national park’, sanparks report, scientific services, skukuza, south african national parks. paterson, d. & steenekamp, p., 2003, soil survey of nkuhlu exclosure, agricultural research council, institute for soil, climate and water, pretoria, south africa. pellegrini, a.f., refsland, t., averill, c., terrer, c., staver, a.c., brockway, d.g. et al., 2021, ‘decadal changes in fire frequencies shift tree communities and functional traits’, nature ecology & evolution 5(4), 504–512. https://doi.org/10.1038/s41559-021-01401-7 rogers, k.h., 2003, ‘adopting a heterogeneity paradigm: implications for management of protected savannas’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience. ecology and management of savanna heterogeneity, pp. 41–58, island press, washington, dc. rowan, j. & faith, j.t., 2019, ‘the paleoecological impact of grazing and browsing: consequences of the late quaternary large herbivore extinctions’, in i.j gordon & h.h.t prins (eds.), the ecology of browsing and grazing ii, ecological studies (analysis and synthesis), p. 451, springer, cham scogings, p.f., 2011, ‘stem growth of woody species at the nkuhlu exclosures, kruger national park: 2006–2010’, koedoe 53(1), 01–09. https://doi.org/10.4102/koedoe.v53i1.1035 scogings, p.f., 2014, ‘large herbivores and season independently affect woody stem circumference increment in a semi-arid african savanna’, plant ecology 215(12), 1433–1443. https://doi.org/10.1007/s11258-014-0400-5 scogings, p.f., hattas, d., skarpe, c., hjältén, j., dziba, l., zobolo, a. et al., 2015, ‘seasonal variations in nutrients and secondary metabolites in semi-arid savannas depend on year and species’, journal of arid environments 114, 54–61. https://doi.org/10.1016/j.jaridenv.2014.11.003 scogings, p.f., hjältén, j. & skarpe, c., 2011, ‘secondary metabolites and nutrients of woody plants in relation to browsing intensity in african savannas’, oecologia 167(4), 1063–1073. https://doi.org/10.1007/s00442-011-2042-9 scogings, p.f., hjältén, j. & skarpe, c., 2013, ‘does large herbivore removal affect secondary metabolites, nutrients and shoot length in woody species in semi-arid savannas?’, journal of arid environments 88, 4–8. https://doi.org/10.1016/j.jaridenv.2012.08.010 scogings, p.f., hjältén, j., skarpe, c., hattas, d., zobolo, a., dziba, l. et al., 2014, ‘nutrient and secondary metabolite concentrations in a savanna are independently affected by large herbivores and shoot growth rate’, plant ecology 215(1), 73–82. https://doi.org/10.1007/s11258-013-0279-6 scogings, p.f., johansson, t., hjältén, j. & kruger, j., 2012, ‘responses of woody vegetation to exclusion of large herbivores in semi-arid savannas’, austral ecology 37(1), 56–66. https://doi.org/10.1111/j.1442-9993.2011.02249.x scogings, p.f., mamashela, t.c. & zobolo, a.m., ‘deciduous sapling responses to season and large herbivores in a semi-arid african savanna’, austral ecology 38(5), 548–556. https://doi.org/10.1111/j.1442-9993.2012.02454.x siebert, f. & eckhardt, h.c., 2008, ‘the vegetation and floristics of the nkhuhlu exclosures, kruger national park’, koedoe: african protected area conservation and science 50(1), 126–144. https://doi.org/10.4102/koedoe.v50i1.138 siebert, f., siebert, s.j. & eckhardt, h.c., 2010, ‘the vegetation and floristics of the letaba exclosures, kruger national park, south africa’, koedoe: african protected area conservation and science 52(1), 1–12. https://doi.org/10.4102/koedoe.v52i1.777 siebert, f. & scogings, p., 2015, ‘browsing intensity of herbaceous forbs across a semi-arid savanna catenal sequence’, south african journal of botany 100, 69–74. https://doi.org/10.1016/j.sajb.2015.05.007 sisson, d., hufschmid, j., jolles, a., beechler, b. & jabbar, a., 2017, ‘molecular characterisation of anaplasma species from african buffalo (syncerus caffer) in kruger national park, south africa’, ticks and tick-borne diseases 8(3), 400–406. https://doi.org/10.1016/j.ttbdis.2017.01.003 smit, i.p., peel, m.j., ferreira, s.m., greaver, c. & pienaar, d.j., 2020, ‘megaherbivore response to droughts under different management regimes: lessons from a large african savanna’, african journal of range & forage science 37(1), 65–80. https://doi.org/10.2989/10220119.2019.1700161 smit, i.p. & prins, h.h., 2015, ‘predicting the effects of woody encroachment on mammal communities, grazing biomass and fire frequency in african savannas’, plos one 10(9), e0137857. https://doi.org/10.1371/journal.pone.0137857 smit, i.p., roux, d.j., swemmer, l.k., boshoff, n. & novellie, p., 2017, ‘protected areas as outdoor classrooms and global laboratories: intellectual ecosystem services flowing to-and-from a national park’, ecosystem services 28(part b), 238–250. https://doi.org/10.1016/j.ecoser.2017.05.003 smith, f.a., doughty, c.e., svenning, j.-c. & terborgh, j.w., 2016a, ‘megafauna in the earth system’, ecography 39(2), 99–108. https://doi.org/10.1111/ecog.02156 smith, m.d., knapp, a.k., collins, s.l., burkepile, d.e., kirkman, k.p., koerner, s.e. et al., 2016b, ‘shared drivers but divergent ecological responses: insights from long-term experiments in mesic savanna grasslands’, bioscience 66(8), 666–682. https://doi.org/10.1093/biosci/biw077 smith, m.k.s., smit, i.p., swemmer, l.k., mokhatla, m.m., freitag, s., roux, d.j. et al., 2021, ‘sustainability of protected areas: vulnerabilities and opportunities as revealed by covid-19 in a national park management agency’, biological conservation 255, 108985. https://doi.org/10.1016/j.biocon.2021.108985 steenekamp, p., 2003, soil survey of nkuhlu exclosure, agricultural research council, institute for soil, climate and water, pretoria, south africa. stanton, r.a. jr., boone iv, w.w., soto-shoender, j., fletcher jr, r.j., blaum, n. & mccleery, r.a., 2018, ‘shrub encroachment and vertebrate diversity: a global meta-analysis’, global ecology and biogeography 27(3), 368–379. https://doi.org/10.1111/geb.12675 staver, a.c., abraham, j.o., hempson, g.p., karp, a.t. & faith, j.t., 2021, ‘the past, present, and future of herbivore impacts on savanna vegetation’, journal of ecology 109(8), 2804–2822. https://doi.org/10.1111/1365-2745.13685 swemmer, a.m., bond, w.j., donaldson, j., hempson, g.p., malherbe, j. & smit, i.p., 2018, ‘the ecology of drought-a workshop report’, south african journal of science 114(9–10), 1–3. https://doi.org/10.17159/sajs.2018/5098 thoresen, j., vermeire, m.-l., venter, z., wolfaard, g., krumins, j., cramer, m. & hawkins, h.j., 2021, ‘fire and herbivory shape soil arthropod communities through habitat heterogeneity and nutrient cycling in savannas’, global ecology and conservation 25, e01413. https://doi.org/10.1016/j.gecco.2020.e01413 treydte, a.c., baumgartner, s., heitkönig, i.m., grant, c.c. & getz, w.m., 2013, ‘herbaceous forage and selection patterns by ungulates across varying herbivore assemblages in a south african savanna’, plos one 8(12), e82831. https://doi.org/10.1371/journal.pone.0082831 van coller, h. & siebert, f., 2015, ‘herbaceous biomass–species diversity relationships in nutrient hotspots of a semi-arid african riparian ecosystem’, african journal of range & forage science 32(3), 213–223. https://doi.org/10.2989/10220119.2014.951394 van coller, h. & siebert, f., 2020, ‘the impact of herbivore exclusion on forb diversity: comparing species and functional responses during a drought’, african journal of ecology 58(2), 236–250. https://doi.org/10.1111/aje.12676 van coller, h., siebert, f., scogings, p.f. & ellis, s., 2018, ‘herbaceous responses to herbivory, fire and rainfall variability differ between grasses and forbs’, south african journal of botany 119, 94–103. https://doi.org/10.1016/j.sajb.2018.08.024 van coller, h., siebert, f. & siebert, s.j., 2013, ‘herbaceous species diversity patterns across various treatments of herbivory and fire along the sodic zone of the nkuhlu exclosures, kruger national park’, koedoe 55(1), 01–06. https://doi.org/10.4102/koedoe.v55i1.1112 vermeire, m.-l., thoresen, j., lennard, k., vikram, s., kirkman, k., swemmer, a.m. et al., 2021, ‘fire and herbivory drive fungal and bacterial communities through distinct above-and belowground mechanisms’, science of the total environment 785, 147189. https://doi.org/10.1016/j.scitotenv.2021.147189 voysey, m.d., archibald, s., bond, w.j., donaldson, j.e., carla staver, a. & greve, m., 2021, ‘the role of browsers in maintaining the openness of savanna grazing lawns’, journal of ecology 109(2), 913–926. https://doi.org/10.1111/1365-2745.13518 wigley, b.j., fritz, h., coetsee, c. & bond, w.j., 2014, ‘herbivores shape woody plant communities in the kruger national park: lessons from three long-term exclosures’, koedoe 56(1), 1–12. https://doi.org/10.4102/koedoe.v56i1.1165 wigley-coetsee, c. & staver, a.c., 2020, ‘grass community responses to drought in an african savanna’, african journal of range & forage science 37(1), 43–52. https://doi.org/10.2989/10220119.2020.1716072 wilcox, k.r., koerner, s.e., hoover, d.l., borkenhagen, a.k., burkepile, d.e., collins, s.l. et al., 2020, ‘rapid recovery of ecosystem function following extreme drought in a south african savanna grassland’, ecology 101(4), e02983. https://doi.org/10.1002/ecy.2983 filelist convert a pdf file! introduction methods results discussion acknowledgements references about the author(s) marika e. van niekerk department of genetics, university of the free state, bloemfontein, south africa francois deacon department of animal, wildlife and grassland sciences, university of the free state, bloemfontein, south africa paul j. grobler department of genetics, university of the free state, bloemfontein, south africa citation van niekerk, m.e., deacon, f. & grobler, p.j., 2019, ‘the genetic status of the introduced giraffe population in central south africa’, koedoe 61(1), a1570. https://doi.org/10.4102/koedoe.v61i1.1570 short communication the genetic status of the introduced giraffe population in central south africa marika e. van niekerk, francois deacon, paul j. grobler received: 22 mar. 2019; accepted: 05 june 2019; published: 16 sept. 2019 copyright: © 2019. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. introduction there has been no reliable historical evidence of giraffe occurring naturally in the free state province in central south africa (dagg 1962; deacon & parker 2016; sydney 1965). although lynch (1983) mentioned the possibility of the natural distribution of giraffe in the east and western free state, there is no concrete evidence that giraffe existed earlier than this in the free state (ansell 1968). according to griesel (1961), hirst (1966), lambrechts (1974) and terblanche and kok (1995), translocations of the species into protected areas and private land in the free state province happened in any case, sometimes regardless of the natural habitat preferences of giraffe (castley, boshof & kerley 2001; deacon & tutchings 2019). these authors also affirmed that giraffe had, however, occurred naturally westwards of the free state, in the northern cape province. the introduced status of the giraffe population in central south africa, and in particular the free state province, thus raises questions regarding the origin and the taxonomic status of these animals. although giraffe is an introduced species in the free state province, it is also a reality that extralimital populations of large mammals add value to ecotourism in self-financing private protected areas (maciejewski & kerley 2014) and are therefore kept on many properties. extralimital animals can also still add conservation value by contributing to long-term population survival, while extralimital populations could in some cases nevertheless be a reservoir of valuable genetic resources. the newly created ownership in wildlife species not only contributes to diversity in wildlife species for ecotourism, but also assists to protect valuable and key resource areas and habitat for all fauna and flora. giraffe taxonomy and subspecies status have been the subject of much debate over the last century. it should also be noted that there have been only limited representation from south african populations in the reported studies. bercovitch and deacon (2015, 2017) stated that there are currently at least four different taxonomic classifications for giraffe with the current global acceptance of nine subspecies (muller et al. 2016). most recently, fennessy et al. (2016) and winter et al. (2018) suggested a new taxonomy for giraffe with four genetically distinct giraffe species instead of one, but this has not been accepted by the international union for conservation of nature (iucn) who requested more evidence and a larger sample size; and the status quo at least for now is one species with nine subspecies (bercovitch et al. 2017). it should also be noted that the aforementioned genetic studies have only limited representation from south african populations. irrespective of the exact classification system used (one or four species), all the current classification systems include recognition of a ‘south african giraffe’ (giraffa camelopardalis giraffe/g. giraffa giraffa) and the ‘angolan giraffe’ (giraffa camelopardalis angolensis [g. c. angolensis]/giraffa giraffa angolensis [g. g. angolensis]). fennessy et al. (2016) suggested that south africa harbours only the south african giraffe. however, deacon and parker (2016) state that the angolan giraffe has been translocated from namibia to countries such as botswana and south africa, and some game ranches might contain a mixture of subspecies, or even hybrid giraffe. the extent of historical natural migration of animals between regions is also not known. the available data on historical and current distribution ranges of south african giraffe are shown in figure 1 (based on the iucn red list assessment of muller et al. 2018). figure 1: the current and historical distribution of giraffe in south africa, regions sampled and localities referred to in the text. in addition to taxonomic considerations, genetic diversity within giraffe populations in the region is also a potential cause for concern (deacon & tutchings 2019). in the free state province, giraffe are almost invariably found in small populations, for reasons of habitat availability or because they may not be utilised as intensively as some other wildlife species. the number of giraffe found on farms thus ranges from populations numbering 10–30 down to populations with single pairs of animals, with the potential risk of decreased genetic diversity and ultimately reduced fitness or adaptability. management strategies to maintain diversity is needed, but, conversely, the necessary translocations needed to conserve genetic diversity could hold risks if the taxa involved are significantly different, in line with theory on the mixing of distinct taxonomic units (crandall et al. 2000; moritz 2002) and outbreeding depression (edmands 2006). with little being known about the effect of extralimital introductions and fragmentation in giraffe populations in central south africa, the aim of the current project was to determine the genetic structure of giraffe in central south africa relative to the wider southern african population, and also obtain an estimate of genetic diversity within and between individual giraffe populations. methods study sites and sample collection we collected samples from three groups: (1) a total of 34 giraffe were sampled across 10 privately owned herds in the free state province (fs-private). the census population size (n) of these groups ranged from 2 to 30+; and the ultimate origins of the animals in these populations are unknown other than the fact that founding involved several sources and that translocations among populations have occurred in recent times. (2) as representative of giraffe in public protected herds from the province, nine giraffe were sampled from the willem pretorius provincial nature reserve (n=21). two groups of founders sourced from the limpopo province and one group from the kwazulu-natal province were originally used to start this population (fs-protected area). (3) finally, four giraffe were sampled from a privately owned herd in the adjacent northern cape province (n = 20). eight individuals were translocated to this locality from namibia in 1971, with another six individuals translocated from the then transvaal province in northern-eastern south africa in 1973. we used dung as the preferred sample type, to minimise stress on the individuals, as well as for cost purposes, ethical considerations and ease of access to populations. the procedure followed at each locality was as follows: once a giraffe was found, it was observed until defecation had occurred. the exact spot was noted; the giraffe was characterised using size, sex and patterns on its body to prevent duplicate sampling; and the sample retrieved. samples were stored in 96% ethanol and at -20 °c as soon as possible after collection. a small number of skeletal muscle and blood samples available opportunistically for specific localities were also included. genetic analysis deoxyribonucleic acid (dna) was extracted using the zymo research zr fecal dna miniprep™ kit used for dung samples, the zymo research quick-dna™ ffpe miniprep kit for tissue samples and the roche high pure pcr template preparation kit used for blood samples. a nanodrop® nd-1000 spectrophotometer was used to determine the quantity and quality of extracted dna. we sequenced sections of the cytochrome b (cytb) and displacement loop (d-loop) mitochondrial dna (mtdna) regions. the cytb primers used were from a study done by bock et al. (2014): forward – gtg gaa ggc gaa gaa tcg; reverse – gaa aaa cca tcg ttg tcg t; with the sequences of primers used for the d-loop region sourced from a study done by seymour (2001): forward – ccc aaa gct gaa gtt cta tt; reverse – caa taa ctg tat gta cta tg-3′. pcr reactions were based on the kapa2g™ robust hotstart readymix pcr kit for the cytb region, with the ampliqon tempase hot master mix a kit used for the d-loop region. annealing temperatures were 62 °c and 50 °c, respectively. polymerase chain reaction (pcr) products were cleaned using the biospin pcr purification kit (bio-rad), followed by sequencing (in both directions) using the abi prism® bigdye® terminator version 3.1 cycle sequencing kit and standard conditions. products were cleaned using the zr dna sequencing clean-up™ kit, before running on either an abi3130 or 3500 genetic analyser. data analysis geneious pro version 4.7.4 software (kearse et al. 2012) was used to view, assemble and align all sequences. mega v7.0.26 software (kumar, stecher & tamura 2016) was utilised to identify the best model of substitution for each mtdna region separately. the haplotypes for cytb and the d-loop region were then combined into a concatenated dataset for all subsequent analyses. phylogenetic trees were constructed using a maximum likelihood (ml) approach, with 1000 bootstrap replications (in mega software). to complement the phylogenetic analysis, network version 5.0.1.1 software (fluxus engineering 2019, www.fluxux-engineering.com) was used to construct haplotype networks, using a median-joining approach. genetic divergence between populations was quantified as the average number of nucleotide differences between populations (dxy), from dna sequence polymorphism (dnasp) version 5 (rozas et al. 2003), and genetic diversity within each group using haplotype diversity (hd) and nucleotide diversity (π) from dnasp software. as a reference database for g. c. giraffa and g. c. angolensis, the concatenated mtdna d-loop and cytb sequences used by winter et al. (2018) were made available by these authors. the latter authors also provided sequences for g. c. peralta and g. c. tippelskilchi, to use as an out-group. ethical considerations the methods employed during this study were approved by the interfaculty animal ethics committee of the university of the free state (numbers: ufs-aed2015/0050 and ufs-aed2015/0066). collection of samples was sanctioned under permits issued by destea (permit number: 01/30305) and the northern cape department of environment and nature conservation (permit number: fauna 0729/2017 and fauna 0730/2017). section 20 veterinary approval for the collection, transport and storage of samples was obtained from the national department of agriculture, forestry and fisheries (daff), with approval number 12/11/1/4. results after alignment of sequences and trimming to equal length, a sequence length of 405 base pairs (bp) was used for further analysis of the cytb gene, with 275 bp available for the d-loop region. the two cytb haplotypes observed have been submitted to the genbank database as mh033837 and mh033838, and the 10 haplotypes for the d-loop region as mh033839 to mh033848. our concatenated sequences were aligned with the reference sequences (total 1556 bp and all sequences trimmed to the shared 680 bp. the data for the two mtdna regions were tested separately to determine the model of substitution, and both regions were best described by a tamura 3-parameter model of substitution. a phylogenetic tree, based on the concatenated dataset and an ml approach, is presented in figure 2a. all the g. c. giraffa reference sequences form a clade with 75% bootstrap support, separate from g. c. angolensis. this cluster also contains all the giraffe samples from the free state protected area and 14 giraffe samples from private farms in the free state province. all other giraffe samples from the private farms in the free state province, comprising 21 individuals, cluster with g. c. angolensis, as do the four samples from the northern cape. the reference g. c. peralta and g. c. tippelskirchi haplotypes formed a distinct cluster with 100% bootstrap support. the trend from the ml tree is confirmed in the haplotype network presented in figure 2b. haplotypes of g. c. angolensis and g. c. giraffa form distinct clusters, with giraffe from a free state protected area clustering with g. c. giraffa and with giraffe from private populations grouping with one or the other subspecies. figure 2: status of giraffe from central south africa, relative to reference giraffe populations, based on a concatenated mitochondrial dna displacement loop (d-loop) and cytochrome b (cytb) sequence dataset. (a) phylogeny from maximum likelihood topology of haplotypes. the haplotype names from giraffe analysed for the current study are underlined, with reference samples indicated in [square brackets]. sample sizes are also indicated. numbers above branches indicate the percentage of bootstrap replicates where the clade was found. (b) minimum spanning network. nodes represent haplotypes, with the size of each node showing the number of individuals that share that haplotype. crossbars reflect the number of mutational events between specific haplotype pairs. the average number of nucleotide differences between reference sample sets of g. c. giraffa and g. c. angolensis was dxy = 0.033. this value was regarded as representative of the difference between pure populations of the two subspecies and was therefore used as a yardstick to gauge the relative magnitude of values obtained for pairwise comparisons between each of the studied populations and the two subspecies. the population from the free state protected area showed comparatively low differentiation from g. c. giraffa, with dxy = 0.014, but with dxy = 0.020 when compared with g. c. angolensis. the northern cape population showed close identity with g. c. angolensis, with dxy = 0.004, but dxy between the northern cape group and g. c. giraffa was much higher at 0.034, thus equalling the distance between g. c. giraffa and g. c. angolensis. the dxy value between the privately owned giraffes and g. c. giraffa and g. c. angolensis was 0.028 and 0.009, respectively. levels of genetic diversity within groups were as follows (with format – number of haplotypes/number of polymorphic sites/haplotype diversity [hd] and nucleotide diversity [π]): fs-private populations – 10/19/0.814/0.011; fs-protected area – 2/1/0.389/0.001 and northern cape – 2/1/0.667/0.008. by comparison, values for the reference datasets were g. c. giraffa – 7/6/0.508/0.001 and g. c. angolensis – 7/6/0.844/π = 0.003. discussion although we were not able to sequence the full length of mtdna used by winter et al. (2018), results from the current study provided valuable new data on the status of the giraffe population in central south africa. based on the concatenated mtdna cytb and d-loop dataset used, a total of 23 individuals classified as g. c. giraffa. these animals originated from a provincial nature reserve and six private game farms. this is in line with expectations that locally sampled individuals would have mtdna lineages of what is currently regarded as the south african giraffe subspecies. unexpectedly, 24 individuals from the present study classified as g. c. angolensis. this outcome suggests that a significant number of individuals with mtdna lineages of the angolan giraffe rather than the south african giraffe are present in the central south african giraffe population. we note that those individuals displaying g. c. angolensis haplotypes may also be hybrids, either first generation or more advanced generations, resulting from initial crosses involving g. c. giraffa males and g. c. angolensis females. (conversely, some of the individuals classified as g. c. giraffa may also be hybrids, with male g. c. angolensis individuals in their pedigrees.) population-genetic analysis based on the average number of nucleotide differences between groups confirmed that the overall privately owned giraffe herds in central south africa show an unexpected identity with g. c. angolensis. there are two possible scenarios to explain the observation of a g. c. angolensis lineage in south african populations, based on artificial translocations and unrecorded historical migrations. the giraffe is a popular game ranching species, and there has been substantial commercial movement of the animal around the sub-continent (deacon & parker 2016). the findings that some of the giraffe sampled in the free state are potentially angolan giraffe may exemplify how the subspecies has been given a more southern distribution to what was previously thought, through translocations. this can be partly explained by the close historical and current link between south africa and namibia, with substantial translocation of game animals between the two countries, and individuals of g. c. angolensis may have found their way into south africa along this route. for example, the kglagadi transfrontier park contains both introduced g. c. angolensis and g. c. giraffa (deacon & tutchings 2019). we note that game breeders may not be aware of the subspecies status of the giraffe on their properties because this is not currently a publicised conservation concern, and rather focus on maintaining viable populations through introductions from readily available sources. in an alternative scenario, the current results may suggest that a re-evaluation of subspecies distribution in the region is necessary. in this scenario, g. c. angolensis detected in south africa may in fact be part of a natural continuum of diversity of lineages existing within and between the two subspecies. winter et al. (2018) discuss how the historically assumed distribution of the subspecies ranged, and that the current distribution of g. c. angolensis ranges over a wider span than what was previously thought. bock et al. (2014) found several deviations from expected subspecies distribution. for example, two giraffe that had been assumed to be south african giraffe grouped with angolan. a large number of giraffe sampled were from the chobe national park, nxai pans, vumbura concession and moremi game reserve in northern botswana, and bwabwata national park in northeastern namibia. namibian localities assumed to have g. c. angolensis grouped with g. c. giraffa from the khamab kalahari reserve in south africa. these authors state that the assignment of giraffe to the incorrect subspecies could be because of either natural migration or human-induced translocation. in their study, bock et al. (2014) also found that the south african giraffe were also distributed further north than the previously assumed range. furthermore, winter et al. (2018) found that the angolan giraffe had a more eastern distribution than the previously known distribution. there is thus a need for a finer grained study of natural populations, to determine the true borders of subspecies and lineages, and to better understand the spectrum of pure genotypes for both the south african and angolan giraffe. within the south african giraffe population, differences between historic origin populations (kruger national park) in comparison to translocated populations such as the individuals in central south africa should be investigated. the observed levels of genetic diversity supported our conclusion on the taxonomic status of the groups studied. the population from a public protected area showed least diversity of all groups studied, despite being a sizeable population in the context of this study. however, this population also showed close identity with g. c. giraffa, which has lower diversity compared to g. c. angolensis. conversely, a small sample of animals from the northern cape showed high diversity and also showed close identity to g. c. angolensis, which has higher diversity compared to g. c. giraffa. the highest level of genetic diversity recorded in this study – higher than values estimated for either pure g. c. giraffa or g. c. angolensis – was found in the pooled sample set for privately owned giraffe in the free state province. grobler et al. (2018) showed that hybridisation may result in artificially elevated levels of genetic diversity. the high extant levels of diversity found in the privately owned herds may thus reflect a mix of haplotypes of both g. c. giraffa and g. c. angolensis. in addition to providing new data on subspecies status, the results from the current study have significant implications for the conservation of genetic diversity in small giraffe populations in central south africa. some of the individual populations sampled consisted of four or fewer giraffe, sometimes founded using animals bought from a single source population and with all animals bought at the same time. these, and many similar small giraffe populations in central south africa, cannot reasonably be expected to add significant numbers of additional giraffe to their populations because of the size of the areas and habitat availability (carrying capacity). exchange of animals among populations may thus have significant advantages in inducing gene flow. however, this suggested management approach is based on the premise that uncontrolled exchanges are permissible from a taxonomic point of view, that is, that all extant animals classify to the same subspecies or another appropriate unit for conservation. the latter requirement is, however, clearly not met by the extant free state giraffe populations, based on the trends reported in the preceding sections. the data presented here provide evidence that both subspecies from southern africa are present in populations sampled from central south africa. this could reflect either translocations or the fact that the borders of the subspecies’ ranges are not clearly defined. for future study, we suggest that sampling and analysis be expanded to provide further data on subspecies’ distribution and possible admixture. in this regard, museum samples could also be used to elucidate the range of each mtdna haplotype, and microsatellite makers will provide more precise data on admixture, especially where male mtdna is not detected in hybrids. finally, the adaptive significance of genetic differences between subspecies should be investigated. acknowledgements the authors thank all private landowners and the free state department of economic, small business development, tourism and environmental affairs (destea) for permission to sample populations. the authors thank two anonymous reviewers for their constructive comments that improved the article. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions f.d. and p.j.g. conceptualised the study. m.e.v.n. conducted fieldwork and performed all laboratory analysis. p.j.g. and m.e.v.n. conducted statistical analysis. all the authors contributed to writing this article. funding information the study was partly financed through the national research foundation incentive funding to p.j.g. data availability statement sequences generated for this study are available from genbank. disclaimer the views and opinions expressed in this article are the authors’ own and not an official position of the institution or funder. references ansell, w.f.h., 1968, ‘part 15. order artiodactyla’, in j. meester & h.w. setzer (eds.), the mammals of africa: an identification manual, pp. 1–84, smithsonian institution press, washington, dc, https://doi.org/10.1111/j.1096-3642.1965.tb00081.x bercovitch, f.b., berry, p.s.m., dagg, a., deacon, f., doherty, j.b., lee, d. et al., 2017, ‘how many giraffe species exist?’ current biology 27, r123–r138, https://doi:10.1016/j.cub.2016.12.039 bercovitch, f.b. & deacon, f., 2015, ‘gazing at a giraffe gyroscope: where are we going?’ african journal of ecology 53, 135–146. https://doi.org/10.1111/aje.12222 bock, f., fennessy, j., bidon, t., tutchings, a., marais, a., deacon, f. et al., 2014, ‘mitochondrial sequences reveal a clear separation between angolan and south african giraffe along a cryptic rift valley’, bmc evolutionary biology 14, 219. https://doi.org/10.1186/s12862-014-0219-7 castley, j.g., boshof, f, a.f. & kerley, g.i.h., 2001, ‘compromising south africa’s natural biodiversity – inappropriate herbivore introductions’, south african journal of science 97, 334–348. https://hdl.handle.net/10520/ejc97379 crandall, k.a., bininda-edmonds, o.r.p., mace, g.m. & wayne, r.k., 2000, ‘considering evolutionary processes in conservation biology: an alternative to “evolutionarily significant units”’, trends in ecology and evolution 15, 290–295. https://doi.org/10.1016/s0169-5347001876-0 dagg, a.i., 1962, ‘the distribution of the giraffe in africa’, mammalia 26, 497–505. https://doi.org/10.1515/mamm-1962-0405 deacon, f. & parker, d., 2016, ‘a conservation assessment of giraffa camelopardalis giraffa’, in m.f. child, l. roxburgh, e. do linh san, d. raimondo & h.t. davies-mostert (eds.), the red list of mammals of south africa, swaziland and lesotho, pp. 1–9, south african national biodiversity institute and endangered wildlife trust, south africa. deacon, f. & tutchings, a., 2019, ‘the south african giraffe giraffa camelopardalis giraffa: a conservation success story’, oryx 53(1), 45–48. https://doi.org/10.1017/s0030605317001612 edmands, s., 2006, ‘between a rock and a hard place: evaluating the relative risks of inbreeding and outbreeding for conservation and management’, molecular ecology 16, 463–475. https://doi.org/10.1111/j.1365-294x.2006.03148.x fennessy, j., bidon, t., reuss, f., kumar, v., elkan, p., nilsson, m.a. et al., 2016, ‘multi-locus analyses reveal four giraffe species instead of one’, current biology 26(18), 2543–2549. https://doi:10.1016/j.cub.2016.07.036 griesel, j., 1961, ‘a new game sanctuary for the orange free state, the willem pretorius game reserve’, african wildlife 15(2), 121–125. grobler, j.p., van wyk, a.m., dalton, d.l., jansen van vurren, b. & kotze, a., 2018, ‘assessing introgressive hybridization between blue wildebeest (connochaetes taurinus) and black wildebeest (connochaetes gnou) from south africa’, conservation genetics 19, 981–993. https://doi.org/10.1007/s10592-018-1071-x hirst, s.m., 1966, immobilization of the transvaal giraffe giraffa camelopardalis giraffa using an oripavin derivative’, journal of the south african veterinary association 37, 85. kearse, m., moir, r., wilson, a., stones-havas, s., cheung, m., sturrock, s. et al., 2012, ‘geneious basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data’, bioinformatics 28(12), 1647–1649. https://doi.org/10.1093/bioinformatics/bts199 kumar, s., stecher, g. & tamura, k., 2016, ‘mega7: molecular evolutionary genetics analysis version 7.0.’, molecular biology and evolution 33(7), 1870–1874. https://doi.org/10.1093/molbev/msw054 lambrechts, a.w., 1974, ‘the numerical status of sixteen game species in the transvaal, excluding the kruger national park’, journal of the south african wildlife management association 4, 95–102. https://hdl.handle.net/10520/aja03794369 lynch, c.d., 1983, ‘the mammals of the orange free state’, memoirs van die nasionale museum bloemfontein 18, 1–218. maciejewski, k. & kerley, g.i.h., 2014, ‘understanding tourists’ preference for mammal species in private protected areas: is there a case for extralimital species for ecotourism?’, plos one 9(2), e88192. https://doi.org/10.1371/journal.pone.0088192 moritz, c., 2002, ‘strategies to protect biological diversity and the evolutionary processes that sustain it’, systematic biology 51, 238–254. https://doi:10.1080/10635150252899752 muller, z., bercovitch, f., brand, r., brown, d., brown, m., bolger, d. et al., 2018, giraffa amelopardalis (amended version of 2016 assessment), the iucn red list of threatened species 2018: e.t9194a136266699, viewed 28 june 2018 from https://www.iucnredlist.org/species/9194/136266699n. muller, z., bercovitch, f., fennessy, j., brown, d., brand, r., brown, m. et al., 2016, giraffa camelopardalis, the iucn red list of threatened species 2016: e.t9194a51140239, viewed 08 december 2016 from www.iucnredlist.org. rozas, j., sánchez-delbarrio, j.c., messeguer, x. & rozas, r., 2003, ‘dnasp, dna polymorphism analyses by the coalescent and other methods’, bioinformatics 19, 2496–2497. https://doi.org/0.1093/bioinformatics/btg359 seymour, r.s., 2001, ‘patterns of subspecies diversity in the giraffe, giraffa camelopardalis (l. 1758): comparison of systematic methods and their implications for conservation policy’, phd thesis, university of kent, canterbury. https://ethos.bl.uk/orderdetails.do?uin=uk.bl.ethos.275008 sydney, j., 1965, ‘the past and present distribution of some african ungulates’, proceeding of the zoological society of london 30, 139–167. https://doi.org/10.1017/s0030605300006815 terblanche, h.j. & kok, o.b., 1995, ‘die voorkoms van wild in die oranje-vrystaat’, navorsinge van die nasionale museum bloemfontein 11(4), 61–100. https://hdl.handle.net/10520/aja00679208_1465 winter, s., fennessy, j., fennessy, s. & janke, a., 2018, ‘matrilineal population structure and distribution of the angolan giraffe in the namib desert and beyond’, ecological genetics and genomics 7–8, 1–5. https://doi.org/10.1016/j.egg.2018.03.003 filelist convert a pdf file! page 1 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 page 4 http://www.koedoe.co.za open access koedoe african protected area conservation and science issn: (online) 2071-0771, (print) 0075-6458 page 1 of 1 corrigendum read online: scan this qr code with your smart phone or mobile device to read online. authors: kerry-ann van der walt1,2 ernst r. swartz3 darragh j. woodford3,4 olaf l.f. weyl1,2,3 affiliations: 1department of ichthyology and fisheries science, rhodes university, south africa 2centre for invasion biology, south african institute for aquatic biodiversity, south africa 3south african institute for aquatic biodiversity, grahamstown, south africa 4school of animal, plant and environmental sciences, university of the witwatersrand, south africa corresponding author: kerry-ann van der walt, k.vanderwalt@saiab.ac.za dates: published: 30 nov. 2017 how to cite this article: van der walt, k-a., swartz, e.r., woodford, d.j. & weyl, o.l.f., 2017, ‘corrigendum: using genetics to prioritise headwater stream fish populations of the marico barb, enteromius motebensis steindachner 1894, for conservation action’, koedoe 59(1), a1506. https://doi. org/10.4102/koedoe. v59i1.1506 copyright: © 2017. the authors. licensee: aosis. this work is licensed under the creative commons attribution license. in the version of this article initially published, kerry-ann van der walt’s and darragh j. woodford’s second affiliation, and olaf l.f. weyl’s first affiliation was omitted. the initials for darragh j. woodford and olaf l.f. weyl was also omitted. the author list and affiliations are hereby corrected to: authors: kerry-ann van der walt1,2 ernst r. swartz3 darragh j. woodford3,4 olaf l.f. weyl1,2,3 affiliations: 1department of ichthyology and fisheries science, rhodes university, south africa 2centre for invasion biology, south african institute for aquatic biodiversity, south africa 3south african institute for aquatic biodiversity, grahamstown, south africa 4school of animal, plant and environmental sciences, university of the witwatersrand, south africa the errors have been corrected in the pdf version of the article. the corresponding author apologises for any inconvenience that this omission may have caused. corrigendum: using genetics to prioritise headwater stream fish populations of the marico barb, enteromius motebensis steindachner 1894, for conservation action read online: scan this qr code with your smart phone or mobile device to read online. note: doi of original article: https://doi.org/10.4102/koedoe.v59i1.1375 http://www.koedoe.co.za mailto:k.vanderwalt@saiab.ac.za https://doi.org/10.4102/koedoe.v59i1.1506 https://doi.org/10.4102/koedoe.v59i1.1506 https://doi.org/10.4102/koedoe.v59i1.1506 http://crossmark.crossref.org/dialog/?doi=10.4102/koedoe.v59i1.1506=pdf&date_stamp=2017-11-30 https://doi.org/10.4102/koedoe.v59i1.1375 abstract introduction methods and materials results discussion conclusion acknowledgements references appendix 1 about the author(s) mathew a. harris department of plant and soil sciences, university of pretoria, pretoria, south africa michael d. voysey department of plant and soil sciences, university of pretoria, pretoria, south africa department of animal, plant and environmental sciences, university of the witwatersrand, johannesburg, south africa samantha-leigh jamison department of plant and soil sciences, university of pretoria, pretoria, south africa michelle greve department of plant and soil sciences, university of pretoria, pretoria, south africa citation harris, m.a., voysey, m.d., jamison, s.-l. & greve, m., 2019, ‘changes in bird assemblages because of vegetation homogenisation in communal livestock systems’, koedoe 61(1), a1543. https://doi.org/10.4102/koedoe.v61i1.1543 original research changes in bird assemblages because of vegetation homogenisation in communal livestock systems mathew a. harris, michael d. voysey, samantha-leigh jamison, michelle greve received: 21 may 2018; accepted: 14 feb. 2019; published: 17 july 2019 copyright: © 2019. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract protected areas are intended to promote biodiversity representation and persistence; yet, whether they are effective in degraded landscapes where much of the original vegetation structure remains intact has received relatively little attention. we test whether avian assemblages in communal rangelands in savannas differ from savannas supporting a full complement of native herbivores and predators. birds were surveyed in 36 transect counts conducted over 18 days. we also compare the vegetation structure between the two land-use types to assess whether differences in bird assemblages could be attributed to changes in vegetation structure. bird assemblages were richer, had greater abundances and different compositions inside protected areas than rangelands. the median body mass of birds was larger inside than outside protected areas, and rangelands supported fewer grassland specialists, but more closed-canopy specialists. however, no differences in feeding guild composition were found between protected areas and communal rangelands. additionally, vegetation structure, but not richness, differed between protected areas and communal rangelands: communal rangelands had higher densities of woody vegetation and shorter grass height than the protected areas. our findings suggest that the altered vegetation structure in communal grazing camps has led to changes in the species richness and composition of bird communities and has been selected by closed-canopy specialists at the cost of open grassy specialists. hunting in communal rangelands is likely to have resulted in the loss of large birds and in reductions in bird abundance in the rangelands. therefore, land-use management that does not lead to irreversible landscape transformation can nevertheless result in changes in the diversity, composition and functioning of native assemblages. conservation implications: savanna landscapes that are degraded, but not transformed, support fewer bird species, fewer open habitat specialists and smaller birds because of vegetation homogenisation. keywords: avifauna; encroachment; functional composition; hunting; vegetation structure; conservation. introduction it has been estimated that 52% of the earth’s surface has been modified by human activities such as food production, timber plantations and urban areas (roser & ritchie 2018). this has resulted in alterations in the structure and functioning of these systems (chown 2010; foley et al. 2005). with human population growth unlikely to stabilise in the 21st century (gerland et al. 2014), the protection of natural habitats remains paramount for the conservation of biodiversity and the ecosystem services they provide (dudley, hockings & stolton 2010). even where primary components of habitats have been retained, unprotected landscapes have often been degraded and community assemblages have been altered via either direct (e.g. harvesting) or indirect (e.g. light pollution) impacts by humans, or both (chown 2010; gaston et al. 2008; longcore & rich 2004). therefore, the designation and maintenance of protected areas (pas) remains a key strategy for protecting biodiversity from such pressures globally (gaston et al. 2008; joppa & pfaff 2009; pimm et al. 2014). a substantial proportion (~40%) of the estimated annual global conservation budget is utilised for establishing, maintaining and improving pas (balmford et al. 2015; chape et al. 2005; james, gaston & balmford 1999; waldron et al. 2013). yet, despite the enormous investment in pas annually (~$10 billion, waldron et al. 2013), only a limited number of studies have explicitly attempted to assess pa ecological effectiveness (e.g. coetzee, gaston & chown 2014; gray et al. 2016; greve et al. 2011), with most studies evaluating pa effectiveness in terms of how well the pa prevents landscape cover change (e.g. andam et al. 2008; joppa, loarie & pimm 2008). however, this measure serves only as a proxy for pa performance, as it cannot capture the implications of land-use change for biodiversity features (coetzee et al. 2014), especially when considering that the effectiveness of any given pa at conserving features of biodiversity is context specific and often depends on the extent and intensity of local pressures, for example fires or poaching. thus, individual pas often show localised differences in measures of their ecological effectiveness (caro et al. 2009; gaston et al. 2008; gray et al. 2016; laurance et al. 2012). the best measure of pa ecological effectiveness would be a comparison of the current overall biodiversity status for a pa with the biodiversity status that would have been if the pa had not been designated (coetzee et al. 2014). however, a comparison of this nature cannot be achieved, as it is impossible to know what the biodiversity status of a particular area would have been had a pa not been established. thus, an alternative is to assess biodiversity before and after pas were established; but baseline data quantifying biodiversity before pa declaration are rare (see laurance et al. 2011; wegge et al. 2009). as a result, assessments of the effectiveness of pas are assessed differently. for example pa effectiveness is measured based on management decisions and their effects on biodiversity within pas. others compare biodiversity features within pas to areas in the immediate vicinity or to non-protected areas with similar characteristics, that is, using space for time substitutions (geldmann et al. 2013). essentially, biodiversity features of areas inside pas are compared to biodiversity features outside pas, as these areas would be similar were it not for their land-use designation (coetzee et al. 2014; gray et al. 2016; greve et al. 2011). the assumption is that the areas inside pas would have experienced land-use changes similar to those outside the pas, had the pa not been established, and is thus used as a proxy measure of pa ecological effectiveness. the ecological effectiveness of pas can be considered at different hierarchical and taxonomic levels of organisation of biodiversity, for example species (abundance), communities (richness and assemblage composition) and the functions they perform (gaston et al. 2006). to maintain biodiversity, pas must display representation, that is, support a diversity of species (or functions) within their boundaries, and they must allow for persistence of biodiversity and function, that is buffer biodiversity from perturbations (margules & pressey 2000) such as climate change (stevens et al. 2016) or exploitation (cullen, bodmer & pádua 2000). the literature on quantifying pa ecological effectiveness has often focused on comparing biodiversity in pas to that of adjacent heavily transformed areas, for example agricultural fields, plantations and human settlements (gardner et al. 2007; jackson, evans & gaston 2009; hoffmann et al. 2010; greve et al. 2011; geldmann et al. 2013). the impacts on biodiversity of areas that have undergone comparatively little habitat transformation remain less well understood (see greve et al. 2011; rayner et al. 2014; sinclair, mduma & arcese 2002). therefore, this study aims to compare the biodiversity features, species assemblages and functional composition of avian fauna within and outside neighbouring pas, across what once was a homogenous vegetation type within the savanna biome. the land-use outside these pas is communal rangelands (crs), which are nearly completely uncultivated semi-natural rangelands, where collective ranching of livestock takes place (palmer & bennett 2013). land-use within the pas is categorised by minimal anthropogenic disturbance, with the areas being managed for the persistence of biodiversity. the crs are extractive reserves where resources such as firewood are harvested, and the native herbivores have largely been replaced with livestock, mainly cattle. several different ecological measures are compared between pas and the surrounding crs: species richness, abundance, species assemblage composition and functional differences. this work thus provides valuable insights into understanding how well pas conserve various measures of biodiversity compared to slightly degraded unprotected habitats. it aids in the growing understanding of how pas and their surrounding matrix areas can function in unison to conserve various biodiversity features and highlights some of the pressures faced by biodiversity in and around the exemplar kruger national park. methods and materials study region surveys were conducted in the lowveld savanna of south africa around the orpen region of the kruger national park and manyeleti game reserve, and adjacent crs belonging to the mnisi tribal authority (figure 1). three sites were chosen for each of two different land-use types: pas and crs. of the three study sites in the pas, two were in the kruger national park – one near the orpen rest camp and the other in the kingfisherspruit section – and the third in the manyeleti game reserve near the khokomya dam. communal rangeland sites were located south-west of kruger and manyeleti and consisted of three cattle grazing camps: shorty, athol and gottenburg. each cr site is associated with a village and stocked with varying densities of cattle, and occasionally goats are allowed into the camps to browse (table 1-a1). unlike the surrounding villages, the cr grazing camps have experienced no habitat loss per se. they are solely used for grazing and almost exclusively support native vegetation. we therefore consider the crs to be degraded and not transformed (rouget et al. 2006). figure 1: map showing the study area. (a) south africa, (b) the north-eastern region of south africa around the kruger national park (in grey) and (c) the locations of the three sites within the protected areas (kruger national park and manyeleti game reserve) and outside the protected areas in the communal rangelands (athol, gottenburg and shorty). the brown colour represents the extent of gabbro soil intrusion in the area (mpumalanga department of agriculture and land administration 1997). at each site, three transects were set up to sample avian diversity and vegetation characteristics. dark grey areas represent government-run protected areas, light grey areas represent privately run protected areas and areas in grey hatching represent cattle camps in communal rangelands used in this study. areas in white represent communally managed areas. all sites were located in the gabbro grassy bushveld vegetation type, characterised by rich clayey soils, open-savanna, dense low-growing grass cover and scattered trees and shrubs (mucina & rutherford 2006). all sites in pas and crs were selected to occur on the timbavati gabbro intrusion; this was confirmed by using google earth imagery (figure 1). all site selection was primarily based on the presence of entry gates within crs and waterpoints within pas. communal rangeland sites were selected based on whether tribal leaders granted the researchers access to their rangeland. pa site selection was chosen based on proximity to the orpen gate to ensure bird surveys were completed before 08:30 each morning. the direction of each transect varied depending on the positions of the gates (in crs), the position of roads (in pas) and to ensure positioning on the gabbro soils. the sites in the pas represent relatively pristine ecosystems that harbour a full set of native herbivores, and regular fires are used as a management strategy. in contrast, crs are characterised by natural vegetation where wild herbivores have been excluded and replaced with cattle. low densities of goats occasionally stray into the camps, where their activities are restricted to areas close to the gates. fire has largely been excluded from crs, although some wood extraction takes place in the area. the maximum distance between two study sites, namely kingfisherspruit and athol, was 30 km and the minimum distance was 4 km (between athol and shorty); thus all sites experience similar climatic conditions. bird sampling all surveys were conducted during january and february 2017; therefore, our findings are representative of summer bird communities. at each of the six sites, three 1-km transects were laid out (bibby, jones & marsden 2000). at each site, the three transects sampled radiated outwards from a focal area of high herbivore activity (i.e. a water point within pas and a gate from which cattle could actively enter the crs) (figure 1-a1). the starting points of transects within a site were at least 150 m apart from one another, and distances between transects increased as one moved further along the transects, as the transects radiated out from one another (figure 1-a1). the transects were set up to radiate out from focal points, which experience high herbivore activity, to areas of lower herbivore activity, to ensure the highest possible similarity in herbivore density for all transects. each transect was walked twice on two different mornings and all birds along the transects were recorded. before starting a transect, the observers waited at the start of the transect for 2 minutes to allow birds to become accustomed to their presence (bibby et al. 2000). then a steady pace was walked for the length of the transect (bibby et al. 2000). the time it took to walk each transect was standardised to not exceed 60 minutes to ensure an equal sampling effort. the average time per transect was 51 minutes. a compass was used to ensure a constant direction was maintained, especially when the surrounding vegetation became thick. all bird observations were made by two observers (m.a.h. and m.d.v.). all birds that were seen or heard on either side of the transect were recorded (bibby et al. 2000). the distance from the observer to the bird was visually estimated and the angle between the bird and the transect was measured using a compass (bibby et al. 2000). both observers’ estimates were used to come to an estimated consensus on distance and angle. when birds of the same species occurred in parties, all individuals of the party were counted and recorded as one sighting, with the distance and angle being estimated to the centre of the party to ensure independence of observations (bibby et al. 2000). the estimated distance from the observer and the angle from the transect line were used to calculate the perpendicular distance from the bird to the transect using pythagoras theorem (bibby et al. 2000). only sightings within the 50 m limit of the transect line were retained (bibby et al. 2000). surveys were conducted on mornings between 06:00 and 08:30, during peak bird activity and vocalisation. surveys were only conducted in mild weather; no transects were surveyed on mornings with high winds or heavy rain. birds flying over the transect were not recorded (bibby et al. 2000). the order of transect surveys was randomised as far as possible by land-use type. in addition, repeat samples of the same transect were temporally spaced out over the sampling period (i.e. not conducted on consecutive mornings) between the sampling dates of 09 january and 02 february 2017. vegetation sampling the composition of avian assemblage in south african savannas is affected by differences in grass and woody vegetation structure (hudson & bouwman 2007; krook, bond & hockey 2007; skowno & bond 2003). thus, structural vegetation attributes were recorded for the first 500 m of each transect. because of time constraints, only the first 500 m of each transect could be sampled. structural changes in vegetation along the transects happened mostly at the start of the transects close to the water source or gate so that the vegetation measures generated within the first 500 m were fairly representative of the entire transect (figures 2-a1 and figure 3-a1). every 10 m along the transect, a 0.5 m × 0.5 m plot was placed on the ground and the maximum leaf table height (the height of the tallest grass leaf material) of each grass species was recorded, from which the average maximum leaf table height per transect was calculated. in addition, at each plot along the transect all woody individuals greater than 0.5 m in height, within a 2 m radius from the centre of the plot, were recorded, from which the average density of woody individuals per transect was calculated (voysey 2018). analyses sampling adequacy and species diversity the data for the two repeat surveys of each transect were pooled before all analyses. to determine sampling adequacy, transect-based species accumulation curves were constructed (gotelli & colwell 2001). jackknife 1 and jackknife 2 estimated species richness of the pas and crs were calculated to get an estimate of the total species richness by accounting for undetected species. generalised linear mixed models (glmms) were used to test whether land-use type, that is pas versus crs, affect the species richness and overall abundance of birds. structural vegetation variables (average maximum grass leaf table height of the first 500 m of the transect, and average density of woody vegetation above 0.5 m in height of the first 500 m of the transect) were incorporated as predictor variables in the models. interaction terms of the two structural vegetation variables with land-use were included in the models to determine if their effect depended on land-use. site was included as a random variable (mcculloch 1996). models were fitted using a poisson distribution and a log-link function (zuur et al. 2009). a full model using all predictor variables was created (global model), from which the best subset model, based on the lowest akaike information criterion (aic), was obtained using the dredge function in r (burnham & anderson 2002). bias in abundance or density estimates may occur because of differences in species detectability between land-use types; we thus tested whether land-use influenced species detectability (coetzee & chown 2016; greve et al. 2011; thomas et al. 2010) (figures 4-a1 and 5-a1). since several species were recorded rarely, it was impossible to create species detectability curves for every species. therefore, a surrogate species approach, where similar species were grouped together, was used (coetzee & chown 2016; greve et al. 2011; thomas et al. 2010), using the same seven groups identified for the region by coetzee and chown (2016). a species detectability curve could not be fit for surrogate group 1 in crs, that is large-bodied ground nesters, because only two observations were made in crs. a pearson correlation analysis was conducted to assess the goodness of fit between true and estimated density for each surrogate bird group, in each of the land-use types, following coetzee and chown (2016). this enabled us to determine if there were any differences in detectability between land-uses or the different surrogate bird species groups. assemblage composition to compare bird species composition between the two land-use types, three approaches were used. first, the number of species that were shared and were unique to each land-use was calculated. secondly, differences in bird composition between pas and crs were explored using non-metric multi-dimensional scaling (nmds) plots based on species identity and abundance per transect. species recorded only once were removed and the remaining data were plotted using a bray–curtis distance measure. random restarts were used to obtain the best possible fit for the data in reduced dimensions. a stress plot was created to determine if the representation of the data in reduced dimensions was good or not. finally, a permutational multivariate analysis of variance (permanova) was used to test whether land-use was a good predictor of bird species composition. functional characteristics body mass is an important characteristic of animals that correlates with many traits, such as reproduction rates and population densities (brose et al. 2017), and often determines the level of exploitation experienced by a species (brose et al. 2017; cullen et al. 2000; pimm et al. 2014). therefore, we tested whether the median body mass of birds differed between land-uses (pas vs. crs). as the body mass distribution for birds is typically skewed, the median body mass is a more appropriate measure than the mean for analysing body mass data of assemblages (meiri & thomas 2007). therefore, median bird mass was calculated per transect-based on the mass of each individual bird observed per transect in both surveys. adult body masses for all species were obtained from greve et al. (2008). we then tested whether median bird body mass per transect differed between land-use types (i.e. pas vs. crs) using a glmm that was fitted with a gaussian distribution and a log-link function (zuur et al. 2009). site was included as a random variable. woody and grass structural vegetation attributes were additionally included as predictor variables in the model. the best subset model selection followed the same procedure as above (zuur et al. 2009). next, birds were assigned to one of five feeding guilds (carnivore, frugivore, mixed feeder, granivore or insectivore) based on diet information that was obtained from hockey, dean and ryan (2005) and categorised by jamison (2017). additionally, species were categorised into four habitat preference classes based on whether they spend the majority of their time in open grassy areas, closed wooded areas or a mixture of both of these habitats. category 1 represents grassland/open-savanna specialists and category 4 represents closed-savanna/thicket/forest specialists. category 2 and 3 species associate with a mixture of the two types of habitats, however, associating with more open (category 2) or more closed (category 3) habitats, respectively. this categorisation was based on habitat association information from hockey et al. (2005), following the categorisation of péron and altwegg (2015), except that categories were scaled the other way around. compositional analyses (aitchison 1982) were performed to test whether the compositions of the feeding guilds and habitat preference categories differed between pas and crs. in this analysis, each transect was treated as a vector (i.e. all five classes of feeding guilds [or habitat preference categories] of one transect represented a single composition). the composition, that is the proportion of each class within feeding guilds (or habitat preference categories), was calculated for each transect, per land-use type, within a multi-dimensional space, rather than as an arithmetic mean of each class individually. separate cluster analyses were conducted for feeding guild and habitat preference category compositions to compare how similar pa and cr transects were in terms of their feeding guild and habitat preference category compositions, respectively. cluster analyses grouped transects based on the similarity of their functional compositions in the multi-dimensional euclidean space. vegetation structure general linear mixed models (lmms) were run to assess whether tree density and maximum leaf table height differed between land-use types (zuur et al. 2009). for these analyses, tree density and maximum leaf table height were averaged per transect prior to analyses. site was included as a random variable. all analyses were conducted in r, version 3.2.6 (r core team 2018), using the following packages for the various analyses: vegan (oksanen et al. 2017) for rarefaction curves, species richness estimates and nmds plot; distance (miller 2017) for detection curves and abundance estimates; lme4 (bates et al. 2015) for lmms and glmms; mumin (barton 2017) for best subset model based on aic; and compositions (van den boogaart, tolosana & bren 2014) for the compositional analysis framework. ethical considerations the authors do not have any ethical considerations that need to be disclosed for the publication of this article. results species diversity the transect-based species accumulation curves for both pa and cr transects began flattening off, though neither reached an asymptote (figure 6-a1), indicating that several species remained unsampled. therefore, the raw species richness values for both pas and crs should be treated with some caution. recorded species richness was larger for pas (94) than for the crs (74). jackknife species richness estimates indicated that estimated total species richness was higher within pas (jackknife 1: 124.2 ± 11.7; jackknife 2: 137.1) than within crs (jackknife 1: 105.1 ± 13.1; jackknife 2: 124.8). a significant correlation was seen between the observed density and modelled density of birds for each surrogate group and each land-use (pearson’s r = 0.989; p < 0.001; n = 13) (figure 7-a1). as a result, we consider the influence of species detectability and the detectability between land-uses as negligible in our study; thus, abundances were not adjusted for detectability (see coetzee & chown 2016; greve et al. 2011; thomas et al. 2010). the effect of land-use type on species richness was significant, but dependent on grass height (table 1). species richness did not change with grass height in pas, but in crs it increased with grass height (figure 8-a1). in addition, species richness was significantly inversely related to increased levels of woody plant density greater than 0.5 m in height, for both crs and pas (table 1, figure 9-a1). the abundance of birds was significantly higher in pas than in crs (figure 2a; table 2-a1). in addition, bird abundance was significantly inversely related to increased grass height (figure 10-a1; table 2-a1). figure 2: (a) boxplot comparing the abundances (counts) of birds per transect survey between land-use types (protected areas vs. communal rangelands). (b) non-metric multi-dimensional scaling ordination plot of bird assemblages in different land-use types. the stress value for the ordination was 0.15, indicating that the projected distances in the ordination diagram were reliable. each point represents one transect; different letters represent different sites. numbers represent the three different transects of each site. (c) boxplot comparing median body masses of birds recorded per transect between land-use types. median body mass was calculated by pooling both bird surveys of each transect. table 1: results from the best subset generalised linear mixed effects model assessing the effect of land-use types (protected areas versus communal rangelands) on raw avian species richness, while taking vegetation structure variables (maximum grass leaf table height and the density of woody vegetation > 0.5 m) into consideration. assemblage composition of the 117 species recorded, 48 were common to both land-uses, of which the most common species were the rattling cisticola (cisticola chiniana), cape turtle dove (streptopelia capicola) and white-browed scrub robin (cercotrichas leucophrys). forty-four species were unique to pas, most commonly senegal lapwing (vanellus lugubris), swainson’s spurfowl (pternistes swainsonii) and magpie shrike (corvinella melanoleuca). in contrast, only 25 species were unique to crs (figure 3; table 3-a1), most commonly white-bellied sunbird (cinnyris talatala), western cattle egret (bubulcus ibis) and sombre greenbul (andropadus importunus). avian species assemblage composition differed substantially between pas and crs (figure 2b). the permanova showed that pas and crs had significantly different avian species assemblages (global r = 0.335; p < 0.001). figure 3: venn diagram showing the number of bird species that are shared and are unique to protected areas and communal rangelands. additionally, the percentage of birds in each habitat preference category is shown for each land-use type. category 1 = birds that associate with open grassy areas; category 2 = birds that associate with open grassy and closed-canopy wooded areas, but have a greater affinity for open grassy areas; category 3 = birds that associate with open grassy areas and closed-canopy wooded areas, but have a greater affinity for closed-canopy wooded areas and category 4 = birds that associate with closed-canopy wooded habitats. an almost equal number of plant species were shared between pas and crs as were unique to either pas or crs (figure 11-a1). however, of the 88 species unique to either pas or crs, 30 were recorded only once. functional composition the median body mass of birds was greater in pas than in crs (figure 2c, table 4-a1). no clear compositional grouping of transects by land-use types (pas vs. crs) existed based on feeding guild compositions (figure 4a). in contrast, cluster analysis grouped communities from pas and crs into two distinct clusters based on bird habitat preference composition (figure 4b). protected areas had higher proportions of category 1 birds (open grassy habitat specialists, e.g. senegal lapwing [vanellus lugubris] and african pipit [anthus cinnamomeus]), while crs had higher proportions of category 4 birds (closed-canopy thicket/woodland habitat specialists, e.g. white-browed scrub robin [cercotrichas leucophrys] and sombre greenbul [andropadus importunus]) (figure 12-a1). indeed, of the species unique to pas, 37% were category 1 birds, while in crs this was only 16% (figure 3). in contrast, pas had 15% and crs had 36% of their unique species classified as category 4 species. figure 4: result of the cluster analysis based on (a) feeding guild composition and (b) habitat preference category compositions. groupings were based on euclidean distances between transects. protected areas are indicated with grey diamonds and communal rangelands are indicated with black circles. vegetation structure tree density was significantly higher in crs (mean = 3050 trees per hectare) than in pas (mean = 884 trees per hectare) (chi-square = 5.62, p = 0.018, df = 1). however, maximum leaf table height did not differ between land-use types (chi-square = 0.42, p = 0.52, df = 1). maximum leaf table height in crs was 12.66 cm ± 0.31 (mean ± se) and in pas it was 13.23 ± 0.43. discussion we show that even when habitat change outside pas involves habitat degradation instead of habitat loss because of transformation, it can still affect bird assemblages. bird assemblages in pas were richer, with greater abundances and different composition than assemblages in adjacent crs. in addition, pas supported larger bodied species and birds of different habitat preference categories, although there were no differences in the composition of feeding guilds between pas and crs. the differences in bird assemblages inside and outside pas indicate either a significant relationship between formal conservation designation and assemblage structure or reflect declines in the favourability of conditions outside pas (e.g. homogenisation of vegetation structure). in the kruger to canyons biosphere reserve, within which this study was conducted, human settlements have expanded by 36%, which has led to a 27% loss of intact savanna vegetation and a 30% increase in thicket vegetation (coetzer et al. 2010). this will have affected bird assemblage composition. floristically, areas within and outside the pas surveyed here, are similar (voysey 2018), yet structurally the areas outside the pas have experienced significant levels of woody thickening over the last 70 years (stevens et al. 2016), which has resulted in a homogenisation of the vegetation structure in crs. elsewhere, structural variation of both woody and grassy vegetation is the most important determinant of savanna avian assemblages (hudson & bouwman 2007; krook et al. 2007; sirami et al. 2009; skowno & bond 2003), and may explain why the more open and structurally heterogeneous vegetation of the pas in this study supported a greater bird species richness, distinct species assemblage composition and a greater incidence of open habitat bird species. the greater avian diversity seen within the structurally more diverse pas may partially be because of the availability of more niches associated with variable vegetation structure (hudson & bouwman 2007). the pas showed more variation in grass height compared to crs, where grass was constantly kept short, and showed little variation in grass height because of constant grazing by cattle in these areas (figure 2-a1) (voysey 2018). this may explain why increasing grass height had a positive impact on species richness outside pas, but no effect within pas: increasing grass height in crs could provide new niches for birds, but more of the same niches in pas. other studies have shown that areas with heterogeneous grass height support more and different grassland bird species than areas that are heavily grazed to homogeneous short grass heights (dias, bastazini, & gianuca 2014; jacoboski, paulsen & hartz 2017). increased woody vegetation density resulted in declines in the proportion of birds that associate with open grassy areas (category 1) and an increase in the proportion of birds that associate with closed-canopy woody vegetation (category 4). the loss of large open grassy patches outside the pas because of the corresponding increases in woody vegetation and its effect on homogenising vegetation structure with crs may explain the observed differences in the proportions of habitat preference categories between the two land-uses. this is in accordance with other studies that have found that woody thickening results in losses of grassland bird species which require open vegetation patches, while benefiting woodland species (sirami et al. 2009; skowno & bond 2003). indeed, closed-habitat specialists have increased across south africa and its bordering countries: passerines that are open-savanna specialists have been declining, while closed-savanna specialists have increased across the region, particularly in the eastern parts of south africa (péron & altwegg 2015), where bush encroachment is considered a serious threat (stevens et al. 2016). increased pressures on the avifauna (especially in the form of hunting) may have further contributed to the differences in bird assemblages inside and outside pas. the crs are extractive reserves for the surrounding villages, as such birds are actively hunted both with slingshots and traps (vincent khosa, pers. obs.; pers. comm., 18 january 2017). elsewhere, hunting in communal areas has caused significant declines in bird numbers (krook et al. 2007). it appears that people preferentially hunt for large-bodied ground nesters (surrogate species group 1, sensu coetzee & chown 2016) (e.g. crested francolin [peliperdix sephaena] and helmeted guineafowl [numida meleagris]) as only two observations of this species group were made within crs, compared to the 117 observations in pas. indeed, larger bodied species are usually targeted disproportionately by hunting (coetzee & chown 2016; cullen et al. 2000; pimm et al. 2014). thus, hunting pressure may not only contribute to a reduction in avian species richness and abundances, but may also explain why the median body mass of birds was higher in pas than in crs. our results suggest that, because of the reduced human pressures within their boundaries, the pas in this study are acting as important population source areas for some bird species that are exploited for hunting in the crs, and possibly also for open habitat bird species (hansen 2011). however, the crs also support a rich diversity of savanna bird species (table 3-a1). indeed, heavier degradation and more intensive land-use have much greater detrimental impacts on biodiversity (greve et al. 2011; newbold et al. 2015). therefore, the contribution of these areas, along with other unprotected areas globally, to the conservation of birds should not be underestimated (cox & underwood 2011). while our study assessed the ecological effectiveness of pas, it did not assess effectiveness in the broader sense of the word. overall effectiveness of pas not only assesses pa contribution to biodiversity conservation, but also considers ecosystem services and social and cultural benefits to communities in and around pas (coad et al. 2008; pringle 2017). adjacent to the pas considered in this study, community benefits of pas are fairly limited; as a matter of fact, some communities incur costs as a result of the pas (anthony 2007; spenceley & goodwin 2008; swemmer, mmethi & twine 2017). therefore, ecological effectiveness does not automatically translate into other aspects of pa effectiveness. in a world of increasing population size and per capita food consumption, the need for agriculture, including pastoralism, has never been greater to ensure the well-being of rural communities. hence, the need to ensure both biodiversity conservation and livelihoods is often considered in assessing pa effectiveness (swemmer et al. 2017). upgrading existing protected areas has been suggested as one of the mechanisms by which both biodiversity conservation and livelihood goals can be addressed in and around pas (pringle 2017). some of the eight principles for upgrading pas are already, or could be, implemented in this area. for example conservation policies already incorporate elements of being long-term and local: government provides long-term funding, and some people from the mnisi community are employed by businesses associated with the pas (v. khosa, pers. comm., 18 january 2017), and involving the youth could raise awareness among younger generations. however, the community remains fairly impoverished, and other principles would be more difficult to implement. these include upsizing the kruger national park (knp) (pringle 2017). around the crs sampled here, the number and extent of human settlements, and the density of people living in settlements, has grown substantially over the last three decades (coetzer et al. 2010; giannecchini, twine & vogel 2007). land remains very important for the livelihood of the local community. therefore, it would be difficult to expand the knp in this area because of possible wildlife–human conflicts (anthony, scott & antypas 2010). however, these crs could be viewed as a matrix or buffer area that serves to reduce external pressures on the pa network (pringle 2017). for example, although there are some changes in bird and plant functional groups, composition and structure between the crs and pas, many species are also retained, and this region probably serves as an important area for gene flow and population sustainability for these species. in summary, as upgrading the pa network in this instance might not be a viable option, continued efforts to engage local people from the surrounding areas may be extremely important to see the benefits of, and create ownership for, the pa network in the region (pringle 2017). this may help to ensure that the wider regional portfolio of birds and their habitats are maintained, so that the entire area, not just the pas, serves to represent bird biodiversity and helps ensure the persistence of viable populations into the future. while we found that all measures of ecological effectiveness were higher in the pas compared to surrounding crs, other studies have found that some of these measures, such as species richness, are higher outside than inside pas (coetzee & chown 2016; greve et al. 2011). this does not necessarily mean that these pas are not performing well to represent and conserve biodiversity within their boundaries, but rather that measures of ecological effectiveness are context-and scale-dependent. for example coetzee and chown (2016) suggest that a higher observed bird richness outside the knp is because of the increased vegetation productivity of croplands which experience year-round irrigation, supporting higher numbers of birds than the pa, which experiences large drops in productivity in the dry season. greve et al. (2011) find higher richness of birds outside than inside one pa, but also significant decreases in specialist guilds, at the expense of generalist guilds, indicating that changes in ecological effectiveness should be considered as a product of all measures. even pas that were not necessarily established to protect all of the biodiversity of an area still function as valuable repositories for biodiversity conservation (greve et al. 2011). finally, scale can affect the direction of differences in diversity between regions: for example areas with high spatial and temporal beta diversity will show low diversity if measured at local scale, but high diversity if measured at regional scale (chiarucci et al. 2012). a number of limitations should be considered for this study. we have mainly attributed differences in bird assemblages in pas and crs to different land management types. we are confident that land-use plays an important role in explaining these differences. however, some differences in bird assemblages and vegetation characteristics of pas and crs may have been driven by the fact that animal densities differed between these land-use types (voysey 2018), which was impossible to fully control in this study. furthermore, our study was conducted in one location only; therefore, other regions or land-use types may have different outcomes (coetzee & chown 2016; greve et al. 2011). additionally, we assessed impacts of degradation in only one location. other regions may show different trends, depending on land-use changes and community characteristics (coetzee & chown 2016; greve et al. 2011). finally, we assessed only mid-summer bird assemblages; winter and spring assemblages may show different responses to land-use. of all the species we recorded, 17% were migrants. conclusion in summary, landscape degradation, though not as extreme a process as landscape transformation, leads to changes in bird assemblages. nevertheless, degraded habitats remain important habitats for a range of bird species, and their contribution to the protection of avian diversity should not be underestimated. acknowledgements the authors thank the south african national parks and mpumalanga tourism and parks agency for providing them with access to the research sites within the protected areas. a special thanks to field rangers thomas rikombe and happy magena. tribal leaders of the mnisi tribal authority are thanked for granting access to the three communal rangeland sites. a special thanks to vincent khosa for his support as field assistant in the communal rangeland sites. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions m.a.h. led all aspects of the project. the project was conceptualised by m.g., who was involved in all aspects of the project. m.d.v. contributed to experimental design, advice regarding the project, fieldwork and analyses. s.-l.j. contributed to the data. all authors contributed to the writing of the manuscript. funding information this work was funded by the south african national research foundation (grant no. 98889 and 110734). data availability statement data is available on request from the corresponding author. disclaimer the views and opinions expressed in this article are those of the author(s) and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references aitchison, j., 1982, ‘the statistical analysis of compositional data’, journal of the royal statistical society: series b 44(2), 139–177. https://doi.org/10.1111/j.2517-6161.1982.tb01195.x andam, k.s., ferraro, p.j., pfaff, a., sanchez-azofeifa, g.a. & robalino, j.a., 2008, ‘measuring the effectivness of protected area networks in reducing deforestation’, proceedings of the national academy of sciences of the united states of america 105(42), 16089–16094. https://doi.org/10.1073/pnas.0800437105 anthony, b.p., 2007, ‘the dual nature of parks: attitudes of neighbouring communities towards kruger national park, south africa’, environmental conservation 34(3), 236–245. https://doi.org/10.1017/s0376892907004018 anthony, b.p., scott, p. & antypas, a., 2010, ‘sitting on the fence? policies and practices in managing human-wildlife conflict in limpopo province, south africa’, conservation and society 8(3), 225–240. https://doi.org/10.4103/0972-4923.73812 balmford, a., green, j.m.h., anderson, m., beresford, j., huang, c., naidoo, r. et al., 2015, ‘walk on the wild side: estimating the global magnitude of visits to protected areas’, plos biology, 13(2), e1002074. https://doi.org/10.1371/journal.pbio.1002074 barton, k., 2017, mumin: multi-model inference, r package, version 1.40.0., viewed 15 october 2017, from https://cran.r-project.org/package=mumin bates, d., maechler, m., bolker, b. & walker, s., 2015, ‘fitting linear mixed-effects models using lme4’, journal of statistical software 67(2015), 1–48. https://doi.org/10.18637/jss.v067.i01 bibby, c., jones, m. & marsden, s., 2000, expedition field techniques: bird surveys, birdlife international, cambridge. brose, u., blanchard, j.l., eklof, a., galiana, n., hartvig, m., kalinkat, r.h.m.g. et al., 2017, ‘predicting the consequences of species loss using size-structured biodiversity approaches’, biological reviews 92(2), 684–697. https://doi.org/10.1111/brv.12250 burnham, k.p. & anderson, d.r., 2002, model selection and mulitmodel inference, 2nd edn., springer-verlag, new york. caro, t., gardner, t.a., stoner, c., fitzherbert, e. & davenport, t.r.b., 2009, ‘assessing the effectiveness of protected areas: paradoxes call for pluralism in evaluating conservation performance’, diversity and distributions 15(1), 178–182. https://doi.org/10.1111/j.1472-4642.2008.00522.x chape, s., harrison, j., spalding, m. & lysenko, i., 2005, ‘measuring the extent and effectiveness of protected areas as an indicator for meeting global biodiversity targets’, philosophical transactions of the royal society b 360(1454), 443–455. https://doi.org/10.1098/rstb.2004.1592 chiarucci, a., bacaro, g., filibeck, g., landi, s., machcherini, s. & scoppola, a., 2012, ‘scale dependence of plant species richness in a network of protected areas’, biodiversity and conservation 21(2), 503–516. https://doi.org/10.1007/s10531-011-0196-8 chown, s.l., 2010, ‘temporal biodiversity change in transformed landscapes: a southern african perspective’, philosophical transactions of the royal society b 365(1558), 3729–3742. https://doi.org/10.1098/rstb.2010.0274 coad, l., campbell, a., miles, l. & humphries, k., 2008, the costs and benefits of protected areas for livelihoods: a review of the current literature, working paper, unep world conservation monitoring centre, cambridge, uk. coetzee, b.w.t. & chown, s.l., 2016, ‘land-use change promotes avian diversity at the expense of species with unique traits’, ecology and evolution 6(21), 7610–7622. https://doi.org/10.1002/ece3.2389 coetzee, b.w.t., gaston, k.j. & chown, s.l., 2014, ‘local scale comparisons of biodiversity as a test for global protected area ecological performance: a meta-analysis’, plos one 9(8), e105824. https://doi.org/10.1371/journal.pone.0105824 coetzer, k.l., erasmus, b.f.n., witkowski, e.t.f. & bachoo, a.k., 2010, ‘land-cover change in the kruger to canyons biosphere reserve (1993–2006): a first step towards creating a conservation plan for the subregion, south african journal of science 106(7/8), 1–10. https://doi.org/10.4102/sajs.v106i7/8.221 cox, r.l. & underwood, e.c., 2011, ‘the importance of conserving biodiversity outside protected areas in mediterranean ecosystems’, plos one 6(1), e14508. https://doi.org/10.1371/journal.pone.0014508 cullen, l., bodmer, r.e. & pádua, c.v., 2000, ‘effects of hunting in habitat fragments of the atlantic forests, brazil’, biological conservation 95(1), 49–56. https://doi.org/10.1016/s0006-3207(00)00011-2 dias, r.a., bastazini, v.a.g. & gianuca, a.t., 2014, ‘bird-habitat associations in costal rangelands of southern brazil’, iheringia série zoologia 104(1), 200–208. https://doi.org/10.1590/1678-476620141042200208 dudley, n., hockings, m. & stolton, s., 2010, ‘precious places: getting the arguments right’, in s. stolton & n. dudley (eds.), arguments for protected areas: multiple benefits for conservation and use, pp. 253–265, earthscan ltd, london. foley, j.a., defries, r., asner, j.p., barford, c., bonan, g., carpenter, s.r. et al., 2005, ‘global consequences of land use’, science 309(5737), 570–574. https://doi.org/10.1126/science.1111772 gardner, t. a., caro, t., fitzherbert, e.b., banda, t. & lalbhai p., 2007, ‘conservation value of multiple-use areas in east africa’, conservation biology 21(6), 1516–1525. https://doi.org/10.1111/j.1523-1739.2007.00794.x gaston, k.j., charman, k., jackson, s.f., armsworth, p.r., bonn, a., briers, r.a. et al., 2006, ‘the ecological effectiveness of protected areas: the united kingdom’, biological conservation 132(1), 76–87. https://doi.org/10.1016/j.biocon.2006.03.013 gaston, k.j., jackson, s.f., cantú-salazar, l. & cruz-piñón, g., 2008, ‘the ecological performance of protected areas’, annual review of ecology, evolution, and systematics 39(1), 93–113. https://doi.org/10.1146/annurev.ecolsys.39.110707.173529 geldmann, j., barnes, m., coad, l., craigie, i.d., hockings, m. & burgess, n.d., 2013, ‘effectivness of terrestrial protected areas in reducing habitat loss and population declines’, biological conservation 16(1), 230–238. https://doi.org/10.1016/j.biocon.2013.02.018 gerland, p., raftery, a.e., ševčíková, h., li, n., gu, d., spoorenberg, t. et al., 2014, ‘world population stabilization unlikely this century’, science 346(6206), 234–239. https://doi.org/10.1126/science.1257469 giannecchini, m., twine, w. & vogel, c., 2007, ‘land-cover change and human-environment interactions in a rural cultural landscape in south africa’, the geographical journal 173(1), 26–42. https://doi.org/10.1111/j.1475-4959.2007.00227.x gotelli, n. & colwell, r.k., 2001, ‘quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness’, ecology letters 4(4), 379–391. https://doi.org/10.1046/j.1461-0248.2001.00230.x gray, c.l., hill, s.l., newbold, t., hudson, l.n., borger, l., contu, s. et al., 2016, ‘local biodiversity is higher inside than outside terrestrial protected areas worldwide’, nature communications 7, 12306. https://doi.org/10.1038/ncomms12306 greve, m., chown, s.l., van rensburg, b.j., dallimer, m. & gaston, k.j., 2011, ‘the ecological effectiveness of protected areas: a case study for south african birds’, animal conservation 14(3), 295–305. https://doi.org/10.1111/j.1469-1795.2010.00429.x greve, m., gaston, k.j., van rensburg, b.j. & chown, s.l., 2008, ‘environmental factors, regional body size distributions and spatial variation in body size of local avian assemblages’, global ecology and biogeography 17(4), 514–523. https://doi.org/10.1111/j.1466-8238.2008.00388.x hansen, a.j., 2011, ‘contribution of source-sink theory to protected area science’, in j. liu, v. hall, a. morzillo & j. wiens (eds.), sources, sinks and sustainability across landscapes, pp. 339–360, cambridge university press, cambridge. hockey, p.a.r., dean, w.r.j. & ryan, p.g., 2005, roberts birds of southern africa, trustees of the john voelcker bird book fund, cape town. hoffmann, m., hilton-taylor, c., angulo, a., bohm, m., brooks, t.m., butchart, s.h. et al., 2010. ‘the impact of conservation on the status of the world’s vertebrates’, science 330(6010), 1503–1509. https://doi.org/10.1126/science.1194442 hudson, a. & bouwman, h., 2007, ‘different land-use types affect bird communities in the kalahari, south africa’, african journal of ecology 45(3), 423–430. https://doi.org/10.1111/j.1365-2028.2006.00750.x jackson, s.f., evans, k.l. & gaston, k.j., 2009, ‘statutory protected areas and avian species richness in britain’, biodiversity and conservation 18(8), 2143–2151. https://doi.org/10.1007/s10531-009-9578-6 jacoboski, l.i., paulsen, r.k. & hartz, s.m., 2017, ‘bird-grassland associations in protected and non-protected areas in southern brazil’, perspectives in ecology and conservation, 15(2), 109–114. https://doi.org/10.1016/j.pecon.2017.05.002 james, a.n., gaston, k.j. & balmford, a., 1999, ‘balancing the earth’s accounts’, nature 401(6751), 323–324. https://doi.org/10.1038/43774 jamison, s.-l. 2017, ‘unlocking bird-tree relationships: exploring biogeographical patterns across southern africa’, msc thesis, dept. of plant and soil sciences, university of pretoria. joppa, l.n., loarie, s.r. & pimm, s.l., 2008, ‘on the protection of “protected areas”’, proceedings of the national academy of sciences of the united states of america 105(18), 6673–6678. https://doi.org/10.1073/pnas.0802471105 joppa, l.n. & pfaff, a., 2009, ‘high and far: biases in the location of protected areas’, plos one 4(12):e8273. https://doi.org/10.1371/journal.pone.0008273 krook, k., bond, w.j. & hockey, p.a.r., 2007, ‘the effect of grassland shifts on the avifauna of a south african savanna’, ostrich 78(2), 271–279. https://doi.org/10.2989/ostrich.2007.78.2.24.104 laurance, w.f., camargo, j.l.c., luizão, r.c.c., laurance, s.g., pimm, s.l., bruna, e.m. et al., 2011, ‘the fate of amazonian forest fragments: a 32-year investigation’, biological conservation 144(1), 56–67. https://doi.org/10.1016/j.biocon.2010.09.021 laurance, w.f., useche, d.c., rendeiro, j., kalka, m., bradshaw, c.j., sloan, s.p. et al, 2012, ‘averting biodiversity collapse in tropical forest protected areas’, nature 489(7415), 290–294. longcore, t. & rich, c., 2004, ‘ecological light pollution’, frontiers in ecology and the environment 2(4), 191–198. https://doi.org/10.1890/1540-9295(2004)002[0191:elp]2.0.co;2 margules, c.r. & pressey, r.l., 2000, ‘systematic conservation planning’, nature 405(6783), 243–254. https://doi.org/10.1038/35012251 mcculloch, c.e., 1996, ‘an introduction to generalized linear mixed models’, applied statistics in agriculture 10(1), 1–17. https://doi.org/10.4148/2475-7772.1314 meiri, s. & thomas, g., 2007, ‘the geography of body size – challenges of the interspecific approach’, global ecology and biogeography 16(6), 689–693. https://doi.org/10.1111/j.1466-8238.2007.00343.x miller, d.l., 2017, distance: distance sampling detection function and abundance estimation, r package, version 0.9.7., viewed 03 august 2017, from https://cran.r-project.org/package=distance mpumalanga department of agriculture and land administration, 1997, institute for soil, climate and water – soils clay layer for mpumalanga part of the kruger national park (knp), received from sanparks gis lab, september 2018. mucina, l. & rutherford, m.c. (eds.), 2006, the vegetation of south africa, lesotho and swaziland, strelitzia 19, pretoria. newbold, t., hudson, l.n., hill, s.l.l., contu, s., lysenko, i., senior, r.a. et al. 2015, ‘global effects of land use on local terrestrial biodiversity’, nature 520(7545), 45–50. https://doi.org/10.1038/nature14324 oksanen, j., blanchet, f.g., friendly, m., kindt, r., solymos, p., henry, m. et al., 2017, vegan: community ecology package, r package, version 2.4–3., viewed 18 july 2017, from https://cran.r-project.org/package=vegan palmer, a.r. & bennett, j.e., 2013, ‘degradation of communal rangelands in south africa towards an improved understanding to inform policy’, african journal of range and forage science 30(1–2), 57–63. https://doi.org/10.2989/10220119.2013.779596 péron, g. & altwegg, r., 2015, ‘twenty-five years of change in southern african passerine diversity: nonclimatic factors of change’, global change biology 21(9), 3347–3355. https://doi.org/10.1111/gcb.12909 pimm, s. l., jenkins, c.n., abell, r., brooks, t.m., gittleman, j.l., joppa, l.n. et al., 2014, ‘the biodiversity of species and their rates of extinction, distribution, and protection’, science 344(6187), 987–999. https://doi.org/10.1126/science.1246752 pringle, r.m., 2017, ‘upgrading protected areas to conserve wild biodiversity’, nature 546(7656), 91–99. https://doi.org/10.1038/nature22902 r core team, 2018, r: a language and environment for statistical computing, version 3.5.0, computer program, r foundation for statistical computing, vienna, austria. rayner, l., lindenmayer, d.b., wood, j.t., gibbons, p. & manning, a.d., 2014, ‘are protected areas maintaining bird diversity?’, ecography 37(1), 43–53. https://doi.org/10.1111/j.1600-0587.2013.00388.x roser, m. & ritchie, h., 2018, ‘yields and land use in agriculture’, in ourworldindata.org, viewed 29 november 2018, from https://ourworldindata.org/yields-and-land-use-in-agriculture rouget, m., jonas, z., cowling, r.m., desmet, p.g., driver, a., mucina, l. et al. 2006, ‘ecosystem status and protection levels of vegetation types’, in l. mucina & m.c. rutherford (eds.), the vegetation of south africa, lesotho and swaziland, pp. 724–737, strelitzia 19, pretoria. sinclair, a.r., mduma, s.a. & arcese, p., 2002, ‘protected areas as biodiversity benchmarks for human impact: agriculture and the serengeti avifauna’, proceedings: biological sciences 269(1508), 2401–2405. https://doi.org/10.1098/rspb.2002.2116 sirami, c., seymour, c., midgley, g. & barnard, p., 2009, ‘the impact of shrub encroachment on savanna bird diversity from local to regional scale’, diversity and distributions 15(6), 948–957. https://doi.org/10.1111/j.1472-4642.2009.00612.x skowno, a.l. & bond, w.j., 2003, ‘bird community composition in an actively managed savanna reserve, importance of vegetation structure and vegetation composition’, biodiversity and conservation 12(11), 2279–2294. https://doi.org/10.1023/a:1024545531463 spenceley, a. & goodwin, h., 2008, ‘nature-based tourism and poverty alleviation: impacts of private sector and parastatal enterprises in and around kruger national park, south africa’, current issues in tourism 10(2–3), 255–277, https://doi.org/10.2167/cit305.0 stevens, n., erasmus, b.f., archibald, s. & bond, w.j., 2016, ‘woody encroachment over 70 years in south african savannahs: overgrazing, global change or extinction aftershock?’, philosophical transactions of the royal society b 371(1703), 20150437. https://doi.org/10.1098/rstb.2015.0437 swemmer, l., mmethi, h. & twine, w., 2017, ‘tracing the cost/benefit pathway of protected areas: a case study of the kruger national park, south africa’, ecosystem services 28, 162–172. https://doi.org/10.1016/j.ecoser.2017.09.002 thomas, l., buckland, s.t., rexstad, e.a., laake, j.l., strindberg, s. & hedley, s.l., 2010, ‘distance software: design and analysis of distance sampling surveys for estimating population size’, journal of applied ecology 47(5), 5–14. https://doi.org/10.1111/j.1365-2664.2009.01737.x van den boogaart, k.g., tolosana, r. & bren, m., 2014, compositions: compositional data analysis, r package, version 1.40–1, viewed 22 september 2017, from https://cran.r-project.org/package=compositions voysey, m.d., 2018, ‘the influence of wildlife and livestock herbivore assemblages on grazing lawn flora’, msc thesis, dept. of plant and soil sciences, university of pretoria. waldron, a., mooers, a.o., miller, d.c., nibbelink, n., redding, d., kuhn, t.s. et al., 2013, ‘targeting global conservation funding to limit immediate biodiversity declines’, proceedings of the national academy of sciences of the united states of america 10(29), 12144–12148. https://doi.org/10.1073/pnas.1221370110 wegge, p., odden, m., pokharel, c.p. & storaas, t., 2009, ‘predator-prey relationships and responses of ungulates and their predators to the establishment of protected areas: a case study of tigers, leopards and their prey in bardia national park, nepal’, biological conservation 142, 189–202. https://doi.org/10.1016/j.biocon.2008.10.020 zuur, a., ieno, e.n., walker, n., saveliev, a.a. & smith, g.m., 2009, mixed effects models and extensions in ecology with r, springer-verlag, new york. appendix 1 table 1-a1: livestock densities of the three communal rangeland sites. data on livestock densities were obtained from the state veterinary department, originally collected by the mnisi community programme, and represent a five-year mean ± standard error (se) (2012–2016). table 2-a1: results from the generalised linear mixed effects models comparing avian abundances between land-use types (protected areas versus communal rangelands), while taking vegetation structural variables (maximum grass leaf table height and the density of woody vegetation greater than 0.5 meters) into consideration. table 3-a1: cumulative counts of birds surveyed per land-use type in alphabetical order. table 4-a1: results from generalised linear mixed effects models comparing median avian body mass between land-use types (protected areas versus communal rangelands), while taking vegetation structure variables (maximum grass leaf table height and the density of woody vegetation greater than 0.5 meters) into consideration. figure 1-a1: diagram showing layout of three transects at each site. at each site, the three transects were set up within 40 meters from a water point (in protected areas) or a livestock camp gate (in communal rangelands). transects started 150 meters from one another and radiated out from on another. transects were 1000 meters in length. figure 2-a1: scatter plot showing the relationship of average maximum leaf table per plot number of each land-use type (protected areas versus communal rangelands). plot number increases with distance from water source (protected area) or gate (communal rangelands). figure 3-a1: scatter plot showing the average woody density of trees > 0.5 m per plot number of each land-use type (protected areas versus communal rangelands) plot number increases with distance from water source (protected area) or gate (communal rangelands). figure 4-a1: detectability curves for the seven surrogate bird species groups fit for protected areas. surrogate group numbers are indicated in the top right of each figure. (a) group 1 (large-bodied, ground nesters), (b) group 2 (small to medium bodied, mixed feeders, frugivores, hole nesters), (c) group 3 (small to medium bodied, granivores, mainly platform nesters), (d) group 4 (small to medium bodied, parasitic breeders or cup nesters, insectivores, nectarivores), (e) group 5 (small to medium bodied, mainly insectivores, mixed feeders, mixed breeders), (f) group 6 (small bodied, granivores, cup nesters) and (g) group 7 (medium bodied, insectivore/frugivores, mainly hole nesters). see coetzee and chown. (2016:2610-2622) for more details. figure 5-a1: detectability curves for six of the seven surrogate bird species groups fit for communal rangelands. a detectability curve could not be fit for surrogate species (a) group 1 (large-bodied, ground nesters) as only two observations were made for this group in communal rangelands. surrogate species groups are indicated in the top right of each figure. (b) group 2 (small to medium bodied, mixed feeders, frugivores, hole nesters), (c) group 3 (small to medium bodied, granivores, mainly platform nesters), (d) group 4 (small to medium bodied, parasitic breeders or cup nesters, insectivores, nectarivores), (e) group 5 (small to medium bodied, mainly insectivores, mixed feeders, mixed breeders), (f) group 6 (small bodied, granivores, cup nesters) and (g) group 7 (medium bodied, insectivore/frugivores, mainly hole nesters). figure 6-a1: transect-based species accumulation curves using the random method, which assigns transects in a random order to quantify sampling adequacy, for (a) the nine transects in communal rangelands and (b) the nine transects in protected areas. grey shading represents the 95% confidence interval for the extrapolated richness. figure 7-a1: the relationship between the observed and modelled bird densities in protected areas and communal rangelands for seven surrogate species groups after coetzee et al. (2016). black represents surrogate bird species group 1, red – group 2, green – group 3, blue – group 4, turquoise – group 5, pink – group 6, and yellow – group 7. see figure 5-a1 for the definition of groups. the two different land-use types are indicated by different symbols, as shown in the legend. the diagonal line with a slope of one to one shows the hypothetical line of best fit if observed density = modelled density. (pearson’s r = 0.989; p < 0.001). figure 8-a1: interaction plot showing the effects of maximum leaf table height (proxy for grass height), on avian species richness in different land-use types (protected areas versus communal rangelands). light blue shading represents the 95% confidence interval for the fitted model. figure 9-a1: effects plot showing how avian species richness is affected by the density of woody vegetation above 0.5 meters in height. light shaded blue represents the 95% confidence interval for the fitted model. figure 10-a1: effects plot showing the relationship between avian abundance and maximum leaf table height (proxy for grass height). abundances represent pooled abundances of both bird surveys conducted for each transect. light shaded blue represents the 95% confidence interval for the fitted model. figure 11-a1: a venn diagram showing the number of plant (trees, grass and forb) species shared between and unique to protected areas and communal rangelands. figure 12-a1: the compositional mean of each habitat preference category between the two land-uses (protected areas versus communal rangelands). category 1 = birds that associate with open grassy areas, category 2 = birds that associate with open grassy and closed canopy wooded areas, but have a greater affinity for open grassy areas, category 3 = birds that associate with open grassy areas and closed canopy wooded areas, but have a greater affinity for closed canopy wooded areas and category 4 = birds that associate with closed canopy wooded habitats. russell.qxd freshwater fishes of tsitsikamma national park i.a. russell russell, i.a. 2002. freshwater fishes of tsitsikamma national park. koedoe 45(2): 13–17. pretoria. issn 0075-6458. this study aimed to determine the distribution and relative abundance of freshwater fishes in the tsitsikamma national park. fish assemblages in six river systems were sampled in 2001, with a total of 323 fish from eight species recorded. indigenous fish collected included four freshwater species (pseudobarbus afer, pseudobarbus tenuis, sandelia capensis, anguilla mossambica), three estuarine species (monodactylus falciformis, caffrogobius gilchristi, myxus capensis), and one alien (micropterus salmoides). one additional indigenous species (galaxias zebratus) and two aliens (salmo trutta, oncorhynchus mykiss) could potentially occur within the park. the topography and locality of the park presents a unique opportunity to meaningfully conserve the endangered p. tenuis as well as other fish characteristic of the eastern reaches of the cape floristic region. management action is required to minimise opportunities for further establishment and spread of alien fish species and to conserve indigenous fish assemblages within the park. key words: freshwater fish, diversity, alien, conservation, tsitsikamma, keurbooms. i.a. russell, south african national parks, p.o. box 176, sedgefield, 6573, republic of south africa. issn 0075-6458 13 koedoe 45/2 (2002) introduction rivers in the cape floristic region (cfr) of the western and eastern cape provinces of south africa harbour a distinctive fish fauna, characterised with a high level of endemicity (16 of 19 species) (skelton 1987). the freshwater fishes of the cfr are also highly threatened, with 15 species (79 %) being red-data listed, nine of which are classed as either endangered or critically endangered (hilton-taylor 2000), and are reliant on effective conservation of the region for their survival (impson et al. 2001). the tsitsikamma national park (tnp) is located in the eastern reaches of the cfr, with recently acquired and contractually managed areas containing river systems which drain catchments supporting predominantly afromontane forest (von breitenbach 1974) and both wet and mesic mountain fynbos (moll & bossi 1984). the tnp thus has the potential to contribute to the conservation of the ichthyofauna of the cape floristic region. the only published account of the freshwater fishes of tnp lists three species (anguilla mossambica, pseudobarbus asper and sandelia capensis) as occurring in the park (smith & smith 1966). sampling undertaken in rivers in the tsitsikamma area outside the park’s boundaries has revealed a more diverse ichthyofauna (le roi le riche & hey 1947; scott & hamman 1984; skelton 1980; skelton 1988; unpublished records from albany museum and south african institute for aquatic biodiversity). of particular conservation interest is the occurrence of pseudobarbus tenuis, classified as endangered (hilton-taylor 2000), in rivers of the keurbooms system outside the current national park boundaries. the objective of this study was to describe the distribution and relative abundance of freshwater fish in tnp, with particular attention being given to assessing the status of p. tenuis. methods fish sampling was undertaken in february and march 2001. rivers in the tnp are generally small, shallow and with a stony substratum, thus all samrussell.qxd 2005/12/09 10:12 page 13 pling was undertaken using an electro-fishing apparatus. a hand-held electric fish shocker, powered by a 220v ac, 2-kva portable generator was used at sample sites that were located close to access roads. at sites that were far removed from access roads a portable 12v dc electric fish shocker (deka 3000) was used to sample fish, which although less effective than the ac powered unit, was more easily portaged for long distances over broken terrain. all fish collected were identified using identification keys in skelton (1993). voucher specimens were preserved in either 10 % formalin or 70 % alcohol, and housed at the south african institute for aquatic biodiversity in grahamstown (caffrogobius gilchristi = rusi 64253; monodactylus falciformis = rusi 64254; myxus capensis = rusi 64255; anguilla mossambica = rusi 64256; micropterus salmoides = rusi 64257; pseudobarbus asper group = rusi 64258; pseudobarbus afer = rusi 64259-60; pseudobarbus tenuis = rusi 64261-66). results a total of 323 fish from at least eight species was recorded (table 1). all specimens classified as pseudobarbus spp. (table 1) were juveniles that were too small to enable identification to species level based on anatomical features. all unidentified specimens were collected in the lower reaches of the groot (west) river in the presence of adult p. afer. of the eight positively identified species, three (m. falciformis, c. gilchristi, m. capensis) occur predominantly in estuaries or inshore coastal waters although are occasionally found in coastal rivers (skelton 1993), and one species (m. salmoides) is an alien. pseudobarbus afer and s. capensis were collected only in the groot (west) river, whereas p. tenuis was found to be restricted to the keurbooms system. anguilla mossambica was the only freshwater fish recorded in both the sout and brak rivers. discussion published accounts of fish in the keurbooms system, and collections of preserved fish specimens from the groot (west) and keurbooms systems, housed in the albany museum and the south african institute for aquatic biodiversity, indicate the possible occurrence of three additional species in the tsitsikamma national park. the first is the cape galaxias galaxias zebratus, which has been recorded in several tributaries of the keurbooms system, including the palmiet river westward of the soetkraal portion of tnp (fig. 1). skelton (1993) describes g. zebratus as being able to tolerate a wide range of water and temperature conditions, and there is no obvious reason why it should not occur in the eastern tributaries of the keurbooms system in the tsitsikamma national park. cambray et al. (1995) recorded g. zebratus from the upper reaches of the koedoe 45/2 (2002) 14 issn 0075-6458 table 1 number of fish specimens per species collected per site in the tsitsikamma national park during february and march 2001. site numbers as per figure 1. sampling effort = minutes electro-fishing site number 1 2 3 4 5 6 7 8 9 10 11 sampling effort 155 60 10 30 120 45 30 20 40 20 20 total anguilla mossambica longfin eel 12 1 1 4 3 1 22 pseudobarbus afer eastern cape redfin 27 27 pseudobarbus spp. redfin species 124 124 pseudobarbus tenuis slender redfin 16 3 35 54 micropterus salmoides largemouth bass 33 1 34 sandelia capensis cape kurper 2 2 myxus capensis freshwater mullet 20 20 monodactylus falciformis cape moony 33 33 caffrogobius gilchristi prison goby 7 7 total number of individuals 165 17 3 36 64 3 1 0 33 1 0 323 total number of species 3 2 1 2 4 1 1 0 1 1 0 9 russell.qxd 2005/12/09 10:12 page 14 issn 0075-6458 15 koedoe 45/2 (2002) fig 1. location of the tsitsikamma national park relative to major centres (a) and geographical features (b), and location of study sites in the keurbooms, salt, brak and groot (west) rivers (c) and two unnamed rivers in the vicinity of the storms river (d). russell.qxd 2005/12/09 10:12 page 15 krom and gamtoos rivers eastward of tnp. it is thus possible that g. zebratus could also occur in rivers eastward of the keurbooms system in the park. the other two species, which could also potentially occur in tnp by virtue of their having previously been collected in the keurbooms system, are the brown trout salmo trutta, and rainbow trout oncorhynchus mykiss. salmo trutta was described by demoor & bruton (1988) as “thriving” in the keurbooms river despite the supposed intolerance of this species of low ph, peat-stained rivers characteristic of the southern cape coastline. skelton (1994) recorded o. mykiss in the mainstream of the upper reaches of the keurbooms river. the current status of both s. trutta and o. mykiss in the keurbooms river is unknown, though their preference for mountain or upland streams (skelton 1993) means that they pose a threat to invasion of the upper reaches of the palmiet river in the park. this study confirms the findings of surveys undertaken in 1925 (hey 1926) and 2000/2001 (a. bok pers. comm.) that no freshwater fish other than eels occur in the salt river, and suggests that a similar absence occurs in the smaller brak river. the absence of cyprinids from the brak river is possibly a consequence of episodic flow, with at the time of sampling the brak river consisting of a series of isolated stagnant pools, conditions not ideal for pseudobarbus species indigenous to the area (skelton 1993). the absence of most expected freshwater fish, and in particular p. afer, from the salt river, however, is from an ecological perspective difficult to explain. local knowledge indicates that the salt river is perennial, and aquatic habitats in the river appear highly suitable for pseudobarbus species, which are found in similar adjacent rivers. it is unknown what influence this largely ‘fish-less’ state of the salt river has had on other aquatic biota. preliminary surveys of aquatic invertebrates have indicated a diverse community with several endemics, and at least three previously undescribed genera and 11 undescibed species (f. de moor pers. comm.). from a conservation standpoint the temptation to introduce fish, particularly aliens, into the salt river should be resisted in the interests of conserving what is potentially a unique aquatic ecosystem. impson et al. (2001) suggest that national parks in the cfr make a negligible contribution to the conservation of indigenous fishes, with only two species recorded in parks. this study along with recent surveys (russell, 1998, 1999, 2001), however, indicate that rivers in national parks contain populations of at least eight of the 19 fish species indigenous to the cfr, and hence make a moderate contribution to the conservation of these biota. the tnp in particular provides a unique opportunity to meaningfully conserve the endangered (hilton-taylor 2000) p. tenuis. with respect to rivers in the soetkraal section, tnp has several of the attributes identified by skelton et al. (1995) as necessary for effective conservation of riverine fishes, namely (i) it is positioned high in the catchment, (ii) encompasses the entire upper catchment of the affected area, in this case the palmiet river, and (iii) highimpact alien predators do not occur within the reserve. the remaining requirements for effective conservation of (iv) maintaining natural hydrological cycles, and (v) securing the minimum water quantity and quality requirements of the entire biotic community are either already being achieved, or could potentially be achieved through continued eradication of alien plants in the upper palmiet river catchment area. an alien plant removal program operating under the auspices of the working for water program of the south african department of water affairs and forestry has been active in the keurbooms river catchment since 1995, and plans have been formulated to continue the program till at least 2008 (gelderblom & rowlinson 1999) demonstrating an ongoing commitment to rehabilitate this catchment. possibly one of the greatest threats to p. tenuis and other indigenous fishes in the keurbooms system is the introduction or spread of alien predatory fish. salmo trutta and o. mykiss have previously been recorded in the keurbooms river (unpublished koedoe 45/2 (2002) 16 issn 0075-6458 russell.qxd 2005/12/09 10:12 page 16 records from the south african institute for aquatic biodiversity; skelton 1994), and this study has demonstrated that largemouth black bass m. salmoides have become established in adjacent streams. a threat is also posed by the interest shown by landowners adjacent to tnp to continue stocking rivers with o. mykiss (pers obs.). every effort should be made to prevent the further invasion of rivers in tnp by alien fish. precautionary steps taken should include prohibition of all alien species introductions into rivers in the area, and wherever possible, the undertaking of rehabilitation projects involving alien fish removal. furthermore, a barrier weir should be constructed in the palmiet river where it leaves tnp to prevent the movement of alien fish into the park from lower in the river system. acknowledgements conservation staff of tnp are thanked for their assistance during sample collection. special thanks are due to mr. j. allan who facilitated sampling in remote areas in the tsitsikamma mountains, mr. e. kivett for assistance in the field, mr. r. bills for the loan of field equipment, and identification and curation of specimens. staff of the south african institute for aquatic biodiversity, and albany museum, are thanked for the provision of museum records on fish distribution in the keurbooms river catchment. references cambray, j.a., a. bok & r. smith. 1995. range extensions for galaxias zebratus castelnau, 1861 (galaxiidae), krom and gamtoos river systems, eastern cape. south african journal of aquatic science 21 (1/2): 99-102. demoor, i.j. & m.n. bruton. 1988. atlas of alien and translocated indigenous aquatic animals in southern africa. pretoria: council for scientific and industrial research, national scientific programmes unit. (south african national scientific programmes report; no. 144.) gelderblom, c. & l. rowlinson. 1999. management plan for alien vegetation in the keurbooms catchment. pretoria: council for scientific and industrial research. hey, s.a. 1926. fisheries survey 1925-6. inland waters. report no. 1. department of mines and industries. pretoria: government printers. hilton-taylor, c. 2000. 2000 iucn red list of threatened species. gland, switzerland: iucn. impson, n.d., i.r. bills & j.a. cambray. 2001. a conservation plan for the unique and highly threatened freshwater fishes of the cape floral kingdom. pp 432-440. in: collares-pereira, m.j., i.g. cowx & m.m. coelho (eds.). conservation of freshwater fish: options for the future. oxford: blackwell science. le roi le riche, h. & d. hey. 1947. survey of the south western districts. cape town: inland fisheries department, provincial administration of the cape of good hope. (report no 4: 19-27). moll, e.j. & l. bossi. 1984. vegetation map of the fynbos biome. mowbray: director of survey and mapping. russell, i.a. 1998. freshwater fish of the addo elephant national park. south african journal of aquatic science 24 (1/2): 57-70. russell, i.a. 1999. freshwater fish of the wilderness national park. koedoe 42(1): 73-78. russell, i.a. 2001. freshwater fishes of bontebok national park. koedoe. 44(2): 71-77. scott, h.a. & k.c.d. hamman. 1984. freshwater fishes of the cape. cape town: cape department of nature and environmental conservation. (cape conservation series no. 5.) skelton, p.h. 1980. systematics and biogeography of the redfin barbus species (pisces: cyprinidae) from southern africa. unpublished ph.d. thesis. rhodes university, grahamstown. skelton, p.h. 1987. south african red data book – fishes. pretoria: council for scientific and industrial research, national scientific programmes unit. (south african national scientific programmes report; no. 137.) skelton, p.h. 1988. a taxonomic revision of the redfin minnows (pisces, cyprinidae) from southern africa. annals of the cape provincial museum 16(10): 201-307. skelton, p.h. 1993. a complete guide to the freshwater fishes of southern africa. halfway house: southern book publishers. skelton, p.h. 1994. baseline survey of fishes in the keurbooms/bietou and piesang rivers, southern cape. pretoria: department of water affairs and forestry. (plettenberg bay coastal catchment study). skelton, p.h., j.a. cambray, a. lombard & g.a. benn. 1995. patterns of distribution and conservation status of freshwater fishes in south africa. south african journal of zoology 30(3): 711-781. smith, j.l.b. & m.m. smith. 1966. fishes of the tsitsikamma coastal national park. pretoria: national parks board of trustees. von breitenbach, f. 1974. southern cape forests and trees. pretoria: government printers. issn 0075-6458 17 koedoe 45/2 (2002) russell.qxd 2005/12/09 10:12 page 17 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice boshoff.qxd the potential distributions, and estimated spatial requirements and population sizes, of the medium to large-sized mammals in the planning domain of the greater addo elephant national park project a.f. boshoff, g.i.h. kerley, r.m. cowling and s.l. wilson boshoff, a.f., g.i.h. kerley, r.m. cowling and s.l. wilson. 2002. the potential distributions, and estimated spatial requirements and population sizes, of the medium to large-sized mammals in the planning domain of the greater addo elephant national park project. koedoe 45(2): 85–116. pretoria. issn 0075-6458. the greater addo elephant national park project (gaenp) involves the establishment of a mega biodiversity reserve in the eastern cape, south africa. conservation planning in the gaenp planning domain requires systematic information on the potential distributions and estimated spatial requirements, and population sizes of the medium to largesized mammals. the potential distribution of each species is based on a combination of literature survey, a review of their ecological requirements, and consultation with conservation scientists and managers. spatial requirements were estimated within 21 mammal habitat classes derived from 43 land classes delineated by expert-based vegetation and river mapping procedures. these estimates were derived from spreadsheet models based on forage availability estimates and the metabolic requirements of the respective mammal species, and that incorporate modifications of the agriculture-based large stock unit approach. the potential population size of each species was calculated by multiplying its density estimate with the area of suitable habitat. population sizes were calculated for pristine, or near pristine, habitats alone, and then for these habitats together with potentially restorable habitats for two park planning domain scenarios. these data will enable (a) the measurement of the effectiveness of the gaenp in achieving predetermined demographic, genetic and evolutionary targets for mammals that can potentially occur in selected park sizes and configurations, (b) decisions regarding acquisition of additional land to achieve these targets to be informed, (c) the identification of species for which targets can only be met through metapopulation management, (d) park managers to be guided regarding the re-introduction of appropriate species, and (e) the application of realistic stocking rates. where possible, the model predictions were tested by comparison with empirical data, which in general corroborated the predictions. all estimates should be considered as testable hypotheses. key words: conservation planning, mammals, distribution, density, population estimates, addo, south africa. a.f. boshoff, g.i.h. kerley and s.l. wilson, terrestrial ecology research unit, department of zoology, university of port elizabeth, port elizabeth, 6013, republic of south africa; r.m. cowling, terrestrial ecology research unit, department of botany, university of port elizabeth, port elizabeth, 6013, republic of south africa. issn 0075-6458 85 koedoe 45/2 (2002) introduction in november 2000, the global environment facility (gef) approved a grant to south african national parks (sanparks) to research and prepare a full proposal to the gef for the planning and establishment of a “greater” addo elephant national park (gaenp). the sw boundary of the addo elephant national park (aenp) is some 35 km east of the city of port elizabeth (33°58's, 25°31'e), south africa. the vision for an expanded addo elephant national boshoff.qxd 2005/12/09 09:46 page 85 park was developed and documented by kerley & boshoff (1997, 2002, www.zoo. upe.ac.za/teru). systematic conservation planning forms an integral and critical component of the implementation of the gaenp project, managed by south african national parks (“http://www.addoelephantpark.com” www.addoelephantpark.com). note that the original boundary of the proposed expansion to the aenp (kerley & boshoff 1997) has been modified, by sanparks, for the purposes of the conservation planning exercise for gaenp, by the addition of a 5-km buffer that follows cadastral boundaries. while the impressive plant diversity remains a major focus of conservation planning in the establishment of a gaenp (kerley & boshoff 1997), other biota and ecological processes which impact on the park’s biodiversity must be taken into account in attempting to achieve its broad conservation objectives. the species’ patterns and ecological and evolutionary processes in the gaenp planning domain include the medium to large-sized mammals and the processes that they drive, many of which (a) are in need of conservation intervention, and (b) may have an important impact on the park’s biota, at the species, community and ecosystem functioning levels. herbivory is known to have an impact on the species composition, structure and dynamics of fynbos vegetation (campbell 1986; johnson 1992) and thicket vegetation (barratt & hall-martin 1991; johnson et al. 1999; lombard et al. 2001; moolman & cowling 1994; penzhorn et al. 1974; stuart-hill 1992; stuart-hill & aucamp 1993). the important role that the proposed mega reserve will play in conserving a diverse array of larger mammals, including the top predators and a number of megaherbivores, is emphasised by kerley & boshoff (1997). the numerous ecological processes that are mediated by the larger mammals, or that they participate in, are reported on elsewhere (boshoff et al. 2001a). the medium to large-sized mammals were selected as “target” species (sensu wilcox 1982) for the gaenp planning exercise because it is likely that if their minimum area requirements are met, adequate survival conditions will be simultaneously met for other biota. in this regard, many of these mammals qualify as “umbrella” species (sensu wilcox 1982) since their minimum area requirements are likely to be at least as comprehensive as those for the remainder of the community. mammals with a large body size (e.g., some ungulates) or which occupy a high trophic level (e.g., carnivores) are regarded as good candidates for target species acting as “umbrella” species (wilcox 1982). in addition, the distributions and spatial requirements/densities of the larger mammals are probably better known, or can be better estimated, than those of the smallsized mammals in the gaenp planning domain. in any case, realistic data for these two population parameters are essential for any conservation exercise that deals with the establishment and maintenance of minimum viable populations of the larger mammalian fauna (caughley 1994; caughley & sinclair 1994; lande & barrowclough 1987). an additional consideration for determining minimum area requirements for preserving biological diversity is that of the estimation of minimum viable populations (mvp) for “target” species (wilcox 1982; soulé 1987). the mvp is a set of specifications concerning the size and structure of the populations of a species that is necessary to provide a margin of safety from extinction. the mvp for a species can be translated into the minimum area requirements by determining the amount and type of habitat that will satisfy the mvp. in view of this, it is necessary for realistic estimates of the spatial requirements/densities of each the selected species in the gaenp planning domain to be obtained. in summary, systematic data and information are required to enable conservation planners to calculate the potential numbers of individuals of each mammal species, within the mammal habitats within various park configuration scenarios. these data will enable planners to measure the effectiveness of the proposed gaenp in achieving predeterkoedoe 45/2 (2002) 86 issn 0075-6458 boshoff.qxd 2005/12/09 09:46 page 86 mined demographic, genetic and evolutionary targets for medium to large-sized mammals that can potentially occur in the park. in addition, they will inform decisions regarding acquisition of additional land, where necessary, to achieve these targets, and help identify species for which targets can only be met through metapopulation management. approach the indigenous mammal species included in this study (table 1) are those with a mass greater than ca. 2 kg (cf. chew 1978), that are the most prominent on the landscape, and which are generally amenable to direct management. as part of a separate exercise, 43 land classes were delineated through field mapping by kruger & sykes (2002), using as a basis the hierarchical classification of subtropical thicket by vlok & eustonbrown (2002). it was considered impractical to use this detailed classification for deriving the potential distributions and estimated spatial requirements/densities of the larger mammals. it was consequently decided to collapse the 43 land classes into a practical number (21) of mammal habitat classes (mhcs) and to use these as the biodiversity surrogates for the mammal conservation planning component of the greater addo elephant national park. only those land classes that exhibited a generally high degree of similarity, in terms of vegetation structure (and hence mammal habitat) and productivity (determining mammal densities), and for which any differences that exist are considered unlikely to impact significantly on the known and potential presence and densities of mammal species, were combined. the potential distributions and estimated spatial requirements of the two otter species are based exclusively on aquatic habitats, i.e., coastline and rivers, where appropriate. distributions methods two steps were followed in determining the potential distribution of each species, within each mhc in the gaenp planning domain. 1. collation and interpretation of evidence that a species occurred, or could potentially occur in all, or in a specific part, of the gaenp planning domain. the early and recent published literature was consulted, as were conservation scientists and managers with a good knowledge of the macro fauna of the existing addo elephant national park (aenp) and close environs (see boshoff & kerley 2001 and boshoff et al. 2001b for details of the methods used). the mammal checklist for the aenp was also consulted, as were the mammal collection registers of the amatole museum in king william’s town, where the terrestrial mammal collections from the four provincial museums in the eastern cape are now housed. the present study attempts to reconstruct the distributions of indigenous herbivores in the period prior to arrival of european settlers, in the gaenp planning domain, in the mid 17th century. these distributions thus represent a situation where the patterns and processes exhibited by the mammals of the region were presumably still fairly intact. thus, domestic herbivores, maintained by khoi pastoralists in the period prior to european settlement, have not been taken into account in this analysis, owing to a lack of information on their distributions, nomadic movements and densities. zoological and explorer’s records from the 17th, 18th and 19th centuries have been well reviewed by du plessis (1969), rookmaaker (1989) and skead (1987). these reviews were useful in determining the general presence or absence of most species in all or parts of the gaenp planning domain, but they generally proved to be vague in terms of the exact areas and habitats occupied by the various species. this resulted mainly from the fact that most early hunters and naturalists only recorded mammal occurrences along well travelled, or passable, routes, and few travelled at night, thereby missing the nocturnal species. other problems arose with interpreting the early, published accounts with regard to the accurate identification of some species (see skead 1987). issn 0075-6458 87 koedoe 45/2 (2002) boshoff.qxd 2005/12/09 09:46 page 87 the following additional sources were consulted for information on the historical occurrence of mammals in the broader area around the gaenp: coetzee (1979); hewitt (1931); lloyd & millar (1983); shortridge (1942); skinner & smithers (1990); smithers (1986); stuart (1981); stuart (1985); stuart et al. (1985). a review of the recent (20th century) literature revealed that surveys are incomplete in terms of species and/or area covered and tend to use political boundaries rather than ecological zones as the basic mapping units. the scale of the distribution maps in the standard account of the mammals in the southern african sub-region (skinner & smithers 1990) allows only generalised ranges (extents of occurrence) to be determined. similarly, distributions of threatened mammal species are illustrated on a broad regional basis (smithers 1986). museum specimens and records provide useful point data but are biased in that they only provide “presence” data, i.e., they do not represent the results of systematic data collection throughout the gaenp planning domain, and they do not take into account the possible migratory or nomadic patterns of some species. 2. estimation of potential presence of the species, based on their ecological requirements the potential presence/absence of each species in each mhc was determined according to our understanding of their ecological requirements, including a review of published habitat requirements (in the general gaenp area and further afield), our personal field knowledge, and the respective habitat characteristics of each mhc. these characteristics included dominant plant species, vegetation structure, grass component, soil nutrients, geology, topography, modal altitude, mean rainfall and rainfall seasonality). see boshoff & kerley (2001) and boshoff et al. (2001b) for details of the methods used. as part of this exercise, aenp conservation scientists and managers, with ecological knowledge of mammals of the area, were consulted. the potential distribution of each species is presented according to three categories: mhcs with the potential to sustain significant resident (i.e., present all year round and breeding) populations. in these mhcs the animals are generally homogeneously distributed across the landscape; mhcs which may be used on a seasonal basis, or which may carry small populations in habitat refugia (i.e., patchy basis). in these mhcs the animals are generally not homogeneously distributed, temporally and spatially, across the landscape; mhcs where the species is unlikely to occur, except perhaps for vagrants or during rare and short incursions. in such cases the species was considered to be absent, and the mhc in question could not be relied upon to contribute to the conservation of that species. the hippopotamus potentially occurs, in suitable habitat, in and along major rivers and dams. these waterbodies must be perennial in nature and must contain pools at least 1.5 m deep. the distance travelled from watercourses to feeding grounds depends on forage availability and can vary widely. hippopotamus generally forage within about 1.5 km from waterbodies but will move freely up to eight or 10 km, and are known to move much further when forage is scarce (skinner & smithers 1990). potential hippopotamus habitat was marked on a digital terrain map produced by csir-environmentek. for practical reasons, the overall distribution of this species is presented, rather than its distribution according to mammal habitat class. using gis, those parts of mhcs that overlap with potential hippopotamus habitat are considered as additional (i.e., additional to the original 21 mhcs) mhcs and are treated as such for the calculation of estimated spatial requirements and densities (see “spatial requirements”). the potential distributions (that are linear in nature) of the two otter species were marked on a digital terrain map that identifies the major rivers and dams. a conservative approach was adopted in determining these distributions; only waterbodies that can be confidently classified as being perennial were included. the approach described above, which involves a simple model based on the estimated range of each species and its association with mappable environmental features expressed as a series of polygons, is broadly similar to that used in other studies (e.g., butterfield et al. 1994). results of the 44 indigenous, non-marine mammal species that occur, or can potentially occur in the proposed gaenp (table 1), 41 species occur exclusively in terrestrial habitats, whereas three species, the cape clawless and spotted-necked otters and the hippopotamus are associated with aquatic habitats. three species are omnivores, 18 are carnivores and 23 are herbivores. of the 44 species, 35 are koedoe 45/2 (2002) 88 issn 0075-6458 boshoff.qxd 2005/12/09 09:46 page 88 issn 0075-6458 89 koedoe 45/2 (2002) table 1 the common and scientific names, foraging guild classifications and current (2001) presence of potentially occurring mediumto large-sized omnivorous, carnivorous and herbivorous mammals in the gaenp planning domain. p = present in 2001. taxonomic order (except for aardvark – see text) and nomenclature (scientific and common names) follow skinner & smithers (1990) common name scientific name foraging guild presence in 2001 omnivores chacma baboon papio cynocephalus p vervet monkey cercopithecus aethiops p porcupine hystrix africaeaustralis p aardvark orycteropus afer p carnivores aardwolf proteles cristatus p brown hyaena hyaenna brunnea spotted hyaena crocuta crocuta cheetah acinonyx jubatus leopard panthera pardus p lion panthera leo caracal felis caracal p african wild cat felis lybica p small spotted cat felis nigripes p serval felis serval bat-eared fox otocyon megalotis p wild dog lycaon pictus cape fox vulpes chama p black-backed jackal canis mesomelas p cape clawless otter aonyx capensis p spotted-necked otter lutra maculicollis p honey badger mellivora capensis p herbivores african elephant loxodonta africana mixed feeder p black rhinoceros diceros bicornis browser p cape mountain zebra equus zebra zebra bulk grazer p burchell’s zebra equus burchelli bulk grazer p bushpig potamochoerus porcus mixed feeder p warthog phacochoerus aethiopicus concentrate grazer p hippopotamus hippopotamus amphibius bulk grazer p black wildebeest connochaetes gnou concentrate grazer red hartebeest alcelaphus buselaphus concentrate grazer p blue duiker philantomba monticola browser p common duiker sylvicapra grimmia browser p springbok antidorcas marsupialis mixed feeder p klipspringer oreotragus oreotragus browser p oribi ourebia ourebi concentrate grazer steenbok raphicerus campestris browser p grysbok raphicerus melanotis browser p grey rhebok pelea capreolus concentrate grazer p african buffalo syncerus caffer bulk grazer p kudu tragelaphus strepsiceros browser p bushbuck tragelaphus scriptus browser p eland taurotragus oryx mixed feeder p reedbuck redunca arundinum concentrate grazer mountain reedbuck redunca fulvorufula concentrate grazer p boshoff.qxd 2005/12/09 09:46 page 89 koedoe 45/2 (2002) 90 issn 0075-6458 already present and nine could be considered for re-introduction. the 21 mhcs delineated for this study are mapped in fig. 1 and listed in table 2. the potential occurrence of each species in each mhc, on a “resident” or “seasonal/patchy” basis (table 2), is illustrated in a series of distribution maps (figs. 2–44). due to a lack of detailed habitat information, the hippopotamus and the two otter species are considered to be potentially resident in all habitats mapped for these species. discussion notwithstanding the constraints inherent in the approach used here, the maps provided in this report are considered to represent realistic potential distributions of the medium to large-sized mammals in the gaenp planning domain. we stress, however, that these data are underpinned by putative habitatmammal relationships that are testable in the future. nonetheless, the data provide new information that is essential for effective conservation planning in the gaenp, and for developing a greater understanding of the larger terrestrial vertebrates as indicators of environmental change in the proposed park (macdonald 1992). the black wildebeest is the only species for which the greater part of its distribution range within the gaenp planning domain falls within the 5-km buffer zone. for two other species, namely oribi and reedbuck, a significant proportion of their distribution range within the planning domain falls within the 5-km buffer. it is emphasised that the allocation of species to specific mhcs should not be interpreted to imply that the distributions of the mammals are spatially and temporally fixed in the planning domain. because of the dearth of ecological information from the region, any reconstruction of the demographics and dynamics of the medium to large mammal populations must be based on the collection of new information. owing, in part, to the expansion of the aenp, there are currently seven extralimital species in the park, namely gemsbok oryx gazella, impala aepyceros melampus, waterbuck kobus ellipsiprymnus, blesbok damaliscus dorcas phillipsi, blue wildebeest connochaetes taurinus, red lechwe kobus leche and nyala tragelaphus angasii. it is recommended that these species be removed from the aenp, in view of the real and potential ecological and economic costs of keeping them in the park (castley et al. 2001). spatial requirements methods the estimated spatial requirements of each species, and the associated density estimates, refer exclusively to those mhcs in the gaenp planning domain where the species is likely to occur, on a “resident” or “seasonal/patchy” basis. omnivores and carnivores the overall lack of information from the gaenp domain precluded an estimation of the spatial requirements of the omnivores and carnivores according to individual mammal habitat classes. consequently, the planning domain was treated as a homogeneous unit for this purpose. this is likely to be more appropriate for the smaller species than for the larger ones; the abundance of the latter will generally reflect the abundance and spatial distribution of the larger herbivores. estimates of the spatial requirements of each species in each mhc were based on a review of available information on densities, social structures, breeding units, territory sizes and home ranges. however, since published ecological information for the region is not available (cf. boshoff et al. 2001b) for any of the species that can potentially occur there, estimates based on the interpretation and extrapolation of information on the relevant species from other regions in south africa, mainly the nama-karoo, grassland and savanna biomes (sensu low & rebelo 1996), were used as surrogates. in the case of the carnivores (especially the large predators and scavengers such as lion and spotted hyaena) the assumption is made that predator-prey systems are in operation and that sufficient food is available. for the sake boshoff.qxd 2005/12/09 09:46 page 90 issn 0075-6458 91 koedoe 45/2 (2002) table 2 the presence/absence of the mediumto large-sized mammals in the gaenp planning domain, according to mammal habitat class (r = resident, sp = seasonal/patchy) common name mammal habitat class fo re st t hi ck et f or es t m os ai c t hi ck et s av an na m os ai c z uu rb er g m es ic t hi ck et a dd o h ei gh ts m es ic t hi ck et su cc ul en t t hi ck et sp ek bo om ve ld e as te rn s pe kb oo m n oo rs ve ld w es te rn s pe kb oo m n oo rs ve ld n oo rs ve ld g ra ss y b on tv el d k ar ro id b on tv el d k ar oo t hi ck et m os ai c k ar ro id d w ar f sh ru bl an d k ar ro id b ro ke n v el d so ur g ra ss la nd m ix ed g ra ss y sh ru bl an d fy nb os r ip ar ia n w oo dl an d d un ef ie ld su nd ay s sa ltm ar sh chacma baboon r r r r r r sp r r r r r vervet monkey r r r r r r r r r sp sp sp r sp sp r sp porcupine r r r r r r r r r r r r r r r r r r r sp aardwolf sp sp sp sp sp sp sp sp r r r r r r r sp sp brown hyaena sp r r r r r r r r r r r r r r r r r r sp spotted hyaena sp sp sp sp sp r r r r r r r r sp r sp r sp sp cheetah r r r r r leopard r r r r r r r r r r r r r r r r r r r sp sp lion r sp sp r sp r r r r r r r r sp r sp r sp sp caracal sp sp r r r r r r r r r r r r r sp r sp r sp sp african wild cat sp r r r r r r r r r r r r r sp r sp r sp sp small spotted cat sp r r r r r r r r r serval sp sp sp sp sp sp sp sp r bat-eared fox sp sp sp r r r r r r r r sp sp wild dog r r r r r r r r r r r r r sp r sp r sp sp cape fox sp sp r r r r r r r r sp sp black-backed jackal sp r r r r r r r r r r r r r r sp r sp r sp sp honey badger sp sp r r r r r r r r r r r r r r r r r sp sp aardvark r sp sp r r r r r r r r r r sp r r african elephant sp r r r r r r sp sp sp r r r sp sp sp r black rhinoceros sp r r r r r r r r r r r r r r r mountain zebra sp sp sp sp sp sp sp sp r r r plains zebra sp sp sp r r r r r r r sp bushpig r r sp r r r r sp sp sp r sp warthog sp sp sp sp r r r r r sp r r r black wildebeest sp sp red hartebeest sp sp r r r r r sp r r sp r sp sp blue duiker r r sp r r sp sp sp common duiker sp sp r r r r r r r r sp sp r sp sp sp r sp r sp springbok sp r sp r sp sp sp r r sp klipspringer r sp sp r sp sp r oribi sp sp sp steenbok sp r r r sp sp r r sp p grysbok r sp r r sp r sp sp sp r r sp grey rhebok r r r cape buffalo sp r sp sp sp sp sp sp sp r r r sp sp sp sp r kudu r r r r r r r r sp r r sp r r r bushbuck r r r r r r r sp sp sp sp sp eland sp sp sp sp sp sp sp sp sp sp sp sp sp sp sp sp sp reedbuck sp mountain reedbuck r r sp r r r sp boshoff.qxd 2005/12/09 09:46 page 91 koedoe 45/2 (2002) 92 issn 0075-6458 fi g. 1 . t he m am m al h ab ita t c la ss es in th e pl an ni ng d om ai n fo r th e g re at er a dd o n at io na l p ar k in iti at iv e. s ee te xt f or d et ai ls . boshoff.qxd 2005/12/09 09:46 page 92 issn 0075-6458 93 koedoe 45/2 (2002) of brevity, the sources of the data used to estimate the spatial requirements have not been included in this paper. given that they occur along rivers or along the coastline, the density estimates for the two otter species are expressed in linear terms (individuals/km). where the distribution of the cape clawless otter potentially overlaps with that of the spotted-necked otter, the food resources have been equally apportioned between them, thereby reducing the potential density of each species. a conservative approach to the estimation of the spatial requirements of the omnivores and carnivores in the gaenp planning domain was adopted because of the naturally, and relatively, low herbivore carrying capacity in some habitats, and a generally poor understanding of the ecology of the species concerned. this was achieved by: (a) usually adopting the lowest densities or largest territories or home ranges provided in the literature; (b) using the home range when territory size is not known; (c) basing, in appropriate cases, the estimates only on the sizes of the territories or home ranges of breeding adults—in these cases effective densities may be higher when non-territorial individuals (e.g., sub-adults, immatures and juveniles) are taken into account; and (d) reducing the densities in the seasonal/patchy habitats to 20 % of those calculated for the “core” habitats (boshoff et al. 2001b). herbivores given the virtual absence of information on the spatial requirements of herbivores in the gaenp planning domain, we followed a pragmatic approach in the derivation of the necessary estimates. this involves a spreadsheet model, based on forage availability estimates and the metabolic requirements of the mammal species in question. the approach followed is very similar to that described by boshoff et al. (2001b) but some adjustments have been made to accommodate the gaenp requirements and characteristics. although the porcupine is predominantly a herbivore, we have treated it as an omnivore and excluded it from the spreadsheet model, since it does not fit in the conventional grazer/browser classification. the six sequential components of the model are described below: 1. allocation of species to foraging guilds each herbivore species was classified according to one of four foraging guilds (table 1, adapted from collinson & goodman 1982), namely: bulk grazer; concentrate grazer; mixed feeder (grazer/browser); and browser. 2. adjustment of the agricultural stocking rate the recommended agricultural stocking rates (srs) for the respective land/agricultural units, as calculated by the south african department of agriculture on the basis of large stock units (lsus) (anon. 1985), were used as guidelines for estimating forage production, and ultimately the spatial requirements of herbivores within each mammal habitat class. it must be emphasised that the term “spatial requirements” normally refers to an ecological response, whereas the term “stocking rates” normally refers to an operator/manager response. the definition and use of the lsu concept to determine stocking rates for livestock and wildlife is discussed in some detail by boshoff et al. (2001b). where available data (cf. stuart-hill & aucamp 1993) have permitted a comparison, the agricultural stocking rate broadly agrees with published empirical data. agricultural management is usually aimed at maximising production (morris et al. 1999), and therefore we adopted a highly conservative approach in the calculations for the indigenous ungulates, for the purpose of sustaining populations and protecting biodiversity. this took the form of adjusting (i.e., reducing) the department of agriculture stocking rate applicable to each mhc by a proportion which was estimated following a subjective assessment of the biophysical attributes, as surrogates for the productivity of forage, for the mhc in question. key surrogates here are dominant vegetation, grass component, soil nutrient status, mean annual rainfall, rainfall seasonality, modal altitude and general topography. in this way, the agricultural srs of mhcs characterised by low productivity, low nutrient soils and a limited grass component, were reduced by a higher percentage than those mhcs characterised by a higher productivity, relatively higher soil nutrient status and a relatively high grass component. thus: adjsr = x (1+y) (1) where adjsr = adjusted stocking rate, x = agricultural carrying capacity/stocking rate (ha/lsu), y = adjustment value (where, e.g., 60 % = 0.4), and lsu = large stock unit. boshoff.qxd 2005/12/09 09:46 page 93 koedoe 45/2 (2002) 94 issn 0075-6458 department of agriculture stocking rates were not available for some mhcs, nor could they be determined, owing to mapping scale differences. in these cases, stocking rates were estimated according to: an interpretation of the key biophysical attributes (as listed above); the stocking rates for similar mhcs; and the stocking rates for neighbouring mammal habitat classes. for the purposes of the model, these adjusted stocking rates (ha/lsu) were expressed as animal unit densities (lsu/ha). 3. allocation of animal units to foraging guilds, within mhcs the available animal units, per hectare, within each mhc (expressed as adjusted lsu/ha) were allocated to each of the four foraging guilds, where appropriate (i.e., for each guild that was represented in that mhc). to achieve this, allocations of forage (as percentages) were made for each guild within each mhc, based on subjective estimations of the proportions and nature (e.g., sweet or sour grassveld) of graze and browse, as suggested by the mhc biophysical descriptions and figs. 2–44. the potential distribution of the different mammals in the greater addo elephant national park planning domain, according to mammal habitat class (mhc). solid shading denotes mhcs with the potential to sustain significant resident (i.e., present all year round and breeding) populations; grey shading denotes mhcs which may be used on an ephemeral (i.e., seasonal) basis, or which may carry small populations in habitat refugia (patchy basis), and no shading denotes mhcs where the species is unlikely to occur, except perhaps for vagrants or during rare and short-lived incursions. fig. 2. fig. 3. fig. 4. fig. 5. fig. 6. boshoff.qxd 2005/12/09 09:47 page 94 issn 0075-6458 95 koedoe 45/2 (2002) our personal knowledge of these habitats (appendix 1). these allocations were then corroborated with the guild structures of the herbivores occurring in each mammal habitat class. for example, a check was made that the distribution patterns described earlier indicated that grazers were the dominant herbivores in mhcs dominated by grass. for pragmatic reasons, no distinction was made between the preand postdarlington dam scenarios, i.e., the dam was considered to be a permanent feature. since the dam itself (water area) it does not contribute any forage, it has been subtracted from the total area of riparian woodland that provides suitable hippopotamus habitat. 4. allocation of available animal units to individual species within foraging guilds, within mhcs for each mhc the available animal units, calculated in step 3 above and expressed as adjusted lsu/ha, were allocated to the herbivore species within each foraging guild. thus, where more than one species occurs within a single foraging guild within an mhc, the lsus accorded to that guild are allocated to these species in equal proportions. this course was chosen owing to the paucity of information on resource partitioning within these guilds. 5. adjustment for seasonality/ patchiness species that are resident in a mhc will most likely have different forage requirements (and possibly other ecological requirements, e.g., availability of surface water, shelter/cover) than species that are highly spatially localised or that may only be present for a limited part of a year (i.e., nomads or migrants). therefore, there was a requirement for the model to incorporate seasonality and habitat patchiness. this was addressed by reducing by 60 % the amount of forage fig. 7. fig. 8. fig. 9. fig. 10. fig. 11. boshoff.qxd 2005/12/09 09:47 page 95 koedoe 45/2 (2002) 96 issn 0075-6458 allocated (expressed as adjusted lsu/ha) to seasonal/patchy species. we assumed that the amount, and indeed quality, of resources was limiting, rather than their seasonal availability or total absence. the basis for using a value of 60 % is the same as that used for a similar study in the cape floristic region, where a value of 90 % was used (boshoff et al. 2001b). a value of 60 % was used here to reflect the probable higher and more reliable year-round forage productivity. thus, each species in each mhc is classified as “resident” or “seasonal/ patchy” (see “distributions”). the lsus that were “released” by a “seasonal/patchy” species were reallocated, in equal proportions, to other species within the same foraging guild. this gives the recalculated number of lsus available to each species within a mammal habitat class. in cases where other species are not present in the same guild, the “released” animal equivalents (lsu/ha) were considered as “floaters” within that mhc—to be utilised across the graze/browse spectrum by the remaining species in the mammal habitat class. 6. calculation of species specific densities and spatial requirements, within each mhc the number of individuals of a species per ha (density), within each mhc, was calculated as follows: d = lsurec /sequ (2) where d = density (number of individuals/ ha), lsurec = recalculated lsus per species (as calculated in steps 1-5 above) and sequ = species’ lsu equivalent. the lsu equivalents for the species follow grossman (1991); that for african elephant follows meissner (1982). the estimated spatial requirement for an individual of each species, within each mhc, is calculated as follows: fig. 12. fig. 13 fig. 14. fig. 15. fig. 16. boshoff.qxd 2005/12/09 09:47 page 96 issn 0075-6458 97 koedoe 45/2 (2002) sprqi = 1/d (3) where sprqi = spatial requirement (ha/individual) of an individual, d = density (individuals/ha from equation 2). constraints a limitation on the spatial requirements of some herbivores is provided by social interaction, namely intolerance of conspecifics, as well as a number of other constraints, e.g., presence of surface water, seasonal food availability. it is known that, irrespective of the availability of forage, social and other constraints can limit the densities of ungulates (e.g., see moen 1973), and for some species the availability of food, water and shelter is superseded by social factors in determining densities. in this regard, the spatial requirements predicted by our model were compared, where possible, with available information to investigate whether species’ social constraints had been violated. model testing the outputs of the model were tested by comparing spatial requirement estimates derived from the model with published, empirically derived observations of densities of species for which appropriate data are available. such data are not available for complete species assemblages. hippopotamus for those parts of mhcs where hippopotamus can potentially occur, a separate spreadsheet model was constructed. it differs from the model for the mhcs without hippopotamus in that hippopotamus has been inserted as an additional herbivore (it is a bulk grazer). this results in adjustments to the allocation of forage between bulk grazers within these parts of mhcs, and ultimately the densities of all species within this foraging guild. there is no published information on the densities and spatial requirements of hippopotamus in the gaenp domain, or even in the eastern cape, fig. 17. fig. 18. fig. 19. fig. 20. fig. 21. boshoff.qxd 2005/12/09 09:47 page 97 koedoe 45/2 (2002) 98 issn 0075-6458 to validate the estimates provided by the model. results omnivores and carnivores the estimates of the spatial requirements/densities for the omnivores and carnivores are provided in table 3. as an example of the basis for the estimation of the spatial requirements, the case of the chacma baboon is given (box 1; cf. table 3). the fig. 22. fig. 23. fig. 24. fig. 25. fig. 26. box 1: estimation of the spatial requirements of the chacma baboon. breeding unit/social structure baboons are highly social, living in female bonded troops of between four and around 100130 individuals, with one adult male in small troops and up to 12 males in large troops; average troop size is 40 (skinner & smithers 1990, apps 1996) and troop size is apparently correlated with habitat quality. breeding density/home range/ territory size troops have home ranges but they are not territorial and rather tend to avoid other troops (apps 1996). in the good hope section of the cape peninsula national park home ranges of three troops of 20, 35 and 80 baboons were 9.1, 14.8 and 33.7 km², respectively, with home range being related to size of troop (devore & hall 1965). home ranges of 4004000 ha have been recorded. boshoff.qxd 2005/12/09 09:47 page 98 issn 0075-6458 99 koedoe 45/2 (2002) ta bl e 3 e st im at ed d en si tie s an d sp at ia l r eq ui re m en ts fo r se le ct ed m ed iu m to la rg e om ni vo re s an d ca rn iv or es in ( a) “ se as on al /p at ch y” a nd ( b) “ re si de nt ” oc cu rr en ce c at ego ri es in th e g a e n p pl an ni ng d om ai n. s ee te xt a nd b os ho ff et a l. (2 00 1a ) fo r as su m pt io ns a nd c al cu la tio ns . s ci en tif ic n am es in t ab le 1 sp ec ie s e xt ra po la te d de ns iti es a nd s pa tia l r eq ui re m en ts e st . d en si ty ( in d. /h a) e st . s pa t./ re q. ( ha /in d. ) se as ./p at ch y r es id en t se as ./p at ch y r es id en t c ha cm a ba bo on 1 tr oo p of 8 0 us es 3 40 0 ha . 0. 01 29 9 0. 02 32 6 77 43 v er ve t m on ke y 25 /tr oo p, 8 tr oo ps r eq ui re d fo r 20 0 in di vi du al s at c a. 8 0 ha /tr oo p. 0. 18 51 9 0. 33 33 3 5. 4 3 po rc up in e fa m ily g ro up o f 5 in di vi du al s; te rr ito ry s iz e of 8 0 ha 0. 03 44 8 0. 06 25 0 29 16 a ar dw ol f m al es a nd f em al es s ha re te rr ito ri es o f up to 8 00 h a. 0. 00 13 9 0. 00 25 0 72 0 40 0 b ro w n hy ae na c la n of 4 m em be rs h as a te rr ito ry s iz e of a bo ut 2 5 00 0 ha . 0. 00 00 9 0. 00 01 6 11 25 0 62 50 sp ot te d hy ae na a cl an o f 9 w ou ld r eq ui re a te rr ito ry o f ar ou nd 4 0 00 0 ha . 0. 00 01 3 0. 00 02 3 79 99 44 44 c he et ah e st . h om e ra ng e fo r 5 an im al s (2 m , 3 f) a t 1 00 0 00 h a, 0. 00 00 3 0. 00 00 5 33 48 0 18 60 0 w ith 7 5% o ve rl ap . 5 0 an im al s = 10 0 00 0 + 25 % f or 1 0 ite ra tio ns l eo pa rd 1 pa ir r eq ui re s a ho m e ra ng e of a bo ut 2 0 00 0 ha 0. 00 00 6 0. 00 01 0 18 00 0 10 00 0 l io n a pr id e of 1 0 an im al s (a du lts , s ub -a du lts a nd y ou ng ) 0. 00 01 2 0. 00 02 2 81 00 45 00 m ay r eq ui re a te rr ito ry o f ab ou t 4 5 00 0 ha . c ar ac al pa ir s ha ve o ve rl ap pi ng ( by u p to 2 0% ) ho m e ra ng es o f ab ou t 6 6 00 h a. 0. 00 02 1 0. 00 03 8 47 52 26 40 a fr ic an w ild c at 1 pa ir h as a h om e ra ng e of a pp ro xi m at el y 25 0 ha . 0. 00 44 4 0. 00 80 0 22 5 12 5 sm al l s po tte d ca t m al es a nd f em al es h av e ov er la pp in g (c a 20 % ) te rr ito ri es o f 0. 00 07 4 0. 00 13 3 13 50 75 0 ab ou t 9 00 h a. t hu s, 1 p ai r ha s a te rr ito ry o f ab ou t 1 50 0 ha . se rv al m al es a nd f em al es o cc ur in h om e ra ng es o f up to 3 00 0 ha , 0. 00 03 7 0. 00 06 7 27 00 15 00 ta ki ng o ve rl ap in to a cc ou nt . b at -e ar ed f ox d en si ty o f 3 an im al s/ 10 0 ha 0. 01 66 7 0. 02 94 1 60 34 w ild d og 0. 01 7 an im al s pe r 10 0h a 0. 00 00 9 0. 00 01 7 10 58 8 58 82 c ap e fo x o nl y in fo rm at io n fo r hu nt in g ra ng e (u p to 5 00 h a) ; o ve rl ap pi ng 0. 00 14 8 0. 00 26 7 67 5 37 5 ho m e ra ng es . s ay 1 p r ne ed s 75 0 ha . b la ck -b ac ke d ja ck al te rr ito ry /h om e ra ng e of a bo ut 1 10 0 ha /p r. 0. 00 10 1 0. 00 18 2 99 0 55 0 c ap e cl aw le ss o tte r 1 pe r 3. 5 km o f ri ve r; 1 p er 2 k m o f co as tli ne sp ot te dne ck ed o tte r 1 pe r 2. 5 km o f ri ve r h on ey b ad ge r m al e an d fe m al e ha ve o ve rl ap pi ng h om e ra ng es o f ab ou t 1 0 00 0 ha . 0. 00 01 1 0. 00 02 0 90 00 50 00 a ar dv ar k e st . h om e ra ng es f or 1 m al e an d 1 fe m al e of 7 5 00 h a. 0. 00 01 5 0. 00 02 7 67 50 37 50 boshoff.qxd 2005/12/09 09:47 page 99 koedoe 45/2 (2002) 100 issn 0075-6458 table 4 estimated densities and spatial requirements of the larger mammalian herbivores in areas of 20 mammalian habitat classes in the gaenp planning domain that do not contain hippopotamus habitat. data derived from a spreadsheet model. see text for calculations and assumptions. scientific names in table 1 mammal species density est. spat. species density est. spat. habitat (ind./ha) req. (ind./ha) req. class (ha/ind.) (ha/ind.) forest african elephant 0.00035 2876 common duiker 0.01704 59 bushpig 0.01758 57 bushbuck 0.03833 26 blue duiker 0.16611 6 thicket forest african elephant 0.00540 185 common duiker 0.06617 15 mosaic black rhinoceros 0.00361 277 grysbok 0.34740 3 bushpig 0.06818 15 cape buffalo 0.00208 482 blue duiker 0.69481 1 bushbuck 0.16034 6 thicket savanna african elephant 0.00540 185 oribi 0.04082 25 mosaic black rhinoceros 0.00807 124 steenbok 0.06122 16 burchell’s zebra 0.00866 116 grysbok 0.06122 16 bushpig 0.00974 103 cape buffalo 0.02136 47 warthog 0.01143 88 kudu 0.02466 41 red hartebeest 0.00772 130 bushbuck 0.10243 10 blue duiker 0.12245 8 eland 0.00198 504 common duiker 0.14796 7 reedbuck 0.01143 88 springbok 0.01429 70 zuurberg mesic african elephant 0.00229 436 grysbok 0.10621 9 thicket black rhinoceros 0.00386 259 cape buffalo 0.00099 1011 cape mtn. zebra 0.00168 595 kudu 0.01180 85 bushpig 0.02897 35 bushbuck 0.04902 20 blue duiker 0.21242 5 eland 0.00182 551 common duiker 0.07081 14 addo heights african elephant 0.00229 436 common duiker 0.07081 14 mesic thicket black rhinoceros 0.00386 259 grysbok 0.10621 9 burchell’s zebra 0.00160 623 cape buffalo 0.00099 1011 bushpig 0.02897 35 kudu 0.01180 85 warthog 0.00094 1063 bushbuck 0.04902 20 blue duiker 0.21242 5 eland 0.00182 551 succulent african elephant 0.00111 898 common duiker 0.06092 16 thicket black rhinoceros 0.00332 301 klipspringer 0.07833 13 cape mtn. zebra 0.00076 1323 grysbok 0.02948 34 bushpig 0.01407 71 cape buffalo 0.00045 2247 warthog 0.00571 175 kudu 0.01015 98 red hartebeest 0.00386 259 bushbuck 0.04218 24 blue duiker 0.05896 17 eland 0.00088 1134 spekboomveld african elephant 0.00142 706 common duiker 0.06128 16 black rhinoceros 0.00334 299 grysbok 0.09192 11 burchell’s zebra 0.00069 1452 cape buffalo 0.00042 2354 bushpig 0.01791 56 kudu 0.01021 98 warthog 0.00727 138 bushbuck 0.04242 24 blue duiker 0.06566 15 eland 0.00112 891 eastern african elephant 0.00027 3753 springbok 0.02716 37 spekboom black rhinoceros 0.00387 258 klipspringer 0.03175 32 noorsveld cape mtn. zebra 0.00353 284 steenbok 0.10648 9 boshoff.qxd 2005/12/09 09:47 page 100 issn 0075-6458 101 koedoe 45/2 (2002) burchell’s zebra 0.01852 54 cape buffalo 0.00208 482 warthog 0.00667 150 kudu 0.01183 85 black wildebeest 0.00121 828 eland 0.00069 1458 red hartebeest 0.00450 222 mtn. reedbuck 0.01282 78 common duiker 0.07099 14 western african elephant 0.0002 24587 springbok 0.00404 248 spekboom black rhinoceros 0.00348 287 klipspringer 0.02857 35 noorsveld cape mtn. zebra 0.00241 416 steenbok 0.09583 10 burchell’s zebra 0.01263 79 cape buffalo 0.00142 706 warthog 0.00606 165 kudu 0.01065 94 red hartebeest 0.00410 244 eland 0.00056 1782 common duiker 0.06389 16 mtn. reedbuck 0.01166 86 noorsveld african elephant 0.00020 5004 common duiker 0.06366 16 black rhinoceros 0.00347 288 springbok 0.02037 49 cape mtn. zebra 0.00220 454 steenbok 0.09549 10 burchell’s zebra 0.01157 86 cape buffalo 0.00130 770 warthog 0.00833 120 kudu 0.01061 94 red hartebeest 0.00563 178 eland 0.00051 1944 grassy bontveld african elephant 0.00252 397 springbok 0.00667 150 black rhinoceros 0.00801 125 oribi 0.01905 53 burchell’s zebra 0.01221 82 grysbok 0.02593 39 bushpig 0.00455 220 cape buffalo 0.00753 133 warthog 0.01733 58 kudu 0.00288 347 red hartebeest 0.01171 85 bushbuck 0.01197 84 common duiker 0.01728 58 eland 0.00093 1080 karroid bontveld african elephant 0.00210 477 springbok 0.00556 180 black rhinoceros 0.00617 162 steenbok 0.03086 32 burchell’s zebra 0.00842 119 grysbok 0.03086 32 bushpig 0.00379 264 cape buffalo 0.00519 193 warthog 0.01667 60 kudu 0.01886 53 red hartebeest 0.01126 89 bushbuck 0.01425 70 common duiker 0.02058 49 eland 0.00077 1296 karoo thicket african elephant 0.00157 635 klipspringer 0.07093 14 mosaic black rhinoceros 0.00301 332 steenbok 0.02546 39 cape mtn. zebra 0.00198 504 cape buffalo 0.00467 214 bushpig 0.00284 352 kudu 0.00919 109 warthog 0.00333 300 bushbuck 0.01175 85 red hartebeest 0.00225 444 eland 0.00058 1728 common duiker 0.05517 18 mtn. reedbuck 0.00641 156 springbok 0.00417 240 karroid dwarf african elephant 0.00030 3336 springbok 0.03055 33 shrubland black rhinoceros 0.00480 208 klipspringer 0.02381 42 cape mtn. zebra 0.00176 567 steenbok 0.13194 8 burchell’s zebra 0.00926 108 cape buffalo 0.00104 963 warthog 0.01250 80 kudu 0.00309 324 red hartebeest 0.00845 118 eland 0.00077 1296 common duiker 0.01852 54 table 4 (continued) mammal species density est. spat. species density est. spat. habitat (ind./ha) req. (ind./ha) req. class (ha/ind.) (ha/ind.) boshoff.qxd 2005/12/09 09:47 page 101 koedoe 45/2 (2002) 102 issn 0075-6458 table 4 (continued) mammal species density est. spat. species density est. spat. habitat (ind./ha) req. (ind./ha) req. class (ha/ind.) (ha/ind.) karroid broken african elephant 0.00034 2919 springbok 0.03492 29 veld black rhinoceros 0.00364 275 klipspringer 0.02449 41 cape mtn. zebra 0.00252 397 steenbok 0.10000 10 burchell’s zebra 0.01323 76 cape buffalo 0.00148 674 warthog 0.00857 117 kudu 0.01111 90 black wildebeest 0.00155 644 eland 0.00088 1134 red hartebeest 0.00579 173 mtn. reedbuck 0.01648 61 common duiker 0.01905 53 sour grassland cape mtn. zebra 0.01905 53 grey rhebok 0.06000 17 red hartebeest 0.00405 247 cape buffalo 0.00280 357 common duiker 0.02222 45 eland 0.00370 270 oribi 0.02143 47 mtn. reedbuck 0.04615 22 grysbok 0.03333 30 mixed grassy african elephant 0.00085 1182 grysbok 0.07320 14 shrubland black rhinoceros 0.00266 376 grey rhebok 0.03922 26 cape mtn. zebra 0.01494 67 cape buffalo 0.00220 455 red hartebeest 0.01060 94 kudu 0.00813 123 common duiker 0.04880 20 eland 0.00218 459 klipspringer 0.06274 16 mtn. reedbuck 0.03017 33 steenbok 0.02614 38 fynbos cape mtn. zebra 0.00756 132 grey rhebok 0.03492 29 red hartebeest 0.00172 583 eland 0.00353 284 common duiker 0.06349 16 mtn. reedbuck 0.00488 205 grysbok 0.38094 3 riparian african elephant 0.00308 324 common duiker 0.16931 6 woodland black rhinoceros 0.00924 108 springbok 0.01429 70 burchell’s zebra 0.01299 77 steenbok 0.06349 16 bushpig 0.03896 26 cape buffalo 0.03204 31 warthog 0.06857 15 kudu 0.02822 35 red hartebeest 0.01158 86 bushbuck 0.02930 34 blue duiker 0.12698 8 eland 0.00198 504 dunefield bushpig 0.00135 743 grysbok 0.00658 152 common duiker 0.00439 228 bushbuck 0.00304 329 estimated requirement of 3400 ha for a troop of 80 individuals is derived from this information. herbivores the model’s estimations of the spatial requirements (and densities) for all species, except hippopotamus, in a single mhc are listed in table 4. the estimations for mhcs that can potentially carry hippopotamus are listed in table 5. the inclusion of hippopotamus within certain mhcs has, predictably, had the effect of lowering the densities and increasing the spatial requirements of all the bulk grazers in these habitats. thicket forest mosaic and thicket savanna mosaic can potentially carry the highest densities of elephant (0.54 individuals/km²). riparian woodland and thicket savanna mosaic can potentially carry the highest densities (0.8–0.9 individuals/km²) of black rhinoceros, as they can cape buffalo boshoff.qxd 2005/12/09 09:47 page 102 (2.1–3.2 individuals/km²) and common duiker (14.7–16.9 individuals/km²). in the case of the african elephant, a megaherbivore, social constraints (e.g., inter-bull aggression–kerley & boshoff 1997; whitehouse & kerley 2002) have not been violated by the model’s predictions. interaction between individuals of another megaherbivore, the black rhinoceros, provides a major constraint (adcock 1994) and a general minimum spatial (social) requirement of 200 ha/ animal has been suggested (see hall-martin & knight 1994). only four of the 16 density estimates provided by the model are above this suggested maximum density for this species; given the nature of the habitats in question, these estimates require field testing. it is noteworthy that in xeric succulent thicket in the andries vosloo kudu reserve, a density of as high as 1 male rhino/50 ha has been recorded (adcock 1994). it is difficult to obtain empirical data to test the estimates provided by the spreadsheet models. there is virtually no information for the study area, and where information is available, it is normally unsuitable due to a number of constraints. for example, populations of the larger mammal species in the current aenp (including the zuurberg portion), and indeed in other national parks and nature reserves in the region, are not natural and are influenced by factors such as presence of fencing (affecting population numbers, and limiting natural movements), and the absence of the large predators (influencing population structure and causing behavioural aberrations). thus, densities may be relatively high in protected areas due to a combination of pristine, or near-pristine, habitats and absence of the larger predators. in addition, the spatial requirements of the herbivores are known to vary between habitats, owing to spatial variation in forage quality and availability, and shelter. thus, until empirical studies have been conducted in the mhcs in the gaenp planning domain, testing of the model’s predictions is always going to be problematic. notwithstanding the contraints mentioned above, an attempt was made to compare empirically obtained spatial requirement data with the predictions from the model (table 6). with few exceptions, the data derived from the model were broadly corroborated for those herbivore species for which published information is available, thereby indicating that realistic values were generated by the model. it is again emphasised that the predictions from the model should be regarded as hypotheses and should be tested through field studies and modified where necessary (see also general discussion). discussion the spatial requirement and density data generated by the model described here are considered to be realistic. they can therefore be meaningfully used in the conservation planning exercise for the larger mammals in the gaenp planning domain. these data also provide useful information for guiding conservation management decisions, for example, determining multi-species assemblages and preliminary stocking rates of herbivores in the proposed gaenp. it is emphasised that the estimated densities or spatial requirements refer to a situation where the entire suite of potentially occurring species is available and present, and the habitats in which they can occur are in an “intact” or “potentially restorable” state. any deviation from this scenario, e.g., due to the unavailability of a species (for various reasons) or total habitat transformation, will require manipulation of the data and rerunning of the model. we recognise, however, that the model greatly oversimplifies the highly complex intraspecific and inter-specific mammal interactions, and the equally complex animal-plant relationships, the latter often being influenced by seasonality. there are, however, no alternatives when working at this scale, and with so little ecological information available for the species concerned. issn 0075-6458 103 koedoe 45/2 (2002) boshoff.qxd 2005/12/09 09:47 page 103 koedoe 45/2 (2002) 104 issn 0075-6458 table 5 estimated densities and spatial requirements values of the larger mammalian herbivores in areas of nine mammalian habitat classes in the gaenp planning domain that contain hippopotamus habitat. data derived from a spreadsheet model. see text for calculations and assumptions. scientific names in table 1 mammal species density est. spat. species density est. spat. habitat (ind./ha) req. (ind./ha) req. class (ha/ind.) (ha/ind.) thicket forest african elephant 0.00540 185 common duiker 0.06617 15 mosaic black rhinoceros 0.00361 277 grysbok 0.34740 3 bushpig 0.06818 15 cape buffalo 0.00104 963 hippopotamus 0.00198 504 bushbuck 0.16034 6 blue duiker 0.69481 1 thicket african elephant 0.005395 185 springbok 0.014286 70 savanna black rhinoceros 0.00807 124 oribi 0.040816 25 mosaic burchell’s zebra 0.005772 173 steenbok 0.061224 16 bushpig 0.00974 103 grysbok 0.061224 16 warthog 0.011429 88 cape buffalo 0.011571 86 hippopotamus 0.005527 181 kudu 0.02466 41 red hartebeest 0.007722 130 bushbuck 0.102432 10 blue duiker 0.122449 8 eland 0.001984 504 common duiker 0.147958 7 reedbuck 0.011429 88 zuurberg mesic african elephant 0.00229 436 common duiker 0.07081 14 thicket black rhinoceros 0.00386 259 grysbok 0.10621 9 cape mtn. zebra 0.00112 893 cape buffalo 0.00066 1516 bushpig 0.02897 35 kudu 0.01180 85 hippopotamus 0.00173 577 bushbuck 0.04902 20 blue duiker 0.21242 5 eland 0.00182 551 succulent african elephant 0.00111 898 common duiker 0.06092 16 thicket black rhinoceros 0.00332 301 klipspringer 0.07833 13 cape mtn. zebra 0.00050 1985 grysbok 0.02948 34 bushpig 0.01407 71 cape buffalo 0.00030 3371 warthog 0.00571 175 kudu 0.01015 98 hippopotamus 0.00078 1283 bushbuck 0.04218 24 red hartebeest 0.00386 259 eland 0.00088 1134 blue duiker 0.05896 17 spekboomveld african elephant 0.00142 706 common duiker 0.06128 16 black rhinoceros 0.00334 299 grysbok 0.09192 11 burchell’s zebra 0.00046 2178 cape buffalo 0.00028 3531 bushpig 0.01791 56 kudu 0.01021 98 warthog 0.00727 138 bushbuck 0.04242 24 hippopotamus 0.00074 1344 eland 0.00112 891 blue duiker 0.06566 15 western african elephant 0.00022 4587 springbok 0.00404 248 spekboom black rhinoceros 0.00348 287 klipspringer 0.02857 35 noorsveld cape mtn. zebra 0.00180 554 steenbok 0.09583 10 burchell’s zebra 0.00689 145 cape buffalo 0.00106 942 warthog 0.00606 165 kudu 0.01065 94 hippopotamus 0.00203 493 eland 0.00056 1782 red hartebeest 0.00410 244 mtn. reedbuck 0.01166 86 common duiker 0.06389 16 noorsveld african elephant 0.00020 5004 common duiker 0.06366 16 black rhinoceros 0.00347 288 springbok 0.02037 49 boshoff.qxd 2005/12/09 09:47 page 104 table 5 (continued) mammal species density est. spat. species density est. spat. habitat (ind./ha) req. (ind./ha) req. class (ha/ind.) (ha/ind.) issn 0075-6458 105 koedoe 45/2 (2002) cape mtn. zebra 0.00165 605 steenbok 0.09549 10 burchell’s zebra 0.00631 158 cape buffalo 0.00097 1027 warthog 0.00833 120 kudu 0.01061 94 hippopotamus 0.00186 538 eland 0.00051 1944 red hartebeest 0.00563 178 karroid african elephant 0.00210 477 springbok 0.00556 180 bontveld black rhinoceros 0.00617 162 steenbok 0.03086 32 burchell’s zebra 0.00561 178 grysbok 0.03086 32 bushpig 0.00379 264 cape buffalo 0.00346 289 warthog 0.01667 60 kudu 0.01886 53 hippopotamus 0.00165 605 bushbuck 0.01425 70 red hartebeest 0.01126 89 eland 0.00077 1296 common duiker 0.02058 49 riparian african elephant 0.00308 324 common duiker 0.16931 6 woodland black rhinoceros 0.00924 108 springbok 0.01429 70 burchell’s zebra 0.00866 116 steenbok 0.06349 16 bushpig 0.03896 26 cape buffalo 0.01736 58 warthog 0.06857 15 kudu 0.02822 35 hippopotamus 0.00829 121 bushbuck 0.02930 34 red hartebeest 0.01158 86 eland 0.00198 504 blue duiker 0.12698 8 the advantages and disadvantages of using the lsu approach to estimate stocking rates, is discussed in some detail by boshoff et al. (2001b) and boshoff et al. (2002). we contend that the lsu-based approach is appropriate for estimating densities of medium to large-sized wild mammals at a mega-reserve (e.g., gaenp) scale, and that realistic values, that can be used for systematic conservation planning in the gaenp planning domain, have been generated. an alternative to the lsu approach for calculating stocking rates is the use of a standing crop biomass of animals as an index of carrying capacity. in savanna regions, often exhibiting high rainfall and nutrient rich soils, primary production and animal density are generally positively correlated with mean annual rainfall (coe et al. 1976). however, soil type influences and further complicates this relationship, even in the savannas, and the biomass of large ungulates can be as much as 20 times lower on nutrient poor soils (fritz & duncan 1994). the fact that savannas with nutrient rich soils support different kinds of vegetation and also different types and densities of herbivores from those with nutrient poor soils has been emphasised by bell (1982). given the high regional variation in rainfall, soil type (ranging from nutrient poor to nutrient rich soils) and presumably primary productivity, in the gaenp planning domain, this approach was not attempted in the present study. there is strong evidence that a high density of elephants in the “addo bush” habitat (spekboomveld mhc) has a negative impact on the cover, architecture and diversity of the plants (barratt & hall-martin 1991; johnson et al. 1999; lombard et al. 2001; moolman & cowling 1994; penzhorn et al. 1974; stuart-hill 1992; stuart-hill & aucamp 1993), summarised in cowling & kerley (2002). the only published recommended density for elephant in the aenp, and specifically boshoff.qxd 2005/12/09 09:47 page 105 koedoe 45/2 (2002) 106 issn 0075-6458 ta bl e 6 c om pa ri so ns b et w ee n th e pr ed ic tio ns o f s pa tia l r eq ui re m en ts fr om th e sp re ad sh ee t m od el ( se e te xt ) an d fr om a va ila bl e em pi ri ca l d at a fo r si m ila r ha bi ta ts b ey on d th e g a e n p pl an ni ng d om ai n. s ci en tif ic n am es in t ab le 1 m am m al s pe ci es b la ck r hi no ce ro s c ap e m ou nt ai n ze br a w ar th og b lu e du ik er c om m on d ui ke r k lip sp ri ng er o ri bi e m pi ri ca l d at a( ha /in di vi du al ) a nd ri es v os lo o k ud u r es er ve : 50 h a/ in di vi du al to 2 00 h a/ in di vi du al ( a dc oc k 19 94 ). m ea n of 4 .7 i nd iv id ua ls p er b re ed in g he rd i n m z n p (p en zh or n 19 75 ); m ea n ho m e ra ng e pe r br ee di ng h er d in m z n p is 9 10 h a (p en zh or n 19 82 ). = 19 4 ha /in di vi du al a nd ri es v os lo o k ud u r es er ve ( so m er s 19 92 ) 12 .3 2 an im al s/ km ² ( fr om 8 00 a ni m al s pe r 64 93 h a) = 8 ha /in di vi du al 0. 5– 1 ha /in di vi du al ( a pp s 19 96 ) 5. 5– 8 ha /in di vi du al ( h an ek om & w ils on 1 99 1) 1. 8– 11 h a/ in di vi du al ( v on g ad ow 1 97 8) 17 h a/ in di vi du al , an d as l ow a s 20 –5 0 ha /in di vi du al (a lle nr ow la nd so n 19 86 ). 20 –5 0 ha /in di vi du al ( r ow er ow e 19 91 ). e as te rn c ap e th ic ke t: es t. le ss th an 3 ha /in di vi da l (f ur st en bu rg & k le yn ha ns 1 99 6) . 11 –1 5 ha /in di vi du al ( n or to n 19 80 ) e as te rn c ap e: 1 2. 5 ha /in di vi du al ( in cr ea si ng p op ul at io n so c ou ld b e hi gh er d en si ty ) (v an t ey lin ge n & k er le y 19 95 ). 8 to 3 0 ha /in di vi du al ( r ow er ow e 19 88 ) an d 6. 3 to 5 7. 6 ha /in di vi du al ( e ve re tt et a l. 19 91 ). pr ed ic tio ns f ro m th e m od el ( ha /in di vi du al ) r an ge : 1 08 -3 76 r an ge : 5 313 73 r an ge : 1 510 63 r an ge : 1 -1 7 r an ge : 6 -2 28 r an ge : 1 342 r an ge : 3 363 n ot es t he e st im at es f ro m t he m od el o ve rl ap w ith e m pi ri ca l da ta . t he e m pi ri ca l v al ue s fa ll w ith in th e m od el 's ra ng e. a v k r d en si ty c on si de re d to b e ar tif ic ia lly h ig h du e to ab se nc e of th e la rg e pr ed at or s, e sp ec ia lly li on . t he e m pi ri ca l va lu es c or re sp on d w el l w ith t he m od el 's ra ng e. t he e m pi ri ca l va lu es f al l w ith in t he m od el 's ra ng e. t he em pi ri ca l v al ue s fa ll w ith in th e m od el 's ra ng e. t he e m pi ric al v al ue is s im pl y an e st im at e. t he e m pi ri ca l va lu es b ro ad ly c or re sp on d w ith t he l ow er en d (i .e . h ig he st d en si ty ) of th e m od el 's ra ng e. t hi s em pi ri ca l s tu dy w as c on du ct ed in p ri m e ha bi ta t a nd on e w ith a re la tiv el y hi gh ra in fa ll. t he em pi ri ca lly ob ta in ed v al ue s ov er la p or c or re sp on d w ith t he m od el 's pr ed ic tio ns . boshoff.qxd 2005/12/09 09:47 page 106 issn 0075-6458 107 koedoe 45/2 (2002) ta bl e 6 (c on tin ue d) g ry sb ok g re y rh eb ok k ud u b us hb uc k m ou nt ai n re ed bu ck r an ge : 3 -1 52 r an ge : 1 729 r an ge : 3 332 4 r an ge : 6 -3 29 r an ge : 2 220 5 1. 3– 9. 4 ha /in di vi du al ( m an so n 19 74 ). 15 –1 52 h a/ in di vi du al ( fe rr ei ra 1 98 4) . 15 h a/ in di vi du al ( b eu ke s 19 87 ). a dd o e le ph an t n at io na l pa rk : m ea n of 5 27 a ni m al s in 13 6 42 h a = 26 h a/ in di vi da l ( fr om s a n p) . m z n p: c a 10 0 an im al s in 6 50 0 ha = 6 5 ha /in di vi du al (f ro m s a n p) . a nd ri es v os lo o k ud u r es er ve (a lle n r ow la nd so n 19 80 ). = 17 k ud u/ km ² o r 5. 9 ha /in di vi du al . 20 h a/ in di vi du al ( a lle nr ow la nd so n 19 86 ) s c ap e: 1 4– 20 h a/ in di vi du al ( se yd ac k 19 84 ). 33 h a/ in di vi du al ( o de nd aa l & b ig al ke 1 97 9) e c ap e: 7 7 ha /in di vi du al ( st ua rt -h ill & d an ck w er ts 19 88 ). m z n p: 4 09 a ni m al s in 6 50 0 ha = 16 h a/ in di vi du al ( fr om s a n p) . t he e m pi ri ca lly o bt ai ne d va lu es o ve rl ap w ith th e m od el 's pr ed ic tio ns . t he e m pi ri ca lly o bt ai ne d va lu es o ve rl ap o r co rr es po nd cl os el y w ith th e m od el 's pr ed ic tio ns . t he m od el p re di ct s re la tiv el y lo w d en si tie s; i t m ay b e ov er es tim at in g th e sp at ia l re qu ir em en ts , fo r re as on s no t un de rs to od b ut p os si bl y lin ke d to k ud u fe ed in g ec ol og y, or th e ku du d en si tie s in th es e re se rv es m ay b e ar tif ic ia lly hi gh d ue to a n ab se nc e of la rg e pr ed at or s. in a ll ca se s qu ot ed , th e em pi ri ca l va lu es f al l w ith in t he m od el 's ra ng e. t he m z n p co ns er ve s pr im e ha bi ta t f or th is s pe ci es . boshoff.qxd 2005/12/09 09:47 page 107 koedoe 45/2 (2002) 108 issn 0075-6458 for the spekboomveld habitat class, is derived from research by national parks scientists during the early 1970s, when a density of 0.4 elephants per km² was proposed (penzhorn et al. 1974). a significantly lower density of 0.142 elephants per km² was estimated for this habitat by our spreadsheet model. the highest densities predicted by the model, namely 0.54 elephants/km², are for the thicket forest mosaic and the thicket savanna mosaic. the basis for the fourfold higher density of 2 elephants/km² recommended by hall-martin & barratt (1991), and adopted by knight et al. (2002), is questionable (cowling & kerley 2002). it needs to be emphasised that even though the model has attempted to address the issue of seasonality for certain species (by reducing the amount of allocated forage), it is important that the gaenp be managed as a single spatial unit, in order to provide maximum opportunity for movements by nomadic or migratory species, on a yearround basis. this will cater for ecological factors such as presence of surface water, seasonal food availability and the possible negative effects of selective foraging on threatened plants. population sizes methods the potential population sizes of the 44 mammal species were calculated by simply multiplying the density estimate for each species with the area (in hectares) of the terrestrial habitats (i.e., mhcs), or length (km), in the case of rivers or coastline. fig. 27. fig. 28. fig. 29. fig. 30. fig. 31. boshoff.qxd 2005/12/09 09:47 page 108 issn 0075-6458 109 koedoe 45/2 (2002) these data were calculated for two park planning scenarios, namely: slightly modified kerley & boshoff (1997) planning domain; and the gaenp planning domain; and according to two habitat transformation categories, namely: “intact” and “restorable”. fig. 32. fig. 33. fig. 34. fig. 35. fig. 36. box 2: estimated population sizes of the two otter species based on the information given in table 3, it was decided to use a linear density of 1 individual/3 km of river or coastal habitat for both species of otter, namely cape clawless otter and spotted-necked otter. note that the transformation of landscapes did not affect the availability of rivers for otter conservation, i.e., all rivers were considered as “intact” habitat for otters. see “spacial requirements” for an explanation of methodology followed. cape clawless otter 183 km of potentially suitable river and coastal habitat for cape clawless otter alone = 61 individuals 342 km of potentially suitable river habitat equally shared with spottednecked otter; results in 171 km available for cape clawless otter = 57 individuals. grand total = 118 individuals spotted-necked otter 342 km of potentially suitable river habitat equally shared with cape clawless otter; results in 171 km available for spotted-necked otter = 57 individuals. grand total = 57 individuals boshoff.qxd 2005/12/09 09:47 page 109 koedoe 45/2 (2002) 110 issn 0075-6458 results the potential population sizes, according to two park planning domain scenarios are listed for all species in table 7. the population estimates for the two otter species are presented in box 2. discussion the estimates of populations sizes provided in table 7 and in box 2, can be used to measure the degree to which mammal population targets, that are set as part of a separate conservation planning exercise (e.g., kerley et al. in prep.), are met by the various planning domain scenarios, taking into account different transformation categories. the estimated population sizes for two species require comment. first, the estimates for the klipspringer require closer scrutiny, as the values appear to be somewhat high. in the mhcs where this species has been marked as being “resident” it may be more appropriate, in a future study, to map individual habitat patches. second, the estimates for the kudu may be too low. the reasons for this are not known but may be linked with this species’ particular feeding ecology. a similar pattern was observed in the analysis for the cape floristic region (boshoff et al. 2002). the estimates for some of the other smaller ungulates may at face value also appear to be high (e.g., grysbok: jg castley pers. comm.). we again emphasise that the model’s outputs represent potential densities in intact habitats, and fig. 37. fig. 38. fig. 39. fig. 40. fig. 41. boshoff.qxd 2005/12/09 09:47 page 110 issn 0075-6458 111 koedoe 45/2 (2002) table 7 the estimated potential total population sizes of the mediumto large-sized mammals, according to two gaenp planning domain scenarios, and according to two transformation categories. scientific names in table 1 estimated number of animals species modified kerley & boshoff (1997) gaenp planning domain planning domain intact restorable total intact restorable total chacma baboon 3885 1504 5388 5477 2233 7710 vervet monkey 60440 28480 88920 90504 46297 136801 porcupine 15915 6328 22242 22957 10250 33207 aardwolf 356 134 490 536 227 763 brown hyaena 37 15 52 55 24 79 spotted hyaena 37 17 55 56 28 84 cheetah 3 3 6 6 4 10 leopard 25 10 36 37 16 53 lion 40 19 58 60 30 90 caracal 85 36 121 126 59 184 african wild cat 1696 720 2416 2554 1189 3743 small spotted cat 130 83 213 205 129 334 serval 36 18 54 49 28 78 bat-eared fox 3280 2137 5416 5291 3472 8763 wild dog 35 15 50 52 25 77 cape fox 282 177 458 448 282 731 black-backed jackal 418 176 594 619 287 907 honey badger 48 19 67 70 31 101 aardvark 45 23 68 70 38 108 african elephant 322 141 464 523 248 771 black rhinoceros 766 401 1167 1250 677 1927 cape mountain zebra 851 161 1012 1083 245 1328 burchell’s zebra 996 805 1801 1724 1324 3048 bushpig 3316 958 4274 4845 1557 6402 warthog 1325 820 2145 2206 1362 3569 hippopotamus 27 29 55 32 33 64 black wildebeest 24 8 32 43 21 64 red hartebeest 874 460 1334 1369 755 2124 blue duiker 24849 7824 32673 36277 12352 48628 common duiker 14479 6843 21322 21777 11204 32981 springbok 1666 1022 2689 2929 1815 4744 klipspringer 3574 1293 4867 4836 1954 6790 oribi 702 510 1213 1234 929 2162 steenbok 7967 6213 14181 13288 9958 23246 grysbok 21964 3693 25657 28569 6296 34865 grey rhebok 2122 88 2210 2393 106 2499 cape buffalo 582 414 996 1058 732 1789 kudu 2023 1120 3143 3277 1892 5169 bushbuck 7684 2965 10649 11577 5074 16652 eland 340 84 424 470 144 614 reedbuck 103 128 231 219 241 460 mountain reedbuck 1528 423 1951 1980 639 2618 boshoff.qxd 2005/12/09 09:47 page 111 therefore cannot be equated with current densities in transformed habitats. the data in table 7 indicate that potential populations of herbivore species that are more prevalent in the “upland” areas are less impacted by transformed (but potentially restorable) habitats than are herbivore species that are more prevalent in “lowland” areas. for example, the steenbok is markedly affected by transformation in the “lowland” areas, whereas the grey rhebok is not. this pattern is understandable, given that most of the transformation has occurred in the “lowland” areas. general discussion the information generated by the present study provides realistic guidelines for the testing of population targets for medium to large-sized mammals that can potentially occur in the gaenp planning domain, and for the identification of species for which metapopulation management may be required for their conservation. it will also guide park managers regarding species that are no longer present in the general gaenp domain but that can be considered for reintroduction, and in the maintenance of realistic densities. the information in this report koedoe 45/2 (2002) 112 issn 0075-6458 fig. 42. fig. 43. fig. 44. boshoff.qxd 2005/12/09 09:47 page 112 will therefore make a significant contribution to achieving the overall conservation goals of the gaenp. it is important that the estimates derived by this study be treated as hypothetical guidelines at this stage. thus, any management action based on these estimates should be considered experimental, should be tested through adaptive management strategies and should be closely monitored. the need to test indigenous herbivore spatial requirement/density estimates in practice, and to adapt them in the light of field experience, has been mentioned elsewhere (trollope 1990). in addition, the final stocking rates for these herbivores should be conservative, in order to cope with unfavourable conditions (trollope 1990). we thus advocate a “management by hypothesis” approach, with assumptions and predictions being explicitly tested. a major advantage of the estimates presented here is that the assumptions are explicitly quantitative and can be modified as these ideas are tested, allowing adaptive management principles and actions to be employed. the concepts of “management by hypothesis” and “adaptive management” are a generally accepted approach to dealing with management challenges associated with a paucity of information (bowman 1995; macnab 1983; may 1991). acknowledgements we are grateful to the following persons for providing insights, information and assistance in the development of this report: rebecca sims-castley (teru, university of port elizabeth); amanda lombard (conservation systems, knysna); mike knight, guy castley, lucius moolman, john adendorff (south african national parks); lloyd wingate and fred kigozi (amatole museum, king william’s town). guy castley provided additional valuable comments on a draft of this paper. the gis tasks involved in collapsing the 43 original land classes to form 21 mammal habitat classes (mhc), and the calculation of the areas of the mhcs that comprised, or did not comprise, habitat for hippopotamus, and also the areas of “intact” and “restorable” habitat, were conducted by csirenvironmentek, in a consortium with teru, as part of a gef co-funded contract to south african parks to provide conservation planning services. we thank jeanne nel, inge kotze, sarah davies and dirk roux, all of csir-environmentek, for their patience and co-operation in providing teru with these gis outputs. south african national parks is thanked for permission to publish this material. references adcock, k. 1994. the relevance of “territorial” behaviour in black rhino to their population management. pp. 82-86. in: penzhorn, b.l. & n.p.j. kriel (eds.). proceedings of a symposium on rhinos as game ranch animals. pretoria: onderstepoort. allen-rowlandson, t.s. 1980. the social and spatial organisation of the greater kudu (tragelaphus strepsiceros pallas, 1766) in the andries vosloo kudu reserve, eastern cape. msc dissertation, rhodes university, grahamstown. allen-rowlandson, t.s. 1986. an autecological study of bushbuck and common duiker in relation to forest management. ph.d. thesis, universtity of natal, pietermaritzburg. anonymous, 1985. the national grazing strategy. pretoria: department of agriculture. apps, p. 1996. smithers’ mammals of southern africa: a field guide. halfway house: southern book publishers. barratt, d.g. & a. hall-martin. 1991. the effect of indigenous browsers on valley bushveld in the addo elephant national park. pp. 14-16. in: zacharias, p.j.k., g.c. stuart-hill & j. midgley (eds.). proceedings of the first valley bushveld symposium. grassland society of south africa. howick. bell, r.h.v. 1982. the effect of soil nutrient availability on community structure in african ecosystems. pp. 193-216. in: huntley, b.j. & b.h. walker (eds.). ecology of tropical savannas. berlin: springer-verlag. beukes, p.c. 1987. responses of grey rhebuck and bontebok to controlled fires in coastal renosterveld. south african journal of wildlife research 17: 103-108. boshoff, a.f. & g.i.h. kerley. 2001. potential distributions of the mediumto large-sized mammals of the cape floristic region based on historical accounts and habitat requirements. african zoology 36: 245-273. boshoff, a.f., g.i.h. kerley, r.m. cowling & s.l. wilson. 2001a. conservation planning in the greater addo national park: a review of the species-, populationand spatially-driven issn 0075-6458 113 koedoe 45/2 (2002) boshoff.qxd 2005/12/09 09:47 page 113 processes involving the mediumto large-sized mammals. terrestrial ecology research unit (university of port elizabeth), report no. 34. boshoff, a.f., g.i.h. kerley & r.m. cowling. 2001b. a pragmatic approach to estimating the distributions and spatial requirements of the mediumto large-sized mammals in the cape floristic region, based on historical accounts and habitat requirements. diversity and distributions 7: 29-43. boshoff, a.f., g.i.h. kerley & r.m. cowling. 2002. estimated spatial requirements of the mediumto large-sized mammals, according to broad habitat units, in the cape floristic region, south africa. african journal of range and forage science 19: 27-42. bowman, d. 1995. down in the forest something stirred. new scientist (9 december) 2007: 54. butterfield, b.r., b. csuti & j.m. scott. 1994. modeling vertebrate distributions for gap analysis. pp. 53-68. in: miller, r.i. (ed.). mapping the diversity of nature. london: chapman & hall. campbell, b.m. 1986. plant spinescence and herbivory in a nutrient-poor ecosystem. oikos 47: 168-172. castley, j.g., a.f. boshoff & g.i.h. kerley. 2001. compromising south africa’s natural biodiversity—inappropriate herbivore introductions. south african journal of science 97: 344-348. caughley, g. 1994. directions in conservation biology. journal of animal ecology 63: 215-244. caughley, g. & a.r.e. sinclair. 1994. wildlife ecology and management. london: blackwell science. chew, r.m. 1978. the impact of small mammals on ecosystem structure and function. pp. 167-180. in: snyder, d.p. (ed.) pymatuning symposium on populations of small mammals under natural conditions. vol. 5. spec. publ. ser. pymatuning laboratory of ecology. university of pittsburgh. coe, m.j., d.m. cumming & j. phillipson. 1976. biomass and production of large african herbivores in relation to rainfall and primary production. oecologia 22: 341-354. coetzee, p.w. 1979. present distributions and status of some of the mammals of albany. grahamstown: albany divisional council. collinson, r.f.h. & p.s. goodman. 1982. an assessment of range conditions and large herbivore carrying capacity of the pilanesberg game reserve, with guidelines and recommendations for management. inkwe 1: 1-55. cowling, r.m. & g.i.h. kerley. 2002. impacts of elephants on the flora and vegetation of subtropical thicket in the eastern cape. pp. 55-72. in: kerley, g.i.h., s.l. wilson & a. massey (eds.). elephant conservation and management in the eastern cape: workshop proceedings. terrestrial ecology research unit (university of port elizabeth) report no. 35: devore, i. & k.r.l. hall. 1965. baboon ecology. in: de vore, i. (ed.) primate behaviour: field studies of monkeys and apes. new york: holt, rinehart & winston. du plessis, s.f. 1969. the past and present geographical distribution of the perissodactyla and artiodactyla in southern africa. m.sc. thesis, university of pretoria, pretoria. everett, p.s., m.r. perrin & d.t. rowe-rowe. 1991. responses by oribi to different range management practises. south african journal of wildlife research 21:114-118. ferreira, n.a. 1984. various aspects of the ecology of the grey rhebuck. pelea 3: 7-13. fritz, h. & p. duncan. 1994. on the carrying capacity for large ungulates of african savanna ecosystems. proceedings of the royal society, london 256: 77-82. furstenburg, d. & m. kleynhans. 1996. kudu and bushbuck in boergoat production systems in valley bushveld: ecological interactions and constraints. pelea 15:5-13. grossman, d. 1991. carrying capacity, game numbers and optimal utilisation. seminar: game ranch management. potchefstroom: potchefstroom university. hall-martin, a.j. & d.g. barratt. 1991. the effect of elephants loxodonta africana on thicket vegetation in the addo elephant national park. internal report, south african national parks, pretoria. hall-martin, a.j. & m.h. knight. 1994. conservation and management of black rhinoceros in south african national parks. pp. 11-19. in: penzhorn, b.l. & n.p.j. kriel (eds.). proceedings of a symposium on rhinos as game ranch animals. pretoria: onderstepoort. hanekom, n. & v. wilson. 1991. blue duiker philantomba monticola densities in the tsitsikamma national park, and probable factors limiting these populations. koedoe 34: 107-120. hewitt, j. 1931. a guide to the vertebrate fauna of the eastern cape province. grahamstown: albany museum. johnson, s.d. 1992. plant-animal relationships. pp. 175-205. in: cowling, r.m. (ed.) the ecology of fynbos: nutrients, fire and diversity. cape town: oxford university press. johnson, c.f., r.m. cowling & p.b. phillipson. 1999. the flora of the addo elephant national park, south africa: are threatened species vulnerable to elephant damage. biodiversity and conservation 8: 1447-1456. kerley, g.i.h. & a.f. boshoff. 1997. a proposal for a greater addo national park: a regional and national conservation and development opportukoedoe 45/2 (2002) 114 issn 0075-6458 boshoff.qxd 2005/12/09 09:47 page 114 nity. terrestrial ecology research unit, university of port elizabeth; (internal report no. 17). kerley, g.i.h. & a.f. boshoff. 2002. the greater addo national park initiative: linking biodiversity to socio-economic development. pp. 57-66. in: pierce, s.m., r.m cowling, & t. sandwith. (eds.). mainstreaming biodiversity in development: case studies from south africa. washington, dc: international bank for reconstruction and development/the world bank. knight, m., g. castley, l. moolman & j. adendorff. 2002. elephant management in addo elephant national park. pp. 32-40. in: kerley, g.i.h., s.l. wilson & a. massey (eds.). elephant conservation and management in the eastern cape: workshop proceedings. (terrestrial ecology research unit, university of port elizabeth; internal report no. 35). kruger, l. & c. sykes. 2002. field mapping of land classes and land transformation. pp. 218262. in: conservation planning framework for the greater addo elephant national park (part 1) – final (internal) report. south african national parks. lande, r. & g.f. barrowclough. 1987. effective population size, genetic variation, and their use in population management. pp. 87-123. in: soulé, m.e. (ed.). viable populations for conservation. cambridge: cambridge university press. lloyd, p.h. & j.c.g. millar. 1983. a questionnaire survey (1969-1974) of some of the larger mammals of the cape province. bontebok 3: 1-49. lombard, a.t., c.f. johnson, r.m. cowling & r.l. pressey. 2001. protecting plants from elephants: botanical reserve scenarios within the addo elephant national park, south africa. biological conservation. 102: 191-203. low, a. b. & a. g. rebelo. 1996. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. macdonald, i.a.w. 1992. vertebrate populations as indicators of change in southern africa. transactions of the royal society of south africa 48: 87-122. macnab, j. 1983. wildlife management as scientific experimentation. wildlife society bulletin 11: 397-401. manson, j. 1974. aspekte van die biologie en gedrag van die kaapse grysbok raphicerus melanotis. m.sc. dissertation. university of stellenbosch. stellenbosch. may, r.m. 1991. the role of ecological theory in planning re-introduction of endangered species. symposium of the zoological society of london 62: 145-163. meissner, h.h. 1982. theory and application of a method to calculate forage intake of wild southern african ungulates for purposes of estimating carrying capacity. south african journal of wildlife research 12: 41-47. moen, a.n. 1973. wildlife ecology. san francisco: freeman. moolman, h.j. & r.m. cowling. 1994. the impact of elephant and goat grazing on the endemic flora of south african succulent thicket. biological conservation 68: 53-61. morris, c.d., m.b. hardy & p.e. bartholomew. 1999. stocking rate. pp. 186-193. in: tainton, n. (ed.). veld management in south africa. pietermaritzburg: university of natal press. norton, p.m. 1980. the habitat and feeding ecology of the klipspringer oreotragus oreotragus in two areas of the cape province. m.sc. dissertation. university of pretoria. pretoria. odendaal, p.b. & r.c. bigalke. 1979. home range and groupings of bushbuck in the southern cape. south african journal of wildlife research 9:96101. penzhorn, b.l. 1975. behaviour and population ecology of the cape mountain zebra in the mountain zebra national park. d.sc. thesis. university of pretoria. penzhorn, b.l. 1982. home range sizes of cape mountain zebras equus zebra zebra in the mountain zebra national park. koedoe 25: 103108. penzhorn, b.l., p.j. robbertse & m.c. olivier. 1974. the influence of the african elephant on the vegetation of the addo elephant national park. koedoe 17: 137-158. rookmaaker, l.c. 1989. the zoological exploration of southern africa 1650-1790. rotterdam: balkema. rowe-rowe, d.t. 1988. oribi natal’s most threatened antelope. pietermaritzburg: natal parks board. (wildlife management technical guides for farmers no. 20.) rowe-rowe, d.t. 1991. the ungulates of natal. pietermaritzburg: natal parks board. seydack, a.h.w. 1984. application of a photorecording device in the census of larger rain-forest mammals. south african journal of wildlife research 14: 10-14. shortridge, g.c. 1942. field notes on the first and second expeditions of the cape museums mammal survey of the cape province and descriptions of some new subgenera and species. annals of the south african museum 36: 27-100. skead, c.j. 1987. historical mammal incidence in the cape province: vol. 2 – the eastern half of the cape province, including the ciskei, transkei and east griqualand. cape town: dept nature & environmental conservation. skinner, j.d. & r.h.n. smithers. 1990. the mammals of the southern african subregion. pretoria: university of pretoria. issn 0075-6458 115 koedoe 45/2 (2002) boshoff.qxd 2005/12/09 09:47 page 115 smithers, r.h.n. 1986. south african red data book: terrestrial mammals. pretoria: council for scientific and industrial research, national scientific programmes unit. (south african national science programmes report; no. 125). somers, m. 1992. the implications of social structure for the conservation and control of a warthog phacochoerus aethiopicus population in the andries vosloo kudu reserve, eastern cape province. m.sc. dissertation. university of pretoria, pretoria. soulé, m. (ed.) 1987. viable populations for conservation. cambridge: cambridge university press. stuart, c.t. 1981. notes on the mammalian carnivores of the cape province, south africa. bontebok 1: 1-58. stuart, c.t. 1985. the status of two endangered carnivores occurring in the cape province, south africa, felis serval and lutra maculicollis. biological conservation 32: 375-382. stuart-hill, g. c. 1992. effects of elephants and goats on the kaffrarian succulent thicket of the eastern cape, south africa. journal of applied ecology 29: 699-710. stuart-hill, g. c. & a.j. aucamp. 1993. carrying capacity of the succulent valley bushveld of the eastern cape. african journal of range and forage science 10: 1-10. stuart-hill, g. & j. danckwerts. 1988. influence of domestic and wild animals on the future of succulent valley bushveld. pelea 7: 45-46. stuart, c.t., i.a.w. macdonald & m.g.l. mills. 1985. history, current status and conservation of large mammalian predators in cape province, republic of south africa. biological conservation 31: 7-19. trollope, w.s.w. 1990. veld management with specific reference to game ranching in the grassland and savanna areas of south africa. koedoe 33: 77-87. van teylingen, k. & g.i.h. kerley. 1995. habitat characteristics of increasing and decreasing oribi subpopulations in the eastern cape province, south africa. south african journal of wildlife research 25:118-122. vlok, j.h. & d. euston-brown. 2002. subtropical thicket ecosystem planning (step) project: biological survey report (plants and birds). unpublished report, terrestrial ecology research unit, university of port elizabeth. von gadow, k. 1978. a pellet count of blue duiker and bushbuck in the knysna forests. south african forestry journal 107: 77-81. wilcox, b.a. 1982. in situ conservation of genetic resources: determinants of minimum area requirements. pp. 639-647. in: mcneely, j.a. & k.r. miller (eds.). national parks, conservation and development—the role of protected areas in sustaining society. washington dc: smithsonian institution. whitehouse, a.m. & g.i.h. kerley. 2002. retrospective assessment of long-term conservation management of elephants in addo elephant national park, south africa. oryx 36: 243-248. koedoe 45/2 (2002) 116 issn 0075-6458 fo re st t hi ck et f or es t m os ai c t hi ck et s av an na m os ai c z uu rb er g m es ic t hi ck et a dd o h ei gh ts m es ic t hi ck et su cc ul en t t hi ck et sp ek bo om ve ld e as te rn s pe kb oo m n oo rs ve ld w es te rn s pe kb oo m n oo rs ve ld n oo rs ve ld g ra ss y b on tv el d k ar ro id b on tv el d k ar oo t hi ck et m os ai c k ar ro id d w ar f sh ru bl an d k ar ro id b ro ke n v el d so ur g ra ss la nd m ix ed g ra ss y sh ru bl an d fy nb os r ip ar ia n w oo dl an d d un ef ie ld su nd ay s sa ltm ar sh appendix 1 adjusting stocking rates and proportional allocation of stocking opportunities to foraging guilds (see text for methods) adjusted stocking rate (lsu/ha) 0.017 0.111 0.143 0.059 0.059 0.048 0.045 0.056 0.045 0.042 0.056 0.056 0.042 0.042 0.048 0.050 0.059 0.048 0.143 0.004 0.000 bulk grazer 1 5 20 9 9 5 5 30 25 25 29 20 15 20 25 30 20 10 30 0 0 concentrate grazer 1 1 20 1 1 15 10 10 10 10 18 15 15 15 15 30 20 10 15 0 0 mixed feeder 29 27 15 25 25 15 20 10 10 10 18 15 15 15 15 20 20 20 15 20 0 browser 69 67 45 65 65 65 65 50 55 55 35 50 55 50 45 20 40 60 40 80 0 boshoff.qxd 2005/12/09 09:47 page 116 filelist convert a pdf file! page 1 page 2 page 3 abstract introduction materials and methods results discussion conclusion acknowledgements references about the author(s) peter chatanga school of agricultural, earth and environmental sciences, university of kwazulu-natal, westville, durban, south africa department of biology, faculty of science and technology, national university of lesotho, roma, lesotho erwin j.j. sieben school of agricultural, earth and environmental sciences, university of kwazulu-natal, westville, durban, south africa citation chatanga, p., sieben, e.j.j., 2019, ‘ecology of palustrine wetlands in lesotho: vegetation classification, description and environmental factors’, koedoe 61(1), a1574. https://doi.org/10.4102/koedoe.v61i1.1574 note: additional supporting information may be found in the online version of this article. online appendix 1; online appendix 2; online appendix 3; online appendix 4 and online appendix 5. original research ecology of palustrine wetlands in lesotho: vegetation classification, description and environmental factors peter chatanga, erwin j.j. sieben received: 18 apr. 2019; accepted: 30 aug. 2019; published: 31 oct. 2019 copyright: © 2019. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the description and classification of wetland vegetation is important for water resource management and biodiversity conservation as it provides an understanding of the wetland vegetation–environment relationships and information to interpret spatial variation in plant communities. this study discusses the vegetation of the palustrine wetlands of lesotho based on a phytosociological approach. data on vegetation and various environmental variables were collected using the braun-blanquet method and a standardised protocol developed for environmental information of wetlands in south africa. the data were analysed mainly by clustering and ordination techniques. twenty-two communities were found by the classification of the wetland vegetation. these communities were found to be diverse in terms of species richness. the ordination revealed that the wetland vegetation is mainly influenced by altitude, longitude, slope, soil parent material, landscape, inundation, potassium content, soil texture, total organic carbon, nitrogen, electrical conductivity and latitude. regarding species composition and diversity, plant communities in the highlands were more diverse and were distinctively different from those in the lowlands. high-altitude communities were also found to be dominated mainly by c3 plants, while those at low altitudes exhibited the dominance of c4 species. some communities were either restricted to the highlands or lowlands but others exhibited a wide ecological amplitude and occurred over an extensive altitudinal range. the diversity of most of the wetlands, coupled with their restricted habitat, distribution at high altitudes and their role in supplying ecosystem services that include water resources, highlights the high conservation value associated with these wetlands, particularly in the face of climate change and loss of biodiversity. conservation implications: the study can be invaluable to wetland scientists, managers, biodiversity conservationists, water resource managers and planners and vegetation ecologists in southern africa. about 70% of lesotho falls in the maloti-drakensberg, accounting for about 60% of the region, and this makes the study important in biodiversity conservation planning, particularly in the highlands. the wetlands in lesotho face severe anthropogenic pressures that include overgrazing and economic development. given that the lesotho highlands as a water catchment is not only important for lesotho, but also for south africa and namibia, the conservation of the associated wetlands and this critical water resource is indispensable. keywords: anthropogenic; biodiversity conservation; canonical ordination; climate change; maloti-drakensberg; plant community; palustrine wetlands; phytosociology; vegetation classification. introduction palustrine wetlands, which cover about 6% of the earth’s land surface, are among the most ecologically sensitive ecosystems and very important globally because of their unique role in biogeochemical cycles (junk et al. 2013; mitsch & gosselink 2015). because they support azonal vegetation that is distinct from the surrounding vegetation (mucina & rutherford 2006; sieben et al. 2016), wetlands are ecological ‘islands’ within terrestrial environments in different landscapes across the globe. the distinction results from the prolonged water logging that causes oxygen deficiency (hypoxia) or its total absence (anoxia) in the wetland soil, with subsequent chemical changes in soil characteristics (gopal 2015; mitsch & gosselink 2015). mucina and rutherford (2006) observed that the presence of water, whether seasonal or permanent, is the primary factor in creating wetland habitats and associated vegetation. nonetheless, it is not wetness per se that primarily influences the geochemistry and morphology of wetland soils, but rather the anaerobic conditions that result from prolonged soil saturation or flooding (collins 2005; kotze et al. 1996). montane palustrine wetlands are a special sub-division of freshwater palustrine wetlands that are embedded within terrestrial ecosystems in mountain areas (mucina & rutherford 2006). these wetlands are often rich in endemics because many species remain isolated in high-altitude habitats (junk et al. 2013). by providing a wide spectrum of ecosystem services, these ecosystems are of high ecological and socio-economic importance (chatterjee et al. 2010). high-altitude wetlands provide essential ecosystem services which help to sustain human life, conserve biodiversity, preserve the hydrological regime and combat the impacts of climate change and are thereby key to water, food and ecological security (singha 2011). montane wetlands are located in the headwaters of major river basins, providing water for many trans-boundary rivers and play an important role in the ecology and hydrology of the local environment and downstream systems. they also play a fundamental role in the overall global and regional water cycles (russi et al. 2013; sieben, mtshali & janks 2014). the montane wetlands of lesotho are important for the supply of ecosystem services, including biodiversity conservation, water resources, livestock grazing, harvestable plant products and environmental regulation. lesotho forms an important hydrological reservoir and watershed for several countries in southern africa (nüsser & grab 2002). for example, the montane wetlands in the country form part of the headwaters of one of the most important international watercourses in southern africa, the senqu-orange river (orasecom 2015), which is not only important for lesotho, but also for south africa and namibia. lesotho falls entirely within the catchment of this river system (orasecom 2015) and the country is drained by the river and its tributaries. moreover, the high-altitude wetlands of lesotho are critical in maintaining the water levels not only in dams supporting the lesotho highlands water project (lhwp), which transfers water to south africa (grab & deschamps 2004), but also in those supplying the local population. directly and indirectly, wetlands in lesotho are estimated to contribute about 22% of the country’s gross domestic products (gdps) and 30% of the total employment in the country (department of environment 2014). furthermore, by providing freshwater that is transferred to the most densely populated and industrialised area of gauteng, the wetlands in lesotho also play a key role in the south african economy. the most ubiquitous and conspicuous feature of the wetland environment, which also plays a critical role in wetland ecosystem functioning, is the vegetation (cronk & fennessy 2001; gopal 2016). as a result, the description of wetland habitats based on vegetation and the associated classifications are useful in strategic planning for the conservation of wetlands (mitsch & gosselink 2015). plants are such a critical component of wetlands that they are the most widely supported biotic indicator for assessing wetland condition (collins 2005). accordingly, vegetation is the most suitable feature to consider over a broad range of wetland habitats (sieben et al. 2014). within a single wetland, a large diversity of habitats may occur, and as a result, large differences in the types of vegetation may be observed in the same wetland (mitsch & gosselink 2000). therefore, wetland habitats can be classified based on the plant communities in the system (collins 2005; sieben, kotze & morris 2010a). plant species occurring in a wetland can be used as indicators of environmental conditions and ecological changes taking place in the system (mitsch & gosselink 2015) because wetland vegetation is azonal (brand, du preez & brown 2013) and responds quickly to local environmental changes (cronk & fennessy 2001). because the plant community gives a characterisation of habitat units within a wetland and also serves as habitat for associated animals, a detailed wetland vegetation description for a given wetland can be used as a proxy for the system’s biodiversity in general, highlighting the conservation value of the wetland (sieben et al. 2016). accordingly, the determination of wetland plants that are useful as biological indicators has become increasingly important for use in monitoring the integrity of specific wetlands (sieben et al. 2014). wetland typology provides useful information for water resource management and biodiversity conservation (sieben et al. 2014). given the importance of wetlands, recent studies (moor et al. 2017; sieben et al. 2016) have emphasised the need for more information about their species composition, ecology and distribution. although many studies have been carried out in the wetlands of surrounding south africa (e.g. brand et al. 2013; sieben et al. 2014, 2016), no recent detailed countrywide survey has been conducted on the wetland vegetation of lesotho. the current study is the first to provide a characterisation of the wetland vegetation in detail, covering many wetlands in the country. the aims of this study for the palustrine wetlands of lesotho are: to produce a phytosociological classification and name the wetland plant communities to map the distribution of the vegetation of the palustrine wetlands using the phytosociological method to explore and describe the wetland vegetation–environment relationships. materials and methods study design the vegetation of the palustrine wetlands of lesotho was characterised using a phytosociological approach. the selection of wetlands for sampling was performed in such a way that as much wetland variation as possible in the country was captured with the available time and resources. an attempt was also made to include wetlands in protected areas such as national parks and nature reserves. fieldwork was carried out during the wet (summer) season, which commenced in february 2017 and ended in march 2018. the study focused on accessible wetlands. in general, the method followed sieben et al. (2014) to make the results compatible with the south african national wetland vegetation database. although many wetlands are showing signs of degradation, where wetland density was high and the systems showed little observable variation, large or near-pristine wetlands were selected for the assessment. depending on the location, each wetland was classified as pristine, rural or urban (table 1). a 3 × 3 m representative sample plot (quadrat) was then located randomly in each visually distinct and homogenous vegetation type of each selected wetland where different attributes of the vegetation and environment were measured and recorded. this plot size has been recommended for grassy wetlands in southern africa (brown et al. 2013; sieben et al. 2014). the same plot size has also been recommended for overgrazed grasslands (brown et al. 2013), a situation that is common in some parts of lesotho where wetland disturbance by livestock grazing and trampling is widespread. the number of plots per wetland was dependent on the number of observable distinct and homogenous vegetation units within the wetland. table 1: environmental variables that were measured or assessed and included in the analysis of the vegetation of the palustrine wetlands of lesotho. assessment of vegetation and environmental variables in each plot, the braun-blanquet method, a protocol often used for collecting vegetation data in south africa (brown et al. 2013; sieben et al. 2014), was used for vegetation assessment. because this method has been applied in south africa for several decades, it is possible to make comparisons between current and historical data (brown et al. 2013), which allows for effective plant community comparisons. the method involves assessing wetland vegetation in a stratified manner where plots are placed randomly in each distinct plant community and the species composition is recorded by determining the species present, as well as estimating the cover for each species using a cover-abundance scale. placing vegetation plots in a stratified manner ensures that as much variation as possible in the wetland is captured, including the different categories of indicators species – obligate wetland (obl), facultative wetland (facw), facultative (fac), facultative upland (facu) and obligate upland (upl) (mitsch & gosselink 2015). vegetation structure was described by making estimations of the proportion of the plot covered by vegetation and visually estimating the average height of the vegetation. in case of inundation, the average vegetation height was estimated from the soil surface (sieben et al. 2014). during the survey, the plant species were identified with the help of botanical field guides (pooley 2003; van oudtshoorn 2014) and those that could not be identified in the field were collected and later taken to the roma herbarium (roml) at the national university of lesotho for identification. for each plot, in addition to the vegetation attributes, a standardised protocol developed for environmental information of wetlands in south africa was also used to systematically measure or assess a number of environmental variables that have been recommended for wetlands (sieben et al. 2014). in at least one plot per wetland, a soil sample was collected from the top 15 cm of the soil (vegetation rooting zone) using a soil auger. both vegetation and environmental data were recorded in a wetland field data collection form (online appendix 5). because of the limited funding for the study, soil sampling was limited to one plot per wetland. the plot chosen for soil sampling would be the one representing the most widespread distinct community in the wetland. the samples were packaged in airtight (zipped) plastic bags for further analysis as recommended by stohlgren et al. (1998). the soil samples were air-dried for at least 48 h and later analysed for different variables (table 1). the soil analyses were performed by the analytical laboratory service of the institute for commercial forestry research in pietermaritzburg, south africa. the environmental and soil variables included in the study, as well as the methods used for their measurement or assessment, are briefly described (table 1). while most variables were measured or assessed on site in all vegetation plots, additional soil variables (indicated with an asterisk in table 1) were measured in the laboratory and only on those plots where soil samples were collected. after describing the plots in each wetland, as much of the wetland would be surveyed to record species occurring within the wetland system but were not encountered in the plots. this was in an attempt to ensure a species list that is as complete as possible for the wetland system for additional floristic analysis. data analysis both vegetation and environmental data were captured in microsoft excel spreadsheets from which they were imported into pc-ord (mjm software design, gleneden beach. or, united states) and canoco (microcomputer power, ithaca, ny, united states) programmes for analysis. prior to importing into these programs for analysis, the braun-blanquet vegetation cover values were converted into percentage cover (mueller-dombois & ellenberg 1974; omar, maroyi & van tol 2016; van der maarel 1979). arcgis version 10.2 was used to map the distribution range of the wetland vegetation plots in the study area and to determine the distribution range of the wetland plant communities in lesotho. based on the recommendations by brown et al. (2013), species richness, diversity and evenness were determined. for each plot, these were determined by calculating shannon–weiner index (h′) and evenness index (e), as well as listing the species. the fraction of the plot that was occupied by species i was referred to as pi, and this value is an indication of the relative abundance of species i. the pi was used to calculate the h′ and e, which are surrogates of species diversity, by the following formulae (ludwig & reynolds 1988): where pi is the proportion of species i and ln is the natural logarithm. where s is the species richness (number of species). to elucidate the ecological patterns in the vegetation of the palustrine wetlands of lesotho, a number of multivariate statistical techniques were employed. for the data analyses of this multivariate ecological community, three main types of analyses were employed: (1) hierarchical cluster analysis (hca), (2) indicator species analysis (isa) and (3) canonical correspondence analysis (cca). these were performed mainly following sieben et al. (2014). classification and indicator species analysis cluster analysis was used to classify sites with respect to similarity or dissimilarity (van tongeren 1995). classification makes the use of similarity between vegetation plots such that plots which are more similar in species composition are grouped together. classification facilitates the description and ecological interpretation of plant communities from a large amount of vegetation data. classification and description of vegetation not only provide an understanding of the vegetation–environment relationships, but also information to interpret the spatial variation between plant communities (clegg & o’connor 2012). accordingly, to obtain vegetation typology of the palustrine wetlands of lesotho, agglomerative hca was performed on vegetation data to identify homogenous plant communities in the wetlands. this would enable vegetation plots that are similar in species composition to be grouped together, with emphasis on the relationships between them (mccune & mefford 2011; van tongeren 1995). the classification was performed by grouping vegetation plots based on species composition and abundance data. pc-ord version 6.0 was used for the classification (mccune & mefford 2011). non-transformed percentage plant cover data were used for the clustering. the best interpretable classification was produced by the combination of the sørenson’s (bray–curtis) similarity index and ward’s linkage method. the ward’s method has been reported to have an advantage of producing clearly defined clusters (pla et al. 2012). the plant communities obtained were named following the guidelines by brown et al. (2013). however, in cases where there was no clear dominant, the name was derived from the description (structure and distribution) of the community. indicator species analysis was used as an objective criterion for determining the optimal number of clusters in the final dendrogram. this was achieved by repeating the clustering algorithm while varying the number of clusters (dufrêne & legendre 1997) and the number that gave the lowest average p-value of the indicator species was used in the final dendrogram (peck 2010). the isa was also used in characterising different wetland plant communities obtained from the final clusters. indicator species with indicator values (ivs) greater than 20 and significant (p ≤ 0.05) in the monte carlo permutation test were considered real indicators (sieben et al. 2016) and were therefore listed for each cluster. the isa is often used to test the fidelity (faithfulness) of a species to a given community. the monte carlo permutation test (also available in pc-ord) was also used to test for the statistical significance of the fidelity of the indicator species to the communities (dufrêne & legendre 1997). the isa was also conducted in pc-ord. for each community obtained from the classification, the median and range for species richness, diversity index, and evenness and means for vegetation height and cover were determined. ordination another important feature of the analysis was the relationship between wetland plant communities and explanatory variables. consequently, to examine the influence of environmental variables and gradients on wetland vegetation, the vegetation and environmental variable data and the plant communities obtained from the classification were subjected to canonical ordination (ter braak & šmilauer 1998). this was performed twice: (1) constrained cca on all vegetation plots using species abundance data and environmental variables and (2) constrained cca only on those vegetation plots where soil samples had been taken, using species abundance data and more detailed explanatory data. data of log-transformed percentage plant cover were used for the ordination. the choice for cca for the ordination was based on the fact that the vegetation data were compositional and the gradient was greater than four units (lepš & šmilauer 2003). the canonical ordination was performed using canoco version 5.11 (ter braak & šmilauer 1998). the statistical significance of the constrained ordination (relationship of communities with environmental variables) was tested using the unrestricted monte carlo permutation test available in canoco (ter braak 1995). the primary objective of canonical ordination is to detect the main pattern in the species–environment or community–environment relationships (ter braak 1995). the cca helps to detect patterns of variation in the species data that can be explained best by the supplied environmental variables (mcgarigal, cushman & stafford 2000). in the ordination output, the total variation in the data set is the sum of all the eigenvalues of all axes. the proportion of this total variation that is explained by the supplied explanatory variables is described as the variation explained and is the sum of all canonical eigenvalues divided by the total variation (ter braak 1995). in the ordination diagram, each arrow points in the direction of the steepest increase in the explanatory variable with its length proportional to its importance in explaining the variation, while the angle between the arrows indicates the correlation between individual variables (ter braak & šmilauer 1998). results overall, 150 vegetation plots from 30 palustrine wetlands in lesotho were analysed in this study (figure 1). a total of 312 plant species belonging to 51 families occurred in the wetlands (online appendix 1 – table 1-a1[a] and [b]). of the 312 species, 276 were encountered in the 150 plots and the remaining 36 species occurred within the wetlands but outside the plots (online appendix 1 – table 1-a1[a]; online appendix 2 – table 1-a2). five most dominant families, accounting for most of the species occurring in the wetlands, were poaceae (20.19%), asteraceae (19.23%), cyperaceae (14.10%), scrophulariaceae (4.17%) and polygonaceae (3.85%) (online appendix 1 – table 1-a1 [b]). from these results, it can be observed that three (poaceae, asteraceae and cyperaceae) out of the 51 families account for 53.52% of all the species occurring in these wetlands. while the species richness of whole wetland units ranged between 10 and 53, the number of species per 3 × 3 m plot ranged from 1 (monospecific communities, e.g. community 19) to 27 (highly diverse communities, e.g. communities 3 and 4), with a median species richness of about 14. the highest median number of species for whole wetland units (41.5) was recorded in the highlands and the lowest (26) in the lowlands. the five most frequently occurring species in the wetlands all represent different families (ranunculaceae, leguminosae, asteraceae, poaceae and cyperaceae) (online appendix 1 – table 1-a1[a] and [b]). the type of metabolism (c3/c4) is also indicated in online appendix 1 – table 1-a1[a] for some of the species. the species that were collected during fieldwork were deposited in the roml at the national university of lesotho. figure 1: distribution of the vegetation plots in the palustrine wetlands of lesotho. classification clustering of all the vegetation plots produced 22 distinct communities (figure 2). the 22 communities are described in terms of their dominant species, indicator species and the associated ivs (table 2). these communities were also grouped into five main clusters (wetland types), four highland wetland types (wetland types 1–4) and one lowland type (wetland type 5). while most of the high-altitude communities exhibited the dominance of c3 plants, most of the dominants in the low-altitude communities were c4 species. exceptions are communities such as schoenoplectus paludicola, potamogeton thunbergii and typha domingensis–phragmites australis, which, although dominated by c3 plants, occurred at low altitudes. however, a mixture of c3 and c4 plants dominated the communities with a wide ecological amplitude (see online appendix 1 – table 1-a1[a] for metabolic pathways of some of the species). the synoptic table for the classification of the wetland vegetation is provided in online appendix 2 – table 1-a2. figure 2: dendrogram showing plant communities of the palustrine wetlands of lesotho. numbers in brackets are the numbers of plots in that community. table 2: indicator species of the palustrine wetland communities of lesotho. only species with indicator values of more than 20 and p-values less than 0.05 are presented. all p-values less than 0.001 were given as < 0.001. ordination the cca ordination diagram for all vegetation plots is presented in figure 3. in this ordination, the total variation is 25.483 and the environmental variables supplied account for 18.11% of this. the first axis of the ordination is positively correlated with altitude, longitude, soil parent material and peat but negatively associated with landscape, inundation and aspect. communities located on the right side of the ordination diagram (i.e. communities 1, 3, 5, 7, 10 and 20) are associated with near-pristine and high-altitude areas that are underlain by basalt and located in the eastern part of the country. communities on the left side of the diagram (i.e. communities 6, 15, 16, 17, 18, 21 and 22) are associated with more inundated wetlands in the western lowland and more urbanised areas of the country that are underlain by sandstone. these communities are also associated with a shallower water column. the second axis of the ordination is best explained by slope but wetness and soil depth are also important factors. however, some communities occur on a wide range of altitudinal gradient and these include communities 12, 13 and 14. figure 3: canonical correspondence analysis ordination diagram showing plant communities in the palustrine wetlands of lesotho, with all plots and using the same classification presented in figure 2. thirty of the 150 vegetation plots had detailed soil data and this subset of the vegetation plots represents 15 of the 22 plant communities presented in figure 2. figure 4 presents the cca ordination diagram for this subset of the vegetation plots. the total variation was 10.051 and 65.40% of this could be explained by the supplied explanatory variables. the first axis is positively correlated with potassium and percentage clay but is negatively correlated with altitude, longitude, total organic carbon, nitrogen, sulphur, percentage sand and sodium. while potassium and percentage clay have a strong positive correlation, total organic carbon, nitrogen, sulphur, percentage sand and longitude also have a strong positive correlation. the second axis is negatively correlated with latitude and positively associated with electrical conductivity, soil depth, magnesium and calcium content. communities on the left side of the ordination diagram (i.e. communities 1, 3, 4, 8, 19 and 20) are in high-altitude areas and are associated with sand soils with high organic carbon, nitrogen, sulphur and sodium content and those on the right side (i.e. communities 9, 13, 17, 18, 21 and 22) are in the lowlands and on soils with high potassium and clay levels. those communities on the upper part of the ordination diagram, including communities 8, 9, 11, 12, 17, 19 and 20, are associated with high levels of electrical conductivity, calcium and magnesium, as well as deeper soils. figure 4: canonical correspondence analysis ordination diagram showing plant communities in the palustrine wetlands of lesotho, with only plots with soil data and using the same classification presented in figure 2. description and distribution of wetland plant communities the 22 plant communities were further described in terms of their most dominant species, structure, diversity and environmental conditions (table 3). species that are protected by law in the country are also highlighted in table 3. online appendix 3 – figure 5 presents some of these communities and online appendix 4 provides a further description of all the communities. the distribution range of the wetland vegetation in lesotho is presented in figure 1. table 3: description and environmental conditions of plant communities in the palustrine wetlands of lesotho. table 3 (continues…): description and environmental conditions of plant communities in the palustrine wetlands of lesotho. table 3 (continues…): description and environmental conditions of plant communities in the palustrine wetlands of lesotho. discussion the earliest studies on the palustrine wetlands of lesotho were carried out in the early 1960s and 1970s (guillarmod 1962; van zinderen bakker & werger 1974). since then, other studies conducted in the country (du preez & brown 2011; grab & deschamps 2004; meakins & duckett 1993) were mainly carried out on small areas, focusing on specific or a few wetlands. the current study has provided the most recent and more comprehensive assessment of the palustrine wetlands in the country by providing a classification and description of the wetland vegetation. the wetland vegetation has been classified into 22 communities, which are influenced mainly by altitude, longitude, slope, soil parent material, landscape, inundation, peat, soil potassium, clay, total organic carbon, nitrogen, sulphur, electrical conductivity, calcium, soil depth, wetness, magnesium, aspect and latitude. the wetlands were found to be occurring mainly in those areas where topography allows wetland conditions to develop (mostly in flat areas); mainly on the summit plateau and in the lowlands. nonetheless, it is noteworthy that there may be more communities in the country than the number reported here because only 150 plots from 30 wetlands were analysed in this study, although this forms an important starting point for documenting and classifying the wetland vegetation for lesotho. despite lesotho being entirely afromontane, the study found a clear distinction between wetlands that are found in the highlands and those in the lowlands, in terms of diversity, species making up the communities and structure of the communities (table 3). while the highland wetlands (types 1–4) are mainly south-facing, those in the lowlands (type 5) are mainly north-facing. differences exist between the northand south-facing slopes because of distinct solar radiation patterns. the resulting ecological differences that include snow cover duration and moisture conditions between the moister and colder south-facing slopes and the drier and warmer north-facing slopes contribute to the observed differences in vegetation types (nüsser 2002). indicators of wetland type 5 are typical lowland wetland plants. this wetland type is comparable to the most widespread and common type of wetlands in south africa, the temperate grassy wetlands (sieben et al. 2014). one of the highland wetland types (type 4) comprises wetlands that are found in the small high-altitude area underlain by sandstone and limited to the eastern edge of lesotho (sehlabathebe national park). these wetlands are also located on very steep slopes. however, this wetland type was under-sampled in this study as it was represented by only two wetlands and nine vegetation plots. despite the country’s high altitude (1388–3482 m asl) and rugged terrain, qualifying it to be entirely afromontane (carbutt & edwards 2015), some of the communities in the lowlands also fit into the temperate grassy wetland vegetation of sieben et al. (2017a) and the eastern temperate freshwater wetlands of mucina and rutherford (2006). all the wetlands surveyed in the current study can broadly be classified as the freshwater wetland vegetation type of mucina and rutherford (2006), which is further divided into eastern temperate freshwater wetlands (azf 3), drakensberg wetlands (azf 4) and lesotho mires (azf 5). the dominance of poaceae, asteraceae, cyperaceae and scrophulariaceae families observed in the current study has also been reported in the entire maloti-drakensberg region and the surrounding areas, although asteraceae was the most dominant in the latter (chatanga et al. 2019). the current study also concurs with sieben, glen and muasya (2017b) who report that poaceae is the most common plant family in the south african wetlands based on species richness. however, of the five most dominant plant families recorded in the current study, two (asteraceae and scrophulariaceae) have higher than average levels of endemism in the maloti-drakensberg region (cowling & hilton-taylor 1994). high levels of endemism in the high-altitude environments such as the maloti-drakensberg are attributed to the many small-scale microhabitats and a wide range of edaphic conditions that develop as a result of slope and aspect (brand, scott-shaw & o’connor 2019). the five most common species in the study all represent different families. this implies that the wetland vegetation in the country is phylogenetically diverse and this is unlike the situation in south africa where the five most common wetland plant species are all grasses (poaceae) (sieben et al. 2014). the high altitude that characterises the greater part of lesotho could account for this high diversity, and montane wetlands have been identified as some of the most species-rich in south africa (sieben et al. 2014). while the high-altitude montane wetland communities in lesotho are more diverse and are virtually never a monoculture, the lowland wetland communities are generally less diverse and sometimes monospecific communities. in lowland wetlands, it is common for just one or two species to dominate the entire wetland plant community (boutin & keddy 1993; sieben et al. 2010a). this trend, also exhibited in the current study, is contrary to the decline in species richness with altitude, which has widely been recognised as a general law of ecology (rosenzweig 1995). however, the large number of species and communities recorded in this study generally reflects the high diversity of wetland habitats in the country, although the highest diversity is associated with higher altitudes. it is noteworthy that exceptions such as kniphofia caulescens and carex cognata communities usually exhibit lower diversity than other communities at such higher elevations. this suggests that the dominant species in these communities are so adapted to the harsh high-altitude conditions that they have developed a very large competitive effect on local resources and act as a habitat filter by excluding the less competitive species (maire et al. 2012). it is also noteworthy that the lowlands is the area subjected to more anthropogenic pressures that include cultivation, urbanisation and conversion to other forms of land use and this threatens the wetlands in this part of the country. the higher species richness and diversity in the highlands highlights that these wetland habitats are more diverse than those in the lowlands. this could be attributable, in part, to the fact that most of the dominant species in the high-altitude montane wetlands of the country are non-clonal (table 3). clonal plants are species that can reproduce vegetatively by sprouting from stolons or rhizomes, forming stands of individuals of that species (song & dong 2002). typical clonal plants include phragmites and typha spp. that form dense rhizome mats and outcompete other non-clonal species. these findings corroborate sieben et al. (2010b, 2017b) who suggest that, unlike lowland wetlands, high-altitude wetlands are unusual in that they are richer in species and particularly non-clonal species. moreover, because the usually dominant wetland plants cannot cope well with the low temperatures characterising high-altitude environments, they cannot be as dominant as usual, leaving many vacant niches that then become available for colonisation by other plants (sieben et al. 2010b). furthermore, in wetland environments, the abundance of clonal plants has been reported to be negatively associated with the overall plant species diversity, as well as with altitude (song & dong 2002). the high diversity can also be attributed to the steep gradients in the landscape and harsh climatic conditions that create unique habitats in the mountains of lesotho (pooley 2003). the high diversity can further be ascribed to the underlying basaltic parent material. mucina and rutherford (2006) observe that the seepage water in these high-altitude wetlands is eutrophic as the underlying basalt is rich in nutrients. this also influences the floristic composition and community structure of the wetlands. these results are consistent with observations by sieben et al. (2010a) who acknowledge the significant number of wetland community types in the montane areas of lesotho. the high floristic diversity observed in the current study is also consistent with findings from the high-altitude montane wetlands of alborz mountains, iran (kamrani et al. 2011; naqinezhad et al. 2009). the ordination of the vegetation data for all plots reveals that the explanatory environmental variables supplied could explain only about 18.11% of the total variation. this highlights that the remaining variation could be explained by the environmental factors which were not included in the analysis, such as precipitation (assessed only indirectly through longitude) and wetland water quality. it may also be that plants colonise wetland habitats by chance (chesson 2000). however, the degree of variation explained in the ordination is comparable to similar studies in the wetlands of south africa (sieben et al. 2016, 2017a). moreover, brand et al. (2013) highlight that substrate and hydrogeological conditions play a bigger role in influencing the floristic composition, structure and dynamics in high-altitude montane wetlands than microclimate. the inclusion of soil variables in the analysis increased the proportion of the total variation explained by the supplied variables from 18.11% to 65.40%. thus, the inclusion of soil data in the analysis significantly improves wetland vegetation–environment assessments. the degree of variation explained in the ordination with soil data is significantly higher than that reported for the south african wetlands (sieben et al. 2016, 2017a). nevertheless, the amount of variation explained after the inclusion of soil data could also have improved because the data set was smaller (30 vegetation plots). while the second axis is positively associated with slope, it is negatively correlated with the wetness and soil depth. this implies that communities on the lower part of the ordination diagram are associated with wetter habitats with deeper soils, while those on the upper part are associated with steeper slopes. the former include communities such as c. cognata–juncus effusus, while the latter include communities such as merxmuellera macowanii, which are often associated with hillslope seepages. the correlation between wetness and soil depth, which are both negatively correlated with slope, could be attributed to the soil deposition and water accumulation that is often consistent with fairly flat habitats. altitude, longitude, slope, wetness, soil parent material, landscape, peat, inundation, aspect, soil depth and wetness have been observed to be very important factors explaining the distribution and variation of the wetland vegetation in this study. abundance of peat is also strongly correlated with altitude and longitude. because most of these afromontane wetlands are often located on a slope at high altitudes, they are unique (mucina & rutherford 2006; sieben et al. 2014). a temperature drop of 1 °c has been estimated for every 125 m gain in altitude in lesotho and the maloti-drakensberg region (pomela et al. 2000). such steep environmental gradients over short distances (körner, paulsen & spehn 2011) in afromontane areas are associated with huge spatial variation in physical features and this results in remarkable variation in terms of species diversity and distribution (kotze & o’connor 2000). as a result, increasing altitude corresponds with a decrease in temperature and an increase in rainfall (mucina & rutherford 2006) and the greater part of lesotho is generally much higher and colder than the surrounding areas (sieben et al. 2014). the hypoxia or anoxia in the wetlands, coupled with the low temperature and ph, often associated with these high-altitude wetlands, reduces the rate of decomposition and favours the accumulation of organic matter and peat formation (chatterjee et al. 2010; gopal 2016). therefore, lower temperatures, higher rainfall and other environmental conditions associated with high altitudes also create habitats that can harbour unique vegetation (sieben et al. 2014). through its influence on temperature and rainfall, altitude is a suitable surrogate measure for climate in lesotho and the maloti-drakensberg region, which represents an indirect gradient (sieben et al. 2010a). the strong correlation between longitude and altitude can therefore be explained by the fact that altitude generally increases on moving from west to east in lesotho and rainfall also increases with altitude and with increasing longitude (cowling & hilton-taylor 1994; mucina & rutherford 2006). the high altitudes of lesotho are mainly associated with abundant orographic rainfall, which results in many springs and seepage zones (mucina & rutherford 2006; sieben et al. 2014). the influence of altitude and wetness on high-altitude wetland vegetation has also been reported in south africa (kotze & o’connor 2000; mucina & rutherford 2006; sieben et al. 2010a, 2010b), southern brazil (rolon & maltchik 2006) and in cumbria, uk (jones, li & maberly 2003). the importance of both altitude and slope gradients on the floristic composition of high-altitude montane wetlands has also been reported in bulgaria, south-eastern europe (hájková, hájek & apostolova 2006) and in the alborz mountains, india (kamrani et al. 2011; naqinezhad et al. 2009). although a negative correlation between altitude and wetness was observed in this study, altitude operates at a larger scale, whereas wetness operates on a local scale. it is nevertheless noteworthy that, while some communities are restricted to either the highlands or lowlands, others seem to have a wide ecological amplitude. these include eragrostis plana–pennisetum sphacelatum, eleocharis dregeana and eragrostis planiculmis communities. furthermore, some species, such as ranunculus meyeri harv., have been reported to occur at low abundances at lower altitudes but achieve greater cover at higher altitudes as they gain a competitive advantage because of the lower temperatures that reduce the vigour of the usually more competitive species (sieben et al. 2010b). because some of the plant communities recorded in this study are restricted to the highest altitudes, occurring at the summit plateaus, they are likely to disappear in the face of climate change as they cannot migrate any further up (bentley, robertson & barker 2019; lee et al. 2015). while high-altitude communities (e.g. limosella grandiflora–haplocarpha nervosa, agrostis bergiana and cotula paludosa– r. meyeri communities) were also associated with soils that are high in total organic carbon, nitrogen and sulphur, the habitat soils for the low-altitude communities (e.g. cynodon incompletus, eleocharis limosa, s. paludicola and e. dregeana communities) were high in potassium and clay percentage. however, the negative correlation of potassium with altitude is contrary to mucina and rutherford (2006) who report that the high-altitude areas are associated with high potassium levels. in the current study, communities such as c. cognata, gunnera perpensa and e. plana–p. sphacelatum were also found to be associated with high soil magnesium, calcium and electrical conductivity. the importance of potassium, electrical conductivity and clay percentage in influencing high-altitude montane wetland vegetation is consistent with findings from a similar study on the wetlands of the alborz mountains, india (kamrani et al. 2011). because of the low temperature and basaltic parent material, the soils in the mountains of lesotho are high in organic matter and generally nutrient-rich, with a high water-retention capacity (mucina & rutherford 2006). however, the soils found in the lowlands are considered to be generally nutrient-poor (maro 2011). conservation value and impacts of climate change some of the species encountered in this study are protected by law in lesotho (table 3), which highlights the fact that they are threatened. the wetlands of lesotho contribute significantly not only to biodiversity, but also provide a wide spectrum of ecosystem services, particularly in terms of water resources and livestock grazing. they are the headwaters of the five major, economically important rivers in the country, namely the maliba-matšo, senqu-orange, mohokare (caledon), makhaleng and senqunyane, which also feed the lhwp dams (department of environment 2014). they play a major role in sustaining the perennial flow of water and regulating the water quality of the rivers that flow into the atlantic ocean (pooley 2003). however, all the mountain areas in the country are drained through the senqu-orange river and its tributaries (backéus & grab 1995), the most important and most developed shared river system in southern africa. for south africa, the water is tapped mainly through the lhwp. in fact, the lesotho highlands water is of national importance to the south african population as a significant portion of the agriculture in the arid regions of the country is watered from the aqueducts from the dams on the senqu-orange river. because of their high carbon sequestration and storage capacity, the conservation of these wetlands can also play a role in mitigating global climate change. peatlands are the second most important reservoir of carbon on earth after oceans (russi et al. 2013). climate change predictions highlight that much of southern africa will become drier (mitchell 2013). based on climate change modelling, by 2025, namibia will probably experience problems of water quality and availability, lesotho will be water-stressed and south africa will be facing absolute water scarcity (sadc 2008). accordingly, the conservation of these wetlands becomes indispensable. the wetlands of lesotho are a critical resource for livestock grazing, especially in summer when thousands of cattle, sheep and goats are seen grazing on these sensitive ecosystems (du preez & brown 2011; van zinderen bakker & werger 1974). in fact, much of the grazing in the mountains of lesotho takes place within wetlands because they harbour the most palatable vegetation (grab & deschamps 2004). the basotho are now sometimes observed keeping their livestock in the mountains throughout the year. this is contrary to their tradition of mostly inhabiting the lowlands and taking their herds of livestock to the mountains in summer and returning them to the lowlands during winter (meakins & duckett 1993). thus, the value of these wetlands as a grazing resource is increasingly becoming greater. the widespread degradation and loss of wetlands, mainly because of grazing and trampling by cattle, sheep and goats, has been reported quite extensively since the 1960s (e.g. backéus & grab 1995; du preez & brown 2011; guillarmod 1962; van zinderen bakker & werger 1974). most of the communities described here are threatened by grazing and trampling except for the m. macowanii community, which is unpalatable. it is painfully unpleasant to walk through the community because of the sharp tips of the dominant m. macowanii (stapf) conert. at least eight invasive alien species have been recorded in the wetlands and these include paspalum dilatatum poir., cirsium vulgare (savi) ten., cyperus esculentus l., hypochaeris radicata l. and p. clandestinum hochst. ex chiov. while some of them occurred more frequently (e.g. p. dilatatum occurred in 16% of the plots and was the 11th most frequent species in the study), others occurred in relatively low frequencies (online appendix 1 – table 1-a1[a]) but in large stands in a given wetland (e.g. c. vulgare). paspalum dilatatum was found to be the fourth most dominant wetland species in south africa, occurring in 9.1% of all the wetland vegetation plots in the country’s database (sieben et al. 2014). invasive alien plants are not only among the major drivers of wetland degradation and loss in sub-saharan africa (mitchell 2013), but also use wetlands as corridors to penetrate even terrestrial plant communities (mucina & rutherford 2006). the wetlands are further threatened mainly by the economic development associated with damming, mining and infrastructure development. therefore, the unsustainable utilisation of the water and other resources, as well as other anthropogenic activities, in the different catchments of the country, would have an adverse effect on the wetlands and concomitantly diminish their capacity to supply the water resources. nevertheless, montane wetlands generally remain less impacted than those in the lowlands where human pressure is higher (lee et al. 2015), although wetlands in general are among the most threatened ecosystems globally (millennium ecosystem assessment 2005; russi et al. 2013). consequently, montane wetlands also serve as repositories and refugia for biodiversity and are regarded as biodiversity hot spots (chatterjee et al. 2010). these wetlands can also potentially be threatened by climate change, which has been predicted to significantly alter the availability of water for individual montane wetlands and give rise to substantial shifts in the distribution and composition of wetlands in montane landscapes (lee et al. 2015). widespread ecological transitions have been reported to be likely at higher elevations because montane regions exhibit greater sensitivity to climate change (bentley et al. 2019; ryan et al. 2014). this will concomitantly give rise to widespread changes in the functioning of these wetlands and their delivery of ecosystem services. projections have also indicated that climate change will cause potentially severe threats on peat-forming wetlands (essl et al. 2012; lee et al. 2015). the maloti-drakensberg area has been reported to be highly vulnerable to climate change (brand et al. 2019). in the maloti-drakensberg, environmental domains that are defined by climate variables are projected to change substantially as a result of climate change, particularly an increase in temperature and a concomitant decline in water availability (brand et al. 2019). most of the species in the maloti-drakensberg are already at the highest altitudes and cannot shift any higher (bentley et al. 2019). furthermore, the pattern of increasing dominance of c3 plants in communities with increasing altitude, which is evident in this study and consistent with the findings of kotze and o’connor (2000) in kwazulu-natal, south africa, highlights a threat to the high-altitude communities because c3 will be replaced with c4 plants as the climate changes. conclusion given their important role in water resources, livestock grazing and harbouring rare and endemic species, as well as unique biodiversity, the wetlands described in the current study are of high conservation value in southern africa, particularly in the face of increased water scarcity, climate change and unprecedented loss of biodiversity. accordingly, the wetlands of lesotho play a critical role in providing ecosystem services at the local, national and regional scales. as a result, the government of lesotho and the orange-senqu river commission (orasecom) have a responsibility in terms of scaling-up the efforts towards the protection of these wetlands for the sustainable provision of the benefits obtained from them. because the wetland vegetation of a particular wetland can be used as a proxy for biodiversity of the wetland, understanding the wetland vegetation–environment patterns is important for the successful conservation planning of these ecosystems. therefore, the current study has provided baseline information, which can be useful for monitoring the wetland vegetation and concomitantly the wetlands, which are vital for the water resources of lesotho, south africa and namibia. acknowledgements the authors would like to thank khotso kobisi and moretloa j. polaki who helped in identifying some of the species. the authors also thank tumelo tjale who helped in collecting part of the data. competing interests the authors declare that they have no competing interests that may have inappropriately influenced them in writing this article. authors’ contributions p.c. led the conceptualisation of the study and conducted fieldwork, laboratory analysis and data analysis and wrote the manuscript. e.j.j.s. helped in conceptualising the study, conducting fieldwork and manuscript writing. ethical considerations this article followed all ethical standards for research without direct contact with human or animal subjects. funding information the study was funded by the university of kwazulu-natal and the national university of lesotho. data availability statement permission to conduct the study was obtained from the lesotho department of environment. disclaimer the views expressed in this article are the authors’ own and not an official position of their institutions or funders. references backéus, i. & grab, s., 1995, ‘mires in lesotho’, gunneria 70, 243–250. bentley, l.k., robertson, m.p. & barker, n.p., 2019, ‘range contraction to a higher elevation: the likely future of the montane vegetation in south africa and lesotho’, biodiversity and conservation 28(1), 131–153. https://doi.org/10.1007/s10531-018-1643-6 boutin, c. & keddy, p.a., 1993, ‘a functional classification of wetland plants’, journal of vegetation science 4(5), 591–600. https://doi.org/10.2307/3236124 brand, r.f., du preez, p.j. & brown, l.r., 2013, ‘high altitude montane wetland vegetation classification of the eastern free state, south africa’, south african journal of botany 88(september), 223–236. https://doi.org/10.1016/j.sajb.2013.07.011 brand, r.f., scott-shaw, c.r. & o’connor, t.g., 2019, ‘the alpine flora on inselberg summits in the maloti–drakensberg park, kwazulu-natal, south africa’, bothalia–african biodiversity & conservation 49(1), 1–15. https://doi.org/10.4102/abc.v49i1.2386 brown, l.r., du preez, p.j., bezuidenhout, h., bredenkamp, g.j., mostert, t.h.c. & collins, n.b., 2013, ‘guidelines for phytosociological classifications and descriptions of vegetation in southern africa’, koedoe 55(1), art. #1103, 1–10. https://doi:10.4102/koedoe.v55i1.1103 carbutt, c. & edwards, t.j., 2015, ‘reconciling ecological and phytogeographical spatial boundaries to clarify the limits of the montane and alpine regions of sub-sahelian africa’, south african journal of botany 98(may), 64–75. https://doi.org/10.1016/j.sajb.2015.01.014 chatanga, p., kotze, d.c., janks, m. & sieben, e.j.j., 2019, ‘classification, description and environmental factors of montane wetland vegetation of the maloti-drakensberg region and the surrounding areas’, south african journal of botany 125(september), 221–233. https://doi.org/10.1016/j.sajb.2019.04.028 chatterjee, a., blom, e., gujja, b., jacimovic, r., beevers, l., o’keeffe, j. et al., 2010, ‘wwf initiatives to study the impact of climate change on himalayan high-altitude wetlands (haws)’, mountain research and development 30(1), 42–52. https://doi.org/10.1659/mrd-journal-d-09-00091.1 chesson, p., 2000, ‘general theory of competitive coexistence in spatially-varying environments’, theoretical population biology 58(3), 211–237. https://doi.org/10.1006/tpbi.2000.1486 clegg, b.w. & o’connor, t.g., 2012, ‘the vegetation of malilangwe wildlife reserve, south-eastern zimbabwe’, african journal of range & forage science 29(3), 109–131. https://doi.org/10.2989/10220119.2012.744352 collins, n.b., 2005, wetlands: the basics and some more, free state department of tourism, environmental and economic affairs, bloemfontein, viewed 07 april 2017, from https://www.dwa.gov.za/iwqs/rhp/provinces/freestate/wetlands_basics&more.pdf. cowling, r.m. & hilton-taylor, c., 1994, ‘patterns of plant diversity and endemism in southern africa: an overview’, in b.j. huntley (ed.), botanical diversity in southern africa, pp. 31–52, botanical research institute, pretoria. cronk, j.k. & fennessy, m.s., 2001, wetland plants: biology and ecology, lewis publishers, boca raton, fl. department of environment, 2014, lesotho environment outlook report 2014: environment for national prosperity, ministry of tourism, environment and culture, maseru, lesotho, 280 p. dufrêne, m. & legendre, p., 1997, ‘species assemblages and indicator species: the need for a flexible asymmetrical approach’, ecological monographs 67(3), 345–366. https://doi.org/10.2307/2963459 du preez, p.j. & brown, l.r., 2011, ‘impact of domestic animals on ecosystem integrity of lesotho high altitude peatlands’, in o. grillo & g. venora (eds.), ecosystems biodiversity, pp. 249–271, intech, rijeka, croatia. essl, f., dullinger, s., moser, d., rabitsch, w. & kleinbauer, i., 2012, ‘vulnerability of mires under climate change: implications for nature conservation and climate change adaptation’, biodiversity and conservation 21(3), 655–669. https://doi.org/10.1007/s10531-011-0206-x gopal, b. (ed.), 2015, guidelines for rapid assessment of biodiversity and ecosystem services of wetlands, version 1.0, asia-pacific network for global change research (apn-gcr), kobe, japan, and national institute of ecology, new delhi, 134 p. gopal, b., 2016, ‘should “wetlands” cover all aquatic ecosystems and do macrophytes make a difference to their ecosystem services?’, folia geobotanica 51(3), 209–226. https://doi.org/10.1007/s12224-016-9248-x grab, s.w. & deschamps, c.l., 2004, ‘geomorphological and geoecological controls and processes following gully development in alpine mires, lesotho’, arctic, antarctic, and alpine research 36(1), 49–58. https://doi.org/10.1657/1523-0430(2004)036[0049:gagcap]2.0.co;2 guillarmod, a.j., 1962, ‘the bogs and sponges of the basutoland mountains’, south african journal of science 58(6), 179–182. hájková, p., hájek, m. & apostolova, i., 2006, ‘diversity of wetland vegetation in the bulgarian high mountains, main gradients and context-dependence of the ph role’, plant ecology 184(1), 111–130. https://doi.org/10.1007/s11258-005-9056-5 jones, j.i., li, w. & maberly, s.c., 2003, ‘area, altitude and aquatic plant diversity’, ecography 26(4), 411–420. https://doi.org/10.1034/j.1600-0587.2003.03554.x junk, w.j., an, s., finlayson, c.m., gopal, b., květ, j., mitchell, s.a. et al., 2013, ‘current state of knowledge regarding the world’s wetlands and their future under global climate change: a synthesis’, aquatic sciences 75(1), 151–167. https://doi.org/10.1007/s00027-012-0278-z kamrani, a., jalili, a., naqinezhad, a., attar, f., maassoumi, a.a. & shaw, s.c., 2011, ‘relationships between environmental variables and vegetation across mountain wetland sites, n. iran’, biologia 66(1), 76–87. https://doi.org/10.2478/s11756-010-0127-2 körner, c., paulsen, j. & spehn, e.m., 2011, ‘a definition of mountains and their bioclimatic belts for global comparisons of biodiversity data’, alpine botany 121(2), 73–78. https://doi.org/10.1007/s00035-011-0094-4 kotze, d.c. & o’connor, t.g., 2000, ‘vegetation variation within and among palustrine wetlands along an altitudinal gradient in kwazulu-natal, south africa’, plant ecology 146(1), 77–96. https://doi.org/10.1023/a:1009812300843 kotze, d.c., klug, j.r., hughes, j.c. & breen, c.m., 1996, ‘improved criteria for classifying hydric soils in south africa’, south african journal of plant and soil 13(3), 67–73. https://doi.org/10.1080/02571862.1996.10634378 lee, s.-y., ryan, m.e., hamlet, a.f., palen, w.j., lawler, j.j. & halabisky, m., 2015, ‘projecting the hydrologic impacts of climate change on montane wetlands’, plos one 10(9), e0136385. https://doi.org/10.1371/journal.pone.0136385 leketa, k., migwi, m., crane, e., upton, k., ó dochartaigh, b. & bellwood-howard, i., 2018, africa groundwater atlas: hydrogeology of lesotho. british geological survey, viewed 06 august 2018, from http://earthwise.bgs.ac.uk/index.php/hydrogeology_of_lesotho. lepš, j. & šmilauer, p., 2003, multivariate analysis of ecological data using canoco, cambridge university press, cambridge. ludwig, j.a. & reynolds, j.f., 1988, statistical ecology: a primer on methods and computing, john wiley and sons, new york. maire, v., gross, n., börger, l., proulx, r., wirth, c., da silveira pontes, l. et al., 2012, ‘habitat filtering and niche differentiation jointly explain species relative abundance within grassland communities along fertility and disturbance gradients’, new phytologist 196(2), 497–509. https://doi.org/10.1111/j.1469-8137.2012.04287.x maro, p., 2011, environmental change in lesotho: an analysis of the causes and consequences of land-use change in the lowland region, springer, london. mccune, b. & mefford, m.j., 2011, pc-ord. multivariate analysis of ecological data. version 6.0, mjm software design, gleneden beach, or. mcgarigal, k., cushman, s. & stafford, s., 2000, multivariate statistics for wildlife and ecology research, springer-verlag inc., new york. meakins, r.h. & duckett, j.d., 1993, ‘vanishing bogs of the mountain kingdom’, veld and flora 79(2), 49–51. millennium ecosystem assessment, 2005, ecosystems and human well-being: wetlands and water synthesis, world resources institute, washington, dc. mitchell, s.a., 2013, ‘the status of wetlands, threats and the predicted effect of global climate change: the situation in sub-saharan africa’, aquatic sciences 75(1), 95–112. https://doi.org/10.1007/s00027-012-0259-2 mitsch, w.j. & gosselink, j.g., 2000, ‘the value of wetlands: importance of scale and landscape setting’, ecological economics 35(1), 25–33. https://doi.org/10.1016/s0921-8009(00)00165-8 mitsch, w.j. & gosselink, j.g., 2015, wetlands, 5th edn., wiley, hoboken, nj. moor, h., rydin, h., hylander, k., nilsson, m.b., lindborg, r. & norberg, j., 2017, ‘towards a trait-based ecology of wetland vegetation’, journal of ecology 105(6), 1623–1635. https://doi.org/10.1111/1365-2745.12734 mucina, l. & rutherford, m.c. (eds.), 2006, the vegetation of south africa, lesotho and swaziland, strelitzia 19, south african national biodiversity institute (sanbi), pretoria. mueller-dombois, d. & ellenberg, h., 1974, aims and methods of vegetation ecology, john wiley and sons, new york. mugomeri, e., chatanga, p., raditladi, t. & tarirai, c., 2016, ‘ethnobotanical study and conservation status of local medicinal plants: towards a repository and monograph of herbal medicines in lesotho’, african journal of traditional, complementary & alternative medicines 13(1), 138–151. https://doi.org/10.4314/ajtcam.v13i1.20 naqinezhad, a., jalili, a., attar, f., ghahreman, a., wheeler, b.d., hodgson, j.g. et al., 2009, ‘floristic characteristics of the wetland sites on dry southern slopes of the alborz mts., n. iran: the role of altitude in floristic composition’, flora – morphology, distribution, functional ecology of plants 204(4), 254–269. https://doi.org/10.1016/j.flora.2008.02.004 nüsser, m., 2002, ‘pastoral utilization and land cover change: a case study from the sanqebethu valley, eastern lesotho’, erdkunde 56(2), 207–221. https://doi.org/10.3112/erdkunde.2002.02.07 nüsser, m. & grab, s., 2002, ‘land degradation and soil erosion in the eastern highlands of lesotho, southern africa’, die erde 133, 291–311. ollis, d., snaddon, k., job, n. & mbona, n., 2013, classification system for wetlands and other aquatic ecosystems in south africa. user manual: inland systems. sanbi biodiversity series 22, south african national biodiversity institute, pretoria. omar, m.y., maroyi, a. & van tol, j.j., 2016, ‘floral diversity, composition and distribution in a montane wetland in hogsback, the eastern cape province, south africa’, pakistan journal of botany 48(5), 1861–1870. orasecom, 2015, orange-senqu river commission 15 years 2000–2015, orange-senqu river commission, gauteng, 60 p. peck, j.e., 2010, step-by-step using pc-ord. multivariate analysis for community ecologists, mjm software design, gleneden beach, or. pla, l., casanoves, f. & di rienzo, j., 2012, functional diversity indices, springer, london. pomela, e.m., mokuku, c., maabataung, s., phororo, r., makoae, m., majoro, m. et al., 2000, biological diversity in lesotho: a country study, national environmental secratariat, government of lesotho, maseru. pooley, e., 2003, mountain flowers: a field guide to the flora of the drakensberg and lesotho, 1st edn., flora publications trust, durban, 320 p.-illus., col. illus. rolon, a.s. & maltchik, l., 2006, ‘environmental factors as predictors of aquatic macrophyte richness and composition in wetlands of southern brazil’, hydrobiologia 556(1), 221–231. https://doi.org/10.1007/s10750-005-1364-1 rosenzweig, m.l., 1995, species diversity in space and time, cambridge university press, cambridge. russi, d., ten brink, p., farmer, a., badura, t., coates, d., förster, j. et al., 2013, the economics of ecosystems and biodiversity for water and wetlands, ieep, london. ryan, m.e., palen, w.j., adams, m.j. & rochefort, r.m., 2014, ‘amphibians in the climate vise: loss and restoration of resilience of montane wetland ecosystems in the western us’, frontiers in ecology and the environment 12(4), 232–240. https://doi.org/10.1890/130145 sadc, 2008, southern africa environment outlook, sadc, sardc, iucn & unep, gaborone. sieben, e.j.j., collins, n.b., kotze, d.c., mofutsanyana, s.s. & janks, m., 2017a, ‘temperate grassy wetlands of south africa: description, classification and explanatory environmental factors’, south african journal of botany 113(november), 68–76. https://doi.org/10.1016/j.sajb.2017.07.009 sieben, e.j.j., glen, r.p. & muasya, a.m., 2017b, ‘the wetland flora of south africa: occurrence patterns, frequency and dominance across a diverse landscape’, aquatic botany 142(september), 112–118. https://doi.org/10.1016/j.aquabot.2017.03.003 sieben, e.j.j., kotze, d.c. & morris, c.d., 2010a, ‘floristic composition of wetlands of the south african section of the maloti-drakensberg transfrontier park’, bothalia 40(1), 117–134. https://doi.org/10.4102/abc.v40i1.201 sieben, e.j.j., morris, c.d., kotze, d.c. & muasya, a.m., 2010b, ‘changes in plant form and function across altitudinal and wetness gradients in the wetlands of the maloti-drakensberg, south africa’, plant ecology 207(1), 107–119. https://doi.org/10.1007/s11258-009-9657-5 sieben, e.j.j., mtshali, h. & janks, m., 2014, national wetland vegetation database: classification and analysis of wetland vegetation types for conservation planning and monitoring, report no. wrc 1980/1/14, water research commission (wrc), pretoria, south africa, 255 p. sieben, e.j.j., nyambeni, t., mtshali, h., corry, f.t.j., venter, c.e., mackenzie, d.r. et al., 2016, ‘the herbaceous vegetation of subtropical freshwater wetlands in south africa: classification, description and explanatory environmental factors’, south african journal of botany 104(may), 158–166. https://doi.org/10.1016/j.sajb.2015.11.005 singha, v., 2011, high altitude wetlands project: a retrospective, wwf, kathmandu, nepal. song, m. & dong, m., 2002, ‘clonal plants and plant species diversity in wetland ecosystems in china’, journal of vegetation science 13(2), 237–244. https://doi.org/10.1111/j.1654-1103.2002.tb02044.x stohlgren, t.j., bull, k.a., otsuki, y., villa, c.a. & lee, m., 1998, ‘riparian zones as havens for exotic plant species in the central grasslands’, plant ecology 138(1), 113–125. https://doi.org/10.1023/a:1009764909413 ter braak, c.j.f., 1995, ‘ordination’, in r.h.g. jongman, c.j.f. ter braak & o.f.r. van tongeren (eds.), data analysis in community and landscape ecology, pp. 91–173, cambridge university press, cambridge. ter braak, c.j.f. & šmilauer, p., 1998, canoco reference manual and user’s guide to canoco for windows: software for canonical community ordination (version 4), microcomputer power, ithaca, ny. van der maarel, e., 1979, ‘transformation of cover-abundance values in phytosociology and its effects on community similarity’, vegetatio 39(2), 97–114. https://doi.org/10.1007/bf00052021 van oudtshoorn, f., 2014, guide to grasses of southern africa, 3rd edn., briza publications, pretoria. van tongeren, o.f.r., 1995, ‘cluster analysis’, in r.h.g. jongman, c.j.f. ter braak & o.f.r. van tongeren (eds.), data analysis in community and landscape ecology, pp. 174–212, cambridge university press, cambridge. van zinderen bakker, e.m. & werger, m.j.a., 1974, ‘environment, vegetation and phytogeography of the high-altitude bogs of lesotho’, vegetatio 29(1), 37–49. https://doi.org/10.1007/bf02390894 introduction the sabie river sabie river geomorphology and sediment dynamics discussion conclusions acknowledgements references about the author(s) jasper knight school of geography, archaeology and environmental studies, university of the witwatersrand, johannesburg, south africa mary evans school of geography, archaeology and environmental studies, university of the witwatersrand, johannesburg, south africa citation knight, j. & evans, m., 2022, ‘characterising the geomorphic dynamics of river systems: an example of the sabie river, south africa’, koedoe 64(1), a1700. https://doi.org/10.4102/koedoe.v64i1.1700 short communication characterising the geomorphic dynamics of river systems: an example of the sabie river, south africa jasper knight, mary evans received: 03 dec. 2021; accepted: 17 feb. 2022; published: 29 mar. 2022 copyright: © 2022. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. introduction semiarid climatic conditions prevail over much of southern africa, with variations in rainfall over seasonal and interannual timescales (saraiva okello et al. 2015). this rainfall pattern provides rise to highly variable river discharge regimes from high-energy floods in the wet season to low perennial flow, or isolated remnant pools or dry river beds, in the dry season (heritage et al. 2015; rountree, rogers & heritage 2000). these discharge regimes have important implications for river properties as a whole, including water quality, maintenance of ecosystems and biodiversity, pollutant dispersal, and changes in river geomorphology (seanego & moyo 2013). the latter has been proposed as a means of mapping and classifying river systems in southern africa, especially in areas with seasonally variable river regimes (e.g. eze & knight 2018; knight in press; rountree et al. 2000; van niekerk, heritage & moon 1995). the major river reach types identified through such mapping are mixed anastomosing, alluvial braided, mixed pool-rapid and alluvial single thread (heritage, van niekerk & moon 1997; moon & heritage 2001) that describe different combinations of bedrock versus loose sediment within and around the river channel(s) (rosgen 1994). the substrate type and its properties and dynamics have implications for turnover of ecosystem biodiversity, persistence of endemic species, spread of invasive species and reducing flood risk (entwistle et al. 2014; pettit et al. 2005; rountree et al. 2000). despite such applications, many river systems in southern africa have not been fully or accurately mapped at this scale, and individual reaches even along the same river can be highly variable in character. for example, along the limpopo river system, the most common river reach category is low/moderate sinuosity with planform-controlled sand beds (37% of total river length), followed by meandering sand beds (26%) and bedrock-forced meanders (17%) (knight in press). however, different tributaries of the limpopo have different geomorphic characters, with bedrock-forced meanders being dominant (84%) on the nzhelele river, meandering sand beds (69%) on the elephantes river and low-sinuosity fine-grained beds (41%) on the lotsane river (knight in press). although river geomorphology can be mapped relatively easy, especially by remote sensing, the dynamics of many semiarid river systems are unknown. here, ‘dynamics’ refers to how the geomorphology of the river system changes over time and space, especially in response to variations in rainfall and thus river discharge. understanding the dynamics of river systems has implications for understanding floodwater and sediment flows and pathways, identifying which areas are likely to experience flood impacts, restoring eroded or polluted river reaches, and producing flood risk maps for human settlements and infrastructure (gericke & du plessis 2012). although this analysis has been undertaken for individual rivers (e.g. heritage et al. 2015; knight & evans 2018; milan et al. 2018) many rivers across southern africa remain unmapped for these purposes. the aim of this study is to use geomorphology, sediments and dating evidence from the sabie river in northeast south africa in order to characterise its dynamics. this is located in a semiarid environment but subject to significant seasonal variations in river discharge, giving rise to repeated historical flood events. this article (1) describes the evidence for river system dynamics based on its (a) geomorphology, (b) sediment patterns and (c) dating evidence for its flood history; (2) proposes a model for the nature of flood events along different river reaches and (3) discusses how this information can be applied to river management. the sabie river the sabie river catchment (7096 km2) has its source areas in the eastern escarpment of mpumalanga province, south africa, and drains eastwards into mozambique and the indian ocean, traversing the lowveld and lebombo topographical zones (400 m and 200 m above see level (a.s.l.), respectively) (figure 1a). bedrock geology ranges from precambrian igneous and metasedimentary rocks in headwater areas of the eastern escarpment to rhyolite, basalt and sandstones (jurassic) found in middle and lower reaches of the lowveld within kruger national park (knp) (schutte 1986). the variable topography results in a precipitation gradient from wetter conditions (~2000 mm yr−1) in westward mountains to dryer conditions (~600 mm yr−1) in eastward lowlands. most rainfall takes place during the austral summer. sabie river discharge is therefore highly seasonal with a winter average low flow of approximately 2 m3 s−1 – 3 m3 s−1 and a summer average high flow of approximately 20 m3 s−1 – 50 m3 s−1 (data for the period 1995–2014 from station x3h015, figure 1a). peak seasonal flood events commonly reach discharges of 300 m3 s−1 – 400 m3 s−1 in its middle and lower reaches. figure 1: (a) location of the sabie river in northeast south africa, showing the locations of sites named in the text (red dots), river gauging station x3h015 and the three river geomorphic zones (marked after eze & knight 2018) of (i) highland zone of gorges and bedrock-forced meanders; (ii) low to moderately sinuous planform controlled mixed beds; (iii) low sinuosity, fine grained beds and floodouts. zones are separated by green lines on the map. (b) photo of the sabie river within zone i at bergvliet (site 1 on panel a); (c) photo within zone ii at kruger gate (site 2 on panel a); (d) photo within zone iii near the sabie-sand confluence (site 3 on panel a). sabie river geomorphology and sediment dynamics several lines of evidence exist for geomorphic change and sediment dynamics along the sabie river and these are examined in turn. integration of such evidence allows for the tempo or pacing of river geomorphology and sediment dynamics to be identified and this is a key step in developing a river-specific flood management model. mapping geomorphic patterns and geomorphic change several stretches of the sabie have been mapped from aerial photographs or satellite images, and based on this, spatial patterns of bedrock versus sediment-dominated reaches have been described (e.g. entwistle et al. 2014; eze & knight 2018; heritage & moon 2000) although in reality most reaches are of mixed type. three geomorphological zones are identified (marked i–iii in figure 1a). zone i: headwater areas of the eastern escarpment sector are bedrock dominated, with pool and riffle systems present, limited loose sand available but often stacked fluvially transported boulders adjacent to the single channel and a restricted or narrow floodplain (figure 1b). zone ii: a transitional zone exists in mid-catchment locations such as east of hazyview where bedrock elements are separated by more sediment-dominant river stretches. here, the single river channel is located within a narrow and incised bedrock-defined valley but valley side sediment accumulations are often present as either unpaired river terraces or as welded lateral bars with overbank deposits (figure 1c). zone iii: in lowland reaches of the sabie river, the main river basin is sediment dominated throughout and the main river channel, which may be braided or multichannel, extends across a wide floodplain (several 100 m) with shallow bedrock slopes on either side. for most of the year, the main channel is underfit but episodic floods cause reactivation of abandoned channels, enhanced mobility of in-channel bars and phases of overbank deposition that commonly extends across the width of the floodplain. rapid geomorphic change takes place in response to seasonal flood events. such recent events coincide with indian ocean cyclones dando (january 2012), dineo (february 2017) and idai (march 2019). high river velocity, basal shear stress and discharge during floods result in differential sediment erosion and deposition, leading to changes in the distributions of sand and exposed bedrock and changes in land surface elevation. substrate types and elevation changes have been mapped in studies that compare preand post-flood river conditions. for example, using light detection and ranging (lidar) data, milan et al. (2018) calculated 66 880 m3/km−1 of erosion and 24 380 m3/km−1 of deposition during cyclone dando in the middle/lower sabie. broadhurst and heritage (1998) showed that river channel responses to floods depend on antecedent sediment distributions, single or multichannels and floodplain width. this results in the development of distinctive landforms in different places along the river system (heritage & moon 2000). bedrock and sediment substrates also show different responses to variations in river discharge. sediment is easily able to erode and accrete, developing and destroying landforms and changing land surface elevation; bedrock only undergoes erosion and at much slower rates. a sediment cover can protect underlying bedrock from erosion but bedrock erosion rates may rapidly increase if the sediment cover is stripped away. these feedbacks highlight the variable rates and styles of geomorphic change in different reaches in response to floods (heritage et al. 2015). sediment dynamics and mobility although changes in the areal sediment thickness can be evaluated using lidar data, this cannot discern changes in sediment type, properties or depositional structures. on the sabie, field observations can inform on such patterns. heritage and moon (2000) showed some examples of different river landform types, including bars and boulder deposits. knight and evans (2017) described the characteristics of flood deposits in the lower reaches of the sabie and a schematic ‘flood stratigraphy’ model comprising (from base to top): 2d and 3d sandy subaqueous dunes (migrating in-channel bedforms) overlying an erosional surface, overlain in turn by climbing rippled sand (saltation transport at the base of the flow), transitional to massive sand deposited by suspension under waning flow conditions, overlain by laminated silts, deposited under stagnant water conditions, topped by fragmented woody debris indicative of high-energy erosion during the flood and deposited when any remaining water in the floodplain or channel is lost. several studies have also shown that different parts of the river channel – and therefore different types of bars – became active at different flow stages and different seasons, depending on the flow regime (heritage, broadhurst & birkhead 2001; heritage et al. 2015). this means that higher elevation former channels will only become active under higher flow conditions and will be abandoned first when the water level falls. they will also act as sites for sediment deposition during waning flow stages (cunningham, evans & knight 2015). patterns of overbank and floodplain deposition by flood events on the sabie have also been examined (knight & evans 2017, 2018; milan et al. 2018), showing that there are different sedimentary expressions in different parts of the occupied floodplain depending on water depth, water velocity (thus shear stress) and local topography of the bedrock or sediment substrate. there is as yet no clear understanding of spatial patterns of sediment properties (grain size, sorting) along the sabie or how in-channel bedforms change during flood stages, owing to a lack of field data. dating periods of sediment deposition organic-poor sandy sediments can be dated using luminescence methods, and this has been carried out extensively throughout southern african rivers, including on the sabie (e.g. colarossi et al. 2015; cunningham et al. 2015; heritage et al. 2015; knight & evans 2018). the basis behind this method is that the calculated luminescence age of a quartz sand grain is related to the time period as the grain was last exposed to sunlight (murray & wintle 2003). in a river system, sand grains are bleached (luminescence signal reset to zero) by exposure to sunlight during active erosion and transport, and grains become locked away from the surface when they are deposited and buried. calculating the luminescence age of a sample can, therefore, inform on the timing of sediment erosion and deposition events, which, along the sabie and similar rivers, take place during floods. on the sabie, sediment samples around or in the present channel at 10 cm – 45 cm depth have yielded ages from 26 ± 7 to 131 ± 19 years (knight & evans 2018). samples from 1.49 m–3.50 m depth have yielded ages from 188 ± 19 to 910 ± 278 years (heritage et al. 2015). however, ages with a wide error margin may reflect incomplete bleaching of the luminescence signal within sediment grains during their transport (colarossi et al. 2015; cunningham et al. 2015). this means that individual grains can contain inherited luminescence signals from previous transport and deposition events. several innovative studies have examined the nature of this inherited signal from the sabie river. cunningham et al. (2015) showed that of flood samples recovered from higher elevations along the sabie, only 20% – 70% of grains are well bleached compared with > 80% of grains from low-flow positions. based on luminescence signals of samples from different sites across the floodplain, knight and evans (2018) showed that sediments suspended in floodwaters at the centre of the channel are well mixed, and thus, when deposited, have a consistent luminescence signal throughout. by contrast, suspended sediment settling from slack water across the floodplain takes place more slowly, leading to increased bleaching and thus younger ages over time. this depositional model can explain relationships between the degree of sediment reworking (and luminescence signal bleaching) during floods and resulting river geomorphic and sedimentary signatures. discussion understanding the dynamics of river systems through considering their geomorphology, sediment patterns and dated periods of activity have implications for identifying the timescale of cyclic behaviour that typifies any river reach. with respect to the sabie river, although significant cyclonic rainfall takes place in summer annually, this does not always give rise to a proportional flood response. this means that sediment transport and geomorphic change does not always take place outside of the main river channel, such that higher areas of the floodplain may not be affected by floods for (possibly) decades. heritage et al. (2001, figure 6) showed that ephemeral channels along the sabie are only affected by floods that have a return period of > 10 years, and based on river gauge data they estimated the february 2000 flood to have a return period of > 60 years. luminescence ages from river sand samples are older than the timing of significant recent flood events (e.g. from 2017, 2012, 2006, 2000), meaning that despite being very large events, these floods were possibly too rapid and carrying too much sand to allow the grains to become fully bleached, thus leading to inherited luminescence ages being present (knight & evans 2018). this also depends on the nature of the river reach itself. for example, in downstream reaches, floodwater can be accommodated across the width of the shallow floodplain. in upstream reaches where the river channel is constrained within a bedrock valley, accommodation space for floodwater is much less, leading to net erosion along constrained reaches and enhanced flooding and deposition immediately downstream at the transition into lowland areas (e.g. from geomorphic zones i to ii; figure 1a). this variability in behaviour takes place according to the characteristics of the river reach in question: no river exhibits the same behaviour patterns of change throughout its length. this is manifested clearly in different combinations of bedrock and sediment substrates, different river channel types and different in-channel and floodplain landforms (knight & evans 2017, 2018). based on the foregoing evidence, we propose a model of river system behaviour in which bedrockand sand-dominated reaches respond differently during flood events because of different initial sediment availability within and around the active channel (figure 2). under low-flow conditions, relict (inactive) sediment is found at higher elevations in bedrock hollows or across the floodplain (e.g. figure 1d). flood conditions result in excavation of sediment from these hollows, revealing a hard bedrock substrate. however, where a sediment floodplain exists, flooding leads to development of a scoured surface and active sediment deposition and erosion phases that extend across the floodplain (figure 2). higher sediment concentrations within the water column, over sandy substrates, lead to rapid development of migrating bedforms that are sustained by traction and saltation (knight & evans 2018). these sandy bedforms are generally absent in bedrock-dominated areas, even within active channels. figure 2: conceptual process model of the evolution of bedrock-dominated (top) and sand-dominated river channels (bottom) in the sabie river during low flow and then flood conditions. note that this then returns to low flow conditions afterwards. a schematic cross-section of the river profile is presented, with bedrock (brown) and sediment areas (yellow) shown. this model highlights that bedrockand sand-dominated river reaches can exhibit different behaviours during and after flood events (figure 2), and is consistent with previous work on sabie river responses to floods (e.g. heritage et al. 2015, their figure 4). the luminescence dating does not show a clear difference between the sediment depositional histories of bedrockand sand-dominated reaches (because luminescence samples are obtained from sediment, not bedrock); however, cunningham et al. (2015) argued that sediments deposited in hollows in bedrock-dominated reaches have quite different luminescence signals because of the episodic nature of their deposition. taken together, this shows that there is a decadal-scale tempo of river system responses to flood events along the sabie and likely other rivers in the region (colarossi et al. 2015; milan et al. 2018). bedrock-dominated substrates are less dynamic and show less interannual variation compared with sediment-dominated substrates, and this is simply controlled by sediment availability. understanding the nature of river system responses to flood events can inform on the most effective strategies for flood risk management. this is because different river reaches show different sensitivities to discharge variations, largely determined by their sediment availability and their landscape setting (figure 1a). river management, especially in areas with significant human presence, tends to focus on managing every flood event, no matter how small or large, rather than just the most geomorphically effective floods that result in greatest impacts. the political imperative for management means that annual floods, which are most important for ecosystems and biogeochemical cycling within rivers, are ‘over-managed’ rather than allowing for natural river flooding to take place across the width of the floodplain. this study suggests that bedrock and sediment reaches respond differently to flood events, and that they are located in different landscape contexts. this means that a single river management strategy that ignores differences between individual reaches is not likely to be successful. river management strategies that allow ‘space for water’ let the river systems respond naturally to variations in discharge, including allowing riparian and floodplain elements to be flooded (e.g. biron et al. 2014; buffin-bélanger et al. 2015). this more integrated approach to flood management works with, rather than against, the dynamics of the river system. it is the major (decadal-scale return period) floods that are most significant in geomorphic change along the sabie river, whereas smaller annual floods require spatial planning action (land-use zoning, infrastructure design, and riparian vegetation planting) rather than flood management action. on the sabie and other rivers in the inkomati catchment, this may mean that different flood management strategies are needed inside (in knp) and outside of formally protected areas (in headwaters) in response to both differences in river geomorphology and dynamics in these areas, and to differing community needs. conclusions the semiarid sabie river is affected by seasonal floods but their impacts vary between bedrockand sediment-dominated reaches. evidence for different river reach behaviour and their varied tempos of change can be evaluated based on river geomorphology, sediment patterns and informed by luminescence dating of phases of sediment deposition. the study results show that bedrockand sediment-dominated reaches are quite different in terms of their properties and dynamics, and this has implications for strategies towards flood management. acknowledgements competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions j.k. designed this study, j.k. and m.e. collected field data, j.k. and m.e. undertook lab analysis on samples, j.k. wrote the first draft of this article, and j.k. and m.e. contributed to the final draft of the submitted article. ethical considerations this study followed all ethical standards for research without direct contact with human or animal subjects. funding information this study is supported by national research foundation grant 91344 (to j.k.). the authors are very grateful to sanparks for permission to undertake research in kruger national park (permit knigj1225) and for logistic support. data availability the authors confirm that the data supporting the findings of this study are available within the article and in published articles cited in the text. disclaimer the views expressed in the submitted article are of the authors and not an official position of the institution or funder. references biron, p.m., buffin-bélanger, t., larocque, m., choné, g., cloutier, c.-a., ouellet, m.-a. et al., 2014, ‘freedom space for rivers: a sustainable management approach to enhance river resilience’, environmental management 54(5), 1056–1073. https://doi.org/10.1007/s00267-014-0366-z broadhurst, l.j. & heritage, g.l., 1998, ‘modelling stage-discharge relationships in anastomosing bedrock-influenced sections of the sabie river system’, earth surface processes and landforms 23(5), 455–465. https://doi.org/10.1002/(sici)1096-9837(199805)23:5<455::aid-esp860>3.0.co;2-2 buffin-bélanger, t., biron, p.m., larocque, m., demers, s., olsen, t., choné, g. et al., 2015, ‘freedom space for rivers: an economically viable river management concept in a changing climate’, geomorphology 251, 137–148. https://doi.org/10.1016/j.geomorph.2015.05.013 colarossi, d., duller, g.a.t., roberts, h.m., tooth, s. & lyons, r., 2015, ‘comparison of paired quartz osl and feldspar post-ir irsl dose distributions in poorly bleached fluvial sediments from south africa’, quaternary geochronology 30(part b), 233–238. https://doi.org/10.1016/j.quageo.2015.02.015 cunningham, a.c., evans, m. & knight, j., 2015, ‘quantifying bleaching for zero-age fluvial sediment: a bayesian approach’, radiation measurements 81, 55–61. https://doi.org/10.1016/j.radmeas.2015.04.007 entwistle, n., heritage, g., tooth, s. & milan, d., 2014, ‘anastomosing reach control on hydraulics and sediment distribution on the sabie river, south africa’, iahs publication 367, 215–219. https://doi.org/10.5194/piahs-367-215-2015 eze, p.n. & knight, j., 2018, ‘a geomorphological characterisation of river systems in south africa: a case study of the sabie river’, physics and chemistry of the earth 105, 196–205. https://doi.org/10.1016/j.pce.2018.01.001 gericke, o.j. & du plessis, j.a., 2012, ‘evaluation of the standard design flood method in selected basins in south africa’, journal of the south african institution of civil engineering 54(2), 2–14. heritage, g., tooth, s., entwhistle, n. & milan, d., 2015, ‘long-term flood controls on semi-arid river form: evidence from the sabie and olifants rivers, eastern south africa’, iahs publication 367, 141–146. https://doi.org/10.5194/piahs-367-141-2015 heritage, g.l., broadhurst, l.j. & birkhead, a.l., 2001, ‘the influence of contemporary flow regime on the geomorphology of the sabie river, south africa’, geomorphology 38(3–4), 197–211. https://doi.org/10.1016/s0169-555x(00)00090-8 heritage, g.l. & moon, b.p., 2000, ‘the contemporary geomorphology of the sabie river in the kruger national park’, koedoe 43(1), 39–55. https://doi.org/10.4102/koedoe.v43i1.207 heritage, g.l., van niekerk, a.w. & moon, b.p., 1997, ‘a comprehensive hierarchical river classification system’, geoöko plus 4, 75–84. knight, j., in press, ‘geomorphology and evolution of the limpopo river’, in f. eckardt (ed.), landforms and landscapes of botswana, springer. knight, j. & evans, m., 2017, ‘the sediment stratigraphy of a flood event: an example from the sabie river, south africa’, catena 151, 87–97. https://doi.org/10.1016/j.catena.2016.12.015 knight, j. & evans, m., 2018, ‘luminescence dating, sediment analyses, and flood dynamics on the sabie river, south africa’, geomorphology 319, 1–14. https://doi.org/10.1016/j.geomorph.2018.07.011 milan, d., heritage, g., tooth, s. & entwistle, n., 2018, ‘morphodynamics of bedrock-influenced dryland rivers during extreme floods: insights from the kruger national park, south africa’, gsa bulletin 130(11–12), 1825–1841. https://doi.org/10.1130/b31839.1 moon, b.p. & heritage, g.l., 2001, ‘the contemporary geomorphology of the letaba river in the kruger national park’, koedoe 44(1), 45–55. https://doi.org/10.4102/koedoe.v44i1.185 murray, a.s. & wintle, a.g., 2003, ‘the single aliquot regenerative dose protocol: potential for improvements in reliability’, radiation measurements 37(4–5), 377–381. https://doi.org/10.1016/s1350-4487(03)00053-2 pettit, n.e., naiman, r.j., rogers, k.h. & little, j.e., 2005, ‘post-flooding distribution and characteristics of large woody debris piles along the semi-arid sabie river, south africa’, river research and applications 21(1), 27–38. https://doi.org/10.1002/rra.812 rosgen, d.l., 1994, ‘a classification of natural rivers’, catena 22(3), 169–199. https://doi.org/10.1016/0341-8162(94)90001-9 rountree, m.w., rogers, k.h. & heritage, g.l., 2000, ‘landscape state change in the semi-arid sabie river, kruger national park, in response to flood and drought’, south african geographical journal 82(3), 173–181. https://doi.org/10.1080/03736245.2000.9713711 saraiva okello, a.m.l., masih, i., uhlenbrook, s., jewitt, g.p.w., van der zaag, p. & riddell, e., 2015, ‘drivers of spatial and temporal variability of streamflow in the incomati river basin’, hydrology and earth system sciences 19(2), 657–673. https://doi.org/10.5194/hess-19-657-2015 schutte, i.c., 1986, ‘the general geology of the kruger national park’, koedoe 29(1), 13–37. https://doi.org/10.4102/koedoe.v29i1.517 seanego, k.g. & moyo, n.a.g., 2013, ‘the effect of sewage effluent on the physico-chemical and biological characteristics of the sand river, limpopo, south africa’, physics and chemistry of the earth 66, 75–82. https://doi.org/10.1016/j.pce.2013.08.008 van niekerk, a.w., heritage, g.l. & moon, b.p., 1995, ‘river classification for management: the geomorphology of the sabie river in the eastern transvaal’, south african geographical journal 77(2), 68–76. https://doi.org/10.1080/03736245.1995.9713594 filelist convert a pdf file! page 1 page 2 page 3 foord.qxd a check list of the spider fauna of the western soutpansberg, south africa (arachnida: araneae) s.h. foord, anna s. dippenaar-schoeman and m. van der merwe foord, s.h., anna s. dippenaar-schoeman and m. van der merwe. 2002. a check list of the spider fauna of the western soutpansberg, south africa (arachnida: araneae). koedoe 45(2): 35–43. pretoria. issn 0075-6458. by virtue of its geological history and geographical location the soutpansberg constitutes a refuge for a high diversity of organisms. the western soutpansberg forms part of the savanna biome and is presently the area with the highest concentration of natural heritage sites in south africa. a unique private initiative is under way to improve its national and international conservation status in a bid to conserve the mountain. a checklist of the spider species of the western soutpansberg collected over a five-year period is presented. forty-six families, represented by 109 genera and 127 species have been collected. of the species collected, 81 (64 %) were wandering spiders and 46 (36 %) web builders. the thomisidae have the highest number of species (15) followed by the araneidae and the salticidae with 10 species each. ninety-six genera are represented by a single species. ninety six percent of the species collected are new records for the area. this survey is the first for the area and forms part of the south african national survey of arachnida (sansa). keywords: biodiversity, guilds, conservancy. s.h. foord ( ) and m. van der merwe, department of biological sciences, university of venda, thohoyandou, 0950, republic of south africa; anna s. dippenaar-schoeman, arc-plant protection research institute, private bag x134, pretoria, 0001 republic of south africa. issn 0075-6458 35 koedoe 45/2 (2002) introduction in ratifying agenda 21 at the earth summit in rio de janeiro (1992) and the convention on biological diversity in 1995, south africa is obliged to develop a strategic plan for the conservation and sustainable use of biodiversity. however, meaningful conservation cannot take place if the species involved are not known. although araneae constitutes an abundant and highly successful group of invertebrates, little is known about their diversity in large parts of south africa. compared with areas in the northern hemisphere ecological surveys of african spiders are sparse (dippenaar-schoeman & jocqué 1997), this is also true for spider fauna in conserved areas with the only check lists compiled being those for the mountain zebra national park (dippenaar-schoeman 1988), roodeplaat dam nature reserve (dippenaar-schoeman et al. 1989), karoo national park (dippenaar-schoeman et al. 1999) and makelali nature reserve in the limpopo province (whitmore et al. 2001). in 1997, the south african national survey (sansa) was initiated. one of its aims is to make inventories of the arachnid fauna of south africa (dippenaar-schoeman & craemer 2000). as part of sansa, various projects are in progress, such as inventories of the spider fauna in different floral biomes and to determine the number of species that are already protected in existing conservation areas. the limpopo province is often perceived as savanna and bushveld, however, the soutpansberg mistbelt has a totally unique composition of plants, animals, and invertebrates. by virtue of its geological history and geographical location the soutpansberg mounfoord.qxd 2005/12/09 10:03 page 35 tain archipelago constitutes a refuge for a disproportionately high diversity of organisms. the floral environment is very diverse, with a higher generic diversity than that of the cape flora and include the following: savannah bushveld, indigenous forests with ancient yellowwood trees, very 'sensitive wetlands', and soutpansberg fynbos similar to that found in the western cape (hahn pers. comm.). the western soutpansberg is presently the area with the highest concentration of natural heritage sites in south africa and is part of a unique private initiative to improve its national and international conservation status in a bid to conserve the mountain (rosmarin et al. 1998). this involves the establishment of various reserves and a conservancy. a process is also in motion to get the soutpansberg recognized as a world heritage site and also to include it in unesco's man and biosphere program as a biosphere reserve. the eastern part of the soutpansberg, especially the area around the hanglip forest, has been fairy well sampled and numerous species described. however, very little is known about the western part. in this study an inventory was made of spider species present in the western soutpansberg. spiders were collected from various habitats over a 5-year period. this is the first inventory of the spiders from the western soutpansberg. it is a contribution towards the soutpansberg conservancy objectives and also forms part of the south african national survey of arachnida (sansa) for the limpopo province and savanna biome. study area the soutpansberg is the northernmost mountain range in south africa, situated within the limpopo province and stretches about 100 km in an east-west direction. the western soutpansberg, which this survey deals with, stretches from the sand river, which is near louis trichardt to the town of vivo about 70 km west (fig. 1). the area consists of 4 physiognomic units: (1) the southern plains; (2) western plains (1200 m koedoe 45/2 (2002) 36 issn 0075-6458 fig.1 position of the study area within south africa. above sea level (a.s.l.)); (3) the middle plateau (1200-1300 m a.s.l.) the mountain range (1300-1900 m a.s.l.); and (4) the northern upper plateau (1600-1900 m a.s.l.). the highest peak of the soutpansberg, letjume (23º02's, 29º26'e) at 1747 m a.s.l., falls within this area. materials and methods sporadic collecting was mainly undertaken from 1996–2000. spiders were sampled by hand (plant and ground search, turning rocks, and sifting of leaf litter) or using a sweepnet or beating tray for grass and low shrubs. pitfall trapping was undertaken for a 10-week period (from 3 may 1997–13 july 1997), with 50 pitfall traps in an old and secondary growth forest at the farm lajuma. since collecting for the current study was on an ad hoc basis for relatively short periods, the results may not reflect the true diversity and species richness of the area, but nevertheless, they give an indication of species present (minimum community). it is also important to note that most of the collecting took place on the farm lajuma, restricting most of the collecting to an area < 500 ha. the material collected was identified by the second author and is housed in the national collection of arachnida (nca) at the arc-plant protection research institute in pretoria. general ecology ecological and behavioural classification follows dippenaar-schoeman et al. (1999) and is summarised in table 1. foord.qxd 2005/12/09 10:03 page 36 results and discussion numbers present during the five-year survey a total of 46 families represented by 109 genera and 127 species were collected (table 3). of the species collected 81 (64%) were wandering spiders and 46 (36%) web builders. all the species are new records for the western soutpansberg. the thomisidae had the highest number of species (15) followed by the araneidae and salticidae each with 10 species, and the gnaphosidae with eight. ninety-six genera are each represented by a single species. one new species, the miturgid, cheiromiona lajuma, has been described, and some undetermined species might be new to science e.g. poecilomigas sp. and mimetus sp. however, only after revisionary studies could this be determined. ground layer twenty-eight families, represented by 53 species are associated with the ground layer (tables 2 & 3). ground web dwellers: 15 species make their webs on or close to the ground surface. the following web dwellers were found: the funnel-webs of the agelenidae and sheetwebs of the hahniidae are made mainly in open grass areas; retreat-webs of the families amaurobiidae (pseudauximus), eresidae (penestomus), hersiliidae (tama), oecobiidae (uroecobius) and phyxelididae (vidole) are made on or below rocks; space-webs of the pholcidae is made in old mammal-made holes and in dark crevices and the gum-foot web of a steatoda sp. of the theridiidae are made under stones. this is the first record of penestomus (eresidae) and tama (hersiliidae) collected outside the cape provinces issn 0075-6458 37 koedoe 45/2 (2002) table 1 ecological and behavioural classification of spider taxa (dippenaar-schoeman et al. 1999) guilds micro-habitats wandering spiders (w) ground wanderers (gw) free living (fgw) burrow living (bgw) plant wanderers (pw) free living (fpw) burrow living (bpw) web builders (wb) adapted orb-webs (aowb) funnel-web (fwb) gumfoot-web gwb) orb-web (owb) retreat-web (rwb) sheet-web (shwb) space-web (swb) table 2 spider species associated with the ground and field layers of the western soutpansberg layer webs % burrows % free % total living ground 15 28 4 8 34 64 53 field 31 42 2 3 41 55 74 total 46 6 75 127 of south africa (dippenaar-schoeman 1989; smithers 1945). ground burrow dwellers: the burrow living ground dwellers are all members of the mygalomorphae and represent two trapdoor spider species of the families cyrtaucheniidae, idiopidae, and two baboon spiders of the family theraphosidae. homostola pardalina (hewitt) (cyrtaucheniidae) is the first record of this genus from the limpopo foord.qxd 2005/12/09 10:03 page 37 koedoe 45/2 (2002) 38 issn 0075-6458 table 3 check list of spiders of the western soutpansberg region collected from 1996 to 2000 family, genus, species guilds habitat abundance 1. agelenidae benoitia australis (simon, 1896) fwb g 2 olorunia ocellata pocock, 1900 fwb g 2 2. amaurobiidae pseudauximus annulatus purcell, 1908 rwb g 2 3. anapidae metanapis bimaculata (simon, 1895) owb f 1 4. araneidae araniella (undetermined sp.) owb f 1 araneus strupifera simon, 1886 owb f 2 caerostris sexcuspidata (fabricius, 1793) owb f 2 cyclosa insulana (costa , 1834) owb f 3 cyrtophora citricola (forskal, 1775) owb f 3 gasteracantha sanguinolenta c.l. koch,1845 owb f 3 neoscona blondeli (simon, 1885) owb f 3 neoscona quincasea roberts, 1983 owb f 2 neoscona subfusca (c.l. koch, 1837) owb f 3 nemoscolus vigintipunctatus simon, 1897 owb f 2 5. caponiidae caponia sp. (undetermined sp.) fgw g 2 6. clubionidae clubiona sp. (undetermined sp.) fpw f 1 clubiona lawrencei roewer, 1951 fpw f 2 7. corinnidae castianeira fulvipes simon, 1896 fgw g 1 cetonana simoni (lawrence, 1942) fgw g 2 trachelas scopulifer simon, 1896 fgw g 1 8. ctenidae ctenus transvaalensis benoit, 1981 fgw g 2 9. cyatholipidae cyatholipus isolatus griswold, 1987 shwb f 1 10. cyrtaucheniidae homostola pardalina (hewitt, 1913) bgw g 2 11. deinopidae deinopis cornigera gerstaecker, 1873 aowb f 1 12. dictynidae devade sp. (immature) rwb f 2 13. eresidae penestomus sp. (immature) rwb g 1 14. gnaphosidae aphantaulax inornata tucker, 1923 fgw g 3 asemesthes numisma tucker, 1923 fgw g 3 drassodes (undetermined sp.) fgw g echemus erutus tucker, 1923 fgw g 3 megamyrmekion transvaalense tucker, 1923 fgw g 2 setaphis arcus tucker, 1923 fgw g 3 foord.qxd 2005/12/09 10:03 page 38 issn 0075-6458 39 koedoe 45/2 (2002) xerophaeus sp. (undetermined sp.) fgw g 1 zelotes tuckeri roewer, 1951 fgw g 3 15. hahniidae hahnia sp. (immature) swb g 1 16. hersiliidae hersilia sericea pocock, 1898 fpw f 1 tama arida smithers 1945 rwb g 1 17. idiopidae idiops castaneus hewitt, 1913 bgw g 1 18. linyphiidae microlinyphia sterilis (pavesi, 1883) shwb g 2 neriene natalensis van helsdingen, 1969 shwb g 2 linyphiidae genus a shwb f 1 linyphiidae genus b shwb f 1 19. liocranidae hortipes contubernalis bosselaers & jocqué, 2000 fgw g 1 20. lycosidae pardosa crassipalpis purcell, 1903 fgw g 3 pardosa leipoldti purcell, 1903 fgw g 2 proevippa fascicularis (purcell, 1903) fgw g 3 proevippa wanlessi (russell-smith, 1981) fgw g 1 21. migidae poecilomigas (undetermined new sp.) bpw f 1 22. mimetidae mimetus (undetermined new sp.). fpw f 1 23. miturgidae cheiracanthium africanum lessert, 1921 fpw f 3 cheiramiona simplicitarse simon, 1910 fpw f 2 cheiramiona (new sp. in press) fpw f 1 24. mysmenidae undetermined new sp. owb f 1 25. nemesidae entypesa schoutedeni benoit, 1965 bpw f 1 26. oecobiidae uroecobius ecribellatus kullman & zimmerman, 1976 rwb g 2 27. oxyopidae hamataliwa kulczynskii (lessert, 1915) fpw f 2 oxyopes jacksoni lessert, 1915 fpw f 2 oxyopes longispinosus lawrence, 1938 fpw f 2 oxyopes pallidecoratus strand, 1906 fpw f 2 peucetia viridis (blackwall, 1858) fpw f 1 28. palpimanidae palpimanus transvaalicus simon, 1893 fgw g 1 29. philodromidae philodromus browningi lawrence, 1952 fgw f 1 suemus punctatus lawrence, 1938 fgw f 1 thanatus vulgaris simon, 1870 fgw f 2 table 3 (continued) family, genus, species guilds habitat abundance foord.qxd 2005/12/09 10:03 page 39 koedoe 45/2 (2002) 40 issn 0075-6458 30. pholcidae micropholcus (undetermined new sp.) spwb g 1 pholcus ciliatus lawrence, 1938 spwb g 2 smeringopus natalensis lawrence, 1947 spwb g 2 spermophora peninsulae lawrence, 1964 spwb g 2 31. phyxelididae vidole sothoana griswold, 1990. rwb g 2 32. pisauridae euprosthenopsis pulchella (pocock, 1902) fwb f 2 cispius problematicus blandin, 1978 fpw f 2 33. prodidomidae austrodomus sp. (undetermined sp.) fgw g 1 theuma purcelli tucker, 1923 fgw g 2 34. salticidae brancus bevisi lessert, 1925 fpw f 2 cosmophasis australis simon, 1902 fpw f 2 heliophanus orchesta simon, 1885 fgw g 2 festucula sp. (immature) fgw f 1 massagris sp. (undetermined new sp.) fgw g 2 natta horizontalis karsch, 1879 fgw g 2 pachyballus transversus simon, 1900 fpw f 1 stenaelurillus (undetermined sp.) fpw f 2 thyene inflata (gerstaecker, 1875) fpw f 2 thyenula ogdeni (peckham & peckham, 1903) fpw f 3 35. scytodidae scytodes sp. a fgw g 1 scytodes sp. b fgw g 2 scytodes fusca walckenaer, 1837 fgw g 3 36. segestriidae ariadna (undetermined sp.) rwb f 2 37. selenopidae selenops brachycephalus lawrence, 1940 fgw g 2 selenops tenebrosus lawrence, 1940 fgw g 2 selenops zuluanus lawrence, 1940 fgw g 1 38. sicariidae loxosceles spiniceps lawrence, 1952 fgw g 1 39. sparassidae fgw g 1 olios (undetermined sp.). fpw f 1 palystes johnstoni pocock, 1896 fpw f 2 40. thomisidae diaea puncta karsch, 1884 fpw f 2 heriaeus fimbriatus lawrence, 1942 fpw f 1 misumenops rubrodecoratus millot, 1941 fpw f 3 monaeses austrinus simon, 1910 fpw f 2 oxytate argenteooculata (simon, 1886) fpw f 2 oxytate concolor (caporiacco, 1947) fpw f 1 runcinia aethiops simon, 1901 fpw f 2 runcinia flavida simon, 1881 fpw f 3 synema imitator roewer, 1951 fpw f 2 thomisops pupa karsch, 1879 fpw f 2 thomisus daradioides simon, 1890 fpw f 3 table 3 (continued) family, genus, species guilds habitat abundance foord.qxd 2005/12/09 10:03 page 40 province (dippenaar-schoeman 2002). they make their burrows underneath rocks with the trapdoor covered with plant material. idiops castaneus hewitt (idiopidae) was previously collected from newington, in the limpopo province. this record extends its range within the province. members of idiops make their burrows on slightly sloping grassy planes (dippenaar-schoeman 2002). of the two theraphosids, the horned baboon spider ceratogyrus bechuanicus purcell are commonly found throughout the limpopo province, while the occurrence of harpactirella flavipilosa lawrence is a new record for the province. free-living ground wanderers: most ground dwellers (64 %) are free-living species that are not constructing webs to catch their prey (table 2). the families gnaphosidae, with eight species, and lycosidae with four, are issn 0075-6458 41 koedoe 45/2 (2002) thomisus. granulatus karsch, 1880 fpw f 1 thomisus. kalaharinus lawrence, 1936 fpw f 2 tmarus cameliformis millot fpw f 2 xysticus natalensis lawrence, 1938 fgw g 2 41. tetragnathidae glenognatha (undetermined sp.) owb f 1 leucauge festiva (blackwall, 1866) owb f 2 leucauge decorata (blackwall, 1864) owb f 2 nephila pilipes (fabricius, 1793) owb f 2 42. theraphosidae ceratogyrus bechuanicus purcell, 1902 bgw g 1 harpactirella flavipilosa lawrence,1936 bgw g 2 43. theridiidae coscinida tibialis simon, 1895 gwb f 1 crustulina sp. (undetermined sp.). gwb f 1 latrodectus geometricus o.p.-cambridge, 1904 gwb f 2 phoroncidia eburnea (simon, 1895) gwb f 1 steatoda (undetermined sp.). gwb g 2 tidarren (undetermined sp.). gwb f 1 theridion (undetermined sp.). gwb f 3 44. trochanteriidae platyoides walteri (karsch, 1886) fpw f 1 45. uloboridae miagrammopes sp. (immature) aowb f 1 uloborus lugubris berland, 1939 owb f 3 uloborus plumipes lucas, 1845 owb f 2 46. zodariidae diores auricula tucker, 1920 fgw g 2 psammoduon (undetermined sp.) fgw g 1 guilds: fgw = free living ground wanderers; bgw = burrow living ground wanderers; fpw = free living plant wanderers; bpw = burrow living plant wanderers. types of webs: aowb= adapted orb-web; fwb = funnel-web; gwb = gumfootweb; owb = orb-web; rwb = retreat-web; swb = sheetweb; spwb = space-web. habitat: f = foliage (plants); g = ground. abundance: 1 = rare; 2 = common; 3 = abundant. table 3 (continued) family, genus, species guilds habitat abundance foord.qxd 2005/12/09 10:03 page 41 the most abundant. the wolf spider, proevippa wanlessi russell-smith, represented 57 % of the total number of spiders that were caught during the pitfall survey and account for most of the specimens caught in the secondary growth forest. their numbers decreased markedly in the old growth forest. the two gnaphosids, zelotes tuckeri roewer and megamyrmekion transvaalense tucker were more commonly found in the oldgrowth forest than in the second growth forest. field layer plant web dwellers: from the field layer 42 % of the species collected were associated with webs (table 2). the orb-web spiders were represented by five families (16 spp.) with the araneidae the more abundant (10 species). representatives of some rare families, such as anapidae and mysmenidae have also been collected. little is known about these families in south africa and both probably represent new species. several of the orb webs constructed where large and made by members of the tetragnathids, the golden orb-web spiders (nephila) and two silver vlei spider species of the genus leucage. the physical structure of the habitat plays a role in the composition of the spider web dwellers (wise 1993) as it not only provides the necessary support for anchoring webs but also increases the availability of retreat space and lead to the modifification of microclimate, which could have an effect on the spiders as well as their prey. plant burrow dwellers: the nemesid trapdoor spider entypesa schoutedeni benoit was collected from transparent webbing from under bark. the type locality of this species, is soutpansberg, but no exact locality was indicated. this species seems to be endemic to the soutpansberg. the migid also known as the tree trapdoor spider probably belongs to a new species of poecilomigas (griswold pers. comm.). the genus makes sac-like burrows beneath the bark with a wafer-lid trapdoor (dippenaar-schoeman 2002). this is the first record of the genus from the limpopo province. free-living plant wanderers: 55 % of all spiders collected from the field layer were freeliving spiders. they were represented by nine families and 34 species. most species were collected with sweepnets from grass. the more common grass dwellers were the clubionidae, miturgidae, oxyopidae, philodromidae, thomisidae and salticidae. a few species are almost exclusively tree dwellers, such as the hersiliidae (hersilia) and trochanteriidae (platyoides). new species of cheiramiona (miturgidae) and mimetus (mimetidae) were collected. conclusions except for taxonomic descriptions the spider fauna of south africa is poorly known. our knowledge of the spiders of the limpopo province, in particular, is based only on short collecting expeditions. the only long-term survey of spiders from this province was a study of a savanna ecosystem at the makelali nature reserve where a total of 37 families, 147 genera and about 268 species were collected (whitmore et al. 2001). this is the first survey of the spiders of the western soutpansberg. about 96 % of the species collected are new records for the area with 10 possibly new species. the 127 species probably represents only a portion of the spider fauna present. however, it compares well with other more comprehensive surveys that have been undertaken in south africa for example: roodepaat dam nature reserve (98 spp,), rietondale research station (55 spp.), the karoo national park (116 spp.), and kruger national park (139 spp.). at family level, the spider fauna from the western soutpansberg is more diverse than any of the above surveys. a direct positive correlation between family and species diversity has been observed for other taxa (andersen 1995; balmford et al. 1996), pointing to a disproportionately high spider diversity for this area and higher biological koedoe 45/2 (2002) 42 issn 0075-6458 foord.qxd 2005/12/09 10:03 page 42 diversity in terms of kinds and disparity of attributes of species. although the araneae are an abundant and highly successful group of invertebrate animals, little has been done to conserve them in south africa. with this rich fauna of spiders present, it fulfils the objectives of the western soutpansberg conservancy. an awareness of spiders present might stimulate additional collecting and more research. in the past two decades both specialists and laymen have shown considerable interest in spiders (dippenaar-schoeman & van den berg 1988) and it is accepted that eco-tourism will be one of the primary future land uses within the soutpansberg conservancy area. to this end an overall eco-tourism theme that retains and enhances the uniqueness of the area should be striven for. spiders may play an important role here. acknowledgements we thank ian and retha gaigher for allowing us to work on their farm lajuma, providing accommodation, and assisting in the surveys. this work was financially supported by a university of venda grant to the first author. references anderson, a.n. 1995. measuring more diversity: genus richness as a surrogate for species richness in australian ant faunas. biological conservation 73: 39–43. balmford, a., a.h.m. jayasuriya, m.s.b. green. 1996. using higher taxon richness as a surrogate for species richness. ii local applications. proceedings of the royal society of london b 263: 1571–1575. dippenaar-schoeman, a.s. 1988. annotated check list of spiders (araneae) of the mountain zebra national park. koedoe 31: 151–160. dippenaar-schoeman, a.s. 1989. the african species of the sub-family penestominae (araneae: eresidae): with description of two new species. phytophylactica 21: 131–134. dippenaar-schoeman, a.s. 2002. baboon and trapdoor spiders of southern africa: an identification manual. pretoria: plant protection research institute, agricultural research council. (handbook no. 13). dippenaar-schoeman, a.s. & c. craemer. 2000. the south african national survey of arachnida. plant protection news 56: 11–12. dippenaar-schoeman, a.s. & r. jocqué. 1997. african spiders, an identification manual. pretoria: plant protection research institute, agricultural research council. (handbook no. 9). dippenaar-schoeman, a.s., a. leroy, m. de jager & a. van den berg. 1999. spider diversity of the karoo national park, south africa (arachnida: araneae). koedoe 42: 31–42. dippenaar-schoeman, a.s., a.m. van den berg & a. van den berg. 1989. species composition and relative seasonal abundance of spiders from the field and tree layers of the roodeplaat dam nature reserve. koedoe 32: 51–60. rosmarin, j., i.g. gaigher, n. hahn. 1998. the western soutpansberg conservancy: towards a proposed policy framework and implementation strategy. unpublished report. smithers, r.h.n. 1945. the hersiliidae of south africa. transactions of the royal society of south africa 31: 1–18. whitmore, c., r. slotow, t.e. crouch & a.s. dippenaar-schoeman. 2001. checklist of spiders (araneae) from savanna ecosystem, northern province, south africa: including a new family record. durban museum novitates 26: 10–19. issn 0075-6458 43 koedoe 45/2 (2002) foord.qxd 2005/12/09 10:03 page 43 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <feff004f007000740069006f006e007300200070006f0075007200200063007200e900650072002000640065007300200064006f00630075006d0065006e00740073002000500044004600200064006f007400e900730020006400270075006e00650020007200e90073006f006c007500740069006f006e002000e9006c0065007600e9006500200070006f0075007200200075006e00650020007100750061006c0069007400e90020006400270069006d007000720065007300730069006f006e00200070007200e9007000720065007300730065002e0020005500740069006c006900730065007a0020004100630072006f0062006100740020006f00750020005200650061006400650072002c002000760065007200730069006f006e00200035002e00300020006f007500200075006c007400e9007200690065007500720065002c00200070006f007500720020006c006500730020006f00750076007200690072002e0020004c00270069006e0063006f00720070006f0072006100740069006f006e002000640065007300200070006f006c0069006300650073002000650073007400200072006500710075006900730065002e> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <feff004200720075006b00200064006900730073006500200069006e006e007300740069006c006c0069006e00670065006e0065002000740069006c002000e50020006f00700070007200650074007400650020005000440046002d0064006f006b0075006d0065006e0074006500720020006d006500640020006800f80079006500720065002000620069006c00640065006f00700070006c00f80073006e0069006e006700200066006f00720020006800f800790020007500740073006b00720069006600740073006b00760061006c00690074006500740020006600f800720020007400720079006b006b002e0020005000440046002d0064006f006b0075006d0065006e0074006500720020006b0061006e002000e50070006e006500730020006d006500640020004100630072006f0062006100740020006f0067002000520065006100640065007200200035002e00300020006f0067002000730065006e006500720065002e00200044006900730073006500200069006e006e007300740069006c006c0069006e00670065006e00650020006b0072006500760065007200200073006b00720069006600740069006e006e00620079006700670069006e0067002e> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice introduction methods sites results and discussion addo elephant national park gardens addo elephant national park zuurberg section conclusion acknowledgements references about the author(s) hugo bezuidenhout south african national parks, scientific services, kimberley, south africa applied behavioural ecology and ecosystem research unit, university of south africa, florida, south africa citation bezuidenhout, h., 2020, ‘assessment results (2015–2018) of re-established poached cycad trees in addo elephant national park, eastern cape, south africa’, koedoe 62(1), a1626. https://doi.org/10.4102/koedoe.v62i1.1626 short communication assessment results (2015–2018) of re-established poached cycad trees in addo elephant national park, eastern cape, south africa hugo bezuidenhout received: 14 feb. 2020; accepted: 12 may 2020; published: 14 july 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. introduction on 12 february 2015, the south african police services confiscated 39 encephalartos longifolius plants (thunberg’s cycad or zuurberg cycad) and nine encephalartos lehmannii (karoo cycad) plants illegally removed from the wild on a farm adjacent to the darlingtondam section of addo elephant national park (aenp). the confiscated plants were delivered to aenp on saturday, 14 february 2015 (figure 1). figure 1: cycad plants recovered from poachers. (a) and (b) e. longifolius. according to the 1997 and 2006 international union for conservation of nature red list of threatened plant species, 82% of the world’s cycads are listed as threatened and, thus, are facing a major extinction crisis (singh 2012). some species are dying out naturally, but the major extinction contributing factor is human activities (donaldson 2003). in south africa, cycads were declared ‘specially protected plants’ in 1971 in the transvaal (van der walt 2014) and cycad owners had to apply for a permit. since then, all other provinces have implemented strict protective laws and heavy penalties were laid down by the nature conservation ordinances of each province (singh 2012). cycads (encephalartos) are known as the most primitive living seed-bearing plants belonging to an ancient order of cycadales, which flourished in the carboniferous period about 100 million years ago (singh & campbell 2011). cycads belong to a group of plants known as gymnosperms (plants which bear naked seeds) and their reproductive organs are produced in cones (giddy 1974). these cycad plants have survived for over 150 million years with little change in their basic structure, and neither fire nor drought presents any real threat to their survival (giddy 1974). cycads are dioecious, meaning that male and female cones occur on separate plant individuals and, in order for pollination to take place, male and female plants have to cone at the same time and be close to one another (giddy 1974). south africa is considered to be one of the centres of cycad diversity, hosting more than half of the known encephalartos species in africa, with 76% of these species endemic to south africa. sadly, 78% of the south african encephalartos species are threatened with extinction (van der walt 2014). according to van der walt (2014), encephalartos species are collectively the most threatened plant species group in south africa today, and also the most threatened group of plants in the world (department of environmental affairs 2015). twelve of the 37 (32%) encephalartos species are regarded as critically endangered, whilst an additional three are already considered extinct in the wild. there are less than 100 plants left in the wild for seven of the critically endangered species, four species of which are on the brink of extinction. a further four encephalartos species are regarded as endangered. the predominant threat facing cycads is the ongoing illegal removal of adult plant individuals from wild populations to meet the current demand for large cycads for private collections and for landscaping purposes. recently, cycads have also been poached for use in muthi-markets, where these plants are used for traditional purposes. adult plant individuals are also highly valued as parental stock for seedling propagation for both the domestic and international cycad trade. all species of the genus encephalartos are included in appendix i of the convention on international trade in endangered species of wild fauna and flora (cites) (bezuidenhout & hofmeyr 2016). a key mandate for the south african national parks (sanparks) is to continue contributing to the representation of threatened ecosystems and plant species in a national protected area network. the south african national parks’ mandate also extends to ensure conservation and protection of cycads under their jurisdiction (bezuidenhout et al. 2017; department of environmental affairs 2015; staatskoerant 2017). the complex nature of the supply and demand of cycads has led to these plants becoming a human dependent for survival and persistence in its natural habitat (van der walt 2014). south african national parks therefore decided to assist in the re-establishing of the confiscated plants in the aenp. donaldson (1995) noted that conservation practitioners might base their actions on generalisations and assumptions, in the absence of detailed information on how cycad species respond to certain actions. donaldson (1995) classified both cycad species (e. lehmannii and e. longifolius) as type 2, namely as a persister/reproducer, with the following life history traits: (1) aerial stems reaching several meters in height with several basal suckers; (2) a single (often large) cone per plant, whilst the environment of these cycads correlate well with an arid environment of fire climax vegetation and a high incidence of mast seeding in fire climax vegetation. a literature review revealed no published information on how unearthed planted cycads have responded to being re-established. partly related to this subject, boyd (1995) described five ex situ conservation measures in the translocation or relocation of several cycad plant species that provincial conservation authorities were involved with, with mixed success results. the long-term feasibility of reintroduction and translocation has not been determined, as it involves long-term monitoring and active management (boyd 1995). cousins and witkowski (2017) noted that: [i]t appears that most african cycad species are fire-tolerant, and that the fine-leaved, woolly-coned encephalartos species generally show the most pronounced positive response to fire, the phenomenon of fire-stimulated leaf and cone production is still poorly understood, and requires further study. further investigation is needed to determine the actual fire-related cues that trigger leaf and cone production. (p. 166) the main aim of this paper is, therefore, to document the actions taken to re-establish the confiscated cycad plants (e. lehmannii and e. longifolius) in two habitat sites: (1) the aenp gardens; and (2) in the zuurberg section of aenp, where a natural fire regime exists. methods the species were identified, numbered, sexed (if possible), and the plant length, stem diameter, injury (%), cones absent/present, leaves absent/present, single or multi-stemmed were recorded. treatment of the cycads consisted of removing all of the leaves and cones as well as chemical treatment. the cycad individuals were chemically treated with dynaroot hormone powder (no. 2) (to stimulate new root growth), bravo 720 with chlorothalonil (active ingredient mostly to prevent fungal diseases attacking the stressed cycads), 20 ml chloropiriphos to kill any insects, and blue tree seal (to enhance recuperation after replanting by protection against fungi, and to keep roots healthy for maximum water absorption). the cycads were replanted between 23–26 february 2015 in accordance with guidelines proposed by bezuidenhout (2015). in march 2018, all of the localities were re-visited and all of the plants were re-assessed. a photograph of each individual was taken. the health of the individual was assessed in three categories: (1) the presence or absence of leaves (healthy with > 4 leaves), (2) dormant without any damage (standing with firm stem and crown, but no leaves present), and (3) dead by being damaged and/or lying on the ground or nothing left (no leaves, not standing, disintegrated stem most of it not present [> 75%] or pieces of stem lying around at the locality). all of the cycads were checked thoroughly for injuries and insects that might have a negative impact on their condition and recovery. sites after discussions, consensus was reached that, as a result of (1) time constraints, (2) lack of man power, (3) costs, (4) original site of the cycads being outside of the aenp, (5) inhospitability of original site and difficulty to reach the terrain, and (6) ease of plot-established monitoring, the recovered cycad individuals would be planted in the zuurberg section and gardens of the aenp offices. four possible sites for planting the e. longifolius (thunberg’s or zuurberg cycad) cycads in the zuurberg section of aenp were identified. the few individuals of e. lehmannii (karoo cycad) were planted in one garden in aenp, whilst four badly-injured e. longifolius were planted in another section of the gardens in aenp. ethical consideration i confirm that ethical clearance was not required for the study. results and discussion in total, 39 e. longifolius cycad plants were replanted; six of these were young individuals, where either suckers or branches had broken off during handling. the sex ratio was eight female plants to 17 males. because of the absence of cones, the sex of the other 14 plant individuals could not be determined at the time. twenty of these cycads were more than 3 m tall, nine were about 2 m tall, four about 1 m tall and the other six were small individuals (0.2 m to 0.4 m). thirty-two cycads were single stemmed and one was multi-branched, and six young cycad suckers were recorded (table 1) (bezuidenhout 2015). table 1: summary of cycad individuals. all of these cycads were in fairly good condition even though they had been handled roughly. bruises were noted on the individual cycads, indicating that plants were pushed over by a tractor and then pulled across rocks and hard soil surfaces using ropes. termite activity was noticed on one of the cycads and it was treated. nine e. lehmannii individuals were planted, including four young individuals (unknown sex) derived from suckers or branches that had broken off during the handling of these plants. the sex ratio for the five adult individuals was one male, one female and three of unknown sex. the height classes of the nine individuals comprised one that was approximately 2 m tall, four about 1 m tall, and the other four young individuals being 0.2 m to 0.4 m tall (table 1) (bezuidenhout 2015). addo elephant national park gardens the individuals that were selected to be planted in one of the aenp gardens were either e. lehmannii or could not be identified because of the absence of leaves. six suckers were also planted in the aloe species rocky garden. in a separate second aenp garden, four individuals of e. longifolius were planted. all four individuals were seriously damaged either by fire scars or from being injured when removed from the veld. five of the adult monitored individuals had more than four leaves (figure 2), and two had suckers bearing leaves. from the six individual suckers that were planted in the aloe species dominated garden, four had healthy leaves whilst two had died. three adult cycad individuals are standing but without leaves, whilst four are dead and have been removed (table 2). figure 2: healthy planted individual encephalartos lehmannii cycad in addo elephant national park garden. table 2: monitoring result for planted cycads in gardens of addo elephant national park. after 3 years, we concluded that of the 18 cycad individuals planted in the gardens of aenp, nine are in good condition with leaves (50%) and six are dead (33%). because cycads can remain dormant for a long time without bearing leaves, it is difficult to assess whether individuals are dead or alive when the stem is still in seemingly good condition. the condition of three of the above-mentioned cycads could not be determined accurately and was recorded as standing (17%). because the habitat of this locality is a human-made garden, these cycads receive more water on a regular basis, the soil was disturbed (construction of building) and drainage might not be ideal compared to the zuurberg localities. despite these, the overall result indicates a significantly positive response. limited impact by wildlife was noted. addo elephant national park zuurberg section all of the cycads that were replanted in the zuurberg section were e. longifolius individuals. care was taken to plant cycads of different sexes in one stand or group. according to advice from experts, these cycads were planted 8 m – 10 m apart (cousins & witkowski 2017; donaldson 1995). four sites were used for planting cycads. a few months after planting, one individual showed new leaf growth and some coned. ants were observed at one individual, but otherwise all of the cycad individuals appeared to be in good condition (bezuidenhout 2015). site 2 at site 2, six cycad individuals were replanted in february 2015. when inspected in 2018, one adult individual was recorded to have more than four leaves and was in a healthy condition. fire scars were noted on the leaves and stem as a result of a fire during the previous year. one individual was still in an upright position with a sucker that might still grow, but showed extensive damage to the main stem caused by baboons. one other individual was recorded as dormant (still standing, but without leaves). three individuals had died, leaving either no trace of stems, or the remains of stems having been scattered in the veld (table 3). table 3: monitoring result for replanted cycads in zuurberg section, addo. it could thus be concluded that of the six cycad individuals replanted at site 2, one was in good condition bearing leaves (17%), two were inconclusive regarding their condition (33%), and three were dead (50%). site 3 eight cycad individuals were replanted in february 2015 at site 3. when monitored in 2018, three adult individuals were recorded to be in a good condition bearing leaves (more than four). one individual was previously recorded with leaves (no leaves noted in 2018) but even though it was still standing, scars and damage caused by baboons were clearly evident at the time of the survey. four individuals had died leaving either no trace, or the remains of stems were scattered in the veld (table 3). it could thus be concluded that of the eight cycad individuals replanted at site 3, three appeared healthy bearing leaves (38%), one’s condition could not be determined (dormant) (12%), and four had died (50%). site 4 eight cycad individuals were replanted in february 2015 at site 4. when monitored in 2018, one adult individual was recorded with more than four leaves. the particular individual, however, did not seem to be in good condition, with ants noted on the stem and leaves. one individual, was dormant (standing without leaves) but the stem looked healthy and intact. two individuals were uprooted and might still grow from this horizontal position. four individuals were recorded dead and either nothing could be found or loose remains of stems were scattered in the veld (table 3). the baboons impacted on these cycads more than at the other three sites, because it appears to have been planted on their daily foraging route. it could thus be concluded that of the eight cycad individuals replanted at site 4, one of them was healthy with leaves (12% of the planted cycad individuals), for three the condition could not be determined (38%), and four were dead (50%). site 5 eight cycad individuals were replanted in february 2015 at site 5. the monitored result was one adult individual with five leaves recorded, however, this individual did not look healthy. one individual was recorded as dormant (still standing but without leaves) and more than 30% of the stem had been damaged. one individual was uprooted but might still grow from its current position lying on the soil surface. five cycad individuals were recorded as dead and either nothing could be found or loose remains of stems were scattered in the veld (table 3). it could thus be concluded that of the eight cycad individuals replanted at site 5, one was healthy with leaves (12%), for two the condition could not be determined (25%), and five were dead (63%). final results of the march 2018 survey indicated that of the 48 cycad individuals (including broken-off additional suckers), 27% were in a good healthy condition and bearing leaves. twenty-seven percent were standing without leaves but still had the potential to grow (dormant). currently, 46% of the planted cycads have died and have either been removed or loose remains of stems have been found (table 4). table 4: the monitoring results of all the planted cycads in addo elephant national park (march 2018). some localities and sites showed more success than others; for example, 43% of the cycad individuals planted in february 2015 at the aenp garden looked healthy with leaves, whilst at sites 4 and 5 in the zuurberg section, only 12% of the replanted individuals had leaves (tables 1 & 2). an interesting observation was that fire actually stimulated five cycads, which had been standing for 3 years without leaves, to produce new leaves. none of these replanted cycads were killed by the fire. from the zuurberg locality, it became clear that baboons caused further injury to cycads that had existing stem damage. it was noted that the baboons made no impact on the natural cycads occurring in the same area. baboons and other wildlife are part of the natural ecosystem where these cycads occur. both cousins and witkowski (2017) and nowak and lee (2011) observed that the feeding and/or spreading of cycad seeds by baboons, monkeys, elephants, birds, fruit-eating bats, bush pigs and rodents is normal and characteristic of the local floral and faunal ecology. interesting to note is that, according to nowak and lee (2011), monkeys have developed specialised foraging techniques that enable them to reach the newly-emerging soft leaves, which contain less of the toxin hydrogen cyanide, which is found in older leaves. donaldson’s (1995) historical records show that baboon damage to some female cones is insignificant (< 15%), thus ensuring the production of more than enough seeds and seedlings. conclusion undisturbed cycads are very slow growers, usually only producing one set of new leaves annually, but intervals between leaf production events can sometimes span several years (cousins & witkowski 2017). the success or failure of this recovery project will only be visible after another 3 years. in the life cycle of these long-living cycads, 3 years is short, however the results that were recorded were remarkably positive and continued monitoring is imperative to record the long-term success of these individual plants after their re-establishment into aenp. according to dr. john donaldson (south african national biodiversity institute [sanbi] cycad specialist – pers. comm., 18 feburary 2015) and mr. werner voigt (kirstenbosch national botanical garden – pers. comm., 18 feburary 2015), a survival rate of more than 20% (or nine) of cycad individuals will indicate that this project was successful. overall, 13 individuals were classified as healthy and produced leaves, whilst 13 individuals were standing or lying on the soil surface (dormant) and could still grow. unfortunately, 22 cycads died. the preliminary results of this study concur with donaldson’s type 2 (persister/reproducer) classification of these two cycad species (donaldson 1995). the results also indicate that the cycad individuals at the aenp gardens showed the best results. this is partially to be expected – apart from receiving more water in the garden, the climate was also less extreme than in the higher lying zuurberg mountain range. although these now garden cycads are not back in the natural habitat, they will hopefully produce seeds and suckers that could be used for re-introduction into the natural habitat. it is important to note that although we are dealing here with re-established cycads in the gardens and into the wild, the main aim should always be to relocate cycads into their natural environment. these cycads play an important role in the ecosystems’ natural patterns and processes, with many organisms dependent on them (cousins & witkowski 2017; oberprieler 1999). acknowledgements without the dedication, passion, friendliness and hard work of so many people, this project would not have been possible. the previous addo elephant national park manager, mr mzwandile mjadu and the previous nature conservation manager, mr john adendorff, and zelna silcock (now breytenbach) are thanked for immediate actions related to the recovery of these cycads. apart from assisting with the treatment and replanting of the cycads, mr hans von gordon of the addo (port elizabeth) honorary rangers as well as the national executive committee (nec) of the honorary rangers, they were also financially involved with this project. mr louis and queen olivier also contributed material, time as well as expertise. appreciation is extended to dr john donaldson and mr werner voigt of sanbi, who gave valuable advice on the whole processes. roxanne erusan and zelna breytenbach assisted with monitoring and keeping me updated. the author would also like to thank leslie adriaanse of the university of south africa library for assisting with obtaining some difficult references. competing interests the author declares that no competing interests exist. author’s contributions i declare that i am the sole author of this research article. funding information this research received no specific grant from any funding agency in the public, commercial, or not-for-profit sectors. data availability statement it is a short note, data will remain with the author. disclaimer the views and opinions expressed in this article are those of the author’s and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references bezuidenhout, h., 2015, recovery procedure of unearthed cycad trees in the greater addo elephant national park and adjacent areas april 2015, internal report 21/2015, scientific services, kimberley, south african national parks. bezuidenhout, h., freitag-ronaldson, s., ferreira, s. & hofmeyr, m., 2017, identification of proposed sustainability thresholds for key species: encephalartos species (cycads) in sanparks, internal report 05/2017, scientific services, kimberley south african national parks. bezuidenhout, h. & hofmeyr, m., 2016, memo on broad conceptual map on cycads (encephalartos species) in sanparks, internal report 02/2016, scientific services, kimberley conservation services division, south african national park. boyd, w.m., 1995. ‘the translocation and re-establishment of priority encephalartos species in the transvaal’, in the proceedings of the third international conference of cycad biology held in pretoria, pp. 423–434. isbn 9780620192286. cousins, s.r. & witkowski, e.t.f., 2017, ‘african cycad ecology, ethnobotany and conservation: a synthesis’, the botanical review 83, 152–194. https://doi.org/10.1007/s12229-017-9183-4 department of environmental affairs, 2015, national strategy and action plan for the management of cycads, publication of environmental affairs, department: environmental affairs, pretoria. donaldson, j.s., 1995, ‘understanding cycad life histories: an essential basis for successful conservation’, in j.s. donaldson (ed.), cycad society of south africa, cycad conservation in south africa: issues, priorities and actions, pp. 8–13, cycad society of south africa, claremont, isbn 9780620190381. donaldson, j.s. (ed.), 2003, cycads. status survey and conservation action plan, international union for conservation of nature/ssc cycad specialist group, international union for conservation of nature, gland, ix + 86 pp. giddy, c., 1974, cycads of south africa, purnell and sons (s.a.) (pty) ltd, cape town, 122 pp. isbn 360002749. nowak, k. & lee, p.c., 2011, ‘consumption of cycads encephalartos hildebrandtii by zanzibar red colobus procolobus kirkii’, journal of east african natural history 100, 123–131. https://doi.org/10.2982/028.100.0108 oberprieler, r.g., 1999, ‘systematics and evolution of the cycad-associated weevil genus apinotropis jordan (coleoptera: anthribidae)’, african entomology 7(1), 1–33. singh, k., 2012, ‘population dynamics of the zuurberg cycad and the predicted impact of climate change’, msc. dissertation. nelson mandela metropolitan university, port elizabeth. singh, k. & campbell, e.e., 2011, the zuurberg cycads and predicted impacts of climate change, research proposal for addo elephant national park by the department of botany, nelson mandela metropolitan university, port elizabeth. staatskoerant, 2017, ‘national environmental management: biodiversity act, 2004 (act no. 10 of 2004). biodiversity management plan for eleven critically endangered and four endangered encephalartos cycad species’, staatskoerant (40815), 275–349. van der walt, k., 2014, biodiversity management plan for 11 critically endangered and 4 endangered encephalartos species, report: strategic environmental focus (s.e.f.), lynnwood ridge or sanbi, silverton, pretoria. filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 dippenaar.qxd a check list of the spiders of the kruger national park, south africa (arachnida: araneae) anna s. dippenaar-schoeman and a. leroy dippenaar-schoeman, anna s. and a. leroy. 2003. a check list of the spiders of the kruger national park, south africa (arachnida: araneae).koedoe 46(1): 91–100. pretoria. issn 0075-6458. as part of the south african national survey of arachnida (sansa), projects are underway to determine the biodiversity of arachnids present in protected areas in south africa. spiders have been collected over a period of 16 years from the kruger national park, south africa. a check list is provided consisting of 152 species, 116 genera and 40 families. this represents about 7.6 % of the total known south african spider fauna. of the 152 species, 103 are new records for the park. the ground dwelling spiders comprise 58 species from 25 families. of these, 21 % are web dwellers and 62 % free living, while 17 % live in burrows. from the plant layer, 94 species have been collected of which 53 % were web builders and 47 % free living wandering spiders. key words: araneae, biodiversity, check list, kruger national park, south africa, spiders. anna s. dippenaar-schoeman, arc-plant protection research institute, private bag x134, pretoria 0001, republic of south africa; astri leroy, p.o. box 2810, wilropark, 1731, republic of south africa. issn 0075-6458 91 koedoe 46/1 (2003) introduction conservation biologists are starting to recognise the importance of the invertebrate component in the functioning of healthy ecosystems. therefore, any approach to conservation needs to take into account the composition of these invertebrate fauna. inventories, with resulting check lists, provide valuable baseline information on species present and are the first step toward a better understanding of the fauna of the area. in 1997, the south african national survey of arachnida (sansa) was initiated (dippenaar-schoeman & craemer 2000) with its main aim to make an inventory of the arachnid diversity of south africa. one of the projects of sansa is to compile an inventory of spiders presently conserved in parks and reserves in south africa. although spiders constitute an abundant and highly diverse group of invertebrate animals, little is still known about their diversity even in conserved areas. from south africa check lists exist for the spider fauna of the mountain zebra national (dippenaar-schoeman 1988), roodeplaat dam nature reserve (dippenaar-schoeman et al. 1989), karoo national park (dippenaar-schoeman et al. 1999), makelali nature reserve in the limpopo province (whitmore et al. 2001) and the western soutpansberg (foord 2002). the aim of this study was to compile the first check list of the spider species of the kruger national park (knp) and to determine the percentage of species protected. this was not an intensive survey but reflects on collecting done in the park over a period of 16 years. although this survey may not reflect the true diversity and species richness of the area, it does give an indication of species present. material and methods study area the kruger national park (knp) is situated in the lowveld region of the mpumalanga and limpopo provinces of south africa. the park is 350 km long dippenaar.qxd 2005/12/09 10:26 page 91 from north to south with a surface area of 1 948 528 ha. the climate is subtropical, with the annual rainfall varying between 700 mm in the south and 400 mm in the north. it falls within the savanna biome and the vegetation includes mixed bushwillow woodlands, mountain bushveld, thorn thicket, thorn veld, sourveld, scrubveld, sandveld and riverine forest. collecting methods sporadic collecting was mainly undertaken from 1985 to 2001. spiders were sampled by hand (ground and plant search, turning rocks and sifting of leaf litter) or using a sweepnet or beating tray for grass and low shrubs. the new records were identified by the first author and are housed in the national collection of arachnida (nca) at the arc-plant protection research institute in pretoria. the lack of taxonomic research in southern africa within certain families made the identification of some genera to species level impossible. in some families, only immature specimens were collected, hampering identification to species level. this means that, in both instances, only generic names are included in the check list. the species list includes published koedoe 46/1 (2003) 92 issn 0075-6458 table 1 guild classification of spiders collected in the kruger national park. guilds abbreviation guild explanation wandering spiders (w) ground wandering spiders (gw) free living fgw free-living spiders running on the soil surface when active burrow living bgw living in burrows plant wandering spiders (pw) free living fpw free-living spiders running on the plant surface when active web-building spiders (wb) orb-web owb orb-webs consisting of a frame with mooring and bridge lines that anchors the web and radial signal threads arranged like the ribs of a umbrella converging onto the centre of the web with circular spiral threads funnel-web fwb sheet-webs made over soil surface with a funnel-shaped retreat gumfoot-web gwb three-dimensional webs consisting of a central area with or without a retreat. the upper part comprises mooring, signal and catch threads and a lower part with mooring and catch threads. the lower catch threads studded with sticky droplets are attached to the substrate retreat-web rwb silk threads used to catch prey radiating from retreat sheet-web shwb sheet-webs which, usually consists of an upper sheet with mooring, signal and catch threads space-web spwb space-webs which, fill open space and are usually attached with mooring threads to different substrates. dippenaar.qxd 2005/12/09 10:26 page 92 records of spiders previously recorded from the kruger national park. guilds a guild is a group of species that potentially compete for jointly exploited limited resources (polis & mccormick 1986). because most spiders live in a defined environment with limitations set by both physical conditions and biological factors (foelix 1982), species can be grouped into guilds based on available information on their habitat preferences and predatory methods. for the present study two main guilds were recognised, namely wandering spiders (w) and web builders (wb), with further subdivisions based on micro-habitat and general behaviour (table 1). results and discussion numbers present forty families represented by 116 genera and 152 species are presently known from the knp (table 2). of the 152 species listed 103 are new records for the knp (table 2 & 3). the orb-web spiders of the family araneidae are the most diverse and represented by 23 species, followed by the crab spiders (thomisidae) with 15, the lynx spiders (oxyopidae) with 9 and the wolf spiders (lycosidae) and jumping spiders (salticidae) with 8 species each. fifteen families are represented by a single species. a total of 91 species are free living wandering spiders (59 %) while 61 species (41 %) build webs. issn 0075-6458 93 koedoe 46/1 (2003) family genera species nr family genera species nr araneidae 15 23 15 oxyopidae 3 9 4 archaeidae 1 1 0 palpimanidae 1 1 1 barychelidae 1 1 1 philodromidae 3 4 3 caponiidae 1 1 1 pholcidae 3 3 3 corinnidae 2 2 2 phyxelididae 1 1 0 ctenidae 1 1 1 pisauridae 5 5 3 cyrtaucheniidae 1 1 1 prodidomidae 1 2 2 deinopidae 1 1 1 salticidae 8 8 7 dictynidae 2 2 1 scytodidae 1 1 1 dipluridae 1 1 0 segestriidae 1 1 1 eresidae 3 6 2 selenopidae 2 6 1 gnaphosidae 6 6 1 sicariidae 2 2 2 hersiliidae 1 2 0 sparassidae 3 3 2 idiopidae 1 1 1 tetragnathidae 5 7 6 linyphiidae 2 2 1 theraphosidae 4 5 1 liocranidae 1 1 1 theridiidae 5 5 5 lycosidae 8 8 8 thomisidae 9 15 14 mimetidae 1 1 1 uloboridae 1 2 2 miturgidae 2 4 2 zodariidae 4 4 2 oecobiidae 2 2 2 oonopidae 1 1 1 tot: 40 116 152 103 table 2 spider families collected at the kruger national park indicating the number of species and genera in each family and the number of species that are new records (nr) dippenaar.qxd 2005/12/09 10:26 page 93 ground wandering spiders twenty-five families represented by 58 species are associated with the ground layer, 10 species live in burrows made in the ground, 36 species are free living and 12 are web dwellers. web-building spiders twelve species construct their webs on or close to the soil surface. the following web types were found: funnel-webs of the dipluridae (allothele malawi coyle, 1984); retreat-webs of the eresidae (dresserus colsoni tucker, 1920), hersiliidae (tama sp.), oecobiidae (2 spp.) and the phyxelididae (xevioso orthomeles griswold, 1990); space-web of the pholcidae (3 spp.) funnelweb of the lycosid (hippasa australis lawrence, 1927) and the gumfoot-webs of the theridiids (latrodectus geometricus c.l. koch, 1841 and steatoda capensis hann, 1990). free living ground spiders thirty-six species (62 %) are free-living wanderers represented by 15 families. the more diverse families are the wolf spiders (lycosidae) with 8 species, the ground spiders (gnaphosidae) and the flatties (selenopidae) both with 6 species each. burrow living spiders eight of the burrow dwellers found in the park belong to the suborder mygalomorphae. they are represented by: two trapdoor spider families, cyrtaucheniidae (ancylotrypa barbertoni (hewitt, 1913)) and idiopidae (idiops castaneus hewitt, 1913); five baboon spider species of the family theraphosidae; and one lesser baboon spider of the family barychelidae. of the mygalomorphs, the international union for the conservation of nature (iucn) (de wet & schoonbee 1991) finds that it is mainly the larger theraphosidae that are considered commercially threathened. four of the theraphosid species, added to schedule vii of the transvaal provincial nature conservation ordinance of 1983 as protected invertebrate animals, are found in the park (dippenaarschoeman 2002a; gallon 2002), viz., ceratogyrus bechuanicus purcell, 1902, harpactira gigas pocock, 1898, augacephalus breyeri (hewitt, 1919) and a. junodi (simon, 1904). plant wandering spiders web-building spiders from the field layer 52 % of the species (50) collected build webs. the orb-web spiders were the most diverse and represented by three families namely the araneidae (23 spp.), tetragnathidae (7 spp.) and uloboridae (2 spp.). members of the genera argiope, afracantha and gasteracantha of the araneidae and leucauge, nephila, nephilengys and tetragnatha of the tetragnathidae construct large orb-webs between plants and are seen in their webs during the day. the tropical tent-web spider cyrtophora citricola (forskål, 1775) makes horizontal adapted orb-webs in plants like aloes. most of the other orb-web species are nocturnal, e.g., the bark spider caerostris, and remove their webs early each morning. the retreat webs of the dictynidae, eresidae and segestriidae are made on different parts of the plants. two species of the genus stegodyphus, s. dumicola pocock, 1898 and s. mimosarum pavesi, 1883 are communal nest spiders found in large nests attached to the trees. the other web spiders construct gumfoot-webs (theridiidae) and sheet-webs (pisauridae and linyphiidae). the large funnel-webs made vertically next to trees with the retreat part usually concealed in the plants or in abandoned animal holes are made by the large pisaurid euprosthenops australis simon, 1898. free living plant spiders ten families from 44 species are found on plants, with members of four families associated with bark, eight with grass, and four with leaves. the long spinnered bark spider hersilia sericea pocock, 1898 is a permanent inhabitant on bark, while members of the philodromidae, salticidae and thomisidae are found on bark as well as koedoe 46/1 (2003) 94 issn 0075-6458 dippenaar.qxd 2005/12/09 10:26 page 94 issn 0075-6458 95 koedoe 46/1 (2003) table 3 check list of the spiders of the kruger national park family/genus/species guild record 1. family araneidae afracantha camerunensis (thorell, 1899) owb emerit 1973 araneus apricus (karsch, 1884) owb nr argiope australis (walckenaer, 1805) owb bjørn 1997 a. flavipalpis (lucas, 1858) owb bjørn 1997 a. trifasciata (forskål, 1775) owb nr caerostris sexcuspidata (fabricius, 1793) owb nr cyclosa insulana (costa, 1834) owb nr cyphalonotus larvatus (simon, 1881) owb nr cyrtophora citricola (forskål, 1775) owb nr gasteracantha milvoides butler, 1873 owb nr g. sanguinolenta c.l. koch, 1844 owb nr g. versicolor (walckenaer,1842) owb emerit 1973; benoit & emerit 1975 hypsacantha crucimaculata (dahl, 1914) owb nr isoxya stuhlmanni (bösenberg & lenz,1895) owb nr i. tabulata (thorell, 1859) owb nr kilima decens (blackwall, 1866) owb nr nemoscolus vigintipunctatus simon, 1897 owb nr neoscona blondeli (simon, 1885) owb grasshoff 1986 n. moreli (vinson, 1863) owb grasshoff 1986 n. rufipalpis (lucas, 1858) owb nr n. subfusca (c.l. koch, 1837) owb nr poltys furcifer simon, 1881 owb nr singa albodorsata kauri, 1950 owb kauri 1950 2. family archaeidae afrarchaea bergae lotz, 1996 fgw lotz 1996a 3. family barychelidae brachionopus pretoriae purcell, 1904 bgw nr 4. family caponiidae caponia natalensis (o.p.-cambridge, 1874) fgw nr 5. family corinnidae castianeira sp. fgw nr merenius alberti lessert, 1923 fgw nr 6. family ctenidae ctenus transvaalensis benoit, 1981 fgw nr 7. family cyrtaucheniidae ancylotrypa barbertoni (hewitt, 1913) bgw nr 8. family deinopidae menneus camelus pocock, 1902o wb nr 9. family dictynidae mashimo leleupi lehtinen, 1967 rwb nr archaeodictyna ulova griswold & m-griswold, 1987 rwb griswold & m-griswold 1987 10. family dipluridae allothele malawi coyle, 1984 fwb coyle 1984 11. family eresidae dresserus colsoni tucker, 1920 rwb nr dippenaar.qxd 2005/12/09 10:26 page 95 koedoe 46/1 (2003) 96 issn 0075-6458 gandanameno purcelli tucker, 1920 rwb nr stegodyphus africanus (blackwall, 1866) rwb kraus & kraus 1988 s. dumicola pocock, 1898 rwb kraus & kraus 1988 s. mimosarum pavesi, 1883 rwb kraus & kraus 1988 s. sabulosus tullgren, 1910 rwb kraus & kraus 1988 12. family gnaphosidae aphantaulax inornata tucker, 1923 fgw tucker 1923 asemesthes purcelli tucker, 1923 fgw tucker 1923 camillina maun platnick & murphy, 1987 fgw platnick & murphy 1987 megamyrmaekion transvaalense tucker, 1923 fgw tucker 1923 scotophaeus marleyi tucker, 1923 fgw tucker 1923 zelotes tuckeri roewer, 1951 fgw nr 13. family hersiliidae hersilia sericea pocock, 1898 fpw benoit 1967 hersilida sp. (immature) rwb nr 14. family idiopidae idiops castaneus hewitt, 1913 bgw nr 15. family linyphiidae tybaertiella krugeri (simon 1894) shwb simon 1894 meioneta habra locket, 1968 shwb nr 16. family liocranidae rhaeboctesis trinotatus tucker, 1920 fgw nr 17. family lycosidae arctosa transvaalana roewer, 1960 fgw nr evippomma squamulatum (simon, 1898) fgw nr hippasa australis lawrence, 1927 fwb nr hogna transvaalica (simon, 1898) fgw nr lycosa sp. bgw nr pardosa crassipalpis purcell, 1903 fgw nr pirata sp. fgw nr proevippa albiventris (simon, 1898) fgw nr 18. family mimetidae mimetus natalensis lawrence, 1938 fpw nr 19. family miturgidae cheiracanthium africanum lessert, 1921 fpw lotz 1996b* c. furculatum karsch, 1879 fpw lotz 1996b* cheiramiona paradisus lotz, 2002 fpw lotz 2002 c. krugerensis lotz, 2002 fpw lotz 2002 20. family oecobiidae. oecobius navus blackwall, 1859 rwb nr uroecobius sp. (immature) rwb nr 21. family oonopidae opopaea speciosa (lawrence, 1952). fgw nr 22. familyoxyopidae hamataliwa kulczynskii (lessert, 1915) fpw nr oxyopes jacksoni lessert, 1915 fpw nr o. longispinosus lawrence,1938 fpw nr o. pallidecoloratus strand, 1906 fpw nr peucetia pulchra(blackwall, 1865) fpw van niekerk & dippenaarschoeman1994 p. madalenae van niekerk & dippenaar-schoeman, 1994 fpw van niekerk & dippenaarschoeman1994 p. striata karsch, 1878 fpw van niekerk & dippenaardippenaar.qxd 2005/12/09 10:26 page 96 issn 0075-6458 97 koedoe 46/1 (2003) schoeman 1994 p. transvaalica simon, 1896 fpw van niekerk & dippenaar schoeman 1994 p. viridis (blackwall, 1858) fpw van niekerk & dippenaarschoeman1994 23.family palpimanidae palpimanus transvaalicus simon, 1893 fgw nr 24. family philodromidae philodromus rufus walckenaer, 1825 fpw nr thanatus atlanticus berland, 1936 fpw nr t. vulgaris simon, 1870 fpw nr tibellus sunetae van den berg &dippenaarschoeman fpw van den berg & dippenaar1994 schoeman, 1994 25. family pholcidae leptopholcus sp. (immature) spwb nr pholcus leptopholcicus strand, 1909 spwb nr smeringopus natalensis lawrence, 1947 spwb nr 26. family phyxelididae xevioso orthomeles griswold, 1990 rwb griswold 1990 27. family pisauridae cispius problematicus blandin, 1978 fpw sierwald 1997 euprosthenops australis simon, 1898 fwb nr perenethis symmetrica (lawrence, 1927) fpw sierwald 1997 rothus purpurissatus simon, 1898 shwb nr thalassius rossi pocock, 1902 fgw nr 28. family prodidomidae theuma foveolata tucker, 1923 fgw nr t. fusca purcell, 1907 fgw nr 29. family salticidae baryphas ahenus simon, 1902 fpw nr brancus bevisi lessert, 1925 fpw nr heliophanus transvaalicus simon, 1901 fpw nr hyllus brevitarsis simon, 1902 fpw nr myrmarachne laurentina bacelar, 1953 fpw nr phlegra albostriata simon, 1901 fgw nr portia schultzi karsch, 1878 fpw wanless 1978 thyene coccineovittata (simon, 1885) fpw nr 30. family scytodidae scytodes fusca walckenaer, 1837 fgw nr 31. family segestriidae ariadna sp. (immature) rwb nr 32.family selenopidae anyphops rubicundus (lawrence, 1940) fgw lawrence 1940 a. silvicolellus (strand, 1913) fgw corronca 1998 selenops krugeri lawrence, 1940 fgw corronca 2000 s. ovambicus lawrence, 1940 fgw corronca 2000 s. radiatus latreille, 1819 fgw corronca 2000 s. tenebrosus lawrence, 1940 fgw nr 33. family sicariidae loxosceles spiniceps lawrence, 1952* fgw nr sicarius oweni newlands, 1986 * fgw nr dippenaar.qxd 2005/12/09 10:26 page 97 koedoe 46/1 (2003) 98 issn 0075-6458 34. family sparassidae olios correvoni lessert, 1921 fpw nr palystes superciliosus l. koch, 1875 fpw croeser 1996 pseudomicrommata longipes bösenberg & lenz, 1895 fpw nr 35. family tetragnathidae diphya simoni kauri, 1950 owb kauri 1950 leucauge decorata (blackwall, 1864) owb nr nephila pilipes (fabricius, 1793) owb nr n. senegalensis annulata (thorell, 1859) owb nr nephilengys cruentata (fabricius, 1775) owb nr tetragnatha boydi o.p.-cambridge, 1898 owb nr t. subsquamata okuma, 1985 owb okuma & dippenaar-schoeman 1988 36. family theraphosidae augacephalus breyeri (hewitt, 1919) bgw gallon 2002 a. junodi (simon, 1904) bgw gallon 2002 ceratogyrus bechuanicus purcell, 1902 bgw smith 1990 harpactira gigas pocock, 1898 bgw nr idiothele nigrofulva (pocock, 1898) bgw gallon 2002 37. family theridiidae argyrodes convivans (lawrence, 1937) gwb nr latrodectus geometricus c.l. koch, 1841 gwb nr phoroncidia eburnea (simon,1895) gwb nr steatoda capensis hann, 1990 gwb nr theridion purcelli o.p.-cambridge, 1940 gwb nr 38. family thomisidae misumenops rubrodecoratus millot, 1942 fpw nr monaeses pustulosus pavesi, 1895 fpw dippenaar-schoeman 1984 m. quadrituberculatus lawrence, 1927 fpw nr pactactes compactus lawrence, 1947 fpw nr runcinia aethiops (simon, 1901) fpw nr r. flavida (simon, 1881) fpw nr r. johnstoni lessert, 1919 fpw nr simorcus cotti lessert, 1936 fpw nr thomisops pupa karsch, 1879 fpw nr thomisus daradioides simon, 1890 fpw nr t. granulatus karsch, 1880 fpw nr t. kalaharinus lawrence, 1936 fpw nr t. spiculosus pocock, 1901 fpw nr tmarus africanus lessert, 1919 fpw nr xysticus sp. fpw nr 39. family uloboridae uloborus planipedius simon, 1896 owb nr u. plumipes lucas, 1846 owb nr 40. family zodariidae capheris transvaalica hewitt, 1915 bgw nr chariobas sp. fpw nr diores rectus jocqué‚ 1990 fgw jocqué 1990 hermippus tenebrosus jocqué, 1986 fgw jocqué 1986 * from unpublished msc and phd theses. bgw = burrow ground dwellers; fwb = funnel web; fgw = free living ground wanderer; fpw = free living plant wanderers; gwb = gumfoot-web; owb = orb-web; nr = new collecting record; shwb = sheetweb; spwb = space web. dippenaar.qxd 2005/12/09 10:26 page 98 other parts of the plant. the knp, as part of the savanna biome, is represented by about 32 grass-living species. many of the grass dwellers are well camouflaged with elongated bodies, e.g., runcinia spp. (thomisidae), tibellus sunetae van den berg & dippenaarschoeman, 1994 (philodromidae) and pseudomicrommata longipes bösenberg & lenz, 1895 (sparassidae), while others with their green or straw-coloured bodies blend in with the grass, e.g., members of the families oxyopidae and thomisidae. conclusion preliminary investigations into the biodiversity of invertebrate fauna in south africa have highlighted the lack of baseline information on the ecology and diversity of most arachnid groups (dippenaar-schoeman 2002b). this survey of the knp forms part of the south african national survey of arachnida (sansa) and data gathered will be used in the savanna biome project, mpumalanga biobase programme and arachnida in conserved area projects. of the 2000 spiders presently known from south africa (dippenaar-schoeman 2002b), the 152 species of knp represent about 7.6 % of the total spider fauna, with 103 species being new distribution records. although this paper reports on sporadic collecting and probably represents only a portion of the spider fauna present, we hope this information will stimulate further research on this group of animals in the knp. future projects for the park include a key to the spiders and maps showing their distribution patterns. acknowledgements we wish to thank the warden and officials of the kruger national park for their friendliness and assistance. thanks are due to dr. m.k.p. smith meyer and ms. a. van den berg, both of the arc-plant protection research institute for reading the manuscript and marie de jager for additional spiders collected. references benoit, p.l.g. 1967. révision des espèces africaines du genre hersilia sav. et aud. (aran.-hersiliidae). revue de zoologie et de botanique africaines 76: 1-36. benoit, p.l.g. & m. emerit. 1975. mise à jour des connaissances concernant les araneidae-gasteracanthinae africains. revue de zoologie africaine 89: 321-336. bjørn, p. p. 1997. a taxonomic revision of the african part of the orb-weaving genus argiope (araneae: araneidae). entomologica scandinavica 28: 199-239. corronca, j. a.1998. uevos registros de anyphops benoit (araneae, selenopidae) y comentarios sobre su distribución. biogeographica 74: 177182. corronca, j. a. 2000. distribucion y nuevos registros de selenops latreille (araneae, selenopidae) en la región afrotropical. biogeographica 76: 89-94. coyle, e.a. 1984. a revision of the african mygalomorph spider genus allothele (araneae, dipluridae). american museum novitates 2794: 1-20. croeser, p.m.c. 1996. a revision of the african huntsman spider genus palystes l. koch, 1875 (araneae: heteropodidae). annals of the natal museum 37: 1-122. de wet, j.i. & h.j. schoonbee. 1991. the occurrence and conservation status of ceratogyrus bechuanicus and c. brachycephalus in the transvaal, south africa. koedoe 34: 69-75. dippenaar-schoeman, a.s. 1984. the crab-spiders of southern africa (araneae: thomisidae). 4. the genus monaeses thorell, 1869. phytophylactica 16: 101-116. dippenaar-schoeman, a.s. 1988. annotated check list of the spiders (araneae) of the mountain zebra national park. koedoe 31: 151-160. dippenaar-schoeman, a. s. 2002a. baboon and trapdoor spiders of southern africa: an identification manual. pretoria: agricultural research council. (plant protection research institute handbook 13) dippenaar-schoeman, a.s. 2002b. status of south african arachnida fauna. pp: 70-81. in: status of south african species. proceedings of symposium organized by the endangered wildlife trust (ewt) of south africa. dippenaar-schoeman, a.s. & c. craemer. 2000. the south african national survey of arachnida (sansa). plant protection news 56: 11-12 dippenaar-schoeman, a.s., a. leroy, m. de jager & a. van den berg. 1999. a checklist of the spider fauna of the karoo national park, south africa (arachnida: araneae). koedoe 42: 31-42. issn 0075-6458 99 koedoe 46/1 (2003) dippenaar.qxd 2005/12/09 10:26 page 99 dippenaar-schoeman, a.s., a.m. van den berg & a. van den berg. 1989. species composition and relative seasonal abundance of spiders from the field and tree layer of the roodeplaat dam nature reserve. phytophylactica 22: 443-447. emerit, m. 1973. contribution à la connaissance des araneidae gasteracanthinae du sud-est africain: les gastéracanthes du natal museum. annals of the natal museum 21: 675-695. foelix, r.f. 1982. biology of spiders. harvard university press, cambridge, massachusetts. foord, s.h., a.s. dipppenaar-schoeman & m. van der merwe. 2002. a check list of the spider fauna of the western soutpansberg, south africa (arachnida: araneae). koedoe 45: 35-43 gallon, r.c. 2002. revision of the african genera pterinochilus and eucratoscelus (araneae, theraphosidae, harpactirinae) with description of two new genera. bulletin of the british arachnological society 12: 201-232. grasshoff, m. 1986. die radnetzspinnen-gattung neoscona in afrika (arachnida: araneae). annales du musée royal de l'afrique centrale 250: 1-123. griswold, c.e. 1990. a revision and phylogenetic analysis of the spider subfamily phyxelidinae (araneae, amaurobiidae). bulletin of the american museum of natural history 196: 1206. griswold, c.e. & t.c. meikle-griswold. 1987. archaeodictyna ulova, new species (araneae: dictynidae), a remarkable kleptoparasite of group-living eresid spiders (stegodyphus spp., araneidae: eresidae). american museum novitates 2897: 111. jocqué, r. 1986. a revision of the genus hermippus (araneae, zodariidae). journal of natural history 20: 7-22. jocqué, r. 1990. a revision of the afrotropical genus diores (araneae, zodariidae). annales du museé royal de l'afrique centrale 260: 1-81. kauri, h. 1950. on some south african spiders and harvest spiders. kunglige fysiografiska sällskapets 1 lund förhandlingar 20: 64-79. kraus, o. & m. kraus. 1988. the genus stegodyphus (arachnida, araneae). sibling species, species groups, and parallel origin of social living. verhandlungen der naturwissenschaftlichen verein in hamburg 30: 151-254. lawrence, r.f. 1940. the genus selenops (araneae) in south africa. annals of the south african museum 32: 555-608. lotz, l.n. 1996a. afrotropical archaeidae (araneae): 1. new species of afrachaea with notes on afrachaea godfreyi (hewitt, 1919). navorsinge van die nasionale museum, bloemfontein 12: 141-160. lotz, l.n. 1996b. the genus cheiracanthium (araneae: clubionidae) in southern africa. msc thesis, university of the orange free state, bloemfontein. lotz, l.n. 2002. a revision of the afrotropical genus cheiramiona (araneae: miturgidae: eutichurinae). annales du museé royal de l'afrique centrale 290: 39-86. okuma, c. & a.s. dippenaar-schoeman. 1988. tetragnatha (araneae: tetragnathidae) in the collection of the plant protection research institute, pretoria with description of a new species. phytophylactica 20: 219-232. polis, g.a. & s.j. mccormick. 1986. scorpions, spiders and solpugids: predation and competition among distantly related taxa. oecologia 71: 111-116. platnick, n.i. & j.a. murphy. 1987. studies on the malagasy spiders. 3. the zelotine gnaphosidae (araneae, gnaphosoidea) with a review of the genus camillina. american museum novitates 2874: 1-33. sierwald, p. 1997. phylogenetic analysis of pisaurine nursery web spiders, with revisions of tetragonophthalma and perenethis (araneae, lycosoidea, pisauridae). the journal of arachnology 25: 361-407. simon, e. 1894. histoire naturelle des araignées 1(3), paris 1894: 489-760. smith, a.m. 1990. a revision of the theraphosidae family from africa and the middle east. uk: fitzgerald publishing. tucker, r.w.e. 1923. the drassidae of south africa. annals of the south african museum 19: 251-438. van den berg, a. & a.s. dippenaar-schoeman. 1994. a revision of the afrotropical species of the genus tibellus simon (araneae: philodromidae). koedoe 37: 67-114. van niekerk, p. & a.s. dippenaar-schoeman. 1994. a revision of the afrotropical species of peucetia (araneae: oxyopidae). entomology memoir, department of agriculture, south africa 89: 1-50. wanless, e.r. 1978. a revision of the spider genus portia (araneae, salticidae). bulletin of the british museum (natural history), zoology 34: 83-124. whitmore, c., r. slotow, t.e. crouch & a.s. dippenaar-schoeman. 2001. checklist of spiders (araneae) from savanna ecosystem, northern province, south africa: including a new family record. durban museum novitates 26: 10-19. koedoe 46/1 (2003) 100 issn 0075-6458 dippenaar.qxd 2005/12/09 10:26 page 100 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <feff004700650062007200750069006b002000640065007a006500200069006e007300740065006c006c0069006e00670065006e0020006f006d0020005000440046002d0064006f00630075006d0065006e00740065006e0020007400650020006d0061006b0065006e0020006d00650074002000650065006e00200068006f00670065002000610066006200650065006c00640069006e00670073007200650073006f006c007500740069006500200076006f006f0072002000610066006400720075006b006b0065006e0020006d0065007400200068006f006700650020006b00770061006c0069007400650069007400200069006e002000650065006e002000700072006500700072006500730073002d006f006d0067006500760069006e0067002e0020004400650020005000440046002d0064006f00630075006d0065006e00740065006e0020006b0075006e006e0065006e00200077006f007200640065006e002000670065006f00700065006e00640020006d006500740020004100630072006f00620061007400200065006e002000520065006100640065007200200035002e003000200065006e00200068006f006700650072002e002000420069006a002000640065007a006500200069006e007300740065006c006c0069006e00670020006d006f006500740065006e00200066006f006e007400730020007a0069006a006e00200069006e006700650073006c006f00740065006e002e> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! filelist convert a pdf file! page 2 page 3 1.pdf page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 pond.qxd floristic analysis of the mountain zebra national park, eastern cape u. pond, b.b. beesley, l.r. brown & h. bezuidenhout pond, u., b.b. beesley, l.r. brown & h. bezuidenhout. 2002. floristic analysis of the mountain zebra national park, eastern cape. koedoe 45(1): 35–57. pretoria. issn 0075-6458. as part of a larger project to assess the vegetation dynamics and conservation potential of the enlarged mountain zebra national park, a checklist was produced to determine the plant species richness for this area. six hundred and eighty species, represented by 333 genera and 87 families were identified. one hundred and eighty species belong to the monocotyledoneae and 479 species to the dicotyledoneae. by far the largest families are the asteraceae with 129 and the poaceae with 82 species. thirteen red data species were recorded. a number of fynbos elements were encountered, the most noteworthy being two families endemic to the cape floristic region, the penaeaceae and grubbiaceae. a very high species to square kilometre ratio of 5.05 supports the area’s rich floristic composition. key words: floristic analysis, species richness, plant species list, mountain zebra national park. u. pond, b.b. beesley, l.r. brown , applied natural sciences, technikon sa, private bag x6, florida 1710, south africa; h. bezuidenhout, conservation development, south african national parks, p o box 110040, hadison park, kimberley 8306, south africa. issn 0075-6458 35 koedoe 45/1 (2002) introduction the mountain zebra national park (mznp) was proclaimed a protected area in 1937 (wahl & naude 1996). it is situated in a zone transitional between the nama karoo biome in the west and the grassland biome in the east (hoffman 1998), but classified as being part of the nama karoo (low & rebelo 1998). the primary objective for this park is to conserve a viable, genetically uncontaminated, representative population of the cape mountain zebra equus zebra zebra. secondary objectives are the conservation of a representative spectrum of the typical faunal elements under natural conditions, and secodly, vegetation types that are unique to this region (van der walt 1980). in 1996, a process was initiated to incorporate adjacent farmland into the park. this will increase the park’s area substantially, from the original 6 536 ha to approximately 18 000 ha (brown & bezuidenhout 2000), once current negotiations have been completed. two extensive vegetation studies have been conducted: a phytosociological reconnaissance of the original mznp by van der walt (1980); and a phytosociological account, describing and mapping the plant communities of the newly acquired de rust section of the mznp (brown & bezuidenhout 2000). as part of a larger and long-term research project, the latter study aimed at assessing the habitat suitability to establish large herbivores on the new farms. no annotated comprehensive plant species list of mznp is available. surveys of the floristic diversity and species richness of an area are fundamental to any wildlife management programme and conservation policy. a floristic database of an area serves as a permanent reference to establish site uniqueness and species richness, monitoring changes in species occurrence and the distribution of endemic and red data species. this study set out to provide a floristic checklist for the pond.qxd 2005/12/09 11:34 page 35 mznp including the newly acquired farms de rust, doornhoek and welgedacht. study area location the mznp is found in the eastern cape province, situated 24 kilometres west of cradock. the area under discussion extends from 32º06'50''–32º18'50''s and 25º24'00''– 25º30'00''e and includes the former mznp (6 536 ha), de rust (1 712 ha), doornhoek (4 027 ha) and welgedacht (1 189 ha). topography the southern parts of the mznp are characterised by mountainous terrain with steepsided drainage lines from mountaintops koedoe 45/1 (2002) 36 issn 0075-6458 figure 1. location of the study area. (bankberg 1 927 m and rooiplaat 1 360 m above sea level. the wilgerboom river traverses the central valley of the southern section and exits the park at 1 200 m a.s.l. to the north. the northwest (de rust) is dominated by salpeterkop (1 514 m a.s.l.), a typical mesa of the karoo landscape with a steep gradient. this gives way to a mid-slope plateau, footslopes and an undulating plain with seasonally dry river beds towards the eastern and central part of the park, at approximately 1 000 m above sea level. the heterogeneous vegetation is co-dominated by dwarf shrubs and grasses, with patches of incipient forest in drainage lines and the extensive grass plain of rooiplaat. climate the mean rainfall for the area is 382 mm per annum (1962–1998), ranging from the highest rainfall of 651 mm (1977) to the lowest rainfall of 153 mm (1966), as measured at the mznp weather station. the rainy season occurs mostly in late summer and autumn, and the winters are relatively dry. mean monthly minimum and maximum temperatures vary from 6–28 ºc in summer (september–march) and from 0–20 ºc during winter (april– august) (brown & bezuidenhout 2000). extreme temperatures range from –7 ºc to 37 ºc and severe frost is likely during the period may–october (van der walt 1980). van der walt (1980), further states that the vegetation is subjected to continuous climatic variables due to the influence of an arid climate from the east and a more moderate climate from the west. the southern mountainous peaks are exposed to increased cloud cover, extreme temperature and moisture regimes, with local climate variations resulting in a number of microhabitats. bankberg forms a barrier to cold winter fronts, thus a warmer climate is experienced in the sheltered valpond.qxd 2005/12/09 11:34 page 36 ley below whilst regular snowfall occurs on higher lying areas (van der walt 1980). geology and soil mudstone, sandstone and shale of the beaufort group of the karoo supergroup dominate the mznp (keyzer 1997). these mudstones and shales are relatively unstable and subject to mechanical and chemical weathering. the material derived from these processes is usually rich in clay and salts. in addition, the mznp sediments have been penetrated on a large scale by post-karoo dolerite intrusions (keyzer 1997), forming large sheets and a number of dykes. the southern part of the park is characterised by dolerite outcrops that are remnants of the former sheet, of which the resistant bankberg is a prominent feature (van der walt 1980). highly fertile clayey soils are also derived from the fairly resistant doleritic parent material. calcareous bedrock (limestone) lines some river courses (pers.obs.), formed by chemical precipitation from sandstone or mudstone and found in tributaries of the wilgerboom river from kranskop peak (brown & bezuidenhout 2000). methods plant collections were undertaken over a two-year period during 1999 and 2000. the areas covered included de rust, doornhoek, welgedacht and the following areas within the former mznp: rooiplaat, kranskop, the top of bankberg and along the kloofs down to weltevrede and fonteinkloof. collection sites were selected randomly within the extensive area according to topography and plant phenology, ensuring that all variations in habitat were considered and sampled. all specimens were labelled, pressed and dried, then identified by the selmar schönland herbarium in grahamstown and form part of the collection of the kimberley south african national parks herbarium. selected duplicates are kept at the selmar schönland herbarium and technikon sa. the final species list was compiled from various sources: pond and beesley’s collection, van der walt (1980) and a species list supplied by precis (national herbarium pretoria computerised information system) for the mznp. plant names were entered into and consolidated by sas (2002), a database for the cape and karoo floras. while arnold & de wet (1993) served as a basis for nomenclature and arrangement, sas introduced the latest taxonomic changes and appropriate author updates. only identified species were listed. red data status, also produced by sas, follows hilton-taylor (1996). introduced species were included and are followed by an asterisk (*). results the mznp plant species list totals 680 species, which represent 333 genera and 87 families. flowering plants are represented by monocotyledoneae with 180 species in 14 families (16 % of the total number of families) and dicotyledoneae with 479 species in 62 families (71 %). the bryophytes with seven families (8 %) and the pteridophytes with four families (5 %) represent non-flowering plants (fig. 2). the complete species list, separated into the different plant divisions, sorted by family and indicating the species’ red data status where applicable, appears as appendix 1. this study added 100 new species to previous records for the mznp. introduced species were included in all floristic calculations. the mznp flora is represented by 87 families, as reflected in table 1. issn 0075-6458 37 koedoe 45/1 (2002) fig. 2. plant divisions reflected as a percentage of the total number of plant families. pond.qxd 2005/12/09 11:34 page 37 koedoe 45/1 (2002) 38 issn 0075-6458 families genera species bryophytes aytoniaceae 1 1 bryaceae 1 1 fissidentaceae 1 1 grimmiaceae 2 2 pottiaceae 2 2 ptychomitriaceae 1 1 ricciaceae 1 2 pteridophytes anemiaceae 1 1 aspleniaceae 1 2 equisetaceae 1 1 pteridaceae 2 7 monocotyledoneae alliaceae 1 1 amaryllidaceae 5 9 asparagaceae 1 12 asphodelaceae 5 13 commelinaceae 2 3 convallariaceae 2 2 cyperaceae 9 18 hyacinthaceae 7 17 hypoxidaceae 2 3 iridaceae 8 12 juncaceae 1 6 poaceae 47 82 potamogetonaceae 1 1 tecophilaeaceae 1 1 dicotyledoneae acanthaceae 2 3 aizoaceae 3 6 amaranthaceae 5 10 anacardiaceae 2 9 apiaceae 6 6 apocynaceae 12 14 araliaceae 1 1 asteraceae 51 129 bignoniaceae 1 1 boraginaceae 3 4 brassicaceae 10 13 buddlejaceae 2 3 campanulaceae 3 12 caryophyllaceae 3 4 celastraceae 2 4 families genera species commelinaceae 2 2 convolvulaceae 3 3 crassulaceae 2 18 cucurbitaceae 2 3 dipsacaceae 1 2 ebenaceae 2 5 ericaceae 1 5 euphorbiaceae 2 10 fabaceae 13 35 gentianaceae 2 2 geraniaceae 4 21 grubbiaceae 1 1 haloragaceae 1 1 kiggelariaceae 1 1 lamiaceae 7 13 linaceae 1 1 loranthaceae 1 1 malvaceae 5 15 melianthaceae 1 1 mesembryanthemac. 10 22 molluginaceae 3 3 myrsinaceae 2 2 nyctaginaceae 1 1 oleaceae 1 1 orobanchaceae 1 1 oxalidaceae 1 5 papaveraceae 2 2 pedaliaceae 2 2 penaeaceae 1 2 plantaginaceae 1 1 polygalaceae 2 6 polygonaceae 2 2 portulacaceae 1 1 ranunculaceae 3 3 rhamnaceae 1 1 rosaceae 1 1 rubiaceae 4 5 salicaceae 2 2 santalaceae 1 1 sapindaceae 1 1 scrophulariaceae 10 31 solanaceae 5 12 thymelaceae 2 3 urticaceae 1 2 verbenaceae 4 4 viscaceae 1 2 vitaceae 2 2 zygophyllaceae 2 2 total 87 680 table 1 an alphabetical list of plant families collected in the mznp, indicating the number of genera and species in each family pond.qxd 2005/12/09 11:34 page 38 issn 0075-6458 39 koedoe 45/1 (2002) eight families dominate the mznp flora. by far the two largest families are the asteraceae with 129 species, reflecting 19 % of the total flora and the poaceae with 82 species (12 %). these are followed by much smaller, yet significant families, represented by the fabaceae with 35 species (5.2 %), scrophulariaceae 31 (4.6 %), mesembryanthemaceae 22 (3.2 %), geraniaceae 21 (3.1 %), cyperaceae 18 (2.6 %) and the crassulaceae also with 18 species and 2.6 % of the species total (fig. 3). these eight families represent 52.3 % of the mznp flora, while the other 79 families reflect the remaining 47.7 %. twenty-nine genera are represented by five or more species. these are listed in table 2. thirteen red data species were found, representing 1.9 % of the mznp flora (fig. 4). red data species and the categorisation are indicated in the species list (appendix 1). symbols and terms used in appendix 1: fig. 3. dominant plant families reflecting the number of species and percentage of the total flora genus no of species helichrysum 20 senecio 17 pelargonium 17 crassula 14 asparagus 12 euryops 9 euphorbia 9 selago 9 eragrostis 8 rhus 8 wahlenbergia 8 indigofera 7 hermannia 7 jamesbrittania 7 genus no of species juncus 6 lotononis 6 delosperma 6 cheilanthes 6 oxalis 5 polygala 5 pentzia 5 pteronia 5 erica 5 lessertia 5 felicia 5 ornithogalum 5 stachys 5 aristida 5 lycium 5 table 2 dominant genera and number of species per genus pond.qxd 2005/12/09 11:34 page 39 introduced species (*) amount to 46 (6.7 %) (appendix 1). the majority of these belong to the poaceae, represented by the genera: bromus, chloris, eleusine, eragrostis, hordeum, melinis, paspalum, poa, polypogon, setaria, tragus, vulpia; amaranthaceae: alternanthera, atriplex, chenopodium, salsola; asteraceae: anthemis, bidens, cirsium, lactuca, schkuhria, verbesina; brassicaceae: capsella, cardaria, descurainia, sisymbrium; caryophyllaceae: stellaria; convolvulaceae: cuscuta; malvaceae: malva; papaveraceae: argemone; plantaginaceae: plantago; anacardiaceae: schinus; apiaceae: ammis, ciclospermum; cyperaceae: cyperus; polygonaceae: polygonum; salicaceae: populus; solanaceae: datura, nicotiana; urticaceae: urtica; verbenaceae: verbena. discussion the mountain zebra national park boasts 680 species, 333 genera and 87 families. this contrasts with the low plant species diversity for the nama karoo claimed by hall, (1980) some 20 years ago. other nama karoo floras compare as follows: augrabies falls national park with 364 species, karoo national park (822), karoo nature reserve (320) and of the succulent karoo: goegap nature reserve (550), richtersveld national park (537) and tankwa karoo national park with 245 species (sas 2002) (table 3). as illustrated in fig. 3, the high presence of asteraceae (19 %) corresponds with most floras of the sub-region and it is usually the largest family in floras of arid to semi-arid regions (goldblatt & manning 2000). the graminoids are strongly represented by the poaceae (12 %). rutherford & westfall (1986) remark that hemicryptophytes of the nama karoo biome are mainly c4 grasses, which are adapted to high temperatures and low water supply. this is relative, as the eastern mixed nama karoo has the highest rainfall of all the karoo types and is thus ecotonal to grassland (hoffman 1998). the relatively high proportion of fabaceae (5.2 %) is not exceptional, as this family is well developed in most parts of the world. so is the wealth of scrophulariaceae (4.6 %), which is reflected right across africa, especially in the floras of drier areas (goldblatt & manning 2000). there is a gradient of species replacement within the dry karoo from winter to summer rainfall. for example, the mesembryanthemaceae, represented by 3.2 % in the mznp, ranked amongst the dominant families. however, the mesembryanthemaeceae decline from the west (the centre of koedoe 45/1 (2002) 40 issn 0075-6458 fig. 4. number and status of red data species according to hilton-taylor (1996). pond.qxd 2005/12/09 11:34 page 40 endemism and species radiation) to the east, where they are replaced by other groups, such as the crassulaceae, which make up 2.6 % of the mznp flora. of the 87 plant families, 71 % are dicotyledoneae and 16 % monocotyledoneae (fig. 2). monocoteae are generally herbaceous, whereas dicots are woody or herbaceous and in general much more diverse in habitat (jones & luchsinger, 1987), suggesting that they are also accustomed to greater edaphic and climatic variations. a ratio of 3.8 dicots to 1 monocot species for the mznp is slightly higher than that of the cape floristic region and the average for floras across the world at 3:1 (goldblatt & manning 2000). although less diverse in habitat, monocot families have specialised to co-dominate in most floras of the sub-region. graminoids, such as restionaceae, poaceae and cyperaceae are respectively ranked as the 8th, 11th and 12th largest families of the cape flora (goldblatt & manning 2000). whilst the restionaceae depend on winter rainfall and are less prominent east of the cape floral kingdom, the poaceae emerge in the more arid summer rainfall areas and rate as the 2nd largest family in the mznp (12 % of the total flora). hoffman & cowling (1987) noted that an overall increase in grass cover from west to east correlated with an overall increase in rainfall and a decline in winter rainfall. the cyperaceae (generally associated with impoverished sandstone soils of the cape mountain ranges by goldblatt & manning, (2000) amount to 2.6 % of the flora (18 species) in the mountain zebra national park. red data species make up 1.9 % of the mznp flora (fig. 4; appendix 1), which is low compared to fynbos, for example, which displays an average of 3–7 % of red data species. other karoo floras, however, also display low numbers of red data species, e.g., augrabies falls national park with 0.6 %, karoo national park 2.2 %, karoo nature reserve 1.6 %, tankwa karoo national park 2.4 %, goegap nature reserve 2.4 %. richtersveld national park has a contrasting high of 6 % (sas 2002). hall et al. (1980), also found the number of rare and endangered species for the nama karoo biome to be low. a unique feature of mznp is the occurrence of two families that are endemic to fynbos, i.e., the penaeaceae and grubbiaceae. whilst penaea cneorum has been recorded from the kogelberg and riviersonderend mountains to port elizabeth, penaea acutifolia is a new record, previously only recorded from the outeniqua mountains to george. also, grubbia rosmarinifolia, whose current range of distribution is restricted from the koue bokkeveld to the tsitsikamma mountains (goldblatt & manning 2000), is a fynbos element and has been recorded in the park. apart from the two records above, a number of fynbos elements were found in the park. according to low (a.b. low, p o box 370, rondebosch, 7701) (pers. comm. 2000), locally occurring genera, such as eriocephalus, elytropappus, euryops, maytenus and diospyros are normally associated with renosterveld (on shale), while passerina, erica and gnidia as well as the amaryllidaceae are also associated with fynbos. passerina and dodonaea spp. are known to link summer and winter rainfall areas. in addition, arid fynbos is normally characterised by the absence of ericaceae, and yet, here it is found in the more arid karoo. to the same extent, leucosidea sericea that commonly occurs in drakensberg streams has also been recorded in the mountain zebra national park. the floristic region of southern africa (botswana, lesotho, namibia, south africa and swaziland) has about 20 400 native vascular plant species in an area of 2 674 000 km². south africa alone may have some 18 500 species of these, considered a particularly rich and diverse flora for a predominantly temperate region. the cape floristic region (cfr) at 90 000 km², with 9 000 species, is striking by world standards (goldblatt & manning 2000). at a smaller scale, however, the mznp study area (13 464 ha of the entire 18 000 ha) and other karoo floras were compared in table 3 issn 0075-6458 41 koedoe 45/1 (2002) pond.qxd 2005/12/09 11:34 page 41 (zietsman & bezuidenhout 1999; zietsman et al. 1992; van rooyen & bezuidenhout 1997; sas 2002). plant species richness at 5.05 species per 100 ha is considerably higher when compared to the other parks and reserves. thus the mznp displays a rich floristic composition, a result of extreme habitat and substrate diversity, ranging from sandstones to shales and dolerite dykes to calcareous bedrock with extensive grass plains, shrublands, high altitude seeps and riverine thicket. microclimatic conditions play an important role in the germination and establishment of seedlings of karoo plants (roux & theron 1987). geophytes seem best adapted to a seasonally extreme climate with a wet winter and dry summer, in semi-arid conditions (goldblatt & manning 2000). this is evident in the succulent karoo, where geophytes are most abundant, both in terms of species richness and cover (hoffman & cowling 1987). there is a high incidence of geophytes in the mznp (8 % of the total flora) represented by the alliaceae, amaryllidaceae, asphodelaceae, hyacinthaceae, hypoxydaceae and iridaceae. despite the fact that most of the park experiences a semi-arid climate with erratic rainfall that occurs mainly in summer, hoffman & cowling (1987) remark that geophytes are very conspicuous in the eastern karoo. conclusion the eastern mixed nama karoo comprises of 77 784 km², of which only 1.08 % is presently conserved (low & rebelo 1998). complex soils, erratic rainfall and climate, as well as variations in altitude and habitat have driven the processes leading to a relatively high species richness for the mountain zebra national park. it is clear that vegetation is much more than a list of plant species. this floristic account, together with the studies on plant communities (van der walt 1980; brown & bezuidenhout 2000), serves as a basis to develop a habitat management plan for the national park. considered an ecotone between the nama karoo and grassland biomes, this area displays a rich variety of vegetation types and habitats in juxtaposition. the results of this study indicate that the newly enlarged mountain zebra national park forms an important conservation area and is a noteworthy centre of biological diversity, including species richness, ecosystem complexity and genetic variation. it is therefore important that a detailed management plan be developed for the park to ensure the sustainalbe utilisation and conservation of the vegetation of the area. acknowledgements the following individuals and institutions are sincerely thanked: south african national parks (sanparks) and technikon sa (tsa) for initiating this koedoe 45/1 (2002) 42 issn 0075-6458 table 3 comparison of the species/area ratio of selected protected areas in south africa area size (ha) number of species species per 100 ha augrabies falls national park 18 600 364 1.95 goegap nature reserve 14 531 550 3.78 kalahari gemsbok national park 959 300 489 0.05 karoo national park 43 261 822 1.90 karoo nature reserve 16 000 320 2.00 mountain zebra national park 13 464 680 5.05 richtersveld national park 162 445 537 0.33 tankwa karoo national park 27 064 245 0.90 vaalbos national park 22 696 334 1.47 pond.qxd 2005/12/09 11:34 page 42 study; national research foundation (nrf) and technikon sa for financing this study; national botanical institute for the use of data from the national herbarium, pretoria (pre) computerised information system (precis) and for ad hoc identifications; tony dold at the selmar schönland herbarium, grahamstown for plant identifications; barrie low, environmental consultant, for the input and update of the species list via the sas database and for comment on the draft report; johan de klerk, section game ranger at mznp, for his help and support; mountain zebra national park warden paddy gordon (at the time) and staff for accommodating this study. references acocks, j.p.h. 1988. veld types of south africa. 3rd edn. memoirs of the botanical survey of south africa 57: 1–146. arnold, t.h. & b.c. de wet. 1993. plants of southern africa: names and distribution. memoirs of the botanical survey of south africa 62: 5–825. brown, l.r. & h. bezuidenhout. 2000. the phytosociology of the de rust section of the mountain zebra national park, eastern cape. koedoe 43(1): 1–18. goldblatt, p. & j. manning. 2000. cape plants. a conspectus of the cape flora of south africa. strelitzia 9. cape town: missouri botanical gardens usa & national botanical institute, sa. hall, a.v., m. de winter, b. de winter & s.a.m. oosterhout. 1980. description of biomes. pp. 45–65. in: rutherford, m.c. & r.h. westfall. biomes of southern africa—an objective categorization. pretoria: botanical research institute, department of agriculture and water supply. (memoirs of the botanical survey of south africa; no 54.) hilton-taylor, c. 1996. red data list of southern african plants. strelitzia 4. cape town: conservation biology research unit, national botanical institute. hoffman, t. 1998. eastern mixed nama karoo. p. 55. in: low, a.b. & a.g. rebelo (eds.) 1998. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. hoffman, m.t. & r.m. cowling. 1987. plant physiognomy, phenology and demography. pp. 1–34. in: cowling, r.m. & p.w. roux (eds.). the karoo biome: a preliminary synthesis. part 2 – vegetation and history. pretoria: council for scientific and industrial research, national scientific programmes unit. (south african national scientific programmes report; no 142). jones, s.b. & a.e. luchsinger. 1987. plant systematics. 2nd ed. singapore: mcgraw-hill. keyzer, n. 1997. geological map of the republic of south africa and kingdoms of lesotho and swaziland, 1:1 000 000. pretoria: council for geoscience. low, a.b. & a.g. rebelo. (eds.) 1998. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. roux, p.w. & g.k. theron. 1987. vegetation change in the karoo biome. pp. 50–69. in: cowling, r.m. & p.w. roux (eds.). the karoo biome: a preliminary synthesis. part 2 – vegetation and history. pretoria: council for scientific and industrial research, national scientific programmes unit. (south african national scientific programmes report; no 142). rutherford, m.c. & r.h. westfall. 1986. biomes of southern africa – an objective categorization. pretoria: botanical research institute, department of agriculture and water supply. (memoirs of the botanical survey of south africa; no. 54.) sas. 2002. site and species database for the cape and karoo floras. rondebosch: coastec. van der walt, p.t. 1980. a phytosociological reconnaissance of the mountain zebra national park. koedoe 23: 1–32. van rooyen, n. & h. bezuidenhout. 1997. new records of flowering plants and ferns from the kalahari gemsbok national park. koedoe 40(2): 105–116. wahl, m. & k. naude. 1996. national register of protected areas in south africa. pretoria: department of environmental affairs and tourism. zietsman, p.c. & h. bezuidenhout. 1999. flowering plant biodiversity of augrabies falls national park: a comparison between augrabies falls national park, kalahari gemsbok national park, vaalbos national park and goegap nature reserve. koedoe 42(2): 95–112. zietsman, p.c., p.j. du preez & h. bezuidenhout. 1992. a preliminary check list of flowering plants of the vaalbos national park. koedoe 35(1): 89–98. appendix 1 plant species list for the mountain zebra national park issn 0075-6458 43 koedoe 45/1 (2002) appendix 1 .../ pond.qxd 2005/12/09 11:34 page 43 e endangered – taxa in danger of extinction if causal factors continue operating ex extinct – taxa which are no longer known to exist in the wild i indeterminate –taxa known to be extinct, endangered, vulnerable or rare but information is insufficient to categorise them k insufficiently known –taxa that are suspected to belong to any of the above categories but information is lacking r rare – taxa with small world populations, presently not endangered or vulnerable but at risk that a threat could cause a critical decline v vulnerable –taxa believed likely to move into the endangered category in the near future if the factors causing decline continue operating us unspecified –taxa previously threatened but are no longer in any of the above categories due to an increase in population size or to subsequent discoveries * introduced species family species red data pres/past bryophytes aytoniaceae plagiochasma rupestre var. rupestre (g.forst.) steph. bryaceae bryum turbinatum (hedw.) turner fissidentaceae fissidens rufescens hornsch. grimmiaceae grimmia pulvinata (hedw.) sm. grimmiaceae schistidium apocarpum (hedw.) bruch & schimp. pottiaceae microbryum davallianum (sm.) r.h.zander pottiaceae triquetrella tristicha (c.müll.) c.müll. ptychomitriaceae ptychomitrium subcrispatum thér. & p.vard. ricciaceae riccia nigrella dc. ricciaceae riccia simii perold complete list selective list total species: 680 total species: 10 total genera: 333 total genera: 9 total families: 87 total families: 7 total red data species: 13 total red data species: 0 pteridophytes anemiaceae mohria vestita baker aspleniaceae asplenium cordatum (thunb.) sw. aspleniaceae asplenium trichomanes l. equisetaceae equisetum ramosissimum desf. pteridaceae cheilanthes bergiana schlechtd. pteridaceae cheilanthes eckloniana (kunze) mett. pteridaceae cheilanthes hirta sw. pteridaceae cheilanthes induta kunze pteridaceae cheilanthes multifida (sw.) sw. subsp. multifida pteridaceae cheilanthes parviloba (sw.) sw. pteridaceae pellaea calomelanos (sw.) link var. calomelanos complete list selective list koedoe 45/1 (2002) 44 issn 0075-6458 pond.qxd 2005/12/09 11:34 page 44 total species: 680 total species: 11 total genera: 333 total genera: 5 total families: 87 total families: 4 total red data species: 13 total red data species: 0 monocotyledoneae alliaceae tulbaghia galpinii schltr. amaryllidaceae ammocharis coranica (ker gawl.) herb. amaryllidaceae boophone disticha (l.f.) herb. amaryllidaceae cyrtanthus contractus n.e.br. amaryllidaceae cyrtanthus macowanii baker us amaryllidaceae cyrtanthus spiralis burch. ex ker gawl. v amaryllidaceae cyrtanthus suaveolens schönland k amaryllidaceae haemanthus albiflos jacq. amaryllidaceae haemanthus humilis subsp. humilis jacq. amaryllidaceae nerine huttoniae schönland k asparagaceae asparagus acocksii jessop asparagaceae asparagus capensis var. capensis l. asparagaceae asparagus cooperi baker asparagaceae asparagus densiflorus (kunth) jessop asparagaceae asparagus denudatus (kunth) baker asparagaceae asparagus exuvialis burch. forma exuvialis asparagaceae asparagus glaucus kies asparagaceae asparagus mucronatus jessop asparagaceae asparagus multiflorus baker asparagaceae asparagus retrofractus l. asparagaceae asparagus striatus (l.f.) thunb. asparagaceae asparagus suaveolens burch. asphodelaceae aloe striata haw. subsp. striata asphodelaceae aloe striatula haw. var. caesia reynolds asphodelaceae aloe variegata l. asphodelaceae bulbine abyssinica a.rich. asphodelaceae bulbine frutescens (l.) willd. asphodelaceae bulbine narcissifolia salm-dyck asphodelaceae gasteria bicolor haw. var. bicolor asphodelaceae haworthia arachnoidea (l.) duval asphodelaceae haworthia deltoidea (hook.f.) parr var. deltoidea asphodelaceae haworthia herbacea (mill.) stearn var. herbacea asphodelaceae haworthia lockwoodii archibald v asphodelaceae kniphofia acraea codd r asphodelaceae kniphofia uvaria (l.) oken commelinaceae commelina africana l. var. africana commelinaceae commelina africana l. var. lancispatha c.b.clarke commelinaceae cyanotis speciosa (l.f.) hassk. convallariaceae eriospermum corymbosum baker convallariaceae sansevieria aethiopica thunb. cyperaceae bulbostylis humilis (kunth) c.b.clarke cyperaceae carex spicato-paniculata c.b.clarke cyperaceae cyperus denudatus l.f cyperaceae cyperus laevigatus l. cyperaceae cyperus rotundus l. * cyperaceae cyperus usitatus burch. cyperaceae fuirena coerulescens steud. cyperaceae isolepis costata (boeck.) a.rich. var. costata cyperaceae isolepis karroica (c.b.clarke) j.raynal issn 0075-6458 45 koedoe 45/1 (2002) pond.qxd 2005/12/09 11:34 page 45 cyperaceae isolepis setacea (l.) r. br. cyperaceae kyllinga alata nees cyperaceae kyllinga alba nees cyperaceae kyllinga pulchella kunth cyperaceae mariscus congestus (vahl) c.b.clarke cyperaceae mariscus sumatrensis (retz.) j. raynal cyperaceae mariscus uitenhagensis steud. cyperaceae pseudoschoenus inanis (thunb.) oteng-yeboah cyperaceae schoenoxiphium sparteum (wahlenb.) c.b.clarke hyacinthaceae albuca flaccida jacq. hyacinthaceae albuca maxima burm.f. hyacinthaceae albuca shawii baker hyacinthaceae dipcadi ciliare (zeyh. ex harv.) baker hyacinthaceae eucomis comosa (houtt.) h.r.wehrh. var. comosa hyacinthaceae lachenalia karooica w.f.barker ex g.d.duncan hyacinthaceae lachenalia variegata w.f.barker hyacinthaceae ledebouria apertiflora (baker) jessop hyacinthaceae ledebouria cooperi (hook.f.) jessop hyacinthaceae ledebouria undulata (jacq.) jessop hyacinthaceae massonia echinata l.f. hyacinthaceae massonia pustulata jacq. hyacinthaceae ornithogalum graminifolium thunb. hyacinthaceae ornithogalum juncifolium jacq. hyacinthaceae ornithogalum maculatum jacq. hyacinthaceae ornithogalum paludosum baker hyacinthaceae ornithogalum secundum jacq. us hypoxidaceae empodium plicatum (thunb.) garside hypoxidaceae hypoxis angustifolia lam. var. angustifolia hypoxidaceae hypoxis argentea var. argentea harv. ex baker iridaceae dierama pendulum (l.f.) baker iridaceae gladiolus permeabilis d.delaroche subsp. edulis (burch. ex ker gawl.) oberm. iridaceae hesperantha longituba (klatt) baker iridaceae lapeirousia plicata (jacq.) diels subsp. plicata iridaceae moraea elliotii baker iridaceae moraea polystachya (thunb.) ker gawl. iridaceae moraea simulans baker iridaceae romulea atrandra g.j.lewis var. atrandra iridaceae syringodea concolor (baker) m.p.de vos iridaceae tritonia dubia eckl. ex klatt iridaceae tritonia laxifolia benth. ex baker iridaceae tritonia lineata (salisb.) ker gawl. juncaceae juncus dregeanus kunth juncaceae juncus effusus l. juncaceae juncus oxycarpus e.mey. ex kunth juncaceae juncus exsertus buchenau subsp. exsertus juncaceae juncus inflexus l. juncaceae juncus rigidus desf. poaceae agrostis lachnantha nees var. lachnantha poaceae aristida adscensionis l. poaceae aristida congesta roem. & schult. subsp. congesta poaceae aristida diffusa trin. subsp. burkei (stapf) meld. poaceae aristida diffusa trin. subsp. diffusa poaceae aristida junciformis trin. & rupr. poaceae bothriochloa radicans (lehm.) a.camus poaceae bromus catharticus vahl * koedoe 45/1 (2002) 46 issn 0075-6458 pond.qxd 2005/12/09 11:34 page 46 poaceae bromus pectinatus thunb. * poaceae cenchrus ciliaris l. poaceae chloris virgata sw. * poaceae cymbopogon dieterlenii stapf ex e.phillips poaceae cymbopogon marginatus (steud.) stapf ex burtt davy poaceae cymbopogon plurinodis (stapf) stapf ex burtt davy poaceae cynodon dactylon (l.) pers. poaceae cynodon incompletus nees poaceae digitaria eriantha steud. poaceae diplachne fusca (l.) p.beauv. ex roem. & schult. poaceae ehrharta calycina j.e.sm. var. calycina poaceae ehrharta erecta lam. var. erecta poaceae eleusine coracana (l.) gaertn. * poaceae elionurus muticus (spreng.) kunth poaceae enneapogon scoparius stapf poaceae eragrostis barrelieri daveau * poaceae eragrostis bergiana (kunth) trin. poaceae eragrostis chapelieri (kunth) nees poaceae eragrostis chloromelas steud. poaceae eragrostis curvula (schrad.) nees poaceae eragrostis lehmanniana nees var. lehmanniana poaceae eragrostis obtusa munro ex ficalho & hiern poaceae eragrostis plana nees poaceae eustachys paspaloides (vahl) lanza & mattei poaceae festuca scabra vahl poaceae fingerhuthia africana lehm. poaceae fingerhuthia sesleriiformis nees poaceae helictotrichon capense schweick. poaceae heteropogon contortus (l.) p.beauv. ex roem. & schult. poaceae hordeum stenostachys godr. * poaceae hyparrhenia hirta (l.) stapf poaceae koeleria capensis (steud.) nees poaceae melica decumbens thunb. poaceae melica racemosa thunb. poaceae melinis nerviglumis (franch.) zizka poaceae melinis repens (willd.) ziska * poaceae merxmuellera disticha (nees) conert poaceae merxmuellera stricta (schrad.) conert poaceae microchloa caffra nees poaceae miscanthus capensis (nees) andersson poaceae oropetium capense stapf poaceae panicum coloratum l. var. coloratum poaceae panicum deustum thunb. poaceae panicum maximum jacq. poaceae panicum stapfianum fourc. poaceae paspalum dilatatum poir. * poaceae pennisetum macrourum trin. poaceae pennisetum sphacelatum (nees) t.durand & schinz poaceae pentaschistis airoides (nees) stapf subsp. airoides poaceae pentaschistis airoides (nees) stapf subsp. jugorum (stapf) h.p.linder poaceae pentaschistis curvifolia (schrad.) stapf poaceae pentaschistis glandulosa (schrad.) h.p.linder poaceae phragmites australis (cav.) trin. ex steud. poaceae poa annua l. * poaceae polypogon monspeliensis (l.) desf. * poaceae schismus barbatus (loefl. ex l.) thell. issn 0075-6458 47 koedoe 45/1 (2002) pond.qxd 2005/12/09 11:34 page 47 poaceae schismus inermis (stapf) c.e.hubb. poaceae setaria nigrirostris (nees) t.durand & schinz poaceae setaria sphacelata (schum.) stapf & c.e.hubb. ex moss var. sphacelata poaceae setaria verticillata (l.) p.beauv. * poaceae sporobolus africanus (poir.) robyns & tournay poaceae sporobolus discosporus nees poaceae sporobolus fimbriatus (trin.) nees poaceae stipa dregeana steud. var. dregeana poaceae stipa dregeana steud. var. elongata (nees) stapf poaceae stipagrostis ciliata (desf.) de winter poaceae stipagrostis obtusa (del.) nees poaceae tetrachne dregei nees poaceae themeda triandra forssk. poaceae tragus berteronianus schult. * poaceae tragus koelerioides aschers. poaceae tribolium hispidum (thunb.) desv. poaceae urochloa panicoides p.beauv. poaceae vulpia myuros (l.) c.c.gmel. * potamogetonaceae potamogeton pusillus l. tecophilaeaceae cyanella lutea l.f. complete list selective list total species: 680 total species: 180 total genera: 333 total genera: 92 total families: 87 total families: 14 total red data species: 13 total red data species: 5 dicotyledoneae acanthaceae blepharis capensis var. capensis (l.f.) pers. acanthaceae blepharis villosa (nees) c.b.clarke acanthaceae justicia orchioides l.f. subsp. glabrata immelman us aizoaceae aizoon glinoides l.f. aizoaceae galenia africana l. aizoaceae galenia procumbens l.f. aizoaceae galenia pubescens var. pubescens (eckl. & zeyh.) druce aizoaceae galenia sarcophylla fenzl aizoaceae tetragonia echinata aiton amaranthaceae alternanthera pungens h.b.&k. * amaranthaceae atriplex semibaccata r.br. * amaranthaceae atriplex suberecta i.verd. amaranthaceae atriplex vestita (thunb.) aellen amaranthaceae chenopodium mucronatum thunb. * amaranthaceae chenopodium murale l. * amaranthaceae chenopodium schraderianum roem. & schult. * amaranthaceae salsola aphylla l.f. amaranthaceae salsola kali l. * amaranthaceae suaeda fruticosa (l.) forssk. anacardiaceae rhus divaricata eckl. & zeyh. anacardiaceae rhus dregeana sond. anacardiaceae rhus erosa thunb. anacardiaceae rhus gracillima engl. anacardiaceae rhus lancea l.f. koedoe 45/1 (2002) 48 issn 0075-6458 pond.qxd 2005/12/09 11:34 page 48 anacardiaceae rhus lucida l. anacardiaceae rhus pyroides burch var. pyroides. anacardiaceae rhus undulata jacq. anacardiaceae schinus molle l. * apiaceae ammis majus l. var. glaucifolium (l.) godron * apiaceae berula erecta (huds.) coville subsp. thunbergii (dc.) b.l.burtt apiaceae bupleurum mundii cham. & schlechtd. apiaceae ciclospermum leptophyllum (pers.) sprague * apiaceae deverra burchellii (dc.) eckl. & zeyh. apiaceae heteromorpha arborescens (spreng.) cham. & schltdl. apocynaceae asclepias compressidens (n.e.br.) a.nicholas i apocynaceae brachystelma circinatum e.mey. apocynaceae carissa haematocarpa (eckl.) a.dc. apocynaceae duvalia caespitosa (masson) haw. var. caespitosa apocynaceae duvalia modesta n.e.br. apocynaceae gomphocarpus fruticosa l. apocynaceae gomphocarpus tomentosus burch. apocynaceae huernia brevirostris n.e.br. subsp. intermedia n.e.br. apocynaceae microloma armatum (thunb.) schltr. ex gilg var. armatum apocynaceae pachycymbium miscellum (n.e.br.) m.gilbert us apocynaceae pachypodium succulentum (l.f.) a.dc. apocynaceae sarcostemma viminale (l.) r.br. subsp. viminale apocynaceae schizoglossum aschersonianum schltr. var. aschersonianum apocynaceae stenostelma capense schltr. araliaceae cussonia paniculata eckl. & zeyh. subsp. paniculata asteraceae amellus strigosus (thunb.) less. asteraceae anthemis cotula l. * asteraceae arctotheca calendula (l.) levyns asteraceae arctotis arctotoides (l.f.) o.hoffm. asteraceae arctotis microcephala (dc.) p.beauv. asteraceae berkheya armata (vahl) druce asteraceae berkheya decurrens (thunb.) willd. asteraceae bidens pilosa l. * asteraceae chrysanthemoides monilifera (l.) norl. subsp. subcanescens (dc.) norl. asteraceae chrysocoma ciliata l. asteraceae cineraria aspera thunb. asteraceae cineraria geraniifolia dc. asteraceae cineraria lobata l’her. asteraceae cirsium vulgare (savi) ten. * asteraceae conyza podocephala dc. asteraceae conyza scabrida dc. asteraceae cotula heterocarpa dc. asteraceae cotula nigellifolia (dc.) bremer & humphries asteraceae cotula zeyheri fenzl asteraceae cuspidia cernua (l.f.) b.l.burtt asteraceae dimorphotheca cuneata (thunb.) less. asteraceae dimorphotheca zeyheri sond. asteraceae elytropappus rhinocerotis (l.f.) less. asteraceae eriocephalus africanus l. asteraceae eriocephalus ericoides (l.f.) druce asteraceae eriocephalus eximius dc. asteraceae eriocephalus punctulatus dc. asteraceae euryops annae e.phillips asteraceae euryops annuus compton asteraceae euryops anthemoides b.nord. subsp. anthemoides asteraceae euryops dentatus b.nord. r issn 0075-6458 49 koedoe 45/1 (2002) pond.qxd 2005/12/09 11:34 page 49 asteraceae euryops floribundus n.e.br. asteraceae euryops lateriflorus (l.f.) dc. asteraceae euryops oligoglossus dc. subsp. oligoglossus asteraceae euryops spathaceus dc. asteraceae euryops tenuissimus (l.) dc. asteraceae felicia fascicularis dc. asteraceae felicia filifolia (vent.) burtt davy subsp. filifolia asteraceae felicia muricata (thunb.) nees subsp. muricata asteraceae felicia ovata (thunb.) compton asteraceae felicia zeyheri (less.) nees subsp. linifolia (harv.) grau asteraceae foveolina albida (dc.) kallersjo asteraceae garuleum bipinnatum (thunb.) less. asteraceae garuleum pinnatifidum (thunb.) dc. asteraceae gazania krebsiana less. subsp. arctotoides (less.) roessler asteraceae gazania krebsiana less. subsp. krebsiana asteraceae gazania linearis (thunb.) druce asteraceae geigeria ornativa o.hoffm. asteraceae gymnostephium papposum nesom asteraceae helichrysum anomalum less. asteraceae helichrysum asperum (thunb.) hilliard & b.l.burtt var. asperum asteraceae helichrysum cerastioides dc. var. cerastioides asteraceae helichrysum dasycephalum o.hoffm. asteraceae helichrysum dregeanum sond. & harv. asteraceae helichrysum epapposum bolus asteraceae helichrysum hamulosum e.mey. ex dc. asteraceae helichrysum lineatum bolus asteraceae helichrysum lucilioides less. asteraceae helichrysum montanum dc. asteraceae helichrysum nudifolium (l.) less. asteraceae helichrysum odoratissimum (l.) sweet asteraceae helichrysum petiolare hilliard & b.l.burtt asteraceae helichrysum rosum (berg.) less. var. rosum asteraceae helichrysum rugulosum less. asteraceae helichrysum rutilans (l.) d.don us asteraceae helichrysum splendidum (thunb.) less. asteraceae helichrysum trilineatum dc. asteraceae helichrysum umbraculigerum less. asteraceae helichrysum zeyheri less. asteraceae hertia pallens (dc.) kuntze asteraceae ifloga glomerata (harv.) schltr. asteraceae kleinia longiflora dc. asteraceae lactuca inermis forssk. * asteraceae lasiopogon muscoides (desf.) dc. asteraceae lasiospermum bipinnatum (thunb.) druce asteraceae lasiospermum pedunculare lag. asteraceae marasmodes polycephalus dc. asteraceae mikania cordata (burm.f.) b.l.rob. asteraceae oldenburgia paradoxa less. asteraceae oligocarpus calendulaceus (l.f.) less. asteraceae oncosiphon piluliferum (l.f.) kallersjo asteraceae othonna auriculifolia licht. ex less. asteraceae othonna cylindrica (lam.) dc. asteraceae othonna pavonia e.mey. asteraceae pegolettia retrofracta (thunb.) kies asteraceae pentzia cooperi harv. asteraceae pentzia globosa less. koedoe 45/1 (2002) 50 issn 0075-6458 pond.qxd 2005/12/09 11:34 page 50 asteraceae pentzia incana (thunb.) kuntze us asteraceae pentzia punctata harv. asteraceae pentzia sphaerocephala dc. asteraceae phymaspermum aciculare (e.mey. ex harv.) benth. & hook. ex jackson asteraceae phymaspermum parvifolium (dc.) benth. & hook. ex jackson asteraceae pseudognaphalium luteo-album (l.) hilliard & b.l.burtt asteraceae pseudognaphalium undulatum (l.) hilliard & b.l.burtt asteraceae pteronia adenocarpa harv. asteraceae pteronia erythrochaeta dc. asteraceae pteronia glauca thunb. asteraceae pteronia incana (burm.) dc. asteraceae pteronia staehelinoides dc. asteraceae pulicaria scabra (thunb.) druce asteraceae rosenia humilis (less.) bremer asteraceae schkuhria pinnata (lam.) cabr. * asteraceae senecio achilleifolius dc. asteraceae senecio acutifolius dc. asteraceae senecio agapetes c.jeffrey asteraceae senecio asperulus dc. asteraceae senecio burchellii dc. asteraceae senecio erysimoides dc. k asteraceae senecio incomptus dc. asteraceae senecio juniperinus l.f. var. juniperinus asteraceae senecio lanifer mart. ex c.jeffrey asteraceae senecio leptophyllus dc. asteraceae senecio linifolius l. asteraceae senecio oxyodontus dc. asteraceae senecio radicans (l.f.) sch.bip. asteraceae senecio ruwenzoriensis s.moore asteraceae senecio speciosus willd. asteraceae senecio striatifolius dc. asteraceae senecio vimineus dc. asteraceae sonchus dregeanus dc. asteraceae tarchonanthus camphoratus l. asteraceae tolpis capensis (l.) sch.bip. asteraceae tripteris aghillana dc. var. aghillana asteraceae troglophyton capillaceum (thunb.) hilliard & b.l.burtt asteraceae ursinia montana dc. subsp. apiculata (dc.) prassler asteraceae ursinia nana dc. subsp. nana asteraceae ursinia paleacea (l.) moench asteraceae ursinia trifida (thunb.) n.e.br. forma trifida asteraceae verbesina encelioides (cav.) benth. & hook. * bignoniaceae rhigozum obovatum burch. boraginaceae anchusa capensis thunb. boraginaceae ehretia rigida (thunb.) druce boraginaceae lithospermum cinereum dc. boraginaceae lithospermum diversifolium dc. brassicaceae boscia oleoides (burch. ex dc.) toelken brassicaceae cadaba aphylla (thunb.) wild brassicaceae capsella bursa-pastoris (l.) medik. * brassicaceae cardaria draba (l.) desv. * brassicaceae descurainia sophia (l.) webb ex prantl * brassicaceae erucastrum strigosum (thunb.) o.e.schulz brassicaceae heliophila cornuta sond. var. squamata (schltr.) marais brassicaceae heliophila suavissima burch. ex dc. brassicaceae lepidium africanum (burm. f.) dc. subsp. divaricatum (aiton) jonsell issn 0075-6458 51 koedoe 45/1 (2002) pond.qxd 2005/12/09 11:34 page 51 brassicaceae rorippa fluviatilis (e.mey. ex sond.) thell. brassicaceae sisymbrium burchellii dc. var. burchellii brassicaceae sisymbrium capense thunb. brassicaceae sisymbrium orientale l. * buddlejaceae buddleja glomerata h.l.wendl. buddlejaceae buddleja salviifolia (l.) lam. buddlejaceae gomphostigma virgatum (l.f.) baill. campanulaceae cyphia digitata (thunb.) willd. campanulaceae cyphia sylvatica var. sylvatica eckl. campanulaceae lobelia flaccida subsp. flaccida (c.presl) a.dc. campanulaceae lobelia neglecta roem. & schult. campanulaceae wahlenbergia albens (spreng. ex a.dc.) lammers campanulaceae wahlenbergia androsacea a.dc. us campanulaceae wahlenbergia galpiniae schltr. campanulaceae wahlenbergia neorigida lammers campanulaceae wahlenbergia nodosa (h.buek) lammers campanulaceae wahlenbergia pyrophila lammers campanulaceae wahlenbergia uitenhagensis (h.buek) lammers campanulaceae wahlenbergia undulata (l.f.) a.dc. caryophyllaceae dianthus thunbergii hooper caryophyllaceae silene undulata aiton caryophyllaceae stellaria media (l.) vill. * caryophyllaceae stellaria pallida (dumort.) piré celastraceae gymnosporia buxifolia (l.) szyszyl. celastraceae gymnosporia linearis (l.f.) loes. subsp. linearis celastraceae maytenus polyacantha (sond.) marais celastraceae maytenus undata (thunb.) blakelock commelinaceae commelina africana l. commelinaceae cyanotis speciosa (l.f.) hassk. convolvulaceae convolvulus sagittatus thunb. convolvulaceae cuscuta campestris yunck. * convolvulaceae ipomoea oenotheroides (l.f.) raf. ex hallier f. crassulaceae cotyledon campanulata marloth crassulaceae cotyledon orbiculata l. var. oblonga (haw.) dc. us crassulaceae cotyledon orbiculata l. var. orbiculata crassulaceae cotyledon velutina hook.f. us crassulaceae crassula capitella thunb. subsp. capitella crassulaceae crassula capitella thunb. subsp. thyrsiflora (thunb.) toelken crassulaceae crassula corallina thunb. crassulaceae crassula dependens bolus crassulaceae crassula exilis harv. subsp. cooperi (regel) toelken crassulaceae crassula lanuginosa var. pachystemon harv. us crassulaceae crassula mesembryanthoides (haw.) dietr. crassulaceae crassula muscosa l. var. muscosa crassulaceae crassula obovata haw. crassulaceae crassula perforata thunb. crassulaceae crassula pubescens thunb. subsp. rattrayi (schönland & baker f.) toelken crassulaceae crassula sarcocaulis eckl. & zeyh. crassulaceae crassula tetragona l. subsp. acutifolia (lam.) toelken crassulaceae crassula vaillantii (willd.) roth cucurbitaceae cucumis zeyheri sond. cucurbitaceae kedrostis africana (l.) cogn. cucurbitaceae kedrostis capensis (sond.) a.meeuse dipsacaceae scabiosa columbaria l. dipsacaceae scabiosa incisa mill. ebenaceae diospyros austro-africana de winter ebenaceae diospyros lycioides desf. subsp. lycioides koedoe 45/1 (2002) 52 issn 0075-6458 pond.qxd 2005/12/09 11:34 page 52 ebenaceae diospyros scabrida (harv. ex hiern) de winter ebenaceae diospyros whyteana (hiern) f.white ebenaceae euclea crispa (thunb.) guerke subsp. ovata (burch.) f.white ericaceae erica albens l. var. albens ericaceae erica fuscescens (klotzsch) e.g.h.oliv. ericaceae erica lanata andrews ericaceae erica lehmannii klotzsch k ericaceae erica leucopelta tausch euphorbiaceae clutia pulchella l. var. pulchella euphorbiaceae euphorbia aggregata a.berger var. aggregata euphorbiaceae euphorbia arceuthobioides boiss. euphorbiaceae euphorbia caterviflora n.e.br. euphorbiaceae euphorbia clavarioides boiss. var. clavarioides euphorbiaceae euphorbia mauritanica l. var. mauritanica euphorbiaceae euphorbia micracantha boiss. euphorbiaceae euphorbia ornithopus jacq. k euphorbiaceae euphorbia polycephala marloth v euphorbiaceae euphorbia tridentata lam. fabaceae acacia caffra (thunb.) willd. fabaceae acacia fleckii schinz fabaceae acacia karroo hayne fabaceae argyrolobium collinum eckl. & zeyh. fabaceae argyrolobium pauciflorum eckl. & zeyh. var. pauciflorum fabaceae aspalathus acicularis e.mey. subsp. acicularis fabaceae dichilus gracilis eckl. & zeyh. fabaceae dolichos angustifolius eckl. & zeyh. fabaceae indigofera alpina eckl. & zeyh. fabaceae indigofera alternans dc. var. alternans fabaceae indigofera disticha eckl. & zeyh. fabaceae indigofera mauritanica (l.) thunb. fabaceae indigofera procumbens l. fabaceae indigofera rhytidocarpa benth. ex harv. subsp. rhytidocarpa fabaceae indigofera sessilifolia dc. fabaceae lebeckia macrantha harv. fabaceae lebeckia spinescens harv. fabaceae lessertia carnosa eckl. & zeyh. k fabaceae lessertia depressa harv. fabaceae lessertia frutescens (l.) goldblatt & j.c.manning fabaceae lessertia humilis (e.phillips & r.a.dyer) goldblatt & j.c.manning fabaceae lessertia microphylla (burch. ex dc.) goldblatt & j.c.manning fabaceae lotononis caerulescens e.mey. fabaceae lotononis divaricata (eckl. & zeyh.) benth. fabaceae lotononis laxa eckl. & zeyh. fabaceae lotononis pulchella (e.mey.) b.-e.van wyk fabaceae lotononis pumila eckl. & zeyh. fabaceae lotononis pungens eckl. & zeyh. fabaceae medicago laciniata (l.) mill. var. laciniata fabaceae melolobium candicans (e. mey.) eckl. & zeyh. fabaceae melolobium microphyllum (l.f.) eckl. & zeyh. fabaceae melolobium obcordatum harv. fabaceae tephrosia angulata e. mey. fabaceae tephrosia capensis (jacq.) pers. fabaceae trifolium burchellianum ser. subsp. burchellianum gentianaceae chironia jasminoides l. gentianaceae sebaea elongata e.mey. geraniaceae geranium harveyi briq. geraniaceae geranium incanum burm.f. issn 0075-6458 53 koedoe 45/1 (2002) pond.qxd 2005/12/09 11:34 page 53 geraniaceae monsonia emarginata (l.f.) l’her. geraniaceae pelargonium abrotanifolium (l.f.) jacq. geraniaceae pelargonium alchemilloides (l.) l’her. geraniaceae pelargonium aridum r.a.dyer geraniaceae pelargonium cordifolium (cav.) curtis geraniaceae pelargonium dichondrifolium dc. geraniaceae pelargonium glutinosum (jacq.) l’her. geraniaceae pelargonium grossularioides (l.) l’her. geraniaceae pelargonium minimum (cav.) willd. geraniaceae pelargonium odoratissimum (l.) l’her. geraniaceae pelargonium quercifolium (l.f.) l’her. geraniaceae pelargonium ramosissimum (cav.) willd. geraniaceae pelargonium reniforme curtis geraniaceae pelargonium scabrum (l.) l’her. geraniaceae pelargonium sidoides dc. geraniaceae pelargonium tragacanthoides burch. geraniaceae pelargonium vitifolium (l.) l’her. geraniaceae pelargonium zonale (l.) l’hér. geraniaceae sarcocaulon camdeboense moffett grubbiaceae grubbia rosmarinifolia p.j.bergius subsp. rosmarinifolia haloragaceae gunnera perpensa l. kiggelariaceae kiggelaria africana l. lamiaceae ballota africana (l.) benth. lamiaceae becium burchellianum (benth.) n.e.br. lamiaceae leonotis ocymifolia (burm.f.) iwarsson var. ocymifolia lamiaceae mentha longifolia (l.) huds. subsp. capensis (thunb.) briq. lamiaceae salvia repens burch. ex benth. var. repens lamiaceae salvia stenophylla burch. ex benth. lamiaceae salvia verbenaca l. lamiaceae stachys aethiopica l. lamiaceae stachys cymbalaria briq. lamiaceae stachys dregeana benth. lamiaceae stachys linearis burch. ex benth. lamiaceae stachys rugosa aiton lamiaceae teucrium africanum thunb. linaceae linum thunbergii eckl. & zeyh. loranthaceae moquiniella rubra (a.spreng.) balle malvaceae abutilon sonneratianum (cav.) sweet malvaceae grewia occidentalis l. subsp. occidentalis malvaceae grewia robusta burch. malvaceae hermannia coccocarpa (eckl. & zeyh.) kuntze malvaceae hermannia cuneifolia jacq. var. cuneifolia malvaceae hermannia cuneifolia jacq. var. glabrescens (harv.) i.verd. malvaceae hermannia filifolia l.f. var. filifolia malvaceae hermannia glabrata l.f. malvaceae hermannia linearifolia harv. malvaceae hermannia vestita thunb. malvaceae hibiscus aethiopicus l. malvaceae hibiscus pusillus thunb. malvaceae hibiscus trionum l. malvaceae malva neglecta wallr. * malvaceae malva parviflora var. parviflora l. * melianthaceae melianthus comosus vahl mesembryanthemaceae chasmatophyllum musculinum haw.) dinter & schwantes mesembryanthemaceae delosperma brevisepalum l.bolus var. brevisepalum mesembryanthemaceae delosperma congestum l.bolus koedoe 45/1 (2002) 54 issn 0075-6458 pond.qxd 2005/12/09 11:34 page 54 mesembryanthemaceae delosperma floribundum l.bolus mesembryanthemaceae delosperma frutescens l. bolus mesembryanthemaceae delosperma gramineum l.bolus mesembryanthemaceae delosperma subincanum (haw.) schwantes mesembryanthemaceae drosanthemum hispidum (l.) schwantes mesembryanthemaceae drosanthemum jamesii l.bolus mesembryanthemaceae eberlanzia cradockensis (kuntze) schwantes mesembryanthemaceae eberlanzia ferox (l.bolus) l.bolus mesembryanthemaceae malephora crocea (jacq.) schwantes var. crocea mesembryanthemaceae mestoklema elatum n.e.br. ex glen mesembryanthemaceae phyllobolus splendens (l.) gerbaulet subsp. splendens mesembryanthemaceae psilocaulon coriarium (burch. ex n.e.br.) n.e.br. mesembryanthemaceae psilocaulon junceum (haw.) schwantes mesembryanthemaceae psilocaulon liebenbergii l.bolus mesembryanthemaceae ruschia unidens (haw.) schwantes mesembryanthemaceae ruschia utilis (l.bolus) l.bolus var. giftbergensis l.bolus mesembryanthemaceae trichodiadema barbatum (l.) schwantes mesembryanthemaceae trichodiadema pygmaeum l.bolus r mesembryanthemaceae trichodiadema strumosum (haw.) l.bolus molluginaceae hypertelis salsoloides (burch.) adamson var. salsoloides molluginaceae limeum aethiopicum burm. subsp. aethiopicum molluginaceae pharnaceum trigonum eckl. & zeyh. myrsinaceae myrsine africana l. myrsinaceae rapanea melanophloeos (l.) mez nyctaginaceae boerhavia erecta l. oleaceae olea europaea (l.) subsp. africana (mill.) p.s.green orobanchaceae hyobanche sanguinea l. oxalidaceae oxalis bifurca sond. var. angustiloba lodd. oxalidaceae oxalis bifurca lodd. var. bifurca oxalidaceae oxalis depressa eckl. & zeyh. oxalidaceae oxalis stellata eckl. & zeyh. var. stellata oxalidaceae oxalis stenopetala t.m.salter papaveraceae argemone ochroleuca sweet subsp. ochroleuca * papaveraceae papaver aculeatum thunb. pedaliaceae pterodiscus luridus hook.f. pedaliaceae sesamum capense burm.f. penaeaceae penaea acutifolia a.juss. penaeaceae penaea cneorum meerb. subsp. gigantea r.dahlgren plantaginaceae plantago lanceolata l. * polygalaceae muraltia macrocarpa eckl. & zeyh. polygalaceae polygala ericaefolia dc. polygalaceae polygala garcinii dc. polygalaceae polygala leptophylla burch. var. leptophylla polygalaceae polygala uncinata e.mey. ex meisn. polygalaceae polygala virgata thunb. var. virgata polygonaceae polygonum aviculare l. * polygonaceae rumex lanceolatus thunb. portulacaceae anacampseros arachnoides (haw.) sims ranunculaceae clematis brachiata thunb. ranunculaceae ranunculus multifidus forssk. ranunculaceae thalictrum minus l. rhamnaceae rhamnus prinoides l’her. rosaceae leucosidea sericea eckl. & zeyh. rubiaceae anthospermum rigidum eckl. & zeyh. subsp. rigidum rubiaceae galium capense thunb. subsp. capense rubiaceae galium tomentosum thunb. issn 0075-6458 55 koedoe 45/1 (2002) pond.qxd 2005/12/09 11:34 page 55 rubiaceae nenax microphylla (sond.) t.m.salter rubiaceae rubia petiolaris dc. salicaceae populus alba l. var. alba * salicaceae salix mucronata thunb. subsp. capensis (thunb.) immelman santalaceae thesium galioides a.dc. sapindaceae dodonaea angustifolia l.f. scrophulariaceae aptosimum procumbens (lehm.) steud. var. elongatum (hiern) codd scrophulariaceae aptosimum procumbens (lehm.) steud. var. procumbens scrophulariaceae diascia capsularis benth. scrophulariaceae diascia integerrima benth. us scrophulariaceae hebenstretia dura choisy scrophulariaceae jamesbrittenia atropurpurea (benth.) hilliard scrophulariaceae jamesbrittenia aurantiaca (burch.) hilliard scrophulariaceae jamesbrittenia crassicaulis (benth.) hilliard scrophulariaceae jamesbrittenia filicaulis (benth.) hilliard scrophulariaceae jamesbrittenia fruticosa (benth.) hilliard scrophulariaceae jamesbrittenia pinnatifida (l.f.) hilliard scrophulariaceae jamesbrittenia tysonii (hiern) hilliard scrophulariaceae limosella grandiflora benth. scrophulariaceae manulea crassifolia benth. subsp. crassifolia scrophulariaceae nemesia affinis benth. scrophulariaceae nemesia albiflora n.e.br. scrophulariaceae nemesia fruticans (thunb.) benth. us scrophulariaceae selago albida choisy scrophulariaceae selago corymbosa l. scrophulariaceae selago divaricata l.f. scrophulariaceae selago dolosa hilliard scrophulariaceae selago geniculata l.f. scrophulariaceae selago paniculata thunb. scrophulariaceae selago saxatilis e.mey. scrophulariaceae selago speciosa rolfe scrophulariaceae selago zeyheri rolfe scrophulariaceae sutera halimifolia (benth.) kuntze scrophulariaceae sutera laxiflora (benth.) kuntze scrophulariaceae sutera mollis (benth.) hiern scrophulariaceae veronica anagallis-aquatica l. solanaceae datura stramonium l. * solanaceae lycium cinereum thunb. solanaceae lycium ferocissimum miers solanaceae lycium oxycarpum dunal solanaceae lycium prunus-spinosa dunal solanaceae lycium schizocalyx c.h.wright solanaceae nicotiana glauca graham * solanaceae solanum capense l. solanaceae solanum nigrum l. solanaceae solanum retroflexum dunal solanaceae solanum tomentosum l. solanaceae withania somnifera (l.) dunal thymelaeaceae gnidia microphylla meisn. thymelaeaceae gnidia polycephala (c.a.mey.) gilg thymelaeaceae passerina obtusifolia thoday urticaceae urtica dioica l. * urticaceae urtica urens l. * verbenaceae chascanum pinnatifidum (l.f.) e.mey var. pinnatifidum. verbenaceae lantana rugosa thunb. verbenaceae lippia javanica (burm.f.) spreng. koedoe 45/1 (2002) 56 issn 0075-6458 pond.qxd 2005/12/09 11:34 page 56 verbenaceae verbena tenuisecta briq. * viscaceae viscum continuum e.mey. ex sprague viscaceae viscum rotundifolium l.f. vitaceae cyphostemma quinatum (dryand.) desc. ex wild & r.b.drumm. vitaceae rhoicissus tridentata (l.f.) wild & r.b.drumm. zygophyllaceae tribulus terrestris l. zygophyllaceae zygophyllum incrustatum e.mey. ex sond. complete list selective list total species: 680 total species: 479 total genera: 333 total genera: 227 total families: 87 total families: 62 total red data species: 13 total red data species: 8 issn 0075-6458 57 koedoe 45/1 (2002) pond.qxd 2005/12/09 11:34 page 57 koedoe 45/1 (2002) 58 issn 0075-6458 pond.qxd 2005/12/09 11:34 page 58 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <feff004200720075006700200064006900730073006500200069006e0064007300740069006c006c0069006e006700650072002000740069006c0020006100740020006f0070007200650074007400650020005000440046002d0064006f006b0075006d0065006e0074006500720020006d006500640020006800f8006a006500720065002000620069006c006c00650064006f0070006c00f80073006e0069006e0067002000740069006c0020007000720065002d00700072006500730073002d007500640073006b007200690076006e0069006e0067002000690020006800f8006a0020006b00760061006c0069007400650074002e0020005000440046002d0064006f006b0075006d0065006e007400650072006e00650020006b0061006e002000e50062006e006500730020006d006500640020004100630072006f0062006100740020006f0067002000520065006100640065007200200035002e00300020006f00670020006e0079006500720065002e00200044006900730073006500200069006e0064007300740069006c006c0069006e0067006500720020006b007200e600760065007200200069006e0074006500670072006500720069006e006700200061006600200073006b007200690066007400740079007000650072002e> /nld <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> /esp <feff0055007300650020006500730074006100730020006f007000630069006f006e006500730020007000610072006100200063007200650061007200200064006f00630075006d0065006e0074006f0073002000500044004600200063006f006e0020006d00610079006f00720020007200650073006f006c00750063006900f3006e00200064006500200069006d006100670065006e00200071007500650020007000650072006d006900740061006e0020006f006200740065006e0065007200200063006f007000690061007300200064006500200070007200650069006d0070007200650073006900f3006e0020006400650020006d00610079006f0072002000630061006c0069006400610064002e0020004c006f007300200064006f00630075006d0065006e0074006f00730020005000440046002000730065002000700075006500640065006e00200061006200720069007200200063006f006e0020004100630072006f00620061007400200079002000520065006100640065007200200035002e003000200079002000760065007200730069006f006e0065007300200070006f00730074006500720069006f007200650073002e0020004500730074006100200063006f006e0066006900670075007200610063006900f3006e0020007200650071007500690065007200650020006c006100200069006e0063007200750073007400610063006900f3006e0020006400650020006600750065006e007400650073002e> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 abstract introduction materials and methods results discussion conclusion acknowledgements references about the author(s) marcele vermeulen microbial biochemical and food biotechnology department, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa errol d. cason department of animal, wildlife and grassland sciences, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa wijnand j. swart department of plant sciences, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa citation vermeulen, m., cason, e.d. & swart, w.j., 2020, ‘the rhizobiome of herbaceous plants in clovelly and sterkspruit soils of the stevenson–hamilton supersite’, koedoe 62(2), a1596. https://doi.org/10.4102/koedoe.v62i2.1596 note: special issue: connections between abiotic and biotic components of a granite catena ecosystem in kruger national park, sub-edited by beanelri janecke and johan van tol. original research the rhizobiome of herbaceous plants in clovelly and sterkspruit soils of the stevenson–hamilton supersite marcele vermeulen, errol d. cason, wijnand j. swart received: 26 sept. 2019; accepted: 04 sept. 2020; published: 29 oct. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract by attracting microorganisms from the surrounding soil via root exudates, the composition of microbial populations in the rhizosphere of plants is regulated and maintained according to the genotype of the plant and its abiotic soil environment. this project investigated the bacterial diversity of the rhizosphere microbiome (i.e. rhizobiome) of the three most common herbaceous plants (kyphocarpa angustifolia [amaranthaceae, caryophyllales], melhania acuminata [malvaceae, malvales] and sida cordifolia [malvacae, malvales]) growing mutually in two different soil types (clovelly [top] and sterkspruit [bottom]) with differing abiotic characteristics at a granite catenal supersite in the kruger national park, south africa. two plant species (k. angustifolia and s. cordifolia) occurred at both the top and bottom sites, whilst m. acuminata only occurred at the top site. ten rhizosphere samples were collected per plant from both the top and bottom sites, comprising a total of 50 samples. biolog ecoplates™ were used to assess differences in carbon source utilisation patterns by bacteria in the rhizobiome. for next-generation sequencing analysis, the dna from four randomly selected rhizosphere soil samples from each plant species, at both the top and bottom sites, was combined to yield two samples from each locality for each species. targeted metagenomic sequencing of the 16s rrna gene region (v3 and v4 regions) was used to characterise the rhizobiome. actinobacteria and proteobacteria were the most dominant phyla in all rhizobiomes, and unique and shared operational taxonomic units were identified in all the rhizobiomes. principal component analysis of the biolog data revealed no disparity between the five rhizobiomes. conservation implications: the results obtained in this study could play a role in micro-ecological scale conservation and management because microbial diversity in soils plays a vital role in shaping above-ground biodiversity and terrestrial ecosystem dynamics. keywords: kruger national park; microbiome; rhizobiome; 16s diversity; stevenson–hamilton supersite; granite catena. introduction the rhizosphere is characterised by high microbial activity and an array of complex and dynamic physical, chemical and biological interactions termed the ‘rhizosphere effect’ (helliwell et al. 2019; hiltner 1904). these interactions play a crucial role in plant nutrition (raaijmakers et al. 2009; sørensen 1997) and ecosystem functioning in terrestrial ecosystems (singh et al. 2004). microbes associated with the rhizosphere (i.e. the rhizosphere microbiome or rhizobiome) represent a vast reservoir of microbial diversity that includes fungi, bacteria, nematodes, protozoa, algae, viruses and arthropods (berg & smalla 2009; bonkowski et al. 2000; bonkowski, villenave & griffiths 2009; bulgarelli et al. 2013). plant species actively determine the composition of their rhizobiome by secreting root exudates that attract or repel specific bacteria from the surrounding bulk soil (berg & smalla 2009; doornbos, van loon & bakker 2012). the composition of the rhizobiome is thus determined by numerous edaphic variables of the immediately surrounding bulk soil such as the soil mineralogy, soil moisture content and ph (bardgett et al. 2005; fierer & jackson 2006; myers et al. 2001). however, comparative studies on the effect of specific edaphic variables on the composition of the rhizobiome are rare because of a lack of suitable study sites where the influence of different soil types on the same plant species in close proximity can be compared effectively. the kruger national park (knp), located in the north-eastern part of south africa, is one of the largest natural reserves in africa, with vast areas that are mostly pristine (carruthers 2017). this area is mostly subtropical, with deciduous savanna landscapes inhabited by diverse herbivore and carnivore populations, leading to the integrity of most ecosystems within the park (rughöft et al. 2016). to study these pristine areas, four ‘research supersites’ were chosen (figure 1), each representing unique ecological, geological and climatic features within the park (smit et al. 2013), making these sites ideal to study microbial diversity. figure 1: kruger national park supersites and the stevenson–hamilton granite supersite catena. the stevenson–hamilton supersite (s-hs) in the knp is a granite catena (a hydrologically linked hill slope with dynamic solute, particle and colloid transport systems along the slope) (rughöft et al. 2016) located 7 km south of skukuza (smit et al. 2013). the granite catena of the s-hs is characterised by pronounced differences in soil differentiation along the slope of the catena with sandy soils in the upper region, clay accumulation along the slope and clayey, duplex soils with sporadic sodic patches on the foot slope (levick et al. 2010) (figure 1). the site was shown to have a unique bacterial community when compared to other soil ecosystems (rughöft et al. 2016), and therefore it is ideal in terms of conducting a comparative study of the diversity of the rhizobiome of the same plant species growing in differing soil types. in most soil ecosystems, acidobacteria and (alpha-)proteobacteria are the two dominant phyla, usually accompanied by actinobacteria as another abundant group (janssen 2006; lauber et al. 2009; nacke et al. 2011; prober et al. 2015; rughöft et al. 2016). however, the most abundant phyla in the granite soils from the s-hs were actinobacteria, chloroflexi and firmicutes (rughöft et al. 2016). acidobacteria, planctomycetes, alphaproteobacteria and deltaproteobacteria were the phyla that most frequently occurred at a lower relative abundance (rughöft et al. 2016). the first objective of the present study was to assess the structural and functional diversity of bacteria in the respective rhizobiomes of the three most prevalent herbaceous plant species growing in the s-hs in sandy soil at the top (clovelly) and clay soil at the bottom (sterkspruit) of the slope. a second objective was to determine whether the structural diversity of bacteria in each plant species was influenced by the surrounding bulk soil – such as the sandy soil at the top and clay soil at the bottom of the slope. a third objective was to determine the most dominant bacterial phyla in the two respective sites and to compare this with the findings of rughöft et al. (2016). this is the first study to investigate the diversity of the bacterial rhizobiome of kyphocarpa angustifolia (amaranthaceae, caryophyllales), melhania acuminata (malvaceae, malvales) and sida cordifolia (malvaceae, malvales) as well as to ascertain how different soil types affect microbial diversity. materials and methods sampling site and procedure previous work done in the s-hs has shown that the granitic catena differs in physiochemical properties from the top of the slope to the bottom (figure 1) (sandoval-denis, swart & crous 2018). the soil at the top of the slope is a clovelly soil type characterised by a high percentage of sand (90%), a lower cation exchange capacity (cec) (mean sodium concentration of 1062 mg/kg) and low ph (mean 5.85) (sandoval-denis et al. 2018). the bottom of the slope is characterised by a sterkspruit soil type, with a higher clay content, leading to a higher cec (mean sodium concentration of 3802 mg/kg) and higher ph (mean 6.4) (bouwer, le roux & van tol 2020; janecke et al. 2020; sandoval-denis et al. 2018). three species of plants, k. angustifolia, m. acuminata and s. cordifolia, were identified as the most prominent plant species in the catena as per recommendation by a local botanist (j. du preez, pers. comm.). kyphocarpa angustifolia and m. acuminata are not endemic to south africa, whilst s. cordifolia is an invasive plant species that could have major effects on the microbial communities in soil (van der putten, klironomos & wardle 2007). two species (k. angustifolia and s. cordifolia) occurred at both the top and bottom of the slope, whilst m. acuminata only occurred at the top site (figure 1). all soil samples were collected on 15 march 2015, during a period with very frequent rain occurring in the area. the rhizosphere soil of 10 plants of the same species occurring at each top or bottom site was sampled using a core soil sampler. a global positioning system (gps) waypoint reading was recorded for each plant sampled (table 1). because sampling was initiated on three species at the top of the slope and m. acuminata was not found at the bottom of the slope, a total of 50 soil samples were taken. approximately 200 g of soil from each plant was sampled and placed in a zip-lock plastic bag and kept on ice in an insulated container at approximately 5 °c until analysed in the laboratory. rhizosphere soil samples destined for rhizobiome analysis were stored at −80 °c. table 1: plant species and global positioning system location on granite catena for rhizosphere soil samples analysed by means of biolog ecoplates tm and 16s microbial diversity. community-level physiological profiles biolog ecoplates™ investigate the functional diversity or metabolic potential of that part of a bacterial community that is capable of being metabolically active under axenic conditions. a soil suspension was made for the 50 rhizosphere soil samples by suspending 2 g of each soil sample (table 1) in 200 ml of sterilised water. to totally suspend the soil in water, the samples were shaken for 30 min at room temperature. each well of the microtitre plate was then filled with 90 µl of the soil suspension, and the plates were incubated at 25 °c in a labcon growth chamber in the dark for 96 h (4 days). the change in optical density (od) was then measured using a multiwell plate reader at 590 nm. the overall colour development in each biolog ecoplate™ was expressed as average well colour development (awcd) (zak et al. 1994). data were statistically analysed with xlstat by conducting an analysis of variance (single factor), and then means separation was performed using fischer’s least significant difference (lsd) (p ≤ 0.05). two methods of multivariate analysis were applied to the results: cluster analysis and principal component analysis (pca). rhizosphere microbiome characterisation: dna extraction, bacterial 16s amplicon preparation and illumina miseq sequencing because of cost implications, only four rhizosphere soil samples were chosen randomly from each plant species at both the top and bottom sites, to yield a total of 20 samples (table 1). dna was extracted from soil samples using a nucleospin® soil kit with some modifications from the manufacturer’s instructions. ceramic beads from the nucleospin® bead type a were transferred to a 2-ml clip cap eppendorf tube, and sample lysis was done in a tissue lyser, whereafter the instructions of the manufacturer were followed. ten composite dna samples were then prepared by combining two samples from each locality for each plant species. dna concentrations were determined on a nanodrop 2000 and standardised to 10 ng/µl in 5 mm tris buffer (ph 8.5). a portion of the 16s rrna gene region (v3 and v4 regions) was amplified for each dna sample following a modified version of the illumina miseq 16s metagenomics sequencing library preparation guide (http://support.illumina.com). reactions were performed, in duplicate, in a total volume of 25 µl, consisting of 25 ng dna template, 1.5 µl (0.3 µm) of each primer, 12.5 µl kapa hifi hotstart ready mix (1×) and 7 µl nuclease free water. the polymerase chain reaction (pcr) program included an initial denaturation step of 95 °c for 3 min followed by 25 amplification cycles consisting of 98 °c for 30 s, 65 °c for 30 s and 72 °c for 30 s, followed by a final extension step at 72 °c for 5 min. polymerase chain reaction products were viewed with uv light on 2% agarose gels with gelredtm. the pcr products of each soil sample were combined for downstream analyses at the next generation sequencing facility at the university of the free state, following the illumina miseq 16s metagenomics sequencing library preparation guide for the 2 × 300 bp miseq v3 kit (http://support.illumina.com). the quality of sequences was assessed using fastqc (andrews 2010), and the sequences were filtered and trimmed using prinseq-lite v0.20.4 (schmieder & edwards 2011) to have a mean quality score (qc) of >20 and a sequence length of >200 bp. paired-end reads were merged using paired-end read merger (pear) v0.9.6 (zhang et al. 2014). chimeric sequences were identified using usearch v6.1.544 (edgar 2010) against the ribosomal database project (rdp) ‘gold’ database (edgar 2010). the quantitative insight into microbial ecology (qiime) v1.9.1 framework was used for subsequent sequence data analyses (caporaso et al. 2010; lozupone et al. 2010). a closed-reference operational taxonomic unit (otu) picking workflow was followed, and otus were picked against the silva database v132 at 97% sequence identity. operational taxonomic unit tables were normalised, and all analyses were carried out using r (www.r-project.org). operational taxonomic unit richness and phylogenetic diversity (pd) were calculated using phyloseq (mcmurdie & holmes 2013) in r (team rdc 2011). the potential functional capacity of the otus was predicted using functional annotation of prokaryotic taxa (faprotax) (louca, parfrey & doebeli 2016). the taxonomic and functional structures of the microbial community were visualised using principal coordinate analysis (pcoa). permutational analysis of variance (permanova) was used to test for differences in composition and function between and within habitats, whereas permutation dispersion (permdisp) was used to test for differences in habitat dissimilarity; both analyses were performed with the ‘adonis’ and ‘betadisper’ functions in vegan (oksanen et al. 2013) for r. ethical considerations ethical approval for the multidisciplinary project as a whole was obtained from the interfaculty animal ethics committee at the university of the free state (ufs-aed2019/0121). the sanparks permit numbers for the collection of soil for lab analyses and vegetation for identification purposes are sk069, sk2095 and sk054. results community-level physiological profiles principal component analysis and awcd data (mean of the absorbance values for a whole plate) revealed no disparity between the rhizosphere samples from k. angustifolia and s. cordifolia from the sterkspruit site (bottom) and those of k. angustifolia, m. acumi nata and s. cordifolia from the clovelly site (top) (pca data not shown). based on the awcd data, after 96 h there were also no significant differences (p > 0.05) between the five samples (figure 2). figure 2: carbon substrate utilisation activity as average well colour development by the rhizosphere microbial community. rhizosphere microbiome characterisation: dna extraction, bacterial 16s amplicon preparation and illumina miseq sequencing a total of 171 091 sequences were obtained from the 16s library after quality assessment and data filtering ranging from 62 246 (max) sequences to 5955 (min) sequences, with 6223 to 2119 otus (97% similarity cut-off). rarefaction curves (not shown) indicated that deeper sequencing is needed to completely resolve the prokaryotic diversity of the samples. after rarefication there were 3374 sequences per sample with a total of 3952 otus (97% similarity cut-off) for all samples. of the total number of otus, 581 (representing 4.78% of the total number of sequences) were unique to k. angustifolia at the clovelly site (top), 313 (2.02%) were unique to m. acuminata at the clovelly site (top), 263 (1.72%) were unique to s. cordifolia at the clovelly site (top), 234 (1.80%) were unique to k. angustifolia at the sterkspruit site (bottom), 332 (1.63%) were unique to s. cordifolia at the sterkspruit site (bottom) and 315 (47.72%) were shared between all the rhizosphere samples. percentage relative abundance plots (figure 3) showed that the phyla actinobacteria (11.74% – 36.22%) and proteobacteria (10.23% – 30.79%) were amongst the most dominant phyla in all rhizosphere samples. in contrast, the phylum verrucomicrobia was one of the dominant phyla in the rhizosphere samples, from one of the k. angustifolia (22.67%) samples from the sterkspruit site (bottom) and both the samples from k. angustifolia (19.71%, 18.94%), m. acuminata (42.77%, 26.08%) to s. cordifolia (29.31%, 25.49%) from the clovelly site (top). planctomycetes was one of the dominant phyla in the second k. angustifolia (47.69%) rhizosphere samples from the sterkspruit site (bottom) and firmicutes (19.85%) and chloroflexi (17.13%), respectively, in the s. cordifolia rhizosphere samples from the sterkspruit site (bottom) (figure 3). figure 3: taxonomic identification. distinct prokaryotic communities (otu level) were not observed between the five rhizosphere samples, k. angustifolia and s. cordifolia from the sterkspruit site (bottom) to k. angustifolia, m. acuminata and s. cordifolia from the clovelly site (top) (permanova f4.9 = 0.96, r2 = 43%, p > 0.1). however, the pcoa showed that samples from the clovelly site (top) clustered together (figure 4). the functional capacity of the otus (figure 5) showed a distinct association between the five rhizosphere samples, k. angustifolia and s. cordifolia from the sterkspruit site (bottom) and k. angustifolia, m. acuminata and s. cordifolia from the clovelly site (top) (permanova f1.9 = 1.72, r2 = 18%, p ≤ 0.1). samples from the clovelly (top) site clustered together (figure 6). overall, the rhizosphere communities were dominated by chemoheterotrophy (20.05% – 32.25%) and aerobic chemoheterotrophy (16.35% – 30.34%) (figure 5). although present at low percentages of the total rhizosphere community, sulphur respiration (0.01% – 0.18%), iron respiration (0.05% – 0.12%), sulphite respiration (0.01% – 0.12%) and thiosulphate respiration (0.01% – 0.12%) were only present at the sterkspruit site (bottom) for both k. angustifolia and s. cordifolia (figure 5). in contrast, arsenate respiration (0.02% – 0.06%), dissimilatory arsenate reduction (0.02% – 0.06%), nitrate ammonification (0.02% – 0.06%) and nitrite ammonification (0.02% – 0.06%) were only present in k. angustifolia at the sterkspruit site (bottom) and dark hydrogen oxidation (0.05% – 0.08%) and reductive acetogenesis (0.05% – 0.08%) only in s. cordifolia at the sterkspruit site (bottom) (figure 5). figure 4: principal coordinates analysis of the rhizosphere microbiome for kyphocarpa angustifolia and sida cordifolia from the sterkspruit site (bottom) and k. angustifolia, melhania acuminata and s. cordifolia from the clovelly site (top). figure 5: potential functional capacity for each sample. discussion the rhizobiome is shaped by complex interactions between the soil type, plant species and root zone location (marschner et al. 2001). although there were no statistically significant differences between the rhizobiome of the clovelly (top) and sterkspruit (bottom) soil types or between the three different plant species, it was evident that the soil type and plant species did have a role in shaping the rhizobiome, as unique otus were observed in all five different rhizosphere samples. this has previously been observed, with soil type (bachman & kinzel 1992) and plant species (wieland, neumann & backhaus 2001) both being reported as the driving factor for microbial soil and rhizosphere diversity. although the sampling area spanned only 200 m, differences in plant diversity were observed in previous studies of the plant diversity of the s-hs granite catena (theron, van aardt & du preez 2020). this was also reflected in the bulk soil microbiome diversity of the s-hs granite catena (rughöft et al. 2016) along the slope and is thus expected for the rhizobiome of the resident plants. there was, however, a significant amount of shared otus (47.72% of the total number of sequences) regardless of the soil type and specific plant rhizosphere they occurred in. the invasive plant, s. cordifolia, is known to have antimicrobial properties (pallela et al. 2018; thangavel, raveendran & kathirvel 2006); however, there is no information available on its rhizobiome. similarly, now information is available for the rhizobiome of the two native plants, k. angustifolia and m. acuminata. the soil surrounding the rhizosphere serves as the main reservoir for the microorganisms that form part of the rhizobiome (berg & smalla 2009). actinobacteria and proteobacteria were amongst the most dominant phyla in all rhizosphere samples, whilst verrucomicrobia and planctomycetes were the dominant phyla in the rhizosphere samples from k. angustifolia at the sterkspruit site (bottom), verrucomicrobia in k. angustifolia, m. acuminata and s. cordifolia at the clovelly site (top) and firmicutes and chloroflexi in the s. cordifolia rhizosphere samples from the sterkspruit site (bottom) (figure 3). previous studies on the soil in the knp s-hs showed that actinobacteria, chloroflexi and firmicutes were dominant, whilst planctomycetes, proteobacteria and verrucomicrobia most frequently had the lowest relative abundance (rughöft et al. 2016). therefore, although the abundance of specific bacterial phyla may differ between the rhizobiome and bulk soil (rughöft et al. 2016), our results are consistent with those of berg and smalla (2009), who suggested that root exudates in the rhizosphere select specific microbes from the surrounding bulk soil. to our knowledge, there are no studies comparing the bacterial rhizobiome of native and invasive plants within the study site. gryzenhout et al. (2020), however, studied the fungal rhizobiome of the same plant species in s-hs and found that s. cordifolia only attracted half the number of otus compared to the native plants, m. acuminata and k. angustifolia, which could be attributed to the antimicrobial properties of s. cordifolia. however, this was not observed for the bacterial rhizobiome. actinobacteria, chloroflexi, firmicutes and verrucomicrobia are well adapted to survival in the nutrientand water-poor soils of the knp s-hs (rughöft et al. 2016). the increase in abundance of proteobacteria in the rhizosphere compared to the surrounding soil could be because they have been shown to benefit from the higher nutrient levels, especially carbon availability (leff et al. 2015; wang, ji & gao 2016), that will be expected because of root exudates in the rhizosphere (haichar et al. 2014). planctomycetes and verrucomicrobia, although not dominant in all samples, were present (>1%) in all samples and have been reported in other studies on rhizobiomes (gkarmiri et al. 2017; zeng et al. 2018; zhou et al. 2019). similar to proteobacteria, planctomycetes and verrucomicrobia could benefit from the increased availability of nutrients in the rhizosphere. planctomycetes are known to play a role in carbohydrate metabolism (fuerst 1995), and a lineage in the planctomycetes has also been shown to play a role in anaerobic oxidation of ammonium (strous et al. 1999). planctomycetes present in the rhizosphere and bulk soil samples from the s-hs granite catena could therefore play a role in the oxidation of ammonium reported in this soil (rughöft et al. 2016). according to the bacterial functional group analyses, soil type (i.e. clovelly vs. sterkspruit) (figure 6) played a role in the composition of the bacterial functional groups. these two soil types present different environments for the rhizospheres. clovelly soils have low fertility and water-holding capacity, when compared with those of other soils of the same clay content, which would affect the rhizosphere diversity and thus functionality when compared to the sterkspruit soil type. the sterkspruit soil type is characterised by a higher clay content, leading to a higher cec. the most important factors pertaining to these soils that affect the rhizosphere biome are the elevated sodium concentration in the sterkspruit soil type (3802 mg/kg vs. 1062 mg/kg) and more acidic ph in the clovelly soil type (5.85 vs. 6.4) (sandoval-denis et al. 2018; theron et al. 2020). higher sodium could lead to lower osmotic potential, making it more difficult for microbes and plant roots to extract water from soils (yan et al. 2015). differences in ph would also mediate nutrient availability in soils (zhalnina et al. 2015), affecting the rhizobiome. figure 6: principal coordinates analysis of the functional capacity for the rhizosphere microbiome for kyphocarpa angustifolia and sida cordifolia from the sterkspruit site (bottom) and k. angustifolia, melhania acuminata and s. cordifolia from the clovelly site (top). chemoheterotrophy and aerobic chemoheterotrophy were the most abundant identified functions in all rhizobiomes (figure 5). however, this is expected for bacterial communities that use organic carbon as the main food source (heterotrophic bacteria) (zhang et al. 2019). in the absence of oxygen, a wide range of compounds can serve as terminal electron donors (anaerobic respiration). the availability of oxygen in the rhizosphere is one of the most changing factors and anaerobic/anoxic conditions may develop (philippot et al. 2013), leading to bacteria adapting to using various anaerobic respiration strategies. this was observed with iron, sulphur, sulphite and thiosulphate respiration, present at the sterkspruit site (bottom) in both k. angustifolia and s. cordifolia, and arsenate respiration in k. angustifolia (figure 5). it is possible that the higher clay content of the sterkspruit soil could also play a role, as anaerobic respiration was only observed in the rhizobiomes associated with this particular soil type. conclusion soil type (clovelly vs. sterkspruit) and differences in ph and salinity (figure 1) in the two soil types played a role in the rhizobiome bacterial diversity, as unique otus could be identified in all respective rhizosphere soil samples. there was, however, also a large overlap in bacterial diversity in the different soil types and in the rhizobiome of the three plant species. soil type also played a significant role in the bacterial functional groups, with anaerobic respiration only being reported in the sterkspruit soil. acknowledgements the university of the free state (ufs) strategic research fund is acknowledged for funding the multidisciplinary research. mercia coetzee (central university of technology, bloemfontein, south africa) is thanked for her technical support in the field. eddie riddell and navashni govender (sanparks) are acknowledged for their research support in the kruger national park. dr jan-g vermeulen (microbial biochemical and food biotechnology department, university of the free state, bloemfontein, south africa) is thanked for his assistance with graphics. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions m.v., e.d.c. and w.j.s. contributed equally to this article. the authors directly participated in the study design, execution and interpretation of the research. funding information this research received no specific grant from any funding agency in the public, commercial or not-for-profit sectors. data availability data sharing is not applicable to this article as no new data were created or analysed in this article. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references andrews, s., 2010, fastqc: a quality control tool for high throughput sequence data, viewed 01 july 2019, from http://www/bioinformatics.babraham.ac.uk/projects/fastqc. bachman, g. & h. kinzel., 1992, ‘physiological and ecological aspects of the interactions between plant roots and rhizosphere soil’, soil biology and biochemistry 24(6), 5543–5552. https://doi.org/10.1016/0038-0717(92)90079-d bardgett, r.d., bowman, w.d., kaufmann, r. & schmidt, s.k., 2005, ‘a temporal approach to linking aboveground and belowground ecology’, trends in ecology and evolution 20(11), 634–641. https://doi.org/10.1016/j.tree.2005.08.005 berg, g. & smalla, k., 2009, ‘plant species and soil type cooperatively shape the structure and function of microbial communities in the rhizosphere’, fems microbial ecology 68(1), 1–13. https://doi.org/10.1111/j.1574-6941.2009.00654.x bonkowski, m., cheng, w., griffiths, b.s., alphei, j. & scheu, s., 2000, ‘microbial-faunal interactions in the rhizosphere and effects on plant growth’, european journal of soil biology 36(3–4), 135–147. https://doi.org/10.1016/s1164-5563(00)01059-1 bonkowski, m., villenave, c. & griffiths, b., 2009, ‘rhizosphere fauna: the functional and structural diversity of intimate interactions of soil fauna with plant roots’, plant soil 321, 213–233. https://doi.org/10.1007/s11104-009-0013-2 bouwer, d., le roux, p.a.l. & van tol, j., 2020, ‘identification of hydropedological flowpaths in stevenson-hamilton catena from soil morphological, chemical and hydraulic properties’, koedoe 62(2), a1584. https://doi.org/10.4102/koedoe.v62i2.1584 bulgarelli, d., schlaeppi, k., spaepen, s., ver loren van themaat, e. & schulze-lefert, p., 2013, ‘structure and functions of the bacterial microbiota of plants’, annual review of plant biology 64, 807–838. https://doi.org/10.1146/annurev-arplant-050312-120106 caporaso, j.g., kuczynski, j., stombaugh, j., bittinger, k., bushman, f.d. & costello, e.k. et al., 2010, ‘qiime allows analyses of high-throughput community sequencing data’, nature methods 7, 335–336. https://doi.org/10.1038/nmeth.f.303 carruthers, j., 2017, national park science: a century of research in south africa (ecology, biodiversity and conservation), cambridge university press, cambridge. doornbos, r.f., van loon, l.c. & bakker, p.a.h.m., 2012, ‘impact of root exudates and plant defence signalling on bacterial communities in the rhizosphere: a review’, agronomy for sustainable development 32, 227–243. https://doi.org/10.1007/s13593-011-0028-y edgar, r.c., 2010, ‘search and clustering orders of magnitude faster than blast’, bioinformatics 26(19), 2460–2461. https://doi.org/10.1093/bioinformatics/btq461 fierer, n. & jackson, r.b., 2006, ‘the diversity and biogeography of soil bacterial communities’, proceedings of the national academy of sciences of the united states of america 103(3), 626–631. https://doi.org/10.1073/pnas.0507535103 fuerst, j.a., 1995, ‘the planctomycetes – emerging models for microbial ecology, evolution and cell biology’, microbiology 141(7), 1493–1506. https://doi.org/10.1099/13500872-141-7-1493 gkarmiri, k., mahmood, s., ekblad, a., alström, s., högberg, n. & finlay, r., 2017, ‘identifying the active microbiome associated with roots and rhizosphere soil of oilseed rape’, applied and environmental microbiology 83(22), e01938–17. https://doi.org/10.1128/aem.01938-17 gryzenhout, m., cason, e.d., vermeulen, m., kloppers, g.a.e., bailey, b. & ghosh, s., 2020, ‘fungal community structure variability between the root rhizosphere and endosphere in a granite catena system in kruger national park, south africa’, koedoe 62(2), a1597. https://doi.org/10.4102/koedoe.v62i2.1597 haichar, f.e.z., santaella, c., heulin, t. & achouak, w., 2014, ‘root exudates mediated interactions belowground’, soil biology and biochemistry 77, 69–80. https://doi.org/10.1016/j.soilbio.2014.06.017 helliwell, j.r., sturrock, c.j., miller, a.j., whalley, w.r. & mooney, s.j., 2019, ‘the role of plant species and soil condition in the structural development of the rhizosphere’, plant cell and environment 42(6), 1974–1986. https://doi.org/10.1111/pce.13529 hiltner, l., 1904, ‘uber neuere erfahrungen und probleme auf dem gebiete der bodenbakteriologie unter besonderden berucksichtigung und brache’, arbeiten der deutschen landwirtschaftlichen gesellschaf 98, 59–78. janecke, b.b., van tol, j., smit, i.p.j., van aardt, a.c., riddell, e.s., seaman, m.t. et al., 2020, ‘biotic and abiotic connections on a granitic catena: framework for multidisciplinary research’, koedoe 62(2), a1600. https://doi.org/10.4102/koedoe.v62i2.1600 janssen, p.h., 2006, ‘identifying the dominant soil bacterial taxa in libraries of 16s rrna and 16s rrna genes’, applied and environmental microbiology 72(3), 1719–1728. https://doi.org/10.1128/aem.72.3.1719-1728.2006 lauber, c.l., hamady, m., knight, r. & fierer, n., 2009, ‘pyrosequencingbased assessment of soil ph as a predictor of soil bacterial community structure at the continental scale’, applied and environmental microbiology 75(15), 5111–5120. https://doi.org/10.1128/aem.00335-09 leff, j.w., jones, s.e., prober, s.m., barberan, a., borer, e.t. & firn, j.l. et al., 2015, ‘consistent response of soil microbial communities to elevated nutrient inputs in grasslands across the globe’, proceedings of the national academy of sciences of the united states of america 112(35), 10967–10972. https://doi.org/10.1073/pnas.1508382112 levick, s.r., asner, g.p., chadwick, o.a., khomo, l.m., rogers, k.h. & hartshorn, a.s. et al., 2010, ‘regional insight into savanna hydrogeomorphology from termite mounds’, nature communications 1, 65. https://doi.org/10.1038/ncomms1066 louca, s., parfrey, l.w. & doebeli, m., 2016, ‘decoupling function and taxonomy in the global ocean microbiome’, science 353(6305), 1272–1277. https://doi.org/10.1126/science.aaf4507 lozupone, c.a., mcdonald, d., muegge, b.d., pirrung, m., reeder, j. & sevinsky, j.r. et al., 2010, ‘qiime allows analyses of high-throughput community sequencing data’, nature methods 7, 335–336. https://doi.org/10.1038/nmeth.f.303 marschner, p., yang, c.h., lieberei, r. & crowley, d.e., 2001, ‘soil and plant specific effects on bacterial community composition in the rhizosphere’, soil biology and biochemistry 33(11), 1437–1445. https://doi.org/10.1016/s0038-0717(01)00052-9 mcmurdie, p.j. & holmes, s., 2013, ‘phyloseq: an r package for reproducible interactive analysis and graphics of microbiome census data’, plos one 8(4), e61217. https://doi.org/10.1371/journal.pone.0061217 myers, r.t.m., zak, d.r., white, d.c. & peacock, a., 2001, ‘landscape-level patterns of microbial community composition and substrate use in upland forest ecosystems’, soil science society of america journal 65(2), 359–367. https://doi.org/10.2136/sssaj2001.652359x nacke, h., thürmer, a., wollherr, a., will, c., hodac, l. & herold, n. et al., 2011, ‘pyrosequencing-based assessment of bacterial community structure along different management types in german forest and grassland soils’, plos one 6(2), e17000. https://doi.org/10.1371/journal.pone.0017000 oksanen, j., blanchet, f.g., kindt, r., legendre, p., minchin, p.r. & o’hara, r.b. et al., 2013, vegan: community ecology package, r package version 2.0-2, 01 viewed july 2019, from http://cran.r-project.org/package=vegan. pallela, p.n.v.k., ummey, s., ruddaraju, l.k., pammi, s.v.n. & yoon, s.-g., 2018, ‘ultra small, mono dispersed green synthesized silver nanoparticles using aqueous extract of sida cordifolia plant and investigation of antibacterial activity’, microbial pathogenesis 124, 63–69. https://doi.org/10.1016/j.micpath.2018.08.026 philippot, l., raaijmakers, j.m., lemanceau, p. & van der putten, w.h., 2013, ‘going back to the roots: the microbial ecology of the rhizosphere’, nature review microbiology 11(11), 789–799. https://doi.org/10.1038/nrmicro3109 prober, s.m., leff, j.w., bates, s.t., borer, e.t., firn, j. & harpole, w.s. et al., 2015, ‘plant diversity predicts beta but not alpha diversity of soil microbes across grasslands worldwide’, ecology letters 18(1), 85–95. https://doi.org/10.1111/ele.12381 raaijmakers, j.m., paulitz, t.c., steinberg, c., alabouvette, c. & moenne-loccoz, y., 2009, ‘the rhizosphere: a playground and battlefield for soilborne pathogens and beneficial microorganisms’, plant soil 321, 341–361. https://doi.org/10.1007/s11104-008-9568-6 rughöft, s., herrmann, m., lazar, c.s., cezar, s., levick, s.r. & trumbore, s.e. et al., 2016, ‘community composition and abundance of bacterial, archaeal and nitrifying populations in savana soils of contrasting bedrock material in kruger national park, south africa’, frontiers in microbiology 7, 1638. https://doi.org/10.3389/fmicb.2016.01638 sandoval-denis, m., swart, w.j. & crous, p.w., 2018, ‘new fusarium species from the kruger national park, south africa’, mycokeys 34, 63–92. https://doi.org/10.3897/mycokeys.34.25974 schmieder, r. & edwards, r., 2011, ‘quality control and preprocessing of metagenomic datasets’, bioinformatics 27(6), 863–864. https://doi.org/10.1093/bioinformatics/btr026 singh, b.k., millard, p., whiteley, a.s. & murrell, j.c., 2004, ‘unravelling rhizosphere-microbial interactions: opportunities and limitations’, trends in microbiology 12(8), 386–393. https://doi.org/10.1016/j.tim.2004.06.008 smit, i.p.j., riddell, s.e., cullum, c. & petersen, r., 2013, ‘kruger national park research supersites: establishing long-term research sites for cross-disciplinary, multiscaled learning’, koedoe 55(1), 1107. https://doi.org/10.4102/koedoe.v55i1.1107 sørensen, j., 1997, ‘the rhizosphere as a habitat for soil microorganisms’, in j.d. van elsas, j.t. trevors & e.m.h. wellington (eds.), modern soil microbiology, pp. 21–45, marcel dekker inc., new york, ny. strous, m., fuerst, j.a., kramer, e.h.m., logemann, s., muyzer, g. & van de pas-schoonen, k.t. et al., 1999, ‘missing lithotroph identified as new planctomycete’, nature 400, 446–449. https://doi.org/10.1038/22749 team rdc, 2011, ‘r: a language and environment for statistical computing’, r foundation for statistical computing, viewed 01 july 2019, from http://www.r-project.org/. thangavel, m., raveendran, m. & kathirvel, m., 2006, ‘a comparative study on the effect of plant extracts with the antibiotics on organisms of hospital origin’, ancient science of life 26(1), 65–72. theron, e., van aardt, a.c. & du preez, p.j., 2020, ‘vegetation distribution along a granite catena, southern kruger national park’, koedoe 62(2), a1588. https://doi.org/10.4102/koedoe.v62i2.1588 van der putten, w.h., klironomos, j.n. & wardle, d.a., 2007, ‘microbial ecology of biological invasions’, the isme journal 1, 28–37. https://doi.org/10.1038/ismej.2007.9 wang, y., ji, h. & gao, c., 2016, ‘differential response of soil bacterial taxa to long-term p, n and organic manure application’, journal of soils and sediments 16, 1046–1058. https://doi.org/10.1007/s11368-015-1320-2 wieland, g., neumann, r. & backhaus, h., 2001, ‘variation of microbial communities in soil, rhizosphere, and rhizoplane in response to crop species, soil type, and crop development’, applied and environmental microbiology 67(12), 5849–5854. https://doi.org/10.1128/aem.67.12.5849-5854.2001 yan, n., marschner, p., cao, w., zuo, c. & qina, w., 2015, ‘influence of salinity and water content on soil microorganisms’, international soil and water conservation research 3(4), 316–23. https://doi.org/10.1016/j.iswcr.2015.11.003 zak, j.c., willig, m.r., moorhead, d.l. & wildman, h.g., 1994, ‘functional diversity of microbial communities: a quantitative approach’, soil biology and biochemistry 26(9), 1101–1108. https://doi.org/10.1016/0038-0717(94)90131-7 zeng, m., zhong, y., cai, s. & diao, y., 2018, ‘deciphering the bacterial composition in the rhizosphere of baphicacanthus cusia (nees) bremek’, scientific reports 8, 15831. https://doi.org/10.1038/s41598-018-34177-1 zhalnina, k., dias, r., de quadros, p.d., davis-richardson, a., camargo, f.a. & clark, i.m. et al., 2015, ‘soil ph determines microbial diversity and composition in the park grass experiment’, microbial ecology 69, 395–406. https://doi.org/10.1007/s00248-014-0530-2 zhang, j., kobert, k., flouri, t. & stamatakis, a., 2014, ‘pear: a fast and accurate illumina paired-end read merger’, bioinformatics 30(5), 614–620. https://doi.org/10.1093/bioinformatics/btt593 zhang, x.w., qu, y.y., ma, q., li, s.z., dai, c.x. & lian, s.y. et al., 2019, ‘performance and microbial community analysis of bioaugmented activated sludge system for indigo production from indole’, applied biochemistry and biotechnology 187, 1437–1447. https://doi.org/10.1007/s12010-018-2879-z zhou, d., jing, j., chen, y., wang, f., qi, d. & feng, r. et al., 2019, ‘deciphering microbial diversity associated with fusarium wilt-diseased and disease-free banana rhizosphere soil’, bmc microbiology 19, 161. https://doi.org/10.1186/s12866-019-1531-6 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 filelist convert a pdf file! page 1 page 2 abstract introduction research methods and design results discussion conclusions acknowledgements references about the author(s) claudine roos unit for environmental sciences and management, faculty of natural and agricultural sciences, north-west university, potchefstroom, south africa reece c. alberts unit for environmental sciences and management, faculty of natural and agricultural sciences, north-west university, potchefstroom, south africa francois p. retief unit for environmental sciences and management, faculty of natural and agricultural sciences, north-west university, potchefstroom, south africa dirk p. cilliers unit for environmental sciences and management, faculty of natural and agricultural sciences, north-west university, potchefstroom, south africa william hodgson sabi sand pfunanani trust, sabi sand wildtuin, skukuza, south africa iain olivier sabi sand pfunanani trust, sabi sand wildtuin, skukuza, south africa citation roos, c., alberts, r.c., retief, f.p., cilliers, d.p., hodgson, w. & olivier, l., 2022, ‘challenges and opportunities for sustainable solid waste management in private nature reserves: the case of sabi sand wildtuin, south africa’, koedoe 64(1), a1710. https://doi.org/10.4102/koedoe.v64i1.1710 original research challenges and opportunities for sustainable solid waste management in private nature reserves: the case of sabi sand wildtuin, south africa claudine roos, reece c. alberts, francois p. retief, dirk p. cilliers, william hodgson, iain olivier received: 12 jan. 2022; accepted: 14 apr. 2022; published: 26 may 2022 copyright: © 2022. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the mismanagement of waste in protected areas may lead to significant and irreversible environmental, economic and social impacts, such as land degradation, resource depletion, surface and groundwater pollution, loss of biodiversity and impacts on the aesthetic value of these areas. this paper aims to identify the challenges and opportunities for sustainable solid waste management in privately protected areas, given the limited research conducted on this topic. a case study approach was followed, which focused on the sabi sand wildtuin, a private nature reserve (pnr) in south africa. interviews were conducted with 30 participants, which included representatives from the management authority, commercial lodges, non-commercial properties and a waste service provider. several challenges have been identified by interviewees. behaviour was the most frequently mentioned challenge, where interviewees raised concerns about negative attitudes, unwillingness to implement waste management measures and a possible lack of support. other frequently mentioned challenges included foreseen difficulties due to the size and location of the reserve and concerns around funding of waste management measures, especially given the financial implications of coronavirus disease 2019 (covid-19) travel and tourism restrictions. the most frequently identified opportunities included creating jobs and improving livelihoods, providing assurance and transparency of what happens to waste ‘beyond the gate’ and improving awareness, knowledge and skills related to waste management. to optimise the opportunities towards sustainable solid waste management, pnrs should focus on aligning their strategic direction to achieve legal compliance and support community initiatives to establish waste-related infrastructure and services that cannot be implemented within the reserve. conservation implications: the pursuit of waste-related opportunities within privately protected areas could enhance the implementation of sustainable solid waste management in pnrs, whilst also contributing to pollution prevention, community upliftment and other secondary benefits, which could ultimately result in increased conservation efforts. keywords: sustainable solid waste management; protected areas; private nature reserves; challenges; opportunities; sabi sand wildtuin. introduction protected areas are of international importance due to their natural, historical, cultural and ecotourism value (alberts et al. 2021; sandbrook et al. 2019). ecotourism in protected areas has rapidly increased globally, which has several adverse impacts, including increased waste generation (rodríguez-rodríguez 2012). the mismanagement of waste in protected areas may lead to significant and irreversible environmental, economic and social impacts, such as land degradation, resource depletion, surface and groundwater pollution, loss of biodiversity and impacts on the aesthetic value of tourism locations (dunjić et al. 2017; steg & vlek 2009). hong and chan (2010:140) have identified solid waste management as the ‘most severe threat’ affecting penang national park in malaysia. this was mainly due to the irresponsible disposal of waste, absence of formal waste management policies and systems and limited environmental and waste-related awareness amongst visitors. in south africa, research by du plessis, van der merwe and saayman (2013) which focused on tourists’ perceptions on the environmental friendliness of south african national parks found that ‘waste management and the recycling of waste in national parks require attention’. sustainable waste management is suggested as an important component of sustainable ecotourism indicator and rating systems (li 2004; van der merwe, saayman & bothma 2017), and it also ranks amongst the top responsible tourism expectations of visitors to national parks (de witt 2015; du plessis et al. 2013; morrison-saunders et al. 2019). chang and pires (2015) argue that the adverse impacts of waste can be mitigated, or entirely avoided, through sustainable solid waste management. sustainable solid waste management is explained as the management of waste in such a manner that ‘resources are sufficient to fulfil the demand for daily consumption whilst guaranteeing ecosystem sustainability by using appropriate waste collection, handling, reuse, recycling and resource conservation’ (chang & pires 2015:12), whilst also providing for multiple triple bottom line dimensions, including social, environmental and economic aspects (yadav & karmakar 2020). limited research has been done on the challenges and opportunities of achieving sustainable solid waste management in protected areas, specifically within the developing country context (przydatek 2019). to this end, this paper aims to identify the challenges and opportunities for sustainable solid waste management in privately protected areas, following a case study approach by focusing on a private nature reserve (pnr) in south africa. mitchell et al. (2018) and przydatek (2019) highlight the importance of privately protected areas such as pnrs in contributing to conservation and protected area strategies, whilst the importance of responsible waste management in pnrs has been highlighted by roos et al. (2021). this research builds on the work of roos et al. (2021) that investigated waste behaviour in pnrs through the application of the theory of planned behaviour, using a south african pnr, sabi sand wildtuin (ssw), as a case study. this research employed surveys where ordinal scales were used to indicate the level of agreement or disagreement with waste behaviour and practice statements. the study did not investigate the reasons for specific responses; follow-up research was thus recommended by the authors. the scope of this research therefore expands on the initial research of roos et al. (2021) by identifying challenges and opportunities for sustainable solid waste management in pnrs. research methods and design this research follows an exploratory research design following a case study approach. qualitative data was collected by conducting interviews with various waste management stakeholders within a selected south african pnr. purposive sampling was used to target the respondents who participated in the earlier survey done by roos et al. (2021). the case study: sabi sand wildtuin the ssw was considered as a suitable case study to explore challenges and opportunities for sustainable solid waste management in pnrs because it is a well-established pnr and the management authority is currently in the process of developing an integrated waste management strategy for the reserve. understanding the challenges and opportunities for waste management, as perceived by different stakeholders, would provide valuable insights towards implementation of this strategy. the ssw, which is located in the greater kruger national park, is the oldest private game reserve in south africa (figure 1), with internationally recognised high-end luxury tourist products and infrastructure resulting in waste management challenges (roos et al. 2021). the reserve includes 24 commercial properties with lodges and 30 non-commercial private properties. a total of 32 lodges are located on the commercial properties. the estimated total amount of waste generated by ssw per year is approximately 180 tonnes and consists of mainly general waste (such as plastic, glass, paper and packaging and metals) and organic waste (such as food waste and garden waste), with limited quantities of hazardous waste being generated. figure 1: sabi sand wildtuin in south africa. data capturing and analysis semi-structured face-to-face interviews were administered in april to july 2021. two questions were posed during the interviews: (1) what are the challenges for solid waste management in the ssw pnr? (2) what are the opportunities for solid waste management in the ssw pnr? interviews lasted approximately 45 min to 1 h and were conducted with 30 participants. they included the following: eleven members of the management authority representatives from 13 of the commercial lodges representatives from five non-commercial private properties one of the two waste service providers rendering services to ssw. these participants represent approximately 50% of the total population of management authority, commercial property and non-commercial property members of the ssw. the research was subjected to ethics approval and was classified within the minimal risk category. no vulnerable communities, children or incapacitated adults were included in the interviewee sample. all participants were informed of the interview procedure and their right to withdraw from the interview. interviewees were required to provide informed consent prior to commencement of the interview. results are presented anonymously. voice recorded interviewee responses were transcribed. the framework suggested by mwangi and thuo (2014) for conceptualising and identifying problems, challenges and mechanisms for waste management in developing countries informed the categorisation of responses. the framework provides for (1) technical, (2) financial and economic, (3) social and cultural, (4) environmental health, (5) institutional and political and (6) legal and policy categories. this framework was ideally suited for the categorisation of responses based on the research scope and the type of responses expected during the interview. responses were thematically analysed through coding (c1 to c18 in table 1; o1 to o19 in table 2) and deductive reasoning (braun & clarke 2006:79). this process involved familiarisation with the data, generation of initial codes and finally collating of codes into themes according to the categories proposed by mwangi and thuo (2014). because the number of respondents per stakeholder group are too small to attempt to draw associations or conclusions between statements of the individual stakeholder groups, the responses of all stakeholder groups were consolidated. responses were ranked based on their frequency of mention, and the 10 most frequently mentioned challenges and opportunities across the themes are highlighted in tables 1 and 2, respectively. table 1: the number (and percentage out of a total of 30 participants) of responses per code as it relates to the perceived challenges for sustainable solid waste management in the private nature reserve. table 2: the number (and percentage out of a total of 30 participants) of responses per code as it relates to the perceived opportunities for sustainable solid waste management in the private nature reserve. ethical considerations the faculty of natural and agricultural sciences ethics committee (fnasrec) approved this study and the north-west university senate committee for research ethics (nwu-scre) granted its permission for the study to be initiated, using the reference number nwu-00500-21-a9. results challenges for sustainable solid waste management the perceived challenges for sustainable solid waste management mentioned by interview respondents are outlined in table 1. opportunities for sustainable solid waste management the perceived opportunities for sustainable solid waste management mentioned by interview respondents are outlined in table 2. discussion sustainable solid waste management challenges table 1 outlines the perceived challenges for sustainable solid waste management, as identified by interviewees. according to the four identified themes (a–d in table 1), key findings are discussed below. institutional and financial challenges financial provision for waste management (c1) was noted by 43% of participants and was ranked as the third most frequently mentioned challenge for sustainable solid waste management (table 1). this finding agreed with what other authors (przydatek 2019; rodríguez-rodríguez 2012) have found for protected areas, where insufficient funds and inadequate budget provision were regarded as two of the most significant challenges for sustainable waste management. interviewees were sensitive towards the costs related to waste management, especially considering the financial strain that the lodges have suffered due to coronavirus disease 2019 (covid-19) tourism-related restrictions. one of the participants reported that: ‘we are prepared to pay for waste management, but it should not be prohibitively expensive. we cannot pay much more than we are paying now given our financial situation due to covid-19 restrictions.’ the challenges related to covid-19, conservation and ecotourism were also highlighted by lindsey et al. (2020). insufficient human resources and capacity (c2) were ranked as the fourth most mentioned challenge (mentioned by 40% of interviewees). coetzee and nell (2019) agree that inadequate human resources may hinder the implementation of sound waste management measures in protected areas. approximately one third of the participants indicated that waste management is not a business priority (c3). this code was ranked as the seventh most frequently mentioned challenge. interviewees indicated that waste is often ‘not prioritised’ because it does not form part of the ‘core business of the reserve’. participants also indicated that they perceived certain waste management activities as ‘inconvenient to guests’ and that they ‘did not want to impact negatively on the ecotourism experience of visitors to the reserve’. the eighth most frequently mentioned challenge was inadequate municipal support and a lack of service delivery (c4), which 30% of participants highlighted. the ssw pnr relies on private waste collection and transportation services, whilst the adjacent communities are largely reliant on municipal services. inadequate municipal services have been highlighted as one of the biggest challenges towards effective waste management in south africa in the 2018 state of waste report (department of environmental affairs [dea] 2018). insufficient waste collection services lead to littering and illegal disposal or dumping of waste in adjacent communities. it leads to pollution and undesirable aesthetic impacts, resulting in negative experiences of reserve visitors. it was therefore not surprising that the participants considered creating cleaner communities (o16) to be one of the most significant opportunities related to waste management. insufficient awareness, knowledge and skills (c5), limited justification for a (waste-to-worth) business case (c6), and time and timing challenges (c7) were also noted by interviewees (table 1). challenges related to ‘time’ referred to the amount of time that waste management actions such as separation at source may take up, whilst ‘timing’ referred to poor timing of required waste management actions, caused by restrictions to ecotourism and travel due to the covid-19 pandemic. technical and operational challenges code c8, which related to separated waste being mixed during transportation, was mentioned by 37% of the interviewees. an interviewee stated that ‘we have tried to implement waste separation at our lodge, but all of the waste ends up being mixed again when it is transported together’. lubsanova and maksanova (2020) have also highlighted the ineffective separation of waste as one of the main challenges facing ecotourism resorts and national parks, respectively. other technical and operational challenges perceived by participants included: inadequate infrastructure, machinery and equipment (c9) inadequate services to manage certain waste streams (c10) limited standardisation and coordination of waste management requirements (c11) within the reserve insufficient and infrequent waste collection services (c12) by private waste service providers insufficient data and information on waste quantities and management (c13). these are common issues for waste management within the south african and developing country contexts, also highlighted by authors such as de witt (2015) and dea (2018). social and cultural challenges the most frequently mentioned challenge related to sustainable waste management, raised by 90% of participants, was code c14, challenging behaviour. waste-related behaviour plays a significant role towards integrated waste management in pnrs (roos et al. 2021). negative attitudes, perceived behavioural control elements, unwillingness to engage in waste management activities and a lack of support for waste management activities could all contribute to poor waste management behaviour and unsound waste management practices in protected areas (chen et al. 2020). in the context of the ssw pnr, interviewees raised concerns regarding ‘aversion and reluctance to implement waste management practices’, ‘role players not being willing to change behaviour and practice’, ‘negative attitudes towards waste’ and ‘a lack of buy-in and support from role players’, as well as poor ‘community participation’, ‘landowner support’ and ‘insufficient cooperation’. similarly, ogato (2014) identified poor culture, attitudes and habits of different role players in protected areas towards pollution control and waste management as one of the significant challenges impacting on sustainable ecotourism development in addis ababa, ethiopia. code c15 ranked as the 10th most frequently mentioned challenge. approximately 30% of the interviewees noted that creating unrealistic expectations of what ssw could reasonably contribute to communities with respect to waste management (c15) may be a challenge. a participant from the management authority noted that ‘a risk or challenge could be that the surrounding community might expect more than what ssw could contribute to waste management’. as mentioned earlier, coetzee and nell (2019) have highlighted that in the south african context, available resources, capacity and expertise of protected areas play an important role in determining the extent to which these areas can realistically contribute to a community development agenda. these authors caution against protected areas creating unrealistic expectations, and they emphasise that protected areas can ‘realistically only address some of the identified community needs significantly’. lastly, challenges related to community conflicts and competition (c16) for waste management business opportunities were raised. according to meletis and campbell (2009), communities in the developing country context generally fail to unite efforts to address waste management issues and that community conflicts remain a concern. similarly, one of the participants of this research stated that: ‘we have twelve different community groups around the reserve. what would their expectations be? who will benefit and how? the process of community involvement needs to be fair to provide equal opportunities to all; otherwise, it may lead to competition and social conflict.’ there were some concerns raised by participants about competition amongst community members leading to social conflicts and that social unrests may negatively influence the pnr. challenges due to the characteristics of the reserve some of the challenges brought up by interviewees were related to the characteristics of the reserve. waste management in protected areas is often complicated because many of these areas are located far from waste-related services and infrastructure, and are therefore likely to have limited waste disposal or treatment options available to them (meletis & campbell 2007). approximately 67% of research participants highlighted challenges related to code c17 (which was regarded as the second most frequently mentioned challenge): the size of the reserve: the ssw pnr consists of 49 481 hectares, with commercial and non-commercial properties located in the reserve. the size of the reserve makes it difficult to coordinate waste management measures such as waste collection and makes the establishment of centralised infrastructure for communal use almost impossible. the location of the reserve: the reserve is located in a rural area, with limited waste management facilities and infrastructure located within a 100 km radius from the reserve. compatibility of waste options with protected areas expectations: incineration was identified as a possible means to divert waste from landfill disposal by some stakeholders. the question was, however, posed regarding the compatibility of the technology (with potential air quality impacts) in a protected area. similarly, composting and disposal of waste within the reserve were regarded as challenges because of practical issues such as smells, breeding of vermin and unwanted lure of animals. finally, the unavailability of land within the reserve for waste-related activities and infrastructure was raised as a challenge by 37% of interviewees and ranked as the sixth most frequently mentioned code. participants voiced their reluctance to use valuable conservation land for waste management purposes and added discussions on the complexities of securing privately owned land for communal waste management purposes. the interviewees argued that off-site management of waste would be preferred to on-site waste management. opportunities for sustainable solid waste management table 2 outlines the perceived opportunities for sustainable solid waste management, as identified by interviewees. according to the four identified themes (a – d in table 2), key findings are discussed below. institutional and legal or policy opportunities providing assurance and transparency (o1) was the most frequently mentioned institutional and legal or policy opportunity. code o1 ranked second overall, with 67% of participants regarding it as an opportunity for sustainable solid waste management. the statements of interviewees indicated that they were concerned about what happens to waste ‘beyond the gate’, once it leaves the reserve. providing assurance and transparency are important aspects of demonstrating legal compliance as far as waste management throughout the entire life cycle (from cradle to grave) is concerned. providing assurance of sound waste management beyond the gate may include documented information such as waste manifests and waybills, which outline the ‘chain of custody’ from the point of generation to the final destination of the waste. this information could ultimately be included as part of waste information reporting (code o6) and sustainability reports of lodges. providing strategic direction and standardisation (o7) were noted by 43% of respondents, and ranked as the seventh most mentioned opportunity for sustainable solid waste management. this code captured the participants’ responses related to the opportunities and benefits of having a common strategic vision with goals for waste management within the reserve across all the properties. interviewees made reference to the advantages of having ‘common goals’, ‘a shared vision’ and ‘aligned approaches, policies and plans’ towards ‘driving coordinated efforts in achieving sustainable solid waste management’. przydatek (2019) emphasises the advantages of synergised waste management in protected areas, whilst sandbrook et al. (2019) highlight the importance of having strategic direction with long-term targets towards waste management. coordination could also support efficiency, optimisation of resources and economies of scale, with meaningful potential financial benefits over time. improving the reputation and brand of the reserve and providing marketing opportunities were mentioned by 40% of the interviewees and ranked as the ninth most frequently mentioned opportunity for sustainable solid waste management. an interviewee from one of the commercial lodges highlighted that: ‘demonstrating that we are “green” and legally compliant are important for our reputation and it has marketing value. the public sees sabi sand wildtuin as one entity – if one lodge does not perform well, it may impact negatively on the entire reserve. the flip side may also be true; if a couple of us perform well, it impacts positively on the image and brand of the entire reserve.’ the value of environmental management performance towards improving reputation, image and brand is recognised by reis et al. (2018), who reported that an improvement in image, with associated marketing value, is perceived as one of the most significant benefits of iso 14001 implementation by organisations. considering that the commercial properties in ssw target mainly the high-end luxury ecotourism market, reputation and brand value are important. the 10th most frequently mentioned code (by 33% of interviewees) was o4, where interviewees highlighted the importance of ‘coordinated’, ‘reliable’, ‘efficient’ and ‘effective’ ‘waste services’, which provide for ‘the entire waste management life cycle’ from ‘cradle to grave’ (linking to code o1). respondents did, however, emphasise that waste management services should be sensitive towards cost. funding of waste management services was ranked as the third most significant challenge (code c3) for solid waste management in ssw and was mentioned by interviewees representing both commercial and non-commercial properties. other institutional and legal or policy opportunities included the following: contributing towards legal compliance (o5); improving monitoring, collection and recording of data and improving communication and reporting (of waste data) (o6); and providing business opportunities within ssw (o7). it seems as if the interviewees regarded business opportunities for waste management as a better prospect for local communities than for the reserve itself (as indicated in code o13); however, nobody could point to a specific proven business case (c6) for community benefits outside the reserve. this remains an area of future research, but at this stage it seems that the amount of waste and the particular waste streams from the reserve provide limited commercial benefits and opportunities. this is important to highlight because unrealistic expectations by communities of the commercialisation potential of reserve waste streams could lead to unintended weakening of relations (as discussed in section 4.1.3). environmental health and ecotourism opportunities codes related to environmental health and ecotourism opportunities were not as frequently mentioned as the codes of the other three themes. protecting and preserving the environment (o8) ranked sixth, overall, being mentioned by 43% of interviewees. the responses indicated that participants felt that protection and preservation of the environment formed part of their ethical and moral obligations. one participant said that ‘conservation is our core business’ and ‘we have a moral obligation to protect the environment’. another respondent added that ‘our ethos is “leave the world a better place”. from a moral point of view, we should all want to engage in responsible waste management’. the contribution of sustainable solid waste management towards protecting and preserving the environment is well-recognised by other authors (dunjić et al. 2017; hong & chan 2010; rodríguez-rodríguez 2012; przydatek 2019). meeting visitor expectations (o9) was mentioned by 40% of the participants and ranked as the eighth most frequently mentioned opportunity for sustainable waste management. du plessis et al. (2013) and morrison-saunders et al. (2019) found that responsible waste management ranked amongst the top expectations of visitors to protected areas in south africa. other health and ecotourism-related opportunities of sustainable solid waste management included addressing visual and aesthetic impacts (o10) related to waste and littering, preventing pollution (o11) and contributing to sustainable environmental management (o12). birendra (2021) reported that improved waste management has resulted in reduced pollution and positive aesthetic impacts in communities surrounding the bardia national park in nepal, which improved the environmental conditions and enhanced the ecotourism experience of visitors. social and cultural opportunities the most frequently mentioned opportunity related to sustainable solid waste management (ranked as the most significant opportunity overall) was creating jobs and contributing towards livelihoods (o13), which was mentioned by 77% of interviewees. opportunities raised by participants focused on benefitting and supporting local communities through ‘creating local job opportunities’, ‘creating employment outside tourism activities’, ‘partnering with the private waste sector’ and ‘creating small, medium and micro enterprises (smmes)’, with the aim of ‘contributing to livelihoods’ to ultimately ‘improve their quality of life’. this could, in the long run, contribute to community upliftment (o15) and creating cleaner communities (o16). meletis and campbell (2009) highlight the role that the ecotourism sector could play in community support in assisting communities with waste management problems, especially where municipal waste services are limited. palfrey, oldekop and holmes (2021) highlight the social outcomes of private protected areas, such as local communities benefitting from increased employment, training and community-wide development. furthermore, the contributions of waste towards the green and circular economies is well-recognised (márquez & rutkowski 2020; taleb & farooque 2021). in the developing country context, a significant number of jobs have been created from the informal waste sector, where the harvesting of waste contributes to the diversion of waste away from landfilling (yu, blaauw & schenck 2020). there are, however, several challenges, such as limited justification for a business case (c6), inadequate infrastructure (c9) and services (c10) for waste management and recycling, challenges related to the location of the reserve (c17) and the unavailability of land for the establishment of waste separation and recycling infrastructure within the reserve (c18) (table 1), which need to be addressed in order for job and livelihoods opportunities to be realised. it is therefore important to reflect on the level of support and job creation that the ecotourism sector, and more specifically protected areas, could reasonably and feasibly contribute to communities. coetzee and nell (2019) have cautioned about the risk of creating unrealistic expectations in terms of the level of community support that protected areas could contribute towards sound waste management (also see c15 in table 1). the improvement of awareness, knowledge and skills regarding waste management (o14) was also regarded as a significant opportunity related to waste management, which was ranked third and mentioned by 57% of interviewees. hong and chan (2010), and dunjić et al. (2017) highlight the importance of awareness and knowledge of waste, to ultimately improve waste management behaviour in protected areas. technical opportunities technical opportunities mainly focused on alternatives to landfill disposal of waste. code o17, focusing on implementing the waste management hierarchy (i.e. avoidance, reduction, separation at source, re-use and recycling), was ranked fourth, with 57% of interviewees mentioning phrases related to this code. implementing alternatives to landfilling may lead to socio-economic opportunities, as discussed earlier. although separation of waste at source, re-use, recycling and recovery of waste were highlighted as opportunities, participants also highlighted certain challenges related to implementing the waste management hierarchy (table 1). in particular, composting (o18) and waste-to-energy (o19) options were mentioned as alternatives to landfilling by a number of participants (table 2). conclusions this research aimed to identify the challenges and opportunities for sustainable solid waste management in privately protected areas by focusing on ssw, a pnr located in south africa. several challenges have been identified by interviewees. behaviour was the most frequently mentioned challenge, where interviewees raised concerns about negative attitudes, unwillingness to implement waste management measures and a possible lack of support. other frequently mentioned challenges included foreseen difficulties due to the size and location of the reserve and concerns around funding of waste management measures, especially given the financial implications of covid-19 travel and tourism restrictions. the most frequently identified opportunities included creating jobs and improving livelihoods, providing assurance and transparency of what happens to waste ‘beyond the gate’, and improving awareness, knowledge and skills related to waste management. private nature reserves should, as far as possible, aim to address the challenges and optimise the opportunities related to waste management by focusing on addressing the following aspects: must do – achieve legal compliance: pnrs must align their strategic direction to focus on achieving legal compliance, as a minimum. may do – go beyond legal compliance: pnrs may support communities and invest in services and infrastructure outside the reserve to enhance waste management opportunities beyond legal compliance. nice to have – supporting cleaner communities: it would be beneficial if pnrs support and contribute to cleaner communities with less environmental pollution and reduced aesthetic impacts, which will have direct benefits for communities and secondary improved aesthetic impacts for the pnr, which would ultimately lead to improved conservation efforts. acknowledgements the authors acknowledge the contributions of the sabi sand wildtuin and sabi sand pfunanani trust towards this research. the participation of the various interviewees is acknowledged and highly appreciated. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions c.r., r.c.a. and f.p.r. conceptualised the project and were responsible for the data collection, analysis and interpretation. w.h. played an important role in scheduling interviews and accompanying the research team during data collection. all authors played a role in drafting and critically revising the manuscript for important intellectual content and approved the final version to be published. funding information the sabi sand wildtuin pfunanani trust funded a project which was related to this research; however, this research received no specific grant from any funding agency in the public, commercial or not-for-profit sectors. data availability the authors confirm that the data supporting the findings of this study are available within the article (tables 1 and 2). disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references alberts, r.c., retief, f.p, cilliers, d.p., roos, c. & hauptfleisch, m., 2021, ‘environmental impact assessment (eia) effectiveness in protected areas’, impact assessment and project appraisal 39(4), 290–303. https://doi.org/10.1080/14615517.2021.1904377 birendra, k.c., 2021, ‘ecotourism for wildlife conservation and sustainable livelihood via community-based homestay: a formula to success or a quagmire?’, current issues in tourism 24(9), 1227–1243. https://doi.org/10.1080/13683500.2020.1772206 braun, v. & clarke, v., 2006, ‘using thematic analysis in psychology’, qualitative research in psychology 3(2), 77–101. https://doi.org/10.1191/1478088706qp063oa chang, n.b. & pires, a., 2015, sustainable solid waste management: a systems engineering approach, john wiley & sons, ieee press, hoboken, nj. chen, f., chen, h., liu, s., li, w., li, q. & long, r., 2020, ‘formation and recurrence mechanism of residents’ waste separation behaviour under the intervention of an information interaction’, resources, conservation and recycling 162, 105027. https://doi.org/10.3390/ijerph16101859 coetzee, h. & nell, w., 2019, ‘the feasibility of national parks in south africa endorsing a community development agenda: the case of mokala national park and two neighbouring rural communities’, koedoe 61(1), a1470. https://doi.org/10.4102/koedoe.v61i1.1470 department of environmetnal affairs (dea), 2018, south africa state of waste. a report on the state of the environment, second draft report, department of environmental affairs, pretoria, 105 pp. de witt, l., 2015, ‘key environmental management factors in protected areas: an eco-tourist perspective’, african journal of hospitality, tourism and leisure 4(2), 1–12. dunjić, j., stojanović, v., solarević, m. & kicošev, v., 2017, ‘sustainable waste management in protected areas of vojvodina’, in conference proceeding: contemporary trends in tourism and hospitality: new spaces in cultural tourism, ctth 2017, serbia, novi sad, 1st–2nd september 2017. du plessis, l., van der merwe, p. & saayman, m., 2013, ‘tourists’ perceptions on whether south african national parks are environmentally friendly’, acta academica 45(1), 187–208. hong, c.w. & chan, n.w., 2010, ‘strength-weakness-opportunities-threats: analysis of penang national park for strategic ecotourism management’, world applied sciences journal 10(special issue of tourism & hospitality), 136–145. li, w., 2004, ‘environmental management indicators for ecotourism in china’s nature reserves: a case study in tianmushan nature reserve’, tourism management 25, 559–564. https://doi.org/10.1016/j.tourman.2003.06.001 lindsey, p., allan, j., brehony, p., dickman, a., robson, a., begg, c. et al., 2020, ‘conserving africa’s wildlife and wildlands through the covid-19 crisis and beyond’, nature ecology & evolution 4(10), 1300–1310. https://doi.org/10.1038/s41559-020-1275-6 lubsanova, n.b. & maksanova, l.b.-z., 2020, ‘prospects for the implementation of a separate waste collection system in specially protected areas by the example of the tunkinsky national park’, iop conference series: materials science and engineering 941, 012016. https://doi.org/10.1088/1757-899x/941/1/012016 márquez, a.j.c. & rutkowski, e.w., 2020, ‘waste management drivers towards a circular economy in the global south – the colombian case’, waste management 110, 53–65. https://doi.org/10.1016/j.wasman.2020.05.016 meletis, z.a. & campbell, l.m., 2007, ‘call it consumption! re-conceptualizing ecotourism as consumption and consumptive’, geography compass 1(4), 850–870. https://doi.org/10.1111/j.1749-8198.2007.00048.x meletis, z.a. & campbell, l.m., 2009, ‘benevolent and benign? using environmental justice to investigate waste-related impacts of ecotourism in destination communities’, antipode 41(4), 741–780. https://doi.org/10.1111/j.1467-8330.2009.00696.x mitchell, b.a., stolton, s., bezaury-creel, j., bingham, h.c., cumming, t.l., dudley, n. et al., 2018, guidelines for privately protected areas. best practice protected area guidelines series no. 29, iucn, gland, switzerland, viewed 12 december 2021, from https://mpg.eurosite.org/wp-content/uploads/iucn-guidance-on-management-of-privately-owned-pas.pdf. morrison-saunders, a., hughes, m., pope, j., douglas, a. & wessels, j., 2019, ‘understanding visitor expectations for responsible tourism in an iconic national park: differences between local and international visitors’, journal of ecotourism 18(3), 284–294. https://doi.org/10.1080/14724049.2019.1567740 mwangi, m. & thuo, a.d.m, 2014, ‘towards conceptual and theoretical foundation for identifying problems, challenges and mechanisms for municipal waste management in developing countries’, international journal of innovation and scientific research 2(2), 230–251. ogato, g.s., 2014, ‘planning for sustainable tourism: challenges and opportunities for ecotourism development in addis ababa, ethiopia’, american journal of human ecology 3(2), 20–26. https://doi.org/10.11634/216796221403570 palfrey, r., oldekop, j. & holmes, g., 2021, ‘conservation and social outcomes of private protected areas’, conservation biology 35(4), 1098–1110. przydatek, g., 2019, ‘waste management in selected national parks – a review’, journal of ecological engineering 20(4), 14–22. https://doi.org/10.12911/22998993/102609 reis, a., neves, f., hikichi, s., salgado, e. & beijo, l., 2018, ‘is iso 14001 certification really good to the company? a critical analysis’, production 28, e20180073. https://doi.org/10.1590/0103-6513.20180073 rodríguez-rodríguez, d., 2012, ‘littering in protected areas: a conservation and management challenge – a case study from the autonomous region of madrid, spain’, journal of sustainable tourism 20(7), 1011–1024. https://doi.org/10.1080/09669582.2011.651221 roos, c., retief, f., alberts, a., cilliers, d., hodgson, w. & olivier, i., 2021, ‘understanding waste management behaviour in private nature reserves through the theory of planned behaviour’, parks 27(2), 47–56. https://doi.org/10.2305/iucn.ch.2021.parks-27-2cr.en sandbrook, c., fisher, j.a., holmes, g., luque-lora, r. & keane, a., 2019, ‘the global conservation movement is diverse but not divided’, nature sustainability 2, 316–323. https://doi.org/10.1038/s41893-019-0267-5 steg, l. & vlek, c., 2009, ‘encouraging pro-environmental behaviour: an integrative review and research agenda’, journal of environmental psychology 29, 309–317. https://doi.org/10.1016/j.jenvp.2008.10.004 taleb, m.a. & farooque, o.a., 2021, ‘towards a circular economy for sustainable development: an application of full cost accounting to municipal waste recyclables’, journal of cleaner production 280(2), 124047. https://doi.org/10.1016/j.jclepro.2020.124047 van der merwe, p., saayman, m. & bothma, l.a., 2017, ‘what should be included in an ecotourism rating system for protected areas?’, african journal of hospitality, tourism and leisure 6(3), 1–22. yadav, v. & karmakar, s., 2020, ‘sustainable collection and transportation of municipal solid waste in urban centers’, sustainable cities and society 53, 101937. https://doi.org/10.1016/j.scs.2019.101937 yu, d., blaauw, d. & schenck, r., 2020, ‘waste pickers in informal self-employment: over-worked and on the breadline’, development southern africa 37(6), 971–996. https://doi.org/10.1080/0376835x.2020.1770578 kruger.qxd the effect of neighbouring klipspringer on the scent–marking behaviour of a group of klipspringer in the kruger national park m. kruger, j. du p. bothma and j.m. kruger m. kruger, j. du p. bothma and j.m. kruger. 2002. the effect of neighbouring klipspringer on the scent–marking behaviour of a group of klipspringer in the kruger national park. koedoe 45(1): 87–92. pretoria. issn 0075-6458. both the male and the female klipspringer scent-mark their ranges. a pair of pre-orbital glands below the eyes produces the scent. the secretion is a sticky, substance that is deposited on a suitable twig. klipspringer scent marks were surveyed in a specific klipspringer range in the kruger national park with the use of a strip transect method. the results showed that klipspringer in the kruger national park scent-mark more frequently on the boundaries of their ranges and also more on those sides where there is another resident klipspringer group. key-words: klipspringer, scent-marking, behaviour, range, neighbours. m. kruger, j. du p. bothma, centre for wildlife management, university of pretoria, pretoria 0002 south africa; j.m. kruger, conservation services: south african national parks, kruger national park, private bag x402, skukuza, 1350 south africa. (m. kruger, current address: conservation services, south african national parks, private bag x402, skukuza, 1350 south africa.) issn 0075-6458 87 koedoe 45/1 (2002) introduction scent-marking behaviour has long been regarded as an important component of animal communication (thiessen & rice 1976). a number of hypotheses to explain why animals scent-mark have been put forward. these range from deterrence of intruders (hediger 1949) to the establishment of a familiar smell within the range (johnson 1973). one such hypothesis proposes that the scent advertises the identity of a range inhabitant to an intruder, enabling the latter to assess the holder’s status, thus avoiding costly fights (gosling 1982). scent-marking is also considered to be associated with dominance in many animal species (ralls 1971; stoddard 1976; brown & mcdonald 1985; kappeler 1980; somers et al. 1990). like other small antelope (cohen & gerneke 1976) both the male and female klipspringer scent-mark their ranges. this chemical communication functions as an unmistakable cue to ownership of a specific range. the scent is produced as a secretion by a pair of preorbital glands just below the eyes. the animal positions its head at a certain angle and rubs it against a suitable twig that normally points in an upward direction so that the twig is inserted directly into the gland. a milky substance is secreted that has a sweet, aniseed-like scent which turns brown once dried. the hypothesis for this preliminary study of scent-marking in the klipspringer in one study area in the kruger national park was that klipspringer would mark more frequently towards the boundaries of their territories, and also more frequently on those sides where there were resident neighbouring klipspringer in their proximity. study area the kruger national park is situated in the lowveld of the mpumalanga and northern provinces along the northeastern boundary of the republic of south africa. this park lies between 22º19'–25º32's latitude and 30º54' kruger.qxd 2005/12/09 11:33 page 87 –32º02'e longitude. this study was conducted in the nwatindlopfu spruit area, approximately 25 km north-east of skukuza in the central district of the kruger national park. this area was selected because it had more than 10 known resident groups of klipspringer in close proximity. this area can briefly be described as follows. according to the landscape classification of the kruger national park by gertenbach (1983), this area includes two landscapes. they are the thickets of the sabie and crocodile rivers and the mixed combretum species / terminalia sericea woodland. the area is underlain by granite and gneiss, with numerous dolerite intrusions that are more than 10 m wide (schutte 1986). the altitude of the landscape varies from 200 m to 500 m above sea level with gently sloping hills and valleys. vegetation the vegetation of this area can be described as mixed combretum savanna woodland with dense thorn bush thickets on the banks of the spruits and rivers (pienaar 1963). the dominant woody species are acacia nigrescens, combretum apiculatum, grewia bicolor, grewia flavescens, sclerocarya birrea, euclea divinorum, spirostachys africana, dichrostachys cinerea, ziziphus mucronata, lonchocarpus capassa, strychnos madagascariensis, cissus cornefolia, ormocarpum trichocarpum and dalbergia melanoxylon. the herbaceous grass layer in the area is dominated by pogonarthria squarrosa, perotis patens, melinis repens, digitaria eriantha, panicum maximum, aristida congesta, schmidtia pappophoroides, urochloa mosambicensis, heteropogon contortus, enneapogon cenchroides, bothriochloa radicans and themeda triandra. the dominant forbs are waltheria indica, evolvulus alsinoides, heliotropium steudneri, dyschoriste rogersii, abutilon astro-africanum, justicia flava, sansevieria hyacinthoides, rhynchosia totta, cassia mimosoides, commelina bengalensis and hibiscus micranthus. koedoe 45/1 (2002) 88 issn 0075-6458 fig. 1. schematic presentation of a klipspringer range that was surveyed for scent marks, and the proximity of the nearest neighbours. mammals the largest population of impala aepyceros melampus in the kruger national park most probably occurs in this area. other mammals include the kudu tragelaphus strepsiceros, common duiker sylvicapra grimmia, steenbok raphicerus campestris, bushbuck tragelaphus scriptus, giraffe girraffa camelopardalis, sable antelope hippotragus niger, warthog phacochoerus aethiopicus, elephant loxodonta africana, white rhinoceros ceratotherium simum and black rhinoceros diceros bicornis. the predators that occur in the area are the lion panthera leo, leopard panthera pardus, cheetah acinonyx jubatus, wild dog lycaon pictus and spotted hyaena crocuta crocuta. kruger.qxd 2005/12/09 11:33 page 88 materials and methods klipspringer scent marks were surveyed with the use of a strip transect method. the transects were placed out from the centre of the range of a specific klipspringer group in the nwatindlopfu spruit area in the four directions, viz., north, south, east and west. each transect was 100 m wide and the survey was done from the centre to somewhat beyond the known boundary of the range of that group. each plant in the transect was inspected carefully for scent marks. when scent marks were found, the type of plant, the position of it in the transect and the number of marks present were recorded. the distance between this scent mark and the known territory boundary of the nearest klipspringer group was also recorded. figure 1 shows the schematic range of klipspringer surveyed, and its position relative to the closest neighbours. the distances between the klipspringer range and the neighbours were obtained by calculating the distance between the known territory boundaries of the studied klipspringer group and the observed klipspringer surrounding them. data analysis a two-fold approach was used to analyse the scentmarking data. the first approach was to investigate the spatial distribution of these marks so as to determine whether they were distributed at random or were clumped within the transects. the null hypothesis that was tested was that a klipspringer scentmarks randomly throughout its range. should this be true, then the next approach was to use log-linear regression techniques in an effort to explain the pattern found. since the variable that was tested was the number of scent marks, normal linear regression methods of analysis were, however, not suitable because this linear method may lead to negative counts, and the variance will not be constant (crawley 1996). loglinear regression techniques or the poisson distribution were therefore used to analyse the data. the poisson distribution is used in cases where it is known how many times an event happened but where it is not known how many times an event did not happen. here it meant how many times a klipspringer scent-marked within its range (crawley 1996). variables recorded the following variables were recorded: the number of scent marks. these were counted in four directions from the centre of the range as described above; issn 0075-6458 89 koedoe 45/1 (2002) the distance between a scent mark and the nearest neighbouring klipspringer; the distance from the centre of the range to the nearest neighbouring klipspringer; the number of neighbouring klipspringer bordering the range in all directions. factors affecting scent-marking: model fitting the full model that was fitted included the variables indicated above. the only interaction term that was included in the model was the interaction between the variable distance between the scent marks and the neighbouring klipspringer and the variable number of neighbouring klipspringer. when the full model was fitted it was determined that the data were over dispersed which means that the probability that we were attempting to model was not constant within each cell, but behaved like a random variable. to correct for this we assumed that the variance was proportional to the binomial variance and not equal to it. we calculated the ratio of scaled deviance to degrees of freedom and used the scale directive to alter the scaled deviance (crawley 1996). in order to evaluate the extent to which an adopted model fits a set of data the distribution of deviance is needed. the deviance is asymptotically distributed as χ² with (n–p) degrees of freedom, where n is the number of observations and p is the number of unknown parameters included in the model. in order to obtain the best model the constant was first fitted and then each variable was included in the model one at a time. a variable was considered significant when the change in deviance associated with including that variable exceeded the chisquare critical value at p < 0.05 for the number of degrees of freedom that the variable conferred (crawley 1996). results spatial distribution figure 2 shows the distribution of the scent marks as a three-dimensional map in which the third vertical dimension reflects the value of the quantitative data. the latter dimension represents the density of the scent marks. it is clear that the scent marks appear to be clumped in specific areas, mainly along the border of the range, with hardly kruger.qxd 2005/12/09 11:33 page 89 any scent-marking occurring in the centre of the range. when this representative pattern was subjected to a nearest neighbour analysis (krebbs 1989), and was compared with an expected random distribution, the index of aggregation (r) was 0.36128. this ratio indicates a deviation of the observed pattern from an expected random pattern. when the spatial pattern is random, then r = 1, and when clumping occurs then r approaches zero. therefore it can be concluded that the scent marks where indeed clumped in specific areas. to test the significance of the deviation from a random pattern, the z statistic was calculated, where z = ra–re/se, ra = the mean distance to the nearest neighbour, re = the expected distance to the nearest neighbour, and se = the standard error of the expected distance to nearest neighbour. this test yielded a z value of 20.79144, which is greater than 1.96 (p = 0.05). meaning the null hypothesis that the scent marks are randomly spaced is rejected (krebs 1989), therefore, they show a clumped pattern. koedoe 45/1 (2002) 90 issn 0075-6458 table 1 poisson regression analyses of the effect of neighbouring klipspringer and the distance of scent marks from the centre of the range on the number of scent marks in klipspringer in the kruger national park. star notation in the table represents the interaction between 2 variables variable degrees change probability of freedom in deviance value number of neighbouring klipspringer 1 -4.511 0.0337 distance from centre of range to nearest neighbouring klipspringer 1 -8.884 0.0029 distance between a scent mark and nearest the neighbouring klipspringer 1 -7.854 0.0050 distance between a scent -mark and the nearest neighbour. * number of neighbours 1 0.020 0.8875 figure 2. three-dimensional representation of klipspringer scent marks in the kruger national park to show the distribution of scent marks. the black dot shows the centre of the range. kruger.qxd 2005/12/09 11:33 page 90 log-linear regression the variable that had the most influence on the number of scent marks was the distance to the nearest neighbouring klipspringer, because it resulted in the greatest change in deviance when it was excluded from the model—and the probability value associated with it was the smallest. the variable of the number of neighbouring klipspringer had the least effect on the number of scent marks because the drop in deviance was the smallest and the probability associated with it was the largest of the significant variables. the interaction between the distance between a scent mark and the nearest neighbouring klipspringer, and between the number of neighbouring klipspringer was, however, not significant (p > 0.05) (table 1). the change in deviance and the corresponding degrees of freedom for each of the variables tested are shown in table 1. the effect of distance to the nearest neighbouring klipspringer and the distance to a given scent mark are illustrated in figs. 3 & 4. issn 0075-6458 91 koedoe 45/1 (2002) fig. 3. the effect of the distances (m) from the scent mark to the nearest neighbouring klipspringer on the number of scent marks produced by the klipspringer group studied in the kruger national park as obtained from log–linear regression techniques. figure 4 the effect of the distance from the centre of the studied klipspringer range on the number of scent marks produced by these klipspringer in the kruger national park. discussion it is clear that as the distance between the scent marks and the nearest neighbouring klipspringer increases, the number of scent marks decreases (fig. 3). therefore the closer a klipspringer is to a neighbouring klipspringer the more scent-marks will occur. also, the further away that the scent marks are from the centre of the range, the more scent marks there are (fig. 4). therefore, the number of scent-marking increases near the boundaries of a range. it is also clear that the number of neighbours that a klipspringer have will affect the number of scent-marking (table 1). however, the data are not conclusive because there were limited klipspringer concentrations. nevertheless, the data does indicate that all the variables except the interaction between the distance between a scent mark and the nearest neighbouring klipspringer and the number of neighbouring klipspringer have a significant effect on the number of scent-marks in klipspringer in the kruger national park. kruger.qxd 2005/12/09 11:33 page 91 conclusions roberts (1998) observed a similar scentmarking pattern in klipspringer in the limpopo river valley of zimbabwe as was found here with the use of a nearest-neighbour mapping technique. in the study by roberts (1998) the scent marks were placed in a rough ring just within the range boundaries of a resident group. the result of the current preliminary study on scent-marking in the klipspringer in the kruger national park, supports the hypothesis that klipspringer will scent-mark more frequently on the boundaries of their ranges, and also more on those sides of the range where there is another resident klipspringer group. we therefore conclude that scent-marking in klipspringer is used as a spacing behaviour. acknowledgements we would like to thank the south african national parks, kodak sa and the csir for their financial assistance. references brown, r.e & d.w. macdonald. 1985. social odours in mammals. oxford: clarendon press. cohen, m. & w.h. gerneke. 1976. preliminary report on the inter-mandibular cutaneous area and the infra-orbital gland of the steenbok. journal of south african veterinary association 47(1):35–37. crawley, m.j. 1996. glim for ecologists. cambridge: university press. gertenbach, w.p.d. 1983. landscapes of the kruger national park. koedoe 26 :9–121. gosling, l.m. 1982. a reassessment of the function of scentmarking in territories. zeitschrift für tierpsychologie 60: 89–118. hediger, h. 1949. säugetier-territorien und ihre makierung. bijdragen tot de dierkunde 28: 172–184 johnson, r.p. 1973. scent marking in mammals. animal behaviour 21: 521–535 kappeler, p.m. 1990. social status and scent marking behaviour in lemur catta. animal behaviour 40: 774–788. krebbs, j.c. 1989. ecological methodology. new york: harper collins. pienaar, u. de v. 1963.the large mammals of the kruger national park—their distribution and present-day status. koedoe 6:1–37. ralls, k. 1971. mammalian scent marking. science 171: 443–449 schutte, i.c.1986. the general geology of the kruger national park. koedoe 29 : 13–37. somers, m., o.a.e. rasa & p.j. apps. 1990. marking behaviour and dominance in suni antelope (neotragus moschatus). zeitschrift für säugetierkunde 55: 340–352. stoddart, d.m. 1976. mammalian odours and pheromones. london: edward arnold. thiessen, d. & m. rice. 1976. mammalian scent gland marking and social behaviour. psychological bulletin 83: 505–539. koedoe 45/1 (2002) 92 issn 0075-6458 kruger.qxd 2005/12/09 11:33 page 92 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <feff004200720075006700200064006900730073006500200069006e0064007300740069006c006c0069006e006700650072002000740069006c0020006100740020006f0070007200650074007400650020005000440046002d0064006f006b0075006d0065006e0074006500720020006d006500640020006800f8006a006500720065002000620069006c006c00650064006f0070006c00f80073006e0069006e0067002000740069006c0020007000720065002d00700072006500730073002d007500640073006b007200690076006e0069006e0067002000690020006800f8006a0020006b00760061006c0069007400650074002e0020005000440046002d0064006f006b0075006d0065006e007400650072006e00650020006b0061006e002000e50062006e006500730020006d006500640020004100630072006f0062006100740020006f0067002000520065006100640065007200200035002e00300020006f00670020006e0079006500720065002e00200044006900730073006500200069006e0064007300740069006c006c0069006e0067006500720020006b007200e600760065007200200069006e0074006500670072006500720069006e006700200061006600200073006b007200690066007400740079007000650072002e> /nld <feff004700650062007200750069006b002000640065007a006500200069006e007300740065006c006c0069006e00670065006e0020006f006d0020005000440046002d0064006f00630075006d0065006e00740065006e0020007400650020006d0061006b0065006e0020006d00650074002000650065006e00200068006f00670065002000610066006200650065006c00640069006e00670073007200650073006f006c007500740069006500200076006f006f0072002000610066006400720075006b006b0065006e0020006d0065007400200068006f006700650020006b00770061006c0069007400650069007400200069006e002000650065006e002000700072006500700072006500730073002d006f006d0067006500760069006e0067002e0020004400650020005000440046002d0064006f00630075006d0065006e00740065006e0020006b0075006e006e0065006e00200077006f007200640065006e002000670065006f00700065006e00640020006d006500740020004100630072006f00620061007400200065006e002000520065006100640065007200200035002e003000200065006e00200068006f006700650072002e002000420069006a002000640065007a006500200069006e007300740065006c006c0069006e00670020006d006f006500740065006e00200066006f006e007400730020007a0069006a006e00200069006e006700650073006c006f00740065006e002e> /esp <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> /suo <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> /ita <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> /nor <feff004200720075006b00200064006900730073006500200069006e006e007300740069006c006c0069006e00670065006e0065002000740069006c002000e50020006f00700070007200650074007400650020005000440046002d0064006f006b0075006d0065006e0074006500720020006d006500640020006800f80079006500720065002000620069006c00640065006f00700070006c00f80073006e0069006e006700200066006f00720020006800f800790020007500740073006b00720069006600740073006b00760061006c00690074006500740020006600f800720020007400720079006b006b002e0020005000440046002d0064006f006b0075006d0065006e0074006500720020006b0061006e002000e50070006e006500730020006d006500640020004100630072006f0062006100740020006f0067002000520065006100640065007200200035002e00300020006f0067002000730065006e006500720065002e00200044006900730073006500200069006e006e007300740069006c006c0069006e00670065006e00650020006b0072006500760065007200200073006b00720069006600740069006e006e00620079006700670069006e0067002e> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 page 4 abstract introduction materials and methods results discussion conclusion acknowledgements references appendices appendix 2 appendix 3 appendix 4 about the author(s) pascal vittoz institute of earth surface dynamics, faculty of geosciences and environment, university of lausanne, lausanne, switzerland federico pellacani institute of earth surface dynamics, faculty of geosciences and environment, university of lausanne, lausanne, switzerland rémy romanens institute of earth surface dynamics, faculty of geosciences and environment, university of lausanne, lausanne, switzerland ali mainga van thuyne ridge research center, chobe enclave, botswana eric p. verrecchia institute of earth surface dynamics, faculty of geosciences and environment, university of lausanne, lausanne, switzerland richard w.s. fynn okavango research institute, maun, botswana citation vittoz, p., pellacani, f., romanens, r., mainga, a., verrecchia, e.p. & fynn, r.w.s., 2020, ‘plant community diversity in the chobe enclave, botswana: insights for functional habitat heterogeneity for herbivores’, koedoe 62(1), a1604. https://doi.org/10.4102/koedoe.v62i1.1604 note: additional supporting information may be found in the online version of this article as online appendix 1 and online appendix 2. original research plant community diversity in the chobe enclave, botswana: insights for functional habitat heterogeneity for herbivores pascal vittoz, federico pellacani, rémy romanens, ali mainga, eric p. verrecchia, richard w.s. fynn received: 22 oct. 2019; accepted: 23 july 2020; published: 21 oct. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract precise vegetation descriptions and maps are essential tools for the management of natural areas, as well as for understanding animal habitat use. the chobe enclave (ce), adjacent to the chobe national park and the chobe forest reserve, forms a critical dry season range for many large herbivores. as a tool for future management and studies about wildlife habitat use and migration, this study proposed to describe the plant communities in the ce and to study their relationships with microtopography and soils. plant species were inventoried in 82 sampling plots (40 x 20 m), covering the vegetation diversity recognised by an unsupervised classification (landsat images, 30-m resolution). a hierarchical clustering classified the inventories in eight plant communities, mapped with a supervised classification. this study was conducted in parallel with a soil study. soil variations and degree of flooding largely determine community composition. floodplains along the linyanti river and dambos (concentrating local run off from rainfall) provide reliable green forage for wildlife during the dry season. adjacent to floodplains, riverine forests also maintain green browse and grazing well into the dry season. in drylands, vegetation is largely determined by soil texture. forests dominated by baikiaea plurijuga occupy the acidic, red sands in the east, while sandveld vegetation grows on deep sands in paleo-river channels. these habitats support dominant grasses, which provide important forage for grazers during the wet season. finally, woodlands dominated by colophospermum mopane, characterised by sodium-rich and alkaline soils, enable herbivores to meet their mineral requirements during reproduction. conservation implications: our soil and vegetation studies provide important insights into factors determining plant communities. their diversity and close vicinity play a critical role in enabling herbivores to adapt to seasonal variations in forage quantity and quality. results will enable researchers to gain insights into animal habitat seasonal use in the chobe enclave. keywords: plant community ecology; vegetation map; soil; water availability; microtopography. introduction biodiversity is threatened worldwide by human activities, directly by its land-use change and introduced species, as well as indirectly by pollution and climate change (sala et al. 2000). although many studies attempted to predict potential future changes (mcclean et al. 2006), only precise and georeferenced data (e.g. species lists) will be able to monitor global change impacts in the future. in plant sciences, exhaustive species inventories are directly beneficial to study their distribution in the landscape and to understand how biodiversity is related to environmental factors. moreover, such exhaustive plant surveys provide information on the dominant species, their co-occurrence with other species and are the basis for the classification of plant communities (ewald 2003). in relation to respective biotic and abiotic factors, they also help to obtain a better knowledge of species autoecology (e.g. ischer et al. 2014). at a landscape scale, the classification of plant communities remains an important tool for improved management and conservation of species and ecosystems and provides the basis for vegetation maps (de cáceres et al. 2015). vegetation maps and descriptions offer an overview of the distribution of ecosystems and associated environmental conditions. they assist in understanding and studying wildlife distribution and migration and improving the management of protected areas (bennitt, bonyongo & harris 2014; fynn et al. 2014; sianga & fynn, 2017). in the process of vegetation map development, field data are still an essential step to get a high-quality classification. in addition, remote sensing data are a powerful aid to prepare field work and to delineate plant communities for large areas (walker et al. 2016). the chobe enclave (ce) is located between two conservation areas in northern botswana (jones 2002), as well as conservancies directly across the border in namibia. situated along the linyanti river, it provides suitable habitat for large populations of mega herbivores all around the year, especially coming from surrounding conservation areas during the dry season. apart from wild herbivores, it is grazed by cattle of a small number of villages (chase 2013). this situation can lead to conflicts between pastoralists and wild fauna. it is therefore important that the seasonal and daily use of the territory by fauna is investigated to ensure optimal management to the benefit of both pastoralists and wildlife. a typology and vegetation map of the region can provide useful information in this context. detailed data about the vegetation of the ce are scarce. a vegetation map covering the much larger region of the savuti–mababe–linyanti ecosystem exists, but it has an insufficient spatial resolution for efficient management at the local scale. our study is novel in the sense that it was accompanied by a detailed soil study (romanens et al. 2019), which provides key insights into mechanisms underlying plant community composition, as well as for understanding herbivore habitat selection. hence, our study provides the first detailed soil-vegetation classification and map. this simultaneous soil study showed an unexpected soil diversity in a relatively flat region on homogeneous sand-dominated bedrock (romanens et al. 2019). this diversity is representative for soil conditions in and around the ce. we can expect that plant communities have a corresponding diversity. therefore, the aims of this study in the ce are (1) to describe the plant communities, (2) to understand their distribution in the landscape in relation to topography, substrate and hydrology and (3) to develop a vegetation map that can be used for animal habitat use studies and conservation planning. study area the ce (17.94° 18.36°s, 23.93° 24.59°e; 930 m above sea level), with an area of 1690 km2, is separated from namibia by the linyanti river (figure 1). it is part of the 440 000 km2 kavango-zambezi trans-frontier conservation area (kaza-tfca; pricope et al. 2015). the ce is managed by the ce community trust (cect; pricope et al. 2015), but it has a lower protection status than the adjacent chobe national park. figure 1: location of the chobe enclave in botswana (delimited by the red rectangle in inset), between namibia in the north and west, chobe national park in the south and chobe forest reserve in the east. the limits of the chobe enclave are delineated by the red line, including the border with namibia, which follows the linyanti river (in blue). the ce is characterised by a wet season between november and march with a mean annual rainfall of 650 mm (maximum daily temperatures 30 °c – 40 °c) and a dry season between april and october (jones 2002). evapotranspiration is higher than rainfall for all months of the year. the ce is part of the kalahari basin dominated by sandy soils. the ce is separated from namibia by the linyanti geological fault (along which the linyanti river is forming swamps), and from the chobe forest reserve by the chobe geological fault with 35 m – 40 m difference in elevation (kinabo et al. 2008; mccarthy 2013). during the late pleistocene and early holocene, the ce was a large delta (burrough & thomas 2008). these conditions locally induced the deposition of lacustrine carbonate and diatomite layers on top of the original sand layers (diaz et al. 2019), and sand dunes were stabilised on the shores of palaeolakes (burrough & thomas 2008). islands probably emerged by accumulation of salts because of plant evapotranspiration – a process comparable to the present-day okavango delta system (mccarthy et al. 2012). the southwestern part of the system became disconnected from the linyanti river and subsequently dried up following the uplift of the linyanti geological fault. then erosion removed the sand around the carbonate layers. these layers are nowadays visible as platforms at about 2 m above the sand plains (diaz et al. 2019). channels became filled with sand. fossil islands and channels, carbonate platforms and sand dunes shape the present-day complex microtopography. apart from this microtopography and faults, the ce is almost completely flat with a mean slope of 0.4% from southwest to northeast and a mean slope of 0.55% from northwest to southeast. a more complete description of the geology and geomorphology is given in the parallel study conducted on soils (romanens et al. 2019). the local human population is limited to five villages distributed in the northeast of the ce, mostly close to the boundary with a total of approximately 3400 inhabitants (figure 1; chase 2013). agriculture mostly consists of extensive cattle breeding, whilst small crop fields are located close to and around the village areas. an estimated 4800 cattle are kept within a 10-km radius around the villages, although only a minority (600–700) would be grazing in the ce (chase 2013). surrounded by species-rich protected areas (chobe national park and chobe forest reserve), the ce shelters many large herbivores as well. chase (2013) estimated the following wildlife populations in the ce during the dry season of 2011: 9003 elephants, 2443 buffaloes, 94 giraffes, 58 hippopotami and 1492 zebras. important populations of roan and sable antelopes, eland, impala and kudu are also living in the area. materials and methods unsupervised classification and vegetation sampling three landsat 8 images of the ce were selected representing different seasons in 2015 (appendix 1): february (wet season), may (beginning of dry season) and september (middle to end of dry season). bands 2–7 were retained for vegetation mapping: blue, green, red, near infrared and two bands of short-wave infrared. the three scenes were stacked into a single raster (30 × 30 m ground resolution), which contained the selected six bands for the three periods. unsupervised classification was run on this raster with the k-means non-hierarchical clustering algorithm (hartigan 1975), according to pixel spectral signature. on the basis of patterns visually identifiable on landsat images and google earth exploration and from previous authors’ observations in the region, 10 classes were retained as a first division in plant communities. roads and tracks were added from google earth images and authors’ experience of the region to select accessible sites. a 250-m buffer around tracks was applied to restrict the time necessary to reach the sampling plots on foot. within this buffer, 10 sampling plots were randomly selected for each class. as a result of weather-related logistic constraints, we had to modify the original sampling strategy. ultimately, most sampling plots were selected directly in the field, in an area closer to base camp (van thuyne ridge research center). in spite of this, the unsupervised classification was used to ensure a balanced sampling between initial classes. only homogeneous areas chosen according to the unsupervised classification were retained for plant surveys (see appendices 2 and 3 for the final distribution of the sampling plots). field descriptions and vegetation classification for each plot, 40 × 20 m of homogeneous vegetation was surveyed between mid-january and mid-march 2017. this area was large enough to contain almost the complete diversity of plant species representative of the plant community, even for tree species with low cover (woodlands), and was similar to former surveys in the same region (sianga & fynn 2017). plot coordinates were recorded by using a global positioning system (gps) device. mean tree height, total tree cover, shrub cover and herb cover were estimated visually in percentage for each plot. all woody, grass and forb species were identified and their respective cover visually estimated in percentage. nomenclature followed the plant list (http://www.theplantlist.org, version 1.1) and kyalangalilwa et al. (2013) for the former acacia. for woody species, specimens exceeding 2 m in height were considered as trees, whereas smaller specimens were considered as shrubs. this separation gave a precise description of the vertical structure, although only the highest of both values (tree or shrub cover) was retained for subsequent analyses. as a result of time constrains, soil profiles were dug and described for the first 36 of 82 plots only. for 16 soil profiles, representing the whole range of observed variations (microtopography, plant communities), laboratory analyses completed the description (phwater, cation exchange capacity [cec], total organic carbon, soil texture). to ensure that these profiles included all soil variations, a total of 27 supplementary soil profiles were described in the field from cores extracted with a soil auger. the most important physicochemical parameters for plant growth are grouped per plant community and reported in the ‘results’ section. statistical analyses were conducted by using r software (r development core team; www.r-project.org). to reduce the weight of dominant species, cover values were square-root transformed (wildi 2010). a similarity matrix was calculated (bray-curtis distance) to build a dendrogram with ward’s minimum variance method. the number of groups was defined by an approach based on the indval method (dufrêne & legendre 1997). this method calculates indicator values for each species based on its fidelity and its relative abundance in a predefined number of groups. a randomisation procedure evaluates the significance of the species indicator values. this was applied iteratively to different numbers of groups, considering that the most meaningful group number corresponds to the highest number of significant indicator species (p < 0.05) and the lower overall mean p-value. eight groups were considered as the most relevant division, coherent with field observations. diagnostic species of each group were extracted by calculating the indicator value based on species presence only (dufrêne & legendre 1997) and evaluated by using a randomisation test. diagnostic species occurring in < 40% of plots were not retained because they are of little help in recognising the plant community in the field (except those with a mean cover > 2%). finally, for each plot, species richness, simpson diversity and evenness were calculated. vegetation mapping once the eight plant communities were defined, vegetation was mapped with a supervised classification (or semi-automatic classification) based on the previously used 18 bands. to provide examples of the variety of spectral signatures that are associated with each plant community, training polygons were manually selected for homogenous areas around each plot (schmidt 2017), based on field knowledge and google earth. the range of spectral values occupied by the respective training sites was used to define a multidimensional parallelepiped (with as many dimensions as bands used) for each plant community, according to the land cover signature classification (perumal & bhaskaran 2010) provided by the semi-automatic classification plugin (gongedo 2017). each pixel was classified depending on the parallelepiped in which it was included. pixels whose signature overlapped two or more parallelepipeds were classified by using the minimum distance algorithm. this algorithm calculates the euclidean distance between the spectral signatures of the pixel and training sites and assigns pixels to the nearest plant community. results a total of 383 species were encountered in the 82 sample plots. the eight plant communities comprise three grassland, three woodland and two forest communities (table 1; figure 2; appendix 4). site 33, originally classified in the wet floodplains (see definition in section wet floodplain), was excluded from subsequent analyses because of its unique composition and structure: a termite mound in a floodplain, but with a heterogeneous composition. figure 2: overview of plant communities in the chobe enclave. (a) aristida junciformis – aristida meridionalis grassland, dominated by a. junciformis and a. meridionalis (plot 71); (b) ipomoea chloroneura – oxygonum alatum woodland, with terminalia sericea, vachellia erioloba and panicum maximum (plot 48); (c) baikiaea plurijuga – baphia massaiensis forest (plot 8); (d) colophospermum mopane – jasminum stenolobum woodland, dominated by c. mopane and aristida adscensionis (plot 36); (e) eragrostis superba – combretum hereroense woodland, dominated by c. hereroense and digitaria eriantha (plot 3); (f) croton megalobotrys – setaria verticillata forest, dominated by vachellia tortilis (plot 16); (g) geigeria schinzii – setaria sphacelata grassland, after a heavy rain; (h) wet floodplain, dominated by andropogon eucomus (plot 81). table 1: mean tree cover, tree height, shrub cover and herb cover for the eight plant communities. aristida junciformis – aristida meridionalis grassland this unit comprises low cover grasslands (mean cover 42.2% ± 15.6%; table 1) established on sandy and thick alluvial deposits. these grasslands are flooded at a low frequency (possibly not each year). however, the observed water whilst digging soil pits and the general absence of trees suggest that they are still under the seasonal influence of a high water table. the very high sand proportion (~98%) of the substrate is related to the lowest content of organic matter (< 0.1%), both explaining the low cation exchange capacity (cec; < 0.4 cmol+ kg−1; table 2). the soils were classified as arenosols by romanens et al. (2019). amongst the diagnostic species, the dominant aristida meridionalis and a. junciformis, schizachyrium jeffreysii and hyperthelia dissoluta are all poaceae with low palatability and nutritional value (roodt 2015). other diagnostic species are hermannia eenii, bulbostylis hispidula, dicerocaryum eriocarpum and pycreus pelophilus (online appendix 2). with a mean of 21.9 ± 4.8 species, plots belonging to this unit were amongst the species poorest (figure 3). this is probably related to three stressing factors: a seasonally high water table, alternating with dry conditions, and very low nutrient availability because of the low cec. figure 3: species richness (a), simpson diversity index (b) and species evenness (c) for the eight plant communities in the chobe enclave. the black line is the median, boxes are limited by 1st and 3rd quartiles and the length of the whiskers corresponds to 1.5 sd. the number of sample plots included in the group is indicated above the whiskers in (a). table 2: ranges of selected physicochemical parameters measured in soil samples between 0–15 cm and 15–30 cm of profiles described by romanens et al. (2019), and grouped by plant communities. ipomoea chloroneura – oxygonum alatum woodland this woodland community has a mean tree cover of 15.8% ± 17.9% and a herb cover generally exceeding 80%. it is found on river-washed sands, mainly in fossil alluvial channels or on stabilised sand dunes. although soils are dominated by sand and classified as arenosols (romanens et al. 2019), they contain more silt (2.5% – 7.2%) and organic carbon than aristida junciformis – aristida meridionalis grasslands and have a slightly higher cec (table 2). another major difference is the higher topographic position, resulting in the absence of a visible water table during the wet season. diagnostic species are the woody species grewia retinervis and senegalia cinerea and the forbs acanthosicyos naudinianus, spermacoce senensis, indigofera flavicans, gisekia africana, cleome hirta and hermbstaedtia linearis. many of these species are typical of porous, sandy soils (heath & heath 2009; sianga & fynn 2017). panicum maximum, urochloa trichopus and dichrostachys cinerea are constant species. terminalia sericea is largely dominant in some 75% of the plots, indicating deep sandy soils. this unit was previously described by sianga and fynn (2017) with a very similar species composition and ecological conditions. these plots were amongst the most species rich (45.5 ± 7.7 species; figure 3). baikiaea plurijuga – baphia massaiensis forest these forests are fairly homogeneous in terms of structure and composition with dense woody strata: tree cover generally > 30% and a shrub cover > 8% (table 1). they are established on thick, reddish kalahari sands, and soils are classified as arenosols. these sands were deposited by wind (aeolian). their red colour is because of an ironand aluminum-rich oxyhydroxide coating, which contributes to acidifying the soil (romanens et al. 2019), with the lowest soil ph in the ce (5.7–6.0; table 2). because of the elevation in the east of the geological fault, this plant community is completely disconnected from the hydrology that characterises the ce. as a result, plants have to rely on rainwater, which is quickly drained. the main diagnostic species are the woody species combretum elaeagnoides, baikiaea plurijuga, baphia massaiensis, combretum apiculatum, croton gratissimus and bauhinia petersiana, the grass dactyloctenium giganteum and the forbs jacquemontia tamnifolia, ipomoea pes-tigridis, syncolostemon bracteosus, erlangea misera, chamaecrista falcinella and euphorbia crotonoides. panicum maximum, urochloa trichopus and digitaria eriantha are constant and dominant grasses, and phyllanthus pentandrus, vigna unguiculata, chamaecrista absus and thunbergia aurea are constant forb species. according to sianga and fynn (2017), this plant community occupies large areas of the chobe forest reserve. its presence in northern botswana is part of the western fringe of a very large distribution range in neighbouring zimbabwe. colophospermum mopane – jasminum stenolobum woodland this woodland is dominated by colophospermum mopane in both tree and shrub layers, but with a large variability in size. some areas are characterised by tall specimens, whereas in other places c. mopane is limited to short specimens with a tree cover of < 1%. most of plots were located at slightly higher elevations, corresponding to fossil islands surrounded by sandy, alluvial deposits (mccarthy et al. 2012). the soil data indicate that c. mopane size corresponded to soil conditions. shrubby individuals were found on soils with a silty and hard upper horizon characterised by a high cec saturated by sodium (4.6–11.2 cmol+ kg−1) and with high alkaline ph (9.4–10.2; table 2). these soils were classified as solonchak (romanens et al. 2019). in contrast, tall trees were observed on soils with a sandy upper horizon, with moderate alkaline ph (7.0–8.2), but with the presence of a hard sodic horizon below 30 cm (8.2–11.8 cmol+ kg−1 de na; ph 9.8–11.3). these soils were classified as solonetz or as other soil type with a salic horizon (romanens et al. 2019). all plots were grouped in our clustering, but supplementary surveys may allow a subdivision in two groups corresponding to tree size. besides c. mopane, the other diagnostic species are the poaceae schmidtia pappophoroides and aristida adscensionis, both dominant, the cyperaceae kyllinga buchananii and the forbs ipomoea coptica, clerodendrum ternatum, indigofera trita and talinum caffrum. moreover, dichrostachys cinerea is constant. this community was previously described with a very similar species composition by sianga and fynn (2017), but some of their diagnostic species were rare in the ce. with a mean of 45.6 ± 5.8 species, these plots were amongst the most species rich (figure 3). eragrostis superba – combretum hereroense woodland this woodland is established on a slightly higher elevation than the surrounding grasslands and therefore does not suffer from waterlogging. tree cover is highly variable (range: 0% – 80%, mean: 16.9% ± 21.6%), and shrubs are always present. the soils are particularly fertile, because of the highest clay content of the study area (14.2% – 20.2%), which favours a high cec dominated by calcium (4.7–17.2 cmol+ kg−1), promoting an alkaline ph (table 2). these soils were classified as kastanozems (romanens et al. 2019) and have a particularly low sand proportion (< 35%). the main diagnostic species are the dominant tree combretum hereroense, along with the tree sclerocarya birrea subsp. caffra, the poaceae eragrostis superba, heteropogon contortus, which can be dominant and urochloa oligotricha/brachyuria (two species difficult to distinguish in the field). the main diagnostic forbs are rhynchosia minima, asparagus africanus, pechuel-loeschea leubnitziae, hoslundia opposita and hibiscus caesius (online appendix 2). digitaria eriantha is a constant species, which can have very high cover (up to 70%). other constant species are senegalia nigrescens and solanum panduriforme. cenchrus ciliaris is sometimes highly dominant. the composition of this plant community is variable, as many species can dominate locally but are absent in other plots. with a mean of 47.4 ± 8.6 species, it is particularly species rich, probably because of the heterogeneous tree cover and the regular presence of nitrogen fixing leguminosae (e.g. senegalia nigrescens, philenoptera violacea), inducing a high diversity of conditions on small areas. because a few species are often dominating the community, it has the lowest mean evenness (0.37 ± 0.08). croton megalobotrys – setaria verticillata forest this plant community is fragmented along the floodplains of the linyanti river on substrates located higher than potentially waterlogged lowlands. it has the highest mean tree cover (51.4% ± 27.2%) and the tallest trees (8.4 ± 2.8 m, table 1). compared with other plant communities, the alkaline soil, classified as kastanozem (romanens et al. 2019), showed intermediate values dominated by sand but with 14.9% – 33.2% silt (table 2). species richness is high (47.9 ± 9.8) with 24 diagnostic species (online appendix 2). the main species are the trees berchemia discolor, vachellia tortilis (locally dominant, with a cover up to 50%) and croton megalobotrys, the woody climbers capparis tomentosa and cocculus hirsutus, the shrub grewia flavescens, the poaceae setaria verticillata and setaria sagittifolia, and forbs such as ipomoea dichroa, acalypha fimbriata, justicia heterocarpa, achyranthes aspera, dicliptera paniculata, commelina petersii, cyathula orthacantha, pupalia lappacea, tragia okanyua, leonotis nepetifolia or cocculus hirsutus (online appendix 2). the woody species philenoptera violacea, combretum mossambicense, flueggea virosa and dichrostachys cinerea and the grasses panicum maximum and urochloa trichopus are frequent and locally dominant. finally, the woody species kigelia africana, terminalia prunioides and ficus thonningii are quite rare, but were encountered in this plant community only, all with cover ≥ 10% (online appendix 1). this unit was previously described by sianga and fynn (2017). geigeria schinzii – setaria sphacelata grassland this grassland corresponds to the previously described dambos, defined as seasonally inundated grasslands, in shallow depressions, often characterised by the absence of a stream channel (acres et al. 1985). seasonal waterlogging hinders tree growth (table 1). the low topographic position of these grasslands without a river draining the areas favours the accumulation of fine mineral material. in addition, waterlogging restrains organic matter mineralisation. hence, these soils show the lowest sand content (19.1% – 28.7%), but highest silt (58.8% – 68.2%), organic carbon (3.1% – 5.1%) and cec (21.4–35.9 cmol+ kg−1) compared with other plant communities (table 2). the cec is dominated by calcium (15.6–23.7 cmol+ kg−1), resulting in a neutral to alkaline soil ph. soils were classified as chernozems or phaeozems (romanens et al. 2019). poaceae largely dominate the community, with setaria sphacelata, digitaria milanjiana, hyparrhenia rufa, bothriochloa bladhii, ischaemum polystachyum and leersia hexandra as diagnostic species, completed by the asteraceae geigeria schinzii and linzia glabra (table 2). cynodon dactylon, cymbopogon caesius, panicum repens and trachypogon spicatus are frequent and locally dominant poaceae. species richness is generally low (27.0 ± 3.6). the dominance of a few grass species results in a very low evenness (0.38 ± 0.08). wet floodplain these plots were in areas with frequent or permanent soil waterlogging because of recurring flood events by the linyanti river. these wetlands were insufficiently sampled because deep-flooded areas were difficult to access. instead, plots were selected on the margins of the wetlands. moreover, plots were highly variable, so that no common, good diagnostic species could be found. cynodon dactylon and enicostema axillare have their highest indicator values in wet floodplains, but are also frequent in geigeria schinzii – setaria sphacelata grasslands. therefore, plots were grouped under the generic name of wet floodplains. however, this cannot be considered as a homogenous unit, and supplementary data are needed for a more precise description. the floodplain vegetation could certainly be divided in different plant communities, related to contrasted frequency and flood duration. discussion field sampling of 82 plots allowed a classification by clustering in eight plant communities. these plant communities were subsequently mapped with a supervised classification of remote sensing images. the ce can be divided into four landscape units (figure 4): (1) the floodplains along the linyanti river (north and northwest), fringed with aristida junciformis – aristida meridionalis grasslands, corresponding to the wetlands and dry floodplains on sianga and fynn’s map (2017) and by c. megalobotrys – setaria verticillata forests, mapped as riverine woodland on sianga and fynn’s map (2017); (2) a mosaic of eragrostis superba – combretum hereroense woodlands and geigeria schinzii – setaria sphacelata grasslands in the central area, mapped as acacia grasslands by sianga and fynn (2017); (3) a mosaic of colophospermum mopane – jasminum stenolobum woodlands and ipomoea chloroneura – oxygonum alatum woodlands in the western area, mapped as mopane; and (4) the baikiaea plurijuga – baphia massaiensis forests in the eastern area, extending into the chobe forest reserve. figure 4: vegetation map of the chobe enclave, delineated by the white line (a version in gis file format is available upon request from the corresponding author). floodplains the floodplains are under the influence of two major river systems: the linyanti (called kwando in its upstream part) and the zambezi. the zambezi inflows push water back into the chobe river, as far back as the ce (pricope 2013). these rivers originate in angolan highlands, 750 km further north. the amount of precipitations in both basins during the wet season controls the intensity of flooding and water table levels. peak of flood pulses of both systems occur at different seasons: march–april for the zambezi and june–july for the linyanti (pricope 2013). these flood peaks correspond to the end of the rainy season and dry season, respectively. similar dynamics are observed in the okavango delta, where timing and duration of seasonal flooding are one of the important factors determining vegetation composition (murray-hudson et al. 2011). seasonal flooding creates a zonation from permanent water (lake liambezi in the north) to areas with irregular flooding only. as stated above, our data set in the wetlands is insufficient to completely translate the diversity of plant communities. supplementary data would allow the subdivision of the wet floodplains in various plant communities, as observed in the okavango delta (murray-hudson et al. 2011). a better description of these floodplains would be especially necessary as they are highly important for water and fodder availability to the wildlife during the dry season, with the gradient of flooding corresponding to variations in foraging time (fynn et al. 2015). although floods are rare and shorter in the aristida junciformis – aristida meridionalis grasslands (higher topographical position), the water table is regularly high enough to reach plant roots. for plant growth, these dry floodplains present quite stressful conditions, which alternate between waterlogged soils at high water and very dry soils (well-draining sand) at the end of the dry season. woody species of this plant community cannot cope with waterlogged periods, whereas with similar soils but clearly above the water table, trees and shrubs are an important component of the ipomoea chloroneura – oxygonum alatum woodlands. in dry ecosystems such as savannahs, trees are susceptible to facilitate grass growth by hydraulic lift (priyadarshini et al. 2016), although the importance of this facilitation for the understorey is discussed (see ludwig et al. 2004). however, facilitation does not occur in aristida grasslands because of the absence of trees. this could explain the low species richness. an unmistakable characteristic of these dry floodplains is the numerous evenly distributed termite mounds (mujinya et al. 2014), often occupied by trees (mccarthy et al. 1998). these mounds were excluded from our inventory because local ecological conditions were deemed too heterogeneous to fit into our sampling design. they rise above the floodplain, are rich in silt, clay, organic matter and nutrients and have a ph of 8–10 (jouquet et al. 2011; romanens et al. 2019). some tree species occur almost exclusively on termite mounds (e.g. phoenix reclinata, imperata cylindrica; roodt 1998). supplementary smaller plots would be necessary to get a better knowledge of their species composition and variability. at the margins of the floodplains are the riverine forests, classified as croton megalobotrys – setaria verticillata forest. these forests are situated high enough to avoid flooding but low enough for riparian trees to reach the water table (ellery & tacheba 2003). this ensures that trees have permanent access to water. these ideal conditions allow a high tree cover and the presence of many grasses and forbs. this results in a high species richness and α-diversity (figure 3). many woody species are exclusively or mainly encountered in these forests. termite mounds are also common, probably explaining the higher proportion of clay and silt in the soil, its higher cec and alkaline ph (romanens et al. 2019). similar to wet floodplains, riverine vegetation provides an important dry season habitat for browsers and grazers owing to its ability to maintain green leaves over the dry season (kleynhans et al. 2011). central area in the central area, both distance to river and slope decreases flood frequency. the floodplains are replaced by a mosaic of depressions and slightly more elevated land. this microrelief creates contrasted ecological conditions, leading to the establishment of two distinct plant communities. the geigeria schinzii – setaria sphacelata grasslands (dambos) occupy the depressions, whereas the eragrostis superba – combretum hereroense woodlands are located on higher-lying grounds. the woodland substrate is dominated by a silty loam soil, rich in calcium carbonate (romanens et al. 2019). this substrate, with high calcium and clay content in a sand-dominated landscape, probably originates from sedimentation in palaeolakes (burrough & thomas 2008; diaz et al. 2019). the dambos do not have trees and are regarded as shallow, seasonally waterlogged, tropical and subtropical african wetlands. they are common in central and southern africa, where an annual rainfall of 500 mm – 1500 mm is concentrated over a wet season (acres et al. 1985). although they share the seasonal waterlogging with floodplains, the origin and timing of waterlogging differ. in dambos, the clay content of soils is responsible for retaining water and saturating the soil during the rainy season (bullock 1992), whereas hydrological inflow of the floodplains originates from the angolan highlands and arrives in the dry season. in dambos, waterlogging persists long enough to restrain organic matter mineralisation (romanens et al. 2019). the high organic carbon and clay content explain the high cec value, which induces a higher soil fertility than most of the other plant communities (weil & brady 2017). these conditions allow tall grasses, mostly setaria sphacelata, cymbopogon caesius and hyparrhenia rufa, to densely grow. this high density may explain the low species richness, as productive habitats have been shown to have the lowest species richness because of competitive exclusion of smaller species by the large dominants (borer et al. 2014). these grasslands represent an important grazing resource by providing reliable green forage for zebra and other large herbivores during the dry season, especially after burning of old stubble (fynn et al. 2014). indeed, herbivores can graze the young, green grasses whilst the lack of trees provides high visibility around them as security against predators. the eragrostis superba – combretum hereroense woodlands have the same high clay and calcium content with a high cec, and we can expect that these woodlands offer good quality forage also and shelter to the herbivores. western area in the western part of the ce, a mosaic can be observed of colophospermum mopane – jasminum stenolobum and ipomoea chloroneura – oxygonum alatum woodlands. the latter corresponds to sandveld. from a bird’s-eye view, this mosaic resembles a fossil delta with islands of colophospermum woodlands separated by channels of sandveld. old river meanders are locally clearly visible. this geomorphological pattern is inherited from ancient hydrological dynamics, with infilling of the channels by kalahari sands (fynn et al. 2014; kinabo et al. 2008). colophospermum mopane is a deciduous tree on calcareous or sodic soils with a high clay content (mapaure 1994; roodt 1998). its shallow root system enables growth on poorly drained substrates (smit & rethman 1998). in the ce, the soil conditions of these woodlands (high ph and cec dominated by sodium) are similar to those of the islands of the okavango delta, where island centres are devoid of trees and shrubs because of a high sodium content (ramberg & wolski 2008). this sodium constitutes a major osmotic stress for plants, reduces nutrient absorption and rainwater infiltration and tends to form a crust (zhu 2001). nevertheless, c. mopane tolerates these high alkaline conditions (werger & coetzee 1978). this woodland provides an ideal habitat for large herbivores such as buffalo, roan and sable antelopes or elephants. the leaves of c. mopane have a high nutritional value, with a high protein and phosphorus content, even when they have fallen on the ground (roodt 1998). moreover, the high clay content induces seasonal waterholes, and the salty soils under the patches of shrubby c. mopane provide saline grazing for wildlife (sianga et al. 2017; sianga & fynn 2017). dye and walker (1980), mapaure (1994) and mlambo (2007) reported that c. mopane was characterised by a shrub structure on shallow, sodium-rich soils. in the ce, the soil under these stands had a hard upper horizon to dig, and crusts were observed. conversely, tall mopanes grow on a 30 cm – 75 cm deep layer of sand covering a hard sodium-rich horizon (romanens et al. 2019; roodt 1998), in line with fraser, van rooyen and verster’s (1987) observations. other studies assigned the short stature to elephant browsing (smallie & o’connor 2000) or frequent fires (mlambo & mapaure 2006), but the frequent presence of both forms close to each other, with distinct boundaries between them, indicates that soil chemistry is an important factor. however, our observations are based on a limited number of soil descriptions. supplementary information would be necessary to confirm this separation. in contrast to the colophospermum mopane – jasminum stenolobum woodlands, the ipomoea chloroneura – oxygonum alatum woodlands are characterised by very poor nutrient conditions. the scarcity of silt and clay induces dry and nutrient-poor conditions for plants. indeed, the sandy structure has lower water retention and cec than silty or clayey textures. in addition, nutrients are leached during the wet season. consequently, soils are deficient in phosphorus and nitrogen (roodt 1998; sianga & fynn 2017). in the fossil palaeo-river channels, the proximity of very alkaline and fine material from colophospermum woodland soils can explain the approximately neutral ph. in contrast, the baikiaea plurijuga – baphia massaiensis forests are characterised by acidic soils because of aluminium (romanens et al. 2019). these differences in ph induce different species compositions, but both units are attractive for grazers because they share a high cover of the palatable grasses digitaria eriantha and panicum maximum (sianga et al. 2017; sianga & fynn 2017). functional habitat heterogeneity for herbivores our detailed vegetation study, in combination with the parallel soil study, provides insight into the functional diversity of these habitats. vegetation heterogeneity at a regional scale enables herbivores to adapt to changing conditions throughout the year (sianga & fynn 2017). this study adds extra knowledge by (1) linking vegetation to soil characteristics and (2) showing the presence of previously undescribed plant communities for northern botswana, such as dambos (geigeria schinzii – setaria sphacelata grassland) or eragrostis superba – combretum hereroense woodland on calciumand clay-rich substrates and (3) by showing some interesting soil chemistry–vegetation community relationships, such as colophospermum mopane – jasminum stenolobum woodland on saline soils. herbivore population growth and reproduction critically depend on access to forage with high concentrations of energy, protein and key minerals such as calcium, sodium or phosphorus (murray 1995; parker, barboza & gillingham 2009). herbivores can access these resources by migrating to saline palaeolake systems (nxai pan, makgadikgadi pans and the mababe depression; bartlam-brooks, bonyongo & harris 2011; fynn et al. 2014; naidoo et al. 2016). it is less clear, however, how non-migratory herbivores, such as buffalo, roan or sable antelopes (bennitt et al. 2014; haveman 2014; hensman et al. 2013; sianga et al. 2017), are able to access forage with sufficient concentrations of minerals for reproductive purposes. we can now hypothesise that the mosaic of c. mopane woodlands and sandveld provides a possible answer to this question. the high abundance of leafy grasses in sandveld, such as digitaria eriantha and panicum maximum, likely provide sufficient intake of digestible forage to meet energy and protein requirements, whilst plants and drinking water in pans in the c. mopane woodlands meet their mineral requirements. these two habitats occur in an extensive mosaic across the region so that herbivores can easily move between these functionally different habitats. during the dry season, when herbivores requires less resources (no lactation or rapid growth requirements), the availability of forage and its quality declines greatly (owen-smith 2008). without access to functional key resource habitats to provide reserves of adequate-quality forage for survival, herbivore populations would collapse during drought years (illius & o’connor 2000; walker et al. 1987). in the ce, the floodplains, dambos and riverine forests provide access to grasses and a shallow water table that enables the maintenance of green forage during the dry season. the simultaneous flood of the linyanti river ensures a reliable source of drinking water (fynn et al. 2014). the vegetation map limitations based on field experience and a lower resolution map (sianga & fynn 2017), we are confident that our vegetation map (figure 4) corresponds to the distribution of forests, woodlands and grasslands. moreover, it reflects the general distribution of the various plant communities. compared with sianga and fynn (2017), its higher resolution enabled the elucidation of small-sized plant communities that are locally restricted to the ce, such as geigeria schinzii – setaria sphacelata grasslands. however, some pixels of the grasslands can be misinterpreted. for example, the presence of geigeria schinzii – setaria sphacelata grassland on the edges of the floodplains does not correspond to their distribution and ecological conditions. the light blue areas shown in northern part of figure 4 are more probably variations amongst wet floodplains. conversely, the dark blue patches (wet floodplains) around the centre and south of the map ought probably to be light blue (figure 4), as these sites are too far away and too isolated from the river to be flooded, but only stay wet during the rainy season. a possible explanation is that the similar vegetation structure of these grasslands, with common species, and their wet soils resulted in too close a reflectance in remote sensing data for accurate discrimination. transition zones can be expected amongst woodlands with intermediate conditions from eragrostis superba – combretum hereroense woodland to colophospermum mopane – jasminum stenolobum woodland (romanens et al. 2019). finally, isolated pixels attributed to croton megalobotrys – setaria verticillata forest or to baikiaea plurijuga – baphia massaiensis forest in the central part of the map are artefacts because of an insufficient number of pixels covering small patches of trees (figure 4). conclusion at the landscape scale, the ce is flat and dominated by sand. however, over time, water and tectonic activity diversified the microtopography and chemistry of the substrates. this microtopography also induced a gradient of soil texture, parallel to the flooding potential. all these variations create a mosaic of soil units, with their respective nutrient availability (romanens et al. 2019). the same parameters and their corresponding soils are the environmental drivers for a diversity of plant communities. these plant communities offer a diversity of resources, allowing some herbivores to live in the area all the year round. supplementary data of the wet floodplain and in areas of permanent water are needed to classify these plant communities at a similar level of detail. the vegetation classification and mapping of this spatially heterogenous landscape will provide an excellent baseline for future studies on seasonal habitat use by herbivores in the ce. acknowledgements we express our sincere thanks to the ministry of environment, natural resources conservation and tourism of the republic of botswana as well as the chobe enclave conservation trust for the research permits, without which this study would not have been possible. we are grateful to the three anonymous reviewers and to jesse m. kalwij, whose comments helped us to improve our manuscript, and to karin verrecchia, who kindly edited the revised version of the manuscript. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced this article. authors’ contributions p.v., f.p., e.p.v. and r.f. designed the study; f.p., r.r., a.m., p.v. and r.f. realised the field work; f.p. and p.v. analysed the data; and all authors contributed to the writing of the manuscript. ethical consideration no ethical clearance was required for this study. funding information this work has been partly supported by a swiss national foundation grant no. 200021_172944 to eric p. verrecchia. data availability statement all data are available in the online version of this article as online appendix 1 (table of the 82 plant inventory plots) and online appendix 2 (synoptic tables for the 8 plant communities). disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references acres, b.d., blair rains, a., king, r.b., lawton, r.m., mitchell, a.j.b. & rackham, l.j., 1985, ‘african dambos: their distribution, characteristics and use’, zeitschrift fur geomorphologie 52(1), 62–83. bartlam-brooks, h.l.a., bonyongo, m.c. & harris, s., 2011, ‘will reconnecting ecosystems allow long-distance mammal migrations to resume? a case study of a zebra equus burchelli migration in botswana’, oryx 45(2), 210–216. https://doi.org/10.1017/s0030605310000414 bennitt, e., bonyongo, m.c. & harris, s., 2014, ‘habitat selection by african buffalo (syncerus caffer) in response to landscape-level fluctuations in water availability on two temporal scales’, plos one 9(7), e101346. https://doi.org/10.1371/journal.pone.0101346 borer, e.t., seabloom, e.w., gruner, d.s., harpole, w.s., hillebrand, h., lind, e.m., et al., 2014, ‘herbivores and nutrients control grassland plant diversity via light limitation’, nature 508(7497), 517–520. https://doi.org/10.1038/nature13144 bullock, a., 1992, ‘dambo hydrology in southern africa – review and reassessment’, journal of hydrology 134(1–4), 373–396. https://doi.org/10.1016/0022-1694(92)90043-u burrough, s.l. & thomas, d.s.g., 2008, ‘late quaternary lake-level fluctuations in the mababe depression: middle kalahari palaeolakes and the role of zambezi inflows’, quaternary research 69(3), 388–403. https://doi.org/10.1016/j.yqres.2008.02.003 chase, m., 2013, status of wildlife populations and land degradation in botswana’s forest reserves and chobe district, unpublished report, forest conservation botswana, elephants without borders, zoological society of san diego. de cáceres, m., chytrý, m., agrillo, e., attorre, f., botta-dukát, z., capelo, j. et al., 2015, ‘a comparative framework for broad-scale plot-based vegetation classification’, applied vegetation science 18(4), 543–560. https://doi.org/10.1111/avsc.12179 diaz, n., armitage, s.j., verrecchia, e.p. & herman, f., 2019, ‘osl dating of a carbonate island in the chobe enclave, nw botswana’, quaternary geochronology 49(1), 172–176. https://doi.org/10.1016/j.quageo.2018.03.001 dufrêne, m. & legendre, p., 1997, ‘species assemblages and indicator species: the need for a flexible asymmetrical approach’, ecological monographs 67(3), 345–366. https://doi.org/10.1890/0012-9615(1997)067[0345:saaist]2.0.co;2 dye, p. & walker, b., 1980, ‘vegetation-environment relations on sodic soils of zimbabwe-rhodesia’, journal of ecology 68(2), 589–606. https://doi.org/10.2307/2259424 ellery, w.n. & tacheba, b., 2003, ‘floristic diversity of the okavango delta, botswana. a rapid biological assessment of the aquatic ecosystems of the okavango delta, botswana: high water survey’, rap-bulletin of biological assessment 25(1), 69–96. esri, 2019, world imagery and national geographic base maps, sources: esri, digitalglobe, geoeye, earthstar geographics, cnes/airbus ds, usda, usgs, aerogrid, ign, and the gis user community. ewald, j., 2003, ‘a critique for phytosociology’, journal of vegetation science 14(2), 291–296. https://doi.org/10.1111/j.1654-1103.2003.tb02154.x fraser, s.w., van rooyen, t.h. & verster, e., 1987, ‘soil-plant relationships in the central kruger national park’, koedoe 30(1), 19–34. https://doi.org/10.4102/koedoe.v30i1.499 fynn, r.w.s., chase, m. & röder, a., 2014, ‘functional habitat heterogeneity and large herbivore seasonal habitat selection in northern botswana’, south african journal of wildlife research 44(1), 1–15. https://doi.org/10.3957/056.044.0103 fynn, r.w.s., murray-hudson, m., dhliwayo, m. & scholte, p., 2015, ‘african wetlands and their seasonal use by wild and domestic herbivores’, wetlands ecology and management 23(1), 559–581. https://doi.org/10.1007/s11273-015-9430-6 gongedo, l., 2017, semi-automatic classification plugin documentation, release 5.3.6.1, viewed 28 september 2020, from https://semiautomaticclassificationmanual-v5.readthedocs.io/en/latest/ hartigan, j.a., 1975, clustering algorithms, wiley, new york, ny. haveman, c.p., 2014, ‘population dynamics and foraging ecology of roan antelope in northern botswana’, msc thesis, university of pretoria. heath, r. & heath, a., 2009, field guide to the plants of northern botswana including the okavango delta, royal botanic gardens, richmond. hensman, m.c., owen-smith, n., parrini, f. & bonyongo, m.c., 2013, ‘resource use and the nutritional status of sable antelope in the okavango delta region of northern botswana’, african journal of ecology 52(3), 237–245. https://doi.org/10.1111/aje.12113 illius, a.w. & o’connor, t.g., 2000, ‘resource heterogeneity and ungulate population dynamics’, oikos 89(2), 283–294. https://doi.org/10.1034/j.1600-0706.2000.890209.x ischer, m., dubuis, a., keller, r. & vittoz, p., 2014, ‘a better understanding of ecological conditions for leontopodium alpinum cassini in the swiss alps’, folia geobotanica 49(4), 541–558. iuss working group wrb, 2014, world reference base for soil resources 2014: international soil classification system for naming soils and creating legends for soil maps, fao, rome. jones, b., 2002, chobe enclave, botswana – lessons learnt from a cbnrm project 1993–2002, cbnrm support project, gaborone. jouquet, p., traoré, s., choosai, c., hartmann, c. & bignell, d., 2011, ‘influence of termites on ecosystem functioning. ecosystem services provided by termites’, european journal of soil biology 47(4), 215–222. https://doi.org/10.1016/j.ejsobi.2011.05.005 kinabo, b.d., hogan, j.p., atekwana, e.a., abdelsalam, m.g. & modisi, m.p., 2008, ‘fault growth and propagation during incipient continental rifting: insights from a combined aeromagnetic and shuttle radar topography mission digital elevation model investigation of the okavango rift zone, northwest botswana’, tectonics 27(3), tc3013. https://doi.org/10.1029/2007tc002154 kleynhans, e.j., jolles, a.e., bos, m.r.e. & olff, h., 2011, ‘resource partitioning along multiple niche dimensions in differently sized african savanna grazers’, oikos 120(4), 591–600. https://doi.org/10.1111/j.1600-0706.2010.18712.x kyalangalilwa, b., boatwright, j.s., daru, b.h., maurin, o. & van der bank, m., 2013, ‘phylogenetic position and revised classification of acacia s.l. (fabaceae: mimosoideae) in africa, including new combinations in vachellia and senegalia’, botanical journal of the linnean society 172(4), 500–523. https://doi.org/10.1111/boj.12047 ludwig, f., dawson, t.e., prins, h.h.t., berendse, f. & de kroon, h., 2004, ‘below-ground competition between trees and grasses may overwhelm the facilitative effects of hydraulic lift’, ecology letters 7(8), 623–631. https://doi.org/10.1111/j.1461-0248.2004.00615.x mapaure, i., 1994, ‘the distribution of colophospermum mopane (leguminosae-caesalpinioideae) in africa’, kirkia 15(1), 1–5. mccarthy, t.s., 2013, ‘the okavango delta and its place in the geomorphological evolution of southern africa’, south african journal of geology 116(1), 1–54. https://doi.org/10.2113/gssajg.116.1.1 mccarthy, t.s., ellery, w.n. & dangerfield, j.m., 1998, ‘the role of biota in the initiation and growth of islands on the floodplain of the okavango alluvial fan, botswana’, earth surface processes and landforms 23(4), 291–316. mccarthy, t.s., humphries, m.s., mahomed, i., le roux, p. & verhagen, b.th., 2012, ‘island forming processes in the okavango delta, botswana’, geomorphology 179(1), 249–257. https://doi.org/10.1016/j.geomorph.2012.08.016 mcclean, c.j., doswald, n., kuper, w., sommer, j.h., barnard, p. & lovett, j.c., 2006, ‘potential impacts of climate change on sub-saharan african plant priority area selection’, diversity and distributions 12(6), 645–655. https://doi.org/10.1111/j.1472-4642.2006.00290.x mlambo, d., 2007, ‘influence of soil fertility on the physiognomy of the african savanna tree colophospermum mopane’, african journal of ecology 45(1), 109–111. https://doi.org/10.1111/j.1365-2028.2006.00676.x mlambo, d. & mapaure, i., 2006, ‘post-fire resprouting of colophospermum mopane saplings in a southern african savanna’, journal of tropical ecology 22(2), 231–234. https://doi.org/10.1017/s026646740500297x mujinya, b.b., adam, m., mees, f., bogaert, j., vranken, i., erens, h. et al., 2014, ‘spatial patterns and morphology of termite (macrotermes falciger) mounds in the upper katanga, dr congo’, catena 114(1), 97–106. https://doi.org/10.1016/j.catena.2013.10.015 murray, m.g., 1995, ‘specific nutrient requirements and migration of wildebeest’, in a.r.e. sinclair & p. arcese (eds.) serengeti ii: dynamics, management, and conservation of an ecosystem, pp. 231–256, the university of chicago press, chicago, il. murray-hudson, m., combs, f., wolski, p. & brown, m.t., 2011, ‘a vegetation-based hierarchical classification for seasonally pulsed floodplains in the okavango delta, botswana’, african journal of aquatic science 36(3), 223–234. https://doi.org/10.2989/16085914.2011.636904 naidoo, r., chase, m.j., beytell, p., du preez, p., landen, k., stuart-hill, g. et al., 2016, ‘a newly discovered wildlife migration in namibia and botswana is the longest in africa’, oryx 50(1), 138–146. https://doi.org/10.1017/s0030605314000222 owen-smith, n., 2008, ‘effects of temporal variability in resources on foraging behaviour’, in h.h.t. prins & f. van langevelde (eds.), resource ecology: spatial and temporal dynamics of foraging, pp. 159–181, springer, dordrecht. parker, k.l., barboza, p.s. & gillingham, m.p., 2009, ‘nutrition integrates environmental responses of ungulates’, functional ecology 23(1), 57–69. https://doi.org/10.1111/j.1365-2435.2009.01528.x perumal, k. & bhaskaran, r., 2010, ‘supervised classification performance of multispectral images’, journal of computing 2(2), 124–129. pricope, n.g., 2013, ‘variable-source flood pulsing in a semi-arid transboundary watershed: the chobe river, botswana and namibia’, environmental monitoring and assessment 185(2), 1883–1906. https://doi.org/10.1007/s10661-012-2675-0 pricope, n.g., gaughan, a.e., all, j.d., binford, m.w. & rutina, l.p., 2015, ‘spatio-temporal analysis of vegetation dynamics in relation to shifting inundation and fire regimes: disentangling environmental variability from land management decisions in a southern african transboundary watershed’, land 4(3), 627–655. https://doi.org/10.3390/land4030627 priyadarshini, k.v.r., prins, h.h.t., de bie, s., heitkonig, i.m.a., woodborne, s., gort, g. et al., 2016, ‘seasonality of hydraulic redistribution by trees to grasses and changes in their water-source use that change tree-grass interactions’, ecohydrology 9(2), 218–228. https://doi.org/10.1002/eco.1624 ramberg, l. & wolski, p., 2008, ‘growing islands and sinking solutes: processes maintaining the endorheic okavango delta as a freshwater system’, plant ecology 196(2), 215–231. https://doi.org/10.1007/s11258-007-9346-1 romanens, r., pellacani, f., mahinga, a., fynn, r., vittoz, p. & verrecchia, e.p., 2019, ‘soil diversity and major soil processes in the kalahari basin, botswana’, geoderma regional 19(1), e00236. https://doi.org/10.1016/j.geodrs.2019.e00236 roodt, v., 1998, trees & shrubs of the okavango delta: medicinal uses and nutritional value, shell oil botswana (pty) ltd, maun. roodt, v., 2015, grasses & grazers of botswana and the surrounding savanna, struik nature, cape town. sala, o.e., chapin, f.s., armesto, j.j., berlow, e., bloomfield, j., dirzo, r. et al., 2000, ‘biodiversity – global biodiversity scenarios for the year 2100’, science 287(5459), 1770–1774. https://doi.org/10.1126/science.287.5459.1770 schmidt, c., 2017, advanced land cover classification webinar series, applied remote sensing training (arset), nasa, viewed 28 september 2020, from https://arset.gsfc.nasa.gov/land/webinars/advanced-land-classification. sianga, k. & fynn, r., 2017, ‘the vegetation and wildlife habitats of the savuti-mababe-linyanti ecosystem, northern botswana’, koedoe 59(2), 1–16. https://doi.org/10.4102/koedoe.v59i2.1406 sianga, k., van telgen, m., vrooman, j., fynn, r.w.s. & van langevelde, f., 2017, ‘spatial refuges buffer landscapes against homogenisation and degradation by large herbivore populations and facilitate vegetation heterogeneity’, koedoe 59(2), a1434. https://doi.org/10.4102/koedoe.v59i2.1434 smallie, j.j. & o’connor, t.g., 2000, ‘elephant utilization of colophospermum mopane: possible benefits of hedging’, african journal of ecology 38(4), 352–359. https://doi.org/10.1046/j.1365-2028.2000.00258.x smit, g.n. & rethman, n.f.g., 1998, ‘root biomass, depth distribution and relations with leaf biomass of colophospermum mopane’, south african journal of botany 64(1), 38–43. https://doi.org/10.1016/s0254-6299(15)30825-5 walker, b.h., emslie, r.h., owen-smith, r.n. & scholes, r.j., 1987, ‘to cull or not to cull: lessons from a southern african drought’, journal of applied ecology 24(2), 381–401. https://doi.org/10.2307/2403882 walker, d.a., daniels, f.j.a., alsos, i., bhatt, u.s., breen, a.l., buchhorn, m. et al., 2016, ‘circumpolar arctic vegetation: a hierarchic review and roadmap toward an internationally consistent approach to survey, archive and classify tundra plot data’, environmental research letters 11(5), 055005. https://doi.org/10.1088/1748-9326/11/5/055005 weil, r.r. & brady, n.c., 2017, the nature and properties of soils, 15th edn., pearson, harlow. werger, m.j.a. & coetzee, b.j., 1978, ‘the sudano-zambezian region’, in m.j.a. werger (ed.), biogeography and ecology of southern africa, pp. 301–462, springer, dordrecht. wildi, o., 2010, data analysis in vegetation ecology, wiley-blackwell, chichester. zhu, j.k., 2001, ‘plant salt tolerance’, trends in plant science 6(2), 66–71. https://doi.org.10.1016/s1360-1385(00)01838-0 appendices appendix 1 figure 1-a1: true colour landsat 8 images of the chobe enclave for the different seasons: february is in the middle of the rainy season, may at the end of the rainy season and september in the dry season. appendix 2 figure 1-a2: distribution of the 82 sampling plots in the chobe enclave. the inset on the right is an enlargement of the central rectangle. appendix 3 table 1-a3: coordinates of the 82 sampling plots in the chobe enclave. measurements refer to the centre of the plot and were taken with a garmin gps (precision 3–5 m). coordinates are indicated according to the utm zone 35south (wgs 84). appendix 4 figure 1-a4: hierarchical clustering of the plant inventory plots (n = 82) in the chobe enclave based on the bray-curtis distance and the ward’s minimum variance method. filelist convert a pdf file! page 1 page 2 page 3 page 4 abstract introduction methods results discussion conclusion acknowledgements references about the author(s) johan a. baard garden route national park, south african national parks, south africa tineke kraaij school of natural resource management, nelson mandela university, south africa citation baard, j.a. & kraaij, t., 2019, ‘use of a rapid roadside survey to detect potentially invasive plant species along the garden route, south africa’, koedoe 61(1), a1515. https://doi.org/10.4102/koedoe.v61i1.1515 original research use of a rapid roadside survey to detect potentially invasive plant species along the garden route, south africa johan a. baard, tineke kraaij received: 17 jan. 2018; accepted: 07 oct. 2018; published: 28 jan. 2019 copyright: © 2019. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract early detection of emerging or sleeper weeds and monitoring of alien plant invasions forms the foundation of effective invasive alien plant management. using a rapid roadside survey technique, we aimed to (1) establish a baseline of alien plant distribution and abundance along roads in the garden route, south africa, and (2) identify priority species (particularly sleeper weeds) and priority areas to inform appropriate management action. we surveyed along 530 km of roads and recorded 109 alien and/or extralimital species across 1942 point locations. of these, 35 species were considered to be sleeper weeds on account of displaying estimated dispersal distances distinctive of invasive plants and not being listed by the south african plant invaders atlas (sapia) or regulated by south african legislation. roads along natural forest and fynbos vegetation (often within the garden route national park) displayed lower incidences of alien plants than those associated with degraded or transformed land, with roads along farmland associated with the highest incidences of alien plants. roads in the southern cape region had more species and higher densities of alien plants than roads in the tsitsikamma region, and a few species were exclusive to either. conservation implications: our inventory contributed significant new records and range extensions to sapia, while the identified sleeper weeds offered suggestions for species that may be considered for regulation under the national environmental management: biodiversity act of south africa. we furthermore provided information to facilitate (1) timely management of emerging invasive alien plants, (2) prioritisation of species and areas for management action, and (3) future monitoring of alien plants in the garden route national park and surrounds. introduction worldwide, invasive alien plants are considered to be a major threat to biodiversity and ecosystem services, with the threat deemed to be increasing (maxwell et al. 2016). emerging alien species, also called sleeper weeds, are species previously thought to be benign but that may turn invasive; or species that have naturalised but not yet expanded their populations exponentially; or species currently only present in a small area but having the potential to spread widely (boy & witt 2013; cunningham et al. 2003; groves 1999; thuiller et al. 2005; williams & west 2000). these species should be closely monitored and assessed for their invasion capacity and timeous decisions made on management actions to be taken. early detection forms a key component of invasive plant management (brooks & klinger 2009) with rapid response in the form of eradication of single plants or small invasions being among the best investments that can be made (rejmánek & pitcairn 2002). rapid response is more effective than delayed implementation of alien plant management strategies (millar, stephenson & stephens 2007; wilson et al. 2014). knowledge of the incidence of alien plants and monitoring of changes in their status underpin sound management interventions (blackburn et al. 2014; wilson et al. 2014) and as such are mandated by the national environmental management: biodiversity act (nemba, act 10 of 2004) in south african protected areas (foxcroft et al. 2017; wilson et al. 2017). roadside surveys are a timeand cost-effective technique that may be used to detect alien plants as they enable relatively rapid collection of substantial quantities of data (henderson 2007; henderson & musil 1984; kalwij, milton & mcgeoch 2008; shackleton et al. 2017). roadside surveys have the disadvantage that only plants in the road verge and immediate surrounds can be detected, and they are biased towards larger, conspicuous growth forms (henderson & wilson 2017). in contrast, walking transects are more precise but cover smaller areas per unit of time and effort. aerial surveys on the other hand are comparatively expensive, and with both aerial surveys and remote sensing techniques, spatial resolution is coarse (vitousek, d’antonio & asner 2010). detection with these techniques is also limited to the most abundant and largest taxa, which will not enable the recording of a diversity of potentially uncommon species, including sleeper weeds. roads and roadsides present disturbances that are known to facilitate invasion (christen & matlack 2006; kalwij et al. 2008; mortensen et al. 2009), in addition to acting as accidental or unintentional pathways for alien plant dispersal (pickering & mount 2010; woodward & quinn 2011). roadsides may accordingly be expected to harbour more alien plants than undisturbed habitat. roadside surveys thus present as a potentially timeand cost-efficient technique to detect alien plants over extensive areas, and they have the additional advantage that they may be combined with other activities (henderson & musil 1984). species able to disperse widely have an increased chance of invasion success (alpert, bone & holzapfel 2000; kolar & lodge 2001; lloret et al. 2004); accordingly, small seed mass is often an indicator of invasiveness (rejmánek & richardson 1996). light seeds have the potential to spread much further by wind or other means than larger seeds or fruit (rejmánek & richardson 1996). however, birds are particularly effective long-distance seed vectors, including the seed of larger fruit (lack & evans 2005). understanding an alien plant species’ dispersal strategy and abilities can inform the appropriate approach to, and priority of, its control. dispersal rates are also used to classify species’ invasion status (pyšek et al. 2004). invasive alien plants are considered to be the leading ecological threat to the garden route national park (grnp) (sanparks 2010) along the southern cape coast of south africa. the grnp is a long and fragmented protected area made up of 29 discontinuous land parcels spreading over an area of 150 km × 35 km. it has a large edge-to-area ratio with various adjacent and protruding land uses, such as urban settlements, agriculture, commercial forestry and conservation (kraaij, cowling & van wilgen 2011). fragmentation and accessibility associated with a well-developed road network within and outside of the grnp make the park particularly susceptible to the introduction and spread of alien plant species (alston & richardson 2006; davies & sheley 2007; jodoin et al. 2008; mortensen et al. 2009; pickering & mount 2010). baard and kraaij (2014) compiled an inventory of 244 alien plant species in the grnp and assigned invasion status to species based on expert opinion, but they did not systematically assess the geographic distribution and abundances of these species in the area. using a rapid roadside survey technique, we aimed to (1) establish a baseline of alien plant status along roads in the area of the grnp that would allow for assessment of future changes in the status of invasions and the effectiveness of management interventions; (2) improve understanding of the spatial distribution of alien plants (including previously unrecorded species) within and outside of the grnp to inform priority areas for management and future risks; and (3) identify sleeper weeds (i.e. invasive species that have not yet been recognised as such in the region) to facilitate timely management responses to be taken by the grnp. methods study area the study was conducted along roads in and around the grnp (33.80°s 22.50°e to 34.15°s 24.20°e; 145 000 ha; approximate east–west extent of 150 km) between wilderness village in the west and storms river village in the east, with the outeniqua and tsitsikamma mountains to the north and the indian ocean to the south (figure 1). the area is divided into two regions, the southern cape in the west and tsitsikamma in the east. the pretransformed terrestrial vegetation of the study area was primarily fynbos shrublands and afrotemperate forest (vlok, euston-brown & wolf 2008), much of which is now farmland, commercial timber plantations, alien plant invasions and towns (wilderness, sedgefield, knysna, plettenberg bay and storms river) that surround much of the grnp (vromans et al. 2010). figure 1: the study area showing the garden route national park, the southern cape and tsitsikamma regions, the roads surveyed and the land cover types adjacent to roads. the topography of the study area is characterised by rugged mountains, foothills and coastal plains (altitudinal range 0–1675 meters above sea level). deep ravines are cut through the landscape by north–south flowing rivers. the cape fold belt dominates the geology and largely consists of table mountain sandstone with smaller areas of the bokkeveld group, enon formation, george granites and kaaimans formation (tham & johnson 2006). aeolian sands deposited during the quaternary cover the coastal plateau (illenberger 1996). mean annual rainfall ranges between 800 mm and 1100 mm and increases from west to east and from south to north with an increase in altitude. rain falls throughout the year but peaks in spring and autumn (tyson & preston-whyte 2000). in winter, frost occurs in some areas and snow on the higher mountain peaks. summers are mildly warm (22 °c – 25 °c) and winters mild (18 °c – 21 °c). data collection and analysis five hundred and thirty kilometres of roads were surveyed for the presence of alien plants over a period of 11 days during the period december 2016 to february 2017. the total cost of our roadside surveys amounted to approximately r29 000.00 (r22 000.00 for observer salary and r7000.00 for transport). the n2 national road, all main and secondary roads outside of urban areas or villages, and roads accessible to the public inside the park (hereafter ‘park roads’) were surveyed (figure 1). road classification followed the roads’ geographical information system (gis) dataset in sa explorer version 3.0 (2004). one driver in a vehicle recorded the presence of all alien plant species visible within or immediately adjacent to road reserves. the national road was tarred, the main and secondary roads tarred or gravelled, and the park roads gravelled. on the n2 speeds of 20 km/h – 40 km/h or less was maintained compared to only 10 km/h – 20 km/h on lesser roads. the focus taxa were alien trees, shrubs, conspicuous creepers (including climbers and scramblers), conspicuous ferns and succulents, whereas inconspicuous herbs (including inconspicuous grasses) were not considered. the most common transformer species in the region – acacia mearnsii (black wattle), a. melanoxylon (blackwood), a. cyclops (redeye), a. saligna (port jackson willow), solanum mauritianum (bugweed), pinus (pine) species and eucalyptus (gum) species were not recorded. extralimital species (i.e. occurring outside of their native distribution ranges indicated by manning & goldblatt 2012) were also recorded. a trimble juno 3b gps was used with a recording sequence set up in advance, using cybertracker software. information recorded included species, locality, abundance, age class, whether the alien was planted, and direction from the road (north, south, east or west). species identifications were verified against the grnp’s herbarium collection. species for which matches could not be found were sent for identification by taxonomists at the national (pretoria) or compton (cape town) herbarium of the south african national biodiversity institute. nomenclature follows the plant list (2013). abundance was recorded as 1, 2, 3 or 4 plants or estimated as the number of plants per location (5–10, 11–20, 21–50 or 51–100), and for subsequent calculations of plant abundances the median values of abundance class ranges (e.g. 15 for 11–20) were used. abundances of species occurring in extensive homogenous infestations were estimated for subsampled areas and extrapolated over the total extent of the infestation. in the case of clonal species, abundance values signify stem count estimates. age classes distinguished were young (i.e. immature), small-mature (i.e. sexually mature plants that were much smaller than the species’ potential size) and mature (i.e. sexually mature plants that were close to the species’ potential size). this differentiation allowed us to relate young or small-mature plants to their potential parent plants in order to calculate dispersal distances (see below). we noted if plants appeared to have been planted, such as inside fenced properties, in the form of rows or hedges, or visibly pruned inside gardens. locations were captured in a gis (arcmap 10.4) for further analyses. point locations were snapped to the road layer and then displaced to the correct side of the road in order to assign each location to an adjacent land cover type. land cover types (figure 1) were derived from the vegetation map of vlok et al. (2008) and the transformation map of vromans et al. (2010), both mapped at 1:50 000 scale during the mid-2000s. native vegetation in a pristine or near-pristine state (i.e. the ‘natural’ category of vromans et al. [2010]) was subdivided into fynbos and forest after the biome classification of vlok et al. (2008). non-natural land cover types included degraded (including the categories degraded, heavy alien degraded and alien transformed of vromans et al. [2010]), plantations (afforested with alien pinus or eucalyptus species) and farms (irrigated crop agriculture or pastures, and farm infrastructure), categorised as such by vromans et al. (2010). we calculated three measures of alien plant incidence: species richness (number of species), species density (number of species expressed per 10 km of road length) and plant density (number of plants expressed per 10 km of road length). we compared the incidence of alien plants in terms of these measures between (1) the southern cape and tsitsikamma regions, (2) inside and outside of the grnp and (3) land cover types. dispersal distances were estimated by means of distance analysis in gis for selected species – largely shrubs and trees that reproduce sexually. dispersal distances were measured from mature individuals to small-mature or young individuals of the same species between 100 m and 3 km from the mature individual. the distances to the 200 closest conspecific plants were calculated in microsoft excel, from the gis-derived data, for each mature plant location of the select species. dispersal distances were used to assess whether species complied with dispersal rates specified by pyšek et al. (2004) for a species to be classified as invasive (i.e. plants producing reproductive offspring by seed more than 100 m from parent plants in less than 50 years). our approach was aimed at deriving approximations of spread ability rather than at proving particular relationships between young and parent plants. to this end, the occurrence of small-mature or young plants (apparently not planted by humans) more than 100 m away from a potential parent signified spread unaided by humans. the study area stretched across 11 quarter degree squares (qdss). we compared our records of alien plant locations per qds with those of the march 2017 south african plant invader atlas (sapia) (henderson & wilson 2017). we also checked whether the alien plant species we recorded were listed as invasive species by nemba. in line with the definitions of sleeper weeds referred to earlier, we aimed to identify species that to date have been considered as benign (i.e. absent from sapia and nemba) but that were showing signs of spread. accordingly, we identified species as sleeper weeds if a species complied with criteria 1 and 3, or 2 and 3, of the following: (1) species has not been recorded by sapia in the study region; (2) species is not listed by nemba; (3) species displayed dispersal distances of > 100 m and is thus potentially invasive (after pyšek et al. 2004). results in this rapid assessment 109 exotic plant species were recorded across 1942 locations (table 1). thirty-five of these species were not nemba-listed and 35 species have not been recorded by sapia for the study area. importantly, absence of a species from our data does not prove absence of the species from the area, as inconspicuous species and individuals may be overlooked during a rapid survey. the most abundant species by numbers were nephrolepis cordifolia (sword fern) estimated at 4871 plants, ammi visnaga (false queen anne’s lace, toothpick weed) 2766 plants and lantana camara (lantana) 1333 plants. the highest abundance class used (50–100 plants) likely underestimated numbers of certain species, such as n. cordifolia and l. camara, which may have occurred at abundances of up to 100–500 plants per location, although in the case of clonal plants, abundance values rather signified stem count estimates. the species that occurred at the largest number of locations (indicative of wide distributions) were acacia elata (pepper tree wattle) at 130 and c. selloana or c. jubata (pampas grass) at 102 locations. table 1: alien and extralimital plant species recorded in the roadside survey and associated information. the southern cape region had higher alien species richness than the tsitsikamma region, but given that the former had double the extent of roads, it had lower species density (table 2). the southern cape also exhibited higher alien plant density than tsitsikamma. prominent species that were only recorded in the southern cape included l. camara, whereas alnus sp., liquidambar styraciflua (american storax) and solanum chrysotrichum (giant devil’s fig) were exclusive to the (easternmost part of) tsitsikamma (table 1). some species occurred as a few large disjunct populations throughout the study area, such as ammi visnaga, which had five large populations, each 2.5 km to 12.0 km long. table 2: extent of roads and alien plant species richness, species density and plant density compared between regions, inside versus outside of the garden route national park and land cover types. alien species richness and plant density were lower inside the grnp than outside (table 2). species density (number of species per unit of road length) was however higher inside the park than outside because of the limited extent of roads inside the park. species recorded at the largest numbers of locations (i.e. widely distributed) inside the park included paraserianthes lophantha (stink bean), melia azedarach (syringa) and hakea salicifolia (willow hakea), whereas those that were most abundant were n. cordifolia, sesbania punicea (red sesbania) and l. camara. among the five land cover types, alien plants were over-represented adjacent to farms in terms of species richness, species density and plant abundance (table 2). alien species richness and plant density were considerably higher in altered land cover types (degraded, farm and plantation) than in natural vegetation (forest and fynbos). forest had almost double the alien species richness of fynbos, but lower species densities and plant densities. the most noticeable addition to sapia records for the region was the herb ammi visnaga, recorded at 74 locations across eight qdss (table 1). we recorded sphaeropteris cooperi (australian tree fern) in eight qdss, as opposed to sapia’s two. other additions to sapia records included alnus sp., s. chrysotrichum, banksia ericifolia (heath-leaved banksia) and cupressus arizonica (arizona cypress), the latter at eight locations across three qdss. ten extralimital species were recorded, the most abundant of which was tecoma capensis (cape honeysuckle) in eight qdss, dais cotinifolia (pompon tree) and setaria cf. megaphylla (bush buffalo grass) in three qdss, cyperus papyrus (papyrus), syzygium cordatum (water berry) and carissa macrocarpa (num-num) in two qdss, and erythrina cf. lysistemon (coral tree) and podocarpus henkelii (henkel’s yellowwood) in one qds (table 1). domestic fruit trees – prunus persica (peach), eriobotrya japonica (loquat), cydonia oblonga (quince), citrus limon (lemon) and pyrus communis (pear) – were observed as young plants that appeared to have recruited from seeds. prunus persica was recorded at 22 locations across seven qdss and e. japonica at four locations in one qds. we identified 35 species as sleeper weeds (i.e. invasive), of which five were extralimital species. ammi visnaga, quercus robur (english oak), t. capensis, corymbia ficifolia (red flowering gum), syzygium paniculatum (bush cherry), quercus palustris (pin oak) and yucca aloifolia (yucca) occurred at the highest densities and were most widely distributed (table 1). ten of these species occurred in more than half of the qdss comprising the study area. quercus robur occurred in all 11 qdss and appeared to have dispersed > 100 m in 31 instances. large numbers of variously aged young of quercus acutissima and q. palustris growing close to the planted parent trees were recorded at witelsbos in the easternmost part of tsitsikamma. such proliferation of quercus acutissima (sawtooth oak) and q. palustris was not evident in the southern cape. numerous y. aloifolia seedlings were furthermore observed in the witelsbos area, whereas in the southern cape the species only reproduced vegetatively. other sleeper weeds identified in this study that were not regarded as invasive by baard and kraaij (2014) included schinus terebinthifolius (brazilian pepper), furcraea foetida (mauritius hemp), opuntia monacantha (drooping prickly pear), prunus persica and quercus acutissima. our observations of occurrences of young plants, at distances of 100 m – 3000 m from mature or planted conspecific plants, furthermore confirmed the invasive status of m. azedarach (87 instances), a. elata (26), c. selloana or c. jubata (17), grevillea robusta (silky oak; 14) and acacia podalyriifolia (pearl acacia; 4). species that commonly occurred as planted individuals but showed no sign of spread or regeneration were also worth noting and included the nemba-listed species acacia baileyana (bailey’s wattle), morus alba (white mulberry), nerium oleander (oleander) and phytolacca dioica (ombú), as well as species not listed by nemba for this region, agave attenuata (lion’s tail), jacaranda mimosifolia (jacaranda), metrosideros excelsa (new zealand christmas tree) and syncarpia glomulifera (turpentine tree). discussion spatial distribution of alien plants our comparison of alien plant incidence among land cover types showed that roads associated with natural vegetation (largely coinciding with roads inside the grnp) had lower species richness and densities of alien plants than those associated with degraded or transformed land (largely coinciding with roads outside the grnp). intact natural vegetation, and in particular tall dense vegetation such as forest, is known to present barriers to invasion (duncan 2011). in addition, narrower verges pertaining to park roads and alien plant management routinely undertaken by the grnp (kraaij et al. 2017; van wilgen et al. 2016) likely contributed to this finding. reduced alien plant invasions along roads inside the park (relative to outside) further attest to some level of success achieved by protected area agencies with alien plant management. roads along farmland had the highest incidence of alien plants and species. farmland represented the most complete transformation among the land cover types and is associated with relatively high human occupation and regular disturbance, all of which facilitate alien plant invasion (kalwij et al. 2008; richardson 2001). ammi visnaga was most abundant along roads associated with dairy farming and in mowed road verges. plantations were associated with lesser infestations of alien plants than farmland and other degraded land, possibly because of lower human occupation, better management and awareness of weeds (plantations are mostly certified by the forestry stewardship council; scotcher 2010) and potential out-shading of weeds by tall plantation trees. although tsitsikamma had fewer species and lower densities of alien plants along roads than the southern cape, it had higher species density and a few species (e.g. alnus sp. and s. chrysotrichum) exclusive to the east of the region. some species furthermore displayed different reproductive behaviour (e.g. yucca aloifolia apparently reproducing from seed) to that observed in the west. among other factors, more mesic conditions may facilitate invasion (richardson et al. 2000) in the eastern tsitsikamma, and monitoring for new weeds needs to be done regularly. the southern cape on the other hand has an expansive road network, transformed landscape and high species richness and densities of alien plants. this region thus faces high propagule pressure and ongoing alien plant management challenges. species priorities we identified 35 species as sleeper weeds on account of displaying (assumed) dispersal distances distinctive of invasive plants and not having been recorded or listed by sapia or nemba previously. effective seed dispersal is notoriously hard to measure (lack & evans 2005), and although it was impossible to prove relationships between young plants and deemed parent plants, the presence of juveniles additionally suggested spread. our approach of calculating distances between mature and younger plants provided valuable approximations of spread abilities, which could be used to assess invasiveness of species. several sleeper weeds occurred widely distributed along roads in the study area. of these, ammi visnaga occurred most abundantly and its linear spread along roads suggests that its seeds may be swept along by wind drag of passing vehicles (cf. maslo 2016; von der lippe & kowarik 2007). although sapia has not recorded this species, a conspecific, ammi majus (queen anne’s lace, large bullwort), has been recorded for the study area by bromilow (2010) and is included in sapia in habitat other than fynbos or forest. ammi visnaga may easily have been overlooked. ruščić and nikolić (2011) reported that this species was initially not recognised in a survey of ammi species in croatia. further investigation revealed that this species has been cultivated and used for medicinal purposes for > 120 years in croatia. recently, naturalised or invasive populations of the species were observed in bosnia and herzegovina, in natural and disturbed habitat (maslo 2016). sphaeropteris cooperi is not listed by nemba and was found over a much larger area than indicated by sapia. this fern sometimes occurred in colonies but very often as single plants with no apparent parent plant nearby, suggesting long distance dispersal via spores. it also occurred in a range of habitat types, that is, away from roads in pristine closed forest (pers. obs.) and in open pastures. unless overlooked in previous surveys, s. cooperi and a. visnaga have spread most markedly compared to sapia records. quercus robur is not nemba-listed and has been considered a casual alien in the grnp (baard & kraaij 2014). however, despite its large, heavy fruits (acorns), we recorded 31 instances of dispersal exceeding 100 m in addition to reproduction in the immediate surrounds of parent trees and thus consider the species to be invasive. olrik, hauser and kjaer (2012) also noted long distance dispersal in this species. corymbia ficifolia also occurred widely and abundantly but is not nemba-listed and has not been indicated by sapia for the study area. although most of the c. ficifolia we observed occurred as planted avenues, groups or rows of trees (47 localities), the species was unlikely to have been planted at another 18 localities, and young plants close to mature or planted individuals were also observed. five sleeper weeds were extralimital species, of which t. capensis, c. macrocarpa and setaria cf. megaphylla appeared most abundant. the natural range of t. capensis is uitenhage to subtropical africa and that of c. macrocarpa from humansdorp to mozambique (manning & goldblatt 2012). tecoma capensis, prevalent in horticulture, was often found close to human settlements but also in disturbed lowland fynbos and scrub forest, possibly discarded as garden refuse. carissa macrocarpa is bird-dispersed and commonly planted in gardens and on road islands. we recorded it in abundance in coastal fynbos between sedgefield and wilderness, which corresponds with observations of this species’ tolerance for saline coastal soils in texas (mink, singhurst & holmes 2015). setaria megaphylla is indigenous to south-eastern africa (van oudtshoorn 2010) but is considered a weed in commercial pine plantations in south africa (rolando & little 2009) and naturalised or invasive in parts of the united states (riefner & boyd 2007). horticultural use of these extralimital species should thus be discouraged outside of their native ranges. wilson et al. (2014) elucidate the conundrum of extralimital introductions (e.g. practical problems for legislators) and its potential negative consequences, such as its competition with indigenous species through exploitation and habitat change, the introduction of pathogens and parasites, and possible hybridisation. our findings on dispersal distances furthermore confirmed the invasive status of bird-dispersed species such as m. azedarach (voigt, farwig & johnson 2011) and wind dispersed species such as g. robusta (smith 1998) and c. selloana or c. jubata (melcher, bouman & cleef 2004; saura-mas & lloret 2005). in addition to sleeper weeds, a few invasive species (indicated by sapia and nemba) that currently do not occur inside the grnp or in low numbers have large numbers of young outside the park (e.g. acer negundo – ash-leaved maple, a. elata, p. lophantha and m. azedarach). these species pose a threat of invasion into the park and should be prioritised for control. conclusion relatively low work effort yielded insights into the diversity of alien plants, their spatial distribution and invasive tendencies in the area of the grnp. the rapid roadside survey provided a baseline dataset of easily detectable alien species and reaffirmed the importance of monitoring to detect sleeper weeds and changes in the status of alien plant populations. our comprehensive dataset contributed significant new records and range extensions to sapia, while the identified sleeper weeds offered suggestions for species to be considered for listing under nemba. we furthermore provided information to be used by the grnp for invasive alien plant management planning and prioritisation of species and areas, as well as future monitoring. if these surveys are to be repeated, it is important that travel speed be replicated as the level of effort affects the precision of observations. acknowledgements we thank lesley henderson for supplying sapia data and the south african weather service climate databank for rainfall data. the south african national parks and nelson mandela university enabled this study. comments from s. freitag-ronaldson, g. durrheim and r. scholtz on earlier drafts and suggestions from anonymous reviewers resulted in improvements to the manuscript. competing interests the authors declare that they have no competing interests with regard to the writing of this article. authors’ contributions j.a.b. and t.k. conceptualised the project. j.a.b. was responsible for data collection, gis data processing and map, and data analyses. j.a.b. and t.k. wrote the manuscript. funding information the south african national parks provided operational cost for this study. references alpert, p., bone, e. & holzapfel, c., 2000, ‘invasiveness, invasibility and the role of environmental stress in the spread of non-native plants’, perspectives in plant ecology, evolution and systematics 3, 52–66. https://doi.org/10.1078/1433-8319-00004 alston, k.p. & richardson, d.m., 2006, ‘the roles of habitat features, disturbance, and distance from putative source populations in structuring alien plant invasions at the urban/wildland interface on the cape peninsula, south africa’, biological conservation 132, 183–198. https://doi.org/10.1016/j.biocon.2006.03.023 baard, j. & kraaij, t., 2014, ‘alien flora of the garden route national park, south africa’, south african journal of botany 94, 51–63. https://doi.org/10.1016/j.sajb.2014.05.010 blackburn, t.m., essl, f., evans, t., hulme, p.e. & jeschke, j.m., 2014, ‘a unified classification of alien species based on the magnitude of their environmental impacts’, plos biology 12(5), e1001850. https://doi.org/10.1371/journal.pbio.1001850 boy, g. & witt, a., 2013, invasive alien plants and their management in africa, unep/gef removing barriers to invasive plant management project, cabi africa. bromilow, c., 2010, problem plants and alien weeds of south africa, 3rd edn., briza publications cc, arcadia. brooks, m.l. & klinger, r.c., 2009, ‘practical considerations for early detection monitoring of plant invasions’, in inderjit (ed.), management of invasive weeds, pp. 9–33, springer, new york. christen, d. & matlack, g., 2006, ‘the role of roadsides in plant invasions: a demographic approach’, conservation biology 20, 385–391. https://doi.org/10.1111/j.1523-1739.2006.00315.x cunningham, d.c., woldendorp, g., burgess, m.b. & barry, s.c., 2003, prioritising sleeper weeds for eradication: selection of species based on potential impacts on agriculture and feasibility of eradication, bureau of resource sciences, canberra, australia. davies, k.w. & sheley, r.l., 2007, ‘a conceptual framework for preventing the spatial dispersal of invasive plants’, weed science 55, 178–184. https://doi.org/10.1614/ws-06-161 duncan, r.p., 2011, ‘propagule pressure’, in d. simberloff & m. rejmánek (eds.), encyclopedia of biological invasions, pp. 561–563, university of california, berkeley, ca. foxcroft, l.c., van wilgen, n.j., baard, j.a. & cole, n.s., 2017, ‘biological invasions in south african national parks’, bothalia 47(2), a2158. https://doi.org/10.4102/abc.v47i2.2158 groves, r., 1999, ‘sleeper weeds’, in a.c. bishop, m. boersma & c.d. barnes (eds.), proceedings of the twelfth australian weeds conference, pp. 632–636, tasmanian weed society, hobart. henderson, l., 2007, ‘invasive, naturalized and casual alien plants in southern africa: a summary based on the southern african plant invaders atlas (sapia)’, bothalia 37(2), 215–248. https://doi.org/10.4102/abc.v37i2.322 henderson, l. & musil, k.j., 1984, ‘exotic woody invaders of transvaal’, bothalia 15(1&2), 297–313. https://doi.org/10.4102/abc.v15i1/2.1128 henderson, l. & wilson, j.r.u., 2017, ‘changes in the composition and distribution of alien plants in south africa: an update from the southern african plant invaders atlas (sapia)’, bothalia 47(2), a2172. https://doi.org/10.4102/abc.v47i2.2172 illenberger, w.k., 1996, ‘the geomorphic evolution of the wilderness dune cordons, south africa’, quaternary international 33, 11–20. https://doi.org/10.1016/1040-6182(95)00099-2 jodoin, y., lavoie, c., villeneuve, p., theriault, m., beaulieu, j. & belzile, f., 2008, ‘highways as corridors and habitats for the invasive common reed phragmites australis in quebec, canada’, journal of applied ecology 45, 459–466. https://doi.org/10.1111/j.1365-2664.2007.01362.x kalwij, j.m., milton, s.j. & mcgeoch, m.a., 2008, ‘road verges as invasion corridors? a spatial hierarchical test in an arid ecosystem’, landscape ecology 23, 439–451. https://doi.org/10.1007/s10980-008-9201-3 kolar, c.s. & lodge, d.m., 2001, ‘progress in invasion biology: predicting invaders’, trends in ecology & evolution 16, 199–204. https://doi.org/10.1016/s0169-5347(01)02101-2 kraaij, t., baard, j.a., rikhotso, d.r., cole, n.s. & van wilgen, b.w., 2017, ‘assessing the effectiveness of invasive alien plant management in a large fynbos protected area’, bothalia 47(2), a2105. https://doi.org/10.4102/abc.v47i2.2105 kraaij, t., cowling, r.m. & van wilgen, b.w., 2011, ‘past approaches and future challenges to the management of fire and invasive alien plants in the new garden route national park’, south african journal of science 107(9/10), art. #633. https://doi.org/10.4102/sajs.v107i9/10.633 lack, a. & evans, d., 2005, instant notes: plant biology, 2nd edn., taylor and francis, oxford. lloret, f., médail, f., brundu, g. & hulme, p.e., 2004, ‘local and regional abundance of exotic plant species on mediterranean islands: are species traits important?’, global ecology and biogeography 13, 37–45. https://doi.org/10.1111/j.1466-882x.2004.00064.x manning, j. & goldblatt, p., 2012, plants of the greater cape floristic region 1: the core cape flora, strelitzia 29, south african national biodiversity institute, pretoria. maslo, s., 2016, ‘contribution to the flora of bosnia & herzegovina (new neophytes in the flora of bosnia and herzegovina)’, herald of the national museum of bosnia and herzegovina ns 36, 43–61. maxwell, s.l., fuller, r.a., brooks, t.m. & watson, j.e.m., 2016, ‘the ravages of guns, nets and bulldozers’, nature 536, 143–145. https://doi.org/10.1038/536143a melcher, i.m., bouman, f. & cleef, a.m., 2004, ‘seed atlas of the monocotyledonous genera of the páramo’, flora 199, 286–308. https://doi.org/10.1078/0367-2530-00157 millar, c.i., stephenson, n.l. & stephens, s.l., 2007, ‘climate change and forests of the future: managing in the face of uncertainty’, ecological applications 17, 2145–2151. https://doi.org/10.1890/06-1715.1 mink, j.n., singhurst, j.r. & holmes, w.c., 2015, ‘jasminum laurifolium (oleaceae) adventive in texas, with observations on alien plant invasions and distribution on the texas gulf coast by passerines’, phytoneuron 36, 1–5. mortensen, d.a., rauschert, e.s.j., nord, a.n. & jones, b.p., 2009, ‘forest roads facilitate the spread of invasive plants’, invasive plant science and management 2, 191–199. https://doi.org/10.1614/ipsm-08-125.1 olrik, d.c., hauser, t.p. & kjaer, e.d., 2012, ‘natural colonisation of an open area by quercus robur l.–from where did the vectors disperse the seed?’, scandinavian journal of forest research 27, 350–360. https://doi.org/10.1080/02827581.2011.644318 pickering, c. & mount, a., 2010, ‘do tourists disperse weed seed? a global review of unintentional human-mediated terrestrial seed dispersal on clothing, vehicles and horses’, journal of sustainable tourism 18(2), 239–256. https://doi.org/10.1080/09669580903406613 pyšek, p., richardson, d.m., rejmánek, m., webster, g.l., williamson, m. & kirschner, j., 2004, ‘alien plants in checklists and floras: towards better communication between taxonomists and ecologists’, taxon 53(1), 131–143. https://doi.org/10.2307/4135498 rejmánek, m. & pitcairn, m.j., 2002, ‘when is eradication of exotic pest plants a realistic goal?’, in c.r. veitch & m.n. clout (eds.), turning the tide: the eradication of invasive species, pp. 249–253, iucn ssc invasive species specialist group, gland. rejmánek, m. & richardson, d.m., 1996, ‘what attributes make some plant species more invasive?’, ecology 77, 1655–1661. richardson, d.m., 2001, ‘plant invasions’, in levin, s.a. (ed.), encyclopedia of biodiversity, vol. 4, pp. 677–688, academic press, san diego, ca. richardson, d.m., bond, w.j., dean, w.r.j., higgins, s.i., midgley, g.f., milton, s.j. et al., 2000, ‘invasive alien organisms and global change: a south african perspective’, in h.a. mooney & r.j. hobbs (eds.), invasive species in a changing world, pp. 303–349, island press, washington, dc. riefner, r.e., jr. & boyd, s., 2007, ‘new records of wetland and riparian plants in southern california, with recommendations and additions to the national list of plant species that occur in wetlands’, journal of the botanical research institute of texas 1, 709–730. rolando, c.a. & little, k.m., 2009, ‘regional vegetation management standards for commercial pine plantations in south africa’, southern forests 71(3), 187–199. https://doi.org/10.2989/sf.2009.71.3.3.915 ruščić, m. & nikolić, t., 2011, ‘ammi visnaga (l.) lam. (apiaceae), a new taxon in croatian flora’, acta botanica croatica 70(2), 301–306. https://doi.org/10.2478/v10184-010-0023-4 sa explorer version 3.0, 2004, cd-rom, municipal demarcation board, pretoria, south africa. sanparks, 2010, the management plan of the garden route national park, unpublished document, south african national parks, knysna, south africa. saura-mas, s. & lloret, f., 2005, ‘wind effects on dispersal patterns of the invasive alien cortaderia selloana in mediterranean wetlands’, acta oecologica 27, 129–133. https://doi.org/10.1016/j.actao.2004.12.001 scotcher, j.s.b., 2010, ‘guest editorial. forest certification in south africa – the next phase’, southern forests: a journal of forest science 72(1), 1–4. https://doi.org/10.2989/20702620.2010.484899 shackleton, r.t., witt, a.b.r., nunda, w. & richardson, d.m., 2017, ‘chromolaena odorata (siam weed) in eastern africa: distribution and socio-ecological impacts’, biological invasions 19, 1285–1298. https://doi.org/10.1007/s10530-016-1338-4 smith, c.w., 1998, ‘hawaiian alien plant studies’. department of botany, university of hawaii, usa, viewed 20 august 2018, from http://www.botany.hawaii.edu/faculty/cw_smith/aliens.htm. tham, a.g. & johnson, m.r., 2006. ‘the cape supergroup’, in m.r. johnson, c.r. anhaeusser & r.j. thomas (eds.), the geology of south africa, pp. 443–457, council of geoscience, pretoria. the plant list, 2013, version 1.1, published on the internet, viewed 06 june 2018, from http://www.theplantlist.org/. thuiller, w., richardson, d.m., pyšek, p., midgley, g.f., hughes, g. & rouget, m., 2005, ‘niche-based modelling as a tool for predicting the risk of alien plant invasions at a global scale’, global change biology 11, 2234–2250. https://doi.org/10.1111/j.1365-2486.2005.001018.x tyson, p.d. & preston-whyte, r.a., 2000, the weather and climate of southern africa, oxford university press, cape town. van oudtshoorn, f., 2010, guide to grasses of southern africa, briza publications, pretoria, south africa. van wilgen, b.w., fill, j.m., baard, j.a., cheney, c., forsyth, a.t. & kraaij, t., 2016, ‘historical costs and projected future scenarios for the management of invasive alien plants in protected areas in the cape floristic region’, biological conservation 200, 168–177. https://doi.org/10.1016/j.biocon.2016.06.008 vitousek, p.m., d’antonio, c.m. & asner, g.p., 2010, ‘invasions and ecosystems: vulnerabilities and the contribution of new technologies’, in d.m. richardson (ed.), fifty years of invasion ecology: the legacy of charles elton, pp. 227–288, wiley-blackwell, sussex. vlok, j.h.j., euston-brown, d.i.w. & wolf, t., 2008, a vegetation map for the garden route initiative, unpublished 1:50 000 maps and report supported by cape fsp task team. voigt, f.a., farwig, n. & johnson, s.d., 2011, ‘interactions between the invasive tree melia azedarach (meliaceae) and native frugivores in south africa’, journal of tropical ecology 27, 355–363. https://doi.org/10.1017/s0266467410000702 von der lippe, m. & kowarik, i., 2007, ‘long-distance dispersal of plants by vehicles as a driver of plant invasions’, conservation biology 21(4), 986–996. https://doi.org/10.1111/j.1523-1739.2007.00722.x vromans, d.c., maree, k.s., holness, s., job, n. & brown, a.e., 2010, the garden route biodiversity sector plan for the george, knysna and bitou municipalities: supporting land-use planning and decision making in critical biodiversity areas and ecological support areas for sustainable development, garden route initiative and south african national parks, knysna, south africa. williams, j.a. & west, c.j., 2000, ‘environmental weeds in australia and new zealand: issues and approaches to management’, austral ecology 25, 425–444. https://doi.org/10.1046/j.1442-9993.2000.01081.x wilson, j.r., gaertner, m., griffiths, c.l., kotzé, i., le maitre, d.c. & marr, s.m., 2014, ‘biological invasions in the cape floristic region: history, current patterns, impacts, and management challenges’, in n. allsopp, j.f. colville & g.a. verboom (eds.), fynbos: ecology, evolution, and conservation of a megadiverse region, pp. 273–298, oxford university press, oxford. wilson, j.r.u., gaertner, m., richardson, d.m. & van wilgen, b.w., 2017, ‘contributions to the national status report on biological invasions in south africa’, bothalia 47(2), a2207. https://doi.org/10.4102/abc.v47i2.2207 wilson, j.r., panetta, d.f. & lindgren, c., 2017, detecting and responding to alien plant incursions, cambridge university press, cambridge, uk. woodward, s.l. & quinn, j.a., 2011, encyclopedia of invasive species: from africanized honey bees to zebra mussels, abc-clio, santa barbara, ca. filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 filelist convert a pdf file! page 1 page 2 hipondoka.qxd monitoring the vegetation structure of south-central etosha national park using terrestrial photographs m.h.t. hipondoka and w.d. versfeld hipondoka, m.h.t. and w.d. versfeld. 2003. monitoring the vegetation structure of south-central etosha national park using terrestrial photographs. koedoe 46(1): 17–27. pretoria. issn 0075-6458. in 1984, a ground-based, photographic library was initiated for monitoring the vegetation dynamics in etosha national park. over 400 photographic points were selected. at these points, panchromatic photographs were taken at intervals of six years. this study was undertaken to assess the applicability of this terrestrial photographic library in characterising the dynamics of the vegetation structure in south-central etosha national park. the methods employed include field validation, visual photograph interpretations, spatial analysis, and aggregated two-dimensional tables. results, though patchy in nature, show that the vegetation structure at 63 % of the sites covered remained either unchanged or increased over the 15-year study period. the patchiness of the photo points can be exploited and be treated as training sites for corresponding satellite image classifications to provide continuous ground coverage results. key words: etosha, photographs, savanna, vegetation. m.h.t. hipondoka and w. d. versfeld , etosha ecological institute, po box 6, okaukuejo via outjo, namibia (mhipondoka@yahoo.com / eei.staff@mweb.com.na). issn 0075-6458 17 koedoe 46/1 (2003) introduction management of a national park of etosha’s size (22 270 km²) often presents challenges. constant and up-to-date scientific feedback is essential to enable timely and adaptive management practices. before the relatively recent widespread application of satellite remote sensing technologies that can be used to survey and monitor the environment, etosha national park (etosha np), due to its size, was confronted with the challenge of adopting a reliable and consistent method of monitoring its vegetation dynamics. consequently, a terrestrial photo-point system was initiated in 1984 (etosha ecological institute 1984). the terrestrial photo-point system forms a visual record of short, medium and long-term changes taking place in the vegetation properties (joubert 1983). graetz & pech (1987), however, heavily criticised this method when they tested it for land cover changes in remote arid lands of australia. they concluded that it did not afford any possibility to accurately estimate the dynamics of land cover. yet, these authors acknowledged its potential for collecting records of re-vegetation at established permanent sites. generally, in spatial analyses vertical aerial photographs are preferred to oblique photographs, particularly, in terms of geometric fidelity (lillesand & kiefer 1994; campbell 1996). oblique photographs, however, offer the advantage of a more natural view-angle, and the ability to view objects that occur under the canopy (campbell 1996; fahsi 2001). of equal importance, is the fact that because oblique aerial photos are not used for photogrammetric and precision purposes, given their poor geometric fidelity, they may use inexpensive cameras. the terrestrial photo point system was therefore deemed as a cost effective and easily applicable method that would meet the demand for vegetation data collection of etosha np. this paper presents the results of applying and assessing this method as a ground-based system for hipondoka.qxd 2005/12/09 10:27 page 17 monitoring the vegetation dynamics of etosha national park. methods in this paper, the term ‘photo point’ is used to describe specific plots where photographs were taken. at each photo point, sets of four photographs were usually taken to capture the northern, southern, eastern and western view of the plot. the term ‘site’ is used to refer to these four views of the photo point. analysis was carried out at the site level, as the vegetation structure of the same plot was not necessarily always homogeneous. study area etosha national park is located in northern namibia and comprises a surface area of 22 270 km² (lindaque & archibald 1991). etosha pan, a saline depression covering a total of 4500 km² or 20 % of the park, is a prominent feature of the area. fourteen (14) vegetation structure classes are recognised in etosha np, ranging from high tree savanna to grassland (sannier et al. in prep.). the park receives summer rainfall and decreases along an east-west gradient, the east being wetter (engert 1997). figure 1 presents the annual rainfall received between 1980 and 1999 at okaukuejo and ombika, both located within the study area. although the two stations are only 17 km apart, comparatively, ombika receives higher rainfall than okaukuejo. the long-term average annual rainfall at okaukuejo and ombika is 320 mm and 382 mm, respectively. essentially, the study area is characterised by erratic rainfall, both in space and time (engert 1997). as a test phase, this study was limited to the southcentral part of etosha np (fig. 2), covering a surface area of 1936 km². all vegetation structure classes, but ‘high tree savanna high’, occur in the study area. the area extends from the park’s southern boundary to adamax pan in the north, and from kapupuhedi waterhole in the east to grunewald waterhole in the west. the study area is covered by a total of 60 photo points, generating approximately 230 photographs each survey year. data acquisition due to the risk of being unable to re-identify the photo points in subsequent visits, and the absence of koedoe 46/1 (2003) 18 issn 0075-6458 fig. 1. annual rainfall received at okaukuejo and ombika between 1980 and 1999. hipondoka.qxd 2005/12/09 10:27 page 18 issn 0075-6458 19 koedoe 46/1 (2003) fig. 2. location of the study area and photo points in etosha national park hipondoka.qxd 2005/12/09 10:27 page 19 data processing and analysis before the analysis and interpretation of the photographs commenced, field verification was conducted in 1999, during the dry season. essentially, this exercise entailed familiarization of the interpreter with the sites to be interpreted (estes 2001). the sites were located with the help of a gps (garmin 45xl) and the photographs of 1996. at each site, a classification of the vegetation structure was made, based on bougey’s (1957, cited by sannier et al. 1995) classification scheme, which was adapted to the semi-arid conditions of etosha by sannier et al. (1995). two vegetation attributes, height and crown cover, were used to determine the vegetation structure of each site. the decision tree used for the structural vegetation description was adapted from du plessis (1999), in which the model and details are provided. briefly, this method defines the vegetation structures based upon combinations of plant height and canopy cover. examples of height descriptions are: high tree savanna, low tree savanna, shrub savanna, grass savanna, steppe and grassland; whilst the canopy cover was categorised in qualitative terms, e.g. low, medium and high. thus, the vegetation type ‘steppe’ could be, for example, subdivided into ‘steppe very low’, ‘steppe low’ and ‘steppe medium’. exceptions in the classes used, were for sites that were devoid of vegetation, due to human activities, such as gravel pits, which were classified accordingly. the results of field verification were summarised as the structural vegetation classification of 1999. global positioning system (gps) technologies at the time, photographic points were out of necessity subjectively selected. monitoring sites were therefore located near places or structures such as roads or track junctions, waterholes, gravel pits, road-signs, horse camps, along pan margins and the park’s boundary fence (etosha ecological institute 1984). however, these sites were not necessarily representative of the rangeland, given their proximity to both anthropogenic impact and higher game concentrations. milton et al. (1998), for example, suggest that rangeland monitoring sites should be located at least 20 m from a track or fence and 100 m from watering points. the goal, however, was to detect change in the vegetation and in taking into consideration the fact that both man and beast are the primary landusers of the park, the justification of establishing the photo points at these localities was warranted. photographs were first taken in 1984 and repeated in 1990 and 1996. a total of 442 photo points are currently in use, including a fraction of photo points that could not be re-identified in successive visits and new ones that were subsequently created to replace them in the same localities. the photo points are spread across the park and cover all the vegetation types. the major pans, such as etosha and adamax, were however not covered. a 35 mm single lens reflex fm nikon camera body with a standard 50 mm lens and a panchromatic 100asa film was used consistently. at each site, photographs were taken from a vehicle’s rooftop, a windmill or road sign-block platform. photographs were taken towards each of the four cardinal compass bearings (i.e. north, east, south and west) to ensure a panoramic, representative and holistic view of the entire plot. consistency was maintained for capturing 7/8th of the subject matter covering the land surface, and the remainder (1/8th) contained the horizon. photographs were taken during the dry, winter months of july and august, when tree leaves are off. although photographs taken when trees have their leaves out have an advantage of tree species identification (lathrop 1999), the timing of these photographs was primarily dictated by accessibility to all parts of the park (not possible during the rainy season) and to coincide with the season that the national aerial photograph series are taken. an oversight, however, was that successive photographs of the same plot were not taken at the same time of day, given that the routes taken and time of exposure were not recorded. photographs were processed locally, using standard processing procedures. koedoe 46/1 (2003) 20 issn 0075-6458 fig. 3. geo-coding of sites as derived from the original coordinates of the photo point. hipondoka.qxd 2005/12/09 10:27 page 20 issn 0075-6458 21 koedoe 46/1 (2003) subsequently, each photograph taken in each of the three study year’s, 1984, 1990 and 1996, was interpreted using the above classification. to maximise consistency, the same author carried out all visual photo interpretations. the other author crosschecked a 30 % sample, randomly selected from the interpretation results. this crosschecking resulted in a high degree (91 %) of agreement. the interpretation results were recorded in a spreadsheet. to facilitate spatial analysis of the interpretation results under a geographic information system (gis), sites were geo-coded using universal transverse mercator (utm) coordinates. the utm grid system was favoured for its measuring units (meters), which simplified the process of geo-coding sites. moreover, a map grid is more appropriate than geographical coordinates for large-scale maps (sannier et al. 2002). to avoid sites of the same plot overlapping in a map format, the appropriate utm coordinate values of each photo point were offset by a 30 m distance towards the respective direction of each plot view. for example, the easting and northing of photo point 100, presented in fig. 3, is 578550 and 7882850, respectively. the coordinates of site 100 n (north) were obtained by adding 30 to 7882850 (northing value), while the easting figure remains unchanged. the coordinates for the centre point of site 100 n would therefore read 578550 and 7882880 for the easting and northing, respectively. the same technique was then used to derive the coordinates for the sites in the south, east and west. integrated land and water information system (ilwis) software (itc, enschede, the netherlands) was used for spatial analysis. the interpretation results were imported into ilwis in a table format and subsequently converted to point maps, in both vector and raster format. raster maps were resampled to a 30 m pixel size. data query, retrieval and display were done in both map and table formats. these included identifying sites where the vegetation structure remained consistently unchanged during the study period, using the equation: mapout:=iff(((map84=map90)and(map84=map96) and(map84=map99)and(map90=map96)and(map90 =map99)and(map96=map99)),map84,"?") ... (equation 1) where mapout is the name of the output map, map84, map90 and map96 is the vegetation structure classes of 1984, 1990 and 1996, in that order, and map99 is the field validation results. fig. 4. distributions and fluctuation patterns of the vegetation structure during the survey years. (note: photo points that could not be re-identified in subsequent visits and new ones created to replace them are grouped under "no photo" in the figure.) hipondoka.qxd 2005/12/09 10:27 page 21 in order to capture the conversion of the vegetation structure classes between any two study years, the “map crossing” function was applied. the results, in table format, were transformed into matrixes. results figure 4 depicts the recorded distributions and fluctuation patterns of the vegetation structure for all survey years. during the study years, ‘steppe low’ was the dominant vegetation structure, followed by ‘shrub savanna low’. few sites were classified as ‘bare ground’, while a fraction of others, were characterised by man-made features, such as gravel pits, tracks and roads. fluctuations among survey years were largest for ‘low tree savanna low’ (coefficient of variation (cv) =59.6 %), ‘steppe low’ (cv=48.69 %) and ‘steppe medium’ (cv=43.5 %) and lowest for ‘bare ground’ (cv=0.2 %), low tree savanna medium’ (cv=0.2 %), ‘high tree savanna low’ (cv=0.8) and ‘high tree savanna medium’ (cv=0.8). using the vegetation structural classes of 1984 as the baseline condition, variations in vegetation structural classes between 1984 and subsequent survey years were obtained (fig. 5). in 1990, the vegetation structure types at 64 % of the sites remained unchanged, whereas 23 % declined and 13 % increased from the previous classification of 1984. the number of sites, where the vegetation structure remained unchanged when compared with the baseline data, decreased by 10 % in 1996, to 54 %. in contrast, the number of sites with increased and declined vegetation structure raised to 17 % and 29 %, respectively, in 1996. the variations between the field results and the baseline show that sites with the same classification and those that declined in their vegetation structure between the two dates, tied at 37.3 %, whilst 25.4 % of the sites experienced growth. over the entire study period, the number of sites that remained unchanged declined steadily, whilst a steady increase occurred in sites with a decline and incline of the vegetation structure. koedoe 46/1 (2003) 22 issn 0075-6458 fig. 5. overall changes of the vegetation structure compared with the baseline condition. the preceded changes identified after 1990, do not make a distinction between sites with change in the vegetation structure between one or two survey years, but returned to the baseline condition in later survey years and those that remained consistently unchanged and progressively declined or increased. for example, site 200-n, classified as ‘steppe very low’ in 1984, was categorised as ‘steppe low’ in 1990 and in 1996 its vegetation was recorded to have returned to the baseline condition. it was thus accounted for having its vegetation structure declined in fig. 6. trend of sites with consistently unchanged, and progressively declined or increased vegetation structural classes. hipondoka.qxd 2005/12/09 10:27 page 22 1 1 4 4 issn 0075-6458 23 koedoe 46/1 (2003) table 1 conversion matrix of the vegetation structural classes at individual sites between 1984 and 1999 no photo road/gravel pit, etc. bare ground grassland steppe very low steppe low steppe medium grass savanna shrub savanna low shrub savanna medium low tree savanna low low tree savanna medium high tree savanna low high tree savanna medium total (1999) 0 2 1 1 3 7 4 9 16 3 32 3 11 2 6 22 2 2 1 10 2 17 1 3 4 2 2 23 5 4 6 11 40 31 21 41 11 34 4 3 0 234 r oa d, g ra ve l p it, e tc . b ar e g ro un d g ra ss la nd st ep pe v l st ep pe l st ep pe m g ra ss s av an na sh ru b sa va nn a l sh ru b sa va nn a m l ow tr ee s av an na l l ow tr ee s av an na m h ig h tr ee s av an na l h ig h tr ee s av an na m to ta l ( 19 84 ) 1999 1984 = number of sites with vegetation structure unchanged = number of sites with vegetation structure increased = number of sites with vegetation structure declined = new sites after 1984. 4 4 4 4 4 13 6 1 1 25 10 3 5 18 9 2 1 2 50 6 1 5 8 1 4 1 26 1 4 1 7 17 5 8 1 44 1990 and remained unchanged under the 1996 classification. tracking of these dynamics show a distinct trend in the structural vegetation classes of sites that consistently remained unchanged, progressively declined or increased over the entire study period. this trend is captured in fig. 6, where a negative linear relationship with time appears to emerge, although more data is needed for that purpose. by 1999, only one site, located some 2.5 km north of ombika waterhole, where the vegetation structure increased progressively. in contrast, only two sites experienced progressive decline in their vegetation structure during the same period. both sites are located hipondoka.qxd 2005/12/09 10:27 page 23 within one kilometre radius from either leeubron waterhole (closed since 1974) or okaukuejo waterhole. moreover, the vegetation structure at 47 or 20 % of the sites remained consistently unchanged over the entire study period. these sites are scattered in all vegetation structure types and at varying distances from areas of high human impact and/or animal concentrations. for example, the status of three sites (two sites with ‘steppe low’ and one with ‘grassland’ classification) located within 400 m radius from okondeka waterhole, remained unchanged since 1984. conversion of the vegetation structure between the survey years is another aspect that was investigated. table 1 presents the matrix of the vegetation structural class conversion between 1984 and 1999. of particular interest in this analysis are 25 % of sites where the vegetation structure increased. among these sites, five of them, originally influenced by human activities i.e. gravel pits, gate or track, in 1984, were classified as either ‘shrub savanna low’ or ‘low tree savanna medium’ in 1999. acacia newbrownii, a renowned natural invasive tree species, dominates one of the sites located in the periphery of a gravel pit, while another site is located along the edge of a tar road. two other sites, where the vegetation structure increased, are located within 400 m radius from waterholes (leeubron or gemsbokvlakte). one of these sites shifted from ‘grassland’ to ‘shrub savanna low’, and the other from ‘steppe very low’ to ‘grass savanna’. at the other end of the spectrum, there are four sites where the vegetation structure declined from either ‘shrub savanna low’ (one site) or ‘low tree savanna low’ (three sites) in 1984 to ‘grassland’ in 1999. these sites are located some tens of meters from waterholes (okaukuejo, ombika or leeubron). in addition, the structural class at five sites declined to ‘steppe low’ from ‘grass savanna’ (two sites), ‘shrub savanna low’ (one site) or ‘shrub savanna medium’ (two sites). of all the sites with declined status in 1999, 59.5 % of them were within one class margin relative to their original baseline conditions. discussions the main objective of this study was to evaluate the potential of terrestrial photographs as an approach for monitoring the vegetation structure through time. our results obtained from the photo interpretations indicate that they are comparable with the data collected from field validations. although noticeable differences, such as a low number of sites classified as ‘low tree savanna low’ in 1999 in relation to the other three years, occurred between the two methods, we do not consider such differences as a consequence of wrongly interpreting the vegetation structure from either the photographs or field validations. rather, as it will be discussed later, these differences are more likely to be related to the response of the vegetation to the accumulated effect of natural factors, such as rainfall and herbivores. with a relatively good agreement obtained between field validations and terrestrial photo interpretations, a photo point system of etosha np shows a promising role to play in supporting the use of satellite remote sensing data, which, as trodd & dougill (1998) noted, are currently not successful in estimating the vegetation structure in a semiarid environment. this is primarily attributed to the limited dimensionality of reflectance data that causes a weak relationship between spectral reflectance and vegetation structure (trodd & dougill 1998; dramstad et al. 2002). moreover, the savanna ecosystems, characterised by spatial heterogeneity introduced through the co-existence of herbaceous and woody plants (scholes & walker 1993), increase this challenge for the use of satellite data, such as landsat tm, in assessing the dynamics of vegetation structure (hudak & wessman 1998). in spite of these limitations, satellite data have the advantages of high temporal resolution and large area coverage. therefore, in conjunction with satellite data, particularly those with high spatial resolution, a koedoe 46/1 (2003) 24 issn 0075-6458 hipondoka.qxd 2005/12/09 10:27 page 24 photo-point system has the potential of serving as a relevant long-term ground-based monitoring method for the vegetation dynamics of a semi-arid environment. however, the selection of monitoring sites in etosha np requires careful consideration, given the fact that the photo-points were subjectively selected and also that they overrepresent areas of high impact. analysis of vegetation structure the results of this study reveal that the vegetation structure of south-central etosha np is continuously changing with time. over the 15-year study period, only a fifth of the sites did not experience change in their vegetation structure. at the same time, the changes experienced at the rest of the sites are not particularly dramatic. there is no obvious trend betraying any preference of change towards or away from any particular structural vegetation class or a group of classes. this condition confirms the notion that stability is not a major feature of savanna ecosystems, while at the same time changes of great magnitude do not occur (walker & noy-meir 1982; kempf & busche 2002). the savanna ecosystems are noted for being constantly changing in their vegetation properties in response to anthropogenic activities and the ecological “driving forces” of rainfall, fire regime and grazing level (lamprey 1979; walker & noy-meir 1982; joubert 1983; scholes & walker 1993; moleele et al. 2001). although human activity is taking place in the study area, a number of sites that were characterized by anthropogenic impact, such as gravel pits, was small in 1984 and has subsequently decreased in successive study years. this suggests that direct human interference in the park is minimal. the role of herbivores in the study area is difficult to assess, due to a lack of appropriate historical data on game distributions, density and range utilisations. however, elephants were often observed by the authors causing significant damage to the vegetation, particularly at and around waterholes, such as okaukuejo. rainfall in the study area between 1980 and 1999 was generally below the long-term average annual rainfall, with only six out of 20 years the precipitation exceeded the longterm annual average. however, the rainfall variability of this area is illustrated by the fact that years with rainfall above or below the long-term annual average, do not necessarily coincide between the two stations, okaukuejo and ombika, both located within the study area. this has practical implications in relating changes in the vegetation structure to rainfall. this study has revealed that, the changes recorded in the vegetation structure have little resemblance to the rainfall pattern. for example, a low number of sites classified as ‘low tree savanna low’ in 1999 (fig. 4), relative to the results of photo interpretations, can be attributed to a very low rainfall received in the two consecutive years, 1998 and 1999 (fig. 1). the higher number of sites classified as ‘shrub savanna medium’ in 1999, could therefore be accounted for being converted down from the neighbouring class ‘low tree savanna low’. this assumption, however, does not apply to fluctuations that occurred in classes ranging from ‘steppe very low’ to ‘shrub savanna low’, which all lack a predictable signature. it is therefore probable that the observed lack of relationship between fluctuations in the vegetation structure between years and rain pattern is a direct consequence of rainfall variability, both in space and time, within the study area. this theory of rainfall variability and its bearing on the vegetation trend needs empirical testing. this will be done using data from a dense network of field rain gauges initiated in etosha np in 1984 (du plessis 1997; engert 1997), and will be dealt with elsewhere. the occurrence of fire in the study area during the period under review was infrequent. the last recorded fire occurred in 1976 in the northern part of the study area and in 1990 (after photographs were already taken) and 1991, in the southern part. none of these fire effects were apparent in the results. in a study aimed at detecting land cover changes in a forested area using tm imagery, rogan et al. (2002) observed that although there issn 0075-6458 25 koedoe 46/1 (2003) hipondoka.qxd 2005/12/09 10:27 page 25 was a relationship between fire scars and the decline of the vegetation, not all wildfires were detected using a five-year temporal resolution images. it is therefore likely that a temporal resolution of this scale contains an intrinsic gap in change detection studies, particularly in areas that are subjected to fire. conclusions the photo point system provided results comparable to field validations. with such a capability, it has the potential to play a supportive role, alongside satellite data, as a parallel, ground-based monitoring system for monitoring the vegetation structure. acknowledgements we thank w. du plessis for his encouragement, insights and field calibration of the interpretation method. n. knox, dr v. myers and j. lates made invaluable remarks on the first draft. we also thank two anonymous reviewers, who made useful comments, which in turn help improved the manuscript. the namibian ministry of environment and tourism supported all phases of the project in every form. references campbell, j. b. 1996. introduction to remote sensing. new york: guilford press. dramstad, w.e., w.j. fjellstad, g.h. strand, h.f. mathiesen, g. engan & j.n. stokland. 2002. development and implementation of the norwegian monitoring programme for agricultural landscapes. journal of environmental management 64: 49–63. du plessis, w.p. 1997. refinements to the burning strategy in the etosha national park, namibia. koedoe 40: 63–76. du plessis, w.p. 1999. linear regression relationships between ndvi, vegetation and rainfall in etosha national park, namibia. journal of arid environment 42: 235–260. etosha ecological institute. 1984. motivation for photo point and rain gauge layout system in etosha national park. okaukuejo. unpublished internal report (n 18/2/1/30). engert, s. 1997. spatial variability and temporal periodicity of rainfall in the etosha national park and surrounding areas in northern namibia. madoqua 20 (1): 115–120. estes, j.e. 2001. elements, aids, techniques and methods of photographic/image interpretation. [online]. in: estes, j.e., k.c mcgwire & k.d. kline (eds.). air photo intepretation and photogrammetry. volume 1. available: “http:// umbc7.umbc.edu/~tbenja1/santabar/vol1/lec2/2l ecture.html” http://umbc7.umbc.edu/~tbenja1/ santabar/vol1/lec2/2lecture.html [accessed 2 july 2002]. fahsi, a. 2001. geometry of aerial photography. [online]. in: estes, j.e., k.c mcgwire & k.d. kline (eds.). air photo intepretation and photogrammetry. volume 1. available: http:// umbc7.umbc.edu/~tbenja1/santabar/vol1/lec6/ 6lecture.html [accessed 2 july 2002]. graetz, r.d. & r.p. pech. 1987. detecting and monitoring impacts of ecological importance in remote arid lands: a case study in southern simpson desert of south australia. journal of arid environments 12: 269–284. hudak, a.t. & c.a. wessman. 1998. textural analysis of historical aerial photography to characterize woody plant encroachment in south african savanna. remote sensing of environment 66: 317–330. joubert, s.c.j. 1983. a monitoring program for an extensive national park. pp. 201–213. in: owen-smith, r.n. (ed.). management of large mammals in african conservation areas. pretoria: haum. kempf, j. & d. busche. 2002. modelling environmental history in central namibia since the late tertiary. zeitschrift für geound umweltwissenschaften 146: 8–15. lamprey, h.f. 1979: structure and functioning of the semi-arid grazing land ecosystem of the serengeti region (tanzania). pp 562–601. in: unesco/unep/fao (eds.). tropical grazing land ecosystems. paris: unesco lathrop, r. 1999. acquisition of aerial photographs [online]. available: hyperlink “http://crssa. rutgers.edu/courses/airphoto/airphoto8/sld007. htm” (accessed 13 january 2003) lillesand, t.m & r.w. kiefer. 1994. remote sensing and image interpretation. new york: john wiley. lindaque, m. & t.j. archibald. 1991. seasonal wetlands in owambo and etosha national park. madoqua 17 (2): 129–133. milton, s.j., w.r.j. dean & r.p. ellis. 1998. rangeland health assessment: a practical guide for ranches in arid karoo shrublands. journal of arid environment 39: 253–265 moleele, n.m, s. ringrose, w. matheson & c. vanderpost. 2001. more woody plants? the status of bush encroachment in botswana’s grazing areas. journal of arid environment 64: 3–11. koedoe 46/1 (2003) 26 issn 0075-6458 hipondoka.qxd 2005/12/09 10:27 page 26 rogan, j., j. franklin & d.a. roberts. 2002. a comparison of methods for monitoring multitemporal vegetation change using thematic mapper imagery. remote sensing of environment 80:143–156. sannier, c.a.d, j.c. taylor, w.p. du plessis & k. campbell. 1995. application of remote sensing and gis for monitoring vegetation in etosha national park. paper presented at: remote sensing and gis for natural resource management rss workshop, 19 december 1995, chatham, uk. sannier, c.a.d, j.c. taylor & w. du plessis. 2002. real-time monitoring of vegetation biomass with noaaa-avhrr in etosha national park, namibia, for fire risk assessment. international journal of remote sensing 23(1): 71–89. scholes, r.j. & b.h. walker. 1993. an african savanna: synthesis of the nylsvley study. cambridge: cambridge university press. trodd, n.m. & a.j. dougill. 1998. monitoring vegetation dynamics in semi-arid african rangelands. applied geography 18 (4): 315–330. walker, b.h & i. noy-meir. 1982. aspects of the stability and resilience of savanna ecosystems. pp. 577–590. in: huntley, b.j, & b.h. walker, (eds.). ecology of tropical savannas. berlin: springer-verlag. issn 0075-6458 27 koedoe 46/1 (2003) hipondoka.qxd 2005/12/09 10:27 page 27 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <feff0041006e007600e4006e00640020006400650020006800e4007200200069006e0073007400e4006c006c006e0069006e006700610072006e00610020006e00e40072002000640075002000760069006c006c00200073006b0061007000610020005000440046002d0064006f006b0075006d0065006e00740020006d006500640020006800f6006700720065002000620069006c0064007500700070006c00f60073006e0069006e00670020006600f60072002000700072006500700072006500730073007500740073006b0072006900660074006500720020006100760020006800f600670020006b00760061006c0069007400650074002e0020005000440046002d0064006f006b0075006d0065006e00740065006e0020006b0061006e002000f600700070006e006100730020006d006500640020004100630072006f0062006100740020006f00630068002000520065006100640065007200200035002e003000200065006c006c00650072002000730065006e006100720065002e00200044006500730073006100200069006e0073007400e4006c006c006e0069006e0067006100720020006b007200e400760065007200200069006e006b006c00750064006500720069006e00670020006100760020007400650063006b0065006e0073006e006900740074002e> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 381---26 october 2009.indd [ final version].indd a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe original research a rticle #381 (page number not for citation purposes) 49 why travel motivation and socio-demographics matter in managing a national park abstract the addo elephant national park is one of only a few national parks in the world that offers the big 7 experience and is therefore one of south africa’s prime tourism destinations. the park plays an important role in the regional economy and has become a hub for tourism development. the aim of this article is to determine the extent to which socio-demographic and behavioural and motivational indicators infl uence the spending of tourists to the park. a better understanding of the latter could help marketers and planners to increase the economic impact of the park. since 2001, surveys have been conducted among tourists to the park and have included a number of socio-demographic, behavioural and motivational questions. in this analysis, 537 questionnaires were used. the methodology used includes factor analysis, cross-sectional regression analysis and pseudo-panel data analysis to determine and compare possible infl uences on spending. the research identifi es six motives for tourists travelling to the addo elephant national park; these are nature, activities, family and socialisation, escape, attractions and photography. the research found that a combination of socio-demographic and motivational factors infl uences visitor spending decisions. added to this, the research confi rms that tourist attractions, including national parks, differ from one another and that the variables that infl uence spending therefore also differ. conservation implications: in order for national parks to fulfi l their conservation mandate, they require funding. one of the main sources of income for national parks is tourist spending. this article identifi es the socio-demographic and motivational factors that infl uence tourist spending. hence, park management can use these results in order to market and create opportunities for tourists to spend more thereby benefi ting conservation directly. authors: melville saayman1 andrea saayman1 affi liations: 1institute for tourism, wildlife economics & leisure studies and school of economics, north-west university, south africa correspondence to: melville saayman e-mail: melville.saayman@nwu. ac.za postal address: private bag x6001, potchefstroom, 2520, south africa keywords: conservation and tourism; sociodemographics; travel motivation; travel behaviour; addo elephant national park dates: received: 18 sep. 2008 accepted: 07 may 2009 published: 25 june 2009 how to cite this article: saayman, m. & saayman, a., 2009, why travel motivation and sociodemographics matter in managing a national park, koedoe 51(1), art. #381, 9 pages. doi: 10.4102/koedoe.v51i1.381 this article is available at: http://www.koedoe.co.za © 2009. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. vol. 51 no. 1 page 1 of 9 introduction addo elephant national park (aenp) was proclaimed in 1931 to protect the 11 remaining eastern cape elephants and a few free-roaming buffalo that also remained in the area. in 1954, an elephant-proof fence was constructed to prevent the animals from wandering onto surrounding farmland. the protection programme proved to be successful and the numbers of the animals increased from only 18 in 1954 to well over 100 in 2007. addo is situated close to port elizabeth in the eastern cape in south africa (see figure 1) and lies in gently undulating country, where 90% of vegetation consists of spekboom and other woody species (known locally as addo bush), with karoo scrub and grassland making up the remainder. figure 1 addo elephant national park koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research saayman & saayman a rt ic le # 38 1 (page number not for citation purposes) 50 vol. 51 no. 1 page 2 of 9 today, the park hosts a variety of game species besides elephants. there are more than 160 bird species and the park is also home to the flightless dung beetle. plans to expand the 125 000 ha (309 000 acre) addo elephant national park to a 492 000 ha (1.2 million acre) ‘greater addo’ park are progressing at an exciting pace. the uniqueness of this park lies in the fact that it is one of the few parks in the world that offers the big 7 (elephant, lion, black rhino, buffalo, leopard, the great white shark and seasonal southern right whale in the marine section complete the big 7), which implies that the park is also a marine reserve and that it includes a few islands (sanparks 2009). a socio-economic impact analysis by saayman and saayman (2006) on the aenp indicates that the park plays a significant role in employment, production and general income creation in the region and that 35% of businesses in the region have been established directly as a result of the park. this is due to the growing number of tourists visiting the park and therefore the area and to investments by the private sector. for the aenp to attract the right market as well as to develop the right products and services, it is important that marketers understand the spending behaviour of visitors, for this has direct bearing on the economic impact. south african national parks (sanparks), which is the conservation authority managing national parks in south africa, has, as one of its functions, the important role of creating benefits for local communities by means of job creation and of improvement in the quality of life of local inhabitants. according to van der merwe, saayman and krugell (2006), economic impact is influenced by the length of stay, the number of tourists, the multiplier effect and the amount spent by tourists. hence, a greater understanding of spending behaviour could assist marketers and product developers, firstly in targeting the right market and, secondly, in developing the right products and services. the purpose of this paper is to determine the extent to which travel motivation and socio-demographic indicators influence the spending of tourists to the aenp. lamb, hair and mcdaniel (2002) state that tourists or consumers do not make purchase decisions in isolation. the mix of cultural, social, personal and psychological factors and previous experiences, all of which influence behaviour, is largely uncontrollable. because of the influence exerted on buying patterns, it is essential that as much effort as possible is put into understanding how these factors interact and, ultimately, how they influence these decisions. from the theory of consumer behaviour, personal factors refer to the socio-demographic characteristics of individuals (gender, age, level of education, family life cycle, social class, place of residence etc.) and to those of a psychological nature (motivations, values, personality, lifestyle etc.). these personal factors affect an individual’s cognitive organisation or evaluation of stimuli and therefore also influence the perceptions of the environment and the resulting image (baker & crompton 2000; beerli, josefa & martin 2003). in the socio-demographic profile, behaviour can be described as the mental, emotional and physical activities in which people engage when selecting, purchasing, using and disposing of a product or service to satisfy needs and desires (wilkie 1994). in support of the notion that many factors influence purchase decisions, researchers such as cai, hong and morrison (1995), walker, scott-melnyk and sherwood (1996), lu and pas (1999), jang et al. (2004), saayman and saayman (2006) and van der merwe et al. (2006) all support the notion that socio-demographic determinants have an effect on activity, participation and travel behaviour. for example jang et al. (2004), fish and waggle (1996) and van der merwe et al. (2006) find that socio-demographic variables can be used to explain travel behaviour and the relationship between variables. the only other study that identifies the socio-demographic determinants of spending for a national park in south africa is by saayman and saayman (2007). their results show that a combination of demographic, behavioural and motivational factors influences spending at aenp. behavioural indicators are the most significant in the case of the kruger national park (knp) and include the number of days spent, the size of a travel group, the frequency of visits and catering preferences. their research contradicts findings by downward and lumsdon (2002) and skuras, simara and petrou (2005), who find that an increase in the size of a travel group leads to increased spending. oppermann (1996) finds that repeat visitors spend less than firsttime visitors, although gyte and phelps (1998) find the opposite, while jang et al. (2004) conclude that the frequency of visitation is an influencing factor in visitor expenditure. in the case of the knp, saayman and saayman (2007) show similar results to those of oppermann (1996). language and the province of origin are found by saayman and saayman (2006) to be significant in the case of arts festivals in south africa. the role of age on spending is not conclusive. studies by, for example, mok and iverson (2000) and kastenholz (2005) find a positive relationship between age and spending, while mumdambi and baum (1997) indicate an inverse relationship between age and spending. van der merwe et al. (2006) and letho et al. (2004) find that older people tend to spend more. the reason for or purpose of travel, according to letho et al. (2004) and sakai (1988), has a definite impact on expenditure levels. saayman and saayman (2006) report that attracting high spenders instead of crowds is desirable not only from an economic-impact point of view but also from an environmental point of view. in the context of an attraction such as the aenp, this is important because conservation areas have to create income but with as little environmental impact as possible. with regard to motivational factors, van der merwe and saayman (2008) conducted research on the travel motives of visitors to the knp in south africa. similar research includes that by tao, eagles and smith (2004) for the taroko national park in taiwan and by uysal, mcdonald and martin (1994) for visitors to a national park in the usa. the conclusion of the study by uysal et al. is that visitor motives in visiting various parks differ. to escape from routine was the only motive that repeated itself. this can therefore be regarded as the most common motive for travelling to a tourist destination. no study, however, could be found that combines a motivational and a socio-demographic analysis in trying to get a better understanding of spending by visitors to national parks. from the literature review, it is clear that the determinants of spending differ from destination to destination and from product to product. the issue that then arises is that, if this is indeed the case, there would surely be differences among various national parks and, if so, the issue then is what these differences would be, since the only other comparable study done in national parks that can be used as a reference is that of the knp. method to gather data from visitors to the addo elephant national park, a visitor questionnaire was administered from 2001 to 2007. the method of research is discussed under three headings: (i) the questionnaire, (ii) the samples and (iii) the methods. questionnaire the visitor questionnaire administered at the park has been used in previous national parks research, including the kruger national park, the tsitsikamma national park, the karoo national park and the kgalagadi transfrontier park. the questionnaires were handed out at the camp-sites and chalets in the early evening by fieldworkers and collected later that evening or early the following morning by the same fieldworkers. only overnight visitors are therefore included in the survey. the questionnaire remained fairly consistent over the years from 2001 to 2007, with only minor adjustments made over this time. the questionnaire administered in 2001 can be viewed as the why travel motivation and socio-demographics matter in managing a national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #381 (page number not for citation purposes) 51vol. 51 no. 1 page 3 of 9 testing phase of the questionnaire and most of the problems experienced with data analysis from the 2001 questionnaire were corrected in subsequent years. one example of such a problem is that spending data were gathered in spending categories (e.g. r0–r250, r250–r500 etc.) in 2001, which proved to have limited use in the analysis. the first section of the questionnaire deals with the sociodemographic information of the respondents, including age, marital status and qualification. in the early years of the questionnaire (2001 and 2002), income categories were also included. the response rate on these was very low and this category was subsequently dropped from the questionnaire. the second section of the questionnaire focuses on spending behaviour during the visit to the park and on motivations for the visit. the amount spent on various items is asked, while motivations for the visit to the park are answered on a 5-point likert scale (maree 2007), where 1 = unimportant and 5 = very important. the qualitative questions were coded to enable them to be used in the analysis. the questions used and the codes assigned to them are indicated in table 1. it should be noted that the province variable was coded to reflect income per capita in the province, with 1 = highest income per capita and 10 = lowest income per capita. one would therefore expect the sign of the coefficient to be positive, since people from richer provinces would be expected to spend more. another variable that may need clarification is that of the wildcard. in 2005, sanparks introduced a loyalty card known as the ‘wildcard’. this card can be bought from sanparks and gives the holder free entry to all the national parks in south africa. it thus constitutes a discount on entry and conservation fees. this could therefore either have a negative impact on spending while wildcard holders are at the park or it could be that visitors decide to spend extra on other items, since they are saving on conservation fees. the dependent variable is spending per person, which was calculated by adding the spending of the respondents on the various components asked and subtracting transport cost to the park from the number obtained. this gave total spending excluding transport, which was then divided by the number of people whom the respondents were paying for on the trip to give spending per person. the reasons why transport cost is excluded are that spending on transport does not normally take place in the park and that transport from an origin further away would inflate spending per person. the natural logarithm of all variables was taken to standardise the data and ease interpretation. sample the survey was conducted once yearly from 2001 to 2005 and two surveys have been conducted annually since 2006, which leads to more respondents and could improve conclusions drawn from the survey. the number of questionnaires administered and the months surveyed are indicated in table 2. this table also indicates the total number of visitors to the park during these years, which includes both overnight and day visitors. to have a better idea of the proportion of visitors who are overnight visitors, the unit nights sold (including camping nights) are also indicated. again, this is only an approximation, since most visitors stay for more than one night. overnight travel parties during one year can be derived by dividing the unit nights sold by the average nights spent in the park for that year (according to every year’s survey). if this number is equally divided by 12 months, visitor groups per month can be guesstimated. it can be clearly seen that the response rate for each month of the survey ranges between a low of 4% to a high of 7.4%. this low response could lead to response bias, which should be noted. the surveys are therefore, however, analysed both in crosssection and as a pseudo-panel. table 1 visitor-survey questions used and their descriptions category question description variable socio-demographic home language: english = 1, afrikaans = 2, other = 3 age: < 19 = 1, 20–24 = 2, 25–34 = 3, 35–49 = 4, 50–64 = 5, 65+ = 6 marital status: married = 1, unmarried = 2, divorced = 3, widow/er = 4, living together = 5 residing province: gauteng = 2, western cape = 3, northern cape = 4, mpumalanga = 5, north west = 6, kwazulu-natal = 7, free state = 8, eastern cape = 9, limpopo = 10, non-sa = 1 highest qualification: no school = 1, matric = 2, diploma/degree = 3, postgrad = 4, professional = 5, other = 6 lang. age marry prov. qual. behavioural group size (fill in) number of visits to national parks over past three years (fill in) number of nights (fill in) wildcard (only 2006–2007): yes = 1, no = 2 preference for catering (only 2004–2007): self-catering = 1, dine out & self-catering = 2, b&b = 3, dinner, bed & breakfast = 4 people visits days wild-card pref. motivational scale importance of reasons for visiting park from 1 to 5 (1 = not important; 5 = very important): to get away from my regular routine to relax to explore a new destination to spend time with friends for the benefit of my children for family recreation to learn about wildlife to develop appreciation for endangered species for educational reasons (to increase knowledge) to learn about animals in general to learn about endangered species to learn about plants to learn about specific animals to photograph animals to photograph plants because i grew up with the park it’s a well-known brand the park has great accommodation facilities i prefer this area because of the climate to do the hiking trails for conferences (only since 2003) for events in the area (only since 2003) routine relax explore friends child. family wildlife endang. edu. animals species plants sanim. photoa. photop. grewup brand acc. climate hiking conf. events original research saayman & saayman koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 38 1 (page number not for citation purposes) 52 vol. 51 no. 1 page 4 of 9 statistical analysis to reduce the number of motivations and aid in the interpretation of data, a factor analysis was firstly undertaken. one additional advantage of such an analysis is that it reveals relationships that were not previously suspected (johnson & wichern 2002). the purpose of factor analysis is normally to describe the covariance relationships among many variables in terms of a few underlying but unobservable random quantities known as ‘factors’. according to johnson and wichern (2002), the factor model can be motivated by the following argument: suppose that all variables in a particular group are highly correlated among themselves and can therefore be grouped according to their correlations. the variables in a group do, however, have small correlations with variables in other groups. this being the case, it is conceivable that each group of variables represents a single underlying factor that is responsible for the observed correlations. using the results from the factor analysis, the motivational variables were grouped into the factors (based purely on averages for each factor) and included in the subsequent analyses. to determine the influence of these factors on spending behaviour, regression analysis was undertaken. regressions were first undertaken for each year, in other words cross-sectional regression analysis was undertaken. the model estimated is as follows: yi =α + xi β + εi , with i = 1, 2, 3 . . . n where yi = spending per person, xi = vector of explanatory variables (as indicated in table 1) and εi = i.d.d. error term wooldridge (2002) notes that, in a random sampling context, errors are always independent and identically distributed, regardless of how they are related to x i . to avoid the inclusion of too many variables, a stepwise least squares regression was used. eviews 6 was used in all the estimations. the method used in selecting the variables to be included is the stepwiseforwards, which starts with no additional regressors in the regression and then adds variables based on their p-values. the variable with the lowest p-value is chosen and added, and this process continues until there is no variable left with a p-value less than the included variables (qms 2007). the results of the stepwise regression were then subjected to diagnostic analyses and, where necessary, changes were made to reach the final regression model. since the survey was repeated over a number of years, the data obtained can also be viewed as a panel consisting of both a crosssection and a time component. yet inoue (2008) notes that repeated cross-sections from survey data cannot be treated as genuine panels and a pseudo-panel should therefore be constructed. pseudo-panels are constructed by grouping individuals together according to some characteristic and averaging the observations in each group or cohort (saayman & saayman 2007). since this is done for every cross-section, a time dimension is again obtained (cottrell & gaubert 2003). the pseudo-panel data model that is then estimated is as follows (inoue 2008): yst = αst +δs+θ΄wst+ εst, for s = 1, . . . s, t = 1, . . . , t where s indicates the different cohorts and t indicates time the bar above the variable denotes that it is an average observation, since cohorts are created via averaging individual observations. αst denotes the intercept and captures the average of the individual specific effects that constitute group s, δs captures the group-specific effects and w st captures the grouptime-specific explanatory variables (see table 1 for variables used in the analysis) and the individual-specific characters included in each group. a characteristic often employed to create cohorts is age or date of birth (cottrell & gaubert 2003). thus, for every cross-section, individuals were grouped into the six age groups described in table 1. age group 1 had very few observations in all years, however, and age groups 1 and 2 had to be merged, since it is a prerequisite for pseudo-panel data that the number of individuals in each group must be large relative to the number of groups and time periods. to ensure homoskedasticity in random errors, matas and raymond (2007) suggest that variables are weighted by the square root of the number of individuals in each cohort. this procedure was followed, which rendered all variables continuous. the same procedure was followed as with the cross-section analysis, where a stepwise regression indicated the independent variables that should be included before diagnostic tests were completed. inoue (2008) shows that using fixed effects when estimating the pseudo-panel model accounts for group effects. this is also taken into consideration in the final estimates. results since the questionnaire responses were subjected to three types of analyses, the results are described accordingly. results of the factor analysis the data used in the factor analysis comprised only the motivational factors (as described in table 1). all missing values were identified and any respondent who did not complete at least two-thirds of the motivational questions was omitted from this analysis. to determine the number of factors that should be used in analysis, the rule of thumb is normally that all factors with eigenvalues greater than unity must be extracted (johnson & wichern 2002). using spss 16.0 for windows® (spss inc. 2007), six factors were therefore extracted with eigenvalues greater than unity. together, these six factors explain 64.384% of the variance. the rotated method was chosen, since it provides a simpler structure and more detailed focus. according to both the varimax and promax rotated method, the same structure was found. table 3 indicates the structure according to the varimax procedure. although the variables do not all load very strongly on the factors, the authors tried to keep as many motivational variables as possible for further analysis. the general rule applied is a loading of more than 0.4 on the factor or a loading of less than 0.4 when it loads almost double on one factor than on any other factor. using these basic guidelines, only the motivational factor ‘explore’ does not load clearly onto any factor and it is thus the only motivational factor that is ignored in further analyses. table 2 total number of questionnaires administered to tourists during a marketing survey at addo elephant national park between 2001 and 2007 year 2001 2002 2003 2004 2005 2006 2007 survey month may july december november november july november june november # of questionnaires 64 35 59 82 67 91 50 68 91 total guests 122,123 117,037 132,734 140,178 160,810 162,884 unit nights 26,426 32,021 37,364 43,458 45,749 46,546 visitor groups 8,008 13,922 13,344 12,782 15,250 18,041 groups per month 667 1,160 1,112 1,065 1,271 1,271 1,503 1,503 compiled from visitor statistics received from sanparks (2002–2008) why travel motivation and socio-demographics matter in managing a national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #381 (page number not for citation purposes) 53vol. 51 no. 1 page 5 of 9 given the content of these factors, the motivations for visiting the aenp can be described as follows: factor 1: nature• this motive includes aspects such as educational reasons, endangered species and seeing animals and plants. it has the fourth highest mean value (2.88). the motive is confirmed by researchers such as van der merwe and saayman (2008), oh, uysal and weaver (1995) and swanson and horridge (2006) as a motive for travel. similar research by van der merwe and saayman (2008) completed for the knp in south africa identifies this factor as a primary motive for visiting national parks in south africa. factor 2: activities• ‘activities’ includes the photography of plants, hiking, conferences and events. this motive has also been found as a reason why visitors travel to the knp in south africa (van der merwe & saayman 2008). oh et al. (1995) have also found activities to be an important travel motivation in their research, although, in this case, it has the lowest mean value of all the motives (1.74). factor 3: family and socialisation• this motive includes being with one’s family and friends, and experiencing wildlife and different species with family members – hence the aspect of socialisation. research on national parks in the usa conducted by uysal et al. (1994) identifies a similar motive, namely the enhancement of kinship relations. yet this aspect has not been found to be a priority in knp research. at the knp, the motive of nostalgia is strong, which confirms many other studies in the field (backman et al. 1995; lee, lee & wicks 2004; schneider & backman 1996). possible reasons for this could table 3 results of factor analysis conducted on motivational factors factor 1: nature 2: activities 3: family 4: escape 5: attractions 6: photography mean value (out of 5) 2.88 1.74 2.98 3.83 2.81 3.11 routine 0.049 -0.045 0.047 0.686 0.125 0.031 relax -0.068 -0.012 0.168 0.867 0.105 0.008 explore 0.150 -0.022 0.137 0.184 0.108 0.053 friends 0.030 0.188 0.353 0.084 0.086 0.045 children 0.005 0.155 0.585 0.097 0.022 -0.106 family -0.002 -0.053 0.470 0.393 0.152 0.034 wildlife 0.162 -0.006 0.900 -0.002 0.101 0.061 species 0.348 0.032 0.703 0.088 -0.048 -0.065 education 0.584 0.187 0.291 -0.058 0.033 0.005 animals 0.801 -0.023 0.010 -0.018 0.048 0.114 endanger 0.903 0.018 0.067 0.119 0.059 0.102 plants 0.718 0.250 0.085 0.046 -0.033 0.012 sanim. 0.656 0.219 0.118 0.008 0.058 0.150 photoa. 0.300 0.054 -0.020 0.110 0.079 0.942 photop. 0.316 0.540 -0.097 -0.012 -0.136 0.346 grewup 0.083 0.663 0.065 0.026 0.103 -0.123 brand 0.059 0.531 0.193 0.011 0.271 -0.009 acc. -0.009 0.118 0.173 0.256 0.864 0.010 climate 0.174 0.261 0.031 0.272 0.424 0.068 hiking 0.167 0.598 0.148 0.043 0.101 0.014 conf. 0.096 0.787 0.104 -0.061 -0.024 -0.038 events 0.010 0.478 -0.030 -0.076 0.001 0.128 for a full description of the statements, see table 1 be because it is not only a well-known and established park but also the oldest national park in south africa. this is not the case for the aenp. the motive has a mean value of 2.98, which indicates high relative importance. factor 4: escape• this motive consists of two aspects, namely to relax and to break away from routine. this motive is confirmed by a great number of researchers, such as kim, borges and chon (2006), uysal et al. (1994), swanson and horridge (2006) and van der merwe and saayman (2008). it also has the highest mean value, indicating that it is the most important motive for tourists visiting the aenp. factor 5: attractions• attractions include accommodation and climate, and this motive has a mean value of 2.81. kim et al. (2006) and van der merwe and saayman (2008) also find attractions to be a motive for visiting national parks, although bansal and eislet (2004) find climate to be a separate motive for the visiting of destinations. factor 6: photography• this motive entails the photography of animals; no other research identifies a similar motive. this aspect can therefore be regarded as a unique motive for travel to the aenp. it is also rated second highest, with a mean value of 3.11. results of the cross-sectional analyses based on the motivational factors identified above, the average for each factor was determined for each respondent. these factors (motives 1 to 6 described above) were then included in the regression models. for the purpose of the regression models, only respondents who sufficiently answered the spending section original research saayman & saayman koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 38 1 (page number not for citation purposes) 54 vol. 51 no. 1 page 6 of 9 (thus, have spending data) and data for people for whom she or he was paying could be used in the analysis, since spending per person is the dependent variable. in addition, 2001 spending data were categorised and could not be used, and the crosssection and pseudo-panel analyses were therefore conducted on the data from 2002 to 2007 only. this led to an adjusted sample for each year: (i) 2002, 28 respondents; (ii) 2003, 48 respondents; (iii) 2004, 66 respondents; (iv) 2005, 43 respondents; (v) 2006, 123 respondents; (vi) 2007, 128 respondents. the variables identified for each year by the stepwise regression are indicated in table 4. it is clear from table 4 that both sociodemographic and motivational factors play a role in the spending patterns of visitors to the aenp. these variables were used to determine the final model for each year. in the final models, adjustments to account for heteroskedasticity were made and the models were tested for over-specification. the final cross-sectional models are indicated in table 5. from table 5, the following is evident: a combination of socio-demographic and motivational • factors influences visitor spending decisions. this research therefore confirms similar findings by cai et al. (1995), walker et al. (1996) and lu and pas (1999). the weak proxies for income, namely qualification • and province of origin, appear in all the cross-section regressions. yet they are not significant everywhere. where significant, they show the expected sign (positive), except for 2007, where province of origin has a negative sign. a reason for this may be that most respondents are from the eastern cape province and that, even though it is a relatively poor province, close vicinity to the park may have a positive impact on spending. although age appears in some of the regressions, nowhere • is it a significant determinant of spending, which again supports the notion that the role of age is not conclusive. the marital status of respondents is significant only in 2002. • the negative coefficient indicates that single people tend to spend more per person than married couples. this research supports findings by saayman and saayman (2007) for the knp. a possible reason for this is cost sharing by couples. although language appears as an explanatory variable in • three years, it is significant only in 2004. an interesting result is that the sign is negative, indicating that englishspeaking people spend more than afrikaans-speaking and overseas visitors. this contradicts previous research completed in south africa on arts festivals (saayman & saayman 2006). as expected, the larger the number of tourists who travel • together, the lower the spending per person, since costs are shared. tourists with more elaborate catering needs tend to spend more, which is to be expected. the same applies to tourists who stay longer. in this regard, this research confirms findings by saayman and saayman (2007). an interesting finding is that repeat visitors tend to spend • less, although this is true only for two of the years and the finding is significant only for 2003. this finding supports opperman (1996), who also finds that repeat visitors tend to spend less than first-time visitors, but contradicts findings by gyte and phelps (1998) and jang et al. (2004). while all regressions include motivational factors that • were identified via the stepwise regressions as having an influence on spending, only the motives of nature, photography, escape and attractions are significant. these are also motives with a high mean value. park management should take note of this. results of the pseudo-panel analysis to construct the pseudo-panel, the data were divided into age cohorts. since limited data were available, especially for the early years, only four age cohorts were formed for each year. these were (1) under 24 years, (2) 25 to 34 years, (3) 35 to 49 years and (4) older than 50 years. the data in each cohort represent the average for the group, which was weighted with the square root of the number of observations. finally, the natural logarithm of the variables was taken to standardise the data. the analysis process followed was similar to that of the crosssection analysis. the results of the stepwise regression indicate that all the motivational factors are identified as factors that explain the variance in spending magnitudes. a number of behavioural variables is also important. these variables are the number of people travelling together, the number of times that the respondents visit national parks and the number of nights spent at the park. socio-demographic variables that influenced spending over the years from 2002 to 2007 include the marital status of the respondents, their language and their province of origin. the model was also subjected to various diagnostic tests, including the breusch-pagan-godfrey heteroskedasticity test and normality tests, before the final model was estimated (asteriou & hall 2007, gujarati 2003). in the final model, fixed effects are used to account for differences across various cross sections. the results of the final model are indicated in table 6. from table 6, the following can be concluded: more people travelling together in a group are associated • with lower spending levels per person, since costs are shared. single people tend to spend more than married people. • again, this may be due to married couples sharing costs. more frequent visitors to the park tend to spend more while • at the park. repeat visitation thus has a positive effect. higher spending is associated with people who travel to • the park to relax and get away from the normal routine (escape motive – [4]). motive 6 (the photography of animals) shows a negative • relationship with spending, which may indicate that tourists who focus on photographing animals tend to spend more time driving and searching for the perfect photo rather than spending money at the restaurant or on souvenirs. management implications based on the question of why motivation and socio-demographics matter in managing a national park, the following management implications can be identified from this research: firstly, marketers need to analyse both socio-demographic and motivational influences on spending, since a combination of the former impacts on spending. in the case of national parks, this is imperative because national parks have a socio-economic mandate to fulfil concerning the communities bordering the parks. marketing decisions should therefore be influenced by knowledge of not only the profile of visitors but also the reasons that they visit the parks. to be able to make informed decisions, quality research is a requirement. this implies more than just a normal visitor-opinion survey. table 4 variables identified with stepwise regression from 2002 to 2007 2002 2003 2004 2005 2006 2007 motive 6 people motive 1 marry age qual. motive 5 motive 2 prov. motive 6 visits people prov. pref. days motive 1 age motive 5 motive 4 visits qual. lang. motive 2 prov. prov. qual. motive 6 lang. motive 2 motive 3 motive 1 motive 4 people marry prov. days motive 5 motive 6 motive 1 qual. marry wildcard visits pref. days pref. prov. wildcard qual. lang. motive 1 motive 4 for a full description of the variables used, see tables 1 and 3 why travel motivation and socio-demographics matter in managing a national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #381 (page number not for citation purposes) 2002 2003 2004 2005 2006 2007 variable coeff & prob variable coeff & prob variable coeff & prob variable coeff & prob variable coeff & prob variable coeff & prob constant motive 6 motive 1 people marry age qual motive 2 motive 5 prov 6.912 (0.007)*** -0.615 (0.091)* 1.207 (0.033)** -1.126 (0.051)** -1.294 (0.008)*** -1.773 (0.209) 1.199 (0.052)** 0.647 (0.239) -0.754 (0.092)* 0.222 (0.429) constant visits motive 6 pref people prov days motive 1 age motive 5 3.876 (0.001)*** -0.979 (0.012)*** 1.311 (0.009)*** 1.116 (0.011)** -1.210 (0.008)*** 0.738 (0.005)*** -0.717 (0.0179) 0.576 (0.207) 0.543 (0.140) -0.507 (0.372) constant motive 4 qual lang visits 7.760 (<0.001)*** -0.826 (0.044)** -0.136 (0.755) -0.457 (0.096)* -0.219 (0.179) constant prov qual motive 6 lang motive 2 motive 3 motive 1 5.479 (0.000)*** 0.260 (0.081)* 1.239 (0.068)* -0.534 (0.063)* -0.162 (0.676) -.214 (0.631) -0.651 (0.175) 0.460 (0.208) constant prov days motive 5 motive 6 motive 1 qual 5.944 (<0.001)*** -0.170 (0.219) 0.414 (0.017)*** -0.277 (0.306) -0.091 (0.612) -0.138 (0.636) 0.444 (0.129) constant days pref prov wildcard qual lang motive 1 motive 4 6.369 (<0.001)*** 0.863 (<0.001)*** 0.368 (0.025)** -0.194 (0.033)** -0.151 (0.476) 0.123 (0.507) -0.201 (0.236) -0.119 (0.497) -0.222 (0.354) r-squared adj r-squared f-stat aik sc 0.632 0.412 2.872 2.164 2.651 r-squared adj r-squared f-stat aik sc 0.515 0.387 4.018 3.149 3.555 r-squared adj r-squared f-stat aik sc 0.119 0.037 1.454 2.666 2.861 r-squared adj r-squared f-stat aik sc 0.222 0.013 1.062 2.998 3.357 r-squared adj r-squared f-stat aik sc 0.113 0.062 2.231 2.834 3.004 r-squared adj r-squared f-stat aik sc 0.316 0.262 5.840 2.183 2.404 values in ( ) = probabilities; *** = significant at 1% level; ** = significant at 5% level; * = significant at 10% level. for a full description of the variables used, see tables 1 and 3 55vol. 51 no. 1 page 7 of 9 secondly, parks have to create an environment where people can spend money. according to the results of this research, the motive of ‘activities’, which entails aspects such as hiking, conferences and events, has the lowest mean average. it is therefore an aspect where management can do more in terms of product development and promotions to enhance spending on these items and to improve performance as a motive for visits to the park. photography of animals was identified as the second most-important motive and is also an aspect that has a significant negative impact on current spending. management therefore needs to create workshops on animal photography and promote opportunities such as a photo shop, events and competitions to reap positive benefits from this important motive. thirdly, marketing for the aenp should focus on the motive of escape, since not only does it have the highest mean value but the results also indicate that tourists visiting the park for this motive are high spenders. current marketing efforts focus strongly on the nature motive, specifically elephants and the big 7, while the research indicates that this is only the fourth most-important motive. focusing on the escape motive should be easy to achieve in the setting of national parks and could lead to an increase in high-spending visitors. escape as a motive has been ignored by marketers in national parks, regardless of the fact that all research, both nationally and internationally, identifies escape as a strong and primary motive for visiting tourist destinations and the purchase of tourist products. fourthly, ensuring quality service and products guarantees repeat visits and, coupled to that, the changing needs of tourists have to be taken into account. the pseudo-panel results indicate that, over the six years under investigation, repeat visitors spend more than first-time visitors. conclusion the purpose of this research was to determine the extent to which socio-demographic, behavioural and motivational indicators influence the spending behaviour of tourists to the aenp. the research identifies six motives for tourists travelling to the park. these are nature, activities, family and socialisation, escape, attractions and photography. the research also finds that a combination of socio-demographic and motivational factors influences visitor spending decisions. added to this, the research confirms that tourist attractions, including national parks, differ from one another and that the variables that influence spending therefore also differ. the final regression models indicate that the main motivational factors that have a significant influence on spending are nature, photography, escape and attractions. an interesting finding is that, although photography is regarded as an important motivational factor, it is negatively related to spending, which has significant management implications. this research confirms previous research by saayman and saayman (2007) and van der merwe and saayman (2008) conducted on the knp in south africa by identifying similar motives that influence spending. the motive of escape is also found to be significant for research completed in other parts of the world (e.g. the usa and taiwan). photography as a motive, however, could not be found in any other research. contrary to research by mok and iverson (2000), kastenholz (2005) and letho et al. (2004), age was not found to have a significant influence on spending during visits to the aenp. the finding by saayman and saayman (2007) that more tourists in a travel party leads to lower spending per person at the knp is confirmed for the aenp. greater support for the notion that repeat visitors tend to spend more is also found in the pseudo-panel analysis, thereby supporting previous research by gyte and phelps (1998) and jang et al. (2004). and, while the results for language are not always significant, the coefficient always seems to be negative, indicating that english-speaking tourists tend to spend more at the aenp. this is the opposite to what is found for spending at arts festivals in south africa (saayman & saayman 2006), where afrikaans-speaking visitors tend to spend more. this article contributes to current literature concerning motivational factors that influence the visiting of tourist attractions and events. this is the first time that factor analysis table 5 results of the cross-section estimations (dependent: log of spending per person) original research saayman & saayman koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 38 1 (page number not for citation purposes) table 6 pseudo panel results variable coefficient std. error t-statistic prob. c 4.862458 0.618292 7.864344 0.0002*** motive 2 1.186568 0.616428 1.924909 0.1026 people -1.621337 0.413536 -3.920672 0.0078*** motive 3 -0.145531 1.246690 -0.116734 0.9109 marry 1.401964 0.487521 2.875698 0.0282** motive 5 1.468112 0.906699 1.619183 0.1565 motive 1 -0.555628 1.129816 -0.491786 0.6403 motive 6 -2.273708 0.734315 -3.096364 0.0212** visits 0.789413 0.266143 2.966128 0.0251** motive 4 2.294685 1.132977 2.025359 0.0892* lang. -1.413100 0.802130 -1.761685 0.1286 prov. 0.047514 0.352555 0.134770 0.8972 days 0.109996 0.272923 0.403029 0.7009 effects specification cross-section fixed (dummy variables) cross-section1 -0.1783 cross-section 2 0.2556 cross-section 3 0.0386 cross-section 4 -0.1753 r-squared 0.972223 mean dependent var 7.690098 adj. r-squared 0.902780 s.d. dependent var 0.697375 s.e. of regression 0.217443 akaike info criterion -0.058500 sum squared resid 0.283688 schwarz criterion 0.734985 log likelihood 16.64350 f-statistic 14.00029 durbin-watson stat 1.644189 prob(f-statistic) 0.001888 dependent variable; spendpp method; panel least squares sample; 2002 2007 periods included; 6 cross-sections included; 4 56 vol. 51 no. 1 page 8 of 9 has been used together with regression analysis to determine the influence of different motivational factors on visitor spending. in doing this, sensible management conclusions have been reached. an important methodological aspect of the article that should be highlighted is that cross-section regression results tend to vary significantly for the different years. some results also show very low levels of reliability in terms of r-square and adjusted r-square. the results from the pseudo-panel are more reliable in these terms. one should therefore guard against the reaching of conclusions on one-off surveys and more effort should be taken to repeat surveys over a number of years to test the reliability of results. references asteriou, d. & hall, s.g., 2007, applied econometrics: a modern approach, rev. edn., palgrave, new york. backman, k.f., backman, s.j., uysal, m. & sunshine, k.m., 1995, 'event tourism: an examination of motivations and activities', festival management and event tourism 3(1), 15–24. baker, d.a. & crompton, j.l., 2000, 'quality, satisfaction and behavioural intentions', annals of tourism research 27(3), 785–804. bansal, h. & eislet, h.a., 2004, 'exploratory research of tourist motivations and planning', tourism management 25, 387– 396. beerli, a., josefa, j.d. & martin, d., 2003, tourists’ characteristics and the perceived image of tourist destinations: a quantitative analysis – a case study of lanzarote, spain, university of las palmas de gran canaria, gran canaria. cai, l.a., hong, g. & morrison, a.m., 1995, 'household expenditure patterns for tourism products and services', journal of travel and tourism marketing 4(4), 15–40. cottrell, m. & gaubert, p., 2003, efficient estimators: the use of neural networks to construct pseudo panels, proceedings of the conference wsom 2003, kitakiushu, japan, 11–14 september, n.p. downward, p.m. & lumsdon, l., 2002, 'beyond the demand for day-visits', tourism economics 9(1), 67–76. fish, m. & waggle, d., 1996, 'current income versus total expenditure measures in regression models of vacation and pleasure travel', journal of travel research 35(2), 70–74. gujarati, d.n., 2003, basic econometrics, 4th edn., mcgraw-hill, new york. gyte, d. & phelps, a., 1998, 'patterns of destination repeat business: british tourists in mallorca, spain', journal of travel research 28(1), 24–28. inoue, a., 2008, 'efficient estimation and inference in linear pseudo-panel data models', journal of econometrics 142, 449– 466. jang, s., bai, b., hong, g.s. & o’leary, j.t., 2004, 'understanding travel expenditure patterns: a study of japanese pleasure travellers to the united states by income level', tourism management 25, 331–341. johnson, r.a. & wichern, d.w., 2002, applied multivariate statistical analysis, 5th edn., prentice hall, upper saddle river. kastenholz, e., 2005, 'analysis of determinants of visitor spending for the rural tourist market in north portugal', tourism economics 11(4), 555–569. kim, h., borges, m.c. & chon, j., 2006, 'impacts of environmental values on tourism motivations: the case of fica, brazil', tourism management 27, 957–967. lamb, c.w., hair, j.f. & mcdaniel, c., 2002, marketing, 6th edn., thompson learning, connecticut. lee, c.k., lee, y.k. & wicks, b.e., 2004, 'segmentation of festival motivation by nationality and satisfaction', tourism management 25, 61–70. letho, x.y., cai, l.a., o’leary, j.t. & huan, t.c., 2004, 'tourism shopping preferences and expenditure behaviours: the case of the taiwanese outbound market', journal of vacation marketing 10(4), 320–332. lu, x. & pas, e.i., 1999, 'socio-demographics, activity participation and travel behavior', transportation research part a 33, 1–18. maree, k, 2007, first steps in research, van schaik publishers, pretoria. matas, a. & raymond, j-l., 2007, cross-section data, disequilibrium situations and estimated coefficients: evidence from car ownership demand, document de treball, xreap2007-04: xarxa de referéncia en economia aplicada. mok, c. & iverson, t.j., 2000, 'expenditure-based segmentation: taiwanese tourists to gaum', tourism management 21(3), 299–305. mumdambi, r. & baum, t., 1997, 'strategic segmentation: an empirical analysis of tourist expenditures in turkey', journal of travel research 36(1), 29–34. oh, h.c., uysal, m. & weaver, p.a., 1995, 'product bundles and market segmentation based on travel motivations: a canonical correlation approach', hospitality management 14(2), 123–137. oppermann, m., 1996, 'visitation of tourism attractions and tourist expenditure patterns: repeat versus first-time visitors', asia pacific journal of tourism research 1(1), 61–68. qms (quantitative micro software), 2007, eviews6 user guide ii, qms, irvine. saayman, a. & saayman, m., 2006, 'the socio-demographics and visitation patterns of arts festivals in south africa', events management 9(4), 211–222. saayman, m. & saayman, a., 2007, 'socio-demographic and behavioural determinants of visitor spending at the kruger national park in south africa', paper presented at the first international association for tourism economics (iate) conference, palma de mallorca, spain, 26–29 october. sakai, m.y., 1988, a micro-analysis of business travel demand, applied economics 20(11), 1481–1495. sanparks, 2009, 'greater addo conservation project', viewed 18 may 2009, from http://www.sanparks.org/parks/addo/ library/documents.php. schneider, i.e. & backman, s.j., 1996, 'cross-cultural equivalence of festival motivations: a study in jordan', festival management and event tourism 4(3/4), 139–144. why travel motivation and socio-demographics matter in managing a national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #381 (page number not for citation purposes) 57vol. 51 no. 1 page 9 of 9 skuras, d., simara, e. & petrou, a., 2005, 'rural tourism and visitor expenditure for local food products', regional studies 40(7), 767–779. spss inc., 2007, spss 16.0 for windows®, spss inc, chicago. swanson, k.k. & horridge, p.e., 2006, 'travel motivations as souvenir purchase indicators', tourism management 27, 671– 683. tao, c.h., eagles, p.f.j. & smith, s.l.j., 2004, 'profiling taiwanese ecotourists using a self-definition approach', journal of sustainable tourism 12(2), 149–168. van der merwe, p. & saayman, m., 2008, 'travel motivations of tourists visiting kruger national park', koedoe 50(1), 154– 159. van der merwe, p., saayman, m. & krugell, w.f., 2006, the determinants of the spending of biltong hunters, paper presented at the second australian wildlife tourism conference in western australia, fremantle, australia, 13-15 august. walker, c., scott-melnyk, s. & sherwood, k., 1996, reggae to rachmaninoff: how and why people participate in arts and culture, urban institute, washington dc. wilkie, w.l., 1994, consumer behavior, 3rd edn., wiley, new york. wooldridge, j.m., 2002, econometric analysis of cross-section and panel data, mit press, cambridge ma. uysal, m., mcdonald, c.d. & martin, b.s., 1994, 'australian visitors to us national parks and natural areas', international journal of contemporary hospitality management 6(3), 18–24. abstract introduction study area methods results conclusion acknowledgements references about the author(s) ettienne j. theron department of plant sciences, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa andri c. van aardt department of plant sciences, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa pieter j. du preez † department of plant sciences, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa citation theron, e.j., van aardt, a.c. & du preez, p.j., 2020, ‘vegetation distribution along a granite catena, southern kruger national park, south africa’, koedoe 62(2), a1588. https://doi.org/10.4102/koedoe.v62i2.1588 note: additional supporting information may be found in the online version of this article. online appendix 1. note: special issue: connections between abiotic and biotic components of a granite catena ecosystem in kruger national park, sub-edited by beanelri janecke and johan van tol. †, 1960-2019. original research vegetation distribution along a granite catena, southern kruger national park, south africa ettienne j. theron, andri c. van aardt, pieter j. du preez received: 04 sept. 2019; accepted: 05 may 2020; published: 29 oct. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract this study aimed to investigate how environmental factors drive the assemblage of vegetation within a landscape at various scales, particularly to which extent a sequence of soil forms (catena) influences plant community distribution in a savanna of kruger national park, south africa. on a regional scale, the geology and associated soil forms correspond to the granite lowveld vegetation type. these associations were studied at the scale of a single hillslope, comprising concurrent soil and vegetation surveys from the crest to footslopes. from chemical and physical soil analyses, strong correlations between the distribution of soil moisture content, soil forms and plant species at community, sub-community and variant levels were found. a vachellia exuvialis–pogonarthria squarrosa-dominated savanna is restricted to the well-drained, nutrient-poor, acidic and sandy soils of the crests and upper-midslopes of the granite domes. on sodic sites along lower-midslopes, dactyloctenium aegyptium–sporobolus nitens grasslands dominate the clay-rich soils. the footslopes are characterised by the grass themeda triandra and the shrub flueggea virosa. the grass panicum maximum growing under tall trees such as diospyros mespiliformis and spirostachys africana typifies riparian vegetation along seasonal streams on deep alluvial soil. the association between plant communities and soil forms exemplifies the interdependency of biotic and abiotic components that maintain heterogeneity within the ecosystem from biome to community scale. conservation implications: this article contributes to understanding plant species distribution along a granite catena; an integral part of which are sodic sites that become overutilized by game, which, albeit natural, could severely impact these sites during drought situations. keywords: vegetation classification; vegetation ordination; catenas; sodic sites; soil factors; environmental gradient. introduction vegetation forms an integral part of any ecosystem and can only be fully studied when the role it plays in the ecosystem is explored (kent 2012). identification of vegetation types and plant communities that provide a habitat for animals, insects and birds is, in turn, associated with certain environmental controls and human or animal interactions. understanding the interaction that vegetation has with environmental factors can lead to improved monitoring of management practices and possibly predict future changes (kent 2012). in kruger national park (knp), several vegetation studies were conducted over the past 60 years. however, most were focused on areas where environmental conditions such as fire and herbivory were manipulated. smit et al. (2013) noted that research on areas under natural conditions and where the geology and rainfall vary should be a priority. they identified four supersites that represent the typical rainfall gradient and geological contrast of knp. one of these sites is the stevenson–hamilton research supersite where this study on a catena took place. brady and weil (2002) define a catena as a soil sequence in which each particular soil group occurs on the same parent material, but in different arrays from foothills to crest zones. the study site with its undulating landscape is known to occur on granite, which shows a prevalence of fineto coarse-grained biotite (schutte 1974) (figure 1) and expresses a general soil sequence where sand and clay content are inversely proportional. the catena includes sandy soils that dominate crest zones, clayey soils that are prevalent on midslopes and alluvial sand in the riparian corridors. the soil properties of both sand and clay affect hillslope hydrology and subsequently also vegetation distribution and species composition (venter, scholes & eckhardt 2003). figure 1: location and elevation (meter above mean sea level) of the study site within stevenson–hamilton research supersite (shrs), kruger national park, south africa. in the southern granite regions of knp, catenas can include well, moderately or poorly drained soil forms, depending on the relief. this facilitates the transfer of solutes, colloids or other particles from upslope areas downhill along an environmental gradient, resulting in soil differentiation (conacher & dalrymple 1977; huggett 2007; khomo et al. 2011). khomo et al. (2011) further state that crest soils act as a source of clay particles and ions for downslope areas and can become leached during this process. furthermore, differences in elevation, combined with differences in the water flow pattern in a landscape, can result in soluble salts being transported from higher to lower lying areas. when the salt solute reaches an impermeable clay barrier at the midor footslopes, it can cause surface seepage which eventually evaporates (brady & weil 2002). because salt is not lost through evaporation, it may accumulate in certain areas where rainfall is insufficient to facilitate leaching. in areas with a semi-arid climate, such as the skukuza region, knp, the above-mentioned process often results in the formation of sodic sites on the lower ends of granitic catenas (khomo & rogers 2005). sodic sites are characterised by the hyper-accumulation of exchangeable sodium (na), which impacts the infiltration and flow of water within the affected soils (khomo & rogers 2005; naidu & rengasamy 1993). in the studied part of the park, specific plant communities are associated with lowland sites and elevated clay and colloidal particle content in sodic soils. in addition to these factors, the hyper-accumulation of sodium along with other solutes promotes the establishment of salt-tolerant species. the vegetation structure is typified by short grasses and a lack of trees, fringed by low-growing shrubs. however, the increased nutritional quality and resistance to grazing of the grass sward often leads to over-utilisation by grazers (mcnaughton 1988; moxley 2013). intense grazing or extreme accumulation of salts may indirectly result in the ultimate denudation of a sodic site’s vegetation (khomo & rogers 2005). this apparent degradation of an area would appear to warrant restoration or rehabilitation efforts. however, khomo and rogers (2005) argue that a holistic understanding of the ecology of sodic sites will contribute to improved management thereof, which is essential in a preserved area such as knp. several research papers (smit et al. 2013; van tol et al. 2015; van zijl & le roux 2014) indicated that the sandy crests are dominated by combretum apiculatum and combretum zeyhri, the seeplines by terminalia sericea, the clayey midslopes by vachellia nilotica and the footslopes by euclea divinorium. we investigated the community composition of the different terrain types along the catena aiming to provide a baseline vegetation description and its association with the different soil forms. this forms part of a multidisciplinary study investigating the interactions between biotic and abiotic components along a catena or hillslope in the area. study area from a geological perspective, knp can be divided lengthwise into eastern basaltic formations and western granitoid formations; the latter constituting roughly 60% of the park’s surface (barton, bristow & venter 1986). the study area in the stevenson–hamilton research supersite (figure 1) forms part of the nelspruit suite and consists of archaean granite and gneiss (alard 2009; smit et al. 2013; van zijl & le roux 2014). it is located at 25.111°s and 31.579°e, about 7 km south of skukuza (bredenkamp 1982; mccarthy & rubidge 2005; venter et al. 2003). the prevalence of the granite and gneiss parent material in the undulating landscapes results in shallow, nutrient-poor soils that vary from grey to red to brown in colour (venter et al. 2003). the stevenson–hamilton research supersite forms part of the granite lowveld (svi3) vegetation type that belongs to the savanna biome, which constitutes a large portion of south africa’s vegetation (eds. mucina & rutherford 2006). this vegetation type is characterised by a ground layer dominated by tall grass swards with intermittent trees or other woody species (naidoo et al. 2012; rutherford & westfall 1986). the semi-arid regional climate falls within a summer seasonal rainfall regime, with an average annual precipitation of 650 mm that ranges from a maximum of 1000 mm to as little as 235 mm per annum (naidoo et al. 2012; smit et al. 2013) falling mostly during december and january (kruger, makamo & shongwe 2002). skukuza and its surroundings have a mean annual air temperature of 21°c – 22°c (khomo et al. 2011; scholes, bond & eckhardt 2003), and the seasonal and diurnal temperature fluctuations were shown by kruger et al. (2002) to be largely statistically insignificant, except in the month of march. however, extreme periods of both inundation and aridity are known to occur. methods data collection to measure marked changes in vegetation and spatial environmental gradients, 49 plots of 10 m2 were laid out next to each other to form a 490 m long (straight) belt transect across the study area (brown et al. 2013; kent 2012). the transect stretched from the crest of the hillslope and included plots within the midslope, the ecotone between the midslope and sodic site, the sodic site itself, the ecotone between the sodic site and the riparian area and, finally, the riparian area. this was the only transect where soil moisture and groundwater data were available to aid in the creation of a hydrological soil response map of the study area (figure 2) (van zijl & le roux 2014). however, soil moisture probes were damaged by wildlife during the course of study, rendering the data unavailable for use in the interpretation of the vegetation analysis. the cover abundance of 172 plant species was recorded by means of the modified braun–blanquet cover abundance scale (kent 2012; kent & coker 1992; van der maarel & franklin 2013), which grouped the vegetation in eight categories based on the percentage of ground cover each constituted, respectively (table 1). in selected plots, the vegetation cover was low, and areas of exposed soil were visible. the cover abundance of these exposed areas was recorded as a pseudo-species, indicated as ‘bare soil’. figure 2: plant community, sub-community and variant distribution aligned with the relevés and soil forms (at the bottom) on which they occur along a hillslope gradient. arrows indicate the movement of water through the soil sequence (see bouwer, le roux & van tol 2020 for a detailed description). table 1: braun–blanquet cover abundance scale used – symbols and associated percentage. a soil sample was collected in the middle of each 10 m2 plot laid out along the hillslope from the crest to the drainage line. within each of the 49 sample plots, soil was collected by means of a bucket soil auger to determine the soil depth. all soil samples were examined and analysed independently to determine the soil form and its diagnostic horizons using the soil classification working group’s (1991) classification system. to perform chemical and physical analysis on the soil, modal profiles were sampled at 100 mm depth intervals, dried, sieved and analysed according to standard methods of the non-affiliated soil analysis work committee (1990). particle size distribution was determined using the pipette method (miller & miller 1987). thereafter, soil texture was determined using the texture triangle as indicated in the soil classification working group (1991). the ph (h2o) was measured from 1:2.5 solutions. in addition, soluble and exchangeable cations (ca, mg, k and na) and cation exchange capacity (cec) were determined using nh4oac solution (bouwer et al. 2020). classification vegetation data were captured with vegcap (unpublished database tool designed by n. collins), a macros-enabled excel spreadsheet, and imported to juice© (tichý & holt 2006) where a modified twinspan classification (roleček et al. 2009) analysis was carried out using the pseudo-species cut level (5) and value of cut levels (0, 5, 25, 50 and 75) parameters. the analysis was constrained to a minimum group size of 3–30 clusters with a maximum of 52. division reached an endpoint if dissimilarity went lower than 0.3 based on average jaccard dissimilarity. the resultant clusters were then arranged within both juice© and excel to form the final vegetation communities. no relevés were omitted from the analysis. brown et al. (2013) indicated that a minimum of three relevés per syntaxon is needed for classification. there were, however, instances where only two relevés defined sub-communities with significant differences in terms of the vegetation. this was done to understand the change in vegetation composition along the catena. naming of communities, sub-communities and variants was done according to the guidelines presented in brown et al. (2013). to obtain diagnostic, constant and dominant species, we made use of the analysis of columns of a synoptic table in juice. the frequency thresholds were set at 75, 60 and 50 for the respective diagnostic, constant and dominant species. ordination the species and environmental data were imported to canoco© (ter braak & smilauer 2002) to perform a direct unimodal canonical correspondence analysis. the analysis was set to focus scaling on inter-species distances with biplot scaling. a log transformation was implemented on the data sets with no down weighing of rare species. automatic forward selection and the monte carlo permutation tests were used. lastly, canodraw© was used to create simple ordination plots for the soil and chemical data, respectively, in conjunction with certain selected diagnostic and indicator species. simple ordination plots were also created for the soil and chemical data, respectively, in conjunction with all sample plots. in both cases, the first, second and third axes were used in different combinations, and the most relevant diagrams are shown in the results and discussion. ethical considerations ethical clearance to conduct the study was obtained from the interfaculty animal ethics committee of university of the free state (clearance number: ufs-aed2019/0121). results classification and visualisation hierarchical classification of the stevenson–hamilton research supersite data revealed four distinct plant communities, which include nine sub-communities and two variants (online appendix 1): panicum maximum–pupalia lappacea community themeda triandra–flueggea virosa community 2.1. themeda triandra–flueggea virosa–spirostachys africana sub-community 2.2. themeda triandra–flueggea virosa–cymbopogon nardus sub-community 2.3. themeda triandra–flueggea virosa–ehretia rigida -sub-community vachellia exuvialis–pogonarthria squarrosa community 3.1. vachellia exuvialis–pogonarthria squarrosa–sesamum alatum sub-community 3.2. vachellia exuvialis–pogonarthria squarrosa–vernonia fastigiata sub-community 3.2.1. terminalia sericea variant 3.2.2. grewia flavescens variant dactyloctenium aegyptium–sporobolus nitens community 4.1. dactyloctenium aegyptium–sporobolus nitens–senegalia nigrescens sub-community 4.2. dactyloctenium aegyptium–sporobolus nitens–bothriochloa radicans sub-community 4.3. dactyloctenium aegyptium–sporobolus nitens–urochloa mosambicensis sub-community 4.4. dactyloctenium aegyptium–sporobolus nitens–aristida congesta sub-community these plant communities are distributed from the footslope to the crest with the sodic site indicated as community 4 of the stevenson–hamilton research supersite (figure 2). thus, the communities do not follow the order from the footslope to the crest because of the fact that certain species from the footslope also occur in the crest area. description of plant communities details of the hierarchical vegetation classification are given below with descriptions of each community. 1. panicum maximum–pupalia lappacea community diagnostic species: none constant species: bare soil 100, bidens bipinnata 75, panicum maximum 100, pupalia lappacea 75 dominant species: cynodon dactylon 50, euclea divinorum 25, gymnosporia glaucophylla 25, panicum maximum 100, themeda triandra 25 this community is located in the riparian zone close to the drainage line of the hillside and is found exclusively on dundee soil forms. the soil profiles are relatively deep (600 mm) with a sandy loam texture (avg. phh2o 6.73) and are freely drained; leaching occurs during wet seasons when first-, secondand third-order streams of the study site are in flow. dundee soil forms are characteristically formed by recent sedimentary deposits from streams and rivers and have a diagnostic stratified alluvium horizon (fey 2010), which is consistent with the riparian location of this soil within the study area. community 1 is mainly defined by a grass species panicum maximum (species group u, online appendix 1) and the annual pupalia lappacea (species group a, online appendix 1). although p. maximum occurs steadily throughout all the relevés, it has especially high cover abundance values in the relevés of community 1. bredenkamp (1982) noted in his study that p. maximum was mostly found in the shade of trees. the observation is consistent with this p. maximum community situated in the riparian corridor, canopied by large trees. although not in all relevés, cynodon dactylon (species group a, online appendix 1) occurs in few other relevés and has significant cover abundance within community 1. this is also true for themeda triandra and gymnosporia glaucophylla (species group g, online appendix 1). among the vegetation in this community, there were also patches of bare soil (species group z, online appendix 1). 2. themeda triandra–flueggea virosa community diagnostic species: none constant species: bare soil 100, barleria elegans 67, dichrostachys cinerea 67, euclea divinorum 67, flueggea virosa 83, grewia hexamita 67, panicum maximum 83, senegalia nigrescens 75, stylosanthes fruticosa 67, themeda triandra 83, waltheria indica 75, ziziphus mucronata 67 dominant species: combretum hereroense 8, eragrostis rigidior 8, ipomoea species 17, panicum maximum 17, senegalia nigrescens 8, spirostachys africana 8, themeda triandra 25, ziziphus mucronata 8 community 2 is located between the sodic site and the riparian zone on the footslope. this community is dominated by themeda triandra (species group g, online appendix 1) and flueggea virosa (species group v, online appendix 1). waltheria indica (species group n, online appendix 1) and senegalia nigrescens (kyalangalilwa et al. 2013) (species group s, online appendix 1) are also prominent species present in this community. species such as stylosanthes fruticosa, elaeodendron transvaalense and searsia gueinzii (species group b, online appendix 1) clearly define this community. their cover abundance is very low in this community, and they are mostly absent from the other communities studied. this community is similar to the themeda triandra–senegalia nigrescens tall bushveld communities described by siebert and eckhardt (2008) in terms of species composition and the associated mispah soil form. in terms of species similarity between the studies, prominent species such as themeda triandra, euclea divinorum, waltheria indica and panicum maximum were present in both studies (siebert & eckhardt 2008). community 2 can be divided into the following three sub-communities: 2.1. themeda triandra–flueggea virosa–spirostachys africana sub-community diagnostic species: asparagus cooperi 90.3, carissa bispinosa 100.0, schotia capitata 75.7, spirostachys africana 75.7 constant species: asparagus cooperi 100, bare soil 100, barleria elegans 100, carissa bispinosa 100, commelina benghalensis 100, euclea divinorum 100, ocimum americanum 100, panicum maximum 100, schotia capitata 100, spirostachys africana 100, stylosanthes fruticosa 100 dominant species: spirostachys africana 50 sub-community 2.1 is present on the boundary of the riparian zone and occurs exclusively on the mispah soil form. the soil profiles are relatively shallow (100–200 mm) and consist mainly of sandy clay loam soil, indicative of good drainage (soil classification working group 1991) and the possible onset of salt accumulation (avg. phh2o 6.56). mispah soil forms are relatively youthful because of their close relationship with the parent material, often occurring on diagnostically hard rock (fey 2010) as is consistent with the shallow depths recorded for these relevés. this sub-community is clearly defined by its position as a transitory zone between the riparian zone and lower footslope of sub-communities 2.2 and 2.3 (figure 2). the high cover abundance recorded for spirostachys africana (species group c, online appendix 1) does not occur elsewhere along the transect, except for a single occurrence on the sodic site (community 4). similarly, carissa bispinosa, schotia capitata and asparagus cooperi (species group c, online appendix 1) occur in this sub-community albeit with low cover abundance and are almost absent or show a very low cover abundance in other communities and sub-communities in this study. the species from species group c are thus all regarded as diagnostic species which indicate the uniqueness of these species to this sub-community. when compared to the study by siebert and eckhardt (2008), their sub-community with the strong presence of spirostachys africana can only be compared to the woodland and scrub vegetation found on stream and riverbanks of this study. however, there is no correspondence between species found in siebert and eckhardt’s sub-communities and those of this study. this might be because of the fact that nkhuhlu is bordered by a perennial river, whereas the stevenson–hamilton research supersite is not. thus, at the latter site, the vegetation is dominated more by dry upland species than wet riparian species as indicated by siebert and eckhardt (2008). 2.2. themeda triandra–flueggea virosa–cymbopogon nardus sub-community diagnostic species: cymbopogon nardus 85.4, hibiscus vitifolius 80.4, ipomoea species 85.4 constant species: bare soil 100, barleria elegans 75, cymbopogon nardus 75, dichrostachys cinerea 88, flueggea virosa 88, grewia hexamita 88, hermannia modesta 62, hibiscus vitifolius 88, ipomoea species 75, panicum maximum 75, senegalia nigrescens 75, themeda triandra 88, tragia okanyua 62, vachellia nilotica 75, waltheria indica 75, ziziphus mucronata 75 dominant species: combretum hereroense 12, ipomoea species 25, panicum maximum 25, senegalia nigrescens 12, themeda triandra 12, ziziphus mucronata 12 sub-community 2.2 is located in the middle of the footslope between the riparian zone and the sodic site and occurs predominantly on bonheim soil forms. soil profiles are mostly shallow (381 mm) loam soil, which suggests poorer drainage and increased water retention (avg. phh2o 6.21). bonheim soil forms are typically dark with structured melanic a and pedocutanic b diagnostic horizons. often found in semi-arid regions, the soil forms tend to support sweet grazing lawns that are frequented by herbivores (fey 2010; moxley 2013). this sub-community is distinguished by the presence of diagnostic species such as cymbopogon nardus, hibiscus vitifolius and ipomoea species (species group d, online appendix 1) where some species have high cover abundance values and others low cover abundance values. in contrast, the study by siebert and eckhardt (2008) found that combretum hereroense occurred in most of the vegetation found on the hillslope except next to the river. however, on the transect along the stevenson–hamilton research supersite, c. hereroense was mostly restricted to the footslope of the valley. bredenkamp (1982) also indicated that various combretum species (c. apiculatum, c. zeyheri and c. collinum subsp. zuluense) occur in significant numbers on granitic geological formations in the western parts of knp, and these species were not that prominent at the stevenson–hamilton research supersite. other species that contribute towards defining this sub-community include: lippia javanica, philenoptera violacea, tragia okanyua, lantana rugosa, hypoestes forskaolii, ipomoea oblongata, panicum coloratum, gossypium herbaceum, convolvulus species and barleria oxyphylla (species group d, online appendix 1). 2.3. themeda triandra–flueggea virosa–ehretia rigida sub-community diagnostic species: ehretia rigida 93.6 constant species: aristida congesta 100, aristida meridionalis 100, bare soil 100, diospyros mespiliformis 100, ehretia rigida 100, eragrostis rigidior 100, euclea divinorum 100, flueggea virosa 100, gymnosporia glaucophylla 100, kyphocarpa angustifolia 100, litogyne gariepina 100, ocimum americanum 100, panicum maximum 100, searsia gueinzii 100, senegalia nigrescens 100, stylosanthes fruticosa 100, themeda triandra 100, urochloa mosambicensis 100, waltheria indica 100, ziziphus mucronata 100 dominant species: eragrostis rigidior 50, themeda triandra 100 sub-community 2.3 is located on the boundary of the lower end of the sodic site and occurs exclusively on the sterkspruit soil form (soil classification working group 1991). sterkspruit soil profiles are relatively shallow (100 mm) sandy clay loam soil (avg. phh2o 6.44), suggesting better water retention, but decreased infiltration. the sterkspruit soil form is characterised by increased clay content in the prismacutanic b horizon because of illuviation which can result in impeded water movement and root growth, thus diminishing vegetation cover. furthermore, the soil texture may often lead to crusting and a high degree of erosion (fey 2010). this sub-community represents a sharp ecological transition from the open grazing lawns of the sodic site to the bushy vegetation associated with the riparian zone and footslopes. the transition of soil forms – from mostly bonheim soils in sub-community 2.2 to sterkspruit soils in sub-community 2.3 (figure 2) – further supports the difference in ecological parameters for this sub-community. the diagnostic species for sub-community 2.3 is ehretia rigida (species group f, online appendix 1). although this species has very low cover abundance values, it only occurs in this sub-community and nowhere else in the study area. diospyros mespiliformis (species group f, online appendix 1) also define this sub-community, however, does occur in other communities found in the area of study. there are no comparable plant associations in the studies conducted by siebert and eckhardt (2008). 3. vachellia exuvialis–pogonarthria squarrosa community diagnostic species: vachellia exuvialis 83.4, pogonarthria squarrosa 91.7, sporobolus pyramidalis 77.0, tricholaena monachne 78.9 constant species: vachellia exuvialis 81, aristida congesta 81, bare soil 94, bidens bipinnata 69, combretum apiculatum 69, combretum zeyheri 62, digitaria eriantha 75, kyphocarpa angustifolia 75, melhania acuminata 88, panicum maximum 81, perotis patens 62, pogonarthria squarrosa 88, sporobolus pyramidalis 75, tricholaena monachne 69 dominant species: bothriochloa radicans 6, combretum apiculatum 12, combretum zeyheri 12, digitaria eriantha 6, eragrostis rigidior 19, panicum maximum 6, pogonarthria squarrosa 6, sporobolus pyramidalis 19 community 3 is located on the crest zone and upslope beyond the sodic site. the clovelly soil form is prevalent in this community, but the soil forms pinedene, fernwood and estcourt also occur. this community is defined by species from species group h (online appendix 1), with vachellia exuvialis and pogonarthria squarrosa being the diagnostic species. in the study by siebert and eckhardt (2008), the enneapogon scoparius–combretum apiculatum dry mixed savanna community contains species similar to the ones found in this community even though diagnostic species differ between the studies. significant species for community identification that are shared among these two communities include combretum apiculatum, vachellia exuvialis and pogonarthria squarrosa. community 3 can be divided into two sub-communities and two variants. 3.1. vachellia exuvialis–pogonarthria squarrosa–sesamum alatum sub-community diagnostic species: sesamum alatum 77.5 constant species: vachellia exuvialis 62, agathisanthemum bojeri 62, aristida congesta 88, aristida meridionalis 88, bare soil 100, bidens bipinnata 62, combretum apiculatum 62, combretum zeyheri 62, digitaria eriantha 75, eragrostis rigidior 75, grewia flavescens 62, indigofera filipes 75, kyphocarpa angustifolia 75, melhania acuminata 75, panicum maximum 88, perotis patens 62, pogonarthria squarrosa 88, senegalia nigrescens 62, sesamum alatum 62, sporobolus pyramidalis 62, tricholaena monachne 62, waltheria indica 75 dominant species: combretum apiculatum 25, combretum zeyheri 12, eragrostis rigidior 25, pogonarthria squarrosa 12 sub-community 3.1 occurs on both fernwood and clovelly soil forms. the fernwood soil profiles are of intermediate depth (533 mm) and consist of loamy sand (avg. phh2o 5.95). fernwood soil forms characteristically have a diagnostic coarse-sand horizon with no clear structure or stratification and have a loose consistency (fey 2010). the clovelly soil profiles are deep (620 mm) with a loam sandy texture (avg. phh2o 6.2). this soil form characteristically has a yellow to brown apedal b horizon because of iron oxides that accumulate through weathering (fey 2010). species group i (online appendix 1) defines the sub-community. the diagnostic species in the group, sesamum alatum, is restricted to this sub-community and does not occur anywhere else in the area of study. other species with a high occurrence in this sub-community are constant species such as aristida congesta (species group q, online appendix 1), aristida meridionalis (species group r, online appendix 1) and also several bare soil patches (species group z, online appendix 1) (janecke, 2020). other species in this sub-community include: eragrostis rigidior, e. superba, indigofera filipes, cissus cornifolia, balanites maughamii, mundulea sericea, dipcadi gracillimum, cyperus indecorus, hibiscus schinzii, rhynchosia caribaea, e. gummiflua, cymbopogon pospischilii, chamaecrista mimosoides, bulbostylis hispidula and b. burchellii (species group i, online appendix 1). gertenbach (1986) worked on a landscape scale in terms of vegetation classification and described the senegalia nigrescens/combretum apiculatum association as thickets along the sabie and crocodile rivers. in contrast to his study, these species did not dominate the vegetation but were present as components of a larger community. 3.2. vachellia exuvialis–pogonarthria squarrosa–vernonia fastigiata sub-community diagnostic species: chamaecrista absus 75.6, vernonia fastigiata 80.4 constant species: vachellia exuvialis 100, aristida congesta 75, aristida stipitata 62, bare soil 88, bidens bipinnata 75, chamaecrista absus 88, combretum apiculatum 75, combretum zeyheri 62, digitaria eriantha 75, grewia bicolor 75, heteropogon contortus 62, justicia protracta 62, kyphocarpa angustifolia 75, melhania acuminata 100, melinis repens 62, panicum maximum 75, perotis patens 62, pogonarthria squarrosa 88, rhynchosia totta 88, sporobolus pyramidalis 88, terminalia sericea 75, tricholaena monachne 75, trichoneura grandiglumis 75, vernonia fastigiata 88 dominant species: bothriochloa radicans 12, combretum zeyheri 12, digitaria eriantha 12, eragrostis rigidior 12, panicum maximum 12, sporobolus pyramidalis 38 sub-community 3.2 is located on the crest of the hillslope and occurs mostly on the clovelly soil form, with single occurrences of the soil forms mispah and sterkspruit. the clovelly soil profiles are deep (616 mm) and consist of coarse loamy sand (avg. phh2o 6.08). the diagnostic species unique to this sub-community are vernonia fastigiata and chamaecrista absus (species group j, online appendix 1). these species also occur in other sub-communities but with very low cover abundance values. the research of siebert and eckhardt (2008) only had different vernonia species present in different communities; however, none, which were similar to the combination of diagnostic species, were found in this study. other species that also define this sub-community include: heteropogon contortus, rhynchosia totta, melinis repens, justicia protracta, setaria sphacelata, jatropha zeyheri, heliotropium ciliatum, kyllinga alba and vangueria infausta (species group j, online appendix 1). sub-community 3.2 encompasses two variants. the terminalia sericea variant (3.2.1) contains the following species: jatropha zeyheri, heliotropium ciliatum, kyllinga alba, vangueria infausta, aptosimum lineare, rhigozum zambesiacum and setaria megaphylla, whereas the grewia flavescens variant (3.2.2) contains most prominently sclerocarya birrea. 4. dactyloctenium aegyptium–sporobolus nitens community diagnostic species: chloris virgata 86.9, dactyloctenium aegyptium 83.1, schkuhria pinnata 87.8, sporobolus nitens 81.5 constant species: bare soil 94, chloris virgata 89, dactyloctenium aegyptium 89, indigofera tinctoria 61, kyphocarpa angustifolia 78, schkuhria pinnata 100, sporobolus nitens 94, urochloa mosambicensis 78 dominant species: bare soil 6, dactyloctenium aegyptium 11, schkuhria pinnata 11, sporobolus nitens 44, vachellia nilotica 6 community 4 is located on the lower-midslope sodic site and occurs almost exclusively on the sterkspruit soil form, with only two instances of the mispah soil form recorded. this community is defined by species from species group n (online appendix 1) where the grasses dactyloctenium aegyptium and sporobolus nitens have high cover abundance values. within this community, there are also various bare soil patches (species group y, online appendix 1) (janecke, 2020). other species that are prominent in this community include schkuhria pinnata, chloris virgata, vachellia nilotica, indigofera tinctoria, geigeria burkei, justicia flava, ocimum americanum and cyperus austro-africanus (species group n, online appendix 1). siebert and eckhardt (2008) also found a sodic site at the nkhuhlu study area. the sporobolus nitens–euclea divinorum dry sodic savanna community indicated high cover abundance of the grass sporobolus nitens which is similar to that of this study. furthermore, they found several tree species at their site which is probably the reason for the dry sodic savanna designation. however, in this study most of the woody species, if present at all, were small shrubs. the sodic site of this study was relatively open and mostly covered by grasses and forbs. in contrast to the study by siebert and eckhardt (2008) where only two sub-communities were identified, this study identified four different sub-communities. 4.1. dactyloctenium aegyptium–sporobolus nitens–senegalia nigrescens sub-community diagnostic species: none constant species: aristida congesta 100, bare soil 100, chloris virgata 80, dactyloctenium aegyptium 100, digitaria eriantha 80, indigofera tinctoria 80, kyphocarpa angustifolia 100, ocimum americanum 100, schkuhria pinnata 100, senegalia nigrescens 80, sporobolus nitens 80, urochloa mosambicensis 80 dominant species: none sub-community 4.1 occurs mostly on the sterkspruit soil from. the soil profile proved to be of intermediate depth (200–500 mm) and consisted on average of coarse sandy loam soil (avg. phh2o 6.20). this sub-community is defined by ctenium concinnum, eragrostis biflora, ocimum filamentosum and achyranthes aspera (species group o, online appendix 1). these species mostly occur in this sub-community; however, there are instances where they are present in other sub-communities but with very low cover abundance values. senegalia nigrescens (species group r, online appendix 1) is also present in this sub-community and absent from most of the other sub-communities in community 4. 4.2. dactyloctenium aegyptium–sporobolus nitens–bothriochloa radicans sub-community diagnostic species: none constant species: bare soil 83, bothriochloa radicans 67, chloris virgata 100, dactyloctenium aegyptium 83, justicia flava 83, kyphocarpa angustifolia 83, melhania acuminata 67, schkuhria pinnata 100, solanum panduriforme 83, sporobolus nitens 100, urochloa mosambicensis 100 dominant species: dactyloctenium aegyptium 17, sporobolus nitens 83, vachellia nilotica 17 sub-community 4.2 occurs exclusively on the sterkspruit soil form. the soil profile remained shallow (250 mm) and consisted on average of coarse sandy loam soil (avg. phh2o 6.24). there are few other species besides b. radicans (species group s, online appendix 1) that define this sub-community. however, from online appendix 1, it can be seen that species from species groups o, p, q, r and v are totally or mostly absent from this sub-community. 4.3. dactyloctenium aegyptium–sporobolus nitens–urochloa mosambicensis sub-community diagnostic species: none constant species: bare soil 100, blepharis integrifolia 100, chloris virgata 100, dactyloctenium aegyptium 75, litogyne gariepina 75, schkuhria pinnata 100, sporobolus nitens 100, urochloa mosambicensis 75 dominant species: schkuhria pinnata 50, sporobolus nitens 50 sub-community 4.3 occurs exclusively on the sterkspruit soil form, with the exception of one occurrence of the mispah soil form. the soil profiles remained shallow (230 mm) and consisted on average of coarse sandy loam soil (avg. phh2o 6.38). this sub-community is defined by the presence of: blepharis integrifolia, alternanthera pungens, litogyne gariepina and portulaca kermesina (species group v, online appendix 1). although urochloa mosambicensis is present in most of the relevés within community 4, it has the highest cover abundance in this sub-community. this sub-community can further be defined by the absence of species from species groups o, p, q, r, s and t (online appendix 1). 4.4. dactyloctenium aegyptium–sporobolus nitens–aristida congesta sub-community diagnostic species: none constant species: aristida congesta 67, bare soil 100, chloris virgata 67, dactyloctenium aegyptium 100, geigeria burkei 67, indigofera tinctoria 67, kyphocarpa angustifolia 67, ocimum americanum 67, schkuhria pinnata 100, sporobolus nitens 100, vachellia nilotica 100 dominant species: bare soil 33, dactyloctenium aegyptium 33, sporobolus nitens 33 sub-community 4.4 occurs exclusively on the sterkspruit soil form. the soil profile remained shallow (180 mm) and consisted on average of coarse sandy loam soil (avg. phh2o 6.43). there are few other species besides aristida congesta that define this sub-community. this sub-community is mostly defined by the absence of species that define other sub-communities in community 4. ordination of plant communities and selected environmental variables the same species used by bredenkamp (1982) are depicted in figure 3 (shown by black outlined boxes) in relation to soil forms identified along the hillslope. distinct relationships are shown between species and textural soil properties where certain species are related to a certain soil textural class and a specific soil form. vachellia exuvialis is associated with coarse, deep sandy soils of the clovelly form found within the crest zone (sub-community 3.1) (species group h, online appendix 1). panicum maximum is more likely to be associated (higher cover abundance values [online appendix 1]) with the dundee soil form found only in the riparian zone of community 1: panicum maximum–pupalia lappacea community (species group u, online appendix 1). themeda triandra is shown as equidistant from two environmental variables: soil form and increased clay content. from a catenary perspective, the position of community 2: themeda triandra–flueggea virosa supports this relationship. situated downhill from the sodic site, which acts as a source of clay particles, relevés 35–44 show an increased average clay content (figure 4) and alignment with the bonheim soil form (figure 2). figure 3: simple ordination diagram of the canonical correspondence analysis (first and second axes) showing the relationships between selected species’ cover abundance, soil texture and soil forms. the eigenvalues of the respective axes were 0.642 for axis 1 and 0.523 for axis 2. figure 4: sand, silt and clay content per soil sample collected for each relevé along the study transect. the relationship between selected species and soil chemical properties (the non-affiliated soil analysis work committee 1990) is visualised along the first and third axes in figure 5. although the majority of species are grouped to the left of the vertical axis without pronounced associations with chemical soil properties, there are very strong correlations between cations found at the sodic site and the species that occupy this habitat (present on the right of the vertical axis). the grouping of both exchangeable (ions that can be taken up by the plants) and soluble (ions that can be leached out by water) na, k and mg with species representative of the sodic site community (community 4: dactyloctenium aegyptium–sporobolus nitens community) is congruent with the catenary premise that lower midslope sodic sites act as sinks for dissolved particles and salts. figure 5: simple ordination diagram of the canonical correspondence analysis (first and third axes) showing the relationships between selected species’ cover abundance and soil chemical properties. the eigenvalues of the respective axes were 0.659 for axis 1 and 0.100 for axis 3. conclusion this study is the first in knp to directly link the hydropedological conditions of a catena to the distribution of plant communities along the gradient of a granitic supersite. sharp ecotones in the vegetation can be attributed to the differences in soil texture, hydropedological conditions and chemical properties of soil. the classification of vegetation communities and the spatial alignment thereof with soil forms and their physical and chemical properties, in this study, again confirmed the close association between fine-scaled plant communities and the individual soil forms on which they occur. the correlation of the abiotic (soil type) and biotic (plant community) parameters that define plant abundance and distribution within this ecosystem is indicative of the interdependency of different ecosystem components on landscape heterogeneity. the results of this study indicate clear differences between plant communities along a catena. these differences can be ascribed to various ecological drivers of which soil seems to play an important role. as a result, the heterogeneity and species composition as well as suitability for various animal species changes along the catena and probably contributes to ecosystem resilience. acknowledgements the authors thank the south african national parks for providing them with access to the research sites within kruger national park. they are grateful to the field rangers who accompanied them during the surveys. they also thank louis scott and leslie brown for suggestions on the writing of the article. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions e.j.t. and p.j.d.p. were responsible for the fieldwork, data collection during field surveys, and data analysis and interpretation. a.c.v.a. contributed towards the analysis and interpretation of the plant communities. all authors contributed to the writing of the article. funding information the authors are grateful to the university of the free state (ufs) strategic research fund for partially funding this multidisciplinary research. data availability data from all research done within kruger national park is placed within the sanparks repository (not for free, open access). disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references alard, g.f., 2009, ‘a comparison of grass production and utilization in sodic and crest patches on a semi-arid granitic savanna catena in the southern kruger national park, south africa’, msc thesis, faculty of science, university of the witwatersrand. barton, j.m., bristow, j.w. & venter, f.j., 1986, ‘a summary of the precambrian granitoid rocks of the kruger national park’, koedoe 29(1), 39–44. https://doi.org/10.4102/koedoe.v29i1.518 bouwer, d., le roux, p.a.l. & van tol, j., 2020, ‘identification of hydropedological flowpaths in stevenson-hamilton catena from soil morphological, chemical and hydraulic properties’, koedoe 62(2), a1584. https://doi.org/10.4102/koedoe.v62i2.1584 brady, n.c. & weil, r.r., 2002, the nature and properties of soils, 13th edn., prentice hall, upper saddle river, nj. bredenkamp, g.j, 1982, ‘’n plantekologiese studie van die manyeleti-wildtuin’, phd thesis, departement plantkunde, university of pretoria. brown, l.r., du preez, p.j., bezuidenhoudt, h., bredenkamp, g.j, mostert, t.h.c. & collins, n.b., 2013, ‘guidelines for phytosociological classifications and descriptions of vegetation in southern africa’, koedoe 55(1), art. #1103, 10. https://doi.org/10.4102/koedoe.v55i1.1103 conacher, a.j. & dalrymple, j.b., 1977, ‘the nine unit based landsurface model. an approach to pedogeomorphic research’, geoderma 18(1–2), 1–154. https://doi.org/10.1016/0016-7061(77)90087-8 fey, m., 2010, soils of south africa: their distribution, properties, classification, genesis, use and environmental significance, cambridge university press, cape town. gertenbach, w.p.d., 1986, ‘landscapes of the kruger national park’, koedoe 26(9), 9–121. https://doi.org/10.4102/koedoe.v26i1.591 hugget, r.j., 2007, fundamentals of geomorphology, 2nd edn., routledge taylor & francis group, new york, ny. janecke, b.b., 2020, ‘vegetation structure and spatial heterogeneity in the granite supersite, kruger national park’, koedoe 62(2), a1591. https://doi.org/10.4102/koedoe.v62i2.1591. kent, m., 2012, vegetation description and data analysis: a practical approach, 2nd edn., wiley-blackwell publishers, west sussex. kent, m. & coker, c., 1992, vegetation description and analysis: a practical approach, bellhaven press, west sussex. khomo, l.m., hartshorn, a.s., rogers, k.h. & chadwick, o.a., 2011, ‘impact of rainfall and topography on the distribution of clays and major cations in granitic catenas of southern africa’, catena 87(1), 119–128. https://doi.org/10.1016/j.catena.2011.05.017 khomo, l.m. & rogers, k.h., 2005, ‘proposed mechanism for the origin of sodic patches in kruger national park, south africa’, african journal of ecology 43(1), 29–34. https://doi.org/10.1111/j.1365-2028.2004.00532.x kruger, a.c., makamo, l.b. & shongwe, s., 2002, ‘an analysis of skukuza climate data’, koedoe 45(1), 1–7. https://doi.org/10.4102/koedoe.v45i1.16 kyalangalilwa, b., boatwright, j.s., daru, b.h., maurin, o. & van der bank, m., 2013, ‘phylogenetic position and revised classification of acacia s.l. (fabaceae: mimosoideae) in africa, including new combinations in vachellia and senegalia’, botanical journal of the linnean society 172(4), 500–523. mccarthy, t. & rubidge, b., 2005, the story of earth & life: a southern african perspective on a 4.6-billion-year journey, struik nature, cape town. mcnaughton, s.j., 1988, ‘mineral nutrition and spatial concentrations of african ungulates, nature 334, 343–345. https://doi.org/10.1038/334343a0 miller, w.p. & miller, d.m., 1987, ‘a micro-pipette method for soil mechanical analysis’, communications in soil science and plant analysis 18(1), 1–15. https://doi.org/10.1080/00103628709367799 moxley, c., 2013, ‘characterization of biotic and abiotic sodic lawns of the kruger national park using the framework of the positive feedback loop’, honours thesis, department of biological sciences, university of cape town. mucina, l. & rutherford, m.c. (eds.), 2006, the vegetation of south africa, lesotho and swaziland, strelizia 19, south african national biodiversity institute, pretoria. naidoo, l., cho, m.a., mathieu, r. & asner, g., 2012, ‘classification of savanna tree species, in the greater kruger national park region, by integrating hyperspectral and lidar data in a random forest data mining environment’, isprs journal of photogrammetry and remote sensing 69, 167–179. https://doi.org/10.1016/j.isprsjprs.2012.03.005 naidu, r. & rengasamy, p., 1993, ‘ion interactions and constraints to plant nutrition in australian sodic soil’, australian journal of soil research 31(6), 801–819. https://doi.org/10.1071/sr9930801 roleček, j., lubomír, t., david, z. & chytry, m., 2009, ‘modified twinspan classification in which the hierarchy respects cluster heterogeneity’, journal of vegetation science 20(4), 596–602. https://doi.org/10.1111/j.1654-1103.2009.01062.x rutherford, m.c. & westfall, r.h., 1986, ‘biomes of southern africa – an objective categorization’, memoirs of the botanical survey of south africa 54, 1–98. scholes, r.j., bond, w.j. & eckhardt, h.c., 2003, ‘vegetation dynamics in the kruger ecosystem’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 242–262, island press, london. schutte, i.c., 1974, ‘n geologiese verkenningsopname van die noord-sentrale gedeelte van die nasionale krugerwildtuin, geological survey, pretoria. siebert, f. & eckhardt, h.c., 2008, ‘the vegetation and floristics of the nkhuhlu enclosures, kruger national park’, koedoe 50(1), 126–144. https://doi.org/10.4102/koedoe.v50i1.138 smit, i.p.j., riddell, e.s., cullum, c. & petersen, r., 2013, ‘kruger national park research supersites: establishing long-term research site for cross-disciplinary, multiscaled learning’, koedoe 55(1), art. #1107, 7 pages. https://doi.org/10.4102/koedoe.v55i1.1107 soil classification working group, 1991, soil classification: a taxonomic system for south africa, department of agricultural development, pretoria. ter braak, c.j.f. & smilauer, p., 2002, canoco reference manual and canodraw for windows user’s guide: software for canonical community ordination version 4.5, computer software, microcomputer power, ithaca, ny. the non-affiliated soil analysis work committee, 1990, handbook of standard soil testing methods for advisory purposes, soil science society of south africa, pretoria. tichý, l. & holt, j., 2006, juice: a program for management, analysis and classification of ecological data, vegetation science group, masaryk university brno, brno. van der maarel, e. & franklin, j., 2013, vegetation ecology, 2nd edn., john wiley & sons, ltd., west sussex. van tol, j.j., van zijl, g.m., riddell, e.s. & fundisi, d., 2015, ‘application of hydropedological insights in hydrological modelling of the stevenson-hamilton research supersite, kruger national park, south africa’, water sa 41(4), 525–533. https://doi.org/10.4314/wsa.v41i4.12 van zijl, g. & le roux, p., 2014, ‘creating a conceptual hydrological soil response map for the stevenson hamilton research supersite, kruger national park, south africa’, water sa 40(2), 331–336. https://doi.org/10.4314/wsa.v40i2.15 venter, f.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation pattern’, in j.t. du toit, k.h. rogers & h.c biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83–129, island press, london. filelist convert a pdf file! page 1 abstract introduction conceptual pillars for adaptive and transformative learning methods results discussion conclusion acknowledgements references about the author(s) craig a. mcloughlin centre for ecosystem science, faculty of science, university of new south wales, sydney, australia riverine landscapes research laboratory, university of new england, armidale, australia eddie s. riddell conservation management, south african national parks (sanparks), skukuza, south africa centre for water resources research, faculty of agricultural, earth and environmental sciences, university of kwazulu-natal, pietermaritzburg, south africa robin m. petersen scientific services, south african national parks (sanparks), skukuza, south africa jacques venter scientific services, south african national parks (sanparks), phalaborwa, south africa citation mcloughlin, c.a., riddell, e.s., petersen, r.m. & venter, j., 2021, ‘adaptive and transformative learning in environmental water management: implementing the crocodile river’s ecological reserve in kruger national park, south africa’, koedoe 63(1), a1663. https://doi.org/10.4102/koedoe.v63i1.1663 note: additional supporting information may be found in the online version of this article as online appendix 1, 2 and 3. original research adaptive and transformative learning in environmental water management: implementing the crocodile river’s ecological reserve in kruger national park, south africa craig a. mcloughlin, eddie s. riddell, robin m. petersen, jacques venter received: 13 nov. 2020; accepted: 17 feb. 2021; published: 20 apr. 2021 copyright: © 2021. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract freshwater biodiversity loss in the anthropocene escalates the need for successful environmental water management to sustain human benefitting ecosystem services. of the world’s river basins, one-third are now severely water depleted, rendering the quality and quantity of water to maintain or restore freshwater ecosystem integrity increasingly urgent. however, managing environmental water is intricate because of complexity and uncertainty in interacting social and biophysical system components, and trade-offs between costs and benefits of implementing environmental flows. learning enabled adaptive management – embracing the uncertainty – is essential; however, practising adaptive management (worldwide) is challenging; single-, doubleand triple-loop learning is required, along with social learning, to tackle complex problems. there is progressive realisation of environmental flows (ecological reserve) in the crocodile river, south africa, linked to the kruger national park, using strategic adaptive management (sam). in this research article, we reflected on adaptive (singleand double-loop) learning and transformative (triple-loop) learning capacity emergent in sam between 2009 and 2019 whilst also considering social learning potentials. we found evidence of preconditions (e.g. transparency) for social learning within a burgeoning stakeholder ‘community-of-practice’, likely fostering capacities (e.g. information sharing) for sustained social learning. adaptive and transformative learning is enabled by social learning, underpinned by ongoing nested feedbacks supporting assessment and reflection, which facilitates single-, doubleand triple-loop learning. champions exist and are vital for sustaining the adaptive management system. executing adaptive and transformative learning aids in positive change across the range of ecological, social and economic outcomes that are essential for success in environmental water programmes, worldwide. conservation implications: crocodile river ecological reserve implementation, associated with kruger national park, provided an important national precedent (lessons) for protecting the ecological integrity of river systems – obligatory under the national water act (act no 36 of 1998). we demonstrated the importance of ongoing stakeholder learning for successful management of the ecological reserve. keywords: adaptive management; ecological reserve; environmental flows; feedbacks; kruger national park; single-, double-, triple-loop learning; social learning. introduction the anthropocene is characterised by ubiquitous loss of biodiversity at unprecedented rates and scales (kingsford, bino & porter 2017a). globally, this biodiversity loss is severest across the freshwater realm (albert et al. 2021; vorosmarty et al. 2010), which exhibits the highest species diversity per unit area (pittock et al. 2015). over the last half century, alteration to natural flows in rivers – from land-use change, water over-abstraction and building of dams – has contributed towards more than 80% reduction in the freshwater species population (harwood et al. 2017; wwf 2020 – freshwater living planet index). currently, one-third of the world’s river basins are severely water depleted (harwood et al. 2017), thereby, rendering the quality and quantity of water to maintain or restore freshwater ecological integrity (e.g. habitats) increasingly urgent (wwap 2018). setting allocations of water for the environment in environmental flow policy and programmes, and monitoring and evaluation of outcomes are now imperative for sustained delivery of human benefitting ecosystem services (e.g. drinking water, food) emanating from rivers (cf. harwood et al. 2017; nel & roux 2018; tickner et al. 2020; webb et al. 2018). managing freshwater resources in complex, interacting social and biophysical systems (anderson et al. 2019; biggs et al. 2015; cilliers 2008) – achieving diverse values and goals – is an arduous undertaking (harwood et al. 2017; pahl-wostl et al. 2013) fraught with uncertainty (biggs et al. 2015; rogers et al. 2013). uncertainty is exacerbated by poor water governance historically (harwood et al. 2017; pahl-wostl et al. 2013) and impacts from climate change (palmer et al. 2009; pittock & max finlayson 2011). moreover, escalating competition for limited water resources is fuelling growing conflict (nel & roux 2018; wwap 2018) with implicit trade-offs between costs and benefits of environmental water, as evidenced in australia’s murray-darling basin (chen et al. 2020; schoeman, allan & finlayson 2019; thoms, rose & dyer 2020; webb et al. 2018). adaptive management is the foremost approach for effective and successful planning and delivery of environmental water (mcloughlin, thoms & parsons 2020; nel & roux 2018; webb et al. 2018). adaptive, learning-by-doing strategies assume that complex social–ecological systems are in a state of flux, and understanding is always imperfect (biggs & rogers 2003; gunderson 2015; rogers 2003). management expects to face substantial uncertainty (biggs et al. 2015; stankey, clark & bormann 2005), and therefore, seeks to remain flexible in the achievement of goals (anderson et al. 2019; mcloughlin et al. 2020). generically, adaptive management encompasses a series of actions characterised by feedback loops, with deliberate intent to achieve goals through the modification and refinement of hypotheses, objectives, outputs or outcomes and of management actions (kingsford et al. 2017b). this iterative process is supported by strategic monitoring, with feedbacks and learning from the outcome of decisions (mcloughlin & thoms 2015). learning amongst all stakeholders is key to success (pahl-wostl 2009; rogers, roux & biggs 2000; roux et al. 2017), and to be effectual requires three modes: ‘adjusting, improving existing routines’ (single-loop learning), ‘reframing, changing practice’ (double-loop learning), and ‘reviewing norms and values, and transforming governance’ (triple-loop learning) (fabricius & cundill 2014; pahl-wostl 2009; pahl-wostl et al. 2011a). singleand double-loop learning (adaptive), and triple-loop learning (transformative) help to drive the achievement of goals within uncertain contexts, and this is performed by aiding modification and improvement of policies, approaches and actions whilst also transforming governance (pahl-wostl et al. 2013; mcloughlin & thoms 2015). sustaining the practice of adaptive management, with learning, is not without its challenges; limited evidence exists for successful applications in natural resource management (cf. nel & roux 2018; pahl-wostl et al. 2011b; rist, campbell & frost 2013; stankey et al. 2005; susskind, camacho & schenk 2012). main barriers to learning in adaptive management practice include a deficit in trust and cooperation across institutions and organisations, and ingrained norms of action without reflection, which can impede learning (kingsford et al. pahl-wostl et al. 2017b; nel & roux 2018; pahl-wostl et al. 2011a). furthermore, often, there is insufficient stakeholder collaboration with shared learning, as shown in the glen canyon dam (united states [us]) adaptive management project where adverse impacts on species and habitats eventuated despite planned interventions for protection (garmestani & allen 2015; susskind et al. 2012). indeed, successful learning in adaptive management relies on individuals, their perceptions, experiences, social relations and networks that function as the web binding the adaptive system together (nel & roux 2018; pahl-wostl 2009) in stakeholder participation and social learning processes (ernst 2019; reed et al. 2010). social learning is an iterative learning and negotiation process rooted in a specific context, with communication comprising many feedback loops for adaptation to ongoing change (ison & watson 2007; pahl-wostl & hare 2004). active involvement of stakeholders and building of a sense of ownership during decision-making processes strengthen commitment, thus promoting consensus for achieving agreed outcomes (pahl-wostl 2009). this research study demonstrates adaptive (single-loop and double-loop) learning and transformative (triple-loop) learning in practice, and importance of this learning for striving towards successful management of environmental water in water-stressed and contested river systems. we use a case study of the crocodile river in the north-eastern part of south africa, linked to the kruger national park (knp), where implementation of environmental flows (ecological reserve) is being pioneered (harwood et al. 2017; jackson 2015; riddell et al. 2014) using strategic adaptive management (sam) (mcloughlin et al. 2011a; pollard, du toit & biggs 2011; roux & foxcroft 2011). we reflect on adaptive and transformative learning capacity emergent in the crocodile river sam system, over a decade from 2009 to 2019. learning capacity is interpreted through a lens of single-, doubleand triple-loop learning assimilated within a heuristic sam cycle framework (cf. mcloughlin & thoms 2015), in conjunction with a stakeholder social learning assessment. the heuristic framework describes desirable (often theoretically based) feedbacks that drive adaptive and transformative learning, thus nurturing stakeholder social relations (cf. anderson et al. 2019). it affords direction to the learning narrative presented. the authors argue that meaningful progress with ecological reserve delivery in the crocodile river reflects trustworthy, cooperative and flexible management – moving through modes of single-, doubleand triple-loop learning – enabling positive change and objectives achievement. ‘learning-by-doing’ is supported by a rapid response system (rrs), where swift and reliable feedbacks of information allow timely decisions on river flow operations for the adjustments when needed. this article underscores key learning experiences, which are applicable to environmental water management across south africa and further abroad. conceptual pillars for adaptive and transformative learning single-, doubleand triple-loop learning originating from the management theory (hargrove 2002), triple-loop learning builds on the double-loop learning concept developed by argyris and schon (1978). this is by increasing the time scales for change by considering different management and governance levels that provide direction and stability in social contexts (fabricius & cundill 2014; pahl-wostl 2009). it differs from single-loop learning, which results in the incremental advances from action strategies without questioning underlying assumptions (pahl-wostl 2009). single-loop learning involves a continuation of, with concurrent improvements to, established practices and routines in targeting the achievement of goals (mcloughlin & thoms 2015; pahl-wostl 2009). in comparison, double-loop learning refers to a change in the actual frame of reference and includes a re-visitation of initial underlying assumptions of any action (fabricius & cundill 2014; pahl-wostl 2009). social learning processes (see below) – trust, cooperation and buy-in between stakeholders, for example – are vital in double-loop learning (pahl-wostl 2009; pahl-wostl et al. 2011a). the reframing process commonly occurs within stakeholder networks characterising the resource governance regime, and improvements are achieved by experimenting with innovative approaches. stakeholders involved in double-loop learning normally explore reframing in the context of structural constraints of governance systems, such as regulatory frameworks (pahl-wostl 2009). triple-loop learning refers to a change in the structural constraints. it includes transformation of the factors that determine the frame of reference of issues and a transformation of the entire governance regime itself (pahl-wostl 2009). this style of transformation necessitates an acknowledgement that paradigms and structural constraints inhibit effective reframing of resource governance and management practices. thus, triple-loop learning implies a paradigm shift, as well as changes in the norms and values underlying the processes of governance (pahl-wostl 2009; pahl-wostl et al. 2011a). social learning social learning assists natural resource practitioners to deal with complex situations by building a shared understanding amongst stakeholders, cooperation and trust (cundill et al. 2011; daniel & walker 1996). acquiring of new information and experiences, via participation and social interactions, can lead to changes in the perception of individuals (reed et al. 2010). however, such change should go beyond individuals and include relational (e.g. improved sense of community), cognitive (e.g. change of perspectives) and technical (e.g. communication skills) dimensions (muro & jeffrey 2012). there are two prominent constructs to social learning in natural resource management (cf. pahl-wostl et al. 2007; pahl-wostl, mostert & tàbara 2008). firstly, processing of factual information involves a problem task or content management about a specific problem. secondly, engagement in social exchange emphasises social relations, which are inextricably connected to management problems, because managers must consider whose problems to solve and how to frame them. integration of the ‘social’ and ‘content’ components is facilitated via ‘relational practices’, for example, quality of interaction, shared ownership of tasks, transparency for mutual testing of options and contradiction, and suitable opportunities for reflexive sessions (mcloughlin et al. 2020) in problem-solving activities. thus, natural resource management is not only composed of technical qualities, for example, improvement in the condition of ecosystems, but relational qualities too, for example, improved capacity of stakeholders to solve conflicts by achieving consensus through cooperation. a number of key criteria have been found to influence social learning dynamics in a group of stakeholders practising adaptive resource management; these criteria are summarised and described in table 1 across three categories. the first category includes a set of preconditions for social learning to take place (cf. mostert et al. 2007). the second category relates to the emergence of a stakeholder ‘community-of-practice’ (sensu wenger 1998), as a vital mechanism driving social learning. evolution of a community of practice depends on a number of criteria, as displayed in table 1 (cf. iaquinto, ison & faggian 2011; pahl-wostl et al. 2008). the third category relates to key capacities, which enable social learning to occur (cf. cundill et al. 2011; pahl-wostl et al. 2007; pahl-wostl & hare 2004), and often these capacities emerge from within a community of practice. table 1: key criteria expected to influence social learning in a group of stakeholders practicing adaptive resource management, spread across three categories – ‘pre-conditions for social learning’, ‘community-of-practice’ and ‘capacities for social learning’. generally, a context of trust, network building, shared understanding and conflict resolution shape stakeholder’s social learning (ernst 2019), emergent within processes of single-, doubleand triple-loop learning (pahl-wostl 2009). success results from ongoing learning and negotiation through communication within trusted networks, perspective sharing and development of adaptive group strategies (huxham & vangen 2000; pahl-wostl & hare 2004). notably, ‘reframing’ required during doubleinto triple-loop learning would be difficult to achieve without social learning (fabricius & cundill 2014; pahl-wostl et al. 2011a). active involvement of stakeholders and building of a sense of ownership during decision-making processes strengthen commitment, thus promoting consensus for achieving agreed outcomes in natural resource management (pahl-wostl 2009). heuristic strategic adaptive management cycle framework the sam cycle framework (mcloughlin & thoms 2015) is designed primarily to complement the sam (roux & foxcroft 2011) variant of adaptive resource management. each sam cycle has two phases: an adaptive planning and an adaptive implementation phase (figure 1). the adaptive planning phase (represented in the black boxes in figure 1) is focused on developing objectives, culminating in the explicit, measurable end-point goals of the sam cycle. end-point goals are the thresholds of potential concern (tpcs), and they recognise the natural variability of selected response indicators by incorporating upper and/or lower levels (thresholds) of acceptable change (mcloughlin et al. 2011b). thresholds of potential concern are viewed as hypotheses of acceptable change, and open to challenge and refinement forming an inductive approach to adaptive management (biggs & rogers 2003; rogers & biggs 1999). the adaptive implementation phase includes five components (represented in the grey boxes in figure 1). these are concerned with processes of selecting the best intervention options to meet goals, determining inputs for planning (associated with meeting goals), operationalising inputs via implementation of the plans, checking adequacy of plan implementation by response to operational outputs, assessing suitability of the operational outputs by auditing strategic outcomes (against tpcs) and testing achievement of the broader objectives within the adaptive management system. figure 1: the adaptive phases and components of the heuristic strategic adaptive management (sam) cycle framework. the sam cycle has no distinct evaluation step – rather, an ongoing and overlapping feedback system, which drives assessment and reflection facilitating adaptive (singleand double-loop) learning and transformative (triple-loop) learning. the learning component in the heuristic sam cycle framework is structured as an essential nested and ongoing process, rather than a distinct step (figure 1). there are two learning potentials recognised – adaptive and transformative learning – facilitated by the sam cycle adaptive feedback system (mcloughlin & thoms 2015). adaptive learning includes singleand double-loop learning. two types of single-loop learning – lower and upper – allow for, respectively, more immediate responses to check operational outputs (thin, solid arrows in figure 1), and adaptive assessments to audit strategic outcomes against explicit and measurable end-point goals (hashed arrows in figure 1). adaptive feedbacks for single-loop learning are mandatory because this is the actual doing, where progress is made within adaptive management (fabricius & cundill 2014). double-loop learning (dotted arrows in figure 1) is about reframing problems and solutions, which is facilitated by adaptive reflection to test achievement of the agreed objectives, considering the occurrence of any surprises. the skill lies in achieving a balance between the use of single-loop and double-loop learning (fabricius & cundill 2014). reviewing values and objectives along with transformation of governance arrangements (folke et al. 2005) is triple-loop learning (thick, solid arrows in figure 1). transformative (triple-loop) learning is important because if governance remains too rigid, this impedes the reframing potentials of double-loop learning (pahl-wostl 2009). time scales for these feedbacks might include daily or weekly (single loop, lower), 1–3 years (single loop, upper), 4–6 years (double loop) and 8–10 years (triple loop). the main objective of this research study was to explore the role of adaptive (single-loop and double-loop) learning and transformative (triple-loop) learning in crocodile river management, related to the reasonable success with ecological reserve delivery in knp, over a decade since 2009. we deployed the heuristic sam cycle framework as a tool for guiding our learning investigation, integrating adaptive and transformative learning processes into the sam system (roux & foxcroft 2011). another objective was to include a stakeholder social learning assessment, because this learning is integral whilst progressing through the modes of single-, doubleand triple-loop learning (ernst 2019; pahl-wostl 2009). we used a quantitative assessment based on stakeholder responses (from interviews) to statements, concerning the key criteria for social learning taken from the literature (see table 1). assessment results offered valuable insights into ‘potentials’ for social learning amongst the crocodile river stakeholders, although the authors concede that there were no definitive answers – further social learning research is needed. still, improved understanding on social learning potentials helped to support the ecological reserve related learning narrative presented in this article. with this learning narrative, the final objective of this study was to offer lessons from the accompanying experiences for ecological reserve delivery in other catchments and water management areas of south africa. methods study area the crocodile river is one of three river systems of the inkomati water management area (iwma) located in the north-eastern region of south africa (figure 2). the water management area has a catchment size of 28 757 km2, of which the crocodile river catchment is 10 400 km2. rainfall is strongly influenced by topography, varying from over 1200 mm per annum in the west over the drakensberg escarpment to as low as 400 mm per annum over the lowland plains in the east. natural mean annual runoff in the iwma is estimated at 3188 million m3 per annum, whilst present-day conditions yield 2058 million m3 per annum (bailey & pitman 2016). the crocodile river flows eastward and confluences with the komati river just before entering mozambique, when it becomes the incomati river flowing into the indian ocean (figure 2). consumptive water uses associated with the crocodile river include industry, irrigated agriculture, domestic water supply and plantation forestry. the non-consumptive water uses are the specified minimum flow requirement to mozambique and the south african ecological reserve, which is critical for maintaining biodiversity values of the knp (see rountree & rogers 2004; russell 1997; van coller, rogers & heritage 2000). figure 2: the crocodile river catchment and its position in relation to the kruger national park (knp). ecological water requirement (ewr) sites, river flow gauging weirs, ecological monitoring sites in knp and the kwena dam are indicated. the crocodile river is important ecologically as it forms the southern boundary of knp (figure 2), a world-renowned protected area whose river landscapes contribute markedly to aquatic and terrestrial biodiversity (rountree & rogers 2004; russell 1997; van coller et al. 2000). the knp has a well-developed history of sam (pollard & du toit 2007; roux & foxcroft 2011): there are river-related tpcs (e.g. todd & thirion 2011), which are explicit and measurable end points (operational and biodiversity) of an objectives hierarchy (see biggs et al. 2011a; mcloughlin et al. 2011b) linked to knp’s vision for maintaining biodiversity in all its natural facets and fluxes (rogers & bestbier 1997). the tpcs are indicators that assist management to respond to biodiversity change resulting from human-induced pressures within the catchment, predominantly upstream and outside the protected area (biggs & rogers 2003; kingsford et al. 2017b). the majority of the crocodile river catchment is dominated by unregulated river reaches, but the main stem is regulated by the kwena dam in the upper catchment (figure 2). the dam has a gross storage capacity of 158 million m3, and the primary use is to supplement water supplies for irrigated agriculture in the middle and lower reaches (dwa 2010). there are smaller dams, weirs and water supply schemes throughout the catchment, which transfer water to towns, and these alter river flow and water quality, and also increase sedimentation rates. management interventions are designed to maintain or return tpc indicators towards more natural ranges, necessitating cooperation and integration of management activities between different catchment institutions (kingsford et al. 2017b). crocodile river operations committee the national water act (act no. 36 of 1998) supports stakeholder-centred, adaptive and sustainable use and protection of freshwater-linked ecosystems (o’keeffe & rogers 2003; roux & nel 2013). at the level of the water management area, the minister of human settlements, water and sanitation promotes management of freshwater resources by assigning powers and duties to catchment management agencies (cmas) (section 73[4] of the national water act). the inkomati–usuthu cma was established in 2004 to manage the freshwater resources of the iwma, but later including the usuthu-to-mhlathuze water management area. a responsibility of cmas is to develop a catchment management strategy (see dwaf 2007 for a full treatise), and in 2010, the inkomati–usuthu cma drafted its first such strategy in collaboration with all stakeholders (icma 2010). it incorporates sam principles, including development of a vision, mission and objectives (jackson 2015; rogers & luton 2011). set up by the inkomati-usuthu cma, the crocodile river operations committee (crococ) is an institutional arrangement, which oversees management of the freshwater resources linked to the crocodile river (harwood et al. 2017; jackson 2015), and serves as a technical subcommittee to the broader crocodile river forum. the crococ stakeholders include the inkomati-usuthu cma, department of water and sanitation, crocodile river major irrigation board (crmib) and, more recently, those in the tributaries, knp, mbombela municipality, nkomazi municipality, ehlanzeni district municipality, komati basin water authority, and consultants or research-related personnel. the crococ was initially set up to meet every 2 months, with ad hoc meetings if and when required, and the committee plays an active role in integrated planning and operations of the crocodile river system. this includes provision of important decision-making advice about implementation of the ecological reserve using this as one of its performance metrics, along with meeting international flow requirements and maintaining an assurance of supply to consumptive uses. the crocodile river ecological reserve under the national water act, the ecological reserve is defined as the quantity and quality of water required in freshwater resources to protect the ecological functions on which humans depend (o’keeffe & rogers 2003). for river systems, it is the recommended flow regime and water quality expected to maintain a specific agreed river ecological category, rather than reinstating pristine conditions (dwaf 1997). ecological categories are derived via an eco-classification procedure (see kleynhans & louw 2007) and range from a (natural) to f (highly impacted), representing an order with decreasing levels of protection for (or increasing levels of risk to) freshwater aquatic species and habitats (kleynhans & louw 2007). the intention is to create a balance between the environmental, economic and social benefits emanating from river resources. once agreed by catchment stakeholders, the ecological category (included in an overall management class; pollard et al. 2011a) is used to plan for the environmental water requirement of ecological reserves in different river systems. in 2010, the crocodile river’s comprehensive ecological reserve determination study was undertaken (dwa 2010). there have been concerted efforts since then to implement and manage the low-flow requirements of the river’s ecological reserve (maintaining ecological category ‘c’), mainly during the critical dry winter and spring months (april–november). implementation of the low-flow ecological reserve (hereafter, ‘ecological reserve’) is aided by flow gauging stations located along the river (figure 2), which enable remotely accessible collection of near real-time river flow data. furthermore, sophisticated hydrological models (e.g. water resource modelling platform, dwaf 2008) are deployed, with several ecological water requirement sites (figure 2) used to determine flow requirements for the ‘c’ ecological category river. biophysical monitoring is performed at demarcated sites (figure 2) along the knp sections of the crocodile river (and upstream to the headwaters; roux et al. 2018). notably, knp tpcs intend to prioritise biodiversity monitoring along these demarcated sites, and tpc auditing enables appropriateness of the managed ecological reserve to be assessed (mcloughlin et al. pollard et al. 2011b). there is no management plan for the high-flow or flooding requirements of the river, as specified in the ecological reserve determination study (dwa 2010). however, larger flooding events are not affected by the kwena dam, which is located high up in the catchment (figure 2). social learning assessment potential for social learning within the crococ was assessed based on stakeholder values, experiences and perceptions concerning the forum. a total of 12 stakeholders were purposefully sampled from a cross-section of sectors represented in the crococ forum: inkomati–usuthu cma (3), crocodile river major irrigation board (2), south african national parks (3), silulumanzi – mbombela local municipality water supplier (1), komati basin water authority (1) and iwr water resources (2). after obtaining ethics approval and informed consent from all participants, in-depth, semi-structured interviews were conducted with stakeholders in 2013. each stakeholder was interviewed individually where the dialogue about the managed system was initiated. discussions focused on key criteria taken from the literature (table 1), which, for this study, were grouped into three relevant categories, as displayed in table 1: ‘preconditions for social learning’, ‘community-of-practice’ and ‘key capacities for social learning’ often evolving from within a community of practice. on conclusion of each discussion session per social learning criterion (per category), the stakeholder was asked to indicate how much they agreed (or not) with a statement related to that criterion (online appendix 1), based on his or her experiences within the forum. for example, ‘there is mutual engagement between the various stakeholders, and this is ongoing’. quantitative data were collected by employing a likert-scale scoring system ranging from 1 to 10, where 1 = ‘strongly disagree’, 5 = ‘neutral’ and 10 = ‘agree completely’. mean scores were calculated per criterion across the three categories and presented graphically in bar charts. qualitative data were also collected during the 2013 stakeholder interviews; however, these data were not used for the purposes of this research study and its objectives, although valuable for future study (see discussion). single-, doubleand triple-loop learning in order to centre our narrative on adaptive and transformative learning capacity, initially, we selected key ecological reserve-related elements (e.g. tpcs, ecological monitoring and hydrological modelling) from crocodile river management, which existed at the beginning of the period 2009–2019 (online appendix 2). we then assigned the selected elements to applicable components of the sam cycle framework (mcloughlin & thoms 2015; figure 1). adaptive planning: summary objectives of the knp water in the landscape objectives hierarchy (see mcloughlin et al. 2011b) includes maintaining the ‘c’ ecological category river. the objectives culminate in the tpcs (online appendix 2; see mcloughlin et al. 2011b). adaptive implementation: current management interventions and planning inputs to apply the ecological reserve, including operational and strategic monitoring, governance arrangements, actions to deliver the ecological reserve, monitoring of ecological conditions, checking operational outputs and auditing of strategic outcomes or tpcs (online appendix 2). the ecological reserve-related elements selected (online appendix 2) are by no means an exhaustive set. for instance, water quality is excluded, yet recognised and used as an important ecosystem driver, and examples of monitoring (related to biodiversity objectives or tpcs) stem from the knp river reaches, although monitoring occurs upstream (outside) of the protected area (roux et al. 2018). moreover, social and economic objectives of the broader crocodile river sam system (jackson 2015; rogers & luton 2011) are important and recognised, but not taken further in this study (achieving these objectives entails comparable and concurrent learning processes). nonetheless, we consider the selected elements to be sufficient for presenting a succinct but informative learning narrative. we then identified key experiences from within the crocodile river sam system (related to ecological reserve management) over the period 2009–2019 and categorised these into observations of change according to the sam cycle framework: single-loop, double-loop and triple-loop learning (figure 1). we sourced information from journal articles, research or technical reports (e.g. water research commission), internal south african national parks documents or reports (and discussions with staff) and, importantly, from our personal experiences with the sam system. the adaptive and transformative learning capacity narrative was then constructed around these observations (examples) – representing key learning experiences (single-, doubleand triple-loop learning) – and also outcomes from the social learning assessment. notably, any adaptive management system is an ongoing evolutionary process (allan & stankey 2009); hence, the narrative of this study makes reference to promising future single-, doubleand triple-loop learning processes, where appropriate. ethical considerations ethics approval for the study was obtained from the human research ethics committee, ethics office, research development & integrity research division of the university of new england, armidale, nsw, australia – he13-240. results social learning assessment preconditions for social learning by 2013, there were promising preconditions for social learning within the crococ stakeholder group, given that all seven criteria had an average score > 6 (figure 3a) (numbers in brackets below link to figure 3; see table 1 for criterion descriptions). this included five criteria rated 7.5 and higher: ‘role of stakeholder involvement’ (1), ‘type of facilitation’ (4), ‘transparency of the approach’ (5), ‘framing of problems’ (6) and ‘resources’ (7). transparency is vital for promoting social learning amongst stakeholders and was scored the highest by the crococ. this suggests a high legitimacy of the work undertaken by the stakeholder group, which is achieved by developing a system of ongoing feedbacks with joint planning and setting of clear ground rules. however, the criterion ‘representativeness’ (3) has the lowest average score though > 5, suggesting that relatively, increased effort is needed to include pertinent stakeholders in the forum in order to increase organisational representation. figure 3: scores (mean) indicating how much stakeholders agreed (or not) with a statement related to each key criterion influencing social learning potentials within the crocodile river operations committee, across three categories: (a) ‘preconditions for social learning’, (b) ‘community-of-practice’ and (c) ‘capacities for social learning’. the mean scores range from 1 to 10, where 1 = ‘strongly disagree’, 5 = ‘neutral’ and 10 = ‘agree completely’ (standard error bars indicated; white triangles represent minimum-value scores obtained; number placed beside each key criterion link to numbers in brackets within the text; see table 1 for criterion descriptions). community-of-practice there is evidence for an emerging crococ ‘community-of-practice’, as all eight criteria received an average score > 6 (figure 3b) (numbers in brackets below link to figure 3; see table 1 for criterion descriptions). of specific interest is ‘recognition of coordinator’ (6), which has the highest average score at 8.83. social learning requires a coordinator who is well networked and accepted within the stakeholder group. the inkomati-usuthu cma has played a key role in this regard, setting up and harmonising interactions of the crococ stakeholders, and acting as a neutral and trusted coordinator that is championing the management process. other relatively high scoring criteria include ‘stakeholder interaction’ (2), ‘management support’ (7) and ‘networking’ (8). although rated > 5, the lowest scoring criterion includes ‘limitations of the group’ (5), suggesting that stakeholders require a greater ability to identify their limitations in order to improve their actions over time. capacity for social learning the crococ stakeholders have been building their capacity for social learning since prior to 2013, given that 11 criteria have an average score of ≥ 6 (figure 3c) (numbers in brackets below link to figure 3; see table 1 for criterion descriptions). notably, five criteria scored ≥ 8, including ‘cooperation’ (4), ‘trust’ (6), ‘communication’ (8), ‘exchange of ideas’ (11) and ‘information sharing’ (12). this suggests that stakeholders are informed about issues, and their views and opinions are listened to. stakeholders are learning how to work together with group communication likely strong, and they generally view initiatives as a learning process. furthermore, stakeholders realise the value of sharing information, exchanging of ideas and being open to new ways of doing things. respect for one another is also being built up as stakeholders seem to be willing to listen to each other’s point of view, and processes are perceived to be open and fair. this situation promotes collaboration when key decisions must be made. the lowest scoring criterion includes ‘informal interactions’ (7), with an average score of 4.8 (minimum score 1.5), suggesting that emphasis is placed on the more formal interactions. thus, the forum should be looking into ways of promoting informal arrangements, because this is where social learning is often nurtured. the second lowest scoring criterion includes ‘understanding complexity’ (5), although > 6, suggesting that relatively more could be performed to increase stakeholder’s conceptualisation of the social and biophysical components, importantly the complex interactions between these components. this is to help with consensus in decision-making and collaborative solving of complex problems that arise. single-, doubleand triple-loop learning lower-grade single-loop learning (rapid response system) the rrs (figure 4; online appendix 2) drives this learning in the crocodile river sam system. it is facilitated by rapid (daily or weekly) email or telephone messaging amongst the crococ stakeholders, and more recently, through visual communication tools using social media platforms, such as whatsapptm (e.g. for real-time gauge verification and incident reporting). the rrs was designed to include different ‘worry-levels’ relative to the low-flow requirements of the ecological reserve (benchmarks are calculated each week). there is an increased urgency if and when the different ‘worry-levels’ are breached (figure 4), as measured at the tenbosch gauging weir (figure 2). stakeholders work together to determine triggers for the deteriorating river flows, thereby coming up with mitigation actions. for example, river flows may be augmented via dam releases and/or water restrictions imposed on irrigation. importantly, feedbacks are executed as soon as possible so that actions can ideally be executed prior to the river flow declining unacceptably. figure 4: example of the deployment of the rapid response system, within strategic adaptive management of the crocodile river ecological reserve. in reality, river flow sometimes falls into the ‘medium’ and even ‘high’, worry levels, often explained by unprecedented weather conditions, such as unusually high temperatures and/or evaporation rates. one of the early associated learnings was realisation of the knp staff that meeting daily requirements downstream becomes increasingly challenging during critical low-flow periods. simply, it is because the nature of water abstractions increase as one moves downstream, compounded by energy tariffs from the national electricity utility, electricity supply commission (eskom). this means that irrigators seek to optimise for access to water and energy, with a ‘3-day average rule’ being accepted. notably during the year 2017, this factor prompted crococ stakeholders to engage with eskom in order to negotiate a workable solution but, unfortunately, to no avail. recommendations for management of the ecological reserve and actual rrs operations emanate from bimonthly meetings of the crococ. a monthly management log (see mcloughlin et al. 2011b) provides valuable feedback on actions and measures undertaken (e.g. percentage of river flows per ‘worry-level’), used to decide on successes and/or failures enabling adaptations and improvements to the rrs. for example, daily hydrological data received at the tenbosch gauging weir became increasingly unreliable, indeed, because of lack of maintenance on its data loggers, a situation compromising trust relations built up. although working cooperatively, stakeholders agreed to purchase and install dependable data loggers and upkeep them regularly, thereby reducing river gauge measurement uncertainties, which occasionally lead to shortfalls in the various short-term allocations (even under restrictions) during times of water stress. a largely self-reliant community of practice was nurtured across disparate sectors, that is, without the need for relying on government interventions. the above-mentioned example fosters confidence in river flow monitoring and is important for meaningful engagement within the rrs approach to ecological reserve compliance. upper-grade single-loop learning (adaptive assessment) this learning is driven by adaptive assessment linked to the tpcs and associated monitoring processes in the crocodile river sam system (see mcloughlin et al. 2011b). post monitoring, tpc audit reports (e.g. fish tpcs) are completed using data and information collected at crocodile river sites along the knp boundary. the tpc process is a ‘red-flag’ concept and may call into question the appropriateness of river flows in line with achieving amongst others the knp ecological objectives. ultimately, tpc auditing informs management about risks of losing the ‘c’ ecological category river, thus supporting longer term decisions associated with operational adjustments to ecological reserve implementation. for example, the risk of deteriorating fish assemblages (i.e. exceedance of fish tpcs) contributed towards a decision to alter the frequency of rrs benchmark calculations (ecological reserve, using data from tenbosch gauging weir; figure 2) from an original 3-month interval to a weekly calculation. the higher temporal resolution in the calculation allows for judicious stakeholder responses to undesirable river flows in the crocodile river. the rrs ecological reserve benchmark has undergone further refinement in recent years following the gazetting of the ecological reserve to allow for a compliance certainty band around the targeted flows. the benchmark currently allows for a 70% assurance (maintenance flows), with some variation for drought (up to 90% assurance), including the absolute limit of the 99% assurance rule. this in itself was cross-institutional learning, applying a less rigid (dynamic) and cumbersome (reduced worry level triggers) compliance reporting system, to meet the institutional needs of inkomati-usuthu cma. notably, this operating rule now includes compliance with the international flow obligations towards the republic of mozambique (figure 2). over time, such refinement to the ecological reserve benchmark allows for increased flexibility in the approach to achieving the crocodile river’s flow targets. ongoing management decisions are needed in conjunction with progressive implementation of the ecological reserve in the crocodile river, particularly if or when tpcs become exceeded, or deemed to be close to exceedance. for instance, the volume of water released from kwena dam is altered relative to the different rrs ‘worry-levels’ to avoid undesirable declines in the river’s flow regime. importantly, to foster upper-grade single-loop learning crococ stakeholders must keep perusing the summary statistics emanating from the rrs (see mcloughlin et al. 2011b), because its purpose was to enhance stakeholder feedbacks and actions related to delivery of the ecological reserve, that is, meeting the calculated benchmarks each week. double-loop learning (adaptive reflection) two key types of feedback and adaptive reflection opportunity arise from within the crocodile river sam system, and these facilitate the ‘reframing’ of complex management problems. firstly, assumptions (hypotheses and/or models) associated with prevailing tpcs can and should be altered based on any newly acquired information and understanding. for example, although based on robust research, there was much subjectivity involved in application of the original geomorphic tpcs developed for the knp mixed alluvial-bedrock controlled rivers (see rogers & bestbier 1997; rountree & rogers 2004), particularly by non-geomorphologists. furthermore, the focus was on a ‘gradual, incremental change’ over time in the macro-morphology of the river channel, but this proved to be unrealistic for red-flagging problems before these arise. new strategies suggested monitoring to be carried out only in the critical in-stream habitat, that is, within the most sensitive sites where suspect sedimentation build-up can be detected quickly. using aerial imagery, the tpc process includes assessing change in the active channel width (instream habitat availability) and functioning of exposed bedrock habitats important for biota (mcloughlin et al. 2011b). the new tpcs are cost-effective and logistically more feasible to implement, which are measured against actual ecological categories of the managed ecological reserve. further double-loop learning examples related to tpcs exist, for instance, dominant understanding – about biotic indicator responses to unnatural sedimentation in mixed alluvial-bedrock controlled rivers – was questioned (e.g. breonadia salicina tpcs; see mackenzie, van coller & rogers 1999; mcloughlin et al. 2011b for a full treatise). all such change, concurrent with experience and knowledge progression, help to test (realistically and efficiently) and adapt the low-flow ecological reserve being implemented. secondly, ‘reframing’ of planning inputs in the crocodile river sam system occurs when tpc monitoring protocols are reflected upon, often as a consequence of scarce resources (people, time and costs). for example, the ecological water resource monitoring framework (ewrmf; kleynhans et al. 2009) was modified into a pragmatic approach for testing effectiveness of the ecological reserve river flows. four levels of monitoring intensity were envisaged: frequent but cheaper or feasible monitoring occurs earlier on prior to higher level and increasingly more costly or intensive monitoring, which is applied based on relevant tpc exceedances. ultimately, this sequence of monitoring aids in more efficient management decision-making about how, where and when to intervene, or not. owing to the logistical burden and ease of sampling encountered, the ewrmf was never implemented in its original form; however, its underlying principles helped to tailor a new approach currently in use. one of the sites, 50 km upstream of the tenbosch gauging weir, is now used for full ecological benchmarking where the full suite of aquatic variables and habitat types are sampled over a 100-m stretch of the crocodile river. replication is achieved at five additional sites solely by macro-invertebrate sampling using the sass-5 method (dickens & graham 2002). this approach, although in its infancy and despite there are problems associated with pseudo-replication, is showing promise. it more explicitly determines the drivers of change in the aquatic environment (bain & stevenson 1999; kleynhans & louw 2009) and dynamics of the response indicators of change to be measured (clarke & warwick 2001; jewitt et al. 1998; weeks et al. 1996), which promotes decision-making. future ‘reframing’ exercises might include reconsideration of the modelling techniques employed for successful delivery of ecological reserve, specifically during the critical dry season period (august–september) when the system is most vulnerable to low flows and a high relative water demand from irrigation. experimentation with the timing of kwena dam water releases is another option, for instance, in conjunction with the international flow requirements into mozambique; however, irrigation farmers most likely will need to coordinate their irrigation activities using water more efficiently. meanwhile, new endeavours are emerging in close cooperation with the irrigation sector to improve the assurance of water supply to both consumptive uses and the ecological reserve, with the 7–10 day lag being realised at the lower end of the river system in response to kwena dam water releases. notably, during the severity of the 2015–2016 drought, the crmib disaggregated the river along the boundary of the knp into three irrigation reaches, with alternating abstractions to ensure continuity of flow, albeit very low towards the komati river confluence. this, in turn, allowed the knp staff to broaden their understanding of environmental water allocation issues in a previously contested section of river. furthermore, there was cooperation in initiating research, for example, related to the unaccounted-for water transmission losses in the river, and a socio-economic review of drought impacts on neighbouring communities dependent on commercial agriculture. such ongoing activities enable (conscious) change, which is vital for harmonious implementation of the ecological reserve in the crocodile river. triple-loop learning (review and transformation) triple-loop learning is about changing norms and values whilst transforming existing governance arrangements, and we identify four key experiences in crocodile river sam. firstly, there is the development and instigation of the rrs and its communication and feedback arrangements, uniquely involving stakeholders collaboratively (figure 4). secondly, there are attempts to broaden the stakeholder networking system, that is, expanding participation and representation in both the crococ (with local municipalities and irrigation boards situated along the crocodile river tributaries) and the more widely constituted crocodile river forum (especially connecting with key sectors in broader freshwater management, for example, mining). thirdly, stakeholder roles and tasks have been identified and agreed upon, which increases the efficiency and effectiveness of ecological reserve management (appendix 3). importantly, such a change in the process of dividing up resources and responsibilities requires (and builds) trust and cooperation between different stakeholders. for instance, there is an informal understanding between the inkomati-usuthu cma, knp, crmib and other stakeholders of the crococ where knp monitors river flows against the ecological reserve benchmarks and reports on ecological outcomes (mcloughlin et al. 2011a, petersen et al. 2019) (figure 5). fourthly, flexible networking between crococ stakeholders is encouraged. for example, an informal fieldtrip was commenced by stakeholders at the kwena dam, who then visited a sugar cane farm (irrigation pump station) and a knp river reach (ecological monitoring site). the fieldtrip allowed fruitful discussions amongst stakeholders in a relaxed atmosphere, for instance, on ecological monitoring methods, water extraction needs and other key aspects of crocodile river management. furthermore, during the year 2015 in the early part of the severe drought, a smaller informal working group was established to explore various low-flow abstraction options for implementation. more recently, there has been concerted collaborative research with a recently established local university on river aquatic connectivity issues, and also problems concerning exotic aquatic species. participation and collaboration in such informal activities help to foster key social learning potentials in adaptive management of the river system (cf. pahl-wostl 2009). figure 5: governance, information, and feedback arrangements and agreed responsibilities in managing the crocodile river ecological reserve between kruger national park, inkomati-usuthu catchment management agency and stakeholders of the crocodile river operations committee (e.g. crocodile river major irrigation board). future triple-loop learning in crocodile river sam should include a review of values and high-level objectives. there are key questions to ask, for example, if the ecological reserve is being implemented as planned, then what undue impacts, if any, will this have on sugar cane farming outside of knp, and therefore, on the region’s economic outcomes? resolving such issues will require innovative thinking; for example, the irrigation farmers are finding ways to use water more efficiently. in addition, the decision made to maintain the ‘c’ ecological category river should be reviewed from time to time, and perhaps, also the broader knp biodiversity objectives based on stakeholder values. by and large, the balance between social, economic and ecological objectives of the managed river system needs to come up for reflection, as knowledge advances along with values change. discussion the crocodile river sam system linked to the knp is setting an important precedent for implementation of ecological reserve in south africa via learning in adaptive management. in order to explain this reasonable success with the ecological reserve, the authors weigh three factors heavily in this regard. firstly, there is evidence of preconditions (e.g. transparency) for social learning, as sensed by the stakeholders themselves; thus, there is valid potential for this learning. the crococ forum has had a trusted and recognised coordinator and is itself a developing ‘community-of-practice’ exhibiting fruitful stakeholder interactions, a situation likely fostering capacities (e.g. information sharing) for sustained social learning. secondly, capacity for adaptive and transformative learning is underpinned by ongoing and overlapping feedbacks supporting adaptive assessment and reflection (biggs et al. 2011b), which help to lubricate the cogs of single-, doubleand triple-loop learning. at this point in the evolution of the adaptive management system, we do not suggest that these feedbacks and ensuing learnings are part of any rigorous predetermined systematic plan. they likely occur as opportune ‘real world happenings’ (cf. biggs et al. 2017), for instance, key motivated or informed staff, funding timing, stakeholder attributes and responses to crises, and outside research interests or collaborations. thirdly, champions exist and are the glue that binds the adaptive management system together without which it is more likely to falter (mcloughlin & thoms 2015; nel & roux 2018). they are enlightened individuals within relevant agencies who are motivated and adaptable, with a grasp of complexity and a desire to learn and integrate knowledge (roux, murray & hill 2010; stirzaker, roux & biggs 2011). we now turn to discussing the learning experiences (lessons) emanating from the crocodile river sam system. lower-grade single-loop learning (rapid response system) a large part of this learning occurs because of operations of the rrs in the crocodile river sam, which gets feedbacks working as quickly as possible ideally before river flows decline to unacceptable levels (figure 4). these feedbacks are vital because uncertainties exist; for instance, the hydrological models are built on imperfect data and assumptions; therefore, errors will always exist when predicting river flow behaviour. typically, lower-grade single-loop learning improves understanding on the quantity and timing of the kwena dam water releases for implementing the ecological reserve in conjunction with irrigation and water restrictions imposed. whilst this article is framed around ecological reserve implementation, there is a concomitant process undertaken by the commercial irrigation sector, who themselves proactively monitor river flows at their specific target points, implement restrictions or strategies where necessary, and/or request releases from kwena dam. critically, rrs-related understanding is growing whilst ecological reserve flows are actually being implemented, and this ‘learning-by-doing’ is likely the key factor promoting ability to make timely decisions on river flow operations and the adjustments when needed. in the authors’ view, success of the rrs would not be possible without champions – people who initiate and facilitate significant stakeholder interactions via more informal interactions and operational feedbacks, often using pragmatic communication mechanisms (e.g. via whatsapptm). the champion role has been taken up by the river systems planning and operations manager of the inkomati-usuthu cma and the knp freshwater team, and more recently the crmib themselves who value the ecological reserve as a sustainability metric. by streamlining operations of the rrs and the now quarterly crococ meetings, these champions are arguably the most important catalyst building social learning potentials (e.g. exchange of ideas, trust, cooperation) in crocodile river sam. this point is significant, given typical (formal) structural constraints imposed on ecological reserve practitioners and stakeholders at the catchment-scale in south africa, from national water policy (see pollard & du toit 2011a). overall, the rrs is an operational conduit illustrating how progressive and adaptive implementation of the ecological reserve can be realised in south african rivers. upper-grade single-loop learning (adaptive assessment) the tpc-related auditing process (cf. mcloughlin et al. 2011b) is the prominent instigator of feedbacks for this learning in crocodile river sam; its intention is to support stakeholders in making purposeful decisions related to implementation of ecological reserve. the process involves collating tpc audit reports via the tpc maintenance system where tpcs are tabled if or when they are exceeded, or close to being exceeded (see biggs & rogers 2003). the tpcs are measurable end points ultimately alerting stakeholders to risk of the ‘c’ ecological category river being lost. if so, increased effort is directed towards technicalities (e.g. rrs procedures) for getting expected ecological reserves delivered. to this end, the reporting of the ecological reserve ‘worry levels’ has to be amended from time to time, as mentioned previously. this is to account for new compliance targets, including international flow obligations (the triple-loop related change, see below), and that in itself is an adaptive process. attention often skews towards the ecological reserve operations (rrs), that is, at the expense of the upper-grade single-loop learning feedbacks. however, implementing the ecological reserve is not an end in itself but a recommended flow regime (hypothesis) to maintain the ‘c’ ecological category river based on the best available knowledge. this is then one constructive critique of crocodile river sam, because monitoring outcomes are needed to test the generally precautionary estimates of required flows – against the real ecological categories which result – thus, enhancing understanding of the relationship between river flow patterns and ecological conditions (mcloughlin et al. 2011a). a greater appreciation of such feedbacks and outcomes can be strengthened by improved group understanding of system complexity (vis-à-vis low-relative score – figure 3c), in particular appreciation of the range of interwoven interests, influences and needs. such a favourable outcome can be achieved if stakeholders co-construct conceptual ‘systemic’ diagrams of the managed system (cf. biggs et al. 2008; pollard, biggs & du toit 2014) and/or with mental model analyses (see adams et al. 2018; biggs et al. 2011c). importantly, learning processes in crocodile river sam need to accommodate changing circumstances across water-use sectors, and factor any change into future medium to long-term river operations and decision-making. overall, single-loop learning (lowerand upper-grade) is the basis for double-loop (reframing; changing practice) into triple-loop (reviewing; transforming governance) learning in the crocodile river sam (mcloughlin & thoms 2015). notably, in order to avoid the trap of ‘learning for the sake of learning’ (fabricius & cundill 2014), there must not be a disproportionate amount of time spent on doubleand/or triple-loop learning (e.g. biggs et al. 2011b; pollard et al. 2011a; varady et al. 2013), that is, at the expense of the ‘actual doing’ (single-loop learning). hence, within environmental water programmes, finding an appropriate balance between modes of single-, doubleand triple-loop learning is encouraged (mcloughlin & thoms 2015). double-loop learning (adaptive reflection) adaptive reflection-related feedbacks in crocodile river sam foster a more deliberate and lengthy evaluation (cf. biggs et al. 2011b) concerning the ecological reserve, which is linked to biodiversity objectives and the explicit measurable end-points or tpcs. construction of tpcs uses existing knowledge (formal scientific and local/tacit), however, incomplete this is (biggs et al. 2011a; mcloughlin et al. 2011b) based on stakeholder values. following principles of requisite simplicity (see stirzaker et al. 2010) – necessitated when managing in complex and uncertain contexts – means that within the inductive approach to tpc development, stakeholders never attempt to finalise indicators and their associated thresholds. hence tpcs are often developed as ‘first generation’ tpcs (biggs et al. 2011a), and open to challenge and refinement as new knowledge becomes available and/or stakeholder values change. for example, the reframed knp geomorphic tpcs (mcloughlin et al. 2011b) are cost effective, feasible and relevant to low-flow ecological reserve implementation and ongoing management (though still to be deployed). generally, reframing should not occur too often; otherwise change at the expense of the ‘actual doing’ may well overwhelm management (fabricius & cundill 2014). another double-loop learning experience was facilitated when adaptive reflection processes led crococ stakeholders to identify and then acknowledge an approximate 10% inaccuracy in the ecological reserve benchmark calculated each week (s. mallory [iwr water resources pty (ltd)] pers. comm., 13 august 2013). effective communication and the build-up of trust amongst stakeholders enabled new thinking to emerge, and this translated into a decision to allow the ecological reserve to fluctuate within an ‘envelope of acceptability’ – rather than an exact amount – defined by the ‘low-worry’ to ‘medium-worry’ rrs levels (figure 4). still, if gauged river flow ever breaches the ‘medium-worry’ level, then management urgency is definitely stepped up. such change in practice – emanating from this more informal stakeholder network – is important, although in direct contrast to more rigid top-down constraints imposed by legislation on cmas and stakeholders (e.g. expecting exact or legally defendable compliances with gazetted ecological reserves), making ecological reserves difficult to achieve in practice (pollard & du toit 2011a). it is within these informal stakeholder networks (figure 4) where innovation and learnings are supported, albeit under the necessary stabilising context of the formal water policy (cf. pahl-wostl 2015). overall, broadening social learning potential in the crococ forum (e.g. awareness of interdependence, information sharing and joint problem solving) helps to foster the reframing requirements of double-loop learning, which is important for resolving complex management problems. here, the stakeholder network proceeds to deal with specific problems and is open to experimentation involving different approaches (pahl-wostl 2009). such innovation and flexible management are crucial factors for progressive implementation of the ecological reserve in the crocodile river. triple-loop learning (review and transformation) triple-loop learning encompasses a review of values, norms and governance arrangements associated with ecological reserve management; it is transformative change over longer time frames. a key example is the crocodile river’s rrs, with already a decade of deployment and further refinement. the rrs reinvigorated (practical) ways for negotiations to occur in a highly contested and water-stressed catchment situation with a history of ecological reserve noncompliance (pollard & du toit 2011b; pollard et al. 2011b; riddell & jewitt 2010; riddell et al. 2014). with collaborative decision-making when river flow declines below different ‘worry-levels’ (figure 4), stakeholders are building trust as they cooperate and together explore best options and solutions for management. this was clearly demonstrated during the peak of the severe 2015–2016 drought. crocodile river flows recorded at the tenbosch gauging weir were consistently higher than that was observed previously during the previous significant droughts of 1991–1992 and 1982–1983, despite lower catchment rainfall and higher temperatures occurring in the 2015–2016 period (smit & bond 2020). furthermore, partnerships and interdependent roles have been entrusted to, and between, different stakeholders, for example, the monitoring and ecosystem reporting role of the knp (figure 5) and the dam and water restriction operations of the crmib. these kinds of innovation, and also cooperation, are vital for effectual ecological reserve implementation. the degree of ‘agency’ – extent to which stakeholders as a group act independently, making free choices (cf. pahl-wostl 2015) – is pivotal for adaptive management of environmental water. agency supports progression of transformative (i.e. double into triple loop) learning and innovation (pahl-wostl 2009) amongst practitioners and stakeholders, in conjunction with top-down (national) influences. indeed, there is now significant national government interest in crocodile river sam, with nascent bottom-up learnings expected to help step up countrywide ecological reserve applications (e.s. riddell [south african national parks] pers. comm., 18 december 2019). besides, national-level transformative change will certainly influence sam of the crocodile river, for example, a review of and amendment to the national water act itself including any legislated water resource protection mechanisms. in addition, recent review of the knp’s vision and mission (www.sanparks.org/assets/docs/conservation/park_man/knp/draft-plan.pdf), reflecting a change in values, demands holistic transformation of governance arrangements, for instance, refocusing management on the entire freshwater resource base, especially the current priority values of equity and redress (pollard & du toit 2011b). in the future, a conscious review is required of all constraints and opportunities associated with the crocodile river’s gazetted ecological reserve. altogether, these bottom-up and top-down processes impact the effectiveness of environmental flow planning and implementation, and hence, they are significant in shaping the delivery of ecological reserve in south african rivers. achieving progress with the crocodile river ecological reserve with emergent social learning – progressing through modes of single-, doubleand triple-loop learning – since 2009 and a decade of managing the crocodile river, growing capacity for adaptive and transformative learning helps stakeholders to deliver the ecological reserve. importantly, this learning has been concurrent and in combination with meaningful change happening in the crocodile river system between 2009 and 2019 (see table 2). learning is by no means absolute; it is ongoing and evolving. after the 2013 interviews, the authors have witnessed further potential for crococ social learning. for example, group problem solving particularly concerning the low-flow river abstraction problems and early warning solutions implemented, applicable not only to the irrigation sector but also to the municipal sector now. stakeholders generally communicate well, exchange ideas and share information, which assists them to deal with joint problems. cooperation is growing within a dynamic environment of increasing trust and shared understanding, demonstrated via stakeholder consensus being reached to avoid, at all costs, river flows dipping into the very high (now extreme) rrs ‘worry level’. social learning-related advantages were evident during the severe drought crisis of 2015–2016. future analysis of the qualitative data also collected during the 2013 interviews will be highly beneficial in understanding crococ social learning processes. furthermore, eliciting stakeholder mental models (cf. biggs et al. 2008; lynam et al. 2012) can make explicit the implicit assumptions individuals hold and how they understand a managed system (adams et al. 2018; moon & adams 2016). besides, mental model analyses offer insights into how single-, doubleand triple-loop learning transpires and proceeds over time (cf. biggs et al. 2011c). table 2: timeline of change events (2009–2019) associated with the crocodile river sam system, concurrent with crocodile river operations committee stakeholder learning (single-, doubleand triple-loop) whilst implementing the ecological reserve in the kruger national park. learning experiences from the knp-linked crocodile river sam system thus far have helped to tailor ecological reserve implementation in other perennial rivers traversing knp (e.g. olifants river; biggs et al. 2017; table 2) via translation of the social learning processes. such experiences (and lessons) are applicable to environmental water programmes throughout south africa and, the authors believe, further abroad too. as climate change uncertainties manifest, particularly frequency and intensity of droughts and floods, increased resilience with stakeholder collaboration, learning and ability to adapt and change will become increasingly important for sustainable management and use of rivers, worldwide. conclusion management of environmental water requires ongoing adaptive and transformative learning. capacity for this learning is built progressively over time through quality stakeholder interactions wherein emergent social learning promotes opportune iterations of single-, doubleand triple-loop learning (fabricius & cundill 2014; pahl-wostl 2009). for effective and successful delivery of the ecological reserve in south african rivers, cmas and stakeholders need to engage within processes of continuing social learning. these processes must entail the generation of trust, cooperation, shared understanding and consensus building because this fosters continual rethinking and negotiation of options to solve complex management problems (vis-à-vis ernst 2019; pahl-wostl et al. 2007). this research study shows that execution of this learning relies critically on champions – people who with their enlightenment can motivate stakeholders whilst they coordinate, share and disseminate information and knowledge (stirzaker et al. 2011). movement away from ‘command and control’ management styles is also vital, and in particular the tendency to seek and implement optimal ‘once-off technical’ solutions to problems. emerging capacity for adaptive and transformative learning emboldens cmas and stakeholders to adapt their management and transform their governance. thus, this guides the complex and uncertain process of ecological reserve planning and implementation to achieve agreed objectives. findings in this article support the case for multiple cmas across south african water management areas, overbids for a single (centralised) cma. whilst managing environmental water, it is imperative that practitioners and stakeholders retain their autonomy – utilising their unique set of resources they keep distinguishing interdependent roles, self-organise and pursue opportunities to experiment, learn, innovate and adapt. full delegation of all ecological reserve management-related responsibilities (e.g. national water act: water resource protection mechanisms), to the applicable regional cma and stakeholder base, must, therefore, be considered in south africa. such an arrangement engenders decentralised, polycentric, flexible and more informal governance (folke et al. 2005; gunderson, holling & light 1995), which promotes adaptive and transformative learning in uncertain contexts (pahl-wostl 2009). this learning enables a positive change across the range of ecological, social and economic outcomes that are essential for success in environmental water programmes worldwide. acknowledgements the authors acknowledge the inkomati-usuthu catchment management agency (iucma) for supporting this work. the authors thank stakeholders of the crocodile river operations committee for their active involvement and also for the time taken to complete the interviews. they are also grateful to brian jackson for enabling their involvement within the broader crocodile river sam project led by iucma and for his continued support. the authors also acknowledge tendai sawunyama (iucma) for the support rendered. melissa parsons provided useful inputs for an early version of the research article. the authors also thank two anonymous reviewers for their constructive critique and suggestions on the article, thereby improving it greatly. competing interests the authors declare that they have no competing interests with regard to the writing of this article. authors’ contributions e.s.r., r.m.p. and j.v. were involved extensively with implementation of crocodile river strategic adaptive management (sam) over the study period, working with monitoring, feedback and learning processes related to water resource management and the ecological reserve. c.a.m. conceptualised and compiled the article, and wrote it in conjunction with significant writing inputs from e.s.r. and finalising the manuscript data requirements and editing. r.m.p. assisted with writing components of the results section. c.a.m. designed and conducted the social learning stakeholder interviews, and completed the analysis. e.s.r. and r.m.p. assisted with figure 5, table 2 and compiled online appendix 3. funding information the south african water research commission (associated with sanparks) and iucma provided funding for completing applied components of the work. early conceptualisation of the article occurred during a phd study funded via an australian government phd scholarship (university of new england). a keith and dorothy travelling scholarship allowed for investigation into the social learning component. data availability statement the data provided in this article may be used, however, any raw data cannot be provided as it is protected via rules of the ethics office, research development and integrity division, university of new england. disclaimer the views and opinions expressed in this article are those of the authors, and do not necessarily reflect official policy or position of any affiliated agency of the authors. references adams, v.m., moon, k., álvarez-romero, j.g., bodin, ö., spencer, m. & blackman, d., 2018, ‘using multiple methods to understand the nature of relationships in social networks’, society and natural resources 31(7), 755–772. https://doi.org/10.1080/08941920.2018.1425514 albert, j.s., destouni, g., duke-sylvester, s.m., magurran, a.e., oberdorff, t., reis, r.e. et al., 2021, ‘scientists’ warning to humanity on the freshwater biodiversity crisis’, ambio 50, 85–94. https://doi.org/10.1007/s13280-020-01318-8 allan, c. & stankey, g.h., 2009, ‘synthesis of lessons’, in c. allan & g.h. stankey (eds.), adaptive environmental management: a practitioner’s guide, pp. 341–346, springer, collingwood. anderson, e.a., jackson, s., tharme, r.e., douglas, m., flotemersch, j.e., zwarteveen, m. et al., 2019, ‘understanding rivers and their social relations: a critical step to advance environmental water management’, water 6(6), e1381. https://doi.org/10.1002/wat2.1381 argyris, c. & schon, d., 1978, organizational learning: a theory of action perspective, addison wesley, reading, ma. bailey, a. & pitman, w., 2016, water resources of south africa 2012 (wr2012): executive summary, research report, project no. k5/2143/1, water research commission, pretoria. bain, m.b. & stevenson, n.j., 1999, aquatic habitat assessment: common methods, american fisheries society, bethesda, md. biggs, h., du toit, d., etienne, m., jones, n., leitch, a., lynam, t. et al., 2008, a preliminary explanation of two approaches for documenting ’mental models’ held by stakeholders in the crocodile catchment, south africa, research report, water research commission, pretoria. biggs, h., ferreira, s., freitag-ronaldson, s. & grant-biggs, r., 2011a, ‘taking stock after a decade: does the “thresholds of potential concern” concept need a socio-ecological revamp’, koedoe 53(2), art1002. https://doi.org/10.4102/koedoe.v53i2.1002 biggs, h., breen, c., slotow, r., freitag-ronaldson, s. & hickings, m., 2011b, ‘how assessment and reflection relate to more effective learning in adaptive management’, koedoe 53(2), art. #1001, 13 pages. https://doi.org/10.4102/koedoe.v53i2.1001 biggs, d., abel, n., knight, a.t., leitch, a., langston, a. & ban, n.c., 2011c, ‘the implementation crisis in conservation planning: could mental models help’, conservation letters 4(3), 169–183. https://doi.org/10.1111/j.1755-263x.2011.00170.x biggs, h. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs, h.c. (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59−80, island press, washington, dc. biggs, h.c., clifford-holmes, j.k., freitag, s. & venter, f.j., 2017, ‘cross-scale governance and ecosystem service delivery: a case narrative from the olifants river in north-eastern south africa’, ecosystem services 28(part b), 173–184. https://doi.org/10.1016/j.ecoser.2017.03.008 biggs, r., rhode, c., archibald, s., kunene, l.m., mutanga, s.s., nkuna, n. et al., 2015, ‘strategies for managing complex social-ecological systems in the face of uncertainty: examples from south africa and beyond’, ecology and society 20(1), 52. https://doi.org/10.5751/es-07380-200152 chen, y., colloff, m.j., lukasiewicz, a. & pittock, j., 2020, ‘trickle, not a flood: environmental watering in the murray–darling basin, australia’, marine and freshwater research. https://doi.org/10.1071/mf20172 cilliers, p., 2008, ‘complexity theory as a general framework for sustainability science’, in m. burns & a. weaver (eds.), exploring sustainability science – a southern african perspective, pp. 9–26, african sun media, stellenbosch. clarke, k.r. & warwick, r., 2001, change in marine communities: an approach to statistical analysis and interpretation, 2nd edn., plymouth marine laboratory primer-e ltd, plymouth. cundill, g., cumming, g.s., biggs, d. & fabricius, c., 2011, ‘soft systems thinking and social learning for adaptive management’, conservation biology 26(1), 13–20. https://doi.org/10.1111/j.1523-1739.2011.01755.x daniel, s.e. & walker, g.b., 1996, ‘collaborative learning: improving public deliberations in ecosystem-based management’, environmental impact assessment review 16(2), 71–102. https://doi.org/10.1016/0195-9255(96)00003-0 dickens, c.w.s. & graham, p.m., 2002, ‘the south african scoring system (sass) version 5 rapid bioassessment method for rivers’, african journal of aquatic science 27(1), 1–10. https://doi.org/10.2989/16085914.2002.9626569 dwa (department of water affairs), 2010, comprehensive reserve determination study for selected water resources (rivers, groundwater and wetlands) in the inkomati water management area, mpumalanga, report no 26/8/3/10/12/015, department of water affairs, pretoria. dwaf (department of water affairs and forestry), 1997, white paper on a national water policy for south africa, department of water affairs and forestry, pretoria. dwaf (department of water affairs and forestry), 2007, guidelines for the development of catchment management strategies: towards equity, efficiency and sustainability in water resources management, department of water affairs and forestry, pretoria. dwaf (department of water affairs and forestry), 2008, the development of a real-time decision support system (dss) for the crocodile east river system – inception report. project no wp 9429, clear pure water with water for africa and dormehl technology, on behalf of the directorate: water resource planning systems, pretoria. ernst, a., 2019, ‘review of factors influencing social learning within participatory environmental governance’, ecology and society 24(1), 3. https://doi.org/10.5751/es-10599-240103 fabricius, c. & cundill, g., 2014, ‘learning in adaptive management: insights from published practice’, ecology and society 19(1), 29. https://doi.org/10.5751/es-06263-190129 folke, c., hahn, t., olsson, p. & norberg, j., 2005, ‘adaptive governance of social-ecological systems’, annual review of environment and resources 30, 441–473. https://doi.org/10.1146/annurev.energy.30.050504.144511 garmestani, a.s. & allen, c.r., 2015, ‘adaptive management of social-ecological systems: the path forward’, in c.r. allen & a.s. garmestani (eds.), adaptive management of social-ecological systems, pp. 255–260, springer, dordrecht. gunderson, l., 2015, ‘lessons from adaptive management: obstacles and outcomes’, in c.r. allen & a.s. garmestani (eds.), adaptive management of social-ecological systems, pp. 27–38, springer, dordrecht. gunderson, l.h., holling, c.s. & light, s.s. (eds.), 1995, barriers and bridges to the renewal of ecosystems and institutions, columbia press, new york, ny. hargrove, r., 2002, masterful coaching, rev. edn., jossey-bass/pfeiffer, wiley, hoboken, new jersey. harwood, a., johnson, s., richter, b., locke, a., yu, x. & tickner, d., 2017, listen to the river: lessons from a global review of environmental flow success stories, wwf-uk, woking. huxham, c. & vangen, s., 2000, ‘leadership in the shaping and implementation of collaboration agendas: how things happen in a (not quite) joined-up world’, academy of management journal 43(6), 1154–1175. https://doi.org/10.5465/1556343 iaquinto, b., ison, r. & faggian, r., 2011, ‘creating communities of practice: scoping purposeful design’, journal of knowledge management 15(1), 4–21. https://doi.org/10.1108/13673271111108666 inkomati catchment management agency (icma), 2010, the inkomati catchment management strategy. a first-generation catchment management strategy for the inkomati water management area. ison, r. & watson, d., 2007, ‘illuminating the possibilities for social learning in the management of scotland’s water’, ecology and society 12(1), 21. https://doi.org/10.5751/es-02020-120121 jackson, b., 2015, ‘an adaptive operational water resources management framework for the crocodile river catchment, south africa’, msc thesis, university of kwazulu-natal, pietermaritzburg. jewitt, g., heritage, g., weeks, d., mackenzie, j., van niekerk, a., görgens, a. et al., 1998, modelling abiotic-biotic links in the sabie river, report no 777/1/9, water research commission, pretoria. kingsford, r.t., bino, g. & porter, j.l., 2017a, ‘continental impacts of water development on water birds, contrasting two australian river basins: global implications for sustainable water use’, global change biology 23(11), 4958–4969. https://doi.org/10.1111/gcb.13743 kingsford, r.t., roux, d.j., mcloughlin, c.a., conallin, j. & norris, v., 2017b, ‘strategic adaptive management (sam) of intermittent rivers and ephemeral streams (ires) – juggling temporal and spatial scales, cultures and governance’, in t. datry, n. bonada & a. boulton (eds.), intermittent rivers and ephemeral streams: ecology and management, pp. 535–562, academic press, burlington. kleynhans, c.j. & louw, m.d., 2007, module a: ecoclassification and ecostatus determination in river ecoclassification: manual for ecostatus determination (version 2), report. joint water research commission and department of water affairs and forestry, pretoria. kleynhans, c.j., louw, m.d., birkhead, a., thirion, c., deacon, a. & angliss, m., 2009, on the way to implementation: ecological reserve monitoring, unpublished report. lynam, t., mathevet, r., etienne, m., stone-jovicich, s., leitch, a., jones, n. et al., 2012, ‘waypoints on a journey of discovery: mental models in human-environment interactions’, ecology and society 17(3), 23. https://doi.org/10.5751/es-05118-170323 mackenzie, j.a., van coller, a.l. & rogers, k.h., 1999, rule based modelling for management of riparian systems, report no. 813/1/99, water research commission, pretoria. mcloughlin, c.a., mackenzie, j., rountree, m. & grant, r., 2011a, implementation of strategic adaptive management for freshwater protection under the south african national water policy, research report number 1797/1/11, water research commission, pretoria. mcloughlin, c.a., deacon, a., sithole, h. & gyedu-ababio, t., 2011b, ‘history, rationale, and lessons learned: thresholds of potential concern in kruger national park river adaptive management’, koedoe 53(2), art. #996, 27 pages. https://doi.org/10.4102/koedoe.v53i2.996 mcloughlin, c.a. & thoms, m.c., 2015, ‘integrative learning for practicing adaptive resource management’, ecology and society 20(1), 34. https:/doi.org/10.5751/es-07303-200134 mcloughlin, c.a., thoms, m.c. & parsons, m., 2020, ‘reflexive learning in adaptive management: a case study of environmental water management in the murray darling basin, australia’, river research and applications 36(4), 681–694. https://doi.org/10.1002/rra.3607 millennium ecosystem assessment, 2005, ecosystems and human wellbeing: biodiversity synthesis, world resources institute, washington, dc. moon, k. & adams, v.m., 2016, ‘using quantitative influence diagrams to map natural resource managers’ mental models of invasive species management’, land use policy 50, 341–351. https://doi.org/10.1016/j.landusepol.2015.10.013 mostert, e., pahl-wostl, c., rees, y., searle, b., tabara, d. & tippett, j., 2007, ‘social learning in european river-basin management: barriers and fostering mechanisms from 10 river basins’, ecology and society 12(1), 19. https://doi.org/10.5751/es-01960-120119 muro, m. & jeffrey, p., 2012, ‘time to talk how the structure of dialog processes shapes stakeholder learning in participatory water resources management’, ecology and society 17(1), 3. https://doi.org/10.5751/es-04476-170103 nel, j.l. & roux, d.j., 2018, ‘planning for the protection and management of freshwater ecosystems inside and outside protected areas’, in c.m. finlayson, a.h. arthington & j. pittock (eds.), freshwater ecosystems in protected areas: conservation and management, pp. 204–220, routledge, new york, ny. o’keeffe, j. & rogers, k.h., 2003, ‘heterogeneity and management of the lowveld rivers’, in j.t.d. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 447–468, island press, washington, dc. pahl-wostl, c., 2009, ‘a conceptual framework for analysing adaptive capacity and multi-level learning processes in resource governance regimes’, global environmental change 19(3), 354–365. https://doi.org/10.1016/j.gloenvcha.2009.06.001 pahl-wostl, c., 2015, water governance in the face of global change: from understanding to transformation, springer, cham. pahl-wostl, c., becker, g., knieper, c. & sendzimir, j., 2013, ‘how multilevel societal learning processes facilitate transformative change: a comparative case-study analysis on flood management’, ecology and society 18(4), 58. https://doi.org/10.5751/es-05779-180458 pahl-wostl, c., craps, m., dewulf, a., mostert, e., tabara, d. & tailliue, t., 2007, ‘social learning and water resources management’, ecology and society 12(2), 5. https://doi.org/10.5751/es-02037-120205 pahl-wostl, c. & hare, m., 2004, ‘processes of social learning in integrated resources management’, journal of community and applied psychology 14(3), 193–206. https://doi.org/10.1002/casp.774 pahl-wostl, c., mostert, e. & tàbara, d., 2008, ‘the growing importance of social learning in water resources management and sustainability science’, ecology and society 13(1), 24. https://doi.org/10.5751/es-02352-130124 pahl-wostl, c., nilsson, c., gupta, j. & tockner, k., 2011a, ‘societal learning needed to face the water challenge’, ambio 40, 549–553. https://doi.org/10.1007/s13280-011-0149-1 pahl-wostl, c., jeffrey, p., isendahl, n. & brugnach, m., 2011b, ‘maturing the new water management paradigm: progressing from aspiration to practice’, water resources management 25, 837–856. https://doi.org/10.1007/s11269-010-9729-2 palmer, m.a., lettenmaier, d.p., leroy poff, n., postel, s.l., richter, b. & warner, r., 2009, ‘climate change and river ecosystems: protection and adaptation options’, environmental management 44, 1053–1068. https://doi.org/10.1007/s00267-009-9329-1 petersen, r., riddell, e., sithole, h., shikwambana, p., venter, j. & nzima, n., 2019, status report of the hydrology and aquatic ecosystems in the kruger national park, unpublished internal report 11/2020, scientific services, south african national parks, skukuza. pittock, j., finlayson, m., arthington, a.h., roux, d., matthews, j.h., biggs, h. et al., 2015, ‘managing freshwater, river, wetland and estuarine protected areas’, in g.l. worboys, m. lockwood, a. kothari, s. feary & i. pulsford (eds.), protected area governance and management, pp. 569–608, anu press, canberra. pittock, j. & max finlayson, c., 2011, ‘australia’s murray-darling basin: freshwater ecosystem conservation options in an era of climate change’, marine and freshwater research 62(3), 232–243. https://doi.org/10.1071/mf09319 pollard, s., biggs, h. & du toit, d.r., 2014, ‘a systemic framework for context-based decision making in natural resource management: reflections on an integrative assessment of water and livelihood security outcomes following policy reform in south africa’, ecology and society 19(2), 63. https://doi.org/10.5751/es-06312-190263 pollard, s. & du toit, d., 2007, guidelines for strategic adaptive management: experiences from managing the rivers of the kruger national park, unep/gef project no. gf/2713-03-4679, epp project, iucn, gland. pollard, s. & du toit, d., 2011a, towards the sustainability of freshwater systems in south africa: an exploration of factors that enable or constrain meeting the ecological reserve within the context of integrated water resources management in the catchments of the lowveld, research report no. tt 477/10, water research commission, pretoria. pollard, s. & du toit, d., 2011b, the shared rivers initiative phase 1. towards the sustainability of freshwater systems in south africa: an exploration of factors that enable or constrain meeting the ecological reserve within the context of integrated water resources management in the catchments of the lowveld, report number tt 477/10, water research commission, pretoria. pollard, s., du toit, d. & biggs, h., 2011a, ‘river management under transformation: the emergence of strategic adaptive management of river systems in the kruger national park’, koedoe 53(2), art1011. https://doi.org/10.4102/koedoe.v54i1.1011 pollard, s., mallory, s., riddell, e. & sawunyama, t., 2011b, towards improving the assessment and implementation of the reserve: real-time assessment and implementation of the ecological reserve, final report, report number k8/881/2, water research commission, pretoria. reed, m.s., evely, a.c., cundill, g., fazey, i., glass, j., laing, a. et al., 2010, ‘what is social learning’, ecology and society 15(4), r1. https://doi.org/10.5751/es-03564-1504r01 riddell, e.s. & jewitt, g.p.w., 2010, deliverable no. 1: project k5/1935 report providing updated catchment information and identifying study focus areas. a management tool for the inkomati basin with focus on improved hydrological understanding for risk-based operational water management, unpublished report, school of bioresources engineering and environmental hydrology, university of kwazulu-natal, pietermaritzburg. riddell, e.s., pollard, s.r., mallory, s. & sawunyama, t., 2014, ‘a methodology for historical assessment of compliance with environmental water allocations: lessons from the crocodile (east) river, south africa’, hydrological sciences journal 59(3–4), 1–13. https://doi.org/10.1080/02626667.2013.853123 rist, l., campbell, b.m. & frost, p., 2013, ‘adaptive management: where are we now’, environmental conservation 40(1), 5–18. https://doi.org/10.1017/s0376892912000240 rogers, k.h., 2003, ‘adopting a heterogeneity paradigm: implications for management of protected areas’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 41–58, island press, washington, dc. rogers, k.h. & bestbier, r., 1997, development of a protocol for the definition of the desired state of riverine systems in south africa, department of environmental affairs and tourism, pretoria. rogers, k.h. & biggs, h., 1999, ‘integrating indicators, end points and value systems in the strategic management of the rivers of the kruger national park’, freshwater biology 41(2), 439−451. https://doi.org/10.1046/j.1365-2427.1999.00441.x rogers, k.h. & luton, r., 2011, strategic adaptive management as a framework for implementing integrated water resources management in south africa, report no. kv 245/10, water research commission, pretoria. rogers, k.h., luton, r., biggs, h., biggs, r., blignaut, s., choles, a.g. et al., 2013, ‘fostering complexity thinking in action research for change in social-ecological systems’, ecology and society 18(2), 31. https://doi.org/10.5751/es-05330-180231 rogers, k.h., roux, d. & biggs, h., 2000, ‘challenges for catchment management agencies: lessons from bureaucracies, business and resource management’, water sa 26(4), 505–512. rountree, m.w. & rogers, k.h., 2004, ‘channel pattern changes in the mixed bedrock/alluvial sabie river, south africa: response to and recovery from large infrequent floods’, in d.g. de jalon & p. vizcaino (eds.), proceedings of the fifth international symposium on ecohydraulics, pp. 318−324, international association for hydro-environment engineering and research, madrid. roux, f., diedericks, g., hoffmann, a.c., selepe, m., scherman, p. & riddell, e., 2018, ecostatus of the crocodile river catchment, inkomati river system, phase ii (2017), report prepared for the inkomati-usuthu catchment management agency, mpumalanga tourism and parks agency, nelspruit. roux, d.j. & foxcroft, l.c., 2011, ‘the development and application of strategic adaptive management within south african national parks’, koedoe 53(2), art1049. https://doi.org/10.4102/koedoe.v54i1.1049 roux, d.j., murray, k. & hill, l., 2010, learning strategy framework for the inkomati catchment management agency, water research commission, pretoria. roux, d.j., nel, j.l., cundill, g., o’farrell, p. & fabricius, c., 2017, ‘transdisciplinary research for systemic change: who to learn with, what to learn about and how to learn’, sustainability science 12, 711. https://doi.org/10.1007/s11625-017-0446-0 roux, d.j. & nel, j.l., 2013, ‘freshwater conservation planning in south africa: milestones to date and catalysts for implementation’, water sa 39(1), 15. https://doi.org/10.4314/wsa.v39i1.15 russell, i.s., 1997, monitoring the conservation status and diversity of fish assemblages in the major rivers of the kruger national park, phd thesis, university of the witwatersrand, johannesburg. schoeman, j., allan, c. & finlayson, c.m., 2019, ‘exploring the multiple meanings of adaptive management: a case study of the lachlan catchment in the murray-darling basin’, environmental management 64, 470–482. https://doi.org/10.1007/s00267-019-01203-5 smit, i.p.j. & bond, w.j., 2020, ‘observations on the natural history of a savanna drought’, african journal of range and forage science 37(1), 119–136. https://doi.org/10.2989/10220119.2020.1723695 stankey, g.h., clark, r.n. & bormann, b.t., 2005, adaptive management of natural resources: theory, concepts and management institutions, general technical report pnw-gtr-654, u.s. forest service, pacific northwest research station, portland, or. stirzaker, r., biggs, h., roux, d. & cilliers, p., 2010, ‘requisite simplicities to help negotiate complex problems’, ambio 39, 600–607. https://doi.org/10.1007/s13280-010-0075-7 stirzaker, r.j., roux, d.j. & biggs, h.c., 2011, ‘learning to bridge the gap between adaptive management and organisational culture’, koedoe 53(2), art. #1007, 6 pages. https://doi.org/10.4102/koedoe.v53i2.1007 susskind, l., camacho, a.e. & schenk, t., 2012, ‘a critical assessment of collaborative adaptive management in practice’, journal of applied ecology 49(1), 47–51. https://doi.org/10.1111/j.1365-2664.2011.02070.x thoms, m.c., rose, t. & dyer, f., 2020, ‘riverine landscapes, water resource development and management: a view from down under’, river research and applications 36(4), 505–511. https://doi.org/10.1002/rra.3622 tickner, d., opperman, j.j., bell, r.a., acreman, m., arthington, a.h., bunn, s.e. et al., 2020, ‘bending the curve of global freshwater biodiversity loss: an emergency recovery plan’, bioscience 70(4), 330–342. https://doi.org/10.1093/biosci/biaa002 todd, c. & thirion, c., 2011, macroinvertebrate thresholds of potential concern (tpcs) and ecological specifications (ecospecs) in the sabie and crocodile rivers, kruger national park, unpublished report. van coller, a.l., rogers, k.h. & heritage, g.l., 2000, ‘riparian vegetation-environment relationships: complimentarity of gradients versus patch hierarchy approaches’, journal of vegetation science 11(3), 337−350. https://doi.org/10.2307/3236626 varady, r.g., scott, c.a., wilder, m., morehouse, b., pablos, n.p. & garfin, g.m., 2013, ‘transboundary adaptive management to reduce climate-change vulnerability in the western u.s. – mexico border region’, environmental science & policy 26, 102–112. https://doi.org/10.1016/j.envsci.2012.07.006 vorosmarty, c.j., mcintyre, p.b., gessner, m.o., dudgeon, d., prusevich, a., green, p. et al., 2010, ‘global threats to human water security and river biodiversity’, nature 467, 555–561. https://doi.org/10.1038/nature09440 webb, j.a., watts, r.j., allan, c. & conallin, j.c., 2018, ‘adaptive management of environmental flows’, environmental management 61, 339. https://doi.org/10.1007/s00267-017-0981-6 weeks, d., o’keeffe, j., fourie, a. & davies, b., 1996, a pre-impoundment study of the sabie-sand river system, mpumalanga with special reference to predicted impacts on the kruger national park, vol. 3, report number 294/3/96, water research commission, pretoria. wenger, e., 1998, communities of practice: learning, meaning, and identity, cambridge university press, cambridge. https://doi.org/10.1017/cbo9780511803932 wwap (united nationals world water assessment programme/un-water), 2018, the united nations world water development report 2018: nature-based solutions for water, united nations educational, scientific and cultural organization (unesco), paris. wwf, 2020, living planet report 2020 – bending the curve of biodiversity loss, r.e.a. almond, m. grooten & t. petersen (eds.), world wide fund for nature (wwf), gland. filelist convert a pdf file! page 1 abstract introduction research methods and design results discussion conclusion acknowledgements references appendix 1 about the author(s) albertus s. louw department of mathematical sciences, faculty of natural sciences, stellenbosch university, stellenbosch, south africa sandra macfadyen department of mathematical sciences, faculty of natural sciences, stellenbosch university, stellenbosch, south africa sam ferreira conservation services, south african national parks, skukuza, south africa cang hui department of mathematical sciences, faculty of natural sciences, stellenbosch university, stellenbosch, south africa department of theoretical ecology group, african institute for mathematical sciences, cape town, south africa citation louw, a.s., macfadyen, s., ferreira, s. & hui, c., 2021, ‘elephant population responses to increased density in kruger national park’, koedoe 63(1), a1660. https://doi.org/10.4102/koedoe.v63i1.1660 original research elephant population responses to increased density in kruger national park albertus s. louw, sandra macfadyen, sam ferreira, cang hui received: 17 oct. 2020; accepted: 23 mar. 2021; published: 10 june 2021 copyright: © 2021. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the elephant population in the kruger national park (knp) has been increasing since the cessation of culling in the mid-1990s. this contrasts with recent trends in elephant populations in many parts of africa where poaching continues to decrease numbers. logistic growth theory predicts that increased competition for vital resources when densities increase should serve to constrain population growth, implying a negative density-growth correlation. we tested this prediction using 28 years of elephant census data to investigate how the growth of the knp’s elephant population responds to increasing elephant density from the period 1985 to 2012. we expected a spatially variable population growth pattern in response to the distribution of elephant densities in the park and thus classified the park into zones with low, medium or high long-term (28 years) average, dry-season elephant density. zones were named ‘peripheral’, ‘semi-peripheral’ and ‘core’ zones, respectively, and represent proxies of resource availability to elephant herds. using a stochastic ricker growth model, we tested for the presence of negative density-dependence in population growth in the core versus peripheral zones. in response, we only detected density-dependent growth in the core zone. overall the population grew at 4.1% per year, coupled with local recruitment rates that increased over time, particularly in the peripheral zones. these density-dependent trends support previous observations of homogenisation of elephant distribution and density across the knp landscapes. conservation implications: density-dependent changes to elephant growth rates are scale-dependent (local vs. park level). only core areas with long-term high density show signs of density-dependent growth. overall, the distributions of elephants are homogenising in the knp. conservation authorities should monitor the impact of such homogenisation to landscape heterogeneity. the spatial variation of the negative density-growth correlation, especially between the core and peripheral zones, can be considered when developing effective strategies to manage the knp elephant population. keywords: african elephants loxodonta africana; population growth; density dependence; spatial distributions; conservation management. introduction despite threats of poaching and habitat loss, trends in the numbers of african elephants (loxodonta africana) in southern africa contrast with their declining numbers in the rest of the continent (chase et al. 2016). south africa epitomises these trends with populations increasing in trans-frontier (selier et al. 2016) and fence protected areas (pretorius, garaï & bates 2019). apart from being a key component of south africa’s natural heritage, elephants are ecological engineers or habitat modifiers because of the substantial influence they exert on the habitats they share with other co-occurring species (valeix et al. 2011). that is, elephants can change the structure of vegetation whilst browsing and can therefore become a catalyst for landscape state change (eckhardt, van wilgen & biggs 2000; trollope et al. 1998). as a result, the influence of elephants on ecosystems has stimulated much debate (van aarde, whyte & pimm 1999; van wyk & fairall 1969). the kruger national park (knp) in south africa is a large, protected area where elephant numbers increased from an estimated 10 animals in the early 1900s to over 17 000 elephants in 2015 (ferreira, greaver & simms 2017). historically, authorities linked increasing elephant population size to vegetation impact, resulting in action being taken to control elephant numbers through culling (van aarde, whyte & pimm 1999). changes in conservation philosophy led to the subsequent cessation of elephant culling in 1995, in favour of allowing density-dependent processes to guide population dynamics over the longer term (whyte et al. 1999). in the same period (1997), the systematic removal of many artificial water points in the park was implemented to regulate game distributions by returning surface water availability to a more natural state (pienaar et al. 1997). the authorities also increased efforts to provide more land for elephants by removing fences from neighbouring private protected areas and international trans-frontier parks since 1996 (venter et al. 2008). in general, these management actions aim to let the spatial heterogeneity of resources, such as water and vegetation, drive elephant distributions and regulate the impacts elephants exerted on landscape heterogeneity in lieu of directly regulating population size (owen-smith et al. 2006; van aarde et al. 1999). to date, the effectiveness of changing elephant management strategies to achieve different biodiversity conservation goals remains open to debate. elephant distribution in the knp has become less clumped and more homogeneous as their density has increased (macfadyen et al.2019). this is likely because of elephants’ responses to local resource and space constraints from crowding and consequent movement into relatively lower quality habitats (macfadyen et al. 2019; robson & van aarde 2018) or could be a result of artificial water holes still enabling elephants to utilise areas far from natural water sources (purdon & van aarde 2017). many have argued that this movement into lower quality habitats should result in decreased survival and birth rates and eventually serve to regulate population growth dynamics (owen-smith et al. 2006; robson & van aarde 2018). others have speculated that this has already contributed to lower population growth rates as new management interventions were introduced (ferreira et al. 2017). increased elephant density and the spatial variation of climate and vegetation in the park are expected to jointly affect elephant population growth (robson 2015). on the other hand, as elephants spread further into the landscape, they will also utilise and potentially disturb a much larger part of the vegetation in the park (macfadyen et al. 2019; young, ferreira & van aarde 2009a). this will compromise and even counter management goals of letting landscape heterogeneity drive and regulate the distribution of elephants. population change rate is dependent on population density, whilst the strength of density-growth correlation should vary across landscapes (owen-smith et al. 2006). large vertebrates typically display convex curves of density-dependent population growth patterns (sibly et al. 2005). this means that density-dependent growth only becomes evident once the population is close to the carrying capacity of the habitat (fowler 1981), as is often seen in smaller, fenced reserves. such density dependence originates from the way large vertebrates respond to aggravating resource constraints. for example, when density increases juvenile survival rates typically decrease followed by reduced reproductive outputs, whilst adult survival rates decrease, culminating in reduced population growth (eberhardt 2002). like most large herbivores, elephants choose landscapes based on water, food, comfort and safety (boult et al. 2019; chamaillé-jammes, valeix & fritz 2007; kinahan, pimm & van aarde 2007; purdon & van aarde 2017; wittemyer et al. 2017) and make large seasonal movements (e.g. cook, henley & parrini 2015). during the dry season, the distribution of water is a primary driver of elephant space use (chamaillé-jammes et al. 2007; purdon & van aarde 2017). this need for water often exceeds the selection of food resources in water-scarce ecosystems and thus elephants are restricted to habitats near water sources. as elephant breeding herds normally use the same dry season home ranges from year to year (young, ferreira & van aarde 2009b), we assume the dry season is the key period during which density-dependent processes will activate. as negative density-dependent growth only occurs in areas with high elephant density, we might not be able to detect it at a park-wide scale. as a result, we focus on how local population growth varies spatially in different long-term density zones throughout the park. negative density-dependence refers to the decline in population growth rate in response to increasing population size or density (hixon & johnson 2009), which is typically associated with the effect of intense resource competition from overcrowding. to identify the effects of density-dependent growth, we consider two lines of evidence. firstly, we present a statistical test for negative density-dependent growth described by dennis and taper (1994) that incorporates logistic growth theory (tsoularis & wallace 2002; verhulst 1845). this approach differs from earlier works that identify changes to growth trends by comparing goodness-of-fit of growth models over different time periods (ferreira et al. 2017) or identify factors influencing population growth with regression models (robson 2015). secondly, we quantify trends in the recruitment rates of elephant calves. mainstream large vertebrate models often predict juvenile survival as the first demographic variable to respond to resource constraints (eberhardt 2002; trimble, ferreira & van aarde 2009). the calves counted during the annual dry season censuses, represent those born and also survived between the previous and present dry season. this recruitment serves as a proxy to evaluate early demographic responses to population density (eberhardt 2002; trimble et al. 2009). here we expect a negative density-dependent recruitment rate from density increases and the resulting resource constraints in the dry season. research methods and design study area the knp is situated in the north-eastern part of south africa, bordered by mozambique and zimbabwe to the east and north, respectively (figure 1a). it covers an area of approximately 2 million hectares and is comprised of 35 predominantly savanna and woodland ecosystems (gertenbach 1983) including a number of artificial water points at scattered locations, permanent and seasonal rivers pass through the park and water supply network. the knp has a summer rainfall season (gertenbach 1983), with the south-western part receiving higher annual rainfall (figure 1a; macfadyen et al. 2018). figure 1: (a) the kruger national park, south africa, lies between latitudes 22°19’40” s to 25°31’44” s and longitudes 30°53’18” e to 32°01’59” e. main rivers (in blue) supply water throughout the year. rainfall occurs mainly in summer and varies in the park. the long-term average monthly rainfall for areas is indicated in the figure by a colour sequence from red (dry) to blue (wet). (b) average elephant density observed in each 25 km2 grid-cell throughout the study period (1985–2012). grid-cell colour indicates density of elephants per square kilometre. high density cells are coloured in purple, whilst medium and low-density cells are in green and yellow. the data are from yearly elephant censuses. data collection for this study, we used census data from annual aerial counts conducted in the knp between 1985 and 2012. all censuses were flown during the dry season (july/august/september), when reduced foliage makes elephant spotting easier (van aarde et al. 1999). counts were conducted using a spotting crew and helicopter that systematically flew over 21 designated census blocks to record all elephants sighted. each observed herd is recorded with the geolocation, number of elephants in the herd and group type (i.e. bull group or herd group). bull groups consist of only bull animals whilst herd groups are a mix of cows and bulls of different age classes. in these mixed groups, the number of calves younger than 1 year was also recorded. no information was recorded on elephant deaths, immigrations or emigrations, which may result in an underestimate of birth rates. these data only represent a winter account of elephant distributions. however, water availability and foraging range are limited to a larger degree in these drier winter months (codron et al. 2006; macfadyen et al.2019; thomas, holland & minot 2008), and it is still the best representation of long-term elephant population distributions in the knp. see whyte (2001) for a full description of the census methodology. data preparation all statistical analyses were carried out using r version 3.6.1 (r core team 2019) and associated r packages referenced in the text. firstly, we divided the knp into a grid with 3738 cells of 5 km × 5 km and aggregated the total number of elephants observed in each grid cell for each census year, distinguishing elephants in bull groups, elephants in mixed herd groups and number of calves present. we then calculated: (1) the density of elephants per square kilometre in each grid cell for each year (equation 1) and (2) the proportion of calves per herd born since the previous census using raster (hijmans 2020). we defined local population density as follows: where di(t) is the density in the i-th grid cell at year t; ai the surface area of the i-th grid cell (25 km2); and nij(t) the total number of elephants in the j-th point observation of an elephant group in grid cell i in year t. we then determined the long-term average density of elephants for each grid cell in the knp from 1985 to 2012. that is, for each cell we estimated the average density from annual census records; for cell i, it is the average of di(t) over t. grid cells were then assigned into three zones according to this long-term average density pattern: periphery (0.0–0.5), semi-periphery (0.5–1.0) and core zone (> 1) (units are elephants per square kilometre). we then summarised the growth patterns in grid cells for each zone and produced a map that illustrates the three zones. throughout our investigation, we made the distinction between time periods when the population was constrained by culling (1985–1995) and when culling was stopped (1996–2012). we did this because culling aimed to maintain the knp’s elephant population at a roughly constant size and would thus have obstructed normal population growth rates. growth rates to estimate the overall growth trends of elephant population numbers, we fitted an exponential growth model to the average, park-wide elephant density from 1985 to 2012 using the least-squares method. we fitted separate exponential models for the two periods: (1) when the population was maintained by culling (1985–1995) and (2) when the population was released from culling (1996–2012): where d (t) is the total elephant population density at time t; d0 the initial population density; and r the per-capita annual growth rate. to identify the effect of elephant population size on growth rate, we used a statistical test for density-dependent growth described by dennis and taper (1994). we applied this test first to the time-series of park-wide total population size and then to the time-series of total population size in each of the three zones (core, semi-periphery and periphery). this test assumes growth happens according to a stochastic ricker model, that is, a discrete-time model of logistic population growth, described as follows: where nt is population size at time t; a and b are growth rate parameters describing the density-independent and density-dependent components of growth, respectively and zt is a noise component reflecting added stochasticity in the model. therefore, if b < 0, this indicates a negative density-dependent growth in the population dynamics, suggesting a decline in growth rate in response to the increase of population density (hixon & johnson 2009). we tested whether b is significantly less than zero by assessing whether a model assuming b < 0 fits the observed population growth significantly better than the null model assuming b = 0. we conducted the test separately for the period when the population size was maintained through culling (1985–1995) and the period when the population had no culling (1996–2012). to assess how well growth models described the observed population growth, we compared our estimated model to observed time-series data. to further uncover any potential patterns in the population growth of different zones, we fitted a smooth line to the time series using a smoothed conditional mean function in ggplot2 (wickham 2016). recruitment rates for each observed mixed herd, the number of calves that were born and survived since the previous census year was documented. in this context, we quantified recruitment rate as the ratio of calves to other animals in each group. the subsequent metric represents how productive or fertile cows in each elephant herd and specific grid cell were, as well as how well calves from the previous year survived for a specific year. however, it is worth noting that the calf survival might have been overestimated because of the fact that the census was conducted during the dry season when resources are limited, so some calves could still succumb to resources limitations after the census. we specifically do not refer to birth rate, because we have no data on actual calf births or deaths. to estimate yearly recruitment rate, we calculated the proportion of the herd made up of calves and reported the average for herds in each grid cell as follows: where ri is the average recruitment rate per herd group in the i-th cell, cij the number of calves in the j-th elephant herd in cell i, and hij the number of elephants in the rest of the herd. the number of herd-groups in a grid cell is ni. in this calculation, we excluded bull groups as calves are present only in mixed herds. to check whether recruitment rates differed significantly between grid cells belonging to the three different zones, we used the non-parametric mann–whitney u test. this statistical test is robust for data that is not normally distributed, for example, our data had unequal number of cells in the different zones. furthermore, to reduce skewness in the distribution of recruitment rates, we transformed the recruitment rate by its cube root. we then fitted a smoothed conditional mean to the time series (wickham 2016) to display recruitment rate trends in the core, semi-peripheral and peripheral zones. a non-parametric mann-kendall test was then conducted to test for the presence of a significant trend in elephant recruitment over the study period (between 1985 and 2012). we used this test because elephant recruitment rate was not normally distributed according to a shapiro–wilk test. results growth and density-dependence during the period elephants were culled (1985–1995), the population was maintained at approximately 0.36 elephants per square kilometre (figure 2; appendix 1 table 1-a1; r2 = 0.34, f(1,9) = 6.16, r = 0.006, p = 0.035). in the period after 1995, growth can be described by an exponential model with per capita annual growth rate of 4.1% (figure 2; appendix 1 table 1-a1; r2 = 0.95, f(1,15) = 278.4, r = 0.041, p < 0.001). figure 2: time series of park-wide elephant density in elephants per square kilometre, as obtained from annual census (circles). lines represent exponential growth models (eqn 2) fitted to the census time series for the period with culling (1985–1995) in blue and without culling (1996–2012) in red. during the culling period (1985–1995), the overall population and the zoned populations showed significant negative density-dependent growth (appendix 1 table 2-a1, appendix 1 figure 1-a1). for the post-culling period, no negative density-dependent growth was observed in the overall population (figure 3a). that is, b was > 0 in the stochastic ricker growth model (eqn 3) and thus we could not reject the null hypothesis (b = 0) (see appendix 1 table 2-a1). however, we did find a significant (α = 0.05) negative density-dependent component for elephant growth in the core zone (b = −1.4 × 10−4). in the semi-peripheral and peripheral zones, the bootstrap confidence intervals for the parameter b included zero, indicating no density-dependent effects. figure 3: deterministic component of stochastic ricker growth models (eqn 3) fitted to time series of elephant population in the kruger national park (1996–2012). models shown for overall population (a), and separately for the three zones in the park; peripheral (b), semi-peripheral (c) and core (d). test for negative density dependent growth reported significant for the core zone (d). model parameter estimates in appendix 1 (table 2–a1). elephant densities increased in the core zone after culling practices (red line in figure 4a). after 2008, elephant densities in these core areas began to stabilise, whereas in the semi-peripheral and peripheral zones elephant densities sharply increased even after 2008 (figure 4a). figure 4: time series of (a) elephant density per grid cell and (b) recruitment rate. the solid lines represent a smoothed conditional-means fit to the data in ggplot2. filled regions show 95% confidence intervals. grid cells were classified into three zones according to the average elephant density observed in them through the study period. colour distinguishes the zones (grid cells that had a low average elephant density were classified as ‘peripheral’ and are shown in yellow, grid cells with medium average density were ‘semi-peripheral’ and are shown in blue and grid cells with high elephant density were defined as the ‘core zone’ for elephants and are in purple). a cube root transform is used to reduce skewness in the distribution of the recruitment rates. recruitment rate is defined as the proportion of calves in a herd and is reported as an average for each grid cell (eqn 4). recruitment in the peripheral and semi-peripheral zones had significant upward trend, by mann-kendall test for trend (table 4-a1). recruitment rates elephant recruitment rates in the peripheral and semi-peripheral zones had a significant upward trend between 1985 and 2012 (mann-kendall tests for trends, p = 0.017 and p = 0.034, respectively; table 4-a1). recruitment rates in the core zone did not have a significant trend (p = 0.277) but were in general higher than the rates in the semi-peripheral and peripheral zones. recruitment rates in the semi-peripheral zone were also higher than in the peripheral zone (mann–whitney u tests, table 3-a1). over time, however, the trends in recruitment appear to converge and all zones experienced similar recruitment rates by 2012 (figure 4b). discussion scale and density-dependent growth elephant population growth rates in the knp were expected to drop in response to management approaches that enhance habitat variability in the landscape (ferreira et al. 2017). over time, these reduced growth rates would manifest as reduced birth rates and increased mortality because of increased spatial variability of resources and thus increased competition for space and resources as population density increases (van aarde et al. 1999). we observed, however, that the knp’s population has been growing exponentially at an annual rate of 4.1% since culling stopped in 1995. these results are similar to those reported by ferreira et al. (2017) but lower than the intrinsic growth rate reported during the time of culling (whyte 2001). african elephant populations, when left undisturbed, typically experience exponential growth (slotow et al. 2006). with persistent poaching, however, elephant population growth can be disrupted (chase et al. 2016) to such an extent that many places across the continent now have only 25% of expected elephant numbers (robson et al. 2017). different management strategies can also influence natural population dynamics and can disrupt natural density-dependent processes in protected areas. for instance, no density-dependent population growth has been recorded in elephants in the addo elephant national park (aenp) (gough & kerley 2006) as various land acquisitions have increased the area available to elephants (kerley, boshoff & knight 2002) and thereby potentially delayed the effect of resource constraints associated with increasing densities. during the culling era in the knp, authorities aimed to maintain the elephant population size to around 7500 animals (van aarde et al. 1999). in practice, to achieve this annual culling, quotas were set based on the number of elephants counted the previous year. in other words, years with high elephant numbers had high culling quotas set the next year (whyte 2001). in some ways, management’s response was itself a density-dependent response. it is thus not surprising that we observed negative density-dependent population growth during the period when the elephant population was maintained at a target population size through regular culling (appendix 1 figure 1-a1; table 2-a1). however, this ‘density-dependent’ population growth during the culling periods was not in response to resource constraints and once the population was released from culling the population growth rate switched to an exponential pattern (figure 2). this is similar to observations elsewhere in south africa (slotow et al. 2006). exponential growth models describe the growth of populations in the absence of density-dependent growth constraints. as the elephant population growth was well described by an exponential growth model, it suggests that the knp elephant population has not yet reached the point where resource constraints curtail the overall population growth in the park. that is, the population has not yet reached the total carrying capacity of the park (van dyken & zhang 2019). for large vertebrates such as elephants, the effect of density on growth rates might only play out and become notable when population abundances have reached levels close to or above the carrying capacity (fowler 1981; sibly et al. 2005). furthermore, in a large protected area, the total carrying capacity of the habitat becomes difficult to establish as it is then a function of local population growth parameters, habitat heterogeneity and the strength of dispersal (van dyken & zhang 2019). in such cases, the population dynamics and presence of localised density dependence might differ from the dynamics observed for the population as a whole. the effect of density dependence can therefore be scale-dependent. we failed to observe density-dependent growth at the scale of the entire park (i.e. the overall population growth; figure 2 and figure 3a), but we detected density-dependence effects on elephant growth at local scales (i.e. when growth was considered as spatially explicit across 25 km2 grid-cells). in particular, in the core zone we detected significant density-dependent growth (figure 3d). these significant local-scale density-dependent growths were masked in the overall park-scale growth because of the spatial variation in local density-dependent growth (figure 3b, 3c). the fact that density dependence was spatially variable suggests that the carrying capacity also varied across the landscape. previous predictions that variations in local water and food availability, thus local carrying capacity, should influence the distribution and dynamics of elephants in the landscape are thus indirectly supported by our findings (harris et al. 2008; owen-smith et al. 2006; smit & ferreira 2010; young et al. 2009a). furthermore, the density-dependent growth in the core zone aligns well with the expected convex shape suggested by sibly et al. (2005). that is, density-dependent effects are only realised at high density levels where elephants are likely to experience increased intraspecific competition for space and food resources (macfadyen et al. 2019). elephants respond to variation in resources and thus move extensively across the knp (cook et al. 2015). elephants choosing favoured habitats in the core zone could therefore result in increased local intraspecific competition for space/territory (i.e. between elephant herds), which will limit the number of groups that can be present in an area (macfadyen et al. 2019). this is known to occur when one herd dominates over another (wittemyer & getz 2007; wittemyer et al. 2007), thereby displacing subordinate herds to suboptimal habitats. such social dynamics could also accelerate herds’ response to leave an area before resource conditions deteriorate to the point where a heuristic learning ‘good-stay, bad-disperse’ dispersal strategy is employed (hui et al. 2012). this might explain elephant numbers rapidly increasing in the peripheral and semi-peripheral zones at roughly the same time (2008/2009) that elephant growth in the core zone slowed down. we speculate that the peripheral and semi-peripheral zones in our study represent suboptimal habitats, as those areas had lower dry-season herd occurrences over the long term. we did not directly evaluate habitat suitability for elephants in the different density areas, as performed in previous studies (harris et al. 2008; young et al. 2009a). the population distribution that results from these responses is a less clumped and more homogenised pattern as elephant densities increase (macfadyen et al. 2019; young et al. 2009a). how elephant herds with calves distribute in the landscape may also be explained by the aforementioned social dynamics and the distribution of resources. we observed that per-herd recruitment rates (the ratio of new calves to other herd members) were highest in the core zone (figure 4b; wilcoxon rank sum test table 3-a1). we expect that herds prefer to keep young calves in areas with good quality habitat, that is, close to water sources and lush vegetation, as noted elsewhere in the region (harris et al. 2008). yet, recruitment in peripheral and semi-peripheral zones had an upward trend (mann-kendall tests: table 4-a1). recruitment patterns that initially differed in the core and peripheral zones, thus began to converge around 2012 (figure 4b). this most likely reflects how herd groups with calves moved into suboptimal habitat as intraspecific competition for resources and space intensified (macfadyen et al. 2019). recruitment rates also fluctuated over time, a pattern expected given the association of first year survival with environmental conditions, for example, rainfall (trimble et al.2009). indeed, robson (2015) found this to be true as variation in the proportion of elephants less than 1 year old was related to changes in vegetation productivity (ndvi), which itself was driven by rainfall in the knp. for example, the recruitment rates increased sharply after 2000, which was also a year of exceptional rainfall in the park (macfadyen et al. 2018) (figure 4b). years with high rainfall might also have an effect on recruitment in following years and future work could investigate the dynamic relationship between recruitment and rainfall. the spatial pattern of herds with calves raises the question of how fecundity and young calf survival respond to resource constraints in areas with suboptimal habitat. owen-smith et al. (2006) suggested that variation in resource availability can put pressure on herds in areas where resources are more limiting. this can lead to reduced population growth through elevated mortality or hampered recruitment rates. african elephants may respond to resource constraints by reducing birth rates (trimble et al. 2009). however, we observed that over time the proportion of calves in herds increased in semi-peripheral and peripheral zones. this suggests that conditions and resource availability in lower quality landscapes have yet to hamper fecundity and calf survival, as was predicted by trimble et al. (2009). purdon and van aarde (2017) have suggested that artificial water holes could have influenced the movements of elephants in the park and possibly reduced the effect that natural resource variation had on the elephant distribution. the effects of water holes on the population dynamics of elephants should thus be investigated in future studies. an alternative theory is that increasing density does not reduce the survival of calves until only after they are weaned (young & van aarde 2010) at 3–4 years of age (lee & moss 1986). if this is the case, calves may be most vulnerable to resource constraints shortly after weaning (young & van aarde 2010). comparative results between several populations elsewhere in africa highlighted that with increased daily walking distances and higher density, mortality of weaned calves does increase (young & van aarde 2010). our census dataset did not report weaned calf mortalities, so we could not test this prediction. in addition to the age-specific fecundity and survival data, we acknowledge the limitations of the census data as winter snapshots of elephant distributions and densities. these annual surveys (whyte 2001) do not account for how elephants may move around throughout the year in response to seasonal changes of resources, an aspect well recorded using collared animals in the knp (loarie, van aarde & pimm 2009) and elsewhere (cook et al. 2015; roever, van aarde & leggett 2012). all results reflect a dry-season snapshot of elephant distribution and abundance patterns. however, these dry-season months are expected to limit water availability and foraging range so it may also reflect to some degree territorial distributions (codron et al. 2006; macfadyen et al. 2019; thomas et al. 2008). we report our findings as the long-term changes in dry-season elephant distributions and offer insights into how elephants are responding to changing conditions over time, as illustrated in previous studies using the same dataset (e.g. macfadyen et al. 2019; young et al. 2009a, 2009b). as restoring resource gradients are expected to have park-wide effects on elephant responses (venter et al. 2008), thus a key management approach is to focus on approaches that induce spatial variations in resources that affect how elephants use landscapes at scales within the core, semi-peripheral and peripheral zones. these include, restoring food, water and comfort (e.g. shade) resource gradients (owen-smith et al. 2006; purdon & van aarde 2017), as well as safety or danger gradients (douglas-hamilton, krink & vollrath 2005; goldenberg, douglas-hamilton & wittemyer 2018; ihwagi et al. 2018). as discussed in carruthers (1995), a key element of how intensely elephants use a landscape also includes human disturbance as predicted from the theories such as ‘landscape of fear’ (cromsigt et al. 2013). adding to this, incorporating movement and migration patterns may help further elucidate the role of local versus regional habitat constraints on the spatial distribution and recruitment dynamics of the elephant herds in the knp. conclusion although density dependence in elephant population growth was realised in the core zone, it coincided with the increase of elephants in the peripheral zones. this pattern could lead to a situation where elephants are more homogeneously distributed throughout the park, as also mentioned in previous studies (e.g. macfadyen et al. 2019). a homogenised distribution of elephants in response to density-dependent factors is in direct contrast with a conservation paradigm that promotes landscape heterogeneity as an important regulator of biodiversity maintenance (katayama et al. 2014). our results, however, only present the population dynamics up to 2012 and could be extended to include recent census data (conducted at a biennial pace, 2015, 2017 and 2019). our analyses of annual census data highlighted that exponential population growth typifies elephants in the knp at a park-wide scale after authorities stopped culling up to 2012. per capita recruitment rate of calves less than 1 year old into the population varied over the study period but was higher in the core areas. at the park-wide scale, the expected pattern of negative density-dependent growth did not occur. we, however, detected significant negative density-dependence in elephant population growth in the core zone. the mechanism of this density dependence can be demographic responses of fecundity and survival rates and/or movements of elephants to the local density. the spatially varied growth patterns and the apparent change in density trends after 2008, suggest a more pronounced future density-dependent response by elephants to local resource constraints. such responses reflect a gradual spread of elephants into less optimal habitats during the dry season and are likely to have consequences for other species in the park. acknowledgements competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions a.s.l., s.m. and c.h. developed the ideas, a.s.l. ran analyses and wrote the article under the guidance of c.h., s.m. and s.f.a.s.l., s.m. and c.h. and s.f. helped improve the final manuscript. ethical considerations this article followed all ethical standards for research without direct contact with human or animal subjects. funding information we are grateful for the financial support received from the following sources: department of mathematical sciences bursary (stellenbosch university); nrf grant holder-linked bursary (nrf 89967). data availability the authors confirm that the data supporting the findings of this study are available within the article. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references boult, v.l., sibly, r.m., quaife, t., fishlock, v., moss, c. & lee, p.c., 2019, ‘modelling large herbivore movement decisions: beyond food availability as a predictor of ranging patterns’, african journal of ecology 57(1), 10–19. https://doi.org/10.1111/aje.12553 carruthers, j., 1995, the kruger national park: a social and political history, university of natal press, pietermaritzburg. chamaillé-jammes, s., valeix, m. & fritz, h., 2007, ‘managing heterogeneity in elephant distribution: interactions between elephant population density and surface-water availability’, journal of applied ecology 44(3), 625–633. https://doi.org/10.1111/j.1365-2664.2007.01300.x chase, m.j., schlossberg, s., griffin, c.r., bouché, p.j., djene, s.w., elkan, p.w., et al., 2016, ‘continent-wide survey reveals massive decline in african savannah elephants’, peerj 4, e2354. https://doi.org/10.7717/peerj.2354 codron, j., lee-thorp, j.a., sponheimer, m., codron, d., grant, r.c., de ruiter, d.j., 2006, ‘elephant (loxodonta africana) diets in kruger national park, south africa: spatial and landscape differences’, journal of mammalogy 87(1), 27–34. https://doi.org/10.1644/05-mamm-a-017r1.1 cook, r.m., henley, m.d. & parrini, f., 2015, ‘elephant movement patterns in relation to human inhabitants in and around the great limpopo transfrontier park’, koedoe 57(1), 1–7. https://doi.org/10.4102/koedoe.v57i1.1298 cromsigt, j.p., kuijper, d.p., adam, m., beschta, r.l., churski, m., eycott, a. et al., 2013, ‘hunting for fear: innovating management of human–wildlife conflicts’, journal of applied ecology 50(3), 544–549. https://doi.org/10.1111/1365-2664.12076 dennis, b. & taper, m.l., 1994, ‘density dependence in time series observations of natural populations: estimation and testing’, ecological monographs 64(2), 205–224. https://doi.org/10.2307/2937041 douglas-hamilton, i., krink, t. & vollrath, f., 2005, ‘movements and corridors of african elephants in relation to protected areas’, naturwissenschaften 92(4), 158–163. https://doi.org/10.1007/s00114-004-0606-9 eberhardt, l.l., 2002, ‘a paradigm for population analysis of long-lived vertebrates’, ecology 83(10), 2841–2854. https://doi.org/10.1890/0012-9658(2002)083[2841:apfpao]2.0.co;2 eckhardt, h.c., van wilgen, b.w. & biggs, h.c., 2000, ‘trends in woody vegetation cover in the kruger national park, south africa, between 1940 and 1998’, african journal of ecology 38(2), 108–115. https://doi.org/10.1046/j.1365-2028.2000.00217.x ferreira, s.m., greaver, c. & simms, c., 2017, ‘elephant population growth in kruger national park, south africa, under a landscape management approach’, koedoe 59(1), 1–6. https://doi.org/10.4102/koedoe.v59i1.1427 fowler, c.w., 1981, ‘density dependence as related to life history strategy’, ecology 62, 602–610. https://doi.org/10.2307/1937727 gertenbach, w.p.d., 1983, ‘landscapes of the kruger national park’, koedoe 26(1), 9–121. https://doi.org/10.4102/koedoe.v26i1.591 goldenberg, s.z., douglas-hamilton, i. & wittemyer, g., 2018, ‘inter-generational change in african elephant range use is associated with poaching risk, primary productivity and adult mortality’, proceedings of the royal society b: biological sciences 285(1879), 20180286. https://doi.org/10.1098/rspb.2018.0286 gough, k.f. & kerley, g.i., 2006, ‘demography and population dynamics in the elephants loxodonta africana of addo elephant national park, south africa: is there evidence of density-dependent regulation?’, oryx 40(4), 434–441. https://doi.org/10.1017/s0030605306001189 harris, g.m., russell, g.j., van aarde, r.i. & pimm, s.l., 2008, ‘rules of habitat use by elephants loxodonta africana in southern africa: insights for regional management’, oryx 42(1), 66–75. https://doi.org/10.1017/s0030605308000483 hijmans, r.j., 2020, raster: geographic data analysis and modelling, version 3.3-13, computer software, viewed 27 february 2021, from https://cran.r-project.org/package=raster. hixon, m.a. & johnson, d.w., 2009, ‘density dependence and independence’, encyclopedia of life sciences (els), john wiley & sons, chichester. https://doi.org/10.1002/9780470015902.a0021219 hui, c., roura-pascual, n., brotons, l., robinson, r.a. & evans, k.l., 2012, ‘flexible dispersal strategies in native and non-native ranges: environmental quality and the ‘good–stay, bad–disperse’ rule’, ecography 35(11), 1024–1032. https://doi.org/10.1111/j.1600-0587.2012.07697.x ihwagi, f.w., thouless, c., wang, t., skidmore, a.k., omondi, p. & douglas-hamilton, i., 2018, ‘night-day speed ratio of elephants as indicator of poaching levels’, ecological indicators 84, 38–44. https://doi.org/10.1016/j.ecolind.2017.08.039 katayama, n., amano, t., naoe, s., yamakita, t., komatsu, i., takagawa, s.i. et al., 2014, ‘landscape heterogeneity–biodiversity relationship: effect of range size’, plos one 9(3), e93359. https://doi.org/10.1371/journal.pone.0093359 kerley, g., boshoff, a. & knight, m.h., 2002, ‘the greater addo national park, south africa: biodiversity conservation as the basis for a healthy ecosystem and human development opportunities’, in d.j. rapport, b.l. lasley, d.e. rolston, n.o. nielsen, c.o. qualset & a.b damania (eds.), managing for healthy ecosystems, p. 16, crc press, boca raton, fl. kinahan, a.a., pimm, s.l. & van aarde, r.j., 2007, ‘ambient temperature as a determinant of landscape use in the savanna elephant, loxodonta africana’, journal of thermal biology 32(1), 47–58. https://doi.org/10.1016/j.jtherbio.2006.09.002 lee, p.c. & moss, c.j., 1986, ‘early maternal investment in male and female african elephant calves’, behavioral ecology and sociobiology 18(5), 353–361. https://doi.org/10.1007/bf00299666 loarie, s.r., van aarde, r.j. & pimm, s.l., 2009, ‘elephant seasonal vegetation preferences across dry and wet savannas’, biological conservation 142(12), 3099–3107. https://doi.org/10.1016/j.biocon.2009.08.021 macfadyen, s., hui, c., verburg, p.h. & van teeffelen, a.j.a., 2019, ‘spatiotemporal distribution dynamics of elephants in response to density, rainfall, rivers and fire in kruger national park, south africa’, diversity and distributions 25(6), 880–894. https://doi.org/10.1111/ddi.12907 macfadyen s., zambatis n., van teeffelen a.j.a. & hui, c., 2018, ‘long-term rainfall regression surfaces for the kruger national park, south africa: a spatio-temporal review of patterns from 1981 to 2015’, international journal of climatology 38(5), 2506–2519. https://doi.org/10.1002/joc.5394 owen-smith, n., kerley, g.i.h., page, b., slotow, r. & van aarde, r.j., 2006, ‘a scientific perspective on the management of elephants in the kruger national park and elsewhere’, south african journal of science 102(9–10), 389–394. pienaar, d., biggs, h., deacon, a., gertenbach, w., joubert, s., nel, f. et al., 1997, a revised water-distribution policy for biodiversity maintenance in the kruger national park, internal report south african national parks, skukuza. pretorius, y., garaï, m.e. & bates, l.a., 2019, ‘the status of african elephant loxodonta africana populations in south africa’, oryx 53(4), 757–763. https://doi.org/10.1017/s0030605317001454 purdon, a. & van aarde, r.j., 2017, ‘water provisioning in kruger national park alters elephant spatial utilisation patterns’, journal of arid environments 141, 45–51. https://doi.org/10.1016/j.jaridenv.2017.01.014 r core team, 2019, r: a language and environment for statistical computing, computer software, viewed 14 august 2019, from https://www.r-project.org/. robson, a.s., 2015, ‘demographic responses to changes in conservation management: a case study on elephants in the kruger national park’, msc-thesis, university of pretoria, pretoria. robson, a.s., trimble, m.j., purdon, a., young-overton, k.d., pimm, s.l. & van aarde, r.j., 2017, ‘savanna elephant numbers are only a quarter of their expected values’, plos one 12(4), e0175942. https://doi.org/10.1371/journal.pone.0175942 robson, a.s. & van aarde, r.j., 2018, ‘changes in elephant conservation management promote density-dependent habitat selection in the kruger national park’, animal conservation 21(4), 302–312. https://doi.org/10.1111/acv.12393 roever, c.l., van aarde, r.j. & leggett, k., 2012, ‘functional responses in the habitat selection of a generalist mega-herbivore, the african savannah elephant’, ecography 35(11), 972–982. https://doi.org/10.1111/j.1600-0587.2012.07359.x selier, s.a.j., henley, m., pretorius, y. & garai, m., 2016, ‘a conservation assessment of loxodonta africana’, in m.f. child, l. roxburgh, e. do linh san, d. raimondo, j. selier & h.t. davies-mostert (eds.), the red list of mammals of south africa, swaziland and lesotho, south african national biodiversity institute and endangered wildlife trust, south africa. sibly, r.m., barker, d., denham, m.c., hone, j. & pagel, m., 2005, ‘on the regulation of populations of mammals, birds, fish, and insects’, science 309(5734), 607–610. https://doi.org/10.1126/science.1110760 slotow, r., page, b.r., mackey, r.l. & duffy, k.j., 2006, ‘modelling elephant population growth in small, fenced, south african reserves’, south african journal of wildlife research 36(1), 33–43. smit, i.p. & ferreira, s.m., 2010, ‘management intervention affects river-bound spatial dynamics of elephants’, biological conservation 143(9), 2172–2181. https://doi.org/10.1016/j.biocon.2010.06.001 thomas, b., holland, j.d., minot, e.o., 2008, ‘elephant (loxodonta africana) home ranges in sabisand reserve and kruger national park: a five-year satellite tracking study’, plos one3(12), e3902. https://doi.org/10.1371/journal.pone.0003902 trimble, m.j., ferreira, s.m. & van aarde, r.j., 2009, ‘drivers of megaherbivore demographic fluctuations: inference from elephants’, journal of zoology 279(1), 18–26. https://doi.org/10.1111/j.1469-7998.2009.00560.x trollope, w.s.w., trollope, l.a., biggs, h.c., pienaar, d. & potgieter, a.l.f., 1998, ‘long-term changes in the woody vegetation of the kruger national park, with special reference to the effects of elephants and fire’, koedoe 41(2), 103–112. https://doi.org/10.4102/koedoe.v41i2.255 tsoularis, a.n. & wallace, j., 2002, ‘analysis of logistic growth models’, mathematical biosciences179(1), 21–55. https://doi.org/10.1016/s0025-5564(02)00096-2 valeix, m., fritz, h., sabatier, r., murindagomo, f., cumming, d. & duncan, p., 2011, ‘elephant-induced structural changes in the vegetation and habitat selection by large herbivores in an african savanna’, biological conservation 144(2), 902–912. https://doi.org/10.1016/j.biocon.2010.10.029 van aarde, r.j., jackson, t.p. & ferreira, s.m., 2006, ‘conservation science and elephant management in southern africa: elephant conservation’, south african journal of science 102(9–10), 385–388. van aarde, r.j., whyte, i. & pimm, s., 1999, ‘culling and the dynamics of the kruger national park african elephant population’, animal conservation 2(4), 287–294. https://doi.org/10.1111/j.1469-1795.1999.tb00075.x van dyken, j.d. & zhang, b., 2019, ‘carrying capacity of a spatially-structured population: disentangling the effects of dispersal, growth parameters, habitat heterogeneity and habitat clustering’, journal of theoretical biology 460, 115–124. https://doi.org/10.1016/j.jtbi.2018.09.015 van wyk, p. & fairall, n., 1969, ‘the influence of the african elephant on the vegetation of the kruger national park’, koedoe 12(1), a747. https://doi.org/10.4102/koedoe.v12i1.747 venter, f.j., naiman, r.j., biggs, h.c. & pienaar d.j., 2008, ‘the evolution of conservation management philosophy: science, environmental change and social adjustments in kruger national park’, ecosystems 11, 173–192. https://doi.org/10.1007/s10021-007-9116-x verhulst, p.-f., 1845, ‘recherchesmathématiques sur la loid’accroissement de la population [mathematical researches into the law of population growth increase]’, nouveaux mémoires de l’académieroyale des sciences et belles-lettres de bruxelles 18, 14–50. whyte, i.j., 2001, ‘the conservation management of the kruger national park elephant population’, phd thesis, university of pretoria, pretoria. whyte, i.j., biggs, h.c., gaylard, a. & braack, l.e.o., 1999, ‘a new policy for the management of the kruger national park’s elephant population’, koedoe 42(1), 111–132. https://doi.org/10.4102/koedoe.v42i1.228 wickham, h., 2016, ggplot2: elegant graphics for data analysis, springer-verlag, new york, ny. wittemyer, g. & getz, w.m., 2007, ‘hierarchical dominance structure and social organization in african elephants, loxodontaafricana’, animal behaviour 73(4), 671–681. https://doi.org/10.1016/j.anbehav.2006.10.008 wittemyer, g., getz, w.m., vollrath, f. & douglas-hamilton, i., 2007, ‘social dominance, seasonal movements, and spatial segregation in african elephants: a contribution to conservation behavior’, behavioral ecology and sociobiology 61(12), 1919–1931. https://doi.org/10.1007/s00265-007-0432-0 wittemyer, g., keating, l.m., vollrath, f. & douglas-hamilton, i., 2017, ‘graph theory illustrates spatial and temporal features that structure elephant rest locations and reflect risk perception’, ecography 40(5), 598–605. https://doi.org/10.1111/ecog.02379 young, k.d., ferreira, s.m. & van aarde, r.j., 2009a, ‘the influence of increasing population size and vegetation productivity on elephant distribution in the kruger national park’, austral ecology 34(3), 329–342. https://doi.org/10.1111/j.1442-9993.2009.01934 young, k.d., ferreira, s.m. & van aarde, r.j., 2009b, ‘elephant spatial use in wet and dry savannas of southern africa’, journal of zoology 278(3), 189–205. https://doi.org/10.1111/j.1469-7998.2009.00568.x young, k.d. & van aarde, r.j., 2010, ‘density as an explanatory variable of movements and calf recruitment in savannah elephants across southern africa’, journal of animal ecology 79(3), 662–673. https://doi.org/10.1111/j.1365-2656.2010.01667.x appendix 1 table 1-a1: exponential growth model parameter estimates and model statistics (eqn 2). table 2-a1: parameter estimates of stochastic ricker model fitted to the time series of elephant population size, as part of test for negative density-dependent growth. table 3-a1: test statistics of mann–whitney u tests for difference in recruitment rates in the three long-term density zones. p-values corrected for multiple comparison by holm method. table 4-a1: test statistics of mann-kendall test for trend in the average elephant recruitment rates in areas belonging to different zones. figure 1-a1: (a) overall, (b) peripheral zone, (c) semi-peripheral zone and (d) core zone. deterministic component of ricker growth models (eqn 3) fitted to time-series of elephant population in kruger national park during the period of regular culling (1985–2012) model parameter estimates in appendix 1 (table 2-a1). models shown for overall population, and separately for three zones in the park having low medium or high long-term average elephant density (denoted as ‘peripheral’, ‘semi-peripheral’ and ‘core’ zones for elephants). test for negative density-dependent growth reported significant for all the zones, and the population overall. figure 2-a1: distribution plots of recruitment rate observations. (a) shows recruitment rates are skewed and (b) shows a cube root transformation of the recruitment rates, to reduce skewness. book review review of bats of southern and central africa, second edition book title: bats of southern and central africa: a biogeographic and taxonomic synthesis authors: a. monadjem; p.j. taylor; f. cotterill; m.c. schoeman isbn: 978-1-77614-582-9 publisher: wits university press, 2020, r600* *book price at time of review review title: review of bats of southern and central africa, second edition reviewer: leigh r. richards1 affiliation: 1mammal department, durban natural science museum, durban, south africa corresponding author: leigh richards, leigh.richards@durban.gov.za how to cite this article: richards, l.r., 2021, ‘review of bats of southern and central africa, second edition’, koedoe 63(1), a1678. https://doi.org/10.4102/koedoe.v63i1.1678 copyright notice: © 2021. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. the southern africa subregion harbours almost 10% of the global bat diversity (mammal diversity database 2020). the previous edition of bats of southern and central africa provided the first comprehensive account of the biogeographical and taxonomic information on the bat species of this highly diverse afrotropical region. it understandably became an essential reference for bat biologists and enthusiasts worldwide. the second edition of this landmark publication provides an extensive update of the biogeography, echolocation, ecology, evolution and taxonomy of the 125 bat species detailed in the book. the explanatory text in the first five chapters has mostly remained the same, with a few exceptions, such as the expanded sections ‘classification and taxonomy’, ‘foraging ecology and associated ecological services’ and ‘bat detectors’. the authors have also included a precautionary note on acceptable bat-handling practices. as the authors point out, taxonomies have undergone radical change since the first edition, published in september 2010. the introductory text for each bat family and identification matrices are updated with the latest scientific evidence. information on two new families, cistugidae and rhinonycteridae, is contained within unique colour-coded sections – a signature style of the publication. the revision includes 10 new species accounts; nine of these represent newly described or recognised species. the species accounts are unpacked in superb detail and augmented by the customary distribution maps, sonograms and colour photographs of the animal and its cranial and dental anatomy. revised taxonomies are presented for 11 species. care has gone into updating species distribution records and indicating instances of taxonomic uncertainty. the museum specimen list has expanded, with 300 new records (pp. 651–694). impressively, 73% of the 6300+ records were verified by the authors. it is unclear why sampling dates were added to some records and not to others. the 13-page reference list contains over 700 articles – a valuable scientific resource. from an aesthetic perspective, the typography has improved, with the use of darker fonts and more prominent diagram labels, making the text easier to read. a major issue of the previous edition was the subpar softcover binding. the current hardcover format is a welcome improvement, especially as the publication will often be used in the field. species distribution maps are cleanly edited and presented in vibrant colours. there is no longer a distinction between verified (red dots) and unverified occurrence records (blue dots). i miss the predicted species distribution ranges in the former edition. recent surveys in under-sampled regions of south africa have confirmed the extended ranges for epomophorus crypturus, kerivoula argentata, laephotis botswanae and neoromicia anchietae as predicted in the first edition (moir et al. 2020). the authors cite the obstruction of occurrence points and geopolitical boundaries as reasons for excluding species distribution modelling outputs from the second edition. this could possibly be corrected by overlaying coloured boundary lines and points on the predicted distribution range. some minor issues were noted: the back cover synopsis erroneously states that the book covers 124 taxa with eight new species. otomops martiensseni is classified regionally as ‘near threatened’ and not ‘least concern’ (child et al. 2016). the chromosomal number of the scotoecus hindei/albigula complex is not unknown (nagorsen et al. 1976). stanley (2013) highlighted the unsatisfactory images of the lancet or connecting processes of some rhinolophid bats in the previous edition. most of these images have been replaced with more suitable photographs. stanley (2013) also noted low-resolution photographs that rendered certain diagnostic traits, relating to dental morphology, difficult to discern. unfortunately, this remains an impediment in the revised version. however, with global efforts towards digitising museum collections, it is almost certain that subsequent editions will contain crania and mandible photographs of higher resolution. lastly, rapidly advancing taxonomic research means that this edition was slightly outdated by the time of publication (see the taxonomic revision of pipistrelle-like bats by monadjem et al. 2020). these minor matters do not detract from the impact of this revised masterpiece. i applaud the authors for their monumental efforts in producing an invaluable resource that will undoubtedly continue to enhance bat research and conservation within the region. references mammal diversity database, 2020, mammal diversity database (version 1.31), zenodo, viewed n.d., from http://doi.org/10.5281/zenodo.4139818 moir, m., richards, l.r., rambau, r.v. & cherry, m.i., 2020, ‘bats of eastern cape and southern kwazulu-natal forests, south africa: diversity, call library and range extensions’, acta chiropterologica 22(2), 365–381. https://doi.org/10.3161/15081109acc2020.22.2.011 monadjem, a., demos, t.c., dalton, d.l., webala, p.w., musila, s., kerbis peterhans, j.c. et al., 2020, ‘a revision of pipistrelle-like bats (mammalia: chiroptera: vespertilionidae) in east africa with the description of new genera and species’, zoological journal of the linnean society 191(4), 1114–1146. https://doi.org/10.1093/zoolinnean/zlaa087 nagorsen, d.w., eger, j.l. & petersen, r.l., 1976, ‘somatic chromosomes of three african species of bats (chiroptera), scotoecus hindei, tadarida aegyptiaca and tadarida bemmeleni’, mammalian chromosome newsletter 17(3), 9–12. child, m.f., roxburgh, l., do linh san, d., raimondo, d. & davies-mostert, h.t., 2016, ‘the red list of mammals of south africa, swaziland and lesotho’, south african national biodiversity institute and endangered wildlife trust, south africa. stanley, w.t., 2013, ‘bats of southern and central africa: a biogeographic and taxonomic synthesis, second edition, ara monadjem, peter j. taylor, fpd (woody) cotterill & m. corrie schoeman: book review’, journal of mammalogy 94(2), 518–551. https://doi.org/10.1644/12-mamm-r-184.1 filelist convert a pdf file! page 1 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 abstract introduction methods statistical analyses results discussion conclusion acknowledgements references appendix 1 about the author(s) edmund february department of biological sciences, university of cape town, south africa eleanor shadwell department of biological sciences, university of cape town, south africa storme viljoen department of biological sciences, university of cape town, south africa dawood hattas department of biological sciences, university of cape town, south africa citation february, e., shadwell, e., viljoen, s. & hattas, d., 2017, ‘feeding choices and impacts of extralimital giraffe on two keystone tree species in the kgalagadi national park’, koedoe 59(1), a1454. https://doi.org/10.4102/koedoe.v59i1.1454 original research feeding choices and impacts of extralimital giraffe on two keystone tree species in the kgalagadi national park edmund february, eleanor shadwell, storme viljoen, dawood hattas received: 17 dec. 2016; accepted: 07 apr. 2017; published: 29 may 2017 copyright: © 2017. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract in this article we determine the effect of an extralimital megaherbivore on the reproductive potential and vegetation structure of two keystone tree species in the auob river in the south western kalahari desert of southern africa. using spoor and dung counts we establish the presence of giraffe in three predetermined density zones by walking 50 transects across the river in each zone. we also photographed six trees from each species in each zone and use these photographs to determine browse impact on reproductive potential, canopy volume as well as the percentage dieback on the extremities of the canopy. we then perform stable carbon and nitrogen isotope analysis on the leaves of the trees and compare these relative to the isotope ratios of giraffe dung to ascertain dietary preference. crude protein was determined as a guide to nutritive value. finally, we determine both chemical and physical defences for the two species. our results show a significant negative impact of giraffe browse on tree canopies, no significant differences in recruitment and a noticeable decrease in flowers and pods at the giraffe browse height of 2 m – 5 m. no significant differences in crude protein or condensed tannins were found but significant differences in spinescence. giraffe are not endemic to the auob river and our study shows that the introduction of these animals is having a negative impact on the canopies of vachellia haematoxylon. while there are, as yet, no significant impacts on reproductive potential we speculate that this will happen with time. conservation implications: our study shows that giraffe are significantly impacting the canopies of two common tree species in the auob river in the arid kgalagadi transfrontier park. without management intervention an increasing population of giraffe will result in substantial changes to the plant community vegetation structure of the river. introduction for many tourists, the primary reason for visiting an african national park is to see a diversity of large mammals (grünewald, schleuning & böhning-gaese 2016). to meet this expectation, giraffe (giraffe camelopardalis) have been introduced into several reserves in south africa (bond & loffell 2001; parker & bernard 2005). in 1991 8 giraffe were introduced into an enclosure near the craig lockhart borehole on the auob river in the kgalagadi transfrontier park in southern africa, and in 1998 35 animals were released into the park from this enclosure (bezuidenhout, herbst & ferreira 2010; rowland 1991). while the historic distribution of giraffe probably included the southern kalahari, there are no historic records for resident populations of these animals in the southern sector of the kgalagadi transfrontier park, where they are considered extralimital (hall-martin & de graaff 1978; skinner & chimimba 2005). several studies have demonstrated negative impacts on vegetation structure with extralimital introductions (bond & loffell 2001; castley, boshoff & kerley 2001; parker & bernard 2005). giraffe in particular have been shown to exert extreme browse pressure on some vachellia species to the extent that these may be eliminated from the environment (bond & loffell 2001). giraffe are the only exclusively browsing megaherbivore in the kgalagadi transfrontier park, with a distinctive browse range of typically between 2 m and 5 m, which is higher than all other browsing species such as eland (taurotragus oryx) and kudu (tragelaphus strepsiceros) (owen-smith 1992). while giraffe may feed on leaves and shoots almost exclusively, flowers, fruits and pods are favoured when available (owen-smith 1992). as the food intake of a female giraffe is around 2.1% of its body mass and 1.6% for a male, an average 800 kg female will need approximately 16.8 kg of browse per day and an average 1200 kg male 19.2 kg (pellew 1984). in 2010 the number of giraffe restricted to the auob river was estimated to be around 50, with between 1 and 55 giraffe sightings per month in 2013 (sanparks [south african national parks] 2014 unpublished data). from our calculations, 50 giraffe would need around 900 kg of browse per day in an arid environment where a lack of resources would limit the potential for trees to replace lost biomass (bloom, chapin & mooney 1985; chapin 1980). large trees in arid savannahs perform essential ecosystem services by providing food, shade, nesting sites and increased nutrients for many other plant and animal species (dean, milton & jeltsch 1999; milton & dean 1995). two such species are vachellia erioloba and vachellia haematoxylon, both biogeographically endemic to the kalahari and making up almost all the large trees in the auob river (van rooyen et al. 2008). these two vachellia species have evolved in this arid environment where the paucity of substrate nutrients result in slow growth rates (chapin 1980; dean et al. 1999). as a result, the cost of herbivory is felt more acutely because plants have to draw on finite resources to recuperate biomass losses (grime 1977). here we examine the effects of giraffe browse on the canopies of two important tree species in the auob river, v. erioloba and v. haematoxylon. we do this through an analysis of a series of photographs of trees taken in high, medium and low giraffe density zones. we propose that an increase in browse by giraffe has had a detrimental effect on the plant community vegetation structure of the auob river by impacting on canopy structure and the reproductive potential of the two tree species through the removal of flowers and seed pods. in this arid environment, trees have slow growth rates, and as a result, we expected increased defences against herbivory (chapin 1980; coley, bryant & chapin 1985; grime 1977). we hypothesise that the tree species with the least defences against herbivory will be most targeted by giraffe, and/or that the most nutritionally valuable species will be favoured as forage. we test this hypothesis through a determination of the relative differences in chemical and physical defences of v. erioloba and v. haematoxylon as well as the nutritional quality of the leaves in the dry season. finally, we use the stable isotope ratios of giraffe dung to establish the relative contribution of the two tree species to the diet of giraffe in the auob river (codron et al. 2007). by implementing strategic adaptive management, sanparks have integrated science and management (biggs & rogers 2003). for such management to succeed in the kgalagadi transfrontier park, there has to be some understanding of the spatio-temporal heterogeneity of the rivers in the park as these are the centres for biodiversity (dean et al. 1999). elucidating the effects that an extralimital megaherbivore may have on this heterogeneity is critical for such an understanding. methods study site the study site is located along the ephemeral auob river in the kgalagadi transfrontier park in southern africa between −25.79944°, 20.0354° and −26.4216°, 20.6234°. giraffe were originally introduced into the auob at the craig lockhart borehole (−25.8655°, 20.1031°). the results of monthly surveys between february 2012 and march 2014 conducted by sanparks show that the highest densities of giraffe are still in this area decreasing in number down to the confluence with the nossob river where giraffe are rarely seen (sanparks unpublished data). the auob river has a narrow channel between 100 m and 500 m wide consisting of fine-grained silts of the goeboe goeboe formation set in 30 m to 50 m high banks of calcrete of the mokalanen formation (malherbe 1984; mills & retief 1984). while there is no surface water, the auob has an aquifer at a depth of between 38 m and 46 m below the surface that has been tapped at intervals to supply 15 permanent water holes for animals (mills & retief 1984; van wyk & le riche 1984). in this narrow channel, the vegetation is dominated by v. erioloba and v. haematoxylon as the only tree species and the low shrub rhigozum trichotomum. common grasses are schmidtia kalahariensis, stipagrostis obtusa and eragrostis porosa (van rooyen et al. 2008). the annual rainfall for the region is distinctly seasonal occurring as erratic and highly localised thunderstorms between november and april at the hottest time of the year with the coldest season also being the driest. mean annual rainfall (1984–2014) is 220 mm at twee rivieren (−26.4721°, 20.6116°) and mean maximum and minimum temperatures 36.7 °c and 0.1 °c (south african weather bureau). giraffe and tree density based on monthly large-mammal surveys conducted by sanparks between 2012 and 2014, we divided the auob river downstream of the mata mata rest camp (−25.7680, 20.0005) into three consecutive zones, ± 30 km in length. these zones start as high giraffe density from just below mata mata (−25.7944°, 20.0354° to −25.9636°, 20.2446°) through medium-density (−26.9989°, 20.3468° to −26.1760°, 20.5432°) to low-density finishing near the confluence of the auob and nossob rivers (−26.1926°, 20.5488° to −26.4216°, 20.6234°). to confirm our assessment based on the monthly large-mammal surveys, we walked 50 transects in each zone perpendicular to the riverbed from mid-slope of one bank to mid-slope of the opposite bank. each transect was done as a pair, with 150 m between each transect and 1 km between each pair of transects. if a borehole (waterhole) were to occur between a pair, a kilometre margin was given on either side to discount the piosphere effect on vegetation and animal density (chamaillé-jammes, fritz & madzikanda et al. 2009). in each transect, every occurrence of giraffe spoor and dung was counted by two people 5 m apart for a total width of 10 m on each transect. to determine the effect that giraffe browse may have on tree density, each observer also noted the occurrence of any v. erioloba and v. haematoxylon in three specific height classes, 0 m – 0.3 m, 0.3 m – 2 m and above 2 m. impact of giraffe browse to determine the impact of giraffe browse on canopy dieback and reproductive potential (flowers and seed pods), we photographed two trees from each of three height classes (2 m – 4 m, 4.1 m – 6 m and above 6 m) in each giraffe density zone for each species (36 trees). each tree was photographed twice with each photograph at 90° to the other. from each photograph, we then determined browse impact for each tree at every metre from 2 m above ground level to the top of the canopy. we did this from 2 m above ground level to 5 m because several studies have shown that while a male giraffe may feed at up to 6 m, the average giraffe browse height for both male and female is between 2 m and 4.5 m (birkett 2002; du toit 1990; young & isbell 1991). all photographs were taken in january 2013 using a nikon d60 camera (nikon, ayuthaya, thailand) with the entire tree just inside the field of view through an 18 mm – 200 mm f/35 to 6.3-hsm dc lens (sigma, fukushima, japan) fitted with a pro 1 d uv (w) filter (kenko, tokyo, japan). the aperture was fixed at 52 mm, f8 and a 5 m retractable aluminium ranging rod (levelling staff; leica geosystems, st gallen, switzerland) was held vertically at the edge of the canopy for calibration. the photographs were taken in raw format and adjusted into tiff images when analysed in adobe photoshop cs5 v12.0 × 32© (adobe systems software ltd, ireland). each photograph was overlaid by a calibrated grid (50 cm boxes, subdivided by 10 cm; figure 1-a1). for an assessment of canopy structure, we measured the width of the canopy from edge to edge at every vertical metre starting at 2 m above ground level. canopy dieback was determined by marking all living (green leaves) and dead material (brown leaves or twigs or branches < 5 cm thick) at each grid intercept (11 points) occurring within each of the 50 cm lengths starting from the outer edge of the canopy moving inwards along the metre-line. we did this for both photographs for each tree. intercepts occurring in branch or leaf free spaces or with main branches were disregarded. canopy dieback at each metre-line was calculated as the number of intercepts denoting dead material over the total number of intercepts (living + dead material) and averaged over two photographs. canopy dieback was also calculated for two sections of each tree, within browse height (2 m – 5 m) and above browse height (> 5 m) by averaging the percentage of canopy dieback for each metre-line within each section. to determine the effects of giraffe browse on reproduction, the numbers of visible flowers and pods were counted in a 50-cm square box; 25 cm on either side of each horizontal metre-line and 50 cm inwards from the outer edges of the canopy. the number of boxes depended on how many edges of the canopy there were. we averaged the counts for all the boxes for both photographs of each tree. we do this for each metre-line before averaging all values for above and within browse height for each tree. dietary proportion we used the stable carbon and nitrogen isotope ratios of leaves and giraffe dung to determine dietary proportion of the two tree species (codron et al. 2007). for this we collected ± 100 g of fully mature, whole leaves from 2 m – 5 m above ground on the northern aspect of 10 trees from each of our two vachellia species in the high giraffe density zone. along our transects, we also opportunistically collected 20 fresh giraffe dung samples. both leaf and dung samples were dried to constant weight at 70 °c before grinding to a fine powder using a retsch mm 200 ball mill (retsch, haan, germany). for both leaves and dung, we determined percentage nitrogen as well as isotopic ratios of 12c/13c and 15n/14n using a thermo finnigan delta plus xp mass spectrometer coupled with a conflo iii device to a thermo finnigan flash ea 1112 elemental analyser (thermo electron corporation, milan, italy). we calibrated the results relative to atmospheric n2 for nitrogen and pee-dee belemnite for carbon as well as to correct for drift in our reference gas. deviation from the standard is denoted by the term δ, and the results expressed as parts per thousand (‰). precision of duplicate analysis is 0.1‰ for carbon and 0.2‰ for nitrogen (february et al. 2011). crude protein there is a strong positive correlation between protein content of the leaf and animal preference (cooper, owen-smith & bryant 1988; heady 1964). to assess preference of the foliage of our two tree species to giraffe, we determined differences in crude protein of the leaves by multiplying the foliar percentage nitrogen by 6.25 (cooper et al. 1988). plant defences plants may defend themselves against mammalian herbivory either chemically with high concentrations of condensed tannins and insoluble fibre or structurally with spines or thorns (hanley et al. 2007; hattas et al. 2011). condensed tannins have a characteristic affinity for protein, forming insoluble complexes, which render protein less digestible and thereby reduces forage quality, whereas spines and thorns affect bite size and feeding efficiency (hattas et al. 2011; mueller-harvey 2006). we quantified condensed tannins (proanthocyanidins) while also deriving an index of spinescence for v. erioloba and v. haematoxylon. both determinations used the distal 20 cm of seven branches per tree on which we also counted the number of broken thorns. for this, branches located at 2 m – 5 m above ground level were randomly selected from seven trees per species in june 2013. we determined condensed tannins using the acid-butanol assay as described by hagerman (2002) using sorghum tannin as a standard (hattas & julkunen-tiitto 2012). plant fibre content was determined using the acid detergent fibre (adf) method in an ankom 220 fiber analyzer using adf method 5 (ankom technology). we counted the number of broken thorns on each branch before measuring thorn length and basal diameter of each thorn, as well as the distance between sets of paired thorns (inter-thorn distance) and leaf length, defined as the distance from the petiole intersection on the branch to the tip of the leaf (midgley, botha & balfour 2001). we used these measurements to develop a spinescence index (t1) to compare the relative degree of physical defence for each species (adapted from midgley et al. [2001]): statistical analyses giraffe densities between zones from spoor and dung counts were each tested using a chi-squared test followed by a post-hoc paired t-test. differences in the densities of large trees/ha between zone were tested with a poisson generalised linear model using an offset of log (transect area) to account for any differences in area covered. the densities of juveniles/large tree/ha and intermediates/large tree/ha of each species were both tested with a generalised linear model. transects with juveniles/intermediates but no large trees were removed from the analyses. as giraffe browse at heights between 2 m and 5 m, we compared every metre of canopy width measurement with the 5-m level across different giraffe density zones using a linear mixed-effects model. average percentage canopy dieback was arcsine transformed, and all height classes were compared relative to each other and between zones and species using a generalised linear mixed effects model. the same test was used to compare average number of flowers and pods between browse height and above browse height for v. haematoxylon (v. erioloba was not flowering) after log transformation of the data. for every analysis, the variance between individual trees was accounted for by including tree number as a random effect. if the interactions between zone and species were not significant, then the analyses were run separately on each species. all tests on the data were assessed in r© v3.1.2 (r_development_core_team 2014), and a value of p < 0.05 was required for significance. we used the siar package v4.2 in r v3.1.2 to determine the dietary proportion of each species (parnell et al. 2010). this package fits a bayesian markov chain monte carlo (mcmc) model to the dietary habits of giraffe based on the similarity of isotopic ratios between food sources and giraffe dung. percentage carbon and nitrogen for each food source (v. erioloba and v. haematoxylon) along with the δ13c and δ15n values of food sources and dung is incorporated into a linear mixing model. calorific content, condensed tannin concentrations, percentages of adf, carbon and nitrogen, and the proportion of crude protein were compared between the two species using a t-test after being tested for normality using a kolmogorov–smirnoff test. thorn length was averaged for each pair and again for each branch. leaf length and distance between thorn pairs were averaged for each branch. significant differences were determined using mann–whitney u-tests for mean thorn length (only non-zero values included), mean basal diameter and the means for the spinescence index, as these data exhibit a non-parametric distribution according to a shapiro–wilk normality test. t-tests were used to compare means for leaf length and distance between thorn pairs. results giraffe and tree density our transect results confirmed our assumptions based on the sanparks monthly large-mammal surveys. there was a significant increase in both spoor and dung (p < 0.001, in both cases) from the low giraffe density zone to the high-density zone. giraffe spoor and dung were present in, respectively, 28% and 8% of transects in the low-density zone, 56% and 46% of transects in the medium-density zone and 94% and 72% of transects in the high-density zone. there were significantly more large (> 2 m high) v. erioloba than large v. haematoxylon trees in the auob river (p < 0.001, table 1). there was also a significant increase in the density of large v. erioloba from 4.35 trees/ha in the high giraffe density zone to 7.11 trees/ha in the medium-density zone (p < 0.01) and a significant decrease to 1.15 trees/ha in the low-density zone. this trend is also reflected non-significantly in the relative proportion of juveniles and intermediates (table 1). table 1: relative tree densities in three specific height classes: juvenile, intermediate and large (0 m – 0.3 m, 0.3 m – 2 m, and above 2 m, respectively) in each of the three giraffe density zones, low, medium and high. for v. haematoxylon, there are significantly fewer large trees (1.30 trees/ha) in the high giraffe density zone relative to the medium(3.23 trees/ha) and low-density zones (3.03 trees/ha, p < 0.001). there were however no significant differences in the number of juveniles or intermediates between the different zones (table 1). impact of giraffe browse there were no significant effects of giraffe browse on canopy width for v. erioloba between the different zones. there were also no significant differences in canopy width for v. haematoxylon between the mediumand high-density zones, but there were significant differences in canopy width between the high/medium-density zones and the low-density zone (p < 0.05). across all density zones v. haematoxylon canopies were significantly narrower below 5 m relative to the 5-m browse height (figure 1, p < 0.05). figure 1: average canopy width for (a) vachellia erioloba and (b) vachellia haematoxylon in the low, medium and high giraffe density zones. our results show high levels of canopy dieback in v. haematoxylon relative to v. erioloba in all giraffe density zones (p < 0.001) with average percentage dieback for v. haematoxylon twice that of v. erioloba (figure 2). in the high-density zone, there was more canopy dieback for v. erioloba between 2 m and 5 m than in the other two zones increasing from 6.5% in the low-density zone through 12% in the medium-density zone to 15.3% in the high-density zone. for v. haematoxylon, there was an increase in canopy dieback as giraffe density increases with the greatest percentage dieback in the high-density zone between 2 m and 4 m (p < 0.05, figure 2). figure 2: average canopy dieback for (a) vachellia erioloba and (b) vachellia haematoxylon in each of the giraffe density zones. for v. haematoxylon, there were significantly more flowers per tree in the low-density zone (2.03 ± 0.93 mean ± standard error [se]) than in the medium-density (0.17 ± 0.11 mean ± se) and no flowers in the high giraffe density zone (p < 0.05). there were however no significant differences in the number of flowers in the browse range (2 m – 5 m) and above browse range. there were also no significant differences in the number of pods above and below giraffe browse height for either species (figure 2-a1). stable isotope ratios there was no significant difference in δ13c values between v. erioloba (−26.3‰) and v. haematoxylon (−25.7‰). there was however a significant difference in δ15n values (p < 0.01), which allows these two vachellia species to be differentiated isotopically. giraffe dung δ15n values (5.5‰) were more similar to v. haematoxylon (6.1‰) than v. erioloba (8.4‰). the results from our model show that up to 100% of the diet of giraffe is v. haematoxylon (79%, ± 20.5%, mean ± se) with only a small proportion of v. erioloba (21%, ± 20.5%, mean ± se, figure 3). figure 3: the relative proportion of vachellia erioloba and vachellia haematoxylon in the diet of giraffe in the auob river as derived from stable isotope analysis of dung. crude protein there was a significantly higher concentration of crude protein (p < 0.05) and lower c:n ratio (p < 0.01) in v. erioloba (13.8% and 22.1%) than in v. haematoxylon (12.2% and 25.8%). plant defences there were no significant differences between the two tree species in condensed tannin concentrations (v. erioloba 3.3% ± 0.32 and v. haematoxylon 2.9% ± 0.29, mean ± se) but there are significant (p < 0.001) differences in thorn length and thorn basal diameter, with v. erioloba having longer (21.6 mm ± 11.9 mm and 19.0 mm ± 11.8 mm) and thicker (2.5 mm ± 1.4 mm and 0.7 mm ± 0.3 mm) thorns than v. haematoxylon. percentage adf in v. haematoxylon was significantly higher (11.3%) than that of v. erioloba (50.1 ± 0.81% and 44.4 ± 0.61%, mean ± se, p < 0.0001). of the two trees, v. haematoxylon also had a significantly (p < 0.001) greater proportion of broken thorns per branch (22.4 ± 13.2 and 11.8 ± 13.8, mean ± se). the results for our spinescence index shows that v. erioloba is significantly better physically defended than v. haematoxylon (figure 4, p < 0.05). figure 4: means and interquartile range for our spinescence indices showing that vachellia erioloba is significantly more spinescent that vachellia haematoxylon. discussion giraffe densities are highest in the zone closest to the point of release near the craig lockhart borehole on the auob river decreasing downstream towards the confluence with the nossob. this distribution coincides with the densities of large trees with v. erioloba highest and v. haematoxylon lowest where the densities of giraffes are highest. the density of large v. haematoxylon trees increases from the high giraffe density zone to the low-density zone. this increase is coincidental with our results for browse impact, which shows that while the highest impact of giraffe browse is in the high-density zone, this impact is not on v. erioloba but rather on v. haematoxylon. when compared relative to trees in the mediumand low-density zones, there was no significant change in canopy structure for v. erioloba but much narrower canopies and significantly higher canopy dieback on v. haematoxylon in the high-density zone. while the canopy dieback reported here may be attributed to drought or natural mortality, these results are closely correlated with giraffe density and it is therefore most likely that giraffe are the cause. short periods of drought should also not result in canopy dieback because these trees are sourcing water from a deep aquifer and are therefore not reliant on immediate rainfall (schachtschneider & february 2013; shadwell & february 2017). vachellia erioloba flowers between july and september and v. haematoxylon between november and january (coates palgrave 2002; sekhwela & yates 2007). as our photographs were taken in january, which is a month after the flowering period for v. erioloba and at the end of flowering for v. haematoxylon, it is not surprising that we saw no flowers on v. erioloba trees. our results showing no flowers on v. haematoxylon in the high giraffe density zone and significantly fewer flowers in the medium zone relative to the low-density zone may also be the result of a combination of both flowering phenology and giraffe browse. more research has to be done on this to determine the exact effect of giraffe browse on the amount of flowers. the pods of both v. erioloba and v. haematoxylon are indehiscent and sought after by mammals, with passage through the gut aiding germination (leistner 1961). vachellia erioloba sets seed from december to march and v. haematoxylon from january to april (coates palgrave 2002; sekhwela & yates 2007). the very few pods on v. haematoxylon relative to v. erioloba across all giraffe density zones in our study may therefore relate to seed set rather than giraffe browse (coates palgrave 2002; sekhwela & yates 2007). research has however shown a distinct browse line for pods on v. erioloba resulting from giraffe browse (coe 1998). as seed pods that fall to the ground are readily infected by bruchid beetle, germination success is higher for ingested seed taken from the tree (hoffman et al. 1989). giraffe browse of the seed pod may therefore aid rather than retard germination for our two study species. our transect results show no correlation between the number of juvenile v. haematoxylon trees and giraffe density. the reason for this could be that savannah trees have a high resilience to stem mortality and may persist as juveniles for many decades (higgins et al. 2007). the drivers for vachellia seed germination and establishment at our study site are not clear but once established those trees top killed by fire, herbivory or drought can persist for many decades resprouting from the base using stored resources (schutz, bond & cramer 2011). the increased impact of giraffe on the canopy of v. haematoxylon rather than v. erioloba as demonstrated by our photographic analysis is corroborated by our stable isotope analysis that shows 80% of giraffe diet to be v. haematoxylon. there were no significant differences in the concentration of condensed tannins between the two species demonstrating that neither species were better chemically defended (cooper & owen-smith 1985). these concentrations, for both species, were less than the frequently reported 5% threshold above which condensed tannins are purported to negatively affect diet selection (cooper & owen-smith 1985). calorific content also did not differ between the two species, but there was a significant difference in c:n ratios making v. erioloba with lower c:n ratios more preferable as a food source (pérez-harguindeguy et al. 2003). our spinescence index showed that v. erioloba with thicker, longer and denser thorns was considerably better physically defended against herbivory than v. haematoxylon. on several occasions, we witnessed giraffe with their heads inside the canopy of a v. haematoxylon, which is likely what caused the greater degree of broken thorns on v. haematoxylon branches in our study. several studies have noted that giraffe are impervious to physical defences by thorns (cooper & owen-smith 1986; pellew 1984), however the results of our study would suggest that the longer, thicker, more prolific thorns on v. erioloba may explain why giraffe have a dietary preference for v. haematoxylon even though v. erioloba is more nutritious. conclusion our study is equivocal in demonstrating a relationship between giraffe and seedling establishment at our study site. it does however show that giraffe are significantly reducing the canopies of both common tree species in the auob river. this impact is far greater for v. haematoxylon than for v. erioloba and if allowed to continue could lead to substantial changes in the plant community vegetation structure of the auob river. it has been proposed that large trees in arid savannahs perform essential ecosystem services by providing food, shade, nesting sites and increased nutrients for many other plant and animal species (dean et al. 1999; milton & dean 1995). any decline of these large trees will have considerable implications for biodiversity conservation. giraffe are regarded as an extralimital in the kgalagadi transfrontier park and were primarily introduced to promote tourism (hall-martin & de graaff 1978; rowland 1991; skinner & chimimba 2005). while the economic benefits of tourism are important for socio-economic development, management has to consider the implications of an increasing giraffe population on biodiversity conservation. our study is the first step in implementing strategic adaptive management that will allow for a compromise between increased tourism revenue and biodiversity conservation (biggs & rogers 2003). such adaptive management could consider future research on why giraffe have not moved away from the area of highest impact into the medium and low-density zones where the densities of large v. haematoxylon are higher. future research could also focus on the effect of giraffe on seedling establishment of our two study species. acknowledgements the authors thank sanparks for allowing them to do the research in the kgalagadi transfrontier park. the authors are grateful to graeme ellis, paola vimercati, chantal elston and amy betzelberger for their help with the fieldwork. the data are available at the south african national park data repository http://dataknp.sanparks.org/sanparks/metacat/judithk.111224.3/sanparks. funding for the project was through a grant from the mellon foundation of new york. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions e.f. originated the study. e.f., e.s. and s.v. designed the experiments and conducted the field work. e.s. and s.v. performed the statistical analysis. s.v. conducted the dietary proportion, and d.h. and s.v. did the plant defences analysis. e.s. conducted the impact of giraffe browse and giraffe density analysis. e.f., e.s. and s.v. wrote the first draft, and all authors contributed equally to the final draft. references bezuidenhout, h., herbst, m. & ferreira, s., 2010, an assessment of the proposed giraffe translocation from the mata mata section to unions end in the kalahari gemsbok national park, sanparks internal report, south african national parks, kimberley, south africa. biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in h.c. biggs, k.h. rogers & j. du toit (eds.), the kruger experience. ecology and management of savanna heterogeneity, pp. 59–80, island press, washington, dc. birkett, a., 2002, ‘the impact of giraffe, rhino and elephant on the habitat of a black rhino sanctuary in kenya’, african journal of ecology 40, 276–282. https://doi.org/10.1046/j.1365-2028.2002.00373.x bloom, a.j., chapin, f.s. & mooney, h.a., 1985, ‘resource limitation in plants – an economic analogy’, annual review of ecology and systematics 16, 363–392. https://doi.org/10.1146/annurev.es.16.110185.002051 bond, w.j. & loffell, d., 2001, ‘introduction of giraffe changes acacia distribution in a south african savanna’, african journal of ecology 39, 286–294. https://doi.org/10.1046/j.1365-2028.2001.00319.x castley, j., boshoff, a. & kerley, g., 2001, ‘compromising south africa’s natural biodiversity: inappropriate herbivore introductions’, south african journal of science 96, 365–378. chamaillé-jammes, s., fritz, h. & madzikanda, h., 2009, ‘piosphere contribution to landscape heterogeneity: a case study of remote-sensed woody cover in a high elephant density landscape’, ecography 32, 871–880. https://doi.org/10.1111/j.1600-0587.2009.05785.x chapin, f.s., 1980, ‘the mineral nutrition of wild plants’, annual review of ecology and systematics 11(1), 233–260. https://doi.org/10.1146/annurev.es.11.110180.001313 coates palgrave, k., 2002, trees of southern africa, struik, cape town. codron, d., codron, j., lee-thorp, j., sponheimer, m., de ruiter, d., sealy, j. et al., 2007, ‘diets of savanna ungulates from stable carbon isotope composition of faeces’, journal of zoology 273, 21–29. https://doi.org/10.1111/j.1469-7998.2007.00292.x coe, m., 1998, ‘some aspects of the interaction between mammalian herbivores and acacia eriobola e. mey’, transactions of the royal society of south africa 53, 141–147. https://doi.org/10.1080/00359199809520382 coley, p.d., bryant, j.p. & chapin, f.s., 1985, ‘resource availability and plant antiherbivore defense’, science 230, 895–899. cooper, s.m. & owen-smith, n., 1985, ‘condensed tannins deter feeding by browsing ruminants in a south african savanna’, oecologia 67, 142–146. https://doi.org/10.1007/bf00378466 cooper, s.m. & owen-smith, n., 1986, ‘effects of plant spinescence on large mammalian herbivores’, oecologia 68, 446–455. https://doi.org/10.1007/bf00376934 cooper, s.m., owen-smith, n. & bryant, j.p., 1988, ‘foliage acceptability to browsing ruminants in relation to seasonal changes in the leaf chemistry of woody plants in a south african savanna’, oecologia 75, 336–342. https://doi.org/10.1007/bf01036753 dean, w., milton, s. & jeltsch, f., 1999, ‘large trees, fertile islands, and birds in arid savanna’, journal of arid environments 41, 61–78. https://doi.org/10.1006/jare.1998.0455 du toit, j., 1990, ‘feeding-height stratification among african browsing ruminants’, african journal of ecology 28, 55–61. https://doi.org/10.1111/j.1365-2028.1990.tb01136.x february, e.c., allsopp, n., shabane, t. & hattas, d., 2011, ‘coexistence of a c4 grass and a leaf succulent shrub in an arid ecosystem. the relationship between rooting depth, water and nitrogen’, plant and soil 349(1–2), 253–260. https://doi.org/10.1007/s11104-011-0867-y grime, j.p., 1977, ‘evidence for the existence of three primary strategies in plants and its relevance to ecological and evolutionary theory’, american naturalist 111(982), 1169–1194. https://doi.org/10.1086/283244 grünewald, c., schleuning, m. & böhning-gaese, k., 2016, ‘biodiversity, scenery and infrastructure: factors driving wildlife tourism in an african savannah national park’, biological conservation 201, 60–68. https://doi.org/10.1016/j.biocon.2016.05.036 hagerman, a.e., 2002, the tannin handbook (updated: 2011), department of chemistry & biochemistry, miami university, coral gables, fl. hall-martin, a., & de graaff, g., 1978, ‘a note on the feasibility of introducing giraffe to the kalahari gemsbok national park’, koedoe 21, 191–193. https://doi.org/10.4102/koedoe.v21i1.973 hanley, m.e., lamont, b.b., fairbanks, m.m. & rafferty, c.m., 2007, ‘plant structural traits and their role in anti-herbivore defence’, perspectives in plant ecology, evolution and systematics 8, 157–178. https://doi.org/10.1016/j.ppees.2007.01.001 hattas, d., hjältén, j., julkunen-tiitto, r., scogings, p.f. & rooke, t., 2011, ‘differential phenolic profiles in six african savanna woody species in relation to antiherbivore defense’, phytochemistry 72, 1796–1803. https://doi.org/10.1016/j.phytochem.2011.05.007 hattas, d. & julkunen-tiitto, r., 2012, ‘the quantification of condensed tannins in african savanna tree species’, phytochemistry letters 5, 329–334. https://doi.org/10.1016/j.phytol.2012.02.013 heady, h.f., 1964, ‘palatability of herbage and animal preference’, journal of range management 17, 76–82. https://doi.org/10.2307/3895315 higgins, s.i., bond, w.j., february, e.c., bronn, a., euston-brown, d.i.w., enslin, b. et al., 2007, ‘effects of four decades of fire manipulation on woody vegetation structure in savanna’, ecology 88, 1119–1125. https://doi.org/10.1890/06-1664 hoffman, m., cowling, r., douie, c. & pierce, s., 1989, ‘seed predation and germination of acacia erioloba in the kuiseb river valley, namib desert’, south african journal of botany 55, 103–106. https://doi.org/10.1016/s0254-6299(16)31237-6 leistner, o., 1961, ‘on the dispersal of acacia giraffae by game’, koedoe 4, 101–104. https://doi.org/10.4102/koedoe.v4i1.831 malherbe, s., 1984, ‘the geology of the kalahari gemsbok national park’, koedoe 27, 33–44. https://doi.org/10.4102/koedoe.v27i2.567 midgley, j., botha, m. & balfour, d., 2001, ‘patterns of thorn length, density, type and colour in african acacias’, african journal of range and forage science 18, 59–61. https://doi.org/10.2989/10220110109485756 mills, m. & retief, p., 1984, ‘the response of ungulates to rainfall along the riverbeds of the southern kalahari’, koedoe 27, 129–141. https://doi.org/10.4102/koedoe.v27i2.574 milton, s. & dean, w., 1995, ‘how useful is the keystone species concept, and can it be applied to acacia erioloba in the kalahari desert?’, zeitschrift fuer oekologie und naturschutz 3, 147–156. mueller-harvey, i., 2006, ‘unravelling the conundrum of tannins in animal nutrition and health’, journal of the science of food and agriculture 86, 2010–2037. https://doi.org/10.1002/jsfa.2577 owen-smith, r.n., 1992, megaherbivores: the influence of very large body size on ecology, cambridge university press, new york. parker, d. & bernard, r., 2005, ‘the diet and ecological role of giraffe (giraffa camelopardalis) introduced to the eastern cape, south africa’, journal of zoology 267, 203–210. https://doi.org/10.1017/s0952836905007399 parnell, a.c., inger, r., bearhop, s. & jackson, a.l., 2010, ‘source partitioning using stable isotopes: coping with too much variation’, plos one 5, e9672. https://doi.org/10.1371/journal.pone.0009672 pellew, r.a., 1984, ‘food consumption and energy budgets of the giraffe’, journal of applied ecology 21(1), 141–159. https://doi.org/10.2307/2403043 pérez-harguindeguy, n., díaz, s., vendramini, f., cornelissen, j.h., gurvich, d.e. & cabido, m., 2003, ‘leaf traits and herbivore selection in the field and in cafeteria experiments’, austral ecology 28, 642–650. https://doi.org/10.1046/j.1442-9993.2003.01321.x r_development_core_team., 2014, r: a language and environment for statistical computing, r foundation for statistical computing, vienna, austria. rowland, f., 1991, kameelperde pas goed aan in kalahari, pp. 31–35, custos, south african national parks, pretoria. schachtschneider, k. & february, e.c., 2013, ‘impact of prosopis invasion on a keystone tree species in the kalahari desert’, plant ecology 214, 597–605. https://doi.org/10.1007/s11258-013-0192-z schutz, a.e., bond, w.j. & cramer, m.d., 2011, ‘defoliation depletes the carbohydrate reserves of resprouting acacia saplings in an african savanna’, plant ecology 212, 2047–2055. https://doi.org/10.1007/s11258-010-9883-x sekhwela, m. & yates, d., 2007, ‘a phenological study of dominant acacia tree species in areas with different rainfall regimes in the kalahari of botswana’, journal of arid environments 70, 1–17. https://doi.org/10.1016/j.jaridenv.2006.12.006 shadwell, e. & february, e., 2017, ‘effects of groundwater abstraction on two keystone tree species in an arid savanna national park’, peerj 5, e2923. https://doi.org/10.7717/peerj.2923 skinner, j.d. & chimimba, c.t., 2005, the mammals of the southern african sub-region, cambridge university press, cape town. van rooyen, m.w., van rooyen, n., bothma, j.d.p. & van den berg, h.m, 2008, ‘landscapes in the kalahari gemsbok national park, south africa’, koedoe 50, 99–112. https://doi.org/10.4102/koedoe.v50i1.154 van wyk, p., & le riche, e., 1984, ‘the kalahari gemsbok national park: 1931–1981’, koedoe 27, 21–31. https://doi.org/10.4102/koedoe.v27i2.566 young, t.p. & isbell, l.a., 1991, ‘sex differences in giraffe feeding ecology: energetic and social constraints’, ethology 87, 79–89. https://doi.org/10.1111/j.1439-0310.1991.tb01190.x appendix 1 figure 1-a1: an illustration of the grid overlay and intercept counting method used for determining the amount of live or dead canopy material and the number of flowers and pods in the browse range and above browse range. figure 2-a1: average number of pods (± standard error) for (a) vachellia erioloba and (b) vachellia haematoxylon in each of the giraffe density zones (low, medium and high). filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 filelist convert a pdf file! page 1 abstract introduction literature background research method results discussion conclusions acknowledgements references about the author(s) zoëgné luyt school of tourism management, faculty of economic and management sciences, north-west university, potchefstroom, south africa peet van der merwe school of tourism management, faculty of economic and management sciences, north-west university, potchefstroom, south africa citation luyt, z. & van der merwe, p., 2022, ‘the potential of frogging as an ecotourism product for south african national parks’, koedoe 64(1), a1725. https://doi.org/10.4102/koedoe.v64i1.1725 project research number: ems2016/11/04-0213 original research the potential of frogging as an ecotourism product for south african national parks zoëgné luyt, peet van der merwe received: 29 mar. 2022; accepted: 23 june 2022; published: 21 july 2022 copyright: © 2022. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract amid global biodiversity loss, it is important to find practical tools and solutions in order to protect biodiversity. ecotourism is the fastest-growing sector of the international travel industry and can be a powerful conservation tool that encourages people to protect the natural environment. traditionally, frogs have not generated much attention among ecotourists, partly because they are easily overshadowed by other more charismatic species or habitat attractions. with almost a third of the nearly 7000 known amphibian species listed as threatened by the international union for conservation of nature (iucn), their protection is crucial. frogging is a well-known term within the frog conservation society, describing the activity of searching for frogs in the wild. this can be combined with other ecotourism activities to attract tourists and create an interest in the conservation of frogs while having fun at the same time. the aim was to determine the ecotourism potential of frogs in south africa, primarily by distributing questionnaires to tourists to retrieve information on whether they would be interested in participating in frog-related ecotourism activities within the south african national parks. for this research, a quantitative research approach was followed, namely non-probability sampling, to which convenience sampling was applied. an online survey (questionnaire) was designed to collect the data for the research. the survey outcome was satisfactory, as potential tourists indicated that they would like to participate in frog-related activities. the project offers the opportunity to conserve frogs, educate tourists, and create job opportunities within the local communities. it will also create a new tourism product for the south african national parks. conservation implications: the contribution of this research to conservation lies in the opportunity to benefit frog conservation through ecotourism. keywords: frogs; ecotourism; conservation; frogging; south africa; natural area tourism; sanparks. introduction one of the main reasons for travelling the world is to view the world’s natural wonders and experience and use different features of nature for enjoyment. national parks and other protected areas play a pivotal role in providing tourists with nature-based attractions to visit (newsome, moore & dowling 2013), and tourism worldwide plays an essential role in protected areas as a mechanism to promote and fund conservation in these areas, especially in africa (morrison et al. 2012). south africa’s natural attractions, both fauna and flora, are vibrant and diverse and have led to a continuous increase in visitors to south africa (department of environmental affairs and tourism [deat] 2008). in 2018, there were 29 million overnight trips within south africa, with 18.6 million of these trips made by domestic tourists and 10.4 million by international travellers (south african tourism 2019a). nature-based tourism products in south africa have been exemplified by the big five, cape fynbos, wildlife safaris, bird watching and geological sites such as table mountain (deat 2008). these products are well promoted and established within the south african tourism industry and generate the same income as the combined total income of fisheries, forestry and farming (metin 2019; sat 2019b). among south africa’s most significant nature-based tourism products are its 19 national parks, which fall under the management of sanparks (2020). these national parks are spread over seven of the nine provinces of south africa (covering an area of just over 4 million ha), represent 67% of the protected areas managed by the state (sanparks 2020) and receive approximately 6.3 million visitors per year (sanparks 2020). statistics from 2016 to 2017 indicate that 1.8 million tourists visited the kruger national park, one of south africa’s flagship conservation areas (brett 2018). in south africa, the national parks were created firstly to conserve natural resources (thomas & middleton 2003) and secondly to introduce tourism into these areas to create opportunities for tourists to learn about the natural resources and engage in nature-based activities, as well as to fund park operations (van der merwe & saayman 2008). therefore, income generated by tourism is used to manage and conserve these natural areas, as government funding for the national parks in south africa has been reduced over the years; the funds generated by tourism services thus provide a vital supplementary income for conservation actions (phillips 2009). creating more ways to attract tourists to the national parks will increase the parks’ revenue. however, the natural area tourism industry in south africa needs to redevelop continuously to stay up to date with current trends, ensure return visits and expand the visits of nature tourists. ‘frogging’ is a niche nature tourism product related to frogs, offering a unique attraction to the current range of south african wildlife and ecotourism products (loubser 2016; sat 2022). it describes the activity of searching for frogs in nature to admire and learn about these species and their relationship with humans, conserve the species and use this tourism product to sustain the local community’s livelihoods (carruthers & du preez 2011; sat 2022). south africa’s network of national parks represents a diversity of fauna and flora (coastal and inland), including (in many cases) a wide variety of frog species (du preez & carruthers 2017; eds. minter et al. 2004). still, these smaller and lesser-known species are sometimes neglected and ignored within national parks (fennell & weaver 1997; tisdell 2007; tolley et al. 2011). morrison et al. (2012) stated that frog tourism has significant potential to contribute to global frog conservation efforts and can be a drawcard for tourists visiting national parks or protected areas. tisdell (2007) found that the public is more willing to get actively involved in conservation efforts of a particular species once the species becomes more endangered or well known (tisdell & wilson 2012). according to the endangered wildlife trust (ewt 2018), frog species’ compositions will differ between biomes across south africa (du preez & carruthers 2017; eds. minter et al. 2004), and all the south african national parks represent the nine different biomes of south africa. creating frogging hotspot destinations in these national parks could help put frogs and their conservation plight into the public eye and generate income for sanparks (the management body of south african national parks) to manage their conservation operations. therefore, this study aims to evaluate the potential of frog tourism in south african national parks. literature background natural area tourism provides opportunities for tourists to learn about and appreciate the natural environment (newsome et al. 2013; weaver 2001). this type of tourism started because of the diversification of the tourism industry and the desire of tourists to experience wildlife and the natural environment (wearing & neil 2009). it consists of four key pillars: ecotourism, wildlife tourism, adventure tourism and geotourism (mckinney 2016; metin 2019; roxana 2012). in this study, frogging tourism is placed under the ecotourism pillar of natural area tourism as a result of its educational, conservational and community qualities (newsome et al. 2013). héctor ceballos-lascuráin (1992, 1996) defined ecotourism as visiting fragile, unspoiled, protected areas (mckinney 2016; saayman 2009). the complete experience helps in educating tourists (in this context, about frogs), provides funds for nature and cultural conservation and raises respect for the environments and cultures found in these areas (newsome et al. 2013; orams 1995; tisdell & wilson 2012; von solms & van der merwe 2020; willemen et al. 2015). ecotourism is an alternative form of tourism that involves conserving resources (biological diversity), maintaining sustainable use of resources, bringing an ecological experience to tourists, conserving the ecological environment and gaining economic benefit (mckinney 2016; saayman 2009; tisdell & wilson 2012). overall, ecotourism can be seen as a more responsible and sustainable form of tourism, if practised correctly (metin 2019; newsome et al. 2013). importance of frog conservation frogs have been under severe pressure since the industrial revolution, with almost a third of the more than 7000 known amphibian species listed as threatened by the iucn (amphibiaweb 2017; bishop et al. 2012; iucn 2017). according to the endangered wildlife trust (ewt) (2018, 2021), south africa has 135 frog species, of which 30% are considered threatened by the iucn. the ewt (2021) is currently running projects across three provinces (western cape, eastern cape and kwazulu-natal) to protect the nine most endangered frog species of south africa. an entire class of vertebrates face a mass extinction spasm, and conservation actions are needed to save them. the first and primary threats to frogs are invasive species, problematic species and gene composition, affecting 37% of all south african frog species. this approximation is noticeably higher than the global average of 15.7% (angulo, hoffmann & measey 2011). the second major threat to south african species is biological resource usage (46%), whereas pollution, together with residential and commercial development, are listed as the third threat, affecting 14% of frogs, which is almost a third of south african frog species (angulo et al. 2011; ewt 2021; harrison et al. 2000). the fourth factor affecting nearly 26% of all south african frog species comprises several natural system modification processes (fire, water quality and weather) (ewt 2018). almost 50% of all frog species in south africa are affected by agriculture and aquaculture, which result in habitat loss (angulo et al. 2011). it is imperative to protect frogs because of the ecological function they perform (ewt 2018). frogs help control insect populations, keep waterways clear and serve as a food source in ecosystems (angulo et al. 2011; du preez & carruthers 2017; west 2018). to minimise the existing amphibian extinction crisis, the global community must respond with an innovative and multidisciplinary approach to protect amphibians at an unprecedented scale (angulo et al. 2011; mittermeier, gascon & andreone 2008). unfortunately, the conservation of frogs does not receive the same attention from the public and conservation authorities as does the conservation of more charismatic endangered species such as rhinos (estren 2012). these attitudes are ascribed mainly to a lack of knowledge about frogs and to the negative connotations of most native folklore and mythology about frogs (ceriaco 2012). the question can be asked whether the public might become more involved in the conservation of frogs once it is emphasised that they are threatened and need urgent conservation action. frogging tourism can be a solution to this problem. with its thriving ecotourism industry and diverse environment, south africa and its provincial and national parks provide the ideal opportunity to study the potential of frogs within ecotourism. by involving tourists in the conservation and education of frogs, it is vital to understand the market of ecotourism and the needs of tourists properly. such information can be used to better plan and manage this supplemental frog conservation approach, thus contributing to its sustainability (angulo et al. 2011; mittermeier et al. 2008). south african national parks cover a wide area across south africa (sanparks 2020) and provide the ideal destination for frogging activities, as numerous endangered and endemic frog species occur within these national parks (eds. minter et al. 2004). the western cape hosts the highest number of endemic species in south africa (du preez & carruthers 2017; eds. minter et al. 2004), for example, the strawberry rain frog (breviceps acutirostris), cape mountain rain frog (breviceps montanus), endangered knysna leaf-folding frog (afrixalus knysnae), near-threatened cape peninsula moss frog (arthroleptella lightfooti) and the critically endangered table mountain ghost frog (heleophryne rosei); some of these species are found within national parks situated in the western cape. the kruger national park, which is the most preferred park by national and international tourists (van der merwe & saayman 2008), hosts a total of 28 known frog species (du preez & carruthers 2017; eds. minter et al. 2004), for example, the charismatic golden leaf-folding frog (afrixalus aureus), water lily frog (hyperolius pusillus) and banded rubber frog (phrynomantis bifasciatus). with their unique colours and features, these species will pose great photographic opportunities for ecotourists. frog-related tourism activities tourists constantly seek unique experiences, especially activities that bring them close to nature. frogging is a fun and easy family or individual activity that can aid in the protection of wetland habitats and the species found within these ecosystems (naturewatch 2022). tourists will learn more about these beautiful creatures’ lifestyles, calls and physical features by participating in frogging activities. a desktop study was conducted to determine the size and scope of frogging tourism, specifically in south africa. table 1 summarises the eight frog-related tourism activities found during the study. within south africa, none of these frogging destinations is found within any of the south african national parks. currently, there are no known frogging activities hosted by national parks or at least listed on their websites as an activity (sanparks 2021). the closest to this is the privately owned jock safari lodge next to the kruger national park (south african lodges.com 2022). out of the eight identified destinations listed in table 1, two are managed by national nonprofit conservation organisations of south africa (i.e. birdlife south africa and the endangered wildlife trust), one by a local nonprofit conservation organisation (matotoland eco-tourism association), two by private nonprofit conservation organisations, two by public tour operators and one by the owners of the destination. this indicates a lack of national parks to develop frogging tourism activities that can conserve frogs in south africa. table 1: south african tourism destinations hosting frogging events. research method the sampling and sample population, questionnaire and statistical analysis will be discussed in the research method section. sampling and sample population for this research, a quantitative research approach was followed, namely nonprobability sampling, to which convenience sampling was applied (mathers, fox & hunn 2007; patel 2009). the research sample population was tourists who had previously visited south african national parks. permission was obtained from the sanparks research office to conduct the investigation. the consent letter accompanying the questionnaire clearly stated that respondents should have visited south african national parks in the past to participate in the research. the questionnaire was hosted on the south african national parks website via a link to google forms. the questionnaire was accessible to the public for six weeks. the data were collected using google forms, which offered live and instant access to the feedback. for the period, 356 (n) completed questionnaires were received back. questionnaire the web-based questionnaire was developed in cooperation with the tourism research in economics, environs and society (trees) of the nwu, sanparks (2016a) and the african amphibian research group of the nwu, in line with a study performed by fricker and schonlau (2002). the questions covered various aspects of frog conservation and ecotourism to generate information on how tourists feel about participating in frog-related activities. the focus of this questionnaire was on the demographic profile of frogging tourists, motivations for frogging, views on the protection and conservation of frogs and current frog identification and call identification knowledge. a five-point likert scale was used to measure the degree to which tourists are aware of frog conservation, their perception of frogs and frog conservation, participation in frog-related ecotourism activities and their main reasons for visiting a destination. statistical analysis the data received from the questionnaires were captured in google forms and exported to a microsoft excel file format for further use and interpretation (factor analysis). an exploratory factor analysis was conducted with the kaiser–meyer–olkin (kmo) test to measure if the data from the questionnaires were suited for factor analysis (glen 2016). spss 24 (statistical package for the social sciences) was used to analyse the data. ethical considerations ethical approval to conduct this study was obtained from the ethics committee of nwu, faculty of economic and management sciences (reference numbers ems2016/11/04-0213 and ems2016/11/04-0214 [online]). results the research results are presented in five stages: firstly, the sociodemographics of respondents; secondly, frog encounters; thirdly, the importance and interest in frog conservation and activities; fourthly, the travel motives of respondents; and lastly, possible activities for frogging tourism. sociodemographics the results (table 2) indicate that the respondents were mainly female (56%), english-speaking, married and between the ages of 25 and 34 years (26%). the gauteng province in south africa was their main province of residence (41%). the most significant percentage (42%) of the respondents had a degree or diploma and were employed (54%), earning a salary higher than r552 001 per annum (32%). most respondents were south african citizens, with only 5% being international. the profile obtained from this research on travellers to national parks in south africa is similar to previous research conducted in national parks by engelbrecht (2011), kruger et al. (2018), mouton (2009) and van tonder (2012), who found that visitors to national parks are mostly married, english-speaking and residing predominantly in gauteng, with an average age of 45 years and a university or secondary qualification. table 2: sociodemographic information of the 356 participants. frog encounters in this section, respondents were asked to respond to questions on when they were first exposed to nature-based tourism, the feelings they experienced during their first frog encounter (if they had one), their preference when visiting a nature-based destination and their preferred holiday activity to participate in. these questions are needed to build the profile of potential frogging tourists to south african national parks. most respondents (56%) started visiting nature-based destinations from the age of three; only 12% of the respondents indicated that they were exposed to natural areas for the first time as an adult (older than 20 years). importance of frog conservation respondents were asked to indicate on a five-point likert scale (where ‘1’ = strongly disagree and ‘5’ = strongly agree) to what extent they agreed with a list of frog protection statements (figure 1). the following three statements obtained the highest values: frogs play an important role in nature (4.75); we must protect frogs and conserve their habitats (4.83); and future generations and your children will benefit if frogs are protected (4.68). most of the respondents were aware that frogs play an important role in nature, that they should be conserved and that people should be educated about frogs. it is thus imperative to send out the message that the global community should get involved with the conservation of frogs and that it is not only the responsibility of the research and conservation community. figure 1: conservation of and educational responses about frogs by respondents. the following results discuss the participants’ awareness and interest regarding frogs and participating in frog-related tourism activities (figure 2). questions were asked to determine whether the participants felt that frogging activities could be offered at nature-based destinations such as national parks. again, respondents had to rate their response on a five-point likert scale (where ‘1’ = not at all and ‘5’ = a great deal). looking at the mean values of each question, the answers to the following five questions stood out: do you think frogging activities should be offered at nature reserves and south african national parks? (4.4) do you think tourism, especially nature-based or ecotourism, can help conserve frogs? (4.35) if you have children or younger siblings, would you like to expose them to frog-related activities? (4.17) do you find frogs interesting? (4.13) would you like to become more educated regarding frogs? (4.07) figure 2: respondents’ awareness and attitudes towards frogs. the results show that most respondents are aware that several frog species are endangered and that frog-related tourism activities can benefit from their conservation. what also stood out is that most respondents find frogs interesting, have been educated about frogs and would like to partake in frog-related activities. when asked if respondents would travel to specific destinations for specific frogging activities, almost half of the respondents were unsure (54%) and 31% stated that they would. a significant percentage of respondents indicated that they would be willing to travel regionally (27%) to visit a frog-related tourism destination, whereas 22% would only travel locally. a total of 21% indicated that they would be willing to travel to neighbouring provinces. travel motives of potential frog tourists the following section discusses the results of the travel motives of the respondents of this research. an exploratory factor analysis was conducted with the kmo test to measure if the data from the questionnaires were suited for factor analysis (glen 2016). the cronbach’s alpha score for the kmo tests was between 0.92 and 0.7 for the various factors, which is acceptable (field 2006; glen 2016). the 22 constructs delivered (table 3) five factors: social aspects, relaxation, biodiversity, conservation and new experiences. table 3: pattern matrix and inter-item correlations for travel motives. inter-item correlations were also calculated and provided the following results (table 3): the total variance declared was above 50%, which shows an appropriate fit of the selected components (pietersen & maree 2007). the mean value of each factor was calculated as the average of all items contributing to a specific factor to interpret the mean scores of the original five-point likert scale and ranged from 3.13 to 4.26. relaxation (factor 2) and conservation (factor 4) are regarded as the most important travel motives, having a mean value of 4.21 (relaxation) and 4.26 (conservation) and a cronbach’s alpha value of 0.92 (relaxation) and 0.87 (conservation). the factors are discussed next. factor 1: socialising socialising can be described as the degree to which participants like to socialise with family, friends or other enthusiasts during activities. socialising is the least important motive for travelling, with a mean value of 3.13 and a cronbach’s alpha value of 0.82. the constructs for the social factors are the following: to make use of appealing accommodation, to meet new people and to socialise with other nature enthusiasts. these results are contradictory to a study performed by kruger and saayman (2010), in which it was observed that one of the most prominent motives for travelling is socialising. nevertheless, as it is a motive, it can be used to attract tourists to frogging destinations, which can be advertised as social events where groups of people join and go on frogging excursions (kotzé 2011; mckibbin 2017; ormond 2022). factor 2: relaxation the relaxation factor is the second most important motive for travelling amongst the respondents, with a mean value of 4.21 and a cronbach’s alpha value of 0.92. all the constructs’ values are close enough to be significant and the constructs are to relax, find relief, escape routine and experience peace and tranquillity, primarily within a nature-based destination. according to uysal, mcdonald and martin (1994) and saayman and saayman (2009), tourists travel to national parks to escape from their daily routine and to relax and enjoy the scenic beauty and wildlife that the park has to offer (kruger & saayman 2010). factor 3: biodiversity with a mean value of 3.85, biodiversity is the respondents’ third most important travel motive. it includes all the animals and plants, also known as destination attributes, which tourists can view when travelling to a specific destination. this is the first time frogs (biodiversity) were identified as a pull factor for travellers to natural areas and can therefore be used as a unique selling point for tourism. factor 4: conservation the respondents indicated that conservation is the most important factor when making travel decisions. the mean value is 4.26 and the cronbach’s alpha value is 0.87. tourists want to feel that they contribute to biodiversity conservation and practise sustainable tourism (herzl 2019). conservation is one of the primary components of ecotourism (newsome et al. 2013; tisdell & wilson 2012; von solms & van der merwe 2020), and the finding that it is a factor is important as one would like tourists to visit protected areas to support conservation, which can improve frog conservation too. factor 5: new experience the factor has a mean value of 3.74, which indicates that it is also a deciding factor when tourists make travelling plans. for potential travellers to experience new adventures, they first need to be open to the idea of doing something new (chaiken & ledgerwood 2012). this is an important finding, as frogging tourism can provide ecotourists with new experiences. frog activity respondents were asked to indicate on a five-point likert scale (where ‘1’ = not at all and ‘5’ = a great deal) to what extent they would participate in specific frog-related activities (table 4). the results show that 50% of participants were interested in doing night walks during which they could look for and listen to frogs, and 40% of participants were interested in doing a frog identification-related activity. the participants were not very keen on doing frog-related activities where entertainment for children was included or where they must watch educational films. their additional suggestions for activities included assisting with the re-establishment of frogs in nature and combining frogging trips with other nocturnal animal expeditions. the activities that obtained the highest mean values out of five are presented in the following table. table 4: the top five frogging activities represented in a likert scale. discussion this study identified frogging as a possible ecotourism product for south african national parks. numerous researchers (amakhosi 2022; gaisford 2021; khalo 2020; kotzé 2011; kouga baviaans 2022; mckibbin 2017; ormond 2022) have concluded that frog tourism and research about critically endangered frog species are important. morrison et al. (2012) indicated that frogs are much less common or popular than tourism products focusing on, for example, mammals or birds (biggs et al. 2011). furthermore, few protected areas in the world were established and explicitly managed for frog conservation and frog-related tourism. this is also the case within south african national parks. for example, mountain zebra national parks were established to protect the mountain zebra (capenature 2022; sanparks 2016b), and addo elephant national park was found to protect the addo elephants (sanparks 2015). research by morrison et al. (2012) confirmed that despite frogs not being focal species for nature-based tourism, their results indicated that tourism contributes to protecting more than half of the critically endangered frog species worldwide. therefore, the south african national parks’ management body, sanparks, should consider a process of identifying possible parks for frogging tourism. it is recommended that more scenic parks be used, for example, golden gate highlands, west coast and the garden route national park or parks where endangered frog species and nonendangered species will sell as a niche product. these parks do not offer traditional wildlife and big five products that can compete against frogging tourism. morrison et al. (2012) also indicated the need for tourism, as responsible tourism funds conservation. their study revealed that tourism represents a significant proportion of protected areas’ budgets, particularly in developing regions such as south africa. developing countries can potentially increase the contribution of tourism to protected areas by introducing new products such as frogging. in the case of south african national parks, frogging tourism can also be used to generate extra funding. from a conservation perspective, frog tourism in south african national parks must not negatively impact the environment. possible negative impacts include facilities being built to observe frogs, litter, noise, direct killing or injury, disturbance of activities of frogs and habitat alteration, to name a few (green & giese 2004). therefore, south african national parks must ensure compliance strategies and regulations are in place before developing frog tourism. the second finding of the research is that the frogging tourism market has not only similar (relaxation and socialising) but also different travel motives to those of the traditional ecotourist market to south african national parks. this is proved by the two new identified travel motives, conservation and biodiversity. when examining the research conducted on travel motives of visitors to protected areas (unrelated to frogging tourism), one sees that the primary motives are relaxation (also found in this research), socialising (also present in this research), to get away, photography of wildlife, novelty, activities, nostalgia and adventure, to name but a few (kruger & saayman 2010; meng, tepanon & uysal 2008; saayman, saayman & ferreira 2009). streicher and saayman (2010) indicated that different destinations feed different motives to travel. therefore, south african national parks cannot use the travel motives determined for previous research, for example, in kruger national park and others (kruger & saayman 2010; saayman & saayman 2009; van der merwe & saayman 2008) with a different market focus, namely the traditional ecotourist market for wildlife mammal species. consequently, the implication is that if sanparks develop frogging ecotourism products, emphasis must be placed on the education of tourists on frogging conservation, the biodiversity of these species and their habitats. education of visitors in protected areas remains a key aspect of protection; as baba dioum (1968) stated, ‘in the end, we will conserve only what we love, we will love only what we understand and we will understand only what we are taught’. the last finding is the identification of potential activities for frogging ecotourism. night walks, drives and frog identification were essential activities (table 4). this research concurs with menbere and admassu (2020) and sánchez-prieto et al. (2021), who identified nature walks, hiking and night drives as important ecotourism activities. these activities can easily be introduced into some of the parks mentioned here as a niche market, such as birding, where people travel thousands of miles to see certain species (u.s. fish and wildlife service 2011). it can further assist in new skills development for south african national parks, as field guides will be needed to guide and educate tourists on frogging. field guides can be recruited from communities adjacent to these protected areas, which will assist in poverty alleviation and skills development in these communities. research by saayman et al. (2009) determining the socio-economic impact of karoo national park on beaufort west communities found that national parks benefit adjacent communities in various ways, such as employment and upliftment of communities. the element of activities is essential. tourism-related activities in national parks have different roles: firstly, a conservation role; secondly, an educational role to educate visitors on specific animals or species (in this case, frogs); thirdly, to generate income; and lastly, to support the existence of protected areas. it is well known that south african national parks’ primary income source lies in tourism-related activities (sanparks 2020). research by bunghez (2016:8) expressed that the sum of tourism commitment to a protected area’s economy depends mainly on the activities provided at the destination. conclusions this research aimed to evaluate the potential of frog tourism in south african national parks. south african national parks are spread all over the country and already host most of the frog species found in south africa (du preez & carruthers, 2017; eds. minter et al. 2004), therefore making it favourable for frogging tourism. the main contribution of this novel research is that this research identified frogging tourism as a potential tourism product in south african national parks. this was also the first time research in this regard was conducted in south african national parks. acknowledgements the authors would like to acknowledge and thank dr l. slabbert from sanparks for assistance in permission to carry out this study and prof. ché weldon for his contribution in serving as study leader for the duration of the project’s development. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions z.l. is the master’s student (currently phd student) whose research was published and involved in the draft article. p.v.m. is the study leader, who was responsible for writing, review and editing and who wrote the findings and implications. funding information this research received no specific grant from any funding agency in the public, commercial or not-for-profit sectors. data availability no data are available, as they belong to nwu. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references amakhosi, 2022, amakhosi safari lodge: frogging safaris, viewed 28 january 2022, from https://www.amakhosi.com/activities. amphibiaweb, 2017, amphibiaweb: species of the week, viewed 28 october 2017, from https://amphibiaweb.org/. angulo, a., hoffmann, m. & measey, g.j., 2011, ‘introduction: conservation assessments of the amphibians of south africa and the world’, in g.j. measey (eds), ensuring a future for south africa’s frogs: a strategy for conservation research, sanbi biodiversity series 19, pp. 1–9, south african national biodiversity institute, pretoria. biggs, d., turpie, j., fabricius, c. & spenceley, a., 2011, ‘the value of avitourism for conservation and job creation – an analysis from south africa’, conservation and society 9(1), 80–90. http://doi.org/10.4103/0972-4923.79198 bishop, p.j., angulo, a., lewis, j.p., moore, r.d., rabb, g.b. & garcia moreno, j., 2012, ‘the amphibian extinction crisis – what will it take to put the action into the amphibian conservation action plan?’, s.a.p.i.e.n.s iucn commissions 5(2), 97–111. brett, r.m., 2018, ‘tourism in the kruger national park: past development, present determinants, and future constraints’, african journal of hospitality, tourism and leisure 7(2), 1–28. bunghez, c.l., 2016, ‘the importance of tourism to a destination’s economy’, journal of eastern europe research in business and economics 1, 1–9. https://doi.org/10.5171/2016.1434495 capenature, 2022, cape mountain zebra project, viewed 15 june 2022, from https://www.capenature.co.za/projects/cape-mountain-zebra-project. carruthers, v. & du preez, l., 2011, frogs & frogging in south africa, p. 112, penguin random house, struik nature, cape town. ceballos-lascuráin, h. (ed.), 1992, ‘tourism, ecotourism, and protected areas: the state of nature-based tourism around the world and guidelines for its development’, iucn world conference on national parks and protected areas, island press, pp. 1–301, caracas, venezuela, february 10–21, 1992. ceballos-lascuráin, h., 1996, tourism, ecotourism, and protected areas: the state of nature-based tourism around the world and guidelines for its development, p. 301, iucn publications, cambridge. ceriaco, l.m.p., 2012, ‘human attitudes towards herpetofauna: the influence of cultural beliefs and negative values on the conservation of amphibians and reptiles in portugal’, journal of ethnobiology and ethnomedicine 8, 8. https://doi.org/10.1186/1746-4269-8-8 chaiken, s. & ledgerwood, a., 2012, ‘a theory of heuristic and systematic information processing’, in a. paul, m. van lange, a.w. kruglanski, & e.t. higgins (eds), the handbook of theories of social psychology, pp. 246–267, sage, thousand oaks, ca. dioum, b., 1968, paper presented at the general assembly of the international union for the conservation of nature and natural resources, seattle public library archive, new delhi. du preez, l. & carruthers, v., 2017, frogs of southern africa, a complete guide, p. 448, struik nature publishers, cape town. endangered wildlife trust (ewt), 2018, frogs in the classroom: frog habitats, viewed 13 may 2021, from https://www.ewt.org.za/wp-content/uploads/2020/05/frogs-in-the-classroom_-habitats-2018.pdf. endangered wildlife trust (ewt), 2021, threatened amphibian programme: saving habitats, viewed 13 may 2021, from https://www.ewt.org.za/what-we-do/conserving-habitats/threatened-amphibian-programme/. engelbrecht, w.h., 2011, ‘critical success factors for management the visitor experience at kruger national park’, ma thesis, north-west university potchefstroom, potchefstroom. estren, m.j., 2012, ‘seeking respect for non-cute’, journal of animal ethics 2(1), 6–11. https://doi.org/10.5406/janimalethics.2.1.0006 fennell, d.a. & weaver, d.b., 1997, ‘vacation farms and ecotourism in saskatchewan, canada’, journal of rural studies 13(4), 467–475. https://doi.org/10.1016/s0743-0167(97)00032-6 field, a., 2006, discovering statistics using spss, 2nd edn., p. 779, sage, thousand oaks, ca. fricker, r.d. & schonlau, m., 2002, ‘advantages and disadvantages of internet research surveys: evidence from the literature’, rand: field methods 14(4), 347–367. https://doi.org/10.1177/152582202237725 gaisford, l. 2021, fabulous frogging after the recent rains, viewed 28 january 2022, from https://www.durbantv.net/fabulous-frogging-after-the-recent-rains/. glen, s., 2016, kaiser-meyer-olkin (kmo) test for sampling adequacy, viewed 02 november 2017, from http://www.statisticshowto.com/kaiser-meyer-olkin/. green, r. & giese, m., 2004, ‘negative effects of wildlife tourism on wildlife’, in k. higginbottom (eds.), wildlife tourism impacts, management and planning, pp. 81–97, common ground publishing, australia. harrison, j.a., burger, m., minter, l.r., de villiers, a.l., baard, e.h.w., scott, e., et al., 2000, conservation assessment and management plan for southern african frogs: final report, p. 202, iucn/ssc conservation breeding specialist group, apple valley, ma. herzl, r., 2019, ethical ecotourism is good for conservation, viewed 23 february 2022, from https://wildnet.org/ethical-ecotourism-is-good-for-conservation/. international union for conservation of nature (iucn), 2017, the iucn red list of threatened species, viewed 20 june 2017, from http://www.iucnredlist.org/initiatives/amphibians/analysis. khalo, j., 2020, kenilworth racecourse conservation area (krca): thrilling guided frog walks, viewed 28 january 2022, from https://natureconnect.earth/2020/11/02/thrilling-guided-frog-walks/. kotzé, p., 2011, ‘hands-on education a hit in biodiversity hotspot: aquatic environment’, the water wheel 10(2), 30–34. kouga baviaans, 2022, frog safari st. francis bay, viewed 28 january 2022, from https://www.surfandsafari.co.za/listing/frog-safari. kruger, m. & saayman, m., 2010, ‘travel motivation of tourists to kruger and tsitsikamma national parks: a comparative study’, south african journal of wildlife research 40(1), 93–102. https://doi.org/10.3957/056.040.0106 kruger, m., van der merwe, p., bosch, z.j. & saayman, m., 2018, ‘adventure activity preferences in south african national parks’, south african journal for research in sport, physical education and recreation 40(1), 1–23. loubser, c., 2016, 10 fascinating facts about frogging and frogs, viewed 02 march 2022, from https://wilderness-safaris.com/blog/posts/10-fascinating-facts-about-frogging-and-frogs. mathers, n., fox, n. & hunn, a., 2007, surveys and questionnaires, p. 48, nihr rds, yorkshire. mckibbin, r., 2017, birdlife south africa; frogging, ‘herping’ and rock-flipping in wakkerstroom, viewed 28 january 2022, from https://www.birdlife.org.za/frogging-herping-and-rock-flipping-in-wakkerstroom/. mckinney, t., 2016, ‘ecotourism’, in a. fuentes (ed.), the international encyclopaedia of primatology, john wiley & sons, inc, hoboken, nj. menbere, i.p. & admassu, f., 2020, ‘challenges and opportunities for ecotourism development: a case study in dilla university botanical and ecotourism garden, south ethiopia’, global journal of ecology 5(1), 154–163. https://doi.org/10.17352/gje.000035 meng, f., tepanon, y. & uysal, m., 2008, ‘measuring tourist satisfaction by attribute and motivation: the case of a nature-based resort’, journal of vacation marketing 14(1), 41–56. https://doi.org/10.1177%2f1356766707084218 metin, t., 2019, nature-based tourism, nature based tourism destinations’ attributes and nature based tourists’ motivations, viewed 01 march 2022, from https://www.researchgate.net/publication/331982531_nature-based_tourism_nature_based_tourism_destinations’_attributes_and_nature_based_tourists’_motivations. minter, l.r., burger, m., harrison, j.a., braack, h.h., bishop, p.j. & kloepfer, d. (eds.), 2004, atlas and red data book of the frogs of south africa, lesotho and swaziland, p. 360, smithsonian institution press, washington, dc. mittermeier, r.a., gascon, c. & andreone, f., 2008, a conservation strategy for the amphibians of madagascar: foreword – monografie xlv, p. 12, museo regionale di scienze, torino. morrison, c., simpkins, c., castley, j.g. & buckley, r.c., 2012, ‘tourism and the conservation of critically endangered frogs’, plos one 7(9), e43757. https://doi.org/10.1371/journal.pone.0043757 mouton, m.e., 2009, ‘socio-economic impact of an urban park: the case of wilderness national park’, ma thesis, north-west university potchefstroom, potchefstroom. naturewatch, 2022, frogwatch: engaging citizens in science, viewed 28 january 2022, from https://www.naturewatch.ca/frogwatch/. newsome, d., moore, s. & dowling, r., 2013, natural area tourism: ecology, impacts and management, 2nd edn., channel view publications, bristol. orams, m.b., 1995, ‘towards a more desirable form of ecotourism’, tourism management 16(1), 3–8. https://doi.org/10.1016/0261-5177(94)00001-q ormond, m., 2022, a friends of kloofendal nature reserve event: frog evening, viewed 28 january 2022, from https://kloofendalfriends.org.za/frog-evening/. patel, p., 2009, introduction to quantitative methods, viewed 28 september 2017, from http://hls.harvard.edu/content/uploads/2011/12/quantitative_methods.pdf. phillips, g., 2009, ‘current status of south african national parks’, paper presented at third annual international conference of tourism competence network (icnt), october 22, 2009, potchefstroom. pietersen, j. & maree, k., 2007, ‘statistical analysis ii: inferential statistics’, in k. maree (ed.), first steps in research, pp. 197–213, van schaik, pretoria. roxana, d.m., 2012, ‘considerations about ecotourism and nature-based tourism-realities and perspectives’, international journal of academic research in economics and management sciences 1(5), 215. saayman, m., 2009, ecotourism: getting back to basics, leisure consultants and publications, potchefstroom. saayman, m. & saayman, a., 2009, ‘why travel motivations and socio-demographics matter in managing a national park’, koedoe 51(1), 381. https://doi.org/10.4102/koedoe.v51i1.381 saayman, m., saayman, a. & ferreira, m., 2009, ‘the socio-economic impact of the karoo national park’, koedoe: african protected area conservation and science 51(1), 26–35. https://doi.org/10.4102/koedoe.v51i1.158 sánchez-prieto, m.c., luna-gonzález, a., espinoza-tenorio, a., & gonzález-ocampo, h.a., 2021, ‘planning ecotourism in coastal protected areas: projecting temporal management scenarios’, sustainability 13(14), 7528. https://doi.org/10.3390/su13147528 sanparks, 2015, addo elephant national park: park management plan, viewed 27 june 2022, from https://www.sanparks.org/assets/docs/conservation/park_man/aenp_plan.pdf sanparks, 2016a, frogging – a new activity in south africa’s national parks?, viewed 3 november 2021, from https://www.tourismupdate.co.za/article/frogging-new-activity-south-africas-national-parks?page=10. sanparks, 2016b, mountain zebra national park: park management plan, viewed 27 june 2022, from https://www.sanparks.org/assets/docs/conservation/park_man/mznp-plan.pdf sanparks, 2020, south african national parks annual report 2019/20, viewed 19 january 2022, from https://www.sanparks.org/assets/docs/general/annual-report-2020.pdf. sanparks, 2021, south african national parks, viewed 19 january 2022, from https://www.sanparks.org/. south africa, department of environmental affairs and tourism (deat), 2008, annual report 2007/2008, viewed 31 may 2017, from http://www.deat.gov.zal/documents/previousannuaireports/2008jan7/annualreview. south african lodges.com, 2022, jock safari lodge, viewed 28 january 2022, from https://www.south-african-lodges.com/lodges/jock-safari-lodge/. south african tourism (sat), 2019a, south african tourism annual report 2018/2019, viewed 02 november 2020, from https://nationalgovernment.co.za/entity_annual/1979/2019-south-african-tourism-annual-report.pdf. south african tourism (sat), 2019b, tourism performance report: january-december 2019, viewed 08 february 2022, from https://live.southafrica.net/media/276660/tourism-perfomance-report-2019_v12062020.pdf?downloadid=333390. south african tourism (sat), 2022, frogging safaris: saving amphibians from croaking, viewed 28 january 2022, from https://www.southafrica.net/gl/en/travel/article/frogging-safaris-in-south-africa. streicher, h. & saayman, m., 2010, ‘travel motives of participants in the cape argus pick n pay cycle tour’, south african journal for research in sport liggaamlike opvoedkunde en ontspanning 32(1), 121–132. https://doi.org/10.4314/sajrs.v32i1.54105 thomas, l. & middleton, j., 2003, guidelines for management planning of protected areas, p. 79, iucn, cambridge. tisdell, c., 2007, ‘valuing the otago peninsula: the economic benefits of conservation’, working papers on economics, ecology and environment, p. 145, school of economics, university of queensland. tisdell, c.a. & wilson, c., 2012, nature-based tourism and conservation: new economic insights and case studies, p. 520, edward elgar publishing, cheltenham. tolley, k.a., minter, l.r., harvey, j., tarrant, j. & measey, g.j. 2011, ‘education, awareness and capacity building’, in g.j. measey (ed.), ensuring a future for south africa’s frogs: a strategy for conservation research, sanbi biodiversity series 19, pp. 37–41, south african national biodiversity institute, pretoria. u.s. fish and wildlife service, 2018, 2011 national survey of fishing, hunting, and wildlife-associated recreation, p. 82, create space, texas. uysal, m., mcdonald, c.d. & martin, b.s, 1994, ‘australian visitors to us national parks and natural areas’, international journal of contemporary hospitality management 6(3), 18–24. https://doi.org/10.1108/09596119410059209 van der merwe, p. & saayman, m., 2008, ‘travel motivations of tourists visiting kruger national park’, koedoe: african protected area conservation and science 50(1), 154–159. https://doi.org/10.4102/koedoe.v50i1.140 von solms, w. & van der merwe, p., 2020, ‘farm size and its impact on land use: the case of the south african private wildlife industry’, open agriculture 5(1), 844–856. https://doi.org/10.1515/opag-2020-0081 van tonder, c., 2012, ‘determining tourists’ valuation of the big five’, ma thesis, north-west university potchefstroom. wearing, s. & neil, j., 2009, ecotourism: impact, potentials and possibilities, 2nd edn., elsevier, oxford. weaver, d.b., 2001, ‘ecotourism as mass tourism: contradiction or reality?’, cornell hospitality quarterly 42(2), 104–112. https://doi.org/10.1016/s0010-8804(01)80022-7 west, j., 2018, ‘importance of amphibians: a synthesis of their environmental functions, benefits to humans, and need for conservation’, honors thesis, bridgewater state university, bridgewater, ma. willemen, l., cottam, a., drakou, e. & burgess, n., 2015., ‘using social media to measure the contribution of red list species to the nature-based tourism potential of african protected areas’, plos one 10(6), e0129785. https://doi.org/10.1371/journal.pone.0129785 filelist convert a pdf file! page 1 filelist convert a pdf file! page 1 abstract introduction research methods and design results discussion conclusion acknowledgements references about the author(s) leana rossouw veterinary wildlife services, south african national parks, skukuza, south africa carin boshoff department of biomedical sciences, tshwane university of technology, pretoria, south africa claude sabeta oie rabies reference laboratory, agriculture research council-onderstepoort veterinary institute, pretoria, south africa johann kotzé msd animal health malelane research unit, malelane, south africa citation rossouw, l., boshoff, c., sabeta, c. & kotzé, j., 2021, ‘a preliminary investigation of exposure to rabies virus in selected wildlife in the kruger national park, south africa’, koedoe 63(1), a1651. https://doi.org/10.4102/koedoe.v63i1.1651 original research a preliminary investigation of exposure to rabies virus in selected wildlife in the kruger national park, south africa leana rossouw, carin boshoff, claude sabeta, johann kotzé received: 22 july 2020; accepted: 24 jan. 2021; published: 12 mar. 2021 copyright: © 2021. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract rabies is a zoonotic disease caused by members of the genus lyssavirus and causes fatal encephalitis in warm-blooded vertebrates. rabies has been previously confirmed in domestic dog populations in close proximity to the kruger national park (knp) and can potentially threaten conservation efforts. domestic dogs infected with rabies virus occasionally enter the knp and may be a source of rabies exposure to wildlife. therefore, the aim of this study was to determine if wild carnivores in the knp have been exposed to rabies virus, based on the presence of antibodies. serum samples from the african wild dog (lycaon pictus), spotted hyena (crocuta crocuta), lion (panthera leo), leopard (panthera pardus) and banded mongoose (mungos mungo) were tested for the presence of rabies-specific antibodies using the biopro enzyme-linked immunoassay kit (biopro elisa kit). selected sera were tested in parallel with the fluorescent antibody virus neutralisation test (favnt). of the 168 carnivore serum samples screened, eight (4.8%) had a percentage blocking (pb) ≥ 40, indicating the presence of rabies-binding antibodies and confirmed with the favnt to be very low levels of rabies virus neutralising antibodies (range 0.00 iu/ml – 0.22 iu/ml). rabies-binding antibodies detected by the biopro elisa kit and rabies virus neutralising antibodies shown by the favnt should however be interpreted with caution because of the lack of validation and species-specific cut-off values for wild carnivores. conservation implications: the results of this study will assist in understanding the epidemiology of rabies in the knp carnivores, especially exposure risk. the use of rabies diagnostic tools developed for domestic animals for disease surveillance in the knp carnivores was also evaluated and the outcomes will further support research on rabies in free-ranging wildlife populations. keywords: rabies; biopro elisa; antibody detection; conservation; virology. introduction rabies is a disease caused by members of the genus lyssavirus in the family rhabdoviridae. the disease is fatal to humans and other warm-blooded vertebrates, following infection, once clinical signs are apparent. late in infection rabies virus is found in the saliva and is usually transmitted by the bite of an infected animal (swanepoel 2004). in south africa, two genetically distinct and epizootiological groups of rabies virus are found, namely the canid rabies biotype and mongoose rabies biotype (mansvelt 1962; nel et al. 1997). the first outbreak of rabies in south africa occurred at port elizabeth (in the eastern cape province) in 1893 involving domestic dogs (canis familiaris), domestic cats (felis catus) and a few cattle (bos taurus) (swanepoel et al. 1993). in 1928, rabies was confirmed in two human cases who had incidentally been bitten by a yellow mongoose (cynictis penicillata) (herzenberg 1928). in 1950, rabies was identified in black-backed jackals (canis mesomelas) and cattle (bos taurus), the disease spread from the northern border district of the limpopo province, and by 1952, was confirmed in mozambique, swaziland and mpumalanga province (swanepoel et al. 1993). in 2008, an outbreak in dogs occurred in the mpumalanga province and subsequently spread to areas surrounding the kruger national park (knp) (mkhize et al. 2010). densely populated towns and villages around the knp are associated with a high number of domestic dogs that enter and interact with wildlife within the boundaries of the knp (grover 2015). cases of domestic dogs with rabies have been reported entering the knp (l. de klerk-lorist [state veterinarian skukuza, daff] pers. comm., 2014). all of these positive cases were confirmed with the fluorescent antibody test (fat) on brain tissue samples (agriculture council-onderstepoort veterinary research [arc-ovr] reference: 2014-d-8316; 2014-d12860; 2014-d-17045; 2014-d-8312; 2014-d-15228). in 2013, an african wild dog (lycaon pictus) (awd) from a pack that was previously seen interacting with a domestic dog (canis familiaris) was tested for exposure to rabies using a fluorescent antibody virus neutralisation test (favnt). this animal had a rabies virus neutralising antibody titre of 0.5 international units per millilitre (iu/ml), which is considered to be adequate seroconversion against immunisation with a rabies vaccine. as awds have not yet been routinely vaccinated against rabies virus in the knp, this result suggested exposure of the carnivore to a lyssavirus. the extent and nature of rabies exposure in the knp wildlife is currently unknown, and the most recent research conducted was in 1995 and 2012. van heerden et al. (1995) investigated the health and disease status of awd; 31 serum samples were tested with a blocking enzyme-linked immunoassay (elisa) method and no rabies-binding antibodies were detected. prager et al. (2012) tested another 26 awd using a rapid fluorescent focus inhibition test (rffit), and no rabies neutralising antibodies were detected. many densely populated towns and villages exist adjacent to the borders of the knp, which host numerous owned and free-roaming domestic dogs (conan et al. 2015). in 2008, a rabies outbreak occurred in the nkomazi local municipality (southern boundary of the knp), spreading westwards to the city of mbombela and northwards to bushbuckridge (western boundary of the knp) and other areas surrounding the knp (mkhize et al. 2010). from 2006 until 2016, a total of 586 domestic dog positive rabies cases were laboratory-confirmed, for both limpopoand mpumalanga-province municipality bordering knp (daff n.d.). between 2009 and 2014, grover (2015) investigated rabies occurrence in both wildlife and domestic animals in private game reserves bordering the knp and the bushbuckridge municipality and found that 42.4% of animals sampled tested positive for rabies (344 animals of 28 different species were sampled, of which 236 were domestic dogs). as the knp is surrounded by a high population of domestic dogs with known rabies status, research investigating the exposure to the knp wildlife is important. exposure can be detected by the presence of serum antibodies in unvaccinated animals. several serological methods are available; one such method detects rabies-binding antibodies, such as the elisa test, and another method detects rabies neutralising antibodies, such as the favnt. test performance is expressed by sensitivity and specificity values, using a recognised reference or gold standard method (altman & bland 1994). the favnt is the gold standard method for detection of rabies neutralising antibodies, with high levels of sensitivity and specificity (cliquet, aubert & sagné 1998). although the favnt is considered the gold standard, it requires specialised reagents, equipment and containment facilities to handle virus; therefore, laboratories prefer an elisa platform for serological testing. the biopro enzyme-linked immunoassay kit (biopro elisa kit) is a blocking elisa method that has been used to detect rabies antibodies in domestic dogs and cats (wasniewski & cliquet 2012) and wild carnivores, including red foxes (vulpes vulpes) and raccoon dogs (nyctereutes procyonoides) (wasniewski et al. 2013). therefore, the aim of this preliminary study was to detect potential exposure of certain wildlife carnivore species to rabies virus in the knp using serological test methods. research methods and design sample collection a total of 168 serum samples were obtained from the veterinary wildlife services (vws) biobank in the knp. samples were selected on the basis of species, age and sample collection date. preference was given to lion, spotted hyena, leopard, awd and banded mongoose samples. sera from the older individuals within each species were chosen with the assumption that the opportunity for rabies exposure would increase with age. samples with collection dates after 2009 were used because a rabies outbreak in domestic dogs occurred in 2009 in the surrounding areas of the knp (mkhize et al. 2010). geographical location of where the animals were sampled was taken into consideration in order to concentrate on animals sampled closer to villages and towns bordering the knp. however, because of the limited samples available in the biobank, this cannot be considered a preference. as a positive control, seven serum samples from previously vaccinated awd, lion and leopard were received from the biobank of the national zoological gardens of south africa (nzg) (pretoria). enzyme-linked immunoassay the biopro elisa kit is a blocking elisa test for detection of rabies glycoprotein g antibodies in fox sera (mojžiš, korytár jerg 2008). serum samples, stored at –80 °c, were thawed and heat inactivated for 30 min at 56 °c to reduce possible background that may interfere with product formation thereby ensuring more consistent results (namekar et al. 2012). a 96-well microplate layout template was designed to include positive, negative and test kit controls according to manufacturer’s instruction. test sera were diluted 1:1 as prescribed in the manufacturer manual using the diluent provided. test sera (100 µl/well) were added to the microtitre plates pre-coated with rabies antigen, the plates were incubated at 2 °c to 8 °c with gentle shaking overnight (18 h to 24 h). positive and negative controls and 100 µl of control sera provided in the kit were added to designated wells. plates were washed six times using the washing solution provided (1:10 dilution), afterwards 100 µl biotinylated anti-rabies antibody diluted 1:100 was added to each well and plates were incubated for 30 min at 37 °c with gentle shaking. plates were washed four times with diluted washing solution and 100 µl streptavidin peroxidase conjugate, diluted 1:100, was added to each well and incubated for 30 min at 37 °c with gentle shaking. tetramethylbenzidine (tmb) substrate (100 µl/well) was added to the test plates, and the plates were placed in a box, to avoid direct light and incubated for 15 min to 30 min at room temperature with gentle shaking. after incubation, 50 µl of the supplied stop solution was added to each well and the optical density (od) was measured at a wavelength of 450 nm using an elisa microplate reader (original multiskan ex; labsystems inc.). the elisa results were calculated by using the od measurement obtained from the elisa microplate reader and presented as a percentage of blocking (pb) value. according to the manufacturer’s instructions, samples with a pb < 40 are considered negative for rabies-binding antibodies and samples with pb ≥ 40 are considered to have rabies-specific antibodies present. samples with a pb ≥ 70 are considered positive with an equivalent antibody level of ≥ 0.5 iu/ml (o.k. servis biopro s.r.o 2014). fluorescent antibody virus neutralisation test rabies virus neutralising antibody titres were determined at the oie rabies reference laboratory (arc-ovr) using the favnt as described previously (cliquet et al. 1998). the serum samples for favnt were selected based on the results obtained from the biopro elisa kit. as the samples used in this study were primarily from unvaccinated animals, very low levels of antibodies, if any were expected to be present and therefore any samples with a pb of ≥ 35 were tested using the favnt. serum samples were thawed and heat-inactivated for 30 min at 56 °c and thereafter kept at 4 °c to 6 °c. all the works were carried out with sterile techniques in a biosafety cabinet (bsc, baker, united states of america), and appropriate personal protective equipment was worn at all times. the favnt was conducted in 96-well microtitre tissue culture plates, using challenge rabies virus (cvs), atcc® vr-959™cvs-11 strain (anses, france) and baby hamster kidney cells. the cvs stock was diluted to a working concentration of 100tcid50/50 µl. each test included an oie positive serum sample prepared in dogs and a negative control reference serum (both procured from anses, france). the positive control serum is an oie positive reference serum diluted to 0.5 iu/ml, the negative control serum is a pool of naïve laboratory dog sera (oie reference laboratory for rabies, nancy, france). dulbecco’s modified eagle’s medium (dmem)/nutrient mixture f12 supplemented with 10% heat-inactivated foetal bovine serum (fbs) and antibiotic antimycotic solution (10 000 units penicillin, 10 µg amphotericin b per ml [sigma-aldrich catalogue number a5955, united states of america) was used for diluting the positive and negative controls, cvs and test sera. one hundred microlitres of reconstituted dmem/f12 was added to all the wells of the control plate and all the wells for the test sera. fifty microlitres of each test serum sample, as well as both positive and negative control sera were added into four consecutive wells, and then serially diluted from 1:3 to 1:432. following a virus back-titration (in the control plate), 50 µl of the diluted cvs was added to each well (with the exception of the cell control), after which the plates were incubated at 37 °c in an incubator with 5% co2 for 1 h. after incubation, 50 µl of bhk cell suspension (with approximately 4 × 105 cells/ml) were added to each well and the plates were incubated for 48 h at 37 °c in 5% co2. at the end of the incubation, the spent medium from the plates was discarded into a virucidal solution (f10, health and hygiene, south africa), and the plates were lightly inverted on absorbent towel to drain off residual media. the plates were then fixed by adding 200 µl of 80% cold acetone to each well, and left to stand for 10 min. the used acetone was discarded into a virucidal solution and the plates were left to dry (maximum 5 min). after the plates had dried sufficiently, 50 µl of fluorescein isothiocyanate (fitc) anti-rabies monoclonal globulin (light diagnostics tm rabies dfa reagent 5100, emd millipore corp., united states of america), diluted at 1:100, was added to each well. the plates were incubated for 45 min in a humidified environment at 37 °c. the contents of the plates were discarded and the plates were rinsed twice with phosphate buffer saline (pbs) ph 7.2. excess pbs was removed by briefly inverting the plates on absorbent paper and the total area of each well was observed for the presence of fluorescence using a fluorescent microscope. if no fluorescence was observed, the well was considered to be negative, and if one or more fluorescing particles were seen, the well was considered to be positive for fluorescence. the logd50 titres of the cvs titration, the naïve reference serum, the positive reference serum and the test sera were calculated according to the spearman–kärber (1931). data analysis and statistical methods data from carnivores tested for rabies antibodies were handled in microsoft excel 2007 using a spreadsheet, recording the following variables: vws laboratory number, date of sampling, species, microchip number (if applicable), age class (old adult [oad], adult, young adult [yad] and juvenile [juv]), sex (male [m] and female [f]), place of capture (according to the knp designated sections), gps (global positioning system) of sampling point, elisa plate, od, elisa pb result, favnt titration dilution result, favnt result in iu/ml and distance to closest community. distance between sampling points and nearest towns and villages were determined through gdistance function of rgeos package (bivand & rundel 2017), using community point reference from chief surveyor general, 1:50 000 topographic layers (including farm parcel boundaries), 2008. a distribution map was compiled using arcgis desktop (esri 2017) using the gps position of sampling with the distinguishing between different species and pb results of the elisa, where pb ≥ 40 is positive. wilcoxon rank-sum test (one-sided) was run to compare vaccinated and unvaccinated pb results. statistical analyses were carried out with r core team software (r core team 2017). data were analysed by logistic regressing using a generalised linear model (glm) to determine an association between animals with a positive rabies-binding antibody result (pb ≥ 40) for any variables that might affect positive serology. the following variables were assessed: species, age, year, month, place of capture and distance to closest community (formula = status ~ distance to closest community + species + age + sex + year + month + place). a p-value < 0.05 was considered indicative of a statistically significant association. ethical consideration ethical approval to conduct the study was obtained from the tshwane university of technology, tut senate committee for research ethics (reference no. arec2016/03/001). results all carnivores with a rabies vaccination history, the positive control group, showed an elisa pb ≥ 70 and a favnt result ≥ 0.5 iu/ml (table 1). results of the elisa with a pb ≥ 70 are considered positive with an equivalent antibody level ≥ 0.5 iu/ml (o.k. servis biopro s.r.o 2014). according to the oie and who, a titre of 0.5 iu/ml is the minimum neutralising antibody titre required for a level of immunity, which allows the host to protect itself against rabies infection. the same measurement is used to confirm a satisfactory response to vaccination (oie 2013). table 1: enzyme-linked immunoassay and fluorescent antibody virus neutralisation test results of carnivores with a rabies vaccination history. percentage blocking less than 40 is considered negative for rabies-binding antibodies and samples with pb ≥ 40 are considered to have rabies-specific antibodies present. samples with a pb ≥ 70 are considered positive with an equivalent antibody level of ≥ 0.5 iu/ml (o.k. servis biopro s.r.o 2014). a titre of 0.5 iu/ml is the minimum neutralising antibody titre required for a level of immunity that allows the host to protect itself against rabies infection. the same measurement is used to confirm a satisfactory response to vaccination (oie 2013). of the 168 carnivore serum samples tested, eight (4.8%) had results with pb ≥ 40. none of the samples showed a pb value of ≥ 70 or the equivalent favnt result of ≥ 0.5 iu/ml (table 2). table 2: results of carnivores with biopro enzyme-linked immunoassay kit pb greater than or equal to 35. a total of 12 awd serum samples were tested and one (8.3%) showed pb values ≥ 40. the one sample with a pb of 42.3 had a neutralising antibody titre of 0.00 iu/ml. two awd samples with pb < 40 but ≥ 35 had neutralising antibody titre levels of 0.17 iu/ml and 0.22 iu/ml. none of the 10 banded mongoose samples showed pb values ≥ 40.a total of 25 leopard serum samples were tested and one (4%) showed a pb value ≥ 40, with a neutralising antibody titre of 0.02 iu/ml. three leopard samples with pb < 40 but ≥ 35 had neutralising antibody titre levels of 0.01 iu/ml, 0.06 iu/ml and 0.07 iu/ml. a total of 84 lion serum samples were tested and four (4.8%) showed pb values ≥ 40; these four samples had neutralising antibody titres ranging from 0.00 iu/ml 0.13 iu/ml. the two lion samples with pb < 40 but ≥ 35 both had a neutralising antibody titre of 0.17 iu/ml. a total of 37 spotted hyena serum samples were tested and two (5.4%) showed pb values ≥ 40, with neutralising antibody titre levels of 0.00 iu/ml and 0.06 iu/ml. three spotted hyena samples with pb < 40 but ≥ 35 had neutralising antibody titre levels of 0.02 iu/ml, 0.01 ui/ml and 0.06 iu/ml. of the 70 carnivores from skukuza tested, only four showed a pb ≥ 40. a total of nine carnivores from tshokwane, 10 from crocodile bridge, 12 from stolsnek and 15 from satara were tested and one carnivore in each section showed pb ≥ 40 (figure 1). figure 1: distribution of elisa results from sera of free-ranging wild carnivores in kruger national park. the wilcoxon rank-sum test (one-sided) was run to compare vaccinated and unvaccinated pb results and showed a clear distinction between vaccinated and unvaccinated carnivores (p < 0.005). interpreting a result above 40 pb as positive, a logistical regression model was built to search for any variables that might affect positive serology. variables included species, age, year, month, place of capture and distance to closest community. none of the variables evaluated had a significant impact on the outcome as evaluated by a p-value below 0.05. discussion in this study, a panel of wild carnivore serum samples was used to determine the presence of rabies-specific antibodies using a rabies elisa test to assess exposure of wildlife in knp to lyssaviruses. rabies-binding antibodies were detected in samples of awds, leopard, lions and spotted hyenas using a pb cut-off of ≥ 40. however, the eight positive carnivore serum samples on the elisa test showed little correlation to the favnt used to detect rabies virus neutralising antibodies. sera from vaccinated carnivores were subjected to both the biopro elisa kit and the favnt analyses. the results indicated that awd, lion and leopard develop high levels of rabies-binding antibodies and high titres of rabies virus neutralising antibodies. the biopro elisa kit was developed and validated as a blocking elisa test for detection of rabies glycoprotein g antibodies in fox sera and has not been validated for the african carnivore species tested in this study. the biopro elisa kit test results suggest 4.8% of the carnivores tested in this study had been exposed to a lyssavirus. one can assume that the exposure to carnivores to lyssavirus would be from rabies-positive domestic dogs from villages and towns bordering the knp. however, there was no association found between a carnivore with rabies-binding antibodies and that carnivore’s distance to the closest village or town. a carnivore closer to the community will have a higher possibility to make contact with rabies-positive domestic dogs; therefore, this could suggest that the antibodies found in the study are false positive. however, the gps location where the carnivore was immobilised for sampling might not be in close proximity to where the animal was possibly exposed to rabies. for example, lion home ranges vary between 50 km2 and 107 km2, awd between 150 km2 and 1110 km2, leopard between 14.8 km2 and 76.2 km2, spotted hyena between 25 km2 and 130 km2 and banded mongoose between 0.8 km2 and 1.0 km2 (mills & bester 2005). although findings of naturally acquired antibodies in wildlife, clinically unaffected by rabies, have also been reported in tanzania (east et al. 2001), bolivia (deem, davis & pacheco 2004), brazil (araujo et al. 2014; jorge et al. 2010), namibia (thalwitzer et al. 2010) and zambia (berentsen et al. 2013). neutralising antibody detection tests were used in these studies rather than elisa, which detect rabies-binding antibodies and therefore results are difficult to compare. the disassociation between the elisa and favnt results could also suggest an occurrence of false positive. on initial comparison, the elisa may appear to be less sensitive and specific than the favnt (cliquet et al. 1998). however, the favnt detects neutralising antibodies and the elisa detects rabies-binding antibodies (barton & campbell 1988; moore et al. 2016; vengušt et al. 2011). as the two methods detect different humoral responses, cut-off values should be independently determined and results considered separately. the elisa method detects all antibodies against rabies viral antigens coated on the elisa plate, which may include non-neutralising antibodies (vengušt et al. 2011). the elisa measures the ability of an antibody to bind to the rabies glycoprotein and parts of the antigen (epitopes) may prompt the production of non-neutralising antibodies (cliquet et al. 2003). the favnt detects biologically active serum neutralising antibodies, which may be a subset of all the binding antibodies. studies evaluating test performance of the biopro elisa kit, using favnt as the gold standard observed differing sensitivities, specificities, level of agreement and cut-off values. the biopro elisa kit has been used to detect rabies antibodies in unvaccinated domestic dogs and cats, calculating an assay specificity of 100% using 315 sera samples (wasniewski & cliquet 2012). a total of 701 sera samples from vaccinated dogs and cats were tested and a sensitivity of 84.7%, specificity of 97.6% and 86.2% level of agreement with favnt were calculated. based on the wasniewski and cliquet (2012) study, a true negative result was designated as any sample with < 0.5 iu/ml on favnt and 70 pb on the biopro elisa kit. wasniewski et al. (2013) determined the specificity of the biopro elisa kit in comparison to the favnt, using serum from 91 unvaccinated red foxes (vulpes vulpes) and 117 unvaccinated raccoon dogs (nyctereutes procyonoides). a specificity of 100% was calculated for the biopro elisa kit, using cut-off values of 40 pb and 0.24 iu/ml for the favnt. using similar cut-off values, 408 red fox sera and 274 raccoon dog sera collected from areas where oral vaccination campaigns had been conducted were tested to calculate a level of agreement between the biopro elisa kit and favnt. an overall level of agreement of 95.0% was calculated, 95.1% for red fox sera and 94.9% for sera from raccoon dogs. bedeković et al. (2016) used the biopro elisa kit for testing thoracic liquid and muscle extraction of orally vaccinated foxes to identify rabies antibodies in these samples. a total of 147 samples were tested, either as heat-inactivated or not-treated samples, using cut-off values of 0.1 iu/ml for the favnt and 40 pb for the biopro elisa kit. the biopro elisa kit had a calculated sensitivity of 93.36%, specificity of 79.1% and diagnostic viability of 94.3% using heat-inactivated sera. when non-treated sera were tested, the biopro elisa kit had a sensitivity of 79.2%, specificity of 92.98% and diagnostic viability of 89.16%. cut-off values as low as 0.1 iu/ml (jorge et al. 2010) and 0.11 iu/ml (araujo et al. 2014) have been used for neutralising antibody tests when screening for rabies exposure in wildlife. if a cut-off value of 0.2 iu/ml (berentsen et al. 2013) is used, the favnt results of this study would also suggest that there is no evidence of rabies virus neutralising antibodies in the selected carnivores from the knp. a single case of an awd with a favnt titre of 0.22 iu/ml and pb < 40 (interpreted as negative on elisa) was observed in this study. a possible explanation could be non-specific factors such as cytotoxicity or other virus neutralising activities of the sample that resulted in overestimation of rabies virus neutralising antibodies when using the favnt (barton & campbell 1988; cliquet et al. 2003). the 4.8% samples considered positive for the presence of rabies-binding antibodies by the biopro elisa kit are all low pb values in comparison to the high pb values obtained from the rabies vaccinated carnivores. whether or not this is true, evidence of rabies exposure will require further investigation. firstly, limitations of this study are the number of samples tested that represents only a very small percentage of the total population size. secondly, neither of the two serological tests has been validated for use in wildlife carnivores as was done here. another possible limitation of the study is the locality where the samples were taken, as these are not all in close proximity to the boundary of knp. therefore, in order to make future studies more meaningful, it is crucial that an extensive population-based validation of both the biopro elisa kit and the favnt be performed in order to obtain species-specific cut-off values. future research should include proactive surveillance by collecting brain samples from carnivore carcasses in order to get a more accurate picture of the impact rabies have on the knp wildlife. research to improve the non-lethal diagnostic tools we do have need to focus on finding a negative control population with zero rabies exposure, including no vaccination history and no contact with domestic animals or the possibility of contact with other rabies infected wildlife. conclusion serological testing of samples opportunistically collected from the awd, leopard, lion, spotted hyena and banded mongoose in knp showed the presence of rabies-binding antibodies in 4.8% of the samples, with very low values of rabies virus neutralising antibodies. rabies-binding antibodies detected by the biopro elisa kit should be interpreted with caution and would require further investigation. an extensive population-based validation for both the biopro elisa kit and the favnt should be performed in order to obtain species-specific cut-off values. it is recommended that continuation of surveillance be carried out to evaluate exposure to rabies virus because infection pressure may change, especially as towns and villages continue to grow along the borders of knp. for now, the surveillance would have to rely on using the validated diagnostic tools that are available (e.g. fats on tissue samples) and for future surveillance to develop the validation of serological methods, such as the ones used in this study. successful domestic dog rabies vaccination campaigns for towns and villages bordering knp should enhance protection for wildlife from rabies. although sample numbers were limited, the preliminary results suggest that rabies vaccination of awd, lion and leopard can induce high levels of protective antibodies, which can be detected using currently available assays. this is valuable information that requires further research as it may provide additional management options for future control of this disease in wild carnivores. in conclusion, this preliminary study showed that carnivores in the knp have potentially been exposed to rabies but a validation of the tests used and continued surveillance is recommended. acknowledgements the national zoological garden of south africa and south african national parks are thanked for supplying samples from their respective biobank facilities. the authors appreciate the information provided by grant beverley, senior field worker: endangered wildlife trust (ewt), scientific services, sanparks and the state veterinary services, skukuza, department of agriculture, forestry and fisheries. the authors would also like to thank the agricultural research council-onderstepoort veterinary research for utilising the rabies reference laboratory facilities. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions the publication draws substantially from the dissertation of l.r. c.b. and j.k. acted as supervisors to the dissertation. c.s. critically revised the manuscript for important intellectual content. all authors approved the final version to be published. funding information tshwane university of technology funded the publication fees and the biopro elisa kit. agricultural research council-onderstepoort veterinary research (project #p10000029) provided funding for the favnt. data availability data sharing is not applicable to this article as no new data were created or analysed in this study. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. this research was published as conference proceedings of the 10th sava veterinary and paraveterinary congress on the 16th of july 2019. references arcgis desktop version 10.5.1, 2017, computer software, esri, redlands, ca. altman, d.g. & bland, j.m., 1994, ‘diagnostic tests 1: sensitivity and specificity’, british medical journal 308(6943), 1552. araujo, d.b., martorelli, l.a., kataoka, a.p.g., campos, a.c., rodrigues, c.s., sanfilippo, l.f. et al., 2014, ‘antibodies to rabies virus in terrestrial wild mammals in native rainforest on the north coast of são paulo state, brazil’, journal of wildlife diseases 50(3), 469–477. https://doi.org/10.7589/2013-04-099 barton, l.d. & campbell, j.b., 1988, ‘measurement of rabies-specific antibodies in carnivores by an enzyme-linked immunosorbent assay’, journal of wildlife diseases 24(2), 246–258. https://doi.org/10.7589/0090-3558-24.2.246 bedeković, t., šimić, i., krešić, n., lojkić, i., mihaljević, ž., sučec, i. et al., 2016, ‘evaluation of elisa for the detection of rabies virus antibodies from the thoracic liquid and muscle extract samples in the monitoring of fox oral vaccination campaigns’, bmc veterinary research 12(76), 1–7. https://doi.org/10.1186/s12917-016-0701-0 berentsen, a.r., dunbar, m.r., becker, m.s., m’soka, j., droge, e., sakuya, n.m. et al., 2013, ‘rabies, canine distemper, and canine parvovirus exposure in large carnivore communities from two zambian ecosystems’, vector-borne and zoonotic diseases 13(9), 643–649. https://doi.org/10.1089/vbz.2012.1233 bivand, r. & rundel, c., 2017, rgeos: interface to geometry engine – open source (‘geos’), version 0.3-26, viewed 26 march 2018, from https://cran.r-project.org/package=rgeos. cliquet, f., aubert, m. & sagné, l., 1998, ‘development of a fluorescent antibody virus neutralisation test (favn test) for the quantification of rabies neutralising antibody’, journal of immunological methods 212(1), 79–87. https://doi.org/10.1016/s0022-1759(97)00212-3 cliquet, f., muller, t., mutinelli, f., geronutti, s., brochier, b., selhorst, t. et al., 2003, ‘standardisation and establishment of a rabies elisa test in european laboratories for assessing the efficacy of oral fox vaccination campaigns’, vaccine 21, 2986–2993. https://doi.org/10.1016/s0264-410x(03)00102-6 conan, a., akerele, o., simpson, g., reininghaus, b., van rooyen, j. & knobel, d., 2015, ‘population dynamics of owned, free-roaming dogs: implications for rabies control’, plos neglected tropical diseases 9(11), e0004177. https://doi.org/10.1371/journal.pntd.0004177 daff, n.d., query on animal diseases in the rsa, viewed 03 december 2016, from http://www.nda.agric.za/vetweb/epidemiology/disease%20database/oiedata/oie_query_criteria.asp. deem, s.l., davis, r. & pacheco, l.f., 2004, ‘serologic evidence of nonfatal rabies exposure in a free-ranging oncilla (leopardus tigrinus) in cotapata national park, bolivia’, journal of wildlife diseases 40(4), 811–815. east, m.l., hofer, h., cox, j.h., wulle, u., wiik, h. & pitra, c., 2001, ‘regular exposure to rabies virus and lack of symptomatic disease in serengeti spotted hyenas’, proceedings of the national academy of sciences 98(26), 15026–15031. grover, m., 2015, ‘spatio-temporal analysis of dog ecology and rabies epidemiology at a wildlife interface in the lowveld region of south africa’, msc dissertation, department of veterinary tropical diseases, university of pretoria. herzenberg, l., 1928, ‘two cases of hydrophobia’, south african medical journal 2(24), 659–661. jorge, r.s.p., pereira, m.s., morato, r.g., scheffer, k.c., carnielijr, p., ferreira, f. et al., 2010, ‘detection of rabies virus antibodies in brazilian free-ranging wild carnivores’, journal of wildlife diseases 46(4), 1310–1315. https://doi.org/10.7589/0090-3558-46.4.1310 kärber, g., 1931, ‘beitragzurkollektiven behandlungpharmakologischer reihenversuche’ [contribution to the collective treatment of series pharmacological experiments], naunyn-schmiedebergs archiv fü rexperimentelle pathologie und pharmakologie [naunyn-schmiedeberg’s archive for experimental pathology and pharmacology] 162(4), 480–483. mansvelt, p.r., 1962, ‘rabies in south africa field control of the disease’, journal of the south african veterinary medical association 33(3), 313–219. https://hdl.handle.net/10520/aja00382809_2530 mills, m.g. & bester, m.n., 2005, ‘order carnivora’, the mammals of the southern african sub-region, pp. 412–423, cambridge university press, cape town. mkhize, g.c., ngoepe, e.c., du plessis, j.a., reininghaus, b. & sabeta, c.t., 2010, ‘re-emergence of dog rabies in mpumalanga province, south africa’, vector-borne and zoonotic diseases 10(9), 921–926. https://doi.org/10.1089/vbz.2009.0109 mojžiš, m., korytár & jerg, s., 2008, ‘development and validation of elisa test for detection of rabies anti-glycoprotein antibodies’, poster presented at the international conference on rabies in the americas (rita xix), atlanta, ga, 28th september–3rd october. moore, s.m., pralle, s., engelman, l., hartschuh, h. & smith, m., 2016, ‘rabies vaccine response measurement is assay dependent’, biologicals 44(6), 481–486. https://doi.org/10.1016/j.biologicals.2016.09.007 namekar, m., kumar, m., o’connell, m. & nerurkar, v.r., 2012, ‘effect of serum heat-inactivation and dilution on detection of anti-wnv antibodies in mice by west nile virus e-protein microsphere immunoassay’, plos one 7(9), 45851. https://doi.org/10.1371/journal.pone.0045851 nel, l., jacobs, j., jaftha, j. & mererdith, c., 1997, ‘natural spillover of a distinctly canidae-associated biotype of rabies virus into an expanded wildlife host range in southern africa’, virus genes 15(1), 79–82. https://doi.org/10.1023/a:1007979502 oie, 2013, ‘world organisation for animal health’, rabies (infection with rabies virus). o.k. servis biopro s.r.o., 2014, elisa diagnostic kit for detection of rabies antibodies, leaflet, czech republic. prager, k.c., mazet, j.a.k., munson, l., cleaveland, s., donnelly, c.a., dubovi, e.j. et al., 2012, ‘the effect of protected areas on pathogen exposure in endangered african wild dog (lycaon pictus) population’, biological conservation 150, 15–22. https://doi.org/10.1016/j.biocon.2012.03.005 r core team, 2017, r: a language and environment for statistical computing. r foundation for statistical computing, vienna, austria, version 3.4.3, https://www.r-project.org/. swanepoel, r., 2004, ‘rabies’, in j.a.w. coetzer & r.c. tustin (eds.), infectious diseases of livestock, 2nd edn., pp. 1123–1182, oxford university press, new york, ny. swanepoel, r., barnard, b.j.h., meredith, c.d., bishop, g.c., brückner, g.k., foggin, c.m. et al., 1993, ‘rabies in southern africa’, onderstepoort journal of veterinary research 60, 325–346. thalwitzer, s., wachter, b., robert, n., wibbelt, g., müller, t., lonzer, j. et al., 2010, ‘seroprevalences to viral pathogens in free-ranging and captive cheetahs (acinonyx jubatus) on namibian farmland’, clinical and vaccine immunology 17(2), 232–238. van heerden, j., mills, m.g.l., van vuuren, m.j., kelly, p.j. & dreyer, m.j., 1995, ‘an investigation into the health status and diseases of wild dogs (lycaonpictus) in the kruger national park’, journal of the south african veterinary association 66(1), 18–27. vengušt, g., hostnik, p., cerovšek, m., cilenšek, p. & malovrh, t., 2011, ‘presence of antibodies against rabies in wild boars’, acta veterinaria hungarica 59(1), 149–154. https://doi.org/10.1556/avet.59.2011.1.14 wasniewski, m. & cliquet, f., 2012, ‘evaluation of elisa for detection of rabies antibodies in domestic carnivores’, journal of virological methods 179(1), 166–175. https://doi.org/10.1016/j.jviromet.2011.10.019 wasniewski, m., guiot, a.l., schereffer, j.l., tribout, l., mähar, k. & cliquet, f., 2013, ‘evaluation of an elisa to detect rabies antibodies in orally vaccinated foxes and raccoon dogs sampled in the field’, journal of virological methods 187(2), 264–270. https://doi.org/10.1016/j.jviromet.2012.11.022 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 abstract introduction methodology results and discussion conclusion and recommendations acknowledgements references footnotes about the author(s) luke a. sandham school of geo and spatial sciences, faculty of natural and agricultural sciences, north-west university, potchefstroom, south africa carla huysamen school of geo and spatial sciences, faculty of natural and agricultural sciences, north-west university, potchefstroom, south africa francois p. retief unit for environmental sciences and management, faculty of natural and agricultural sciences, north-west university, potchefstroom, south africa angus morrison-saunders unit for environmental sciences and management, faculty of natural and agricultural sciences, north-west university, potchefstroom, south africa centre for ecosystem management, school of sciences, edith cowan university, perth, australia alan j. bond unit for environmental sciences and management, faculty of natural and agricultural sciences, north-west university, potchefstroom, south africa school of environmental sciences, faculty of sciences, university of east anglia, norwich, united kingdom jenny pope unit for environmental science and management, faculty of natural and agricultural sciences, north-west university, potchefstroom, south africa integral sustainability, perth, australia reece c. alberts unit for environmental science and management, faculty of natural and agricultural sciences, north-west university, potchefstroom, south africa citation sandham, l.a., huysamen, c., retief, f.p., morrison-saunders, a., bond, a.j., pope, j. et al., 2020, ‘evaluating environmental impact assessment report quality in south african national parks’, koedoe 62(1), a1631. https://doi.org/10.4102/koedoe.v62i1.1631 original research evaluating environmental impact assessment report quality in south african national parks luke a. sandham, carla huysamen, francois p. retief, angus morrison-saunders, alan j. bond, jenny pope, reece c. alberts received: 26 mar. 2020; accepted: 14 july 2020; published: 16 sept. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract this research evaluates environmental impact assessment (eia) report quality for a selected sample of development projects in south african national parks. it applies an adapted version of an international eia report quality review package to 24 developments within 10 national parks, across three eia regimes. the results suggest good eia report quality across all four quality review areas, with improvement over time, but also highlight particular weaknesses in terms of dealing with waste and, to a lesser extent, with significance and mitigation. to build on this research, the development of a sector-specific eia report quality review package is recommended, with more emphasis on the strategic context, waste and water-related aspects. the conclusion is that eia is well positioned to remain an important decision support instrument for developments within national parks. conservation implications: the results show that eia reports for developments in south african national parks are generally of sufficient quality for decision-making that benefits conservation. however, weaker performance regarding waste, significance determination, water-related impacts and a lack of consideration of strategic context requires a conservation-specific eia report quality review package to improve report quality in the areas of weakness and thereby increase the value of eia as an instrument for environmental governance and sustainable development in conservation areas. keywords: environmental impact assessment; report quality; protected areas; national parks; south africa. introduction environmental impact assessment (eia) has been conducted internationally and in south africa since the 1970s and is recognised as one of the most successful environmental policy interventions of the 20th century (kidd, retief & alberts 2018; morgan 2012; roos et al. 2020). originally conceived as an instrument to protect and conserve the biophysical environment, its mandate has since been broadened in many countries to also include the promotion of sustainable development more broadly (cilliers et al. 2020). today over 200 countries have some form of mandatory impact assessment system (bond et al. 2020; yang 2019). the overall aim of eia is to inform decision-making by pro-actively considering and predicting consequences of actions on the environment before decisions are made (international association for impact assessment and institute of environmental assessment [iaia] 1999). the actions for which mandatory eia is required differ between countries. in south africa, eia is required for the so-called listed activities described in the eia regulations (most recently amended in 2017). once an activity is listed, it means that actions in relation to that legally defined activity are subject to government authorisation and therefore must undergo an eia process to inform the eventual authorisation decision. in south africa, infrastructure development in protected areas has been subject to mandatory eia for more than two decades. with a rapidly increasing global population and a steadily growing middle class, protected areas are increasingly under pressure as tourist destinations (morrison-saunders et al. 2019; steffen et al. 2015). governments, especially in the global south, are understandably keen to exploit the opportunity for economic development and job creation. this is particularly true for a country such as south africa, which boasts arguably some of the best and most diverse nature-based tourism opportunities in the world (de witt, van der merwe & saayman 2014). the country has a complex system of protected areas, managed by agencies in the municipal, provincial and national spheres of government. however, the 21 national parks managed by the south african national parks (sanparks) agency are the flagship of south africa’s protected areas network. with an increasing demand for tourism and tourism-related infrastructure development in protected areas, effective functioning of existing governance mechanisms aimed at planning and implementing infrastructure in a sustainable way is essential (arrow et al. 1995; swemmer & taljaard 2011). a suite of governance mechanisms for protected areas includes strategic plans, conservation plans, park management plans, species management plans and norms and standards. many developments in protected areas in south africa trigger mandatory eia, and therefore, eia is an example of an already existing governance instrument to assist with planning and implementing development in protected areas in a responsible and sustainable manner. the output of the eia process in south africa is a so-called eia report. because the eia report provides the basis for decision-making, the quality of the content of the eia report is critical if good authorisation decisions are to be made. the underlying assumption is that good quality eia reports enhance the likelihood of achieving good decisions (alberts et al. 2020; bond et al. 2016). for this reason, the quality of eia reports has been a particular focus of international and south african eia research (barker & jones 2013; bond et al. 2018; jalava et al. 2010; sandham et al. 2008a). evaluating eia quality should not be confused with eia effectiveness or efficiency, which rely on good quality eia reports in order for eia to achieve its objectives (effectiveness) within available time and resources (efficiency) (pope et al. 2018; retief 2010). a wealth of literature exists on eia report quality in south africa for different national and regional contexts as well as for different sectors such as mining, housing, agriculture, energy and tourism (boshoff 2019; kidd et al. 2018; sandham et al. 2008a, 2008b; sandham et al. 2013b; sandham & pretorius 2008; wylie, bhattacharjee & rampedi 2019). furthermore, the quality of biodiversity specialist inputs to eia in areas with high biodiversity value has also been researched (hallatt, retief & sandham 2015; swanepoel et al. 2019). however, there has been scant overall reflection on the contribution of eia to the conservation sector and more specifically protected areas. as a first step towards gaining a better understanding of the potentially broader contribution of eia to responsible decision making in protected areas, this research aims to evaluate the quality of eia reports for a selected sample of developments in sanparks. methodology environmental impact assessment report selection this research is concerned with eias for developments within the boundaries of national parks. however, sanparks does not maintain a central database for eias conducted in national parks, which means that an accurate representative sample cannot be identified. for this reason, it was necessary to apply the so-called replication logic that relies on a general point of saturation where the addition of more reports is unlikely to change the general patterns that emerge from the data (yin 2003). based on other quality review studies identified in the literature, a sample of between 20 and 30 reports is usually sufficient to reach a valid level of saturation (hallatt et al. 2015; sandham et al. 2013a; swanepoel et al. 2019). ultimately, 24 eia reports across 10 national parks were included in this study (see figure 1). figure 1: location of national parks included in the sample of environmental impact assessment reports. the 24 eia reports included in the evaluation reflect a range of activities and development types, over different eia regimes spanning 20 years (1997–2017) (see table 1). to simplify the data analysis, the five periods of legal changes to the eia legislation over the past two decades are grouped into three regime periods, namely the eca 1997 regime, nema 2006/10 regime and the nema 2014/17 regime (see kidd et al. 2018). six reports reflect the eca 1997 regime, of which two are the so-called beefed-up scoping reports1 akin to an eia report; 10 reports reflect the nema 2006/10 regime; and eight reports reflect the nema 2014/17 regime. table 1: summary of environmental impact assessment reports included in the evaluation. environmental impact assessment report quality review package quality review packages are widely used to evaluate the quality of eia documentation such as scoping reports, environmental statements and specialist studies across different sectors and in different contexts (barker & jones 2013; canelas et al. 2005; cashmore et al. 2002; lee et al. 1999; mcgrath & bond 1997; sandham et al. 2008a, 2008b, 2013a; sandham, carrol & retief 2010). a package typically consists of different report quality review criteria against which the quality of eia reports can be evaluated. the lee and colley review package is probably the most widely applied internationally and consists of multiple criteria arranged in a four-level hierarchical structure that consists of an overall report grade, review areas, categories and sub-categories (see figure 2; lee et al. 1999). the lee and colley review package was adapted by sandham et al. (2013a) to create a generic eia report quality review package suitable for the south african eia system. as the original lee and colley review package evaluates best practice as well as eia report quality (lee et al. 1999), the adapted package retained all the original lee and colley review topics and added a number of sub-categories to cater for the unique features of the south african eia system. it was not deemed desirable for this research to further adapt the sandham et al. (2013a) review package specifically for conservation or protected areas because this would have made comparison with other report quality and longitudinal2 studies difficult. figure 2: hierarchical structure of the lee and colley environmental impact assessment report review package. the review of an eia report starts at the lowest level of the hierarchy, that is, the sub-categories that consist of simple criteria related to specific tasks and procedures for which grades are given. following the review of the sub-categories, the review categories (rc) are evaluated, followed in turn by the review areas (ra) and finally the overall eia report grade. for each level, the review is based on the review grades of the previous level. a summary description of the eia report quality review areas and criteria is provided in table 3. grades are awarded ranging from a to f, depending on how well a specific task is judged by the reviewer to have been performed (lee et al. 1999; sandham et al. 2013a; see table 2). the review relied on an inter-comparison and double reviewer approach to calibrate markers as recommended by lee et al. (1999) and põder and lukki (2011). two reviewers initially reviewed two of the reports independently, and the findings were recorded on a collation sheet using the assessment system of table 2. after completing the review independently, the results were compared; differences were identified, re-examined and discussed; and a consensus was reached. several small differences in allocated grades occurred at sub-category level, fewer at the category level, minimal differences at the level of review areas and none at the overall report level. after becoming familiar with the use of the adapted review package in the first two reports, the remaining eia reports were then reviewed by a single reviewer. ethical consideration this article followed all ethical standards for a research without direct contact with human or animal subjects. results and discussion environmental impact assessment report quality table 3 provides a summary of the overall grades, review area (ra) grades, review category (rc) and sub-category grades achieved by the 24 eia reports, i.e. without consideration of the three different eia regimes. the analysis of the overall quality of the eia reports for sanparks projects shows that 92% (n = 22) of the reports were graded as satisfactory (a–c) and 25% (n = 6) could be described as ‘well performed’ (a). however, the majority of the reports, 58% (n = 14), were ‘generally satisfactory’ (b), and 8% (n = 2) were graded as ‘just satisfactory’ (c) despite omissions and/or inadequacies. the remaining 8% (n = 2) of the reports were graded as ‘just unsatisfactory’ (d) because of omissions and/or inadequacies. none of the reports obtained an e (unsatisfactory) or f (very unsatisfactory) overall grading. table 2: list of assessment symbols. figure 3 provides a summary of the quality review results for the different review areas. the best performance was achieved in review area 4 (presentation and communication) with all of the reports being satisfactory (a–c). this was followed by review area 1 (description of project and environment) and review area 3 (alternatives and mitigation) with 96% satisfactory. however, for review area 1, 88% of the reports were graded as good (a–b grades), compared with 75% for review area 3. the weakest performance was in review area 2 (impact identification and evaluation), which had the lowest frequency of satisfactory grades, with 88% of the reports achieving a c or higher. figure 3: grades for review areas of the sample of environmental impact assessment reports in national parks. review area 1: description of the project and the environment the analysis (table 3) suggests three main conclusions: overall, projects were well described in the eia reports. this could be expected for developments in national parks because of the national and even international interest in and importance of these areas. this requires the eia report to accurately communicate the development proposal to well-informed, interested and affected parties. moreover, the types of developments such as visitor facilities, accommodation and access gates are generally not complex developments per se and are easily described. conversely, other research has shown that project descriptions are weaker for more complex and large-scale developments such as those related to mining (sandham et al. 2008a), explosives manufacturing (sandham et al. 2013b) and large developments impacting on wetlands (sandham et al. 2008b). the description of waste-related aspects was often unsatisfactorily dealt with in the eia reports, with only 46% of the reports being satisfactory (a–c) in terms of sub-category 1.3.3. the analysis found weak performance with respect to the description of the types and quantities of waste, treatment, disposal and disposal routes, and description of methods to obtain types and quantities of waste. these results suggest that eias for national parks need to significantly strengthen the consideration of waste-related aspects, in order to align with international trends of emphasising and promoting best practice waste management in protected areas (dunjić et al. 2017; hockings et al. 2006). failing in this area could result in omitting waste-related pollution impacts, which must be a critical consideration for any development in a protected area, particularly in national parks. morrison-saunders et al. (2019) showed that waste management ranks highly in terms of visitor expectations in the kruger national park. properly dealing with waste management should therefore underpin developmental thinking for protected areas. it is worth noting that dealing with waste also performed poorly in eia report quality in other sectors (see, e.g., sandham et al. 2013a, 2013b). the environment to be affected and the baseline were generally well described. this is not surprising for eias in national parks, especially in the south african context, in which there is a long history of generating scientific data and knowledge to inform conservation management. national parks in south africa are therefore data rich in terms of environmental and baseline data. table 3: summary of results: overall grades, review areas and review categories (n = 24). review area 2: impact identification and evaluation based on the quality evaluation results, the following conclusions are made: impact identification and scoping were generally well conducted. this could most likely be attributed to the perceived high profile and sensitive nature of developments in national parks, resulting in high levels of public interest and participation. this, combined with the availability of good quality data and information, seems to result in good quality impact identification and scoping, which is a typical for south african eia report quality, as public participation is not always well conducted and project impacts are not always well scoped (kidd et al. 2018). significance prediction and assessment scored the lowest of the five sub-categories in review area 2, notwithstanding that quality was found to be relatively high when compared to the more general performance with respect to impact significance in south africa (see kidd et al. 2018; sandham et al. 2013a). therefore, dealing with the prediction of impact magnitude, data to estimate the magnitude of main impacts and methods used to predict impact magnitude were generally satisfactory. moreover, the assessment of impact significance on affected communities and society in general and the justification of proposed methods of assessing significance were also satisfactory. however, two sub-categories scored significantly lower. sub-category 2.4.3 dealing with the predictions of impact in measurable quantities achieved a c score for 54% of the reports, which highlights the subjective nature of significance ratings and the difficulties in providing quantitative justification for such ratings (ehrlich & ross 2015). furthermore, sub-category 2.5.2 dealing with significance in relation to national and international quality standards produced a c grade for more than half (58%) of the reports. this may be attributed to the fact that such standards do not always exist, especially for more subjective aspects such as aesthetics, visual, heritage, sense of place and social impacts. review area 3: alternatives and mitigation analysis of the review grades lead to the following conclusions: alternatives were generally well described and considered. although the description of alternative sites in sub-category 3.1.1 had the lowest satisfactory score of the research area, it still achieved a 75% satisfactory rating overall. the location of developments in national parks is a critical consideration from a mitigation perspective, and related aspects of aesthetics, architecture and building design are highly rated considerations of visitors (morrison-saunders et al. 2019) as well as being important for place attachment (douglas et al. 2019). therefore, this result is encouraging. ideally, the location for infrastructure development should be guided by strong strategic planning, promoting the location of development preferably outside of or in buffer areas surrounding parks. furthermore, the research showed that architectural design alternatives were generally well considered for tourist facilities, accommodation and access gate developments, which supports attempts to mitigate aesthetic and sense of place impacts. mitigation was generally dealt with satisfactorily. in the context of national parks, all impacts are potentially significant, making the identification and implementation of effective mitigation measures particularly important. moreover, continual monitoring is critical to be able to gauge the effectiveness of mitigation measures over time. the commitment to the implementation of mitigation measures was rated high, which is to be expected, especially in cases where sanparks was the proponent. public interest also most likely drives mitigation performance in national parks. review area 4: presentation and communication of results the communication of eia information in an accessible format to stakeholders and decision-makers is a critical component of a good quality eia report. all but one report scored satisfactorily in all categories in this review area, which can be explained by the following: rigorous public scrutiny, the oversight role by sanparks and the quality of available baseline data and information (see review area 1). these results suggest that the high quality of the report presentation and communication should place the competent authority in a position to digest, interpret and consider the content towards making an informed and justified decision. temporal trends in environmental impact assessment report quality in terms of eias for national parks, figure 4 shows that four out of six of the reports for the eca 1997 regime were graded as satisfactory (a–c), while all of the eia reports under the other regimes were found to be satisfactory. moreover, four of the six eca 1997 reports obtained a b grading (generally satisfactory) and the remaining two were graded as ‘just unsatisfactory’ (d). in the nema 2006/10 sample, three reports were ‘well performed’ (a), five were ‘generally satisfactory’ and two were graded as ‘just satisfactory’(c). in the nema 2014/17 sample, three reports were graded as ‘well performed’ (a) and the remaining five were graded as ‘generally satisfactory’ (b). in comparison to the eca 1997 regime, the nema regimes (2006/10 and 2014/17) achieved a higher percentage of satisfactory grades, as well as an increase in a and b grades. figure 4: overall quality across environmental impact assessment regimes. therefore, an increase in overall report quality is noticeable from the eca regime to the nema regimes. comparing the nema regimes, an increase in overall report quality is also observed between the nema 2006/10 regime and the nema 2014/17 regime, with the nema 2006/10 regime achieving two c grades, while the nema 2014/17 co-regime only achieved a and b grades. it is encouraging that eia report quality for developments in national parks is increasing over time, in contrast to the negative trend identified by sandham et al. (2013a) in comparing report quality from eca to nema 2006. conclusion and recommendations this research aimed, for the first time, to determine the quality of eia reports for a selected sample of developments in sanparks. an adapted version of the well-known lee and colley eia report quality review package (sandham et al. 2013a) was applied to a sample of 24 eia reports. overall, the eia report quality for developments in national parks was high, which is encouraging. however, certain weaknesses were still evident, especially in relation to the consideration of waste and, to a lesser extent, in dealing with significance and mitigation. the overall good quality results most likely are indicative of certain positive key underlying inputs to the eia process such as: a strong oversight role by the environmental authority and sanparks, quality of data and information, meaningful public interest and participation and eia practitioner competency, but further research is needed to test this hypothesis. nevertheless, despite these positive results, there are still several cautionary aspects to consider given the methodology and the quality review package applied. while the use of the particular review package is fully justified as it allowed for comparative analysis with previous research, it is also limited in terms of its lack of a sector-specific focus. a protected area sector-specific quality review package would potentially include certain important aspects not addressed by this research. for example, consideration of context in the review package applied is limited to one sub-category (1.5.3) specifically focused on alignment with local land-use plans and policies, which does not explicitly reflect the strategic conservation context or other strategic objectives relevant to tourism in protected areas, such as alignment with responsible tourism practices (pope et al. 2019). the strategic context for national parks is critical to set cumulative limits and thresholds for consideration of significance in relation to, for example, expanding the number of beds, day visitors, vehicles and services infrastructure. an emphasis on the strategic context would also allow consideration of cumulative impacts and the extent to which aspects such as climate and land use change have been considered in eia. it is therefore recommended that a protected area-specific review package should include additional sub-categories reflecting other important components of the strategic context for development in protected areas. in view of the weak performance around waste, whilst a revised package might include waste as a separate review area, the results suggest a pressing need for capacity development in terms of waste management in national parks. this assumes that quality grades are poor because of a lack of understanding of how best to manage waste, rather than any other explanation, although this assumption is untested. in south africa, which is a water-stressed country, more emphasis might also be required specifically in relation to water use impacts and mitigation. ideally, the sector-specific quality review package should be designed to still allow for comparison to other report quality research findings. ultimately, good eia report quality could significantly support the potential of eia to contribute to informed decision-making for developments in national parks. with the expected increase in development pressure on national parks, eia is therefore well positioned to continue to inform development decisions in national parks towards more sustainable outcomes. acknowledgements the contributions of anonymous referees are gratefully acknowledged. competing interests the authors have declared that no competing interests exist. authors’ contributions l.a.s. and c.h. conceptualised the project and were responsible for the data collection, analysis and interpretation. l.a.s., f.p.r., a.m.-s., a.j.b., j.p. and r.c.a. drafted and critically revised the manuscript for important intellectual content and approved the final version to be published. funding information this research received no specific grant from any funding agency in the public, commercial or not-for-profit sectors. data availability statement data sharing is not applicable to this article as no new data were created or analysed in this study. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references alberts, r., retief, f., roos, c., cilliers, d. & arakele, m., 2020, ‘re-thinking the fundamentals of eia through the identification of key assumptions for evaluation’, impact assessment and project appraisal 38(3), 205–213. https://doi.org/10.1080/14615517.2019.1676069 arrow, k.a., bolin, b., costanza, r., dasgupta, p., folke, c., holling, c.s. et al., 1995, ‘economic growth, carrying capacity, and the environment’, science 268(1), 520–521. barker, a. & jones, c., 2013, ‘a critique of the performance of eia within the offshore oil and gas sector’, environmental impact assessment review 43, 31–39. bond, a., pope, j., morrison-saunders, a. & retief, f., 2016, ‘a game theory perspective on environmental assessment: what games are played and what does this tell us about decision making rationality and legitimacy?’, environmental impact assessment review 57(1), 187–194. https://doi.org/10.1016/j.eiar.2016.01.002 bond, a., pope, j., retief, f., morrison-saunders, a., fundings land, m. & hauptfleisch, m., 2020, ‘explaining the political nature of environmental impact assessment (eia): a neo-gramscian perspective’, journal of cleaner production 244(1), 118694. https://doi.org/10.1016/j.jclepro.2019.118694 bond, a., retief, f., cave, b., fundings land, m., duinker, p.n., verheem, r. et al., 2018, ‘a contribution to the conceptualisation of quality in impact assessment’, environmental impact assessment review 68(1), 49–58. https://doi.org/10.1016/j.eiar.2017.10.006 boshoff, d.s., 2019, ‘of smoke and mirrors: (mis)communicating eia results of solar energy projects in south africa’, journal of environmental assessment policy and management 21(3), 1950014. https://doi.org/10.1142/s1464333219500145 canelas, l., almansa, p., merchan, m. & cifuentes, p., 2005, ‘quality of environmental impact statements in portugal and spain’, environmental impact assessment review 25(3), 217–225. cashmore, m., christophilopoulos, e. & cobb, d., 2002, ‘an evaluation of the quality of environmental impact statements in thessaloniki, greece’, journal of environmental assessment policy and management 4(4), 371–395. christensen, p., kørnøv, l. & nielsen, e.h., 2005, ‘eia as regulation: does it work?’, journal of environmental planning and management 48(3), 393–412. https://doi.org/10.1080/09640560500067491 cilliers, d., van staden, i., roos, c., alberts, r. & retief, f., 2020, ‘the perceived benefits of eia for government: a regulators perspective’, impact assessment and project appraisal 38(5). https://doi.org/10.1080/14615517.2020.1734403 de witt, l., van der merwe, p. & saayman, m., 2014, ‘critical ecotourism factors applicable to national parks: a visitor perspective’, tourism review international 17(3), 179–194. douglas, a., wessels, j-a., pope, j., morrison-saunders, a. & hughes, m., 2019, ‘measuring kruger visitors’ place attachment to specific camps’, koedoe 61(1), a159, 1–11. https://doi.org/10.4102/koedoe.v61i1.1559 dunjić, j., stojanović, v., solarević, m. & kicošev, v., 2017, ‘sustainable waste management in protected areas of vojvodina’, new spaces in cultural tourism. conference proceedings: ctth 2017, serbia, novi sad, september 1–2, 2017. ehrlich, a. & ross, w., 2015, ‘the significance spectrum and eia significance determinations’, impact assessment and project appraisal 33(2), 87–97. https://doi.org/10.1080/14615517.2014.981023 hallatt, t., retief, f. & sandham, l.a., 2015, ‘a critical evaluation of the quality of biodiversity inputs to eia in areas with high biodiversity value – experience from the cape floristic region, south africa’, journal of environmental assessment policy and management17(3), 1550025. hockings, m., stolton, s., leverington, f., dudley, n. & courrau, j., 2006, evaluating effectiveness: a framework for assessing management effectiveness of protected areas, 2nd edn., iucn, gland. international association for impact assessment and institute of environmental assessment (iaia), 1999, principles of environmental impact assessment best practice, viewed 13 december 2018, from http://iaia.org/publicdocuments/special-publications/principles%20of%20ia_web.pdf jalava, k., pasanen, s., saalasti, m. & kuitunen, m., 2010, ‘quality of environmental impact assessment: finnish eiss and the opinions of eia professionals’, impact assessment project appraisal 28(1), 15–27. https://doi.org/10.3152/146155110x488826 kidd, m., retief, f. & alberts, r., 2018, ‘integrated environmental assessment and management’, in h. strydom, n. king & f. retief (eds.), fuggle and rabie’s environmental management in south africa, pp. 1213–1275, juta, cape town. lee, n., colley, r., bonde, j. & simpson, j., 1999, reviewing the quality of environmental statements and environmental appraisals, occasional paper number 55, eia centre, department of planning and landscape, university of manchester, manchester. mcgrath, c. & bond, a., 1997, ‘the quality of environmental impact statements: a review of those submitted in cork, eire from 1988-1993’, project appraisal 12(1), 43–52. morgan, r.k., 2012, ‘environmental impact assessment: the state of the art’, impact assessment and project appraisal 30(1), 5–14. https://doi.org/10.1080/14615517.2012.661557 morrison-saunders, a., hughes, m., pope, j., douglas, a. & wessels, j-a., 2019, ‘understanding visitor expectations for responsible tourism in an iconic national park: differences between local and international visitors’, journal of ecotourism 18(3), 284–294. https://doi.org/10.1080/14724049.2019.1567740 pope, j., bond, a., morrison-saunders, a., cameron, c. & retief, f., 2018, ‘are current effectiveness criteria fit for purpose? using a controversial strategic assessment as a test case’, environmental impact assessment review 70(1), 34–44. https://doi.org/10.1016/j.eiar.2018.01.004 pope, j., wessels, j.a., douglas, a., hughes, m. & morrison-saunders, a., 2019, ‘the potential contribution of environmental impact assessment (eia) to responsible tourism: the case of the kruger national park’, tourism management perspectives 32(1), 100557. https://doi.org/10.1016/j.tmp.2019.100557 põder, t. & lukki, t., 2011, ‘a critical review of checklist-based evaluation of environmental impact statements’, impact assessment and project appraisal, 29(1), 27–38. retief, f., 2010, ‘the evolution of environmental assessment debates – critical perspectives from south africa’, journal of environmental assessment policy and management 12(4), 1–23. https://doi.org/10.1142/s146433321000370x roos, c., cilliers, d., retief, f., alberts, r. & bond a., 2020, ‘regulators’ perceptions of environmental impact assessment (eia) benefits in a sustainable development context’, environmental impact assessment review 81(1), 106360. https://doi.org/10.1016/j.eiar.2019.106360 sandham, l.a., carrol, t. & retief, f., 2010, ‘the contribution of environmental impact assessment (eia) to decision making for biological pest control in south africa – the case of lantana camara’, biological control 55(2), 141–149. https://doi.org/10.1016/j.biocontrol.2009.12.010 sandham, l.a., hoffmann, a.r. & retief, f.p., 2008a, ‘reflections on the quality of mining eia reports in south africa’, the journal of the southern african institute of mining and metallurgy 108(1), 701–706. sandham, l.a., moloto, m. & retief, f., 2008b, ‘the quality of environmental impact assessment reports for projects with the potential of affecting wetlands’, water sa 34(2),155–162. sandham, l.a. & pretorius, h.m., 2008, ‘a review of eia report quality in the north west province of south africa’, environmental impact assessment review 28(4–5), 229–240. https://doi.org/10.1016/j.eiar.2007.07.002 sandham, l.a., van heerden, a.j., jones, c.e., retief, f.p. & morrison-saunders, a.n., 2013a, ‘does enhanced regulation improve eia report quality? lessons from south africa’, environmental impact assessment review 38(1), 155–162. https://doi.org/10.1016/j.eiar.2012.08.001 sandham, l.a., van der vyver, f. & retief, f.p., 2013b, ‘the performance of environmental impact assessment in the explosives manufacturing industry in south africa’, journal of environmental assessment policy and management 15(3), 1350013. https://doi.org/10.1142/s1464333213500130 steffen, w., richardson, k., rockström., cornell, s.e., fetzer, i., bennett, e.m. et al., 2015, ‘planetary boundaries: guiding human development on a changing planet’, science 347(6223), 1259855. https://doi.org/10.1126/science.1259855 swanepoel, f., retief, f., bond, a., pope, j., morrison-saunders, a., hauptfleisch, m. et al., 2019, ‘evaluating the quality of biodiversity inputs to environmental impact assessments (eia) in areas with high biodiversity value’, journal of environmental assessment, policy and management 21(2), https://doi.org/10.1142/s1464333219500091 swemmer, l.k. & taljaard, s., 2011, ‘sanparks, people and adaptive management: understanding a diverse field of practice during changing times’, koedoe 53(2), 1–7. https://doi.org/10.4102/koedoe.v53i2.1017 wylie, d.k., bhattacharjee, s. & rampedi, i.t., 2018, ‘evaluating the quality of environmental impact reporting for proposed tourism-related infrastructure in the protected areas of south africa: a case study on selected eia reports’, african journal of hospitality, tourism and leisure7(3), 1–14. yang, t., 2019, ‘the emergence of the environmental impact assessment duty as a global legal norm and general principle of law’, hastings law journal 70(2), 525–572. yin, r., 2003, case study research: design and methods, 3rd edn., applied social research methods series, vol. 5, sage, london. footnotes 1. sandham and pretorius (2008) provide an explanation of the origin and nature of ‘beefed-up’ scoping reports. 2. this refers to quality review studies that compare quality over time. filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 filelist convert a pdf file! page 1 page 2 page 3 abstract introduction study area methods results discussion conclusions and management implications acknowledgements references appendix 1 about the author(s) helga van der merwe south african environmental observation network, kimberley, south africa plant conservation unit, university of cape town, cape town, south africa noel van rooyen south african environmental observation network, kimberley, south africa hugo bezuidenhout south african national parks, kimberley, south africa applied behavioural ecology and ecosystem research unit, university of south africa, florida, south africa jacobus du p. bothma centre for wildlife management, university of pretoria, pretoria, south africa margaretha w. van rooyen south african environmental observation network, kimberley, south africa department of plant and soil science, university of pretoria, pretoria, south africa citation van der merwe, h., van rooyen, n., bezuidenhout, h., bothma, j. du p. & van rooyen, m.w., 2019, ‘vachellia erioloba dynamics over 38 years in the kalahari gemsbok national park, south africa’, koedoe 61(1), a1534. https://doi.org/10.4102/koedoe.v61i1.1534 original research vachellia erioloba dynamics over 38 years in the kalahari gemsbok national park, south africa helga van der merwe, noel van rooyen, hugo bezuidenhout, jacobus du p. bothma, margaretha w. van rooyen received: 16 mar. 2018; accepted: 21 nov. 2018; published: 29 apr. 2019 copyright: © 2019. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract vachellia erioloba is a keystone tree species in the southern kalahari. this long-term study over nearly four decades tracks two populations in different landscapes (the interior sandy duneveld versus the clayey nossob riverbed) of a large conservation area and offers valuable data on this species under natural soil moisture conditions and with limited anthropogenic influences. in 1978, 18 trees were permanently marked in a 1 ha plot in the interior duneveld of the kalahari gemsbok national park (dankbaar site). in the nossob riverbed all trees in a 1 ha plot were surveyed in 1979 (grootkolk site). at both sites, tree height and stem circumference were subsequently measured at irregular intervals until 2016 in order to investigate growth rates and population structure. of the 18 marked trees at dankbaar, six died and three showed coppice regrowth following substantial dieback after a fire. a mean height increase of 60 mm/year was recorded and the mean height of the remaining uncoppiced trees was 6.8 m in 2016. stem diameter growth rate per year varied widely between trees and between years with a mean value of 2.5 mm/year over the 38-year period. growth rate calculated for three 10-year intervals varied. using the mean growth rate derived in the current study and stem size of the dead trees, the mean age of the trees when they died was estimated. at the grootkolk site, the position of the centroid in relation to the midpoint of the diameter class range suggests that this population is gradually becoming a mature to old population with limited recruitment. this was supported by the size class distribution curves. however, no differences between slopes or intercepts of the stem diameter size class distributions were found. conservation implications: this study was conducted in a large conservation area, that is, a natural ecosystem excluding most of the anthropogenic threats that are present outside of the park. the study illustrated that in the duneveld the population studied was self-sustaining, with recruitment occurring and large individuals presumably dying of old age. although fire caused a few individuals to coppice, no fire-related deaths were reported. in the nossob riverbed, surveys started in a stand of predominantly young trees and the size class distribution at that stage already showed a lack of recruitment. this stand is ageing and will likely disappear at this site; however, new young stands are appearing at other sites in the nossob riverbed. under the current conditions with negligible anthropogenic influences, it therefore appears that some v. erioloba populations in the park are increasing in size while others are decreasing, but that overall the species will persist. the impact of global climate change on this species is, however, unknown. keywords: conservation area; growth rate; kgalagadi transfrontier park; long-term study; size class distribution; tree age. introduction ecological patterns generated from a single snapshot in time should be used carefully when trying to identify the underlying processes driving the system (jeltsch, moloney & milton 1999), and long-term data can be very useful to assist with the interpretation of ecosystem patterns and processes. however, appropriate long-term data are difficult to find in the case of long-lived organisms such as trees, whose lifetimes are usually considerably longer than those of researchers (menges 2000). in arid systems, this is further exacerbated by low rates of biotic change (cody 2000; lawley et al. 2013; van rooyen et al. 2015). in such systems, to accommodate climatic fluctuations, the monitoring of woody species should take place over a timespan in excess of 20 years (o’connor 1985). in arid savannas, the tree component may seem to be constituted by even-aged cohorts (midgley & bond 2001; wiegand, ward & saltz 2005); however, what appear to be cohorts among adult trees may not be even-aged individuals (seymour 2008). even-aged stands are interpreted as evidence of the importance of abiotic factors, particularly rainfall, to produce pulsed recruitment. apparent even-aged stands signify the release from fire or herbivory whereby different-aged saplings are allowed to recruit simultaneously into larger height classes, thus appearing of equal age (midgley & bond 2001). in the southern kalahari, vachellia erioloba is regarded as a keystone species because it provides food, shelter, perches, nests and roost sites to many types of animals and harbours a distinct assemblage of plant species below its canopy (barnes 2001; leistner 1967; milton & dean 1995; seymour 2008; steenkamp et al. 2008). it is listed as a protected tree species in south africa (government notice no. 1602, 2016). there are, however, growing concerns over increased mortality in mature v. erioloba in the southwestern kalahari (schachtschneider & february 2013). the reasons proposed for the increase in mortality include an increase in fire frequency (seymour & huyser 2008), increasing groundwater abstraction (powell 2005; shadwell & february 2017), harvesting (seymour 2008), diseases (slippers et al. 2014), arboricides (powell 2005) and competition for resources from alien invasive species such as prosopis (robertson & woodborne 2002; schachtschneider & february 2013; zachariades, hoffman & roberts 2011). these threats are especially relevant outside of conservation areas, emphasising the importance of protected environments in which v. erioloba individuals, exposed only to natural threats such as fire, can be studied. vachellia erioloba is a long-lived tree with a lifespan in excess of 240 years (steenkamp et al. 2008). mature individuals of this species are reportedly not easily affected by short-term climatic variations (barnes 2001), possibly because individuals have some of the deepest roots of any known species (the maximum recorded is 68 m), allowing them access to deep groundwater sources (jennings 1974 in moustakas et al. 2006). quantitative information on growth rates and lifespan of the species are, however, still inadequate (steenkamp et al. 2008). from 1978, surveys were initiated in the kalahari gemsbok national park (kgnp) to gather information on the growth rate, lifespan and population dynamics of v. erioloba in the interior duneveld (dankbaar) and in the nossob riverbed habitats (grootkolk). the objectives of this paper were (1) to report on the mean annual increase in stem diameter and tree height of the 18 marked v. erioloba trees at the dankbaar site; and (2) to document the mean age of those trees that died in the interior duneveld of the park; and (3) at the grootkolk site to describe changes in the size class distribution of a v. erioloba population in the nossob riverbed. the publication of these findings contributes to our understanding of v. erioloba population dynamics within large conservation areas without many of the anthropogenic factors proposed to increase mortality in this species. study area the kgnp is situated in the southern kalahari (24°15’s–26°30’s and 20°00’e–20°45’e) and covers approximately 9600 km² (figure 1). it forms part of the kgalagadi transfrontier park, together with the gemsbok national park in botswana. figure 1: location of the two vachellia erioloba study sites, dankbaar situated in the interior duneveld and grootkolk located in the nossob riverbed, within the kalahari gemsbok national park. the mean annual rainfall recorded from 1976 to 2015 at nossob rest camp in the park is 194 mm (figure 2), with most rain falling between january and april. the annual rainfall has a high coefficient of variation and the rain often falls as short-duration, high-intensity thunderstorms. temperatures show a large range, with winter lows reaching -10.3 °c and summer highs reaching 45.4 °c (van rooyen & van rooyen 1998). figure 2: total annual rainfall (january to december) at nossob rest camp from 1976 until 2016. the orange line represents the mean annual rainfall of 194 mm. sampling was conducted at two sites (figure 1). the first site is situated 2.7 km east of dankbaar (1040 m above sea level) in the interior duneveld in the acacia (vachellia) erioloba – schmidtia kalahariensis low duneveld landscape (van rooyen et al. 2008). this open tree savanna, dominated by v. erioloba (tree) and s. kalahariensis (grass), occurs in the interior duneveld in the northern part of the park. the aeolian sands of the low duneveld landscape are piled into a gently undulating landscape. beneath the sand lies a vast sheet of calcareous or silicified sand or sandstone that contains grits and minor conglomerates (malherbe 1984). van rooyen (1984) describes the duneveld soils as deep, red, eutrophic, fine and medium sandy soils of the hutton soil form, or yellow-brown, eutrophic and calcareous, fine to medium sandy soils of the clovelly soil form (soil classification working group 1991). the second site, at grootkolk (1041 m above sea level), is situated in the nossob riverbed in the acacia (vachellia) erioloba – acacia (vachellia) haematoxylon riverbeds landscape (van rooyen et al. 2008). this landscape is characterised by large scattered v. erioloba trees with a dense grass stratum dominated by panicum coloratum. van rooyen (1984) described the nossob riverbed’s soil as deep, calcareous, reddish-brown and grey-brown loam with clay soils of the oakleaf, dundee and valsrivier soil forms. upstream from the grootkolk site, shallow, brown and yellow-brown calcareous sands and loams of the mispah soil form occur (soil classification working group 1991). fine-grained soils in arid areas show the so-called inverse texture effect (noy-meir 1973), with the result that there is less available soil water and consequently lower production on fine-textured soils than on coarse-textured soils (van rooyen et al. 1984). methods field surveys two study sites (dankbaar site and grootkolk site) were surveyed on numerous occasions; however, survey methods used at these two sites differed. in 1978, all v. erioloba trees were marked in a 1 ha plot at the dankbaar site. tree height and stem circumference were recorded for each of the 18 permanently marked trees at irregular intervals from 1978 to 2016 (survey years 1978, 1980, 1983, 1985, 1988, 1989, 1991, 1994, 1998, 1999, 2008 and 2016). in 2016, for the first time since the original survey in 1978, all additional trees within the plot were measured and permanently marked. at the 1 hectare (ha) grootkolk site, height and stem circumference of all v. erioloba individuals were measured at irregular intervals from 1979 to 2016 (survey years 1979, 1984, 1986, 1989, 1994, 1996, 1999, 2008 and 2016). the stem circumferences of all dead trees were also recorded at grootkolk in the year of the survey when the tree was found to have died. analysis because of the different survey methods used at the two sites; the data collected could not be analysed in the same manner for both the sites. for example, only the 18 marked trees at dankbaar were surveyed throughout the 38-year period; thus an analysis of the population structure is inappropriate as the entire population in the 1 ha plot was not surveyed. an estimate of the growth rate per year over the entire study period (38 years) was calculated for each marked tree at the dankbaar site by subtracting the first measured stem diameter (1978 survey) from the last live measured stem diameter and dividing by the number of years between these two measurements. similarly, this method was used to calculate changes in height of a tree. a linear regression using graphpad 7.02 (graphpad software, www.graphpad.com) of stem diameter versus annual growth rate was used to investigate the relationship between these two variables. additionally, stem diameter growth rates over the first 10-year period (1978–1988), second 10-year period (1988–1998) and third 10-year period (1988–2008) were calculated for each marked individual. this was calculated in the same way as for the entire period, but using the first and last measured diameters for the appropriate periods and calculating a mean across all the individuals. dead and coppiced individuals were excluded from the calculations from the survey year in which the death or coppicing was recorded for the first time. the mean stem diameter in 1978 and 2016 of all marked trees was used to estimate the average age using the growth rate determined in the current study. the average age of all the marked trees in 1978 and in 2016 were calculated by dividing the mean stem diameter for the particular year by the growth rate determined in the current study. for comparative purposes the mean age of the trees was also calculated by the same method using the mean diameter growth rate for only the interior duneveld (3.2 mm/year) and for the interior duneveld and riverbed combined (3.6 mm/year) reported by steenkamp et al. (2008). age was also determined using the growth rate for the interior duneveld and riverbed combined (5.1 mm/year) based on the equation derived for the relationship between radiocarbon age and stem circumference (steenkamp et al. 2008). these calculations were made using the mean stem diameter of the live trees in 2016 as well as the last ‘alive’ measured stem diameter for the dead trees. these calculations were also performed for the grootkolk population using the mean stem diameter of the stem circumference classes. the stem circumferences for trees at the grootkolk 1 ha plot were classified into 12 equal stem circumference classes (0–20, > 20–40, > 40–60, > 60–80, > 80–100, > 100–120, > 120–140, > 140–160, > 160–180, > 180–200, > 200–220, > 220–240 starting with class 1 and ending with class 12) for each of the survey years. the size class distribution was examined visually to classify the population into one of four population structure types (gaugris & van rooyen 2007; figure 3). in this model, type i size class distribution represents a growing population and follows an inverse j-shaped curve, which represents the ideal curve shape for a growing population in a natural environment. there is a constant decrease in the number of individuals as the size classes increase, and the curve spans the whole range of size classes described for the species. populations with a type ii curve have a near ideal population structure that differs from a type i only by having a lower abundance of smaller size classes. populations with type iii size class distributions are typically associated with disturbance. the type iiia curve broadly resembles a type i size class distribution but applies to populations where a section of the curve spanning one or several size classes is missing or where abundance is lower than expected. the typical large pool of small individuals is generally missing in type iiib and the size class distribution shows a succession of minor peaks, even with some gaps where one or several size classes may be missing altogether. in the type iv curve, one or two peaks are observed over a restricted range of size classes, and this curve thus represents an abnormal size class distribution. the curve is either bell-shaped with a peak more or less in the middle of the size class range, or it shows a high frequency of large size classes, with little or no representation of smaller classes. figure 3: idealised shapes of the four different population curve types. for the 1 ha grootkolk site the number of individuals recorded per stem circumference size class was calculated in order to obtain the density of trees per size class. regressions were performed on the density (di) of individuals against size class (mi) (gaugris & van rooyen 2007). the diameter class midpoint (mi) was calculated as the mean of the upper and lower limit of each size class (condit et al. 1998). logarithmic transformations, ln (di + 1) and ln (mi), were used to standardise the data (lykke 1998; niklas, midgley & rand 2003) before performing least square linear regressions, that is, ln (di + 1) for the y-axis and ln (mi) for the x-axis. the value of +1 was added because some size class bins were empty (lykke 1998). the linear regressions were evaluated in terms of their slopes and y-axis intercepts (gaugris, vasicek & van rooyen 2012; van der merwe & geldenhuys 2017). strongly negative slopes typically represent regenerating populations (condit et al. 1998), and a high y-axis intercept indicates many small individuals in the population. a y-axis intercept near to zero indicates that few small individuals are present (everard, midgley & van wyk 1994). both slopes and y-axis intercepts of regressions were compared statistically across years by means of an analysis of covariance using graphpad 7.02 (graphpad software, www.graphpad.com. two additional mean values were calculated for the stem diameter distribution for each survey year: (1) the ‘centroid’ was calculated as the arithmetic mean of all stem diameters recorded; and (2) the ‘midpoint of stem diameter range’ was calculated as the mean of the upper and lower limits of the diameter range included in the size class distribution. niklas et al. (2003) found a relationship between the position of the centroid and the diameter class range midpoint for a species. if a centroid value is smaller than the midpoint value, the population is young and growing. when the centroid value is larger than the midpoint value, the population is mature to old. in general, as the mean stem diameter increases in a population, self-thinning occurs and the density decreases, in such a way that an increasing part of the biomass is found in fewer individuals (niklas et al. 2003; silvertown & charlesworth 2001). as self-thinning occurs the centroid of a population shifts towards the right within the size class distribution. results growth rate over the period of 38 years, 6 of the 18 originally marked trees at the dankbaar site died (33%) and three showed coppice regrowth following substantial dieback because of a fire (table 1-a1). in 1978 the mean height of the 18 trees was 5.1 m and mean stem diameter 235.85 mm. in 2016 the mean height of the remaining nine originally marked trees that had not experienced dieback was 6.8 m and mean stem diameter 282.7 mm. the mean height for the six dead individuals at the onset of the study was 6.1 m and at the time of last measurement before death 6.8 m. a mean height increase of 60 mm/year was calculated. the 2016 survey measured the remaining 12 originally marked trees as well as 12 additional trees now present; thus, after 38 years, the population increased by six trees in the 1 ha plot. table 1: age calculations for vachellia erioloba trees in the interior duneveld at the dankbaar site using mean stem diameter of the 18 originally marked trees in 1978, the nine remaining live trees recorded in 2016 and the six dead trees last measured stem diameter when last surveyed as alive, derived by different methods. a linear regression of growth rate against stem diameter (figure 4) indicated that growth rate decreases significantly as stem diameter increases, that is, as the individuals age (p = 0.022). the mean growth rate for the first 10-year period (1978–1988) was 2.99 mm/year, the second 10-year period (1988–1998) 1.01 mm/year and the third 10-year period (1998–2008) 2.84 mm/year, indicating a variable growth rate over time. stem diameter growth rate per year varied widely between the nine remaining trees (0.91 mm – 4.86 mm/year), with a mean value of 2.5 mm/year. depending on the choice of mean annual growth rate or application of the regression between stem circumference and age, the mean age of the six dead trees could be estimated at between 71 and 142 years (table 1). figure 4: linear regression of stem diameter versus annual stem diameter growth rate for the 18 marked vachellia erioloba trees at the dankbaar site in the kalahari gemsbok national park. population structure at the grootkolk site, 61 live and 14 dead individuals were recorded in the initial survey in 1979. the number of live individuals at grootkolk in the nossob riverbed gradually decreased over the survey period (figure 5). initially the size class distribution of the trees from 1979 to 1999 could visually be classified as type iiib curves; however, there was a gradual transition to a type iv curve from 2008 to 2016 (figures 3 and 5). in spite of the decreasing tree numbers and the change of curve shape, linear regressions on the density of individuals against the diameter class midpoint (figure 1-a1) revealed no significant differences between the slopes (table 2-a1) or intercepts (table 3-a1) of the fitted lines. figure 5: size class distribution curves of vachellia erioloba individuals at the grootkolk site in the various survey years. (a) 1986; (b) 1984; (c) 1986; (d) 1989; (e) 1994; (f) 1996; (g) 1999; (g) 2008; (i) 2016. using calculations from steenkamp et al. (2008) for the interior duneveld and riverbed, the largest live tree at grootkolk was aged between 195 and 212 years, while the relationship between stem circumference and radiocarbon age for the interior duneveld and riverbed combined aged the tree between 136 and 148 years (table 2). when using the same two methods of calculation, most of the dead trees fell in the age categories 25–53, 37–71 and 49–89 years. table 2: age calculations, derived by different methods, for vachellia erioloba trees at the grootkolk site in the nossob riverbed using mean stem diameter of the circumference class for the largest live tree; and for circumference size classes that recorded the majority of the dead trees. in all survey years the position of the centroid (arithmetic mean of all stem diameters recorded per site) was to the left of the midpoint of the size class distribution. however, the difference between these two values decreased over time and in 2016 the two values were nearly the same (table 4-a1), indicating that the population was changing from a young and growing population to a mature to old population. discussion growth rate marked v. erioloba tree stem diameters at dankbaar in the interior duneveld increased at a mean rate of 2.50 mm/year (range 0.91 mm – 4.86 mm/year) over the monitored period. this mean annual diameter increase is less than the 3.6 mm/year established by steenkamp et al. (2008) and substantially lower than the 12 mm – 14 mm/year previously reported for acacia species growing in arid and semi-arid environments (gourlay 1992). the steenkamp et al. (2008) study used annual growth ring and radiocarbon ageing to determine growth rates. a mean growth rate was derived from individuals in the interior duneveld as well as in the nossob riverbed. if only the individuals in the interior duneveld in the steenkamp et al. (2008) study are considered, then the mean annual diameter increment is 3.2 mm/year. although steenkamp et al. (2008) concluded that the growth rate did not differ significantly between trees growing in the dry nossob riverbed and those growing in the interior duneveld, growth rate varied substantially between individual trees. variability in the growth rates of other tree species has been reported and the suggested underlying causes for this phenomenon include microsite conditions and/or genetic potential of individual trees (stahle et al. 1996). depending on the equation used to determine the age of the dead trees there could be a twofold or greater difference in calculated age at both the dankbaar (table 1) (71 years as opposed to 142 years) and grootkolk (table 2) (25 years as opposed to 127 years) sites. the high variability in estimated age at which 6 of the 18 marked trees died (48, 103, 163, 178, 181 and 191 years, using the mean annual diameter increase of 2.5 mm/year calculated for this current study) was unexpected. moustakas et al. (2008), using a combination of aerial photographs and satellite images covering a period of 61 years to provide spatial data on mortality, found that v. erioloba mortality risk was greatest for trees of intermediate size. this does not seem to be the case for the dankbaar population, where mortality was found to be predominantly among the large trees. at grootkolk, the data seem to indicate that mortality was highest in smaller size classes and gradually declined for larger size classes. the density of trees at grootkolk was low, and consequently self-thinning (silvertown & charlesworth 2001) is unlikely to have occurred. population structure on the basis of the position of the centroid in relation to the midpoint of the stem diameter class range, the grootkolk population could be considered as a population gradually changing from young and growing to mature to old. this is supported by a visual assessment of the size class distribution, which showed a gradual transition from a type iiib to a type iv curve shape over the 38 years. the large peak of young individuals in size class 4 that was evident at the first survey gradually became smaller and shifted to size class 5, and a prominent second peak emerged at size class 8 at the last survey. at grootkolk, few seedlings were recorded and few individuals appeared to be recruiting into the population. the low survival of size class 2, 3 and 4 individuals over the monitored period could possibly be ascribed to individuals progressing to larger size classes over the monitored period without being replenished. this is supported by the fact that size classes 5 and 8 had more individuals at the end of the monitoring period than when monitoring commenced. overall, the survival of individuals in the larger size classes (e.g. size classes 6, 7 and 9) exceeded that in the smaller size classes (e.g. size classes 1, 2 and 3). in spite of the fact that the grootkolk population was almost halved over the 38-year period, the slopes and intercepts of the size class distribution regressions showed no differences between years (figure 1-a1). in their study, steenkamp et al. (2008) reported an ageing size class distribution with poor recruitment at some sites in the nossob riverbed. at other sites, many young individuals were found with only a few mature or dying trees in these populations. the grootkolk site situated in the same riverbed illustrated an ageing population with few young individuals. it has been suggested that flooding of the nossob river and/or consecutive years of above-average rainfall were necessary for regeneration of v. erioloba. steenkamp et al. (2008) were, however, unable to find support to substantiate the need for flooding. seed germination of v. erioloba is best after good rain events, and seedling survival depends on the availability of moisture during the next few seasons. mortality rates during seedling establishment are generally higher than during the adult stages (silvertown & charlesworth 2001). seedlings and saplings suffer high mortality in dry years (barnes 2001; theron, van rooyen & van rooyen 1985; van rooyen & van rooyen 1998; van rooyen et al. 1984). the high density of wildlife in the nossob riverbed could also have contributed to the ageing of the grootkolk population, with losses in young individuals because of trampling and browsing by herbivores and occasional toppling and breaking of larger trees by wind. likewise, insects and rodents that impact on the seed pool may be more numerous in the riverbed habitat (steenkamp et al. 2008). furthermore, competition from a well-established grass layer may inhibit tree seedling establishment and growth in the riverbed (steenkamp et al. 2008); however, this was not evident at the grootkolk site. fire also significantly impacts v. erioloba populations. after good rainfall, sufficient fuel accumulates and enables fire to spread through the landscape. in general, up to one-third of individuals in an area that experiences a fire are killed by the fire, with most of the individuals subsequently resprouting from the base (van der walt & le riche 1984). large old trees, especially those with hollow stems, are most sensitive to fire (milton & dean 1995). during the study period, fire only occurred at the dankbaar study site (large fires in 1995 and 2012). no tree mortality resulting from the fire was evident at the site; however, three of the individuals displayed coppiced growth subsequent to fire. vachellia erioloba population dynamics differed at the two study sites over the last 38 years. considering all tree mortalities and recruitments, the dankbaar population can be considered a self-sustaining population with an increase of 12 new individuals (net increase of six trees) at the site over the 38-year period. the density (number of individuals per the 1 ha plot) of the grootkolk population is declining and the size classes of the remaining individuals increasing as the population ages. conclusions and management implications there is growing concern that mortality rates in v. erioloba, a keystone species of the kalahari, are increasing as a result of threats such as increasing fire frequency, groundwater abstraction, competition from alien invasive species, diseases, harvesting and the effect of arboricides. this study in the kgnp provided an ideal opportunity to investigate v. erioloba dynamics in a natural environment, under different soil types and consequently water availability, where threats were negligible or absent. the different survey methods used at the two sites and alternative data analysis techniques have provided findings on the growth of individual trees as well as on population dynamics. additionally, the study provided evidence that the population studied in the interior duneveld was self-sustaining and although the population in the nossob riverbed was declining, new young populations were being established elsewhere in the riverbed (steenkamp et al. 2008). the continued monitoring of the population at the two sites, where baseline data are already available, is highly recommended. however, monitoring sites should also be established at additional sites outside of park borders to incorporate sites in which various threats to this species can be investigated. it would also be imperative to determine growth rates in the riverbeds. rainfall stations should be erected at all sites to enable the investigation of the influence of significant rainfall events and drought on seed germination, seedling survival and adult mortality. acknowledgements the authors gratefully acknowledge the financial assistance of the university of pretoria, centre for wildlife management and the national research foundation (gun 2053522, grant numbers 61277, 81089 and 91434). the numerous honours students in wildlife management and botany are thanked for their assistance in the field. south african national parks (sanparks) are thanked for the opportunity to conduct the research in the kalahari gemsbok national park. the field surveys of 2016 were successfully conducted with the assistance of sanparks and south african environmental observation network employees. the anonymous reviewers and editor are also thanked for their valuable contributions to this manuscript. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions n.v.r. donated the data set to south african environmental observation network. the authors provided financial and/or institutional support and conducted numerous field surveys over the years. h.v.d.m. and m.w.v.r. compiled the manuscript with valuable input from the remaining co-authors. funding information funding for this study was provided by the university of pretoria, centre for wildlife management and the national research foundation. references barnes, m.e., 2001, ‘effects of large herbivores and fire on the regeneration of acacia erioloba in chobe national park, botswana’, african journal of ecology 39, 340–350. https://doi.org/10.1046/j.1365-2028.2001.00325.x cody, m.l., 2000, ‘slow-motion population dynamics in mojave desert perennial plants’, journal of vegetation science 11, 351–358. https://doi.org/10.2307/3236627 condit, r., sukumar, r., hubbel, s. & foster, r., 1998, ‘predicting population trends from size distributions: a direct test in a tropical tree community’, american naturalist 152, 495–509. https://doi.org/10.1086/286186 everard, d.a., midgley, j.j. & van wyk, g.f., 1994, ‘disturbance and the diversity of forests in natal, south africa: lessons for their utilization’, strelitzia 1, 275–285. gaugris, j.y. & van rooyen, m.w., 2007, ‘the structure and harvesting potential of the sand forest in tshanini game reserve, south africa’, south african journal of botany 73, 611–622. https://doi.org/10.1016/j.sajb.2007.06.004 gaugris, j.y., vasicek, c.a. & van rooyen, m.w., 2012, ‘herbivore and humans impact on woody species dynamics in maputaland, south africa’, forestry 4, 497–512. https://doi.org/10.1093/forestry/cps046 gourlay, i.d., 1992, ‘age and growth rate determination of some african acacia species’, msc dissertation, university of oxford, london. government notice no. 1602, 2016, ‘notice of the list of protected tree species under the national forests act (1998) (act no. 84 of 1998)’, government printer, republic of south africa. jeltsch, f., moloney, k.a. & milton, s.j., 1999, ‘detecting processes from snap-shot pattern – lessons from tree spacing in the southern kalahari’, oikos 85, 451–466. https://doi.org/10.2307/3546695 lawley, v., parrot, l., lewis, m., sinclair, r. & ostendorf, b., 2013, ‘self-organization and complex dynamics of regenerating vegetation in an arid ecosystem: 82 years of recovery after grazing’, journal of arid environments 88, 156–164. https://doi.org/10.1016/j.jaridenv.2012.08.014 leistner, o.a., 1967, ‘the plant ecology of the southern kalahari’, memoirs of the botanical survey of south africa 38, 1–172. lykke, a.m., 1998, ‘assessment of species composition change in savanna vegetation by means of woody plants’ size class distributions and local information’, biodiversity and conservation 7, 1261–1275. https://doi.org/10.1023/a:1008877819286 malherbe, s.j., 1984, ‘the geology of the kalahari gemsbok national park’, koedoe 33–44. menges, e.s., 2000, ‘population viability analyses in plants: challenges and opportunities’, trends in ecology & evolution 15, 51–56. https://doi.org/10.1016/s0169-5347(99)01763-2 midgley, j.j. & bond, w.j., 2001, ‘a synthesis of the demography of african acacias’, journal of tropical ecology 17, 871–886. https://doi.org/10.1017/s026646740100164x milton, s.j. & dean, w.r.j., 1995, ‘how useful is the keystone species concept, and can it be applied to acacia erioloba in the kalahari desert?’, zeitschrift für ökologie und naturschutz 4, 147–156. moustakas, a., guenther, m., wiegand, k., mueller, k-h., ward, d., meyer, k.m. et al., 2006, ‘long-term mortality patterns of the deep-rooted acacia erioloba: the middle class shall die!’, journal of vegetation science 17, 473–480. https://doi.org/10.1658/1100-9233(2006)17[473:lmpotd]2.0.co;2 moustakas, a., wiegand, k., getzina, s., ward, d., meyer, k.m., guenther, m. et al., 2008, ‘spacing patterns of an acacia tree in the kalahari over a 61-year period: how clumped becomes regular and vice versa’, acta oecologica 33, 355–364. https://doi.org/10.1016/j.actao.2008.01.008 niklas, k.j., midgley, j.j. & rand, r.h., 2003, ‘tree size frequency distributions, plant diversity, age and community disturbance’, ecological letters 6, 405–411. https://doi.org/10.1046/j.1461-0248.2003.00440.x noy-meir, i., 1973, ‘desert ecosystems: environment and producers’, annual review of ecology and systematics 4, 25–51. https://doi.org/10.1146/annurev.es.04.110173.000325 o’connor, t.g., 1985, a synthesis of field experiments concerning the grass layer in the savanna regions of southern africa, south african national programmes report no. 114, f.r.d., csir, pretoria. powell, e., 2005, what is happening to the camel thorns of kathu?, veld and flora, cape town, pp. 34–35. robertson, i. & woodborne, s., 2002, carbon isotopes confirm the competitive advantages of prosopis over acacia erioloba. study of environmental change using isotope techniques, international atomic energy agency, vienna. iaea-csp-13/p. schachtschneider, k. & february, e.c., 2013, ‘impact of prosopis invasion on a keystone tree species in the kalahari desert’, plant ecology 4, 597–605. https://doi.org/10.1007/s11258-013-0192-z seymour, c., 2008, ‘grass, rainfall and herbivores as determinants of acacia erioloba (meyer) recruitment in an african savanna’, plant ecology 197, 131–138. https://doi.org/10.1007/s11258-007-9366-x seymour, c.l. & huyser, o., 2008, ‘fire and the demography of camelthorn (acacia erioloba meyer) in the southern kalahari—evidence for a bonfire effect?’, african journal ecology 46, 594–601. https://doi.org/10.1111/j.1365-2028.2007.00909.x shadwell, e. & february, e., 2017, ‘effects of groundwater abstraction on two keystone tree species in an arid savanna national park’, peer j 5, e2923. https://doi.org/10.7717/peerj.2923 silvertown, j. & charlesworth, d., 2001, introduction to plant population biology, 4th edn., blackwell science, oxford, london. slippers, b., roux, j., wingfield, m.j., van der walt, f.j.j., jami, f., mehl, j.w.m. et al., 2014, ‘confronting the constraints of morphological taxonomy in the botryosphaeriales’, persoonia 33, 155–168. https://doi.org/10.3767/003158514x684780 soil classification working group, 1991, ‘soil classification: a taxonomic system for south africa’, memoirs on the agricultural natural resources of south africa 15, 1–262. stahle, d.w., cleaveland, m.k., haynes, g.a., klimowicz, j., ngwenya, p. & mushove, p., 1996, ‘preliminary tree-ring investigations of indigenous trees in zimbabwe’, in p.t. mushove, e.m. shumba & f. matose (eds.), sustainable management of indigenous forests in the dry tropics. proceedings of an international conference in kadoma, zimbabwe, may 28–june 01, 1996, zimbabwe forestry commission, harare, pp. 57–61. steenkamp, c.j., vogel, j.c., fuls, a., van rooyen, n. & van rooyen, m.w., 2008, ‘age determination of acacia erioloba trees in the kalahari’, journal of arid environments 72, 302–313. https://doi.org/10.1016/j.jaridenv.2007.07.015 theron, g.k., van rooyen, n. & van rooyen, m.w., 1985, ‘vegetation structure and vitality in the lower kuiseb’, in b.j. huntley (ed.), the kuiseb environment: the development of a monitoring baseline, south african scientific programmes report 106, pp. 81–91, council for scientific and industrial research, pretoria. van der merwe, h. & geldenhuys, c., 2017, ‘proposed long-term monitoring protocol and applications for aloidendron dichotomum populations’, south african journal of botany 109, 253–262. https://doi.org/10.1016/j.sajb.2017.01.008 van der walt, p.t &, le riche, e.a.n., 1984, ‘the influence of veld fire on an acacia erioloba community in the kalahari gemsbok national park’, koedoe 103–106. van rooyen, t.h., 1984, ‘the soils of the kalahari gemsbok national park’, koedoe 27(suppl.) 45–61. https://doi.org/10.4102/koedoe.v27i2.568 van rooyen, m.w., le roux, a., geldenhuys, c., van rooyen, n., broodryk, n.l. & van der merwe, h., 2015, ‘long-term vegetation dynamics (40yr) in the succulent karoo, south africa: effects of rainfall and grazing’, applied vegetation science 18, 311–322. https://doi.org/10.1111/avsc.12150 van rooyen, n., van rensburg, d.j., theron, g.k. & bothma, j.d.p., 1984, ‘a preliminary report on the dynamics of the vegetation of the kalahari gemsbok national park’, koedoe 27(suppl.) 83–102. https://doi.org/10.4102/koedoe.v27i2.570 van rooyen, n. & van rooyen, m.w., 1998, ‘vegetation of the south-western arid kalahari: an overview’, transactions of the royal society of south africa 53, 113–140. https://doi.org/10.1080/00359199809520381 van rooyen, m.w., van rooyen, n., bothma, j.d.p. & van den berg, h., 2008, ‘landscapes in the kalahari gemsbok national park, south africa’, koedoe 50, 99–112. https://doi.org/10.4102/koedoe.v50i1.154 wiegand, k., ward, d. & saltz, d., 2005, ‘multi-scale patterns and bush encroachment in an arid savanna with a shallow soil layer’, journal of vegetation science 16, 311–320. https://doi.org/10.1111/j.1654-1103.2005.tb02369.x zachariades, c., hoffman, j.h. & roberts, a.p., 2011, ‘biological control of mesquite (prosopis species) (fabaceae) in south africa’, african entomology 19, 402–415. https://doi.org/10.4001/003.019.0230 appendix 1 table 1-a1: changes in stem diameter (mm) from 1978 to 2016 of 18 marked vachellia erioloba trees at the dankbaar site in the interior duneveld of the kalahari gemsbok national park. table 2-a1: statistical comparison of slope values for linear regressions fitted on the density of individuals against the diameter class midpoint of live trees at grootkolk in the kalahari gemsbok national park using an analysis of covariance. table 3-a1: statistical comparison of intercept values for linear regressions fitted on the density of individuals against the diameter class midpoint of live trees at grootkolk in the kalahari gemsbok national park using an analysis of covariance. table 4-a1: calculated centroid (arithmetic mean of all stem diameters recorded per site in mm) and the midpoint (mm) of the size class distribution for each year surveyed at the grootkolk site in the nossob riverbed. figure 1-a1: linear regressions fitted on the ln (mi) against ln (di + 1) of live trees at grootkolk in the kalahari gemsbok national park in (a) 1979, (b) 1984, (c) 1986, (d) 1989, (e) 1994, (f) 1996, (g) 1999, (h) 2008 and (i) 2016, where mi, midpoint of the size class and di, density of individuals. filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 filelist convert a pdf file! page 1 page 2 page 3 sponnheimer.qxd diets of impala from kruger national park: evidence from stable carbon isotopes m. sponheimer, c.c. grant, d.j. de ruiter, j.a lee-thorp, d.m. codron and j. codron sponheimer, m., c.c. grant, d.j. de ruiter, j.a. lee-thorp, d.m. codron and j. codron. 2003. diets of impala from kruger national park: evidence from stable carbon isotopes. koedoe 46(1): 101–106. pretoria. issn 0075-6458. impala are known to exhibit dietary flexibility, relying primarily on browse in some areas and graze in others. in this study we use stable isotope analysis of faeces and hair to examine the diets of impala in kruger national park. as expected, the data show that impala are mixed-feeders and highly distinct from grazing buffalo and browsing kudu. moreover, impala, buffalo, and kudu faeces contain 2.1 %, 1.4 %, and 2.9 % nitrogen respectively, suggesting that impala diets are of intermediate quality. there are also marked differences between impala populations in the northern and southern regions of the park. the northern impala graze less than their southern counterparts. this difference probably reflects decreased availability of herbaceous forage in the mopane-dominated north. males and females also have different diets, with males grazing more than females. key words: impala, carbon isotopes, faeces, hair, diet. sponheimer, m., department of archaeology, university of cape town, private bag, rondebosch, 7700, rsa ( current address: department of biology, university of utah, salt lake city, utah, 84112, usa ); c.c. grant, school of animal plant and environmental sciences, university of the witwatersrand, private bag 3, johannesburg, 2050, rsa; d.j. de ruiter, palaeoanthropology unit for research and exploration, university of the witwatersrand, private bag 3, johannesburg, 2050, rsa; j.a. leethorp, d.m. codron and j. codron, department of archaeology, university of cape town, private bag, rondebosch, 7700, republic of south africa. issn 0075-6458 101 koedoe 46/1 (2003) introduction impala (aepyceros melampus) are known to exhibit great dietary flexibility. they can make large changes in the percentages of browse and graze consumed seasonally (monro 1980; dunham 1982; smithers 1983; meissner et al. 1996; wronski 2002). one study suggested that grasses make up 90 % of impala diets during the rainy season, but only 33 % during the dry season (meissner et al. 1996). impala diets are also believed to differ markedly between localities reflecting changes in local vegetation. for instance, it has been reported that some tanzanian impala populations eat over 90 % grass (lamprey 1963), whereas some zimbabwean populations have diets dominated by browse (smithers 1983). here, we investigate the diets of impala from kruger national park, south africa, using stable carbon isotope analysis of faeces and hair. stable carbon isotopes of hair or faeces have been used to investigate the diets of wildlife (tieszen & imbaba 1980; van der merwe et al. 1988; witt & beeton 1997; wit et al. 1998; schoeninger et al. 1999), but such studies are few in number. furthermore, these materials have never been analysed in tandem. stable carbon isotope analysis is particularly useful for determining the percentages of browse and graze in the diets of african herbivores (vogel 1978; ambrose & deniro 1986). grasses (graze) in tropical environments use the c4 photosynthetic pathway, while trees, shrubs, and forbs (browse) use the c3 pathway. this results in these food sponnheimer.qxd 2005/12/09 10:30 page 101 types having highly divergent, non-overlapping ratios of 13c/12c (smith & epstein 1971). because the carbon from these food sources is incorporated into faeces and hair in well-understood ways (deniro & epstein 1978; jones et al. 1981; cerling & harris 1999; o’connell & hedges 1999), we can then use the 13c/12c ratios of these materials to determine the percentages of graze and browse in an animal’s diet. methods the kruger national park is broadly divided into granite-derived soils in the west and basalt-derived soils in the east. the rainfall decreases from south to north and to a lesser extent from west to east (joubert 1986). tail hair samples were obtained from darted animals in december, 2001 and faeces were collected in june, 2002. our collections included animals from the northern and southern regions of the park, to see if the different rainfall regimes led to divergent trophic behavior. in order to provide a comparative baseline, we also gathered tail hair and faeces samples from the grazing cape buffalo (syncerus caffer) and browsing kudu (tragelaphus strepsiceros). the darted animals were from healthy populations that were undergoing routine veterinary study by kruger national park staff. thus, our sampling strategy was perforce opportunistic. about 6 cm of tail hair was homogenised so that analysis would provide a long-term dietary average. hair grows at the rate of about 1 cm per month (saitoh et al. 1969), so the samples we analysed represented about 6 months’ growth. faecal samples were dried, homogenised and ground. hair and faecal samples were combusted in an automated carloerba device (carlo-erba, milan) and stable carbon isotopes were analysed using a flow-through inlet system on a continuous flow isotope ratio mass spectrometer (finnigan, bremen). this provided us with the 13c/12c ratios and percentage nitrogen of each sample. the 13c/12c ratios are expressed here as δ values in parts per thousand (‰) relative to the pdb standard. the standard deviation for replicate measurements of a yeast standard was < 0.1 ‰. data from grazing buffalo and browsing kudu are included to provide a comparative baseline. we used oneway analysis of variance to look for species, region, and sex differences. the δ13c values were converted to % c4 consumed using –12.5 ‰ and –27.0 ‰ as pure grazing and browsing dietary endpoints (vogel et al. 1978), and assuming isotopic fractionations of –0.9 ‰ and +3.0 for faeces and hair respectively (jones et al. 1981; cerling & harris 1999; sponheimer pers. obs.). for example, an animal with hair δ13c of –24.0 ‰ would have a dietary δ13c of –27.0 ‰, and thus be koedoe 46/1 (2003) 102 issn 0075-6458 table 1 sample size, δ13c and % c4 grass consumed for all groups in this study. percent c4 grass consumption was determined assuming fractionations of –0.9 ‰ and +3.0 ‰ for faeces and hair respectively (e.g., jones et al., 1981; cerling & harris 1999). percent nitrogen is also included for faeces species n δ13c % n % c4 faeces impala combined 39 -20.5±2.8 2.1±0.5 53±20 impala north 18 -22.2±2.6 2.1±0.7 41±19 impala south 21 -19.2±2.2 2.1±0.4 63±16 buffalo 19 -14.7±0.5 1.4±0.3 95±3 kudu 19 -26.8±0.4 2.9±0.5 8±3 hair impala combined 36 -15.2±3.4 na 63±24 impala north 18 -17.8±2.8 na 44±20 impala south 18 -12.6±1.3 na 82±10 impala female 13 -16.6±3.8 na 53±27 impala male 22 -14.6±3.0 na 67±21 buffalo 51 -10.2±0.4 na 98±2 kudu 19 -23.5±0.5 na 4±3 sponnheimer.qxd 2005/12/09 10:30 page 102 a pure browser. similarly, faeces with a δ13c of –13.4 ‰ would indicate a pure grazing diet with a δ13c of –12.5 ‰. results and discussion both faeces and hair data show impala to be mixed-feeders and significantly different from buffalo and kudu (p < 0.0001) (table 1; fig. 1). the faeces data suggest impala have a diet of 53 % c4 vegetation (grass), whereas the hair data suggest a diet of 63 % grass. these results are consistent with previous studies of impala in kruger national park and elsewhere (van zyl 1965; hofmann & stewart 1973; monro 1980; smithers 1983; van rooyen 1992; meissner et al. 1996; vogel 1978; wronski 2002). the faecal nitrogen contents of impala, buffalo, and kudu are also significantly different (p < 0.0001) (table 1). befitting a mixed-feeder, impala faecal nitrogen concentrations are higher than those of buffalo, but lower than those of kudu. these findings are also consistent with a multiple year study of ungulate faecal nitrogen concentrations in kruger national park (grant et al. 2000). both faeces and hair data also show strong dietary differences between impala from the northern and southern regions of the park (p < 0.0001) (table 1; fig. 2), with southern impala grazing more than their northern counterparts. the faeces data suggest a diet of 41 % grass in the north and 63 % grass in the south. similarly, the hair data suggest a 44 % grass diet in the north and a 82 % grass diet in the south. the difference between the faeces and hair data in the south may reflect the fact that the hair data represent a long-term dietary average, whereas the faeces data largely reflect the season when they were collected (the beginning of the dry season). nonetheless, both data sets concur that impala have different diets in the mopane-dominated north than they do in the marula-knobthorn and bushwillow issn 0075-6458 103 koedoe 46/1 (2003) fig. 1. boxplot with the median 25-75 % and standard deviation of the dietary percentage of c4 grass consumed by impala, buffalo, and kudu in kruger national park. buffalo eat nearly 100 % c4 grass, kudu eat nearly 100 % browse, and impala clearly eat both. note the strong similarity between the hair and faeces data. fig. 2. boxplot with the median, 25-75 %, and standard deviation of the dietary percentage of c4 grass consumed by impala in the northern and southern regions of kruger national park. note that both hair and faeces data agree that impala graze more in the south. woodlands of the south (gertenbach 1983). intriguingly, there is no difference in the faecal nitrogen concentrations of impala in the north and south of the park (table 1). this supports the contention of meissner et al. (1996) that impala optimise their diets for quality forage irrespective of plant composition. sponnheimer.qxd 2005/12/09 10:30 page 103 because the hair samples were collected from darted individuals, we were also able to look for sex differences in diet. throughout the park males grazed 14 % more than females (table 1; fig. 3); however, this difference was only statistically significant in the southern portion of the park (p = 0.03). this provides further evidence for sex based dietary differences as observed by other researchers (wronski 2002). in future, it might also prove fruitful to explore possible dietary differences associated with disparate social structures. for instance, do males in bachelor herds and those with harems have divergent diets? conclusion in this study we were able to show regional and sex based dietary differences of impala in kruger national park. further stable isotope analysis should prove a powerful tool for supplementing traditional studies of wildlife diets. observational studies are very time-consuming, which necessarily limits koedoe 46/1 (2003) 104 issn 0075-6458 fig. 3. boxplot with the median, 25–75 %, and standard deviation of the dietary percentage of c4 grass consumed by male and female impala in the kruger national park. males clearly graze more than females throughout the park, although the difference is only significant in the south. the number of populations that can be observed. furthermore, although these studies provide rich detail about the variety of plants consumed and the time spent feeding on each diurnally, they cannot easily quantify the actual amounts of browse and graze consumed both day and night. in contrast, stable isotope analysis is very fast and can provide precise information on the amount of browse and graze digested. once samples have been obtained, preparation and analysis of 100 samples can be completed in a single day. thus, although stable isotope studies cannot be used to identify individual plant species consumed, they are ideal for quickly comparing the percentages of browse and graze consumed by different groups, whether they are intraor inter-populational. stable isotope analysis also has the advantage that it can be used to study extinct populations. the hair of mounted trophies and even fossilised teeth can be used to glean something of the diets of animals long since dead (e.g., lee-thorp & van der merwe 1987; macko et al. 1999; sponheimer et al. 1999). thus, we can use this tool not only to study the nutritional ecology of modern wildlife, but to study the development of modern dietary adaptations through time. moreover, emerging techniques will make it possible to derive even more ecological information from such analyses. for instance, sampling tail hair in small increments (< 1 cm), makes it possible to trace dietary change through time (o’connell & hedges 1999). this should prove particularly useful for documenting seasonal changes in browse and graze consumption. it has also been suggested that nitrogen isotopes in hair can be used as indicators of nutritional stress (hobson et al. 1993). given present capabilities and the increasing potential offered by constant technical advances in the field, stable isotope analysis should prove a valuable supplement to traditional studies in mammalian dietary ecology in the future. sponnheimer.qxd 2005/12/09 10:30 page 104 acknowledgments the authors would like to thank the rangers and veterinary staff of kruger national park for help in obtaining specimens. we would especially like to thank charles trennery, retha jansen van vuuren, obert mathebula, wilson sondelya, josiah baloyi, shela patrickson, adam west and yasmin rahman for help in the field, and john lanham for his invaluable assistance with the mass spectrometer. we also thank two anonymous reviewers for their helpful comments on the manuscript. references ambrose, s.h. & m.j. deniro. 1986. the isotopic ecology of east african mammals. oecologia 69: 395–406. cerling. t.e. & j.m. harris. 1999. carbon isotope fractionation between diet and bioapatite in ungulate mammals and implications for ecological and paleoecological studies. oecologia 120: 347–363. deniro, m.j., & s. epstein. 1978. influence of diet on the distribution of carbon isotopes in animals. geochimica et cosmochimica acta 42: 495–506. dunham, k.m. 1982. the foraging behaviour of impala aepyceros melampus. south african journal of wildlife research 12: 36–40. gertenbach, w.p.d. 1983. landscapes of the kruger national park. koedoe 26: 9–122. grant,c.c., m.j.s. peel, n. zambatis & j.b.j. van ryssen. 2000. nitrogen and phosphorus concentration in faeces: an indicator of range quality as a practical adjunct to existing evaluation methods. african journal of range and forage science 17: 81–92. hobson, k.a., r.t. alisauskas & r.g. clark. 1993. stable nitrogen isotope enrichment in avian tissues due to fasting and nutritional stress: implications for isotopic analysis of diet. condor 95: 388–394. hofmann, r. r. & d.r.m. stewart. 1972. grazer or browser: a classification based on the stomach structure and feeding habits of east african ruminants. mammalia 36: 226–240. jones, r.j., m.m. ludlow, j.h. troughton & c.g. blunt. 1981. changes in natural carbon isotope ratio of the hair from steers fed diets of c4, c3 and c4 species in sequence. search 12: 85–87. joubert, s.c.j. 1986. the kruger national park – an introduction. koedoe 29: 1–11. lamprey, h.f. 1963. ecological separation of the large mammal species in the tarangire game reserve, tanganyika. east african wildlife journal 1: 63–92. lee-thorp, j.a. & n.j. van der merwe. 1987. carbon isotope analysis of fossil bone apatite. south african journal of science 83: 712–715. macko, s. a., m. h. engel, v. andrusevich, g. lubec, t. c. o’connell & r. e. m. hedges. 1999. documenting the diet of ancient human populations through stable isotope analysis of hair. philosophical transactions of the royal society of london b 354: 65–76. meissner, h.h., e. pieterse & j.h.j. potgieter. 1996. seasonal food selection and intake by male impala aepyceros melampus in two habitats. south african journal of wildlife research 26: 56–63. minson, d.j. 1990. forage in ruminant nutrition. san diego: academic press. monro, r.h. 1980. observations on the feeding ecology of impala. south african journal of zoology 15: 107–110. o’connell, t.c. & r.e.m. hedges. 1999. investigations into the effect of diet on modern human hair isotopic values. american journal of physical anthropology 108: 409–425. saitoh, m., m. uzuka, m. sakamoto & t. kobori. 1969. rate of hair growth. pp. 183–201. in: w. montagna & r. l. dobson (eds.). hair growth. oxford: pergamon press. schoeninger, m.j., j. moore & j.m. sept. 1999. subsistence strategies of two savanna chimpanzee populations: the stable isotope evidence. american journal of primatology 49: 297–314. smith, b.n. & s. epstein. 1971. two categories of 13c/12c ratios for higher plants. plant physiology 47: 380–384. smithers, r.h.n. 1983. the mammals of the southern african subregion. pretoria: university of pretoria. sponheimer, m., k. reed & j.a. lee-thorp. 1999. combining isotopic and ecomorphological data to refine bovid paleodietary reconstruction: a case study from the makapansgat limeworks hominin locality. journal of human evolution 36: 705–718. tieszen, l.l. & s.k. imbaba. 1980. photosynthetic systems, carbon isotope discrimination, and herbivore selectivity in kenya. african journal of ecology 18: 237–242. van der merwe, n.j., j.a. lee-thorp & r.h.v. bell. 1988. carbon isotopes as indicators of elephant diets and african environments. african journal of ecology 26: 163–172. van zyl, j.h.m. 1965. the vegetation of the s.a. lombard nature reserve and its utilization by certain antelope. zoologica africana 1: 55–71. van rooyen, a.f. 1992. diets of impala and nyala in two game reserves in natal, south africa. south african journal of wildlife research 22: 98–101. issn 0075-6458 105 koedoe 46/1 (2003) sponnheimer.qxd 2005/12/09 10:30 page 105 vogel, j. 1978. isotopic assessment of the dietary habits of ungulates. south african journal of science 74: 298–301. vogel, j.c., a. fuls & r.p. ellis. 1978. the geographical distribution of kranz grasses in south africa. south african journal of science 74: 209–215. witt, g.b. & r.j. beeton. 1997. sheep faeces under shearing sheds: a documentary of vegetation change using stable carbon isotope analysis. rangeland journal 19: 109–115. witt, g.b., e.j. moll, r.j. beeton & p.j. murray. 1998. isotopes, wool and rangeland monitoring: let the sheep do the sampling. environmental management 22: 145–152. wronski, t. 2002. feeding ecology and foraging behaviour of impala aepyceros melampus in lake mburu national park, uganda. african journal of ecology 40: 205–211. koedoe 46/1 (2003) 106 issn 0075-6458 sponnheimer.qxd 2005/12/09 10:30 page 106 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <feff00550073006100720065002000710075006500730074006500200069006d0070006f007300740061007a0069006f006e00690020007000650072002000630072006500610072006500200064006f00630075006d0065006e00740069002000500044004600200063006f006e00200075006e00610020007200690073006f006c0075007a0069006f006e00650020006d0061006700670069006f00720065002000700065007200200075006e00610020007100750061006c0069007400e00020006400690020007000720065007300740061006d007000610020006d00690067006c0069006f00720065002e0020004900200064006f00630075006d0065006e00740069002000500044004600200070006f00730073006f006e006f0020006500730073006500720065002000610070006500720074006900200063006f006e0020004100630072006f00620061007400200065002000520065006100640065007200200035002e003000200065002000760065007200730069006f006e006900200073007500630063006500730073006900760065002e002000510075006500730074006500200069006d0070006f007300740061007a0069006f006e006900200072006900630068006900650064006f006e006f0020006c002700750073006f00200064006900200066006f006e007400200069006e0063006f00720070006f0072006100740069002e> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 kyle.qxd the status of and some notes on the gaboon adder bitis gabonica in northern kwazulu/natal, south africa and southern mozambique r. kyle issn 0075-6458 113 koedoe 46/2 (2003) a lot of work has been carried out on the gaboon adder (bitis gabonica) in kwazulu/ natal (see botbijl 1994a, 1994b), but the status of the snake has remained unclear. the distribution of this snake is associated with tropical coastal forest and thickets, woodland, forest-savanna mosaic, wellwooded savanna and forests (spawls et al. 2002) extending from nigeria through central to east africa and southwards to northern kwazulu/natal (branch 1998). this distribution, especially towards the south, is very disjunct and the closest record to the north of the southern african population is from about 450 km in central mozambique (broadley 1983). the snake is reported ‘rare’ in the region (branch 1998) and is described as ‘vulnerable’ in the south african reptile red data book (branch 1988). in south africa, it has been reported from localities between mtubatuba in the south and kosi bay in the north (fig. 1) (bruton 1982; bruton & haacke 1980; broadley1983; branch 1988, 1998; spawls & branch 1995). on the one hand, this area has been subject to large-scale forest destruction and on the other, has become part of the greater st. lucia world heritage site. botbijl (1994b) considered b. gabonica threatened by large-scale habitat destruction as well as persistent collection. most of the intense work focussed on the dukuduku forest, which is only a small part of its known range (botbijl 1994a & 1994b.). in 1995, ezemvelo kwazulu/natal wildlife (ekznw) approved a project to translocate as many b. gabonica as possible from dukuduku forest, to umlalazi nature reserve, since dukuduku forest was under serious threat from habitat degradation. although outside its known range, umlalazi was considered suitable habitat and was under conservation management (botbijl 1996). initially the aims of the project were to capture and translocate 80 snakes (botbijl 1994b). the translocations were initially infrequent, but increased in frequency until the project was stopped, by ekznw (armstrong, a., ezemvelo kwazulu/natal wildlife, po box 13053, cascades, 3201 pers. comm.) in 1998. by this time approximately 200 snakes had been caught, paid for and translocated. the project was terminated due to, among others, concerns about the suitability of umlalazi, the survival of translocated snakes, and the creation of a market (at r200 each) for a rare animal. the remaining snakes in captivity were released on the eastern shores of lake st. lucia. bitis gabonica is a cryptically coloured snake that, despite its size, is extremely rarely seen, even though it is probably relatively abundant in some areas. although it often moves off when encountered, it can bury in leaf litter or even sand (pers. obs.) with only its head exposed. many local residents in the kosi bay area, who spend much of their lives in the forests, are not familiar with the species, and are surprised when informed of its presence near to their homes. natural history note kyle.qxd 2005/12/09 11:18 page 113 man has markedly modified much of dukuduku forest, the southern known locality in the distribution of b. gabonica and an area of high snake density, and snake numbers may be drastically reduced. although 200 individuals have been released in umlalazi nature reserve, no sightings of b. gabonica have been reported in the reserve subsequent to the project. there have been no confirmed reports of the snakes between umlalazi and dukuduku, despite large-scale dune mining operations in the area. the area north of dukuduku, koedoe 46/2 (2003) 114 issn 0075-6458 from st. lucia to the mozambique border, has progressively improved in conservation status over the last few decades. almost all known sightings of b. gabonica have been made in this coastal strip. surveying for the presence of the snake has proved particularly unsuccessful (botbijl 1994a) and the only information available is from accidental or incidental sightings. these are clearly biased in favour of areas of high human activity. most records were initially from dukuduku forest, through which a major road was cut, resulting in many snakes being killed or collected on the road. recent habitat destruction of the forest also resulted in many encounters. sightings have, however, also been made at virtually all the locations where conservation staff have been active between st lucia town and the international border (bruton & haacke 1980; broadley 1983; branch 1988 & branch 1998; spawls & branch 1995). other sightings have been made on roads through the coastal strip and during alien plant eradication. there have been four sightings outside the coastal strip. in 1983, workers fencing manguzi forest killed an adult. three separate sightings have been made subsequently in the garden of a manguzi shop owner (pers. obs.). i have lived at kosi bay for twenty-three years and have never seen a ‘wild’ b. gabonica, and yet they have been found in the campsite less than three hundred metres from my residence. since this species has been recorded throughout the area wherever conservation staff has been active, it seems likely that it occurs in suitable habitat throughout the coastal strip. it also seems probable that within this range, there will be areas of relative high abundance. fig. 1. map of northern kwazulu/natal showing the main conservation protected areas and localities important in the known distribution of b. gabonica. kyle.qxd 2005/12/09 11:18 page 114 residents of enkovukeni, a region on the eastern shores of kosi bay where there are no confirmed records, also report this species from several locations in the area. when a photograph of b. gabonica was shown to people clearing the invasive alien plant pereskia aculeata from the area south west of kosi bay, they not only recognised the snake, but also reported killing one while clearing exotic vegetation. relying on identification of snakes, particularly venomous species, by lay people is generally not dependable. in this instance, however, although young b. gabonica could be confused with puffadders, bitis arietans, adult b. gabonica should be easily distinguished as they are much larger than b. arietans. north of the southern mozambique border there were no known records. however, the habitat appears to be suitable and to continue uninterrupted to just south of inhaca island. recently, there have been unconfirmed reports from the dive camp at ponta do ouro and further north (pers. obs.). as more of the coastal bush is destroyed in southern mozambique, it is likely that further records, extending its known distribution to the north, will be made. botbijl (1994a) suggests that b. gabonica may occur at low densities and gave the total south african population an estimated upper limit of 500. however, the 200 snakes that were caught during the translocation project, almost all from dukuduku, suggests that population densities are much higher than botbijl's estimations. the intrusions into and progressive destruction of dukuduku forest opened a window and focussed attention on b. gabonica in that area. it seems unlikely that the areas that form the peripheral of the distribution contain the majority of its animals. recent b. gabonica sightings from st. lucia, cape vidal, manzengwenya, sodwana bay and kosi bay suggest the continued presence and breeding of the snake throughout the almost 200 km coastal strip. the report from mozambique further suggests that the species may well extend significantly into that country. recent evidence, such as the may 2002 capture of a hatchling in a st. lucia town garden (dickson, e., ezemvelo kwazulu/natal wildlife, po box 1, kwangwanase, 3973 pers. comm.), the continued manguzi town sightings, and two recent records from inside sodwana bay camp, suggest that the species shows an amazing capacity to survive unnoticed very close to human activity. in summary, it appears that there is probably a secure, though possibly often low density, population of b. gabonica along almost the entire coastal strip from st. lucia mouth to the mozambique border. attempts to ‘save’ the species in south africa by relocating all wild specimens to umlalazi were thus possibly misdirected and indicative of our tendency to over-manage in situations where data are lacking. habitat destruction, unnatural fires and road kills remain important threats to the conservation status of b. gabonica, collection for the international pet and traditional medicine trade are also likely threats. although i do not suggest lowering the conservation status of the snake in south africa by its removal from the ‘red data list’ (branch 1988), i am of the opinion that previous population and range estimates seriously underestimate the distribution and abundance of this species. acknowledgements i thank ezemvelo kwazulu/natal wildlife for permission to publish this paper. references botbijl, t. 1994a. the autecology of the gaboon adder, bitis gabonica in zululand. msc. thesis. university of natal pietermaritzburg. botbijl, t. 1994b. proceedings of the gaboon adder seminar st. lucia, 22 april 1994. natal parks board. unpublished internal report. botbijl, t. 1996. report on gaboon adder habitat assessment. natal parks board. unpublished internal report. issn 0075-6458 115 koedoe 46/2 (2003) kyle.qxd 2005/12/09 11:18 page 115 branch, w.r. (ed.). 1988. south african red data book reptiles & amphibians. pretoria: council for scientific and industrial research, national scientific programmes unit. (south african national scientific programmes report; no. 151). branch, w.r. 1998. field guide to snakes and other reptiles of southern africa. cape town: struik. broadley, d.g. 1983. fitzsimon's snakes of southern africa. cape town: delta. bruton m.n. 1982. uncommon and rare reptiles of maputaland. african wildlife 36 (4/5) 184-185. bruton m.n. & w.d. haacke. 1980. the reptiles of maputaland. pp 251-287. in: bruton, m.n. & k.h. cooper (eds.). studies on the ecology of maputaland. rhodes university and natal branch of the wildlife society of southern africa. spawls, s. & w. r. branch. 1995. dangerous snakes of africa. halfway house: southern books. spawls, s., k. howell, r. drewes & j. ashe. 2002. a field guide to the reptiles of east africa. london: academic press. r. kyle, p.o. box 43, kwangwanase, 3973, south africa. (ezemvelo kwazulu natal wildlife; rkyle@ iafrica.com). koedoe 46/2 (2003) 116 issn 0075-6458 kyle.qxd 2005/12/09 11:18 page 116 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <feff004700650062007200750069006b002000640065007a006500200069006e007300740065006c006c0069006e00670065006e0020006f006d0020005000440046002d0064006f00630075006d0065006e00740065006e0020007400650020006d0061006b0065006e0020006d00650074002000650065006e00200068006f00670065002000610066006200650065006c00640069006e00670073007200650073006f006c007500740069006500200076006f006f0072002000610066006400720075006b006b0065006e0020006d0065007400200068006f006700650020006b00770061006c0069007400650069007400200069006e002000650065006e002000700072006500700072006500730073002d006f006d0067006500760069006e0067002e0020004400650020005000440046002d0064006f00630075006d0065006e00740065006e0020006b0075006e006e0065006e00200077006f007200640065006e002000670065006f00700065006e00640020006d006500740020004100630072006f00620061007400200065006e002000520065006100640065007200200035002e003000200065006e00200068006f006700650072002e002000420069006a002000640065007a006500200069006e007300740065006c006c0069006e00670020006d006f006500740065006e00200066006f006e007400730020007a0069006a006e00200069006e006700650073006c006f00740065006e002e> /esp <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> /suo <feff004e00e4006900640065006e002000610073006500740075007300740065006e0020006100760075006c006c006100200076006f0069006400610061006e0020006c0075006f006400610020005000440046002d0061007300690061006b00690072006a006f006a0061002c0020006a006f006900640065006e002000740075006c006f0073007400750073006c00610061007400750020006f006e0020006b006f0072006b006500610020006a00610020006b007500760061006e0020007400610072006b006b007500750073002000730075007500720069002e0020005000440046002d0061007300690061006b00690072006a0061007400200076006f0069006400610061006e0020006100760061007400610020004100630072006f006200610074002d0020006a0061002000520065006100640065007200200035002e00300020002d006f0068006a0065006c006d0061006c006c0061002000740061006900200075007500640065006d006d0061006c006c0061002000760065007200730069006f006c006c0061002e0020004e00e4006d00e4002000610073006500740075006b0073006500740020006500640065006c006c00790074007400e4007600e4007400200066006f006e0074007400690065006e002000750070006f00740075007300740061002e> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! abstract introduction methods conceptual framework of interconnectivity of the catena ecosystem synthesis of research findings future research conclusion acknowledgements references about the author(s) beanelri b. janecke department of animal, wildlife and grassland sciences, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa johan van tol department of soil, crop and climate sciences, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa izak p.j. smit south african national parks, scientific services, skukuza, south africa centre for african ecology, school of animal, plant and environmental sciences, university of the witwatersrand, johannesburg, south africa andri c. van aardt department of plant sciences, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa edward s. riddell south african national parks, conservation management, skukuza, south africa centre for water resources research, university of kwazulu-natal, pietermaritzburg, south africa maitland t. seaman centre for environmental management, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa wijnand j. swart department of plant sciences, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa pieter j. du preez † department of plant sciences, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa pieter a.l. le roux digital soils africa, south africa previously institute for groundwater studies, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa citation janecke, b.b., van tol, j., smit, i.p.j., van aardt, a.c., riddell, e.s., seaman, m.t. et al., 2020, ‘biotic and abiotic connections on a granitic catena: framework for multidisciplinary research’, koedoe 62(2), a1600. https://doi.org/10.4102/koedoe.v62i2.1600 note: special issue: connections between abiotic and biotic components of a granite catena ecosystem in kruger national park, sub-edited by beanelri janecke and johan van tol. †, 1960–2019. original research biotic and abiotic connections on a granitic catena: framework for multidisciplinary research beanelri b. janecke, johan van tol, izak p.j. smit, andri c. van aardt, edward s. riddell, maitland t. seaman, wijnand j. swart, pieter j. du preez, pieter a.l. le roux received: 01 oct. 2019; accepted: 17 apr. 2020; published: 29 oct. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract local environmental gradients on a catenal scale create ecological patterns from the crest to the stream of the hillslope. bottom-up drivers interact with top-down controls to give rise to these patterns. a multidisciplinary project was conducted to study the processes that govern functioning, structure and heterogeneity on a catena in a third-order catchment in the southern granite supersite in the kruger national park. the project included abiotic components (e.g. groundwater-surface water interactions, soil chemical and physical properties) as well as biotic components (e.g. soil microbes, small aquatic organisms in ephemeral pools, plant communities, vegetation structure and mammal diversity). each of these components was investigated in detail along the catenal gradient and reported on in separate articles in this special issue. the drought of 2015–2016 occurred during the sampling period of the study and information on the response of vegetation and mammals to the drought were included. in this article, a synthesis of findings from the separate components or disciplines is provided to highlight the interactive functioning and ecological patterns of the catena. these findings were then used to develop a framework for multidisciplinary studies in similar environments. the framework highlights the interactive relationships between various components of the ecosystem and the importance of a multidisciplinary approach. conservation implications: the findings of this study were used to develop a conceptual framework outlining how a range of biotic and abiotic patterns and processes interact along the catenal gradient. the framework highlights the importance of recognising these interactions in a multidisciplinary approach focused on one supersite. keywords: climate; hydrology; interdisciplinary studies; mammals and mud wallows; trophic levels. introduction one of the key concepts linking ecology and biodiversity management is environmental heterogeneity. in the words of pickett, cadenasso and benning (2003): the general concept of heterogeneity becomes ecologically meaningful when it is given operational life by specifying what agents modify what substrates, what controls the modification, and what organisms or processes respond to the resulting spatial template. (p. 25) the concept of savanna heterogeneity has been fundamental in how this has informed the management of the kruger national park (knp), and has influenced, for example, how fire, artificial surface water and large herbivores are managed to maintain, mimic or, in some cases, restore inherent heterogeneity. for an overview of the concepts of heterogeneity in the savanna context on different levels from a fine to a broader scale, and how that has influenced thinking and management of the knp, see du toit, rogers and biggs (ed. 2003) and rogers (2003). ‘heterogeneity’ is a broad term, but can comprise differences and interactions between soil types and properties, vegetation composition and structure, patchiness and patterns, sub-habitats, animal presence and so on. at the scale of our study (in this instance, a third-order catchment in a granitic landscape), heterogeneity is the outcome of local environmental gradients along a hillslope (from the crest to the valley bottom). this spatial heterogeneity typically exhibits a common form of organisation and symmetry – termed the catena (chain) concept. the term was originally used by milne (1936) to describe the soil distribution patterns in east africa, but can be expanded to include processes causing differentiation along the hillslope and processes causing vertical differentiation in ecological properties. although the patterns are normally associated with gradients (e.g. gradual increase in moisture downslope), it can also include sharp transitions caused by geology and geological features (e.g. seepline and alluvial deposits). bottom-up drivers (like climate and soils), together with top-down controls (like fire and herbivory), interact to give rise to the emergent heterogeneity and patterning. this is illustrated in the article by sankaran et al. (2005), where mean annual precipitation (bottom-up environmental gradient) influences the upper asymptote of tree cover, whilst top-down disturbance factors (like fire and herbivory) and other bottom-up factors (i.e. soil and climate) explain the large variability observed under the asymptote. soils interact with biology, the soil is a habitat for the biota and the biota is controlled by the macroand microclimate above ground and water and nutrient supply from below ground. these relationships between environmental controls create soil distribution patterns with specific characteristics (e.g. hydrological, structural and chemical), which partly act as abiotic templates determining the vegetation above ground on a catena. the resulting vegetation patterns provide heterogeneity in terms of food and habitat for a range of animals (e.g. from open grassy areas for grazers to dense wooded areas for browsers and including everything in between) (smit & prins 2015). this model of the biodiversity–heterogeneity nexus rests on ideas about the nature of the ecological community, species assemblage processes and niche construction. whilst savanna science has generated a broad conceptual framework of the hierarchy and processes of heterogeneity and its management in the knp, there is still a need to improve our understanding of ecological patterns and interactions between different components of the ecosystem in granitic catenas. with this as background, a multidisciplinary project was conducted that specifically focused on hydrology, soil, microbial, small aquatic organisms (in ephemeral pans and mud wallows), vegetation, mammal diversity and heterogeneity linked to below ground–above ground relationships along the catena or hillslope. each of these different components of the ecosystem was researched separately and published as individual articles in a special issue, which this article (as part of the same special issue) aims to integrate. the objectives of this article were (1) to provide a framework of ecological patterns and interactions linking certain abiotic and biotic components of the catena ecosystem and (2) to briefly synthesise the main findings from the individual multidisciplinary research articles published in this special issue. a brief overview of the study area is provided together with a concise report of methodological approaches followed during the multidisciplinary project. detailed descriptions of materials and methods can be found in the individual articles. methods study area and surveys the study was conducted in knp on the southern granite supersite (stevenson-hamilton supersite) (smit et al. 2013). the supersite is considered a ‘typical’ granite ecosystem in southern knp and has previously been identified as a focal site for multi-scaled and multi-disciplinary research to understand abiotic and biotic patterns and processes of these granitic landscapes (smit 2020). the project focused on one specific catena of a third-order catchment between 25°06’28.6s, 31°34’41.9e and 25°06’25.7s, 31°34’33.7e near skukuza. this catchment eventually drains into the perennial sabie river further to the north. the multidisciplinary project continued on earlier work conducted by riddell et al. (2014) focusing on groundwater-surface water interactions and included the same catena as part of the third-order catchment. the catena is approximately 500 m from top (crest) to bottom (drainage line) with four hydrometric monitoring stations that recorded soil matric potential at different depths (riddell et al. 2014; riddell et al. 2020). a hydropedological survey was conducted along the belt transect, from crest to drainage line, which included 49 soil observations (10 m intervals). the hydropedological survey included detailed morphological description and classification of the soils as well as in situ measuring of hydraulic properties. this was supplemented with sampling for laboratory determination of physical and chemical properties of the soils. soil information was used to derive a conceptual model of hillslope hydrological response (bouwer, le roux & van tol 2020). plants were classified based on species composition and abundance and grouped into communities for the same 49 plots in the belt transect where the soil observations were made (theron, van aardt & du preez 2020). vegetation composition and structure were also determined (janecke 2020), as well as wildlife habitat and mammal species presence that were noted through camera trapping (janecke & bolton 2020). temporary surface water sources (i.e. small ephemeral pans, animal-created wallows and other vernal pools) sustained by hillslope interflow were sampled for small aquatic organisms and their resting stages, while soil samples were taken of mud in depressions, of soils in the belt transect on the catena, and of soil associated with plant roots. next-generation sequencing and isolations were used to explore potential differences in fungal (gryzenhout et al. 2020) and bacterial communities (vermeulen, cason & swart 2020) in the rhizosperes (root area) across different catenal positions with different resulting soil chemical compositions. extreme drought conditions coincided with the study period during one of the strongest el niño southern oscillations in the historical record (hu & fedorov 2017; malherbe et al. 2020; swemmer et al. 2018). the impacts of this drought started during late 2014 when rainfall at the nearest rainfall station (skukuza; approximately 13 km north of study site) for the 2014–2015 rainfall season was 350.4 mm (63.7% of long-term average). it became severe from 2016 onwards when total rainfall was very low during the 2015–2016 rainfall season (194.2 mm, 35.3% of long-term average) (sanparks unpublished weather data). in march 2016, after a big rainfall event in the midst of the drought, some soil, mud and water samples were taken. during october 2016, barely any herbaceous vegetation remained and there was no standing water at the study site. herbaceous vegetation cover, plant species composition and large animal presence (on a species basis) were, however, still recorded annually from 2015 to 2017 (janecke 2020, janecke & bolton 2020, theron et al. 2020). although the drought did present challenges in terms of achievement of our original objectives, it also created an opportunity to study such an event, with a general paucity in drought-related studies in savannas (i.e. smit & bond 2020, theron et al. 2020). as such, an opportunity arose to study how the vegetation on the catena would recover after the drought (van aardt et al. 2020). the vegetation at the study site was therefore surveyed in the growing seasons of 2015 (beginning of drought) to 2018 (post-drought). developing the framework of interconnectivity the framework to highlight the interconnectivity between the various components of this ecosystem was developed through several discussions and workshops between the various researchers. where available, the literature on similar studies was consulted to identify relationships between various components. although some of the main components of ecosystem connectivity were addressed in this study, the framework also identified other relationships which should be addressed in future research. conceptual framework of interconnectivity of the catena ecosystem a catena is a soil sequence where each soil group occurs on similar parent material, but in arrays from the crest to the footslope along a hillslope (brady & weil 2002). differences in elevation and in water flow patterns can lead to the transport of dissolved (i.e. salts) and suspended substances from higherto lower-lying areas. at the midslope (localised) and/or footslope of these catenas, a clay layer occurs as a less permeable barrier that significantly retards flow, where water seeps through the saprolite on the surface which evaporates after some time – these areas are called ‘seeps’ or ‘seep zones’. downslope of the seeps, sodium accumulates to create sodic areas if rainfall is not sufficient to cause leaching of the accumulated salts (brady & weil 2002; khomo & rogers 2005). the southern granites of knp can have well, moderately or poorly drained soil forms. the spatial distribution is dominantly associated with terrain morphology and depends on a combination of soil forming factors (e.g. parent material and topography). all of this results in varying types of fertility along the catena that forms different gradients of supply to the soil biota. the heterogeneity in soil fulfils different hydrological functions. firstly, it controls infiltration rate that determines runoff. secondly, it controls plant water-holding capacity. thirdly, it regulates sub-surface lateral water flow in sandy crests with low clay content. fourthly, it serves as a clay plug and creates surface water flow in clayey areas down the slope where permeability of soil restricts drainage and infiltration (khomo et al. 2011). these effects of soil texture, soil moisture and soil water-holding capacity are expressed in distribution and composition of vegetation along a catena (theron et al. 2020). the catena thus forms an ecosystem that consists of abiotic (gradient of soil types, soil properties and nutrients; ground water flow; surface water; topography; micro-climate; etc.) and biotic components (micro-organisms, plants and animals) that are connected and interact in the different zones present along the hillslope. the focus of this section is to describe the interconnectivity between some of the components of the catena ecosystem that formed part of a multidisciplinary study conducted on one catena or hillslope. this article summarises only the basic connections between soil, hydrology, micro-organisms in the rhizosphere and roots of dominant vegetation, vegetation structure, mammals, permanent and temporary water sources, and drought in the catena ecosystem. the readers are referred to the specific articles in the special issue for further detail. figure 1 shows a simplified overview of how the abiotic and biotic components that formed part of this study are interlinked. other important taxa (e.g. detritus feeders, decomposers, birds, reptiles, etc.) and processes (e.g. fire) that did not form part of the study, but are deemed critical in the savanna biome, were included using dotted line blocks in figure 1. note, however, that no connections between these taxa and other components were indicated as it falls beyond the scope of what was studied here (figure 1). no fire went through the study area during the time of this study and the mean fire return interval is 5.8 years for the area (smit et al. 2013). the impact of elephants is also included in the discussion, but not indicated separately in the figure as it is implicitly part of the ‘herbivores’ and ‘mammals’ components. some of the ecosystem components have been lumped together in figure 2 to illustrate the interconnectivity in more detail (i.e. figure 1 focuses on ecosystem components in some detail, whilst figure 2 highlights ecosystem connections). figure 1: links between abiotic and biotic components of the savanna granitic catena ecosystem. ecosystem components that did not form part of this study are indicated in dotted line blocks, but only fire has been linked to other components. a connection to soil or plants implies its sub-components as well. figure 2: interconnectivity of components that formed part of this study on a granitic catena ecosystem in the savanna. the following discussion refers to the numbered arrows in figure 2 – the arrows represent relationships between the various ecosystem components covered in this study. climate is one of the key drivers of patterns, processes and heterogeneity. climate, which includes inter alia precipitation (volume and intensity), temperature, radiation and humidity, determines hydrological processes (1), as well as soil properties (10), micro-organism activity (they may become dormant during dry conditions) (11), vegetation types (19), presence of mammals (20) and standing water in the temporary mud wallows and vernal, ephemeral pools (15). soil is a first-order control of hydrological processes as it governs infiltration rates, storage and drainage (2). the relationship between soil and hydrological processes is, however, interactive, as water plays a primary role in the formation of the morphological, chemical and biological properties of soil (2). these interactions between soil and water are known as hydropedology. the soil distribution pattern is determined by five soil-forming factors of which geology and topography are key factors at the hillslope scale (3). in combination with the soil distribution, the topography affects hydrological processes (4). nutrients, water and anchorage are key provisioning functions of soils to plants (5). different soils have specific abilities to support plant species and vitality of growth; thus, the soil distribution pattern is a key determinant in the distribution of various vegetation communities at catenal or hillslope scale. plants, on the contrary, influence soil processes like infiltration and provide soil with organic material and nutrients through their leaves and root systems. mining of nutrients from the subsoil and fractured rock is an important biocycling process that brings nutrients from deep soil layers and returns it to the surface, which determines the chemical, physical and biological properties of soil. in combination with the soil distribution pattern (as impacted by topography, geology and hydrology), micro-climate and micro-topography can influence soil water availability that governs vegetation distribution in savannas (5). the patchiness of the vegetation is a common occurrence in savannas and is observed at specific scales. vegetation are grouped in patterns because of species composition and plant communities growing in the same local patch, presence of bare and vegetated patches, sub-canopied habitats that differ from open uncanopied areas, varying vegetation structure, etc. spatial patterns can be described based on the size of plants and their canopies (vegetation structure), as an example. sub-canopy shaded habitats are not only important for thermal regulation of animals, but also are preferred areas of grazing (figure 2, number 7) because large woody plants change the availability of resources to the herbaceous layer by mostly creating patches of higher nutrition and moisture under the canopies of especially large trees (treydte et al. 2011). large trees (> 5 m) are important in the environment, but declines in their numbers have been reported in knp (eckhardt, van wilgen & biggs 2000). density of large trees on granites has decreased by 15% between 1984 and 1996 (whyte, van aarde & pimm 2003). whyte et al. (2003) described a change in the structural diversity (but not in woody species diversity) in which woody vegetation was converted to a short woodland (with higher density of small trees and shrubs), because of a declining density of large trees. according to trollope et al. (1988) and whyte et al. (2003), elephants were primarily responsible for killing trees larger than 3 m and large tree recruitment was prevented by increase in frequency of fires. in the nkuhlu exclosures (also on a granite catena but next to the perennial sabie river) where herbivores are excluded through fences, the density and number of big trees are higher (siebert & eckardt 2008). the proximity to the river may also play a role in the presence of big trees because of higher probability of sub-surface water. using aerial light detection and ranging (lidar) collected over time, it was determined that the rate of treefall was sixfold higher in areas outside of herbivore exclosures where elephants had direct access, as compared to areas within the exclosures which was protected from elephants (asner & levick 2012). this pattern seemingly also plays out over larger landscape scales (asner et al. 2016). not only does vegetation influence herbivore spatio-temporal patterns, but herbivores also influence vegetation patterns. number 8 in figure 2 implies the impact of all mammalian herbivores (including grazers, browsers, mixed feeders and the mega-herbivores) on vegetation structure and species composition. the impact of herbivores on vegetation composition and structure is not restricted to woody vegetation as described above. increased and frequent grazing of the herbaceous layer often result in changing grass species composition and structure (i.e. from tall bunch grasses to short stoloniferous grazing lawns; grant et al. 2019; hempson et al. 2015). the variation in spatial distribution of quality and quantity of forage contributes to the different ways that various large herbivores feed at a local scale. the density of animals, social structure, animal behaviour, palatability of plants, competition and forage regrowth patterns may also play a role. grazing that varies spatially can potentially limit the expression of vegetation distribution patterns that are related to edaphic factors that operate at larger scales (adler, raff & lauenroth 2001; augustine 2003). the relationship between vegetation and mammals is also interactive (7 and 8, figure 2): vegetation (plant species, palatability and abundance) is not only the source of food for herbivores, but also limits their numbers (plant defence mechanisms, availability of food – bottom-up control), which will determine the occurrence and behaviour of animals on different areas on the catena (7). herbivores can stimulate plant growth through pruning or inhibit growth through overutilisation (i.e. mcnaughton 1983; milewski & madden 2006). vegetation also provides shelter against elements and against predators, or it can increase predation risk (limited visibility). the strategy of some animals, such as bushbuck, is probably to ‘hide’ from predators in dense undergrowth, whereas other species like wildebeest prefer open areas to see predators. vegetation thus plays an important role when considering the landscape of fear created by the possible presence of predators in a denser area with lower visibility (i.e. laundre, hernandez & ripple 2010; nasar & jones 1997). as the vegetation (as food and habitat) would impact the occurrence and abundance of herbivores, it is likely that it would also impact the occurrence of carnivores that feed on the former. the presence of carnivores can have a top-down effect on the ecosystem (figure 1) and thus can impact the assemblage structure and space-use patterns of herbivores in an area (through creating landscapes of fear in areas that are mostly avoided by herbivores versus areas with good visibility frequented by herbivores) (e.g. le roux, kerley & cromsigt 2018). herbivores feed on and interact with the vegetation and impact the soil (9) (through trampling, nutrient enrichment through dung, digging holes and causing erosion through intense utilisation of vegetation); therefore, lower herbivore numbers (regulated top-down by carnivores), or changed space-use patterns (e.g. herbivores aggregating in open areas at night to reduce predation risk – burkepile et al. 2013), can have cascading effects on vegetation and soil. large herbivores (especially elephants) can have a direct impact on the vegetation structure (e.g. uproot trees) and aid in creating micro-habitats (augustine 2003) underneath fallen trees or scattered branches left behind after feeding (8). uprooting of trees by elephants would also impact the soil properties (9, figure 2) through changing the physical properties of soil (e.g. decrease bulk density, increase infiltration rates and bring rocks up to the topsoil). herds of elephant and rhino could result in soil compaction, whilst hoofed animals can often break soil crusting and therefore actually loosen soil. these animals can also provide valuable nutrients in the form of excretions and decomposing carcasses. soil further provides a habitat for soil organisms and a substrate for plants to grow in as food source for smaller animals (mice, moles, hares, etc.), whilst the animals, in turn, increase macro-porosity of the soils through tunnelling (9) (these processes were not specifically researched in the current study). spatial patterns consisting of vegetated patches that alternate with bare soil areas are characteristic in savannas (augustine 2003; rietkerk et al. 2000). these patches can vary greatly in size and can be attributed to interactions between biotic (trampling, bulk and selective grazing, animal densities, competition, seed dispersal, etc.) and abiotic factors (variation in topography, surface water run-off, soil types and depth, soil properties, rainfall patterns that vary, microclimate, etc.) (augustine 2003). sodic patches are another example of large open areas that are more sparsely vegetated than surrounding areas on the catena. khomo and rogers (2005) used evapotranspiration models for upland-based catena and riparian-based areas to predict vegetation zonation and found that zonation of catena vegetation is dictated by a gradient in salt tolerance, whilst an increase in the surface area of the sodic patch is evident over time as a result of this progressive salt accumulation. rietkerk et al. (1997) described a model where a feedback occurs between vegetation cover and water infiltration, or retention of nutrients. thus, with increased grazing intensity, which reduces plant cover beyond a certain threshold, the feedbacks between plants and soil ‘can create a stable, degraded bare state which cannot be reversed simply by reducing grazing pressure’ (augustine 2003:320). however, the opposite is also true under certain conditions; for example, on sodic sites, certain conditions give rise to highly productive grazing lawns that are dependent on high-intensity grazing for their formation and maintenance. these grazing lawns and/or other regularly utilised patches can be important for certain animal species (see grant et al. 2019). the properties of soils on these nutrient hotspots and the processes of the catena could, however, also explain the bare soil areas, plant species composition and herbaceous plant density found in the sodic zone on the lower midslope (number 5, figure 2). climate, especially temperature and moisture, has a direct impact on soil biochemical reactions (whalen & sampedro 2010), such as mineralisation and/or accumulation of organic matter contents (carbon sequestration; brady & weil 2002) (10, figure 2). these reactions are largely driven by the microbial composition of the soil, which, in turn, is also directly influenced by the climate (11) (i.e. classen et al. 2015) as well as the soil properties (12) and fire (figure 1). the activity of fungi and bacteria would be impacted inter alia by the organic matter content, ph and the soil water content. the microbial abundance and activity would further be impacted by the plant community which provides specific conditions in the rhizosphere where certain microbial species can flourish (13). microbes, together with the vegetation in which root zones they occur, can be impacted by soil moisture and nitrogen availability. february and higgins (2010) monitored soil moisture levels continuously at satara and pretoriuskop in knp and found that surface layers are drier and on average exceed the wilting point for fewer days per year than the deeper layers. the negative correlation of root biomass with soil moisture is a representation of the dynamic process of removal of water through evaporation in upper soil layers and moisture that might actually be lower where highest root biomass and uptake capacity occur. they also reported a decrease in total soil nitrogen for the top 40 cm of the soil profile. thus, the distribution of roots may primarily be responding to nitrogen availability and not necessarily to water availability (february & higgins 2010). according to augustine (2003), lower water-holding capacity and decreased nitrogen in soil can increase the susceptibility of communities on the ridgetop or crest to grazing, and grazing intensity can vary with topography. this also seems to be consistent with larger aggregations of bare soil and annual plant communities at upper topographic positions, compared to lower positions (augustine 2003). all these aspects are important to describe the presence of specific grass plants and their root zones, used in the study to indicate the bacteria associated with these rhizospheres (numbers 12 and 13, figure 2). vernal pools and mud wallows are important components of the savanna ecosystem. a vernal pool is a seasonal, shallow, usually small, ephemeral water body, with no permanent inlet or outlet. their occurrence and frequency of occurrence are directly determined by the precipitation (15, figure 2). the high clay content of the soils (16) in combination with micro-topography and some of the geomorphological aspects determine their location on the catena (17). various mammals also use some of these ephemeral water bodies for drinking and as mud wallows (18). climate (together with geology) is one of the main drivers of long-term vegetation patterns and processes in knp (scholtz et al. 2014). however, climate also affects vegetation response in the short term, as evidenced by the severe 2015–2016 drought experienced during our study period (see study area description) when one of the most severe droughts on record for the area in which the study site falls was experienced (swemmer et al. 2018). drought is not separately indicated, but included indirectly under the label ‘climate’ on figures 1 and 2. during october 2016, barely any herbaceous vegetation was visible on the catena as a result of drought (19, figure 2) and was compounded by associated grazing pressure (i.e. staver, wigley-coetsee & botha 2019). this specific drought affected various plant functional groups (van aardt et al. 2020), including forbs (siebert, klem & van coller 2020), grasses (janecke 2020; wigley-coetsee & staver 2020) and woody vegetation (case et al. 2020; swemmer 2020). the vegetation response during and after the drought was often dependent on past land use (siebert et al. 2020) and/or catenal position (e.g. swemmer 2020). mammals are also influenced by climate (20, figure 2) in terms of temperature (see, e.g. shrestha et al. 2014), humidity and indirectly through rainfall that leads to vegetation growth (food for herbivores) and the presence of natural, surface drinking water. although climate includes many other factors also, only temperature and rainfall have been included in the discussion. many factors on the catena are impacted indirectly by the climate (figure 1 and numbers 19 and 20 on figure 2), by the veld condition and by the intensity and frequency of fire in the ecosystem. the framework discussed in this section (figures 1 and 2) highlighted the interconnectivity of a variety of abiotic and biotic components, and how this may have contributed towards shaping the heterogeneity in patterns and processes along our study catena. we acknowledge that there are numerous factors not mentioned here but that also form part of the processes, heterogeneity and functioning of savanna catena ecosystems, and these could be added in future efforts to refine the framework. synthesis of research findings the focus of this section will be on providing a short, introductory synthesis of the findings published in the articles in this special issue that fits into the framework provided in the previous section. the catena in the study area consists of a low relief crest and upper midslope (slope = 1%), sodic site, shrubveld inside a riparian area (footslope), a small floodplain area and drainage line. the geology, different soil types, some soil properties, the hydrology, vegetation communities, the number of trees and mammals are summarised in figure 3. figure 3: hydropedological response model, topography, soil types and properties, plant communities, vegetation structure (percentages and numbers of individual plants per 100 m transect) and number of mammal species in each zone on the catena. numbers of mammals may seem similar between catenal zones, but do not necessarily fully overlap in species composition (janecke & bolton 2020). the hydropedological response is described in detail in bouwer et al. (2020) and van tol et al. (2020), soil forms (blocks indicate modal profiles where more detailed analyses were done) are described by bouwer et al. (2020), the plant communities are described in theron et al. (2020) and the vegetation structure is described in janecke (2020). hydrological processes hydrological processes included all three catchment orders, with a focus on the third-order catena, whereupon it was shown that lateral interflow along the hillslope at the first-order augment streamflows, whereas the secondand third-order stream reaches are conduits for groundwater recharge to the fractured rock aquifer and thereby show transmission loss from the stream network. recharge soils are typical of the crests and footslopes through preferential vertical flow, whilst the midslope sodic regions are responsive in nature with strong lateral flow contributions at the surface and shallow sub-surface. hillslope soil water balances revealed large variations in the riparian zones resulting from upslope contributions and evapotranspiration losses (riddell et al. 2020). soil distribution four main soil forms were found from the crest to the valley bottom, namely, clovelly, sterkspruit, bonheim and dundee (bouwer et al. 2020). the clovelly soil dominates the crest position; this soil is relatively deep, sandy and freely drained, resulting in low inherent fertility and slightly acidic conditions. below the clovelly soil, a prominent seepline exists (figure 3). this seepline marks the transition from crest to the midslope and the beginning of a sodic site. sterkspruit soils dominate the sodic site with strong duplex character. there is an increase in clay, exchangeable cations (especially sodium [na]) and ph on the sodic site when compared with the crest soils. these soils have a low hydraulic conductivity and will promote overlandflow (van tol et al. 2020). on the lower-midslope, bonheim soils occur. although they also have a relatively strong structure, there is a decline in the dominance of na compared to other cations. in the valley bottom, dundee soils formed from alluvial deposits dominate. these soils are coarse and freely drained with relatively low ph and nutrient status (bouwer et al. 2020). vegetation differences between the four main soil types (chemical and physical properties) led to different vegetation communities and sub-communities that are associated with each soil type (figure 3). smaller differences between soil forms were also observed along the gradient as indicated in the plots where mispah (ms) (the themeda triandra–flueggea virosa–spirostachys africana sub-community in the riparian area occurs exclusively on this soil form [theron et al. 2020]), fernwood (fw), pinedine (pn), estcourt (es) and milkwood (mw) were found (figure 3). furthermore, the themeda triandra–flueggea virosa–ehretia rigida sub-community on the midslope; the dactyloctenium aegyptium–sporobolus nitens–bothriochloa radicans sub-community and the dactyloctenium aegyptium–sporobolus nitens–aristida congesta sub-community of the riparian areas exclusively occur on sterkspruit soils. the other sub-communities occur on a combination of different soil forms (theron et al. 2020). differences in soil properties, moisture content, nutrients and the depth of the hard rock bed can all play a role in the number of trees (excluding other factors such as the impact of animals), especially large trees in different catenal zones. in the study area (figure 3), most of the woody plants had canopy sizes less than 2 m wide, with a low number of big trees (> 5 m) which occurred mostly on the banks of the drainage line and midslope (less than four big trees were found in a 100 m × 3.5 m transect per zone). the canopy cover of woody plants ranged from 4% on the sodic site to 17% on the crest and midslope, and 33% in the riparian area at the drainage line (janecke 2020). the soil type and depth present on the midslope, riparian shrubveld and drainage line banks (figure 3) can support big trees, whilst possible access to ground water in the riparian zone might also explain their presence. just before the drought conditions peaked, the shrubveld inside the riparian area on the catena studied had a higher density of shrubs and of grass plants than the other vegetation zones (janecke 2020, theron et al. 2020). it might be that the shrubs created protected areas that led to increased grass densities, or that through the process of hydraulic lift more water was available closer to the soil surface, which enabled these shrubs and grasses to flourish. it may also be the fact that because of the denser woody conditions and associated reduction in visibility in the shrubveld, herbivores are avoiding those areas to reduce their predation risk (landscape of fear) and in this way grazing pressure was also reduced. furthermore, the soil present in this zone is loamier and could, therefore, also hold more water. bottomlands also have higher clay content, which is more nutrient rich and gives rise to higher productivity. these could be possible explanations for the presence of shrubs and a different herbaceous community than the rest of the catenal zones in the study area. soil and plant associated microbes on a microscale the different ecotypes and soil conditions influence microbial communities in the rhizospheres of associated plants. although the same plants may occur across vegetation zones, the microbes associated with the rhizospheres differ because of chemical differences (vermeulen et al. 2020). bacterial and fungal diversity of the rhizospheres of three species of herbaceous plants (kyphocarpa angustifolia, melhania acuminata and sida cordifolia) growing mutually in two different soil types (clovelly and sterkspruit – figure 3) of the catenal supersite was studied (gryzenhout et al. 2020, vermeulen et al. 2020). carbon source utilisation patterns of bacteria in the rhizospheres of these plants were assessed in addition to illumina sequencing of the 16s rrna (v3 and v4 region), which was used to characterise the rhizobiome (vermeulen et al. 2020). principal component analysis (pca) of biolog data revealed no disparity between the five rhizobiomes based on carbon source utilisation of bacteria. actinobacteria and proteobacteria were the most dominant phyla in all rhizobiomes. unique and shared operational taxonomic units were also identified in all the rhizobiomes (vermeulen et al. 2020). soil type and plant species play an important role in shaping the rhizosphere as unique operational taxonomic units were observed in all rhizospheres. for the fungi, differences between the two soil type rhizosphere communities were found, as well as for those within the roots (gryzenhout et al. 2020). mammals the vegetation creates habitat and provides food for herbivores. available space for larger herds of herbivores and for larger sized mammals (mega-herbivores), as well as competition for food resources, predators, shelter and other factors, could influence animal densities in certain areas that can again affect their impact on vegetation and soil (figure 1) (janecke 2020). the total number of mammal species observed through camera trapping (herbivores + carnivores + other; figure 3) in each catenal zone was as follows: crest and upper midslope (19 species), sodic patch (17 species), shrubveld (14 species), riparian area (15 species) and at the mud wallows (13 species). there were no obvious differences between mammal species observed in the different catenal zones. common mammal species present on the catena, in order of frequency of observations, included impala, elephant, spotted hyena, kudu, giraffe, buffalo, zebra, warthog and grey duiker (janecke and bolton 2020). a total of 31 mammal species were observed on the catena over three camera trapping periods. group sizes varied from a single individual to less than 10 individuals for most species, whilst for elephant it went up to 21 and for impala up to 90 individuals in a herd at certain times (janecke & bolton 2020). the drought indirectly affected the presence or absence of mammal species and impacted on the numbers of the group or herd. mud wallows and ephemeral pans in the study area, the temporary water-filled depressions (mud wallows and vernal pools) were found on the midslope-sodic ecotone (close to the seepline) and next to the drainage line (figure 3). this position is interpreted as being related to the wetter part of the hillslope. these vernal pools are important habitats not only for small aquatic organisms, such as euphyllopod crustaceans (that formed part of this study), but also for other organisms. the drought is probably to blame for lack of crustaceans in these depressions at the time of study. various mammals used some of these temporal water bodies as mud wallows for thermal regulation, but it became especially heavily utilised during the intense heat of the extreme drought. the soils of the sodic site and riparian area have a higher clay content (figure 3) with lower hydraulic conductivity, thus increasing its potential to retain water longer in holes created by animals than the more sandy soils of the midslope and crest. these depressions probably form because of the fact that water collects and remains for longer periods in these clayey soils, which then attracts the mammals to start wallowing in it. according to janecke and bolton (2020), these holes ‘might fill up with rain water, surface run-off and there is a possibility that water seepage from underground may also play a role (depending on the position of the mud hole on the catena)’ and based on the hydrology of the catena (figure 3). severe drought of 2015–2016 as is common during drought periods, isolated small-scale rainfall events occurred during the drought and the study area seemingly received more rain during the study period than the surrounding landscapes, resulting in some grass plants retaining above-ground biomass and experiencing limited flushing, with trees retaining green leaves for longer. anecdotal evidence during the study period suggested occurrence of some precipitation on the watershed between the sabie and crocodile river, where the supersite is situated, which is likely to have contributed to the seeming ‘greenness’ experienced. the veld in the study area appeared like a green island in the surrounding drought-stricken area during march 2016 and might have acted as an important critical resource area to sustain herbivores. the severity of the drought eventually took its toll on the vegetation and during october 2016, almost no herbaceous vegetation was observed, only bare soil (pers. obs. october 2016.). the presence of temporary surface water in holes created by large mammals was used for drinking and as mud wallows during 2015 and 2016 drought period (janecke & bolton 2020) because it retained the rainwater for longer periods. this also contributed to sustaining mammals after water became scarce and is one of the reasons why certain mammal species were present in the study area. plant community and species composition changed in the crest and sodic site zones during the different sampling seasons as the drought progressed. these changes could not be seen in the richness and diversity of plants in these mentioned zones. changes in the riparian areas were, however, not that clear and can be ascribed to the possible hydraulic lift from deeper soil layers, decreasing the impact of the drought (van aardt et al. 2020). the visible changes in the community compositions over the 2015–2018 period could possibly indicate that the granite catena is a relatively resilient ecosystem in terms of abiotic factors affecting the biotic factors. future research this study provides baseline data and biotic–abiotic associations that can be built upon and the hypotheses presented by our conceptual framework can be further tested in future. future research can inter alia focus on various other aspects, such as (1) comparing the geohydrology of a true wet, flowing catena to the dry and semi-dry catena that we experienced because of drought conditions; (2) determining the presence of animals over different seasons and over extended periods, including years with different rainfall regimes; (3) comparing the ecology, diversity and processes of other hillslopes in the supersite with our results, or (4) with other supersites which have different rainfall regimes and underlying geology; (5) including other animals (reptiles, birds, insects, invertebrates and other non-mammals) in the study and understanding how they may respond to catenal heterogeneity. lidar and/or terrestrial scanning of the study area is also a useful data set to acquire for the study area in future. according to naiman et al. (2003), it must be kept in mind that the environment system is complex and full of non-linearity, amplifications, thresholds, damping effects and more that make prediction difficult. however, the formulation of the general principles is not impossible; it just poses a major challenge that ecologists must accomplish. conclusion biodiversity and heterogeneity associated with the catena ecosystem processes are important components in understanding the dynamics of the savanna ecosystem as a whole. most research projects focus on only one or a few of the biotic and abiotic components, and possibly one or two connections between these components, but studies considering a multitude of biotic and abiotic components and their interaction, feedback and integration are rare. this research aimed to contribute towards the knowledge gap by studying the biotic and abiotic components during the same period and in the same local area, to make the interactions, relationships and processes of the ecosystem more explicit and, hence, providing a systemic conceptual understanding of the patterns and connections within the study system. to better understand the complexity of these ecosystems, there will always be opportunity for research to gain more knowledge and a better understanding of these unique ecosystems. furthermore, we trust that this article, and all the studies forming part of the special issue, will contribute valuable baseline information and provide a useful first stab at a conceptual understanding of the patterns, processes and interactions on the knp southern granite supersite. we envisage that this will stimulate further research on the supersites to become pivotal research sites for multi-disciplinary studies of savanna ecosystems. acknowledgements the authors would like to acknowledge the university of the free state (ufs) strategic research fund for funding the multidisciplinary research; sanparks scientific services and game guards for their friendly assistance; dr marcele vermeulen from microbial, biochemical, and food biotechnology (ufs) for her contributions to the article; and the late dr fred kruger (ots & ufs) who was very enthusiastic and excited about this multidisciplinary project but passed away in 2017 before the publication of the special issue. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions all authors directly participated in the study design and interpretation of the multidisciplinary research. the first three authors wrote the main part of the article, whilst b.b.j. and j.v.t. also created the figures with some input from the entire team. the other authors contributed mostly to their expert research field in the article. p.a.l.l.r. is acknowledged for the vision and concept of this research project. m.t.s. and a.c.v.a assisted with language editing. ethical considerations ethical approval for the multidisciplinary project as a whole was obtained from the interfaculty animal ethics committee at the university of the free state (ufs-aed2019/0121). declaration: for project number lropal1255 (lropal1345 extension addendum) as on the contract between the principal investigator and sanparks scientific services, the following permits were issued to collect and remove soil samples for laboratory analyses, vegetation samples for identification purposes, mud samples to identify small aquatic organisms and soil samples from rhizospheres for microbe analyses: sk069; sk2095; sk054. funding information the authors are grateful to the university of the free state (ufs) strategic research fund for largely funding this multidisciplinary research. data availability data sharing is not applicable to this article as no new data were created or analysed during this study. disclaimer the views and opinions expressed in this article are the authors’ own and not an official position of the institution or funder. references adler, p.b., raff, d.a. & lauenroth, w.k., 2001, ‘the effect of grazing on the spatial heterogeneity of vegetation’, oecologia 128, 465–479. https://doi.org/10.1007/s004420100737 asner, g.p. & levick, s.r., 2012, ‘landscape-scale effects of herbivores on treefall in african savannas’, ecology letters 15(11), 1211–1217. https://doi.org/10.1111/j.1461-0248.2012.01842.x asner, g.p., vaughn, n., smit, i.p.j. & levick, s., 2016, ‘ecosystem-scale effects of megafauna in african savannas’, ecography 39(2), 240–252. https://doi.org/10.1111/ecog.01640 augustine, d.j., 2003, ‘spatial heterogeneity in the herbaceous layer of a semi-arid savanna ecosystem’, plant ecology 167, 319–332. https://doi.org/10.1023/a:1023927512590 bouwer, d., le roux, p.a.l. & van tol, j., 2020, ‘identification of hydropedological flowpaths in stevenson-hamilton catena from soil morphological, chemical and hydraulic properties’, koedoe 62(2), a1584. https://doi.org/10.4102/koedoe.v62i2.1584 brady, n.c. & weil, r.r., 2002, the nature and properties of soils, 13th edn., prentice hall, nj. burkepile, d.e., burns, c.e., tambling, c.j., amendola, e., buis, g.m., govender, n. et al., 2013, ‘habitat selection by large herbivores in a southern african savanna: the relative roles of bottom-up and top-down forces’, ecosphere 4(11), 1–19. https://doi.org/10.1890/es13-00078.1 case, m.f., wigley, b.j., wigley-coetsee, c. & carla staver, a., 2020, ‘could drought constrain woody encroachers in savannas?’ african journal of range & forage science 37(1), 19–29. https://doi.org/10.2989/10220119.2019.1697363 classen, a.t., sundqvist, m.k., henning, j.a., newman, g.s., moore, j.a.m., cregger, m.a. et al., 2015, ‘direct and indirect effects of climate change on soil microbial and soil microbial-plant interactions: what lies ahead?’, ecosphere 6(8), 1–21. https://doi.org/10.1890/es15-00217.1 du toit, j.t., rogers, k.h. & biggs, h.c. (eds.), 2003, the kruger experience: ecology and management of savanna heterogeneity, island press, washington, dc. eckhardt, h.c., van wilgen, b.w. & biggs, h.c., 2000, ‘trends in woody vegetation cover in the kruger national park, south africa, between 1940 and 1988’, african journal of ecology 38(2), 108–115. https://doi.org/10.1046/j.1365-2028.2000.00217.x february, e.c. & higgins, s.i., 2010, ‘the distribution of tree and grass roots in savannas in relation to soil nitrogen and water’, south african journal of botany 76(3), 517–523. https://doi.org/10.1016/j.sajb.2010.04.001 grant, c.c., botha, j., grant, t.c., peel, m.j.s. & smit, i.p.j., 2019. ‘when less is more: heterogeneity in grass patch height supports herbivores in counter-intuitive ways’, african journal of range & forage science 36(1), 1–8. https://doi.org/10.2989/10220119.2018.1543209 gryzenhout, m., cason, e.d., vermeulen, m., kloppers, g.a.e., bailey, b. & ghosh, s., 2020, ‘fungal community structure variability between the root rhizosphere and endosphere in a granite catena system in the kruger national park, south africa’, koedoe 62(2), a1597. https://doi.org/10.4102/koedoe.v62i2.1597 hempson, g.p., archibald, s., bond, w.j., ellis, r.p., grant, c.c., kruger, f.j. et al., 2015, ‘ecology of grazing lawns in africa’, biological reviews 90(3), 979–994. https://doi.org/10.1111/brv.12145 hu, s. & fedorov, a.v., 2017, ‘the extreme el niño of 2015–2016: the role of westerly and easterly wind bursts, and preconditioning by the failed 2014 event’, climate dynamics 48, 1–19. https://doi.org/10.1007/s00382-017-3531-2 janecke, b.b., 2020, ‘vegetation structure and spatial heterogeneity in the granite supersite, kruger national park’, koedoe 62(2), a1591. https://doi.org/10.4102/koedoe.v62i2.1591 janecke, b.b. & bolton, j., 2020, ‘variation in mammal diversity and habitat affect heterogeneity and processes of a granite catena’, koedoe 62(2), a1592. https://doi.org/10.4102/koedoe.v62i2.1592 khomo, l.m., hartshorn, a.s., rogers, k.h. & chadwick, o.a., 2011, ‘impact of rainfall and topography on the distribution of clays and major cations in granitic catenas of southern africa’, catena 87(1), 119–128. https://doi.org/10.1016/j.catena.2011.05.017 khomo, l.m. & rogers, k.h., 2005, ‘proposed mechanism for the origin of sodic patches in kruger national park, south africa’, african journal of ecology 43(1), 29–34. https://doi.org/10.1111/j.1365-2028.2004.00532.x laundre, j.w., hernandez, l. & ripple, w.j., 2010, ‘the landscape of fear: ecological implications of being afraid’, the open ecology journal 3(3), 1–7. https://doi.org/10.2174/1874213001003030001 le roux, e., kerley, g.i.j. & cromsigt, j.p.g.m., 2018, ‘megaherbivores modify trophic cascades triggered by fear of predation in an african savanna ecosystem’, current biology 28(15), 2493–2499. https://doi.org/10.1016/j.cub.2018.05.088 malherbe, j., smit, i.p.j., wessels, k.j. & beukes, p.j., 2020, ‘recent droughts in the kruger national park as reflected in the extreme climate index’, african journal of range & forage science 37(1), 1–17. https://doi.org/10.2989/10220119.2020.1718755 mcnaughton, s.j. 1983, ‘compensatory plant growth as a response to herbivory’, oikos 40(3), 329–336. https://doi.org/10.2307/3544305 milewski, a.v. & madden, d. 2006, ‘interactions between large african browsers and thorny acacia on a wildlife ranch in kenya’, african journal of ecology 44(4), 515–522. https://doi.org/10.1111/j.1365-2028.2006.00665.x milne, g., 1936, ‘a provisional soil map of east africa’, geograf review 26(3), 522–523. https://doi.org/10.2307/209064 naiman, r.j., braack, l., grant, r., kemp, a.c., du toit, j.t. & venter, f., 2003, ‘interactions between species and ecosystem characteristics’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 221–241, island press, washington, dc. nasar, j.l. & jones, k.m., 1997, ‘landscapes of fear and stress’, environment and behavior 29(3), 291–323. https://doi.org/10.1177/001391659702900301 pickett, s.t.a., cadenasso, m.l. & benning, t.l., 2003, ‘biotic and abiotic variability as key determinants of savanna heterogeneity at multiple spatiotemporal scales’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 22–40, island press, washington, dc. riddell, e.s., nel, j., fundisi, d., jumbi, f., van niekerk, a. & lorentz, s., 2014, ephemeral hydrological processes in savannas, wrc report no. tt619/14, water research commission, pretoria. riddell, e.s., nel, j.m., van tol, j., fundisi, d., jumbi, f., van niekerk, a. et al., 2020, ‘groundwater–surface water interactions in an ephemeral savanna catchment, kruger national park’, koedoe 62(2), a1583. https://doi.org/10.4102/koedoe.v62i2.1583 rietkerk, m., ketner, p., burger, j., hoorens, b. & olff, h., 2000, ‘multiscale soil and vegetation patchiness along a gradient of herbivore impact in a semi-arid grazing system in west africa’, plant ecology 148, 207–224. https://doi.org/10.1023/a:1009828432690 rietkerk, m., van den bosch, f. & van de koppel, j., 1997, ‘sitespecific properties and irreversible vegetation changes in semiarid grazing systems’, oikos 80(2), 241–252. https://doi.org/10.2307/3546592 rogers, k.h., 2003, ‘adopting a heterogeneity paradigm: implications for management of protected savannas’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience – ecology and management of savanna heterogeneity, pp. 41–58, island press, washington, dc. sankaran, m., hanan, n.p., scholes, r.j., ratnam, j., augustine, d.j., cade, b.s. et al., 2005, ‘determinants of woody cover in african savannas’, nature 438(7069), 846. https://doi.org/10.1038/nature04070 scholtz, r., kiker, g.a., smit, i.p.j. & venter, f.j., 2014, ‘identifying drivers that influence the spatial distribution of woody vegetation in kruger national park, south africa’, ecosphere 5(6), 1–12. https://doi.org/10.1890/es14-00034.1 siebert, f. & eckhardt, h.c., 2008, ‘the vegetation and floristics of the nkuhlu exclosures, kruger national park’, koedoe 50(1), 126–144. https://doi.org/10.4102/koedoe.v50i1.138 siebert, f., klem, j. & van coller, h., 2020, ‘forb community responses to an extensive drought in two contrasting land-use types of a semi-arid lowveld savanna’, african journal of range & forage science 37(1), 53–64. https://doi.org/10.2989/10220119.2020.1726464 shrestha, a.k., van wieren, s.e., van langevelde, f., fuller, a., hetem, r.s., meyer, l. et al., 2014, ‘larger antelopes are sensitive to heat stress throughout all seasons but smaller antelopes only during summer in an african semi-arid environment’, international journal of biometeorology 58(1), 41–49. https://doi.org/10.1007/s00484-012-0622-y smit, i.p.j., 2020, ‘integrating multi-scaled and multi-disciplinary studies: a critical reflection on the kruger national park research supersites’, koedoe 62(2), a1586. https://doi.org/10.4102/koedoe.v62i2.1586 smit, i.p.j. & bond, w.j., 2020, ‘observations on the natural history of a savanna drought’, african journal of range & forage science 37(1), 119–136. https://doi.org/10.2989/10220119.2020.1723695 smit, i.p.j. & prins, h.h.t., 2015, ‘predicting the effects of woody encroachment on mammal communities, grazing biomass and fire frequency in african savannas’, plos one 10(9), e0137857. https://doi.org/10.1371/journal.pone.0137857 smit, i.p.j., riddell, e.s., cullum, c. & petersen, r., 2013, ‘kruger national park research supersites: establishing long-term research sites for cross-disciplinary, multiscaled learning’, koedoe 55(1), art#1107, 7 pages. https://doi.org/10.4102/koedoe.v55i1.1107 staver, a.c., wigley-coetsee, c. & botha, j., 2019, ‘grazer movements exacerbate grass declines during drought in an african savanna’, journal of ecology 107(3), 1482–1491. https://doi.org/10.1111/1365-2745.13106 swemmer, a.m., 2020, ‘locally high, but regionally low: the impact of the 2014–2016 drought on the trees of semi-arid savannas, south africa’, african journal of range & forage science 37(1), 31–42. https://doi.org/10.2989/10220119.2020.1723696 swemmer, a.m., bond, w.j., donaldson, j., hempson, g.p., malherbe, j. & smit, i.p.j., 2018, ‘the ecology of drought a workshop report’, south african journal of science 114(9–10), 1–3. https://doi.org/10.17159/sajs.2018/5098 theron, e., van aardt, a.c. & du preez, p.j., 2020, ‘vegetation distribution along a granite catena, southern kruger national park, south africa’, koedoe 62(2), a1588. https://doi.org/10.4102/koedoe.v62i2.1588 treydte, a.c., van der beek, j.g.m., perdok, a.a. & van wieren, s.e., 2011, ‘grazing ungulates select for grasses growing beneath trees in african savannas’, mammalian biology 76(3), 345–350. https://doi.org/10.1016/j.mambio.2010.09.003 trollope, w.s.w., trollope, l.a., biggs, h.c., pienaar, d. & potgieter, a.l.f., 1988, ‘long-term changes in the woody vegetation of the kruger national park with special reference to the effects of elephants and fire’, koedoe 41(2), 103–112. https://doi.org/10.4102/koedoe.v41i2.255 van aardt, a.c., codron, d., theron, e. & du preez, p.j., 2020, ‘plant community structure and possible vegetation changes after drought on a granite catena in the kruger national park, south africa’, koedoe 62(2), a1585. https://doi.org/10.4102/koedoe.v62i2.1585 van tol, j., julich, s., bouwer, d. & riddell, e.s., 2020, ‘hydrological response in a savanna hillslope under different rainfall regimes’, koedoe 62(2), a1602. https://doi.org/10.4102/koedoe.v62i2.1602 vermeulen, m., cason, e.d. & swart, w.j., 2020, ‘the rhizobiome of herbaceous plants in clovelly and sterkspruit soils of the stevenson-hamilton supersite’, koedoe 62(2), a1596. https://doi.org/10.4102/koedoe.v62i2.1596 whalen, j.k & sampedro, l, 2010, soil ecology and management, cambridge university press, cambridge. whyte, i.j., van aarde, r.j. & pimm, s.l., 2003, ‘kruger’s elephant population: its size and consequences for ecosystem heterogeneity’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience – ecology and management of savanna heterogeneity, pp. 332–348, island press, washington, dc. wigley-coetsee, c. & staver, a.c., 2020, ‘grass community responses to drought in an african savanna’, african journal of range & forage science 37(1), 43–52. https://doi.org/10.2989/10220119.2020.1716072 koedoe 62-1_2020_contents.indd http://www.koedoe.co.za open access table of contents short communication probing hidden diversity to enhance conservation of the endangered narrow-range endemic eastern cape rocky, sandelia bainsii (castelnau 1861) albert chakona, gavin gouws, wilbert t. kadye, pule p. mpopetsi, paul h. skelton koedoe | vol 62, no 1 | a1627 | 29 september 2020 review article using palaeoecology to explore the resilience of southern african savannas lindsey gillson, anneli ekblom koedoe | vol 62, no 1 | a1576 | 15 june 2020 essay no-take estuarine-protected areas: the missing armour for the conservation of fishes alan k. whitfield, colin g. attwood, paul d. cowley, stephen j. lamberth, bruce q. mann koedoe | vol 62, no 1 | a1648 | 05 november 2020 checklist a list of spider species found in the addo elephant national park, eastern cape province, south africa anna s. dippenaar-schoeman, linda wiese, stefan h. foord, charles r. haddad koedoe | vol 62, no 1 | a1578 | 02 april 2020 original research sampling bias in reptile occurrence data for the kruger national park jody m. barends, darren w. pietersen, guinevere zambatis, donovan r.c. tye, bryan maritz koedoe | vol 62, no 1 | a1579 | 11 may 2020 original research climate change and the ownership of game: a concern for fenced wildlife areas andy blackmore koedoe | vol 62, no 1 | a1594 | 22 june 2020 original research community-based ecotourism project in communities adjacent to the addo elephant national park: will households pay for it? babatope e. akinyemi, abbyssinia mushunje koedoe | vol 62, no 1 | a1571 | 22 july 2020 original research interpretation needs and preferences of visitors to kgalagadi transfrontier park petrus van der merwe, melville saayman, elricke botha koedoe | vol 62, no 1 | a1613 | 11 august 2020 40 46 58 65 78 87 92 102 page i of iii table of contents editorial koedoe: changing of the guard llewellyn c. foxcroft koedoe | vol 62, no 1 | a1645 | 30 october 2020 book review biography of marjorie courtenay-latimer – book review peter a. novellie koedoe | vol 62, no 1 | a1607 | 20 january 2020 book review eden lost, and regained – book review peter a. novellie koedoe | vol 62, no 1 | a1628 | 30 april 2020 book review review: conservation biology in sub-saharan africa marna herbst koedoe | vol 62, no 1 | a1635 | 30 june 2020 short communication reconnaissance of epigeal ants at the degraded and control sites of mountain zebra and mokala national parks hendrik sithole, nolubabalo tantsi, hugo bezuidenhout, thinandavha c. munyai, lufuno munyai koedoe | vol 62, no 1 | a1542 | 25 may 2020 short communication assessment results (2015–2018) of re-established poached cycad trees in addo elephant national park, eastern cape, south africa hugo bezuidenhout koedoe | vol 62, no 1 | a1626 | 14 july 2020 short communication woody cover mapping in the savanna ecosystem of the kruger national park using sentinel-1 c-band time series data marcel urban, kai heckel, christian berger, patrick schratz, izak p.j. smit, tercia strydom, jussi baade, christiane schmullius koedoe | vol 62, no 1 | a1621 | 04 august 2020 short communication assessing water conditions for heleophryne rosei tadpoles and the conservation relevance zishan ebrahim, atherton de villiers, john measey koedoe | vol 62, no 1 | a1581 | 29 june 2020 short communication morphometric regressions for the endangered knysna seahorse, hippocampus capensis, in the swartvlei estuary from mass stranding events clement j. arendse, ian a. russell koedoe | vol 62, no 1 | a1632 | 28 september 2020 1 4 6 8 10 16 22 28 34 vol 62, no 1 (2020) issn: 0075-6458 (print) | issn: 2071-0771 (online)koedoe http://www.koedoe.co.za open access table of contents original research a preliminary assessment of the presence and distribution of invasive and potentially invasive alien plant species in laikipia county, kenya, a biodiversity hotspot arne b.r. witt, winnie nunda, tim beale, darren j. kriticos koedoe | vol 62, no 1 | a1605 | 09 september 2020 original research evaluating environmental impact assessment report quality in south african national parks luke a. sandham, carla huysamen, francois p. retief, angus morrisonsaunders, alan j. bond, jenny pope, reece c. alberts koedoe | vol 62, no 1 | a1631 | 16 september 2020 original research effects of fire frequency on savanna butterfly diversity and composition: a preliminary study elie gaget, catherine l. parr, clélia sirami koedoe | vol 62, no 1 | a1617 | 21 september 2020 original research plant community diversity in the chobe enclave, botswana: insights for functional habitat heterogeneity for herbivores pascal vittoz, federico pellacani, rémy romanens, ali mainga, eric p. verrecchia, richard w.s. fynn koedoe | vol 62, no 1 | a1604 | 21 october 2020 original research differential herbivore occupancy of fire-manipulated savannas in the satara region of the kruger national park, south africa thobile b. dlamini, brian k. reilly, dave i. thompson, deron e. burkepile, judith m. botha, anthony g. rebelo koedoe | vol 62, no 1 | a1603 | 26 october 2020 113 123 132 141 158 vol 62, no 2 (2020) special collection: granite catena ecosystem, sub-edited by beanelri janecke and johan van tol editorial connections between abiotic and biotic components of a granite catena ecosystem in kruger national park beanelri b. janecke, johan van tol koedoe | vol 62, no 2 | a1638 | 29 october 2020 tribute a tribute to frederick (fred) j. kruger wijnand j. swart, maitland t. seaman, pieter a.l. le roux, beanelri b. janecke koedoe | vol 62, no 2 | a1639 | 29 october 2020 tribute a tribute to pieter johannes (johann) du preez leslie r. brown, andri c. van aardt, beanelri b. janecke koedoe | vol 62, no 2 | a1640 | 29 october 2020 short communication first report of various fusarium species from the stevenson-hamilton supersite granite catena system in the kruger national park, south africa marieka gryzenhout, marcele vermeulen, gilmore pambuka, riana jacobs koedoe | vol 62, no 2 | a1599 | 29 october 2020 204 206 207 208 page ii of iii original research potential impacts of changing climate on nature-based tourism: a case study of south africa’s national parks kevin m. coldrey, jane k. turpie koedoe | vol 62, no 1 | a1629 | 29 october 2020 original research analysis of three decades of land cover changes in the maputaland coastal plain, south africa manish ramjeawon, molla demlie, michele l. toucher, susan janse van rensburg koedoe | vol 62, no 1 | a1642 | 09 november 2020 original research assessing the effect of tagging and the vulnerability to predation in tigerfish (hydrocynus vittatus, castelnau 1861) in a water-stressed system using telemetry methods bayanda sonamzi, matthew burnett, robin petersen, gordon o’brien, colleen t. downs koedoe | vol 62, no 1 | a1649 | 17 december 2020 correction erratum: climate change and the ownership of game: a concern for fenced wildlife areas andy blackmore koedoe | vol 62, no 1 | a1659 | 11 december 2020 reviewer acknowledgement koedoe | vol 62, no 1 | a1662 | 22 december 2020 166 178 190 202 203 essay integrating multi-scaled and multidisciplinary studies: a critical reflection on the kruger national park research supersites izak p.j. smit koedoe | vol 62, no 2 | a1586 | 29 october 2020 original research biotic and abiotic connections on a granitic catena: framework for multidisciplinary research beanelri b. janecke, johan van tol, izak p.j. smit, andri c. van aardt, edward s. riddell, maitland t. seaman, wijnand j. swart, pieter j. du preez, pieter a.l. le roux koedoe | vol 62, no 2 | a1600 | 29 october 2020 original research identification of hydropedological flowpaths in stevenson–hamilton catena from soil morphological, chemical and hydraulic properties darren bouwer, pieter a.l. le roux, johan van tol koedoe | vol 62, no 2 | a1584 | 29 october 2020 216 221 232 http://www.koedoe.co.za open access table of contents original research groundwater–surface water interactions in an ephemeral savanna catchment, kruger national park edward s. riddell, jaco nel, johan van tol, daniel fundisi, faith jumbi, ashton van niekerk, simon lorentz koedoe | vol 62, no 2 | a1583 | 29 october 2020 original research hydrological response in a savanna hillslope under different rainfall regimes johan van tol, stefan julich, darren bouwer, edward s. riddell koedoe | vol 62, no 2 | a1602 | 29 october 2020 original research the rhizobiome of herbaceous plants in clovelly and sterkspruit soils of the stevenson–hamilton supersite marcele vermeulen, errol d. cason, wijnand j. swart koedoe | vol 62, no 2 | a1596 | 29 october 2020 original research fungal community structure variability between the root rhizosphere and endosphere in a granite catena system in kruger national park, south africa marieka gryzenhout, errol d. cason, marcele vermeulen, george a.e. kloppers, brooke bailey, soumya ghosh koedoe | vol 62, no 2 | a1597 | 29 october 2020 243 257 267 276 page iii of iii original research vegetation distribution along a granite catena, southern kruger national park, south africa ettienne j. theron, andri c. van aardt, pieter j. du preez koedoe | vol 62, no 2 | a1588 | 29 october 2020 original research vegetation structure and spatial heterogeneity in the granite supersite, kruger national park beanelri b. janecke koedoe | vol 62, no 2 | a1591 | 29 october 2020 original research plant community structure and possible vegetation changes after drought on a granite catena in the kruger national park, south africa andri c. van aardt, daryl codron, ettienne j. theron, pieter j. du preez koedoe | vol 62, no 2 | a1585 | 29 october 2020 original research variation in mammal diversity and habitat affect heterogeneity and processes of a granite catena beanelri b. janecke, jeremy g. bolton koedoe | vol 62, no 2 | a1592 | 29 october 2020 287 298 310 321 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research vol. 50 no. 1 pp. 99 112 landscapes in the kalahari gemsbok national park, south africa abstract a landscape map of the kalahari gemsbok national park is presented. mapping is at a finer scale than previous vegetation and habitat maps for the same area. the landscapes were grouped into seven large classes and a total of 20 landscapes were mapped. a description of the terrain morphology, soil and vegetation of each landscape is provided. landscapes that are focal points for the large animals of the region include the calcrete outcrops, riverbeds and pans. these landscapes cover only about 10% of the total area of the region. this map can be used as basis for park planning, management, research and other applications. keywords: arid savanna, dune vegetation, kgalagadi, shrubby grassland, vegetation map margaretha w. van rooyen noel van rooyen department of plant science university of pretoria south africa jacobus du p. bothma centre for wildlife management university of pretoria south africa hendrik m. van den berg iris international south africa correspondence to: margaretha w. van rooyen e-mail: gretel.vanrooyen@up.ac.za postal address: department of plant science, university of pretoria, pretoria, 0002, south africa 99 the classification, description and mapping of plant communities and their associated abiotic features are vital first steps in building a framework to understand, protect, conserve and manage our natural resources. mapping of physiographicphysiognomic relatively homogeneous spatial landscape units at an appropriate scale is therefore a basic requirement for objective resource assessment of any study area. such a landscape map can amongst others be used to determine the geographical distribution and extent of different habitats; identify areas that are ecologically sensitive (disturbed or high in diversity); assess diversity and the status of rare and endemic species; provide a baseline for assessing range condition and grazing and browsing capacities; interpret animal movements and distribution; and position roads, lodges, hiking and offroad trails. the vegetation of the southern kalahari as defined by thomas and shaw (1991), or parts thereof, has been described by pole-evans (1936), acocks (1953), story (1958), brynard (1958), leistner (1959a, 1959b, 1967), giess (1971), mostert et al. (1971), bothma and de graaff (1973), leistner and werger (1973), coetzee and werger (1975), werger and coetzee (1978), gubb (1980), werger (1986), van rooyen et al. (1988), van rooyen (1996) and van rooyen and van rooyen (1998). interestingly, the vegetation of the kalahari gemsbok national park (kgnp) was not included in the vegetation map of south africa compiled by acocks in 1953. in the refined mapping of the vegetation of south africa (low & rebelo 1998), seven vegetation types were distinguished and mapped for the southern kalahari savanna in south africa of which two occurred within the boundaries of the park, i.e. the thorny kalahari dune bushveld and the shrubby kalahari dune bushveld. in the latest vegetation map of south africa (mucina et al. 2005, mucina & rutherford 2006), seven vegetation types were identified in the kgnp. these seven types are the nossob bushveld, gordonia kameeldoring bushveld, gordonia duneveld, auob duneveld, southern kalahari mekgacha, gordonia plains shrubland and southern kalahari salt pans. three phytosociological studies have been conducted in the kgnp. the first classification was done by leistner and werger (1973) and 12 plant communities were distinguished and described but not mapped. they concluded that the pattern of the plant communities was determined by soil characteristics, such as soil-water regime and nutrient availability (werger 1986). in 1999, lubbinge (1999) did a more detailed study and distinguished 24 plant communities in the kgnp, which he combined into 14 mapping units. van rooyen (2000) expanded lubbinge’s floristic analysis to include the entire kgalagadi transfrontier park. in the resultant vegetation map nine vegetation units were confined to the botswana side of the kgalagadi transfrontier park and 14 units were mapped for the south african side (kgnp) (van rooyen 2000). bothma and de graaff (1973) produced the first habitat map of the kgnp. six major habitat categories were distinguished, viz. the riverbeds of the auob and nossob rivers; pans; tree savanna; dunes with treeor shrub-covered crests; dunes superficially devoid of trees and shrubs; and flat, open grassy plains. the main objective of the current project was to produce a more detailed map of the kgnp that could be used for park planning and management. such a map can be used to aid in the selection of sites for vegetation monitoring and serve as a basis to determine animal-habitat relationships. it can also be related to many other ecological spatial data sets, and by so doing it will help to identify knowledge gaps so that efforts to acquire additional data can be prioritised and coordinated. the units identified in the present study were termed landscapes following the definition of gertenbach (1983): “a landscape is 2008 kalahari gemsbok national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 99 112 figure 1 location of the kgalagadi transfrontier park (shaded area) within southern africa 100 original research van rooyen, van rooyen, bothma & van den berg vol. 50 no. 1 pp. 99 112koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za figure 2 landscapes of the kalahari gemsbok national park 101 kalahari gemsbok national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 99 112 figure 2 (cont...) between the units can often only be ascribed to terrain morphology, e.g. dune height and pattern, or soil characteristics, which relate to soil-water and nutrient supply (werger 1986), or changes in the dominance and structure of the plant species. the landscapes and their associated vegetation identified in fig. 2 were grouped into seven large classes and a total of 20 landscapes were mapped (table 1). a key has been provided in the form of a dendrogram (fig. 3) that can be used to classify landscapes in the field. an overview of the main corresponding vegetation units in the latest vegetation map of south africa (mucina & rutherford 2006) is provided in fig. 4. in general, the mapping units of mucina and rutherford are heterogeneous assemblages compared to the units presented here. an electronic arcview-compatible version (raster or vector format) of the landscape map is available from the authors on request. a printable version at a scale of 1:150 000 when printed to media 841 mm by 1 400 mm is also available. the pdf version provided here should best be printed to a3. a. high dunes 1. acacia haematoxylon parallel high duneveld these grass-covered, relatively parallel (linear) high dunes occupy an area of 40 951 ha (3.75% of the mapped area; fig. 2) and occur in the south in the vicinity of twee rivieren. fig. 5a illustrates the parallel (linear) dunes that are up to 20 m in height in this unit. the coarse, red, loose sand of the dune crests is dominated by the tall grass stipagrostis amabilis while the shrub acacia haematoxylon is the most conspicuous and widespread woody species on the grassy dunes and in the valleys. besides stipagrostis amabilis, the other prominent grass species are schmidtia kalahariensis, centropodia glauca, eragrostis trichophora, e. lehmanniana, brachiaria glomerata and stipagrostis uniplumis. the most common dwarf shrubs or forbs include hermannia tomentosa, plinthus sericeus, requienia sphaerosperma, bulbostylis hispidula, limeum arenicolum, oxygonum dregeanum and sesamum triphyllum. the prostrate creepers acanthosicyos naudinianus and citrullus lanatus may be seasonally prominent on exposed dune crests. in isolated patches within this landscape unit, the shrub lebeckia linearifolia is a prominent species found on dunes near pans and rivers. 2. acacia haematoxylon irregular high duneveld this extensive dune landscape (fig. 5b) occurs south of the auob river and between the nossob and auob rivers in the south and occupies an area of 127 532 ha (11.67% of the mapped area). the unit is characterised by scattered acacia haematoxylon trees and shrubs, while the presence of boscia albitrunca and acacia mellifera differentiates this unit from the previous one. the prominent grasses are stipagrostis amabilis, eragrostis trichophora, brachiaria glomerata, stipagrostis uniplumis, s. ciliata, centropodia glauca and schmidtia kalahariensis. rhigozum trichotomum dominates the whitish calcareous soils in the dune valleys and depressions. in these patches harpagophytum procumbens, jatropha erythropoda and dipcadi gracillimum are the diagnostic species. 3. acacia erioloba-acacia mellifera parallel high duneveld this open tree savanna (fig. 5c) occurs widespread on the deep, reddish sand in the duneveld and near rivers and pans, mostly along a strip on the northern side of the auob river and on both sides of the nossob river from samevloeiing in the south to langklaas in the north. the landscape unit also occurs south of grootkolk in the north. it is the third largest unit in the kgnp and covers an area of 175 923 ha (16.09% of the mapped area). the prominent woody species are acacia erioloba, a. haematoxylon, boscia albitrunca and grewia flava. acacia mellifera is particularly prominent against north to northeast-facing slopes. the diagnostic species on the dune slopes are searsia tenuinervis (= rhus tenuinervis), lycium bosciifolium, l. villosum an area with a specific geomorphology, macroclimate, soil and vegetation pattern and associated fauna”. study area the study area comprises the south african portion of the kgalagadi transfrontier park (fig. 1), as well as a narrow zone along the eastern side of the nossob river in botswana. the mapped area covers approximately 10 932 km². the southern kalahari is a summer rainfall region and the mean annual rainfall ranges from 150 mm in the southwest to 450 mm in the northeast in botswana. in the study area the mean annual rainfall ranges from approximately 185 mm to 230 mm. rainfall is highly erratic (skarpe 1986, tyson 1986) and at any specific locality in the park can vary from less than 100 mm up to more than 700 mm per annum. the highest rainfall occurs in the four months january to april, with a peak in march. temperatures show a large amplitude with winter lows reaching -10.3ºc and summer highs reaching 45.4ºc (van rooyen et al. 1990). methods stratification and interpretation of the area was done with landsat etm satellite images (bands 4, 5 and 3 [r,g,b]) 1:5000 orthophoto maps and previous habitat and/or vegetation maps and/or descriptions produced by leistner (1967), bothma and de graaff (1973), leistner and werger (1973), skarpe (1986), bothma and swart (1990), bothma et al. (1993), van rooyen and van rooyen (1998), lubbinge (1999) and van rooyen (2000). in addition to the floristic data of more than 500 phytosociological relevés, qualitative vegetation and habitat surveys were conducted throughout the park based on the stratification of the available imagery. the data collected at each sample site were a global positioning system (gps) coordinate, terrain morphology, soil type, soil texture, soil colour, dominant plant species and vegetation structure. results the southern kalahari environment is relatively homogeneous because of the predominance of the sand mantle covering the area. the differences between the landscape units that were distinguished in this study are therefore often subtle. the same plant species occur in most of the units, and the differences 102 original research van rooyen, van rooyen, bothma & van den berg vol. 50 no. 1 pp. 99 112koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za and ehretia alba. conspicuous dwarf shrubs are asparagus nelsii, gnidia polycephala and rhigozum trichotomum. the most common grass species include schmidtia kalahariensis, eragrostis lehmanniana, centropodia glauca, stipagrostis uniplumis and s. ciliata. common herb species are crotalaria sphaerocarpa, pollichia campestris, bulbostylis hispidula, sesamum triphyllum, limeum fenestratum, l. argute-carinatum and cleome kalachariensis. b. low dunes 4. acacia erioloba-schmidtia kalahariensis low duneveld this open tree savanna occupies 38 993 ha (3.57% of the mapped area) and occurs in the interior duneveld in the north in the vicinity of loffiesdraai, dankbaar, gharagab and bayip and west of the nossob river near grootbrak. the dunes are typically from 6 m to 9 m high in the interior and from 9 m to 15 m high along the rivers. the landscape is relatively species poor and is dominated by acacia erioloba and schmidtia kalahariensis (fig. 6a). other prominent woody species are rhigozum trichotomum, boscia albitrunca and acacia mellifera and occasionally searsia tenuinervis, cadaba aphylla and lycium spp. terminalia sericea is locally prominent on the higher dunes. besides schmidtia kalahariensis, the grass layer is characterised by eragrostis lehmanniana, centropodia glauca, stipagrostis amabilis, s. uniplumis, brachiaria glomerata and stipagrostis ciliata. herbs that are found in this unit include hermannia tomentosa, hermbstaedtia fleckii, requienia sphaerosperma, pupalia lappacea, chrysocoma obtusata, cynanchum orangeanum and the climber pergularia daemia. isolated patches of either acacia luederitzii or albizia anthelmintica occur on dunes and slopes within this unit in the northern parts of the park, especially west of the nossob river in the vicinity of grootbrak and bayip pan (fig. 6b). although many of these patches are too small to be mapped individually, the larger ones have been mapped (unit 4a in fig. 2) and in total cover 532 ha (0.05% of the mapped area). other woody species associated table 1 landscapes of the kalahari gemsbok national park based on the dominant plant species, terrain morphology and soil types landscape unit area (ha) % of total area mapped % per landscape group a. 1. 2. 3. high dunes acacia haematoxylon parallel high duneveld acacia haematoxylon irregular high duneveld acacia erioloba-acacia mellifera parallel high duneveld 40 950.8 127 532.4 175 922.5 3.75 11.67 16.09 31.51 b. 4. 4a 5. 6. low dunes acacia erioloba-schmidtia kalahariensis low duneveld acacia luederitzii or albizia anthelmintica patches acacia haematoxylon-stipagrostis ciliata irregular low duneveld acacia haematoxylon-centropodia glauca undulating low duneveld 38 992.5 532.3 243 154.8 182 888.7 3.57 0.05 22.24 16.73 42.59 c. 7. 8. plains on deep reddish sand acacia haematoxylon-stipagrostis amabilis undulating plains stipagrostis amabilis plains 115 478.9 51 198.5 10.56 4.68 15.24 d. 9 calcretes calcrete outcrops 5 826.1 0.53 0.53 e. 10 11 12 13 plains and terraces along riverbeds and grass pans on compact whitish sand acacia erioloba-rhigozum trichotomum-stipagrostis obtusa river terraces acacia mellifera-rhigozum trichotomum-stipagrostis obtusa river terraces salsola spp.-eriocephalus decussatus river terraces and pan edges stipagrostis obtusa river terraces 38 763.8 34 431.3 1 593.7 2 667.3 3.55 3.15 0.15 0.24 7.09 f. 14 15 16 17 18 19 riverbeds and pans sporobolus rangei pan floors panicum coloratum riverbeds galenia africana-lagerra decurrens riverbeds lebeckia linearifolia riverbeds acacia erioloba-acacia haematoxylon riverbeds grassy pans 3 024.3 6 917.4 6 320.5 910.7 4 350.6 10 781.8 0.26 0.63 0.58 0.08 0.40 0.99 2.94 g. 20 highly irregular dunes highly irregular dunes 917.1 0.08 0.08 total 1 093 155.9 100.00 100.00 with these patches include acacia erioloba, a. mellifera, boscia albitrunca and grewia flava, with schmidtia kalahariensis the dominant grass species. 5. acacia haematoxylon-stipagrostis ciliata irregular low duneveld this is the largest landscape unit in the park, covering an area of 243 155 ha or approximately 22% of the mapped area (fig. 2). it encompasses the grass-covered irregular dunes (fig. 6c) in the southwestern section, east of mata-mata and north of the auob river in the strathmore, nu quap, copeng and lammermoor areas, as well as from the south to the far north in the eastern interior duneveld. a large number of pans are scattered throughout this landscape. acacia haematoxylon is the most conspicuous woody species with stipagrostis ciliata, centropodia glauca and stipagrostis amabilis the prominent grass species. other widespread grass species in this unit include stipagrostis uniplumis, brachiaria glomerata, schmidtia kalahariensis and eragrostis lehmanniana. the forb layer is characterised by requienia sphaerosperma, sesamum triphyllum, hermannia tomentosa, plinthus sericeus, oxygonum dregeanum and limeum arenicolum. the prostrate cucurbits citrullus lanatus and acanthosicyos naudinianus are sometimes prominent. the vegetation on the dune slopes often differs from that in the interdune valleys. the prominent woody species on the dune slopes are acacia mellifera, boscia albitrunca and acacia haematoxylon with centropodia glauca, schmidtia kalahariensis, eragrostis lehmanniana, requienia sphaerosperma and senna italica the conspicuous grass and forb species. the prominent woody species in the grass-covered dune valleys on deep greyish to reddish sand are acacia mellifera, rhigozum trichotomum and acacia haematoxylon. the more common grass and forb species include stipagrostis uniplumis, centropodia glauca, eragrostis lehmanniana, schmidtia kalahariensis, chascanum pumilum, hermannia burchellii, senna italica, limeum argute103 kalahari gemsbok national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 99 112 irregular 2, acacia haematoxylon irregular high dunevelddunes high acacia haematoxylon 1, acacia haematoxylon parallel high dunevelddunes tall shrubland parallel dunes acacia erioloba 3, acacia erioloba-acacia mellifera parallel high duneveldsavanna tree acacia erioloba 4, acacia erioloba-schmidtia kalahariensis low duneveldsavanna savanna dunes and low sandy plains dunes irregular dune 5, acacia haematoxylon-stipagrostis ciliata irregular low duneveld low dunes shrub topography and plains savanna undulating 6, acacia haematoxylon-centropodia glauca undulating low duneveldtopography undulating 7, acacia haematoxylon-stipagrostis amabilis undulating plains plains shrubland flat grassy 8, stipagrostis amabilis plains plains parabolic 20, highly irregular dunes dunes open tree 10, acacia erioloba-rhigozum trichotomumstipagrostis obtusa river terracessavanna river acacia mellifera 11, acacia mellifera-rhigozum trichotomumstipagrostis obtusa river terracesterraces shrubland salsola spp. brackish 12, salsola spp.-eriocephalus decussatus river terraces and pan edgesdwarf shrubland stipagrostis obtusa 13, stipagrostis obtusa river terraces riverbeds grassland & terraces panicum coloratum 15, panicum coloratum riverbeds grassland galenia africana 16, galenia africana-lagerra decurrens riverbeds riverbed shrubland lebeckia linearifolia 17, lebeckia linearifolia riverbeds river & pan shrubland systems riverine 18, acacia erioloba-acacia haematoxylon riverbedssavanna bare pan 14, sporobolus rangei pan floors floor pans grassy 19, grassy pans pans calcrete 9, calcrete outcrops outcrops figure 3 relationships between the landscape units identified in the kalahari gemsbok national park 104 original research van rooyen, van rooyen, bothma & van den berg vol. 50 no. 1 pp. 99 112koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za acacia haematoxylon parallel high duneveld auob duneveld acacia haematoxylon irregular high duneveld acacia erioloba-acacia mellifera parallel high duneveld acacia erioloba-schmidtia kalahariensis low duneveld gordonia kameeldoring bushveld acacia haematoxylon-stipagrostis ciliata irregular low duneveld acacia haematoxylon-centropodia glauca undulating low duneveld acacia haematoxylon-stipagrostis amabilis undulating plains nossob bushveld stipagrostis amabilis plains calcrete outcrops acacia erioloba-rhigozum trichotomum-stipagrostis obtusa river terraces gordonia plains shrubland acacia mellifera-rhigozum trichotomum-stipagrostis obtusa river terraces salsola spp.-eriocephalus decussatus river terraces and pan edges stipagrostis obtusa river terraces southern kalahari mekgacha sporobolus rangei pan floors panicum coloratum riverbeds galenia africana-lagerra decurrens riverbeds gordonia duneveld lebeckia linearifolia riverbeds acacia erioloba-acacia haematoxylon riverbeds grassy pans southern kalahari salt pans highly irregular dunes figure 4 an overview of the correspondence between the present landscapes and the vegetation units in the latest vegetation map of south africa by mucina and rutherford (2006) 105 kalahari gemsbok national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 99 112 carinatum and ipomoea hackeliana. in the dune valleys, near pans and rivers on slightly compact calcareous and whitish sand, open to dense stands of rhigozum trichotomum, monechma incanum and stipagrostis obtusa dominate. aptosimum albomarginatum, talinum caffrum, stipagrostis ciliata and tribulus zeyheri are some of the other prominent species on these calcareous soils. 6. acacia haematoxylon-centropodia glauca undulating low duneveld this undulating shrubby grassland covers the second largest area in the kgnp (182 889 ha or 16.73%) and occurs throughout the interior duneveld from kij gamies in the south to union’s end in the north. the prominent woody species in this undulating landscape are acacia haematoxylon, acacia erioloba and boscia albitrunca, with centropodia glauca, aristida meridionalis, schmidtia kalahariensis, eragrostis lehmanniana and stipagrostis uniplumis the conspicuous grass species (fig. 6d). the characteristic dwarf shrubs and forbs include requienia sphaerosperma, chascanum pumilum, hermannia burchellii, limeum argute-carinatum, ipomoea hackeliana, chascanum garipense and senna italica. a b c figure 5 landscapes of the high dunes: (a) the acacia haematoxylon parallel high duneveld (a1); (b) the acacia haematoxylon irregular high duneveld (a2); and (c) the acacia erioloba-acacia mellifera parallel high duneveld (a3) in a zone along the rivers figure 6 landscapes of the low dunes: (a) the acacia erioloba-schmidtia kalahariensis low duneveld (b4) near loffiesdraai; (b) an albizia anthelmintica patch (b4a) around bayip pan; (c) the acacia haematoxylon-stipagrostis ciliata irregular low duneveld (b5); and (d) the acacia haematoxylon-centropodia glauca undulating low duneveld (b6) c b a d c. plains on deep reddish sand 7. acacia haematoxylon-stipagrostis amabilis undulating plains this shrubby grassland on deep sand of gently undulating plains covers a large area (115 479 ha or 10.56% of the mapped area) and occurs in large patches from south to north throughout the interior duneveld. the dunes are not prominent, and if they occur they seldom reach 6 m in height. acacia haematoxylon is the 106 original research van rooyen, van rooyen, bothma & van den berg vol. 50 no. 1 pp. 99 112koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a b figure 7 landscapes of the plains on deep reddish sand in the interior duneveld: (a) the acacia haematoxylon-stipagrostis amabilis undulating plains (c7); and (b) the flat landscape of the stipagrostis amabilis plains (c8) conspicuous woody species, usually as shrub, with stipagrostis amabilis the most prominent grass species (fig. 7a). other grass species of note are schmidtia kalahariensis, centropodia glauca, eragrostis lehmanniana, stipagrostis uniplumis and brachiaria glomerata. the most common dwarf shrubs and forbs include requienia sphaerosperma, bulbostylis hispidula, acanthosicyos naudinianus, citrullus lanatus, hermannia tomentosa, plinthus sericeus, oxygonum dregeanum, sesamum triphyllum and limeum arenicolum. 8. stipagrostis amabilis plains these flat, grassy plains (fig. 7b) with scattered shrubs of acacia haematoxylon are found in the central interior duneveld, with undulations of less than 2 m. the unit covers 4.68% of the total mapped area (51 199 ha). the grass stipagrostis amabilis is dominant together with prominent species such as aristida meridionalis, centropodia glauca, eragrostis lehmanniana, e. pallens, stipagrostis uniplumis and brachiaria glomerata. d. calcretes 9. calcrete outcrops the calcrete outcrops, terraces and floodplains along the auob and nossob rivers and at pans (fig. 8) where whitish calcareous and compact sandy soils occur are characterised by a specific plant assemblage. this small unit covers only 0.53% of the mapped area (5 826 ha). scattered individuals of acacia erioloba, a. mellifera, boscia albitrunca, lycium bosciifolium and grewia flava occur in this unit, which is dominated by dwarf shrubs such as rhigozum trichotomum, monechma genistifolium subsp. australe, zygophyllum pubescens, ehretia alba, leucosphaera bainesii, plinthus cryptocarpus, salsola tuberculata and aizoon schellenbergii. the grass layer is poorly developed and is characterised by enneapogon desvauxii, e. scaber, stipagrostis hirtiglumma, cenchrus ciliaris, fingerhuthia africana, stipagrostis ciliata, s. obtusa and the ever-present schmidtia kalahariensis. other prominent dwarf shrubs, forbs and succulents are indigofera auricoma, barleria rigida, chamaesyce glanduligera, ptycholobium biflorum, tribulus zeyheri, cleome angustifolia, adenium oleifolium and euphorbia braunsii. e. plains and terraces along riverbeds and grass pans on compact whitish sand 10. acacia erioloba-rhigozum trichotomum-stipagrostis obtusa river terraces this landscape is characterised by scattered large acacia erioloba trees on slightly compact, calcareous and whitish sand in the dune valleys, near pans and along the auob and nossob rivers throughout the park (fig. 9a). it covers an area of 38 764 ha (3.55% of the mapped area). the most noteworthy woody species are acacia erioloba, a. luederitzii, a. mellifera, boscia albitrunca, grewia flava, searsia tenuinervis, lycium bosciifolium, l. hirsutum and ehretia alba. open to dense stands of rhigozum trichotomum, monechma incanum and aptosimum albomarginatum often develop. the dominant grass species are stipagrostis obtusa, schmidtia kalahariensis, eragrostis lehmanniana, stipagrostis ciliata and s. uniplumis. herbs include talinum caffrum, requienia sphaerosperma, sesamum triphyllum, phyllanthus maderaspatensis, limeum aethiopicum and tribulus zeyheri. 11. acacia mellifera-rhigozum trichotomum-stipagrostis obtusa river terraces this shrubland is related to unit 10 and can be found on calcareous terraces near pans and rivers throughout the park (fig. 9b). it covers an area of 34 431 ha (3.15% of the mapped area). the dominant woody species in this unit is acacia mellifera, with other less frequent woody species being boscia albitrunca, acacia erioloba, grewia flava, lycium bosciifolium, searsia tenuinervis, ehretia alba and rhigozum trichotomum. the dominant grass figure 8 calcrete outcrop (d9) along the nossob river. species are stipagrostis obtusa, s. ciliata, schmidtia kalahariensis and stipagrostis uniplumis. 12. salsola spperiocephalus decussatus river terraces and pan edges this small unit is characterised by dwarf shrubs on the edges of pans and calcareous outcrops along the nossob river and is well-developed in the vicinity of polentswa (fig. 9c). it covers an area of only 1 594 ha, constituting 0.15% of the mapped area (fig. 2). the diagnostic species are salsola tuberculata, eriocephalus decussata, monechma genistifolium subsp. australe and plinthus karrooicus. other conspicuous species are rhigozum trichotomum, salsola etoshensis, eriocephalus pubescens, zygophyllum simplex, leucosphaera bainesii, hypertelis salsoloides, pseudogaltonia clavata and rosenia humilis. grasses are represented by enneapogon desvauxii, stipagrostis obtusa, eragrostis truncata, e. echinochloidea, sporobolus rangei and schmidtia kalahariensis. 107 kalahari gemsbok national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 99 112 13. stipagrostis obtusa river terraces this species-poor unit is characterised by an almost uniform and dense monoculture of the grass stipagrostis obtusa and occurs locally on whitish, compact sandy soils on terraces and floodplains next to the nossob river (fig. 9d). other infrequent species are rhigozum trichotomum, schmidtia kalahariensis, gisekia africana and eragrostis annulata. the unit covers 2 667 ha (0.24% of the mapped area). f . riverbeds and pans 14. sporobolus rangei pan floors essentially two types of pan can be found, i.e. those with a bare pan surface (landscape 14) and grassy pans (landscape 19). bare pan surfaces are commonly referred to as salt pans (leistner 1967, parris 1984, van rooyen 1984). they occur on whitish, calcareous, strongly alkaline and silty soils (fig. 10a) and cover an area of 3 024 ha (0.28% of the mapped area). the vegetation of the pans differs noticeably from the surrounding savannas and often shows a concentric zonation. the vegetation of the pan floor and periphery is characterised by grasses such as sporobolus coromandelianus, s. rangei, eragrostis echinochloidea and enneapogon desvauxii and dwarf shrubs such as zygophyllum tenue, salsola etoshensis, s. namaqualandica, eriocephalus aspalathoides and lycium cinereum. further out from the pans is a zone of scattered trees of acacia erioloba and boscia albitrunca with dwarf shrubs such as rhigozum trichotomum, monechma incanum, m. genistifolium subsp. australe, barleria rigida, asparagus bechuanicus and aptosimum albomarginatum. prominent grass species in this zone include eragrostis truncata, tragus racemosa, eragrostis porosa, stipagrostis ciliata, s. obtusa a b c d figure 9 landscapes of the plains and terraces along riverbeds and grass pans on compact whitish sand: (a) the acacia erioloba-rhigozum trichotomum-stipagrostis obtusa river terraces (e10) along the nossob river; (b) the acacia mellifera-rhigozum trichotomum-stipagrostis obtusa river terraces (e11); (c) the salsola spp.-eriocephalus decussatus river terraces and pan edges (e12); and (d) a monoculture of stipagrostis obtusa on a stipagrostis obtusa river terrace (e13) and schmidtia kalahariensis. around some of the northern pans in the park, e.g. bayip, stands of albizia anthelmintica trees are prominent. 15. panicum coloratum riverbeds this riverbed unit occurs especially in the northern parts of the nossob river and is characterised by a dense grass stratum and large scattered trees of acacia erioloba (fig. 10b). it covers an area of 6 917 ha (0.63% of the mapped area). the dominant grass species are panicum coloratum, eragrostis bicolor, chloris virgata, eragrostis rotifer, e. annulata, e. porosa, schmidtia kalahariensis and enneapogon desvauxii. the unit is rich in forb species, e.g. cullen obtusifolia, hermannia modesta, cucumis africanus, platycarpha carlinioides, heliotropium lineare, boerhavia repens, stachys spathulata, radyera urens, cucumis africana, aptosimum spinescens, geigeria pectidea and g. ornativa. setaria verticillata is a prominent grass species beneath trees in the riverbeds. 16. galenia africana-lagerra decurrens riverbeds this unit occurs locally in the nossob riverbed from the confluence with the auob river northwards to nossob camp and covers 6 321 ha (0.58% of the mapped area). it is differentiated by the shrubs galenia africana, lagerra decurrens and deverra denudata (fig. 10c). scattered acacia erioloba trees occur while chloris virgata, panicum coloratum, schmidtia kalahariensis and eragrostis porosa are the prominent grass species. 17. lebeckia linearifolia riverbeds in the lower auob and nossob rivers near twee rivieren the riverbed is dominated by the shrub lebeckia linearifolia (fig. 10d). this small unit covers only 911 ha (0.08% of the mapped area). lebeckia linearifolia also occurs on dunes in landscape unit 1. 108 original research van rooyen, van rooyen, bothma & van den berg vol. 50 no. 1 pp. 99 112koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a b c d e f figure 10 landscapes of riverbeds and pans: (a) a sporobolus rangei pan floor (f14); (b) the panicum coloratum riverbed near kwang (f15); (c) the galenia africana-lagerra decurrens riverbed (f16) near leeudril; (d) a lebeckia linearifolia riverbed (f17); (e) a large acacia haematoxylon in the auob river in the acacia erioloba-acacia haematoxylon riverbed (f18); and (f ) an example of a grassy pan (f19) in the kalahari 18. acacia erioloba-acacia haematoxylon riverbeds this savanna occurs as a narrow zone on moderately compact calcareous sandy soils on the banks of the auob river and occupies 4 351 ha or 0.40% of the mapped area (fig. 2). it is characterised by large trees of acacia erioloba and a. haematoxylon (fig. 10e) and the shrub rhigozum trichotomum. the prominent grass species are schmidtia kalahariensis, stipagrostis obtusa and eragrostis porosa, and the forb tribulus zeyheri also occurs. 19. grassy pans these pans occur throughout the park and in total they cover approximately 1% of the mapped area (10 782 ha) (fig. 10f). these pans differ from the bare pans in having a lower concentration of minerals, mainly sodium chloride, on the pan floor (parris 1984). the most prominent grass species are sporobolus rangei, stipagrostis ciliata, s. obtusa, eragrostis echinochloidea, enneapogon desvauxii, eragrostis truncata, tragus racemosa and eragrostis 109 kalahari gemsbok national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 99 112 porosa. this landscape is characterised by dwarf shrubs such as salsola etoshensis, eriocephalus pubescens, zygophyllum simplex, z. tenue, rhigozum trichotomum, hypertelis salsoloides and rosenia humilis on the edge of pans. forbs such as aptosimum lineare, trianthema parvifolia, geigeria ornativa, heliotropium lineare, cucumis africanus and indigofera alternans and geophytes such as nerine laticoma and pseudogaltonia clavata are found in this landscape. g. highly irregular dunes 20. highly irregular dunes this striking landscape with ellipsoidal parabolic dunes (eriksson et al. 1989) is found in a few isolated patches in the interior duneveld of the park (fig. 11). the unit covers only 917 ha (0.08% of the mapped area) and is characterised by scattered trees and shrubs of boscia albitrunca and acacia mellifera. the prominent grass species include stipagrostis amabilis, eragrostis trichophora, brachiaria glomerata and schmidtia kalahariensis. discussion the whole area of the kgnp is fairly homogeneous and for the most part belongs to the kalahari duneveld bioregion (mucina & rutherford 2006), characterised by acacia erioloba, rhigozum trichotomum and schmidtia kalahariensis. the vegetation structure is predominantly that of a shrubby grassland, except in the north and along the auob and nossob rivers where an open tree savanna is found. topographically the area can be divided into dunes (dune crests and dune valleys); sandy plains with red to pinkish sand; and rivers (including terraces and calcrete outcrops) and pans on whitish, compact calcareous sand and clay. the vegetation on fine-textured soils, such as those found in the pans, frequently shows a karroid nature (leistner 1967). the southern kalahari is described as a floristically impoverished area. the homogeneity of the relatively recent sand deposits leads to reduced landscape differentiation and thus low niche differentiation. the kalahari is therefore not a ‘hotspot’ in terms of plant species richness or rarity. the vegetation and landscapes around pans and along the rivers are the most diverse (van rooyen & van rooyen 1998). however, there are a number of plant species that are of special interest in the kgnp. these species are all common and abundant species elsewhere but are of rare occurrence in the park. for example, parkinsonia africana is known only by a small population that occurs in the lower nossob river. other woody species that are relatively rare in the park include boscia foetida subsp. foetida on calcrete ridges in the south of the park, cadaba aphylla, diospyros austro-africana and ehretia alba. a few individuals of dichrostachys cinerea are found in the north near grootbrak and union’s end in landscape unit 6. likewise, catophractes alexandrii is limited to the extreme northern part of the kgnp on calcrete ridges adjacent to the nossob river, although it is abundant on the botswana side of the kgalagadi transfrontier park. albizia anthelmintica and to a large extent also acacia luederitzii have restricted distributions on the south african side of the kgalagadi transfrontier park. albizia anthelmintica reaches its southwestern distribution limit in south africa in the park whereas acacia luederitzii is one of the few species endemic to the kalahari. terminalia sericea is locally prominent on the higher dunes in the northern part of the park, especially in landscape unit 4. the false camel thorn acacia erioloba x acacia haematoxylon is relatively rare but is found in the southern parts of the park along the auob and nossob rivers. woody species that have been recorded on the botswana side of the kgalagadi transfrontier park and not yet on the south african side include several commiphora species, grewia flavescens, ochna pulchra, vangueria infausta, tylosema esculentum and ziziphus mucronata. the shrub phaeoptilum spinosum was recorded south of the park but not yet in the kgalagadi transfrontier park. the characteristic tree species in several of the landscapes, i.e. acacia erioloba, acacia haematoxylon and boscia albitrunca, are figure 11 an isolated patch of ellipsoidal parabolic dunes or highly irregular dunes (g21) in the kalahari gemsbok national park listed as protected trees through the national forestry act by the department of water affairs and forestry. acacia erioloba is regarded as a keystone species and the survival of many other animal and plant species depends on it (dean et al. 1999). although wood harvesting, especially that of acacia erioloba, is a serious problem outside the boundaries of the park, wood harvesting is not permitted inside the park. alien invasive plant species are not a big problem in the kgnp. large prosopis glandulosa trees occur in the camps but have not yet been noticed spreading into the natural vegetation. other species that have been listed as invasive in the kalahari environment include prosopis chilensis, verbesina enceloides and argemone ochroleuca. the landscapes most prone to invasion by alien species are the riverbeds (landscapes 15 to 18). the aims of the present map were to identify landscapes and their associated vegetation to assist in vegetation and animal conservation and tourist management. landscapes that play an important role in the ecological relationships of the region include the calcretes, riverbeds and pans (landscape units 9 to 19). these landscapes cover only about 10% of the total area but are focal points for the large animals of the region (parris 1970, 1984, child et al. 1971, bothma & mills 1977, mills & retief 1984, bergstrom & skarpe 1999). most of the animal sightings are concentrated in the riverbeds and pans that cover less than 4% of the total area. the interior dunes and plains (landscape units 1 to 8) are usually used by wildlife only in the wet season when the food plants are not as nutrient deficient as during the dry season (stapelberg 2007). although animals tend to congregate in the river systems, grazing impacts are localised and controlled by a natural rotational grazing system, with the availability and nutritional quality of food rather than water controlling animal numbers (van rooyen & van rooyen 1998). the seasonal migratory movements of wildlife is essential to maintain good veld condition and ensures that a long-term ecological balance prevails, which should continue if the present large size of this conservation area is maintained (van der walt et al. 1984). artificial water provision does affect some aspects of the ecology of the system (van rooyen et al. 1994), but these watering points are deemed necessary from a tourist point of view (knight & knight-eloff 1988). the landscapes most affected by artificial water provision are landscapes 9 to 18. one of the major uses envisaged for the present landscape map is that it would serve as basis for planning monitoring programmes. aspects of the vegetation that need to be monitored include the following: veld condition and the ecological capacity for large • herbivores within each landscape; the impact of the provision of artificial watering points • on landscapes 9 to 18; 110 original research van rooyen, van rooyen, bothma & van den berg vol. 50 no. 1 pp. 99 112koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za trends in population density, frequency and cover • of possible encroacher plant species, e.g. rhigozum trichotomum, acacia mellifera and geigeria ornativa (all landscapes); the population structure of keystone species such as • acacia erioloba, acacia haematoxylon, acacia luederitzi and boscia albitrunca; and the occurrence of alien plant species such as • prosopis spp. and argemone ochroleuca in the riverbeds and pans (landscapes 10 to 19). these monitoring efforts should determine the dynamics and trends of the ecosystem attributes and the extent to which the conservation objectives are being met. geographic information systems (gis) would be an ideal tool for managing such spatially related databases (kotzé & fairall 2006). it is imperative that the results of monitoring be used to adapt management policies when necessary. by basing the monitoring efforts on the present landscape units, this map will assist in resource management that is based on the inherent ecological capacity of the ecosystem. acknowledgements we gratefully acknowledge the assistance of south african national parks and specifically dries engelbrecht, nico van der walt, nardus du plessis and hugo bezuidenhout. johan kriek and the van rooyen family assisted with the field work. references acocks, j.p.h. 1953. veld types of south africa. memoirs of the botanical survey of south africa, 57: 1–146. bergström, r. & skarpe, c. 1999. the abundance of large wild herbivores in a semi-arid savanna in relation to seasons, pans and livestock. african journal of ecology, 37(1): 12–26. bothma, j. du p. & de graaff, g. 1973. a habitat map of the kalahari gemsbok national park. koedoe, 16: 181–188. bothma, j. du p. & mills, m.g.l. 1977. ungulate abundance in the nossob river valley, kalahari desert. proceedings of the xiiith international congress of game biologists, atlanta, pp. 90–102. bothma, j. du p. & swart, d.r. 1990. landsat-based remapping of the southwestern kalahari. koedoe, 33: 89–93. bothma, j. du p., van hoven, s. & le riche, e.a.n. 1993. gps corrected and gis based remapping of the kalahari gemsbok national park and the adjacent area in botswana. koedoe, 36: 105–108. brynard, a.m. 1958. verslag insake voorlopige ondersoek rakende toestande in die nasionale kalahari-gemsbokpark. koedoe, 1: 162–183. child, g., parris, r. & le riche, e. 1971. use of mineralised water by kalahari wildlife and its effects on habitats. east african wildlife journal, 9(1): 125–142. coetzee, b.j. & werger, m.j.a. 1975. an east-west transect through africa south of the tropic of capricorn. bothalia, 11: 539–560. dean, w.r.j., milton, s.j. & jeltsch, f. 1999. large trees, fertile islands and birds in arid savanna. journal of arid environments, 41(1): 61–78. eriksson, p.g., nixon, n. snyman, c.p. & bothma, j. du p. 1989. ellipsoidal parabolic dune patches in the southern kalahari desert. journal of arid environments, 16: 111–124. gertenbach, w.p.d. 1983. landscapes of the kruger national park. koedoe, 26: 9–21. giess, w. 1971. a preliminary vegetation map of south west africa. dinteria, 4: 5–114. gubb, a. 1980. vegetation map of the kalahari thornveld. unpublished mimeograph. kimberley: mcgregor museum. knight, m.h. & knight-eloff, a.k. 1988. the importance of borehole water and lick sites for kalahari south africa ungulates. journal of arid environments, 15: 269–282. kotze, i.j.d.f. & fairall, n. 2006. using landsat tn imagery to map fynbos plant communities: a case study. journal of wildlife research, 36: 75–87. leistner, o.a. 1959a. notes on the vegetation of the kalahari gemsbok national park with special reference to its influence on the distribution of antelopes. koedoe, 2: 128– 151. leistner, o.a. 1959b. preliminary list of plants found in the kalahari gemsbok national park. koedoe, 2: 152–172. leistner, o.a. 1967. the plant ecology of the southern kalahari. memoirs of the botanical survey of south africa, 38: 1–172. leistner, o.a. & werger, m.j.a. 1973. southern kalahari phytosociology. vegetatio, 28(5/6): 353–397. low, a.b. & rebelo, a.g. 1998. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. lubbinge, j.w. 1999. ‘n fitososiologiese studie van die suidelike kalahari duineveld. m.sc. dissertation. pretoria: university of pretoria. mills, m.g.l. & retief, p.f. 1984. the response of ungulates to rainfall along the riverbeds of the southern kalahari. koedoe (suppl.), 27: 129–141. mostert, j.w.c., roberts, b.r., heslinga, d.f. & coetzee, p.g.f. 1971. veld management in the ofs region. pretoria: government printer. mucina, l. & rutherford, m.c. 2006. the vegetation of south africa, lesotho and swaziland. strelitzia, 19, pretoria: south african national biodiversity institute. mucina, l., rutherford, m.c. & powrie, l.w. 2005. vegetation map of south africa, lesotho and swaziland. pretoria: south african national biodiversity institute. parris, r. 1970. important role of the kalahari pans. african wildlife, 24: 234–237. parris, r. 1984. pans, rivers and artificial waterholes in the protected areas of the south-western kalahari. koedoe (suppl.), 27: 63–82. pole-evans, i.b. 1936. a vegetation map of south africa. memoirs of the botanical survey of south africa, 15: 1–23. skarpe, c. 1986. plant community structure in relation to grazing and environmental changes along a north-south transect in the western kalahari. vegetatio, 68: 3–18. stapelberg, f.h. 2007. feeding ecology of the kalahari springbok antidorcas marsupialis in the kgalagadi transfrontier park, south africa. msc dissertation, pretoria: university of pretoria. story, r. 1958. some plants used by the bushmen in obtaining food and water. memoirs of the botanical survey of south africa, 30: 1–155. thomas, d.s.g. & shaw, p.a. 1991. the kalahari environment. cambridge: cambridge university press. tyson, p.d. 1986. climatic change and variability in southern africa. cape town: oxford university press. van der walt, p.t., retief, p.t., le riche, e.a.n., mills, m.g.l. & de graaff, g. 1984. features of habitat selection by large herbivorous mammals and the ostrich in the southern kalahari conservation areas. koedoe (suppl.), 27: 119–128. van rooyen, n. 1996. phytosociology, vegetation dynamics and conservation of the southern kalahari. unpublished report on a project executed on behalf of the department of environmental affairs and tourism. pretoria: university of pretoria. van rooyen, n. 2000. vegetation survey of the gemsbok national park, botswana and mapping of the kgalagadi transfrontier park. stellenbosch: peace parks foundation. van rooyen, n., bezuidenhout, d.j., theron, g.k. & bothma, j. du p. 1990. monitoring of the vegetation around artificial watering points (windmills) in the kalahari gemsbok national park. koedoe, 33: 63–88. van rooyen, n., bredenkamp, g.j., theron, g.k., bothma, j. du p. & le riche, e.a.n. 1994. vegetation gradients around artificial watering points in the kalahari gemsbok national park. journal of arid environments, 26: 349–361. 111 kalahari gemsbok national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 99 112 van rooyen, n., van rensburg, d.j., theron, g.k. & bothma, j. du p. 1988. a check list of flowering plants of the kalahari gemsbok national park. koedoe, 31: 115–135. van rooyen, n. & van rooyen, m.w. 1998. vegetation of the southwestern arid kalahari: an overview. transactions of the royal society of south africa, 53(2): 113–140. van rooyen, t.h. 1984. the soils of the kalahari gemsbok national park. koedoe (suppl.), 27: 45–62. werger, m.j.a. 1986. the karoo and southern kalahari. in: evenari, m., noy-meir, i. & goodall, d.w. (eds.). hot desert and arid shrublands. amsterdam: elsevier, pp. 283–359. werger, m.j.a. & coetzee, b.j. 1978. the sudano-zambezian region. in: werger, m.j.a. & van bruggen, a.c. (eds.). biogeography and ecology of southern africa. the hague: junk, pp. 301–462 112 abstract introduction and background research study site, design and methods results discussion conclusion acknowledgements references about the author(s) reitumetse m.h. sethaba school of animal, plant and environmental sciences, faculty of science, university of the witwatersrand, johannesburg, south africa mary scholes school of animal, plant and environmental sciences, faculty of science, university of the witwatersrand, johannesburg, south africa citation sethaba, r.m.h. & scholes, m., 2021, ‘management guidelines for the conservation of heritage resources in wakkerstroom, mpumalanga’, koedoe 63(1), a1647. https://doi.org/10.4102/koedoe.v63i1.1647 original research management guidelines for the conservation of heritage resources in wakkerstroom, mpumalanga reitumetse m.h. sethaba, mary scholes received: 03 july 2020; accepted: 02 nov. 2020; published: 20 jan. 2021 copyright: © 2021. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract conservation of heritage resources, both architectural and cultural, focuses on the important of historical context, identity and to provide livelihoods for people. management and stakeholder engagement are fundamental for conservation. this article describes the contribution of stakeholder perceptions in the development of management guidelines for the conservation of heritage resources in the town of wakkerstroom, mpumalanga. the concept of conservation is usually used with reference to the natural heritage; however, this should also include architectural and cultural conservation, in the same location, as is done in many other parts of the world. the study of the combined architectural and cultural heritage in wakkerstroom is one of the first of such studies in south africa. the natural heritage resources were identified and a stakeholder map developed after conducting interviews with 27 participants in the township of esizameleni and 15 participants in the town. resources identified included the provincial heritage sites of the wakkerstroom wetland, the paul kruger bridge, st mark’s anglican church and the old courthouse building. heritage value definitions in the town area were centred on inheritance, whilst township stakeholders valued the identity aspect of heritage. a lack of communication, participation and awareness around heritage conservation was identified as a key issue amongst the communities. in addition, unlike natural heritage conservation, where the wakkerstroom natural heritage association takes care of the wetland, no organisation was found to be responsible for cultural heritage conservation in the town. the municipality urgently needs to address these concerns by developing an integrated management plan for heritage resource conservation. conservation implications: conservation includes more than just the conservation of natural heritage. the wakkerstroom natural heritage association (wnha) manages natural heritage, but there is no organisation managing cultural heritage. heritage conservation would benefit from the development of an integrated management plan lead by dr pixley ka isaka seme local municipality together with the identified stakeholders. keywords: heritage; conservation; resource management; stakeholder engagement; sustainable communities. introduction and background the term ‘heritage’ is used loosely to describe everything that is inherited or handed down to future generations from the past (lowenthal 2005). natural heritage resources usually refer to land, water, animals and plants. cultural heritage resources include buildings, art, traditions and practices of cultural and historical value (lowenthal 2005). heritage can also be described as tangible ‘a monument, group of buildings, or site of historical, aesthetic, archaeological, scientific, ethnological or anthropological value’ (kirshenblatt-gimblett 2004); or intangible such as ‘all forms of traditional and popular culture such as collective works originating in a given community, and based on tradition’ including music, dance and traditional medicine (united nations educational, scientific and cultural organisation (civallero 2007; unesco 1989). heritage resources have been shown to be important in, amongst other factors, issues of history, identity and maintenance of livelihoods for people in south africa and the world (nzama 2009; parts et al. 2011; su et al. 2018). conservation of both natural and cultural heritage provides the benefit of encouraging economic activity through an integrated development plan, one aspect of which could be tourism. conservation of the natural heritage is often focussed on biodiversity, ecosystem integrity and ecosystem goods and services and seems more tangible than conserving cultural heritage, which has many more components. resource management serves as a tool for resource conservation, enabling the protection of resources through the monitoring of use, preventing damage to, and over-exploitation of, valued resources (cleere 2010). plans for sustainable resource management must consider human–environment interactions. understanding these interactions and what influences them is an important component of effective management strategies. stakeholder engagement is an important process in resource management, representing inclusive management strategies and has been on an incremental trajectory in the environmental sector since the 1980s (young et al. 2013). the united nations conference on environment and development (unced) identified stakeholder engagement in management and decision-making as one of the steps instrumental in the move towards sustainable development (unced 1992). the argument for this has been mainly around bringing different perspectives in the form of knowledge and values into the processes of decision-making, in order to make better decisions where people and development are concerned (chirikure et al. 2008; renn 2006). studies have shown that where effective stakeholder engagement was carried out, trust is built, communication and decision-making processes are improved (munton 2003) and there is rapid adoption of management strategies in communities (beierle & konisky 2001; sultana & abeyasekera 2008). effective communication and consultation between management and other stakeholders are also important considerations for successful stakeholder engagement. this is especially true in areas where activities such as tourism intersect with heritage conservation, potentially creating conflicts. communication and consultation are vital to the successful and sustainable execution of activities (aas, ladkin & fletcher 2005; keitumetse 2011; peters 1999). in heritage conservation and resource management, stakeholders include those affected by, or interacting with, the heritage resources in question (blundo et al. 2018). the perceptions of stakeholders are a tool used in stakeholder engagement to understand how people value their resources (carter & bramley 2002). understanding perceptions gives an indication of the actions that should be taken towards resource conservation and management (bakri et al. 2015). incorporating perceptions of people into management plans facilitates effective management and subsequently supports the move towards the goal of sustainable development through resource maintenance and conservation (jaafar, noor & rasoolimanesh 2015). in addition to an improved or maintained state of the environment, well-managed resources may also lead to wider community involvement and greater social cohesion, contributing to sustainable communities (hribar, bole & pipan 2015). the significance of resources is attributed according to the value placed by people on heritage resources (the burra charter 1999). heritage value is defined by díaz-andreu (2017) as ‘the meanings and values that individuals or groups of people bestow on heritage’. heritage value definitions may differ according to the communities to which people belong, race, gender and other background information (deacon & smeets 2013). heritage-management plans are important in identifying important resources, understanding their value and significance, as well as provisioning for their protection, use and maintenance in the future. goal 11 of the sustainable development goals, adopted by the united nations in 2015, states the aim to ‘make cities and human settlements inclusive, safe, resilient and sustainable’. one of the targets under this goal is to ’strengthen efforts to protect and safeguard the world’s cultural and natural heritage’ (patel et al. 2017). sustainable communities require a just and inclusive decision-making process (caffyn & jobbins 2003). this study aims to contribute to this goal by using stakeholder perceptions in an assessment of the natural and cultural heritage assets for the town of wakkerstroom, mpumalanga. this was done in order to make suggestions towards the development of management guidelines for their conservation. there are 10 world heritage sites in south africa and numerous national and provincial sites (unesco n.d.). one of these provincial sites is the museum town of pilgrim’s rest in mpumalanga, which could be referred to as a heritage town (rahman 2013). locations such as the horseshoe waterfall, joubert bridge and the natural rock bridge are all declared as provincial heritage sites in this town by the south african heritage resources agency (sahra 2020). by identifying resources, their uses, conservation issues, as well as stakeholders, stakeholder issues and interests, the development of future management actions aimed at resource conservation and sustainability that can be facilitated. no studies of this sort have been conducted for the town, making outputs of the study valuable for environmental heritage protection in wakkerstroom, and for other heritage-rich locations. this study aimed to use stakeholder perceptions to contribute to the development of management guidelines for the conservation of heritage resources in the town. in south africa, the concept of conservation is mostly focused on components of the natural heritage, but this concept can, and should, be extended to include architectural and cultural conservation, in the same location. a seminal work was published by stubbs (2009) in which he proposed a global view of conservation. this approach has been adopted in several case studies in the usa, the uk, malaysia and italy (de medici & senia 2007; harun 2011; lapadula & quiroga 2012; pereira & marker 2016). this study explicitly includes the conservation of the natural and the architectural and cultural heritage in wakkerstroom and is one of the first of such studies in south africa. research study site, design and methods study site the study was conducted in wakkerstroom, a small town in the mpumalanga province bordering the province of kwazulu-natal, south africa, which is part of the dr pixley ka isaka seme (dpkis) local municipality. the town was established in 1859 (theal 2010), occupies an area of 87.68 square kilometres (km2) and has a population of approximately 6800 residents (dpkis local municipality idp 2011). the primary industry is maize, sheep and cattle farming which are conducted at commercial and subsistence levels. the town is one of the oldest towns in the region, and is subsequently rich in heritage, both natural and cultural. the area is home to the wakkerstroom wetland reserve (commonly known as the vlei) which has a natural heritage status and provides habitats for numerous bird species. the grasslands are also protected by the national environmental management biodiversity act (act no. 10 of 2004). the town is inhabited by people of diverse heritage, including the ndebele, zulu, afrikaaner, british and their descendants. there are many farming families (e.g. uys, oosthuizen and greyling) which have been on the same land for seven generations, making it a suitable site for heritage perception exploration (gardiner 2017). the town consists of a ‘township’ and ‘town’ area as a result of segregation during the apartheid era. the township area is known as esizameleni, and comprises approximately 1650 ndebele and zulu households (dpkis local municipality idp 2011). this area is primarily characterised by underdevelopment, unemployment, low education levels and low access to adequate sanitation (jürgens et al. 2013; kalichman & simbayi 2003; salo 2003; wood et al. 2010). the town area conversely displays a higher level of development, with a larger number of white afrikaaner inhabitants (statssa 2011). esizameleni is about 4 km from the centre of the town. study design and methods this study was divided into three parts: (1) a desktop study to identify the heritage resources; (2) a stakeholder map created to identify groups of stakeholders and the structure of the community and (3) interviews with stakeholders to assess their perceptions that were used to develop management guidelines for conservation. the desktop study used a variety of sources, including the wakkerstroom official website (https://www.wakkerstroom.co.za/) and south african history online (https://www.sahistory.org.za/) to obtain baseline information about cultural and natural heritage resources in the town. the register of the south african heritage resources agency (sahra declared sites 2020) was also consulted for a list of declared sites. publications from the town including ‘wakkerstroom: jewel of mpumalanga’ (hofmeyer, smith & smith 2009) as well as the ‘wakkerstroom; bird and nature guide’ (tarboton 2004) were used to identify sites of heritage importance. information relating to the location, history and the known uses of the resources was gathered in order to understand the relevance of the heritage resources and aspects identified. a publication titled ‘wakkerstroom: a conservation study’ (peters 1995) was also consulted for information on the architecture in the town. a table was constructed summarising the resources identified, the type of resource and the importance or value attributed to the resource. in order to identify stakeholders who should be interviewed, a stakeholder map was developed with the help of managers of conservation organisations in the town by identifying users of the resources in the communities who would be affected by, or be involved in, changes in the condition of the resources, as well as any management practices implemented. managers responsible for the protection and monitoring of heritage resources were also identified through various organisations assigned with the task of management of resources in the town. the identified stakeholders were then grouped according to categories that would represent the structure of the community of wakkerstroom, and ensure that stakeholder groups were adequately represented where resource conservation is concerned according to the best practice of stakeholder engagement as suggested by hare and pahl-wostl (2002). categories were developed according to where stakeholders lived, and where possible, the economic and organisational role of stakeholders in the community. a stakeholder map was used to document the stakeholders identified and the strength of stakeholder linkages in relation to the main management body which was the dpkis local municipality. this, in conjunction with subsequent interview data also enabled assessment of roles in resource protection, influence in resource conservation and resource-conservation plan execution (varvasovszky & brugha 2000). two semi-structured interview questionnaires were used because they allowed for more open-ended questions and the collection of a wider range of information. one questionnaire was used for interviews with community members including business owners, residents from the town and esizameleni areas of wakkerstroom. interviews were conducted in english and isizulu according to the choice of the participant. the interviews addressed questions related to demographic information, the understanding of heritage and heritage values, identification of resources valued and the reasons for the perceived importance. use of the resources and participation in heritage activities were also investigated including the contribution of participants to the protection of these resources. sources of information regarding heritage resource aspects and threats perceived to resources were also addressed by the interview questions. the second interview questionnaire was addressed to the caretakers and managers of organisations tasked with the preservation of heritage resources in the town in order to gather more information from these stakeholders of the town region regarding resource management. organisations included birdlife south africa, working on fire and the wakkerstroom natural heritage association (wnha). the second interview instrument was similar to the first, differing only in the addition of questions related to management actions. this was in order to establish current management practices and to use the information gathered in developing suggestions for future management plans. in both the town and township areas, key informants were initially identified and then snowball sampling was used as described by sadler et al. (2010). in the town area, a total of 15 people were interviewed (including three managers of the aforementioned conservation organisations), whilst 27 people were interviewed from the township area. interviews were 30 min long; interviews people in the town area took place at a local hotel restaurant, and interviews in the township were conducted at participant households. a classical ‘stakeholder map’ as defined in the literature was very difficult to achieve in this small village, as many people, especially those in the town, played multiple roles. they were simultaneously owners of businesses, chair-people of volunteer organisations and part-time residents. the downside of this may be offset by the fact that many people understood the need for conservation as they were embedded in the activities. the municipality was also contacted for interviews as it was important that they be interviewed; however, they did not respond. officials in the municipality (e.g. town manager, mayor and chief financial officer) were all struggling with financial compliance in order to prevent intervention by the authorities and had no interest in service delivery or in the conservation of the natural and heritage resources (municipal money profile: dr pixley ka isaka seme 2020). to analyse the data, content analysis was carried out – a method identified to be most effective in describing the qualitative data gathered in this study. content analysis also lends itself to a quantitative description of the data. responses from town and township groups of stakeholders were transcribed into two microsoft excel spreadsheets. where applicable, categories were developed with similar themes or answers falling under the same category. responses were coded according to categories and expressed as percentages within and between categories. bar graphs were produced for each question, in each area (town or township). bar graphs of demographic data (average age of participants, education levels and average time participants have lived in the town) were also produced. no statistical tests were conducted on these data as sample sizes were too small. results inventory of heritage resources natural and cultural heritage resources, both intangible and tangible, were identified. buildings and natural features in the environment were identified (table 1). significance was mainly attributed to architectural quality preserved over time and the history of the buildings conferring cultural value. table 1: heritage resources and activities identified in the town of wakkerstroom. three structures listed under national monument status are located in the town. one of these is the paul kruger bridge, referred to as ‘the old road bridge’, marthinus wesselstroom, under the sahra monuments register. the bridge is a steel structure spanning 12.2 metres (m) in length built with material imported from europe (peters et al. 2001), and is archived under national monument status (site reference: 9/2/279/0006), declared as a provincial heritage site under sahra and listed under grade ii heritage protection (resources significant within the context of a province – administered by the relevant provincial heritage authority [phra]) (sahra declared sites 2020). the second structure is the old court house building (old magistrates office, engelbrecht street, wakkerstroom) which dates back to 1897. similar to the paul kruger bridge, the building is archived under national monument status. the building is also declared under grade ii provincial heritage protection as a provincial heritage site (site reference 9/2/279/0002) (sahra declared sites 2020). the third structure is the st mark’s anglican church, also known as the old apostolic church of africa, dating back to the 1870s. the building, gazetted in 1982, is the third building in the town archived under national monument status and currently declared as a grade ii, provincial heritage site (site reference: 9/2/279/0006-001). the building was erected following the anglo-boer war, serving as a residence for an officer who led the british army (south african heritage resources information systems n.d.). the church is located on erf 82 kerk street, it is privately owned, but local anglicans still conduct services in the church. the wakkerstroom wetland is one of the most notable natural heritage resources mentioned in the literature on the town. the wetland measures about 1000 hectares (ha) (begg 1989) and is located at the upper reaches of the tugela catchment. the wetland does not have a high floral diversity (kotze, breen & klug 1994) but is dominated by a dense growth of carex acutiformis (ehrh.), phragmites australis (cav.) trin. ex steud. and typha capensis (rohrb.) (joubert & ellery 2013). this wetland is however important as a habitat, and a nesting and foraging area for rare birds such as the white-winged flufftail (sarothrura ayresi) (tarboton 2004), as well as the grey crowned crane (balearica regulorum), both of which are some of the rarest birds in africa (khan 2010; oellermann, darroch & klug 1994; reid et al. 2005). the wetland attracts multiple bird species from around the world, including migratory birds from eurasia and intra-african migrants (joubert & ellery 2013). invertebrate species are well represented, as well as species of fresh-water otters (including the spotted-necked otter [lutra maculicollis]) (wakkerstroom bird club 2016; zhang et al. 2011). in addition, the wakkerstroom vlei is also used as a grazing site for cattle by local farmers (kotze 2013). harvesting of plants for medicinal purposes as well as water supply are valued services provided by the vlei. the surrounding grasslands and forests are highly diverse with the species list tally currently at 1300 and further work is under progress (sanbi 2020a). stakeholders stakeholders identified included the organisations tasked with the conservation and management of heritage resources in the town of wakkerstroom. the wnha together with the working on fire programme and birdlife sa are the main organisations tasked with resource conservation. these organisations mainly focus on the protection of natural heritage resources in the town, including bird and bird habitat conservation, wetland ecosystem protection and fire control in the wetland and grassland areas. these organisations are part of the stakeholders of the heritage resources identified (figure 1). although, they are also, part of the town community (as explained earlier), the leaders of these organisations were interviewed separately as they served a management function abrogated by the municipality. figure 1: stakeholder map. two main stakeholder groups were identified: the esizameleni township and the town areas of wakkerstroom. in the town region, stakeholders included businessmen, residents and tourist stakeholder groups because of the role they played as dominant stakeholder subgroups within the town area. tourists were included because of the role tourism plays in motivating conservation of the town. these stakeholder groups interact directly with resources, are involved in, impacted and affected by, resource management and conservation in the town, and subsequently have resource management and conservation interests. in the township area of esizameleni, the low levels of tourism activity, business operations and interactions with resource conservation activities in wakkerstroom resulted in no clear subgroups being identified. one group of stakeholders was therefore formed and labelled as ‘residents’. establishing a ‘business people’ subgroup similar to that of the town area could not take place in the township stakeholder group. this is because, although there are individuals that manage businesses such as spazas and shebeens in the township, the ownership, longevity and location of these businesses change frequently, making it difficult to identify actors as stakeholder subgroups. a ‘residents’ group was therefore perceived as a representative stakeholder grouping for the purpose of assessing conservation linkages, roles and influence in resource conservation and management. the dpkis local municipality is included as the main actor in management of resources in the town. the municipality, through its role as government, should serve as the main body to which all other stakeholders are directly or indirectly linked. according to interviews conducted with managers of conservation organisations in wakkerstroom, people employed by the municipality in the town can, however, be described as relatively inactive, paying little attention to heritage resource conservation in the way of financing and supporting town conservation activities. this leaves the bulk of management to town non-governmental organisations such the wnha and volunteer work by members of the community, despite the management responsibility of the municipality, by virtue of it being elected. stakeholder linkages, their influence and involvement in the conservation of heritage resources although conservation decisions made regarding heritage resources affect all stakeholders identified, the involvement, influence and linkages of stakeholders, and subsequently their roles in resource preservation and management differ (figure 1). through interviews, the community of the town area was shown to have a greater involvement in resource conservation and management actions taken, with the residents, those running businesses within the town and part-time residents (people who have holiday homes and visit at different times of the year) playing the biggest role. this means that there is a greater influence of stakeholders in the town area on resources compared to those in the community of esizameleni. from interviews conducted, it was evident that the esizameleni township community plays a small role in conservation actions implemented by the town. this is reiterated by the low representation of this stakeholder group in the heritage conservation organisations of the town. stakeholders in the esizameleni area were found to have little knowledge or involvement in resource conservation activities (figure 1). connections between township stakeholders and stakeholders of the town area are also shown to be weak because of the lack of communication and mutual activities between the two stakeholder groups. perceptions towards heritage the highest level of education amongst participants of the esizameleni township was found to be a high school qualification (figure 2a) in contrast to the town area where the highest level of education reported amongst participants reached postgraduate degree levels, and all participants had some level of education (figure 2b). township participants linked heritage definitions mainly to the theme of ‘identity’ with references to cultural and traditional identities, followed by ‘inheritance’ alluding to customs and traditional practices passed down through generations (figure 3a). town participants associated heritage mainly with the theme of ‘inheritance’ which denoted the passing down of tangible resources (e.g. natural resources, farms or buildings), generationally (figure 3b). township participants identified threats to heritage under the category of ‘people and livestock’ (livestock referring to the farming practices associated with keeping livestock) including the negative attitudes towards conservation by town participants as the greatest threats to heritage resources (figure 4a). town participants identified mining as the top threat to natural heritage resources (the wetland) followed by concerns about farming practices related to the number of cattle grazing in the wetland, which may lead to overgrazing. a lack of interest in conservation and general pollution were also mentioned as threats (figure 4b). figure 2: levels of education of participants. the highest levels of education reported by participants in esizameleni and town communities in wakkerstroom, mpumalanga. proportions of responses (%) (y-axis) against education level (x-axis) are illustrated in the graphs. ‘late high school’ (grade 10–12), ‘early high school’ (grade 8–9), ‘late primary’ (grade 4–7), ‘higher education’ (diplomas and post schooling courses) and ‘postgraduate degrees’ were educational categories used for participants. figure 3: heritage definitions by frequency. the heritage definitions provided for the term ‘heritage’ by participants in the esizameleni township and the town area. proportions of responses (y-axis) amongst participants are shown against the themes of heritage definitions provided (x-axis). figure 4: threats identified. threats to heritage resources identified by participants in the township and town are proportions of threats mentioned (%) (y-axis), against the threats mentioned by participants (x-axis) are shown. in the township region, ‘people and livestock’ are of greatest concern. this includes grazing by livestock and littering. in the town region, ‘mining’ is stated as the greatest concern amongst participants. the development of management guidelines for heritage resource conservation the aim of this study was to understand how stakeholder perceptions (presented above) may contribute to the development of management guidelines for the conservation of heritage resources in the town of wakkerstroom, mpumalanga. the interview responses from the wnha, working on fire and birdlife sa organisations showed that there is no integrated management plan for the conservation of heritage resources. the municipality, as part of the government, is the main body responsible for the management of heritage resources within towns and townships located within the dpkis local municipality (rautenbach, hart & naudé 2015) but it has abrogated its responsibility for conservation and management of the wetland to the wnha, which remains legislatively compliant under the national environmental act (act no. 107 of 1998). interviews also revealed that there are no ongoing conservation activities or management plans associated with the cultural heritage. the only activity, which the wnha organises for heritage day, is a series of guided walks and information brochures to select buildings. there are several conservation projects conducted by the wnha, but most of these are carried out on an ad hoc basis and do not form part of any management plan which should be developed using a standard framework of setting objectives, time frames, budgets, monitoring and evaluating planned actions and activities. in terms of some current management actions for conservation, the wnha reported conservation projects associated with the wetland including monitoring the number of cattle grazing in the vlei, the amount and removal of litter, the removal of invasive alien plants and limited spraying with herbicide to maintain patch diversity. fires occur frequently in wakkerstroom. fire management in the wetland is also undertaken by the wnha working closely with the working on fire organisation. in winter months, fire-breaks prevent the spread of fires and fire damage. patch burning in the vlei will be monitored to limit the dominance of phragmites australis. interviews also revealed that the wnha and birdlife sa are advocating the wetland be declared a ramsar site. the ramsar convention is targeted towards promoting the conservation and wise use of all wetlands through local and national actions and international cooperation, as a contribution towards achieving sustainable development throughout the world. in south africa, a wetland can however only be included as a ramsar site when declared a protected area under the national environmental management: protected areas act (act no. 57 of 2003). currently, the wetland is classified as an informally protected area, not yet formally protected by legislation (sanbi 2020). discussion the study identified a combination of natural and cultural heritage resources in the town of wakkerstroom; the rich heritage is revealed in the diverse assemblage of heritage-related sites and activities, linked in many ways to the town’s history. the results provide an interesting test case as to whether a single location could be used to show the benefit of a single integrated management plan for the conservation of both the natural and cultural heritage resources. this study provides some preliminary information which would be needed to develop such a plan. the resources are relatively well-defined, which is a first and very important step. the wakkerstroom wetland was identified as the main natural heritage resource of importance in the town. in terms of cultural heritage, three sites classified as national heritage resources were identified (the paul kruger bridge, the old magistrates court and st mark’s anglican church), along with two annual town heritage festivals of music and crafts which celebrate, through artistic display, the cultures of various inhabitants of the region and beyond. this diversity in cultural and natural heritage, and high historic value of the town, support the proposed classification of the town under the definition of a ‘heritage town’, as used by rahman (2013). the study also highlighted the opportunities and limitations that would need to be considered and included in the development of a management plan. these included value definitions, stakeholder participation in decision-making and engagement in activities, styles of communication, language and awareness; trade-offs needed to achieve sustainable development. value definitions were found to be different amongst the town and township communities interviewed. the value attributed to heritage amongst township participants related to the importance of heritage in defining identity. this was coupled to the mention of cultural practices and traditions at personal and household levels as important heritage aspects. in the town area, the natural heritage was valued most, with the wetland mentioned as the most important heritage resource. this may be because of the greater interaction the people in the town have with the economic benefits of the wetland through tourism, even though many of the ‘bird guides’ trained by birdlife, live in the township. the generational transfer of values was shared across the communities. both the town and township communities valued the transfer of heritage from one generation to the next despite the differences in the elements of heritage transferred. clear divisions were seen in the communities with respect to heritage conservation in the town, and the level of contribution of the various stakeholders to resource conservation efforts. conservation of heritage resources was found to be focused in the town area with conservation organisations and members of these organisations primarily based in this area. this created a disconnect between township stakeholders and current heritage resource conservation, perpetuating the unequal participation of township stakeholders in conservation efforts by organisations. this supports findings by chirikure et al. (2010), stating that local communities are not active in conservation efforts. a situation where stakeholders are spectators of resource conservation, where awareness of conservation efforts taking place exists at all, is created (ndoro, mumma & abungu 2008). this means that stakeholders do not participate in conservation or the decisions surrounding conservation and rather stand back and watch whilst managers or other small groups of authority make important conservation-related decisions and implement these independent of stakeholders. the unequal participation highlighted in the township negatively impacts the move towards achieving sustainability. sustainable communities require greater integration in heritage conservation and decision-making in this regard, in order to ensure that efforts taken are effective and far-reaching. caffyn and jobbins (2003) also mention the importance of incorporating stakeholders in the establishment of plans before plan implementation. this emphasises the importance of stakeholder participation at all stages of decision-making and management plan development. greater participation, and therefore, possibly greater conservation of heritage resources would result in better conservation of both natural and cultural heritage types, ensuring their sustainability over time. the involvement of township communities would encourage a more cohesive community approach to conservation, and possibly effect more conscious and careful interactions of township residents with resources such as the natural environment, also promoting sustainability in the future of wakkerstroom. stakeholder perceptions in this study highlighted a lack of communication amongst the two communities. the people in the town valued written communication, whereas the people in the township preferred oral communication in their traditional languages. communication is an important step towards inclusiveness, sustainable development and effective heritage resource conservation (aas et al. 2005). a lack of communication has been linked to hampered development in tourism and other developmental activities where towns are concerned (peters 1999). in order to ensure progression towards a sustainable town where communication is concerned, it is vital that the language differences amongst the two stakeholder communities are considered. the creation of awareness, possibly through the education system, and incorporation of more languages in information communication and transfer (i.e. more of the local languages including isizulu and siswati) where heritage conservation is concerned is vital for the success of conservation efforts and effective management. effective communication of heritage conservation-related plans and conservation issues in both the township and town area would facilitate greater community cohesion and facilitate better participatory engagement, contributing to the aim of a sustainable town, and sustainable development (keitumetse 2011). a balance between the need for development in the town and the need for heritage conservation is also highlighted by threats such as those of mining mentioned by stakeholders. whilst some township stakeholders viewed mining as a source of much-needed job creation and development, concerns of damage to the natural environment including the wetland were mentioned by other stakeholders. this creates a conflict between development and heritage protection and presents a challenge to management (chirikure 2013; chirikure et al. 2008). local economic development is mentioned in the integrated development plan, including the prospect of mining the natural resources (mainly coal) in the district and using this as a source of economic development (dpkis local municipality idp 2017). the idp recognises under development principle 1 the need to achieve a balance between areas of environmental conservation and activities such as mining and tourism (dpkis local municipality 2017). this idp should be reviewed considering the information collected in this study. discussions are ongoing with the municipality. future participation of communities is important in heritage management and the success of conservation efforts (yung & chan 2011). through greater participation and inclusion of township stakeholders in conservation actions in the town, the needs of various stakeholders, the differences in values and heritage definitions can be addressed and incorporated into future conservation plans (richards, carter & sherlock 2004). incorporating the perceptions of local community stakeholders is important for successful efforts towards sustainable development and sustainable communities to be implemented (gursoy, jurowski & uysal 2002). explicit presentation of benefits of conservation to stakeholders could result in a greater appreciation and participation in conservation efforts. the value of public education has been recognised and should be incorporated in this regard (azman et al. 2010). education influences the behaviour of people and actions taken towards conservation (azman, halim & komoo 2009). awareness and understanding of heritage have been recognised as important factors to effective conservation, as these facilitate value appreciation. consideration and actions towards the declaration of wakkerstroom as a heritage town could also be undertaken. heritage towns are places of great historical value, with multiple historic buildings and areas, and historical events associated with the town. according to unesco, a heritage town is characterised by continued use of the town for living purposes, and subsequently, the maintenance of resources, preventing deterioration (schröder-esch 2006). classification of wakkerstroom as a heritage town will lead to a greater pool of resources becoming available for the conservation of not only the natural heritage in the town, but also the cultural heritage including the town buildings which are national monuments. heritage management plans are an important tool and essential for effective conservation. in order to achieve sustainable towns, it is important that all stakeholders take responsibility. the municipality needs to take the leadership in developing the management plan for both natural and cultural heritage conservation, allocating roles for effective implementation. conclusion the wakkerstroom area has diverse natural and cultural heritage resources including an internationally recognised wetland with high bird, mammal and reptile diversity as well as it being one of the oldest towns in the region, established in 1859. these attributes suggest that the concept of conservation should embrace both the natural heritage as well as the architectural and cultural heritage, in the same location. this is the trend in many parts of the world, and the results provide an interesting test case as to whether a single location could be used to show the benefit of an integrated management plan for the conservation of both the natural and cultural heritage resources. this study provides some preliminary information which would be needed to develop such a plan, which does seem feasible. the resources are relatively well-defined, which is a first and very important step. there are three buildings that are listed on the south african monuments register. the study also highlighted the opportunities and limitations which would need to be included in the development of a management plan. these included value definitions, stakeholder participation in decision-making and engagement in activities, styles of communication, language and awareness, trade-offs needed to achieve sustainable development. this study also clearly highlighted the fact that the responsibility for the conservation of these resources is vested in the authoritative power of the elected municipality, but they have abrogated this responsibility. interestingly this responsibility has been adopted by ngos. these bodies should work together, using the findings of this study, to turn wakkerstroom into a ‘heritage town’ with a sustainable development plan that includes the conservation of both the natural and cultural heritage resources. this would lead to the overall uplift of the community. acknowledgements the participants of the town and township are thanked for their contributions and hospitality. competing interests the authors have declared that no competing interests exist. authors’ contributions m.s. conceived and helped develop the original project. r.m.h.s. carried out the research required on the project under the supervision of m.s. both authors discussed the results and contributed to the final article. ethical consideration ethical clearance for the study was issued by the university of the witwatersrand (protocol number: ha1905). informed consent forms were also provided for all participants of the study. funding information the project was funded by the national research foundation and the department of science and technology, through the funding of prof scholes’ sarchi chair in global change and systems analysis (grant number 101057). data availability statement the data provided in the paper may be used but the raw data cannot be provided as that is protected by the ethics rules of the university. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references aas, c., ladkin, a. & fletcher, j., 2005, ‘stakeholder collaboration and heritage management’, annals of tourism research 32(1), 28–48. https://doi.org/10.1016/j.annals.2004.04.005 azman, n., halim, s. & komoo, i., 2009, ‘integrated public education for heritage conservation: a case for langkawi global geopark’, in g. ainsworth & s. garnett (eds.), dalam rimba: sustainable forest livelihoods in malaysia and australia, pp. 151–156, universiti kebangsaan malaysia, bangi malaysia. azman, n., halim, s., liu, o., saidin, s. & komoo, i., 2010, ‘public education in heritage conservation for geopark community’, procedia-social and behavioral sciences 7, 504–511. https://doi.org/10.1016/j.sbspro.2010.10.068 bakri, a., ibrahim, n., ahmad, s. & zaman, n., 2015, ‘public perception on the cultural significance of heritage buildings in kuala lumpur’, procedia – social and behavioral sciences 202, 294–302. https://doi.org/10.1016/j.sbspro.2015.08.233 begg, g., 1989, the wetlands of natal: the distribution, extent and status of wetlands in the mfolozi catchment, 68, natal town and regional planning commission, pietermaritzburg. beierle, t. & konisky, d., 2001, ‘what are we gaining from stakeholder involvement? observations from environmental planning in the great lakes’, environment, planning and conservation 19(4), 515–527. https://doi.org/10.1068/c5s blundo, d., ferrari, a., del hoyo, a., riccardi, m. & muina, f., 2018, ‘improving sustainable cultural heritage restoration work through life cycle assessment-based model’, journal of cultural heritage 32, 221–231. https://doi.org/10.1016/j.culher.2018.01.008 caffyn, a. & jobbins, g., 2003, governance capacity and stakeholder interactions in the development and management of coastal tourism: examples from morocco and tunisia, journal of sustainable tourism 11(2–3), 224–245. https://doi.org/10.1080/09669580308667204 carter, r. & bramley, r., 2002, ‘defining heritage values and significance for improved resource management: an application to australian tourism’, international journal of heritage studies 8(3), 175–199. https://doi.org/10.1080/13527250220000/18895 chirikure, s., 2013, ‘heritage conservation in africa: the good, the bad, and the challenges’, south african journal of science 109(1–2), 1–3. chirikure, s., manyanga, m., ndoro, w. & pwiti, g., 2010, ‘unfulfilled promises? heritage management and community participation at some of africa’s cultural heritage sites’, international journal of heritage studies 16(1–2), 30–44. https://doi.org/10.1080/13527250903441739 chirikure, s., pwiti, g., damm, c., folorunso, c.a., hughes, d.m., phillips, c. et al., 2008, ‘community involvement in archaeology and cultural heritage management: an assessment from case studies in southern africa and elsewhere’, current anthropology 49(3), 467–485. civallero, e., 2007, ‘traditional games, music and oral tradition: intangible tools in multicultural libraries’, in ifla satellite meeting 2007, conference on innovative multicultural library services for all, national university of cordoba, pretoria, south africa, august 15–17, 2007, pp. 15–17. cleere, h., 2010, ‘management plans for archaeological sites: a world heritage template’, conservation and management of archaeological sites 12(1), 4–12. https://doi.org/10.1179/175355210x12791900195025 deacon, h. & smeets, r., 2013, ‘authenticity, value and community involvement in heritage management under the world heritage and intangible heritage conventions’, heritage & society 6(2), 129–143. https://doi.org/10.1179/2159032x13z.0000000009 de medici, s. & senia, c., 2007, ‘sustainable requalification of architectural and natural resources: the coastal village of marzamemi’, ecosystems and sustainable development 4(6), 175. díaz-andreu, m., 2017, ‘heritage values and the public’, journal of community archaeology & heritage 4(1), 2–6. https://doi.org/10.1080/20518196.2016.1228213 dr pixley ka isaka seme local municipality, 2011, idp, dr pixley ka isaka seme local municipality, volksrust, mpumalanga. dr pixley ka isaka seme local municipality, 2017, dr pixley ka isaka seme local municipality final 2017–2022 integrated development plan (idp), pp. 2–188, dr pixley ka isaka seme local municipality, volksrust, mpumalanga. gardiner, n., 2017, a walk through history in wakkerstroom by: nicolette gardiner – wakkerstroom. wakkerstroom official website, viewed 26 october 2020, from https://www.wakkerstroom.co.za/walk-history-wakkerstroom-nicolette-gardiner/. gursoy, d., jurowski, c. & uysal, m., 2002, ‘resident attitudes: a structural modeling approach’, annals of tourism research 29(1), 79–105. https://doi.org/10.1016/s0160-7383(01)00028-7 hare, m. & pahl-wostl, c., 2002, ‘stakeholder categorisation in participatory integrated assessment processes’, integrated assessment 3(1), 50–62. harun, s., 2011, ‘heritage building conservation in malaysia: experience and challenges’, procedia engineering 20, 41–53. https://doi.org/10.1016/j.proeng.2011.11.137 hofmeyer, h., smith, k. & smith, c., 2009, wakkerstroom jewel of mpumalanga, pp. 1–191, 1st edn., mediakor, johannesburg. hribar, m., bole, d. & pipan, p., 2015, ‘sustainable heritage management: social, economic and other potentials of culture in local development’, social and behavioral sciences 188, 103–110. https://doi.org/10.1016/j.sbspro.2015.03.344 jaafar, m., noor, s. & rasoolimanesh, s., 2015, ‘perception of young local residents toward sustainable conservation programmes: a case study of the lenggong world cultural heritage site’, tourism management 48, 154–163. https://doi.org/10.1016/j.tourman.2014.10.018 joubert, r. & ellery, w., 2013, controls on the formation of wakkerstroom vlei, mpumalanga province, south africa’, african journal of aquatic science 38(2), 135–151. https://doi.org/10.2989/16085914.2012.762897 jürgens, u., donaldson, r., rule, s. & bähr, j., 2013, ‘townships in south african cities–literature review and research perspectives’, habitat international 39, 256–260. https://doi.org/10.1016/j.habitatint.2012.10.011 kalichman, s. & simbayi, l., 2003, ‘hiv testing attitudes, aids stigma, and voluntary hiv counselling and testing in a black township in cape town, south africa’, sexually transmitted infections 79(6), 442–447. https://doi.org/10.1136/sti.79.6.442 keitumetse, s., 2011, ‘sustainable development and cultural heritage management in botswana: towards sustainable communities’, sustainable development 19(1), 49–59. https://doi.org/10.1002/sd.419 khan, f., 2010, ‘verlorenvlei: where past and present collide’, veld & flora 96(4), 182–183. kirshenblatt-gimblett, b., 2004, ‘intangible heritage as metacultural production’, museum international 56(1–2), 52–65. https://doi.org/10.1111/j.1350-0775.2004.00458.x kotze, d., 2013, ‘a system for supporting wetland management decisions’, doctoral dissertation, university of natal, pietermaritzurg. kotze, d., breen, c. & klug, j., 1994, ‘a management plan for wakkerstroom vlei’, in r.g. oellermann, m.a.g. darroch, j.r. klug & d.c. kotze (eds.), wetland preservation valuation, and management practices applied to wetlands: south african case studies. wrc report no. 501/5/94, water research commission, pretoria. lapadula, m. & quiroga, c., 2012, ‘heritage as a pedagogical resource and platform for exploration in architectural design education’, the journal of architecture 17(4), 591–607. https://doi.org/10.1080/13602365.2012.709028 lowenthal, d., 2005, ‘natural and cultural heritage’, international journal of heritage studies 11(1), 81–92. https://doi.org/10.1080/13527250500037088 municipal money, 2020, municipal money profile: dr pixley ka isaka seme, viewed 05 september 2020, from https://municipalmoney.gov.za/profiles/municipality-mp304-dr-pixley-ka-isaka-seme/. munton, r., 2003, ‘deliberative democracy and environmental decision-making’, in f. berkhout, m. leach & i. scoones (eds.), negotiating change: advances in environmental social science, pp. 109–136, edward elgar, cheltenham. ndoro, w., mumma, a. & abungu, g., 2008, cultural heritage and the law: protecting immovable cultural heritage in english speaking sub-saharan africa, iccrom studies in conservation, 8, pp. 1–129, international center for the study of the preservation and restoration of cultural property, rome italy. nzama, a., 2009, ‘the nexus between sustainable livelihoods and ecological management of the world heritage sites: lessons from isimangaliso world heritage park, south africa’, inkanyiso: journal of humanities and social sciences, 1(1), 34–42. https://doi.org/10.4314/ijhss.v1i1.62107 oellermann, r., darroch, m. & klug, j., 1994, ‘valuing preferences for wetland preservation: a wakkerstroom case study’, african journal of range & forage science 11(3), 89–95. https://doi.org/10.1080/10220119.1994.9647853 parts, p., rennu, m., jääts, l., matsin, a. & metslang, j., 2011, ‘developing sustainable heritage-based livelihoods: an initial study of artisans and their crafts in viljandi county, estonia’, international journal of heritage studies 17(5), 401–425. https://doi.org/10.1080/13527258.2011.589199 patel, z., greyling, s., simon, d., arfvidsson, h., moodley, n., primo, n. et al., 2017, ‘local responses to global sustainability agendas: learning from experimenting with the urban sustainable development goal in cape town’, sustainability science 12(5), 785–797. https://doi.org/10.1007/s11625-017-0500-y pereira, d. & marker, b., 2016, ‘the value of original natural stone in the context of architectural heritage’, geosciences 6(1), 13. https://doi.org/10.3390/geosciences6010013 peters, h., 1999, ‘making tourism work for heritage preservation: lijiang, a case study’, international conference on anthropology, chinese society and tourism, at yunnan university, kunming, yunnan from, september, 1999, p. 28. peters, w., 1995, wakkerstroom: a conservation study, 1st edn., pp. 1–175, department of architecture, university of natal, durban. peters, w., vicari, p., nortjie, s., hewitt, g., garach, h., naroth, m. et al., 2001, wakkerstroom: a re-assessment in urban conservation, 1st edn., pp. 1–131, school of architecture, university of natal, durban. rahman, s., 2013, ‘heritage management challenges in historic town of ludlow, england’, world applied sciences journal 24(12), 1589–1596. rautenbach, c., hart, d. & naudé, m., 2015, ‘heritage resources management’, in a. du plessis (ed.), environmental law and local government in south africa, pp. 853–885, juta and company, potchefstroom. reid, w., mooney, h., cropper, a., capistrano, d., carpenter, s., chopra, k. et al., 2005, ecosystems and human well-being-synthesis: a report of the millennium ecosystem assessment, island press, washington d.c. renn, o., 2006, ‘participatory processes for designing environmental policies’, land use policy 23, 34–43. https://doi.org/10.1016/j.landusepol.2004.08.005 richards, c., carter, c. & sherlock, k., 2004, practical approaches to participation, macaulay institute, aberdeen. sadler, g., lee, h., lim, r. & fullerton, j., 2010, ‘recruitment of hard-to-reach population subgroups via adaptations of the snowball sampling strategy, nursing & health sciences 12(3), 369–374. https://doi.org/10.1111/j.1442-2018.2010.00541.x sahra, 2020, sahris.sahra.org.za. 2020, declared sites | sahra, viewed 26 october 2020, from https://sahris.sahra.org.za/declaredsites. salo, e., 2003, ‘negotiating gender and personhood in the new south africa: adolescent women and gangsters in manenberg township on the cape flats’, european journal of cultural studies 6(3), 345–365. https://doi.org/10.1177/13675494030063005 sanbi, 2020a, plants of southern africa: floristic information. viewed 3 december 2020, from http://newposa.sanbi.org/sanbi/flora. sanbi, 2020b, geocortex viewer for html5, viewed 17 august 2020, from https://bgisviewer.sanbi.org/html5viewer/index.html?configbase=http://bgisviewer.sanbi.org/geocortex/essentials/rest/sites/mbcp_themes/viewers/mbsp/virtualdirectory/resources/. schröder-esch, s., 2006, practical aspects of cultural heritage: presentation, revaluation, development; contributions presented on the 28th and 29th october 2005 as part of the 2nd hermes symposium in krakow, poland, bauhaus-university. south african heritage resources information systems, n.d., old apostolic church of africa, church street, wakkerstroom | sahra, viewed 21 february 2020, from https://sahris.sahra.org.za/node/33735. statssa, 2011, main place: wakkerstroom, viewed 25 october 2020, from https://www.statssa.gov.za/?page_id=4286&id=11493. stubbs, j., 2009, time honored: a global view of architectural conservation, john wiley & sons, hoboken new jersey. su, m., sun, y., min, q. & jiao, w., 2018, ‘a community livelihood approach to agricultural heritage system conservation and tourism development: xuanhua grape garden urban agricultural heritage site, hebei province of china’, sustainability 10(2), 361. sultana, p. & abeyasekera, s., 2008, ‘effectiveness of participatory planning for community management of fisheries in bangladesh’, journal of environmental management 86, 201–213. https://doi.org/10.1016/j.jenvman.2006.12.027 tarboton, w., 2004, wakkerstroom bird & nature guide, s.n., wakkerstroom. theal, g., 2010, history of south africa since september 1795, cambridge university press, cambridge, england. the burra charter, 1999, the australia icomos charter for places of cultural significance, australia icomos, burwood. unced, 1992, agenda 21: programme for action for sustainable development, united nations conference on the environment and development, rio de janeiro. unesco, n.d., south africa – unesco world heritage centre, viewed 25 october 2020, from https://whc.unesco.org/en/statesparties/za. united nations educational, scientific and cultural organisation (unesco), 1989, legal instruments. recommendation on the safeguarding of traditional culture and folklore, paris, 25th general conference, viewed 25 october 2020, from https://portal.unesco.org/en/ev.php-url_id=13141&url_do=do_topic&url_section=201.html. varvasovszky, z. & brugha, r., 2000, ‘how to do (or not to do). a stakeholder analysis’, health, policy and planning 15(3), 338–345. https://doi.org/10.1093/heapol/15.3.338 wakkerstroom bird club, 2016, newsletter, 44, viewed 03 september 2020, from https://www.wakkerstroom.co.za/wp-content/uploads/2014/10/newsletter-no-44.pdf. wood, r., liang, h., wu, h., middelkoop, k., oni, t., rangaka, m. et al., 2010, ‘changing prevalence of tuberculosis infection with increasing age in high-burden townships in south africa’, the international journal of tuberculosis and lung disease 14(4), 406–412. young, j., jordon, a., searle, r., butler, a., chapman, d., simmons, p. et al., 2013, ‘does stakeholder involvement really benefit biodiversity conservation?’, biological conservation 158, 59–70. https://doi.org/10.1016/j.biocon.2012.08.018 yung, e. & chan, e., 2011, ‘problem issues of public participation in built-heritage conservation: two controversial cases in hong kong’, habitat international 35(3), 457–466. https://doi.org/10.1016/j.habitatint.2010.12.004 zhang, y., lu, d., yang, b., sun, c. & sun, m., 2011, ‘coastal wetland vegetation classification with a landsat thematic mapper image’, international journal of remote sensing 32(2), 545–561. https://doi.org/10.1080/01431160903475241 filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 venter.qxd short-term vegetation change on rehabilitated peatland on rietvlei nature reserve c.e. venter, g.j. bredenkamp and p-l. grundlingh venter, c.e., g.j. bredenkamp and p-l. grundlingh. 2003. short-term vegetation change on rehabilitated peatland on rietvlei nature reserve. koedoe 46(1): 53–63. pretoria. issn 0075-6458. natural peatlands occur on the rietvlei nature reserve. before the pretoria city council acquired the land, these peatlands were mined by private land-owners. ditches were constructed to drain the area for mining and the peatlands became desiccated. later the area was proclaimed as a nature reserve and has since then been managed as such. rehabilitation of the drained peatland on rietvlei nature reserve first started in 2000 as a working for water project. the aim of the rehabilitation was to close the ditches and rewet the peatland, to enable possible revival of the peatland. a baseline vegetation survey was undertaken during the summer (march to april) of 2001 to determine the nature of the pioneer communities that established on the rehabilitated area. this survey was repeated during the summer (march to april) of 2002 to detect changes in the vegetation. the same sample plots were used on both occasions. the initial pioneer vegetation was mostly composed of weedy annuals. three communities were identified from the 2001 data set using the classification algorithm twinspan. decorana ordination showed a clear distinction between the community on the mined and the communities on the drained areas. a specific pioneer community dominated by persicaria species established on the rehabilitated ditches. phragmites australis reed communities, with phytolacca octandra, were prominent on the least disturbed sites. cyperus dominated communities, without reeds (phragmites australis), occurred on areas that were mined extensively, but not rehabilitated. the 2002 floristic data was added to the baseline data and the ordination procedure as performed on the 2001 data set was repeated on this new data set. the results indicate that within a single year the vegetation already started to change in the direction of the climax communities. key words: vegetation survey; peatland; succession; rehabilitation. c.e. venter, g.j. bredenkamp and p-l. grundlingh, african vegetation and plant diversity research center, department of botany, university of pretoria, pretoria, 0002 republic of south africa. issn 0075-6458 53 koedoe 46/1 (2003) introduction wetlands are very important in nature, because they form unique habitats for certain aquatic and hydrophilic plant species and also result in an increase in the number of waterfowl and other fauna typically associated with wetlands. wetlands lessen the devastating effects of floods and are responsible for clearer and healthier surface water. they act as a filter for debris and certain toxic substances, such as heavy metals, and absorb large quantities of water that are eventually released over time into natural drainage channels (hey & philippi 1999). wetlands in south africa do not have high plant species diversity but some of the species that do occur in wetlands are restricted to these habitats (bloem et al. 1993; myburgh et al. 1995; eckhardt et al. 1993; smit et al. 1995). additional to these functional and biological values, wetlands also have consumptive uses, e.g. they provide water to man and animals, they generate energy and can be used for fish farming. non-consumptive uses include recreation, tourism and transport (rand water scientific services 1998). venter.qxd 2005/12/09 10:34 page 53 the goal of a rehabilitation project may be to a particular species, or may include the rehabilitaion of a whole ecosystem or landscape, or to restore the ecosystem functions (ehrenfeld 2000). south africa ratified the ramsar convention (cowan & marneweck 1996) and are therefore under an obligation to conserve wetlands. the rehabilitation of wetlands play an important role in wetland conservation, as mentioned in the ramsar convention. when rehabilitating wetlands it is important to understand wetland functions and to give the system time to react. some rehabilitation projects may take up to 50 years before rehabilitation has been accomplished (middleton 1999; mitsch & wilson 1996). a wetland ecosystem is not a stable system, because many disturbances such as flood pulsing, fire and herbivory occur naturally in many wetlands (middleton 1999; mitsch & wilson 1996), especially in a country such as south africa with its high frequency of thunder storms, fire-prone grasslands and large variety of large herbivores. a restored wetland should be able to deal with these kinds of disturbances. only a few peatlands have been rehabilitated in south africa and information is limited. in 1995 it has been proposed that the orange river mouth salt marsh should be rehabilitated (marneweck 1995), but in 1998 almost no progress has been made and many problems were experienced with the project (dini 1998). another similar project was the rehabilitation of blesbokspruit (dini 1998). in 1996 the free state department of environmental affairs and tourism and rand water started the rehabilitation of the seekoeivlei as well as other wetlands in the northeastern free state (collins 1998). the rehabilitation of the wilge river started as a rand water and working for water project in the same year (rand water & working for water programme 1998). these projects take time and it is therefore difficult to determine if the projects have been successful. the orange river mouth, blesbokspruit and seekoeivlei are ramsar sites (cowan & marneweck 1996). on the contrary, there is much information available on the rehabilitation of european peatlands, but those are completely different ecosystems, which developed under very different climatic conditions. during peat mining operations in the past on rietvlei nature reserve, ditches with sharp edges were constructed to drain the peatland. some of the water was used for irrigation of farms in the area. there are four dolomite springs on the reserve. the water from these springs is piped to the rietvlei water works. a large amount of water was therefore drained from the peatland, which subsequently became dry and resulted in the vegetation being altered. rehabilitation of the rietvlei peatland started as a "working for water" project. the aim of the rehabilitation project was mainly to rewet the peatland to allow hydrophytic vegetation to re-establish. the sharp edges of the main channel were sloped and gabions were placed in the main channel to stop the peat from rapidly drying out and to allow floodwater to easily overspill onto the adjacent peatland, thus enhancing the rewetting process. the peat recovered from the sloping process was used to fill some of the smaller ditches. soil and straw gabions were placed in the ditches to stabilise them and to retain the water. rock filled gabions were placed in the main channel to slow the flow, increase the flooding frequency and duration and to promote the deposition of sediment. it was necessary to conduct a baseline vegetation survey to determine the pioneer plant communities that established naturally on the rehabilitated peatland. the communities closer to the climax, that occur on areas with less disturbance, were also surveyed, to gain knowledge on the potential climax vegetation. study area the rietvlei dam nature reserve is located at about 25º50's and 28º20'e. the reserve became established as a result of the rietvlei water scheme to provide drinking water to the people of pretoria. the rietvlei dam is koedoe 46/1 (2003) 54 issn 0075-6458 venter.qxd 2005/12/09 10:34 page 54 located in the northern portion of the reserve and the peatland is located upstream from the dam in the southern portion of the reserve. the city council of pretoria acquired the farm in 1929. the dam was built in the ses myl spruit during the great depression and it was completed in 1934. the catchment area of the ses myl spruit is to the south of pretoria, towards kempton park. the area was declared the rietvlei game reserve in 1937. in 1948 it became the rietvlei reserve for game and indigenous flora. in 1954 the area became the van riebeeck nature reserve and in 1992 the name was changed again to the rietvlei nature reserve (culture and recreation department 1997). to drain the peatland, a deep ditch was dug through it, and this now forms part of the ses myl spruit main channel. the main channel has been severely eroded and the water flows very fast, with high energy levels. the main channel and the peatland on the western side of the main channel have been rehabilitated. certain portions of the peatland, on the eastern side of the main channel, were extensively mined and almost all the peat was removed. this area has not been rehabilitated. the underlying clay soil was exposed. on the western side of the main channel deep deposits of peat still remained intact, but they were severely disturbed by drainage ditches. the vegetation was disturbed during the construction of the ditches and the ditches altered the hydrological regime of the peatland. methods the peatland area was stratified into the following units: ditches filled with peat; ditches filled with soil; sloped main channel edges; reed areas on old peatdeposits; and the mined area east of the main channel. during the summer of 2001, 22 sample plots of 4 m x 4 m were placed randomly in these different units. the exact locality of each plot was recorded using a global positioning system (gps). in each plot all plant species present were listed and each assigned the approriate braun-blanquet cover-abundance value (werger 1974). some aspects of the habitat were also noted such as soil type (peat or clay), degree of disturbance and rehabilitation measures. the floristic data were entered into the turboveg data basis (hennekens 1996a) and the data were subjected to classification and ordination procedures to identify plant communities. twinspan (hill 1979b) was used for the initial classification of the vegetation, and the result was refined by using megatab (hennekens 1996b). the final classification is presented in a phytosociological table. the procedure of detrended correspondence analysis was performed by applying the decorana software (hill 1979a). this ordination was used to indentify environmental gradients responsible for plant species distribution, and hence the occurrence of plant communities. to determine a probable change in vegetation (path of succession) after one year, the floristic survey was repeated during the summer of 2002 on the same sample plots as 2001. the data of the two surveys were pooled and to determine the direction of change in vegetation, the ordination was re-applied to the pooled data set. results classification the result of the classification is presented in the phytosociological table (table 1). the classification of the plant communities found in the baseline survey (2001 data set) is as follows: 1. amaranthus hibridus setaria ustilata community on filled ditches. 1.1. cynodon dactylon solanum sisymbrifolium sub-community on soil filled ditch. 1.2. persicaria species amaranthus hibridus sub-community on peat filled ditch. 2. phytolacca octandra solanum nigrum wetland community. 2.1. phytolacca octandra datura stramonium disturbed wetland sub-community. issn 0075-6458 55 koedoe 46/1 (2003) venter.qxd 2005/12/09 10:34 page 55 koedoe 46/1 (2003) 56 issn 0075-6458 table 1 phytosociological table of pioneer vegetation on the rehabilitated peatland at rietvlei nature reserve venter.qxd 2005/12/09 10:34 page 56 issn 0075-6458 57 koedoe 46/1 (2003) table 1 (continued) the following species were encountered in the peatland, but were not of any significant value to the phytosociological table hyparrhenia hirta setaria verticillata alisma plantago-aquatica schoenoplectus corymbosus bidens formosa panicum schinzii hemarthria altissima digitaria sanguinalis askidiosperma albo-aristatum ranunculus multifidus cyperus eragrostis sambucus nigra helichrysum species trifolium africanum sisymbrium thellungii pseudognaphalium species xanthium strumarium 2.2. phragmites australis chenopodium album wetland sub-community. 3. cyperus species oenothera rosea wetland community on clay. 3.1. cyperus species paspalum dilatatum moist wetland sub-community on clay. 3.2. typha capensis cyperus fastigiatus wet wetland sub-community on clay. a hierarchical and habitat interpretation of the recognised plant communities is given in fig. 1. community description 1. amaranthus hibridus setaria ustilata community on filled ditches. the plant community occurs on the ditches that were filled with peat or soil during the rehabilitation of the peatland. the vegetation represents a pioneer stage, which established directly after the new disturbance caused by the rehabilitation. the vegetation is short, mostly below 0.5 m, and young. some parts of the community have a dense vegetation (aerial) cover (80 %) while other parts have open patches with bare peat. the community is characterised by species group a. persicaria species is dominant and amarathus hibridus and setaria ustilata are conspicuously present. all these species are annual weeds, common in disturbed areas and are indicators of disturbance. a total of 38 species were noted in the community and an average of 14 species per relevé. 1.1. cynodon dactylon solanum sisymbrifolium sub-community on soil filled ditches. the sub-community is located on soil (not peat) filled ditches at the edge of the wetland. this habitat is much drier that the rest of the wetland. the vegetation is open (about 30 % aerial cover) with large patches of bare soil surface visible. this sub-community is characterised by species group b. the stoloniferous perennial grass cynodon dactylon is the dominant species and the annual weeds solanum sisymbrifolium and datura stramonium are conspicuous in this sub-community. both these weeds occur in disturbed areas. cynodon dactylon occurs in disturbed places on all soil types and is often found in damp areas (van oudtshoorn 1999). an average of 15 species per relevé and a total of 15 species were found in this subcommunity. 1.2. persicaria species amaranthus hibridus sub-community on peat filled ditches. this sub-community is located on old drainage ditches filled with peat during the rehabilitation. the aerial cover of the vegation is about 50 %. species group c characterises this sub-community and the dominant species is persicaria. furthermore amaranthus hibridus, persicaria lapathifolia and cyperus fastigiatus are also conspicuous. phragmites australis is starting to establish within this sub-community. persicaria lapventer.qxd 2005/12/09 10:34 page 57 athifolia is a weed that often occurs in wet areas and amarathus hibridus is common in disturbed areas (van wyk & malan 1988). a total of 32 species were noted in this subcommunity with an average of 14 species per relevé. 2. phytolacca octandra solanum nigrum wetland community. this is a weedy community on peat, probably representing a further stage in the succession. it is closer to the main channel and is moister than community 1. the community is characterised by species group d and shares species group f with community 1. the dominant species is phytolacca octandra, which is a tall-growing annual weed (>2 m) restricted to disturbed areas (van wyk & malan 1988). the community has a species diversity of 26 and an average of nine species per relevé. 2.1. phytolacca octandra datura stramonium disturbed wetland subcommunity. the sub-community has been disturbed and the vegetation consists mainly of weeds. the vegetation is dense, with an aerial cover of almost 100 % and up to 2.5 m tall. this subcommunity is characterised by the absence of species group e. the dominant species are phytolacca octandra and datura stramonium, while persicaria lapathifolia and solanum nigrum are also very conspicuous. phytolacca octandra is a weed restricted to disturbed areas and persicaria lapathifolia is a weed that often occurs in wet areas (van wyk & malan 1988). the sub-community with 13 species is relatively poor in species koedoe 46/1 (2003) 58 issn 0075-6458 subcommunity 1.1. 1 relevé subcommunity 1.2 6 relevés subcommunity 2.1 3 relevés subcommunity 2.2 6 relevés clay soil peat weed dominated phragmites dominated rietvlei rehabilitated peatlands filled ditches disturbed area drier area wetter area rehabilitated drained area unrehabilitated mined area subcommunity 3.1 3 relevés subcommunity 3.2 3 relevés community 1 community 2 community 3 fig. 1. dendrogram showing the hierarchy of twinspan divisions for the vegetation data of the rehabilitated peatlands on rietvlei nature reserve on the 2001 dataset. venter.qxd 2005/12/09 10:34 page 58 composition, probably due to the shading effect of the dominant species. seven species on average were found in each relevé. 2.2. phragmites australis chenopodium album wetland sub-community the sub-community represents a further successional stage, probably close to the reed dominated climax. it is located close to the main channel and is quite wet. species group e characterises the sub-community. phragmites australis is the dominant species and the weedy annuals chenopodium album and solanum nigrum are often present. phytolacca octandra is present, but is relatively small in size in this sub-community, probably due to the competition from phragmites australis. a total of 23 species were noted in this sub-community with an average of nine species per relevé. 3. cyperus species oenothera rosea wetland community on clay. this community is restricted to the eastern side of the main channel. the community is on a part of the wetland where the peat was mined extensively, and where the mined area is relatively flat without any drainage ditches. all peat was removed and the community is therefore on a clay soil. this area was not actively rehabilitated. species group h characterises the community and the dominant species is cyperus species. oenothera rosea is a weed that occurs in moist disturbed areas and is an indicator of the past disturbance that occurred at the site (van wyk & malan 1988). only 14 species were recorded in this community with an average of six species per relevé. 3.1. cyperus species paspalum dilatatum moist wetland sub-community on clay. this sub-community is drier than sub-community 3.2, because is is on slightly higher ground, and the vegetation is shorter, mostly lower than 1 m. the sub-community is characterised by the absence of species group i and it is dominated by cyperus species. oenothera rosea and cirsium vulgare are conspicuous in the vegetation. paspalum dilatatum occurs mostly on clay and loam soil and in wet areas and is often a weed in disturbed areas (van oudtshoorn 1999). cirsium vulgare also occurs in moist disturbed areas (van wyk & malan 1988). these species are indicators of the past disturbance, as well as the moist clayey soil. the subcommunity has an average of six species in each relevé and a species diversity of only nine species. this is the sub-community with the lowest species diversity. 3.2. typha capensis cyperus fastigiatus wet wetland sub-community on clay. the sub-community occurs on moist clay soil and is located in a lower area of the wetland. the vegetation is about 2 m tall. species group i is characteristic of the subcommunity and the dominant species is the perennial typha capensis. epilobium hirsutum, persicaria lapathifolia, cyperus sp., cyperus fastigiatus and oenothera rosea are conspicuously present. typha capensis occurs in wet areas and epilobium hirsutum occur in moist grassland and are often found in vlei areas (van wyk & malan 1988). these species are indicators of the wet soil. the sub-community has 10 species and an average of seven species per relevé. ordination in the ordination on the 2001 data set (fig. 2), the separation of communities 1 and 2 on the drained, rehabilitated side of the channel and community 3 on the mined side of the channel is clear. most of the relevès in a sub-community are close together but there are a few exeptions where the relevès are in a state of transition between two communities. although relevè 2 is still in sub-community 1.2 it is also close to sub-community 2.2. it is therefore closer to the climax (subcommunity 2.2) than the other relevès in sub-community 1.2. the environmental gradient from dry to wet can be seen in the ordination as well as the gradients from the pioneer community to the two different closeissn 0075-6458 59 koedoe 46/1 (2003) venter.qxd 2005/12/09 10:34 page 59 to-climax communities. these communities occur in the areas with less recent disturbance and are more similar to climax vegetation than the pioneer communities. the succession of community 1 will be to either one of them. the one close-to-climax community should not change to become like the other close-to-climax community. monitoring vegetation change the following is the result of the ordination as performed on the pooled data set for the specific purpose to monitor vegetation change from 2001 to 2002. the ordination (fig. 3) indicates that the vegetation of subcommunity 1.1 moved away (to the top left in the diagram) from the other communities. this is probably because it is situated at the edge of the wetland and has more terrestrial species than wetland species and is becoming more terrestrial, like the adjacent grassland community. the vegetation of sub-community 1.2 changed (moved downwards in the diagram) in the direction of the of the reed-dominated community 2.2. this is probably an indication of succession form a pioneer community towards the climax phragmites australis dominated vegetation. the vegetation of sub-community 2.1 stayed almost the same and although reeds are present it is dominated by the tall weed phytolacca octandra. it does however share a lot of species with the reed-dominated sub-community 2.2. relevé 11 moved closer to 2.1 and it therefore seems that it has degraded, due to later disturbance by the floods. sub-community 2.2 represents the close-toclimax vegetation dominated by phragmites australis. this vegetation stayed almost the same, however, the vegetation of sample plots 14 and 15 seems to have degraded, and moved towards sub-community 1.2, which is an earlier successional stage. community 3 stayed almost the same, but the vegetation at relevé 22 moved towards the more reed-dominated group. this may be because it is the only relevé in community 3 with phragmites australis. relevé 20 in community 3 was probably more disturbed koedoe 46/1 (2003) 60 issn 0075-6458 fig. 2. scatter diagram of a decorana ordination illustrating the difference between the communities along an environmental gradient using the 2001 dataset. axis 1 was plotted against axis 2. venter.qxd 2005/12/09 10:34 page 60 than the rest, because it is close to the rehabilitation area and has been trampled. it moved closer to the rest of community 3. the two sub-communities in community 3 seem to become more similar, since it is more difficult to distinguish them from the ordination diagram. discussion the baseline survey the first division of twinspan on the 2001 baseline data separated the un-rehabilitated, mined areas and areas disturbed by the rehabilitation. the rehabilitation disturbed the sites because of trampling, the construction of gabions and the movement of peat from one site to another. the mined area is on the eastern side of the main channel and the disturbed rehabilitated area is on the western side. these communities differ largely and have few species in common. these species are in species groups g and j and include persicaria lapathifolia, tagetus minuta and cyperus fastigiatus, mostly pioneer, annual species. the decorana ordination diagram also shows significant separation between these plant communities. there are two communities on the western, rehabilitated, side of the main channel and each of these have two sub-communities. there is only one community on the eastern side, with two sub-communities. each community and subcommunity occurs on a particular habitat. community 1 is on the filled ditches that were rehabilitated recently and the weedy pioneer plant species of the two sub-communities are mostly the same. most of the pioneer species are exotic weeds. the vegetation of sub-community 1.1 differs from 1.2, due to the dominance of the perennial stoloniferous grass cynodon dactylon, while the annual weedy solanum sisymbrifolium is very conspicuous in 1.1. persicaria species and amaranthus hibridus are more dominant in sub-community 1.2 and the annual grass urochloa panicoides occurs in this sub-community. phragmites australis is becoming established in sub-community 1.2, but not in 1.1. the differences in the vegetation of the issn 0075-6458 61 koedoe 46/1 (2003) fig. 3. scatter diagram of a decorana ordination illustrating the change of the vegetation of the communities after one year using the pooled dataset of the 2001 and 2002 data. axis 1 was plotted against axis 2. venter.qxd 2005/12/09 10:34 page 61 two sub-communities are probably a result of the difference in soil. sub-community 1.1 is on relatively dry clay and sub-community 1.2 on moist peat. the two sub-communities of community 2 differ in the dominant vegetation and probably the degree of disturbance. the vegetation of sub-community 2.1 consists mainly of tall growing weeds, mostly phytolacca octandra. the vegetation of sub-community 2.2 is dominated by phragmites australis which includes the older, well developed stands, probably closer to the climax vegetation, as well as the recently sloped area next to the river. the older and pioneer sub-communities are floristically therefore combined in a single community. sub-community 2.2 occurs on both peat and clay. it seems that a community completely different from the filled ditches (community 2.1) is therefore developing on the sloped area (part of community 2.2). the difference in disturbance may be responsible for the difference in subcommunities 2.1 and 2.2. another possibility may be that they are at different successional stages, or moving towards different new climaxes. reeds were established on the sloped areas before and the reeds are growing back rapidly. this may explain the separation of community 2.2 from community 1. community 3 on the eastern side of the main channel is completely different from the communities on the western side. the area was mined and the peat removed and not rehabilitated. the community occurs on exposed clay, that was previously underlying the peat. the community that established is dominated by cyperus species. this may be an example of a climax, or close to climax, community dominated by cyperus and typha. the wetter (slightly lower) sub-community (3.2) is dominated by typha capensis and the drier sub-community (3.1) is dominated by cyperus species. there are very few general species. communities 1 and 2 share some species, but they do not share many species with community 3. community 1 has the highest species richness and community 3 has the lowest species richness. the pioneer communities therefore contribute more to species diversity than the close-to-climax communities, which are dominated by tall groups of reeds or sedges, with few other species present. community 3 is already stable and there is probably more competition between dominant species, while community 1 is a pioneer community with many weeds and open spaces for plants to establish. few individuals of these weeds occur in the established communities and they should disappear in time. in contrast, the flood regime of the wetland may cause sufficient disturbance for the weedy species to persist in the pioneer communities for some time. wetland species are adapted to these disturbances, but they need time to establish in a disturbed wetland and the flooding regime still need to stabilise. vegetation change the study was conducted over a short period with only two data sets one year apart. this is important to take into consideration, since wetlands take a long time to rehabilitate. the study is at the start of rehabilitation and it is difficult to predict what the vegetation will look like after a few years. this study will hopefully give us some indication. after the flooding disturbance at the end of the year, that destroyed most of the pioneer vegetation, the peat was wetter. this flooding was caused by the rehabilitation measures. it seems that the rehabilitation was successful and that succession is taking place towards the climax vegetation, even after only a single year. sub-community 2.1 may be a problem, because it does not seem to advance towards a reed-dominated community, but stays the same. this is a problem because this community are dominated by weeds. it would be interesting to see if this community would change over a longer period of time. further monitoring would confirm whether the changes in vegetation is of a permanent nature, or whether the vegetation is merely fluctuating due to yearly changes in the moisture regime. the moisture regime is changing due to the rehabilikoedoe 46/1 (2003) 62 issn 0075-6458 venter.qxd 2005/12/09 10:34 page 62 taion, but yearly fluctuations in the amount of rainfall also plays a role. although it seems as if the peatland is recovering the time frame of the study is to short to confirm this as a fact. other disturbances in the peatland are grazing, by the animals occuring in the reserve as well as trampling by the buffalo. any of the disturbances could have caused the degeneration of some of the relevés, for example relevé 15. acknowledgements i would like to thank the university of pretoria for funding and the use of equipment, and the management of rietvlei nature reserve for allowing the study on the reserve. references bloem, k.j., g.k. theron & n. van rooyen. 1993. wetland plant communities of the verlorenvalei nature reserve in the north-eastern sandy highveld, transvaal. south african journal of botany 59: 281-286. collins, n. 1998. joint venture implemented between free state department of environmental affairs and tourism and rand water. south african wetlands. newsletter on activities relating to the ramsar convention in south africa 9: 5. cowan, g.i. & g.c. marneweck. 1996. south african national report to the ramsar convention 1996. pretoria: department of environmental affairs and tourism. culture and recreation department. 1997. rietvlei nature reserve. pretoria: city council of pretoria. dini, j. 1998. montreux record update. south african wetlands. newsletter on activities relating to the ramsar convention in south africa 9: 8. eckhardt, h.g., n. van rooyen & g.j. bredenkamp. 1993. wetland plant communities of the vrede-memel-warden area, northeastern orange free state. navorsinge van die nasionale museum, bloemfontein 9: 245-262. ehrenfeld, j.g. 2000. defining the limits of restoration: the need for realistic goals. restoration ecology 8(1): 2-9. hennekens, s.m. 1996a. turbo(veg): software package for input, processing, and presentation of phytosociological data. user's guide. version july 1996. wageningen: ibn-dlo. hennekens, s.m. 1996b. megatab: a visual editor for phytosociological data. user's guide. version october 1996. wageningen: ibn-dlo. hey, d.l. & n.s. philippi. 1999. a case for wetland restoration. new york: john wiley. hill, m.o. 1979a. decorana a fortran program for detrended correspondence analysis and reciprocal averaging. ithaca, new york: cornell university. hill, m.o. 1979b. twinspan a fortran program for arranging multivariate data in a ordered two-way table by classification of individuals and attributes. ithaca, new york: cornell university. marneweck, g.j. 1995. a first for south africa. joining efforts to manage and rehabilitate the first transborder ramsar site. south african wetlands. newsletter on activities relating to the ramsar convention in south africa 5: 1. middleton, b. 1999. wetland restoration. flood pulsing and disturbance dynamics. new york: john wiley. mitsch, w.j. & r.f. wilson. 1996. improving the success of wetland creation and restoration with know-how, time, and self-design. ecological applications 6(1): 77-83. myburgh, w.j., p.j.j. breytenbach, g.k. theron & g.j. bredenkamp. 1995. die fitososiologie van die vleie in die grootvlei-omgewing, suidtransvaal. suid-afrikaanse tydskrif vir wetenskap en tegnologie 14(2): 48-54. rand water scientific services. 1998. the socioeconnomic value of wetlands in highly industrialized catchments. development of a programme for the klip river catchment 2: 1. (unpublished). rand water & wording for water programme. 1998. wetland rehabilitation programme for the wilge river. harrismith: department of environmental affairs & tourism, free state conservation. smit, c.m., g.j. bredenkamp & n. van rooyen. 1995. the vegetation of the upper klip river valley in the north-eastern orange free state. navorsinge van die nasionale museum, bloemfontein 11: 37-58. van oudtshoorn, f. 1999. guide to grasses of southern africa. pretoria: briza. van wyk, b. & s. malan. 1988. veldgids tot die veldblomme van die witwatersrand& pretoriagebied, insluitend magaliesberg & suikerbosrand. kaastad: struik. werger, m.j.a. 1974. on concepts and techniques applied in the zürich-montpellier method of vegetation survey. bothalia 11(3): 309-32. issn 0075-6458 63 koedoe 46/1 (2003) venter.qxd 2005/12/09 10:34 page 63 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <feff0055007300650020006500730074006100730020006f007000630069006f006e006500730020007000610072006100200063007200650061007200200064006f00630075006d0065006e0074006f0073002000500044004600200063006f006e0020006d00610079006f00720020007200650073006f006c00750063006900f3006e00200064006500200069006d006100670065006e00200071007500650020007000650072006d006900740061006e0020006f006200740065006e0065007200200063006f007000690061007300200064006500200070007200650069006d0070007200650073006900f3006e0020006400650020006d00610079006f0072002000630061006c0069006400610064002e0020004c006f007300200064006f00630075006d0065006e0074006f00730020005000440046002000730065002000700075006500640065006e00200061006200720069007200200063006f006e0020004100630072006f00620061007400200079002000520065006100640065007200200035002e003000200079002000760065007200730069006f006e0065007300200070006f00730074006500720069006f007200650073002e0020004500730074006100200063006f006e0066006900670075007200610063006900f3006e0020007200650071007500690065007200650020006c006100200069006e0063007200750073007400610063006900f3006e0020006400650020006600750065006e007400650073002e> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 abstract introduction materials and methods discussion acknowledgements references about the author(s) elie gaget centre d’ecologie fonctionnelle et evolutive, montpellier, france department of biology, university of turku, turku, finland catherine l. parr department of earth, ocean and ecological sciences, school of environmental sciences, university of liverpool, liverpool, united kingdom school of animal, plant and environmental sciences, university of the witwatersrand, johannesburg, south africa department of zoology and entomology, university of pretoria, pretoria, south africa clélia sirami centre d’ecologie fonctionnelle et evolutive, montpellier, france climate change and bioadaptation division, south african national biodiversity institute, cape town, south africa université de toulouse, castanet-tolosan, france citation gaget, e., parr, c.l. & sirami, c., 2020, ‘effects of fire frequency on savanna butterfly diversity and composition: a preliminary study’, koedoe 62(1), a1617. https://doi.org/10.4102/koedoe.v62i1.1617 note: additional supporting information may be found in the online version of this article as online appendix 1. original research effects of fire frequency on savanna butterfly diversity and composition: a preliminary study elie gaget, catherine l. parr, clélia sirami received: 06 jan. 2020; accepted: 13 july 2020; published: 21 sept. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract fire plays a major role in many biomes, is widely used as a management tool and is likely to be affected by climate change. for effective conservation management, it is essential to understand how fire regimes affect different taxa, yet responses of invertebrates are particularly poorly documented. we tested how different fire frequencies influence savanna butterfly diversity and composition by using a long-term savanna fire experiment initiated in 1954 in the kruger national park (south africa). we compared butterfly abundance, species richness and community composition across three fire frequencies: high (burnt annually), medium (burnt triennially) and low (burnt twice in 60 years). plots with high fire frequency hosted higher abundance than mediumor low-frequency plots. fire frequencies did not affect species richness, but they led to distinct communities of butterflies. our findings suggest that, in view of the three fire frequencies tested, a spatial diversity of fire frequencies may increase butterfly diversity at the landscape level in wet savannas. managers may need to promote a greater diversity of fire frequencies by increasing fire frequency in some areas to provide habitat for species requiring high fire frequency, and by decreasing fire frequency in a large proportion of the landscape to provide fire refuges. this study provides new insights for butterfly conservation in savannas and highlights several knowledge gaps, which further studies should address for insect responses to be given adequate consideration in fire management strategies. conservation implications: a spatial diversity of fire frequencies may increase butterfly diversity. managers may need to promote a greater diversity of fire frequencies by increasing fire frequency in some areas to provide habitat for species requiring high fire frequency, and by decreasing fire frequency in other areas to provide fire refuges. keywords: climate change; conservation management; fire ecology; fire refugia; invertebrate pyrodiversity. introduction fire is a frequent and widespread disturbance in many biomes (pausas & keeley 2009) and shapes landscapes by modifying vegetation structure and composition (bond & keeley 2005). the recent increase in large uncontrolled fire incidences (e.g. in the western-mediterranean basin; pausas & fernández-muñoz 2012; and in the north american boreal region; kasischke & turetsky 2006) and predictions indicating that the frequency of high-intensity fires will increase in the future as a result of climate change (pechony & shindell 2010) have triggered growing concerns about the impacts of changes in long-term fire regimes. fire management has become increasingly important as both human population densities and pressure to manage fuels for asset protection increase (gill & stephens 2009). in parallel, fire has also become one of the primary tools for biodiversity conservation (andersen et al. 1998; parr & chown 2003). yet, numerous knowledge gaps are still currently impeding informed decision-making in fire management (driscoll et al. 2010). fire management practices for biodiversity conservation are often based on limited information, usually focussing on the response of plant communities in short-term studies or natural experiments by using space-for-time substitution (driscoll et al. 2010). however, data on the long-term ecological response of multiple taxa to fire regimes are crucial to design appropriate fire management strategies for biodiversity conservation (andersen et al. 1998). despite the fact that invertebrates represent the largest component of global biodiversity and ecosystem functioning (lavelle et al. 2006; losey & vaughan 2006), they are relatively poorly studied in fire ecology research (new et al. 2010; parr & chown 2003). in savannas, one of the world’s most fire-prone biomes, arthropods are considered to be generally well adapted to frequent fires with several recent studies across taxonomic groups highlighting their resilience (andersen & müller 2000; andersen et al. 2014; barrow, parr & kohen 2007; davies et al. 2012; parr et al. 2004). most studies have observed a rapid recovery of communities after fire, between 2 and 13 months after fire, depending on the insect order and savanna type (diniz, higgins & morais 2011; parr et al. 2004). the few existing studies on long-term savanna fire regimes observed only a weak overall effect of fire regime on insect diversity and composition, for example, for ants (maravalhas & vasconcelos 2014; parr et al. 2004) and for termites (davies et al. 2012). studies to date therefore suggest that little pyrodiversity (i.e. variability or diversity of fire regimes, parr & andersen 2006) may be required to conserve savanna insect diversity (but see maravalhas & vasconcelos 2014). however, beale et al. (2018) recently showed that the effects of pyrodiversity on birds and mammals across african savannas are rainfall contingent, with positive effects on these taxa particularly pronounced in wet savannas (> 650 mm/year). beale et al.’s (2018) findings support studies suggesting that fire regime may have a greater effect on ants and termites in mesic savannas (> 650 mm/year; davies et al. 2012; parr et al. 2004). it therefore seems likely that invertebrates may also be more responsive to pyrodiversity in wet savannas. furthermore, the role of fire regime may also depend on species traits, in particular specific habitat and dietary requirements. as a result, although most studies highlight the high resilience of invertebrate groups to single fire events and the weak effect of fire regime, this may not be true in wet savannas for invertebrate groups that have more specific habitat and dietary requirements. butterflies (lepidoptera) are herbivorous species, commonly used as indicator species to reveal impacts of environmental change on biodiversity (e.g. devictor et al. 2012). butterflies might be greatly influenced by changes in vegetation structure and composition caused by changes in fire frequency (pryke & samways 2012) because of their food specialisation at the larval (gilbert & singer 1975) and adult (mevi-schütz & erhardt 2003) stages. indeed, their distributions and population dynamics could be structured by the various plant species composition and structure shaped by different fire regimes (smit et al. 2010; smith et al. 2012). because of their ecological requirements, butterfly species can be positively affected by frequent fires (gardiner & terblanche 2010) or conversely negatively impacted by them (de andrade et al. 2017). as a result, we expect savanna butterflies to be significantly affected by habitat differences that can develop as a result of long-term fire regimes in wetter savannas, from closed to open savanna depending of the fire frequency (smit et al. 2010). we studied butterfly communities associated with different fire frequencies by using a long-term fire experiment in a wet savanna (> 650 mm/year) of kruger national park, south africa. fire regimes can be characterised by five components (fuel consumption and spread patterns, fire intensity, severity, frequency and seasonality; bond & keeley 2005), which are often correlated (díaz-delgado, lloret & pons 2003; govender, trollope & van wilgen 2006). we here focussed on the effect of fire frequency on butterfly communities. we sampled plots burnt at different frequencies from annual fires with open savanna to quasi-fire exclusion with closed savanna (smit et al. 2010). firstly, based on results observed for vertebrates (beale et al. 2018), we expected butterfly species richness to be promoted with more frequent fires. secondly, because changes in fire frequency induce changes in plant diversity and composition as well as vegetation structure (as already shown in the same study area by smit et al. 2010; smith et al. 2012), we expected fire frequency to influence both butterfly diversity and composition. furthermore, we also expected butterfly species to show some degree of specialisation to specific fire frequencies. these last two hypotheses are related to the pyrodiversity–biodiversity hypothesis, which predicts that a spatial diversity of fire regimes should increase biodiversity at the landscape level (parr & andersen 2006). finally, because butterfly communities are likely to show dynamic patterns with time since fire, we sampled butterfly communities across three sampling periods. we expected butterfly diversity to increase with time since fire, in line with the recovery of butterfly communities, particularly in plots burnt at intermediate and high frequencies. materials and methods study site the study was conducted in the pretoriuskop region of kruger national park, south africa (knp; 31°10’e, 25°10’s). the topography is moderately undulating, varies in altitude from 560 to 640 m above sea level and is dominated by nutrient-poor soils (granite-derived sandy soils). pretoriuskop has a warm subtropical climate with annual temperature averaging 27 °c (ranging from 4 °c to 39 °c) and moderately high annual rainfall (744 mm/year on average; gertenbach 1983). the wet season occurs during summer, with 85% of total annual rainfall occurring between october and march (gertenbach 1980). the vegetation corresponds to a mesic savanna characterised by the predominance of broadleaved thornless tree species, mainly silver cluster-leaf (terminalia sericea), marula (sclerocarya birrea), sickle bush (dichrostachys cinerea) and bush willow (combretum collinum; mucina & rutherford 2006). dominant grasses include setaria sphacelata, digitaria eriantha and loudetia simplex (davies et al. 2014). fires are typically surface fires occurring during the dry winter months with a mean return period of 3.5 years in areas that receive a mean annual rainfaxll of over 700 mm (van wilgen et al. 2000). study design the experimental burn plot (ebp) trial was initiated in 1954 by using a pseudo-randomised block design (biggs et al. 2003). butterfly communities were sampled in three replicate sites in the pretoriuskop area: the numbi, shabeni and kambeni sites (figure 1). we compared butterfly communities among three fire treatments. high fire frequency plots (hereafter referred to as high) were burnt every year in winter, and characterised by an open savanna dominated by grass species, although some tall trees were still present (govender et al. 2006; smith et al. 2010). medium fire frequency plots (hereafter referred to as medium) were burnt every 3 years in winter (august 2009) and were associated with an open savanna and sparse woody vegetation (smith et al. 2010). low fire frequency plots (hereafter referred to as low) were subjected to less than two burn events since 1954 and were characterised by a closed savanna, dominated by woody vegetation (smith et al. 2010). plots associated with low fire frequency correspond to unburnt plots within the ebp where accidental fires occurred (in 2011 in shabeni and in 2002 and 2010 in numbi). each plot measured approximately 380 × 180 m (7 ha) and was separated from adjacent plots by firebreaks and roads. wild megafauna was free to browse and graze on all plots. browsing and grazing are known to interact with fire frequency (archibald et al. 2005). the present study does not aim to disentangle the respective effect of fire frequency and browsing or grazing, a topic already investigated in other studies (see burkepile et al. 2016; trollope et al. 2014). we studied the three fire treatments at each of the three sites, that is a total of nine plots; replication was limited because of experimental design constraints, but the uniqueness of this fire experiment – specifically its long-term and large-scale nature – makes the results valuable (see also parr et al. 2004). figure 1: the pretoriuskop study area in the south-western part of the kruger national park (lower insert, 31°10’e, 25°10’s), south africa (upper insert). black rectangles correspond to the long-term experimental burn plots set up in 1954 and red rectangles represent the nine plots sampled within the three sites (numbi, shabeni and kambeni). butterfly sampling butterfly communities were surveyed in each plot by using the pollard walk transect method commonly used in the literature (pollard & yates 1993). surveys were conducted by a team of local butterfly experts led by i.a. sharp during the austral summer (december–march), which corresponds to the peak flight period for most butterfly species (woodhall 2005). each plot was visited during three sampling periods in december 2009, march 2010 and january 2012. consequently, in plots with medium fire frequency, sampling was conducted 4, 7 and 29 months after burning, whereas high-frequency plots were sampled at 4, 5 and 7 months after burning. during each sampling period, two transects of 380 m were sampled between opposite corners of the plot (transects started 20 m away from the border of the plot to limit edge effects). for each sampling period, we considered species richness and abundance per plot as the total number of species and the total number of individuals recorded on the two transects, respectively. butterflies were recorded by walking along each transect at a steady pace (around 12 m/min), thus taking 30 min to be completed. our sampling effort was therefore similar to those commonly used in the literature and can be considered sufficient to compare species diversity and composition between plots (e.g. davis, debinski & danielson 2007; koh & sodhi 2004; vasconcelos et al. 2015). in our study, we observed 78 species (see below), which corresponds to 72% of the 108 species occurring in our study area (pretoriuskop; based on otto 2014). considering that our sampling sites only include open-closed savanna (smit et al. 2010), but not other habitats found in this area, this suggests that our sampling provides a good representation of the local butterfly community (online appendix 1 figure s2). nevertheless, we acknowledge that sampling in december, january and march resulted in the exclusion of at least 15 species observed only during the rest of the year, based on the atlas of african lepidoptera (fitzpatrick institute of african ornithology 2020). surveys were restricted to clear, windless days between 10:00 and 14:00. all butterflies seen within 5 m of the transect line were recorded (pollard 1977). most butterflies were identified to species level; when this was not possible, the recorder would net the butterfly and identify the species in the field. individuals flying from behind the recorders were excluded to avoid double counting (pollard 1977). statistical analyses to evaluate differences in species richness and abundance across the different fire frequencies, the cumulative species richness and abundance over the three sampling periods was calculated. a generalised linear mixed-effects model (glmm, poisson error distribution) was applied to data with species richness and abundance set as response variables and fire frequency (i.e. high, medium and low) set as explanatory variables. site identity (numbi, shabeni and kambeni) was added as random factor (see online appendix 1 figure s1 for the effect of site identity on cumulative species richness and cumulative abundance). the likelihood-ratio chi-square was computed to interpret the significance of each model, and data were checked for residual homoscedasticity. differences in community composition between the three fire frequencies, based on cumulative presence or absence and abundance data over the three sampling periods, were evaluated by using the jaccard dissimilarity on butterfly species presence or absence and bray–curtis dissimilarity on abundance. both these indices exclude joint absences and do not consider double absences as a factor of similarity (legendre & legendre 1998). a principal coordinates analysis (pcoa, package ‘ade4’, dray & dufour 2007) was constructed to visually represent differences in community composition between the three fire frequencies based on jaccard (presence or absence) and bray–curtis (abundance) distance matrices. principal coordinates analysis is used to represent objects from a distance matrix by preserving the distance relationships among objects in the full-dimensional principal coordinate euclidean space (gower 1966). we also performed a permutational multivariate analysis of variance (permanova, function ‘adonis’ in package ‘vegan’, oksanen et al. 2013) on the jaccard and bray–curtis distance matrix with fire frequency as an explanatory variable. additionally, we conducted an indicator species analysis (indval; dufrêne & legendre 1997) to determine whether certain butterfly associated more strongly with particular fire frequencies. indval was applied to identify species associations with fire frequencies for each sampling period. we used the function ‘multipatt’ (package ‘indicspecies’, de caceres & jansen 2016). based on mcgeoch, van rensburg and botes (2002), we retained only indicator species with significant indvals greater than 70%. the recovery of butterfly diversity with time since fire in plots burnt at high and medium fire frequencies was determined by analysing the effect of time since fire on species richness and abundance for these two fire frequencies. to this end, a glmm (poisson error distribution, package ‘lme4’, bates et al. 2014) was applied for each fire frequency (high and medium), with species richness and abundance as a response variable and sampling period as an explanatory variable (december 2009, march 2010, january 2012). site identity (numbi, shabeni and kambeni) was added as a random factor. the likelihood-ratio chi-square was computed to interpret the significance of each model, and we checked for residual homoscedasticity. all statistical analyses and graphic representations were conducted in r 3.4.3 (r development core team 2018). ethical consideration this article followed all ethical standards for a research without direct contact with human or animal subjects. results a total of 1809 individuals from 78 species were recorded, with an average of 67 ± 53.3 individuals (mean ± standard deviation) and 14.8 ± 4.9 species per plot per sampling period. twelve of these species were recorded in more than 10 sampling periods, although 39 species (50% of all species) were recorded in three or less of the sampling periods (online appendix 1). effect of fire frequency on butterfly species richness and abundance species richness did not differ significantly between the three fire frequencies (figure 2, χ² = 0.65, degree of freedom [df] = 5, p = 0.7). however, butterfly abundance was significantly higher in high-frequency plots, compared with the medium (figure 2, β = 0.53 ± 0.06, df = 2, z = 9.21, p < 0.001) and low fire frequency plots (figure 2, β = 0.52 ± 0.06, df = 2, z = 9.06, p < 0.001). abundance did not differ significantly between medium and low fire frequency plots (figure 2, p > 0.05). the random effect associated with site identity was not highly informative in the applied model (deviance < 0.0001). however, abundance varied significantly between sites, whereas species richness did not reveal any significant variation between sites (online appendix 1 figure s1). figure 2: the effect of fire frequency on cumulative species richness (a) and cumulative abundance (b) over the three sampling periods. overall significant differences (95% confidence interval) are indicated by *** (p < 0.001). fire frequencies associated with distinct letters are significantly different. effect of fire frequency on community composition the first two axes of the analysis based on the presence or absence data (jaccard) explained 40.6% and 20.4% of the inertia (figure 3a), whereas the first two axes based on abundance data (bray–curtis) explained 20.9% and 20.5% of the inertia (figure 3b). community composition based on abundance data (bray–curtis) was significantly affected by fire frequency (f = 2.45, r² = 0.43, p = 0.006). community composition based on the presence or absence was not significantly different between fire frequencies, despite a low p-value (f = 1.54, r² = 0.34, p = 0.07). both analyses revealed a heavily structured distribution of plots burnt at different frequencies along the first axis (figure 3a and b). figure 3: position of fire frequencies on the axes 1–2 of the principal coordinates analysis performed on the distance matrix of the presence or absence (a, jaccard) the abundance (b, bray–curtis). indicator species five species were strongly associated with a specific fire frequency (table 1; see abundance per fire frequency, per period and per site for these five species in table 2). among these species, cupreous blue (eicochrysops messapus) and broad-bordered grass yellow (eurema brigitta brigitta) were indicators of most frequently burnt plots (high and medium), whilst common blue (leptotes pirithous pirithous), clear-spotted acraea (acraea aglaonice) and zulu shadefly (coenyra hebe) were indicators of infrequently burnt plots (low) (online appendix 1 figure s3). table 1: percentage of species association with each fire frequency (high, medium and low) during each sampling period (december, january and march) and associated p-value based on the indval analysis. table 2: abundance of each identified indicator species per fire frequency, per sampling period and per site. recovery of butterfly diversity after fire in plots burnt at high fire frequency, species richness did not differ significantly between 4, 5 and 7 months after fire (figure 4, χ² = 2.15, p = 0.3). however, abundance was significantly higher 5 months after fire, compared with 4 and 7 months after fire (β = 1.43 ± 0.10, z = 14.83, p < 0.001 and β = 1.40 ± 0.10, z = 14.70, p < 0.001, respectively). abundance did not differ between 4 and 7 months after fire (β = 0.30 ± 0.10, z = 0.24, p = 0.9). the random effect associated with site identity was not highly informative in each model (deviance < 0.0001). figure 4: effect of time since fire on butterfly species richness and abundance for high fire frequency (annual burning in august) and medium fire frequency (triennial burning in august). overall significant differences (95% confidence interval) are indicated by ** (p < 0.01) or *** (p < 0.001). significant time effects on the species richness and abundance are indicated by lowercase letters. in plots burnt at medium fire frequency, butterfly species richness and abundance were significantly affected by time since fire (figure 4, χ² = 10.39, p = 0.006; χ² = 60.04, p < 0.001, respectively). species richness was lower 4 months after burning than 7 and 29 months after burning (figure 4, β = −0.56 ± 0.26, z = −2.19, p = 0.03; β = −0.79 ± 0.25, z = −3.22, p = 0.001, respectively). similarly, abundance was lower 4 months after burning than 7 and 29 months after burning (figure 4, β = −0.89 ± 0.13, z = −6.69, p < 0.001; β = −0.99 ± 0.13, z = −7.53, p < 0.001, respectively). however, neither the species richness nor the abundance differs between 7 and 29 months after fire (figure 4, β = 0.23 ± 0.21, z = 1.13, p = 0.2; β = 0.10 ± 0.10, z = 0.99, p = 0.3, respectively). the random effect associated with site identity was not highly informative in each model (deviance < 0.0001). discussion this study represents a first assessment on the effect of differences in long-term fire frequencies on butterfly communities in wet african savannas. we observed significant differences in species abundance and species composition between fire frequencies. the high abundance of butterflies observed in high fire frequency plots lends support to the idea that most savanna butterfly species are resilient to, or favoured by, some level of disturbance compared with no disturbance (wilkerson, roche & young 2013). butterflies are sensitive to changes in plant diversity, which may influence their distribution at both larval and adult stage (gilbert & singer 1975; mevi-schütz & erhardt 2003). the fact that herbaceous species diversity is greater in plots burnt at high frequency than in plots burnt at medium frequency in the kruger national park (burkepile et al. 2016) may contribute to increased butterfly species abundances. the results partially confirm the specialisation hypothesis that predicts that different fire frequencies hosting different plant species composition will be associated with different butterfly communities (gilbert & singer 1975). significant differences across fire frequencies (figures 3a and b) is in accord with the fact that differences in fire frequency result in differences in plant species composition, as already shown in the same study area (smith et al. 2012). in this long-term experiment, herbaceous vegetation has been shown to be heavily influenced by fire frequency, with lower herbaceous diversity in plots burnt at low frequency than in plots burnt at high or medium frequencies (smith et al. 2012). this was true even it is accidental fire occurred in the low fire frequency plots, where the differences in vegetation structure and composition compared with frequently burnt plots remain very high. such differences in host plants for the larval stage or flowers for the adult stage are likely to explain differences in butterfly community composition observed among ebps (swengel 2001). accurate monitoring of the diversity of host plants and data on butterfly functional traits would, however, be necessary to confirm this preliminary finding. we identified indicator species for each fire frequency, suggesting that butterfly species respond differently to variations in fire frequency. literature suggests that butterfly forest specialists are generally sensitive to fire and can be negatively affected by frequent burning (de andrade et al. 2017). accordingly, we found that low fire frequency benefits the species that prefer shaded areas in mixed woodland and forest, like the zulu shadefly (online appendix 1 figure s3). conversely, medium and high fire frequencies had positive effects on some open savanna species, such as the broad-bordered grass yellow and the cupreous blue (online appendix 1 figure s3), which are resilient to fire and promoted by frequent burning (adamidis et al. 2019). this study furthermore suggests that butterfly diversity may be able to recover quickly after a fire. indeed, butterfly species richness and abundance were similar 7 months and 29 months after the last fire in plots burnt triennially. this high speed of recovery has also been observed with ants (parr et al. 2004) – likely linked to rapid recovery of vegetation. the recovery of butterfly communities could be attributed to an immigration effect from the surrounding landscape (moranz, fuhlendorf & engle 2014; robinson et al. 2013). the small size of experimental plots means that recovery time may be longer in natural landscapes where bigger fires occur. further research is necessary to address variations in recovery time between species (vogel, koford & debinski 2010). this study was conducted in one of the largest and oldest fire experiments globally. despite its many advantages, it is important to acknowledge the limitations with this long-term experiment, especially for mobile invertebrates such as butterflies. in their adult stage, butterflies can cover vast distances (hundreds of metres) (pollard 1977) and are therefore able to move from one 7 ha plot to another. although we did not include individual butterflies along the border of plots (20 m), it is possible that some of the individuals recorded during our surveys came from outside the plots. the presence of non-local vagrant butterflies may have influenced our ability to accurately assess species diversity (avuletey & niba 2014). this may explain the lack of difference in community composition based on species occurrence between fire frequencies in our study. more research should be conducted on transient individuals, notably on their ability to cross different vegetation structures, to assess differences in community composition and species richness between ebps with higher certainty levels (wood & samways 1991). additionally, even long-term experiment may be affected by inter-annual variations, such as, for instance, different climatic variations between years impacting either fire intensity or vegetation recovery after fire. for example, some species were very abundant 5 months after fire compared with 4 or 7 months after fire, in the high frequently burnt plot (figure 2). indeed, 4 and 7 months after fire correspond to the summer 2009–2010, although 5 months after fire correspond to 2012. lastly, although we appreciate the sample size is relatively low, the size and duration of this experiment are valuable. moreover, nevertheless, we were able to detect differences among the treatments suggesting that in this habitat, fire regimes can have a significant impact on butterfly communities. this preliminary assessment focusses on fire frequency, an easy parameter for conservation managers to measure (biggs et al. 2003). in the pretoriuskop area of kruger national park, no part of the landscape currently has a fire return period of 2 years or less (sanparks, unpublished) and only 6% of the area burns infrequently (fire return period of > 5 years; sanparks, unpublished). this suggests that managers may need to promote a greater diversity of fire frequencies by (1) increasing fire frequency in some areas to provide habitat for species requiring high fire frequency; and (2) decreasing fire frequency in a larger proportion of the area. our study highlights that insect responses should be given more consideration to determine whether the current fire strategy in kruger national park is adequate in the long term for biodiversity conservation. further studies should be conducted to explore the role of other components of fire regime on wet savanna biodiversity (beale et al. 2018; hempson et al. 2017) in relation with butterfly dispersal capacities and the ability of larvae to escape fire, to explicitly address colonisation and extinction processes after fire. acknowledgements we are especially grateful to ian and allison sharp and their family for conducting all the butterfly surveys as well as all the volunteers who assisted them, in particular valentina lupano, richard mercer and julie wolhunter. we thank south african national parks, and especially navashni govender, for logistical support. we gratefully acknowledge funding from the rufford small grants, national geographic and oxford university john fell fund. we thank the two anonymous reviewers for helpful comments. competing interests the authors have declared that no competing interest exists. authors’ contributions e.g., c.s. and c.l.p. conceived and designed the study, e.g. and c.s. performed the analyses, e.g. and c.s. wrote the manuscript and all authors edited and approved it. funding information funding for the study was received from the rufford small grants, national geographic and oxford university john fell fund. data availability statement the data used in this study will be archived in the dryad digital repository. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references adamidis, g.c., swartz, m.t., zografou, k. & sewall, b.j., 2019, ‘prescribed fire maintains host plants of a rare grassland butterfly’, scientific reports 9(1), 1–12. https://doi.org/10.1038/s41598-019-53400-1 andersen, a.n., braithwaite, r.w., cook, g.d., corbett, l.k., williams, r.j., douglas, m.m. et al., 1998, ‘fire research for conservation management in tropical savannas: introducing the kapalga fire experiment’, australian journal of ecology 23(2), 95–110. https://doi.org/10.1111/j.1442-9993.1998.tb00708.x andersen, a.n. & müller, w.j., 2000, ‘arthropod responses to experimental fire regimes in an australian tropical savannah: ordinal-level analysis’, austral ecology 25(2), 199–209. https://doi.org/10.1046/j.1442-9993.2000.01038.x andersen, a.n., ribbons, r.r., pettit, m. & parr, c.l., 2014, ‘burning for biodiversity: highly resilient ant communities respond only to strongly contrasting fire regimes in australia’s seasonal tropics’, journal of applied ecology 51(5), 1406–1413. https://doi.org/10.1111/1365-2664.12307 archibald, s., bond, w.j., stock, w.d. & fairbanks, d.h.k., 2005, ‘shaping the landscape: fire–grazer interactions in an african savanna’, ecological applications 15(1), 96–109. https://doi.org/10.1890/03-5210 avuletey, r. & niba, a.s., 2014, ‘butterfly (lepidoptera) assemblage composition and distribution patterns in king sabata dalindyebo municipality, eastern cape, south africa’, african entomology 22(1), 57–67. https://doi.org/10.4001/003.022.0103 barrow, l., parr, c.l. & kohen, j.l., 2007, ‘habitat type influences fire resilience of ant assemblages in the semi-arid tropics of northern australia’, journal of arid environments 69(1), 80–95. https://doi.org/10.1016/j.jaridenv.2006.08.005 bates, d., maechler, m., bolker, b. & walker, s., 2014, ‘lme4: linear mixed-effects models using eigen and s4’, r package version 1(7), 1–23. https://doi.org/10.18637/jss.v067.i01 beale, c.m., courtney mustaphi, c.j., morrison, t.a., archibald, s., anderson, t.m., dobson, a.p. et al., 2018, ‘pyrodiversity interacts with rainfall to increase bird and mammal richness in african savannas’, ecology letters 21(4), 557–567. https://doi.org/10.1111/ele.12921 biggs, r., biggs, h.c., dunne, t.t., govender, n. & potgieter, a.l.f., 2003, ‘experimental burn plot trial in the kruger national park: history, experimental design and suggestions for data analysis’, koedoe – african protected area conservation and science 46(1), 1–15. https://doi.org/10.4102/koedoe.v46i1.35 bond, w.j. & keeley, j.e., 2005, ‘fire as a global “herbivore”: the ecology and evolution of flammable ecosystems’, trends in ecology & evolution 20(7), 387–394. https://doi.org/10.1016/j.tree.2005.04.025 burkepile, d.e., thompson, d.i., fynn, r.w.s., koerner, s.e., eby, s., govender, n. et al., 2016, ‘fire frequency drives habitat selection by a diverse herbivore guild impacting top–down control of plant communities in an african savanna’, oikos 125(11), 1636–1646. https://doi.org/10.1111/oik.02987 davies, a.b., eggleton, p., van rensburg, b.j. & parr, c.l., 2012, ‘the pyrodiversity–biodiversity hypothesis: a test with savanna termite assemblages’, journal of applied ecology 49(2), 422–430. https://doi.org/10.1111/j.1365-2664.2012.02107.x davies, a.b., robertson, m.p., levick, s.r., asner, g.p., van rensburg, b.j. & parr, c.l., 2014, ‘variable effects of termite mounds on african savanna grass communities across a rainfall gradient’, journal of vegetation science 25(6), 1405–1416. https://doi.org/10.1111/jvs.12200 davis, j.d., debinski, d.m. & danielson, b.j., 2007, ‘local and landscape effects on the butterfly community in fragmented midwest usa prairie habitats’, landscape ecology 22(9), 1341–1354. https://doi.org/10.1007/s10980-007-9111-9 de andrade, r.b., balch, j.k., carreira, j.y., brando, p.m. & freitas, a.v., 2017, ‘the impacts of recurrent fires on diversity of fruit-feeding butterflies in a south-eastern amazon forest’, journal of tropical ecology 33(1), 22. https://doi.org/10.1017/s0266467416000559 de caceres, m. & jansen, f., 2016, ‘package “indicspecies”’, indicators 8, 1. devictor, v., van swaay, c., brereton, t., brotons, l., chamberlain, d., heliölä, j. et al., 2012, ‘differences in the climatic debts of birds and butterflies at a continental scale’, nature climate change 2(2), 121–124. https://doi.org/10.1038/nclimate1347 díaz-delgado, r., lloret, f. & pons, x., 2003, ‘influence of fire severity on plant regeneration by means of remote sensing imagery’, international journal of remote sensing 24(8), 1751–1763. https://doi.org/10.1080/01431160210144732 diniz, i.r., higgins, b. & morais, h.c., 2011, ‘how do frequent fires in the cerrado alter the lepidopteran community?’, biodiversity and conservation 20(7), 1415–1426. https://doi.org/10.1007/s10531-011-0034-z dray, s. & dufour, a.b., 2007, ‘the ade4 package: implementing the duality diagram for ecologists’, journal of statistical software 22(4), 1–20. https://doi.org/10.18637/jss.v022.i04 driscoll, d.a., lindenmayer, d.b., bennett, a.f., bode, m., bradstock, r.a., cary, g.j. et al., 2010, ‘fire management for biodiversity conservation: key research questions and our capacity to answer them’, biological conservation 143(9), 1928–1939. https://doi.org/10.1016/j.biocon.2010.05.026 dufrêne, m. & legendre, p., 1997, ‘species assemblages and indicator species: the need for a flexible asymmetrical approach’, ecological monographs 67(3), 345–366. https://doi.org/10.2307/2963459 fitzpatrick institute of african ornithology, 2020, lepimap: atlas of african lepidoptera, viewed 24 june 2020, from http://vmus.adu.org.za/?vm=lepimap. gardiner, a.j. & terblanche, r.f., 2010, ‘taxonomy, biology, biogeography, evolution and conservation of the genus erikssonia trimen (lepidoptera: lycaenidae)’, african entomology 18(1), 171–191. https://doi.org/10.4001/003.018.0114 gertenbach, w.p.d., 1980, ‘rainfall patterns in the kruger national park’, koedoe – african protected area conservation and science 23(1), 35–43. https://doi.org/10.4102/koedoe.v23i1.634 gertenbach, w.p.d., 1983, ‘landscapes of the kruger national park’, koedoe – african protected area conservation and science 26(1), 9–121. https://doi.org/10.4102/koedoe.v26i1.591 gilbert, l.e. & singer, m.c., 1975, ‘butterfly ecology’, annual review of ecology and systematics 6(1), 365–395. https://doi.org/10.1146/annurev.es.06.110175.002053 gill, a.m. & stephens, s.l., 2009, ‘scientific and social challenges for the management of fire-prone wildland–urban interfaces’, environmental research letters 4(3), 034014. https://doi.org/10.1088/1748-9326/4/3/034014 govender, n., trollope, w.s. & van wilgen, b.w., 2006, ‘the effect of fire season, fire frequency, rainfall and management on fire intensity in savanna vegetation in south africa’, journal of applied ecology 43(4), 748–758. https://doi.org/10.1111/j.1365-2664.2006.01184.x gower, j.c., 1966, ‘some distance properties of latent root and vector methods used in multivariate analysis’, biometrika 53(3–4), 325–338. https://doi.org/10.1093/biomet/53.3-4.325 hempson, g.p., parr, c.l., archibald, s., anderson, t.m., mustaphi, c.j.c., dobson, a.p. et al., 2017, ‘continent-level drivers of african pyrodiversity’, ecography 41(6), 889–899. https://doi.org/10.1111/ecog.03109 kasischke, e.s. & turetsky, m.r., 2006, ‘recent changes in the fire regime across the north american boreal region – spatial and temporal patterns of burning across canada and alaska’, geophysical research letters 33(9), 1–5. https://doi.org/10.1029/2006gl025677 koh, l.p. & sodhi, n.s., 2004, ‘importance of reserves, fragments, and parks for butterfly conservation in a tropical urban landscape’, ecological applications 14(6), 1695–1708. https://doi.org/10.1890/03-5269 lavelle, p., decaëns, t., aubert, m., barot, s., blouin, m., bureau, f. et al., 2006, ‘soil invertebrates and ecosystem services’, european journal of soil biology 42(suppl 1), s3–s15. https://doi.org/10.1016/j.ejsobi.2006.10.002 legendre, p. & legendre, l., 1998, numerical ecology, volume 24, (developments in environmental modelling), elsevier, québec. losey, j.e. & vaughan, m., 2006, ‘the economic value of ecological services provided by insects’, bioscience 56(4), 311–323. https://doi.org/10.1641/0006-3568(2006)56[311:tevoes]2.0.co;2 maravalhas, j. & vasconcelos, h.l., 2014, ‘revisiting the pyrodiversity–biodiversity hypothesis: long-term fire regimes and the structure of ant communities in a neotropical savanna hotspot’, journal of applied ecology 51(6), 1661–1668. https://doi.org/10.1111/1365-2664.12338 mcgeoch, m.a., van rensburg, b.j. & botes, a., 2002, ‘the verification and application of bioindicators: a case study of dung beetles in a savanna ecosystem’, journal of applied ecology 39(4), 661–672. https://doi.org/10.1046/j.1365-2664.2002.00743.x mevi-schütz, j. & erhardt, a., 2003, ‘larval nutrition affects female nectar amino acid preference in the map butterfly (arashnia levana)’, ecology 84(10), 2788–2794. https://doi.org/10.1890/02-3108 moranz, r.a., fuhlendorf, s.d. & engle, d.m., 2014, ‘making sense of a prairie butterfly paradox: the effects of grazing, time since fire, and sampling period on regal fritillary abundance’, biological conservation 173(2014), 32–41. https://doi.org/10.1016/j.biocon.2014.03.003 mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, south african national biodiversity institute, pretoria. new, t.r., yen, a.l., sands, d.p.a., greenslade, p., neville, p.j., york, a. et al., 2010, ‘planned fires and invertebrate conservation in south east australia’, journal of insect conservation 14(5), 567–574. https://doi.org/10.1007/s10841-010-9284-4 oksanen, j., blanchet, f.g., kindt, r., legendre, p., minchin, p.r., o’hara, r.b. et al., 2013, ‘package “vegan”’, community ecology package version 2(9), 1-295. otto, h.h., 2014, butterflies of the kruger national park & surrounds, herbert otto, barberton. parr, c.l. & andersen, a.n., 2006, ‘patch mosaic burning for biodiversity conservation: a critique of the pyrodiversity paradigm’, conservation biology 20(6), 1610–1619. https://doi.org/10.1111/j.1523-1739.2006.00492.x parr, c.l. & chown, s.l., 2003, ‘burning issues for conservation: a critique of faunal fire research in southern africa’, austral ecology 28(4), 384–395. https://doi.org/10.1046/j.1442-9993.2003.01296.x parr, c.l., robertson, h.g., biggs, h.c. & chown, s.l., 2004, ‘response of african savanna ants to long-term fire regimes’, journal of applied ecology 41(4), 630–642. https://doi.org/10.1111/j.0021-8901.2004.00920.x pausas, j.g. & fernández-muñoz, s., 2012, ‘fire regime changes in the western mediterranean basin: from fuel-limited to drought-driven fire regime’, climatic change 110(1), 215–226. https://doi.org/10.1007/s10584-011-0060-6 pausas, j.g. & keeley, j.e., 2009, ‘a burning story: the role of fire in the history of life’, bioscience 59(7), 593–601. https://doi.org/10.1525/bio.2009.59.7.10 pechony, o. & shindell, d.t., 2010, ‘driving forces of global wildfires over the past millennium and the forthcoming century’, proceedings of the national academy of sciences 107(45), 19167–19170. https://doi.org/10.1073/pnas.1003669107 pollard, e., 1977, ‘a method for assessing changes in the abundance of butterflies’, biological conservation 12(2), 115–134. https://doi.org/10.1016/0006-3207(77)90065-9 pollard, e. & yates, t.j., 1993, monitoring butterflies for ecology and conservation, chapman & hall, london. pryke, j.s. & samways, m.j., 2012, ‘differential resilience of invertebrates to fire’, austral ecology 37(4), 460–469. https://doi.org/10.1111/j.1442-9993.2011.02307.x r development core team, 2018, r: a language and environment for statistical computing, r foundation for statistical computing, vienna. robinson, n.m., leonard, s.w.j., ritchie, e.g., bassett, m., chia, e.k., buckingham, s. et al., 2013, ‘refuges for fauna in fire-prone landscapes: their ecological function and importance’, journal of applied ecology 50(6), 1321–1329. https://doi.org/10.1111/1365-2664.12153 smit, i.p.j., asner, g.p., govender, n., kennedy-bowdoin, t., knapp, d.e. & jacobson, j., 2010, ‘effects of fire on woody vegetation structure in african savanna’, ecological applications 20(7), 1865–1875. https://doi.org/10.1890/09-0929.1 smith, m.d., wilgen, b.w., van, burns, c.e., govender, n., potgieter, a.l.f., andelman, s. et al., 2012, ‘long-term effects of fire frequency and season on herbaceous vegetation in savannas of the kruger national park, south africa’, journal of plant ecology 6(1), 71–83. https://doi.org/10.1093/jpe/rts014 swengel, a.b., 2001, ‘a literature review of insect responses to fire, compared to other conservation managements of open habitat’, biodiversity and conservation 10(7), 1141–1169. https://doi.org/10.1023/a:1016683807033 trollope, w., van wilgen, b., trollope, l.a., govender, n. & potgieter, a.l., 2014, ‘the long-term effect of fire and grazing by wildlife on range condition in moist and arid savannas in the kruger national park’, african journal of range & forage science 31(3), 199–208. https://doi.org/10.2989/10220119.2014.884511 van wilgen, b.w., biggs, h., o’regan, s.p. & mare, n., 2000, ‘fire history of the savanna ecosystems in the kruger national park, south africa, between 1941 and 1996’, south african journal of science 96(4), 167–178. vasconcelos, s., rodrigues, p., palma, l., mendes, l.f., palminha, a., catarino, l. et al., 2015, ‘through the eye of a butterfly: assessing biodiversity impacts of cashew expansion in west africa’, biological conservation 191, 779–786. https://doi.org/10.1016/j.biocon.2015.08.032 vogel, j.a., koford, r.r. & debinski, d.m., 2010, ‘direct and indirect responses of tallgrass prairie butterflies to prescribed burning’, journal of insect conservation 14(6), 663–677. https://doi.org/10.1007/s10841-010-9295-1 wilkerson, m.l., roche, l.m. & young, t.p., 2013, ‘indirect effects of domestic and wild herbivores on butterflies in an african savanna’, ecology and evolution 3(11), 3672–3682. https://doi.org/10.1002/ece3.744 wood, p.a. & samways, m.j., 1991, ‘landscape element pattern and continuity of butterfly flight paths in an ecologically landscaped botanic garden, natal, south africa’, biological conservation 58(2), 149–166. https://doi.org/10.1016/0006-3207(91)90117-r woodhall, s., 2005, field guide to butterflies of south africa, struik, cape town. abstract introduction methods results discussion conclusion acknowledgements references about the author(s) darren bouwer department of soil, crop and climate sciences, faculty of science and agriculture, university of the free state, bloemfontein, south africa pieter a.l. le roux department of soil, crop and climate sciences, faculty of science and agriculture, university of the free state, bloemfontein, south africa johan van tol department of soil, crop and climate sciences, faculty of science and agriculture, university of the free state, bloemfontein, south africa citation bouwer, d., le roux, p.a.l. & van tol, j., 2020, ‘identification of hydropedological flowpaths in stevenson–hamilton catena from soil morphological, chemical and hydraulic properties’, koedoe 62(2), a1584. https://doi.org/10.4102/koedoe.v62i2.1584 note: special issue: connections between abiotic and biotic components of a granite catena ecosystem in kruger national park, sub-edited by beanelri janecke and johan van tol. original research identification of hydropedological flowpaths in stevenson–hamilton catena from soil morphological, chemical and hydraulic properties darren bouwer, pieter a.l. le roux, jacobus j. van tol received: 30 aug. 2019; accepted: 05 may 2020; published: 29 oct. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract in highly variable water regimes of semi-arid savannahs, water is the key driving force for biotic and abiotic processes. understanding and measuring components of the hydrological cycle at landscape scale is however difficult because of the spatiotemporal variation of these processes. hydropedology is a new interdisciplinary research field aiming to use soil information to conceptualise hydrological processes at different scales. in this study, a hydropedological approach was used to identify key hydrological flowpaths on a granitic catena in the stevenson–hamilton supersite in the kruger national park. soils from 49 plots, spaced 10 m apart along a catena, were classified, and their morphology was interpreted in relation to the dominant hydrological response. soil samples were taken at 10-cm-depth intervals for chemical and physical analysis to assess the relationship between their expected hydrological behaviour and physiochemical properties. the hydropedological survey indicated that the crest is dominated by freely drained recharge soils where infiltration and vertical drainage are dominant. on the midslope, the underlying bedrock has restricted permeability; this promotes lateral flow at the soil/bedrock interface. on the upper footslope, high clay content soils (sodic) restricts further lateral drainage, resulting in return flow (seepage). overland flow is dominant on the upper and lower footslope. the valley bottom is occupied by freely drained alluvial soils, which act as a recharge zone. the chemical and physical analyses of soil support the interpretations of the hydropedological interpretation of the soil morphology. conservation implications: understanding hydrological processes is important for sustainable water resource management, especially in the areas with highly variable water regimes. a hydropedological approach provides an efficient method to characterise dominant flowpaths at landscape scale. this aids the estimation of the hydrological sensitivity of the landscape to climate and land use changes. keywords: hillslope; hydrology; soil morphology; semi-arid environments; hydrological processes; savanna ecosystems. introduction soil acts as a first-order control in the portioning of hydrological flowpaths and governing residence periods of water in a landscape. water, on the other hand, is a primary agent in soil formation and leaves unique soil morphological signatures of water movement and storage. this interactive relationship between soil and water serves as the basis for a relatively new, interdisciplinary field of study called hydropedology (lin 2010; lin et al. 2006a). this synergistic research field promotes a more integrated approach to address complex issues regarding water resource management in variable water regimes of semi-arid landscapes (kutílek & nielsen 2007; lin 2003; lin et al. 2006b). hydropedological studies at landscape scale typically include various components of the hydrological cycle and recognise the intimate relationship between surface and groundwater at various spatiotemporal scales (van tol & lorentz 2018). in the recent past, hydropedological studies done on landscape scale in south africa have focussed on the interpretation of soil morphological properties, and their spatial distribution, in order to derive the hydrological behaviour of hillslopes and catchments (e.g. van tol et al. 2010; kuenene et al. 2013). these qualitative descriptions of the dominant hydrological behaviour were then used to construct conceptual flowpath models that were used in hydrological modelling (van zijl et al. 2016) to determine the migration paths of pollutants (mamera & van tol 2018), wetland rehabilitation (riddell, lorentz & kotze 2012) and the impact of land-use changes (e.g. open-cast mining) on water resources (van tol et al. 2018). the underlying assumption of these hydrological interpretations of soil morphology is that soil morphological properties are reflecting the current soil–water regime. the irreversible nature and slow reaction of some soil properties to soil-forming processes (especially climate) and the lack of morphological expression in immature soils question the correctness of interpretations derived from morphological properties alone (lin et al. 2005). the process of soil formation is relatively long (102–104 years), and soil morphology might, therefore, represent relict environmental conditions (macewan & fitzpatrick 1996). although there is sufficient evidence that soil morphology correlates well to current soil water regimes in wetter parts of south africa (e.g. van huyssteen et al. 2007; van tol et al. 2010), there is still a need to establish these correlations in drier climates. soil chemical properties are less buffered to environmental changes than morphological properties (macewan 1997). the factors influencing chemical processes are similar to those influencing pedological process (kutílek & nielsen 2007), water being the prime mediator of chemical reactions in soils (essington 2004). soil chemical properties and their spatial distribution (vertically and laterally) can, therefore, be interpreted to identify hydrological flowpaths (bouwer et al. 2015) and verify the correctness of interpretations and conceptual models of hydrological behaviour that were derived from soil morphology alone. this work forms part of a larger interdisciplinary project that focused on the aboveground–belowground ecological patterns and their relationship with granitic catenal processes in the kruger national park (knp). in most arid and semi-arid areas, water is the primary driver of ecological patterns and processes. in this article, we aimed to characterise the dominant hydropedological response of the granite catena using different levels of soil information. the objective was firstly to use soil morphological properties and relate them with soil-forming processes to deduct the dominant hydrological processes in different soils along the catena. secondly, soil chemical properties and their spatial distribution were interpreted to verify whether the morphological properties in this catena are in sync with the current climate. the last objective was to measure selected hydraulic properties to quantify dominant flow processes and mechanisms. methods study area the site is a granite catena of third-order stream in the southern granite supersite (smit et al. 2013) in the kruger national park, south africa (figure 1). the mean annual precipitation is 560 mm/a (smit et al. 2013) and the geology comprises granites and gneissic migmatites of the nelspruit suite (dippenaar & van rooy 2014; robb et al. 2006). figure 1: (a) location of the southern granite supersite in the kruger national park in south africa and (b) location of soil observations on the identified catena. the crest to the midslope is associated with gentle gradients and comprises apedal soils frequented by terminalia sericea (riddell et al. 2014). in the lower midslope positions, there is an abrupt transition to sodic duplex soils. woody vegetation dominates the crest (combretum apiculatum and combretum zeyheri), whilst acacia nilotica dominates the lower slopes and euclea divinorium is associated with the duplex sodic soils (smit et al. 2013). data collection as part of a greater study, botanists laid out 49 plots of 10 m2 from the crest of the hillslope to the valley bottom at 10-m intervals. soil observations were made to refusal using a hand auger in the centre of these plots, effectively creating a 10-m-interval transect. the soils were classified according to the soil classification working group (scwg 1991) and the international union of soil sciences working group world reference base (wrb 2015). the soil properties and their spatial distribution were interpreted and related to their hydropedological significance and reclassified into hydrological soil types (van tol & le roux 2019). soil samples were collected from the top and subsoil horizons of 49 observations. four modal profiles were identified based on terrain position and soil type. these modal profiles were located on the crest, upper footslope, lower footslope and valley bottom terrain positions (figure 2). these profiles were sampled at 100mmdepth intervals. figure 2: soil distribution pattern and location of observations, soil types and modal profiles. analysis the samples were dried, sieved and analysed according to standard methods (the non-affiliated soil analysis work committee 1990). samples were analysed for particle distribution using the pipette method and ph (h2o and kcl) with 1:2.5 solution. the soluble and exchangeable cations and cation exchange capacity were determined using nh4oac solution. selected laboratory analyses were presented as average values of the entire profiles for 49 observations as well as the vertical distribution of selected elements for the modal profiles. the s-value was used to express the total content of basic cations (expressed as equivalent values): and the exchangeable sodium percentage (esp) was calculated as follows: soil hydraulic properties were determined on three profiles representing diagnostic horizons dominating the upslope, midslope and valley bottom positions. the saturated hydraulic conductivity of each diagnostic horizon in the modal profiles was measured in duplicate using the modified bouwer and rice (1976) double-ring falling-head method: where ks is the saturated hydraulic conductivity (mm h−1), l is the thickness of the horizon, t is the time until constant infiltration and h0 and h1 are the heads of water above the surface before and after t respectively. water retention characteristics were measured for each horizon of the modal profiles on undisturbed core samples. the samples were de-aired at −70 kpa at room temperature for 24 h. water was then allowed to gradually flow out of the chambers. the saturated samples were mounted on a hanging water column setup in accordance with the procedure shown by dirksen (1999). the suction levels (h) were set at 0 mm, 38 mm, 50 mm, 100 mm, 200 mm, 400 mm, 600 mm and 800 mm. the gravimetric water content was determined at each level. the samples were then oven-dried at 105 °c to determine the bulk density to calculate the volumetric water content (θv). ethical considerations ethical approval to conduct the study was obtained from the interfaculty animal ethics committee of the university of the free state (reference no. ufs-aed2019/0121). results soil classification and morphology the soil types range from sandy and apedal soils on the crest to strongly structured soils on the footslope with alluvial deposits on the banks on the river (figure 2). the soil forms and their associated hydropedological grouping are presented in table 1. clovelly soils dominate the crest position. there is a small sodic area, covered by sterkspuit soils on the transition from the crest to the midslope position. fernwood and pinedene (arenosols) soil forms are found on the midslope. these soils are relatively deep (>750 mm) when compared to the soils on the footslope. the upper part of the footslope is dominated by the sterkspruit (solonetz) soils. the change in soil type (from sandy to clayey duplex) is correlated with a change in vegetation, that is, from woody vegetation to small shrubs. topsoil horizons of these duplex soils are shallow and often eroded completely (table 1). the lower footslope position is characterised by the pedocutanic horizons of the bonheim (luvisol) having a strongly developed blocky structure. the textural discontinuity is, however, not as dominant in these soils as in the sterkspruit soils. dundee soil (fluvisol) is found along the streambeds (valley bottom). these soils are characterised by alluvial layering. table 1: soil forms, diagnostic horizons, world reference base groups and hydropedological response types on the catena. soil chemistry and physical properties catenal distribution of soil properties profile average particle size distribution and chemical trends along the catena are presented in figure 3. drastic differences in clay and silt+clay percentages are observed between apedal horizons and the sodic zones (prismacutanic horizons; figure 3b). the upperand lower-footslope positions are marked by relatively high silt+clay contents (>30%). in the alluvial soils of the valley bottom, there is a marked decrease in the clay contents. the highest average clay content (30%) was recorded in the milkwood soil form (plot 38; figure 2) and the lowest (1.2%) in the mispah soil form of plot 7 (figure 2). the average contents of basic cations follow the same trends as the clay and silt+clay contents (figure 3c). when the impact of the clay content is neutralised, it is clear that na is dominant in the sodic sites of the upper-footslope, whereas ca is more prominent in the valley bottom (figure 3d – note the difference in scale between primary and secondary y-axis). average ph values are below 6 in the crest and midslope positions (apedal horizons) but tend to increase downslope. considerable fluctuation is observed in the ph values. the esp is relatively low in the upper sections of the slope, increases above 6% (the threshold for classification of sodic soils) in the upper-footslope position and decreases notably below the sodic sites (figure 3e). figure 3: profile of average physical and chemical properties along the catena. (a) topography/elevation of the catena, (b) particle size distribution, (c) basic cations, (d) selected cation to clay ratios and (e) ph and exchangeable sodium percentage. figure 3 (continues...): profile of average physical and chemical properties along the catena. (a) topography/elevation of the catena, (b) particle size distribution, (c) basic cations, (d) selected cation to clay ratios and (e) ph and exchangeable sodium percentage. vertical distribution of properties in modal profiles the clovelly soil forms of the crest have a relatively uniform distribution of silt+clay with depth (figure 4a). bonheim soils show a typical increase in silt+clay up to a maximum value of 53% at 550 mm depth. the topsoil of this sterkspruit soil was eroded, but the silt+clay contents were generally high (>45%). dundee soils develop from alluvial deposits and are therefore prone to textural variation, and this is reflected in figure 4(a) in the distribution of silt+clay with depth. figure 4: vertical distribution of selected properties in modal profiles. (a) silt+clay contents, (b) ph profiles and (c) s-value. the ph ranges from 5.1 in the subsoil of the clovelly to 7.8 in the pedocutanic horizon of the bonheim soil (figure 4b). ph values of the surface horizons are similar for all soil forms in figure 4(a). all profiles show an initial decline in ph with depth. this decline is more pronounced in the clovelly soil. the initial decline is followed by an increase in ph of more than 1 and 2 units in the sterkspruit and the bonheim soils respectively. the dundee soil again shows considerable variation because of the nature of alluvial deposits (figure 4b). the s-value ranges from 0.48 cmolc.kg−1 in the clovelly soil to more than 37 cmolc.kg−1 in the pedocutanic horizon of the bonheim form (figure 4c). the s-value is relatively constant throughout the clovelly soil. the s-value decreases drastically from 50 to 150 mm in the bo soil, from where it increases to the 550 mm layer. in the sterkspruit soil form, an increase in the s-value with depth is observed whereas the opposite trend is observed in the dundee soil form (figure 4c). the soluble ca contents range from 0.9 mg kg−1 in the dundee soil to 78.9 mg kg−1 in the sterkspruit soil (figure 5a). the magnitude of soluble ca profiles is within the same order as that of the clovelly, dundee and bonheim soil forms. this trend is remarkably different from the s-values calculated from the exchangeable cation concentrations. the magnitude of soluble ca profiles in the sterkspruit soil form is greater than that of the other soil forms. soluble mg profiles are very similar to that of ca profiles; however, the concentrations are lower and the difference is smaller between the sterkspruit and other soil forms (figure 5b). figure 5: soluble cation modal profiles: (a) calcium (ca), (b) magnesium (mg) and (c) sodium (na). soluble na contents range from 1.1 mg kg−1 in surface horizons of the clovelly to 6.1 mg kg−1 in the subsoil of the sterkspruit form (figure 5c). structured soils (sterkspruit and bonheim) generally have higher soluble na concentrations than the apedal soils. all soils show an increase in na with depth, although this increase is variable in the bonheim and dundee soils. hydraulic parameters the bulk density, water content at different suction levels and measured saturated hydraulic conductivity for the modal profiles indicate a large degree of variation in the physical properties (table 2). the yellow–brown apedal b (ye) horizon of the clovelly soil has the highest bulk density (1.73 g cm−3) with the lowest bulk density recorded in the soil/bedrock transition (on) horizon of the bonheim soil (1.32 g cm−3). the high bulk density of the clovelly soil translates to relatively low water content at saturation (low porosity) when compared to the higher clay content horizons. water content decreases rapidly in the sandy soils, with 0.19 mm mm−1 and 0.20 mm mm−1 available at 800 mm of suction. the water holding capacity of the structured horizons is considerably higher. at 800 mm suction, the water content in the sterkspruit soil is around 0.35 mm.mm−1 and ranges from 0.28 to 0.40 in the bonheim soil. table 2: physical measurements of three modal profiles. saturated hydraulic conductivities were very high in the clovelly soil and exceeded 50 mm h−1 for all horizons (table 2). the sterkspruit soil form had the lowest conductivity that ranges between 3 mm h−1 and 5 mm h−1. in the bonheim soil, the conductivity decreased steadily from the surface to the soil/bedrock interface with 44, 28 and 12 mm h−1 conductivity for the melanic, pedocutanic and unconsolidated horizons respectively (table 2). discussion hydropedological response based on soil morphology at the crest and midslope, the soil morphology of the sandy clovelly and mispah forms indicates that soil weathering of granitic parent material is the dominant soil-forming process. horizonisation is visible as yellowing of the b horizon and slight darkening of the orthic horizon because of accumulation of organic matter. the low clay contents indicate that silicate clay, which is formed during weathering of the granite, is eluviated (figures 3b and 4a). the yellow colour of the clovelly soils is related to the presence of iron oxides, especially goethite, which are stable and do not occur in soils saturated for significant periods (schwertmann 1985). hydrologically, it implies that a perched water table is not formed in these soils, and deep drainage into the saprolite and fracture rock is the dominant process. the occurence of a bleached (grey colours as defined in scwg 1991) b2 horizon in the pinedene soil form (table 1 and figure 2) in the lower-midslope indicates redoximorphic process in the deep subsoil. bleaching is dependent on reduction associated with energy supplied to microbes in the form of organic matter and prolonged saturation (vepraskas, huffman & kreiser 2006). in environments subjected to high evaporative demands, prolonged saturation requires that water is fed to the soil through a sustained flowpath of water. in this hillslope, this implies that flow at the soil/bedrock interface could occur in the saprolite horizon of the clovelly soil or that the water which infiltrated into the fractured rock below the clovelly and mispah soils returns to the solum under the influence of gravitation to the pinedene soil downslope. these indicators are supported by the occurrence of a fernwood soil below the pinedene soil. in simple terms, the bleached horizon at the soil/bedrock interface of the pinedene soil becomes deeper downslope until the entire subsoil is bleached. the lateral flowing water continues to accumulate until it reaches the prismatcutanic horizon in the estcourt soil. estcourt soils are known for the generation of lateral flow (van tol et al. 2013). the prismacutanic horizon acts as a clay plug and governs the lateral flowing water to return to the surface, forming a seepline. the grey colour of the e-horizon of the estcourt soil further supports this lateral flowpath. the grey colour is the result of removal of iron oxide under reducing (saturated) conditions (schwertmann 1985; van huyssteen et al. 2007). topsoil erosion, high clay contents and the presence of prismacutanic horizons with strong structure signify that overland flow is the dominant hydrological flowpath in this part of the hillslope. soil morphology indicates soil formation by clay enrichment due to either, or a combination of, physical (illuviation) or chemical (neo-formation) processes (owliaie, abtahi & heck 2006). the bonheim and milkwood soils of the lower-footslope indicate that weathering is more prominent than in the upper-footslope. angular blocky structured pedocutanic horizons are in a more advanced stage of weathering than the primary prismatic and secondary angular blocky structures of the prismacutanic soils in the upper-footslopes. the dominant flowpath is still overland flow towards the valley bottom with limited vertical flow. this is expected because of the high clay contents and strong structure of the melanic and pedocutanic horizons. dundee soils occurring in the valley bottom/riverbed are formed because of alluvial deposits from upstream. the coarse texture of the dundee soils serves as an area of infiltration and recharge of overland flow from upslope positions as well as during streamflow episodes, that is, the stream is losing water to the groundwater. the distribution of the soils in the catena indicates four distinctly different zones of soil formation related to different sets of hillslope flowpaths and storage mechanisms. firstly, soils on the crest indicate soil formation by weathering of granitic parent material with a dominant vertical flowpath recharging the underlying fractured rock. secondly, in the midslope position, soil morphology indicates subsoil saturation and associated reducing conditions. this results in gleysation and bleaching at the soil/bedrock interface because of removal of iron oxides. the dominant flowpaths are interflow at the soil/bedrock interface which returns to the surface to form a hillslope seep-line. thirdly, on the footslope, soil morphology indicates soil degradation in the form of water erosion. overland flow dominates these soils and leaching is subdominant. the strong prismatic structure of the soils of the upper-footslope is an indication that solonisation (accumulation of na) dominates. in the lower-footslope, the weathering of the prismatic structure into angular blocky structure is because of slightly wetter moisture regimes. lastly, in the valley bottom, recharge occurs in coarse materials that were deposited during flood events. hydropedological response inferred from soil chemical trends the change in subsoil chemistry from the crest to the valley bottom is correlated to changes in soil types (figures 2 and 3). low concentrations of base cations and low ph in the apedal soils at the crest and midslope positions (plots 1 to 7) indicate that eluviation is a dominant process. the almost constant low s-value (figure 4c) and soluble ca, mg and na cations (figure 5) at different depths in the clovelly soil support the deduction made from morphological properties, that is, the vertical flow through and out of these profiles is dominant. bouwer et al. (2015) found similar trends in freely drained soils in the weatherley research catchment, south africa. at plot 8, there was a marked increase in silt+clay contents and cations associated with the sterkspruit soil form. the accumulation of cations, especially ca, suggests that leaching is limited. significant increase in the ca:cl ratio (figure 3d) signifies the presence of 2:1 clay minerals formed by neo-formation (wilson 1999). very high soluble concentrations of ca, mg and na cations in the subsoil of plot 17 (ss – figure 5) suggest that a considerable amount of these cations accumulate in this ‘clay plug’. the interpretation is that lateral flow occurs at the soil/bedrock interface from plot 11 to 15, carrying dissolved cations which accumulate in the prismacutanic layer of the sterkspruit soil form. this supports the morphological interpretation that plots 11 to 15 are interflow soils. water presumably dams up behind the clay plug and, as water is lost to evapotranspiration (et), cations accumulate in the soil resulting in an increase in the ph. the change from apedal soils to structured soils (figure 2) is marked by an increase in clay contents (figure 3b) as well as an increase in esp (figure 3e). this marks the change from acidic to sodic soils. there is a change from leaching environment in the apedal soils to an accumulation in the structured soils, as clearly shown by the increase of base cations at the sodic site (note the increase in na:cl ratios as shown in figure 3d). plot 36 marks the lower boundary of the sodic site; the soils change from sterkspruit to bonheim forms (figures 2 and 3a). the relatively high ph and s-value of the surface horizon (50 mm) of the bonheim (bo) soil, compared to the 150 mm layer (figure 4), suggest that surface accumulations of cations are dominant. this indicates that overland flow water, rich in dissolved cations, will accumulate at the break of slope. below plot 36, prismacutanic horizons are degrading into pedocutanic horizons (figure 2). although the absolute concentrations of exchangeable cations are still relatively high, the cation to clay ratio decreases notably (figure 3d). this implies that vertical drainage becomes slightly more important than in the sodic soils. the combination of more gentle slopes and blocky structures (compared to prismatic) could be attributed to enhanced infiltration (kutílek 2004; leeds-harrison et al. 2007). the variation in concentrations in plots 47–49 is associated with the deposition of different materials in the dundee soil on the flood banks of the streams (figure 3). vertical variation in chemical properties is clearly depicted in figures 4 and 5, which demonstrate the variation in deposited materials. the decreasing s-value with depth suggests that leaching is dominant in this profile, also supported by the morphological interpretation of soil properties. hydraulic property support for the interpretation of morphology and pedochemistry the water contents at different suction levels and saturated hydraulic conductivity of the clovelly soil support the morphological and chemical deductions that apedal horizons are a fast flowpath (table 2). the high ks value in the orthic horizon suggests that soluble cations could easily be leached from the horizon. the steep decrease in water retention suggests that macroporosity in this horizon is high (leeds-harrison et. al. 2007). this implies that conditions for complete saturation are hardly met because of the fast drainage of macropores. the water retention data and ks of the sterkspruit soil are in accordance with morphological and chemical interpretations that water movement is very slow in the prismacutanic horizon (table 2). dispersion of clays, associated with the high na content, could clog soil pores resulting in the low conductivity. the gradual decrease in water retention of the sterkspruit indicates that macroporosity is relatively low under saturated conditions, that is, most of the water is held in the soil matrix. the water retention data and ks of the bonheim soil support the morphological and chemical interpretations that water movement is faster than that observed in the sterkspruit soil (table 2). the decrease in water content with an accompanying increase in pressure is considerately less than in the pinedene soil. the structure of the pedocutanic horizon promotes the presence of more macropores, as is evident by the water retention data. in spite of the greater number of macropores, compared to the sterkspruit, the soil is dominated by smaller pores. conceptual hydropedological response model from morphology, chemistry and hydraulic measurements the conceptual hydrological response model of this hillslope is presented in figure 6. this model was constructed from the interpretation of soil morphological properties and then refined and improved using measurements and interpretations of soil chemical and physical properties. the numbered arrows in figure 6 reflect different dominant and subdominant hydrological flowpaths/processes as discussed below: infiltration and vertical flow are dominant in the clovelly soils of the crest position. this is supported by the apedal structure, absence of grey colours, sandy texture, low ph, low base cation concentrations and high ks values. in the valley bottom, infiltration is also dominant in the streambeds and through coarse alluvial material. re-infiltration of overland flowing water is likely to occur in the gentle slopes of this terrain position. in this semi-arid environment, evapotranspiration is also dominant (see, for example, van tol et al. 2015). a great portion of infiltrated water is likely to evaporate or transpire back into the atmosphere. evapotranspiration is likely to be high in the valley bottom-associated riparian vegetation. the majority of the hillslope is covered by soils with relatively high clay contents and low hydraulic conductivity. these soils are also characterised by a high base status and ph, which is associated with limited leaching. overland flow is considered to be the dominant hydrological process. erosion of surface horizons is additional confirmation that overland flow dominates in the upperand lower-footslope positions. the presence of hydromorphic b2 horizons suggests that saturated conditions exist during wet periods. because these hydromorphic signatures become more prominent downslope (up to the start of the sodic site), this is an indication that lateral flow occurs here. this is supported by an increase in soluble base cations in a downslope direction. when this lateral flowing water in (4) reaches the clay-enriched sterkspruit soils, the water accumulates until the storage capacity of the soils is exceeded and exfiltration in the form of a seepage face takes place. this return flowpath contributes to overland flow. infiltration into fractures of the granitic parent material (recharge) is possible, although considered a subdominant process in the greater part of this hillslope. on the crest and in the valley bottom, significant recharge could take place during extreme rainfall events. figure 6: conceptual hydrological response model of a third-order stream in the kruger national park. vertical axis exaggerated for display purposes. conclusion terrain morphology is a common representative of homogeneous ecological units in pristine granite landscapes of the kruger national park. this catenal effect is an interaction between water, topography and geology as primary components and relates to a hydropedological composition. this composition is both an indication of partitioning and a control factor of movement and storage of water in the catena. soil morphology is strongly associated with the hydrological response. the degree to which chemical properties are in sync with current pedological process emphasises the fact that the soils are mature and chemistry and morphology are useful indicators of hydrological response. chemistry must be interpreted from several observations and with depth, as it responds to the environmental impact within the phase of pedogenesis. chemical indicators of leaching were used as indicators for flowpaths, although the prediction of flow direction depends on slope and indicators of return flow. the accumulation of cations was an indication of a slow flowpath. low subsoil ph, and therefore low cation concentration, was used as an indicator of ferrolysis because of periodic saturation. the accumulation of cations in apedal horizons improved morphological interpretation, as the horizons did not respond as fast flowpaths but rather as a storage mechanism induced by the poor drainage of the underlying rock. the chemical properties confirmed the degrading of sterkspruit to bonheim soil. the melanic and pedocutanic horizons had chemical trends similar to orthic and prismacutanic horizons, except in lower concentrations. the c horizon of the bonheim soil has similar chemical properties as those of the prismacutanic horizons. pedon scale hydropedology contributed to the understanding of the physical catena and predicted a strong correlation with the biological catena. the use of morphology, chemistry and physics resulted in improving the conceptual understanding of a conceptual model and soil types. thus, the technique to use soil chemical properties to identify flowpaths in soils has improved the conceptual hydrological models compared to those where only soil morphology was used. acknowledgements competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions d.b. and p.a.l. led the research design and conducted fieldwork. d.b. was responsible for lab and data analysis. j.v.t. and d.b. were responsible for data interpretation and the write-up of the article. funding information the authors are grateful to the university of the free state for funding this multidisciplinary research and the water research commission (wrc) of south africa under project number k5/2051. data availability data are available upon request to the corresponding author. data from all research done within kruger national park is placed within the sanparks repository (not for free, open access). disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references bouwer, d., le roux, p.a., van tol, j.j. & van huyssteen, c.w., 2015, ‘using ancient and recent soil properties to design a conceptual hydrological response model’, geoderma 241–242, 1–11. https://doi.org/10.1016/j.geoderma.2014.10.002 bouwer, h. & rice, r.c., 1976, ‘a slug test for determining hydraulic conductivity of unconfined aquifers with completely or partially penetrating wells’, water resources research 12(3), 423–428. https://doi.org/10.1029/wr012i003p00423 dippenaar, m.a. & van rooy, j.l., 2014, ‘review of engineering, hydrogeological and vadose zone hydrological aspects of the lanseria gneiss, goudplaats-hout river gneiss and nelspruit suite granite (south africa)’, journal of african earth sciences 91, 12–31. https://doi.org/10.1016/j.jafrearsci.2013.11.019 dirksen, c., 1999, soil physics measurements, catena verlag, reiskirchen. essington, m.e., 2004, soil and water chemistry: an integrative approach, crc press, new york, ny. international union of soil sciences working group world reference base (wrb), 2015, world reference base for soil resources 2014: international soil classification system for naming soils and creating legends for soil maps, world soil resources reports no. 106, food and agriculture organization of the united nations, rome. kuenene b.t., le roux p.a.l., van huyssteen c.w. & hensley m., 2013, ‘selected soil properties as indicators of soil water regime in the cathedral peak vi catchment of kwazulu-natal, south africa’, south african journal of plant & soil 30(1), 1–6. https://doi.org/10.1080/02571862.2013.767387 kutílek, m., 2004, ‘soil hydraulic properties as related to soil structure’, soil tillage research 79(2), 175–184. https://doi.org/10.1016/j.still.2004.07.006 kutílek, m. & nielsen, d.r., 2007, ‘interdisciplinarity of hydropedology’, geoderma 138(3–4), 252–260. https://doi.org/10.1016/j.geoderma.2006.11.015 leeds-harrison, p.b., shipway, c.j.p., jarvus, n.j. & youngs, e.g., 2007, ‘the influence of soil macroporosity on water retention, transmission and drainage in a clay soil’, soil use and management 2(2), 47–50. https://doi.org/10.1111/j.1475-2743.1986.tb00678.x lin, h.s., 2003, ‘hydropedology: bridging disciplines, scales, and data’, vadose zone journal 2(1), 1–11. https://doi.org/10.2136/vzj2003.1000 lin, h.s., 2010, ‘earth’s critical zone and hydropedology: concepts, characteristics, and advances’, hydrology and earth system sciences 14(1), 25–45. https://doi.org/10.5194/hess-14-25-2010 lin, h.s., bouma, j., pachepsky, y., western, a.w., thompson, j., van genuchten, r. et al., 2006a, ‘hydropedology: synergistic integration of pedology and hydrology’, water resources research 42(5), 1–13. https://doi.org/10.1029/2005wr004085 lin, h.s., kogelmann, w., walker, c. & bruns, m.a., 2006b, ‘soil moisture patterns in a forested catchment: a hydropedological perspective’, geoderma 131(3–4), 345–368. https://doi.org/10.1016/j.geoderma.2005.03.013 lin, h.s., wheeler, d., bell, j.c. & wilding, l.p., 2005, ‘assessment of soil spatial variability at multiple scales’, ecological modelling 182(3–4), 271–290. https://doi.org/10.1016/j.ecolmodel.2004.04.006 macewan, r.j., 1997, ‘soil quality indicators: pedological aspects’, in e.g. gregorich & m.r. carter (eds.), soil quality for crop production and ecosystem health, pp. 143–166, elsevier, new york, ny. macewan, r.j. & fitzpatrick, r.w., 1996, ‘the pedological context for assessment of soil quality’, in r.j. macewan & m.r. carter (eds.), soil quality is in the hands of the land manager. proceedings of an international symposium on advances in soil quality for land management: science, practice and policy, pp. 10–16, university of ballarat, ballarat. mamera, m. & van tol, j.j., 2018, ‘application of hydropedological information to conceptualise pollution migration from dry sanitation systems in the ntabelanga area, south africa’, air, soil and water research 11, 1–12. https://doi.org/10.1177/1178622118795485. owliaie, h.r., abtahi, a. & heck, r.j., 2006, ‘pedogenesis and clay mineralogical investigation of soils formed on gypsiferous and calcareous materials, on a transect, south-western iran’, geoderma 134(1–2), 62–81. https://doi.org/10.1016/j.geoderma.2005.08.015 riddell, e.s., lorentz, s.a. & kotze, d.c., 2012, ‘the hydrodynamic response of a semi-arid headwater wetland to technical rehabilitation interventions’, water sa 38(1), 55–66. https://doi.org/10.4314/wsa.v38i1.8 riddell, e.s., nel, j., fundisi, d., jumbi, f., van niekerk, a. & lorentz, s., 2014, ephemeral hydrological processes in savannas, water research commission report no. tt619/14, water research commission, pretoria. robb, l.j., brandl, g., anhaeusser, c.r. & poujol, m., 2006, ‘archaean granitoid intrusions’, in m.r. johnson, c.r. annhaeusser & r.j. thomas (eds.), the geology of south africa, pp. 57–94, geological society of south africa, johannesburg. schwertmann, u., 1985, ‘the effect of pedogenic environments on iron oxide minerals’, advances in soil science 1, 171–200. https://doi.org/10.1007/978-1-4612-5046-3_5 smit, i.p.j., riddell, e.s., cullum, c. & petersen, r., 2013. ‘kruger national park research supersites: establishing long-term research sites for cross-disciplinary, multiscaled learning’, koedoe 55(1), art. #1107, 7 pages. https://doi.org/10.4102/koedoe.v55i1.1107 soil classification working group (scwg), 1991, soil classification: a taxonomic system for south africa, department of agricultural development, pretoria. the non-affiliated soil analysis work committee, 1990, handbook of standard soil testing methods for advisory purposes, soil science society of south africa, bloemfontein. van huyssteen c.w., le roux p.a.l., hensley m. & zere t.b., 2007, ‘duration of water saturation in selected soils of weatherley, south africa’, south african journal of plant & soil 24(3), 152–160. https://doi.org/10.1080/02571862.2007.10634799 van tol, j.j., hensley, m. & le roux, p.a., 2013, ‘pedological criteria for estimating the importance of subsurface lateral flow in e horizons in south african soils’, water sa 39(1), 47–56. https://doi.org/10.4314/wsa.v39i1.7 van tol, j.j. & le roux, p.a.l., 2019, ‘hydropedological grouping of south african soil forms’, south african journal of plant and soil. 36(3), 233–235. https://doi.org/10.1080/02571862.2018.1537012. van tol, j.j., le roux, p.a., lorentz, s.a. & hensley, m., 2010, ‘soil as indicator of hillslope hydrological behaviour in the weatherley catchment, eastern cape, south africa’, water sa 36(5), 513–520. https://doi.org/10.4314/wsa.v36i5.61985 van tol, j.j. & lorentz, s.a., 2018, ‘hydropedological interpretation of soil distribution patterns to characterise groundwater/surface-water interactions’, vadose zone journal 17(1), 1–13. https://doi.org/10.2136/vzj2017.05.0097 van tol, j.j., lorentz, s.a., van zijl, g.m. & le roux, p.a.l., 2018, ‘the contribution of hydropedological assessments to the availability and sustainable water, for all (sdg#6)’, in r. lal, r. horn & t. kosaki (eds.), soil and sustainable development goals, pp. 102–117, catena-schweizerbart, stuttgart. van tol, j.j., van zijl, g.m., riddell, e.s. & fundisi, d., 2015, ‘application of hydropedological insights in hydrological modelling of the stevenson hamilton research supersite, kruger national park, south africa’, watersa 41(4), 525–533. https://doi.org/10.4314/wsa.v41i4.12 van zijl, g.m., van tol, j.j. & riddell, e.s., 2016, ‘digital mapping for hydrological modelling’, in g. zhang, d. brus, f. liu, x. song & p. lagacherie (eds.), digital soil mapping across paradigms, scales and boundaries, pp. 115–129, springer, singapore. vepraskas, m., huffman, r.l. & kreiser, g.s., 2006, ‘hydrologic models for altered landscapes’, geoderma 131(3–4), 287–298. https://doi.org/10.1016/j.geoderma.2005.03.010 wilson, m.j., 1999, ‘the origin and formation of clay minerals in soils: past, present and future perspectives’, clay minerals 34(1), 7–25. https://doi.org/10.1180/000985599545957 filelist convert a pdf file! page 1 page 2 page 3 page 4 687-table1-21 july 2009.indd [final table vesion].indd a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe online supplement table 1 a rticle #687 1vol. 51 no. 1 page 1 of 4 table 1 phytosociological table of the plant communities of the adansonia digitata–acacia nigrescens soutpansberg arid northern bushveld major vegetation type koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za online supplement table 1 mostert, bredenkamp & mostert a rt ic le # 68 7 2 vol. 51 no. 1 page 2 of 4 table 1 (cont.) phytosociological table of the plant communities of the adansonia digitata–acacia nigrescens soutpansberg arid northern bushveld major vegetation type plant communities of the soutpansberg arid northern bushveld online supplement table 1 a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #687 3vol. 51 no. 1 page 3 of 4 table 1 (cont.) phytosociological table of the plant communities of the adansonia digitata–acacia nigrescens soutpansberg arid northern bushveld major vegetation type online supplement table 1 mostert, bredenkamp & mostert koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 68 7 4 vol. 51 no. 1 page 4 of 4 table 1 (cont.) phytosociological table of the plant communities of the adansonia digitata–acacia nigrescens soutpansberg arid northern bushveld major vegetation type filelist convert a pdf file! page 1 page 2 page 3 page 4 abstract introduction methods results: alternate stable states in savannas discussion and conclusions: implications for ecosystem services and management options acknowledgements references appendix 1 about the author(s) lindsey gillson plant conservation unit, department of biological sciences, university of cape town, cape town, south africa anneli ekblom department of archaeology and ancient history, uppsala university, uppsala, sweden citation gillson l. & ekblom, a., 2020, ‘using palaeoecology to explore the resilience of southern african savannas’, koedoe 62(1), a1576. https://doi.org/10.4102/koedoe.v62i1.1576 review article using palaeoecology to explore the resilience of southern african savannas lindsey gillson, anneli ekblom received: 20 june 2019; accepted: 18 mar. 2020; published: 15 june 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract savannas are dynamic and heterogeneous environments with highly variable vegetation that responds to a multitude of interacting drivers. rainfall, soils, herbivory, fire and land use all effect land cover in savannas. in addition, savannas have a long history of human use. setting management goals is therefore complex. understanding long-term variability in savannas using palaeoecology provides a context for interpretation of recent changes in vegetation and can help to inform management based on acceptable or historical ranges of variability. in this article, we review and synthesise palaeoecological data from southern african savannas and use resilience theory as a framework for structuring and understanding of vegetation dynamics in savannas. we identify thresholds between alternate stable states, which have different ecological properties, suites of species and ecosystem services. multi-proxy palaeoecological records can assist in identifying alternate states in savanna vegetation, as well as showing how different drivers (fire, herbivory, nutrients and climate) interact to drive transitions between states. conservation implications: the ecological thresholds identified from palaeoecological data can be used to inform the development of management thresholds, known as thresholds of potential concern. thresholds of potential concern are designed to facilitate or impede transitions between states by manipulation of those variables (e.g. fire and herbivory) that can be controlled at the landscape scale. keywords: resilience; thresholds; thresholds of potential concern; palaeoecology; alternate stable states. introduction in savannas, interactions between fire, herbivory and climate can alter ecosystem structure and function, potentially resulting in shifts between grassand tree-dominated states (hempson et al. 2019; sankaran, ratnam & hanan 2008). these transitions may occur on any spatial scale from local (hundreds of metres) to regional (subcontinental) and timescales of decades to centuries. the heterogeneity and dynamism of savanna vegetation and the complexity of drivers and scales raise challenges both theoretically and practically, especially in view of climate change and possible vegetation responses. resilience theory provides a framework for integrating the knowledge of savanna dynamics that can be applied in biodiversity conservation and ecosystem management, as will be explored in this article. two aspects of resilience are commonly described in the literature: (1) ecological resilience – which is defined as the capacity of an ecosystem to withstand disturbance without changing structure and function and (2) engineering resilience – which is defined as the capacity of an ecosystem to recover from a disturbance that causes change in structure and function (holling 1996; oliver et al. 2015; sundstrom et al. 2016). understanding of holocene vegetation dynamics can elucidate how interactions between fire, herbivory and climate alter ecosystem structure and function over different timescales (venter, hawkins & cramer 2017) – knowledge that is fundamental for the formulation of management strategies today and in the future. in addition, studies of areas with different management histories illustrate possible effects of different combinations of fire and herbivory over time. here we will focus on palaeoecological sequences from three protected areas – kruger national park (knp) and hluhlwe-imfolozi parks (hip) in south africa and limpopo national park (pnl) in mozambique – to analyse how different drivers (e.g. fire, herbivory, nutrients and climate) interact to drive transitions between alternate states in savanna vegetation. the focus on protected areas is motivated by the specific challenges of conservation management, in defining the range of variability that can sustain biodiversity without leading to irreversible or undesirable shifts in vegetation with changing environmental parameters (rogers 2003). non-equilibrium ecology and resilience thinking have led to innovations in savanna management and conservation, with the emergence of thresholds of potential concern (tpcs) as operational goals that define the upper and lower limits of acceptable change in ecosystem parameters (rogers 1997, 2003; rogers & o’keeffe 2003). the formulation of tpcs in defining spatiotemporal heterogeneity raises both ecological and social questions (biggs et al. 2011). most of africa’s parks and protected areas were founded under atypical conditions that nevertheless continue to shape expectations to this day. hunting of large herbivores and carnivores, as well as policies of fire suppression and outbreaks of disease such as rinderpest during the 18th–19th century, likely transformed ecosystem structure and function, yet we have limited knowledge of how ecosystems looked prior to this time. stakeholder preferences are informed also by how they perceive current changes in ecosystem state relative to their own personal baselines. we have argued elsewhere (forbes et al. 2018) that baselines shift depending on personal experience and the lifespan of projects, and highlight the valuable role of long-term data in contextualising change that has occurred over recent decades. palaeoecological data may therefore help managers and ecologists to envisage scenarios to help that are outside of their personal experience (gillson & marchant 2014; gillson 2015a). in this article, we interpret changes in vegetation structure inferred from palaeoecological data in terms of resilience and thresholds, and attempt to identify the range of factors that drive transitions between alternate stable states, or which promote resistance and stability of vegetation, despite environmental variability. to translate the palaeoecological data into a form that can be utilised in ecosystem management, we then compare long-term (palaeo) ecological thresholds with management thresholds (tpcs). we anticipate that such a framework could inform future quantitative analysis and modelling that could allow savanna managers and conservationists to explore the effects of manipulating fire and herbivory under various scenarios of future rainfall, temperature and carbon dioxide (co2). methods the basis for our discussion is data generated from multiple palaeoecological records from knp and hip in south africa, and limpopo national park in mozambique (pnl [see figure 1 and table 1–a1]) (ekblom & gillson 2010a, 2010b, 2010c; ekblom, gillson & notelid 2011; ekblom et al. 2012; gillson 2015b; gillson & duffin 2007; gillson & ekblom 2009, 2009b). the period discussed here, the last ca. 6000 years, has experienced both periods of warming and cooling combined with shifts in rainfall (scott 2016; scott et al. 2012). the time period covers the end of the mid-holocene warming 4000 years ago when there was an increase in rainfall in the summer rainfall region. this was followed by a period of variable but overall cooler and drier conditions. overall wetter conditions took place ca. 1200–800 years ago associated with globally warmer climate (sometimes referred to as the medieval warming) (holmgren et al. 2003; humphries, green & finch 2016; lee-thorp et al. 2001; neumann, botha & scott 2014; scott & lee-thorp 2004; scott et al. 2012; scott 2016). shorter cool–dry events took place in ad 1100, 1350, 1450 and 1500 interspersed with periods of higher rainfall. a prolonged trend of cool–dry conditions took place after ad 1600, especially severe at ad 1700, with a return of more rainfall only 200 years ago (holmgren et al. 2003, 2012; lee-thorp et al. 2001; scott et al. 2012; sundqvist et al. 2013; woodborne et al. 2015, 2016). although the dynamics of these changes, and if these climate shifts are analogous to future climate shifts, are still debated (fitchett et al. 2017), the knowledge of the scale and rate of vegetation changes are important to understand for informed management decisions. figure 1: map showing location of sediment cores and protected areas mentioned in the text. changes in vegetation are here reconstructed using the relative changes of pollen types in dated sediments. transitions from grass-dominated to woody vegetation are studied using pollen from sediments and stable carbon isotopes in soil profiles. the pollen record is compared with the amount of charcoal as an indicator of fire frequency/intensity and dung loving spores as a possible indicator of herbivore abundance, while nitrogen availability is also reconstructed based on δ15n isotopes. transitions from grass-dominated to woody vegetation can also be studied using stable carbon isotopes in soil profiles. grasses and trees in savannas use different photosynthetic pathways, resulting in distinctive isotopic signatures, where grasses are enriched in heavier carbon. by analysing δ13c in soil profiles, changes in the relative abundance of grasses and trees over time can be estimated, with more recent vegetation reflected in the uppermost soil or sediment profiles. although the palaeoecological records from small wetlands, pans and swamps presented here reflect local changes, and sampling has been limited by suitable sediment deposits and available proxies, we are still able to construct a general understanding of vegetation dynamic in terms of the factors that might drive transitions between stable states, or that promote resistance or resilience. ethical considerations this article followed all ethical standards for a research without direct contact with human or animal subjects. results: alternate stable states in savannas water-limited grassland the most stable vegetation states identified in our pollen records are water-limited grasslands. although annual rainfall (600 mm/year) is presently sufficient for supporting a more dense woody cover, trees are scarce or absent because of water limitations imposed by local hydrology (i.e. lack of connection to ground water fed by perennial rivers). such a landscape is exemplified by radio pan in pnl (ekblom & gillson 2010c). the resistance or resilience of this vegetation state is demonstrated by the lack of response of tree cover (represented by arboreal pollen) to climate, fire and nutrients. on the basis of the time series data, we did simple regression analyses between tree cover, charcoal (fire influence), dung loving spores (possible grazing influence) and δ15n isotopes (nitrogen availability). we expected arboreal pollen abundance to have a significant negative relationship with spores and charcoal, reflecting the negative impact of herbivory and fire on tree recruitment. we expected to find a positive correlation between nitrogen availability and tree abundance, reflecting a positive relationship between tree abundance and nitrogen. however, at radio pan, there was no significant correlation between arboreal pollen and nitrogen availability (based on δ15n), fire (based on charcoal abundance) or herbivory (based on dung fungal spore abundance) (figure 2a) (ekblom & gillson 2010c). in contrast, a nearby site, chixuludzi, with connections to perennial water, that is, limpopo river, showed significant correlations between tree abundance, fire and nitrogen availability (figure 2b) (ekblom & gillson 2010c). figure 2: summary diagrams of the radio (a) and chixuludzi (b) sedimentary sequences. colored silhouettes show distribution of the main vegetation groups in percentages based on terrestrial pollen (c. 300–500 pollen); black bars show charcoal concentration (estimated as concentration cm2/cm3). in the radio diagram (a) the red lines shows the distribution of coprophilous dung fungal spores (converted in percentages in relation to the pollen sum) and the blue lines changes in the δ15n values. the blue line shown in the chixuludzi diagram (b) shows changes in the δ15n values and the grey line the distribution of coprophilous dung fungal spores (converted in percentages in relation to the pollen sum). the complete dominance of grassland surrounding radio pan persisted throughout a 600-year history, despite changes in rainfall and in contrast to chixuludzi, which showed variations in tree cover over time (figures 2a and 2b). we suggest that at radio pan, water is the limiting factor that constrains tree recruitment, meaning that changes in fire and herbivory had no effect on tree abundance. it is not clear from the pollen data whether the resistance of grassland around radio pan is conferred by persistence of grassland composition or by internal dynamism (i.e. turnover of grass species within the grass community). recent work from highland grasslands mpumalanga (breman et al. 2019), based on phytoliths, suggests that resilience of grasslands might be enhanced through internal turnover within the grassland community, which can buffer the effects of changing climate, thereby enhancing resistance. further analysis of phytoliths and biomarkers from water-limited grasslands such as radio pan might help to resolve this question. herbivore-limited grassland (grazing lawn) transitioning to wooded savanna grazing lawns are maintained by high levels of herbivory, especially by mega-herbivores such as white rhino, which limit tree recruitment and also prevent fire by reducing grass biomass levels below those which can support fire (waldram, bond & stock 2007). herbivores can contribute to fast nutrient cycling through addition of faeces and urine to the soil, although they may also promote competition for nitrogen by removal of biomass. competition for nitrogen can also help maintain grazing lawns as grasses grow quickly and the rapid uptake of nitrogen makes them the superior competitor, able to exclude trees. these grazing lawns can persist over long time periods but might be de-stabilised by sudden losses of herbivory (e.g. because of drought, disease or over-hunting) (staver & bond 2014; waldram et al. 2007). decreased herbivory in combination with increased water availability could therefore facilitate transitions from grazing lawns to wooded (open-canopy) savannas, especially if increased water in the landscape allowed dispersal of water-dependent herbivores, reducing pressure on vegetation around water sources (redfern et al. 2005). we observed this phenomenon in the pollen record from malahlapanga (knp), our oldest record covering a long period of climate variability. here, an open grassland (indicated by very scarce tree pollen and absence of charcoal abundance) transitioned to fire-prone wooded savanna during the period of increasing rainfall around 800 years ago (figure 3a) (gillson & ekblom 2009a). figure 3: summary diagrams of the malahlapanga (a) and the mapimbi (b) sedimentary sequences. colored silhouettes show distribution of the main vegetation groups in percentages based on terrestrial pollen (c. 300-500 pollen); black bars show charcoal concentration (estimated as concentration cm2/cm3). the blue lines in the malahlapanga sequence (a) show the δ15n values and the red lines the c/n (eg. carbon to nitrogen) ratio. as mega-herbivores such as rhino continue to decline in african savannas because of illegal hunting (bennett 2014; lubbe et al. 2017; wittemyer et al. 2014), we might expect the number of grazing lawns to decrease (archibald et al. 2005; waldram et al. 2007). beyond a threshold of woody plant density, densely wooded areas may also create barriers for fire by shading out grass biomass and creating discontinuities in the fuel layer, which would further enhance recruitment of woody taxa (higgins & scheiter 2012). if the atmospheric forcing resulting from global warming is analogous to the past, we would expect more rainfall with higher temperatures. however, current climate models suggest overall drier conditions with higher incidence of extreme rainfall events (engelbrecht, engelbrecht & dyson 2013; engelbrecht, mcgregor & engelbrecht 2009; pinto, jack & hewitson 2018) and/or more unpredictability (haensler 2010; haensler, hagemann & jacob 2011a; pinto et al. 2016). regardless of rainfall changes, the added effects of co2 fertilisation (see below) might further enhance the competitiveness of trees, shifting the balance away from open grassland habitats (bond & midgley 2012). unstable wooded savannas the above examples are savannas where open grass cover is persistent as trees are constrained either by water availability, herbivory and/or nitrogen availability. below we will describe savannas that are potentially unstable and that can change from open grassland to wooded savanna because of one or the combination of drivers. on a landscape scale, savannas are constituted by patches of dense or lightly wooded areas interspersed with grasslands. overall, the stability of grass or woody cover is maintained when the number of dense and lightly wooded patches remains constant at the landscape scale, although each patch will go through asynchronous cycles of tree recruitment, thickening and senescence, a process known as patch dynamics (see gillson 2004a). however, drivers such as dramatic, region-wide changes in herbivore density might lead to mass tree recruitment, as occurred at the end of the 19th century during the rinderpest pandemic (staver & bond 2014). furthermore, policies of fire suppression might also facilitate tree recruitment, as was the case with the fire suppression policy implemented in south africa during the early 20th century (e.g. carruthers 1995; van wilgen et al. 2004). in both of these examples, the decline in herbivory and fire suppression resulted in a mass recruitment of woody cover, which led to an overall loss of patch structure through the development of even-aged tree or shrub stands. the increase in woody cover could limit the effects of fire by creating discontinuity in the fuel (grass biomass) layer, further enhancing the possibilities for woody recruitment. we see evidence for mass closure of savanna tree canopies, for example, in stable isotope data from soil profiles in hip, where δ13c consistently decreases in the upper layers of the soil profile, indicating a transition from mixed c3–c4 vegetation to c3-dominated vegetation over time (gillson 2015b). furthermore, when the frequency distribution of δ13c is analysed, the stable isotope data show some evidence of bimodal distribution, clustered around δ13c values typical of grasses and trees. this bimodality is a possible signature of transition between alternate stable states; if woody cover was a continuum between more and less grassy landscapes, we would expect the stable isotope data to be normally distributed and not bimodally distributed as here (gillson 2015b). similarly, analysis of aerial photographs covering the period from the 1930s to the 21st century shows consistent increase in tree canopy cover across different land-use types that differ in grazing and fire management history (conservation area, communally managed farm and commercial farm), providing evidence for a global rather than a local driver (wigley, bond & hoffman 2009, 2010). wooded savanna transitioning to closed canopy forest while savanna trees are able to persist in fire-prone environments through re-seeding or re-sprouting, forest trees cannot tolerate fire. closure of tree canopies in savannas will shade out the herbaceous layer, potentially excluding fire and allowing fire-sensitive, shade-tolerant forest trees to penetrate (charles-dominique et al. 2015; higgins & scheiter 2012; hoffmann et al. 2012). once these trees have established, they would eventually replace the shade-intolerant savanna species and would continue to exclude fire through shading out of the herbaceous layer. this feedback between vegetation and fire exclusion will continuously promote forest trees and associated taxa into an alternate stable state (see figure 5). because perennial rivers in savannas are typically bordered by riverine forests, and patches of true forest can co-exist alongside wooded savanna, we might expect the closure of tree canopies in savannas to lead to transitions to forest vegetation that are difficult to reverse. such a hysteretic mechanism has been modelled using an adaptive dynamic global vegetation model (adgvm) and is predicted to lead to an expansion of forest biomes at the expense of savanna and grassland biomes by 2100 (higgins & scheiter 2012). forest transitions would be difficult to reverse because forest ecosystems maintain themselves through the feedback mechanism of shading and fire exclusion, as described above (staver, archibald & levin 2011). a transition back to open savanna might be expected only in the case of severe fire-storms, which are rare events where intense fires spread to the forest canopy (archibald et al. 2017; charles-dominique et al. 2015). another situation is in case of very extreme and prolonged or repeated droughts which may kill off trees. at mapimbi, in the extreme north of knp, forest taxa declined during the 18th century, which has been connected with the extreme droughts in this period (figure 3b) (ekblom et al. 2012; holmgren et al. 2003, 2012; lee-thorp et al. 2001; scott et al. 2012; sundqvist et al. 2013; woodborne et al. 2015, 2016). an analogous situation is where humans clear patches of forest, for example, for crop cultivation, which may interact with drought effects; however, there is little evidence of historical forest clearing on a larger scale (ekblom et al. 2012; gillson & ekblom 2009b). riverine forest recovered within a century from these climatically induced disturbances. riverine forest also showed relatively rapid recovery over a few decades following the extreme flood events in the 20th century (o’connor 2010; rountree, rogers & heritage 2000). even though riverine forests in mapimbi recovered, the centennial scale of recovery from droughts is problematic from a managerial point of view and may have far reaching consequences that are difficult to predict because of the uncertainty in the climate model projections. again, negative effects on forest vegetation from more droughts or more catastrophic events may be countered by co2 fertilisation, but more data are needed to assess this. discussion and conclusions: implications for ecosystem services and management options the palaeoecological record shows evidence of different alternate stable states and mechanisms of resilience and resistance in the vegetation of southern african savannas (figure 4a). the most resistant form of vegetation that we encountered in our data is water-scarce open grasslands. these are maintained by a combination of local hydrology and fire despite climate variability. water scarcity overrides other factors such as fire, herbivory and nutrients (beckage et al. 2019) that we would normally expect to correlate with tree abundance in landscapes that are less water limited (e.g. see gillson & ekblom 2009a). the effects of global warming on rainfall patterns currently are debated (fitchett et al. 2017) and there is no consensus here other than that conditions will become more unpredictable (see above). thus, we will leave open the question of effects of rainfall changes to look at other dynamics in this discussion. however, the managerial decisions discussed below will be relevant also for observed effects of rainfall changes. figure 4: (a) hypothetical model showing transitions between alternate states with increasing plant available moisture, showing the role of vegetation-fire feedbacks in maintaining alternate stable states. (b) alternate stable states in savanna landscapes as a function of tree density and water availability, indicating both ecological and management thresholds. ecological states are bounded by solid lines. dotted lines represent management thresholds (tpc) where tree density could be manipulated through changes in fire or ivory. three grassland states are possible, depending on levels of water availability. they could transition to fire-maintained wooded savannas if water availability increased and/or herbivory decreased. wooded savannas could transition to forest ecosystems if canopies closed leading to the exclusion of fire. management intervention in the form of manipulation of fire and herbivory could be triggered when ecosystems reach the tpc. grazing lawns are open habitats similar to water-scarce grasslands, but they are maintained by mega-herbivores, particularly white rhino. their continued existence will therefore be threatened by the ongoing illegal killing of rhino (lubbe et al. 2017; waldram et al. 2007). the loss of open, grassy landscapes has implications for grazers and other flora and fauna adapted to these environments (higgins & scheiter 2012; parr et al. 2014). reduction in open grazing lawns may also be detrimental to wildlife tourism as it will decrease game viewing potential. management would need to focus on monitoring of open areas, through on the ground and remotely sensed data, and rapid management action where tree encroachment reaches a tpc (gillson & duffin 2007; rogers & biggs 1999) (figure 4b). for example, intense fires have been shown to effectively control shrub encroachment (van wilgen et al. 2014) and could be used alongside increased herbivore density in an attempt to curb tree recruitment. furthermore, the capacity to support mega-herbivores may increase under conditions of high co2 (scheiter & higgins 2012). the most variable and unstable type of vegetation that we found in the palaeoecological record is the wooded, heterogeneous savannas that typify most of the savanna biome. these landscapes can exist in a multitude of forms with varying levels of tree cover (e.g. see sankaran et al. 2005, 2008). from a conservation management perspective, there are many ecologically possible options, and decisions about the desirable state of these landscapes will have different trade-offs, for example, between carbon storage and game viewing and/or grazing mentioned above. decisions will therefore need to consider the perspectives of multiple stakeholder groups (biggs et al. 2011). in this context, it is also important to consider the landscape history of individual sites and the acceptance of landscape transformation (gillson 2015a; gillson & marchant 2014). woody plant thickening may be seen as undesirable in ancient grassland sites (parr et al. 2014), but may be tolerated in sites that have been historically cleared but which are recovering because of changing land uses, for example, land abandonment. therefore, site-specific tpcs would be needed, where stakeholders decide on the acceptable limits of change in woody cover, depending on the landscape history and their land-use needs. the degree of change in vegetation cover in savannas over decadal to millennial timescales is poorly understood. yet, in deciding management thresholds, it is important to understand how the vegetation cover has changed before, during and after the main environmental and historical events that have shaped the southern african region since the mid-holocene. these include changes in human land use, including the arrival of pastoralists from ca. 2000 years ago and increased management of fire, as also warmer conditions of the mediaeval warm period, the little ice age. more recent changes that may have influenced vegetation changes are the growth of the international trade in ivory as well as the rise of the use of firearms in hunting for sport and trade, colonial settlement and associated changes in faunal assemblages (e.g. extermination of predators, local extirpation of elephants) and land management (e.g. replacement of traditional fire management with policies of fire suppression) (carruthers 1995; holmgren et al. 2001, 2003, van wilgen et al. 2004). in addition to this, there are 20th-century changes in warming temperatures, changing rainfall patterns, co2 fertilisation, which interact with the effects of past and present land-use changes, including the effects of urbanisation, land abandonment and the growth of the private conservation sector. by examining the historical range of variability under different climatic and land-use scenarios, stakeholders can decide on the acceptable range of variability and thereby better define the tpcs in tree cover, which is appropriate to their local context (figure 4b) (biggs et al. 2011; gillson & duffin 2007). in the future, the resistance of these landscapes might be compromised by the effects of co2 fertilisation, which will increase the growth rates of trees, allowing more rapid access to water in deep soil layers and enhancing the water use efficiency of c3 savanna trees to the detriment of the grasses (bond & midgley 2012; midgley & bond 2015). from the mid-20th century, increasing co2 has had a fertilising effect on savanna trees, causing them to grow more rapidly, making them (1) less vulnerable to drought, as roots reach moist soil layers more quickly and (2) more able to escape the fire trap – that is, they can quickly reach a height that is higher than the flame zone, allowing them to recruit to larger size classes (midgley & bond 2015). in the absence of co2 fertilisation, savanna tree seedlings can become trapped in the herbaceous layer, being repeatedly burned to the ground and re-sprouting only from their root stock (bond & midgley 2012). with faster growth under higher co2 conditions, trees can grow more rapidly, escape the fire trap and join the ranks of adult tree classes (midgley & bond 2015). this effect is currently being exacerbated by escalating illegal hunting of mega-herbivores, historically important in keeping tree canopies open (bennett 2014; lubbe et al. 2017; wittemyer et al. 2014). riverine forest responds to changing climatic and disturbance drivers but, as suggested from the mapimbi record discussed above, recovers from disturbance over decades (e.g. ekblom et al. 2012; gillson & ekblom 2009b). although more records are needed to analyse this process in detail, similar observations have been made in connection recovery of riverine forest after the 20th-century flood events as discussed above. this resilience is likely to be further enhanced by co2 fertilisation, which will facilitate rapid colonisation by trees. these gallery forests and other forest patches could act as a reservoir of forest species that could colonise closed canopy savannas, leading to hysteretic transitions to forest cover, as predicted by higgins and scheiter (2012) (figure 5). however, the effects of increasingly severe drought and/or more seasonal rainfall (davis-reddy & vincent 2017; engelbrecht & engelbrecht 2016; haensler, hagemann & jacob 2011b; pinto et al. 2016) might act in opposition by hampering the recolonisation of fireand drought-sensitive forest taxa. forest expansion would again lead to loss of grazing adapted suites of species, loss of grazing resources and game-viewing activities. remotely sensed data will be critical in monitoring forest expansion, but more detailed palaeoecological studies assessing rate of change in forest communities will be needed to put recent changes into context. decisions as to whether this transition should be permitted or prevented must be negotiated based on the trade-offs between ecosystem services, such as carbon storage, grazing, game viewing and biodiversity (biggs et al. 2011). in this respect, it is important that carbon storage potential of open ecosystems is not underestimated (veldman et al. 2019) and that the urgent need to increase carbon storage does not overwhelm other concerns such as the biodiversity and ecosystems services provided by open ecosystems (midgley 2018). therefore, where the extent of riverine forests is changing, site-specific tpcs will be needed depending on what is ecologically possible and socially preferable. figure 5: possible feedback mechanisms that maintain savanna and forests as alternate stable states. the grass layer in savannas supports fires, perpetuating an open canopy of fire-tolerant savanna trees. forest trees shade out understory and prevent fires, perpetuating the recruitment of shade-tolerant, fire-sensitive forest trees. transitions from savanna to forest can be facilitated by the fertilising effects of co2, fire suppression, changes in rainfall, loss of megaherbivores, or interactions between any or all of these factors. transitions between stable states have consequences for biodiversity and the provision of ecosystem services (e.g. see figure 4). savanna vegetation responds to a range of factors that operate over a range of spatial and temporal scales, and their heterogeneous and patchy nature means that local-scale changes may not reflect overall vegetation dynamics at landscape or regional scales (e.g. see gillson 2004a, ekblom & gillson 2009; 2010c). although global drivers such as co2 and regional effects such as rinderpest have had major effects on the composition of southern african savannas, the palaeoecological record also shows the importance of local factors, specifically fire and herbivory, in maintaining alternate stable states and in driving transitions between them (e.g. gillson 2004a, 2004b; gillson & ekblom 2009a). this knowledge is of great significance for biodiversity conservation because local factors can be manipulated by conservation managers to maintain desirable states, prevent transitions to undesirable states and even in some cases attempt to reverse transitions. informed local management can thus have major effects on the future of biodiversity and ecosystem services. greater understanding of the mechanisms that drive transitions between stable states is needed to identify leverage points where managers can manipulate local drivers, such as fire and herbivory, to develop appropriate tpcs and maintain favoured vegetation states. such analyses could lead to the development of co-designed models that allow managers to test the effects of different fire and herbivory regimes under varying future scenarios of climate and co2, possibly utilising palaeoecological records from past warm periods such as the medieval warm period and mid-holocene altithermal to provide clues of how savannas change in warmer climates and linked rainfall changes. as has been briefly discussed here, there is still plenty of work needed to refine climate models using palaeoecological data (see the discussion in fitchett et al. 2017). the decision as to what constitutes a desirable state is to some extent subjective, and we argue here that the knowledge of palaeoecology and the impacts of interacting environmental and social drivers can help stakeholders to develop tpcs for current use, as well as to explore future scenarios of vegetation change under different scenarios of rainfall, drought, fire and herbivory. the complexity of savanna ecology and the feedbacks between climate, disturbance and vegetation mean that various future states are possible. for example, increasing drought may reduce fire occurrence because of reduced grass biomass (van wilgen et al. 2004) and could also reduce the population of herbivores, reducing grazing pressure. processes such as system dynamics modelling will assist stakeholders in exploring these feedback loops, helping them to make informed decisions about desirable outcomes and the best management practices to achieve them, both now and in the future. acknowledgements the authors would like to thank hana petersen for producing figure 1. the authors acknowledge the south africa national parks for their permission to carry out the research, as well as for the comments by the anonymous reviewers. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. author’s contributions both authors were involved in generating and interpreting data, and in drafting of the manuscript. funding information the authors acknowledge funding from the applied centre for climate and earth system science (access), the higgins-trapnell family foundation, the andrew w. mellon foundation and the national research foundation (nrf) of south africa. data availability statement data sharing is not applicable to this article as no new data were created or analysed in this study. disclaimer the views and opinions expressed in this article are the authors’ own and not an official position of the institution or funder. references archibald, s., beckett, h., bond, w.j., coetsee, c., drucer, d.j. & staver, c., 2017, ‘10interactions between fire and ecosystem processes’, in j. cromsigt, s. archibald & n. owen-smith (eds.), conserving africa’s mega-diversity in the anthropocene: the hluhluwe-imfolozi park story, pp. 233–262, cambridge university press, cambridge. archibald, s., bond, w.j., stock, w.d. & fairbanks, d.h.k., 2005, ‘shaping the landscape: fire-grazer interactions in an african savanna’, ecological applications 15(1), 96–109. https://doi.org/10.1890/03-5210 beckage, b., bucini, g., gross, l.j., platt, w.j., higgins, s.i., fowler, n.l. et al., 2019, ‘water limitation, fire, and savanna persistence: a conceptual model’, in p.f. scogings & m. sankaran (eds.), savanna woody plants and large herbivores, pp. 643–659, wiley, hoboken, nj. bennett, e.l., 2014, ‘legal ivory trade in a corrupt world and its impact on african elephant populations’, conservation biology 29(1), 54–60. https://doi.org/10.1111/cobi.12377 biggs, h., ferreira, s., freitag-ronaldson, s. & grant-biggs, r., 2011, ‘taking stock after a decade: does the “thresholds of potential concern” concept need a socioecological revamp?’, koedoe 53(2), 60–68. https://doi.org/10.4102/koedoe.v53i2.1002 bond, w.j. & midgley, g.f., 2012, ‘carbon dioxide and the uneasy interactions of trees and savannah grasses’, philosophical transactions of the royal society b 367(1588), 601–612. https://doi.org/10.1098/rstb.2011.0182 breman, e., ekblom, a., gillson, l. & norström, e., 2019, ‘phytolith-based environmental reconstruction from an altitudinal gradient in mpumalanga, south africa, 10,600 bp–present’, review of paleobotany and palynology 263(2019), 104–116. https://doi.org/10.1016/j.revpalbo.2019.01.001 carruthers, j., 1995, the kruger national park: a social and poltical history, university of natal press, pietermaritzburg. charles-dominique, t., staver, a., midgley, g. & bond, w., 2015, ‘functional differentiation of biomes in an african savanna/forest mosaic’, south african journal of botany 101(2015), 82–90. https://doi.org/10.1016/j.sajb.2015.05.005 davis-reddy, c.l. & vincent, k., 2017, climate risk and vulnerability: a handbook for southern africa, 2nd edn., csir, pretoria. ekblom, a. & gillson, l., 2010a, ‘dung fungi as indicators of past herbivore abundance, kruger and limpopo national park’, palaeogeography, palaeoclimatology, palaeoecology 296(1–2), 14–27. https://doi.org/10.1016/j.palaeo.2010.06.009 ekblom, a. & gillson, l., 2010b, ‘fire history and fire ecology of northern kruger (knp) and limpopo national park (pnl), southern africa’, the holocene 20(7), 1063–1077. https://doi.org/10.1177/0959683610369499 ekblom, a. & gillson, l., 2010c, ‘hierarchy and scale: testing the long term role of water, grazing and nitrogen in the savanna landscape of limpopo national park (mozambique)’, landscape ecology 25, 1529–1546. ekblom, a., gillson, l. & notelid, m., 2011, ‘a historical ecology of the limpopo and kruger national parks and lower limpopo valley’, journal of archaeology and ancient history 1(1), 1–29. ekblom, a., gillson, l., risberg, j., holmgren, k. & chidoub, z., 2012, ‘rainfall variability and vegetation dynamics of the lower limpopo valley, southern africa, 500 ad to present’, palaeogeography, palaeoclimatology, palaeoecology 363–364(2012), 69–78. https://doi.org/10.1016/j.palaeo.2012.08.015 engelbrecht, c.j. & engelbrecht, f.a., 2016, ‘shifts in köppen-geiger climate zones over southern africa in relation to key global temperature goals’, theoretical and applied climatology 123, 247–261. https://doi.org/10.1007/s00704-014-1354-1 engelbrecht, c.j., engelbrecht, f.a. & dyson, l.l., 2013, ‘high-resolution model-projected changes in mid-tropospheric closed-lows and extreme rainfall events over southern africa’, international journal of climatology 33(1), 173–187. https://doi.org/10.1002/joc.3420 engelbrecht, f., mcgregor, j. & engelbrecht, c., 2009, ‘dynamics of the conformal-cubic atmospheric model projected climate-change signal over southern africa’, international journal of climatology: a journal of the royal meteorological society 29(7), 1013–1033. https://doi.org/10.1002/joc.1742 fitchett, j.m., grab, s.w., bamford, m.k. & mackay, a.w., 2017, ‘late quaternary research in southern africa: progress, challenges and future trajectories’, transactions of the royal society of south africa 72(3), 280–293. https://doi.org/10.1080/0035919x.2017.1297966 forbes, c. j., gillson, l. & hoffman, m. t., 2018, ‘shifting baselines in a changing world: identifying management targets in endangered heathlands of the cape floristic region, south africa’, anthropocene 22, 81–93. https://doi.org/10.1016/j.ancene.2018.05.001 gillson, l., 2004a, ‘evidence of hierarchical patch dynamics in an east african savanna?’, landscape ecology 19(2004):883–894. https://doi.org/10.1007/s10980-004-0248-5 gillson, l., 2004b, ‘testing non-equilibrium theories in savannas: 1400 years of vegetation change in tsavo national park, kenya’, ecological complexity 1(4), 281–298. https://doi.org/10.1016/j.ecocom.2004.06.001 gillson, l., 2015a, biodiversity conservation and environmental change: using palaeoecology to manage dynamic landscapes in the anthropocene, oxford university press, oxford. gillson, l., 2015b, ‘evidence of a tipping point in a southern african savanna?’, ecological complexity 21(2015), 78–86. https://doi.org/10.1016/j.ecocom.2014.12.005 gillson, l. & duffin, k.i., 2007, ‘thresholds of potential concern as benchmarks in the management of african savannahs’, philosophical transactions of the royal society of london series b-biological sciences 362(1478), 309–319. https://doi.org/10.1098/rstb.2006.1988 gillson, l. & ekblom, a., 2009a, ‘resilience and thresholds in savannas: nitrogen and fire as drivers and responders of egetation transition’, ecosystems 12(2009), 1189–1203. https://doi.org/10.1007/s10021-009-9284-y gillson, l. & ekblom, a., 2009b, ‘untangling anthropogenic and climatic influence on riverine forest in the kruger national park, south africa’, vegetation history and archaeobotany 18(2009), 171–185. https://doi.org/10.1007/s00334-008-0202-6 gillson, l. & marchant, r., 2014, ‘from myopia to clarity: sharpening the focus of ecosystem management through the lens of palaeoecology’, trends in ecology & evolution 29, 317–325. https://doi.org/10.1016/j.tree.2014.03.010 haensler, a., 2010, how the future climate of the southern african region might look like: results of a high resolution regional climate change projection, pik report 23, potsdam-institut für klimafolgenforschung, potsdam. haensler, a., hagemann, s. & jacob, d., 2011a, ‘dynamical downscaling of era40 reanalysis data over southern africa: added value in the simulation of the seasonal rainfall characteristics’, international journal of climatology 31(15), 2338–2349. https://doi.org/10.1002/joc.2242 haensler, a., hagemann, s. & jacob, d., 2011b, ‘the role of the simulation setup in a long-term high-resolution climate change projection for the southern african region’, theoretical and applied climatology 106(2011), 153–169. https://doi.org/10.1007/s00704-011-0420-1 hempson, g.p., archibald, s., donaldson, j.e. & lehmann, c.e., 2019, ‘alternate grassy ecosystem states are determined by palatability–flammability trade-offs’, trends in ecology & evolution 34(4), 286–290. https://doi.org/10.1016/j.tree.2019.01.007 higgins, s.i. & scheiter, s., 2012, ‘atmospheric co2 forces abrupt vegetation shifts locally, but not globally’, nature 488(2012), 209–212. https://doi.org/10.1038/nature11238 hoffmann, w.a., jaconis, s.a., mckinley, k.l., geiger, e.l., gotsch, s.g. & franco, a.c., 2012, ‘fuels or microclimate? understanding the drivers of fire feedbacks at savanna–forest boundaries’, austral ecology 37(6), 634–643. https://doi.org/10.1111/j.1442-9993.2011.02324.x holmgren, k., lee-thorp, j.a., cooper, g.r.j., lundblad, k., partridge, t.c., scott, l. et al., 2003, ‘persistent millenial-scale climatic variability over the past 25,000 years in southern africa’, quaternary science reviews 22(21–22), 2311–2326. https://doi.org/10.1016/s0277-3791(03)00204-x holling, c.s., 1996, ‘engineering resilience versus ecological resilience’, in p. schulze, (ed.), engineering within ecological constraints, pp. 31–44, national academy press, washington. holmgren, k., moberg, a., svanered, o. & tyson, p.d., 2001, ‘a preliminary 3000-year regional temperature reconstruction for south africa’, south african journal of science 97(1–2), 49–51. holmgren, k., risberg, j., freudendahl, j., achimo, m., ekblom, a., mugabe, j. et al., 2012, ‘water-level variations in lake nhauhache, mozambique, during the last 2,300 years’, journal of paleolimnology 48, 311–322. https://doi.org/10.1007/s10933-012-9596-3 humphries, m., green, a. & finch, j., 2016, ‘evidence of el niño driven desiccation cycles in a shallow estuarine lake: the evolution and fate of africa’s largest estuarine system, lake st lucia’, global and planetary change 147, 97–105. https://doi.org/10.1016/j.gloplacha.2016.11.002 lee-thorp, j., holmgren, k., lauritzen, s.e., linge, h., moberg, a., partridge, t. et al., 2001, ‘rapid climate shifts in the southern african interior throughout the mid to late holocene’, geophysical research letters 28(23), 4507–4510. https://doi.org/10.1029/2000gl012728 lubbe, b.a., du preez, e.a., douglas, a. & fairer-wessels, f., 2017, ‘the impact of rhino poaching on tourist experiences and future visitation to national parks in south africa’, current issues in tourism 22(1), 8–15. https://doi.org/10.1080/13683500.2017.1343807 midgley, g., 2018, ‘narrowing pathways to a sustainable future’, science 360(6390), 714–715. https://doi.org/10.1126/science.aat6671 midgley, g.f. & bond, w.j., 2015, ‘future of african terrestrial biodiversity and ecosystems under anthropogenic climate change’, nature climate change 5(2015), 823–829. https://doi.org/10.1038/nclimate2753 neumann, f.h., botha, g.a. & scott, l., 2014, ‘18,000 years of grassland evolution in the summer rainfall region of south africa: evidence from mahwaqa mountain, kwazulu-natal’, vegetation history and archaeobotany 23, 665–681. https://doi.org/10.1007/s00334-014-0445-3 o’connor, t.g., 2010, ‘transformation of riparian forest to woodland in mapungubwe national park, south africa, between 1990 and 2007’, austral ecology 35(7),778–786. https://doi.org/10.1111/j.1442-9993.2009.02084.x oliver, t.h., heard, m.s., isaac, n.j., roy, d.b., procter d., eigenbrod, f. et al., 2015, ‘biodiversity and resilience of ecosystem functions’, trends in ecology & evolution 30, 673–684. https://doi.org/10.1016/j.tree.2015.08.009 parr, c.l., lehmann, c.e., bond, w.j., hoffmann, w.a. & andersen, a.n., 2014, ‘tropical grassy biomes: misunderstood, neglected, and under threat’, trends in ecology & evolution 29(4), 205–213. https://doi.org/10.1016/j.tree.2014.02.004 pinto, i., jack, c. & hewitson, b., 2018, ‘process-based model evaluation and projections over southern africa from coordinated regional climate downscaling experiment and coupled model intercomparison project phase 5 models’, international journal of climatology 38(11), 4251–4261. https://doi.org/10.1002/joc.5666 pinto, i., lennard, c., tadross, m., hewitson, b., dosio, a., nikulin, g. et al., 2016, ‘evaluation and projections of extreme precipitation over southern africa from two cordex models’, climatic change 135, 655–668. https://doi.org/10.1007/s10584-015-1573-1 purdon, j., parr, c.l. & somers, m.j., 2019, ‘grazing by large savanna herbivores indirectly alters ant diversity and promotes resource monopolisation’, peerjournl 7, e6226. https://doi.org/10.7717/peerj.6226 redfern, j., grant, c., gaylard, a. & getz, w., 2005, ‘surface water availability and the management of herbivore distributions in an african savanna ecosystem’, journal of arid environments 63(2),406–424. https://doi.org/10.1016/j.jaridenv.2005.03.016 rogers, k. & biggs, h., 1999, ‘integrating indicators, endpoints and value systems in strategic management of the rivers of the kruger national park’, freshwater biology 41(2), 439–451. rogers, k.h., 1997, ‘operationalizing ecology under a new paradigm: an african perspective’, in s.t.a. pickett, r.s. ostfeld, m. shachak & g.e. likens (eds.), the ecological basis for conservation: heterogeneity, ecosystems, and biodiversity, pp. 60–77, chapman and hall, new york, ny. rogers, k.h., 2003, ‘adopting a heterogeneity paradigm: implications for management of protected savannas’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 41–58, island press, washington, dc. rogers, k.h. & o’keeffe, j., 2003, ‘river heterogeneity: ecosystem structure, function, and management’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 189–219, island press, washington, dc. rountree, m., rogers, k. & heritage, g., 2000, ‘landscape state change in the semi-arid sabie river, kruger national park, in response to flood and drought’, south african geographical journal 82(3), 173–181. https://doi.org/10.1080/03736245.2000.9713711 sankaran, m., hanan, n.p., scholes, r.j., ratnam, j., augustine, d.j., cade, b.s. et al., 2005, ‘determinants of woody cover in african savannas’, nature 438, 846–849. https://doi.org/10.1038/nature04070 sankaran, m., ratnam, j. & hanan, n., 2008, ‘woody cover in african savannas: the role of resources, fire and herbivory’, global ecology and biogeography 17(2), 236–245. https://doi.org/10.1111/j.1466-8238.2007.00360.x scheiter, s. & higgins, s.i., 2012, ‘how many elephants can you fit into a conservation area’, conservation letters 5(3), 176–185. https://doi.org/10.1111/j.1755-263x.2012.00225.x scott, l., 2016, ‘fluctuations of vegetation and climate over the last 75 000 years in the savanna biome, south africa: tswaing crater and wonderkrater pollen sequences reviewed’, quaternary science reviews 145, 117–133. https://doi.org/10.1016/j.quascirev.2016.05.035 scott, l. & lee-thorp, j.a., 2004, ‘holocene climatic trends and rhythms in southern africa’, in r.w. battarbee, f. gasse & c.e. stickley (eds.), past climate variability through europe and africa, pp. 69–91, vol. 6., 69–91, springer, dordrecht. scott, l., neumann, f.h., brook, g.a., bousman, c.b., norström, e. & metwally, a., 2012, ‘terrestrial fossil-pollen evidence of climate change during the last 26 thousand years in southern africa’, quaternary science reviews 32, 100–118. https://doi.org/10.1016/j.quascirev.2011.11.010 staver, a.c., archibald, s. & levin, s., 2011, ‘tree cover in sub-saharan africa: rainfall and fire constrain forest and savanna as alternative stable states’, ecology 92(5), 1063–1072. https://doi.org/10.1890/10-1684.1 staver, a.c. & bond, w.j., 2014, ‘is there a “browse trap?” dynamics of herbivore impacts on trees and grasses in an african savanna’, journal of ecology 102(3), 595–602. https://doi.org/10.1111/1365-2745.12230 sundqvist, h.s., holmgren, k., fohlmeister, j., zhang, q., matthews, m.b., spötl, c. & körnich, h., 2013, ‘evidence of a large cooling between 1690 and 1740 ad in southern africa’, scientific reports 3, 1767. https://doi.org/10.1038/srep01767 sundstrom, sm., allen, c.r. & gunderson, l., 2016, resisting resilience theory: a response to connell and ghedini’, trends in ecology & evolution 31(6), 412–413. https://doi.org/10.1016/j.tree.2016.03.014 van wilgen, b.w., govender, n., smit, i.p. & macfadyen, s., 2014, ‘the ongoing development of a pragmatic and adaptive fire management policy in a large african savanna protected area’, journal of environmental management 132, 358–368. https://doi.org/10.1016/j.jenvman.2013.11.003 van wilgen, b.w., govener, n., biggs, h.c., ntsala, d. & funda, x.n., 2004, ‘response of savanna fire regimes to changing fire management policies in a large african national park’, conservation biology 18(6), 1533–1540. https://doi.org/10.1111/j.1523-1739.2004.00362.x veldman, j.w., aleman, j.c., alvarado, s.t., anderson, t.m., archibald, s., bond, w.j. et al., 2019, ‘comment on “the global tree restoration potential”’, science 366(6463), eaay7976. https://doi.org/10.1126/science.aay7976 venter, z.s., hawkins, h.j. & cramer, m.d., 2017, ‘implications of historical interactions between herbivory and fire for rangeland management in african savannas’, ecosphere 8(10), e01946. https://doi.org/10.1002/ecs2.1946 waldram, m., bond, w.j. & stock, w.d., 2007, ‘ecological engineering by a mega-grazer: white rhino impacts on a south african savanna’, ecosystems https://doi.org/10.1007/s10021-007-9109-9 wigley, b.j., bond, w.j. & hoffman, m.t., 2009, ‘bush encroachment under three contrasting land-use practices in a mesic south african savanna’, african journal of ecology 47(s1), 62–70. https://doi.org/10.1111/j.1365-2028.2008.01051.x wigley, b.j., bond, w.j. & hoffman, m.t., 2010, ‘thicket expansion in a south african savanna under divergent land use: local vs. global drivers?’, global change biology 16(3), 964–976. https://doi.org/10.1111/j.1365-2486.2009.02030.x wittemyer, g., northrup, j.m., blanc, j., douglas-hamilton, i., omondi, p. & burnham, k.p., 2014, ‘illegal killing for ivory drives global decline in african elephants’, proceedings of the national academy of sciences 111(36), 13117–13121. https://doi.org/10.1073/pnas.1403984111 woodborne, s., gandiwa, p., hall, g., patrut, a. & finch, j., 2016, ‘a regional stable carbon isotope dendro-climatology from the south african summer rainfall area’, plos one 11(7), e0159361. https://doi.org/10.1371/journal.pone.0159361 woodborne, s., hall, g., robertson, i., patrut, a., rouault, m., loader, n.j. et al. 2015, ‘a 1000-year carbon isotope rainfall proxy record from south african baobab trees (adansonia digitata l.)’, plos one 10(5), e0124202. https://doi.org/10.1371/journal.pone.0124202 appendix 1 table 1-a1: site locations, ecology and sampling details for sites presented in the article. filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 abstract introduction study area materials and methods results description of the plant communities discussion conclusion acknowledgements references about the author(s) petrus j. van staden department of botany, university of pretoria, pretoria, south africa george j. bredenkamp department of botany, university of pretoria, pretoria, south africa hugo bezuidenhout scientific services, sanparks, kimberley, south africa applied behavioural ecology and ecosystem research unit, faculty of environmental sciences, university of south africa, florida, south africa leslie r. brown applied behavioural ecology and ecosystem research unit, faculty of environmental sciences, university of south africa, florida, south africa citation van staden, p.j., bredenkamp, g.j., bezuidenhout, h. & brown, l.r., 2021, ‘a reclassification and description of the waterberg mountain vegetation of the marakele national park, limpopo province, south africa’, koedoe 63(1), a1689. https://doi.org/10.4102/koedoe.v63i1.1689 note: additional supporting information may be found in the online version of this article as online appendix 1. original research a reclassification and description of the waterberg mountain vegetation of the marakele national park, limpopo province, south africa petrus j. van staden, george j. bredenkamp, hugo bezuidenhout, leslie r. brown received: 25 june 2021; accepted: 08 oct. 2021; published: 10 dec. 2021 copyright: © 2021. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the description and classification of vegetation are important for conservation and resource management. the aim of this study was to identify, reclassify and describe the plant communities present in the waterberg mountain vegetation of the marakele national park in the limpopo province, south africa. a phytosociological classification, mapping and description of sections of the waterberg mountain vegetation in the park were done in 1995. since 1995, various farms adjacent to the park have been bought and incorporated into it. little is known about the vegetation and habitat status of these newly acquired areas, which led to this study. the floristic data were analysed according to the braun-blanquet procedure using the braun blanquet personal computer (bbpc) suite as well as the juice software package, whilst the diversity of the plant communities was determined using the shannon–wiener and gini–simpson indices. a total of 12 plant communities were identified and are described according to their diagnostic, constant and dominant plant species as determined from the synoptic table analysis as well as their characteristic species as derived from the phytosociological table. based on the topography and plant species composition, the vegetation can be grouped into five major groups, namely the: (1) lower midslope and plateau shruband woodlands, (2) high altitude midslope woodland, (3) high-lying plateau and midslope grass-, shruband woodlands, (4) ravine, footslope and drainage line forests and woodland, and (5) higher-lying plateau wetlands and forblands. the high altitude midslope grassland and shrubland and the lower midslope and plateau areas have the highest diversity. the high-lying vegetation has affinity with bankenveld and drakensberg vegetation, whilst the relatively low-lying plateaus and midslope vegetation are typical of the bushveld areas. conservation implications: this reclassification, mapping and description of the waterberg mountain vegetation have been incorporated into the management plan for the park. it will enable managers to make scientifically based decisions on the management of the different ecosystems to ensure biodiversity conservation. this vegetation study also provides baseline information that allows for vegetation assessments to determine veld condition, carrying capacity and stocking density for the park. keywords: marakele national park; juice; phytosociology; diversity; vegetation structure; nature conservation. introduction marakele national park including marakele park (pty) ltd (referred to as marakele in the text) is managed under the jurisdiction of south african national parks. the park was established in 1988 and was proclaimed as a national park in february 1994 (van staden 2002). since 1995, various farms adjacent to the original marakele have been bought and incorporated into the park. these new areas have not only resulted in the park more than doubling its size from its previous 290.51 km² (29 051 ha) to approximately 700.00 km² (70 000 ha) but included new ecosystems and made the re-introduction of various large mammals such as elephant, buffalo, blackand white rhinoceros and predators such as lion possible (van staden 2007). the necessity for a vegetation classification, description and mapping of conservation areas was stated by bezuidenhout (2009). in order to manage and conserve our natural heritage for current and future generations, a profound knowledge of the different ecosystems is a prerequisite. to achieve this, an inventory of the biotic and abiotic components of the different ecosystems needs to be compiled to assist with informed management decision making (brown et al. 2013; van staden 2002). various studies on sections of the mountain vegetation of the original marakele national park were undertaken in the past by van staden (2002) and van staden and bredenkamp (2005). these studies described the vegetation of selected sections of the mountainous areas of the original park only. this study was undertaken to provide a detailed classification and ecological interpretation of the mountain vegetation of marakele (marakele national park including marakele park [pty] ltd). study area location and size marakele is situated approximately 13 km east of the town of thabazimbi, in the limpopo province, south africa (see figure 1). it extends from latitude 24.54081° to 24.49407°s and longitude 027.44802° to 027.76378°e (van staden 2015). figure 1: the location of the marakele national park. geology marakele is underlain by various geological formations. the geology of the mountainous area of marakele consists of rocks belonging to the kransberg subgroup, which consists of yellowish, coarse-grained sandstone, locally gritty, with ferruginous laminae on cross-bedding planes of the sandriviersberg stage, kransberg series of the sandriviersberg sandstone formation with a thickness of approximately 1250 m, and covers the largest part of the study area in the waterberg mountain in marakele (van staden 2015). the waterberg rocks were faulted and intruded by diabase sills and dykes of the post waterberg to the southern and northern areas of marakele (van staden 2015). physiography, soil, land types and vegetation the mountainous landscape includes upper and low-lying valleys and drainage lines, together with steep to moderately steep mountain slopes and relatively flat to undulating plateaus. altitude ranges from 1200 m to 2100 m above the mean sea level. the following topographical terrain units are distinguished in marakele: plateau (1), scarp (2), midslopes (3), footslopes (4), wetlands and drainage lines (5) (land type survey staff 1988). the matlabas-, sterkstroom-, sondagsand mamba rivers form the core of the drainage system of the area. the study area comprises three land types, namely ad, fa and ib (land type survey staff 1988). the soil varies from deep (> 0.8 m) red-brown and yellow-brown sands (avalon, champagne, clovelly, hutton, katspruit, oakleaf and westleigh soil forms) to shallow (< 0.3 m) and stony (mispah and glenrosa soil forms) (soil classification working group 1991; van staden 2015). mucina and rutherford (2006) classified the vegetation of marakele as the western sandy bushveld (svcb16) and the waterberg mountain bushveld (svcb17) vegetation types in the savanna biome, the waterberg-magaliesberg summit sourveld (gm29) vegetation type in the grassland biome and the marakele afromontane forest (foz2) in the northern afromontane forest vegetation type in the afro-temperate, subtropical and azonal forests biome of south africa. climate the rainfall, mainly during the spring and summer (september–march), can range from as high as 1196.9 to lower than 400 mm per year. the mean annual rainfall for marakele based on data collected in marakele is ca. 704.5 mm per annum (van staden 2015). the park recorded winter temperatures (coldest months june–july) as low as –3.7 °c, whilst the summer temperatures (warmest months december to february) can reach a maximum temperature of 40.0 °c. mean frost days per annum is 16 days with the earliest occurrence 14 may and the latest occurrence 23 july (van staden 2015). materials and methods using 1:50 000 stereo aerial photographs (task 874 of 1984), the study area was stratified into relatively uniform physiographic-physiognomic units. after a reconnaissance of the area, sample plots were placed in a stratified random manner (brown et al. 2013) within the different homogeneous units identified. the number of sample plots for each delineated physiographic-physiognomic unit was determined according to the size of each delineated unit. the larger the area, the higher the number of sampling plots allocated to the unit. a total of 136 sample plots were located within these units to ensure that all variations in the vegetation were considered and sampled. global positioning system (gps) locations of all sample plots were noted. plot sizes were fixed at approximately 200 m² according to van staden (2002). the floristic data were analysed according to the braun–blanquet procedure. the braun blanquet personal computer (bbpc) suite (bezuidenhout, biggs & bredenkamp 1996) was used to capture the data from where it was imported into the juice software package (roleček et al. 2009; tichy & holt 2006). the numerical classification method twinspan (hill 1979) was used to provide a hierarchical classification of the mountainous vegetation of the study area. the classification results were used to compile a phytosociological table in which the species were manually rearranged into species groups as described by brown et al. (2013), but no relevés or clusters were modified. a synoptic table was compiled for the different plant communities using the juice 7.1.21 software package (tichy & holt 2006). a combined synoptic table analysis of columns was performed with the lower threshold values set as follows: fidelity – 70; frequency – 80 and cover – 30. the different plant communities are described according to their diagnostic, constant and dominant plant species (tichy & holt 2006) as determined from the synoptic table analysis as well as their characteristic species as derived from the phytosociological table. a vegetation map was compiled by overlaying the gps locations of the sample plots and their classification on georeferenced google earth images. the structural classification for this study was based on the broad scale structural classification of edwards (1983). the tree stratum includes all singleand multi-stemmed woody, self-supporting plants 3 m and taller, and the shrub stratum is classified as singleor multi-stemmed woody, self-supporting plants < 3 m high (adapted from edwards 1983). species richness was determined for each plant community by determining the number of species present in each. the shannon–wiener index of diversity (h) (shannon & weaver 1949) and the rich–gini–simpson index of diversity (rgs) (guiasu & guiasu 2010) were calculated for each of the major plant communities. where pi = (number of individuals of one species)/(total number of all individuals) the cover-abundance data collected using the braun–blanquet surveys were transformed to a numerical scale (r = 0.5; + = 1; 1 = 2; 2a = 8.5; 2b = 17.5; 3 = 35; 4 = 70; 5 = 140) to calculate species diversity as described by van der maarel (2007). the diversity results from the indices were converted to true diversities (effective numbers) as described by jost (2006) to give them common behaviours and properties that allow for appropriate comparisons between plant communities. ethical considerations this article followed all ethical standards for research without direct contact with human or animal subjects. results classification the analysis of the waterberg mountain vegetation in marakele resulted in the identification of 12 plant communities, 16 sub-communities and two variants that are indicated in the phytosociological table and vegetation map (online appendix 1 & figure 2) and are listed below (the sub-communities and the variants were omitted from the vegetation map as their distribution is too limited to be visible at the scale on the vegetation map). themeda triandra–senegalia caffra woodland 1.1 themeda triandra–senegalia caffra–eragrostis curvula sub-community 1.2 themeda triandra–senegalia caffra–eragrostis chloromelas sub-community 1.2.1 themeda triandra variant 1.2.2 triumfetta sonderi variant tristachya leucothrix–protea welwitschii shrubland monocymbium ceresiiforme–andropogon schirensis grassland 3.1 monocymbium ceresiiforme–andropogon schirensis–protea gaguedi sub-community 3.2 monocymbium ceresiiforme–andropogon schirensis–helichrysum kraussii sub-community indigofera mollicoma–encephalartos eugene–maraisii shrubland tephrosia elongata–protea caffra woodland 5.1 tephrosia elongata–protea caffra–gerbera viridifolia sub-community 5.2 tephrosia elongata–protea caffra–brachiaria serrata sub-community combretum molle–setaria lindenbergiana woodland 6.1 combretum molle–setaria lindenbergiana–brachylaena rotundata sub-community 6.2 combretum molle–setaria lindenbergiana–diplorhynchus condylocarpon sub-community perotis patens–burkea africana woodland 7.1 perotis patens–burkea africana–fadogia homblei sub-community 7.2 perotis patens–burkea africana–gardenia volkensii sub-community kirkia wilmsii–buxus macowanii forest diospyros whyteana–calodendron capensis forest cyperus albostriatus–podocarpus latifolius forest 10.1 cyperus albostriatus–podocarpus latifolius–rothmannia capensis sub-community 10.2 cyperus albostriatus–podocarpus latifolius–widdringtonia nodiflora sub-community syzygium cordatum–ischaemum fasciculatum woodland 11.1 syzygium cordatum–ischaemum fasciculatum–schoenoplectus corymbosus sub-community 11.2 syzygium cordatum–ischaemum fasciculatum–osmunda regalis sub-community leptochloa fusca–fuirena pubescens forbland 12.1 leptochloa fusca–fuirena pubescens–aristida junciformis sub-community 12.2 leptochloa fusca–fuirena pubescens–drosera madagascariensis sub-community figure 2: plant communities of the mountainous vegetation of marakele national park. table 1 provides a summary of the various soil, land type, terrain units and geology associated with each plant community, whilst table 2 lists the mean height classes (m) and percentage canopy cover for the tree, shrub and herbaceous (grasses and forbs) layers. table 1: summary of soil parameters relating to each plant community. table 2: summary of the average height classes (m) and – cover abundance (%) relating to each plant community. description of the plant communities all references to species groups in the descriptions of the various plant communities refer to the phytosociological table (see online appendix 1), whilst the diagnostic, constant and dominant species were obtained from the analysis of the synoptic table as described in the methods. themeda triandra–senegalia caffra woodland this community is situated on the midslopes and plateaus of marakele (see figure 2). altitude ranges from 1300 m to 1600 m above the mean sea level (mamsl). the slopes are gentle to moderate and vary from 4° to 16°. rock cover varies between 10% and 75% and occurs as loose rocks and boulders. species from species group a are characteristic of this woodland. the tree senegalia caffra (species group a) and the grasses themeda triandra and setaria sphacelata (species group v) dominate the woody and herbaceous layers, respectively. this plant community has a total of 129 plant species recorded and is divided into the following two sub-communities. themeda triandra–senegalia caffra–eragrostis curvula sub-community this woodland sub-community is situated on plateau areas and midslopes (slope 4° – 16°) of marakele. the altitude ranges from 1300 mamsl to 1600 mamsl. rock cover is 10% – 35% and comprises large boulders and loose rocks. this community is associated with the ib280 land type and occurs mostly on the mispah soil form that is shallow with the soil depth of less than 150 mm (see table 1). species from species group b characterise this sub-community. the following are the important species as determined from the synoptic table analysis: diagnostic species: none constant species: pearsonia cajanifolia, raphionacme burkei, senegalia caffra, setaria sphacelata, themeda triandra, trachypogon spicatus dominant species: senegalia caffra, setaria sphacelata, themeda triandra the shrub dombeya rotundifolia (species group v) and the grasses melinis repens (species group v), melinis nerviglumis (species group i) and heteropogon contortus (species group t) are prominent locally. the tree (4 m tall) and herbaceous layers (0.8 m tall) are well developed and the most prominent layers in this sub-community covering 80% and 70%, respectively. the shrub layer (1.1 m tall) is poorly developed with a cover of 19% (see table 2). themeda triandra–senegalia caffra–eragrostis chloromelas sub-community this woodland sub-community is situated on the northern to north-eastern lower midslopes and plateaus of the mountainous areas of marakele. the altitude ranges from 1163 mamsl to 1420 mamsl with the steepness of the slopes from 4° to 23°. this woodland sub-community is covered with medium to large boulders and large rocks with an approximate 36% coverage. this sub-community is associated with the ib280 land type and occurs mostly on the hutton soil form that ranges in depth from 400 mm to 1200 mm (see table 1). species from species group c characterise this sub-community. the following are the important species as determined from the synoptic table analysis: diagnostic species: none constant species: asparagus transvaalensis, bewsia biflora, elionurus muticus, eragrostis racemosa, melinis repens, raphionacme divaricatum, senegalia caffra, setaria sphacelata, themeda triandra, vangueria infausta dominant species: faurea saligna, senegalia caffra, themeda triandra the tree layer is 5 m tall and covers 53% and is the most prominent layer in this sub-community, whereas the shrub and herbaceous layers (1.1 m and 0.8 m tall) are not well developed with relatively low cover of 15% and 29%, respectively (see table 2). this sub-community is divided into two variants: themeda triandra variant: this variant is characterised by the absence of species from species group d (characteristic species for variant 1.2.2) and include species such as the tree lannea discolor (species group n), the grass heteropogon contortus (species group t), the grass themeda triandra (species group v) and the tree vangueria infausta (species group v). triumfetta sonderi variant: this variant is characterised by the presence of species from species group d. the shrub ozoroa paniculosa (species group o), the grasses eragrostis racemosa (species group v), aristida meridionalis and the forbs triumfetta sonderi (species group d) and pseudopegolettia tenella (species group l) are locally prominent within this variant. tristachya leucothrix–protea welwitschii shrubland this community occurs in the eastern section of marakele (see figure 2) and occurs on footslope at altitudes ranging from 1480 mamsl to 1500 mamsl. the slope varies from 1° to 5° and no rocks are present. this shrubland is associated with the ib286 land type and occurs mostly on hutton soil form that ranges in depth from 400 mm to 1200 mm (see table 1). species from species group e characterise this community. the following are the important species as determined from the synoptic table analysis: diagnostic species: cymbopogon caesius, cynodon dactylon, dicoma zeyheri, elephantorrhiza elephantina, eragrostis plana, gladiolus pretoriensis, lasiosiphon kraussianus, helichrysum coriaceum, hyparrhenia hirta, protea welwitschii, schizocarphus nervosus, senecio coronatus, triraphis andropogonoides, tristachya leucothrix, hilliardiella aristata, selago densiflora constant species: aeschynomene rehmannii, anthospermum hispidulum, aristida congesta subsp. congesta, brachiaria serrata, eragrostis capensis, eragrostis gummiflua, eragrostis racemosa, helichrysum kraussii, heteropogon contortus, hypoxis obtusa, melinis repens, parinari capensis, pygmaeothamnus zeyheri, themeda triandra, thesium utile, trachypogon spicatus, urelytrum agropyroides dominant species: elephantorrhiza elephantina, hyparrhenia hirta, urelytrum agropyroides other species that are prominent in this community are the shrub protea welwitschii (species group e), the dwarf shrub parinari capensis (species group m), the grasses tristachya leucothrix, cynodon dactylon and the forb lasiosiphon kraussianus (species group e). a total number of 77 plant species was recorded for this community. this community has a sparse vegetation cover with the shrub and herbaceous layers covering 10% and 18%, respectively. the shrub layer has an average height of 0.8 m whilst the herbaceous layer is 1 m tall (see table 2). monocymbium ceresiiforme–andropogon schirensis grassland this community is situated on the plateaus of the mountainous areas of marakele (see figure 2). altitude ranges 1300 mamsl – 1600 mamsl and slopes are gentle to moderate (4° – 16°). rock cover is 10% – 75% and occurs as loose rocks and boulders. species from species group f are characteristic of this grassland. the grasses monocymbium ceresiiforme (species group f), loudetia simplex and eragrostis racemosa (species group v) and the tree englerophytum magalismontanum (species group u) dominate the herbaceous and woody layers, respectively. this plant community has a total of 129 plant species recorded and is divided into the following two sub-communities: monocymbium ceresiiforme–andropogon schirensis–protea gaguedi sub-community this shrubland sub-community occurs on the high mountain plateau in the southern section of marakele at an altitude from 1620 mamsl to 1690 mamsl with the slope not exceeding 2°. it is covered with large rocky sheets with a cover of 20% – 35%. this sub-community is associated with the ad20 land type and occurs predominantly on mispah and glenrosa soil forms that are from 100 mm to 600 mm deep (see table 1). species from species group g are characteristic of this sub-community. the following are the important species as determined from the synoptic table analysis: diagnostic species: cleome maculata, cyanotis speciosa, cyperus rupestris, thunbergia atriplicifolia constant species: andropogon schirensis, anthospermum hispidulum, aristida transvaalensis, bulbine burchellii, commelina africana, eragrostis racemosa, fadogia homblei, indigofera mollicoma, loudetia simplex, melinis repens, parinari capensis, pygmaeothamnus zeyheri, schizachyrium sanguineum, sphenostylis angustifolia, trachypogon spicatus dominant species: andropogon schirensis the shrub protea gaguedi (species group g) and the grasses loudetia simplex, eragrostis racemosa (species group v) and aristida transvaalensis (species group u) are prominent throughout this sub-community. the rocky sheet cover has resulted in a low vegetation cover with a sparse shrub layer covering only 3% and the herbaceous layer 30% of the area. the dwarf shrub layer has an average height of 0.8 m compared to the taller grasses that are 1 m tall (see≈table 2). monocymbium ceresiiforme–andropogon schirensis–helichrysum kraussii sub-community the monocymbium ceresiiforme–andropogon schirensis–helichrysum krausii sub-community is situated on the southern section of the high mountain plateau at a higher altitude than the monocymbium ceresiiforme–andropogon schirensis–protea gaguedi sub-community. the altitude ranges from 1640 mamsl to 2000 mamsl. the slope does not exceed 3°, and the rock cover is 30% – 65% consisting of rocky outcrops. this sub-community is associated with the fa286 and ib310 land types and occurs predominantly on rock or the mispah soil form that is 100 mm to 600 mm deep (see table 1). species from species group h are characteristic of this sub-community. the following are the important species as determined from the synoptic table analysis: diagnostic species: none constant species: acalypha angustata, andropogon schirensis, anthospermum hispidulum, bulbine burchellii, dicoma anomala, eragrostis racemosa, fadogia homblei, helichrysum kraussii, loudetia simplex, monocymbium ceresiiforme, trachypogon spicatus dominant species: andropogon schirensis, loudetia simplex, schizachyrium sanguineum the grasses monocymbium ceresiiforme (species group f), eragrostis racemosa (species group v) and the forbs dicoma anomala (species group f) and helichrysum krausii (species group h) are prominent throughout this sub-community. the woody layer (trees and shrubs) is not well developed and covers ≤ 2% of the area and can grow up to 3 m tall, whilst the herbaceous layer covers 23% and has an average height of 0.8 m (see table 2). indigofera mollicoma–encephalartos eugene-maraisii shrubland this community occurs on the scarp of the mountainous areas of marakele (see figure 2). this shrubland community occurs on steep slopes (3° – 30°) at altitudes ranging from 1160 mamsl to 1785 mamsl. rocky outcrops cover 25% – 70% of the area. this community is associated with the fa286 land type with shallow (100 mm – 300 mm) mispah soil form dominant (see table 1). species from species group i are characteristic of this community. the following are the important species as determined from the synoptic table analysis: diagnostic species: none constant species: andropogon schirensis, diheteropogon amplectens, eragrostis racemosa, loudetia simplex, panicum natalense dominant species: andropogon schirensis, combretum apiculatum, cyperus leptocladus, diheteropogon amplectens, elephantorrhiza burkei, encephalartos eugene-maraisii, loudetia simplex, protea caffra, trachypogon spicatus this community is characterised by the presence of the endangered waterberg cycad, encephalartos eugene-maraisii (species group i), which occurs in selected areas within this community. the tree heteropyxis natalensis (species group y), the shrub protea caffra (species group m) and the grasses themeda triandra, loudetia simplex and trachypogon spicatus (species group v) dominate the herbaceous layer in the areas where encephalartos eugene-maraisii occurs. the vegetation has a relatively high cover with the trees (5 m tall) covering 26%, the shrubs (2 m tall) covering 13% and the herbaceous layer (1.8 m tall) covering 43% of the area (table 2). this plant community has 126 plant species recorded. tephrosia elongata–protea caffra woodland this short woodland is found on moderate to steep, southern, south-eastern and south-western midslopes of the mountainous area of marakele (figure 2). the altitude ranges from 1240 mamsl to 1880 mamsl with large boulders, rocky sheets and rocks covering 20% – 70% of the soil. this community is associated with the ib310 land type and is associated with shallow (50 mm – 150 mm) mispah soil form (see table 1). species from species group j are characteristic of this community. the following are the important species as determined from the synoptic table analysis: diagnostic species: erica drakensbergensis, kohautia amatymbica, tephrosia elongata constant species: bulbine burchellii, eragrostis racemosa, loudetia simplex, panicum natalense, themeda triandra dominant species: englerophytum magalismontanum, loudetia simplex, monocymbium ceresiiforme, panicum natalense, podocarpus latifolius, protea caffra, rhynchosia nitens, senecio venosus, tephrosia elongata, themeda triandra, trachypogon spicatus, urelytrum agropyroides the shrub searsia dentata (species group m), scandent shrub searsia magalismontana (species group j), the grass diheteropogon amplectens (species group u) and the forb helichrysum setosum (species group j) are prominent within this community. a high number of 139 plant species was recorded in this community. this community is divided into the following two sub-communities. tephrosia elongata–protea caffra–gerbera viridifolia woodland sub-community this woodland sub-community is situated on steep (16° – 32°) southern, south-eastern and south-western midslopes of marakele. the altitude ranges from 1240 mamsl to 1880 mamsl and large boulders, rocky outcrops and loose rocks cover 20% – 70% of the area (see table 1). the tree (3 m tall), shrub (1.5 m tall) and herbaceous (1.6 m tall) layers have mean cover values of 13%, 7% and 36%, respectively (see table 2). the characteristic species include the grass cymbopogon pospischilii and the forbs scabiosa columbaria, becium obovatum, gerbera viridifolia and berkheya zeyheri (species group k). tephrosia elongata–protea caffra–brachiaria serrata sub-community the brachiaria serrata-protea caffra sub-community is found on the higher midslopes (3) of the mountainous areas of marakele. the gradient ranges from 20° to 31° and the altitude from 1470 mamsl to 1880 mamsl. the area comprises largeto medium-sized rocks covering 40% – 70% of the area with the vegetation growing between the rocks (see table 1). the tree (3 m tall), shrub (1.8 m tall) and herbaceous (0.8 m tall) layers have mean cover values of 42%, 14% and 44%, respectively (see table 2). species from species group l are characteristic for this sub-community and include the grasses brachiaria serrata, aristida aequiglumis, cymbopogon validus and the forbs pseudopegolettia tenella, polygala uncinata, athrixia elata, berkheya insignis, castalis spectabilis and helichrysum cephaloideum. combretum molle–setaria lindenbergiana woodland this woodland is situated on moderate-to-steep midslopes at an altitude of 1360 mamsl – 1620 mamsl with largeand medium-sized rocks covering 7% – 80% of the area (figure 2). this community is associated with fa286 and ib305 land types and is associated with moderately deep (50 mm – 600 mm) mispah soil form (see table 1). species from species group n are characteristic of this woodland community. the tree vangueria infausta and the grasses setaria lindenbergiana (species group n), trachypogon spicatus (species group v) and diheteropogon amplectens (species group u) dominate the woody and herbaceous layers, respectively. this plant community has the highest species richness with 155 different plant species recorded. this community is divided into the following two sub-communities: combretum molle–setaria lindenbergiana–brachylaena rotundata sub-community this short open woodland sub-community is situated on the steep north-eastern and south-western midslopes (20°–33°) of the mountainous areas of marakele altitude ranges from 1400 mamsl to 1540 mamsl with largeand medium-sized rocks covering 7% – 45% of the area. species from species group o are characteristic of this sub-community. the following are the important species as determined from the synoptic table analysis: diagnostic species: none constant species: andropogon schirensis, asparagus transvaalensis, burkea africana, combretum molle, commelina africana, cyperus leptocladus, elephantorrhiza burkei, englerophytum magalismontanum, heteropyxis natalensis, ozoroa paniculosa, pellaea calomelanos, phyllanthus parvulus, rhoicissus revoilii, setaria lindenbergiana, vangueria infausta dominant species: andropogon schirensis, diheteropogon amplectens, elephantorrhiza burkei, heteropyxis natalensis, protea caffra, setaria lindenbergiana the trees have an average height of 4 m, shrubs 1.5 m and the herbaceous layer 0.8 m. the cover of the woody layer (trees and shrubs) is less than 14% and the herbaceous layer 30% (see table 2). combretum molle–setaria lindenbergiana–diplorhynchus condylocarpon sub-community this sub-community occurs on moderate-to-steep (11°–30°) midslopes (3) at an altitude that ranges from 1038 mamsl to 1620 mamsl. a total of 22 relevés were recorded of which seven were new relevés recorded on the newly acquired land. largeto medium-sized rocks and boulders as well as loose rock occur in this sub-community with a rock cover of 25% – 80%. species from species group p are characteristic of this sub-community. the following are the important species as determined from the synoptic table analysis: diagnostic species: diplorhynchus condylocarpon constant species: burkea africana, combretum molle, commelina africana, englerophytum magalismontanum, pellaea calomelanos, setaria lindenbergiana, vangueria infausta dominant species: andropogon schirensis, brachiaria serrata, burkea africana, combretum apiculatum, combretum molle, croton gratissimus, diplorhynchus condylocarpon, elephantorrhiza burkei, heteropogon contortus, ochna pulchra, pseudolachnostylis maprouneifolia, schizachyrium sanguineum, setaria lindenbergiana the tall shrub diplorhynchus condylocarpon (species group p) is diagnostic for this sub-community and occurs exclusively in this sub-community. the shrub ochna pulchra (species group n) and the grasses aristida scabrivalvis (species group p), brachiaria serrata (species group l) and the tree pseudolachnostylis maprouneifolia (species group n) are prominent throughout this community. the vegetation has a moderate to high cover with the trees (4 m tall) covering 34%, the shrubs (1.5 m tall) covering 29% and the herbaceous layer (0.8 m tall) covering 49% (see table 2). perotis patens–burkea africana woodland this woodland is situated on the midslopes and plateaus of the mountainous areas in marakele. species from species group q are characteristic of this woodland community (see figure 2). the tree burkea africana (species group t) and the grasses setaria sphacelata (species group v), heteropogon contortus (species group t) and schizachyrium sanguineum (species group u) dominate the woody and herbaceous layers, respectively. this plant community has a total of 150 plant species and has the second highest species richness and has one of the highest cover of all the plant communities. this community is divided into the following two sub-communities: perotis patens–burkea africana–fadogia homblei sub-community this tall shrubland sub-community is situated on the southern and north-eastern midslopes and plateaus of marakele. the slope ranges from 2° to 23° and altitude from 1180 mamsl to 1500 mamsl with no rock cover on the plateaus to a rock cover of 60% – 70% on the midslopes. the sub-community is associated with the ib305 land type with moderately deep (50 mm – 600 mm) soil of the mispah and hutton soil forms (see table 1). species from species group r are characteristic of this sub-community. the following are the important species as determined from the synoptic table analysis: diagnostic species: none constant species: burkea africana, faurea saligna, schizachyrium sanguineum, senecio venosus dominant species: burkea africana, calpurnia aurea, hypoestes forskaolii the tree faurea saligna (species group a), the grasses heteropogon contortus (species group t), andropogon schirensis (species group v) and the forb fadogia homblei (species group u) are prominent throughout this sub-community. the tree layer is tall (6 m tall) and together with the herbaceous layer (0.8 m tall) covers 43% each of the area. the shrubs are 1.5 m tall and cover 23% (see table 2). perotis patens–burkea africana–gardenia volkensii sub-community this tall woodland sub-community is found on the lower midslopes (3) and plateaus (1) with a slope of 1° – 7° at an altitude that of 1150 mamsl – 1400 mamsl. rock cover 5% – 70% of the area. mispah and hutton soil forms are characteristic whilst the vegetation is associated with the fa286 land type with moderate deep (50 mm – 600 mm) soil of the mispah and clovelly soil forms (see table 1). species from species group s are characteristic of this sub-community. the following are the important species as determined from the synoptic table analysis: diagnostic species: none constant species: heteropogon contortus, pellaea calomelanos, rhoicissus revoilii dominant species: andropogon schirensis, burkea africana, combretum apiculatum, heteropogon contortus, setaria sphacelata, themeda triandra the grasses schizachyrium sanguineum (species group u), eragrostis curvula (species group b) and the forb kohautia cynanchica (species group i) are prominent locally. this sub-community has a well-developed tree layer that is 6 m tall covering 64% of the area, whilst the shrub layer (1 m tall) is not well developed and covers 13%. the herbaceous layer consists of medium tall grasses (0.8 m tall) that cover 33% of the area (see table 2). kirkia wilmsii–buxus macowanii forest the tall forest community is restricted to relatively high, hot, dry ravine areas on steep midslopes (slope 2° – 29°) in marakele (see figure 2). the altitude varies from 1110 mamsl to 1265 mamsl. rocks cover 40% – 60% of the area. this community is associated with the ib305 land type with shallow (10 mm – 300 mm) soil of the mispah soil form (see table 1). species from species group w are characteristic of this community. the following are the important species as determined from the synoptic table analysis: diagnostic species: buxus macowanii, croton gratissimus, kirkia wilmsii, pouzolzia mixta constant species: cheilanthes viridus, cryptolepis transvaalensis, mimusops zeyheri, olea europaea subsp. cuspidata, pellaea calomelanos dominant species: buxus macowanii, kirkia wilmsii, mimusops zeyheri the trees olea europaea subsp. cuspidata (species group y), berchemia zeyheri, ficus thonningii (species group w) and the woody climber cryptolepis transvaalensis (species group y) are prominent within this community. the tree (7 m tall), shrub (2 m tall) and herbaceous (0.5 m tall) layers have a mean cover of 35%, 60% and 12%, respectively (see table 2). a total number of 58 different plant species was recorded for this community. diospyros whyteana–calodendron capense forest this tall forest is found on midslopes with steep (3° – 30°), dry ravines of the western mountainous section in marakele, at an altitude of 1180 mamsl – 1530 mamsl (see figure 2). rock cover is > 80% where rockslides occurred in the past; however, an important variant, with the same plant species composition, occurs on the floodplain (4) where four drainage lines converse with water always present at the confluence. the alluvial dundee soil form is deeper than > 1.2 m and with no rocks present. this community is located within the ib305 land type (see table 1). species from species group x are characteristic of this community. the following are the important species as determined from the synoptic table analysis: diagnostic species: none constant species: diospyros whyteana, mimusops zeyheri dominant species: calodendron capense, calpurnia aurea, chaetacanthus aristata, combretum moggii, diospyros whyteana, gymnosporia polyacantha, hypoestes forskaolii, maytenus undata, mimusops zeyheri, nuxia congesta, obetia tenax, olea europaea subsp. cuspidata, podocarpus latifolius, strychnos usambarensis, tetradenia brevispicata, trema orientalis, tricalysia lanceolata the trees celtis africana (species group x), pappea capensis (species group y), syzygium guineense (species group ac) and the shrubs grewia occidentalis (species group x) and cryptolepis transvaalensis (species group y) are prominent within this community. the woody layer is dominant and well developed with the tree layer (10 m tall) covering 65%, the shrubs (2 m tall) covering 42%. as expected, because of the dense woody layer, the herbaceous layer is sparse and short (1 m tall) and covers 23% (see table 2). compared to the other plant communities, this plant community has a moderate species richness with 86 different plant species recorded. cyperus albostriatus–podocarpus latifolius forest this forest is situated on moderate to steep midslopes at altitudes of 1380 mamsl – 1730 mamsl. species from species group z are characteristic of this forest community (see figure 2). the trees podocarpus latifolius, curtisia dentata, the small shrub myrsine africana and the forb cyperus albostriatus (species group z) dominate the woody and herbaceous layers, respectively. with a total of 57 different plant species recorded, this plant community has a low species richness. this community is divided into the following two sub-communities: cyperus albostriatus–podocarpus latifolius–rothmannia capensis sub-community the cyperus albostriatus-podocarpus latifolius-rothmannia capensis sub-community is situated on gentle to steep, north, northeast and northwest facing midslopes (4° – 22°) at an altitude of 1380 mamsl – 1660 mamsl. rocks cover from 60% to 80% of the area and consist of large blocks, boulders and loose rocks. this tall forest occurs in the ib305 land type with the mispah soil form that ranges from 10 mm to 300 mm deep (see table 1). species from species group aa are characteristic for this sub-community. the following are the important species as determined from the synoptic table analysis: diagnostic species: tetradenia brevispicata constant species: curtisia dentata, diospyros whyteana, myrsine africana, podocarpus latifolius, rothmannia capensis dominant species: diospyros whyteana, myrsine africana, pittosporum viridiflorum, podocarpus latifolius, pterocelastrus echinatus the tree ilex mitis, the scandent shrub secamone alpini and the forb cyperus albostriatus (species group z) are present throughout this sub-community. the tree layer is very tall (14 m tall) and well developed with a canopy cover of 67%, whereas the shrub layer is 2 m tall and covers 36%. the herbaceous layer is relatively tall (1.3 m) and covers 20% of the area (see table 2). cyperus albostriatus–podocarpus latifolius–widdringtonia nodiflora sub-community the cyperus albostriatus-podocarpus latifolius-widdringtonia nodiflora sub-community is situated on moderate to steep, south and east facing midslopes (3) (slope 6° – 22°) in marakele at an altitude of 1500 mamsl – 1730 mamsl. this tall forest community occurs along perennial watercourses in the protected, moist ravines of the mountainous areas of marakele. because of the high percentage rock cover and poor herbaceous layer, this forest community is protected from fire. rocks cover from 65% to 85% of the area. this community occurs on the ib305 land type associated with the shallow (10 mm – 300 mm) mispah soil form (see table 1). species from species group ab are characteristic for this sub-community. the following are the important species as determined from the synoptic table analysis: diagnostic species: blechnum attenuatum, phylica paniculata, widdringtonia nodiflora constant species: cyperus albostriatus, myrsine africana, podocarpus latifolius dominant species: blechnum attenuatum, widdringtonia nodiflora the trees pittosporum viridiflorum (species group ab), ilex mitis, the scandent shrub secamone alpini (species group z) and the fern pteridium aquilinum (species group ab) are prominent within this community. the tall tree layer (9 m) has a 78% cover, the shrubs (1.3 m tall) cover 22% and the short herbaceous layer (0.5 m) covers 10 % (see table 2). syzygium cordatum–ischaemum fasciculatum woodland this woodland is situated along the upper reaches of the matlabas river in marakele (see figure 2). species from species group ac are characteristic for this woodland community. the trees syzygium cordatum, s. guineense and the grasses ischaemum fasciculatum, andropogon eucomus and phragmites australis (species group ac) dominate the woody and herbaceous layers, respectively. this plant community has a moderate plant species richness with 75 plant species identified and is divided into the following two sub-communities: syzygium cordatum–ischaemum fasciculatum–schoenoplectus corymbosus sub-community the syzygium cordatum–ischaemum fasciculatum–schoenoplectus corymbosus sub-community occurs along the floodplains (4) of the matlabas river at an altitude of 1036 mamsl – 1320 mamsl with the slope not > 3°. rocks cover from 10% to 48% of the area with most of the rocks forming part of the riverbed. this sub-community is associated with the fa286 land type with deep (> 1200 mm) hutton, clovelly and avalon soil forms (see table 1). species from species group ad are characteristic of this sub-community. the following are the important species as determined from the synoptic table analysis: diagnostic species: pseudognaphalium luteo-album, schoenoplectus corymbosus constant species: ischaemum fasciculatum, miscanthus junceus, syzygium cordatum dominant species: ischaemum fasciculatum, miscanthus junceus, phragmites australis, pseudognaphalium luteo-album, syzygium cordatum the sedges xyris capensis and cyperus leptocladus (species group ai) are present throughout this sub-community. the woody layer in this riverine vegetation is well developed with trees (5 m tall) covering 80%, shrubs (2 m) covering 60% and the herbaceous layer (2 m) covering 60% of the area (see table 2). syzygium cordatum–ischaemum fasciculatum–osmunda regalis sub-community the osmunda regalis–syzygium guineense woodland is found on floodplains along the fast-flowing upstream tributaries of the matlabas river (5) at an altitude of 1036 mamsl – 1420 mamsl with the slope not exceeding 2°. because of the higher altitude and narrower streambeds, the rock cover is higher, viz. 40% – 90% and consists of large blocks and boulders along as well as in the streambed. this sub-community is associated with the fa286 land type with deep (> 1200 mm) hutton, clovelly and avalon soil forms (see table 1). species from species group ae are characteristic for this sub-community. the following are the important species as determined from the synoptic table analysis: diagnostic species: osmunda regalis, setaria pallide-fusca constant species: cyperus leptocladus, ischaemum fasciculatum, miscanthus junceus, syzygium cordatum, xyris capensis dominant species: ilex mitis, ischaemum fasciculatum, miscanthus junceus, osmunda regalis, phragmites australis, syzygium cordatum, syzygium guineense the tree, shrub and grass layers cover 50%, 38% and 45%, respectively, with the trees 7 m tall, the shrubs 2 m tall and the herbaceous layer 1.5 m tall (see table 2). leptochloa fusca–fuirena pubescens forbland the leptochloa fusca–fuirena pubescens forbland occurs in the waterlogged and seasonal waterlogged sponges (wetlands) in the eastern and southern high mountain catchment areas of the matlabas-, sterkstroomand sondags rivers (see figure 2). species from species group af are characteristic for this wetland community that has a low species richness of 50 plant species being recorded and is divided into the following two sub-communities. leptochloa fusca–fuirena pubescens–aristida junciformis sub-community the leptochloa fusca–fuirena pubescens–aristida junciformis sub-community is situated as waterlogged to seasonal waterlogged sponges (5) in the southern section of the mountainous catchments area of the matlabas river. this sub-community occurs at an altitude that ranges from 1460 mamsl to 1480 mamsl, with the slope not exceeding 5°. no rocks are present in this sub-community. the vegetation occurs in the ib310 land type with moderately deep (100 mm – 600 mm) soil belonging to the mispah and oakleaf soil forms (see table 1). species from species group ag are characteristic for this sub-community. the following are the important species as determined from the synoptic table analysis: diagnostic species: arundinella nepalensis, ascolepis capensis, berkheya radula, carex cernua, cyperus denudatus, leptochloa fusca, eriochrysis pallida, helichrysum epapposum, pennisetum sphacelatum, pycnostachys reticulata constant species: andropogon huillensis, aristida junciformis, cyperus leptocladus, fuirena pubescens dominant species: aristida junciformis, cyperus denudatus, leptochloa fusca, fuirena pubescens, miscanthus junceus, xyris capensis there are no woody species, and the herbaceous layer is 1.5 m tall covering 75% of the area (see table 2). leptochloa fusca–fuirena pubescens–drosera madagascariensis sub-community this sub-community is situated in seasonal sponges on the eastern and southern section of marakele that feed the sterkstroom and sondags rivers. it occurs at an altitude of 1420 mamsl with the slope not exceeding 3°. no rocks are present in this forbland sub-community. this community occurs in the fa286 land type with hutton, clovelly, katspruit and avalon soil forms with the soil from 50 mm to 1200 mm deep (see table 1). species from species group ah are characteristic for this sub-community. the following are the important species as determined from the synoptic table analysis: diagnostic species: aristida bipartita, chironia purpurescens, cyperus denudatus, cyperus thorncroftii, dierama medium, disa woodii, drosera madagascariensis, eragrostis inamoena, helichrysum aureonitens, kyllinga alba, lobelia erinus, monopsis decipiens, panicum dregeanum, senecio affinis, senecio erubescens, senecio polyodon, verbena bonariensis constant species: andropogon huillensis, fuirena pubescens, hypericum lalandii, kohautia virgata, xyris capensis dominant species: fuirena pubescens, hypericum lalandii, monopsis decipiens, xyris capensis there are no woody species with the herbaceous layer 0.5 m tall covering 85% of the area (see table 2). plant communities 3, 4 and 6 have the highest diversity followed by plant communities 2, 5 and 7. plant community 6 has the highest species richness (155) followed by plant community 7 with 150 species. plant communities 8, 10, 11 and 12 have the lowest diversity and also the lowest species richness of 58, 57, 75 and 50 species, respectively (see figures 2 & 3). discussion this study resulted in the identification and classification of 16 sub-communities and two variants that can be grouped into 12 main plant communities. based on the topography, plant species composition and vegetation structure the vegetation can be grouped into five major groups, namely the: (1) lower midslope and plateau shruband woodlands, (2) high altitude midslope woodland, (3) high-lying plateau and midslope grass-, shruband woodlands, (4) ravine, footslope and drainage line forests and woodland and (5) higher-lying plateau wetlands and forblands. the lower midslope and plateau shrub and woodlands group is represented by the themeda triandra–senegalia caffra woodland (1), the tristachya leucothrix–protea welwitschii shrubland (2) and the perotis patens–burkea africana woodland (7). these plant communities are located at low to medium altitudes on plateaus and lower midslopes with moderate slopes (1° – 16°). the soils are shallow to moderately deep with rock cover that varies from low to high (see table 1). the plant communities belonging to this group have a moderate to high diversity and moderate species richness, indicating them to be stable with a high ecosystem functioning (allan et al. 2011) (see figure 3). the themeda triandra–senegalia caffra woodland (1) corresponds to the acacia caffra–setaria sphacelata short closed woodland as described by van staden (2002) and van staden and bredenkamp (2005). elephants were introduced to marakele national park in 1996 and a monitoring study established that the impact of the elephants on this community is negligible (van staden et al. 2016). figure 3: species diversity (shannon-wiener & gini simpson index) for the main plant communities of the mountainous waterberg section of marakele national park. the vegetation of the tristachya leucothrix–protea welwitschii shrubland (2) is characterised by several woody suffrutex species having traits that allow them to survive low temperatures and fires, which are prevalent on these lower plateaus and midslopes. the vegetation of this community shows affinity with similar plant communities in the bushveld area as described by brown, bredenkamp and van rooyen (1996) in the borakalalo national park. the perotis patens–burkea africana woodland (7) consists of plateaus as well as steep slopes (1° – 23°) (see figure 2). animals tend to utilise the plateaus with deeper soil with palatable grass species often, and consequently some sections were heavily overgrazed in the past, which lead to degradation of the veld and created bare patches and erosion in some areas. although this plant community has the second highest species richness, it has a moderate to high diversity (see figure 3). the high species richness can be ascribed to the various pioneer species present in the degraded areas. the high altitude midslope grass-, shrubland and woodland is represented by plant community 6. the combretum molle–setaria lindenbergiana woodland (6) occurs on steep upper midslopes (see figure 2). the soils are shallow with moderate to high rock cover, consisting of large boulders and rocky sheets, covering 11% – 80%. this community has the highest diversity and species richness of all the communities in the study area (see figure 3). the steep gradients helps to maintain this plant community in a very good to excellent condition because the steep gradient makes it difficult to be affected by wildlife. fire plays a role to maintain the build-up of moribund material. this community has some similarity in terms of species composition with the diplorhynchus condylocarpon–combretum zeyheri woodland as described by brown et al. (1996) in the borakalalo national park and the combretum molle–xerophyta retinervis open to closed woodland in the loskopdam nature reserve (filmalter 2010) and the burkea africana–diplorhynchus condylocarpon variation in marakele national park (van staden & bredenkamp 2005). all of these communities have a similar structure and species composition (see online appendix 1 & table 2). the high-lying plateau and midslope grass-, shruband woodlands group is represented by plant communities 3, 4 and 5. the monocymbium ceresiiforme–andropogon schirensis grassland (3) is located on the plateaus on flat rocky plateaus with a high rock cover of 20% – 45% on shallow sandy to sandy-loam soils (see table 1). this group is typical of high altitude vegetation dominated by grasses with few woody species present. these high-lying areas are exposed to low temperatures and together with the large rocky outcrops provide suitable habitat for grass species. this group has affinity with the protea gaguedi–monocymbium ceresiiforme short open shrubland of the rustenburg nature reserve as described by coetzee (1975), the monocymbium ceresiiforme–loudetia simplex grassland as described by bredenkamp and brown (2003) in the bankenveld in gauteng and shares species such as eragrostis racemosa, loudetia simplex, parinari capensis, pygmaeothamnus zeyheri, aristida transvaalensis, monocymbium ceresiiforme, andropogon schirensis, sphenostylis angustifolia and trachypogon spicatus. the plant species composition of this group has affinity to drakensberg vegetation and is typical of high-altitude cooler aspects although it can also occur on warmer north-facing high-altitude rocky slopes. this group has the second highest diversity and a high species richness (see figure 3) that can be ascribed to the high altitude and little disturbance experienced by wildlife. this group is thought to be in a pristine condition. the indigofera mollicoma–encephalartos eugene-maraisii shrubland (4) of this group is located at high altitudes on moderate to very steep midslopes with shallow soil and moderate to high rock cover (see table 1). the vegetation is mostly high-altitude grassland and shrubland and as such exposed to low winter temperatures. the presence of the endangered waterberg cycad, encephalartos eugene-maraisii along steep scarp of this plant community makes this shrubland an important area from a management perspective to ensure the conservation of this species. it needs full sunlight and is adapted to hot summers and cold winters (grobbelaar 2002). this plant community has high diversity and species richness indicating it to be in a pristine condition with a unique habitat for different species (see figure 3). the tephrosia elongata–protea caffra woodland (5) of this group is characterised by the dominance of especially the tall shrub protea caffra. the plant community occurs at relatively high altitudes on moderate to steep midslopes with shallow soil and medium to high rock cover (table 1). this community is similar to the protea caffra–rhus dentata low closed woodland (van staden & bredenkamp 2005) as well as the protea caffra–loudetia simplex community of the walter sisulu botanical gardens (behr & bredenkamp 1988), the tristachya biseriata–protea caffra cool temperate mountain bushveld and the protea roupelliae cool mountain bushveld of the bankenveld as described by bredenkamp and brown (2003). the group occurs on steep and mostly southern midslopes with high rock cover and moist cool habitats and has affinity with drakensberg vegetation. according to scott (1982) and partridge et al. (1993), certain interglacial periods in the late and middle pleistocene provided opportunity for afromontane (drakensberg) vegetation to establish on the magaliesberg and waterberg. during the last glacial maximum, this vegetation became restricted because of aridification and survive only in protected mountain valleys in the interior of south africa, including the bankenveld. this group has a moderate-high diversity and species richness. the grass, themeda triandra, which is the dominant grass species in this group, is influenced by fire. during a season without fire in the mountainous areas, the occurrence of themeda triandra is lower than just after an intensive fire in the mountainous areas and good rain. themeda triandra is a decreaser grass species (novellie & kraaij 2010) that is utilized well by animals in the mountainous areas of marakele, especially after a fire. the ravine, footslope and drainage line forests and woodland group are represented by plant communities 8, 9, 10 and 11 (figure 2). the kirkia wilmsii–buxus macowanii forest (8) and the diospyros whyteana–calodendron capense forest (9) are located at low to high altitudes on flat to very steep, hot, dry ravine areas and represent the dry forests. the soils are shallow to deep, and the rock cover ranges from 0% to 80% (table 1). the woody layer is well developed and dominates the vegetation. trees range in height from 7 m to 10 m, whilst the shrubs are approximately 1.5 m tall. because of the dense canopies, dry conditions and variable rock cover, the herbaceous layer is not well developed and has a low canopy cover (table 2). similar ravine vegetation is found in other parts of the bushveld areas (brown et al. 1996). these communities have a low-moderate diversity and species richness (figure 3). the cyperus albostriatus–podocarpus latifolius forest (10) of this group represents the wet forests on steep slopes mostly in protected ravines. the soils are shallow, and the rock cover ranges from 60% to 85% (table 1). the cyperus albostriatus–podocarpus latifolius–widdringtonia nodiflora forest sub-community (10.2) of this group is not utilised by wildlife, but fire plays an important role in the pattern and processes of it. because of this area being a forest, there is a regular build-up of moribund leaf material. unfortunately, hot fires occur in the surrounding plant communities every second year, which then escape into this sub-community (van staden 2002). as a result, the damage to the vegetation of this forest is extensive after each fire causing a decrease in woody species. if not effectively managed, it could lead to the encroachment of grass species that will increase fire damage to this forest sub-community and could potentially lead to this sub-community disappearing. this plant community is similar to the podocarpus latifolius–rothmannia capensis tall forest described by van staden (2002), mucina and van staden (2003), van staden and bredenkamp (2006). podocarpus latifolius is a protected tree in south africa and the limpopo province, which makes this plant community even more important. the forest plant communities, including the termitaria bush clumps, were damaged when the elephants were introduced, but the impact on some of the forest communities changed when the northern section (marakele pty ltd.), consisting mainly of valley bottomland and plains with more palatable plant species, was incorporated into marakele and the elephants moved into that area (van staden et al. 2016). these termitaria bush clumps are susceptible to fire and frequent fires could lead to their degradation. the fact that some of these forest communities occur in sheltered ravines and are protected by large boulders protects them from excessive fire damage. it is, however, important that management plans are put in place to prevent accidental fires in these communities that would negatively affect the vegetation composition and structure. the syzygium cordatum–ischaemum fasciculatum woodland (11) corresponds with the syzygium cordatum–miscanthus junceus short thicket as described by van staden and bredenkamp (2005). this community occurs in the narrow ravines along three large tributaries of the matlabas river in the mountainous areas of marakele as well as along the source of the mamba-, sterkstroom– and matlabas rivers and is regarded as being in an excellent condition in terms of plant species composition with mostly climax species present (online appendix 1) and with respect to vegetation structure (table 2). plant community 12 (leptochloa fusca–fuirena pubescens forbland) represents the higher-lying plateau wetlands and forblands group (figure 2) and comprises seasonal waterlogged sponges (wetlands) that feed the various river/stream systems of the park. these relatively high-lying sub-communities are situated on fertile soils with the rock cover ranging from 0% to 90%, with rocks occurring in the fast-flowing upper sections drainage line streams and no rocks in the wetland forbland areas. this group (wetlands and forblands) is susceptible to fire. frequent fires in these wetland communities could lead to the degradation, drying out and compacting of the soil that would cause a serious loss of ecosystem functioning. if fire is, however, kept out completely, it could lead to bush encroachment, which would be detrimental for this group. a three-year burning cycle could be investigated for these wetland plant communities. this group has a low diversity and species richness because of them being relatively homogeneous (figure 3). conclusion the braun–blanquet approach proved to be an efficient way to identify, classify and map floristically defined plant communities in the field. this study resulted in the identification, classification and description of 16 different and clearly recognisable sub-communities and two variants that can be grouped into 12 main plant communities that can be ecologically interpreted. the high-lying vegetation has clear affinities with the bankenveld and drakensberg vegetation and can be classified as inselbergs of high-altitude vegetation. the low-lying and midslope vegetation is typical of the bushveld areas and shows clear affinities to these areas. overall, the plant communities of the high-altitude grassland and shrubland in the midslope areas have the highest diversity. these rocky areas provide various microhabitats to a number of different plant species. according to brown and bezuidenhout (2018), rocky, high-lying areas also act as refuge sites for plants from adjacent areas that are affected by animal utilisation. the forests have a moderate to low diversity and species richness. this can be ascribed to the well-developed tree and shrub layer preventing the growth of large numbers of herbaceous species. the wetland and river plant communities are stable and relatively homogeneous ecosystems with low species richness compared to the other plant communities. sambare et al. (2011) found that the tree species diversity along riverine areas was low, similar to what was found in this study. the mountain vegetation of the marakele has a high species richness and diversity of species that add to the conservation value of the park. this study incorporated previous data as well as new data collected in the study area and has resulted in a more detailed and comprehensive classification of the vegetation than previous studies. this research therefore provides valuable data on the mountainous ecosystems present in marakele. an understanding of the plant communities and their associated habitats is of fundamental importance for compiling sound management and conservation strategies for marakele. this vegetation survey and description provide baseline information that allows for vegetation assessments to conduct carrying capacity as well as similar surveys to be conducted in the future. the expansion of marakele not only contributed to the floristic diversity of the park but also to faunal diversity because of the inclusion of new habitats into the park that made it suitable for the introduction of lions and larger herbivore species that historically occurred in this region. acknowledgements south african national parks, the park managers and staff of marakele national park are thanked for allowing and encouraging the research. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions p.j.v.s. collected the data, did the initial analysis and drafted the manuscript. all authors contributed to further data analyses, data interpretation, further writing, review and editing of the manuscript. funding information this research was financially supported by the university of south africa (unisa) and sanparks. data availability the raw data is available from the first author, p.j.v.s. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors, and the publisher. references allan, e., weisser, w., weigelt, a., roscher, c., fisher, m. & hillebrand, h., 2011, ‘more diverse plant communities have higher functioning over time due to turnover in complementary dominant species’, proceedings of the national academy of sciences 108, 17034–17039. https://doi.org/10.1073/pnas.1104015108 behr, c.m. & bredenkamp, g.j., 1988, ‘a phytosociological classification of the witwatersrand national botanical garden’, south african journal of botany 54, 525–533. https://doi.org/10.1016/s0254-6299(16)31248-0 bezuidenhout, h., 2009, ‘the classification, mapping and description of the vegetation of the rooipoort nature reserve, northern cape, south africa’, koedoe 51(1), a695. https://doi.org/10.4102/koedoe.v51i1.695 bezuidenhout, h., biggs, h.c. & bredenkamp, g.j., 1996, ‘a process supported by the utility bbpc for analyzing braun blanquet data on a personal computer’, koedoe 39, 107–112. bredenkamp, g.j & brown, l.r., 2003, ‘a reappraisal of acocks’ bankenveld: origin and diversity of vegetation types’, south african journal of botany 69, 7–26. https://doi.org/10.1016/s0254-6299(15)30357-4 brown, l.r., bredenkamp, g.j. & van rooyen, n., 1996, ‘the phytosociology of the northern section of the borakalalo nature reserve, south africa’, koedoe 39, 9–24. https://doi.org/10.4102/koedoe.v39i1.279 brown, l.r., du preez p.j., bezuidenhout, h., bredenkamp, g.j., mostert, t.h.c. & collins, n.b., 2013, ‘guideline for phytosociological classifications and descriptions of vegetation in southern africa’, koedoe 55, 1–10. https://doi.org/10.4102/koedoe.v55i1.1103 brown, l.r. & bezuidenhout, h., 2018, ‘ecosystem description and diversity of the jurisdam seekoegat section of the mountain zebra national park, south africa’, south african journal of botany 118, 166–178. https://doi.org/10.1016/j.sajb.2018.07.013 coetzee, b.j., 1975, ‘a phytosociological classification of the rustenburg nature reserve’, bothalia 11, 561–580. https://doi.org/10.4102/abc.v11i4.1502 edwards, d., 1983, ‘a broad-scale structural classification of vegetation for practical purposes’, bothalia 14, 705–712. https://doi.org/10.4102/abc.v14i3/4.1231 filmalter, n., 2010, ‘a vegetation classification and management plan for the hondekraal section of the loskopdam nature reserve’, m tech dissertation, university of south africa, pretoria. grobbelaar, n., 2002, cycads of southern africa, four images bureau & printers, pretoria, p. 334. guiasu, r.c. & guiasu, s., 2010, ‘the rich-gini-simpson quadratic index of biodiversity’, natural science 02, 1130–1137. https://doi.org/10.4236/ns.2010.210140 hill, m.o., 1979, ‘twinspan – a fortran program for arranging multivariate data in an ordered two-way table by classification of individuals and attributes’, cornell university, ithaca, ny. jost, l., 2006, ‘entropy and diversity’, oikos 113, 363–376. https://doi.org/10.1111/j.2006.0030-1299.14714.x land type survey staff, 1988, ‘land types of the maps 2426 thabazimbi, 2428 nylstroom’, memoirs of the agricultural natural resources of south africa no.10, pp. 1–431, department of agriculture and water supply, pretoria. mucina, l. & van staden, p.j., 2003, ‘marakele afromontane forests’, in g. von maltitz (ed.). classification system for south african indigenous forests: an objective classification for the department of water affairs and forestry, council for scientific and industrial research, environmentek report env-p-c 2003-017. mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, strelitzia 19, south african national biodiversity institute, pretoria. novellie, p. & kraaij, t., 2010, ‘evaluation of themeda triandra as an indicator for monitoring the effects of grazing and fire in the bontebok national park’, koedoe 52, 5 pages, https://doi.org/10.4102/koedoe.v52i1.977 partridge, t.c., kerr, s.j., metcalfe, s.e., scott, l., talma, a.s. & vogel, j.c., 1993, ‘the pretoria saltpan: a 200000 year southern african lacustrine sequence’, palaeogeography palaeoclimatology palaeoecology 101, 317–337. https://doi.org/10.1016/0031-0182(93)90022-b roleček, j., tichý, l., zelený, d. & chytrý, m., 2009, ‘modified twinspan 986 classification in which the hierarchy respects cluster heterogeneity’, journal of vegetation science 20, 596–602. https://doi.org/10.1111/j.1654-1103.2009.01062.x sambare, o., bognounou, f., wittig, r. & thiombiano, a., 2011, ‘woody species composition, diversity and structure of riparian forests of four watercourses types in burkina faso’, journal of forestry research 22, 145–158. https://doi.org/10.1007/s11676-011-0143-2 scott, l., 1982, ‘a late quarternary pollen record from the transvaal bushveld, south africa’, quaternary research 17, 339–370. https://doi.org/10.1016/0033-5894(82)90028-x shannon, c.e. & weaver, w., 1949, the mathematical theory of communication, p. 144, university of illinois press, champaign, il. soil classification working group, 1991, ‘soil classification: a taxonomic system for south africa’, memoirs on the agricultural natural resources of south africa 15, 1–262. tichy, l. & holt, j., 2006, ‘juice program for management and classification of ecological data; program manual’, vegetation science group, czech republic, p. 98. van der maarel, e., 2007, ‘transformation of cover-abundance values for appropriate numerical treatment – alternatives to the proposals by podani’, journal of vegetation science 18, 767–770. https://doi.org/10.1111/j.1654-1103.2007.tb02592.x van staden, p.j., 2002, ‘an ecological study of the plant communities of marakele national park’, msc dissertation, university of pretoria, pretoria. van staden, p.j., 2007, a phytosociological study of the newly acquired land as well as the existing land of marakele national park and the contractual section, marakele park (pty) ltd in the limpopo province, south africa, unpublished report, sanparks. van staden, p.j., 2015, ‘aspects of the vegetation ecology of the marakele national park and marakele park (pty) ltd, limpopo province, south africa’, phd thesis, university of pretoria, pretoria. van staden, p.j. & bredenkamp, g.j., 2005, ‘major plant communities of the marakele national park’, koedoe 48, 59–70. https://doi.org/10.4102/koedoe.v48i2.101 van staden, p.j. & bredenkamp, g.j., 2006, ‘a floristic analysis of forest and thicket vegetation of the marakele national park’, koedoe 49, 15–32. https://doi.org/10.4102/koedoe.v49i1.109 van staden, p.j., bredenkamp, g.j., bezuidenhout, h. & ferreira, s., 2016, ‘the effects of elephants and fire on vegetation at marakele national park, south africa’, pachyderm 58, 107–122. filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page138.pdf page 1 matipano.qxd a comparison of woody browse selection by hand-raised, boma-adapted and wild black rhinoceros, diceros bicornis, l. in matusadona national park, zimbabwe g. matipano matipano, g. 2003. a comparison of woody browse selection by hand-raised, bomaadapted and wild black rhinoceros, diceros bicornis, l. in matusadona national park, zimbabwe. koedoe 46(2): 83–96. pretoria. issn 0075-6458. differences in woody browse selection between hand-raised (and subsequently released), boma-adapted and wild black rhinoceros, diceros bicornis, l. were studied in matusadona national park between december 1999 and july 2000. boma-adapted rhinoceros were animals that were subjected to hand-raising and were kept in bomas (enclosures) over night. the feeding behaviour was different between the three rhinoceros groups. all rhinoceros groups utilised and selected for a few browse species in common, at different preference levels in the same habitat types according to season. wild rhinoceroses browsed most in colophospermum-terminalia-combretum woodland in the wet season and in thicket in the dry season. hand-raised rhinoceroses browsed most in colophospermum-terminalia-combretum woodland and boma-adated rhinos in thickets in both the wet and the early dry seasons. hand-raised and bomaadapted rhinos changed their habitats less for browse selection than wild rhinos. this can be ascribed to a relative restriction of home range in the hand-raised group and a herding effect for the boma-adapted animals. these situations might have accounted for differences in seasonal browse selection by the rhinoceros groups. key words: black rhino, hand-raised, browse selection. g. matipano, matusadona national park, department of national parks and wildlife management, p. bag 2003, kariba, zimbabwe. present address: national unversity of science and technology, department of forest resources and wildlife management, p.o. box ac 939, ascot, bulawayo, zimbabwe (gmatipano@nust.ac.zw). issn 0075-6458 83 koedoe 46/2 (2003) introduction the black rhinoceros is predominantly a browser, concentrating on forbs and lowgrowing shrubs; most browsing (ca 60 %) takes place within a 2-m height zone from the ground (owen-smith 1988). black rhinoceroses eat woody plants, forbs, creepers and succulents. however, grass and forbs constitute only a very small proportion of the overall diet, realtive to woody matter (atkinson 1995; hall-martin et al. 1982). the contribution of each type of plant to the diet varies both seasonally and regionally (atkinson 1995). the black rhinoceros select a wide range of plant species according to region and climatic condition (goddard 1968, 1970; hall-martin et al. 1982; oloo et al., 1994; atkinson 1995; muya & oguge 2000). the number of food species eaten decreases with increasing aridity. for example, woody browse species increased in number from 74 in the desert of namibia (loutit et al. 1987), to 113 species in semi-arid regions like sinamatella, hwange national park in zimbabwe (atkinson 1995), and to 191 in the moist ngorongoro crater of tanzania (goddard 1968). although a wide range of species are eaten, black rhinoceros are highly selective for both species type and size class (emslie & adcock 1994). acceptability depends on the level of plant spinescence and the presence of chemical defences (atkinson 1995). since a black rhinoceros is a hind-gut fermenter, it does not benefit from bacterial detoxification of chemicals after ingestion. consequently, it has become adapted to matipano.qxd 2005/12/09 11:20 page 83 selecting for subtoxic levels of phytochemicals by selection from a diversity of food species (muya & oguge 2000). although the rhinoceros can be highly selective for a few food species and plant sizes (emslie & adcock 1994), the species has the ability to feed on a variety of plants, at least in small quantities (oloo et al. 1994). monro (1982) asserted that tame animals have similar food habits to those of their wild counter-parts. however, monro did not distinguish between wild captured and tamed adult animals or young hand-raised animals. atkinson (1995) found that wild captured and captive adult and subadult black rhinos had similar food preferences to those shown by their wild free-ranging counter-parts. thus, the role of domestication in determining food selection is not clear from the literature. this paper aims at studying the influence of hand raising on diet selection by rhinoceroses. it is important to identify species of plants that are important in the diet of the animals in order to improve their management in bomas and to assess suitability of habitats for reintroduction purposes in the park. methods research design matusadona national park stretches from 28º23'e to 28º51'e and from 16º41's to 17º13's. the park is about 1 407 km² in area. an escarpment divides the park into two major geomorphologic landscapes, the semi-arid eutrophic valley floor lying 485–600 m above sea level (a.s.l.) and the wet dystrophic rugged highland section lying 600–1200 m a.s.l. the lowland area is dominated by semi-arid vegetation, mainly colophosperemum mopane woodland while the highland area supports brachystegia-julbernadia woodlands. the study was restricted to the valley floor section of the park. three rhinoceros groups were used to study food selection. the groups were: wild, hand-raised, and boma-adapted rhinos. hand-raised rhinos were animals that had been raised by man and then released into the wild. the hand-raised sample comprised two adults (a male and female) and two subadults (a male and female). two of these hand-raised rhinos, an adult male and a subadult female, were collared. boma-adapted rhinoceroses are defined as animals under semi-captive conditions, where they are kept in bomas (enclosures or kraals) every night. two males and three females, aged between two and four years represented this group. they were herded during the day to browse within a 3–4 km radius of the bomas, and were also fed known quantities of locally collected browse species over-night in the bomas. in addition, they received supplementary artificial feed in the form of horse cubes. habitat classification was based on taylor (1985) who identified seven overall vegetation types. a modification was done to the vegetation classification because the scrub savanna and mixed escarpment ecotone were not easy to define on the ground. the scrub savanna was placed together with the colophospermum mopane vegetation, and the mixed escarpment ecotone woodland was grouped together with the colophospermum-terminalia woodland. in this study, the term ‘thicket’ excludes riverine thickets, which were grouped with riverine woodland as riverine habitats. the lakeshore grassland was excluded because it was not considered important for woody plant browsing. five vegetation types were identified for browse selection studies; colophospermum-terminalia-combretum woodland, riverine, thickets, combretum woodland and mopane vegetation types. the study was limited to assessment of browse quantity due to time constraints. a more comprehensive analysis of food selection would require assessment of both quality and quantity. information was collected between december 1999 and july 2000 inclusively. two seasons, the wet season (december-march) and the early dry season (may-july) were used for data collection. the late dry season (august-early november) was not studied because of time constraints. data were collected only during the day. measurement of browse utilisation and determination of principal food species (pfs) an indirect observation method, involving measurement of previously browsed vegetation was used for both wild and hand-raised groups. two hand-raised rhinoceroses were radio-tracked and located to study their food habits. the uncollared hand-raised rhinoceroses were spoor-tracked and positively identified to confirm their hand-raised status, before information was collected. the indirect method was chosen for reasons highlighted by kotze & zacharias (1993), including the fact that free-ranging animals are not easy to locate, that detection of the observer may influence feeding koedoe 46/2 (2003) 84 issn 0075-6458 matipano.qxd 2005/12/09 11:20 page 84 behaviour, that black rhinoceroses are partly nocturnal, and that vegetation may obscure feeding animals. a random search was made for fresh spoor from which to track wild rhinoceroses. the actual procedure to collect data was the same as for the direct method described below. a direct observation method was used for bomaadapted rhinoceroses when they were being herded during the day. the same method was also used with hand-raised animals when they did not run away from the researcher. a feeding station (quadrat) was defined as a 5-m radius circle, with a browsed plant species as its centre, along the feeding path. the centre of the quadrat was at the base of the first plant that was identified as browsed. all plants that were browsed within the quadrat were considered part of the same feeding station. in each quadrat, the following variables were recorded: habitat type, the plant species selected, the number of fresh bites taken per plant species, and height of the plant. a ‘bite’ was defined as any isolated cut twig or branch; where multiple twigs had been bitten, the definition included all twigs less than five millimetres in diameter and within a hypothetical circle with a diameter of five centimetres (hallmartin et al. 1982). to a certain extent, feeding by other browsers could be excluded because black rhinoceroses feed in a characteristic manner. they prune large twigs (joubert & eloff 1971), severing them at a 30-45º angle (atkinson 1995). plant heights were assigned to height classes. a > 1 m, b > 1 m and <2 m, c > 2 m and <2.5 m, and d < 2.5 m the cut-off point of 2.5 m in classes c & d was chosen because it was assumed that no feeding would take place above 2.5 m from the ground. other information collected at the feeding site was the presence or absence of grazing by rhino and forb browsing. grazing was defined as the actually eating of grass and sedges by rhinos. this allowed a level of herb eating to be defined as the total number of quadrats in which herbs were recorded divided by the total number of quadrats sampled for a given rhino group. the proportional usage (proportion of bites taken) for a given plant species, pu, is given by the number of bites taken from that species divided by the total number of bites taken from all species in that habitat for a given rhino group. for each habitat type, and for each rhino group, pu values were calculated. the relative percentages of bites on woody plants in each vegetation type by a given rhino group by season, were shown graphically. a principal food species (pfs) is defined as that food species consumed in greatest quantities irrespective of its availability or proportional abundance (petrides 1975). pfs represent species that are eaten most by rhinos. in this study a pfs had a pu equal or more than 0.1. chi-square tests were used to analyse the differences in grazing frequencies between different rhino groups. the same tests were used to analyse the total number of bites on woody plants in different vegetation types by season. the chi-square tests were also used to analyse the number of bites taken on woody species common to rhinoceros groups. measurement of browse availability. animals were tracked until a feeding site was encountered. at every fifth quadrat along the feeding path, and only where a woody plant species had been eaten, browse plants (together with other species present within a 5-m radius of the browsed species), were recorded. the fifth quadrat where only grass, sedge and young forb material was consumed was not used to collect data on availability because this aspect of the study was limited to woody browse selection. all woody plants in the fifth quadrat were recorded and identified. every fifth quadrat was equivalent to an availability plot. woody plants were classified by height as described in the above section. plants whose canopies wholly fell above 2.5 m from the ground were regarded as unavailable to rhinos for feeding and were ignored. the proportional availability, pa, of browse plant species in the foraging path was then calculated by dividing the number of times that a species occurred in all the availability plots by the total of the number of occurrences for all species in that habitat type. the most available woody species were defined as those that had a proportional value of equal or greater than 0.1. measurement of food preference indices (fpi) and determination of food selection the fpi of each woody species, whose browseable material fell whithin the 2.5 m zone from the ground, is defined as the proportional utilisation of that species divided by its proportional availability (petrides 1975), i.e. fpi = pu/pa. a preferred or selected species is proportionally more frequent in the diet than it is available to an animal. a rejected species occurs in the diet in a lower proportion than it occurs in the habitat (emslie & adcock 1994). a comparative study on the influence of woody plant height on browse selection by hand-raised, boma and issn 0075-6458 85 koedoe 46/2 (2003) matipano.qxd 2005/12/09 11:20 page 85 wild rhinos (matipano in prep.) was delibarately made subject of another paper. results diet composition the numbers of quadrats in which eating of herbs (grasses, sedges and young forbs) and browsing by hand-raised, wild and bomaadapted rhino groups were recorded by season are shown in table 1. the frequency of browsing was higher than for herb eating for all rhino group by season. both wild and boma-adapted rhinos selected more herb matter in the wet season than in the early dry season but this trend was reversed for handraised rhinos. in the wet season wild rhinoceros ate the highest overall proportion (20.4 %) of herbs while hand-raised rhinos ate the least proportion (12.9 %). in the early dry season the condition reversed with handraised rhinos eating the highest overall proportion (27.8 %) of herb matter while wild rhinos ate the least (7.4 %). the frequency of woody browsing was above 70 % for all rhino groups in the wet and in the early dry seasons. the highest frequency for woody browsing was over 90 % for the wild group in both seasons, and least (82.5 %) for boma-adapted rhinos in the wet season and 74 % for the hand-raised group in the early dry season. the numbers of quadrats in which rhino selected for herbs (grasses, sedges and young forbs combined) and woody species differed significantly among all the three rhino groups (χ2 > 36; p < 0.001; df = 2; table 2). these differences were also found between pairs of rhino groups (χ2 > 4.773; p < 0.05; df = 1) except for forbs between wild koedoe 46/2 (2003) 86 issn 0075-6458 table 1 the frequency of eating herbs (grass, sedge and young forbs) and browsing by different black rhino groups in mnp as expressed by the number of quadrats in which grazing or browsing was recorded (wwild; hhand-raised ; bboma) season herbs woody plants % no. of quadrats % no. of quadrats w h b w h b wet 20.4 12.9 17.5 93 87.1 82.5 early dry 7.4 27.8 13.3 98.4 74.2 86.5 table 2 chi-square tests on the frequency of herbs (grass, sedges and young forbs) and of browsing by different black rhino groups in the wet and early dry seasons, mnp. (significance level is 0.05; ssignificant; nsnot significant; wwild; hhand-raised; b-boma) season rhino groups comments herbaceous plants woody plants wet season w vs h vs b s s w vs h s s w vs b s s h vs b s s early dry w vs h vs b s s season w vs h ns s w vs b s s h vs b s s matipano.qxd 2005/12/09 11:20 page 86 and hand-raised groups in the early dry season (χ2 = 0.067; 0.75 < p < 0.9; df = 1). habitat use for herb eating and browsing wild rhinos fed most on herbs in thickets and in riverine habitats in the wet and early dry seasons, respectively (figs. 1 & 2). an example of a grass species that was grazed in wooded vegetation was panicum maximum. wild rhinos were not observed feeding on herbs in the lakeshore areas. hand-raised rhinos fed most on herbs in the lakeshores and mopane in the wet and early dry season respectively. lakeshore areas were the most important for herb eating for boma-adapted rhinos in both seasons. examples of plants that were grazed in the lakeshore area included the grass panicum repens and sedges. forb browsing included hibiscus sp. and sida cordifolia. the total numbers of bites on woody species in different vegetation types were significantly different among rhino groups for both the wet and in the early dry seasons (χ2 >500, p = 0.00, df = 8). figures 3 & 4 show that all rhino groups browsed most in the colophospermum-terminalia combretum woodland and thickets in both the wet and early dry seasons. hand-raised did not browse in the combretum woodland in the wet season, and boma-adapted rhinos did not browse in the riverine woodland in the early dry seasons. only boma-adapted rhino browsed in the lakeshore area (not shown on the graphs); fed on young plants like colophospermum mopane and acacia sp. seedlings. rhino groups adjusted habitat usage for browsing according to season. wild rhinos used most colophospermum-terminaliacombretum woodland in the wet season and thickets in the early dry season (figs. 3 & 4). they browsed least in combretum woodland and mopane vegetation in both seasons. wild rhinos increased feeding in thickets and riverine areas in the early dry season compared to the wet season. issn 0075-6458 87 koedoe 46/2 (2003) 1 colophospermum-terminalia-combretum woodland 2 thickets 3 combretum woodland 4 riverine vegetation 5 mopane vegetation 6 lakeshore fig. 1. percentage of quadrats in which grazing by rhino groups was recorded in different vegetation types in the wet season. 1 colophospermum-terminalia-combretum woodland 2 thichets 3 combretum woodland 4 riverine vegetation 5 mopane vegetation 6 lakeshore fig. 2. percentage of quadrats in which grazing by rhino groups was recorded in different vegetation. hand-raised rhinos browsed most in colophospermum-terminalia-combretum woodland in both seasons. they browsed least in combretum woodland in the wet season and least in mopane vegetation in the early dry season. hand-raised rhinos increased browsing in thickets and combretum woodland in the early dry season compared to the wet season. boma-adapted rhinos browsed most in thickets and least in riverine habitats in both seasons. they increased browsing in colophospermummatipano.qxd 2005/12/09 11:20 page 87 terminalia-combretum woodland in the early dry season compared to the wet season. the boma-adapted rhino group showed the least flexible and the wild group the most flexible pattern of habitat use for browsing with season. woody browse utilisation and principal food species (pfs) all rhino groups utilised 87 and 68 woody browse species in the wet and early dry seasons, respectively. rhino groups shared a few woody browse species within different habitats both in the wet (table 3) and early dry seasons (table 4). differences in the levels of browsing on common woody species were tested between rhino groups for the wet season (table 5) and for the early dry season (table 6). all rhino groups fed on most of the common woody browse species at different level of selection in both seasons (χ2 >8; p < 0.005, df = 2), except for baphia massaiensis in thickets in the wet season, which showed no difference in levels of selection between groups (χ2 =2.489; 0.05<p<0.1; df = 1). in most cases, any pair of tested rhino group was shown to select differently for common woody species (χ2 >3.956; p<0.05; df = 1). however, the level of feeding by wild and hand-raised rhino groups did not differ sigkoedoe 46/2 (2003) 88 issn 0075-6458 table 3 the numbers of bites on woody species common to rhino groups in different habitats in the wet season (w-wild; hhand-raised; bboma) colophospermum-terminalia-combretum habitat thicket vegetation species rhino group species rhino group w h b w h b c. zeyheri 105 31 88 c. zeyheri 10 102 c. apiculatum 12 71 136 c. apiculatum 9 93 k. tettensis 32 23 350 k. tettensis 185a 306a 915a d. quiloensis 51 14 26 d. quiloensis 11 15 c. spinosa 7 144 36 c. celastroides 8 12 a. nigrescens 24 5 15 t. stuhlmannii 26 11 c. pubscens 24 8 37 b. massiensis 10 12 6 c. celastroides 11 12 b. massaiensis 19 6 combretum woodlands riverine woodland species rhino group species rhino group w h b w h b k. tettensis 59 142 d. ceneria 9 6 d. condylocarpon 6 14 mopane vegetation species rhino group w h b c. mopane 13 40 k. tettensis 11 25 c. gratissimus 28 12 t. stuhlmannii 26 6 c. apiculatum 36 14 a species qualifying as pfs and are common to two or all rhino groups. matipano.qxd 2005/12/09 11:20 page 88 issn 0075-6458 89 koedoe 46/2 (2003) table 4 the number of bites on woody species common to rhino groups in different habitats in the early dry season (w-wild; h-hand-raised; b-boma) colophospermum-terminalia-combretum habitat thicket vegetation species rhino group species rhino group w h b w h b c. zeyheri 20 21 283 c. zeyheri 7 50 129 c. apiculatum 57a 207a 383a c. apiculatum 10 162 k. tettensis 6 6 182 k. tettensis 205a 337a 930a d. quiloensis 168 48 10 d. quiloensis 80a 12a 14 c. spinosa 66a 270a 6 b. massaiensis 24 85 63 s. spinosa 7 58 s. trichoclada 67a 110a c. pubscens 8 10 t. stuhlmannii 28 25 combretum woodlands riverine vegetation species rhino group species rhino group w h b w h b c. zeyheri 7 21 60 d. quiloensis 124 45 c. apiculatum 6 207 41 s. kunthianum 18 8 k. tettensis 6a 82a b. massaiensis 66 48 c. celastroides 7 17 mopane vegetation species rhino group w h b c. apiculatum 7a 54a d. quiloensis 78a 30a a species qualifying as pfs and are common to two or all rhino groups nificantly for the following: karomia tettensis in colophospermum-terminalia-combretum woodland vegetation, baphia massaiensis in thickets, and dichrostachys ceneria in riverine habitat in the wet season (χ2 < 2.91; p > 0.05; df = 1; table 5). these rhino groups did not differ significantly on their feeding on combretum zeyheri, k. tettensis, carphalea pubescens in colophospermumterminalia-combretum woodland and on b. massaiensis in combretum woodland in the early dry season (χ2 < 2.25; df = 1; table 6). wild and boma-adapted rhino groups had no significant differences when feeding on c. zeyheri, a. nigrescens and c. pubescens in colophospermum-terminalia-combretum woodland vegetation, and on d. quiloensis, b. massaiensis and combretum celastroides in thickets in the wet season (χ2 < 1.509; p > 0.05; df = 1; table 5). the feeding behaviour was completely different between the wild and boma-adapted rhinos in the early dry season (table 6). hand-raised and boma-adapted rhino groups did not differ significantly in their feeding levels on combretum celastroides in colophospermum-terminalia-combretum woodland and on baphia massaiensis in thickets in the wet season (χ2 < 1.09; p > 0.05; df = 1; table 5). these rhino groups did not differ significantly on their feeding on terminalia stuhlmannii in colophospermum-terminalia-combretum woodland and on diospyros quiloensis and b. massaiensis in thickets in the early dry season (χ2 < 2.25; matipano.qxd 2005/12/09 11:20 page 89 koedoe 46/2 (2003) 90 issn 0075-6458 table 5 chi-square tests on the number of bites on woody species common for black rhinos groups in different habitats in the wet season (significant level is 0.05; wwild; h-hand-raised; b-boma rhinos) habitat type species rhino group significant (s)/ not significant (ns) colophospermum-terminaliac. zeyheri w vs h vs b s combretum woodland w vs h s w vs b ns h vs b s c. apiculatum w vs h vs b s w vs h s w vs b s hvs b s k. tettensis w vs h vs b s w vs h ns w vs b s h vs b s d. quiloensis w vs h vs b s w vs h s w vs b s h vs b s c. spinosa w vs h vs b s w vs h s w vs b s h vs b s a nigrescens w vs h vs b s w vs h s w vs b ns h vs b s c. pubescens w vs h vs b s w vs h s w vs b ns h vs b s c. celastroides h vs b ns b. massaiensis w vs b s thickets k. tettensis w vs h vs b s w vs h s w vs b s h vs b s b. massaiensis w vs h vs b ns w vs h ns w vs b ns h vs b ns d. quiloensis w vs b ns c. zeyheri w vs b s c. apiculatum h vs b s t. stulhmannii h vs b s c. celastroides w vs b ns combretum woodland k. tettensis w vs b s riverine vegetation d. ceneria w vs b ns d. condylocarpon w vs b s c. mopane w vs b s k. tettensis w vs b s mopane c. gratissimus h vs b s t. stuhlmannii h vs b s c. apiculatum h vs b s matipano.qxd 2005/12/09 11:20 page 90 p > 0.05; df = 1; table 6). the most utilised woody plant species for the wet and early dry seasons (pu > 0.10) were ragarded as principal food species (pfs). rhino groups shared a few common pfs including k. tettensis in thicket vegetation in the wet season and early dry season (tables 3 & 4) and combretum apiculatum in colophospermum-terminalia-combretum woodland in the early dry season (table 4). hand–raised and wild groups shared the following additional pfs: catunaregan spinosa in colophospermum-terminalia-combretum woodland, d. duiloensis in mopane vegetation and thickets, and s. trichoclada in thickets. hand-raised and boma-adapted rhinos shared the following additional pfs: k. tettensis in combretum woodland and c. apiculatum in mopane. some species that registered as pfs in the lower height categories were not in the d height class. for example, acacia nigrescens, carphalea pubscens and baphia massaiensis were utilised at levels ranging between 11 % and 13 % in the lower height categories (< 1 m height) but were utilised at lower levels (from 6 % to 8 %) in the d height class. this was for the colophospermum-terminalia-combretum woodland. woody browse preference and selection all rhino groups preferred totals of 85 and 70 woody species in the wet and early dry seasons, respectively. the fpis were expressed with species-size. many of the species preferred by one group received low fpi scores for another; thus there was little similarities in preferences between groups in a given vegetation type. some species were highly selected or highly preferred in the lower height categories, but rejected in the d height class. examples of such species were bauhinia tomentosa and baphia massaiensis which had preference indices of 1.465 and 4.421 in the lower height categories, compared to indices of 0.991 and 0.892 in the d height class, respectively, in the colophospermumterminalia-combretum vegetation and in thickets. issn 0075-6458 91 koedoe 46/2 (2003) fig. 3. percentage of bites on woody plants in different vegetation types by rhino groups in the wet season. fig. 4. percentage of bites on woody plant species in different vegetation type by rhino groups in the early dry season. 1-colophospermum-terminaliacombretum woodland 2-thichets 3-combretum woodland 4-riverine vegetation 5 mopane vegetation 1-colophospermum-terminaliacombretum woodland 2-thickets 3combretum woodland 4-riverine vegetation 5-mopane vegetation matipano.qxd 2005/12/09 11:20 page 91 table 6 chi-square tests on the number of bites on woody species common to black rhino groups in different habitats in the early dry season in mnp (significance level is 0.05; wwild rhinos; h-hand-raised rhinos; b-boma rhinos) habitat type species rhino group significant (s)/ not significant (ns) colophospermum-terminaliac. zeyheri w vs h vs b s combretum woodland w vs h ns w vs b s h vs b s d. quiloensis w vs h vs b s w vs h s w vs b s h vs b s c. apiculatum w vs h vs b s w vs h s w vs b s h vs b s c. spinosa w vs h vs b s w vs h s w vs b s h vs b s k. tettensis w vs h vs b s w vs h ns w vs b s h vs b s c. pubscens w vs h ns s. spinosa w vs h s t. stuhlmannii h vs b ns thickets k. tettensis w vs h vs b s w vs h s w vs b s h vs b s c. zeyheri w vs h vs b s w vs h s w vs b s h vs b s b. massaiensis w vs h vs b s w vs h s w vs b s h vs b ns d. quiloensis w vs h vs b s w vs h s w vs b s h vs b ns c. apiculatum h vs b s s. trichoclada w vs h s combretum woodland c. zeyheri w vs h vs b s w vs h s w vs b s h vs b s c. apiculatum w vs h vs b s w vs h s w vs b s h vs b s k. tettensis h vs b s c. celastroides h vs b s b. massaiensis w vs b ns riverine d. quiloensis w vs h s s. kunthianum w vs h s mopane c. apiculatum h vs b s d. quiloensis w vs h s koedoe 46/2 (2003) 92 issn 0075-6458 matipano.qxd 2005/12/09 11:20 page 92 discussion and conclusion summary of group differences patterns of habitat use for grazing and for browsing were different for all rhino groups. in general, the three rhino groups shared different feeding behaviours for the same habitat according to season: the percentages of browsing and grazing were different, food species selected differed, or a narrow range of species was shared, or the number of bites on shared species differed, and the preference indices on common species differed hand-raised rhinos avoided using combretum woodlands in the wet season, could not make a selection of food species in thickets in the early dry season. thickets are important dry season habitats for wild rhinos (atkinson 1995). hand-raised rhinos selected some species like erythroxylum zambesiacum and cauboura glauca that are not normally favoured by rhinos. boma and handraised rhinos selected for colophospermum mopane and euclea divinorum in the wet season. wild rhino preferred euclea divinorum during the dry season (atkinson 1995). hand-raised and boma-adapted rhinos showed anomalous feeding behaviour. a male hand-raised animal was observed feeding on impala carcass (woodfine pers. comm.). plastic material was recorded in the dung of the same rhino, which could indicate anomalous feeding behaviour. on imire game ranch, hand-raised rhinos licked animal carcass and bones (poole 1995). this might mean that hand-raised rhinos were failing to obtain a diet with adequate minerals and they resorted to eating bones. bones provide minerals, including calcium. explaining differences in feeding behaviour many factors affect dietary selection, including season, resource distribution (atkinson 1995; pellew 1984), plant phenology and size, and the management history of the animal. all rhino groups changed their home ranges spatially and in turn adjusted habitat use (matipano 2000), resulting in differing food utilisation and especially selection with season within habitats. however, the nature of this adjustment differed for each group. oloo et al. (1994) highlighted reasons for this seasonal variability in food species in the diet. in this study, seasonal requirement for succulence (high-moisture food) could have been a major factor governing diet selection; choice of riverine habitats in the dry season would have exposed wild rhinos to a ‘new’ range of plant species from which to select. riverine vegetation has a higher moisture content providing more ‘green bite’ during the dry season. herding restricted the movement of boma-adapted rhinos such that they could not use riverine areas in the early dry. herding of boma-adapted rhinos partly explained dietary differences between them and wild rhinos. secondly, rhinos changed browse selection in response to browse phenological changes. reduced palatability of browse in the dry season will induce a shift to more palatable species, as well as to other plant parts of the same species (atkinson 1995). the valley floor is semi-arid and most species are deciduous. as drier conditions prevail, evergreen species contribute more to the diet than deciduous species (atkinson 1995). species like cleistochlamys kirkii were eaten by wild rhinos, and euphorbia sp. and euclea divinorum by hand-raised and boma-adapted rhinos in the early dry season. some species that dropped leaves later in the dry season like strychnos spinosa, strychnos madagascariensis, catunaregan spinosa and diospyros quiloensis were favoured in the early dry season. in this study, hand-raised rhinos took stems (ca 15 mm diameter) from strychnos spinosa in the early dry season. in this study, the quality of food eaten was not measured. thus, it was not possible to confirm the observation of hall-martin et al. (1982) that black rhinos modified their diet to suit environmental conditions by reducing the intake of high fibre plants in favour of more succuissn 0075-6458 93 koedoe 46/2 (2003) matipano.qxd 2005/12/09 11:20 page 93 lent and nutritive species. however, plant phenology might have influenced diet selection through change in quality. browse quality is influenced by toxic phytochemical and nutrient content in plants. plant height was another factor influencing selection. some shrub species, only appeared as pfss or were selected in the lower height classes and not in the d height class. examples were the shrubs such as dichrostachys ceneria and carphalea pubscens in colophospermum-terminalia-combretum vegetation, holarrhena pubescens and croton gratissimus in thickets. black rhinos feed mostly on those plants within the 50-120 cm height category from the ground (owensmith 1988). individual plants < 20 cm in height were included in the study to determine availability. this inclusion probably resulted in underestimating preference for those species that were most used when above 20 cm in height. in this study boma and hand-raised rhinos selected for more species than the wild group. other studies have shown a high number of food species for wild black rhino (goddard 1970; hall-martin et al. 1982; oloo et al.; 1994; atkinson 1995). in this study, all species that could be utilised were not measured since data for the late dry season were not collected. further, some plant species were only recorded as utilised and preferred, but as not available. this means that such species could have been highly sought for and selected, e.g. euphorbia spp. alternatively, these were rare species only taken by chance, and were not of any dietary importance, e.g. crossopteryx febrifuga and manilkara mochisa in combretum woodlands, bridelia carthertica, pterocarpus brenanii and combretum collinum in mopane in the case of boma-adapted rhinos. a third explanation is simply that sampling effort was too low to provide meaningful data for spatially scattered species. the distribution of food resources is not the same for a given habitat type. individual rhinos probably perceive resource patchiness differently based on management history, which modifies feeding behaviour. it is likely that the boma group fed on most of the food patches encountered. the reasons for this are that animals: were restricted to the same small feeding ground; had limited nocturnal feeding in bomas, and could have been permanently hungry. they fed for longer time in the afternoons than the wild group, probably to compensate for the time they are put in bomas before sunset (poole 1995); had reduced ability to select food because of some reliance on supplementary feed; browsed in a group, whereby the feeding of one rhino encouraged that of adjacent animal. this would account for the high intensity of feeding by boma animals, both in terms of the number of species per station and in terms of the number of bites per plant; and after weaned young calves are stressed and hungry, and may feed on poisonous plants (tyler 1999 pers. comm.). wild rhinos select highly for a few species (emslie & adcock 1994) and feed in a spatially uneven manner (kotze & zacharias 1992). this study, showed that boma-adapted rhinos were less selective for browse species than either hand-raised or wild rhino group. boma-adapted rhinos, and to an extent the hand-raised group, apparently browsed at sites closer to each other and took more bites per feeding station than wild rhinos. however, this feeding behaviour was to a large extent imposed upon the boma group, because they were restricted to certain habitats by herding. for hand-raised rhinos, the feeding behaviour was influenced by the habit of restricting home ranges close to human settlement (matipano 2000). black rhinos have high dietary selection for a few species and high dietary diversity by taking other species at lower preferences. this feeding strategy can be explained by the need to obtain the full complement of food quality requirements, and at the same time limit the amount of each species taken at one time in order to keep intoxication tolerable koedoe 46/2 (2003) 94 issn 0075-6458 matipano.qxd 2005/12/09 11:20 page 94 (emslie & adcock 1994; muya & oguge 2000). however, this does not adequately explain the higher diversity of food species selected by both boma and hand-raised rhinos. one speculative explanation is that, having been exposed initially to a narrower range of browse species, these groups could have been used to relatively higher levels of phytotoxic chemicals, and could perhaps tolerate feeding for longer on a single species, as well as on a wider range of species encountered. this reduced selectivity is also shown by hand-raised rhinos which settled in the vicinity of human settlement, where they were exposed to human litter. plastic material was recorded in the dung of such rhinos, which could indicate reduced selective capacity. a second explanation is that rhinos reduce the number of bites per plant with the increasing times of feeding on the same species by day (poole 1995). the boma group, therefore, ingested only small amounts from the narrow range of species they were given in bomas overnight and relied on artificial feed. hand-raised rhinos at imire most preferred cubes to other food items in both the wet and dry seasons (poole 1995). horse cubes, compared with natural browse were easily digested and rhinos may be feeding on more species simply to obtain roughage. boma-adapted rhinos did not show signs of poor body condition. there might have been an associative effect of foods. the supplements might have improved the efficiency of utilising of browse. wild rhinos might have eaten a relatively low diversity of food species as a strategy to reduce the intake of toxins. bias in data collection might be an alternative reason for recording a low diversity of browse species taken by wild rhinos. it was difficult to locate and follow tracks for reasonable distances, compared to the ease of direct observation on boma-adapted rhinos, and radio tracking of hand-raised rhinos. in addition, lists of plants utilised were probably incomplete because studies using more than one technique yield longer lists (muya & oguge 2000). these investigations on selected aspects of the behavioural ecology of hand-raised rhinos were seasonally biased. the late dry season studies, especially on preference, were not carried out due to time constraints. further, studies were limited to daytime, ignoring nocturnal activities of rhinos. the results obtained were not representative of the annual requirements for rhinos. if a home range shifted, available habitats would also change, together with available browse species. monitoring or studying animals during periods of resource scarcity is vital to provide knowledge on the ecology and management of rhinos. the method used to collect data ensured that it was the rhino not the researcher determining availability of browse and that plants included were within the reach of a rhino (atkinson 1995). this made it possible to study availability and utilisation at the same time. however, with appropriately designed studies, separate utilisation and availability studies can be done (du toit pers. comm.). du toit’s proposal would be done with the assumption that the researcher can determine feeding sites in a manner equivalent to that of a rhino. the problem with this proposed approach is that any differences between the perception of browse availability by the researcher and the rhinos would not be established. conclusions in this study, semi-tamed hand-raised rhinos had different and anomalous feeding behaviour, compared to wild rhinos. hand-raised rhinos selected fewer browse plant species than boma-adapted rhinos, but more than wild rhinos. however, plant species selection may, in fact, be less important than nutritional levels. rhinos may take different plants but without significantly affecting nutritional intake. there is, thus, a need for chemical analysis of plants utilised by rhino groups, especially in the dry season when food quality is critical. information is issn 0075-6458 95 koedoe 46/2 (2003) matipano.qxd 2005/12/09 11:20 page 95 required on levels of toxins, protein, lignin and nutrients, e.g., na, ca and zn. herding boma-adapted rhinos during the day is importatnt as it gives them opportunity to learn about their release environment prior to reintroduction. the approach at mnp of providing young rhinos with both artificial feed and indigenous browse whilst they are in bomas, and of herding them in the field during the day, allows animals to acquire some necessary information on habitat and food selection for survival in the wild after release. this approach to raising rhinos differs from, and is superior to, complete captive management. under conditions of total captivity, exposure to natural conditions does not occur. on the other hand, complete freemanagement in conservancies and other larger areas is better than hand-raising. acknowledgements financial support from the european commission is greatly appreciated. i would like to thank dr. c.a.m. attwell and mr. r.f. du toit for their guidance. i also thank mr. e. chidziya of the department of national parks and wildlife management for his support during the study. i am grateful to e. muchuchutiti and t. nyakashaya for their assistance during field work. finally, i thank the department of national parks and wildlife management for granting permission to carry out the study in matusadona national park. references atkinson, s.j. 1995. maintenance of captive black rhinos (diceros bicornis) on indigenous browse in zimbabwe: energetics, nutrition and implications for conservation. msc. thesis. university of zimbabwe. emslie, h. r. & k. adcock. 1994. feeding ecology of the black rhinoceros. pp. 65–81. in: penzhorn, b.l. & n.p.l. kriek (eds.). proceedings of symposium on rhinos as game ranching animals. onderstepoort. 9–10 september 1994. wildlife group of the south african veterinary association. republic of south africa. goddard, j. 1968. food preferences of black rhinoceros populations. east african wildlife journal 6: 1–18. goddard, j. 1970. food preferences of black rhino in the tsavo national park. east african wildlife journal 8: 145–161. hall-martin, a.j., t. erasmus & b.p. botha. 1982. seasonal variation in the diet and faeces composition of black rhino, diceros bicornis in addo elephant national park. koedoe 25: 63–82. joubert, e. & f.c.eloff. 1971. notes on the ecology and behaviour of the black rhinoceros, diceros bicornis, l. 1758 in south west africa. madoqua ser. 1(3): 5–53. kotze, d.c. & p.j.k. zacharias. 1993. utilisation of woody browse and habitat by the black rhino (diceros bicornis) in western itala game reserve. african journal of forest science 10(1): 36–40. loutit, b. d., g.w. louw & m.k. seely. 1987. first approximation of food preferences and the chemical composition of the diet of the desert dwelling diceros bicornis bicornis l. madoqua 15: 35–54. matipano, g. 2000. effects of hand raising on balck rhinoceros, diceros bicornis, l. in matusadona national park, zimbabwe: an evaluation of ranging behaviour, habitat use and browse selection. msc. thesis, university of zimbabwe, harare. monro, r.h. 1982. an appraisal of some techniques to investigate the feeding ecology of large herbivores with reference to study of impala in the northern transvaal. african journal of ecology 20: 71–80. muya, s. m. & n.o. oguge. 2000. effects of browse availability and quality on black rhino (diceros bicornis michaeli groves 1967) diet in nairobi national park, kenya. journal of african ecology 38(1): 62–71. oloo, t.w., r. brett & t.p. young 1994. seasonal variation in the feeding ecology of black rhinos (diceros bicornis l.) in laikipia, kenya. african journal of ecology 32: 142–157. pellew, r. 1984. food consumption and energy budgets of the giraffe. journal of applied ecology 21: 141–159. petrides, g. a. 1975. principal foods versus preferred foods and their relations to stocking rate and range conditions. biological conservation 7: 161–169. poole, a. 1995. feeding ecology of the black rhino, diceros bicornis in a highveld habitat. bsc. (hons.) thesis. bangor university, uk. taylor, r.d. 1985. the response of buffalo, syncerus caffer (sparrman) to the kariba lake grassland (panicum repens l.) in matusadona national park. phd thesis, university of zimbabwe, harare. koedoe 46/2 (2003) 96 issn 0075-6458 matipano.qxd 2005/12/09 11:20 page 96 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <feff004e00e4006900640065006e002000610073006500740075007300740065006e0020006100760075006c006c006100200076006f0069006400610061006e0020006c0075006f006400610020005000440046002d0061007300690061006b00690072006a006f006a0061002c0020006a006f006900640065006e002000740075006c006f0073007400750073006c00610061007400750020006f006e0020006b006f0072006b006500610020006a00610020006b007500760061006e0020007400610072006b006b007500750073002000730075007500720069002e0020005000440046002d0061007300690061006b00690072006a0061007400200076006f0069006400610061006e0020006100760061007400610020004100630072006f006200610074002d0020006a0061002000520065006100640065007200200035002e00300020002d006f0068006a0065006c006d0061006c006c0061002000740061006900200075007500640065006d006d0061006c006c0061002000760065007200730069006f006c006c0061002e0020004e00e4006d00e4002000610073006500740075006b0073006500740020006500640065006c006c00790074007400e4007600e4007400200066006f006e0074007400690065006e002000750070006f00740075007300740061002e> /ita <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> /nor <feff004200720075006b00200064006900730073006500200069006e006e007300740069006c006c0069006e00670065006e0065002000740069006c002000e50020006f00700070007200650074007400650020005000440046002d0064006f006b0075006d0065006e0074006500720020006d006500640020006800f80079006500720065002000620069006c00640065006f00700070006c00f80073006e0069006e006700200066006f00720020006800f800790020007500740073006b00720069006600740073006b00760061006c00690074006500740020006600f800720020007400720079006b006b002e0020005000440046002d0064006f006b0075006d0065006e0074006500720020006b0061006e002000e50070006e006500730020006d006500640020004100630072006f0062006100740020006f0067002000520065006100640065007200200035002e00300020006f0067002000730065006e006500720065002e00200044006900730073006500200069006e006e007300740069006c006c0069006e00670065006e00650020006b0072006500760065007200200073006b00720069006600740069006e006e00620079006700670069006e0067002e> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 about the author(s) anna hübner german corporation for international development, dong hoi, vietnam tuan phon ly school of hotel and tourism management, hong kong polytechnic university, hong kong trương s.h. châu german corporation for international development, dong hoi, vietnam citation hübner, a., ly t.p. & châu, t.s.h., 2019, ‘corrigendum: good governance and tourism development in protected areas: the case of phong nha-ke bang national park, central vietnam’, koedoe 61(1), a1601. https://doi.org/10.4102/koedoe.v61i1.1601 note: doi of original article: http://dx.doi.org/10.4102/koedoe.v56i2.1146 corrigendum corrigendum: good governance and tourism development in protected areas: the case of phong nha-ke bang national park, central vietnam anna hübner, tuan phon ly, trương s.h. châu published: 06 dec. 2019 copyright: © 2019. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. in the version of this article published earlier, the name of the second author was given incorrectly. the correct name should be tuan phon ly instead of lý t. phong in the ‘author’ and ‘how to cite this article’ sections. the author apologises for any inconvenience caused. abstract introduction place, sense of place and place attachment how attachment levels differ based on demographics research methods and design results discussion conclusion acknowledgements references about the author(s) anneli douglas division of tourism management, university of pretoria, pretoria, south africa jan-albert wessels department of environmental sciences, school of ecological and human sustainability, university of south africa, pretoria, south africa jenny pope school of science, edith cowan university, joondalup, australia research unit for environmental science and management, north-west university, potchefstroom, south africa integral sustainability, south fremantle, australia angus morrison-saunders school of science, edith cowan university, joondalup, australia research unit for environmental science and management, north-west university, potchefstroom, south africa mike hughes environmental and conservation sciences, murdoch university, perth, australia citation douglas, a., wessels j.-a., pope, j., morrison-saunders, a. & hughes, m., 2019, ‘measuring kruger visitors’ place attachment to specific camps’, koedoe 61(1), a1559. https://doi.org/10.4102/koedoe.v61i1.1559 original research measuring kruger visitors’ place attachment to specific camps anneli douglas, jan-albert wessels, jenny pope, angus morrison-saunders, mike hughes received: 25 oct. 2018; accepted: 12 june 2019; published: 17 sept. 2019 copyright: © 2019. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract tourists become emotionally, physically and socially attached to national parks as they become familiar with the park’s settings and endow it with value. researchers have pointed out that place attachment leads to environmentally responsible behaviour and higher levels of visitor satisfaction. therefore, increasing the level of attachment that visitors feel is vital for park and camp managers, and to do so a greater understanding of the various dimensions of it is needed. while attachment to parks has been evaluated previously, attachment to specific camps in parks has not been done. the main purpose of this research study was to measure the extent to which visitors to the tamboti and satara camps in the kruger national park feel attached to these camps. we also determined whether differences exist between visitors in terms of the level of attachment that they experience towards these camps. finally, we established the variables that influence place attachment. a self-administered paper-based questionnaire was distributed to visitors to the tamboti and satara camps, with 201 questionnaires completed. the results show that visitors generally have a neutral feeling towards the camps. furthermore, the differences in visitors’ levels of attachment could be attributed to their nationality, wild card membership and frequency of visits. various managerial implications are drawn and recommendations made on how to increase place attachment to these camps. conservation implications: this results indicate that visitors do not show particularly strong attachment towards tamboti and satara. recommendations are given for camp managers to increase place attachment to the camps. if camp managers can succeed in fostering stronger levels of attachment to these camps, visitors are more likely to display environmentally responsible behaviour in the camps, with positive conservation implications. keywords: place attachment; kruger national park; camps; south africa; south african national parks. introduction national parks hold varied and often distinctive features (reimann, lamp & palang 2011) and provide the ideal setting for social and psychological exchanges to take place between people and the environment (ramkissoon, weiler & smith 2012). when these exchanges tie an individual to a park, they become attached to the park, as they familiarise themselves with the park’s settings and place a value on it (kyle, graefe & manning 2005). research into place attachment in the context of south african national parks remains scant, even though south africa is home to some of the most well-known national parks globally. place attachment helps us understand visitor behaviour (kyle et al. 2004). klenosky et al. (2007) state that negative place attachment occurs when specific elements of a location are in conflict with an individual’s self-identity or do not satisfy an individual’s needs. negative place attachment will likely prevent an individual from visiting a location, whereas positive place attachment will encourage visitation to a location. walker and chapman (2003) show that positive place attachment may influence an individual’s willingness to take part in protecting a place, while vaske and kobrin (2001) speculate that positive attachment may significantly influence environmentally beneficial behaviours (e.g. picking up litter, conserving water and recycling), especially in a nature-based context such as a national park. previous studies have shown that tourists who are highly attached to a place will even persuade others to adopt behaviours that benefit the environment (ramkissoon, smith & weiler 2013a). to assist parks in fostering place attachment, a greater understanding of it is needed (ramkissoon & mavondo 2014). consequently, various calls have been made for more investigation into place attachment (dredge 2010; tsai 2012; yuksel, yuksel & bilim 2010). to date, place attachment to specific national parks has been measured (hwang, lee & chen 2005; ramkissoon et al. 2013a), but not to specific accommodation settings in these parks. the purpose of this article was thus to measure place attachment to specific camps, and not to the park in general, as this has been done before. in south african national parks, facilities are provided at camps within parks, with these camps owned and run by the park. it would thus make sense that visitors could become attached to a specific camp setting and that this level of attachment should be measured, rather than place attachment to the park in general. the main purpose of this study was thus to measure the extent to which visitors are attached to the tamboti and satara camps in the kruger national park. in addition, we assess whether visitors’ levels of place attachment differ across age groups, nationalities, gender, level of education and others. finally, we establish whether certain variables have a stronger influence on levels of place attachment than other variables. the remainder of the article is structured as follows: firstly, we discuss the concepts of place, sense of place (sop) and place attachment, after which place attachment and the dimensions thereof are clarified. next we explain the methodology used whereafter we discuss the results. finally, conclusions are drawn and managerial recommendations are given. place, sense of place and place attachment ‘place’, as a concept, has both tangible and intangible dimensions; place is more than simply the location of a site. according to halpenny (2010), the value and meaning of place are given by individuals and society, and presented in groups, cultures and individuals. researchers are increasingly acknowledging the value of the less quantifiable and less tangible advantages that individuals get from nature and places such as protected areas (barendse et al. 2016), for example the recreational, spiritual, experiential and educational exchanges with nature that add to the well-being of a human (millennium ecosystem assessment 2005). more importantly, the extent to which one appreciates such benefits is often dependent on one’s ability to engage with or form an association with the natural environment (hinds & sparks 2008). ramkissoon, smith and weiler (2013b) noted the overabundance of terms in the literature explaining the association between people and spatial settings, including connectedness to nature (gosling & williams 2010), community attachment (perkins & long 2002), place attachment (altman & low 1992), sop (jorgensen & stedman 2001) and neighbourhood attachment (lewicka 2010), among others. authors such as yuksel et al. (2010) opine that sop, place identify and place dependence are forms of place attachment, whereas others such as kyle et al. (2004b) propose that sop is the extensive term and place attachment is a subdimension. chen, dwyer and firth (2014) explain the difference between sop and place attachment as follows: sop is made up of two components: relationship to place, which entails all of the various ways that people relate to places, or the kinds of ties individuals can form with a setting, and place attachment, which entails the depth and sorts of attachments to one specific place (cross in chen et al., 2014). relationship to place reveals the individual–place connection in relation to how this connection is made. for example, an individual is connected to a place if he or she was born there. the relationship to place fluctuates according to the nature of the relationship rather than for psychological reasons. on the other hand, the level of attachment between a person and a place differs and may be impacted by other aspects such as memorable events, level of satisfaction and length of residence. place attachment can reveal a person’s psychological change in their connection with a specific place, and is a vital question to ask if we want to understand tourists after holidaying at a destination (chen et al. 2014). consequently, this study focused on the variable component of sop: place attachment. place attachment place attachment originates from attachment theory (bowlby 1969) and is drawing substantial attention from tourism researchers (mcleod & busser 2012; ramkissoon et al. 2013b) who utilise it to discover recreationists’ or tourists’ attachment behaviour and feelings (hwang et al. 2005). williams et al. (1992) define place attachment as the emotional bond that is formed between an individual and a specific setting. jorgensen and stedman (2001) go further by explaining place attachment as not only an emotional bond, but also a cognitive and functional bond with a location. in recreation and leisure, hidalgo and hernandez (2001) are of the opinion that place attachment is personified in the feelings and emotions linked to a recreational setting. some authors claim that when tourists become attached to tourism destinations, they display affective identification and dependence (schultz 2000) and grow an inseparable connection with the location (kals, schumacher & montada 1999). moore and graefe (1994) further state that while place attachment links people with their natural environment, it also induces identification, gratification and concern for a distinctive area (harris, brown & werner 1996). gosling and williams (2010) have found that when people grow attached to a specific location (place), they demonstrate care and concern for the protection of the environment, and become more aware of current matters affecting the environment (lee 2011). this then increases their commitment to the growth and conservation of natural resources (scannell & gifford 2010), while at the same time leading them to exhibit environmentally responsible behaviours, such as willingly picking up litter (halpenny 2006), recycling as well as conserving water (vaske & kobrin 2001) and preventing environmental damage (stedman 2002). cheng, wu and huang (2013) assert that most leisure tourism researchers assess place attachment with two constructs: place dependence, which is linked to the usefulness of a location for a leisure pastime, and place identification, which is a symbolic or emotional connection to a location (kyle absher & graefe 2003). ramkissoon et al. (2013a) conversely see place attachment as a multidimensional construct including place affect (kals & maes 2002), place dependence (stokols & shumacker 1981), place social bonding (scannell & gifford 2010) and place identity (prohansky 1978), with each construct significantly different from the other (kyle et al. 2005; ramkissoon et al. 2012). in our study, we also view place attachment as a multidimensional construct as ramkissoon et al. (2013a) and devine-wright and clayton (2010) found that construing place attachment, as a singular concept, is rather deceptive. they emphasised the need for future research to see place attachment as a multidimensional construct, as this would aid in developing research questions that stay true to vital theoretical concepts (stedman 2002). these four subconstructs of place attachment – place dependence, place identity, place affect and place social bonding – are defined next. place dependence according to ramkissoon et al. (2012), national parks are theoretically the perfect setting to foster place dependence. place dependence can be defined as ‘how well a setting serves goal achievement given an existing range of alternatives’ (jorgensen & stedman 2001:234). individuals and groups measure the functionality of places, that is, the degree to which they assist the accomplishment of specific actions. the physical characteristics of the area or destination (williams & vaske 2003) personify this functional attachment and are significantly linked to the distinctive qualities that the setting is perceived to have (williams et al. 1992). place dependence also points to the location’s relative quality when compared to other locations (halpenny 2010). scannell and gifford (2010) opine that the more someone associates with the physical characteristics of a setting, the less enthusiastic he or she will be to substitute the setting for another. according to hammitt, backlund and bixler (2006), place dependence is also a form of bonding, where places that gratify numerous needs generally result in a more entrenched, profound and all-encompassing place dependence, than places where fewer needs are satisfied (stokols & shumaker 1981). moore and graefe (1994) opine that extensive interaction with a place because of place dependence may produce place identity. place identity research studies have shown that experiences with nature create place identity (clayton & opotow 2003; prohansky 1978). place identity is defined as ‘an individual’s cognitions, beliefs, perceptions or thoughts that the self is invested in a particular spatial setting’ (jorgensen & stedman 2001:238). place identity describes the symbolic link between a person’s self-identity and his or her physical environment (prohansky 1978:155; stedman 2002). people usually create a sense of identity with a setting (budruk, thomas & tyrrell 2009; halpenny 2010) because of its distinctiveness or uniqueness from other settings (twigger-ross & uzzell 1996), resulting in a psychological investment with the place as time passes (williams & patterson 1999). place affect place affect mainly depends on emotions, allowing individuals to develop their feelings towards a place and giving significance to it (tuan 1977). in the past, place affect used to be combined with place identity measures, but more recently, researchers such as halpenny (2010) and ramkissoon et al. (2013b) have started to regard it as a separate subdimension of place attachment. ramkissoon et al. (2013b) show that it is likely for natural environments, such as national parks, to create feelings of psychological well-being for visitors, thus further stimulating positive emotions in visitors (hartig et al. 1996), leading to increased levels of emotional attachment (hinds & sparks 2008). natural environments are likely to increase positive emotions (hartig et al. 1996), leading to stronger affective connections with those environments (hinds & sparks 2008; ramkissoon et al. 2013a). furthermore, vining (1992) linked place affect with nature protective behaviours. place social bonding places form an essential part of social relationships. socially based place bonds refer to the experiences individuals get from social exchanges at a specific site (scannell & gifford 2010). consequently, ‘social bonding’ has been developed as a dimension of place attachment, to better express the emotional and social parts of place attachment (ramkissoon et al. 2012). as places form an essential part of social relationships, social bonding comes from the exchanges between friends and family that are reliant on a specific site (hidalgo & hernandez 2001; ramkissoon et al. 2012). according to tonge et al. (2015), we attach meaning to a place because of the recollection of experiences in the places that we shared with our loved ones (kyle et al. 2005), which often results in a feeling of group belonging (low & altman 1992). in their assessment of social and physical place attachment, hidalgo and hernandez (2001) determined that the social attachments were stronger than setting attachments. how attachment levels differ based on demographics research can help us in predicting the behaviour of groups and individuals in agreement with the meanings, values and feelings that they attach to a place (cass & walker 2009). people’s connection with nature is always a function of their value systems (chan, satterfield & goldstein 2012), which are particular to their context and constantly changing with time. place attachment can vary according to certain demographical variables (rollero & de piccoli 2010). according to ednie, daigle and leahy (2010), the addition of sociodemographic variables is vital for research directed towards establishing management implications as it is easier to gear actions towards members of a specific sociographic group than towards those with high levels of place attachment. when considering gender, results differ. in a number of studies, men and women demonstrate similar levels of place attachment (brown, perkins & brown 2003; ednie et al. 2010; lewicka 2005), while in others women show stronger connections to places (hidalgo & hernandez 2001; rollero & de piccolli 2010). in the study of kyle, graefe and manning (2004), there were significant differences between men and women, with men showing a greater place attachment than women. age also plays a vital role in place attachment (ng, kam & pong 2005; pretty, chipuer & bramston 2003). ng et al. (2005) reported a positive influence of older age on the place belonging dimension of place attachment. in their study, kyle et al. (2004a) also found older respondents’ level of attachment higher than younger respondents. similarly, lewicka (2008) found higher levels of attachment in older generations. this was confirmed by ednie et al. (2010), who found a significant difference in terms of age where respondents in their low-attachment cluster were significantly younger than respondents in their high-attachment cluster. the role of level of education in place attachment has not been studied sufficiently. lewicka (2005) and rollero and de piccoli (2010) established that the level of education was a negative forecaster of place attachment, explaining that people with higher levels of education are more geographically moveable and thus less reliant on a specific place. kyle et al. (2004), on the other hand, found no significant difference in terms of education, while ednie et al. (2010) also reported no significant differences in levels of education. furthermore, lewicka (2008) is of the opinion that education is of less importance in predicting place attachment. in terms of travel parties, ednie et al. (2010) reported that respondents with high place attachments were more probable to be travelling as part of a group with family and/or friends and less probable to be part of a guided group or organisation. also, moore and scott (2003) found a relationship between frequency of use and positive attachment. from the above discussion, it is clear that place attachment is made up of different dimensions, and that individuals differ in terms of their levels of attachment to a specific place. the same seems to be true in the context of south african national parks, and barendse et al. (2016) raised specific questions that still remain unanswered in terms of place attachment (sense of place) experiences in south african national parks, for example, how place attachment experiences vary across groups of visitors. for this reason, we hypothesise that: h1: groups of visitors differ in the levels of attachment that they experience towards specific camps in the kruger. another question that remains unanswered is the extent to which certain variables influence the level of place attachment that visitors experience towards specific camps. it is thus hypothesised that: h2: variables such as age, wild card membership, camp visited, travelling party, number of visits and gender influence the level of place attachment that visitors experience towards specific camps in the kruger. the methodology used to test these hypotheses is explained next. research methods and design study site the kruger national park (knp) is the flagship conservation and tourism product offering within the south african national parks system. hausmann et al. (2017) found that visitors gave high ratings (between 4 = perceived and 5 = highly perceived) to the sop dimensions in three parks, namely, knp, isimangaliso wetland park and table mountain national park. even so, research into place attachment in the context of specific camps within this park remains scarce. some studies have been conducted in terms of sop, with the results suggesting that although sop is accounted for in national parks and environmental management, it remains an underdeveloped concept denoting a substantial void in the way that we understand the link between park management and visitor experiences (ament et al. 2017; barendse et al. 2016; hausmann et al. 2017). to measure place attachment to camps, the selected camps had to comply with specific criteria. firstly, the selected camps should have the capacity to accommodate enough visitors so that comparisons between camps could be drawn. secondly, camps in the southern kruger were preferred because this would allow for the collection of more data (both the capacity and occupancy levels of camps in the southern part of the kruger are higher than camps in the northern part of the kruger). thirdly, we wanted to use contrasting camps to assess if place attachment differed between the camps (the camps chosen should not only be different in name, but also in all other aspects). if similar camps were chosen, we would not have been able to attribute the differences to anything. fourthly, we wanted to include one main camp and one satellite camp. lastly, attachment to the camp should not be obvious. for example, given the lower sabie’s popularity, one would assume high levels of place attachment, and thus no need to measure it. the camps that met all these criteria were tamboti and satara. satara can be described as an older, more established (in the traditional knp style) camp with permanent chalets, and ample infrastructure and facilities. on the other hand, tamboti is a newer (more modern) camp, with rustic, semi-structured eco-tents and limited infrastructure and facilities. satara is open to day visitors, whereas tamboti does not allow day visitors. interestingly, in terms of overnight guests’ overall satisfaction scores with these camps (for the period may to june 2017), we see that satara’s overall satisfaction score is slightly lower than the average satisfaction score of visitors to all camps in the marula region of the kruger. in contrast, the overall satisfaction score of visitors to tamboti is higher than the average satisfaction score of visitors to all camps in the marula region (sanparks 2019). questionnaire development, sampling, data collection and analysis a paper-based questionnaire was distributed to a convenience sample drawn from both day and overnight visitors, domestic and international, to satara and tamboti camps in the kruger national park. the questionnaires were distributed from 23 to 28 july 2017. the first section of the questionnaire asked some demographic questions, including age, gender, education level, nationality, travelling party, wild card membership and frequency of visits. the next section measured place attachment (adapted from ramkissoon et al. 2013b). for the place attachment construct, four dimensions were included in this study: place dependence (three items), place identity (three items), place affect (three items) and place social bonding (three items). each item was measured on a five-point likert scale (1 = strongly disagree, 5 = strongly agree). tourists were approached in both the accommodation areas as well as the public areas of the two camps. fieldworkers were instructed to attempt to vary the age, gender and nationality of respondents. a total of 201 responses were obtained (day visitors collected at satara = 34; overnight in satara = 124; overnight in tamboti = 43). in order to meet the purpose of the study satisfactorily, diverse techniques for data analysis were used. the descriptive methods contributed in describing the data in terms of age groups, gender representations and levels of education, while inferential methods permitted us to draw certain deductions about the larger population of travellers and their place attachment to two specific camps in the kruger. because of modifications necessary to customise ramkissoon et al.’s (2013b) scale to our context, the place attachment scale was subjected to an exploratory factor analysis (efa). t-tests and analyses of variance (anovas) were used to determine how groups of visitors differ in terms of their place attachment. lastly, multivariate analysis of variance was used to explore the effect of the key identified variables (age, camp visited, nationality, gender, wild card membership, level of education, travelling party and number of visits) as well as their possible interaction effects in explaining the variance in the two factors of place attachment. multivariate analysis of variance (manova) is a technique that is used to test for the difference in two or more vectors of means. the multivariate tests (pillai’ trace, wilks’ lambda, hotelling’s trace and roy’s largest root) all test the manova null hypothesis, namely, that the mean on the composite variable is the same across groups. the test thus determines the equality of a composite of the means (optimised to yield the maximum possible f-ratio) across groups. the focus of the identification of meaningful effects will be in using wilks’ lambda in conjunction with the partial eta squared value. although the pillai–bartlett criterion is considered the most robust and powerful test statistic, the wilks’ lambda is used as it provides an indication of the variance not accounted for by the combined dependent variables with (1 – λ) the variance that is accounted for by the best linear combination of dependent variables, which enables the explorative understanding of key effects. table 1 provides the demographic profile of the respondents. from the table, it is clear that almost an equal number of men (48%) and women (52%) completed the questionnaire. almost two-thirds of respondents indicated that they are wild card members (a loyalty card that entitles members to reduced entrance fees at all national parks in south africa). the biggest age group responding to the questionnaire were the 31–50 year-olds. forty-five percent of respondents indicated that they hold a postgraduate degree. nearly two-thirds of respondents visited the park with family and 60% of respondents have visited the park more than three times. official sanparks data on the gender of visitors were not available; however, in a large sample survey (n = 4369), conducted by the agency in 2018 on overnight visitors to the knp, the gender split of respondents was 59% men and 41% women. according to sanparks, however, this is not necessarily an indication of actual visitation, because the online survey that they use to collect it is sent to the email address of the person making the booking, which is dominated by men. our sample is similar to the sample of kruger, viljoen and saayman (2017). the biggest age group (35%) in their sample was between the ages of 45 and 59 years. again, according to kruger et al. (2017), visitors to kruger generally hold a degree or diploma, and in our sample this was the same. in terms of frequency of visits, sanparks survey data show that 61% of their visitors have been to the kruger for one to three times in the preceding 3 years, and 23% between four and nine times. this is similar to our sample. table 1: demographic profiles of respondents. ethical considerations ethical approval for undertaking the research was obtained from the university of pretoria, faculty of economic and management sciences ethics committee (protocol no. ems014/17). the research was also conducted under the consent and approval of tourism development and marketing division of south african national parks. all the participants in the study took part voluntarily after they were informed of the objectives of the study and the completion of an informed consent agreement. all participants were entitled to withdraw from the study at any point. the completed questionnaires were also completed anonymously and confidentially. results descriptive statistics visitors were asked to indicate the extent to which they agreed with a number of statements regarding the specific camps at which they were surveyed. place attachment was measured on a five-point scale (1 = strongly agree and 5 = strongly disagree). results are shown in table 2. it is interesting to note that visitors to the two camps did not have a particularly strong place attachment to the specific camps, even though some preferred the particular camps for their specific settings and facilities. of the four dimensions measuring place attachment, respondents seemed to agree the strongest with the place dependence dimension, followed by place identity and place affect. place social bonding scored the lowest mean. when compared, day visitors to satara felt a consistent stronger attachment to the camp than overnight visitors to satara and tamboti. a plausible explanation could be that day visitors could include repeat visitors who come to the camp because they are attached to it, for example, people living near the knp. only in terms of place dependence did tamboti visitors score higher agreement means than day and overnight visitors to satara. the low agreement mean given to place social bonding is also interesting, especially given the fact that 95% of respondents indicated that they visit the park with friends and family. the most probable explanation could perhaps be that social bonding does not adequately explain a visitor’s attachment to tamboti and satara. table 2: place attachment. exploratory factor analysis because of the modifications necessary to customise ramkissoon et al.’s (2013b) scale to our context (ramkissoon et al. 2013b) measured place attachment in the context of national parks, whereas our study measured place attachment in the context of camps in the parks), the place attachment scale was subjected to an exploratory factor analysis (efa) using maximum likelihood extraction and direct oblimin rotation (a confirmatory factor analysis was conducted first; however, model fit indicated an inadequate fit, and therefore an efa was conducted to determine the underlying dimensional structure). the purpose of the efa was to ensure unidimensionality and internal consistency of this construct in the present context. the kaiser–meyer–olkin measure of sampling adequacy (0.908) and the bartlett’s test of sphericity which was significant (p = 0.000) both indicate that a factor analysis is appropriate. the efa was conducted using a sample of 201 respondents who were intercepted at different locations in the two camps. no items were eliminated from the scale, with the number of items remaining at 12. these 12 items used to measure the place attachment construct were loaded onto two factors (see table 2). based on the items, which showed that respondents would continue their attachment to specific camps, factor 1 was labelled ‘continued attachment’, while factor 2 was labelled ‘interrupted attachment’, because respondents indicated that if their visitation to the camp would stop, they would disappoint or lose contact with their friends and family, and hence interrupt their attachment to the specific camp. these two factors thus become composites of specific variables and include specific items that are a facet of the broader place attachment dimension (hair et al. 2014). internal consistency was evaluated using cronbach’s alpha and both factors showed a measurement greater than 0.9, indicating strong levels of internal consistency (nunnally 1978). together, these two factors explain 75.42% of the variance. t-tests and analyses of variance differences in levels of attachment between groups were measured in terms of age, the camp visited, nationality, gender, wild card membership, level of education, travelling party and number of visits. only those where significant differences were shown in at least one of the factors are given in table 3. from the table it is clear that south africans have a higher level of continued attachment than international visitors have, confirming the results of hausmann et al. (2017), who found that more experienced national tourists have a higher sop perception. as expected, wild card members showed a stronger attachment than non-wild card members, for both factors of place attachment. those visitors with only a matric or high school qualification had the highest levels of place attachment, followed by those with a postgraduate degree. visitors with a matric or high school qualification felt a stronger continued attachment than visitors with a degree, while visitors with a postgraduate qualification showed a stronger continued attachment than visitors with a degree. this contradicts the results of kyle et al. (2004) and ednie et al. (2010) who reported no significant differences between levels of education. in terms of number of visits, those who have visited most frequently showed a stronger continued attachment than those who have only visited 1–3 times. at the same time, those who have visited most frequently also had a stronger attachment than those who have visited between 4 and 20 times. interestingly, day visitors to satara were more attached to the camp than overnight visitors to satara. at the same time, overnight visitors to tamboti showed a stronger continued attachment than overnight visitors to satara. furthermore, day visitors to satara felt a stronger interrupted attachment to the camp than overnight visitors to tamboti. the results thus support hypothesis 1 in that groups of visitors differ in the levels of attachment that they experience towards specific camps in the kruger. table 3: analysis of variance and t-tests. multivariate analysis of variance multivariate analysis of variance was used to explore the effect of the key identified variables, namely, age, the camp visited, nationality, gender, wild card membership, level of education, travelling party and number of visits as well as their possible interaction effects in explaining the variance in the two factors of place attachment, namely, continued attachment and interrupted attachment. multivariate analysis of variance has greater power to identify an effect because it can identify whether groups are different along a combination of variables, whereas anova can identify only if groups are different along a single variable field. the manova measures whether or not the independent grouping variable simultaneously explains a statistically significant amount of variance in the dependent variable (continued attachment and interrupted attachment). unfortunately, the manova cannot predict which groups are significantly different from each other, it only tells us that at least two groups are different and that the independent variables influence some patterning of response on the dependent variable. the set of two dependent variables considered were not highly correlated (> 0.7); thus, multi-collinearity was not present and thus were adequate for the purposes of the manova analysis. the analysis revealed the following key statistical significant multivariate main and interaction effects: wild card membership: wilks’ λ = 0.839, f = 4.211, p <. 0.021, partial eta squared = 0.161. power to detect the effect was 1. camp visited: wilks’ λ = 0.797, f = 2.639, p <.0.039, partial eta squared = 0.107. power to detect the effect was 1. wild card membership, age and gender: wilks’ λ = 0.870, f = 3.288, p < 0.047, partial eta squared = 0.130. power to detect the effect was 1. from the above it is evident that a number of key variables influence the set of attachment factors, with the wild card membership, camp visited and the interaction between wild card membership, age and gender being the most influential. the fact that wild card membership explains the variance in the two factors of place attachment is not surprising, as one would expect members to have different levels of attachment than non-members. the factor camp visited also influences some patterning of the response on the dependent variable, again providing support for hypothesis 1, that visitors differ in the levels of attachment they feel towards specific camps. even though alone age and gender do not influence the variance in the two factors, it is interesting that the interaction between wild card membership, age and gender simultaneously explains a statistically significant amount of variance in the two attachment factors. the results from the manova thus support the second hypothesis that certain variables influence the level of place attachment that visitors experience towards specific camps in the kruger. the fact that wild card membership showed significant differences in place attachment in both the anova and manova results should encourage sanparks to find ways of boosting their membership sales. yet, visitors will only become loyal to a park (and camp) if they are satisfied with the overall experience, which should provide more motivation for camp and park management to scrutinise the overall satisfaction levels of visitors, and create strategies to improve the scores of items with which visitors are not particularly happy. discussion bearing in mind that place attachment leads to more environmentally responsible visitors and increases visitor satisfaction, determining how to increase levels of place attachment has to be an important topic for kruger national park management. for this reason, the main purpose of this research study was to measure the extent to which visitors to the tamboti and satara camps in the kruger national park feel attached to these camps, while also determining whether differences exist between groups in terms of the level of attachment that they experience towards these camps. finally, we established whether certain variables have a stronger influence on place attachment than other variables. the results show that respondents did not have a particularly strong place attachment to tamboti and satara, even though some preferred the particular camps for their specific settings and facilities. it is expected that place attachment to the park in general will be higher as hausmann et al. (2017) found in their study. thus, attachment to the camps does not equate to attachment to the park, and visitors could be attached to the park in general, but not to specific camps. the kruger national park provides a variety of camps that may fulfil visitors’ needs and some visitors might be attached to one camp, whereas others might be attached to another camp. our results offer possible answers to the questions raised by barendse et al. (2016) in a previous article. in answer to their question on the link between individual and shared experiences in fostering place attachment, social bonding (shared experiences) scored the lowest mean of all four attachment dimensions in our results, contradicting the results of hidalgo and hernandez (2001) who determined that social attachments are stronger than setting attachments. in response to their (barendse et al. 2016) question relating to how attachment (sop) experiences vary across nationalities of tourists, our results showed that south africans have a higher level of continued attachment than international visitors. as expected, wild card members showed a stronger place attachment than non-wild card members because it is obvious that loyal park visitors will be more attached to the camps. this confirms previous research studies showing that frequency of use is a significant predictor of place attachment (lewicka 2005; moore & scott 2003). interestingly, those visitors with only a matric or high school qualification had the highest levels of place attachment, confirming the results of lewicka (2005) and rollera and de piccoli (2010). the manova tests showed a number of key variables influencing the set of attachment factors, with wild card membership, the camp visited and the interaction between wild card membership, age and gender being the most influential. this confirms the research of poira, reichel and biran (2006) who found that the same place might have diverse meanings for diverse individuals. a number of suggestions are made to increase visitors’ attachment to tamboti and satara. firstly, place attachment should be developed through long-term, frequent and positive experiences with the camps. camp managers should ensure that visitors are satisfied when they depart from the camp to ensure future visitation. the more frequent the visits, the more likely visitors are to become attached (lewicka 2005; williams & patterson 1999). when taking the overall customer satisfaction scores for tamboti and satara into account, it is evident that two of the items with which visitors experienced the lowest satisfaction were ‘nature experience’ and ‘caravan, camping and accommodation’ at the camps. camp management should thus show concerted efforts in trying to increase satisfaction with these items (sanparks 2019). engagements with other people also affect place attachment (eisenhauer, krannich & blahna 2000). this is of particular importance given the low rating to place social bonding in our results. camp management should encourage participation and social interaction in touristic activities in these camps to produce increased levels of attachment (prayag & ryan 2012). in terms of place dependence, high scores imply that visitors are dependent on the resources of the camp to enjoy their specific tourist activities (kyle et al. 2004). in our study, place dependence scored higher than the other three dimensions, even though the score was still neutral. camp managers should thus endeavour to fulfil and meet tourists’ real needs and services so that they can develop a sense of dependence on the camp (cheng et al. 2013). efforts should thus be made to improve tourist experiences by ensuring that visitors are satisfied with the infrastructure provided and intangible qualities (exoticness and reputation) (prayag & ryan 2012). various authors have shown that place attachment leads to more environmentally responsible behaviour. at the same time, the higher visitors perceive the value of the camp experience, the more environmentally responsible they become. when tourists feel that they benefit from the experience, they are more likely to identify more strongly with the environment. this, in turn, will stimulate their sensitivity towards and concern for the environment, which will shape their environmentally responsible behaviour. it is thus suggested that camp management should increase the satisfaction with the camp experience in order to promote environmentally responsible behaviour in the camp, decreasing the damage to the environment (chiu, lee & chen 2014). this could be attained by ensuring that the camp shows proof of good management, support for biodiversity and reinforcement of sustainable and responsible consumption, which are likely to be valued by visitors (ramkissoon & mavondo 2014). if the tourists see the camp’s commitment to conservation, it is likely to encourage them to show environmentally responsible behaviour (lee 2011). the theoretical contribution of this article lies in the fact that it adds to the debate on whether place attachment should be seen as a multidimensional or bi-dimensional construct. as opposed to previous studies indicating four dimensions (devine-wright & clayton 2010; ramkissoon et al. 2013), our results only showed two, with the one dimension, labelled interrupted attachment, showing very low levels of attachment. this article is not without limitations. the results of this study cannot be generalised to all kruger visitors as the sample was non-random. future research should look at how place attachment experiences differ for each camp and each national park (based on feedback from visitors) and whether place attachment experiences should be taken into account in the design, establishment and management of protected areas (barendse et al. 2016). furthermore, hausmann et al. (2017) mentioned the lack of understanding about how biodiversity, and experiences related to biodiversity, influence tourists’ place attachment when visiting protected areas. future research should look into this. conclusion the purpose of this study was to measure the extent to which visitors are attached to the tamboti and satara camps in the kruger national park. in addition, we assessed whether visitors’ levels of place attachment differ. finally, we established whether certain variables have a stronger influence on levels of place attachment than other variables. the research identifies two factors of place attachment, confirming the research of cheng et al. (2013) and contradicting the results of ramkissoon et al. (2013a). the research also finds that visitors to tamboti and satara have neutral levels of attachment to the camps. furthermore, anova results show that levels of attachment between visitors differ in terms of nationality, wild card membership, level of education, number of visits and the camp visited. the manova results indicate that the main variables that have an influence on place attachment are wild card membership, the camp visited and the interaction between wild card membership, age and gender. our results confirm the results of hausmann et al. (2017) showing differences between nationalities. the results also confirm the results of moore and scott (2003), who showed differences in place attachment in terms of frequency of visits. contrary to research by ednie et al. (2010), our research did not show differences in levels of place attachment in terms of the group that the respondents travel with. also, kyle et al. (2004) found no significant difference in terms of level of education, whereas our results show significant differences. this article contributes to the current literature regarding place attachment, and specifically place attachment in the context of camps in the kruger national park. this is the first time that place attachment is measured in the context of camps within parks. in doing so, sensible management conclusions have been reached. in conclusion, place attachment (sense of place) is a vital feature of conservation from both legal and conceptual viewpoints (barendse et al. 2016). as such, results such as ours could be used to inform park management plans and conservation action. the study responds to a call from barendse et al. (2016) who suggested that place attachment should be accounted for from both social and natural lenses to enable debates of the ‘desired future conditions’ of conservation areas from both social and ecological viewpoints (williams & stewart 1998). our results provide the social lens and viewpoints and thus stimulate communication and interdisciplinary learning not only between natural and social scientists but also between management, science and all stakeholders (chapin iii & knapp 2015). acknowledgements the authors would like to thank the tourism development and marketing division of south african national parks for their support of the project. competing interests the authors declare that they have no conflicts of interest. authors’ contributions a.d. was the lead author of this article. the remainder of the authors all contributed in reviewing the article, developing the measurement instrument and collecting the data. funding information this work is based on the research supported in part by the national research foundation of south africa (grant number 114916). data availability statement data are available upon request from the corresponding author. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references altman, l. & low, s., 1992, place attachment, plenum press, new york. ament, j., moore, c., herbst, m. & cumming, g., 2017, ‘cultural ecosystem services in protected areas: understanding bundles, trade-offs, and synergies’, conservation letters 10(4), 440–450. https://doi.org/10.1111/conl.12283 barendse, j., roux, d., erfmann, w., baard, j., kraaij, t. & nieuwoudt, c., 2016, ‘view shed and sense of place as conservation features: a case study and research agenda for south africa’s national parks’, koedoe 58(1), a1357. https://doi.org/10.4102/koedoe.v58i1.1357 bowlby, j., 1969, attachment and loss: attachment, vol. 1, basic books, new york. brown, b., perkins, d. & brown, g., 2003, ‘place attachment in a revitalizing neighbourhood: individual and block levels of analysis’, journal of environmental psychology 23, 259–271. https://doi.org/10.1016/s0272-4944(02)00117-2 budruk, m., thomas, h. & tyrrell, t., 2009, ‘urban green spaces: place attachment and environmental attitudes in india’, society and natural resource 22, 824–839. https://doi.org/10.1080/08941920802628515 cass, n. & walker, g., 2009, ‘emotion and rationality: the characterisation and evaluation of opposition to renewable energy projects’, emotion, space and society 2(1), 62–69. https://doi.org/10.1016/j.emospa.2009.05.006 chan, k., satterfield, t. & goldstein, j., 2012, ‘rethinking ecosystem services to better address and navigate cultural values’, ecological economics 74, 8–18. https://doi.org/10.1016/j.ecolecon.2011.11.011 chapin, iii, f.s. & knapp, c.n., 2015, ‘sense of place: a process for identifying and negotiating potentially contested visions of sustainability’, environmental science & policy 53, 38–46. https://doi.org/10.1016/j.envsci.2015.04.012 chen, n., dwyer, l. & firth, t., 2014, ‘conceptualization and measurement of dimensionality of place attachment’, tourism analysis 19, 323–338. https://doi.org/10.3727/108354214x14029467968529 cheng, t., wu, h. & huang, l., 2013, ‘the influence of place attachment on the relationship between destination attractiveness and environmentally responsible behavior for island tourism in penghu, taiwan’, journal of sustainable tourism 21(8), 1166–1187. https://doi.org/10.1080/09669582.2014.965177 chiu, y., lee, w. & chen, t., 2014, ‘environmentally responsible behaviour in ecotourism: antecedents and implications’, tourism management 40, 321–329. https://doi.org/10.1016/j.tourman.2013.06.013 clayton, s. & opotow, s., 2003, ‘introduction: identity and the natural environment’, in s. clayton & s. opotow (eds.), identity and the natural environment: the psychological significance of nature, pp. 1–24, mit press, cambridge, ma. cross, j., 2001, ‘what is “sense of place”?’, paper presented at the twelfth headwaters conference, western state college, november 2–4. devine-wright, p. & clayton, s., 2010, ‘introduction to the special issue: place, identity and environmental behavior’, journal of environmental psychology 30, 267–270. https://doi.org/10.1016/s0272-4944(10)00078-2 dredge, d., 2010, ‘place change and tourism development conflict: evaluating public interest’, tourism management 31(1), 104–112. https://doi.org/10.1016/j.tourman.2009.01.004 ednie, a.j., daigle, j.j. & leahy, j.e., 2010, ‘the development of recreation place attachment on the maine coast: user characteristics and reasons for visiting’, journal of park and recreation administration 28(1), 36–51. eisenhauer, b.w., r.s. krannich & d.j. blahna., 2000, ‘attachment to special places on public lands: an analysis of activities, reasons for attachments, and community connections’, society and natural resources 13(5), 421–441. https://doi.org/10.1080/089419200403848 gosling, e. & williams, k., 2010, ‘connectedness to nature, place attachment and conservation behavior: testing connectedness theory among farmers’, journal of environmental psychology 30(3), 298–304. https://doi.org/10.1016/j.jenvp.2010.01.005 hair, j.f., black, w.c., babin, b.j. & anderson, r.e., 2014, multivariate data analysis, pearson education limited, essex. halpenny, e., 2010, ‘pro-environmental behaviours and park visitors: the effect of place attachment’, journal of environmental psychology 30, 409–421. https://doi.org/10.1016/j.jenvp.2010.04.006 halpenny, e.a., 2006, ‘environmental behaviour, place attachment and park visitation: a case study of visitors to point pele national park’, unpublished doctoral dissertation, university of waterloo, ontario, canada. hammitt, w., backlund, e. & bixler, r., 2006, ‘place bonding for recreational places: conceptual and empirical development’, leisure studies 25(1), 17–41. https://doi.org/10.1080/02614360500098100 harris, b., brown, b. & werner, m., 1996, ‘privacy regulation and place attachment: predicting attachments to a student family housing facility’, journal of environmental psychology 16, 287–301. https://doi.org/10.1006/jevp.1996.0025 hartig, t., book, a., garvill, j., olsson, t. & garling, t., 1996, ‘environmental influences on psychological restoration’, scandinavian journal of psychology 37(4), 378–393. https://doi.org/10.1111/j.1467-9450.1996.tb00670.x hausmann, a., slotow, r., fraser, i. & di minin, e., 2017, ‘ecotourism marketing alternative to charismatic megafauna can also support biodiversity conservation’, animal conservation 20, 91–100. https://doi.org/10.1111/acv.12292 hidalgo, m. & hernandez, b., 2001, ‘place attachment: conceptual and empirical questions’, journal of environmental psychology 21(3), 273–281. https://doi.org/10.1006/jevp.2001.0221 hinds, j. & sparks, p., 2008, ‘engaging with the natural environment: the role of affective connection and identity’, journal of environmental psychology 28, 109–120. https://doi.org/10.1016/j.jenvp.2007.11.001 hwang, s., lee, c. & chen, h., 2005, ‘the relationship among tourists’ involvement, place attachment and interpretation satisfaction in taiwan’s national parks’, tourism management 26(2), 143–156. https://doi.org/10.1016/j.tourman.2003.11.006 jorgensen, b. & stedman, r., 2001, ‘sense of place as an attitude: lakeshore owners’ attitudes toward their properties’, journal of environmental psychology 21, 233–248. https://doi.org/10.1006/jevp.2001.0226 kals, e. & maes, j., 2002, ‘sustainable development and emotions’, in p. schmuck & w. schultz (eds.), psychology of sustainable development, pp. 97–122, kluwer, norwell, ma. kals, e., shumaker, d. & montada, l., 1999, ‘emotional affinity toward nature as a motivational basis to protect nature’, environment and behaviour 31, 178–202. https://doi.org/10.1177/00139169921972056 klenosky, d.b., leblanc, c.l., vogt, c.a. & schroeder, h.w., 2007, ‘factors that attract and repel visitation to urban recreation sites: a framework for research’, paper presented at the 2007 northeastern recreation research symposium. bolton landing, new york. kruger, m., viljoen, a. & saayman, m., 2017, ‘who visits the kruger national park, and why? identifying target markets’, journal of travel & tourism marketing 34(3), 312–340. https://doi.org/10.1080/10548408.2016.1156618 kyle, g., absher, j. & graefe, a., 2003, ‘the moderating role of place attachment on the relationship between attitudes toward fees and spending preference’, leisure sciences 25(1), 33–50. kyle, g., graefe, a. & manning, r., 2004a, ‘attached recreationists … who are they?’, journal of park and recreation administration 22(2), 65–84. kyle, g., graefe, a. & manning, r., 2005, ‘testing the dimensionality of place attachment in recreational settings’, environment and behavior 37(2), 153–177. https://doi.org/10.1177/0013916504269654 kyle, g., graefe, a., manning, r. & bacon, j., 2004b, ‘effects of place attachment on users’ perceptions of social and environmental conditions in a natural setting’, journal of environmental psychology 24(2), 213–225. https://doi.org/10.1016/j.jenvp.2003.12.006 lee, t., 2011, ‘how recreation involvement, place attachment and conservation commitment affect environmentally responsible behavior’, journal of sustainable tourism 19(7), 895–915. https://doi.org/10.1080/09669582.2011.570345 lewicka, m., 2005, ‘ways to make people active: the role of place attachment, cultural capital, and neighbourhood ties’, journal of environmental psychology 25, 381–395. https://doi.org/10.1016/j.jenvp.2005.10.004 lewicka, m., 2008, ‘place attachment, place identity, and place memory: restoring the forgotten city past’, journal of environmental psychology 28, 209–231. https://doi.org/10.1016/j.jenvp.2008.02.001 lewicka, m., 2010, ‘what makes neighborhood different from home and city? effects of place scale on place attachment’, journal of environmental psychology 30, 35–51. https://doi.org/10.1016/j.jenvp.2009.05.004 low, s.m. & altman, i., 1992, ‘place attachment: a conceptual inquiry’, in i. altman & s.m. low (eds.), place attachment, pp. 1–12, plenum press, new york. mcleod, b. & busser, j., 2012, ‘second-home ownership and place attachment: drivers of visitation, word-of-mouth promotion, and hosting’, tourism analysis 17(5), 601–616. https://doi.org/10.3727/108354212x13485873913886 millennium ecosystem assessment (mea), 2005, ecosystems and human well-being: synthesis, island press, washington, dc. moore, r. & graefe, a., 1994, ‘attachments to recreation settings’, leisure sciences 16, 17–31. https://doi.org/10.1080/01490409409513214 moore, r.l. & d. scott., 2003, ‘place attachment and context: comparing a park and a trail within’, forest science 49(6), 877–884. ng, s., kam, p. & pong, r., 2005, ‘people living in ageing buildings: their quality of life and sense of belonging’, journal of environmental psychology 25, 347–360. https://doi.org/10.1016/j.jenvp.2005.08.005 nunnally, j.c., 1978, psychometric theory, mcgraw-hill, new york. perkins, d. & long, a., 2002, ‘neighbourhood sense of community and social capital: a multi-level analysis’, in a. fischer, c. sonn & b. bishop (eds.), psychological sense of community: research, applications and implications, pp. 291–318, plenum press, new york. poira, y., butler, r. & airey, d., 2001, ‘tourism sub-groups: do they exist?’, tourism today 1(1), 14–22. prayag, g. & ryan, d., 2012, ‘antecedents of tourists’ loyalty to mauritius: the role and influence of destination image, place attachment, personal involvement, and satisfaction’, journal of travel research 51(3), 342–356. https://doi.org/10.1177/0047287511410321 pretty, g., chipuer, h. & bramston, p., 2003, ‘sense of place amongst adolescents and adults in two rural australian towns: the discriminating features of place attachment, sense of community and place dependence in relation to place identity’, journal of environmental psychology 23, 273–287. https://doi.org/10.1016/s0272-4944(02)00079-8 prohansky, h., 1978, ‘the city and self-identity’, environment and behaviour 10, 147–169. https://doi.org/10.1177/0013916578102002 ramkissoon, h. & mavondo, f., 2014, ‘proenvironmental behavior: the link between place attachment and place satisfaction’, tourism analysis 19, 673–688. https://doi.org/10.3727/108354214x14146846679286 ramkissoon, h., smith, l. & weiler, b., 2013a, ‘relationships between place attachment, place satisfaction and pro-environmental behaviour in an australian national park’, journal of sustainable tourism 21(3), 434–457. https://doi.org/10.1080/09669582.2012.708042 ramkissoon, h., smith, l. & weiler, b., 2013b, ‘testing the dimensionality of place attachment and its relationships with place satisfaction and pro-environmental behaviours: a structural equation modelling approach’, tourism management 36, 552–566. https://doi.org/10.1016/j.tourman.2012.09.003 ramkissoon, h., weiler, b. & smith, l., 2012, ‘place attachment and pro-environmental behavior in national parks: the development of a conceptual framework’, journal of sustainable tourism 20(2), 257–276. https://doi.org/10.1080/09669582.2011.602194 reimann, m., lamp, m. & palang, h., 2011, ‘tourism impacts and local communities in estonian national parks’, scandinavian journal of hospitality & tourism 11(1), 87–99. https://doi.org/10.1080/15022250.2011.638206 rollero, c. & de piccoli, n., 2010, ‘place attachment, identification and environment perception: an empirical study’, journal of environmental psychology 30, 196–205. https://doi.org/10.1016/j.jenvp.2009.12.003 sanparks, 2019, knp and satara and tamboti customer satisfaction may to july 2017. scannell, l. & gifford, r., 2010, ‘the relations between natural and civic place attachment and pro-environmental behavior’, journal of environmental psychology 30, 289–297. https://doi.org/10.1016/j.jenvp.2010.01.010 schultz, p., 2000, ‘empathizing with nature: the effects of perspective taking on concern for environmental issues’, journal of social issues 56(3), 391–406. https://doi.org/10.1111/0022-4537.00174 stedman, r., 2002, ‘toward a social psychology of place: predicting behaviour from place-based cognitions, attitudes, and identity’, environment and behaviour 34, 561–581. https://doi.org/10.1177/0013916502034005001 stokols, d. & shumacker, s., 1981, ‘people in places: a transactional view of settings’, in j. harvey (ed.), cognition, social behavior, and the environment, pp. 441e448, erlbaum, hillsdale, nj. tonge, j., ryan, m., moore, s. & beckley, l., 2015, ‘the effect of place attachment on pro-environment behavioral intentions of visitors to coastal natural area tourist destinations’, journal of travel research 54(6), 730–743. https://doi.org/10.1177/0047287514533010 tsai, s., 2012, ‘place attachment and tourism marketing: investigating international tourists in singapore’, international journal of tourism research 14(2), 139–152. https://doi.org/10.1002/jtr.842 tuan, y., 1977, space and place: the perspective of experience, university of minnesota press, minneapolis, mn. twigger-ross, c. & uzzell, d., 1996, ‘place and identity processes’, journal of environmental psychology 16(3), 205–220. https://doi.org/10.1006/jevp.1996.0017 vaske, j. & kobrin, k., 2001, ‘place attachment and environmentally responsible behavior’, the journal of environmental education 32(4), 16–21. https://doi.org/10.1080/00958960109598658 vining, j., 1992, ‘environmental emotions and decisions’, environment and behavior 24(1), 3–34. https://doi.org/10.1177/0013916592241001 walker, g. & chapman, r., 2003, ‘thinking like a park: the effects of sense of place, perspective-taking, and empathy on pro-environment intentions’, journal of park and recreation administration 21(4), 71–86. williams, d. & patterson, m., 1999, ‘environmental psychology: mapping landscape meanings for ecosystem management’, in h.k. cordell & j.c. bergstrom (eds.), integrating social sciences and ecosystem management: human dimensions in assessment, policy and management, pp. 141–160, sagamore press, champaign, il. williams, d., patterson, m., roggenbuck, j. & watson, a., 1992, ‘beyond the commodity metaphor: examining emotional and symbolic attachment to place’, leisure sciences 14, 29–46. https://doi.org/10.1080/01490409209513155 williams, d. & stewart, s., 1998, ‘sense of place: an elusive concept that is finding a home in ecosystem management’, journal of forestry 96, 18–23. williams, d. & vaske, j., 2003, ‘the measurement of place attachment: validity and generalizability of a psychometric approach’, forest science 49(6), 830–840. yuksel, a., yuksel, f. & bilim, y., 2010, ‘destination attachment: effects on customer satisfaction and cognitive, affective and connative loyalty’, tourism management 31(2), 274–284. https://doi.org/10.1016/j.tourman.2009.03.007 abstract introduction research methods and design results discussion conclusions acknowledgements references about the author(s) katlego k. mashiane department of geography, faculty of natural and agricultural sciences, university of the free state, qwaqwa, south africa abel ramoelo department of geography, geoinformatics and meteorology, faculty of natural and agricultural sciences, university of pretoria, hatfield, south africa samuel adelebu department of geography, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa ernest daemane department of geography, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa citation mashiane, k.k., ramoelo, a., adelebu, s. & daemane, e., 2023, ‘assessing environmental factors contributing to plant species richness in mountainous mesic grasslands’, koedoe 65(1), a1732. https://doi.org/10.4102/koedoe.v65i1.1732 original research assessing environmental factors contributing to plant species richness in mountainous mesic grasslands katlego k. mashiane, abel ramoelo, samuel adelebu, ernest daemane received: 27 may 2022; accepted: 22 june 2023; published: 04 aug. 2023 copyright: © 2023. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract southern african mountain ranges are characterised by rich and diverse plant species thriving in different habitats with pronounced endemic species. however, globally, biodiversity at the species level is deteriorating rapidly because of environmental change leading to habitat degradation and fragmentation. mountainous grassland communities are particularly vulnerable to rapid ecosystem changes because of their specialised niches and sensitivity to global warming. understanding the determinants of vegetation is necessary for effective and efficient management. this study aimed to determine significant environmental drivers influencing plant species richness in mountainous mesic grasslands. topographical variables (slope and elevation) were derived by using a 30 m resolution digital elevation model. soil variables such as bulk density, silt fragments, ph, coarse fragments, soil organic carbon, sand and nitrogen were acquired from the international soil conference and information centre (isci), and species richness and diversity were derived from vascular plant species inventory data compiled using a field survey. species richness was influenced by soil bulk density, and the interaction between elevation and soil bulk density; higher species richness was associated with lower bulk density and higher elevations. similarly, species composition changed as edaphic factors and elevation changed. conservation implications: species richness increased with soil bulk density, which decreased with elevation. fire severity had little effect on species richness and diversity, which may indicate that management actions do not affect species. however, the impact of grazers still needs to be better understood at this time. moreover, the strong positive relationship between species richness and diversity in this study shows that species richness can be used as a surrogate for diversity and conservation monitoring, especially in mesic highland grasslands. keywords: mountains; subalpine; edaphic; topography; protected grasslands. introduction plant diversity controls processes are pivotal to the functioning of ecosystems (cardinale et al. 2011). however, species-level biodiversity is deteriorating rapidly because of global environmental change drivers such as climate change, pollution, invasive species and land-use changes (hooper et al. 2012). it has been stressed that reduced species richness and concomitant changes in species composition could alter ecosystem services (tilman, reich & isbell 2012). in addition, studies showed that reducing plant diversity in grasslands may lead to lower production of above-ground plant biomass (tilman, wedin & knops 1996). modelling the distribution of mountainous species diversity is essential for identifying priority areas for conservation (barros et al. 2015), mainly because of the sensitivity of these species to climate change (picó & van groenendael 2007; white et al. 2012). for example, subtropical highland grasslands in brazil are home to several endemic groups of plants (iganci et al. 2011), and some taxa are represented by an island-like distribution (lorenz-lemke et al. 2010). furthermore, in mountainous areas of europe, plant diversity is increased by disturbance regimes such as grazing and soil erosion; as a result, the subalpine grasslands are mosaics comprising varying herbaceous communities (armas-herrera et al. 2020). this enhances species composition for grazing while restricting the expansion of unwanted species, that is, shrub encroachment. understanding of biodiversity indicators and factors driving heterogeneity is crucial for conservation interventions and modelling future distributions in response to global change (barros et al. 2015; kuzemko et al. 2016). in addition, the composition of grassland species is controlled by interacting environmental factors such as moisture regime, soil characteristics and other disturbances, including fire. despite interventions such as policy reform and increasing protected areas, most pressures on biodiversity are intensifying (butchart et al. 2010; hooper et al. 2012). as a result, biodiversity loss is accelerating, and thus, this loss of species could be detrimental to the ecosystem and its functions (butchart et al. 2010; meyer et al. 2018; oliver et al. 2015). in addition, richness in grassland species is controlled by parameters that increase heterogeneity such as disturbances, management practices and topography (zulka et al. 2014). spatio-temporal studies of grassland vegetation are required to examine species’ response to heterogeneity and ecosystem processes (lundholm 2009). besides evidence supporting a positive relationship between environmental heterogeneity and species richness, some studies report negative and monotonous relationships (lundholm 2009; yang et al. 2015). establishing heterogeneity-diversity relationships (hdrs) is fundamental for developing management strategies and abating biodiversity loss (filibeck et al. 2019). for example, forage nutrition and spatiotemporal vegetation variations because of geographic rainfall gradients influence the occurrence of large mammals in serengeti national park, tanzania (mcnaughton 1985, 1988). soil catena characteristics control species composition in sub-alpine grassland communities of monte perdido national park in spain (armas-herrera et al. 2020). although potentially susceptible to global environmental change, the vegetation of temperate mountains in southern africa is influenced by ecological, physical and anthropogenic processes (brown & du preez 2020). information and studies on the mountains in southern africa still need to be documented. therefore, by intensifying research and monitoring in this region, information on the vegetation and resilience of this ecosystem would become available (brown & du preez 2020). studies that combine multiple datasets based on ecological traits are encouraged to better understand what determines spatial patterns of biodiversity at explicit scales (barros et al. 2015). grassland ecosystems rank among the most threatened ecosystems in south africa. thus, conservation plans should be put in place to abate the loss of rare and endangered species. in mesic afromontane grasslands of southern africa, vegetation is subjected to intense land-use activities such as grazing by livestock and wildlife (o’connor 2008). the golden gate highlands national park (gghnp) is a protected conservation area experiencing intense grazing by livestock and wild animals, with the potential to change the composition of grass species. the gghnp is highly variable in topography and climate, which results in a complex mosaic of plant communities (kay, bredenkamp & theron 1993). the existing monitoring programmes for grassland vegetation and data ought to be evaluated and analysed for effective conservation policies. the objective of the study was to determine significant environmental drivers influencing plant species richness in the mountainous mesic grasslands of gghnp. research methods and design study area the study was conducted in gghnp, situated in the north-eastern part of the free state province, south africa (figure 1). the park covers 32758.35 ha and lies between 28°27ʹ s – 28°37ʹ s and 28°33ʹ e – 28°42ʹ e. the gghnp is a mountainous grassland located at the foothills of the drakensberg and forms part of the mesic highveld grassland with marked variation in geology and topography. the following soil types were identified in the study area: shallow rocky soils (glenrosa and mispah), deep soil along drainage lines (oakleaf), well-developed sandy soils (hutton and clovelly), as well as clayey structured soils (milkwood and tambakulu) (sanparks 2020). the park has a rich geology divided into stratigraphic units; four sedimentary units (tarkastad subgroup, molteno, elliot and clarens formations) and an igneous drakensberg group that dates 195–210 million years ago. golden gate highlands national park is characterised by summer rainfall, temperate summers and cold winters. the rainfall season stretches from september to april with a mean annual ranging from 1800 mm to 2000 mm (kay et al. 1993). the gghnp lies between 1654 m and 2829 m above sea level (figure 1) and comprises the following grasslands units: eastern free state sandy grasslands (gm 4), basotho montane shrubland (gm 5), lesotho highveld basalt grassland (gd 8) and northern drakensberg highveld (gd5) (mucina & rutherford 2006). figure 1: a map of the golden gate highlands national park study area (the park) with names of sampling sites, plots and elevation (metres above sea level [masl]). sampling design and field data collection the researchers stratified gghnp according to old crop farming fields that are distributed evenly, these sites have quadrats, were sampled in the study area. data were collected in march 2019, april 2019 and may 2019 during the peak growing season. at each quadrat, all herbaceous plant species were identified to species level where possible. the 1 m × 1 m was gridded into 100 cells of 1 cm2 each (representing 1% cover) and summed different land types and elevation (figure 1). golden gate highlands national park was proclaimed in 1962 with additional proclamation of eight farms on the border of qwaqwa in 1989 (sanparks 2020). vegetation sampling was undertaken in 30 m × 30 m plots placed purposively within homogeneous patches of grass communities. each sampling plot was divided into four transects of 30 m. within each 30 m transect, a nested 1 m × 1 m quadrat was sampled at 10 m intervals. in total 142 plots each with 16 quadrats, species cover was averaged per species from each plot. ancillary data acquisition and preparation chemical and physical properties of soil such as soil bulk density (bd), silt fragments (sf), ph, coarse fragments (cfs), soil organic carbon (soc), sand (sd) and nitrogen of topsoil (15 cm) were downloaded from the international soil conference and information centre (https://soilgrids.org); the soil data were validated using ground data from across the globe (hengl et al. 2017). elevation data and shuttle radar topography mission (srtm), digital elevation model (dem) at 30 m resolution were obtained from us geological survey’s eros data centre (https://earthexplorer.usgs.gov). the slope was derived from a dem using the spatial analyst tool in arcgis 10. 4. fire severity data were acquired from a study conducted in the study area (adagbasa, adelabu & okello 2018). the study estimated fire severity using the normalized burn ratio (nbr) index by analysing pre and post fire season (april 2017–september 2017) from remote sensing images with burnt and unburnt pixels. to prepare the data for analysis, the raster images were clipped to the extent of the study area and imported into r to extract pixel values representing environmental variables in each plot using the ‘extract’ function. the taxonomic composition of the vegetation and its cover were used to derive species richness and diversity for each plot. the former was determined by averaging the 1 m2 quadrats to obtain mean plot species richness. total species diversity was calculated using the shannon–wiener index: where pi is the proportion of the species within the sampling unit, these units were averaged to obtain plot level value of species diversity (alpha diversity), which was used as a predictor of species richness. the species richness and diversity were computed using ‘vegan’ and ‘plyr’ found in r statistical software packages (oksanen 2017) by implementing the diversity and apply function for species diversity and richness, respectively. data analysis the function ‘stat. desc’ (in pastecs package) in r was used to determine descriptive statistics of all the variables in this study. exploratory analysis, that is, a correlogram, was used to test for intercorrelations and the relationships between biotic (species richness and diversity), abiotic (bd, sf, sand, soil ph, soc, cf), fire severity, topographic data (elevation and slope) and an interaction of soil and topo-edaphic variables. simple multiple linear regression was used to determine the influence of each environmental variable on species richness. to ensure that multicollinearity was low inflation factors (vifs) were calculated and the variable with the highest vif was removed. this was repeated until the vif of all variables in the final model was at acceptable levels (vif < 5). hierarchical partitioning was used to estimate the contribution of each explanatory variable in the multivariate linear models because the goal was to identify variables with strong direct influence. hierarchical partitioning calculates all possible regression subsets and averages model improvements across all hierarchies, thus permitting a ranking of variables by their independent effects. this analysis was calculated using the r package ‘hier.part’. permutational multivariate analysis of variance (permanova) was used to determine the influence of environmental predictor variables on species composition among different sites in golden gate highlands national park (gghnp). subsequently, environmental variables were fitted onto an ordination of species composition to determine variables that significantly affected species composition across 9 of 13 sites based on the ample species composition data > 5 species per quadrat. these relationships were displayed using non-metric multidimensional scaling (nmds). ethical considerations this article followed all ethical standards for research without direct contact with human or animal subjects. results descriptive statistics the maximum number of species recorded in a single plot was 17 and the lowest was 3. the highest record of species diversity was 2.45 while the lowest was 0.64 in gghnp (table 1). out of the ecological variables, species richness had the highest coefficient of variation (38.9%) compared with fire severity (25.9%) and species diversity (24.7%) (table 1). among all edaphic variables, soc showed the highest coefficient of variation (31.6%). slope (80.4%) had the highest coefficient of variation out of the topographic variables when compared with elevation (5.72%) (table 1). table 1: descriptive statistics including species richness, species diversity, soil variables (list them), topographic variables (list them) and fire severity. relationship between species richness and environmental variables there was a significantly negative relationship between grass species richness and sf, soil organic content and cf; however, there was a positive relationship between species richness and sand. although significant, the other variables had a weak relationship with species richness. species diversity was not significantly related to topographical and soil variables (see figure 2). figure 2: correlogram for 142 herbaceous samples in the golden gate highlands national park. the multiple linear regression showed that species richness was positively and significantly associated with species diversity within the biotic-based model (table 2). however, after removing species diversity, the model with abiotic variables showed soil bd as the only significant variable associated with species richness, albeit negatively. with the topo-edaphic-based model, interaction variables of elevation with bd, cf and sand fragments contributed significantly to variation. only soil bd was positively associated with species richness. table 2: herbaceous species richness versus biotic, abiotic and topo-edaphic variables. hierarchical partitioning showed species diversity as the variable having the highest independent effects followed by edaphic variables (figure 3). the abiotic-based model exhibited silt fragments as the variable with the highest explanation of variation in species richness. the interaction term of elevation and bd was more pronounced as the variable with the strongest direct influence on species richness. figure 3: hierarchical partitioning regression analysis for herbaceous species richness of (a) biotic-based (b) abiotic-based and (c) topo-edaphic-based variables. grass species composition was significantly related to the environmental predictors, that is, bd, sf, sand fragments, ph and elevation. annashope communities were associated with low sand and ph values. maluti and heuwelkop (figure 4) communities separated from the other sites and thrived on soils with high bd. figure 4: non-metric multidimensional scaling ordination of golden gate highlands national park, grass species composition at different sites. discussion in this study, species richness was not significantly related to any of the topographic variables. although a positive hdr was expected, these are common but not universal. a metanalysis conducted by lundholm (2009) found positive hdrs to be mostly lacking. the lack of a relationship may have been because of the low levels of topographic variation across the sites. for instance, the variation in elevation range was very low, this may be a result of our sampling technique of ‘stratified random’, which used old farming lands as a stratum and not elevation. the full mountain range in gghnp has highly variable and unique topography, which results in diverse habitats (kay et al. 1993). therefore, stratification based on elevation maybe important for testing positive hdr of mountainous areas. the variations in mountain topography create micro-environmental conditions that influence habitat heterogeneity and fragmentation (brown & du preez 2020; gaury & devi 2017). consequently, these micro-habitats harbour and shelter a diversity of species serving as essential biodiversity hotspots, providing ecosystem services such as pollination and controlling biological invasion (zulka et al. 2014). studies investigating the influence of elevational gradients on species composition would be a boon for conservation management. a model incorporating species diversity and a set of environmental variables performed well in predicting species richness. in addition, species diversity was strongly related to species richness in spearman’s correlation analysis. the results highlight the importance of incorporating biotic variables in establishing hdrs, which require an understanding of individual species’ response to gradients of spatial environmental heterogeneity factors (lundholm 2009). for example, local patterns in soil moisture seem to be affected by topography through more mechanisms than other topographically controlled factors and with a strong influence on plant diversity. lundholm (2009) suggests that species diversity allows for compartmentalisation and complementary resource use because no single species can fully exploit the conditions of the entire ecosystem, a concept known as niche differentiation (tilman 1999). therefore, species richness is merely the number of species in an ecosystem, while diversity considers the evenness; thus, considering species diversity and richness as independent and dependent variables helps to understand their influence on the gghnp ecosystems’ resilience. soil bd was the only variable negatively affecting species richness within the abiotic-based model and the results of the topo-edaphic-based model showed that elevation and bd jointly affected species richness. this is not surprising because the mountain soil is affected by topography and thus influences local terrestrial plant diversity patterns (moeslund et al. 2013). edaphic features such as soil bd can impair or amplify water loss and spatial accumulation patterns (moeslund et al. 2013). as such, the extent to which portions of a landscape capture or retain water and nutrient resources is an essential determinant of vegetation patterns (moeslund et al. 2013). this emphasises the role of topographically controlled edaphic features on landscapes in gghnp. this could indicate that soil with a lower bd can be vital for plant biodiversity in grassland terrestrial plant species. the results suggest that stable landscapes at high elevations harbour niche species diversity. mid and foot-slope catena are often the recipients of all upslope water and nutrient resources. this causes enrichment that either limits or enhances species richness and diversity, depending on soil quality (mcnaughton 1985). steep slopes are prone to soil erosion, a disturbance could either promote the growth of early-succession species or preclude disturbance-intolerant species in mountainous grassland ecosystems (armas-herrera et al. 2020; filibeck et al. 2019). species composition was significantly related to the environmental predictors of soil variables and elevation. the ordination showed that species composition turned over along sand, ph and bd gradients. this is consistent with the results of armas-herrera et al. (2020), who suggested that grassland vegetation mosaics are maintained by soil–plant feedbacks, especially in subalpine grassland communities where soil disturbances and inevitable succession drive plant co-existence. this finding emphasises the importance of soil drainage in maintaining species diversity in african mountain vegetation (brown & du preez 2020). a species composition shift will likely occur associated with increasing soil resources such as water and nutrients. as defined by particle size, soil texture, is important for controlling soil–plant relationships, especially seedling germination and establishment. soil texture especially, silt and clay content, strongly influences the establishment of many plant species (fair, peters & lauenroth 2001). the risk of species extinction because of global environmental change is imminent, propelling the modelling and identification of conservation hotspots (barros et al. 2015). subtropical highland grasslands have co-evolved with climate and specific microhabitat conditions (iganci et al. 2011). changes in climatic conditions may lead to environmental changes in microhabitats with dire consequences for species diversity, especially in mountainous grasslands, which are sensitive and thus susceptible to environmental changes. the conservation of topographically controlled soils that support and enhance plant diversity can be crucial for maintaining grassland plant communities. in fact, mcnaughton (1985) showed that the resilience for grass species to environmental changes will likely depend on their physiological traits in rangelands. monitoring aspects of biodiversity, that is, richness and diversity as well as species composition are critical for managing ecosystem functions and services for rangelands. these parameters of diversity are crucial for determining carrying capacity and stocking rates to ensure sustainable management of natural resources. conclusions the research corroborates the fact that drivers of species diversity in grassland communities are multidimensional by nature (shoemaker et al. 2020). this is evident by the variations in importance and direction of multiple environmental drivers and their influence on species richness. to understand community dynamics, the incorporation of biotic and abiotic factors is beneficial (shoemaker et al. 2020). the notion that diverse habitats, which are influenced by unique and heterogeneous topography in mountainous biomes, enhance species composition should be investigated. these habitats could be meta-populations fragmented into islands of similar species composition. the research shows that slope and soil interactions are important influencers of species richness in this montane grassland, particularly slope and bd. soil conservation practices such as revegetation and erosion control blankets on bare soil should therefore be a priority, especially in mesic mountainous grasslands where the soil is prone to erosion and nutrient leaching. acknowledgements the authors would like to thank the south african national parks for affording the opportunity to conduct this research. they also acknowledge scientific services intern mpho mbedzi, and the staff at the golden gate highlands national park, especially the rangers who aided immensely in the field data collection. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions all authors conceived and designed the research. k.k.m. conducted the fieldwork, analysed the data and wrote the article. all authors reviewed and edited the manuscript. funding information funding for this research was provided by the university of the free state and sanparks scientific service division. data availability data used in the analysis for the results are available upon request. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references adagbasa, g.e., adelabu, s.a. & okello, t.w., 2018, ‘spatio-temporal assessment of fire severity in a protected and mountainous ecosystem’, igarss 2018 2018 ieee international geoscience and remote sensing symposium, valencia, spain, 2018, pp. 6572–6575. https://doi.org/10.1109/igarss.2018.8518268 armas-herrera, c.m., badía-villas, d., mora, j.l. & gómez, d., 2020, ‘plant-topsoil relationships underlying subalpine grassland patchiness’, science of the total environment 712, 134483. https://doi.org/10.1016/j.scitotenv.2019.134483 barros, m.j.f., silva-arias, g.a., fregonezi, j.n., turchetto-zolet, a.c., iganci, j.r.v., diniz-filho, j.a.f. et al., 2015, ‘environmental drivers of diversity in subtropical highland grasslands’, perspectives in plant ecology, evolution and systematics 17(5), 360–368. https://doi.org/10.1016/j.ppees.2015.08.001 brown, l.r. & du preez, j., 2020, ‘alpine vegetation of temperate mountains of southern africa’, in m.i. goldstein & d.a. dellasala (eds.), encyclopedia of the world’s biomes, pp. 395–404, elsevier, amsterdam. butchart, s.h., walpole, m., collen, b., van strien, a., scharlemann, j.p., almond, r.e. et al., 2010, ‘global biodiversity: indicators of recent declines’, science 328(may), 1164–1169. cardinale, b.j., matulich, k.l., hooper, d.u., byrnes, j.e., duffy, e., gamfeldt, l. et al., 2011, ‘the functional role of producer diversity in ecosystems’, american journal of botany 98(3), 572–592. https://doi.org/10.3732/ajb.1000364 fair, j.l., peters, d.p.c. & lauenroth, w.k., 2001, ‘response of individual bouteloua gracilis (gramineae) plants and tillers to small disturbances’, american midland naturalist 145(1), 147–158. https://doi.org/10.1674/0003-0031(2001)145[0147:roibgg]2.0.co;2 filibeck, g., sperandii, m.g., bazzichetto, m., mancini, l.d., rossini, f. & cancellieri, l., 2019, ‘exploring the drivers of vascular plant richness at very fine spatial scale in sub-mediterranean limestone grasslands (central apennines, italy)’, biodiversity and conservation 28(10), 2701–2725. https://doi.org/10.1007/s10531-019-01788-7 gaury, p.k. & devi, r., 2017, ‘plant species composition and diversity at the aravalli mountain range in haryana, india’, journal of biodiversity 8(1), 34–43. https://doi.org/10.1080/09766901.2017.1336306 hengl, t., mendes de jesus, j., heuvelink, g.b., ruiperez gonzalez, m., kilibarda, m., blagotić, a. et al., 2017, ‘soilgrids250m: global gridded soil information based on machine learning’, plos one 12(2), e0169748. https://doi.org/10.1371/journal.pone.0169748 hooper, d.u., adair, e.c., cardinale, b.j., byrnes, j.e., hungate, b.a., matulich, k.l. et al., 2012, ‘a global synthesis reveals biodiversity loss as a major driver of ecosystem change’, nature 486(7401), 105–108. https://doi.org/10.1038/nature11118 iganci, j.r.v., heiden, g., miotto, s.t.s. & pennington, r.t., 2011, ‘campos de cima da serra: the brazilian subtropical highland grasslands show an unexpected level of plant endemism’, botanical journal of the linnean society 167(4), 378–393. https://doi.org/10.1111/j.1095-8339.2011.01182.x kay, c., bredenkamp, g.j. & theron, g.k., 1993, ‘the plant communities of the golden gate highlands national park in the north-eastern orange free state’, south african journal of botany 59(4), 442–449. https://doi.org/10.1016/s0254-6299(16)30717-7 kuzemko, a.a., steinbauer, m.j., becker, t., didukh, y.p., dolnik, c., jeschke, m. et al., 2016, ‘patterns and drivers of phytodiversity in steppe grasslands of central podolia (ukraine)’, biodiversity and conservation 25(12), 2233–2250. https://doi.org/10.1007/s10531-016-1060-7 lorenz-lemke, a.p., togni, p.d., mäder, g., kriedt, r.a., stehmann, j.r., salzano, f.m. et al., 2010, ‘diversification of plant species in a subtropical region of eastern south american highlands: a phylogeographic perspective on native petunia (solanaceae)’, molecular ecology 19(23), 5240–5251. https://doi.org/10.1111/j.1365-294x.2010.04871.x lundholm, j.t., 2009, ‘plant species diversity and environmental heterogeneity: spatial scale and competing hypotheses’, journal of vegetation science 20(3), 377–391. https://doi.org/10.1111/j.1654-1103.2009.05577.x mcnaughton, s.j., 1985, ‘ecology of a grazing ecosystem: the serengeti’, ecological monographs 55(3), 259–294. https://doi.org/10.2307/1942578 mcnaughton, s.j., 1988, ‘mineral nutrition and spatial concentrations of african ungulates’, nature 334(6180), 343–345. https://doi.org/10.1038/334343a0 mashiane, k., ramoelo, a. & adelabu, s., 2023, ‘diversifying modeling techniques to disentangle the complex patterns of species richness and diversity in the protected afromontane grasslands’, biodiversity and conservation 32, 1423–1436. https://doi.org/10.1007/s10531-023-02560-8 meyer, s.t., ptacnik, r., hillebrand, h., bessler, h., buchmann, n., ebeling, a. et al., 2018, ‘biodiversity-multifunctionality relationships depend on identity and number of measured functions’, nature ecology and evolution 2(1), 44–49. https://doi.org/10.1038/s41559-017-0391-4 moeslund, j.e., arge, l., bøcher, p.k., dalgaard, t. & svenning, j.-c., 2013, ‘topography as a driver of local terrestrial vascular plant diversity patterns’, nordic journal of botany 31(2), 129–144. https://doi.org/10.1111/j.1756-1051.2013.00082.x mucina, l. & rutherford, m.c., 2006, ‘the vegetation of south africa, lesotho and swaziland’, strelitzia 19(december), 1–30, viewed 23 february 2022, from http://ebooks.cambridge.org/ref/id/cbo9781107415324a009. o’connor, t., 2008, ‘influence of land use on phytomass accumulation in highland sourveld grassland in the southern drakensberg, south africa’, african journal of range & forage science 25(1), 17–27. https://doi.org/10.2989/ajrfs.2008.25.1.3.381 oksanen, j., 2017, vegan: ecological diversity, r package version 2.4-4, p. 11, viewed 10 january 2022, from https://cran.r-project.org/package=vegan. oliver, t.h., heard, m.s., isaac, n.j.b., roy, d.b., procter, d., eigenbrod, f. et al., 2015, ‘biodiversity and resilience of ecosystem functions’, trends in ecology & evolution 30(11), 673–684. https://doi.org/10.1016/j.tree.2015.08.009 picó, f.x. & van groenendael, j., 2007, ‘large-scale plant conservation in european semi-natural grasslands: a population genetic perspective’, diversity and distributions 13(6), 920–926. https://doi.org/10.1111/j.1472-4642.2007.00349.x sanparks, 2020, golden gate highlands national park park management plan, south african national parks. shoemaker, l.g., sullivan, l.l., donohue, i., cabral, j.s., williams, r.j., mayfield, m.m. et al., 2020, ‘integrating the underlying structure of stochasticity into community ecology’, ecology 101(2), 1–17. https://doi.org/10.1002/ecy.2922 tilman, d., wedin, d. & knops, j., 1996, ‘productivity and sustainability in grassland ecosystems’, nature, 379(3), 718–720. https://doi.org/10.1038/379718a0 tilman, d., 1999, ‘the ecological consequences of changes in biodiversity: a search for general principles’, ecology 80(5), 1455–1474. https://doi.org/10.1890/0012-9658(1999)080[1455:tecoci]2.0.co;2 tilman, d., reich, p.b. & isbell, f., 2012, ‘biodiversity impacts ecosystem productivity as much as resources, disturbance, or herbivory’, proceedings of the national academy of sciences 109(26), 10394–10397. https://doi.org/10.1073/pnas.1208240109 white, s.r., carlyle, c.n., fraser, l.h. & cahill, jr. j.f., 2012, ‘climate change experiments in temperate grasslands: synthesis and future directions’, biology letters 8(4), 484–487. https://doi.org/10.1098/rsbl.2011.0956 yang, z., liu, x., zhou, m., ai, d., wang, g., wang, y. et al., 2015, ‘the effect of environmental heterogeneity on species richness depends on community position along the environmental gradient’, scientific reports 5(1), 15723. https://doi.org/10.1038/srep15723 zulka, k.p., abensperg-traun, m., milasowszky, n., bieringer, g., gereben-krenn, b.a., holzinger, w. et al., 2014, ‘species richness in dry grassland patches of eastern austria: a multi-taxon study on the role of local, landscape and habitat quality variables’, agriculture, ecosystems and environment 182, 25–36. https://doi.org/10.1016/j.agee.2013.11.016 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 page 14 page 15 page 16 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 abstract introduction methods results discussion conclusion acknowledgements references appendix 1 about the author(s) nabil majdi department of animal ecology, faculty of biology, university of bielefeld, bielefeld, germany lizaan de necker unit for environmental sciences and management, north-west university, potchefstroom, south africa south african institute for aquatic biodiversity, makhanda, south africa hendrika fourie unit for environmental sciences and management, north-west university, potchefstroom, south africa ané loggenberg unit for environmental sciences and management, north-west university, potchefstroom, south africa edward c. netherlands unit for environmental sciences and management, north-west university, potchefstroom, south africa jasmine bunte-tschikin department of animal ecology, faculty of biology, university of bielefeld, bielefeld, germany walter traunspurger department of animal ecology, faculty of biology, university of bielefeld, bielefeld, germany gerhard c. du preez unit for environmental sciences and management, north-west university, potchefstroom, south africa citation majdi, n., de necker, l., fourie, h., loggenberg, a., netherlands, e.c., bunte-tschikin, j. et al., 2022, ‘diversity and distribution of benthic invertebrates dwelling rivers of the kruger national park, south africa’, koedoe 64(1), a1702. https://doi.org/10.4102/koedoe.v64i1.1702 original research diversity and distribution of benthic invertebrates dwelling rivers of the kruger national park, south africa nabil majdi, lizaan de necker, hendrika fourie, ané loggenberg, edward c. netherlands, jasmine bunte-tschikin, walter traunspurger, gerhard c. du preez received: 04 dec. 2021; accepted: 29 apr. 2022; published: 27 june 2022 copyright: © 2022. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract meiobenthos (or meiofauna) are microscopic invertebrates that inhabit biofilms and interstitial spaces in rivers. they are diverse and extremely abundant, and they perform essential ecological functions by linking microbial production to higher trophic levels (e.g. macrobenthic invertebrates and fishes). however, meiobenthic communities remain poorly studied in africa. here, we sampled meioand macrobenthic invertebrate communities associated with biofilms and sediments across an upstream–downstream gradient along the olifants, sabie and crocodile rivers flowing through the kruger national park (knp). we expected to link differences in community structure to environmental gradients as those rivers show different degrees of anthropogenic stress as they enter the park. both meioand macrobenthic communities differed across rivers and also structured along an upstream–downstream gradient. the upstream sites, which were the closest to the park borders, consistently showed a lower diversity in all three rivers. there, the invasive snail tarebia granifera strongly dominated (making up 73% – 87% of the macrobenthos), crowding hard substrates, while concomitantly the abundances of biofilm-dwelling meiobenthos like nematodes and rotifers were substantially reduced. nevertheless, the diversity and evenness of communities then tended to increase as water flowed downstream through the park, suggesting a beneficial effect of protected river reaches on benthic invertebrate diversity. however, for the crocodile river, which makes up the southern border of the park, this trend was less conspicuous, suggesting that this river may experience the greatest threats. more generally, benthic invertebrate communities were driven by the concentrations of phosphates, sulphates, ammonium and organic matter and by substrate characteristics. conservation implications: meiobenthic organisms are very abundant in knp rivers and react to environmental gradients; thus, they should be more considered for bio-monitoring or conservation of comprehensive assemblages of animals. interestingly, protected reaches tended to show a reduced dominance of the invasive t. granifera and a higher diversity of benthic invertebrates. keywords: biodiversity; aquatic ecology; community structure; meiofauna; macrofauna; invasive species; pollution. introduction rivers represent only a minor part of earth’s water resources, but they play a key role for the connectivity of ecosystems and the maintenance of biological diversity at a global scale (dudgeon et al. 2006). however, most river ecosystems have been degraded by human activities throughout history (e.g. fisheries, commercial routes, irrigation, production of energy, receptacles for wastewater and introduction of invasive species), resulting in a current situation where rivers are one of the most threatened ecosystems worldwide (malmqvist & rundle 2002). as an example, south african rivers are intensively exploited to ensure water security for urban areas, agriculture and industry, where water use often exceeds natural water availability (o’keeffe 1989). those rivers further serve as outlets for urban wastewater, excess nutrients and xenobiotics from agriculture, as well as metals draining from mine exploitation or industrial activities (e.g. jackson et al. 2013; rimayi et al. 2018; riddell et al. 2019). nel et al. (2007) analysed the health status of south african rivers and found that 84% of river reaches (and especially those located in the largest, permanently flowing river systems) were degraded to the point they could be considered as highly threatened. nature conservation regions including relatively large portions of river catchments can help mitigate pollution inputs and restore water quality (nel et al. 2007). although some xenobiotics like organochlorine pesticides can still drift over long distances, their bio-accumulation potential in aquatic organisms of conservation areas is correlated with their proximity to pollution sources (wolmarans et al. 2021). this means that river reaches further downstream from pollution sources should show a better ecological status. to reduce threats and develop proactive management practices, one requires an in-depth understanding of the response of natural communities and ecosystem processes to water quality. however, in the case of south african rivers, management of water resources is often decided in a context of incomplete knowledge of the response of the different biological communities present in the field, leading to uncertainty in decision-making (see, e.g., roux, kleynhans & thirion 1999a; roux et al. 1999b). for example, management decisions concerning the conservation actions applicable in the large protected area of the kruger national park (knp) have been traditionally based on the assessment of abiotic rather than biotic factors (solomon et al. 1999). this is based on the assumption that resultant biotic patterns are likely to be correlated with abiotic components. however, to measure, protect and restore natural river integrity in the knp, it would be more relevant to include multiple biological indicators of water quality (rogers & biggs 1999). pollutants not only alter the chemistry of the water column but also persist in sediments (gerber et al. 2015) and within tissues of organisms (gerber et al. 2016; seymore 2014; wolmarans et al. 2021). the consequences of altered environmental conditions have been found to translate into conspicuous modifications of the structure of bacteria, diatom, fish and insect communities in knp water (farrell et al. 2019; rasifudi et al. 2018; riddell et al. 2019; shikwambana et al. 2021). another cause of stress to the native aquatic communities of knp is the accidental introductions followed by problematic blooms of invasive pathogenic micro-organisms, fishes, crayfishes and molluscs species (e.g. jones et al. 2017; macdonald 1988; petersen et al. 2017). while the response of vertebrates to abiotic and biotic threats is conspicuous, the response of benthic invertebrates (living hidden in biofilm matrices and aquatic sediments) is mostly overlooked when it comes to assess ecosystem health status. among those benthic invertebrates, one may further make a distinction between the macrobenthos, such as aquatic insects, snails and leeches that are large enough to be visible and usually are retained on 500 µm meshes, and the meiobenthos, such as nematodes, rotifers, copepods and tardigrades that are so tiny that they are invisible and usually pass through 500 µm-meshes but are retained on 20 µm meshes (ptatscheck, gehner & traunspurger 2020a). meiobenthic invertebrates are little studied although they show complex behaviours and extraordinary physiologies that allow them to colonise most, if not all, benthic habitats (brüchner-hüttemann, ptatscheck & traunspurger 2020; rebecchi, boschetti & nelson 2020) where they can reach remarkable abundances (between 105–106 ind. m−2 on any submerged substrate) (majdi, schmid-araya & traunspurger 2020; traunspurger, wilden & majdi 2020). meiobenthos play an important role in riverine food webs as intermediaries: grazing on microbes (e.g. bacteria, protozoans and micro-algae) and serving as prey for a variety of macro-invertebrates and fish juveniles (majdi & traunspurger 2015; ptatscheck et al. 2020b; schmid-araya et al. 2002). some studies have highlighted the potential of meiobenthic invertebrates to reflect aquatic ecosystem health in south africa: for instance, gyedu-abadio et al. (1999) observed that the density and diversity of nematode assemblages were affected by the concentration of metals in a subtidal portion of the swartkops estuary. further experimentation in the laboratory showed that nematode genera like axonolaimus, theristus and paramonhystera were tolerant to metal pollution (gyedu-ababio & baird 2006). in europe, a nematode ‘species at risk’ index has been developed to monitor river sediment pollution (höss et al. 2011). this index adds value to routinely used macrobenthic-based indices because nematodes are ubiquitous and experience the effects of pollutants during their whole-life cycle within the sediment (brüchner-hüttemann et al. 2021). whether such meiobenthos-based indices could be tested, developed and added to the bio-indication toolbox outside europe is a question of general interest for ecologists and stakeholders, but lacks support from an insufficient collection of field data so far. the specific objective of this study is to provide fresh insights into the diversity and distribution of benthic invertebrates (meioand macrobenthos) in three large rivers (olifants, sabie and crocodile) flowing through an emblematic protected area of austral africa, the knp. we sampled the main benthic habitats in those rivers (i.e. biofilms associated with hard substrates and surface sediment) to examine the structure of resident communities. the three rivers were expected to show a gradient of environmental stress as they enter the park: olifants and crocodile rivers experience different types of pressures (mostly related to mining practices for olifants and to agricultural practices for crocodile), while sabie is quite pristine in comparison. thus, we expected that the structure of benthic invertebrate communities could reflect gradients of pollution experienced by the three rivers. finally, an upstream–downstream gradient was also expected to be manifest through a beneficial effect of preserved areas of the park on the taxonomic richness of benthic assemblages. methods study sites the landscape in the southern part of the knp consists of plains showing a gentle eastward slope and being drained by three major river systems (crocodile, sabie and olifants rivers; figure 1). the crocodile (catchment area: 10 455 km2) and sabie rivers (6252 km2) start off in the drakensberg region. the olifants river has the largest catchment area (54 434 km2) and starts in the highveld region (muller & villet 2004; venter & bristow 1986). it generally shows the highest annual run-off followed by the crocodile and sabie rivers (muller & villet 2004) but, at the time of sampling, all rivers had similar, relatively low discharges (3.82 m3 s−1, 3.87 m3 s−1 and 4.11 m3 s−1 in olifants, crocodile and sabie rivers, respectively). this was the result of a steady decrease in flow over 1 month prior to sampling (figure 1-a1). figure 1: map of the southern part of the kruger national park showing residential and agricultural areas, the different ecozones and river systems with the location of sampling sites. the three rivers drain residential and agricultural areas and then flow eastwards through well-preserved ecozones in the park, except for the crocodile river that delineates the southern border of the park, one of its bank draining the large agricultural zone of the lowveld (figure 1). the crocodile river is considered highly threatened, draining untreated industrial and household wastewater from large cities such as johannesburg and tshwane, its water being increasingly used for irrigation and receiving agricultural effluents. the olifants river drains the industrial fertiliser complex of phalaborwa as well as phosphate rock mining facilities (foskorite and pyroxenite) before entering the park and is considered one of the most polluted rivers in south africa (gerber et al. 2015 and references cited therein). in contrast, the sabie river is recognised as a remarkable hotspot of aquatic biodiversity and one of the most pristine river systems of south africa (riddell et al. 2019). three sites were selected in each river system, with the help of south african national park officials and knp game rangers. each site was chosen according to its accessibility, the availability of riverine microhabitats (hard and soft substrates) and its coherence regarding our aim to examine the effects of an environmental gradient through the park. as far as possible, we selected river reaches that showed the most similar hydro-morphologies and without riparian canopy cover (e.g. figure 2-a1). upstream sites were located close (< 500 m) to the park’s border, and to ensure substantial spatial coverage, each sampling site was located 10 km – 60 km apart from each other (figure 1). sampling samples were collected over a 4-day sampling campaign in may 2019 during austral autumn, a period of low flow conditions (figure 1-a1). to homogenise sampling, at each site, a 25-m reach was selected along which five 1 m2 shallow (< 30 cm water depth) plots located approximately 5 m apart were set (e.g. figure 3-a1). each plot comprised both hard substrates (in the form of large bedrock or cobble stones) and soft substrates (mostly in the form of fine, sandy sediment). to avoid trampling disturbance, plots were always sampled from downstream to upstream (figure 3-a1). samples were placed at 4°c in the dark upon collection and further preserved with fixatives or frozen until further laboratory analyses could take place. at each plot, 500 ml of water was collected for nutrient analyses; and water temperature, ph, conductivity, dissolved inorganic salts, total dissolved solids and turbidity were measured in situ with a portable probe (hi9813-5, hanna instruments inc., bedfordview, south africa). sediments were sampled to measure total organic carbon by pushing one 2-cm diameter perspex corer (figure 1-a1) through 5 cm of the upper sediment layer at one location chosen randomly within each plot. sediments were sampled for meiobenthos by pushing two 2-cm diameter perspex corers through 5 cm of the upper sediment layer at two different locations within each plot. the two sediment cores collected were merged in the same tube and the resulting sample was fixed in a final concentration of 4% buffered formaldehyde. a calibrated brush-sampler (figure 3-a1; peters et al. 2005) was used for underwater sampling of 3.14 cm2 areas of biofilm (and its associated meiobenthos) growing on hard substrates such as rocks and cobbles. this process was repeated three times at different random locations within each plot, using a 20-µm-aperture nylon mesh sieve each time to concentrate the collected subsamples. the contents of the sieve were then poured into a tube and the resulting total sample (representing an area of 9.42 cm2) was fixed in 4% buffered formaldehyde. after meiobenthos sampling, a ‘kick-sampling’ procedure was used to obtain a semi-quantitative sample representative of the macrobenthic community dwelling in each plot. stones, macrophytes and superficial sediments were thoroughly agitated by hand and foot all over the plot for 3 min. the resulting suspended organisms were collected in a sturdy hand net (500 µm meshes) held downstream of the plot. the content of the net was poured in a jar and preserved in 70% ethanol. sample processing water chemistry variables were measured in the laboratory using a merck spectroquant pharo 300 spectrophotometer and appropriate test kits following standard protocols (merck kgaa 2014). variables were measured on defrosted, unfiltered water samples and included cl(further referred to as chloride) (merck protocol #14897), p-po43-(phosphate) (#14848), so42(sulphate) (#14791), n-nh4+ (ammonium) (#14752), n-no3(nitrate) (#14773). subsamples of frozen sediments were defrosted, dried in an oven at 60°c for 96 h, weighted, burned to ashes at 600°c for 6 h, then weighted once again to measure the percentage of total organic content (toc) as a ratio between ash-free dry weight and dry weight of the sediment. to extract meiobenthos from sediment samples, a density-centrifugation procedure was used in the laboratory involving the flotation of organic particles on ludox hs-40 (specific gravity set at 1.14), following the method of pfannkuche and thiel (1988). the supernatant, containing the meiobenthic invertebrates, was poured through 20-µm meshes, preserved in 4% formaldehyde and stained with a few drops of rose bengal. the pellet (i.e. inorganic sediment particles from which the meiobenthos had been extracted) was further passed through a series of stacked sieves (1000 µm, 500 µm, 250 µm, 125 µm, 63 µm and 32-µm-aperture meshes). each sediment size fraction retained on the respective sieve (further coined f1000, f500, f250, f125, f63 and f32) was dried in an oven at 60°c for 96 h and weighted to estimate the sediment particle size distribution of each sample. meiobenthic invertebrates were counted and identified under a 40–80x magnification stereomicroscope (nikon smz1500). morpho-taxonomic features were used to classify the meiobenthic individuals into coarse taxonomic groups: nematodes, rotifers, gastrotrichs, copepods and their nauplii larvae, ostracods, oligochaetes, tardigrades, mites, molluscs (mostly veliger and pediveliger larval stages) and larval stages of plecopters, ephemeropters and dipters (detailing chironomidae and ceratopogonidae families). the weight of sediment from which meiobenthos had been extracted was used to express abundances per gram sediment dry weight. meiobenthic organisms dwelling on hard substrates were not extracted using the ludox procedure and they were directly counted and identified as described above, except that abundances were expressed per area of biofilm. individuals of macrobenthic invertebrates were counted in each sample under a 1–5x magnification dissection microscope (nikon c-leds) and identified to lowest possible taxonomic level based on morphological characteristics and regional identification key for aquatic macro-invertebrates. total abundances were expressed as number of individuals per 1 m2 plot. however, relative abundances were used for community structure analyses to avoid potential misinterpretations because of the semi-quantitative nature of the ‘kick-sampling’ procedure used to sample (table 1-a1). data analysis statistical analyses were performed in r (v 4.0.3.; r development core team 2020) using packages ‘vegan’ and ‘indicspecies’ (de cáceres & legendre 2009). effects of river identity (olifants, crocodile or sabie) and reach location (upstream, midstream or downstream) on abiotic parameters, faunal abundances and macrobenthic diversity indices were tested using a two-way analysis of variance (anova) method followed by a post-hoc tukey’s hsd test for pairwise comparisons. the univariate data were checked for normality and homoscedasticity using shapiro–wilk test and levene’s test, respectively. if data were not normally distributed and homoscedastic, the non-parametric kruskal–wallis rank sum test followed by a multiple comparison test was used instead. for macrobenthos data, effects of river identity and reach were tested on shannon’s diversity index (h’) and pielou’s evenness (j’). multivariate effects of river and reach on the benthic community structure were tested using permutational multivariate analysis of variance using distance matrices (permanova: adonis function with 9999 permutations). the assumption of multivariate homogeneity of variances was first checked using the permdisp2 procedure (betadisper function), which is a multivariate analogous to levene’s test. the function rankindex was also used before tests to rank the performance of different dissimilarity indices. in our case, bray–curtis distance was the most relevant dissimilarity index. non-metric multidimensional scaling (nmds) based on bray–curtis distances (metamds function) was used to ordinate samples and species scores. to highlight the most important environmental factors describing the variability in community structure of macrobenthos and meiobenthos, envfit function (that fits vectors of continuous environmental variables) was used. the significance of environmental variables (vectors) fitted onto the ordination was assessed using a goodness-of-fit (r2) statistics and empirical p-values based on a permutation test (n = 999 permutations). the direction of vector showed the direction of the gradient, and the length of the arrow was proportional to the correlation between the variable score and the ordination space. only significant vectors were further displayed on nmds plot using ‘p.max=0.05’ as a filter argument when plotting vectors on the ordination space. the response of individual taxon abundances to river identity and reach location was examined using multi-level pattern analysis after de cáceres and legendre (2009), which is an extended method of indicator species analysis. multi-level pattern analysis (multipatt function) is a permutation test examining which taxa are significantly responsible for differences among a group of samples. multi-level pattern analysis was chosen over other species indicator methods because it is less sensitive to the weight of over-dominant species, and it is not based on the relative contribution to differences (de cáceres & legendre 2009). ethical considerations this article followed all ethical standards for research without direct contact with human or animal subjects. results water and sediment environment overall, water ph was slightly alkaline but did not differ significantly across rivers or reaches (table 1). however, conductivity was significantly higher in the olifants river (on average 663 µs cm−1) in comparison to crocodile (450 µs cm−1) and sabie (104 µs cm−1) rivers (kruskal–wallis test: χ2 = 38.7, p < 0.001). the concentrations of sulphate and chloride were likely the dominant contributors to variance in conductivity values as strong positive correlations were observed between conductivity and sulphate and chloride concentrations (figure 4-a1). turbidity was significantly lower in sabie in comparison to crocodile and olifants rivers (anova, f2,36 = 31, p < 0.001), and in the latter two more turbid rivers, turbidity was found to be significantly higher at upstream sites (anova, f2,36 = 9.8, p < 0.001). nitrate and ammonium concentrations did not show significant associations with river or with reach, although they were found to be positively correlated with phosphate concentration (figure 4-a1). phosphate was significantly more concentrated in the crocodile river (anova, f2,36 = 37.1, p < 0.001) and interestingly its concentration tended to increase in downstream areas in all rivers (anova, f2,36 = 187.7, p < 0.001). table 1: water and sediment environmental parameters measured at three sites along three rivers flowing through the kruger national park. the sediment of the olifants river showed a significantly greater proportion of organic matter in comparison to the other rivers (kruskal–wallis test: χ2 = 8.2, p = 0.015). in terms of granulometry, relatively coarse sediment dominated (fractions > 500 µm represented on average 63.5% of the sediment grain size distribution). however, it is worth noting that the downstream site on olifants river showed unusually low proportions of the largest grain size categories (table 1). the proportion of coarser sediment was negatively correlated with the proportion of finer grain size categories (figure 4-a1). the coarsest sediment fraction (> 1000 µm) showed a significant river*reach interaction (anova, f4,36 = 6.2, p < 0.001), essentially confirming that sabie’s riverbed showed coarser sediment as we moved downstream. abundance and diversity of benthic invertebrates macrobenthos was at least three orders of magnitude less abundant than the meiobenthos: on average, 130 macrobenthic individuals per m2 against 332 000 ind. m−2 for biofilm-dwelling meiobenthos (table 2). macrobenthos abundance appeared significantly lower in the sabie river (anova, f2,36 = 8.4, p < 0.001). the macrobenthic community consisted of 82 taxa from 53 families or coarser taxonomic groups. fifty-seven taxa could be further resolved to genus or species level (table 1-a1). the taxon dominating the macrobenthic community was tarebia granifera, making up 36.95% of the macrobenthic community on average. however, t. granifera made up 73% – 87% of individuals in upstream reaches (figure 2a). as a result, reach position had a significant effect on the relative abundance of t. granifera (anova, f2,36 = 101, p < 0.001). the larval stage of the ephemeropteran genus caenis sp. was the second most common macrobenthic taxon, representing 16.8% of the assemblage on average and being especially abundant in midand downstream reaches of the sabie and crocodile rivers (figure 2a, table 1-a1). other macrobenthic taxa collectively represented 46.2% of the assemblage on average (figure 2a), with only 14 taxa making up individually more than 1% of the assemblage (tricorythus sp., corbicula fluminalis, paragomphus sp., acanthiops sp., leptophlebiidae, cheumatopsyche sp., notogomphus sp., hydroglyphus sp., laccocoris sp., pseudagrion sp., simulium sp., chironominae anisops sp., elminae larvae; see table 1-a1). on average, 9.2 different macrobenthic taxa were identified from each sample (table 2); however, taxon richness was significantly lower in upstream reaches in comparison to midor downstream reaches (table 2, anova, f2,36 = 12.7, p < 0.001). shannon’s diversity index (h’) was also lower in upstream reaches (table 2) but was further found to be significantly lower in the crocodile river as well (anova, f2,36 = 11, p < 0.001). pielou’s equitability index followed a similar pattern as h’ with communities generally more equitable in midand downstream reaches (anova, f2,36 > 11.2, p < 0.001), except for the downstream reach of the crocodile river, which was uneven and strongly dominated (87%) by caenis sp. (table 2, figure 2a). figure 2: composition of the benthic invertebrate community in three rivers flowing through the kruger national park, south africa. (a) mean (n = 5) relative composition of the macrobenthic community at each sampling site. only the first 51 most abundant taxa out of 82 have assigned coloured legend. taxa abbreviations are detailed in table 1-a1. (b) mean (n = 5) abundance and composition of the meiobenthic community in the sediment. (c) mean (n = 5) abundance and composition of the meiobenthic community in biofilms. examples of sampling site. table 2: abundance, diversity and taxonomic composition of benthic invertebrate communities in three rivers flowing through the kruger national park (detailed taxonomic composition for macrobenthic invertebrates is presented in table 1-a1). meiobenthos abundance in sediment did not show any significant trend (table 2). however, biofilm-dwelling meiobenthic organisms were significantly less abundant in upstream reaches (anova, f2,36 = 9.3, p < 0.001). nematodes and rotifers were the numerically dominant representatives of the meiobenthos in all rivers and reaches (figure 2b and figure 2c), but while nematodes made up on average 21% of the meiobenthic assemblage both in the sediment and in biofilms, rotifers were relatively more abundant in biofilms (45.7% of individuals) than in sediments (31.9%; table 2). structural responses of the macrobenthic community to environmental gradients the structure of the macrobenthic community was significantly affected by river identity (permanova, f2,42 = 29, r2 = 0.12, p = 0.002). multivariate homoscedasticity was met without data transformation (permdisp2, p = 0.72); however, the effects of reach location on macrobenthic community structure could not be tested as multivariate homoscedasticity of variances was not met, even after reasonable data transformation techniques. this was caused by a largely heterogeneous distribution of distances to centroids among reaches: samples from upstream reaches had a very skewed community structure because of the large dominance of t. granifera (figure 2a), resulting in a very low dispersion of sample scores around group centroids (n = 15, average distance to centroid: 0.15 for upstream samples), while for midand downstream sections, average distances of samples to centroids were 0.51 and 0.54, respectively. phosphate was found to be the first driver of macrobenthic community structure (r2 = 0.29, p < 0.001), followed by variables associated with sediment granulometry: f125 (r2 = 0.24, p = 0.002), f1000 (r2 = 0.21, p = 0.009), f250 (r2 = 0.20, p = 0.01) and f63 (r2 = 0.15, p = 0.034). sulphate was also a significant driver (r2 = 0.14, p = 0.036) although its direction was opposed to that of phosphate (figure 3). figure 3: non-metric multidimensional scaling ordination of the macrobenthic community. ellipses show the 50% confidence regions for the locations of group of samples’ centroids. ‘spider webs’ show distances of samples to group centroids they belong. species scores are directly ordinated on the biplot as text abbreviations (in table 1-a1). stars highlight taxa significantly associated with one or two river systems after multi-level pattern analysis. environmental fitting was used to determine significant effects of environmental factors: the direction of vector showed the direction of the gradient, and the length of the arrow was proportional to correlation strength. only significant environmental variables were displayed. other abbreviations include nmds, non-metric multidimensional scaling; po4, concentration of phosphate in the water; so4, concentration of sulphate in the water; f1000, fraction of sediment particles > 1000 µm; f250, fraction of sediment particles of size 250 µm – 500 µm; f125, fraction of sediment particles of size 125 µm – 250 µm; f63, fraction of sediment particles of size 63 µm – 125 µm. the multi-level pattern analysis identified 14 taxa significantly associated with one river group. those groups are highlighted with stars on the nmds biplot (figure 3). namely, only macrobenthic nematodes were found to be significantly associated with the crocodile river. the ephemeropteran larvae tricorythus sp. and elassoneuria sp., the trichopteran larvae cheumatopsyche sp. and macrostemum sp., the dipteran larvae of simulium sp., the dragonfly larvae of pseudagrion sp. and coleopteran adults of berosus sp. were significantly associated with the olifants river. the ephemeropteran larvae of machadorythus maculatus, the hemipteran laccocoris sp., the dipteran larvae of subfamily chironominae, the dragonfly larvae of lestinogomphus sp. and notogomphus sp., and the coleopteran larvae of subfamily larainae were significantly associated with the sabie river. the clam corbicula fluminalis was associated with both crocodile and olifants rivers. dragonfly larvae coenagrionidae and paragomphus sp., and ephemeropteran larvae of the leptophlebiidae family were associated with both crocodile and sabie rivers (figure 3). structural responses of the meiobenthic community to environmental gradients the structure of the meiobenthic community dwelling sediments was significantly affected by river identity (permanova, f2,36 = 2.3, r2 = 0.08, p = 0.003), multivariate homoscedasticity being met without data transformation (permdisp2river, p = 0.21; permdisp2reach, p = 0.4). it was significantly affected by the interaction river*reach (permanova, f4,36 = 2.3, r2 = 0.17, p < 0.001). this was caused by the high nematode and rotifer abundances found in the downstream reach of olifants river (figure 2b). sediment toc was found to be the first driver of sediment-dwelling meiobenthic community structure (r2 = 0.38, p < 0.001; figure 4), followed by variables associated with fine sediment granulometry: f63 (r2 = 0.34, p < 0.001), f32 (r2 = 0.28, p < 0.001). water quality variables such as total dissolved solids (tds) and conductivity (cond) were also significant drivers (r2 = 0.18, p = 0.012, and r2 = 0.17, p = 0.017, respectively). the multi-level pattern analysis identified only one taxon: nematodes, as being significantly associated with the olifants and sabie rivers (figures 2b, figure 4 and table 2). when performed to examine the associations with a specific river reach, the multi-level pattern analysis identified two taxa, namely, the larvae of ceratopogonidae and copepods being associated with both midand downstream reaches (table 2). figure 4: non-metric multidimensional scaling ordination sediment-dwelling meiobenthic community. ordination is based on untransformed absolute abundances (ind. gseddm−1). the structure of the meiobenthic community dwelling biofilms growing on hard substrates was significantly affected by river identity (permanova, f2,36 = 4.9, r2 = 0.15, p < 0.001), river reach (permanova, f2,36 = 5.1, r2 = 0.16, p < 0.001) and by the river*reach interaction (permanova, f4,36 = 1.8, r2 = 0.11, p = 0.023). multivariate homoscedasticity was achieved without data transformation (permdisp2river, p = 0.79; permdisp2reach, p = 0.65). this may have been caused by the relatively coherent pattern of lower abundances in upstream reaches coupled with relatively higher abundances of chironomids in the sabie river (figure 2b and table 2). concentration of ammonium in the water was found to be the strongest driver of biofilm-dwelling meiobenthic community structure (r2 = 0.17, p =0.02; figure 5), followed by tds and conductivity (r2 = 0.145, p = 0.035 and r2 = 0.146, p = 0.032, respectively). the multi-level pattern analysis identified three taxa significantly associated with the sabie river, namely, oligochaetes, tardigrades and gastrotrichs; while one taxon (chironomids) was significantly associated with both olifants and sabie rivers (figure 5). when performed to examine associations with a specific river reach, the multi-level pattern analysis identified four taxa being associated with both midand downstream reaches, namely, ostracods and larval stages of ceratopogonidae, chironomidae and ephemeropterans. figure 5: non-metric multidimensional scaling ordination of the biofilm-dwelling meiobenthic community. ordination is based on untransformed abundances (ind. cm−2). discussion here we studied for the first time the entire assemblage of benthic invertebrates dwelling hard and soft substrates of three major rivers flowing through the knp, south africa. overall, we observed a great diversity of benthic invertebrates of various sizes and morphologies belonging to a total of seven major animal groups: arthropoda, mollusca, nematoda, rotifera, gastrotricha, tardigrada and annelida. we found the highest diversity of families in the sabie river (42 families), followed by the olifants (36 families) and crocodile river (25 families). these results generally confirmed a better biodiversity status for the sabie river as was already observed in previous studies focusing on macrobenthic invertebrates dwelling rivers of the knp. for example, muller and villet (2004) identified 58, 51 and 46 families in the sabie, crocodile and olifants rivers, respectively. the crocodile and olifants rivers are known to experience substantial anthropogenic pressures in comparison to the sabie river (riddell et al. 2019). we found different structures for both meioand macrobenthic communities across the rivers, confirming our hypothesis that benthic communities would react to environmental gradients across rivers. crocodile river’s water had more phosphates and showed the largest alterations of benthic communities, suggesting that this system faced acute stress. olifants river’s water showed more sulphates and a higher conductivity and turbidity, and the sediment texture was finer housing higher amounts of organic material. but as olifants ran through the knp, diversity and evenness tended to improve. sabie showed lower concentrations of nutrients and the most diverse assemblages. however, we found that upstream sites located at the park border were consistently less diverse, supporting that the richness of benthic assemblages benefits from the course of the river through protected areas of the park. the crocodile river, making the southern border of the park, did not show such improvement presumably because it receives effluents from intensive agriculture plots all along its southern catchment outside the knp (van der laan, van antwerpen & bristow 2012). one of the most conspicuous effects of the upstream-downstream gradient was the alteration of community structure because of the strong dominance of t. granifera at all upstream sites. this snail species originates from asia and was first reported from south africa in 1999 in northern kwazulu-natal (appleton & nadasan 2002). it has further rapidly spread into the mpumalanga province, knp and swaziland and will probably continue its expansion to northern sub-tropical lowlands in zimbabwe and mozambique (appleton, forbes & demetriades 2009). fifteen years ago, wolmarans and de kock (2006) sampled t. granifera from the sabie and crocodile rivers, but did not find it in olifants. in our study we found substantial numbers of t. granifera in all river sites, including all river sites along the olifants. t. granifera can attain high densities in invaded areas, thus impacting other benthic fauna and flora (appleton et al. 2009; jones et al. 2017). in invaded estuarine reaches, it has been shown that t. granifera mostly exploits algal biofilms and detritus (miranda, perissinotto & appleton 2011). thus, their grazing activity on hard substrates might particularly affect resource availability for biofilm-dwelling meiobenthos, as was evidenced empirically in the case of other aquatic snails dampening biofilm-associated assemblages (peters, hillebrand & traunspurger 2007). here, we found a significant reduction of biofilm-dwelling meiobenthos in upstream reaches, reduction that could have been caused by a more severe biofilm grazing activity by t. granifera (we observed that ostracods and small instars of insect larvae were rare in the locations strongly dominated by t. granifera). however, stronger evidences are needed to show direct causality so we recommend experimental evaluation of the direct top-down or indirect bottom-up effect of the invasive t. granifera on local biofilm-dwelling meiobenthos. interestingly, our results showed that t. granifera did not over-dominate other downstream reaches in the knp. a possible rationale is that the more diverse communities in midand downstream reaches could have reduced the invasion success of t. granifera through increased competition, predation, or parasitism (miranda et al. 2016). this would also need further experimental evaluation. nevertheless, our results contribute to support that the success of invasive species is all the more increased when indigenous fauna is impoverished because of stress like habitat deterioration (van der velde et al. 2002). the positive association of sulphates with the olifants river is not surprising as this river has historically suffered from salt enrichment because of sulphates that enter the river from effluent discharge which is one of the main contributors to the high salinity measurements in this river (goetsch & palmer 1997; riddell et al. 2019). furthermore, we found a positive association of phosphates in the crocodile river as this system receives a great deal of organic effluents through fertilisers from agricultural activities taking place along its catchment outside the knp (van der laan et al. 2012). similar results were obtained by riddell et al. (2019) who attributed impacts of orthophosphates on rivers in knp to agricultural activities taking place immediately adjacent to the park. in our study, increased sulphate and phosphate levels, along with other organic and inorganic variables emerged as significant drivers of the meioand macrobenthic community structure. biota such as nematodes, the water scavenger beetles berosus spp., the hydropsychid caddisfly cheumatopsyche spp., caenis spp. mayflies, dipteran simulidae and chironominae are adapted to tolerate a wide range of water quality parameters, such as organic enrichment and high salinity (erasmus et al. 2021; malherbe, wepener & van vuren 2010), which is likely why they were associated with the olifants and crocodile rivers. habitat likely also played a role in the distribution of macrobenthos in the rivers as most of the biota specifically associated with either the olifants or the sabie river also have particular habitat and water flow preferences. for instance, the mayflies elassoneuria sp. and tricorythus sp. and the caddisflies macrostemum sp. were significantly associated with the olifants river presumably because of their habitat preference for rocky surfaces in fast-flowing water (gerber & gabriel 2002), a habitat that was more represented at sampling sites in this river. similarly, the sabie river habitats consisted of dense reed vegetation (figure 2-a1), and most of the macrobenthos that were significantly associated with the sabie river have affinities with such dense vegetation patches, particularly laccocoris, or show affinities with the sandy to muddy substrate in slower flowing water at the edges of streams, particularly the gomphids lestinogomphus sp. and notogomphus sp. (gerber & gabriel 2002). although they are omnipresent in limnetic habitats, meiobenthic communities are poorly considered in ecology and conservation studies (majdi et al. 2020). this knowledge gap is even more preoccupying for tropical and subtropical regions because studies about freshwater meiobenthos ecology and taxonomy have traditionally focused on temperate regions from the northern hemisphere (e.g. fontaneto et al. 2012; traunspurger et al. 2020; zullini 2014). here we found an abundant meiobenthic community dominated by nematodes and rotifers, and the taxonomic composition and abundance values reported in the present study are quite comparable to meiobenthic communities found in streams and rivers in europe (e.g. brüchner-hüttemann et al. 2020; majdi et al. 2012; majdi, threis & traunspurger 2017). some meiobenthic groups have been previously investigated in the knp: for example, 30 years ago, botha and heyns (1991, 1992, 1993) identified a total of 33 nematode species (including species new to science) in sediment samples collected from crocodile and olifants rivers. here we also observed an abundant nematode community, particularly in fine sediments of the olifants river. in south african estuaries, pillay and perissinotto (2009) and nozais, perissinotto and tita (2005) have observed that the abundance and community structure of the meiobenthos respond to river inflow and drought conditions (mostly through a decrease in physiologically sensitive taxa under harsher conditions). here we found that biofilmand sediment-dwelling meiobenthos exhibited different responses: nutrient and water quality proxies affected the biofilm-dwelling organisms, while sediment-dwelling meiofauna was also shaped by sediment texture and the availability of benthic organic material. overall, the observed patterns are in agreement with previous studies showing that freshwater meiobenthic communities can become relevant bio-indicators of sediment pollution, water eutrophication or sediment texture (e.g. haegerbaeumer et al. 2017; schenk et al. 2020; traunspurger et al. 2020). however, further studies should detail the response of dominant meiobenthic phyla (e.g. nematodes or rotifers) to the genus or species level in order to identify sensitive and tolerant species and develop subtler bio-indication tools. conclusion the knp is known worldwide for its unique and emblematic terrestrial megafauna but less so for its remarkable freshwater fauna of benthic invertebrates. in this study we observed that the diversity of those invertebrates increased as we moved away from park borders, highlighting a potential beneficial effect of protected river reaches. furthermore, in protected reaches, the assemblage was more even, less affected by the domination of the invasive snail t. granifera. the effects of this species on native assemblages should be further assessed, our results suggesting that t. granifera could affect the structure of biofilm-dwelling communities. other drivers (like the concentration of nutrients in water and sediment granulometry) also showed significant effects on the distribution of benthic invertebrates. we recommend that the interplay between biotic and abiotic drivers should be further studied for a more comprehensive management and conservation of aquatic resources. finally, our results stress that the crocodile river showed the poorest and most unbalanced communities all the way. this suggests that further environment-damaging project on the crocodile river (such as current applications for large-scale coal-mining on the southern boundary of the park) could have catastrophic impacts on an already stressed river system. acknowledgements the authors are grateful to south african national park game guards and officials for providing kind support during sampling and stay at phalaborwa research house and skukuza research camp. they thank robin petersen, mike silberbauer, harold van niekerk, jacques venter, marié du toit and stefanie gehner for their helpful technical support and expertise. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions all authors contributed to the conception and design of the study. material preparation, data collection and analysis were performed by n.m., a.l., e.c.n., j.b.-t. and g.c.d.p. the first draft of the manuscript was written by n.m. and all authors commented on previous versions of the manuscript. the final manuscript has been read and approved by all the authors. funding information the authors declare that no funds, grants or other support were received during the preparation of this manuscript. data availability the data sets generated during the current study are available from the corresponding author upon reasonable request. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. the research complies with ethical standards and sample collection, and the experimental procedures were approved in strict accordance with institutional guidelines. sampling in the park was authorised under strict guidance and after permissions were accorded by sanparks officials. sampling only concerned invertebrates, and reasonable sample sizes were collected. references appleton, c.c., forbes, a.t. & demetriades, n.t., 2009, ‘the occurrence, bionomics and potential impacts of the invasive freshwater snail tarebia granifera (lamarck, 1822) (gastropoda: thiaridae) in south africa’, zoologische mededelingen 83, 525–536. appleton, c.c. & nadasan, d.s., 2002, ‘first report of tarebia granifera (lamarck, 1822) (gastropoda: thiaridae) from africa’, journal of molluscan studies 68(4), 399–402. https://doi.org/10.1093/mollus/68.4.399 botha, a. & heyns, j., 1991, ‘dorylaimoidea (nematoda) from rivers in the kruger national park’, koedoe 34(2), 1–24. https://doi.org/10.4102/koedoe.v34i2.420 botha, a. & heyns, j., 1992, ‘freshwater nematodes of the genera thornenema and mesodorylaimus from the kruger national park with a diagnostic species compendium for south african species of the genus mesodorylaimus (nematoda: dorylaimida)’, koedoe 35(1), 25–42. https://doi.org/10.4102/koedoe.v35i1.387 botha, a. & heyns, j., 1993, ‘species of the genera oxydirus, dorylaimellus (axodorylaimellus), laimydorus and rhabdolaimus from rivers in the kruger national park (nematoda: dorylaimida and araeolaimida)’, koedoe 36(1), 49–60. https://doi.org/10.4102/koedoe.v36i1.361 brüchner-hüttemann, h., höss, s., ptatscheck, c., brinke, m., schenk, j. & traunspurger, w., 2021, ‘added value of the nemaspear [%]-index to routinely used macrofauna-based indices for assessing the quality of freshwater sediments’, ecological indicators 121, 107015. https://doi.org/10.1016/j.ecolind.2020.107015 brüchner-hüttemann, h., ptatscheck, c. & traunspurger, w., 2020, ‘meiofauna in stream habitats: temporal dynamics of abundance, biomass and secondary production in different substrate microhabitats in a first-order stream’, aquatic ecology 54, 1079–1095. https://doi.org/10.1007/s10452-020-09795-5 de cáceres, m. & legendre, p., 2009, ‘associations between species and groups of sites: indices and statistical inference’, ecology 90(12), 3566–3574. https://doi.org/10.1890/08-1823.1 dudgeon, d., arthington, a.h., gessner, m.o., kawabata, z.i., knowler, d.j., lévèque, c. et al., 2006, ‘freshwater biodiversity: importance, threats, status and conservation challenges’, biological reviews 81(2), 163–182. https://doi.org/10.1017/s1464793105006950 erasmus, j.h., lorenz, a.w., zimmermann, s., wepener, v., sures, b., smit, n.j. et al., 2021, ‘a diversity and functional approach to evaluate the macroinvertebrate responses to multiple stressors in a small subtropical austral river’, ecological indicators 131, 108206. https://doi.org/10.1016/j.ecolind.2021.108206 farrell, m.j., govender, d., hajibabaei, m., van der bank, m. & davies, t.j., 2019, ‘bacterial diversity in the waterholes of the kruger national park: an edna metabarcoding approach’, genome 62(3), 229–242. https://doi.org/10.1139/gen-2018-0064 fontaneto, d., barbosa, a.m., segers, h. & pautasso, m., 2012, ‘the ‘rotiferologist’effect and other global correlates of species richness in monogonont rotifers’, ecography 35, 174–182. https://doi.org/10.1111/j.1600-0587.2011.06850.x gerber, a. & gabriel, m.j.m., 2002, aquatic invertebrates of south african rivers field guide, institute for water quality studies department of water affairs and forestry, pretoria, south africa. gerber, r., smit, n.j., van vuren, j.h., nakayama, s.m., yohannes, y.b., ikenaka, y. et al., 2015, ‘application of a sediment quality index for the assessment and monitoring of metals and organochlorines in a premier conservation area’, environmental science and pollution research 22, 19971–19989. https://doi.org/10.1007/s11356-015-5206-z gerber, r., smit, n.j., van vuren, j.h., nakayama, s.m., yohannes, y.b., ikenaka, y. et al., 2016, ‘bioaccumulation and human health risk assessment of ddt and other organochlorine pesticides in an apex aquatic predator from a premier conservation area’, science of the total environment 550, 522–533. https://doi.org/10.1016/j.scitotenv.2016.01.129 goetsch, p.-a. & palmer, c.g., 1997, ‘salinity tolerances of selected macroinvertebrates of the sabie river, kruger national park, south africa’, archives of environmental contamination and toxicology 32, 32–41. https://doi.org/10.1007/s002449900152 gyedu-ababio, t.k. & baird, d., 2006, ‘response of meiofauna and nematode communities to increased levels of contaminants in a laboratory microcosm experiment’, ecotoxicology and environmental safety 63(3), 443–450. https://doi.org/10.1016/j.ecoenv.2005.01.010 gyedu-ababio, t.k., furstenberg, j.p., baird, d. & vanreusel, a., 1999, ‘nematodes as indicators of pollution: a case study from the swartkops river system, south africa’, hydrobiologia 397, 155–169. https://doi.org/10.1023/a:1003617825985 haegerbaeumer, a., höss, s., ristau, k., claus, e., heininger, p. & traunspurger, w., 2017, ‘the use of meiofauna in freshwater sediment assessments: structural and functional responses of meiobenthic communities to metal and organics contamination’, ecological indicators 78, 512–525. https://doi.org/10.1016/j.ecolind.2017.03.048 höss, s., claus, e., von der ohe, p.c., brinke, m., güde, h., heininger, p. et al., 2011, ‘nematode species at risk – a metric to assess pollution in soft sediments of freshwaters’, environment international 37(5), 940–949. https://doi.org/10.1016/j.envint.2011.03.013 jackson, v.a., paulse, a.n., odendaal, j.p. & khan, w., 2013, ‘identification of point sources of metal pollution in the berg river, western cape, south africa’, water, air, & soil pollution 224, 1–8. https://doi.org/10.1007/s11270-013-1477-5 jones, r.w., hill, j.m., coetzee, j.a., avery, t.s., weyl, o.l.f. & hill, m.p., 2017, ‘the abundance of an invasive freshwater snail tarebia granifera (lamarck, 1822) in the nseleni river, south africa’, african journal of aquatic science 42(1), 75–81. https://doi.org/10.2989/16085914.2017.1298984 macdonald, i.a.w., 1988, ‘the history, impacts and control of introduced species in the kruger national park, south africa’, transactions of the royal society of south africa 46(4), 251–276. https://doi.org/10.1080/00359198809520128 majdi, n., mialet, b., boyer, s., tackx, m., leflaive, j., boulêtreau, s. et al., 2012, ‘the relationship between epilithic biofilm stability and its associated meiofauna under two patterns of flood disturbance’, freshwater science 31(1), 38–50. https://doi.org/10.1899/11-073.1 majdi, n., schmid-araya, j.m. & traunspurger, w., 2020, ‘preface: patterns and processes of meiofauna in freshwater ecosystems’, hydrobiologia 847, 2587–2595. https://doi.org/10.1007/s10750-020-04301-2 majdi, n., threis, i. & traunspurger, w., 2017, ‘it’s the little things that count: meiofaunal density and production in the sediment of two headwater streams’, limnology and oceanography 62(1), 151–163. https://doi.org/10.1002/lno.10382 majdi, n. & traunspurger, w., 2015, ‘free-living nematodes in the freshwater food web: a review’, journal of nematology 47, 28–44. malherbe, w., wepener, v. & van vuren, j.h.j., 2010, ‘anthropogenic spatial and temporal changes in the aquatic macro invertebrate assemblages of the lower mvoti river, kwazulu-natal, south africa’, african journal of aquatic science 35(1), 13–20. https://doi.org/10.2989/16085914.2010.466638 malmqvist, b. & rundle, s., 2002, ‘threats to the running water ecosystems of the world’, environmental conservation 29(2), 134–153. https://doi.org/10.1017/s0376892902000097 miranda, n.a., measey, g.j., peer, n., raw, j.l., perissinotto, r. & appleton, c.c., 2016, ‘shell crushing resistance of alien and native thiarid gastropods to predatory crabs in south africa’, aquatic invasions 11, 303–311. https://doi.org/10.3391/ai.2016.11.3.08 miranda, n.a., perissinotto, r. & appleton, c.c., 2011, ‘feeding dynamics of the invasive gastropod tarebia granifera in coastal and estuarine lakes of northern kwazulu-natal, south africa’, estuarine, coastal and shelf science 91(3), 442–449. https://doi.org/10.1016/j.ecss.2010.11.007 muller, w.j. & villet, m.h., 2004, similarities and differences between rivers of the kruger national park, wrc report, water research commission, grahamstown, south africa. nel, j.l., roux, d.j., maree, g., kleynhans, c.j., moolman, j., reyers, b. et al., 2007, ‘rivers in peril inside and outside protected areas: a systematic approach to conservation assessment of river ecosystems’, diversity and distributions 13(3), 341–352. https://doi.org/10.1111/j.1472-4642.2007.00308.x nozais, c., perissinotto, r. & tita, g., 2005, ‘seasonal dynamics of meiofauna in a south african temporarily open/closed estuary (mdloti estuary, indian ocean)’, estuarine, coastal and shelf science 62(1–2), 325–338. https://doi.org/10.1016/j.ecss.2004.09.020 o’keeffe, j.h., 1989, ‘conserving rivers in southern africa’, biological conservation 49(4), 255–274. https://doi.org/10.1016/0006-3207(89)90047-5 peters, l., hillebrand, h. & traunspurger, w., 2007, ‘spatial variation of grazer effects on epilithic meiofauna and algae’, journal of the north american benthological society 26(1), 78–91. https://doi.org/10.1899/0887-3593(2007)26[78:svogeo]2.0.co;2 peters, l., scheifhacken, n., kahlert, m. & rothhaupt, k.o., 2005, ‘note: an efficient in situ method for sampling periphyton in lakes and streams’, archiv für hydrobiologie 163(1), 133–141. https://doi.org/10.1127/0003-9136/2005/0163-0133 petersen, r.m., hoffman, a.c., kotze, p. & marr, s.m., 2017, ‘first record of the invasive australian redclaw crayfish cherax quadricarinatus (von martens, 1868) in the crocodile river, kruger national park, south africa’, koedoe 59(1), 1–3. https://doi.org/10.4102/koedoe.v59i1.1435 pfannkuche, o. & thiel, h., 1988, ‘sample processing’, in r.p. higgins & h. thiel (eds.), introduction to the study of meiofauna, pp. 134–145, smithsonian institution press, washington, dc. pillay, d. & perissinotto, r., 2009, ‘community structure of epibenthic meiofauna in the st. lucia estuarine lake (south africa) during a drought phase’, estuarine, coastal and shelf science 81(1), 94–104. https://doi.org/10.1016/j.ecss.2008.10.014 ptatscheck, c., gehner, s. & traunspurger, w., 2020a, ‘should we redefine meiofaunal organisms? the impact of mesh size on collection of meiofauna with special regard to nematodes’, aquatic ecology 54, 1135–1143. https://doi.org/10.1007/s10452-020-09798-2 ptatscheck, c., brüchner-hüttemann, h., kreuzinger-janik, b., weber, s. & traunspurger, w., 2020b, ‘are meiofauna a standard meal for macroinvertebrates and juvenile fish?’, hydrobiologia 847, 2755–2778. https://doi.org/10.1007/s10750-020-04189-y rasifudi, l., addo-bediako, a., bal, k. & swemmer, t.m., 2018, ‘distribution of benthic macroinvertebrates in the selati river of the olifants river system, south africa’, african entomology 26(2), 398–406. https://doi.org/10.4001/003.026.0398 r development core team, 2020, r: a language and environment for statistical computing, r foundation for statistical computing, vienna, viewed n.d., from http://r-project.org. rebecchi, l., boschetti, c. & nelson, d.r., 2020, ‘extreme-tolerance in meiofaunal organisms: a case study with tardigrades, rotifers and nematodes’, hydrobiologia 847, 2779. https://doi.org/10.1007/s10750-019-04144-6 riddell, e.s., govender, d., botha, j., sithole, h., petersen, r.m. & shikwambana, p., 2019, ‘pollution impacts on the aquatic ecosystems of the kruger national park, south africa’, scientific african 6, e00195. https://doi.org/10.1016/j.sciaf.2019.e00195 rimayi, c., odusanya, d., weiss, j.m., de boer, j. & chimuka, l., 2018, ‘contaminants of emerging concern in the hartbeespoort dam catchment and the umngeni river estuary 2016 pollution incident, south africa’, science of the total environment 627, 1008–1017. https://doi.org/10.1016/j.scitotenv.2018.01.263 rogers, k. & biggs, h., 1999, ‘integrating indicators, endpoints and value systems in strategic management of the rivers of the kruger national park’, freshwater biology 41(2), 439–451. https://doi.org/10.1046/j.1365-2427.1999.00441.x roux, d.j., kleynhans, c.j. & thirion, c., 1999a, ‘biological monitoring and assessment of rivers as a basis for identifying and prioritising river management options’, water science and technology 39(10–11), 207–210. https://doi.org/10.2166/wst.1999.0657 roux, d.j., kleynhans, c.j., thirion, c., hill, l., engelbrecht, j.s., deacon, a.r. et al., 1999b, ‘adaptive assessment and management of riverine ecosystems: the crocodile/elands river case study’, water sa 25, 501–511. schenk, j., höss, s., brinke, m., kleinbölting, n., brüchner-hüttemann, h. & traunspurger, w., 2020, ‘nematodes as bioindicators of polluted sediments using metabarcoding and microscopic taxonomy’, environment international 143, 105922. https://doi.org/10.1016/j.envint.2020.105922 schmid-araya, j.m., hildrew, a.g., robertson, a., schmid, p.e. & winterbottom, j., 2002, ‘the importance of meiofauna in food webs: evidence from an acid stream’, ecology 83(5), 1271–1285. https://doi.org/10.1890/0012-9658(2002)083[1271:tiomif]2.0.co;2 seymore, t., 2014, ‘bioaccumulation of metals in barbus marequensis from the olifants river, kruger national park and lethal levels of manganese to juvenile oreochromis mossambicus’, phd thesis, university of johannesburg. shikwambana, p., taylor, j.c., govender, d. & botha, j., 2021, ‘diatom responses to river water quality in the kruger national park, south africa’, bothalia-african biodiversity & conservation 51(1), 1–11. https://doi.org/10.38201/btha.abc.v51.i1.5 solomon, m., zambatis, n., biggs, h.c. & mare, n., 1999, ‘comparison of classifications commonly used as templates for management, scientific and gis work in the kruger national park’, koedoe 42(2), 131–142. https://doi.org/10.4102/koedoe.v42i2.238 traunspurger, w., wilden, b. & majdi, n., 2020, ‘an overview of meiofaunal and nematode distribution patterns in lake ecosystems differing in their trophic state’, hydrobiologia 847, 2665–2679. https://doi.org/10.1007/s10750-019-04092-1 van der laan, m., van antwerpen, r. & bristow, k.l., 2012, ‘river water quality in the northern sugarcane-producing regions of south africa and implications for irrigation: a scoping study’, water sa 38(1), 87–96. https://doi.org/10.4314/wsa.v38i1.11 van der velde, g., nagelkerken, i., rajagopal, s. & de vaate, a.b., 2002, ‘invasions by alien species in inland freshwater bodies in western europe: the rhine delta’, in e. leppäkoski, s. gollasch & s. olenin (eds.), invasive aquatic species of europe. distribution, impacts and management, pp. 360–372, springer, dordrecht. venter, f.j. & bristow, j.w., 1986, ‘an account of the geomorphology and drainage of the kruger national park’, koedoe 29(1), 117–124. https://doi.org/10.4102/koedoe.v29i1.524 wolmarans, n.j., bervoets, l., gerber, r., yohannes, y.b., nakayama, s.m., ikenaka, y. et al., 2021, ‘bioaccumulation of ddt and other organochlorine pesticides in amphibians from two conservation areas within malaria risk regions of south africa’, chemosphere 274, 129956. https://doi.org/10.1016/j.chemosphere.2021.129956 zullini, a., 2014, ‘is a biogeography of freshwater nematodes possible?’, nematology 16(1), 1–18. https://doi.org/10.1163/15685411-00002779 appendix 1 figure 1-a1: mean daily discharge of olifants, sabie and crocodile rivers before sampling over the period january–may 2019. figure 2-a1: upstream sampling sites and surrounding landscape. figure 3-a1: (a) material used to sample meiobenthos: brush-sampler to scrape stones and cobbles underwater, perspex corer to sample sediment, sieve, funnel and tubes. (b) localisation of sampling plots along a reach of the crocodile river. the downstream plot (#1) was sampled first figure 4-a1: correlation plot showing upper triangular matrix of environmental variables measured in the water and sediment of three south african rivers. correlation coefficients are displayed and clustered. non-significant correlations are not shown. table 1-a1: relative taxonomic composition of the macrobenthic invertebrate community dwelling the riverbed of three rivers flowing through the kruger national park. filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 page 14 page 15 filelist convert a pdf file! abstract introduction methods results discussion appropriate management responses acknowledgements references about the author(s) arne b.r. witt centre for agriculture and biosciences international (cabi), nairobi, kenya winnie nunda centre for agriculture and biosciences international (cabi), nairobi, kenya tim beale centre for agriculture and biosciences international (cabi), oxfordshire, united kingdom darren j. kriticos commonwealth scientific and industrial research organisation (csiro), canberra, australia school of biological science, university of queensland, lucia, australia citation witt, a.b.r., nunda, w., beale, t. & kriticos, d.j., 2020, ‘a preliminary assessment of the presence and distribution of invasive and potentially invasive alien plant species in laikipia county, kenya, a biodiversity hotspot’, koedoe 62(1), a1605. https://doi.org/10.4102/koedoe.v62i1.1605 note: additional supporting information may be found in the online version of this article as online appendix 1 original research a preliminary assessment of the presence and distribution of invasive and potentially invasive alien plant species in laikipia county, kenya, a biodiversity hotspot arne b.r. witt, winnie nunda, tim beale, darren j. kriticos received: 18 oct. 2019; accepted: 23 june 2020; published: 09 sept. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract this is the first assessment of naturalised, invasive and potentially invasive alien plant species present in laikipia county, kenya, which hosts the highest populations of endangered large mammals in the country. we undertook broad-scale roadside surveys in laikipia, recording all naturalised and invasive species, and based on an extensive literature review, also compiled a list of those alien species present that are known to threaten biodiversity and livelihoods elsewhere in the world. the data were supplemented by climex eco-climatic niche models of nine species that we consider to pose the biggest threat to conservation initiatives in the east african region. of the 145 alien plant species recorded, 67 and 37 (including four species of uncertain origin) were considered to be already naturalised or invasive, respectively, and a further 41 species had been recorded as being naturalised or invasive outside of laikipia. most (141) of these species were introduced as ornamentals only or had uses in addition to being ornamentals, with the majority (77) having their origins in tropical america. widespread species in the county included opuntia stricta, o. ficus-indica, austrocylindropuntia subulata and other succulents. based on the current eco-climatic conditions, most of laikipia is unsuitable for chromolaena odorata, marginally suitable for mimosa pigra and lantana camara, and a better climatic match, ranked from least to most favourable, for tithonia diversifolia, cryptostegia grandiflora, parthenium hysterophorus, prosopis juliflora, o. stricta and parkinsonia aculeata. conservation implications: invasive alien plants are known to have negative impacts on biodiversity, and as such pose significant threats to protected area ecosystems worldwide. without efforts to eradicate, contain or control invasive plant species in laikipia, one of the most important conservation areas in eastern africa many rare and iconic wildlife species may be lost. keywords: invasive alien plants; distribution; management; protected areas; alien plant species. introduction invasive alien species are those plants and animals that have been introduced by people, either intentionally or unintentionally, outside of their natural range or outside of their natural dispersal potential, and are destructive to the environment in which they have established and proliferated (unep 2002; witt & luke 2017). invasive alien species (plants and animals) pose a significant threat to biodiversity (pyšek et al. 2012; randall 1996; vilà et al. 2011). for example, a global meta-analysis by vilà et al. (2011) found that invasive plants decrease native plant species diversity and abundance. these plant invasions may have cascading trophic effects (bailey, schweitzer & whitham 2001; sakai et al. 2001; valentine, roberts & schwartzkopf 2007) by decreasing animal fitness and abundance (vilà et al. 2011). this is especially an issue for protected areas where the primary goal is biodiversity conservation (eds. foxcroft et al. 2013; funk & vitousek 2007; hobbs & humphries 1995). de poorter (2007) identified 487 protected areas worldwide in which invasive alien species (plants and animals) were recorded as a threat. allen, brown and stohlgren (2009) reported 20 305 alien plant species invasions in 218 national parks in the united states. invasive plant species have also been reported from protected areas in australia (setterfield et al. 2013), south america (pauchard et al. 2013), europe (pyšek et al. 2013), india (hiremath & sundaram 2013) and elsewhere (see eds. foxcroft et al. 2013). more than 60% of managers in united states national parks indicated that alien plant invasions were of moderate or major concern (randall 2011). goodman (2003) found that invasive plants pose the biggest threat to protected areas in the province of kwazulu-natal, south africa, and protected area managers in europe perceive invasive species as the second greatest threat to biodiversity (pyšek et al. 2013). invasions also impact on communities that are dependent on natural resources for their survival as reported by mwangi and swallow (2008), maundu et al. (2009), kebede and coppock (2015), shackleton et al. (2017a, 2017b, 2017c) and witt, beale and van wilgen (2018). there is therefore a global imperative to manage these species to protect biodiversity and improve livelihoods, especially in mixed-use landscapes, where the main goals are biodiversity conservation and livestock production. most plant species that are now invasive in protected areas were initially intentionally introduced for ornamental purposes, accidentally by tourists or staff, whereas others may have invaded the protected area through natural dispersal from surrounding areas (allen et al. 2009; meyerson & pyšek 2013). tourist facilities, including staff villages, and villages interspersed within conservation areas can be an important source of invasive alien plant species. foxcroft, richardson and wilson (2008) surveyed 36 tourist camps and staff villages in the kruger national park (knp), south africa, and identified 258 alien plant species, several of which had already escaped cultivation and become invasive. in the garden route national park (grnp), also in south africa, baard and kraaij (2014) recorded 244 species of alien plants of which 59% were invasive. witt et al. (2017) recorded 245 alien plant species in the serengeti-mara ecosystem, east africa, of which 212 were intentionally introduced into gardens. of these 212 species, 23 had escaped cultivation, and were recorded as being invasive outside of gardens. the first step in facilitating the management of these invasive alien plants is to gain a better understanding of their presence, distribution and impacts (shackleton et al. 2017a, 2017b, 2017c; witt et al. (2018). here, we report on the naturalised, invasive and potentially invasive alien plant species in laikipia county, kenya, one of the most important multiple-use conservation areas in eastern africa (sundaresan & riginos 2010). despite only 2% of the land in laikipia having been set aside exclusively for wildlife conservation (georgiadis et al. 2007), the county is home to the second highest abundance of wildlife in east africa, after the mara-serengeti ecosystem, and hosts the highest populations of endangered large mammals in kenya, including half of the country’s rhino population, together with significant populations of elephants, grevy’s zebra, reticulated giraffe and wild dogs (sundaresan & riginos 2010). in fact, the county is home to a higher diversity of large mammals than either the serengeti national park in tanzania or knp in south africa (sundaresan & riginos 2010), with the highest diversity of large mammal species of similar size anywhere in the world (butynski & de jong 2014). of the 62 large mammal species present in the county, one is ‘critically endangered’, two are ‘endangered’, four are ‘vulnerable’ and six are ‘near threatened’ (butynski & de jong 2014). moreover, 50% of kenya’s bird species (i.e. more than 560 species) have been recorded in laikipia (butynski & de jong 2014). the only known previous study on the naturalised and invasive plants present in this county was a field guide produced by witt (2017), which did not include any detailed analyses of the data collected. other studies on invasive plants conducted in the area focussed on the invasion of opuntia stricta (haw.) haw. (cactaceae) (strum, stirling & mutunga 2015) or the associated impacts of o. stricta invasion (dudenhoeffer & hodge 2018; dyck 2017). we provide a list of naturalised and/or invasive alien plant species recorded in laikipia and include alien plant species present that are known to be naturalised or invasive elsewhere in the world, but have not been recorded as such in laikipia at the time of the survey. we also provide distribution data, based on roadside surveys, for the most invasive species. we also assess the eco-climatic suitability of laikipia to invasions by some of the worst invasive alien plant species in eastern africa, a few of which are already present in laikipia. this information will be useful in prioritising species for management to protect biodiversity and enhance livelihoods. methods study site approximately 9500 km2 in extent, laikipia county in central kenya is a mix of grasslands, savanna woodland and forest, lying between the aberdares range (4000 m asl) to the south and southwest, mount kenya (5200 m asl) to the east and southeast, eastern (gregory) rift valley (c. 970 m asl) to the west, karisia hills (2580 m asl) to the northwest, mathews range (2688 m asl) to the north and buffalo springs national reserve and samburu national reserve (c. 900 m asl) to the northeast (butynski & de jong 2014; figure 1). figure 1: map showing the location of kenya in africa (inset) and the location of laikipia county within kenya. laikipia experiences a dry and cool climate, which is influenced by the presence of mount kenya and the aberdare mountain range. daily maximum temperatures are around 25 °c, except for the northern part, which is a little warmer, with december and january being the warmest months (lwf 2012). mean annual rainfall increases with elevation, from 400 mm in the northeast to 1000 mm in the southwest on the slopes of mount kenya and the aberdares (lwf 2012). there are two main rainy seasons with the ‘long rains’ falling from march to may, with april being the wettest month, followed by the ‘short rains’ in november. this range of temperatures and rainfall provides habitats for a large number of native and introduced plant and animal species. laikipia is in a transition zone for three major vegetation types: ‘somalia-masai semi-desert grassland and shrubland’, ‘somalia-masai acacia-commiphora bushland and thicket’ and ‘afromontane undifferentiated montane vegetation’ (butynski & de jong 2014). this diversity of vegetation types accounts in part, for the high biological diversity of laikipia. land uses in the county include: (1) mix of ranching (livestock farming) and wildlife conservation, which is the most dominant land use, followed by (2) pastoralism and wildlife, (3) cultivation, (4) pastoralism and cultivation, (5) forests, (6) wildlife ranching and (7) urban settlements (lwf 2012). the county is a unique combination of large-scale ranches that make up about 40% of the landscape, with the remainder consisting mainly of community-owned lands. the large-scale ranches focus mainly on wildlife conservation, tourism and raising beef cattle, while the communities are mainly pastoralists. in fact, more than 80% of people in laikipia are dependent on livestock (butynski & de jong 2014). species surveys we recorded all alien plant species during roadside surveys similar to those undertaken by henderson (2007), rejmánek et al. (2017), shackleton et al. (2017a, 2017b, 2017c), witt and luke (2017) and witt et al. (2018) over 2 years from 2014 to 2015. driving on all accessible roads, including jeep tracks, we recorded the location (using a handheld global positioning system device) and status (present, naturalised and/or invasive), of all alien species that are also known to be naturalised or invasive elsewhere (outside of laikipia county), based on a review of global databases (cabi 2019; issg 2015), and other sources (witt & luke 2017; witt et al. 2018). alien species were recorded as naturalised if they reproduced consistently, and had established self-sustaining populations that had not yet spread widely, whereas invasive species are those that produce large numbers of reproductive offspring that have spread over substantial distances (blackburn et al. 2011; richardson et al. 2000). alien species present in lodge or other gardens, in areas where the main land use was conservation, were surveyed on foot. we only recorded those alien species that are known to be transformers with the potential to have a major impact on the structure and functioning of ecosystems. this information, together with data on the species growth form, origin and uses, was largely gleaned from the same sources described above. we did not record any alien ruderal or agricultural weeds that are not considered to have a significant impact on biodiversity or rangeland productivity. no surveys of alien species were undertaken in towns because it was logistically too complex to survey a large number of gardens when most home owners were not present during the day. a new locality for any particular species was only recorded if it was seen at least 1 km from the previous record. in situations where a species could not be immediately identified, specimens were collected or photographed for later identification by specialists. naturalised and invasive grass species were not recorded, whereas morus, bougainvillea and eucalyptus species were only recorded to genus level because of difficulties in identifying individual species within these genera; they were included in the analysis as ‘species’. vines and many herbaceous plant species are often difficult to observe in the field, especially when not in flower and as such, may have been under-recorded or in some cases not recorded at all. so the absence of a record in a particular area does not mean that the species is not present, just that it was not seen during our surveys. locality data acquired through surveys were entered into a database, and distributions were then mapped at 1/16 degree grid cells (~11 km × 11 km) for the most widespread and abundant invasive alien plant species, based on the number of grid cells in which the species was recorded. if a plant species was found to be present, naturalised and invasive at various localities in the same cell, then the latter took precedence in the species map, indicating that it was found to be invasive in at least one locality within that particular cell. eco-climatic suitability and impacts of selected species there are numerous invasive and potentially invasive plant species already present in laikipia (witt 2017). in addition, there are a number of problematic species that are abundant and widespread outside of laikipia, which could potentially invade the county (witt & luke 2017). we adapted only published eco-climatic suitability models or developed new models for those species currently present in laikipia, or absent yet present in the eastern african region, which pose disproportionate threats to biodiversity and rangeland productivity. these are aggressive invaders that are known to displace valuable forage species, reducing carrying capacities of wildlife and livestock, and ultimately impacting on the welfare of communities. the species of most concern in the eastern african region are prosopis juliflora (sw.) dc (fabaceae), lantana camara l. (verbenaceae), tithonia diversifolia (hemsl.) gray (asteraceae), parthenium hysterophorus l. (asteraceae), o. stricta, chromolaena odorata (l.) r.m. king & h. rob (asteraceae), mimosa pigra l. (fabaceae), parkinsonia aculeata l. (fabaceae) and cryptostegia grandiflora roxb. ex r. br (asclepiadaceae) (witt & luke 2017; witt et al. 2018). although some of these species such as l. camara, t. diversifolia, p. hysterophorus (one grid cell) and o. stricta are already present in laikipia, the others have not been recorded there yet. to estimate whether climatic conditions in laikipia will support further invasions of species already present in the county, and those that are currently absent, yet present in the region, climex eco-climatic models (kriticos & randall 2001) were applied for c. odorata (kriticos et al. 2005), p. hysterophorus (kriticos et al. 2015), l. camara (taylor et al. 2012), m. pigra (walden et al. 2002), p. aculeata (van klinken et al. 2009) or developed de novo for o. stricta (d.j. kriticos unpublished data), p. juliflora (d.j. kriticos unpublished data) and t. diversifolia (j.m. kriticos unpublished data). climex is used to fit eco-climatic niche models to estimate the potential distribution or phenology of organisms based on distribution data for the target organism, and additional information about the response of the organism to weather variables drawn from experiments or phenological observations (kriticos et al. 2015; sutherst & maywald 1985). the resulting models can then be applied to climatic data to explore the climatic suitability of new regions, in this case east africa, and more specifically laikipia county. the distribution data used in the unpublished models were obtained from the global biodiversity information facility (gbif) and witt and luke (2017). specific sources of locality data are described in the relevant model publications (kriticos et al. 2005, 2015; taylor et al. 2012; van klinken et al. 2009; walden et al. 2002). we became aware that the known distribution of c. grandiflora in south africa exceeded its niche as modelled using climex. therefore, the published model of c. grandiflora was modified to fit distribution data from the south africa plant invaders atlas (sapia) database (henderson & wilson 2017), which had been acquired subsequent to the development of the original model developed by kriticos et al. (2003). the only parameter that needed adjustment was the minimum annual heat sum for reproduction (pdd), which was reduced to 1200 °c days, allowing the model results to agree with the distribution data. ethical consideration this article followed all ethical standards for a research without direct contact with human or animal subjects. results species surveys almost 50% of the grid cells in laikipia were surveyed to some extent (figure 2). it was not possible, because of logistic and other reasons, to survey every single garden, even in areas where the predominant land use was conservation. one-hundred and forty-five alien plant species were seen and recorded during our surveys (online appendix 1). this includes calotropis procera (aiton) dryand. (apocynaceae), ipomoea cairica (l.) sweet (convolvulaceae), ricinus communis l. (euphorbiaceae), senna didymobotrya (fresen.) h.s. irwin & barneby (fabaceae) and solanum campylacanthum a. rich (solanaceae), which have an uncertain origin, although considered by some to be native to eastern africa (see witt 2017; witt & luke 2017). they were considered to be naturalised and/or invasive in our analysis. there was also uncertainty as to the identification of vinca major l. (apocynaceae), azolla filiculoides lam. (azollaceae) and crocosmia x crocosmiiflora (lemoine) n.e.br. (iridaceae), but these were nevertheless also included as such in the analysis. this uncertainty occurred because v. major and v. minor l. are morphologically very similar to each other, whereas a. filiculoides could be confused with a. cristata kaulf. (salviniaceae), which is more common in tropical regions, or the native a. pinnata subsp. africana (desv.) saunders and fowler. both argemone mexicana l. and a. ochroleuca sweet were recorded as a single taxon. figure 2: map showing the areas surveyed in laikipia county, kenya (surveyed area shown in 1/16 degree grid squares; ~ 11 km × 11 km). most of the alien species recorded were in the families fabaceae (16 species), asteraceae (12), crassulaceae (11), cactaceae (10) and solanaceae (8). of the 145 alien plant species recorded, 67 were considered to be naturalised in laikipia, although there was uncertainty with regard to the invasion status of cosmos bipinnatus cav. (asterceae), which was included as naturalised for the purposes of this study. most naturalised species recorded belonged to the families cactaceae (nine species), crassulaceae (nine), asteraceae (eight), fabaceae (six) and solanaceae (five). thirty-seven species were regarded as being invasive in laikipia, belonging mainly to the asteraceae (six species), followed by five species in each of fabaceae and solanaceae, and four in each of crassulaceae and cactaceae. the dominant growth forms of all alien species in laikipia considered to be naturalised, invasive or potentially invasive included trees or shrubs (59 species), followed by herbs (31) and climbers (16) (table 1). naturalised species were dominated by herbs (20 species) and trees or shrubs (19), with invasive plants following a similar pattern. most of these alien species were intentionally introduced as ornamentals, although some ornamentals were also used for other purposes (table 2). most of the naturalised and invasive plant species were used, among others, for ornamental, barrier or agricultural purposes. the majority of naturalised species (27) were only used for ornamental purposes, and 14 of the invasive plant species had no other uses other than ornamental. the vast majority of aliens included in this study originated from tropical america (74), followed by species from temperate africa (17) and madagascar (11), and most of those considered to be naturalised and invasive also had a tropical american origin (table 3). table 1: growth forms of alien species seen in laikipia county, kenya, considered to be naturalised, invasive or potentially invasive. table 2: uses of alien species seen in laikipia county, kenya, considered to be naturalised, invasive or potentially invasive. table 3: regions of origin of species seen in laikipia county, kenya, considered to be naturalised, invasive or potentially invasive. distribution opuntia ficus-indica was seen (recorded as present) in 43% of the grid cells surveyed, followed by austrocylindropuntia subulata (41%), datura stramonium l. (solanaceae) (38%), agave sisalana (37%), o. stricta (31%) and agave americana (30%). opuntia ficus-indica was the most widely naturalised species, followed by a. americana, a. subulata, a. sisalana and o. stricta. datura stramonium l. (solanaceae) was the most widespread invasive plant species seen, recorded as such in 28 of the 111 grid cells surveyed, followed by o. stricta (19), o. ficus-indica (18), cirsium vulgare (savi) ten. (asteraceae) (14), a. subulata (11), xanthium strumarium l. (asteraceae) (9), bryophyllum delagoense (eckl. & zeyh.) schinz (crassulaceae) (9), verbena bonariensis l. (verbenaceae) (6) and acacia mearnsii de wild. (fabaceae) (5) (figure 3). datura stramonium was scattered and widespread throughout laikipia, present wherever there was significant disturbance, especially along roadsides, whereas most other invasive plant species had a clumped distribution. although species such as b. delagoense and o. engelmannii (salm-dyck ex engelm. (cactaceae) were not widespread in laikipia, they were invasive in 75% or more of the grid cells in which they were recorded. invasive species such as pistia stratiotes l. (araceae), p. aculeata, p. juliflora, c. grandiflora, m. pigra and passiflora subpeltata ortega (passifloraceae), which are widespread elsewhere in kenya and/or eastern africa, were not seen in laikipia during surveys. figure 3: maps showing the distribution of nine of the most widespread invasive plant species in laikipia county, kenya. eco-climatic modelling the climex eco-climatic niche models indicate that much of laikipia is climatically very suitable for further invasions by o. stricta (figure 4). climatic conditions may also favour the establishment and proliferation of p. aculeata and p. juliflora, and p. hysterophorus is likely to establish and spread throughout most of laikipia. however, we postulate that it is unlikely to be as problematic as in other areas in eastern africa that are more climatically suitable (e.g. south-western and southern kenya). the present climate in laikipia does not appear to be suitable for c. odorata, although l. camara may expand its current distribution but is unlikely to proliferate (figure 4). climatic conditions are suitable for c. grandiflora and t. diversifolia. with no extensive floodplains or swamps, m. pigra is unlikely to invade in lakipia, although plants may establish along some water bodies. figure 4: the eco-climatic suitability of laikipia county, kenya, for nine plant species known to be invasive in eastern africa based on climex eco-climatic niche models. shading depicts the eco-climatic indices (suitability of each location); the darker the shading, the more suitable the climate in that area is for these species to establish and proliferate. discussion the presence of a large number of alien species, mainly ornamentals, a range of climatic regimes (lwf 2012), vegetation types (butynski & de jong 2014) and land-use practices ranging from crop production to conservation (lwf 2012) increase the risk of plant invasions (catford, jansson & nilsson 2009; saunders, hobbs & margules 1991) across the laikipia landscape. data from protected areas in south africa support this assertion (baard & kraaij 2014; eds. foxcroft et al. 2017). the grnp and table mountain national park, both in south africa, are similar in some respects in that they are both fragmented, and nestled among a range of different land-use types, with comparable numbers of alien plant species (baard & kraaij 2014). the grnp consists of approximately 30 detached portions, with farmland, plantations and towns dispersed along its boundaries, making it highly susceptible to invasions (baard & kraaij 2014). of the 244 species of alien plants recorded outside of gardens in the grnp, 23 were casual aliens, 66 were naturalised, 144 were invasive and 12 were transformers (baard & kraaij 2014). these figures are comparable to those of table mountain np (baard & kraaij 2014). the only national park in south africa which has more plant species listed as invasive, based on the national environmental management: biodiversity act (nem:ba) than either the grnp (98 species) or table mountain np (114), is the considerably larger knp with 130 species (baard & kraaij 2014; eds. van wilgen & herbst 2017). however, 96 species recorded as invasive in grnp are not listed by nem:ba as requiring regulation (baard & kraaij 2014), which supports the contention that fragmented conservation areas within mixed-use landscapes may be at higher risk of invasions. there is no similar comparative data for eastern africa, other than that from the serengeti-mara ecosystem, which also consists of multiple land-use types, but has two large contiguous conservation areas, far larger than in laikipia, in the serengeti np and masai-mara national reserve. witt et al. (2017) recorded 245 alien plant species in this ecosystem, of which 212 were intentionally introduced. of these 212 species, 23 were invasive (witt et al. 2017) compared with 67 naturalised and 37 invasive plant species in laikipia. according to spear et al. (2013), high human populations and their associated activities, which may include gardening, in areas surrounding protected areas, may be driving these invasions. plants in cultivation are often the main source of invasions (bucharova & van kleunen 2009; hulme et al. 2008, 2014, 2018; van kleunen, bossdorf & dawson 2018). according to van kleunen et al. (2018), at least 75% and 93% of the globalised naturalised alien flora is grown in domestic and botanical gardens, respectively. the substantial o. stricta invasion in kruger np originated from plants in the staff village in skukuza (foxcroft at al. 2004). we also assume that the o. stricta invasion in laikipia reportedly originated from plantings in the colonial administrators residence in doldol (0°24′00.0″n; 37°10′00.0″e), a small town in the east of laikipia county, although invasions of o. engelmannii on loisaba conservancy (0°21′38.1″n; 36°46′55.3″e) originated from hedge plants that had been discarded in a quarry from where they subsequently spread. many of the bryophyllum and crassula species spreading in laikipia are cultivated in gardens, largely because they are so well adapted to local conditions. the escape and subsequent establishment of cereus jamacaru dc. (cactaceae) on ol jogi conservancy (0°18′54.78″n; 36°58′32.15″e) in laikipia can also be directly linked to plants grown in lodge gardens on the property (witt 2017; witt & luke 2017). austrocylindropuntia subulata has escaped cultivation and has established widely, mainly along water channels (witt 2017; witt & luke 2017). despite being invasive, it is still actively being planted as a hedge, and in some cases, even used on earthen dam walls on conservancies to prevent elephant damage (a.b.r. witt pers. observ.). the threat of naturalised, invasive and potentially invasive succulent species, with the exception of o. stricta and o. engelmannii, is largely being ignored in laikipia, despite their known negative impacts (witt 2017; witt & luke 2017). for example, similar to other invasive cactus species, c. jamacaru can form dense stands, displacing native plants and preventing access to forage by grazers and browsers, resulting in reduced livestockand/or wildlife-carrying capacities. thickets may also impede the movement of livestock and wildlife, and the spines may cause injuries to people and animals (see witt 2017; witt & luke 2017). other potentially invasive cacti, such as cylindropuntia imbricata (haw.) knuth (cactaceae) and opuntia microdasys (lehm.) pfeiff., are also present in the town of doldol. according to issa (2016), the spiny cladodes of c. imbricata adhere to ‘passing animals and the barbed spines can penetrate their skin and feet causing severe injuries’ (n.p.). the succulent herb b. delagoense is another aggressive invader that is expanding its range rapidly in laikipia. it is allelopathic, so it can readily displace grasses and legumes, forming dense monotypic stands (groner 1975). it is also highly toxic (mckenzie & armstrong 1986). in 1997, 125 head of cattle died after eating this species on a travelling stock reserve near moree in new south wales, australia (mckenzie, franke & dunster 1987). no activities are being undertaken to manage any of these invasive and potentially invasive plant species. appropriate management responses alien plant invasions pose significant threats to conservation and livelihoods in laikipia county (shackleton et al. 2017c; witt 2017; witt & luke 2017). as such, it would be prudent to develop and implement management strategies to reduce the threats of all invasive and potentially invasive plant species. to that end, it is imperative that all naturalised, invasive and potentially invasive alien plant species be removed from the grounds of all tourist facilities and possibly also villages that fall within areas where the main land-use practice is livestock production and conservation. those plants that have already escaped cultivation should be eradicated, if possible, or their further spread contained. finally, biological control solutions for widespread and abundant species should be implemented wherever possible, as has been performed for o. stricta and initiated for o. engelmannii (witt et al. 2020). many plant invasions in protected areas have originated from tourism facilities and staff villages (foxcroft & freitag-ronaldson 2007; witt et al. 2017). although attempts to remove these species may be resisted by many residents (foxcroft et al. 2008), this opposition could largely be overcome by implementing a more gradual and nuanced approach. for example, strategies implemented in the kruger np included the removal of high-risk species first, followed by the removal of low-risk species at a later stage, and the clearing of staff gardens whenever a house was vacated (foxcroft at al. 2008). another approach may be to replace alien species with native species, facilitated through the establishment of nurseries focussing on indigenous plantings. actions can also be supported by undertaking weed risk assessments, or similar, which should ideally include eco-climatic maps to determine the climatic suitability of laikipia to invasions by selected species (kriticos, beautrais & dodd 2018). cost–benefit analyses (cbas) should ideally also be undertaken to consider issues around those species that have benefits but are also known to be invasive – the so-called conflict species such as prosopis juliflora (wise, van wilgen & le maitre 2012). if no scientific evidence is available to support these actions, then the precautionary principle (principle 15 of the 1992 rio declaration) which states that ‘where there are threats of serious or irreversible damage, lack of full scientific certainty shall not be used as a reason for postponing cost-effective measures to prevent environmental degradation’ (n.p.) (un 1992) should be invoked. finally, there is also legislation, supporting the removal of invasive and even exotic species from protected areas (see witt et al. 2017). failure to remove invasive or potentially invasive species will merely increase management costs as they escape cultivation and proliferate (eds. wittenberg & cock 2001). once alien species have escaped cultivation and established in natural habitats all efforts should be made to eradicate populations, if possible. this can only be achieved if new incursions are detected early and populations are small and localised (eds. wittenberg & cock 2001). this requires the establishment of surveillance teams or units that are well versed in the identification of alien plant species. resources should also be available at short notice to implement any interventions. cereus jamacaru is currently a good target for local eradication because it has only recently escaped cultivation in laikipia (witt 2017). if this species is not targeted as a matter of urgency, control costs will increase over time as will the impacts on biodiversity. for widespread and abundant species, we strongly advocate the use of biological control, if effective agents are available (day, witt & winston 2020; winston et al. 2014; witt 2017; eds. wittenberg & cock 2001). reviews have indicated that this is a very safe management intervention (see hinz, winston & schwarzländer 2019). ideally, biological control should be integrated with other control practices, wherever possible. biological control is cost-effective, sustainable and environmentally friendly (day & witt 2019; van wilgen & de lange 2011; eds. wittenberg & cock 2001). there are many additional benefits associated with biological control including the fact that agents establish self-perpetuating populations, often across the whole range of the target species (greathead 1995). in addition, most biological control projects only require a one-off investment, and benefits can be reaped by many stakeholders independent of their financial status and irrespective of the fact that if they contributed to the initial research (greathead 1995). the economic returns from biological control projects have also been phenomenal with estimated benefit–cost ratios ranging from 8:1 up to 3726:1 in south africa (van wilgen & de lange 2011). although rarely implemented in kenya, biological control has widely been used at a global level with 1555 separate and intentional releases of 469 species of biological control agents against 175 invasive plant species across 90 countries (winston et al. 2014). there are a number of widespread and abundant invasive plant species in laikipia that could be targeted for biological control (winston et al. 2014). the cochineal dactylopius opuntiae (cockerell) ‘stricta’ biotype (dactylopiidae), recently introduced for the control of o. stricta, is already established in laikipia (witt et al. 2020). species such as o. ficus-indica and o. monacantha have been brought under good control through the introduction of cochineal in the last century (winston et al. 2014). permission is currently being sought from the regulatory authorities to introduce another biotype of d. opuntiae for the control of o. engelmannii. cereus jamacaru has also been brought under good biological control in south africa (zachariades et al. 2017), an option should this species become invasive, although populations are currently such that it can still be eradicated in laikipia. although p. hysterophorus populations are currently localised, biological control agents could also be introduced (strathie, mcconnachie & retief 2011), should the species expand its range in laikipia. additional agents are also available for l. camara (urban et al. 2011), and agents were recently released for the control of t. diversifolia in south africa (simelane, mawela & fourie 2011). a number of agents are also available for other alien plants present in laikipia that could potentially become invasive, provided that they pose no risk to native plants. however, there are a number of targets for which no effective or host-specific agents have been found. for example, despite the sourcing of a number of potential agents for the control of b. delagoense, none are suitably host-specific for release in africa. in this case, concerted efforts will need to be made using conventional means to stop its further spread and reduce the density of current invasions. intervention strategies will need to be developed and implemented for every species based on the control methodologies available locally and internationally. failure to manage invasive alien plants in laikipia will lead to the demise of biodiversity and erode rangeland productivity to the detriment of its people. acknowledgements centre for agriculture and biosciences international (cabi) gratefully acknowledges the core financial support from our member countries (and lead agencies) including the united kingdom (department for international development), china (chinese ministry of agriculture), australia (australian centre for international agricultural research), canada (agriculture and agri-food canada), the netherlands (directorate-general for international cooperation) and switzerland (swiss agency for development and cooperation). see https://www.cabi.org/about-cabi/who-we-work-with/key-donors/ for details. the authors are grateful to the united nations development programme (undp) for providing resources for surveys throughout laikipia county. we also acknowledge the support of ol jogi conservancy and thank lesley henderson for reviewing a previous draft of the manuscript. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions a.b.r.w was responsible for undertaking the field surveys, assisted by w.n. who also entered the field data. t.b. undertook the spatial analysis and compiled the maps, and d.j.k. contributed to the development of the climex niche models. the interpretation of the results and writing of the article was undertaken by a.b.r.w. with inputs from the other authors. funding information this research received no specific grant from any funding agency in the public, commercial, or not-for-profit sectors. data availability statement much of the data has been uploaded to the gbif website witt, a.b.r. & beale, t., 2018, cabi africa invasive and alien species data. cab international. occurrence dataset can be accessed at https://doi.org/10.15468/pkgevu via gbif.org. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references allen, j.a., brown, c.s., stohlgren, t.j., 2009, ‘non-native plant invasions of the united states national parks’, biological invasions 11, 2195–2207. https://doi.org/10.1007/s10530-008-9376-1 baard, j.a. & kraaij, t., 2014, ‘alien flora of the garden route national park, south africa’, south african journal of botany 94, 51–63. https://doi.org/10.1016/j.sajb.2014.05.010 bailey, j., schweitzer, j. & whitham, t., 2001, ‘salt cedar negatively affects biodiversity of aquatic macroinvertebrates’, wetlands 21(3), 442–447. https://doi.org/10.1672/0277-5212(2001)021[0442:scnabo]2.0.co;2 blackburn, t.m., pyšek, p., bacher, s., carlton, j.t., duncan, r.p., jarošík, v. et al., 2011, ‘a proposed unified framework for biological invasions’, trends in ecology and evolution 26, 333–339. https://doi.org/10.1016/j.tree.2011.03.023 bucharova, a. & van kleunen, m., 2009, ‘introduction history and species characteristics partly explain naturalization success of north american woody species in europe’, journal of ecology 97(2), 230–238. https://doi.org/10.1111/j.1365-2745.2008.01469.x butynski, t.m. & de jong, y.a., 2014, ‘primate conservation in the rangeland agroecosystem of laikipia county, central kenya’, primate conservation 28, 117–128. https://doi.org/10.1896/052.028.0104 cabi, 2019, invasive species compendium, cabi international, wallingford, uk, viewed 23 may 2019, from https://www.cabi.org/isc. catford, j.a., jansson, r. & nilsson, c., 2009, ‘reducing redundancy in invasion ecology by integrating hypotheses into a single theoretical framework’, diversity and distributions 15(1), 22–40. https://doi.org/10.1111/j.1472-4642.2008.00521.x day, m.d. & witt, a.b.r., 2019, ‘weed biological control: challenges and opportunities’, journal of asian-pacific weed science society 1, 34–44. day, m.d., witt, a.b.r. & winston, r., 2020, ‘weed biological control in lowand middle-income countries’, current opinion in insect science 38, 92–98. https://doi.org/10.1016/j.cois.2020.02.004 de poorter, m., 2007, invasive alien species and protected areas: a scoping report part. 1. scoping the scale and nature of invasive alien species threats to protected areas impediments to invasive alien species management and means to adress those impediments, global invasive species programme, invasive species specialist group, viewed 01 july 2019, from http://issg.org/gisp_publications_reports.htm. dudenhoeffer, m. & hodge, a-m.c., 2018, ‘opposing forces of seed dispersal and seed predation by mammals for an invasive cactus in central kenya’, african journal of ecology 56(2), 179–184. https://doi.org/10.1111/aje.12504 dyck, m.a., 2017, ‘restoration of native baboon-plant mutualisms following biocontrol of the invasive prickly pear cactus (opuntia stricta) in kenya’, honors theses ay 16/17. 44, viewed 22 july 2018, from http://repository.uwyo.edu/honors_theses_16-17/44. foxcroft, l.c. & downey, p.o., 2008, ‘protecting biodiversity by managing alien plants in national parks: perspectives from south africa and australia’, in b. tokarska-gizic, j. brock, g. brundu, l. child, c. daehler & p. pyšek (eds.), plant invasions: human perception, ecological impacts and management, pp. 387–403, backhuys, leiden. foxcroft, l.c. & freitag-ronaldson, s., 2007, ‘seven decades of institutional learning: managing alien plant invasions in the kruger national park, south africa’, oryx 41(2), 160–167. https://doi.org/10.1017/s0030605307001871 foxcroft, l.c., pyšek, p., richardson, d.m. & genovesi, p. (eds.), 2013, plant invasions in protected areas: patterns, problems and challenges, springer, dordrecht. foxcroft, l.c., richardson, d.m. & wilson, j.r.u., 2008, ‘ornamental plants as invasive aliens: problems and solutions in kruger national park, south africa’, environmental management 41(1), 32–51. https://doi.org/10.1007/s00267-007-9027-9 foxcroft, l.c., rouget, m., richardson, d.m. & mac fadyen, s., 2004, ‘reconstructing 50 years of opuntia stricta invasion in the kruger national park, south africa: environmental determinants and propagule pressure’, diversity and distributions 10(5–6), 427–437. https://doi.org/10.1111/j.1366-9516.2004.00117.x foxcroft, l.c., van wilgen, n.j., baard, j.a. & cole, n.s., 2017, ‘biological invasions in south african national parks’, bothalia 47(2), 1–12. https://doi.org/10.4102/abc.v47i2.2158 funk, j.l. & vitousek, p.m., 2007, ‘resource‑use efficiency and plant invasion in low‑resource systems’, nature 446 (7139), 1079–1081. https://doi.org/10.1038/nature05719 georgiadis, n.j., olwero, j.g.n., ojwang’, g. & romañach, s.s., 2007, ‘savanna herbivore dynamics in a livestock-dominated landscape: i. dependence on land use, rainfall, density, and time’, biological conservation 137(3), 461–472. https://doi.org/10.1016/j.biocon.2007.03.005 goodman, p.s., 2003, ‘assessing management effectiveness and setting priorities in protected areas in kwazulu-natal’, bioscience 53(9), 843–850. https://doi.org/10.1641/0006-3568(2003)053[0843:ameasp]2.0.co;2 greathead, d.j., 1995, ‘benefits and risks of classical biological control’, in h.m.t. hokkanen & j.m. lynch (eds.), biological control, pp. 53–63, cambridge university press, cambridge. groner, m.g., 1975, ‘allelopathic influence of kalanchoe daigremontiana on other species of plants’, botanical gazette 136(2), 207–211. https://doi.org/10.1086/336804 henderson, l. & wilson, j.r.u., 2017, ‘changes in the composition and distribution of alien plants in south africa: an update from the southern african plant invaders atlas (sapia)’, bothalia 47(2), a2172. https://doi.org/10.4102/abc.v47i2.2172 henderson, l., 2007, ‘invasive, naturalised and casual alien plants in southern africa: a summary based on the southern african plant invaders atlas (sapia)’, bothalia 37(2), 215–248. https://doi.org/10.4102/abc.v37i2.322 hinz, h.l., winston, r.l. & schwarzländer, m., 2019, ‘how safe is weed biological control? a global review of direct nontarget attack’, the quarterly review of biology 94(1), 1–27. https://doi.org/10.1086/702340 hiremath, a.j. & sundaram, b., 2013, ‘invasive plant species in indian protected areas: conserving biodiversity in cultural landscapes’ in l.c. foxcroft, p. pyšek, d.m. richardson & p. genovesi (eds.), plant invasions in protected areas: patterns, problems and challenges, pp. 241–266, springer, dordrecht. hobbs, r.j. & humphries, s.e., 1995, ‘an integrated approach to the ecology and management of plant invasions’, conservation biology 9(4), 761–770. https://doi.org/10.1046/j.1523-1739.1995.09040761.x hulme, p.e., bacher, s., kenis, m., klotz, s., kühn, i., minchin, d., et al., 2008, ‘grasping at the routes of biological invasions: a framework for integrating pathways into policy’, journal of applied ecology 45(2), 403–414. https://doi.org/10.1111/j.1365-2664.2007.01442.x hulme, p.e., brundu, g., carboni, m., dehnen-schmutz, k., dullinger, s., early, r., et al., 2018, ‘integrating invasive species policies across ornamental horticulture supply chains to prevent plant invasions’, journal of applied ecology 55(1), 92–98. https://doi.org/10.1111/1365-2664.12953 hulme, p.e., pyšek, p., pergl, j., jarošík, v., schaffner, u. & vilà, m., 2014, ‘greater focus needed on alien plant impacts in protected areas’, conservation letters 7(5), 459–466. https://doi.org/10.1111/conl.12061 issa (invasive species south africa), 2016, factsheet for cylindropuntia imbricata, viewed 01 july 2019, from https://www.invasives.org.za. issg (invasive species specialist group), 2015, the global invasive species database, version 2015.1, viewed 01 july 2019, from https://www.iucngisd.org/gisd/. kebede, a.t. & coppock, d.l., 2015, ‘livestock-mediated dispersal of prosopis juliflora imperils grasslands and the endangered grevy’s zebra in northeastern ethiopia’, rangeland ecology and management 68(5), 402–407. https://doi.org/10.1016/j.rama.2015.07.002 kriticos, d., sutherst, r., brown, j., adkins, s. & maywald, g., 2003, ‘climate change and the potential distribution of an invasive alien plant: acacia nilotica ssp. indica in australia’, journal of applied ecology 40(1), 111–124. https://doi.org/10.1046/j.1365-2664.2003.00777.x kriticos, d.j. & randall, r.p., 2001, ‘a comparison of systems to analyse potential weed distributions’, in r.h. groves, f.d. panetta & j.g. virtue (eds.), weed risk assessment, pp. 61–79, csiro publishing, melbourne. kriticos, d.j., beautrais, j.r. & dodd, m.b., 2018, ‘wrasp: a spatial strategic weed risk analysis tool reveals important subnational variations in weed risks’, weed research 58(6), 398–412. https://doi.org/10.1111/wre.12327 kriticos, d.j., brunel, s., ota, n., fried, g., oude lansink, a.g.j.m., panetta, f.d., et al., 2015, ‘downscaling pest risk analyses: identifying current and future potentially suitable habitats for parthenium hysterophorus with particular reference to europe and north africa’, plos one 10(9), e0132807. https://doi.org/10.1371/journal.pone.0132807 kriticos, d.j., yonow, t. & mcfadyen, r.e., 2005, ‘the potential distribution of chromolaena odorata (siam weed) in relation to climate’, weed research 45(4), 246–254. https://doi.org/10.1111/j.1365-3180.2005.00458.x lwf (laikipia wildlife forum), 2012, a wildlife conservation strategy for laikipia county (2012-2030), 1st edn., laikipia wildlife forum, nanyuki, viewed 21 august 2018, from http://spaceforgiants.org/testsite/wp-content/uploads/2016/02/strat_lwf2012-2030email.pdf. maundu, p., kibet, s., morimoto, y., imbumi, m. & adeka, r., 2009, ‘impact of prosopis juliflora on kenya’s semi-arid and arid ecosystems and local livelihoods’, biodiversity 10(2–3), 33–50. https://doi.org/10.1080/14888386.2009.9712842 mckenzie, r.a. & armstrong, t.r., 1986, ‘poisoning of cattle by bryophyllum plants’, queensland agricultural journal 112(3), 105–108. mckenzie, r.a., franke, f.p. & dunster, p.j., 1987, ‘the toxicity to cattle and bufadienolide content of six species’, australian veterinary journal 66(11), 374–376. https://doi.org/10.1111/j.1751-0813.1989.tb09741.x meyerson, l.a. & pyšek, p., 2013, ‘manipulating alien plant species propagule pressure as a prevention strategy for protected areas’, in l.c. foxcroft, p. pyšek, d.m. richardson & p. genovesi (eds.), plant invasions in protected areas: patterns, problems and challenges, pp. 473–486, springer, dordrecht. mwangi, e. & swallow, b., 2008, ‘prosopis julifora invasion and rural livelihoods in the lake baringo area of kenya’, conservation and society 6(2), 130–140. https://doi.org/10.4103/0972-4923.49207 pauchard, a., fuentes, n., jiménez, a., bustamante, r. & marticorena, a., 2013, ‘alien plants homogenise protected areas: evidence from the landscape and regional scales in south central chile’, in l.c. foxcroft, p. pyšek, d.m. richardson & p. genovesi (eds.), plant invasions in protected areas: patterns, problems and challenges, pp. 191–208, springer, dordrecht. pyšek, p., genovesi, p., pergl, j., monaco, a. & wild, j., 2013, ‘plant invasions of protected areas in europe: an old continent facing new problems’, in l.c. foxcroft, p. pyšek, d.m. richardson & p. genovesi (eds.), plant invasions in protected areas: patterns, problems and challenges, pp. 209–240, springer, dordrecht. pyšek, p., jarošík, v., hulme, p., pergl, j., hejda, m., schaffner, u. et al., 2012, ‘a global assessment of invasive plant impacts on resident species, communities and ecosystems: the interaction of impact measures, invading species’ traits and environment’, global change biology 18(5), 725–1737. https://doi.org/10.1111/j.1365-2486.2011.02636.x randall, j.m., 1996, ‘weed control for the preservation of biological diversity’, weed technology 10(2), 370–383. https://doi.org/10.1017/s0890037x00040124 randall, j.m., 2011, ‘protected areas’, in d. simberloff & m. rejmének (eds.), encyclopaedia of biological invasions, pp. 563–567, university of california press, berkeley, ca. rejmánek, m., huntley, b.j., le roux, j.j. & richardson, d.m., 2017, ‘rapid assessment survey of the invasive plant species in western angola’, african journal of ecology 55(1), 56–69. https://doi.org/10.1111/aje.12315 richardson, d.m., pyšek, p., rejmánek, m., barbour, m.g., dane panetta, f. & west, c.j., 2000, ‘naturalization and invasion of alien plants: concepts and definitions’, diversity and distributions 6(2), 93–107. https://doi.org/10.1046/j.1472-4642.2000.00083.x sakai, a.k., allendorf, f.w., holt, j.s., lodge, d.m., molofsky, j., with, k.a., et al., 2001, ‘the population biology of invasive species’, annual review of ecology and systematics 32, 305–332. https://doi.org/10.1146/annurev.ecolsys.32.081501.114037 saunders, d.a, hobbs, r.j. & margules, c.r., 1991, ‘biological consequences of ecosystem fragmentation: a review’, conservation biology 5(1), 18–32. https://doi.org/10.1111/j.1523-1739.1991.tb00384.x setterfield, s.a., douglas, m.m., petty, a.m., bayliss, p., ferdinands, k.b. & winderlich, s., 2013, ‘invasive plants in the floodplains of australia’s kakadu national park’, in l.c. foxcroft, p. pyšek, d.m. richardson & p. genovesi (eds.), plant invasions in protected areas: patterns, problems and challenges, pp. 167–190, springer, dordrecht. shackleton, r.t., witt, a.b.r., aool, w. & pratt, c.f., 2017a, ‘distribution of the invasive alien weed, lantana camara, and its ecological and livelihood impacts in eastern africa’, african journal of range and forage science 34(1), 1–11. https://doi.org/10.2989/10220119.2017.1301551 shackleton, r.t., witt, a.b.r., nunda, w. & richardson, d.m., 2017b, ‘chromolaena odorata (siam weed) in eastern africa: distribution and socio-ecological impacts’, biological invasions 19, 1285–1298. https://doi.org/10.1007/s20530-016-1338-4 shackleton, r.t., witt, a.b.r., piroris, f.m. & van wilgen, b.w., 2017c, ‘distribution and socio-ecological impacts of the invasive alien cactus opuntia stricta in eastern africa’, biological invasions 19(8), 2427–2441. https://doi.org/10.1007/s10530-017-1453-x simelane, d.o., mawela, k.v. & fourie, a., 2011, ‘prospective agents for the biological control of tithonia rotundifolia (mill.) s.f. blake and tithonia diversifolia (hemsl.) a. gray (asteraceae) in south africa’, african entomology 19(2), 443–450. https://doi.org/10.4001/003.019.0223 spear, d., foxcroft, l.c., bezuidenhout, h. & mcgeoch, m.a., 2013, ‘human population density explains alien species richness in protected areas’, biological conservation 159, 137–147. https://doi.org/10.1016/j.biocon.2012.11.022 strathie, l.w., mcconnachie, a. & retief, e., 2011, ‘initiation of biological control against parthenium hysterophorus l. (asteraceae) in south africa’, african entomology 19(2), 378–392. https://doi.org/10.4001/003.019.0224 strum, s.c., stirling, g. & mutunga, s.k., 2015, ‘the perfect storm: land use change promotes opuntia stricta’s invasion of pastoral rangelands in kenya’, journal of arid environments 118, 37–47. https://doi.org/10.1016/j.jaridenv.2015.02.015 sundaresan, s.r. & riginos, c., 2010, ‘lessons learned from biodiversity conservation in the private lands of laikipia, kenya’, great plains research 20, 17–28. sutherst, r.w. & maywald, g.f., 1985, ‘a computerised system for matching climates in ecology’, agriculture, ecosystems & environment 13(4), 281–299. https://doi.org/10.1016/0167-8809(85)90016-7 taylor, s., kumar, l., reid, n. & kriticos, d.j., 2012, ‘climate change and the potential distribution of an invasive shrub, lantana camara l’, plos one 7(4), e35565. https://doi.org/10.1371/journal.pone.0035565 un (united nations), 1992, rio declaration on environment and development, principle 15, viewed 18 october 2019, from https://www.cbd.int/doc/ref/rio-declaration.shtml. unep (united nations environmental programme), 2002, alien species that threaten ecosystems, habitats or species, unep, the hague. urban, a.j., simelane, d.o., retief, e., heystek, f., williams, h.e. & madire, l.g., 2011, ‘the invasive “lantana camara l.” hybrid complex (verbenaceae): a review of research into its identity and biological control in south africa’, african entomology 19(2), 315–348. https://doi.org/10.4001/003.019.0225 valentine, l., roberts, b. & schwarzkopf, l., 2007, ‘mechanisms driving avoidance of non‑native plants by lizards’, journal of applied ecology 44(1), 228–237. https://doi.org/10.1111/j.1365-2664.2006.01244.x van kleunen, m., bossdorf, o. & dawson, w., 2018, ‘the ecology and evolution of alien plants’, annual review of ecology, evolution, and systematics 49, 25–47. https://doi.org/10.1146/annurev-ecolsys-110617-062654 van kleunen, m., essl, f., pergl, j., brundu, g., carboni, m., dullinger, s. et al., 2018, ‘the changing role of ornamental horticulture in alien plant invasions’, biological reviews 1(3), 0–0. https://doi.org/10.1111/brv.12402 van klinken, r.d., lawson, b.e. & zalucki, m.p., 2009, ‘predicting invasions in australia by a neotropical shrub under climate change: the challenge of novel climates and parameter estimation’, global ecology and biogeography 18(6), 688–700. https://doi.org/10.1111/j.1466-8238.2009.00483.x van wilgen, b.w. & de lange, w.j., 2011, ‘the costs and benefits of biological control of invasive alien plants in south africa’, african entomology 19(2), 504–514. https://doi.org/10.4001/003.019.0228 van wilgen, n.j. & herbst, m. (eds.), 2017, taking stock of parks in a changing world: the sanparks global environmental change assessment, sanparks, cape town. vilà, m., espinar, j.l., hejda, m., hulme, p.e., jarošík, v., maron, j.l., et al., 2011, ‘ecological impacts of invasive alien plants: a meta-analysis of their effects on species, communities and ecosystems’, ecology letters 14(7), 702–708. https://doi.org/10.1111/j.1461-0248.2011.01628.x walden, d., van dam, r., finlayson, m., storrs, m., lowry, j. & kriticos, d.j., 2004, ‘a risk assessment of the tropical wetland weed mimosa pigra in northern australia’, in m. julien, g. flanagan, t. heard, b. hennecke, q. paynter & c. wilson (eds.), research and management of mimosa pigra, pp. 11–21, csiro entomology, canberra. winston, r., schwartzländer, m., hinz, h.l., day, m.d., cock, m.j.w., julien, m.h., 2014, biological control of weeds: a world catalogue of agents and their target weeds, 5th edn., usda forest service, morgantown, wv. wise, r., van wilgen, b. & le maitre, d.c., 2012, ‘costs, benefits and management options for an invasive alien tree species: the case of mesquite in the northern cape, south africa’, journal of arid environments 84, 80–90. https://doi.org/10.1016/j.jaridenv.2012.03.001 witt, a. & luke, q., 2017, guide to the naturalised and invasive plants of eastern africa, cab international, oxfordshire. witt, a., 2017, guide to the naturalised and invasive plants of laikipia, cab international, oxfordshire. witt, a., beale, t. & van wilgen, b.w., 2018, ‘an assessment of the distribution and potential ecological impacts of invasive alien plant species in eastern africa’, transactions of the royal society of south africa 73(3), 217–236. https://doi.org/10.1080/0035919x.2018.1529003 witt, a.b.r., kiambi, s., beale, t. & van wilgen, b.w., 2017, ‘a preliminary assessment of the extent and potential impacts of alien plant invasions in the serengeti-mara ecosystem, east africa’, koedoe 59(1), 1–16. https://doi.org/10.4102/koedoe.v59i1.1426 witt, a.b.r., nunda, w., makale, f., reynolds, k., 2020, ‘a preliminary analysis of the costs and benefits of the biological control agent dactylopius opuntiae on opuntia stricta in laikipia county, kenya’, biocontrol. https://doi.org/10.1007/s10526-020-10018-x wittenberg, r. & cock, m.j.w. (eds.), 2001, invasive alien species: a toolkit of best prevention and management practices, cab international, wallingford. zachariades, c., paterson, i.d., strathie, l.w., hill, m.p. & van wilgen, b.w., 2017, ‘assessing the status of biological control as a management tool for suppression of invasive alien plants in south africa’, bothalia – african biodiversity and conservation 47(2), a2142. https://doi.org/10.4102/abc.v47i2.2142 filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 about the author(s) albertus s. louw department of mathematical sciences, faculty of natural sciences, stellenbosch university, stellenbosch, south africa sandra macfadyen department of mathematical sciences, faculty of natural sciences, stellenbosch university, stellenbosch, south africa sam ferreira conservation services, south african national parks, skukuza, south africa cang hui department of mathematical sciences, faculty of natural sciences, stellenbosch university, stellenbosch, south africa department of theoretical ecology group, african institute for mathematical sciences, cape town, south africa citation louw, a.s., macfadyen, s., ferreira, s. & hui, c., 2021, ‘erratum: elephant population responses to increased density in kruger national park’, koedoe 63(1), a1691. https://doi.org/10.4102/koedoe.v63i1.1691 note: doi of original article published: https://doi.org/10.4102/koedoe.v63i1.1660 correction erratum: elephant population responses to increased density in kruger national park albertus s. louw, sandra macfadyen, sam ferreira, cang hui published: 13 dec. 2021 copyright: © 2021. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. in the published article, louw, a.s., macfadyen, s., ferreira, s. & hui, c., 2021, ‘elephant population responses to increased density in kruger national park’, koedoe 63(1), a1660. https://doi.org/10.4102/koedoe.v63i1.1660, there was a mistake in the legend for figure 1b as published. the colour labels of the legend in figure 1b was replaced with text from the legend of figure 1a by mistake. the original incorrect legend for figure 1b: the revised and updated legend for figure 1b: the publisher apologise for this error. the correction does not change the study’s findings of significance or overall interpretation of the study’s results or the scientific conclusions of the article in any way. brown.qxd sanparks plant communities of the ebenhaezer section of the mountain zebra national park j. de klerk, l.r. brown and h. bezuidenhout de klerk, j., l.r. brown and h. bezuidenhout. 2003. plant communities of the ebenhaezer section of the mountain zebra national park. koedoe 46(2): 9-20. pretoria. issn 0075-6458. the long-term conservation of viable ecosystems requires a broader understanding of the ecological processes involved. because ecosystems react differently to different management practices, it is important to have a description and classification of the vegetation of an area available. as part of a vegetation survey programme for the newly acquired farms to be incorporated into the mountain zebra national park, the vegetation of the ebenhaezer section was investigated. a hierarchical classification, vegetation map, description and ecological interpretation of the plant communities of the study area are presented. a twinspan classification, refined by braun-blanquet procedures revealed eight distinct plant communities. habitat factors associated with differences in vegetation include topography, soil form and grazing. descriptions of the plant communities include diagnostic species as well as prominent and less conspicuous species of the tree, shrub and herbaceous layers. key words: braun-blanquet procedures, conservation area, plant communities, twinspan, vegetation classification. j. de klerk and l.r. brown, applied natural sciences, technikon sa, private bag x6, florida, 1710 republic of south africa; h. bezuidenhout, conservation services, south african national parks, p.o. box 110040, hadison park, kimberley, 8306 republic of south africa. issn 0075-6458 9 koedoe 46/2 (2003) introduction the mountain zebra national park (mznp), situated in the eastern cape province of south africa, was proclaimed in 1937 (wahl & naude 1996) to preserve the last mountain zebra (equus zebra zebra l., 1758) population in the bankberg area of the park. the first proclaimed section was only 1 712 ha in size and too small to carry a viable mountain zebra population. in order to accommodate a more viable mountain zebra population, it was decided during the 1960s to enlarge the park. neighbouring farms were purchased and the park was enlarged to 6 536 ha. a detailed phytosociological study on the vegetation of the enlarged park was conducted by van der walt in 1980. in the early 1990s, a further long-term project to enlarge the park was initiated. from 1996, various farms adjacent to the mznp were acquired and incorporated into the park. one of the farms, ebenhaezer, also known as wendover, became part of the greater mountain zebra national park during the 1999/2000 period. vegetation classification and mapping forms an integral part of ecological planning in a conservation area (edwards 1972). according to harper & hawksworth (1995) biological inventories are needed if full quantitative measures of biodiversity are to be used in making conservation decisions. certain parts of mznp have already been studied in this respect. brown & bezuidenhout (2000) have described fourteen plant communities in the de rust area. a study by pond et al. (2002) indicated that the species richness (680 plant species) in mznp is considerably higher when compared to other parks and reserves in the more arid regions of south africa. brown.qxd 2005/12/09 10:36 page 9 sanparks descriptions of the plant communities (inventories) on the new farms incorporated into mznp are therefore necessary study area the ebenhaezer section of mountain zebra national park is situated approximately 18 km west of the town of cradock in the eastern cape (fig. 1). the study area comprises 1 800 ha and lies directly north of the previously proclaimed section of mznp, where it borders onto the rooiplaat plateau. it extends from latitude 32º09'–32º14's and longitude 25º25'–25º31'e. hoffman (1996) classified this section as eastern mixed nama karoo (52), while acocks (1988) classified the area as false karroid broken veld (37). the wilgerboom river acts as the main functional drainage system, that extends from the old park into the study area. the eastern and western areas are mountainous and consist of moderate to steep slopes. the vegetation comprises mainly of a complex mixture of grass-and-shrub-dominated vegetation types (hoffman 1996). the central area comprises lower lying drainage lines characterised by mixed acacia karroo veld, as well as disturbed fields, previously used for cultivation purposes. however, this cultivated field covers less than 1 % of the total study area. land types three land types, namely fc, ib and da are prominent in the study area. a remarkable association between the major plant communities and the different land types have been observed. the land type concept has frequently provided a useful basis for descripkoedoe 46/2 (2003) 10 issn 0075-6458 fig. 1. location of the study area in mountain zebra national park (compiled by s. holness). km brown.qxd 2005/12/09 10:36 page 10 sanparks tion of vegetation (bezuidenhout 1993; eckhardt 1993; götze 2002). the geology in the fc land type consists of mudstone, shale and sandstone of the beaufort group of the karoo sequence with rare dolerite intrusions. the main topographical positions present in this land type are footslopes, midslopes and valley bottoms. glenrosa and oakleaf soil forms are characteristic of this land type. underlying limestone is present in the valley bottoms (land type survey staff 1999). the ib land type can be associated with midslopes in mountainous areas. the rock cover in this land type is approximately 50–60 % and consists of exposed sandstone rocks. the geology depicts dolerite with mudstone, shale and sandstone of the beaufort group of the karoo sequence (land type survey staff 1986). the mispah soil form is prominent in this land type (land type survey staff 1999). the upper plateau area in the south western section of the study area includes the da land type with a rock cover of less than 50 %, with mudstone, shale and sandstone of the balfour formation in the beaufort group of the karoo sequence as the underlying geology (land type survey staff 1999). climate the study area is located within the summer rainfall area of south africa, with an average rainfall of 389 mm per annum. the highest annual rainfall recorded was 651 mm in 1977, while the lowest rainfall recorded was 153 mm, in 1966. the average monthly rainfall varies between 11 mm in the winter to 61 mm in summer. during summer (september–march) the average maximum temperatures vary between 23–28 ºc, while in the winter (april–august), the average minimum varies between 0–8 ºc (brown & bezuidenhout 2000). methods the method applied in this study was based on the method described and applied by brown & bezuidenhout (2000) in the de rust section of the mznp. to incorporate this study with similar studies in mznp it is crucial that the same procedure is followed. the study area was stratified into physiognomic-physiographic units by using 1:50 000 stereo aerial photographs. relevés were compiled at 47 stratified random plots. plot sizes were fixed at 400 m² (brown 1997). all plant species were recorded in each sample plot and a cover-abundance score was allocated to each species using the braun-blanquet cover-abundance scale (mueller-dombois & ellenberg 1974). an estimation of the percentage canopy cover and average height of the herbaceous, shrub, and tree layer in each sample plot was also noted. shrubs are defined as woody species varying in height between >0-3 m and trees as woody species taller than 3 m (brown 1997; brown & bezuidenhout 2000). topographical characteristics such as aspect, slope, rockiness and soil surface were also noted. the bbpc suite (bezuidenhout et al. 1996) was used to analyse the floristic data. by applying a two-way indicator species analysis (twinspan) (hill 1979) to the floristic data, a first approximation of the main plant communities was derived. refinement of the classification was achieved by applying braun-blanquet procedures. soil nomenclature follows the classification of the soil classification working group (1991). although certain taxon names have changed, the taxon names of this publication conform to those of arnold & de wet (1993). structural terminology is according to edwards (1983). no attempt was made to formally fix syntaxa names as this is normally avoided in detailed local studies (coetzee 1983). results classification the following eight plant communities that can be grouped in five major community types were derived from the analysis (table 1): 1. buddleja glomerata–rhus lucida woodland 2. themeda triandra–merxmuellera disticha grassland issn 0075-6458 11 koedoe 46/2 (2003) brown.qxd 2005/12/09 10:36 page 11 sanparks 2.1 aloe striatula–merxmuellera disticha sub-community 2.2 walafrida saxatillis – merxmuellera disticha sub-community 3. acacia karroo–pentzia globosa woodland 3.1 setaria sphacelata – rhus longispina sub-community 3.2 enneapogon scoparius – rhigozum obovatum sub-community 3.3 eragrostis obtusa–pentzia globosa sub-community 4. panicum maximum–acacia karroo woodland 5. salsola kali–medicago sativa old cultivated fields description of the plant communities 1. buddleja glomerata–rhus lucida woodland this plant community is found on the southern and south-east facing slopes in the western corner of the study area (fig. 2). characteristic of this community is steep slopes with scattered large rocks (sandstone). it can also be associated with the ib land type and mispah soil form. rockiness was estimated at approximately 60 %. the cover of the tree layer (buddleja glomerata and acacia karroo) was estimated at 30 %, the shrub layer at 60 % (rhus lucida and tarchonanthus camphoratus) and the herbaceous layer at 10 %. tree height varies between 4-5 m while the shrub height varies between 2-3 m. koedoe 46/2 (2003) 12 issn 0075-6458 fig. 2. the plant communities of the ebenhaezer section of the mountain zebra national park. brown.qxd 2005/12/09 10:36 page 12 sanparks this community is characterised by the presence of the tree buddleja glomerata, the grass melinis nerviglumis and the forbs solanum supinum, stachys cymbalaria, stachys linearis, moraea polystachya and hibiscus pusillus (species group a, table 1). the vegetation is dominated by the trees buddleja glomerata (species group a) and rhus lucida (species group g). the grasses digitaria eriantha, sporobolus fimbriatus (species group g) and enneapogon scoparius (species group e) are prominent locally, while the shrubs grewia occidentalis, maytenus heterophylla (species group g) and the dwarf shrub chrysocoma ciliata (species group g) are also present in this community (table 1). this community shows affinity with the rhus lucida-buddleja glomerata shrubland described by brown & bezuidenhout (2000) in the de rust section of mznp. 2 themeda triandra–merxmuellera disticha grassland the themeda triandra–merxmuellera disticha grassland community is found in the south-western corner of the study area (fig. 2). this community is situated on the plateau and midslopes (ib and da land types) with a gradient of approximately 8–25 º. the habitat consists of moist shallow soil on a mispah soil form and a rock cover of approximately 40–55 %. tree cover is estimated at 1 %, while the shrub stratum covers between 1–30 % of the area (fig. 2). the herbaceous cover is estimated at between 40–55 %. the average tree height varies between 1.5–3 m, while the average height of the shrubs are 0.5 m. the herbaceous layer is approximately 0.3 m tall. this community is characterised by the presence of the grasses merxmuellera disticha and themeda triandra (species group b, table 1) that also dominates the vegetation and can be divided into the following two sub communities: 2.1 aloe striatula–merxmuellera disticha sub-community this sub-community is found on steep south facing slopes in the southern section of the study area (fig. 2). the sub-community can be found on steep slopes (25 º gradient) in moist soil, which is covered with large rocks (sandstone) and has an estimated 50 % rock cover. a mispah soil form with underlying sandstone and shale is dominant in this community and situated on an ib land type. the cover of the tree layer is estimated at only 1% with an average height of 2.5 m while shrub layer with an average height of 1.5 m has an estimated 25–30 % cover. the herbaceous layer is well represented and covers an area of approximately 55 %. this community is characterised by the presence of species group c (table 1) and includes the succulent aloe striatula, the tree cussonia paniculata, the shrubs rhus erosa, myrsine africana, diospyros austroafricanum, rhus dregeanum, euclea crispa, tarchonanthus camphoratus, asparagus retrofractus, the grass eragrostis curvula, the fern cheilanthes hirta, and the forb euryops nodosus. the vegetation is dominated by the grasses themeda triandra, merxmuellera disticha (species group b) and the succulent aloe striatula (species group c, table 1). other prominent species associated with this vegetation type includes the tree cussonia paniculata and the shrub rhus erosa (species group c, table 1). although this is the smallest community identified, less than one hectare in size, it contributes to the species richness of the area. 2.2 walafrida saxatillis–merxmuellera disticha sub-community this vegetation is localised in the southern section of the study area, situated on the upper midslopes to the plateau area (fig. 2). this sub-community is associated with the da land type with shallow soil. rock cover is estimated at 40–55 % with medium size rocks present on an 8 º slope. issn 0075-6458 13 koedoe 46/2 (2003) brown.qxd 2005/12/09 10:36 page 13 sanparks koedoe 46/2 (2003) 14 issn 0075-6458 ta bl e 1 a ph yt os oc io lo gi ca l t ab le o f t he e be nh ae ze r se ct io n of th e m ou nt ai n ze br a n at io na l p ar k (n on -d ia gn os tic s pe ci es a re o m itt ed fr om th e ta bl e) . brown.qxd 2005/12/09 10:36 page 14 sanparks issn 0075-6458 15 koedoe 46/2 (2003) ta bl e 1 (c on tin ue d) brown.qxd 2005/12/09 10:36 page 15 sanparks koedoe 46/2 (2003) 16 issn 0075-6458 ta bl e 1 (c on tin ue d) brown.qxd 2005/12/09 10:37 page 16 sanparks issn 0075-6458 17 koedoe 46/2 (2003) the cover of the tree layer is estimated at only 1% of the area, with an average height ranging between 2.5–3 m, and is dominated by acacia karroo. the shrub layer has an estimated 1 % coverage with an average height of 1.5 m, while the herbaceous layer with an average height of 0.6 m has an estimated 45–65 % cover. this community is characterised by the presence of the dwarf shrubs walafrida saxatillis and felicia fascicularis (species group d). the vegetation is dominated by the grasses themeda triandra and merxmuellera disticha that covers approximately 55% of the area. other prominent species include the tree acacia karroo (species group j) and the grass eragrostis chloromelas (species group i, table 1). the grass component showed indications that the grass sward may become moribund. 3. acacia karroo – pentzia globosa woodland the acacia karroo–pentzia globosa woodland is found on all warm and dry footslopes and upper midslopes and includes the majority of plateau areas in this section (fig. 2). this community is associated with the ib and fc land types, with mispah (midslope) and glenrosa (footslope) as the dominant soil types. soil depth is limited by underlying limestone in the glenrosa soil form. rock cover varies between 15–60 % with underlying sandstone, shale and limestone. the cover of the tree layer is estimated at 1–20 %, while the shrub stratum covers approximately 5–45 % of the area. the cover of the herbaceous layer is estimated at 20–60 %. tree height averages 2–3.5 m, while shrubs are on average between 1.5–3.5 m in height. the herbaceous layer is approximately 0.3 m tall. this community is characterised by the presence of the tree ehretia rigida, the dwarf shrubs helichrysum dregeanum, limeum aethiopicum, the grasses aristida diffusa, aristida adscensionis, aristida junciformis, aristida congesta, cymbopogon plurinodis, enneapogon scoparius and the forbs sutera virgulosa, blepharis capensis, hermannia vestita, solanum sisymbrifolium and ledebouria marginata (species group e, table 1). the vegetation is dominated by the dwarf shrub pentzia globosa (species group k) and the grass aristida diffusa (species group e). the dwarf shrub eriocephalus ericoides (species group k) and the grasses enneapogon scoparius and aristida adscensionis (species group e) are co-dominant within this community (table 1). other species also prominent include the tree acacia karroo (species group l), the shrub rhus longispina (species group k), the grasses setaria sphacelata, heteropogon contortus (species group k), aristida congesta subsp barbicollis (species group l) and the forb walafrida geniculata (species group l, table 1). the abundance of the pioneer grass aristida adscensionis (species group e) in large sections of this community indicates localised overgrazing that can be ascribed to previous farming activities in the area. this community is divided into three subcommunities namely the setaria sphacelata –rhus longispina sub-community, the enneapogon scoparius–rhigozum obovatum sub-community and the eragrostis obtusapentzia globosa sub-community: 3.1 setaria sphacelata–rhus longispina sub-community this woodland is found on warm, dry footslopes to midslopes and plateaux (fig. 2). this community is associated with the ib land type. the surface is covered with small to medium sized rocks covering an estimated 15–60 % of the area. the dominant soil-rock complex is a mispah soil with underlying sandstone and shale. tree cover is estimated at 15 % with an average height of 2.5–3 m, while the shrub layer covers between 5–10 % of the area and is approximately 1.5–3.5 m in height. the cover of the herbaceous layer is estimated at 60 % of the area with an average height of 0.3 m. brown.qxd 2005/12/09 10:37 page 17 sanparks koedoe 46/2 (2003) 18 issn 0075-6458 this community is characterised by the absence of a diagnostic species group and is dominated by the dwarf shrub pentzia globosa (species group k) and the tree acacia karroo (species group l) while the shrub rhus longispina and the grass setaria sphacelata (species group k) are co-dominant (table 1). prominent species are the dwarf shrub eriocephalus ericoides (species group k) and the grasses aristida congesta subsp barbicollis (species group l), heteropogon contortus (species group k) and digitaria eriantha (species group g, table 1). 3.2 enneapogon scoparius–rhigozum obovatum sub-community this vegetation type can be found on lower and upper midslopes throughout the section (fig. 2). the fc land type is associated with this community. underlying limestone is characteristic of this land type on the footslope and valley bottom areas (land type survey staff 1999). the geology consists of mudstone and sandstone while the soil is classified as the glenrosa soil form. soil depth is limited by the under lying limestone and averages 150 mm. small to medium sized rocks are scattered over the area with an average cover of 20–40 %. the tree layer covers 1–5 % of the area with an average height of 2–3 m, while shrub cover is estimated at 5–20 % and varies in height between 1.5–3 m. the herbaceous layer covers an estimated 45 % of the area with an average height of 0.3 m. this community is characterised by species from species group f (table 1) and include the shrubs rhigozum obovatum, carissa macrocarpa, colpoon compressum, euclea crispa and the forb gallenia spp. the vegetation is dominated by the shrubs rhus longispina (species group k), rhigozum obovatum (species group f) and the grasses aristida diffusa and enneapogon scoparius (species group e, table 1). other species also prominent include the tree acacia karroo and the dwarf shrubs pentzia globosa, eriocephalus ericoides and felicia filifolia (species group k, table 1). though not listed in table 1, the grass cenchrus ciliaris was found to be locally dominant within this sub-community at higher points below cliffs. this community was previously heavily grazed and some places are severely disturbed with signs of sheet and donga erosion present. the enneapogon scoparius–rhigozum obovatum sub-community shows affinity with the enneapogon scoparius–ehretia rigida sub-community of the de rust area, described by brown & bezuidenhout (2000). 3.3 eragrostis obtusa–pentzia globosa sub-community this sub-community can be found on plateaux and midslopes throughout the study area (fig. 2). this community is associated with the ib land type and comprises approximately 32 % of the study area. this habitat consists of shallow soil on a mispah soil form and a rock cover that varies between 5–55 %. the habitat is also found on plateaux and upper midslopes facing east and west, as well as footslopes leading into drainage lines. trees and shrubs comprise only 1–2 % of the vegetation and are seldom taller than 2.5 m. the cover of the herbaceous layer varies between 20–45 % with an average height of 0.3 m. the presence of the dwarf shrub helichrysum dregeanum, the grasses oropetium capense, urochloa panicoides and the forb blepharis mitrata is characteristic for this large sub-community (species group h, table 1). the vegetation is dominated by the dwarf shrubs pentzia globosa and eriocephalus ericoides (species group k) together with the grass eragrostis obtusa (species group i, table 1). other species also prominent include the dwarf shrub walafrida geniculata (species group k) and the grasses aristida diffusa, aristida adscensionis, enneapogon scoparius (species group e), heteropogon contortus, eragrostis lehmanniana (species group k) and tragus koelerioides (species group l, table 1). brown.qxd 2005/12/09 10:37 page 18 sanparks issn 0075-6458 19 koedoe 46/2 (2003) locally some patches are dominated by the grasses cynodon incompletus and chloris virgata (species group l, table 1). these patches have been heavily grazed in the past by goats and sheep, leading to the presence of these pioneer grasses. bush encroachment by the tree acacia karroo (species group l, table 1) was also noted in some areas within this sub-community. 4. panicum maximum–acacia karroo woodland this community consists of the drainage lines and the wilgerboom river, and comprises approximately 30 % of the study area (fig. 2). the wilgerboom river is perennial while the drainage lines only flow after heavy thunderstorms. this area has been seriously mismanaged and some areas are disturbed with erosion patches visible. the habitat is situated on the fc land type with hutton, oakleaf and swartland soil forms present. rock cover is less than 5 % and consists mainly of small rocks. tree height varies between 3.5–5 m while tree cover varies between 20 % in the upper catchment areas to 60 % in the drainage lines and river. the average shrub height is approximately 2.5 m and covers 15 % of the area. the herbaceous layer covers an estimated 35 % of the area with an average height of 0.25 m. this woodland is characterised by the presence of the dwarf shrub zygophyllum incrustatum and the grass panicum maximum (species group j, table 1). the vegetation is dominated by the tree acacia karroo (species group l) and the grass panicum maximum (species group j, table 1). other prominent species include the tree lycium oxycarpum (species group k), the dwarf shrubs zygophyllum incrustatum (species group j), pentzia globosa and the grasses chloris virgata and cynodon incompletus (species group k, table 1). pioneers species like atriplex semibaccata and salsola kali (species group n) are prominent on disturbed soil. this community has affinity with the becium burchellianum-acacia karroo woodland and sporobolus africanus-acacia karroo woodland described by brown & bezuidenhout (2000). 5. salsola kali–medicago sativa old cultivated fields this small community (< 1 % of total area) consists of old cultivated fields situated in the drainage line of the study area (fig. 2). rock cover is 0 % and slope 0–1 %. the vegetation consists only of a herbaceous component with the grasses and forbs covering an estimated 80 % of the area with an average height of 0.4 m. this community is characterised by the presence of the forbs medicago sativa (lucerne) amaranthus hybridus, conyza albida and alternanthera pungens (species group m, table 1). this old field community is dominated by lucerne medicago sativa (species group m), while the forbs salsola kali and atriplex semibaccata are also prominent locally. due to the disturbed nature of this community it does not contribute to the conservation or biological value of this study area. discussion and conclusion from the eight communities identified in the study area, three communities have affinities with four communities described by brown & bezuidenhout (2000) for the de rust section of mountain zebra national park. a further two communities indicate affinity with two communities described by van der walt (1980) in the old proclaimed section of mznp. this study proves that the extended land incorporated into the park contributes to the biological diversity of the park by adding more vegetation communities and habitat types to the area. the expansion of mznp does not only contribute floristically to the diversity of the park but also faunally due to the inclusion of new habitats into the park which make it suitable for the introduction of brown.qxd 2005/12/09 10:37 page 19 sanparks koedoe 46/2 (2003) 20 issn 0075-6458 larger herbivores that occurred in this region before. data obtained from this study will be incorporated into the draft management plan for the larger mznp. this will be in line with the mission and vision statement of south african national parks, for example, to conserve the faunal and floral assemblages and ecological processes that characterise the northeastern karoo areas, and to actively preserve this for the appreciation by visitors. acknowledgements the following individuals and institutions are sincerely thanked: national research foundation (nrf) and technikon sa for financing this study; south african national parks, especially park management of mountain zebra, for allowing and encouraging the research; dr stephen holness for the assistance with the maps and mr ernest daemane (kimberley south african national parks herbarium ksan) for verifying the plant species names. references acocks, j.p.h. 1988. veld types of south africa. 3rd ed. memoirs of the botanical survey of south africa 57: 1-146. arnold, t.h. & b.c. de wet. 1993. plants of southern africa: names and distribution. memoirs of the botanical survey of south africa. 62: 1-825. bezuidenhout, h. 1993. syntaxonomy and synecology of western transvaal grasslands. ph.d dissertation. university of pretoria, pretoria. bezuidenhout, h., h.c. biggs & g.j. bredenkamp. 1996. a process supported by the utility bbpc for analysing braun-blanquet data on a personal computer. koedoe 39: 107-112. brown, l.r. 1997. a plant ecological and wildlife management plan of the borakalalo nature reserve, north-west province. ph.d. dissertation. university of pretoria, pretoria. brown, l.r. & h. bezuidenhout. 2000. the phytosociology of the de rust section of the mountain zebra national park, eastern cape. koedoe 43(1): 1-18 coetzee, b.j. 1983. phytosociology, vegetation structure and landscapes of the central district, kruger national park. dissertationes botanicae 69: 1-456. eckhardt, h.c. 1993. a synecological study of the vegetation of the north-eastern orange free state. m.sc. thesis. university of pretoria, pretoria. edwards, d. 1972. botanical survey and agriculture. proceedings of the grassland society of southern africa 7: 15-19. edwards, d. 1983. a broad-scale structural classification of vegetation for practical purposes. bothalia 14: 705-712. götze, a.r. 2002. the classification of plant communities and the evaluation of restoration technologies in different land-use areas in the vhembe-dongola national park, northern province, south africa. m. sc. thesis. potchefstroom university for christian higher education, potchefstroom. harper, j.l. & d.l. hawksworth. 1995. preface. p.10. in: hawksworth, d.l. (ed) biodiversity, measurement and estimation. chapman & hall. london. hill, m.o. 1979. twinspan: a fortran program for arranging multivariate data in an ordered twoway table by classification of individuals and attributes. new york: cornell university. hoffman, t. 1996. eastern mixed nama-karoo. p.55. in: low, a.b. & a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: departement of environmental affairs & tourism. land type survey staff. 1986. land types of the map se27/20 witdraai, 2720 noenieput, 2722 kuruman, 2724 christiana, 2820 upington, 2822 postmasburg. menoirs of the agricutural natural resources of southern africa 3: 1-185. land type survey staff. 1999. land types occurrence (maps) and areas of 3224 graaff-reinet. unpublished information obtained from institute for soil, climate and water, pretoria. mueller-dombois, d. & h. ellenberg. 1974. aims and methods of vegetation ecology. new york; wiley. pond, u., b.b. beesley, l.r. brown & h. bezuidenhout. 2002. floristic analysis of the mountain zebra national park, eastern cape. koedoe 45 (1): 35-37. soil classification working group. 1991. soil classification: a taxonomic system for south africa. memoirs on the agriculrural natural resources of south africa 15: 1-262. van der walt, p.t. 1980. a phytosociological reconnaissance of the mountain zebra national park. koedoe 23:1-32. wahl, m. & k. naude. 1996. national register of protected areas in south africa. pretoria: department of environmental affairs and tourism. brown.qxd 2005/12/09 10:37 page 20 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <feff004200720075006700200064006900730073006500200069006e0064007300740069006c006c0069006e006700650072002000740069006c0020006100740020006f0070007200650074007400650020005000440046002d0064006f006b0075006d0065006e0074006500720020006d006500640020006800f8006a006500720065002000620069006c006c00650064006f0070006c00f80073006e0069006e0067002000740069006c0020007000720065002d00700072006500730073002d007500640073006b007200690076006e0069006e0067002000690020006800f8006a0020006b00760061006c0069007400650074002e0020005000440046002d0064006f006b0075006d0065006e007400650072006e00650020006b0061006e002000e50062006e006500730020006d006500640020004100630072006f0062006100740020006f0067002000520065006100640065007200200035002e00300020006f00670020006e0079006500720065002e00200044006900730073006500200069006e0064007300740069006c006c0069006e0067006500720020006b007200e600760065007200200069006e0074006500670072006500720069006e006700200061006600200073006b007200690066007400740079007000650072002e> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 terblance 2.qxd the vegetation of three localities of the threatened butterfly species chrysoritis aureus (lepidoptera: lycaenidae) r.f. terblanche, t.l. morgenthal and s.s. cilliers terblanche, r.f., t.l. morgenthal and s.s. cilliers. 2003. the vegetation of three localities of the threatened butterfly species chrysoritis aureus (lepidoptera: lycaenidae). koedoe 46(1): 73–90. pretoria. issn 0075-6458. the vegetation and habitat characteristics of three localities of chrysoritis aureus at the alice glockner nature reserve, suikerbosrand nature reserve and malanskraal farm near heidelberg in south africa, were compared. a numerical classification technique, twinspan, was used and refined by using braun blanquet procedures to classify the vegetation at the different localities. a dca ordination was applied to confirm the results of the classification. although the general vegetation structure at the three habitats of chrysoritis aureus were found to be similar, marked differences in the floristic composition were evidenced. a different sub-community, compared to the vegetation at suikerbosrand and alice glockner nature reserve, was recorded at the malanskraal habitat of chrysoritis aureus. these differences in floristic composition, but with similarities in vegetation structure, indicate the possible importance of fire for the ultimate survival of these butterflies in the rocky highveld grassland. the host plant of chrysoritis aureus, clutia pulchella, collected at malanskraal differed markedly and consistently in their morphology, compared to the individuals from the habitats at suikerbosrand and alice glockner nature reserve. these differences in the floristic composition of one of the habitats compared to the others, raise research questions concerning the butterfly metapopulation structure, since the subpopulations seem to be adapted to slightly different habitat conditions. the variation in the habitat suggests that the “last remaining locality scenario” for other localised butterflies in south africa, such as orachrysops niobe, needs to be redressed. management strategies are addressed while the importance of conserving rare, localised ecosystems are highlighted by the phytosociological study. key words: myrmecophilous, lycaenidae, chrysoritis, clutia, crematogaster, habitat, phytosociology, conservation r.f. terblanche (drkrft@puknet.puk.ac.za), t.l. morgenthal & s.s. cilliers, school of environmental sciences and development, private bag x6001, potchefstroom university for che, potchefstroom, 2520 republic of south africa. issn 0075-6458 73 koedoe 46/1 (2003) introduction the butterfly chrysoritis aureus—previously known as poecilmitis aureus before the new combination was designated by heath (1997) —is considered to be rare in the red data book of butterflies (henning & henning 1989) and the updated list of the red data book of butterfly species (henning & henning 1995). c.r. barrett and f. coetzee discovered chrysoritis aureus on christmas day 1959 on a rocky slope below the water tower near the natal road of heidelberg in gauteng (pringle et al. 1994). during 1983, chrysoritis aureus was added to the protected wild animal list of the former transvaal under ordinance 12, since the type locality had been the only known area where the butterfly ocurred (de wet 1987). g.a. henning found another locality near greylingstad (pringle et al. 1994). members of the lepidopterists’ society of africa found another locality in suikerbosrand nature reserve (henning & henning 1989) after teams of terblance 2.qxd 2005/12/09 10:31 page 73 lepidopterists carried out organised extensive exploration in this reserve (fig. 1). the colony at suikerbosrand seems to be limited in numbers (henning & henning 1989; roos & henning 2000) though the numbers tend to remain stable (the word ‘colony’ refers to the subpopulations of these butterflies, that is where they actually breed. the adults usually also fly at or close to these restricted areas). in the suikerbosrand nature reserve, the only recorded habitat of the butterfly is limited to the north eastern extreme of the reserve, even though the lepidopterists’s society of africa organised surveys that covered many different habitats in the reserve (g.a. henning 1986a, 1986b, 1987). after decades of exploring by a number of lepidopterists in the hilly areas of gauteng and the adjacent mpumalanga (table 1), only five confirmed localities of chrysoritis aureus were reported. the distribution of the butterfly can be described as patchy and koedoe 46/1 (2003) 74 issn 0075-6458 table 1 the locality, altitude and geological system present at the confirmed localities of chrysoritis aureus locality province locality * altitude (m) geological supergroup alice glockner nr 8 km s heidelberg gauteng 26º34's, 28º22'e(2628cb) 1690 witwatersrand hill n of greylingstad mpumalanga 26º44's, 28º44'e(2628da) 1810 ventersdorp malanskraal locality 14 km w balfour mpumalanga 26º39's, 28º26'e(2628cb) 1780 ventersdorp suikerbosrand nr gauteng 26º29's, 28º18'e(2628ad) 1720 witwatersrand type locality at heidelberg gauteng 26º29's, 28º21'e(2628ad) 1600 witwatersrand * the grid reference reading is from the north western corner of each minute by minute square. **the malanskraal locality is close to the borderline between the ventersdorp supergroup and the witwatersrand supergroup. fig. 1 the location of the habitats of chrysoritis aureus that were studied. terblance 2.qxd 2005/12/09 10:31 page 74 chrysoritis aureus is very localised at each of these localities. the three habitats studied each cover approximately 200 m² – 400 m². all the recorded localities of chrysoritis aureus are situated at sites where formal or informal urban development pose an everincreasing threat to the habitat. habitat destruction across the world as the main cause for the extinction of populations of localised butterflies, is well recognised, thereby emphasising the primary importance of habitat conservation (s.f. henning 1987; larsen 1995; kudrna 1995; munguira 1995; new 1995; oates 1995; opler 1995; pullin et al. 1995; thomas 1995; warren 1995). an ongoing comparative study of the chrysoritis aureus habitats (vegetation) is therefore imperative to gain knowledge for developing strategies for the conservation of these habitats. a description of the vegetation of the chrysoritis aureus habitats is needed if management practices such as translocation of butterflies or habitat restoration is considered. vegetation studies of the habitats of the threatened golden sun moth in australia (o’dwyer & attiwill 2000), provided essential information for the restoration of its habitat. a vegetation analysis of the then only known habitat of a threatened butterfly, aloeides dentatis subsp. dentatis, in south africa, (deutschländer & bredenkamp 1999) was very applicable for its conservation management. for this study, the vegetation of three habitats were compared to identify possible key factors and to detect possible differences between these habitats. this study also forms part of an ongoing investigation into the reasons why the subpopulations of the butterfly are confined to such small areas. in south africa, studies on myrmecophilous butterflies, such as chrysoritis aureus, are limited to life history descriptions or reporting of information on the host plant and host ant. some clues for the confinement of chrysoritis aureus to its habitats can be found in its life history (henning 1983). the host plant of the larvae is the small shrub clutia pulchella l. sens. lat. (euphorbiaceae) which is widely distributed in south africa (arnold & de wet issn 0075-6458 75 koedoe 46/1 (2003) fig. 2. clutia pulchella ("heidelberg morphotype"), the host plant of chrysoritis aureus. 1993; retief & herman 1997) (fig. 2). the larvae of chrysoritis aureus are also constantly tended by ants of the genus crematogaster (henning 1983). during the day, the larvae shelter for protection and only appear at night to feed on their host plants (henning 1987a). well developed dorsal nectary organs (cottrell 1984) or honey glands (henning 1983; 1987a) are present on the larvae and the ants are attracted to feed on the secretions of these organs (henning 1983, 1987a). in the absence of the ants, a fungal infection occurs in the dorsal nectary organ and the larvae often die after a few days (henning 1983) (fig. 3). since both the ant and the host plant are needed for survival of chrysoritis aureus at any locality, the habitat requirements are complicated. these include specific local vegetation communities. we therefore investigated the vegetation, to conterblance 2.qxd 2005/12/09 10:31 page 75 tribute to a niche/habitat profile of chrysoritis aureus. study area three localities inhabited by chrysoritis aureus were studied: one at the suikerbosrand nature reserve, one at the alice glockner nature reserve near heidelberg, gauteng and one at the farm malanskraal (fig. 1). the habitats of chrysoritis aureus are situated on the south-facing slopes of rocky ridges and hills. the first two habitats studied are situated on the quartzites of the wiwatersrand supergroup while the habitat at malanskraal habitat is situated on the lavas of the ventersdorp supergroup. the landscape is rugged and broken in appearance, since the witwatersrand system was subjected to considerable folding, faulting and erosion in the past (du toit 1954). the study sites are in the summer rainfall area, with temperate summers and cold koedoe 46/1 (2003) 76 issn 0075-6458 fig. 3. the larva of chrysoritis aureus. the two white structures at the anterior (head) area, are tubercle organs. note the fungal growth at the posterior end at the dorsal nectary organ. photo: r.f. terblanche. frosty winters. according to information obtained from the south african weather service, the long-term average annual rainfall data from nearby weather stations vary considerably between 620-750 mm per annum (south african weather service, private bag x097, pretoria, 0001). variation in topography, the rock cover and vegetation structure results in different mesoand microclimates. the sites that were investigated are situated in the grassland biome (rutherford & westfall 1994; low & rebelo 1996), the bankenveld (acocks 1988) or rocky highveld grassland (bredenkamp & van rooyen 1996). materials and methods sample plots (5 m x 5 m) were placed on the habitats of chrysoritis aureus. these plots fitted well on the restricted habitats of chrysoritis aureus at the alice glockner nature reserve and malanskraal, while four sample plots were adequate for the habitat at suikerbosrand. since the range of occurence of chrysoritis aureus is so small, the samples are thought to be representative. a transect of sample plots over the habitat from the summit to the midslopes and down to a valley bottom, was surveyed at the alice glockner nature reserve to compare the vegetation on the slopes close to the habitats with the habitats of chrysoritis aureus (fig. 4). the floristic composition and cover abundance according to the braun-blanquet scale (mueller-dombois & ellenberg 1974) were noted in each plot during surveys on 12 and 13 february 1999. the number of clutia pulchella individuals that occurred in each sample plot were determined. plant specimens were deposited in the a.p. goossens herbarium of the potchefstroom university for c.h.e. the height of trees (von breitenbach 1990; von breitenbach et al. 2001) was noted. these species will, therefore, be referred to as trees, even if they are physionomically not regarded as trees. the height classes as described by edwards (1983) were used. habitat features such as slope, aspect and rockiness were noted, though not quantitatively measured. notes on the presence of chrysoritis aureus and other butterflies were continuously made by means of direct observations on 7 and 13 november, 1998 as well as 12 and 13 february 1999. the local distribution of chrysoritis aureus was confirmed by p. roos and g.a. henning who have visited the localities for a terblance 2.qxd 2005/12/09 10:31 page 76 number of years (roos & henning 2000). the same procedure was also followed for the breeding sites, i.e., where pupae, larvae, and oviposition by the female were evidenced. the ants associated with chrysoritis aureus have been identified by dr. h.g. robertson, south african museum, cape town, 8000. the twinspan classification algorithm (hill 1979a) was used to analyse the floristic data and the result was refined by using braun blanquet procedures by means of the bbpc-suite (bezuidenhout et al. 1996). a dca ordination was applied (hill 1979b) to confirm the results of the classification. results and discussion the host plant mr. i. bampton discovered the host plant of chrysoritis aureus in january 1976 (owenjohnston 1991). it is sometimes reported to be clutia galpinii (owen-johnston 1991; heath 2001). clutia galpinii is considered to be a synonym of clutia pulchella (retief & herman 1997). the entity clutia galpinii refers to a relatively smaller plant of 91 cm – 152 cm, whereas clutia pulchella reaches heights up to 244 cm (prain 1925). clutia pulchella individuals that have been measured near or at the habitats of chrysoritis aureus, ranged from 13 cm to more than 200 cm tall (in more dense vegetation) and fits the descriptions of both entities overviewed by prain (1925). the plant specimens from the localities of chrysoritis aureus were compared to the material at the national herbarium. it does not fit clutia monticola of which the leaves are sessile (retief & herman 1997). the host plant of chrysoritis aureus was confirmed to be only clutia pulchella at all the localities. clutia pulchella constantly showed specific variation from different localities. individuals from malanskraal have more lanceolate leaves, are of a less bluish-green colour and the upper surface of the leaves are covered with velvety hairs. this is in contrast to the more ovate, more glaucous and much less hairy leaves of individuals from the alice glockner nature reserve and suikerbosrand nature reserve. the reasons for these differences could be due to the different geological systems where these plants occur. these morphotypes may be the subject of more detailed study on the taxonomic status of the host plant. larvae of the different localities issn 0075-6458 77 koedoe 46/1 (2003) fig. 4. position of the 5m x 5m sample plots from the summit (ac 14) to the lower valley bottom (ac 8, ac 9) of the hill (southern aspect) at alice glockner nature reserve. the seven plots (ac 1–ac 7) that are inhabited by chrysoritis aureus were put out horizontally next to each other along the rocky ridge. terblance 2.qxd 2005/12/09 10:31 page 77 seems to be adapted to feed on these different morphotypes of clutia pulchella. the number of clutia pulchella individuals at the sample plots where chrysoritis aureus is present varies (2-14 indiv/25m²) with an average of 7.8 individuals/25m² for all the habitats combined—that means an average of 0.31 clutia pulchella indiv/m². the locality at the alice glockner nature reserve showed the lowest average of clutia pulchella individuals namely 6.7 indiv/25m². though chrysoritis aureus was not present at sample plots ac 10, ac 11 and ac 14, host plant individuals were recorded at these sites. the number of host plants at sample plots ac 10 and ac 11 (table 2) seems to be adequate to serve as possible breeding areas. all the sites where clutia pulchella individkoedoe 46/1 (2003) 78 issn 0075-6458 table 2 the number of clutia pulchella counted as well as the status of chrysoritis aureus at the plots (5m x 5m) plot number of status of number clutia pulchella c. aureus individuals 1 = present 0 = absent ac 1 4 1 ac 2 3 1 ac 3 4 1 ac 4 7 1 ac 5 13 1 ac 6 14 1 ac 7 2 1 ac 8 0 0 ac 9 0 0 ac 10 5 0 ac 11 4 0 ac 12 0 0 ac 13 0 0 ac 14 2 0 mk 15 7 1 mk 16 8 1 mk 17 10 1 sb 18 6 1 sb 19 10 1 sb 20 8 1 sb 21 13 1 uals were present are characterised by the presence of large rocks and this seems to be one of the key habitat requirements of the host plant. the factors that can play a role in excluding chrysoritis aureus from establishing at sample plots ac 10 and ac 11, are not known. however, both sample plots ac 10 and ac 11 are situated on the lower midslopes compared to the other sites where clutia pulchella occurs at higher midslopes in the alice glockner nature reserve (fig. 4). another subject for further research is the local distribution and vitality of clutia pulchella in the vicinity of chrysoritis aureus habitats. it was observed that clutia pulchella may have higher densities and seemed to have better vitality (higher plants) in some of the surrounding areas, where no plots were put out, as chrysoritis aureus was not present here. a quantitative survey may shed more light on the above observation. vegetation classification the floristic composition of the plant communities identified is represented in a phytosociological table (table 3) and the results of the dca-ordination (fig. 5). the phytosociological analysis resulted in the following two communities with their sub-communities and variants: 1. melinis nerviglumis-aristida transvaalensis community 1.1. melinis nerviglumis-aristida transvaalensis–eragrostis curvula subcommunity 1.2 melinis nerviglumis-aristida transvaalensis-loudetia simplex sub-community 1.2.1. anthospermum hispidulum variant 1.2.2 digitaria monodactyla variant 1.3. melinis nerviglumis-aristida transvaalensis-englerophytum magalismontanum sub-community 2. elionurus muticus-eragrostis trichophora community terblance 2.qxd 2005/12/09 10:31 page 78 community description 1. melinis nerviglumis aristida transvaalensis community this community is found on the south-facing slopes and on the terraces within the slopes, and includes a representation of all the habitats of chrysoritis aureus that were studied. apart from the terraces, large rocks (often exceeding one metre in length) are conspicuous on all the sites. the community is characterised and dominated by the grasses melinis nerviglumis, aristida transvaalensis and cymbopogon plurinodis (species group a, table 3). three subcommunities are recognised: 1.1 melinis nerviglumis-aristida transvaalensis -eragrostis curvula subcommunity this sub-community occurs on the southfacing slopes at malanskraal on the ventersdorp supergroup (table 1). it is characterised by the absence of loudetia simplex and the constant high cover-abundance values of eragrostis curvula, helichrysum nudifolium and the constant presence of garuleum pinnatifidum and tephrosia capensis (species group b, table 3). this sub-community is further characterised by a number of plant species with a low constancy which are exclusive to this subcommunity. these include the forbs calpurnia glabrata, pelargonium sidoides and the dwarf shrub rhus discolor. three tree species, namely leucosidea sericea, buddleja salviifolia and gymnosporia buxifolia were recorded in this sub-community with a low cover-abundance and constancy. none of these tree species exceeded 2 m in height and therefore the woody layer is represented only by shrubs lower than 2 m. the malanskraal habitat of chrysoritis aureus is also the exclusive habitat to this sub-community. issn 0075-6458 79 koedoe 46/1 (2003) fig. 5. dca-ordination of the sample plots at the alice glockner nature reserve (ac), malanskraal (mk) and suikerbosrand nature reserve (sbr). terblance 2.qxd 2005/12/09 10:31 page 79 koedoe 46/1 (2003) 80 issn 0075-6458 table 3 a phytosociological table of the surveys done at the alice glockner nature reserve, malanskraal and suikerbosrand terblance 2.qxd 2005/12/09 10:31 page 80 issn 0075-6458 81 koedoe 46/1 (2003) table 3 (continued) terblance 2.qxd 2005/12/09 10:31 page 81 koedoe 46/1 (2003) 82 issn 0075-6458 table 3 (continued) terblance 2.qxd 2005/12/09 10:31 page 82 issn 0075-6458 83 koedoe 46/1 (2003) table 3 (continued) 1.2. melinis nerviglumis-aristida transvaalensis-loudetia simplex sub-community this sub-community was recorded on the quartzite slopes of alice glockner and suikerbosrand on the witwatersrand supergroup (table 1). the diagnostic species include the grasses loudetia simplex, diheteropogon amplectens, tristachya leucothrix and the forbs vernonia staehelinoides and senecio venosus. (species group c, table 3). it is interesting to note that the forbs vernonia staehelinoides and senecio venosus (species group c, table 3) are absent from the sites of all the other communities, but are found scattered throughout this community. this community can be divided into two variants: 1.2.1 anthospermum hispidulum variant this variant is recorded from the southfacing rocky slopes of the alice glockner nature reserve and the suikerbosrand nature reserve. the constant presence of anthospermum hispidulum characterises this variant while clutia pulchella occurs abundantly on most of the sites that were surveyed in this community. the bracken fern pteridium aquilinum was only found at the suikerterblance 2.qxd 2005/12/09 10:31 page 83 koedoe 46/1 (2003) 84 issn 0075-6458 bosrand habitat (table 3) which at first glance gave the suikerbosrand habitat a different appearance. the similarities with the habitats at alice glockner nature reserve were only revealed after the vegetation classification. almost no tree species were found at all these sites. the woody layer is represented by canthium gilfillanii, myrsine africana and diospyros lyciodes subsp. guerkei (table 3). these woody species occurred sporadically, had low cover-abundance values, and none were taller than 1 m. chrysoritis aureus occurs on most of these sites, excluding sites ac 10 and ac 11 that are present on the lower midslopes (fig..4). of interest is that chrysoritis lycegenes, a closely related species of chrysoritis aureus, uses myrsine africana as host plant (henning 1983). chrysoritis lycegenes, however, is mainly restricted to central kwa-zulu natal and the eastern free state. 1.2.2 digitaria monodactyla variant this variant was observed on two flat terraces on the slopes at alice glockner nature reserve (fig. 4). the diagnostic species include the grasses aristida aequiglumis, digitaria monodatyla and schizachyrium sanguineum (table 3, species group e). the terraces are narrow and form a mini-plateau, whereas the huge exposed rocks that occur elsewhere on the slopes are absent. chrysoritis aureus does not colonise these flat terraces, and this coincides with the absence of the host plant clutia pulchella (table 2, table 3). 1.3 melinis nerviglumis-aristida transvaalensis-englerophytum magalismontanum sub-community the summit of the ridge at the alice glockner nature reserve is the habitat of this subcommunity which is dominated by shrubs rather than grasses and forbs. a small patch of vegetation, measuring only approximately 10 m x 10 m, which differs physiognomically from the surrounding areas, is present. the diagnostic species is the shrub englerophytum magalismontanum. other dominant shrubs present are myrsine africana, ancylobotrys capensis and rhus magalismontana. no chrysoritis aureus occur at this site since the habitat is not suitable. a number of other butterflies namely coeliades forestan, junonia archesia, junonia oenone, cigaritis natalensis and anthene butleri subsp. livida showed typical hilltopping behaviour at this site. hilltopping indicates a type of territorial behaviour where males of butterfly species gather at tops of hills, often at midday—individuals will choose a site which they will defend by flying after “intruders”. no hilltopping of chrysoritis aureus was observed. 2. elionurus muticus – eragrostis trichophora community. this community is found in the valley bottom on the southern hillside below the rocky ridges at the alice glockner nature reserve. it is characterised by the diagnostic species namely the grasses elionurus muticus, eragrostis trichophora, digitaria tricholaenoides and heteropogon contortus as well as the forbs indigofera comosa and vernonia oligocephala (species group g, table 3). very little exposed rocks occur in this area and the plant species cover abundance is relatively higher than in all the other sample plots. chrysoritis aureus is absent in this community. dca-ordination the results of the dca-ordination (fig. 4) support the classification of the communities identified from the phytosociological table. the difference between the floristic composition of the habitats of chrysoritis aureus at malanskraal and those at suikerbosrand nature reserve and alice glockner nature reserve, is confirmed by the results of the dca ordination – a topographical gradient from the valley bottom to the upper slopes are reflected by the configuration of the ordination. the position of the sample plots on ordination axis two can be related to the geology, with habitats on the ventersdorp lava supergroup at the top and habitats on terblance 2.qxd 2005/12/09 10:31 page 84 issn 0075-6458 85 koedoe 46/1 (2003) the witwatersrand supergroup in the middle and at the bottom. the discontinuous position of ac 14 on the scatter diagram may partly be due to its location on the summit of a ridge in contrast with all the other sample plots that are on south-facing slopes. the elionurus muticus-eragrostis trichophora community is at the left extreme along ordination axis 1 of the scatter diagram and the melinis nerviglumis-aristida transvaalensis community (community 1.1., 1.2 and 1.3) to the middle and right of the scatter diagram. the melinis nerviglumis-aristida transvaalensis-eragrostis curvula subcommunity (community 2) that occurs at malanskraal is situated at the top right end of the dca scatter diagram and the melinis nerviglumis-aristida transvaalensis-loudetia simplex sub-community in the middle. most of the sample plots, of which the plant species composition, geology and topography favour the presence of chrysoritis aureus, are to the middle-right and top end of the scatter diagram. this confirms the importance of open vegetation on south-facing rocky upper slopes for the survival of the butterfly. discussion the habitat of chrysoritis aureus table 4 provides a matrix in which some of the important environmental habitat characteristics that became apparent in this study, are listed. the altitude at the investigated habitats varies between 1690 m–1780 m, table 4 a table with specified habitat factors/ observations that apply to a relevé (= 1) or not (= 0). if a number 1 is in bold it indicates that the presence of chrysoritis aureus coincides with the factor. the presence of chrysoritis aureus is indicated in the first row of the matrix terblance 2.qxd 2005/12/09 10:31 page 85 koedoe 46/1 (2003) 86 issn 0075-6458 indicating that the butterfly only occurs in a certain climate regime (table 2). the butterfly colonies are only found at locations that are covered with large rocks. these rocks seem to be a habitat requirement for the host plant. in addition, steep upper midslopes seem to be essential even on the most local scale (the butterfly does not breed on the terraces). only the cool southern-facing slopes are chosen by the colonies of the butterflies for reasons which are not yet clear. this small-scale study showed that chrysoritis aureus is indeed specific on a very local scale with regard to its choice of habitat. two related sub-communities with different floristic compositions, but similar vegetation structures were recorded as habitats of chrysoritis aureus. the habitats of chrysoritis aureus are dominated by grass species though a high forb diversity was observed. many of these forbs are perennials that can survive fire. these include clutia pulchella (the host plant) as well as nectar plants for the adult butterfly, such as tetraselago wilmsii (species-group d; table 3). despite dense stands of trees or shrubs observed to be growing on the same hill of each of the localities studied, the tree component was almost absent where chrysoritis aureus is present. since dense and relatively diverse communities of trees and shrubs are particular to the rocky highveld grassland (bredenkamp & van rooyen 1996)—compared to other types of grassland—it is believed that the more open vegetation noted at the habitats of chrysoritis aureus are important for conservation management. fire and frost are two factors that probably play a key role in the maintenance of a suitable habitat for chrysoritis aureus since both are needed to maintain the grasslands in these rocky highveld grassland areas. acocks (1988) interpreted bankenveld (now included in the rocky highveld grassland) as a fire-maintained false grassland which would develop into savanna if fire was excluded. the vegetation of the suikerbosrand nature reserve that was studied by bredenkamp & theron (1978, 1980) provides a further context within which the vegetation studies at the chrysoritis aureus habitat at suikerbosrand could be seen. bredenkamp & theron (1978) described a protea caffra–helichrysum setosum savanna community that is also found on rocky steep southern slopes of the witwatersrand supergroup in the suikerbosrand nature reserve. a protea caffra-clutia pulchella variant was also distinguished by bredenkamp & theron (1978). an open tree stratum was observed in this variant consisting of a number of trees of which protea caffra was the most conspicuous (bredenkamp & theron 1978). no chrysoritis aureus were found in the past (including surveys by members of the lepidopterists’ society) in the vegetation where protea caffra individuals are an important component of the vegetation. the absence of protea caffra and any tree layer at all the sites where chrysoritis aureus occur is therefore a confirmation of the more open nature of the vegetation in the habitats of the butterfly. these open patches in the locations where clutia pulchella occur and which are concomitant with an absence of protea caffra seems to be localised, and perhaps marginal. a larger study area should reveal more information about the significance of the absence of a tree stratum at the sites where chrysoritis aureus occurs. fire may play an important role in the regulation of the chrysoritis aureus habitats. the timing and intensity of fires in the vegetation among the rocky ridges where clutia pulchella grow should be continually monitored to promote the survival of chrysoritis aureus. frost is also mentioned in literature to play an important role in the distribution of the woody elements in the rocky highveld grassland, as mentioned by bredenkamp & van rooyen (1996). grassland vegetation in the rocky highveld grassland is restricted to exposed cooler sites (bredenkamp & van rooyen 1996). all the habitats of chrysoritis aureus are at exposed sites below the summit of these hills (on the upper slopes). chrysoritis aureus have not been found in any of the kloofs in these areas where large numbers of clutia pulchella were observed, though unfortunately no sample plots were placed there for quantitative comparisons. terblance 2.qxd 2005/12/09 10:31 page 86 issn 0075-6458 87 koedoe 46/1 (2003) since chrysoritis aureus is also associated with an ant species, crematogaster liengmei, the structure of the vegetation and the role of factors such as fire may be part of the specific habitat requirements of the ants. the present ant community studies may reveal more insight into these habitat requirements, especially against the background of the vegetation analysis provided in this study. the open nature of the vegetation at the habitats of chrysoritis aureus has further implications for the control of invasive plants in these areas. cover-abundances of the pteridium aquilinum (bracken fern) stands at the chrysoritis aureus habitat at the suikerbosrand nature reserve should be monitored, since the bracken fern can be regarded as invasive. this habitat occurs at the border fence of the reserve and is regularly burnt as part of a fire break. the best method to control bracken, if deemed necessary, is by cutting (marrs et al. 1998). bracken ferns possess a number of underground rhizomes and are well adapted to fire. oinonen (1967) found that bracken established best from spores that are present on sites that had been burnt. another consideration for the management at all the sites is to prevent bush encroachment and the establishment of any exotic tree species, since this will clearly destroy the localised habitats. continued control of the invasive acacia mearnsii, which is a declared invader (with commercial value) (bromilow 2001; henderson 2001) at the alice glockner nature reserve, is imperative since this will save the butterfly habitats from a possible catastrophe in future. conclusion the vegetation at the three localities where chrysoritis aureus is present, is similar with regard to general structure, but there are differences in species composition on the subcommunity level. a remarkable coincidence in the malanskraal habitats, being a different sub-community, is that the clutia pulchella individuals collected there also differ markedly from individuals from the localities at alice glockner nature reserve and the suikerbosrand nature reserve. the malanskraal habitat is also on the ventersdorp supergroup in contrast with the habitats at the alice glockner nature reserve and suikerbosrand nature reserve. an hypothesis to be tested is that the genetic make-up of the chrysoritis aureus populations at the three localities are different, since they seem to be adapted to habitats that differ to some extent. these findings would be valuable for the conservation of chrysoritis aureus if translocation of the butterfly or restoration of habitats are considered. securing corridors that link habitats is an aspect that deserves attention, especially in the sense that these must include adequate nectar sources. the value of a phytosociological survey to support conservation research and management strategies for a threatened butterfly are confirmed by this study and by deutschländer & bredenkamp (1999). the vegetation structure would be an important aspect to consider in the management of areas where chrysoritis aureus occurs and could even be studied in more detail on a larger scale. due to the open nature of the vegetation and localised distribution of the habitats of chrysoritis aureus the application of firemanagement should be applied with care while bush encroachment, especially of any tree layer, should be avoided. alien vegetation in these areas should also be monitored since the establishment of exotic tree species will destroy the habitats of chrysoritis aureus. further research priorities that were identified, include a study of ant communities locally at the habitats of chrysoritis aureus. to study the nectar sources of chrysoritis aureus in more detail a larger study area than the one used here, would be needed since it seems that the butterfly occasionally wanders of from its central breeding ground in search of nectar. larger sample plots (100 m²) and a larger study area should therefore be included for such studies. for the studies on the optimal density of the host terblance 2.qxd 2005/12/09 10:31 page 87 koedoe 46/1 (2003) 88 issn 0075-6458 plant, aspects other than southern aspects, should also be included. it is hoped that more and diverse research on the rare and localised butterflies of africa can be continued since it is believed that a holistic approach in the research will lead to more discoveries with regard to the intrigued habitat requirements of these organisms. it is recommended that the various sites where chrysoritis aureus occurs, should receive priority for conservation purposes. in the light of the variation in the habitats of chrysoritis aureus encountered during this study, the securing of one habitat for the conservation of a local habitat-specific butterfly might be a risk. the implications for butterflies such as orachrysops niobe (brenton blue), where only one locality is known, is that such butterflies might be in a precarious situation even if the remaining habitat is conserved, since the meta-population dynamics may be limited. the threat of urbanisation to the conservation of the rocky highveld grassland in general is mentioned by bredenkamp & van rooyen (1996) and it seems to be prophetic regarding the conservation of chrysoritis aureus. rocky highveld grassland is poorly conserved (bredenkamp & van rooyen 1996) and if the number and extent of conservation areas are considered the development and conservation of the alice glockner nature reserve will prove to be an important asset for biodiversity conservation and ecotourism in south africa. acknowledgements we wish to thank henning & roos cc as well as michelle pfab and marianne forsyth (gauteng department of agriculture, conservation and environment) for their professional support and assistance. this research would not have been possible if it had not been for the financial support of the gauteng department of agriculture, conservation and environment, via roos & henning cc. references acocks, j.p.h. 1988. veld types of south africa. memoirs of the botanical survey of south africa no. 57. arnold, t.h. & b.c. de wet (eds). 1993. plants of southern africa: names and distribution. memoirs of the botanical survey of south africa no. 62. bezuidenhout, h., h.c. biggs & g.j. bredenkamp. 1996. a process supported by the utility bbpc for analysing braun-blanquet data on a personal computor. koedoe 39(1): 107-112. bredenkamp, g.j. & g.k. theron. 1978. a synecological account of the suikerbosrand nature reserve i. the phytosociology of the witwatersrand geological system. bothalia 12(3): 513529. bredenkamp, g.j. & g.k. theron. 1980. a synecological account of the suikerbosrand nature reserve ii. the phytosociology of the ventersdorp geological system. bothalia 13(1&2): 199-216. bredenkamp, g.j. & n. van rooyen. 1996. rocky highveld grassland. in: low, a.b. & a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. bromilow, c. 2001. problem plants of south africa. pretoria: briza. cottrell, c.b. 1984. aphytophagy in butterflies: its relationships to myrmecophily. zoological journal of the linnean society 79: 1-57. deutschländer, m.s. & c.j. bredenkamp. 1999. importance of vegetation analysis in the conservation management of the endangered butterfly aloeides dentatis subsp. dentatis (swierstra) (lepidoptera: lycaenidae). koedoe 42(2): 1-12. de wet, k. 1987. die bewaring van die skoenlappers van die genus charaxes en van erikssonia acraeina in die transvaal. proceedings of the first lepidoptera conservation symposium, roodepoort: lepidopterists' society of southern africa: 5-7. du toit, a.c. 1954. the geology of south africa edinburgh: oliver & boyd. edwards, d. 1983. a broad-scale structural classification of vegetation for practical purposes. bothalia 14(3): 705-712. heath, a. 1997. a review of african genera of the tribe aphnaeini (lepidoptera: lycaenidae). metamorphosis: occasional supplement no. 2: 1-60. heath, a. 2001. new synonymies and taxonomic notes on the genus chrysoritis butler (lepidoptera: lycaenidae). metamorphosis 12 (3): 85-98. henderson, l. 2001. alien weeds and invasive plants: a complete guide to declared weeds and invaders in south africa. cape town: agriculterblance 2.qxd 2005/12/09 10:31 page 88 issn 0075-6458 89 koedoe 46/1 (2003) tural research council. (plant protection research institute handbook no. 12.) henning, g.a. 1986a. research at suikerbosrand nature reserve, 7 september 1986. transvaal nature conservation. unpublished report. henning, g.a. 1986b. research at suikerbosrand nature reserve, 2 december 1986. transvaal nature conservation. unpublished report henning, g.a. 1987. research at suikerbosrand nature reserve, 1st quarter 1987. transvaal nature conservation. unpublished report. henning, g.a. & s.f. henning. 1995. updating the status of south african red data butterfly species. metamorphosis 6(2): 96-98. henning, s.f. 1983. biological groups within the lycaenidae (lepidoptera). journal of the entomological society of southern africa 46(1): 6585. henning, s.f. 1987a. myrmecophilous lycaenidae (or how ants help butterflies). south african journal of science 83: 8-9. henning, s.f. 1987b. outline of lepidoptera conservation with special reference to ant associated lycaenidae. proceedings of the first lepidoptera conservation symposium, roodepoort: lepidopterists' society of southern africa: 5-7. henning, s.f. & g.a. henning. 1989. south african red data book: butterflies. pretoria: council for scientific and industrial research, national scientific programmes unit. (south african national scientific programmes report; no. 158.) hill, m.o. 1979a. twinspan. a fortran programme for arranging multivariate data in an ordered two way table by classification of individuals and attributes. ithaca, new york: cornell university. hill, m.o. 1979b. decorana. a fortran programme for detrended correspondence analysis and reciprocal averaging. ithaca, new york: cornell university. kudrna, o. 1995. conservation of butterflies in central europe. pp. 248 257. in: pullin, a.s. (ed.). ecology and conservation of butterflies. london: chapman & hall. larsen, t.b. 1995. butterfly biodiversity and conservation in the afrotropical region. pp. 290303. in: pullin, a.s. (ed.). ecology and conservation of butterflies. london: chapman & hall. low, a.b. & a.g. rebelo. (eds.). 1996. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. marrs, r.h., s.w. johnson & m.g. le duc. 1998. control of bracken and restoration of heathland vii. the response of bracken rhizomes to 18 years of continued bracken control or 6 years of control followed by recovery. journal of applied ecology 35: 748-757. mueller-dombois, d. & h. ellenberg. 1974. aims and methods of vegetation ecology. new york: wiley. munguira, m.l. 1995. conservation of butterfy habitats and diversity in european mediterranean countries. pp. 277 289. in pullin, a.s. (ed.). ecology and conservation of butterflies. london: chapman & hall. new, t.r. 1995. butterfly conservation in australasia an emerging awareness and an increasing need. pp. 304 315. in pullin, a.s. (ed.). ecology and conservation of butterflies. london: chapman & hall. oates, m.r. 1995. butterfly conservation within the management of grassland habitats. pp. 98-112. in: pullin, a.s. (ed.). ecology and conservation of butterflies. london: chapman & hall. o'dwyer, c. & p.m. attiwill. 2000. restoration of a native grassland as habitat for the golden sun moth synemon plana walker (lepidoptera: castniidae) at mount piper, australia. restoration ecology 8 (2):170-174. oinonen, e. 1967. sporal regeneration of bracken (pteridium aquilinum) (l. kuhn.) in finland in the light of the dimensions and age of its clones. acta forestalia fennica 83(2): 1-51. opler, p.a. 1995. conservation and management of butterfly diversity in north america. pp. 316 324. in: pullin, a.s. (ed.). ecology and conservation of butterflies. london: chapman & hall. owen-johnston, n.k. 1991. thoughts and observations on the genus poecilmitis butler (lepidoptera: lycaenidae) in the transvaal. metamorphosis 2(3): 49-50. prain, d. 1925. cluytia. pp. 427-456 in: thiseltondyer, w.t. (ed.). flora capensis: part iii. london: reeve. pringle, e.l., g.a. henning & j.b. ball (eds.). 1994. pennington's butterflies of southern africa. kaapstad: struik winchester. pullin, a.s. ed. ecology and conservation of butterflies. london: chapman & hall. pullin, a.s., i.f.g. mclean & m.r. webb. 1995. ecology and conservation of lycaena dispar: british and european perspectives. pp. 150-164. in: pullin, a.s. (ed.). ecology and conservation of butterflies. london: chapman & hall. retief, e. & p.p.j. herman. 1997. plants of the northern provinces of south africa: keys and diagnostic characters. pretoria: national botanical institute. roos, p. & g.a. henning. 2000. the heidelberg copper butterfly. african wildlife 54(2): 24 -27. rutherford, m.c. & r.h. westfall. 1994. biomes of southern africa: an objective categorization. memoirs of the botanical survey of south africa no. 63. terblance 2.qxd 2005/12/09 10:31 page 89 thomas, c.d. 1995. ecology and conservation of butterfly metapopulations in the fragmented british landscape. pp. 46-64. in: pullin, a.s. (ed.). ecology and conservation of butterflies. london: chapman & hall. van oudtshoorn, f.p., w.s.w., trollope, d.m. scotney & p.j. mcphee. 1991. guide to the grasses of south africa. pretoria: briza. von breitenbach, f. 1990. national list of indigenous trees. pretoria: dendrological foundation. von breitenbach, j., b. de winter, r. poynton, e. van der berg, b. van wyk & e. van wyk. 2001. pocket list of southern african indigenous trees: including selected shrubs and woody climbers. pretoria: briza. warren, m.s. 1995. managing local microclimates for the high brown fritillary, argynnis adipe. pp. 198–210. in: pullin, a.s. (ed.). ecology and conservation of butterflies. london: chapman & hall. koedoe 46/1 (2003) 90 issn 0075-6458 terblance 2.qxd 2005/12/09 10:31 page 90 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <feff004e00e4006900640065006e002000610073006500740075007300740065006e0020006100760075006c006c006100200076006f0069006400610061006e0020006c0075006f006400610020005000440046002d0061007300690061006b00690072006a006f006a0061002c0020006a006f006900640065006e002000740075006c006f0073007400750073006c00610061007400750020006f006e0020006b006f0072006b006500610020006a00610020006b007500760061006e0020007400610072006b006b007500750073002000730075007500720069002e0020005000440046002d0061007300690061006b00690072006a0061007400200076006f0069006400610061006e0020006100760061007400610020004100630072006f006200610074002d0020006a0061002000520065006100640065007200200035002e00300020002d006f0068006a0065006c006d0061006c006c0061002000740061006900200075007500640065006d006d0061006c006c0061002000760065007200730069006f006c006c0061002e0020004e00e4006d00e4002000610073006500740075006b0073006500740020006500640065006c006c00790074007400e4007600e4007400200066006f006e0074007400690065006e002000750070006f00740075007300740061002e> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 matipano ranging.qxd post-release ranging behaviour of hand-raised black rhinoceros, diceros bicornis, l. in matusadona national park, zimbabwe with recommendations for management of introduction to the wild g. matipano matipano, g. 2004. post-release ranging behaviour of hand-raised black rhinoceros, diceros bicornis, l. in matusadona national park, zimbabwe with recommendations for management of introduction to the wild. koedoe 47(1): 89–101. pretoria. issn 0075-6458. hand-raised rhinos would preferentially settle at sites close to areas where human activity was significant. hand-raised rhinos that attached themselves to sites of human activity tended to move less widely and to have smaller seasonal ranges than those that moved away from human settlements. these rhinos had no fear of humans and some animals would move along established roads into hostile areas outside the park. these factors made hand-raised rhinos more vulnerable to poaching than their wild counterparts. management of hand-raised rhinos with regards to ranging behaviour is discussed in the text. key words: black rhino, hand-raised, ranging behaviour, reintroduction. g. matipano, matusadona national park, parks and wildlife management authority, private bag 2003, kariba, zimbabwe. present address: national university of science and technology, department of forest resources and wildlife management, p. o. box ac 939, ascot, bulawayo, zimbabwe (gmatipano@nust.ac.zw). issn 0075-6458 89 koedoe 47/1 (2004) introduction reintroduction is an important wildlife management technique for conservation of endangered species including the black rhinoceros. the first months after release in a new area are naturally a stressful period for introduced rhino (adcock et al. 1998). wild black rhino are known to be a relatively poor dispersing species after introduction to a new area. reintroduction of black rhino met with problems, including poor habitat quality and hence poor reproductive performance (emslie 1994). other problems are related to mass release of animals at one site at the same time and the resultant mortality due to intra-specific fights among introduced animals (emslie 1994) and intra-specific aggression from resident individuals (brett 1998). this paper focused on seasonal ranges rather than home ranges. this was done because some of the hand-raised rhinos were too young to hold home ranges. others, although they were adults on release, had not had enough time to established home ranges by the time of the study. home range is that area traversed by an individual animal in its normal activities of food gathering, mating and caring for young excluding the area outside its normal area (burt 1943). many factors influence the size of home ranges of black rhino. home range areas vary with an animal’s requirements: ranges tend to be larger when food, water and cover are scarce or scattered (mukinya 1973). black rhinos have larger home ranges in woodlands than in thickets (hitchins 1969). home ranges tend to increase in area with decreasing browse availability. the social status (group member, loner, resident, transient), age, sex and reproductive condition may influence range sizes (bekoff & mech 1984). generally, adult male rhinos tend to have smaller home ranges than adult females (owensmith 1988), but in some cases they are equal (mukinya 1973). ranges of immature black rhinos and females with calves are larger than those of single adult males or females (joubert & eloff 1971). for cows, matipano ranging.qxd 2004/04/14 02:03 page 89 this difference may be due to the need to meet food requirements and to acquaint the calf with the home range (mukinya 1973). typical black rhino mean home range sizes are 7–35 km² (owen-smith 1988). animals do not use home ranges in a uniform manner. core areas denote space of high home range usage. an animal can have one or more core areas in its home range (dixon & chapman 1980). core areas may provide a more clear measure of the changing pattern of range use than home range area (harris et al. 1990). in general, core areas of conspecific animals are mutually exclusive (harris et al. 1990) but can overlap for rhinos, specifically (tatman et al. 2000). ranging behaviour studies of black rhinos have been carried out on wild free-ranging populations (e.g. goddard 1967; mukinya 1973; frame 1980; kiwia 1989), and on relocated or reintroduced populations (e.g. hitchins 1969; adcock et al. 1998; tatman et al. 2000). however, information on the ranging behaviour of hand-raised and released black rhinos is scarce and is not directly related to matusadona national park (mnp). objectives of the study are to determine effects of hand-raising on the ranging behaviour of black rhinos, and secondly to contribute information towards the development of guidelines for sound reintroduction process and post-release management of hand-raised black rhino. methods study area matusadona national park (mnp) stretches from 28º23'e–28º51'e and from 16º41's–17º13's. the park is about 1407 km² in area. an escarpment divides the park into two major geomorphologic landscapes, the semi-arid eutrophic valley floor lying between 485 m and 600 m above sea level (a.s.l.) and the wet dystrophic highland section lying between 600 m and 1200 m a.s.l. the lowland area is dominated by semi-arid vegetation mainly colophospermum mopane (kirk ex benth.) kirk ex j. léonard woodland while the highland area supports brachystegia benth.-julbernadia pellegr. woodlands. the study was restricted to the valley floor section of the park. mnp has a semi-arid climate in the valley floor where rainfall ranges between 400 and 800 mm per year with an average mean annual of 729.5 mm. study rhinos no rhinos were collared specifically for this project. two adult female rhinos were collared in 1998. four hand-raised rhinos (two adults: male and female, and two subadults: male and female) also had been radiocollared before the research project started. sex, origin, duration stayed in bomas, season of release, age on release and age during the time of study of the hand-raised rhinos are shown in table 1. the ages of rhinos during the time of the study ranged between 4.5 and 13 years. these animals had been kept in bomas for 4-12 years before final release. handraised animals were first released into mnp in march 1998. during the study the park was supporting about 55 black rhinos (wild, hand-raised and released and those that were still under hand-raising). collared animals were radio-tracked by telemetry and their positions were recorded using a gps between march 1998 and july 2000. routine monitoring data (department of national parks and wildlife management files) on two wild adult female rhinos were used to calculate seasonal range sizes for 1998. during fieldwork, the target was to locate each collared animal at least once a week, using a telonics model tr-2 receiver. radio-collared animals were initially approached by vehicle, and final contact was made on foot to avoid disturbance. an animal’s position on first sighting was recorded using a gps garmin 40. data collection was limited to diurnal observations. information was collated by season, as follows: wet season (december-march), early dry season (may-july) and late dry season (septemberearly november). calculation of seasonal range sizes the vegetation map of mnp compiled by taylor (1985) was scanned and digitised into the tntmips gis software programme operated by the wwf multispecies project. positions of individual rhinos were plotted on the vegetation map. the 100 % minimum convex polygon (mohr 1947) was used to draw seasonal ranges and to calculate their sizes. in this study, seasonal range was investigated in the wet, early dry and late dry seasons. the 100 % mcp method was chosen for its ease of use. radio-locations were assumed to be non-autokoedoe 47/1 (2004) 90 issn 0075-6458 matipano ranging.qxd 2004/04/14 02:03 page 90 correlated because they were collected at least daily for a given rhino. auto-correlation refers to two or more fixes that are not independent in terms of time. the mcp method, is not affected by temporal autocorrelation of fixes (harris et al. 1990; dexter 1999). this study only required an index of seasonal range, and not absolute sizes. the mcp is the smallest convex polygon containing all the observed positions; the area within this polygon is the estimated seasonal range size (anderson 1982). landscapes not known to be used by rhinos, such as water bodies, were excluded from the seasonal range area. calculation of core area sizes the 70 % harmonic method (dixon & chapman 1980) was used to calculate core areas for individual seasonal ranges. the relationships between seasonal range and core area sizes for both rhino groups were graphically presented. statistical treatments the sample size was small and data were therefore pooled for analysis. the chi-square tests were used to analyse the differences between mean seasonal range and core area sizes for hand-raised and wild rhinos. the same tests were used to analyse the within-group variation in seasonal range and core area sizes. the regression analysis was used to determine the relationship between seasonal range and core area sizes. results movements of hand-raised rhinos no rhino settled at its release site. the longest period of monitoring (march 1998 – july 2000) by radio tracking was for chewore, a subadult female rhino. the animal was immobilised, translocated and released at mhukadzapera river (fig. 1) on the same day in march, 1998. it first left the park in november 1999, into the nyaminyami district communal lands, about two years after release (fig. 2). the animal was recaptured and released at nyamoni river. thereafter, chewore repeatedly moved out (once in each of the months january, april, may, june, and july 2000), not only out of the study area itself but also out of the park. the longest distance moved from the park was about 40 km, and from the study area was about 70 km. each time the animal moved out of the park, it walked along established roads. in communal lands, the animal tended to reduce movements. the animal used the same general route going out of the park and passing close to or through a township. at one time the rhino was put in an enclosure at a growth point township outside the park, awaiting collection by national parks. every time it was out of the park, the animal was walked back into the conservation area. this habit of hand-raised rhinos, of following roads, was also observed in the midlands area of zimbabwe (pers. obs.). chewore was relocated to sinamatella (hwange national park) in january 2002. soon after arrival it crossed to botswana from where it was later taken back. cuckoo, an adult female, was released by walking it out of the gubu bomas (fig. 3) in the early dry season, june 1998. the animal moved widely soon after release and in the issn 0075-6458 91 koedoe 47/1 (2004) table 1 history of hand-raised rhino in mnp name of sex source duration age on season of date rhino in boma release final of release (years) (years) release pfumbe male mana pools 12 12 early dry april 1999 via imire cuckoo female mana pools 11 11 dry june 1998 via imire chewore female chewore 5 5 wet march 1998 mgofu male imire 4 4 wet dec 1999 matipano ranging.qxd 2004/04/14 02:03 page 91 koedoe 47/1 (2004) 92 issn 0075-6458 fig. 2. map showing movements of hand-raised rhino outside matusadona national park. fig. 1. map of matusadona national park (mnp) showing seasonal ranges of a hand-raised rhino, ‘chewore’. 1. first release site (mhukadzapera river) 2. second release site (nyamoni river). matipano ranging.qxd 2004/04/14 02:04 page 92 subsequent late dry season. in the 1998/99 wet season, the animal then stayed briefly close to muuyu and changachirere camps on the lakeshore (where there was marked tourist activity). later, she went up the escarpment, in the following early dry season, leaving the study area (fig. 2). an introduced adult male, pfumbe, was released by walking it out from gubu bomas in june 1998 (fig. 4). the animal stayed briefly in the park and then crossed the park boundary into the nyaminyami and gokwe north district communal lands (fig. 2). in communal lands the animal did not to move widely and could visit homesteads. once the animal broke window panes at a villager’s homestead. after staying for about two weeks in the communal lands, pfumbe was recaptured and placed in the boma at tashinga camp in july 1998. the animal was released again in january 1999 from the tashinga bomas by letting it walk out on its own. after engaging in two serious fights with a wild resident bull, pfumbe returned into the boma of its own accord. pfumbe was immobilised and transported and finally released at lower jenje river, in april 1999 (fig. 4). by may 2003, the animal seemed to issn 0075-6458 93 koedoe 47/1 (2004) fig. 3. map of mnp indicating seasonal ranges of a hand raised rhino,’cuckoo’. 1 = release site. fig. 4. map of mnp indicating seasonal ranges of a hand-raised rhino, ‘pfumbe’. 1-3 = release sites 1-3. fig. 5. map of mnp indicating seasonal ranges of a hand-raised rhino, ‘mgofu’. 1 = release site. have settled near changachirere camp where it settled since 2000. mgofu was walked to a satellite boma at upper jenje river (site 1), about 18 km from the raising boma at tashinga. it stayed there for about one and half months before final release into the wild in december 1999. the animal moved for about 12 km to a headland in the lake kariba, where muuyu camp is matipano ranging.qxd 2004/04/14 02:04 page 93 koedoe 47/1 (2004) 94 issn 0075-6458 fig. 6. map of mnp indicating seasonal ranges of a female wild rhino number 7. fig. 7. map of mnp indicating seasonal ranges of a wild female rhino number 21. table 2 seasonal ranges and core areas of black rhinos in mnp year and rhino rhino no. of seasonal bmean core mean core season group identity fixes range seasonal range area size area size (no./name) used size (km²) size (km²) (km²) (km²) 1998 wet wild 7 5 20.8 7.3 season 21 6 32.9 26.9 14.1 10.7 hand-raised chewore 6 0.8 0.8 0.2 0.2 1998 early wild 7 5 10.4 0.8 dry season 21 4 2.1 6.3 1.9 1.4 hand-raised chewore 8 2.3 2.3 cuckoo 4 32.6 11.2 pfumbe 4 4 13.0 1.2 4.9 1998 late wild 7 6 13.7 0.6 dry season 21 6 1.3 7.5 0.4 0.5 hand-raised chewore 8 15.6 6.4 cuckoo 7 58.7 37.2 55.5 31.0 1998/99 hand-raised chewore 4 7.3 0.6 wet season cuckoo 6 7.5 7.4 0.6 0.6 1999/2000 hand-raised chewore 8 22 1.5 wet season pfumbe 6 1.3 0.6 mgofu a 1.5 8.0 a 1.4 2000 early hand-raised chewore 6 9.8 1.5 dry season pfumbe 8 1.2 0.2 mgofu 7 1.2 4.1 1 0.9 a area was estimated based on the island the animal was occupying in that season b mean seasonal range sizes (column six) are not for individual rhinos but for a group of rhinos, i.e. wild and hand-raised rhino groups. matipano ranging.qxd 2004/04/14 02:04 page 94 situated (fig. 5). when the headland became an island, mgofu moved to the mainland in the early dry season. by may 2003, the animal remained in the same area it had settled in previously, in 1999. figures 6 & 7 show movements of two wild adult females during the time of the study. seasonal ranges’ sizes seasonal ranges of rhino groups are shown in table 2. the mean seasonal range areas were significantly different from each other between hand-raised and wild rhinos in the wet and late dry seasons (χ² > 19; p< 0.001) and not in the early dry season (χ² =2.350; 0.1<p<0.25, table 3) in 1998. in the wet season the mean seasonal range area for handraised rhinos was smaller than that for wild rhinos and the situation was vice versa in the dry seasons in 1998. between 1999 and 2000, the mean seasonal range areas for hand-raised rhinos varied with season and ranged between 4.1 km² and 8.0 km². the within-group seasonal range variation in size was significant for hand-raised rhino (χ² >5; p<0.025), except between chewore and pfumbe in the 1998 early dry season, and pfumbe and mgofu in the 2000 wet and early dry seasons (χ² <2.727; p>0.1; table 3). pfumbe and mgofu had the smallest seasonal ranges, as these animals had settled close to human settlements. seasonal ranges of hand-raised rhinos that settled close to human activity were both smaller than those of hand-raised rhinos that settled further away from man, and than those of their wild counterparts (table 2). seasonal ranges of rhinos were variable in terms of spatial location with season (figs. 1 & 3–7). there was some overlap in seasonal ranges of a given individual animal except pfumbe. core area sizes the mean core areas for both hand-raised and two wild rhinos are shown in table 2. issn 0075-6458 95 koedoe 47/1 (2004) table 3 chi-square tests on the differences in seasonal range and core area sizes within hand-raised and wild rhino groups year and season rhino group rhino name or significant (s) or core area number not significant (ns) seasonal range 1998 wet season wild 7 x 21 ns ns wild 7 x 21 s ns 1998 early dry season hand-raised chewore x cuckoo s s chewore x pfumbe ns ns cuckoo x pfumbe s s 1998 late dry season wild 7 x 21 s ns hand-raised chewore x cuckoo s s 1998-99 wet season chewore x cuckoo s ns chewore x pfumbe s ns 99/2000 wet season hand-raised chewore x mgofu s ns pfumbe x mgofu ns ns chewore x pfumbe s ns 2000 early dry season hand-raised chewore x mgofu s ns pfumbe x mgofu ns ns matipano ranging.qxd 2004/04/14 02:04 page 95 the mean core area sizes for hand-raised and wild rhinos were significantly different from each other in the wet and late dry seasons (χ² >10; p<0.05) and not significantly different in the early dry season (χ² =2.016; 0.1<p<0.25) in 1998. in the wet season, the mean core area for hand-raised rhinos was smaller than that for the wild group, and the situation was vice versa in the dry season. in 1998, the within-group variations in core area size of hand-raised rhino were not significantly different (χ² <2.166; p>0.1)— except between chewore and cuckoo and pfumbe and cuckoo in the early dry season, and chewore and cuckoo in the late dry season (χ² >8.066; p<0.05; table 3). cuckoo appeared to move widely and to be responsible for the major variations in seasonal ranges of hand-raised rhinos. the relationships between seasonal range and core area sizes the relationships between seasonal range and core area sizes are shown in fig. 8. the sizes of core areas increased significantly with the increasing seasonal range areas for both rhino groups (r² >0.67; p<0.03). the relationships between seasonal range and core area sizes were represented by the following equations for wild rhinos (i) and hand-raised rhinos (ii): seasonal range area = -1.35 + 0.394 x core area size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .(i) seasonal range area = -3.87 + 0.81 x core area size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .(ii) there was a threshold below which the decrease in seasonal range sizes did not result in significant decrease in core area sizes (r² =0.0; p>0.63). the threshold was lower for hand-raised than for wild rhino rhinos, i.e., 15.6 km² and 20.8 km², respectively. below the threshold, core area size remained below 2.4 km² for hand-raised rhinos, and below 2 km² for wild rhinos and they were not significantly different from each other (χ² =0.036, p> 0.75). discussion the study did not focus on the concept of home range, but on seasonal range, because of reasons discussed in the introduction and below. analysis of seasonal ranges was appropriate because short-term seasonal ranges are necessary for studying dispersing animals and when monitoring change in range use over time (harris et al. 1990). analyses at the level of the annual range under-utilise available data and do not show the within-year variations. thus, seasonal ranges can yield additional information compared to annual ranges. problems common to use of seasonal and annual ranges include that of distinguishing abnormal from normal movements of the animal. animal locations outside the study area were disregarded in the analysis. seasonal ranges have potential for additional errors compared to annual ranges. seasonal range shift is a series of continuous changes in location and time. it is represented by a series of overlapping ranges. that period of shifting from one seasonal range to another might not be identifiable on the ground. the researcher defined the time frame for seasonal ranges. the defined time frame for seasonal ranges might not be biologically meaningful to the species, i.e., the season of the range that has been determined by the researcher might not coincide with that for the animal. a trial run of the experiment should have been carried out to establish the time interval to collect data such that serial correlation of fixes does not occur. a test of independence of the fixes will reveal whether an animal is shifting its seasonal range or not. serially correlated subsets of data may indicate the spatio-temporal transitional phase between seasonal ranges (swihart & slade 1985). major components of the behavioural ecology of the species should be considered to establish whether the shift in ranges is attributable to season or not. it was difficult to find out whether handraised rhinos were moving within the seasonal ranges or were changing seasonal koedoe 47/1 (2004) 96 issn 0075-6458 matipano ranging.qxd 2004/04/14 02:04 page 96 ranges or moving about without defined seasonal ranges. seasonal ranges were highly variable both within and between groups of wild and hand-raised rhinos. the variation was in size and in location, according to season. wild rhinos less than eight years old are too young to hold home ranges (adcock 1994). half of the hand-raised rhinos, therefore, had not reached this critical age. in the wild, calves of both sexes sever relationship with their mothers at 2-4 years old and can move widely for up to another four years before establishing a home range (adcock 1994). in areas with a high density of adult bulls, young males remain with their mothers for protection, up to the age of eight years (adcock 1994). in kenya and south africa, wild captured rhinos (not hand-raised) had the highest mortality due to intra-specific fights within two years of release (brett 1998) and could take up to three years to settle in terms of home range sizes (adcock et al. 1998). it is, therefore, normal for rhinos to wander about soon after weaning or reintroduction into the wild. a matter of concern was the extent of movements by hand-raised rhinos, especially into potentially hostile areas, after reintroduction. in the wild, black rhino are relatively solitary and sedentary, confining their normal issn 0075-6458 97 koedoe 47/1 (2004) fig. 8. the relationship between black rhino core area and home range sizes. movements to their home ranges (owensmith 1988). movements of hand-raised rhinos were influenced by the presence of human settlements and, probably, by dominant wild bulls. seasonal ranges were smaller for hand-raised rhinos that stayed close to human settlements than those that were further away. the mean seasonal range areas for hand-raised rhinos decreased with time after release as the animals were prevented from further northward movement by lake kariba waters and settled near tourist camps. staying close to human settlements was a direct result of hand-raising on the behaviour of rhinos. rhinos released in the dry season moved widely soon after release compared to those that were released in the wet season. the wide movements might have been in response to low availability and poor interspersion of resources in the dry season compared to the wet season conditions. in reintroduction, animals should be released in the wet season or just after the rains when resources are still abundant. for various reasons, sample sizes were too small to be meaningful for both within and between wild-release group comparisons. the problem could not be resolved by pooling data because the within-group variance led to pseudoreplication. the small sample size therefore did not allow for meaningful conclusions on the roles of social status (resident or transient), sex-age status, reproductive condition and even season in determining home range size (bekoff & mech 1984). further, it was not possible to assess how the following variables affected seasonal range sizes: the period the animal had spent in the wild after release, the year or season of release, and the age of the rhino on release. an adult female rhino, cuckoo was responsible for the variation in core area within the hand-raised group. the number of fixes used for each rhino was not adequate for seasonal range analysis and could have resulted in bias by under-estimating range sizes. the mcp method is sensitive to sample size. home range sizes matipano ranging.qxd 2004/04/14 02:04 page 97 increase as the number of fixes increase towards the asymptote (bekoff & mech 1984), and the asymptotic level is reached when a further addition of fixes does not result in increase in the size of the home range. in this study (table 2), seasonal ranges for hand-raised rhino fell in the lower range of wild counter-parts in comparable semi-arid areas of africa—including masaimara in kenya (mukinya 1973) and hluhluwe in south africa (hitchins 1971) —probably due to small numbers of fixes that were used and fixes recorded outside the study area were not used in the analysis. the number of fixes used for seasonal ranges were not the same for all rhinos, which prevented comparison of range attributes. as long as the number of fixes achieve asymptote, attributes whose range have been obtained using unequal number of locations can be compared. many researchers (e.g. bekoff & mech 1984; robertson et al. 1998), recommended a total of 100-200 fixes for large mammals in home range analysis. in kenya, tatman et al. (2000) found that 20 to 30 fixes were adequate for home range in a small and confined black rhino population. these extremes in the number of required fixes arise from different situations, where the large figure is for free-ranging population and the smaller for those small fenced ranches. individual animals reach asymptotic number of fixes at different values, and some animals may not even reach an asymptote. for example, transient adults or dispersing subadult may not attain asymptote values (harris et al. 1990). an inappropriate interval in collecting data may also govern failure to achieve asymptote (harris et al. 1990). all hand-raised rhinos dispersed after release, which meant that the constantly changing localities did not always allow for asymptotic fixes. the 100 % mcp method, as an index to measure the ranging behaviour of rhinos, showed that hand-raised and wild rhinos were different from each other. the method includes portions of the study area that were not normally used by the animals. this results in overestimating the seasonal range size. the method, therefore, is biologically not suitable to study range sizes and habitat requirements unless it has been smoothed to correct for the increase in range size (white & garrott 1990; conybeare 1991). smoothing involves removal of outliers, i.e., those points which stand alone. the 70 % harmonic mean method removed some of the outliers. although smoothing drastically reduced the within-group variations, the mean core areas of hand-raised and wild rhinos became more significantly different from each other than their mean seasonal range areas. hand-raised rhinos had a lower threshold than wild animals, below which the decrease in seasonal range areas did not result in significant decrease in core area sizes. on the other hand, the size of the core area below the threshold point was larger for hand-raised rhinos than for wild rhinos, though not significantly different from each other. these statements seemed to indicate that hand-raised rhinos tended to wander about until they establish a seasonal range. findings of this study partially tended to agree with harris et al. (1990) that home ranges vary greatly during dispersal period of animals while core areas remain constant for both dispersing and non-dispersing animals. conclusion the ranging behaviour for hand-raised rhinos seemed to be partly depended on whether there was a human near the release site. hand-raised rhinos would preferentially settle at sites close to areas where human activity was significant. further, seasonal ranges established in the vicinity of human settlements were smaller than for those established independent of settlements. in this study, there was no conclusive data to show that any of the hand-raised rhinos, including adults of over 10 years old, had stabilised their seasonal range sizes 2.5 years after release. in some cases, hand-raised rhinos used established roads when moving to new sites outside mnp. physical barriers koedoe 47/1 (2004) 98 issn 0075-6458 matipano ranging.qxd 2004/04/14 02:04 page 98 issn 0075-6458 99 koedoe 47/1 (2004) were important to prevent hand-raised rhinos from moving about widely. an understanding of ranging behaviour is critical in managing hand-raised rhinos. ranging behaviour and reaction to human presence have a bearing on the security of hand-raised rhinos. having no fear of humans, hand-raised rhinos were vulnerable to poaching. management issues and recommendations there is no guarantee that a rhino will remain at a release site. hand-raised rhinos became attached to sites where they had been raised, and treated humans as surrogate mothers. those rhinos that did not return to raising sites or to human settlements made extensive post-release movements. these wide movements could result in reduced survival and reproduction as animals either entered areas of potential conflict with humans, or were killed (linell et al. 1997) by humans in communal lands. the season of release for rhinos is critical. rhinos that were released in the dry season tended to move more widely soon after release compared to those that were released in the wet season. rhinos moved widely in the dry season immediately after release probably in search of water and food. rhinos should be released during the wet season when the resources are abundant. both the season and area selected for release will govern resource availability. animals released in unsuitable habitats will move widely looking for water, food and refuge. in this study hand-raised rhino behaved in the opposite way to wild rhino which mukinya (1973), found to move widely where there were humans. in this study, the presence of humans seemed to be a factor for hand-raised rhinos to remain in the release area and not to move widely. animals that were released further from human settlement tended to reduce movements once they had found one. in reintroduction of rhinos, the aim is to reduce human influence and to allow animals to naturalise fast. hand-raised rhinos released close to humans will remain used to people and will fail to adapt to local conditions quickly. on the other hand rhinos that stay close to humans tend to survive for longer periods of time than those that live in areas where poachers can operate with little chance of detection (du toit, pers. comm.). however, if animals are released away from humans, they will have a higher probability of moving out of the protected area into high-risk areas. in reintroduction programmes hand-raised rhinos should be released away from human settlements, provided there are barriers to movement into unprotected areas. whether released adjacent to human settlement or not, monitoring is important for postrelease management. the time hand-raised rhinos took to settle after release was not established, and hand-raised rhinos were still ranging widely two and a half years after release. on release, hand-raised rhinos should be marked to allow identification and monitoring of individual animals and record dispersal patterns. monitoring of hand-raised rhinos should be continued for several years after release, until the animals have settled down. hand-raised rhinos are more vulnerable to poaching than their wild counterparts. for at least two and half years after release, handraised rhinos sometimes did not run away from trackers, and would advance to within a few metres of researchers. in due course, hand-raised rhinos will make a significant contribution to the black rhino population in mnp. currently (october, 2000), the combined total number of hand-raised rhinos and those still in bomas makes up about 6 % of the mnp black rhino population. if there is an outbreak of poaching, hand-raised rhinos will be the first to suffer. having a motivated, well-trained and adequately equipped antipoaching personnel is an important conservation measure for rhinos. hand-raised rhinos were released when they were more than three years old; age at which their wild counter-parts are weaned and when they are not old enough to establish home ranges. at the time hand raised rhino were released the resident population density was matipano ranging.qxd 2004/04/14 02:04 page 99 below 0.25 animals/km². by the time this study was done there were no deaths of hand-raised rhino caused by intraspecific fights. hand-raised rhinos should continued to be released when they are aged above three years especially in areas where the resident population is at a low density. it is to be expected that on weaning, animals will extend their ranges. the extent of ranging after weaning is important as rhinos might enter into hostile areas. the age at which hand-raised rhinos establish home ranges is not known. hand-raised rhinos should be released before they are sexually mature and before they reach the age at which wild rhinos establish home ranges to give them time to learn about their environment. there is also need to determine the density of the resident population at which intraspecific fights would cause significant loss of hand-raised rhinos. in this study, those rhinos that moved northwards were resident for some time in habitats next to the lake. the reason why most of the animals that went to the lakeshore stayed in habitats close to the lake is not clear. reasons why hand-raised rhino were found on the lakeshore might include that the rhino were taking refuge from aggressive wild counterparts resident in the interior of the study area and or were attracted to sites of high human activity at tourist camps or house-boats in bays. physical barriers, e.g., a strong perimeter fence can prevent extensive movements especially where the release area is small. it would be appropriate to release rhinos in areas large enough to meet ecological and social requirements of the animals. the exact size of the release area for handraised rhinos to meet ecological and social requirements depends on factors including habitat quality, density of the resident population, history of the animal and existence of the boundary game fence. unfenced release areas for hand-raised rhinos should be at least 1000 km². in unfenced areas, animals should be released at sites from where there is the greatest distance to the edge of the conservation area. for animals that have learnt how to repeatedly move out of a large release area, relocation to a totally new area might be a solution. such rhinos should be relocated to fenced areas that offer animals better security. most of the department of national parks and wildlife management areas designated for release of rhinos are surrounded by communal lands. if rhinos move into communal lands there will be potential conflict with local communities. nevertheless, some residents in the communal lands assist by reporting the presence of stray rhinos in their areas. an education programme is necessary to allow local communities to appreciate efforts to conserve endangered species. this includes the safari industry within and outside the park, as well as local residents in the surrounding communal lands. in the case of mnp, the adjacent nyaminyami district is a designated communal area management programme for indigenous resources (campfire) area with a wildlife section that can jointly work with mnp staff to carry out antipoaching and monitoring activities. acknowledgements this article was produced as part of my msc. thesis. financial support from the european commission is greatly appreciated. i would like to thank my msc project internal supervisor dr. c. a. m. attwell and external technical supervisor mr. r. f. du toit. i thank the chief ecologist (terrestrial ecology) mr. e. chidziya of the department of national parks and wildlife management for his support during the study. i am grateful to matusadona staff, especially senior scout e. muchuchutiti and scout t. nyakashaya for their assistance during field work. finally, i thank the acting director, brigadier e. kanhanga of the department of national parks and wildlife management for granting permission to carry out the study in matusadona national park. references adcock, k. 1994. the relevance of territorial behaviour in black rhino to the population management. pp. 82–86. in: penzhorn, b.l. & n.p. kriek (eds.). proceedings of a symposium on rhinoceros as game ranch animals, onderstepoort. september, 1994. adcock, k., h. hansen & h.j. lindemann. 1998. lesson from the introduced black rhino populakoedoe 47/1 (2004) 100 issn 0075-6458 matipano ranging.qxd 2004/04/14 02:04 page 100 tion in pilanesberg national park. pachyderm 26: 40–68. aebischer, n.j., p.a. robertson & r.e. kenward. 1993. compositional analysis of habitat use from animal radio-tracking data. ecology 75(5): 1313–1325. anderson, d.j. 1982. the home range: a new nonparametric estimation technique. ecology 63: 103–112. atkinson, s.j. 1995 maintenance of captive black rhinos (diceros bicornis) on indigenous browse in zimbabwe: energetics, nutrition and implications for conservation. msc. thesis. university of zimbabwe. bekoff, m. & l.d. mech. 1984. simulation and analyses of space use: home range estimates, variability, and sample size. behaviour research methods, instruments, and computers 16(1): 32–37. brett, r. 1998. mortality factors and breeding performance of translocated black rhinos in kenya: 1984–1985. pachyderm 26:69–82. burt, w.h. 1943. territoriality and home range concepts as applied to mammals. journal of mammalogy 24: 346–353. conybeare, a.m. 1991. elephant occupancy and vegetation change in relation to artificial water points in a kalahari sand area of hwange national park. phd thesis. university of zimbabwe, harare. dexter, n. 1999. the influence of pasture distribution, temperature and sex on home range sizes of feral pigs in a semi-arid environment. wildlife research 26: 755–762. dixon, k.r. & j.a. chapman. 1980. harmonic mean measure of animal activity areas. ecology 61: 1040–1044. frame, g.w. 1980. black rhinoceros (diceros bicornis l.) sub-populations on the serengeti plains, tanzania. african journal of ecology 18: 155–166. goddard, j. 1967. home range, behaviour and recruitment rates of two black rhinoceros populations. east african wildlife journal 5: 133–150. harris, s., w.j. cresswell, p.g. forde, w.j. trewhella, t. woollard & s. wray. 1990. home range analysis using radio-tracking data—a review of problems and techniques particularly applied in the study of mammals. mammal review 20 (2/3): 97–123. hitchins, p.m. 1969. the influence of vegetation types on the sizes of home ranges of black rhinos in hluhluwe game reserve, zululand. lammergeyer 10: 81–85. hitchins, p.m. 1971. preliminary findings of a radiotracking study on the black rhinoceros in hluhluwe game reserve, zululand. a symposium in biometry, pretoria. (council for scientific and industrial research, s57: 95–100). joubert, e. & f.c. eloff. 1971. notes on the ecology and behaviour of the black rhinoceros, diceros bicornis, l. 1758 in south west africa. madoqua ser. 1(3): 5–53. kiwia, h.y.d. 1989. ranging patterns of the black rhinoceros (diceros bicornis, (l)) in ngorongoro crater, tanzania. african journal of ecology 27(4): 305–312. linell, j.d., r. aanes, j.e. swenson, j.odden & m.e. smith. 1997. translocation of carnivores as a method for managing problem animals: a review. biodiversity and conservation 6: 1245–1257. loutit, b.d., g.w. louw & m.k. seely. 1987. preliminary observation of food preferences and chemical composition of the diet of the desertdwelling black rhino. madoqua 15: 35–54. mohr, c.o. 1947. table of equivalent populations of north american small mammals. american midlands naturalist 37: 223–249. monro, r.h. 1982. an appraisal of some techniques to investigate the feeding ecology of large herbivores with reference to study of impala in the northern transvaal. african journal of ecology 20: 71–80. mukinya, j.g. 1973. density, distribution, population structure and social organisation of the black rhino in masai mara game reserve. east african wildlife journal 11(3 & 4): 385–400. owen-smith, r.h. 1988. megaherbivores. the influence of very large body size on ecology. cambridge: cambridge university press. robertson, p.a., n.j. aebischer, r.e. kenward, i.k. hanski & n.p. williams. 1998. simulation and jack-knifing assessment of home-range indices based on underlying trajectories. journal of applied ecology 35(6): 928–940. smithers, r.h.n. & v.j. wilson. 1979. checklist and atlas of the mammals of zimbabwe-rhodesia. museum memoir 9: 1–147. swihart, r.k. & n.a. slade. 1985. testing for independence of observations in animal movements. ecology 66: 1176–1184. tatman, s.c., b. stevens-wood & v.b.t. smith. 2000. ranging behaviour and habitat usage in black rhinoceros, diceros bicornis, in a kenyan sanctuary. journal of african ecology 38(2): 163–172. tufto, j., r. andersen & j. linnell. 1996. habitat use and ecological correlates of home size in a small cervid: the roe deer. journal of applied ecology 65: 715–724. issn 0075-6458 101 koedoe 47/1 (2004) matipano ranging.qxd 2004/04/14 02:04 page 101 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 saayman.qxd determining the economic value of game farm tourism p. van der merwe, and m. saayman van der merwe, p. and m. saayman. 2003. determining the economic value of game farm tourism. koedoe 46(2): 103–112. pretoria. issn 0075-6458. internationally tourism is accepted as one of the world's fastest-growing industries. the world tourism organisation (wto) indicated that tourist arrivals in 1998 grew by 2.4 % worldwide. the wto has forecast that the number of people travelling internationally will increase from 613 million in 1997 to 1.6 billion by the year 2020. ecotourism, which according to the wto is any form of tourism to an unspoilt nature area, is responsible for 20 % of the world's total tourism expenditure and is also rated the fastest growing of all tourism sectors. it is also a fact that 80 % of nature conservation in south africa is taking place on privately owned land such as game farms, and this forms part of ecotourism. the above endorses that ecotourism is an important product for south africa and a drawcard for international as well as local tourists. the main objective of this study was to determine the economic value of game farm tourism. this will be done by determining the economic value of each of the four pillars on which game farming is based, namely hunting, ecotourism, breeding rare game species and venison sales. data collation was done in two ways. firstly, research was conducted in the form of questionnaires. game farms were randomly sampled from the database of registered game farms. the aim of this questionnaire was threefold and determined the economic contribution of game farm tourism to ecotourism. secondly, a literature study was conducted that included the latest data by phasa (professional hunters association of south africa), nature conservation and trophy hunting in south africa. this paper will argue that game farm tourism makes a significant economic contribution to the economy of south africa, apart from the substantial economic contribution game farm tourism already makes to conservation. this paper will be organised as follows: the first section deals with the introduction, which indicates the growth of game farm tourism; the second section explains the methodology; the third section discusses the results; and the last section concludes the paper. key words: ecotourism, economic contribution, hunting, game farm tourism. p. merwe, and m. saayman, institute for tourism and leisure studies, potchefstroom university for che, private bag x 6001, potchefstroom, 2520 republic of south africa (ontpvdm@puknet.puk.ac.za; ontms@puknet.puk.ac.za issn 0075-6458 103 koedoe 46/2 (2003) introduction tourism has been accepted by the south african government, business and labour as one of the key drivers for the creation of wealth and economic empowerment (gcis 1999). this is supported by the fact that internationally tourism is acknowledge as one of the world's fastest-growing industries (shaw & williams 1994; cooper et al. 1993; kerzner 1995; anon 1998). as further support to the latter, the world tourism organisation (wto) forecasts that the number of people travelling internationally would increase to 1.6 billion by the year 2020 (world bank 1998; wto 1998, 2001a). if the target as predicted by the wto is reached it would result in an increase in the next seventeen years of almost $2 000 billion a year (van zyl 2000). fraser (1999) forecasts that south africa's tourism industry will grow by more than 5.5 % between 1998 and 2010, which is betsaayman.qxd 2005/12/09 11:24 page 103 ter than the predicted 4.1 % of the global market. it is also estimated that arrivals will increase from 6 million international tourists arrivals in 2000 to 30.5 million in 2020 (wto 2001b). the african continent can treble the size of its tourism industry by 2020 if proper efforts are made. this will result in forecast tourist arrivals of 77.3 million to africa. on the african continent, south africa attracts the most overseas tourists, which makes tourism one of the largest industries in south africa (wto 2001b). research indicated that tourism in south africa contributed approximately 4.6 % to the gross domestic product (gdp) the past four years. in 2002 tourism arrivals increased by 16 % ranking it one of the highest growth rates in the world (thornton & feinstein 2002; gcis 1998). for the overseas as well as local tourists scenic beauty and wildlife remain the major tourism attractions that south africa has to offer (gcis 1998; gcis 1999). it is a fact that 80 % of nature conservation in south africa is taking place on privately owned land such as game farms (eloff 2000; fox & du plessis 2000), and according to bothma (2002) there was a 2.5 % increase in land utilised for game farming from 1998 to 1999. the latter translates into a 300 000 ha per year increase for the purpose of game farming tourism. flack (2002a) states that it is estimated that the conversion rate from cattle to game farming in 2002 was nearly 500 000 ha, which is 200 000 ha more than the 1998 to 1999 increase. table 1 indicates the number of exempted game farms in south africa (exempted in this context implies that these game farms have permission to hunt within their boundaries), as well as the surface area. from table 1 it is clear that the limpopo province has the most game farms (2 484), followed by the northern cape (986) and thirdly the eastern cape (624). when considering the total surface area that is exempted, the northern cape outperforms the other provinces by 4 852 053 ha. the reason for this is that the northern cape's carrying capacity is lower, therefore more land is needed for the same number of head of game as for example in the eastern cape. in 2000, it was estimated that there were 7 000 privately owned game farms in south africa with a total surface area of 16 million ha (ebedes 2002), of which only 5 061 were exempted game farms, with a surface area of 10 364 154 ha (erasmus 2000). research conducted in 1993 indicated that the surface area of exempted game farms constituted 8.5 % of the total agricultural land in south africa, which increased to 12.5 % in 2000, with a total of 1.7 million head of game (eloff 2002a; erasmus 2000). the 2002 koedoe 46/2 (2003) 104 issn 0075-6458 table 1 number of exempted game farms in south africa province(year 2000) free state limpopo north west mpumalanga gauteng kwa zulu-natal eastern cape northern cape western cape total number of exempted game farms 180 2482 340 205 72 90 624 986 82 5061 surface area of exempted farms 147 743 3 325 652 364 935 276 016 82 076 168 841 881 633 4 852 0532 65 205 10 364 154 % of exempted farms 3.56 % 49.04 % 6.72 % 4.05 % 1.42 % 1.78 % 12.33 % 19.48 % 1.62 % 100 % % of exempted farms according to hectares 1.43 % 32.10 % 3.51 % 2.66 % 0.79 % 1.63 % 8.51 % 46.81 % 2.56 % 100 % source: eloff 2002a saayman.qxd 2005/12/09 11:24 page 104 statistics showed that 13.3 % of the agricultural land was used for game farming, while national and provincial game reserves only covered 6 % or 6.1 million ha of land in south africa (flack 2002; van der walt 2002). if one adds up the total land used for wildlife tourism it is almost 20 % of south africa's total surface area. as a significant surface area is set aside for the purpose of game farm tourism, the question arises whether this is productive land use. therefore the aim of this paper is to determine the economic value of game farm tourism in south africa, and to achieve the aim the paper is organised in four sections. the first section deals with an overview of game farm tourism. in the second section the methodology is discussed, and thirdly the economic value of the four pillars of game farm tourism is analysed. section four sets out the conclusions about the economic value of game farm tourism drawn from the study. methods methodology used in this research consists of a literature study and a survey. literature study the literature study was done in order to collate the latest data on game farm tourism provided by phasa (professional hunters association of south africa), nature conservation and trophy hunting in south africa respectively. in other words all existing data, which included the growth of hunting and game sales, were consulted. previous research done in the eastern cape on game farms was also included (radder et al. 2000). the literature study was done to determine the economic value of the four pillars of game farm tourism, which include breeding of rare game species and live game sales, hunting, processed game products and ecotourism expenditure (van der merwe & saayman 2002), and these are described next. breeding of scarce game species and live game sales eloff (1999) describes game farming with scarce game species as game farming where the emphasis is more on the production of the scarce game species with the aim of selling them again to other game farmers. animals that are classified as scarce game species include buffalo, black rhino, sable antelope and the endangered roan antelope, just to name but a few. live game sales or auctions imply the selling of game (endangered and not endangered) at auctions. here farmers have an opportunity to bid on game species that they are interested in. hunting hunting is defined by nature conservation as follows: in relation to any wild animal, it means by any means whatsoever to hunt or search for, to kill, capture or attempt to kill or capture, or pursue, follow or drive with intent to kill or capture, or to shoot at, poison, lie and wait for or wilfully disturb (eastern cape province 1974). hunting can be broken down into trophy hunting, biltong hunting, bow hunting and bird hunting. however, for the purpose of this paper only trophy hunting and biltong hunting (of which bow hunting forms part) will be discussed. the reason is that hunting is the single biggest economic contributor to game farm tourism. processed game products venison or game products are currently mentioned regularly as potential market resources for game farm owners in south africa (bothma 2002). processed game products imply that excess game is killed for their meat (venison) to produce meat products such as, biltong, salami, chilli bites, dry wors and venison for restaurants. ecotourism koch (1995) describes ecotourism as ‘purposeful travel to natural areas to understand the cultural and natural history of the environment, taking care not to alter the integrity of the ecosystem, and producing economic opportunities that make conservation of natural resources beneficial to local people’. van wyk (1995) defines ecotourism as ‘an enlightening nature travel experience that contributes to conservation of the ecosystem, while respecting the integrity of the host’. the latter emphasises the concept of an experience. this forms the backbone of tourism and tourism products should be developed in order to provide for a special experience. the following are some ecotourism activities that can be provided to tourists on a game farm: · hiking trails, bird watching, photographic safaris, 4x4 trails, canoeing and abseiling for more adventurous tourists; · conference facilities; · promotion of unusual attractions, like caves, bushmen paintings, waterfalls and ruins of previous civilisations; issn 0075-6458 105 koedoe 46/2 (2003) saayman.qxd 2005/12/09 11:24 page 105 · hot-air balloon trips, horse safaris, walking safaris, mountain bike trails and donkey cart trips; and · educational tours where the tourist can learn more about nature, which means it does not have to include the big five (van der merwe & saayman 2002). survey secondly, research was conducted in the form of questionnaires. game farms were randomly sampled from the database of the north west parks and tourism board. two surveys were conducted, one among game farm owners and one among tourists visiting game farms. of the 340 exempted game farms in the north west province (according to the north west parks and tourism board's database) it was decided to select every ninth game farm in order to obtain a 10 % sample. these game farms were then visited in person during july 2002, and all questionnaires were hand-delivered and collected. the second questionnaire targeted tourists visiting a game farm. a random sample was also drawn from the database of the north west parks and tourism board, of which ten game farms were prepared to participate in the research. the criterion used was that these game farms had to have a well-developed tourism infrastructure. the latter includes proper accommodation and ecotourism facilities in order to attract tourists. an average of five questionnaires per game farm were personally handed out to visitors during the weekends of 30 august and 6 september 2002. 50 (n = 50) questionnaires were received back from the respondents. research by radder et al. (2000) done in the eastern cape was also used. radder et al. (2000) conducted a survey of 616 questionnaires among local hunters, 107 of which could be used (17.4 % response rate). results the results as mentioned earlier will be based on the four pillars of game farm tourism, namely hunting, breeding scarce game species and live game sales, processed game products and ecotourism expenditure in order to achieve the aim of the study. breeding scarce species of game and live animal sales live game sales and the breeding of scarce species are the second biggest generator of revenue for game farm tourism. during the 2000 season live game sales generated r180 million of the gross income of the south african game farm industry (eloff 2002a; eloff 2003). this represents a total of game sales and not only sales at auctions. analysis of game auctions over the past ten years revealed an average increase of 9 % per year. koedoe 46/2 (2003) 106 issn 0075-6458 table 2 auction sales over past ten years year number of % change from turnover % change from game sold previous year (tax excluded) previous year in terms of turnover 1991 8 292 8 999 871 1992 9 546 (+)15 % 10 059 969 (+)20.66 % 1993 11 499 (+)19.93 % 11 732 596 (+)8.04 % 1994 11 096 (-)3.08 % 11 705 605 (-)0.23 % 1995 9 171 (-)17.35 % 14 335 894 (+)22.47 % 1996 11 340 (+)23.65 % 26 559 557 (+)85.26 % 1997 12 077 (+)6.5 % 28 526 052 (+)7.4 % 1998 14 354 (+)18.85 % 40 017 964 (+)40.29 % 1999 15 455 (+)7.67 % 53 705 823 (+)34.2 % 2000 17 702 (+)14.54 % 62 960 451 (+)17.23 % 2001 17 282 (-)2.37 % 87 000 473 (+)38.18 % 2002 20 022 (+)15.85 % 105192180 (+)20.91 % source: eloff 2003 saayman.qxd 2005/12/09 11:24 page 106 issn 0075-6458 107 koedoe 46/2 (2003) table 3 percentage price increase (+) or decrease (-) in average auctioned game prices over the past ten years species % price increase average % price % increase (+) or in ten years increase per year decrease (-) in 2000 in ten years compared with 1999 roan antelope 408.12 % 40.81 % 8.2 % (-) tsessebe 210.76 % 21.08 % 49.25 % (+) blesbok: white 134.78 % 13.48 % 19.50 % (-) common 150.37 % 15.04 % 3.84 % (+) blue wildebeest 199.19 % 19.92 % 0.98 % (-) bontebok 215.9 % 21.59 % 15.9 % (+) bushbuck 284.9 % 28.49 % 23.92 % (+) buffalo (disease free) 296.52 % 29.65 % 53.13 % (+) duiker 187.25 % 18.73 % 32.07 % (+) eland (cape) 105.8 % 10.58 % 13.44 % (+) gemsbok 164.02 % 16.40 % 4.74 % (+) giraffe 91.04 % 9.10 % 13.79 % (+) klipspringer 143.48 % 14.35 % 6.45 % (-) kudu 142.51 % 14.25 % 16.11 % (+) lion 5.42 %(-) 0.54 %(-) 54.86 % (-) nyala 210.32 % 21.03 % 80.33 % (+) reedbuck 492.33 % 49.23 % 49.65 % (+) impala 198 % 19.8 % 5.11 % (+) red hartebeest 157.17 % 15.72 % 8.74 % (+) mountain reedbuck 298.80 % 29.88 % 22.14 % (+) burchell's zebra 81.82 % 8.18 % 18.97 % (+) springbok (common) 223.73 % 22.37 % 6.83 % (-) heartwater 74 % 7.4 % 30.64 % (+) black 203 % 20.3 % 13.47 % (+) white 384.38 % 38.44 % 155.2 % (+) steenbok 233.40 % 22.34 % 53.61 % (+) black rhinoceros 53.06 % 5.31 % 69.68 % (+) black wildebeest 499.78 % 49.98 % 5.1 % (+) sable antelope 111.90 % 11.19 % 9.85 % (+) fallow deer 288.28 % 28.83 % 1.64 % (+) grey rhebuck 826.83 % 82.68 % 12.72 % (-) warthog 112.36 % 11.24 % 8.7 % (-) ostrich 105.82 % 10.58 % 39.46 % (+) waterbuck 129.90 % 12.00 % 36.5 % (+) white rhinoceros 303.66 % 30.37 % 39.67 % (+) total game 113.53 % 11.35 % 14.54 % (+) total turnover 600.18 % 60.02 % 17.23 % (+) (source: eloff. 2002(a):81) the turnover per year grew from almost r9 m in 1991 to r105 m in 2002, with an average growth rate of r38 million per year over 12 years (table 2). live game sales stabilised during the 2001 season but there is a growing demand, especially for scarce game species. the more common species did show signs of price maturity, although new record prices were recorded (bothma 2002). the fact that the demand for scarce game species is still on the increase leads to an encouragingly intensive system of production of these animals. it is interesting to note that the prices of male animals are generally lower than those of females, the reason is simply that the latter can be used better for breeding purposes, unless they are of exceptional trophy value. as the number of scarce animals increases, it will eventually lead to a saayman.qxd 2005/12/09 11:24 page 107 lower market price but they will still have a high trophy value, much higher than their meat value (eloff 2002a, 2002b). table 3 indicates the average price increase or decrease of the different game species over the past ten years. flack (2002b) indicates that in 1992 there were only nine auctions per year, with 9 546 head of game auctioned. this grew to an outstanding 48 auctions in 2000 (17 702 head of game were auctioned), and increased by a further 38 % in 2001. in 2002, it again increased by nearly 21 % with a total of 20 022 head of game auctioned (table 2). the five game species that showed the largest increase in sales in 2000-2001 were as follows; tsessebe (60.9 %), springbok (31.2 %), sable antelope (24.2 %), nyala (20.2 %) and burchell's zebra (13.9 %) (eloff 2002a; table 3). hunting hunting is the biggest generator of income for game farms and for the purpose of this paper can be divided into two types, namely trophy hunting and biltong hunting. trophy hunting according to table 4 there were 2 671 active hunting outfitters/operators and 3 589 active professional hunters registered with nature conservation during 2000 and 2001 season. during the same period south africa received a total of 5 304 trophy hunters (excluding biltong hunters), which is 1 284 more than during the 1999–2000 season (4 020 trophy hunters). from table 4 it is clear that the two leading provinces in trophy hunting in 2001 were limpopo and eastern cape. according to koedoe 46/2 (2003) 108 issn 0075-6458 table 4 trophy hunting statistics 2001 province ec nc fs kzn nw gp lm mp total active hunting outfitters 99 33 250 113 99 32 1972 73 2671 active professional hunters 229 179 250 287 261 155 1972 256 3589 clients 1183 537 688 588 465 62 1644 137 5304 total animals 8941 4034 3143 2531 2696 185 10287 429 32246 animals per client 7.56 7.51 4.57 4.3 5.8 2.98 6.26 3.13 6.08 clients per hunting outfitter 11.95 16.27 2.75 5.2 4.7 1.94 0.83 1.88 clients per professional hunter 5.17 3 2.75 2.05 1.78 0.4 0.83 0.54 total hunting days 9223 4134 4425 3017 3024 165 28432 1112 53532 average. length of hunt (days) 7.8 7.7 6.43 5.13 6.5 2.66 17.29 8.12 10.09 wc=western cape; ec=eastern cape; nc=northern cape; fs=free state; kzn=kwazulu-natal; nw=north west; gp=gauteng; lm=limpopo; mp=mpumalanga (source: phasa. 2001) table 5 income of foreign trophy hunters year amount generated 1996 4000 (hunters and non-hunters) r117 653 190 (daily rates and trophy fees) 1997 7500 (hunters and non-hunters) r175 063 050 (daily rates and trophy fees) 1998 7500 (hunters and non-hunters) r156 871 650 (daily rates and trophy fees) 1999 8604 (hunters and non-hunters) r238 158 000 (daily rates and trophy fees) saayman.qxd 2005/12/09 11:24 page 108 table 5, the amount that hunting generated was r117 m in 1996, and it increased to r237 m in 1999, which includes only trophy hunters that made use of phasa-registered professional hunters. therefore, these statistics are regarded as the official trophy hunting statistics. it is interesting to note that the number of trophy hunters and non-hunters—person who accompany the hunter but does not hunt—remained the same, i.e. 7500, in 1997 and 1998, although the amounts spent were less in 1998. the reason for this may be that the price of game increased from 1997 to 1998. the amount generated then increased from r156 871 650 in 1998 to r 238 158 000 in 1999. it is also important to take into consideration that the rand weakened against the dollar during this period and that only phasa outfitters were recorded (phasa 2000). during the 2001 hunting season, trophy hunters hunted a total of 32 246 animals over a total of 53 532 hunting days sold (table 6). when the average daily rate of $350 is taken and multiplied by the number of hunting days sold, it adds up to almost r225 million. the total for accommodation and hunted species combined adds up to r532 million (table 6). the high demand among foreign trophy hunters for hunting in south africa might be due to a favourable exchange rate as well as an increase in the wildlife population following the establishment of wellmanaged game farms. thirty seven percent of all rowland ward hunting records have been recorded in south africa since 1975, as well as 92.9 % of all safari club international records (van rooyen & van rooyen 2002). biltong hunting according to radder et al. (2000) biltong hunters tend to spend the amounts listed in table 7 when visiting game farms in the eastern cape. radder et al. (2000) indicated that the average length of stay of biltong hunters is two days and that the average size of a group consists of four hunters. table 7 clearly illustrates that most money spent on a hunting trip is spent on game, followed by accommodation and refreshments. hunters tend to spend r300 on daily tariffs and r200 on issn 0075-6458 109 koedoe 46/2 (2003) table 6 income: trophy hunting and accommodation season hunting days daily rate total total animals sold (excluding trophy prize) and accommodation 2000 32 652 $350 $11 428 200 $30 818 710 (r85 711 500) (r231 140 325) 2001 53 532 $350 $18,736,200 $44 325 965 (r224 834 400) (r532 000 000) (source: phasa, 2002) table 7 biltong hunting expenditure accommodation, meals and refreshments per hunting trip r200.00 accommodation expenses per hunting trip considered reasonable r200.00 transport costs per hunting trip considered reasonable r200.00 daily tariffs considered reasonable per hunting trip r300.00 game prices per hunting trip considered reasonable r1 100.00 licence expenses considered reasonable r22.50 saayman.qxd 2005/12/09 11:24 page 109 transport per biltong hunting trip. add to this the licence fee and it gives an average total of r2022.50 per biltong hunter per hunting trip (radder et al. 2000). based on the 18 000 registered members of the south african hunters and game conservation association, the total amount generated by them for one hunting trip per year is approximately r36 m. this is just the tip of the iceberg, the reason being that most of the biltong hunters in south africa are not yet registered members of the latter association and that each province also has its own hunting association, apart from the lattermentioned national body. therefore, more research still needs to be done to determine the real economic value of biltong hunters to game farm tourism in south africa (a. van dyk pers. comm.). processed game products the potential for the production of venison as a legitimate form of wildlife utilisation, especially in the more arid regions of africa, has been recognised for a long time (bothma 2002). surplus game, usually females, which were not hunted during the season, can be used to supply processed game products to exclusive markets, both overseas and in south africa. it is important to make these products exotic for the customer. the fact that there is a global movement among humans to live healthier can be an important marketing factor to help sell venison, because venison has a very low fat percentage. both smoked and fresh processed game products are sold to restaurants and hotels (gouws 1999; brits & herselman 1988). when game (venison) is compared with domestic herbivores for meat production, game cannot compete because domesticated herbivores have been genetically bred and selected for many centuries specifically for meat production. the advantage that game has over domestic herbivores is the fact that they offer a wider diversity of products. there are at least 45 types of wild herbivore in southern africa, of which many occur more or less in the same ecosystem (bothma koedoe 46/2 (2003) 110 issn 0075-6458 table 8 tourist spending category amount spent accommodation r1699.38 entrance fee r18.21 restaurants r408.36 food r335.77 beverages r171.63 tobacco products r15.30 clothing and shoes r44.00 transport r411.00 recreation r81.00 medicine r1.80 toiletry r 6.20 total r3 192.65 2002). meat production also generates the lowest income of all the pillars on which game farm tourism is based. the gross income for meat production sales was estimated at r20 million in 2000 (eloff 2002a). ecotourism expenditure the literature study confirmed that little research has been conducted on ecotourism expenditure in relation to game farm tourism. it is therefore difficult to determine the real economic value of ecotourism on game farms. however, a pilot study done in the north west province as part of this paper revealed the spending of tourists to game farms set out in table 8. research done in the north west province indicated that the average size of groups travelling to/visiting game farms was three persons and that they tended to stay an average of three days. it is clear that most money spent (table 8) by tourists was on accommodation and transport per ecotourist group per trip. the least money was spent on clothing, tobacco, medicine, toiletries and recreation activities. the reason for this is that game farms tend not to have tourist or curio shops where visitors can spend money. recreation spending is also low as it was found that saayman.qxd 2005/12/09 11:24 page 110 only few game farms made provision for recreational activities. the average spending per group (3 persons) for three days is r3 192.65. each person would then spend an average of r1 064.21 per trip. hence one person spends on average r354.73 per day over a three-day period, so that if 100 000 tourists took one trip to a game farm during a year and spent r1 064.21 per trip, this would add up to approximately r106 million. this clearly indicates the economic value of ecotourism to game farm tourism. conclusion the purpose of this paper was to determine the economic value of game farm tourism. even though this was the purpose, one cannot but mention also the conservation value of game farm tourism. the latter entails more land for consumptive utilisation, even though some conservationists still cannot perceive the contribution of the consumptive utilisation of game in the form of hunting to both the economy and the tourism industry in south africa. something else that also has to be kept in mind is that as a consequence more breeding centres are established throughout south africa, which has a positive effect in terms of the number of species available. this paper also showed that even though there are gaps in the statistics of some of the aspects (pillars) of game farm tourism, the latter nevertheless has a significant economic value, as can be seen from the 2000 statistics: hunting (trophy and biltong) r568 m game sales r180 m game products r20 m ecotourism r106 m total r874 m over and above the fact that game farm tourism in south africa generated approximately r874 m, the 7 000 game farms also employ approximately 63 000 people. in order to grow this sector, this research recommends that game farm owners/managers must, as far as possible, implement all four pillars. the ecotourism component should be exploited and opportunities where tourists can spend more money should be created. this implies, for example, providing recreational activities, restaurants and the selling of curios and services. it is also a recommendation of this paper that more research be done, especially into the value of ecotourism and into biltong hunting needs. acknowledgements we thank dr andrea saayman for the assistance given. references anon. 1998. tourism leading global economic driver of the 21st century. satour news 3, december.. bothma, j. du p. 2002(a). some economics of wildlife ranching. wildlife group symposium on game ranch management, s.a. veterinary association, onderstepoort, 1-2 november 2002 brits, d. & w. herselman. 1988. wildbenutting. fauna & flora 44:18-24. cooper, c., j. fletcher, d. gilbert & s. wanhill. 1993. tourism principles & practice. london: pitman. eastern cape province (south africa). 1974. ordonnansie op natuuren omgewingsbewaring, no. 19 van 1974. kaapstad. ebedes, h. 2002. preface. in: ebedes, h., b. reilly, w. van hoven & b. penzhorn (eds.). sustainable utilisation-conservation in practice. proceedings of the 5th international wildlife ranching symposium 2001. pretoria. eloff, t. 1999. professionalisme in die wildbedryf. klk nuus 29(1): 9. eloff, t. 2000. die omvang van die wildbedryf in suid-afrika. potchestroom: pu vir cho. eloff, t. 2002(a). the economic realities of the game industry in south africa. in: ebedes, h., b. reilly, w. van hoven & b. penzhorn (eds.). sustainable utilisation-conservation in practice. proceedings of the 5th international wildlife ranching symposium 2001. pretoria. eloff, t. 2002(b). boerdery met skaars groter spesies. game & hunt 8(2): 21. eloff, t. 2003. gemiddelde wildveilingpryse 2002. game & hunt 9(2): 21. issn 0075-6458 111 koedoe 46/2 (2003) saayman.qxd 2005/12/09 11:24 page 111 erasmus, s.d. 2000. die ekonomiese lewensvatbaarheid van 'n wildvleisbewerkingsaanleg in suid-afrika. m.com. verhandeling, potchefstroomse universiteit vir cho. flack, p.h. 2002. the conservation revolution. game & hunt 8(10): 29, 31,33. fox, t. & p. du plessis. 2000. hunting in south africa. africa indigo in touch with the african renaissance 3(3): 40. fraser, j. 1999. the travel industry. 2nd ed. new york: van nostrand reinhold. gouws, a. 1999. smul gevaar van surpluswild weg. landbouweekblad 1124: 16-17. gcis (government communication and information system). 1998. south africa official yearbook. cape town: rustica. gcis (government communication and information system). 1999. south africa official yearbook. cape town: rustica. kerzner, s. 1995. foreword. in: bennett, j.a., (ed.). managing tourism services. pretoria: van schaik. koch, e. 1995. solutions conservation and rural community development in south africa. optima 41(2): 35. phasa. 2000. south african hunting statistics. pretoria: professional hunters association of south africa. phasa. 2001. south african hunting statistics. pretoria: professional hunters association of south africa. phasa. 2002. south african hunting statistics. pretoria: professional hunters association of south africa. radder, l. & p. van niekerk & a. nagel. 2000. matching the hunting experience stage by selected farmers in the eastern cape to the value expectations of hunters: research report. port elizabeth: port elizabeth technikon. shaw, s. & a.m. williams. 1994. critical issues in tourism: a geographical perspective. oxford: blackwell. thornton, g. & k. feinstein. 2002. tourism talk: a summary of the most important data and statistics relevant to the regional tourism industry. journal of southern african tourism 1(3): 4853. van der merwe, p. & m. saayman. 2002. managing game farms from a tourism perspective. potchefstroom: leisure c publications. van der walt, j. 2002. proliferation of game ranches. game & hunt (10)8: 7, oct. van rooyen, n. & j. van rooyen. 2002. handling and measuring trophies. pp. 513-546. in: bothma, j. du p., (ed.). game ranch management. 4th ed. pretoria: van schaik. van wyk, m.l. 1995. die invloed van ekotoerisme op lewenskwaliteit in ontwikkelende gemeenskappe: 'n ontwikkelingskommunikasieperspektief. m.com. verhandeling. potchefstroomse universiteit vir cho. van zyl, j. 2000. 'n boer maak 'n plan: wildbedryf: fokus. finansies & tegniek: 22. apr. 14. world bank. 1998. world bank group and world tourism organisation examine role of tourism in development. http://www.world-tourism.org/ pressrel/wbwto [date of access: 3 dec. 1998]. wto (world tourism organisation). 1998. results prove strength of tourism. http://www.worldtourism.org/newslett/febmar99/1998results [date of access: 19 mar. 1999]. wto (world tourism organisation). 2001(a). wto long-term forecast tourism 2020 vision. [web:] h t t p : / / w w w. w o r l d t o u r i s m . o r g / m a r k e t _ research/data/forecast.html [date of access: 19 mar. 1999]. wto (world tourism organisation). 2001(b). africa to triple tourist numbers. http://www. world-tourism.org/newsroom/releases/more _releases/r010607.html [date of access: 19 mar. 1999]. koedoe 46/2 (2003) 112 issn 0075-6458 saayman.qxd 2005/12/09 11:24 page 112 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 abstract introduction materials and methods results discussion conclusion acknowledgements references about the author(s) marieka gryzenhout department of genetics, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa errol d. cason department of animal and wildlife and grassland sciences, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa marcele vermeulen department of microbial, biochemical and food biotechnology, faculty of agricultural and natural sciences, university of the free state, bloemfontein, south africa george a.e. kloppers department of genetics, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa brooke bailey department of genetics, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa soumya ghosh department of genetics, faculty of natural and agricultural sciences, university of the free state, bloemfontein, south africa citation gryzenhout, m., cason, e.d., vermeulen, m., kloppers, g.a.e., bailey, b. & ghosh, s., 2020, ‘fungal community structure variability between the root rhizosphere and endosphere in a granite catena system in kruger national park, south africa’, koedoe 62(2), a1597. https://doi.org/10.4102/koedoe.v62i2.1597 note: special issue: connections between abiotic and biotic components of a granite catena ecosystem in kruger national park, sub-edited by beanelri janecke and johan van tol. original research fungal community structure variability between the root rhizosphere and endosphere in a granite catena system in kruger national park, south africa marieka gryzenhout, errol d. cason, marcele vermeulen, george a.e. kloppers, brooke bailey, soumya ghosh received: 26 sept. 2019; accepted: 28 apr. 2020; published: 29 oct. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract fungi colonise various substrates such as organic matter (dead or alive) from plants or animals. these fungi can be specialists (i.e. belonging to a substrate) or generalists (i.e. surviving on different types of organisms). fungi fulfil various functions in specialised niches, for example, acting as plant pathogens, helping in plant growth from the root systems or decomposing organic matter and fertilising soil. species are specialised to occur in only one niche, or others can utilise or occur in various niches. for example, certain species occur only within certain plant tissues (endophytes), on the exterior surface of the plants growing above the ground (epiphytes) or below the ground in the sphere surrounding the roots (rhizosphere). different soil types or conditions can favour certain species. this study used environmental sequencing to characterise the fungal communities associated with the root exterior and interior of sida cordifolia, a plant growing across the varying soil conditions of the catena system. fungal rhizosphere communities between three commonly occurring plant species – s. cordifolia, melhania acuminata (both malvaceae) and kyphocarpa angustifolia (amaranthaceae) – in one of the soil types were also studied to compare and contrast the fungal rhizosphere communities of these herbs. molecular operational taxonomic units co-occurred between niches, soil conditions and the rhizospheres of three plants at the same location, whilst others were restricted to only one niche or plant species. results showed that soil conditions in a catena can influence the associations of fungal species between different catena zones, on the outside and inside of the roots, and that these communities also differ between plant species. conservation implications: this study showed that complex and sensitive fungal communities are associated with plant roots in different zones of the catena. this is most likely also true between different habitats and soil types on a larger scale. this study emphasises the need to also manage a catena system on the micro-ecological scale whilst framing conservation and management plans of the kruger national park. keywords: fungi; sodic soil; grazing lawn; rhizosphere; root endophytes; catena. introduction soils harbour a great diversity of fungal species that have various ecological functions (bridge & spooner 2001; havlicek & mitchell 2014). saprophytic fungi break down dead organic matter and, in turn, fertilise the soil (setala & mclean 2004). certain plant fungal pathogens are specifically adapted to infect plants through roots and to spread or survive in soils, while some pathogens affecting tissues of plants growing above the soil also have the ability to survive in soils. propagules of more specialised below-ground fungi, such as mycorrhiza that form specialised root associations benefitting plant health, can also be found in surrounding soils (moore, robson & trinci 2011). similarly, propagules of various fungi occurring in different niches and substrates above ground can also be found in soils (aylor 2003; taylor & bruns 1999). soil conditions can permit specific fungal species to grow and produce fruiting structures. soil qualities such as ph and nutrient concentrations, soil types, temperature and humidity may influence optimal growth conditions for species (rousk et al. 2010). furthermore, types of vegetation in a particular area such as grasslands and forests, which harbour their own unique fungal communities, serve as inoculum sources for associated soils (talbot et al. 2014). similarly, propagules of coprophilous fungi (those closely associated with animal dung) can also be isolated from soil (richardson 2001). fungi occurring asymptomatically within plant tissues are named endophytes and they play diverse roles (rodriguez et al. 2009). these vary from highly symbiotic relationships, for example, conferring traits such as herbivore resistance to their hosts or boosting growth, to detrimental as is the case with latent plant pathogens. other fungi do not have any specific known function, or simply have the ability to overcome plant resistance and infect the tissues (rodriguez et al. 2009). endophyte communities can differ between plant tissues; for instance, some endophytes are better specialised to infect leaves (khare, mishra & arora 2018). roots also often harbour very specialised endophytes that have functions beneficial to their host (sabra et al. 2018). root endophytes can include fungi that infect through aerial tissues or from surrounding soils and often play beneficial roles for plant health (glynou et al. 2016; rodriguez et al. 2009). however, certain fungi have the ability to occur across various plant tissues. plant pathogens have varying biological compositions (agrios 2005). as already mentioned, latent pathogens can reside inside plant tissues without causing disease symptoms until triggered by specific stimuli such as stress conditions (slippers et al. 2007). other pathogens do not follow this strategy and can only cause disease symptoms. certain pathogens are also adapted to only infect aerial tissues or below-ground tissues such as roots. however, survival strategies of pathogens may enable them to survive in other substrates such as soil or water (hargreaves et al. 2015; havlicek & mitchell 2014; lareen, burton & schafer 2016). the rhizosphere is an interface consisting of a zone of soil that is associated with the roots (hiltner 1904). often this area contains exudates of the plant, which select for specific fungal communities to be present (lareen et al. 2016; sasse, martinoia & northen 2018). this area has an important function where fungi, as well as other microbial organisms, can facilitate the movement of nutrients (mommer, kirkegaard & van ruijven 2016; schippers, bakker & bakker 1987). other functions include protection against organisms feeding on roots, or plant pathogens (moore et al. 2011). grazing lawns are unique ecological phenomena (mcnaughton 1984). areas of grazing lawns contain natural vegetation significantly different from the surrounding area. the vegetation has a high nutritional value and productivity, attracting animals that in turn fertilise the soil again (verweij et al. 2006). it is also known that these areas are self-sustaining although the mechanisms behind this phenomenon are still unknown (archibald 2007; hempson et al. 2015). grazing lawns can form part of a catena system, such as the one forming the topic of this special issue. a catena consists of a serial soil group sequence from foothills to the crest, and that originated from the same geological material (brady & weil 2002). the kruger national park (knp) is one of the largest game reserves in africa (carruthers 2017). it covers the north-eastern part of south africa and now forms part of the great limpopo transfrontier park, linking it with the gonarezhou national park in zimbabwe, and the limpopo national park in mozambique. recently, four research ‘supersites’ have been identified and established in knp, with each of these supersites representing catenas with distinct geological, ecological and climatic features (rughöft et al. 2016; smit et al. 2013). such a site represents an ideal opportunity to determine if fungal communities will be uniform over niches and soil conditions within the system, or if the catena will harbour a complex and differing fungal community. the study focuses on characterisation of the fungal communities from the root ectoand endo-environments of one plant species which occurred within the different soils and soil conditions of the grazing lawn zone and the neighbouring hillside zone of the catena (theron, van aardt & du preez 2020; vermeulen, casson & swart 2020). whether different plants in the catena will have similar specialised rhizosphere fungal communities was also investigated, as it would add to the complexity to properly conserve and manage the catena site should variations be observed. materials and methods study site and sampling the study was conducted at the southern granite supersite catena close to the stevenson-hamilton memorial (smit et al. 2013) in april 2017 (autumn), during which time it still rained frequently. sida cordifolia (malvacae, malvales) or flannel weed is an invasive herbaceous plant (jain et al. 2011) and grows across the grazing lawn zone (sterkspruit soil type, high clay content, mean sodium concentration 3802 mg/kg, ph mean 6.4) and hillside zone (clovelly soil type, high sand percentage [90%], mean sodium concentration 1062 mg/kg, ph mean 5.85) of the catena (sandoval-denis, swart & crous 2019; theron et al. 2020; vermeulen et al. 2020). this plant was selected for sampling for rhizosphere and endophytic fungal communities from the two zones (vermeulen et al. 2020). two additional native herbaceous species, namely, melhania acuminata (also malvaceae) and kyphocarpa angustifolia (amaranthaceae, caryophyllales) were collected from the hillside zone for rhizosphere comparisons (vermeulen et al. 2020). ten plants from the grazing lawn and 10 plants from the hillside zones were excavated for each plant species, respectively. the plants occurred randomly throughout the two sites. the plant and soil samples were transferred aseptically to paper bags, and kept cold while being transported to the laboratory. illumina sequencing and analysis the plant roots were submerged in sterile water and shaken to dislodge soil from the rhizosphere that was attached to the root surfaces. consequently, the roots were surface sterilised in a standard sodium hypochlorite (3%), ethanol (70%) and sterile distilled water series. the roots were cut into small pieces, freeze dried and pulverised with 2-mm-diameter metal beads in a qiagen tissuelyser ii cell disrupter (qiagen, germantown, usa). the soil suspension was dried so that only the soils remained, and were subsequently also lysed. for each sample, 0.1 g of the pulverised plant and soil samples were used. the genomic deoxynuceic acid (dna) from the soil suspension and sterile root samples was extracted by using the soil and plant ii nucleospin® kits (machery-nagel gmbh and co kg, duren, germany), respectively, as per the user manual. the dna concentrations, determined by using a nanodrop lite spectrophotometer (thermo scientific, usa), were all adjusted to 10 ng/µl using sterile water. the internal transcribed spacer 2 (its2) region was targeted for the illumina sequencing study using primers its3 and its4 fitted with illumina adapters following the protocol described in tonjock et al. (2019). generated amplicons were pooled on the basis of plant, zone, rhizosphere or root endophyte categories, normalised and sequenced using an illumina miseq (illumina, usa) based on the procedure described by tonjock et al. (2019), using miseq v3 reagents and as paired 300 bp reads, at the next generation sequencing facility at the department of health sciences, university of the free state, south africa. sequences were run through a quality control pipeline as described by tonjock et al. (2019) with the final step being taxonomic assignment against the unite database at 99% similarity (nilsson et al. 2011) and did not differ from results obtained at 97% similarity. in this pipeline, chimeric sequences were identified, using usearch 6.1.544 (edgar 2010) as the chimera detection method (edgar 2010), and filtered out of the quality trimmed reads by using identify_chimeric_seqs.py and filter_fasta.py commands, respectively. taxonomic assignments were verified according to current taxonomic classifications. molecular operational taxonomic unit (motu) tables were normalised. prior to the analysis, the otu-table was normalised using normalize_table.py in qiime with the css normalisation option (paulson et al. 2013) and all analyses were carried out using r (www.r-project). shared and unique motus between communities were plotted using the venn function in gplots (https://cran.r-project.org/package=gplots). the dna reads were deposited in genbank under the bioproject accession no. prjna624016 (accession numbers: s. cordifolia roots grazing lawn samn14564480, s. cordifolia roots slope samn14564481, s. cordifolia rhizosphere grazing lawn samn14564484, s. cordifolia rhizosphere slope samn14564483, m. acuminata rhizosphere slope samn14564482, k. angustifolia rhizosphere slope samn14564485). ethical considerations ethical approval for the multidisciplinary project as a whole was obtained from the interfaculty animal ethics committee at the university of the free state (ufs-aed2019/0121). sanparks permit numbers for collection of soil for lab analyses and vegetation for identification purposes are, respectively, sk069, sk2095 and sk054. results illumina sequencing and analysis after quality control analyses, a substantial number of reads remained (table 1). the majority of the motus that could be assigned with names in the pipeline (table 2, figures 1–6) belonged to ascomycetes (41 of 54), while the remainder 12 motus belonged to basidiomycetes and one to zygomycete. no other types of fungi were detected by using the particular experimental design, dna extraction protocol and polymerase chain reaction (pcr) setup. a large percentage of motus that could not be named by the pipeline were detected. the ascomycete and basidiomycete groups included diverse orders. the number of named motus per niche (namely, endophytes vs. rhizosphere communities of s. cordifolia in two zones, and rhizosphere communities of different plant species in one zone) varied between 16 and 26, with the lowest number being that of the grazing lawn endophytes of s. cordifolia and the highest being that of the rhizosphere community of k. angustifolia. figure 1: percentage relative abundance of fungi (up to genus level) with a frequency higher than 1% at 97% level of sequence similarity. results were obtained for endophytes and rhizosphere fungi sida cordifolia growing in the grazing lawn and slope zones, from the grazing lawn and slope zones, respectively. figure 2: percentage relative abundance of fungi (up to genus level) with a frequency lower than 1% at 97% level of sequence similarity. results were obtained for endophytes and rhizosphere fungi from sida cordifolia growing in the grazing lawn and slope zones, respectively. figure 3: venn diagram of the fungal molecular operational taxonomic units obtained from the outside (rhizosphere) and inside (endophytes) of sida cordifolia roots growing on the grazing lawn and slope zones, respectively. figure 4: percentage relative abundance of fungi (up to genus level) with a frequency higher than 1% at 97% level of sequence similarity. results were obtained for rhizosphere fungi from three plant species growing in the slope zone. figure 5: percentage relative abundance of fungi (up to genus level) with a frequency lower than 1% at 97% level of sequence similarity. results were obtained for rhizosphere fungi from three plant species growing in the slope zone. figure 6: venn diagram of the fungal molecular operational taxonomic units obtained from the rhizosphere of three plant species growing in the slope zone. table 1: summary of number of reads before and after quality control analyses. four motus occurred in all of the niches or were present in both types of soils, but showed interesting shifts in frequency (table 2, figures 1, 2, 4 and 5). an motu with the assigned name of alternaria was dominant in all of the rhizosphere communities, but was present only in low numbers as a root endophyte. similarly, fusarium was present with frequency >1% as an endophyte and in all of the rhizosphere samples, but shifted to a lower presence in the s. cordifolia rhizosphere in the slope vegetation. rhodotorula was always present in frequencies higher than 1% as an endophyte in s. cordifolia in both soils, but had low frequencies in the rhizospheres of all plants. interestingly, in s. cordifolia rhizosphere samples, it was present as a root endophyte in the slope but not present in the slope rhizosphere. the only motu present across all niches and soils, but in low numbers, was one presented as microdiplodia. table 2: summary of the most prominent molecular operational taxonomic units that were assigned names from the rhizospheres and roots of three plant species in two zones of the catena. for the rhizosphere communities, there was a high degree of motu overlap, and also numbers of unique motus (table 2, figures 1–6). fifty motus were shared between rhizosphere soils of s. cordifolia in the grazing lawn and slope conditions, while there were 57 unique motus in the sodic grazing lawn site, and 63 in the non-sodic slope site (figure 3). for the comparisons between the rhizospheres of the three plant species in the slope site, 144 motus were shared between all, while the native m. acuminata and k. angustifolia plants had 183 and 178 motus, respectively; unique motus were compared to the 63 unique motus of the invasive s. cordifolia (figure 6). in total, m. acuminata had 482 motus, k. angustifolia had 451 and s. cordifolia had 291 motus. when comparing the number of motus of the s. cordifolia rhizospheres and root endophyte samples in the grazing lawn and slope sites (figure 3), no motu was shared between all the rhizosphere and root samples. very low numbers were shared between the rhizosphere and root collections from both sites. however, more motus were shared within the rhizosphere samples for the two sites, as well as between the root samples, as previously stated. the rhizosphere samples had almost half of the number of motus that the root samples had. the most dominant motus in the rhizosphere (>1% frequency) for all the plant species and in both zones (table 2, figures 2 and 4) were assigned as alternaria and boletus. fusarium was also dominant in the rhizospheres of k. angustifolia, m. acuminata and for s. cordifolia in the grazing lawn zone, while it was less dominant in the hill slope zone (table 2, figures 2 and 4). k. angustifolia had the most diverse rhizosphere community. molecular operational taxonomic units (<1% frequency) that were shared included candida, lodderomyces, agaricus, amanita and the plant pathogenic genus ganoderma. the remainder of the motus occurred only in one plant species or zone, or were shared between species or the endophyte and rhizosphere niches. endophyte profiles from s. cordifolia roots were quite different from the rhizophere profiles of the same plant (table 2, figures 2 and 3). profiles of the dominant motus (>1%) included different fungi than those present in the rhizosphere, while the endophyte patterns between the grazing lawn and slope vegetation zones were also different. however, motus assigned as humicola, fusarium and rhodotorula were present in both zones at a frequency >1%. some motus were only found as endophytes, including those assigned as pestalotiopsis and phaeoacremonium, members of pleosporales (curvularia, fusculina) and myrmecridium. in most cases, endophyte motus could be found in the rhizosphere, including those of other plants, albeit at different levels of frequency. microdiplia was the only motu found in similar levels as an endophyte and in the rhizospheres. comparisons between the rhizosphere and root endophyte communities in the soils of the grazing lawn and slope vegetation zones showed distinct differences in presence or absence, or shifts in frequency (table 2, figures 2 and 4). out of 27 detected motus, 19 were unique to the rhizosphere soils, while only four were unique to the internal tissues of the roots. some motus, such as alternaria, boletus, microdiplodia, chaetomium, penicillium, agaricus and amanita, occurred in the rhizosphere of both zones at similar levels, while there were shifts in the frequencies of others, such as those of fusarium. for the endophyte community, fusarium and humicola were present in roots of both zones at frequencies >1%. humicola were absent from all rhizosphere soils but were found in the rhizosphere of s. cordifolia in the grazing lawn zone. microdiplodia, aureobasidium, phoma, curvularia and fusculina were present at frequencies <1% in roots from both zones. lasiodiplodia and botryosphaeria (both in the botryosphaeriaceae) were only present in the hill slope zone and at frequencies higher than 1%, while other motus with frequencies <1% were also unique to a zone. molecular operational taxonomic units showing shifts in frequency, included pestalotiopsis that was more frequent in the hill slope zone, and exophiala and rhyzopycnis that were more frequent in the grazing lawn roots. discussion a catena system, such as the stevenson–hamilton granite supersite in the knp, represents an ideal system to study the fungal communities that are associated with plant roots in different soil types and conditions and from different plant species. within only approximately 500 m, a range of soil types and conditions are found that was also reflected by differences in the plant community (theron et al. 2020). similarly, marked differences in the fungal communities existed in the plant–soil interface. unique motus and shifts in frequencies were observed between the inside (endophytes) and outside (rhizosphere) communities of s. cordifolia roots, and also between two different soils. furthermore, differences were observed between the rhizosphere soils of three different plant species in one zone, although the dominant motus were relatively similar. fungi specialised to occur in different substrates, for example, soil compared to plant tissues, are different because they require different dissemination strategies and may also have different functional roles (bayman et al. 1997; carroll 1988; hardoim et al. 2015, moore et al. 2011; talbot et al. 2014; yang et al. 2018). the root rhizosphere is also known to usually harbour selected species either from the general soil environment or the root, and are often specialised according to the plant species (berlanas et al. 2019; mohanram & kumar 2019). nonetheless, levels of overlap do occur (tedersoo et al. 2016). our results thus reflected previous literature and represent the first such study for the knp. it also represents the first evidence regarding how fungal communities differ within a catena system. alternaria and fusarium include species that are endophytic, saprobic in soils and pathogenic or beneficial to plants, and that occur in numerous habitats (leslie & summerell 2006; woudenberg et al. 2013). such cosmopolitan fungi can be dominant or omnipresent in communities because of their ability to easily colonise or disperse, as opposed to more specialised fungi that may not always have a rapid growth rate and that have evolved with their hosts or niche (vujanovic et al. 2006). while such dominant motus of alternaria and fusarium were detected, shifts in their frequencies occurred between different substrates (e.g. alternaria was dominant in rhizospheres but less so as an endophyte) and soils (fusarium was one of the dominant groups in all collections except in the slope rhizosphere of s. cordifolia; fusarium was also almost five times as dominant as a root endophyte of s. cordifolia in the grazing lawn area compared to the slope). plants that were selected were all healthy, and the presence of alternaria within plants was latent and in low numbers, and the higher abundance of fusarium could be because of non-pathogenic fusarium species. an interesting result was that other fungi that can occur both in plants and soils, for example, curvularia, penicillium and aureobasidium (domsch, anderson & gams 1980; haas et al. 2016) were selectively detected, and others were not detected at all, such as cladosporiod fungi. knowledge regarding the ecological functions of fungi associated with different soil types and those associated with plants is virtually non-existent for natural and pristine ecosystems in south africa. the ecological functions of these fungi can only be guessed from the existing literature. intensive studies are needed to elucidate the functions of the various fungi characterised in this study, and how changes will impact them and their associated hosts. the more rare fungal groups could be more specialised and sensitive, but without more surveys it will be difficult to identify them as important in the ecosystem or if they are promoting plant and soil health (frac et al. 2018), as opposed to just being able to infect at a low frequency. sida cordifolia is not native to south africa, but a declared invasive weed (morris, witkowski & coetee 2011). it was chosen because of its ability to grow across the different soils of the catena system. nonetheless, the communities involved with the rhizospheres and roots in the two sites of the catena showed telling variation. the invasive plant s. cordifolia appeared to attract only half of the number of motus present in its rhizosphere compared to those in the native relative m. acuminata, and the other native plant k. angustifolia. however, more extensive sampling will have to be performed to really prove this statistically. various factors could possibly explain the differences observed between s. cordifolia and the two native herbs. certain plants have been shown to have a stronger rhizosphere effect than others (gomes et al. 2003), and what has been observed in this study could merely be such an effect. no studies have been conducted for comparing the microbial rhizosphere community of invasive plants, or even non-native plants, with those of native plants in the same area. nonetheless, it could be hypothesised that the ability to attract a functional rhizosphere microbial community that benefits the plant will aid the establishment or invasiveness of a plant (coats & rumpho 2014), similar to what has been found with other types of microbes such as mycorrhiza (nuñez & dickie 2014; rodríguez-echeverría et al. 2009). furthermore, it has been shown that some invasive plants can change the soil microbiome around their roots to their benefit (dawkins & esiobu 2017; policelli et al. 2019). what has also been found in other groups of plant-associated microbes, such as endophytes, was that the endophytic community of non-native plants were composed of endophytes with wide host ranges and large geographical locations (cosmopolitan fungi), while the native plants had more specialised and diverse endophytes including those with narrow host ranges (clay et al. 2016; hoffman & arnold 2008; newcombe et al. 2009). whether the native plant species in the knp have closer evolved endophytic and rhizosphere fungi compared to introduced plants will have to be studied further. many of the motus detected in all the samples were assigned names of genera known to include plant pathogens, such as fusarium and alternaria. however, these genera include a wide diversity of species with diverse ecological roles and life strategies, including pathogens, non-pathogens or latent pathogens (pathogens that can hide as endophytes) and endophytes. it is difficult to assess if the dnas detected from the samples represented pathogenic species or not based on the limited sequence data, especially because the its regions commonly used in next generation sequencing (ngs) studies, are known to be incapable of differentiating between species of these genera (schoch et al. 2012). an interesting result is that members of other important pathogen groups have been detected. molecular operational taxonomic units in the botryosphaeriaceae and an motu from the teratosphaeriaceae were detected as root endophytes. these fungi are not recorded as soil fungi and are mostly known from aerial tissues of woody plants, where they have latent pathogenic life stages before causing various disease symptoms (crous, wingfield & groenewald 2009; finlay & clay 2007; marsberg et al. 2014; pillay et al. 2013; sieber 2007; slippers & wingfield 2007). while their presence in the rhizosphere could be attributed to the presence of spores, the prominence of especially two motus of the botryosphaeriaceae (botryosphaeria and lasiodiplodia) from the roots of the herbaceous plant s. cordifolia is interesting. an motu that was prominent in all of the rhizosphere samples had the assigned name of boletus (boletaceae). the motu was not present in the root endophyte samples. members of boletaceae are important ecto-mycorrhizal partners of plant roots and can easily be seen by the production of mushroom-like fruiting bodies (goldman & gryzenhout 2019). south africa does not have reported indigenous ectomycorrhizal fungi, while the only indigenous bolete species is phaeogyroporus sudanicus or the bushveld bolete (goldman & gryzenhout 2019; gryzenhout 2010; tonjock et al. 2020; van der westhuizen & eicker 1994). while the fungus is known to be ectomycorrhizal (thoen & ducousso 1990, as phlebopus sudanicus), its host range throughout south africa, including the knp, has not really been fully established. this bolete has been observed in the knp by the first author, and it is thus possible that the motu could represent this species, especially so because there are no phaeogyroporus sequences currently present in the unite database (https://unite.ut.ee/). the absence of the motu from inside surface sterilised roots could indicate that it does not have ectomycorrhizal associations with the three plant species. ecto-mycorrhizal fungi such as boletes are more easily observed than other groups of mycorrhiza because they produce visible fruiting bodies above soil. the detection of a bolete sequence raises the question if other native ectomycorrhizal partners to indigenous plants in the knp do not in actual effect exist. this is especially so because extensive surveys for the presence of such fungi have not yet been conducted in the knp. furthermore, species of some plant genera that are known to have ectomycorrhizal partners, such as albizia, terminalia and combretum in neighbouring countries such as zimbabwe (tsamba et al. 2015) and even up to central africa (eneke, njoh & egbe 2018; härkönen, niemelä & mwasumbi 2003; härkönen et al. 2015), occur in the knp. related species such as colophospermum mopane occur in the knp, which belong to detariodeae (fabaceae). this family includes keystone genera known to have ecto-mycorrhizal partners in the iconic and widespread miombo woodlands that are spread from zimbabwe further north into central africa (palgrave 2015). in fact, members of these named genera are present in the catena (theron et al. 2020). studying these special fungi is vital because they play an integral role in the health and resilience of their associated plants, and should they disappear for some reason, it will have grave consequences on plant survival in the knp. other macrofungal motus (fungi that can be seen without the need of a microscope) that were detected from all of the rhizosphere samples included the mushroom genera agaricus and amanita. agaricus is known to be saprophytic while many species of amanita are known ecto-mycorrhizal plant partners (goldman & gryzenhout 2019). however, certain amanita species, including south african native species such as a. veldiei, are saprophytes (goldman & gryzenhout 2019). an motu assigned as ganoderma was present in the rhizosphere samples of the three plant species from the slope. ganoderma species are common throughout south africa and include wood rotting bracket fungi that often are pathogens of plants (goldman & gryzenhout 2019; tchoumi et al. 2019). a number of species of these genera occur in south africa (tonjock et al. 2020), and conducting more detailed surveys in the knp to ascertain what species occur in the park will prove to be useful. for the data analysis pipeline used in this study, the unite database (nilsson et al. 2019) was used for taxonomic assignment. this database is better curated than general databases such as genbank, and widely used for fungal environmental sequencing (nilsson et al. 2019). however, taxonomic assignments must always be verified because the assignment process may not truly reflect the correct taxonomic identity even at the genus level, or may not reflect current taxonomies (tonjock et al. 2019). the its region is also known to not always distinguish species in some genera, although it is still widely used as the barcode and minibarcode for fungi (schoch et al. 2012). in practice, complicated taxonomic groups such as phoma and epicoccum cannot be distinguished based only on its data in routine its-based environmental sequencing studies (tonjock et al. 2019). the conclusions from results of this study must thus be made with care considering that even generic names may not be accurate. however, in some cases, associations can be made if members of a particular family or genus have similar ecological attributes, such as the boletaceae. in other cases, such as for fusarium (leslie & summerell 2006), different species often have certain ecological roles (e.g. pathogen or latent pathogen or non-pathogenic, or pathogen with saprophytic survival phase). it is thus possible that an motu assigned as fusarium (or previously used sexual names such as gibberella and haematonectria) from different niches and substrates may in fact represent different fusarium species with different ecological roles. conclusion results from this study revealed a complex fungal community associated with the plant–soil interface within a catena system. this is despite the fact that the two sites occurred within a 500 m area. results from other studies showed the effect of the different geological and hydrological processes on the plant and animal communities, while the biological communities also affect each other (janecke 2020; janecke & bolton 2020; theron et al. 2020; vermeulen et al. 2020). this study also showed the need to include fungi in studying these interactions more thoroughly. fungi provide fundamental ecosystem services as key saprophytes, plant degraders, pathogens and those providing benefits to the plant as mycorrhiza, endophytes or rhizosphere partners. the complexity of fungal communities differing already in a small component of the entire ecosystem of the catena indicates that such a system with its differing communities will be sensitive to change and disturbances, both in the short and long runs. similar results have also been obtained by looking at the bacterial communities (vermeulen et al. 2020). this is despite the fact that the overall animal and plant communities may appear more uniform and thus do not indicate the importance to specifically also define differences of microbial communities on a micro-ecological scale. the same will be true for fungal communities associated with other plant species and soil types of the knp. acknowledgements the authors thank the university of the free state (ufs) strategic research fund for providing funding for the multidisciplinary research. they are also thankful to the sanparks scientific services for the assistance provided during field sampling. they also acknowledge the leadership role of dr beanelri janecke along with the rest of the research team for providing insights. prof. wijnand swart (university of the free state) is thanked for the choice of plant species in the catena system and for providing the root samples. mr paul kgoare (university of the free state) assisted with sequencing of the samples. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions the authors directly participated in the study design, execution and interpretation of the research. funding information the ufs strategic research fund is acknowledged for funding of the multidisciplinary research. data availability data are available from the corresponding author, on request. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references agrios, g.n., 2005, plant pathology, 5th edn., elsevier academic press, london. archibald, s., 2007, ‘african grazing lawns: how fire, rainfall, and grazer numbers interact to affect grass community states’, journal of wildlife management 22(2), 492–500. https://doi.org/10.2193/2007-045 aylor, d.e., 2003, ‘spread of plant disease on a continental scale: role of aerial dispersal of pathogens’, ecology 84(8), 1989–1997. https://doi.org/10.1890/01-0619 bayman, p., lebron, l.l., tremblay, r.l. & lodge, d.j., 1997, ‘endophytic fungi in roots and leaves of lepanthes (orchidaceae)’, new phytologist 135(1), 143–149. https://doi.org/10.1046/j.1469-8137.1997.00618.x berlanas, c., berbegal, m., elena, g., laidani, m., cibriain, j.f., sagues, a. & gramaje, d., 2019, ‘the fungal and bacterial rhizosphere microbiome associated with grapevine rootstock genotypes in mature and young vineyards’, frontiers in microbiology 10, 1142. https://doi.org/10.3389/fmicb.2019.01142 brady, n.c. & weil, r.r., 2002, the nature and properties of soils, 13th edn., prentice hall, england cliffs, nj. bridge, p. & spooner, b., 2001, ‘soil fungi: diversity and detection’, plant and soil 232, 147–154. https://doi.org/10.1023/a:1010346305799 carroll, g., 1988, ‘fungal endophytes in stems and leaves: from latent pathogen to mutualistic symbiont’, ecology 69(1), 2–9. https://doi.org/10.2307/1943154 carruthers, j., 2017, national park science: a century of research in south africa, cambridge university press, cambridge. clay, k., shearin, z.r.c., bourke, k.a. & kowalski, k.p., 2016, ‘diversity of fungal endophytes in non-native phragmites australis in the great lakes’, biological invasions 18, 2703–2716. https://doi.org/10.1007/s10530-016-1137-y coats, v.c. & rumpho, m.e., 2014, ‘the rhizosphere microbiota of plant invaders: an overview of recent advances in the microbiomics of invasive plants’, frontiers in microbiology 5, 1–10. https://doi.org/10.3389/fmicb.2014.00368 crous, p.w., wingfield, m.j. & groenewald, j.z., 2009, ‘niche sharing reflects a poorly understood biodiversity phenomenon’, persoonia 22, 83–94. https://doi.org/10.3767/003158509x439364 dawkins, k. & esiobu, n., 2017, ‘arbuscular and ectomycorrhizal fungi associated with the invasive brazilian pepper tree (schinus terebinthifolius) and two native plants in south florida’, frontiers in microbiology 8, 665. https://doi.org/10.3389/fmicb.2017.00665 domsch, k.h., anderson, t. & gams, w., 1980, compendium of soil fungi, academic press ltd., london. edgar, r.c., 2010, ‘search and clustering orders of magnitude faster than blast’, bioinformatics 26(1), 2460–2461. https://doi.org/10.1093/bioinformatics/btq461 finlay, r.d. & clay, k., 2007, ‘fungal endophytes in forests, woody plants and grassland ecosystems: diversity, functional ecology and evolution’, fungal biology reviews 21(2–3), 49–50. https://doi.org/10.1016/j.fbr.2007.06.001 frac, m., hannula, s.e., belka, m. & jedryczka, m., 2018, ‘fungal biodiversity and their role in soil health’, frontiers in microbiology 9, 707. https://doi.org/10.3389/fmicb.2018.00707 glynou, k., ali, t., buch, a., kia, s.h., ploch, s., xia, x. et al., 2016, ‘the local environment determines the assembly of root endophytic fungi at a continental scale’, environmental microbiology 18(8), 2418–2434. https://doi.org/10.1111/1462-2920.13112 goldman, g.b. & gryzenhout, m., 2019, field guide to mushrooms and other fungi of south africa, penguin random house struik, cape town. gomes, n.c.m., fagbola, o., costa, r., rumjanek, n.m., buchner, a., mendona-hagler, l. et al., 2003, ‘dynamics of fungal communities in bulk and maize rhizosphere soil in the tropics’, applied and environmental microbiology 69(7), 3758–3766. https://doi.org/10.1128/aem.69.7.3758-3766.2003 gryzenhout, m., 2010, pocket guide: mushrooms of south africa, randomhouse-struik, cape town. haas, d., lesch, s., buzina, w., galler, h., gutschi, a.m., habib, j. et al., 2016, ‘culturable fungi in potting soils and compost’, medical mycology 54(8), 825–834. https://doi.org/10.1093/mmy/myw047 hardoim, p.r., van overbeek, l.s., berg, g., pirttila, a.m., compant, s., campisano, a. et al., 2015, ‘the hidden world within plants: ecological and evolutionary considerations for defining functioning of microbial endophytes’, microbiology and molecular biology reviews 79(3), 293–320. https://doi.org/10.1128/mmbr.00050-14 hargreaves, s.k., williams, r.j. & hofmockel, k.s., 2015, ‘environmental filtering of microbial communities in agricultural soil shifts with crop growth’, plos one 10(7), e0134345. https://doi.org/10.1371/journal.pone.0134345 härkönen, m., niemelä, t., mbindo, k., kotiranta, h. & piearce, g., 2015, zambian mushrooms and mycology, botanical museum, finnish museum of natural history, helizinki. härkönen, m., niemelä, t. & mwasumbi, l., 2003, tanzanian mushrooms: edible, harmful and fungi, botanical museum, finnish museum of natural history, helizinki. havlicek, e. & mitchell, e.a.d., 2014, ‘interactions in soil: promoting plant growth, biodiversity, community and ecosystems’, in j. dighton & j.a. krumins (eds.), soils supporting biodiversity, pp. 27–58, springer, dordrecht. hempson, g.p., archibald, s., bond, w.j., ellis, r.p., grant, c.c., kruger, f.j. et al., 2015, ‘ecology of grazing lawns in africa’, biological reviews of the cambridge philosophical society 90(3), 979–994. https://doi.org/10.1111/brv.12145 hiltner, l., 1904, ‘über neuere erfahrungen und probleme auf dem gebiet der bodenbakteriologie und unter besonderer berucksichtigung der gründüngung und brache’, arbeiten der deutschen landwirtschaftlichen gesellschaf 98, 59–78. hoffman, m.t. & arnold, a.e., 2008, ‘geographic locality and host identity shape fungal endophyte communities in cupressaceous trees’, mycological research 118(3), 331–344. https://doi.org/10.1016/j.mycres.2007.10.014 jain, a., choubey, s., singour, p.k., rajak, h. & pawar, r.s., 2011, ‘sida cordifolia (linn) – an overview’, journal of applied pharmaceutical science 1, 23–31. janecke, b.b., 2020, ‘vegetation structure and spatial heterogeneity in the granite supersite, kruger national park’, koedoe 62(2), a1591. https://doi.org/10.4102/koedoe.v62i2.1591 janecke, b.b. & bolton, j.g., 2020, ‘variation in mammal diversity and habitat affect heterogeneity and processes of a granite catena’, koedoe 62(2), a1592. https://doi.org/10.4102/koedoe.v62i2.1592 khare, e., mishra, j. & arora, n.k., 2018, ‘multifaceted interactions between endophytes and plant: developments and prospects’, frontiers in microbiology 9, 2732. https://doi.org/10.3389/fmicb.2018.02732 lareen, a., burton, f. & schafer, p., 2016, ‘plant root-microbe communication in shaping root microbiomes’, plant molecular biology 90, 575–587. https://doi.org/10.1007/s11103-015-0417-8 leslie, j.f. & summerell, b.a., 2006, the fusarium laboratory manual, blackwell, ames, ia. marsberg, a., slippers, b., wingfield, m.j. & gryzenhout m., 2014, ‘endophyte isolations from syzygium cordatum and a eucalyptus grandis clone (myrtaceae) reveal new host and geographical reports for the mycosphaerellaceae and teratosphaeriaceae’, australasian journal of plant pathology 43, 503–512. https://doi.org/10.1007/s13313-014-0290-y mcnaughton, s.j., 1984, ‘grazing lawns: animals in herds, plant form, and coevolution’, the american naturalist 124(6), 863–886. https://doi.org/10.1086/284321 mohanram, s. & kumar, p., 2019, ‘rhizosphere microbiome: revisiting the synergy of plant-microbe interactions’, annual microbiology 69, 307–320. https://doi.org/10.1007/s13213-019-01448-9 mommer, l., kirkegaard, j. & van ruijven, j., 2016, ‘root–root interactions: towards a rhizosphere framework’, trends in plant sciences 21(3), 209–217. https://doi.org/10.1016/j.tplants.2016.01.009 moore, d.m., robson, g.d., trinci, a.p.j., 2011, 21st century guidebook to fungi, cambridge university press, cambridge. morris, t.l., witkowski, e.t.f. & coetee, j.a., 2011, ‘initial response of riparian plant community structure to clearing of invasive alien plants in kruger national park, south africa’, south african journal of botany 74(3), 485–494. https://doi.org/10.1016/j.sajb.2008.01.177 newcombe, g., shipunov, a., eigenbrode, s.d., raghavendra, a.k.h., ding, h., anderson, c.l. et al., 2009, ‘endophytes influence protection and growth of an invasive plant’, communicative and integrative biology 2(1), 29–31. https://doi.org/10.4161/cib.2.1.7393 nilsson, r., tedersoo, l., lindahl, b.d., kjoller, r., carlsen, t., quince, c. et al., 2011, ‘towards standardization of the description and publication of next generation sequencing datasets of fungal comm-unities’, new phytologist 191(2), 314–318. https://doi.org/10.1111/j.1469-8137.2011.03755.x nilsson, r.h., larsson, k.h., taylor, a.f.s., bengtsson-palme, j., jeppesen, t.s., schigel, d. et al., 2019, ‘the unite database for molecular identification of fungi: handling dark taxa and parallel taxonomic classifications’, nucleic acids research 47(d1), d259–d264. https://doi.org/10.1093/nar/gky1022 nuñez, m.a. & dickie, i.a., 2014, ‘invasive belowground mutualists of woody plants’, biological invasions 16, 645–661. https://doi.org/10.1007/s10530-013-0612-y palgrave, k.c., 2015, palgrave’s trees of southern africa, struik nature, cape town. paulson, j.n., stine, o.c., bravo, h.c. & pop, m., 2013, ‘robust methods for differential abundance analysis in marker gene surveys’, nature methods 10, 1200–1202. https://doi.org/10.1038/nmeth.2658 pillay, k., slippers, b., wingfield, m.j. & gryzenhout, m., 2013, ‘diversity and distribution of co-infecting botryosphaeriaceae from eucalyptus grandis and syzygium cordatum in south africa’, south african journal of botany 84, 38–43. https://doi.org/10.1016/j.sajb.2012.09.003 policelli, n., bruns, t.d., vilgalys, r. & nuñez, m.a., 2019, ‘suilloid fungi as global drivers of pine invasions’, new phytologist 222(2), 714–725. https://doi.org/10.1111/nph.15660 richardson, m.j., 2001, ‘diversity and occurrence of coprophilous fungi’, mycological research 105(4), 387–402. https://doi.org/10.1017/s0953756201003884 rodriguez, r.j., white, j.f., jr., arnold, a.e. & redman, r.s., 2009, ‘fungal endophytes: diversity and functional roles’, new phytoogistl 182(2), 314–330. https://doi.org/10.1111/j.1469-8137.2009.02773.x rodríguez-echeverría, s., crisóstomo, j.a., nabais, c. & freitas, a., 2009, ‘belowground mutualists and the invasive ability of acacia longifolia in coastal dunes of portugal’, biological invasions 11, 651–661. https://doi.org/10.1007/s10530-008-9280-8 rousk, j., baath, e., brookes, p.c., lauber, c.l., lozupone, c., caporaso, j.g. et al., 2010, ‘soil bacterial and fungal communities across a ph gradient in an arable soil’, isme journal 4, 1340–1351. https://doi.org/10.1038/ismej.2010.58 rughöft, s., herrmann, m., lazar, c.s., cesarz, s., levick, s.r., trumbore, s.e. & küsel, k., 2016, ‘community composition and abundance of bacterial, archaeal and nitrifying populations in savanna soils on contrasting bedrock material in kruger national park, south africa’, frontiers in microbiology 7, 1638. sabra, m., aboulnasr, a., franken, p., perreca, e., wright, l.p. & camehl, i., 2018, ‘beneficial root endophytic fungi increase growth and quality parameters of sweet basil in heavy metal contaminated soil’, frontiers in plant sciences 9, 1726. https://doi.org/10.3389/fpls.2018.01726 sandoval-denis, m., swart, w.j. & crous, p.w., 2018, ‘new fusarium species from the kruger national park, south africa’, mycokeys 34, 63–92. https://doi.org/10.3897/mycokeys.34.25974 sasse, j., martinoia, e. & northen, t., 2018, ‘feed your friends: do plant exudates shape the root microbiome?’, trends in plant sciences 23(1), 25–41. https://doi.org/10.1016/j.tplants.2017.09.003 schippers, b., bakker, a.w. & bakker, p.a.h.m., 1987, ‘interactions of deleterious and beneficial rhizosphere microorganisms and the effect of cropping practices’, annual review of phytopathology 25, 339–358. https://doi.org/10.1146/annurev.py.25.090187.002011 schoch, c.l., seifert, k.a., huhndorf, s., robert, v., spouge, j.l., levesque, c.a. et al., 2012, ‘nuclear ribosomal internal transcribed spacer (its) region as a universal dna barcode marker for fungi’, proceedings of the national academy of science, usa 109, 6241–6246. setala, h. & mclean, m.a., 2004, ‘decomposition rate of organic substrates in relation to the species diversity of soil saprophytic fungi’, oecologia 139, 98–107. https://doi.org/10.1007/s00442-003-1478-y sieber, t.n., 2007, ‘endophytic fungi in forest trees: are they mutualists?’, fungal biology reviews 21(2–3), 75–89. https://doi.org/10.1016/j.fbr.2007.05.004 slippers, b & wingfield, m.j., 2007, ‘botryosphaeriaceae as endophytes and latent pathogens of woody plants: diversity, ecology and impact’, fungal biology reviews 21(2–3), 90–106. https://doi.org/10.1016/j.fbr.2007.06.002 smit, i.p.j., riddell, e.s., cullum, c. & petersen, r., 2013, ‘kruger national park research supersites: establishing long-term research sites for cross-disciplinary, multiscaled learning’, koedoe 55(1), art. #1107, 7 pages. https://doi.org/10.4102/koedoe.v55i1.1107 talbot, j.m., bruns, t.d., taylor, j.w., smith, d.p., branco, s., glassman, s.i. et al., 2014, ‘endemism and functional convergence across the north american soil mycobiome’, proceedings of the national academy of science, usa 111(17), 6341–6346. https://doi.org/10.1073/pnas.1402584111 taylor, d.l. & bruns, t.d., 1999, ‘community structure of ectomycorrhizal fungi in a pinus muricata forest: minimal overlap between the mature forest and resistant propagule communities’, molecular ecology 8(11), 1837–1850. https://doi.org/10.1046/j.1365-294x.1999.00773.x tchotet tchoumi, j.m., coetzee, m.p.a., rajchenberg, m. & roux, j., 2019, ‘taxonomy and species diversity of ganoderma species in the garden route national park of south africa inferred from morphology and multilocus phylogenies’, mycologia 111(5), 730–747. https://doi.org/10.1080/00275514.2019.1635387 tedersoo, l., bahram, m., cajthaml, t., polme, s., hiiesalu, i., anslan, s. et al., 2016, ‘tree diversity and species identity effects on soil fungi, protists and animals are context dependent’, the isme journal 10, 346–362. https://doi.org/10.1038/ismej.2015.116 theron, e.j., van aardt, a.c. & du preez, p.j., 2020, ‘vegetation distribution along a granite catena, southern kruger national park, south africa’, koedoe 62(2), a1588. https://doi.org/10.4102/koedoe.v62i2.1588 thoen, d. & ducousso, m., 1990, ‘mycorrhizal habit and sclerogenesis of phlebopus sudanicus (gyrodontaceae) in senegal’, agriculture, ecosystems & environment 28(1–4), 519–523. https://doi.org/10.1016/0167-8809(90)90091-q tonjock, r.k., cason, e.d., valverde, a., nyaga, m. & gryzenhout, m., 2019, ‘endophytic seed mycobiomes of six sorghum (sorghum bicolor) cultivars from commercial seedlots using an illumina sequencing approach’, mycosphere 10(1), 739–756. https://doi.org/10.5943/mycosphere/10/1/16 tonjock kinge, r., goldman, g., jacobs, a., nderitu, g. & gryzenhout, m., 2020, ‘the first checklist for macrofungi in south africa’, mycokeys 63, 1–40. https://doi.org/10.3897/mycokeys.63.36566 tsamba, j., kativu, s. & sithole-niang, i., 2015, ‘diversity and host associations of ectomycorrhizae fungi in fallow lands of the mid-zambezi valley area, zimbabwe’, transactions of the royal society of south africa 70, 71–77. van der westhuizen, g.c.a. & eicker, a., 1994, field guide: mushrooms of southern africa, struik, cape town. vermeulen, m., casson, e.d. & swart, w.k., 2020, ‘the rhizobiome of herbaceous plants in clovelly and sterkspruit soils of the stevenson-hamilton supersite’, koedoe 62(2), a1596. https://doi.org/10.4102/koedoe.v62i2.1596 verweij, r.j.t., verrelst, j., loth, p.e., heitkönig, i.m.a. & brunsting, a.m.h., 2006, ‘grazing lawns contribute to the subsistence of mesoherbivores on dystrophic savannas’, oikos 114(1), 108–116. https://doi.org/10.1111/j.2006.0030-1299.14209.x vujanovic, v., hamel, c., yergeau, e. & st-arnaud, m., 2006, ‘biodiversity and biogeography of fusarium species from northeastern north american asparagus fields based on microbiological and molecular approaches’, microbial ecology 51, 242–255. https://doi.org/10.1007/s00248-005-0046-x woudenberg, j.h., groenewald, j.z., binder, m. & crous, p.w., 2013, ‘alternaria redefined’, studies in mycology 75, 171–212. https://doi.org/10.3114/sim0015 yang, h., ye, w., ma, j., zeng, d., rong, z., xu, m. et al., 2018, ‘endophytic fungal communities associated with field-grown soybean roots and seeds in the huang-huai region of china’, peer journal 6, e4713. https://doi.org/10.7717/peerj.4713 vol. 50 no. 1 pp. 202 213koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research a floristic description of the afromontane fynbos communities on platberg, eastern free state, south africa abstract within the platberg area and the wider drakensberg region, the shrinking natural resources and the threat posed to biodiversity are of concern to conservation management and require an understanding of long-term ecological processes. the vegetation of platberg was investigated as part of an ecological survey to establish afromontane floristic links to the drakensberg as well as for the management of natural resources. from a twinspan classification, refined by the braun-blanquet method, four main plant communities were identified, which were subdivided into fynbos, wetland, a woody/ shrub community and grassland. a classification and description of the fynbos are presented in this article. the analysis showed the fynbos divided into two communities comprising four sub-communities and seven variants. the fynbos community had an average of 28.34 species per relevé, ranging from 14 to 54 species per sample plot. twenty-four endemic or near-endemic drakensberg alpine centre (dac) species and 22 exotic (alien-invasive) species were recorded. numerous floristic links with the dac, cape flora fynbos and grassland bioregions to the north and west were also found. the description of the fynbos plant communities can serve as a basis for the formulation of management plans for the area. keywords: free state province, drakensberg alpine centre (dac) of endemism, braun-blanquet method, afromontane fynbos communities, platberg indigenous biodiversity is under increasing pressure from numerous forces that threaten a loss of species and the depaupering of natural resources. platberg, in the eastern free state, overlooking the town of harrismith and is an island surrounded by plains of grassland. this inselberg represents a refuge for indigenous afromontane plants and animals. prior to this study, no extensive vegetation surveys had been undertaken on platberg. limited opportunistic floristic collections had been done in the mid-1960s by mrs ml jacobs, and some 50 relevés had been sampled by prof. hjt venter of the university of the free state (ufs) between 1975 and 1977 (vouchers from these collectors are housed at the geo potts herbarium (blfu) of the ufs). due to the proposed commercial development of parts of the platberg conservancy as well as the aims of the maluti drakensberg transfrontier project, it was therefore considered necessary to undertake a more detailed and extensive ecological survey to describe and classify the vegetation of platberg. these data will be used by the free state provincial department of tourism, economic and environmental affairs as part of its biological-diversity survey and conservation-management strategies. the data may also be incorporated into the maluti drakensberg transfrontier-project vegetation maps for future land use and conservation management. study area the town of harrismith is located at the foot of platberg (latitude s 29° 10’ and longitude e 28° 16’) on the n3 highway between johannesburg and durban (fig. 1). platberg’s altitude ranges from 1 900 m to 2 394 m; its surface area covers approximately 3 000 ha. the slopes are steep, with numerous vegetated gullies and boulder-scree slopes below the vertical cliffs, which are 20 to 45 m high. waterfalls cascade down the southern cliffs after rain. a permanent stream arising from the manmade dam (gibson dam) flows off the escarpment, cascading into a waterfall. woody patches of the genera leucosidea, buddleja, kiggelaria, polygala, heteromorpha and rhus shrubs as well as the only indigenous mountain bamboo, thamnocalamus tessellatus, grow along the bases of the cliffs, on sandstone of the clarens formation, in the gullies, on the scree slopes, in the mobile boulder beds, on the rocky ridges and as riparian vegetation in the cleft, in which the only road ascends steeply to the summit. locally, the pass is aptly named donkey pass for its steepness robert f. brand department of plant sciences, university of the free state south africa pieter j. du preez department of plant sciences university of the free state south africa leslie r. brown applied behavioural ecology and ecosystem research unit department of environmental sciences university of south africa south africa correspondence to: pieter j. du preez e-mail: dpreezpj.sci@ufs.ac.za postal address: department of plant sciences, ufs, po box 339, bloemfontein 202 2008 afromontane fynbos in platberg original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 202 213 and sheer sides. it bypasses a waterfall that drains the gibson dam to the east. from a distance, platberg appears to have a distinct flat top; once on the summit plateau, however, it is obvious that it has undulating, rolling, grass-covered plains instead. the vegetation of the plateau is dominated by grassland – with a few rocky ridges, sheet rock and rubble patches – by numerous seasonal wetlands and by a few permanent open playas (pans) on the far-western side of the platberg plateau. acocks (1952, 1988) classifies the vegetation of the drakensberg region into the following veld types: highveld sourveld (44a), themeda–festuca alpine veld (58), stormberg plateau sweetveld (59) and sclerophyllous bush types, and fynbos (69). as a result of the 1:1 500 000 scale of acocks’s map (1988), however, platberg does not appear as a distinct vegetation unit and is therefore included in the larger grassland biome. low and rebelo (1996) give a more detailed description of the vegetation of the region, indicating the following vegetation types: wet, cold highveld grassland (41), moist upland grassland (42), afro mountain grassland (45) and alti mountain grassland (46). platberg falls within the grassland biome, which generally contains short to tall sour grasses (acocks, 1988; low and rebelo, 1996). mucina and rutherford (2006) designate the study area as part of the drakensberg grassland bioregion. it also has elements of fynbos, false karroo (acocks, 1988) or succulent karoo (low and rebelo, 1996) as well as elements of temperate and transitional forest, specifically 44a highland sourveld (acocks, 1952, 1988) veld types. the most recent and detailed vegetation description presented by mucina and rutherford (2006) shows that the study area falls within the eastern free state sandy grassland (gm 4), which is part of the broader mesic highveld grassland bioregion. the platberg plateau is shown as an island that belongs to the drakensberg grassland bioregion. embedded within this bioregion are a series of isolated patches of fynbos, designated by mucina and rutherford (2006) as drakensberg-amathole afromontane fynbos (gd6). mucina and rutherford (2006) additionally recognise two structurally similar but floristically different afromontane fynbos shrublands in the drakensberg and surrounding area. a species of poor, fynbos-like shrubland with passerina montana is found on the edges of some of the high eastern free state inselbergs (mucina & rutherford, 2006). floristically, fynbos-like shrubland elements found on platberg show strong affinities with passerina montana, a sub-unit of the gd 8 fynbos-type vegetation, which, according to mucina and rutherford (2006) still has hundreds of patches to be mapped. geology platberg consists of layers of the karoo supergroup, which, stratagraphically, lies immediately below the volcanics of the drakensberg formation, the youngest unit of the stormberg group. the 1 900 m contour forms the start to the footslopes of platberg and is constituted by the aeolian and water-borne sediments of the clarens formation (du toit, 1954; loock, praekelt, van der westhuizen & welman, 1991; norman & whitfield, 1998; truswell, 1970). several dolerite dykes intersect platberg, with a close-set sequence of parallel dykes forming the woody/shrub community on the northern and southern slopes. a dolerite dyke is evident just below the gibson dam, shown in a borrow pit used for quarrying. numerous other dykes are visible on aerial photographs. these dyke swarms occur mostly on the flat terrain at lower altitudes, with only a few intersecting platberg itself. some of the dykes are also visible in plantation road-cuts on the southern side of platberg. topography and soils fig. 2 shows a terrain-form sketch of the study area. the steep slopes are composed of mudstone and mostly sandstone of the clarens formation, intersected by dolerite dykes, which results in benches, deep ravines and terraces. numerous sandstone and basalt boulders form scree and boulder slopes, resulting 203 figure 1 map of region original research brand, du preez & brown koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 202 213 in miscellaneous soils forms characteristic of the ib land type, specifically mispah (ms) and glenrosa (gs) soil forms (soil classification working group 1991 du preez, 1991; mucina and rutherford, 2006). there are affinities with fa land types, undulating rocky landscape dominated by shallow ms, and gs soil forms (land type survey staff, 1991; smit, bredenkamp and van rooyen, 1992). the summit plateau is composed of igneous rock. the soils are generally shallow (20 to 200 mm), with the deepest being < 500 mm. they are derived from basalt and are shown to have even proportions of course sand, fine sand, clay, silt and moderate to high organic-matter content. the main land type is ea (clay vertic, melanic or red diagnostic horizons) (erickson 1985, soil classification working group 1991; mucina and rutherford, 2006; smit et al., 1992; taylor, 1996), which has a clay content of 8 to 60%, depending on the soil series (land type survey staff, 1991). climate the average annual precipitation for the study area ranges between 800 and 1 200 mm per annum. daily mean maximum temperatures fluctuate between 22 and 24°c for the hottest month of january; during the coldest months (june and july), the daily mean temperature ranges from minus 2 to 0°c, respectively. the daily mean relative humidity for the most humid month of march varies between 68 and 72% and the daily minimum relative humidity for july and august between 32 and 38% (schulze, 1997; van zinderen bakker, 1973). snow is recorded to cover the top of platberg during some winters and can stay on the ground for days. water in southfacing gullies may be frozen for several weeks at a time (van zinderen bakker, 1973). possible heave soils are present on mtabazwe, the highest point on platberg (hilliard and burtt, 1987), and may be significant factors affecting vegetation and structure at high altitudes (hilliard and burtt, 1987). the climate of platberg and the drakensberg is part of a much longer, continental, north-south trending austro-afroalpine climate, with noticeably higher precipitation than the surrounding lowland. precipitation along the drakensberg escarpment is over 1 400 mm in some areas, with a gradual decrease both to the north and the south as well as westwards (560 mm) to the drier, sub-humid, continental-climate interior towards bloemfontein, some 400 km away (van zinderen bakker, 1973; scott, 1988). precipitation is orographic, not only from violent thunderstorms in summer but also from dense mists and fogs (smit et al., 1992) as well as snow. climatic heterogeneity associated with montane landscapes is, for cowling, holmes and rebelo (1992), a more important determinant of plant diversity than geological heterogeneity. inselbergs: their importance and uniqueness plant communities that occur on inselbergs form unique phytosociological associations (porembski and brown, 1995; parmentier, oumorou, porembski, lejoly & decocq, 2006; sarthou and villiers, 1998). however, inselberg plant associations also have numerous species that occur in the vegetation matrix surrounding the inselbergs (mutke, kier, braun, schultz and barhlott, 2001) and in which the inselberg flora is embedded (platberg and korannaberg fall within the grassland biome). the unique, high-altitude conditions found above 2 000 m lead to high levels of endemism in organisms, plants, animals and bryophytes (carbutt and edwards, 2006; haore and bredenkamp, 2001; hilliard and burtt, 1987; mucina and rutherford, 2006; smit et al., 1992; van wyk and smith, 2001). this is due to the compression of climatic life zones over a short distance that makes mountains hot spots for biological diversity (körner, 2003). mountains may be regarded as analogous with an archipelago of islands in an “ocean” of low-level vegetation types that act as an isolation factor (macarthur and wilson, 2001; taylor, 1996). this, in turn, precludes plant species with less mobile seed-dispersal mechanisms from propagating over wide ranges and allows for high levels of endemism to develop (hilliard and burtt, 1987; taylor, 1996). high levels of endemism mean that a large proportion of the available gene pool is unique to that site, and inselbergs and mountains therefore have an important role to play in the maintenance of genetic diversity (mucina and rutherford, 2006; taylor, 1996). methods platberg was chosen as the study site as it is the largest inselberg situated approximately 60 km north of the main massif of the maluti/drakensberg range (chief directorate: survey and mapping, 2000). it is also the highest peak in the area, at 2 394,5 m (survey beacon 44), as well as the largest inselberg capped by igneous rock (du toit, 1954; erickson, 1985; king, 1963; norman and whitfield, 1998). it forms a distinctive flattopped, l-shaped plateau, with clear-cut contact between the stormberg volcanics and the cave sandstone of the clarens formation, clearly visible from afar. the analysis of 38 relevés, which were extracted from a total of 393 relevés for the entire platberg, was used to constitute the fynbos community on platberg. the scope of the study was to sample above the 1 800 m contour in order to work within the limits set by killick (1978), who regarded the region in the drakensberg above 1 800 m as a distinct floristic region: the afroalpine region. the topography of the region is relatively flat and rises abruptly at the 1 900 m contour – this being the start of the footslopes of platberg – to the highest point of 2 394 m. plot sizes were fixed at 6 x 5 m to give a total surface area of 30 m² (du preez, 1991; malan, 1998; whittaker, 1980). in all the sample plots, each species was recorded, all the woody plants were counted and all the cover/abundance was calculated with the braun-blanquet cover-abundance scale (behr and bredenkamp, 1988; brown, marais, henzi and barrett, 2005; clever, brown and bredenkamp, 2005; mueller-dombois and ellenberg, 1974; westhoff and van der maarel, 1980; whittaker, 1980). ad hoc floristic collections were undertaken in 2004, with the main fieldwork starting in february/march 2005, and continued during the following summers in the growing season from october to december 2005 through to the end of march 2006. final follow-up fieldwork was done in january 2007. over 1 500 floristic collections were made and identified at either the geo potts herbarium (blfu) or the national herbarium (pre) in pretoria. germishuizen and meyer (2003) were used figure 2 landscape sketch showing the principal terrain units. (after land types survey staff 1991) 204 land type: ea 199 platberg, harrismith 2828. terrain type: b 3 terrain form sketch. terrain units: 1 plateau/scarp 2 scarp/cliff 3 midslope 4 footslope 5 valley bottom afromontane fynbos in platberg original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 202 213 to confirm the names of the taxa. the specimens are currently housed at the geo potts herbarium of the ufs. environmental data recorded included aspect, slope, exposure, the size of the rocks, latitude and longitude, altitude, locality, geology, the per cent of area covered by rock, topography, the degree of surface erosion, the degree of trampling, drainage, soil depth and notes on management and utilisation as well as total percentage canopy cover. numerous phytosociological studies (barrett, brown, barrett and henzi, 2006; bredenkamp and brown, 2003; du preez, 1991, 1992; eckhardt et al., 1995; eckhardt, van rooyen and bredenkamp, 1993; grobler, bredenkamp and brown, 2002; hoare and bredenkamp, 2001; malan, 1998; pond, beesley, brown and bezuidenhout, 2002; smit, bredenkamp and van rooyen, 1995) have shown that the braun-blanquet classification (kent and coker, 1992; whittaker, 1980) is the most efficient phytosociological method to use. habitat as well as floristic data were processed with turboveg (hennekens, 1996a). a first approximation was done with the twinspan (two-way indicator-species analysis) algorithm of hill (1979a). megatab (hennekens, 1996b) was used to generate a phytosociological table, which was refined by braun-blanquet procedures. a decorana (hill, 1979b) analysis was also applied to the floristic data to determine the relationship between the various vegetation units and the environmental gradients. erosion was estimated with the three-scale numerical system, where 1 = no erosion, 2 = moderate and 3 = high. slope was estimated in the following scale in degrees: 0 to 3° = flat, 3 to 8° = gradual, 8 to 16° = moderate, 16 to 26° = steep, 26 to 45° = very steep, and greater than 45°. soil depths were measured with a probe graded for 5 cm intervals to a maximum of 30 cm. hand-held garmin magellan 2x and etrex global positioningsystem instruments were used to give latitude, longitude, aspect and altitude for the sample plots. altitudes were also read from 1:50 000 topographical maps. results a total of 299 species (table 1) was recorded in 38 sample plots. the fynbos community has an average of 28.34 species per relevé, ranging from 14 to 54 species per sample plot. the fynbos is dominated by several low, sclerophyllous shrubs approximately 1 m high on the rocky outcrops, summit plateau, vertical cliffs, steep scree slopes and cave sandstone of the clarens formation. a mixture of low shrubs, grasses, forbs and succulents characterises the fynbos community for the study site. vegetation classification the analysis resulted in the identification of two communities, four sub-communities and seven variants. the phytosociological classification is presented in table 1 and is as follows: 1. tristachya leucothrix-–eragrostis curvula grassland community 1.1 eragrostis curvula-–themeda triandra grassland subcommunity 2.2 eragrostis curvula-–eragrostis racemosa grassland subcommunity 2.2.1 passerina montana variant 2.2.2 harpochloa falx variant 2. helichrysum dasycephalum-–digitaria monodactyla grassland community 1.1 bulbostylis humilis-–passerina montana subcommunity 1.1.1 watsonia lepida variant 1.1.2 cliffortia ramosissima variant 1.1.3 ischyrolepis schoenoides variant 1.1.4 eragrostis capensis variant 1.1.5 cliffortia nitidula variant 2.2 crassula sarcocaulis-–cliffortia nitidula subcommunity description of plant communities the study area is mostly open grassland, with scattered fynbos communities either on rocky outcrops, on the summit plateau area or concentrated on the steep slopes and cliffs of platberg on the sandstone of the clarens formation. little disturbance from grazing is evident, with no record of agricultural practice on the plateau of platberg. there are some clusters and scattered individuals of the declared alien invasive tree species pinus patula (species group e), having escaped from plantations established at lower altitudes and now being firmly ensconced on the footslopes and scattered on the summit plateau area. the sclerophyllous vegetation is characterised by passerina montana (species group m, table 1), cliffortia ramosissima (species group f, table 1) and cliffortia nitidula (species group p, table 1). common species throughout the study area include the forbs hebenstretia dura, watsonia lepida, commelina africana, oxalis obliquifolia, senecio glaberrimus, sutera floribunda and polygala amatymbica, the xerophytic fern cheilanthes quadripinnata, the perennial tufted grasses eragrostis curvula and melinis nerviglumis, the two sedges carex zuluensis and ficinia stolonifera, the succulent crassula lanceolata subsp. transvaalensis and the geophyte merwilla plumbea (species group p, table 1). the geophyte agapanthus campanulatus and the climber cyphia elata (species group o, table 1) occur in all the communities. 1. tristachya leucothrix–eragrostis curvula grassland community the tristachya leucothrix–eragrostis curvula grassland is located on flat to moderately steep rocky terrain on the ridge lines and steep slopes. rock cover varies between 10 and 40%, with sandy soil between 40 and 250 mm deep. on the steep slopes, high erosion is present, with no erosion occurring on the flat areas. this community is characterised by the single-bunch grass tristachya leucothrix (species group a, table 1). vegetation is dominated by the perennial grass eragrostis curvula (species group p), the low fynbos shrub passerina montana (species group m) and the dwarf shrub selago galpinii (species group f). other species of importance include the perennial grass tristachya leucothrix (species group a), the restio ischyrolepis schoenoides (species group g), eragrostis curvula (species group p), the forb watsonia lepida and the sedge ficinia stolonifera (species group p, table 1). the total species count is 338, which gives an average of 30.7 species per 30 m². there are two sub-communities and two variants: 1.1 eragrostis curvula-–themeda triandra grassland subcommunity the eragrostis curvula–themeda triandra sub-community is located on the northern and north-western slopes, with gradients varying between 8 and 26°, in full sun and in soil depths varying between 40 and 60 mm. the vegetation occurs on the contact between the cave sandstone of the clarens formation and the igneous cap at approximately 2 200 m. surface erosion is high, with rock cover estimated at 15%. the area is overgrazed, with concomitant high levels of erosion due to impact by cattle. species belonging to species group b, table 1, are diagnostic for this community and include the geophyte chlorophytum cooperi, the forbs cyanotis speciosa and acalypha punctata and the perennial grass themeda triandra. 205 original research brand, du preez & brown koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 202 213 number of species per relevé 2 2 2 2 1 2 3 4 2 4 4 3 3 2 2 5 3 2 2 2 1 3 3 2 2 2 4 2 2 3 2 3 1 2 1 3 1 2 5 7 2 1 6 8 1 1 9 5 0 3 8 3 8 4 6 5 3 6 7 1 2 0 0 2 6 9 7 2 2 6 7 6 3 5 4 7 community 1 2 sub-community 1.1 1.2 2.1 2.2 variant 1.2.1 1.2.2 2.1.1 2.1.2 2.1.3 2.1.4 2.1.5 species group a tristachya leucothrix + . . . b . . a b . . + . . . . . . . . . . . . . . . . . . . + . . . species group b themeda triandra b . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . cyanotis speciosa . + + + . r . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . + . chlorophytum cooperi . + . . r . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . acalypha punctata + . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . species group c eragrostis racemosa . . . . . a a b . . . . . . . . . . b . . . . . . . . . . . . . . . heteropogon contortus . . . . . b b a b . . . . . . . + + . . . . . . . . . . . . . + . species group d harpochloa falx . . . . . + . b b + + . . . . . . . . . . . + . . . . . . . . . . + . . helichrysum nudifolium . + . . . . . + . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . wahlenbergia depressa . . . . . . . . . r r . . . . . . . . . . . . . . . . . . . . . . . . . . . senecio paucicalyculatus . . . . . . . . r r . . . . . . . . . . . . . . . . . . . . . . . . . . . . species group e aristida monticola . . . . . . b . . . a . . b . . . . . . . . . . . . . . . . + . a . b cymbopogon dieterlenii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinus pinaster . . . b . . . r . . . . . . . + . . . . . . . . . . . . . . . . . . . . . eragrostis chloromelas . . . . . . . . . . . + . . + . . . . . . . . . . . . . . . . . . . . . eucomis bicolor . . . . + . . . . . . . r . . . r . . . . . . . . . . . . . . + . . . . . . rhus dentata . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . clutia affinis + . . . . . . . . + . . . + . . . . . . . . . . . . . . . . . . . . . . . . rhus discolor a . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . species group f selago galpinii + + + . + + + + + + . + + + . . . . . . . . . . . . . + . . . . cliffortia ramosissima . . . . . b . . . . . . . b a . . . b . . a . . . . . . . . . . . senecio subrubriflorus . . . . . . . . . + . . + . . . . + . . . . . . . . . . . . . . . . . . . species group g ischyrolepis schoenoides . . . . . a . . . . . . . . b b b . . . . . . . . . . . . . . crassula peploides . . + . . . . . + . . . + . . . . . + . + r + r . . . . . + . . . . . . . . macowania glandulosa . . . . . b . . . . . . . . . . . . . . . . + + . . + . . . . . . . . . . . aster perfoliatus . + . . . . . + + . . . . . . . + . . r . . . . . . . . . . . . . . . . . . helichrysum sutherlandii . + . . . . . . . . . . . . . + . . + . . . . r . . . . . . . . . . . . . . species group h digitaria monodactyla + . . . . . b . . . . a a . b + b . a . b . a . . . . . helichrysum dasycephalum . + . . . . + . . . . + + . + . . + . + b a . . . . + + b . . . . cyperus schlechteri . . . . . . . . . . . + . + + + . . . . . . . . + . + . . . . + . . crassula dependens . . . . . . . + . . . . . . + + + r + . . . + + . . . . . + . . b . . . . . schizocarphus nervosa . . . . . . . . . . . . . . . + . + + . . + . . + + . . + + . + . + + . + ruschia putterillii . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . psammotropha mucronata . . . . . . . . . + . . . . + . . + r . + + + . . . . . r + r . . . + + . + helichrysum ecklonis . . . . . . . . . . . . . . . . . . . r . + + + . . . . . . . + . . . + . + stoebe vulgaris . . . . . . . . . . . . . + . . . . b . . . . . . . . . . . . + . . . . dimorphotheca jucunda . . . . . . . . . . . + + . . . . . + r . . . . . . . . . . . . + . . . . muraltia saxicola . + . . . . . . . . . . + . . . . + . . . . . . . . . . . . . . . . . . . lotononis lotononoides . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . species group i bulbostylis humilis . . . . . . . . . + . + + . + + + + + . . . + + a + r . + . . . . + . . . crassula lanceolata . . . . . . . r . r + . . . . . . . r . . . r . . r . . . + r + . . . . . . senecio othonniflorus . . . . . . . . . . + . + . . . . . . . . r + . . . . . . + . . . + . . . . cycnium racemosum . . . . . . . . . . . . . + . . . . . . . . + . . . + . + . . . . . . . . . sebaea natalensis . . . . . . . . . . . . + . + r . . . . . + . . . . . . . . . . . . . + . . table 1 fynbos communities of platberg, eastern free state plant species with a presence of 4 or less have been omitted. 2 3 2 1 1 3 3 2 2 3 2 2 3 1 2 3 2 2 2 2 2 3 2 2 1 3 6 2 5 6 3 5 6 8 3 4 4 6 8 2 7 9 2 7 7 3 3 7 9 9 2 2 5 5 7 7 2 1 5 3 8 7 3 8 4 9 0 1 0 2 6 5 2 5 7 7 9 9 9 1 8 7 6 3 1 6 4 3 6 3 2 5 3 2 7 9 5 0 7 9 9 0 database no. 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 4 2 2 2 2 5 4 3 3 5 5 4 4 5 4 4 5 3 4 5 2 2 4 4 4 4 4 5 4 2 4 2 3 5 2 2 2 2 6 3 5 6 3 5 7 8 3 4 5 7 9 3 8 9 3 8 8 3 3 8 9 9 3 2 5 5 7 7 3 2 5 3 9 8 4 8 8 3 4 5 4 6 0 9 6 9 1 1 3 3 3 5 2 1 0 7 5 0 8 7 0 7 6 9 7 6 1 3 9 4 1 3 3 4 206 afromontane fynbos in platberg original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 202 213 number of species per relevé 2 2 2 2 1 2 3 4 2 4 4 3 3 2 2 5 3 2 2 2 1 3 3 2 2 2 4 2 2 3 2 3 1 2 1 3 1 2 5 7 2 1 6 8 1 1 9 5 0 3 8 3 8 4 6 5 3 6 7 1 2 0 0 2 6 9 7 2 2 6 7 6 3 5 4 7 community 1 2 sub-community 1.1 1.2 2.1 2.2 variant 1.2.1 1.2.2 2.1.1 2.1.2 2.1.3 2.1.4 2.1.5 cotula socialis . . . . . . . . . . . . + . . + . . . . . . . . . . r . . . . . . . . . . . panicum eckloniiwahlenbergia cuspidata .. . . . . . . . . + . + . . b . . . + . . . . . . . . . . . . . . . . species group j crassula biplanata x ericoides . . . . . . . . . . . r + . + + + . . . . . . . . . . . . . . . . . . . . . hesperantha baurii . . . . . r . . . . . + + . r + r . . . . . r + . . . . . . . . . + . . . . merxmuellera drakensbergensis . . . . . . . . . . . . . . b . + . . . . . . . . . . . . . . . . . . . . aeollanthus buchnerianus . . . . . . . . . . . + . . . + . . r . . . . . . . . . . . . . . . . . . . helichrysum album . . . . . . . . . . . . + . + . . . . . . . . . . . . . . . . . . . . . . . species group k eragrostis capensis + . . . . . . . . . . . . . . . . . . . . . . . b + . . . . . . . . . . . hypoxis costata . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . ledebouria cooperi . . . . . . . . . . . . . . . . . . . . . . + . + + r . . . . . . . + + . . oxalis depressa . . . . . . . . . . . r + . . . . . . . . . . . + + + . . . + . . . . . . . anthospermum herbaceum + r . . . r . . . . + . + . . . . . . . . . . . r r r . . . . . . . . + . . felicia filifolia . . . . . . . . . . . . . . . . . . . . . . a . . . . a . . . . . . . pennisetum sphacelatum . . . . . . . + . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . hermannia gerrardii . . . . . . . . . . . . . . . . . . . . . . . . r + . . . . . . . . . . . . nemesia caerulea . . . + . . . . . . r . . . . . . . . . . . . . b + . . . . r . . . . . . . species group l crassula depressa . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . b . . . . . . . cyperus rupestris . . . . . . . . . . . . . . . . . . . . . . . . . . . b . . . . . . . eucomis montana . . . . . . . . . . . . . . . + . . . . . . . . . . . . . + + . . . . r . . polygala virgata . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . r . . . . . . species group m passerina montana a + + + + + b + b . b b b + b a b + b + . . . . . . . delosperma sutherlandii . + . . . r r . . r . . . . . . . r r r + . . r . . . . . r . . . . + . . . helictotrichon longifolium . . . . . a . . b . . . . . . . . . . . . . . . + . b . . . . . . . . . senecio inornatus . . . + . . . . . . . . . . . + + . . . . . . . . . . . + . . . . . . rhus pyroides + . + . . . . . . . . . . . . . . . . . . . . . . b . . . . . . . . . . . euclea coriacea . . + . . . . . . + . . . . . . . . . . . . . . . . . + . . . . . . . . . . chrysocoma ciliata + . . . . . . . . . . . . . . . . . . + . . . . . . b . . . . . . . . . . species group n crassula sarcocaulis . . . . . . . . . . . . . . . . . . . . . . . . . + b . . . . . + . + + rhodohypoxis baurii . . . . . . + . . . . . + . . . . . . . . . . . . . . . . + . . . . + r . + drimia sphaerocephala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . + . . + albuca humilis . . . . . . . . . r r . . . . . . . . . . . . . . . . . . + . . + . . . . + helichrysum chionosphaerum . . . . . . . . + . . . . . . . . . . . . + . . . . . . . . . . . . + + . . senecio inaequidens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . + species group o senecio rhomboideus . . . . . . r + . r . r r . . + + . . + + . r + . . + + + . r . + + r . + agapanthus campanulatus . . . . . . . . r . + . . . + + + + . . . + . . . . + + . + . + . + cyphia elata r . . . . . r . . . . + + . . + + . . . . + + . . . + . + + . . . r + + . + eriospermum ornithogalum . . . . . . . r . + . . + . + . + r + r . + + . . . . . + + . r . + . . . + gazania krebsiana . . . . . + . . . + . . . . + . . r . . . + + . . . . . . . . . . . + + . . moraea brevistyla . . . . . + r . r r . . . . . . . . r . . . + . r . r . r . . . + . . . . + species group p cliffortia nitidula . . + . b . . . . . . b . a b . . . . . b . . . . . . + a watsonia lepida . . . b . b b . + + + . . . . b . . . . . . . . . + . . + hebenstretia dura . . . + . . . . . . + + . . + + . + . . . . . + . . . . + + . . . . + . + cheilanthes quadripinnata . . + . . . + + + + + + . + + + + + + . . . . . + + . . . + . + . + commelina africana + . . . . . + + + + + + . + . + + . . + . . + . + + + . . + . + + . . . . oxalis obliquifolia r . . . . . r + + . r r r . . r + + + . . + r . . . . + + + . . . + + + . + carex zuluensis . . . + . . . b + . . . + . + . . + . . . + . + + . b . . . + . . . . eragrostis curvula . . . . a b . . . . . . . . . . . + + a . . . . . . . . . cyperus semitrifidus . . + . . . . . a . . . . . . . + + . . + . + . . . . . . . . . . merwilla plumbea . . . + . . + r + . . . . . . . . . . . . . . . a + . . . . . + . . + . crassula lanceolata sp. transvaalensis . + . . . . r r . . . r . + . r + . . . + . . . . . . + . . . . + . + . + . senecio glaberrimus . . . . + . . + + . . . + . . . . . . + . . . . . . . . . . . + . . . + . . ficinia gracilis . . . . + . . b . . . . + . . . . . . + . b . . . . + . . . + . . melinis nerviglumis + . . . . . . . + b . . . b + . . . . . . . . . + . . . . . . . . ficinia stolonifera . . . . . . b b . . . . . . . . . . . . . . . . . . . . . sutera floribunda . . . . . . r + . . . . r . . . . . . . . . . . . . . + . . . . . . . . + . polygala amatymbica . r . . + . . + . . . . . . . . . . . . . + . . . . . . . . . . . . . + . . table 1 (cont...) plant species with a presence of 4 or less have been omitted. 207 original research brand, du preez & brown koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 202 213 the vegetation has no dominant species but it does have significant cover abundance by the low shrubs passerina montana (species group m) and selago galpinii (species group f), the perennial grasses themeda triandra (species group b) and eragrostis curvula (species group p) and the forb watsonia lepida (species group p), with a dense stand of the invader exotic tree pinus patula (species group e, table 1). the total species count is 111, which gives an average of 22.2 species per 30 m². 1.2 eragrostis curvula-–eragrostis racemosa grassland subcommunity the eragrostis curvula–eragrostis racemosa sub-community is located on rocky terrain, on steep to very steep slopes (16 to 45°), in the sandstone of the clarens formation, and on the cool, southern or south-western slopes. the community is in full sun on rocky outcrops of between 10 and 40%. soil depths are shallow (100 mm) or > 300 mm deep, a mix of sandstone and basalt, and well drained. the short tufted grasses eragrostis racemosa and heteropogon contortus (species group c, table 1) are diagnostic and also the dominant species for this sub-community. the sclerophyllous shrub passerina montana (species group m, table 1) is also codominant. other species of importance include the grass tristachya leucothrix (species group a), aristida monticola (species group e), eragrostis curvula, (species group p), the restio ischyrolepis schoenoides (species group g), watsonia lepida, the sedge ficinia stolonifera (species group p) and the dwarf shrub selago galpinii (species group f). the total species count is 222, which gives an average of 37 species per 30 m². 1.2.1 passerina montana variant this variant occurs on the southern aspects, on the steep to very steep (16 to 45°) mid-slopes, on a series of rocky ledges and semi-vertical cliffs (with 10 to 25% exposure), and in the cave sandstones of the clarens formation. soils are shallow (40 to 100 mm) and well drained, and show no trampling or erosion. the vegetation is dominated by the sclerophyllous shrub passerina montana (species group m), with the restio ischyrolepis schoenoides (species group g) and the perennial grass eragrostis racemosa (species group c) being co-dominant. other species with significant cover abundance include the perennial grasses tristachya leucothrix (species group a), heteropogon contortus (species group c) and helictotrichon longifolium (species group m), and the fynbos shrubs cliffortia ramosissima (species group f), macowania glandulosa (species group g) and cliffortia nitidula (species group p). the community is depaupered in geophytes, herbs and forbs, with an average species per 30 m² of 28.1. 1.2.2 harpochloa falx variant this variant is located on the mid-slope at 2 100 m on moderate to steep (28 to 45°) slopes. rockiness is between 10 and 50%, with dry, shallow soils 40 to 120 mm deep. no trampling or erosion was seen, despite the steep slope. the grass harpochloa falx and the forbs helichrysum nudifolium, wahlenbergia depressa and senecio paucicalyculatus (species group d) are diagnostic for this variant. the vegetation is dominated by the grasses eragrostis racemosa and heteropogon contortus (species group c). important species include the grasses tristachya leucothrix (species group a), harpochloa falx (species group d) and aristida monticola (species group e), eragrostis curvula, melinus nerviglumis, the sedges carex zuluensis, cyperus semitrifidus and ficinia stolonifera (species group p), the restio ischyrolepis schoenoides (species group g), the shrubs passerina montana and selago galpinii (species group f) and the fern chelianthes quadripinnata (species group p). the total species count is 165, which gives an average of 41.2 species per 30 m². 2. helichrysum dasycephalum–digitaria monodactyla grassland community the helichrysum dasycephalum–digitaria monodactyla community is the largest fynbos community on platberg. it is located on the southern aspects, on the flat, rocky ridge line on the summit plateau and on the steep, exposed mid-slope along the cave sandstone. soils are well drained and shallow (40 to 180 mm) or > 300 mm deep. rock cover varies between 10 and 50%, with little or no erosion or trampling visible. species belonging to group h are diagnostic for this community and include the grass digitaria monodactyla, the forbs helichrysum dasycephalum, psammotropha mucronata, helichrysum eckloni and dimorphotheca jucunda, the sedge cyperus schlecteri, the succulents crassula dependens and ruschia putterllii, the geophyte schisocarpus nervosa, the fynbos dwarf shrub muraltia saxicola and the shrub lotononis lotononoides. the community is dominated by the grass digitaria monodactyla (species group h) and the sclerophyllous shrub passerina montana (species group m). important species include the forb helichrysum dasycephalum, the sedge cyperus schlecteri (species group h), the shrub cliffortia nitidula, the bulb watsonia lepida, the sedges and grasses ficinia gracilis and f. stolonifera, eragrostis curvula and melinis nerviglumis (species group p), respectively. the total species count is 752, which gives an average of 27.9 species per 30 m². 2.1 bulbostylis humilis–passerina montana sub-community the bulbostylis humilis-–passerina montana sub-community occurs in full sun on the gently sloping ridge line (8°) on the summit basalt and on the steep slopes (26 to 45°) of the cave sandstone. rock exposure varies between 15 and 30%. little or no erosion, with moderate to no trampling, was noted. soils are either shallow (40 to 160 mm) or > 300 mm deep, well drained and a mix of weathered basalt, sandstone and dolerite. canopy cover for the shrubs is < 2 m high and is 5 to 15%, with the grasses, forbs and herbs forming a low sub-layer. the species diagnostic for this community include the small sedge bulbostylis humilis, the small erect succulent crassula lanceolata, the forb senecio othonniflorus, the root parasite cycnium racemosa, sebaea natalensis, cotula socialis, wahlenbergia cuspidata and the grass panicum ecklonii (species group i). the vegetation is dominated by the grass digitaria monodactyla and the forb helichrysum dasycephalum, with co-dominance of the fynbos shrubs passerina montana (species group m). other significant species include cliffortia nitidula, the bulb watsonia lepida, the grasses eragrostis curvula and melinis nerviglumis, the sedges ficinia gracilis and f. stolonifera and the geophyte merwilla plumbea (species group p). the total species count is 620, which gives an average of 29.5 species per 30 m². 1.1.1 watsonia lepida variant this variant is located in full sun, on the hot western and northern aspects, on the moderate to very steep slopes (16 to 45°) and on the mid-slope and plateau area. altitudes vary from 2 100 to 2 200 m 208 afromontane fynbos in platberg original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 202 213 the low shrub cliffortia nitidula, the forb hebenstretia dura and the sedge ficinia gracillis (species group p). the total species count is 100, which gives an average of 25 species for 30 m². 1.1.4 eragrostis capensis variant this variant is located on the dry northern and north-western aspects on gentle slopes (3 to 8°) between 2 250 and 2 320 m on the basalt summit in full sun. rock exposure is less than 5%. soils are well drained and depths vary from medium (100 to 110 mm) to (> 300 mm) deep. no trampling or surface erosion was seen. the variant is low, flat grassland with limited shrub presence. the variant is defined by species from species group k, which includes the perennial grasses eragrostis capensis and pennisetum sphacelatum, plus the forbs hypoxis costata, oxalis depressa, anthospermum herbaceum, felicia filifolia, hermannia gerrardii and nemesia caerulea and the geophyte ledebouria cooperi. the vegetation is further dominated by the grasses digitaria monodactyla (species group h) and eragrostis capensis, the forb felicia filiformis (species group k) and the shrub passerina montana (species group m). species with significant cover abundance include the small sedge bulbostylis humilis (species group i) and the forbs hypoxis costat and nemesia caerulea (species group k), along with the shrubs cliffortia ramosissima (species group f) and rhus pyroides (species group m) and the geophyte merwilla plumbea (species group p). the total species count is 88, which gives an average of 29.3 species per 30 m². 1.1.5 cliffortia nitidula variant this cliffortia nitidula variant occurs in full sun and is scattered over all aspects on moderate (0 to 8°) slopes at 2 291 to 2 310 m on the summit basalt or at 2 151 to 2 176 m on very steep > 45° slopes on the scree slopes below the cliffs. rock cover varies between 15 and 25%, with very shallow, well-drained soils between 10 and 50 mm deep. no trampling was seen and zero erosion was evident, despite the boulder-scree terrain. canopy cover for shrubs < 2 m tall is between 5 and 20%. the community forms the distinct horizontal vegetation line growing halfway up the cliffs. geologically, it is a dolerite sill that forms a narrow (5 to 10 m) ledge, visible as a bench on the south side of platberg. this variant is defined by the succulent crassula depressa, the sedge cyperus rupestris, the geophyte eucomis montana and the slender shrub polygala virgata (species group l). no single species dominates this community. there is significant presence of the shrubs cliffortia nitidula (species group p) and passerina montana (species group m). the graminoid/sedge layer includes the perennial grasses themeda triandra (species group b), digitaria monodactyla, helictotrichon longifolium (species group m), eragrostis curvula and melinis nerviglumis (species group p) and the sedges cyperus rupestris (species group l), carex zuluensis and ficinia gracilis (species group p). other species that are locally prominent are the forbs helichrysum dasycephalum (species group h) and felicia filiformis (species group k), the succulent crassula depressa (species group l), the low karroid shrub chrysocoma ciliata (species group m), the succulent shrub crassula sarcocaulis (species group n) and the geophyte agapanthus campanulatus (species group o). the total species count is 146, which gives an average of 29.2 species per 30 m². 2.2 crassula sarcocaulis–cliffortia nitidula sub-community this sub-community covers basalt sheets and boulders on moderate slopes of between 8 and 26° in full sun on the plateau for the slopes and 2 300 m for the plateau. rock exposure varies between 10 and 25%. no trampling or erosion was noted. soil depths vary between 20 and 200 mm. the variant was located either on basalt of the drakensberg volcanics or cave sandstone of the clarens formation. the diagnostic species for this variant are the geophyte hesperantha baurii and the succulent crassula biplanata x ericoides, with the tall mountain grass merxmuellera drakensbergensis and the forbs aeolanthus buchnerianus and helichrysum album (species group j). the grass digitaria monodactyla (species group h), together with the fynbos shrub passerina montana (species group m), dominates this vegetation. species with significant cover include the shrubs cliffortia ramosissima (species group f) and cliffortia nitidula (species group p), the perennial grasses cymbopogon dieterlenii and aristida monticola (species group e), the restio ischyrolepis schoenoides (species group g), the sedge cyperus schlecteri (species group h), eragrostis curvula, melinis nerviglumis, the geophyte schisocarpus nervosa (species group p) and the forb senecio inornatus (species group m). the total species count is 212, which gives an average of 35.3 species per 30 m². 1.1.2 cliffortia ramosissima variant this variant is located on the hot and dry western and northwestern aspects on moderate (8°) to steep (> 45°) inclines on the summit plateau. rock exposure is 15 to 30%. soils are shallow, from 40 to 100 mm deep, and well drained. no trampling was noted and there was moderate to zero soil erosion. the low sclerophylous shrub passerina montana (species group m) covers 20 to 50% of the variant and grows at altitudes of between 2 224 and 2 367 m. the dwarf shrubs selago galpinii and cliffortia ramosissima, along with the forb senecio subrubrifloris (species group f), are diagnostic for this variant. the vegetation is dominated by the sclerophyllous shrubs cliffortia ramosissima (species group f) and passerina montana (species group m) and the short bunch grass digitaria monodactyla (species group h). only two species have any significance: the low shrub selago galpinii and the forb senecio subrubriflorus (species group f). the total species count is 74, which gives an average of 25 species per 30 m². 1.1.3 ischyrolepis schoenoides variant this variant is located in full sun on the northern and eastern aspects with flat or moderate (8 to 16°) slopes on the rocky summit ridge line at altitudes of 2 100 to 2 279 m. rock exposure varies between 15 and 25% and is either basalt or cave sandstone. no trampling or surface erosion was seen. soil depths are shallow, between 40 and 180 mm, well drained and of mixed ib land type. total canopy cover is 10 and 40% for the low shrubs cliffortia ramosissima (species group f) and passerina montana (species group m) and less than 2 m high. this variant has no defining species group, although it is defined by the high cover abundance of the restio ischyrolepis schoenoides (species group g). the vegetation is dominated by the short, leafless, grass-like restio ischyrolepis schoenoides (species group g) and the low shrub passerina montana (species group m). other species of importance include the grasses digitaria monodactyla (species group h), the forb helichrysum dasycephalum (species group h), 209 original research brand, du preez & brown koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 202 213 ridge line. it is predominantly located on the north-eastern aspects at 2 290 to 2 315 m. soils are shallow (40 to 120 mm), they are seasonally damp to waterlogged, the sand and gravel < 10 mm fine, and the soils exhibit no trampling or erosion. rock exposure is 10 to 20%. species belonging to species group n are diagnostic for this subcommunity and include the succulent shrub crassula sarcocaulis, the geophytes rhodohypoxis baurii, drimia sphaerocephala and albuca humilis and the forbs helichrysum chionosphaerum and senecio inaequidens. the sub-community has no dominant species but the significant presence of the fynbos shrub cliffortia nitidula (species group p) and the perennial grasses aristida monticola (species group e), digitaria monodactyla (species group h) and panicum ecklonii (species group i) is notable. other important species include the caespitose forb helichrysum dasycephalum, the sedge cyperus schlecteri and the decumbent succulent crassula depressa (species group h), the forbs senecio rhomboideus and agapanthus campanulatus (species group o), the fern cheilanthus quadripinnata and the sedges cyperus rupestrus, ficinia gracilis and f. stolonifera (species group h). this sub-community has a high proportion of geophytes. the total species count is 132, which gives an average of 22 species per 30 m². ordination and environmental gradients the distribution of fynbos plant communities for platberg along the first and third axes of the decorana (hill, 1979b) ordination diagram is shown in fig. 3. soil moisture decreases from left to right, with soil depth reflecting this pattern along axis 1 (eigen value = 0.779). the decrease in soil depth may reflect an increase in rockiness and shallow soil. the presence of pine and wattle trees on the extreme left (relevé 37265) of the ordination diagram and of well-developed grassland (sub-community 1.1, themeda triandra) on the far right may indicate an environmental gradient of disturbance (invasion by alien species), decreasing along axis 1 from left to right. fire is an important factor in fynbos distribution and this may be reflected by irregular occurrence on the left and more regular on the right, as seen in sub-communities 1.1, via restio, to 2.2 and passerina montana. the environmental gradient along axis 1 may be related to exposure and frost effects and increases from left to right. rockiness and exposure also increase as the altitude increases from left to right. this may be indicated by the physiognomy of the habitat trend on this axis. the environmental gradient along axis 3 (eigen value = 0.498) may be a combination of decreasing soil depth from the bottom to the top of axis 3 as well as of soil texture and composition, with shallow, course-grained, leached quartzitic soils at the top of the ordination diagram and with deeper, higher-organic content, finer soils at the bottom of the axis. the vegetation gradient for axis 3 from the community circled as passerina montana through to 2.1, with the inclusion of variant 2.1.4, can be explained by the general habitat gradient. an examination of the ordination diagram for axes 1 and 2 (eigen value = 0.580) as well as for axes 2 and 3 does not clearly show related community clusters (table 1) along clearly discernible environmental gradients. it may be that a diagonal environmental gradient exists along axes 1 and 3, from bottom left to top right, which better explains the combination of the aspect habitat and environmental gradient of physiognomy, of fire, of soil depth and composition, of exposure and frost, of soil moisture and of rockiness. discussion prior to this survey, no phytosociological classification had been completed for platberg. the description of the fynbos is part of a larger study on the entire vegetation of platberg and has been identified as one of four distinct communities occurring on this inselberg (the others are wetlands, a woody/shrubland community and grassland). figure 3 decorana ordination of the fynbos communities of platberg 210 afromontane fynbos in platberg original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 202 213 the sclerophyllous vegetation of the afromontane fynbos communities of the drakensberg region is dominated by the low shrubs cliffortia nitidula and passerina montana, with a less dominant but significant cover abundance of selago galpinii and cliffortia ramosissima (du preez, 1992). the geophyte watsonia lepida, the fern cheilanthus quadripinnata, the grasses eragrostis curvula and melinis nerviglumis and the two sedges carex zuluensis and ficinia stolonifera are common throughout. of note is that no protea shrubs occur on platberg, although some protea species have been recorded in the vicinity (the golden gate national park, the qwa qwa national park, the southern drakensberg and kwazulu-natal as well as the northern drakensberg). the study resulted in the identification of 13 different fynbos vegetation units, which can be grouped into two communities, four sub-communities and seven variants. at least 16 fynbos genera occur on platberg, with 22 endemic or near-endemic species belonging to the drakensberg alpine centre (dac) and with 22 exotic species (alien invasives). fynbos on platberg shows a species richness of 28.34 per 30 m² (with a total 43.67 per km² for the whole of platberg), which is comparable with the cape and which is moderately higher than other floristic regions in south africa (pond et al., 2002). affinities with platberg at plant-community level, the fynbos element on platberg is embedded within the broader grassland biome. numerous phytosociological and floristic studies on the grassland biome elucidate the strong floristic links that exist within several of the putative grassland bioregions (mucina and rutherford, 2006) and with the dac (bredenkamp and brown, 2003; grobler, bredenkamp and brown, 2005). the floristic links with the fynbos on platberg and the cape floral kingdom are shown by the following genera, which are considered typical of the cape centres: cliffortia, erica, metalasia, muraltia, passerina, pentaschistis, restio, schoenoxiphium and watsonia (bester, 1998; cowling et al., 1992; du preez, 1991; haaksma and linder, 2002; pond et al., 2002). all these genera, except for the restio, are present on platberg (table 2) and form main communities or sub-communities (table 1). the family restionaceace is so dominant in the cape floristic region that it is of great importance (haaksma & linder, 2002). it is considered to be one of the diagnostic families used as a distinguishing part of fynbos. the restionaceae genus ischyrolepis schoenoides is found on platberg as well as other inselbergs in the free state (du preez, 1991, 1992; du preez and bredenkamp, 1991; malan, 1998) and the main drakensberg (carbutt and edwards, 2004; hilliard and burtt, 1987; körner, 2003; pooley, 2003). it can be regarded as a replacement genus for the genus restio, as its growth form is similar and phylogenetically these two genera are closely related (haaksma & linder, 2002). taxonomically, it was formally named restio schoenoides to indicate its links with the genus restio (germishuizen and meyer, 2003; haaksma and linder, 2002). two fynbos species on platberg, eriocephalus and euryops, as well as the two woody/shrub species maytenus and diospyros are found in association with renosterveld (pond et al., 2002) in the western cape. passerina, erica and gnidia are also associated with fynbos (goldblatt and manning, 2000; pond et al., 2002). proteaceae and the common genus protea are a major component of cape flora (cowling et al., 1992; goldblatt and manning, 2000; van wyk and smith, 2001). of note is that no protea shrubs grow on platberg. they have, however, been recorded in the proximity at the golden gate national park (roberts, 1969) and 60 km to the south in qwa qwa (moffett, daemane, pitso, lentsoane, and taoana, 2001), in kwazulu-natal, (robbeson, 1998) and to the northern drakensberg (matthews, 1991) and southern drakensberg (bester, 1998; hill, 1996; smit et al., 1992). additionally, floristic collections of protea caffra, p. subvestita and p. roupelliae have been made 20 to 30 km to the south of platberg (in 2003, 2004 and 2005, currently housed at the geo potts herbarium of the ufs). no evidence, such as excessive fire, overgrazing or the cutting of proteas, can be found to explain this anomaly on platberg. affinities with the dac the fynbos genera given above also occur throughout the drakensberg (hill, 1996; hilliard and burtt, 1987; killick, 1963, 1978; van wyk and smith, 2001), with carbutt and edwards (2004; 2006) providing the most current and complete species list and floral analysis illustrating these phytogeographic links. the fynbos species found on platberg is, according to acocks (1988), “different in origin and nature from the tropical vegetation” and is a vegetation type associated with poor, white, sandy soils. for platberg, it would seem that the response to these poor soils and this habitat is the existence of the fynbos vegetation type. it is also a response to climate change over an extended period, leaving relict populations in mountainous areas (acocks, 1988; hilliard and burtt, 1987; white, 1983). fire, grazing and conservation studies by everard (1986), seabloom and richards (2003) and heini, silva and tacheba. (2004) show that fire is a key factor influencing vegetation composition and structure, and fynbos propagation and control, with these having been extensively documented and researched (acocks, 1988; cowling et al., 1992; mucina and rutherford, 2006). the ordination for fynbos on platberg (fig. 3) highlights these environmental influences on the composition and structure of the sclerophyllous vegetation type. grazers and herbivores are important factors controlling the spread of woody vegetation (mucina and rutherford, 2006; seabloom and richards, 2003). they show that the dispersal and foraging behaviour of herbivores has the potential to structure plant communities over wide spatial and temporal ranges. on platberg, fynbos exists on the steep sides of the clarens formation, on rocky outcrops and on the exposed rim above the cliffs, were its location greatly reduces herbivory and trampling. more importantly, the effects of cold temperatures below freezing and high altitudes over 2 200 m (cowling, 1992; mucina and rutherford, 2006; van wyk and smith, 2001) would suggest these as the main mechanism responsible for maintaining the fynbos community as well as preventing the invasion of grassland by fynbos species. by fynbos. conservation the fynbos community on platberg has moderate species richness and low alpha diversity (table 1). however, it is embedded within the grassland biome, which has high species richness (the total floral count for platberg is 687) (mucina and rutherford, 2006), although this is rapidly being fragmented, with as much as 89% transformed and between 0 and 12.53% conserved (low and rebelo, 1996). as a biome, it is furthermore under enormous pressure for the little remaining untransformed grassland to be developed (bester, 1998; low and rebelo, 1996; malan, 1998; mucina and rutherford, 2006; perkins, bredenkamp & granger, 1999). platberg has high terrain heterogeneity, with steep contours, deeply incised gullies and numerous slope orientations. according to cowling et al. (1992), low and rebelo (1996), mucina and rutherford (2006), perkins et al., (1999) and van wyk and smith (2001), this provides more habitat for species and is more important than other factors in determining high biodiversity. in addition, platberg is the single largest, bestpreserved, high-altitude grassland in the free state (mucina and rutherford, 2006). 211 original research brand, du preez & brown koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 202 213 other, immediate threats are from the invasion of pinus patula on the footslopes and plateau area of platberg to the south and east. this poses a considerable conservation problem. pinus patula is in the process of rapidly out-competing local indigenous vegetation, particularly on the cool southern slope and in the stream flowing down the gully on the south side of platberg. for conservation, the threat to genetic diversity (anderson, 2001) as well as global warming are key considerations. these highlight the importance of mountains and associated inselbergs and stress the great need for and urgency of their protection. additionally, high-altitude vegetation, specifically alpine flora as defined by white (1983) and körner (2003), is the only biogeographic unit on land with a global distribution that accounts for nearly 3% of land surface and includes about 10 000 species or about 4% of higher-order plants (körner, 2003). the threat posed by global warming to the world’s alpine regions, including those in africa, was predicted by peters (1992), who indicated that as little as a 3ºc increase in temperature for 100 years would be equivalent to a 500 m upward shift in altitudinal zones. for the afromontane and afroalpine biota, this would cause a significant reduction in the distribution of plants and change their structure and composition, forcing some taxa to higher altitudes (taylor, 1996). in some cases, such as that of platberg, a significant reduction in taxa could also occur where the altitude is under 2 500 m. conclusions fynbos is defined as having proteaceae, ericaceae and restionaceae (goldblatt & manning, 2000). no proteaceae grow on platberg, although they have been recorded in the region. one member of the restionaceae, namely, ischyrolepis schoenoides, and eight members of the ericaceae occur on platberg. the vegetation (table 1) shows more grass species than fynbos but, despite this, it is suggested that platberg fynbos falls within drakensberg-amathole afromontane fynbos (gd6) vegetation as defined by mucina and rutherford (2006) and that it should be regarded as the species-poor passerina montana subunit as described by mucina and rutherford (2006), forming an outlier of fynbos-like shrubland embedded within the broader grassland biome. the floristic collections plus the vegetation-community analysis shows platberg to be an important centre for biological diversity, with high species richness and a variety of habitats and complex ecosystems. it is important then that it should be given priority as a conservation site, with the development of a management plan to carry out this task. acknowledgements national geographic (grant number 7920-05) for the generous funds, without which the fieldwork and study would not have been possible. references acocks, j.p.h. 1952. veld types of south africa. 1st ed. memoirs of the botanical survey of south africa, 28: 1–192. acocks, j.p.h. 1988. veld types of south africa. 3rd ed. memoirs of the botanical survey of south africa, 57: 1–146. anderson, j.m. (ed.). 2001. towards gondwana alive. pretoria: national botanical institute. barrett, a.s., brown, l.r., barrett, l. and henzi, s.p. 2006. phytosociology and plant community utilisation by vervet monkeys of the blydeberg conservancy, limpopo province. koedoe, 49(1): 49–68. behr, c.m. and bredenkamp, g.j. 1988. an analysis of the flowering plants and ferns of the witwatersrand national botanic garden. south african journal of botany, 54(6): 515– 524. bester, s.p. 1998. vegetation and flora of the southern drakensberg escarpment and adjacent areas. unpublished msc thesis, university of pretoria, pretoria. bredenkamp, g.j. and brown, l.r. 2003. a reappraisal of acocks’ bankenveld: origin and diversity of vegetation types. south african journal of botany, 69(1): 7–26. brown, l.r., marais, h., henzi, s.p. and barrett, l. 2005. vegetation classification as the basis for baboon management in the bourke’s luck section of the blyde canyon nature reserve, mpumalanga. koedoe, 48(2): 71–92. carbutt, c. and edwards, t.j. 2004. the flora of the drakensberg alpine centre. edinburgh journal of botany, 60(3): 581–607. carbutt, c. and edwards, t.j. 2006. the endemic and nearendemic angiosperms of the drakensberg alpine centre. south african journal of botany, 72: 105–132. clever, g., brown, l.r. and bredenkamp, g.j. 2005. the phytosociology of the vermaaks, markewicks and buffelsklip valleys of the kammanassie nature reserve, western cape. koedoe, 48(1): 1–16. cowling, r.m. 1992. (ed.). the ecology of fynbos. nutrients, fire and diversity. cape town: oxford university press. cowling, r.m., holmes, p.m. and rebelo, a.g. 1992. plant diversity and endemism, in r.m. cowling (ed.). the ecology of fynbos. nutrients, fire and diversity. cape town: oxford university press. du preez, p.j. 1991. a syntaxonomical and synecological study of the vegetation of the south-eastern orange free state and related areas with special reference to korannaberg. unpublished phd thesis, university of the orange free state, bloemfontein. du preez, p.j. 1992. the classification of the vegetation of korannaberg, eastern orange free state. i. afromontane fynbos communities. south african journal of botany, 58(3): 165–172. du preez, p. j. and bredenkamp, g.j. 1991. vegetation classes of the southern and eastern orange free state, (republic of south africa) and the highlands of lesotho. navors. nas. mus. bloemfontein 7: 477 – 526. du toit, a.l. 1956. the geology of south africa. edinburgh: oliver and boyd. eckhardt, h.c.n., van rooyen, n. and bredenkamp, g.j. 1993. the vegetation of the north-eastern orange free state, south africa: physical environment and plant communities of the ea land type. bothalia, 23(1): 117–127. eckhardt, h.c.n., van rooyen, n. and bredenkamp, g.j. 1995. the grassland communities of the slopes and plains of the north-eastern orange free state. phytocoenologia, 25(1): 1–21. eriksson, p.g. 1985. the depositional palaeoenvironment of the elliot formation in the natal drakensberg and northeastern orange free state. geology society, 88: 19–26. everard, d.a. 1986. the effects of fire on the podocarpus latifolius forests of the royal natal national park, natal drakensberg. south african journal of botany, 52: 60–66. germishuizen, g. and meyer, n.l. (eds.). 2003. plants of southern africa: an annotated checklist. strelitzia, 14: 1–1231. goldblatt, p. and manning, j. 2000. cape plants. a conspectus of the cape flora of south africa. strelitzia, 9: 1–743. grobler, c.h., bredenkamp, g.j. and brown, l.r. 2002. natural woodland vegetation and plant species richness of the urban open spaces in gauteng, south africa. koedoe, 45(1): 19–34. grobler, c.h., bredenkamp, g.j. and brown, l.r. 2005. primary grassland communities of the urban open spaces in gauteng, south africa. south african journal of botany, 72: 367–377. haaksma, e.d. and linder, h.p. 2000. restios of the fynbos. cape town: botanical society of south africa. heini, m., silva, j. and tacheba, b. 2004. vegetation changes after a single fire-event in the okavango delta wetland, botswana. south african journal of botany, 70(5): 695–704. 212 afromontane fynbos in platberg original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 202 213 hennekens, s.m. 1996a. turboveg. a software package for input, processing and presentation of phytosociological data. user’s guide, version 2001. wageningen: ibn-dlo. hennekens, s.m. 1996b. megatab. a visual editor for phytosociological tables. ulft: giesen hill, m.o. 1979a. twinspan: a fortran program for arranging multivariate data in an ordered two-way table by classification of the individuals and attributes. new york: cornell university, ithaca. hill, m.o. 1979b. decorana: a fortran program for detrended correspondence and analysis and reciprocal averaging. new york: cornell university, ithaca. hill, t.r. 1996. description, classification and ordination of the dominant vegetation communities, cathedral peak, kwazulu-natal, drakensberg. south african journal of botany, 62: 263–269. hilliard, o.m. and burtt, b.l. 1987. the botany of southern natal drakensberg. cape town: national botanical gardens. hoare, d.b. and bredenkamp, g.j. 2001. syntaxonomy and environmental gradients of the grassland of the stormberg/ drakensberg mountain region of the eastern cape, south africa. south african journal of botany, 67: 595–608. kent, m. and coker, p. 1992. vegetation description and analysis: a practical approach. chichester: j. wiley & sons. killick, d.j.b. 1963. an account of the plant ecology of the cathedral peak area of the natal drakensberg. memoirs of the botanical survey of south africa, 34: 1–178. killick, d.j.b. 1978. the afro-alpine region. in m.j.a. werger (ed.). biogeography and ecology of southern africa. the hague: junk, pp. 515–560. körner, c. 2003. alpine plant life. functional plant ecology of high mountain ecosystems. berlin: springer-verlag. land type survey staff. 1991. land types of the map 2828 harrismith. pretoria: department of agriculture. loock, j.c., praekelt, h.e., van der westhuizen, w.a. and welman, j. 1991. the dewetsdorp-thaba nchu mountain excursion. bloemfontein: national museum. low, a.b. and rebelo, a.g. (eds.). 1996. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. macarthur, r.h. and wilson, e.o. 2001. the theory of island biogeography. princeton: princeton university press. malan, p.w. 1998. vegetation ecology of the southern free state. unpublished phd thesis, university of the orange free state, bloemfontein. matthews, w.s. 1991. phytosociology of the north-eastern mountain sourveld. unpublished msc thesis, university of pretoria, pretoria. moffett, r.o., daemane, m.e. pitso, t.r., lentsoane, r. and taoana, t.r.n. 2001. a checklist of the vascular plants of qwa-qwa and notes on the flora and vegetation of the area. uniqwa research chronicles, 3(2): 32–83. moon, b.p. and darids, g.f. (eds.). 1992. the geomorphology of southern africa. halfway house: southern book publishers. mucina, l. and rutherford, m.c. 2006. the vegetation of south africa, lesotho and swaziland. strelitzia, 19: 1–807. mueller-dombois, d. & ellenberg, h. 1974. aims and methods of vegetation ecology. new york: j. wiley & sons. mutke, j., kier, g., braun, g., schultz, c.h.r. and barhlott, w. 2001. patterns of african vascular plant diversity: a gisbased analysis. syst. geogr. pl., 71: 1125–1136. parmentier, i., oumorou, m., porembski, s., lejoly, j. and decocq, g. 2006. ecology, distribution and classification of xeric monocotyledonous mats on inselbergs in west africa and atlantic central africa. phytocoenology, 36(4): 547–564. perkins, l., bredenkamp, g.j. and granger, j.e. 1999. the phytosociology of the incised river valleys and dry upland savannah of the southern kwazulu-natal. south african journal of botany, 65(5/6): 321–330. peters, r.l. 1992. conservation of biological diversity in the face of climatic change. in r.l. peters, & t.e. lovejoy (eds.) global warming and biological diversity. new haven: yale university press, pp. 15–30. pond, u., beesley, b.b., brown, l.r. and bezuidenhout, h. 2002. floristic analysis of the mountain zebra national park, eastern cape. koedoe, 45(1): 35–58. pooley, e. 2003. mountain flowers. a field guide to he flora of the drakensberg and lesotho. the flora publishing trust, durban. porembski, s. and brown, g. 1995. the vegetation of inselbergs in the comoé national park (ivory park). conservatoire et jardin botanicues de geneve, 50(2): 351–365. robbeson, r.a.j. 1998. phytosociology of northwestern kwazulunatal. msc thesis, university of pretoria, pretoria. roberts, b.r. 1969. the vegetation of the golden gate highlands national park. koedoe, 12: 15–28. sarthou, c. and villiers, j-f. 1998. epilithic plant communities on inselbergs in french guiana. journal of vegetation science, 9: 847–860. schulze, r.e. 1997. south african atlas of agro hydrology and climatology. pretoria: water research commission. scott, l. 1988. late quaternary vegetation history and climatic change in the eastern orange free state, south africa. south african journal of botany, 55(1): 107–116. seabloom, e.w. and richards, s.a. 2003. multiple stable equilibrium in grassland mediated by herbivore population dynamics and foraging behaviour. ecology, 84(11): 2891–2904. segal, s. 1969. ecological notes on wall vegetation. phd thesis, university of amsterdam, amsterdam. soil classification working group, 1991. soil classification. a taxonomic system for south africa. memoirs on the agricultural natural resources of south africa. no 15. dept. of agricultural development, pretoria, pp. 1–262. smit, c.m., bredenkamp, g.j. and van rooyen, n. 1992. phytosociology of the ac land type in the foothills of the low drakensberg in north-western natal. south african journal of botany, 59: 203–214. smit, c.m., bredenkamp, g.j. and van rooyen, n. 1995. the grassland vegetation of the low drakensberg escarpment in the north-western kwazulu-natal and north-eastern orange free state border area. south african journal of botany, 61: 9–17. south africa weather services. 2007. climate of south africa. climate statistics for bethlehem from 1961-1990. wb 40. pretoria: government printer. taylor, d. 1996. mountains. in: w.m. adams, a.s. goudie & a.r. orme (eds.) the physical geography of africa. oxford: oxford university press, pp. 287–306. truswell, j.f. 1970. an introduction to the historical geology of south africa. london: purnell press. van wyk, a.e. and smith, g.f. 2001. regions of floristic endemism in southern africa: a review with emphasis on succulents. pretoria: umdaus press. van zinderen bakker jr, e.m. 1973. ecological investigation of forest communities in the eastern orange free state and the adjacent natal drakensberg. vegetatio, 28: 299–334. westhoff, v. and van der maarel, e. 1980. the braun-blanquet approach. in: r.h. whitaker, (ed.) classification of plant communities. the hague. junk: kluwer academic publisher, pp. 287–378. whittaker, r.h. (ed.). 1980. classification of plant communities. the hague: junk: kluwer academic publisher. white, f. 1983. the vegetation of africa. volume xx, a descriptive memoir to accompany the unesco/aetfat/unso vegetation map of africa. paris: unesco. 213 abstract introduction data description methods results discussion conclusion acknowledgements references about the author(s) kai heckel department for earth observation, faculty of chemistry and earth sciences, friedrich schiller university, jena, germany earth observation services (eos), jena, germany marcel urban department for earth observation, faculty of chemistry and earth sciences, friedrich schiller university, jena, germany jean-sébastien bouffard catalyst, ottawa, canada jussi baade department for physical geography, faculty of chemistry and earth sciences, friedrich schiller university, jena, germany peter boucher department of organismic and evolutionary biology, harvard university, cambridge, united states of america andrew davies department of organismic and evolutionary biology, harvard university, cambridge, united states of america evan g. hockridge department of organismic and evolutionary biology, harvard university, cambridge, united states of america wolfgang lück pinkmatter north america, victoria, canada jonas ziemer department for earth observation, faculty of chemistry and earth sciences, friedrich schiller university, jena, germany izak smit scientific services, south african national parks (sanparks), skukuza, south africa centre for african ecology, school of animal, plant and environmental sciences, university of the witwatersrand, johannesburg, south africa bernhard jacobs geospace international, pretoria, south africa mark norris-rogers rastermatics international, umgungundlova, south africa christiane schmullius department for earth observation, faculty of chemistry and earth sciences, friedrich schiller university, jena, germany citation heckel, k., urban, m., bouffard, j.-s., baade, j., boucher, p., davies, a. et al., 2021, ‘the first sub-meter resolution digital elevation model of the kruger national park, south africa’, koedoe 63(1), a1679. https://doi.org/10.4102/koedoe.v63i1.1679 original research the first sub-meter resolution digital elevation model of the kruger national park, south africa kai heckel, marcel urban, jean-sébastien bouffard, jussi baade, peter boucher, andrew davies, evan g. hockridge, wolfgang lück, jonas ziemer, izak smit, bernhard jacobs, mark norris-rogers, christiane schmullius received: 20 mar. 2021; accepted: 01 oct. 2021; published: 17 dec. 2021 copyright: © 2021. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the use of digital elevation models has proven to be crucial in numerous studies related to savanna ecosystem research. however, the insufficient spatial resolution of the chosen input data is often considered to be a limiting factor when conducting local to regional scale ecosystem analysis. the elevation models and orthorectified imagery created in this study represent the first wall-to-wall digital elevation data sets produced for the kruger national park (knp), south africa, at very high spatial resolution. using colour-infrared (cir) aerial imagery from the archives of the chief directorate: national geo-spatial information (cdngi), department of agriculture, land reform and rural development (dalrrd) aerial acquisition programme, we created digital surface models (dsms), digital terrain models (dtms) and cir orthomosaics covering the entire knp with a nominal ground sampling distance of 0.25 m. elevation information was derived using state-of-the-art stereo matching algorithms that utilised semi-global matching (sgm) as a cost aggregation function throughout the image pairing, using the enterprise software from catalyst. the final products were validated against reference products, and showed excellent agreement with r² values of 0.99. further, the validation of the dtm and dsm revealed median absolute vertical height error (le90) across all sites of 1.02 m and 2.58 m, respectively. the orthomosaics were validated with in situ ground control points (gcps) exhibiting a horizontal circular probable error (cpe) of 1.37 m. the data resulting from this work will be distributed freely with the aim of fostering more scientific studies in the african science community and beyond. conservation implications: accurate information about terrain and surface height are crucial inputs to a variety of scientific analysis, which are essential in protected areas, such as flood prediction or fire hazard estimation. elevation data sets and orthomosaics in very high resolution can therefore serve as a crucial tool to improve park management and foster positive implications on conservation efforts. keywords: kruger national park; elevation; aerial imagery; semi-global matching; stereoscopy; 3d; protected areas. introduction since their introduction in 1958, digital elevation models (dems) have been used widely in fields such as ecology, hydrology, other earth-related disciplines, modelling and remote sensing (miller & laflamme 1958). a dem is a digital representation of a topographic surface that is defined through a two-dimensional discrete function of a morphometric variable and consists of xyz coordinates (florinsky 2012). their accurate derivation is relevant not only from a scientific but also a socio-economic point of view. originally, dems were created through classic field surveys. in recent decades, height structure analyses were mostly carried out using elevation information derived using remote sensing techniques such as photogrammetry based on stereoscopic imagery (pieczonka, bolch & buchroithner 2011; pulighe & fava 2013), light detection and ranging (lidar) (lim et al. 2003; lindberg & holmgren 2017) or air-/spaceborne interferometric synthetic aperture radar (insar) (hirt, filmer & featherstone 2010; nelson, reuter & gessler 2009). on local to regional scales, aerial photos and lidar collected by unmanned aerial vehicles (uavs) or airborne instruments are, because of their comparably lower degree of complexity, more commonly used than insar procedures from satellite-derived imagery. since lidar data sets represent a direct physical measurement of height information, they are known to be more accurate than dems derived from aerial images (baltsavias 1999). this arises mainly from the ability of full-waveform laser scanning systems to detect entire vertical structures of objects situated aboveground and to penetrate tree canopies (reitberger, krzystek & stilla 2008). major disadvantages of lidar campaigns are the costs of data acquisition, which are significantly higher than for flight missions acquiring aerial imagery (white et al. 2013), as well as the missing physical relation between plant structures and the radiation in comparison to true colour (rgb) or colour-infrared (cir) data. photogrammetric methods have been used for more than a century to retrieve height information from aerial imagery. while they are now processed digitally, these methods still use the same fundamental principles of parallax measurement, taking into account different viewing angles of overlapping image pairs (straub et al. 2013). because of the lower flight altitude compared to satellite imagery, airborne flight acquisitions are able to reach higher spatial resolutions with adjustable overlap, and thus are less impacted by vertical as well as horizontal uncertainties. the high degree of overlap presents multiple measurements of the same point at different parallaxes with multi-angular observations, leading to more precise elevation measurements, and a more rigorous triangulation via a block bundle adjustment. furthermore, the development of innovative techniques has increased the accuracy of elevation models that were calculated from airborne imagery in recent years (reitberger et al. 2008; stepper, straub & pretzsch 2015). these include innovations in computer vision technologies, such as the improvement of dense image matching algorithms, fostering new research using stereoscopic imagery from uavs and airborne sensors for forest applications (haala et al. 2010; straub et al. 2013; tian et al. 2017). global-scale data sets from dems have existed for decades, and many are now freely available. however, these global data sets provide only medium to coarse resolution products (≥ 30 m cell size), and fine-resolution dems are required to study local topographic phenomena at the regional level. high-resolution products are usually not freely available in most areas of the world. in protected areas, such as the kruger national park (knp), reference data are usually scarce, yet the need for high precision and (very) high resolution elevation models is great. several studies have indicated the need for such data products to minimise biases in vegetation modelling (o’loughlin et al. 2016) and flood management (wu et al. 2017). this study aims to deliver the first publicly available very high resolution (vhr) elevation models and orthomosaics of the knp derived from photogrammetric algorithms using freely available aerial imagery. we introduce and discuss three products: (1) a digital surface model (dsm), (2) a digital terrain model (dtm), and (3) an orthomosaic data set. a dsm represents the earth’s surface including all aboveground features (such as vegetation or buildings), while a dtm characterises the elevation of bare ground underneath these features. the orthomosaic data sets represent a mosaic of aerial images that were corrected for geometric effects, such as perspective distortions and topographic influences. the final products were produced with pixel postings between 0.25 m and 5 m to provide the basis for a variety of ecosystem analysis applications. in addition, to allow users to utilise data products at a lower resolution, we resampled the data to 1 m and 5 m, as lower resolution products may suffice for specific types of applications (e.g. modelling vs. habitat mapping). data description study area the knp comprises protected land areas to the extent of approximately 19 500 km², making it south africa’s largest protected area. it is located in the northeastern part of south africa, bordering mozambique in the west and zimbabwe in the north. elevations vary between 260 m and 839 m above sea level. the knp has a mean annual precipitation of around 600 mm, and exhibits a gradient of rainfall that increases from north to south (gertenbach 1980). precipitation occurs heterogeneously over time and space. the climate alternates between contrasting seasons, with a wet and hot season (usually october to march) as well as a dry and mild season (april to september). geologically, the park can be divided into two parts: (1) the basaltic formations (nutrient-rich) in the east, and (2) the granitoid bedrock (nutrient-poor) in the west (venter 1990). the characteristic flora of the knp is dominated by woody vegetation and herbaceous plants, with canopy cover ranging between 10% and 80% (venter, scholes & eckhardt 2003). vegetation is largely influenced by geology and rainfall, as well as disturbances like fire and herbivory. aerial imagery to cover the entire knp, an area of approximately 19 500 km², image data from 12 image acquisition tiles, also referred to as ‘entities’, were collected through flight campaigns accomplished at the end of the dry season in 2018. entities comprise between 500-700 aerial images, each covering an area of roughly 20 km² adding up to an extent of approximately 60 by 60 km per tile, which can differ for some areas. the cir raster data sets were stored in geotiff file format and included three channels: near-infrared (nir) (830 nm, red (652 nm) and green (544 nm). an overview of the aerial campaign entities is provided in figure 1. figure 1: leica digital mapping camera coverage of the kruger national park, south africa represented as individual flight strip centroids for each acquisition tile. the aerial images were acquired with the leica digital mapping camera (dmciii) instrument which was utilised within the flight campaigns of the south african national mapping agency chief directorate: national geo-spatial information (cdngi), department of agriculture, land reform and rural development (dalrrd) aerial acquisition programme. the dmciii consists of eight individual monochromatic cameras, four pan cameras, and one for each spectral band. each camera features its individual interior orientation (io). during the calibration processing (level 1 processing), images were projected from their io to that of an idealised and common io (or camera model). pan bands were mosaicked, and multispectral data were stacked. pan and multispectral data were then fused to create a pan-sharpened product. the level 1 processing software uses a brovery image fusion approach, allowing that only three spectral bands can be fused at a time and multispectral bands poorly correlated with the pan band get radiometrically distorted. the imagery gets captured in 12 bit and delivered in either 8 or 16 bit (16 bit imagery was used in this study). data sets were taken from an altitude of approximately 5500 m – 6000 m above ground. the flight campaigns were carried out by geospace international (pty) ltd. latest data for the whole country of south africa have been acquired at 0.5 m and 0.25 m pixel posting for the periods 2008–2016 and 2017–present, respectively. the imagery was acquired in september/october 2018 using a cir channel combination using the nir, red and green bands. analysis was then carried out at a pixel posting of 0.25 m. validation data to validate the products that were derived in this study, various independent data sources were used. for the elevation models, we utilised different types of height measurements as reference. in order to validate the height at ground level (dtm), we used differential global navigation satellite system (gnss)-based ground survey points that were collected during field campaigns between february 2014 and march 2016 using leica gs10 and gs15 gnss receivers (baade & schmullius 2016). the roughly 10 000 ground measurements were stratified over various land cover types following a transect sampling strategy and filtered to prevent the usage of points that were situated inside water bodies, which were dried out during the time of acquisition. for the validation of the dsm, we utilised vhr lidar uav data. data sets were collected at 100 m altitude above ground over numerous sites across the knp, primarily focusing on experimental burn and herbivore exclosure plots, during january and february 2020 using a riegl vux-1 lr instrument attached to a dji m600 pro uav. these sites covered a diverse set of vegetation types and structures, ranging from open to dense and from short to tall, thus providing an ideal data set for validation. the average size of each site was 110 ha with a flight overlap of 60% and a pulse repetition rate of 820 khz, resulting in an average point density of ~150 points per m². terrain information from the shuttle radar topography (srtm) mission was used in flight planning to homogenise lidar point density and beam divergence across the study area. in post-processing, uav trajectories were refined with base station data, denoised, classified (axelsson 2000), and matched together. rasterised dsms were created by calculating the maximum elevation of all 1st return points within each 0.25 m cell. after processing, the accuracy of the point clouds, and thus the dsm elevations, was expected to be ~0.10 m or less. the point clouds were pre-processed to calculate dsms at a cell resolution of 0.25 m. the locations of all sites that were used to validate the dmc surface model are shown in figure 2. figure 2: distribution of reference data sets used to validate the elevation models for kruger national park, south africa. in order to validate the horizontal precision of the orthomosaics, we relied on the geolocation of triangulation stations, or ‘trig’ beacons. shapefiles were provided by cdngi with the exact easting and northing position, as well as the appearance and height of the trig object (e.g. pole). in total, more than 50 locations were used to evaluate the accuracy of the orthorectified aerial imagery (see figure 2). methods image processing with catalyst enterprise to extract height information from the stereoscopic dmc data, we utilised catalyst’s (formerly known as pci geomatics) commercial enterprise software environment, which is a geospatial image-processing software designed to leverage complex computing hardware and to be scalable, using the available computational resources efficiently. a major advantage is its underlying python programming framework, enabling the development of new tools and the optimised management of resources (catalyst 2020). using catalyst professional’s archive of algorithms, a wide variety of data manipulation methods can be used for elevation extraction and orthomosaic creation. thus, we created a workflow to implement different processing and quality assessment steps. these are automated within enterprise, and thus could be easily applied to other study sites and data sets. to derive height information from stereoscopic aerial imagery, we followed the workflow outlined in figure 3. the processing steps made use of state-of-the-art routines that are applied to carry out height extraction algorithms on stereoscopic images from uavs, airplanes, or satellites. figure 3: workflow to extract elevation information from digital mapping camera aerial imagery using catalyst enterprise. data preparation as visualised in figure 3, the ingestion of aerial image data into the enterprise production system could only be accomplished once metadata had been prepared. the metadata provide crucial information derived from the physical characteristics of the acquisition platform, in this case inertial measurement unit (imu) and differential global positioning system (gps) of an airplane. they include image centre position (easting & northing), flight altitude, as well as image orientation (ѡ), phi (ϕ) and kappa (к), which determine the orientation of images through three different axes. these variables are essential inputs for all aerial image processing. for this study, the exterior orientations (eo) calculated by cdngi were used. once all available input information has been properly prepared, the data can be ingested into the enterprise production system. bundle adjustments and tie point collection the eo model of each image was refined for the purpose of improving the accuracy of the extracted dsms and the ortho images. this was accomplished by collecting tie points and performing block bundle adjustments. the selection of tie points was carried out using a gridded sampling approach with 150 samples per overlapping area using the nir channel as the matching variable between image pairs. this bandwidth was chosen because it is less sensitive to illumination differences, and thus more robust in terms of atmospheric effects (zhu et al. 2019). a fast fourier transform phase matching (fftp) approach (brigham 1988) was utilised to automatically identify tie points between aerial images. this approach identifies points based on the relative shift between images (the phase difference in the frequency domain) as the criteria for point matching. if the correlation coefficient between two matching points was below a given threshold (0.85) within a search radius of five pixels, these points were rejected for the adjustment process, which further increased the quality of the matching process between adjacent images. extraction and filtering of a digital elevation model to generate dems from airborne imagery, images needed to be projected from sensor to an epipolar geometry, with y coordinates of overlapping pairs perfectly aligned and the remaining parallax represented in the x coordinates of the resulting data set, from which a measured parallax is then translated into elevation. the threshold for the minimum overlap between images was chosen to be at least 40% and the nir channel was selected as the input for the epipolar generation. this bandwidth was chosen because the nir band preserves contrast, and thus continues to exhibit bright reflectances over both bare surfaces and vegetation. the nir channel is also less affected by haze and atmospheric effects, although its effective resolution is poorer than that of the shorter wavelength because of diffraction. any remaining misalignment identified in the epipolar geometry was modelled and quantified to improve the subsequent dsm extraction procedure. generally, height extraction from airborne data was carried out through several steps including matching cost computation, cost aggregation, disparity optimisation and refinement procedures. in recent years, semi-global matching (sgm) approaches have proven to be amongst the most popular and successful algorithms in the fields of stereo vision and photogrammetry (klette et al. 2011; michael et al. 2013). to extract the height information from the aerial imagery, we used this matching approach, which utilises intensity differences, mutual information (as the cost function) and an approximation of the global energy function that is being optimised path-wise (16 paths in this study) from all directions over the image. the cost function is significantly influenced by the use of penalty values, which were chosen based on performance tests and represent varying magnitudes of disparity changes. these variables have a strong impact on the matching performance and the robustness that is related to this processing step. the term ‘semi-global’ arises from the combination of both global and local methods in a way that the complexity of the process is lowered, and the quality of the matching is drastically improved. while the computation time for global methods is often considerably higher, the overall performance increases compared to local matching algorithms. further, pixel-wise calculated matching cost, contrary to the calculation along image paths, poses negative effects of insufficient correspondences related to low texture and ambiguity (hirschmüller 2007). to minimise undesirable features in the elevation model, filtering is a necessary step in the extraction of height information. to minimise noise originating from correlator accuracy and to improve the accuracy of the extracted dsm, a series of spatial filters were applied. an intermediate level of filtering strength was used, this includes: noise filter, hybrid median filter, and a bi-level smoothing filter. the noise filter performs two operations: (1) a neighbourhood analysis to identify blunders by examining statistics, and (2) analysis to identify blunders by pinpointing pixels surrounded by failed pixel matches. the hybrid median filter is used to smooth noise after the interpolation and rasterisation of the correlation point cloud. this version of the median filter has been modified to preserve natural edge features and corners (e.g. trees, buildings), while still reducing noise. the bi-level smoothing is applied to further reduce noise, while preserving abrupt and steep elevation changes throughout the dsm. in addition to these spatial filters, a pattern suppression technique was applied to minimise artefacts caused by strong patterns in the landscape (e.g. rows in agricultural fields). this method examines characteristics of the accumulated costs to infer a blunder. after filtering, the dem extraction for each tile was completed. the impact of these filtering algorithms on the final spatial resolution of the dsm is believed to be minimal. this assumption is based on the evidence that the noise filter has no impact, and the hybrid median and bi-level smoothing may have only a minimal impact because of their small neighbourhood size and feature preserving attributes. digital terrain model conversion the derivation of bare earth elevation data requires the removal of all elements that are above the ground surface (such as buildings or vegetation) of the dem. an adaptive slope-based filtering approach was applied to remove surface features from the raster data set. to allow maximum flexibility in the elimination of aboveground elements, the filtering process was customised by defining object sizes (100 pixels) to remove surface features, as well as a small gradient threshold to preserve hills and natural slopes (slopes above five degrees of gradient were filtered). additionally, filtering steps were introduced to remove unwanted bumps and pits (filter window of 15 pixels and a slope of five degrees) in the data that may persist from prior object removals. thresholds for these filters and surface object extents were fixed for the entire knp to retrieve height information with consistent parameters. the application of filters to smoothen out remaining objects that are situated above ground (above dtm height) led to a certain degree of blending of the height statistics of some of the pixels within their neighbourhood in the magnitude of the described window size. during this filtering step, it can be assumed that the height of the terrain surface was not altered as much as that of the bumps and pits, which were the main objective of filtering in this processing step. orthorectification of aerial photos to create orthorectified aerial images with exact geolocations, the previously created height models were used to correct the flight strips for effects of terrain. in order to use the cir aerial imagery for land cover and classification analysis, the aerial imagery needed to be orthorectified and prepared for image mosaicking. because of its high spatial detail, it was essential to geolocate the data as precisely as possible. the main goal of orthorectification is the correction of view angle and relief-induced effects, as well as the georeferencing of the aerial scenes. thus, the elevation models with a pixel size of 0.25 m served as elevation input to remove topographical effects from the imagery. the conical rays in unrectified aerial images were converted to parallel rays, which made them orthogonal to the earth’s surface (jensen 2007). mosaicking for each entity presented in figure 1, all available flight paths were merged to create a single seamless mosaic of elevation models and orthomosaics. the so-called ‘image stitching’ of the elevation models was based on two main components, cutline computation and colour balancing, which minimised edge effects that could have occurred in the combined data set between the borders of neighbouring tiles. firstly, cutlines were created between all adjacent images using a criterion of minimum relative difference in grey value levels in the overlapping area of neighbouring images. these boundaries define at which location adjacent images are combined to form seamless mosaics. next, the radiometric differences within the image were reduced by applying colour balancing algorithms, which balanced histograms of the areas of overlap between the flight strips. the colour balancing process is important to remove possible patchwork effects from the mosaic. thus, contrasts are levelled out and seams were removed, which positively impacted the image statistics. artefact elimination to remove artefacts from the elevation models that remained after height extraction, filter techniques were applied to the data. larger water bodies such as dams, lakes, or other standing water bodies led to faulty values in the elevation models, which were located in the greater negative feature space. this effect occurs because the algorithm is not able to detect tie points to connect adjacent aerial images over water surfaces, resulting in incorrect elevation values in these areas. because of the changing appearance of the water surface and the very low contrast, elevation estimates over water are unreliable and highly variable. to account for these effects and remove them from the data sets, we used an edge filter to fill pits within water bodies using the elevation value of the surrounding bank areas. besides the removal of water bodies, which could not be captured by the sgm algorithm, no post-processing filtering techniques were applied to the data. validation elevation models a comparison between the highly precise in situ samples (from the gnss ground points and uav-lidar data) with the modelled elevation (dtm and dsm) from the dmc data sets was carried out using the absolute pixel-wise height difference δh, which quantifies the deviation between the reference height information (hr) and the respective target elevation model (hdmc). thus, a vertical error is measured relative to the known height. consequently, the height differences δhdtm and δhdsm for dtm and dsm, respectively, are given as: since the analysis of the height deviation underlies a number of sources of uncertainty, further height measures were calculated to provide a substantial representation of the differences between data sets. besides the pixel-specific height deviation, we calculated the median δhp50 of the height deviation, the standard deviation (sd), the 95th percentile, and the median of the 90th percentile of the absolute linear vertical error (le90) of all individual reference sites. additionally, we calculated the confidence intervals δhci (95%) for δh. orthomosaics to quantify the horizontal accuracy of the geolocation of the orthorectified aerial imagery we used trigonometrical beacons as reference data. these are relatively evenly distributed across the knp, allowing a fairly good representation of the area. in total, 89 trig stations were utilised, which vary in their appearance from beacons installed on hilltops to well-known man-made structures, such as outlooks or buildings. they are part of the physical framework for south africa’s geodetic reference, and will be replaced with 50 gps base stations in the next few years. however, these trig stations represented the most evenly distributed and precise source for geolocation accuracy assessments available for the knp available for this study. in order to quantify the horizontal accuracy of the ortho imagery, we computed several statistical measures. first, the single ortho images were merged by creating a virtual raster (vrt) data set using geospatial data abstraction library (gdal) for the knp. the vrt file format is a gdal driver, allowing the computation of fully functional (statistically) raster stacks without the need to write and export the extensive data set to common raster file formats such as ‘.tif’ or ‘.pix’. this way, valuable disk space can be saved, while also speeding up the validation process (gdal/ogr contributors 2021). second, the vrt’s estimated location of the trig stations was acquired. finally, several statistics were calculated to quantify the horizontal error in x and y direction, using the geographic coordinates. the equations for these statistics are provided in table 1 and visualised in figure 4. figure 4: overview of circular error measures; error measures given: three-five sigma error, circular map accuracy standard, mean square positional error, circular probable error, mean circular standard error, directional standard error of x and y. table 1: statistical measures used for the validation of the orthomosaics. results products digital terrain model, dsm and orthomosaics were created covering the entire knp. cell sizes of the products are 0.25 m, 1 m and 5 m. all products are projected to utm zone 36, wgs84 using the wgs84 ellipsoid also as the vertical datum (if vertical information is stored). digital elevation models and orthomosaics were partitioned according to the cdngi 1:50 000 map sheet units for more convenient download. each unit covers an area of maximum 640 km² or 25 km in x and y direction (depending on the overlay with knp borders). all data sets are freely available through the website of the centre for environmental data analysis (ceda), and can be accessed as described in the data availability statement. digital elevation models as described earlier, two types of dems were calculated: dsms and dtms. both products were filtered using the same parameters in the post-processing phase and cropped to the boundary of the knp. elevation spanned from 105 m to 832 m above sea level as well as 106 m to 833 m above sea level for the dtm and dsm, respectively. the median height value for the knp dtm and dsm was 333.3 m and 333.4 m, respectively. from the surface height model, features below the horizontal extent of four pixels can be identified. this allows for a minimum mapping unit (mmu) up to a meter or less. examples of objects which can be located in the height data set are smaller trees as well as distinctive bushes, which are typical for the savanna environment. figure 5 visualises both elevation models using an identical colour range as well as according to their orthomosaic subset. as visualised in the exemplary zoomed windows in the figure, the dsm allows the identification of individual trees and bushes, especially when they are foliated. subset 2, which displays the village of skukuza, demonstrates the precise delineation of single houses and other artificial structures. the dtm displays a very good representation of the terrain height after the removal of objects above the ground. because of the large extent of the study area (~19 500 km²), some artefacts may not have been removed by the dtm filtering algorithms completely, and thus remain in the final products partially or to their full extent. figure 5: orthomosaic (upper), digital surface model (middle) and digital terrain model (lower) generated from aerial imagery for four different sites across the kruger national park at a scale of 1:5000. orthorectified aerial imagery the orthomosaics were created using cir ortho images as well as the previously described dsm at 0.25 m cell size. after the mosaicking process, the ortho images were combined to form a seamless orthomosaic of the original leica dmciii tiles. thus, the results enable the analysis of savanna ecosystems at the scale of an individual tree or shrub. an exemplary visualisation of the great level of detail of the 0.25 m orthomosaic, located along a central stretch of the limpopo river close to the northern knp border, displaying different scales at which the data can be used is shown in figure 6. figure 6: orthorectified colour-infrared aerial imagery at different scales enabling local to regional analysis; single bushes and trees can easily be delineated. orthorectified imagery of this scale can be used in a variety of scientific analysis, including vegetation cover analysis (le bris, tassin & chehata 2013; lucas et al. 2000), biomass estimation (meng, pang & zhang 2014; shin et al. 2016), or the evaluation of the effects of severe flood events (ichim & popescu 2019). validation digital terrain model the results of the validation of the 0.25 m dtm are shown in figure 7. here, the relation between the dgnss-derived ground points and the modelled surface height from vhr aerial imagery is visualised. for the various locations across which dgnss points were collected, the agreement with the estimated elevation was found to be very strong, with a coefficient of determination (r²) of 0.99. local variations were found between the different acquisition areas. the median of the le90 for all in situ points was 1.02 m with a site-based (averaged for each location) minimum of 0.12 m, a maximum of 2.06 m, and a 95th percentile of 1.79 m. strongest deviations from the dmc-based terrain estimation were found near some of the reservoir sites. these sites were either dried out or at least only partially filled with water at the time of the dgnss data acquisition. the best agreement with the dgnss height estimations was found around the dried-out hartbeesfontein reservoir, north of the timbavati river, as well as in the sites that were located in and around skukuza. here, flat surfaces, such as a sports field or a runway, were measured during the field campaigns. figure 7: validation of the modelled terrain (digital terrain model) elevation from stereoscopic aerial imagery with ground measurements for entire area and lugmag reservoir sub-site. more than ten study sites were included in the validation (see figure 2 for the distribution of the sites). digital surface model similar to the validation of the dtm, the dsm was validated using a pixel posting of 0.25 m. the results are visualised in figure 8. the validation of the dsm was carried out using vhr lidar data that were acquired in january and february 2020, representing the end of the wet season. across all sample sites (with 250 validation points at each site), a trend of small underestimation of vegetation height was found in the modelled dsm. while this effect varied by location, it was present among all sites. similar to the derived dtm, the calculated dsm exhibited a very good overall agreement with the reference data exhibiting a coefficient of determination (r²) value of 0.99. however, similar to the 95th percentile of height deviation of 5.61 m, the le90 was found to be larger than the dtm error reference value with 2.58 m averaged (median) over all uav acquisition locations that are visualised in figure 2. here, the median of the deviation varied between 0.5 m and 2.6 m across the various lidar validation sites. the largest deviation between the uav-based reference data and the modelled surface height was found at the kambeni experimental burn plots (ebp), which are located near numbi gate, with a le90 height difference of 6.8 m. the best fit was found at the makhohlola exclosure plot near crocodile bridge, close to the mozambique border, exhibiting a deviation between reference and modelled data of 1.6 m. figure 8: validation of the modelled surface elevation (digital surface model) from stereoscopic aerial imagery with unmanned aerial vehicle-based lidar measurements for entire area and shabeni experimental burn plot sub-site. seven different locations were used for the validation. orthomosaic the visual inspection of the final orthorectified aerial images revealed very high agreement between existing and freely available ortho products. as described in the orthomosaics section, the geolocation accuracy of the orthomosaic was assessed using the precise location of several ground control points (gcps) (trig stations) that are distributed across the knp. the trig stations were solely used for a horizontal accuracy validation, not for the correction of the aerial images. in total, 58 validation points were used to analyse the geolocation. as visualised in figure 4, the individual circular error measures represent different levels of confidence (circular precision probability [cpp]). table 2 summarises the individual levels of error probabilities as described in the orthomosaics section. all validation points were found to be within a 2.52 m radius of the associated pixel in the orthomosaic with a probability of 90%. the accuracies at different in situ locations varied significantly because of multiple reasons (explained in discussion section). table 2: statistical measures describing the horizontal geolocation accuracy. discussion the elevation models derived in this study were found to exhibit a very strong agreement with the reference data sets described in data description section. both models exhibited r² values of 0.99 and sd of 0.76 m and 2.03 m for the dtm and dsm, respectively. thus, the different ecosystems and landscapes of the knp did not significantly influence the correlation between the reference data and the estimated elevation models. visual interpretation as well as statistical analysis of the vertical error in the derived dems showed a fairly good agreement between the high-resolution height information with the available reference data sets. while the in situ samples used for the validation of the elevation models were well-distributed across the entire knp, some areas were not covered sufficiently (e.g. mountainous areas). to further improve on the characterisation of the performance of the height estimations, more reference data would be desirable in the future, especially in areas with steep slopes. comparing the different types of elevation models, the dtm was found to feature a lower vertical deviation from the in situ dgnss data with a median le90 of 1.02 m. in contrast, the derived dsm, which was validated using uav-based lidar data, displayed a median le90 of 2.58 m. especially in areas with no or only sparse vegetation cover, the estimated dtm performed fairly well against the ground points. likewise, the dsm effectively captured the open savanna vegetation present in the makholola exclosure plot in the southeast of the knp, exhibiting a le90 of 1.6 m (open savannas on basaltic soils). the largest deviations of the dtm from the reference ground points were found around a reservoir south of the olifants river (central knp), with a le90 of 2.06 m. a number of possible sources of uncertainty underlie the accuracy of the dtm. for example, one source of uncertainty could be seasonal variation occurring between the acquisition date of the aerial imagery (end of the dry season) and the collection of terrain dgnss points (end of the wet season). also, seasonal events such as floods could have altered ground height between aerial and dgnss acquisitions. in open areas, such as those located around the knp water reservoirs, the presence of vegetation (e.g. grasses) can also negatively influence the comparison between the two acquisitions, as some objects above the ground may have not been filtered out properly in the dtm conversion process. the conversion from the dsm to the dtm was carried out using several filtering techniques and smoothing parameters, including gradient thresholds and moving-window bump and pit filters. the type and aggressiveness of filtering grids was tested and selected after visual and statistical interpretation. however, large structures, such as long walls, buildings, or bridges, might not have been completely removed from the dsm, thus transforming the dtm ground height as well. such remaining artefacts were likely to introduce differences in height values compared to the original gnss points. in addition to affecting the vertical accuracy, the filtering mechanisms also altered the horizontal dimensions of the data sets. as a result, the pixel posting does not necessarily equal the spatial resolution, as some of the filters calculated neighbourhood-based statistics, averaging the height values of single pixels in relation to their neighbouring pixels using the thresholds and parameters described earlier. further, the location of some dgnss samples on the tops or edges of dams possibly led to incorrect values in the estimated dtm, because of misinterpretations in the epipolar matching algorithms. savanna ecosystems are vulnerable to shifts in vegetation, as well as to the loss of biodiversity because of natural or anthropogenic causes (smit et al. 2010). thus, savannas can be attributed as dynamic environments, where seasonal differences, often reinforced by wildfires, hugely impact the growth of mostly woody vegetation. in contrast to the pure terrain elevation, the height of vegetated areas can change drastically over time, such as after extreme droughts, exceptionally moist wet seasons or bushfires. these seasonal events combined with herbivory effects (e.g. elephant impacts) foster resprouting, regrowth, and coppicing from stems and roots (asner & levick 2012; clarke et al. 2005; mograbi et al. 2017). consequently, seasonal differences can lead to errors in the deviation estimation between reference data and calculated height models, as data were collected 18 months apart. thus, the growth of woody vegetation, which varies between different acquisition dates of the reference data and the aerial imagery, can lead to height differences. the seasonal differences become even more visible when analysing δh in the estimated dsms that were located on or around the ebp, which are scattered across the knp. since these sites are frequently burned (burnings every one to six years and for some plots up to decades), vegetation patterns here are highly dynamic (chown 2010). however, the impact of these management practices also depends on environmental conditions, such as varying amounts of fire fuel or rainfall (biggs et al. 2003). the quality of the dsm could also have been affected as a result of the aerial imagery being acquired during the late dry season, when vegetation cover is generally sparse. open canopies, which are common for the deciduous plants of knp at that time of the year, can create challenges for the sgm matching algorithms. the algorithms will occasionally miss elevated pixels from open canopy vegetation during leaf-off season, instead capturing the ground underneath sparse canopies. thus, the algorithm may fail to capture accurate height information for certain trees during leaf-off season. these circumstances led to larger height reference deviations in the dsm, especially in the structurally more complex sites such as the kambeni burn plot, where the le90 was found to be highest with 6.8 m. while a data acquisition campaign during the wet season would be preferable in terms of image-matching success and, consequently, vegetation structure and plant vitality mapping (e.g. through chlorophyll content – nir interactions), wet season acquisitions are often not feasible as cloud cover heavily impedes flight missions carrying optical sensors (hoekman & varekamp 2001). besides artefacts that originated in the dsm filtering process, other effects, such as visible cutlines between entity borders or flight path edges, are still visible in some parts of the mosaic. this is especially true in areas with topographic features such as hills (or koppies) in the elevated parts of the knp, where some image paths can cause height differences between neighbouring pixels (but these differences rarely exceed 0.5 m). because of increased slope conditions in those areas, illumination effects, such as shadows from mountain ridges or hilltops, can severely impact the performance of the process of matching epipolar image pairs. these effects are only visible in very small portions of the elevation models and are, in most cases, negligible. however, in the case of fine-scale analyses, such as local watershed modelling, these small errors should be taken into account. further filtering steps should also take into account the spatial variability and textural heterogeneity of the knp. to provide the basis for the analysis of the high-resolution aerial imagery without terrain effects, orthomosaics at the original cell size of 0.25 m were derived. the comparison to the geolocation of trig stations, as described in the orthomosaics section, revealed a very good overall agreement between the orthorectified aerial imagery and the reference ground information. major sources of uncertainty for the validation of the orthorectified aerial imagery arise from the way trig stations were identified in the 0.25 m dmc aerial imagery. since the locations were estimated from the trig station metadata in the 0.25 m imagery, locating these landmarks was a challenging process that could not be automated. the difficulty of detecting reference points in the orthomosaics could be mainly attributed to the appearance of the triangulation stations, which range from windmill towers to platforms, vary in height, and can be with or without beacons. while locating tall trig stations with large poles that cast visible shadows was fairly simple, the identification of smaller stations was challenging and subject to a certain amount of subjectiveness. however, these circumstances were kept to a minimum during the selection of gcps, which were mostly visible in the dmc imagery, which is why only 58 of the 89 available trig stations were used for the validation. conclusion with the data sets created and published through this study, we present the first wall-to-wall spatial vhr elevation models and orthomosaics derived from aerial imagery for the knp, south africa. we produced elevation data sets (dtm and dsm) that are publicly and freely available at three different pixel resolutions (0.25 m, 1 m and 5 m). orthomosaics (cir) were generated at the original cell size of 0.25 m. these data sets can be used as reference information for a broad variety of savanna ecosystem analyses, including vegetation structure analysis (vertical and horizontal), biomass estimation, hydrological modelling (including erosion and flood modelling), habitat suitability modelling, or as a reference data set for other elevation models of lower resolution. to further improve data quality, we will continuously review and investigate extraction and filtering algorithms for photogrammetric elevation data in the knp. our goal is to derive an optimal top-of-canopy model, especially in open savanna woodlands. for future validation work, more extensive elevation data sets would be beneficial to further assess the accuracy of the models in different contexts and to improve the matching precision in the sgm algorithm. in order to make the products presented in this study even more statistically comparable to existing height derivatives, greater quantities of better stratified in situ data sets need to be utilised. this would allow for more standardised validation strategies, such as those proposed in the american society for photogrammetry and remote sensing (asprs) positional accuracy standards for digital geospatial data (asprs 2015). to this point, collaborative efforts in conjunction with the greater savanna science community can assist in evaluating the existing products and further refining future products. acknowledgements the authors would like to thank geospace intl. for the personal provision of digital mapping camera (dmc) data and thus enabling an early start of this project. the authors also acknowledge the support of the chief directorate: national geo-spatial information (cdngi) in providing the validation data for the orthomosaics. thanks also to the information technology (it) support team of catalyst led by guillaume morin, who supported us during the setup of our high-performance cluster-based processing workflow. we would also like to express our thankfulness towards catalyst for financially supporting the presentation of results of this study at international conferences. furthermore, this study was supported by the spaces ii funded projects ecosystem management support for climate change in southern africa (emsafrica) and south africa land degradation monitor (saldi). competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions processing and evaluation of the presented products as well as the preparation of the manuscript were carried out by k.h. and m.u.. major technical support to derive the final datasets provided by j.s.b., j.b., p.b., a.d., e.h. and j.z. supported the validation strategy that was accomplished. w.l. and i.s. provided necessary methodological input for the study. b.j. provided the initial aerial imagery. m.n.-r. initiated the data delivery process. w.l. and c.s. contributed to the conceptualisation of the study. all co-authors contributed towards and/or commented on the manuscript. ethical considerations this article followed all ethical standards for research without direct contact with human or animal subjects. funding information this research received no specific grant from any funding agency in the public, commercial or not-for-profit sectors. data availability the data sets derived in this study are freely accessible through the centre for environmental data analysis (ceda), and are available for download via the following link: http://dx.doi.org/10.5285/deab4235f1ef4cd79b73d0cbf2655bd7. updates of these products will be continuously added and accordingly announced on the platform. users of the data products should reference this article as source of the data. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the author. references american society for photogrammetry and remote sensing (asprs), 2015, ‘asprs positional accuracy standards for digital geospatial data’, photogrammetric engineering & remote sensing 81(3), 1–26. https://doi.org/10.14358/pers.81.3.a1-a26 asner, g.p. & levick, s.r., 2012, ‘landscape-scale effects of herbivores on treefall in african savannas’, ecology letters 15(11), 1211–1217. https://doi.org/10.1111/j.1461-0248.2012.01842.x axelsson, p., 2000, ‘dem generation from laser scanner data using adaptive tin models’, international archives of the photogrammetry, remote sensing and spatial information sciences – isprs archives 33, 110–117. baade, j. & schmullius, c., 2016, ‘tandem-x idem precision and accuracy assessment based on a large assembly of differential gnss measurements in kruger national park, south africa’, isprs journal of photogrammetry and remote sensing 119, 496–508. https://doi.org/10.1016/j.isprsjprs.2016.05.005 baltsavias, e.p., 1999, ‘a comparison between photogrammetry and laser scanning’, isprs journal of photogrammetry and remote sensing 54(2–3), 83–94. https://doi.org/10.1016/s0924-2716(99)00014-3 biggs, r., biggs, h.c., dunne, t.t., govender, n. & potgieter, a.l.f., 2003, ‘experimental burn plot trial in the kruger national park: history, experimental design and suggestions for data analysis’, koedoe 46(1), 35. https://doi.org/10.4102/koedoe.v46i1.35 brigham, e.o., 1988, the fast fourier transform and its applications, prentice hall, hoboken, nj. catalyst pci geomatics brand, 2020, software: catalyst enterprise, viewed 01 february 2021, from https://catalyst.earth/products/catalyst-enterprise/. chown, s., 2010, ‘temporal biodiversity change in transformed landscapes: a southern-africa perspective’, philosophical transactions of the royal society 365(1558), 3729–3742. https://doi.org/10.1098/rstb.2010.0274 clarke, p.j., know, k.j.e., wills, k.e. & campbell, m., 2005, ‘landscape patterns of woody plant response to crown fire: disturbance and productivity influence sprouting ability’, journal of ecology 93(3), 544–555. https://doi.org/10.1111/j.1365-2745.2005.00971.x florinsky, i.v., 2012, digital terrain analysis in soil science and geology, academic press, oxford. gdal/ogr contributors, 2021, ‘gdal/ogr geospatial data abstraction software library’, open source geospatial foundation, viewed 21 january 2021, from https://gdal.org. gertenbach, w.p.d., 1980, ‘rainfall patterns in the kruger national park’, koedoe 23(1), 35–43. https://doi.org/10.4102/koedoe.v23i1.634 greenwalt, c.r. & shultz, m.e., 1968, principles of error theory and cartographic applications, aeronautical chart & information center technical report no, 96, st. louis, missouri, united states of america, viewed n.d., from https://www.fgdc.gov/standards/projects/accuracy/part3/tr96 haala, n., hastedt, h., wolf, k., ressl, c. & baltrusch, s., 2010, ‘digital photogrammetric camera evaluation – generation of digital elevation models’, photogrammetrie – fernerkundung – geoinformation jahrgang 2, 99–115. https://doi.org/10.1127/1432-8364/2010/0043 hirschmüller, h., 2007, ‘stereo processing by semi-global matching and mutual information’, ieee transactions on pattern analysis and machine intelligence 30(2), 328–341. https://doi.org/10.1109/tpami.2007.1166 hirt, c., filmer, m.s. & featherstone, w.e., 2010, ‘comparison and validation of recent freely-available aster-gdem ver 1, srtm ver4.1 and geodata dem-9s ver3 digital elevation models over australia’, australian journal of earth sciences 57(3), 337–347, https://doi.org/10.1080/08120091003677553 hoekman, d.h. & varekamp, c., 2001, ‘observation of tropical rain forest trees by airborne high-resolution interferometric radar’, ieee transactions on geoscience and remote sensing 39(3), 584–594. https://doi.org/10.1109/36.911116 ichim, l. & popescu, d., 2019, ‘flooded areas evaluation from aerial images based on convolutional neural network’, in international geoscience and remote sensing symposium (igarss), yokohama, japan, july 28 – august 02, 2019. jensen, j.r., 2007, remote sensing of the environment: an earth resource perspective, prentice hall, hoboken, nj. klette, r., kruger, n., vaudrey, n.t., pauwels, k., van hulle, m., morales, s. et al., 2011, ‘performance of correspondence algorithms in vision-based driver assistance using an online image sequence database’, ieee transactions on vehicular technology 60, 2012–2026. https://doi.org/10.1109/tvt.2011.2148134 le bris, a., tassin, f. & chehata, n., 2013, ‘contribution of texture and red-edge band for vegetated areas detection and identification’, in international geoscience and remote sensing symposium (igarss), melbourne, australia, july 21–26, 2013. lim, k., treitz, p., wulder, m., st. onge, b. & flood, m., 2003, ‘lidar remote sensing of forest structure’, progress in physical geography: earth and environment 27(1), 88–106. https://doi.org/10.1191/0309133303pp360ra lindberg, e. & holmgren, j., 2017, ‘individual tree crown methods for 3d data from remote sensing’, current forestry reports 3, 19–31. https://doi.org/10.1007/s40725-017-0051-6 lucas, r.m., milne, a.k., mitchell, a., donnelly, b. & ellison, j., 2000, ‘use of stereo aerial photography for assessing changes in the extent and height of mangrove canopies in tropical australia’, in international geoscience and remote sensing symposium (igarss), honolulu, july 24–28, 2000. meng, s., pang, y. & zhang, z., 2014, ‘forest biomass estimation using fourier-based textural ordination of high resolution airborne optical image’, in third international workshop on earth observation and remote sensing applications (eorsa), changsha, china, june 11–14, 2014. michael, m., salmen, j., stallkamp, j. & schlipsing, m., 2013, ‘real-time stereo vision: optimizing semi-global matching’, in ieee intelligent vehicles symposium, gold coast, australia, june 23–26, 2013. miller, c.l. & laflamme, r.a., 1958, the digital terrain model – theory & application, mit photogrammetry laboratory, cambridge, ma. mograbi, p.j., asner, g.p., witkowski, e.t.f., erasmus, b.f.n., wessels, k.j., mathieu, r. et al., 2017, ‘humans and elephants as treefall drivers in african savannas’, ecography 40, 1274–1284. https://doi.org/10.1111/ecog.02549 nelson, a., reuter, h.i. & gessler, p., 2009, ‘dem production methods and sources’, in t. hengl & h. reuter (eds.), geomorphology, concepts, software, applications: developments in soil science, p. 33, elsevier, oxford. o’loughlin, f.e., paiva, r.c.d., durand, m., alsdorf, d.e. & bates, p.d., 2016 ‘a multi-sensor approach towards a global vegetation corrected srtm dem product’, remote sensing of environment 182, 49–59. https://doi.org/10.1016/j.rse.2016.04.018 pieczonka, t., bolch, t. & buchroithner, m.f., 2011, ‘generation and evaluation of multitemporal digital terrain models of the mt. everest area from difference optical sensors’, isprs journal of photogrammetry and remote sensing 66(6), 927–940. https://doi.org/10.1016/j.isprsjprs.2011.07.003 pulighe, g. & fava, f., 2013, ‘dem extraction from archive aerial photos: accuracy assessment in areas of complex topography’, european journal of remote sensing 46(1), 363–378. https://doi.org/10.5721/eujrs20134621 reitberger, j., krzystek, p. & stilla, u., 2008, ‘analysis of full waveform lidar data for the classification of deciduous and coniferous trees’, international journal of remote sensing 29(5), 1407–1431. https://doi.org/10.1080/01431160701736448 shin, x., cao, l., liu, k., she, g. & ruan, h., 2016, ‘forest biomass estimation using fourier-based textural ordination of high resolution airborne optical image’, in fourth international workshop on earth observation and remote sensing applications (eorsa), guangzhou, china, july 04–06, 2016. smit, i.p.j., gregory, p.a., govender, n., kennedy-bowdoin, t., knapp, d.e. & jacobson, j., 2010, ‘effects of fire on woody vegetation structure in african savanna’, ecological applications 20(7), 1865–1875. https://doi.org/10.1890/09-0929.1 stepper, c., straub, c. & pretzsch, h., 2015, ‘assessing height changes in a highly structured forest using regularly acquired aerial image data’, forestry 88(3), 304–316. https://doi.org/10.1093/forestry/cpu050 straub, c., stepper, c., seitz, r., & waser, l.t., 2013, ‘potential of ultracamx stereo images for estimating timber volume and basal area at the plot level in mixed european forests’, canadian journal of forest research 43(8), 731–741. https://doi.org/10.1139/cjfr-2013-0125 tian, j., schneider, t., straub, c., kugler, f. & reinartz, p., 2017, ‘exploring digital surface models from nine different sensors for forest monitoring and change detection’, remote sensing 9(287), 1–26. https://doi.org/10.3390/rs9030287 venter, f.j., 1990, ‘a classification of land for management planning in the kruger national park’, phd thesis, university of south africa, pretoria. venter, k.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation pattern’, in j. du toit, h. biggs & k.h. rogers (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83–129, island press, london. white, j.c., wulder, m.a., vastaranta, m., coops, n.c., pitt, d. & woods, m., 2013, ‘the utility of image-based point clouds for forest inventory: a comparison with airborne laser scanning’, forests 4(3), 518–536. https://doi.org/10.3390/f4030518 wu, m., shi, p., chen, a., shen, c. & wang, p., 2017, ‘impacts of dem resolution and area threshold value uncertainty on the drainage network derived using swat’, water sa 43(3), 450–463. https://doi.org/10.4314/wsa.v43i3.10 zhu, r., yu, d., ji, s. & lu, m., 2019, ‘matching rgb and infrared remote sensing images with densely-connected convolutional neural networks’, remote sensing 11(23), 2836. https://doi.org/10.3390/rs11232836 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 filelist convert a pdf file! filelist convert a pdf file! page 1 page 2 abstract introduction research method and design results and discussion conclusion acknowledgements references appendix 1 about the author(s) anna s. dippenaar-schoeman biosystematics: arachnology, arc – plant health and protection, queenswood, south africa department of zoology and entomology, university of pretoria, pretoria, south africa linda wiese south african national survey of arachnida (sansa) team eastern cape, jeffrey’s bay, south africa stefan h. foord department of zoology, university of venda, thohoyandou, south africa charles r. haddad department of zoology and entomology, university of the free state, bloemfontein, south africa citation dippenaar-schoeman, a.s., wiese, l., foord, s.h. & haddad, c.r., 2020, ‘a list of spider species found in the addo elephant national park, eastern cape province, south africa’, koedoe 62(1), a1578. https://doi.org/10.4102/koedoe.v62i1.1578 checklist a list of spider species found in the addo elephant national park, eastern cape province, south africa anna s. dippenaar-schoeman, linda wiese, stefan h. foord, charles r. haddad received: 11 july 2019; accepted: 17 feb. 2020; published: 02 apr. 2020 copyright: © 2020. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the knowledge of spiders in the eastern cape province lags behind that of most other south african provinces. the eastern cape province is renowned for its conservation areas, as the largest part of the albany centre of endemism falls within this province. this article provides a checklist for the spider fauna of the addo elephant national park, one of the most prominent conservation areas of the eastern cape, to detail the species found in the park and determine their conservation status and level of endemicity based on their known distribution. various collecting methods were used to sample spiders between 1974 and 2016. forty-seven families that include 184 genera and 276 species were recorded. thomisidae (39 spp.), araneidae (39 spp.), salticidae (35 spp.) and theridiidae (25 spp.) were the most species-rich families, while 14 families were only represented by a single species. conservation implications: a total of 12.7% of the south african spider fauna and 32.9% of the eastern cape fauna are protected in the park; 26.4% are south african endemics, and of these, 3.6% are eastern cape endemics. approximately, 4% of the species are possibly new to science, and 240 species are recorded from the park for the first time. keywords: arachnida; conservation; endemism; biome; records. introduction one of the core research areas of the south african national survey of arachnida (sansa) is to determine the number of arachnid species presently conserved in protected areas, including the south african national parks (sanparks) (dippenaar-schoeman & haddad 2006; dippenaar-schoeman 2016). the species distribution data generated through sanparks and other surveys feeds into the conservation assessments used to compile the red data list of the arachnida of south africa (lyle & dippenaar-schoeman 2015). compared to some of the better surveyed provinces of south africa, for example, kwazulu-natal and limpopo, the eastern cape has received far less attention, but harbours relatively more endemic taxa than the limpopo province, particularly as a function of its size (foord et al. 2011). this serves as a possible indicator of the unique spider assemblages of the region. although the addo elephant national park (aenp) is the largest national park in the eastern cape and the third largest national park in south africa, no documented surveys of spiders exist for this protected area. work in the province has largely been limited to the mountain zebra national park, the first national park for which a spider species list was published (dippenaar-schoeman 1988, 2006), and a study on the subsocial eresid stegodyphus tentoriicola purcell, 1904 (ruch et al. 2009a, 2009b, 2012). further work in the eastern cape includes aspects of the spider fauna in the mkambati nature reserve (dippenaar-schoeman, hamer & haddad 2011; hamer & slotow 2017) and the silaka nature reserve (forbanka & niba 2013), while surveys in the asante sana nature reserve were the focus of a doctoral study on epigaeic invertebrates (midgley 2012). the only records of spiders from the aenp (30 spp.) were included in the first atlas of south african spiders (dippenaar-schoeman et al. 2010), and six species have been referred to in taxonomic papers in the past (dippenaar-schoeman 1980; jocqué 1984; lyle & haddad 2018; wesołowska & haddad 2013, 2018). here, we document all spider species recorded from the aenp over the last 50 years, with efforts being concentrated over the last decade. information is provided on the global distribution, endemicity and conservation status of each species. research method and design study area and period the aenp is situated in the sundays river region, between the city of port elizabeth and the town of makhanda (grahamstown) (figure 1). although originally proclaimed in 1931 to protect one of the four elephant populations that survived the start of the 20th century in south africa, it is very significant in the conservation of the albany thicket, which is predominantly found in the eastern cape (johnson, cowling & phillipson 1999) and is the dominant biome of the region (mucina & rutherford 2006). the woody cape nature reserve and some portions of privately owned land were recently added to the park, bringing its total area to approximately 179 000 hectares (sanparks 2015). figure 1: map of south africa, showing the locality of the addo elephant national park in the eastern cape province (inset) and the different sites where spiders were collected. rainfall in the park is mainly concentrated in spring and autumn, varying between 350 mm and 900 mm per annum, depending on topography and biomes (sanparks 2015). the region seldom experiences temperatures below 0 °c, while summer extremes of 45 °c are quite common (lombard et al. 2001). sampling methods and identification the first spider survey in the aenp took place in 1974, when the members of the agricultural research council (arc) – plant protection research institute (ppri) (now arc – plant health and protection) sampled there. in 2009, a second survey was undertaken by the members of the arc-ppri. between these events, there was occasional collecting by various museum researchers and members of the public who deposited specimens in several museums in south africa and internationally. in 2010, more regular sampling started in the park after a formal research agreement was established between sansa and sanparks, to stimulate more intensive spider sampling in the national parks. as a result, more regular sampling was undertaken in five different parts of the park (figure 2), which included the semi-arid sundays river succulent thicket around the darlington dam; the fynbos and afromontane forests of the zuurberg mountains; the subtropical thickets in the kabouga area just north of kirkwood; the alexandria coastal forest and dune thickets of the woody cape area; and the albany thickets of the main camp in the sundays river valley (figure 2). figure 2: habitat types in the addo elephant national park: (a) darlington dam (succulent karoo thicket); (b) main camp (albany thickets); (c) and (d) kabouga (subtropical thicket); (e) and (f) woody cape (coastal forest and dune thicket); (g) and (h) zuurberg (fynbos and afromontane forest). conventional sampling techniques, as described by dippenaar-schoeman et al. (2015), were used. this included sweepnet sampling, beating, leaf-litter sifting and active searches during the day and night. sampling did not follow a standardised protocol but was undertaken sporadically and ad hoc, depending on locality and time constraints. voucher specimens were predominantly deposited in the national collection of arachnida (nca) at the arc–plant health and protection in pretoria. taxonomic constraints, juveniles and undescribed species necessitated the use of morphospecies in some instances (appendix 1). endemicity value and conservation status a detailed explanation of the indices to assess the endemicity value and conservation status of each species can be found at the end of appendix 1, but it distinguishes between species only known from the type locality (6), the eastern cape (5), two adjoining provinces (4), south africa (3), southern africa (2), afrotropical region (1) and finally areas outside the afrotropical region (0). the conservation status was derived from a recent national red list assessment of spiders in south africa. ethical considerations this research followed all ethical standards for research without direct contact with human or animal subjects. results and discussion numbers present in total, 276 species in 184 genera and 47 families were recorded (table 1; appendix 1). seven species are probably new and 16 species are undetermined, because of the lack of resolved taxonomy of their genera (appendix 1). possible new species belong to large families that require comprehensive revision to determine the taxonomic status and identity of these taxa. table 1: spider diversity of the addo elephant national park, with the total number of families, genera and species sampled. family diversity of the 47 spider families collected from aenp (table 1; appendix 1), the thomisidae and araneidae with 39 spp. are the most species-rich, followed by the salticidae (35 spp.) and theridiidae (25 spp.). for comparison, at the mkambati nature reserve, the theridiidae and araneidae were the most species-rich families (dippenaar-schoeman et al. 2011), and at the mountain zebra national park, it was the thomisidae (dippenaar-schoeman 1988, 2006). thomisidae: thomisids are mainly plant dwellers and are easily dispersed by wind. consequently, most species have a wide distribution. only five species were previously reported from aenp in the spider atlas (dippenaar-schoeman et al. 2010); therefore, 34 spp. are newly recorded from the park. twenty-one species have a wide distribution throughout africa, while four species are also known from europe and nine occur throughout southern africa. four species are south african endemics, with two – holopelus almiae dippenaar-schoeman, 1986 and mystaria lindaicapensis lewis & dippenaar-schoeman, 2014 (figure 3l) – being near endemics of the eastern cape. both species are also known from the western cape. the species listed as monaeses sp. 2 is new to science, and the same species was also recorded at the mkambati nature reserve (dippenaar-schoeman et al. 2011). diaea dorsata (fabricius, 1777) (figure 3k) is known from europe to japan (world spider catalog 2019) and is reported from south africa for the first time. figure 3: selected spiders of the addo elephant national park: (a) araneus detrimentosus (o. pickard-cambridge, 1889) (araneidae); (b) undescribed cyclosa sp. (araneidae); (c) ideocaira transversa simon, 1903 (araneidae); (d) zygiella sp. (araneidae); (e) ancylotrypa cornuta purcell, 1904 (cyrtaucheniidae); (f) allothele australis (purcell, 1903) (dipluridae); (g) moggridgea rupicoloides hewitt, 1914 (migidae); (h) hermacha grahami (hewitt, 1915) (nemesiidae); (i) new genus (philodromidae); (j) harpactirella magna purcell, 1903 (theraphosidae); (k) diaea dorsata (fabricius, 1777) (thomisidae); (l) mystaria lindaicapensis lewis & dippenaar-schoeman, 2014 (thomisidae). araneidae: the araneidae are web builders and produce typical orb webs. of the 39 species recorded, only four were previously known from the park (dippenaar-schoeman et al. 2010). twenty of the species have a wide african distribution, three species are endemic to southern africa, and only three are south african endemics that include ideocaira transversa simon, 1903 (figure 3c), nemoscolus elongatus lawrence, 1947 and ursa turbinata simon, 1895. three genera, acanthepeira, larinioides and zygiella (figure 3d), represented by undetermined species, are newly recorded for south africa, as well as the species araneus detrimentosus (o. pickard-cambridge, 1889) (figure 3a) and chorizopesoides orientalis (simon, 1909). species of six genera could not be determined (e.g. figure 3b) and might represent new records. salticidae: the salticidae are free-living spiders found on vegetation and the soil surface. only cyrba nigrimana simon, 1900 (dippenaar-schoeman et al. 2010), heliophanus (heliophanus) gramineus wesołowska & haddad, 2013, massagris mirifica peckham & peckham, 1903 (wesołowska & haddad 2018) and pseudicius maculatus haddad & wesołowska, 2011 (wesołowska & haddad 2013) were previously recorded from the park. eighteen species are widely known throughout africa, and one, menemerus bivittatus (dufour, 1831), has a pantropical distribution. six species are endemic to southern africa, and 10 species are south african endemics; five species are near endemic to the eastern cape. only one tanzania species is possibly new to science. theridiidae: the theridiids construct different types of webs known as cobwebs or gumfoot webs. no theridiids were previously reported from the park. unfortunately, the taxonomy of most theridiid genera in africa is still unresolved. seven species (28%) could not be identified to species level. this corresponds with the mkambati nature reserve where 67% of the species were unresolved (dippenaar-schoeman et al. 2011). eight species have a wide global distribution and five species are south african endemics, with only one species (phoroncidia capensis [simon, 1895]) being endemic to the eastern cape. some of the undetermined species might be new to science. species endemicity and conservation status of the 276 species sampled, 17 spp. (6.2%) are data deficient (dd) and lack taxonomic or distribution data, while 31 spp. (11.2%) were not evaluated (ne) (table 2; appendix 1). the majority of the species (227 spp., 82.2%) sampled are listed as least concern (lc). ninety-seven of the species (35.1%) are found throughout africa, while 50 species (18.1%) are endemic to southern africa. one species, mystaria lindaicapensis, is considered to be vulnerable, because of its restricted distribution in the eastern and western cape provinces (table 3). table 2: conservation status and endemicity of the spider species sampled at the addo elephant national park. table 3: species of special concern recorded from the addo elephant national park. of the species identified to species level, only 73 spp. (26.4%) are south african endemics (table 2; appendix 1). there are presently no endemic species known only from the aenp, but 10 species from eight families are eastern cape endemics (ece) and a further species is near endemic. these species are of special concern because of their small distribution ranges (table 3), and need more sampling and/or taxonomic study to improve knowledge of their distributions. the 10 ece are listed as data deficient, of which only ancylotrypa cornuta purcell, 1904 (figure 3e) is known from both sexes. the vu near-endemic species, m. lindaicapensis, is only known from three south coast localities, including aenp. the ece species include three trapdoor spider species (a. cornuta, figure 3e; hermacha grahami [hewitt, 1915], figure 2e; moggridgea rupicoloides hewitt, 1914, figure 3g) and two baboon spider species (harpactira curvipes pocock, 1897; harpactirella magna purcell, 1903, figure 3j). twenty-five of the aenp spiders have a wide global distribution (table 2; appendix 1), and most of them have been recorded from several continents (world spider catalog 2019). some of these species listed from aenp have not been recorded from south africa before, including araneus detrimentosus, chorizopesoides orientalis and diaea dorsata. conclusion a total of 240 species have been newly recorded from the aenp, bringing the total number of species known from the park to 276. all these new data have been made available for the spider red listing project. the aenp spider fauna represents 12.7% of south african species and 32.9% of eastern cape species, of which 10 species are ece that are of special concern and need additional collecting. acknowledgements this survey forms part of the south african national survey of arachnida (sansa) inventories of the south african national parks. the agricultural research council (arc) and the south african national biodiversity institute (sanbi) threatened species programme are thanked for generously funding sansa phase ii. the authors would like to thank the park manager and officials of the addo elephant national park for their friendliness and assistance, and all the people who sampled in the park and donated their specimens. the authors are grateful to dr w. wiese, husband of the second author, who assisted with the fieldwork. the authors also received financial support from the university of pretoria. the authors also thank the staff of the arachnology unit of the biosystematics programme, arc – plant health and protection, for their assistance with processing and databasing the material collected. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions all the authors are team members of sansa and contributed towards planning this national survey. l.w. participated in fieldwork, while the rest of the authors assisted in the identification of specimens and the preparation and editing of the manuscript. funding information a.s.d.-s. s.h.f. and c.r.h. are grateful for the financial support from the national research foundation of south africa. data availability statement all specimens sampled in this study were deposited in the national collection of arachnida (nca) at the arc – plant health and protection in pretoria, south africa. the specimen data are available from the nca and sansa databases on request. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references benoit, p.l.g., 1968, ‘les selenopidae africains au nord du 17e parallèle sud et reclassement des espèces africaines de la famille (araneae)’, revue de zoologie et de botanique africaines 77(1–2), 113–141. dippenaar-schoeman, a.s., 1980, ‘the crab-spiders of southern africa; araneae: thomisidae). 1. the genus runcinia simon, 1875’, journal of the entomological society of south africa 43(2), 303–326. dippenaar-schoeman, a.s., 1988, ‘annotated check list of the spiders (araneae) of the mountain zebra national park’, koedoe 31(1), 151–160. https://doi.org/10.4102/koedoe.v31i1.492 dippenaar-schoeman, a.s., 2006, ‘new records of 43 spider species from the mountain zebra national park, south africa (arachnida: araneae)’, koedoe 49(2), 23–28. https://doi.org/10.4102/koedoe.v49i2.113 dippenaar-schoeman, a.s., 2016, ‘sansa: protected areas: national parks’, sansa news 25, 5–6. dippenaar-schoeman, a.s. & haddad, c.r., 2006, ‘what is the south african national survey of arachnida (sansa) all about?’, sansa news 1, 1–3. dippenaar-schoeman, a.s., haddad, c.r., foord, s.h., lyle, r., lotz, l.n. & marais, p., 2015, ‘south african national survey of arachnida (sansa): review of current knowledge, constraints and future needs for documenting spider diversity (arachnida: araneae)’, transactions of the royal society of south africa 70(3), 245–277. https://doi.org/10.1080/0035919x.2015.1088486 dippenaar-schoeman, a.s., haddad, c.r., foord, s.h., lyle, r., helberg, l. & mathebula, s. et al., 2010, first atlas of the spiders of south africa (arachnida: araneae), arc – plant protection research institute, pretoria. dippenaar-schoeman, a.s., hamer, m. & haddad, c.r., 2011, ‘spiders (arachnida: araneae) of the vegetation layer of the mkambati nature reserve, eastern cape, south africa’, koedoe 53(1), 1058. https://doi.org/10.4102/koedoe.v53i1.1058 foord, s.h., dippenaar-schoeman, a.s. & haddad, c.r., 2011, ‘the faunistic diversity of spiders (arachnida, araneae) of the savanna biome in south africa’, transactions of the royal society of south africa 66(3), 170–201. https://doi.org/10.1080/0035919x.2011.639406 forbanka, d.n. & niba, a.s., 2013, ‘distribution and diversity of epigaiec invertebrate assemblages in silaka nature reserve, eastern cape, south africa’, african entomology 21(1), 58–69. https://doi.org/10.4001/003.021.0111 griswold, c.e., 1987, ‘the african members of the trap-door spider family migidae (araneae: mygalomorphae) 1: the genus moggridgea o.p.-cambridge, 1875’, annals of the natal museum 28(1), 1–118. hamer, m.l. & slotow, r., 2017, ‘a conservation assessment of the terrestrial invertebrate fauna of mkambati nature reserve in the pondoland centre of endemism’, koedoe 59(1), a1428. https://doi.org/10.4102/koedoe.v59i1.1428 huber, b.a., 2003, ‘southern african pholcid spiders: revision and cladistic analysis of quamtana gen. nov. and spermophora hentz (araneae: pholcidae), with notes on male-female covariation’, zoological journal of the linnean society 139(4), 477–527. https://doi.org/10.1046/j.0024-4082.2003.00082.x jocqué, r., 1984, ‘linyphiidae (araneae) from south africa. part i: the collection of the plant protection research institute, pretoria’, journal of the entomological society of south africa 47(1), 121–146. johnson, c.f., cowling, r.m. & phillipson, p.b., 1999, ‘the flora of the addo elephant national park, south africa: are threatened species vulnerable to elephant damage?’, biodiversity & conservation 8(11), 1447–1456. https://doi.org/10.1023/a:1008980120379 lewis, a.s.h. & dippenaar-schoeman, a.s., 2014, ‘revision of the spider genus mystaria simon, 1895 (araneae: thomisidae) and the description of a new genus from the afrotropical region’, zootaxa 3873(2), 101–144. https://doi.org/10.11646/zootaxa.3873.2.1 lombard, a.t., johnson, c.f., cowling, r.m. & pressey, r.l., 2001, ‘protecting plants from elephants: botanical reserve scenarios within the addo elephant national park, south africa’, biological conservation 102(2), 191–203. https://doi.org/10.1016/s0006-3207(01)00056-8 lyle, r. & dippenaar-schoeman, a.s., 2015, ‘red listing of south african spiders’, sansa newsletter 23, 1. lyle, r. & haddad, c.r., 2018, ‘jocquestus, a new genus of trachelid sac spiders from the afrotropical region (arachnida: araneae)’, zootaxa 4471(2), 309–333. https://doi.org/10.11646/zootaxa.4471.2.4 midgley, j.m., 2012, ‘quantifying the structure of epigaeic invertebrate communities along an altitudinal gradient on the great escarpment, south africa’, unpublished phd thesis, rhodes university, grahamstown. mucina, l. & rutherford, m.c., 2006, the vegetation of south africa, lesotho and swaziland, strelitzia 19, south african national biodiversity institute, pretoria. ruch, j., heinrich, l., bilde, t. & schneider, j.m., 2009a, ‘relatedness facilitates cooperation in the subsocial spider, stegodyphus tentoriicola’, bmc evolutionary biology 9(257), 1–7. https://doi.org/10.1186/1471-2148-9-257 ruch, j., heinrich, l., bilde, t. & schneider, j.m., 2009b, ‘the evolution of social inbreeding mating systems in spiders: limited male mating dispersal and lack of pre-copulatory inbreeding avoidance in a subsocial predecessor’, biological journal of the linnean society 98(4), 851–859. https://doi.org/10.1111/j.1095-8312.2009.01322.x ruch, j., heinrich, l., bilde, t. & schneider, j.m., 2012, ‘site selection and foraging in the eresid spider stegodyphus tentoriicola’, journal of insect behaviour 25(1), 1–11. https://doi.org/10.1007/s10905-011-9273-9 south african national parks (sanparks), 2015, addo elephant national park management plan for the period 2015–2025, south african national parks, pretoria. wesołowska, w. & haddad, c.r., 2013, ‘new data on the jumping spiders of south africa (araneae: salticidae)’, african invertebrates 54(1), 177–240. https://doi.org/10.5733/afin.054.0111 wesołowska, w. & haddad, c.r., 2018, ‘further additions to the jumping spider fauna of south africa (araneae: salticidae)’, annales zoologici, warszawa 68(4), 879–908. https://doi.org/10.3161/00034541anz2018.68.4.011 appendix 1 table 1-a1: spiders of the addo elephant national park, listing their endemicity, conservation status, distribution and areas sampled in the park. table 1-a1 (continues): spiders of the addo elephant national park, listing their endemicity, conservation status, distribution and areas sampled in the park. table 1-a1 (continues): spiders of the addo elephant national park, listing their endemicity, conservation status, distribution and areas sampled in the park. table 1-a1 (continues): spiders of the addo elephant national park, listing their endemicity, conservation status, distribution and areas sampled in the park. table 1-a1 (continues): spiders of the addo elephant national park, listing their endemicity, conservation status, distribution and areas sampled in the park. table 1-a1 (continues): spiders of the addo elephant national park, listing their endemicity, conservation status, distribution and areas sampled in the park. filelist convert a pdf file! page 1 page 2 page 3 158--26 oct 2009.indd [final version].indd a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe original research a rticle #158 (page number not for citation purposes) 26 the socio-economic impact of the karoo national park authors: melville saayman1 andrea saayman1 madelien ferreira1 affi liations: 1institute for tourism and leisure studies, school of economics, north west university, south africa correspondence to: melville saayman e-mail: melville.saayman@nwu. ac.za postal address: private bag x6001, potchefstroom, 2520 keywords: sanparks; south africa; tourism; economic value; community perceptions dates: received: 10 aug. 2008 accepted: 11 nov. 2008 published: 05 may 2009 how to cite this article: saayman, m., saayman, a. & ferreira, m., 2009, ‘the socio-economic impact of the karoo national park’, koedoe 51(1), art. #158, 10 pages. doi: 10.4102/ koedoe.v51i1.158 this article is available at: http://www.koedoe.co.za © 2009. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. vol. 51 no. 1 page 1 of 10 introduction national parks in south africa have three fundamental spheres: fi rstly, to conserve the biodiversity of the country; secondly, to maintain a relationship of community upliftment and capacity building with people living in the areas in and around the parks; and, lastly, to provide a recreational outlet for people to experience and enjoy the wonders of the parks (saayman & saayman 2006a). while the main emphasis of national parks has traditionally been on conservation, there has been a shift towards economic sustainability and the upliftment of local communities (myburgh & saayman 2002). it is the latter that provides the rationale for this research and, although south africa has some of the greatest protected areas in the world, little is known about the socio-economic impact of these parks. socio-economics is the study of the relationship between economic activity and social life (van der merwe 2008). the goal of a socio-economic study is generally to assess socio-economic development, usually in terms of improvements in metrics such as gross domestic product (gdp), life expectancy, literacy and levels of employment (van der merwe 2008), with the aim of enhancing the benefi ts received by the society. a socio-economic impact study therefore goes beyond assessing income generated by the park, and also involves the contribution of the park to the quality of life of the community (saayman & saayman 2006a). eadington and redman (1991, cited by walpole & goodwin 2000) state that tourism, as a development tool, holds many potential economic benefi ts for host communities, including increased employment opportunities, improved socio-economic conditions and greater market stability than are provided by traditional commodity exports. the magnitude of the economic benefi ts is dependent on the amount that remains in the area. adding to this, saayman and saayman (2004) express their concern about leakages, both in the form of imported goods and imported skills, which will reduce the benefi t for the local community because most national parks are situated in rural areas. page (2007) identifi ed four economic benefi ts of tourism, namely the generation of income for the local community; the creation of new employment opportunities; improvements to the structure and balance of economic activities in the locality; and the encouragement of entrepreneurial activity. the attitudes of residents towards tourism represent an important way in which participants contribute to policy and public support for, or dissent towards, tourism (page 2007). studies that were conducted on socio-economic and economic impacts include those of loader (1994) and pelser (2003), who argue that social involvement is an integral part of conservation in south africa. macleod (2001) and fauzi and buchary (2002) suggest that the alleviation of poverty and marginality in the case of residents of the kepulauan seribu marine park, del este in the dominican republic, should receive priority, and that park management should be based on consensus building and participation by all stakeholders. therefore, the community should play an important role in maximising the local socio-economic benefi ts (govender et al. 2005). an example of how the local community has focused efforts on maximising benefi ts from tourism activity as the result of a national park can be found in mpumalanga province. there, towns adjacent to the kruger national park have started to focus on offering additional tourism products, which contribute to income and employment and thereby increase the impact of the park in the region (saayman & saayman 2006b). the karoo national park (karoo np) is located in the western cape province of south africa, on the outskirts of the town of beaufort west, and can be accessed via the n1 highway. the karoo np abstract national parks in south africa are seen as major tourism assets due to the wildlife and various activities for international and local visitors. little is known of the socio-economic contribution of these parks to their respective local economies. the purpose of this research was to determine the socio-economic impact of the karoo national park (karoo np) in south africa, especially the economic impact of the karoo np on the local economy, the impact of tourism business development in the karoo district, and how the park affects the community. three surveys were used to determine the socio-economic impact: a community survey, a business survey and a tourist survey. the results show that the park has an impact in terms of production, income generation and employment in the area, but this impact is not as signifi cant as that of other national parks in south africa. a small percentage (4%) of businesses in beaufort west owe their existence to the karoo np, but most rely on tourist spending. for the park to have a greater impact, it is imperative to increase accommodation capacity, offer more activities and promote activities and attractions in the region. conservation implication: the importance of this article lies in the economic value that conservation management generates as well as identifying the benefi ts that communities derive from the existence of a national park. it also supports the notion that conservation entails more than just conserving fauna and fl ora and highlights the interdependence of conservation, tourism and community participation. koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research saayman, saayman & ferreira a rt ic le # 15 8 (page number not for citation purposes) serves mainly as an overnight park for visitors and tourists en route from or to the western cape/gauteng province (figure 1). during the late 1950s, mr william quinton, a local farmer and keen birdwatcher, began campaigning to have a conservation area declared in the vicinity of beaufort west. in the 1970s, the south african national parks board put forward plans to establish a national park that would be representative of the karoo biome. after considering a number of potential suitable areas, it was decided that this new park would be established in the vicinity of beaufort west (south african national parks 2007). the town council of beaufort west donated 7 209 ha of vacant communal land northwest of the town to the south african national parks board (currently known as sanparks). this formed the nucleus of the karoo np, proclaimed in 1979. since then, additional land purchases have increased the karoo np to its present size of approximately 96 000 ha (south african national parks 2007). the aim of this research was to estimate the socio-economic contribution of the karoo np to the surrounding communities, with an additional focus on the community’s involvement in the karoo np, guided by the following questions: what is the economic impact of the karoo np on the local • economy? what percentage of business development in the region is • as a result of the park? how many people are employed as a result of tourism • activity due to the karoo np? how satisfi ed is the community with regard to the karoo • np? methods saayman & saayman (2006a) created a framework to assess the socio-economic impact of national parks in south africa (see figure 2). they identifi ed fi ve issues of importance when assessing the socio-economic impacts of parks. firstly, there must be an understanding of the area and the main economic activities in the area – the spatial area must therefore be specifi ed and all the communities in the specifi ed area should be included in the research. secondly, the magnitude of visitor spending and the distribution of the spending throughout the local economy should be determined. this could be done via visitor (demand) and business (supply) surveys, especially in local areas, where business and spending information is not readily available. thirdly, and if possible, these should be complemented by a community survey. fourthly, the spending by the national park on employees and local products should be included to obtain a complete picture of spending activities resulting from the existence of the park. fifthly, the social impact of the park on the community must be determined. in areas where no local input-output model is available (which is usually the case), a proportional multiplier analysis could be used to determine the values of the income and sales multipliers, and to estimate the economic impact of the national park on income and employment. finally, the change in the nature of business and employment opportunities in the area due to the national park should be determined by comparing it to previous business surveys of the area (saayman & saayman 2006a). based on the above, three surveys were conducted: a community survey, a business survey in beaufort west and a visitor survey of tourists visiting the karoo np. secondly, a partial input-output model was developed to estimate partial multipliers and thus the economic impact of the karoo np. in general, multipliers indicate the magnitude of economic benefi ts in terms of sales, income and employment generated by the initial spending in the economy due to the tourism activity. input-output models are often used to generate these multipliers. yet regional inputoutput tables are often not available (also in south africa) and are expensive to develop because of the extensive data required. because of these constraints, this research employs partial multiplier analysis, and the partial multipliers are derived from a simplifi ed input-output table that was compiled from survey data – not regional estimates based on national input-output 27 figure 1 location of karoo national park (source: south african national parks 2007) vol. 51 no. 1 page 2 of 10 the socio-economic impact of the karoo national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #158 (page number not for citation purposes) 28vol. 51 no. 1 page 3 of 10 tables. thirdly, descriptive analysis was used to determine how the community feels about the karoo np and to determine its effect on the community of beaufort west. the surveys were completed between 27 march 2007 and 1 april 2007. seven postgraduate students were recruited to assist with the surveys. these fieldworkers distributed the questionnaires and collected the completed questionnaires later in the evening. microsoft excel was used to capture the data and data analyses were conducted using both excel and spss. the statistical techniques that were used include descriptive analyses. these descriptive analyses helped to provide preliminary insights into the nature of the responses obtained, as reflected in the distribution of the values for each variable of interest (tustin et al. 2005:341). the surveys survey one: community survey a structured questionnaire served as the instrument to collect the data, which contained both descriptive and causal questions. according to trochim (2006), descriptive research primarily describes what is occurring or what exists, whereas causal research investigates the effects of variables on one another. the questionnaire was based on the social impact measuring instrument developed by fredline et al. (2003). the questionnaire was adapted slightly to focus on the needs of the study. the main variables (residents’ perceptions of the impacts of the national park) were measured using a threeor four-point likert scale (fredline et al. 2003). questions were used to measure the independent variables and demographic information as well. these variables included participation, community attachment and demographics (fredline et al.2003). according to the 2006 census (statssa 2007) and the department of local government and housing (2006), there are approximately 5 000 households in a 30 km radius in beaufort west. stratified sampling of 300 households was undertaken, figure 2 framework to assess socio-economic impacts (from saayman & saayman 2006a) and 240 questionnaires were usable. for the purpose of the article, only the relevant descriptive data are used. survey two: business survey the questionnaire used for the business survey was similar to the one used by saayman and saayman (2006a) to determine the socio-economic impact of the addo elephant national park. research boundaries were determined for the survey, which were as follows: a 30 km radius around the karoo national park, which included the town of beaufort west. according to the municipal demarcation board, south africa (2006) there are a total of 757 businesses in beaufort west. most of the businesses that were surveyed were located in the town (see figure 1). convenience sampling was used for the business survey. in a convenience sample, the sample members are chosen on the basis of being readily accessible and willing to complete the questionnaire (tustin et al. 2005). a total of 86 questionnaires were received, which gives an 11% response rate. survey three: visitor survey the questionnaire that was used has been applied in various national park visitor surveys in south africa since 2001. in 2007, an availability sample was drawn from the visitors to the karoo np for the period (27 march 2007 to 1 april 2007). it must be noted that, for the purpose of this article, only overnight visitors formed part of the survey. a total of 104 questionnaires were received in this period, which is greater than the sample sizes of the previous years because the research team used a longer data collection period in 2007. in 2007, 31 469 people visited the karoo np. this visitor number includes both day visitors and overnight visitors. since the karoo np offers camping facilities and chalets, the overnight visitors were further divided into campers and visitors that spent the night in chalets. in terms of overnight visitors, 13 449 camping person nights and 9 373 chalet unit nights were sold. if it is taken into account that the average size of a visiting group original research saayman, saayman & ferreira koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 15 8 (page number not for citation purposes) to be made. the profit was calculated as a mark-up over stock and ranged from 20% (transport to, petroleum) to 50% (accommodation, curios). second, to trace the purchases made by businesses, each business had to list its main suppliers of stock, services and other operational expenses. since a further breakdown of the percentage of stock (for example) bought from a certain supplier was not given by the respondents, the assumption is made that the suppliers supply equal percentages. third, since consumption data for beaufort west is not available, the 2005 national input-output table consumption percentages are used to determine the income multipliers. care was taken to include the coefficients of the industries that are pertinent in beaufort west. lastly, the municipal allocations were made in accordance with the beaufort west integrated development plan for 2007 (nkungwana 2007). the visitor and business surveys provided the information for the economic impact assessment. yet, to get a clear picture of all spending as a result of the karoo np, sanparks provided the income and expense account for the karoo np for the year ending march 2007. according to this account, expenditure excluding depreciation amounted to approximately r5 million. a rough breakdown of these costs reveals that the highest spending item is personnel (56.6%), followed by operational costs (26.1%). to allocate these cost items to the various sectors in the model, some assumptions were again made, including that 30% of operational expenses are not incurred in the local area; maintenance cost is divided equally between retail, wholesale and manufacturing; all finance cost accrues to the local area; the remaining operational cost is divided between retail (40%), municipal (20%), transport (20%), wholesale (10%), food (5%) and manufacturing (5%); and 85.5% of personnel costs accrue to the local area (excluded are uif and other insurance). results the results are discussed in three sections. in the first section, a brief overview is given of the descriptive results of the surveys. the second section contains the results of the economic impact study and section three contains the results of the social impact study. descriptive results of the surveys the host community is mainly afrikaans speaking and tourists are predominantly english speaking (table 1). both the host community and the visitors are in their mid to late forties. the community is not that well educated relative to the visitors. both visitors and community members feel that the facilities at the karoo np meet their expectations. the visitors visit the karoo np for relaxation, whereas the community visits the karoo np to learn more about the animals and plants (wildlife, nature). the karoo np is also well known amongst various schools in beaufort west, especially the previously disadvantaged schools, for the opportunities the learners have to visit the park for educational reasons. four important pieces of information obtained from the visitor survey serve as input into the economic analysis. these are: number of tourists (number of tourist groups) the visitor survey revealed that visitors to the karoo np travel in groups of 3.14, with campers travelling in larger groups (3.37) than visitors staying in chalets (2.89). length of stay the average length of stay in the karoo np is 1.7 nights according to the visitor survey. campers again tend to stay a bit longer – 1.94 nights, while visitors in chalets stay on average 1.49 nights. this is in line with previous surveys and the karoo np is therefore often seen as a stop-over destination between cape town and gauteng. expenditure by tourists according to the visitor survey, camping groups spend on average r570.92 while staying in the karoo np (see table 2). to the karoo np was 3.14, the 104 questionnaires received represent 327 visitors to the park during the year. factors that have an influence on the number of questionnaires completed during the survey period include that the park has only 36 chalets and the bed occupancy varies between seasons, which makes it difficult to collect large samples during a short span of time. a larger sample requires longer survey periods, and this has significant budgetary implications. the surveys conducted since 2001 were done in different months and therefore different seasons. coupled to that, saayman and fouché (2007) indicated in an analysis of all surveys conducted at national parks in south africa since 2001 that the profile stayed consistent, especially in the case of the karoo np. therefore, the sample can be seen as being representative of the total population. economic impact analysis as mentioned previously, the economic impact analysis consisted of compiling a partial input-output model and partial multipliers. in general, an input-output table formalises the relationship between industry inputs and its outputs into technical relationships. leontief (united nations 1999) introduced the fixed-coefficient linear production function, where the output of an industry is related to its inputs, assuming that the inputs are used in fixed amounts (therefore the fixed coefficients). every sector’s total output is a linear function of its intermediate output (i.e. products used by other industries) and final consumption output (i.e. products used by consumers). mathematically, this can be expressed as follows: 112121111 fxaxaxax nn +++= eqn 1 where 1x = output of sector 1; naaa 11211 ,, = the amounts of these outputs that are used in other sectors as inputs (thus, the fixed coefficients), and 1f = the amount of final output that is used by consumers. equation 1 can also be shuffled to show how final demand, 1f , relates to total output and intermediate output: nn xaxaxaxf 121211111 −−−= eqn 2 if the values of the coefficients are known, it is possible to determine the effect that a change in final demand will have on the level of output in the industry in which the demand changes. via the linkages with other industries (the intermediate inputs used from other industries to produce the additional output), it can also be determined by how much the other industries’ output will change. by nature, an input-output model makes certain assumptions. these assumptions include (un 1999): total input is equal to total output in each sector.• each coefficient is smaller than 1.• the sum of all the coefficients should add up to 1 for every • sector. the partial input-output model compiled for the beaufort west economy consists of only nine sectors – (i) accommodation, (ii) restaurants and food, (iii) curios, (iv) tourism services, (v) municipal services, (vi) transport services (including petroleum), (vii) retail, (viii) wholesale and (ix) manufacturing – to reflect the main spending items of tourists to the national park. since the partial inputoutput model was built based on survey data and not in-depth industry data, a number of simplifying assumptions were made. first, since businesses are reluctant to give sensitive information regarding income and profit, some profit assumptions had 29 vol. 51 no. 1 page 4 of 10 the socio-economic impact of the karoo national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #158 (page number not for citation purposes) 30vol. 51 no. 1 page 5 of 10 the spending by chalet tourists is on average r882.58 per group, exceeding that of campers. it is interesting to note that the spending patterns of campers differ substantially from those of visitors in chalets – this is evident from the percentages spent in each category. transport to the karoo np is excluded from the analysis, as it cannot be ascertained where the car has been filled with fuel. spending in the beaufort west area just over seventy (70.8) per cent of the respondents indicated that they spent money outside the karoo np in beaufort west. among the most prominent businesses visited by the respondents are petrol stations and supermarkets (retail/ food), followed by a variety of other businesses, including the butchery (food), restaurants, curio shops, other retailers (chemist, liquor store, bank), car repairs and the wholesaler fruit and veg. table 3 presents a breakdown of the businesses in the area according to the business survey, as well as an indication of the employment levels and the number of permanent versus part-time personnel. economic impact to derive total visitor expenditure the visitor numbers were used. since unit nights already exclude the problem of “spending per person” versus “spending per group”, only the number of campers is divided by the average group size for campers (3.37) to estimate the number of camper groups. according to this method, 3 991 camper groups visited the park in 2006/7. table 2 indicates the magnitude of visitor spending, which was calculated by making use of the above estimates and the average spending per visitor group, as indicated in table 4. it can be seen that visitor spending is estimated at approximately r10.5 million. proportional multiplier analysis as suggested by vaughan, farr and slee (2000), two methods can be employed to determine the economic impact on the local economy, namely iteration and matrix inversion. this research used the second method, matrix inversion, by which the nineby-nine matrix that was created was inverted to determine the effects of a change in demand on output (see table 5). in terms of output, one additional tourist group (change in demand) in each category created the following additional output (see eqn 1). to determine the total effect on income (direct, indirect and induced), the input-output table has to be expanded to include the effect of household spending and wages. in other words, a ten-by-ten matrix was created. the assumption was made that the consumption pattern of the locals in the study area is similar to that of the national consumption pattern (as defined table 1 profile of visitors and community members variable overnight visitors community gender n/a female (63%) and male (37%) language afrikaans (45%), english (53%), other (2%) afrikaans (73%), english (15%), other (12%) average age 48 years 42 years education 98% have post-matric qualifications 56% have a post-matric qualification 40% have matric province of origin gauteng (35%), western cape (26%); north-west (10%); kzn (10%) western cape times visited the park on average, five times over the past three years 39% visited the park at least once, and 61% had not visited the park during march 2006 to march 2007 karoo np effect on personal quality of life n/a 67% felt very positive; 30% indicated the park had no effect facilities 50% had high expectations of the facilities in the park 46% felt the availability of facilities to local residents had increased, 25% indicated no change, 12% indicated a decrease, 17% were not sure reason for visiting the karoo np 76% indicated that they visited the park for relaxation 83% visited the park to learn more about animals and plants table 2 spending per visitor group according to input-output categories spending category visitors camping % visitors in chalets % accommodation r327.20 57.3% r673.35 76.3% curios r12.42 2.2% r8.04 0.9% food and restaurants r107.20 18.8% r119.53 13.5% tourism services r65.40 11.5% r36.76 4.2% retail r25.90 4.5% r5.68 0.6% transport r32.80 5.7% r39.22 4.4% municipal r0.00 0.0% r0.00 0.0% wholesale r0.00 0.0% r0.00 0.0% manufacturing r0.00 0.0% r0.00 0.0% total r570.92 100% r882.58 100% original research saayman, saayman & ferreira koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 15 8 (page number not for citation purposes) gives a good indication of how business has developed in the region. the karoo np was founded in 1979. none of the accommodation units that are still available, curio shops or food and restaurant businesses are older than 28 years. most are less than 15 years old, with only one curio shop and one restaurant being between 15 and 20 years old. it is also interesting that 78% of the retail stores are also younger than the karoo np, and that all the wholesalers and manufacturers that were surveyed have existed for less than 28 years. only one transport business exceeds 28 years of existence. only 4% of the businesses indicated that they owed their existence to the karoo np. these businesses are all retailers. many businesses indicated that they relied heavily on tourist spending (see table 9), which might be an indication that the influence of the karoo np on tourism activity in the area is not taken into account by these businesses. it would be an interesting future study to examine whether the existence of the karoo np has changed tourism patterns to beaufort west substantially (given that data would be available). intuitively it has, since for many beaufort west was initially just an overnight stop on the route between cape town and johannesburg. yet the visitor survey revealed that visitors to the karoo np spent on average almost two days in the beaufort west area and most (86%) of these visitors indicated that they also shopped in town. as indicated in table 3, the accommodation industry is, as expected, very reliant on labour, with the five accommodation units providing employment for 78 people. this figure is put into perspective when it is compared to the 308 people working in the 55 surveyed retailers (see table 3). it should be noted that, in 98% of the cases, the employed workers live in the area, which is encouraging, since employment opportunities are being created in beaufort west, which has one of the highest rates of unemployment in the western cape. on average, accommodation establishments sell 68% of their business to tourists and 32% to locals. the latter entails hiring of venues for meetings and conferences, as well as for catering. tourists are also the main customers of tourism and recreation services, while transport businesses, restaurants and food stores, and curio shops are also very reliant on tourism for their existence. this is also indicated in table 9, which shows that the beaufort west economy is, in many respects, dependent on tourism for income and employment. it is difficult to determine the exact number of jobs that can be ascribed to the existence of the karoo np. the karoo np itself employs 40 people, which is relatively few compared to other parks such as the addo, kruger and tsitsikamma national parks. the businesses also indicated the percentage of their turnover that they ascribed to the karoo np being in the vicinity. guest houses and bed-and-breakfast establishments indicated that they owed 12% of their turnover to the karoo np. in the 2005 nationwide input-output table). it is evident that spending directly translates into income for a business and, as the business expands its production, the income of other businesses also expands. this process captures the direct and indirect effects of income for businesses. not all income that businesses receive translates into income for consumers. the income is used to buy stock, pay for operational expenses and services, pay labourers, and for profits. what is more interesting to know is the latter – in other words, how does this translate into income for consumers (households) in the beaufort west area? therefore, we were only interested in finding the induced income effect of the additional spending. the results of the analysis are shown in table 5. it is also noteworthy that the income and output multipliers for the camping groups are lower than those of the chalets. this is to be expected, however, since campers spend less on accommodation (which is a labour-intensive activity) than groups in chalets and more on retail items and food – which is more capital-intensive and requires more stock purchases. the change in output due to visitor spending in each industry in the proportional multiplier model is indicated in table 6. also indicated in table 6 is the business income (direct and indirect effect) generated due to one additional group. if the above multipliers are applied to total spending by all visitors, the production and income effect of their spending would be as follows (table 7): while these estimates reflect the spending by visitors, the spending of the karoo np in its operations has not yet been reflected. if a similar process is repeated for karoo np expenditure (based on the allocations described above), the output multiplier for karoo np expenditure is 1.26, while the income multiplier is 0.88. the total impact of the karoo np on the local economy is indicated in table 8. employment and business development the years that the different businesses have been in existence 31 vol. 51 no. 1 page 6 of 10 table 3 breakdown of businesses and employment in the study area type number average number of employees permanent part-time accommodation units 5 15.6 14.2 1.4 curio shop 5 7.8 5 2.8 food and restaurant 11 12.7 9.5 3.2 tourism services 2 3.5 3.5 0 retailers 55 5.6 4.9 0.8 transport and petrol stations 4 34.3 27.3 7 wholesalers 3 25.7 22.7 3 manufacturers 4 5.7 4.7 1 table 4 estimated total expenditure by visitor groups, 2007 category number of groups average spending per group total spending chalets 9 373 r882.58 r8 272 422 camping 3 991 r570.92 r2 278 542 total 13 364 r10 550 964 (source: saayman & bothma 2007) the socio-economic impact of the karoo national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #158 (page number not for citation purposes) 32 table 5 multipliers category change in demand change in output change in income output multiplier income multiplier chalet r882.58 r1 300.74 r353.42 1.47 0.40 camp r570.92 r 818.34 r211.35 1.43 0.39 table 6 sectoral changes in output and income sector chalet: change in output chalet: change in income camp: change in output camp: change in income accommodation 674.52 674.79 327.90 328.07 curios 8.04 13.34 12.42 15.74 food and restaurants 121.87 204.05 108.61 160.08 tourism/recreation services 37.34 60.80 65.75 80.45 retail 235.98 312.06 160.33 207.98 transport 82.44 115.93 55.42 76.39 municipal 84.45 104.07 50.97 63.26 wholesale 47.78 58.61 28.77 35.55 manufacturing 8.32 12.09 8.16 10.52 total 1 300.74 1 555.75 818.34 978.06 table 7 impact of visitor spending on production and income category total spending output effect income effect chalet r8 272 422 r12 160 460 r3 308 969 camp r2 278 542 r3 258 315 r888 631 total r10 550 964 r15 418 775 r4 197 600 table 8 total impact of the karoo national park on the local economy category total spending output effect income effect chalet r8 272 422 r12 160 460 r3 308 969 camp r2 278 542 r3 258 315 r888 631 park r4 187 378 r5 276 096 r3 684 893 total r14 738 342 r20 694 871 r7 882 493 curio shops indicated that 8.8% of their business was due the karoo np, with restaurants indicating a figure of 15%. retailers and service stations also indicated that they respectively owed 8% and 20% of their turnover to the karoo np, and tourism and recreation services estimated this contribution at 15%. if this is taken as an indication of the number of employment opportunities created due to the karoo np, it can be concluded that the karoo np led to only 102 job opportunities in the businesses surveyed (a linear relationship between turnover and employment is assumed). the survey captured 142 beds as part of the accommodation sector, while the beaufort west tourism association indicates that 300 beds are available. thus, approximately 50% of the accommodation products were surveyed. if this proportion is true for all the business sectors, extrapolating the employment results would indicate that 204 additional jobs were created by the karoo np. with the direct employment by the karoo np, total employment is thus 244. seen in the light that there are 7 306 households in beaufort west, it can be concluded that the karoo np, directly and indirectly, provides employment opportunities to 3.34% of all the households. social impact the analysis of the social impacts of tourism is, however, affected by a number of elements, for example collective lifestyles, individual behaviour and family relationships. the following section focuses on the influence of the karoo np on the community and the role of the karoo np in beaufort west. table 10 gives the results of the influence of the park on the community by using a scale ranging from -3, being very negative, to 3, being very positive. according to the respondents, the karoo np has a very positive impact (72%) on the beaufort west community. the karoo np also affects the community’s personal quality of life positively (67%) (table 10). therefore, the perception by the community of the way the karoo np influences them is overall very positive. if one analyses the role that the karoo np plays in the beaufort west community (table 11), the respondents indicated that the appearance of the area had improved (70%) due to the karoo np; the number of tourists visiting beaufort west had increased (68%); and the pride of the residents towards beaufort west was positive (65%). in general, the karoo np is seen as contributing to the upliftment of the area by adding to the tourism product offered by the area and improving people’s perceptions of the area. all these aspects are positive. discussion based on the results of the research conducted at the karoo np and among the businesses and community of beaufort west, the following results were identified. vol. 51 no. 1 page 7 of 10 original research saayman, saayman & ferreira koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 15 8 (page number not for citation purposes) 33 vol. 51 no. 1 page 8 of 10 table 9 employment and other ratios type number total employment % turnover due to park tourist/local ratio accommodation units 5 78 12% 68:32 curio shops 5 39 8.8% 37:63 food and restaurants 11 140 15% 32:68 tourism services 2 7 15% 60:40 retailers 55 308 8% 20:80 transport and petrol stations 4 138 20% 40:60 wholesalers 3 77 16.7% 20:80 manufacturers 4 23 9.3% 11:89 table 10 influence of the karoo np on the community statement very negative no effect very positive -3 -2 -1 0 1 2 3 personal quality of life 1% 2% 1% 21% 8% 25% 42% beaufort west community 4% 3% 2% 10% 10% 28% 44% table 11 community perceptions of the role of the karoo np in beaufort west social indicators decreased/ worse increased/ better no change do not know the appearance of the area 4% 70% 10% 16% employment opportunities in beaufort west 10% 60% 18% 12% the range of things to do in beaufort west 9% 55% 22% 14% the number of people in the area 3% 46% 21% 30% property values in the area 5% 49% 23% 23% crime levels 17% 26% 36% 21% participation in community activities 8% 45% 27% 20% entertainment opportunities 15% 41% 25% 19% prices of some goods and services 14% 35% 31% 20% the pride that beaufort west residents have in their town 8% 65% 13% 14% the overall cost of living 13% 37% 28% 22% the opportunities to meet new people 7% 54% 25% 14% opportunities for local business 6% 55% 19% 20% the number of tourists visiting beaufort west at other times of the year 7% 68% 7% 18% public funding for community activities 11% 35% 21% 33% the rights and civil liberties of local residents 11% 37% 25% 27% interaction between locals and tourists 10% 41% 20% 29% facilities available to local residents 12% 46% 25% 17% social and moral values 13% 37% 26% 24% the skill base for events management in beaufort west 7% 48% 20% 25% the turnover for local businesses 8% 45% 18% 29% the socio-economic impact of the karoo national park original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #158 (page number not for citation purposes) 36 34 first, if one compares this research with the study conducted by saayman & saayman (2006a) in the addo elephant national park, which is the only other case study in south africa in which the same method of research was applied, the following differences become apparent: businesses in beaufort west clearly cannot distinguish between tourists and local shoppers, and this is because many tourists pass through beaufort west when travelling and most tourists only make use of the karoo np as a ‘stop-over’ destination. in the case of addo elephant national park, the main drawcard in the area is the park, which is not the case in beaufort west. added to this is the fact that the addo elephant national park is surrounded by smaller villages, for example, addo and paterson, where the locals know each other, which is less of a feature in beaufort west. second, the multipliers created by the matrix inversion process are higher than those of the addo national park, which is to be expected given that beaufort west is a much larger town than addo, paterson or kirkwood (see saayman & saayman, 2006a). the income multiplier generated by the matrix inversion process for groups holidaying in chalets is again in line, but slightly higher for camping groups than that found for the addo elephant park. the rationale behind the higher multipliers is intuitively correct, since beaufort west is not situated close to any larger town that creates more extensive leakages in terms of employment creation and spending by households. third, the fact that the park borders on beaufort west and that karoo np staff are living in the community has a positive impact on community perceptions of the park. fourth, the smaller the operation (for example, accommodation, tourist numbers and activities), the lesser the socio-economic impact. hence the size of operations definitely plays a role. national parks, and specifically the karoo np, should offer more activities, more species and more accommodation facilities, where possible, to get tourists to stay longer and spend more money. this will increase the magnitude of the socio-economic impact. fifth, the socio-economic impact of a tourist product is influenced mainly by length of stay and magnitude of spending, as indicated by page (2007), dwyer, forsyth and spurr (2005) and mathieson and wall (1982). as has been mentioned, these aspects are influenced by aspects such as variety and scope of activities, in other words what does the park offer or what can the region offer tourists to do? the fewer species and variety of species the park offers, the shorter the stay. this applies to activities as well. in the case of the karoo np, the number of species is significantly lower than in other national parks, such as the addo elephant park, the kruger national park and the mokala national park. lastly, improved communication can also increase awareness among tourists, businesses and other product owners of what the park wants to achieve and how the community needs to work together to achieve this. according to andriotis (2006), gursoy, jurowski and uysal, (2002), page et al. (2002), slabbert (2007), saayman (2000), fredline and faulkner (2001), and urtasun and gutiérrez (2005), the following are the most important in social impact analysis: involvement in the park, community attachment, contact with tourists and state of local economy. therefore, to grow the socio-economic impact of the park it is of paramount importance that tourism products in the region are developed and promoted, and that the community must be informed regularly. added to this, coetzee (2004) has pointed out that, from a socio-economic and sustainability point of view, community perceptions are important. in this regard, communication plays an important role, since it creates perceptions (streuders 2008). conclusion the purpose of this research was to determine the socioeconomic impact of the karoo national park (karoo np). this was done by estimating the economic impact of the karoo np on the local economy. secondly, the impact of tourism business development in the karoo district was estimated, as well as the way in which these businesses benefited the community. thirdly, the employment generated as a result of the karoo np and tourism activities in the area was determined, as well as the community’s satisfaction with the karoo np. the results indicated that the karoo np has an impact on employment, production and general income in the region, but not to as significant an extent as documented for the addo elephant national park. only 4% of the businesses owe their existence to the karoo np, yet many of these businesses rely on tourists’ spending – which might be an indication that the influence of the karoo np on tourism activity in the area was not taken into account by these businesses. guest houses and bed-and-breakfast establishments indicated that they owed 12% of their turnover to the karoo np. curio shops indicated that 8.8% of their business was due the karoo np, with restaurants indicating a 15% contribution by the park. retailers and service stations indicated that they owed 8% and 20% respectively of their turnover to the karoo np, and tourism and recreation services estimated this contribution at 15%. it would be beneficial for park management to increase communication between businesses in the area, as well as with the community by means of a park forum. both businesses and the community need to understand what the aims and objectives of the park are. the results also support the concern that rural areas usually face high leakages, and thus the local economy does not benefit to its full potential. however, the impact is smaller than in the case of the addo elephant national park, as beaufort west is not situated close to any larger towns, such as port elizabeth and george, where more extensive leakages are created in terms of employment and spending by households. however, in terms of employment creation, the fact that 98% of all workers live in the area is an indication that the benefits of the karoo np reach the local community and that it certainly contributes towards social upliftment in the vicinity. park management could also consider a local ‘wildcard’ (the latter is a loyalty card that tourists buy and gives free access to national parks for a period of one year that can be renewed) in order to increase the number of day visitors. added to this, the park could consider open days when communities are invited to visit the park free of charge and park management can then interact with the visitors. with regard to future research, it is important to examine whether the existence of the karoo np changed tourism patterns to beaufort west. the same research should also be conducted in other national parks, since this research clearly showed how two national parks, namely the karoo national park and addo elephant national park, differ in their socio-economic impacts, and thus offer important lessons to be learned. acknowledgements the authors would like to acknowledge glenn phillips from sanparks, for financial assistance, as well as the karoo national park manager. the nrf provided financial assistance. the anonymous reviewers are thanked for their invaluable input and comments. references andriotis, k., 2006, ‘hosts, guests and politics: coastal resorts morphological change’, annals of tourism research 33, 1079– 1098. coetzee, w.j.l., 2004, ‘an integrated sustainable tourism development strategy for the vredefort dome as a world heritage site’, phd dissertation, north-west university. vol. 51 no. 1 page 9 of 10 original research saayman, saayman & ferreira koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 15 8 (page number not for citation purposes) saayman, m. & bothma, l., 2007, a marketing profile of visitors to the karoo national park, north-west university, potchefstroom. saayman, m. & fouché, m., 2007, executive summary of research profiles of selected national parks in south africa, north-west university, potchefstroom. saayman, m. & saayman, a., 2004, ‘economic impact of cultural events’, south african journal for economic and management sciences 7(4), 629–642. saayman, m. & saayman, a., 2006a, ‘creating a framework to assess the economic contribution of national parks in south africa: the case of the addo elephant national park’, tourism economics 12(4), 619–633. saayman, m. & saayman, a., 2006b, ‘estimating the economic contribution of visitor spending in the kruger national park to the regional economy’, journal of sustainable tourism 14(1), 67–81. slabbert, e., 2007, ‘the social impact of tourism’, in m. saayman (ed.), en route with tourism, 3rd edn., pp. 149–177, leisure publications, potchefstroom. south african national parks, 2007, ‘south african national park – karoo national park’, viewed 17 january 2008 from http://www.sanparks.org/parks/karoo. statssa, 2007, ‘statistics south africa’, viewed 28 june 2007 from http://www.statssa.gov.za. streuders, c., 2008, ‘communication efficacy of karoo national park’, master’s thesis, north-west university. trochim, w.m., 2006, the research methods knowledge base, 2nd edn., atomic dog publishing, cincinnati. tustin, d.h., ligthelm, a.a., martins, j.h. & van wyk, h., 2005, marketing research, university of south africa press, pretoria. urtasun, a. & gutiérrez, i., 2005, ‘tourism agglomeration and its impact on social welfare: an empirical approach to the spanish case’, tourism management 27, 901–912. united nations (un), 1999, studies in methods: handbook of national accounting. handbook of input-output table compilation and analysis, series f, no. 74, united nations, new york. van der merwe, l.h., 2008, ‘the socio-economic impact of klein karoo national arts festival in oudtshoorn’, mcom thesis, north-west university. vaughan, d.r., farr, h. & slee, r.w., 2000, ‘estimating and interpreting the local economic benefits of visitor spending: an explanation’, leisure studies 19, 95–118. walpole, m.j. & goodwin, h.j., 2000, ‘local economic impacts of dragon tourism in indonesia’, annals of tourism research 27(3), 559–576. department of local government and housing, 2006, national imbizo focus week. pre-imbizo report, beaufort west municipality, beaufort west. dwyer, l. forsyth, p. & spurr, r., 2005, ‘economic impacts and benefits of sport events: a cge perspective’, in j. allen (ed.), the impacts of events: international event research conference proceedings, sydney, australia, july 13–14, 2005, n.p. fauzi, a. & buchary, e.s., 2002, ‘a socio-economic perspective of environmental degradation at kepulauan seribu marine national park, indonesia’, coastal management 30, 167–181. fredline, e. & faulkner, b., 2001, ‘residents’ reactions to the staging of major motor sport events within their communities: a cluster analysis’, event management 7, 103– 114. fredline, l., jago, l. & deery, m., 2003, ‘the development of a generic scale to measure the social impact of events’, event management 8, 23–37. govender, y., jury, m.r., mthembu, a., hatesse, s. & bulfoni, e., 2005, ‘socio-economic status and development potential for a rural community on the maputaland coast of south africa’, south african geographical journal 87(1), 37–42. gursoy, d., jurowski, c. & uysal, m., 2002, ‘resident attitudes: a structural modeling approach’, annals of tourism research 31(3), 495–516. loader, j.a., 1994, ‘national parks and social involvement – an argument’, koedoe 37(1), 137–148. macleod, d.v.l., 2001, ‘parks or people: national parks and the case of del este, dominican republic’, progress in development studies 1(3), 221–235. mathieson, a. & wall, g., 1982, tourism: economic, physical and social impacts, longman group limited, london. municipal demarcation board, south africa, 2006, ‘municipal profile’, viewed from http://www.demarcation.org.za/ mu n ic prof i leon l i ne/websiterouter.a spx?ht t p://w w w. demarcation.org.za/municprofileonline/index.aspx. myburgh, e. & saayman, m., 2002, ecotourism in action: guidelines and principals, 2nd edn., pu for che, potchefstroom. nkungwana, m., 2007, beaufort west integrated development plan final document, beaufort west municipality, beaufort west. page, s.j., 2007, tourism management: managing for change, 2nd edn., butterworth-heinemann, oxford. page, s.j., brunt, p., busby, g. & connell, j., 2002, tourism: a modern synthesis, thomson, london. pelser, a., 2003, ‘omgewingsbewaring in ‘n nuwe era: ‘n nis vir die geestesen sosiale wetenskappe’, tydskrif vir geestesweetenskappe 43(3/4), 164–176. saayman, m., 2000, en route with tourism, 2nd edn., leisure publications, potchefstroom. vol. 51 no. 1 page 10 of 1035 gotze.qxd analysis of the riparian vegetation (ia land type) of the proposed vhembe-dongola national park, limpopo province, south africa a.r. götze, s.s. cilliers, h. bezuidenhout and k. kellner a.r. götze, s.s. cilliers, h. bezuidenhout and k. kellner. 2003. analysis of the riparian vegetation (ia land type) of the proposed vhembe-dongola national park, limpopo province, south africa. koedoe 46(2): 45-64. pretoria. issn 0075-6458. the establishment of the vhembe-dongola national park has been an objective of several conservationists for many years. the ultimate objective is that this park would become a major component of a transfrontier park shared by botswana, zimbabwe and south africa. the aim of this study was to identify, classify and describe the plant communities present in the ia land type of the proposed area for the park. sampling was done by means of the braun-blanquet method. a total of 70 stratified random relevés were sampled in the ia land type. all relevé data was imported into the database turboveg after which the numerical classification technique twinspan was used as a first approximation. subsequently braun-blanquet procedures were used to refine data and a phytosociological table was constructed, using the visual editor, megatab. from the phytosociological table four plant communities were identified and described in the ia land type. the ordination algorithm, decorana, was applied to the floristic data in order to illustrate floristic relationships between plant communities, to detect possible gradients in and between communities and to detect possible habitat gradients and/or disturbance gradients associated with vegetation gradients. key words: national parks, phytosociology, vegetation classification, braun-blanquet procedures, savanna biome. a.r. götze, s.s. cilliers and k. kellner, school of environmnetal sciences and development, section botany, rivate bag x6001, potchefstroom university for c.h.e., potchefstroom, 2531; h. bezuidenhout, conservation services, s.a. national parks, p.o. box 110040, hadison park, kimberley, 8306 republic of south africa. issn 0075-6458 45 koedoe 46/2 (2003) introduction the establishment of the vhembe-dongola national park has been an objective of several conservationists for many years and was finally approved by south african national parks in 1994 (robinson 1996). this action follows after a long history of proposals to this regard, starting in 1922 from an initiative of general j.c. smuts to form the dongola botanical reserve (carruthers 1992). the conservation value of this area lies in it’s rich biodiversity, it’s great scenic beauty and the cultural importance of the archaeological treasures of mapungubwe which is included in the proposed area for this national park. the artifacts found at mapungubwe rank among the most important pieces of ancient history yet found in sub-saharan africa (willcox 1966). the significance of this proposed park and its surrounding areas is further enhanced by the potential role of the area as a sanctuary for some of the most endangered mammals on earth, such as the black rhino and the african wild dog (robinson 1996). the ultimate objective is that the vhembedongola national park would become a major component of a transfrontier park shared by botswana, zimbabwe and south africa. the total area of this transfrontier park could potentially add up to 2530 km² (robinson 1996). certain parts of the proposed area for the vhembe-dongola national park is, however, extremely degraded because of a combination of anthropogenic influences, such as mining, cultivation, overgotze.qxd 2005/12/09 10:52 page 45 grazing and trampling by livestock as well as other management malpractices (götze 2002). a large military base of the former south african defence force was also situated on the farm greefswald, and contributed to the disturbance and degradation of the area. old campsites litter the proposed area and the scars left by forgotten roads, quarries and dumpsites are a cruel sight to the eye (götze 2002). it is, therefore, evident that specific restoration and rehabilitation practices should be implemented in an attempt to restore the land to a higher potential for the conservation of our precious fauna and flora. of all the land types in the vhembe-dongola national park the ia land type which include the riparian areas seems to be the most disturbed, mainly because of the extensiveness of this linear landscape, the high fertility of the soils and the consequent establishment of irrigation lands (götze 2002). the riparian zone can be identified as the area of land adjacent to a stream or river, which is, at least periodically, influenced by fluctuations of the water level (rogers 1995). although the ia land type is overutilised by small and large livestock, the destruction caused by migratory tuli elephants, from neighboring botswana, is an additional cause for concern. furthermore, riparian zones warrant special attention since their positioning in the landscape ensures that they play direct roles in the functioning of both the river system and the terrestrial landscape and are one of the major centers of biodiversity in a global context (naiman et al. 1993). the narrow spatial dimensions and open endedness of riparian wetlands make them highly sensitive to landscape changes, with major consequences to the river ecosystems which they buffer from terrestrial influences (naiman & décamps 1990; rogers 1995). the current state of the proposed area for the vhembe-dongola national park and the uniqueness of the area with respect to diversity and cultural importance warrant the establishment of detailed management practices. in the past decade conservation management has moved more in the direction of total environmental management. emphasis in conservation is not on strictly policed, protected areas, primarily for large mammals anymore, but on sustainable resource use, maintenance of ecological processes and conservation of genetic diversity (winterbach 1998). a sound knowledge of the vegetation ecology of areas of conservation significance is essential for the establishment of efficient wildlife and environmental management programs and the compilation of conservation policies (bredenkamp & theron 1978; bredenkamp et al. 1993; bezuidenhout 1996; van rooyen et al. 1981). different ecosystems react differently to certain management practices (bredenkamp & theron 1976) and therefore, in order to formulate a management policy, where proper land use is emphasised, the classification of the vegetation is essential (van rooyen et al. 1981; bezuidenhout 1993). it is widely recognised that a detailed description, identification, classification and mapping of the vegetation, form the basis for sound land-use planning and management (fuls 1993; bezuidenhout 1996; brown 1997). the aim of this study is to identify, classify and describe the plant communities present in the ia land type of the proposed area for the vhembe-dongola national park. this study forms part of a bigger study on the vegetation of all the land types included in the proposed vhembe-dongola national park. study area location the proposed area for the vhembe-dongola national park is centered on the confluence of the limpopo and shashi rivers, in the limpopo province, on the international borders between botswana, zimbabwe and south africa (robinson 1996) (fig. 1). the primary core area spans from pontdrif in the west to weipe in the east incorporating 22 koedoe 46/2 (2003) 46 issn 0075-6458 gotze.qxd 2005/12/09 10:52 page 46 issn 0075-6458 47 koedoe 46/2 (2003) fi g. 1 r el at iv e lo ca tio n of th e ia la nd ty pe in th e pr op os ed a re a of th e v he m be -d on go la n at io na l p ar k gotze.qxd 2005/12/09 10:52 page 47 farms with a surface area of 28 000 ha (fig. 1). land types in south africa land types have been used as a map unit denoting land, mappable at 1:250 000 scale, over which there is marked uniformity of climate, terrain form and soil pattern. the ia land type (fig. 1) includes the riverine and non-perennial wetland vegetation with deep to moderately deep soils with a high silt and clay content. other land types which occur within the proposed area of the vhembe-dongola national park are the ae, db, fb and ib land types (fig. 1) (land type survey staff 2000). climate the climate is semi-arid with the long-term mean annual rainfall ranging from 350 mm to 400 mm per annum (willcox 1966; robinson 1996). rainfall is highly variable and usually falls during the summer months between october and march. evaporation from free water surfaces is in excess of 2 500 mm per annum for the largest part of the area. in summer the temperatures rise as high as 45 °c (weather bureau 1986). the winters are mild, although frost may occur in low-lying areas (robinson 1996). hydrology surface drainage is mostly in a northerly direction towards the limpopo river. none of the rivers in the area, including the limpopo, are perennial. ground water supplies are generally poor in the region except where these occur along well-developed fault lines (van den heever 1983; robinson 1996). geology, geomorphology and soils the area is between 300 m and 780 m above sea level. south of the limpopo river the ground tends to be flat with sandstone and conglomerate ridges and koppies. closer to the limpopo river the flats give way to a more rugged, hilly terrain. the soils of the limpopo valley are derived from rocks of the archaean granite formations in the south and, more generally, from the karoo-system (stormberg, ecca and beaufort series). the soils vary from red-brown sandy loam to dark brown clays with high silt content. large areas are characterised by sandy, limerich soils (robinson 1996). the geology of the ia land type consists of sand and alluvium from the quaternary system and deep red/brown, alluvial soil forms (oakleaf) and some duplex soils (valsrivier & arcadia) dominate the soils of this land type (land type survey staff 2000). vegetation according to van rooyen & bredenkamp (1996), the area is situated in vegetation type no. 10, mopane bushveld, while acocks (1988) placed it in veld type no.15, also mopane veld. according to acocks (1988), there are two major blocks of colophospermum mopane veld. one of these blocks of colophospermum mopane is situated in the limpopo valley north of the soutpansberg and the other to the east of the soutpansberg. the conservation status of the eastern block, which occurs in the kruger national park is sound with 89% of the area protected. the western block is not well represented in conservation areas with only 0,36% being formally protected (berry 1994; robinson 1996). the former transvaal provincial administration (transvaal provincial administration 1989) recognised three main vegetation units in the region: the riparian fringe along the limpopo river and its tributaries, the acacia-salvadora communities of the limpopo flats (including flood plains) and wetland areas, and the mixed mopane veld on ridges and flats south of the riparian fringe and flood plains. other vegetation studies done in the vhembe-dongola national park and surrounding areas in the past, include several unpublished studies, namely a terrain and vegetation evaluation by the former south koedoe 46/2 (2003) 48 issn 0075-6458 gotze.qxd 2005/12/09 10:52 page 48 african defence force (south african defence force 1986a, 1986b), a study of vegetation types in the limpopo-venetia nature reserve (o’connor 1991) and a vegetation survey of the maramani/tuli area in southwestern zimbabwe (timberlake & mapaure 1999). cultural historical assets robinson (1996) stated that the proposed area of the vhembe-dongola national park has numerous archaeological sites dating from the early stone age to the present. many of these sites, which are concentrated in the area of the confluence of the limpopo and shashi rivers, are of major importance and scientific value. of particular interest are the zhizo site (ad 700-900) on the farm schroda, and mapungubwe hill and the adjoining bambandyanalo (ad 1100-1250) situated on greefswald. according to voigt & plug (1981) the mapungubwe site is considered to be of major importance in subsaharan africa and is the most remarkable iron age site in south africa. additional features of importance are the numerous rock paintings and petroglyphs found in the area. methods stratification with the aid of 1:50 000 topographical maps, geological maps and vegetation and soil maps compiled by the old south african defence force 1986 a,b), a land type map of the area (land type survey staff 2000) and aerial photographs, a preliminary evaluation of the study area was made. in conjunction with the maps, a detailed reconnaissance of the area was done to determine homogeneous areas in the vegetation, before sampling commenced. land types were used as a means of primary stratification of the study area. sampling making visual estimates, sampling was done by means of the braun-blanquet method (muellerdombois & ellenberg 1974). a total of 70 stratified random relevés were sampled in the ia land type. braun-blanquet cover abundance values were appointed to all plant species encountered in the relevés (table 1). plant species identification was done by following arnold & de wet (1993), but are updated according to the precis floristic database of south africa, managed by the national botanical institute in pretoria. soil classification is according to the soil classification work group (1991). trees and shrubs were distinguished from each other using the guidelines set by edwards (1983). trees are classified as rooted, woody, self-supporting plants over 2 m high with one or up to three definite trunks and shrubs are classified as rooted, woody, self-supporting, multistemmed or single-stemmed plants less than 2 m high (edwards 1983). the habitat conditions including geology, soil forms and other edaphic factors, slope, aspect and the rockiness of the soil surface, of each relevé, were qualitatively described. this information was used in the description of the different plant communities. the coordinates of each relevé were determined and noted with the use of a gps. data processing all relevé data has been imported into the database turboveg (hennekens 1996a). the numerical classification technique twinspan (hill 1979a), which is regarded as a successful approach for classification by several phytosociologists (mucina & van der maarel 1989; bredenkamp & bezuidenhout 1995; cilliers 1998) was used as a first approximation for the floristic data. subsequently braun-blanquet procedures (bezuidenhout et al. 1996) were used to refine data and construct a phytosociological table (table 1) using a visual editor, megatab (hennekens 1996b). uncommon species not diagnostic for any of the communities are not included in the table, but can be witnessed in götze (2002). using the phytosociological table and the habitat information gathered during the sampling period, the different plant communities were identified and described. no attempt was made in this study to formally name the plant communities. this will only follow after extensive studies on savanna vegetation in south africa to prevent synonymy of syntaxa names. an ordination algorithm, decorana (hill 1979b), was applied to the floristic data in order to illustrate floristic relationships between plant communities and to detect possible habitat gradients and/or disturbance gradients associated with vegetation gradients. issn 0075-6458 49 koedoe 46/2 (2003) gotze.qxd 2005/12/09 10:52 page 49 koedoe 46/2 (2003) 50 issn 0075-6458 ta bl e 1 p hy to so ci ol og ic al ta bl e of th e ve ge ta tio n of th e ia la nd ty pe o f t he v he m be -d on go la n at io na l p ar k. gotze.qxd 2005/12/09 10:53 page 50 issn 0075-6458 51 koedoe 46/2 (2003) ta bl e 1 (c on tin ue d) gotze.qxd 2005/12/09 10:53 page 51 koedoe 46/2 (2003) 52 issn 0075-6458 ta bl e 1 (c on tin ue d) gotze.qxd 2005/12/09 10:53 page 52 issn 0075-6458 53 koedoe 46/2 (2003) ta bl e 1 (c on tin ue d) gotze.qxd 2005/12/09 10:53 page 53 koedoe 46/2 (2003) 54 issn 0075-6458 ta bl e 1 (c on tin ue d) gotze.qxd 2005/12/09 10:53 page 54 results and discussion classification from the phytosociological table (table 1) the following plant communities, sub-communities and variants were recognised: 1. salvadora australis-cucumis zeyheri community 1.1 colophospermum mopane-eragrostis trichophora sub-community 1.2 indigastrum costatum subsp. maculata-setaria verticillata sub-community. 1.2.1 aristida congesta-leucas sexdentata variant. 1.2.2 eragrostis racemosa-acacia tortilis variant 1.3 combretum imberbe-abutilon ramosum sub-community 2. hyphaene petersiana acacia tortilis community 2.1 ximenia americana-flueggea virosa sub-community 2.2 cordia monoica sub-community 3. croton megalobotrys-combretum microphyllum community 3.1 cenchrus ciliaris-faidherbia albida sub-community. 3.2 acacia schweinfurthii maytenus senegalensis sub-community. 4. diplachne fusca acacia xanthophloea community description of the plant communities 1. salvadora australis-cucumis zeyheri community this community mostly occurs on the floodplain, toward the outskirts of the lush riverine forest belt found along the main river systems in the study area. a wide spectrum of soil types was found, with oakleaf being the dominant soil form and hutton, and valsrivier soil forms in close support. with the exception of one sub-community, the soils that are found in this community have a high silt and moderate clay content. this community is situated higher above the annual flood level than its neighbouring communities, and as a result, appears to be drier. the slight slope is generally in the main direction leading towards the limpopo river. a further observation was that the soil surface is compacted through years of overgrazing by cattle and a large number of game that frequent the area, and therefore a high degree of surface runoff is apparent. in some extreme cases this has led to a degree of surface erosion. the vegetation of the salvadora australis – cucumis zeyheri community varies between open woodland, thickets and forests. the diagnostic species of this community are found in species group a (table 1), and include the forbs cucumis zeyheri, boerhavia erecta, heliotropium ovalifolium, phyllanthus parvulus and ipomoea sinensis subsp. blepharosepala. salvadora australis (species group k, table 1) and the herbaceous cucumis zeyheri, boerhavia erecta (species group a, table 1) and setaria verticillata (species group p, table 1) dominate the vegetation of this community. the absence of the species that are found in species group o (table 1) differentiates this community from most of the other communities in the ia land type. three sub-communities were identified in this community: 1.1 colophospermum mopane-eragrostis trichophora sub-community this sub-community is situated on the very outskirts of the floodplain. it is situated on sandy hutton soils with surface erosion clearly visible as there are usually drainage systems running off towards the limpopo river, bisecting this vegetation unit. diagnostic species of this sub-community are found in species group b (table 1), which includes the tree species colophospermum mopane the grass aristida adscensionis and the forbs phyllanthus angolensis, issn 0075-6458 55 koedoe 46/2 (2003) gotze.qxd 2005/12/09 10:53 page 55 becium filamentosum and melhania rehmannii. the tree layer is dominated by colophospermum mopane (species group b, table 1) and to a lesser extent salvadora australis (species group k, table 1). the shrub layer is sparse. the grass species eragrostis trichophora (species group k, table 1) and the annual grass aristida congesta subsp. congesta (species group d, table 1) dominate the herbaceous layer. the tree layer of this sub-community reaches heights of between 2.5 m and 4.0 m and has an average crown cover of 35 %. the shrub layer has an average height of 1.4 m and the cover varies between 10 % and 35 %. the herbaceous layer reaches an average height of 0.5 m and a crown cover of between 45 % and 65 %. 1.2 indigastrum costatum subsp. maculata-setaria verticillata sub-community. the soil forms occurring in this sub-community are hutton, oakleaf and valsrivier. the sub-community is situated on a level floodplain of which the soil surface gets trampled to a fine dust during the dry season. the area is targeted by high degree of grazing by game and, therefore the area shows signs of degradation through the appearance of bare patches in the grass layer. due to flooding in high rainfall years, as well as the high resilience of this floodplain area, any restoration practices are uncertain at this stage. the vegetation of this sub-community varies between open woodlands and thickets. the species that are diagnostic to this sub-community are found in species group c (table 1) and include indigastrum costatum subsp. maculata, tribulus terrestris and the scarce herbaceous climber kedrostis limpompensis. the species of species group f (table 1), indicates a similarity between this sub-community and the colophospermum mopane – eragrostis trichophora sub-community (1.1). the tree layer of this sub-community reaches heights of between 2.2 m and 8 m (in some cases > 8 m). the crown cover of the tree layer varies between 20 % and 45 %. the shrub layer has an average height of 1.4 m and the crown cover varies between 5 % and 25%. the herbaceous layer is well developed with an average crown cover of 80% and heights varying from 0.2 m to 0.9 m. the indigastrum costatum subsp. maculata – setaria verticillata sub-community has two variants, which differ from each other with respect to soil types and vegetation structure. 1.2.1 aristida congesta-leucas sexdentata variant. this variant comes in the form of open woodlands with salvadora australis (species group k, table 1) occurring in bush-clumps. the soils that are found in this variant have a high silt content and consist of the hutton and oakleaf soil forms. there are no diagnostic species to this variant. the absence of species groups e and h (table 1) as well as a low frequency of the tree species acacia tortilis and the shrub abutilon austro-africana (species group k, table 1) do, however, differentiate this variant. it is further distinguishable from the eragrostis racemosa – acacia tortilis variant (1.2.2) by species group d (table 1) and the presence of the woody species acacia stuhlmannii, and a high abundance of the grass eragrostis trichophora (species group k, table 1). the tree cover of this variant is between 20 % and 35 %, and the height of the trees alternating between 2.2 m to 4.0 m. in some isolated places there is no tree cover at all. the shrub layer reaches an average height of 1.3 m and the crown cover varies between 5 % and 20 %. the herbaceous layer covers between 60 % and 90% of the variant area and reaches an average height of 0.5 m. 1.2.2 eragrostis racemosa-acacia tortilis variant koedoe 46/2 (2003) 56 issn 0075-6458 gotze.qxd 2005/12/09 10:53 page 56 this thicket variant of the indigastrum costatum subsp. maculata – setaria verticillata sub-community forms a boundary between the open woodlands of the aristida congesta – leucas sexdentata variant (1.2.1) and the other riverine forest communities and sub-communities. the valsrivier and oakleaf soil forms found in this variant have a high clay content and appear to be less dry compared to the open woodland communities. the species of species group e (table 1) are diagnostic to this variant, which include the grass species eragrostis racemosa and the forb ctenolepis cerasiformis. a species with high abundance that further differentiate this variant from the aristida congesta-leucas sexdentata variant (1.2.1) is the woody species acacia tortilis (species group k, table 1). the vegetation cover of this variant is markedly higher than that of the aristida congesta-leucas sexdentata variant (1.2.1). salvadora australis and acacia tortilis (species group k, table 3) dominate the tree layer. the tree cover is 40 % on average and it reaches heights of between 3.5 m and > 8 m. the shrub layer has an average height of 1.4 m with an average crown cover of 20 %. the herbaceous layer has an average crown cover of 82 % and reaches an average height of 0.5 m. 1.3 combretum imberbe-abutilon ramosum sub-community this sub-community is situated in the riverine forest and in some cases on the fringes of the acacia xanthophloea diplachne fusca community (4) on the floodplain of this land type. soil forms include valsrivier, oakleaf and arcadia. in most cases the aspect is towards the middle of the adjacent water system, the slope is usually not to steep (2 -5º) and the soil surface is normally free of rocks. tall combretum imberbe (species group g, table 1), lonchocarpus capassa (species group p, table 1) and to a lesser extent, acacia tortilis (species group k, table 1) trees and a high abundance of the shrub abutilon ramosum (species group h, table 1) and the grass urochloa mosambicensis (species group k, table 1) dominate the vegetation in this sub-community. diagnostic species of the combretum imberbe-abutilon ramosum sub-community is found in species group g (table 1), which include the tree species combretum imberbe and the herbaceous solanum species. the absence of the species that are found in species group f (table 1) differentiates this sub-community from the other two in the salvadora australiscucumis zeyheri community (1). species with high cover abundance values that further differentiate this sub-community from the colophospermum mopane-eragrostis trichophora sub-community (1.1) and the indigastrum costatum subsp. maculatasetaria verticillata sub-community (1.2) are lycium cinereum (species group h, table 1) in the woody component and the shrub abutilon austro-africanum (species group k, table 1). the grasses echinochloa colona (species group h, table 1) and panicum maximum (species group p, table 1) and the forbs cucumis metuliferus and dicliptera eenii (species group h, table 1) also differentiate this sub-community. trees with an average height of more than 7 m and a crown cover of approximately 50 % dominate the vegetation of this subcommunity. the shrub layer reaches an average height of 1.4 m and the shrub cover varies between 5 % and 30 %. the herbaceous layer covers between 30 % and 70 % of the sub-community with an average height of 0.4 m. 2. hyphaene petersiana acacia tortilis community this community occurs on the flood plains; adjacent to the dense forest belt that occurs on the banks of the main rivers in the area. soils of the oakleaf form is the main soil type found in this community, with arcadia and valsrivier soils occurring on a more sporadic basis. the aspect and extremely gradual slope are mostly in a northerly direction as this is the general flow direction of the limpopo river in respect to this community. issn 0075-6458 57 koedoe 46/2 (2003) gotze.qxd 2005/12/09 10:53 page 57 rockiness of the soil surface is 1–6 % and the rocks and/or pebbles are often of a calcareous origin. this area is frequented by elephants and as a result is heavily utilised. the diagnostic plant species of this community are found in species group i (table 1) and include the woody species hyphaene petersiana and the forb trianthema triquetra. other species with high cover abundance values are acacia tortilis, salvadora australis (species group k, table 1) and grewia bicolor (species group o, table 1) from the woody component as well as the grasses enneapogon cenchroides, urochloa mosambicensis (species group k, table 1) and setaria verticillata (species group p, table 1). the tree layer of this community covers 10–60 % of the area. the height of the tree layer varies between 3 m and 9 m. the height of the shrub layer varies between 1.2 m and 2.0 m and the crown cover between 10 % and 60 %. grasses dominate the herbaceous layer. this layer has an average height of 0.5 m and a crown cover of between 37 % and 82 %. this community is divided into two subcommunities based on differences in vegetation structure. in the ximenia americana flueggea virosa sub-community (2.1) the shrub layer is dominant and in the cordia monoica sub-community (2.2) the tree layer is dominant. 2.1 ximenia americana-flueggea virosa sub-community this sub-community covers the largest part of the hyphaene petersiana acacia tortilis community (2). the dominant soil forms are oakleaf and valsrivier with high silt content and a small number of the arcadia soil form, which is a duplex soil with high clay content. diagnostic species of this sub-community are found in species group j (table 1) and include the woody species flueggea virosa subsp. virosa and ximenia americana. the dominant species are the tree species hyphaene petersiana (species group i, table 1) and acacia tortilis (species group k, table 1) as well as the grass species urochloa mosambicensis (species group k, table 1). other species distinguishing this sub-community from the cordia monoica sub-community (2.2) is the woody acacia nigrescens (species group j, table 1), the grass panicum maximum (species group p, table 1) and the annual forb tribulus zeyheri subsp. zeyheri (species group j, table 1). the tree layer has an average height of 5 m (in some cases as high as 9 m) and the crown cover varies between 10 % and 55 %. the shrub layer on the other hand enjoys a crown cover of between 36 % and 63 % and an average height of 1.7 m. the herbaceous layer has an average height of 0.5 m and covers 38 % to 80 % of the sub-community area. 2.2 cordia monoica sub-community the habitat of this sub-community is in many ways similar to that of the ximenia americana flueggea virosa sub-community (2.1). the only difference being that this sub-community has established under drier conditions and mainly occurs on somewhat higher lying areas further away from the limpopo river. the soils are also of the oakleaf and valsrivier forms with less occurrences of the arcadia soil form. there are no diagnostic species, which define this sub-community. the dominant species are acacia tortilis and urochloa mosambicensis (species group k, table 1) with hyphaene petersiana (species group i, table 1) not as dominant as in the ximenia americana flueggea virosa sub-community (3.1). species that further differentiate this sub-community are the woody species cordia monoica (species group i, table 1) and the absence of the tree species lonchocarpus capassa (species group p, table 1) and the grasses panicum deustum (species group o, table 1) and p. maximum (species group p, table 1). in the cordia monoica sub-community (2.2) the tree layer dominates the shrub layer. the tree layer reaches an average height of 5 m koedoe 46/2 (2003) 58 issn 0075-6458 gotze.qxd 2005/12/09 10:53 page 58 and a crown cover of between 30 % and 60 % while the crown cover of the shrub layer varies between 10 % and 45 % and reach an average height of 1.6 m. the herbaceous layer covers between 37 % and 82 % of the sub-community and has an average height of 0.4 m. 3. croton megalobotrys-combretum microphyllum community this riverine community is mainly situated on the banks of the limpopo river and to a lesser extent on the floodplain. it is characterised by tall trees and sometimes a dense undergrowth of shrubs, climbers and various other forbs and grasses. the utilisation by the tuli elephants is of a big, but not totally destructive measure. the elephants trample the area as they frequent it, mainly for shelter at night. the soils of this unit consist of oakleaf and valsrivier soil forms. the aspect is once more chiefly in a northerly direction and the slope varies between gradual slopes (1-5 º), leading from the flood-plain unit to the river, and short steep inclines (10-20 º) leading to the river itself. the diagnostic species is found in species group l (table 1) and in this case it is the woody species croton megalobotrys, which occurs in tree and shrub form. the woody species croton megalobotrys (species group l, table 1), the creeper combretum microphyllum (species group o, table 1) and the grass species panicum maximum (species group p, table 1) dominate the vegetation of this community. other species with high cover abundance values are the tree species acacia xanthophloea (species group r, table 1) and the grass species setaria verticillata (species group p, table 1) and panicum schinzii (species group o, table 1). the community is further characterised by the absence of the species of species groups i and k (table 1) as well as the herbaceous amaranthus sp. (species group r, table 1). the croton megalobotrys combretum microphyllum community has a high tree layer with the height of the trees varying between 7.8 m and 20.0 m and a crown cover of between 50 % and 90 %. the shrub layer has an average height of 1.6 m and covers between 18 % and 68 % of the total community area. the herbaceous layer reaches an average height of 0.7 m and a crown cover of between 8 % and 90 %. this community is divided into two subcommunities: 3.1 cenchrus ciliaris-faidherbia albida sub-community. this sub-community is found on the riverbank of the ia land type. the dominant soil form is valsrivier and to a lesser extent the oakleaf soil form. the habitat differs from the acacia schweinfurthii – maytenus senegalensis sub-community (3.2) because of it being more associated with low-lying areas that are frequently flooded during the annual flooding of the limpopo river, and its side branches. in some areas a layer of alluvial sand and flood-debris cover the soil surface only to be disturbed and washed away during the next flooding. the tree species faidherbia albida, the grass species cenchrus ciliaris and bothriochloa insculpta and the herbaceous species datura sp. and phragmites australis are diagnostic to this sub-community (species group m, table 1). tall, sometimes-dense trees dominate the vegetation of this sub-community, of which croton megalobotrys (species group l, table 1) is by far the most dominant. other species, which distinguish this sub-community from the acacia schweinfurthii – maytenus senegalensis sub-community (3.2) through their absence or low abundance, are the tree species grewia bicolor, grewia flavescens (species group o, table 1) and lonchocarpus capassa (species group p, table 1) and the grass species panicum deustum (species group o, table 1). the tree layer covers between 58 % and 90 % of this community and reaches a height of between 7.8 m and 17.0 m. the shrub layer has an average height of 1.5 m and a crown cover of between 18 % and 60 %. the issn 0075-6458 59 koedoe 46/2 (2003) gotze.qxd 2005/12/09 10:53 page 59 herbaceous layer covers between 10 % and 80 % and has an average height of 0.5 m. 3.2 acacia schweinfurthii maytenus senegalensis sub-community. the habitat of this sub-community is somewhat drier than that of the cenchrus ciliaris – faidherbia albida sub-community (3.1). it is situated on the floodplain and the riverbank of the ia land type on areas that are less frequently flooded. the soil forms are basically the same as in the faidherbia albida – acacia xanthophloea sub-community (3.1) only with the oakleaf soil form being more dominant than the valsrivier soil form. diagnostic species to the acacia schweinfurthii – maytenus senegalensis sub-community are found in species group n (table 1). this includes the rambling woody species acacia schweinfurthii, and the tree species xanthocercis zambesiaca, maytenus senegalensis and phyllanthus reticulatus as well as the creeper cocculus hirsutus. tall trees with undergrowth, which proves to be impenetrable in places, dominate the vegetation of this sub-community. croton megalobotrys (species group l, table 1) is less dominant in this sub-community and have lower cover abundance values than in the faidherbia albida – acacia xanthophloea sub-community (3.1). species, which further distinguish the acacia schweinfurthii – maytenus senegalensis sub-community (3.2) from the cenchrus ciliaris-faidherbia albida sub-community (3.1), are the tree species grewia bicolor, grewia flavescens (species group o, table 3) and lonchocarpus capassa (species group p, table 1). the tree layer of this sub-community covers between 50 % and 88 % of the area and the trees reach a height of between 8.0 m and 20.0 m. the shrub layer has more crown cover in this sub-community than is the case in the cenchrus ciliaris – faidherbia albida sub-community (3.1). the shrub layer covers between 30 % and 63 % of the sub-community and reaches an average height of 1.7 m. the herbaceous layer has an average height of 0.8 m and a crown cover varying between 8 % and 90 %. 4. diplachne fusca acacia xanthophloea community the diplachne fusca – acacia xanthophloea community commonly occurs on heavy clayey duplex soils in non-perennial pans and wetlands in the ia land type. soil types commonly found in these areas belong to the rensburg, arcadia, valsrivier and sepane soil forms. the soil surface, which is covered by cracks in the dry season, is mostly clear of rocks. aspect and slope is of no great significance in this community. it can however be stressed that most of the wetland/pan systems in the area drain towards the limpopo river. the major degradation problems of this community lie outside the current boundaries of the vhemeb-dongola national park, where large areas of these wetlands have been cleared for cultivation purposes. large dams have also been built for irrigation purposes and large-scale fish farming is also practiced in these dams. the diagnostic species to this community are found in species group q (table 1). they are the grasses diplachne fusca, dinebra retroflexa, echinochloa pyramidalis and the forbs corbichonia decumbens and abutilon sonneratianum. dominant species are the woody species acacia xanthophloea (species group r, table 1) and the grass diplachne fusca (species group q, table 1). the diplachne fusca – acacia xanthophloea community is further characterised by the absence of the species found in species groups o and p (table 1), which occur in most of the other communities in this land type. the tree layer has an average height of 8.2 m and a crown cover varying between 25 % and 35 %. the shrub layer is poorly developed with an average crown cover of only 3 % and an average height of 1.0 m. the herbaceous layer on the other hand is well developed with an average height of 1.3 m and a crown cover varying between 60 % and 90 %. koedoe 46/2 (2003) 60 issn 0075-6458 gotze.qxd 2005/12/09 10:53 page 60 ordination the distribution of the relevés of the ia land type indicates a distinct discontinuity between the four communities along ordination axis one of the scatter diagram (fig. 2). the gradient illustrated along the first ordination axis could be related to soil moisture and to a lesser extent clay content and soil depth . the salvadora australis – cucumis zeyheri community (1) is associated with flood plain areas on the outskirts of the riverine vegetation. on the scatter diagram (fig. 2) it is clear that the habitat of this community is virtually opposite to that of the acacia xanthophloea diplachne fusca community (4), with regard to the gradients stipulated along the first ordination axis. the acacia xanthophloea diplachne fusca community (4) is found in and around pans and other wetland systems in the vhembe-dongola national park on soils that are waterlogged for a long period during the year. the croton megalobotrys combretum microphyllum community (3) is found on the very edge of the limpopo river and its tributaries. in these parts the water table should certainly be higher than is the case further away from the river in the salvadora australis – cucumis zeyheri community (1) and the hyphaene petersiana – acacia tortilis community (2). this latter community is found on an area between the drier salvadora australis – cucumis zeyheri community (1) and the wetter croton megalobotrys combretum microphyllum community (3). concluding remarks some of the vegetation types described in the ia land type are similar to plant communities previously described in other studies. the vegetation and habitat of the salvadora australis – cucumis zeyheri community (1) are similar to the salvadora australis veld described in a vegetation report of the area by the former south african defence force (1986b), but different communities of riverine vegetation have not been described (south african defence force 1986 a,b). the hyphaene petersiana –acacia tortilis community (2) and the croton megalobotryscombretum microphyllum community (3) thus occurs in the same vegetation unit, issn 0075-6458 61 koedoe 46/2 (2003) fig. 2. the relative positions of the communities and sub-communities along the first two axes of the ordination of the relevés sampled in the vhembe-dongola national park on the ia land type. gotze.qxd 2005/12/09 10:53 page 61 which is referred to as riverine vegetation in the vegetation report of the south african defence force (1986a,b). an eragrostis vlei grass vegetation unit of which the habitat and vegetation compares well with the acacia xanthophloea diplachne fusca community (4) have also been described in the studies completed by the south african defence force (1986 a,b) in the area proposed for the vhembe-dongola national park. the riverine vegetation is more accurately described in the study of timberlake & mapaure (1999). the acacia/faidherbia woodland type described by timberlake & mapaure (1999) is further sub-divided into four sub-types namely the faidherbia albida woodland, the xanthocercis-schotia woodland, the acacia xanthophloea woodland and the acacia tortilis woodland. the faidherbia albida woodland shows similarities with the cenchrus ciliaris-faidherbia albida sub-community (3.1) and the xanthocercis-scotia woodland with the acacia schweinfurthii – maytenus senegalensis sub-community (3.2), both are sub-communities of the croton megalobotrys-combretum microphyllum community (3). in a phytosociological synthesis of the mopane veld, du plessis (2001) described the croton megalobotrys-colophospermum mopane vegetation type. this vegetation type and the croton megalobotrys-combretum microphyllum community (3) described in the current study show similarities with respect to habitat and floristic composition. the hyphaene petersiana–acacia tortilis community (2) is comparable with the hyphaene shrubland and to a lesser extent to the acacia tortilis woodland described by timberlake & mapaure (1999). the dry pan and wetland areas in which the acacia xanthophloea diplachne fusca community (4) occurs was not specifically described by timberlake & mapaure (1999). it does, however, to a certain degree, resemble the acacia xanthophloea woodland described by these researchers. the habitats described in different vegetation types by timberlake & mapaure (1999) and the south african defence force (1986 a,b) show similarities with the habitat descriptions of the different plant communities of the ia land type in this study. as a result of land clearing by intensive irrigation farming practices, as well as over-utilisation by small and large livestock and game (especially elephants), most communities of the ia land type are degraded. overutilisation and destruction caused by migratory tuli-elephants, from neighbouring botswana, is a large cause for concern. the south african national parks and the management of the vhembe-dongola national park consider this in a serious light and a number of monitoring sites have been established. another issue of concern observed in the ia land-type was the damage caused by elephants to large trees through the stripping of bark which will eventually lead to the death of these trees. the trees mostly affected include acacia xanthophloea, faidherbia albida and ficus sycomorus. plant communities described in this study and in other land types in the vhembe-dongola national park (götze 2002) should eventually be used in the compilation of a vegetation map of the park that can be used in the delineation of management units and the establishment of management plans. acknowledgements south african national parks and the peace parks foundation are thanked for technical and financial assistance and the south african defence force for providing soil and vegetation maps of the area. this study forms part of a msc.-dissertation at the potchefstroom university for christian higher education. references acocks, j.p.h. 1988. veld types of south africa, 3rd ed. memoirs of the botanical survey of south africa 57: 1-146. arnold, t.h. & b.c. de wet. 1993. plants of southern africa: names and distribution. memoirs of the botanical survey of south africa 62: 1-825. koedoe 46/2 (2003) 62 issn 0075-6458 gotze.qxd 2005/12/09 10:53 page 62 berry, m.p.s. 1994. visit to venetia by dr. john hanks of the south african nature foundation. unpublished report, de beers consolidated mines ltd. bezuidenhout, h. 1993. syntaxonomy and synecology of western transvaal grasslands. ph.d. thesis, university of pretoria, pretoria. bezuidenhout, h. 1996. the major vegetation communities of the augrabies falls national park, northern cape. 1. the southern section. koedoe 39(2): 7-24. bezuidenhout, h., h.c. biggs & g.j. bredenkamp. 1996. a process supported by the utility bbpc for analyzing braun-blanquet data on a personal computer. koedoe 39(1): 107-112. bredenkamp, g.j. & h. bezuidenhout. 1995. a proposed procedure for the analysis of large data sets in the classification of south african grasslands. koedoe 38(1): 33 – 39. bredenkamp, g.j. & g.k. theron. 1976. vegetation units for management of grasslands of the suikerbosrand nature reserve. south african journal of wildlife research 6: 113 – 122. bredenkamp, g.j. & g.k. theron. 1978. a synecological account of the suikerbosrand nature reserve 1. the phytosociology of the witwatersrand geological system. bothalia 12: 513-529. bredenkamp, g.j., m.s. deutschländer & g.k. theron. 1993. a phytosociological analysis of the albizio harveyi eucleetum divinorum from sodic bottom land clay soils of the manyeleti game reserve, gazankulu, south africa. south african journal of botany 59(1): 57-64. brown, l.r. 1997. a plant ecological study and wildlife management plan of the borakalalo nature reserve, north-west province. ph.d. thesis, university of pretoria, pretoria. carruthers, j. 1992. the dongola wild life sanctuary: ‘psychological blunder, economic folly and political monstrosity’ or ‘more valuable than rubies and gold’? kleio 24: 82 – 100. cilliers, s.s. 1998. phytosociological studies of urban open spaces in potchefstroom, north west province, south africa. ph.d. thesis, potchefstroom university for che, potchefstroom. du plessis, f. 2001. a phytosociological synthesis of mopaneveld. m.sc. thesis, university of pretoria, pretoria. edwards, d., 1983. a broad-scale structural classification of vegetation for practical purposes. bothalia 14, 3 & 4: 705–712. fuls, e.r., 1993. vegetation ecology of the northern orange free state. ph.d. thesis. university of pretoria, pretoria. götze, a.r. 2002. the classification of plant communities and the evaluation of restoration technologies in different land-use areas in the vhembe-dongola national park, northern province, south africa. m.sc. dissertation. potchefstroom university for c.h.e., potchefstroom. hennekens, s.m. 1996a. turboveg: software package for input, processing and presentation of phytosociological data. university of lancaster: ibn-dlo. hennekens, s.m. 1996b. megatab: a visual editor for phytosociological tables. ulft: giesen. hill, m.o. 1979a. twinspan: a fortran program for arranging multivariate data in an ordered twoway table by classification of individuals and attributes. ithaca, new york: cornell university. hill, m.o. 1979b. decorana: a fortran program for detrended correspondence analysis and reciprocal averaging. ithaca, new york: cornell university. land type survey staff. 2000. information on the land types of the map 2228-alldays. pretoria: directorate of agricultural information. mucina, l. & e. van der maarel. 1989. twenty years of numerical syntaxonomy. vegetatio 81: 1 – 15. mueller-dombois, d. & h. ellenberg. 1974. aims and methods of vegetation ecology. new york: wiley. naiman, r.j. & h. décamps. 1990. aquatic-terrestrial ecotones: summary and recommendations. pp. 295-301. in: naiman, r.j. & h. décamps (eds.). the ecology and management of aquaticterrestrial ecotones. paris: unesco. naiman, r.j., h. décamps & m. pollock. 1993. the role of riparian corridors in maintaining regional biodiversity. ecological applications 3(2): 209 – 212. o’connor, t.g. 1991. the vegetation types of the limpopo-venetia nature reserve. unpublished report, de beers consolidated mines, pretoria. robinson, g.a. 1996. limpopo valley national park: towards transfrontier conservation in south africa. unpublished report, south african national parks. rogers, k.h., 1995. riparian wetlands. pp. 41-52. in: cowan, g.i. (ed.) wetlands of south africa. pretoria: dept. environmental affairs & tourism, s.a. (wetlands conservation program series). south african defence force. 1986a. vegetation of the map areas 2229aa, pontdrif and 2229ac, evangelina. unpublished report, south african defence force. south african defence force. 1986b. vegetation of the map areas 2229ab, mapungubwe and 2229ad, coila. unpublished report, south african defence force. soil classification work group. 1991. soil classification—a taxonomic system for south africa. 2nd edition. memoirs on the agricultural natural resources of south africa 15: 1-262. issn 0075-6458 63 koedoe 46/2 (2003) gotze.qxd 2005/12/09 10:53 page 63 timberlake, j. & i. mapaure. 1999. vegetation survey of the maramani/tuli area, southwest zimbabwe. unpublished report on the cesvi lowveld project. transvaal provincial administration. 1989. verslag van die interdepartementele werksgroep na haalbaarheid van ‘n limpopo-bewaringsgebied, desember 1989. unpublished report, transvaal provincial administration, directorate nature and environmental protection. van den heever, j.l. 1983. grensgebied – geologiese verslag. unpublished report, geological survey, pretoria. van rooyen, n. & g.j. bredenkamp. 1996. mopane bushveld. pp. 20-21. in: low, a.b. & a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: dept. environmental affairs & tourism. van rooyen, n., g.k. theron & n. grobbelaar. 1981. a floristic description and structural analysis of the plant communities of the punda milia-pafuri-wambia area in the kruger national park, republic of south africa. 1. the hygrophilous communities. south african journal of botany 47: 213-246. voigt, e.a. & i. plug. 1981. early iron-age herders of the limpopo valley. unpublished report, transvaal museum, pretoria. weather bureau. 1986. climate of south africa. climate statistics up to 1984. weather bureau report 40: 1 – 474. winterbach, r. 1998. a phytosociological synthesis of acacia tortilis communities in the northwestern savanna of south africa. m.sc. thesis, university of pretoria, pretoria. willcox, a.r., 1966. memorandum concerning a suggested nature reserve and archaeological reserve in the limpopo valley. unpublished report, national parks board. koedoe 46/2 (2003) 64 issn 0075-6458 gotze.qxd 2005/12/09 10:53 page 64 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <feff004200720075006700200064006900730073006500200069006e0064007300740069006c006c0069006e006700650072002000740069006c0020006100740020006f0070007200650074007400650020005000440046002d0064006f006b0075006d0065006e0074006500720020006d006500640020006800f8006a006500720065002000620069006c006c00650064006f0070006c00f80073006e0069006e0067002000740069006c0020007000720065002d00700072006500730073002d007500640073006b007200690076006e0069006e0067002000690020006800f8006a0020006b00760061006c0069007400650074002e0020005000440046002d0064006f006b0075006d0065006e007400650072006e00650020006b0061006e002000e50062006e006500730020006d006500640020004100630072006f0062006100740020006f0067002000520065006100640065007200200035002e00300020006f00670020006e0079006500720065002e00200044006900730073006500200069006e0064007300740069006c006c0069006e0067006500720020006b007200e600760065007200200069006e0074006500670072006500720069006e006700200061006600200073006b007200690066007400740079007000650072002e> /nld <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> /esp <feff0055007300650020006500730074006100730020006f007000630069006f006e006500730020007000610072006100200063007200650061007200200064006f00630075006d0065006e0074006f0073002000500044004600200063006f006e0020006d00610079006f00720020007200650073006f006c00750063006900f3006e00200064006500200069006d006100670065006e00200071007500650020007000650072006d006900740061006e0020006f006200740065006e0065007200200063006f007000690061007300200064006500200070007200650069006d0070007200650073006900f3006e0020006400650020006d00610079006f0072002000630061006c0069006400610064002e0020004c006f007300200064006f00630075006d0065006e0074006f00730020005000440046002000730065002000700075006500640065006e00200061006200720069007200200063006f006e0020004100630072006f00620061007400200079002000520065006100640065007200200035002e003000200079002000760065007200730069006f006e0065007300200070006f00730074006500720069006f007200650073002e0020004500730074006100200063006f006e0066006900670075007200610063006900f3006e0020007200650071007500690065007200650020006c006100200069006e0063007200750073007400610063006900f3006e0020006400650020006600750065006e007400650073002e> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! 171.pdf page 1 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 page 4 marais.qxd plant nematodes in south africa. 7. a check list of plant nematodes from the fynbos biome, with a description of helicotylenchus curatus sp. n. m. marais, e. van den berg, a. swart and l.r. tiedt marais, m., e. van den berg, a. swart and l.r. tiedt. 2004. plant nematodes in south africa. 7. a check list of plant nematodes from the fynbos biome, with a description of helicotylenchus curatus sp. n. koedoe 47(1): 67–78. pretoria. issn 0075-6458. plant nematodes recorded during surveys in the fynbos biome are listed and a new helicotylenchus species is described. helicotylenchus curatus sp. n. is characterised by stylet length (42-46 µm in females, 37-40 µm in males), presence of two rudimentary subdorsal and two rudimentary subventral lobes on the labial disc, first lip annulus divided into six sectors, presence of fasciculi and presence of males. nine families represented by 32 genera and 152 species were identified from the fynbos biome. the genera criconema, helicotylenchus, hemicycliophora, rotylenchus, scutellonema and xiphinema were found in more than 30 % of the localities, whereas caloosia, criconemoides, ditylenchus, geocenamus, hemicriconemoides, heterodera, hoplolaimus, longidorus, meloidogyne, mesocriconema, ogma, paralongidorus, paratrichodorus, paratylenchus, pratylenchoides, pratylenchus, rotylenchulus, trichodorus and tylenchorhynchus were found at fewer localities. the genera anguina, hirschmaniella, histotylenchus and zygotylenchus were each identified from a single locality. key words: biodiversity, fynbos, helicotylenchus, plant nematodes, new species, south africa, taxonomy. m. marais, e. van den berg, a. swart, nematology unit, biosystematics division, arcplant protection research institute, private bag x134, queenswood, 0121 republic of south africa; l.r. tiedt, laboratory for electron microscopy, north west university, potchefstroom campus, potchefstroom, 2520 republic of south africa. issn 0075-6458 67 koedoe 47/1 (2004) introduction in 1987, the south african plant-parasitic nematode survey (sappns) programme was initiated with the aim to make a comprehensive assessment of the nematode biodiversity resources of south africa. one of the objectives of the sappns is to compile an inventory of the plant-parasitic nematodes of south africa. although nematodes constitute an abundant and highly diverse group of invertebrates, very little is known about their diversity in fynbos. the checklist reflects on collecting done by the agricultural research council, western cape nature conservation board, rand afrikaans university and university of stellenbosch (heyns 1971; kleynhans et al. 1996; marais & swart 1999; van den berg et al. 2003; van den berg & tiedt 2000) during the past 30 years. the fynbos biome is one of the biomes defined by low & rebelo (1996). in south africa, vegetation structure and climate mostly define these ecological zones. the fynbos biome is considered by many to be synonymous with the cape floral kingdom, but this biome refers only to the two vegetation groups, viz., fynbos and renosterveld within the region (low & rebelo 1996). renosterveld is characterised by the dominance of members of the asteraceae, specifically one species elytropappus rhinocerotis (renosterbos), from which the vegetation type gets its name. although renosterbos is the characteristic dominant plant, other plants are also prominent. these plants are all shrubs, characterised by small, tough grey leaves (rebelo 1996a). the various fynbos vegetation types marais.qxd 2004/04/15 07:56 page 67 comprise most of the area of the fynbos biome. fynbos is characterised by the presence of three elements, viz., a component belonging to the cape reed family (restionaceae), an ericoid or heath component and a proteoid component. fynbos is also characterised by the presence of a number of endemic or near-endemic plant families, viz., bruniaceae (blacktips), geissolomaceae (guyalone), grubbiaceae (sillyberry), penaeaceae (brickleaf), retziaceae (buttbush), roridulaceae (dewstick) and stilbaceae (candlestick) (rebelo 1996b). helicotylenchus is a cosmopolitan genus with more than 200 species described (marais 2001). this genus is found in all the biomes of south africa. thirty-one species of helicotylenchus have been reported from south africa. this paper is the result of an ongoing investigation on the genus helicotylenchus (marais 1993, 1998, 2001; marais & buckley 1992; marais & quénéhervé 1996; marais & quénéhervé 1999; marais et al. 2000; van den berg & marais 1995; van den berg et al. 2003). material and methods soil samples were collected with a garden trowel, soil auger or spade to a depth of about 20-25 cm at each locality. samples were collected in unprotected areas such as farms and plantations but also in nature reserves, wilderness areas and national parks. nematodes were extracted from 250 cm3 of soil, killed in water, preserved in faa, taf or fpg and mounted in anhydrous glycerine (hooper & evans 1993; jenkins 1964; kleynhans 1997; koen & furstenberg 1970; netscher & seinhorst 1969; seinhorst 1959). to obtain females of meloidogyne for identification, subsamples of each soil sample, in which root-knot nematode juveniles were found, were planted to tomato seedlings (cv. uc82b or roma vf) in a greenhouse. after six weeks, females were extracted from the roots and mounted in anhydrous glycerine (kleynhans 1991). for scanning electron microscopy, fpg-preserved specimens were used after dehydration in increasing concentrations of acetone. following conventional critical-point drying and gold/palladium-coating (15 nm), specimens were viewed with a quanta 200 esem. results and discussion the classification of nematodes followed is based on maggenti et al. (1988) for tylenchina, hunt (1993) for aphelenchina and longidoridae and decraemer (1995) for trichodoriae. nine families represented by 32 genera and 152 species were identified from the fynbos biome (table 1). the genera criconema, helicotylenchus, scutellonema, rotylenchus and xiphinema were the most common and were found at 40 %, 35 %, 31 %, 30 % and 30 % of the localities, respectively. the predominance of records in the mountain fynbos vegetation type is a consequence of the number of samples collected in this vegetation type (table 1). mountain fynbos is also the vegetation type with the highest total area (27 461 km2) of the ten fynbos vegetation types (low & rebelo 1996). taxonomy helicotylenchus curatus sp. n. figs. 1 & 2 measurements holotype female: l = 933 µm; a = 26.4; c = 43.8; c' = 1.2; stylet length = 42 µm; m = 53 %; v = 62 %. paratype females (n = 9): l = 1040 ± 110.1 (839-1262) µm; a = 23.0 ± 2.9 (17.5–27.5); c = 44.1 ± 4.5 (38.1–52.0); c' = 1.0 ± 0.2 (0.8–1.2); stylet length = 44 ± 1.7 (42–46); m = 54 ± 1.4 (52–56) %; v = 61 ± 1.8 (58–64) %. paratype males (n = 5): l = 864 ± 69.6 (760–944) µm; a = 36.2 ± 3.4 (32.5–39.2); c = 30.0 ± 2.9 (27.0–34.5); c' = 1.8 ± 0.4 (1.4– 2.6); stylet length = 39 ± 1.4 (37–40) µm; m = 57 ± 3.5 (52–60) %; spicule length = 33 ± 0.9 (32–34) µm; gubernaculum length = 10 ± 0.7 (9–11) µm. female: habitus spiral. lip region 7 ± 0.5 (6–8) µm high and 10 ± 0.9 (10–13) µm wide, truncate with distinct labial disc, six to seven annuli, basal annuli bulging out. koedoe 47/1 (2004) 68 issn 0075-6458 marais.qxd 2004/04/15 07:56 page 68 issn 0075-6458 69 koedoe 47/1 (2004) table 1 plant nematodes found associated with the fynbos biome 1a 2 3 4 5 6 7 8 % incidenceb number of localities samples 1 12 16 28 159 2 1 7 anguinidae anguina agrostis (steinbuch, 1799) filip'ev, 1936 + <1 ditylenchus spp. + <1 ditylenchus africanus wendt, swart, vrain & webster, 1995 + <1 belonolaimidae tylenchorhynchus spp. + + + + + 15 tylenchorhynchus avaricus (kleynhans, 1975) fortuner & luc, 1987 + <1 tylenchorhynchus capitatus allen, 1955 + + 1 tylenchorhynchus clarus allen, 1955 + <1 tylenchorhynchus dewaelei kleynhans, 1992 + <1 tylenchorhynchus estherae kleynhans, 1992 + + 1 tylenchorhynchus indicus (siddiqi, 1960) fortuner & luc, 1987 + <1 tylenchorhynchus mashoodi siddiqi & basir, 1959 + <1 tylenchorhynchus namibiensis rashid & heyns, 1990 + <1 tylenchorhynchus phaseoli sethi & swarup, 1968 + <1 histotylenchus spp. + <1 geocenamus brevidens (allen, 1955) brzeski, 1991 + + 1 pratylenchidae hirschmaniella spp. + <1 pratylenchoides spp. + + <1 pratylenchus spp. + + + + + 11 pratylenchus crenatus loof, 1960 + <1 pratylenchus delattrei luc, 1958 + <1 pratylenchus neglectus (rensch, 1924) filip'ev & schuurmans stekhoven, 1941 + + 1 pratylenchus penetrans (cobb, 1917) filip'ev & schuurmans stekhoven, 1941 + + 3 pratylenchus pratensis (de man, 1880) filip'ev, 1936 + <1 pratylenchus vulnus allen & jensen, 1951 + <1 pratylenchus zeae graham, 1951 + 1 zygotylenchus guevarai (tobar jiménez, 1963) braun & loof, 1966 + <1 hoplolaimidae hoplolaimus spp. + 3 hoplolaimus capensis van den berg & heyns, 1970 + + + 4 hoplolaimus pararobustus (schuurmans stekhoven & teunissen, 1938) sher, 1963 + 1 rotylenchus spp. + + 5 rotylenchus alius van den berg, 1986c + + + 6 rotylenchus brevicaudatus colbran, 1962 + + + 8 rotylenchus catharinae van den berg & heyns, 1974 + + 1 rotylenchus caudaphasmidius sher, 1965 + 2 rotylenchus incultus sher, 1965 + + 3 rotylenchus karooensis van den berg, 1986 + 1 rotylenchus kenti van den berg, 1989 + + + 2 scutellonema minutus (sher, 1964) germani, baldwin & wu, 1986 + <1 marais.qxd 2004/04/15 07:56 page 69 koedoe 47/1 (2004) 70 issn 0075-6458 rotylenchus unisexus sher, 1965 + + + 6 rotylenchus usitatus van den berg & heyns, 1974 + <1 helicotylenchus spp. + + + + + 22 helicotylenchus californicus sher, 1966 + + 2 helicotylenchus cavenessi sher, 1966 + <1 helicotylenchus curatus sp. n. + <1 helicotylenchus digonicus perry in perry, darling & thorne, 1959 + + 2 helicotylenchus dihystera (cobb, 1893) sher, 1961 + + + 4 helicotylenchus exallus sher, 1966 + + 6 helicotylenchus hydrophilus sher, 1966 + <1 helicotylenchus minzi sher, 1966 + 4 helicotylenchus paraplatyurus siddiqi, 1972 + + 1 helicotylenchus pseudorobustus (steiner, 1914) golden, 1956 + 1 helicotylenchus serenus siddiqi, 1963 + + 1 scutellonema spp. + + + + 8 scutellonema bizanae van den berg & heyns, 1973 + + + + 13 scutellonema brachyurus (steiner, 1938) andrássy, 1958 + + + + 9 scutellonema nigermontanum van den berg, 1990 + <1 scutellonema tsitsikamense van den berg, 1976 + 1 scutellonema unum sher, 1964 + <1 rotylenchulus spp. + 1 rotylenchulus borealis loof & oostenbrink, 1962 + 2 rotylenchulus macrodoratus dasgupta, raski & sher, 1968 + 1 rotylenchulus parvus (williams, 1960) sher, 1961 + + + + 5 rotylenchulus reniformis linford & oliveira, 1940 + heteroderidae heterodera spp. + 1 heterodera trifolii goffart, 1932 + <1 meloidogyne spp. + + + 5 meloidogyne hapla chitwood, 1949 + + 3 meloidogyne incognita (kofoid & white, 1919) chitwood, 1949 + 3 meloidogyne javanica (treub, 1885) chitwood, 1949 + + + 3 criconematidae criconema spp. + + + + 13 criconema ananas (heyns, 1970) siddiqi, 1986 + <1 criconema corbetti (de grisse, 1967) raski & luc, 1985 + 3 criconema crassianulatum (de guiran, 1963) raski & luc, 1985) + 1 criconema duplicivestitum (andrássy, 1963) raski & luc, 1985 + 11 criconema indigenae van den berg & meyer, 1991c + 1 criconema mutabile (taylor, 1936) raski & luc, 1987 + + + + + 4 criconema proteae van den berg & meyer, 1991c + 1 criconema sanctifrancisci (van den berg & heyns, 1977) raski & luc, 1985 + + 11 criconema simplex marais & van den berg, 1996c + 1 table 1 (continued) 1a 2 3 4 5 6 7 8 % incidenceb number of localities samples 1 12 16 28 159 2 1 7 marais.qxd 2004/04/15 07:56 page 70 issn 0075-6458 71 koedoe 47/1 (2004) criconema sirgeli van den berg & meyer, 1987c + <1 criconemoides parvus raski, 1952 + + 2 hemicriconemoides spp. + <1 hemicriconemoides brachyurus (loos, 1949) chitwood & birchfield, 1957 + + + + 6 hemicriconemoides capensis van den berg, 1990 + <1 hemicriconemoides cedrusmontanus van den berg & meyer, 1991c + <1 mesocriconema spp. + + + + + + 12 mesocriconema ferniae (luc, 1959) loof & de grisse, 1989 + 3 mesocriconema obtusicaudatum (heyns, 1962) loof & de grisse, 1989 + + + 3 mesocriconema sphaerocephalum (taylor, 1936) loof & de grisse, 1989 + 1 mesocriconema thabaum van den berg, 1996c + <1 mesocriconema xenoplax (raski, 1952) loof & de grisse, 1989 + + + 1 ogma spp. + <1 ogma civellae civellae reay & davies, 1998 + + ogma decalineatum (chitwood, 1957) andrássy, 1979 + 1 ogma inornatum (van den berg, 1983) siddiqi, 1986c + <1 ogma rhombosquamatum (mehta & raski, 1971) andrássy, 1979 + + 1 ogma squamiferum (heyns, 1970) andrássy, 1979c + + 1 caloosia exigua van den berg, marais & tiedt, 2003c + caloosia peculiaris van den berg & meyer, 1991c + <1 hemicycliophora spp. + + + 9 hemicycliophora demani edward & rai, 1971 + 2 hemicycliophora epicharoides loof, 1968 + <1 hemicycliophora halophila yeates, 1967 + + 1 hemicycliophora labiata colbran, 1960 + + + 7 hemicycliophora natalensis loof & heyns, 1969 + + 1 hemicycliophora nullinca van den berg, 1987 + <1 hemicycliophora peca van den berg, 1987c + <1 hemicycliophora stiaani van den berg & tiedt, 1999c + <1 hemicycliophora typica de man, 1921 + 1 hemicycliophora wesca van den berg & meyer, 1987c + <1 tylenchulidae paratylenchus spp. + + <1 paratylenchus arculatus luc & de guiran, 1962 + <1 paratylenchus elachistus steiner, 1949 + <1 paratylenchus projectus jenkins, 1956 + + 1 paratylenchus vandenbrandei de grisse, 1962 + <1 trichodoridae trichodorus spp. + 5 trichodorus iuventus decraemer & marais, 2000c + <1 table 1 (continued) 1a 2 3 4 5 6 7 8 % incidenceb number of localities samples 1 12 16 28 159 2 1 7 marais.qxd 2004/04/15 07:56 page 71 koedoe 47/1 (2004) 72 issn 0075-6458 trichodorus philipi de waele & van mieghem, 1990c + 1 trichodorus vandenbergae de waele & kilian, 1992 + 2 paratrichodorus spp. + + 4 paratrichodorus lobatus (colbran. 1956) siddiqi, 1974 + + + 3 paratrichodorus minor (colbran. 1956) siddiqi, 1974 + + + 3 paratrichodorus porosus (allen, 1957) siddiqi, 1974 + <1 longidoridae paralongidorus spp. + 4 paralongidorus capensis (heyns, 1967) liebenberg, heyns & swart, 1993 + <1 paralongidorus christiani liebenberg, heyns & swart, 1993c + 1 paralongidorus costatus (jacobs & heyns, 1987) siddiqi, baujard & mounport, 1993 + <1 paralongidorus deborae (jacobs & heyns, 1982) luc & doucet, 1984 + <1 paralongidorus spasskii heyns, 1972 + 1 longidorus spp. + + 2 longidorus fursti heyns, coomans, hutsebaut & swart, 1987 + <1 longidorus jagerae heyns & swart, 1998 + <1 longidorus juvenilis dalmasso, 1969 + + 1 longidorus pisi edward, misra & singh, 1964 6 xiphinema spp. + + + + 8 xiphinema americanum cobb, 1913 + 1 xiphinema barbercheckae coomans & heyns, 1985 + + + + 3 xiphinema bolandium coomans & heyns, 1985c + + + 5 xiphinema capense coomans & heyns, 1985 + 2 xiphinema diffusum lamberti & bleve-zacheo, 1979 + + 1 xiphinema elongatum schuurmans stekhoven & teunissen, 1938 + + + 1 xiphinema hardingi joubert, kruger & heyns, 1988 + 1 xiphinema krugi lordello, 1955 + + 1 xiphinema mampara heyns, 1979 + <1 xiphinema meridianum heyns, 1971 + + + 4 xiphinema mluci heyns, 1976 + <1 xiphinema pachtaicum (tulaganov, 1938) kir'yanova, 1951 + + 2 xiphinema parvistilus heyns, 1971 + + 2 xiphinema ripogranum hutsebaut, heyns & coomans, 1988c + + + 6 xiphinema vanderlindei heyns, 1962 + + + 1 xiphinema variabile heyns, 1966 + <1 xiphinema vitis heyns, 1974 + + 1 xiphinema zulu heyns, 1965 + <1 a vegetation types of the fynbos biome according to low & rebelo (1996) (1 = escarpment mountain renosterveld, 2 = central mountain renosterveld, 3 = west coast renosterveld, 4 = south and south-west coast renosterveld, 5 = mountain fynbos, 6 = grassy fynbos, 7 = laterite fynbos, 9 = sand plain fynbos) b percentage incidence in total number of samples collected during surveys. c nematodes described from the fynbos biome. table 1 (continued) 1a 2 3 4 5 6 7 8 % incidenceb number of localities samples 1 12 16 28 159 2 1 7 marais.qxd 2004/04/15 07:56 page 72 issn 0075-6458 73 koedoe 47/1 (2004) fig. 1. helictylenchus curatus sp. n. female. (a) habitus. (b) en face view, lip region. (c) lateral view, anterior part of body. (d) lateral field. (e-f) posterior part of body. (scale bar: a = 100 µm, b-e = 4 µm). a b c d e f marais.qxd 2004/04/15 07:56 page 73 koedoe 47/1 (2004) 74 issn 0075-6458 labial disc rectangular in en face view, with two rudimentary subdorsal and two rudimentary subventral lobes. first lip annulus divided into six sectors, two lateral, two subventral and two subdorsal. outer margins of the well-developed labial framework extend 4 ± 0.2 (3–4) µm backward from basal plate. anterior cephalids three to four annuli posterior to basal plate, posterior cephalids 11 to 13 annuli posterior to anterior cephalids. stylet slender, stylet knobs 4 ± 0.5 (3–4) µm high and 7 ± 1.3 (6–10) µm wide; anterior faces rounded (14 %), flattened (29 %), indented (14 %) or flattened and slightly inclined backward (43 %). position of dorsal gland opening (dgo) 5 ± 1.7 (3–9) µm behind stylet knobs. median bulb oval, 16 ± 1.4 (14–18) µm long and 12 ± 1.7 (10–15) µm wide; valve 4 ± 0.4 (3–4) µm long and 3 µm wide. length of oesophagus 151 ± 13.0 (134–175) µm, with length to end of gland 172 ± 19.0 (154–212) µm. oesophagus with 22 ± 4.3 (16–28) µm long ventral and dorsal overlap. excretory pore 154 ± 11.0 (142–172) µm from front, i.e. at 15 ± 1.3 (14–17) % of body length. hemizonid two annuli long located one to three annuli anterior to excretory pore. hemizonion half an annulus long, located 10 annuli posterior to excretory pore (n = 1). fasciculi present. width of annulus at midbody 2 µm. body width at excretory pore 33 ± 4.7 (26–40) µm, at midbody 46 ± 8.2 (35–59) µm and at anus 24 ± 4.8 (18–29) µm. two branches of the reproductive system both functional, length of posterior branch 81 ± 16.6 (57–94) % of corresponding anterior branch length; anterior branch 276 ± 59.8 (165–334) µm and posterior branch 199 ± 38.5 (156–235) µm long. spermatheca set off, empty or filled with sperm. epiptygma folded into vagina. lateral field 10 ± 1.7 (8–13) µm wide; outer two lines areolated opposite oesophagus, crenate or incompletely areolated on rest of body; inner two lines end on tail in a u-shaped (80 %) or open m-shaped pattern (20 %). phasmids located from five annuli posterior to opposite anus. tail 24 ± 2.3 (20–28) µm long, asymmetrical, more curved dorsally, with rounded end, with 13 to 18 ventral annuli, some annuli irregular. male: habitus c-shaped (20 %) to spiral (80 %). lip region 6 ± 0.5 (5–6) µm high and 8 ± 0.4 (8–9) µm wide; with six to seven annuli, basal annuli bulging out. anterior cephalids three to four annuli posterior to basal ring, posterior cephalids 12 to 14 annuli posterior to anterior cephalids. stylet more slender than that of female; stylet knobs 2 ± 0.4 (1–2) µm high and 4 ± 0.8 (4–5) µm wide, anterior faces flattened (50 %) or indented (50 %). position of dgo 4 ± 0.6 (3–4) µm behind stylet knobs. median bulb oval, 15 ± 2.3 (13–19) µm long and 10 µm wide; valve 3 ± 0.3 (3–4) µm long and 3 ± 0.8 (2–4) µm wide. length of oesophagus 146 ± 9.3 (133–153) µm, with length to end of glands 169 ± 15.4 (147– 184) µm. oesophagus with 29 ± 8.9 (18– 39) µm long ventral or ventro-lateral overlap. excretory pore 129 ± 5.4 (121–136) µm from front, i.e. at 15 ± 0.7 (14–16) % of body length. hemizonid one to three annuli long, located two to three annuli anterior to excretory pore. hemizonion half to one annuli long, located 10 to 13 annuli posterior to excretory pore. fasciculi present. width of annuli at midbody 2 µm. body width at excretory pore 20 ± 1.8 (18–23) µm, at midbody 24 ± 3.3 (21–29) µm and at cloaca 17 ± 3.3 (15–24) µm. lateral field 6 ± 0.3 (6–7) µm wide; outer two lines areolated anterior to and opposite bursa, crenate or incompletely areolated on body. phasmids located posterior to cloaca. margin of bursa almost straight, flattened. tail 31 ± 4.9 (26–39) µm long, conical with rounded projection. diagnosis h. curatus sp. n. is characterised by stylet length (42–46 µm in females, 37–40 µm in males), presence of two rudimentary subdorsal and two rudimentary subventral lobes on the labial disc, first lip annulus divided into six sectors, presence of fasciculi and presence of males. relationships a conspicuous character of h. curatus sp. n. is the short dgo (5–9 µm) in females and (3–4 µm) in males. this short dgo fit into marais.qxd 2004/04/15 07:56 page 74 issn 0075-6458 75 koedoe 47/1 (2004) fig. 2. helictylenchus curatus sp. n. female. (a) oesophageal region. (d) lateral field and fasciculi. (e) vulval region. (f-h) posterior part of body. male. (b) oesophageal region. (c) posterior part of body. a b c d e f g h 20 µm marais.qxd 2004/04/15 07:56 page 75 the diagnosis for the genus where the dgo varies from 3 µm (h. labiatus in yeates & wouts 1992) to 23 µm (h. erythrinae in marais 2001). according to fortuner (1984), this character has little value for identification, therefor the dgo was not used as a differentiating character. the stylet length has the smallest coefficient of variation among the quantitative characters (fortuner 1979; fortuner et al. 1981). there is a group of species in helicotylenchus characterised by long bodies and stylets, the respective measurements ranging from 800 µm to 1300 µm and from 35 µm to 46 µm. nine species are included in this group: h. arliani khan, singh & lal, 1998, h. canalis sher, 1966, h. coomansi sharafati-ali & loof, 1975, h. dolichodoryphorus sher, 1966, h. kashmirensis fotedar & handoo, 1974, h. macrostylus marais & quénéhervé, 1996, h. orthosomaticus siddiqi, 1972, h. paracanalis sauer & winoto, 1975, h. rohtangus jairajpuri & baqri, 1973 and h. tunisiensis siddiqi, 1963 (jairajpuri & baqri 1973; fotedar & handoo 1974; kepenekci & ökten 1996; khan et al. 1998; marais 1998; marais & quénéhervé 1996; marais et al. 2003; sharafati-ali & loof 1975; sher 1966; siddiqi 1972; van den berg & kirby 1979). h. curatus sp. n. was compared with three of these species, where the stylet lengths are more than 40 µm i.e. h. canalis, h. coomansi and h. macrostylus. the new species can be separated from h. canalis in lip region shape (truncate with basal annuli bulging out vs flattened to rounded, not set off), labial disc characteristics (rectangular in en face view, with two rudimentary subdorsal and two rudimentary subventral lobes vs oval in en face view without any lobes), first lip annulus divided vs not divided, mean body length (1040 µm vs 859 µm), position of excretory pore from front (142–172 µm vs 121–143 µm), mean tail length (24 µm vs 20 µm), position of phasmids (opposite anus to five annuli posterior to anus vs two to seven annuli anterior to anus) and presence of males vs absence of males. females of the new species can be distinguished from h. coomansi females in lip region shape (truncate with basal annuli bulging out vs continuous), labial disc characteristics (with two rudimentary subdorsal and two rudimentary subventral lobes present vs without any lobes), number of lip annuli (six to seven vs four to five), presence vs absence of fasciculi, a-value (17.5–27.5 vs 35–40), stylet length (42–46 µm vs 39–42 µm), m-value (52–56 % vs 48–50 %), position of phasmids (opposite anus to five annuli posterior to anus vs five to eight annuli anterior to anus) and tail form (asymmetrical, more curved dorsally, with rounded end vs straight, tapering dorsally with unstriated ventral portion). the males differ in body length (760– 944 µm vs 1190–1260 µm), position of excretory pore from front (121–126 µm vs 161 µm) and tail length (26–39 µm vs 41 µm — calculated from paratype material.). the new species can be separated from h. macrostylus in presence of fasciculi vs absence, m-value (52–56 % vs 42–45 %), lip region shape (truncate with basal annuli bulging out vs continuous), labial disc characteristics (rectangular, with two rudimentary subdorsal and two rudimentary subventral lobes vs round with no lobes), first lip annulus divided vs not divided. type locality and habitat collected from cyclopia plicata kies at haarlem, langkloof, western cape province (33°44's, 23°20'e) in a mountain fynbos vegetation type by h. hugo. type material holotype female (slide number 36524), six female paratypes and seven male paratypes (slide 36525–36528) deposited in the national collection of nematodes, biosystematics division, arc-plant protection research institute, pretoria, south africa. etymology from the latin cura, meaning to care, in recognition of liezel scheepers for her invaluable assistance. koedoe 47/1 (2004) 76 issn 0075-6458 marais.qxd 2004/04/15 07:56 page 76 acknowledgements we thank n.h. buckley and e. van niekerk (arcppri) for technical assistance, the various farmers, foresters and officers in charge for their help in collecting the soil samples and p.a.a. loof (landbouwuniversiteit wageningen) for the loan of h. coomansi paratypes. references decraemer, w. 1995. the family trichodoridae: stubby root and virus vector nematodes. dorderecht: kluwer. fortuner, r. 1979. morphometrical variability in helicotylenchus steiner, 1945. i. the progeny of a single female. revue de nématologie 2: 179–202. fortuner, r. 1984. morphometrical variability in helicotylenchus steiner, 1945. 6: value of the characters used for specific identification. revue de nématologie 7: 245–264. fortuner, r., g. merny & c. roux. 1981. morphometrical variability in helicotylenchus steiner, 1945. 3: observations on african populations of helicotylenchus dihystera and considerations on related species. revue de nématologie 4: 235–260. fotedar, d.n. & z.a. handoo. 1974. two new species of helicotylenchus (hoplolaiminae: nematoda) from kashmir, india. journal of science kashmir university 2: 57–62. heyns, j. 1971. a guide to the plant and soil nematodes of south africa. cape town: balkema. hooper, d.j. & k. evans. 1993. extraction, identification and control of plant nematodes. pp. 1–59. in: evans, k., d.l. trudgill & j.m. webster (eds.). plant parasitic nematodes in temperate agriculture. wallingford: cab international. hunt, d.j. 1993. aphelenchida, longidoridae and trichodoridae: their systematics and bionomics. wallingford: cab international. jairajpuri, m.s. & q.h. baqri. 1973. nematodes of high altitudes in india. i. four new species of tylenchida. nematologica 19: 19–30. jenkins, w.r. 1964. a rapid centrifugal-flotation technique for separating nematodes from soil. plant disease reporter 48: 692. kepenekci, i. & m.e. ökten. 1996. beypazari (ankara) ilçesi'nde havuf ile münabeye giren domates ekiliú alanlarinda saptanan helicotylenchus (tylenchida, hoplolaimidae) cinsine bagli türler. türk entomoloji dergisi 20: 137–148. khan, e., m. singh & m. lal. 1998. four new species of tylenchids (nematoda: tylenchida) from nepal. international journal of nematology 8: 27–32. koen, h. & j.p. furstenberg. 1970. 'n beknopte laboratoriumhandleiding vir nematologie. pretoria: departement van landbou-tegniese dienste. (tegniese mededeling 92.) kleynhans, k.p.n. 1991. the root-knot nematodes of south africa. pretoria: department of agricultural development, south africa. (technical communications no. 195.) kleynhans, k.p.n. 1997. collecting and preserving nematodes. a manual for a safrinet course in practical nematology. arc-plant protection research institute. pretoria. kleynhans, k.p.n., e. van den berg, a. swart, m. marais & n.h. buckley. 1996. plant nematodes in south africa. pretoria: arc-plant protection research institute. (plant protection research institute handbook; no. 8.) low, a.b. & a.g. rebelo. 1996. vegetation of south africa, lesotho and swaziland. a companion to the vegetation map of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. maggenti, a.r., m. luc, d.j. raski, r. fortuner & e. geraert. 1988. a reappraisal of tylenchina (nemata). 11. list of generic and suprageneric taxa, with their junior synonyms. revue de nématologie 11: 177–188. marais, m. 1993. on some helicotylenchus steiner, 1945 from south africa. phytophylactica 25: 21–35. marais, m. 1998. some species of helicotylenchus steiner, 1945 from south africa (nematoda: hoplolaimidae). fundamental and applied nematology 21: 327–352. marais, m. 2001. a monograph of the genus helicotylenchus steiner, 1945 (nemata: hoplolaimidae). ph.d. agric. dissertation, university of stellenbosch, stellenbosch. marais, m. & n.h. buckley. 1992. external morphology of eight south african helicotylenchus species (hoplolaimidae: nemata). phytophylactica 24: 297–306. marais, m. & p. quénéhervé. 1996. helicotylenchus macrostylus n.sp. (hoplolaimidae: nemata) from french guiana with notes on three known species. nematropica 26: 39–45. marais, m. & p. quénéhervé. 1999. a new species of helicotylenchus from french guiana, with notes on two known species (nemata: hoplolaimidae). journal for nematode morphology and systematics 2: 81–88. marais, m., p. quénéhervé, l.r. tiedt & a.j. meyer. 2003. description of helicotylenchus marethae n.sp., with redescriptions of some helicotylenchus species (nemata: hoplolaimidae) from south africa and martinique. journal issn 0075-6458 77 koedoe 47/1 (2004) marais.qxd 2004/04/15 07:56 page 77 koedoe 47/1 (2004) 78 issn 0075-6458 of nematode morphology and systematics 5: 1–19. (2002 published in 2003) marais, m. & a. swart. 1999. plant nematodes in the lottering plantation, eastern cape province. south african forestry journal 186: 77–79. marais, m. & e. van den berg, p. quénéhervé & l.r. tiedt. 2000. description of helicotylenchus kermarreci n.sp., with notes on some helicotylenchus steiner, 1945 and a rotylenchus filip'ev, 1936 species (nemata: hoplolaimidae) from the guadeloupe islands, french west indies. journal for nematode morphology and systematics 2: 159–172. (1999 published in 2000). netscher, c. & j.w. seinhorst. 1969. propionic acid better than acetic acid for killing nematodes. nematologica 15: 286. rebelo, a.g. 1996a. renosterveld. pp. 63. in: low, a.b. & a.g. rebelo. vegetation of south africa, lesotho and swaziland. a companion to the vegetation map of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. rebelo, a.g. 1996b. fynbos. pp. 68–70. in: low, a.b. & a.g. rebelo. vegetation of south africa, lesotho and swaziland. a companion to the vegetation map of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. sauer, m. r. & r. winoto. 1975. the genus helicotylenchus steiner, 1945 in west malaysia. nematologica 21: 341–350. seinhorst, j.w. 1959. a rapid method for the transfer of nematodes from fixative to anhydrous glycerin. nematologica 4: 67–69. sharafati-ali, s. & p.a.a. loof. 1975. two new species of helicotylenchus steiner, 1945 (nematoda: hoplolaimidae). nematologica 21: 207–212. sher, s.a. 1966. revision of the hoplolaiminae (nematoda) vi. helicotylenchus steiner, 1945. nematologica 12: 1–56. siddiqi, m.r. 1972. on the genus helicotylenchus steiner, 1945 (nematoda: tylenchida), with descriptions of nine new species. nematologica 18: 74–91. van den berg, e. & m. f. kirby. 1979. some spiral nematodes from the fiji islands (hoplolaimidae: nematoda). phytophylactica 11: 99–109. van den berg, e. & m. marais. 1995. new species of tylenchina chitwood (nemata) from the amazonas province, brazil. african plant protection 1: 25–39. van den berg, e., m. marais, s. gaidashova & l.r. tiedt. 2003. hoplolaimidae filip’ev, 1934 (nemata) from rwandan banana fields. african plant protection 9: 31–42. van den berg, e., m. marais & l.r. tiedt. 2003. plant nematodes in south africa. 5. plant nematodes from the gouekrans area of the swartberg nature reserve, with descriptions of one new and one known hemicycliophorionae (nemata). african plant protection 9: 43–52. van den berg, e. & l.r. tiedt. 2000. hemicycliophora stiaani sp.n. with notes on three tylenchid (nemata) species from south africa. journal of nematode morphology and systematics 2: 137–148. (1999 published 2000). yeates, g.w. & w.m. wouts. 1992. helicotylenchus spp. (nematoda: tylenchida) from managed soils in new zealand. new zealand journal of zoology 19: 13–23. marais.qxd 2004/04/15 07:57 page 78 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 515 --26 october 2009 [final version].indd a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe original research a rticle #515 (page number not for citation purposes) 80 alien plant species list and distribution for camdeboo national park, eastern cape province, south africa abstract protected areas globally are threatened by the potential negative impacts that invasive alien plants pose, and camdeboo national park (cnp), south africa, is no exception. alien plants have been recorded in the cnp since 1981, before it was proclaimed a national park by south african national parks in 2005. this is the fi rst publication of a list of alien plants in and around the cnp. distribution maps of some of the fi rst recorded alien plant species are also presented and discussed. to date, 39 species of alien plants have been recorded, of which 13 are invasive and one is a transformer weed. the majority of alien plant species in the park are herbaceous (39%) and succulent (24%) species. the most widespread alien plant species in the cnp are atriplex infl ata (= a. lindleyi subsp. infl ata), salsola tragus (= s. australis) and cacti species, especially opuntia fi cus-indica. eradication and control measures that have been used for specifi c problematic alien plant species are described. conservation implications: this article represents the fi rst step in managing invasive alien plants and includes the collation of a species list and basic information on their distribution in and around the protected area. this is important for enabling effective monitoring of both new introductions and the distribution of species already present. we present the fi rst species list and distribution information for camdeboo national park. authors: mmoto l. masubelele1 llewellyn c. foxcroft1 suzanne j. milton2 affi liations: 1sanparks scientifi c services, skukuza, south africa 2fitzpatrick institute of african ornithology, university of cape town, south africa correspondence to: mmoto l. masubelele e-mail: mmotom@sanparks.org postal address: sanparks scientifi c services, private bag x 402, skukuza 1350, south africa keywords: alien plants; biological invasion; habitat; threats; weeds dates: received: 12 nov. 2008 accepted: 22 july 2009 published: 28 sept. 2009 how to cite this article: masubelele, m.l., foxcroft, l.c. & milton, s.j., 2009, ‘alien plant species list and distribution for camdeboo national park, eastern cape province, south africa’, koedoe 51(1), art. #515, 10 pages. doi: 10.4102/koedoe.v51i1.515 this article is available at: http://www.koedoe.co.za © 2009. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. vol. 51 no. 1 page 1 of 10 introduction protected areas globally are under threat from invasive alien plants (pauchard & alaback 2004), as they change habitats and threaten resources, ecosystem services and indigenous species. alien plants enter parks in various ways; they are brought into the parks by people, wildlife, wind, water and vehicles (national park service 1996). further, alien plant species with established populations on park boundaries frequently encroach into the parks. for these reasons, a number of south africa’s national parks, for example table mountain national park (alston & richardson 2006) and kruger national park (foxcroft 2007), have been shown to be threatened by invasive alien plants. the eastern cape province of south africa falls within the albany centre for endemism, which has the highest number of plant extinctions, mainly as a result of agriculture, overgrazing, urbanisation and invasive alien plants (smith & wilson 2002). of the 4.7% of formally protected land in the eastern cape, 2.1% is located in the camdeboo municipality (smith & wilson 2002). although there are major data defi ciencies, invasive alien plants are believed to cover between 0.15% and 0.79% of the eastern cape surface area (csir 2004). climate change and invasive alien plants are regarded as the major threats to camdeboo national park (cnp), the largest protected area within the albany centre of endemism (camdeboo national park 2006). very few records of alien plants existed prior to 1981, and it is thus important to document alien plant introductions into the park to enable park management to prevent or mitigate the impact of alien species. of serious concern to cnp management is the fact that it is surrounded by a variety of land use types, including suburban areas, garbage disposal sites, quarries, and agricultural areas, all of which are likely sources of propagules of invasive alien plant species. moreover, a river runs from the town of graaffreinet through the cnp, and the park includes an artifi cial impoundment (nqweba dam). these water bodies and associated disturbances are also likely to transport and promote alien plant invasion. a species list provides a baseline for monitoring and managing alien plants that may threaten conservation management. species checklists are used by biologists to keep records of the numbers, types and categories of species groups in a defi ned area, and are a starting point towards effective management of either problematic indigenous or alien species (foxcroft et al. 2003) in a specifi c habitat, area, region, ecosystem, biome or country. the aims of this article are to present a species list of alien plants for the cnp, together with distribution maps for 15 alien plant species recorded between 1981 and 2008, and to examine changes in species distribution patterns over this period. methods study site cnp surrounds the town of graaff-reinet (figure 1), located in the camdeboo municipality of the eastern cape province of south africa. the cnp lies at the foothills of the sneeuberg range, with a small section of low-lying plains included within the boundaries, and ranges topographically between 740 and 1480 metres above sea level. the climate is semi-arid, with 32% of the average annual rainfall of 336 mm occurring during the hottest months of the year (february–april). the cnp also experiences snow, fog and frost, with a maximum summer temperature of 43ºc and a minimum temperature of –3 ºc in winter. the hydrology of cnp is determined by its location at the edge of the great escarpment, which has six seasonal rivers (sundays, gats, melk, camdeboo, pienaars and erasmuskloof rivers) draining into the nqweba dam in the central area of the park. koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research masubelele, foxcroft & milton a rt ic le # 51 5 (page number not for citation purposes) 81 vol. 51 no. 1 page 2 of 10 proclaimed in 2005, the cnp is one of 22 protected areas proclaimed under the management of south african national parks. however, it was first proclaimed as a provincial reserve (karoo nature reserve) in 1979. prior to 1979, the cnp area was used as town commonage with tenants grazing their livestock, thus contributing to overgrazing and erosion of some areas (burdett 1995). prior to colonial settlement, the land was used by the early, mid and late stone age people, as well as the khoisan hunters and herders in the late stone age. the inqua tribe occupied the park area in the mid-1600s with their cattle and fat-tailed sheep. white farmers settled on the camdeboo plains and sneeuberg in 1770 and introduced merino sheep and angora goats, as well as alien plants (burdett 1995). in the ensuing years, overgrazing and the effects of alien plants resulted in soil erosion and an increase in woody species and unpalatable plants (cnp 2006). the geological systems of cnp consist of very thick layers of near horizontal strata of sedimentary rocks, with the largest parts covered with alluvium, gravel, sand, mud and wash stone of recent origin. these tertiary to quaternary deposits are an important feature influencing the vegetation of the nama karoo biome, and they comprise the growth medium for many dwarf shrubs in the region (lovegrove 1993). the soils are generally calcareous duplex forms of secondary nature, having been deposited as alluvium on the impermeable sandstone. they are subject to sheet and gully erosion, which is aggravated by a reduction in vegetation cover. the vegetation of the cnp falls into three biomes, namely the albany thicket, grassland and nama karoo (mucina & rutherford 2006; palmer 1989). there is also azonal alluvial vegetation around the nqweba dam. the vegetation has been divided into three distinct physiognomic classes of vegetation, figure 1 location and extent of the cnp and graaff-reinet in the eastern cape province of south africa note: the insert places cnp in relation to the eastern cape province (shaded), and the rest of south africa. the n9 and r63 are the main roads that run through graaff-reinet and cnp alien plant species list and distribution for camdeboo national park, eastern cape province, south africa original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #515 (page number not for citation purposes) 82vol. 51 no. 1 page 3 of 10 namely the shrubland, succulent thicket and dwarf shrubland (palmer 1989). the shrubland is dominant in sandstone uplands above 1300 meters elevation. grasslands separate the shrubs growing in moist conditions from the dwarf shrub vegetation in the drier areas. the succulent thicket is a distinctive vegetation class dominated by shrubs and succulents of subtropical affinity, while the dwarf shrubland is restricted to bottomlands which may be grassy, succulent or degraded, depending on the surface substrate, the frequency of precipitation and recent land use history. to date, 336 plant species from 71 families of flowering plants have been recorded (palmer 1989). the following families are important components of the region’s flora: asteraceae (daisy family with 55 species), poaceae (grass family with 36 species), liliaceae (lily family with 25 species) and crassulaceae (crassula family with 16 species). other life forms include 43 species of mammals, consisting of, amongst others, disease-free buffalo (syncerus caffer), kudu (tragelaphus strepsiceros), springbok (antidorcas marsupialis) and blesbuck (damaliscus dorcas phillipsi), and thirteen carnivore species (cnp 2006). there are also 225 species of birds. the herpetofauna includes five frog, five tortoise, 19 lizard and 10 snake species. there are also ten species of fish, including two alien species of angling value, namely, carp (cyprinus carpio) and barbell (clarias gariepinus). the invertebrate fauna is largely unrecorded, but there are brown locust (locustana pardalina) outbreaks, and karoo caterpillar (loxostege frutalis) and harvester termite (hodotermes mossambicus) eruptions. data collection the data used in this study originates primarily from two sources. the first is an unpublished alien plant species list that was compiled in 1981, containing 15 species recorded within the park boundaries. each of these species was plotted on a 1:50 000 map by the park manager and digitised in 2008. the data contained in the hand-plotted maps were digitised onto a 1:50 000 map to improve the visibility and quality of data, using arcview gis version 3.1. the second dataset is based on ranger patrols that took place between april and december 2008 inside the cnp (figure 1). routine patrols were conducted by field rangers, on foot and by vehicle, using airis picket pcs with customised cybertracker software (see foxcroft et al. 2009 for a discussion on the cybertracker programme). the collection of these and other environmental data by park rangers is ongoing. in addition, in may 2008 a general survey was carried out and all new species encountered were added to the species list for the cnp. presentation of the list families and species, respectively, are listed alphabetically within each of the major plant groupings of pteridophyta (ferns and fern allies), gymnospermae (gymnosperms) and angiospermae (monocotyledons and dicotyledons). the date of the first record of the species indicates the year recorded by park authorities. the list further provides information on the status of the weed and its current impact in the cnp, methods for control, and legal status in south africa. weed status is presented according to terminology described by pyšek et al. (2004), who defined and classified weed status as follows. casual alien plants (c) are those that may flourish and even reproduce occasionally outside cultivation in an area but eventually die out because they do not form self-replacing populations and rely on repeated introductions for their persistence. invaders (i) are a subset of naturalised plants that produce reproductive offspring, often in very large numbers, at considerable distances from the parent plants, and thus have the potential to spread over a large area. naturalised plants (n) are alien plants that sustain self-replacing populations for at least 10 years without direct intervention by people, or by recruitment from seed or ramets capable of independent growth. potential invaders (p) means alien plants with the potential to reach invader status. the habitat types include all habitats within the park boundaries, and rivers, urban and rural areas within a 1-km zone from the park boundary. almost all areas in the park have been exposed to agriculture, either through cultivation or livestock grazing, or both. the species observed and recorded in the cnp were found either in disturbed areas and roadsides (denoted by dr), villages and picnic areas in the cnp, urban and rural areas adjacent to the cnp (v), aquatic habitats (including nqweba dam, denoted by a), riparian or riverine areas in and adjacent to cnp (r), and terrestrial or dryland habitats (td). control mechanisms utilised in cnp include chemical and mechanical techniques; there are no records of biological control. mechanical clearing for the first, second and third time, respectively, is indicated by me1–me3. chemical control is indicated in a manner similar to mechanical control (ce1–ce3), with the active ingredient of the herbicide used provided in brackets. legal status refers to the declared status of the species in terms of the conservation of agricultural resources act (act 43 of 1983) regulations (as amended in 2001). species are divided into three categories. category 1 species are ‘declared weeds’, which are prohibited, and must be controlled or eradicated (dw1). category 2 species are ‘declared invaders’, which comprise mainly commercial species that must be grown under controlled conditions by permit holders (di2). category 3 species are ‘ornamental species’, which may no longer be planted, except with a special permit, and must be prevented from spreading (di3). for a full description of the regulations and details regarding individual alien plants see henderson (2001). results the first list compiled by the park manager contained 15 alien plant species, including one grass species (pennisetum clandestinum), four herbaceous species (argemone ochroleuca subsp. ochroleuca, cirsium vulgare, salsola tragus (= s. australis) [see 6 27 17 20 12 0 5 10 15 20 25 30 35 40 aquatic disturbed/ roadsides terrestrial/dryland riparian/riverine villages, picnic areas and urban areas nearest to cnp habitat type n u m b er o f s p ec ie s figure 2 number of alien plant species per habitat type recorded during the 2008 surveys (some species occur in more than one habitat type) figure 3 families of alien plant species represented in camdeboo national park ������������ � ���������� � �������� � ���������� � ��������� � ������� �� ������������� � ���������� �� ��������� �� �������������� � ������������� � ������� �� !�������� � !"������� # ������������ � $������� � ���������� # �������� �������%�&%�����%�����%�������%�����������%��%��������%'�������%���( original research masubelele, foxcroft & milton koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 51 5 (page number not for citation purposes) 83 vol. 51 no. 1 page 4 of 10 figure 4 distribution of alien plant species in camdeboo national park, compiled from data collected in 1981 alien plant species list and distribution for camdeboo national park, eastern cape province, south africa original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #515 (page number not for citation purposes) figure 5 distribution of alien plant species in the camdeboo national park, compiled from patrol data collected using cybertracker, april–december 2008 6 vol. 51 no. 1 page 5 of 10 84 original research masubelele, foxcroft & milton koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 51 5 (page number not for citation purposes) discussion on s. tragus by hrusa & gaskin 2008] and xanthium spinosum), three shrubs (nicotiana glauca, ricinus communis and solanum elaeagnifolium), three trees (eucalyptus camaldulensis, prosopis glandulosa var. torreyana and schinus molle) and four cacti: opuntia aurantiaca, opuntia ficus-indica, cylindropuntia tunicata (previously misidentified as o. fulgida or o. rosea) and cylindropuntia imbricata (= o. imbricata). although e. globulus was indicated in the early park records, this was most likely a misidentification, as the only eucalyptus species from the area has been identified as e. camaldulensis. during the 2008 surveys a total of 24 additional alien plant species were added to the original alien species list (table 1). when the alien plants on the list are ranked according to weed status, the majority of the species (17) are invasive (46%) and 15 are potential invaders (41%). two species are recorded as naturalised (5%), and the remaining three have a casual status (8%). therefore, over 80% of the alien plants recorded either are, or pose, a potential threat to the vegetation types and ecosystems around cnp. among the successful invaders in and around cnp, the following are prominent: cirsium vulgare, xanthium spinosum, opuntia aurantiaca, o. ficus-indica, cylindropuntia imbricata, c. tunicata and eucalyptus camaldulensis. tamarix ramosissima is an invader with the potential to transform the aquatic edge habitat in cnp (especially the nqweba dam shoreline). atriplex inflata and salsola tragus are also widespread invaders in terrestrial drylands and disturbed areas in large sections of cnp. this is possibly due to earlier ploughing in almost all areas in and around the park. most of the alien plants have invaded disturbed areas, including areas that were previously heavily grazed and ploughed, as well as cleared areas along roadsides (figure 2). riparian habitat is the next most vulnerable to invasion, with the terrestrial or dryland areas appearing to be least vulnerable. when the nqweba dam overflows it results in an increase in the presence of especially tamarix ramosissima and cirsium vulgare (charlotte vermeulen [wfw manager] pers. comm., 16 may 2008). solanum elaeagnifolium occurs in all habitat types, and xanthium spinosum, atriplex inflata, a. nummularia subsp. nummularia, salsola tragus, opuntia aurantiaca, o. ficus-indica, cylindropuntia imbricata and datura stramonium occur in four habitat types. the most highly invasive families of alien plants in the cnp (figure 3) are the cactaceae (21%), then asteraceae (10%), poaceae (10%), fabaceae (10%), chenopodiaceae (8%) and solanaceae (8%). of the cacti species, opuntia aurantiaca, o. ficusindica, cylindropuntia imbricata and c. tunicata are highly invasive within a 1-km zone from the park boundary and within the park. cylindropuntia tunicata and c. imbricata appear to be spreading from the areas where people live, into the disturbed areas and roadsides at the edge of the park. the most widespread invader in the cnp is opuntia ficus-indica. it was already widespread in the early distribution records obtained from 1981 (figure 4) and is still currently widespread (figure 5). some alien plant species that are also known to be widespread but have not been comprehensively mapped are atriplex inflata, salsola tragus and tamarix ramosissima. the current distribution data (figure 5) implies that some alien plant species have decreased in distribution when compared to the situation in 1981 (figure 4). this is most likely due to the fact that the data collected for 2008 might not be extensive enough to have covered the entire area of the park, which was handmapped in 1981. it might also be because some individual plants have been mechanically or chemically controlled, and thus they now occur in lower abundance. alien plant species that appear to have decreased in distribution are cirsium vulgare, eucalyptus camaldulensis, nicotiana glauca, cylindropuntia imbricata, solanum elaeagnifolium and xanthium spinosum (figures 4 and 5). of vol. 51 no. 1 page 6 of 10 figure 6b a selection of cacti at a cactus nursery in graaff-reinet figure 6a a selection of cacti at a cactus nursery in graaff-reinet figure 6d a cactus collection in an open space in graaff-reinet (photographs: l.c. foxcroft) figure 6c various cacti along the street bordering the nursery (pennisetum setaceum is also visible in the foreground) note larger figures are available on the online journal 85 alien plant species list and distribution for camdeboo national park, eastern cape province, south africa original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #515 (page number not for citation purposes) 86vol. 51 no. 1 page 7 of 10 table 1 alien plant species list of camdeboo national park order and families species common name date of first record status habitat control legal status order: pteridophyta family: azollaceae azolla filiculoides lam. red water fern 2001 i a r the biological control agent stenopelmus rufinasus, while not intentionally released into cnp, is most likely present dw 1 order: gymnospermae family: pinaceae pinus halepensis mill. aleppo pine 2000 c td none di 2 order: angiospermae: monocotyledonae family: agavaceae agave sisalana perrine sisal 2006 n v td r ce1 (msma) me1 di 2 family: poaceae arundo donax l. giant reed/spanish reed 2008 i r none dw 1 nassella trichotoma (nees) arechav. nassella tussock 2008 p dr td none dw 1 pennisetum clandestinum chiov. kikuyu grass 2006 i dr v td none proposed dw 2 pennisetum setaceum (forssk) chiov. fountain grass 2008 p dr r none dw 1 order: angiospermae: dicotyledonae family: anacardiaceae schinus molle l. pepper tree 1986 c dr v me 1 proposed di 3 family: asteraceae bidens pilosa l. common blackjack 2008 n dr v none na cirsium vulgare (sari) ten spear thistle/ scotch thistle 1986 i a r ce1 (imazapyr) dw 1 xanthium spinosum l. spiny cocklebur 1981 i a dr td r ce1 unknown dw 1 xanthium strumarium l. large cocklebur 1981 p dr none dw 1 family: cactaceae cereus jamacaru dc. queen of the night 2008 p dr unknown dw 1 cylindropuntia imbricata (haw.) knuth (= opuntia imbricata (haw.) dc. imbricate prickly pear 1981 i dr td r cei msma dw 1 cylindropuntia tunicata (lehm.) f.m. knuth thistle cholla 1986 i dr td v ce3 msma na echinopsis spachiana (lem.) friedrich & rowley torch cactus 2008 p dr r ce1 dw 1 opuntia aurantiaca lindl. jointed cactus 1981 i dr v td r ce3 msma dw 1 opuntia ficus-indica (l.) mill sweet prickly pear 1981 i dr v td r ce3 msma dw 1 opuntia humifusa (raf.) raf. large flowered or creeping prickly pear 2008 p td v cei msma dw 1 opuntia stricta (haw.) haw. sour prickly pear 2008 i dr td none dw 1 family: chenopodiaceae atriplex inflata f. muell. (= a. lindleyi subsp. inflata (f. muell.) p.g. wilson sponge-fruit salt bush 1986 i dr td ce1 (fluroxypyrmeptyl) di 3 atriplex nummularia lindl. subsp. nummularia old man salt bush 1989 p dr td v none di 2 salsola tragus l. (s. kali l. misapplied in sa) russian tumbleweed 1986 i dr td r none na family: euphorbiaceae ricinus communis l. castor oil plant 1986 i dr r none di 2 family: fabaceae gleditsia triacanthos l. honey locust 2008 p dr r me1 di 2 parkinsonia aculeata l. jerusalem thorn 2008 p dr r none na prosopis glandulosa torr. var. torreyana (l.d. benson) m.c. johnst. honey mesquite 1986 t dr me1 di 2 robinia pseudacacia l. black locust 2008 p dr r none di 2 family: meliaceae melia azedarach l. seringa, syringa, persian lilac 2006 c v r none di 3 family: myrtaceae eucalyptus camaldulensis dehn. red river gum 1986 i v r me1 di 2 family: papaveraceae argemone ochroleuca sweet subsp. ochroleuca (a. subfusiformis g.b. ownbey misapplied in sa) white-flowered mexican poppy 1986 i dr none dw 1 family: rosaceae pyracantha angustifolia (franch.) c.k. schneid. yellow firethorn 2008 p dr none di 3 family: salicaceae populus x canescens (aiton) sm. grey poplar 2008 p r me1 di 2 salix babylonica l. weeping willow 2008 p r none di 2 family: solanaceae datura stramonium l. common thorn apple 2008 p a dr td r none dw 1 original research masubelele, foxcroft & milton koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 51 5 (page number not for citation purposes) 87 vol. 51 no. 1 page 8 of 10 these species, cirsium vulgare, s. elaeagnifolium, cylindropuntia imbricata and x. spinosum have been chemically controlled while the other two have been mechanically removed. the chemicals used here include monosodium methanearsonate (msma) on cylindropuntia imbricata, and imazapyr on cirsium vulgare and s. elaeagnifolium. there was an apparent increase in the distribution of o. ficusindica, o. aurantiaca, c. tunicata and schinus molle (figures 4 and 5). schinus molle has increased in distribution although some individual species have been mechanically removed. the three cactus species (o. ficus-indica, o. aurantiaca, c. tunicata) are persistent invaders, as efforts to reduce these species have been unsuccessful, even though there have been more than two chemical controls and follow-up operations (table 1). the fact that different species of cactaceae utilise a variety of methods to invade (easily dispersed seeds and vegetative dispersal by ramets) makes them difficult to eradicate. alien plant species that were recorded in 1981 but were not found in the 2008 general survey data include argemone ochroleuca subsp. ochroleuca, pennisetum clandestinum, prosopis glandulosa var. torreyana, salsola tragus and ricinus communis. plants recorded only in 2008 include echinopsis spachiana and pennisetum setaceum, both of which have invaded the eastern section of the cnp, presumably having spread from the nearby urban area. it is anticipated that more alien plant species will be recorded at the edges of the park as further surveys are carried out. the rivers that flow through the town and into the park are a source of many woody and shrubby alien plant species, and present a substantial management challenge. discussion our knowledge on the current distribution of invasive alien plant species in the cnp is still relatively limited; the focus has mainly been on individual plant species and the distribution has not been frequently updated. this article reports on efforts that have been made to improve our knowledge of the extent of invasive alien species by assessing the distribution of 18 of the most important and persistent species. while this is a start, it should be recognised that monitoring and recording the distribution of alien plants should remain an ongoing process. this paper only provides a picture of where specific alien plant species are common; detailed structured maps at relevant spatial scales should be compiled in the near future. effective management of invasive plants is based on thorough knowledge of the species’ locations and distribution, modes and rates of spread, potential and known effects, and control methods (crimmins et al. 2008). an inventory of the invasive species, invasion processes and management history provides management with a valuable baseline. this updated list contains a total of 39 alien plant species, an increase of 24 species from the 15 initially recorded and mapped before the cnp was proclaimed. cactaceae as invaders in the karoo most of the important alien plant invaders of the karoo biomes are succulents (richardson et al. 1997). the unique shapes and sizes of cacti results in these species being planted widely in gardens by people. this is done mainly for ornamental and aesthetic reasons. there is a large cactus nursery in graaffreinet, on the edge of the sundays river (which runs from the town into the park), which sells many types, shapes and forms of cacti (see figures 6a–d). a number of cacti species appear to have been dumped into the adjacent river, which will promote invasion in the camdeboo municipality. many species spread through vegetative reproduction, and their fleshy fruits are consumed by a host of animals, which assists their distribution into natural areas. prickly pear (e.g. opuntia ficus-indica) seeds are often dispersed to perch sites by birds (particularly crows) and to riverbeds and woodlands by primates (humans, vervet monkeys and baboons) that eat the sweet, watery fruit (milton & dean 1998; richardson et al. 2000). cylindropuntia imbricata and o. aurantiaca also appear to be dispersed by similar agents. birds are major contributors to the spread of cacti to shaded sites below perches on trees and cliffs (milton et al. 2007). opuntia ficus-indica can be seen at the valley of desolation hanging from cliffs as well as under acacia karroo in drainage lines. opuntia aurantiaca is common only under a. karroo and pentzia incana. however, as acacia karroo and pentzia incana are the most common indigenous plant species in the karoo (palmer 1989), these cacti species pose a risk to the species richness and diversity in the acacia karroo-pentzia incana plant communities if nothing is done to reverse the situation. the four major cacti invaders are opuntia aurantiaca, o. ficusindica, cylindropuntia imbricata and c. tunicata. the worst invader in cnp and the surrounding municipality remains o. ficus-indica. this species was originally introduced in the early 1700s as fodder for sheep and cattle. it survived and spread effectively to become not only a problem plant in the semi-arid karoo and savanna, but to cause one of the worst agricultural catastrophes in the history of south africa (annecke & moran 1978; moran & annecke 1979). the extent of the problem can be seen in old photographs, and its invasion of protected areas in the savanna (foxcroft 2007; macdonald & frame 1988; wells et al. 1986; zimmermann and moran 1982; milton et al. 2007) and the karoo (dean & milton 2000; milton & dean 1998) is discussed in various places in the literature. records for cnp indicate that o. ficus-indica has been treated mechanically and chemically using msma until it was visibly reduced. ongoing follow-up treatment is important for the successful eradication or more likely, maintenance at low levels, of the species. the fact that follow-up control operations were not carried out continuously has resulted in the species returning in large numbers and still occurring as the most widespread species in the area. although not specifically released in the cnp, the biological control agent dactylopius opuntiae, a cochineal insect, has substantially reduced the density of o. ficus-indica in south order and families species common name date of first record status habitat control legal status nicotiana glauca graham. wild tobacco 1986 p dr me1 dw 1 solanum elaeagnifolium cav. silver-leaf bitter apple 1986 i a dr td ce1 (imazapyr) dw 1 family: tamaricaceae tamarix ramosissima ledeb. pink tamarisk 2006 i a r me1 ce1 (imazapyr) dw 1 codes used: date: first year of record in the park status: as in cnp currently; c=casual; i=invader; n=naturalized; p=potential invader habitat type: includes rivers, urban and rural areas adjacent to cnp; a=aquatic, includes nqweba dam; dr=disturbed area/roadside; r=riparian/riverine; td=terrestrial/dryland; v=urban areas around graaff-reinet adjacent to cnp control: mechanical stage of clearing from 1st to 3rd=me1 until me3; chemical stage of application from 1st to 3rd with the name of the chemical in bracket=ce1 until ce3 legal status according to cara (act 43 of 1983, as amended in 2001): dw 1=declared weed category 1; di 2=declared invader category 2; di 3=declared invader category 3 table 1 (continued...) alien plant species list of the camdeboo national park 7 alien plant species list and distribution for camdeboo national park, eastern cape province, south africa original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #515 (page number not for citation purposes) 88vol. 51 no. 1 page 9 of 10 africa (moran and zimmermann 1991; zimmermann et al. 1986). the other biological control agent stenopelmus rufinasus, also not specifically released into cnp, is most likely controlling the red water fern (azolla filiculoides) infestations in dams, ponds and rivers on the camdeboo municipality. invasive trees, shrubs and herbs alien plant species recorded in the karoo nature reserve (now cnp) by palmer (1989) include tamarix ramosissima, salsola tragus (s. kali misapplied), atriplex inflata (= a. lindleyi subsp. inflata), and argemone ochroleuca subsp. ochroleuca (a. mexicana misapplied by palmer). tamarix ramosissima is a major invader of rivers throughout the world (loope et al. 1988), and has invaded the shores of nqweba dam in cnp. the rivers in the camdeboo municipality have further been invaded by a range of alien plant species, with eucalyptus camaldulensis already producing large numbers of seedlings and saplings. the other alien trees invading semi-arid areas are schinus molle (iponga et al. 2008) and prosopis glandulosa var. torreyana. these species, although present, do not appear to be highly invasive in the cnp at present. however, according to iponga et al. (2008), s. molle has the ability to out compete acacia tortilis and thus poses a considerable threat to cnp. a higher proportion of herbaceous alien plants are to be expected in the karoo (brown & gubb 1986; milton & dean 1998; richardson et al. 1997; dean & milton 2000) and the common invasive herbaceous alien plant species found in this study were consistent with other studies in the karoo (lloyd 1999; milton et al. 2008; milton et al. 1999; weiersbye et al., 2006;). these included atriplex inflata, salsola tragus, pennisetum clandestinum and p. setaceum. although present in cnp, p. setaceum has not yet spread along the major rivers, as it has done in the karoo national park (milton et al. 2008). however, it clearly has the potential to become problematic, which may further promote fire, thereby threatening biodiversity among indigenous succulents. nassella trichotoma, a grass species, is spreading from nqweba dam into the surrounding natural vegetation. the herbaceous species atriplex inflata and salsola tragus were found across different habitat types and sections of the park. these species were common in almost all areas or habitats, including along the rivers running through graaff-reinet. this is most likely because most sections of the park were previously ploughed. what needs to be done? ongoing distribution data collection is required to assist efforts to prioritise and manage the threats posed by alien plants to ecosystem function and structure in the karoo. the precise point locality data collected through the use of the cybertracker system can be used at a later stage to determine frequency or abundance per unit area. the unit area, or resolution of the grid cell in which abundance can be assessed, can then be determined relative to the extent of the area under consideration (foxcroft et al. 2009). very little is known about the impacts of alien plants on the biodiversity (whether compositionally, structurally or functionally) of cnp and this should be given urgent attention. as understanding of the impacts improves, the modes of invasion used by species with the highest impact in cnp should also be examined and used to inform control measures. the influence of the different land use types on invasion by these plants will help identify areas of higher risk. human impacts and the lack of sufficient knowledge and awareness of alien plants should be investigated for the camdeboo municipality. control and rehabilitation to improve the natural vegetation condition will be of utmost importance for areas that are highly invaded. in areas where transformation by alien plants has persisted for a long period, the assumption that the system would self-repair following alien plant removal does not always hold true (esler et al. 2008). management should attempt to restore basic ecosystem functions through providing vegetation cover that is resistant to further invasion (holmes et al. 2008). for example, in the case of cnp, the indigenous malephora sp. might out-compete atriplex inflata and salsola tragus. this mat-forming succulent plant may therefore stabilise the soil to prevent erosion and further spread of other herbaceous alien plant species, in the areas or communities where it naturally occurs. conclusion the first step has been taken by collating a species list and distribution maps to monitor both new introductions of alien plants and the distribution thereof. it is highly likely that the area invaded and densities of alien plants will increase in the cnp and adjacent municipal areas. the aggressively invasive cacti, herbaceous weeds and certain woody species in the rivers will also present serious management challenges to cnp. a control programme will require concerted efforts from cnp management, working for water (invasive alien plant clearing programme) and the people of graaff-reinet. acknowledgements we are grateful to the manager of the cnp, peter burdett, for his assistance and contributing much of the alien plant distribution data used in this paper (with the assistance of cnp field rangers and students). working for water in the camdeboo municipality supplied the eradication and control information for some of the species (charlotte vermuelen). sandra macfadyen assisted with the digitising of the 1981 data, and also created the figures (maps). vutomi mdlhovu also assisted with digitising the 1981 maps. we also thank sanparks and the junior scientist programme (m.l. masubelele was funded by the andrew w. mellon foundation) for financial support. d.m. richardson, m.t. makgohlo and m.t. hoffmann are thanked for their input. we also thank three anonymous reviewers for their constructive comments. our sincerest appreciation goes to m.w. (gretel) van rooyen who was the responsible editor for this article. references alston, k.p. & richardson, d.m., 2006, ‘the roles of habitat feature, disturbance and distance from putative source populations in structuring alien plant invasion at the urban/wildlife interface on the cape peninsula, south africa’, biological conservation 132, 183–198. annecke, d.p. & moran, v.c., 1978, ‘critical reviews of biological pest control in south africa. 2. the prickly pear opuntia ficus-indica (l) miller’, journal of the entomological society of southern africa 41, 161–188. brown, c.j. & gubb, a.a., 1986, ‘invasive alien organisms in the namib desert, upper karoo and the arid and semi-arid savannas of western southern africa’, in i.a.w. macdonald, f.j. kruger & a.a. ferrar (eds.), the ecology and management of biological invasions in southern africa, pp. 93–108, oxford university press, cape town. burdett, p.d., 1995, karoo nature reserve management plan, unpublished report, eastern cape nature conservation, bisho, south africa. camdeboo national park (cnp), 2006, the management plan of camdeboo national park, unpublished report, south african national parks, graaff-reinet, south africa. crimmins, t.m., mauzy, m.s. & studd, s.e., 2008, ‘assessing exotic plant distribution, abundance, and impact at montezuma castle and tuzigoot national monuments in arizona’, ecological restoration 26(1), 44–50. csir, 2004, eastern cape state of the environment report, csir division of water, environment and technology, durban, south africa, produced on behalf of the eastern cape department of economic affairs, environment and tourism, bisho, south africa. dean, w.r.j. & milton, s.j., 2000, ‘directed dispersal of opuntia species in the karoo, south africa: are crows the responsible agents?’ journal of arid environments 45, 305–314. esler, k.j., holmes, p.m, richardson, d.m. & witkowski, e.t.f., 2008, ‘riparian vegetation management in landscapes invaded by alien plants: insights from south africa’, south african journal of botany 74(3), 397–400. doi: 10.1016/j. sajb.2008.01.168. original research masubelele, foxcroft & milton koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 51 5 (page number not for citation purposes) foxcroft, l.c., 2007, ‘patterns and processes of plant invasion in an african savanna ecosystem, with emphasis on multiple spatial and temporal scales’, phd thesis, university of cape town. foxcroft, l.c., henderson, l., nichols, g. & martin, b., 2003, ‘a revised list of alien plants for the kruger national park’, koedoe 46(2), 21–44. foxcroft, l.c., richardson, d.m., rouget, m. & macfadyen, s., 2009, ‘patterns of alien plant distribution at multiple spatial scales in a large national park: implications for ecology, management and monitoring’, diversity and distributions 15, 367–378. doi: 10.1111/j.1472-4642.2008.00544.x. henderson, l., 2001, alien weeds and invasive plants, plant protection research institute handbook, no. 12, plant protection research institute, agricultural research council, pretoria. holmes, p.m., esler, k.j., richardson, d.m. & witkowski, e.t.f., 2008, ‘guidelines for improved management of riparian zones invaded by alien plants in south africa’, south african journal of botany 74, 538–552. doi: 10.1016/j.sajb.2008.01.182. hrusa, g.f. & gaskin, j.f., 2008, ‘the salsola tragus complex in california (chenopodiaceae): characterization and status of salsola australis and the autochthonous allopolyploid salsola ryanii sp. nov. madroňo’, 55(2), 113–131. iponga, d.m., milton, s.j. & richardson, d.m., 2008, ‘soil type, microsite, and herbivory influence growth and survival of schinus molle (peruvian pepper tree) invading semiarid african savanna’, biological invasions 11, 159–169. doi 10.1007/s10530-008-9221-6. lloyd, j.w., 1999, ‘nama karoo’ in j. knobel (ed.), the magnificent natural heritage of south africa, pp. 84–93, sunbird publishing, cape town. loope, l.l., sanchez, p.g., tarr, p.w., loope, w.l. & anderson, r.l., 1988, ‘biological invasions of arid land nature reserves’, biological conservation 44, 95–118. lovegrove, b., 1993, the living deserts of southern africa, fernwood press, cape town. macdonald, i.a.w & frame, g.w., 1988, ‘the invasion of introduced species into nature reserves in tropical savannas and dry woodlands’, biological conservation 44, 67–93. milton, s.j. & dean, w.r.j., 1998, ‘alien plant assemblages near roads in arid and semi-arid south africa’, diversity and distributions 4(4), 175–187. milton, s.j., dean, w.r.j. & rahlao, s.j., 2008, ‘evidence for induced pseudo-vivipary in pennisetum setaceum (fountain grass) invading a dry river, arid karoo, south africa’, south african journal of botany 74, 348–349. milton, s.j., wilson, j.r.u., richardson, d.m., seymour, c.l., dean, w.r.j., iponga, d.m. & procheş, ş., 2007, ‘invasive alien plants infiltrate bird-mediated shrub nucleation processes in arid savanna’, journal of ecology 95(4), 648–661. milton, s.j., zimmermann, h.g. & hoffmann, j.h., 1999, ‘alien plant invaders of the karoo: attributes, impact and control’, in r.j. dean & and s.j. milton (eds.), the karoo. ecological patterns and processes, pp. 274–287, cambridge university press, cambridge. moran, v.c. & annecke, d.p., 1979, ‘critical reviews of biological pest control in south africa. 3. the jointed cactus opuntia aurantiaca lindley’, journal of the entomological society of southern africa 42, 299–329. moran, v.c., hoffmann, j.h. & zimmermann, h.g., 1993, ‘objectives, constraints and tactics in the biological control of mesquite weeds (prosopis) in south africa’, biological control 3, 80–93. moran, v.c. & zimmermann, h.g., 1991. ‘biological control of cactus weeds of minor importance in south africa’, agriculture ecosystems and environment 37, 37–55. mucina, l. & rutherford, m.c., 2006, ‘the vegetation of south africa, lesotho and swaziland’, strelitzia 19, south african national biodiversity institute, pretoria. national park service (nps), 1996, a strategic plan for managing invasive non-native plants on national park system land, viewed 10 august 2007, from http://www.nature.nps.gov/biology/ invasivespecies/strat_pl.htm. palmer, a.r., 1989, ‘the vegetation of the karoo nature reserve, cape province. 1. a phytosociological reconnaissance’, south african journal of botany 55(2), 215–230. pauchard, a. & alaback, p.b., 2004, ‘influence of elevation, land use, and landscape context on patterns of alien plant invasions along roadsides in protected areas of southcentral chile’, conservation biology 18, 238–248. pyšek, p., richardson, d.m., rejmánek, m., webster, g.l., williamson, m. & kirschner, j., 2004, ‘alien plants in checklists and floras: towards better communication between taxonomists and ecologists’, taxon 53(1), 131–143. richardson, d.m., bond, w.j., dean, w.r.j., higgins, s.i., midgley, g.f., milton, s.j., powrie, l.w., rutherford, m.c., samways, m.j. & schulze, r.e., 2000, ‘invasive alien organisms and global change: a south african perspective’, in h.a mooney & r.j. hobbs (eds.), invasive species in a changing world, pp. 303–349, island press, washington d.c. richardson, d.m., macdonald, i.a.w., hoffmann, j.j. & henderson, l., 1997, ‘alien plant invasions’, in r.m. cowling, d.m. richardson & s.m. pierce (eds.), vegetation of southern africa, pp. 535–570, cambridge university press, cambridge. smith, n. & wilson, s.l., 2002, changing land use trends in the thicket biome: pastoralism to game farming, terrestrial ecosystem research unit (teru), report no. 38, nelson mandela metropolitan university, south africa. weiersbye, i.m., witkowski, e.t.f. & reichardt, m., 2006, ‘floristic composition of gold and uranium tailings dams, and adjacent polluted areas, on south africa’s deep level mines’, bothalia 36, 101–127. wells, m.j., balsinhas, a.a., joffe, h., engelbrecht, v.m., harding, g.b. & stirton, s.h., 1986, ‘a catalogue of problem plants in southern africa’, memoirs of botanical survey of south africa, 53, 1–685. zimmermann, h.g., 1991, ‘biological control of mesquite prosopis spp. (fabaceae) in south africa’, agricultural ecosystems and environment 37, 175–186. zimmermann, h.g. & moran, v.c., 1982, ‘ecology and management of cactus weeds in south africa’, south african journal of science 78, 314–320. zimmermann, h.g., moran, v.c. & hoffmann, j.h., 1986, ‘insect herbivores as determinants of the present distribution and abundance of invasive cacti in south africa’ in i.a.w. macdonald, f.j. kruger & a.a. ferrar (eds.), the ecology and management of biological invasions in southern africa, pp. 269– 274, oxford university press, cape town. vol. 51 no. 1 page 10 of 1089 druce.qxd population demography and spatial ecology of a reintroduced lion population in the greater makalali conservancy, south africa d. druce, heleen genis, j. braak, sophie greatwood, audrey delsink, r. kettles, l. hunter and r. slotow druce, d., heleen genis, j. braak, sophie greatwood, audrey delsink, r. kettles, l. hunter and r. slotow. 2004. population demography and spatial ecology of a reintroduced lion population in the greater makalali conservancy, south africa. koedoe 47(1): 103–118. pretoria. issn 0075-6458. many recently established small game reserves in southern africa have introduced lion, and have had to actively manage their populations as the lion are in relatively small (<1000 km²), enclosed areas. this study was undertaken on the greater makalali conservancy (140 km²), limpopo province, south africa to describe population demography and lion movement patterns in order to enhance management decisions. a pride of five lion were introduced to makalali in december 1994 and since then 35 cubs have been born (11.6 % population growth per year over 7.5 years), 31 of which have been translocated to other reserves. to increase genetic diversity, the two male coalition was removed in may 1999 and another two males introduced during july 1999. during the study, the reserve area was increased by approximately 23 %. although the makalali lion utilised a smaller home range during winter (75.3 km²) than summer (106.8 km²), this difference was not significant. the core ranges also shifted between seasons, with 46 % of the summer core being utilised during winter. when the new males were introduced, they centred their core range near their release boma. the females also shifted their core range to this region after their young male cubs were moved into the boma. after the makalali area was increased, the lion increased their home range area, while their core range became reduced and more dispersed. there was no significant difference in the distances the lion moved in a 24 hour period between the two seasons, or before and after the two management actions. the makalali lion also spent more time than expected in low open woodland and grassland habitats and less than expected in low thickets. we conclude that lion populations can successfully exist within small reserves (<100 km²) and that actions to manage the population can be successful if certain criteria are considered. key words: core range, home range, reproduction, territory, movement patterns. d. druce, heleen genis, sophie greatwood, audrey delsink, l. hunter and r. slotow (slotow@ukzn.ac.za), school of biological sciences, university of kwazulu-natal, durban, 4041 republic of south africa; j. braak and r. kettles, makalali private game reserve, p.o. box 809, hoedspruit, 1380 republic of south africa. issn 0075-6458 103 koedoe 47/1 (2004) introduction during the last few years there has been a rapid increase in the number of small reserves (<1000 km²) in south africa. although some of these reserves have been developed to support the hunting industry, many of them have been formed purely as eco-tourism ventures and to enhance biodiversity conservation. as many of these reserves do not contain all species of animals that would have occurred there in the past, some species have been reintroduced. many of these reserves have introduced lion in order to attract guests and to enhance the wilderness experience (cotterill 1995; vartan 2002). however, due to the small size of these reserves, many are only able to sustain small numbers of lion and may even only be able to support one pride. active management is, therefore, required in these reserves for a number of ecological reasons as well as to enhance tourist sightings. druce.qxd 2004/04/15 08:18 page 103 as many of the lion populations are isolated, little or no immigration or emigration occurs, with the result that populations may become inbred (packer 1996) and need to be manipulated in order to manage genetic diversity (grubbich 2001). most managers believe that lion numbers have to be controlled in order reduce their impact on prey populations and the risk of lion breaking out. managers generally aim to maintain lion populations without the likely population peaks and crashes in both lion and prey species that may occur if they were left to carry on without manipulation. a number of studies have shown that lion predation may negatively affect prey numbers (hunter 1998; peel & montagu 1999; harrington et al. 1999). a management tool that can have an effect on the ungulate distribution, and therefore lion predation and distribution, is that of the provision of artificial waterpoints (smuts 1976; mills & retief 1984; kalikawa 1990; stander 1991; owen-smith 1996). as a result, by manipulating lion numbers and factors such as water distribution, managers may be able to reduce predation of certain important species. however, in larger reserves, managers also need to consider the effect of rainfall on the distribution of lion. east (1984) has shown that in arid/eutrophic savannas, total large carnivore biomass is positively correlated with rainfall and that in all areas the biomass of each carnivore species is most closely correlated with the biomass of its preferred size class of prey. by ensuring that lion numbers are kept relatively low, the risk of lion breaking out may also be reduced. a high population density facilitates sub-adult emigration from a pride (hanby & bygott 1987), with the result that a high lion density within a small, enclosed reserve may increase the risk of lion, especially sub-adults, breaking out. because these lion populations are contained within small reserves, even a minor management decision could have a large impact on the lion and/or the reserve ecology, e.g., through a negative impact on prey populations, which could then possibly result in less prey for other predators. as a result, in order to generate sustainable management strategies and to manage effectively, managers need to understand the ecology of lion in small, enclosed systems and how lion populations may respond to various management strategies. however, there is very little information available on the ecology of lion within these small reserves and to their response to management actions. two major management interventions were implemented during the study: (1) the only male coalition in the reserve was removed and a new coalition introduced in its place and (2) the area of the reserve was increased by 23 % through the addition of part of a neighbouring reserve. the aim of this study was to monitor and to determine the effect of these two management interventions on the population demography and movement patterns of the makalali lion pride. study area this study was undertaken in the greater makalali conservancy, limpopo province, south africa (29º09's, 30º42'e) between february 1998 and december 2001. makalali is situated on the lowveld plain at an altitude of between 300 m and 500 m above sea-level. the landscape is a combination of undulating terrain and rocky outcrops. makalali is found within the savanna biome of southern africa, with mixed lowveld bushveld (low & rebelo 1996; type 19) and mopane bushveld (low & rebelo 1996; type 10) as the dominant vegetation types. makalali contains only one river, the makhutswi river, a perennial tributary of the olifants river. within this riverine belt there is a high diversity of tree species. this river splits makalali and runs from west to east, through makalali. artificial waterpoints have been created in makalali with some of them being supplied with borehole water, especially during the dry winter months. makalali is a relatively dry area with an average annual rainfall of 450 mm. most of the rains fall in the summer months between october and march. temperatures in the koedoe 47/1 (2004) 104 issn 0075-6458 druce.qxd 2004/04/15 08:18 page 104 reserve vary between 3 °c in winter to above 36 °c in summer. makalali was formed in 1993 by the initial purchase of 7500 ha of cattle ranchland. it has been extended by the acquisition of adjacent farms and the co-operation of surrounding game reserve owners has led to fences being removed between neighboring reserves, with the result that the conservancy is now approximately 14000 ha (fig. 1). large mammals that would previously have been in the area were introduced, including a pride of lion introduced during december 1994. this pride, which originated from the kruger national park, consisted of five individuals, a mother and her four cubs (two males and two females). the pride settled in well and after only three years the first cubs were born, from a mother-son mating. methods a lion database has been maintained since the lion were introduced to makalali in 1994. individuals were identified by the patterns of dots above their rows of whiskers as well as features such as facial scars and ear tears. the database lists an identity number, name, mother and father, date of birth and death, origin and location (makalali or where they have been translocated) for each lion. fathers were assigned to the male coalition that was present in makalali during that time (there was only one male coalition present at any given time), while the age of the cubs and their association with lionesses, alone or in a group, were used to determine their mother. data from the date of reintroduction until the end of january 2002 were used to determine patterns in reproduction. detailed lion data were recorded on makalali between 7 february 1998 and 13 december 2001. most of the information was collected by rangers during game drives, with some of the information being collected by researchers and other staff working on makalali. as makalali is an eco-tourism destination with game-viewing offered as a guest activity, rangers track lions on foot if they are not on the roads. once they have located the lions they return to the vehicle and drive the guests off-road into the sighting. as a result, there were a large number of sightings that did not occur along roads. lion were generally located between 05:30 and 10:30 in the issn 0075-6458 105 koedoe 47/1 (2004) fig. 1. the greater makalali conservancy with the pidwa section before the fences were removed (dashed line) in october 2000. the black line through the conservancy indicates the makhutswi river and the circle near the centre of the conservancy indicates the location of the lion release boma. druce.qxd 2004/04/15 08:18 page 105 morning and 15:30 and 20:30 in the evening, however, there were periods (such as immediately after the introduction of the new male coalition) that lion movement was monitored outside these hours. lion data that were recorded included the date, time, individuals present, position (road and gps), behaviour and when there was a kill, what the prey was. prey data are presented elsewhere (druce et al. in press). movement patterns — introduction of new pride males on 21 may 1999, the two makalali pride males (mak 1 and mak 2) were removed from the pride and translocated to kapama game reserve. the same day the two new males (mak 15 & mak 16) were translocated from kapama and released into the makalali boma (fig. 1). this was done in order to introduce new genes into the makalali pride and to prevent the males from breeding with their offspring. in order to determine if the introduction of the new males had any effect on the makalali pride's ranging patterns, we used gps positions for both the females and the males for the last six months before the removal of the old males and the first six months after the release of the new males to estimate the 50 % kernel ranges and 95 % kernel ranges during these time periods. the animal movement analysis arcview extension (hooge & eichenlaub 1997) was used in all gis analyses to estimate the ranges, with the 95 % kernel being used as the estimate of home range and the 50 % kernel as an estimate of core range. a least squares cross-validation (lscv) smoothing factor of 500 m was also used throughout all gis analyses. fifty percent was used to denote core area (mizutani & jewell 1998), while 95 % was used to denote home ranges as this excludes occasional sallies outside their normal range (burt 1943). by including these occasional sallies in the home range, one may generate range estimates that are of a large and bizarre shape (burt 1943). as a result, most studies define the home range as the smallest area containing 95 % of the distribution (seaman & powell 1996; apps 1999). separate maps were produced for the females before and after the introduction of the new males and for the old and new males. we assessed shifts in range by contrasting the old and new core ranges and home ranges. we determined if males changed the distance that they covered between consecutive days by contrasting distances moved over the last six months the old males were on makalali with the first six months post-release of the new males. the males and females were considered separately, with the male coalition being considered as one group and the females as individuals. only the four females that survived the ‘take-over’ by the new males were used in the analysis as this allowed direct comparison between the two time periods. the animal movement analysis arcview extension (hooge & eichenlaub 1997) was used to determine the distance between consecutive sightings for the male coalition and each of the females. the number of days between each sighting was determined for the male coalition and the four females and only those sightings that occurred on consecutive days were used in the analysis. when one of the males was not at a sighting (very rarely) the number of days between sightings was taken to be the number of days from the last sighting of that particular male, whether with the other male or alone. the number of days between sightings for each female was recorded as the number of days from the last sighting of that particular female, whether in a group or alone. the distances moved by the old and new male coalitions during the two six month periods were compared, while the mean distances moved by each individual female were calculated for each six month period, with these means then being compared. as these distances were straight line distances between two points recorded on consecutive days and not recorded by continuous tracking, they cannot be viewed as actual distances moved and should rather be used as an indication of whether change took place or not. — removal of pidwa fence during october 2000 the fence between makalali and pidwa game reserve was removed enlarging the size of the conservancy from 11089 ha to 13600 ha. fence removal was started at both the northeastern and northwestern sections of the fence simultaneously. the whole fence was systematically removed from the northern boundary on both sides, with the southern section of the fence being removed last. fence removal was completed within a month. in order to determine if there was any change in range utilisation before and after reserve expansion, we produced kernel ranges for the males and females for the six-month period before fence removal was started and the first six months after fence removal was begun. gps positions of all the lion, for these six-month periods were used to produce core and home ranges for the male coalition and the females. the core ranges and home ranges, after fence removal, were compared to those before reserve expansion to determine the percentage overlap. koedoe 47/1 (2004) 106 issn 0075-6458 druce.qxd 2004/04/15 08:18 page 106 with reserve expansion, one may expect the lion to increase the distance that they cover between consecutive days in order to investigate the new area. as a result, we determined the distance that the male coalition and the individual females moved during consecutive days for the six months before and after the removal of the fence was begun. the same methods used in determining the distances moved in consecutive days for the introduction of the new males were employed. the distances moved by the male coalitions between the two time periods were compared, while the mean distances moved by each of the four individual females were calculated for each time period and these means compared. — summer and winter patterns various studies have demonstrated that there is an expansion in lion home range during the wet season (van orsdol et al. 1985; hunter 1998) while another reported an expansion during the dry season (viljoen 1993). although these changes in home range size could be brought about by purely ecological conditions and changes in herbivore distribution, home range expansion might not be evident in a small reserve especially if lion are using all the area available to them. in order to determine what occurs in makalali, the year was divided up into two seasons, each of six months. the summer months refer to the beginning of november through to the end of april, while the winter months refer to the beginning of may through to the end of october. this split was chosen as the most rains fall in makalali between november and april (summer rainfall pattern) and this is also the period that the highest temperatures were recorded. the sighting data were grouped based on these periods and core and home ranges estimated in arcview. the core and home ranges were compared between winter and summer to determine the percentage overlap between the two time periods. if lion change their home ranges between seasons, they may also be expected to change the distance that they move between consecutive days. as a result, we calculated the distances that the male coalition and the individual females moved during consecutive days for the summer and winter periods using the same methods used in determining the distances moved between consecutive days during the introduction of the new males. the distances moved by the male coalitions between the two 'seasons' were compared. as all five of the lionesses were present during both summer and winter, the mean distances moved by each of the five individual females were calculated for both 'seasons' and these means compared. habitat influences lion select habitats based on factors such as prey availability (east 1984; creel & macdonald 1995; funston et al. 1998), cover for hunting (van orsdol 1984) and protection for cubs (donkin 2000). in order to determine in what habitat type the makalali lion spent the greatest proportion of their time, the vegetation map of the greater makalali conservancy was reduced to four habitat types, determined by their degree of openness. the nine vegetation types with their sub-communities and variants, as described by druce (2000), were reduced to four structural habitat types, the characteristics of which are summarised in table 1. as no vegetation classification for pidwa was available, only the sightings recorded before the fence removal were used in the analysis. these sightings were used in the geoprocessing extension of arcview to determine the number of sightings per vegetation type and ultimately the number of sightings per structural habitat type. the total number of sightings within each habitat was then compared to the expected number of sightings based on the area each habitat type occupied within the conservancy. issn 0075-6458 107 koedoe 47/1 (2004) table 1 characteristics of the four structural habitat types descriptor tree density grass and shrub density area (km²) low closed woodland moderate to dense moderate to sparse 79.06 low open woodland moderate to open moderate 8.1 low thicket dense sparse 18.16 grasslands open dense 5.34 druce.qxd 2004/04/15 08:18 page 107 results population demography until 31/06/02 since the introduction of the lion, 35 cubs have been born (table 2) at a birth rate of 11.6 % per year over 7.5 years. thirty one of these cubs have been translocated to other reserves (fig. 2). the only cubs born on the conservancy that have not been translocated are the two female cubs that were born first on the property (mak 6 & mak 7) and the two cubs born in january 2002 (mak 34 & mak 35). the only other translocations that have taken place were the exchange of the two males that were introduced first to the reserve. these males were exchanged with two other males from kapama during 1999. there has only been one lion death on the property. this was the oldest original female who was killed by the new males when they were introduced. she was defending cubs at the time and formed an obstacle to the ‘takeover’ by the new males. one nomadic male (mak 30) came through the fence into the conservancy towards the end of november 2000 and was last seen on 20 february 2001. it was presumed that he broke out through the fence shortly afterwards, as tracks were seen going out through the fence. koedoe 47/1 (2004) 108 issn 0075-6458 table 2 number of cubs born to each female on makalali until 31/06/02 number name male female total birth rate cubs cubs /female /month1 mak 3 stickynyawu 0 2 2 0.07 mak 4 one-eye 8 4 12 0.18 mak 5 bertha 5 2 7 0.13 mak 6 gertrude 4 3 7 0.28 mak 7 dorris 1 6 7 0.29 total 18 17 35 1note: birth rate per female per month was calculated by dividing the number of cubs born to each female by the number of months between their first litter and the end of june 2002 or the date that they left makalali. all of the litters (n = 12 litters) ranged in size from two to four individuals, with all but two of the litters a single sex litter. after cubs were removed (mean age = 16.6 months, n = 20 cubs), there was an average of 6.1 months (approximately 189 days) before their next litter was born (n = 5 litters). assuming a gestation period of 110 days (mills & hes 1997), the average interval from cub removal to conception was 79 days (approximately 2.5 months; n = 5 litters). the average interval between litters when cubs were removed from the conservancy was 22.8 months (n = 5 litters) and when the litters were not removed the average inter-birth interval was 15 months (n = 2 litters). however, only two litters were born while previous litters were still on the conservancy and both litters were born to the same female, indicating that she may have had an inter-birth interval that was shorter than that of the other lionesses. the age at which the makalali females had their first successful litters ranged from 39 to 51 months with an average of 43.25 months (n = 4 females). introduction of new pride males the two new males from kapama were kept in the boma from the 21 may 1999 until their release on the morning of 13 july 1999. immediately they were out of the boma the three smallest male cubs (mak 12, 13 & 14), who were 15 months old, were darted, removed from the pride and placed in the boma to ensure that the new males did not kill them as occurs during natural pride takeovers (packer et al. 1988). they had to be placed in the boma for approximately three months, as the permits for their translocation had not yet been received. for the first 11 days post-release, intense monitoring was carried out, with most of the lions being located every day. on the second day after their release, the new males met the pride for the first time. initial contact was violent, resulting in the pride of nine scattering in different directions. seven days after the initial contact (21 july 1999), the males located female mak 3 and killed her. the following druce.qxd 2004/04/15 08:18 page 108 issn 0075-6458 109 koedoe 47/1 (2004) fi g. 2 . r ep ro du ct iv e ou tp ut o f t he li on s af te r i nt ro du ct io n to th e g re at er m ak al al i c on se rv an cy . p oi nt s on th e le ft -h an d si de o f t he li ne s in di ca te th e da te a t w hi ch th e in di vi du al e nt er ed th e sy st em , w hi le th os e on th e ri gh tha nd s id e ar e w he n th e in di vi du al d ie d or w as tr an sl oc at ed . n um be rs to th e le ft o f ea ch s ym bo l a re th e st ud bo ok n um be rs . c ub s bo rn a re s ho w n ab ov e th ei r m ot he r, w hi le th e da m a nd s ir e fo r ea ch li tte r ar e gi ve n on th e ri gh tha nd s id e of th e fi gu re . druce.qxd 2004/04/15 08:18 page 109 day (22 july 1999) the males located the other females for only the second time postrelease. initially the females were not worried but soon ran off, without any aggression from the males. seventeen days after the males were released from the boma they were seen again with the females, with mak 15 mating with mak 5 and mak 16 mating with mak 6. for the six-month period before the new pride males were introduced into the conservancy, the females concentrated their core range (50 % kernel) in the north western section of the conservancy (fig. 3a). however, they utilised most of the conservancy with the exception of the far north eastern and far south eastern sections near the boundary fence. directly after the new males had been introduced, the females’ core range shifted further east into the central areas of the conservancy and along the river (fig. 3b). twenty three percent of the old core range was included in the new range. after the introduction of the new males, the females did not use the north eastern section of makalali at all, while 45 % of their old home range (95 % kernel) was included within the new range. the core range (50 % kernel) of the old males (fig. 3c) was similar to that of the females over the same time period. however, they spent a greater proportion of their time towards the centre of makalali and their home range (95 % kernel) was not as extensive as that of the females. the new males concentrated their home range along the river for the first six months after release (fig. 3d). their core range was near the boma where they had been held prior to release, as well as towards the eastern boundary fence. during this time, the new males’ core range only overlapped that of the old males by 12.5 %, while their home range overlapped that of the old males by approximately 37 %. anovas were used to compare the distances moved by the male lion. although the males covered greater distances than the original males on a few occasions (fig. 4a), there was no significant difference in distance moved in consecutive days between the two male coalitions (f1, 42 = 0.003, p = 0.959). the female distance data were analysed using a wilcoxon paired signed ranks test to compare the mean distance for each of the females before and after introduction of the new males. as with the males, there was no significant difference in the distance moved between consecutive days before and after introduction of the new males (z = 0.000, p = 1.0). as a result, although the females shifted their home range post-release of the new males, they did not change the distances they were moving between consecutive days. the first six months after the removal of the old males and the introduction of the new males saw a concentration of the home range for both the new males and the females around the boma from where the males were released. at the same time as the new makalali males were released, the remaining cubs (± 15 months old) that had not yet been translocated, were removed from the pride and put in the boma to ensure that they were not killed by the new males during a pride take-over. they remained in the boma for the first four months post-release of the new males, while the conservancy manager awaited permits to translocate them. as a result, the new males spent a large portion of their time near the boma trying to get at the young males possibly either because they were males or because they were still cubs. removal of pidwa fence for the six months prior to the pidwa fence removal, the lion core range (50 % kernel) was situated in the centre of makalali, while their home range (95 % kernel) was concentrated in the central and northwestern areas (fig. 3e) with only a few sightings in the far northeastern section. after the fence was removed, the lion did not take long to move into the pidwa section, with the females making use of the new area within the first week after the fences were removed. however, after fence removal the core range became dispersed into much smaller pockets in the centre of makalali and towards the koedoe 47/1 (2004) 110 issn 0075-6458 druce.qxd 2004/04/15 08:18 page 110 issn 0075-6458 111 koedoe 47/1 (2004) fig. 3. the effect of two management interventions on the ranging behaviour of the makalali lions between consecutive days. core ranges (dark grey shading) are the 50 % kernel ranges while the home ranges (light grey shading) are the 95 % kernel ranges. (a) female ranges before the introduction of the new males; (b) female ranges after the introduction of the new males; (c) old male ranges; d) new male ranges; (e) all lion ranges (males and females combined) before fence removal; (f) all lion ranges (males and females combined) after fence removal. druce.qxd 2004/04/15 08:19 page 111 north western and western boundaries (fig. 3f). this, as well as an increase and dispersal in the home range, indicates that the lion did not spend much time in any areas in particular, but were constantly shifting position and utilising different areas. the core range after the fence removal overlapped the old core range by 5 %, while the new home range overlapped the old by 84 %. an interesting point to note is that after the fence was removed the lion tended to use much more of the north eastern and south eastern section than prior to reserve expansion. although the core range for the makalali lion became much more dispersed after makalali’s area was increased, both the anova for the males (f1,222 = 0.054, p = 0.816) and the wilcoxon paired signed ranks test for the females (z = -1.826, p = 0.068) indicate that there was no significant difference between the distances they moved before and after the fence was removed (fig. 4b). the core ranges and home ranges for the makalali lion for the various management actions and for the summer and winter seasons are relatively small. the core ranges vary from 1.9 km² (first six months after fence removal) to 11.1 km² (first six months before fence removal), less than 10 % of the area available to them. this indicates that the makalali lion do not need an extensive area in which to meet most of their requirements. the home ranges (24.9 km² to 106.8 km²) also do not extend to the entire reserve area (140 km²). summer and winter patterns during the summer months, the makalali lion utilise a large proportion of the reserve (fig. 5a), with their core range (50 % kernel) concentrated in the central to western areas of makalali. however, during winter (fig. 5b) their home range (95 % kernel) is much reduced, with little of the northern, south western and south eastern areas of makalali being used. their core range also becomes slightly more concentrated and is koedoe 47/1 (2004) 112 issn 0075-6458 fig. 4. the effect of management action on daily movement of the makalali lions. (a) the effect of the introduction of new males to the makalali conservancy on the distance moved by the original lions in consecutive days. distances moved over the first six months post-release of the new males are indicated by hatched boxes, while the distances moved over the last six months the original lions were on the property are indicated by the clear boxes. (b) effect of reserve area expansion on the distances moved by lions in consecutive days. the six month period immediately before reserve expansion is shown by the clear boxes, while the first six months after fence removal was started are given by the hatched boxes. (c) the effect of season on distances moved by the makalali lions in consecutive days. summer is shown by the clear boxes, while winter is shown by hatched boxes. druce.qxd 2004/04/15 08:19 page 112 found in the central area along the river, as this is the only permanent water supply towards the end of winter. forty six percent of the summer core range is utilised during winter, while 59 % of the summer home range is used during winter. although the kernel ranges increased during the summer months, there was no significant increase in the distance that the males (anova; f1, 167 = 0.83, p = 0.701) or females (wilcoxon paired signed ranks test; z = -1.461, p = 0.144) moved in consecutive days between the two seasons (fig. 4c). habitat influences the low closed woodland constituted the greatest area in the conservancy with the grassland habitat being the smallest (table 1). as lion were tracked on foot if they were not found on the roads, one would not expect a significant difference between the numbers of sightings in each habitat type and the expected based on the area of each habitat type. however, there was a significant difference (χ² 0.05, 3 = 172.53, p <0.05), with the lion spending more time than expected in the low open woodland and grassland habitats and less than expected in the low thickets (fig. 6). discussion in makalali, the age at first conception, interval between litters, and interval between loss of previous litter and birth of next (in makalali loss was due to sub-adult cub removal) was within the range reported from both the serengeti national park, tanzania (schaller 1972; packer et al. 1988) and nairobi national park, kenya (rudnai 1973), but was less than half the interval (40 months) of litters within two prides in the kruger national park (funston et al. 2003). the makalali litter sizes showed a similar trend to those of large reserves in east africa such as nairobi national park (rudnai 1973), serengeti national park (bertram 1975; packer & pusey 1995) and the issn 0075-6458 113 koedoe 47/1 (2004) fig. 5. the effect of season on lion spatial use. core ranges (50 % kernel ranges) are shown in dark grey, while home ranges (95 % kernel ranges) are shown in light grey. (a) all lion ranges during the summer months; (b) all lion ranges during the winter months. druce.qxd 2004/04/15 08:19 page 113 ngorongoro crater (packer & pusey 1995). however, although the sex ratios of individual litters on makalali were not 1:1 males to females as reported from nairobi national park (rudnai 1973) or between 1:5 to 1:7 (kruger national park; smuts 1976), the sex ratio of all the makalali litters combined was very close to 1:1. this is very similar to the mean ratio of 1:1.1 males to females that funston et al. (2003) reported from all litters within their study area in the kruger national park and suggests that although lion in smaller reserves may not face competition from other prides, as occurs in these larger reserves, reproductive patterns and frequency may still be similar as a result of constant management intervention (i.e., removal of cubs). when introducing lion into a reserve, managers need to consider pride structure. an unnatural situation, as occurred in the greater makalali conservancy, where mother son matings occurred needs to be avoided and females need to be free to mate with unrelated males in order to ensure that no inbreeding occurs. it was noted that the makalali lion home range did not completely fill the reserve, which suggests that the lion were satisfying their needs within the reserve and as far as area is concerned, makalali could possibly support a greater lion population. this was koedoe 47/1 (2004) 114 issn 0075-6458 fig. 6. habitat selection by lion in makalali. the actual number of sightings (clear bars) and expected (black bars) number of sightings in the four habitat types investigated. further emphasised by the relatively small core range used by the makalali lion. this suggests that small conservation areas are able to successfully support lion. however, lion and prey populations within these small areas will need to be constantly monitored and possibly managed to ensure that lion do not have a negative impact on prey populations. the makalali lion had a smaller kernel range during the winter months than in the summer, concentrating their activity around the makhutswi river. although this reduction in home range in makalali was small, home range reduction during winter has been shown in other reserves containing a greater number of prides (van orsdol et al. 1985; hunter 1998), while in others, expansion in home range occurs during the dry seasons (viljoen 1993). in the savuti area in northern botswana, ranges increased during winter probably as a result of prey biomass decreasing within the lion ranges forcing them to expand their range in search of prey species (viljoen 1993). however, in phinda game reserve, south africa, reduced winter ranges were concentrated around artificially supplied waterpoints (hunter 1998). in the makalali conservancy, most of the waterpoints throughout the conservancy dry up during the winter period, with the makhutswi river being the only permanent water supply throughout the dry season. as a result, most of the prey species concentrate along the length of the river, resulting in the lion reducing their winter range and concentrating their activity along the river. the provision of artificially supplied water during the dry period in reserves may affect the movement patterns of ungulates (western 1975; mills & retief 1984; owen-smith 1996) and as a result the distribution and ranging patterns of predators such as lion. a study in the kruger national park also showed that the number of lion prides in an area increased with an increase of permanent water (smuts 1976). therefore, managers of small, enclosed reserves may need to consider the supply of artificial waterpoints within the home ranges of lion in order to ensure there is enough opportunity for prey druce.qxd 2004/04/15 08:19 page 114 species to escape predation (hunter 1998) by providing a number of alternative water sources during the dry months. as with the makalali lion, the distance travelled by both the serengeti and ngorongoro crater lion was not significantly different between wet and dry seasons (hanby et al. 1995), however, the makalali lion did not travel as far, on average, within consecutive days as the east african lion. there was also no difference in distance moved when the conservancy area was increased or when the new males were introduced. the increase in makalali area also resulted in a shift of the core range (50 % kernel) indicating that an increase in reserve area could result in lion spending less time in certain areas, but rather spreading their activity in order to make use of the new area available to them. this is important with respect to tourist sightings, because if lion spend less time in certain areas, it might become more difficult to locate them. it is interesting to note, however, that although they may increase their range, the distances that they covered in consecutive days did not change. this indicates that although a larger area may be made available to them, males may not increase the distances moved during territory patrols. hanby et al. (1995) suggested that the lack of water and cover in the serengeti ecosystem resulted in the serengeti lion covering greater distances than the ngorongoro crater prides in search of those factors. it would appear, therefore, that the availability of cover, water and prey within a system would allow the lions to cover less distance within a 24-hour period. the lionesses concentrated their activity in the boma area after the release of the males possibly because the cubs had been put in the boma. it has been noted in makalali on a number of other occasions when cubs have been removed from the pride and put into the boma whilst awaiting translocation, the pride females also moved into the area near the boma. in order to ensure that the lionesses do not shift their movement patterns in response to the capture of their cubs (i.e. ensuring a minimum amount of manipulation), cubs should be translocated immediately after capture and not retained in a boma. a further factor that may have influenced the position of the core area for the first six months postrelease is that this area near the boma is usually used as the core area during winter and the males were released just before the middle of the winter period. in phinda, hunter (1998) noted that lion and cheetah remained near their release site for at least one week post-release and thereafter utilised the area near their release site as part of their home range. lion released from a boma in zimbabwe were also reported to have remained within 5 km of the boma for the first 16 days post-release (hoare & williamson 2001). as a result, the concentration of male lion activity around the boma area after their release is in accordance with other studies, but their continued use of the boma area could have been caused by the fact that the females concentrated their activity over this period in the boma area as well or because of the presence of the young male cubs in the boma. translocating the cubs immediately may have resulted in the males moving out of the area sooner than they did. however, if managers wanted to shift lion activity to a particular area within a reserve, they could leave cubs awaiting translocation, in a boma within that particular area. this may not be a desirable management technique, however, as it involves a large amount of manipulation and could, therefore, be seen as unethical. a further reason why the male lion could have been concentrating their activity around the boma was because the lionesses concentrated their activity in this area and many would have been ready to mate. this was further supported in that many of the male lion sightings within the first six months post-release where there were females present, only a single lioness was present, with mating often being witnessed. seventeen days after the new male lion were released from the boma, they were observed mating with one of the females. these results are similar to those reported from zimbabwe by hoare & williamson (2001), where on issn 0075-6458 115 koedoe 47/1 (2004) druce.qxd 2004/04/15 08:19 page 115 day 16 to 20 post-release of a female with sub-adult cubs, she was observed mating with resident males. the makalali lion appeared to show a preference for the grassland and low open woodland habitat types and spent less time than expected in low thickets. this trend has also been shown in phinda game reserve, south africa, where hunter (1998) demonstrated that lion showed a greater than expected utilisation of the grassland habitat type and less than expected use of dense vegetation types. however, in the kruger national park, south africa, lion select for open tree savannas (similar in structure to the low open woodland in makalali) and thickets (donkin 2000). although areas of denser vegetation may provide better cover for cubs (during most of the study period, there were cubs and sub-adults on the conservancy), food supply has been shown to affect cub survival (bertram 1975). open woodlands provide protected areas for cubs, while at the same time provide a good environment for hunting, while grasslands provide much fewer hiding places but better visibility for hunting (donkin, 2000). grasslands will also, however, provide greater visibility for prey species when trying to avoid predation, although this advantage may be reduced during the night. this may provide answers as to why the makalali lion preferred these habitat types. this study has highlighted the importance of collecting detailed information on lion movements and activities in order to gain a better understanding of lion ecology on a small, enclosed reserve. we conclude that it is possible to exchange male lion coalitions in order to introduce fresh genes into a population without breakouts from the reserve. however, careful note needs to be made of the reproductive state of all the lion on the reserve in order to prevent any mortality through infanticide. although infanticide occurs naturally in large systems, private game reserves often aim to generate income through the sale of excess animals and as a result, do not want to loose potential income through the death of cubs. increasing a reserve’s area can result in lion successfully utilising the new area, however, this may also result in dispersion of the lion ranges, making it more difficult to locate them. however, managers would need to weigh up benefits gained from increasing a conservation area and allowing an ecological system to operate with less human intervention, with the possible costs of not having as regular lion sightings. our results also indicate that small reserves can adequately house lion, with core ranges being as small as 1.9 km² for a six month period (core range after fence removal) and home ranges as small as 24.9 km² (home range of first six months of new males). therefore, depending on prey density, lion could be adequately housed on reserves smaller than 100 km². acknowledgements the rangers and trackers of makalali private game reserve and garonga safari camp are thanked for providing data over the years. martin bornman and the various students of africa conservation experience are also thanked for providing additional data. roelof nieman is thanked for providing information relating to management carried out within the greater makalali conservancy. yvonne ebertson and paul woods are thanked for their generous financial support and interest in the project, through their donations to the makalali land & wildlife trust. craig packer is thanked for providing funding from the university of minnesota and mgm grand. financial support was also provided by the makalali land & wildlife trust, a thrip grant (no. 1897) and a nrf grant (2053623), while l.h. was a nrf post-doctoral fellow. references apps, c.d. 1999. space-use, diet, demographics and topographic associations of lynx in the canadian rocky mountains: a study. usda forest service research papers 1-11. bertram, b.c.r. 1975. social factors influencing reproduction in wild lions. journal of zoology, london 177: 463-482. burt, w.h. 1943. territoriality and home range concepts as applied to mammals. journal of mammalogy 24: 346-352. koedoe 47/1 (2004) 116 issn 0075-6458 druce.qxd 2004/04/15 08:19 page 116 cotterill, a.h. 1995. lions (panthera leo) on a commercial wildlife ranch: a zimbabwean casestudy and feasibility assessment. msc thesis, university of zimbabwe. creel, s. & d. macdonald. 1995. sociality, group size and reproductive suppression among carnivores. advances in the study of behavior 24: 203-257. donkin, d.a. 2000. lion spatial ecology: the effect of habitat on lion group dynamics. msc thesis, university of natal, durban. druce, d.j. 2000. factors affecting millipede, centipede and scorpion diversity in a savanna environment. msc thesis, university of natal, durban. druce, d.j., h. genis, j. braak, s. greatwood, a. delsink, r. kettles, l. hunter & r. slotow. in press. prey selection of a reintroduced lion popultion in the greater makalali conservancy, south africa. african zoology. east, r. 1984. rainfall, soil nutrient status and biomass of large african savanna mammals. african journal of ecology 22: 245-270. funston, p.j., m.g.l. mills, h.c. biggs & p.k.r. richardson. 1998. hunting by male lions: ecological influences and socioecological implications. animal behaviour 56: 1333-1345. funston, p.j., m.g.l. mills, p.k.r. richardson & a.s. van jaarsveld. 2003. reduced dispersal and opportunistic territory acquisition in male lions. journal of zoology, london 259: 131-142. grubbich, j.d. 2001. genetic variation within and among fragmented populations of south african lions panthera leo: implications for management. msc thesis, university of pretoria. hanby, j.p. & j.d. bygott. 1987. emigration of subadult lions. animal behavior 35: 161-169. hanby, j.p., j.d. bygott & c. packer. 1995. ecology, demography and behaviour of lions in two contrasting habitats: ngorongoro crater and the serengeti plains. pp. 315-331. in: sinclair, a.r.e. & p. arcese (eds.). serengeti ii. chicago: chicago press. harrington, r., n. owen-smith, p.c. viljoen, h.c. biggs, d.r. mason & p. funston. 1999. establishing the causes of the roan antelope decline in the kruger national park, south africa. biological conservation 90: 69-78. hoare, r.e. & j. williamson. 2001. assisted reestablishment of a resident pride of lions from a largely itinerant population. south african journal of wildlife research 31: 179-182. hooge, p.n. & b. eichenlaub. 1997. animal movement extension to arcview version 1.1. alaska biological science center. anchorage: u.s. geological survey. hunter, l.t.b. 1998. the behavioural ecology of reintroduced lions and cheetahs in the phinda resource reserve, kwazulu-natal, south africa. phd thesis, university of pretoria. kalikawa, m.c. 1990. baseline vegetation description at artificial watering points of central kalahari game reserve. african journal of ecology 28: 253-256. low, a.b. & a.g. rebelo. 1996. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. mills, g. & l. hes. 1997. the complete book of southern african mammals. cape town: struik winchester. mills, m.g.l. & p.f. retief. 1984. the effect of windmill closure on the movement patterns of ungulates along the auob riverbed. koedoe 27: 107-118. mizutani, f. & p.a. jewell. 1998. home-range and movements of leopards on a livestock ranch in kenya. journal zoology, london 244: 269-286. owen-smith, n. 1996. ecological guidelines for waterpoints in extensive protected areas. south african journal of wildlife research 26: 107112. packer, c. 1996. who rules the park? wildlife conservation may/june: 36-39. packer, c., l. herbst, a.e. pusey, j.d. bygott, j.p. hanby, s.j. cairns & m.b. mulder. 1988. reproductive success of lions. in: cluttonbrock, t.h. (ed.). reproductive success. chicago: university of chigago press. packer, c. & a.e. pusey. 1995. the lack clutch in a communal breeder: lion litter size is a mixed evolutionary stable strategy. the american naturalist 145: 833-841. peel, m.j.s. & g.p. montagu. 1999. modelling predator-prey interactions on a northern province game ranch. south african journal of wildlife research 29: 31-34. rudnai, j. 1973. reproductive biology of lions (panthera leo massaica neumann) in nairobi national park. east african wildlife journal 11: 241-253. schaller, g.b. 1972. the serengeti lion. chicago: university of chigago press. seaman, d.r. & r.a. powell. 1996. an evaluation of the accuracy of kernel density estimators for home range analysis. ecology 77: 2075-2085. smuts, g.l. 1976. population characteristics and recent history of lions in two parts of the kruger national park. koedoe 19: 153-164. stander, p.e. 1991. demography of lions in the etosha national park, namibia. madoqua 18: 1-9. issn 0075-6458 117 koedoe 47/1 (2004) druce.qxd 2004/04/15 08:19 page 117 van orsdol, k.g. 1984. foraging behaviour and hunting success of lions in queen elizabeth national park, uganda. african journal of ecology 22: 79-99. van orsdol, k.g., j.p. hanby & j.d.bygott. 1985. ecological correlates of lion social organisation. journal of zoology, london 206: 97-112. vartan, s. 2002. overpopulation and inbreeding in small game reserves: the lion panthera leo as a case study. msc thesis, university of cape town, rondebosch. viljoen, p.c. 1993. the effects of changes in prey availability on lion predation in a large natural ecosystem in northern botswana. symposium zoological society, london 65: 193-213. western, d. 1975. water availability and its influence on the structure and dynamics of a savannah large mammal community. east african wildlife journal 13: 265-286. koedoe 47/1 (2004) 118 issn 0075-6458 druce.qxd 2004/04/15 08:19 page 118 filelist convert a pdf file! page 1 myburgh & bredenkamp.qxd macro-channel riparian vegetation of the olifants river system in the grassland biome, mpumalanga w.j. myburgh and g.j. bredenkamp myburgh, w.j. and g.j. bredenkamp. 2004. macro-channel riparian vegetation of the olifants river system in the grassland biome, mpumalanga. koedoe 47(1): 41–54. pretoria. issn 0075-6458. the vegetation associated with the macro-channel of the olifants river system was investigated to identify plant communities mappable at a spatial scale of 1:250 000. the results obtained by using the phytotab pc-classification and mapping program package, revealed eight distinct plant communities associated with the macro-channel of this river system that occurs within the grassland biome. key words: macro-channel, olifants river system, plant communities, riparian vegetation. w.j. myburgh , private bag x680, department of nature conservation, tshwane university of technogoly, pretoria, 0001 republic of south africa; g.j. bredenkamp, department of botany, university of pretoria, pretoria, 0002 republic of south africa. issn 0075-6458 41 koedoe 47/1 (2004) introduction the olifants river, the second largest river in the former transvaal (now the limpopo province and mpumalanga), is one of the most severely polluted rivers in the region (olifants river forum 1995). this project was initiated, inter alia, to obtain a holistic perspective of the current state of the vegetation associated with the macro-channel, including the extent and distribution of exotic vegetation within the macro channel of the river system (myburgh 2000, 2001). in the study, which was conducted over a period of four years, the vegetation was monitored prior to and after flooding events. this report focuses on the species composition and structure of the plant communities associated with the macro-channel of the olifants river system within the grassland biome. study area the grassland biome section of the olifants river (fig. 1) stretches downstream from the river's origin in the vicinity of the town breyten, to the witbank dam (rutherford & westfall 1986). the vegetation of this area is broadly referred to as bankenveld (veld type 61) (acocks 1988). the width of the macro-channel varies from only a few metres (< 5 m), with no distinct riparian vegetation zone on the river banks, to 32 m wide with a well-developed riparian zone quite distinct from the adjacent zonal grassland. this section of the olifants river flows through a landscape that may, in general, be described as flat to rolling plains dominated by grassland, with woody plant species restricted to the rocky river banks. the catchment area surrounding the macrochannel is mainly used for agricultural and industrial purposes, with mining activities being the most prominent. methods various researchers (vannote et al. 1980; ward & stanford 1983; naiman et al. 1988; townsend 1989) indicated that riparian vegetation changes downstream from the origin of a river system, as the macro-channel develops and changes. in this study, it was decided to use a spatial scale of 1:250 000, in order to include the olifants river system from its origin to the western border of the kruger national park. geological data were the most practical empirical data set available at this particular scale and were myburgh & bredenkamp.qxd 2004/04/08 01:57 page 41 therefore used as baseline template to stratify the landscapes through which the olifants river flows. the width of the riparian zone varies in correspondence to the slope and height of the banks at any given locality (nilsson et al. 1994). an area-based vegetation sampling method was used to survey the vegetation. variable-sized belt transects were placed within the stratified units. the number of sample plots placed in each stratified unit was calculated proportionally depending on the length of the unit. the width and length of the belt transects were influenced by the width of the channel bank and the species richness and growth forms present at a specific site. floristic and habitat data were collected at 18 geo-referenced localities representing 36 relevés (>200 m²), since both macro-channel banks were surveyed as separate relevés. the floristic data recorded at each of these relevés included species composition, growth forms and canopy cover using the plant number scale (westfall & panagos 1988). the floristic data were analysed using the phytotab-pc computer program package (westfall 1997). results and discussion a total of eight clearly recognisable plant communities, representative of the vegetation of this section of the macro-channel of the olifants river system (myburgh 2000, 2001), were identified, mapped and described koedoe 47/1 (2004) 42 issn 0075-6458 fig. 1. distribution of the plant communities of the olifants river associated with the grassland biome. myburgh & bredenkamp.qxd 2004/04/08 01:57 page 42 at a spatial scale of 1:250 000 (fig. 1, table 1). 1. the eragrostis plana-monopsis decipiens grassland the eragrostis plana-monopsis decipiens grassland is represented by two relevés only. this grassland community represents primarily the grassland along the macro-channel banks of that section of the olifants river that stretches from its origin to the nooitgedacht farm (fig. 1). the distribution is limited to the areas associated with the ea land type (land type series 1985a), which are characterised by rocks from the karoo sequence, at an altitude of about 1 750 m a.s.l. the soil depth at the top of the banks varies between 600 mm and 640 mm. this section of the river has a single active channel in the macro-channel floor that is 1–6 m wide (fig. 2). the tops of the banks are flat, with a slope gradient of ≤1º, and there is no clear riparian zone. the vegetation on top of the banks may be described as typically terrestrial eragrostis plana grassland and it cannot be visually distinguished from the surrounding grassland. there are no issn 0075-6458 43 koedoe 47/1 (2004) table 1 mean canopy cover (m.c.c) of the plant communities identified in the grassland biome section of the olifants river plant mean canopy cover (%) total community trees shrubs dwarf forbs grasses m.c.c. number shrubs (%) 1 3 30 33 2 10 32 42 3 7 4 26 37 4 <1 <1 5 50 56 5 1 5 33 39 6 <1 4 2 9 23 38 7.1 2 <1 6 39 48 7.2 12 9 2 21 24 68 8 4 <1 <1 12 46 62 fig. 2. river profile of the eragrostis plana-monopsis decipiens grassland myburgh & bredenkamp.qxd 2004/04/08 01:57 page 43 stones or rocks in the channel bed or on the banks. this grassland is characterised by two diagnostic species (table 2). on average, the canopy cover of both the forb monopsis decipiens and the sedge cyperus esculentus is <1 %. dominant grasses are eragrostis plana, eragrostis curvula and cynodon dactylon. the total average canopy cover of this grassland is 33 % (table 1). 2. the themeda triandra-fingerhuthia sesleriiformis grassland the themeda triandra-fingerhuthia sesleriiformis grassland is represented by four relevés and describes the floristics of both the macro-channel banks and the vegetation of the surrounding terrestrial grassland. as is the case with plant community 1, this plant community (fig. 1) is associated with the ea koedoe 47/1 (2004) 44 issn 0075-6458 table 2 diagnostic species of the eragrostis plana-monopsis decipiens grassland (m.c.c. mean canopy cover) species growth consistency m.c.c. name form (%) (%) monopsis decipiens forb 100 <1 cyperus esculentus grass 50 <1 land type (land type series 1985a) at an altitude of about 1 750 m a.s.l., but here the rock is typically dolerite. the soil depth varies between 700 mm and 1 000 mm, and the soil has a high clay content throughout (>55 % clay). this section of the olifants river has a single active channel that is < 17 m wide (fig. 3). the river has cut into the landscape and consequently has steep macro-channel banks of about 2–3 m high. the vegetation above the macro-channel banks may be described as typical terrestrial themeda triandra grassland on flat plains. the presence of large, isolated sheets of rock characterises parts of the active channel bed. the grass species, fingerhuthia sesleriiformis, brachiaria eruciformis, aristida bipartita, and the forb species, hemizygia sp., haplocarpha scaposa, falckia oblonga, ledebouria sp., anthospermum pumilum subsp. rigidum, scabiosa columbaria and jamesbrittenia montana are diagnostic for this community (table 3). grass species, such as themeda triandra and eragrostis plana, and the forb haplocarpha scaposa are dominant and have the highest average canopy cover. these plant species also occur in the adjacent terrestrial veld. fingerhuthia sesleriiformis, brachiaria eruciformis, cirsium vulgare, tagetes minuta and cyperus longus var. tenuiflorus are mostly limited to fig. 3. river profile of the themeda triandra-fingerhuthia sesleriiformis grassland. myburgh & bredenkamp.qxd 2004/04/08 01:57 page 44 the macro-channel banks and active channel bed. a woody component is absent in this grassland community. the total average canopy cover of the themeda triandra-fingerhuthia sesleriiformis grassland is 42 % (table 1). 3. the clutia natalensis-panicum dregeanum grassland the clutia natalensis-panicum dregeanum grassland is represented by two relevés and occurs on both the macro-channel banks and the terrestrial area above the banks. this plant community is situated in the vicinity of the frischgewaagd farm (fig. 1). it is a grassland community found where the olifants river runs through the bb land type (land type series 1985a), associated with the karoo sequence and aeolic sand at an altitude of about 1 650 m a.s.l. the deep (≥ 1 200 mm) soil on the macro-channel banks is typically a sand-deposit covering clayey subsoil (>55 % clay). issn 0075-6458 45 koedoe 47/1 (2004) table 3 diagnostic spesies of the themeda triandra-fingerhuthia sesleriiformis grassland (m.c.c. mean canopy cover) species name growth form consistency m.c.c. (%) (%) fingerhuthia sesleriiformis grass 100 3 hemizygia sp. forb 100 <1 haplocarpha scaposa forb 75 3 falcforbia oblonga forb 75 1 ledebouria sp. forb 75 <1 brachiaria eruciformis grass 75 1 anthospermum pumilum subsp. rigidum forb 50 <1 scabiosa columbaria forb 50 >1 jamesbrittenia montana forb 50 <1 aristida bipartita grass 50 2 fig. 4. river profile of the clutia natalensis-panicum dregeanum grassland. myburgh & bredenkamp.qxd 2004/04/08 01:57 page 45 the macro-channel in this section of the river is, on average, 14 m wide and the steep macro-channel banks are similar to the themeda triandra-fingerhuthia sesleriiformis grassland (fig. 4). the surrounding grassland above the banks is situated on flat to convex plains, with a clear riparian zone limited to the macro-channel banks. this grassland community is characterised by the presence of a woody component, with the dwarf shrub species clutia natalensis, artemisia afra and gomphocarpus fruticosus limited to the steep macro-channel banks, while the reed phragmites australis and forb persicaria lapathifolia are found at the waterside in the active channel. koedoe 47/1 (2004) 46 issn 0075-6458 table 4 diagnostic species of the clutia natalensis-panicum dregeanum grassland (m.c.c. mean canopy cover) species name growth consistency m.c.c. form (%) (%) panicum dregeanum grass 100 2 artemisia afra dwarf 100 7 shrub oenothera erythrosepala forb 100 <1 the grass panicum dregeanum, forb oenothera erythrosepala, and dwarf shrub artemisia afra are diagnostic for this community (table 4). dominant dwarf shrub species are clutia natalensis and artemisia afra, while themeda triandra, eragrostis curvula, setaria sphacelata var. sphacelata, andropogon appendiculatus and hyparrhenia tamba mostly dominate the grass layer. the total average canopy cover of this grassland is 37 % (table 1). 4. the heteropogon contortus-cyperus longus var. tenuiflorus grassland the heteropogon contortus-cyperus longus var. tenuiflorus grassland is represented by seven relevés and occurs on both the steep macro-channel banks and the terrestrial area above the banks. this plant community (fig. 1) is associated with that section of the olifants river that is situated in the bb land type (land type series 1985a) with sandstones and shales from the karoo sequence. deep soil (1 000–1 200 mm) is common and the soil texture varies from sandy loam (11 % to 15 % clay) to clayey soil (>55 % clay). the width of the macro-channel varies from 15–27 m (fig. 5). the river has cut deeply into the landscape, and steep macro-channel fig. 5. river profile of the heteropogon contortus-cyperus longus var. tenuiflorus grassland. myburgh & bredenkamp.qxd 2004/04/08 01:57 page 46 banks, up to 3.5 m high, are common. the landscape above the banks is flat to convex and has slopes with gradients of 1–8 º. riparian vegetation is limited to the macrochannel banks. woody vegetation is mostly limited to rocky areas along the macro-channel banks. the active channel bed has alluvial deposits or sand banks (depositional bars), and large rocks are quite common on both the macrochannel banks and in the active channel. this section of the active channel is characterised by pools, and a few rapids. in certain areas mining activities have seriously disturbed the adjacent terrestrial vegetation, while planted pastures that stretch down to the macro-channel replaced the natural grassland. diagnostic species are, inter alia, the grasses heteropogon contortus, eragrostis gummiflua, elionurus muticus, alloteropsis semialata, aristida congesta subsp. congesta, aristida sp., the forbs bidens formosa, acalypha caperonoides, and the fern pteridium aquilinum (table 5). the grass layer is completely dominated by themeda triandra, followed by cynodon dactylon, aristida sp., patches of phragmites australis and hyparrhenia tamba. dominant sedges are cyperus longus var. tenuiflorus and cyperus latiissn 0075-6458 47 koedoe 47/1 (2004) table 5 diagnostic species of the heteropogon contortuscyperus longus var. tenuiflorus grassland (m.c.c. mean canopy cover) species name growth constancy m.c.c. form (%) (%) heteropogon contortus grass 71 2 eragrostis gummiflua grass 71 <1 elionurus muticus grass 43 <1 bidens formosa forb 43 <1 alloteropsis semialata grass 29 <1 acalypha caperonioides forb 29 <1 aristida congesta subsp. congesta grass 29 <1 aristida sp. grass 29 3 pteridium aquilinum forb 29 <1 folius. these species, and phragmites australis, are, however, limited to the waterside and alluvial deposits (depositional bars) in the active channel. the grasses brachiaria brizantha, hemarthria altissima and the forb persicaria lapathifolia are also associated with the lower-lying, moister sections of the macro-channel banks. the shrub component is represented by rubus fruticosus, rhus gerrardii and diospyros lycioides subsp. sericea. the latter shrub is associated with stony and rocky substrates, similar to the surrounding terrestrial bankenveld substrates. the average canopy cover of these three species is less than 1 %, while the total average canopy cover of the heteropogon contortus-cyperus longus var. tenuiflorus grassland is 56 % (table 1). 5. the eragrostis plana-cyperus fastigiatus grassland the eragrostis plana-cyperus fastigiatus grassland is represented by five relevés and occurs both on the macro-channel banks and the terrestrial area above the banks. this plant community is associated with the olifants river in the vicinity of the middelkraal farm and it also occurs in the vicinity of the vandyksdrift farm (fig. 1). this grassland is associated with the bb and fa land types (land type series 1985a) at an altitude of about 1 600 m a.s.l. typical underlying rocks are, inter alia, sandstone, shale, layered mudstone and rhyolite from the karoo and transvaal sequences. the soil depth varies (1 000–1 200 mm), and the soil texture ranges from sandy clay loam to loamy clays throughout this community. soil texture varies from 21 % to 55 % clay. the macro-channel of this section of the olifants river is, on average, 16 m wide with a single active channel (fig. 6). the river typically has steep macro-channel banks. above ground rock occurs only in a few isolated areas, either on the banks or in the active channel. locally, alluvial deposits or sand banks (depositional bars) are found in the active channel. myburgh & bredenkamp.qxd 2004/04/08 01:57 page 47 the grass bromus catharticus and forbs cirsium vulgare and crabbea acaulis are diagnostic for this grassland community (table 6). the grasses eragrostis planiculmis, brachiaria brizantha, bromus cartharticus, the sedge cyperus fastigiatus and the forbs rumex crispus and persicaria lapathifolia are dominant on the steep macro-channel banks and at the waterside. the grass eragrostis plana and to a lesser degree eragrostis curvula and themeda triandra are dominant above the banks. only three dwarf shrub species are found in this community and clutia natalensis has the highest average canopy cover. the dwarf shrubs gomphostigma virgatum and gomphocarpus physocarpus were recorded only in one relevé and are poorly represented. the average total canopy cover of the eragrostis plana-cyperus fastigiatus grassland is 39 % (table 1). 6. the rhus gerrardii-hemarthria altissima grassland the rhus gerrardii-hemarthria altissima grassland is, as is the case with the eragrostis plana-cyperus fastigiatus grassland (community 5), associated with the bb and fa land types (land type series 1985a), but it occurs at an altitude of about 1 550 m a.s.l. (fig. 1). the karoo and transvaal sequences represent the geology of this area. soil depth varies from 600 mm to 1 200 mm with a clay content of higher than 55 %. this vegetation is represented by six relevés. koedoe 47/1 (2004) 48 issn 0075-6458 fig. 6. river profile of the eragrostis plana-cyperus fastigiatus grassland. table 6 diagnostic species of the eragrostis plana-cyperus fastigiatus grassland (m.c.c. mean canopy cover) species name growth constancy m.c.c. form (%) (%) bromus catharticus grass 40 <1 cirsium vulgare forb 60 <1 crabbea acaulis forb 40 <1 the macro-channel has a single active channel varying between 19 m and 22 m in width, with pools and rapids succeeding each other (fig. 7). the river flows through rocky ridges, consequently the above-ground rock cover is high in both the active channel bed and on the macro-channel banks. stone sizes vary from large stones (> 250 mm) to boulders (>1 000 mm). it may be assumed that this high rock cover would play a stabilising role on both the macro-channel banks and in the active channel bed, and that it would myburgh & bredenkamp.qxd 2004/04/08 01:57 page 48 influence the water flow pattern, and consequently, that this section of the river would not be influenced as much as other sections by floods. the riparian zone is characterised by the presence of a shrub component. the exotic, though naturalised, tree salix babylonica and forbs galium capense subsp. garipense, argyrolobium tuberosum, helichrysum mundtii and polygala hottentotta are diagnostic for this grassland community (table 7). the dwarf shrubs, gomphostigma virgatum and gomphocarpus physocarpus, the grasses, miscanthus junceus, brachiaria brizantha and hemarthria altissima and the forb persicaria lapathifolia are dominant at the waterside and in the lower-lying parts of the macro-channel bank. rhus gerrardii is the only shrub recorded. it has an average canopy cover of 3.91 %. the tree salix babylonica was recorded at only two sites. it has an average canopy cover of issn 0075-6458 49 koedoe 47/1 (2004) fig. 7. river profile of the rhus gerrardii-hemarthria altissima grassland. table 7 diagnostic species of the rhus gerrardiihemarthria altissima grassland (m.c.c. mean canopy cover) species name growth constancy m.c.c. form (%) (%) galium capense subsp. garipense forb 50 2 argyrolobium tuberosum forb 67 <1 helichrysum mundtii forb 33 <1 polygala hottentotta forb 33 <1 salix babylonica tree 33 <1 < 1 %. digitaria eriantha, brachiaria brizantha, eragrostis curvula, hemarthria altissima, miscanthus junceus, themeda triandra and eragrostis plana are dominant in the grass stratum while the forb species galium capense subsp. garipense, persicaria lapathifolia and tagetes minuta are commonly found. the average total canopy cover of the rhus gerrardii-hemarthria altissima grassland is 38 % (table 1). 7. the salix mucronata subsp. wilmsiieragrostis curvula grassland the salix mucronata subsp. wilmsii-eragrostis curvula grassland is represented by six relevés. this plant community is associated with the bb land type (land type series 1985b) at altitudes ranging between myburgh & bredenkamp.qxd 2004/04/08 01:57 page 49 1 425 m and 1 450 m a.s.l. this grassland is limited to narrow, easily distinguishable diabase dykes and intrusions crossing the olifants river, immediately north of the doringpoort dam, in the vicinity of the town witbank (fig. 1). the shrub salix mucronata subsp. wilmsii and the grass ischaemum fasciculatum are diagnostic for this community. the community is divided into two variants, namely the paspalum scrobiculatum-miscanthus junceus variant (variant 7.1) and the acacia dealbata-eragrostis curvula variant (variant 7.2). the most significant difference in the habitats of these two variants lies in soil depth and above ground rock cover. the paspalum scrobiculatum-miscanthus junceus variant is found on deep soil with a low rock cover, koedoe 47/1 (2004) 50 issn 0075-6458 fig. 8. river profile of the paspalum scrobiculatum-miscanthus junceus variant. table 8 diagnostic species of the paspalum scrobiculatummiscanthus junceus variant (m.c.c. mean canopy cover) species name growth constancy m.c.c. form (%) (%) paspalum scrobiculatum grass 100 2 chamaecrista comosa forb 100 <1 imperata cylindrica grass 50 7 plantago virginica forb 50 <1 ipomoea crassipes forb 50 <1 cynoglossum lanceolatum forb 50 <1 while the acacia dealbata-eragrostis curvula variant is found on shallow soil with a high rock cover. there is also a great difference in the vegetation structure of these two variants. the woody component of variant 7.1 is limited to a few dwarf shrub and shrub species with a low average canopy cover, in comparison with the woody component of variant 7.2, which consists of tree, shrub and dwarf shrub species. the tree acacia dealbata (a declared alien invader,) and shrub sesbania punicea (a declared weed), dominate the macro-channel banks of variant 7.2 and contribute largely to the total canopy cover of the woody component of this variant. however, these two alien problem species are not found in the paspalum scrobiculatum-miscanthus junceus variant. there is also a huge difference in the floristic composition between myburgh & bredenkamp.qxd 2004/04/08 01:57 page 50 these variants that are limited to the diabase intrusions. 7.1 paspalum scrobiculatum-miscanthus junceus variant the paspalum scrobiculatum-miscanthus junceus variant is represented by two relevés only, and the extent of this unit is limited. the macro-channel is up to 22 m wide and deep, stagnant pools of water are predominant (fig. 8). the macro-channel banks form flat to convex terraces and the above ground rock cover is limited to a few pebbles in the active channel bed. the sandy clay loam soil is deep (≥ 1 200 mm) and has a clay content of 21 % to 35 %. this variant has six diagnostic species (table 8), and the grass paspalum scrobiculatum and forb chamaecrista comosa, have been recorded only on the macro-channel banks. the shrub layer of this variant is dominated by salix mucronata subsp. wilmsii. the shrubs diospyros lycioides subsp. sericea and rhus gerrardii are poorly represented in this variant. hyparrhenia hirta, paspalum scrobiculatum, ischaemum fasciculatum, imperata cylindrica, setaria sphacelata var. sphacelata, hemarthria altissima and miscanthus junceus dominate the grass layer. the grass species miscanthus junceus together with the sedge species, cyperus latifolius, cyperus marginatus, cyperus longus var. tenuiflorus are limited to the lower-lying terraces on the macro-channel banks and the areas bordering the active channel. the variant represents the vegetation associated with the macro-channel bank as well as the vegetation above the macro-channel bank. there is no single dominant forb species. all the forb species present have an average canopy cover of less than 1 %. however, the joint, average canopy cover of the forbs is 6 %. forb species that make the largest contribution are crabbea acaulis, berkheya radula, ipomoea bathycolpos, ipomoea crassipes, helichrysum rugulosum and tagetes minuta. the total average canopy cover of the paspalum scrobiculatum-miscanthus junceus variant is 48 % (table 1). 7.2 acacia dealbata-eragrostis curvula variant the acacia dealbata-eragrostis curvula variant is represented by four relevés. the macro-channel divides in certain areas to form two active channels that are between 23 m and 25 m wide (fig. 9). these active channels are characterised by shallow pools issn 0075-6458 51 koedoe 47/1 (2004) fig. 9. river profile of the acacia dealbata-eragrostis curvula variant. myburgh & bredenkamp.qxd 2004/04/08 01:57 page 51 followed by rapids, and by the presence of a few islands on which woody plant species are found. the riparian zone is clearly visible with a strong woody component. the soil on the macro-channel banks is 150 mm to 200 mm deep and the soil texture varies from loamy sand (11 % -15 % clay) to clayey soil (>55 % clay). this section of the river is characterised by a high percentage rock cover. the stones vary from pebbles limited to the active channel bed to large stones and rocks that occur mostly on the macro-channel banks. this variant has 14 diagnostic species (table 9). the declared alien invader acacia dealbata and the declared alien weed sesbania punicea have the highest average canopy cover. these species should be removed, since they both grow near the waterside and are a source of seed that may spread to downstream areas. the acacia dealbata-eragrostis curvula variant is characterised by the presence of a strong woody component. dominant woody species are the tree acacia dealbata, the shrubs sesbania punicea, diospyros lycioides subsp. sericea, salix mucronata subsp. wilmsii, rhus gerrardii, and the dwarf koedoe 47/1 (2004) 52 issn 0075-6458 table 9 diagnostic species of the acacia dealbata-eragrostis curvula variant (m.c.c. mean canopy cover) species name growth constancy m.c.c. form (%) (%) acacia dealbata tree 75 12 sesbania punicea shrub 100 2 deverra sp. forb 75 <1 acacia karroo dwarf shrub 75 <1 bidens pilosa forb 50 2 dicliptera clinopodia forb 75 1 lepidium virginicum forb 75 <1 ranunculus multifidus forb 50 <1 sida rhombifolia dwarf shrub 50 <1 vangueria cyanescens shrub 50 <1 maytenus heterophylla shrub 50 <1 chenopodium ambrosioides forb 50 <1 typha capensis grass 50 <1 achyranthes aspera var. aspera forb 50 <1 shrubs clutia natalensis and artemisia afra. the grasses eragrostis plana, miscanthus junceus, hemarthria altissima, eragrostis curvula and paspalum distichum, and the sedges cyperus marginatus and cyperus latifolius dominate the grass layer. the grass species miscanthus junceus, hemarthria altissima, paspalum distichum, phragmites australis, and sedge species cyperus marginatus and cyperus latifolius are mostly limited to the lower-lying parts of the macro-channel bank close to the active channels. the macro-channel banks are disturbed in local areas. the high average canopy cover of the opportunistic weedy forbs tagetes minuta, verbena bonariensis and bidens pilosa confirms that disturbance. the high weed cover probably results from the high woody cover and its shadow effect—the alien tree acacia dealbata, with an average canopy cover of 12 %, being the greatest contributor. the negative shadow effect is clearly visible on the grass layer. the total average canopy cover of the acacia dealbata-eragrostis curvula variant is 68 % (table 1). 8. the echinochloa crus-galli-paspalum distichum grassland the echinochloa crus-galli-paspalum distichum grassland is represented by four relevés and it occurs south of witbank dam in the vicinity of the wolwekrans farm and downstream of the doringpoort dam (fig. 1). the plant community is associated with the bb land type (land type series 1985b) and is found at altitudes of 1 450 m to 1 500 m a.s.l. the typical underlying geology is derived from the loskop formation and the karoo sequence, group ecca, and mostly consists of sandstone, shale and conglomerate. the soil is mostly 1 200 mm deep and varies from sandy clay loam (21 %– 35 % clay) to heavy clayey soil (>55 % clay). myburgh & bredenkamp.qxd 2004/04/08 01:57 page 52 the width of the macro-channel varies between 26 m and 32 m with steep banks up to 4 m high (fig. 10). a single active channel has a rocky appearance in certain areas, and divides to form two active channels with an island covered by a dwarf shrub component. the grasses echinochloa crus-galli and echinochloa jubata, the sedge juncus effusus, the forb cotula anthemoides and the alien tree morus alba are diagnostic for this grassland community (table 10). the tree component of this community is represented by acacia dealbata, morus alba and rhus lancea. however, the latter two species have low average canopy covers. the woody species have a low constancy throughout. the grasses eragrostis plana, phragmites australis, setaria sphacelata var. sphacelata, echinochloa jubata, cyperus latifolius and paspalum distichum and the sedge cyperus fastigiatus dominate the grass layer, while verbena bonariensis, persicaria lapathifolia issn 0075-6458 53 koedoe 47/1 (2004) fig. 10. river profile of the echinochloa crus-galli-paspalum distichum grassland. table 10 diagnostic species of the echinochloa crus-gallipaspalum distichum grassland (m.c.c. mean canopy cover) species name growth constancy m.c.c. form (%) (%) echinochloa crus-galli grass 75 <1 echinochloa jubata grass 50 10 juncus effusus grass 50 <1 morus alba tree 50 <1 cotula anthemoides forb 50 <1 and tagetes minuta dominate the forb layer. the sedges cyperus latifolius and cyperus fastigiatus, the reed, phragmites australis, the grasses echinochloa jubata, paspalum distichum and hemarthria altissima and the forb persicaria lapathifolia are mostly limited to the watersides of the macro-channels. phragmites australis is also associated with the active channel bed and islands. the total average canopy cover of the echinochloa crus-galli-paspalum distichum grassland is 62 %, with grass species making the largest contribution (table 1). myburgh & bredenkamp.qxd 2004/04/08 01:57 page 53 discussion and conclusion the spatial scale of 1:250 000 at which the stratification and sampling took place, caused the vegetation on the macro-channel banks and the vegetation on top of the banks to be seen and described as a single unit. the species composition, representing both macro channel and terrestrial vegetation, therefore, may seem strange, unless it is taken into account that specific plant species are associated with the lower-lying sections bordering the waterside and that other plant species are associated with the banks or even on top of the banks. the locality and distribution of the species were however recorded and described. the riparian vegetation of the macro-channel in the grassland biome is not clearly distinguishable from the surrounding vegetation and is mainly represented by terrestrial grass species. a few indigenous woody species occur in the rocky areas associated with the macro-channel. floristic differences occur within and between communities due to changes in habitat but also as a result of different land use practices and intensities of utilisation. in certain areas the river was diverted due to mining activities and the natural vegetation replaced with planted pastures. the alteration of riverbanks and the occurrence of exotic woody species in certain areas modify the stabilising role of the indigenous vegetation and influence hydrological processes. acacia dealbata and the declared alien weed sesbania punicea pose a threat to the biodiversity of the macro-channel and should be removed. these species grow near the waterside and are a source of seed, spreading to the downstream areas. although time consuming and costly, it is important to obtain geo-referenced baseline data for these longitudinally narrow ecosystems, in order to manage and monitor vegetation change over time. acknowledgements this project was made possible by funds donated by the water research commission and the use of the facilities at the agricultural research council. references acocks, j.p.h. 1988. veld types of southern africa. (3rd. ed) memoirs of the botanical survey of south africa 57:1–146. land type series. 1985a. land type map 2628 east rand (1:250 000). pretoria: government printer. land type series. 1985b. land type map 2528 pretoria (1:250 000). pretoria: government printer. myburgh, w.j. 2000. oewerplantegroei van die olifantsriviersisteem—'n ekologiese perspektief. water research commission. isbn no 186845 5688. myburgh, w.j. 2001. die ekologie en plantegroeibestuur van die olifantsriviersisteem. phd thesis, university of pretoria, pretoria. naiman, r.j., h. decamps, j. pastor & a.c. johnston. 1988. the potential importance of boundaries to fluvial ecosystems. journal of the north american benthological society 7:289–306. nillson, c., a. ekblad, m. dynesius, s. backe, m. gardfjell, b. carlberg, s. hellqvist & r. jansson. 1994. a comparison of species richness and traits of riparian plants between a main river channel and its tributaries. journal of ecology 82:281–295. olifants river forum. 1995. save water—or face wars. on track (october/november): 60–61. rutherford, m.c. & r.h. westfall. 1986. biomes of southern africa—an objective categorization. memoirs of the botanical survey of south africa 54: 45–65. townsend, c.r. 1989. the patch dynamics concept of stream community ecology. journal of the north american benthological society 8:36–50. vannote, r.l., g.w. minshall, k.w. cummins, j.r. sedell & c.e. cushing. 1980. the river continuum concept. canadian journal of fisheries and aquatic science 37:130–137. ward, j.v. & j.a. stanford. 1987. the ecology of regulated streams: past accomplishments and directions for future research. pp. 391–409. in: craig, j.f. & j.b. kemper (eds.). regulated streams. advances in ecology. new york: plenum press. westfall, r.h. & m.d. panagos. 1988. the plant number scale—an improved method for cover estimation using variable belt transects. bothalia 18(2): 289–291. westfall, r.h., g.k. theron & n. van rooyen. 1997. objective classification and analysis of vegetation data. plant ecology 132: 137–154. koedoe 47/1 (2004) 54 issn 0075-6458 myburgh & bredenkamp.qxd 2004/04/08 01:57 page 54 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 foxcroft.qxd a revised list of alien plants for the kruger national park l.c. foxcroft, l. henderson, g. r. nichols and b.w. martin foxcroft, l.c., l. henderson, g. r. nichols and b.w. martin. 2003. a revised list of alien plants for the kruger national park. koedoe 46(2): 21-44. pretoria. issn 00756458. alien plants recorded in the kruger national park are listed, providing an update of species that have been recorded since the last published list in 1988. the serious consequences that invasive alien plants pose are widely recognised internationally and are regarded as the greatest threat to the kruger national park. it is extremely important to maintain an updated list of species to use as a reference and from which to monitor changes in the influx of invasive species. the list contains the invasive status of the plants, current impact, habitat types, biological control status and status according to the conservation of agricultural resources act (cara), act 43 of 1983, as amended in march 2001. the list comprises 370 species, of which 121 are invaders and two are transformer weeds, which may cause considerable damage and alterations to the indigenous biodiversity. a total of 91 species listed herein are also listed in the cara regulations or are proposed additions to the cara regulations. key words: alien plants, invasive, species list, status, kruger national park. l.c. foxcroft and b.w. martin, invasive alien species section, conservation services, kruger national park, private bag x 402, skukuza, 1350; l. henderson, arc-plant protection research institute, national botanical institute, private bag x 101, pretoria, 0001; g. r. nichols, geoff nichols horticultural services, 8 larch road, durban, 4001 republic of south africa. issn 0075-6458 21 koedoe 46/2 (2003) introduction the species list in this paper contains reference to lists previously published by a number of authors of alien plant species recorded in the kruger national park (knp). the first published list was by obermeijer (1937), who listed six alien plant species ("troublesome weeds": chenopodium ambrosioides l., tagetes minuta l., argemone mexicana l., gomphrena celosiodes mart., boerhavia diffusa l., and cocculus hirsutus (l.) diels. (foxcroft & richardson 2003), thereafter codd (1951) indicated 32 species, van der schijff (1957 & 1969) indicated 43 and 76 species, respectively, and macdonald & gertenbach (1988) listed 150 alien plant species. between 1988 and 1997, the species list increased to 216 species (knp alien plant section records). in february 1999, nichols and foxcroft undertook a survey in the skukuza personnel village listing approximately 243 species. the number of alien plant species in the knp then totalled approximately 360 species, prompting a revision of the knp alien plant list to provide a more accurate reflection of the records. figure 1 indicates the increase in alien plant species in the knp. the long-term average increase in alien plant species, since the first record of alien plants was made in 1937, is 5.6 alien species per year. thirteen species have been removed from the list presented by macdonald & gertenbach (1988), as information now indicates that these species are probably indigenous (appendix 1). presentation of the list families and species, respectively, are listed alphabetically within each of the major plant groupings of pteridophyta (ferns and fern allies), gymnosperms, angiosperms (monocotyledons and dicotyledons). foxcroft.qxd 2005/12/09 10:40 page 21 the list further provides information on the status of the weed and its current impact in the knp, as well as its biological control and legal status for south africa. the date provided is that of the first record for the species or when the species was first listed in publications. ‘m&g’ is added to the date column to indicate plants listed in the last published list (macdonald & gertenbach 1988). all plants not indigenous to the knp lowveld have been listed in a manner similar to macdonald & gertenbach (1988). weed status (as is currently observed in the knp) is described using the terminology proposed by richardson et al. (2000): alien plants: plant taxa in a given area whose presence there is due to intentional or accidental introduction as a result of human activity (synonyms: non-native plants, non-indigenous plants). casual alien plants: alien plants that may flourish and even reproduce occasionally in an area, but which do not form self-replacing populations and which rely on repeated introductions for their persistence (denoted by c). naturalised plants: alien plants that reproduce consistently and sustain populations over many life cycles without direct intervention by humans (or in spite of human intervention); they often recruit offspring freely, usually close to adult plants, and do not necessarily invade natural, semi-natural or human-made ecosystems (denoted by n). invasive plants: naturalised plants that produce reproductive offspring, often in very large numbers, at considerable distances from parent plants and thus have the potential to spread over a considerable area (denoted by i). where the species is considered as a potential invader, pi is added. transformers: a subset of invasive plants which change the character, condition, form or nature of ecosystems over a substantial area relative to koedoe 46/2 (2003) 22 issn 0075-6458 fig. 1: the number of invasive alien plant species recorded in the kruger national park between 1930 and 2001. the list includes species listed according to published and other lists, as well as according to dates of collections/observations of species. for published and other species lists, the authors are indicated: 1. obermeijer (1937); 2. codd (1951); 3. van der schijff (1957); 4. van der schijff (1969); 5. gertenbach (1985); 6. macdonald & gertenbach (1988); 7. anon (1995); 8. anon (1996); 9. foxcroft (1999); 10. foxcroft (2000); 11. foxcroft (2001). foxcroft.qxd 2005/12/09 10:40 page 22 the extent of that ecosystem (denoted by t). where the species is considered as a potential transformer, pt is added. impact on the knp is noted as the current known impact, and rated as high (h), moderate (m) or low (l). this information is an estimate based on records of the species in the knp and the authors' personal experience. the habitat types where the plants are currently observed in the knp are: disturbed areas and roadsides denoted by the symbol (dr), villages and rest camps (v), aquatic habitats (a), riparian/riverine (r) and terrestrial/dryland habitats (td). status of biological control is indicated at two levels; that taken from the national biocontrol list (olckers & hill 1999); and, that for the knp (martin & foxcroft 2001) indicated by (*). legal status refers to the recently revised regulation 15 in terms of the conservation of agricultural resources act, 1983 (act no. 43 of 1983): dw1: declared weed (category 1) are prohibited plants which must be controlled, or eradicated where possible (except in biocontrol reserves, which are areas designated for the breeding of biocontrol agents); di2: declared invader (category 2) are mainly commercial plantation species but also plants for woodlots, animal fodder, soil stabilisation, etc., allowed only in demarcated areas, by permit holders, under controlled conditions and in biocontrol reserves; di3: declared invader (category 3) are mainly ornamental species that may no longer be planted (except with special written permission), nor may there be trade in propagative material. existing plants may remain but must be prevented from spreading. all three categories of plants are prohibited within 30 m of the 1:50 year floodline of watercourses or wetlands, unless exemption is obtained. the full regulations and species list are given in henderson (2001). discussion a total of 370 alien plant species are listed and contain some subspecies, forms and varieties. of the list, 2 species (0.5 %) (lantana camara l. and opuntia stricta (haworth.) haworth.) are transformers, 125 (33.8 %) are invasive and 223 (60.2 %) are either casual aliens or naturalised species, as issn 0075-6458 23 koedoe 46/2 (2003) is indicated by fig. 2. the basic under-riding principle of the south african national parks (sanparks), "to preserve biodiversity in all its natural facets and fluxes" (braack 1997) is directly violated by the introduction, whether intentional or accidental, of alien organisms, and according to definition, indicates the requirement to remove or control all alien species. in practice, the control of prioritised transformer and invader species, to acceptable limits of abundance, is the most likely scenario. at this minimal level of abundance, the impact on biodiversity (as described by noss 1990) is hoped to be minimal. macdonald & gertenbach (1988) stated that the increase in number of alien plant species recorded for the knp was probably due to increased collecting and awareness between 1937 and 1983. further increase may also be ascribed to increased tourism (macdonald 1988), which has unintentionally brought in seeds from other infested parts of the country as well as increased infestation of the lowveld. infestations and new species may also have arisen due to the considerable increase in size of towns such as nelspruit and others along the escarpment, over the past 50 years. although macdonald & gertenbach (1988) state that 10 species had been eradicated, at least four of these (senna didymobotrya, fig. 2: weed status of alien plant species. foxcroft.qxd 2005/12/09 10:40 page 23 koedoe 46/2 (2003) 24 issn 0075-6458 ta bl e 1 r ev is ed a lie n pl an t l is t f or th e k ru ge r n at io na l p ar k foxcroft.qxd 2005/12/09 10:40 page 24 issn 0075-6458 25 koedoe 46/2 (2003) ta bl e 1 (c on tin ue d) o rd er a nd f am ili es sp ec ie s c om m on n am e d at e of sa tu s & h ab ita t t yp e b io l eg al 1s t r ec or d im pa ct co nt ro l st at us foxcroft.qxd 2005/12/09 10:40 page 25 koedoe 46/2 (2003) 26 issn 0075-6458 ta bl e 1 (c on tin ue d) o rd er a nd f am ili es sp ec ie s c om m on n am e d at e of sa tu s & h ab ita t t yp e b io l eg al 1s t r ec or d im pa ct co nt ro l st at us foxcroft.qxd 2005/12/09 10:40 page 26 issn 0075-6458 27 koedoe 46/2 (2003) ta bl e 1 (c on tin ue d) o rd er a nd f am ili es sp ec ie s c om m on n am e d at e of sa tu s & h ab ita t t yp e b io l eg al 1s t r ec or d im pa ct co nt ro l st at us foxcroft.qxd 2005/12/09 10:40 page 27 koedoe 46/2 (2003) 28 issn 0075-6458 ta bl e 1 (c on tin ue d) o rd er a nd f am ili es sp ec ie s c om m on n am e d at e of sa tu s & h ab ita t t yp e b io l eg al 1s t r ec or d im pa ct co nt ro l st at us foxcroft.qxd 2005/12/09 10:40 page 28 issn 0075-6458 29 koedoe 46/2 (2003) ta bl e 1 (c on tin ue d) o rd er a nd f am ili es sp ec ie s c om m on n am e d at e of sa tu s & h ab ita t t yp e b io l eg al 1s t r ec or d im pa ct co nt ro l st at us foxcroft.qxd 2005/12/09 10:40 page 29 koedoe 46/2 (2003) 30 issn 0075-6458 ta bl e 1 (c on tin ue d) o rd er a nd f am ili es sp ec ie s c om m on n am e d at e of sa tu s & h ab ita t t yp e b io l eg al 1s t r ec or d im pa ct co nt ro l st at us foxcroft.qxd 2005/12/09 10:41 page 30 issn 0075-6458 31 koedoe 46/2 (2003) ta bl e 1 (c on tin ue d) o rd er a nd f am ili es sp ec ie s c om m on n am e d at e of sa tu s & h ab ita t t yp e b io l eg al 1s t r ec or d im pa ct co nt ro l st at us foxcroft.qxd 2005/12/09 10:41 page 31 koedoe 46/2 (2003) 32 issn 0075-6458 ta bl e 1 (c on tin ue d) o rd er a nd f am ili es sp ec ie s c om m on n am e d at e of sa tu s & h ab ita t t yp e b io l eg al 1s t r ec or d im pa ct co nt ro l st at us foxcroft.qxd 2005/12/09 10:41 page 32 issn 0075-6458 33 koedoe 46/2 (2003) ta bl e 1 (c on tin ue d) o rd er a nd f am ili es sp ec ie s c om m on n am e d at e of sa tu s & h ab ita t t yp e b io l eg al 1s t r ec or d im pa ct co nt ro l st at us foxcroft.qxd 2005/12/09 10:41 page 33 koedoe 46/2 (2003) 34 issn 0075-6458 ta bl e 1 (c on tin ue d) o rd er a nd f am ili es sp ec ie s c om m on n am e d at e of sa tu s & h ab ita t t yp e b io l eg al 1s t r ec or d im pa ct co nt ro l st at us foxcroft.qxd 2005/12/09 10:41 page 34 issn 0075-6458 35 koedoe 46/2 (2003) ta bl e 1 (c on tin ue d) o rd er a nd f am ili es sp ec ie s c om m on n am e d at e of sa tu s & h ab ita t t yp e b io l eg al 1s t r ec or d im pa ct co nt ro l st at us foxcroft.qxd 2005/12/09 10:41 page 35 koedoe 46/2 (2003) 36 issn 0075-6458 ta bl e 1 (c on tin ue d) o rd er a nd f am ili es sp ec ie s c om m on n am e d at e of sa tu s & h ab ita t t yp e b io l eg al 1s t r ec or d im pa ct co nt ro l st at us foxcroft.qxd 2005/12/09 10:42 page 36 issn 0075-6458 37 koedoe 46/2 (2003) ta bl e 1 (c on tin ue d) o rd er a nd f am ili es sp ec ie s c om m on n am e d at e of sa tu s & h ab ita t t yp e b io l eg al 1s t r ec or d im pa ct co nt ro l st at us foxcroft.qxd 2005/12/09 10:42 page 37 koedoe 46/2 (2003) 38 issn 0075-6458 ta bl e 1 (c on tin ue d) o rd er a nd f am ili es sp ec ie s c om m on n am e d at e of sa tu s & h ab ita t t yp e b io l eg al 1s t r ec or d im pa ct co nt ro l st at us foxcroft.qxd 2005/12/09 10:42 page 38 issn 0075-6458 39 koedoe 46/2 (2003) ta bl e 1 (c on tin ue d) o rd er a nd f am ili es sp ec ie s c om m on n am e d at e of sa tu s & h ab ita t t yp e b io l eg al 1s t r ec or d im pa ct co nt ro l st at us foxcroft.qxd 2005/12/09 10:42 page 39 koedoe 46/2 (2003) 40 issn 0075-6458 ta bl e 1 (c on tin ue d) c od es u se d ab ov e: d at e: f ir st r ec or d: m & g 1 98 8: p re vi ou s lis t st at us : a s in k n p cu re nt ly c = c as ua l; i = in va de r; n = n at ur al is ed ; p i = po te nt ia l i nv ad er ; p t = po te nt ia l t ra ns fo rm er ; pn = p ot en tia l t o be co m e na tu ra lis ed ; t = tr an sf or m er im pa ct : k n p cu rr en tly h = h ig h; m = m od er at e; l = lo w h ab ita tt yp e: a = aq ua tic ; d r = d is tu rb ed a re a/ r oa ds id e; r = r ip ar ia n/ ri ve ri ne ; t d = te rr es tr ia l / d ry la nd ; v = vi lla ge a nd r es tc am ps b io co nt ro l: c = c om pl et e; s = s ub st an tia l; n = n eg lig ab le ; u = u nk no w n; b io co nt ro l i m pl em en te d el se w he re in s a bu t s ta tu s in k n p un kn ow n; u i = un de r in ve st ig at io n le ga l s ta tu s: d w 1 = d ec la re d w ee d ca t 1 ; d i = de cl ar ed in va de r ca t 2 ; d i = de cl ar ed in va de r ca t 3 foxcroft.qxd 2005/12/09 10:42 page 40 issn 0075-6458 41 koedoe 46/2 (2003) nicotiana glauca, tecoma stans and ipomoea purpurea) are frequently found in the rivers. they have either re-invaded the knp or were never eradicated in the first instance. nicotiana glauca appears to have long-term cycles, where plants flourish along the olifants river and then for a number of years disappear. two more species are present despite having been reported as eradicated. salvinia molesta, first recorded in the knp in 1974 (joubert 1986) was chemically controlled over a long period, and despite being recorded as eradicated a number of times (joubert 1986) was reported in 1985 on the sabie river at lower sabie rest camp. biological control was released on the sabie river in 1992, and the infestation is now under complete biological control. the infestation recorded at mtshawu dam (near pretoriuskop) in 1974 had biological control released in 1995 (lotter 1997), but was sprayed chemically soon afterwards, and then aggressively controlled using herbicide from 1997 to 2000, with small isolated plants still occasionally being found. however, during 2000-2001 the plants re-infested the pool below the mtshawu dam wall and a decision was taken to release biological control agents on the infestation. eight months after releasing the biocontrol agent cyrtobagous salviniae below the mtshawu dam wall, the plant population crashed and remains clear of s. molesta. macdonald & gertenbach (1988) also reported that solanum mauritianum was first reported along the crocodile river and was either eradicated or later failed to be reported as present along the river by officials from skukuza. this indicates that, for a reserve the size of the knp, the likelihood of a species being eradicated is remote once listed, and it may appear in another area at a later stage. furthermore, the large infestations of most of these weeds on the boundaries of the knp make it unlikely that only a few plants will enter the knp and re-invasion by the plants is inevitable. evidence to prove the lag phase and our limited knowledge of the ability of alien plants to become invasive is borne by the listing of 43 species by macdonald & gertenbach (1988) as non-invasive. of these species, 17 are now well-known invasive species, while seven have since become invasive in the park. in a national park, which aims at the protection of indigenous biodiversity, policy should rather err on the side of caution. the removal of all alien species accompanied by the replacement thereof with indigenous species should be promoted. although a species may not, at a single point in time, exhibit a tendency to invade, the lack of knowledge of the species and the long phase that is often found to prelude an explosion in the distribution of invasive species, places most species in an unknown category. this may result in serious, often immense, insidious and irreversible problems at a later stage (mcneely 2000), as species enter a rapid growth phase. the habitat invaded indicates that the vast majority of the species occur, or occurred, in the staff villages or rest camps (320 species) and is of concern (fig. 3). disturbed/roadside areas are invaded by 40 species, terrestrial/dryland by 27 species, riparian by 191 species and aquatic areas are invaded by seven species. a notable example in the knp of a species intentionally introduced and then escaping into the surrounding areas is that of o. stricta. assumed to have been introduced into the knp during the 1950s, probably as an ornamental garden plant in fig. 3: number of alien plant species per habitat type (some species may occur in more than one habitat type) foxcroft.qxd 2005/12/09 10:42 page 41 koedoe 46/2 (2003) 42 issn 0075-6458 skukuza personnel village, o. stricta was reported by lotter & hoffmann (1998) to cover an area of 30 000 ha around skukuza. currently, the management area of o. stricta is approximately 66 000 hectares, with a number of outlying invasions in the tshokwane area, nyamundwa dam and near the mtshawu and sabie river confluence. opuntia stricta is also an example of the problems associated with poor record keeping and incorrect identification. thought to have been first recorded in the knp by obermeijer in 1953 (lotter & hoffmann 1998, cite zeller 1996), the first record of o. stricta being recorded by name was in the knp alien plant sections records in 1995. no indication was made of the observer or date. however, brynard & pienaar (1960) refer to experiments on o. vulgaris (= o. monacantha) in the skukuza area, and it is likely that these plants were in fact o. stricta. opuntia monacantha (= o. vulgaris) was however recorded by zeller (knp alien plant sections records, 1996) in orpen rest camp in 1992, and also by van der schijff (1957) at the komati river and shilova (mooiplaas ranger section). examination of the plant list indicates the total number of plant families represented is 98, of which eight families collectively contain 125 species (or 35 % of the total) (fig. 4). the other 90 families contribute to the remaining 245 species. impact ranking assigned to the species indicates that 87 % of the species are rated as having a low impact, 7 % medium and 4 % as having a high impact (fig. 5). although the perception may be created that invasive species therefore represent a minimal impact to the knp, the combined effects of only a few invasive species can have major consequences and result in dense stands that cover vast areas. conclusion invasive alien plants have been recognised as one of the greatest threats to the biodiversity of the knp (braack 1997). in order to address this, proactive work on the prevention of species reaching and infesting the knp is necessary. early detection is the single most important step in combating the plants. while still in the early stages of invasion, the plants may be removed before they are able to set seed and become established. fig. 4: families of alien plant species in nkp. fig. 5: impact of alien plant species in knp. foxcroft.qxd 2005/12/09 10:42 page 42 issn 0075-6458 43 koedoe 46/2 (2003) updated species lists provide a firm base from which to work and are necessary to determine new invasions in an area as well as the rate of invasion. it is however only a starting point as spatial data is needed to determine the areas invaded, the extent to which they are invaded and the densities of invasions, in order to make meaningful interpretations and recommendations. acknowledgements mrs. guin zambatis is thanked for identifying numerous plant specimens (over a number of years) for the alien plant section as well as for checking the plant list and adding valuable comments and corrections. my co-authors are thanked for the continued support with often tedious re-checking of the lists. references anon. 1995. alien biota section records. scientific services department, skukuza. unpublished records, south african national parks. anon. 1996. alien biota section records. scientific services department, skukuza. unpublished records, south african national parks. braack, l.e.o. 1997. vol vii. an objectives hierarchy for the management of the kruger national park. a revision of parts of the management plan for the kruger national park. http://www. parks-sa.co.za. codd, l.e.w. 1951. trees and shrubs of the kruger national park. memoirs of the botanical survey of south africa 26: 1-192. foxcroft, l.c. 1999. alien biota section records. scientific services department, skukuza. unpublished records, south african national parks. foxcroft, l.c. 2000. alien biota section records. scientific services department, skukuza. unpublished records, south african national parks. foxcroft, l.c. 2001. alien biota section records. scientific services department, skukuza. unpublished records, south african national parks. foxcroft, l.c. & d.m. richardson. 2003. managing alien plant invasions in the kruger national park, south africa. pp. 385-403. in: child, l.e., j.h. brock, g. brundu, k. prach, p. pyšek, p.m. wade & m. williamson (eds.). plant invasions: ecological threats and management solutions. leiden: backhuys. gertenbach, w.p.d. 1985. alien plant section records. scientific services department, skukuza. unpublished records, south african national parks. henderson, l. 2001. alien weeds and invasive plants. pretoria: plant protection research institute. (handbook no. 12. agricultural research council). joubert, s.c.j. 1986. master plan for the management of the kruger national park. national parks board, skukuza, south africa. http://www.parks-sa.co.za lotter, w.d. 1997. management proposals for the alien aquatic biological invasions of the kruger national park. scientific report 10/97. national parks board, republic of south africa. lotter, w.d. & j.h. hoffmann. 1998. an integrated management plan for the control of opuntia stricta (cactaceae) in the kruger national park, south africa. koedoe 41(1): 63–68. macdonald, i.a.w. & w.p.d. gertenbach. 1988. a list of alien plants in the kruger national park. koedoe 31: 137-150. mcneely, j.a. 2000. global strategy for addressing the problem of invasive alien species. a result of the global invasive species programme (gisp). unpublished report. iucnthe world conservation union, switzerland. noss, r.f. 1990. indicators for monitoring biodiversity: a hierarchical approach. conservation biology 4: 355-364. obermeijer, a.a. 1937. a preliminary list of the plants found in the kruger national park. annals of the transvaal museum 17(4): 185-227. olckers, t & m.p. hill (eds). 1999. biological control of weeds in south africa (1990-1998). african entomology memoir no.1. richardson, d.m., p. pysek, m. rejmanek, m.g. barbour, f.d. panetta & c.j. west. 2000. naturalization and invasion of alien plants: concepts and definitions. diversity and distributions 6: 93-107. van der schijff, h.p. 1957. ekologiese studie van die flora van die nasionale krugerwildtuin. d.sc. thesis, potchefstroom university for christian higher education, potchefstroom. van der schijff, h.p. 1969. a checklist of the plants of the kruger national park. publikasies van die universiteit van pretoria, nuwe reeks 53: 1-100. foxcroft.qxd 2005/12/09 10:42 page 43 koedoe 46/2 (2003) 44 issn 0075-6458 appendix 1 list of plants not classed as aliens 1. centella asiatica (l.) urb. 2. trichodesma zeylanicum (burm.) r. br. 3. ocimum americanum l. var. americanum 4. hibiscus cannabinus l. var. cannabinus 5. sida alba l. 6. sida cordifolia l. 7. crotalaria sphaerocarpa perr. ex dc. subsp. sphaerocarpa 8. vigna oblongifolia a. rich. var. oblongifolia 9. dodonaea angustifolia l.f. 10. striga asiatica (l.) kuntze 11. withania somnifera (l.) dunal 12. triumfetta pentandra a. rich. var. pentandra 13. triumfetta rhomboidea jacq. var. rhomboidea foxcroft.qxd 2005/12/09 10:42 page 44 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <feff004e00e4006900640065006e002000610073006500740075007300740065006e0020006100760075006c006c006100200076006f0069006400610061006e0020006c0075006f006400610020005000440046002d0061007300690061006b00690072006a006f006a0061002c0020006a006f006900640065006e002000740075006c006f0073007400750073006c00610061007400750020006f006e0020006b006f0072006b006500610020006a00610020006b007500760061006e0020007400610072006b006b007500750073002000730075007500720069002e0020005000440046002d0061007300690061006b00690072006a0061007400200076006f0069006400610061006e0020006100760061007400610020004100630072006f006200610074002d0020006a0061002000520065006100640065007200200035002e00300020002d006f0068006a0065006c006d0061006c006c0061002000740061006900200075007500640065006d006d0061006c006c0061002000760065007200730069006f006c006c0061002e0020004e00e4006d00e4002000610073006500740075006b0073006500740020006500640065006c006c00790074007400e4007600e4007400200066006f006e0074007400690065006e002000750070006f00740075007300740061002e> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 abstract introduction methods results discussion acknowledgements references about the author(s) willem a. nieman department of conservation ecology and entomology, faculty of agrisciences, stellenbosch university, stellenbosch, south africa majete research programme, majete wildlife reserve, chikwawa, malawi brian w. van wilgen centre for invasion biology, department of botany and zoology, faculty of science, stellenbosch university, stellenbosch, south africa alison j. leslie department of conservation ecology and entomology, faculty of agrisciences, stellenbosch university, stellenbosch, south africa citation nieman, w.a., van wilgen, b.w. & leslie, a.j., 2021, ‘a review of fire management practices in african savanna-protected areas’, koedoe 63(1), a1655. https://doi.org/10.4102/koedoe.v63i1.1655 review article a review of fire management practices in african savanna-protected areas willem a. nieman, brian w. van wilgen, alison j. leslie received: 04 sept. 2020; accepted: 27 nov. 2020; published: 15 feb. 2021 copyright: © 2021. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract the level of understanding of the ecological effects of fires has improved over the past century, but comprehensive information on the practical application of fire remains restricted to a few well-studied areas, and management information is scattered. this article reviews the goals of fire management practices in african savanna-protected areas, and the approaches that have been adopted to achieve them. we identified 15 distinct fire management practices described in 107 papers from 19 african countries. fire management has evolved in response to changing ecological understanding, as well as the shifting goals of protected areas. currently, fire management practices can be divided into those that use fire to achieve specific ecological outcomes, those where fire is applied to promote diverse fire patterns across the landscape without necessarily having a specific ecological outcome in mind, and those that use fire to achieve specific, non-ecological or social goals. in larger, heterogeneous protected areas, fire management practices may vary at different sites in order to achieve a range of goals. we compared the effectiveness of each practice in terms of achieving 10 broad goals. these included ecological goals, for example, reversing woody and social goals (e.g. maintaining community relationships). conservation implications: fire management remains an important ecosystem process that can be manipulated to achieve particular goals in protected areas. the choice of a particular approach, or approaches, will depend on the circumstances pertaining to a particular protected area, and we provide examples of situations where each practice could be most appropriate. keywords: biodiversity; bush encroachment; fire regime; global change; grass fuel; co2-enrichment; environmental management; prescribed burning. introduction savannas are characterised by the co-dominance of woody plants and grasses, in which trees that are at least 2 metres (m) tall occur above a grassy layer between 0.5 m and 2 m (scholes & walker 1993). frequent natural fires have historically been reported in african savannas during the dry season, and have played an important role in shaping the composition and structure of these ecosystems (bond 2019). the importance of fire in maintaining vegetation in savanna is thus well recognised (archibald et al. 2017), especially in terms of ensuring the continued co-existence of woody and herbaceous plants (archibald et al. 2013). approaches to fire management have evolved over the past century, and some protected areas have undergone multiple changes in fire management over time (archibald et al. 2017; van wilgen et al. 2014). these changes came about in response to increases in understanding of the role and impacts of fire in natural ecosystems, as well as to changing societal needs and increasing challenges associated with global change, including climate change, increasing concentrations of atmospheric co2 and invasion by alien plants (bond & midgley 2012; sala et al. 2000). african savannas are dynamic ecosystems, and fire interacts with climatic cycles, varying levels of herbivory, and increasing pressures brought about by rising human populations (smit & archibald 2019). managers of african savanna-protected areas have to make decisions on the use of fire, including whether, where and how to apply fire, the conditions under which fire should be applied and the goals that such use is intended to achieve. whilst there is a growing literature on the ecological effects of fire, comprehensive information on the practical application of fire is restricted to a few well-studied areas. the remaining information remains scattered, and a review of the goals of fire management, and the means by which they can be achieved is, therefore, needed. in this research article, we report on a review of the literature relevant to the management of fire in savanna-protected areas in sub-saharan africa. the interest of this review is in conservation management, and the purpose of this study was to identify the various goals of fire management in protected areas and to review the practices that have been put forward to achieve them. the focus of this review was on studies that dealt with the practical application of fire in african savanna-protected areas, and the assessment of this study was based on these practical applications, and not on the many papers that dealt with the ecological effects of fire, or with the fire management of other african ecosystems, including forests, grasslands and shrublands, or agricultural land uses. methods a review of the relevant literature was initiated by examining the scopus and web of science databases in april 2020 to identify papers or book chapters published between 1960 and the present (2020) that were relevant to this study. we used the following selection terms in the title, abstract or keywords of the research article: ‘fire’ and ‘africa’ and ‘manag*’ or ‘prescribed’ or ‘policy’ or ‘plan’ or ‘control’ and ‘protected’ or ‘reserve’ or ‘park’ or ‘conserv*’. for all of the papers identified, we read the abstracts and excluded papers that were not relevant to the specific aims of this study (e.g. papers that were not from africa, papers that did not deal in sufficient detail with fire, papers that dealt with vegetation types other than savannas, papers that were not relevant to protected areas or their management or papers that addressed the ecological effects of fire, but not the management of fire). we then identified additional sources based on the personal knowledge and on references cited in papers identified in the search (these included studies or management plans not covered by scopus or web of science). for each of the papers that were retained or added, we noted the country where the study was conducted, the goals of fire management, the practices that were implemented or proposed to achieve the goals, the time periods in which the practices were implemented, and the principal findings or recommendations. we used this information to examine how and why management practices changed during the 20th century and beyond, and how different practices were implemented to achieve specific conservation goals. ethical consideration this article followed all ethical standards for research without direct contact with human or animal subjects. results literature identified the systematic search revealed 903 papers and book chapters, of which 848 were excluded for one or more of the reasons outlined above. we also identified 52 additional sources, and thus, had a sample of 107 papers based on the review. papers were sourced from 19 african countries, with over half (58) being from south africa (figure 1). several important early papers arose from a landmark conference dedicated to fire in africa, which was held at tallahassee, florida in 1971, with contributions from 20 authors covering seven african countries in southern, eastern and western africa (komarek 1972). following this, the rate of publication remained below one paper per year until the year 2000, after which there was a steady increase in published studies related to fire management in africa (figure 2). figure 1: origin of published sources on fire management in african savanna-protected areas. figure 2: the cumulative number of published studies related to fire management in african savanna-protected areas between 1922 and 2020. evolution of fire management fire management has evolved during the past century in response to changing ecological understanding, as well as to the shifting goals of protected areas (bond & archibald 2003; mentis & bailey 1990) (figure 3). in the early 20th century, colonial foresters often influenced the fire policy for large areas of woodland across africa. foresters believed that fire prevented vegetation from attaining its full potential (‘climax’) through succession after disturbance (as set out by clements 1916), and that it led to landscape degradation in wooded areas (aubréville 1947; chipp 1922; rains 1963; stebbing 1937). some examples of interpretations of fire effects in terms of clements’ succession model include regular burning kept savannas in a ‘non-climax’ (phillips 1930) or ‘sub-climax’ (roux 1969) condition, ‘climax’ savanna woodlands could persist in the absence of fire (austen 1972) and climax vegetation was a ‘theoretical ideal’ towards which savannas would develop in the absence of fire (west 1972). figure 3: examples of factors that influenced changes in fire management policies and management responses in selected african savanna-protected areas over the past 100 years. these interpretations led, in turn, either to attempts to exclude fire or to the promotion of early dry season fires of relatively low intensity that would minimise impacts on trees (geldenhuys 1977; laris & wardell 2006; trapnell 1959). examples include a total ban on fires in etosha national park in namibia (from its establishment in 1907 up to the 1980s) (stander, nott & mentis 1993), in the kruger national park in south africa (1948–1955) (joubert 2007), and protection from fire over large parts of hwange national park in zimbabwe in the 1960s and beyond (austen 1972). in west africa, fire was regarded as damaging in the early 20th century, and periodic attempts were made to ban fires. however, the inevitability of fire led to the acceptance of early dry season burning in the mid-1950s (wardell et al. 2004). the policy was underpinned by the findings of burning experiments, which established that early dry season burns would allow trees to regenerate and would also limit the more destructive effects of late dry season fires (charter & keay 1960; trapnell 1959). in contrast, the influential ecologist jfv phillips also noted in 1930 that ‘[c]ontrolled firing is a useful and oftentimes necessary agent in veld management’ (phillips 1930). mid-20th century pasture scientists followed this lead, noting that fires played an important role in the management of grassy rangelands (including savannas), serving to remove unpalatable material and rejuvenate the grass sward, as well as retarding encroachment by woody plants (scott 1955; west 1955). scott (1955) proposed that early wet season burns (applied in spring after the first rain) would be least harmful to the vegetation. managers of protected areas thus found themselves conflicted. for example, austen (1972) noted that ‘good forest policy (early burning) is not good wildlife policy’, as early burning was less detrimental to trees but encouraged bush encroachment. the recognition that fire was inevitable, and often necessary, led to the increasing adoption of fire as a management practice in many protected areas. in 1955, prescribed burning was initiated in the hluhluwe-imfolozi park, south africa, inter alia to counter bush increase and to improve visibility for tourists (vincent 1970). in the kruger national park, the application of early wet season fires on a 3-year cycle on fixed areas (‘blocks’) across the whole park was initiated in 1957 (brynard 1958). this policy change revealed that fires were both inevitable and ecologically important, and was to stay in place until ‘proven incorrect’. initially, fire management was aimed at maintaining protected areas in a desired (stable) state by applying fire at regular intervals, in line with ecological theories of equilibrium at the time (mentis & bailey 1990). these theories predicted that grass-tree coexistence was possible, for example, because of niche separation in the rooting depths of grasses and trees that allowed differential access to water in shallow and deeper layers of the soil (walter 1971), and that a range of stable states were possible (scholes & archer 1997). ecologists subsequently developed disequilibrium models, which suggested that stable states could not exist, and that wet and dry climatic cycles, fluctuating levels of grazing and browsing, and differences in fire frequency and intensity all combined to continually change the relative mix of grasses and trees (mentis & bailey 1990; scholes & archer 1997). this led to the encouragement of more variability in the application of fire. a growing recognition of the importance of biodiversity also influenced fire management in the later 20th century. the convention on biological diversity entered into force on 29 december 1993 and was initially signed by 168 countries. member countries undertook, as per article 8, to establish and manage protected areas that could maintain viable populations of all species. managers of african savanna-protected areas began to shift their focus from creating forage for large mammals or protecting trees to catering for the needs of all species. given the lack of scientific evidence to underpin the formulation of appropriate fire regimes, managers opted for variable fire regimes that would presumably mimic natural processes and provide a wide range of conditions to support all facets of biodiversity. one such approach was patch mosaic burning, which was initially developed in australia (saxon 1984) and which aims to create diversity in all elements of the fire regime (brockett, biggs & van wilgen 2001; parr & brockett 1999). the approach assumes that spatial and temporal heterogeneity in fire patterns will promote biodiversity conservation. a policy of patch-mosaic burning was implemented in mkhuze game reserve in 1985 (mulqueeny, goodman & o’connor 2010), in pilanesberg national park in 1989 (brockett et al. 2001) and in the kruger national park in 2005 (van wilgen, govender & macfadyen 2008). siegfried (1981) also noted that ‘a primary objective of a fire-management strategy for a nature reserve should be an imitation of the local natural fire regime’. this reasoning led to the adoption of policies of deliberately burning to simulate lightning-ignited fire patterns in etosha national park in 1985, du plessis (1997), or sought to restrict all fires to those ignited by lightning (in the kruger national park in 1999, biggs & potgieter 1999). in the early 1990s, the kruger national park also adopted adaptive management as an overarching framework to guide all of its activities, including fire management (van wilgen & biggs 2011). van wilgen, biggs and potgieter (1998) proposed a set of thresholds of potential concern (tpcs) in which upper and lower limits were defined for the area burnt, and the distribution of fire seasonality, size and intensity. variability in fire patterns was thus tolerated within limits, and if any tpc was exceeded, then either the management approach could be altered or the threshold could be recalibrated (biggs & rogers 2003). there was little in the way of scientific evidence to guide the degree of variability in fire regimes required to conserve biodiversity in any of these policies (parr & andersen 2006), and the approaches were essentially an insurance policy designed to capture enough variability in fire regimes as might be necessary to cater for all species. in the 21st century, technological approaches such as remote sensing and fire behaviour prediction models have been increasingly proposed to support the implementation of prescribed burns. for example, it was suggested that remotely sensed estimates of herbaceous biomass (verbesselt et al. 2006) or fuel moisture content (verbesselt et al. 2007) could be used to evaluate fire danger, and subsequently assist managers in the identification of areas and timings for prescribed burning. similarly, real-time monitoring of vegetation biomass with national oceanic and atmospheric administration advanced very high resolution radiometer (noaa-avhrr) has been proposed for use in arid ecosystems, such as the etosha national park (sannier, taylor & du plessis 2002) and the tswalu kalahari reserve, south africa (tokura et al. 2018), to assist prescribed burning as a form of adaptive fire management. fire management has historically been adjusted in individual protected areas several times as understanding developed, but there are few well-documented accounts of these multiple changes outside of south africa. changing fire policies are well recorded for the kruger national park (biggs & potgieter 1999; joubert 2007; van wilgen et al. 2014) and the hluhluwe-imfolozi park (archibald et al. 2017; vincent 1970). in the serengeti national park in tanzania, fires were essentially tolerated for decades before a formal fire management plan was adopted. in the serengeti, norton-griffiths (1979) noted that sources of ignition included lightning, fires ignited by pastoralists, honey-hunters and poachers, and that park wardens used fires to make it easier to apprehend poachers and honey-hunters. in 1988, a fire management plan was adopted for the serengeti, with the aim of reducing the impact of fire on regenerating woodlands, preventing large fires in the late dry season and improving visibility in tourist areas (stronach 1988). in the following sections, we discuss the approaches that have been adopted to achieve broad management goals. broad categories of fire management fire can be manipulated to achieve a wide range of goals in protected areas, and these goals can be divided into three broad categories: firstly, fire is used to achieve specific ecological outcomes, where fire is applied at a particular frequency, season or intensity, or where fire is intentionally avoided or supressed to directly influence clearly identified aspects of vegetation structure and composition. secondly, fire is applied to promote a diversity of fire patterns across the landscape (e.g. a mosaic of fire return periods, seasons, intensities and sizes), without necessarily having a specific ecological outcome in mind. the rationale for such approaches has been either that diversity would promote the continued co-existence of co-occurring species that differed in their response or tolerance to elements of the fire regime, or, alternatively, that diversity would mimic natural variations that would have occurred in the absence of human interference. thirdly, fire is used to achieve specific goals that are non-ecological or social in nature, and these include issues relating to safety or other human benefits (table 1). each of these broad categories is discussed below. table 1: fire management goals and associated practices that have been proposed, or are used, for the management of fire in african savanna-protected areas. fire management to achieve specific ecological outcomes reducing tree mortality up to the mid-20th century, the practice of not using fire, or actively suppressing any fires that may occur, was widely promoted to reduce the putative negative effects of fire, especially on large trees, or to allow for the development of climax vegetation (west 1972). these fire protection policies were common across african savanna areas (bond & archibald 2003; du plessis 1997; egunjobi 2018; laris & wardell 2006; pricope & binford 2012), but these largely failed to achieve their intended purpose, as large fires persisted despite fire suppression (biggs & potgieter 1999; pricope & binford 2012). additionally, fire suppression may lead to an increased bush encroachment, particularly when coupled with overgrazing, as shown in madikwe game reserve, south africa (hudak et al. 1998, 2004). fire suppression policies have, therefore, become largely outdated, but some countries, particularly in west africa (eriksen 2007; goldammer & de ronde 2004), remain opposed to fire. in more recent years, managers wishing to reduce tree mortality rates or promote tree recovery (especially in miombo woodlands, ryan & williams 2011) opted for low-intensity, early dry season fires under mild weather conditions (govender et al. 2006; ribeiro et al. 2017; smit et al. 2016), or for reduced fire frequencies (especially in areas with high elephant densities; eckhardt et al. 2000). improve forage quality and remove moribund or unpalatable grass material burning to improve forage quality (i.e. range condition), or to remove moribund or unpalatable grass for the benefit of grazers, usually requires managers to base their decision of whether to burn on the species composition and biomass of the grass sward (bond & archibald 2003; van wilgen et al. 2003). the main goal is often not necessarily to improve range condition but to encourage uniform grazing, and to decrease unpalatable grasses and bare ground patches. as a result, the forage quality for wildlife is improved, thus enabling larger numbers of grazing animals to be supported. a balanced woody to grass ratio is also established, and wildlife is encouraged to move to less preferred areas in order to prevent overutilisation (edwards 1984; trollope 2007). in order to achieve these goals, burning can take place either (1) in the early dry season to produce a flush of green grass regrowth or (2) in the early wet season to remove moribund grass that remains after the dry season with minimal damage to the grass sward (scott 1955; trollope 2007). burning in the early dry season would force grasses to regrow through the dry season without the benefit of moisture, thus removing forage that could sustain grazing mammals through the dry season (archibald et al. 2017; scholes & walker 1993). fires in either the early dry or early wet season need to be of sufficient intensity to avoid bush encroachment (o’connor, puttick & hoffman 2014). areas of relatively high rainfall (> 650 millimetres [mm] map) can usually be burnt more regularly, but in more arid sites fire should only be applied in years when the grass biomass is unusually high, normally following a period of above-average rainfall, and when the grass species composition warrants the use of fire to remove unpalatable species (trollope et al. 2014). reverse woody encroachment in instances where the density of woody plants has increased to undesirable levels, relatively high-intensity fires can be used to reverse woody encroachment and limit tree recruitment (ryan & williams 2011; smit et al. 2016; van wilgen et al. 2014). high-intensity fires can also have unintended consequences, such as increased mortality rates of very large trees (govender et al. 2006; smit et al. 2016), compromising the safety of burning crews (kamminga 2001), and increasing greenhouse gas emissions (van der werf et al. 2010). the effect of high-intensity fires on tree mortality is also more pronounced in drier ecosystems (holdo 2005), in years when grass fuels are high (govender et al. 2006), and in miombo woodlands, where they inhibit the natural regeneration of miombo trees (ribeiro et al. 2017). burning in the late dry season is, therefore, not widely used, although it has been shown to be effective in reducing the cover of shrubs on an experimental basis in the kruger national park (smit et al. 2016) and has been included in the fire management plan of the okavango delta ramsar site (trollope et al. 2006). fires in the late dry season nonetheless remain illegal or are actively discouraged in many african countries, particularly in west africa (laris & wardell 2006). early wet season fires can also reverse woody encroachment, provided that grass fuel loads are high enough to support a relatively high-intensity fire (see, e.g., crowley, garnett & shephard 2009). control invasive alien plants invasive alien plants are a substantially growing problem in protected areas (foxcroft et al. 2013), including african savannas (te beest et al. 2012; van wilgen et al. 2017; witt et al. 2017), and fire is often involved in both their spread and their management (dew et al. 2017). fires of high intensity can be used to clear invasive alien plants and reduce their seedling densities (te beest et al. 2012). te beest et al. (2017) suggested that frequent fires of high intensity would be necessary to control the invasive alien shrub chromolaena odorata. similarly, bukombe et al. (2018) found that alien species were more abundant in areas with infrequent or no fires. fires, however, also open up the landscape, providing opportunities for alien species to establish (te beest et al. 2015), and a deliberate reduction in fire frequency has, therefore, been suggested for some protected areas to limit the colonisation potential of alien invasive species. this applies especially to mesic savannas, where alien forbs readily establish on disturbed areas (masocha et al. 2011). because responses to fire are species-specific, a species-specific approach to managing alien invasions would be required. protect rare or fire-sensitive species or habitats within protected areas, fires may be excluded from the habitats of rare or fire-sensitive animal species, such as reptiles, rhinoceros and nesting birds (eby et al. 2015), or vegetative communities to maintain their presence in the landscape, such as regenerating woodlands, tree copses, rocky hill vegetation, forests, and thickets (eby et al. 2015; nangendo et al. 2005; pfab & witkowski 2000; stronach & mcnaughton 1989). furthermore, in many african-protected areas, forest, woodland and savanna vegetation types coexist, and fire management should accommodate this variation. for example, nangendo et al. (2006) proposed that forest landscapes in the budongo forest reserve, uganda, should not be burned, whilst the adjacent wooded grassland should be burnt regularly. similarly, a variety of approaches were recommended for murchison falls national park, uganda, because of the large variety of vegetation types (nangendo et al. 2005). te beest et al. (2012) recommended selective fire suppression in riverine vegetation and margins of scarp forests to protect fire-sensitive endemic species. excluding fires from selected areas is, however, difficult in protected areas that are prone to unplanned fires, or that have limited resources to manage fires (nangendo et al. 2005). control disease vectors fire can be a useful tool for controlling an array of disease vectors. it is most commonly used for controlling ticks (acari: ixodidae), which may cause wildlife mortalities when excessively abundant (trollope et al. 2003). numerous studies have validated this approach by demonstrating a decline in tick abundance for up to 3 years after a fire (fyumagwa et al. 2007; goodenough et al. 2017). tick numbers not only are reduced by the actual fire but also remain low long after the fire because of less abundant grass (trollope et al. 2003). similarly, managers of protected areas have used fire to reduce the risk of exposure to african trypanosomiasis (sleeping sickness) by reducing the incidence of tsetse flies (glossina spp.) (eby et al. 2015). fire management to promote diverse fire patterns in the past three decades, the importance of heterogeneous fire patterns (i.e. pyrodiversity) to accommodate all facets of biodiversity has increasingly been highlighted as an important management consideration (parr & andersen 2006), resulting in attempts to promote diverse fire regimes in many savanna-protected areas. the promotion of pyrodiversity is believed to provide an array of habitats through space and time, which should allow biota with different life histories to coexist at the landscape scale (beale et al. 2018). these policies, known as patch mosaic burning, aim to create heterogeneity in fire distributions, size classes, return intervals, seasons and intensity (brockett et al. 2001; mentis & bailey 1990). the reported outcomes of patch-mosaic burning policies implemented in south african-protected areas (archibald et al. 2017; balfour & howison 2001; brockett et al. 2001; mulqueeny et al. 2010) most often include a shift in a more even distribution of fires over the dry season, an increase in the number of fires and a reduction in mean fire sizes because of the breaking up of continuous grass fuels in relatively small early dry season burns. both increased fire frequencies (mulqueeny et al. 2010) and spatial heterogeneity (brockett et al. 2001) have also resulted from these approaches in smaller (< 100 000 hectares [ha]) protected areas. in larger protected areas, managers have been able to manipulate the spatial heterogeneity of fires, but fire return periods were correlated with annual rainfall rather than by management interventions (van wilgen et al. 2004). the majority of fires reported in african savanna-protected areas often occur during the late dry season as unplanned wildfires (lipsett-moore et al. 2018) because fires are ignited outside of protected areas (eriksen 2007). fires in the early dry season are thus widely considered to be the only practical means to prevent wildfires in the late dry season by breaking up of continuous fuel beds (owen 1972), and patch mosaic burning could achieve this goal by igniting fires in the early dry season. patch burning is sometimes closely associated with natural fire policies and has been implemented as a hybrid system in the kruger national park (van wilgen et al. 2014). the promotion of a natural fire regime is based on the belief that ‘nature knows best’ (siegfried 1981; van wilgen et al. 1998), and human interference should thus be kept to a minimum. implementing a natural fire regime thus entails allowing all fires ignited by lightning to spread unhindered, whilst suppressing any fires ignited by human beings. this policy was, however, abandoned in the kruger national park after 10 years of implementation, when it became apparent that it was unworkable, because the vast majority of the area continued to be burnt by fires ignited by people (van wilgen & biggs 2011). the etosha national park in namibia sought to replicate lightning fire patterns by applying prescribed burns to selected areas that had received above-average rainfall in the preceding year and by limiting burning to < 12% of the protected area (du plessis 1997; stander et al. 1993). fires should be applied in the early wet season, as this is thought to mimic a fire regime that would have been driven by lightning storms (goldammer & de ronde 2004). fire management to promote non-ecological or social goals greenhouse gas emissions and carbon stocks global emissions from wildfires (mainly methane [ch4] and nitrous oxide [n2o]) (landry & matthews 2016) have become a cause of concern in recent years (van der werf et al. 2017). it has thus been proposed that burning in the early dry season in africa’s protected areas could offer an opportunity to reduce these gas emissions. a switch from late to early dry season fires can potentially reduce gas emissions by as much as 37% (64.2 mtco2-e yr−1) (lipsett-moore et al. 2018; russell-smith et al. 2013). simultaneously, this could provide stakeholders in protected areas with substantial revenue from the green climate fund (green climate fund [gcf] pilot programme 2017). for this effect to become apparent, however, grass fuels will have to be fully cured before burning occurs, as moist grass would produce smoldering fires with enhanced ch4 and carbon monoxide (co) emissions (meyer et al. 2012). avoiding conflict with local communities most protected areas in african savannas exist in close proximity to indigenous communities, and often, the responsibility for land management is shared by conservation and local authorities. conflicts may arise between the two if an appropriate approach to fire management is not agreed upon (laris & wardell 2006; moura et al. 2019). local communities may consider regular burning in the early dry season to be necessary for, inter alia, improving both pasture and hunting landscapes, clearing fields for crops, or preventing destructive fires later in the dry season (eriksen 2007; laris & wardell 2006), whilst the conservation goals of protected areas may not be aligned with this (eriksen 2007; laris 2002). in instances where fire policies were implemented without due consideration of the resource needs and traditional beliefs of local communities, large-scale illegal burning has occurred in retaliation (hough 1993; kull 2002). for example, kamau and medley (2014) listed ‘protest against the kenya wildlife service’ as a reason given by communities for burning, whilst laris and wardell (2006) noted that west african communities were ‘setting fires covertly and as a form of protest’ against punishment for practising traditional burning. it is thus often necessary for managers of protected areas and representatives of local communities to work together to design appropriate fire regimes that incorporate modern conservation goals and traditional practices (kamau & medley 2014; moura et al. 2019; shaffer et al. 2010) in order to avoid alienating indigenous communities (eriksen 2007). this will likely result in increases in fires in the early dry season, which may placate local communities, as well as reduce greenhouse gas emissions (moura et al. 2019). however, in instances where local communities are permitted to collect resources (most often thatch grass, but also bamboo, reeds and medicinal plants) from protected areas, fires may have to be postponed until the late dry season to provide access to thatch grass and other subsistence goods that need to be collected before burning (hough 1993; nieman 2020), which may prevent managers from planning fire regimes to achieve desired goals (archibald et al. 2017). substantial progress has been made in connecting local communities and protected area managers through the development of community-based fire management and integrated fire management concepts during the late 1990s (goldammer & de ronde 2004; pricope & binford 2012). additional non-ecological goals low-intensity fires in the early dry season are more easily controlled than high-intensity fires in the late dry season, and managers, therefore, often prefer applying prescribed fires in the early dry season to promote the safety of people, wildlife and infrastructure (kamminga 2001). fires may also be entirely excluded from areas with high human population or infrastructure density by establishing firebreaks before the onset of the fire season (nieman 2020), and in the hluhluwe-imfolozi park, active fire suppression was practised to prevent fires from destroying harris traps that were used to reduce populations of tsetse flies (archibald et al. 2017). in addition, in order to enhance tourist experiences, managers may use fires to improve visibility for game viewing or to minimise large, unattractive burnt areas by applying low-intensity burns in the early dry season (eby et al. 2015; hudak 1999; nieman 2020). likewise, managers may use fires to create conditions for effective policing, such as burning areas with high poaching incidence, or where there are security concerns, to improve visibility and accessibility (eby et al. 2015). finally, in order to avoid wasting scarce resources, managers may simply allow fires to burn unhindered (laissez faire) in cases where there is no evidence to suggest detrimental effects on ecosystems, or where there is insufficient capacity to implement alternative approaches. ribeiro et al. (2017) recommended this approach in areas reported with high frequency of fire to allow for healthy persistence of the vegetation (3–4 years). riparian zones in the kruger national park are also simply allowed to burn in infrequent fires (van wilgen et al. 2014). discussion options for fire management this review has confirmed the widespread use of fire in the management of savannah-protected areas in africa, reflecting on a growing body of research to evaluate past fire management approaches and to make proposals for novel ways to manage fire in the face of global change. although a number of fire management approaches have become largely outdated (e.g. fire suppression policies), and novel methods of monitoring fires (e.g. remote sensing) have improved the ability to monitor and understand fire, there is still little consensus on a broad fire management approach that would adequately cater for different ecological or societal circumstances. this can leave managers without clear guidelines for implementing appropriate fire regimes, especially in the face of unprecedented global change. the seven distinct fire management approaches identified here have been used to achieve 10 ecological and socio-economic goals that were sometimes confined to specific time periods and locations (table 2). application of fires in the early dry and wet season fires is arguably the most practical approach for balancing both ecological (e.g. maintaining good range condition and controlling disease vectors) and societal (e.g. maintaining good relationships with communities and promoting safe burning conditions) goals, and is thus appropriate for an array of savanna-protected areas, particularly those that are heavily influenced by local communities. conversely, whilst application of fires in the late dry season may provide the same ecological and social benefits as early dry or early wet season regimes, they can compromise the safety of people, wildlife and infrastructure (kamminga 2001), and are, therefore, normally only considered for use in protected areas with exceedingly high levels of woody encroachment. fires in the late dry season may also be useful for controlling invasive alien plants (te beest et al. 2012), but specific guidelines need to be developed for the alien species concerned. fire exclusion would protect tall trees (eckhardt et al. 2000), but would be costly and impractical for most fire-prone african savanna-protected areas (berry & macdonald 1979). fire exclusion may nonetheless be suited for the protection of discrete fire-sensitive habitats within protected areas (eby et al. 2015). natural, lightning-driven regimes are similarly difficult to maintain because of the influence of human ignitions, but can be simulated in large protected areas. patch-mosaic burning regimes, in contrast, are more ideally suited for smaller (< 100 000 ha) protected areas; however, brockett et al. (2001) noted that this policy is not appropriate for areas smaller than 20 000 ha, but many gaps remain in our understanding of this approach. for example, although pyrodiversity is argued to beget biodiversity (martin & sapsis 1992), it remains unclear exactly what degree of pyrodiversity would be needed to conserve biodiversity (parr & andersen 2006). finally, hands-off (laissez faire) approaches to fire management are not ideal, but may be adopted if there is no evidence that the prevailing fire regime has detrimental consequences. a laissez faire approach may also be necessitated in protected areas that lack fire management resources, or in instances where fire management funds have been redirected to address other issues of conservation policy. for example, the imperative to curb rampant poaching of rhinoceros in south africa’s protected areas has placed undue pressure on managers (ferreira, pfab & knight 2014), resulting in funds being redirected to anti-rhinoceros poaching activities and reducing the capacity for inter alia fire management. similarly, fire management has become much less of a priority in hwange national park since the onset of the ongoing economic recession in zimbabwe (pricope & binford 2012). table 2: goals of fire management in african savanna-protected areas, with an assessment of the suitability of different practices for achieving the goal concerned. fire management trade-offs and conflicts managers of protected areas normally use fire to influence a conservation-orientated outcome, and in many instances, a particular type of fire may be used to achieve the desired goal, but it could at the same time also precipitate unintended additional outcomes. furthermore, a single protected area may have multiple desired goals for fire management that could contradict each other. managers may, therefore, have to consciously consider making trade-offs when selecting a particular approach for fire management. for example, high-intensity fires are normally used to reduce the cover of woody plants, which is expected to accelerate because of increasing co2 levels in the 21st century (bond & midgley 2012). however, the effectiveness of high-intensity fires may have to be reduced to accommodate safety needs. in addition, smit et al. (2016) noted that the use of repeated high-intensity fires simultaneously causes both a positive (reduction in cover of short shrubs) and a negative (loss of tall trees) outcome, and that managers would need to consider trade-offs. case and staver (2017) also noted that grazing mammals and fire are competitors for grass fuel, and because high-intensity fires require high fuel loads, managers would face a stark choice between the maintenance of short-term grazer populations (which would reduce fuel loads) and prevention of woody encroachment (which requires a build-up of high fuel loads). another example includes the trade-offs necessitated by modern threats to ecosystem health. many approaches to fire management were designed to promote conservation goals in pristine ecosystems unaffected by the need to reduce greenhouse gas emissions, to sequester carbon (which would require the encouragement of woody encroachment, lipsett-moore et al. 2018), to combat invasive alien species (sala et al. 2000) or to accommodate traditional burning practices, which may be at odds with conservation objectives (ryan & williams 2011; shaffer et al. 2010). these pressures are increasingly brought to bear on fire managers, and fire management policies will, therefore, have to be developed taking potential conflicting goals into account, and trade-offs may be needed to accommodate an optimal range of outcomes. imposing desired fire regimes once a particular approach to fire management is chosen to promote desired outcomes, managers should attempt to maintain elements of the fire regime within certain limits by manipulating the season, frequency, intensity and size of fires. however, implementing and maintaining any chosen fire regime in the long term remain notoriously challenging, particularly in large conservation areas (smit et al. 2016). managers often have to contend with unplanned fires, which alter predefined burn-area targets or seasonal distributions (eriksen 2007; van wilgen et al. 2004). for example, a substantial number of fires were reported in the hluhluwe-imfolozi park during the time when fires were actively suppressed (1930–1940) (berry & macdonald 1979), and most areas in the kruger national park that had been burnt during the first 9 years of the lightning fire policy were ignited by sources other than lightning (van wilgen et al. 2007). furthermore, a study in the 2 million hectare kruger national park (van wilgen et al. 2004) found that four major changes to the fire management policy over six decades had little effect on the total area burnt, which was strongly related to rainfall (and, therefore, grass fuel loads) in the preceding 2 years. management did, however, alter the spatial heterogeneity of fires and their seasonal distribution over this period. similarly, in the smaller (20 000 ha) pilanesberg national park, brockett et al. (2001) observed that the spatial heterogeneity of fire patterns increased a few years after the introduction of the patch-mosaic burning policy. it appears, therefore, that managers should be able to influence the fire regime in terms of season and size distributions (and possibly in intensity, smit et al. 2016), but not in terms of fire frequency. despite the difficulty of maintaining desired fire return periods, it remains an important consideration for managers, irrespective of the chosen fire regime, as both too frequent and too infrequent fires can result in undesired consequences (mccleery et al., 2018). typically, areas of higher rainfall can be burnt more frequently, whilst areas of lower rainfall (< 650 mm map) should only be burnt in years of above-average rainfall and when the botanical composition suggests that a fire would be beneficial (trollope et al. 2014). acknowledgements the authors would like to thank marie theron and nicola van wilgen for assistance in sourcing the publications cited in this review, and jeremy russel-smith for comments on an earlier draft. w.a.n. acknowledges logistical support received from african parks ltd. competing interests the authors have declared that no competing interests exist. authors’ contributions all authors conceptualised the study. w.a.n. and b.w.v.w. jointly sourced and interpreted the relevant literature and co-wrote the research article. all authors approved the final text. funding information this study was funded by the earthwatch institute (grant to a.j.l.) under the majete wildlife research programme, malawi, the dsi-nrf centre for invasion biology, stellenbosch university, and the national research foundation of south africa (grant 109467 to b.w.v.w.). none of the funding agencies played any role in the design of the study, collection, analysis, interpretation of data or in writing the article. data availability statement data sharing is not applicable to this research article as no new data were created or analysed in this study. disclaimer the views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors. references archibald, s., beckett, h., bond, w.j., coetsee, c., druce, d.j. & staver, c.a., 2017, ‘interactions between fire and ecosystem processes’, in conserving africa’s mega-diversity in the anthropocene: the hluhluwe-imfolozi park story, pp. 233–262, cambridge university press, cambridge. archibald, s., lehmann, c.e.r., gómez-dans, j.l. & bradstock, r.a., 2013, ‘defining pyromes and global syndromes of fire regimes’, proceedings of the national academy of sciences of the united states of america 110(16), 6442–6447. https://doi.org/10.1073/pnas.1211466110 aubréville, a.m.a., 1947, ‘the disappearance of the tropical forests of africa’, unasylva 1(1), 5–11. reprinted in fire ecology 9(2), 3–13, (2013). https://doi.org/10.1007/bf03400624 austen, b., 1972, ‘the history of veld burning in the wankie national park, rhodesia’, in proceedings of the annual tall timbers fire ecology conference, 11, tall timbers research station, june 8–9, 1972, tallahassee, fl, pp. 277–296. balfour, d.a. & howison, o.e., 2001, ‘spatial and temporal variation in a mesic savanna fire regime: responses to variation in annual rainfall’, african journal of range & forage science 19(1), 45–53. https://doi.org/10.2989/10220110209485773 beale, c.m., courtney mustaphi, c.j., morrison, t.a., archibald, s., anderson, t.m., dobson, a.p. et al., 2018, ‘pyrodiversity interacts with rainfall to increase bird and mammal richness in african savannas’, ecology letters 21(4), 557–567. https://doi.org/10.1111/ele.12921 berry, a. & macdonald, i.a.w., 1979, fire regime characteristics in the hluhluwe-corridor-umfolozi game reserve complex in zululand. area description and an analysis of causal factors and seasonal incidence of fire in the central complex with particular reference to the period 1955 to 1978, unpublished report, natal parks board, pietermaritzburg. biggs, h.c. & potgieter, a.l.f., 1999, ‘overview of the fire management policy of the kruger national park’, koedoe 42(1), 101–110. https://doi.org/10.4102/koedoe.v42i1.227 biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59–80, island press, washington, dc. bond, w.j., 2019, open ecosystems: ecology and evolution beyond the forest edge, oxford university press, oxford. https://doi.org/10.1093/oso/9780198812456.001.0001 bond, w.j. & archibald, s., 2003, ‘confronting complexity: fire policy choices in south african savanna parks’, international journal of wildland fire 12(4), 381–389. https://doi.org/10.1071/wf03024 bond, w.j. & midgley, g.f., 2012, ‘carbon dioxide and the uneasy interactions of trees and savannah grasses’, philosophical transactions of the royal society of london b: biological sciences 367(1588), 601–612. https://doi.org/10.1098/rstb.2011.0182 brockett, b.h., biggs, h.c. & van wilgen, b.w., 2001, ‘a patch mosaic burning system for conservation areas in southern african savannas’, international journal of wildland fire 10(2), 169–183. https://doi.org/10.1071/wf01024 browne, c. & bond, w., 2011, ‘firestorms in savanna and forest ecosytems: curse or cure?’, veld & flora 97(2), 62–63. brynard, a.m., 1958, memorandum in verband met die beleid van driejaarlikse rotasiebrand in die nationale krugerwildtuin, addendum k 1, board of curators of national parks, pretoria. brynard, a.m., 1972, ‘controlled burning in the kruger national park-history and development of a veld burning policy’, tall timbers fire ecology conference proceedings 838, viewed 22 april 2020, from http://agris.fao.org/agris-search/search.do?recordid=us201302337593. bukombe, j., smith, s.w., kija, h., loishooki, a., sumay, g., mwita, m. et al., 2018, ‘fire regulates the abundance of alien plant species around roads and settlements in the serengeti national park’, management of biological invasions 9(3), 357–367. https://doi.org/10.3391/mbi.2018.9.3.17 case, m.f. & staver, a.c., 2017, ‘fire prevents woody encroachment only at higher-than-historical frequencies in a south african savanna’, journal of applied ecology 64(3), 955–962. https://doi.org/10.1111/1365-2664.12805 charter, j.r. & keay, r.w., 1960, ‘assessment of the olokemeji fire-control experiment (investigation 254) 28 years after institution’, nigerian forestry information bulletin 3, 1–32. chipp, t.f., 1922, forest officers’ handbook of the gold coast, ashanti and the northern territories, gold coast colony government printer, accra. clements, f.e., 1916, plant succession: an analysis of the development of vegetation, carnegie institution of washington, washington, d.c. crowley, g., garnett, s. & shephard, s., 2009, ‘impact of storm-burning on melaleuca viridiflora invasion of grasslands and grassy woodlands on cape york peninsula, australia’, austral ecology 34(2), 196–209. https://doi.org/10.1111/j.1442-9993.2008.01921.x dew, l.a., rozen-rechels, d., le roux, e., cromsigt, j.p.g.m. & te beest, m., 2017, ‘evaluating the efficacy of invasive plant control in response to ecological factors’, south african journal of botany 109, 203–213. https://doi.org/10.1016/j.sajb.2016.12.007 docherty, t.d.s., hethcoat, m.g., mactavish, l.m., mactavish, d., dell, s., stephens, p.a. et al., 2020, ‘burning savanna for avian species richness and functional diversity’, ecological applications 30(4), e02091. https://doi.org/10.1002/eap.2091 du plessis, w.p., 1997, ‘refinements to the burning strategy in the etosha national park, namibia’, koedoe 40(1), 63–76. https://doi.org/10.4102/koedoe.v40i1.264 eby, s.l., dempewolf, j., holdo, r.m. & metzger, k.l., 2015, ‘fire in the serengeti ecosystem: history, drivers, and consequences’, in a.r.e. sinclair, k.l. metzger, s.a.r. mduma & j.m. fryxell (eds.), serengeti iv: sustaining biodiversity in a coupled human-natural system, pp. 73–104, the university of chicago press, chicago, u.s. eckhardt, h.c., van wilgen, b.w. & biggs, h.c., 2000, ‘trends in woody vegetation cover in the kruger national park, south africa, between 1940 and 1998’, african journal of ecology 38(2), 108–115. https://doi.org/10.1046/j.1365-2028.2000.00217.x edwards, d., 1984, ‘fire regimes in the biomes of south africa’, in p. de v. booysen & n.m. tainton (eds.), ecological effects of fire in south african ecosystems, pp. 19–37, springer, heidelberg, berlin. egunjobi, j.k., 2018, ‘an appraisel of the savanna burning problem’, in s.s. ajayi & l.b. halstead (eds.), wildlife management in savannah woodland, pp. 161–164, routledge revivals, taylor and francis ltd., london. eriksen, c., 2007, ‘why do they burn the “bush”? fire, rural livelihoods, and conservation in zambia’, geographical journal 173(3), 242–256. https://doi.org/10.1111/j.1475-4959.2007.00239.x ferreira, s.m., pfab, m. & knight, m.h., 2014, ‘management strategies to curb rhino poaching: an exploration of alternative options’, south african journal of science 110(5), 6. https://doi.org/10.1590/sajs.2014/20120055 foxcroft, l.c., pyšek, p., richardson, d.m. & genovesi, p., 2013, plant invasions in protected areas: patterns, problems and challenges, vol. 7, springer science & business media, berlin. https://doi.org/10.1007/978-94-007-7750-7 fyumagwa, r.d., runyoro, v., horak, i.g. & hoare, r., 2007, ‘ecology and control of ticks as disease vectors in wildlife of the ngorongoro crater, tanzania’, african journal of wildlife research 37(1), 79–90. https://doi.org/10.3957/0379-4369-37.1.79 geldenhuys, c.j., 1977, ‘the effect of different regimes of annual burning on two woodland communities in kavango’, south african forestry journal 103(1), 32–42. https://doi.org/10.1080/00382167.1977.9629467 goldammer, j.g. & de ronde, c. (eds.), 2004, wildland fire management handbook for sub-sahara africa, african minds, global fire monitoring center (gfmc), freiburg. goodenough, a.e., harrell, a.n., keating, r.l., rolfe, r.n., stubbs, h., mactavish, l. et al., 2017, ‘managing grassland for wildlife: the effects of rotational burning on tick presence and abundance in african savannah habitat’, wildlife biology 2017(4), 1–9. https://doi.org/10.2981/wlb.00318 govender, n., mutanga, o. & ntsala, d., 2012, ‘veld fire reporting and mapping techniques in the kruger national park, south africa, from 1941 to 2011’, african journal of range and forage science 29(2), 63–73. https://doi.org/10.2989/10220119.2012.697918 govender, n., trollope, w.s.w. & van wilgen, b.w., 2006, ‘the effect of fire season, fire frequency, rainfall and management on fire intensity in savanna vegetation in south africa’, journal of applied ecology 43(4), 748–758. https://doi.org/10.1111/j.1365-2664.2006.01184.x green climate fund (gcf), 2017, terms of reference for the pilot programme for redd+ results-based payments, green climate fund incheon. gureja, n. & owen-smith, n., 2002, ‘comparative use of burnt grassland by rare antelope species in a lowveld game ranch, south africa’, south african journal of wildlife research 32(1), 31–38. holdo, r.m., 2005, ‘stem mortality following fire in kalahari sand vegetation: effects of frost, prior damage, and tree neighbourhoods’, plant ecology 180(1), 77–86. https://doi.org/10.1007/s11258-005-2796-4 hough, j.l., 1993, ‘why burn the bush? social approaches to bush-fire management in west african national parks’, biological conservation 65(1), 23–28. https://doi.org/10.1016/0006-3207(93)90192-4 hudak, a.t., 1999, ‘rangeland mismanagement in south africa: failure to apply ecological knowledge’, human ecology 27, 55–78. https://doi.org/10.1023/a:1018705300730 hudak, a.t., brockett, b.h. & wessman, c.a., 1998, ‘fire scar mapping in a southern african savanna’, international geoscience and remote sensing symposium (igarss) 3, 1608–1610. https://doi.org/10.1109/igarss.1998.691638 hudak, a.t., fairbanks, d.h.k. & brockett, b.h., 2004, ‘trends in fire patterns in a southern african savanna under alternative land use practices’, agriculture, ecosystems and environment 101(2–3), 307–325. https://doi.org/10.1016/j.agee.2003.09.010 joubert, s., 2007, the kruger national park: a history, vol. 2, high branching, johannesburg. kamau, p.n. & medley, k.e., 2014, ‘anthropogenic fires and local livelihoods at chyulu hills, kenya’, landscape and urban planning 124, 76–84. https://doi.org/10.1016/j.landurbplan.2014.01.010 kamminga, e., 2001, impact of the integrated forest fire management program on rural livelihoods in east caprivi region, namibia, pp. 1–43 namibia finland forestry program, directorate of forestry, ministry of environment and tourism, windhoek, namibia. komarek, e., 1972, ‘fire in africa’, in proceeding of the annual tall timbers fire ecology conference, 11, tall timbers research station, june 8–9, 1972, tallahassee, fl. kull, c.a., 2002, ‘madagascar a flame: landscape burning as peasant protest, resistance, or a resource management tool?’, political geography 21(7), 927–953. https://doi.org/10.1016/s0962-6298(02)00054-9 landry, j.s. & matthews, h.d., 2016, ‘non-deforestation fire vs. fossil fuel combustion: the source of co2 emissions affects the global carbon cycle and climate responses’, biogeosciences 13(7), 2137–2149. https://doi.org/10.5194/bg-13-2137-2016 laris, p., 2002, ‘burning the seasonal mosaic: preventative burning strategies in the wooded savanna of southern mali’, human ecology 30(2), 155–186. https://doi.org/10.1023/a:1015685529180 laris, p. & wardell, d.a., 2006, ‘good, bad or ‘necessary evil’? reinterpreting the colonial burning experiments in the savanna landscapes of west africa’, the geographic journal 172(4), 271–290. https://doi.org/10.1111/j.1475-4959.2006.00215.x lipsett-moore, g.j., wolff, n.h. & game, e.t., 2018, ‘emissions mitigation opportunities for savanna countries from early dry season fire management’, nature communications 9(1), 1–8. https://doi.org/10.1038/s41467-018-04687-7 martin, r.e. & sapsis, d.b., 1992, ‘fires as agents of biodiversity: pyrodiversity promotes biodiversity’, in proceedings of the conference on biodiversity of northwest california ecosystems, cooperative extension, university of california, berkeley, ca, october 28–30, 1991, pp. 150–157. masocha, m., skidmore, a.k., poshiwa, x. & prins, h.h.t., 2011, ‘frequent burning promotes invasions of alien plants into a mesic african savanna’, biological invasions 13(7), 1641–1648. https://doi.org/10.1007/s10530-010-9921-6 mccleery, r., monadjem, a., baiser, b., fletcher, r., vickers, k., kruger, l., 2018, ‘animal diversity declines with broad-scale homogenization of canopy cover in african savannas’, biological conservation 226, 54–62. mcgranahan, d.a. & kirkman, k.p., 2013, ‘multifunctional rangeland in southern africa: managing for production, conservation, and resilience with fire and grazing’, land 2(2), 176–193. https://doi.org/10.3390/land2020176 mentis, m.t. & bailey, a.w., 1990, ‘changing perceptions of fire management in savanna parks’, journal of the grassland society of southern africa 7(2), 81–85. https://doi.org/10.1080/02566702.1990.9648211 messina, m., cunliffe, r., farcomeni, a., malatesta, l., smit, i.p.j., testolin, r. et al., 2018, ‘an innovative approach to disentangling the effect of management and environment on tree cover and density of protected areas in african savanna’, forest ecology and management 419–420, 1–9. https://doi.org/10.1016/j.foreco.2018.03.019 meyer, c.p., cook, g.d., reisen, f., smith, t.e.l., tattaris, m., russell-smith, j. et al., 2012, ‘direct measurements of the seasonality of emission factors from savanna fires in northern australia’, journal of geophysical research: atmospheres 117(d20), 1–14. https://doi.org/10.1029/2012jd017671 moura, l.c., scariot, a.o., schmidt, i.b., beatty, r. & russell-smith, j., 2019, ‘the legacy of colonial fire management policies on traditional livelihoods and ecological sustainability in savannas: impacts, consequences, new directions’, journal of environmental management 232, 600–606. https://doi.org/10.1016/j.jenvman.2018.11.057 mulqueeny, c.m., goodman, p.s. & o’connor, t.g., 2010, ‘landscape-level differences in fire regime between block and patch-mosaic burning strategies in mkuzi game reserve, south africa’, african journal of range & forage science 27(3), 143–150. https://doi.org/10.2989/10220119.2010.527300 mulqueeny, c.m., goodman, p.s. & o’connor, t.g., 2011, ‘determinants of inter-annual variation in the area burnt in a semiarid african savanna’, international journal of wildland fire 20(4), 532–539. https://doi.org/10.1071/wf08141 munyati, c., shaker, p. & phasha, m.g., 2011, ‘using remotely sensed imagery to monitor savanna rangeland deterioration through woody plant proliferation: a case study from communal and biodiversity conservation rangeland sites in mokopane, south africa’, environmental monitoring and assessment 176(1–4), 293–311. https://doi.org/10.1007/s10661-010-1583-4 nangendo, g., stein, a., ter steege, h. & bongers, f., 2005, ‘changes in woody plant composition of three vegetation types exposed to a similar fire regime for over 46 years’, forest ecology and management 217(2–3), 351–364. https://doi.org/10.1016/j.foreco.2005.07.007 nangendo, g., ter steege, h. & bongers, f., 2006, ‘composition of woody species in a dynamic forest-woodland-savannah mosaic in uganda: implications for conservation and management’, biodiversity and conservation 15(4), 1467–1495. https://doi.org/10.1007/s10531-005-1876-z nieman, w.a., 2020, integrated fire management plan: majete wildlife reserve, malawi, technical report, african parks ltd, johannesburg. norton-griffiths, m., 1979, ‘the influence of grazing, browsing, and fire on the vegetation dynamics of the serengeti’, in a.r.e. sinclair & m. norton-griffiths (eds.), serengeti: dynamics of an ecosystem, pp. 310–352, university of chicago press, chicago, il. o’connor, t.g., puttick, j.r. & hoffman, m.t., 2014, ‘bush encroachment in southern africa: changes and causes’, african journal of range & forage science 31(2), 67–88. https://doi.org/10.2989/10220119.2014.939996 owen, j.s., 1972, ‘fire management in the tanzania national parks’, in tall timbers fire ecology conference proceedings, 11, tall timbers research station, june 8–9, 1972, tallahassee, fl. parr, c.l. & andersen, a.n., 2006, ‘patch mosaic burning for biodiversity conservation: a critique of the pyrodiversity paradigm’, conservation biology 20(6), 1610–1619. https://doi.org/10.1111/j.1523-1739.2006.00492.x parr, c.l. & brockett, b.h., 1999, ‘patch-mosaic burning: a new paradigm for savanna fire management in protected areas?’, koedoe 42(2), 117–130. https://doi.org/10.4102/koedoe.v42i2.237 parr, c.l., woinarski, j.c.z. & pienaar, d.j., 2009, ‘cornerstones of biodiversity conservation? comparing the management effectiveness of kruger and kakadu national parks, two key savanna reserves’, biodiversity and conservation 18(13), 3643–3662. https://doi.org/10.1007/s10531-009-9669-4 pfab, m.f. & witkowski, e.t.f., 2000, ‘a simple population viability analysis of the critically endangered euphorbia clivicola under four management scenarios’, biological conservation 96(3), 263–270. https://doi.org/10.1016/s0006-3207(00)00088-4 phillips, j.f.v., 1930, ‘fire: its influence on biotic communities and physical factors in south and east africa’, african journal of science 27, 352–367. pricope, n.g. & binford, m.w., 2012, ‘a spatio-temporal analysis of fire recurrence and extent for semi-arid savanna ecosystems in southern africa using moderate-resolution satellite imagery’, journal of environmental management 100, 72–85. https://doi.org/10.1016/j.jenvman.2012.01.024 rains, a.b., 1963, grassland research in northern nigeria 1952/62, samaru misc, paper no. 1, institute for agricultural research, ahmadu bello university, zaria. ribeiro, n.s., cangela, a., chauque, a., bandeira, r.r. & ribeiro-barros, a.i., 2017, ‘characterisation of spatial and temporal distribution of the fire regime in niassa national reserve, northern mozambique’, international journal of wildland fire 26(12), 1021–1029. https://doi.org/10.1071/wf17085 ross, i.c., field, c.r. & harrington, g.n., 1976, ‘the savanna ecology of kidepo valley national park, uganda: iii, animal populations and park management recommendations’, african journal of ecology 14(1), 35–48. https://doi.org/10.1111/j.1365-2028.1976.tb00150.x roux, e., 1969, grass, a story of frankenwald, oxford university press, cape town. russell-smith, j., monagle, c., jacobsohn, m., beatty, r.l., bilbao, b., millán, a. et al., 2013, ‘can savanna burning projects deliver measurable greenhouse emissions reductions and sustainable livelihood opportunities in fire-prone settings?’, climatic change 140(1), 47–61. https://doi.org/10.1007/s10584-013-0910-5 ryan, c.m. & williams, m., 2011, ‘how does fire intensity and frequency affect miombo woodland tree populations and biomass?’, ecological applications 21(1), 48–60. https://doi.org/10.1890/09-1489.1 sala, o.e., iii, f.s.c., armesto, j.j., berlow, e., bloomfield, j., dirzo, r. et al., 2000, ‘global biodiversity scenarios for the year 2100’, science 287, 1770–1774. https://doi.org/10.1126/science.287.5459.1770 sannier, c.a.d., taylor, j.c. & du plessis, w., 2002, ‘real-time monitoring of vegetation biomass with noaa-avhrr in etosha national park, namibia, for fire risk assessment’, international journal of remote sensing 23(1), 71–89. https://doi.org/10.1080/01431160010006863 saxon, e.c., 1984, anticipating the inevitable: a patch-burn strategy for fire management at uluru (ayers rock-mount olga) national park (nt), commonwealth scientific and industrial research organization, melbourne, australia. scholes, r.j. & archer, s.r., 1997, ‘tree-grass interactions in savannas’, annual review of ecology and systematics 28, 517–544. https://doi.org/10.1146/annurev.ecolsys.28.1.517 scholes, r.j. & walker, b.h., 1993, an african savanna: synthesis of the nylsvley study, cambridge university press, cambridge. scott, j.d., 1955, ‘principles of pasture management’, in d. meridith (ed.), the grasses and pastures of south africa, pp. 601–623, central news agency ltd., johannesburg. shaffer, l.j., hall, b. & street, j., 2010, ‘indigenous fire use to manage savanna landscapes in southern mozambique’, fire ecology 6(2), 43–59. https://doi.org/10.4996/fireecolgy.0602043 siegfried, w.r., 1981, ‘the incidence of veld-fire in the etosha national park, 1970–1979’, madoqua 12(4), 225–230. smit, i.p.j. & archibald, s., 2019, ‘herbivore culling influences spatio-temporal patterns of fire in a semiarid savanna’, journal of applied ecology 56(3), 711–721. https://doi.org/10.1111/1365-2664.13312 smit, i.p.j., asner, g.p., govender, n., vaughn, n.r. & van wilgen, b.w., 2016, ‘an examination of the potential efficacy of high-intensity fires for reversing woody encroachment in savannas’, journal of applied ecology 53(5), 1623–1633. https://doi.org/10.1111/1365-2664.12738 smit, i.p.j., smit, c.f., govender, n., van der linde, m. & macfadyen, s., 2013, ‘rainfall, geology and landscape position generate large-scale spatiotemporal fire pattern heterogeneity in an african savanna’, ecography 36(4), 447–459. https://doi.org/10.1111/j.1600-0587.2012.07555.x stander, p.e., nott, t.b. & mentis, m.t., 1993, ‘proposed burning strategy for a semi-arid african savanna’, african journal of ecology 31(4), 282–289. https://doi.org/10.1111/j.1365-2028.1993.tb00542.x stebbing, e.p., 1937, the forests of west africa and the sahara: a study of modern conditions, w & r chambers, london. stronach, n.r.h., 1988, the management of fire in serengeti national park, objectives and prescriptions, unpublished manuscript, tanzania national parks, arusha. stronach, n.r.h. & mcnaughton, s.j., 1989, ‘grassland fire dynamics in the serengeti ecosystem, and a potential method of retrospectively estimating fire energy’, journal of applied ecology 26(3), 1025–1033. https://doi.org/10.2307/2403709 te beest, m., cromsigt, j.p.g.m., ngobese, j. & olff, h., 2012, ‘managing invasions at the cost of native habitat? an experimental test of the impact of fire on the invasion of chromolaena odorata in a south african savanna’, biological invasions 14, 607–618. https://doi.org/10.1007/s10530-011-0102-z te beest, m., howison, o.e., howison, r.a., dew, l.a., poswa, m.m., dumalisle, l. et al., 2017, ‘successful control of the invasive shrub chromolaena odorata in hluhluwe-imfolozi park’, in j.p.g.m. cromsight, s. archibald & n. owen-smith (eds.), conserving africa’s mega-diversity in the anthropocene: the hluhluwe-imfolozi park story, p. 358, cambridge university press, cambridge. te beest, m., mpandza, n.j. & olff, h., 2015, ‘fire and simulated herbivory have antagonistic effects on resistance of savanna grasslands to alien shrub invasion’, journal of vegetation science 26(1), 114–122. https://doi.org/10.1111/jvs.12208 tokura, w., jack, s.l., anderson, t. & hoffman, m.t., 2018, ‘long-term variability in vegetation productivity in relation to rainfall, herbivory and fire in tswalu kalahari reserve’, koedoe 60(1), 1–18. https://doi.org/10.4102/koedoe.v60i1.1473 trapnell, c.g., 1959, ‘ecological results of woodland burning in northern rhodesia’, the journal of ecology 47(1), 129–168. https://doi.org/10.2307/2257252 trollope, l.a. & trollope, l.a., 2010, ‘fire effects and management in african grasslands and savannas’, range and animal sciences and resources management 2, 121–145. trollope, w., van wilgen, b., trollope, l.a., govender, n. & potgieter, a.l., 2014, ‘the long-term effect of fire and grazing by wildlife on range condition in moist and arid savannas in the kruger national park’, african journal of range & forage science 31(3), 199–208. https://doi.org/10.2989/10220119.2014.884511 trollope, w.s.w., 2007, ‘fire – a key factor in the ecology and management of african grasslands and savannas’, in proceedings of the 23rd tall timbers fire ecology conference: fire in grassland and shrubland ecosystems, tall timbers research station, tallahassee, florida, pp. 2–14. trollope, w.s.w., biggs, h.c., potgieter, a.l.f. & zambatis, n., 1995, ‘a structured vs a wilderness approach to burning in the kruger national park in south africa’, in n.e. west (ed.), rangelands in a sustainable biosphere, pp. 574–575, society for range management, denver. trollope, w.s.w., fyumagwa, r. & trollope, l.a., 2003, ‘relation between range condition and the incidence of ticks in the ngorongoro crater, tanzania’, in proceedings of the vii international rangeland conference, grassland society of southern africa, 26 july–1 august 2003, durban, south africa, pp. 531–533. trollope, w.s.w., trollope, l.a., de bruno austin, c., held, a.c., emery, a. & hines, c.j.h., 2006, a fire management plan for the okavango delta ramsar site in botswana, botswana department of forestry and range resources, gabarone. van der werf, g.r., randerson, j.t., giglio, l., collatz, g.j., mu, m., kasibhatla, p.s. et al., 2010, ‘global fire emissions and the contribution of deforestation, savanna, forest, agricultural, and peat fires (1997–2009)’, atmospheric chemistry and physics 10(23), 11707–11735. https://doi.org/10.5194/acp-10-11707-2010 van der werf, g.r., randerson, j.t., giglio, l., van leeuwen, t.t., chen, y., rogers, b.m. et al., 2017, ‘global fire emissions estimates during 1997–2016’, earth system science data 9(2), 697–720. https://doi.org/10.5194/essd-9-697-2017 van wilgen, b.w., 2009, ‘the evolution of fire management practices in savanna protected areas in south africa’, south african journal of science 5(9–10), 343–349. https://doi.org/10.4102/sajs.v105i9/10.107 van wilgen, b.w. & biggs, h.c., 2011, ‘a critical assessment of adaptive ecosystem management in a large savanna protected area in south africa’, biological conservation 144(4), 1179–1187. https://doi.org/10.1016/j.biocon.2010.05.006 van wilgen, b.w., biggs, h.c. & potgieter, a.l.f., 1998, ‘fire management and research in the kruger national park, with suggestions on the detection of thresholds of potential concern’, koedoe 41(1), 69–87. https://doi.org/10.4102/koedoe.v41i1.248 van wilgen, b.w, fill, j.m., govender, n. & foxcroft, l.c., 2017, ‘an assessment of the evolution, costs and effectiveness of alien plant control operations in kruger national park, south africa’, neobiota 35, 35–59. https://doi.org/10.3897/neobiota.35.12391 van wilgen, b.w., govender, n. & biggs, h.c., 2007, ‘the contribution of fire research to fire management: a critical review of a long-term experiment in the kruger national park, south africa’, international journal of wildland fire 16(2), 519–530. https://doi.org/10.1071/wf06115 van wilgen, b.w., govender, n., ntsala, d. & funda, x.n., 2004, ‘response of savanna fire regimes to changing fire-management policies in a large african national park’, conservation biology 18(6), 1533–1540. https://doi.org/10.1111/j.1523-1739.2004.00362.x van wilgen, b.w., govender, n., smit, i.p.j. & macfadyen, s., 2014, ‘the ongoing development of a pragmatic and adaptive fire management policy in a large african savanna protected area’, journal of environmental management 132, 358–368. https://doi.org/10.1016/j.jenvman.2013.11.003 van wilgen, b.w., govender, n. & macfadyen, s., 2008, ‘an assessment of the implementation and outcomes of recent changes to fire management in the kruger national park’, koedoe 50(1), 22–31. https://doi.org/10.4102/koedoe.v50i1.135 van wilgen, b.w., trollope, w.s.w., biggs, h.c., potgieter, a.l.f. & brockett, b.h., 2003, ‘fire as a driver of ecosystem variability’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 149–170, island press, washington, dc. verbesselt, j., somers, b., lhermitte, s., jonckheere, i., van aardt, j. & coppin, p., 2007, ‘monitoring herbaceous fuel moisture content with spot vegetation time-series for fire risk prediction in savanna ecosystems’, remote sensing of environment 108(4), 357–368. https://doi.org/10.1016/j.rse.2006.11.019 verbesselt, j., somers, b., van aardt, j., jonckheere, i. & coppin, p., 2006, ‘monitoring herbaceous biomass and water content with spot vegetation time-series to improve fire risk assessment in savanna ecosystems’, remote sensing of environment 101(3), 399–414. https://doi.org/10.1016/j.rse.2006.01.005 vincent, j., 1970, ‘the history of umfolozi game reserve, zululand, as it relates to management’, lammergeyer 11, 7–48. walter, h., 1971, ecology of tropical and subtropical vegetation, oliver & boyd, edinburgh. wardell, a., nielsen, t.t., rasmussen, k. & mbow, c., 2004, ‘fire history, fire regimes and fire management in west africa: an overview’, in j. godammer & c. de ronde (eds.), wildland fire management handbook for sub-sahara africa, pp. 350–382, global fire monitoring center, freiburg. west, o., 1955, ‘veld management in the dry, summer-rainfall bushveld’, in d. meridith (ed.), the grasses and pastures of south africa, pp. 624–636, central news agency ltd., johannesburg. west, o., 1972, ‘fire, man and wildlife as interacting factors limiting the development of vegetation in rhodesia’, in proceedings of the tall timbers fire ecology conference, 11, tall timbers research station, june 8–9, 1972, tallahassee, fl, pp. 121–146. witt, a.b.r., kiambi, s., beale, t. & van wilgen, b.w., 2017, ‘a preliminary assessment of the extent and potential impacts of alien plant invasions in the serengeti-mara ecosystem, east africa’, koedoe 59(1), 1–16. https://doi.org/10.4102/koedoe.v59i1.1426 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 grant.qxd challenges faced in the conservation of rare antelope: a case study on the northern basalt plains of the kruger national park c.c. grant, t. davidson, p.j. funston and d.j. pienaar grant, c.c., t. davidson, p.j. funston and d.j. pienaar. 2002. challenges faced in the conservation of rare antelope: a case study on the northern basalt plains of the kruger national park. koedoe 45(2): 45–62. pretoria. issn 0075-6458. the conservation of rare antelope has long been one of the goals of the kruger national park. the roan antelope hippotragus equinus, and to a lesser extent the tsessebe damaliscus lunatus, represent low-density species or rare antelope in the park. specific management approaches representing the older equilibrium approach, have been employed to conserve these antelope. of these, the supply of artificial water over many decades was the most resource intensive. the sudden, severe drop in the roan antelope population towards the end of the 1980s was unexpected and, retrospectively, attributed to the development of a high density of perennial waterpoints. the postulated mechanism was that the perennial presence of water allowed burchell’s zebra equus burchelli to stay permanently in an area that was previously only seasonally accessible. the combined effect of a long, dry climatic cycle, high numbers of zebra and their associated predators was proposed to be the cause of this decline. as part of the new nature evolving or ecosystem resilience approach, twelve artificial waterpoints were closed in the prime roan antelope habitat in 1994 in an attempt to move the zebra out of this area. the zebra numbers declined as the rainfall increased. closure of waterholes clearly led to redistribution of zebra numbers on the northern plains, zebra tending to avoid areas within several kilometres of closed waterpoints. however, at a larger scale, regional densities appeared similar in areas with and without closed waterpoints. there was an initial drop in the lion numbers in 1995, after which they stabilised. in spite of an improvement in the grass species composition and an increase in biomass the roan antelope population did not increase. the complexity of maintaining a population at the edge of their distribution and the problems associated with the conservation of such populations are discussed in terms of management options and monitoring approaches that may be employed in this process. key words: artificial waterpoints, management, predation, drought, zebra, rare antelope. c.c. grant and t davidson northern plains programme of the kruger national park, private bag x402, skukuza 1350 republic of south africa; p.j. funston, department of nature conservation, pretoria technikon, private bag x680, pretoria, 0001 republic of south africa; d.j. pienaar, south african national parks, conservation services, private bag x402, skukuza 1350 republic of south africa. issn 0075-6458 45 koedoe 45/2 (2002) introduction in a review of the large mammal distribution and status in the kruger national park (knp), pienaar (1963) described the roan antelope as one of the few species that had not reacted favourably to conservation actions. the population of roan antelope hippotragus equinus had oscillated between 150 and 300 over about 60 years since the proclamation of the kruger national park. sable antelope hippotragus niger were found to be very susceptible to drought conditions and tsessebe populations damaliscus lunatus were only growing favourably in the area north of the letaba river at the time of this review. these species have received special grant.qxd 2005/12/09 10:17 page 45 koedoe 45/2 (2002) 46 issn 0075-6458 table 1 summary of two management approaches the nature balanced pre-1994 and nature evolving post1994(holling & gunderson 2002). most of these were employed over the whole knp, although the effects on the northern basalt plains (nbp) is emphasised especially for the post-1994 period management option water provision disease control fire animal control management action pre-1994 to animals whose migration routes out of the knp were cut off by fencing (pienaar 1985). in forage-rich areas that would otherwise be inaccessible to water-dependent species (pienaar 1963). as a tool to increase and stabilise numbers of rare antelope (nature conservation year report of 1961/1962 (anon 1962) roan antelope vaccinated against anthrax 1971 – 1992 on the nbp only (de vos et al. 1973) the first boundary fence was erected in 1957 and the last elephant proof fence was completed in 1976. no management fires till 1912 biennial burns till 1946 five yearly burns till 1958 triennial burns till 1970 according to biomass till 1992 (van wilgen et al. 2000) carnivore control at various stages 1912 to late 1920s, late 1940s to 1961, mid 1970s (when the zebra and wildebeest numbers dropped severely) (joubert 1986). zebra and wildebeest culled in 1960s (joubert 1986). culling of buffalo 1966 – 1991. culling of elephant 1967 – 1994. expected outcome stabilising numbers of migrating animals especially zebra and wildebeest. wider utilisation of available forage and an increase in herbivore numbers. increase and stabilisation of numbers of rare antelope. decrease losses of roan antelope due to anthrax epidemics. fence to keep diseases out and to avoid diseases being transferred to domestic stock (joubert 1986). provide green grazing for animals. to allow herbivore numbers to increase. to avoid overutilisation of the vegetation during low rainfall years. to avoid overutilisation of the vegetation. to avoid overulitlisation of the vegetation. water provision / ..... grant.qxd 2005/12/09 10:17 page 46 attention from management since 1963 (grant & van der walt 2000). the different scenarios that could have led to the poor population growth of the roan antelope were discussed in detail in the nature conservation year report of 1961–1962 (anon 1962). the unsuitability of the habitat was cited as the most likely limitation to population growth. disease, specifically anthrax, and a decrease in the vitality of animals due to inbreeding were also mentioned, as was the possible role of predation. roan antelope in the knp have always been concentrated on the northern basalt plains (nbp) at the northeastern tip of the park. a substantial herd of 10–20 roan antelope was also present in the southern granites in the pretoriuskop area from 1960 to 1985, according to ranger reports. the area to the north of the nbp has some species more characteristic of the mesic savannas, such as nyala tregalaphus angasi (pienaar 1974), which commonly occur along large drainage lines and rivers. seasonal rivers and the broad, marshy drainage lines form corridors connecting the nbp to similar habitats to the north and east. these low-lying areas mostly consist of dambo-like grasslands similar to the habitats utilised by roan antelope and tsessebe in mozambique and zimbabwe (joubert 1976). smithers (1983), states that the habitat requirements of tall grass cover and available surface water are essential to roan antelope and their occurrence is confined to areas with suitable habitat. tsessebe issn 0075-6458 47 koedoe 45/2 (2002) table 1 (continued) management option water provision disease control fire animal control management action post 1994 close 12 waterpoints on the nbp (grant 1999) stopped anthrax vaccination in 1991 western boundary opened in the south of the knp start with a patch mosaic burning system in 2002 (van wilgen et al. 2000) stopped all culling in 1998 (elephant are still being removed) second enclosure was built in 2002. expected outcome zebra will move out of the area. this will allow an increase in roan antelope due to improved vegetation. predators will decrease following the decline in zebra decrease the splitting of the roan antelope herds in response to the darting and hence enhance the growth of the population a larger area available to the rare antelope may enhance their populations provide green forage to tsessebe and sable (gureja 2001) animal populations will settle at optimal densities for the landscape and rainfall. this will increase the breeding potential of the captive roan antelope and thus the number that can be released in a group (grant & van der walt 2000). grant.qxd 2005/12/09 10:17 page 47 generally prefer short grassland (child et al. 1972; joubert & bronkhorst 1977) and are attracted to the green shoots appearing after fires in the presence of sufficient soil moisture (garstang 1982). tsessebe are highly dependent on surface water within open woodland. bush encroachment accounts for reductions in their range and numbers (smithers 1983). tsessebe, and to a lesser extent, roan antelope, show a specific preference for the more open ecotones between the tall woodland and the open grassland of the vleis (joubert 1970). the ecotone area also forms an important forage area for the large variety of other grazers that occur in the area, of which burchell's zebra equus burchellii and buffalo syncerus caffer make up the largest percentage (pienaar 1963). sable antelope are a savanna woodland species, dependent on cover and availability of water (smithers 1983), preferring the combretum/mopane woodland with high trees and large canopies in the granitic western half of the knp (pienaar 1974). the nature conservation year report of 1961–1962 (anon 1962) discusses possible management approaches to enhancing the roan antelope population and provides a good illustration of the equilibrium resilience approach (holling & meffe 1996) of the time. these were supplementation of the knp roan from namibia, the development of dams along the eastern border to attract some of the remaining large roan herds from the adjacent part of mozambique, koedoe 45/2 (2002) 48 issn 0075-6458 fig. 1. location of geographical zones in the granite and basalt landscapes relative to the main rivers. ������ ������ ������ ������ ��� ��� ��� ��� � �� � � � �� ������ ���������������������� ������ � ������� � � ������� ������� � �������������� � ����������� �� � �� ��� ����� �� �������������� � ��� � �������� � ��� � �������� � ��������� � ��� � �������� � ��������� � ��� �� grant.qxd 2005/12/09 10:17 page 48 and the creation of more suitable habitat by clearing the drainage lines of mopane trees. this paper aims at providing an historical overview of management approaches used in the knp and northern basalt plains (nbp) from 1960 to 2000, and the associated changes in animal numbers and vegetation. the definition of holling & gunderson (2002) of two basic management approaches is used (table 1). 1. a structurally static view of "nature balanced", which concentrated on conserving the elements of the ecosystem by assuming overall equilibrium (holling & meffe 1996). this approach was followed from about 1960 to 1994 (joubert 1985). we postulate that as predicted by holling & meffe (1996) this approach led to a reduction in the resilience of the system as a result of limiting natural variation. 2. from 1994 onwards, the more dynamic view of "nature evolving" was adopted (braack 1997) and was aimed at conserving the ability to adapt to change, thus "creating options to buffer disturbance and to create novelty". only the initial results of this approach on the nbp of the knp are discussed here. as part of this discussion; the importance of monitoring and adaptive management are highlighted. although it is true that management was adapting their approach before 1994, this year was taken as the dividing line because it was then that the first consciously "destabilising" management action was taken (one promoting variability), viz., the closing of waterpoints study area the kruger national park is coarsely divided longitudinally into basaltic (eastern) and granitic (western) zones (gertenbach 1983), and the vegetation is broadly defined as an arid/eutrophic savanna (huntley 1982). the area north of the olifants river is dominated by mopane, while the vegetation in the southern half of the park consists mainly of broad-leaved combretum species on the granite crests and acacias in the valleys. the southern basalts are dominated by knobthorn acacia nigrescens and marula sclerocarya birrea (fig. 1). the study area of the nbp stretches over an area of 700 km² with the shingwedzi river in the south, the punda hills in the north, the basalt/granite division in the west, with the nwambiya sandveld in the east. the vegetation is classified as mopane (colophospermum mopane) shrubveld on basalt (gertenbach 1983). for comparative purposes, the knp is divided into four granitic and four basaltic zones, with the large rivers (shingwedzi, olifants and sabie) forming the zone boundaries (fig. 1). issn 0075-6458 49 koedoe 45/2 (2002) fig. 2. percentage deviation from average rainfall at shingwedzi between 1971 and 1999. year on x-axis is the year the rainfall season ends 30 june. grant.qxd 2005/12/09 10:17 page 49 this summer rainfall area has a mean annual rainfall of 477 mm north of the olifants river, measured over 32 years at 10 rainfall stations, and 560 mm south of the olifants river measured at 12 rainfall stations over 46 years (zambatis & biggs 1995). in fig. 2, the percentage deviation from the mean annual rainfall is shown at shingwedzi over the station's recording period. at shingwedzi, a wet cycle occurred in the seventies followed by a drier cycle in the eighties and nineties. here the average rainfall decreased from 551 ± 65 mm between 1977 and 1986 to 338 ± 60 mm between 1987 and 1993, with 344 ± 76 mm between 1994 and 1998. the mean knp rainfall was also significantly higher during the period 1977 to 1986 (561 ± 18 mm) than between 1987 and 1993 (423 ± 18 mm) and between 1994 and 1998 (357 ± 39 mm) (p < 0.0005). development of waterpoints during the four decades between 1939 and 1980 there was a steady increase in artificial water supply, with the largest number of windmills erected between 1960 and 1980 koedoe 45/2 (2002) 50 issn 0075-6458 fig. 3. development of waterpoints in different geographical zones between 1950 and 2000. (fig. 3). on the nbp a total of 35 active boreholes plus six dams were established by 1975. this development resulted in a significantly higher density of waterpoints on the nbp than all the other areas over 18 years (p < 0.0005) with an average of 41 km² per borehole over that period. the density of boreholes present in 1986 at the start of the roan antelope decline in the different geographical zones is listed in table 2. closure of waterpoints during 1994, 12 waterpoints were closed, over an area of 412 km² in the northern part of the nbp, where the only remaining roan herds were left. the windmills in a comparative area of 360 km² to the south remained open. one critically placed dam was breached in june 1996. in march 1998 another waterpoint further south was closed, and in march 1999 another two. fences the boundary fences precluded any movement of animals from or towards habitats in grant.qxd 2005/12/09 10:17 page 50 mozambique and zimbabwe where roan antelope, sable antelope and tsessebe were once common (anon 1962). thus within the knp the rare antelope were restricted to the southern boundary of their distribution. anecdotal information provided in a report by the park biologist (anon 1961) indicate that there was movement across the knp boundary before the fences were erected, some of the roan herds further south being split by the western boundary fence. in 1995, the south-western boundary fence between the knp and the private reserves was removed, and in 2002, negotiations to remove the fence between the north-eastern border of the knp and mozambique were finalised (gaza-kruger-gonarezhou transboundary natural resources management initiative 2002). anthrax vaccination anthrax caused the death of at least 83 roan antelope (over 11 years, during epidemics in 1959, 1960 and 1970) out of an estimated mean population of 250 north of the olifants river (de vos et al. 1973). during the same period only eight sable were recorded to have died of anthrax. anthrax was thus idenissn 0075-6458 51 koedoe 45/2 (2002) table 2 the area of the different geographical zones and the density of boreholes in 1986 description size borehole in km² density in km²/borehole in 1986 far northern granites 1777.4 62 northern basalt plains (nbp) 1328.1 37 northern granites 2003.1 68 mooiplaas basalts 1838.5 61 olifants transition zones 1791.4 74 central basalts 3001.9 132 central granites 2570.1 64 southern basalts 1739.7 140 southern granites 1811.7 50 tified as a major threat to the small roan population and the decision was taken to vaccinate them. roan antelope were immunised annually between 1971 and 1992, on the nbp, but not on the mooiplaas basalts which were further from the main focus of infection along the levuvhu river. projectile syringes were used from 1971 until 1984, after which bio-bullets were used. vaccination was stopped in 1992 because it was believed that the immunisation procedure scattered the few remaining roan herds and made them more vulnerable. roan enclosure because of the possibility of losing all the roan antelope in the park, seven roan antelope including three males (one adult) and four females (two adult) were placed in the existing n'washitshumbe enclosure between june and august 1994. a single calf was born in december 1994. this enclosure of 254 ha (which excludes large predators and all other herbivores) was erected roughly in the centre of the nbp in 1967 (joubert 1970) and expanded in 1986 with 48 ha to include a part of the adjacent watercourse and associated ecotone vegetation. initiation of a specific research programme (northern plains programme). this programme was initiated in 1994 and aimed at evaluating the effects of the closure of waterpoints (harrington & pienaar 1994). it was postulated that the decline in roan antelope was related to the increase in the zebra population that was responding to an increased density of artificial waterpoints, and that predation of adult roan antelope also played a significant role in the decline (mills et al. 1995a; owen-smith 1996; harrington et al. 1999). the initial studies thus concentrated on the effect of closure of waterpoints on vegetation, roan antelope, zebra and lion panthero leo populations. grant.qxd 2005/12/09 10:17 page 51 monitoring aerial census from 1980 until 1994, a fixed-wing aircraft was used to provide a total aerial coverage of the knp between may and august every year, using four observers. from a height of 65–70 m above ground level, parallel strips of about 800 m wide (400 m each side) were flown (viljoen 1996), covering the entire kruger national park. on the nbp this census was continued between 1994 and 1998 while only incomplete census counts were done in the rest of the knp. in 1999 and 2000 a 50 % sample of the nbp and a 17 % sample of the rest of the knp was counted using the distance sampling method (buckland et al. 1993). buffalo counts were done using a helicopter, by splitting herds into smaller groups and then photographing them. animals on the photographs were then counted with visual aids. vegetation condition assessment range condition has been assessed yearly at 18 specific localities on the nbp since 1989, using a standard range condition assessment (vca) technique (trollope et al. 1989; trollope 1990). at each point the grass species were recorded, and the frequency of each species was expressed as the percentage of the total number of grasses. grasses were grouped into decreasers (grasses that decrease with intense utilisation by herbivores) and increasers (species that increase with intense utilisation by herbivores). fifteen new vca localities were added on the nbp in 1994 to increase the sampling intensity in the area where the waterpoints were closed. forage utilisation by roan antelope roan antelope were located by radio-tracking between march and october 1996. starting at the point where they were recently found to be grazing, a standard range assessment (trollope et al. 1989) was done at 60 sites along four diagonal transects of 15 m each in quadrats of 20 cm². in addition to the standard measurements, leaf height, compressed height as determined with the disc pasture meter, grass species utilised, and percentage utilisation was recorded. the effect of artificial waterpoints on surrounding vegetation the effect of the presence of artificial water on the surrounding vegetation was examined by comparing the vegetation in the area of open windmills to that in the area of closed windmills. recovery of the herbaceous layer was aimed at determining how the herbaceous layer around three artificial waterpoints improved after decreasing herbivore pressure, brought about by the closure of the waterpoints (davidson 1996). this study was expanded to look at the spatial structure of the herbaceous vegetation under different grazing intensities, as well as the species association (jacoby 1999). a further study was undertaken to examine changes in woody composition around waterpoints. for this study vegetation was examined around four open waterpoints, and four closed (fruhauf 1997). herbivore studies rare antelope studies studies on roan antelope (joubert 1976), tsessebe (joubert 1972) and sable (joubert 1976) were undertaken to establish their habitat preferences. joubert (1970) investigated the social behaviour of roan antelope. in 1994, two cows in each of the two largest roan antelope herds were fitted with radio collars. once a signal was detected from an elevated point, the animal was followed by vehicle and its position recorded. the distance to the nearest waterpoint, and the position in the landscape according to the land unit classification (venter 1990) was also recorded. as far as possible, the number, sex, and age-class distribution was determined for each located group or herd. due to poor visibility the whole herd was often not seen, and some young calves could have been missed. monthly reports of rangers with observations of rare antelope were used for data up to 1984, after which data came from mason (1990) up to 1990. no data is available for 1990–1994. zebra observations zebra activity and position in the landscape was noted when they were encountered during roan radio-tracking sessions. in the case of zebra, the position in the landscape was recorded in only two categories—upland and lowland. thirty zebra females from separate herds in the study area were fitted with colour-coded collars in november 1993. visitors and staff members, whose koedoe 45/2 (2002) 52 issn 0075-6458 grant.qxd 2005/12/09 10:17 page 52 involvement was encouraged by posters, recorded sightings of collared zebra on forms that were placed in the reception area of major rest camps. predator studies lion census in order to determine lion densities, the mass capture technique of smuts et al. (1977a, 1977b) was employed to attract lions to calling stations in 1989. this census was repeated in 1993, 1996, 1998 and 2002, with slight changes in the locality of the survey points. the area covered by this census technique is not constant as lions are drawn from territories outside the designated study area. also, no statistical tests can be used to compare surveys (mills pers. comm.). data analysis herbivore numbers animal counts and densities at different periods and in different geographical zones were normalised for statistical analysis by natural logarithms. animal populations in different areas were compared in a multiple anova, correcting for the effect of rainfall by including the annual rainfall for each geographiissn 0075-6458 53 koedoe 45/2 (2002) fig. 4. zebra and buffalo numbers in the knp from total aerial counts for buffalo and for zebra between 1977 and 1993. (counts between 1998 and 2001 are estimated from a 17 % and 22 % sampling, respectively, according to the distance procedure (buckland et al. 1993). fig. 5. numbers of common herbivores on the northern basalt plains from total counts between 1980 and 1999. cal zone as covariate. standard error is presented in brackets in tables and text. area per animal and area per borehole were calculated by dividing the area size by the number of animals or boreholes. roan herd totals, according to ranger reports, mason's reports, and data collected by the research programme since 1994, were grouped as follows: 1950-1964; 1965-1969; 1975-1985; 1985-1994; 1986-1994; and 1995-2001. vegetation the chi-square test was used to test the difference in abundance of species on the vca plots compared to the areas where roan antegrant.qxd 2005/12/09 10:17 page 53 lope preferred to graze. the selection of specific habitats by roan antelope compared to the general area covered by each habitat as estimated in venter's (1990) land unit classification was tested in the same way. differences in vegetation condition parameters between geographical zones north of the olifants river were determined in a multiple anova using standing crop as indicator of biomass and percentage of decreasers and increasers as indicators of forage quality. percentage of decreasers and increasers were normalised, using the square root arcsine transformation. to test the effect of the closure of waterpoints on the vegetation condition parameters, the 1999 vca results were corrected for the 1994 situation by using the 1994 data as a covariate. koedoe 45/2 (2002) 54 issn 0075-6458 fig. 6. rare antelope numbers in the knp between 1977 and the last total aerial census in1995 and on the nbp between 1977 and 1999. bars are roan on nbp; solid symbols are rare antelope on nbp; open symbols are rare antelope in knp. table 3 the mean density in the geographical zones between 1980-1993 of zebra in the knp and the density of rare antelope north of the olifants river (standard error in brackets) zone n km²/zebra km²/roan km²/sable km²/tsessebe far northern granites 13 0.84 abc (0.07) 186 ab (74) 8.09 a (13.8) 61.96 ab (41.8) northern basalt plains 22 0.67 ab (0.07) 22 a (57) 13.5 ab (11.4) 14.5 ab (32.1) northern granites 19 0.99 abc (0.07) 190 abc (61) 7.4 a (11.38) 55 ab (34.5) mooiplaas basalts 19 0.56 b (0.07) 35 ab (61) 67.9 abc (11.39) 5.5 a (34.6) olifants transition zone 18 0.60 b (0.07) 21.9 abc (11.7) central basalts 16 0.51 a (0.074) central granites 16 0.48 a (0.074) southern basalts 16 1.39 abc (0.076) southern granites 16 1.69 abc (0.074) significance within column p < 0.0005 p < 0.0005 p < 0.0005 p < 0.0005 significance of rainfall p < 0.0005 p = 0.003 ns ns as covariate letters in superscript and upper case indicate a significant difference with the same letter in superscript and lower case. thus value a differs significantly form value a in the same column. grant.qxd 2005/12/09 10:17 page 54 results of management strategy up to 1994 herbivore numbers zebra numbers in the knp showed a steady increase since 1977 when the aerial counts started, levelling off after 1985. the buffalo population was fairly stable due to culling and then declined sharply after the 1991 drought (fig. 4). there was a corresponding steady increase in the zebra population on the nbp until 1991, a year in which rainfall was very low (fig. 5). the population of blue wildebeest connochaetes taurinus did not increase as much as the zebra, and issn 0075-6458 55 koedoe 45/2 (2002) fig. 7. changes in biomass (standing crop) and percentage palatable, perennial decreaser species between 1989 at the start of the vegetation surveys and 1999. table 4 difference in vegetation parameters in 1994 in the different geographical zones n = 435 geographical zone biomass kg/ha % decreasers % increasers far northern granites 1935 ac 13.73 b 85.2 abc northern basalt plains 3002 abc 22.6 ab 66.5 ac northern granites 739 a 6.8 a 90 abc mooiplaas basalts 1191a 12.96 b 83.1 abc olifants transition zone 736 a 10.3 a 82 abc central basalts 1865 ac 17.72 ac 71.7 ac central granites 1743 ab 19.25 ac 65.8 ab southern basalts 2389 ab 23.43 ab 71.5 ac southern granites 1926 ac 26.28 abc 54.3 a letters in superscript and upper case indicate a significant difference with the same letter in superscript and lower case. thus value a differs significantly form value a in the same column. biomass (p = 0.0000), percentage decreaser species (p = 0.0000) and percentage increaser species (p = 0.000) differed significantly. grant.qxd 2005/12/09 10:17 page 55 decreased substantially after 1991. the buffalo population on the nbp showed wide fluctuations due to movement in and out of the study area, high numbers being present during the census period in 1991. the numbers of the strongly water-associated waterbuck kobus ellipsiprymnus varied with the rainfall, but stayed low after the 1991–1992 drought. sable, tsessebe and especially roan antelope declined dramatically since 1986 in the kruger national park. even in the south where the dry season rainfall is generally higher (zambatis & biggs 1995), the pretoriuskop herd disappeared in 1985. this decline was as evident in the nbp, the prime habitat of roan antelope and tsessebe (fig. 6), although the nbp still remained one of the areas with the highest rare antelope populations (table 3). overall the high zebra numbers did not show the expected negative correlation with the roan antelope population; indeed there was a small but significant positive correlation (r = 0.18, n = 168, p = 0.018) between roan numbers and zebra numbers in different zones of the park. on the nbp, this relationship seemed numerically stronger but not significant (r = 0.31, n = 31, p = 0.08). vegetation the northern and southern basalts had the highest biomass. percentage decreasers were the highest in the southern granites and basalts, and in the northern basalts (table 4). the low rainfall in 1992–1993 resulted in the lowest recorded biomass and percentage decreaser species (fig. 7). disease management the effect of vaccination was examined by looking at the density of roan antelope numbers on the plains, compared to the density in the adjacent mooiplaas basalts, where the roan were not vaccinated at any stage. the density of roan antelope on the nbp of 14.8 (± 40) km²/roan between 1980 and 1992 did not differ significantly from the 28 (± 40) km²/roan in the mooiplaas basalts during this vaccination period. the density of roan antelope after the decline between 1993 and 1995 did not differ significantly. the density was 51 (± 58) km²/roan after vaccination stopped on the nbp, versus 98 (± 82) km²/roan on the mooiplaas basalts. koedoe 45/2 (2002) 56 issn 0075-6458 fig. 8. herd size of roan antelope (bars) and juvenile to adult female ratios (line) in the northern knp from 1984 to 1999 for two sets of data. grant.qxd 2005/12/09 10:17 page 56 issn 0075-6458 57 koedoe 45/2 (2002) � �� ���� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� ���� �� ���� ���� �� �� �� �� �� �� � ��� �� �� �� �� �� �� ���� �� �� �� �� �� �� �� ���� �� �� ���� �� �� �� � � �� �� �������� ������ �� �� ���� �� �� �� �� �� ���� �� �� �� � ������� � � ���� �� �� �� �� �� �� ���� �� �� ���� �� ���� ���� ������ ���� �� �� �� �� �� �� � � ������ �� ��� � �� �� �� ���� �� �� �� �� � � ������ �� �� �� �� �� ������ �� � � �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� ������ �� �� �� ���� �� � ��� �� �� �� ���� � � �� �� ���� �� �� �� �� �� �� �� �� ���� �� �� �� �� �� �� �� �� �� ������ �� �� � � � � � ��� � ��� �� �� �� ���� �� �� �� �� �� �� � ��� �� �� �� �� �� �� �� �� �� �� � � �� �� �� �� �� �� �� �� �� �� ���� �� �� �� �� �� ���� ���� �� �� �� �� �� � � �� ���� �� � ��� �� �� �� �� �� �� �� �� �� ���� �� �� �� �� � � � � � � � � � � � � � �� � � � � � � �� �� �� �� �� � � � � � � � � � � � � � � � � � � � � �� �� �� �� �� �� � �� �� �� �� �� �� �� �� � �� � ��� �� �� �� � � �� �� �� �� ��� �� � � �� ��� �� �� � � � � �� �� � �� � �� � � �� �� �� �� � �� �� � � � � �� �� � �� �� � �� � � � ���� ���� �� �� �� ���� �� �� �� �� ���� �� �� ���� ���� �� �� �� � ��� �� � � ���� �� � � �� �� ���� �� �� �� �� �� �� ���� �� �� �� �� ���� ���� �� �� �� �� �� ���� �� �� �� �� �� ���� �� ���� �� � � �� �� ���� ���� �� �� ���� �� �� ���� �� �� �� �� �� �� ���� �� � � � � � � � � � � � � � �� � � � � � � �� �� �� �� �� � � � � � � � � � � � � � � � � � � � � �� �� �� �� �� �� �� �� �� �� �� �� �� � � �� �� ���� �� �� �� ���� �� ��� � �� �� �� �� ��� � �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� ���� � � ���� �� � ��� �� � � �� �� ���� �� �� �� �� � ���� � �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� ���� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� ���� �� ���� �� �� �� �� �� �� �� �� �� �� �� �� �� �� � � �� �� �� �� �� � � �� �� �� �� �� ���� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� ���� ���� �� �� �� �� ���� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� ���� �� �� �� ���� �� �� �� �� �� �� �� ���� �� �� �� �� �� �� ���� �� �� �� �� �� �� �� �� �� �� �� �� ���� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� ���� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� �� � � � � � � � � � � � � � �� � � � � � � �� �� �� �� �� � � � � � � � � � � � � � � � � � � � � �� �� �� �� �� �� �� �� ��� �� �� ��� � �� �� ��� �� � � �� ��� �� �� ��� �� � � � fi g. 9 . d is tr ib ut io n of z eb ra in 1 99 3, b ef or e cl os ur e of w at er po in ts in 1 99 4. t he s ub se qu en t m ov em en t a w ay f ro m th e ce nt ra l a re a w he re w at er po in ts w er e cl os ed is il lu st ra te d— tw o ye ar s af te r cl os ur e in 1 99 6 an d fo ur y ea rs a ft er c lo su re in 1 99 8. t he m ov em en t o f co lo ur -c ol la re d ze br a to w ar ds th e so ut h is in di ca te d on th e 19 96 m ap . grant.qxd 2005/12/09 10:17 page 57 koedoe 45/2 (2002) 58 issn 0075-6458 fig. 10. density of zebra in the open and closed area of the northern plains. recorded between 1980 and 1985, while the average was 8.8 between 1957 and 1964, and 6.3 between 1986 and 1994. the juvenile to adult female ratios of the roan antelope were very high between 1984 and 1989, but then decreased to levels recorded before 1984 (fig. 8). results of management actions after 1994 on the northern basalt plains the effect of waterpoint closure on the herbivores there was a general decline in herbivores on the nbp that, however, started before the closure of waterpoints in 1994 (fig. 5). thus the total number of herbivores (zebra, buffalo, wildebeest and waterbuck) counted (2573 ± 458) between 1995 and 2000 is significantly less than the numbers of these her� � � �� �� �� � �� � � � � � � � � � � �� � � � � � � � � �� �� � � � � �� � � � �� �� � � � � � � � � � � � �� � � � � � � � � �� � ������ � ������ �� �� ���� � � � � �� �� � � ��� ����� �� � � �� �� �� �� �� �� �� �� �� �� �� ���� �� �� �� �� � �� � �� � �� �� �� �� �� �� �� �� �� �� �� �� �� � � ���� �� �� � �� �� � �� � � �� � � � � ���� �� ���� ��� � � � �� �� ��� �� � � � ������� ��� ��� ���� ������ � � �� � � � � � � �� � � � �� � � � �� � ������ � ������ �� �� ���� � � � � �� ��� � � ������� �� � � � � � � � � � � � � � � � � � � �� � �� �� ������ � � � �� ����� �� ��� ���� ���� ���� ���� ���� � � � � � ���� � � ��� � � �� ������ �� ���� � ������ �� � ����� �� �� ������� ������ � � fig. 11. the relation of open and closed waterpoints on the nbp to the distribution of radio-collared roan antelope between 1995 and 1999, and the activity zones described in 1976. rare antelope on the plains, the number of breeding herds of roan antelope—with more than six individuals per herd—had decreased from four in 1970 (joubert 1970), to two in 1984. herd sizes differed significantly between the approximate ten-year periods since 1957 (p = 0.003). the largest average herd size in this area was 9.4 (fig. 8) in the north, grant.qxd 2005/12/09 10:17 page 58 bivores on the nbp between 1980 and 1994 (6552 ± 348) (p < 0.0005) (fig. 5). zebra continued their decline with the increase in rainfall, as expected (mills et al. 1995), and moved away from the core area where waterpoints were closed towards the open waterpoints and concentrated along drainage lines (fig. 9). however, the decline in the open and closed areas was very similar (fig. 10). tsessebe populations declined sharply after closure of waterpoints in 1994, from 98 on the nbp in 1993, to between 20 and 40 in 1998 (fig. 6). during the same period the sable decreased from 60 in 1993, to 3 in 1998 (fig. 6), probably due to movements westward to the granites. the roan antelope population stabilised at about 21 animals after 1994, but did not increase (fig. 6). positions where radio-collared roan antelope were recorded during the study period mostissn 0075-6458 59 koedoe 45/2 (2002) table 5 selection of habitats by roan relative to the area covered by the land type habitat % roan habitat as % encounters of mooiplaas land typea crest 35 77 midslope 25.8 10 low lying area 19.5 7 drainage lines, 19.5 6 vleis and ecotones aaccording to venter (1990). ly fell within the home range boundaries of territories established by joubert (1976). very few animals were sighted in the southern part where waterpoints were still open (fig. 11). furthermore, the herd of 20 roan antelope that was recorded in 1981 near the langtoon dam (in the south-eastern corner of the closed area) with a permanent zebra herd, had declined to only five animals in 1994, and disappeared towards the end of 1997. growth of population in the n’washitshumbe enclosure compared to that in the free-ranging herds. the number of the animals in the enclosure increased from 7 in 1994 to 39 in may 2001. the breeding herd in the enclosure then consisted of: 1 adult bull; 7 adult cows; 4 young animals 20–30 months old; 5 calves of 10–12 months; and 8 calves of 6 weeks; the other 14 animals did not form part of the breeding herd. the animals in the enclosure showed a steady increase, in contrast to the free-ranging population. there did not seem to be a link between the dry season rainfall (july to september) of each year and the population growth or the number of calves per cow produced in the enclosure or outside as suggested by owen-smith & ogutu (in press) (fig. 12). fig. 12. calves produced per cow in the free roan antelope herds as well as in the n'washitshumbe enclosure, compared to the dry season rainfall. fig. 13. lion numbers on the northern basalt plains. grant.qxd 2005/12/09 10:18 page 59 overlap in habitat selection of roan antelope and zebra the mean home range of nine colourcollared zebra calculated from 487 sightings, between november 1993 and may 1995, was 267 ± 103 km². this is much larger than the home range calculated for the four radio-collared roan antelope (60 ± 4.5 km²). zebra home ranges thus include more waterpoints than those of the roan antelope, which restrict their movements to one or two waterpoints. of the 234 sightings of zebra during radio tracking sessions of roan antelope during 1996, 165 were in the vicinity of waterpoints. of the 146 sightings of zebra grazing 85.8 % were grazing in the more open, lowlying areas rather than on the denser crests (13.2 % of encounters). radio-tracked roan antelope were an average of 2.19 km from koedoe 45/2 (2002) 60 issn 0075-6458 table 6 difference in vegetation in the areas north of the olifants river in 1999 correcting for the vegetation status of 1994 biomass % decreasers geographical zone kg/ha sd n % sd n far northern granites 1898 ab 163 41 14.4 a 2.5 35 northern basalt plains 2970 a 148 50 22.3 a 2.2 47 northern granites 772 ab 157 44 7.9 a 2.3 41 mooiplaas basalts 1167 abc 142 54 12.7 a 2.0 52 olifants transition zone 735 abc 155 45 10.5 a 2.2 6.1 letters in superscript and upper case indicate a significant difference with the same letter in superscript and lower case. thus valuea differs significantly form valuea in the same column. (biomass (p = <0.0001); percentage decreaser species (p <0.0001). table 7 vegetation parameters in the closed and open area of the nbp compared to the mooiplaas basalts geographical biomass % % zone kg/ha decreasers increasers closed 5261 34 69 open 4777 31 64 mooiplaas basalts 5531 22 75 water (55 sightings), also showing a significant selection for grazing in the low-lying areas (p < 0.0001) (table 5). vegetation responses with the increase in rainfall after 1993 there was a general increase in biomass and in percentage decreaser species (fig.7). the closure of waterpoints on the nbp was associated with a significantly higher biomass and percentage of palatable, decreaser species on the nbp than the other areas north of the olifants river (table 6). the biomass, however, did not differ significantly (p = 0.22) between the closed and open areas and the adjacent mooiplaas basalts. the percentage decreasers were significantly higher in the closed area (p = 0.041) than in the mooiplaas basalts (table 7). the detailed vegetation studies on the effect of closure of waterpoints indicated that the area around closed waterpoints changed from annual to perennial grasslands. these areas had a high abundance of palatable, perennial, decreaser species. the grassland around open waterpoints was still dominated by annuals, of which most were unpalatable increaser species (jacoby 1999). decreaser species such as themeda triandra and digitaria eriantha were also present at closed, and not open, waterpoints (davidson 1996). grant.qxd 2005/12/09 10:18 page 60 issn 0075-6458 61 koedoe 45/2 (2002) table 8 tuft and grazing height of grass species selected by roan from available species in 1996 grass species average average % of % of leaf grazing available selected height cm height cm tufts tufts cenchrus ciliaris 71.3 54.6 9.63 21.99 themeda triandra 73.9 53.2 5.96 21.23 panicum coloratum 77.2 32.6 41.06 20.46 panicum maximum 88.3 42.9 2.52 16.11 eragrostis superba 55.8 40.5 2.29 6.65 urochloa mosambicensis 53.4 14.4 9.63 5.12 heteropogon contortus 49.8 30.4 2.75 4.86 setaria incrassata 79.2 31 1.15 1.53 schmidtia poppophoroides 34 34.8 12.39 1.02 enneapogon cenchroides 25.4 10.5 3.67 0.51 digitaria eriantha 56.7 25 1.38 0.26 table 9 numbers of lions in different sex and age classes on the nbp before and after closure of waterpoints october 1993 december 1996 adult male 33 5 adult female 36 24 subadult male 6 10 subadult female 7 7 cub 14 5 the number of mopane trees, with branches broken by elephant loxodonta africana, decreased with increasing distance from open waterpoints, to a level of damage that was the same as around closed waterpoints at 1600 m. other species that were browsed by elephant around waterpoints, such as dichrostachys cinerea, lonchocarpus capassa, and combretum imberbe were utilised up to 100 m, 200 m and 800 m, respectively, from the waterpoints (fruhauf 1997) . the roan antelope selected different grass species during the 1996 study, compared to the species selected in 1976 (joubert 1976). themeda triandra was the most preferred species of roan antelope in 1976, even though panicum coloratum was the most abundant of the selected grass species. in 1996 panicum coloratum was by far the most abundant of the species that the roan antelope utilised, but themeda triandra and cenchrus ciliaris, that were far less abundant than during the 1976 study made up an equal proportion of their diet (table 8). according to 480 observations of tufts utilised, roan antelope selected to feed at an average height of 41 cm, which was below the average leaf height of 69.3 cm of the decreaser species, indicating that the grass height was suitable for roan to select the preferred parts. change in the lion population results of four lion-calling censuses suggested an initial decrease in 1995 after the closure of waterpoints. population numbers stayed more or less constant towards 2002 (fig. 13). after the closure of waterpoints, the mean pride size on the nbp of 6.4 (n = 39) was significantly larger than the 3.54 (n = 87) recorded when the waterpoints were still open (p = 0.0006). this was associated with changes in pride composition with a larger number of subadult males (table 9). grant.qxd 2005/12/09 10:18 page 61 very few incidences of predation of rare antelope were recorded on the nbp, in spite of intense observation. there was only one record of a tsessebe calf caught by lions in 1996 and an adult roan antelope cow probably caught by a lion in april 2001. discussion the rainfall during the dry cycle of the 1980s and early 1990s was the lowest in recorded history in 10 out of the 22 rainfall measuring stations in the knp, and followed on a very wet cycle in the 1970s (zambatis & biggs 1995). vegetation monitoring only started in 1989, but by using remote sensing techniques (botha 1997) found that the area of degraded or bare veld had increased from 6 % of the northern plains area in 1985 to 40 % in 1989. during the same period the area of healthy vegetation (by his criteria) declined from 10 % in 1985, to 3 % in 1989. the vegetation monitoring after 1989 reflected the effect of the dry cycle, with the decline of palatable, decreaser species being one of the main consequences. the vegetation surveys in 1999 indicated that the percentage decreaser grass species have not increased to the same point as in 1989 and can be presumed to have been much lower than in 1985. the changes observed in animal numbers are most likely the result of the change in rainfall. animal biomass would have been high at the end the very wet cycle of the 1970s (coe et al. 1976) when animal counts were started, with mesic species such as tsessebe, sable and roan responding favourably to the high rainfall (pienaar 1974; mills et al. 1995b). with the onset of the dry cycle, buffalo and rare antelope numbers could be expected to decline, with zebra and wildebeest increasing (mills et al. 1995b). the initial drop in roan numbers was thus not unexpected as they are one of the species most sensitive to habitat changes (smithers 1983), and to competition from other species (wilson & hirst 1977). the presence of artificial waterpoints would have further favoured water-dependent species such as zebra above the rare antelope species during a period of severe drought (owen-smith 1996). with the increase in rainfall after 1994 zebra and wildebeest numbers would again be expected to drop, as was indeed observed, with a slow rise in the numbers of the mesic species of which only the buffalo numbers showed some evidence (fig. 5). smuts (1972) commented that in spite of the fact that zebra had not over-utilised the northern plains, their large numbers led to competition with the rare antelope species such as roan. he stated that zebra numbers would have to be checked especially if the eastern boundary was to be fenced. the vegetation index maps created by botha (1997) for 1989 show that the drainage lines (and densely wooded sandveld, which is not readily utilised by zebra or roan antelope) were the only areas with readily available biomass in 1989. observations of radio-collared roan in 1996 show that roan and zebra preferably foraged in the low-lying drainage areas. one could thus assume that both species would have been utilising these areas during the drier periods and that roan antelope as selective feeders would have been the more disadvantaged (wilson & hirst 1977). the decision to close waterpoints coincided with an increase in rainfall from 1994 onwards. this increase in rainfall and the expected resultant decline in zebra numbers in the area (mills et al. 1995b), resulted in an improved field layer around the windmills as well as in the general area, seen as an increase in biomass (standing crop) and palatable decreaser species in the area of closed windmills. in spite of the improvement of the vegetation after the closure of waterpoints and the increase in rainfall, even dry season rainfall, the habitat had not recovered totally. this is reflected by the fact that even though roan were able to feed on nutritious species such as panicum coloratum they were not able to select the same percentage of themeda triandra as in the 1976 study (joubert 1976). at that time this was the most preferred koedoe 45/2 (2002) 62 issn 0075-6458 grant.qxd 2005/12/09 10:18 page 62 species, and was identified as good indicator of where to find roan. rather than the expected slow increase of rare antelope in response to the improvement in vegetation, roan stabilised but did not increase, while sable and tsessebe numbers declined further after the closure of waterpoints. sable antelope that were scattered along the drainage lines on the nbp in 1993 and 1996, had all but disappeared from the nbp in 1998, while the remaining larger herds of tsessebe were more strongly associated with waterpoints. although the general decline of tsessebe was found to be related to the decline in rainfall (dunham & robertson 2001), they seemed to have reacted negatively to the closure of waterpoints, showing a substantial decline from 70 on the plains in 1993 to 40 in 1998, and 20 in 2000. eckhardt (2000 pers. comm.) also found evidence of a large increase in the woody vegetation cover on the basalts and granites of northern knp over five years, which would have made the habitat less suitable for tsessebe (garstang 1982). unfortunately, the response of the rare antelope to the increase in rainfall in the rest of the knp does not lend itself to examination, as the approach to the collection of rare antelope numbers was being adapted between 1995 and 1999. in a survey during may 1999, rangers in the knp estimated the sable population at 507, the tsessebe population at 158, and the roan population (including 33 animals in the enclosure) at 63. these numbers are all much lower than reported in 1994 and the closure of waterpoints may not be the only reason for the decline in sable and tsessebe. one can only speculate on the effect of the boundary fence (since its erection in 1976) in further limiting movement of roan to the southern limit of their distribution. the roan numbers started dropping 10 years after the effective closure of the area as the rainfall dropped below the mean. the restricted chance for dispersal of the dambo-loving roan to other suitable habitats was first associated with a build-up of numbers during a wet period, and then with a severe drop in the roan antelope population during the drought. genetic links with the eastern and northern population, rather than the western population (mathee & robinson 1999) seem to indicate that the roan in the knp must have had historical links with these eastern populations. this indicates likely movement between the adjacent areas of mozambique and zimbabwe before the presence of fences. even though severe losses of roan were reported during the anthrax epidemics, the protective effect of vaccination on the population is difficult to evaluate, as there had not been similar outbreaks in the area during the vaccination campaign or thereafter. it does not seem that the vaccination procedure had a negative affect on the roan antelope, as the only remaining roan herds are still on the nbp, where they were vaccinated. densities on the nbp remained higher than in the mooiplaas basalts where there is also suitable habitat and where they were not vaccinated. the possible role of predation, especially on the small remaining population, cannot be ignored. in the enclosure where the roan antelope were safe from predators, the population grew from 7 to 49 in 7 years, while the free-living populations hardly increased during the same period. roan had been reported to fall prey to lions even during the previous wet cycle, and over 22 years, rangers reported 90 roan killed by lion which formed 1.11 % of the lion diet. from these reports it was estimated that 4–12 % of adult roan fall prey to lions every year. harrington et al. (1999) developed a model that explained the initial steep decline in the roan population by predation. in a subsequent study, mcloughlin (2001) found that at predation rates of 15 % the viability of the current small freeranging population over the next 50 years is low. issn 0075-6458 63 koedoe 45/2 (2002) grant.qxd 2005/12/09 10:18 page 63 conclusion over the span of several decades, the equilibrium-centred "command and control" approach aimed at the conservation of elements such as species, of which the rare antelope was of specific concern, seems to have been very successful. however, the increased availability of perennial water and limitation of movement to other habitats removed important sources of variation. thus the system may not have been able to adapt to the severe and prolonged drought, as its ability to do so had been reduced resulting in a reduction in system variation (holling & gunderson 2002). the new management approach involving closure of waterpoints, the opening of boundary fences as well as the readjustment of the fire management to a less controlled regime, should again increase the ecosystem resilience (holling & gunderson 2002). however, the time scale for this resilience to take effect is probably several decades and possible negative short-term results have to be considered carefully. this study also illustrates the difficulties involved in conserving low numbers of a species at the limit of their distribution. the habitats of these rare antelope in the knp may be marginalised even further, as global climate models mostly predict drier and warmer conditions in the decades ahead, with an associated increase in woody biomass (rutherford et al. 1999). in "nature evolving" management approaches emphasising ecosystem resilience, an adaptive management approach needs to be adopted (rogers et al. 2000; holling & gunderson 2002). parameters to indicate thresholds of potential concern (tpcs) were developed for herbivores and vegetation and were incorporated in the new knp management plan (braack 1997). these limits were implemented to elicit further management responses to the persistent small roan antelope population and the declining tsessebe population and led to the decision to expand the captive breeding facilities (grant & van der walt 2000). the challenge for rare antelope conservation will be to decide how to balance the need for resilient ecosystems with the specialised requirements of marginal species. acknowledgements nick zambatis, wikus van der walt, ian whyte, judith kruger and steven whitfield are acknowledged for providing or collecting data used in this publication and dewald keet is thanked for providing the lion data for the last two censuses. the kruger national park and the kruger park marathon club is thanked for financial and logistic support and assistance. j. t. du toit, gus mills, n. owen-smith and harry biggs are thanked for their advice on the preparation of the manuscript, and reinette biggs is thanked for her assistance with the analysis of the data. references anon. 1961. biologist year report 1960-1961. scientific services, skukuza, kruger national park. anon. 1962. nature conservation year report 19611962. scientific services, skukuza, kruger national park botha, j.h. 1997. remote change detection of vegetation as a method for investigating the declining herbivore populations in the northern plains, kruger national park, south africa. scientific report, scientific services, skukuza, kruger national park braack, l.e.o. 1997. an objectives hierarchy for the management of the knp. a review of parts of the management plan for the knp. vol. vii. skukuza: kruger national park. buckland, s.t., d.r. anderson, k.p. burnham & j.l. laake. 1993. distance sampling. estimating abundance of biological populations. london: chapman. child, g., h. robbell & c.p. hepburn. 1972. observations on the biology of tsessebe, (damaliscus lunatus lunatus) in northern botswana. mammalia 36: 131–141. coe, m.j., d.h. cumming & j. phillipson. 1976. biomass and production of large african herbivores in relation to rainfall and primary production. oecologia 22: 341–354. davidson, t. 1996. recovery of the herbaceous vegetation around three closed waterhols on the northern plains of the kruger national park. honours thesis, university of the witwatersrand, johannesburg. davidson, t. 2002. response of the zebra population on the northern plains to resource distribukoedoe 45/2 (2002) 64 issn 0075-6458 grant.qxd 2005/12/09 10:18 page 64 tion in the landscape. msc. thesis, university of the witwatersrand, johannesburg. de vos, v., g.l. van rooyen & j.j. kloppers. 1973. anthrax immunization of free-ranging roan antelope hippotragus equinus in the kruger national park. koedoe 16: 11–25. dunham, k.m. & e.f. robertson. 2001. trends in rainfall explain the rise and fall of tsessebe in the kruger national park. internal scientific report 01/01, scientific services, skukuza, kruger national park. fruhauf, n. 1997. pattern of elephant impacts around artificial water points in the northern region of the kruger national park. hons. thesis, university of the witwatersrand, johannesburg. garstang, r. 1982. an analysis of home range utilization by the tsessebe damaliscus lunatus lunatus (burchell), in the p.w. willis private nature reserve. msc. thesis, university of pretoria, pretoria. gaza-kruger-gonarezhou transboundary natural resources management initiative. 2002. great limpoppo transfrontier park. joint management plan draft january 2002. gertenbach, w. 1983. landscapes of the kruger national park. koedoe 26: 9–121. grant, c.c. & j.l. van der walt. 2000. towards an adaptive management approach for the conservation of rare antelope in the kruger national park – outcome of a workshop held in may 2000. koedoe 43: 103–112. gureja, n. 2001. the comparative use of burnt areaas by four species of grazing antelope in a northern province game farm, south africa. msc thesis, university of the witwatersrand: johannesburg. harrington, r., r.n. owen-smith, p.c. viljoen, d.r. mason & p.j. funston. 1999. establishing the causes of the roan antelope decline in the kruger national park, south africa. biological conservation 90: 69–78. harrington, r. & d.j. pienaar. 1994. report on the roan antelope workshop skukuza 26 july 1994. internal report, scientific services, skukuza, kruger national park holling, c.s. & l.h. gunderson. 2002. 2. resilience and adapative cycles. pp. 25–62. in: gunderson, l.h. & c.s. holling (eds). panarchy. understanding transformations in human and natural systems. washington: island press. holling, c.s. & g.k. meffe. 1996. command and control pathology of natural resource management. conservation biology 10: 328–337. huntley, b.j. 1982. southern african savannas. pp. 101–119. in: huntley, b.j. & b.h. walker (eds). ecology of tropical savannas. heidelberg, n.y: springer verlag berlin. jacoby, d.l. 1999. responses of herbaceous vegetation to reduced disturbance on the northern plains of the kruger national park. msc thesis, university of the witwatersrand, johannesburg. joubert, s.c.j. 1970. a study of the social behaviour of the roan antelope, hippotragus equinus equinus (desmarest, 1804), in the kruger national park. m.sc. thesis, university of pretoria, pretoria. joubert, s.c.j. 1972. territorial behaviour of the tsessebe (damaliscus lunatus lunatus burchell) in the kruger national park. zoologica africana 7: 141–156. joubert, s.c.j. 1976. the population ecology of the roan antelope (hippotragus equinus equinus) (desmarest, 1804) in the kruger national park. dsc thesis, university of pretoria, pretoria. joubert, s.c.j. 1985. masterplan for the management of the kruger national park. skukuza: kruger national park. internal report, scientific services, skukuza, kruger national park joubert, s.c.j. 1986. the kruger national park an introduction. koedoe 29: 1–11. joubert, s.c.j. & p.j.l. bronkhorst. 1977. some aspects of the history and population ecology of the tsessebe damaliscus lunatus lunatus in the kruger national park. koedoe 20: 125–145. mason, d.r. 1990. monitoring age and sex ratios in ungulate populations of the kruger national park by ground survey. koedoe 33: 19–28. mathee, c.a. & t.j. robinson. 1999. mitochondrial dna population structure of roan and sable antelope: implications for the traanslocation and conservation of the species. molecular ecology 8: 227–238. mcloughlin, c.a. 2001. population viability of rare antelope in the kruger national park. msc thesis, university of the witwatersrand. johannesburg mills, m.g.l., h.c. biggs, p.j. funston, j.l. van der walt, p.c. viljoen & a.. viljoen. 1995. annual report on the northern plains project june 1995. scientific services, skukuza, kruger national park. mills, m.g.l., h.c. biggs, & i.j. whyte. 1995. the relationship between rainfall, lion predation and population trends in african herbivores. wildlife research 22: 75–88. owen-smith, r.n. 1996. ecological guidelines for waterpoints in extensive protected areas. south african journal of wildlife research 26: 107–112. owen-smith, r.n. & j. ogutu (in press). rainfall influences on ungulate population dynamics in the kruger national park. in: du toit, j.t., k.h. rogers & h.c. biggs (eds). the kruger experience: ecology and management of savanna heterogeneity. island press. pienaar, u. de v. 1963. the large mammals of the kruger national park – their distribution and present-day status. koedoe 6: 1–37. issn 0075-6458 65 koedoe 45/2 (2002) grant.qxd 2005/12/09 10:18 page 65 pienaar, u. de v. 1974. habitat-preference in south african antelope species and its significance in natural and artificial distribution patterns. koedoe 17: 185–195. pienaar, u. de v.. 1985. indication of progressive dessication of the transvaal lowveld over the past 100 years, and implications for the water stabilization programme in the kruger national park. koedoe 28: 93–165. rogers, k., d. roux & h.c. biggs. 2000. challenges for catchment management agencies: lessons from bureaucracies, business and resource management. water sa 26: 505–511. rutherford, m.c., g.f. midgeley, w.j. bond, l.w. powrie, r. roberts & j. allsop. 2000. plant biodiversity. in: kiker, g. (ed.). climate change impacts in southern africa. report to the national climate change committee, department of environmental affairs and tourism, pretoria, south africa. smithers, r.h.n. 1983. the mammals of the southern african subregion. pretoria: university of pretoria. smuts, g.l. 1972. seasonal movements, migration and age determination of burchell’s zebra (equus burchelli antiquorum, h. smith, 1841) in the kruger national park. m.sc. thesis, university of pretoria, pretoria. trollope, w.s.w. 1990. development of a technique for assesing veld condition in the kruger national park. journal of the grassland society southern africa 7: 46–51. trollope, w.s.w., a.l.f. potgieter & n. zambatis.. 1989. assessing veld condition in the kruger national park using key grass species. koedoe 32: 67–93. van wilgen, b.w., h.c. biggs, s.p. o’regan & n. maré. 2000. a fire history of the savanna ecosystems in the kruger national park, south africa, between 1941 and 1996. south african journal of science 96: 167–178. venter, f.j. 1990. a classification of land use for management planning in the kruger national park. washington: island press. viljoen, p.c. 1996. ecological aerial surveys in the kruger national park. 1995 no 4/95. scientific services, skukuza, kruger national park. wilson, d.e. & s.m. hirst. 1977. ecology and factors limiting roan and sable antelope populations in south africa. wildlife monographs 54: 1–109. zambatis, n. & h.c. biggs. 1995. rainfall and temperatures during the 1991/92 drought in the kruger national park. koedoe 38: 1–16. koedoe 45/2 (2002) 66 issn 0075-6458 grant.qxd 2005/12/09 10:18 page 66 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice reilly audit.qxd auditing wildlife b.k. reilly and y. reilly reilly b.k. and y. reilly. 2003. auditing wildlife. koedoe 46(2): 97–102. pretoria. issn 0075-6458. accountants and auditors are increasingly confronted with the problem of auditing wildlife populations on game ranches as their clients' asset base expands into this industry. this paper aims to provide guidelines on these actions based on case study data and research in the field of wildlife monitoring. parties entering into dispute on numbers of animals on a property often resort to their auditors for advice. this paper tracks a method of deciding on whether or not to audit the population based on wildlife value and an initial sample count. this will act as a guideline for the accounting profession when confronted by this problem. keywords: game counting, game ranches, decision support, operational auditing, helicopter counts. b.k.reilly , department of nature conservation, technikon pretoria, private bag x680, pretoria, 0001, south africa; y. reilly, department of auditing, university of south africa, p.o.box 392, pretoria, 0001 republic of south africa. issn 0075-6458 97 koedoe 46/2 (2003) introduction the current number of game ranches in south africa is estimated at 8000 (eloff 2000). business people who are in part time participants in wildlife own a significant number of these or nature based undertakings. many of these operations do not have ecological management plans with clearly defined goals and the necessary monitoring actions in place to track their progress. from time to time, on sale of property, expansion, and consolidation or for ad hoc management, owners and their appointed managers resort to once-off game counts. even more disturbing is the fact that many disreputable operators in the wildlife industry are only too willing to oblige and provide poor data resulting in poor management and strategic decisionmaking. the position is often exacerbated by poor advice from neighbours, relatives and government extension officers, both from conservation and agricultural agencies. internal auditing all of the above accepted scientific/biological definitions that are essentially within the gambit of what is known as an operational audit. the operational audit is used to judge the quality of operations and make operational improvements in any undertaking (reider 1994). because of today's multifaceted and multi-disciplined economic environment, management began to place increasing emphasis on evaluation of the organization's operations with regards to economy, efficiency, effectiveness and impact on the environment. operational auditing can thus be seen as a service to management as well as improving organizational performance because it involves a systematic review of an entities activities in relation to specified objectives. performance appraisal involves a comparison of the way in which an entity performs its activities with: objectives established by management or any other party asked to do so; other appropriate measurement criteria. operational auditing as a subset of internal auditing, operational auditing reviews undertakings' activities for efficiency and effectiveness (table 1) and reilly audit.qxd 2005/12/09 11:22 page 97 may evaluate any type of activity at any level within the organization. the overall objective is maximizing organizational welfare. efficiency may be viewed in this definition as an input measure that relates to cost control and is concerned with the performance of recurring functions at a minimum of cost to the undertaking. effectiveness is output orientated; it is viewed as a measure of productivity in utilizing the undertaking's resources and in terms of long term profitability. one might also view this approach as riskbased operational auditing because of its focusing on helping management to improve the way the undertakings activities are performed. performance objectives are very important— if they are non-existent, direction, measurement, evaluation and management of performance cannot be achieved. these objectives provide direction as to what management is trying to achieve, a means of measuring the level of achievement as well as benchmarks against which performance can be evaluated. this performance evaluation provides the trigger to begin actions to correct performance problems or to improving performance by raising standards. according to reider (1994) operational auditing has many benefits and depending on its scope it can be of significant benefit to management in some or all of the following ways: identifying problem areas, related causes koedoe 46/2 (2003) 98 issn 0075-6458 table 1 the relationship between economy, efficiency and effectiveness input activity output economy efficiency effectiveness resources objectives doing things right doing the right thing and alternatives for improvement; locating opportunities for eliminating waste and inefficiency thence cost reduction; locating opportunities to increase revenues—improvement of income; identifying undefined organizational goals, objectives, policies and procedures; identifying criteria for measuring the achievement of organizational goals and objectives; recommending improvement in policies, procedure and organizational structure; providing checks on performance by individuals and organizational units; reviewing compliance with legal requirements and organizational goals objectives and procedures; testing for existence of unauthorized fraudulent or otherwise irregular acts; assessing management information and central systems; identifying possible trouble spots in future operations; providing an additional channel for communication between operating levels and top management; providing an independent objective evaluation of operations. what, you may wonder does this have to do with gamecounting? currently, monitoring of wildlife populations is central to decision making surrounding a resource that provides the backbone of the ecotourism industry. where wildlife population monitoring still takes place, budgets are shrinking and managers responsible are failing to meet all the requirements of monitoring, biological, statistical and financial. there is a growing perception within the financial management component of organizations that valuable wildlife assets need to be better accounted for. wildlife on state land is not being adequately accounted for and an operational audit approach may be an avenue for adequate accounting under internal audit responsibility instead of that of the conservation biologist. internal audit components of government may be the area of responsibility required to adequately fund and conreilly audit.qxd 2005/12/09 11:22 page 98 trol wildlife resources. the time of external operational audit of government conservation agencies and statutory bodies may have arrived with regard to the wildlife assets of which they are custodians. this in the light of increasing risk to wildlife populations from various quarters. an underlying principle of auditing is, however, that sampling procedures answer to rigorous statistical requirements in particular error probabilities (reider 1994). authors in wildlife management concur that estimates of precision are essential outputs of gamecounts. the relationship between operational auditing and wildlife management in statistical terms is laid out in table 2. collinson (1985) observed that there were limitations with helicopter-based surveys, which form the backbone of gamecounting in southern africa. the results of reilly (2000) directly support collinson's (1985) contention that the method has severe limitations where repeatable estimates over time are required unless more than four counts are contemplated. reilly (2000) draws the following conclusions on helicopter total counts in his study: the method is incapable of consistently detecting small population (10 20%) changes at acceptable power and significance under moderate replication (3-3) or high significance (talpha.1); the precision and power attained in this limited study is entirely site specific. it is difficult to make any general statements about either; the assumption of constant variance from t1 to t2 cannot be validated; operating time for the method should be limited with operations over 2,5 hours being corrected for decline in observations over time; cognizance should be taken of the differing observation profiles of observers in the initial choice of observers for gamecounts. failures of techniques, be they helicopter, fixed-wing or ground-based methods, combined with declining financial resources have forced monitoring programmes back to a point of serious review by both management and biologists. the increasing value of wildlife, on the other hand, combined with its pivotal role in the hunting and ecotourism industry, necessitates a new approach to justifying costing and implementing such programmes. clearly the value of wildlife has to enter into determination of monitoring efficiency, effectiveness and budgeted cost. the proposed audit hypothesis model forms the basis of decision making in whether or not the count should take place. this is based on a sample which can be extracted from previous counts. the operational audit route statistical sampling the internal auditor can assist in two ways, viz., determination of monetary value of the unit to be audited, and consideration of the statistical sampling technique. normally wildlife managers select the appropriate counting technique for monitoring wildlife and very often ignore the statistical considerations. biologists generally take a different view of statistical sampling to that of managers. reilly (2000) and thompson et al. (1998) point to the fact that wildlife managers and some biologists ignore the rules of inference in determining large ungulate numbers. it appears that the audit profession concurs and the internal auditors have found the use of certain statistical sampling techniques advantageous in certain instances. the use of statistical sampling does not reduce the use of judgement by the auditor but provides certain statistical measurements as to the results of audit tests, which measurements may not otherwise be available. when an auditor is faced with a situation requiring the projection of a total quantity or total rand value, based on a mean of a sample, accounting estimation is the appropriate tool to use (guy et al. 1998). by using estimation sampling, a projection can be made of the estimated population value that becomes the booked amount. the estimated population value plus and minus calculated precision is expected to contain the true but unknown value at a defined reliability level. issn 0075-6458 99 koedoe 46/2 (2003) reilly audit.qxd 2005/12/09 11:22 page 99 the audit hypothesis model in statistics the risk of incorrect rejection is called the alpha risk and the risk of incorrect acceptance is called the beta risk. the relationship between audit terms and statistical terms are as follows: alpha risk is the chance that the statistical evidence might fail to support a materially correct book value. this type of error usually results in testing additional sample items. decreasing or increasing reliability controls this risk. for reliability in auditing read precision in wildlife monitoring. in wildlife management terms a decision based on population change shown in a sample that did not occur in reality. beta risk is the probability that the statistical evidence might support a lack of material. beta risk is controlled by adjusting the ratio of precision a to materiality. again in wildlife terms increasing precision reduces this risk and in management terms this is a scenario of no change shown by the sample when in reality change has occurred. audit hypotheses testing when the auditor's primary concern is controlling beta risk, effectiveness—(output orientated) thus utilizing the resources in terms of long-run profitability—as discussed under operational auditing, he will use this method: in the previous method discussed beta risk was not considered because we were not trying to decide if a book value was materially misstated. if an auditor rejects the recorded inventory value the client will generally insist on an increase in sample size or a search for additional evidence to show that the inventory balance is misstated. in this method the audit hypothesis model differs from the previous method as follows: the calculated desired beta risk must be calculated taking into consideration audit risk, internal control and other audit procedures results. once precision is determined, the substantive test sample size is then calculated. after the sample items selected are audited, the auditor ascertains whether the statistical evidence supports material correctness of the client's book value. the audit hypothesis model (ahm) is ideally suited to wildlife management undertakings. it consists of a 22 step approach, beginning with assessment of internal control and ending with an accept or reject decision as to material misstatement of a balance—in this case the number of animals, grasses or trees, i.e., the wildlife asset. the model will be explained by way of an example that commences with table 3. from the operational audit standpoint, this reserve has a stock of 1289 items of varying value. a counting (audit) programme is in place. based on (1) tests of control of the inventory count (2) observation that the gamecount instructions are followed, and (3) the evaluation of the competence of personnel undertaking the count, it is decided that the control risk (cr) is no more than 20%. management decides that the inventory valuation of r 9 755 805 is acceptable if actual inventory is within 5% (95% confident) of this value. the statistical test will be based on recounting, repricing and extending each item in the sample (part of a count). other audit procedures (ap) consist of inventory turnover calculations (population dynamics), comparison to previous years and cutoff tests to ensure that purchases and sales are reflected in the inventory for the period. at a guess these procedures are estimated at 80% risk. substantive test planning now begins with the calculation of sample size using the following equation: n' = (ur.sd.n / a)2 ur is already defined as 1,96 (95% confidence), sd is calculated from the value of the sample, n is the 1289 items and a needs to be calculated. firstly beta (type ii error) risk is calculated based on audit risk of 5% (95% confidence) assessed control koedoe 46/2 (2003) 100 issn 0075-6458 table 2 relationship between statistical terms, operational auditing and wildlife monitoring statistical terms audit terms gamecount risk of incorrect rejection alpha risk audit efficiency significance/risk of type i error risk of incorrect acceptance beta risk audit effectiveness 1 power/risk of type ii error reilly audit.qxd 2005/12/09 11:22 page 100 risk (cr) of 20% and other procedural risk (ap) of 80%, beta is: beta(planned) = (0.05 / 0.20 x 0.80) = 0.3125 the beta risk coefficient is 0,5–0,3125 which is 0,1875. using a normal area curve table we see that this corresponds to a beta risk coefficient (zbeta) of 0,49. beta is controlled by varying the precision of a in relation to tolerable misstatement (tm) (rand value ± 5 %). a = tm.(ur / ur + zbeta) = r390232 this is acceptable precision as it falls inside the rand value ± 5%. the sample size calculation is now: n' = (ur.sd.n / a)2 = 4679 this is the sample size with replacement, the samples size without replacement is: n' = (4679 / 1 + (4679/1289) = 1011 at this point if the sample appears representative the sample is audited. a' is now calculated for which the sd of the sample (table 3) and the sample is required: a' = (ur.se.n √ 1 n/n) = r2158694 if a' is not equal to a then a'' must be calculated: a'' = (a' + tm(1 a'/a) = r47434 estimated audited value (eav) is the mean of the sample value times n or r5 306 555.20. a decision interval of book value (r9 755 805) ± a'' or r9 755 805 ± r47 434. clearly the eav falls outside this decision interval, therefore the statistical evidence does not support material correctness. the implications are that the entire stock should be audited. the effectiveness of a count or audit can be measured using statistical power as an index. reilly & haskins (1999) have amply demonstrated this for counts on suikerbosrand nature reserve. the issue of efficiency, viz., the ratio of output to input can also be illustrated using statistical power. the relationship between power per number of replicates calculated against the cost of those replicates allows decision making on optimizing in this area. table 4 shows this relationship. it is evident from table 4 that as replicates increase efficiency improves—it is cheaper to count more often. finally, how much to spend. the audit profession has no fixed guidelines in this regard, relying on the objective evaluation of the auditor. with what we have seen so far, the auditor will definitely decide on a full audit. in the absence of fixed formulae, the authors propose that 1 % of the value of the stock be allocated to the audit. issn 0075-6458 101 koedoe 46/2 (2003) table 3 mean number of animals counted from 4 replicate counts on loskop dam nature reserve (1991), mean rand value (eloff, 2000), total value and values from a random sample count of 1 hour species mean no. mean value (2000) total value no in sample sample value kudu 259 r2 105.00 r544 142.50 3 r6 315.00 zebra 239 r2 640.00 r630 300.00 8 r21 120.00 wildebeest 186 r2 226.00 r414 036.00 26 r57 876.00 impala 317 r596.00 r188 783.00 52 r30 992.00 warthog 39 r567.00 r21 971.25 11 r6 237.00 sable 41 r53 580.00 r2 210 175.00 r0.00 white rhino 30 r176 801.00 r5 304 030.00 2 r353 602.00 mountain reedbuck 156 r993.00 r154 659.75 18 r17 874.00 giraffe 23 r12 509.00 r287 707.00 r0.00 total 1289 r252 017 r9 755 805 120 r494 016 standard deviation (sd) r58 360 r1 712 094 r113 508 standard error (se) r1 836.34 reilly audit.qxd 2005/12/09 11:22 page 101 summary and conclusions in this paper we have seen that it is possible to provide concrete decision support in terms of the key questions that surround the ubiquitous gamecount. these questions are becoming more relevant as conservation agencies, in particular, struggle for funding. considering the custodianship entrusted to these organizations and the monetary value of the wildlife coupled with the fact that the biologists have to perpetually seek funding to undertake the gamecounts, a shift of responsibility in this regard to an internal audit component is indicated. with a random sample from a previous count, a known book value of each ungulate, published mean power values and a little bit of borrowed technology a desktop exercise provides answers to all the posed questions. the time has come for the competent, realistically funded audit of wildlife with commensurate audit responsibilities while the biologists get on with the field techniques. references collinson, r.h.f. 1985. selecting wildlife census techniques. monograph 6, institute of natural resources, university of natal, pietermaritzburg. eloff, t. 2000. gemiddelde wildveilingpryse vir suid afrika, 2000. unpublished report, university of potchefstroom. goldsmith, f.b. 1991. monitoring for conservation and ecology. london: chapman. guy, d.m., d.r. carmichael & o.r. whittington. 1998. audit sampling. an introduction. new york. john wiley. reider, h.r. 1994. the complete guide to operational auditing. new york: john wiley. reilly, b.k. & c.i. haskins. 1999. comparative efficiency of two game counting techniques as applied to suikerbosrand nature reserve. south african journal of wildlife research 29(3): 89 97. reilly, b.k. 2000. the statistics of helicopter total counts of large ungulates in sourish mixed bushveld, northwest arid bushveld and mopane veld, republic of south africa. unpublished thesis, university of stellenbosch. thompson, w.l., g.c. white & c. gowan. 1998. monitoring vertebrate populations. san diego: academic press. koedoe 46/2 (2003) 102 issn 0075-6458 table 4 cost per percent statistical power (1-beta) to show 15% population change at 10% significance for populations (stock) on loskop dam nature reserve (power analysis based on actual counted replicates) using helicopter based surveys at current prices species kudu zebra wildebeest impala warthog sable warthog mt reedbuck r832.76 r1 097.73 r1 271.05 r1 420.59 r1 610.00 r1 271.05 r1 150.00 r3 450.00 r402.50 r589.02 r690.00 r805.00 r1 006.25 r710.29 r619.23 r2 683.33 r305.70 r431.25 r525.00 r603.75 r754.69 r525.00 r464.42 r2 415.00 r473.53 r710.29 r862.50 r928.85 r1 150.00 r832.76 r731.82 r3 018.75 r271.35 r355.15 r423.68 r473.53 r603.75 r409.32 r371.54 r2 195.45 r383.33 r575.00 r690.00 r779.03 r966.00 r670.83 r603.75 r2 683.33 r280.81 r383.33 r455.66 r503.13 r652.70 r439.09 r395.90 r2 415.00 r256.91 r317.76 r377.34 r416.38 r525.00 r360.45 r330.82 r2 012.50 r248.97 r290.96 r330.82 r365.91 r464.42 r322.00 r298.15 r1 857.69 reilly audit.qxd 2005/12/09 11:22 page 102 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice robinson.qxd distribution and status of marine invasive species in and bordering the west coast national park t.b. robinson, c.l. griffiths and n. kruger robinson, t.b., c.l. griffiths and n. kruger. 2004. distribution and status of marine invasive species in and bordering the west coast national park. koedoe 47(1): 79–87. pretoria. issn 0075-6458. this study describes the present distribution and abundance of alien marine species within the west coast national park. the mediterranean mussel mytilus galloprovincialis was recorded in 79 % of sampling areas that provided suitable rocky-shore habitat. the highest biomass (25.4 ± 18.7 kg/m²) was recorded in the mid-shore on marcus island, and total intertidal biomass in the langebaan/saldanha system was estimated at 281.6 tons. including farmed and subtidal stocks, the area supports 4 362 tons m. galloprovincialis in summer and 959 tons in winter. no live individuals of the european shore crab carcinus maenas were recorded, but a single dead carapace was found. the european periwinkle littorina saxatilis had a limited distribution within langebaan lagoon and supported a total population of 2.4 million individuals. the alien anemone sagartia ornata was recorded as locally abundant (up to 426 ± 81 individuals/m²) and was also restricted in distribution. despite the extensive invasion of the saldanha bay–langebaan lagoon system by m. galloprovincialis, the threat to the west coast national park appears to be limited by low wave action and low nutrient supply to the rocky shores within the park. although not present in detectable numbers, c. maenas poses a major threat to the park, should it become established there. the threats posed by l. saxatilis and s. ornata are presently unquantified but appear slight. key words: west coast national park, marine alien species, mytilus galloprovincialis, carcinus maenas, littorina saxatilis, sagartia ornata. t.b. robinson, c.l. griffiths and n. kruger, zoology department and marine biology research institute, university of cape town, rondebosch, 7700 republic of south africa. issn 0075-6458 79 koedoe 47/1 (2004) introduction on a global scale, species are constantly being moved from their areas of origin to new locales. such range extensions can occur naturally, but are frequently aided (intentionally or non-intentionally) by humans (mack et al. 2000). as a result, the marked world-wide increase in animal, plant, and microbial immigrations, has been found to roughly track the increase in human commerce (mack et al. 2000). the principal vectors of human-mediated marine invasions are ballast water (williams et al. 1988), mariculture (minchin 1996), sediment held in ballast tanks (carlton 1985), and ship hull fouling (minchin 1996). as south africa's second largest bulk port (unit for maritime studies 2000), the port of saldanha faces an elevated risk of inoculation by alien species. this, in combination with the increasing rate at which invasions are occurring globally, and the threats posed by marine invasive species (berman et al. 1992; griffiths et al. 1992; grosholz & ruiz 1996; carlton 1996), has led to concerns regarding the protection of indigenous fauna of areas surrounding the port. these concerns have been heightened by the location of the port within saldanha bay and langebaan lagoon, parts of which form the west coast national park (wcnp). established in 1985, the wcnp is the only major marine reserve robinson.qxd 2004/04/10 09:49 page 79 along the west coast of south africa. the park includes langebaan lagoon, jutten, malgas and marcus islands and an extensive terrestrial area surrounding most of the lagoon. the lagoon itself supports a high diversity of marine taxa (day 1959), while the surrounding sand flats and salt marshes host numerous species of palaearctic migrant waders during the southern summer (summers et al. 1977). the lagoon is recognised under both the ramsar and bonn conventions and, as such, holds the status of a wetland of international importance. of the 17 introduced marine species recorded in south africa by griffiths (2000), nine are listed to have occurred, or to presently occur, within saldanha bay. they are the bivalves tapes philippinarum and mytilus galloprovincialis, the gastropods haliotis koedoe 47/1 (2004) 80 issn 0075-6458 fig. 1. map of saldanha bay and langebaan lagoon, showing the boundaries of the west coast national park and the 28 areas which were sampled. robinson.qxd 2004/04/10 09:49 page 80 rufescenens and littorina saxatilis, the crabs pilumnus hirsutus and carcinus maenas, and the ascidians diplosoma listerianum, ciona intestinalis and ascidiella aspersa. of these, t. philippinarum and h. rufescenens populations have become extinct, while d. listerianum, c. intestinalis and a. aspersa are restricted to harbour environments. the introduced status of p. hirsutus is dubious. thus, only the mussel m. galloprovincialis, the gastropod littorina saxatilis and the crab c. maenas, are known to have established naturalised populations outside of harbours in the saldanha bay region. in 2002, acuña et al. (pers. comm.) recorded the non-native anemone sagartia ornata within langebaan lagoon. there are thus four alien marine species currently found within the saldanha bay–langebaan lagoon system. in response to the growing global awareness of the threats posed by invasive species, the international maritime organisation (imo) (a specialised agency of the united nations), in conjunction with the global environment facility (gef) and the united nations development programme (undp), has set up the global ballast water management programme (globallast). incorporated within globallast is an initiative to undertake port baseline surveys of active shipping harbours. at present only six demonstration ports have been selected worldwide, with saldanha bay being representative of the african continent. in view of the threat posed by invading species to the wcnp, and the international focus on the status of alien species in this region, the objective of this study is to set a benchmark regarding the status of marine alien species within and surrounding the west coast national park. with this in place, it will be possible to resurvey the area in future years and determine whether the existing alien species have spread or increased in density, or whether new invasive species have colonised the area. methods mytilus galloprovincialis the shoreline of saldanha bay and langebaan lagoon was divided into 28 areas, each approximately 3 km in length (fig. 1). area 1 (east shore of marcus island) and area 10s (schaapen island) covered less than 3 km, but due to their distinct nature were sampled as separate areas. within each of the areas where rocky shores occurred, three rocky shore sites were sampled, and at each of these, triplicate transects were surveyed. areas of vertical wharf were not sampled. a 0.5 m² quadrat was rolled up each transect from mean low water of spring tide (mlws) to mean high water of spring tide (mhws) and the cover accounted for by each mussel species estimated and summed in square metres. three quadrats (one each in the high, mid, and low shore zones), each measuring 0.016 m², were sampled within areas of 100 % mussel cover along each transect. the various mussel species were separated out and the whole wet mass of individuals of each species weighed separately, using a spring balance. together the whole wet weight and cover of mytilus galloprovincialis were used to calculate the biomass present per meter square (m²) and per linear meter (m). the total linear distance covered by rocky shore within each 3 km area was calculated from the south african navy chart san sc 2, or estimated while working in the field if the rocky shore was too small to be shown on the map. mussel biomass per linear meter, and length of rocky shore within each sector were then used to calculate the total mussel biomass supported in each 3 km area. carcinus maenas within each sampling area in which rocky shore occurred, a five-minute intertidal search was conducted by four researchers, during which rocks were overturned in an effort to locate c. maenas. crab traps (18.8 l volume) made of 1.5 cm mesh were also set at 17 subtidal sites around the perimeter of the bay and mouth of the lagoon (fig. 1). six baited traps were set at each site at 6 pm and were retrieved at 10 am the following morning. the trap design was previously tested in an area known to be heavily invaded by c. maenas (table bay harbour), and proved to be extremely effective (up to 50 crabs being caught per trap). twelve subtidal areas within the bay (fig. 1) were also dredged using a day dredge dragged for five to ten minutes, in order to detect subtidal populations. littorina saxatilis the same basic sampling procedure applied to m. galloprovincialis was used to determine the density of l. saxatilis within the survey area. in each of the issn 0075-6458 81 koedoe 47/1 (2004) robinson.qxd 2004/04/10 09:49 page 81 3 km sampling areas containing beds of spartina maritima (the known habitat of l. saxatilis), three sites were visited. at each site three transects were layed between mlws and mhws. counts were done within a 0.5 m² quadrat to determine the number of l. saxatilis individuals per m² and per linear meter of s. maritima bed. the total length of s. maritima bed in each 3 km sampling area was then used to calculate the total density of l. saxatilis. sagartia ornata within each 3 km sampling area, all sites offering suitable habitat (i.e. s. maritima beds or sandy shores with shallow underlying rock) were surveyed for s. ornata. at each site where s. ornata was detected, at least two transects were run from mlws to mhws. by rolling a 0.5 m² quadrat up each transect, the number of s. ornata individuals per m² in the high, mid and low shore were determined, as were the number of individuals per linear meter. the length of suitable habitat in each 3 km sector was then used to calculate overall density of s. ornata. results mytilus galloprovincialis mytilus galloprovincialis was found to be widely distributed across all areas of potenkoedoe 47/1 (2004) 82 issn 0075-6458 fig. 2. total biomass supported by mytlius galloprovincialis in each sampling area. tial habitat within saldanha bay and langebaan lagoon. this species was recorded in sampling areas 1–9 in saldanha bay, was absent throughout most of the lagoon, then reoccurred on the south side of the entrance to the bay. m. galloprovincialis was present in 11 of the 14 sampling areas (79 %) that had rocky shores. the indigenous mussels choromytilus meridionalis and aulacomya ater were recorded in only three and four sampling areas respectively (fig. 2). total standing stocks of m. galloprovincialis per 3 km sampling area are shown in fig. 2. although occurring through out the intertidal, the biomass of this species was centered in the mid-shore. the highest biomass of m. galloprovincialis (25.4 ± 18.7 (sd) kg/m²) was recorded in the mid-shore on marcus island, followed by the mid shore of sample area 3 (23.9 ± 77 (sd) kg/m²). significant biomasses of 8.3–18.7 ± 4.7 (sd) kg/m² were also supported in the mid shore of sampling areas 6, 8 and 9. although m. galloprovincialis was detected within langebaan lagoon (sampling area 23), and on the relatively sheltered south side of the bay (sampling areas 26, 27 and 28), only a few scattered individuals were present here, representing minimal biomass (0.01 ± 0.23, 0.06 ± 0.21, 0.26 ± 0.1 (sd) kg/m² respectively). mytilus galloprovincialis had a total intertidal biomass of 281.6 tons within the saldanha bay–langebaan lagoon system (table 1). this constituted 79.7 % of the total 353.2 tons of intertidal mussels within the system. carcinus maenas no live c. maenas were recorded in any of the rocky shore sampling areas, despite the fact that many of the sample sites contained what appeared to be ideal habitat. areas 22–28 in particular offered protected rocky robinson.qxd 2004/04/10 09:49 page 82 shores with boulders and numerous potential food species for c. maenas. a single carapace measuring 3.5 cm was, however, found in sampling area 3. no c. maenas individuals were recorded during trapping, although the crab plagusia chabrus, and the whelks burnupena cincta and nassarius vinctus were common in the traps. no subtidal populations of c. maenas were recorded at sites that were dredged. littorina saxatilis the periwinkle l. saxatilis was only recorded in areas 14–20 (fig. 3), and was restricted to s. maritima beds in the high shore zone. area 19 supported the highest average densities at 433 ± 123 (sd) individuals/m². however, as this area supported few spartina maritima beds, the overall density of l. saxatilis was much lower than in areas 19 and 20 which support vast beds of s. maritima. a total population of 2.4 million individuals was estimated to occur within the park. sagartia ornata very high numbers of the anemone s. ornata were recorded throughout the intertidal zone in areas 21–23, issn 0075-6458 83 koedoe 47/1 (2004) table 1 total rocky intertidal wet biomass (tons) of mytilus galloprovincialis, choromytilus meridionalis and aulacomya ater within saldanha bay and langebaan lagoon. the percentage contribution made by each species to the total mussel biomass of the system is given biomass (tons) % contribution mytilus galloprovincialis 281.6 9.7 choromytilus meridionalis 52.5 14.9 aulacomya ater 19.1 5.4 total 353.2 fig. 3. total number of individuals of (a) littorina saxatilis and (b) sagartia ornata recorded in each sampling area. (a) (b) robinson.qxd 2004/04/10 09:49 page 83 with highest number occurring in the midshore zone. the highest densities were recoded in the mid-shore of area 21 and peaked at 426 ± 81 (sd) individuals/m². much lower densities were recorded in areas 11, 18 and 19. despite limited suitable habitat being found in areas 13 and 14, no s. ornata were recorded here (fig. 3). discussion mytilus galloprovincialis mytilus galloprovincialis was first recorded along the west coast of south africa in the early 1980s (grant & cherry 1985), but by 1990 had become the dominant intertidal mussel in the area (van erkom schurink & griffiths 1990). as with all aggressive invaders, m. galloprovincialis possesses several characteristics that enable its rapid and successful spread at the expense of indigenous species. these features include a high resistance to desiccation, high fecundity and high growth rate (hockey & van erkom schurink 1992). mytilus galloprovincialis has a widespread distribution across the rocky shores of the area. being filter feeders, mussels tend to achieve maximal biomass, growth and condition in areas of moderately high wave action (steffani & branch 2003). thus we would expect m. galloprovincialis to achieve maximal biomass within the wave-exposed bay, and lower biomass within the sheltered lagoon. this was generally the case, with mussel biomass being low in sheltered areas along the west side of the lagoon (areas 23–28), and high opposite the mouth of the bay (areas 6, 8 and 9). by far the highest mussel biomass, in particular of m. galloprovincialis, was recorded on marcus island (fig. 2). the low biomass within langebaan lagoon itself, i.e. sampling areas 11–23, can be related to both the low wave exposure experienced in this region and the depleted nutrient supply. shannon & stander (1977) found chlorophyll concentrations within the lagoon to be significantly lower than in saldanha bay. this is a consequence of the low rate of water interchange between the bay and lagoon (shannon & stander 1977), coupled with the uptake of nutrients by macrophytes. the low wave action and turbulence experienced along these shores compound the above, resulting in the low mussel biomass/m² observed (fig. 3). although m. galloprovincialis had a large overall biomass of 281.6 tons within the rocky intertidal zone of saldanha bay and langebaan lagoon, these shores have not always been the only ones to support populations. in the mid 1990s m. galloprovincialis established beds on the sand flats of the center banks of langebaan lagoon. these supported a biomass of 77.64 tons by 1998 (robinson & griffiths 2002). these beds, however, underwent a dramatic reduction during 2000 and 2001 (hanekom & nel 2002), and by april 2003 had completely disappeared (griffiths pers. obs.). thus, for the purpose of calculating the present total intertidal mussel biomass of the system, only rocky shore populations have been taken into account. in addition to intertidal stocks, the saldanha bay–langebaan lagoon supports a highly productive mussel culture industry (fig. 1), where m. galloprovincialis is grown from ropes strung from rafts. the area supports 60 rafts with an average of 332 ropes per raft (grant et al. 1998). using the biomass per rope given by grant et al. (1998), the total biomass of cultured mussels varies between 4 080 tons in summer and 677 tons in winter. consequently, the overall biomass of m. galloprovincialis within the saldanha bay–langebaan lagoon system is estimated to range between 4 362 tons in summer and 959 tons in winter. carcinus maenas carcinus maenas was first detected along the south african coast in 1983, within table bay harbour (joska & branch 1986). by 1984, this globally recognised invader (welch 1968; zeildler 1978; yamada 2001) had established breeding populations both in table bay and bloubergstrand. by 1990, it had been recorded at eight sites along the koedoe 47/1 (2004) 84 issn 0075-6458 robinson.qxd 2004/04/10 09:49 page 84 west coast, including saldanha bay (le roux et al. 1990). the record from saldanha bay was of a single mating pair found in the dam next to the iron ore loading jetty (sample area 6). the european shore-crab is well known predator of molluscs (welch 1968; hughes 1979) and has caused dramatic declines in mollusc populations in other parts of the world (yamada 2001). to date, no such effects have been recorded in south africa, but this may be due to lack of research. in south africa, the factor limiting the spread of c. maenas is a paucity of suitable sheltered habitat (yamada 2001). as c. maenas is restricted to wave-protected habitats along the south african coast, le roux et al. (1990) predicted that the detrimental impacts the c. maenas invasion would be most strongly felt in sheltered areas such as harbours, bays and lagoons. thus the wcnp is particularly at risk. although two c. maenas individuals were recorded in saldanha bay in 1990 (le roux et al. 1990), no live individuals have subsequently been recorded (despite extensive sampling throughout the area during annual university of cape town student camps). no live animals were captured during the directed trapping operations carried out in this study either. due to the vigour with which c. maenas has invaded other areas and the high level of shipping traffic passing through saldanha bay, the total absence of live individuals was unexpected. the occurrence of a single carapace is also curious. one possibility is that a population exists at such low densities that no live individuals could be found. this would be surprising considering that at least some individuals were present 11 years previously. littorina saxatilis this small intertidal periwinkle was first recorded in south africa in 1974 (day 1974). the only known south african populations are in the langebaan and knysna lagoons (hughes 1979), although the present status of the knysna population is unknown. on the south african coast this species is restricted to sheltered salt marshes and lagoons, where it occurs on the stems of the cord grass spartina maritima (hughes 1979), although in its home range it occurs in crevices on rocky shores (gibson et al. 2001). despite establishing dense local populations, l. saxatilis is geographically restricted within the wcnp (fig. 3), and has not spread dramatically in the more than 20 years that it has colonized this area. no ecological effects of the invasion are known, although this species could form an abundant food source for predators such as wading birds and crabs. sagartia ornata this anemone is widely distributed throughout western europe, britain and the mediterranean (manuel 1981), and was first recorded in south africa in 2002 (acuña et al. in press). this record was the first in the southern hemisphere. at present, the species appears to be restricted to langebaan lagoon, where it occurs intertidally in s. maritima beds and on rocks covered by sand. along british coasts sagartia ornata is, however, known to occur in crevices on rocky shores and on kelp holdfasts (gibson et al. 2001). as such, there is potential for this species to spread widely along the south african west coast, which offers cold waters and vast kelp beds which typify its home range. it is thought that this species was unintentionally introduced into the bay via shipping. the distribution of s. ornata within the wcnp is presently restricted (fig. 3). however, follow-up surveys are needed to ascertain whether it is still spreading within the park, or whether populations have stabilised. the presence of a total of 3.1 million individuals within the wcnp is cause for concern. the ecological impacts of this invasion are likely to be restricted to local effects on small invertebrate prey species. issn 0075-6458 85 koedoe 47/1 (2004) robinson.qxd 2004/04/10 09:49 page 85 conclusions mytilus galloprovincialis has a wide distribution within the saldanha bay–langebaan lagoon system, but densities are low within the wcnp itself. it is thought that the invasion of the wcnp and surrounding areas by m. galloprovincialis has reached saturation, and that the biomass supported within the intertidal zone will not increase significantly. although no live individuals of carcinus maenas were recorded during this study, the potential threat posed by this crab to the natural biota of the wcnp is of considerable concern and the failure of this species to become established remains a paradox. littorina saxatilis has not spread significantly since it was first recorded in langebaan lagoon in the late 1970s and, due to its benign habits, poses little threat to indigenous biota. due to the paucity of attachment surfaces it is unlikely that sagartia ornata will extend its range within the lagoon. however, suitable habitat within saldanha bay and along the adjoining coast may see the spread of this species, although it is felt that strong wave action in this region may moderate this range extension. it is interesting to note that although the saldanha bay–langebaan lagoon system hosts more alien marine species than any other site along the south african coast, the number is still substantially lower than similar international sites such as san francisco bay (carlton et al. 1990). in order to ensure the future protection of the natural fauna of the wcnp against the threat of invasive alien marine species, conservation authorities need to be proactive. many developed countries, such as australia and the united states of america, employ a preemptive approach to marine invasions. in these systems conservation officials are equipped with a list and full description of invasive species which could potentially invade the area. this list includes known global invaders and problem species which are known to occur in similar areas elsewhere. together with routine surveys, this approach has allowed early detection of a number of invasions. this is important, as it is only in the early stages of marine invasions that removal is a viable management option (bax 2000). it is thus suggested that a booklet of known and potential marine invasive species be compiled for the south african coast, and that this be made available to south african national parks officials in marine parks. in addition, periodic surveys such as the one done in this study should be undertaken to monitor the spread and population densities of established invaders. references bax, n. 2000. control options. pp. 248–256. in: preston, g., g. brown & e. van wyk (eds.). best management practices for preventing and controlling invasive alien species. cape town: the working for water programme. berman, j., harris, l., lambert, w., m. buttrick, & m. dufresne. 1992. recent invasions of the gulf of maine: three contrasting ecological histories. conservation biology 6: 435–441. carlton, j.t. 1985. transoceanic and interoceanic dispersal of coastal marine organisms: the biology of ballast water. oceanography and marine biology annual review 23:313–371. carlton, j.t. 1996. marine bioinvasions: the alteration of marine ecosystems by nonindigenous species. oceanography 9: 36–43. carlton, j.t., thompson, j.k., l.e. schemel, & f.h. nichols. 1990. remarkable invasion of san francisco bay (california, usa) by the asian clam potamocorbula amurensis. i. introduction and dispersal. marine ecology progress series 66: 81–94. day, j.h. 1959. the biology of langebaan lagoon: a study of the effect of shelter from wave action. transactions of the royal society of south africa 26: 475–545. day, j.h. 1974. a guide to the marine life on south african shores. cape town: a.a. balkema. gibson, r., b. hextall & a. rogers. 2001. photographic guide to the sea and shore life of britain and north-west europe. new york: oxford university press. grant, w.s. & m.i. cherry. 1985. mytilus galloprovincialis lmk. in southern africa. journal of experimental marine biology and ecology 90: 179–191. grant, j., stenton-dozey, j., monteiro, p.m.s., g. pitcher & k. heasman. 1998. shellfish culture in the benguela system: a carbon budget of saldanha bay for raft culture of mytilus gallokoedoe 47/1 (2004) 86 issn 0075-6458 robinson.qxd 2004/04/10 09:49 page 86 provincialis. journal of shellfish research 17: 41-49. griffiths, c.l. 2000. overview on current problems and future risks. pp: 235–240. in: preston, g., g. brown & e. van wyk (eds.). best management practices for preventing and controlling invasive alien species. cape town: the working for water programme. griffiths, c.l., hockey, p.a.r., c. van erkom schurink & p.j. le roux. 1992. marine invasive aliens on south african shores: implications for community structure and trophic functioning. south african journal of marine science 12: 713–722. grosholz, e.d. & g.m. ruiz. 1996. predicting the impact of introduced marine species: lessons from the multiple invasions of the european green crab carcinus maenas. biological conservation 78: 59–66. hanekom, n. & p. nel. 2002. invasion of sandflats in langebaan lagoon, south africa, by the alien mussel mytilus galloprovincialis: size, composition and decline of the populations. african zoology 37: 197–208. hockey, p.a.r. & c. van erkom schurink. 1992. the invasive biology of the mussel mytilus galloprovincialis on the southern african coast. transactions of the royal society of south africa 48: 123–139. hughes, r.n. 1979. south african populations of littorina rudis. zoological journal of the linnean society 65: 119–126. joska, m.a.p. & g.m. branch. 1986. the european shore-crab–another alien invader? african wildlife 40(2): 63–65. le roux, p.j., g.m. branch & m.a.p. joska. 1990. on the distribution, diet and possible impact of the invasive european shore crab carcinus maenas (l.) along the south african coast. south african journal of marine science 9: 85–93. mack, r.n., simberloff, d., lonsdale, w.m., evans, h., m. clout & f.a. bazzaz. 2000. biotic invasions: causes, epidemiology, global consequences, and control. ecological applications 10: 689–710. manuel, r.l. 1981. british anthozoa. london: linnean society minchin, d. 1996. management of the introduction and transfer of marine mollusks. aquatic conservation of marine and freshwater ecosysteyms 6: 229–244. robinson, t.b. & c.l. griffiths. 2002. invasion of langebaan lagoon, south africa, by mytilus galloprovincialis–effects on natural communities. african zoology 37: 151–158. shannon, l.v. & g.h. stander. 1977. physical and chemical characteristics of water in saldanha bay and langebaan lagoon. transactions of the royal society of south africa 42 (3 &4): 441–459. steffani, c.n. & g.m. branch. 2003. growth rate, condition, and shell shape of m. galloprovincialis: responses to wave exposure. marine ecological progress series 246: 197–209. summers, r.w. 1977. distribution, abundance and energy relationships of waders (aves: charadrii) at langebaan lagoon. transactions of the royal society of south africa 42: 483–494. unit for marine studies. 2000. south africa's maritime industries. cape town: m.e.r.i.t van erkom schurink, c. & c.l. griffiths. 1990. marine mussels of southern africa—their distribution patterns, standing stocks, exploitation and culture. journal of shellfish research 9: 75–85. welch, w.r. 1968. changes in abundance of the green shore crab, carcinus maenas (l.), in relation to recent temperature changes. fishery bulletin 67 (2): 337–345. williams, r.j., griffiths, f.b., e.j. van der wal & j. kelly. 1988. cargo vessel ballast water as a vector for the transport of non-indigenous marine species. estuarine, coastal and shelf science 26: 409–420. yamada, s.b. 2001. global invader: the european green crab. oregon: corvallis. zeidler, w. 1978. note on the occurrence of the european shore crab carcinus maenas (linn., 1758) in australia. south australian naturalist 53 (1): 11–12. issn 0075-6458 87 koedoe 47/1 (2004) robinson.qxd 2004/04/10 09:49 page 87 filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 siebert2.qxd semi-arid savanna of the potlake nature reserve and surrounding areas in sekhukhuneland, south africa s.j. siebert, m. matthee and a.e. van wyk siebert, s.j., m. matthee and a.e. van wyk. 2003. semi-arid savanna of the potlake nature reserve and surrounding areas in sekhukhuneland, south africa. koedoe 46(1): 29–52. pretoria. issn 0075-6458. a hierarchical classification, description, and ecological and floristic interpretations are presented on the vegetation types of the semi-arid northern savanna of the sekhukhuneland centre of plant endemism. relevés were compiled in 47 stratified random plots. a twinspan classification, refined by braun-blanquet procedures, revealed eight plant communities, classified as four associations, one of which is subdivided into five sub-associations. for each plant community, the floristic richness, endemism and conservation status was determined. much of the plant community distribution can be ascribed to environmental factors and anthropogenic disturbance. an ordination (decorana), based on floristic data, showed environmental gradients that possibly exist between plant communities and associated habitats. the floristic information, proposed classification, general description and vegetation key, can be used for the identification and monitoring of protected areas, land-use planning, and further conservation research. key words: biodiversity, conservation, limpopo, phytosociology, savanna, sekhukhuneland, syntaxonomy, ultramafic. s.j. siebert, m. matthee and a.e. van wyk, department of botany, university of pretoria, pretoria, 0002 republic of south africa. issn 0075-6458 29 koedoe 46/1 (2003) introduction phytosociological studies have been conducted on savanna vegetation types of the bushveld complex in limpopo province (bredenkamp & van vuuren 1977; breebaart & deutschlander 1997; siebert et al. 2002b, 2002c). however, the vegetation of most areas still remains to be investigated and described, both on reconnaissance level and in more detail. such an area is the semi-arid plains and hills of the potlake nature reserve and surrounding mecklenburg region, previously described as the northern dry mixed bushveld (siebert et al. 2002a) of the sekhukhuneland centre of plant endemism (scpe) (siebert & van wyk 2001; van wyk & smith 2001). the area is situated to the north of the steelpoort subcentre of the scpe, which is floristically noteworthy in that many rare and endemic species with distributions correlated with the geological substrate occur here (siebert et al. 2001, 2002d). this paper forms part of a wider investigation into the vegetation of the scpe (siebert et al. 2002a). it is envisaged that the identification, classification and description of the various vegetation units of conservation areas will contribute to the knowledge of the plant diversity and biological intricacies of the region. classification of the vegetation is essential for the formulation of a management policy and for proper land use planning. this paper shows which plant communities (and rare species) are presently conserved in sekhukhuneland, and by comparison with data from siebert et al. (2002b, 2002c, 2002d), it will identify ecosystems not presently conserved. therefore, the paper aims to provide ecological and floristic data of the region’s semiarid bushveld and the associated habitats, by characterising and interpreting the vegetation units, as well as constructing a vegetation key to assist with the identification of plant communities in this homogeneous vegetation siebert2.qxd 2005/12/09 10:28 page 29 type. an assessment of the plant diversity, endemism/near-endemism and red data list taxa in the plant communities of the study area is supplied for future conservation planning initiatives in the region. the vegetation described here only includes those plant communities that are part of the acacia tortilis–eragrostis barbinodis northern dry mixed bushveld (siebert et al. 2002a). a synthesis in the form of a phytosociological table is presented as a summary of the major vegetation types. study area the study area is situated directly south of the strydpoort mountains in limpopo province between latitude 24º15'00''– 24º30'00''s and longitude 29º30'00''– 30º10'00''e (fig. 1). this northern part of the scpe is restricted to the eastern bushveld complex, situated on top of the lower zone of the rustenburg layered suite (visser et al. 1989). the lower zone stretches around the eastern side of the leolo mountains – from lebowakgomo eastward and then southward towards steelpoort where it fades. koedoe 46/1 (2003) 30 issn 0075-6458 fig. 1. location of the semi-arid savanna of the sekhukhuneland centre in the limpopo province. siebert2.qxd 2005/12/09 10:28 page 30 issn 0075-6458 31 koedoe 46/1 (2003) the study area (potlake nature reserve and surrounding areas) covers approximately 1250 km² and comprises a moderately heterogeneous physiography (land type survey staff 1987), which is underlain by a heterogeneous geology of pyroxenite belts covered by surficial deposits of alluvium and scree (visser et al. 1989). other prominent rocks include jagdlust harzburgite, shelter norite and karoo dolerite. potlake nature reserve is small (approximately 30 km²) and conserves the vegetation of the largest serpentinised ultramafic outcrop in sekhukhuneland. sekhukhuneland lies in the summer rainfall region, with the semi-arid northern parts receiving mean annual precipitation of 430 mm (south african weather bureau 1998). the rainfall pattern is strongly influenced by the area’s topography (siebert 1998), varying from 416 mm in the east to 499 mm in the west, and from 500 mm in the south to 456 mm in the north (erasmus 1985). daily temperatures range from a minimum of 8 °c in winter to a maximum of 38 °c in summer, with a mean annual temperature of 20 °c (south african weather bureau 1998). temperatures vary at different localities within the study area, also correlating strongly with physiographic regions, being higher for lowlying valleys and lower for high-lying plateaus (buckle 1996). minimum temperatures of below freezing point are extremely rare, even at higher altitudes. the difference in altitude between the two most extreme locations is approximately 200 metres. bredenkamp & van vuuren (1977) recognised various savanna vegetation types on the adjacent pietersburg plateau, which show a definite floristic affinity with the semi-arid savanna of the scpe (siebert et al. 2002a). in this semi-arid region of sequential bands of hills and plains, the predominant characteristic vegetation feature is low open savanna, which is mostly restricted to the ultramafic areas between the leolo mountains and the strydpoort mountains. acocks (1988) mapped the vegetation of the study area as tropical bush and savanna veld types, namely mixed bushveld and sourish mixed bushveld. acocks (1988) accurately distinguished the major vegetation type of the study area as mixed bushveld, which is an acacia-dominated savanna of relatively drier plains (siebert et al. 2002a). according to acocks (1988) the vegetation of the mountain slopes of the leolo mountains’ northern ranges is sourish mixed bushveld, which links with the vegetation on the southern slopes of the serala subcentre and the western slopes of the blyde river subcentre (matthews 1991). however, it is not a typical sour bushveld and needs to be studied further to determine its true identity, because the area under discussion is semi-arid (mean annual rainfall between 350 and 450 mm per annum). in the 1940s, sekhukhuneland was already highlighted as an area that is rapidly becoming a semi-desert (codd 1949). methods floristic and habitat data of the study area were obtained from 47 stratified random plots, sampled by matthee in 1978, and verified by siebert in 1998. longitude and latitude readings were recorded for the 1978 sample units during the 1998 survey. stratification was based on soil form, terrain type and aspect. to standardise the plot size for the sekhukhuneland survey, plots in the savanna were fixed at 20 m x 20 m (siebert et al. 2002a). within each sample plot, all species were recorded and a cover-abundance value assigned to each species according to the braun-blanquet scale (muellerdombois & ellenberg 1974). plant species names conform to those of retief & herman (1997). terminology to describe vegetation structure follows edwards (1983). environmental data recorded in each sample plot include terrain type, aspect, slope, geology, soil form and rockiness of soil surface. all relevé data are stored in the turboveg database (hennekens 1996a), and managed by the department of botany, university of pretoria (mucina et al. 2000). a first approximation of a vegetation classification for the sekhukhuneland region in general, based on the total floristic data set of 415 relevès (siebert et al. 2002a), was obtained by the application of the twoway indicator species analysis (twinspan) (hill 1979a). this first step identified several major vegetation types for the scpe (siebert et al. 2002a) and was used to subdivide the data set into five phytosociological tables. the table representing northern siebert2.qxd 2005/12/09 10:28 page 31 koedoe 46/1 (2003) 32 issn 0075-6458 ta bl e 1 p hy to so ci ol og ic al ta bl e of th e se m iar id s av an na o f t he p ot la ke n at ur e r es er ve a nd s ur ro un di ng a re as siebert2.qxd 2005/12/09 10:28 page 32 issn 0075-6458 33 koedoe 46/1 (2003) ta bl e 1 (c on tin ue d) siebert2.qxd 2005/12/09 10:28 page 33 koedoe 46/1 (2003) 34 issn 0075-6458 ta bl e 1 (c on tin ue d) siebert2.qxd 2005/12/09 10:28 page 34 issn 0075-6458 35 koedoe 46/1 (2003) ta bl e 1 (c on tin ue d) siebert2.qxd 2005/12/09 10:28 page 35 koedoe 46/1 (2003) 36 issn 0075-6458 ta bl e 1 (c on tin ue d) siebert2.qxd 2005/12/09 10:28 page 36 issn 0075-6458 37 koedoe 46/1 (2003) ta bl e 1 (c on tin ue d) siebert2.qxd 2005/12/09 10:28 page 37 koedoe 46/1 (2003) 38 issn 0075-6458 dry mixed bushveld was again subjected to twinspan. braun-blanquet procedures were used to further refine the resultant classification in the megatab computer programme (hennekens 1996b) (table 1). all syntaxa were subsequently hierarchically classified and described according to the code of phytosociological nomenclature (weber et al. 2000). the ordination algorithm detrended correspondence analysis (decorana) (hill 1979b) (fig. 2) was applied to illustrate possible floristic relationships between communities and to confirm possible gradients between the vegetation and the physical environment. in order to facilitate the identification of areas of high conservation potential, the alpha diversities of the different plant communities were calculated. the alpha diversity (plant species richness) is defined as the number of species per unit area within a homogeneous community or the total number of species per community (whittaker 1977). a 400 m² sample plot was taken as the unit area within a homogeneous community. distribution ranges of all the taxa were verified at the national herbarium [pre], pretoria, to identify any taxa endemic to the region (siebert et al. 2002d). all taxa were also checked against red data lists of southern african plants (hilton-taylor 1996; victor 2002) to determine their conservation status in south africa. mean vegetation cover percentage (mcp) was determined for each species in every community. for the purpose of this study, mcp is the sum of a species’ percentage cover in all the relevés of a community, divided by the total number of relevés for that community (species cover was determined using the following conversion from the braun-blanquet scale: r = 0.5%; + = 1%; 1 = 3%; a = 8%; b = 18%; 3 = 37%). species were grouped into their respective growth forms, namely trees, forbs or grasses, and the total vegetation cover percentage (tcp) was determined by adding the mcps for each growth form in each community. results and discussion classification since the study area lies in the geological and climatically uniform dry and warm northern region of the sekhukhuneland centre (siebert et al. 2002a), no major climatic variation plays a role in local differentiation of plant communities. communities were not always distinctive in the field. this might be attributed to the homogeneity of the environmental factors, causing a complex transition pattern of habitats and associated vegetation. the hierarchical classification of the vegetation reinforces the relationship between habitat and plant communities (fig. 2). major plant communities relate to soil properties, aspect and terrain type. a summary of selected community attributes is given in table 2. distribution of the 16 identified sekhukhuneland endemic/near-endemic and rare/threatened species (siebert et al. 2002d) among the various plant communities of this semi-arid savanna is listed in table 3. analyses resulted in the identification of eight plant communities, classified as four associations, one of which is subdivided into five sub-associations (table 1). plant communities of the scpe’s semi-arid savanna (acacia tortilis–eragrostis barbinodis northern dry mixed bushveld) are classified as follows: urochloo mosambicencis–acacion tortilis 1. panico colorati–crotonetum menyhartii 2. schmidtio pappophoroidis–acacietum tortilis 2.1 schmidtio pappophoroidis–acacietum tortilis grewietosum bicoloris 2.2 schmidtio pappophoroidis–acacietum tortilis rhigozetosum obovati 2.3 schmidtio pappophoroidis–acacietum tortilis diospyretosum lycioidis 2.4 schmidtio pappophoroidis–acacietum tortilis acacietosum niloticae 2.5 schmidtio pappophoroidis–acacietum tortilis indigoferetosum rhytidocarpae 3. enneapogono cenchroidis–salvadoretum australis 4. urochloo panicoidis–agavetum americanae siebert2.qxd 2005/12/09 10:28 page 38 issn 0075-6458 39 koedoe 46/1 (2003) table 2 environmental factors and selected attributes associated with the different plant communities of the potlake nature reserve and surrounding areas factors/attributes syntaxa 1 2.1 2.2 2.3 2.4 2.5 3 4 number of relevés 6 5 4 7 11 7 4 3 total number of species 59 110 92 87 110 109 98 55 mean number of spp. per relevé 25 50 46 34 38 39 48 24 grass cover (%) 14 9 12 17 19 18 15 6 forb cover (%) 8 16 15 19 16 12 20 8 tree cover (%) 10 28 27 2 8 3 10 3 total vegetation cover (%) 32 53 54 38 43 33 45 17 geology1 p p p p/sh p/sh/q sh/q sh/q q topographic position2 f/m f/m f/m f/m p/f p/f p/f p slope (°) 1–3 3–5 1–5 1–5 1–3 1–3 1–3 1–3 aspect e nw se s ns n n nesw predominant soil form3 gs sn/mw va/gs sd/gs va bo sn rock cover (%) 15–50 10–15 10–20 10–20 05–10 05–10 05–15 05–10 mean rock size (mm) 250–500 100–400 300–500 100–400 200–400 300–400 250–400 50–100 1p = pyroxenite; sh = serpentinized harzburgite; q = alluvium 2m = midslope; f = footslope; p = plain 3bo = bonheim; gs = glenrosa; mw = milkwood; sd = shortlands; sn = steendal; va = valsrivier fig. 2. relative positions of all the relevés of the semi-arid savanna of the sekhukhuneland centre along the first and second axis of an ordination. numbers correspond with the plant communities in table 1. siebert2.qxd 2005/12/09 10:28 page 39 koedoe 46/1 (2003) 40 issn 0075-6458 a vegetation key is presented to facilitate plant community identification (table 4). the definitions are broad indications of typical groups and should be seen as a guideline. a diagnostic characteristic of the vegetation or habitat is given, followed by the diagnostic and most conspicuous species of a group. the first species is restricted to the specific group only, and the second is dominant in the group, but also occurs in other groups. where one species is given, no species was restricted to the group only. description acacia tortilis–eragrostis barbinodis northern dry mixed bushveld of the scpe is predominantly restricted to the plains and lower slopes of undulating hills. surface rocks are common and abundant in many of the communities, with soil clay percentages varying from 15 % to 35 %. according to the edwards (1983) classification, the structure of the vegetation is mainly closed tree savanna (undisturbed vegetation), shrubland thicktable 3 sekhukhuneland centre endemic/near-endemic and red data list plant taxa recorded for the potlake nature reserve and surrounding areas taxon family syntaxa 1 2.1 2.2 2.3 2.4 2.5 3 4 aloe castanea lili . . . . . . . #+ aloe burgersfortensis lili . . $+ . . . . . boscia foetida subsp. minima capp rr rr . rr . r+ r+ . euphorbia sp. nov. (a pers. comm.) euph . $r . . . . . . gossypium herbaceum malv . . . . n+ . . . grewia vernicosa tili . . . . . . . #r hermbstaedtia rogersii amar . . . . $+ $r . . heurnia stapelioides ascl . #+ . . . . . . hibiscus barnardii malv r$ r$ r$ . . . . . leucas capensis [form] (ws 13007) lami . $r $r $+ $1 $+ $+ $+ pegolettia senegalensis aste . . . . . nr n+ . petalidium oblongifolium acan #1 . . . . . . . phyllanthus sp. nov. (s 470) euph $r . . . $r $r . . plinthus rehmannii aizo v#+ v#+ v#r v#r v#+ v#+ . . polygala sp. nov. (s 449) poly $+ . . . $r . . . rhus engleri anac 1 #+ #r #r #1 . #r scpe endemics 3 3 3 1 4 3 1 1 scpe near-endemics 3 3 2 2 2 1 1 2 red data list 3 3 2 2 2 3 2 0 restricted to syntaxon 1 2 1 0 1 0 0 2 restricted to association 1 3 0 2 total for syntaxon 7 6 5 4 7 6 4 3 restricted to association 7 13 4 3 endemism: $ = endemic, # = near-endemic; red data list: r = rare, v = vulnerable, n = not threatened in the northern provinces of south africa, but in other areas of southern africa; abundance in communities: 1 = abundant, + = frequent, r = rare, . = absent; collectors: a = archer, s = siebert, w = van wyk; bold blocks represent community/syntaxon specific taxa. siebert2.qxd 2005/12/09 10:28 page 40 issn 0075-6458 41 koedoe 46/1 (2003) et (bush encroachment) and open tree savanna (man made grassland). urochloo mosambicencis–acacion tortilis all. nova hoc loco nomenclatural type: schmidtio pappophoroidis–acacietum tortilis (holotypus), association 2 described in this paper. this alliance is classified under the panico maximi–acacietea tortilis (winterbach et al. 2000). it is related to the colophospermum mopane–euclea divinorum tree savanna (van rooyen et al. 1981) and the urochloo mosambicensis–eucleion divinorum (siebert et al. 2002c). habitat. vegetation representing this alliance could be interpreted as a semi-arid vegetatable 4 vegetation key to the syntaxa of potlake nature reserve and surrounding areas in the sekhukhuneland centre leads/description go to/syntaxon 1a glenrosa soils only 1. panico colorati–crotonetum menyhartii (panicum coloratum & croton menyhartii) 1b various soil forms ... 2 (acacia grandicornuta & panicum maximum) 2a mean rock size < 100 mm 4. urochloo panicoidis–agavetum americanae (agave americana & seddera suffruticosa) 2b mean rock size > 100 mm … 3 (albizia anthelmintica & acacia tortilis) 3a steendal soils only 3. enneapogono cenchroidis–salvadoretum (salvadora australis & tragus berteronianus) australis 3b various soils … 4 (schmidtia pappophoroides & eragrostis barbinodis) 4a pyroxenite only … 5 (acacia luederitzii & tribulus terrestris) 4b pyroxenite and/or harzburgite … 6 (monechma divaricatum) 5a mainly northern aspects 2.1 schmidtio pappophoroidis–acacietum tortilis (grewia bicolor & lantana rugosa) grewetosum bicoloris 5b mainly southern aspects 2.2 schmidtio pappophoroidis–acacietum tortilis (rhigozum obovatum & balanites maughamii) rhigozetosum obovati 6a footslopes/midslopes 2.3 schmidtio pappophoroidis–acacietum tortilis (diospyros lycioides & acacia mellifera) diospyretosum lycioidis 6b footslopes/plains (cadaba termitaria) … 7 7a valsrivier soils (acacia nilotica & kleinia longiflora) 2.4 schmidtio pappophoroidis–acacietum tortil is acacietosum niloticae 7b bonheim soils 2.5 schmidtio pappophoroidis–acacietum tortilis (indigofera rhytidocarpa & boscia foetida) indigoferetosum rhytidocarpae siebert2.qxd 2005/12/09 10:28 page 41 koedoe 46/1 (2003) 42 issn 0075-6458 tion unit driven by rural livelihood systems (this includes overgrazing by goats, irregularly planted fields and intensive wood harvesting), a highly erodable soil profile (weakly structured upper horizons that wash away during heavy rains), and periodic droughts. these habitats occur on gentle footslopes (1–5°) on all aspects (table 2) of the potlake nature reserve and surrounding areas in sekhukhuneland. the dominant soil forms are the steendal and valsrivier forms. average rock size varies from 100 mm to 400 mm and covers 10–50 % of the soil surface (table 2). vegetation structure. the structure represents a transition from pristine savanna to man made grassland to degraded systems with bush encroachment. the alliance is characterised by species group u (table 1). the forb seddera suffruticosa, grasses eragrostis barbinodis and urochloa mosambicensis, and tree acacia tortilis are the most abundant diagnostic species of this syntaxon. acacias are the most distinctive, dominant and largest group of trees for this vegetation type and are generally associated with typical mixed bushveld landscapes (acocks 1988). other prominent herbaceous species include the forbs felicia clavipilosa and kleinia longiflora, and grasses aristida congesta, enneapogon cenchroides, panicum maximum and tragus berteronianus. prominent woody taxa of the alliance are acacia grandicornuta, albizia anthelmintica, ehretia rigida and rhus engleri. floristic diversity. it is speculated that much of the vegetation distribution in the semi-arid savanna of the potlake area can be attributed to the vegetation dynamics and historic evolution of the region, which is influenced by the surrounding flora, continual disturbance by over grazing/harvesting and aridity. plant species migrations from especially the pietersburg plateau (bredenkamp & van vuuren 1977) and disjunct local occurrences with areas even further north (van rooyen et al. 1981), had a considerable influence on the local patterns and species composition of plant communities. loreau & mouquet (1999) demonstrated that local diversity, community patterns and ecosystem processes, are strongly influenced by plant migration. this is further supported by the presence of species typically from higher rainfall areas in the lowveld, and includes balanites maughamii, combretum imberbe, diospyros mespiliformis, philenoptera violacea and ptaeroxylon obliquum. these taxa were recorded in the vicinity of the banks of the olifants river in sekhukhuneland. 1. panico colorati–crotonetum menyhartii ass. nova hoc loco nomenclatural type: relevé 383 (holotypus), table 1 this association is related to the loudetio simplicis–eucleetum linearis and the enteropogono macrostachyos–sclerocaryetum birreae (siebert et al. 2002b). habitat. this association occurs on the lower midslopes and footslopes of hills. the habitat is predominantly restricted to eastern aspects that are gently sloped and slightly eroded (table 2). loam soils are predominant, mostly the glenrosa form, with a black coloured ortic a-horizon over a lithocutanic b-horizon. the surface rock cover percentage is high and the mean rock diameter is between 250 mm and 500 mm (table 2). vegetation structure. the association represents short, open to closed shrubland of which the diagnostic species are presented in species group a (table 1). diagnostic small trees/shrubs of the association are maerua cafra and mundulea sericea. diagnostic forbs include blepharis pruinosa, cleome angustifolia, helichrysum cerastioides, petalidium oblongifolium and polygala sp. nov. (van wyk & siebert 13311). diagnostic grasses of this association are typical for gravel soils and include eustachys paspaloides, fingerhuthia africana and panicum coloratum. prominent woody species of the tree cover of 10 % are acacia mellifera, balanites maughamii, croton menyhartii and rhus engleri. forb cover is sparse and of the lowest recorded for the study area. conspicuous grasses of the association siebert2.qxd 2005/12/09 10:28 page 42 issn 0075-6458 43 koedoe 46/1 (2003) include enneapogon scoparius and eragrostis barbinodis, which contribute substantially towards the grass cover of 14 %. floristic diversity. this is the most pristine plant community of the urochloo mosambicencis–acacion tortilis. floristically the association shows an affinity with association 2 in species groups f, h and m (table 1). the average number of plant species encountered per sample plot is 25, with the total number for this association being 59 (six relevés) (table 2). the scpe near-endemic petalidium oblongifolium is restricted to this association. a further three endemics, two near-endemics and three red data list taxa were recorded for this association (table 3). 2. schmidtio pappophoroidis–acacietum tortilis ass. nova hoc loco nomenclatural type: relevé 348 (holotypus), table 1 this association is related to the colophospermum mopane–acacia tortilis–urochloa mosambicensis tree savanna of van rooyen et al. (1981) and the enneapogono cenchroides–acacietum leiorachis (siebert et al. 2002c). habitat. the association is typical of plains, footslopes and lower midslopes of ultramafic hills. it is managed for grazing purposes in and adjacent to the potlake nature reserve. it is situated on relatively sloped areas (1–5°) with a surface rock cover of 5–15 % (table 2). mean rock diameter is approximately 100–500 mm (table 2). vegetation structure. vegetation is short, sparse to open shrubland that is characterised by the diagnostic taxa presented in species group b (table 1). the association is typified by a high diversity of herbaceous taxa. corchorus asplenifolius, corbichonia decumbens, limeum viscosum, melhania rehmannii, ptycholobium contortum, solanum coccineum and s. panduriforme are the diagnostic forbs that contribute significantly towards the forb cover of 16 %. diagnostic grasses typical of gravel and/or lime-rich soils are common, namely schmidtia pappophoroides and tricholaena monachme. grass cover is 15%, with the most abundant grasses including eragrostis barbinodis, panicum maximum and tragus berteronianus. prominent woody species, typical of the association and contributing substantially to the tree cover of 14 % (the highest for the urochloo mosambicencis–acacion tortilis), include the small trees acacia mellifera, balanites maughamii, commiphora pyracanthoides and euclea undulata. acacia grandicornuta, a. tortilis, dichrostachys cinerea, ehretia rigida and rhus engleri are the dominant small trees of the association. floristic diversity. the association shows strong floristic affinities with the other associations of the study area in especially species groups m, r and t (table 1). the average number of plant species encountered per sample plot is 40, with the total number of species for this association being 164 (34 relevés). of the four taxa restricted to this association, three are scpe endemics, namely aloe burgersfortensis, euphorbia sp. nov. (archer pers.comm.) and hermbstaedtia rogersii, one is a scpe near-endemic, huernia stapelioides, and one a red list taxon, gossypium herbaceum (table 3). the highest number of endemics (7), near-endemics (4) and red list taxa (4) occur in this association, which is also the community with the highest numbers of taxa (13) with conservation value. 2.1 schmidtio pappophoroidis–acacietum tortilis grewetosum bicoloris subass. nova hoc loco nomenclatural type: relevé 354 (holotypus), table 1 this sub-association represents short thicket on black loam soils underlain by gypsumrich lower horizons. the dominant soil forms are melanic, namely the steendal form (soft carbonate b-horizon) interspersed by the milkwood form (a-horizon underlain with hard rock). the community occurs on northern aspects of moderate footslopes and lower midslopes of hills (table 2). rock cover is siebert2.qxd 2005/12/09 10:28 page 43 koedoe 46/1 (2003) 44 issn 0075-6458 sparse. in table 1, species group c contains the diagnostic species, which is dominated by the diagnostic woody taxa cadaba aphylla and grewia bicolor. diagnostic forbs are hybanthus enneaspermus, indigofera tristoides, limeum pterocarpum, and tephrosia burchellii, and succulents are huernia stapelioides, opuntia ficus-indica (naturalised alien tree) and talinum arnotii. with the highest tree cover for the study area (28 %), prominent taxa of the sub-association include acacia grandicornuta, a. mellifera, albizia anthelmintica, commiphora pyracanthoides, croton menyhartii and ptaeroxylon obliquum. forb cover is average and includes frequently occurring taxa such as becium filamentosum, hibiscus praeteritus, lantana rugosa, melhania rehmannii and seddera suffruticosa. grass cover is low (9 %) and is dominated by aristida congesta, eragrostis barbinodis, panicum maximum, schmidtia pappophoroides and tragus berteronianus. floristically the sub-association is noteworthy in that, together with subassociation 2.2, it shares a unique link of small forb diversity with association 3 (species group q; table 1). this plant community is floristically the richest of the urochloo mosambicencis–acacion tortilis, with the highest number of plant species encountered per sample plot (50) and the highest total number recorded per community, namely 110 (five relevés) (table 2). one scpe near-endemic, huernia stapelioides, and one scpe endemic, euphorbia sp. nov., are restricted to this sub-association (table 3). 2.2 schmidtio pappophoroidis–acacietum tortilis rhigozetosum obovati subass. nova hoc loco nomenclatural type: relevé 368 (holotypus), table 1 this sub-association comprises tall, closed shrubland on footslopes and midslopes. it occurs on relatively deep loam soils of the valsrivier form (ortic a-horizon over a pedocutanic b-horizon), which is interspersed with shallow soils of the glenrosa form (ortic a-horizon over a lithocutanic b-horizon). it prefers south-easterly aspects of hills which are characterised by gentle slopes and a rocky surface (table 2). species group d (table 1) includes rhigozum obovatum, the only diagnostic woody species, and the diagnostic forbs aneilema hockii, barleria prionitis, commelina livingstonii, melhania virescens, mollugo nudicaulis and polygala hottentotta. other diagnostic species include the succulents portulaca quatrifida and stapelia gettliffei. grasses are common (12% cover) and aristida congesta, eragrostis barbinodis, panicum maximum and tragus berteronianus dominate this layer. frequently occurring forbs are felicia clavipilosa, melhania rehmannii, phyllanthus maderaspatensis and seddera suffruticosa. conspicuous trees/shrubs that contribute significantly to the relatively high tree cover of 27 % include acacia luderitzii, a. tortilis, croton menyhartii and ptaeroxylon obliquum. floristically the sub-association shows the same relationships as sub-association 2.1 (table 1). a sekhukhuneland centre endemic, aloe burgersfortensis, is restricted to this community within the study area (table 3), but can also be found in the mountain bushveld of the maandagshoek, burgersfort, steelpoort and dwarsrivier areas. 2.3 schmidtio pappophoroidis–acacietum tortilis diospyretosum lycioidis subass. nova hoc loco nomenclatural type: relevé 360 (holotypus), table 1 the short, sparse and open tree savanna of this sub-association occurs on footslopes and lower midslopes with red loam soils. soils are predominantly of the shortlands form (ortic a-horizon with a red-structured bhorizon), interspersed with soils of the glenrosa form. it lies on gently sloped, southern aspects of undulating, ultramafic hills (table 2). rock surface cover and size are relatively high (table 2). diagnostic species are listed in species group g (table 1). diospyros lycioides is the only diagnostic woody species and eragrostis trichophora siebert2.qxd 2005/12/09 10:28 page 44 issn 0075-6458 45 koedoe 46/1 (2003) (along the footpaths) the only diagnostic grass. however, the syntaxon is rich in diagnostic forbs, including coccinia rehmannii, dipcadi viride, melhania acuminata, phyllanthus incurvus and pollichia campestris. dominant grasses such as aristida congesta, cenchrus ciliaris, eragrostis barbinodis, schmidtia pappophoroides, tragus berteronianus and urochloa mosambicensis contribute to a high grass cover of 17 %. forb cover is also relatively high (19 %) and includes the prominent species blepharis integrifolia, monechma divaricatum, pechuel-loeschea leubnitzia, phyllanthus maderaspatensis and seddera suffruticosa. tree cover is the lowest for the study area (2 %) and includes conspicuous small trees, namely acacia tortilis, a. mellifera and ehretia rigida. species groups h and k show the major floristic relationship between this vegetation unit and the other sub-associations of association 2 (table 1). no taxa with conservation value are restricted to the sub-association, but four taxa of conservation value were recorded (table 3). 2.4 schmidtio pappophoroidis–acacietum tortilis acacietosum niloticae subass. nova hoc loco nomenclatural type: relevé 348 (holotypus), table 1 a short, closed to open secondary savanna is characteristic of this widespread sub-association. it occurs on heavily grazed, deep, loam/clay soils (500–1000 mm) of the valsrivier form (ortic a-horizon). it prefers gently sloped footslopes and plains, and can be found on either northern or southern aspects of ultramafic hills (table 2). the geology is heterogeneous and rock size is relatively large, but rock cover is sparse (table 2). diagnostic species are listed in species group i (table 1) and include the trees acacia nilotica and ziziphus mucronata. diagnostic forbs are plentiful and include gossypium herbaceum, hibiscus palmatus, kohautia cynanchica, lycium cinereum, polygala uncinata, seddera capensis and tragia rupestris. eragrostis rigidior is the only diagnostic grass and is characteristic of bare patches in the overgrazed veld. although disturbed, the grass cover is dense and the highest for the study area (19 %), including unpalatable grasses such as aristida adscensionis, a. congesta, enneapogon cenchroides, eragrostis barbinodis and tragus berteronianus, and palatable grasses such as panicum maximum and urochloa mosambicensis. the sub-association is dominated by encroaching acacia tortilis trees. other conspicuous trees include albizia anthelmintica, cadaba termitaria, dichrostachys cinerea and ehretia rigida. forbs are abundant (16 %) and this growth form is dominated by a rich diversity (corchorus asplenifolius, gisekia africana, hermannia odorata, hibiscus praeterius, indigastrum costatum, lantana rugosa, leucas capensis, melhania rehmannii, monechma divaricatum, phyllanthus maderaspatensis and solanum panduriforme), which contributes substantially to its recognition as one of the communities with the highest total number of species, namely 110 (11 relevés) (table 1). gossypium herbaceum subsp. africanum, a red data list taxon described as insufficiently known (k) for swaziland, is the only species restricted to the sub-association (table 3). 2.5 schmidtio pappophoroidis–acacietum tortilis indigoferetosum rhytidocarpae subass. nova hoc loco nomenclatural type: relevé 370 (holotypus), table 1 this sub-association of short, open, disturbed savanna in zones of previously cultivated land is common on soils characterised by melanic loams of the bonheim form (pedocutanic b-horizon). this syntaxon occurs on the plains and on lower footslopes of warmer northern aspects (table 2). mean rock size is large, but rock cover is sparse (table 2). species group l contains the diagnostic species for this sub-association (table 1) and includes two grasses, eragrostis biflora in the shade and the alien dactyloctenium aegyptium where water collects. no diagnostic woody species were siebert2.qxd 2005/12/09 10:28 page 45 koedoe 46/1 (2003) 46 issn 0075-6458 recorded, but diagnostic forbs are frequent, and include indigofera circinnata, i. rhytidocarpa, ipomoea sinensis, leucas sexdentata, limeum sulcatum, peliostomum leucorrhiza and trianthema salsoloides. certain alien species are diagnostic for this sub-association, namely acalypha segetalis, amaranthus thunbergii and schkuhria pinnata. corallocarpus bainesii, dipcadi gracillimum, kohautia aspera and phyllanthus burchellii are rare indigenous species that were also recorded. dominant grasses of the sub-association, which contribute substantially to the relatively high grass cover of 18%, include aristida congesta, chloris virgata, eragrostis barbinodis, sporobolus ioclados, tragus berteronianus and urochloa mosambicensis. prominent forbs are becium filamentosum, corchorus asplenifolius, felicia clavipilosa, gisekia africana, hermannia modesta, phyllanthus maderaspatensis and seddera suffruticosa. acacia tortilis dominates the sparse tree cover (3 %), and a. grandicornuta and dichrostachys cinerea encroaches to a lesser degree. the sub-association is the last of the strong floristic relationship between associations 1 and 2 in species group m (table 1). a sole relationship, possibly indicating an ecotone, is shared with association 3 in species group o (table 1). the plant species diversity per community is high (table 2) and includes taxa of conservation value (table 3). 3. enneapogono cenchroidis–salvadoretum australis ass. nova hoc loco nomenclatural type: relevé 356 (holotypus), table 1 habitat. this association occurs on predominantly warm, but locally moister, northerly aspects of ultramafic hills. mean rock size is high and varies between 250 mm and 400 mm in diameter, and covers 5–15 % of the soil surface (table 2). it is characterised by gentle footslopes and undulating plains (table 2). the dominant soil form is brackish alluvium of the steendal form (table 2). vegetation structure. species group n (table 1) contains the diagnostic species for this short, closed or open tree savanna in the semi-arid savanna of the scpe. it is characterised by diagnostic tree species such as acacia robusta, cordia monoica, maerua edulis and salvadora australis. diagnostic grasses are plentiful and include brachiaria deflexa, diplachne eleusine, enneapogon desvauxii, eragrostis curvula and setaria verticillata. in these relatively undisturbed vegetation units, abutilon grandiflorum, hibiscus micranthus, justicia odora, pegolettia senegalensis and the succulent sansevieria hyacinthoides, are the diagnostic forbs of the association. prominent, frequently occurring trees with a cover of 10 % include acacia grandicornuta, boscia foetida and cadaba termitaria. aristida adscensionis, cenchrus ciliaris, chloris virgata, enneapogon cenchroides, e. scoparius, sporobolus ioclados and tragus berteronianus dominate the grass cover of 15 %. forbs are abundant and the forb layer is well developed with species such as barleria senensis, b. virgula, becium filamentosum, blepharis integrifolia, hermannia modesta, ruellia patula and tribulus terrestris. the forb cover of 20 % is the highest for the study area. floristic diversity. floristically this association shows strong affinities with most of the other associations of the semi-arid savanna of the scpe, however not including association 1 (table 1). a very close floristic relationship with association 2 is visible in species group r (table 1) on average, a high mean number of 48 species were recorded per sample plot for the association, with a total of 98 different plant species overall (four relevés) (table 2). no taxa of conservation value are restricted to this association; however, one scpe endemic, one scpe near-endemic and two red data list taxa were recorded (table 3). siebert2.qxd 2005/12/09 10:28 page 46 issn 0075-6458 47 koedoe 46/1 (2003) 4. urochloo panicoidis–agavetum americanae all. nova hoc loco nomenclatural type: relevé 390 (holotypus), table 1 habitat. this association occurs on all aspects along gently sloped undulating plains (table 2). soils are divers and may include turf soils such as the arcadia form (vertic a-horizon), clay soils such as the steendal form (melanic a-horizon), loam soils such as the shortlands form (ortic ahorizon, red-structured b-horizon) or sandy soils such as the hutton form (ortic a-horizon, red-apedale b-horizon). the soil surface is sparsely (5–10%) covered by stones with a small mean size of 50–100 mm (table 2). vegetation structure. this plant community is described as sparse, open, species-poor savanna on disturbed, compacted soils adjacent to natural footpaths of domestic livestock, around settlements of local people, and along natural migration routes of game in the reserve. diagnostic species are presented in species group s (table 1). a sparse herb layer dominates the vegetation unit (8 % forb cover), with the dominant diagnostic forbs including aptosimum lineare, chamaesyce prostrata (alien species), dicoma tomentosa, felicia mossamedensis, geigeria burkei, indigofera spicata, melhania forbesii, phyllanthus parvulus, sida dregei and withania somnifera. diagnostic grasses are aristida canescens, eragrostis racemosa and urochloa panicoides. the large alien succulent, agave americana, is the most conspicuous diagnostic species in the association. large individuals of schotia brachypetala are scattered in this continually disturbed system and dense clumps of the succulent euphorbia tirucalli are also common. the woody layer is species-poor and sparse (3 % tree cover) due to long-term wood harvesting, overgrazing and trampling by livestock and game. aristida congesta and eragrostis barbinodis are other dominant species of the 6 % grass cover of the association. no other common species of the semi-arid savanna of the study area occur frequently in this association. floristic diversity. floristically the association shows a slight relationship with the other associations in species groups t and u (table 1). however, this association is distinct and is only grouped with the other associations due to its locality in the semi-arid savanna of the scpe and lack of a better placement as a true anthropogenic plant community. plant species encountered per sample plot are on average 24 and the total number recorded for this association was 55 (three relevés) (table 2). two of the recorded taxa of conservation value are restricted to this association for the study area, namely the scpe near-endemics aloe castanea and grewia vernicosa (table 3). ordination on a regional scale, the area occupied by the urochloo mosambicencis–acacion tortilis of the scpe is characterised as naturally sparsely vegetated due to low rainfall. it contains many syntaxa that are locally typical for this habitat, occurring nowhere else in the scpe and possibly even the world. when compared with other habitats of the scpe, the major environmental factors such as climate and geology are relatively homogeneous for this major vegetation group. however, the ordination indicated possible gradients that are mainly influenced by factors such as rock cover, soil moisture and soil depth. all these gradients correlate closely with each other and have a strong influence on the vegetation structure and species composition of the different plant communities (table 1). the ordination scatter diagram displays the distribution of relevés along the first and second axes (fig. 2). the vegetation units are represented as groups, their distribution on the diagram corresponding with certain physical environmental conditions. the rockiness and soil properties determine a definite vegetation gradient that is depicted by both the first (eigen value = 0.629) and second axis (eigen value = 0.344). rockiness, soil moisture and soil depth determines siebert2.qxd 2005/12/09 10:28 page 47 koedoe 46/1 (2003) 48 issn 0075-6458 moisture retention and drainage of any habitat. the gradient on the first axis expresses rock cover as a percentage of the soil surface, with the left of the scatter diagram representing rocky foot slopes and the right depicting the open valley plains. this relates closely to soil moisture, as rock covered soils have reserves of retained moisture in the topsoil which is available for plant uptake. on the second axis, deeper soils located at the bottom of the graph have prolonged water availability, because deep clayey soils retain moisture for a longer period. steep slopes with shallow soils dry out quickly and are at the top of the diagram. overall, the scatter diagram expresses a gradient from the shallow, rocky soils in the top, left corner (high available soil moisture, mainly in the wet season) to deep, clay soils in the bottom, right corner (low available soil moisture due to water retention in wet season). structure to assist future land-use planning, the current relationships between the tree cover and herbaceous layer of the plant communities of the potlake nature reserve and surrounds were investigated and documented. co-existence graphs can be applied in the future management of the reserve to determine the state of the grass-forb-tree relationship. briske & hendrickson (1998) explain changes in vegetation structure, as a result of over-utilization, as an ecological consequence to minimise the effect of selective grazing/harvesting. change in vegetation structure will reduce the probability of localised population extinction caused by long-term selective grazing/harvesting. the fig. 3. total cover percentage (tcp) for selected growth forms, as a function of co-existence: (a) grass tcp plotted against forb tcp, and (b) herbaceous (forbs and grasses) tcp plotted against tree tcp. siebert2.qxd 2005/12/09 10:28 page 48 issn 0075-6458 49 koedoe 46/1 (2003) graphs of co-existence can be further applied by establishing control plots to monitor the state of the ecosystem by comparing it to the 20-year old information provided in this paper. the trend depicted in fig. 3a is that of soil quality, with grass cover percentage decreasing as the soil quality decreases. to the left of the diagram the soils have been altered by erosion and compacting, which has made it unsuitable for colonisation by forbs and grasses. grass cover is a maximum on old fields and heavily grazed areas probably due to a history of fertilisation (sub-associations 2.4 and 2.5). to the right of the diagram, the soil chemistry has been changed by high salt concentrations deposited by non-perennial streams, favouring colonisation by salt tolerant forbs and a decrease in grass cover. the optimum condition for current land-use activities would be one with both the forb and grass tcps at 15% each, which means allowing the land to revert back to an optimal system for grazing. figure 3b illustrates the relationship between tree cover and the understorey herbaceous layer. tree and herbaceous cover increases linearly, but once the tree cover reaches 15 %, there is a noticeable drop in the herbaceous cover due to competition for resources such as light. the communities to the left of the diagram are typically open tree savanna and the communities to the right represent closed thickets. both the open savanna and the thickets are in some cases a result of anthropogenic and/or natural disturbance. when considering current land-use such as grazing, the optimum condition would be a tree tcp of 15 %, which would allow the herbaceous layer to attain a tcp of 30 % and higher. a short, semi-closed savanna would be the ideal structure to protect the soil, and provide enough light for forbs and grasses to co-exist in these semi-arid conditions and, at the same time, provide optimal grazing conditions. dynamics classification of the vegetation indicated that intensive long-term grazing and harvesting has modified the vegetation structure of this semi-arid savanna type of the scpe in certain areas, resulting in disturbed plant communities. altered species composition, decreased biomass and decreased species richness have previously been recorded in the savanna communities of other southern african communal grazing lands (higgins et al. 1999). highly degraded ecosystems, especially those in semi-arid regions, do not recover to their original state once the stress loads are lessened (rapport & whitford 1999), for these systems are event-driven (ellis & swift 1988). the semi-arid savanna of the scpe is probably an event-driven system as it is floristically related to mopaneveld (van rooyen et al. 1981), and should the disturbance be removed, its species composition will return to a type of savanna that will be determined by the environmental factors at that given time (du plessis 2001). if the disturbance was drought, the species composition alternates between this event and the one during wetter periods, but will remain stable. however, if the species composition is changed as a result of persistent disturbance, the vegetation seems unable to recover to its so-called stable state. this is the case for the over-utilised areas of semiarid savanna in sekhukhuneland, with old fields, heavily grazed rangelands, open cast mines and mine dumps covered by a grass layer of annual and alien species infestations (siebert & van wyk 2001). ongoing unsustainable utilisation of the woody canopy of the plant communities in the scpe semi-arid savanna is detrimental to its future recovery, as it has been shown that the woody canopy of communities in the scpe has a nurse plant effect on younger seedlings of trees and especially succulents (thrash 1998). furthermore, due to continuous intensive grazing in this region, no more protection is given to seedlings against fires and herbivores. hence, the main factor for vegetation structure, species distribution and depletion during the past 50 years have possibly been anthropogenical. small-scale disturbances act to increase the range of environmental conditions under which trees and grasses coexist as savanna (jeltsch et al. siebert2.qxd 2005/12/09 10:28 page 49 koedoe 46/1 (2003) 50 issn 0075-6458 1998). however, large-scale disturbance, such as the over-harvesting of trees in the scpe, result in grasslands dominated by annuals or bush encroachment by certain woody species. according to roth (1999), thorn scrub, a vegetation type of the dominican republic which the authors consider as structurally similar to the dominated acacia tortilis shrublands of this northern part of the scpe, is a form of secondary climax created through past land-use activities in areas once bearing more diverse dry woodlands. in semi-arid regions where event driven systems prevail, sustained disturbance will alter species composition to such an extent that the original, viable composition may never be regained (westoby et al. 1989). conservation an analysis of the size structure in southern african communal lands suggested that land used for grazing have a reduced capacity for regeneration that consequently predicts future species losses (higgins et al. 1999). large areas of the study area have already reached this stage and there are now certain vegetation units or areas that need special attention and should be considered for rehabilitation purposes. throughout the scpe the remaining wild populations of plants and animals are under intense pressure from the rapid developing mining industry and fast growing human population in the region (joubert & dreyer 2000; knoll 2002; siebert et al. 2002d). certain plant endemics and rare taxa of this semi-arid region of the scpe are restricted to specific communities. areas containing these communities require immediate attention as conservation priorities. furthermore, like in other centres of plant diversity of the world (stohlgren 1999), alien plant invasions pose a significant challenge to the unique species composition of habitats and distinctive plant communities in this part of the scpe. to further protect these unique species assemblages, attention needs to be given to the protection of important species with conservation value, such as local endemic, near-endemic and threatened taxa (table 2). the creation of nature reserves and/or resource reserves is one solution to this problem. if its size could be increased, the potlake nature reserve could be the solution to protect a large part of this unique semi-arid savanna. however, financial constraints, frequent droughts, illegal plant harvesting and unsustainable former land-use have left its scars in this reserve and its surrounds. conclusions the twinspan classification and its subsequent refinement by braun-blanquet procedures resulted in the delineation of eight plant communities. these communities can be related to certain environmental factors, the gradients of which are proposed in the decorana scatter diagram. moisture availability plays a major role in the survival of the plant species and therefore soil depth and rock cover are major determinants of the species composition of the communities. different events related to land-use history and current management practices, as well as natural factors such as soil structure, determine the current structure and distribution of these savanna plant communities. description and classification of the different vegetation units of potlake nature reserve and its surrounds makes a significant contribution towards the understanding of the semi-arid regions of the scpe. however, further more detailed ecological studies need to be done on old fields and degraded systems of different ages to derive hypothetical succession pathways for future grazing management in especially the potlake nature reserve and its surrounding grazing areas. proper and sound assessments of the region’s vegetation (including aspects such as species richness and threats) to determine its suitability for conservation are a prerequisite before suggestions concerning sustainable development and conservation can be made (bedward et al. 1992). the information supplied in this paper can be meaningfully applied to develop management and conservation plans for this species rich, but threatened, area. siebert2.qxd 2005/12/09 10:28 page 50 issn 0075-6458 51 koedoe 46/1 (2003) acknowledgements our appreciation to ms martie dednam, h.g.w.j. schweikerdt herbarium (pru), university of pretoria, for processing the plant specimens and to dr elizabeth retief, national herbarium (pre), pretoria, for assistance with plant identifications. a special thanks to staff at the department of botany, university of pretoria, namely ms franci du plessis for assistance with numerical analysis, prof. george bredenkamp for guidance with the descriptions and prof. noel van rooyen for supervising the fieldwork of ms maryna matthee in potlake nature reserve during 1978. the national research foundation, university of pretoria, andrew mellon foundation and department of environmental affairs and tourism, financially supported the research. references acocks, j.p.h. 1988. veld types of south africa. memoirs of the botanical survey of south africa 57: 1-146. bedward, m., r.l. pressey & d.a. keith. 1992. a new approach for selecting fully representative reserve networks: addressing efficiency, reserve design and land suitability with an iterative analysis. biological conservation 62: 115-125. bredenkamp, g.j. & d.r.j. van vuuren. 1977. the plant communities below turfloop dam, lebowa. pietersburg: university of the north. (series a17) breebaart, l. & m. deutschländer. 1997. the vegetation types and management units of goedverwacht farm in the mixed bushveld of the northern province, south africa. koedoe 40: 19-33. briske, d.d. & j.r. hendrickson. 1998. does selective defoliation mediate competitive interactions in a semiarid savanna? a demographic evaluation. journal of vegetation science 9: 611-622. buckle, c. 1996. weather and climate in africa. harlow: addison wesley longman. codd, l.e.w. 1949. the application of ecology to agricultural problems in south africa. statement and communication of the african regional scientific conference, pp. 115-119. du plessis, f. 2001. a phytosociological synthesis of mopaneveld. m.sc. dissertation, university of pretoria, pretoria. edwards, d. 1983. a broad-scale structural classification of vegetation for practical purposes. bothalia 14: 705-712. ellis, j.e. & d.m. swift. 1988. stability of african pastoral systems: alternate paradigms and implications for development. journal of range management 41: 450-459. erasmus, j.f. 1985. rainfall deciles for the transvaal region. pretoria: soil and irrigation research institute. (report gb/a/87/18) hennekens, s. 1996a. turboveg: software package for input, processing, and presentation of phytosociological data. user's guide. university of lancaster: ibn-dlo. hennekens, s. 1996b. megatab: a visual editor for phytosociological tables. version 1.0. user's guide. ulft: giesen & geurts. higgins, s.i., c.m. shackleton & e.r. robinson. 1999. changes in woody community structure and composition under contrasting landuse systems in a semi-arid savanna, south africa. journal of biogeography 26: 619-627. hill, m.o. 1979a. twinspan – a fortran program for arranging multivariate data in an ordered two way table by classification of individuals and attributes. new york: cornell university, ithaca. hill, m.o. 1979b. decorana – a fortran program for detrended correspondence analysis and reciprocal averaging. new york: cornell university, ithaca. hilton-taylor, c. 1996. red data list of southern african plants. pretoria: national botanical institute. (strelitzia 4) jeltsch, f., s.j. milton, w.r.j. dean, n. van rooyen & k.a. moloney. 1998. modelling the impact of small-scale heterogeneities on treegrass coexistence in semi-arid savannas. journal of ecology 86: 780-793. joubert, h. & a. dreyer. 2000. rocky road for rare endemics: granite mining threatens a rich mpumalanga environment. african wildlife 54: 18-19. knoll, c. 2002. operation wildflower: plant rescue at andalusite mine. urban green file 7: 19-21. land type survey staff. 1987. land types of the maps 2526 rustenburg, 2528 pretoria. pretoria: department of agriculture and water supply. (memoirs on the agricultural natural resources of south africa no. 8) loreau, m. & n. mouquet. 1999. immigration and the maintenance of local species diversity. the american naturalist 154: 427-440. matthews, w.s. 1991. phytosociology of the northeastern mountain sourveld. m.sc. dissertation, university of pretoria, pretoria. mucina, l., g.j. bredenkamp, d.b. hoare & d.j. mcdonald. 2000. a national vegetation database for south africa. south african journal of science 96: 497-498. mueller-dombois, d. & h. ellenberg. 1974. aims and methods of vegetation ecology. new york: wiley. siebert2.qxd 2005/12/09 10:28 page 51 retief, e. & p.p.j. herman. 1997. plants of the northern provinces of south africa: keys and diagnostic characters. pretoria: national botanical institute. (strelitzia 6) roth, l.c. 1999. anthropogenic change in subtropical dry forest during a century of settlement in jaiqui picado, santiago province, dominican republic. journal of biogeography 26: 739-759. siebert, s.j. 1998. ultramafic substrates and floristic patterns in sekhukhuneland, south africa. m.sc. dissertation, university of pretoria, pretoria. siebert, s.j. & a.e. van wyk. 2001. sekhukhuneland: floristic wealth versus platinum and chromium riches. veld & flora 87: 168-173. siebert, s.j., a.e. van wyk & g.j. bredenkamp. 2001. endemism in the flora of ultramafic areas of sekhukhuneland, south africa. south african journal of science 97: 529-532. siebert, s.j., a.e. van wyk & g.j. bredenkamp. 2002a. the physical environment and major vegetation types of sekhukhuneland, south africa. south african journal of botany 68: 127142. siebert, s.j., a.e. van wyk & g.j. bredenkamp. 2002b. vegetation ecology of sekhukhuneland, south africa: combretum hereroense-grewia vernicosa open mountain bushveld. south african journal of botany 68: 475-496. siebert, s.j., a.e. van wyk & g.j. bredenkamp. 2002c. vegetation ecology of sekhukhuneland, south africa: kirkia wilmsii-terminalia prunioides closed mountain bushveld. south african journal of botany 68: 497-517. siebert, s.j., j.e. victor, a.e. van wyk & g.j. bredenkamp. 2002d. an assessment of threatened plants and conservation in sekhukhuneland. plantlife 26: 7-18. south african weather bureau. 1998. unpublished report. department of environmental affairs and tourism, pretoria. stohlgren, t.j., d. binkley & g.w. chong. 1999. exotic plant species invade hot spots of native plant diversity. ecological monographs 69: 2546. thrash, i. 1998. association of three succulent plant species with woody canopy in the mixed bushveld, south africa. koedoe 41(1): 95-101. van rooyen, n., g.k. theron & n. grobbelaar. 1981. a floristic description and structural analysis of the plant communities of the punda milia-pafuri-wambiya area in the kruger national park. journal of south african botany 47: 585-626. van wyk, a.e. & g.f. smith. 2001. regions of floristic endemism in southern africa: a review with emphasis on succulents. pretoria: umdaus press. victor, j.e. 2002. south africa. pp. 93-120. in: golding, j.s. (ed). southern african plant red data lists. pretoria: sabonet. (sabonet report series no. 14) visser, d.j.l., f.j. coertze & f. walraven. 1989. explanation of the 1:1000000 geological map, fourth edition, 1984: the geology of the republics of south africa, transkei, bophuthatswana, venda and ciskei and the kingdoms of lesotho and swaziland. pretoria: the government printer. weber, h.e., j. moravec & j-p. theurillat. 2000. international code of phytosociological nomenclature. journal of vegetation science 11: 739768. westoby, m., b. walker & i. noy-meir. 1989. opportunistic management for rangelands at equilibrium. journal of range management 42: 266-274. whittaker, r.h. 1977. evolution of species diversity in land communities. evolutionary biology 10: 1-67. winterbach, r., g.j. bredenkamp, m.s. deutschländer & l. mucina. 2000. preliminary syntaxonomic scheme of vegetation classes for the central bushveld of south africa. pp. 123-127. in: white, p.s., l. mucina, j.s. leps & e. van der maarel (eds.). proceedings iavs symposium. uppsala: opulus press. koedoe 46/1 (2003) 52 issn 0075-6458 siebert2.qxd 2005/12/09 10:28 page 52 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <feff004700650062007200750069006b002000640065007a006500200069006e007300740065006c006c0069006e00670065006e0020006f006d0020005000440046002d0064006f00630075006d0065006e00740065006e0020007400650020006d0061006b0065006e0020006d00650074002000650065006e00200068006f00670065002000610066006200650065006c00640069006e00670073007200650073006f006c007500740069006500200076006f006f0072002000610066006400720075006b006b0065006e0020006d0065007400200068006f006700650020006b00770061006c0069007400650069007400200069006e002000650065006e002000700072006500700072006500730073002d006f006d0067006500760069006e0067002e0020004400650020005000440046002d0064006f00630075006d0065006e00740065006e0020006b0075006e006e0065006e00200077006f007200640065006e002000670065006f00700065006e00640020006d006500740020004100630072006f00620061007400200065006e002000520065006100640065007200200035002e003000200065006e00200068006f006700650072002e002000420069006a002000640065007a006500200069006e007300740065006c006c0069006e00670020006d006f006500740065006e00200066006f006e007400730020007a0069006a006e00200069006e006700650073006c006f00740065006e002e> /esp <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> /suo <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> /ita <feff00550073006100720065002000710075006500730074006500200069006d0070006f007300740061007a0069006f006e00690020007000650072002000630072006500610072006500200064006f00630075006d0065006e00740069002000500044004600200063006f006e00200075006e00610020007200690073006f006c0075007a0069006f006e00650020006d0061006700670069006f00720065002000700065007200200075006e00610020007100750061006c0069007400e00020006400690020007000720065007300740061006d007000610020006d00690067006c0069006f00720065002e0020004900200064006f00630075006d0065006e00740069002000500044004600200070006f00730073006f006e006f0020006500730073006500720065002000610070006500720074006900200063006f006e0020004100630072006f00620061007400200065002000520065006100640065007200200035002e003000200065002000760065007200730069006f006e006900200073007500630063006500730073006900760065002e002000510075006500730074006500200069006d0070006f007300740061007a0069006f006e006900200072006900630068006900650064006f006e006f0020006c002700750073006f00200064006900200066006f006e007400200069006e0063006f00720070006f0072006100740069002e> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 filelist convert a pdf file! page 1 page 2 page 3 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! shackleton.qxd fuelwood availability and use in the richtersveld national park, south africa c.m. shackleton, g. guthrie, j. keirungi and j. stewart shackleton, c.m., g. guthrie, j. keirungi and j. stewart. 2003. fuelwood availability and use in the richtersveld national park, south africa. koedoe 46(2): 1-8. pretoria. issn 0075-6458. concern has been voiced about the possible over-use of fuelwood from the riparian fringe by pastoralist herders in the richtersveld national park (rnp). this coincided with the current examination and modelling of the supply and use of ecosystem goods and services in the gariep basin as part of the southern african millennium assessment (safma). this paper reports on a study to index the current availability of deadwood within the riparian zone of rnp, its relationship with proximity to human habitation, and species preferences of the local herders. deadwood availability was assessed per woody species and on the ground in 12 transects within the riparian fringe. herders were interviewed regarding their species preferences, and the composition of woodpiles was examined. there was no relationship between the percentage of attached deadwood on the tree, or the percentage deadwood ground cover, and the distance from herder stockposts. euclea pseudobenus and tamarix usneoides were the dominant species in the riparian fringe. there was strong selection for ziziphus mucronata as a fuelwood species and only marginal or random selection for e. pseudobenus. tamarix usneoides and prosopis sp. were abundant in the riparian zone, but were not used for fuelwood. there was a significant difference between species with respect to the mean proportion of the stem that was dead, the highest being z. mucronata (± 28 % deadwood), followed by t. usneoides (± 12 %). most of the prosopis trees had no deadwood. across all species, the mean percentage dead per tree was approximately 15 %. additionally, detached deadwood covered just less than 9 % of ground area, averaged across all plots and transects. all the variables measured indicated that there seems to be little need for concern over the current fuelwood extraction activities of pastoralists within the rnp. there was no clear evidence of cutting of branches or deadwood. the abundance of both attached and detached deadwood was not depleted close to human habitation. there was still abundant deadwood, even on preferred species. the most preferred species (ziziphus mucronata) had the greatest mean proportion of deadwood. the herders stated that they only used deadwood, and that there was plenty. they rarely used driftwood washed down the river, although there was an abundance of it. thus, overall, there was no sign of depletion of the deadwood resource, even on favoured species, or degradation of the productive capacity for deadwood. key words: deadwood; fuelwood; preference; riparian fringe; selection c.m. shackleton (c.shackleton@ru.ac.za), g. guthrie, j. keirungi and j. stewart, department of environmental science, rhodes university, grahamstown, 6140 south africa. issn 0075-6458 1 koedoe 46/2 (2003) introduction fuelwood is the dominant energy form throughout southern africa and the rest of the developing world (gielink 1991; hall 1991; pandey 2002). this has consequences for human well-being as well as environmental quality. in terms of the latter, large-scale use of fuelwood has often been implicated in deforestation and desertification, with many historical official reports demanding interventions to limit perceived environmental damage, before total denudation occurred (leach & mearns 1996; sullivan 1999; dahlberg 2000). this became known as the fuelwood crisis; based on fuelwood gap shackleton.qxd 2005/12/09 11:24 page 1 models portraying that fuelwood supply was diminishing in the face of increasing demand from growing human populations, and that ultimately there would be insufficient fuelwood to meet human needs with attendant severe environmental degradation. the notion of a fuelwood crisis has been questioned in recent times (e.g. leach & mearns 1989; benjaminsen 1997; nagothu 2002; pandey 2002), primarily because the predictions of the models frequently failed to materialize. the failure of model predictions can be ascribed to a number of limitations in the original models, the primary ones, amongst others, being (i) failure to account for changes in consumer patterns with increasing scarcity, (ii) overlooking the contribution of wood from arable and residential areas, (iii) the strong coppicing ability of most woodland species and the faster growth rates of regrowth from cut stumps than from seedlings, (iv) tree yield and growth models taken from commercial forestry equations for harvestable timber, and therefore excluding smaller branches and twigs, and wood that would be unsuitable for timber, but acceptable as fuelwood, and (v) unrealistic time frames to models. the further the model was required to project into the future, the greater was the magnification of weak measurement precision and model assumptions. within south africa the majority of the rural population make extensive of fuelwood, despite increased electrification in the last decade. annual demand is approximately 11 million tons (dme 1996), well within sustainable supply at a national level (williams & shackleton 2002). however, in many areas there is a severe imbalance between demand and supply for fuelwood at a localised scale resulting in progressive deforestation. this is most apparent in areas of high human population density, although not ubiquitous to them all (banks et al. 1996). within this context, some conservationists have raised concerns about the possible over-use of fuelwood as the primary energy source of pastoralist herders in the richtersveld national park (rnp) (hendricks pers. comm.). within south africa the rnp is unusual in that, despite being a protected area, there are 26 permanent households who graze 6600 small livestock and who depend upon the environmental goods and services offered by the habitats of the rnp for many of their daily needs (hendricks pers. comm.). this includes fuelwood. whilst the rnp is a contractual park (a 30 year lease agreement from the nama community by the south african national parks (sanp) for conservation purposes), sanp is mandated with biodiversity conservation, and therefore seeks to ensure that landuses within rnp have minimal impact. hence the need to examine the use of fuelwood. this coincided with the current assessment and modelling of the supply and use of ecosystem goods and services in the gariep basin as part of the southern african millennium assessment (safma). the rnp represents one of the core study sites for the local level assessments of the safma, and fuelwood is one of the indicators of ecosystem status across all safma sites. for the rnp pastoralists, the riparian fringe of the gariep river, which forms the northern boundary of the park, is a key resource area in an extremely arid environment. during summer, the herders temporarily occupy stockposts within one kilometre of the river. they rotate from one stockpost to another every two to four months. the location of stockposts is fixed and permanent, but use thereof is periodic. in the winter months the herders move to the ‘buiteveld’; towards the centre of the park and away from the gariep river. during this time the bulk, but not all, of their fuelwood is still collected from the riparian fringe and transported to the buiteveld stockposts (hendricks pers. comm.). this is supplemented with biomass from dry shrubs, especially zygophyllum prismatocarpum. because of the concentration of fuelwood collection along the riparian fringe in rnp it can be assumed that the environmental impacts of wood use would be most apparent along the riparian zone. additionally, since it koedoe 46/2 (2003) 2 issn 0075-6458 shackleton.qxd 2005/12/09 11:24 page 2 has been shown in numerous studies (e.g. grundy et al. 1993; banks et al. 1996; luoga et al. 2001) that fuelwood collection is most intense close to human settlements rather than further away, it can be hypothesized that if there is insufficient fuelwood for herders in rnp, then a zone of depletion would be apparent radiating out from the stockposts. such a zone of depletion would be manifest as a reduced standing crop or abundance of fuelwood close to stockposts, as well as possibly increased signs of chopping or breakage on preferred species. the presence of such a zone would indicate that deadwood production was insufficient to meet localized demand, and hence the spatial extent of environmental impacts could be expected to gradually increase in the near term. this would not, however, imply that all wood would ultimately be depleted, because human and livestock population numbers within the rnp are limited by the contractual agreement. this paper reports on a study to index the current availability of deadwood (as the favoured fuelwood resource) within the riparian zone of rnp, its relationship with proximity to stockposts, and species preferences of the local herders. it sought to index the harvesting impact of local communities on the deadwood resource, without developing dataand timeintensive supply and demand models. as such the study attempted to provide support or negation for sanp's concerns regarding potential negative impacts of fuelwood extraction activities. study area the rnp is a contractual park located in the semi-arid region of the namaqualand magisterial district in the northwestern part of south africa, bordering onto namibia. it is regarded as the only true mountain desert in south africa, and is managed jointly by the local nama people and the south african national parks. the richtersveld experiences an arid climate. most of the region receives winter rainfall (may to september), but summer rainfall is not uncommon. between 1995-2001, the mean annual rainfall in the richtersveld national park was 82 mm, with rainfall variability tending to decrease from east to west and from south to north, although the varied topography increases the spatial variability in rainfall (hendricks pers. comm.). as is typical in arid environments, the rnp experiences massive temperature fluctuations from over 50 ºc in the summer to freezing point on some winter nights. the rnp is widely known as one of the world's most succulent-rich areas. this richness can be attributed to the large number and diversity of habitats in the area (van der walt 1992). the vegetation of the park has evolved within a water-stressed environment and is thus mainly succulent, comprising more than 700 species of herbs, shrubs and trees, geophytes and annuals (williamson 2000). it is estimated that 50 genera out of the total of 160 from the mesembryanthemaceae family occur in rnp (van der walt 1992). none of the preferred fuelwood species have a significant conservation status, in that none are listed as rare, endangered or endemic. there are 26 households resident within the rnp. they are nomadic pastoralists, rotating their herds of goats and sheep within the park in response to availability of forage. wood is their primary energy source for cooking and warmth. fires are made in the open or behind makeshift windbreaks. there is no trade in fuelwood. visitors to the rnp are expected to supply their own energy requirements. methods deadwood availability the availability of both attached (still on tree) and detached (on ground) deadwood was assessed along 12 line transects in the riparian fringe parallel to the gariep river and radiating away from eight herder stockposts. each line transect consisted of 20 points 50 m apart (hence 1 km in total length), with the first point being 50 m from the stock post. at each point issn 0075-6458 3 koedoe 46/2 (2003) shackleton.qxd 2005/12/09 11:24 page 3 the nearest tree was examined and records made of (i) the species, and (ii) a visual estimation of the proportion dead. the proportion was first estimated in terms of one of eight classes using the scale of walker (1976). once the class was established, which provided upper and lower limits for the visual estimation thereby reducing variability, a final percentage dead was estimated. in addition, a five by five metre quadrat was marked out, to the left and front (in the direction of the transect) of the sample point. a visual estimation of the percentage of detached deadwood cover (branches on the ground) was made in these quadrats. detached deadwood included that from the trees in the riparian fringe, as well as driftwood debris deposited by the gariep river. sampling was restricted to the riparian fringe since tree biomass is relatively limited throughout other areas of the rnp. this is well appreciated by the resident pastoralists, who stated that even when they moved away from the river in winter, fuelwood is transported from the riparian fringe by motor vehicle (hendricks pers. comm.). in terms of fuelwood species preferences expressed by the herders, the majority are found only in the riparian fringe, and not inland in rnp, other than zygophyllum prismatocarpum. species preferences four pastoralist herders within the rnp were opportunistically interviewed to determine the species preferences with respect to fuelwood. a semi-structured interview was used to gather data on preferred species, species used and wood collection methods. the interview approach had no fixed wording or ordering of questions (case 1990). this allowed for flexibility in that it is possible to seek clarification and elaboration on some of the answers given (martin 1995). three fuelwood piles were inspected and the species identity of each stick/log was recorded to provide quantitative data regarding preferences. data analyses woody plant species composition along the riparian fringe was determined as the percentage occurrence (as the proportion of total stems sampled) of different species along the transects. the relative proportion of species present was compared to the relative proportion of species found in the herders' woodpiles. a ratio of availability to use was then used to indicate selection or avoidance of specific species. if the ratio was greater than one for a specific species, then active selection was indicated, if less than one, then avoidance, and a ratio of approximately one koedoe 46/2 (2003) 4 issn 0075-6458 fig. 1. mean percentage deadwood per tree in relation to distance from stockpost (n = 12 transects). fig. 2. mean percentage deadwood cover on ground in relation to distance from stockpost (n = 12 transects). indicated use was random for that species (shackleton et al. 1994). the influence of distance from stockposts on attached and detached deadwood availability was examined via linear regression. differences in the proportion of attached deadwood per species were tested using a kruskal-wallis anova after arcsine transformation of the original percentage data. only those species constituting 5 % or more of the frequency data set were included as the occurrence of species with lower frequency was superfluous because of their low sample number. results there was no relationship (r² = 0.02; n = 20; p>0.05) between the percentage of attached deadwood on the tree, and the distance from the stockpost (fig. 1). similarly, there was no relationship (r² = 0.0003; n = 20; p>0.05) between the percentage cover of detached deadwood on the ground and the distance from the stockpost (fig. 2). shackleton.qxd 2005/12/09 11:24 page 4 damage to trees (cutting or breakage) was observed on only six of the sampled trees, where a branch had been bent back. it was not possible to tell the cause of the break, whether human or perhaps baboons. species selection ziziphus mucronata and e. pseudobenus were the preferred species for fuelwood collection, dominating with 45% and 34% respectively (table 1). overall, eleven tree species were encountered during the survey, but several were represented by only a single individual. those representing greater than 5 % of the stems sampled are presented in table 2. euclea pseudobenus and t. usneoides were the dominant species in the area, comprising 31 % and 32 %, respectively. there was strong selection for z. mucronata and only random selection for e. pseudobenus (table 3). ziziphus mucronata was also the principal fuelwood species. t. usneoides and prosopis sp. were common species in the riparian zone, but were absent from woodpiles, indicating that they were actively avoided. the interview responses confirmed that harvesters did not make use of these two species. proportion of deadwood per species there was a significant difference (s = 43.2; p< 0.001) between species with respect to the mean proportion of the stem that was dead. on average, more than one quarter of issn 0075-6458 5 koedoe 46/2 (2003) table 3 preference ratios species % in wood % in ratio preference piles environment ziziphus mucronata 45 20 2.25 preferred euclea pseudobenus 35 31 1.13 random tamarix usneoides 0 32 0 avoided prosopis sp. 0 6 0 avoided table 1 composition of fuelwood piles species percentage ziziphus mucronata 45 euclea pseudobenus 34 unknown # 21 # the unknown species was one unidentifiable species, and not a combination of more than one species. table 2 species composition along the riparian fringe species percentage tamarix usneoides 32 euclea pseudobenus 31 ziziphus mucronata 20 prosopis sp. 6 all others combined (each less than 5 %) (acacia karroo, maytenus linearis, nicotiana glauca, rhus pendulina, plus unidentified species) 11 z. mucronata trees were dead, which was more than double the amount of deadwood of the next closest species, i.e. t. usneoides (table 4). most of the prosopis trees had no deadwood at all. across all sites and species, the mean percentage dead per tree was approximately 15 %. additionally, detached shackleton.qxd 2005/12/09 11:24 page 5 deadwood covered just less than 9 % of ground area, average across all plots and transects. table 4 mean percentage (± se) deadwood on each tree species (unlike letters are significantly different) species mean % deadwood per stem ziziphus mucronata 27.8 ± 5.4 a euclea pseudobenus 8.8 ± 2.4 b tamarix usneoides 12.2 ± 2.9 b prosopis sp. 0.3 ± 0.2 b discussion there is some concern among conservationists regarding the threat that the local pastoralists pose to conservation in the rnp (h. hendricks pers. comm.). the pastoralists use a variety of natural resources for their livelihood, from grazing their herds, to the use of wood for construction and fuel, use of medicinal plants, and fishing. in terms of fuelwood extraction, we have demonstrated that, despite the rudimentary nature of the study, all the variables measured indicated that there seems to be little cause for concern over the current fuelwood extraction activities of pastoralists within the rnp. there was no evidence of cutting of branches or deadwood. the availability of both attached and detached deadwood was not depleted close to human settlement (stockposts), relative to further away. there was still abundant deadwood, even of preferred species. the most preferred species (ziziphus mucronata) had the greatest mean proportion of deadwood. moreover, of the two most common species in the riparian zone, one was not a favoured species (tamarix unsneoides) and the other (euclea pseudobenus) was not actively selected for, as indexed through the ratio of the proportions in the environment and fuelwood piles. the four herders interviewed stated that they only used deadwood, and that there was plenty. they did not have to walk far to collect deadwood for their daily needs. none of the four interviewees felt that there had been any decline in the availability of deadwood over the last five to ten years. they rarely used driftwood washed down the river, although there was an abundance of it. thus, across all these indices, there was no sign of depletion of the deadwood resource, even on favoured species, or degradation of the productive capacity for deadwood. this finding conforms to other recent work in arid environments questioning the notion and severity of degradation by low density populations of pastoralists (reid & ellis 1995; sullivan 1999; ward et al. 2000; mcneely 2003). it also mirrors the rationale and findings of the study by abbot & homewood (1999) of fuelwood use in lake malawi national park. there too, conservation management considered domestic fuelwood use by people residing in the park to be the primary cause of miombo woodland decline within the park. but, quantitative analysis of supply and demand demonstrated that domestic fuelwood demand was well below the sustainable annual production of deadwood in lake malawi national park, and demand for wood for commercially driven fish smoking enterprises was the primary cause. whilst our study has shown there to be little cause for concern at present, it is possible this may change in the future, although unlikely in the near term because of the low and controlled number of residents in the park and the seemingly high abundance of deadwood relative to current demand. however, since the residents in the park practice a nomadic lifestyle it is likely that fuelwood will remain their primary energy source for years to come. therefore, it may be desirable for conservation management to engage in a process of defining, with the residents, thresholds of potential concern and indicators of decline, which could be monitored at an appropriate frequency. a more detailed study of deadwood production and use may also be desirable. koedoe 46/2 (2003) 6 issn 0075-6458 shackleton.qxd 2005/12/09 11:24 page 6 overall, only 11 tree species were recorded in the line transects, providing only limited choice to the fuelwood collectors. of these, two (e. pseudobenus and z. mucronata) contributed the bulk of the wood in the fuelwood piles, as well as mentioned during the interviews. both of these have relatively dense wood, which is ideal for cooking fires (tietema et al. 1991; abbot & lowore 1999; kataki & konwer 2002), since it has a higher energy content, burns slowly but with even heat, and makes good coals. in comparison, although t. usneoides was common and had more deadwood per stem than e. pseudobenus, it was actively avoided, which may be a consequence of its wood being light and brittle and therefore unsuitable for cooking fires. interestingly, goel & behl (2001), in a study assessing the relative potential of various hardwood species for fuelwood production, found prosopis juliflora to be the most promising species in terms of biomass productivity and fuel value index. additionally prosopis spp. are widely used in other parts of the northern cape province as fuelwood (c. fabricius pers. comm.) yet, in rnp, prosopis sp. was not used, as measured in the fuelwood bundles and supported by the interviews. this is possibly related to the negligible amounts of deadwood found on prosopis sp. in rnp (< 1 %). deadwood is invariably favoured over live wood as a fuelwood resource where a choice is possible, i.e. where deadwood has not been totally harvested, which was the case in rnp. the difference in deadwood availability between species was noteworthy, and little studied in the literature pertaining to fuelwood. typically, the fuelwood production capacity of different species has been tacitly assumed as uniform, or simply to mirror the growth rates of the live tree. previously, estimates of production of deadwood have been area based rather than tree based (shackleton 1998; abbot & homewood 1999). in comparison, mudekwe (1997) examined species differences in both standing crop and production of deadwood in the limpopo lowveld and found marked differences between tree size classes and the five species investigated. this needs to be examined at other sites. additionally, the productive capacity of fuelwood in rnp has a patchy distribution, being largely limited to disjunct strips along the riparian fringe. this means that coarse-scale studies and models would underestimate the availability of fuelwood, which must also influence subjective impressions about the sustainability of fuelwood use. the study focussed solely on the availability of deadwood. we did not examine aspects relating to its role in ecosystem function, such as nutrient cycling or provision of habitats for microfauna and flora, although the importance of such a role has been questioned in southern african savanna ecosystems where collection is by hand (williams & shackleton 2002). nonetheless, the clear abundance of deadwood with rnp under the current harvesting arrangements, allows us to hypothesize that impacts relating to ecosystem function would be well within acceptable limits, but this needs to be tested. this is so even from a biodiversity conservation perspective in light of the need to maintain a cooperative and beneficial relationship with the richtersveld people, from whom the land is leased for conservation purposes. in summary, this study could not substantiate verbal concerns relating to the possible overexploitation of deadwood within rnp. it appears that current extraction levels are low relative to the availability, and hence can be accommodated within the broader conditions pertaining to the lease arrangements and management of this key biodiversity area. acknowledgements we gratefully acknowledge the assistance of howie hendricks in orientating us within rnp and facilitating interviews with pastoralists. this study formed part of, and was partially funded by, the south african component of the southern african millennium assessment, which focuses on ecosystem goods and services in the gariep basin. useful comments were received from christo fabricius and three anonymous reviewers on earlier drafts of this paper, for which we are appreciative. issn 0075-6458 7 koedoe 46/2 (2003) shackleton.qxd 2005/12/09 11:24 page 7 references abbot, j.i.o. & k. homewood. 1999. a history of change: causes of woodland decline in a protected area in malawi. journal of applied ecology 36: 422-433. abbot, p.g. & j.d. lowore. 1999. characteristics and management of some indigenous firewood species in malawi. forest ecology and management 119: 111-121. banks, d.i. griffin, n.j., shackleton, c.m., shackleton, s.e. & j.m. mavrandonis. 1996. wood supply and demand around two rural settlements in a semi-arid savanna, south africa. biomass & bioenergy 11: 319-331. benjaminsen, t.a. 1997. is there a fuelwood crisis in mali? geojournal 43: 163-174. case, d.d. 1990. the community's toolbox: the idea, methods and tools for participatory assessment, monitoring and evaluation in community forestry. rome: food and agricultural organization (fao). dahlberg, a.c. 2000. interpretations of environmental change and diversity: a critical approach to indications of degradation: the case of kalakamate, northeast botswana. land degradation & development 11: 549-562. department of minerals and energy affairs, 1996. energy in south africa. pretoria: government printer. gielink, m.i. 1991. energy in southern and eastern sub-saharan africa. journal of energy research & development in south africa 2: 3-11. goel, v.l. & h.m. behl. 2001.genetic selection and improvement of hard wood tree species for fuelwood production on sodic soil with particular reference to prosopis juliflora. biomass & bioenergy 20: 9-15. grundy, i.m., campbell, b.m., balebereho, s., cunliffe, r., tafangenyasha, c., fergusson, r. & d. parry. 1993. availability and use of trees in mutanda resettlement area, zimbabwe. forest ecology & management 56: 243-266. hall, d.o. 1991. biomass energy. energy policy 19: 711-737. kataki, r. & d. konwer. 2002. fuelwood characteristics of indigenous tree species of north-east india. biomass & bioenergy 22: 433-437. leach, g. & r. mearns. 1989. beyond the fuelwood crisis: people, land and trees in africa. london: earthscan. leach, m. & r. mearns. 1996. the lie of the land: challenging received wisdom on the african environment. oxford: currey & heinemann. luoga, e.j., witkowski, e.t.f. & k. balkwill. 2001. harvested and standing wood stocks in protected and communal miombo woodlands of eastern tanzania. forest ecology & management 164: 15-30. martin, g. 1995. ethnobotany: a methods manual. london: chapmans & hall. mcneely, j.a. 2003. biodiversity in arid regions: values and perceptions. journal of arid environments 54: 61-70. mudekwe, j. 1997. assessing the effect of tree size on the production of harvesting deadwood for fuelwood. m.sc thesis, university of the witwatersrand, johannesburg. nagothu, u.s. 2001. fuelwood and fodder extraction and deforestation: mainstream views in india discussed on the basis of data from the semi-arid region of rajasthan. geoforum 32: 319-332. pandey, d. 2002. fuelwood studies in india: myth and reality. bogor: cifor. reid, r.s. & j.e. ellis. 1995. impacts of pastoralists on woodlands in south turkana, kenya: livestock mediated tree recruitment. ecological applications 5: 978-992. shackleton, c.m., griffin, n.j., banks, d.i., mavrandonis, j.m. & s.e. shackleton. 1994. community structure and species composition along a disturbance gradient in a communally managed south african savanna. vegetatio 115: 157-167. shackleton, c.m. 1998. annual production of harvestable deadwood in semi-arid savannas, south africa. forest ecology & management 112: 139-144. sullivan, s. 1999. the impacts of people and livestock on topographically diverse open woodand shrub-lands of arid north-west namibia. global ecology & biogeography 8: 257-277. tietema, t. dithogo, m. tibone, c. & n. mathalaza. 1991. characteristics of eight firewood species of botswana. biomass & bioenergy 1: 41-46. van der walt, p.t.1992. the richtersveld—a desert paradise. airlink 2 (1): 4-5. walker, b.h. 1976. an approach to the monitoring of changes in the composition and utilization of woodland and savanna vegetation. south african journal of wildlife research 6: 1-32. ward, d., b.t. ngairorue, a. apollus & h. tjiveze. 2000. perceptions and realities of land degradation in arid otjimbingwe, namibia. journal of arid environments 45: 337-356. williams, a. & c.m. shackleton. 2002. fuelwood use in south africa: where to in the 21st century? southern african forestry journal 196: 1-7. williamson, g. 2000. notes on the vegetation and climate of the richtersveld national park. aloe 31: 83-87. koedoe 46/2 (2003) 8 issn 0075-6458 shackleton.qxd 2005/12/09 11:24 page 8 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <feff004f007000740069006f006e007300200070006f0075007200200063007200e900650072002000640065007300200064006f00630075006d0065006e00740073002000500044004600200064006f007400e900730020006400270075006e00650020007200e90073006f006c007500740069006f006e002000e9006c0065007600e9006500200070006f0075007200200075006e00650020007100750061006c0069007400e90020006400270069006d007000720065007300730069006f006e00200070007200e9007000720065007300730065002e0020005500740069006c006900730065007a0020004100630072006f0062006100740020006f00750020005200650061006400650072002c002000760065007200730069006f006e00200035002e00300020006f007500200075006c007400e9007200690065007500720065002c00200070006f007500720020006c006500730020006f00750076007200690072002e0020004c00270069006e0063006f00720070006f0072006100740069006f006e002000640065007300200070006f006c0069006300650073002000650073007400200072006500710075006900730065002e> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 van rooyen.qxd impact of harvesting and fire on phragmites australis reed quality in tembe elephant park, maputaland m.w. van rooyen, c.a. tosh, n. van rooyen, w.s. matthews and m.j.s. kellerman van rooyen, m.w., c.a. tosh, n. van rooyen, w.s. matthews and m.j.s. kellerman. 2004. impact of harvesting and fire on phragmites australis reed quality in tembe elephant park, maputaland. koedoe 47(1): 31–40. pretoria. issn 0075-6458. in maputaland, south africa, the common reed (phragmites australis) is used extensively for hut building, fencing, craftwork and thatching. as a result of over-harvesting most reed beds in communal areas have been degraded and are no longer producing reeds of the desired quality. at present the most productive reed beds are all found in conservation areas. the kwamsomi area of the muzi swamp in the tembe elephant park has been allocated to the sibonisweni community for harvesting purposes. this community has recently requested ezemvelo kwazulu-natal wildlife for additional areas for harvesting on the grounds that the current site was no longer yielding reeds of suitable quality. the main objective of this study was therefore to determine whether there was a decline in reed quality in the kwamsomi harvested area. the results of this study suggest that harvested areas contained more thin, short reeds than unharvested areas. fire can be used to increase reed diameter in harvested areas, but will not significantly affect reed height. ideally, reeds should only be harvested after the active growth period, when most of the nutrient reserves have been translocated to the rhizomes and the buds are still dormant. to improve reed quality a three-year rotational harvesting programme should be implemented to allow the reeds to recover fully before being harvested again. keywords: common reed, fire, harvesting, kwazulu-natal, muzi swamp, phragmites australis, sustainable utilisation. m.w. van rooyen, c.a. tosh, n. van rooyen and m.j.s. kellerman, department of botany, university of pretoria, pretoria, 0002 republic of south africa; w.s. matthews, ezemvelo kwazulu-natal wildlife, p o box 356, kwangwanase, 3973 republic of south africa. issn 0075-6458 31 koedoe 47/1 (2004) introduction thousands of rural south africans in kwazulu-natal depend on biological resources for their day-to-day survival. access to this natural capital provides a crucial contribution to their livelihood and is a buffer against poverty and an opportunity for self-employment (wynberg 2002). the common reed, phragmites australis (cav.) trin. ex steud., is a widespread, morphologically highly-variable key wetland species that provides many essential ecosystem services. it is a major component of freshwater bodies in temperate regions throughout the world and often forms large monospecific stands at the interface between land and water (ingram et al. 1980; clevering 1999; clevering & lissner 1999; karunaratne & asaeda 2000; asaeda & karunaratne 2000). large-scale decline of phragmites australis stands in european freshwater bodies has stimulated intensive research on this species and a large body of literature has been accumulated over the past decades. the effects of environmental factors such as chemical composition of the soil and water, altitude, temperature stress and mineral uptake (weisner 1996; klimes et al. 1999; cízková & lukavska 1999; cízková et al. 2001b; batty et al. 2002) and management practices such as mowing, cutting and burning on biomass van rooyen.qxd 2004/04/15 03:12 page 31 production of phragmites australis have been well documented for european conditions (haslam 1970; van der toorn & mook 1982; mook & van der toorn 1982; ostendorp 1995, 1999). autecological studies on phragmites australis have not been conducted in south african conditions. several studies have reported on changes in the distribution and extent of phragmites australis stands in south africa (e.g., weisser 1981; kotschy et al. 2000). two studies have investigated the impact of reed harvesting in the fuyeni reed bed in kwazulu-natal (frankland 1982; mckean 2001). in southern africa, the common reed is used extensively for non-food commodities. the thick hollow stems are widely used for building walls, fences and bomas (van wyk & gericke 2000). stems are woven into traditional sitting mats and winnowing baskets and also used for arrow shafts, tobacco pipestems, flutes and parts of musical instruments. the common reed is also used for traditional medicine (van wyk & gericke 2000). in maputaland, northern kwazulu-natal, reeds are in demand for hut building, fencing, craftwork and thatching (begg 1988). however, due to over-harvesting, most reed beds in communal areas have been degraded and are no longer producing reeds of the desired quality. at present, the most productive reed beds are all situated in conservation areas. to prevent the degradation of reed beds in conservation areas there is an urgent need to establish levels of sustainable utilisation for this resource. when the tembe elephant park was proclaimed, part of the agreement with the neighbouring communities was that they would be allowed to harvest particular renewable natural resources inside the park at sustainable levels. one particular area of the muzi swamp was specifically allocated to the sibonisweni community to harvest phragmites australis for non-commercial use. the harvesters of this community, enter the park at kwamsomi gate and may harvest reeds for approximately 1.5 km northwards along the swamp. when the park was fenced in 1983 the fenceline bisected a monoculture of phragmites australis across the muzi swamp at kwamsomi. the reed beds on the outside of the park have since been almost destroyed leading to increased pressure for additional collection inside the park. other areas of the muzi swamp have been set aside for harvesting by other communities and access to those areas is obtained via other gates. the sibonisweni community have recently requested ezemvelo kwazulu-natal wildlife for additional areas for harvesting on the grounds that the current site was no longer yielding reeds of suitable quality. however, conservation authorities are concerned that this may lead to non-sustainable levels of harvest because the reeds are being sold commercially and not merely for local use as stipulated by the original agreement. the primary aim of this study was to determine whether there was a decline in reed quality in the kwamsomi harvested areas. however, because accidental fires had swept through sections of both the harvested and non-harvested areas of the muzi swamp in the previous winter, an opportunity existed to investigate the use of fire to improve reed quality. the second aim of the study was therefore to determine the effect of fire on reed quality. based on these results suggestions for sustainable harvesting are made. study area the tembe elephant park is located in northern kwazulu-natal with its northern boundary bordering mozambique. the park covers approximately 30 000 ha and was proclaimed in 1983 as a haven for the maputaland elephant population in an effort to reduce the human-elephant conflict. the park is situated in the core area of the maputaland centre of endemism (van wyk & smith 2001) and the conservation of many rare and endemic plant and animal species is one of the park's main objectives. koedoe 47/1 (2004) 32 issn 0075-6458 van rooyen.qxd 2004/04/15 03:12 page 32 the muzi swamp covers about 560 ha in the eastern part of the tembe elephant park. this is the most important wetland system in the park and is the only permanent water supply in the area. it stretches for 25 km from south to north, running in a slightly easterly direction. for most of its length it is covered by the phragmites australis community (matthews et al. 2001). the climate of the area can be described as subtropical, with hot summers and cool to warm winters (schulze 1982). rainfall is mainly in summer, but winter showers are known to occur. mean annual rainfall at sihangwane (in tembe elephant park) is 722 mm, but ranges from 245 mm to a maximum of 2105 mm per annum (matthews et al. 2001). the period preceding the harvest was exceptionally wet and 1541 mm of rain fell from august 1999 to june 2000. material and methods the effects of two factors, viz. utilisation and burning on reed quality were investigated. reed quality was determined in terms of reed diameter, reed height and reed density. four sites, differing in utilisation/burning status were selected. these sites were: (a) unutilised, unburnt; (b) unutilised, burnt; (c) utilised, unburnt and (d) utilised, burnt. burnt and unburnt sites within a utilisation zone were adjacent to each other to minimise possible effects of other environmental factors. the utilised sites were approximately 500 m from kwamsomi gate and represented an area under relatively high utilisation pressure. the unutilised sites were approximately 1.8 km from kwamsomi gate in an area of the swamp that may not be harvested by the neighbouring communities. at each site eight randomly selected samples of 1 m2 were harvested in july. all reeds within the 1 m2 sample area were cut at water level and the height and diameter of each reed measured. total reed height was obtained by correcting for water depth. frequency distributions of height and diameter classes were calculated for each site. data were analysed with statistica 6 (statsoft inc, tulsa oklahoma) by performing a factorial anova and scheffe's post hoc test to determine statistically significant differences between treatments. the sigissn 0075-6458 33 koedoe 47/1 (2004) fig. 1. annual reed harvest from the muzi swamp from 1993 to 1999 (kyle 2000). nificant differences (p < 0.05) are indicated in the figures by different superscripts above the bars. results harvesting pattern annual reed harvest from the muzi swamp increased from 1993 to a maximum off-take in 1996, whereafter reed harvesting decreased (fig. 1) (kyle 2000). until 1996 no harvesting quotas were imposed by conservation authorities in the tembe elephant park. the authorities exercised control over the harvesting only by delimiting the area that may be harvested and controlling the number of reed cutters entering the park at kwamsomi gate per day. the number of bundles harvested during that period was therefore, determined largely by the demand for reeds. monthly reed off-take from the muzi swamp in 1999 illustrates that approximately 65% of all bundles were harvested from march to august (fig. 2). harvesters in the tembe elephant park do not select reeds of only a particular thickness and height to harvest. usually an area is selected and all reeds within that area are harvested (fig. 3). the cut reeds are sorted into bundles containing reeds of even thickvan rooyen.qxd 2004/04/15 03:12 page 33 ness and length. the taller and thicker reeds are the most prized and of the highest quality. for this study reeds of >8 mm in diameter and >2.1 m in height indicate the availability of the most desirable reeds. in his study of phragmites australis harvesting in the fuyeni reed beds mckean (2001) regarded usable reeds in the fuyeni reed bed as >10 mm in diameter and >2.5 m tall. because of the apparent difference in harvesting pattern in the fuyeni reed bed and the muzi swamp the definition used in this study is different to that used by mckean (2001). koedoe 47/1 (2004) 34 issn 0075-6458 fig. 4. phragmites australis reed quality under different utilisation and burning regimes in the muzi swamp, tembe elephant park as determined by harvesting trials in 2000: (a) mean reed density per m²; (b) mean reed diameter; and (c) mean reed height. bars with different superscripts are significantly different (p < 0.05). a b c fig. 2. monthly reed harvest from the muzi swamp in 1999 (kyle 2000). fig. 3. aerial view of a harvested area in the muzi swamp. d en si ty ( re ed s / s qu ar e m et re ) van rooyen.qxd 2004/04/15 03:12 page 34 utilisation density was not significantly affected by utilisation (p = 0.234, table 1) within a burning treatment (fig. 4a). although reed density was higher when the reeds were utilised than when unutilised in the absence of fire, this difference was not statistically significant. unutilised sites produced significantly thicker (p < 0.001, table 1) reeds both in the presence and absence of fire (fig. 4b). the frequency distribution of reed diameter classes (fig. 5) gives a more detailed indication of the effects of utilisation and fire. at the unburnt sites, almost no reeds thicker than 8 mm were produced in the utilised area (2.7 % of all reeds, fig. 5c), whereas 13.0 % of the reeds produced in the unutilised area (fig. 5d) were > 8 mm in diameter. at the burnt sites, utilisation reduced the percentage of reeds > 8 mm thick from 23.9 % at the unutilised site (fig. 5b) to 15.0 % in the utilised site (fig. 5a). in utilised areas no reeds with a diameter > 14 mm were recorded, whereas reeds up to 18 mm in diameter were recorded in unutilised areas. reeds were not only thicker but also significantly taller (p < 0.001, table 1) at unutilised sites than at utilised sites (fig. 4c). in the absence of fire, utilisation reduced the percentage of reeds taller than 2.1 m from 33.0 % (fig. 6d) to only 7.5 % in the utilised area (fig. 6c). after fire, only 11.7 % of the reeds were > 2.1 m tall at the utilised sites (fig. 6a) as against 39.7 % at the unutilised sites (fig. 6b). fire fire significantly reduced mean reed density (p < 0.001, table 1) at both the harvested and non-harvested sites (fig. 4a), but significantly increased mean reed diameter (p < 0.001, table 1) at both harvested and non-harvested sites (fig. 4b). in the unutilised zone stem diameter classes > 8 mm constituted 23.9 % of all reeds after fire (fig. 5b) as against 13.0 % without fire (fig. 5d). however, because of the two-fold difference in reed density, the density of preferred reeds is approximately 15 reeds/m² in both cases. in the utilised area, fire had a greater effect and only 2.7 % of the reeds in the unburnt area (fig. 5c) were > 8 mm thick, whereas 15.0 % were > 8 mm thick after fire (fig. 5a). even taking the difference in density into account, the number of thick reeds is still more after fire, with the burnt area yielding approximately 10 reeds/m² in these size classes as opposed to only 4 reeds/m² in the unburnt area. fire did not significantly affect mean reed height (p = 0.073, table 1) in either the utilised or unutilised zone (fig. 4c). the percentage contribution of reeds > 2.1 m tall were more or less equal in the unutilised zone whether the area had been burnt or not, with 39.7 % of all reeds in the unutilised burnt zone (fig. 6b) as against 33.0 % in the unutilised, unburnt zone (fig. 6d). however, if the difference in density is accounted for, the unburnt treatment will yield more tall reeds to harvest (38 reeds/m² as against 25 reeds/m²). in the utilised area only 7.5 % issn 0075-6458 35 koedoe 47/1 (2004) table 1 results of factorial analyses of variance (anovas) examining the effects of fire and harvesting on reed density, reed diameter and reed height in the muzi swamp, tembe elephant park factor p-values reed density reed diameter reed height fire <0.001 <0.001 0.073 utilisation 0.234 <0.001 <0.001 fire x utilisation interaction 0.410 0.673 0.297 van rooyen.qxd 2004/04/15 03:12 page 35 of the unburnt reeds were > 2.1 m tall (fig. 6c) as opposed to 11.7 % of the reeds produced after fire (fig. 6a). however, after accounting for the differences in density, there will be slightly more tall reeds to harvest in the unburnt area (11 reeds/m²) than in the burnt area (8 reeds/m²). discussion in kwazulu-natal the degradation of wetlands has been extensive and future threats such as agricultural and rural development and global climate change remain high (kotze & o’connor 2000). over-harvesting of reeds has led to the degradation of reed beds and could eventually result in their total loss. to maintain the integrity of these systems and their inherent value to their users, it is crucial to establish levels of sustainable reed harvesting. in ecological terms, sustainable harvesting is defined as harvesting that has no long-term detrimental effect on the reproduction and regeneration of the population being harvested compared with similar non-harvested populations (hall & bawa 1993). natural degeneration of phragmites australis stands is known to occur and has often been associated with the accumulation of its own organic matter (van der putten et al. 1997; clevering 1999; lenssen et al. 2000). in waterlogged soil, microbial decomposition occurs under reduced conditions and koedoe 47/1 (2004) 36 issn 0075-6458 fig. 5. frequency distribution of different diameter size classes under different utilisation regimes in the muzi swamp, tembe elephant park as determined by harvesting trials in 2000. (a) burnt, utilised area; (b) burnt, unutilised area; (c) unburnt, utilised area; and (d) unburnt, unutilised area. (a) (b) (c) (d) van rooyen.qxd 2004/04/15 03:12 page 36 under these conditions of oxygen deficiency organic acids are formed that might be involved in the phytotoxic effect (cízková et al. 1999). in this regard the input of discarded leaf litter to the reed beds by the harvesters might be contributing to the degradation of the swamp in the harvested areas. the timing of reed harvesting is of critical importance. phragmites australis is one of many wetland species that produces an extensive perennial rhizome system, which is able to tolerate long-term oxygen deficiency in flooded soils because of its internal aeration system (cízková & bauer 1998; soukup et al. 2000, 2002; cízková et al. 2001a; strand & weisner 2002). these rhizomes play a major role in the storage of carbohydrate reserves that are the only source of carbohydrates during winter after shoots have died off. buds are produced throughout the year but remain dormant near the soil surface for most of the time. the main burst of aerial annual shoot growth occurs in spring when developing shoots are supplied with stored carbohydrates from the rhizomes. reeds mature during summer and shoot dry matter accumulation stops abruptly at the onset of flowering, which coincides with the start of senescence. at the end of the active growing stage, the accumulated dry matter is translocated to the rhizomes. ideally, reeds should be harvested after the active growth period when most of the reserves have been translocated to the rhizomes, but the buds are still dormant or shoots are still issn 0075-6458 37 koedoe 47/1 (2004) fig. 6. frequency distribution of different height classes under different utilisation regimes in the muzi swamp, tembe elephant park as determined by harvesting trials in 2000. (a) burnt, utilised area; (b) burnt, unutilised area; (c) unburnt, utilised area; and (d) unburnt, unutilised area. van rooyen.qxd 2004/04/15 03:12 page 37 under the soil surface (mook & van der toorn 1982). most of the harvesting done in the muzi swamp (65 %) is done from may to august and therefore falls within this period. however, summer harvesting still accounts for 35 % of the reed off-take, and especially when done year after year, this will deplete rhizome reserves, and is probably a major factor in the decline in reed quality in the kwamsomi area. several previous studies have indicated that when shoots are damaged, either by harvesting, mowing or burning, the density of the replacement crop is increased (frankland 1982; van der toorn & mook 1982; ostendorp 1995, 1999). however, this contrasts with the findings of the present study where burning significantly decreased reed density and harvesting had no significant effect on density. in this study, we found that the response of reeds to harvesting corresponds closely with the findings of mckean (2001) who investigated the productivity and sustainable use of phragmites australis in the fuyeni reed bed in the hluhluwe-umfolozi park, also in kwazulu-natal. in both these studies, harvested treatments showed higher reed densities than non-harvested treatments, but the differences were not statistically significant. previous studies have reported that after shoot damage by burning, cutting or mowing, replacement shoots are shorter and thinner (van der toorn & mook 1982; ostendorp 1995, 1999; mckean 2001). the results of the present study confirmed that harvesting significantly reduced mean reed diameter and reed height, in both the burnt and unburnt areas. consequently, thick and tall reeds are in short supply in the harvested areas. however, fire did not have the same effect. fire significantly increased mean reed diameter while having no effect on mean reed height within a utilisation regime. in utilised areas, fire during the dormant season can, therefore, have a positive effect on reed diameter but not on reed height. in spite of the fact that reed density is reduced after fire, the density of thick reeds is higher after fire in the utilised zone. in unutilised areas, these differences are not as marked or can even be absent and because of the density difference more thick reeds are still found in unburnt areas. in europe, burning or mowing in winter is sometimes applied as countermeasure against the die-back of reeds (ostendorp 1995, 1999). although early season damage retards leaf and shoot growth, recovery is rapid and the maximum biomass production is not significantly lowered (mook & van der toorn 1982; mckean 2001). however, ostendorp (1995, 1999) warns against the long-term negative effects of burning and mowing as management practices. winter harvesting and burning lead to the dominance of weaker and thinner secondary shoots and in conditions where mechanical resistance to waves and washes is important, these management practices are not recommended (ostendorp 1995, 1999). management proposals utilisation should be confined to the winter months (dormant season) when the damage to the reeds is minimal, because nutrient reserves have already been translocated to the rhizomes. likewise, should fire be necessary, it should only be implemented during winter or early spring to minimise negative effects. continuous harvesting leads to the production of more short, thin reeds of limited value and could also deplete the carbohydrate reserves in the rhizomes. a system of rotational harvesting should be implemented even if this requires that an additional area of the muzi swamp should be set aside for harvesting. mckean (2001) investigated the effects of annual and biennial harvesting in the fuyeni reed beds and concluded that it would be necessary to restrict harvesting of a given area to every second or preferably every third year to allow the reed bed to recover a full complement of large reeds. harvesting on a tri-annual basis is therefore suggested for the muzi swamp. koedoe 47/1 (2004) 38 issn 0075-6458 van rooyen.qxd 2004/04/15 03:12 page 38 in unutilised areas, fire can be used to remove moribund material and to yield a larger proportion of tall and thick reeds. this study suggests that though fire will lead to more thick reeds in utilised areas these reeds will be shorter (although the latter is not statistically significant). the long-term effects of fire were not investigated in this study and it is therefore not recommended to burn on an annual basis. conclusions the allegation of the sibonisweni community that the quality of reeds in the harvested area has declined is confirmed. compared with the unharvested area more short, thin reeds are being produced in the harvested area. fire has the potential to increase reed diameter but does not significantly affect reed height. to improve reed quality harvesting should be restricted to the dormant period when damage to the reeds is minimal. a three-year rotational harvesting programme is suggested as this should allow the reeds to recover fully, before being harvested again. such a programme might, however, require an additional area to be set aside for harvesting. acknowledgements the authors would like to acknowledge the assistance of ezemvelo kwazulu-natal wildlife. the material is based on research supported by the national research foundation (gun 2053522). references asaeda, t. & s. karunaratne. 2000. dynamic modelling of the growth of phragmites australis: model description. aquatic botany 67: 301-318. batty, l.c., a.j.m. baker & b.d. wheeler. 2002. aluminium and phosphate uptake by phragmites australis: the role of fe, mn and al root plaques. annals of botany 89: 443-449. begg, g.w. 1988. the wetlands of natal. natal town and regional planning commission, pietermaritzburg. cízková, h. & v. bauer. 1998. rhizome respiration of phragmites australis: effect of rhizome age, temperature, and nutrient status of the habitat. aquatic botany 61: 239-253. cízková, h., h. brix, j. kopecky & j. lukavská. 1999. organic acids in the sediments of wetland dominated by phragmites australis: evidence of phytotoxic concentrations. aquatic botany 64: 303-315. cízková, h. & j. lukavská. 1999 rhizome age structure of three populations of phragmites australis (cav.) trin. ex steud.: biomass and mineral nutrient concentrations. folia geobotanica 34: 209-220. cízková, h., v. istvánovics, v. bauer & l. balázs. 2001a. low levels of reserve carbohydrates in reed (phragmites australis) stands of kis-balatow, hungary. aquatic botany 69: 209216. cízková, h., l. pechar, s. husák, j. kvet, v. bauer, j. radová & k. edwards. 2001b. chemical characteristics of soils and pore waters on three wetland sites dominated by phragmites australis: relation to vegetation composition and reed performance. aquatic botany 69: 235-249. clevering, o.a. 1999. betweenand within-population differences in phragmites australis. 1. the effects of nutrients on seedling growth. oecologia 121: 447-457. clevering, o.a. & j. lissner. 1999. taxonomy, chromosome numbers, clonal diversity and population dynamics of phragmites australis. aquatic botany 64: 185-208. frankland, c. 1982. the impact of annual reed cutting on the ecology of the fuyeni reedbed. honours dissertation, department of biological sciences, university of natal, durban. hall, p. & k. bawa. 1993. methods to assess the impact of extraction of non-timber tropical forest products on plant populations. economic botany 47: 234-247. haslam, s.m. 1970 the development of the annual population in phragmites communis trin. annals of botany 34: 571-591. ingram, h.a.p., a.m. barclay, a.m. cougar, j.g. glover, b.m. lynch & j.i. sprent. 1980. phragmites performance in reedbeds of the tay estuary. proceedings of the royal society of edinburgh 78b: 89-107. karunaratne, s. & t. asaeda. 2000. verification of a mathematical growth model of phragmites australis using field data from two scottish lochs. folia geobotanica 35: 419-432. klimeš, l., j. klimešová & h. cízková. 1999. carbohydrate storage in rhizomes of phragmites australis: the effects of altitude and rhizome age. aquatic botany 64: 105-110. kotschy, k.a., k.h. rogers & a.j. carter. 2000. patterns of change in reed cover and distribution issn 0075-6458 39 koedoe 47/1 (2004) van rooyen.qxd 2004/04/15 03:12 page 39 in a seasonal riverine wetland in south africa. folia geobotanica 35: 363-373. kotze, d.c. & t.g.o. o’connor. 2000. vegetation variation within and among palustrine wetlands along an altitudinal gradient in kwazulu-natal, south africa. plant ecology 146: 77-96. kyle, r. 2000. a summary and some comments on resource utilization in the tembe/ndumo game reserve complex during 1999. internal report kwazulu-natal nature conservation services. lenssen, j.p.m., f.b.j. menting, w.h. van der putten & c.w.p.m. blom. 2000. variation in species composition and species richness within phragmites australis dominated riparian zones. plant ecology 147: 137-146. matthews, w.s., a.e. van wyk, n. van rooyen & g.a. botha. 2001. vegetation of the tembe elephant park, maputaland, south africa. south african journal of botany 67: 573-594. mckean, s.g. 2001. productivity and sustainable use of phragmites in the fuyeni reedbed hluhluwe-umfolozi park management guideline for harvest. south african journal of botany 67: 274-280. mook, j.h. & j. van der toorn. 1982. the influence of environmental factors and management on stands of phragmites australis. 2. effects on yield and its relationships with shoot density. journal of applied ecology 19: 501-517. ostendorp, w. 1995. effect of management on the mechanical stability of lakeside reeds in lake constance-untersee. acta oecologica 16: 277294. ostendorp, w. 1999. management impact on stand structure of lakeshore phragmites reeds. international review of hydrobiology 84: 33-47. schulze, r.e. 1982. agrohydrology and agroclimatology of natal. agricultural catchments research unit, report 14. university of natal, pietermaritzburg, south africa. soukup, a., o. votrubova & h. cízková. 2000. internal segmentation of rhizomes of phragmites australis: protection of the internal aeration system against being flooded. new phytologist 145: 71-75. soukup, a., o. votrubova & h. cízková. 2002. development of anatomical structure of roots of phragmites australis. new phytologist 153: 277287. strand, v.v. & s.e.b. weisner. 2002. interactive effects of pressurized ventilation, water depth and substrate conditions on phragmites australis. oecologia 131: 490-497. van der putten, w.h., b.a.m. peters & m.s. van den berg. 1997. effects of litter on substrate conditions and growth of emergent macrophytes. new phytologist 135: 527-537. van der toorn, j. & j.h. mook. 1982. the influence of environmental factors and management on stands of phragmites australis. 1. effects of burning, frost and insect damage on shoot density and shoot size. journal of applied ecology 19: 477-499. van wyk, a.e. & g.f. smith. 2001. regions of floristic endemism in southern africa: a review with emphasis on succulents. pretoria: umdaus press. van wyk, b.e. & n. gericke. 2000. people's plants. pretoria: briza publications. weisner, s.e.b. 1996. effects of organic sediment on performance of young phragmites australis clones at different water depth treatments. hydrobiologia 330: 189-194. weisser, p.j. & r.j. parsons. 1981. monitoring phragmites australis increases from 1937 to 1976 in the siyai lagoon (natal, south africa) by means of air photo interpretation. bothalia 13: 553-556. wynberg, r. 2002. a decade of biodiversity conservation and use in south africa: tracking progress from the rio earth summit to the johannesburg world summit on sustainable development. south african journal of science 98: 233-243. koedoe 47/1 (2004) 40 issn 0075-6458 van rooyen.qxd 2004/04/15 03:12 page 40 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 96.pdf page 1 696 --online appendix table 1 --14 october 2009.indd a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe online supplement table 1 a rticle #696 1vol. 51 no. 1 page 1 of 5 ��������� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������������������������� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������������������������� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������� � � � � � � � � � � � � � � � � � � ��� � ��� � ��� � ��� � ��� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �� � � � ������� � ����� � ����� � ����� � ����� � � � � � � ����� � ����� � � � � � ����� ����� � � � � � ������ �!� ���� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ��������� ������� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������������������ � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ���������������� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ��������� �������� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���" � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���# � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � $%�� �� & '���&���(&��) ���%�� %��(&��*�� �������� � )�$&��(��' +�&�,-����(�� .��(��� )� ����� �������&�,��(� ������� ��� ��������� ������ ���� ���������� � � �� �������������� ���� ������������ � ������ ������� �������� ������������ �������� ������� ���� ������ ���� ���������� ����� ����������� ���������������� �� �������� ����� ! ������� �������� �� ����� �������� � ���������������� ������������������ �� � �������� ���������� ������ ���������� ������ �������������� "������� ������������� #��������������������� $�������������������� ��� ��������������������� ������������������ ������������������� %���������������� � ������� ����������� #����������� ��� �������� ���� table 1 phytosociological table for the shrubland communities of platberg koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za online supplement table 1 brand, du preez & brown a rt ic le # 69 6 2 vol. 51 no. 1 page 2 of 5 ������ �!� ���! � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���/ � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���0 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���1 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���2 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���3 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���. � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���4 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ���� ���������������� ������ ������������� ���������������������� ������ ��������������� ������������������� &����� ������������� '���������������� ������ ������������ ���� ���������� ���� ������������ ��� �� ����������� %�������� ������ � ��������� ������ �������� ������ ������� (������� ����)����� ��������������� �� � ���������� � ���������������� �������� ��������������� "������� ����� ����� � ������)����� '���������� ���� #������ ������������� �� �� ��������������� ������������ ��������� ����������������� !�������� ���� � ! ���������������� � "�������������� �������� ������ ������ #���������� �������� ������ ��������������� ������������� ���� � �� ������ ������� !��������������� �*������� ���� ��� � �������������� ���� ����� ���� $��������������� +� ���������������� ����������������� ������������ ������ ����������������� �������� ���������� ����� ������ ���� (������� ����������� , ������� ��� ����������� ����������� ���� �������� �� ��� �������-������� table 1 (continues.....) phytosociological table for the shrubland communities of platberg a classification and description of the shrubland vegetation on platberg, eastern free state, south africa online supplement table 1 a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #696 3vol. 51 no. 1 page 3 of 5 ������ �!� ���$ � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���5 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���+ � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���6 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���7 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���8 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ����������������� $����������� #������������������ $���������������� � ,�)������������� ��������������� ����� ��������� ,��������������� &����� ���)������� ������������� ������� �������� ���� $����������� .�� ������������� ,������������ ���� ������ ���/������� ������������ ����� ����� ������ ���� ���������������� ����� . ���������-������� ������ �������� ��������� ���������������� +� �������� ���� $��� ����������� ������������������ ,� ������������ � ���������������������� �� )��������������� %���������������� ���� ��������� ���� &����� ������� �0�� ����� ����������������� $��������)���� ���� � ���� �������� �� ��������������� 1������������������������ ��������������������� ������������������ ��� ������������������ ������� 2������ ����� ����� 1���� ������� � � �������������������/�� ������������ ��������������������� ,�����*������������� table 1 (continues.....) phytosociological table for the shrubland communities of platberg online supplement table 1 brand, du preez & brown koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 69 6 4 vol. 51 no. 1 page 4 of 5 ������ �!� ���9 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ���: � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ����� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ����� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ����� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ����" � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ����� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ����# � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ����! � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ����/ � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ����0 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ������ �!� ����2 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � ���� ��� ����� �� ������ � �� �������� ���� $������ ����� &����� ������������ ����������� �� ���� �� �������������� ���� � ����� ��� �������������� '���������������� ����������������� ����������� �������� ������������������� ��������� ���� %���������� ��� ������������� ������ �� ������)���� �� �������� �������������� ���� ������������� ,�����*���� ����� $�������� ����� ��������� ��������� �������������� ����������� ���� $�������� ������� ���������� ������� ����� �������� ������� �������� �� ���� �� % � ����������������� , ���������������� �� �������������� ������� ���������������� ����������� ��� ��� �� ���� ����������� ���� ����� ����������� �������������� ��� �� $��� �� ����� ����� �� ������ �������������� ������� �!� ����1 ���� � �������� ����� ������������)������� 2������������������� !������������������������ ������������������� ������ ��������������� �� ������������ table 1 (continues.....) phytosociological table for the shrubland communities of platberg a classification and description of the shrubland vegetation on platberg, eastern free state, south africa online supplement table 1 a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #696 ������ �!� ����� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � #������������ ���� ����������� ����� ������������-��� �������� ����������������������� ���� ��������� ������� �� ������� ����� �� �� ���� ���� ���� �������� � ����� ���� ����� ������ ������������������ ��������� ����� vol. 51 no. 1 page 5 of 5 5 table 1 (continues.....) phytosociological table for the shrubland communities of platberg 690--30 oct 2009.indd [final version].indd a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe editorial a rticle #690 (page number not for citation purposes) 1 the status of koedoe one year after changing to an online publication mode author: llewellyn c. foxcroft1,2 affi liations: 1south african national parks, conservation services south africa 2centre for invasion biology, department of botany and zoology, south africa correspondence to: llewellyn c. foxcroft e-mail: llewellynf@sanparks.org postal address: private bag x402, skukuza, 1350, south africa keywords: editorial; koedoe; african protected area conservation and science; future; history dates: received: 27 jan. 2009 accepted: 27 jan. 2009 published: 14 apr. 2009 how to cite this article: foxcroft, l.c., 2009, ‘the status of koedoe one year after changing to an online publication mode’, koedoe 51(1), art. #690, 1 page. doi: 10.4102/koedoe. v51i1.690 this article is available at: http://www.koedoe.co.za © 2009. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. vol. 51 no. 1 page 1 of 1 foreword 2008 represented the start of a new online era for koedoe, which provided us with a number of challenges and opportunities. the challenges lay in developing an entirely new publication and information dissemination system containing a number of new processes. the opportunities however, allowed us to build on koedoes’ 50 year publication history. the main opportunity for koedoe lies in using the open access publication route, where all our articles are freely available via the world wide web. further, all back issues of the journal will be available as pdf downloads by march 2009, additional special interest sections were added (for example, essays and book reviews) and the marketing strategy was expanded to reach a wider audience. as a result, 20 manuscripts were published online in 2008 and we hope they provide for interesting and varied reading. furthermore, we greatly appreciate the patience and support of the authors and readers during this time of refurbishment. below, i have included some statistics for 2008, in order to show the progress that has been made by koedoe in just one year. from the humble beginnings of only 178 web visitors in january 2008, to 2293 visitors in november (the month with the highest records), the koedoe website received a total of 12,602 unique visitors for the year. web traffi c was recorded from over 92 countries, with south africa presenting the highest levels of activity (6.9 gb of information downloaded). next was the united kingdom (595 mb), germany (415 mb), the netherlands (400 mb) and australia (266 mb), with a total 19.59 gb being downloaded. to date, the most viewed articles from 2008 include. a checklist of the spiders (arachnida, araneae) of the polokwane nature reserve, limpopo province, south africa by susan dippenaar, ansie dippenaarschoeman, mogadi modiba and thembile khoza, which has been viewed 307 times; fifty years of koedoe: current status and future directions by llewellyn foxcroft, viewed 231 times; the recent fi re history of the table mountain national park, and implications for fi re management by brian van wilgen and greg forsyth, viewed 220 times; measurement of concentrations of faecal glucocorticoid metabolites in free-ranging african elephants within the kruger national park, by jozua viljoen, andre ganswindt, rupert palme, hendrick reynecke, johan du toit and william langbauer jr, viewed 213 times; and an assessment of the implementation and outcomes of recent changes to the fi re management of the kruger national park, by brian van wilgen, navashni govender and sandra macfadyen, viewed 206 times. in total, the 20 articles published in 2008 were viewed 1577 times collectively. interestingly, the following articles from back issues uploaded during 2008 have also been extensively viewed. the effect of mature elephant bull introductions on ranging patterns of resident bulls: phinda private game reserve, south africa (2006) by heleen druce, k pretorius, d druce and r slotow, has been viewed 260 times; and a check list of the spiders of the kruger national park, south africa (arachnida: araneae) (2003) by anna dippenaarschoeman and a. leroy, has been viewed 258 times. i hope that you will be as encouraged as i am by these fi gures, and ensure your support further, as either a contributing author or reader. llewellyn foxcroft editor–in–chief references dippenaar, s.m., dippenaar-schoeman, a.s., modiba, m.a., & khoza, t.t., 2008, ‘a checklist of the spiders (arachnida, araneae) of the polokwane nature reserve, limpopo province, south africa’, koedoe 50(1), 10–17. dippenaar-schoeman, a.s. & leroy, a., 2003, ‘a checklist of the spiders of the kruger national park, south africa (arachnida: araneae)’, koedoe 46(1), 91–100. druce, h., pretorius, k., druce, d. & slotow, r., 2006, ‘the effect of mature elephant bull introductions on ranging patterns of resident bulls: phinda private game reserve, south africa’, koedoe 49(2), 77–84. foxcroft, l.c., 2008, ‘fifty years of koedoe: current status and future directions’, koedoe 50(1), 1–2. forsyth, g.g. & van wilgen, b.w., 2008, ‘the recent fi re history of the table mountain national park, and implications for fi re management’, koedoe 50(1), 3–9. van wilgen, b.w., govender, n. & macfadyen, s., 2008, ‘an assessment of the implementation and outcomes of recent changes to the fi re management of the kruger national park’, koedoe 50(1), 22–31. viljoen, j.j., ganswindt, a., palme, r., reynecke, h.c., du toit, j.t. & langbauer jr., w.r., 2008, ‘measurement of concentrations of faecal glucocorticoid metabolites in free-ranging african elephants within the kruger national park’, koedoe 50(1), 18–21. filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! gaugris.qxd the vegetation of tshanini game reserve and a comparison with equivalent units in the tembe elephant park in maputaland, south africa j.y. gaugris, w.s. matthews, m.w. van rooyen and j. du p. bothma gaugris, j.y. , w.s. matthews, m.w. van rooyen and j. du p. bothma. 2004. the vegetation of tshanini game reserve and a comparison with equivalent units in the tembe elephant park in maputaland, south africa. koedoe 47(1): 9–29. pretoria. issn 00756458. an analysis of the plant communities of the to be proclaimed tshanini game reserve and their relation with selected similar plant communities of tembe elephant park are presented. the study area lies 6 km due south of tembe elephant park within the maputaland centre of plant endemism, which is part of the maputaland-pondoland region recognised by the iucn as a centre of plant diversity. a braun-blanquet classification on 51 random sample plots revealed eight distinct, mainly woodland plant associations. vegetation distribution follows the same patterns as described in tembe elephant park. plant communities and subcommunities are mapped, described and compared to similar plant communities in tembe elephant park to detect differences between protected and unprotected areas. results from the qualitative comparison showed >50 % similarity between equivalent units and results from the quantitative comparison showed >25 % to 50 % similarity between equivalent units. mueller-dombois and ellenberg (1974) regard 25 % as the lower threshold value to indicate similar associations. the observed differences may relate to animal and human utilisation. key words: plant communities, sand forest, similarity indices, tembe elephant park, tshanini game reserve j.y. gaugris and j. du p. bothma, centre for wildlife management, university of pretoria, pretoria, 0002 republic of south africa; w.s. matthews, tembe elephant park, ezemvelo kwazulu-natal nature wildlife, private bag 356, kwangwanase, 3973 republic of south africa; m.w. van rooyen department of botany, university of pretoria, pretoria, 0002 republic of south africa. issn 0075-6458 9 koedoe 47/1 (2004) introduction the tembe elephant park was proclaimed in 1983 after negotiations between the then kwazulu bureau of natural resources and the tembe tribal authority in consultation with the local communities of northern maputaland, kwazulu-natal, south africa. the park boundaries were subsequently fenced and animal numbers started to increase. the fence has kept the utilisation of renewable natural resources by the local communities at bay for the past 19 years. in this period, the vegetation of the park has been utilised only by the indigenous fauna, but it has been affected by management decisions and possibly also regional environmental changes. the tshanini game reserve was established in 2000 on some land of the community of manqakulane, but it has not yet been registered. the reserve’s creation developed from the initiative of the local community, from as far back as the creation of tembe elephant park. the ideal of the manqakulane people was to create their own reserve on their own land to counteract high unemployment and to raise funds for the development of the community through ecotourism. for the past 20 years, the land where the tshanini game reserve has been established was utilised by the local people for wood harvesting, livestock grazing, hunting, fruit and honey gathering, and subsistence cultivation where this was viable. it is presumed that large ungulate numbers have always been low in tshanini gaugris.qxd 2004/04/15 06:28 page 9 game reserve, due to the absence of standing water. however, hunting pressure has affected the larger mammal populations, and only small, secretive antelope such as the suni neotragus moschatus and red duiker cephalophus natalensis have survived there. the tembe elephant park and the tshanini game reserve are underlain by the same type of soil and they share the same broad vegetation types. moll (1977) distinguished the pallid bushveld and the sand forest zones in the area, whereas low & rebelo (1980) recognised the subhumid lowveld bushveld of the savanna biome and the sand forest of the forest biome. the two reserves lie close to each other with the northern boundary of tshanini game reserve some 6 km due south of the southern boundary of tembe elephant park. both of these conservation areas lie in the core of the maputaland centre of plant endemism as described by van wyk (1996). in both reserves a rare forest type, known as sand forest occurs (van wyk 1996; van rensburg et al. 1999; kirkwood & midgley 1999; matthews et al. 2001; van wyk & smith 2001). the sand forest in this particular area is further defined as the eastern sand forest by macdevette et al. (1989) and kirkwood & midgley (1999). in south africa and in mozambique, this forest type is restricted to koedoe 47/1 (2004) 10 issn 0075-6458 fig. 1. map of maputaland with the locations of tembe elephant park and tshanini game reserve in northern kwazulu-natal, south africa. gaugris.qxd 2004/04/15 06:28 page 10 issn 0075-6458 11 koedoe 47/1 (2004) the ancient coastal dune cordons of northern kwazulu-natal, and it is home to many rare and unusual plant and animal species (kirkwood & midgley 1999; van wyk & smith 2001). sand forest in south africa covers only 354 km², of which 44 % is conserved (low & rebelo 1998) with the largest protected area in south africa in the tembe elephant park (mcgeogh et al. 2002). the major part of this vegetation type is found in mozambique but the extent of it is unknown at present, because of a lack of detailed studies. cattle grazing, firewood demands and elephant loxodonta africana impact are the main identified threats for the sand forest (van wyk 1996; van rensburg et al. 1999). the human population growth rate in maputaland is high, increasing both by natural growth and immigration, despite a high mortality rate from the aids epidemic. roads have been constructed through the communities, and safe drinking water is provided. although these are laudable rural developments, they may become a threat to the conservation of the remote and less exploited areas of northern maputaland. matthews et al. (2001) did a complete vegetation survey of tembe elephant park, and suggested that their data could be used for management and land use planning for other similar areas in maputaland. however, the questions are to what extent has human pressure affected the vegetation in comparison with wildlife use; can the vegetation of tembe elephant park be regarded as representative of similar areas in maputaland and how well did the sand forest fare outside a formal conservation area? in this paper we aim to provide ecological and floristic data specifically for the tshanini game reserve by identifying, characterising, interpreting and mapping the major vegetation units that occur there. further objectives are to compare the vegetation of tshanini game reserve with similar units in tembe elephant park in terms of species composition and abundance, and to relate the perceived changes to differences in utilisation pressure. study area tshanini game reserve is 2 420 ha in extent and its northern boundary lies 6 km due south of the southern boundary of tembe elephant park (fig. 1). the area consists mainly of a sandy plain that is interspersed with ancient littoral dunes, and is covered by an open to closed woodland with patches of short to tall sand forest (fig. 2). the area covered by tshanini game reserve is unfenced and therefore subject to human influence. cattle grazing with small herds, rarely exceeding 20 heads, and wood collecting, either for construction or fire, are the most important forms of natural resource utilisation. other less intensive forms of use include fruit gathering, the collecting of honey and thatching grass, and hunting of birds and mammals. until 1995, the eastern portion of tshanini was still utilised for agriculture, with some large fields cut into the woodland or forest. the manqakulane community resided on the eastern boundary of the reserve until 1990, when they moved to 3 km east of the eastern boundary of tshanini game reserve. the relocation followed the installation of a governmental clean water supply to the local communities. the new location of the human settlements on the southern part of the muzi swamp has a better soil potential for agriculture. the community of manqakulane today controls an area of just over 10 000 ha, with a population of approximately 700 people in 110 households that are concentrated around the muzi swamp. topography tshanini game reserve lies centrally in the southernmost portion of the mozambican coastal plain between the lebombo mountains in the west and the indian ocean in the east. ancient littoral dunes that run in a broadly north to south direction are interspersed with inter-dune depressions. to the west of the reserve a seepage line forms the source of the nhlole river, a non-perennial seepage line winding to makane’s drift and flowing into the pongola river. the seepage gaugris.qxd 2004/04/15 06:28 page 11 koedoe 47/1 (2004) 12 issn 0075-6458 fig. 2. the vegetation of the tshanini game reserve and surrounding area in maputaland, northern kwazulu-natal, south africa. gaugris.qxd 2004/04/15 06:28 page 12 issn 0075-6458 13 koedoe 47/1 (2004) fig. 3. climatogram of the sihangwane weather station, tembe elephant park, maputaland, northern kwazulu-natal, south africa (b = height above sea level, c = duration of observations in years, d = mean annual temperature in °c, e = mean annual precipitation in mm, f = mean daily minimum of the coldest month in °c, g = lowest temperature recorded in °c, h = mean daily maximum of warmest month in °c, i = highest temperature recorded in °c, j = mean daily temperature variation in °c, m = dry season, n = humid season, o = mean monthly rainfall > 100 mm) following walter (cox & moore 1994). line runs from the southeast to the northwest. the tshanini game reserve has a mean altitude of 80 m above sea level, and two main ancient littoral dunes occur in it. the dune on the eastern boundary is the southern extension of nhlela ridge that extends into the western portion of tembe elephant park. a beacon is positioned on top of the ridge at an altitude of 103 m above sea level. a second dune runs along the western boundary at an altitude of 95 m above sea level. between the two main dunes, a third but smaller dune formation is found that starts in the middle of tshanini game reserve and increases in height towards the southern-central part of the reserve. the nhlole river lies to the east of the western dune cordon, and the lowest point of the reserve is 64 m above sea level in the northeastern corner. climate rainfall and temperature data for the weather station at sihangwane at the entrance of tembe elephant park (32.42388 ºe, 27.04316 ºs) appear in fig. 3. maputaland lies in a transition zone between the tropics to the north and subtropics to the south. the summers are hot and the winters cool to warm. the summer months are the wettest, although rainfall occurs throughout the year. in the drier winter months, morning mist is a common phenomenon especially in the swamp areas. the relative humidity of the air in summer is higher than in winter (matthews et al. 2001). geology the underlying geology of tshanini game reserve consists of cretaceous siltstone units that form the base of the mozambican coastal plain. this formation is overlain by sediments of miocene and pleistocene origin. the maputaland group overlies the latter. on top of this formation lie the marine, littoral and coastal dune deposits that were formed by successive marine transgressions and regressions. the oldest dunes of the coastal plain date back approximately 5 million years to the early pleistocene. the recent dunes are as young as 10 000 years, from the late pleistocene to 500 years ago for the current coastal dunes. the dunes of maputaland are among the most recent geological formations that are found in southern africa (hobday 1976). soils the soils of tshanini game reserve are similar to those described by matthews et al. (2001) for tembe elephant park, and specifically so for the dunes and inter-dune depressions. the soils have a thin, organically enriched a-horizon underlain by a sandy subsoil. the clay content is attributed gaugris.qxd 2004/04/15 06:28 page 13 to the weathering of labile minerals over a long period. the soils along the two main dunes are similar to those described for the nhlela ridge in tembe elephant park. the soils are well-drained profiles with less than 5 % clay, have a high base status, and are classified as hutton or clovelly forms (soil classification working group 1991; matthews et al. 2001). the inter-dune depressions are underlain by moderately to well-drained yellowish clovelly or grey fernwood soils (soil classification working group 1991; matthews et al. 2001). methods aerial photographs were obtained from the surveyor general in pretoria, south africa. the appropriate 1:50 000 scale photograph was scanned and digitised by using the geographical information tool cartalinx (the spatial data builder, version 1.1. clarks laboratories, 1998) to create a geographical information system (gis) based geo-referenced map of the homogeneous physiognomic and physiographic units of the reserve. the map was then refined by using the arcview gis tool (arc view gis, version 3.1, environmental systems research institute 1998). in all, 51 sample plots, of 30 m x 30 m each, were distributed in a stratified random manner throughout the study area. vegetation sampling was done from january to may 2001. the terminology that was used to name and describe the vegetation structure of the plant communities follows edwards (1983). for each sample plot, a relevé of the floristic composition and cover-abundance values for each plant species, according to the braun-blanquet coverabundance scale (mueller-dombois & ellenberg 1974), were recorded. environmental and ecological information collected for each sample plot included slope, aspect, altitude, topography, rock cover, erosion percentage, cover for canopy height classes of herbs, grasses, shrubs and trees and biotic influence. unidentified plant species were sampled for later identification at the herbarium of the tembe elephant park. the floristic data were classified with braun-blanquet procedures using the turboveg and megatab computer packages (turboveg for windows version 1.97, hennekens & schaminee 2001). this vegetation classification system was used to refine and interpret the preliminary map of homogeneous physiognomic units to produce a more detailed vegetation map of the tshanini game reserve. as it appeared impossible to digitise every vegetation unit and subunit separately, some units were grouped in mosaics. these mapping units are indicated in fig. 2. the homogeneous vegetation units that are common to tembe elephant park and tshanini game reserve were then analysed further to determine the degree of similarity or dissimilarity in terms of species composition and abundance by using similarity indices as described by mueller-dombois & ellenberg (1974). the phytosociological table created for tshanini game reserve and the table obtained from the study of matthews et al. (2001) for tembe elephant park were used to calculate the indices. sorensen’s qualitative index of similarity (muellerdombois & ellenberg 1974) that is based on the absence or presence of plant species was used to calculate indices of similarity in plant species composition. to be able to do so, a similar number of comparable relevés were compared in each of the vegetation communities. the following equation was used: sorensen’s index of similarity = (2c x 100) / a + b where: c = number of species common to both communities a = total number of species in community a b = total number of species in community b. the similarity index of motyka et al. (mueller-dombois & ellenberg 1974) was used to calculate a quantitative similarity index in terms of species abundances. the equation is: motyka et al.’s index of similarity = (2 mw x 100) / (ma + mb) where: mw refers to the sum of the smaller quantitative values of the species that are common to the two communities; ma is the sum of the quantitative values of all the species in one community; mb is the sum of all the quantitative values of all the species in the other community. to calculate the values of mw, ma and mb, the braun-blanquet cover-abundance values in the phytosociological tables of both reserves were converted into mean cover percentages. to accomplish this each cover-abundance value on the braun-blanquet scale was replaced by a cover value, whereafter a mean for the community could be calculated. although this is a crude method it does allow quankoedoe 47/1 (2004) 14 issn 0075-6458 gaugris.qxd 2004/04/15 06:28 page 14 titative comparisons to be made. when comparing vegetation communities, the similarity in terms of absence or presence of species in two communities is more important than the quantitative contribution of each species to a community (mueller-dombois & ellenberg 1974). the latter can be used to refine the comparisons. results and discussion classification three major plant communities and eight sub-communities were identified for the tshanini game reserve (tables 1 & 2, fig. 2). the tshanini game reserve lies on regic sand and no major geological feature could be identified that plays a role in the differentiation of the plant communities. matthews et al. (2001) postulated that clay content, moisture level in the soil and vegetation dynamics were major determinants of vegetation differentiation in tembe elephant park. however, clay content and moisture level differences in the soil of various areas in tshanini game reserve are slight and could not be directly related to the differentiation between plant communities in tshanini game reserve. except for the sand forest, the plant communities do not have sharp boundaries. the open and closed woodlands form mosaics that grade into each other and the boundaries are difficult to map on an aerial photograph. however, the plant communities are relatively easily recognised in the field. description of plant communities 1. the urelytrum agropyroides-sapium integerrimum open woodland this open woodland is the most widespread community in tshanini game reserve. it is associated with the dunes and inter-dune depressions on grey-brown to orange-grey dystrophic regosols. these soils have a low clay content and are associated with a low ph value of 5.5 (matthews et al. 2001). the open woodland covers 37.7 % of tshanini game reserve and represents a surface area of 912 ha. this plant community is characterised by diagnostic species of species group e. plant species of species group k form the link between this open woodland community and the closed woodland community. diagnostic plant species are the tree albizia adianthifolia, the grasses diheteropogon amplectens and urelytrum agropyroides and the shrub acridocarpus natalitius var. linearifolius. issn 0075-6458 15 koedoe 47/1 (2004) table 1 classification of the vegetation communities for tshanini game reserve in northern kwazulu-natal, south africa 1. the urelytrum agropyroides-sapium integerrimum open woodland 1.1. the albizia versicolor-diheteropogon amplectens sparse woodland 1.2. the indigofera podophylla-albizia adianthifolia ecotonal sparse woodland 1.3. the antidesma venosum-urelytrum agropyroides open woodland 1.4. the fimbristylis complanata-diheteropogon amplectens open woodland 2. the sclerocarya birrea-strychnos madagascariensis closed woodland 2.1. the cordia monoica-sclerocarya birrea open to closed woodland 2.2. the melinis repens-sclerocarya birrea closed woodland 3. the ptaeroxylon obliquum-hymenocardia ulmoides sand forest 3.1. the rothmannia fischeri-ptaeroxylon obliquum low to short sand forest 3.2. the strychnos henningsii-ptaeroxylon obliquum tall sand forest. gaugris.qxd 2004/04/15 06:28 page 15 koedoe 47/1 (2004) 16 issn 0075-6458 ta bl e 2 p hy to so ci ol og ic al ta bl e of th e ve ge ta tio n of t sh an in i g am e r es er ve in n or th er n k w az ul un at al , s ou th a fr ic a gaugris.qxd 2004/04/15 06:28 page 16 issn 0075-6458 17 koedoe 47/1 (2004) ta bl e 2 (c on tin ue d) gaugris.qxd 2004/04/15 06:28 page 17 koedoe 47/1 (2004) 18 issn 0075-6458 ta bl e 2 (c on tin ue d) gaugris.qxd 2004/04/15 06:28 page 18 issn 0075-6458 19 koedoe 47/1 (2004) ta bl e 2 (c on tin ue d) gaugris.qxd 2004/04/15 06:28 page 19 koedoe 47/1 (2004) 20 issn 0075-6458 ta bl e 2 (c on tin ue d) gaugris.qxd 2004/04/15 06:28 page 20 issn 0075-6458 21 koedoe 47/1 (2004) prominent species with a constancy of >50 % include the shrub vangueria infausta, the grass andropogon gayanus and the trees acacia burkei, strychnos madagascariensis, terminalia sericea, combretum molle and sapium integerrimum. the dwarf shrubs pavetta schumanniana, eugenia mossambicensis and the forb corchorus junodii have a high constancy in this community (table 2). a mean of 45 species was recorded per relevé. of the plant species that were recorded in this community, 18 species are endemic to maputaland (van wyk 1996). structurally, according to the classification of edwards (1983) this community is a woodland that varies from sparse to open. the non-continuous canopy can reach heights of 10–12 m, but a height of 6–8 m is the norm. this community is subdivided into four subcommunities based on the density, structure and floristic composition of the vegetation. 1.1 the albizia versicolor-diheteropogon amplectens sparse woodland this subcommunity is found on the dune crests, slopes and inter-dune depressions of the western side of the tshanini game reserve and occurs on acidic orange-grey dystrophic regosols. the subcommunity is characterised by the diagnostic species of species group a and common species of species groups e, j and k. the diagnostic species are the tree albizia versicolor, the forb oxygonum robustum and the shrub grewia inaequilatera. prominent species are the trees albizia adianthifolia, sapium integerrimum and strychnos madagascariensis, the grasses diheteropogon amplectens, urelytrum agropyroides, perotis patens and pogonarthria squarrosa, the shrub acridocarpus natalitius var. linearifolius and the dwarf shrubs pavetta schumanniana, eugenia mossambicensis and albertisia delagoensis. a mean of 45 plant species was recorded per relevé. according to the classification of edwards (1983), this subcommunity is a short sparse woodland with a few tall trees that reach a height of 6–8 m. the tree layer is underlain by a short to tall grassland up to 2 m tall. 1.2 the indigofera podophylla-albizia adianthifolia ecotonal sparse woodland this subcommunity is also found on the dune crests, slopes and inter-dune depressions on the western portion of the tshanini game reserve on acidic orange-grey dystrophic regosols. the subcommunity is not characterised by its own distinct species group but shares species group b with community 1.1. prominent species in this community are found in species groups e, j and k. this subcommunity forms a transition zone linking the sparse and open woodlands. diagnostic species are the tree combretum mkuzense and the shrubs garcinia livingstonei and indigofera podophylla. the prominent species include the trees albizia adianthifolia, sapium integerrimum and strychnos madagascariensis, the grasses diheteropon amplectens, urelytrum agropyroides, perotis patens, digitaria eriantha and pogonarthria squarrosa, the shrub acridocarpus natalitius var. linearifolius and dwarf shrubs pavetta schumanniana, eugenia mossambicensis and albertisia delagoensis. a mean of 46 plant species was recorded per relevé. according to the classification of edwards (1983), this subcommunity is a short sparse woodland, with a few tall trees that reach a height of 6–8 m. the tree layer is underlain by a short to tall grassland up to 2 m tall. 1.3 the antidesma venosum-urelytrum agropyroides open woodland this subcommunity is found on the dune crests, slopes and inter-dune depressions on the western portion of the tshanini game reserve. it occurs on acidic orange-grey dystrophic regosols and is characterised by diagnostic species of species group c with common species of species groups e, i, j and k. the diagnostic species are the tree antidesma venosum, the forbs oxygonum dregeanum and hoslundia opposita, the gaugris.qxd 2004/04/15 06:28 page 21 koedoe 47/1 (2004) 22 issn 0075-6458 sedge cyperus obtusiflorus and geophyte hypoxis hemerocallidea. the prominent species are the trees albizia adianthifolia, sapium integerrimum and strychnos madagascariensis, the grasses diheteropogon amplectens, urelytrum agropyroides, eustachys paspaloides and perotis patens, the shrub acridocarpus natalitius var. linearifolius and dwarf shrubs eugenia mossambicensis and diospyros galpinii. a mean of 52 plant species was recorded per relevé. this makes this plant subcommunity the second richest unit sampled in the tshanini game reserve. according to the classification of edwards (1983), this subcommunity is a short open woodland with a few tall trees that reach a height of 6–8 m. the tree layer is underlain by a short to tall grassland up to 2 m tall. 1.4 the fimbristylis complanatadiheteropogon amplectens open woodland this subcommunity is found on the dune crests, slopes and inter-dune depressions on the western portion of the tshanini game reserve. it occurs on acidic orange-grey dystrophic regosols (matthews et al. 2001). the subcommunity is characterised by the diagnostic species of species group d with common species in species groups e, i, j and k. the diagnostic species are the sedge fimbristylis complanata, the dwarf shrub parinari capensis subsp. incohata, the forb tricliceras mossambicense and the grass alloteropsis semialata. the prominent species are the trees albizia adianthifolia and strychnos madagascariensis, the grasses diheteropogon amplectens, urelytrum agropyroides, eustachys paspaloides, perotis patens and pogonarthria squarrosa, the shrub acridocarpus natalitius var. linearifolius and the dwarf shrub eugenia mossambicensis. a mean of 37 plant species was recorded per relevé. according to the classification of edwards (1983), this subcommunity is a short open woodland with a few tall trees that reach a height of 6–8 m. the tree layer is underlain by a short to tall grassland up to 2 m tall. 2. the sclerocarya birrea-strychnos madagascariensis closed woodland in the tshanini game reserve the closed woodland community is associated with the dunes and inter-dunes depressions and occurs on grey dystrophic regosols. the soils have a low clay content associated with a low ph value of 5.5 (matthews et al. 2001). the closed woodland covers 19.1 % of the tshanini game reserve and represents a surface area of 463 ha. this plant community is characterised by species of species group h. diagnostic plant species are the trees sclerocarya birrea, ziziphus mucronata and bridelia cathartica, the grass eragrostis superba, the shrub gymnosporia senegalensis and forb priva cordifolia. prominent species with a constancy >50 % include the shrubs vangueria infausta and diospyros dichrophylla, the trees strychnos madagascariensis, strychnos spinosa, euclea natalensis and tabernaemontana elegans and the grasses digitaria eriantha, perotis patens and pogonarthria squarrosa (table 2). a mean of 52 plant species was recorded per relevé. of the plant species that were recorded in this community, 15 species are endemic or near endemic to maputaland (van wyk 1996). according to the classification of edwards (1983), this community is a short, open to closed woodland that can be distinguished by a semi-continuous to continuous canopy reaching heights from 6–10 m. a tall to high shrub component, which forms dense clumps around the base of trees, is associated with this community. the tree and shrub canopies are underlain by a continuous mixed layer of short grasses and herbs. this community is subdivided into two subcommunities based on the structure and floristic composition of the vegetation. 2.1 the cordia monoica-sclerocarya birrea open to closed woodland this subcommunity is found on the dune crests, slopes and inter-dune depressions in the eastern half of the tshanini game reserve on acidic grey-brown dystrophic gaugris.qxd 2004/04/15 06:28 page 22 issn 0075-6458 23 koedoe 47/1 (2004) regosols. it is characterised by diagnostic species of species group f with common species of species groups h, i and k. this subcommunity is usually found around ancient settlements and shows a slow recovery from human impact. old settlements can still be located and trees grow back on the former cultivated lands around these emplacements. the outline of the old fields is still distinguishable from surrounding vegetation. the diagnostic species are the shrubs cordia monoica, lantana rugosa and lippia javanica, the herb helichrysum nudifolium and the grass setaria sphacelata. prominent species are the trees sclerocarya birrea, ziziphus mucronata, tabernaemontana elegans and strychnos madagascariensis, the grasses eragrostis superba, digitaria eriantha and perotis patens, the shrubs gymnosporia senegalensis, strychnos spinosa, bridelia cathartica and dwarf shrub eugenia mossambicensis. a mean of 56 plant species was recorded per relevé, which makes it the richest plant subcommunity sampled in the tshanini game reserve. according to the classification of edwards (1983), this subcommunity is a short, open to closed woodland with a semi-continuous canopy, reaching heights of 8–10 m. the tree layer is underlain by tall to high shrubs as well as a continuous grass layer up to 1 m tall. 2.2 the melinis repens-sclerocarya birrea closed woodland this subcommunity is found on the dune crests, slopes and inter-dune depressions in the eastern half of the tshanini game reserve. it occurs on acidic grey-brown dystrophic regosols. this subcommunity is characterised by diagnostic species of species group g, with common species in species groups h, i and k. the diagnostic species are the tree trichilia emetica, the shrub phyllanthus reticulatus, the herb nidorella auriculata and the grasses melinis repens and eragrostis ciliaris. prominent species are the trees sclerocarya birrea, ziziphus mucronata, tabernaemontana elegans and strychnos madagascariensis, the grasses eragrostis superba, digitaria eriantha and perotis patens, the shrubs gymnosporia senegalensis, strychnos spinosa and bridelia cathartica and dwarf shrub eugenia mossambicensis. a mean of 48 plant species was recorded per relevé. according to the classification of edwards (1983), this subcommunity is a short closed woodland with a semi-continuous to continuous canopy reaching a height of 8–10 m. the tree layer is underlain by a mixed short layer of grasses and herbs as well as tall to high shrubs. 3. the ptaeroxylon obliquum-hymenocardia ulmoides sand forest the sand forest is a major community in tshanini game reserve. it is associated with the dunes and inter-dunes depressions on orange-brown arenosols and dystrophic regosols. the soils have a low clay content associated with a low ph value of 5.6 (matthews et al. 2001). the sand forest covers 43.2 % of tshanini game reserve and represents a surface area of 1 045.5 ha. this plant community is characterised by diagnostic species of species group o. consistent diagnostic species are the tree ptaeroxylon obliquum, the shrub or liana uvaria caffra, the understorey shrubs drypetes arguta and toddaliopsis bremekampii. prominent species with a constancy of >50 % include the shrubs croton pseudopulchellus, hyperacanthus microphyllus, monodora junodii, salacia leptoclada and croton steenkampianus, the trees cleistanthus schlechteri, hymenocardia ulmoides, pteleopsis myrtifolia, brachylaena huilensis and balanites maughamii. the understorey shrub synaptolepis kirkii has a high constancy in this community (table 2). a mean of 40 plant species was recorded per relevé. of the plant species that were recorded, 20 species are endemic to maputaland with 10 species restricted to the sand forest type (van wyk 1996; matthews et al. 2001). according to the classification of edwards (1983), this community is classified as a forest, with a closed canopy reaching heights from 5 m for the low sand forest subcommunity to 12 m for the tall sand forest subcommunity. this community is subdivided into gaugris.qxd 2004/04/15 06:28 page 23 koedoe 47/1 (2004) 24 issn 0075-6458 two subcommunities based on the structure and floristic composition of the vegetation. 3.1 the rothmannia fischeri-ptaeroxylon obliquum low to short sand forest this subcommunity is found on the dune crests, slopes and inter-dune depressions throughout the tshanini game reserve on acidic orange-brown arenosols and dystrophic regosols. it is characterised by diagnostic species of species group l and common species of species groups n and o. the diagnostic species are the tree rothmannia fischeri, the creeper tecoma capensis and the shrub tarenna littoralis. prominent species are the trees ptaeroxylon obliquum, balanites maughamii, brachylaena huilensis, pteleopsis myrtifolia and hymenocardia ulmoides, the lianas uvaria caffra, uvaria lucida and hippocratea delagoensis, the shrubs ochna natalitia, boscia filipes and hyperacanthus microphyllus. the understory shrub synaptolepis kirkii has a high constancy in this subcommunity. a mean of 48 plant species was recorded per relevé. according to the classification of edwards (1983), this subcommunity is a low to short forest with a continuous canopy that reaches a height of 5–7 m. the understory is dense and nearly impenetrable with no intermediate strata between the canopy and ground. at ground level a low, semi-continuous layer of mixed grass and herbs is found. there is a controversy regarding the forest status of the low sand forest. the low canopy associated with the absence of secondary intermediate canopy strata suggests that this subcommunity should be classified as tall thicket as defined by low & rebelo (1998). izidine et al. (2003) suggest that such vegetation types should be named licuati thicket. 3.2 the strychnos henningsii-ptaeroxylon obliquum tall sand forest this subcommunity is found on the dune crests, slopes and inter-dune depressions throughout the tshanini game reserve. it occurs on low clay content, acidic orangebrown arenosols and dystrophic regosols. this subcommunity is characterised by diagnostic species of species group m and common species of species groups n and o. the diagnostic species are the trees strychnos henningsii, manilkara concolor and wrightia natalensis, the shrubs dovyalis longispina, commiphora neglecta and tricalysia capensis. prominent species are the trees ptaeroxylon obliquum, newtonia hildebrandtii, haplocoelum gallense, ochna arborea, brachylaena huilensis and monodora junodii, the lianas uvaria caffra, uvaria lucida and hippocratea delagoensis, the shrubs ochna natalitia, drypetes arguta, toddaliopsis bremekampii, salacia leptoclada, monanthotaxis caffra and hyperacanthus microphyllus. a mean of 39 plant species was recorded per relevé. according to the classification of edwards (1983), this subcommunity is structurally a tall forest with a closed canopy that reaches a height of 12–15 m for the largest and oldest trees. an intermediate canopy of smaller trees and high shrubs reaches heights of 5–6 m. the ground level is covered with a semi-continuous layer of low to short herbs. the forest is relatively open and easy to walk through. izidine et al. (2003) suggest that such vegetation types should be named licuati forest. the nhlole seepage line area was not sampled and has to be investigated further. it appears that the seepage line may have a duplex soil and that the vegetation type along the nhlole should be similar to the spirostachys africana-berchemia zeyheri closed woodland on duplex soils described by matthews et al. (2001). this area has been mapped as woodland on clay in fig. 2. similarity indices the concept of some degree of similarity or dissimilarity between plant communities is difficult to quantify. it is however, possible to use mathematical models to describe such relationships, but because of the mathematical nature of these models, arbitrary limits have to be set (mueller-dombois & ellengaugris.qxd 2004/04/15 06:28 page 24 issn 0075-6458 25 koedoe 47/1 (2004) berg 1974). several indices have been used in the past to quantify the degree of similarity between homogeneous vegetation units or plant communities. jaccard’s similarity index is a simple mathematical expression that is based on the ratio of the plant species that are common to two communities, and the total number of species present. sorensen’s similarity index is an evolution of jaccard’s and is also used to compare vegetation communities on the basis of the absence or presence of species. the similarity index of motyka et al. (mueller-dombois & ellenberg 1974) is a modification of the sorensen similarity index and is designed to treat data on vegetation structure by quantifying each species that is present (mueller dombois & ellenberg 1974). to interpret similarity indices based on the absence or presence of species, mueller-dombois & ellenberg (1974) suggest that a 25 % up to 50 % overlap in species indicates similar vegetation communities, with 25 % as the lower threshold below which communities are considered as different. similarity indices exceeding 50 % show that the two vegetation communities are so similar that more fieldwork might be necessary to obtain ecologically meaningful information (mueller dombois & ellenberg 1974). the vegetation units from tembe elephant park that were used in this comparison are described by matthews et al. (2001) as: the drypetes arguta-uvaria lucida subsp. virens sand forest community of the deep sandy areas (dry sands) the acacia burkei-euclea natalensis closed woodland and thicket subcommunity the salacia kraussii-themeda triandra open woodland on sand subcommunity the similarity indices indicate that the three main plant communities of tshanini game reserve are similar to their equivalent plant communities and subcommunities described for tembe elephant park (table 3). these results therefore confirm the prediction made by matthews et al. (2001) that the plant communities that were described for tembe elephant park can be extrapolated to other parts of maputaland and can be used for further planning and development of tshanini game reserve and any other similar areas. the qualitative similarity in species composition between the two sets of data is exceptionally high (table 3). however, the quantitative similarity indices suggest that there are more pronounced differences between the two reserves. the tembe elephant park has been protected from human utilisation since 1983, although various wildlife species were introduced or re-established there. the tshanini game reserve on the other hand has been devoid of most forms of wildlife for a similar period of time, but it has been subject to human utilisation, cattle grazing and the associated fire regime, especially in the sparse to open woodland areas where grassland occurs. the leaders from the community of manqakulane still allow cattle grazing on the land of the tshanini game reserve. table 3 a comparison between the vegetation of tembe elephant park and tshanini game reserve by means of the sorensen similarity index based on species composition and that of motyka et al. (in: mueller-dombois & ellenberg 1974) based on species abundance vegetation type species composition: species abundance: presence-absence data cover-abundance data open woodland 50.9 % 31.8 % closed woodland 53.4 % 26.8 % sand forest 54.6 % 39.0 % gaugris.qxd 2004/04/15 06:28 page 25 koedoe 47/1 (2004) 26 issn 0075-6458 as a result, small cattle herds still roam the reserve, especially in the open woodlands. the cattle are herded to and from the centre of the community through small tracks cut through the bush, usually avoiding the sand forest vegetation type. the cattle herders have also been observed repeatedly burning the open woodland areas in tshanini game reserve to keep the grass green for the cattle. the high similarity in plant species composition between the two reserves, despite the different utilisation regimes over a period of 19 years is noteworthy. although no factual data on human population numbers and the intensity of their use of natural resources are available, various authorities in the northern maputaland region agree that both have increased over the past 19 years. the degree of similarity that still exists in the plant resources suggests that either the human presence and utilisation of these resources in tshanini game reserve has been consistently low, or that the wildlife utilisation in tembe elephant park has been high, or that both forms of utilisation lead to the same pressure on the species. however, the lower similarity between the two reserves when a quantitative measure is used suggests that there are differences in terms of cover of the species. the sand forest community in tshanini game reserve has a significantly higher mean cover value per species than the equivalent community in tembe elephant park (t = 5.65, df = 8, p < 0.001). although the species composition has remained relatively unchanged in both reserves, the sand forest vegetation in the tshanini game reserve is denser than in tembe elephant park. the assumption made by van rensburg et al. (1999) that the sand forest around the park suffers the same from human pressure as from elephant impact inside the park may not be true as the distance from the forest to human settlements increases. the present study suggests that elephants may be affecting the sand forest in tembe elephant park and that they have already changed the plant cover significantly but not the species composition. concerning the closed woodland the similarity index is high in terms of plant species composition but is greatly reduced in terms of species cover indicating a definite difference in closed woodland structure between tshanini game reserve and tembe elephant park. the mean species cover value for the closed woodlands in tshanini game reserve is significantly higher than for tembe elephant park (t = 7.62, df = 9, p < 0.001). the larger mammals found in tembe elephant park are unlikely to have used the tshanini game reserve intensively at any stage in the past due to the absence of standing water. utilisation of tshanini game reserve was probably limited to migratory incursions from larger forms of wildlife, which must have been restricted in time and impact. small cattle herds, which have supplanted the wildlife in tshanini game reserve, prefer the more open woodland areas as opposed to the closed woodland and sand forest. the cattle herds were mostly observed walking through the closed woodland from one open woodland area to another. it therefore appears unlikely that the cattle herds could create a noticeable impact on the vegetation of the closed woodland community. the lower cover values in the closed woodland community in tembe elephant park may indicate that this vegetation type has been subjected to a high pressure from the wildlife since the complete fencing of the park in 1989. a similar phenomenon was reported in grazing exclosures where species diversity remained similar, but plant density and cover increased (lenzi-grillini et al. 1996). in areas that are subjected to animal utilisation and especially elephant presence, a decrease in vegetation cover and density is often observed. this is especially true when the stocking density of animals is higher than the ecological capacity, or the areas are close to natural or artificial watering points (brits et al. 2002, mapaure & campbell 2002, mosugelo et al. 2002). further research is needed to compare utilisation pressure and patterns by cattle herds in tshanini game reserve with wildlife in tembe elephant park. gaugris.qxd 2004/04/15 06:28 page 26 the similarity between the open woodland of the two reserves is once again noticeable in terms of plant species composition, but suggests a difference at the species abundance level. the mean species cover value for the open woodlands in tshanini game reserve is significantly higher (t = 12.45, df = 18, p <0.001) than for tembe elephant park. this difference can possibly be linked to the absence of wildlife in tshanini game reserve combined with the improper use of fire. it is unlikely that the small cattle herds that roam the open woodland of the tshanini game reserve now have had a noticeable impact on the vegetation. however, the higher density of the woody vegetation in tshanini game reserve may also be interpreted as a sign of bush encroachment. the repeated burning of the grass layer of the open and sparse woodland by the cattle herders to stimulate out of season growth might provide conditions favourable for bush encroachment (van rooyen 2002). cattle and wildlife have notoriously different requirements and impact on the vegetation (bothma 2002). moreover, the absence of the whole range of grazers and browsers in tshanini game reserve combined with the absence of tree damaging animals could also provide conditions favourable for the increase in density of the woody component in the open woodland. the improper use of fire and to a lesser extent cattle grazing as opposed to high intensity wildlife utilisation are suggested as probable causes for the difference observed in species cover between the open woodland of the two reserves. the trends observed in this study differ from those that were suggested by van rensburg et al. (1999) and verified by mcgeoch et al. (2002) working with a suite of dung beetle species as bio-indicators. van rensburg et al. (1999) suggested that the impact of elephants on sand forest structure inside tembe elephant park was comparable to the human impact outside the park. when selecting sites outside conserved areas, van rensburg et al. (1999) and mcgeoch et al. (2002) concentrated on the populated areas within close proximity of tembe elephant park. they attributed the change in the sand forest structure immediately around the park to an increase in human utilisation of these resources. the changes recorded mainly involved the opening up of the canopy in the sand forest, an action they deemed was comparable to the opening up of the canopy by elephants in tembe elephant park. in the rural community of manqakulane the situation appears to differ from that described by van rensburg et al. (1999), possibly because of the remoteness of the tshanini game reserve as opposed to areas adjacent to tembe elephant park. the latter areas are served by the main road that has become an extraction route for construction wood and especially for firewood. the vegetation adjacent to the road is also harvested for the sale of wood. human settlements are found up to 2 km south of the main road that passes along the southern boundary of the tembe elephant park. impact on vegetation surrounding human settlements can be observed as far as 2 km from the settlement location. the interpretation of the vegetation data suggests that the present human pressure on the tshanini game reserve in northern maputaland is still low. there is no doubt that the pressure is increasing because of the population increase over the past 10 years (h. els, centre for indigenous knowledge, university of pretoria, 0002 pretoria pers. comm.). however, local people prefer to remain close to the water resources, main roads and arable lands. a low potential for agriculture, the distance from a major road and a lack of water have all kept people away from tshanini game reserve. these factors have therefore contributed to the preservation of this environment. the distance of the tshanini game reserve from the community centre, where most people of the community of manqakulane have settled, is less than 3 km. however, this distance in such a poor community where vehicles and other means of transportation of goods and resources are not readily available, seems to be sufficient to prevent an excessive impact on the reserve from human utilisation. most people have to carry the material which they harvest themissn 0075-6458 27 koedoe 47/1 (2004) gaugris.qxd 2004/04/15 06:28 page 27 selves, often as loads on their head, and this seems to have limited the utilisation of resources further than the immediate vicinity of the settlements. the availability of adequate material for human utilisation in the immediate vicinity of human settlements may also have contributed to the protection of land further away. however, there are indications that a limit of sustainable utilisation has been reached in the vicinity of the human settlements, as suggested by van rensburg et al. (1999), and that the people in this community may soon seek to utilise the plant resources further away (gaugris unpublished data 2003). conclusions a vegetation map for tshanini game reserve and its surrounding areas has been provided. moreover, the plant communities have been described, and were compared with similar vegetation units in tembe elephant park. this comparison between similar vegetation units shows that species composition is very similar but that plant cover values are significantly higher in tshanini game reserve than in tembe elephant park. these differences will have to be investigated in more detail to establish which condition should be regarded as representative of pristine conditions. the present study suggests that the sand forest in tshanini game reserve has not suffered degradation by human pressure, but that elephants may be affecting the sand forest in tembe elephant park and that they have already changed the plant cover significantly but not the species composition. in general the high intensity wildlife utilisation, notably by elephant, in tembe elephant park is suggested as probable cause for the difference observed in species cover between equivalent communities of the two reserves. the manqakulane community is still reasonably small and natural resource use is restricted to the vicinity of the settlements. this has apparently preserved large areas from been overexploited. acknowledgements we are grateful to thabani mthembu, who assisted with the fieldwork, the people from the community of manqakulane for authorising the research to be conducted on their land, the management staff from tembe elephant park and the ezemvelo kwazulunatal wildlife for the use of the research facilities and the logistical support, the centre for wildlife management, prof. h. els and the centre for indigenous knowledge from the university of pretoria for the material and financial support of the project. this material is also based upon work supported by the national research foundation under grant number 2047386. references bothma, j. du. p. 2002. combining wild and domestic herbivores. pp. 201–208. in: bothma, j. du p. (ed.). game ranch management. pretoria: van schaik. brits, j., m.w. van rooyen, & n. van rooyen. 2002. ecological impact of large herbivores on the woody vegetation at selected watering points on the eastern basaltic soils in the kruger national park. african journal of ecology 40: 53–60. cox, c.b. & p.d. moore. 1994. biogeography: an ecological and evolutionary approach. 5th ed. oxford: blackwell scientific publishing. edwards, d. 1983. a broad scale structural classification of vegetation for practical purposes. bothalia 14: 705–712. hennekens, s.k. & h.j. schaminee. 2001. turboveg, a comprehensive data base management system for vegetation data. journal of vegetation science 12: 589–591. hobday, d.k. 1976. quarternary sedimentation and the development of the lagoonal complex, lake st lucia, zululand. annals of the south african museum 71: 93–113. izidine, s., s. siebert & a.e. van wyk. 2003. maputaland's licuati forest and thicket, botanical exploration of the coastal plain south of maputo bay, with an emphasis on the licuati forest reserve. veld & flora 89: 56–61. kirkwood, d. & j.j. midgley. 1999. the floristics of sand forest in northern kwazulu-natal, south africa. bothalia 29: 293–304. lenzi-grillini, c.r., p. viskanic & m. mapesa. 1996. effects of 20 years of grazing exclusion in an area of the queen elizabeth national park, uganda. african journal of ecology 34: 333–341. koedoe 47/1 (2004) 28 issn 0075-6458 gaugris.qxd 2004/04/15 06:28 page 28 low, a.b. & a.g. rebelo. 1998. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. macdevette, d.r., d.k. macdevette, i.g. gordon. & r.l.c. bartholomew. 1989. floristics of the natal indigenous forests. pp. 124–144. in: geldenhuys, e.c.j. (ed.). biogeography of the mixed evergreen forests of southern africa. pretoria: foundation for research development. (ecosystems programmes occasional report; no 45.). mapaure, i. & b.m. campbell. 2002. changes in miombo woodland cover in and around sengwa wildlife research area, zimbabwe, in relation to elephants and fire. african journal of ecology 40: 212–219. matthews, w.s., a.e. van wyk, n. van rooyen. & g.a. botha. 2001. vegetation of the tembe elephant park, maputaland, south africa. south african journal of botany 67: 573–594. mcgeogh, m.a., b.j. van rensburg & a. botes. 2002. the verification and application of bioindicators: a case study of dung beetles in a savanna ecosystem. journal of applied ecology 39: 661–672. moll, e.j. 1977. the vegetation of maputaland—a preliminary report of the plant communities and their future conservation status. trees in south africa 29: 31–58. mosugelo, d.k., s.r. moe, s. ringrose & c. nelleman. 2002. vegetation changes during a 36 year period in northern chobe national park, botswana. african journal of ecology 40: 232–240. mueller-dombois, d. & h. ellenberg. 1974. aims & methods of vegetation ecology. new york: john wiley. soil classification working group. 1991. soil classification a taxonomic system for south africa. memoirs of the agricultural natural resources of south africa no. 15. pretoria: department of agricultural development. van rensburg, b.j., m.a. mcgeogh, s.l. chown & a.s. van jaarsveld. 1999. conservation of heterogeneity among dung beetles in the maputaland centre of endemism, south africa. biological conservation 88: 145–153. van rooyen, n. 2002. veld management in the savannas. pp. 571–620. in: bothma, j. du p. (ed.). game ranch management. pretoria: van schaik. van wyk, a.e. 1996. biodiversity of the maputaland centre. pp. 198–207. in: van der maesen, l.j.g., x.m. van den burgt & j.m. van medenbach de rooy (eds.). the biodiversity of african plants. dordrecht: kluwer academic publishers. van wyk, a.e. & g.f. smith. 2001. regions of floristic endemism in southern africa. a review with emphasis on succulents. pretoria: umdaus. issn 0075-6458 29 koedoe 47/1 (2004) gaugris.qxd 2004/04/15 06:28 page 29 filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 page 14 swanepoel.qxd community involvement in the development of an environmental education programme: the tswaing meteorite crater conservation area as a case study c.h. swanepoel, c.p. loubser and e.m.j.c. schaller swanepoel, c.h., c.p. loubser and e.m.j.c. schaller. 2004. community involvement in the development of an environmental education programme: the tswaing meteorite crater conservation area as a case study. koedoe 47(1): 115–123. pretoria. issn 00756458. a key requirement for the relevance of applied research in education is its actual impact on society. a case study was undertaken to determine how research insights could be implemented by involving a local community in the design and implementation of environmental education programmes in their environment. the tswaing meteorite crater conservation area project was undertaken with the active participation of teachers, learners and education officers from the communities living around tswaing, as well as subject specialists. issues which should be considered in the development of similar programmes were also highlighted. key words: environmental education, community development, community involvement, rural development, environmental management, environmental ownership, preservation and utilisation, social sustainability. c.h. swanepoel, institute for educational research, university of south africa, p.o. box 392, unisa, 0003, south africa; c.p. loubser, department of further teacher education, university of south africa; e.m.j.c. schaller, research fellow, institute for educational research, university of south africa. issn 0075-6458 115 koedoe 47/2 (2004) introduction an issue which has often emerged and will probably always surface when research, especially in the human and social sciences, is the subject of discussion, is the way that research results are utilised for the benefit of human existence. this also applies to research regarding environmental education. an enormous expansion of knowledge concerning environmental education has occurred (mosidi 1999). moreover, environmental education literature tends to be loaded with noble statements such as ‘a concern for the state of the environment and a looming environmental crisis [has] resulted in educators starting to teach with the environment as a focus’ (mosidi 1999) and ‘it can be accepted that education had to move beyond teaching in and about the environment’ (viljoen 1993) to ‘include education for the environment’ (fien 1993; huckle 1991). in other words, knowledge, skills and attitudes must be simultaneously addressed in order to increase the community’s awareness of the environment, to initiate actions to protect the environment in its totality and to achieve sustainable development within communities (stevenson 1993; republic of south africa 1998). however, it is doubtful whether these ideals are actually realised in the daily life of citizens. sustainability education in european primary schools (seeps 1998) has, for example, pointed out that environmental education has failed in many cases and that the environmental crisis has worsened. if the success of the implementation of environmental education is being questioned, then environmental educators should ask themselves what has been done with the knowledge and expertise gained in the past through international and local investigations. a further question deals with how this swanepoel.qxd 2004/10/05 11:30 page 115 knowledge and expertise can be imparted successfully to communities of various age groups, contexts and cultures. in this regard, smyth (1995) mentioned that environmental education calls for increased participation by communities in the management and development of the environment. in south africa, society and the government are putting increasing pressure on environmental educators to transform this information into viable programmes within their respective communities or places of work (republic of south africa 1995; neluvhalani & mphaphuli 2000). therefore, as stated in the white paper for education and training, there is an urgent need for all groups to become involved in the design, planning and implementation of environmental education programmes and projects (republic of south africa 1998). with reference to the aforementioned exposition, the following research question was formulated: how can environmental education knowledge be implemented by involving a local community in the planning and design of its own environmental education programmes and what are the important issues in this regard? method the aim of the research was to address the research problem by conducting a case study in a particular local community. this was done by identifying a community (the cluster of communities living around the tswaing meteorite crater in soshanguve) and a specific environmental opportunity which could be utilised in the development of environmental education programmes; providing the community with an opportunity to become involved in developing outcomes-specific programmes for learners; and helping the community to discover the totality of their environment. identifying a community there is no clear-cut definition for the term community. many definitions are structured in an ecological sense and refer to animal communities, plant communities, etc. defining the notion of ‘human’ communities is different and the specific definition depends on the context of a community. khan (1994) gives an example in which a community was defined as the poor black people in an area in the united states of america and indicated that this group could also be a subgroup of other communities: town, koedoe 47/2 (2004) 116 issn 0075-6458 fig.1. tswaing meteorite crater's location in south africa. country, state, black people, poor people and so forth. people tend to live differently in similar environments and differently at different times in the same environment (bird 1989). it is, therefore, important for the researcher, facilitator and/or the community worker to define the target community. once this has been done, such a person should be prepared to work with all the members/role players in the community (khan 1994). for the purpose of this research, the communities living around the tswaing meteorite crater conservation area in soshanguve were identified as the target community and are referred to as the tswaing community. this community comprises areas of winterveld, soshanguve and hammanskraal. although these areas belong to the same larger geographic area and are characterised by poor living conditions, the inhabitants are diverse in the sense that they speak several different south african languages (english, afrikaans, isizulu, isixhosa, sesotho, setswana, etc.), each with its own cultural identity. the name soshanguve, for example, originated from the language groups swanepoel.qxd 2004/10/05 11:30 page 116 sotho, shangaan, nguni and venda. in this area, the environment and the use of natural resources and agriculture play an important role and influences the way of life. many communities depend only on the environment for their existence. the houses are built from materials derived from the environment, food is obtained from the environment and the inhabitants work as farmers in the immediate environment. this relationship with the environment has been the topic of many projects. furthermore, it has been established that poverty remains an alarming factor in other developing countries where this kind of relationship is dominant (loubser 1998). therefore, the tswaing meteorite crater conservation area was identified with the aim of establishing how the environment could be utilised to the benefit of the surrounding communities. the research reported in this particular article focussed on adults and schools within communities in the larger tswaing meteorite crater environment. identifying an environmental opportunity the second phase in the research process was to identify an environmental opportunity which could be utilised in the development of environmental education programmes. the tswaing meteorite crater conservation area was identified as suitable for this purpose. the crater is a young, well-preserved impact crater within a largely uniform geological setting. it is of considerable international interest as a focus of future scientific research, as the sediment in the crater has a 160 000 year record of climatic and environmental change (partridge 1993). it is a unique environmental phenomenon since the wellpreserved crater is the only example of a lake occupying a meteorite impact crater in southern africa. zooplankton and fish are prevented from colonising the lake because of nocturnal deoxygenation. the few species of invertebrates found in the lake (nematode worms and brine flies) are confined to the surface. several species of waders and water-fowl are found around the edges of the lake. a single species of salt-tolerant sedge grows in and along the lowsalinity feeder stream from the artesian spring. tswaing is situated in an ecological region known as the sourish-mixed bushveld. for many years this ecological system had a bad reputation for its poor agricultural potential, mainly due to the great variation in the palatability of grass types and the occurrence of toxic plants. the result was that areas with more palatable grasses and less toxic plants were inevitably overgrazed. overgrazing and the elimination of the tastier grass types increased the occurrence of more sour and less palatable grasses (http://www.nfi.org.za/tswaing). the surroundings around tswaing were probably periodically inhabited by early stone age people 500 000 years ago. at tswaing stone artefacts from the late stone age, brought there by the ancestors of the san (bushman) people who lived from 30 000 to 2 000 years ago, have been found. the first farmers using iron implements, ancestors of the present indigenous people, migrated to south africa about 1 750 years ago. the migration by iron age communities to the south african highveld (where tswaing is located) took place in more recent times, about 800 to 900 years ago. shards from decorated clay vessels, found on the crater floor, indicate that the latter were early sothoor tswana-speaking communities, known as the moloko. similar pot fragments have been found in the waterberg mountain range, about 60 km north of tswaing. soon after the departure of the matebele in the early 1830s, the tswaing area was claimed by another group of settlers, the voortrekkers. they were white afrikaner farmers, who had emigrated from the eastern cape region to the present-day kwazulu-natal and the interior. the voortrekkers regarded the land abandoned by the matabele, whom they eventually defeated in 1837, as land available for permanent colonial settlement. in the 1850s these farmers divided the region north of the magaliesberg into huge farms. the crater became the centre of one such farm, aptly named zoutpan (dutch for salt pan). during the 1890s, when zoutpan was surveyed properly for the first time, its northeastern quarter became a separate farm, named uitspan (outspan, or resting-place). during the 19th century, tswaing was one of the main sources of natural salt for the region north of pretoria, which was established in 1855 as the capital of the transvaal boer republic. because of its economic importance, zoutpan became state property in 1876 (http://www.nfi.org.za/tswaing). more lately the crater has been developed as an enviro-museum, the first of its kind in south africa (moolman & de jongh 1995). a trail was also previously developed in the crater with the aim of affording tourists the opportunity to explore it. the environmental education potential of tswaing meteorite crater lies in its potential as a popular environmental education community resource centre providing opportunities for recreation and environmental education. it could also generate revenue for the local community through tourism (de jong 1995). to do justice, however, to the ideal of ‘for the people, by the people’, community participation and empowerment are key elements in the development and management of the tswaing meteorite crater opportunity (de jong 1995). issn 0075-6458 117 koedoe 47/2 (2004) swanepoel.qxd 2004/10/05 11:30 page 117 identifying an environmental education programme the tswaing meteorite crater trail provides an opportunity to involve diverse communities living around the crater in the development of relevant environmental education programmes. it was proposed that contextualised and relevant environmental education could be presented to suit the educational, social, economical, cultural, physical and political environments of the community in which it would operate. this could be done by the development of an environmental education programme, based on the crater trail, which could be used by teachers, environmental education officers and tour guides. the development of the aforementioned programme would provide the community with an opportunity to become involved in developing outcomes-specific programmes for learners. this entails incorporating personal experience, participatory and discovery activities to enable the community to experience the tswaing meteorite crater environment in its multiplicity, not only as a natural resource to be utilised, but also to be nurtured and developed responsibly and sustainably. an attempt would also be made to provide opportunities for learners and the community to broaden their sense of understanding and appreciation of the broader environment in which a unique natural resource, like tswaing meteorite crater, exists. results situation analysis when one considers getting a local community involved in a joint venture, it is important to ascertain how the community chooses to define participation, because this will determine which level of participation will satisfy the environmental education goals of the programme. it is necessary, therefore, to learn more about the community concerned beforehand. according to swanepoel (1989), existing obstacles concerning community development, such as illiteracy, customs and traditions, dependency and apathy could be encountered. swanepoel (1989) further states that the very reason for community development acts as an obstacle to successful projects. the poverty of the people and their lack of self-sufficiency and selfreliance make it very difficult to involve them in development efforts. the following aspects were considered as general guidelines during the situation analysis: the first decision of a person who wants to work in a community is to decide how and when to enter the community. the first entrance into a community may be the last opportunity to gain its trust. initial mistakes might influence the researcher, facilitator or community worker permanently. most community workers do not have a specific choice of community because they work for an organisation. they should, therefore, work extensively at establishing their own acceptability in the community and the image of their organisation. if the community worker has a choice of community, the strategy of approaching the community should be carefully planned beforehand. it is useful if the community worker has some existing link with the community. it is also extremely helpful to stay in the community itself. it is valuable for community workers to choose their own initial contacts, but sometimes the organisation they work for prescribes these contacts. community development can be a learning process only if people really participate (swanepoel 1989). the community must be made aware of the exact purpose of the project: which needs should be addressed and who should address them? community contact at tswaing had already been established through community forums and meetings. these structures were operating functionally and positively and the environmental education project could make use of them rather than attempting to establish new links and associations with the communities. using these structures, the community was made aware of the purpose of the project, ascertaining their role in the project and koedoe 47/2 (2004) 118 issn 0075-6458 swanepoel.qxd 2004/10/05 11:30 page 118 the needs it would attempt to address. direct participants of the community included forum members, tour guides, teachers and learners, tswaing museum staff and construction workers from the local community. the broader "community" was indirectly involved through the utilisation of tswaing facilities and programmes. after having made contact with the community, the next most important step was to assess the community structures. it was necessary to obtain both abstract and physical information. abstract information included aspects such as the attitudes, beliefs, religion, habits and culture of the community. physical structures included the population size, housing facilities, job opportunities, school facilities, transport, etc. swanepoel (1989) also regards the determining of family ties in the community as a very important step, because the family is a basic social structure (abstract structure) in communities. the researchers had to be aware of some of the subtleties of the abstract structures such as political positions to enable them to be fexible. it was useful to be familiar with physical structures to determine which role the researcher could play in the improvement of the community’s physical needs. implementation of initial environmental education project the environmental education project at tswaing meteorite crater was carried out in two main phases and was part of a larger project conducted by the former national cultural history museum. the initial project (co-ordinated by an independent researcher) comprised two elements, namely a teacher’s guide and tourist guides. the teacher’s guide was compiled to give the teacher adequate information about tswaing meteorite crater which had been obtained from previous research reports and articles. it included instructions on how to use the guide, details of the trail and possible activities that could be carried out along a specific section. these included preliminaryand follow-up activities. it also contained safety hints, materials needed for the activities, techniques that could be used on the trail, examples of worksheets and lists of fauna and flora found in the vicinity of the crater. teachers could select activities from the teacher’s guide and design their own programmes for their learners. in conjunction with the compilation of the teacher’s guide, a programme was structured and tour guides from the local communities were trained to assist the teacher in facilitating the school groups during the programme. this created jobs for the local communities, which is one of the main purposes of a community project. to ensure that both abstract and physical information had been included in the manual, the development of the teacher’s guide and the training of the tour guides were done in close consultation with the parents, teachers and learners from the communities, as well as subject specialists. these consultations were done through regular forum meetings and training sessions at tswaing. local inputs shaped both the training phase of the tour guides and the development of the teacher’s guide. three trial groups of learners (grades 1 to 6; 7 to 9; and 10 to 12) traversed the trail assisted by the tour guides. it became clear from this exercise that identifiable results of group dynamics come into play and cultural and language differences become significant influencing factors. different environmental features had different meanings to different groups. the different language groups used different names for the same object and had a variety of folk stories to tell about certain objects. these were incorporated into the project. this exercise helped to enrich the project by lending an indigenous flavour and helped to incorporate abstract information. to cope with groups that could represent up to nine different language groups, individuals in a group were used as interpreters. it became evident from the pilot group that adaptations had to be made regarding walking distance, time spent on activities and making activities suitable for the different grades. after having made all the necessary changes to the project, it was launched in 1996. issn 0075-6458 119 koedoe 47/2 (2004) swanepoel.qxd 2004/10/05 11:30 page 119 a major local community issue soon emerged, namely that 300 learners of different age groups from surrounding schools arrived at tswaing meteorite crater conservation area unprepared for the programmes they were about to experience. these groups had no set outcomes or selected activities. thus, it was very difficult for the limited number of tswaing staff who only had basic environmental education skills, to accommodate these groups effectively. this had to be taken into consideration during the next phase of the research. implementation of second phase of environmental education project at the end of 1996 a more structured approach to the project was adopted. the project manager approached an environmental education specialist from the university of south africa middle 1997 to set up a team of environmental educationists who could help with extending the environmental education programmes at tswaing meteorite crater conservation area. this was subsequently done and representatives from the university of south africa, vista university, the south african college for teacher education and staff of the former national cultural history museum met to start the process. it was emphasised that: all the communities living around the crater should be consulted and included in the development of the extended programmes; all programmes should adhere to the new outcomes-based education system; the crater should form the focal point of the programmes; the entire education system should be covered (all the phases and learning areas); and the programmes would operate along the trail. a chairperson for the educational committee was elected at a subsequent meeting and commissioned to proceed with the development of programmes. the educational committee was extended to include other role players such as the gauteng department of nature conservation, teachers from communities around tswaing, as well as community members. the committee explored the trail at tswaing meteorite crater twice. sixteen special features concerning the trail were identified and grouped together into 10 stations. this was further defined into six possible programmes that could be conducted along sections of the longer trail. small teams, including teachers from local schools and community members, worked on the development of these programmes, keeping in mind the different phases and learning areas of the new outcomes-based curriculum, adaptability and exchangeability. moreover, guides had to be trained to facilitate the programmes. a pilot run of the 57 activities in the six programmes was carried out over four days. on day one the activities were piloted for the foundation phase (grades 1 to 3), day two for the intermediate phase (grades 4 to 6), day three for the senior phase (grades 7 to 9) and day four for grades 10 to 12 which form part of the further education phase of the national qualifications framework. the learners who were used had been invited from schools in the local area, together with their teachers, to ensure that the target community assisted in shaping the programme and establishing a true learning process. the teachers were included to ensure an element of certainty in the groups and to facilitate communication. this piloting exercise was experienced positively by the learners, as well as the teachers. however, it took a while for both groups to become comfortable with the programme presenters, because they were unfamiliar with the people, methods and approaches utilised. however, the pilot groups settled down after a while. the programme developers also learned a great deal and further principles emerged which had to be kept in mind when developing an environmental education programme for a specific community and environment. it became clear that: koedoe 47/2 (2004) 120 issn 0075-6458 swanepoel.qxd 2004/10/05 11:30 page 120 the teachers and learners were not familiar with the concept environmental education; the background knowledge of the teachers and learners were not what the environmental education programme developers had assumed it to be; language and terminology differences existed; time was a crucial factor; the mastering of specific skills required repetition; the environment was perceived mainly as physical; and cross-curricular activities were foreign to them. these points merely serve to reinforce the necessity of doing research and consulting with the community in question prior to developing programmes. projects need to be contextualised socially, politically, culturally, educationally and economically within the environment in which the project is to be conducted. this result was no different from the first phase of the project, but the project developers still redesigned their activities to try and accommodate the above-mentioned aspects. these revised activities have been incorporated into a publication which is currently used by the environmental education officer at tswaing to facilitate different school groups regarding environmental education. issues emerging from the case study the tswaing environmental education project was a case study of an attempt to put environmental education research into practice by involving local communities in developing environmental education programmes in their environment. it was found that in the development of similar programmes a number of issues have to be considered, namely, the target audience (community) of the programme, the training of tour guides, educational officers and teachers, the development of specific stations and the writing of the teacher’s guide. regarding the target audience, it is essential to: keep in mind the way in which the target audience interprets the local and broader environment because of indigenous abstract information which is not known to the developer; use the local community in as many aspects of the programme development as possible in order to ensure that the people participate in the development of the direction and context of the programme; use the local community to help construct the buildings and structures necessary for the project, thus contributing to the well-being of the community; to tell stories recorded on tape and to act as tour guides, and so be involved and be part of the action; keep in mind the indigenous background knowledge of the community; keep in mind the local community’s understanding of environmental education; develop pre-visit activities for the programme users to familiarise them with the purpose of the programme and to obtain information from them which could influence the progress of the programme; develop follow-up activities for programme users to help the developer to assess the success of the activity; and keep in mind language and terminology differences and to incorporate these into the programme. the abovementioned principles correspond with the supporting objectives of goal 5 of the white paper on environmental management (republic of south africa 1998). the facet of training is important for preventing possible misunderstandings between the developer and the community. possible strategies are to: train project developers about the customs, traditions, fears and literacy levels of the local community in which they are about to work; issn 0075-6458 121 koedoe 47/2 (2004) swanepoel.qxd 2004/10/05 11:30 page 121 train local tour guides/educational officers to help facilitate programmes; train local teachers how to use programmes; train staff to understand environmental education and its underlying principles; provide training in outcomes-based education for teachers and tour guides/educational officers; provide extensive background information rather than limited information; and use community leaders to assist with training in order to localise and contextualise the programme. if stations or special features are a facet of a project, the following points should be considered: use members from the local community to do construction work and to build toolboxes and equipment needed at the stations. this assists the community in participating actively in the project and provides them with a welcome income. use the stations which are relevant to the teachers’ requests/outcomes. keep safety aspects in mind. incorporate local information, for example use local persons to tell stories, incorporating local customs and traditions. give stations locally-accepted and understood names rather than number features, as this creates an indigenous and familiar environment. use only one feature rather than too many. provide facilities such as water, recycling and toilet points to meet the physical needs of the community. if a teacher’s guide forms part of a project, the following aspects should be considered: use people from the local community to assist with compiling activities and thus eliminate possible biased interpretations; incorporate local interpretation(s) and terminology into the activities; allow for freedom of interpretation and flexibility when programmes are used by different groups/communities. conclusion according to makumbe (1996), robinson & shallcross (1998) and pausewang et al. (1990), there is a definite need for development in communities in developing countries. knowledge gained from work in these communities could inform more developed countries on issues such as who should participate in programme development, and to which extent. the tswaing meteorite crater case study shed light on this particular statement because of the participation of a diverse group of role players from the local area (teachers, learners, community leaders and other individuals) and the surrounding environment (universities and the former national culture history museum). during the tswaing environmental education project, research insights were put into practice with success to a large extent. it would, however, need to be assessed whether the local communities have benefited through the environmental education programmes in the long term and whether these benefits can actually sustain the tswaing environment, thus satisfying conservation, educational and developmental objectives. through the tswaing research project and its unique physical environment and mixture of diverse communities, certain principles were identified from the two development phases for community-based environmental education programmes which could accomplish the abovementioned. it is envisaged that these principles can be applied to similar programmes in other communities. to understand the society or community one is working with, one needs to understand the space or environment in which the community operates (bird 1989). however, it is important to realise that these principles are flexible and should be adapted to the specific environment and communities where they are being implemented, to make environmental education programmes sufficient and beneficial to the community. koedoe 47/2 (2004) 122 issn 0075-6458 swanepoel.qxd 2004/10/05 11:30 page 122 acknowledgements the former national cultural history museum is acknowledged for making this project possible. the settlements in and around tswaing meteorite crater are thanked for their participation in the project and their contribution to this article. references bird, j. 1989. the changing worlds of geography: a critical guide to concepts and methods. oxford: claredon press. de jong, r.c. 1995. the tswaing crater museum. pretoria: national cultural historical museum. fien, j. 1993. education for sustainable living: an international perspective on environmental education. southern african journal for environmental education 7:20. http://www.nfi.org.za/tswaing. http://www.nfi.org.za/tswaing/map.htm. huckle, j. 1991. education for sustainability: assessing pathways to the future. australian journal of environmental education 7:43–62. khan, s. 1994. how people get power. washington: nasw press. loubser, c.p. 1998. empowering rural communities through environmental education and research: draft report. pretoria: university of south africa. makumbe, j.m. 1996. participatory development. the case of zimbabwe. harare: university of zimbabwe publications. moolman, h.j. & r.c. de jongh. 1995. final report of the research project on: the contribution of site museums to the conservation and the interpretation of the environment with special reference to the tswaing crater museum. internal report. national cultural history museum. pretoria. mosidi, s.m. 1999. environmental education in curriculum 2005: a case study in the northern cape. johannesburg: rand afrikaans university. nelvhalani, e. & s. mphaphuli. 2000. environmental educators networking for professional development: a case study of the northern province, far north region. case study 4. educating for social change. case studies of changing practices in south african tertiary institutions, australian institutional links programme. partridge, t. 1993. environmental data relative to the establishment of a museum at the pretoria salt pan. pretoria: national cultural historical museum library. (ta17.) pausewang, s., f. cheru, s. brune & e. chole. 1990. ethiopia. rural development options. london: zed books ltd. republic of south africa. 1995. white paper for education and training. pretoria: department of education, government printer. republic of south africa. 1998. white paper on environmental management. pretoria: department of environmental affairs and tourism, government printer. 395(18894). robinson, j. & t. shallcross. 1998. the socioecological crisis and education for sustainable living: is an essentialist’s standpoint possible? pp. 23-28. in: laubscher, c.p. proceedings of the international best of both worlds conference at dikhololo, brits, september 1998. pretoria: university of south africa. sustainability education in european primary schools. 1998. creating a better environment in our school. edinburgh: moray house institute. smythe, j.c. 1995. environment and education: a view of a changing scene. environmental education research 1(1):3–20. stevenson, r.b. 1993. becoming compatible: curriculum and environmental thought. journal of environmental education 24(2):4–9. swanepoel, h. 1989. community development. putting plans into action. cape town: juta . viljoen, h.j. 1993. environmental education and open spaces. parke en rekreasiebestuur 51(4): 22–24. issn 0075-6458 123 koedoe 47/2 (2004) swanepoel.qxd 2004/10/05 11:30 page 123 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 slotow & van dyk.qxd ranging of older male elephants introduced to an existing small population without older males: pilanesberg national park r. slotow and g. van dyk slotow, r. and g. van dyk. 2004. ranging of older male elephants introduced to an existing small population without older males: pilanesberg national park. koedoe 47(2): 91–104. pretoria. issn 0075-6458. african elephant have been introduced to small reserves in southern africa as rangelands are converted to game reserves. early introductions were limited to very young individuals (<10 years old), and these populations have manifested abnormal behaviour. this project aimed to rectify this by creating a more normal population age structure. six older (25 to 35 year old) male elephant were introduced from kruger national park to pilanesberg national park from february 1998. these radio-collared individuals were followed intensively. immediately after release, the males demonstrated exploratory behaviour before settling down in very small home ranges. core home range size (50 % kernel) was 3–23 km², while total home range (95 % kernel) was 24–139 km² (total reserve size = 500 km²). there was no indication of homing towards the direction of their origin. during musth, males shifted their spatial use to coincide with that of females, and walked significantly longer distances while in musth. these results imply that males were moving from ‘bull areas’ to join the female groups when they came into musth. the limited range used by males for most of the year has very important implications for managers of small reserves. firstly, large adult males can be maintained on very small reserves, and secondly, the impact of large males on their environment will be extremely heterogeneous, with some areas severely impacted, and others with little or no impact. this study highlights the importance of detailed individual behavioural knowledge of large mammals in understanding their potential ecological contribution to the ecosystem. the consequences of heterogeneous spatial use need to be well modelled for any meaningful management plan of elephant to be generated. key words: loxodonta africana, reintroduction, homing, home range, movement r. slotow , school of life & environmental sciences, university of kwazulu-natal, durban, 4041 republic of south africa, (slotow@ukzn.ac.za); g. van dyk, school of life & environmental sciences, university of kwazulu-natal, durban, 4041 republic of south africa (present address: pilanesberg national park, north west parks and tourism board, p.o. box 1201, mogwase, 0314 republic of south africa). issn 0075-6458 91 koedoe 47/2 (2004) introduction the african elephant loxodonta africana is one of the key components of african savanna. not only do they play a crucial role in the ecosystem (dublin et al. 1990; van de vijver et al. 1999), but they also provide one of the integral parts of sustainable conservation through ecotourism (e.g. brown 1993) and consumptive utilisation (taylor 1993). the ever-expanding transformation of savanna land-use through human settlement is resulting in the isolation of elephants into small populations. furthermore, in south africa, the reclamation of ranch land as game areas has resulted in the reintroduction of elephant to a large number of small, isolated, fenced reserves. elephants potentially modify their environment, and managers of small reserves are faced with the ‘elephant problem’ (caughley 1976), the loss of aesthetically important large trees and potential adverse effects on biodiversity. this is particularly pertinent for older male elephant that regularly push over the larger trees in their range (pers. obs.). slotow & van dyk.qxd 2004/10/05 10:42 page 91 however, elephants do not utilise all of their range equally, and such adverse effects are likely to be heterogeneous. to more effectively model the potential effects of elephant, and thus provide managers with more comprehensive data for which to set stocking rates of small reserves, detailed knowledge of spatial behaviour of elephant is required. an aim of this paper is to describe spatial use by older male elephants in a small reserve, and further to describe variation in use in relation to behaviour such as musth (see poole 1987 for definition), and which will provide a useful contrast with a recent study at addo elephant national park (whitehouse & schoeman 2003). before 1998, all small elephant populations in south africa that were founded through reintroduction lacked older males (> 25 yrs). this was because of technological constraints on moving large animals, and trucks large enough to move males were developed for this introduction project. subsequent to the success of this project a large number of reserves have introduced larger males with mixed success (garai & carr 2001), and it is likely that most of the small populations in southern africa will receive older males in the coming years. the second aim of this paper is to provide a description of the spatial use of elephants that are introduced to an existing population. the main reason for the introduction of these older males was that the younger males were displaying abnormal behaviour in that they were killing white rhino ceratotherium simum while in musth (slotow et al. 2001; slotow & van dyk 2001). the young males were entering musth at an early age, and the older males were introduced from kruger national park in order to suppress musth in these young bulls. this proved successful, and the musth patterns of the young males before and after introduction, and of the older bulls is described elsewhere (slotow et al. 2000). methods pilanesberg national park (25º8' –25º22' s; 26º57'– 27º13' e) is situated in the remains of an extinct volcano. it comprises 500 km² of very hilly savanna terrain. the habitat consists of acacia and broad-leaf bushveld which ranges from thickets to open grassland patches. there is one major river system running southeast through the central part of the park, and a large dam in the centre of the park, with a number of smaller dams scattered throughout the reserve. rainfall is approximately 630 mm p.a., and falls in summer. winters are cold (minimum temperature 1–5 ºc), and summers very hot (mean temperature 28–31 ºc). the park was proclaimed in 1979, and since then, about 6000 individuals from a range of species have been reintroduced to the park. african elephants were introduced from kruger national park in 1981 (13 juvenile males; 5 juvenile females), 1983 (13 juvenile males; 11 juvenile females), 1993 (19 males; 17 females) and, from namibia in 1982 (1 juvenile male; 1 juvenile female). at the time of the introduction, the technology of translocations was such that elephants of a maximum shoulder height of 2 m (8–10 years old) could be translocated live. the elephants that were translocated were orphans from culling operations. in 1982, two 19-year-old females were introduced. these were tame elephants that were returned from the usa to africa, and released into the wild. these two older females assumed the role of matriarchs in the pilanesberg herds. two young male elephants were introduced from mabula lodge in 1992 and were removed in march 1993. the first young elephant was born in 1989, indicating that males were breeding at the age of about 18 years old. there were a total of 93 elephant in the population at the start of this study in early 1998, with 17 individually recognizable independent males. these were males that had left the female groups, and were consistently alone or with other males. they ranged in age from 18 to 25 years old. six older male elephants were introduced from kruger national park in early 1998 (table 1). these were captured separately and moved in pairs. each pair was immobilised in the morning from a helicopter, and transported in a specially built trailer to be released into a holding facility in pilanesberg on the evening of the same day. each pair was held for one to three nights and then released by opening the gate in the morning, and the animals moved out of the facility that night. the elephants had vocal contact with the existing population, but no physical contact because the fence of the facility was electrikoedoe 47/2 (2004) 92 issn 0075-6458 slotow & van dyk.qxd 2004/10/05 10:42 page 92 issn 0075-6458 93 koedoe 47/2 (2004) fied inside and out. the park border fence is also electrified and provides a barrier to elephants. all independent male and adult female elephants in pilanesberg are identified through unique ear, trunk, tail and tusk characteristics. all six introduced males were fitted with vhf radio-collars. a dedicated assistant monitored the elephants from introduction until the end of may 1999. from november 1998, a second assistant was working on the rhino population, and included additional sighting information. from june 1999, the latter assistant monitored the elephants, but at a lesser intensity than previously. in july 1998, december 1998 and july 1999 volunteer undergraduates from the university of natal provided additional sightings. several tourist guides driving open viewers in the park and park staff also provided additional information. at each sighting elephants were identified using a master file, and the following information was recorded: date, time, grid location (500 m x 500 m grid), gps location (without correction), ids, total group size, number males, number females, number young, habitat type (one of 22 types). in addition, time budgets were recorded where possible. pilanesberg consists of bushveld, making it difficult to observe animals, and most observations were made from a vehicle from the road. there is an extensive road network in pilanesberg (fig.1). however, there are two large wilderness areas that are traversed by few tracks, and some areas are completely inaccessible. the larger one is in the northwest and the other in the south-east of the reserve. elephants did ‘disappear’ into these ‘black holes’ for varying periods. until may 1999, the assistant attempted to get a daily location on each of the six introduced bulls using telemetry. from june 1999, sightings became more opportunistic, and the frequency of sighting each individual decreased. during the first year, the assistant would walk in on collared animals in order to get group information, but this effort was reduced as the study proceeded. data analysis data were processed in excel (microsoft), and imported to arcview (esri). maps were produced covering six-month periods, and each record was checked on a master spreadsheet against the maps for accuracy. suspected errors and duplicate records were deleted. observations of the same animal on the same day were retained if the records were early morning (before 10:00) or late afternoon (after 16:00) or if there had been a substantial change in location (at least 1 km). data were captured in latitude and longitude, and projected into utm using wgs84 projection. the master file is lodged at the university of kwazulunatal and with the database manager, ecological services, north west parks & tourism board. the master file was processed into separate files that contained selected periods. these periods were selected separately for each individual, and reflected changes in spatial use (settled in one area or moving widely across reserve), or behaviour (in or out of musth, with or without female groups). an individual was considered as settled when the locations could be contained within a single kernel during a kernel analysis, and as moving when the locations ranged outside of the kernel, and were not repeatedly in the same place. although this was a subjective assessment, whether an individual should be included in a specific category was unambiguous in each case. each of these files was then processed in two ways using the arcview extension called spatial movement analysis (hooge & eichenlaub 1997). firstly, if the elephants were moving widely, a sequential route was produced using the ‘create polyline from point file function’. secondly, kernel home ranges were produced using the ‘kernel home range function (with h = 500 m)’ to indicate ranges. we present 95 % (assumed to indicate greatest range extent) and 50 % (assumed to indicate core range area) probability kernels (worton 1989). overlaps in areas were calculated by overlaying maps of the prior behaviour with the selected behaviour, and then dividing the area of overlap by the prior area (multiplied by 100), to give a percentage overlap area. male ranges before and during musth were contrasted with ranges for females. during the period of the study there were three groups of females. the matriarch of the one group of about 16 animals was collared, and the other two groups regularly joined to form a single large group. male #33 spent most of his time with females in this large group. females were therefore sighted as often as the males. we generated home ranges for the females for the periods whenever a male was in musth and assessed overlaps in the same way as above. results each individual was located on average 336 times during the period of the study (table 1). almost two thirds of those sightings were up to the end of 1998 (average 235 sightings over about 330 days) giving an average sighting rate per elephant of greater slotow & van dyk.qxd 2004/10/05 10:42 page 93 koedoe 47/2 (2004) 94 issn 0075-6458 fig. 1. ranging patterns of older male elephants on introduction to pilanesberg national park. (a) male #30: route from introduction until 7 march 1998 and kernel range of first settlement from 8 to 24 march 1998. (b) male #30: route from 26 march to 29 april 1998 and kernel range of second settlement period from 30 april to 31 may 1998. (c) male #30: kernel range 1 to 30 june 1998, solid line showing route from 1 to 13 july 1998, dashed line showing route from 25 july to 10 august. (d) male #31: kernel range from 17 february to 3 march 1998, dashed line showing route from introduction to 17 february 1998, solid line showing route from 3 march to 13 march 1998. (e) male #31: kernel range from 14 march to 1 april 1998, line showing route from 2 april to 14 april 1998. (f) male #32: initial range from introduction to 15 april 1998. (g) male #33: initial range from introduction to 2 april 1998. (h) males #34 & #35: initial range from introduction until 24 april 1998. see appendix a for additional information. slotow & van dyk.qxd 2004/10/05 10:42 page 94 than 2 days in every three for the calendar year 1998. the sightability varied among individuals. we calculated the proportion of all sightings that were made on the same or subsequent day, i.e., within 36 hours of each other as 0.76 for male #30, but this dropped to 0.51 of 284 sightings for male #33 (table 1). there were periods when the elephant were out of sight and others when they were regularly seen. for example, male #35 was accessible over a longer period than male #30, but male #30 was seen more often while he was accessible. details of movements over selected periods for each male are presented in appendix a. initial period following introduction all of the introduced males displayed some form of exploratory movement before settling into restricted areas. there was however variation both in the extent, and the timing of the explorations. male #30 showed the widest initial movement (fig. 1a line) ranging across 2/3 of the reserve over a period of 28 days. male #30 then settled in the southwestern part of the park for the next 16 days (fig 1a kernels). following this there was a further exploration (fig. 1b line) and then male #30 settled in a compact home range in the centre of the reserve (fig. 1b kernels). male #31 was slightly injured in the left front leg during the translocation process. this may have effected his initial movement. male #31 showed a much shorter exploration initially (fig. 1d -dashed line), and settled in a compact range in thickets in the north (fig 1d kernels). a second short exploration (fig 1d solid line) was followed by a return to the north where male #31 settled in a compact home range until 1 april 1998 (fig. 1e kernels). a longer exploration occurred from 1 april to 14 april 1998 when male #31 visited the southeast of the reserve (fig 1e solid line). male #32 showed intermediate exploration, visiting about half of the reserve, before settling in a large range in the centre of the reserve (fig. 1f). male #33 showed the least exploration (fig. 1g) and then joined up with the largest female group, and ranged with them. males #34 and #35 consistently moved together until 25 april 1998. they showed extensive ranging (fig. 1h) before settling in the centre of the park (fig. 2m). none of the elephants showed a tendency to move to, or to settle in the direction of their origin in the kruger national park. the first movement in all cases was in a direction issn 0075-6458 95 koedoe 47/2 (2004) table 1 physical characteristics, release dates, and sample sizes of six large male elephants introduced to pilanesberg national park from kruger national park percent of sightings on: id release est. total number number of same subsequent same or date age of sightings sightings to day day subsequent (1998) end 1998 day 30 7 feb 33 362 274 21 55 76 31 7 feb 29 285 205 15 39 54 32 22 mar 35 335 242 19 44 63 33 22 mar 25 284 192 16 35 51 34 13 mar 31 397 264 20 43 63 35 13 mar 32 353 233 17 42 59 mean 336 235 18 43 61 slotow & van dyk.qxd 2004/10/05 10:42 page 95 koedoe 47/2 (2004) 96 issn 0075-6458 fig. 2. contrast of ranging patterns before and during musth for older bulls introduced to pilanesberg national park. (a) male #30 pre-musth: 4 jan to 17 dec 1999. (b) male #30 musth: 24 dec 1999 to 28 feb 2000. (c) male #31 pre-musth 1: 9 aug to 2 september 1998. (d) male #31 musth 1: 11 sep to 2 nov 1998. (e) male #31 pre-musth 2: 3 to 24 nov 1998. (f) male #31 musth 2: 25 nov 1998 to 4 jan 1999. (g) male #31 premusth 3: 2 mar to 6 oct 1999. (h) male #31 musth 3: 18 oct to 23 nov 1999. (i) male #32 pre-musth 1: 16 apr to 25 nov 1998. (j) male #32 musth 1: 28 nov 1998 to 10 jan 1999. (k) male #32 pre-musth 2: 11 jan to 13 dec 1999. (l) male #32 musth 2: 16 dec 1999 to 16 feb 2000. (m) male #34 pre-musth 1: 26 apr to 5 dec 1998. (n) male #34 musth 1: 7 dec 1998 to 28 jan 1999. (o) male #34 pre-musth 2: 23 jun to 13 dec 1999. (p) male #34 musth 2: 16 dec 1999 to 7 feb 2000. (q) male #35 pre-musth 1: 17 nov 1998 to 1 apr 1999. (r) male #35 musth 1: 10 apr to 20 aug 1999. (s) male #35 pre-musth 2: 21 aug to 19 dec 1999. (t) male #35 musth 2: 10 jan to 7 feb 2000. slotow & van dyk.qxd 2004/10/05 10:42 page 96 other than the kruger national park (situated due east). we calculated the distance moved from observations made on two sequential days. five of the six males showed substantially longer movement during the initial period relative to the subsequent period (table 2). this result was marginally non-significant (wilcoxon test: z = -1.787, n = 6, p = 0.074). this test has extremely low power, and is most probably a biologically significant result. given that all males showed some initial exploration, we interpret the increased movement as extra distances moved in association with exploration. for male #30 there was a substantial difference in the distance moved between observations on subsequent days when contrasting the period when settled (mean = 2.6 km) versus those during when he was moving (3.4 km) (fig 1a, b, c; appendix a). these differences in distance moved per day were marginally non-significant (wilcoxon paired signed ranks test: z = -1.84, n = 4 paired periods, movement period paired with subsequent settled period; p = 0.066). a similar contrast for male #31 indicated the same pattern, but with non-significant differences (mean settled = 2.4 km; mean ranging = 3.1 km; wilcoxon test: z = -0.7, p = 0.47; appendix a) comparable data for the other individuals were not available because they did not show such clearly dichotomous (settled vs nonsettled) behaviour (see below). subsequent ranging behaviour all of the males settled into restricted ranges within the reserve. they spent extended periods in a local area, and then went ‘walkabout’ moving extensively, followed by settlement into another local area. there may have been some shift in the local areas. after the initial movements and settled periods detailed above, from august 1998 through to the end of 1999 male #30 remained in a restricted range in the centre of the reserve (fig. 2a). the major change in ranging patterns came at the end of 1999, when #30 entered musth, and moved into the northeastern section of the park, which is issn 0075-6458 97 koedoe 47/2 (2004) table 2 movement of older adult male elephant introduced from kruger park to pilanesberg. distances are calculated as straight-line distance between points of observation on two sequential days separated by a night period (except first values column) average distance (km) moved between sightings id on the on during during during when with when not same day1 subsequent initial non-musth musth females with days² period periods periods females 30 2.0 2.9 5.4 2.3 4.1 31 1.7 3.4 3.8 3.1 5.5 32 2.0 3.5 3.4 2.7 4.4 33 1.7 4.5 5.5 4.3 4.5 34 2.5 4.1 3.2 3.7 5.2 35 2.7 4.3 3.7 3.3 7.1 mean 2.1 3.8 4.2 3.0 5.3 data are averages of average values for all separate periods for each individual (appendix a). 1 distance between two points on the same day separated by at least 6 h and 1 km (see methods). ² distance between two points on subsequent days. slotow & van dyk.qxd 2004/10/05 10:42 page 97 fig. 3. contrast of spatial use before and during musth. we present area (km²) of (a) core range (50 % kernel), and (b) overall range (95% kernel). we also present percent overlap of the range during musth with that before musth as numbers above each data-pair. non-musth behaviour. the core range (50% kernel) sizes of males in musth were larger than when they were not in musth (fig. 3a). this difference was not statistically significant (wilcoxon paired signed ranks test: z = -1.753: n = 5 males with values averaged when more than one musth period occurred, p = 0.08). given the small samples size of this test, this is probably a biologically significant result. there was no significant difference in the area of overall range of males when in musth relative to the period before musth (fig. 3b: wilcoxon test: z = -0.94: n = 5, p = 0.345). (a) (b) where the largest female groups ranged (fig. 2b). following initial ranging described above, male #31 settled into a restricted range in the northeast of the park (fig. 2c). male #31 did venture out for 2 weeks from 25 july to 8 august 1998 and again from 4 january to 24 february 1999 (appendix a). while not in musth, male #31 spent less time in the centre of the reserve (figs. 2c, 2e, 2g) than the other males (figs. 2a, 2i, 2k, 2m, 2o, 2q, 2s). after an initial foray, male #32 settled into a restricted home range in the centre of the park (fig. 2i). male #32 range while in musth was more extensive, and included the northeast that he had previously not visited (fig. 2j). following his first musth period male #32 returned to a core range in the centre of the reserve, in a similar area to that before musth (fig. 2k). during this period his core range was restricted to only 7.1 km² (50% kernel). during male #32’s second musth period his range did not shift from the centre of the reserve (fig. 2l). male #32 may have been excluded from the northeast by male #30 who was also in musth at that time (appendix a), and spending time in the northeast (fig. 2b). after a very extensive ranging, male #34 settled into a restricted range in the centre of the park (fig. 2m). male #34’s range increased during his first musth period to include the wilderness area to the northwest, where the female groups were located at that time (fig. 2n). after musth, male 34 returned to the centre of the park (fig. 2o), while during his second musth period, the range shifted to the female herds in the northeast (fig. 2p). male #35 spent a lot of time associated with male #34, and also settled in the centre of the reserve during non-musth periods (fig. 2q, s). while in musth, male #35’s range shifted to the northeast (fig. 2r, t). behaviour during musth periods the five males that entered musth (all excluding male #33) showed different behaviour when in musth relative to their koedoe 47/2 (2004) 98 issn 0075-6458 slotow & van dyk.qxd 2004/10/05 10:42 page 98 during musth, the older introduced males shifted their range use, moving into areas that they did not normally frequent. this is indicated by the low overlap in core ranges when in musth relative to out of musth (fig. 3). the median percent overlap of core range (50 % kernel) was only 4 %, while that of overall range was also relatively low at 23 %. this range shift was primarily in response to the ranging patterns of the female groups. we illustrate this shift by contrasting the overlap of male ranges with female ranges for the period before musth and after musth (fig. 4). there was a statistically significant increase in the percentage overlap of core range (50 % kernel) of male range with female range from pre-musth to musth periods (wilcoxon paired signed ranks test: z = -2.023, n = 5 males, p = 0.043). there was no significant difference in the degree of overlap of overall (95 % kernel) male range with female range from premusth to musth periods (wilcoxon paired signed ranks test: z = -0.135, n = 5 males, p = 0.893). males walked significantly longer distances when in musth relative to the period before musth (fig. 5: wilcoxon paired signed ranks test: z = -2.023, n = 5 males, p = 0.043). this increased distance is probably a consequence of their association with females that are moving longer distances, or from moving between female groups, than any causal factor among the males themselves. male #33 showed different behaviour because of the close association with the largest breeding herd. male #33 was the youngest of the introduced males (table 1), and had not yet entered musth by february 2001. discussion this introduction was successful in that it met the objective of reducing the mortality of rhino through suppression of musth in younger males (slotow et al. 2000). this project was also the first to translocate adult male elephant (over the age of 25 years old), issn 0075-6458 99 koedoe 47/2 (2004) fig. 4. association of older introduced males with females before and during musth. we present overlap of ranges of males with those of females over the same period, and present percent overlap of their ranges during musth with those before musth for (a) core range (50 % kernel) and (b) overall range (95 % kernel). fig. 5. contrast of distance moved (km) before and during musth. slotow & van dyk.qxd 2004/10/05 10:42 page 99 koedoe 47/2 (2004) 100 issn 0075-6458 and was successful in that the introduced animals settled well into the park. that there were no breakouts is in contrast to subsequent introductions to other reserves where about half of introductions have resulted in a breakout (garai & carr 2001). the lack of breakouts is not related to the relatively large size of pilanesberg, since all males approached the boundary fence soon after introduction, and some spent extended periods in the area adjacent to the fence. the behaviour of the males following introduction is indicative of the well developed social structure, and high levels of intelligence of elephant. all individuals exhibited some exploratory behaviour, and all except male #31 immediately linked up with at least one other male. in some cases, these were the existing resident elephants. although the males appeared to settle rapidly into the reserve, their musth patterns may indicate that they were stressed by their translocation. the soonest musth was after six months (male #31), but male #33 had not entered musth by the end of 2001 (three years after introduction). male #30 took almost 2 years to enter musth (see slotow et al. 2000 for details of musth periods). although we do not know their previous musth history, as the oldest elephants in the reserve, they should not have been constrained from entering musth, even if only for short periods, so we believe that the delay in entering musth must have been from an environmental stressor such as translocation to a new environment. the elephant used a relatively small home range (mean 95 % kernel = 99.7 km² for non-musth periods). this is especially evident when considering the core range of the animals (mean 50 % kernel = 8.4 km² for non-musth periods). there was no change in the total range size during musth, but there was a small increase in core range size (to 12.1 km²). these range sizes are much smaller than those documented (157–465 km²) for male elephant from the nearest comparable population (reserves adjacent to the kruger national park, i.e., their origin population) (de villiers & kok 1997). hall-martin (1984) gives values of 436 km² for female elephants in kruger national park). the ranges observed in pilanesberg are comparable to some other areas, such as lake manyara (57 km²) and the zambezi valley (156 km²) (dunham 1986). addo elephant national park (aenp) is a fenced area of 103 km² and male elephants there had a mean 100 % kernel range of 52.8 km² (whitehouse & schoeman 2003). at aenp, the core 50 % range of non-musth males was 8.4 km² (whitehouse & schoeman 2003), which is exactly the same as observed by us in pilanesberg. this coincidence may indicate a lower-limit requirement by male elephant, which is unlikely to be related to water (widespread in both reserves) or food (addo has a much higher food quality than pilanesberg (pers. obs.), and see references in whitehouse & schoeman 2003). thouless (1996) suggested that range size is correlated with rainfall, with smaller ranges at higher rainfall. the rainfall of pilanesberg is in fact within the range of the kruger national park, and according to this hypothesis we would predict a range several times larger than manifested (bearing in mind that these animals, in fact, originated from kruger national park). note that elephant home ranges are in the 1000s km² in some populations (lindeque & lindeque 1991; demeke & bekele 2000). the introduced elephants moved, on average, 2.1 km between sightings on the same day. in addo, the most frequently observed daily movement was in all cases <2.4 km (fig. 4 in whitehouse & schoeman 2003). this is less than half the distance noted for elephant in the sengwa valley area of zimbabwe (guy 1976), and much lower than the 14.1 km traveled per day by male elephants in the area adjacent to kruger national park —calculated by dividing the distance traveled by the number of days between observations, i.e., more comparable to our measurements between days (de villiers & kok 1997). continuous studies of elephant indicate rates of movement of about 0.5 km/h (wyatt & eltringham 1974). the disparity in these figures implies that the animals in pilanesberg could be backtracking on themselves, rather than moving in straight lines slotow & van dyk.qxd 2004/10/05 10:42 page 100 issn 0075-6458 101 koedoe 47/2 (2004) over longer distances. this would indeed result in the relatively small ranges of the introduced pilanesberg animals. note that water is abundant in pilanesberg, with the whole of the park being within 2.5 km of a source of water. the lack of a need for water may result in the shorted distances traveled. the range sizes are in fact within the lower range of those from northern botswana, specifically those closest to water (verlinden & gavor 1998). there is a very strong social influence on male behaviour, with the males changing their range and distances moved when they enter musth. essentially males are joining the female groups, as observed in addo (whitehouse & schoeman 2003), which results in a shift in spatial use and an increase in distances moved. this result has an important implication. even in areas as small as pilanesberg national park, the male elephant are spending most of their time in ‘bull areas’, and then move from these to be with the females when in musth. this is an interesting result, because there is a lot of overlap in non-musth male and female ranges. in other words, the areas are not spatially exclusive. however, the areas are temporally exclusive. these results are encouraging because they indicate that even in areas as small as pilanesberg national park, there is the potential for a normal elephant social network to be established. management implications each of the large males are thus using about 20 % of the reserve for their total range, but only 2 % of the reserve for their core range. this implies that large elephant may well be maintained on very small reserves. the small ranges used by these males indicate that vegetation use is going to be extremely heterogeneous, particularly as males associate with each other in small groups, and stocking rates should not be calculated on averages across reserves, but rather on spatial usage patterns. this implies a strong need for research on spatial use by elephant on each reserve where they occur in order to sensibly model their potential ecological impact. ‘bull areas’ exist even on small reserves, which imply a higher probability of encounters between large males in those areas. knowledge of bull areas will allow infrastructure planning to enhance tourism sightings (place lodge in bull area), and reduce destruction of materials (place machinery outside bull area). managers and planners of small reserves must aim to provide the necessary environmental characteristics that encourage the separation of male and female elephant into their natural social system. further study is necessary to elucidate the features and factors that govern spatial and temporal separation of home ranges of nonmusth male and female elephant. this study emphasises the importance of detailed study of individually identified large mammals in fundamental understanding of key management issues, and any management plan for elephant should incorporate such study for effective adaptive management to be implemented. acknowledgements this paper is dedicated to the memory of andre klocke, who was killed by an elephant in botswana in 2000. we thank andre klocke and donald sutton for collecting most of the data used in this paper. we thank also all the others who contributed observations. this study was funded by pretoria portland cement, distell (amarula), total petrolium, the south african broadcasting corporation (programme 50/50), the national research foundation, the university of natal, and north west parks and tourism board. references brown, g. 1993. the viewing value of elephants. pp. 146-156. in: barber, e.b. (ed.) economics and ecology: new frontiers and sustainable development. london: chapman & hall. caughley, g. 1976. the elephant problem—an alternative hypothesis. east african wildlife journal 14: 265-283. cumming, d.h.m., m.b. fenton, i.l. rautenbach, r.d. taylor, g.s. cumming, m.s. cumming, slotow & van dyk.qxd 2004/10/05 10:42 page 101 j.m. dunlop, a.g. ford, m.d. hovorka, d.s. johnston, m. kalcounis, z. mahlangu & c.v.r. portfors. 1997. elephants, woodlands and biodiversity in southern africa. south african journal of science 93: 231-236. demeke, y. & a. bekele. 2000. study on the elephants of mago national park, ethiopia. pachyderm 28: 32-43. de villiers, p.a. & o.b. kok. 1997. home ranges, association and related aspects of elephants in the eastern transvaal lowveld. african journal of ecology 35: 224-236. dublin, h.t., a.r.e. sinclair & j.mcglade. 1990. elephants and fire as causes of multiple stable states in the serengeti-mara woodlands. journal of animal ecology 59: 1147-1164. dunham, k.m. 1986. movements of elephant cows in the unflooded middle zambezi valley, zimbabwe. african journal of ecology 24: 287-291. garai, m.e. & r.d. carr. 2001. unsuccessful introductions of adult elephant bulls to confined areas in south africa. pachyderm 31: 52-57. guy, p.r. 1976. diurnal activity patterns of elephant in the sengwa area, rhodesia. east african wildlife journal 14: 285-295. hall-martin, a.j. 1984. conservation and management of elephants in the kruger national park, south africa. pp. 104-118. in: cumming, d.h.m & p. jackson (eds.). the status and conservation of africa's elephants. proceedings of the joint meeting of iucn/ssc african elephant and african rhino specialist groups, 30 july 7 august, 1981, hwange safari lodge, zimbabwe. gland: iucn. hooge, p.n. & b. eichenlaub. 1997. animal movement extension to arcview, version 1.1. anchorage, alaska: alaska biological science center, u.s. geological survey. lindeque, m. & p.m. lindeque. 1991. satellite tracking of elephants in northern namibia. african journal of ecology 29: 196-206. poole, j.h. 1987. rutting behaviour in african elephants: the phenomenon of musth. behaviour 102: 283-316. slotow, r., d. balfour & o. howison. 2001. killing of black and white rhinoceroses by african elephants in hluhluwe-umfolozi park, south africa. pachyderm 31: 14-20. slotow, r., g. van dyk, j. poole, b. page & a. klocke. 2000. older bull elephants control young males. nature 408: 425-426. slotow, r. & g. van dyk. 2001. role of delinquent young ‘orphan’ male elephants in high mortality of white rhinoceros in pilanesberg national park, south africa. koedoe 44: 85-94. taylor, r.d. 1993. elephant management in nyaminyami district, zimbabwe: turning a liability into an asset. pachyderm 17: 19-29. thouless, c.r. 1996. home ranges and social organization of female elephants in northern kenya. african journal of ecology 34: 284-297. van de vijver, c.a.d.m., c.a. foley & h. olff.1999. changes in the woody component of an east african savanna during 25 years. journal of tropical ecology 15: 545-564. verlinden, a. & i.k.n. gavor. 1998. satellite tracking of elephants in northern botswana. african journal of ecology 36: 105-116. whitehouse, a. m. & d.s. schoeman. 2003. ranging behaviour of elephants within a small fenced area in addo elephant national park, south africa. african zoology 38: 95-108. worton, b.j. 1989. kernel methods for estimating the utilization distribution in home-range studies. ecology 70: 164-168. wyatt, j.r. & s.k. eltringham. 1974. the daily activity of the elephant in rwenzori national park, uganda. east african wildlife journal 12: 273-289. koedoe 47/2 (2004) 102 issn 0075-6458 slotow & van dyk.qxd 2004/10/05 10:42 page 102 issn 0075-6458 103 koedoe 47/2 (2004) appendix a breakdown of distances moved (km) by individuals during different periods depending on each individual's state. sample sizes are number of observations. same day refers to distances between two points on the same day. subsequent day refers to distances between two points on subsequent days. distances are straight line distances averaged for same day or subsequent day within each period. these averages were then averaged for each state for analysis and interpretation. ranges are only illustrated for periods that were analysed same day next day core total id start end state days total n mean n mean range range fig date date n distance distance 50% kernel 95% kernel (km) (km) (km²) (km²) 30 7/2/98 7/3/98 initial 28 24 4 3.0 16 5.4 1a 30 8/3/98 24/3/98 settled 16 17 4 0.7 10 2.4 1a 30 26/3/98 29/4/98 move 34 48 16 2.1 28 2.6 1b 30 1/5/98 31/5/98 settled 30 31 9 1.0 17 2.0 1b 30 1/6/98 30/6/98 settled 29 23 3 1.9 15 2.5 1c 30 1/7/98 13/7/98 move 12 19 6 1.8 13 2.5 1c 30 14/7/98 24/7/98 settled 10 20 9 0.7 11 1.8 30 25/7/98 10/8/98 move 16 44 3 0.8 22 3.0 1c 30 12/10/98 29/12/98 range 78 47 9 1.5 23 2.4 30 4/1/99 17/12/99 range 347 71 4 3.3 21 2.6 8.9 139.4 2a 30 24/12/99 28/2/00 musth 66 17 1 4.5 8 4.1 16.3 114.8 2b 31 7/2/98 17/2/98 initial 10 8 0 0.0 5 3.8 1d 31 18/2/98 3/3/98 settled 13 8 1 1.4 3 4.1 1d 31 4/3/98 13/3/98 move 9 7 1 0.5 5 1.7 1d 31 14/3/98 1/4/98 settled 18 7 0 0.0 3 0.6 1e 31 2/4/98 13/4/98 move 11 9 0 0.0 6 2.6 1e 31 14/5/98 25/7/98 settled 72 42 7 1.8 20 2.7 31 27/7/98 7/8/98 move 11 8 2 0.9 3 4.3 31 9/8/98 2/9/98 settled 24 12 2 2.3 4 2.1 4.6 24 2c 31 11/9/98 2/11/98 musth 52 47 13 2.0 23 4.2 6.8 89.2 2d 31 3/11/98 24/11/98 range 21 18 3 1.5 10 2.1 5 49.4 2e 31 25/11/98 4/1/99 musth 40 29 9 2.8 12 2.9 12.7 102.9 2f 31 6/1/99 23/2/99 range 48 16 2 0.7 4 2.1 31 2/3/99 6/10/99 range 218 44 4 3.7 13 4.0 2.8 118.7 2g 31 18/10/99 23/11/99 musth 36 2 0 0.0 0 3.8 38.9 2h 31 11/11/99 27/12/99 range 46 4 1 0.4 0 31 31/12/99 5/1/00 musth 5 5 2 2.8 2 6.7 31 6/1/00 21/2/00 range 46 7 0 0.0 3 6.8 32 22/3/98 15/4/98 initial 24 30 8 2.1 19 3.4 1f 32 16/4/98 25/11/98 range 223 188 46 1.8 101 2.4 19.1 133.1 2i 32 28/11/98 10/1/99 musth 43 29 6 1.9 15 4.3 23.3 132.1 2j 32 11/1/99 13/12/99 range 336 70 7 1.7 15 3.0 7.1 132.5 2k 32 16/12/99 16/2/00 musth 62 16 2 2.3 5 4.6 14.6 101.1 2l 33 23/3/98 2/4/98 initial 10 7 1 0.7 3 5.5 1g 33 3/4/98 25/6/98 with 83 61 9 2.9 31 3.8 33 27/6/98 18/9/98 away 83 48 8 1.3 20 3.4 33 19/9/98 19/1/99 with 122 93 28 2.0 43 3.6 33 23/1/99 19/3/99 away 55 21 2 1.8 7 5.0 33 21/3/99 22/5/99 with 62 24 4 2.2 8 3.4 33 3/6/99 21/11/99 away 171 15 0 2 5.3 33 7/1/00 28/2/00 with 52 13 3 1.1 2 6.2 slotow & van dyk.qxd 2004/10/05 10:42 page 103 koedoe 47/2 (2004) 104 issn 0075-6458 34 13/3/98 24/4/98 initial 42 62 22 2.1 37 3.2 1h 34 25/4/98 5/12/98 range 224 188 38 1.7 110 2.4 11.8 105.2 2m 34 7/12/98 28/1/99 musth 52 20 5 3.2 4 7.1 6.4 103.3 2n 34 10/2/99 11/6/99 range 121 68 11 1.7 32 4.0 34 23/6/99 13/12/99 range 173 28 1 4.5 6 4.8 3.8 92.4 2o 34 16/12/99 7/2/00 musth 53 22 4 2.0 11 3.3 19.2 106.6 2p 35 13/3/98 24/4/98 initial 42 58 21 2.0 33 3.7 35 26/4/98 30/5/98 range 34 39 14 1.7 19 2.3 35 31/5/98 15/7/98 range 45 25 1 0.8 15 2.0 35 16/7/98 13/11/98 range 120 85 12 1.0 50 2.6 35 17/11/98 1/4/99 range 135 54 9 1.7 17 3.8 10.1 122.9 2q 35 10/4/99 20/8/99 musth 132 64 14 1.0 31 2.2 4.6 114.8 2r 35 21/8/99 19/12/99 range 120 19 1 10.4 4 5.7 11 79.5 2s 35 10/1/00 7/2/00 musth 28 7 0 1 12.0 13.3 65.7 2t status: initial = initial period after introduction, and before the individual settled in an area. settled and move are used for individuals 30 and 31, and indicate periods of settlement in an area followed by periods of exploration (see text). range refers to the period when not in musth and after the initial settleing period. musth refers to the period in musth. appendix a (continued) same day next day core total id start end state days total n mean n mean range range fig date date n distance distance 50% kernel 95% kernel (km) (km) (km²) (km²) slotow & van dyk.qxd 2004/10/05 10:42 page 104 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 botha.qxd harvesting impacts on commonly used medicinal tree species (catha edulis and rapanea melanophloeos) under different land management regimes in the mpumalanga lowveld, south africa j. botha, e.t.f. witkowski and c.m. shackleton botha, j., e.t.f. witkowski and c.m. shackleton. 2004. harvesting impacts on commonly used medicinal tree species (catha edulis and rapanea melanophloeos) under different land management regimes in the mpumalanga lowveld, south africa. koedoe 47(2): 1–18. pretoria. issn 0075-6458. harvesting of products from plants in the wild is widespread throughout southern africa. particularly important products are plant parts used in traditional medicine. however, the impacts of harvesting practices are rarely quantified, with as yet insufficient generic rules across species and life forms. this limits the predictive ability to monitor and manage the affected populations. this paper examines the harvesting impact on two popular woody medicinal species used throughout sub-saharan africa, namely catha edulis (vahl) forssk. ex endl. (bushman’s tea) and rapanea melanophloeos (l.) mez. (cape beech). in both species, basal diameters, heights, and the number of size classes in the harvested populations were lower than in unharvested. densities of harvested populations were higher in both species, including densities of young plants, but the frequency of individuals in larger size classes was lower. the populations of both species being harvested for medicinal products appeared to be withstanding the current levels of harvesting, but the population structure of c. edulis populations being harvested for poles on the roadside and near human settlements had been substantially altered. despite the resilience of the two species to harvesting for medicinal products, loss of habitat or changes in land use pose a threat to a number of the assessed populations. key words: density, disturbance, dominance, harvesting, impact, medicinal plants, size class. j. botha and e.t.f. witkowski, restoration and conservation biology research group, school of animal, plant and environmental sciences, university of the witwatersrand, p.o. wits, 2050 republic of south africa; c.m. shackleton, department of environmental science, rhodes university, grahamstown, 6140 republic of south africa (c.shackleton@ru.ac.za). issn 0075-6458 1 koedoe 47/2 (2004) introduction disturbance factors frequently have greater effects in determining plant community structure, composition and functioning than do the primary abiotic variables (grime 1979). the effects of any single disturbance or stress are multidimensional, being influenced by its nature, intensity, timing, duration, frequency and interaction with other disturbance factors, as well as the characteristics of the community or species being disturbed. consequently, they are frequently difficult to predict. yet, in the face of accelerating land transformation (lambin et al. 2001) there is an increasing need to be able to predict the effects and outcomes of particular disturbance regimens on species, populations and communities, at a range of spatial scales, before potentially irreversible losses occur. in southern africa, past and present research efforts into the effects of disturbance have focussed on changes in species composition at a community level, essentially within a preservationist paradigm. yet changes in community and population structure often occur before major shifts in species composibotha.qxd 2004/10/05 09:53 page 1 tion, or species loss (harper 1977; shackleton 1993), and are a useful indicator of management and disturbance impact. changes in size class profile of key species, or communities, may alert managers to situations of declining recruitment (walker et al. 1986; shackleton et al. 1994; wiegand et al. 2000; wilson & witkowski 2003) and hence can be regarded as a useful threshold of concern. a widespread disturbance to plant populations in southern africa, and elsewhere, is the harvesting of plants from the wild (hoffman 1997; dzerefos & witkowski 2001). this can be for daily subsistence needs of rural communities, such as fuelwood or wild fruits, but increasingly it is being driven by the need of poor rural households to generate cash incomes (mander 1998; braedt & standa-gunda 2000; shackleton & shackleton 2000). much commercialisation happens at a local level, but equally significant is the supply of plant parts and products to external markets, such as to tourists, regional urban centres, and even export markets. the intensity of harvesting of specific plant species is influenced by resource abundance, accessibility and market demand. it often decreases with increasing distance from human settlements, transport routes and decreased accessibility of harvesting sites (konstant et al. 1995; cawe & ntloko 1997; luoga et al. 2002). the techniques used by commercial harvesters, or those not resident in the area, often tend to cause higher damage to target species than local resource users (cunningham 1992; botha 1998; luoga et al. 2000), although impact tends to be lower where land and/or resource tenure is strong (crook & clapp 1998). household and commercial demand for traditional medicines based on wild plants is a particularly large harvesting sector in south africa (mander 1998; williams et al. 2000; botha et al. 2004a, 2004b). this parallels the growth of the trade throughout the rest of the continent with the expansion of urban populations (marshall 1998). it is estimated that almost 70 % of south africa’s 40 million consumers make use of traditional medicine to some degree and approximately 30 000 households derive income from trading in such plants within kwazulu-natal province alone (mander 1998). catha edulis (vahl) forssk. ex endl. (bushman’s tea) (celastraceae) and rapanea melanophloeos (l.) mez. (cape beech) (myrsinaceae) are two popular woody medicinal species used throughout sub-saharan africa. in afro-arabian cultures, the leaves of the former are traditionally chewed to relieve fatigue and induce euphoria, but production and marketing are now generally prohibited due to side effects (van wyk et al. 1997). roots and bark are traded in medicinal plant markets in mpumalanga province of south africa (botha et al. 2001), while bark and leaves have been recorded in the urban markets of gauteng province (williams et al. 2001). bark and roots of r. melanophloeos are utilised medicinally, but only the bark has been recorded in markets in mpumalanga (botha et al. 2001), limpopo province (mander 1997), kwazulunatal (cunningham 1992) and gauteng (williams et al. 2000). although ranked only 65th out of 120 species in a trade study in limpopo (mander 1997), r. melanophloeos was one of ten species most commonly nominated as ‘popular’ in gauteng markets (williams et al. 2000). perceptions of demand in markets in the lowveld of mpumalanga (the low lying plains to the east of the drakensberg escarpment) were divided; 50 % of the respondents believed that this species was in high demand, while 50 % said that there was little demand for it (botha 2001). however, traditional healers listed r. melanophloeos as one of their most frequently used plants. in addition to their medicinal value, the wood of c. edulis is used for building, as wood pulp and to make furniture, and carvings, while that of r. melanophloeos is utilised for building materials and woodwork. although both species occur in many regions of southern africa, both have restricted distributions in mpumalanga, with habitats often under pressure through changing land use. nevertheless, vendors in the lowveld koedoe 47/2 (2004) 2 issn 0075-6458 botha.qxd 2004/10/05 09:53 page 2 reported that both species are readily available (botha et al. 2001). catha edulis is classed on the red data list as vulnerable in the neighbouring province of kwazulunatal (hilton-taylor 1996). thus, because of the perceived continuing high demand, but decline in a neighbouring province, local conservationists deemed it important to obtain greater knowledge of the population status of the species in the mpumalanga lowveld as part of a broader regional study on the medicinal plant trade (botha 2001), and to set the basis for prediction of harvesting impacts. plant demographic surveys are a cost effective means of assessing the impact of harvesting on plant populations, while simultaneously providing baseline data to enable the development of management strategies and for future monitoring (witkowski et al. 1994). within this context, this paper reports on patterns of harvesting of c. edulis and r. melanophloeos populations under different land management regimes in mpumalanga, south africa, as a case example facing conservation and management of popular mediissn 0075-6458 3 koedoe 47/2 (2004) fig. 1. the localities of the assessed catha edulis and rapanea melanophloeos populations. cinal species elsewhere in africa and the developing world. study area populations of both species were identified on the basis of local knowledge and accessibility in the lowveld region of mpumalanga, centred near the towns of hazyview and barberton, approximately 100 km apart (fig. 1). rainfall in this region occurs mainly in the hot summer months (october to may). mean annual rainfall in the two areas is between 900 and 1 200 mm (institute for soil, climate and water 2000, pers. comm.). the mean maximum temperature in the warmest month (january) is over 25 ºc and the minimum is 18.5 ºc at hazyview and 15.9 ºc at barberton. in winter, mean maximum and minimum temperatures are approximately 3–4 degrees lower. frost is rare. the natural vegetation of the lowveld is savanna, being dense under high rainfall sites and less so in more arid sites. being high rainfall sites, both barberton and hazyview fall in the sour lowveld bushveld vegetation type (low & rebelo 1986) dominated by, amongst others, combretum species, terminalia sericea, acacia species, pterocarpus angolensis and parinari curatellifolia. the shrub layer is characterised by taxa such as dichrostachys cinerea, antidesma venosum, pavetta species and maytenus species. methods field sampling for c. edulis, eight populations were sampled; five that were subject to different intensities of harvesting and three unharvested populations. one harvested population occurred on communal land, one at the side of a public road and three on land under commercial forest plantations, of increasing distance away from human settlements. of the three unharvested populations, one was on a private farm in a fragment of natural vegetation surrounded by farmbotha.qxd 2004/10/05 09:54 page 3 land (banana and mango plantations), and two were in a provincial protected area (songimvelo game reserve). one of these was growing within and alongside a perennial streambed subject to occasional flooding. the other occurred on a drainage line, and experienced infrequent fires. five populations of r. melanophloeos were sampled; three harvested and two unharvested. of the three harvested sites, two were on commercial forestry plantation land, and one on communal land. the forestry sites were burnt biennially. both the unharvested populations were on forestry land, one near hazyview and the other near barberton. total counts of a population were carried out in a particular stand or plot, depending on the size of the population; nine plots or populations for c. edulis and seven for r. melanophloeos. plot sizes varied according to the distribution of the target species, but were usually 20 x 40 m, or 20 x 30 m. as heights may be modified through natural or anthropogenic disturbance, both heights and basal diameters were measured, the latter at 30 cm above the ground as some individuals branched below breast height. each stem of multi-stemmed trees was measured and its vigour assessed. desiccated or rotting stems, with no shoots or other signs of life, were classified as dead. stressed stems were those showing signs of disease, for example, rotting or desiccated branches, but part of the plant was still living or it was coppicing. external damage to the stem was assessed on a scale adapted from cunningham (1992), where 0 = no damage; 1 = ≤10 % of the bark below 2 m was damaged; 2 = 11–25 % of the bark below 2 m was damaged; 3 = 26–50 % of the bark below 2 m was damaged; 4 = 51–75 % of the bark below 2 m was damaged; 5 = the stem was ringbarked; 6 = 76–100 % of the bark below 2 m was damaged and 7 = the stem was chopped. the cause of damage was also noted. the number of coppice shoots was recorded as a measure of coppicing vigour. at some sites, the roots of c. edulis had also been harvested. to assess the percentage survival of plants that had been harvested for root material, the extent of root damage was assessed according to the proportion of disturbed surface area radiating out from the base of the stem (botha 2001): 0 = no harvesting; 1 = <10 %; 2 = 11–25 %; 3 = 26–50 %; 4 = 51– 75 %; 5 = >76 %. data and statistical analysis the equivalent basal diameter was calculated for multi-stemmed individuals. size classes were grouped into basal diameters of 5 cm increments, apart from size classes 1 (0–1 cm) and 2 (2–5 cm). size class 1 included seedlings, coppices and stems from the previous season that had died back during the non-growing season (shackleton et al. 1994) and were termed ‘young plants’. size class distributions of harvested and unharvested populations were compared by means of kolmogorov-smirnov goodness of fit tests. the percentages of trees of harvestable size were calculated according to the size classes of populations that were currently being harvested. chi squared (χ²) analyses were performed to compare the proportions of trees of harvestable sizes in harvested and unharvested populations, as well as the proportions of individuals that had survived after harvesting. smaller size classes were pooled to ensure a minimum frequency of five in each class. basal diameter and height variations between harvested and unharvested populations were compared using non-parametric statistics (kruskal wallis and mann whitney u tests). quotients between the numbers of trees in successive size classes were calculated to assess the stability of populations (meyer 1952). fluctuating quotients indicate an unstable population, while the quotient between the numbers of trees in successive diameter classes approaches a constant value in a stable population. stability may also be assessed through simpson’s index of dominance (c) and the permutation index (p). the former describes the probability that any two trees drawn at random from the same community are of the same species or, in this case, from the same size class (pielou 1977, wiegand et al. 2000) (equation 1). the evenness of occupation of size classes is measured, ignoring the order in which size classes are arranged (wiegand et al. 2000). values above 0.1 reveal that the size frequency is steeper than would be expected from an exponentially declining population (i.e. the size classes are arranged in descending order), while values below 0.1 show that the size classes are more evenly distributed. where n is the total number of trees and ni the number of trees in class i, and k is the number of size classes. the permutation index assesses the deviation of a population from a monotonic decline, which would be expected in an "ideal" population. it is the sum of the absolute distances between the expected and real location (rank) of all size classes, and ignores the relative frequency of different size classes (equation 2) (wiegand et al. 2000). the permutation index is koedoe 47/2 (2004) 4 issn 0075-6458 eq. 1 botha.qxd 2004/10/05 09:54 page 4 where ji is the rank of size class i (i = 1 for the smallest trees), with the highest rank (ji = 1) given to the most frequent size class. results population size and density the densities of harvested c. edulis populations were significantly higher than those of the unharvested sites. the former ranged from 767 to 9800 trees/ha (3047 ± 1706 trees/ha), while the unharvested varied from 20 to 85 trees/ha (52 ± 19 trees/ha) (table 1). there was no significant difference between the population sizes of harvested and unharvested populations (393 ± 175 and 313 ± 131 trees, respectively). the density of harvested r. melanophloeos populations ranged from 789–8200 trees/ha while the unharvested populations ranged from 750–1013 trees/ha, but they were not significantly different (3864 ± 1464 & 882 ± 132 trees/ha, respectively). similarly, population sizes of harvested r. melanoissn 0075-6458 5 koedoe 47/2 (2004) eq. 2 table 1 population parameters (mean ± se) of harvested and unharvested catha edulis and rapanea melanophloeos populations (range is provided in brackets below the mean) species site estimated population young plant basal height simpson's permutation treatment population density density diameter (m) dominance index (p) size (no.) (trees/ha) (no./ha) (cm) index (c) catha edulis harvested 393 ± 175 3 047 ±1 706 2 192 ± 1 516 2.6 ± 0.1 1.6 ± 0.1 0.71 4 (96 1 000) (767 9 800) (75 8 188) (0.1 31.8) (0.1 18.0) unharvested 313 ± 131 52 ± 19 149 ± 126 8.3 ± 0.7 2.0 ± 0.1 0.32 2 (60 500) (20 85) (20 402) (0.1 29.9) (0.2 9.0) p values p > 0.05 p < 0.05 p < 0.05 p < 0.01 p < 0.01 rapanea harvested 1 056 ± 973 3 864 ± 1 464 2 783 ± 1 249 3.1 ± 0.2 1.1 ± 0.0 0.67 2 melanophloeos (789 8 200) (367 6 700) (0.1 27.1) (0.5 7.0) unharvested 136 ± 16 882 ± 132 485 ± 425 9.4 ± 0.7 2.2 ± 0.2 0.28 6 (750 -1 013) (60 -910) (0.1 34.1) (0.1 12) p values p > 0.05 p > 0.05 p > 0.05 p < 0.01 p < 0.01 phloeos were higher than unharvested (1056 ± 973 & 136 ± 16, respectively), but this was not significant (table 1). the densities of young plants in harvested c. edulis populations were significantly higher than in unharvested populations (2192 ± 1516 & 149 ± 126 trees/ha, respectively). there was a highly significant difference in the relative proportion of plants occurring in size class 0–1 cm in the harvested populations compared with the unharvested populations (71 % and 7 %, respectively) (fig. 2a). the frequencies of c. edulis in the larger size classes was low in the harvested populations. size class 6–10 cm comprised only 8 % of the population, and size class 11–15 cm 4 %, compared with 21 % and 12 %, respectively, for the unharvested populations. the densities of r. melanophloeos individuals in size class 1 on harvested sites (range 367 to 6700 trees/ha) tended to be higher than those on unharvested (range 60 to 910 trees/ha) but this was not significant (2 783 ± 1249 and 485 ± 425 trees/ha, respectively). size class distribution the size class distribution (basal diameters) of harvested c. edulis populations was significantly different from unharvested ones (d1507,127 = 0.64; p <0.005). the basal diameters of harvested populations were signifihigher in a discontinuous size class distribution than in a continuous, monotonically declining population. botha.qxd 2004/10/05 09:54 page 5 fig. 2. population structure of harvested and unharvested catha edulis populations (a) relative frequencies and (b) quotients between numbers of trees in successive size classes. fig. 3. population structure of harvested and unharvested rapanea melanophloeos populations (a) relative frequencies and (b) quotients between numbers of trees in successive size classes. koedoe 47/2 (2004) 6 issn 0075-6458 cantly lower than those of unharvested populations (2.6 ± 0.1 & 8.3 ± 0.7 cm, respectively) (table 1; fig. 2a). although fig. 2a shows higher basal diameters for the harvested populations, this is due to larger individuals growing on one site, namely, the forestry population near barberton. higher means are due to relatively higher frequencies of larger individuals in a population. similarly, the heights of harvested populations were significantly lower than those of the unharvested (1.6 ± 0.1 m & 2.0 ± 0.1 m, respectively), although the maximum height for harvested populations was higher than that in the unharvested (table 1). individuals were absent from size classes on the roadside c. edulis population near hazyview, and only the first four size classes (0–15 cm) on the communal site had individuals (figs. 4d & 4e). there were no size classes with frequencies of zero in the harvested forestry populations near barberton (figs. 4a-c). there was a significant difference between the size class distributions (basal diameters) of harvested and unharvested r. melanophloeos populations (d704,227 = 0.35; p < 0.005). mean basal diameters of harvested r. melanophloeos were significantly lower than plants in the unharvested populations (3.1 ± 0.2 & 9.4 ± 0.7 cm, respectively) (table 1; fig. 3). similarly, plant height in the harvested populations was significantly lower than in the unharvested (1.1 ± 0.0 m and 2.2 ± 0.2 m, respectively). botha.qxd 2004/10/05 09:54 page 6 issn 0075-6458 7 koedoe 47/2 (2004) fig. 4(a-c). degree of stem damage to harvested catha edulis through harvesting of bark below 2 m on the stem, and chopping of stems in populations situated (a) adjacent to a village near barberton; (b) approximately 3 km from the village; (c) on a forestry plantation near barberton. botha.qxd 2004/10/05 09:54 page 7 there was a significantly higher proportion of individuals in size class 0–1 cm in the harvested compared with the unharvested ones (48 % and 18 %, respectively). in the harvested populations, 13 % and 3 % of the population had survived to size classes 6–10 cm and 11–15 cm, respectively. in the unharvested r. melanophloeos, 19 % of the population had survived to size class 6–10 cm and 11 % to 11–15 cm. quotients were relatively stable in both the harvested and unharvested c. edulis populations (fig. 2b), although they were more variable in the latter. both populations, but particularly the harvested, showed steeper distributions than would be expected from an exponentially declining distribution (table 1). neither the harvested nor the unharvested populations deviated substantially from a monotonic decline. quotients between numbers of trees in successive size classes showed more variation in the unharvested r. melanophloeos populations than in the harvested ones (fig. 3b). both harvested and unharvested populations had steeper size frequency distributions than would be expected from an exponentially declining distribution (simpson’s index of dominance = 0.67 and 0.28, respectively), particularly the former (table 1). neither population deviated substantially from a monotonic decline. harvesting impact there was a highly significant difference in the proportion of stems of harvestable size between the harvested and unharvested c. edulis populations (38 ± 12 % and 54 ± 20 %, respectively) (χ² = 175.9; df = 1; p < 0.0001) (table 2). no c. edulis had been harvested on the private farm near hazyview, and less than 0.1 % of the protected area populations had been utilised. the extent of utilisation of harvested c. edulis populations ranged from 7–76 % of the harvestable stems (36 ± 14 %), 0–42 % of the roots (14 ± 8 %) and 2–24 % (8 ± 4 %) of roots and stems harvested on the same tree. the degree of harvesting was high in the c. edulis populations adjacent to settlements and roadsides, although the harvesting patterns varied. the majority of larger stems had been intensively harvested (>26 % of the koedoe 47/2 (2004) 8 issn 0075-6458 table 2 extent of harvesting of catha edulis and rapanea melanophloeos populations (mean ± se) and condition of the harvested stems. (range in brackets under the mean) (# statistical analysis not performed as n = 0 in one or more cells) c. edulis r. melanophloeos harvested unharvested p harvested unharvested p proportion of stems of 38 ± 12 54 ± 20 p < 0.01 19.2 ± 3.8 59.0 ± 21.0 p< 0.01 harvestable size (%) (9 71) (34 -73) (7 31) (28 80) proportion of stems -trunk 35.6 ± 13.9 0.03 ± 0.02 # 26.8 ± 8.5 1.0 ± 1.0 # harvested (7 76) (0 0.05) (9 57) (0 2.0) -roots 14.0 ± 7.8 0 # 0 0 # (0 -42) -trunk & 7.6 ± 4.4 0 # 0 0 # roots (2 -24) condition of -dead 4.8 ± 3.0 0 # 8.6 ± 5.4 0 harvested trees (2 17) (4 29) -stressed 1.2 ± 1.1 0 # 0.6 ± 0.6 0 (0 9) (0 3) -alive 96.5 ± 3.2 100 # 90.4 ± 55.1 100 (74 100) (71 100) botha.qxd 2004/10/05 09:54 page 8 issn 0075-6458 9 koedoe 47/2 (2004) fig. 4(d-f). degree of stem damage to harvested catha edulis through harvesting of bark below 2 m on the stem, and chopping of stems in populations situated (d) on communal land in nsikazi; (e) on the roadside near hazyview and (f) pooled populations. botha.qxd 2004/10/05 09:54 page 9 stem below 2 m) in the c. edulis populations growing adjacent to the village near barberton, with extensive ringbarking taking place (fig. 4a). of the ringbarked individuals, 80 % had died, and 10 % showed signs of stress. chopping had also occurred across all size classes. both the degree and extent of harvesting was lower in the population situated approximately 3 km from the village, although the level of exploitation was high, with the majority of stems in the three largest size classes (basal diameters 15–25 cm) having been utilised. limited ringbarking had occurred. limited harvesting of bark had occurred in the population on communal lands and the roadside population near hazyview. here, the bulk of harvesting was for poles for building materials (figs. 4d & 4e). in the most remote forestry population, c. edulis had only been harvested for medicinal materials (fig. 4c). only 15 % of the harvestable trees had been utilised, the majority of which had occurred in the highest size classes (basal diameters 36–50 cm). although the degree of harvesting in these size classes was high (26–75 %), no stems had been ringbarked. roots had been utilised at all the harvested sites, with the exception of the one situated approximately 3 km from a village in the barberton area. the majority of individuals in the larger size classes had been harvested at intensities >26 % of their root circumference (fig. 5). c. edulis roots had been harvested most extensively at the roadside site, with 42 % of the population having been affected at degrees of 26–50 % of their circumference (fig. 5d). on the forestry site, roots of all the individuals in the largest size classes (36–50 cm) had been utilised (fig. 5b). stems and roots had also been harvested from the same trees, with the highest percentage occurring on the roadside site (24 %). a significantly higher proportion of r. melanophloeos were of harvestable size in the unharvested populations, compared with those that had been harvested (χ² =190.0; df =1; p < 0.0001) (59 ± 21 % & 19 ± 4 %, respectively; table 2). in the harvested populations, 27 ± 9 % of trees had been utilised. the degree and extent of harvesting was high in both the forestry populations near barberton and the population on communal land near hazyview. the majority of stems in the largest size classes (basal diameters 21– 45 cm) of the forestry populations had been utilised at levels >10 %, with a considerable proportion having been harvested at koedoe 47/2 (2004) 10 issn 0075-6458 table 3 proportion of harvested and unharvested catha edulis and rapanea melanophloeos populations that were coppicing or single stemmed (# statistical analysis not performed as n=0 in one or more cells) species populations individuals % % single p coppicing stemmed c. edulis total population harvested 45 55 p < 0.01 unharvested 86 14 harvested harvested individuals 56 44 p < 0.05 unharvested individuals 40 60 unharvested harvested individuals 100 0 # unharvested individuals 85 15 r. melanophloeos total population harvested 45 55 p < 0.01 unharvested 30 70 harvested harvested individuals 23 77 p < 0.05 unharvested individuals 48 52 unharvested harvested individuals 100 0 # unharvested individuals 31 69 botha.qxd 2004/10/05 09:54 page 10 issn 0075-6458 11 koedoe 47/2 (2004) fig. 5(a-c). the degree of root disturbance through harvesting of catha edulis at sites (a) adjacent to a village and (b) in a forestry plantation near barberton; (c) on communal land populations. botha.qxd 2004/10/05 09:54 page 11 koedoe 47/2 (2004) 12 issn 0075-6458 fig. 5(d-e). the degree of root disturbance through harvesting of catha edulis at sites (d) a roadside site near hazyview, and (e) pooled populations. between 51 and 75 % (figs. 6a & 6b). only one r. melanophloeos stem had been chopped in the forestry populations. harvesting pressure in the population located close to communal lands was high from size class 11–15 cm and larger, with either chopping or ringbarking occurring in all these size classes. no harvesting had occurred in the population located close to hazyview. there was no significant difference between the proportions of dead, alive or stressed c. edulis in the harvested and unharvested populations. c. edulis appears resilient to harvesting, with the percentage survival of harvested trees ranging from 74–100 % (97 ± 3 %; table 2). only a few trees appeared stressed after harvesting (1 ± 1 %) or had died (5 ± 3 %). the majority of the stressed or dead individuals had been ringbarked. the percentage survival of harvested individuals ranged from 71–100 % on the harvested sites (90 ± 55 %; table 2). between 4 % and 29 % of the harvested stems had died (9 ± 5), and 0.6 ± 0.6 % appeared stressed. all the r. melanophloeos on the unharvested sites appeared healthy. coppicing the number of stems per tree in harvested c. edulis was significantly lower than in the unharvested ones (1.4 ± 0.03 & 7.0 ± 0.9, respectively). there was also a highly significant difference in the proportions of multi-stemmed trees growing in harvested and unharvested populations. in the former populations, 45 % of the c. edulis were coppicing, compared with 86 % coppicing in the unharvested populations (table 3). within the harvested populations, a significantly higher proportion of harvested individuals were coppicing (56 %), compared with the proportion of unharvested trees that were multi-stemmed (40 %). in the unharvested populations, all harvested individuals were coppicing (n = 4), as were 85 % of the unharvested . there was no significant difference between the numbers of stems per tree in harvested r. melanophloeos (1.5 ± 0.04 stems) vs unharvested populations (1.6 ± 0.1 stems) (u862,227 = 99 856; p > 0.05). there was, however, a highly significant difference between the proportions of trees coppicing in the harvested and unharvested populations (χ² = 10.9; df =1; p <0.0001). in the latter, 30 % of the trees were coppicing, while 45 % of the harvested population were multi-stemmed (table 3). within the harvested populations, the proportion of utilised individuals that were coppicing was significantly lower than the proportion of botha.qxd 2004/10/05 09:54 page 12 issn 0075-6458 13 koedoe 47/2 (2004) melanophloeos stems, as only five stems had been chopped. discussion species respond to disturbance differently, depending on where they fit along the continuum between k and r strategists (primack 1993; shackleton et al. 1994). significant decreases have been found in stem density, basal area, biomass, height, young plant density and species richness when certain woody species are subject to human disturbance (rao et al. 1990; shackleton et al. 1994; clarke 1997). in others, pioneers for example, the opposite has been found (rao et al. 1990; shackleton et al. 1994). in this study, the densities of harvested populations were higher than for the unharvested, although not statistically significant for r. melanophloeos. fig. 6. degree of stem damage to harvested rapanea melanophloeos through harvesting of bark below 2 m on the stem, and chopping of stems in populations located on (a) and (b) a forestry plantation near barberton and (c) near nsikazi. unharvested individuals that were multistemmed (23 % and 48 %, respectively) (χ² = 4.4; df =1; p <0.05). there was a relationship between the height of chopping of catha edulis stems, and the number of stems per individual (r² = 0.19; p <0.05) (fig. 7), but no relationship between the basal diameter of a chopped stem and the number of coppices (r² = 0.003; p > 0.05). there were insufficient data to test this relationship with rapanea in both species, the densities of young plants were higher in the harvested populations compared with the unharvested. while the frequency of plants in size class 1 was high, numbers in subsequent size classes was limited in populations that had been extensively harvested. c. edulis has been observed in fruit at heights of 2–3m (pers. comm. g. strydom, mpumalanga parks board 2000). in the harvested populations 22 % of the c. edulis were >2 m, compared with 36 % in the unharvested populations. within the heavily harvested population on the commubotha.qxd 2004/10/05 09:54 page 13 koedoe 47/2 (2004) 14 issn 0075-6458 nal site near hazyview, only 4 % of individuals were taller than 2 m. the approximate size at which r. melanophloeos produces seed is unknown, but was estimated to be between 4 and 6 m, depending on growth form (pers. comm. g. strydom, mpumalanga parks board 2000; pers. comm. w. froneman, nelspruit botanic gardens 2000). in the harvested population, 12 % were >2 m, while 24 % of the unharvested population were >2 m. only 3 % of the harvested and 13 % of the unharvested r. melanophloeos were >4 metres. basal diameters and heights were significantly lower in the harvested populations than in the unharvested, with high levels of chopping in certain c. edulis populations. growth may also have been inhibited through physiological stress resulting from harvesting (hall & bawa 1993). a decrease in mature trees diminishes the sexual reproductive potential of a population through a lowering of seed production which, if unchecked, could lead to a total loss of sexual reproduction in a population, although c. edulis and r. melanophloeos are able to regenerate successfully vegetatively (van wyk 1988). the communal and roadside c. edulis populations also had a reduced number of size classes, which also may lower the probability of long-term persistence in a population if harvesting pressure is maintained (harper 1977). fig. 7. relationship between the height of chopping and the number of coppices for catha edulis. as often occurs in woody populations where wide climatic fluctuations, fire, herbivory and harvesting pressures are experienced (walker et al. 1986), both the harvested and unharvested r. melanophloeos populations appeared unstable. the c. edulis population were relatively more stable. the size frequency distributions of all the populations were steeper than would be expected from an exponentially declining population (c values >0.1). the c values of harvested populations were higher than those of the unharvested, showing steeper size frequency distributions in the former through selective harvesting of trees in higher size classes. the harvested r. melanophloeos and unharvested c. edulis populations showed the highest levels of monotonic decline (p = 2), while the size classes of the harvested c. edulis and unharvested r. melanophloeos populations were more discontinuous (p = 4 and 6, respectively), the former due in part to the extensive chopping of poles on the communal and roadside sites. many savanna species have adapted to herbivory and regular dry season fires by coppicing from surviving rootstocks, often resulting in multi-stemmed trees (bond & van wilgen 1996; luoga et al. 2004). a significantly higher proportion of harvested c. edulis individuals were coppicing within the harvested populations, compared with the unharvested trees in the same populations. the high level of coppicing in the unharvested c. edulis populations can be attributed to other forms of disturbance. the c. edulis population at one site in songimvelo game reserve was growing along a stream that was subject to periodic flooding. at the other site, 12 % of the population showed signs of having been burnt. none of the harvested c. edulis sites showed additional signs of disturbance, apart from the sites adjacent to the rural villages. although only 1 % of these stems had been burnt, this area is subject to regular burning by residents to improve the quality of grazing for livestock. similarly, the higher proportion of unharvested r. melanophloeos stems that were coppicing in the harvested populations is botha.qxd 2004/10/05 09:54 page 14 issn 0075-6458 15 koedoe 47/2 (2004) probably due to fire. the forestry plantation sites are currently burned every two years, but were burnt annually until recently. the commercially harvested population growing close to communal lands was relatively protected from fire, with only 0.6 % of the r. melanophloeos individuals showing signs of having been burnt. the unharvested population was also protected from fire, as it grows in a sheltered position beneath a granite outcrop; 80 % of the r. melanophloeos in this population were single stemmed. while all the populations assessed, apart from the c. edulis growing on the roadside and the communal area, appear to be withstanding current levels of harvesting, they may become vulnerable if utilisation increases, which is possible as harvesters from kwazulu-natal attempt to locate new harvesting areas to replace those where populations have become locally extinct (williams et al. 2000). higher levels of bark removal are also likely to decrease the plants’ ability to withstand regular fires. fire also suppresses the growth of young plants and saplings in particulatr. the combined effects of harvesting and fire diminishes the regeneration capacity of both species. in india and tanzania, for example, different harvesting patterns were discernible in plant populations subject to different land management regimes, with distinct gradients in harvesting intensity corresponding to the accessibility of populations to human settlements (chandrashekara & sankar 1998; luoga et al. 2000, 2004). the extent and degree of harvesting in c. edulis populations near settlements and along the roadside was high. the c. edulis population growing adjacent to the village near barberton had been more intensively utilised than the one located approximately 3 km from the village, with both roots and bark being harvested. adjacent to the village, stems had been chopped in 23 % of the population across all size classes, while in the population further from the village, only 3 % of the stems had been chopped. no harvesting for roots had occurred in the latter. the extent and degree of harvesting in the population located within the forestry plantation approximately 15 km from the nearest human settlement was substantially lower and no poles had been extracted. the largest c. edulis individuals assessed during this study were growing on the forestry site, and there were greater proportions of individuals in the larger size classes. limited harvesting of bark had occurred in the c. edulis populations growing in the communal areas and on the roadside near hazyview, although extensive harvesting for roots had occurred, particularly on the latter site. poles had been extensively chopped in both populations. it is unlikely that the r. melanophloeos populations growing on the forestry plantations were being harvested for commercial purposes at the time of the study. although 26–75 % of the bark had been removed below 2 m from the stems in the largest size classes, these comprised a relatively small proportion of the population and the overall extent of harvesting was relatively low. few stems had been chopped or ringbarked. the population near communal lands was being harvested by vendors for bark to trade on local markets. it is more likely that the chopped stems were being utilised for building materials than for medicinal purposes, though, as r. melanophloeos bark was still within easy reach of the harvesters. in kwazulu-natal, species such as ocotea bullata are chopped to enable harvesters to access bark in the higher reaches of the tree, once the lower bark has been harvested. in an eastern cape study, all the trees ringbarked some time prior to the assessment had died—including r. melanophloeos (la cock & briers 1992). the proportion of trees surviving ringbarking in this study was also low, with only 20 % of the ringbarked c. edulis surviving. in a concurrent study, 27 % of the assessed warburgia salutaris had survived ringbarking; the remainder had either died or showed signs of stress (botha et al. 2004b). apart from the ringbarked trees, the percentage survival of harvested c. edulis and r. melanophloeos plants was high, including botha.qxd 2004/10/05 09:54 page 15 koedoe 47/2 (2004) 16 issn 0075-6458 individuals of the former species that had been harvested for both roots and bark. species with good coppicing ability have been shown to be less vulnerable to anthropogenic disturbances than those with poor coppicing ability. both c. edulis and r. melanophloeos exhibited high levels of coppicing after both natural (fire) and anthropogenic disturbance. this corresponds to results obtained for c. edulis in a previous survey conducted at songimvelo (mbewe et al. 1997). in a study on the impact of harvesting on woody species (including dalbergia melanoxylon, spirostachys africana, acacia ataxacantha and combretum imberbe) in limpopo, all the recently harvested trees had survived and over 70 % of the harvested stems of each species were coppicing (clarke 1997). in another study investigating the impact of chopping on savanna trees in the same area, all the chopped stumps had coppiced (shackleton 2000). in this study, 86 % of the chopped c. edulis stems had coppiced. only five r. melanophloeos stems had been chopped, four of which were coppicing. the basal diameter of the stem and the height at which a stem was chopped may affect the coppicing success of certain species. with an increase in harvesting height, there was a decrease in the number of shoots produced by c. imberbe, a. ataxacantha and, to a lesser degree, d. melanoxylon in the central lowveld (clarke 1997). in addition, the higher the harvesting height the longer the shoot length, possibly due to livestock grazing the shoots of the shorter tree stumps (clarke 1997). the opposite occurred with c. edulis in this study; there was an increase in the number of stems with an increase in harvesting height (botha 2001). this corresponds to other observations that coppicing vigour decreases when trees are cut low down (wild & mutebi 1996; shackleton 2000; luoga et al. 2004). there was no relationship between harvesting height and the heights of coppices in either c. edulis or w. salutaris populations, or the basal diameter and number of coppices in either species (botha 2001). this study has highlighted the different harvesting patterns on c. edulis and r. melanophloeos under different management regimes. in both species, basal diameters, heights, and the number of size classes in the harvested populations were lower than in unharvested. densities of harvested populations were higher in both species, including densities of young plants, but the frequency of individuals in higher size classes was lower. the populations of both species being harvested for medicinal products appeared to be withstanding the current levels of harvesting, but the population structure of c. edulis populations being harvested for poles on the roadside and near human settlements had been substantially altered. the c. edulis and r. melanophloeos populations situated within forestry plantations appeared the most vigorous. despite the resilience of the two species to harvesting for medicinal products, loss of habitat or changes in land use pose a threat to a number of the assessed populations. for example, the r. melanophloeos population growing near the communal lands was located on a remnant patch of indigenous vegetation, surrounded by plantations. the c. edulis population growing on communal land in nsikazi was located between two homesteads; it is quite possible that this land could be allocated for settlement or agriculture in the future. on the banana farm near hazyview, a similar site to the remnant patch of open woodland on which the c. edulis was growing was cleared for cultivation two years prior to the survey. although the farmer was conservation conscious (he had granted traditional healers and other local user groups harvesting access approximately one year prior to the bush clearing), the land may be converted if economic conditions dictate. acknowledgements this study was funded by the green trust. the kruger national park and university of the witwatersrand are thanked for logistical support. the following people are thanked for their assistance with field surveys: mr dup du plessis, mr elijah lekhulene, ms josephina hloliwe silinda, mr phillip papbotha.qxd 2004/10/05 09:54 page 16 issn 0075-6458 17 koedoe 47/2 (2004) pin, mr ernst schmidt, mr king khoza, mr andries mathonsi and sipho. references bond, w.j. & b. van wilgen. 1996. fires and plants. cambridge: cambridge university press. botha, j. 1998. developing an understanding of problems being experienced by traditional healers living on the western border of the kruger national park: foundations for an integrated conservation and development programme. development southern africa 15: 621–634. botha, j. 2001. perceptions of species availability and values of medicinal plants traded in areas adjacent to the kruger national park. m.sc thesis, university of the witwatersrand, johannesburg. botha, j., e.t.f. witkowski & c.m. shackleton. 2001. an inventory of the medicinal plants traded on the western boundary of the kruger national park, south africa. koedoe 44(2): 7–46. botha, j., e.t.f. witkowski & c.m. shackleton. 2004a. market profiles and trade in medicinal plants in the lowveld, south africa. environmental conservation 31: 38–46. botha, j., e.t.f. witkowski & c.m. shackleton. 2004b. the impact of commercial harvesting on warburgia salutaris (‘pepper-bark tree’) in mpumalanga, south africa. biodiversity & conservation 13: 1675–1698. botha, j., e.t.f. witkowski & c.m. shackleton & d.h.k. fairbanks (in press). socio-economic differentiation in the trade of wildlife species for traditional medicines in the lowveld, south africa: implications for resource management. the international journal of sustainable development and world ecology. braedt, o. & w. standa-gunda. 2000. woodcraft markets in zimbabwe. international tree crops journal 10: 367–384. cawe, s.g. & m. ntloko. 1997. distribution, uses and exploitation patterns of flagellearia guinensis schumach. with particular reference to port st. johns, south africa. south african journal of botany 63: 233–238. chandrashekara, u.m. & sankar, s. 1998. ecology and management of sacred groves in kerala, india. foresty ecology and management 112: 165–177. clarke, a.b. 1997. sustainability of harvesting seven favoured plant species used in the indigenous wood carving industry in the bushbuckridge district of the northern province lowveld. honours dissertation, university of the witwatersrand, johannesburg. crook, c. & r.a. clapp. 1998. is market-orientated forest conservation a contradiction in terms? environmental conservation. 25: 131–145. cunningham, a.b. 1992. imithi isizulu: the traditional medicine trade in natal/kwazulu. m. soc sci. dissertation, university of natal, pietermaritzburg. godoy, r.a. & k.s. bawa. 1993. the economic value and sustainable harvest of plants and animals from the tropical forest: assumptions, hypotheses and methods. economic botany 47: 215–219. grime, j.p. 1979. plant strategies and vegetation processes. chichester: wiley. hall, p. & bawa, k. 1993. methods to assess the impact of extraction of non timber forest products on plant populations. economic botany 47: 234–247. harper, j.l. 1977. population biology of plants. san diego: academic press. hilton-taylor, c. 1996. red data list of southern african plants. strelitizia 4: 1–117. hoffman, m.t. 1997. human impacts on vegetation. pp 507–534. in: cowling, r.m., richardson, d.m. & pierce, s.m. (eds.). vegetation of southern africa. cambridge: cambridge university press. konstant, t.l., s. sullivan & a.b.cunningham. 1995. the effects of utilization by people and livestock on hyphaene petersiana (arecaceae) basketry resources in the palm savanna of northcentral namibia. economic botany 49: 345–356. la cock, g.d. & j.h. briers. 1992. bark collecting at tootabie nature reserve, eastern cape, south africa. south african journal of botany 58: 505–509. lambin, e.f., b.l. turner, h.j. geist, s.b. agbola, a. angelsen, j.w. bruce, o.t. coomes, r. dirzo, g. fischer, c. folke, p.s. george, k. homewood, j. imbernon, r. leemans, x. li, e.f. moran, m. mortimore, p.s. ramakrishnan, j.f. richards, h. skånes, w. steffen, g.d. stone, u. svedin, t.a. veldkamp, c. vogel & j. xi. 2001. the causes of land-use and land-cover change: moving beyond the myths. global environmental change 11: 261–269. letsela, t., e.t.f. witkowski & k. balkwill. 2003. plant resources used for subsistence in tsehlanyane and bokong in lesotho. economic botany 57: 619–639. low a.b. & a.g. rebelo. 1996. vegetation of south africa, lesotho and swaziland. pretoria: department of environment affairs and tourism. luoga e.j., e.t.f. witkowski & k. balkwill. 2000. economics of charcoal production in miombo woodlands of eastern tanzania: some hidden costs associated with commercialisation of the resources. ecological economics 35: 243–257. botha.qxd 2004/10/05 09:54 page 17 luoga e.j., e.t.f. witkowski & k. balkwill. 2002. harvested and standing wood stocks in protected and communal miombo woodlands of eastern tanzania. forest ecology and management 164: 15–30. luoga e.j., e.t.f. witkowski & k. balkwill. 2004. regeneration by coppicing (resprouting) of miombo (african savanna) trees in relation to land use. forest ecology and management 189: 23–35. mander, m. 1997. medicinal plant marketing in bushbuckridge and mpumalanga: a market survey and recommended strategies for sustaining the supply of plants in the region. nelspruit: darudec/department of water affairs and forestry. mander, m. 1998. marketing of indigenous medicinal plants in south africa. a case study in kwazulu-natal. rome: food and agricultural organisation of the united nations (fao). marshall, n.t. 1998. searching for a cure: conservation of medicinal wildlife resources in east and southern africa. cambridge: traffic international. mbewe, m.n., j.a. cooke & b.r. page. 1997. plant resource availability, utilisation and management in and around songimvelo game reserve, south africa. internal report. centre of indigenous plant use research, dept of biology, university of natal, durban. meyer, h.a. 1952. structure, growth and grain in balanced uneven-aged forests. south african journal of forestry 50: 85–92. pielou, e.c. 1977. mathematical ecology. new york, ny: wiley. primack, r.b. 1993. essentials of conservation biology. massachusetts: sinauer. rao, p., s.k. barik, h.n. pandey & r.s. tripathi. 1990. community composition and tree population structure in a sub-tropical broad-leaved forest along a disturbance gradient. vegetatio 88: 151–162. shackleton, c.m. 1993. demography and dynamics of the dominant woody species in a communal and protected area of the eastern transvaal lowveld. south african journal of botany 59: 569–574. shackleton, c.m. 2000. stump size and the number of coppice shoots for selected savanna tree species. south african journal of botany 66: 124–127. shackleton, c.m., n.j. griffin, d.i. banks, j.m. mavrandonis & s.e. shackleton. 1994. community structure and species composition along a disturbance gradient in a communally managed south african savanna. vegetatio 115: 157–167. shackleton, c.m. & s.e. shackleton. 2000. direct use values of savanna resource harvested from communal savannas in the bushbuckridge lowveld, south africa. journal of tropical forest products 6: 21–40. shackleton, s.e., c.m. shackleton, t.r. netshiluvhi, b.s. geach, a. ballance & d.h.k. fairbanks. 2002. use patterns and value of savanna resources from three rural villages in south africa. economic botany 56: 130–146. van wyk, b., b. van oudtshoorn & n. gericke. 1997. medicinal plants of south africa. pretoria: briza. van wyk, p. 1988. field guide to the trees of the kruger national park. cape town: struik. walker, b.h., l. stone, l. henderson & m. vernede. 1986. size structure analysis of the dominant trees in a south african savanna. south african journal of botany 52: 397–402. wiegand, k., d. ward, h.h. thulke & f. jeltsch. 2000. from snapshot information to long-term population dynamics of acacias by a simulation model. plant ecology 150: 97–114. wild, r.g. & j. mutebi, j. 1996. conservation through community use of plant resources. establishing collaborative management at bwindi impenetrable and mgahinga gorilla national parks, uganda. france: united nations educational, scientific and cultural organisation. williams, v.l., k. balkwill & e.t.f. witkowski. 2000. unravelling the commercial market for medicinal plants and plant parts on the witwatersrand. economic botany 54: 310–327. williams, v.l., k. balkwill & e.t.f. witkowski. 2001. checklist of plants used in the muthi trade on the witwatersrand, south africa. bothalia 31: 71–98. wilson, b.g. & e.t.f. witkowski. 2003. seed banks, bark thickness, and changes in age and size structure (1978–1999) of the african savanna tree, burkea africana. plant ecology 167: 151–162. witkowski e.t.f., b.b. lamont & f.j. obbens. 1994. commercial picking of banksia hookeriana in the wild reduces subsequent shoot, flower and seed production. journal of applied ecology 31: 508–520. koedoe 47/2 (2004) 18 issn 0075-6458 botha.qxd 2004/10/05 09:54 page 18 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 gandy.qxd altertative trophy measuring techniques for african buffalo s.e. gandy and b.k. reilly gandy, s.e. and b.k. reilly. 2004. altertative trophy measuring techniques for african buffalo. koedoe 47(1): 119–124. pretoria. issn 0075-6458. the african buffalo is considered the classic african trophy. it is the choice of many hunters who will never go on to hunt any of the other dangerous game animals on the continent. a good trophy is perceived as that of a mature bull with a hardened boss and horn tips that lengthen into sharply pointed hooks. however, indications are that these are the bulls in their breeding prime and there is concern that the continued targeting of these individuals will negatively impact on the population dynamics of the herds, ultimately affecting the sustainability of buffalo hunting. as they age and become postreproductive, the horns broom down, reducing the trophy score under the current measurement systems. a new measuring system is needed that encourages hunters to target the older post-reproductive bulls, instead of those that are still breeding. a random sample of trophies was divided into broomed and non-broomed sub-samples. all key parameters that can be measured in the trophy were measured with a view to identifying the parameters that would allow broomed-down individuals to compete favourably with the non-broomed “classic trophy” in the primary measurement systems, those of safari club international and rowland ward. an index, created through dividing tip space by the mean of the two individual horn lengths proved to serve the purpose. this factor was then applied to the mean of the sci and rowland ward measurements in the samples. these methods allowed broomed horns to score more points in the record books than non-broomed horns. boss width and boss space are other possible measurement inclusions that could be considered. s.e. gandy and b.k. reilly , department of nature conservation, tshwane university of technology, private bag x680, pretoria, 0001 republic of south africa. issn 0075-6458 119 koedoe 47/1 (2004) introduction the african buffalo has always attracted attention, not only because of its value as a member of the ‘big five’, but also because of its spectacular qualities from the sportsman’s point of view. most hunters regard them as one of the ultimate trophies. by nature, buffalo are unpredictable and therefore difficult and dangerous to get close to, so the true spirit of hunting—fear, fascination, and a test of the hunter’s skills—is the main reason why every dedicated hunter desires to devote a period of his or her hunting career in pursuit of the finest trophy (grobler 1996). trophy hunting is a specialised form of game utilisation, where adult males are selected for their trophy value. buffalo bulls reach sexual maturity at 4–5 years (estes 1997), but the trophy is still ‘green’ at this stage and not of much value as a trophy. the trophy reaches its full potential when the boss hardens to form the so-called armour plates, and the horns are fully grown out to form the characteristic hooks. at this stage the points are still sharp and well formed. authors estimate age at 7 to 12 years in contrast to sinclair's (1977) visual estimate of 5 to 10 years of age. as they become older, their boss closes up completely and the horn tips wear down more and more to form the typical ‘old bull’ shape (grobler 1996). technique gandy.qxd 2004/04/14 06:09 page 119 the dominant trophy measuring systems— safari club international (sci)—encourages hunters to target soft-bossed, “green” bulls and horn tips sharply pointed and well formed, as this is when a trophy scores the highest points in the record books. these bulls are in their prime, however, and at peak breeding status within the herds. it is believed by a number of researchers, veterinarians and the african safari industry (e.g., grellmann, anvo@mweb.com.na pers. comm.; heath, fishunt@samara.co.zw pers. comm.; robertson 1996) that to hunt them at this age cuts short their genetic contribution, whereas the older bulls have already passed on their genes and left the herds to live solitary existences. these individuals and organisations all suggest that by giving a bias to the boss in sci’s measuring system will solve the problem, encouraging trophy hunters to harvest older males. grellmann (pers. comm.) also encourages modifying sci trophy scores for antelope so that those with larger base circumferences ranked the highest, believing this would help select older post-reproductive males with broomed-down horns. sci requires 100 points (fig. 2), reached by adding the two boss width measurements to the total length of the horns from one tip to the other over the forehead (quimby 1993). under the sci measuring system, once the horns broom down to ‘old bull’ status, the trophy scores less points in the record books because of the decline of total width and total length. a new measuring system is needed that allows older, worn-down trophies to compete with bulls in their prime. this will allow enough time for bulls to complete their breeding while simultaneously allowing for a better reflection of the trophy as a function of the age of the animal, which will ultimately contribute to the economic and possibly ecological sustainability of buffalo hunting. rowland ward’s scoring, using the outside spread, is also believed inadequate since different populations of buffalo may have genetically and/or environmentally (e.g., food availability, nutritional constituency of food supply or physical environment) prekoedoe 47/1 (2004) 120 issn 0075-6458 fig. 1. diagrammatical representation of the current measuring methods as applied by safari club international and rowland ward. gandy.qxd 2004/04/14 06:09 page 120 determined tendencies for larger or smaller outside spreads. for instance tarangire and moyowosi buffalo populations in tanzania, on the average, have the reputation for significantly larger outside spreads than okavango and zambezi delta buffalo. a 44-inch buffalo in the moyowosi, tanzania can be soft bossed “green,” scoring well in rowland ward, while a 34-37 inch outside spread buffalo in the zambezi delta may be mature with hard closed bosses. therefore, scoring by the outside spread alone does not guarantee that one shoots an older hard bossed tending towards post-reproductive buffalo. the minimum score for a rowland ward buffalo trophy is 47 inches, taken from the widest outside spread (halse 1998) (fig. 1). today, very few buffalo make it into rowland ward. not only are sci’s record book minimums easily attainable, but also today the majority of hunters follow this scoring system. current measuring system safari club international (fig. 1) tip-to-tip measurement of horns the combined lengths of both horns are measured, from tip to tip across the forehead. using a measuring cable, measurement is begun at one horn tip and follows the hairlike grain of the horn, around its outer curve to its lowest point. the cable is pivoted and the shortest distance along the underside of the horn to the forward edge of the boss measured, keeping the cable in constant contact with the horn. this line will cross the grain of the horn, not run parallel to it. from the forward edge of the boss, the forehead is bridged directly across to the same point on the other boss edge. the cable is kept taut, not pressed into the boss gap. the process is repeated to the opposite horn tip (fig. 1). width of boss the surface width of each horn boss is measured at its widest point. this measurement should be at an approximate right angle to issn 0075-6458 121 koedoe 47/1 (2004) fig. 2. examples of broomed (top) and non-broomed (bottom) buffalo horns. the longitudinal axis of the horn. once the widest point has been determined, the measurement will include the top surface of the boss, the surfaces of the rear edge plus the surface of any front edge that exists. only hard black horn is measured. so-called ‘green’ horn (which will boil away) or skull bone are not included (fig. 2). all measurements are totaled to produce a score that can be submitted for record book purposes. rowland ward (fig. 1) 1. the greatest outside spread in a straight line is measured, at right angles to the axis of the skull (x-y). 2. the greatest inside spread at the widest point between the curves of the horns (c-d). 3. measuring the length of the longest horn on the outside curve begins at the front lower corner of the boss along the front gandy.qxd 2004/04/14 06:09 page 121 edge, inclining to the outside surface of the horn and continues to the tip (a-b). 4. the spread or tip-to-tip distance is measured (c-d). 5. the width of the wider boss, at its widest point, beginning where the horn meets the skull in the front over the top of the boss and ending where the horn meets the skull at the back. the line of measurement is to be parallel to the axis of the skull (g-h) (fig. 1). for scoring purposes, only the outside spread is recorded. method a random sample of 29 trophies from taxidermists around pretoria were selected and subjectively divided into two sub-samples, 14 broomed down (old bull) trophies and 15 non-broomed (prime bull) trophies (fig. 2). taylor (1988) has provided ageing techniques based on tooth morphology but no attempt was made to accurately age the trophies in the sample. the following parameters were measured for each set of horns: outside spread boss width of both horns at widest point tip-to-tip (combined lengths of both horns koedoe 47/1 (2004) 122 issn 0075-6458 table 1 mean horn parameters (inches) for 14 ‘broomed’ and 15 ‘non broomed’ randomly selected buffalo trophies left right outside tip-to-tip left horn right horn tip boss sci rw boss boss spread length (b) length (c) space (a) space non-roomed 12.78 13 37.8 75.5 35.32 35.23 26.13 4.89 101.3 37.17 broomed 13.63 13.86 36 65 30.35 31.2 28.63 3.5 92.5 36 sci safari club international rw rowland ward table 2 various horn parameters (inches) and resulting sci and rowland ward scores after application of an index (dividing the tip space measurements by the mean of the two individual horn lengths squared), the quantity squared designed to advantage broomed-down animals factor outside tip-to-tip left horn right horn sci rw [a/(b+c)/2]2] spread length length non-broomed 0.55 20.70 41.53 19.43 19.37 55.72 20.79 broomed 0.93 33.48 60.45 28.22 29.02 86.03 33.48 sci safari club international rw rowland ward table 3 summarized means (inches) of sampled buffalo trophy measurements of sci, rowland ward and proposed alternative methods sci new sci new rw new rw new sci rw method method/ method method/ boss space boss space non-broomed 101.3 37.8 55.72 11.39 20.79 4.25 broomed 92.5 36 86.03 24.58 33.48 9.57 sci safari club international rw rowland ward gandy.qxd 2004/04/14 06:09 page 122 across forehead) individual horn lengths of both horns tip space (distance between the two horn tips) boss space (space between the two bosses) (fig. 1). data was recorded in inches, to the nearest 1/8. these parameters were considered to be all the parameters that contribute to both the two and threedimensional descriptors of the size of the “trophy”. the sci and rowland ward scores for each trophy were totaled in both samples. the mean was calculated for each measurement and the two samples compared. the additional parameters measured were used to explore indices that would change the profile of “trophies” in favour of older broomed-down individuals. results the importance of this exercise is to demonstrate that trophy measurement parameters can be employed to score sport-hunted trophies that encourage hunters to select for older post-reproductive animals. table 1 shows that the boss width and the tip space are the only two measurements where broomed-horn trophies scored higher than non-broomed. working with the tip space measurement, it was decided to create a factor with this measurement and the mean of the two individual horn lengths. nonbroomed bulls tend to have individual horn lengths longer than the tip space while broomed bulls have horn lengths close to or even shorter than the tip space (figs. 1 & 2). dividing the tip space measurements by the mean of the two individual horn lengths and then squaring the result derived this factor. the non-broomed measurements became fractions with a mean of 0.55 while the broomed measurements showed a mean of 0.93. the higher the value of the factor, therefore, the longer the tip space measurement in relation to the mean of the individual horn lengths. some of the broomed samples even exceeded 1, showing that the tip space measurement was longer than the mean of the individual horn lengths. where nonbroomed horns scored higher than broomed horns under existing measurement systems, all measurements were multiplied by the factor calculated for each trophy. the means for these calculations of the non-broomed horns were significantly lower than the means of the broomed horns (table 2). as prime bulls age, the boss thickens and begins closing up, thus reducing the distance (boss space) between the two horns. to magnify the broomed trophy score even more, the total trophy score can be divided by the boss space measurement (table 3). one shortcoming in the above analysis is that it requires separating out broomed and non-broomed trophies and computing a separate multiplication factor for each category, something that sci and rw may be unwilling and/or are incapable of doing with the data on hand if the goal is to go back and rectify older records in order to come up with a new set of listings based on recognition of age. it is also unlikely that they will be willing to require the separation of “broomed” versus non-broomed animals, as this can be subjective, and might give quite varying results with a wide standard error if many individuals as opposed to one researcher were making this decision. neither sci nor rowland ward currently recordst boss-space measurements, and only rowland ward collects tip-space measurements as supplemental data. however, both scoring systems measure boss width. future research might focus on analysing the use of standard boss-width measurements combined with “a single multiplication factor”, as opposed to two that can be applied to buffalo trophy measurements across-theboard to encourage a similar selective bias for old age in hunted buffalo. this will be required to enable both sci and rowland ward to easily adjust existing data in their trophy record books. similar analyses (i.e., base circumference) are needed for antelope species. of course the other option is to leave the old records stand, while all new records would apply the new measurement scoring method, as happened with the development of a new javelin in track and field back in the 1980s. issn 0075-6458 123 koedoe 47/1 (2004) gandy.qxd 2004/04/14 06:09 page 123 this would also require sci to begin collecting additional data. regardless of the direction taken, future research must be undertaken in close collaboration and negotiation with the record book associations and their sport hunting members to assure that the results from the research will be applied in a manner that improves the management of the buffalo and/or other species for which research had been undertaken. however, the most difficult challenge confronting the scientific community will not be coming up with these conversion figures, but changing the mind-set of the sport hunters who currently dominate the market, that “bigger is not necessarily better” while providing the scientific data to demonstrate that by modifying their current systems of measurement they are enhancing the economic and possibly the ecological sustainability of the species. conclusions from the above calculations it can be concluded that trophies with factors close to or above 1, are generally much older bulls with worn horn tips. they should be scoring more points in the record books than trophies with low factors. in other words, the closer (or even exceeding it) to 1 the factor is, the better the trophy. the proposed new trophy measuring method computes a multiplication factor created by dividing the tip space measurements the mean of the two individual horn lengths and then squaring the result. with this method, a good trophy should have the following attributes: a wide tip space in relation to individual horn lengths a wide outside spread large boss widths a small boss space there are undoubtedly some factors that may influence new scoring methods—current research in the kruger national park is showing that older bulls act as buffers for predators (bowers, p.o.box 76, skukuza, 1350, rsa pers. comm.). the issue of boss space also needs further investigation, as this appears to open again in very old animals (bowers pers. comm.). in this limited experiment we have seen that a minor change to the trophy measuring system can shift the emphasis on different sectors of a population being utilised and lead to a more conservation-minded approach to trophy hunting of buffalo. future research needs to focus on computing a “single” multiplication factor based upon boss-width measurements, which both record scoring systems collect, while providing scientific evidence that by modifying their scoring systems, the international sport hunter is helping to assure quality wildlife trophies for the next generation of hunters and, as importantly, sustain the role hunting plays to rural and national economies in africa. references bothma, j. du p. 1996. game ranch management. pretoria: van schaik. estes, r.d 1997. the behavior guide to african mammals. halfway house, south africa: russel friedman. grobler, d.g 1996. the potential utilization of the african buffalo with regard to hunting and meat production. in: penzhorn, b.l. (ed.). proceedings of a symposium on the african buffalo as a game ranch animal. pretoria: onderstepoort. halse, a.r.d (ed.). 1998. rowland ward’s records of big game. hartam, south africa: rowland ward. quimby, w.r (ed.). 1993. sci record book of trophy animals, african field edition, vol. 1. tucson, arizona: walsworth. robertson, k. 1996. nyati. a guide to hunting zimbabwe’s most dangerous big game. the southern buffalo. by ‘doctari’. the zimbabwe hunter magazine (now african hunter). harare, zimbabwe: magset. taylor, r.d. 1988. age determination of the african buffalo, syncerus caffer (sparrman) in zimbabwe. african journal of ecology 26(3): 207-220. sinclair, a.r.e. 1977. the african buffalo. chicago: university of chicago press. koedoe 47/1 (2004) 124 issn 0075-6458 gandy.qxd 2004/04/14 06:09 page 124 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 page 14 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page8-1969.pdf page 1 page9-1969.pdf page 1 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 last.pdf page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 article information authors: robert f. brand1 leslie r. brown1 pieter j. du preez2 affiliations: 1applied behavioural ecology and ecosystem research unit, department of environmental sciences, university of south africa, florida campus, south africa 2department of plant sciences, university of the free state, south africa correspondance to: leslie r. brown email: lrbrown@unisa.ac.za postal address: department of environmental sciences, unisa, florida campus, private bag x6, florida, 1710, south africa keywords drakensberg alpine centre; endemism; platberg; red data species; species richness dates: received: 18 june 2009 accepted: 11 oct. 2009 published: 29 mar. 2010 how to cite this article: brand, r.f., brown, l.r., du preez, p.j., 2010, ‘a floristic analysis of the vegetation of platberg, eastern free state, south africa’, koedoe 52(1), art. #710, 11 pages, doi: 10.4102/koedoe.v52i1.710 copyright notice: © 2010. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0771 (online) a floristic analysis of the vegetation of platberg, eastern free state, south africa in this original research article... open access • abstract • introduction    • study area       • location       • geology       • topography and soils       • climate • method • results    • floristic analysis of platberg    • largest plant families on platberg • discussion    • platberg: environmental conditions on inselbergs and floristic comparisons    • endemism, species richness and floristic links with the dac    • platberg and affinities with fynbos and other cape floral elements in high-altitude floras (dac)    • conservation implications for platberg • conclusion • references abstract (back to top) a checklist of vascular plants of platberg was compiled to determine species richness, rarity and endemism. the floristic analysis is part of the department of economic, tourism and environmental affairs free state biodiversity assessment programme and conservation management plan for platberg. the analysis identified a total of 669 species belonging to 304 genera and 95 families, with 214 species belonging to the monocotyledoneae and 438 species to the dicotyledoneae. the largest family is asteraceae with 126 species, followed by poaceae with 73 species, cyperaceae with 39 species, fabaceae with 33 species, and scrophulariaceae with 27 species. various fynbos species were found, as well as 26 endemic/near-endemic species belonging to the drakensberg alpine centre or eastern mountain region. the results of this study revealed that platberg shares inselberg floral richness and endemism that can be tracked via the afromontane archipelago-like string of inselbergs and mountains, which stretch north through the chimanimani mountains, into malawi, the eastern arc mountains via tanzania and north through ethiopia, into eurasia. conservation implications: platberg, as an inselberg, is a site of significant biological diversity, with high species richness, vegetation selection and ecosystem complexity. it shares floral richness and endemism via inselbergs and mountains throughout africa. the high species richness, red data species and ecosystems make this area an important conservation site that should be legislated and protected. introduction (back to top) with the human population steadily on the rise, the natural environment is under greater pressure than ever before (huntley 1991). natural areas act as reservoirs for plant and animal populations (iucn 1980). platberg, overlooking the town of harrismith in the free state, is an inselberg that presents a refuge for indigenous plants and animals (burke 2001; gröger et al. 1996; mutke et al. 2001; porembski et al. 1997, 1998; porembski & brown 1995). vegetation surveys provide information on the different plant communities and plant species present and form the basis of any management plan for a specific area (brown et al. 1996). little is known about the different taxa of platberg and hence a detailed floristic and ecological survey was undertaken to quantify threats to the native flora and to establish whether links exist with higher-altitude afro-alpine flora occurring on the drakensberg. no extensive vegetation surveys had been undertaken on platberg prior to this study; only limited opportunistic floristic collections were done in the mid-1960s by mrs. jacobs (these vouchers were mounted and authenticated in 2006 and are now housed at the geo potts herbarium, botany department, university of the free state). similarly, 50 relevés were sampled between 1975 and 1976 by professor h.j.t. venter, department of genetics and plant sciences, university of the free state. study area location the town of harrismith is located at the foot of platberg (29º10’ s and 28º16’ e), along the n3 highway between johannesburg and durban (figure 1). figure 1: location of platberg, free state platberg’s altitude ranges from 1 900 m to 2 394 m. the surface area covers approximately 3 000 ha. the slopes are steep with numerous vegetated gullies and boulder scree slopes below vertical cliffs that are 20 m to 45 m high. waterfalls cascade down the southern cliffs after rain. a permanent stream arising from the gibson dam on the undulating plateau flows off the escarpment and cascades as a waterfall (figure 2). figure 2: topography of the study area from a distance, platberg appears to have a distinct flat top. however, once on the summit the plateau is found to be undulating, with rolling grass-covered slopes. the vegetation of the plateau is dominated by grassland, with a few rocky ridges, sheet rock and rubble patches, as well as numerous seasonal wetlands and a permanent open playa (pan) on its far western side. woody patches of the genera leucosidea, buddleja, kiggelaria, polygala, heteromorpha and rhus shrubs, as well as the indigenous mountain bamboo thamnocalamus tessellatus, grow along the base of the cliffs. the shrubland vegetation is concentrated on the cool side of platberg, on sandstone of the clarens formation, in gullies, on scree slopes, mobile boulder beds, and on rocky ridges. shrubs and trees also occur in a riparian habitat in the south-facing cleft, in which the only road ascends steeply to the summit. platberg falls within the grassland biome, generally containing short to tall sour grasses (acocks 1988; low & rebelo 1996). the vegetation map of mucina and rutherford (2006) shows platberg as a prominent isolated vegetation ‘island’ with affinities to the drakensberg grassland bioregion, embedded in a lower lying matrix of eastern free state sandy grassland (gm 4). platberg also has elements of fynbos (brand et al. 2008; mucina & rutherford 2006), false karoo (acocks 1998) and succulent karoo (low & rebelo 1996), as well as elements of temperate and transitional forest, specifically highland sourveld (44a, acocks 1988) veld types. geology platberg consists of layers of the karoo supergroup, which stratagraphically lie immediately below the volcanic rocks of the drakensberg formation, the youngest unit of the stormberg group. the 1 900 m contour marks the initiation of the footslopes of platberg and is constituted by the aeolian and waterborne sediments of the clarens formation (du toit 1954; loock et al. 1991; norman & whitfield 1998; truswell 1970). several dolerite dykes intersect platberg, with a close-set sequence of parallel dykes forming the woody/shrub community on the northern and southern slopes. a dolerite dyke is evident just below the gibson dam and is exposed by a borrow pit used for quarrying. numerous other dykes are visible on aerial photographs. these dyke swarms occur mostly on the flat terrain at lower altitudes, with only a few intersecting platberg itself. these dykes are also visible in plantation road-cuts on the southern side of platberg. topography and soils the steep slopes are composed of shale and mostly sandstone of the clarens formation, intersected by dolerite dykes, resulting in benches, deep ravines and terraces. numerous sandstone and basalt boulders form scree and boulder slopes, resulting in miscellaneous soils forms, characteristic of the ib land type, specifically mispah (ms) and glenrosa (gs) soil forms (du preez 1991; macvicar et al. 1991; mucina & rutherford 2006). there are affinities with fa land types’ undulating rocky landscape dominated by shallow ms and gs soil forms (land type survey staff 1991; smit et al. 1993). the summit plateau is composed of igneous rock, specifically basalt and dolerite. these soils are generally shallow (20 mm – 200 mm), with the deepest being < 500 mm. soils are derived from basalt and are shown to have even proportions of course sand, fine sand, clay and silt, with moderate to high organic matter content. the main land type is ea (clay vertic, melanic or red diagnostic horizons) (macvicar 1991; mucina & rutherford 2006; smit et al. 1993) with a clay content of 8% to 60%, depending on the soil series (land type survey staff 1991). climate the mean annual precipitation for the study area ranges between 800 mm and 1 200 mm, with the mean daily maximum temperature fluctuating between 22 ºc and 24 ºc for the hottest month of january. during the coldest months (june and july) the daily mean temperature ranges from -2 ºc to 0 ºc. rainfall occurs throughout the year; the highest rainfall is recorded in the summer months (october to march; figure 3). the daily mean relative humidity for the most humid month, march, varies between 68% and 72%, and the daily minimum relative humidity for july and august varies between 32% and 38% (mucina & rutherford 2006; schulze 1997; south african weather service 2007; van zinderen bakker 1973). figure 3: the average rainfall and temperatures for the study area snow occurs on the top of platberg during most winters and can remain on the ground for up to two weeks, while water in south facing gullies may be frozen for several weeks at a time (mucina & rutherford 2006; van zinderen bakker 1973). possible heavy soils are present on mtabazwe, the highest point on platberg (hilliard & burtt 1987) and may be a significant factor affecting vegetation at high altitudes (hilliard & burtt 1987; mucina & rutherford 2006). the climate of platberg and the drakensberg is part of a continental, north-south trending austro–afro-alpine climate, with noticeably higher precipitation than the surrounding lowland. precipitation along the drakensberg escarpment is over 1 400 mm in some areas, with a gradual decrease both to the north and south, as well as westwards (560 mm) to the drier sub-humid continental climate towards bloemfontein, some 400 km away (scott 1988; van zinderen bakker 1973). precipitation is orographic, not only from violent thunderstorms in summer, but also from dense mist and fog (smit et al. 1993) as well as snow. according to cowling et al. (1992), climatic heterogeneity associated with montane landscape is a more important determinant of plant diversity than geological heterogeneity. method (back to top) ad hoc plant collections and reconnaissance surveys commenced in 2003 and 2004. the main collections associated with the braun–blanquet survey on platberg were made from october 2005 through to january 2007. these collections were of plants growing exclusively in the 393 sample plots, which comprised the entire survey for this study. all specimens were pressed, dried and labelled according to standard field and herbaria practices. identification was done at the geo potts herbarium, ufs. problematic material was identified at the national herbarium of the south african national biodiversity institute (sanbi) in pretoria. all floristic material currently resides at the geo potts herbarium. the final species list was compiled from material collected at platberg for this study. this species list was compiled from all collections of platberg housed at the ufs and sanbi–pretoria herbaria, supplemented by information contained in numerous reference books.for trees and wildflowers the following books were consulted: hilliard (1990, 1992), hilliard and burtt (1987), killick (1990), pooley (1997, 1998, 2003), and van wyk and smith (2001); for grasses: gibbs-russell et al. (1991), moffett (1997) and van oudtshoorn (1999); and for the sedges: gordon-grey (1995). information compiled by germishuizen et al. (2006) was used for nomenclature, the latest taxonomic changes and arrangement, as well as an author update. alien species (introduced exotics) are marked with an asterisk (*). the conservation status of plants was checked against the national red list of south african plants (2009), as well as golding (2002), hilton-taylor (1996) and scott-shaw (1999). endemic or near-endemic classification was ascertained from carbutt & edwards (2004, 2006), pooley (2003) and scott-shaw (1999). the complete species list for platberg (tables 2−4 in online appendix) was sorted according to the englerian system (sanbi–pretoria) that conforms to the system used by arnold and de wet (1993). red data status was included, where applicable. throughout this discussion the term drakensberg alpine centre (dac) will be used in line with the terminology and updated nomenclature as followed by carbutt and edwards (2006). results (back to top) an assumption and basic hypothesis has been that platberg is linked, phytosociologically to, and is a floristic extension of, the dac. the phytosociological analysis of platberg as described by brand (2007) and brand et al. (2008) shows strong links with the dac and supports this assumption. the dac is regarded as the only true alpine region in southern africa (carbutt & edwards 2004; killick 1963, 1978a) and, together with the cape floristic region (cfr), contributes significantly to the flora of africa (hilliard & burtt 1987), via east and west coast mountain tracks (mutke et al. 2001). the flora of the dac and, by extension, platberg is an amalgamation of numerous floras from both temperate and subtropical as well as afromontane sources (hilliard & burtt 1987) and contains significant taxonomic influences from the cfr (goldblatt & manning 2000). the dac and cfr link is reported in studies of the eastern cape drakensberg region by hoare and bredenkamp (2001), in studies of the stormberg and drakensberg and the grasslands of southern kwazulu-natal by bester (1998), and in studies of the adjacent drakensberg regions by perkins (1997) and perkins et al. (1999a, 1999b). according to van wyk and smith (2001) the total number of species for the dac is about 2 200, with no endemic/near-endemic families, four endemic/near-endemic genera, and > 400 endemic/near-endemic species/taxa (18.2%), with 5% succulents amongst the endemics. other surveys have covered smaller regions. there exists no complete survey for the entire drakensberg that provides comprehensive and accurate figures. table 1 presents a summary of regional floras of the dac and includes the most recent figures for a total of 2 520 species, with no endemic families and six near-endemic genera (carbutt & edwards 2004). table 1: a summary of regional floras of the drakensberg alpine centre floristic analysis of platberg the platberg vascular plant species list comprised a total of 669 species, represented by 304 genera and 95 families (table 1 in online appendix). the flowering plants are represented by monocotyledoneae with 214 species from 23 families (23% of total number of families) and dicotyledoneae with 438 species from 63 families (67%). pteridophytes are represented by 16 species and eight families (8.4%), and gymnosperms by one family and one species (1%). the complete species list, separated into the different divisions and sorted by family, appears in the online appendix. largest plant families on platberg the 13 largest families comprised 411 species from 158 genera, which represented 50.7% of the angiosperms and 61.4% of the total flora thus far recorded for platberg, while the remaining 82 families reflected the other 38.6%.the two largest families were asteraceae with 126 species from 40 genera, constituting 18.8% of the total flora, and poaceae with 73 species from 39 genera, constituting 10.9 % of the total flora. with a significantly lower number of species, the next 11 largest families (in descending order) were the following: • cyperaceae with 39 species from 18 genera (5.8%) • fabaceae with 33 species from 13 genera (4.9%) • scrophulariaceae with 27 species from 13 genera (4.0%) • hyacinthaceae with 21 species from 10 genera (3.1%) • orchidaceae with 16 species from eight genera (2.4%) • iridaceae with 17 species from six genera (2.5%) • crassulaceae with 13 species from three genera (1.9%) • geraniaceae with 12 species from three genera (1.8%) • thymelaeaceae with 12 species from three genera (1.8%) • lobeliaceae with 12 species from three genera (1.8%) • campanulaceae with 10 species from two genera (1.5%). all other families had fewer than 10 species. the pteridophytes comprised 8.5% of the total number of families for platberg and gymnosperms comprised 1%. angiosperms represented 90.5%, with a breakdown into monocotyledons constituting 26.7% and dicotyledons constituting 73.3% of vascular plants. the largest 17 genera each had five species or more, with the genera helichrysum and senecio each having a considerably larger number of species (31 and 30 species, respectively) than any of the other genera (table 2). twenty-nine red data, endemic or near-endemic species were found to represent 4.3% of the total flora for platberg, all of which were angiosperm (table 3). introduced species (see tables 2−4 in online appendix) amounted to 22, which represented 3.3% of the total plant species. three of the six near-endemic genera for the dac were found on platberg, two of which were monocotyledoneae: the mountain bamboo thamnocalamus (poaceae) and rhodohypoxis (hypoxidaceae), and one the dicotyledoneae craterocapsa (campanulaceae). the other remaining three near-endemic genera, not recorded on platberg, were guthriea (achariaceae), huttonaea (orchidaceae) and glekia, which made a total of 11 endemic or near-endemic genera for the dac (carbutt & edwards 2004). table 2: ranking of the 17 most dominant genera for platberg table 3: a compilation of endemic/near-endemic and threatened taxa, and their conservation status for platberg discussion (back to top) an analysis of the flora for platberg (online appendix 1) shows that the largest family is asteraceae, with 126 species (18.8% of the total species). this is not unusual for the grassland biome or the dac (carbutt & edwards 2004), or even for south africa, where in semi-arid to arid areas asteraceae is the largest family (goldblatt & manning 2000). the poaceae is the next largest family (10.9%), with the majority of grasses using c4 metabolism, which is characteristic of adaptations to high temperatures and low rainfall. this is somewhat analogous for platberg, with its higher altitude and rainfall as well as its close proximity to the drakensberg massif. however, a more detailed analysis shows the composition of grasses to be a mix of c3 and c4, with about a quarter using a combination of the metabolic pathways of both (brand 2007). the position of fabaceae (fourth largest, 4.9%) and scrophulariaceae (fifth largest, 4%) is also a reflection of their position globally, as well as across drier parts of africa (goldblatt & manning 2000). the high value of cyperaceae is not unexpected; it shows similar patterns for south africa and related areas. a comparison of cyperaceae for inselbergs (brand 2007) confirms these high values for cyperaceae, which may be a result of the harsh conditions found on inselbergs that favour cyperaceae development and radiation (gröger & barthlott 1996). a comparison of the ratio of monocotyledoneae to dicotyledoneae for platberg (table 2) is 1:2.15, which is almost identical to that for korannaberg of 1:2.16 (du preez 1992). however, the ratio for mountain zebra national park, eastern cape, is 1:8 (pond et al. 2002), which is higher than the average for the cfr of 1:3 (goldblatt & manning 2000). the latter is the average for floras worldwide (goldblatt & manning 2000). a comparison of the five largest angiosperm families of platberg, the dac and other relevant areas shows striking similarities. table 4: summary of the relationship between numbers of families, genera and species for platberg and korannaberg twenty-nine red data, endemic and near-endemic species (all of which are angiosperms) comprise 4.3% of the total flora of platberg (table 3), which is lower than for the dac at 7.2% (carbutt & edwards 2004). platberg: environmental conditions on inselbergs and floristic comparisons inselbergs represent sites with extreme environmental conditions (gröger & barthlott 1996), which include high levels of daily temperature change, namely 8 °c to 28 °c for december and -2 °c to 18 °c for july (schulze 1997). frost is another important environmental factor for inselbergs; the average number of days with frost is between 61 and 120 days (schulze 1997). shallow soils, characteristic of large sections of inselbergs, have limited water retention, which results in harsh conditions for the vegetation; intermittent water is a major limiting factor, resulting in edaphic and micro-climatic xeric islands (gröger & barthlott 1996; körner 2003).a comparison of the inselberg vegetation of platberg and korannaberg is shown in table 4, with a close correlation of taxa and their numbers. there is a clear topographic diversity pattern associated with inselbergs. climate and topography are only part of the explanation for high biodiversity; it is rather the diversity of plant communities that provides a measure of habitat diversity (cowling & lombard 2002) and is a reflection of the response to climate change (mucina & rutherford 2006; scott et al. 1997). there is a high degree of similarity between the angiosperm flora of platberg and korannaberg (table 4), with the higher numbers for korannaberg possibly the result of its much larger area (cowling & lombard 2002; macarthur & wilson 2001) – as the surface area increases more species can be supported (gröger & barthlott 1996; linder 2003). additionally, it was noted that the higher species numbers for korannaberg could be due to its position, which has strong afromontane, nama karoo and savanna floristic influences (du preez 1991, 1992; du preez et al. 1991; du preez & bredenkamp 1991). compared to other regional floras, platberg has one of the highest species-to-area ratios (figure 4). for the bourke’s luck site, the figure of 5.244 is converted from 524.4 hectares, selected from the total reserve size of 26 818 hectares, with similar rainfall (700 mm – 1 400 mm per annum), slightly higher temperatures, and situated on the escarpment at 1 640 m to 1 900 m (brown et al. 2005; mucina & rutherford 2006). figure 4: a comparison of regional floras and their surface areas endemism, species richness and floristic links with the dac phytosociologically, geologically, topographically, climatically and biogeographically, platberg may be seen as an extension of the dac. it would then be expected that strong floristic affinities would be seen between the two sites (table 5). the dac is seen as a transition zone, a migratory pathway and repository for taxa of diverse regions and biomes (carbutt & edwards 2004; hilliard & burtt 1987; killick 1963, 1978a, 1978b; mucina & rutherford 2006). the dac has close affinities with the cfr. this is particularly seen in the fynbos vegetation unit elements found on platberg and the dac, and described as the gd 9 drakensberg–amathole afromontane fynbos, as well as the northern escarpment afromontane fynbos (gm 24; mucina & rutherford 2006). other floristic affinities are seen with the magaliesberg, with the gm 29 waterberg–magaliesberg summit sourveld (mucina & rutherford 2006), and the surrounding bankenveld, witwatersrand areas (behr & bredenkamp 1988; bredenkamp & brown 2003; grobler et al. 2002, 2006). floristic links between the dac, kwazulu-natal and the cfr are elucidated in studies by bester (1998), hill (1996), hoare and bredenkamp (2001), pond et al. (2002), perkins (1997), perkins et al. (1999a), smit et al. (1993, 1995), and described as the gs 1 northern zululand mistbelt grassland (mucina & rutherford 2006). table 5: a comparison and ranking of the larger families on platberg and the dac that contribute 1% or more species to the total angiosperm flora in both regions the ecological survey undertaken by kay et al. (1993) of the golden gate national park does not give a floristic analysis with the breakdown into the numbers of families, genera and species. the phytosociological table given lists a total of 114 species and makes no reference to a total species count. the paper by roberts (1969) on the vegetation of the golden gate park lists a total of 331 vascular plants and is ‘a list of the more abundant species present’ (roberts 1969). it is thus not possible to make comparisons with the flora of the only other site that has high-altitude grasslands in the proximity of platberg. the floristic composition for platberg and the related dac shows asteraceae as the largest family (table 5). this is also the case for the cape flora, with significant correlation within the top 12 to 20 families (goldblatt & manning 2000). fabaceae is the second largest family, followed by aizoaceae, ericaceae and iridaceae, which is a unique aspect of the cape flora (goldblatt & manning 2000). these families are thus represented in the top 10 for platberg as well as the dac. the second most speciose family, poaceae, followed by cyperaceae, reflects platberg’s position in the grassland biome with similar floral composition to the dac (carbutt & edwards 2004, 2006; mucina & rutherford 2006) and the more grassy regions to the west and north (du preez 1991; du preez & bredenkamp 1991; eckhardt et al. 1993a, 1993b, 1995; fuls 1993; kooij et al. 1990a, 1990b, 1990c; malan 1998). this is in contrast to the cape flora, where poaceae and cyperaceae are poorly represented, with restionaceae filling the environmental and floristic position of the poaceae (table 5; goldblatt & manning 2000). platberg and affinities with fynbos and other cape floral elements in high-altitude floras (dac) south africa has about 20 500 vascular plant taxa (germishuizen et al. 2006; goldblatt & manning 2000; national red list of south african plants 2009). the cfr has the richest flora for its area, with 9 000 species (goldblatt & manning 2000), as it is an important centre of endemic species and radiation for genetic material (linder 2003). as the cfr is confined to the southern tip of the african continent and thus at a ‘dead end’, species are forced to migrate to the north (linder 2003; mucina & rutherford 2006). a floristic comparison with the cfr would elucidate links with high-altitude taxa that may have migrated out of the cfr, through the dac, and further north. the cfr is also the largest contributor to dac flora in the kwazulu-natal drakensberg, with 67 genera (carbutt & edwards 2004). the number of cfr elements decrease further northwards from the dac (carbutt & edwards 2004). however, unlike the cfr (table 6), the largest families for the dac show similar trends with mount mulanje and the nyika plateau, with the exception of orchidaceae (which is the most plentiful family), represented by 122 and 200 species respectively for these centres (burrows & willis 2005). the next three highest species-rich families are asteraceae, poaceae and fabaceae, with rubiaceae the fifth largest family (82 and 90 species respectively). a further floristic comparison of endemics and near-endemics with the dac and the afromontane archipelago, complied by burrows & willis (2005), shows the following: table 6: a comparison of the largest families for platberg, drakensberg alpine centre and cape floral region • nyika plateau (malawi) comprises 1 891 total species, of which 46 are endemic or near-endemic. • mount mulanje (malawi) comprises 1 303 total species, of which 94 are endemic or near-endemic. • chimanimani mounatins (mozambique) comprises 859 total species, of which 70 are endemic or near-endemic. • simien highlands (ethiopia) comprises 550 total species, of which 12 (plus) are endemic or near-endemic. • mount cameroon (cameroon) comprises 2 435 total species, of which 49 are endemic or near-endemic. the largest family for all three regions is asteraceae, with 18.9% for platberg, 17.1% for the dac, and 11.5% for the cape. comparing the density of asteraceae within south africa (11%) to other regions of the world, we find that hawaii comprises 15.9%, the sonora desert 15%, texas 13.4%, eastern north america 12.7%, new zealand 12.5% and europe 12% (goldblatt & manning 2000; hilliard & burtt 1987). the dac has a significantly higher percentage than the cape. the percentage for platberg is higher than for the dac, which is significantly higher than for hawaii (which has the largest percentage globally). asteraceae show significant spread in the dac and more so on platberg. these areas have been geologically stable for a considerable time (mccarthy & rubidge 2005), and have many different abiotic parameters that are an important for high speciation (mutke et al. 2001). terrain heterogeneity and a relatively stable quaternary climate (scott et al. 1997) may have enabled this genus to spread and establish so successfully at altitude (linder 2003). alternatively, this may be a recent phenomenon, where new taxa respond rapidly to a new environment and fill available niches (linder 2003), or perhaps it is a response to climate change over longer periods, possibly from the start of the holocene epoch 30 million years ago. the low species numbers for poaceae in the cfr is explained by it being a replacement genus for restionaceae that fills its niche and thus is more plentiful (carbutt & edwards 2006; haaksma & linder 2001; linder 2003). in the cfr, restoniaceae also displaces cyperaceae (goldblatt & manning 2000), which may partially explain the low number of cyperaceae species and its ranking at twelfth for the cfr, third for platberg and sixth for the dac (table 6) as the environmental parameters for all regions are analogous (stock et al. 2004). conservation implications for platberg the floristic analysis of platberg shows its value as both a site of diversity and endemism. this value is also linked to the dac flora and shows its position as the main inselberg in the chain of such prominent landscape features. the high plant diversity of platberg and its main genetic source, the dac, is because it includes elements of mesic karoo-nama vegetation, the afromontane podocarpus elements, mesic kwazulu-natal escarpment, and drier eastern cape stormberg and lesotho floras. platberg has high terrain heterogeneity, with steep contours, deeply incised gullies and numerous slope orientations. according to cowling et al. (1992), low & rebelo (1996), mucina and rutherford (2006), perkins (1997) and van wyk and smith (2001) this provides more habitats for species and is more important than other factors in determining high biodiversity. in addition, platberg is the single largest and best preserved high-altitude grassland in the free state (mucina & rutherford 2006). immediate threats are from the invasion of pinus patula on the footslopes and plateau area of platberg to the south and east. this poses a considerable conservation problem because pinus patula is in the process of rapidly out-competing the local indigenous vegetation, particularly on the cool south slope and in the stream flowing down the gully on the south side of platberg. the threats to genetic diversity (anderson 2001) also influenced by global warming are key considerations for conservationists and highlight the importance of mountains and the associated inselbergs as areas of refuge to plant species. this, therefore, stresses the urgent need for their protection. additionally, high-altitude vegetation, specifically alpine flora as defined by white (1983) and körner (2003), is the only biogeographic unit on land with a global distribution, accounting for nearly 3% of the land surface and including about 10 000 species, or about 4% of higher order plants (körner 2003). the threat posed by global warming to the world’s alpine regions, including africa, was predicted by peters (1992), who indicated that as little as a 3 ºc increase in temperature over 100 years would be equivalent to a 500 m upward shift in altitudinal zones. in the afromontane and afro-alpine biota, this would cause a significant reduction in the distribution of plants, changing their structure and composition, and forcing some taxa to higher altitudes (taylor 1996). furthermore, in some cases (e.g. platberg), a significant reduction of the taxa could possibly occur where the altitude is under 2 500 m. conclusion (back to top) platberg is centred in the grassland biome, with a lesser mix of fynbos, nama-karoo and forest biomes. platberg consists of 3 000 ha of high-altitude grassland composed of a mix of c3 and c4 grasses and other plant families that use this metabolic pathway. between 30% and 55% of the grassland biome has been transformed, only 5.5% of which is under protection. numerous abiotic environmental factors (soil, altitude, slope, aspect, moisture availability, temperature range, fire, modern and palaeoclimatic factors) and biotic interactions between plants and plants, plants and pollinators, and herbivores and plants have resulted in a relatively high species richness and diversity. the drakensberg alpine centre is the main floristic influence on platberg, with strong affinities with the cape floral region. the result of this study has shown that platberg is a centre of significant biological diversity, with high species richness, vegetation structure and ecosystem complexity, and a centre of genetic diversity and variation. it occurs as an island in the grassland ‘sea’ and shares inselberg floral richness and endemism that can be tracked via the afromontane archipelago-like string of inselbergs and mountains, which stretches north through the chimanimani mountains, into malawi, the eastern arc mountains via tanzania and north through ethiopia, into eurasia. it also shows a western tract via the congo, ivory coast and cameroon inselbergs and mountains. the current vegetation patterns on platberg reflect changes in palaeo-environmental cycles of cooling and warming, which, since miocene times, have had the greatest influence on the texture and composition of plants and speciation on platberg. current trends towards climate change will alter the composition and species numbers of grasses as well as other plant families on platberg. as an important high-altitude grassland, it is imperative that platberg be provided with protection legislated on at least a provincial level. references (back to top) acocks, j.p.h., 1988, ‘veld types of south africa’, 3rd edn., memoirs of the botanical survey of south africa 57, 1–146. anderson, j.m. (ed.), 2001, towards gondwana alive, national botanical institute, pretoria. arnold, t.h. & de wet, b.c. (eds.), 1993, ‘plants of southern africa: names and distribution’, memoirs of the botanical survey of south africa 62. behr, c.m. & bredenkamp, g.j., 1988, ‘an analysis of the flowering plants and ferns of the witwatersrand national botanic garden’, south african journal of botany 54(6), 515–524. bester, s.p., 1998, ‘vegetation and flora of the southern drakensberg escarpment and adjacent areas’, msc thesis, university of pretoria. brand, r.f., 2007, ‘phytosociology of platberg mountain, eastern free state, south africa’, phd thesis, university of the free state. brand, r.f., du preez, p.j., & brown, l.r., 2008, ‘a floristic description of the afromontane fynbos communities on platberg, eastern free state, south africa’, koedoe 50(1), 202–213. bredenkamp, g.j. & brown, l.r., 2003, ‘a reappraisal of acocks’ bankenveld: origin and diversity of vegetation types’, south african journal of botany 69(1), 7–26. brown, l.r., bredenkamp, g.j. & van rooyen, n., 1996, ‘the phytosociology of the northern section of the borakalalo nature reserve, south africa’, koedoe 39(1), 9–24. brown, l.r., marais, h., henzi, s.p. & barrett, l., 2005, ‘vegetation classification as the basis for baboon management in the bourke’s luck section of the blyde canyon nature reserve, mpumalanga’, koedoe 48(2), 71–92. burke, a., 2001, ‘determinants of inselberg floras in arid nama karoo landscape’, journal of biogeography 22, 1211–1220. burrows, j. & willis, c. (eds.), 2005, ‘plants of the nyika plateau: an account of the vegetation of the nyika national parks of malawi and zambia’, sabonet report 31, southern african botanical diversity network, pretoria. carbutt, c. & edwards, t.j., 2004, ‘the flora of the drakensberg alpine centre’, edinburgh journal of botany 60(3), 581–607. carbutt, c. & edwards, t.j., 2006, ‘the endemic and near-endemic angiosperms of the drakensberg alpine centre’, south african journal of botany 72, 105–132. cowling, r.m., holmes, p.m. & rebelo, a.g., 1992, ‘plant diversity and endemism’, in r.m. cowling (ed.), the ecology of fynbos: nutrients, fire and diversity, oxford university press, cape town. cowling, r.m. & lombard, a. t., 2002, heterogeneity, speciation/extinction history and climate: explaining regional plant diversity patterns in the cape floristic region, diversity and distributions 8, 163–179. du preez, p.j., 1991, ‘a syntaxonomical and synecological study of the vegetation of the south-eastern orange free state and related areas with special reference to korannaberg’, phd thesis, university of the orange free state. du preez, p.j., 1992, ‘the classification of the vegetation of korannaberg, eastern orange free state. i. afromontane fynbos communities’, south african journal of botany 58(3), 165–172. du preez, p.j. & bredenkamp, g.j., 1991, ‘vegetation classes of the southern and eastern orange free state (republic of south africa) and the highlands of lesotho’, research of the bloemfontein national museum, bloemfontein 7, 477–526. du preez, p.j., bredenkamp, g.j. & venter, h.j.t., 1991, ‘the syntaxonomy and synecology of the forests in the eastern orange free state, south africa. l. the podocarpetalla latifolii’, south african journal of botany 57, 198–206. du toit, a.l., 1956, the geology of south africa, oliver and boyd, edinburgh. eckhardt, h.c., van rooyen, n. & bredenkamp, g.j., 1993a, ‘the phytosociology of the thicket and woodland vegetation of the north-eastern orange free state’, south african journal of botany 59(4), 401–409. eckhardt, h.c., van rooyen, n. & bredenkamp, g.j., 1993b, ‘the vegetation of the north-eastern orange free state, south africa: physical environment and plant communities of the ea land type’, bothalia 23(1), 117–127. fuls, e.r., 1993, ‘vegetation ecology of the northern orange free state’, phd thesis, university of pretoria. germishuizen, g., meyer, n.l., steenkamp, y. & keith, m. (eds.), 2006, ‘plants of southern africa: an annotated checklist’, sabonet report 41, national botanical institute, pretoria. gibbs-russell, g.e., watson, l., koekemoer, m., smook, l., barker, n.p., anderson, h.m. et al., 1991, ‘grasses of southern africa’, memoirs of the botanical survey of south africa 58. goldblatt, p. & manning, j., 2000, ‘cape plants: a conspectus of the cape flora of south africa’, strelitzia 9. golding, j.s., 2002, southern african plant red data lists, report no. 14, national botanical institute, pretoria. gordon-gray, k.d., 1995, ‘cyperaceae in natal’, strelitzia 2. grobler, c.h., bredenkamp, g.j. & brown, l.r., 2002, ‘natural woodland vegetation and plant species richness of the urban open spaces in gauteng, south africa’, koedoe 45(1), 19–34. grobler, c.h., bredenkamp, g.j. & brown, l.r., 2006, ‘primary grassland communities of the urban open spaces in gauteng, south africa’, south african journal of botany 72, 367–377. gröger, a. & barthlott, w., 1996, ‘biogeography and diversity of the inselbergs (laja) vegetation of southern venezuela’, biodiversity letters 3, 165–179. haaksma, e.d. & linder, h.p., 2000, restios of the fynbos, the botanical society of south africa, cape town. hill, t.r., 1996, ‘description, classification and ordination of the dominant vegetation communities, cathedral peak, kwazulu-natal drakensberg’, south african journal of botany 62, 263–269. hilliard, o.m., 1990, flowers of the natal drakensberg, university of natal press, pietermaritzburg. hilliard, o.m., 1992, trees and shrubs of the natal drakensberg, university of natal press, pietermaritzburg. hilliard, o.m. & burtt, b.l., 1987, the botany of the southern natal drakensberg, national botanical gardens, cape town. hilton-taylor, c., 1996, red data list of southern african plants, national botanical institute, pretoria. hoare, d.b. & bredenkamp, g.j., 2001, ‘syntaxonomy and environmental gradients of the grasslands of the stormberg/drakensberg mountain region of the eastern cape, south africa’, south african journal of botany 67, 595–608. huntley, b.j. (ed.), 1991, ‘challenges to maintaining biotic diversity in a changing world’, in biotic diversity in southern africa, pp. xii–xix, oxford university press, cape town. iucn, 1980, world conservation strategy: living resource conservation for sustainable development, iucn:unep/wwf, gland, switzerland. jacot guillarmod, a., 1971, flora of lesotho, verlag von j. cramer, lehre. kay, c., bredenkamp, g.j. & theron, g.k., 1993, ‘the plant communities of the golden gate highlands national park in the north-eastern orange free state’, south african journal of botany 59(4), 442–449. killick, d.j.b., 1963, ‘an account of the plant ecology of the cathedral peak area of the natal drakensberg’, memoirs of the botanical survey of south africa 34. killick, d.j.b., 1978a, ‘the afro-alpine region’, in m.j.a. werger (ed.), biogeography and ecology of southern africa, pp. 515–560, junk, the hague. killick, d.j.b., 1978b, ‘notes on the vegetation of the sani pass area of the southern drakensberg’, bothalia 12, 537–542. killick, d.j.b., 1990, a field guide to the flora of the natal drakensberg, jonathan ball and ad donker publishers, johannesburg. kooij, m.s., bredenkamp, g.j. & theron, g.k., 1990a, ‘classification of the vegetation of the b land type in the north-western orange free state’, south african journal of botany 56(3), 309–318. kooij, m.s., bredenkamp, g.j. & theron, g.k., 1990b, ‘the vegetation of the deep sandy soils of the a land type in the north-western orange free state, south africa’, botanical bulletin academia sinica 31, 235–243. kooij, m.s., bredenkamp, g.j. & theron, g.k., 1990c, ‘the vegetation of the north-western orange free state, south africa’, bothalia 20(2), 214–248. körner, c., 2003, alpine plant life: functional plant ecology of high mountain ecosystems, springer-verlag, berlin. land type survey staff, 1991, land types of the map 2828, harrismith, soil and irrigation research institute, department of agriculture, pretoria. linder, h.p., 2003, ‘the radiation of the cape flora, southern africa’, biological reviews of the cambridge philosophical society 78, 597–638. loock, j.c., praekelt, h.e., van der westhuizen, w.a. & welman, j., 1991, the dewetsdorp-thaba nchu mountain excursion, bloemfontein national museum, bloemfontein. low, a.b. & rebelo, a.g. (eds.), 1996, vegetation of south africa, lesotho and swaziland, department of environmental affairs & tourism, pretoria. macarthur, r.h. & wilson, e.o., 2001, the theory of island biogeography, princeton university press, princeton. macvicar, c.n., loxton, r.f., lambrechts, j.j.n., le roux, j., de villiers, j.m., verster, e., et al., 1991, soil classification: a taxonomic system for south africa, department of agricultural development, soil and irrigation research institute, pretoria. (memoirs on the agricultural natural resources of south africa 15). malan, p.w., 1998, ‘vegetation ecology of the southern free state’, phd thesis, university of the orange free state. mccarthy, t. & rubidge, b., 2005, the story of earth and life, struik publishing, cape town. moffett, r., 1997, ‘grasses of the eastern free state’, uniqua, qwaqwa campus of the university of the north, phuthaditijhaba. moffett, r.o., daemane, m.e., pitso, t.r., lentsoane r. & taoana, t.r.n., 2001, ‘a checklist of the vascular plants of qwaqwa and notes on the flora and vegetation of the area’, uniqwa research chronicle 3(2), 32–83. mucina, l., & rutherford, m.c., 2006, ‘the vegetation of south africa, lesotho and swaziland’, strelitzia 19. mutke, j., kier, g., braun, g., schultz, c.h.r. & barhlott, w., 2001, ‘patterns of african vascular plant diversity – a gis based analysis’, systematic geography plantae 71, 1125–1136. norman, n. & whitfield, g., 1998, geological journeys: a traveller’s guide to south africa’s rocks and landforms, de beers, south africa. perkins, l., 1997, ‘aspects of the syntaxonomy and synecology of the grasslands of southern kwazulu-natal’, msc thesis, university of pretoria. perkins, l., bredenkamp, g.j. & granger, j.e., 1999a, ‘the phytosociology of the incised river valleys and dry upland savannah of the southern kwazulu-natal’, south african journal of botany 65(5&6), 321–330. perkins, l., bredenkamp, g.j. & granger, j.e., 1999b, ‘wetland vegetation of the southern kwazulu-natal, south africa’, bothalia 30(2), 175–185. peters, r.l., 1992, ’conservation of biological diversity in the face of climatic change’, in r.l. peters & t.e. lovejoy (eds.), global warming and biological diversity, pp. 15–30, new haven. phillips, f.p., 1917, ‘a contribution to the flora of the leribe plateau and environs’, annals of the south african museum 16, 1–379. pond, u., beesley, b.b., brown, l.r. & bredenkamp, h., 2002, ‘floristic analysis of the mountain zebra national park, eastern cape’, koedoe 45(1), 35–57. pooley, e., 1997, a complete field guide to trees of natal zululand and transkei, natal flora publications trust, durban. pooley, e., 1998, a field guide to wild flowers of kwazulu-natal and the eastern region, natal flora publications trust, durban. pooley, e., 2003, mountain flowers: a field guide to the flora of the drakensberg and lesotho, natal flora publications trust, durban. porembski, s. & brown, g., 1995, ‘the vegetation of inselbergs in the comoé national park (ivory coast)’, conservatoire et jardin botanicues de geneve 50(2), 351–365. porembski, s., fisher, e. & biedenger, n., 1997, ‘vegetation of inselbergs, quartzitic outcrops and ferricretes in rwanda and eastern zaire (kivu)’, bulletin du jardin botanique national de belgique 66, 81–95. porembski, s., martinelli, g., ghlemüller, r. & barthlott, w., 1998, ‘diversity and ecology of saxicolous vegetation mats on inselbergs in the brazilian atlantic rainforest’, diversity and distribution 4, 107–119. roberts, b.r., 1969, ‘the vegetation of the golden gate highlands national park’, koedoe 12, 15–28. sanbi, 2009, national red list of south african plants, viewed 23 february 2009, from http://www.sanbi.org/biodiversity/redlist/2003-02-2009 schulze, r.e., 1997, ‘south african atlas of agrohydrology and climatology’, wrc report tt 82/96, water research commission, pretoria. scott, l., 1988, ‘late quaternary vegetation history and climatic change in the eastern orange free state, south africa’, south african journal of botany 55(1), 107–116. scott, l., anderson, h.m. & anderson, j.m., 1997, ‘vegetation history’ in r.m. cowling, d.m. richardson & s.m. pierce, (eds.), vegetation of southern africa, pp. 62–84, cambridge university press, cambridge. scott-shaw, r.c., 1999, rare and threatened plants of kwazulu-natal and neighbouring regions, kwazulu-natal nature conservation services biodiversity division, scientific services directorate, pietermaritzburg. smit, c.m., bredenkamp, g.j. & van rooyen, n., 1993, ‘phytosociology of the ac land type in the foothills of the low drakensberg in north-western natal’, south african journal of botany 59, 203–214. smit, c.m., bredenkamp, g.j. & van rooyen, n., 1995, ‘the grassland vegetation of the low drakensberg escarpment in the north-western kwazulu-natal and north-eastern orange free state border area’, south african journal of botany 61, 9–17. south african weather service, 2007, ‘climate of south africa’. climate statistics for bethlehem from 1961–1990’, wb 40, government printer, pretoria. stock, w.d., chuba, d.k. & verboom, g.a., 2004, ‘distribution of south african c3 and c4 species of cyperaceae in relation to climate and phylogeny’, austral ecology 29, 313–319. taylor, d., 1996, ‘mountains’, in w.m. adams, a.s. goudie & a.r. orme (eds.), the physical geography of africa, pp. 287–306, university press, oxford. truswell, j.f., 1970, an introduction to the historical geology of south africa, purnell press, london. van oudtsthoorn, f., 1999, guide to grasses of southern africa, briza publications, cape town. van wyk, a.e. & smith, g.f., 2001, regions of floristic endemism in southern africa: a review with emphasis on succulents, umdaus press, pretoria. van zinderen bakker jr., e.m., 1973, ‘ecological investigation of forest communities in the eastern orange free state and the adjacent natal drakensberg’, vegetatio 28, 299–334. white, f., 1983, the vegetation of africa: a descriptive memoir to accompany the unesco/aetfat/unso vegetation map of africa, volume xx, unesco, paris. filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 article information author: jane carruthers1 affiliation: 1department of history, university of south africa, south africa correspondance to: jane carruthers email: carruej@unisa.ac.za postal address: department of history, university of south africa, po box 392, unisa 0003, south africa keywords elephant ethics; elephant conservation; southern african literature; discourse analysis dates: received: 21 july 2009 accepted: 10 nov. 2009 published: 20 apr. 2010 how to cite this article: carruthers, j., 2010, ‘romance, reverence, research, rights: writing about elephant hunting and management in southern africa, c.1830s to 2008’, koedoe 52(1), 6 pages. doi: 10.4102/koedoe.v52i1.880 copyright notice: © 2010. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0771 (online) romance, reverence, research, rights: writing about elephant hunting and management in southern africa, c.1830s to 2008 in this review article... open access • abstract • introduction • elephants in africa • nineteenth-century hunters • the early 20th century • after the second world war • writing in the new century • conclusion • acknowledgements • references abstract (back to top) the protection and management of large mammals in africa’s national parks is not a matter to be left solely for the attention of natural science and scientists. the way in which the natural world is conceptualised by the humanities and social sciences is also significant, because nature is cultural as well as scientific. this article is an interdisciplinary appraisal of the manner in which the writing (e.g. discourse, vocabulary) about elephants in various literary and scientific texts has altered over time. it aims to provide an analysis of some of the literature about elephants in order to examine literate society’s changing responses to the hunting and management of elephants in southern africa over the past two centuries. the review suggests that new research questions regarding animal cognition and empathy have been generated by these changing attitudes, in conjunction with fresh directions in ecological understanding. conservation implications: biodiversity conservation is an inexact science, and even the distinction between conservation research and conservation management is not clear-cut. moreover, a degree of emotion is evident in scientific and popular discussions around what should be ‘saved’ and how best this might be achieved. nature is cultural as well as scientific and interdisciplinary insights from the humanities and social sciences may beneficially inform protected area management. introduction (back to top) adams and mcshane (1992:59) have observed of elephants, ‘no other species carries as much symbolic or emotional force. fascination with elephants is hardly a new phenomenon – man has by turns worshipped, idealised, contemplated, or slaughtered elephants, but rarely ignored them.’ elephants seem appropriate as a particular subject for investigation by scholars in many disciplines, including the humanities and the social sciences, because of their status as the largest african land mammals and their enormous economic and ecological significance; they raise emotions and topics surrounding bioethics and animal rights to a greater degree than any other creature (scholes & mennell 2008; wemmer & christen 2008). elephants (and a small number of other animals) are spearheading the global animal rights movement (wemmer & christen 2008). this is a development that environmental historian roderick nash (1989) foresaw many years ago. he argued that, on the basis of the existence of an over-extending network of rights, which started with slaves, women and indigenous people, animals and the environment itself would eventually be liberated. this line of thinking generally resonates with changes in scientific perspective (beinart & hughes 2007; mackenzie 1988; ritvo 1987, 1997; thomas 1984). many natural scientists, in a variety of disciplines today, are considering how to balance rights, pondering questions about whether the rights of biodiversity take precedence over those of individual species, or how best we might attempt to manage the planet for favourable long-term outcomes that benefit the largest number of species. biologists who study elephants are struggling with important questions around the cognitive and self-conscious sentient life of these animals. research in this field might have consequences on future management philosophy (bates, lee, njiraini, poole, sayialel, k., sayialel, s. et al. 2008; bates, poole & byrne 2008; douglas-hamilton et al. 2006; mccomb, baker & moss 2006; mccomb, moss, durant, baker & sayialel 2001). elephants in africa (back to top) ivory has been the major product that has linked africa with the outside world and shaped perceptions of the continent (alpers 1992; carruthers et al. 2008; luxmoore 1991; meredith 2001). modern western writing about elephants generally does not extol the beauty of ivory, perhaps because the demand for ivory is the principal reason why many people in the western world are mourning the diminution of elephant numbers and, as such, it might seem irresponsible, let alone politically incorrect, to harp on ivory’s aesthetic beauty and the satisfaction of ownership. nonetheless, for more than 10 000 years the ‘subtle glowing colour and sensual surface’ of ivory has ensured its prominent position among the luxury goods of the world (luxmoore 1991; ross 1992). this luxury market has existed for centuries and raised concern. in 77 ce, pliny remarked on the reduction of the herds of north africa (alpers 1992; meredith 2001; wickens 1981). a major peak in supply and demand occurred during the 19th century, with the industrialisation of europe and the united states; practical as well as luxury objects made of ivory became very popular with the growing middle class and the market expanded accordingly (oliver & atmore 1967). in the early decades of the 20th century, ivory was used less frequently in the west, but, in the 1970s, demand from asia (for ornaments and signature seals) took its toll, particularly in east africa, leading to a ban on ivory exports from africa (parker & amin 1983; parker 2004; wylie 2008). elephants are not evenly distributed around africa. in many regions they are illegally hunted and their numbers are dwindling to the point of real concern. in southern africa, the debate is intense for the opposite reason; many people believe (sometimes on the basis of incomplete evidence) that there are too many elephants – that there ought to be intervention to limit their numbers and thus minimise their environmental impact (scholes & mennell 2008). the ivory trade has profoundly affected africa and its people. it brought wealth and power, enabling africans to control and exploit each other; it laid the foundations of strong states and encouraged the emergence of hierarchies based on wealth and class (gordon & gordon 1996; thorbahn 1979). initially, ivory was exchanged for iron and other useful metals that contributed to improved methods of cultivation, as well as for cloth, beads and other goods, but, in later centuries, ivory was traded predominantly for firearms and liquor. ivory extraction, however, led to fragile economies based on a single product; it also created an ongoing trading frontier which expanded into the african interior taking with it warfare and slavery, often leaving economic collapse in its wake (curtin et al. 1995; gordon & gordon 1996). in more recent years, the ban on the ivory trade has led to a number of african states becoming financially dependent on donations from animal rights and nature conservation organizations, which has created opportunities for state and private corruption and illegal trade. nineteenth-century hunters (back to top) the texts analysed below are examples of 19th century hunting literature in southern africa, a genre covered by a number of literary critics (gray 1979; hammond & jablow 1970; mckenzie 1988; murray 1993; pratt 1992; ritvo 1987). as colonial settlement increased and expanded during the late-1700s and early 1800s, small parties of trekboers and traders penetrated the northern interior in search of ivory and other products of the hunt for subsistence and trade (guelke 1989; pollock & agnew 1963; ross 1989; van der merwe 1938, 1945). at about the same time, there were scientific expeditions into the interior (e.g. that of andrew smith) and visits by recreational sport hunters, who referred to themselves as ‘naturalists’, (harris 1840, 1852; lye 1975). the voortrekker parties of this period were also significant because they were potential large-scale, permanent settlers, who acted in the belief that ridding the countryside of wildlife was a pioneering necessity in order to create a ‘civilised’ state (carruthers 1995a, 1995b; kruger 1902). the settler commercial hunters – generally boers – have not left literature detailing their exploits; what we know of their activities comes from evidence in legal debates and from literate travellers. one of the earliest and most famous of the visiting hunter-writers was william cornwallis harris. his books initiated a new genre, viz. nature and travel writing on the subject of the ‘excitement’ of hunting wild animals in an ‘untamed’ africa (harris 1840, 1852). harris, a british army officer stationed in india, arrived in south africa in 1836 and devoted a considerable part of his furlough – and his books – to describing how he hunted elephants – for their tusks certainly, but also for ‘science’ and entertainment. his descriptions captured the drama of the hunt and the imagination of his audience. for example, a scene in the magaliesberg of 300 elephants is described as a ‘ grand panorama’, ‘a picture at once soul-stirring and sublime’. after he had ‘attacked’ and killed some of the females in the herd, he realised that the young were deeply attached to their mothers, as one of juveniles walked around a corpse, ‘with touching demonstrations of grief, piping sorrowfully and vainly attempting to raise her with its tiny trunk’. harris later confessed that ‘i had felt compunctions in committing the murder the day before, and now half resolved never to assist in another’ (harris 1852:168–175). despite his ethical sensitivity that what he was doing might be construed as ‘murder’, harris did not desist and his book abounds with descriptions of profligate killing and details of the gory process of extracting tusks. another example of the hunting genre in this period is the well-known, and often quoted, a hunter’s life in south africa, by roualeyn gordon cumming, first published in 1850. cumming longed for an encounter with ‘the noble elephants’ and, when he realised this objective, he killed them at night (cumming n.d.:178–180), torturing them in an experiment to see how long it would take them to die – an incident he described in detached and unemotional language (cumming n.d.:227–228). cumming justified his activities by explaining that the killing of an elephant was ‘so overpoweringly exciting that it almost takes a man’s breath away’ (cumming n.d.:230). nevertheless, he was not insensitive to the fact that the elephant was a ‘wonderful animal’ that had a defined social structure and patterns of behaviour, a well-developed sense of smell and means of communication (cumming n.d.:180). cumming was initially described as ‘bold, enterprising and skilful’, but, a decade or so later, was referred to as ‘an unprincipled man and an indiscriminate slaughterer’ (carruthers 2005:189), as taking pleasure in killing became increasingly morally indefensible behaviour in the west. in addition to the distaste that modern readers have for the reckless and senseless killings perpetrated by these men, it has also become difficult for us to reconcile statements, such as the following, with the actions of the people who made them. delegorgue, for example, writes: what paltry reason can justify the death and destruction of such beautiful, strong and excellent animals? what are a couple of hundred pounds of ivory compared with the long service which such animals might render to man for generations? … i was perfectly conscious of the mischief i was doing but i was a hunter first and foremost … i desired no other; all the animals of creation, whatsoever they may be, are as nothing compared with the elephant. (delegorgue 1997:3–4) perhaps a later writer of the 1930s, robert henriques, in death by moonlight, succinctly expressed what these hunters were experiencing: i hunt big game frankly and brutally because not otherwise can i find the same disappointments and reverses, the same excitements, the same antagonists, and the same exhilarating rewards … above all, i get from it a moment of triumph and of pure emotion which i have never found elsewhere. (henriques, cited in hammond & jablow 1979) hall-martin has suggested that, before white settlement, there may have been around 100 000 elephants in south africa (hall-martin 1992). by 1900, almost all had been exterminated through commercial and recreational hunting, with only four remaining populations totalling fewer than 200 individuals. these were in the knysna area of the western cape coastline (30–50), in the addo district of the eastern cape (130–140), an unknown, but very small, number in the tropical coastal tembe region of maputaland in northern kwazulu-natal, and a few in the mpumalanga lowveld olifants river gorge, on the boundary between the transvaal and mozambique (boshoff, skead & kerley 2002; hall-martin 1992; roche 1996; skead 1980, 2007; whitehouse & hall-martin 2000; whyte 2001). the early 20th century (back to top) the largest remaining population of these four isolated communities of elephants was in the addo area (hoffman 1993). they had survived because of the specific local vegetation, currently referred to as ‘xeric thicket’ (lubke 1996). in order to modernise the economy of the sundays river region after the first world war, the government invested in a large and expensive irrigation scheme to encourage the development of large citrus estates. for the elephants, this was an irresistible attraction: water and food in abundance. local farmers complained bitterly about their losses. after a formal state enquiry, it was decided to kill all of these elephants, despite the opposition of some, like f.w. fitzsimons, director of the port elizabeth museum, whose opinion it was that the deliberate extermination of these elephants, would, upon grounds of deeply-felt general sentiment, and in the interests of science, be received by not only very high and influential circles in south africa, but by the general feeling of the civilised world with condemnation, as a step reflecting no credit upon south africa.(fitzsimons 1920:270–271) however, the decision to exterminate them was taken and the was task assigned to major philip jacobus (jan) pretorius (1877–1945), a descendant of boer hero andries pretorius (1798–1853). major pretorius had led an extraordinarily adventurous life, which he encapsulated in his autobiography (published in england and australia, later translated into dutch and published in amsterdam), entitled jungle man (pretorius 1948). it is noteworthy that jungle man, ostensibly conceived in the literary genre pioneered by harris and cumming, was the first book in this vein written by a local afrikaner. pretorius presented himself as a national hero and the laudatory foreword to the book was written by jan smuts (1870–1950), the famous south african statesman. at the time of the elephant killing, smuts was prime minister of the union of south africa and, somewhat ironically in terms of his praise of the book and of pretorius personally, he was deeply committed to wildlife. smuts wrote: ‘i gladly write a foreword to this amazing book …. i have never seen a more thrilling story of a hunter’s life’ (pretorius 1948:5–6). in the course of just over a year, killing between three and five elephants almost every week, pretorius reduced the number of elephants to about 16. unlike the entertainment and ivory that elephant hunting had offered the sport-hunters, pretorius was obliterating these elephants in a commercial culling operation, merely because they had hampered economic ‘progress’ (hoffman 1993). in the account of killing the addo elephants, pretorius’s book marks a shift in the discourse around elephant hunting in south africa. pretorius’s campaign against the elephants was not a metaphorical ‘military campaign’, it was a real one. the idea was to exterminate systematically an ‘enemy’ in order to reduce their numbers and their threat to human welfare. standing on ladders and injuring oneself on spiny vegetation in the quest of elephants in the addo thicket was not enjoyable, or sporting, by any standard. then too, unlike harris or cumming, pretorius was a paid government official, who earned, probably, around £6000.00, in addition to the sale of elephant and elephant products (hoffman 1993). hunting the addo elephants was, for the first time, placed in the same category as vermin hunting. it was designed only to reduce elephant numbers and to prevent human–elephant contact. it was done, not for sport or ivory, but was a calculated extermination programme. it did not even produce the scientific data which might, at least, have been a potentially useful benefit of the extermination exercise. paradoxically, and for reasons that are not entirely clear, but which almost certainly included pretorius’s own wish, as soon as the numbers of elephant had been drastically reduced, the survivors were given protection within a game reserve, proclaimed by the cape province and upgraded in 1931 to the addo elephant national park (hoffman 1993; kerley & landman 2006). after the second world war (back to top) by the middle of the 20th century, wildlife viewing had replaced wildlife hunting, and south africa’s national parks, in particular, the kruger national park, had become tourist attractions (carruthers 1995b, 2009). values had changed, as had habitats almost throughout the country; ivory was no longer a product from which individuals might make a living. as a result, elephant numbers in certain localities were on the rise. in 1905, the warden of the reserves that later became the kruger national park, estimated that there were 10 elephants living within the reserves, by 1925, this had increased to 100, and continued to rise to 250 by 1936, 450 by 1946, and 1000 by 1956. a helicopter count a decade later (1967) gave a confirmed number of 6 586 (whyte, van aarde & pimm 2003). after the second world war, various changes had an impact on wildlife conservation: for example, the change of government in 1948, the increasing influence of ecology and wildlife biology and the eventual retirement of a park warden who believed that nature would manage itself. consequently, the national parks board employed a growing number of university-trained scientists with the responsibility to manage the fauna and flora scientifically. in terms of the prevailing conservation paradigm of that era (ecological climax and stability) and within the limits of managerial knowledge, the kruger national park was managed on the principle of carrying capacity, in the same way as an extensive cattle ranch (carruthers 2007a, 2007b). it was thus decided in 1965 – on the basis of area size and feed and water requirements – that the optimum number of elephants in the kruger national park ought be stabilised at around 7000 (0.4 elephants/km2) (whyte 2001). surveys and information for tourists linked expanding elephant numbers with the observed loss of large trees and changes to woody cover, which were held to be the result of ‘too many’ elephants, even though these ecological alterations might not have been the consequence of elephant activity alone (whyte, van aarde & pimm 2003). elephants were culled regularly in the kruger national park for almost thirty years. not surprisingly, culling did not generate a literary genre, but it was recorded. the annual reports of the kruger park carried details of the elephants killed every year: as many as 1846 in 1970, as few as 16 in 1981; from 1967 to 1997, some 14 629 in total (whyte, van aarde & pimm 2003). this programme led to improvements in census and tracking techniques, in the operation of darting and drugs and to some elephant population and behavioural studies. the by-products of the elephants that were culled were not wasted. the carcasses were transported to an abattoir near (but not in sight of) the skukuza tourist camp in the kruger park, which processed, canned and dried thousands of culled elephant, as well as buffalo and hippopotamus each year (de vos, bengis & coetzee 1983; meiring 1976:102). many of the products were used as rations for national park employees and mine workers on the mines nearby. the ivory was sold, until the trade became illegal in 1989. in the 1990s, east african conservationist, richard leakey, was shown this abattoir and exclaimed, ‘i watched for a while impressed by the size and scale of the operation, but appalled that this was what wildlife “management” in the late twentieth century had come to’ (leakey 2001:220–221). in the last elephant: an african quest, journalist j. gavron was similarly outraged by sustainable utilisation. he compared the kruger park unfavourably with the parks of east africa, describing the kruger park, with its high fences and good roads as too manicured, run on accounting principles and mass harvesting for profit, rather than conserving a representative african ecosystem (gavron 1993:132–134). local south african writers on the kruger national park approached their subject from another perspective. one of the most prolific was journalist piet meiring, author of behind the scenes in kruger park (1982) and kruger park saga (1976), who also had family ties to the kruger park and people in government. using quotations from the then warden of the park, meiring extolled the kruger national park as an example of ‘untouched nature’, ‘nature’s paradise’, a place that brings the ‘public closer to nature’ in a ‘blessed atmosphere’ (meiring 1982:68–69). some 10 pages later, meiring seems oblivious to the irony of his support of ‘the principle of culling and control’ to maintain the ‘ correct population balance’ in this apparent paradisiacal landscape (meiring 1982:77). however, he was firm in his view that ‘without such control measures the game reserve would soon degenerate into a desolate wilderness’ (meiring 1982:86). this is language reminiscent of nash’s wilderness and the american mind: ‘if paradise was early man’s greatest good, wilderness, as its antipode, was his greatest evil’ (nash 1982:9). this type of discourse makes for fascinating analysis in terms of the concepts and linguistics connected to the ideology of ‘wilderness’ that people who visited the park sought. meiring’s journalistic style was mirrored by the vocabulary of science and certainty. at a conference on managing mammals in african conservation areas held in 1982, kruger’s warden defended the command-and-control model, warning that non-intervention, as was generally the case in east africa, was ‘impractical, fraught with danger, inherently untidy and can have unpredictable and even shocking consequences’ (pienaar 1983:23). he was, presumably, referring to what had happened to tsavo national park, when elephant numbers went unchecked and plummeted, with devastating results, during a serious drought. he also defended taking manipulative action: because southern africa was no longer ‘wild’, conservation areas were relatively small, fenced, and directly influenced by human activity. stability and maintenance of the climax ecosystem was, he believed, the key to maintaining a semblance of ‘the natural’. moreover, the economic alternative was ethical in his perspective, because it integrated national parks within the economy and ensured that culling was not wasteful of the lives that had been taken (pienaar 1983). as adams and mcshane (1992:100) assert, scientists often have the passion of writers of ‘literature’ and this is evident in pienaar’s presentation. by contrast – at the same conference – a paper given by the veterinarians who were actually responsible for conducting the culling was chilling because it was so unemotional: the humane killing methods were described with detachment, the carcasses clinically ‘removed’ and, finally, the slaughterhouse activities outlined in technical detail (de vos et al. 1983). one cannot help but recall cummings’s disturbing detachment in his description of himself putting bullet after bullet into an elephant. it would be tempting – but outside the scope of this paper – to investigate the connection between the culling discourse and apartheid’s discourse of this period in south african history. the connection has, however, been made. in a newspaper article in march 2008, which contested the proposed re-introduction of elephant culling in south africa, a spokesperson for animal rights africa was quoted, the formulation of the government’s policy on elephant management, specifically with regard to culling, has been driven by the rampant chauvinistic mindset of utilisation that is deeply rooted in colonialism and apartheid, which disregards the inherent value of each individual elephant and commodifies them into unfeeling units purely to be assessed for their recreational and economic value. (momberg 2008) writing in the new century (back to top) in 2001, martin meredith published elephant destiny: biography of an endangered species in africa. africa is a large and diverse continent, about which generalising is difficult, usually impossible. certainly, in many parts of africa, elephants are endangered, but this is not in southern africa. this region has the largest number of elephants in africa, accounting for 39% of the total range area. by comparison, central africa has 29%, east africa 26% and west africa 5% (blanc et al. 2007). in 1900, elephant numbers in southern africa were close to zero. currently, it is estimated that there are more than 300 000 in the region: nearly 100 000 in zimbabwe, around 130 000 in botswana, between 14 000 and 26 000 in mozambique, 12 000 to 19 000 in namibia and around 18 000 in south africa (scholes & mennell 2008:3). this growth has occurred because of a combination of the proclamation and fencing of national parks that contained elephants where natural population growth has taken place. in addition, national parks and other protected areas (private, as well as state-owned) have been established or expanded and these have been stocked with ‘excess’ elephants from elsewhere. trans-border immigration of elephants has also occurred in the region. there is debate about whether the number of elephant in southern africa is truly ‘too large’ and whether the numbers impact detrimentally on the vegetation and habitat (kerley & landman 2006). the alternative view is that, having been hunted to near-extinction in certain localities for their ivory, populations are still recovering to previous levels and that vegetation currently considered ‘natural’ (particularly large trees along major river systems) may only have matured since elephant densities were reduced (gillson & lindsay 2003; owen-smith 1988; sharpe et al. 2004). the burgeoning numbers of elephant have become a matter of local and international concern. in 1994, elephant culling ceased in south africa’s national parks for three main reasons. at the time of the transition to democracy in south africa that year and with an appreciation for the new government’s limited financial resources, there was a need to find external sources of funding. together with a growing revulsion against culling from the animal rights movement, a moratorium on culling was put in place to encourage international donors. secondly, there was growing scientific evidence that culling might not be the most appropriate tool by which to manage elephant populations in a dynamic system (page, slotow & van aarde 2006; whyte 2004; whyte & grobler 1998; whyte et al. 2003). thirdly, a new paradigm of non-equilibrium complex ecosystem dynamics (with its practical application in adaptive management, rather than command-and-control) was being advocated in international conservation biology circles (biggs & rogers 2003; rogers 2003). since culling ended, elephant numbers have doubled in the kruger national park and the changes they have wrought to the landscape is of concern to tourists and managers alike (sanparks 2005). biodiversity may be compromised by elephant actions and tourists have complained that the park is aesthetically less attractive to them (whyte 2001; whyte, biggs, gaylard & braack 1999). the government of south africa has been reluctant to re-introduce culling, particularly as the country was the host of the world summit on sustainable development (2002) and the world parks congress (2003). moreover, new conservation legislation has been under consideration during this time, viz. the biodiversity act no. 10 of 2004 and protected areas act no. 57 of 2003, and the matter has not been finally resolved, even with the proclamation of the minister of environmental affairs and tourism of the ‘national norms and standards for the management of elephants in south africa’ (republic of south africa 2008). conclusion (back to top) active public participation and interdisciplinary debate about managing and hunting elephants has been in the forefront of south africa’s animal rights debate. to some extent, the rhetoric of animal rights has become more strident, scientists have become less arrogant, perhaps even more hesitant in explaining the limits of their knowledge, while rural communities have demanded a voice in the matter in order to ensure that if culling occurs, they will benefit financially. in 1979, gray observed that no southern african literary work about hunting and adventure had received its reverse image (gray 1979:111). despitethe plethora of illustrated guidebooks, game ranger reminiscences and dense scientific journal articles on southern african mammals, this still applies. nature writing (which may be defined as non-fiction prose dealing with the natural environment and which relies on scientific factual information) is an extensive and respected genre in europe, australia and the united states of america, dating back to gilbert white, charles darwin and henry thoreau. twentieth-century masters of the genre include aldo leopold, rachel carson, wendell berry, barry lopez and stephen jay gould – some of whom are scientists, others literary and humanities scholars (finch & elder 1990; lyon 1994). apparently, however, this genre is not attractive to the south african reading public. by contrast, one might point to the extremely rich south african literature that race relations and race politics has spawned, of which the extraordinary novels of j.m. coetzee and nadine gordimer have become iconic. ours has been described as the ‘age of the anthropocene’ (crutzen 2002), because human actions have long-term and permanent effects on the planet. how humanity treats its elephants could, from a literary point of view, become a metaphor for how humanity regards itself as a species. descriptions of how elephants – an intelligent animal that shares many characteristics with humans – have over-populated the areas to which they have been confined and the extent to which this has resulted in the destruction of their habitat, the reduction in biodiversity and ‘natural’ behaviour, may be reflective of growing human populations and their impact on the environment at this time, in addition to the effects of climate change. as alexander pope expressed, in an essay on man in 1734, ‘the proper study of mankind is man, but when one regards the elephant, one wonders’ (pope, cited in watson 2003:60). literature certainly has a role to play in identifying how the elephant fits into the various human and cultural worlds, not only into the ecological. this is a matter that one author has recently confronted. in 2003, lyall watson – a biologist with an interest in the supernatural – wrote elephantoms: tracking the elephant. it is an innovative exploration of elephants in south africa, autobiographically based and recounting anecdote and adventure. watson’s novel approach lies in the fact that he combined science, imagination and history in recording his personal journey into the discovery of elephants. he avoids becoming embroiled in the culling debate, by situating his story along the cape coast, a specific location in which culling is not an issue because elephants are merely too few – phantoms in fact. the book is worth noting, for watson suggests that emotion, history, experience, science and all the other ways of knowing, are an integral part of the human-elephant encounter (watson 2003). watson touches upon ideas which biologists now study, including elephant cognition, consciousness and behaviour. moreover, historical sources, like some of those discussed above, can have more practical scientific value too, in terms of rich descriptive evidence of vegetation and the occurrence of certain species and their abundance in certain localities. such benchmarks would – as with all historical sources – have to be mediated through the prism of evidential criticism, but they may nonetheless assist management decisions in understanding elephant population dynamics and pre-settlement ranges. in the above analysis, it is hoped that some of the plethora of voices, and also the variety of discourses that elephants have generated, have been conveyed. these have ranged from the hostile to the sympathetic and from the economic to the ethical, encompassing a gamut of human responses over the past 150 years, straddling the romantic, the reverent, careful research and a discussion about rights. acknowledgements (back to top) this article is based on a paper presented at a workshop on ‘literary studies and environmental studies in africa’ convened at kansas university, lawrence, kansas, in march 2008. i would like thank the organisers, garth a. myers and byron caminero-santangelo, for inviting me to attend, as well as participants at that workshop for their helpful comments and lively discussion. i am also grateful for the substantive and editorial improvements suggested by the two anonymous referees of koedoe. i also express my appreciation to dr bob scholes and kathleen mennell, editors of scientific assessment scientific assessment of elephant management in south africa, and everyone involved in that enjoyable project, in the course of which my knowledge of elephant issues increased immensely. i also wish to express my appreciation to the university of south africa and the south african national research council for financial support. editor: llewellyn c. foxcroft references (back to top) adams, j.s. & mcshane, t.o., 1992, the myth of wild africa: conservation without illusion, university of california press, berkeley. alpers, e., 1992, ‘the ivory trade in africa: an historical review’ in d. ross (ed.), elephant: the animal and its ivory in african culture, pp. 349−360, university of california press, los angeles. bates, l.a., lee, p.c., njiraini, n., poole, j.h., sayialel, k., sayialel, s. et al., 2008,’ do elephants show empathy?’, journal of consciousness studies 15, 204–225. bates, l.a., poole, j.h. & byrne, r.w., 2008, ‘elephant cognition’ current biology 18, r544−r546. beinart, w. & hughes, l., 2007, environment and empire, oxford university press, oxford. biggs, h. & rogers, k., 2003, ‘an adaptive system to link science, monitoring and management in practice’ in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59−80, island press, washington. blanc, j.j., barnes, r.f.w., craig, g.c., dublin, h.t., thouless, c.r., douglas-hamilton, i. et al., 2007, african elephant status report, occasional paper of the iucn species survival commission no. 33. boshoff, a., skead, j. & kerley, g., 2002, ‘elephants in the broader eastern cape – an historical overview’ in g. kerley, s. wilson & a. massey (eds.), elephant conservation and management in the eastern cape, workshop proceedings, university of port elizabeth, 5 february, 2002, pp. 3−15. terrestrial ecology research unit. report no. 35. carruthers, e.j., 1995a, game protection in the transvaal, 1846 to 1926, government printer, pretoria. carruthers, j., 1995b, the kruger national park: a social and political history, natal university press, pietermaritzburg. carruthers, j., 2005, ‘changing perspectives on wildlife in southern africa, c1840 to c1914’, society and animals 13(3), 183−199. carruthers, j., 2007a, ‘conservation and wildlife management in south african national parks 1930s−1960’, journal of the history of biology 41, 203−236. carruthers, j., 2007b, ‘influences on wildlife management and conservation biology in south africa c.1900–c.1940’ south african historical journal 58, 65−90. carruthers, j., boshoff, a., slotow, r., biggs, h., avery g. & matthews, w., 2008, ‘the elephant in south africa: history and distribution’ in r.j. scholes & k.g. mennell (eds.), scientific assessment of elephant management in south africa, pp. 23–83, wits university press, johannesburg. carruthers, j., 2009, ‘”full of rubberneck wagons and tourists”: the development of tourism in south africa’s national parks and protected areas’, in w. frost & c.m. hall (eds.), tourism and national parks: international perspectives on development, history and change, pp. 211−224, routledge, london. connor, s., 2008, ‘final solution to just too many jumbos’, the sunday independent, 2 march, 2008, p. 14. crutzen, p., 2002, ‘geology of mankind’ nature 415, 23. cumming, r.g., n.d., a hunter’s life in south africa, galago, johannesburg. (first published 1850.) curtin, p., feierman, s., thompson, l. & vansina, j. 1995, african history: from earliest times to independence, 2nd edn., longman, london. de vos v., bengis r.c. & coetzee h.j., 1983, ‘population control of large mammals in the kruger national park’, in r.n. owen-smith (ed.), management of large mammals in african conservation areas. proceedings of a symposium held in pretoria, south africa, 29–30 april 1982, pp. 213−231, haum, pretoria. delegorgue, a., 1997, travels in southern africa, vol. 2, killie campbell africana library & university of natal press, pietermaritzburg. douglas-hamilton, i., bhalla, s., wittemyer, g. & vollrath, f., 2006, ‘behavioural reactions of elephants towards a dying and deceased matriarch’ applied animal behaviour science 100(1–2), 87–102. du toit, j.t., rogers, k.h. & biggs, h.c. (eds.), 2003, the kruger experience: ecology and management of savanna heterogeneity, island press, washington d.c. finch, r. & elder, j. (eds.), 1990, the norton book of nature writing, norton, new york. fitzsimons, f.w., 1920, ‘the african elephants’, in f.w. fitzsimons (ed.), mammals. the natural history of south africa, vol. 3, pp. 242−276, longmans green, london. gavron, j., 1993, the last elephant: an african quest, flamingo, london. gillson, l. & lindsay, k., 2003, ‘ivory and ecology-changing perspectives on elephant management and the international trade in ivory’ environmental science and policy 6, 411−419. gordon, a.a. & gordon, d.l., 1996, understanding contemporary africa, lynne reinner, boulder. gray, s., 1979, southern african literature: an introduction, david philip, cape town. guelke, l., 1989, ‘freehold farmers and frontier settlers, 1657−1780’, in r. elphick. & h. giliomee (eds.), the shaping of south african society, 1652-1840, pp. 66−108, maskew miller longman, cape town. hall-martin, a.j., 1992, ‘distribution and status of the african elephant loxodonta africana in south africa, 1652−1992’, koedoe 35, 65−88. hammond, d. & jablow, a., 1970, the africa that never was: four centuries of british writing about africa, twayne, new york. harris, w.c., 1840, portraits of the game and wild animals of southern africa, n.p., london. harris, w.c., 1852, the wild sports of southern africa, henry bohm, london. hoffman, m.t., 1993, ‘major p.j. pretorius and the decimation of the addo elephant herd in 1919−1920: important reassessments’, koedoe 36(2), 23−44. kerley, g.i.h. & landman, m., 2006, ‘the impacts of elephants on biodiversity in the eastern cape subtropical thickets’, south african journal of science 102, 395−402. kruger, p., 1902, the memoirs of paul kruger, 2 vols., t fisher unwin, london. leakey, r.e., 2001, wildlife wars: my fight to save africa’s natural resources, st martin’s press, new york. lubke, r., 1996, ‘thicket biome’, in a.b. low & a.g. rebelo (eds.), vegetation of south africa, lesotho and swaziland, pp. 14−18, department of environmental affairs and tourism, pretoria. luxmoore, r., 1991, ‘the ivory trade’, in s.k. eltringham (consultant), the illustrated encyclopedia of elephants, pp. 148−157, salamander, london. lye, w.f. (ed.), 1975, andrew smith’s journal of his expedition into the interior of south africa, 1834–1836, balkema, cape town. lyon, t.j. (ed.), 1989, this incomperable lande: a book of american nature writing, houghton mifflin, boston. mackenzie, j.m., 1988, the empire of nature: hunting, conservation and british imperialism, manchester university press, manchester. mccomb, k., baker, l., & moss, c., 2006, ‘african elephants show high levels of interest in the skulls and ivory of their own species’ biological letters 2, 26–28. mccomb, k., moss, c., durant, s.m., baker, l. & sayialel, s., 2001, ‘matriarchs as repositories of social knowledge in african elephants’, science 292, 491–494. meiring, p., 1976, kruger park saga, n.p., s.l. meiring, p., 1982, behind the scenes in kruger park, perskor, johannesburg. meredith, m., 2001, elephant destiny: biography of an endangered species in africa, public affairs, new york. momberg, e., 2008, ‘outrage greets lifting of culling ban’, the sunday independent, 2 march, 2008, p. 4. murray, j.a. (ed.), 1993, wild africa: three centuries of nature writing from africa, oxford university press, new york. nash, r., 1982, wilderness and the american mind, 3rd edn., yale university press, new haven. nash, r., 1989, the rights of nature: a history of environmental ethics, university of wisconsin press, madison. oliver, r. & atmore, a., 1967, africa since 1800, cambridge university press, cambridge. owen-smith, r.n., 1988, megaherbivores – the influence of very large body size on ecology, cambridge university press, cambridge. page, b., slotow, r. & van aarde, r., 2006, ‘a scientific perspective on the management of elephants in the kruger national park and elsewhere’, south african journal of science 102, 389−394. parker, i. & amin, m., 1983, ivory crisis, chatto and windus, london. parker, i., 2004, what i tell you three times is true: conservation, ivory, history and politics, librario, moray. pienaar, u. de v., 1983, ‘management by intervention: the pragmatic/economic option’, in n. owen-smith (ed.), management of large mammals in african conservation areas. proceedings of a symposium held in pretoria, south africa, 29-30 april 1982, pp. 23−36, haum, pretoria. pollock, n.c. & agnew, s., 1963, an historical geography of south africa, longmans, london. pratt, m.l., 1992, imperial eyes: travel writing and transculturation, routledge, london. pretorius, p.j. 1948, jungle man: the autobiography of major p.j. pretorius c.m.g. d.s.o. and bar, australasian publishing co., sydney. republic of south africa, 2008, department of environment and tourism, draft national norms and standards for the management of elephants in south africa, government gazette 29674, 2 march, under general notice no. 224. ritvo, h., 1987, the animal estate: the english and other creatures of the victorian age, harvard university press, cambridge. ritvo, h., 1997, the platypus and the mermaid and other figments of the classifying imagination, harvard university press, cambridge. roche, c., 1996, ‘the elephants at knysna’ and ‘the knysna elephants’. from exploitation to conservation: man and elephants at knysna 1856–1920, ba (hons.) thesis, university of cape town. rogers, k.h., 2003, ‘adopting a heterogeneity paradigm: implications for management of protected savannas’, in, j.t. du toit, k.h. rogers & h.c. biggs, h.c. (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 41−58, island press, washington d.c. ross, d.h. (ed.), 1992, elephant: the animal and its ivory in african culture, fowler museum of cultural history, university of california, los angeles. ross, r., 1989, ‘the cape and the world economy, 1652−1835’, in r. elphick & h. giliomee (eds.), the shaping of south african society, 1652–1840, pp. 243−280, maskew miller longman, cape town. scholes, r.j. &. mennell, k.h. (eds.), 2008, scientific assessment of elephant management in south africa, wits university press, johannesburg. sharpe, c., aarrestad, p.a., andreassen, h.p., dhillion, s.s., dimakatso, t., du toit, j.t., et al., 2004, ‘the return of the giants: ecological effects of an increasing elephant population’, ambio 33, 276−282. skead, c.j., 1980, historical mammal incidence in the cape province: vol.1 – the western and northern cape, department of nature and environmental conservation, provincial administration of the cape of good hope, cape town. skead, c.j., 2007, in a.f. boshoff, g.i.h. kerley & p.h. lloyd (eds.), historical incidence of the larger land mammals in the broader eastern cape, 2nd edn., centre for african conservation ecology, nelson mandela metropolitan university, port elizabeth. south african national parks, 2005, the great elephant indaba: finding an african solution to an african problem, south african national parks, pretoria. thomas, k., 1984, man and the natural world: changing attitudes in england, 1500–1800, penguin, harmondsworth. thorbahn, p.f., 1979, ‘the precolonial ivory trade of east africa: reconstruction of a human-elephant ecosystem’, phd thesis, department of anthropology, university of massachusetts, amherst. van der merwe, p.j., 1938, die trekboer in die geskiedenis van die kaapkolonie [the migrant farmer in the history of the cape colony], nasionale pers, cape town. van der merwe, p.j., 1945, trek, nasionale pers, cape town. van wyk, p. & fairall, n., 1969, ‘the influence of the african elephant on the vegetation of the kruger national park’ koedoe 12, 57−89. watson, l., 2003, elephantoms: tracking the elephant, penguin, johannesburg. wemmer, c. & christen, c. (eds.), 2008, elephants and ethics: towards a morality of coexistence, johns hopkins press, baltimore. whitehouse, a.m. & hall-martin, a.j., 2000, ‘elephants in the addo national elephant park, south africa: reconstruction of the population’s history’, oryx 34, 46−55. whyte i.j., 2004, ‘ecological basis of the new elephant management policy for kruger national park and expected outcomes’, pachyderm 36, 99−108. whyte, i.j. & grobler, d.g., 1998, ‘elephant contraception in the kruger national park’, pachyderm 25, 45−52. whyte, i.j., 2001, conservation management of the kruger national park elephant population, phd thesis, department of wildlife management, university of pretoria. whyte, i.j., biggs, h.c., gaylard, a. & braack, l.e.o., 1999, ‘a new policy for the management of the kruger national park’s elephant population’, koedoe 42, 111−131. whyte, i.j., van aarde, r. & pimm, s.l. 2003, ‘kruger’s elephant population: its size and consequences for ecosystem heterogeneity’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 332−348, island press, washington d.c. wickens, p.l., 1981, an economic history of africa from the earliest times to partition, oxford university press, cape town. wylie, d., 2008, elephant, reaktion books, london. filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 terblanche 2 life history.qxd the application of life history information to the conservation management of chrysoritis butterflies (lepidoptera: lycaenidae) in south africa r.f. terblanche and h. van hamburg terblanche, r.f. and h. van hamburg. 2004. the application of life history information to the conservation management of chrysoritis butterflies (lepidoptera: lycaenidae) in south africa. koedoe 47(1): 55–65. pretoria. issn 0075-6458. due to their intricate life histories and the unique wing patterns and colouring the butterflies of the genus chrysoritis are of significant conservation and aesthetic value. this overview probes into practical examples of butterfly life history research applicable to environmental management of this relatively well-known invertebrate group in south africa. despite the pioneer work on life histories of chrysoritis in the past, more should be done to understand the life history of the butterflies in the wild, especially their natural host plants and the behaviour of adults and larvae. a system of voucher specimens of host plants should be introduced in south africa. although various host plant species in nature are used by the members of chrysoritis, including the chrysoritis chrysaor group, the choice of these in nature by each species is significant for conservation management and in the case of chrysoritis aureus perhaps even as a specific characteristic. a revision of the ant genus crematogaster will benefit the conservation management of chrysoritis species since some of these ant species may consist of a number of species with much more restricted distributions than previously thought. rigorous quantified studies of population dynamics of chrysoritis butterflies are absent and the introduction of such studies will benefit conservation management of these localised butterflies extensively. key words: chrysoritis, myrmecophilous, endemic, conservation, life histories, management, taxonomy, ecology, synecology. r.f. terblanche and h. van hamburg, school of environmental sciences and development, private bag x6001, potchefstroom university for che, potchefstroom, 2520 south africa. issn 0075-6458 55 koedoe 47/1 (2004) introduction williams (1996) reported a lack of information on the life histories of south african butterflies despite the most valuable efforts of the past. pioneer lepidopterists (including clark, claassens, dickson, cottrell, g.a. henning, s.f. henning and heath) provided a rich source of life history information in the past, useful for environmental management. further research priorities are identified here. this paper reviews the life history information on the chrysoritis genus to date, and discusses how applicable this information is for conservation management. world-wide, the symbioses of butterflies and ants are found only among members of the riodinidae and lycaenidae (de vries 1997) and at present the riodinidae is often regarded as a subfamily riodininae within the lycaenidae (eliot 1973; eliot 1990; pierce et al. 2002). although the riodininae is poorly represented in africa, the rest of the lycaenidae is well-represented in most habitat types on the continent. judging by the number of species, the majority of south african butterflies belong to the family lycaenidae. from our list of 674 butterfly species, i.e. superfamilies hesperioidea: ‘skipper butterflies’ and papilionoidea: ‘true terblanche 2 life history.qxd 2004/04/08 02:19 page 55 butterflies’ in south africa, 49 % of the species belong to the family lycaenidae. the majority of these lycaenidae species are associated with ants. a diversity of strategies has developed into ant associations among the lycaenidae. extremes, from aphytophagy (cottrell 1984) to facultative ant associations, have been recorded. the term entomophagous is used by pierce et al. (2002) instead of aphytophagy, which alter the normal use of entomophagous as ‘insecteating’ to ‘eating insect-derived food sources’. apart from the term entomophagous, all the other definitions to describe lycaenid-ant associations as defined by pierce et al. are followed here. note that pierce et al. (2002) use the terms obligate and facultative ant associations in terms of survival under field conditions, which is strictly followed here. given the nature and complexity of larvae ant symbioses, there is much to be learnt from their studies that will be relevant to all branches of biology (de vries 1997). the various types of associations with ants (henning 1983a; cottrell 1984; pierce et al. 2002) by lycaenidae butterflies, should be appreciated in that different conservation strategies should probably be applied to conserve these, often localised, butterflies. discussion detailed life history descriptions that entail most stages and some illustrations have only been published for 35.7 % of all the chrysoritis species in south africa (table 1). it should be added that ecophysiological studies of interactions between butterfly larvae and host plants as well as attendant ants are absent, not only for chrysoritis but for almost all the butterflies of south africa. at least one host plant and one attendant ant are known, respectively, for 83.3 % and 88.1 % of the chrysoritis species. these relatively high numbers are due to the contribution of heath (1997a) (table 1). dickson (1940, 1943, 1944, 1945a, 1945b, 1946, 1947, 1948, 1953, 1959 and 1965) gave either detailed and well-illustrated (by clark) descriptions of the life histories of some chrysoritis species or information on the host plants and attendant ants. the first major work on the life histories of lycaenidae butterflies of south africa was that of clark & dickson (1971) that contains a number of detailed life history descriptions. a summary of recent observations of ant associations and life history adaptations of lycaenidae butterflies in south africa is given by heath & claassens (2000). heath (1997) for the first time reported the extensive use of thesium plants (santalaceae) as host plants by chrysoritis species. the importance of the presence of the correct ant as an oviposition stimulus for most chrysoritis species, at least in captivity, is reported (heath & claassens 2000). clark & dickson (1971) concluded that chrysoritis dicksoni is aphytophagous, since larvae refused to feed on any plant that females had laid on. the mature larvae and pupa were found in crematogaster ant nests (cotrell 1984). heath & brinkman (1995), heath (1998) and heath & claassens (2000) demonstrated that at least in some larval stages, the larvae of chrysoritis dicksoni are sustained by trophallaxis. among the chrysoritis genus trophallaxis (ants feeding caterpillars by regurgitation) has thus been reported only for chrysoritis dicksoni by heath (1998). the majority of the chrysoritis species, with the possible exception of chrysoritis dicksoni, seems to be phytophagous accompanied by an obligate ant association (tables 14). no voucher material of host plants has been cited in all the above cases. although the genera and even some of the plant species identified leave little doubt about its identification, the lack of voucher specimens may be a limitation to future research efforts. cottrell (1984) highlighted the problems associated with the fragmented observations of life histories of aphytophagous butterflies. these are also applicable to the ant-dependent phytophagous chrysoritis species. thomas et al. (1989) summarised the difficulties with the accuracy of interpretations of captive observations, the misinterpretation of field data, as well as the misidentification koedoe 47/1 (2004) 56 issn 0075-6458 terblanche 2 life history.qxd 2004/04/08 02:19 page 56 issn 0075-6458 57 koedoe 47/1 (2004) of ant species concerning research on maculinea butterflies and their ant associations in europe. the question is whether quantitative laboratory-controlled observations and careful observations in the field concerning the life histories of chrysoritis species are available or not. most chrysoritis species were bred in captivity (where behaviour and requirements are abnormal). therefore little information exists under controlled conditions or in the field. we discuss the present knowledge of the host plant association and attendant ant association separately for simplicity. the host plant association a summary of the host plant genera, host plant families, and attendant ant genera associated with the species groups of chrysoritis is given in table 2. chrysoritis species have adapted to use a variety of host plants from different genera and families as larval food (table 2). different species groups share some genera and families. the ability of some species to use succulents of the family crassulaceae as host plants gives rise to the hypothesis that such host plants may have been important for the survival of these butterflies especially during dry spells. the full significance of the use of host plants will only become apparent if more research on their use in specific ecosystems were conducted. another research priority is that some host plants for the chrysoritis species need confirmation. chrysoritis chrysaor was observed to oviposit on acacia karroo (heath 1997), but it is not clear whether the larvae use these trees as a host plant. during research on chrysoritis aureus from 1998 to 2002, it was found that the females often oviposit on rocks close to the host plant and in fact close to the attendant ant trails (roos & henning 2000). the proof for the use of a specific host plant remains the larvae eating the host plant in its habitat. the type of observation (egg laying, larva at base of host plant) about the host plant should be stated clearly in publications. the significance of the host plants on a taxonomic and ecological level is discussed by using chrysoritis aureus as an example. heath (2001) demonstrated that the choice of larval host plant is not a major issue within a species group of chrysoritis and should not be regarded as a specific characteristic. the variety of host plants used by chrysoritis, even in the same species group such as the chrysoritis chrysaor species group seems to support this hypothesis. pierce (1984) proposed two ways by which ant association could have enhanced diversification of the lycaenidae, once it evolved. the first is by inducing a higher incidence of host plant switching and the second is by modifying the butterfly population structure. an example in chrysoritis is the observation that chrysoritis aethon larva used a crassula species as host plant, and not only the known host plant rhus zeyheri (anacardiaceae), that was reported by owen-johnston (1991). this phenomenon should be further investigated since it might be ecologically significant. heath (2001) states that chrysoritis aureus has been successfully bred in captivity on another plant, diospyros lycioides, an important observation in the context of the chrysoritis chrysaor species group to which chrysoritis aureus belongs. up to date, clutia pulchella individuals were observed as host plants of chrysoritis aureus larvae in the field. only once (17 march 2004) was diospyros lyciodes observed to be a host plant, despite its presence at many of our research sample plots. clutia pulchella has a wide distribution in the northern provinces of south africa (retief & herman 1997) as well as south africa (arnold & de wet 1993) but is not used by any chrysoritis species other than chrysoritis aureus. the only euphorbiaceae plant known to be utilised by any chrysoritis species is clutia pulchella. it is utilised by larvae of chrysoritis aureus (tables 2 and 3). the unidentified morgenzon chrysoritis entity, as well as chrysoritis lyncurium (closely related to chrysoritis aureus within the chrysoritis chrysaor species group), is also thought to use diospyros as a host plant (owen-johnston 1991). this has been assumed from the adult behaviour and has terblanche 2 life history.qxd 2004/04/08 02:19 page 57 koedoe 47/1 (2004) 58 issn 0075-6458 table 1 a summary of the life-history studies published on chrysoritis. the types of descriptions (a,b,c) include: a a description of all or some of the stages (from egg to adult), b the host plant has been reported, c the attendant ant has been reported. the literature sources are those that reported previously unknown aspects of the life-histories. the literature sources used were, dickson (1940, 1943, 1944, 1945a, 1945b, 1946, 1947, 1948, 1953, 1959, 1965), clark and dickson (1971), dickson and kroon (1978), s.f. henning (1983a), owen-johnston (1991), heath and brinkman (1995), heath (1997a), heath (1998), claassens (2000) and heath (2001) chrysoritis species description literature a b c chrysaor species group chrysoritis aethon 1 1 owen-johnston (1991) (trimen & bowker, 1887) chrysoritis aureus (van son, 1966) 1 1 1 henning (1983a) chrysoritis chrysaor (trimen, 1864) 1 1 1 dickson (1943), dickson (1944), heath(1997a), claassens (2000) chrysoritis lycegenes (trimen, 1874) 1 1 1 clark & dickson (1971), henning (1983a) chrysoritis lyncurium (trimen, 1868) 1 heath (1997a) chrysoritis midas (pennington, 1962) 1 1 heath (1997a) chrysoritis natalensis (van son, 1966) 1 dickson & kroon (1978) chrysoritis phosphor (trimen, 1866) __ __ __ chrysantas species group chrysoritis chrysantas (trimen, 1868) __ __ __ oreas species group chrysoritis dicksoni (gabriel, 1946) 1 1 dickson (1953), dickson (1965), clark & dickson (1971), heath & brinkman (1995), heath (1997a), heath (1998) chrysoritis oreas (trimen, 1891) 1 1 zeuxo species group chrysoritis zeuxo (linnaeus, 1764) 1 1 1 dickson (1953), dickson (1975), clark & dickson (1971), heath (1997a) chrysoritis zonarius (riley, 1938) 1 1 1 heath (1997a), claassens (2000) pyroeis species group chrysoritis felthami (trimen, 1904) 1 1 1 dickson (1940), clark & dickson (1971), heath (1997a), claassens (2000) chrysoritis pyroeis (trimen, 1864) 1 1 1 *dickson (1948), clark & dickson (1971), heath (1997a), claassens (2000) thysbe species group chrysoritis adonis (pennington, 1962) 1 1 1 dickson (1965), clark & dickson (1971), heath (1997a) chrysoritis aridus (pennington, 1953) 1 1 dickson (1965), clark & dickson (1971) chrysoritis azurius (swanepoel, 1975) __ __ __ chrysoritis beaufortius (dickson, 1966) 1 1 heath (1997a) chrysoritis beulah (quickelberge, 1966) __ __ __ chrysoritis blencathrae 1 1 heath (1997a) (heath and ball, 1992) chrysoritis braueri (pennington, 1967) 1 1 1 clark & dickson (1971), heath (1997a) chrysoritis brooksi (riley, 1938) 1 1 dickson (1959), clark & dickson (1971), heath (1997a) chrysoritis daphne (dickson, 1975) 1 1 heath (1997a) chrysoritis endymion (pennington, 1962) 1 1 heath (1997a) chrysoritis irene (pennington, 1968) __ __ __ chrysoritis nigricans (aurivillius, 1924) 1 1 1 dickson (1944), dickson (1947), clark & dickson (1971), heath (1997a), claassens (2000) chrysoritis orientalis (swanepoel, 1976) 1 1 heath (1997a) chrysoritis palmus (stoll, 1781) 1 1 1 dickson (1945a,1945b), dickson (1953), dickson (1965), clark & dickson (1971), claassens (2000) chrysoritis pan (pennington, 1962) 1 1 1 **dickson (1965), **clark & dickson (1971), heath (1997a) chrysoritis pelion (pennington, 1953) __ __ __ chrysoritis penningtoni (riley, 1938) 1 heath (1997a) chrysoritis perseus (henning, 1977) 1 1 heath (1997a) terblanche 2 life history.qxd 2004/04/08 02:19 page 58 not been verified. an alternative explanation is that diospyros lycioides is an ancestral host plant for chrysoritis lyncurium, chrysoritis lycegenes, the morgenzon chrysoritis entity, as well as chrysoritis aureus and chrysoritis aethon—all belonging to the chrysoritis chrysaor species group. in general, it is common practice for lepidopterists to use substitute host plants for breeding butterflies, plants that may not necessarily be used in the wild. although various host plant species are used by the members of chrysoritis, including the chrysoritis chrysaor group, the choice of these, in nature, by each species is significant for conservation management, in the case of chrysoritis aureus perhaps even as a specific characteristic. the proper management of clutia pulchella at the habitats of chrysoritis aureus would be very important for the survival of the butterfly in its increasingly urbanised distribution. in the light of the above it is furthermore significant that chrysoritis aureus is the only chrysoritis species apart from the ubiquitous chrysoritis chrysaor that extended its distribution into the rocky highveld grassland via clutia pulchella and its attendant ant (bredenkamp & van rooyen 1996). the ant association for a number of chrysoritis species, the attendant ant is perhaps of more importance than the host plant as a signal for the female to lay eggs (heath 1997, 2001). various adaptations and strategies exist among the rich myrmecophilous lycaenid fauna of africa, of which various are described by s.f. henning (1983a) and cottrell (1984). during the day, chrysoritis larvae shelter for protection and only venture forth at night to feed on their host plants (henning 1987a). although the morphological and physiological adaptations among the myrmecophilous larvae have been described, the role of these has been more difficult to demonstrate. there are a number of accessory structures that are associated with larvae that adopted a myrmecophilous life style. most lycaenid larvae have a median dorsal organ (honey gland) (henning 1983a), which is referred to as a dorsal nectary organ (dno) by cotrell (1984). these are present in all the chrysoritis larvae studied up to date. a pair of dorsolateral eversible organs is also found on many lycaenid larvae (henning 1983a). these are referred to as tubercle organs (tos) by cottrell (1984) and are also found on all the larvae of chrysoritis studied to issn 0075-6458 59 koedoe 47/1 (2004) table 1 (continued) chrysoritis species description literature a b c chrysoritis plutus (pennington, 1976) 1 1 heath (1997a) chrysoritis pyramus (pennington, 1953) 1 1 heath (1997a) chrysoritis rileyi (dickson, 1966) 1 1 heath (1997a) chrysoritis swanepoeli (dickson, 1965) 1 1 heath (1997a) chrysoritis thysbe (linnaeus, 1764) 1 1 1 dickson (1947), dickson (1965), clark & dickson (1971), heath (1997a), claassens (2000) chrysoritis trimeni (riley, 1938) 1 1 heath (1997a) chrysoritis turneri (riley, 1938) 1 1 dickson (1953); heath (1997a) chrysoritis uranus (pennington, 1962) 1 1 1 dickson (1965), clark & dickson (1971), heath (1997a) chrysoritis violescens (dickson, 1971) 1 1 heath (1997a) total 15 35 37 percentage of total species 35.7 83.3 88.1 * see heath (1997a) that the camponotus associate that dickson reported was never repeated ** note this was observed for chrysoritis lysander (now regarded as synonym of chrysoritis pan by heath (2001). terblanche 2 life history.qxd 2004/04/08 02:19 page 59 koedoe 47/1 (2004) 60 issn 0075-6458 table 2 summary of the host plant genera, host plant families and attendant ant genera associated with the different species groups of chrysoritis. the literature sources: dickson (1940, 1943, 1944, 1945a, 1945b, 1946, 1947, 1948, 1953, 1959, 1965), clark & dickson (1971), dickson & kroon (1978), s.f. henning (1983a), owen-johnston (1991), heath & brinkman (1995), heath (1997a), heath (1998), claassens (2000) and heath (2001). plant genera and families are in alphabetic order no. of species: known host known host host ant heath (2001) plant genera plant families genera c. chrysaor sp. group 8 acacia anacardiaceae crematogaster clutia asteraceae chrysanthemoides crassulaceae cotyledon ebenaceae diospyros euphorbiaceae myrsine fabaceae rhus myrsinaceae tylecodon zygophyllaceae zygophyllum c. chrysantas sp. group 1 unknown unknown unknown c. oreas sp. group 2 thesium1 santalaceae1 crematogaster myrmicaria c. zeuxo sp. group 2 chrysanthemoides asteraceae crematogaster c. pyroeis sp. group 2 zygophyllum zygophyllaceae crematogaster thesium santalaceae myrmicaria c. thysbe sp. group 27 aspalathus apiaceae crematogaster berzelia asteraceae centella bruniaceae chrysanthemoides crassulaceae dimorphotheca fabaceae lebeckia myrsinaceae osteospermum santalaceae thesium zygophyllaceae tylecodon zygophyllum (1perhaps no plant utilised in the case of chrysoritis dicksoni) date. these organs tend to be important with regard to the ant associations as defined by pierce et al. (2002). in the case of chrysoritis aureus, the anterior part of the head of the cremagaster liengmei ant sometimes disappears into the dno in search of the nectar produced by the larvae. furthermore, the inner part of the tubercles is often eversed in the presence of the ants or to ‘attract the attention of the ants’. the above studies emphasise the extent to which these butterfly larvae are adapted for ant associations. in the case of aloeides dentatis henning (1983b) found that epidermal glands (pore copula organs) have a secretion that mimics the brood pheromone of the attendant ant. such detailed studies have not yet been published on any of the chrysoritis. audible squeaking sounds, produced by pupae of chrysoritis brooksi (schlosz & schlosz 1990) and chrysoritis irene (schlosz 1991), might be of ecological significance and should be studied further. drumming sounds from the larvae of chrysoritis dicksoni and chrysoritis thysbe were recorded by heath (1998). de vries (1990, 1992) has provided detailed descriptions and research techniques that detected audible rituals and other accessories that enhance symbiotic terblanche 2 life history.qxd 2004/04/08 02:19 page 60 associations between riodininae larvae and ants. at present, no such detailed research exists for any of the african lycaenidae. a number of instances have been reported where chrysoritis species have been bred in captivity in the absence of ants. nevertheless, in the field the association seems to be vital. the latter confirms the importance of observations in the field—associations that may not be essential for survival in the laboratory might be obligatory in the field. most of the chrysoritis species fall into the category of phytophagous, but with a myrmecophily, where the butterfly is dependent on ants (see henning 1983a). issn 0075-6458 61 koedoe 47/1 (2004) table 3 summary of the host plants used by species belonging to the c. chrysaor species group in the genus chrysoritis and the literature source where the host plant was first reported. a larvae were bred on the plant in captivity or in the laboratory. aa only oviposition on the plant observed (heath 1997). aaa larvae or pupae were observed on or at the base of the host plant in the field species plant species plant family literature source information with indication of about type of observation voucher specimens chrysoritis rhus zeyheri aaa anacardiaceae owen-johnston (1991) no voucher aethon (trimen crassula sp. aaa crassulaceae heath (2001) material sited & bowker, 1887) chrysoritis clutia pulchella aaa euphorbiaceae henning (1983) no voucher aureus diospyros lycioides a ebenaceae heath (2001) material sited (van son, 1966) chrysoritis cotyledon orbiculata l. aaa crassulaceae dickson (1943) no voucher chrysaor rhus sp. aaa anacardiaceae dickson (1943) material sited (trimen, 1864) acacia karroo aa fabaceae heath (1997a zygophyllum retrofractum aa zygophyllaceae heath (1997a) tylecodon paniculata aa crassulaceae. heath (1997a)# chrysoritis royena hirsuta aaa ebenaceae clark & dickson 1971) no voucher lycegenes myrsine africana aaa myrsinaceae clark & dickson (1971) material sited (trimen, 1874) diospyros lycioides aaa ebenaceae henning (1983a) diospyros austro-africana aaa ebenaceae henning (1983a) rhus sp. aaa anacardiaceae henning (1983a) chrysoritis not confirmed not confirmed lyncurium (trimen, 1868) chrysoritis diospyros austro-africana ebenaceae heath (1997a) no voucher midas material sited (pennington, 1962) chrysoritis chrysanthemoides monilifera asteraceae dickson & kroon (1978) no voucher natalensis cotyledon orbiculata crassulaceae dickson & kroon (1978) material sited (van son, 1966) (no reference to observations) chrysoritis unknown unknown phosphor (trimen, 1866) #note this information is for the entity chrysoritis lycia (riley, 1938), which is now regarded as a synonym of chrysoritis chrysaor by heath (2001). terblanche 2 life history.qxd 2004/04/08 02:19 page 61 the reasons for the evolution of ant associations are discussed by cottrell (1984), especially whether these associations are for protection from parasites and predators (thomann 1901) or to avoid ant aggression (lenz 1917). quantitative and controlled observations in this regard are also rare. cottrell (1984) suggests that both factors had differing and probably complementary parts to play in different taxonomic groups within the lycaenidae. evidence that the presence of ants associated with myrmecophilous butterflies reduces the risk of attack from predators and parasites are provided by pierce & mead (1981) and pierce et al. (1987). pierce et al. (2002) provides an overview of costs and benefits of lycaenid-ant interactions. specific detailed experiments of the south african fauna seem to be absent, and seems to be important for conservation since the importance of ant protection seems to vary among species (see pierce et al. 2002). despite the variety of host plants used by the chrysoritis species (tables 2 & 3), only ants of the subfamily myrmicinae are used as attendant ants (table 4). the larvae of chrysoritis aureus are constantly attended by crematogaster ants (henning 1983a). the ant genus crematogaster is a large genus of small, monomorphic, blackish or brownish-yellow ants of which at least 50 species have been described from southern africa. they occur from the cape to ethiopia (prins 1978). it is interesting to note that the chrysoritis genus is confined to the south-western and south-eastern parts of south africa, despite the ability to use various host plants and being associated with crematogaster ants, of which numerous species are widely distributed in africa. perhaps the various other aphnaeini (lycaenidae: theclinae) butterflies that are also associated with crematogaster ants might be better adapted to savanna and forest ecosystems than chrysoritis species. heath (2001) noted the importance of ant associates in the systematics of the genus chrysoritis as well as the possibility that the ant species are themselves composed of species groups that should not be overlooked. the importance of crematogaster liengmei and crematogaster peringueyi as attendant ants can be appreciated for the chrysoritis chrysaor species group (table 4). both these ant species are also attendant ants for chrysoritis species that belong to other species groups (see heath 2001). therefore, the correct crematogaster species in each case, that seems to be essential for the survival of these butterflies may be less widespread than is currently understood. the implication of heath's observation is that a revision of the crematogaster ant species will benefit the conservation management of chrysoritis species. in the absence of ants, a fungal infection occurs in the dorsal nectary organ and the larva usually dies after a few days ( henning 1987; roos & henning 2000; terblanche et al. 2003). all the larval instars of chrysoritis aureus, as well as the pupae in the ant nest were observed to be accompanied by at least two but often more crematogaster liengmei individuals. this was found every time that larvae or pupae were observed during the field work done in the wet seasons of 2000–2004 by r.f. terblanche in the heidelberg district of south africa. when an ant nest underneath a rock contained pupae, it was carefully turned over (fig. 1), the crematogaster liengmei individuals were observed to lift their gasters and some would position themselves on the dorsal part of the pupae. in these cases the ants that rarely bite when disturbed, now did so fiercely. the pupae, and not only larvae, seem to be protected, but the mechanisms of the pupa to enhance protection seem to be poorly studied. the presence of larvae and especially pupae in the ant nests could be a response to a variety of unfavourable factors, such as mammalian grazing and trampling pressures, frost and fire (cottrell 1984). although these factors make sense considering for example the field observations on chrysoritis aureus, cottrell (1984) noted that such explanations should be based on much more detailed quantitative studies of individual lycaenid biology. koedoe 47/1 (2004) 62 issn 0075-6458 terblanche 2 life history.qxd 2004/04/08 02:19 page 62 brussard (1991) and new (1997) verified the importance of detailed auto-ecological information. new (1997) noted the importance of studying butterfly population dynamics at least for a number of seasons, but also mentioned that the duration of the studies or urgency sometimes permits for one flight season only, or sometimes even less. heath & brinkman (1995) have described aspects of the population dynamics of chrysoritis dicksoni based on collections and observations in the field. no published literature could, however, be found where the size or the abundance of the colonies of chrysoritis had been measured or quantified with the aid of sample plots, transects or any well-described mark-andrecapture procedure. in fact, few population studies have been published for any of the myrmecophilous lycaenidae of southern africa. an unpublished report by de wet (1992) on the butterfly erikssonia acraeina contains numbers and sex ratios. conclusions pioneer lepidopterists (including clark, claassens, dickson, cottrell, g.a.henning, s.f. henning and heath) provided a rich source of life history information in the past which in the end is a source of information for environmental management. this paper indicates that much more should be done to understand the life history of the butterflies in the wild, especially their natural host plants and the behaviour of adults and larvae. the circumstances and types of associations with host plants and issn 0075-6458 63 koedoe 47/1 (2004) fig. 1. wall of ant nest with pupae and attendant crematogaster liengmei ants. attendant ants should be more precisely described so that traceability and repeatability could be enhanced. experiments of microclimate on the morphology of chrysoritis species should be described in detail, as well as the results, if one were to prove the relevant hypotheses. no system of voucher specimens of host plants exists in south africa. such a system would enhance the traceability of host plant information, especially if taxonomic changes regarding the plants have taken place. fish (1999) gives clear guidelines on the preparation of herbarium specimens that are recommended to study the life histories of butterflies. the physiological interactions during the life cycle of myrmecophilous butterflies with their attendant ants and host plants in south africa are unknown for chrysoritis species (and also for all the lycaenidae in southern africa). the research will therefore most probably be rewarding. much remains to be done to quantify the population dynamics of chrysoritis species and in fact all the localised butterflies in africa. although it would prove to be very difficult, efforts should also be made to quantify predation and competition, or at least make an effort to do so. the use of alternatives for mark-and-recapture methods, such as transecting methods, is a research field worth exploring in africa. a better idea of dispersal behaviour of chrysoritis butterflies would enhance the terblanche 2 life history.qxd 2004/04/08 02:19 page 63 understanding of distribution, population dynamics and especially metapopulations. references arnold, t.h. & b.c. de wet (eds.). 1993. plants of southern africa: names and distribution. memoirs of the botanical survey of south africa no. 62. bredenkamp, g.j. & n. van rooyen. 1996. rocky highveld grassland. p. 39. in: low, a.b. & a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. brussard, p.f. 1991. the role of ecology in biological conservation. ecological applications 1(1): 6–12. claassens, a.j.m. 2000. the butterflies of the cape peninsula: a comprehensive guide. cape town: tafelberg. clark, g.c. & c.g.c. dickson. 1971. life histories of the south african lycaenid butterflies. cape town: purnell. cottrell, c.b. 1984. aphytophagy in butterflies: its relationships to myrmecophily. zoological journal of the linnean society 79: 1–57. de vries, p.j. 1990. enhancement of symbiosis between butterfly caterpillars and ants by vibrational communication. science 248(4959): 1104–1106. de vries, p.j. 1992. singing caterpillars, ants and symbiosis. scientific american (oct.): 56–62. de vries, p.j. 1997. the butterflies of costa rica and their natural history. vol. ii riodinidae. nj: princeton university press. de wet, j.i. 1992. die bewaring van die skoenlapper erikssonia acraeina in die waterberg. interne verslag. (projek tk 3. eerste vorderingsverslag. hoofdirektoraat: natuuren omgewingsbewaring, invertebraatseksie, lydenburg). dickson, c.g.c. 1940. notes on the early stages of phasis felthami (trim.), a lycaenid butterfly from the cape peninsula, and a list of some recently determined foodplants of some other south african butterflies. annals of the south african museum 32 (6): 545–554. dickson, c.g.c. 1943. the life-history of phasis chrysaor (trim.) (lepidoptera: lycaenidae). journal of the entomological society of southern africa 6: 37–47. dickson, c.g.c. 1944. recently observed foodplants of some cape lepidopterous larvae (1st series). journal of the entomological society of southern africa 7: 96–99. dickson, c.g.c. 1945a. the life-history of phasis palmus (cram.) (lepidoptera: lycaenidae). journal of the entomological society of southern africa 8: 99–110. dickson, c.g.c. 1945b. recently observed foodplants of some cape lepidopterous larvae (2nd series). journal of the entomological society of southern africa 8: 150–153. dickson, c.g.c. 1946. the life-history of phasis thysbe l. var. nigricans aurivillius (lepidoptera: lycaenidae). journal of the entomological society of southern africa 9: 178–192. dickson, c.g.c. 1947. recently observed foodplants of some cape lepidopterous larvae (3rd series). journal of the entomological society of southern africa 10:126–130. dickson, c.g.c. 1948. the life-history of phasis pyroeis (trim.) (lepidoptera: lycaenidae). journal of the entomological society of southern africa 11: 99–110. dickson, c.g.c. 1953. recently observed foodplants of some cape lepidopterous larvae (4th series). journal of the entomological society of southern africa 15(1): 11–20. dickson, c.g.c. 1959. notes on the early stages of poecilmitis brooksi (riley) (lepidoptera: lycaenidae). journal of the entomological society of southern africa 22 (2): 312–315. dickson, c.g.c. 1965. recently observed foodplants of some south african lepidopterous larvae (5th series). journal of the entomological society southern africa 28(1): 11–20. dickson, c.g.c. & d.m. kroon (eds.). 1978. pennington's butterflies of southern africa. johannesburg: ad. donker. elliot, j.n. 1973. the higher classification of the lycaenidae (lepidoptera): a tentative arrangement. bulletin of the british museum of natural history (entomology) 28:371–505. elliot, j.n. 1990. notes on the genus curetis hübner (lepidoptera, lycaenidae). tyô to ga 41: 201–225. fish, l. 1999. preparing herbarium specimens. strelitzia 7: 1–57. heath, a. 1997. a review of african genera of the tribe aphnaeini (lepidoptera: lycaenidae). metamorphosis: occasional supplement 2: 1–60. heath, a. 1998. further aspects on the life history of the myrmecophilous species chrysoritis dicksoni (gabriel), (lepidoptera: lycaenidae). metamorphosis 9(4): 160–172. heath, a. 2001. new synonymies and taxonomic notes on the genus chrysoritis (butler) (lepidoptera: lycaenidae). metamorphosis 12 (3): 85–98. heath, a. & a.k. brinkman. 1995. aspects of the life history, distribution and population fluctuations of oxychaeta dicksoni (gabriel) (lepidoptera lycaenidae). metamorphosis 6(3): 117–127. koedoe 47/1 (2004) 64 issn 0075-6458 terblanche 2 life history.qxd 2004/04/08 02:19 page 64 heath, a. & a.j.m. claassens. 2000. new observations of ant associations and life history adaptations (lepidoptera: lycaenidae) in south africa. metamorphosis 11(1): 3–19. henning, s.f. 1983a. biological groups within the lycaenidae (lepidoptera). journal of the entomological society of southern africa, 46(1): 65–85. henning, s.f. 1983b. chemical communication between lycaenidae larvae (lepidoptera: lycaenidae) and ants (hymenoptera: formicidae). journal of the entomological society of southern africa 46(1): 65–85. henning, s.f. 1987. myrmecophilous lycaenidae (or how ants help butterflies). south african journal of science 83: 8–9. lenz, f. 1917. der erhaltungsgrund der myrmekophilie. zeitschrift für induktive abstammungsund vererbungslehre 18: 44–46, 71–72. low, a.b. & a.g. rebelo (eds.). 1996. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. new, t.r. 1997. butterfly conservation. melbourne: oxford university press. owen-johnston, n.k. 1991. thoughts and observations on the genus poecilmitis butler (lepidoptera: lycaenidae) in the transvaal. metamorphosis 2(3): 49–50. pierce, n.e. 1984. amplified species diversity: a case study of an australian lycaenid butterfly and its attendant ants pp. 198–200. in: ackery, p. & r. vane-wright (eds.). the biology of butterflies. london: academic press. pierce, n.e. & p.s. mead. 1981. parasitoids as selective agents in the symbiosis between lycaenid butterfly larvae and ants. science 211: 1185–1187. pierce, n.e., r.l. kitching, r.c. buckley, m.f.j. taylor & k.f. benbow. 1987. the costs and benefits of co-operation between the australian lycaenid butterfly jalmenus evagoras and its attendant ants. behavioural ecology and sociobiology 21: 237–248. pierce, n.e., m.f. braby, a. heath, d.j. lohman, j. mathew, d.b. rand & m.a. travassos. 2002. the ecology and evolution of ant association in the lycaenidae. annual reviews of entomology 47: 733–771. pringle, e.l., g.a. henning & j.b. ball (eds.). 1994. pennington's butterflies of southern africa. kaapstad: struik. prins, a.j. 1978. hymenoptera. pp. 825–872. in: werger, m.j.a. (ed.). biogeography and ecology of southern africa. the hague: junk. retief, e. & p.p.j. hermann. 1997. plants of the northern provinces of south africa: keys and diagnostic characters. pretoria: national botanical institute. roos, p. & g.a. henning. 2000. the heidelberg copper butterfly. african wildlife 54(2): 24 –27. schlosz, m. & p. schlosz. 1990. a ‘sound’ observation. metamorphosis 27: 24–25. schlosz, m. 1991. the call of pupae. metamorphosis 2(2): 19–20. thomann, h. 1901. schmetterlinge und ameisen. beobachtung über einer symbiose zwischen lycaena argus l. und formica cinerea mayr. jaresbericht der naturforschenden gesselschaft graubündens 44:1–40. thomas, j.a., g.w. elmes, j.c. wardlaw & m. woyciechowski. 1989. host specificity among maculinea butterflies in myrmica ant nests. oecologia 79(4): 452–457. williams, m.c. 1996. studying the early stages of lepidoptera in southern africa. metamorphosis 7(2): 56–64. issn 0075-6458 65 koedoe 47/1 (2004) terblanche 2 life history.qxd 2004/04/08 02:19 page 65 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 dippenaar marion.qxd the biodiversity and species composition of the spider community of marion island, a recent survey (arachnida: araneae) t.t. khoza, s.m. dippenaar and a.s. dippenaar-schoeman khoza, t.t., s.m. dippenaar and a.s. dippenaar-schoeman. 2005. the biodiversity and species composition of the spider community of marion island, a recent survey (arachnida: araneae). koedoe 48(2): 103–107. pretoria. issn 0075-6458. marion island, the larger of the prince edward islands, lies in the sub-antarctic biogeographic region in the southern indian ocean. from previous surveys, four spider species are known from marion. the last survey was undertaken in 1968. during this study a survey was undertaken over a period of four weeks on the island to determine the present spider diversity and to record information about the habitat preferences and general behaviour of the species present. three collection methods (active search, tullgren funnels and pitfall traps) were used, and spiders were sampled from six habitat sites. a total of 430 spiders represented by four families were collected, myro kerguelenesis crozetensis enderlein, 1909 and m. paucispinosus berland, 1947 (desidae), prinerigone vagans (audouin, 1826) (linyphiidae), cheiracanthium furculatum karsch, 1879 (miturgidae) and an immature salticidae. the miturgid and salticid are first records. neomaso antarticus (hickman, 1939) (linyphiidae) was absent from samples, confirming that the species might have been an erroneous record. key words: araneae, biodiversity, desidae, linyphiidae, marion island, miturgidae, salticidae, spiders. t.t. khoza & s.m. dippenaar, school of molecular and life sciences, university of limpopo, private bag x1106, sovenga, 0727 republic of south africa; a.s. dippenaarschoeman, arc-plant protection research institute, private bag x134, queenswood, 0121 republic of south africa/ department of zoology and entomology, university of pretoria, republic of south africa. issn 0075-6458 103 koedoe 48/2 (2005) introduction the prince edward island group has a surprising number of terrestrial invertebrates. most species are specially adapted to survive in the harsh conditions. the islands remained largely pristine, except for marion island. increased human activities led to a number of introduced species being documented. from marion, two arachnid orders have been recorded: acari (mites) represented by 60 species (van pletzen & kok 1971; hänel & chown 1998), and araneae (spiders) represented by four species (lawrence 1971). the first two spider species reported from marion island were myro paucispinosus berland, 1947 (desidae) and porrhomma antartica hickman, 1939 (now neomaso antarticus) (linyphiidae) (berland 1947). during a south african expedition in 1968, a further two species, myro kerguelenensis o.p.-cambridge, 1876 and erigone vagans audouin, 1826 (now prinerigone vagans), were added to the list (lawrence 1971). ledoux (1991), in his revision of the spiders of the french sub-antarctic islands (crozet and kerguelen), a group of islands some 950 km east of marion island, discussed the marion island spiders as well. he transferred porrhomma antartica to neomaso antarticus and recognised two subspecies of m. kerguelenensis, namely m. kerguelenensis crozetensis (the subspecies recorded from marion, prince edward and crozet islands), and m. kerguelenensis kerguelenensis o.p.-cambridge, 1876 that occurs only on kerguelen island. the only other reference to spiders of the southern ocean islands was by pugh (2004), who indicated the occurrence of neomaso antarticus from the prince edward islands as a “problem entry”, as it is based only on published reports (berland 1947) and has never been collected again. during this study a survey was undertaken over a period of four weeks (11 april 30 may 2004) on the island to determine the latest spider diversity and to record their habitat preference, with notes on their general behaviour. study area the southern ocean, stretching from the coast of antarctica to the antarctic polar frontal zone, contains several isolated groups of islands. the prince edward island group lies in the southern indian ocean and consists of two islands, prince edward island and marion island, separated by about 19 km of water. prince edward is uninhabited, while marion, the larger of the prince edward island group, is home to weather and research stations manned year-round, and is managed by south africa as a special nature reserve. marion island is 1770 km southeast of port elizabeth and a little further from antarctica. this island originated through volcanic eruptions and is only about 500 000 years old. it is one of the peaks of a coalescing shield volcano and is about 1300 m above sea level. mean monthly air temperature of the island closely follows that of the ocean and ranges from 3.6 °c (august) to 7.9 °c (february). sunshine averages 3.6 hrs per day. rain occurs on average 317 days per year, more frequently in winter, with a yearly average of 2 500 mm. snow and frost may occur in any month of the year, with snow on average about 80 days per year. northwesterly winds are predominant, frequently reaching gale force (> 55 km/h) and occur about 107 days of the year. the climate can therefore be summarised as cold, wet and windy (http://marion.sanap.org.za/weather. html). there are relatively few influencing factors beyond its maritime climate (joubert 2004). the island consists of a watery terrain with little vegetation and the absence of trees is largely due to the persistent strong westerly winds (http://www.btintorner). vegetation consists mainly of about 187 species of flowering plants, ferns, lichens, mosses, grasses and liverworts (http://cbp.inxcom.com). methods spiders were collected at marion island (46º54's, 37º50'e) over a period of four weeks (11 april 30 may 2004) from six different sites. collecting took place within approximately a 2-km radius of the research station that is situated on the northeastern side of the island. the description of the sites are as follows: skua ridge—a rocky, barren fell field area with only a few plants consisting mostly of azorella selago hook.fil.; tafelberg—similar to skua ridge but with more densely vegetated patches and at a higher altitude; ship’s cove—an area with bare patches and areas covered with tussock grass poa cookii hook.f. and alpine water fern blechnum penna-marina h.christ with some areas exposed to salt spray from the sea; research station—a mire covered with a mixture of b. penna-marina, agrostis magellanica lam., acaena magellanica vahl and uncinia compacta r.br., as well as patches of cotula plumosa hook.f. and b. penna-marina; an area between research station and trypot beach (rstb)—covered with b. penna-marina; trypot beach —an area similar to ship’s cove covered with b. penna-marina, c. plumosa and p. cookii (hänel & chown 1998). three different collecting methods (eardley & dippenaar 1996) were used to catch spiders from the different areas. pitfall trapping (50 ml containers, 5.5 cm height x 3.7 cm diameter) half-filled with 70 % ethanol (etoh) and buried into the ground with the upper rim level with the ground surface was conducted at the trypot beach and research station sites. the first set of pitfall traps at both locations was buried in exposed areas with less cotula growth with five traps in one row, placed one metre apart. the second set of traps at the research station site was buried in thicker growth of cotula, while that at the trypot beach site was planted at the edge of the sea exposed to salt spray. the pitfall traps at research station were kept open during the whole sampling period and were checked and emptied daily. those at trypot beach were kept open for a period of five days and were also checked and emptied daily. spiders were caught by actively searching through the sample sites. active searching was done at tafelberg (both in densely vegetated patches and on gray lava), skua ridge (on both azorella selago koedoe 48/2 (2005) 104 issn 0075-6458 cushions and rocks) and ship’s cove beach (in both p. cookii and b. penna-marina areas). active searching was done daily on a single sample site. plant material (agrostis castellana boiss. & reut., a. stolonifera l., sononia sp. and b. penna-marina) was collected from research station and rs-tb and put into ten tullgren funnels. the tullgren funnels were operated for two weeks. they were checked every morning to remove old plant material while new material was added late in the afternoon. old material was checked before being discarding, and spiders stuck in the plant material were collected by hand. collected spiders were preserved in 70 % ethanol and studied using stereo and light microscopes. voucher specimens are housed in the national collection of arachnida, at the arc-plant protection research institute in pretoria. results and discussion a total of 430 individuals represented by four families and four species were collected. these included: myro kerguelenesis crozetensis and m. paucisipinosus (desidae); prinerigone vagans (linyphiidae), as well as a new record cheiracanthium furculatum (miturgidae) and an immature salticidae, which could unfortunately not be identified to species level (table 1). prinerigone vagans: these are very small brown spiders with a shiny reddish tint. the males are easily recognised by the row of large tubercles on the anterior surface of the chelicerae and the large apophysis on the femur of the palp. their size ranges from issn 0075-6458 105 koedoe 48/2 (2005) table 1 number of spiders of each species collected from marion island during april and may 2004, with locations from which they were collected and method of collection. (imm = immatures, as = active searching, pt = pitfall trapping, tf = tullgren funnel) spider species location method male female imm total linyphiidae prinerigone vagans trypot beach pt 6 16 9 31 ship's cove as 55 49 10 114 research station pt, tf 2 11 9 22 total 63 76 28 167 desidae myro paucispinosus trypot beach tf, as 0 1 2 3 tafelberg as 0 0 6 6 research station pt, tf 0 0 3 3 skua ridge as 0 0 6 6 total 0 1 17 18 myro kerguelenesis crozetensis trypot beach tf, as, pt 1 7 6 14 ship's cove as 0 0 2 2 tafelberg as 5 3 35 43 research station pt, tf 10 4 64 78 skua ridge as 47 9 50 106 total 63 23 157 243 salticidae tafelberg as 0 0 1 1 miturgidae cheiracanthium furculatum skua ridge as 1 0 0 1 total 127 100 203 430 1.3–1.5 mm. a total of 167 specimens were collected comprising of 76 females, 63 males and 28 immatures. they were collected from three of the six locations with all three collecting methods (table 1). pitfall trapping was the least successful, since they spin a web on the upper rim of the pitfall traps. in patches of p. cookii, they occurred in clusters with more than a hundred individuals per cluster. a group of spiders was found at the base of the plant with the delicate sheet webs on top of the grass blades. most specimens caught in c. plumosa patches were collected from disturbed areas occupied by penguins and seals. here the spiders were mostly single and did not form clusters as in the p. cookii patches. prinerigione vagans is an old world species with a wide distribution (platnick 2005). this species is frequently found in disturbed habitats and was introduced by man onto the island. of the two myro species, m. paucisipinosus is less common on marion and only 18 specimens were collected, represented by one adult female and five immature females, two immature males, while the rest were immature. this species can be distinguished by its larger size (10-12 mm), darker colour and especially the presence of fur-like setae on the legs and body. ledoux (1991) provided illustrations of both the male and female genitalia. myro paucisipinosus specimens were collected mainly by hand and with the tullgren funnels from four of the six study sites (table 1). it was absent from ship’s cove. this species is known from marion and crozet (lawrence 1971; platnick 2005) and prince edward islands (lawrence 1971). myro paucispinosus was previously collected from under rocks, and are more common at high altitudes (lawrence 1971). in this study they were mainly collected by active searching and were always found in rocky areas next to cushions of azorella selago at higher altitudes. the egg sac found with the mature female was attached to the under side of rocks and camouflaged with dirt (lawrence 1971). myro kerguelenesis crozetensis is more abundant and a total of 243 specimens were collected represented by 63 females, 23 males and 157 immatures. they have been collected from five of the six sites but were most abundant at skua ridge (106 specimens), research station (78), tafelberg (43) and trypot beach (14) (table 1). only two immature specimens were collected from ship’s cove. they occur sympatrically with m. paucisipinosus and have been collected from the same pit traps. myro k. crozetensis is smaller (4.9-5.5 mm) than m. paucisipinosus and more yellowish in colour without the fur-like setae on the legs and body. iilustrations of the genitalia are provided by ledoux (1991). during this study they were mostly caught from under rocks at lower altitudes by active searching and less frequently with pitfall trapping. immature males were mostly caught by tullgren funnels from blechnum penna-marina and agrostis stolonifera. no webs were observed, only egg sacs (smaller than that of m. paucispinosus) on the ground camouflaged with brownish stripes. cheiracanthium furculatum: one male specimen was collected by hand from skua ridge. the species can be recognised by the creamish-yellow body with the ocular area and mouthparts black-brown; male palp distinctive in having both the cymbial apophysis and retrotibial apophysis long and sharply pointed (lotz 1995). this species is commonly found throughout the southern african region as well as from the cape verde islands, gabon, rwanda, tanzania and zaire (lotz 1995). it is synanthropic and commonly found in houses in south africa where they are also responsible for most of the cytotoxic spider bites (newlands 1975). the immature specimen of the family salticidae was found at tafelberg under rocks. the genitalia are very important in species identification and unfortunately it could not be identified (wesolowska pers. comm.). no specimens of neomaso antarticus were collected. koedoe 48/2 (2005) 106 issn 0075-6458 conclusion during the survey conducted in 1968 a total of 71 specimens were collected from two families (desidae and linyphiidae). during the current survey a total of 430 individuals from four families were collected. spiders were collected from five of the six sampling sites with no spiders found in rs-tb where the only collecting method used was tullgren funnels. this study confirms the observation of plugh (2004) that neomaso antarticus was wrongly reported to occur on marion island. further surveys are needed to locate adult specimens of the salticid species collected during this study to determine its identity. the distribution of cheiracanthium furculatum, a synantropic species that is of medical importance, also needs to be monitored acknowledgements the authors thank mr r mecer (the capacity building program for climate change research (cbpccr)) for field assistance and advice and both stellenbosch university and the stellenbosch university/deat/usaid cbp-ccr for sponsoring the study and the provision of a student bursary and dr wanda wesolowska of the wroclaw university for the assistance with the salticid identification. references berland, l. 1947. croisière du bougainville aux iles australes françaises. memoirs of the museum of natural history, paris 20: 53-64. eardley, c.d. & a.s. dippenaar-schoeman. 1996. collection and preparation of material. i. collecting methods. pp: 23–35. in: uys, v.m. & r.p. urban (eds). how to collect and preserve insects and arachnids. plant protection research institute, handbook 7. pretoria: agricultural research council. hänel, c. & s.l. chown. (1999). an introductory guide to the marion and prince edward island special nature reserves. fifty years after annexation. pretoria: department of environmental affairs & tourism. joubert l. 2004. love and desolation in the land of foo. getaway 15: 105 109. lawrence r.f. 1971. araneida. pp. 301-313. in: van zinderen bakker, e.m., j.m. winterbottom & r.a dyer (eds.). marion and prince edward island: report on the south african biological and geological expedition 1965-1966. cape town: a.a. balkema. ledoux, j. -c. 1991. araignées des îles subantarctiques françaises (crozet et kerguelen). revue arachnologique 9: 119-164. lotz, l.n. 1995. the genus cheiracanthium (araneae: clubionidae) in southern africa. msc thesis, university of the free state, bloemfontein. newlands, g. 1975. review of the medically important spiders of southern africa. south african medical journal 49: 823-826. platnick, n.i. 2005 the world spider catalog, version 5.5. (http://research.amnh.org/entomology/spiders/catalog. pugh, p.j.a. 2004. biogeography of spiders (araneae: arachnida) on the islands of the southern ocean. journal of natural history 38: 1461-1487. van pletzen, r. & d.j. kok. 1971. oribatei. pp. 314-318. in: van zinderen bakker, e.m., j.m winterbottom & r.a. dyer (eds.). marion and prince edward island: report on the south african biological and geological expedition 1965-1966. cape town: balkema. issn 0075-6458 107 koedoe 48/2 (2005) << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <feff00550073006100720065002000710075006500730074006500200069006d0070006f007300740061007a0069006f006e00690020007000650072002000630072006500610072006500200064006f00630075006d0065006e00740069002000500044004600200063006f006e00200075006e00610020007200690073006f006c0075007a0069006f006e00650020006d0061006700670069006f00720065002000700065007200200075006e00610020007100750061006c0069007400e00020006400690020007000720065007300740061006d007000610020006d00690067006c0069006f00720065002e0020004900200064006f00630075006d0065006e00740069002000500044004600200070006f00730073006f006e006f0020006500730073006500720065002000610070006500720074006900200063006f006e0020004100630072006f00620061007400200065002000520065006100640065007200200035002e003000200065002000760065007200730069006f006e006900200073007500630063006500730073006900760065002e002000510075006500730074006500200069006d0070006f007300740061007a0069006f006e006900200072006900630068006900650064006f006e006f0020006c002700750073006f00200064006900200066006f006e007400200069006e0063006f00720070006f0072006100740069002e> /nor <feff004200720075006b00200064006900730073006500200069006e006e007300740069006c006c0069006e00670065006e0065002000740069006c002000e50020006f00700070007200650074007400650020005000440046002d0064006f006b0075006d0065006e0074006500720020006d006500640020006800f80079006500720065002000620069006c00640065006f00700070006c00f80073006e0069006e006700200066006f00720020006800f800790020007500740073006b00720069006600740073006b00760061006c00690074006500740020006600f800720020007400720079006b006b002e0020005000440046002d0064006f006b0075006d0065006e0074006500720020006b0061006e002000e50070006e006500730020006d006500640020004100630072006f0062006100740020006f0067002000520065006100640065007200200035002e00300020006f0067002000730065006e006500720065002e00200044006900730073006500200069006e006e007300740069006c006c0069006e00670065006e00650020006b0072006500760065007200200073006b00720069006600740069006e006e00620079006700670069006e0067002e> /sve <feff0041006e007600e4006e00640020006400650020006800e4007200200069006e0073007400e4006c006c006e0069006e006700610072006e00610020006e00e40072002000640075002000760069006c006c00200073006b0061007000610020005000440046002d0064006f006b0075006d0065006e00740020006d006500640020006800f6006700720065002000620069006c0064007500700070006c00f60073006e0069006e00670020006600f60072002000700072006500700072006500730073007500740073006b0072006900660074006500720020006100760020006800f600670020006b00760061006c0069007400650074002e0020005000440046002d0064006f006b0075006d0065006e00740065006e0020006b0061006e002000f600700070006e006100730020006d006500640020004100630072006f0062006100740020006f00630068002000520065006100640065007200200035002e003000200065006c006c00650072002000730065006e006100720065002e00200044006500730073006100200069006e0073007400e4006c006c006e0069006e0067006100720020006b007200e400760065007200200069006e006b006c00750064006500720069006e00670020006100760020007400650063006b0065006e0073006e006900740074002e> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 filelist convert a pdf file! page 1 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 cleaver.qxd a vegetation description and floristic analyses of the springs on the kammanassie mountain, western cape g. cleaver, l.r. brown and g.j. bredenkamp cleaver,g., l.r. brown and g.j. bredenkamp. 2004. a vegetation description and floristic analyses of the springs on the kammanassie mountain, western cape. koedoe 47(2): 19-36. pretoria. issn 0075-6458. the kammanassie mountain is a declared mountain catchment area and a cape mountain zebra equus zebra zebra population is preserved on the mountain. the high number of springs on the mountain not only provides water for the animal species but also contributes to overall ecosystem functioning. long-term conservation of viable ecosystems requires a broader understanding of the ecological processes involved. it was therefore decided that a classification, description and mapping of the spring vegetation of the kammanassie mountain be undertaken. a twinspan classification, refined by braun-blanquet procedures, revealed 11 major plant communities that could be related to geological origin. habitat factors associated with differences in vegetation include topography, soil type and grazing. descriptions of the plant communities include diagnostic species as well as prominent and less conspicuous species of the tree, shrub and herbaceous layers. the results also indicate a high species richness compared to similar regions and the difference between plant communities of wet and dry springs. this data is important for long-term monitoring of the spring ecosystems as well as for the compilation of management plans. keywords: springs, kammanassie mountain, braun-blanquet, plant communities, twinspan, plant species richness. g. cleaver and l.r. brown (lrbrown@unisa.ac.za), applied behavioural ecology research unit, department of nature conservation, unisa, private bag x6, florida, 1710 republic of south africa; g.j. bredenkamp, african vegetation and plant diversity research centre, department of botany, university of pretoria, pretoria, 0002 republic of south africa. issn 0075-6458 19 koedoe 47/2 (2004) introduction the kammanassie nature reserve was established in 1978 to conserve the cape mountain zebra equus zebra zebra population present in the area and the natural ecosystems as part of the mountain catchment area. the area has a high species richness compared to similar vegetation types (cleaver et al. 2003; cleaver 2004). a large number of springs is found scattered throughout the reserve contributing to the high species richness and different ecosystem functioning while also providing drinking water to various animals, especially the cape mountain zebra (cleaver 2004). a total of 53 springs have been located on the kammanassie mountain. during 1998, it was noticed that certain springs were drying up. this was cause for concern for the survival of cape mountain zebra on the reserve. therefore, spring monitoring was initiated on the kammanassie mountain during 1999. flora occurring at flowing springs were thought to be unique and in danger of being lost should further springs dry up on the mountain. it was decided to determine the floristic composition of springs to give an indication of what flora is currently found at the springs and what species could be lost should further flowing springs dry up on the reserve. this information is important for future management purposes. furthermore, the department of water affairs and forestry (dwaf) abstract approximately 0.65 x 106 m3/a of groundwater from five production boreholes on the cleaver.qxd 2004/10/05 10:02 page 19 kammanassie nature reserve. the klein karoo rural water supply scheme (kkrwss) has been in operation since 1993 and the scheme supplies purified domestic water, at subsidised rates, to the town of dysselsdorp, to farms in the olifants river valley, to tributary valleys downstream of the stompdrift and kammanassie dams, and to the gamka river valley downstream of calitzdorp. the effects of large-scale groundwater abstraction on springs is not clearly understood and this study forms part of a larger research project funded by the water research commission to assess the environmental impacts of large-scale groundwater abstraction from table mountain group aquifers on ecosystems in the kammanassie nature reserve. in order to understand ecosystems and the management thereof (bredenkamp 1982; bezuidenhout 1993; brown & bezuidenhout 2000), it is important to classify, describe and map the different vegetation types found in the spring areas of the kammanassie mountain (van rooyen et al. 1981; brown & bezuidenhout 2000). this data will also be used in the long-term monitoring of the spring ecosystems as well as for compiling a management plan. the study area the kammanassie mountain complex is situated between the towns of uniondale in the east and de rust/dysselsdorp in the northwest and west (fig. 1). the mountain is an inselberg within the little karoo between the swartberg and outeniqua mountains. the total area of the mountain range, managed by the western cape nature conservation board (wcncb), is 49 430 ha of which 21 532 ha are privately owned and a declared mountain catchment. the remaining 27 898 ha are state land, of which 17 661 ha have been declared state forest. the kammanassie nature reserve is situated between koedoe 47/2 (2004) 20 issn 0075-6458 fig. 1. location of the study area. cleaver.qxd 2004/10/05 10:02 page 20 33°33's–33°37's and 22°27'e– 23°01'e and was proclaimed a protected area in 1978 as an important water catchment area (cleaver 2002). according to rebelo (1996) and lubke (1996) the kammanassie mountain falls into the fynbos and thicket biomes with small pockets of the forest biome present in the kloofs on the southern slopes of the kammanassie nature reserve (cleaver 2002). the average monthly rainfall and minimum and maximum temperatures for the study area are indicated in fig. 2. the reserve receives rain throughout the year, with an average annual rainfall of ± 450 mm. the highest annual rainfall of 1216 mm and lowest of 242 mm were recorded in 1981 and 1984 respectively. the hottest months on the issn 0075-6458 21 koedoe 47/2 (2004) kammanassie nature reserve are december to february with maximum temperatures of 35 ºc and the coldest months are june and july with minimum temperatures of -2 ºc (fig. 2). the kammanassie is one of the prominent east-west trending ranges comprising the southern branch of the cape fold belt. it was formed as a result of north-south oriented compressive stress during the cape orogeny 123-200 million years ago (hälbich & greef 1995). the kammanassie nature reserve is an eroded remnant of the kammanassie mega-anticline (hälbich & greef 1995). the kammanassie mountain range comprises almost exclusively the resistant quartz arenites of the table mountain group, overlain on the lower slopes by the shale of the bokkeveld group (kotze 2001). a very important shale marker horizon, the cedarberg formation (50–120 m thick) occurs within the table mountain group, separating formations of the peninsula formation from the lithologies comprising the nardouw formation (kotze 2001). according to kotze (2001), the peninsula formation is a highly competent succession of medium to coarse-grained, thickly bedded, grey sandstone. the nardouw formation weathers more brownish, and thin shale intercalations are more common, than in the peninsula formation. as a result the nardouw formation is more ductile (kotze 2001). soil generally forms a thin (<1 m) veneer of silty sands as a result of the steep slopes of the kammanassie mountain and predominantly quartzitic rocks. locally, clayey soil occurs in association with weathered shale horizons and, in particular, the cedarberg formation (cleaver et al. 2003). methods relevés were compiled at 52 of the known springs on the kammanassie mountain. spring localities were identified by kammanassie nature reserve staff, who monitor these springs (flow rates and water samples) annually during february/march. the coordinates of each spring were determined using an omnistar global positioning system (gps). to facilitate this monitoring, each individual spring was given a number, beginning at 01. the prefix kamm is the standard wcncb monitoring code used for the kammanassie nature reserve while the standard fig. 2. the average monthly rainfall and the mean average minimum and maximum temperatures of the kammanassie nature reserve. cleaver.qxd 2004/10/05 10:02 page 21 wcncb code for water monitoring is indicated by the letter ‘w’ (van der walt 1993). floristic composition of springs (woody species, grasses and forbs) was surveyed at the origin of the spring using the braun-blanquet cover-abundance scale (mueller-dombois & ellenberg 1974). the area sampled varied from spring to spring and was determined to include at least 95% of all floral species present at the site. care was taken to sample only homogeneous spring vegetation and not to include adjacent drier communities. fieldwork was carried out between february and april 2002. environmental data recorded included aspect, altitude, slope, geology, soil texture, percentage rock cover and rock size (small, small/medium, medium, medium/large and large). a first approximation of the main plant communities was derived by applying the two-way indicator species analysis (twinspan) (hill 1979) to the floristic data set. further refinement of the classification was achieved by braun-blanquet procedures (bredenkamp et al. 1989; kooij et al. 1990; bezuidenhout 1993; eckhart 1993; brown & bredenkamp 1994). the results are presented in a phytosociological table produced by using megatab (hennekens 1996a). springs showing similar plant communities were plotted on an arcview (ver. 3.2) geographical information system (gis) generated map, to indicate their localities. plant taxon names conform to those given by goldblatt and manning (2000). these names may therefore differ from the plant species list used by the turboveg database (hennekens 1996b), which is based on the precis database of the national botanical institute (nbi) in pretoria, as on date 1997. red data status follows hilton-taylor (1996). results classification the results are presented in a phytosociological table (table 1). the analysis resulted in the identification of the following fourteen plant communities, which can be grouped into 11 major community types: 1. blechnum tabulare-hippa frutescens fern community 1.1. hippia frutescens-helichrysum cymosum shrubland 1.2. hippia frutescens-helichrysum petiolare shrubland 1.3. hippia frutescens-ehrharta species grassland 2. eragrostis plana-juncus lomatophyllus grassland 3. berzelia intermedia-psoralea verrucosa shrubland 4. cliffortia ilicifolia-stoebe plumosa shrubland 5. ehrharta ramosa-aspalathus kougaensis shrubland 6. conyza canadensis-conyza scabrida shrubland 7. phragmites australis reed community 8. ehrharta erecta-rhus pallens shrubland 9. mentha longifolia-thelypteris confluens shrubland 10. pelargonium radulifolium-salvia namaensis shrubland 11. ehrharta erecta-crassula biplanata shrubland description of spring plant communities in the descriptions of the different spring plant communities, all species groups refer to table 1. the position of the springs (spring numbers) with similar plant communities is indicated in with same colour and symbol in fig. 3. 1. blechnum tabulare-hippa frutescens fern community the springs with vegetation belonging to this community are mainly found on southern slopes on the kammanassie mountain. altitude varies between 837-1394 m above sea level. the soil is loamy to clayey with rock cover varying between 5-40%. all the springs of this community are perennial and undisturbed. diagnostic species include the dwarf shrub hippa frutescens and the ferns blechnum tabulare and blechnum inflexum (species group a). this community can be sub-divided into the following three sub-communities: koedoe 47/2 (2004) 22 issn 0075-6458 cleaver.qxd 2004/10/05 10:02 page 22 1.1 hippia frutescens-helichrysum cymosum shrubland the vegetation of springs kamm/w/22 and 37 (mannetjiesberg), kamm/w/31 (elandsvlakte) and kamm/w/52 (wildealsvlei) fall within this sub-community. altitude varies between 1038-1361 m a.s.l. all springs occur on gentle slopes (1-2°) with a south-southwestern aspect. rock cover varies between 5 % to 40 %. all four springs were flowing but the spring kamm/w/22 had water though not flowing at the time of the survey. the shrub layer comprises 50-60 % and the herbaceous layer 30-40 %. diagnostic species include the shrub helichrysum cymosum, the moss racomitrium lamprocarpum and the restio calopsis paniculata (species group b). the dwarf shrub hippia frutescens (species group a) is dominant while the ferns blechnum tabulare and blechnum inflexum (species group a) are locally dominant in this sub-community. the shrubs helichrysum petiolare (species group c), psoralea verrucosa and small fern hymenophyllum tunbridgense (species group f) are prominent. other prominent species include the cosmopolitan weed spergularia rubra and the moss jamesoniella species. 1.2 hippia frutescens-helichrysum petiolare shrubland the hippia frutescens-helichrysum petiolare sub-community is found at springs kamm/w/35 and kamm/w/36 (mannetjiesberg) which were both flowing during the period of the survey. these springs occur on gentle slopes (1-2°) between 1343-1394 m a.s.l. where the aspect is south-southwest. soil texture is sandyloamy and with small-to-medium rocks, with a coverage of 30-40 % of the soil surface. shrub cover is 70 % and herbaceous cover is 30 %. issn 0075-6458 23 koedoe 47/2 (2004) fig. 3: location of the vegetation communities of springs on the kammanassie nature reserve. cleaver.qxd 2004/10/05 10:02 page 23 koedoe 47/2 (2004) 24 issn 0075-6458 ta bl e 1 p hy to ci ol og ic al ta bl e of th e ve ge ta tio n of s pr in gs o n th e k am m an as si e m ou nt ai n cleaver.qxd 2004/10/05 10:02 page 24 issn 0075-6458 25 koedoe 47/2 (2004) ta bl e 1 (c on tin ue d) cleaver.qxd 2004/10/05 10:02 page 25 koedoe 47/2 (2004) 26 issn 0075-6458 ta bl e 1 (c on tin ue d) cleaver.qxd 2004/10/05 10:02 page 26 issn 0075-6458 27 koedoe 47/2 (2004) ta bl e 1 (c on tin ue d) cleaver.qxd 2004/10/05 10:02 page 27 diagnostic species include the shrubs helichrysum petiolare and cliffortia burchelli (species group c). the shrub hippia frutescens and the fern blechnum tabulare (species group a) are prominantly present. the shrub psoralea verrucosa (species group f) is also conspicuous in this subcommunity. 1.3 hippia frutescens-ehrharta species grassland this sub-community is found at the following springs: kamm/w/05 (buffelsdrif), kamm/w/11 (wildebeesvlakte), kamm/w/33 (elandsvlakte) and kamm/w/46 (upper diepkloof). the spring kamm/w/05 dried up in december 2001 and kamm/w/33 dried up in 2000. at the time of the survey kamm/w/11 had water but was not flowing and kamm/w/46 was flowing. altitude varies between 837-1154 m a.s.l. on a gentle slope. all these springs occur on south-facing slopes. soil is sandy, while rock cover is very low, at most localities less than 5 %. shrubs cover an estimated 60 % of the area and the herbaceous layer 40 %. diagnostic species include the shrubs pelargonium cordifolium and rhus tomentosa, the grass ehrharta species , the restios isolepsis verrucosula and juncus capensis and aquatic plant eleocharis limosa (species group d). the shrub hippia frutescens, ferns blechnum tabulare and blechnum inflexum (species group a) are also prominent. other prominent local species include the shrub helichrysum cymosum (species group b), restio juncus lomatophyllus and grasses panicum ecklonii and pennisetum macrourum (species group e) and watsonia fourcadei (species group f). (fig. 4) 2. eragrostis plana-juncus lomatophyllus grassland the vegetation of springs kamm/w/02 and 03 (kleinfonteinsberg), kamm/w/34 (elandsvlakte) and kamm/w/53 (mannetjiesberg) represent this plant community. some springs dried up, e.g., kamm/w/02 in 1978, kamm/w/03 in 1999, kamm/w/34 in 2000, but kamm/w/53 was flowing at the time of the survey. altitude varies considerably between 8671405 m above sea level, while slope also varies considerably between 1-11°. the koedoe 47/2 (2004) 28 issn 0075-6458 fig. 4. dominant plant families reflecting the percentage of total flora of springs on the kammanassie mountain. cleaver.qxd 2004/10/05 10:02 page 28 aspect is northeast to north-northeast. rock cover is low and is less than 5 % in most cases while the soil texture is loamy-clay. shrub cover is between 10-40 % and the herbaceous layer between 40-60 %. diagnostic species include grasses such as eragrostis plana, panicum ecklonii, pennisetum macrourum, restio juncus lomatophyllus, shrub gunnera perpensa and the forbs pelargonium grossularioides and vellereophyton vellereum (species group e). other prominent species that occur locally include grasses ehrharta species (species group d) and ehrharta erecta (species group k), the tall reed phragmites australis (species group j), and sedge mariscus congestus (species group m). 3. berzelia intermedia-psoralea verrucosa shrubland a total of nine springs, namely; kamm/w/06 (buffelsdrif), kamm/w/07, 32 and 45 (elandsvlakte), kamm/w/23 (mannetjiesberg), kamm/w/40 (kleinberg), kamm/w/43 and 44 (groenplaat) and kamm/w/47 (kleinrivier) form the habitat of the berzelia intermedia-psoralea verrucosa shrubland. the spring kamm/w/06 dried up at the end of 2001, kamm/w/40 and 43 dried up in 2000 and kamm/w/47 is dry but the year its spring dried up is unknown. the springs kamm/w/07, 32, 45 and 23 were all flowing at the time of this survey. altitude ranges between 927–1404 m a.s.l. these springs occur on a slope between 1–17° and aspect varies greatly. rock size is small to medium while rock cover varies between 5–40 %. all these springs are situated on the southern slopes of the kammanassie mountain. shrubs and herbs have a canopy cover of 50 %. the diagnostic species from group f include: the shrubs psoralea verrucosa, berzelia intermedia, erica curviflora, helichrysum anomalum, clutia alaternoides, helichrysum zeyheri and plecostachys polifolia, the corm watsonia fourcadei, the restios platycaulos callistachyus, elegia filacea, ischyrolepis ocreata and juncus dregeanus, the forbs selago glomerata, centella eriantha, melasma scabrum, and the small fern hymenophyllum tunbridgense. other locally dominant species are the shrubs ficinia nigrescens (species group h) and helichrysum cymosum (species group b), the grasses, eragrostis plana and panicum ecklonii (species group e) and ehrharta ramosa (species group h) and the restio juncus lomatophyllus (species group e). further conspicuous species locally present include the shrubs stoebe plumosa and cliffortia ilicifolia (species group g) and anthospermum aethiopicum (species group i). the mosses symphyogyna podophylla, riccardia species, sematophyllum species and campylopus species, are prominent locally. 4. cliffortia ilicifolia-stoebe plumosa shrubland the vegetation of springs kamm/w/08 (elandsvlakte), kamm/w/24, 25 and 38 (paardevlakte), kamm/w/39 (mannetjiesberg), kamm/w/30 (bergplaas) and kamm/w/41 (upper diepkloof) represents a cliffortia ilicifolia-stoebe plumosa shrubland. all these springs were dry during the survey period. most of these springs, excluding kamm/w/08 (2000) and kamm/w/30 (2001)), dried up before 1993. springs belonging to this shrubland are relatively flat with a slight gradient that varies between 1–2°. altitude varies between 758 m–1475 m a.s.l., and aspect varies greatly. soil texture is sandy-loam, while the rock cover varies between 40–50 % with little to no rocks present in certain localities. the shrub and herbaceous layer has a cover of 50 %. diagnostic species include the shrubs stoebe plumosa, cliffortia ilicifolia and the grass pentaschistis species 831, which grows in large tufts (species group g). other locally dominant species include the shrubs issn 0075-6458 29 koedoe 47/2 (2004) cleaver.qxd 2004/10/05 10:02 page 29 helichrysum cymosum (species group b), helichrysum zeyheri (species group f), aspalathus kougaensis (species group h) and the grass eragrostis plana (species group e). the large shrub chrysanthemoides monilifera, tree protea neriifolia and succulent crassula ericoides (not included in table 1) also occur locally. 5. ehrharta ramosa-aspalathus kougaensis shrubland this plant community occurs at the following three springs: kamm/w/21 (leeuwblad) and kamm/w/28 and 29 (bergplaas). all these springs dried up between 1999 and 2000. this community is found at lower-lying areas with altitudes varying between 612 m–797 m a.s.l. on south or west facing aspects. the soil texture is very coarse gravel. medium-to-large rocks with coverage of between 70–80 % are common. shrub and herbaceous coverage is between 30–40%. species group h is diagnostic for this community and includes the shrubs aspalathus kougaensis, phylica paniculata, ficinia ramosissima and the grass ehrharta ramosa. eragrostis plana (species group e) is also present. another locally prominent species is the grass pentaschistis species (species group g). the following species are locally present: the shrubs dodonaea angustifolia, pteronia stricta, anomalanthus anguliger, the grasses panicum repens, pentameris species, the restios tetraria capillacea, juncus oxycarpus and forbs ursinia anthemoides and senecio ilicifolius. 6. conyza canadensis-conyza scabrida shrubland the vegetation of springs kamm/w/10 (wildebeesvlakte), kamm/w/20 (leeuwblad) and kamm/w/42 (ylandsrivier) represents the conyza canadensis-conyza scabrida shrubland. this plant community occurs at a lower altitude (586–927 m a.s.l.), with slopes of 1-3°. the soil texture varies from sandy, loamy to clay, while rock cover varies between 0–50 %. the shrub cover varies between 30–50 % and the herbaceous layer coverage is between 30–40 %. species diagnostic for this community include the invader conyza scabrida, the shrubs conyza canadensis, helichrysum nudifolium and anthospermum aethiopicum (species group i). the shrubs stoebe plumosa (species group g) and rhus pallens (species group k) are locally prominent within this community. 7. phragmites australis reed community the dominant vegetation of two springs kamm/w/26 (wildebeesvlakte) and kamm/ w/49 (rietfontein) is phragmites australis. at present these springs are both dry. kamm/w/26 dried up in 1999 and kamm/w/49 in 2001. altitude varies between 624–862 m a.s.l. both have a western aspect and occur on gentle slopes of between 1-3°. soil varies from clay to gravel and rock cover varies from 60 % to no rocks present in certain localities. the herbaceous layer is dominated by the reed phragmites australis and has a cover of 70 %. shrub coverage varies (10–30 %). this community has the reed phragmites australis as the diagnostic and dominant species (species group j). the tree rhus pallens (species group k) is locally prominent within the community. 8. ehrharta erecta-rhus pallens shrubland the four springs, kamm/w/14 (slawedam i) and kamm/w/16 (slawedam ii) and kamm/w/17 (rooielskloof) and kamm/ w/50 (rietfontein), are the habitat of the ehrharta erecta-rhus pallens shrubland. some springs dried up, viz., kamm/w/14 in koedoe 47/2 (2004) 30 issn 0075-6458 cleaver.qxd 2004/10/05 10:02 page 30 1994, kamm/w/17 in 1999, and kamm/w/16 date unknown. spring kamm/w/50 had water but was not flowing during the period of the survey. altitude is between 553 -602 m a.s.l. aspect varies between north to north-east and west. soil texture is coarse gravel, with mediumto-large rocks present with a coverage of between 60–80 %. this shrubland has a tree coverage of 10–50 %, a shrub coverage of 30–50 % and herbaceous coverage of 10–20 %. species from species group k are diagnostic for this community and include the trees tarchonanthus camphoratus and salix mucronata, the large shrub rhus pallens, the small shrub sutera campanulata, the grass ehrharta erecta, and forb pollichia campestris. the aromatic forb mentha longifolia (species group l) is prominently present. 9. mentha longifolia-thelypteris confluens shrubland the vegetation of springs kamm/w/01 (buffelsklip), kamm/w/09 and 51 (vermaaks river) represent this plant community. the spring kamm/w/01 was standing at the time of the survey, kamm/w/09 dried up in 1999 and kamm/w/51 dried up at the end of 2001. altitude is between 569–711m a.s.l. and aspect varies between west, south and northnorthwest on a gentle slope. soil texture is loamy-sandy with rock cover between 30–60 % and less than 5 % in certain areas. the shrub and herbaceous layer have a 50 % coverage. the tree calpurnia intrusa, the shrubs pelargonium zonale and stachys aethiopica, the forb mentha longifolia and fern thelypteris confluens (species group l) are diagnostic species of this community. the shrub conyza scabrida (species group i) and grasses ehrharta ramosa and ehrharta erecta (species group h and k) are locally dominant. 10. pelargonium radulifolium-salvia namaensis shrubland this plant community occurs at the following springs: kamm/w/12 and 13 (kleingeluk), kamm/w/15 (slawedam ii) and kamm/w/18 (solomanskraal). all the springs dried up before 1996. they are found on the northern slopes of the kammanassie mountain. these springs occur at altitudes between 533–684 m above sea level on gentle north to north-east slopes (2°). soil texture is predominantly sandy to coarse gravel, with rock cover between 20–50 % (medium-to-large). shrub coverage varies between 50–70 % and the herbaceous laver has a cover between 10–30 %. diagnostic species include the shrub salvia namaensis, the forb pelargonium radulifolium, the creepers dipogon lignosus and cissampelos capensis and the sedge mariscus congestus (species group m). the grass ehrharta ramosa (species group h) and tree rhus pallens (species group k) occur locally while the sedge mariscus thunbergii is locally conspicuous. 11. ehrharta erecta-crassula biplanata shrubland the vegetation of the springs (kamm/w/19 and 48) at wagenpadnek represent this plant community. kamm/w/19 dried up in 1999, while kamm/w/48 was standing at the time of the survey. these springs occur on gentle (1°) western or southern slopes at an altitude of 718–732 m a.s.l. soil texture is sandy with small-to-medium rocks with a rock cover of 10–20 %. shrub cover is 60 % and the herbaceous layer has a coverage of 30 %. the forb, species 814 (sent in to herbarium for identification), and succulent crassula biplanata are diagnostic species (species group n). the shrub rhus pallens and grasses pennisetum macrourum and ehrharta issn 0075-6458 31 koedoe 47/2 (2004) cleaver.qxd 2004/10/05 10:02 page 31 erecta (species group e and k) are locally prominent. floristic analysis of springs on the kammanassie mountain a total of 244 plant species were recorded for spring flora, representing 145 genera and 71 families (table 2). flowering plants are represented by monocotyledoneae with 63 species in 7 families and eudicotyledoneae with 156 species in 43 families. the pteridophytes, with 12 species in 8 families and 12 species in 12 families of bryophytes (mosses) represent the non-flowering plants. only one species of gymnosperm was found. a ratio of 6:1 dicotyledon species to monocotyledon species was found for the spring flora. seven families dominate the flora found at springs on the kammanassie mountain. the two largest families are the asteraceae with 41 species, reflecting 14.5 % of the total flora and the poaceae with 25 species (8.8 % of the total flora) (fig. 4). these are followed by much smaller, yet significant families, represented by the cyperaceae with 19 species (6.7 % of the total flora), restionaceae with eight species (2.8 % of the total flora), juncaceae with six species (2.1 % of the total flora), rosaceae with six species (2.1 % of the total flora) and scrophulariaceae also with six species (2.1 % of the total flora) (fig. 4). two red data species were found, representing 1 % of the flora of springs on the kammanassie mountain. the endangered erica sp nov (ericaceae) was sampled at sample plot 18 (kamm/w/19) and the rare agathosma affinis (rutaceae) in sample plot 24 (kamm/w/25). both these two springs dried up before the kkrwss started abstraction. a total of 11 alien species or cosmopolitan weeds were found at springs on the kammanassie mountain. geological origin of spring plant communities the springs within the hippia frutescenshelichrysum shrubland (plant community 1.1) are all springs emanating at a perched water table of the nardouw/cedarberg formation, while springs within the hippia frutesens-helichrysum petiolare shrubland (plant community 1.2) are all emanating at perched water tables on the nardouw/kouga formation. springs of the hippia frutescensehrharta species grassland (plant community 1.3) has its geological origin at shallow springs emanating at perched water tables on cedarberg shale and peninsula formation. thus all the springs within the blechnum tabulare-hippa frutescens fern (plant community 1) have their geological origin at shallow, perched springs. the eragrostis plana-juncus lomatophyllus grassland (plant community 2) has a mixed geological origin, emanating from the nardouw formation and the kouga/baviaanskloof formation that is in close contact with the bokkeveld group. all these springs originates predominantly from the nardouw formation with cedarberg shale contact. the nine springs found in the berzelia intermedia-psoralea verrucosa shrubland (plant community 3) are all shallow and perched, with six emanating from the nardouw/ cedarberg formation, one from the nardouw/kouga formation, and two from the peninsula formation. a total of five of the seven springs of the cliffortia ilicifolia-stoebe plumose shrubland (plant community 4) has a predominantly nardouw/kouga formation geological origin. other origins include the nardouw formation with combined lithological and structural control. therefore, the dominant geological origin is from the nardouw formation. the ehrharta ramosa-aspalatus kougaensis shrubland (plant community 5) springs have their geological origin from the nardouw formation. springs from the conyza canadensis-conyza scabrida shrubland koedoe 47/2 (2004) 32 issn 0075-6458 cleaver.qxd 2004/10/05 10:02 page 32 issn 0075-6458 33 koedoe 47/2 (2004) table 2 an alphabetical list of plant families, collected at springs on the kammanassie mountain, indicating the number of genera and species in each family bryophytes aneuraceae 1 1 bartramiaceae 1 1 brachytheciaceae 1 1 bryaceae 1 1 dicranaceae 1 1 fissidentaceae 1 1 grimmiaceae 1 1 hedwigiaceae 1 1 jungermanniaceae 1 1 pallaviciniaceae 1 1 pottiaceae 1 1 sematophyllaceae 1 1 12 12 pteridophytes aspleniaceae 1 2 blechnaceae 1 2 dennstaedtiaceae 1 1 dryopteridaceae 1 1 gleicheniaceae 1 1 hymenophyllaceae 1 1 pteridaceae 1 3 thelypteridaceae 1 1 8 12 gymnosperm taxodiaceae 1 1 1 1 monocotyledons araceae 1 1 cyperaceae 9 19 dioscoreaceae 1 1 iridaceae 3 3 juncaceae 1 6 poaceae 13 25 restionaceae 6 8 34 63 eudicotyledons aizoaceae 2 2 anacardiaceae 1 3 apiaceae 2 2 .../eudicotyledons apocynaceae 1 1 araliaceae 2 2 asteraceae 19 42 brassicaceae 1 1 bruniaceae 1 1 campanulaceae 2 2 caryophyllaceae 4 4 celastraceae 2 3 convolvulaceae 1 1 crassulaceae 2 5 dipsacaceac 1 1 droseraceae 1 1 ebenaceae 2 3 ericaceae 1 13 euphorbiaceae 1 1 fabaceae 9 14 geraniaceae 1 5 gunneraceae 1 1 lamiaceae 4 5 linaceae 1 1 malvaceae 2 3 menispermaceae 1 1 molluginaceae 1 1 moraceae 1 1 orobanchaceae 1 1 oxalidaceae 1 1 polygalaceae 2 3 primulaceae 1 2 proteaceae 2 3 ranunculaceae 1 1 rhamnaceae 1 1 rosaceae 2 6 rubiaceae 1 1 rutaceae 2 4 salicaceae 2 2 santalaceae 1 1 sapindaceae 1 1 scrophulariaceae 3 6 solanaceae 1 2 urticaceae 1 1 90 156 total 145 244 families genera species families genera species cleaver.qxd 2004/10/05 10:02 page 33 koedoe 47/2 (2004) 34 issn 0075-6458 (plant community 6) have a mixed geological origin, with two springs originating from the nardouw formation and one shallow, perched spring on cedarberg shale. the springs from the phragmites australis reed community (7), the ehrharta erecta-rhus pallens shrubland (plant community 8) and pelargonium radulifolium-salvia namaensis (plant community 10) originate on the nardouw formation-kouga/baviaanskloof formation close to the contact with the bokkeveld group. these plant communities, therefore, have similar geological origins. plant community 9, mentha longifolia-thelypteris confluens shrubland, originates from the nardouw formation while the ehrharta erecta-crassula biplanata shrubland (plant community 11) has two springs, both shallow and perched, on cedarberg shale. discussion a total of 244 plant species were found at 52 springs on the kammanassie mountain, representing 145 genera and 71 families. of the 71 plant families, one gymnosperm, eight families of pteridophytes (11 %), 12 families of bryophytes (mosses)(17 %), seven families of monocotyledoneaes (10 %) and 43 families of dicotyledons (60 %) were recorded for the spring vegetation. the high presence of asteraceae (14.5 % of the total flora) corresponds to goldblatt & manning (2000) and was similar to the vermaaks, marnewicks and buffelsklip valley floristic composition which had 16 % asteraceae present (cleaver 2004). cyperaceae (6.7 % of the total flora) with 19 species and restionaceae (2.8 % of the total flora) were the second and third most dominant families present at the springs on the kammanassie mountain. the ratio of 6:1 dicotyledon to monocotyledon species observed for the spring flora compares well to that of the spring flora at fernkloof nature reserve (four dicotyledons to one monocotyledon species) also situated in mountain fynbos vegetation (mostert 2003). this ratio of spring flora for kammanassie nature reserve (6:1) is also higher than the ratio of 3:1 for the cape floristic region as given by goldblatt & manning (2000). however, the species richness for the kammanassie mountain springs is 47 species/100 m² compared to the much higher 126 species/100 m² for fernkloof nature reserve (mostert 2003). a possible explanation for this could be that the springs of the fernkloof nature reserve are mainly situated on moist southern slopes (mostert 2003) whereas a large number of springs on the kammanassie nature reserve are located on drier northern slopes. this, together with the higher average annual rainfall of 674 mm for the fernkloof nature reserve, compared to the average annual rainfall of 450 mm for the kammanassie mountain can explain the higher species richness found at fernkloof nature reserve. the cliffortia ilicifolia-stoebe plumosa shrubland represents springs that dried up before 1993. these springs are also perched and comprise drier shrubland vegetation, with grasses prominent. both the rare agathosma affinis (rutaceae) and endangered erica sp nov (ericaeae) occur within this plant community, making it an important plant community to manage and conserve. the erica sp nov, an endangered species, also occurs within the ehrharta erecta-rhus pallens community. flowing springs and the ones that have water have very different plant communities to those that are dry. a difference in plant species composition was also found between springs that have been drier longer than others. the species composition of flowing springs differ strongly with those of drier springs in that ferns, mosses and water plants dominate these communities. springs that have been dry for a number of years, have a shrub, grass and restio dominance while springs that have dried up recently (within the last 3 years) show a distinct difference, with no shrub layer present and the herbaceous layer being dominant. there is a strong relationship between spring geological origin and plant communities cleaver.qxd 2004/10/05 10:02 page 34 issn 0075-6458 35 koedoe 47/2 (2004) found at springs, whether shallow perched type springs on the nardouw/cedarberg or nardouw/kouga contacts, nardouw formation with combined litological and structural control or nardouw formation-kouga/baviaanskloof formation close to the bokkeveld group. spring (kamm/w/09) of the mentha longifolia-thelypteris confluens shrubland (plant community 9), dried up in september 1999, but still (three years later) has similar vegetation as the other flowing springs within this community. a possible explanation for this could be that this spring (kamm/w/09, 635 m a.s.l.) occurs in the alluvium of the vermaaks valley and is not a perched spring. the water table, obtained from a monitoring hole close to this spring, was found to be approximately 5 m below surface (cleaver et al. 2003). therefore, although this spring stopped flowing at surface the water table at this spring is at a sufficient level to sustain the wetter plant community found at this site. springs kamm/w/08 (1135 m a.s.l.) and 22 (1404 m a.s.l.) also originate at the cedarberg shale contact between the peninsula and nardouw formations and dried up in 1999. these two perched springs have a drier plant community, completely reliant on rainfall, as the water table is not within reach of the plants at such high-altitude springs. it is interesting that the plant community at spring kamm/w/09 has not changed sufficiently to fall within a drier type community. should the water table at this spring decline it could result in a change of the present plant community and further monitoring is essential at this spring to better understand the relationship between the water table and plant community present. conclusion no similar vegetation descriptions have previously been carried out on the springs of the kammanassie mountains. this research therefore provides valuable data on these ecosystems. the plant species identified from this study can now be included into the kammanassie nature reserve management plan and will result in a more comprehensive plant species list for the reserve. the localities and plant communities within which previously unknown, rare, and vulnerable plant species occur were identified during this survey. the wetness of a spring, period of time the spring had been dry and a spring’s geological origin have influenced plant community composition of the springs on the kammanassie mountain. if more springs dry up on the mountain, plant communities at these springs could change over time and species diversity could be expected to decrease as is the case with the dry and wet communities of this study. important water dependent plants will be lost and ‘wetland’ plant communities will be transformed into dry shrub and grass dominated areas. the time it takes for these changes in plant species composition to form another plant community will, however, have to be determined through long-term monitoring of the different springs on the kammanassie nature reserve. these vegetation surveys and descriptions provide baseline information that allows similar surveys to be conducted in future. these monitoring data could then be compared with this study to determine if changes/shifts in plant communities have occurred. since a change in habitat conditions will result in the loss of not only ‘wetland’ plant communities, but a host of other organisms such as invertebrates, frogs and fish species that depend on these plants for their existence, will also disappear. this will result in a reduction of flora and fauna biodiversity. the wcncb should therefore continue to monitor rare and vulnerable species found during this survey to ensure their survival by applying the correct management strategies. monitoring of spring vegetation should continue to determine changes in plant communities over time related to springs drying up. this could form part of a cleaver.qxd 2004/10/05 10:02 page 35 koedoe 47/2 (2004) 36 issn 0075-6458 long-term monitoring programme on the kammanassie nature reserve. acknowledgements the water research commission, unisa (old technikon sa) and western cape nature conservation board contributed to the funding of this research project. mr mc smart (dwaf-geohydrology department) is thanked for his assistance in classifying spring origins. references bezuidenhout, h. 1993. syntaxonomy and synecology of the western transvaal grasslands. ph.d thesis, university of pretoria, pretoria. bezuidenhout, g.j. 1982. 'n plantekologiese studie van die manyeleti-wildtuin. d.sc thesis. university of pretoria, pretoria. bredenkamp, g.j., a.f., joubert, a.f. & h. bezuidenhout. 1989. a reconnaissance survey of the vegetation of the plains in the potchefstroom-fochville-parys area. south african journal of botany 55: 199-206. brown, l.r & h. bezuidenhout. 2000. the phytosociology of the de rust section of the mountain zebra national park, eastern cape. koedoe 43/1: 1-18. brown, l.r. & g.j. bredenkamp. 1994. the phytosociology of southern section of the borakalalo nature reserve, south africa. koedoe 37: 5972. cleaver g. 2002. the management plan of the kammanassie nature reserve. western cape nature conservation board, uniondale. unpublished. cleaver g. 2004. environmental impacts of largescale groundwater abstraction on ecosystems of the kammanassie mountain. m.tech thesis, unisa, florida. cleaver, g., l.r. brown, g.j. bredenkamp, m.c. smart & c.j. de w. rautenbach. 2003. assessment of environmental impacts of groundwater abstraction from table mountain group (tms) aquifers on ecosystems in the kammanassie nature reserve and environs. pretoria: water research commission (report no. 1115/1/03). eckhart, h.c. 1993. a synecological study of the vegetation of the north-eastern orange free state. m.sc. thesis. university of pretoria, pretoria. goldblatt, p & j. manning. 2000. cape plants : a conspectus of the cape flora of south africa. national botanical institute of south africa, mbg press, missouri botanical garden. (strelitzia 9). hälbich i.w. & g.j. greef. 1995. final report on a structural analysis of the west plunging nose of the kammanassie anticline. geology department, university of stellenbosch. (consulting report to dwaf, directorate geohydrology). hennekens, s.m. 1996a. megatab: a visual editor for phytosociological tables. user’s guide version october 1996, ibn-dlo, wageningen and lancaster university, lancaster. hennekens, s.m. 1996b. turboveg: a software package for input, processing and presentation of phytosociological data. user’s guide, version july 1996, ibn-dlo, wageningen and lancaster university, lancaster. hill, m.o. 1979. twinspan: a fortran programme for arranging multivariate data in an ordered two-way table classification of individuals and attributes. cornell university, new york. hilton-taylor, c. 1996. red data list of southern african plants. pretoria: national botanical institute. lubke, r.a. 1996. eastern mixed nama karoo. p. 55 in: low, a.b. & a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. kooij, m.s., g.j. bredenkamp & g.k. theron. 1990. classification of the vegetation of the b land type in the north-western orange free state. south african journal of botany 56: 309318. kotze j.c. 2001. towards a management tool for groundwater exploration in the table mountain sandstone fractured aquifer. pretoria: water research commission (report no k5/729). rebelo, a.g. 1996. mountain fynbos. p. 55 in: low, a.b. & a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. mostert, c. 2003. environmental impacts of largescale groundwater abstraction on springs of the fernkloof nature reserve. fernkloof nature reserve, hermanus. unpublished report. mueller-dombois, d. & h. ellenburg. 1974. aims and methods of vegetation ecology. new york: wiley. van der walt, j.p.l. 1993. collecting baseline data. cape nature conservation. unpublished. van rooyen, n., g.k. theron & n. grobelaar. 1981. a floristic description and structural analysis of the plant communities of the phinda milia-pafuri-wambiya area in the kruger national park, republic of south africa. 1. the hydrophilous communities. south african journal of botany 47: 213-246. cleaver.qxd 2004/10/05 10:02 page 36 filelist convert a pdf file! page 1 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 filelist convert a pdf file! page 1 page 2 page 3 page 4 filelist convert a pdf file! page 1 page 2 article information authors: marna herbst1,2,3 michael g.l. mills1,4 affiliations: 1mammal research institute, department zoology and entomology, university of pretoria, south africa 2carnivore conservation group, endangered wildlife trust, johannesburg, south africa 3south african national parks, conservation services, kimberley, south africa 4tony and lisette lewis foundation, johannesburg, south africa correspondence to: marna herbst email: marnah@sanparks.org postal address: po box 110040, hadison park, kimberley 8306, south africa keywords african wildcat; capture techniques; chemical immobilisation; habituation; kalahari dates: received: 04 aug. 2009 accepted: 11 dec. 2009 published: 24 may 2010 how to cite this article: herbst, m. & mills, m.g.l., 2010, ‘techniques used in the study of african wildcat, felis silvestris cafra, in the kgalagadi transfrontier park (south africa/botswana)’, koedoe 52(1), art. #939, 6 pages. doi: 10.4102/koedoe.v52i1.939 copyright notice: © 2010. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn:issn: 2071-0771 (online)(online) techniques used in the study of african wildcat, felis silvestris cafra, in the kgalagadi transfrontier park (south africa/botswana) in this original research... open access • abstract • introduction • study area    • kgalagadi transfrontier park • method    • capture techniques       • cage traps / drop door traps       • darting       • radio collars       • habituation • results    • capture success       • cage traps       • darting       • radio collaring       • habituation • discussion • conclusion • acknowledgements • references abstract (back to top) the techniques used for the capture, marking and habituation of african wildcats (felis silvestris cafra) in the kalahari are described and evaluated in this paper. african wildcats were captured, with either baited cage traps or chemical immobilisation through darting. darting proved to be a more efficient and less stressful way of capturing cats. very high frequency (vhf) radio collars fitted with activity monitors were especially effective in the open habitat of the kalahari for locating and maintaining contact with cats; they also aided in determining if the cats were active or resting in dense vegetation. the habituation of individual cats to a 4× 4 vehicle proved to be time consuming, but it provided a unique opportunity to investigate the feeding ecology and spatial organisation of cats through direct visual observations. conservation implications: in describing and comparing the various methods of capture, handling and release of the african wildcats that we followed during our study in the southern kalahari, we recommend the most efficient, least stressful method for researchers to follow – both in relation to time and energy, as well as in terms of the impact on the animals being studied. introduction (back to top) the african wildcat (felis silvestris cafra), is widely distributed throughout the african continent and listed by the international union for conservation of nature (iucn) as least concern (nowell 2008). however, status and density estimates of african wildcats are poorly known throughout most of its range. therefore, the ecological status of wildcat populations is frequently determined from incomplete or unverified data (nowell & jackson 1996). previous research efforts on african wildcats have focused on scat analyses and opportunistic sightings of cats in their natural environment (palmer & fairall 1988; smithers 1971; smithers & wilson 1979; stuart 1977; stuart 1982). the aim of this study was to gain insight into the population genetics and behavioural ecology of african wildcats in the southern kalahari. this required the capture of cats for the fitting of radio collars, taking morphometric measurements and obtaining dna samples. radio telemetry was crucial for locating individual cats for the collection of data on feeding behaviour, home range and movement patterns. investigating the foraging and social behaviour relied on the habituation of certain individuals for direct observations. steel, wire, mesh and tomahawk cage traps are widely used in the live trapping of small mammals, for example in the european wildcat and domestic cats (biró, szemethy & heitai 2005), lynx (breitnemoser & haller 1993), kodkod (dunstone et al. 2002), blanford’s foxes (geffen et al. 1992; geffen & macdonald 1993), leopard cat (grassman & tewes 2005), caracal (marker & dickman 2005; melville 2004), black-footed cats (sliwa 2004, 2006), dhole (grassman et al. 2005), ferrets (norbury, norbury & heyward 1998) and civits (jennings, seymour & dunston 2006). the successful capture and release of an animal is not only determined by the capture of the animal, but also by how the animals are handled, transported and kept after capture (ebedes, du toit & van rooyen 1996). this paper provides detailed information on the methodology involved in capturing, immobilising and habituating of african wildcats in the southern kalahari. study area (back to top) kgalagadi transfrontier park this study was initiated in march 2003 and continued until december 2006 (46 consecutive months) in the kgalagadi transfrontier park (ktp), which comprises the kalahari gemsbok national park (south africa) and the adjacent gemsbok national park in botswana. the ktp is a 37 000 km2 semi-arid wilderness area in the southern kalahari, described as the western form of the kalahari duneveld (mucina & rutherford 2006), consisting of extreme open savannah of acacia erioloba, acacia haemotoxylon and desert grasses. the study was primarily conducted in a 53 km2 area surrounding the leeudril waterhole (26º28’17.7” s, 20º36’ 45.2” e), in the south of the park, and included the nossob riverbed together with adjacent calcrete ridges, rhigozum veld and dune areas (figure 1). method (back to top) all capture, darting and handling of african wildcats were approved by the ethics committee, university of pretoria, (ec 030305-007) and south african national parks animal use and care committee (sanparks aucc). approval to conduct research in the botswanan side of the ktp was obtained from the office of the president: op 46/1 cvii (48), with a supplementary permit from the department wildlife and national parks (9 july 2006). capture techniques cage traps / drop door traps cage traps (50 cm × 50 cm × 150 cm) were constructed from welded mesh, with a single sliding door. a stepping plate mechanism towards the rear end of the cage activated the trap door. the size of the cages permitted cats to enter fully before depressing the plate, causing the door to drop. bait, either locally bought chicken pieces or fresh road kills, suspended from a wire over the plate, was used as lure. additionally, cat urine was collected opportunistically whenever following a focal cat, stored in plastic bags and added to baited traps as a supplementary attractant for other cats (6 out of 12 cats were caught with the use of urine as attractant). cages were sometimes camouflaged by hiding them in vegetation, or covering the sides (only two of the 12 cats were caught when the cages were camouflaged). the stepping plate was covered with soil to give it a more natural feel. the traps were set late in the afternoons and checked daily, early in the mornings. when a cat was found inside the trap, the far end was covered with a blanket in an attempt to provide a measure of security for the cat. a 40 cm × 40 cm crush plate, attached to a steel rod, was inserted at the front of the trap and, slowly and gently, the cat was pushed towards the back of the cage. in this way, the cat could be trapped at the far end of the cage, from where it was possible to hand inject it through the wire mesh. zoletilr (tiletamine hydrochloride with benzodiazephine derivative zolazepam in 1:1 combination). a dosage of approximately 2.5 mg/kg was used for all cats caught by this method. once anaesthetised, cats could be removed from the cages without difficulty, whereupon standard body measurements were taken (table 1). a small skin sample was collected for molecular analysis and, if relevant, a radio collar was fitted. all procedures were conducted as quickly as possible and in the immediate vicinity of the trap. on completion of the necessary procedures, the cat was returned to the shaded cage and left to recover from the anaesthesia. it was released when it had fully recovered. table 1: standard body measurements collected from all african wild cats trapped and darted during 2003–2006 in the kgalagadi transfrontier park figure 1: study site in the kgalagadi transfrontier park, indicating the area around the nossob riverbed and leeudril waterhole, where african wildcats, felis silvestris cafra, were radio collared and monitored darting a co2 rifle (dan-inject jm standard model) was used to propel a standard dart syringe (10.5 mm, 1.5 ml capacity) and fitted with a small rubber stopper to reduce penetration. owing to the small size of the cats, it was necessary to lower the co2 pressure in the rifle as much as possible to reduce the projectile velocity and, in so doing, lessen the impact and therefore the chances of injury to an animal. as a trade-off, the range over which the dart could be propelled had to be reduced. cats were thus always stalked to within 10 m. cats caught by darting were immobilised with a combination of drugs and an appropriate antidote as follows (p. buss & d. govender, pers. comm.): either butorphanol (1.38 mg/kg) and medetomidine (0.4 mg/kg), with the antidote of antipamezole administered at five times the medetomidine dose (mg) intramuscularly and naltrexone administered at 10 times the butorphanol dose (mg) intramuscularly, or zoletil (1.58 mg/kg) and medetomidine (0.07 mg/kg), with the antidote of antipamezole administered at 6.25–12.5 times the medetomidine dose (mg) intramuscularly. zoletil does not have an antidote. radio collars african wildcats were fitted with radio collars from africa wildlife tracking cc, weighing 80 g – 85 g, with external antennae of 20 cm and a battery life of up to 18 months. radio collars were each fitted with an activity monitor to assist in the remote detection of cat activity. cats were detected with a two or three element handheld yagi antenna by traversing the home range of the individual study animal and using the dune crests as high vantage points, using a telonics handheld receiver. habituation the open, clear spaces of the kalahari provide ideal conditions for visual observation of animals (begg 2001; mills 2003), although the stealthy nature of cats, especially at night, required close proximity to the focal animal at all times. all radio collared cats were habituated to the presence of the research vehicle, allowing the researchers to closely follow individual cats without any obvious influence on their behaviour. this was achieved by patiently following cats daily for the first week after initial capture and collaring, at a distance of 50 m – 100 m, while keeping the engine running. habituation appeared to be facilitated by keeping the engine running in the beginning and slowly moving closer to the cats. after 1 week, the following distance was gradually decreased, until the cats could be followed from a distance of 10 m – 30 m without them looking back at the vehicle. wildcats were followed on a rotational system, allowing continuous monitoring of a focal animal every night. cats were located at night by radio tracking, with the initial visual contact being made with a 1 000 000-candle spotlight. once a cat was located, the headlights of the research vehicle were usually sufficient to follow cats, with the spotlight used only periodically to re-establish contact when lost in patches of denser vegetation, or when cresting sand dunes. care was taken to keep the spotlight trained behind the cat – to neither influence their hunting success negatively by blinding them, nor positively, by dazzling prey animals. results (back to top) capture success cage traps african wildcats were frequently spotted during opportunistic searches and cage traps were placed in close vicinity to these spots. seven of the ten cats caught in the study site were trapped after being spotted in a specific area. only three cats were caught by randomly placing the traps in the study site. trapping success for african wildcats in the kalahari was 1.4 cats per 100 trap nights. the trapping frequency between wildcats is highly variable and for african wildcats it was estimated at 73 trap nights per new cat, compared to the results of the european wildcat (felis s. silvestris) (biró et al. 2004; corbett 1979), at 860 and 299 trap nights per new cat, respectively. trapping of feral domestic cats (felis s. catus) ranged between 75 and 823 trap nights per new cat (barratt 1997; biró et al. 2004; bromley 1986; corbett 1979; daniels et al. 2001; molsher 2001, 2006), for lynx (lynx canadensis) it was 67 trap nights per new cat (mech 1980), ocelot (leopardus pardalis) was 116 trap nights per new cat (dillon & kelly 2008) and leopard cat (prionailurus bengalensis) 405 trap nights per new cat (grassman et al. 2005). the main drawback of cage traps appeared to be the reluctance of wildcats to enter, as well as their non-selective nature (table 2). loss of bait could possibly have been attributed to the ineffective setting of cages. bait was stolen on numerous occasions, by smaller mammals such as the yellow mongoose (cynictis penicillata) and rodents; in some instances it was consumed by ants. table 2: the percentage capture success expressed as the total of traps (n = 1244) used during all the trapping days (n = 301) in the kgalagadi transfrontier park darting during two darting expeditions, consisting of four nights each (10 h – 14 h per night), in august 2005 and january 2006, a total of 18 african wildcats were successfully darted, with only one injury reported. cats were spotted by driving up and down the riverbed, constantly scanning with spotlights in two vehicles and looking for retinal reflections. when cats were spotted, the research vehicle slowly moved in the direction of the cat, maintaining visual contact with the vehicle headlights and a spotlight. assistants with spotlights in the second vehicle acted as spotters, and when necessary, pedestrian herders directed the cat towards the darting vehicle. the cat was slowly approached until it stopped and a clear shot was possible. cats were darted from a distance of no more than 10 m. once successfully darted, a cat was followed at a distance of 30 m – 40 m, with spotlights, until it became fully immobilised. this was important, as a premature approach could have caused the cat to flee, leading to a temporary loss of contact with a highly vulnerable animal. within 10 min – 15 min after the drugs were administered, it was possible to walk up to the cat and carefully cover the head and eyes with a blanket. standard body measurements and genetic samples were taken, and in two cases the cats were fitted with radio collars. antidotes were very effective and cats regained full motor control within minutes after administering. african wildcats did not appear to associate the vehicle with the darting procedures, as two cats that were fitted with radio collars were easily habituated to the vehicle afterwards. the majority of cats were darted primarily to collect genetic material for molecular analysis and were not approached again afterwards. owing to the risk of missing the small target area on the thigh of a cat and potentially injuring it, only qualified, experienced wildlife veterinarians were employed in darting. african wildcats were immobilised on 31 occasions (13 cats were hand injected and 18 cats were darted). no fatalities were recorded, although the fate of the injured one is not known. radio collaring radio collaring proved to be invaluable for finding and following cats, as they do not return to a fixed den site and are difficult to find at night. the estimated total home range sizes (100% minimum convex polygon) were: adult male = 13.17 km² ± 7.32 km² (n = 5) and adult female = 11.75 km² ± 2.01 km² (n = 3) (herbst, unpublished). in total, 12 african wildcats were radio collared. only one female cat showed a slight irritation to the radio collar, symptomised by localised hair loss ten days after been collared. symptoms lasted for 4 weeks, with hair growing back gradually. the cat was monitored daily until all symptoms had disappeared. on two occasions, damaged radio collars were retrieved, (3 weeks and 2 months after being fitted) suggesting that the cats had fallen prey to a larger predator (one unknown and one confirmed from tracks as a caracal, caracal caracal). two radio-collared cats disappeared (a young female, two months after being fitted and a young male, two days after), either as a result of malfunctioning radio collars or emigration to an area outside the range searched. external antenna of radio collars broke off within 2–6 months, however, this did not seem to make a difference in the detection of cats, because the cats had known home ranges (herbst, unpublished) and searching for a signal from high dunes was almost always successful. habituation on average, the habituation period took 73.8 h ± 63.9 h (n = 8), although large individual differences occurred (table 3). in general, females were easier to habituate (average 36.7 h ± 5.8 h; n = 3). three radio-collared and habituated females had litters during the study period and dens and kittens could be approached without difficulty. kittens were extremely curious and would investigate the research vehicle of their own accord. male cats were more difficult to habituate (96 h ± 74 h; n = 5), as they move faster and over a much larger area than females, making observations of males more difficult. habituation was lost quickly and maintaining the maximum degree of habituation required that weekly contact with each cat was maintained. table 3: the time (days and hours) invested to habituate three female and fve male african wildcats in the kgalagadi transfrontier park habituated african wildcats were visually observed for 1538 h (males for 657 h, females for 881 h) on a rotational basis. continued observations of selected individuals provided detailed information on sexual and seasonal differences in diet, foraging behaviour, movement patterns, reproduction and inter-specific interactions. discussion (back to top) long-term and intensive field studies on smaller cats are still uncommon and even the common species have not been well studied (macdonald & loveridge, in press; nowell & jackson 1996). the reason for this is the relative difficulty associated with studying small felids. previous research on african wildcats was based on opportunistic sightings, scat and stomach analysis (palmer & fairall 1988; smithers 1971; smithers & wilson 1979; stuart 1977). their nocturnal behaviour and general shy and elusive nature, make it practically impossible to study cats in their natural environment without the aid of radio telemetry. radio telemetry has become more reliable and efficient since the 1980s (nowell & jackson 1996); recently, radio collars have been designed smaller, lighter and reliable enough for the use on smaller cats. however, in spite of the advances in technology, the time required to catch smaller cats for radio collaring purposes poses a challenge. the trapping frequency of african wildcats is comparable with frequencies of the trapping of feral domestic cats (f. s. catus) (barratt 1997; molsher 1999, 2001). this is much lower than the results on european wildcats (f.s. silvestris) (biró et al. 2004; corbett 1979), which are difficult to catch, the reason possibly being that these populations in europe have declined, are fragmented and, in many places, are already extinct (nowell 2008). for black-footed cats (felis nigripes), the trapping frequency was one cat for 100–200 trap nights (including recaptures) (a. sliwa, pers. comm.). african wildcats in the kalahari were regularly spotted during our study period, therefore it is believed that densities are much higher in the kalahari than in europe. the results in this study not only confirm the difficulty of catching african wildcats, but also emphasise the general low success rate of trapping small carnivores in the southern kalahari. mainly trap door cages, with various combinations of bait and urine to attract cats were used. positioning cages in areas of high animal activity should increase the selectivity of the trapping efforts (boddicker 1999). our results suggest that, after an extensive search in the riverbed with a spotlight and placing of traps close to sightings of cats, the success of trapping increased in comparison with randomly placed traps. the use of a co2 dan inject dart gun proved to be the best method in the capture of free range african wildcats. the time and cost effectiveness of this capture method was enhanced with the use of drugs combined with antidotes. once all the data and measurements were collected from the cats, they could be revived with the antidote and the darting operation could continue. special care and qualified personnel (two wildlife veterinarians and four assistants in two vehicles) were needed to assist with darting operations, because the target animal was so small. the cost of qualified veterinarians and personnel needed in a darting operation is high; however, to obtain a representative sample size using only conventional trapping methods might have taken the researcher another few years of intensive fieldwork. it was relatively easy to habituate african wildcats to a research vehicle (590 h were needed to habituate eight cats). the kalahari is the ideal location to study small carnivores, such as african wildcats, because the openness of the environment makes it possible to follow them, even at night (begg 2001; mills 2003). although there were large individual differences between the times needed to habituate individuals (table 3), it was possible to collect data on feeding, hunting, reproduction and mating behaviour of african wildcats (herbst & mills 2010; herbst, unpublished). to achieve this, radio telemetry was essential and because african wildcats do not travel to the same extent than larger felids, it was feasible to traverse the whole study area in a few hours in search of a signal. this was enhanced by using high dunes as a vantage point. conclusion (back to top) for dispersed and elusive animals, radio collaring might be the key to obtaining appropriate data (kenward 2001). despite the advances in the use of satellites for radio tracking – platform transmitter terminals and global positioning system collars – they remain relatively expensive in comparison with the vhf transmitters (kenward 2001). in this study visual observations of habituated cats fitted with vhf transmitters enabled us to record valuable behavioural information on a nocturnal and secretive animal that more sophisticated and expensive tracking devices could not. this is the first report on the methodology of darting of wildcats (f. silvestris), and it proved to be a more efficient and less stressful method than cage trapping of african wildcats in the ktp. acknowledgements (back to top) this research was supported by the endangered wildlife trust’s carnivore conservation group (south africa), the elizabeth wakemen henderson charitable foundation, the mammal research institute and the molecular ecology and evolution programme (department of genetics) at the university of pretoria, the national research foundation (south africa), the kaplan awards program from wildlife conservation society and the skukuza marathon club. we thank sanparks and the department of wildlife and national parks, botswana for permission to conduct the study in the kgalagadi transfrontier park. n. du plessis (sanparks section ranger) is thanked for assistance in the chemical immobilisation of cats and dr p. buss and dr d. govender (sanparks wildlife veterinarians) for darting the cats. m. haupt, dr l. jansen van rensburg, p. odendaal and n. read are also thanked for their support in the darting operations. references (back to top) barratt, d.g., 1997, ’home range size, habitat utilisation and movement patterns of suburban and farm cats, felis catus’, ecography 20, 271−280. begg, c.m., 2001, ‘feeding ecology and social organization of the honey badgers (mellivora capensis) in the southern kalahari’, phd thesis, department of zoology and entomology, university of pretoria. biró, z., szemethy, l. & heitai, m., 2004, ‘home range size of wildcats (felis silvestris) and feral domestic cats (felis silvestris f. catus) in a hilly region of hungary’, mammalian biology 69, 302−310. boddicker, m.l., 1999, catch’n coyotes & other crit’rs, rocky mountain wildlife products, colorado. breitenmoser, u. & haller, h., 1993, ‘patterns of predation by reintroduced european lynx in the swiss alps’, journal of wildlife management 57, 135−144. corbett, l.k., 1979, ‘feeding ecology and social organization of wildcats (felis silvestris) and domestic cats (felis catus) in scotland’, phd thesis, department of zoology, university of aberdeen. daniels, m.j., beaumont, m.a., johnson, p.j., balharry, d., macdonald, d.w. & barratt, e., 2001, ‘ecology and genetics of wild-living cats in the north-east of scotland and the implications for the conservation of the wildcat’, journal of applied ecology 38, 146−161. dillon, a. & kelly, m.j., 2008, ‘ocelot home range, overlap and density: comparing radio telemetry with camera trapping’, journal of zoology (london) 275, 391−398. dunstone, n. durbin, l., wyllie, i., freer, r., jamett, g.a., mazolli, m. et al., 2002, ‘spatial organization, ranging behaviour and habitat use of the kodkod (oncifelis guigna) in southern chili’, journal of zoology (london) 257, 1−11. ebedes, h., du toit, j.g. & van rooyen, j., 2000, ‘capturing wild animals’ in j. du p bothma (ed.), game ranch management, 4th edn., n.p., van schaik publishers, pretoria. geffen, e., degen, a.a., kam, m., hefner, r. & nagy, k.a., 1992, ‘daily energy expenditure and water flux of free-living blanford’s foxes (vulpes cana), a small desert carnivore’, journal of animal ecology 61, 611−617. geffen, e. & macdonald, d.w., 1993, ‘activity and movement patterns of blanford’s foxes’, journal of mammalogy 74, 455–463. grassman, l.i. & tewes, m.e., 2005, ‘spatial ecology and diet of the dhole cuon alpinus (canidae, carnivora) in north central thailand’, journal of mammalogy 85, 29−38. grassman, l.i., tewes, m.e., silvey, n.j. & kreetiyutanont, k., 2005, ‘ecology of three sympatric felids in a mixed evergreen forest in north-central thailand’, journal of mammalogy 86, 29−38. herbst, m., n.d., ‘behavioural ecology and population genetics of the african wild cat, felis silvestris forster 1780, in the southern kalahari’, unpublished phd thesis, department of zoology and entomology, university of pretoria. herbst, m. & mills, m.g.l., 2010, the feeding habits of the southern african wildcat (felis silvestris cafra), a facultative trophic specialist, in the southern kalahari (kgalagadi transfrontier park, south africa/botswana), journal of zoology 280, 403–413. jennings, a.p., seymour, a.s. & dunstone, n., 2006, ‘ranging behaviour, spatial organization and activity of the malay civit (viverra tangalunga) on buton island, sulawesi’, journal of zoology (london) 268, 63−71. kenward, r.e., 2001, a manual for wildlife radio tagging, academic press, london. macdonald, d. & loveridge, a., in press, ‘the biology and conservation of wild felids’, oxford university press, oxford. marker, l. & dickman, a., 2005, ‘notes on the spatial ecology of caracals (felis caracal), with particular reference to namibian farmlands’, african journal of ecology 43, 73−76. mech, l.d., 1980, ‘age, sex, reproduction, and spatial organization of lynxes colonizing north-eastern minnesota’, journal of mammalogy 61, 261−267. melville, h., 2004, ‘behavioural ecology of caracal in the kgalagadi transfrontier park, and its impact on adjacent small stock production units’, msc thesis, department of wildlife management, university of pretoria. mills, m.g.l., 2003, kalahari hyenas: comparative behavioural ecology of two species, the blackburn press, new jersey. molsher, r.l., 1999, ‘the ecology of feral cats, felis catus, in open forests in new south wales: interactions with food resources and foxes’, phd thesis, school of biological sciences, university of sydney. molsher, r.l., 2001, ‘trapping and demographics of feral cats (felis catus) in central new south wales’, wildlife research 28, 631−636. mucina, l. & rutherford, m.c., 2006, ‘the vegetation of south africa, lesotho and swaziland’, strelitzia 19, south african national biodiversity institute, pretoria. norbury, g.l., norbury, d.c. & heyward, r.p., 1998, ‘space use and denning behaviour of wild ferrets (mustela furo) and cats (felis catus)’, new zealand journal of ecology 22, 149−159. nowell, k., 2008, felis silvestris,‘iucn 2008. 2008 iucn red list of threatened species’, viewed 20 april 2010, from http://www.iucnredlist.org nowell, k. & jackson, p., 1996, wild cats. status survey and conservation action plan, iucn, gland. palmer, r. & fairall, n., 1988, ‘caracal and african wild cat diet in the karoo national park and the implications thereof for hyrax’ south african journal of wildlife research 18, 30−34. sliwa, a., 2004, ‘home range size and social organisation of black-footed cats (felis nigripes)’, mammalian biology 69, 96−107. sliwa, a., 2006, ‘seasonal and sex-specific prey-composition of black-footed cats felis nigripes’, acta theriologica 51, 195−206. smithers, r.h.n., 1971, ‘the mammals of botswana’, museum memoirs of the national monument rhodes 4, 1−340. smithers, r.h.n. & wilson, v.j., 1979, checklist and atlas of the mammals of zimbabwe–rhodesia, trustees, national museums and monuments, zimbabwe–rhodesia, salisbury. stuart, c.t., 1977, ‘the distribution, status, feeding and reproduction of carnivores of the cape province’, department of nature and environmental conservation research report, mammals 1977, 91–174. stuart, c.t., 1982, ‘aspects of the biology of the caracal (felis caracal) schreber 1776, in the cape province of south africa’, msc thesis, department of zoology, university of natal. filelist convert a pdf file! page 1 page 2 page 3 page 4 saayman & slabbert.qxd a profile of tourists visiting the kruger national park m. saayman and e. slabbert saayman, m. and e. slabbert. 2004. a profile of tourists visiting the kruger national park. koedoe 47(1): 1–8. pretoria. issn 0075-6458 due to shrinking budgets for conservation and an increase in the number of government and privately owned parks, it has become very important for parks to determine who the tourists are who visit one of south africa’s top tourist attractions. the reason for this is that park management and marketers need to focus their efforts to optimise their limited resources. this can only be done once there is a clear understanding of who the market is, where they come from and what they expect. the literature study clearly showed that market segmentation is essential for the effective marketing of a tourism product or destination. two surveys were conducted, one in 2001 and a follow-up study in 2002, profiling tourists to the kruger national park. different months were chosen to conduct the two surveys in order to get a more comprehensive profile of tourists visiting the park in different seasons. key words: market segmentation, profile, market, tourism, tourist/s, tourism industry, kruger national park. m. saayman and e. slabbert, institute for tourism and leisure studies, potchefstroom university for che, private bag x6001, potchefstroom, 2520 republic of south africa (ontms@puknet.puk.ac.za). issn 0075-6458 1 koedoe 47/1 (2004) introduction one of the world’s best-known and most profitable national parks, the kruger national park (knp) in the republic of south africa, was established in 1898 as the sabie game reserve. the park is one of the largest parks in the world, with the main aim to protect the biodiversity of the region. the latter includes 147 mammal species, 500 species of birds, 116 species of reptiles, 34 species of amphibians, 49 species of fish, 457 species of trees and shrubs, 1500 species of smaller plants and countless species of insects (dennis 2000). the knp covers a total surface area of 19 624 km², with a total length of 350 km and an average width of 60 km. most of the park consists of flat grass and bushcovered plains, interrupted by the low lebombo mountains situated along the border of mozambique. negotiations are under way with neighbouring countries which will significantly enlarge the park to form the limpopo transfrontier conservation area. this will be the largest national park in the world. approximately 950 000 people visit this park each year, of which 80 % are south africans. to ensure continuity and growth, a destination (attraction) is dependent on, inter alia, tourists and the satisfaction of their needs. although total satisfaction of tourists’ needs are not the goal in itself, striving to achieve this enables the attraction to attain its own goals (strydom et al. 2000). to achieve maximum tourist satisfaction, marketing strategists divide the heterogeneous market into fairly homogeneous groups of tourists, a process that is referred to as market segmentation. through market segmentation one can develop a tourist profile that will enable knp to concentrate its resources and efforts so that maximum penetration of that market can be achieved (doole & lowe 2001). however, little is known about tourists who visit the park. the problem that knp must deal with is that the ‘product’ sought by tourists can be satisfied by a number of attractions/destinations. this creates the need to develop an overall competitive advantage for knp (hassan 2000) and makes the prosaayman & slabbert.qxd 2004/04/15 09:08 page 1 filing of tourists crucial (nickels & wood 1997). the number of parks and game reserves in south africa are on the increase, which leads to stiff competition. except for national parks, there are also parks at local, provincial and government levels, as well as more than 8000 privately owned game reserves. all these parks and game reserves are offering an ecotourism experience. other destinations/attractions are winning competitive battles through careful analysis and response to the core values and needs of the segmented travel market place (hassan 2000). knp needs to do the same. with regard to the international travel to south africa, research by south african tourism indicated that more than 80 % of these tourists travel to south africa for game and nature experiences (sat 2002). this creates yet another opportunity for marketers to profile tourists travelling to the knp (sat 2002). it is no longer possible to serve a total market and it is therefore necessary to segment markets and concentrate on a single segment or a number of segments (saayman 2001; kinnear et al. 1995). segments are evaluated according to a number of criteria, but the essence of the approach is to identify the most relevant characteristics of tourists seeking particular sets of benefits from their tourism and leisure purchases (laws 1997; pride & ferrell 1987). in this process, consumer behaviour plays an important role. the tourist does not make purchase decisions in isolation. the mix of cultural, social, personal, psychological factors and previous experiences, all which influence behaviour, is largely uncontrollable. because of the influence exerted upon patterns of buying, it is essential that as much effort as possible is put into understanding how these factors interact and ultimately how they influence decisions (lamb et al. 2002). the attractiveness of knp reflects the feelings, beliefs and opinions that an individual has about the park’s ability to provide satisfaction in relation to his or her special vacation needs (slabbert 2002). successful segmentation is the product of a detailed understanding of the market and will therefore take time (saayman 2001; mcdonald & dunbar 1995). market segmentation is also one of the keys to developing a sustainable competitive advantage based on differentiation, low cost, or a focused strategy (aaker 1998). knowing the profile characteristics of target groups can help marketing strategists to tailor the product or service and promote the product or service more effectively. each group can be targeted and reached with a distinct marketing mix (mcdonald & dunbar 1995). communication effects have a direct bearing on the prospective tourist’s decision to act. the prospective tourist decides whether or not to respond to the advertising by taking action (pritchard 1998). to be of use, markets need to be segmented according to attributes that can relate to the product or service, distribution, price and media (andereck & caldwell 1994). benefits of market segmentation include: long-term relationships can be formed with tourists who are brand loyal (nickels & wood 1997); gauging the park’s current position, as it is perceived by the tourists, relative to the competition; segmentation can help guide the proper allocation and use of marketing resources (strydom et al. 2000; semenik 2002); new markets can be detected and the success of marketing campaigns can be measured; scarce resources are not wasted and the marketing message/communication can be very specific; skills and insights used in segmenting the product market give the knp an important competitive advantage by identifying buyer groups that will respond favourably to the park’s marketing efforts (slabbert 2002); long-term growth can be secured by understanding each tourist market as an individual group of tourists with their distinct cultural make-up (reisinger & turner 1998); koedoe 47/1 (2004) 2 issn 0075-6458 saayman & slabbert.qxd 2004/04/15 09:08 page 2 better service and facilities can be delivered (saayman 2001). it is therefore important for the knp to profile the target market accurately and to understand its diversity. the profiling of a tourist typically involves four components, namely demographic, geographic, psychographic and socio-economic characteristics (saayman 2001 & kotler et al. 1999). based on the above, the purpose of this paper is to determine the profile of the typical tourist visiting the kruger national park. this information will help park management and marketers in the formulation of marketing policies and development strategies in order to maintain a competitive advantage. methods the methodology used in this research was twofold, namely a literature study and two surveys. the literature study focused on similar research as well as marketing literature in order to compile a questionnaire. research by kotler et al. (1999) and saayman (2001) gave insight into what aspects are relevant for the development of a profile of tourists to the park. the surveys were conducted in the knp by means of questionnaires completed by field-workers whilst interviewing tourists. the 2001 survey was completed during the first two weeks of may and the 2002 survey was completed during the first two weeks of july. the reason for this was because the months of may to july are rated as shoulder to high season and it is also the best time of the year to view game in this park. a random sampling method was used to determine the number of tourists who should form part of the survey. interviews with 220 people were conducted in 2001. camps included in the study were: berg en dal (75), satara (68), shingwedzi (34) and olifants (40). a total of 323 questionnaires were completed in 2002, viz., in berg en dal (62), olifants (93), shingwedzi (81) and satara (87) (fig. 1). the number of participants was determined by the availability of tourists and bed capacity. a structured questionnaire was used. the questionnaire dealt with demographic, socio-economic, geographic and psychographic information. respondents had to rate aspects on a five-point likert scale, where: 1 = less important, 5 = very important. issn 0075-6458 3 koedoe 47/1 (2004) fig. 1. map of the kruger national park. saayman & slabbert.qxd 2004/04/15 09:08 page 3 koedoe 47/1 (2004) 4 issn 0075-6458 table 2 socio-economic profile 2001 2002 annual income 24 % r 50 001 r 100 000 16 % r100 001 r150 000 13 % r100 001 r 150 000 15 % r50 001 r 100 000 12 % r150 001 r 200 000 education 35 % diploma/degree 45 % diploma/degree 31 % matric/equivalent 18 % post graduate 18 % professional 20 % matric mode of transport 36 % sedan 30 % sedan 18 % 4 x 4 20 % 4 x 4 15 % leisure vehicle 18 % kombi expenditure r 3500 per travelling r 4440 per travelling group per trip group per trip table 3 geographic profile 2001 2002 province 39% gauteng 42% gauteng 11% northern province 20% western cape 8% free state country of residence 11% other countries 22% usa 22% australia 19% uk table 1 demographic profile 2001 2002 gender 64 % male 66 % male 36 % female 34 % female language 54 % afrikaans 68 % afrikaans 40 % english 26 % english 5 % xhosa 4.7 % other languages marital status 78 % married 85 % married 11 % unmarried 9 % not married 6 % living together 3 % living together size of travelling group 68 % 1 to 2 47 % 3 to 4 people 24 % 3 to 4 30 % 5 to 10 people 8 % 5 or more 18 % 1 to 2 people occupation 27 % professional 43 % professional 22 % other 14 % managerial 17 % managerial 10 % other saayman & slabbert.qxd 2004/04/15 09:08 page 4 issn 0075-6458 5 koedoe 47/1 (2004) for the purpose of this article, certain aspects of the market segmentation model most frequently used in tourism research was applied (saayman 2001; kotler et al. 1999; proctor 1996; pender 1999; nickels & wood 1997). this segmentation model is used in profiling a specific market according to: demographic information, e.g., age, gender, income, marital status, education and nationality; socio-economic information, for example spending patterns; psychographic information, for example reasons for attending and most effective marketing tools; geographic information, e.g., where tourists come from. results from the surveys the following results were obtained. only the three highest percentages in each category are indicated in the results, therefore the total of some of the percentages will not equal 100 %. demographic profile from table 1 it is clear that in both years the respondents are mostly married, afrikaansspeaking tourists who travel in groups of approximately three people and practise a professional occupation. socio-economic profile table 2 shows that the respondents’ annual income varies between r 50 001 and r 150 000. most of the tourists are well-educated and this supports the notion that they are in professional occupations. their most common mode of transport is sedan vehicles, and the second-most common mode of transport is 4 x 4 vehicles. the average expenditure per travelling group in 2002 has increased by 26.8 % compared to 2001. geographic profile tourists to this park are mainly south africans who reside in the gauteng province. however, the 2002 survey indicated an increase in tourists from the limpopo and western cape provinces. this could be due to the school holidays in july, compared to may which is not a school holiday month. in july, international tourists were mostly from the usa (22 %), australia (22 %) and the uk (19 %). psychographic profile more than 50 % of the tourists in both surveys (table 4) indicated that they had visited one of the national parks on average three times per year. this is an indication that most of the respondents are loyal. from a parks management perspective, it is necessary to maintain a high quality of service to persuade these tourists to return annually. the reasons for visiting this park are: to view the variety of animal life; to relax; and to break away from their normal routine. the decision to travel to knp was made at least three weeks in advance and the respondents stayed between four and seven days. from table 5 it may be concluded that family and friends are an important source of information for tourists. word of mouth was followed by magazines, specifically the getaway, caravan and outdoor life, as well as outthere. the newspapers that are read most by respondents are the rapport and beeld. this correlates with the fact that these tourists are mainly afrikaans-speaking. television programmes most enjoyed by respondents are news and current affairs, as well as documentaries. profile of the tourist in order to make sense of the information obtained above, plog (1976) recommended the personification of target groups, so that dull statistics are transposed into a ‘living’ profile to which economists and marketers can respond more easily. using this approach introduced by the plog research group, the typical tourist to the kruger national park is personified as chris and penny steyn. chris and penny steyn are living in pretoria, in gauteng province in the republic of south africa. they are afrikaans-speaking and have a son, chris junior. chris (senior) is saayman & slabbert.qxd 2004/04/15 09:08 page 5 koedoe 47/1 (2004) 6 issn 0075-6458 table 5 media usage 2001 2002 came to hear of the park 51 % family and friends 64 % family and friends 24 % magazines 27 % other 12 % sanparks 5 % sa tourism magazines 54 % getaway 66 % getaway 44 % caravan and outdoor life 68 % outthere 36 % timbila 60 % caravan and outdoor life newspapers 40 % beeld 50 % rapport 36 % rapport radio and television programmes 76 % news and current affairs 69 % documentaries 76 % documentaries 66 % news 58 % movies 56 % sports table 4 personality type 2001 2002 personality type preferred leisure activities 88 % game viewing 90 % game viewing 78 % travel 67 % travel 72 % reading 20 % movies buying behaviour frequency of visits to national parks (per year) 51 % 2 5 times 58 % 2 5 times 20 % once 27 % once 14 % 5 10 times 11 % 5 10 times preference of the park 42 % variety of wildlife 59 % variety of wildlife 22 % beautiful nature 15 % wilderness experience 18 % wilderness experience 9 % other reason for visiting 82 % to relax 61 % to relax 78 % to get away from my 49 % to get away from my regular routine regular routine 66 % to learn about animals 35 % for the benefit of my children in general length of stay 35 % 4 7 days 34 % 7 14 days 26 % 7 14 days 33 % 4 7 days 16 % 2 4 days 24 % 2 4 days private game reserves 64 % no 63 % no 36 % yes 37 % yes booking arrangements 66 % 21+ days 84 % 21+days 19 % 1 7 days 8 % 1 7 days 8 % 15 21 days 4 % 15 21 days saayman & slabbert.qxd 2004/04/15 09:08 page 6 graduated, drives a sedan and works as a manager for a company in pretoria where he earns approximately r150 000-00 per annum. the family loves to visit the kruger national park at least once a year and stays for five days. they heard about the park from their friends and family and regularly see articles in getaway as well as in documentaries on television. chris and penny enjoy watching game as well travelling. they visit the kruger national park to relax and enjoy themselves. they enjoy the kruger national park because of the variety of game. because chris and penny prefer and love this park, they do not visit privately owned game reserves and remain brand loyal. conclusions and recommendations the aim of this paper was to determine the profile of a typical tourist visiting the kruger national park. from this research the following conclusions can be drawn: there is not a notable difference between the results, therefore a change from the months may to july, of the surveys done in 2001 and 2002. what does change, however, is that there are more foreign tourists in this park in july than in may. this might be explained by the fact that it is then the european summer holidays; a specific market segment has been determined, portrayed by chris and penny, which can streamline all marketing efforts and makes it easy to target this particular market; tourists visiting the knp are brand loyal and a quality service should sustain this market well into the future. this creates a situation where marketers can identify other potential markets to focus on them; needs of current tourists are basically the same, which makes it easy for the park to gain a larger part of the market. based on the conclusions the following recommendations can be made: from a marketing point of view, park management knows who the target market is, especially during the months of may to july. the marketing message for the international market should therefore focus on the fact that may to july is the best game viewing months of the year. if the policy is to focus primarily on the domestic market during these months, then the status quo should be maintained. from a management policy point of view, an analysis of certain aspects of the profile, e.g., spending, can give important information on how to create a situation where tourists can spend more money. one way of addressing the latter is by the development of more recreational facilities for children. this will not only lead to more spending in the park but also to investing in a future market for this and other parks. other parks can take note of these results and implement similar research in order to compare results. acknowledgements the authers would like to acknowledge dr suria ellis of statistical services for the statistical analysis. references aaker, d.a. 1998. strategic market management. new york: wiley. andereck, k.l. & l.l. caldwell. 1994. variable selection in tourism market segmentation models. journal of travel research 33: 40-46. dennis, n. 2000. kruger: images of an african park. cape town: struik. doole, i. & r. lowe. 2001. international marketing strategy. london: thomson learning. hassan, s.s. 2000. determinants of market competitiveness in an environmentally sustainable tourism industry. journal of travel research 38: 239-245. kinnear, t.c., k.l. bernard & k.a. krenter. 1995. principles of marketing. new york, ny: harpercollins. kotler, p., j. bowen & j. makens. 1999. marketing for hospitality and tourism. upper saddle river, ny: prentice hall. lamb, c.w., j.f. hair & c. mcdaniel. 2002. marketing. australia: thomson learning. laws, e. 1997. managing package tourism. london: international thomson business press. issn 0075-6458 7 koedoe 47/1 (2004) saayman & slabbert.qxd 2004/04/15 09:08 page 7 mcdonald, m. & i. dunbar. 1995. market segmentation: a step-by-step approach to creating profitable market segments. london: macmillan. nickels, w. g. & m.b. wood. 1997. marketing: relationships, quality, value. new york: worth. pender, l. 1999. marketing management for travel and tourism. cheltenham: stanley thomas. plog, s. 1976. increasing your sales to new and existing markets. florida: plog research, inc. pride, w.m. & o.c. ferrell. 1987. marketing. boston: houghton muffin company. pritchard, m.p. 1998. responses to destination advertising: differentiating inquirers to a short, getaway vacation campaign. journal of travel & tourism marketing 7: 31-68. proctor, t. 1996. marketing management: integrating theory and practice. london: international thomson business press. reisinger, y. & l. turner. 1998. cross differences in tourism: a strategy for tourism marketers. journal of travel & tourism marketing 7: 79106. saayman, m. 2001. tourism marketing in south africa. potchefstroom: institute for tourism and leisure studies. sat (south african tourism). 2002. south african tourism: business plan & budget version. johannesburg: south african tourism. semenik, r.j. 2000. promotion and integrated marketing communications. canada: thomson learning. slabbert, e. 2002. key success factors of tourist market segmentation. ma dissertation. potchefstroomuniversity for christian higher education. strydom, j.w., m.c. cant & c.j. jooste. 2000. marketing management. cape town: juta. koedoe 47/1 (2004) 8 issn 0075-6458 saayman & slabbert.qxd 2004/04/15 09:08 page 8 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 filelist convert a pdf file! page 15 page 16 page 17 page 18 page 19 page 20 page 21 page 22 page 23 page 24 page 25 page 26 page 27 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 turner.qxd the micromorphology of the african buffalo louse haematopinus bufali as observed under the scanning electron microscope m.l. turner, c. labuschagne and e. d. green turner, m.l., c. labuschagne and e. d. green. 2004. the micromorphology of the african buffalo louse haematopinus bufali as observed under the scanning electron microscope. koedoe 47(2): 83-90. pretoria. issn 0075-6458. an extensive literature search revealed that no scanning electron microscopical investigation has, to date, been performed on haematopinus bufali, a sucking louse, host specific to the african buffalo (syncerus caffer). live lice were collected from a diseased african buffalo in the rietvlei nature reserve. the specimens were fixed in 70 % ethanol. graded alcohol was used to achieve dehydration of the lice. the specimens were sonicated. the lice were dried at the critical point of medical wet carbon dioxide, mounted and fixed onto stubs. the lice were then coated with gold and viewed in a leica stereoscan 420 sem at 6–12 kv. micrographs were recorded as tagged information files (*.tif) and later printed onto hewlett packard premium glossy photographic paper with a hewlett packard 1120c professional series printer. the sem investigation revealed several micro-morphological features not previously described in light microscopical investigations of h. bufali. the head was cone shaped and the haustellum was situated on the anterior tip. setae were positioned laterally to the haustellar opening. omatidia were absent. the apex of the distal tip of the antenna showed the sensilla basoconica and sensory pegs. the inner face of the last segment revealed a pore, plate and tuft organs. the fourth segment also showed a pore organ. the pro, mesa and metathorax were completely fused. three pairs of legs with highly modified claws were present. between the distotibial pad at the base of the tarsal claw, a small structure resembling an empodium was found. two mesothoracic spiracles were noted. the abdomens of the male and female specimens differed considerably but were closely related. generally h. bufali differs distinctly from related species due to its elongated, slender, dorso-ventrally flattened and elliptically shaped body and the scalloped appearance of the elongated abdomen. m. l. turner, electron microscope unit, medical university of southern africa, 0204 republic of south africa,; c. labuschagne, rietvlei nature reserve, p.o. box 1454, pretoria, 0001 republic of south africa; e.d. green, department of anatomy, medical university of southern africa, 0204 republic of south africa). issn 0075-6458 83 koedoe 47/2 (2004) introduction the lice of mammalian and avian species are obligatory ectoparasites, mainly host specific and allocated to one of two groups, according to their feeding techniques. lice are either anopluric (sucking lice living off the body fluids of the host) or mallophagic (biting lice feeding mainly off the epidermal tissues or feathers of the host) in their feeding strategies. the superorder psocodea comprises two orders, one of which is the phthiraptera, a group with no free living representatives. it comprises obligate ectoparasites found on almost all avians and approximately a quarter of all mammals (smith 2000; soler cruz 1995). lice are further divided into four groups namely amblycera, ischnocera, rhyncophthirina (a monogeneric group found on elephant and warthog only) and anoplura. the former three are biting lice (mallophaga) whilst the latter (anoplura) are commonly known as sucking lice (smith, 2000). turner.qxd 2004/10/05 12:16 page 83 biting and sucking lice are economically important parasites and play an important role in the success of game ranches, reserves and parks all over the world. heavy lice infestations can lead to lower production yields and lower population numbers in herds of game. their feeding habits can lead to severe skin irritations of the host and the resultant symptomatic scratching aggravates these conditions even further, thus causing severe dermatitis. high infestations of anoplura may also cause harmful immune reactions (hypersensitivity and anaphylaxis), skin necrosis, localised haemorrhages, lowered weight gain or weight loss and so forth (usda 1997). high infestations may lead to anaemia, hypoproteinaemia, secondary infection (green et al. 2001), nutritional deficiencies and reduced vigour as well as secondary bacterial or fungal infections. the spread of lice infestations is promoted by increased population density, as is the case in most infectious situations. in order to design an effective control progamme in the event of high infestations, it is necessary to correctly identify the ectoparasites infesting the game. the african buffalo (syncerus caffer) is ranched on many game farms and is also protected in many game reserves in southern africa. extensive literature searches revealed that no scanning electron microscopical investigations have, to date, been performed on haematopinus bufali (pediculus bufali de geer 1778) (fig. 1), a sucking louse, host specific to the african buffalo (ledger 1980; stimie & van der merwe, 1968; usda 1997). the distribution of h. bufali coincides with that of its host and includes areas of south africa, namibia, zimbabwe, zambia, malawi and angola (stimie & van der merwe 1968). the anoplura are small, wingless and often, eyeless insects. the lice are flattened dorso-ventrally and are extremely well adapted to their blood-sucking ectoparasitic lifestyle. the head is conically shaped and adapted for piercing the skin of the host. this enables the dentate proboscis, situated in the trophic pouch (buccal funnel) of the haustellum, to koedoe 47/2 (2004) 84 issn 0075-6458 function efficiently in assisting with the blood-sucking feeding mechanism (kim, pratt & stojanovich, 1986). the mouthparts are usually retracted into the head whilst the parasite is at rest and not feeding,. eversion of the haustellum reveals dentate structures which rotate in an outward direction and in a hook-like fashion. these assist in attaching to the host and facilitates the function of the robust and well developed claws, in their role of attachment to the host. h. bufali, has to date, not been described under the scanning electron microscope (sem) although authors such as stimie & van der merwe (1968) recorded diagrammatic sketches drawn from light microscopic observations, with little or no ultra-structural detail. nevertheless, they briefly described the head, thorax and abdomen of the female and the genitals of both male and female which, in this investigation, aided in the identification of the species. it was hence concluded that the micro-morphology of h. bufali was incomplete and scanning electron microscopy was used to add to the existing information available in the literature at present. fig. 1. a sem micrograph of the dorsal view of heamatopinus bufali. turner.qxd 2004/10/05 12:16 page 84 materials and methods live lice were collected from a diseased african buffalo at the rietvlei nature reserve which is situated approximately 20 km to the southeast of pretoria. the buffalo was heavily infested and at least 30 lice were collected. the lice were found behind the ears, in the proximity of the genitalia and on the inner surfaces of the fore and hind legs. the lice were processed according to the method as described by turner et al. (2002) for the mallophagean type waterbuck louse (bovicola hilli). reproduceable and consistent results were obtained by following this procedure and it is thus re-documented in order to facilitate the investigations of researchers who wish to obtain similar high quality photo-micrographic results. the specimens were fixed in 70 % ethanol. graded alcohol (70 %, 80 %, 96 % and absolute) was used to achieve dehydration of the lice. after dehydration the specimens were sonicated for approximately 30 seconds. ultrasonic cleaning times may vary according to the toughness of the species and should be experimented with accordingly. the lice were dried at the critical point of medical wet carbon dioxide, after which they were mounted and fixed onto standard sem pin type aluminium stubs with doublesided tape. the lice were sputter-coated with gold for two minutes at 12 ma under vacuum and in the presence of argon gas. the coating procedure was repeated from different angles to reduce the effects of charging when rotating and tilting the specimen stage (holder). the specimens were viewed in a leica stereoscan 420 sem at 6–12 kv with 3–12 mm working distances. although the working distances varied due to the tilting of the specimen stage, good resolution and contrast was obtained by optimally manipulating the beam of electrons. the accelerating voltages varied proportionately to the working distances during tilting. the specimens were tilted from 0–90° and rotated from 0–360° thus covering all planes of the x, y and z axes through the specimens. the tilting and rotation was performed for the sole purpose of viewing the exterior surface of each louse in as many planes as possible. the contrast was set to 14 % whilst the brightness was kept at 32 %. an aperture of 30 µm was placed in the path of the beam to lessen the negating effects of diffraction and interference of the accelerated electrons. micrographs were recorded as tagged information files (*.tif) and submitted as such. results the lice were identified according to the distinct and pronounced morphological features as recorded by stimie & van der merwe (1968) using the light microscope. their observations included the following: female: the head was slightly longer than broad and dorso-ventrally flattened. the fore head and hind head were almost equal in length. the hind head constricts into a ‘neck’. the post-antennal angles of the ocular processes were well developed. the thorax was rectangular in shape. the sternal plate was trapezoidal in shape with an anteriorly directed posterior margin. the legs and claws were large and robust. the abdomen was elongated and flatly elliptical in shape. it had a ‘scalloped’ appearance due to the conical shape and the posterio-lateral angle of projection of the paratergites. the conical shape and angle of projection and the absence of paratergites on segments i and ii were crucial observations during identification of the lice. each tergite had a pair of median and a pair of submarginal sclerotic areas. these were absent on the first and second segments. the submarginal sclerotic areas were heavily sclerotised and formed definite plates. one or two setae were situated caudally to the paratergites. the gonophyses were concave in position and semilunar in shape. male: the male was very similar to the female but varied in length. it showed a typical v-shaped pseudopenis. the genitalia were heavily sclerotised. generally these authors remark that h. bufali differs distinctly from related species because of its elongated, slender body, and the scalloped appearance of the abdomen. the sem investigation showed several micro-morphological features not previously described in h. bufali. the following specializations and anatomical features were revealed: head: the general light microscopic model of the head as described by stimie & van der merwe (1968) was correct but incomplete issn 0075-6458 85 koedoe 47/2 (2004) turner.qxd 2004/10/05 12:16 page 85 koedoe 47/2 (2004) 86 issn 0075-6458 fig. 2. the haustellum (h), two pairs of laterally (l) situated setae and six short transverse (t) setae. fig. 4. the sensilla basoconica comprised 12 sensory pegs (p) and a long seta (s). fig. 3. the labrum (l) was folded in a membranous fashion. when viewed under the sem. this is due to the higher resolution of the sem when compared to that of a light microscope. the head was conically shaped and the haustellum was situated on the anterior tip (fig. 2). dentate processes as seen in the trophic pouch of h. phacochoeri, the warthog louse (green et al. 2000), were not noted due to the orientation of the specimen in the microscope. two pairs of setae were positioned laterally to the haustellar opening, naturally on the left and right side. the labrum was folded in a membranous fashion dorsally (fig. 3). omatidia were absent although prominent ocular processes were noted. the five segments of each of the antennae of the male seemed thicker and more robust than those of the female but this could not be verified by measurement. the apex of the distal tip of the male antenna showed that the sensilla basoconica comprised 12 sensory pegs and a long seta at the rim of the apex (fig. 4). at higher magnifications, the surfaces of the pegs seemed smooth, with no evidence of pores or other structures on their round and blunt tips. the inner face of the fifth segment revealed a complex of one pore organ (sensilla coeloconica) and two adjacent plate organs (fig. 5). a pit organ (tuft organs) was observed within the sensilla coeloconica (fig. 6). the fourth segment showed another pore organ but with no adjacent plate organs. no tuft organ protruded from this pore organ. thorax: the pro, mesa and metathorax were completely fused and the thorax was characterised dorsally by two notal pits on the prothorax and a central notal pit on the mesothorax (fig. 7). two spinous processes originated from the metathorax and extended toward the posterior. three pairs of legs with highly modified claws were present. the three pairs of legs and claws were rather similar in size. each leg had a large claw which gradually acuminated towards the most distal end. the legs showed the usual five segments namely the coxa, trochanter, femur, tibia and tarsus (fig. 8). the tibia and tarsus were highly modified to form powerful grasping organs for attachment to the turner.qxd 2004/10/05 12:17 page 86 issn 0075-6458 87 koedoe 47/2 (2004) fig. 6. the enlarged pit organ (tuft organ)(to) within the sensilla coeloconica. fig. 5. the sensilla coeloconica showing the pore organ (po) and two adjacent plate organs (pl). fig. 7. a dorsal view of the fused pro, mesa and metathorax showing two notal pits (np) on the prothorax and a central notal pit (cnp) on the mesothorax. two spinous processes (sp) protruded towards the rear. fig. 8. each leg had a large claw which gradually acuminated towards the most distal end. fig. 9. the claw with the distiotibial thumb (t), distiotibial pad (p) and empodium (e). host (fig. 9). this grasping was further aided by a distotibial thumb, as well as a distotibial pad-like structure. this seemingly acts as a shock absorbing pad. between the distotibial pad at the base of the tarsal claw, a small structure was found. this structure strongly resembles the spiked empodium found on the claws of pseudolynchia canariensis, a common ectoparasite of pigeons and wild birds (green & turner 2001) and as far as could be established, it has not been recorded in h. bufali until now (fig. 10). two mesothoracic spiracles were noted (fig. 11), each situated laterally on the left and right turner.qxd 2004/10/05 12:18 page 87 side of the thorax. the spiracles were well developed. the spiracular lumen was lined with irregular pentagonal shaped scales (fig. 12). some micrographs suggest that being pedunculated, these scales, besides filtering the air during discontinuous respiration, could purposely restrict entrance to the lumen of the spiracle in order to prevent dust particles and other debris from entering. thoracically there seemed to be no micromorphological differences in the male and female subjects. however, a single distinguishing feature which could suggest sexual dimorphism and which could be of value in future investigations, was the presence of a anterio-dorsal ridge resembling a thoracic collar in the male specimens. this collar was absent in the females. abdomen: the abdomens of the male and female specimens differed considerably but were closely related (fig. 13) (fig. 14). as recorded by stimie & van der merwe (1968), the abdomen was elongated, dorsoventrally flattened and elliptical in shape. the acute posterio-lateral angles of projection and the conical shape of the paratergites led to the scalloped ridge of the outer edge of the abdomen. no paratergites were observed on segments i and ii and this was a crucial observation in the identification of the species. a single slender marginal or postspiracular seta was situated midway between each tergite and paratergite. caudally situated and almost on the apical tip of each of the paratergites a spiracular opening was observed (fig. 15). the spiracles were also lumenally lined with the irregular pentagonal shaped and pedunculated scales closely resembled that of the thoracic spiracle. a dorsal view of the male and female abdomen showed that in the female, the scalloping was more irregular and projected radially as opposed to the caudal direction of projection in the male. the genitalia of the female were concave, semi-lunar protrudences, whilst a row of marginal setae lined a pair of flap-like claspers (fig. 16). two small tufts of setae were situated to the anterior base of the claspers (fig. 17). the anal opening was situated posterio-dorsally and just above the sexual orifice (fig. 18). the koedoe 47/2 (2004) 88 issn 0075-6458 fig. 10. an enlarged view of the empodium (e) fig. 11. a mesothoracic spiracle. fig. 12. the spiracular lumen lined with pedunculated irregular pentagonally shaped scales (s). turner.qxd 2004/10/05 12:19 page 88 male showed the typical v-shaped pseudopenis with the tip of the aedegus (penis) protruding (fig. 19). generally h. bufali distinctly differs from related species due to its elongated, slender, dorso-ventrally flattened, elliptically shaped body and scalloped appearance of the abdomen. this investigation was performed in order to assist entomologists, veterinarians and biologists in the identification of this species epidemiological studies, control as well as for the purpose of comparative studies. the study does, however, not purport to be taxoissn 0075-6458 89 koedoe 47/2 (2004) fig. 13. dorsal view of the male abdomen. fig. 14. dorsal view of the female abdomen. fig. 15. the apical tip of each of each paratergite revealed a spiracular opening (so). fig. 16. the flap-like claspers (c) of the female were lined with setae (s). fig. 17. two small tufts (t) of setae were situated to the anterior base of the claspers. turner.qxd 2004/10/05 12:20 page 89 where the idea for the article originated from and also where the first notes were made. chantelle baker, director of the medunsa electron microscope unit and riaan marais, manager of the rietvlei nature reserve. references green, e.d., m.l. turner & p.j. sebei. 2001. the functional micromorphology of the goat biting louse (bovicola caprae). proceedings of the microscopy society of southern africa 31: 62. green, e.d., f.c. clarke & m.l. turner. 2000. sem observations on the micromorphology of the warthog louse (haematopinus phacochoeri). journal of the south african veterinary association 71(2): 131138. green, e.d. & m.l. turner. 2001. the micromorphological specialisations of the claw of the lousefly. journal of the south african veterinary association 73(2): 124158. kim, k.c., h.d. pratt & c. j. stojanovich. 1986. the sucking lice of north america: an illustrated manual. london: pennsylvania state university press. ledger, j.a. 1980. the arthropod parasites of vertebrates in africa south of the sahara. volume 4: phthiraptera (insecta). johannesburg: south african institute for medical research. (publications of the south african institute for medical research; no. 56: 196-296. ). smith, v.s. 2000. avian louse phylogeny (phthiraptera: ischnocera): a cladystic study based on morphology. phd thesis, university of glasgow. soler cruz, m.d. 1995. antennal sense organs of phthiraptera (insecta). scanning electron microscopy of several species of anoplura. micron 26(1): 7-14. stimie, m. & s. van der merwe. 1968. a revision of the genus haematopinus leach (phtiraptera: anoplura). zoologischer anzeiger 180: 182220. turner, m.l., c. baker & r. marais. 2002. a scanning electron microscope investigation of the waterbuck louse bovicola hilli found at the reitvlei nature reserve. koedoe 45(1): 59-63. usda, 1997. chewing and sucking lice as parasites of mammals and birds. washington dc: united states department of agriculture. (technical bulletin; no. 1849.) koedoe 47/2 (2004) 90 issn 0075-6458 fig. 18. the anal opening (a) above the sexual orifice (s). fig. 19. the male showed the typical v-shaped pseudopenis (a). nomic in nature but rather to serve as a guide to further studies on h. bufali. acknowledgements the chief author would like to express his gratitude to the friends of rietvlei for kindly allowing me to use their private accommodation facility, as this is turner.qxd 2004/10/05 12:20 page 90 carruthers.qxd mapungubwe: an historical and contemporary analysis of a world heritage cultural landscape jane carruthers carruthers, jane. 2006. mapungubwe: an historical and contemporary analysis of a world heritage cultural landscape. koedoe 49(1): 1–13. pretoria. issn 0075-6458. the mapungubwe world heritage cultural landscape, situated on the farm greefswald at the junction of the limpopo and shashi rivers on the border of south africa, zimbabwe and botswana, falls under the management of south african national parks. this article presents a close examination of contemporary and historical issues around the inscription of the site and argues that the value of places is not self-evident but that significance is culturally constructed. this, as is explained here in respect of mapungubwe, has changed over time and has been imposed by the concerns, aspirations and values of society at a specific time and within a specific context. basic facts about mapungubwe are provided and its importance as a contemporary economic and cultural driver is outlined. a brief history of mapungubwe’s inclusion in a national park from 1947 to 1949 is presented, as is the archaeological science that it spawned. the effect that the inscription of mapungubwe may have on elevating south africa’s international profile and on african national pride is described. key words: mapungubwe, world heritage, cultural landscape, national park, archaeology, history. jane carruthers department of history, university of south africa, p.o. box 392, 0003 unisa (carruej@unisa.ac.za). issn 0075-6458 1 koedoe 49/1 (2006) introduction the inscription of a world heritage site is generally greeted with considerable publicity and expressions of national pride and selfcongratulation. while this may have palled a little for some countries that have been involved in the process since the inception of the world heritage convention in 1972, the excitement is still very real in south africa. having been barred for years from many unesco projects on account of apartheid, in 1997 the south african government was able to ratify the world heritage convention and thus became eligible to nominate sites for the list. the legislation governing south african world heritage sites is the world heritage convention act no. 49 of 1999. no site is ipso facto culturally or environmentally valuable. ‘a place is neither automatic nor self-evident. places have to be made both imaginatively and materially’ (hofmeyr 1989: 263). the aim of this article is to interrogate against the grain some of these imaginative and material values that, over time and within context, have together constructed the ‘cultural landscape’ of mapungubwe, a world heritage site in northern limpopo province, south africa. the trajectory from scientific discovery to world heritage value is contextual and, i argue, explains much about changing linkages between south african society and the natural and cultural environment. the twentieth century history of mapungubwe requires remembering in all its complexity. too often heritage sites present sanitised versions of the past because the ease of an oversimplified story is so alluring. the mapungubwe cultural landscape, situated in savanna country at the point at which south africa, zimbabwe and botswana converge at the confluence of the limpopo and shashi rivers, was added to the list of world heritage sites in july 2003 at a meeting held carruthers.qxd 2006/04/17 03:08 pm page 1 at unesco headquarters in paris. mapungubwe itself is a high (30 m) isolated sandstone outcrop, some 323x78 m in extent that overlooks the confluence. its geographical position is significant because some seven hundred years ago mapungubwe straddled the trade routes both to the indian ocean and into the interior. the underlying importance of mapungubwe, which was abandoned after 400 years of settlement in 1290 ad, is that it was for a period the largest ‘kingdom’—to use a eurocentric term—in the sub-continent. unlike many other precolonial sites, it is frozen in time, its remains are almost untouched and the whole place (there are other sites in the vicinity) bears witness to the development and altering landscape of complex social and political structures. the world heritage committee defines cultural and natural criteria to which nominated sites must conform in order to be eligible for inscription onto the world list. cultural landscapes, however, are a later inclusion into world heritage. in october 1992, after years of discussion, the operational guidelines of the convention were revised to include this new category that represented ‘the combined works of nature and man’ that were of ‘outstanding universal value’ as specified in article 1 of the world heritage convention. as far as mapungubwe is concerned, four of the five possible cultural criteria apply, viz., criteria (ii), (iii), (iv) and (v). it also meets article 39 of the operational guidelines of the world heritage convention, falling into the category of an ‘organically evolved landscape’ and the sub-categories of (ii) ‘a relict (or fossil) landscape in which an evolutionary process came to an end at some time in the past, either abruptly or over a period. its significant distinguishing features are, however, still visible in material form’ and (iii) an ‘associative cultural landscape … by virtue of the powerful religious, artistic and cultural associations of the natural elements of the landscape rather than material cultural evidence, which may be insignificant or even absent’, as specified in paragraphs 35 to 42 of the operational guidelines. mapungubwe was south africa’s first cultural landscape and its fifth world heritage site of which (at march 2006) there are seven. discussion mapungubwe: basic facts there are more than four hundred documented archaeological sites in the vicinity of mapungubwe. during the period 900– 1300 ad there were three dense settlements around mapungubwe and each of these has yielded considerable information about the economy and society of this region. the first major centre was on the farm schroda. this zhizo settlement (900–1020 ad) was followed by a site that archaeologists refer to as k2, an extensive midden deposit of the leopard’s kopje stylistic cluster which is quite separate from the zhizo ceramic tradition (1020–1220 ad) and not a continuation of it (maggs 2000:18). mapungubwe, the hill about a kilometre to the northeast of k2, was inhabited between 1220 and 1290 ad and it is here that the first consequences of wealth accumulation become evident. this inequitable distribution of wealth resulted in class distinction and disparity in access to resources, together with the physical separation of commoners from the ruling class or ‘sacred leadership’, the latter living on the top of the hill, the commoners below and serving their interests. when mapungubwe was suddenly abandoned, the result of a combination of factors including the onset of the ‘little ice age’ and perhaps a strong el niño (o’connor & kiker 2004: 49–66), the centre of regional power shifted to great zimbabwe between 1290 and 1450 ad and then to khami (rozwi) from 1450 to 1820 ad, both of which were inscribed on the list of world heritage sites in 1986. mapungubwe: a contemporary ecomonic and cultural driver at mapungubwe, there is no ongoing relationship between people and place. no modern community occupies the site or can lay koedoe 49/1 (2006) 2 issn 0075-6458 carruthers.qxd 2006/04/17 03:08 pm page 2 claim to an organic association with it. no oral evidence survives about the rise or abandonment of the sites during the period 900 to 1300 ad. mapungubwe celebrates an african, but nonetheless an alien, culture for modern black and white south africans alike. it is therefore, and unusually, a spiritually and culturally uncontested landscape, in this regard not unlike another south african world heritage site, the far older hominid fossil deposit at the cradle of humankind near johannesburg. mapungubwe’s symbolism can be appropriated by a number of groups and interested parties and the fact that it can be shared by many stakeholders means that it can serve as an exemplar landscape for south african reconciliation and nationbuilding. it can also be utilised for presentday political and economic outcomes and serve a number of contemporary agendas quite unrelated to its original function. the most urgent of these agendas is economic development. mapungubwe holds the promise of considerable practical benefit for expanding the tourist industry and the accompanying developmental infrastructure into the limpopo valley, one of the more remote parts of south africa. a number of tourist amenities are planned and interpretive visitor centres are being built inside the protected area that has recently been renamed the mapungubwe national park. a stairway enables visitors to climb mapungubwe to see the ‘royal’ quarters and to enjoy what is a spectacular view of three countries. there is a shop and view site at the confluence of the limpopo and shashi rivers, and a canopylevel boardwalk in the riverine trees along the limpopo provides views and opportunities for birding and botanizing (norton 2000; sivest 2002; deat 2002). david lowenthal believes that it is inherent in modern heritage sites to ‘become major sources of employment and revenue alike. eco-tourism and cultural travel are increasingly conjoined’ (lowenthal 2005: 82). among the legal requirements for south african world heritage sites is the obligation that they encourage investment and job creation and promote the development of culturally, environmentally and economically sustainable projects, while facilitating the empowerment and advancement of historically disadvantaged persons. world heritage status thus encourages the commodification of heritage but it also feeds into national pride and cohesion around ‘special places’ (marschall 2005: 103–122). the status of mapungubwe as being of world value is useful to south africa because the resources of limpopo province are extremely limited and it will need substantial marketing and education in order to become attractive to the average international tourist or even to south africans. first, the site cannot be understood without substantial interpretation. second, getting to mapungubwe takes effort, as it is a long drive from any metropolitan area, the climate is tropical and often extremely hot and malaria is a threat. substantial investment will be required in order to promote and publicize the destination (norton 2000; sivest 2002; deat 2002). the mapungubwe cultural landscape specifically elevates the profile of limpopo province and is already being actively used to give that province a distinct and marketable identity. there is a ‘mapungubwe tourism initiative’ and a prominent limpopo provincial leader, romokone moloto, has asserted that ‘limpopo is the home of mapungubwe, the home of civilization’ http://www.anc.org.za/limpopo/ anclimpopo.html 8 july 2005). an annual mapungubwe arts festival has been launched in polokwane and politicians declared the inaugural event ‘a resounding success that showcased many of limpopo’s talents and succeeded in attracting the best artists from all over south africa and the continent. this festival has clearly put limpopo [province] on the arts and cultural map of our continent’ (www.limpopo.gov.za/ dynamic/news/view/speech1.asp?speechid =2 18 july 2005) to have a world heritage site that can facilitate these schemes, and attribute them to mapungubwe, is politically and ideologicalissn 0075-6458 3 koedoe 49/1 (2006) carruthers.qxd 2006/04/17 03:08 pm page 3 ly useful in promoting the image of limpopo province. mapungubwe: repositioning south africa within africa the south african government is anxious to project itself as a stable modern developing nation, worthy of global respect. south africa plays the leading role in the african union (au) and also devised and heads up the new partnership for african development (nepad), a continental economic initiative that stands for good governance and neo-liberal economic management. in furthering this image of south african political and economic modernity and global outreach, mapungubwe is an appropriate symbol because it salutes an early modern, technologically advanced state and economy that existed in the region long before the era of colonization. in common with modern south africa, therefore, mapungubwe had a commercial system with ramifications worldwide. the values encapsulated in this cultural landscape are modern, capitalistic and international. while this jargon might seem strange to use for a site that is precolonial, integral to the agenda of the au and nepad is the intellectual component of the african renaissance. ‘the term “african renaissance” carries great resonance, but for historians and archaeologists the question will inevitably arise: what was the original african “golden age” that will inspire the renaissance—the rebirth of society and culture—in the new millennium? the european renaissance, emerging out of the “dark” middle ages, invoked as its vision the “golden age” drawn from the classical cultures of greece and rome. what is it that southern africa can call upon in the postcolonial era to serve as an appropriate model from the past?’ the simple answer, according to archaeologist tim maggs, is mapungubwe (maggs 2000:4). this is reflected also in the symbolism projected by the order of mapungubwe, a national decoration that recognizes excellence in science and creativity. of this, mapungubwe is an indigenous example. its claim to technological brilliance is the indigenous production of wrought gold, ironically an industry generally much maligned in south african history because its demand for cheap unskilled labour meant a level of coercion that played the major role in the disintegration of traditional african community structures and the consolidation of racial segregation. in may 2005 the south african mint launched a new gold coin (a hippopotamus) in the ‘natura’, ‘giants of africa’ series. symbolically, this was done near mapungubwe on a property belonging to de beers, the company that operates venetia, an opencast diamond mine in the area. once vilified by anti-apartheid activists, de beers has become a partner with the government in the mapungubwe enterprise because the lucrative mine is critical for its contribution to limpopo province’s economy and employs around 760 people (http://www.miningweekly.co.za/min/sector/diamonds/?show=6 9276 creamer’s media mining weekly 20 july 2005). modern south africa is replete with paradoxes and some of them take form around mapungubwe. before coming to power in 1994 the policies of the african national congress, the congress of south african trade unions and the south african communist party were based on a vision of a classless society, entrenching workers’ rights and instituting an anti-capitalist programme of nationalisation. some rhetoric of this kind remains despite the introduction of gear (growth, employment and redistribution) and government policy certainly aims to try to reduce the gap between rich and poor. but as explained, technology, mining, international trade, environmental and human exploitation and capitalist accumulation are integral to the cultural landscape that is world heritage mapungubwe. there is some irony, therefore, in the fact that mapungubwe is celebrated as an example of early, indigenous evidence of class distinction in southern africa. this class division did not come about because one intruding communikoedoe 49/1 (2006) 4 issn 0075-6458 carruthers.qxd 2006/04/17 03:08 pm page 4 ty came to dominate another, it was an organic outgrowth of mapungubwe society emanating from the inequitable control over resources by the few who consolidated and expanded their power over others (huffman 2000: 20–27). this cultural landscape therefore celebrates the emergence of a society with a hierarchy based on wealth and class and moreover, one that erased or assimilated the culture of another. mapungubwe: cultural imperialism mapungubwe does not commemorate a firstnation culture or one with an integral attachment to the landscape as a source of spirituality such as the san/bushman hunter-gatherers of the ukhahlamba-drakensberg, another south african world heritage site —a mixed site combining natural and cultural criteria. mapungubwe is an example of one culture suppressing and supplanting another: farmers squeezing out herders and foragers. in contrast to mapungubwe’s hierarchical and class-based society and the emphasis on material accumulation, the small, kin-based san communities of the area were egalitarian and bonds between people were predicated on a strong sharing ethic and on a spiritual life characterised by shamans and trance expressed in rock paintings and engravings (lewis-williams 1981; 1989; 2002). prior to the rise of mapungubwe and its associated sites, the limpopo area was inhabited by forager groups whose legacy is reflected in the rich rock art of the area. another economy, that of the herders who preceded the farmers, traders and miners of the mapungubwe ‘golden age’, also left its mark. the complicated relations among these groups in the area have been explored by simon hall and ben smith (hall & smith 2000) and karim sadr (sadr 2005). the evidence from the archaeological record is that the accomplishments of the society that is celebrated in the mapungubwe cultural landscape had a range of negative implications for foragers. despite farmers expropriating rock shelters ‘by overwriting, adding to and subtracting from, and recycling huntergatherer deposits and images by imposing their own set of marks … [farmers] assimilated their powerful places at a general level but, at the same time, expunged and rewrote the meaning and power of place in their own terms’. foragers were ‘displaced, out-competed and marginalised as the density of farmer settlements and their social and economic complexity increased’ (hall & smith 2000:30–31). by the time that mapungubwe collapsed in around 1300 ad conventional forager identity had been erased from the region. after mapungubwe was suddenly abandoned, its nine thousand inhabitants scattered and mingled with other iron age cultures. thereafter, changing climatic conditions left the limpopo valley sparsely populated but the region continued to be a corridor for human movement. groups such as the tlokwa and birwa are known to have passed back and forth and over time, fragments of these communities settled in the environs of mapungubwe. the colonial frontier in this region opened with a trickle of frontiersmen. first was the arrival of the buys people from the eastern cape in the 1820s, and they soon became embroiled in local venda contest for power. once the voortrekkers arrived in the late 1830s the balance of power shifted again. the northern boer zoutpansberg republic, with its capital at schoemansdal, was dominated by elephant hunting and the export of many tons of ivory each year. in due course, settler power consolidated in the area in the late nineteenth century. after the south african war (1899–1902) a series of generous land settlement schemes began, but because of the rugged nature of the country and its deleterious climate most farms were owned by absentee landlords or for speculative purposes by mining and land companies. in the 1930s leo fouché was able to describe the region as ‘perhaps the wildest and most desolate in the transvaal. the farms for the most part were unoccupied and only used for a few weeks shooting in the winter’ (fouché 1937: 1). eventually the extension of the railway line to musina and the burgeoning copper industry there (also a issn 0075-6458 5 koedoe 49/1 (2006) carruthers.qxd 2006/04/17 03:08 pm page 5 precolonial mining site) brought a degree of prosperity to the district but the drought and depression of the 1930s shattered the stabilising economy. mapungubwe: a colonial adventure story the rigid and bureaucratic criteria of the nomination procedures for world heritage status are unable to incorporate fully the complexity of the layers of cultural interest around mapungubwe. one of the narratives that is omitted concerns early european perceptions of africa and africans that mythologized the continent as replete with superstitious and barbaric practices, peopled by people who were backward or primitive. until the 1920s there was a general reluctance among europeans to acknowledge that the complex walled sites such as great zimbabwe and others between the limpopo and the zambezi rivers might have been constructed by africans. phoenicians, romans, hebrews, dravidian indians and other outsiders were all credited from time to time with their construction. only with british archaeologist gertrude caton-thompson’s work on zimbabwe in the late 1920s were indigenous africans eventually rightly acknowledged as the creators of these marvellous works (caton-thompson1931; 1939). as explained, in the 1920s the limpopo valley on the south african side consisted of private land, farms each of about 3 000ha that had been sold on extremely favourable terms by the state. the van graan family were settlers and young j.c.o. van graan was a student at the university of pretoria. at home on vacation in 1932 van graan was hunting on a neighbouring farm. the day was hot and the young man was thirsty and went looking for water. at a nearby african homestead he was offered water in an interesting ceramic container. being intrigued by the unusual characteristics of the bowl, van graan offered to buy it. the owner refused to part with it but did, however, divulge that it had come from a ‘sacred hill’ not far way (fouché, 1937: 1–10). his interest piqued, van graan later returned with his father, e.s.j. van graan, and three other local men intent on solving the matter of the interesting container. in trespassing on the farm greefswald, the van graan party commandeered an african, named only ‘mowena’ in the record, to point out mapungubwe, the ‘sacred hill’. it seems first by cajoling and subsequently by threatening and bribing, the man—‘literally shivering with fright’ (fouché 1937: 1) —eventually nodded in the direction of where they should look for the route that gave access to the summit. in the narrow crevice that provided a difficult climb, a large rock fig gave some purchase. the first to ascend was the young van graan. although terrified of falling into the abyss below and frightened by bats that flew into his face, he realised that a stairway had been deliberately carved into the rock. soon the group was atop mapungubwe where they found golden objects in profusion—beads, bangles—and many thousands of ceramic and glass beads and pots and potsherds. as they dug around and disturbed the surface they found more and more. eventually they realised the extent of what they had discovered and, inevitably, discussion ensued about what should be done. while the van graans felt guilty about their grave-robbing and some responsibility to alert the authorities, the others argued for keeping the treasure to themselves. in an interview done during the mid-1980s when he was then an elderly man, van graan recalled ‘i then threatened to go to the police … things were getting really ugly. my father also pleaded: we don’t have the right to these treasures … it belongs to history … south africa’s prehistory’ (van der merwe 1984: 18–37). mapungubwe: a prism of south african archaeology the catalyst for the academic study of mapungubwe was leo fouché, van graan’s history teacher, to whom the student had told his story. at the time, fouché was a founder member of what had become the university of pretoria and its professor of history. koedoe 49/1 (2006) 6 issn 0075-6458 carruthers.qxd 2006/04/17 03:08 pm page 6 unusually for this period he taught precolonial as well as settler history and he believed that archaeology (in which he was well read and relatively expert) had much to tell about the south african past. thoroughly bilingual but english-speaking rather than afrikaans, he was a great friend and admirer of jan smuts and had been smuts’s personal secretary in 1914. once fouché had visited mapungubwe he appreciated the significance of what had been discovered. fouché arranged that the van graans and their neighbours be paid out for the gold objects and these were donated to the university. the professor also located e.e. collins, the absentee owner of the farm greefswald, and negotiated for the government to purchase it. politicians of both parties of the time acted quickly. given his well known and well established scientific interests, opposition leader jan smuts, fouché’s friend, was extremely supportive. at a meeting of the south african association for the advancement of science in 1932, smuts observed that archaeological science was ‘a great field awaiting investigation in south africa’. in 1933 he personally visited mapungubwe (mason 1989:107–8) and the site was the major reason for the establishment of a south african archaeological survey (fouché 1937: 5). mapungubwe was announced internationally in the illustrated london news of 8 april 1933. the first scientific publication relating to the site was an article that year in the south african journal of science entitled ‘trade and mining in the pre-european transvaal’ by f.r. paver. the author discussed the ‘remarkable grave’ on the farm greefswald that directed attention to the ‘older native history of the transvaal’ that was extremely valuable and interesting. paver expressed a keenness to hear what the archaeologists would discover and looked forward to the time when facts would replace speculation (paver 1933: 603–611; van riet lowe 1936: 282–291). the first archaeological team under the leadership of fouché arrived at mapungubwe in april 1933. soon they were joined by local africans who helped with the manual labour. archaeological understanding of the time was limited, the experts differed in their interpretations and the underlying dynamics of the place eluded them. the concept of a southern african iron age did not then exist and most archaeological work had been done on stone age sites. fouché concluded that this was a cultural site of the local community and in his book (published 1937) he even included a photograph of a man in tattered clothes labelled as ‘petty chief tshiwana, successor and reputed descendant of the legendary chief, mapungubwe.’ after fouché left, the dig was led by rhodesian archaeologist dr neville jones and second series of excavations in the 1930s was conducted by captain guy gardner whose monumental work was published only in 1963. institutional politics played a large part in the archaeological history of mapungubwe and more detailed research in this regard would be rewarding. between his adopting the site for his university and the publication of the mapungubwe volumes, fouché had resigned from the university of pretoria in the most unpleasant of circumstances—he was hounded out for his ‘liberal’ and ‘antiafrikaner’ views—and had taken up a post at the university of the witwatersrand. during the 1930s the university of pretoria increasingly came into the grip of the cultural and political values of afrikaner nationalism, an ideology that led to full-blown apartheid in 1948. in 1932 the language policy of the university became exclusively afrikaans with an avowed mission to enter ‘the service of the afrikaner volk and … to pursue the ideals of the voortrekkers’. academic freedom became impossible and only the heroes of afrikanerdom were studied and revered. a culture of intolerance took hold and those who were not politically and ethnically correct—like fouché—were victimised (mouton 1993; university of pretoria 1960). the discipline of history became the afrikaner battleground and, not surprisingly in the paradigm of afrikaner nationalism and the ‘myth of the empty land’, mapungubwe was political anathema. because in fouché’s time there was no department of archaeoloissn 0075-6458 7 koedoe 49/1 (2006) carruthers.qxd 2006/04/17 03:08 pm page 7 gy, the professor was responsible for the establishment of an archaeological committee that operated between1933 to 1947. thereafter, however, archaeology was incorporated into a new department of anthropology (known as volkekunde in afrikaans). this is a questionable discipline in south africa because of the racism that volkekunde espoused in its ‘scientific’ support for atomizing african communities into ‘ethnic’ groups and thus relegating them to bantustans in terms of apartheid policy. in the nomination document for world heritage status, a university of pretoria professor of archaeology is quoted as saying that the slow pace of research at mapungubwe can be explained by the lack of trained archaeologists (deat 2002: 21). to some extent this is entirely correct, but there is little evidence that the university of pretoria went substantially out of its way in the early decades of working the site—this despite the fact that the excavations had deeply ‘stirred the pubic imagination’ (mason 1962: 25)— to prioritize the mapungubwe dig or to bring it within south african national consciousness. this might have been accomplished, for example, by incorporating knowledge of the site (scanty as it might have been at this time) into school textbooks and the public literature as is being done today. in this regard, one might compare the high international profile enjoyed by great zimbabwe at this time. moreover, because at the time they regarded their discipline as one remote from modern society, most archaeologists in south africa distanced themselves and their work from the public domain and the question is moot as to whether any other south african academic institution, perhaps one with a tradition of liberalism or marxism, would have done any more to publicize mapungubwe and create public value and national pride around it. shepherd argues that archaeology has a ‘history of political implication’ and has created more myths than it has dispelled, but he also acknowledges that under the political patronage of jan smuts, ‘the study of prehistory played a key role in an emergent south african national identity’ (shepherd1999; 2002: 189–209; 2003: 826–827).this was accomplished through the establishment of a bureau of archaeology (later the archaeological survey) in the department of the interior. thus archaeology in the 1930s was a directly funded branch of the civil service. this national priority, however, came to an abrupt end in 1948 when smuts was ousted as prime minister and when the ‘settler panafricanism and anglophilia … were replaced by the parochialism of afrikaner nationalism; the strange occluded twilight of prehistory—part fantasy, part brute material artifact—was eclipsed by the narratives of afrikaner sacred history’ (shepherd 2003: 833). with smuts’s patronage ended (he died in 1950) archaeological enthusiasm and enterprise wavered: the second pan-african congress in prehistory, which was to be held in south africa, did not take place there, and the archaeological survey faltered and then closed down in the early 1960s. only after 1994 did mapungubwe serve a national purpose. as explained in the introduction to this article, the place needed to be imaginatively created and its culture validated as nationally significant. apart from the archaeological politics, a comparison of fouché’s1937 work with the goodwin volume of 2000 edited by mary leslie and tim maggs is informative about disciplinary developments in archaeology that have aided reconstruction and appreciation of mapungubwe’s regional and international significance. in the period 1920 to 1950 effort was concentrated on artifact typologies, stratigraphic context, and material sequences. despite the fact that ‘it was possible—in fact, it was entirely normal—to practise african archaeology without knowing, or wanting to know, anything about african people per se’ (shepherd 2003: 838), it is fortunate that early archaeologists had been conservative. consequently their findings can now be reinterpreted as societal patterns and benefit from developments in cognitive archaeology (steyn & nienaber 2000: 112). radio-carbon dating made its useful mark in the 1950s and 1960s and from the 1970s the evidence has been re-examined (maggs 2000: 4–5). the department of koedoe 49/1 (2006) 8 issn 0075-6458 carruthers.qxd 2006/04/17 03:08 pm page 8 volkekunde at the university of pretoria and later, the department of archaeology that separated from it, never ceased working at mapungubwe. there have been a number of spurts in archaeological output. the first was from 1934 until the outbreak of the second world war. some digging was conducted in 1953–4 and again from about 1968 (voigt 1983: 5) to the mid-1980s (http://mapungbuwe.up.ac.za 13 december 2004). it is now well recognized thanks to this work that mapungubwe is part of a regional development and has a place in a far larger picture that includes other stone-walled sites of the region, for example the newly celebrated thulamela site in the kruger national park as well as great zimbabwe, khami and the others. mapungubwe: a protected area mapungubwe is presently within a newly established national park and is part of the proposed transfrontier conservation area that is being negotiated with neighbouring zimbabwe and botswana. important though these developments are, there was, however, a previous life to this protected area and it is integral to the history of mapungubwe and national park policy in the region. it forms another layer in the cultural construction of mapungubwe that needs to be recalled. i have written elsewhere in an article called ‘dongola wild life sanctuary: “psychological blunder, economic folly and political monstrosity” or “more valuable than rubies and gold”? (carruthers 1992) about the details of the genesis, brief existence and demise of the only substantial national park in south africa ever to be abolished. few people are aware that this limpopo valley landscape was formerly a national park with a strong cultural focus in addition to an ecological one. nor do they know that political pressure from local white farmers and afrikaner nationalists keen on vote-grabbing before a general election led to its abolition. it is surprising that the dongola wild life sanctuary fell from public memory so quickly because during the period 1944 to 1949 it received wide publicity in south africa and was known at the time as the ‘battle of dongola’. it led to some of the longest and most acrimonious debates in the south african parliament and the largest select committee report on record. in 1944 when discussion began, the dongola wild life sanctuary (named after a volcanoshaped mountain in the north-east corner of the farm goeree) was to be a vast area of 240 000ha, stretching from five kilometres west of musina to the confluence of the limpopo and macloutsie rivers, an area some 100km long and 36km wide at its widest point. by the time the politicians had compromised, the national park (dongola wild life sanctuary act no. 6 of 1947) had been reduced to 92 000 ha, beginning further westwards and ending not far west of the junction of the limpopo and shashi rivers. a far smaller protected area has been recreated in the mapungubwe national park and this has been at considerable expense to the state because since the 1950s irrigation in the limpopo valley has become feasible and there is an extensive agricultural industry by way of citrus, tomatoes and other crops. the dongola wild life sanctuary would have been south africa’s first national park to be founded on ecological and scientific principles. attention had been drawn to this area in the 1920s by south african botanists who were in the vanguard of the new ecological thinking that was developing in britain. the person most responsible was dr i.b. pole evans who, after an education in wales and at cambridge, took up the post of mycologist and plant pathologist in the department of agriculture in the transvaal colony in 1905. he was soon out of the laboratory and in the field. he initiated vegetation surveys, developed new fodder grasses, described plant species and involved himself in the broader issues of soil and vegetation conservation. pole evans became well acquainted and friendly with jan smuts, himself an amateur botanist of some stature, and thanks to the efforts of these men in 1918 the department of agriculture established a botanical survey of the union. issn 0075-6458 9 koedoe 49/1 (2006) carruthers.qxd 2006/04/17 03:08 pm page 9 to assist the botanical survey a number of botanical reserves were set aside in different ecosystems of the country and one of these was in the mapungubwe area. this block of nine farms was named the dongola botanical reserve. over the next decade, the reserve achieved some of its objectives. being under conservation management rather than cattle ranching, it was not long before wildlife returned to dongola. there was soon a distinct difference between environmental conditions within and outside it, and problems of overgrazing on the neighbouring properties were soon quite clear. because no work had been done on mapungubwe for a number of years, in the early 1940s the government took over the farm greefswald from the university of pretoria and added others to the dongola reserve. pole evans lobbied to have dongola elevated to national park status, not so much on account of the wildlife it contained, but on the basis of the scientific value of a natural research station, the ecological knowledge that could be gained and the proximity of mapungubwe. when smuts became prime minister again in 1939 the scheme really took off and his minister of lands, andrew conroy, became a great proponent. part of the government’s plan was to collaborate with the rhodesian government and the chartered company of bechuanaland to create an international protected area that would straddle the limpopo valley and have mapungubwe as its cultural focus, much as is the plan today. in the event, dongola was politically divisive and highly contested. conroy, pole evans and smuts were taken aback by the animosity towards the scheme from local farmers and the opposition national party which, hoping to come into power in1948, defended white property owners against expropriation. the national parks board, dominated as it was then by nationalists and by the broederbond, refused to discuss the matter. the scheme was vilified in the afrikaner press. but undeterred and undaunted, and apparently without specifically cultivating support, conroy went ahead, announcing his scheme in october 1944 in the government gazette. because private farms were to be expropriated, a select committee had to be appointed. when the matter came before parliament, the debate was vicious and personal and the dongola national park was a major election issue in 1948. the fact that the international community considered this favourably was, at that time of rising south african isolationism, a negative argument not a positive one as world heritage status is today in a different national and global environment. that black africans might have been canvassed for their views and given the national park their support was another negative to the national party. but when parliament voted in 1947 it was strictly on party lines and the national park was written into law. soon trustees were appointed, money raised, farms acquired, negotiations for the transfrontier park begun, and dongola was poised to fulfil the promise which conroy, smuts and pole evans (now warden) believed that it held. but this was not to be. so intense were emotions over dongola that when the national party won the 1948 election the national park was abolished—as voters had been promised. there was a short revival of interest in re-establishing the dongola sanctuary in the mid 1960s by the south african association for the advancement of science which sounded out the minister of agriculture, the national parks board and the university of pretoria. from all three quarters there was a negative response, memories of the previous debacle still being too fresh although three farms became a ‘reserve’ in 1967. during the 1970s and 1980s south africa was involved in a war with its neighbouring states that harboured what were regarded by the then government as ‘terrorists’ intent on destabilising the country. the army built an electric fence along the limpopo boundary and greefswald became a place for ‘rehabilitating’ conscripted gays and drug offenders. army top brass often went out hunting and koedoe 49/1 (2006) 10 issn 0075-6458 carruthers.qxd 2006/04/17 03:08 pm page 10 poached the large game of the district. they even defaced local rock shelters with graffiti (bonner & carruthers 2003: 8, 53). but ironically, the fact that an army detachment was based on greefswald brought renewed attention to the site and in the 1980s both k2 and mapungubwe hill were declared national monuments. by the 1990s the political ground had shifted once more and to most people it had become clear that the end of apartheid was in sight. it was around this time that the venetia diamond mine began operating and diamond giant de beers established its own 26 000ha venetia limpopo nature reserve, including schroda and bordering on greefswald. while recognized for its good record in wildlife conservation and management south african national parks has, however, come in for fierce criticism because of its ethos of ‘fortress conservation’ that marginalized local communities and africans in general, from the national park enterprise. sanparks can also be criticized, however, for neglecting its legal duty in terms of cultural conservation because, by law, it is obliged to value the cultural, historical and archaeological dimensions of its protected areas. the fact that mapungubwe now lies within a national park is of obvious merit to righting the situation. conclusion world heritage consultant peter fowler has emphasised that the absence of any intrusion of unsympathetic development is an essential quality of a world heritage cultural landscape (fowler 2003). despite the fact that the place was once the home to around nine thousand people with their cattle, agricultural fields and iron and gold smelting works the site is considered to be ‘authentic’ because no one lives there. (there are, however, currently a number of land restitution claims in the area and on greefswald (sanparks 2002: 33)). certainly the pre-colonial population at mapungubwe altered this environment substantially, probably removing trees as well as piling tons of soil on the summit of the hill. declaring this a world heritage cultural landscape is, arguably, celebrating environmental usage which was quite unsustainable. lowenthal argues, as i do, that culture and nature are interconnected and indivisible (lowenthal 2005: 81–92). mapungubwe is not natural, but an environment constructed to reflect the society that utilised it. the driver of the original society was the use, exploitation and maximisation of natural resources to reinforce social, political and economic values. the lesson that can be drawn from mapungubwe as a cultural landscape is the recognition that ‘much of the world’s terrestrial surface is, to a greater or lesser extent, “cultural landscape” [and] one of the most important long-term benefits of the inclusion of cultural landscapes under the world heritage convention is that it should help to promote everywhere greater awareness of landscape issues generally’ (fowler 2002). but as well as the physical landscape, the intellectual and political landscape of mapungubwe subsequent to its abandonment needs to be remembered and integrated into its heritage values. mapungubwe has faded out of the ‘history’ of the modern era twice: the first time after 1290 when it was abandoned, and the second after 1949 when it was suppressed. it has now been resurrected and transformed to meet the demands of a new society and has been given a role to play in national identity, national pride, national and transnational economics and politics. acknowledgements i thank the humanities research centre, the australian national university, canberra, and its director, professor ian donaldson, for awarding me a visiting fellowship in 2005 and with it the opportunity to benefit from the ‘cultural landscape’ programme. i am also grateful to the history program of the research school of social sciences at the australian national university for previous visits to canberra that have greatly informed my work on heritage issues. in particular i value assistance from libby robin, tom griffiths, mandy martin, guy fitzhardinge, vincent carruthers, peter norton, phil bonner, richard grove, mike smith, jane lennon, issn 0075-6458 11 koedoe 49/1 (2006) carruthers.qxd 2006/04/17 03:08 pm page 11 kirsty douglas, max bourke, john mulvaney, tom huffman, isabel mcbryde, alex mouton, greg cuthbertson and emily o’gorman, as well as the anonymous readers of this article. references bonner, p. & e.j. carruthers. 2003. the recent history of the mapungubwe area. mapungubwe cultural heritage resources survey. unpublished report commissioned by the department of environmental affairs and tourism: funded by the norwegian government. breedlove, g. 2003. a systematics for the south african cultural landscape with a view to implementation. ph. d. thesis, university of pretoria, pretoria. carruthers, j. 1992. dongola wild life sanctuary: ‘psychological blunder, economic folly and political monstrosity’ or ‘more valuable than rubies and gold’. kleio 24: 82–100. carruthers, j. 2005. wildlife science and livestock disease in south africa, 1900 to 1960. paper presented at the conference ‘science, disease and livestock economies’ at st antony’s college, oxford. caton-thompson, g. 1939. mapungubwe. antiquity 13 (51): 324–341. caton-thompson, g. 1931. the zimbabwe culture. oxford: clarendon press. deacon, j. 1993. archaeological sites as national monuments in south africa: a review of sites declared since 1936. south african historical journal 29: 118–131. department of environmental affairs and tourism (deat). 2002. mapungubwe world heritage site nomination. pretoria: government printer? fouché, l. (ed.) 1937. mapungubwe: ancient bantu civilization on the limpopo. cambridge: university press. fowler, p. 2002. world heritage cultural landscapes, 1992–2002: a review and prospect, cultural landscapes: the challenges of conservation world heritage 2002, shared legacy, common responsibility associated workshops 11–12 november 2002 ferrara—italy. http:// whc.unesco.org/venice2002. fowler, p. 2003. world heritage cultural landscapes 1992-2002, world heritage papers 6, 2003. unesco world heritage centre, paris world heritage cultural landscapes 1992-2002 http://whc.unesco.org. gardner, g.j. 1958. mapungubwe and the second volume. south african archaeological bulletin 13: 123–132. gardner, g.j. 1955. mapungubwe 1935–1940. south african archaeological bulletin 10: 73–77. gardner, g.j. 1963. mapungubwe vol. ii. pretoria: van schaik. hall, s. & b. smith. 2000. empowering places: rock shelters and ritual control in farmer-forager interactions in the northern province. pp. 30– 46. in: leslie, m. & t. maggs (eds.). african naissance: the limpopo valley 1000 years ago. cape town: the south african archaeological society. (goodwin series, vol. 8.) hofmeyr, i. 1989. turning region into narrative: english storytelling in the waterberg. in: bonner, p., i. hofmeyr, d. james & t. lodge (eds.). holding their ground: class locality and culture in 19th and 20th century south africa. johannesburg: witwatersrand university press. huffman, t. m. 2005. mapungubwe: ancient african civilisation on the limpopo. johannesburg: witwatersrand university press. huffman, t.m. 2000. mapungubwe and the origins of the zimbabwe culture. pp.14–29. in: leslie, m. & t. maggs (eds.). african naissance: the limpopo valley 1000 years ago. cape town: the south african archaeological society. (goodwin series, vol. 8.) leslie, m. & t. maggs (eds.). 2000. african naissance: the limpopo valley 1000 years ago. cape town: the south african archaeological society. (goodwin series, vol. 8.) lewis-williams, j.d. 2002. a cosmos in stone: interpreting religion and society through rock art. walnut creek: ca, altamira press. lewis-williams, j.d. 1981. believing and seeing: symbolic meanings in southern african san rock paintings. london: academic press. lewis-williams, j.d. 1989. images of power: understanding bushman rock art. johannesburg: southern book publishers. lowenthal, d. 2005. natural and cultural heritage. international journal of heritage studies 11(1): 81–92. lowenthal, d. 1997. the heritage crusade and the spoils of history. london: viking. lowenthal, d. 1985. the past is a foreign country. cambridge: cambridge university press. maggs, t. 2000. african naissance: an introduction. pp. 1–4. in: leslie, m. & t. maggs (eds.). african naissance: the limpopo valley 1 000 years ago. cape town: the south african archaeological society. (goodwin series, vol. 8.) marschall, s. 2005. making money with memories: the fusion of heritage, tourism and identity formation in south africa. historia 50(1): 103–122. mason, r. 1962. prehistory of the transvaal: a record of human activity. johannesburg: witwatersrand university press. koedoe 49/1 (2006) 12 issn 0075-6458 carruthers.qxd 2006/04/17 03:08 pm page 12 mason, r.j. 1989. south african archaeology 19221988. johannesburg: archaeological research unit university of the witwatersrand. mouton, f.a. 1993. professor leo fouché, the history department and the afrikanerisation of the university of pretoria. historia 38(1): 51–63. munjeri, d. 2002. smart partnerships: cultural landscape issues in africa, cultural landscapes: the challenges of conservation world heritage 2002. shared legacy, common responsibility associated workshops 11-12 november 2002 ferrara – italy http://whc. unesco.org/venice2002. norton, p. 2000. an overview of tourism development potential in the mapungubwe area. johannesburg: dbsa. o’connor, t.g. & g.a. kiker. 2004. collapse of the mapungubwe society: vulnerability of pastoralism to increasing aridity. climatic change 66 (1/2): 49–66. sadr, k. 2005. hunter-gatherers and herders of the kalahari during the late holocene. pp. 206–221. in: veth, p., smith, m. & p. hiscock (eds.). desert peoples: archaeological perspectives. malden: blackwell. shepherd, n. 2003. state of the discipline: science, culture and identity in south african archaeology, 1870–2003. journal of southern african studies 29 (4): 823–844. shepherd, n. 2002. the politics of archaeology in africa. annual review of anthropology 31: 189–209. sivest consortium. 2002. mapungubwe tourism initiative conceptual tourism destination plan. south african national parks. 2002. notes to the annual financial statements for the year ended 31 march 2002. steyn, m. & w. nienaber. 2000. iron age human skeletal remains from the limpopo valley and soutpansberg area. pp. 112–116. in: leslie, m. & t. maggs (eds.). african naissance: the limpopo valley 1 000 years ago. cape town: the south african archaeological society. (goodwin series, vol. 8.) university of pretoria. 1960. ad destinatum: gedenkboek van die universiteit van pretoria. vol. 1. johannesburg: voortrekkerpers. van der merwe, c. 1984. the face of south africa —mapungubwe. flying springbok june, 1984: 18–37. van riet lowe, c. 1936. mapungubwe: first report on excavations in the northern transvaal. antiquity 10(39): 282–291. van warmelo, n.j. 1942.the copper miners of musina and the early history of the zoutpansberg. pretoria: government printer. voigt, e.a. 1983. mapungbwe: an archaeological interpretation of an iron age community. pretoria: transvaal museum monograph no. 1. weinek, c. 2000. the hills are alive: a new documentary uncovers a drama of denied heritage, stolen treasures and botched excavations. mail and guardian 3–9 march: 3. world heritage convention. 1972. convention concerning the protection of the world cultural and natural heritage. paris: unesco. www.limpopo.gov.za/dynamic/news/view/speech1. asp?speechid=2. accessed 18 july 2005 http://www.anc.org.za/limpopo/anclimpopo.html accessed 18 july 2005. h t t p : / / w w w. m i n i n g t e c h n o l o g y. c o m / p r o j e c t s / de_beers/specs.html. http://www.miningweekly.co.za/min/sector/diamonds/?show=69276, creamer’s media mining weekly 20 july 2005. issn 0075-6458 13 koedoe 49/1 (2006) carruthers.qxd 2006/04/17 03:08 pm page 13 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 article information authors: frances siebert1 holger c. eckhardt2 stefan j. siebert1 affiliations: 1school of environmental sciences and development, north-west university, south africa 2scientific services, south african national parks, south africa correspondence to: frances siebert email: frances.siebert@nwu.ac.za postal address: school of environmental sciences and development, north-west university, private bag x6001, potchefstroom 2520, south africa keywords biodiversity; letaba river; mopaneveld; riparian vegetation; savanna; vegetation mapping dates: received: 27 may 2009 accepted: 24 nov. 2009 published: 21 june 2010 how to cite this article: siebert, f., eckhardt, h.c. & siebert, s.j., 2010, ‘the vegetation and floristics of the letaba exclosures, kruger national park, south africa’, koedoe 52(1), art. #777, 12 pages. doi: 10.4102/koedoe.v52i1.777 copyright notice: © 2010. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0771 (online) the vegetation and floristics of the letaba exclosures, kruger national park, south africa in this original research... open access • abstract • introduction • study area • method    • classification and mapping    • floristic analysis • results    • classification and mapping    • floristic analysis • discussion    • description of plant communities       • 1. hemizygia bracteosa – acacia exuvialis crest savanna       • 2. schmidtia pappophoroides – colophospermum mopane lowland savanna       • 3. sporobolus fimbriatus – grewia bicolor seasonally wet savanna       • 4. croton megalobotrys – panicum deustum riverine woodland • conclusion • acknowledgements • references abstract (back to top) the construction of exclosures along two of the most important rivers in the kruger national park was done to investigate how patterns of spatial and temporal heterogeneity of the riparian zone is affected by fire, flood and herbivory. to assist this research programme, vegetation surveys were conducted within exclosures along the letaba river to classify and map the vegetation of the area. the history and experimental design of the letaba exclosures are similar to that of the nkhuhlu exclosures along the sabie river, which is directly related to questions surrounding elephant management. the main difference between the nkhuhlu and letaba exclosures is local heterogeneity, since the latter lies within the mopaneveld, which is floristically and physiognomically much more homogenous than the vegetation of the southern kruger national park. nevertheless, four plant communities, eight sub-communities and six variants were recognised and mapped for the letaba exclosures. the vegetation description was done in relation to prevailing soil forms, differences in species richness, diversity and community structure, and therefore should serve as a basis for further detailed and broad-based botanical studies. vegetation mapping was done to sub-community level and, where possible, to variant level. as expected in mopaneveld vegetation, the plant communities could broadly be related to soil types, although smaller-scale variations correspond to soil moisture availability because the mopaneveld is considered ‘event-driven’, especially in the herbaceous layer. conservation implications: floristic surveying and vegetation mapping of a long-term monitoring site, such as the letaba exclosures, is seen as a baseline inventory to assist natural resource management. linking mapping units to biodiversity strengthens the understanding needed to maintain biodiversity in all its natural facets and fluxes. introduction (back to top) situated on the banks of the letaba river, in the northern half of the kruger national park (knp), the letaba exclosures were constructed in 2002. for the same reason and purpose as the nkhuhlu exclosures, namely to monitor spatial and temporal heterogeneity along the riparian zone and attempt to determine how these are affected by fire and herbivory (siebert & eckhardt 2008). also, the history and experimental design of the letaba exclosures are similar to those of the nkhuhlu exclosures (o’keefe & alard 2002), which is directly related to questions surrounding elephant management. the different treatment blocks at the letaba exclosures vary in size, with 42 ha, 51 ha and 36 ha respectively, for the full exclosure, partial exclosure and unfenced area. the main difference between the nkhuhlu and letaba exclosures is local heterogeneity, because the latter lies within the mopane bioregion (mucina & rutherford 2006), which is floristically and physiognomically much more homogenous than the southern savannas of the knp. animal concentrations are consequently relatively low, exacerbated by the fact that, approximately a decade ago, the letaba river changed from a perennial river to a seasonal river (o’keeffe & rogers 2003). the lower mean annual rainfall (approximately 400 mm), compared to the annual 560 mm at nkhuhlu, drives a different fire regime and mammal concentrations. these not only translate into less obvious differences between the different treatments at the letaba exclosures, but also in a lower rate of changes overall. in 2000, a flood caused the letaba river to rise above its macro-channel banks, but did not cause as dramatica change to the riparian vegetation as was recorded for the sabie river (parsons, mcloughlin, kotschy, rogers & rountree 2005). nevertheless, the changes brought about by this large, infrequent disturbance provide an ideal opportunity and context to study and understand the dynamics of ecosystems subjected to different management options. baseline vegetation surveys were conducted initially and repeated five years later, in 2007, to assess differences between the six treatments in terms of plant species composition and structure for both the woody and herbaceous component (o’keefe & alard 2002). the results from this survey will be published at a later stage. a detailed soil inventory was conducted by paterson and steenekamp (2003), the results of which were used to help delineate plant communities. owing to their remoteness and other logistical reasons, the letaba exclosures have not received the same degree of attention from researchers as the nkhuhlu exclosures. this has led to missed opportunities with regard to the flood legacy. the detailed vegetation classification and floristic description presented here will hopefully attract more interest from scientists, leading to research projects addressing key environmental questions. study area (back to top) the letaba exclosures are situated approximately 5 km upstream of the mingerhout dam, along the northern bank of the letaba river in the kruger national park. the exclosures consist of six treatments: • a fully fenced area (42 ha) excluding all herbivores, divided into, (1) a burn block and (2) a no-burn block • a partially fenced area (51 ha) excluding only elephant and giraffe, also divided into a burn (3) and no-burn (4) block • (5) an unfenced area protected from fire only (36 ha) • (6) a 400 m wide buffer area that is subjected to fire and herbivory. the total experimental area is 129 ha, excluding the area along the river, which forms a narrow strip inside the river bed (the so-called ‘sacrificial zone’) and the buffer zone adjacent to the three inland sides of the project area. this summer rainfall region receives approximately 400 mm per annum, as measured at letaba rest camp, while the mean daily temperature is 23.3 °c, ranging from 7.8 °c in winter, to a maximum of 34.1 °c in summer (weather bureau 1986). the most representative underlying geology is granite and makhutswi gneiss, which contains swaziland rock formations such as amphibolite and migmatite (visser, coertze & walraven 1989). according to paterson and steenekamp (2003), the letaba exclosures have a relatively simple soil distribution, with mispah being the most representative soil form along the midslopes and crests. hutton soils form a strip of deeper apedal soils along the footslopes, whereas alluvial soils of the oakleaf soil form characterises the banks of the letaba river (figure 1). the topography of the area is characterised by a gently undulating landscape without any prominent koppies. figure 1: soil map of the letaba exclosures study site (courtesy of arc-institute of soil, climate and water, paterson and steenekamp 2003) the vegetation of the letaba exclosures is classified as ‘letaba river rugged veld’ (landscape 10, gertenbach 1983). the vegetation of the drier upper savanna areas contains elements of landscape 22, the ‘combretum spp. / colophospermum mopane rugged veld’ (gertenbach 1983) and is floristically linked to the ‘colophospermum mopane – combretum apiculatum – digitaria eriantha open tree savanna’, described by van rooyen, theron and grobbelaar (1981). the riparian zone of the letaba site forms part of subtropical alluvial vegetation (unit aza7 of the inland azonal vegetation), whereas the zonal savanna area forms part of the lowveld rugged mopaneveld (svmp6) in the mopane bioregion (mucina & rutherford 2006). on a larger scale, the vegetation of the letaba exclosures forms part of the combretum apiculatum – colophospermum mopane major plant community on granite and gneiss in the cissus cornifolia – colophospermum mopane vegetation type (siebert, bredenkamp & siebert 2003). mopaneveld on granitic substrates is typically in association with combretum spp., where the latter dominates the sandy crests and colophospermum mopane the more clayey substrates of the bottomlands (du plessis 2001). of the four major plant communities identified and described by bredenkamp and van rooyen (1993), three are represented at the letaba site, of which some are typically found in the study area. species that characterise the riparian zone include woody species such as croton megalobotrys, nuxia oppositifolia, gymnosporia senegalensis, philenoptera violacea, diospyros mespiliformis, combretum microphyllum and trichilia emetica. inland, the vegetation is typically dominated by colophospermum mopane, with variation in species associations that co-exist in the woody layer. woody species, other than colophospermum mopane, that characterise the savanna zone of the letaba site, include combretum apiculatum, terminalia prunioides, dichrostachys cinerea, maerua parvifolia, cissus cornifolia, commiphora mollis, c. africana, grewia bicolor, combretum imberbe, acacia nigrescens, acacia exuvialis and rhigozum zambeziacum. grasses, such as panicum coloratum, schmidtia pappophoroides, melinis repens, bothriochloa radicans, and forbs such as crabbea velutina, ocimum americanum, phyllanthus maderaspatensis, pavonia burchellii, hibiscus micranthus and jasminum stenolobum, characterise the field layer. method (back to top) classification and mapping vegetation sampling, classification and mapping was based on the approach followed in the study of the nkhuhlu exclosures (siebert & eckhardt 2008). a total of 83 plots were identified through randomised stratification, of which 72 plots were sampled in detail using the braun-blanquet scale (mueller-dombois & ellenberg 1974). total floristics were obtained for each 20 m × 20 m plot to allow for comparison of species diversity and richness between the different plant communities. plant species names conform to those of germishuizen and meyer (2003).braun-blanquet procedures were used to refine the classification results of the twinspan (hill 1979) application using the megatab computer program (hennekens 1996). the results are presented in a phytosociological table (see online appendix). results from the classification and subsequent refinement were used to construct a 1:5000 vegetation map for the letaba exclosures (figure 2). mapping was done to the sub-community level in arcmap™ of the esri arcgis 9.1® computer package. to improve the understanding of the map in terms of the topography, the distribution of the plant communities is presented in a profile drawing (figure 3) that was constructed using corel draw suite 9©. figure 2: vegetation map of the letaba exclosures study site figure 3: profile of the vegetation (corresponding to the mapping units of figure 2) along a topographical gradient of the study area from the letaba river riverbed to the crest of the landscape floristic analysis floristic data were analysed to allow floristic comparison across mapping units and were therefore presented on variant level where appropriate. the approach for floristic comparison followed the method used for the nkhuhlu exclosures (siebert & eckhardt 2008), which entailed the use of shanon-wiener diversity indices to combine species richness and relative abundance among species (barbour, burk & pitts 1987) for comparison across mapping units. the braun-blanquet scale was converted to percentage cover and the mean cover per species within the plant community was taken as its abundance value. figure 4: profile of the vegetation (corresponding to the mapping units of figure 2) along a topographical gradient of the study area from the letaba river riverbed to the crest of the landscape results (back to top) classification and mapping first order divisions, created by the hierarchical classification of the letaba data set, separated riparian woodland from the dryland woodland communities, the latter of which are dominated by co lophospermum mopane and differentiated according to elevation and drainage. these two vegetation types represent four main communities, which include eight sub-communities and six variants after refinement:1. hemizygia bracteosa – acacia exuvialis crest savannah 1.1 xerophyta humilis – oldenlandia herbacea seepage 1.2 sclerocarya birrea – bothriochloa radicans ridge bushveld 1.2.1 senna italica variant 1.2.2 typical variant 2. schmidtia pappophoroides – colophospermum mopane lowland savanna 2.1 melhania rehmannii – enneapogon scoparius mixed bushveld 2.1.1 lannea schweinfurthii variant 2.1.2 aristida canescens variant 2.2 maerua parvifolia – agathisanthemum bojeri mopane bushveld 3. sporobolus fimbriatus – grewia bicolor seasonally wet savanna 3.1 eragrostis trichophora – sporobolus stapfianus mud pans 3.2 barleria elegans – acacia karroo streambank woodland 4. croton megalobotrys – panicum deustum riverine woodland 4.1 philenoptera violacea – jasminum fluminense riverbank woodland 4.1.1 hyphaene coriacea variant 4.1.2 nuxia oppositifolia variant 4.2 combretum imberbe – enneapogon cenchroides open woodland. the identified plant assemblages are distributed throughout the letaba exclosures study area and are well-presented in all treatment blocks at plant community level (figure 2). some communities have a limited distribution range due to small-scale disturbances and patchiness, for example the xerophyta humilis – oldenlandia herbacea seepage (1.1) and the eragrostis trichophora – sporobolus stapfianus mud pans (3.1). sub-communities are easily identifiable in the field; some can even be distinguished at variant level, for instance the lannea schweinfurthii (2.1.1) and aristida canescens (2.1.2) variants of the melhania rehmannii – enneapogon scoparius mixed bushveld, as well as the hyphaene coriacea (4.1.1) and nuxia oppositifolia (4.1.2) variants of the philenoptera violacea – jasminum fluminense riverbank woodland. vegetation mapping was therefore done up to variant level where possible, although sub-communities form the most representative units of the vegetation map (figure 2). a schematic presentation of plant community distributions – from the letaba river to the crest of the landscape (figure 3) – illustrates changes in the vegetation along the riparian catena. the riverbed and lower riverbank is characterised by plant species of the nuxia oppositifolia variant (4.1.2). on the slightly elevated riverbank, the hyphaene coriacea variant (4.1.1) can easily be recognised by the presence of prominent trees, such as hyphaene coriacea, philenoptera violacea, croton megalobotrys and combretum hereroense. the hyphaene coriacea variant is often interrupted by drier, open woodland, the combretum imberbe – enneapogon cenchroides open woodland (4.2), on the higher terraces of the letaba river. higher up into the savanna zone, the vegetation varies from a closed, almost homogenous mopane woodland (2.2), via a mixed bushveld (2.1) that is interrupted by narrow green ‘belts’ (2.1.1), to a crest bushveld (1), which includes the sclerocarya birrea – bothriochloa radicans ridge bushveld (1.2) and a unique, localised seepage (1.1) (figure 3). floristic analysis compared to the nkhuhlu exclosures, the plant richness per area of the letaba exclosures is low (figure 4), due to colophospermum mopane dominating the tree and shrub layers (o’connor 1992; du plessis 2001). the overall plant richness per area, of both the nkhuhlu and letaba exclosures, is above average for the kruger national park (figure 4).at community level, the schmidtia pappophoroides – colophospermum mopane lowland savanna (community 2) hosts the highest alpha diversity (species richness per unit area) at 32 species per plot and a total of 141 different plant species. however, the shannon-wiener diversity index, which combines both richness and equitability, revealed it has a low diversity and evenness (table 1), attributable to the almost sole dominance of colophospermum mopane (o’connor 1992; du plessis 2001). the highest alpha diversity on sub-community level was recorded for the philenoptera violacea – jasminum fluminense riverbank woodland (4.1) and the melhania rehmannii – enneapogon scoparius mixed bushveld (2.1) (figure 5). the highest floristic diversity on sub-community level was measured for the streambank woodland (3.2), with a value of 3.9 (figure 5), which can be attributed to the variation in bordering savanna plant communities, from which individuals spread into this sub-community with moist soils. floristically, the study area is dominated by taxa from the fabaceae, poaceae and malvaceae (table 2). the dominant growth forms in the study area are forbs, graminoids, trees and shrubs, although several climber species were recorded (table 3). succulents are less prominent, compared to the nkhuhlu exclosures. figure 5: analysis of species diversity, richness and cover of the eight sub-communities of the letaba exclosures: (a) shannon-wiener index of diversity and evenness, (b) total and mean species richness per unit area (i.e. per 20 m × 20 m plot) and (c) herbaceous versus woody layer cover percentages discussion (back to top) description of plant communities the letaba exclosures experimental site lies in the geological and climatically uniform landscape 10 of gertenbach (1983), namely the ‘letaba river rugged veld’. no major climatic variation occurs that could have an effect on the local differentiation of plant communities. small-scale heterogeneity of environmental factors could contribute to variations in plant assemblages. detailed descriptions of the plant communities are therefore important to explain and provide support for the delineation of each mapping unit in a heterogeneous landscape. 1. hemizygia bracteosa – acacia exuvialis crest savanna this community forms one of four plant communities within the study site and comprises approximately 30% of the total sampling area (including the buffer zone). it is spread out across the exclosure fence lines, with little difference between the three exclosure types in terms of plant composition. it extends over the crests of the savanna landscape inland of the letaba river and is associated with shallow, greyish-brown, gravelly loamy sand to sandy loam topsoil directly overlying hard rock of the mispah soil form (paterson & steenekamp 2003) (figure 1). the hemizygia bracteosa – acacia exuvialis crest savanna varies from open to moderately open shrubveld. the field layer is dominated by forbs, whereas shrubs constitute the dominant growth form in the woody layer. it is floristically characterised by species group 1 (see online appendix), which includes acacia exuvialis in the woody layer, and hemizygia bracteosa and tricliceras laceratum in the forb layer (species group 1). dominant species include the trees colophospermum mopane, terminalia prunioides and combretum apiculatum and the grass panicum coloratum (species group 14). species frequently present include the grasses bothriochloa radicans, tragus berteronianus (species group 3), schmidtia pappophororoides (species group 5) and enneapogon scoparius (species group 10), whereas evolvulus alsinoides, clerodendrum ternatum (species group 10), lantana rugosa, ceratotheca triloba and pavonia burchellii (species group 14) are commonly found in the forb layer. this community is floristically the poorest, with a total of 96 plant species recorded (table 1), which contribute to only 41% of the species richness of the letaba exclosures. diversity and evenness scores are at a moderate level for the study area (table 1). taxa of the fabaceae and poaceae (table 2) floristically dominate the crest savanna. the crest savanna contains elements of the ‘combretum spp. / colophospermum mopane landscape’ (landscape 22, gertenbach 1983) in the field layer, but not typically in the woody layer, except for common trees such as colophospermum mopane, terminalia sericea, acacia exuvialis, combretum apiculatum and acacia nigrescens. this savanna community is too distant to associate its plant assemblages with the riparian vegetation described by bredenkamp and van rooyen (1993), although it does contain elements of the colophospermum mopane – terminalia prunioides dry riparian woodland (community 1.1.2, bredenkamp & van rooyen 1993). two sub-communities were identified for the hemizygia bracteosa – acacia exuvialis crest savanna: 1.1 xerophyta humilis – oldenlandia herbacea seepage: this sub-community is not widely distributed in the letaba exclosures site, nor in the adjacent colophospermum mopane savanna surrounding the study area. it covers only a small patch (1% of the study area) on the crest of the landscape where underground water seeps through the surface, resulting in a seasonally wet area. mispah is the dominant soil form, with a stony soil surface that is often underlain by rock plates where moisture accumulates. this somewhat unique plant community is found within the buffer zone of the full exclosure, extending into the full exclosure. it is an open savanna dominated by low colophospermum mopane shrubs of approximately 3 m in height, which are interrupted by damp, open patches dominated by xerophyta humilis. woody cover percentage is the lowest in this sub-community. cover in the herbaceous layer is approximately 25%. the seepage, which is dominated by herbaceous species (figure 5), is characterised by species group 2 (see online appendix). since this sub-community is not typically found in the study area it is undersampled, represented by only one relevé: sample plot 12. the most conspicuous woody species is colophospermum mopane (species group 14) in its low shrub (± 3 m) form. graminoids such as melinis repens (species group 2), schmidtia pappophoroides (species group 5) and panicum coloratum (species group 14) occur, but with low percentage cover. prominent forbs include, other than xerophyta humilis and oldenlandia herbacea (species group 2), kohautia virgata (species group 9), evolvulus alsinoides (species group 10), tephrosia purpurea and ceratotheca triloba (species group 14). this sub-community has a species richness value of 15, which is also the mean species number per sampling plot (figure 5). 1.2 sclerocarya birrea – bothriochloa radicans ridge bushveld: this sub-community is widespread in the savanna uplands of the letaba exclosures site, especially along the crests of the landscape which predominantly occurs within the 400 m bufferzone (figure 2). soils underlying this sub-community are mainly from the mispah soil form. the vegetation is structurally characterised by an open to moderately open shrubveld. the herbaceous layer has a cover of 34% and the woody layer covers approximately 23% of the area (figure 5). this ridge bushveld is characterised by species group 3 (see online appendix) and include sclerocarya birrea in the woody layer and tragus berteronianus and bothriochloa radicans in the grass layer. the woody layer is dominated by colophospermum mopane (species group 14). prominent woody species comprise rhigozum zambesiacum (species group 10), terminalia prunioides and combretum apiculatum (species group 14). forbs include indigofera nebrowniana (species group 7), evolvulus alsinoides, dicoma tomentosa, talinum caffrum, clerodendrum ternatum (species group 10), ceratotheca triloba, hibiscus micranthus, lantana rugosa, melhania acuminata, ocimum americanum, pavonia burchellii, phyllanthus incurvus and phyllanthus maderaspatensis (species group 14). species richness is 96, whereas mean species number per plot is 30 (figure 5). the diversity for this sub-community is above average, common for plant communities found on ridges (siebert et al. 2003). evenness is average for the letaba site (figure 5). two variants were identified but do not represent different mapping units. these variants were: 1.2.1 senna italica variant: this variant is structurally classified as moderately open to open shrubveld on slightly elevated crests on shallow soils of the mispah soil form. it is less stoney than the typical variant and occurs in the open control area, the partial exclosure and in the buffer zone outside the northeastern border of the partial exclosure. characteristic species of this variant are listed in species group 4 and comprise senna italica, heliotropium ovalifolium, boerhavia diffusa, corbichonia decumbens and pterodiscus speciosus, all of which are in the herbaceous layer. hemizygia bracteosa (species group 1) is less abundant here than the typical variant. dominant species include colophospermum mopane, terminalia prunioides and panicum coloratum (species group 14), whereas combretum apiculatum (species group 14), bothriochloa radicans (species group 3) and enneapogon scoparius (species group 14) are locally prominent. 1.2.2 typical variant: this variant occurs as interrupted patches of moderately open shrubveld and is in general associated with crests on shallow soils covered by granitic pebbles, associated with the mispah soil form. it covers most of the upland savanna vegetation towards the north-eastern border of the partial exclosure, as well as outside the exclosure in the bufferzone. there are no species that typically characterise this variant, although the absence of species group 4 may serve as an indicator of this typical variant. the dominance of hemizygia bracteosa (species group 1) is characteristic. important species in the woody layer, other than colophospermum mopane, include acacia exuvialis (species group 1), sclerocarya birrea (species group 3), rhigozum zambesiacum (species group 10), terminalia prunioides, combretum apiculatum, acacia nigrescens and dichrostachys cinerea (species group 14). other species present in the herbaceous layer are grasses such as bothriochloa radicans (species group 3), schmidtia pappophoroides (species group 5), enneapogon scoparius (species group 10), panicum maximum (species group 21) and forbs such as indigofera nebrowniana (species group 7), ocimum americanum, ceratotheca triloba (species group 14) and waltheria indica (species group 21). 2. schmidtia pappophoroides – colophospermum mopane lowland savanna this lowland savanna typically belongs to the lowveld rugged mopaneveld (svmp6, mucina & rutherford 2006) and covers the largest portion of the letaba exclosures site. it is a typical mopaneveld plant community that is widespread in the knp and adjacent savanna areas. its nearest counterpart in plant communities, as have been described previously, is the colophospermum mopane – combretutum apiculatum – digitaria eriantha open tree savanna (van rooyen et al. 1981), which occurs further north of the letaba exclosures site, in the punda maria region. this typical mopaneveld community occurs on granitic soils of the mispah soil form (siebert et al. 2003). structurally, this savanna type varies from an open shrubveld to closed savanna woodland. the dominant growth form in the field layer is forbs, whereas shrubs dominate the woody layer. this community is floristically characterised by species group 5. the woody layer is dominated by colophospermum mopane and terminalia prunioides (species group 14). the grass layer is dominated by panicum coloratum (species group 14) and schmidtia pappophoroides (species group 5). the grasses enneapogon scoparius (species group 10), eragrostis superba and panicum maximum (species group 21) are prominent. forbs such as crabbea velutina, hibiscus praeteritus, ruellia cordata (species group 5), evolvulus alsinoides, limeum fenestratum, talinum caffrum, clerodendrum ternatum (species group 10), ocimum americanum, phyllanthus maderaspatensis, lantana rugosa, hibiscus micranthus (species group 14), waltheria indica, kyllinga alba and melhania forbesii (species group 21) are present in this community. total species count for this community is 141, which is the highest for the letaba study area (table 1). this community scored the lowest on the diversity index and its low evenness (table 1) can be attributed to the almost complete dominance by colophospermum mopane in both the tree and the shrub layer. despite its low diversity, this community contains the highest richness of forbs, graminoids and annuals for the study area (table 3). half of its species are members of the fabaceae and poaceae, although the highest number of species in the malvaceae has been recorded in this community (table 2). table 1: richness, shanon-wiener diversity and evenness at plant community level of the letaba exclosures study site table 2: species counts for the 11 most dominant plant families recorded in the four major plant communities of the letaba exclosures study site the lowland savanna consists of two sub-communities that are easily recognisable in the field. even the variants are distinguishable and are represented by different mapping units (figure 2). 2.1 melhania rehmannii – enneapogon scoparius mixed bushveld: mixed bushveld of the letaba exclosures site is typically associated with midslopes and footslopes, extending over all exclosure fence lines into the buffer zone. it covers the largest area of the study site. the dominant soil form underlying this sub-community is mispah (paterson & steenekamp 2003). structurally, it varies from open shrubveld to closed high shrubveld, and is dominated alternately by colophospermum mopane and combretum apiculatum. as in most other plant communities of the letaba exclosures site, the highest vegetation cover is in the herbaceous layer (figure 5; table 3). species from species group 6 (see online appendix) characterise the floristic composition of this sub-community and include melhania rehmannii, aptosimum lineare, asparagus setaceus, abutilon angulatum, becium filamentosum, dumasia villosa and coccinia adoensis, all of which are representative of the forb layer. conspicuous woody species include cissus cornifolia (species group 5), colophospermum mopane, combretum apiculatum, dichrostachys cinerea (species group 14), cassia abbreviata (species group 20) and combretum hereroense (species group 21). prominent forbs, other than the character species, are crabbea velutina, ruellia cordata (species group 5), evolvulus alsinoides, dicoma tomentosa, clerodendrum ternatum (species group 10), ocimum americanum, lantana rugosa, melhania acuminata, phyllanthus incurvus, pavonia burchellii, hibiscus micranthus, tragia dioica (specis group 14), kyllinga alba and waltheria indica (species group 21). in the grass layer, pogonarthria squorrosa, schmidtia pappophoroides (species group 9), digitaria eriantha (species group 16), panicum maximum and tragus beteronianus (species group 28) are most common. this sub-community contains the highest number of graminoids, forbs and dwarf shrubs (table 3), which explains its high species richness (115 taxa, figure 5), the second highest for the letaba exclosures site. mean species per sampling plot is 32. diversity is average for the letaba study area, whereas the evenness is below average as a result of the dominance of colophospermum mopane (figure 5). table 3: plant species richness for different growth forms in each of the plant communities and sub-communities of the letaba exclosures study site two variants were identified that are easily be differentiated in the field, due to the difference in vegetation structure. although species composition does not vary significantly between these two variants, they alternate along the midslopes and footslopes of the landscape. 2.1.1 lannea schweinfurthii variant: this mosaic of moderately closed to closed savanna occurs throughout the study area on slightly deeper soils of the mispah soil form. the lannea schweinfurthii variant is very apparent from an aerial view and appears as green ‘tongues’ dissecting the savanna mopaneveld (figure 2). it is conspicuously present in the study area and structurally it becomes more dense towards the letaba river, in particular the area surrounding the mud pans (community 3.1) (figure 2). species group 7 (see online appendix) characterises this variant and include tree species such as lannea schweinfurthii and commiphora africana. the herbaceous layer is characterised by the climbers vigna luteola and the forbs indigofera nebrowniana, i. cryptantha and ocimum gratissimum. the vegetation is dominated by the tree colophospermum mopane (species group 14) and the grasses enneapogon scoparius (species group 10) and schmidtia pappophoroides (species group 5). prominent species include: the trees terminalia prunioides and combretum apiculatum, the grasses, panicum coloratum, (species group 14) and panicum maximum (species group 21) and the forbs, kyllinga alba and waltheria indica (species group 21). other woody species present, include maerua parvifolia (species group 9), philenoptera violacea (species group 16), cassia abbreviata (species group 20) and combretum hereroense (species group 21). forb species such as crabbea velutina (species group 5), evolvulus alsinoides, commelina africana, talinum caffrum and clerodendrum ternatum (species group 10) are also present. 2.1.2 aristida canescens variant: the aristida canescens variant makes up the open to moderately open part of this mixed bushveld, overlying shallow soils of the mispah soil forms (paterson & steenekamp 2003). it is found throughout the study area, stretching over all exclosure types and along the buffer zone in the north and the west (figure 2). species group 8 (see online appendix) floristically characterises the variant, which includes no woody species, but only the grass aristida canescens and the forbs rhynchosia totta, cleome monophylla and mariscus sumatrensis. trees such as colophospermum mopane, terminalia prunioides and combretum apiculatum (species group 14) dominate the woody layer, with prominent species in the forb layer being crabbea velutina (species group 5), ocimum americanum, phyllanthus maderaspatensis, ceratotheca triloba and phyllanthus incurvus (species group 21). schmidtia pappophoroides (species group 5), enneapogon scoparius (species group 10), panicum coloratum (species group 14), eragrostis superba, urochloa mosambicensis and panicum maximum (species group 21) are the most conspicuous grasses. 2.2 maerua parvifolia – agathisanthemum bojeri mopane bushveld: this mopane bushveld occurs along the bottomlands of the landscape, on the elevated terraces above the riparian zone of the letaba river. it forms a continuous band of moderately closed, tall colophospermum mopane shrubland, except in the full exclosure, where it extends higher up into the midslopes where the structure is significantly more open shrubveld. structurally, it resembles the lannea schweinfurthii variant (2.1.1), although the mopane bushveld is weaker in terms of species richness. this sub-community is associated with moderately deep to deep, brown to reddish-brown, unstructured sandy loam topsoil of the hutton soil form (paterson & steenekamp 2003). this moderately closed shrubland is characterised by species group 9 (see online appendix), of which the shrub maerua parvifolia and the forbs acalypha indica and heliotropium steudneri are the most prevalent. it has floristic affinities to the lannea schweinfurthii variant, of which maerua parvifolia, agathisanthemum bojeri and eragrostis rigidior provide evidence (species groups 8 and 9). colophospermum mopane (species group 14) dominates the tree and shrub layer, although other species, such as maerua parvifolia (species group 9), rhigozum zambesiacum (species group 10), terminalia prunioides and dichrostachys cinerea (species group 14) occur, but in low abundance values. other conspicuous species include those in the forb and grass layer, for example, schmidtia pappophoroides, crabbea velutina, hibiscus praeteritus (species group 5), evolvulus alsinoides, commelina africana, limeum fenestratum (species group 10), panicum coloratum, ocimum americanum, phyllanthus maderaspatensis, pavonia burchellii, jasminum stenolobum, (species group 14), eragrostis superba, kyllinga alba, waltheria indica, melhania forbesii and panicum maximum (species group 21). the mopane bushveld has the highest richness and percentage cover of annuals in the study area and the second highest richness of forbs (table 3). the overall species richness is 93 and the number of species recorded per sampling plot is 32, which is an average count for the study area. diversity and evenness is slightly below average for the letaba study area (figure 5), as expected in a community dominated by a single species, colophospermum mopane (see online appendix). 3. sporobolus fimbriatus – grewia bicolor seasonally wet savanna the seasonally wet savanna of the letaba exclosures study site refers to the seasonal streams and mud pans that are habitat for the azonal savanna vegetation, excluding the riparian zones. plant species that differentiate this community are floristically closer to the savanna communities (communities 1 and 2), because they receive water for limited periods of the year only. soil forms underlying this community vary from moderately deep hutton along the mud pans, to sandy mispah along the seasonal stream. structurally, this community is classified as a moderately open to moderately closed woodland, typically found along the edges of the watercourses, including the pans. forbs dominate the field layer, whereas shrubs dominate the woody layer. species group 11 (see online appendix) contains the diagnostic species for this community, including only grewia bicolor in the woody layer and forbs, such as justicia flava, hibiscus calyphyllus and the fern cheilanthes viridis. the grasses sporobolus fimbriatus (species group 11), eragrostis superba and panicum maximum (species group 21) are typically found in the field layer. other prominent species associated with this community include colophospermum mopane (species group 14), that occurs as large individuals on the perimeter of the mud pans and on the streambank. species of lesser abundance, but of value to differentiate this community, include cissus cornifolia (species group 5), terminalia prunioides, combretum apiculatum, the climber jasminum stenolobum (species group 14) and the woody shrub flueggea virosa (species group 21). this community has an average species richness (calculated at 119), although the average number of species per sampling plot is above average (27) (table 1) for the letaba site. it has the second highest diversity and the highest evenness of all communities (table 1). it hosts the highest richness of climber species (table 3). poaceae and fabaceae are the dominant plant families in this community (table 2). 3.1 eragrostis trichophora – sporobolus stapfianus mud pans: this sub-community comprises the vegetation associated with the zoogenic mud pans that are scattered in the letaba exclosure site. the four most prominent mud pans are located in the full exclosure on the bottomlands towards the letaba river (figure 2). these muddy patches, created by large herbivores seeking surface water to create a mud bath, are surrounded by tall mopane trees that, other than the surrounding high shrubland (community 2.1.1), develop into mopane woodland. soils underlying this sub-community, include deep, gravelly sandy loam of the hutton soil form. character species of this community are listed in species group 12, of which eragrostis trichophora is dominant. other character species include an oxalis species, cyperus esculentus, sporobolus stapfianus, hemizygia bracteosa, sporobolus africanus, urochloa panicoides, sesbania bispinosa and polygala sphenoptera. along the edges of the open mud pans are trees and shrubs, for example terminalia prunioides, combretum apiculatum, jasminum stenolobum (species group 14) and flueggea virosa (species group 21), although they occur in low abundances. dominant grass species include eragrostis trichophora (species group 12), and panicum maximum (species group 21). this sub-community consists of a unique composition of graminoids and geophytes, which contribute to the high cover percentage of herbaceous species in the study area (figure 5). it has a below average diversity and evenness (figure 5), due to the significantly high cover abundance and dominance of eragrostis trichophora. 3.2 barleria elegans – acacia karroo streambank woodland: streambank vegetation in the letaba exclosures site is associated with the two seasonal streams that dissect the study area from north to south. the larger flows through the partial exclosure, where the vegetation is also more typical of this community. the streambank vegetation is not well developed along the smaller tributary that flows through the full exclosure that contains many elements of the surrounding savanna areas. structurally, it is classified as moderately closed woodland. the woodland becomes more and more integrated with the surrounding savanna area as the distance away from the stream increases. soils are not the typically shallow mispah, as is the case in the adjacent savanna areas, but is overlain by gravelly sandy sediments of variable depths that are washed down from higher lying areas. diagnostic species for the streambank woodland include trees and shrubs such as manilkara mochisia, gymnosporia buxifolia, acacia karroo, tricalysia lanceolata and euclea divinorum. barleria elegans is a prominent diagnostic species, whereas climbers, such as vigna frutescens and cardiospermum corindum are also present. colophospermum mopane, terminalia prunioides, combretum apiculatum, dichrostachys cinerea, acacia nigrescens, jasminum stenolobum (species group 14) and combretum hereroense (species group 21) are some of the conspicuous species in the woody layer. other important species in the field layer include sporobolus fimbriatus (species group 11), lantana rugosa, pavonia burchellii (species group 14), eragrostis superba and panicum maximum (species group 21). this woodland contains species that are common in the riparian woodland, such as gymnosporia senegalensis (species group 15), philenoptera violacea, diospyros mespiliformis (species group 16), phyllanthus reticulatus, combretum microphyllum, cassia abbreviata, cocculus hirsutus and momordica foetida (species group 20). this moderately closed woodland is rich in tree and liana species. it is the only community in which the woody cover percentage is highest (figure 5). all other growth forms are well represented in this community, although of lower percentage cover (see online appendix). a total of 78 species have been recorded in this woodland type, with an average of 30 species per 400 m2 (figure 5). species diversity and evenness is the highest of all communities that occur at the letaba exclosures site (figure 5) and can be ascribed to the less clear-cut boundary between the streambank vegetation and the surrounding savanna vegetation. 4. croton megalobotrys – panicum deustum riverine woodland this riverine woodland comprises the communities that are separated from the zonal communities at the first hierarchical level of classification. it contains elements of several major communities identified by bredenkamp and van rooyen (1993), for example the ‘lonchocarpus capassa (= philenoptera violacea) –croton megalobotrys – colophospermum mopane dry riparian woodland’, the ‘croton megalobotrys – maytenus senegalensis (= gymnosporia senegalensis) riparian forest’ and the ‘phragmites mauritianus – cynodon dactylon reed communities’. according to the description of the subtropical alluvial vegetation (aza7), given by mucina and rutherford (2006), the croton megalobotrys – panicum deustum riverine woodland described here partially forms part of this inland azonal vegetation type with common species in the woody layer being phoenix reclinata, ziziphus mucronata, acacia nigrescens, colophospermum mopane, combretum hereroense, philenoptera violacea, euclea divinorum and gymnosporia senegalensis and, in the reedbeds, phragmites mauritianus and cynodon dactylon. this community covers approximately 10% of the total letaba site (including the 400 m buffer zone). it stretches along the letaba river bank from west to east, crossing exclosure fence lines. the largest part of this community is associated with deep, greyish-brown, sandy loam topsoil on brown, sandy-loam subsoil of the oakleaf soil form (paterson & steenekamp 2003). the vegetation structure of the croton megalobotrys – panicum deustum riverine woodland varies between an open grassy shrubveld to moderately closed woodland. true to its structural nature, it harbours the highest number of tree and shrub taxa in the study area (table 3). the field layer is dominated by forbs and annuals. species group 15 (see online appendix) floristically characterises this woodland, of which croton megalobotrys and gymnosporia senegalensis are most prolific in the woody layer. panicum deustum and indigofera tinctoria are prominent characteristic species in the field layer. the tree philenoptera violacea (species group 16) is also conspicuously present in this woodland. the total species count for this community is 131, the second highest at the letaba exclosures site. it has average evenness, but scored the highest diversity (table 1), suggesting that there is a lack of dominance of a single or few species and that all species are evenly distributed within the community. shrubs and trees are the dominant growth forms, and this community hosts the highest richness in tree species (table 3). it is dominated by taxa of the fabaceae and poaceae, and is characterised by many taxa of the rubiaceae, asteraceae and euphorbiaceae (table 2). the riverine forest is subdivided into sub-communities that represent small-scale differences in habitat and floristic composition. these sub-communities comprise: 4.1 philenoptera violacea – jasminum fluminense riverbank woodland: the riverbank woodland delineates the riparian zone of the letaba river, as it includes the riverbank, lower riverbank and riverbed. this sub-community is present in all the exclosure types, as well as in the buffer zone. the dominant soil form underlying this sub-community is oakleaf (paterson & steenekamp 2003). it is structurally classified as moderately closed woodland, where the trees and shrubs reach their maximum heights at the letaba exclosures site, albeit areas closer to the riverbed being classified as moderately open woodland and open riverbank vegetation. typical of riverbank vegetation, this sub-community has a high percentage cover by trees and shrubs, although percentage grass cover remains the highest. the riverine woodland community is similar to the ‘croton megalobotrys – maytenus senegalensis (= gymnosporia senegalensis) riparian forest’ community described by bredenkamp and van rooyen (1993). tree species common to both communities include croton megalobotrys, trichilia emetica, philenoptera violacea, nuxia oppositifolia and diospyros mespiliformis. species from species group 16 (see online appendix) characterise the floristic composition of this sub-community and include philenoptera violacea, diospyros mespiliformis, combretum microphyllum, jasminum fluminense, trichilia emetica and lippia javanica. other species that occur, but in lower abundance values, include the succulent shrub gardenia volkensii (species group 17), the grass cynodon dactylon and the tree nuxia oppositifolia, which is conspicuously present on the lower riverbank. taxa of species group 20 are all present in this community, of which some are more conspicous than others. they include phyllanthus reticulatus, combretum mossambicense and cassia abbreviata in the woody layer and lianas such as cocculus hirsutus and momordica balsamina. prominent tree and shrub species, other than the character species, include croton megalobotrys, gymnosporia senegalensis (species group 15), combretum hereroense and flueggea virosa (species group 21). important grasses are eragrostis superba and panicum maximum (species group 21), whereas melhania forbesii and waltheria indica (species group 21) are the only conspicuous forbs in the field layer. this sub-community has the highest species richness (116) of all sub-communities and its diversity is second highest in the study area (figure 5). tree and shrub species richness are also the highest in this sub-community (table 3). 4.1.1 hyphaene coriacea variant: this variant is similar to the ‘croton megalobotrys – hyphaene coriacea riparian and floodplain woodland’ described by bredenkamp and van rooyen (1993). it is a relatively moist woodland that covers the typical evergreen moderately closed riparian woodland zone along the letaba river on deep oakleaf soils. it is well represented in the full and partial exclosures, but discontinuous and fading in the open area and buffer zone. several tall (up to 12 m) riparian forest trees characterise this variant at the letaba exclosures site with its relatively high vegetation cover of trees and shrubs. philenoptera violacea is the most prominent ‘big tree’ species in this variant, whereas several individuals of diospyros mespiliformis is a diagnostic feature. diagnostic species for this variant are listed under species group 17 (see online appendix), of which the succulent shrub or tree gardenia volkensii, the lala-palm hyphaene coriacea and the tree or high shrub euclea natalensis, are the major contributors to the woody cover. in the forb layer thunbergia neglecta and tragia glabrata are most prominent, with no grass species being diagnostic for this variant. panicum deustum (species group 15) and p. maximum (species group 21) are dominant in the field layer. other prominent trees and shrubs are croton megalobotrys (species group 15), cassia abbreviata (species group 20) and combretum hereroense (species group 21). forb species are rare in this community, whereas the climber jasminum fluminense (species group 16) are conspicuously present. 4.1.2 nuxia oppositifolia variant: this variant falls into the ‘croton megalobotrys – nuxia oppositifolia riparian terrace bush’ (bredenkamp & van rooyen 1993). according to bredenkamp and van rooyen (1993), this riverine bush represents typical croton megalobotrys dominated vegetation, the most common and conspicuous vegetation type on the riverbanks and terraces and widespread along most parts of the letaba river. soils underlying this community are mainly of the oakleaf soil form that is associated with deep sandy-loam topsoil on sandy loam subsoil (paterson & steenekamp 2003). this is well represented in all zones, including the sacrificial zone. it is better developed in the full exclosure than in the other zones. this variant varies from an open riverbank in the control site, to moderately open woodland in the full and partial exclosures. it is typically more open nearer to the letaba river, where it is associated with phragmites mauritianus clumps. diagnostic species for this sub-community are listed in species group 18 (see online appendix), of which the grass cynodon dactylon and the tree nuxia oppositifolia being the most conspicuous. other differential species include woody species, such as phyllanthus reticulatus (species group 20), peltophorum africanum and flueggea virosa (species group 21). panicum maximum (species group 21) and cynodon dactylon dominate the grass layer. this variant is in some areas covered by a dense grass sward, especially by cynodon dactylon (species group 18). similar to the ‘lippia javanica – gymnosporia senegalensis riverbank scrub’ of the nkhuhlu exclosures site (siebert & eckhardt 2008), scattered clusters of woody species are found within this variant. these clusters serve as a microhabitat for colonisation by riparian woodland species. many precursors of riparian woodlands were recorded from this variant and include combretum hereroense, peltophorum africanum and cassia abbreviata. 4.2 combretum imberbe – enneapogon cenchroides open woodland: this open woodland sub-community occurs discontinuous within the riverbank woodland. it is best developed in the full exclosure along the lower terraces of the letaba river and is easily recognised in the field as an opening in the riverbank woodland, characterised by a dense grass sward and several scattered trees and shrubs. it is similar to the ‘colophospermum mopane – combretum imberbe dry riparian woodland’ (bredenkamp & van rooyen 1993). the dominant soil form and series underlying this sub-community is oakleaf (paterson & steenekamp 2003). diagnostic species of importance are listed in species group 19 (see online appendix) and include the dominant grass enneapogon cenchroides, the tree combretum imberbe, the shrub pechuel-loeshea leubnitzi and two geophytes, albuca setosa and ledebouria revoluta. croton megalobotrys (species group 15) and combretum hereroense (species group 21) are important contributors to the woody layer, whereas urochloa mosambicensis (species group 21) is an abundant annual grass species. forb species are not abundant, although kyllinga alba (species group 21) is the most prominent. grass cover is high in this sub-community, which contributes to the overall cover by the herbaceous layer of 45%, with only 16% in the woody layer (figure 5). species richness is low and the diversity below average (figure 5). conclusion (back to top) an investigation into the natural vegetation of the study area revealed different plant communities, some of which are unique in its plant assemblage, such as the xerophyta humilis – oldenlandea herbacea (community 1.1), and others that contain little variance within and between. mapping of the vegetation could be done down to sub-community level and, in some cases, even variant level. as expected in mopaneveld vegetation, plant communities of the exclosures are broadly associated with soil types. although not tested here, it is known that smaller scale variations in mopaneveld vegetation corresponds to soil moisture availability, because the mopaneveld is considered ‘event-driven’, especially in the field layer (du plessis 2001). the description of the distinguished plant communities, as well as ecological and floristic interpretations thereof, contribute significantly to the knowledge required for the future management of the vegetation. as ecologically sound plant communities were distinguished during this survey, the description of the vegetation should serve as a basis for further detailed and broad-based botanical studies at this long-term monitoring site to inform knp’s adaptive management approach. acknowledgements (back to top) the authors would like to thank south african national parks (sanparks) for funding the project. our sincere thanks to jacques venter, sanparks for logistic support during field surveying. izak smit, sanparks is thanked for providing spatial images. the post-flood programme is thanked for making available digital aerial maps of the study area. gis assistance was received from ms. marie du toit (north-west university). all field rangers are thanked for keeping a watchful eye over the field surveying team. editor: llewellyn c. foxcroft references (back to top) barbour, m.g., burk, j.h. & pitts, w.d., 1987, terrestrial plant ecology, cummings, california. bredenkamp, g.j. & van rooyen, n., 1993, ‘a survey of the vegetation of the letaba river in the kruger national park’, unpublished report, ekotrust. du plessis, f., 2001, ‘a phytosociological synthesis of mopaneveld’, msc dissertation, university of pretoria. germishuizen, g. & meyer, n.l., 2003, ‘plants of southern africa: an annotated checklist’, strelitzia 14, national botanical institute, pretoria. gertenbach, w.p.d., 1983, ‘landscapes of the kruger national park’, koedoe 26, 9–121. hennekens, s., 1996, megatab: a visual editor for phytosociological tables, computer program, version 1.0, user’s guide, giesen & geurts, ulft. hill, m.o., 1979, twinspan – a fortran program for arranging multivariate data in an ordered two way table by classification of individuals and attributes, computer program, cornell university, ithaca. mucina, l., rutherford, m.c. (eds.), 2006, ‘the vegetation of south africa, lesotho and swaziland’, strelitzia 19, south african national biodiversity institute, pretoria. mueller-dombois, d. & ellenberg, h., 1974, aims and methods of vegetation ecology, wiley, new york. o’connor, t.g., 1992, ‘woody vegetation-environment relations in a semi-arid savanna in the northen transvaal’, south african journal of botany 58(4), 268–274. o’keeffe, t. & alard g., 2002, field operations manual for herbivore & fire exclosures on the sabie and letaba rivers in the kruger national park, viewed n.d. april 2009, from http://www.sanparks.org/parks/kruger/conservation/scientific/exclosures/exclosure_field_manual.pdf o’keeffe, j. & rogers k.h., 2003, ‘heterogeneity and management of the lowveld rivers’ in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 41–58, island press, washington. parsons, m., mcloughlin, c.a., kotschy, k.a., rogers, k.h. & rountree, m.w., 2005, ‘the effects of extreme floods on the biophysical heterogeneity of river landscapes’, frontiers in ecology and the environment 3, 487–494. paterson, d.g. & steenekamp, p.i., 2003, ‘soil survey of letaba exclosure’, iscw report no. gw/a/2003/38, arc – institute for soil, climate & water, pretoria. siebert, f., bredenkamp, g.j. & siebert, s.j., 2003, ‘a comparison of mopaneveld vegetation in south africa, namibia and zimbabwe’, bothalia 33(1), 121–134. siebert, s.j., van wyk, a.e., bredenkamp, g.j. & siebert, f., 2003, ‘vegetation of the rock habitats of the sekhukhuneland centre of plant endemism, south africa’, bothalia 33(2), 207–228. siebert, f. & eckhardt, h.c., 2008, ‘the vegetation and floristics of the nkhuhlu exclosures, kruger national park’, koedoe 50(1), 126–144. van rooyen, n., theron, g.k. & grobbelaar, n., 1981, ‘a floristic description and structural analysis of the plant communities of the punda–milia–wambiya area in the kruger national park, republic of south africa: 3. the colophospermum mopane communities’, journal of south african botany 47(4), 585–626. visser, d.j.l., coertze, f.j. & walraven f., 1989, explanation of the 1:1000000 geological map, fourth edition, 1984: the geology of the republics of south africa, transkei, bophuthatswana, venda and ciskei and the kingdoms of lesotho and swaziland, government printer, pretoria. weather bureau, 1986, climate of south africa. pretoria, south africa, department of environmental affairs, weather bureau publication no. 40. filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 page 14 page 15 page 16 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 page 14 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 page 14 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 filelist convert a pdf file! page 1 vd merwe.qxd factors that determine the price of game p. van der merwe, m. saayman and w. krugell van der merwe, p., m. saayman and w. krugell. 2004. factors that determine the price of game. koedoe 47(2): 105–113. pretoria. issn 0075-6458. today, approximately 50 years after the first game farm was established in south africa, the game industry has grown significantly, and is growing still. there are roughly 9000 game farms and they make a large contribution to both the tourism industry and nature conservation. the price of the majority of game species in south africa shows an upward trend since 1990. lately, a number of species, such as roan antelope, reedbuck, white springbok, black wildebeest, grey rhebuck and white rhinoceros have seen significant increases in value. the price increases can be attributed to a number of factors. the factors include the growth in the number of game farms, that in turn is driven by the growth of the tourism industry in south africa, coupled with the growth of trophy hunting and increased demand for game meat products. farmers have diversified into game farming in areas that are otherwise not suitable for cattle or crop farming. these increases in demand has, however, to some extent been met by the success of game breeding programmes (specifically of endangered and rare species). the question that comes to mind is which factors have the most significant impact on the price of game. answers to this will help game farm owners to manage the fluctuation in game prices better. based on the above the aim of this paper is to determine to what extent the abovementioned factors have an influence on the price of game and which factors have the most significant impact. panel data consisting of the price of different game species, number of game farms, number of tourist arrivals and the number of trophies hunters were analysed for the years 1996-2001. a regression analysis is undertaken to identify the determinants of the price of game in south africa. the above-mentioned factors explain the movement in the price of game, specifically that the demand factors have a significant influence on the price. therefore an increase in the number of trophy and biltong hunters as well as demand for game farms impacts significantly on the price of game. keywords: game sales, game farm, nature conservation, ecotourism, game farm tourism, hunting. p. van der merwe, m. saayman and w. krugell, north-west university, school for entrepreneurship, marketing and tourism management: institute for tourism and leisure studies and the school of economics, risk management and international trade, private bag x6001, potchefstroom, 2520 republic of south africa. (ontpvdm@puk.ac.za). issn 0075-6458 105 koedoe 47/2 (2004) introduction fraser (1999) forecasts that south africa’s tourism industry will grow by more than 5.5 % between 1998 and 2010, which is better than the predicted 4.1% for the global market. it is estimated that arrivals will increase from six million international tourists in 2000 to 30.5 million in 2020 (wto 2001). the african continent can triple the size of its tourism industry by 2020 if proper efforts are made. this will result in forecast tourist arrivals of 77.3 million to africa. on the african continent, south africa attracts the most overseas tourists, which makes tourism one of the largest industries in south africa (wto 2001). research indicates that tourism contributes approximately 4.6 % to the gross domestic product (gdp). in 2002, tourism arrivals increased by 16 %, making south africa one of the highest vd merwe.qxd 2004/10/05 12:54 page 105 growth destinations in the world (thornton & feinstein 2002; gcis 1998). for both local and overseas tourists, scenic beauty and wildlife remain the major attractions (ecotourism) that south africa has to offer (gcis 1998; gcis 1999). it is a fact that 80 % of nature conservation in south africa is taking place on privately-owned land, such as game farms (eloff 2000; fox & du plessis 2000). hence, an increase in the demand for game, and for sustainable management of game farms in all its facets. game farm tourism, according to van der merwe & saayman (2001) is based on four pillars (fig. 1). the pillar of interest for this paper is the first mentioned, namely, breeding game and rare/endangered game which includes game auctions/sales. the reason for this is that researchers such as furstenburg & van niekerk (2004), eloff (2004) and du plessis (2003) have indicated that a great number of aspects influence the price of game. in order for game farm owners to manage their game farms and specifically their stock (game) more effectively it has become important to know which factors are having the most significant impact on the price of game, for if the price of game drops or increases it has an impact on the profitability and viability of game farms. the aim of this paper is to determine the factors that have the most significant impact on the price of game. game farming in south africa the size and scope of game farming in south africa in 2000, it was estimated that there were approximately 7000 privately-owned game farms in south africa, with a total surface area of 16 million ha (ebedes 2002) of which 5061 farms are fenced according to nature conservation regulations. research conducted in 1993 indicated that the surface area of exempted game farms constituted 8.5 % of the total agricultural land in south africa, which increased to 12.5 % in 2000, with a total of 1.7 million head of game (eloff 2002; erasmus 2000; cic 2004). according to bothma (2002) there was a 2.5 % increase in land utilised for game farming from 1998 to 1999. the latter translates into a 300 000 ha per year increase for the purpose of gamefarm tourism. flack (2002a) states that it is estimated that the conversion rate from cattle to game farming in 2002 was nearly 500 000 ha, which is 200 000 ha more than the 1998 to 1999 increase. statistics of 2002 showed that 13.3 % of the agricultural land was used for game farming, while national and provincial game reserves only covered 6 % or 6.1 million ha of land in south africa (flack 2002a; van der walt 2002). over and above the fact that the 7000 game farms in south africa during 2000 generated approximately r1 billion, game farms also employ approximately 63 000 people (van der merwe & saayman 2003). live game sales and the breeding of rare/endangered species is the second biggest generator of koedoe 47/2 (2004) 106 issn 0075-6458 game farm tourism breeding game and rare/endangered game hunting ecotourism processed game products fig. 1. four pillars of game farm tourism. vd merwe.qxd 2004/10/05 12:54 page 106 revenue for game farm tourism and contributed approximately 30 % of the total amount generated by game farms in south africa. eloff (1999) divided the breeding of game into two categories. firstly, breeding common game species, such as kudu, springbok and impala. secondly, breeding endangered and rare game species such as sable antelope, black rhino and roan antelope. in the two categories mentioned above the emphasis is on the production of game species to be auctioned to game farmers. game auctions analysis of game auctions from 1991 to 2001 reveals an average increase of 9 % per year over an 11-year period. the turnover per year grew from almost r9 million in 1991 to r105 million in 2002, with an average growth rate of r38 million per year over 12 years (table 1) (eloff 2002; eloff 2003; eloff 2004). table 1 shows a steady increase in the numbers of game sold over the 13-year period from 1991 to 2003. the values, however, indicates a surge in average game prices with increases in turnover value of 85 % in 1995/1996, 40 % in 1997/1998 and 38 % in 2000/2001. the volume of live game sold stabilised during the 2001 season but there was still an active and growing demand, especially for the rare game species. the more common species have recently shown signs of price maturity, but new record prices have also been set (bothma 2002). in 2003, the value of sales was 2.63 % less than in 2002, going down from r105 million to r102 million. factors that contributed to this reduction were the drought that was experienced during the second part of the year in one of the leading provinces in game farm tourism, namely the limpopo province, and a general cost increase in south africa. table 2 indicates the average increase or decrease in the price of the different game species over the past ten years. flack (2002b) indicates that in 1992, there were only nine auctions per year, with 9 546 heads of game auctioned. this grew to 48 auctions in 2000 and 17 702 heads of game sold. in 2002, this number increased to 20 022 (table 1). it decreased by 1.88 % in 2003, with a total of 19 645 heads of game sold at 59 auctions. therefore, noticably, a significant increase in the number of auctions per year as well as in the price of game issn 0075-6458 107 koedoe 47/2 (2004) table 1 auction sales over the past ten years year number of % change turnover % change game sold in volume in rand in value (tax excluded) 1991 8 292 8 999 871 1992 9 546 (+) 15% 10 059 969 (+) 20.66% 1993 11 499 (+) 19.93% 11 732 596 (+) 8.04% 1994 11 096 (-)3.08% 11 705 605 (-) 0.23% 1995 9 171 (-) 17.35% 14 335 894 (+) 22.47% 1996 11 340 (+) 23.65% 26 559 557 (+) 85.26% 1997 12 077 (+) 6.5% 28 526 052 (+) 7.4% 1998 14 354 (+) 18.85% 40 017 964 (+) 40.29% 1999 15 455 (+) 7.67% 53 705 823 (+) 34.2% 2000 17 702 (+) 14.54% 62 960 451 (+) 17.23% 2001 17 282 (-) 2.37% 87 000 473 (+) 38.18% 2002 20 022 (+) 15.85% 105 192 180 (+) 20.91% 2003 19 645 (-) 1.88% 102 420 445 (-) 2.63% (source: eloff 2002, 2003, 2004) vd merwe.qxd 2004/10/05 12:54 page 107 in general. hence, the question: what has been driving the boom in the price of game? according to table 2, the five game species that had the biggest average percentage price increase per year from 1991 to 2000 were the following; nyala (210.32 %), kudu (142.51 %), grey rhebuck koedoe 47/2 (2004) 108 issn 0075-6458 table 2 percentage price increase (+) or decrease (-) in average auctioned game prices over the past ten years species average % price % increase (+) increase or decrease (-) per year in in 2000 compared ten years with 1999 roan antelope 40.81% (-) 8.2% tsessebe 21.08% (+) 49.25% blesbok: white 13.48% (-) 19.50% blesbok: common 15.04% (+) 3.84% blue wildebeest 19.92% (-) 0.98% bontebok 21.59% (+) 15.9% bushbuck 28.49% (+) 23.92% buffalo/disease free 29.65% (+) 53.13% duiker 18.73% (+) 32.07% eland/cape 10.58% (+) 13.44% gemsbok 16.40% (+) 4.74% giraffe 9.10% (+) 13.79% klipspringer 14.35% (-) 6.45% kudu 142.51% (+)16.11% lion 5.42% (-) 54.86% nyala 210.32% (+) 80.33% reedbuck 49.23% (+) 49.65% impala 19.8% (+) 5.11% red hartebeest 15.72% (+) 8.74% mountain reedbuck 29.88% (+) 22.14% burchell's zebra 8.18% (+) 18.97% springbok: common 22.37% (-) 6.83% springbok: heartwater 7.4% (+) 30.64% springbok: black 20.3% (+) 13.47% springbok: white 38.44% (+) 155.2% steenbok 22.34% (+) 53.61% black rhinoceros 5.31% (+) 69.68% black wildebeest 49.98% (+) 5.1% sable antelope 11.19% (+) 9.85% fallow deer 28.83% (+) 1.64% grey rhebuck 82.68% (-) 12.72% warthog 11.24% (-) 8.7% ostrich 10.58% (+) 39.46% waterbuck 12.00% (+) 36.5% white rhinoceros 30.37% (+) 39.67% total game 11.35% (+) 14.54% total turnover 60.02% (+) 17.23% (source: eloff 2002) (82.68 %), black wildebeest (49.98 %) and reedbuck (49.23 %). the five game species that had the biggest increase in sales in 2000–2001 were as follows; tsessebe (60.9 %), springbok (31.2 %), sable antelope (24.2 %), nyala (20.2 %) and burchell’s zebra (13.9 %) (eloff 2002). determinants of the price of game the price of game is determined by factors that influence the demand for and supply of game. microeconomics explains that price is determined by the interaction of demand and supply. this can be illustrated with a simple diagram. figure 2(a) shows a negative relationship between price and the quantity demanded and a positive relationship between price and quantity supplied. the equilibrium price and quantity are set where the two curves intersect. figure 2(b) abstracts from the situation in the south african game industry where there has been an increase in the equilibrium price and quantity over a number of years. this section examines the determinants of the price of game, by looking at the factors that could have caused the increases in demand and supply. at the most basic level the increase in the demand for game stems from the growth in the number of game farms in south africa, described in section 2 (absa 2003). a number of factors have played a role in the conversion of regular commercial farms into game farms. these major factors are the following: problems that traditional livestock/crop farming are currently experience makes it a good a option such as to compete with other countries that do receive subsidies from their governments; vd merwe.qxd 2004/10/05 12:54 page 108 the deregulation of the agricultural sector, resulting in lower, but also more competitive prices; an increase in the loss of livestock due to stock theft on farms; the control of cattle diseases had become an expensive operation; a decline in farmers’ political power base in parliament since the early 1990s; the number of land claims that increased; bush encroachment that increased (which often resulted from overgrasing which, in turn, reduced the frequency of veldfires); the impact of global warming (absa 2003; eloff 2001; van zyl 2002; du plessis 2003; van der walt 2003; schak 2003a; smith 2004). while there are drivers of growth there are also forces counteracting the growth of the demand for game. these include the following: political developments such as new legislation put forward by political parties can also influence the price of game, proposed translocation act, firearms control act (act 60 of 2000) firearms control aspects regarding overseas hunters and south africa’s crime situation; economic developments, such as the distance that game need to be transported to market, a decrease in discretionary income, population growth and need for land, possible surplus of game in future, stronger rand hampering game meat exports and game capturing technique use to capture game; climate/weather conditions namely, drought and floods; the pressure against the consumptive use of wildlife; diseases such as foot-and-mouth, anthrax, tuberculosis, rinderpest, bovine malignant and african swine fever to mention a few (phasa 2000; phasa 2003; du toit et al. 2002; thomson 2004; schak; 2003b; makhubela 2004; furstenburg & van niekerk 2004; hamman et al. 2002; eloff 2004; damm 2004b; damm 2004a; damm 2004c; scriven & eloff 2003; neethling 2003). while the higher prices discussed in the previous section reflects an increase in the demand for game, table 1 also showed that the demand was to some extent met by increases in the supply. table 1 showed a steady increase in the numbers of game sold over the period. the factors that have been influencing the supply-side of the market include factors such as: issn 0075-6458 109 koedoe 47/2 (2004) fig. 2. interaction of demand and supply vd merwe.qxd 2004/10/05 12:54 page 109 the development of more game farms climate/weather conditions namely good raining seasons the consumptive use of game such as biltongand trophy hunting the success obtained in the breeding programmes of rare/endangered game farmers today are more conservation orientated the economic value that lies in the breeding with game (du plessis 2003; eloff 2001; kzn-wildlife 2003; van der merwe & saayman 2001; van zyl 2002). the following section undertakes econometric analysis of the determinants of the price of game. empirical analysis the empirical analysis of the determinants of the price of game involves regressing the average price on a number of explanatory variables. the estimating equation can be expressed as follows: pit = c + ßqit + εit where pit is the average price of specie i in year t. the intercept, or constant term, is represented by c. the qit represents a vector of determinants of the price of game (where ß denotes the coefficients). unfortunately, data constraints limit the variables that can be included here. many of the determinants discussed in the previous section are particularly difficult to quantify such as drought or good rainy seasons, new legislation and diseases. there is a lack in good research in the game farm tourism industry such as how many biltong hunters hunt per year and how much do they spend to mention only a few. the explanatory variables that can be included are: the quantity of game, the number of game farms in the country, the number of tourists visiting south africa, as well as the number of trophy hunters. following the law of demand one could expect a negative relationship between price and quantity. the numbers of game farms, visiting tourists and trophy hunters are expected to be positively related to the price of game. the error term (ε ) captures the effects of unmeasured attributes that affect the price of game. a panel data set is used to estimate the regression equation. it contains data on the prices and quantities of 43 species of game sold at auction over the period 1996 to 2003. the list of species is included in the appendix. data on the number of game farms, number of tourists and trophy hunters are obtained from phasa (professional hunters association of south africa, south africa tourism and the centre for wildlife economics at north-west university. where possible natural logs were taken. regressions were estimated using stata 8 software. the estimation strategy involves estimating levels and growth rates versions of the estimating equation, using a panel data random effects estimator. the estimator is a weighted average of the between estimator that uses the variation between cross-section observations (in other words, between species), and the within estimator that uses the time variation within each cross-section of observations. table 3 shows the results for the levels and growth rates specifications. in the levels specifications the numbers of farms, tourists and trophy hunters explain the price level. the levels model (2) provides the best results. as expected, there is a negative and significant relationship between the price level and the quantity of game sold at auctions. there is a positive relationship between the price of game and the number of game farms and the number of trophy hunters. the results are similar to that of levels model (1) where the numbers of tourists, rather than the number of trophy hunters, are used, but it has a slightly better overall fit. including both the numbers of tourists and the numbers of trophy hunters as explanatory variables, as is done in the levels model (3), causes the tourists variable to become insignificant. overall, it is clear though that the models have low explanatory power. the levels model (2) has an overall r² of 0.13, koedoe 47/2 (2004) 110 issn 0075-6458 vd merwe.qxd 2004/10/05 12:54 page 110 issn 0075-6458 111 koedoe 47/2 (2004) table 3 estimation results variable levels levels levels growth growth growth model (1) model (2) model (3) rates (4) rates (5) rates (6) game sold -0.036 -0.092 -0.090 -0.005 -0.007 -0.007 (-1.24) (-2.32)* (-2.30)* (-2.18)* (-2.69)* (-2.64)* number of 0.540 1.115 0.844 0.649 1.291 1.046 game farms (3.47)* (11.35)* (4.37)* (3.73)* (11.95)* (4.88)* tourists 1.445 0.790 4.857 2.281 (3.89)* (1.63) (3.67)* (1.32) trophy 0.413 0.286 0.435 0.323 hunters (4.71)* (2.45)* (4.58)* (2.54)* r² within 0.49 0.55 0.56 0.48 0.55 0.56 between 0.11 0.20 0.20 0.07 0.10 0.10 overall 0.07 0.13 0.14 0.08 0.09 0.10 (* significant at a 5 % level) indicating that the conditioning variables explain 13 per cent of the variation in the price level over time and between species. this implies that the other determinants of game prices that are more difficult to quantify, also have a significant influence on the price level for example drought, change in legislation and diseases. the ‘within’ r² is encouraging in this regard. for example, level model (2) does explain 55 % of the ‘within’ (in other words, across time) variation. the growth rate specifications explain the changes in the price of game in terms of changes in the numbers of farms, tourists and trophy hunters. the models yield largely similar results regarding the signs and significance of coefficients and the overall explanatory power of the model. the benefit of also running the growth rates estimations is that one can interpret the coefficients. for example, the growth rates level model (5) indicates that 12 % increase in the number of game farms in the country will lead to a 10 % increase in the price of game. similarly, a four per cent increase in the number of trophy hunters will lead to a 10 % increase in the price of game. implications for game farms in south africa where there is a growing demand for land, one can only foresee that a saturation point will be reached in terms of the availability for new developments and this will have an impact on the price of game species (van der merwe & saayman 2002). therefore game farm owners have to think of alternative methods that will sustain game prices in the long run. alternative methods include. firstly, to start a marketing campaign in order to increase south africa’s market share of trophy hunting, because hunting is a consumptive form of usage. if one can increase the number of overseas trophy hunters one will also increase the demand in game. secondly, a similar process need to be followed with biltong hunting, if one is able to increase the number of biltong hunters then there will also be an increase in the demand for game. thirdly, the game industry need to position itself better as a supplier of game meat (venison) which will imply finding more and bigger markets. these methods can help to sustain a particular level of pricing. vd merwe.qxd 2004/10/05 12:54 page 111 conclusion the aim of this paper was to determine the factors that have the most significant impact on the price of game. it is clear from the paper that game sales had increased over the last twelve years from nine auctions per year (1992) and 9 546 game auctioned, to 59 auctions and 19 645 game auctioned in 2003. the five game species that had the biggest average percentage price increase per year from 1991 to 2000 were the following; nyala (210.32 %), kudu (142.51 %), grey rhebuck (82.68 %), black wildebeest (49.98 %) and reedbuck (49.23 %). the literature also indicated that ecotourism and trophy hunting did become a booming industry in south africa. a number of factors were identified that influence the price of game, viz., a decrease in the development of game farms; drought, or on the other hand, a good raining season; political instability; the rand/dollar exchange rate; and the proposed translocation act. empirical analysis of the determinants of the price of game showed that the number of game farms, tourists and trophy hunters are significant explanatory variables. trophy hunting has been identified as an important determinant of game prices. these factors have an impact on the price of game and the demand for game is important, since a decrease in these industries can harm the game selling industry significantly. references absa. 2003. report: game ranch profitability in south africa. johannesburg: sa financial sector forum. bothma, j. du p. 2002. some economics of wildlife ranching. pp. 23-29. in: penzhorn, b.l. (ed.). game ranch planning and management. symposium held at onderstepoort, pretoria on 1-2 nov. 2002. pretoria: wildlife group of the south african veterinary association. cic (international council for game and wildlife conservation). 2004. tourism and development: the win-win performance. africa indaba 2(2): 2. damm, g.r. 2004a. editor's comment. africa indaba 2(2): 1. damm, g.r. 2004b. iucn welcomes adoption of sustainable use principles by cbd. africa indaba 2(2): 2-4. damm, g.r. 2004c. sustainable lion hunting. africa indaba 2(2): 14-15. du plessis, p. 2003. gevaarseine. wild & jag 9(7):1. du toit, j.g., b.l. penzhorn & j. van heerden. 2002. bacterial, viral and protozoal diseases. pp. 295-330. in: bothma, j. du p. (ed.). game ranch management. 4th edition. pretoria: van schaik. ebedes, h. 2002. preface. pp. 7-9. in: ebedes, h., b. reilly, w. van hoven & b. penzhorn (eds.). sustainable utilization—conservation in practice. proceedings of the 5th international wildlife ranching symposium held at pretoria on 20-23 march 2001. pretoria: south african game ranchers association. eloff, t. 1999. wins uit wildboerdery. sa wild & jag 5(2): 22-25. eloff, t. 2000. die omvang van die wildbedryf in suid-afrika. potchefstroomse universiteit vir cho. ongepubliseerde verslag. eloff, t. 2001. knelpunte en geleenthede. sa game & hunt 2(5): 9. eloff, t. 2002. the economic realities of the game industry in south africa. pp. 78-86. in: ebedes, h., b. reilly, w. van hoven & b. penzhorn (eds.). sustainable utilization—conservation in practice. proceedings of the 5th international wildlife ranching symposium held at pretoria on 20-23 march 2001. pretoria: south african game ranchers association. eloff, t. 2003. gemiddelde wildveilingpryse 2002. sa wild & jag 9(2): 21. eloff, t. 2004. game sales 2003 in south africa. africa indaba 2(2): 5. erasmus, s.d. 2000. die ekonomiese lewensvatbaarheid van 'n wildvleisbewerkingsaanleg in suid-afrika. m.com. verhandeling. potchefstroomse universiteit vir cho. flack, p.h. 2002a. the conservation revolution. sa game & hunt 8(10): 29-33. flack, p. h. 2002b. exotic game—catching up with texas? magnum (oct.): 76-80. fox, t. & p. du plessis. 2000. hunting in southern africa. africa indigo 3(3): 40-47. fraser, j. 1999. tourism is worth r80,6bn to sa. the star (aug. 17): 6. furstenburg, d. & p. van niekerk. 2004. die wildbedryf en die voorgestelde wildhervestigingswet. sa wild & jag 10(6): 30-31. koedoe 47/2 (2004) 112 issn 0075-6458 vd merwe.qxd 2004/10/05 12:54 page 112 gcis (government communication and information system). 1998. south africa official yearbook. cape town: rustica. gcis (government communication and information system). 1999. south africa official yearbook. cape town: rustica. hamman, k. h. stadler & p. lloyd. 2002. is die wildbedryf versoenbaar met natuurbewaring? sa wild & jag 9(4): 35-36. kzn-wildlife. 2003. game auctions. sa game & hunt 9(8): 21-23. makhubela, w. 2004. firearms control act south africa. africa indaba 2(2): 5. neethling, p. 2003. sterk rand knel wildvleisuitvoer. sa wild & jag 9(7): 53. phasa (professional hunters association of south africa). 2000. professional hunters association of south africa. statistics. pretoria: phasa. phasa (professional hunters associations of south africa) 2003. professional hunters association of south africa. statistics. pretoria: phasa. schak, w. 2003a. brusellos. sa wild & jag 9(10): 24-25. schak, w. 2003b. bek-en-klouseer. sa wild & jag 9(5): 24-25. scriven, l. & t. eloff. 2003. markets derived from nature tourism in south africa and kwazulunatal. pp. 247-324. in: aylward, b. & e. lutz (eds.). nature tourism, conservation, and development in kwazulu-natal, south africa. washington, dc: world bank publishers. smith, c. 2004. wild behaal rekordpryse op rooikraal-wildveiling. volksblad, mrt. 27: 3. thomson, r. 2004. national parks in crisis? africa indaba 2(2):16. thornton, g. & k. feinstein. 2002. tourism talk: a summery of the most important data and statistics relevant to the regional tourism industry. the journal of southern africa tourism 1(3): 48-53. van der merwe, p. & m. saayman. 2001. managing game farms from a tourism perspective. potchefstroom: ons drukkers. van der merwe, p. & m. saayman. 2002. a different approach to wildlife tourism and hunting. pp. 128-136. in: ebedes, h., b. reilly, w. van hoven & b. penzhorn (eds.). sustainable utilization—conservation in practice. proceedings of the 5th international wildlife ranching symposium held at pretoria on 20-23 march 2001. pretoria: south african game ranchers association. van der merwe, p. & m. saayman. 2003. determining the economic value of game farm tourism. (paper delivered at the 55th international atlantic economic conference at vienna austria on 13–16 march 2003.) vienna. (unpublished.) van der walt, j. 2002. proliferation of game ranches. sa game & hunt 8(10): 7. van der walt, j. 2003. malignant catarrhal fever. sa game & hunt 9(8): 7. van zyl, j. 2002. wees mak as jy jou somme maak. finansies en tegniek mrt: 22. world tourism organisation (wto). 2001. africa to triple tourist numbers. [web:] h t t p : / / w w w. w o r l d t o u r i s m . o rg / n e w s r o o m /releases/more _releases/r010607.html [date of access: 19 march 1999]. issn 0075-6458 113 koedoe 47/2 (2004) vd merwe.qxd 2004/10/05 12:54 page 113 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 filelist convert a pdf file! page 1 page 2 page 3 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 dippenaar sovenga.qxd a checklist of spiders from sovenga hill, an inselberg in the savanna biome, limpopo province, south africa (arachnida: araneae) m.a. modiba, s.m. dippenaar and a.s. dippenaar-schoeman modiba, m.a., s.m. dippenaar and a.s. dippenaar-schoeman. a checklist of spiders from sovenga hill, an inselberg in the savanna biome, limpopo province, south africa (arachnida: araneae). koedoe 48(2): 109–115. pretoria. issn 0075-6458. the south african national survey of arachnida (sansa) was initiated to make an inventory of the arachnid fauna of south africa. various projects are underway to prepare inventories of the spider fauna of the different floral biomes and provinces of south africa. during april and may 2004 five different collecting methods were used to sample spiders from four slopes on sovenga hill, an inselberg situated in the savanna biome, near polokwane, in the limpopo province of south africa. a total of 793 specimens represented by 29 families, 62 genera and 76 species were recorded over the twomonth period. the thomisidae was the most abundant (n = 167) representing 21.1 % of all spiders sampled, followed by the gnaphosidae (n = 101) with 12.7 % and the lycosidae (n = 77) with 9.7 %. the most abundant species was a thomisid tmarus comellini garcia-neto (n = 82), representing 10.3 % of the total, followed by a clubionid clubiona godfreyi lessert (n = 66) with 8.3 %. the thomisidae was the most species-rich family with 12 species, followed by the gnaphosidae with 11 species and the araneidae with 10 species. of the species collected 83.9 % were wandering spiders and 16.1 % web builders. this is the first quantitative survey of the savanna biome in the polokwane area. keywords: araneae, checklist, diversity, limpopo province, savanna biome, south africa, spiders. m.a. modiba, s.m. dippenaar, school of molecular and life sciences, university of limpopo, private bag x1106, sovenga, 0727, south africa; a.s. dippenaar-schoeman, arc-plant protection research institute, private bag x134, queenswood, 0121, south africa/department of zoology and entomology, university of pretoria, south africa issn 0075-6458 109 koedoe 48/2 (2005) introduction although spiders constitute an abundant and successful group of invertebrates in south africa they are poorly sampled and little is known about their diversity within most ecoregions (dippenaar-schoeman 2002a). this lack of knowledge undermines meaningful conservation (de wet & schoonbee 1991). additionally, the critical lack of professional taxonomic expertise within the country presents significant problems for the understanding of invertebrate biodiversity, which leads to an under-appreciation and estimation of the actual species pool (gibbons 1999; dippenaar-schoeman 2002a). in south africa, more than 2000 spider species have been recorded but only about 30 % of the families have been revised (dippenaarschoeman 2002a). the south african national survey of arachnida (sansa) was initiated in 1997 to address this lack of baseline information by conducting surveys and biodiversity assessments of the arachnid fauna of south africa (dippenaar-schoeman & craemer 2000). one such a project of sansa is to determine spider diversity in the savanna biome of south africa, with the primary aim to gather information on areas still poorly sampled (dippenaar-schoeman & leroy 2003; van den berg et al. 2003). research on the spider fauna of the savanna biome in south africa is presently restricted to surveys of the roodeplaat dam nature reserve (dippenaar-schoeman et al. 1989); makalali game reserve (whitmore et al. 2001; whitmore et al. 2002), the spiders of the western soutpansberg mountains (foord et al. 2002), a survey of the spiders of the kruger national park (dippenaar-schoeman & leroy 2003) and the spiders of the springbok flats (van den berg et al. 2003). there are numerous collecting methods used for sampling arachnids (eardley & dippenaar-schoeman 1996), but there are biases associated with some methods frequently resulting in under-sampling of sites when only one method is used or one habitat is sampled (new 1999). scientists have for years recognised that standard methods are essential to be able to compare biodiversity assessments (coddington et al. 1996). spiders are able to occupy nearly every terrestrial habitat (dippenaar-schoeman & jocqué 1997). it is therefore essential to sample the fauna separately at different layers during biodiversity surveys. areas showing low diversity may simply be a reflection of inadequate sampling (whitmore et al. 2002), therefore sampling protocols are very important. unfortunately, many of the surveys undertaken in south africa have used restricted sampling techniques, sampling only one habitat type. in this study, a survey was undertaken to determine the spider diversity of sovenga hill, an inselberg, situated near polokwane, limpopo province. during a two-month period this inselberg was extensively surveyed using five collecting methods. twenty-four sites were sampled and samples were taken from both the ground, litter, grass and tree layers. the collecting methods were evaluated for surveys to be continued in the polokwane area. this is the first of a series of papers on spider diversity of the savanna biome in the polokwane area. material and methods sovenga hill (23º53's, 29º44'e), an inselberg, was sampled over a two-month period from 1 april to 31 may 2004. this unique landform consists of domeshaped granite rocks and is situated on the campus of the university of limpopo near polokwane in the limpopo province of south africa. it covers an area of approximately 6.3 ha and is surrounded by indigenous flora consisting mostly of euphorbia cooperi n.e.br. ex a. berger var. cooperi, heteropyxis natalensis harv. and croton gratissimus burch. var. gratissimus. this is a closed woodland plant community classified as mamabolo mountain bushveld of which small, outlying, isolated communities are found on the pietersburg plateau (s.a. vegmap project, in prep. p. winter pers. comm.). six sampling sites of 4 m x 4 m were selected on each of the southern, eastern, northern and western slopes, resulting in a total of 24 sites sampled. the first two sites were at an altitude of between 1318 m–1335 m, the second two sites between 1333 m–1354 m and the third two sites between 1347 m–1374 m. sampling was conducted twice a week during april and may 2004. during april sampling was conducted on all the sites while no sampling was done on the sites of the northern slope during may due to vandalism of the sampling sites. the following methods were used during sampling: sweep netting (s) — a sweeping net was used to sweep through the grass and herb layer to dislodge specimens into the net. walking in a straight line, 20 sweeps were taken to form one sample. the contents from the net were emptied into a marked plastic bag and spiders and other organisms were separated from the vegetation in the laboratory. a total of 101 sweepnet samples were taken. this method could only be used at sites where grass and herb layers were present. active searching (a) — active lifting of stones and searching for spiders was done at all sites for 10 minutes per sampling session (eight session during april and nine in may). tree beating (t) — a stick was used to knock spiders from trees while a tray was used to catch all the specimens and plant material knocked off. a total of 20 beats formed one sample. the contents from the tray were placed in a marked plastic bag and spiders and other organisms were separated from the vegetation in the laboratory. a total of 354 samples were collected during the sampling period using this method. different types of trees present on a site were sampled. pitfall trapping (p) — small plastic containers (10.5 cm diameter) were planted into the ground, with the upper rim level with the ground surface. wooden planks of about 30 cm long were placed in three directions towards the trap to increase its effective catch area. a funnel was placed over the opening of the trap to prevent koedoe 48/2 (2005) 110 issn 0075-6458 spiders from escaping and a small container with 70 % ethanol was placed inside to immobilise and preserve the captured specimens. there were 20 pitfall traps per slope, randomly distributed. the traps were kept open for the whole period and were emptied twice a week. leaf litter sifting (l) — a leaf litter sample was collected in a wooden box (53 cm x 20 cm x 18.5 cm) with a chicken wired base and specimens were sieved from the litter. one sample consisted of enough litter to fill the wooden box. two samples of litter were collected from sites where leaf litter was found. captured specimens were preserved in 70 % ethanol and were sorted and counted in the laboratory. all spiders were identified to species level by the third author, where possible. some specimens could not be identified to species owing to the unresolved taxonomy of certain families in africa (for instance the lycosidae) and the immature stages of some specimens collected. voucher specimens were deposited in the national collection of arachnida (nca) at the plant protection research institute in pretoria, an institute of the agricultural research council. results and discussion during the two-month sampling period a total of 793 spiders, represented by 29 families, 62 genera and 76 species were collected (table 1). the few published surveys of spiders from herbaceous and ground layers in africa indicate that the relative abundance of species differs between different areas and different plant biomes (dippenaarschoeman et al. 1989; russell-smith 1981). three surveys of spiders are known from the limpopo province. a study at makalali private game reserve was conducted over 11 months and resulted in a total of 4832 spiders collected, representing 38 families (whitmore et al. 2002). another study, conducted over a two-year period on the springbok flats, resulted in a total of 3139 specimens caught, representing 35 families (van den berg et al. 2003). sporadic collection over a five-year period at lajuma in the western soutpansberg resulted in 127 species belonging to 46 families (foord et al. 2002). from the above it is clear that although the current study was only conducted for two months, on an isoissn 0075-6458 111 koedoe 48/2 (2005) table 1 spider families collected at sovenga hill indicating the number of species and percentage of the total number of specimens collected family genera species specimens % agelenidae 1 1 6 0.8 amaurobiidae 1 1 4 0.5 araneidae 8 10 57 7.2 corinnidae 3 3 42 5.3 clubionidae 1 1 66 8.3 caponiidae 1 1 9 1.1 ctenidae 1 1 7 0.9 deinopidae 1 1 2 0.3 eresidae 1 1 8 1.0 gnaphosidae 5 11 101 12.7 hahniidae 1 1 6 0.8 hersiliidae 1 1 3 0.4 linyphiidae 1 1 7 0.9 lycosidae 2 2 77 9.7 miturgidae 1 1 7 0.9 nemesiidae 1 1 11 1.4 oxyopidae 1 1 18 2.3 palpimanidae 1 1 4 0.5 philodromidae 2 5 51 6.4 pholcidae 1 1 2 0.3 pisauridae 1 1 9 1.1 salticidae 5 5 54 6.8 scytodidae 1 1 16 2.0 segestriidae 1 1 3 0.4 selenopidae 1 1 12 1.5 theridiidae 6 6 21 2.6 thomisidae 9 12 167 21.1 uloboridae 1 1 3 0.4 zodariidae 2 2 20 2.5 total 62 76 793 100 lated hill, a good representation of families (29) were caught. however, because of the short duration of the present study, the results may not reflect the actual abundance and species richness of sovenga hill, but nevertheless provide an indication of the minimum abundance and richness. the family composition of the spider fauna as a whole is shown in tables 1 and 2. the thomisidae was the most abundant family (n = 167) representing 21.1 % of all spiders sampled, followed by the gnaphosidae (n = 101) representing 12.7 %, the lycosidae (n = 77) representing 9.7 % and clukoedoe 48/2 (2005) 112 issn 0075-6458 table 2 checklist of the spiders of sovenga hill, polokwane, south africa. (a: active search; b: beating; l: litter sifting; p: pittrapping; s: sweepnetting; w: wanderer; wd: web dweller) family species nr method guild habitat agelenidae agelena sp. (immature) 6 a wd funnel-web amaurobiidae chresiona sp. (immature) 4 p;l wd retreat-web araneidae argiope sp. (immature) 1 b wd orb-web argiope australis (walckenaer, 1805) 3 b wd orb-web caerostris sexcuspidata (fabricius, 1793) 5 b wd orb-web cyphalonotus larvatus (simon, 1881) 5 b;l wd orb-web cyrtophora citricola (forskal, 1775) 1 b wd orb-web nemoscolus sp. (immature) 1 b wd orb-web neoscona blondeli (simon, 1885) 20 p;b;l wd orb-web neoscona subfusca (c.l. koch, 1837) 19 b;l;a wd orb-web prasonica sp. (immature) 1 b wd orb-web singa sp. (immature) 1 l wd orb-web caponiidae caponia sp. (immature) 9 p;a w soil clubionidae clubiona godfreyi lessert, 1921 66 p;b;l w soil corinnidae thysanina sp. 12 p;l w soil cetonana sp. (immature) 13 p;l w soil copa flavoplumosa simon, 1885 17 b;l w soil ctenidae ctenus sp. (immature) 7 l;b w soil deinopidae menneus camelus pocock, 1902 2 b wd throw-web eresidae gandanameno sp. (immature) 8 p;b;l wd retreat-web gnaphosidae asemesthes sp. (immature) 2 l;b w soil camillina sp. (immature) 6 l;p;b w soil drassodinae (undetermined) 2 b w soil setaphis browni (tucker, 1923) 12 l;a;p w soil zelotes lightfooti (purcell, 1907) 21 l;p w soil zelotes oneili (purcell, 1907) 13 p;l w soil zelotes reduncus (purcell, 1907) 12 b;a;l;p w soil zelotes ungulus tucker, 1923 17 l;p w soil zelotes sp. a 3 p w soil zelotes sp. b 3 b;p w soil hahniidae hahnia tabulicola simon, 1898 6 l wd sheet-web hersiliidae hersilia sericea pocock, 1898 3 b w tree trunk linyphiidae sp. (undetermined) 7 p wd sheet-web lycosidae sp. (undetermined) 69 p w soil pardosa sp. 8 p w soil miturgidae cheiracanthium sp. (immature) 7 b w leaves nemesiidae lepthercus sp. 11 p;b w burrow soil oxyopidae oxyopes schenkeli lessert, 1927 18 p;b w grass palpimanidae palpimanus transvaalicus simon, 1893. 4 p w soil philodromidae philodromus sp. (immature) 10 b;a,s w grass philodromus brachycephalus lawrence, 1952 15 b,s w grass philodromus grosi lessert, 1943 1 b,s w grass philodromus partitus lessert, 1919 3 s w grass suemus punctatus lawrence, 1938 22 b w grass pholcidae smeringopus sp. (immature) 2 b wd space-web pisauridae chiasmopes sp. (immature) 9 s;a;b wd sheet-web salticidae heliophanus sp. (immature) 5 b w leaves natta sp. 18 l w soil portia sp. 4 l w litter thyene ogdeni peckham & peckham, 1903 24 b w leaves bionidae (n = 66) representing 8.3 %. the rest of the families were low in abundance and none exceeded 7 % of the total. these percentages differ from other studies done in the savanna biome, possibly because of the difference in vegetation types. during the study on the makalali game reserve the two vegetation types sampled were mixed lowveld bush and mopane bushveld, and 32 % of the specimens caught belonged to the araneidae, followed by the salticidae (18 %) and thomisidae (10 %) (whitmore et al. 2002). the study of roodeplaat dam nature reserve was done in an open savanna and tetragnathidae with 29.3 % was the most abundant family, followed by the araneidae with 22.7 % and salticidae with 21.4 % (dippenaar-schoeman et al. 1989). on the springbok flats, an open savanna but more disturbed area, lycosidae was the dominant family (30 %), followed by gnaphosidae (21 %) (van den berg et al. 2003). the thomisidae was the most species-rich family with 12 species followed by the gnaphosidae with 11 species and the araneidae with 10 species. twenty families were represented by a single species (table 1). the three most abundant species was a thomisid tmarus comellini garcia-neto (n = 82), representing 10.3 % of the total, issn 0075-6458 113 koedoe 48/2 (2005) scytodidae scytodes sp. 16 l;p w soil segestriidae ariadna sp. (immature) 3 p wd soil selenopidae anyphops sp. 12 b w tree theridiidae chorizopella tragardhi 1 b wd g u m f o o t lawrence, 1947 web dipoena sp. 4 b wd gumfootweb enoplognatha sp. 4 b wd gumfootweb episinus bilineatus simon, 1894 1 b wd gumfootweb euryopis sp. 1 b wd gumfootweb theridion sp. 10 b wd gumfootweb thomisidae heriaeus fimbriatus lawrence, 1942 10 s,p w grass misumenops rubrodecoratus millot, 1941 2 s w grass monaeses austrinus simon, 1910 3 b w tree oxytate argenteooculata (simon, 1886) 4 b w tree runcinia flavida (simon, 1881) 1 s w grass stiphropus bisigillatus lawrence, 1952 5 p w soil thomisus granulatus karsch, 1880 2 s w grass thomisus scrupeus (simon, 1886) 2 s w grass thomisus sp. (immature) 2 s w grass tmarus africanus lessert, 1919 25 s,p w grass tmarus comellini garcia-neto, 1989 82 s,p w grass xysticus lucifugus lawrence, 1937 29 b;p w soil uloboridae miagrammopes brevicaudus o.p.-cambridge, 1882 3 b;p wd vegetation zodariidae cydrela sp. (immature) 3 p w soil diores sp. (immature) 17 b;p w soil total 793 table 2 (continued) followed by the clubionid clubiona godfreyi lessert (n = 66), representing 8.3 % and an undetermined lycosid (n = 69), representing 8.7 %. neither t. comellini nor c. godfreyi have before been collected in high numbers during surveys. tmarus comellini is known from zaire and south africa and has previously been collected from the grass and tree layer. it has a fairly wide distribution throughout south africa and is known from the eastern cape, kwazulu-natal, gauteng and limpopo province. clubiona godfreyi is a species first recorded in uganda. the first record from south africa is from lajuma, soutpansberg. species dominance also differs from area to area. a quantitative survey of the herbaceous layer of coastal dune forest at richards bay, south africa showed the pisaurid charminus atomarius (lawrence) to be the most abundant species, representing 18 % of the total, followed by the salticid thyene ogdeni (peckham & peckham) with 12 % and the araneid caerostris sexcuspidata (fabricius) with 11 % (dippenaar-schoeman & wassenaar in prep.). most spiders live in a defined environment with limitations set by both physical conditions and biological factors (foelix 1982) and species can be grouped into guilds based on available information on their habitat preferences and predatory methods. a guild is a group of species that potentially compete for jointly exploited limited resources (polis & mccormick 1986). for the present study two main guilds were recognised, namely wandering spiders (w) and web builders (wb), with further subdivisions based on microhabitat and web structure (dippenaarschoeman et al. 2005). twelve of the collected families (16.1 %) are web dwellers. three families namely araneidae, deinopidae and uloboridae, construct orb-webs or adapted orb-webs while the amaurobiidae, eresidae and segestriidae build retreat-webs usually low in vegetation, the hahniidae, linyphiidae and pisauridae construct sheetwebs, the pholcidae space-webs, the theridiidae gumfoot-webs and the agelenidae funnel-webs. seventeen of the collected families (83.9 %) are wanderers. of the wanderers 55.6 % (370) are soil dwellers while 44.4 % (295) were collected from plants. the lack of grass on parts of sovenga hill may have contributed to the lower number of plant dwellers collected. only one mygalomorph species, a member of the nemesiidae (wishbone trapdoor spiders) was collected. this is the first record of a lepthercus species from an area outside the eastern cape (dippenaarschoeman 2002b). studies indicated that the physical structure of the habitat could have a distinct effect on the composition of the spider community (wise 1993). vegetation not only provides the necessary support for anchoring webs but it also increases the availability of retreat space and modifies the microclimate, which could have an effect on the spiders as well as their prey. therefore, the structure of the vegetation is expected to influence the diversity of spiders found in a habitat (whitmore et. al. 2002). from a survey conducted at swartberg nature reserve in the succulent karoo biome, 76.5 % of the spiders collected over a 10-year period were wanderers and 23.5 % were web dwellers. of the wanderers 56.7 % were associated with the ground layer (dippenaar-schoeman et al. 2005). since the present study was conducted for only a short period of time (two months), numbers were too low to be able to make any meaningful comparisons amongst spiders collected from different slopes or different altitudes. conclusion in south africa there is a lack of baseline information on most invertebrate taxa. this causes a serious problem in conservation planning. with the ecology and diversity of the spider fauna of the savanna biome still poorly known, this study is a contribution to our knowledge of the geographical distribution of species. as such, it represents new distribution records for all the species recorded and specimens of some undescribed species are now available for syskoedoe 48/2 (2005) 114 issn 0075-6458 tematic research. since the present study was conducted over a relatively short period totaling only 17 sampling sessions over two months, the results may not reflect the actual abundance and species richness of sovenga hill. however, it indicates the importance to sample more than one habitat type in order to obtain a clearer picture of species richness and abundance. acknowledgements we thank the following people for assistance during the study: mrs. a. van den berg (arc-plant protection research institute), mr p. winter (south african national biodiversity institute) and from the university of limpopo (turfloop campus) mr. p. schoeman, mrs. h. du plessis, job matlebjane and mrs. r. olwagen. additionally we acknowledge both stellenbosch university and the stellenbosch university/deat/usaid cbp-ccr for sponsoring the study and the provision of a student bursary. references coddington, j.a., l.h. young & f.a. coyle. 1996. estimating spider species richness in a southern appalachian cove hardwood forest. journal of arachnology 24: 111–128. de wet, j.i. & h.j. schoonbee. 1991. the occurrence and conservation status of ceratogyrus bechuanicus and c. brachycephalus in the transvaal, south africa. koedoe 34: 69–75. dippenaar-schoeman, a.s. 2002a. status of south african arachnida fauna. proceedings of the symposium on the status of south african species organized by the endangered wildlife trust (ewt) of south africa. rosebank. 4-7 september. dippenaar-schoeman, a.s. 2002b. baboon and trapdoor spiders of southern africa: an identification manual. pretoria: agricultural research council. (plant protection research institute handbook no. 13). dippenaar-schoeman, a. s. & c. craemer. 2000. the south african national survey of arachnida. plant protection news 56: 11–12. dippenaar-schoeman, a.s. & r. jocqué. 1997. african spiders, an identification manual. pretoria: agricultural research council. (plant protection research institute handbook no. 9). dippenaar-schoeman, a.s. & a. leroy. 2003. a checklist of the spiders of the kruger national park, south africa (arachnida: araneae). koedoe 46: 91–100. dippenaar-schoeman, a.s., a.m. van den berg, & a. van den berg. 1989. species composition and relative seasonal abundance of spiders from the field and tree layers of the roodeplaat dam nature reserve. koedoe 32: 51–60. dippenaar-schoeman, a.s., a.e. van der walt, m. de jager, e. le roux & a. van den berg. 2005. the spiders of the swartberg nature reserve in south africa (arachnida: araneae). koedoe 48: 77–86. eardley, c.d. & a.s. dippenaar-schoeman. 1996. collection and preparation of material. i. collecting methods. in: uys, v.m. & r.p. urban (eds.). how to collect and preserve insects and arachnids. pretoria: agricultural research council. (plant protection research institute handbook no. 7). foelix, r.f. 1982. biology of spiders. cambridge, massachusetts: harvard university press. foord, s.h., a.s. dippenaar-schoeman & m. van der merwe. 2002. a checklist of the spider fauna of the western soutpansberg, south africa (araneae). koedoe 43: 35–43. gibbons, m.j. 1999. the taxonomic richness of south africa's marine fauna: a crisis at hand. south african journal of science 95: 8–12. new, t.r. 1999. untangling the web: spiders and the challenges of invertebrate conservation. journal of insect conservation 3: 251–256. polis, g.a. & s.j. mccormick. 1986. scorpions, spiders and solfugids: predation and competition among distantly related taxa. oecologia 71: 111–116. russell-smith, a. 1981. seasonal activity and diversity of ground-living spiders in two african savanna habitats. bulletin of the british arachnological society 5: 145–154. van den berg, a., a.s. dippenaar-schoeman, e. ueckermann, m. van jaarsveld & e. van der walt. 2003. diversity of the arachnida fauna of a savanna biome in the limpopo province. proceedings of the 4th congress of the southern african society for systematic biology. pretoria. 9 -11 july 2003. whitmore, c., t.e. crouch, r. slotow & a.s. dippenaar-schoeman. 2001. checklist of spiders (araneae) from makalali private game reserve, northern province, south africa: including a new family record. durban museum noviates 26: 10–19. whitmore, c., r. slotow, t.e. crouch & a.s. dippenaar-schoeman. 2002. diversity of spiders (araneae) in a savanna reserve, northern province, south africa. journal of arachnology 30: 344–356. wise, d.h. 1993. spiders in ecological webs. cambridge: university press. issn 0075-6458 115 koedoe 48/2 (2005) << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <feff0055007300650020006500730074006100730020006f007000630069006f006e006500730020007000610072006100200063007200650061007200200064006f00630075006d0065006e0074006f0073002000500044004600200063006f006e0020006d00610079006f00720020007200650073006f006c00750063006900f3006e00200064006500200069006d006100670065006e00200071007500650020007000650072006d006900740061006e0020006f006200740065006e0065007200200063006f007000690061007300200064006500200070007200650069006d0070007200650073006900f3006e0020006400650020006d00610079006f0072002000630061006c0069006400610064002e0020004c006f007300200064006f00630075006d0065006e0074006f00730020005000440046002000730065002000700075006500640065006e00200061006200720069007200200063006f006e0020004100630072006f00620061007400200079002000520065006100640065007200200035002e003000200079002000760065007200730069006f006e0065007300200070006f00730074006500720069006f007200650073002e0020004500730074006100200063006f006e0066006900670075007200610063006900f3006e0020007200650071007500690065007200650020006c006100200069006e0063007200750073007400610063006900f3006e0020006400650020006600750065006e007400650073002e> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice article information authors: susan oberholzer1 melville saayman1 andrea saayman2 elmarie slabbert1 affiliations: 1institute for tourism and leisure studies, north-west university, potchefstroom campus, south africa 2school of economics, north-west university, potchefstroom campus, south africa correspondence to: melville saayman email: melville.saayman@nwu.ac.za postal address: private bag x6001, potchefstroom 2521, south africa keywords community; marine tourism; national parks; partial multiplier modelling; regional economy; socio-economic impact; tourism dates: received: 06 july 2009 accepted: 29 june 2010 published: 21 sept. 2010 how to cite this article: oberholzer, s., saayman, m., saayman, a. & slabbert, e., 2010, ‘the socio-economic impact of africa’s oldest marine park’, koedoe 52(1), art. #879, 9 pages. doi: 10.4102/koedoe.v52i1.879 copyright notice: © 2010. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0771 (online) the socio-economic impact of africa’s oldest marine park in this original research... open access • abstract • introduction • methods    • multiplier iteration    • community survey    • business survey    • visitors’ survey • results    • social impacts    • economic impacts • discussion • conclusion • acknowledgements • references abstract (back to top) south african national parks (sanparks) plays a major role in the tourism industry and has three primary functions, namely to conserve biodiversity, to create tourism and recreational opportunities and to build strong community relations. these parks, therefore, have a definite socio-economic impact on adjacent communities, although little is known about this impact. the main aim of this study was to determine the socio-economic impact of africa’s oldest marine park, namely tsitsikamma national park, which forms part of the newly created garden route national park. this was done by conducting three surveys during april 2008: a visitor’s survey (156 respondents), a community survey (132 respondents) and a business survey (11 respondents). we found that the park has a positive economic impact on the surrounding area and that the community exhibits a favourable attitude towards tsitsikamma national park. the results also differed when compared to similar studies conducted at other national parks in south arica and one of the main reasons for this was that the park is located in a touristic area. for a greater impact however, the park should expand its marine activities, while communication with the local community could also be improved. conservation implications: good community relations and ecotourism activities are important components of good conservation practices. this research indicates that tourism activities not only generated funds for conservation, but also benefited the local communities of tsitsikamma national park. the positive attitude of local communities makes conservation of biodiversity more sustainable. introduction (back to top) situated on the garden route in the eastern cape province of south africa, tsitsikamma national park (tnp) is africa’s oldest marine park (figure 1). tsitsikamma means ‘place of much water’ in khoi-san and incorporates the wide storms river mouth as well as 80 km of rocky coastline (including marine life and a variety of fauna and flora), spanning into the indian ocean (maree 2007:2–8). the world conservation union (iucn) appealed to governments for the establishment of marine parks and reserves during the first world conference on national parks in seattle in 1962. in response, tnp was proclaimed in december 1964 as the first national marine park in africa by the then national parks board (government gazette 1964). the original coastal park extended approximately 59 km between groot river (east) (24°12’ e, west of oubosstrand) and groot river (west) (23°34’ e, at nature’s valley) and included the areas within the region of 800 m landward and 800 m seaward of the low water mark (the horizontal distance was used and contours were ignored). in september 1983, the seaward boundary of the park between the groot river (east) and the bloukrans river (23°39’ e) was extended to three nautical miles. the remainder from bloukrans to groot river (west), was changed to half a nautical mile offshore (government gazette 1983). in december 1987, the de vasselot reserve (2561 ha) was added to the park (government gazette 1987). the park offers activities such as swimming, diving, snorkelling, hiking, short boat cruises and marine pond explorations. as a very popular ecotourism attraction in south africa, the park attracts a considerable number of tourists annually (155 762 in the year ending march 2009) and has an influence on the economic and social well-being of the surrounding community. south african national parks (sanparks) has three primary objectives, namely to conserve the biodiversity of the country, to promote community upliftment and capacity building among people living in the areas neighbouring the parks and to provide tourism and recreational outlets that allow people to experience and enjoy the wonders of the parks (streuders 2008). this research focuses on the second and third objectives as the socio-economic impact of tnp has not yet been determined. in fact, no socio-economic study has been conducted on marine parks in south africa; the reason being that it is difficult to understand and incorporate the role of local communities in park structures, even though the importance thereof has long been realised. telfer and sharpley (2008:6) refer to socio-economic development as the creation and improvement of wealth and employment, as well as an improvement in the accessibility of resources. the aim of a socio-economic study is therefore to measure socio-economic development (figure 2), usually in terms of improvements in metrics such as gross domestic product (gdp), life expectancy, employment opportunities and how these improvements impact on a community’s quality of life (anon 2006). from a socio-economic point of view, there are four major relationships to be managed, and at a later stage, determined (saayman, saayman, ferreira 2009a:1, massyn 2008:225–236; relly 2008:267–280). these relationships are reflected in figure 2 and add to the importance and relevance of conducting a socio-economic study. streuders (2008:45), ferreira (2008) and van der merwe (2008:44) highlighted that it is of utmost importance that role players seen (figure 2) are continuously in consultation with one another in order to avoid problems and conflict as well as to encourage socio-economic development, which may ultimately contribute to an improvement in the quality of life of communities, tourists and businesses. the economic contribution of a park is thus not the only important factor, but also the contribution the park makes towards the community’s quality of life (saayman & saayman 2006:3) and how these are integrated into sanparks’ three primary objectives. conservation plays a major role in the sustainability of the parks; however, parks have now become more reliant on tourism to sustain their existence; this reliability on tourism has both a positive and negative impact on the community (ferreira 2008; massyn 2008; myburgh & saayman 2002:259; streuders 2008). a higher level of community participation is needed wherein the community interacts directly with the environment and hopes to reap certain benefits from these interactions (streuders 2008:113–114). the rights of the communities need to be acknowledged, including participation in decision-making, sharing in the benefits of development (e.g. job opportunities) and an enhancement of quality of life as a result of environmental improvements (hall & richards 2000:103; spenceley 2008:1–10). streuders (2008:113–114) found that not all communities understand the relevance of a national park and that without the input of the community, park management will not be aware of community needs or know how to fulfil them. in addition, communities also need to understand the economic contribution and value of a national park. when well managed, within the socio-economic framework, tourism has various positive effects on communities, such as economic development, additional employment, the enhancement of community image, improved community–tourist relationships, cultural opportunities and increased community participation (shone & parry 2004:54). however, tiyce and dimmock (2000:223–229) indicate that negative social impacts may include transformation in community characteristics and image as well as an increase in noise pollution and overcrowding. in terms of the economic impact, money spent by tourists has a multiplier effect within the local and regional economy, whereby income generated by the park can help in funding other activities such as nature conservation. on the other hand, the job opportunities created are often poorly paid and may lead to the occurrence of leakages (i.e. skills, products and services obtained from outside the region). tnp is situated in a rural area which means that the degree of economic benefit depends on the amount of money that remains in the region. leakages, in the form of imported skills and goods, pose a serious threat to the region’s economic stability. these leakages will reduce the money remaining in the region and will greatly affect the economic benefits received by local communities. in determining the magnitude of tnp’s economic impact, determinants such as location and size of the park, the number of tourists, their length of stay and how much they spend are important because they directly influence these leakages (van der merwe 2008:49). based on these observations, the following questions need to be asked when conducting a socio-economic study: • what is the economic (monetary) value of the park, in terms of what tourists and park management spend? • how many jobs are created by the park? • what is the multiplier effect – in other words, how does the money spent in this particular region circulate through the region in order to create output and income for the community? • what are communities’ perceptions of the park and do they benefit from the park in general? inevitably, local communities experience social and cultural changes ascribed to tourism development. as quoted by telfer and sharpley (2008:199), previous studies by dogan (1989), ap and crompton (1993), costa and feronne (1995), williams and lawson (2001), kuvan and akan (2005) and spenceley (2008) found that local communities’ attitudes towards tourists range from one of welcoming to one of disregard. the studies investigated how members of small communities perceive tourism, as well as which issues they have identified as being important such as personal values. most found that tourism was regarded in a positive light and that community members exhibit a favourable attitude towards tourism development. however, it was also found that community members were concerned about the negative impacts of tourism. green (2005) and simpson (2008) found that changes within the community as a result of tourism were perceived as negative and that the focus of future tourism development should fall on sustainable development. community involvement was identified as a key success factor for advantageous tourism development. samuelsson and stage (2007), relly (2008), suich, busch and barbancho (2005), mahony and van zyl (2002) and gössling (2001) conducted studies concerning the economic impact of tourism on the community. their findings indicated that tourism had a positive effect on local communities by improving their livelihood. according to their results, small-scale high-value tourism is important in maximising a long term stable income.therefore, the local communities need tourism and tourism needs the local communities’ support in order to be sustainable. other socio-economic studies have been conducted, including investigations into the okavango delta in botswana (mbaiwa 2003, 2004, 2005), a socio-economic impact study of hunting in the northern cape province of south africa (van der merwe, saayman and rossouw 2009), the klein karoo national arts festival (van der merwe 2008), turco et al.’s (2003) investigations into the socio-economic impact of sport tourism in durban, as well as a study by sims-castley et al. (2005), which determined the significance of an ecotourism-based private game reserve in the eastern cape province of south africa. according to these studies, tourism has contributed positively towards the development of communication facilities and infrastructure and has increased tourist-flow. therefore, for successful and sustainable tourism development to occur, it is important to focus on how tourism impacts on social equity, economic efficiency and on ecological sustainability of local communities. loader (1994:143) and pelser (2003:164) state that in south africa social involvement is an integral part of conservation. in the case of the residents of the kepulauan seribu marine park in del este in the dominican republic, macleod (2001:221), fauzi and buchary (2002:167) suggest that poverty should be improved; residents should be the main concern and park management should be based on the agreement and involvement of all stakeholders. the problem with most of these studies is that, firstly, their methodologies differed significantly thereby making it difficult to compare their results. a reason for this is that researchers attempt this kind of research from different perspectives, including: sociology, investment, economics, conservation, pro-poor tourism, responsible tourism, ecotourism and sustainable tourism, according to their different fields of expertise and interest. secondly, the fact that the study areas (i.e. products) differed had significant implications for each set of results; some studies were conducted in wetlands, in national parks, in game farms and in private parks. the importance of these results lies in the aims and objectives of these different offerings (i.e. products). thirdly, few studies had specific multipliers; relly (2008) states that determining multipliers could provide an indication in terms of the impact of a park on the poor. finally, ferreira (2008) states that most so called ‘socio-economic studies’ are, in fact, economic analyses, with little or no social impact analyses other than a reference to employment. again, ferreira (2008) supports the notion that socio-economic studies have to be more comprehensive. only two socio-economic studies have been conducted at national parks in south africa, namely karoo national park (saayman et al. 2009a) and addo elephant national park (saayman & saayman 2006). both of these studies found that these parks had a positive socio-economic impact, although their results differed significantly. these differences were ascribed to aspects such as the size of the park, type of activities available, number of tourists, variety of animals and plants found in the park, involvement of the community in the park management and levels of poverty in communities adjacent to the park. the question remains whether a marine park will have different results and what lessons can be learnt from the socio-economic impact of a marine park. therefore, results from this study could assist sanparks in achieving its objectives more efficiently, as well as contribute to the existing knowledge of socio-economic studies in national parks. the aim of this research was to estimate the socio-economic contribution of tnp to the surrounding communities which involved estimating the economic impact of tnp on the local economy, estimating the impact of tourism business development in the region along with how these businesses and communities benefit from the park and finally, determining the employment generated as a result of the tourism activity generated by the park. figure 1: location of tsitsikamma national park figure 2: conceptual framework for socio-economic research in national parks methods (back to top) when conducting socio-economic studies, three controversial issues need to be addressed (gelan 2003), namely, (1) which monetary spending to include, (2) designating the study area, and (3) determining what multipliers to use. for this study, we assessed both park expenditure and expenditure by visitors, in order to determine the economic value of tnp to the region. secondly, we defined the local economy as the 20-km radius around tnp, which meant that the study area included the communities of nature’s valley and storms river village. storms river village is a fairly poor community, with 25% of the local residents being unemployed and 11% being pensioners or self-employed. nature’s valley, however, is a fairly well-off community, although most of the houses are holiday or second homes. three surveys were conducted to achieve the goal of this study, (1) a community survey among the local residents of nature’s valley and storms river village to measure the social impacts, (2) a business survey, including permanent local businesses in and around the park and (3) a visitor survey of tourists going to the park in order to measure the economic impacts that increase as a result of the park’s presence in the region. partial multipliers were derived through a process of iteration to determine the economic impact of tnp. vaughan, farr and slee (2000) proposed proportional multiplier analysis in determining the local economic impact of visitors' spending on a rural national park. two methods can be used to derive these multipliers, namely a process of iteration or a partial input-output model. because of a limited number of firms in the study area, a process of iteration was used where the spending is traced through the local economy as tourists buy goods and firms buy stock from suppliers and pay their employees. the multipliers thus represent the change in sales and income that results from tourist spending. multiplier iteration the multiplier iteration procedure can occur in a number of ways. one example is when a tourist purchases accommodation services from a hotel, guest house or lodge. the owner of the accommodation service uses the money received to pay for various expenses, including goods such as soap, towels and breakfast provisions. the portion spent on goods is then traced to the providers of these goods (e.g. supermarkets). supermarkets use the received money to make certain payments (e.g. inventory). this portion of the money is then traced towards the providers of inventory (i.e. wholesalers and producers), who use the funds received to pay their own expenses. as an example, producers would use a portion of the money to pay for raw materials, a payment which can then be traced to the provider of those raw materials. the sum of these payments to the various role players constitute the increase in total output due to the purchase of accommodation by the tourist and thus the output multiplier is derived by dividing the sum of these payments by the original payment made by the tourist. this process requires detailed information from businesses, which was provided by the business survey.six postgraduate students assisted in distributing the questionnaires to the selected groups between 25 and 29 march 2008. these field workers distributed the questionnaires and collected the completed questionnaires later in the evening. the data were then coded and captured on microsoft excel 2007, and the analyses were done using both spss v.15 and microsoft excel. descriptive analyses were conducted in order to provide preliminary insights into the nature of the responses obtained, as reflected in the distribution of the values of each variable of interest (tustin et al. 2005:341). community survey community data were obtained by means of a structured questionnaire based on the social impact measuring instrument developed by fredline, deery and jago (2003:29). the reason for using this instrument is twofold: firstly, it is frequently used to determine the social impact of tourism on a community and, secondly, it allows for easier comparison of results from different studies. this instrument was adapted from a festival-based study for which the instrument was originally designed according to the needs of this nature-based product. residents’ perceptions (i.e. the main variables for the purpose of this study) were measured using a 3-point or 4-point likert scale. the scales used by fredline et al. (2003) were retained and independent variables, such as community attachment, participation and length of stay, were also measured by means of closed-ended questions. there are approximately 360 households (with 91 permanent households) in nature’s valley and 470 in storms river village and can therefore be considered small towns. this led to the decision to include all households in the sample frame. the initial part of the study was based on probability sampling, where each element in the population had a known, non-zero probability of being included in the sample (tustin et al. 2005:344). availability sampling was therefore implemented and residents who were willing and available to complete the questionnaire formed part of the final sample. in total, 132 questionnaires were completed even though field workers experienced problems such as encountering inaccessible and/or vacant houses. business survey in this study, a questionnaire similar to the one used for determining the socio-economic impact of the karoo and wilderness national parks was implemented (ferreira 2008). again, availability sampling was applied and all businesses in the specified area were included in the sampling frame. however, only those who were willing and available to complete the questionnaire participated. thirty five questionnaires were distributed to businesses operating within the survey area. in total, only 11 questionnaires were completed and used for our study. visitors’ survey the visitors’ survey was based on a questionnaire that has been used since 2001 in various national parks. questions focused on the demographic aspects and expenditure of the park visitors in order to obtain the socio-economic information needed to conduct this study. a convenience sampling was drawn from the visitors to tnp between 25 and 29 march 2008. field workers distributed the questionnaires among overnight visitors and 156 questionnaires were completed for the purpose of this study: 80 by visitors in campsites and 76 by visitors in chalets. the number of questionnaires was deemed representative of the total population seeing that the surveys conducted since 2001 were done in different months and therefore different seasons. saayman and fouche (2007:26) also indicated that the profile of the visitors to tnp had remained consistent since 2001 and that the sample could be seen as representative. results (back to top) the results are divided into two sections and discussed below. the first section covers the social impact results that were obtained through the community survey and the second section indicates the economic impact results that were obtained from both the business and visitor surveys. social impacts sustainable socio-economic development is influenced by community perceptions. it is important that the community members of nature’s valley and storms river village respond positively towards tnp and the management thereof. statements were rated by using a 7-point likert scale where 1 is very negative and 7 is very positive (table 1). respondents indicated that tnp has a very positive impact on the community of nature’s valley and storms river village (68%). tnp also affects the community’s personal quality of life in a very positive manner (65%). when considering specific perceptions (table 2) of the community, respondents indicated that (i.e. the sum of the percentage of respondents who chose 3 and 4 on the likert scale) tnp contributes to the increased number of tourists visiting the area (81%), sustains the environment (80%) and conserves the natural resources (80%). in contrast, however, community members feel that, because of tnp’s presence, the prices of some goods and services have increased (76%) and that less parking is available during high season (71%). in general, tnp is seen as contributing to the development of the area and community members consider the park as an asset. however, it is imperative to get community members more involved in park management and park management should continue to maintain positive relationships with these communities. table 1: the influence of the tsitsikamma national park on the community table 2: perceptions concerning tsitsikamma national park economic impacts visitors to tnp can be grouped into two broad categories, namely day visitors and overnight visitors. seeing that the park offers both camping facilities and chalets, the overnight visitors are divided further into campers and visitors who overnight in chalets. the magnitude of economic impact depends on (1) the number of visitors to the park, (2) the magnitude of their spending while visiting the park and (3) the effect of their spending on the generation of additional output and income in the area surrounding the park; this occurrence is referred to as the multiplying effect. firstly, there was a steady decline in the number of visitors to tnp between 2003 (203 678 visitors) to 2006 (162 549 visitors). however, visitor numbers appear to have increased since 2007, with a total of 177 722 tourists visiting the park during 2008. this decline can mainly be ascribed to a decline in chalet unit nights sold until 2005 (declining to a low of 22 655 chalet nights sold), while 2006 showed a large decline in camping nights sold (38 890 nights, compared to the 50 876 nights in 2005). in this case, 2006 was clearly an atypical year in the park’s history. in 2008, a total of 60 236 camp person nights were sold, while 24 655 chalet unit nights were sold. the visitor survey revealed that campers visiting tnp travel in groups averaging 3.4 people, while visitors staying in chalets travel in groups averaging 3.6 people. this information is valuable because camping numbers are in ‘person nights’. this is also an interesting observation seeing that campers generally tend to travel in larger groups when visiting other national parks (see saayman, fouché & kruger 2009b). secondly, to determine the magnitude of spending by tourists the visitor survey was again conducted. spending was grouped into the selected sectors used in the iteration process. according to the survey, campers spent, on average, r2707.45 while staying at tnp. this amounted to r530.87 per group per day. spending of chalet tourists was on average r3194.80 per group, exceeding that of campers and resulted in chalet tourists spending r887.44 per day per night per group. the spending patterns of campers differed from those of visitors in chalets, with campers spending a relatively higher percentage on food and restaurants than visitors in chalets (table 3). however, for both campers and chalet visitors, spending on accommodation, food and restaurants amounted to approximately 80% of their total spending, with very little being spent on curios. transport to tnp was excluded from the analysis because it could not be ascertained where or how frequently the visitors filled their car with petrol or which transport company was used. owing to the fact that accommodation is paid to the sanparks head office in pretoria, this figure was also subtracted from the spending of tourists. the portion of accommodation spending that accrues to tnp is included in the budget of park management. approximately 74% of respondents indicated that they also spent money in the area outside tnp. the businesses most visited by the respondents were petrol stations, general dealers (i.e. retail and food), restaurants and liquor stores. sanparks provided the income and expenditure account for tnp for the year ending march 2008. according to this account, expenditure, excluding depreciation, amounted to approximately r15 million. a rough breakdown of these costs revealed that the highest spending item was personnel (56.5%), followed by operational costs (31%) and maintenance (12.5%). to allocate these cost items to the various sectors in the model, some assumptions were made, including that 90% of operational expenses did not take place in the area surrounding the park, with the remainder bought from retailers. only 50% of maintenance cost accrued to the local economy and was equally divided between retail and other service providers (such as electrical services); no finance costs accrued to the local area because the nearest bank branches are located in plettenberg bay in the western cape. as unit nights already excluded the problem associated with ‘spending per person’ versus ‘spending per group’, only the numbers of campers were divided by the average group size for campers (i.e. 3.4 people) and the average length of stay (i.e. 5.1 nights) in order to estimate the camper groups. according to this method, 3474 camper groups visited the park during 2007 and 2008. to determine the number of chalet visitors, the unit nights sold were divided by the number of days that a group stayed (i.e. 3.6 days); the result was 6849 groups or chalet visitors. the magnitude of visitor spending was calculated by multiplying the number of groups with the above estimates and the average spending per visitor group, excluding transport cost (table 3). this calculation revealed that total spending by visitors staying in chalets amounted to approximately r8.6 million, while the total spending by camping visitors amounted to approximately r4.7 million. total visitor spending during 2008 was therefore estimated at approximately r13.3 million. thirdly, the multiplier effect of the spending by visitors was determined. two methods can be employed to determine the multiplier impact on the local economy, namely iteration and matrix inversion (vaughan et al. 2000). the authors decided to use iteration, since the local economy surrounding tsitsikamma national park is limited and has no municipal, manufacturing or wholesale sectors. in terms of output, one additional chalet tourist group (change in demand) in each category created an additional output of r1400.49, indicating an output multiplier of 1.11. one additional camping group created an additional output of r1510.78, indicating an output multiplier of 1.12. the multipliers created by the iteration process are in line with those determined for addo elephant national park, but the multiplier is much smaller than that for karoo national park. the latter was expected because the towns near tnp are less diversified and industrially developed than beaufort west in the case of karoo national park. to determine the total effect on income (i.e. direct, indirect and induced), the process had to be expanded to include the effect of household spending and wages. the assumption was made that the consumption pattern of locals in the study area is similar to that of the national consumption pattern as defined in the 2004 nationwide social accounting matrix (coningarth consultants 2006). it is evident that spending directly translates into income for a business and as the business expands its production, the income of other businesses also expands. this iteration process captures the direct and indirect effects of income for businesses; however, not all income received by businesses translate into income for consumers. income is used to buy stock, to pay for operational expenses and services and to pay labourers. once these payments have been made, it is possible to ascertain whether or not profit has been made. it is integral to reach a point where profit is turned consistently in order to determine how the spending by tourists and park management can be translated into income for consumers (i.e. households) in the tsitsikamma area. the income multiplier generated by the iteration process for people holidaying in chalets was 0.57 and 0.62 for campers, indicating that one additional chalet group generated income of r716.70 for the immediate economy and one additional camping group generated income of r845.92. these multipliers are in line with those found for addo elephant national park, but they are lower than the income multipliers of karoo national park. it is interesting to note that the effect of one camping group is greater than that of tourists staying in chalets. one of the reasons might be that campers spend money on a wider variety of products and services in the local economy. while these estimates reflect the spending by visitors, they do not indicate the operational spending of tnp. if a similar process is repeated for park expenditure, the output multiplier is found to be 1.10, while the income multiplier is 0.27. these income effects now exclude the income paid to its employees by the park; therefore, the total income effect of park expenditure was adapted to include both the direct, indirect and induced income effects of park expenditure. the total impact of the park on the local economy is indicated in table 4. the number of years in which the different businesses have been operational provided a good indication of how businesses evolved in the region. isolating the influence of tnp is a non-accomplishable task, because tourist activity along the garden route (which is the park’s location) has increased significantly; the area is renowned for its scenic beauty and tourism potential. however, it is interesting to note that none of the businesses in the area is older than tnp. it is evident that the tourism service industry (especially recreational activities in the area) only started to boom in the past few years. as could be expected, many businesses indicated that they rely heavily on tourist spending (table 5) seeing that the area boasts a thriving tourism destination for both locals and foreigners. table 5 presents a breakdown of the businesses in the area that completed the business survey, as well as an indication of the employment levels and the number of permanent versus part-time personnel. as indicated in table 5, the accommodation industry is extremely reliant on labour and employs a significant number of people in the area; approximately 118 people were employed by the accommodation units surveyed. it should also be noted that, in all the cases except for one particular retailer with three employees, the workers employed lived in the area. this is encouraging because it is an indication that employment opportunities are created for the local community. on average, accommodation establishments sold 81% of their business to tourists and 19% to locals. it should be kept in mind that some of the accommodation units also offer other products (such as a restaurant or small retail outlets) which may have contributed to the relatively high percentage of turnover ascribed to the area’s locals. tourists are also the main customers of tourism and recreation services, while restaurants, food stores and retailers are reliant on both tourist and local business for their existence. this is indicated in table 5 and again highlights the importance of the tourism industry as a source of employment and income in the local economy surrounding tnp. the accommodation establishments indicated that they owed 35% of their turnover to tnp, while 12% of retail turnover was ascribed to the park. owing to a lack of additional information, we assume that the same percentages indicated in previous studies can be applied to tnp for the restaurant (7.5%) and tourism and recreation services (33.3%). if this is taken as an indication of the number of employment opportunities created because of the park, it can be concluded that the park led to the creation of 44 additional job opportunities within the businesses surveyed (i.e. presuming that a linear relationship between turnover and employment exists); tnp alone employs 86 people. this survey captured 157 beds as part of the accommodation sector, while the tourism association indicated that there were 620 beds available. thus, approximately 25% of the accommodation products were surveyed with the extrapolation of the employment results indicating that 176 additional jobs were created by tnp. including the park employment figure, total employment for the study area was 262. considering the 561 households in storms river village and nature’s valley, it can be concluded that the tourism industry, sparked by the presence of tnp, provides employment opportunities for approximately 46.7% of all the households, which confirms that the area is very reliant on tourism. table 3:the spending patterns of campers and visitors in chalets table 4: total impact of the tsitsikamma national park on the local economy table 5: breakdown of businesses in the area and employment levels discussion (back to top) from the results of this study, it becomes clear that tnp has a positive socio-economic impact on the region. the results, specifically in terms of the perceptions of the local community, confirm research by streuders (2008), fauzi and buchary (2002), macleod (2001), dogan (1989) and kuvan and akan (2005) in that the communities, in general, perceive tourism development (in this case, tnp) as positive. research by spenceley (2008) and simpson (2008) had mixed results in their respective studies concerning community participation. results from this study also indicated that the park has a positive impact on the quality of life of adjacent communities which confirmed research by macleod (2001), saayman and saayman (2006), ferreira (2008), streuders (2008), mbaiwa (2004, 2005) and saayman et al. (2009a). the newly established park forums should contribute to an even better relationship between tnp and adjacent communities. the economic analysis also shows that the park has a positive impact thereby re-confirming the findings of saayman and saayman (2006) in the addo elephant national park, as well as saayman et al. (2009a) in the karoo national park; however, tnp indicated a more substantial economic impact than the karoo national park. mbaiwa (2004, 2005), spenceley (2008) and relly (2008) also confirm that tourism and conservation areas in general have a positive economic impact and is highlighted by relly’s research in madikwe, which clearly illustrated the significant impact of tourism investment in the area (relly 2008). tnp, however, has a significant leakage: tiyce and dimmock (2000) stated that this is an aspect that has to be addressed if the region wants to take full advantage of tnp. the main reason for the high leakage is the dependence of the region on tourism and the absence of other major industries. this implies that most of the money spent in the area, immediately leaves the area when goods and services are purchased. most of the stock in the area is therefore ‘imported’ from other regions. therefore, it seems that the more dependent the regional economy becomes on tourism, the greater this leakage. the leakage in terms of immediate employment is lower because most of the employees live in the area. however, the high leakage of goods causes the positive impact on income generation and employment to be lower than it would be in a more diversified economy. an overlooked opportunity revealed by this research is the low spending on curios. perhaps the production of curious could become a joint venture between the park and the community in order to grow small industries which would assist in curbing leakages. results also showed that tnp has directly contributed to the existence of 25% of the businesses in nature’s valley and storms river village, which is higher than the 4% for karoo national park, but lower than the 35% for addo national park. however, a much higher percentage of businesses indicated that tourism activity contributes significantly to their turnover. this might be an indication that the businesses in nature’s valley and storms river village underestimate the impact of tnp on their activities. a major contradiction revealed in terms of the literature presented in this study is that a larger park and greater variety of tourist activities and species lead to an increased socio-economic impact when compared to terrestrial parks. tnp, however, is small compared to most terrestrial parks and offers few organised activities with only a small variety of species accessible to tourists; this proves that tourists are attracted to the region because of its natural beauty (especially in terms of the sea and shoreline). in addition, tourists stay longer at tnp than at the karoo or addo elephant national parks, even though these parks offer more activities and a wider variety of species. results also indicated that chalet tourists spend more than campers, but that the campers’ spending patterns affect more sectors of the economy. therefore, both types of tourists are important from a socio-economic and marketing point of view. however, tnp should offer more marine activities (such as diving, boat rides, canoeing, etc.) because this could lead to greater tourist spending which will result in a greater socio-economic impact conclusion (back to top) the purpose of this research was to determine the socio-economic impact of tnp on the local communities. from the results, it has been confirmed that tnp has a positive socio-economic impact on the communities of nature’s valley and storms river village. the positive impacts are evident in direct tourist spending, job creation, a wide variety of social indicators and business development in the area. a negative aspect, however, is high leakage from the regional economy. the tnp is therefore achieving its conservation goals, creating economic opportunities and involving the community. an aspect not covered by this research is the cost of economic opportunities that were lost to make way for the park. this would be difficult to determine and should be seen in the context of the government mandate to conserve 10% of land in south africa which includes all types of biomes; however, government’s goal has yet to be achieved. if this level of conservation is not achieved, south africa’s biodiversity will become impoverished and will necessitate the protection of more land, including marine areas. from the results, it appears that tnp has a balanced approach in terms of its three main objectives. the area is dependent on tourism, which leads to high leakages which is an aspect that requires urgent attention. however, solutions will depend on cooperation between the park, communities and businesses in the region. in this regard, the results from this study contradict some findings compared to similar research at terrestrial parks where leakages were lower. the marine attributes of tnp are a strong enough attraction to ensure that tourists stay longer than at karoo and addo elephant national parks, confirming the importance of a marine park. this study suggests that marine conservation is highly valued by communities and should therefore be maintained by ensuring a high quality of water, the creation and maintenance of natural habitats for animals as well as a pollution-free environment. acknowledgements (back to top) we thank sanparks and especially mr glenn phillips and mr joep stevens for funding, as well as information used in this study. the national research foundation is acknowledged for funding. we thank dr anna spenceley, the anonymous reviewers and the language editor for valuable comments and suggestions. references (back to top) anon., 2006, ‘section on socio-economics newsletter of the association of american law schools’, journal of law and socio-economics 27, dec. ap, j. & crompton, j.l., 1993, ‘residents’ strategies for responding to tourism impacts’, journal of travel research 32(1), 47–50. conningarth consultants, 2006, ‘disaggregated social accounting matrix for south africa’, unpublished report delivered as part of a research project commissioned by the development bank of southern africa, pretoria. costa, j. & ferrone, l., 1995, ‘socio-cultural perspectives on tourism planning and development’, international journal of contemporary hospitality management 7(7), 27–35. dogan, h.z., 1989, ‘forms of adjustment: sociocultural impacts of tourism’, annals of tourism research 16(2), 216–236. fauzi, a. & buchary, e.a., 2002, ‘a socioeconomic perspective of environmental degradation at kepulauan seribu marine national park, indonesia’, coastal management 30(2), 167–181. ferreira, m., 2008, ‘the socio-economic impact of tourism in the karoo national park’, mcom thesis, school of business management, north-west university. fredline, e., deery, m. & jago, l., 2003, ‘the development of a generic scale to measure the social impacts of events’, event management 8(1), 23–37. gelan, a., 2003, ‘local economic impacts: the british open’, annals of tourism research 30(2), 406–425. gössling, s., 2001, ‘tourism, economic transition and ecosystem degradation: interacting processes in a tanzanian coastal community’, tourism geographics, 3(4), 430-453. government gazette, 1987, viewed 28 january 2010, from http://www.sanparks.org/parks/garden_route/camps/storms_river/tourism/history.php green, r., 2005, ‘community perceptions of environmental and social change and tourism development on the island of koh samui, thailand’, journal of environmental psychology 25(1), 37–56. hall, d. & richards, g., 2000, tourism and sustainable community development. routledge, abingdon. kuvan, y. & akan, p., 2005, ‘residents’ attitudes toward general and forest-related impacts of tourism: the case of belek, antalye’, tourism management 26(5), 691–706. loader, j.a., 1994, ‘national parks and social involvement – an argument’, koedoe 37(1), 137–148. macleod, d.v.l., 2001, ‘parks or people: national parks and the case of del este, dominican republic’, progress in development studies 1(3), 221–235. mahoney, k. & van zyl, j., 2002., ‘the impacts of tourism investment on rural communities: three case studies in south africa’, development southern africa 19(1), 83–103. maree, m., 2007, ‘an investigation into the antecedents and origins of tsitsikamma national park’, honours mini dissertation, school of business management, north-west university. massyn, p.j., 2008, ‘citizen participation in the lodge sector of the okavango delta’, in a. spenceley, (ed.), responsible tourism: critical issues for conservation and development, p. 225–237, earthscan, london. mbaiwa, j.e., 2003, ‘the socio-economic and environmental impacts of tourism development on the okavango delta, north-western botswana’, journal of arid environments 54(2), 447–446. mbaiwa, j.e., 2004, ‘the socio-economic impacts and challenges of a community-based safari hunting tourism in the okawango delta, botswana’, journal of tourism studies 15(2), 37–50. mbaiwa, j.e., 2005, ‘enclave tourism and its socio-economic impacts in the okavango delta, botswana’, tourism management 26, 157–172. myburgh, e. & saayman, m., 2002, ecotourism in action: guidelines and principals, 2nd edn., institute for tourism and leisure studies, potchefstroom. pelser, a., 2003, ‘omgewingsbewaring in ‘n nuwe era: ‘n nis vir die geestesen sosiale wetenskappe’ [conservation in a new era: a niche for the human and social sciences]’, tydskrif vir geesteswetenskappe 43(3/4), 164-176. relly, p., 2008, 'madikwe game reserve, south africa investment and employment', in a. spenceley, (ed.), responsible tourism: critical issues for conservation and development, pp. 267–282, earthscan, london. saayman, m. & fouche, m., 2007, executive summary of research profiles of selected national parks in south africa, institute for tourism and leisure studies, potchefstroom. saayman, m. & saayman, a., 2006, ‘creating a framework to assess the economic contribution of national parks in south africa: the case study of the addo elephant national park’, tourism economics 12(4), 619–633. saayman, m., saayman, a. & ferreira, m., 2009a, ‘the socio-economic impact of the karoo national park’, koedoe 51(1), 1–10. saayman, m., fouche, m. & kruger, m., 2009b, an analysis of visitors to selected national parks in south africa, institute for tourism and leisure studies, potchefstroom. samuelsson, e. & stage, j., 2007 ‘the size and distribution of the economic impacts of namibian hunting tourism’, south african journal of wildlife research 37(1), 41–52. shone, a. & parry, b., 2004, successful event management, 2nd edn., thomson, london. simpson, m.c., 2008, ‘the impacts of tourism initiatives in rural livelihoods and poverty reduction in south africa: mathenjwa and mqobela’, in a. spenceley (ed.), responsible tourism: critical issues for conservation and development, pp. 239–266 earthscan, london. sims-castley, r., kerley, g.i.h., geacg, b. & langholtz, j., 2005, ‘socio-economic significance of ecotourism-based private game reserves in south africa’s eastern cape province’, parks 15(2), 6–17. spenceley, a. (ed.), 2008, responsible tourism: critical issues for conservation and development. earthscan, london. streuders, c., 2008, ‘communication efficacy of south african national parks: a case study of karoo national park’, mcom thesis, school of business management, north-west university. suich, h., busch, j. & barbancho, n., 2005, ‘economic impacts of transfrontier conservation areas: baseline of tourism in the kavango-zambezi tfca’, paper no. 4., conservation international, cape town. telfer, d.j. & sharpley, r., 2008, tourism and development in the developing world, routledge, abingdon. tiyce, m. & dimmock, k., 2000, ‘nimbin mardi grass festival: the impacts’, in j. allen, r. harris, l.k. jago & a.j. veal, (eds.), events beyond 2000: setting the agenda. proceedings of conference on event evaluation, research and education held in sydney in july 2000, pp. 222–230, australian centre for event management, sydney. turco, d.m., swart, k., bob, u. & moodley, v., 2003, ‘socio-economic impacts of sport tourism in the durban unicity, south africa’, journal of sport tourism 8(4), 223–239. tustin, d.h., ligthelm, a.a., martins, j.h. & van wyk, h.j., 2005, marketing research in practice, unisa press, pretoria. van der merwe, l.h., 2008, ‘the socio-economic impact of the klein karoo national arts festival’, mcom thesis, school of business management, north-west university. van der merwe, p., saayman, m. & rossouw, r., 2009, socio-economic impact of hunting in the northern cape province, institute for tourism and leisure studies, potchefstroom. vaughan, d.r., farr, h. & slee, r.w., 2000, ‘estimating and interpreting the local economic benefits of visitor spending: an explanation’, leisure studies 19(2), 95–118. williams, j. & lawson, r., 2001, 'community issues and resident opinions of tourism', annals of tourism research 28(2), 269-290. filelist convert a pdf file! page 1 page 2 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 filelist convert a pdf file! page 1 page 2 page 3 article information authors: peter novellie1 tineke kraaij2 affiliations: 1south african national parks, conservation services, pretoria, south africa 2south african national parks, scientific services, sedgefield, south africa correspondence to: peter novellie email: petern@sanparks.org postal address: sanparks, po box 787, pretoria 0001, south africa keywords defoliation; fynbos; grass cover; heterogeneity; renosterveld; threshold of potential concern dates: received: 29 oct. 2009 accepted: 25 mar. 2010 published: 13 july 2010 how to cite this article: novellie, p. & kraaij, t., 2010, ‘evaluation of themeda triandra as an indicator for monitoring the effects of grazing and fire in the bontebok national park’, koedoe 52(1), 5 pages. doi: 10.4102/koedoe.v52i1.977 copyright notice: © 2010. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: (print) issn: (online) evaluation of themeda triandra as an indicator for monitoring the effects of grazing and fire in the bontebok national park in this article... open access • abstract • introduction • study area • method • results • discussion • conslusion • references abstract (back to top) up until 2004, the burning regime applied in the bontebok national park was aimed at maintaining grazing conditions suitable for bontebok (damaliscus pygargus pygargus). it was, however, not suitable for maintaining plant species diversity, an increasingly urgent conservation priority for the park. accordingly, the burning regime was changed in 2004 to increase the interval between fires. a possible unintended outcome of the new burning regime is the spread of grazing lawns which is likely to be deleterious for maintaining the diversity of rare plants. red grass, themeda triandra, a species often locally abundant in areas preferred by bontebok, is potentially a good indicator of the anticipated change because, although it persists with moderate grazing, it is sensitive to localised intense grazing and will decrease on grazing lawns. to gauge the potential of this indicator, the canopy spread cover, degree of defoliation and inflorescence production of t. triandra was determined at 13 permanently marked sites in november 2005. the results are compared with a survey conducted 20 years previously (october–november 1984 and october–november 1985) using similar methodology. the results suggest that t. triandra remained abundant over the previous 20 years’ application of the prior burning regime. in 1984–1985, defoliation of t. triandra was high within 1 year after a fire but declined quickly thereafter. in 2005, the tendency for the defoliation level to decline with increasing time after a fire was still apparent, but it was much less marked than in the previous survey period. a likely cause of this was the fact that cape mountain zebra (equus zebra zebra) and red hartebeest (alcelaphus buselaphus caama) were present in 2005 but absent in 1984–1985 and these taller-grass grazers would have contributed to the use of the older veld. provided it is interpreted together with other monitoring programmes, the use of t. triandra cover and defoliation intensity appears promising as an efficient indicator of some of the potentially deleterious outcomes of the interactions between herbivory and the new burning regime. conservation implications: the conservation objectives of maintaining (1) large mammal herbivory as an ecological process and (2) plant species diversity may be difficult to reconcile with each other in the highly fragmented renosterveld and lowland fynbos ecosystems. this paper explores a rapid approach to monitoring impacts of bontebok and other grazing ungulates. introduction (back to top) the bontebok national park (bnp) was established at its current location near the town of swellendam in the western cape province in 1960 to save the bontebok (damaliscus pygargus pygargus) from extinction. in this the park was successful, allowing the bontebok to increase in numbers and strengthening its conservation status. in contrast to the successful regeneration of the bontebok, the vegetation of the bnp, which comprises renosterveld and lowland fynbos (kraaij in press), has become a conservation priority due to near-complete habitat destruction outside the park (von hase et al. 2003). in 2004, in response to the increasing need to prioritise vegetation conservation, a new fire regime was introduced (kraaij 2010). the interval between fires was extended to not less than 8 years for renosterveld vegetation and to not less than 16 years for fynbos. under the previous fire regime, the average fire return period was 5.8 years for renosterveld and 8 years for fynbos (kraaij 2010). the new fire regime for fynbos in the bnp follows general practice (kruger & bigalke 1984; van wilgen et al. 1992; van wilgen et al. in press), in that it is based on the requirements of the serotinous plants in the vegetation (these are slow-maturing plants that depend on canopy-stored seeds to regenerate after fire). under the previous fire regime, a substantial area of newly burnt veld was available every year and, because fire was the main driver of spatial variation in grazing pressure, the burning regime was successful in promoting a rotation of grazing pressure (kraaij & novellie 2010). the new regime prescribes the biennial burning of an area that is comparable in size to that which was previously burnt annually. in effect, newly burnt veld is now available only every second year. evidence from savannahs (archibald et al. 2005; archibald 2008) suggests that the new fire regime may have the unintended outcome of facilitating the spread of grazing lawns in the park. ‘grazing lawns’ is a term coined by mcnaughton (1984) to mean ‘areas created and maintained by heavy grazing with unique assemblages of grazing adapted grass species’. these areas are important determinants of spatial heterogeneity in african grassland ecosystems (archibald et al. 2005; cromsigt & olff 2008). bontebok, like the conspecific blesbok (damaliscus pygargus phillipsi), are short-grass grazers and tend to repeatedly graze localised patches (du plessis 1972; grobler 1983; kraaij & novellie 2010; novellie 1987; novellie 1990). the territorial behaviour of both bontebok and blesbok results in localised accumulations of dung and urine (david 1973), which promote the formation of stoloniferous, lawn-forming grasses such as couch grass (cynodon dactylon). because of the attractiveness of newly burnt veld to grazers, fires play a role in dispersing grazing pressure, thereby preventing the formation of grazing lawns (archibald et al. 2005; archibald 2008). thus, the reduced fire frequency in bnp may lead to increased patch-selective grazing and the spread of grazing lawns. grazing lawns are a widespread natural phenomenon in grassland ecosystems (archibald et al. 2005; mcnaughton 1984; stock et al. 2010) and play an important role in creating and maintaining spatial heterogeneity and habitat diversity (cromsigt & olff 2008; novellie 1990; pickett et al. 2003). grazing lawns may be beneficial in the bnp only if their area remains within stable limits, for then grazing would be concentrated in relatively confined areas, leaving the rest of the park ungrazed or grazed at a low intensity. however, if the total area of grazing lawns were to undergo progressive expansion it would be deleterious for the threatened vegetation types in the park. certain species of red data plants in the bnp (kraaij in press; raimondo et al. 2009) (aspalathus burchelliana, a. grobleri, a. incompta, a. lebeckioides, haworthia heidelbergensis, h. marginata, h. mirabilis, stoebe rugulosa) are found in the general vicinity of intensively grazed areas, but do not occur on, or near, grazing lawns (kraaij pers. obs.). the spread of grazing lawns in the bnp may threaten species such as these, as well as plant species diversity in general, thereby constituting a threshold of potential concern (sensu biggs & rogers 2003). a monitoring system is required that is capable of reflecting a sustained increase in the area of grazing lawns. under ideal circumstances, this would justify an intensive, multi-component monitoring system. however, personnel and resources are too limited for intensive monitoring and, as such, there is a need to develop a simple indicator that could be sampled rapidly to check for the anticipated vegetation changes. in this paper we examine the potential of using a rapid cover assessment of one important grass species, red grass, themeda triandra, as an indicator. the reasons for selecting this species are as follows: • firstly, this grass species grows abundantly in those parts of the bnp favoured by bontebok and thus tends to be subjected to heavy grazing (novellie 1987). • secondly, grazing lawn formation is characterised by the replacement of bunch grasses, such as t. triandra, with grazing-tolerant stoloniferous grasses (archibald et al. 2005; cromsigt & olff 2008; novellie 1990). • thirdly, t. triandra is not easily confused with other grasses in the bnp; observers can quickly learn to recognise it even in a vegetative state. there is reason to expect that t. triandra may decline not only as a result of localised intense grazing but also as a result of very low grazing pressure in association with infrequent burning. this species is known to thrive when it is frequently burnt, but, like many grass species, it declines with prolonged absence of fire and grazing (raitt 2005; tainton 1981). this may occur particularly on the management units that are only burnt at 16-year intervals. therefore, we also examine the degree of defoliation and inflorescence production of t. triandra, on the grounds that these additional observations may be helpful in interpreting the driving factors behind observed trends in cover. study area (back to top) the 3475 ha bnp is situated 8 km from the town of swellendam (figure 1) in the western cape province (34º02’ s, 20º25’ e). the annual rainfall is 511 mm, of which 59% falls in winter. rebelo et al. (2006) classified the vegetation of the bnp as swellendam silcrete fynbos, considering it a poorly known vegetation unit that exhibits floristic features of both fynbos and renosterveld, with very high plant species richness (kraaij in press). figure 1: the bontebok national park showing the locations of the monitoring plots method (back to top) the percentage of canopy spread cover and level of defoliation were determined using the procedure of novellie (1987). however, to achieve a quick and simple indicator-based method, the procedure was adapted as follows: • instead of locating the points at 1-m intervals along a tape measure (novellie 1987), the points were located using the step-point method of evans and love (1957), as modified by mentis (1981), with the points being taken at every second step. • only ‘hits’ on t. triandra were recorded, other plant species were ignored. in both this study and that of novellie (1987), a ‘hit’ was recorded if the point fell within the perimeter of the canopy of the grass tuft. in both studies, the canopy spread cover was calculated as a ratio of hits to total points examined and was expressed as a percentage. the assessment of defoliation of individual tufts was also simplified by distinguishing only three defoliation classes, instead of the five classes of novellie (1987). the three classes were, (1) no defoliation, (2) less than 50% of shoots severed and (3) more than 50% of shoots severed. midpoint values of each class were used to calculate an index of defoliation (i.e. the mean percentage defoliation), as described by novellie (1987). the presence or absence of inflorescences was noted in order to assess the relationships between grazing, burning and seed production. novellie (1987) applied this procedure in eight plots, chosen in patches of renosterveld vegetation that were particularly favoured by bontebok. the corners of each plot were marked with steel fencing standards and its approximate location was indicated on a map of the park. novellie’s surveys were conducted during october and november in both 1984 and 1985. in this study, an attempt was made to relocate these plots, but the markers could be found for only four of them (plots 2, 4, 5 and 6 of novellie 1987). the location of an additional plot (plot 9) could be derived with reasonable certainty from the map. in november 2005, step-point surveys for t. triandra cover were conducted in the five relocated plots and in an additional eight plots that were located in patches of renosterveld favoured by bontebok. a gps reading was taken to mark the corner of each plot. it was necessary to obtain a sufficient sample of hits to determine the degree of defoliation and inflorescence production. judging subjectively that 30 would be enough, the survey within each plot was continued until at least this number of hits on t. triandra had been obtained. the number of step-points necessary to achieve this varied between 100 and 430 (table 1). the date of the last burning was known for all plots surveyed by novellie (1987) and the present study, so the results could be examined in relation to ‘veld age’ (time since last fire). table 1: canopy spread cover, defoliation index (mean percentage defoliation) and presence of inflorescences in themeda triandra, as determined from step-point surveys conducted in 13 monitoring plots in renosterveld, bontebok national park results (back to top) for the 13 plots surveyed in november 2005, the duration of sampling by a single observer of each plot varied from 13 min to 28 min (table 1); the time taken for all 13 plots was just short of 4.5 h. considering the time needed to travel from one plot to the next, it seems reasonable to conclude that the sampling of all plots could be completed by a single observer within two days’ field work. the mean percentage defoliation of t. triandra declined with increasing time since the last fire (table 1 and figure 2). this would be expected from the fact that all large herbivore species in the park tend to concentrate on areas up to 5 years after burning, whereas areas that have not been burnt for 5 years or longer are generally avoided (kraaij & novellie 2010). a small proportion of t. triandra tufts bore inflorescences within the first 2 years after burning, but thereafter inflorescence production increased (figure 3). figure 2: mean percentage defoliation of themeda triandra in relation to years after fire (2005 data) we compared the cover and mean percentage defoliation of this study with that of novellie (1987) for those plots that could be relocated (table 2). the cover values of t. triandra in 2005 were generally higher than those of 1984–1985. rainfall variability can have a major influence on grass cover (fuhlendorf et al. 2001); hence it is possible that the high cover in 2005 was due to the higher level of rainfall that had preceded these surveys (table 2). for the comparison of mean percentages of defoliation (table 2), the raw data of novellie (1987) were grouped into three defoliation classes to match the data collected in 2005. novellie (1987) reported that defoliation of all grass species was very high within 1 year after a fire but declined sharply thereafter. comparison of the mean defoliation between 1984 and 1985 (table 2) shows how this occurs for t. triandra. in contrast, the 2005 surveys did not show the same sharp decline in use with veld age; the mean defoliation of t. triandra was relatively high even several years after fire (table 2, figure 1). cape mountain zebra (equus zebra zebra) and red hartebeest (alcelaphus buselaphus caama) were relatively abundant in the park in 2005 whereas they were absent in 1984–1985. kraaij and novellie (2010) found that zebra and hartebeest do not concentrate on recently burnt veld to the same extent as the bontebok, but more readily use older veld. the presence of zebra and hartebeest in 2005 may therefore explain the higher use of t. triandra on the older veld. in drawing conclusions from table 2 it needs to be recognised that, owing to missing markers, the 2005 surveys could not be done within exactly the same areas as those of 1984 and 1985. furthermore, as noted above, the methods used to locate the points differed between this study and novellie (1987). there may also have been differences between observers; novellie was the sole observer in 1984 and 1985, whereas kraaij conducted all observations in 2005. the extent to which these factors may have influenced results is unknown. nevertheless, the results show that there was no significant decline in t. triandra over the 20 years between 1985 and 2005. figure 3: percentage of themeda triandra tufts bearing inflorescences in relation to years after fire (2005 data) table 2: canopy spread cover and mean percentage defoliation of themeda triandra compared between this study (the year 2005) and that of novellie (1987) (the years 1984 and 1985) for those of novellie’s plots which could be relocated in this study discussion (back to top) 0< i>themeda triandra was one of the dominant forage species during the 1980s, when bontebok was the only grazer in the bnp (novellie 1987) and it was still highly favoured by grazers in 2005. the results suggest that t. triandra persisted well in the bnp for 20 years under the application of the previous burning regime. this may be expected, especially in view of the fact that it is favoured by frequent burning and moderate grazing (the previous regime evidently having rotated grazing pressure, thereby reducing patch selectivity). monitoring of t. triandra cover, defoliation and inflorescence production therefore seems promising as an indicator of at least some of the anticipated outcomes of the revised fire management programme. in areas that are subjected to prolonged heavy grazing the expected trend may be: • a number of successive years with high mean defoliation (in excess of 50%), decreasing little or not at all with time since the last fire • limited inflorescence production, not increasing with time since fire • eventual decline in cover of t. triandra at the expense of lawn-forming grasses such as cynodon dactylon. at these sites we recommend examining the cover of cynodon dactylon during the step-point surveys, in addition to that of t. triandra, as the former grass will be a good indicator of grazing lawn formation. in areas that are infrequently grazed and burnt the trend may consist of a number of successive years with low mean defoliation of t. triandra tufts, which show an accumulation of old leaves and inflorescences, as well as an eventual decline in t. triandra cover at the expense of woody plants. by itself, t. triandra would be an insufficient indicator for management. at most it would provide an early warning of trends that should be examined in more detail. a vital requirement of the monitoring programme is to consider the use of space by the large herbivores in relation to fire. this constitutes an existing herbivore census programme, which has continued in the bnp, although with gaps, over many years (kraaij & novellie 2010). the following five additional monitoring programmes are important: 1. abundance of red data plant species (mostly woody shrubs; kraaij in press) 2. the extent and dynamics of grazing lawn formation 3. performance of the large herbivore populations, for example, the reproduction rates in bontebok (novellie 1986) 4. weather records, especially rainfall 5. incidence of planned and unplanned fires (kraaij 2010; van wilgen et al. in press). programme 1 was introduced in 2003 and is ongoing in partnership with the threatened species programme of the south african national biodiversity institute. programme 2 is currently not implemented, but is considered a priority and programmes 3–5 are currently being kept in operation. conclusion (back to top) in terms of determining the threshold of potential concern (biggs & rogers 2003), the key concern would be to establish whether the area of grazing lawns expands progressively, or whether it remains within acceptable limits from the point of view of maintaining plant species diversity. further work is needed to determine an appropriate regime for sampling the indicator that would be useful for detecting where this threshold has potentially been crossed. an option would be to extend the sample of 13 plots surveyed in this study to represent all of the management units (‘burning blocks’) in the bnp, so that trends in each unit could be examined in relation to the grazing and fire history of each unit. however, given the high variability of the indicator when compared between individual monitoring plots of the same veld age (tables 1 and 2), the sampling intensity necessary to achieve representation of all units may be prohibitively high. an alternative may be to determine the areas of highest and lowest grazing intensities and to sample the indicator over a gradient from high to low grazing intensity. it would also be worth investigating the potential use of satellite imagery to determine the area of lawn and bunch grasses (archibald et al. 2005). editor: llewellyn c. foxcroft references (back to top) archibald, s., 2008, ‘african grazing lawns – how fire, rainfall, and grazer numbers interact to affect grass community states’, journal of wildlife management 72(2), 492−501. archibald, s., bond, w.j., stock, w.d. & fairbanks, d.h.k., 2005, ‘shaping the landscape: fire-grazer interactions in an african savanna’, ecological applications 15, 96−109. biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring, and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59−80, island press, washington dc. cromsigt, p.g.m. & olff h., 2008, ‘dynamics of grazing lawn formation: an experimental test of the role of scale-dependent processes’, oikos 117, 1444−1452. david, j.h.m., 1973, ‘the behaviour of the bontebok, damaliscus dorcas dorcas, (pallas 1766), with special reference to territorial behaviour’, zeitschrift fur tierpsychologie 33, 38−107. du plessis, s.s., 1972, ‘ecology of blesbok with special reference to productivity’, wildlife monographs 30, 6–70. evans, r.a. & love, r.m., 1957, ‘the step-point method of sampling – a practical tool in range research’, journal of range management 10, 208−212. fuhlendorf, s.d., briske, d.d. & smeins, f.e., 2001, ‘herbaceous vegetation change in rangeland environments: the relative contribution of grazing and climatic variability’, applied vegetation science 4, 177−188. grobler, j.h., 1983, ‘feeding habits of the cape mountain zebra’, koedoe 26, 159−168. kraaij, t., 2010, ‘changing the fire management regime in the renosterveld and lowland fynbos of the bontebok national park’, south african journal of botany, doi:10.1016/j.sajb.2010.04.008. kraaij, t., in press, ‘bontebok national park: hotspot of plant diversity within the cape floral kingdom’, south african journal of botany. kraaij, t. & novellie, p.a., 2010, ‘habitat selection by large herbivores in relation to fire at the bontebok national park (1974–2009): the effects of management changes’, african journal of range and forage science 27(1), 21–27. kruger, f.j. & bigalke, r.c., 1984, ‘fire in fynbos’ in p.v. booysen & n.m. tainton (eds.), ecological effects of fire in south african ecosystems (ecological studies 48), pp. 67−114, springer-verlag, berlin. mcnaughton, s.j., 1984, ‘grazing lawns: animals in herds, plant form, and coevolution’, the american naturalist 124(6), 863−886. mentis, m.t., 1981, ‘evaluation of the wheel-point and step-point methods of veld condition assessment’, proceedings of the grassland society of southern africa 16, 89−94. novellie, p., 1986, ‘relationships between rainfall, population density and the size of the bontebok lamb crop in the bontebok national park’, south african journal of wildlife research 16, 39−46. novellie, p., 1987, ‘interrelationships between fire, grazing and grass cover at the bontebok national park’, koedoe 30, 1−17. novellie, p.a., 1990, ‘habitat use by indigenous grazing ungulates in relation to sward structure and veld condition’, journal of the grassland society of southern africa 7, 16−23. pickett, s.t.a., cadenasso, m.l. & benning, t.l., 2003, ‘biotic and abiotic variability as key determinants of savanna heterogeneity at multiple spatiotemporal scales’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience, ecology and management of savanna heterogeneity, pp. 22−40, island press, washington dc. raimondo, d., von staden, l., foden, w., victor, j.e., helme, n.a., turner, r.c. et al. (eds.), 2009, red list of south african plants (strelitzia 25), south african national biodiversity institute, pretoria. raitt, g., 2005, ‘the effect of management on themeda renosterveld grasslands in the groot vaders bosch conservancy’, msc thesis, conservation ecology department, university of stellenbosch. rebelo, a.g., boucher, c., helme, n., mucina, l. & rutherford, m.c., 2006, ‘fynbos biome’, in l. mucina & m.c. rutherford (eds.), the vegetation of south africa, lesotho and swaziland (strelitzia 19), pp. 158−159, south african national biodiversity institute, pretoria. stock, w.d., bond, w.j. & van de vijver, c.a.d.m., 2010, ‘herbivore and nutrient control of lawn and bunch grass distributions in a southern african savanna’, plant ecology 206(1), 15−27. tainton, n.m., 1981, ‘the effect of mowing, burning and grazing on species composition of pastures’, in n.m. tainton (ed.), veld and pasture management in south africa, pp. 265−268, shuter and shooter, pietermaritzburg. van wilgen, b.w., bond, w.j. & richardson, d.m., 1992, ‘ecosystem management’, in r.m. cowling (ed.), the ecology of fynbos. nutrients, fire and diversity, pp. 345−371, oxford university press, oxford. van wilgen, b.w., govender, n., forsyth, g.g. & kraaij, t., in press, ‘the adaptive management of fire regimes for the conservation of biodiversity: examples from south african national parks’, koedoe. von hase, a., rouget, m., maze, k. & helme, n., 2003, a fine-scale conservation plan for cape lowlands renosterveld, technical report no. ccu2/03, cape conservation unit, botanical society of south africa, cape town. 695--26 october 2009.indd [final version].indd a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe original research a rticle #695 (page number not for citation purposes) 69 the classification, mapping and description of the vegetation of the rooipoort nature reserve, northern cape, south africa abstract the need for a scientifi cally-based wildlife management plan and for more knowledge on vegetation led to an investigation into the plant ecology of the rooipoort nature reserve. the main aim of this study was therefore to classify, describe and map the vegetation of the reserve. the fl oristic data were analysed according to the braun-blanquet procedure using the bbpc suite. the data analysis resulted in the identifi cation of 15 communities that can be grouped into ten major community types. this resulted in fi ve ecology-based management units, which could assist with the compilation of an ecologically sound management plan for the reserve in order to achieve sustainable utilisation of the natural resources. the rooipoort nature reserve is one of the oldest and largest private nature reserves in south africa and as such deserves to be conserved and protected. the riverine and pan vegetation communities are considered to be endangered and are in need of special conservation and protection. conservation implication: the results suggest fi ve management units, which will assist in the compilation of an ecologically sound management plan for the rnr, in order to allow sustainable utilization of natural resources. author: hugo bezuidenhout1 affi liation: 1conservation services, south african national parks, south africa correspondence to: hugo bezuidenhout e-mail: hugob@sanparks.org postal address: south african national parks, scientifi c services, po box 110040, hadison park, kimberley, south africa, 8306 keywords: northern cape; rooipoort nature reserve; vegetation classifi cation; inventory; braun-blanquet dates: received: 02 mar. 2009 accepted: 19 may 2009 published: 17 aug. 2009 how to cite this article: bezuidenhout, h., 2009, ‘the classifi cation, mapping and description of the vegetation of the rooipoort nature reserve, northern cape, south africa’, koedoe 51(1), art. #695, 11 pages. doi: 10.4102/koedoe.v51i1.695 this article is available at: http://www.koedoe.co.za © 2009. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. vol. 51 no. 1 page 1 of 11 introduction the rooipoort nature reserve (rnr) is one of the oldest and largest private nature reserves in south africa and has a very interesting utilisation history. in 1985 it was also declared as the fourth south african natural heritage site. according to berry and crowe (1985), the properties comprising the rnr were consolidated in 1893. from 1893 until 1930 the estate was used primarily as a private hunting game reserve, but thereafter the estate was leased (1930–1947) for domestic stock farming, such as horses, goats and cattle. this management system involved over-grazing, which had a signifi cant impact on the vegetation. in 1947 the estate reverted to a hunting game reserve and since the early 1950s has been developed as a private nature reserve (berry & crowe 1985). the rnr is well known for its contribution to archaeological conservation with important bushman rock engravings, which is one of the richest sites of its kind in southern africa. the necessity for ecological classifi cation, description and mapping of the vegetation of conservation areas has been stated by bredenkamp and theron (1978), nakor (1979) and bredenkamp and bezuidenhout (1990). a sound knowledge of the ecology will contribute considerably to the establishment of an effi cient wildlife management programme and conservation policy for the rnr. being an old nature reserve, it should also serve as a long-term reference area for wider reconnaissance surveys in the northern cape region. it has considerable potential to serve as an education centre for environmental sciences and wildlife management. although studies on various ecological aspects of rnr have been conducted by berry (1991), crowe, schijf and gubb (1981) and fabricius and mentis (1990; 1992), a detailed description and mapping of the area’s vegetation has not been attempted. relatively medium and broad-scale vegetation classifi cation of the northern cape was done by gubb (1989), smit (2000), acocks (1953), low and rebelo (1996) and mucina and rutherford (2006). therefore, the main aim of this study was to classify, describe, ecologically interpret and map the vegetation of the rnr using the braun-blanquet procedure (mueller-dombois & ellenberg 1974). the study area location and regional vegetation the rnr is situated approximately 60 km west of kimberley in the northern cape province, south africa. it extends from latitude 280 30’ to 280 40’s and longitude 240 02’ to 240 25’e (figure 1). the rnr comprises 42 647 ha and is divided into an exclusive wildlife area totalling 34 500 ha and an area of approximately 8 000 ha where domestic stock (cattle) are grazed. a game-proof fence bounds the reserve on three sides and the vaal river on the fourth side (berry 1991). the vaal river stretches for about 30 km on the western border of the reserve and comprises of relatively ‘pristine’ riverine habitat. according to acocks (1988), the area can be classifi ed as kalahari thornveld invaded by karoo (veld type 17), while a small section along the banks of the vaal river consists of the false orange river broken veld (veld type 40). mucina and rutherford (2006) mapped three vegetation units, namely the kimberley thornveld (svk4) and schmidtsdrif thornveld (svk6) of the savanna biome and the highveld salt pans (azi10) as part of the inland azonal vegetation. physiography, geology, soils and land types the mountainous landscape varies from upper and lower lying valleys and drainage lines, together with steep to moderately steep mountain slopes and relatively fl at to undulating plateaux. altitude varies from 1 050 m to 1 187 m. the following topographical positions are distinguished in the rnr: crest, scarp, midslope, footslope, plain, fl oodplain, pan and riverbank. the vaal river, which is joined by the harts river just outside the rnr, forms the core of the drainage system of the area. according to berry (1991), the rnr lies at the base and east of the ghaap plateau. the rnr is underlain by various geological formations. outcrops of the andesitic lavas of the ventersdorp supergroup koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research bezuidenhout a rt ic le # 69 5 (page number not for citation purposes) 70 vol. 51 no. 1 page 2 of 11 mainly occur as rocky hills (koppies), with outliers of dolomite (transvaal sequence) also occurring in the area (spaggiari 1993). some low, flat ridges of quartzite (ventersdorp supergroup) are also found in the study area. the northern section of the rnr is mainly underlain by aeolian sand with surface limestones and sometimes by alluvial gravels of tertiary to recent age covering dwyka tillite (helgren 1979). during the 1920s relatively rich diamond deposits were found in the ancient gravel-filled watercourses of the vaal river. at this time the adjacent area (the deproclaimed vaalbos national park) was a hive of activity (bezuidenhout 1994). it produced diamonds to the value of over £300 000 in 1913 (wagner 1914). the heaps of mixed gravel still present in the area attest to the disturbance to which it was subjected. the soil type varies from deep (> 0.8 m) red-brown and yellowbrown sands (hutton and clovelly soil forms) to shallow (< 0.3 m) and stony (mispah, prieska and kimberley soil forms) while the soil on the western floodplain is moderately deep (0.3–0.8 m) and clayey (valsrivier and swartland soil forms). the banks of the vaal river consist of silt-clayey soil (oakleaf soil form) while the soil of old diamond diggings debris is very disturbed. the soil of the pans is moderately deep and very clayey with 35% clay content, and is of the arcadia, rensburg and willowbrook soil forms (soil classification working group 1991). the diversity of the study area is accentuated by the five land types, the ah, ae, dc, fb and fc land types that occur in the study area. according to the land type survey staff (1986) ‘a land type denotes an area that can be shown at 1 : 250 000 scale and that displays a marked degree of uniformity with respect to terrain form, soil pattern and climate’. the a unit refers to yellow and red apedal, freely drained soil without water tables. the ae land type refers to red, high-base status soil, of which the depth varies from 0.1 to > 0.3 m. the ah land type differs from the ae land type in that it includes yellow soil as well and is consistently deeper than 0.3 m. the dc land type indicates land where the soil has structure, such as prismacutanic, pedocutanic, vertic, melanic and red structured (land type survey staff 1986). the f unit indicates land where the dominant soil-forming processes have been rock weathering, typically giving rise to lithocutanic horizons. the fb and fc land types indicate land where lime occurs regularly (it does not need to be present in every soil) in upland and valley bottom soils. the two land types differ, with different geology types that underlie them (land type survey staff 1986). the land type concept has frequently provided a useful basis for description of vegetation (bezuidenhout 1993; breytenbach 1991; kooij 1990). climate the rainfall, mainly during summer (january to march), is erratic and can vary between as high as 700 mm per year, to lower than 300 mm per year (july to june). the average annual (july to june) rainfall for rnr based on data collected in the vaalbos national park (deproclaimed 2006), over nine years (1987–1995), was just over 400 mm per annum (crowe et al. 1981). the temperature is less erratic than the rainfall with cold winter temperatures (coldest months june to july) as low as -4 oc while the summer temperatures (warmest months december to january) are as high as 44 oc. frost occurs, with the earliest date recorded being 27 april and the latest date 23 september, while its duration can be as long as 107 days (land type survey staff 1986). methods by using 1:50 000 stereo aerial photographs, the study area was stratified into physiognomic-physiographic units. after a reconnaissance of the area a total number of 120 sample plots were located on a randomly stratified basis within the different homogeneous units identified from the aerial photographs. the number of sample plots allocated within each homogeneous unit depended on the size of the area; the larger the area the higher the number of sampling plots allocated to the unit. plot sizes were fixed on 900 m2 (30 m x 30 m) based on bezuidenhout’s (1994; 1995) recommendations. fieldwork was done between november 2006 and the end of january 2007. in each sample plot all species were recorded and the cover-abundance of each species was assessed according to the braun-blanquet cover-abundance scale (mueller-dombois & ellenberge 1974). plant species identification was done in accordance with the identification of germishuizen and meyer (2003). structural terminology is based on the work of edwards (1983). the percentage average canopy cover and height of the herbaceous layer (grasses and herbaceous plants), the shrub layer (woody species varying in height between > 0–3 m) and tree layer (woody species higher than 3 m) were also estimated. the habitat information is qualitatively described. edaphic information noted for each relevè included geology, soil forms, soil depth (shallow < 0.3 m; moderate 0.3–0.8 m; deep > 0.8 m), slope, aspect and the rockiness of the soil surface. the floristic data were analysed according to the braunblanquet procedure using the bbpc suite (bezuidenhout, biggs & bredenkamp, 1996). the numerical classification program twinspan (hill 1979), which is regarded as a successful approach for classification by several phytosociologists (cilliers 1998; mucina & van der maarel 1989) was used to derive a first approximation of the floristic data. further refinement of the classification was achieved by the braun-blanquet procedure (bredenkamp, joubert & bezuidenhout, 1989; brown & bezuidenhout 2005). using the phytosociological table and the habitat information gathered during the sampling period, the different plant communities were identified and described. the different plant communities are described according to their dominant species. dominant species are those that are most conspicuous in the community and are high in one or more of the importance values (whittaker 1978), which in this case are cover and frequency. no attempt was made to formally fix syntaxonomic names as this is normally avoided in detailed local studies (barkman, moravec & rauschert, 1986; coetzee 1983). soil nomenclature follows the classification of the soil classification working group (1991). the relatively homogeneous plant communities should form the basis for the compilation of wildlife and ecotourism management plans. benchmark type relevès were identified for the various plant communities on the grounds that these relevès (species composition and habitat) represent the particular community the best. the benchmark type relevès could be used for monitoring purposes. results classification the data analysis resulted in the identification of 15 different communities that can be grouped into ten major community units. the hierarchical classification is as follows: 1. schmidtia pappophoroides – themeda triandra grassland 2. schmidtia pappophoroides – acacia erioloba woodland 3. tarchonanthus camphoratus shrubland 3.1 eragrostis lehmanniana – tarchonanthus camphoratus shrubland 3.2 ziziphus mucronata – tarchonanthus camphoratus shrubland 4. acacia mellifera – acacia tortilis shrubland 5. acacia mellifera shrubland 5.1 tarchonanthus camphoratus – acacia mellifera shrubland 5.2 digitaria eriantha – rhigozum obovatum shrubland 5.3 heteropogon contortus – tarchonanthus camphoratus shrubland 6. diospyros lycioides woodland 6.1 diospyros lycioides – acacia karroo woodland 6.2 salsola rabieana – diospyros lycioides shrubland 7. pentzia globosa – eragrostis truncata forbland rooipoort nature reserve, northern cape, south africa original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #695 (page number not for citation purposes) 71vol. 51 no. 1 page 3 of 11 ve. figure 1 location of the rnr figure 2 vegetation map for the rnrr original research bezuidenhout koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 69 5 (page number not for citation purposes) 72 vol. 51 no. 1 page 4 of 11 8. eragrostis bicolor grassland 8.1 salsola rabieana – eragrostis bicolor grassland 8.2 osteospermum species – eragrostis bicolor grassland 9. cynodon dactylon – sporobolus ioclados grassland 10. scirpus species – diplachne fusca grassland the following four major soil type-cum-habitats were identified as part of the associated environmental descriptions of plant communities: a. deep, well-drained, red-brown or yellow-brown sandy soil b. shallow, well-drained, yellow-brown or red-brown rocky soil c. moderately deep, moderately drained, dark silt-clayey soil d. moderately deep, poorly drained, dark clayey soil description of the plant communities a. deep, well-drained, red-brown or yellow-brown sandy soil 1. schmidtia pappophoroides – themeda triandra grassland benchmark type: relevè 22 the schmidtia pappophoroides – themeda triandra grassland occurs on the plain, mainly in the northern part of the study area (figure 2) and is strongly associated with deep (> 0.8 m) to moderately deep (0.3–0.8 m) well-drained, yellow-brown or red-brown sandy (clay content < 10%) soil. the community is 2 726 ha in size and is underlain by aeolian sand covering dwyka tillite. surface limestone occurs sporadically. no rocks or stones on the soil surface were recorded in the relevès of this community. the dominant soil forms are hutton and clovelly, but the mispah form can also occur. this grassland is associated with the ah land type. this community is characterised by species group a (table 1) and the diagnostic species are the grasses themeda triandra and pogonarthia squarrosa and the forbs elephanthorrhiza elephantina, rhynchosia nervosa and plinthus sericeus and the geophyte moraea verecunda. the poorly developed tree stratum is 5 m tall, with a canopy cover of 1%. sparsely distributed acacia erioloba and acacia tortilis are the prominent trees in this community. the shrub stratum, with species such as grewia flava and tarchonanthus camphoratus, is poorly developed with a canopy cover of 1% and a height of 2 m. the herbaceous stratum is well developed, about 0.3 m tall with a canopy cover of 68%. the dominant grasses are schmidtia pappophoroides, eragrostis lehmanniana, stipagrostis uniplumis and aristida congesta, while the most prominent forb is hermannia tomentosa (table 1 online supplement). 2. schmidtia pappophoroides – acacia erioloba woodland benchmark type: relevè 28 the schmidtia pappophoroides – acacia erioloba woodland is strongly associated with very deep (> 0.8 m) well-drained, yellow-brown or red-brown sandy (clay content < 10%) soil and it occurs on the plain, mainly in the northern part of the study area (figure 2). the community is 2 388 ha in size and is 2. schmidtia pappophoroides – acacia erioloba woodland benchmark type: relevè 28 1. schmidtia pappophoroides – themeda triandra grassland benchmark type: relevè 22 3.2 ziziphus mucronata – tarchonanthus camphoratus shrubland benchmark type: relevè 57 3.1 eragrostis lehmanniana – tarchonanthus camphoratus shrubland benchmark type: relevè 23 rooipoort nature reserve, northern cape, south africa original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #695 (page number not for citation purposes) 73vol. 51 no. 1 page 5 of 11 underlain by aeolian sand covering dwyka tillite. no rocks or stones on the soil surface were recorded in the relevès of this community. the dominant soil forms are hutton and clovelly. this woodland is associated with the ah and ae land types. this community is characterised by species group b (table 1) and the diagnostic species are the tree acacia erioloba, the grasses aristida stipitata and eragrostis pallens and the forbs indigofera daleoides and pollichia campestris. the tree stratum is 8 m tall, with a canopy cover of 6%. the dominant and most prominent tree is acacia erioloba with sparsely distributed acacia tortilis in this community. the shrub stratum, with grewia flava, with a canopy cover of 13% and a height of 2 m, is prominent in this community. the herbaceous stratum is well developed, about 0.2 m tall with a canopy cover of 55%. the dominant grasses apart from the diagnostic grasses are schmidtia pappophoroides, 5.3 heteropogon contortus – tarchonanthus camphoratus shrubland benchmark type: relevè 44 5.2 digitaria eriantha – rhigozum obovatum shrubland benchmark type: relevè 109 5. acacia mellifera shrubland4. acacia mellifera – acacia tortilis shrubland benchmark type: relevè 105 6. diospyros lycioides woodland 6.1 diospyros lycioides – acacia karroo woodland benchmark type: relevè 95 5.1 tarchonanthus camphoratus – acacia mellifera shrubland benchmark type: relevè 76 original research bezuidenhout koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 69 5 (page number not for citation purposes) 74 vol. 51 no. 1 page 6 of 11 eragrostis lehmanniana, stipagrostis uniplumis and aristida congesta subsp. congesta, while the most prominent forb is hermannia tomentosa (table 1). 3. tarchonanthus camphoratus shrubland this shrubland, consisting of two sub-communities, is found throughout the park and is the largest in size, namely 15 088 ha (figure 2). the tarchonanthus camphoratus shrubland occurs on moderately deep, well-drained, yellow-brown or red-brown sandy soil and is associated with midslopes and footslopes of the study area. the parent material of the soil is either aeolian sand of tertiary to recent age or soil that was formed in situ. sporadic outcrops of limestone or andesitic lava may occur. the soil surface that is covered by rocks or stones varies from 2 to 60%. the shrubland is characterised by species group d (table 1) and the diagnostic species are the tree ziziphus mucronata and the grass species cymbopogon plurinodis. 3.1 eragrostis lehmanniana – tarchonanthus camphoratus shrubland benchmark type: relevè 23 this midslope eragrostis lehmanniana – tarchonanthus camphoratus shrubland sub-community is associated with moderately deep (0.3–0.8 m), well-drained sandy soil. the parent material of the soil is aeolian sand of tertiary to recent age. sporadic outcrops of limestone or andesitic lava may occur. rocks or stones cover less than 2% of the soil surface. the shrubland is associated with the ae land type. the dominant soil forms are the hutton and kimberley forms. the size of the sub-community is 9 126 ha (figure 2). the diagnostic species is the conspicuous shrub rhus ciliata (species group e; table 1). the tall (average of 6 m) acacia 6.2 salsola rabieana – diospyros lycioides shrubland benchmark type: relevè 34 7. pentzia globosa – eragrostis truncata forbland benchmark type: relevè 52 8. eragrostis bicolor grassland 8.1 salsola rabieana – eragrostis bicolor grassland benchmark type: relevè 19 8.2 osteospermum species – eragrostis bicolor grassland benchmark type: relevè 56 9. cynodon dactylon – sporobolus ioclados grassland benchmark type: relevè 53 rooipoort nature reserve, northern cape, south africa original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #695 (page number not for citation purposes) 75vol. 51 no. 1 page 7 of 11 tortilis and ziziphus mucronata trees are prominent in the poorly developed tree stratum with a canopy cover of 2%. the welldeveloped shrub stratum, which is dominated by tarchonanthus camphoratus is 2.1 m tall with a canopy cover of 15%. two other prominent woody species in this stratum are rhus ciliata and grewia flava. in isolated patches of this sub-community, where the soil tends to become rocky and shallow, the shrub acacia mellifera is more prominently present and is co-dominant with tarchonanthus camphoratus. the herbaceous layer is 0.3 m tall and has a canopy cover of 60%. the dominant grass species are cymbopogon plurinodis, schmidtia pappophoroides, eragrostis lehmanniana and aristida congesta subsp. congesta. no dominant forbs were recorded in this sub-community. 3.2 ziziphus mucronata – tarchonanthus camphoratus shrubland benchmark type: relevè 57 this sub-community ziziphus mucronata – tarchonanthus camphoratus shrubland (size 5 962 ha) is restricted to the footslopes of the study area and is associated with the ae and fc land types. the mispah soil form is dominant with more than 30% rock cover of the soil surface. outcrops of andesitic lava and limestone were noted and the soil depth varies between 0.3 to 0.6 m. this sub-community shrubland is characterised by the presence of species from species groups d, h, l, o and p, the absence of species from species groups e, f, g and n (table 1). these species include the trees ziziphus mucronata and acacia tortilis as well as the shrubs grewia flava, ehretia rigida and tarchonanthus camphoratus, with the grasses cymbopogon plurinodis and eragrostis lehmanniana. the tree stratum is 4 m tall with a canopy cover of 2%, while the shrub stratum is 2.1 m tall with a canopy cover of 15%. the herbaceous layer is 0.3 m tall with a canopy cover of 60%. b. shallow, well-drained, yellow-brown or red-brown rocky soil 4. acacia mellifera – acacia tortilis shrubland benchmark type: relevè 105 the acacia tortilis – acacia mellifera shrubland is strongly associated with the plains and footslopes of the study area (figure 2). the size of the shrubland is 14 947 ha, which is the second largest community in rnr. the habitat of this shrubland consists of shallow (< 0.3 m), well-drained, rocky soil with andesitic lava, shale and dolomitic rocks (more than 25%) on the soil surface. this shrubland is associated with the ae and fc land types. the soil-rock complex of this shrubland consists of rock and mispah soil form. the diagnostic species for this shrubland are the tree boscia albitrunca, and the shrub phaeoptilum spinosum and asparagus cf. laricinus and the forbs barleria rigida, blepharis furcata and phyllanthus parvulus as well as the grass species aristida meridionalis (species group g; table 1). the poorly developed tree stratum is 5.3 m tall with a canopy cover of 2.3%. only two tree species are prominent in this community, namely boscia albitrunca and acacia tortilis. the well-developed shrub stratum, with dominant shrub species acacia mellifera, a. tortilis, tarchonanthus camphoratus, ehretia rigida and grewia flava, is 1.8 m tall with a canopy cover of 18%. the herbaceous layer is 0.2 m tall with a canopy cover of 38%. except for the grass species aristida meridionalis, a. congesta subsp. barbicollis and eragrostis lehmanniana no other grass or forb species are prominent in this shrubland. the physiognomy of this shrubland sometimes changes to impenetrable acacia tortilis – a. mellifera thickets. 5. acacia mellifera shrubland the acacia mellifera shrubland is strongly correlated with the mountainous areas in the study area. the total size of the community is 5 292 ha, but is distributed throughout the study area (figure 2). the soil-rock complex consists of rock and mispah soil form, with outcrops of andesitic lava and quartzite occurring in the community. rocks and stones cover more than 80% of the soil surface, while the soil is shallow and well drained. the shrubland in the study area is associated with the ae and fc land types. the community is characterised by the diagnostic species from species group i (table 1), the grass species heteropogon contortus and the fern pellaea calomelanos. three sub-communities will be described in detail. 5.1 tarchonanthus camphoratus – acacia mellifera shrubland benchmark type: relevè 76 the tarchonanthus camphoratus – acacia mellifera shrubland occurs on the footslopes and lower midslopes of the andesitic lava hills in the study area (figure 2) and is strongly associated with shallow (< 0.3 m), well-drained, yellow-brown or red-brown rocky soil. the community is 2 129 ha in size. about 80% rocks or stones were recorded on the soil surface of this community. the soil-rock complex of this shrubland consists of rock and mispah soil form. this shrubland is associated with the ae land type. this community is characterised by the presence of species group i, k, l, n, o and p and the absence of species group g, h and j (table 1). the poorly developed tree stratum is 4.7 m tall, with a canopy cover of 2%. the sparsely distributed acacia tortilis is the prominent tree in this sub-community. the shrub stratum, with species such as grewia flava, ehretia rigida, acacia mellifera and tarchonanthus camphoratus is well developed with a canopy cover of 15% and a height of 1.6 m. the herbaceous stratum is well developed, 0.4 m tall with a canopy cover of 49%. the prominent grass species are digitaria eriantha, enneapogon scoparius and heteropogon contortus, while no prominent forbs were recorded (table 1). the drainage lines in this sub-community support woody plant species such as acacia tortilis, a. mellifera and tarchonanthus camphoratus and perennial grass species themeda triandra, eragrostis superba and sporobolus fimbriatus. 5.2 digitaria eriantha – rhigozum obovatum shrubland benchmark type: relevè 109 this sub-community digitaria eriantha – rhigozum obovatum shrubland (size 2 950 ha) is restricted to the upper midslopes and plateau of the andesitic lava hills of the study area and is associated with the ae and fc land types. rock and mispah soil form are dominant in the soil-rock complex of this subcommunity, with more than 87% rock cover of the soil surface. the diagnostic plant species for this sub-community is the shrub rhigozum obovatum (species group j; table 1). the poorly developed tree stratum is dominated by acacia tortilis and is 4.5 m tall with a canopy cover of 2%. the shrub stratum is well developed and apart from the diagnostic shrub species, grewia flava, ehretia rigida, acacia mellifera and tarchonanthus camphoratus also are prominent. the shrub stratum is 1.2 m tall and the canopy cover is 12%, while the herbaceous stratum is 0.35 m tall and the canopy cover is 52%. the prominent grass species are digitaria eriantha, enneapogon scoparius and heteropogon contortus and the inconspicuous forb pellaea calomelanos. 5.3 heteropogon contortus – tarchonanthus camphoratus shrubland benchmark type: relevè 44 this heteropogon contortus – tarchonanthus camphoratus shrubland sub-community is associated with shallow (< 0.3 m), well-drained, rocky soil. this shrubland occurs on the low, flat quartzite outcrops (ventersdorp supergroup) of the study area. original research bezuidenhout koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 69 5 (page number not for citation purposes) 76 vol. 51 no. 1 page 8 of 11 rocks or stones cover more than 80% of the soil surface. the shrubland is associated with the ae land type. the rock-soil complex is dominated by rock (lithosols) and mispah soil form. the size of the sub-community is 213 ha (figure 2). the diagnostic species are the conspicuous forbs pegolettia retrofracta and gomphocarpus fruticosus (species group m; table 1). no trees were noted and although the shrub stratum is present it is poorly developed with a height of 1 m and a canopy cover of 3%. the most prominent shrub species are tarchonanthus camphoratus and acacia mellifera. the herbaceous layer is 0.4 m tall and has a canopy cover of 55%. the dominant grass species are heteropogon contortus, anthephora pubescens and eragrostis lehmanniana. no dominant forbs were recorded in this sub-community. c. moderately deep, moderately drained, dark siltclayey soil 6. diospyros lycioides woodland the woodland is closely associated with the vaal river and its associated floodplain. this dark soil of the woodland is moderately deep (0.3–0.8 m), moderately drained and silt-clayey in texture. the size of the habitat, including the river (67 ha) is 996 ha (figure 2). no or limited rocks or stones covering the soil surface was noted. it is found in the dc and ae land types of the study area. the diagnostic species for the diospyros lycioides woodland are the shrub diospyros lycioides, the shrubby forb lycium hirsutum and the grass species cynodon dactylon (species group q; table 1). the woodland is divided into two sub-communities. 6.1 diospyros lycioides – acacia karroo woodland benchmark type: relevè 95 the diospyros lycioides – acacia karroo woodland sub-community is strongly associated with the moderately drained, moderately deep (0.3–0.8 m), silt-clayey, alluvial soil (clay content >10%) of the vaal river (figure 2). no rocks or stones were recorded on the soil surface. this sub-community associates strongly with the vaal river system and can be related to the dc land type in the study area. the dominant soil form is oakleaf while others such as dundee and mispah are also present. the size of the subcommunity is 329 ha. the tree stratum is well developed and is 9 m tall with a canopy cover of 30%, while the shrub stratum is 3 m tall with a canopy cover of 18%. the diagnostic woody species are the trees acacia karroo, combretum erythrophyllum, rhus lancea and salix mucronata (species group r; table 1). other woody species present in this community are the tree ziziphus mucronata and the shrub diospyros lycioides. the herbaceous layer, which is very disturbed and poorly developed, is 0.2 m tall with a canopy cover of 16%. the prominent grasses are setaria verticillata and cynodon dactylon. the prominent diagnostic forb atriplex semibaccata and the alien species, argemone ochroleuca and datura stramonium, also occur. the diagnostic common reed phragmites australis is fairly abundant (species group r; table 1). 6.2 salsola rabieana – diospyros lycioides shrubland benchmark type: relevè 34 this shrubland sub-community is restricted to the floodplain adjacent to the vaal river, with one isolated floodplain exception near hoffman’s pan (figure 2). it is strongly associated with moderately drained, moderately deep (0.3–0.8 m), silt-clayey (clay content > 10%) soil. the parent material is alluvium. stones or rocks cover less than 2% of the soil surface. this salsola rabieana – diospyros lycioides shrubland sub-community is situated in the dc and ae land types of the study area. the dominant soil form is the valsrivier form, while other soil forms such as mispah and glenrosa are also present. the size of the sub-community is 600 ha. two grass species, chloris virgata and panicum coloratum, and one forb, salsola rabieana, are the diagnostic species for this subcommunity (species group s; table 1). the herbaceous layer is well developed with a canopy cover of 44% and is 0.4 m tall. the diagnostic grass species are eragrostis lehmanniana and aristida congesta subsp. barbicollis and the forb pentzia globosa are prominent in this sub-community. the tree and shrub strata, which occur on the edge of the floodplain, has a tree stratum that is 7 m tall and a canopy cover of 6%, while the shrub stratum is 2.3 m tall and has a canopy cover of 12%. the prominent woody species are the shrub/tree acacia tortilis, tree/shrub a. karroo, the tree/shrub ziziphus mucronata and the shrub diospyros lycioides. d. moderately deep, poorly drained, dark clayey soil 7. pentzia globosa – eragrostis truncata forbland benchmark type: relevè 52 the pentzia globosa – eragrostis truncata forbland is strongly associated with the floodplain east of the hoffman’s pan (figure 2). the soil of the community is moderately deep (0.3–0.8 m), poorly drained with a dark clayey texture, with katspruit and mispah as the dominant soil forms. calcrete stones cover more than 55% of the soil surface. the size of these two forblands is 436 ha and it is associated with the ae land type of the study area. this community is characterised by three grass species, enneapogon desvauxii, eragrostis truncata and oropetium capense and two forbs, pentzia globosa and lycium horridum (species group t; table 1). no trees or shrubs were recorded in this community. the well-developed herbaceous stratum is 0.15 m tall and has a canopy cover of 58%. no other prominent plant species were noted in this community (table 1). 8. eragrostis bicolor grassland the eragrostis bicolor grassland occurs on moderately deep (0.3– 0.8 m), poorly drained, dark clayey soil. it is strongly associated with isolated bare pans and old cultivated lands of the study area (figure 2). the size of the community, which can be divided into two sub-communities, is 520 ha. no or limited rocks or stones covering the soil surface was noted. it is found in the ae and ah land types of the study area. the katspruit, swartland, valsrivier and mispah soil forms are dominant in both subcommunities. the diagnostic species for this grassland is the grass species eragrostis bicolor (species group u; table 1). 8.1 salsola rabieana – eragrostis bicolor grassland benchmark type: relevè 19 the salsola rabieana – eragrostis bicolor grassland occurs as three isolated pans in the eastern section of the study area and two old cultivated lands adjacent to the vaal river in the western section of the study area (figure 2). most of the time the pans are bare, but on the edge of these pans moderately deep (0.3–0.8 m) soil has accumulated and this is where the grassland has established. this links up with the habitat of the old cultivated lands with moderately deep (0.3–0.8 m), poorly drained, dark clayey soil. it is associated with the ah and ae land types of the study area and has a size of 483 ha. no rocks or stones were noted on the soil surface. no diagnostic plant species was identified, but the presence of species group u and the absence of species groups t and v (table 1) characterise this sub-community. the herbaceous rooipoort nature reserve, northern cape, south africa original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #695 (page number not for citation purposes) 36 77vol. 51 no. 1 page 9 of 11 stratum is well developed with a height of 0.25 m and a canopy cover of 60%. the prominent grass species eragrostis bicolor, chloris virgata and the forb salsola rabieana have been noted in this sub-community. individual acacia tortilis trees or shrubs are sparsely distributed on the edge of these pans and in the old cultivated lands. 8.2 osteospermum species – eragrostis bicolor grassland benchmark type: relevè 56 this sub-community is found to the north of hoffman’s pan on an isolated pan, with a size of 37 ha (figure 2). the bare pan’s dark coloured soil is moderately deep (0.3–0.8 m), poorly drained, with a clayey texture. less than 2% rocks or stones cover the soil surface of the pan. the sub-community is associated with the ae land type of the study area. the diagnostic forb is an osteospermum species (species group v; table 1) and the grass species cynodon dactylon is prominent in this pan (species group q; table 1). most of the pan is bare, but on the edge the herbaceous stratum is well developed with a height of 0.05 m and a canopy cover of 60%. no trees or shrubs have been noted. 9. cynodon dactylon – sporobolus ioclados grassland benchmark type: relevè 53 the cynodon dactylon – sporobolus ioclados grassland is restricted to hoffman’s pan (figure 2). the pan’s dark coloured soil is moderately deep (0.3–0.8 m), poorly drained, with a clayey texture. the pan is associated with the ae land type and its size is 214 ha. calcrete rocks and stones cover 5% of the soil surface. micro-habitats have been noted (berry 1991), but for this study a more common broad vegetation-cum-habitat description is being given. the dominant soil forms are katspruit, swartland and mispah. the only diagnostic species for this community is the grass species sporobolus ioclados. no trees or shrubs have been noted, but the herbaceous stratum is 0.25 m tall and has a canopy cover of 55%. apart from the diagnostic grass species no other grass species or forbs were prominent. 10. scirpus species – diplachne fusca grassland benchmark type: relevè 17 this grassland occurs on three isolated pans on the border between the grootkolk section and the larger rnr section (figure 2). the scirpus species – diplachne fusca grassland soil is moderately deep (0.3–0.8 m) and poorly drained, with a clayey texture. the dominant soil forms are katspruit and swartland and the size is 40 ha. no rocks or stones occur on the soil surface. the grassland is associated with the ae land type. the grass species diplachne fusca and forbs vahlia capensis, falkia oblonga, helichrysum species and the sedge scirpus species are diagnostic and prominent for these grassy pan communities (species group x; table 1). no trees or shrubs were noted in these pans. the herbaceous layer is well developed and is 0.6 m tall and has a canopy cover of 88%. discussion brown (1997) describes a nature reserve as an area removed from the development stream in order to conserve and protect nature and its processes. before vegetation can be managed efficiently, relatively homogeneous vegetation communities should be identified, described and mapped, as has been done in this study. 15 plant communities have been identified and grouped into ten major communities. the plant communities are strongly correlated with four major soil type-cum-habitats. the description of the plant communities, together with the vegetation map, can serve as a basis to formulate a management programme for rnr. an understanding of the plant communities and their associated habitats is of fundamental importance for devising sound management and conservation strategies. however, the plant communities occurring in an area do not necessarily represent separate ecological management units that can be used in a management plan for a reserve, since many of these plant communities have similarities regarding species composition and habitat, and may have a complex mosaic distribution pattern. it would therefore be necessary to group these plant communities together in order to identify broad management units that can be incorporated into a management plan from where it can be managed effectively. it is important to notice that the hierarchical classification derived from the braun-blanquet procedure facilitates this grouping of plant communities into related and manageable ecological units. the combination of these management units is based on ecological principles whereby not only the vegetation, but also the abiotic component such as the topography, geology and soil type, must be taken into consideration (brown 1997). it is important to survey each of these management units in order to determine their average veld condition score and tree density (king 1989). in a preliminary exercise the following management units were identified: management unit 1: sandy plains – consisting of communities • 1, 2 and 3.1 management unit 2: rocky plains and footslopes – consisting • of communities 3.2 and 4 management unit 3: rocky hills – consisting of communities • 5.1, 5.2 and 5.3 management unit 4: river and floodplains – consisting of • communities 6.1, 6.2 and 7 management unit 5: pans – consisting of communities 8.1, • 8.2, 9 and 10 veld management can be described as the utilisation and conservation of natural veld in such a way that the activity does not adversely affect the vegetation (brown 1997). the procedure to determine and assess the veld condition of each management unit will be the second phase of the veld management plan, where the description and mapping of the vegetation communities were the first phase. the assessment of veld condition, if repeated at intervals (monitoring), may provide a descriptive measure of direction and change of the veld condition in response to current management practices (brown 1997; hobson 1989). the rich variety of the described vegetation types which inhabit some of the available habitats in the northern cape province is reflected in the rnr faunal diversity, as was also noticed by crowe et al. (1981). unfortunately, due to poor rainfall and high temperatures it was not the ideal time for a floristic survey to compile a species list for the rnr, but the time was, however, suitable for a phytosociological study. by comparing the survey with a study done in the adjacent vaalbos national park (bezuidenhout 1994), which was carried out in a relatively wet year (1993), more annual and geophytic plant species were recorded in vaalbos national park. three different pan communities were described, of which one can be sub-divided into two sub-communities. these pans are characteristic to this part of the northern cape and have the potential of a high carrying capacity (berry 1991). according to berry (1991) ‘these pans act as nutrient sumps and as a result herbaceous growth on these areas is of high quality’. during the ‘wet/good’ years game is fairly well spread over the whole study area, but during ‘dry/poor’ years game tends to concentrate on the pans, which can easily lead to overgrazing. in general, the vegetation of the study area is characterised by the presence and in some areas the dominance of the shrub tarchonanthus camphoratus and the grass species eragrostis lehmanniana. the riverine vegetation has a distinct species composition that is different to that of the pan vegetation. conclusion the objectives set for this study, namely to classify, describe, original research bezuidenhout koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 69 5 (page number not for citation purposes) vol. 51 no. 1 page 10 of 1178 interpret and map the vegetation of the rnr using the braunblanquet procedure, were attained. the braun-blanquet procedure proved to be an accurate and effective way whereby floristically defined plant communities could be classified and identified in the field. the description of the plant communities, together with the vegetation map, can serve as a basis to formulate a management programme for the rnr. it is recommended that the division of the reserve into ecologically sound management units should receive high priority. these management units should be used to determine veld condition and grazing capacity, in order to make recommendations for stocking rates and burning programmes in the reserve. special vegetation communities that are important to be conserved and protected are the riverine and the pan vegetation. if the rnr’s natural resources are not well managed and protected according to ecologically sound management plans, sustainable utilisation and conservation will not be achieved. the resultant loss of the rich flora and fauna of the rnr will have detrimental results for conservation in the northern cape province and south africa. the rnr strives to conserve and protect nature to the betterment of present and future generations. the result of this study together with present management strategies should enable the management of the rnr to be scientifically and ecologically sound. acknowledgements ms shari cade (de beers farms), mr jonathan denga (de beers farms) and mr morgan hauptfleisch (regional manager farms: de beers) are thanked for information, comments and support during the study. dr stephen holness is thanked for his assistance with the maps. references acocks, j.p.h., 1953, ‘veld types of south africa, 1st edn.’, memoirs of the botanical survey of south africa 28, 1–192. acocks, j.p.h., 1988, ‘veld types of south africa, 3rd edn.’, memoirs of the botanical survey of south africa 57, 1–146. barkman, j.j., moravec, j. & rauschert, s., 1986, ‘code of phytosociological nomenclature, 2nd edn.’, vegetatio 67, 145–195. berry, m.p.s., 1991, ‘rainfall orientated management for a multiherbivore grazing system’, dsc thesis, university of the witwatersrand. berry, m.p.s. & crowe, t.m., 1985, ‘effects of monthly and annual rainfall on game bird populations in the northern cape province, south africa’, south african journal of wildlife research 15(3), 69–76. bezuidenhout, h., 1993, ‘syntaxonomy and synecology of western transvaal grasslands’, phd dissertation, university of pretoria. bezuidenhout, h., 1994, ‘an ecological study of the major vegetation communties of the vaalbos national park, northern cape. 1. the than-droogeveldt section’, koedoe 37(2), 19–42. bezuidenhout, h., 1995, ‘an ecological study of the major vegetation communities of the vaalbos national park, northern cape. 2. the graspan-holpan section’, koedoe 38(2), 65–83. bezuidenhout, h., biggs, h.c. & bredenkamp, g.j., 1996, ‘a process supported by the utility bbpc for analysing braunblanquet data on a personal computer’, koedoe 39(1), 107–112. bredenkamp, g.j. & bezuidenhout, h., 1990, ‘the vegetation of the faan meintjes nature reserve, transvaal’, south african journal of botany 56, 54–64. bredenkamp, g.j. & theron, g.k., 1978, ‘a synecological account of the suikerbosrand nature reserve i. the phytosociology of the witwatersrand geological system’, bothalia 12, 313– 329. bredenkamp, g.j., joubert, a.f. & bezuidenhout, h., 1989, ‘a reconnaissance survey of the vegetation of the plains in the potchefstroom-fochville-parys area’, south african journal of botany 55, 199–206. breytenbach, p.j.j., 1991, ‘die fitososiologie van die villiersgrootvlei-omgewing’, msc thesis, university of pretoria. brown, l.r., 1997, ‘a plant ecological study and wildlife management plan of the borakalalo nature reserve, northwest province’, phd thesis, university of pretoria. brown, l.r. & bezuidenhout, h., 2005, ‘a vegetation description and classification of the farms ingleside and welgedacht on the mountain zebra national park, eastern cape’, koedoe 48(2), 23–42. coetzee, b.j., 1983, ‘phytosociology, vegetation structure and landscapes of the central district, kruger national park’, dissertationes botanica 69, 1–456. cilliers, s.s., 1998, ‘phytosociological studies of urban open spaces in potchefstroom, north west province, south africa’, phd thesis, potchefstroom university for christian higher education. crowe, t.m., schijf, j.c. & gubb, a.a., 1981, ‘effects of rainfall variation, fire, vegetation and habitat physiognomy on a northern cape animal community’, south african journal of wildlife research 11, 87–104. edwards, d., 1983, ‘a broad-scale structural classification of vegetation for practical purposes’, bothalia 14, 705–812. fabricius, c. & mentis, m.t., 1990, ‘seasonal habitat selection by eland in arid savanna in south africa’, south african journal of zoology 25, 238–244. fabricius, c. & mentis, m.t., 1992, ‘modelling the habitat relations of kudu in arid savanna’, south african journal of science 88(5), 280–284. germishuizen, g. & meyer, n.l. (eds.), 2003, ‘plants of southern africa: an annotated checklist’, strelitzia 14. gubb, a.a., 1989, ‘an evaluation of landsat mss data for ecological land classification’, msc thesis, university of cape town. helgren, d.m., 1979, rivers of diamonds: an alluvial history of the lower vaal basin, south africa, the university of chicago, chicago. hill, m.o., 1979, twinspan – a fortran program for arranging mul tivariate data in an ordered two-way table by classification of the individuals and attributes, cornell university, new york. hobson, f.o., 1989, ‘veld condition assessment of false karoo’, in j.e. danckwerts & w.r. teague (eds.), veld management in the eastern cape, pp. 108–113, government printers, pretoria. king, p.g., 1989, ‘integrated farm planning’, in j.e. danckwerts & w.r. teague (eds.), veld management in the eastern cape, pp. 114–130, government printers, pretoria. kooij, m.s., 1990, ‘a phytosociological study of the north-western orange free state’, msc thesis, university of pretoria. land type survey staff, 1986, ‘land types of the maps se27/20 witdraai, 2720 noenieput, 2722 kuruman, 2724 christiana, 2820 upington, 2822 postmasburg’, memoirs on the agricultural natural resources of south africa 3, 1–185. low, a.b. & rebelo, a.g. (eds.), 1996, vegetation of south africa, lesotho and swaziland, department of environmental affairs and tourism, pretoria. mucina, l. & rutherford m.c. (eds.), 2006, ‘the vegetation map of south africa, lesotho & swaziland’, strelitzia 19. mucina, l. & van der maarel, e., 1989, ‘twenty years of numerical syntaxonomy’, vegetatio 81, 1–15. mueller-dombois, d. & ellenberg, h., 1974, aims and methods of vegetation ecology, wiley, new york. national committee for nature conservation, 1979, national plan for nature conservation, department of environmental planning, pretoria. smit, j.h.l., 2000, ‘fitososiologie en veldbestuur van die oostelike kalahari doringveld’, msc thesis, university of pretoria. soil classification working group, 1991, ‘soil classification: a taxonomic system for south africa’, memoirs on the agricultural natural resources of south africa 15, 1–262. spaggiari, r.i., 1993, ‘reconstruction of the palaeo-drainage from the gravels on the farm droogeveldt 292, barkly west, rooipoort nature reserve, northern cape, south africa original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #695 (page number not for citation purposes) vol. 51 no. 1 page 11 of 11 79 northern cape province’, bsc (honours) thesis, rhodes university. wagner, p.a., 1914, the diamond fields of southern africa, struik, cape town. whittaker, r.h., 1978, ‘dominance types’, in r.h. whittaker, (ed.), classification of plant communities, pp. 65-79, junk, the hague. filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 page 14 filelist convert a pdf file! page 1 page 2 page 3 article information authors: melodie a. mcgeoch1 mbulelo dopolo1 peter novellie2 howard hendriks2 stefanie freitag3 sam ferreira3 rina grant3 judith kruger3 hugo bezuidenhout4 rodney m. randall5 wessel vermeulen6 tineke kraaij5 ian a. russell5 michael h. knight7,8 stephen holness7,8 ané oosthuizen7,8 affiliations: 1cape research centre, conservation services, steenberg, south african national parks, south africa 2conservation services, south african national parks, south africa 3savanna research unit, conservation services, skukuza, south african national parks, south africa 4savanna research unit, conservation services, kimberley, south african national parks, south africa 5garden route unit, conservation services, sedgefield, south african national parks, south africa 6garden route unit, conservation services, knysna, south african national parks, south africa 7park planning & development, conservation services, port elizabeth, south african national parks, south africa 8department of zoology, nelson mandela metropolitan university, south africa correspondence to: melodie mcgeoch email: melodiem@sanparks.org postal address: po box 216, steenberg 7947, south africa dates: received: 29 mar. 2010 accepted: 04 aug. 2010 published: 13 may 2011 how to cite this article: mcgeoch, m.a., dopolo, m., novellie, p., hendriks, h., freitag, s., ferreira, s., grant, r., kruger, j., bezuidenhout, h., randall, r.m., vermeulen, w., kraaij, t., russell, i.a., knight, m.h., holness, s. & oosthuizen, a., 2011, ‘a strategic framework for biodiversity monitoring in south african national parks’, koedoe 53(2), art. #991, 10 pages. doi:10.4102/koedoe.v53i2.991 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) a strategic framework for biodiversity monitoring in south african national parks in this essay... open access • abstract • introduction    • context • current status • sanparks’ biodiversity monitoring system    • biodiversity monitoring programmes    • resource needs, capacity building and collaboration    • evaluation, feedback and reporting cycles • conclusion • acknowledgements • references abstract (back to top) protected areas are under increasing threat from a range of external and internal pressures on biodiversity. with a primary mandate being the conservation of biodiversity, monitoring is an essential component of measuring the performance of protected areas. here we present a framework for guiding the structure and development of a biodiversity monitoring system (bms) for south african national parks (sanparks). monitoring activities in the organisation are currently unevenly distributed across parks, taxa and key concerns: they do not address the full array of biodiversity objectives, and have largely evolved in the absence of a coherent, overarching framework. the requirement for biodiversity monitoring in national parks is clearly specified in national legislation and international policy, as well as by sanparks’ own adaptive management philosophy. several approaches available for categorising the multitude of monitoring requirements were considered in the development of the bms, and 10 biodiversity monitoring programmes (bmps) were selected that provide broad coverage of higher-level biodiversity objectives of parks. a set of principles was adopted to guide the development of bmps (currently underway), and data management, resource and capacity needs will be considered during their development. it is envisaged that the bms will provide strategic direction for future investment in this core component of biodiversity conservation and management in sanparks. conservation implications: monitoring biodiversity in protected areas is essential to assessing their performance. here we provide a coordinated framework for biodiversity monitoring in south african national parks. the proposed biodiversity monitoring system addresses the broad range of park management plan derived biodiversity objectives. introduction (back to top) protected areas in many parts of the world are under increasing threat from climate change, unsustainable resource use, system impoverishment (e.g. air pollution, encroachment), habitat conversion (e.g. roads, buildings) and isolation as a result of increasingly intense land use along their borders (carey, dudley & stolton 2000; hansen & defries 2007). for example, mammal populations in african protected areas (specifically eastern and western africa) have declined significantly since the 1970s (craigie et al. 2010). although mammal population abundances have generally been stable in southern africa over the same time period (craigie et al. 2010), pressures on ecosystem services in the region continue to grow, including the use of such services for fresh water, food and fuel (van jaarsveld et al. 2005). while many of the impacts of the pressures on ecosystem services are experienced outside of protected areas, protected areas themselves are not immune to the direct and indirect consequences of environmental change. monitoring the status of, and threats to, biodiversity in protected areas is thus becoming increasingly important. the measurement and monitoring of biodiversity in protected areas is generally aimed at, (1) assessing and improving the efficiency and effectiveness of conservation action, (2) informing management action and policy at both local and national levels, (3) providing evidence of conservation success and (4) strengthening the case for conservation among policy makers, funding agencies and land owners (gaston et al. 2006). in addition, biodiversity monitoring systems in protected areas are intended to provide early recognition of unforeseen changes that impact on biodiversity, and to contribute to understanding potential impacts of current and new activities on biodiversity. these data will also feed into national and international assessments of the state of biodiversity (buckley et al. 2008). monitoring systems are thus necessary to both identify where policy or management intervention may be required, and to inform and evaluate the effectiveness of any interventions. monitoring is ‘the collection and analysis of repeated observations or measurements to evaluate changes in condition and progress toward meeting a conservation or management objective’ (elzinga et al. 2001). monitoring systems for assessing the effectiveness of the management of protected areas broadly involves two approaches, (1) quantitative, measurementbased data collection obtained from in-field measurements (or, for example, from remote sensing) of some aspect of biodiversity or ecosystem process (a quantitative approach) and (2) qualitative, perception-based scoring of performance, conducted by managers and other stakeholders (i.e., a scoring approach) (hockings 2003). both scoring and quantitative approaches are valuable in the protected area context (buckley et al. 2008). for example, the world commission on protected areas management effectiveness tracking tool is a scoring system that is widely applied to protected areas across the world (stolton et al. 2007). south african national parks (sanparks) has also developed and implemented a scoring process, that is, state of biodiversity (sob) management reporting, that is used to assess how well management targets within sanparks are being met. it is used as a broad-based, qualitative assessment of the implementation of park objectives (goodman 2003; hockings, leverington & james 2006; knight 2007). however, to date sanparks has not yet implemented a quantitative, biodiversity monitoring system that integrates biodiversity management objectives and activities within and across national parks. here we outline a framework for guiding the development and implementation of a sanparks biodiversity monitoring system (bms). in this context, the term ‘biodiversity’ includes all levels of the biological hierarchy, from genes to ecosystems, and includes structure, function and composition (gaston 1996; noss 1990). it thus encompasses quantitative monitoring of not only the elements, but also the processes and mechanisms that generate, maintain and threaten biodiversity. as such, this framework provides, (1) the rationale for developing a sanparks bms, (2) the organisational, national and international contexts within which the bms is positioned, (3) an overview of the current status of biodiversity monitoring in sanparks, (4) an overarching structure and set of objectives for the bms and (5) guiding principles and requirements for the development of monitoring programmes. stages in the development of a bms and the monitoring programmes that constitute it include scoping, design, testing and implementation phases, and the feedbacks between them (mace et al. 2005; reyers & mcgeoch 2007; the royal society 2003). the strategic framework presented here thus represents part of the scoping and design phase of a monitoring system for sanparks. context sanparks’ monitoring system must be cognisant of national and international biodiversity monitoring systems and programmes, while simultaneously focusing on the organisation’s own mandate and management objectives (reyers & mcgeoch 2007). explicitly demonstrating how the sanparks bms fits within the broader context, as far as is relevant, will ensure that monitoring retains a distinct profile within the organisation, and that it remains relevant and in line with global biodiversity monitoring standards. sanparks’ obligation to monitor is specified in the national environmental management: protected areas act (republic of south africa 2003, act 57 of 2003), in part, as: ‘the management authority of a protected area must monitor the area against the indicators set in terms of established indicators for monitoring performance’. the performance indicators relate to, (1) the management of protected areas and (2) the conservation of biodiversity in those areas. the national environmental management: biodiversity act (republic of south africa, 2004, act 10 of 2004) specifies that the requirements for biodiversity management are aimed at ensuring the long-term survival in nature of species or ecosystems and includes provision for the responsible person, organisation or organ of state to monitor and report on progress with such objectives. south africa has some history of environmental and biodiversity assessment, for example, the national state of the environment report (deat 2006), the national environmental indicators programme (deat 2002), the national state of forests report (dwaf 2007) and the south african environmental observation network (van jaarsveld et al. 2007). further, in keeping with our commitments under the convention on biological diversity (cbd), south africa has developed a national biodiversity strategy and action plan and a national biodiversity framework (deat 2005). the national biodiversity monitoring framework, together with the national spatial biodiversity assessment (driver et al. 2005) and the national protected areas expansion strategy (deat 2008), form the basis for the national biodiversity framework. sanparks’ bms aligns closely with these national initiatives, and will therefore be in a position to inform national reporting on biodiversity (grobler 2009). relevant international monitoring initiatives include, for example, those reporting on various targets set by the cbd (e.g., the 2010 biodiversity target, to achieve a significant reduction in the rate of biodiversity loss [butchart et al. 2010; walpole et al. 2009]). relating specifically to protected areas, the cbd programme of work on protected areas outlines clear goals and targets for protected areas. for example, goal 4.3 of this programme is ‘to assess and monitor protected area status and trends’ (unep-wcmc 2008). the cbd’s programme of work on marine and coastal biological diversity also provides an overview of status and trends, and highlights the 2010 sub-targets for marine and coastal biodiversity. other national protected area agencies have also adopted quantitative biodiversity monitoring systems, such as parks canada’s ecological integrity monitoring framework (timko & innes 2009). australian protected areas have clear monitoring strategies linked to management plans, although monitoring remains largely decentralised to individual parks (buckley et al. 2008). by contrast, in the united kingdom, the absence of systematically collected data in protected areas is currently considered to be a substantial hurdle to assessing their effectiveness (gaston et al. 2006). there are therefore clear international and national imperatives for monitoring the state of, and trends in, biodiversity, together with the success of management actions aimed at its conservation. as a significant role player in south africa’s biodiversity and conservation sector, sanparks’ involvement in assessing the conservation and management of biodiversity in the context of the performance of protected areas is fundamental to the purpose of the organisation (sanparks 2006a). sanparks’ strategic business is underpinned by the principles of adaptive management, that is, ‘a management philosophy that places emphasis on strong goal-setting, integration of design, management and systematic monitoring in order to adapt and learn’ (biggs & rogers 2003). as part of the adaptive management system, each park has a management plan (submitted to the minister of water and environmental affairs) that outlines a series of higher-level biodiversity objectives. these objectives provide broad foci for the rationale and design of a bms for sanparks (table 1). table 1: current, higher level biodiversity objectives common to park management plans (march 2008) that form the basis for identifying monitoring requirements. current status (back to top) monitoring is one of the pillars underpinning sanparks’ strategic adaptive management (sam) approach, where strongly goal-orientated park objective hierarchies are used to link science, monitoring and management (biggs & rogers 2003). monitoring, or ‘regular state-of-the-system measurement’, is pivotal to the success of sam (biggs & rogers 2003). sanparks has a long history of biodiversity monitoring, with several existing biodiversity monitoring projects and activities (bryden & de vos 1994; durrheim 2009). over 170 current and historical sanparks projects qualify as either monitoring projects or research projects that provide relevant baseline information for monitoring. together with sanparks scientists, external agencies and researchers have made a significant contribution to this body of work, specifically to understanding ecosystem processes, compiling species inventories and conducting species-specific studies (bryden & de vos 1994). these monitoring-relevant projects encompass a spectrum of individual researcher and organisational involvement and data ownership arrangements. in a number of instances, therefore, the data lie with external organisations and are not necessarily effectively integrated into sanparks’ knowledge systems and management processes. in addition, little biodiversity monitoring and reporting (quantitative, rather than score-based) currently occurs at either biodiversity estate (i.e., the full suite of biodiversity and ecosystems under sanparks management), regional or organisational levels (figure 1). investment in monitoring has also been unevenly distributed across environments and parks (as has research more generally, bryden & de vos 1994). in most instances monitoring projects are designed for specific parks and in a few cases, for park clusters. this, in part, reflects the history of the organisation and the adoption and proclamation of new terrestrial parks and also marine protected areas. figure 1: the sanparks biodiversity monitoring system (bms) will address objectives at several levels: organisational, biodiversity estate, regional and park-specific. existing monitoring activities have thus developed independently of an integrated organisational monitoring framework, and many of them are not necessarily well linked to park management requirements. furthermore, few existing monitoring activities are directly linked to management objectives via, for example, specific thresholds of potential concern (tpcs, see figure 1) (for tpc examples, see foxcroft 2009 [alien species]; rogers and o’keefe 2003 [rivers]; van wilgen, biggs & potgieter 1998 [fire]). time series information is also limited to selected taxa (mostly mammals) and specific parks, for example, the kruger national park (du toit 2003). the alignment between research projects and sanparks’ biodiversity objectives (table 1) is not always clear, and existing projects do not currently address the full range of biodiversity objectives in park management plans (see also buckley et al. 2008; kapos et al. 2008). for example, the rehabilitation of degraded areas is an important management objective in several parks, particularly more recently proclaimed parks (such as fynbos in plantation exit areas in the garden route national park), but it is currently largely not monitored. by contrast, sustainable indigenous forest monitoring in the garden route national park is well developed, and encompasses monitoring natural ecosystem changes and the effects of management activities (durrheim 2009). the design of a bms for sanparks must therefore be led by strategic organisational objectives, rather than by existing monitoring activities. several gaps exist, and stronger alignment with park and organisational objectives is required (buckley et al. 2008). motivation for biodiversity monitoring is significantly strengthened when diverse monitoring objectives are placed within a coherent, overarching framework, such as that provided here. in addition to the absence of a strategic monitoring framework for the organisation, the most important reason to date for not implementing the required monitoring programmes has been the severe shortage of resources, both in terms of numbers of people to carry out the fieldwork, as well as a lack of the skills required to carry out extensive biodiversity surveys and associated data analyses (field et al. 2007). the situation outlined here, while recognised as undesirable, is not uncommon to national protected area systems (e.g., australia [buckley et al. 2008] and the united kingdom [kapos et al. 2008]). nonetheless, it provides the principle motivation for the development and implementation of a bms for sanparks that addresses and prioritises the full range of key biodiversity concerns, conservation, and reporting commitments and obligations across parks, taxa and environments. sanparks’ biodiversity monitoring system (back to top) monitoring systems must be objective driven, and the rationale for a particular monitoring programme can be expressed as a series of multiple, hierarchical objectives (field et al. 2007). there are many ways of categorising monitoring objectives, for example, by taxon or by threat, by management objective, or by the scale at which the monitoring programme will operate. while there is no universally best way of doing so, grouping monitoring objectives is a critical first step in rationalising the inevitable multitude of monitoring requirements (regan et al. 2008). different approaches to categorising monitoring objectives are usually complementary, and two main approaches (described below) were used to guide the design and development of the sanparks bms and to identify biodiversity monitoring programmes (bmps) (figure 2). figure 2: sanparks’ biodiversity monitoring system (bms) is composed of a number of biodiversity monitoring programmes (bmps); the 10 biodiversity monitoring programmes shown here are those that have been identified for development and adoption. firstly, the bms was primarily designed to encompass the monitoring of key organisational biodiversity management objectives at various scales and across national parks. table 1 provides a synthesis of the range of higher-level management objectives extracted from sanparks’ park management plans (sanparks 2008). all protected areas managed by sanparks explicitly have biodiversity as a management objective, and most include biodiversity representation, process and sustainability in their objective hierarchies. the similarity of broad objectives across parks means that monitoring programmes may be identified and designed that are applicable to more than a single park. this approach has the following advantages, (1) it provides comparative information on the status of, and trends in, biodiversity across parks, (2) it provides information on the biodiversity estate and facilitates the interpretation of monitoring outcomes within a biome and regional context (figure 1) and (3) it is more efficient than repeated, parallel investment in the development of monitoring programmes for individual parks. secondly, the bms for sanparks must necessarily encompass multiple objectives at different levels and different scales. for example, the following three objectives are hierarchically related, (1) to monitor the effectiveness of sanparks in conserving rare and threatened species (organisational level objective), (2) to monitor changes in the conservation status of species of special concern in the fynbos biome (biodiversity estate/regional level objective) and (3) to monitor changes in the conservation status of bontebok in bontebok national park (park level objective) (as shown in figure 1). these levels (organisational, estate, regional and park) do not necessarily form discrete categories, but do illustrate the hierarchical nature of monitoring needs and objectives (figure 1). biodiversity monitoring programmes the sanparks bms comprises a series of bmps (figure 2). a monitoring programme encompasses one or more objectives with a common underlying rationale and usually more than a single measure or indicator (heink & kowarik 2010). together, the set of programmes (i.e. the monitoring system) addresses the broad range of sanparks’ biodiversity monitoring priorities (figure 3). figure 3: relationship between biodiversity management objectives (left) and biodiversity monitoring programmes (right). ten bmps have been identified for development and implementation. the identification and selection of these programmes took place at a workshop attended by 20 sanparks scientists in october 2009, following the development and adoption of the biodiversity monitoring framework (2008–2009, presented here) (figure 2 and table 2). after discussion of the need to group or categorise monitoring requirements, and the various models and approaches available for doing so, a list of monitoring programmes was compiled by soliciting proposals from all scientists present. this list was then examined and related proposals merged, resulting in a final list of 10 programmes (table 2). the list was then put to a vote to prioritise the programmes and potentially reduce their number. however, although there were wide-ranging views on the relative priorities of the programmes, there was consensus that, together, the 10 programmes encompass priority monitoring needs within sanparks. the number of programmes to be developed and adopted thus remained 10. table 2: biodiversity monitoring programmes (bmps, see figure 1), the rationale for their selection and the application of information generated by each programme. the 10 programmes align strongly with park management objectives and are also strongly interrelated (figure 3). for example, the freshwater and estuarine systems programme may rely on the alien and invasive species programme for alien monitoring in freshwater systems. the habitat degradation and rehabilitation programme and the habitat representation and persistence programme may work together, for example, on the impacts of adjacent land use on the park. the climate and climate change programme, in addition to its value in its own right, has the potential (via collation and reporting of weather and climate data) to support most other programmes. this interrelatedness between programmes, and the explicit integration of their design and outputs, is essential to maximise the knowledge gained and to achieve broader and more robust insights into ecosystem change (biggs 2003; buckland et al. 2005; henry et al. 2008; nielsen et al. 2009). indeed, information generated by the bms is designed to be integrated within and across both parks and programmes. such integration will not only add value to the bms, but will also streamline individual programmes and avoid duplication of effort. although one of the 10 programmes selected is environment-specific (the freshwater and estuarine system programme), terrestrial, freshwater and marine environments are integral to all programmes. similarly, most programmes are likely to encompass the monitoring of a cross-section of taxa. table 3: principles guiding the development of the sanparks biodiversity monitoring system (bms) and the biodiversity monitoring programmes (bmps) that it encompasses. each bmp will be designed to adopt, as far as possible, relevant, common approaches, methods and reporting systems across parks. information generated by bmps may be integrated (aggregated and disaggregated) and synthesised for reporting on the biodiversity status of individual parks, park clusters or all parks (table 3). the system is also flexible, such that new programmes may be identified and introduced over time. the bmps will be designed following a set of guiding principles (table 3). these principles were adopted largely to ensure the relevance, efficiency, effectiveness and quality of the bms. the programme design phase will be informed by subject and taxonomic expertise, as well as survey design, sampling design and statistical principles (e.g., buckland et al. 2005; elzinga et al. 2001; field et al. 2007; lovett et al. 2007) (table 3). programmes are intended for mediumto long-term adoption (over several 5-yearly park management plan revision cycles), and must therefore balance priority monitoring needs with logistic feasibility and sustainability. existing monitoring projects and activities will, wherever feasible and appropriate, be integrated into one or more of the 10 programmes. in addition, where monitoring approaches have been developed and tested, or are widely used elsewhere, these may be adopted. for example, well developed sets of indicators exist for marine protected areas and marine ecosystems (moloney & shillington 2007; pomeroy et al. 2005), as well as in the area of sustainable forest management (durrheim 2009; dwaf 2005). established systems such as these may, as appropriate, be adopted or modified for use in sanparks’ bms. each bmp will be formally proposed and independently reviewed prior to adoption. programme proposals will include a description of context and rationale, objectives and methods, research and inventory needs, as well as a critical assessment of the strengths and weaknesses of the programme, and the indicators and measures that underpin it. programme proposals will also include assessments of the capacity and funding requirements necessary to implement them, as well as assessments of relationships, overlaps and linkages between related programmes, and data management protocols. these bmp proposals serve several purposes, (1) they provide a means to solicit input, (2) they provide (in a single document) information that can be accessed readily and referenced, (3) they ensure continuity of implementation with staff changes and (4) they may be used to elicit support for the bms (elzinga et al. 2001). finally, effective data management and reporting is also key to the success of monitoring systems such as this, and data management protocols will be fully integrated into the development of the bms. resource needs, capacity building and collaboration the greatest challenge facing conservation-relevant monitoring is balancing the development of appropriate strategies with personnel, time and budget constraints (buckley et al. 2008; walker 2009). in fact, the monitoring that is carried out in protected areas has been shown to be more strongly correlated with resource availability than any other factor (bruner et al. 2001; wwf 2004). as outlined earlier, the assessment of resource and capacity needs will form part of the proposal development process for each bmp. although existing resources, personnel and infrastructure will be used as far as possible, it is likely that a significant injection of both human and financial resources will be required for the sanparks bms to be fully and effectively implemented. for example, the capacity for data management that is required by the bms is far from sufficient to handle data from all 19 national parks. as is commonly the case elsewhere (kapos et al. 2008), the implementation of biodiversity monitoring in sanparks is constrained by a lack of personnel tasked with and qualified to collect data, to manage and archive data, as well as to analyse and interpret data to inform management.the inadequate state of resources for biodiversity monitoring in sanparks reflects a countrywide shortage of resources, as reported in south africa’s fourth national report to the cbd (deat 2009). this report notes a significant shortfall in funds required for the implementation of south africa’s national biodiversity framework, which includes the priority actions of developing national biodiversity research and monitoring strategies, as well as strategies to address the widespread shortage of human capital in the biodiversity sector. nonetheless, as james, gaston & balmford (2001) demonstrated, the cost of investing in conservation is small in comparison with the lost opportunities and environmental degradation in the absence of such investment (e.g., declining water quality, alien species invasion, human health costs of poor air quality). biodiversity monitoring is essential to ensuring the effectiveness of this investment. a further element critical to the success of the sanparks bms is the continuation, strengthening and expansion of collaborative relationships with other agencies involved in biodiversity monitoring. this includes, for example, the south african national biodiversity institute, national departments, provincial conservation agencies, researchers and research organisations. the use of natural history societies, volunteer groups and citizen scientists in monitoring programmes has been shown to be highly effective, both internationally and in south africa, with volunteers making significant contributions in this area (bell et al. 2008; braschler et al. 2010; podjed & mursic 2008). the honorary rangers of sanparks is one such example, as are the citizen scientists that contribute to bird monitoring in south africa and globally (see, e.g. www.adu.org.za). evaluation, feedback and reporting cycles the approach that will be used to track and evaluate progress in the development and adoption of sanparks’ bms will be based on the evaluation and monitoring principles set for the organisation (sanparks 2006b). this approach adopts a logical series of steps to measure progress with the implementation of the bms. it ensures ongoing assessment of the effectiveness of the framework and its implementation, and ultimately the organisation’s mandate to enable informed and accountable decision-making through monitoring and analysis (kapos et al. 2008). as part of this process, the bms should be regularly reviewed and evaluated, as is the case with park management plans. conclusion (back to top) the biodiversity monitoring framework presented here maps the way forward for biodiversity monitoring in sanparks. as such, it is intended to play a significant role in guiding investment in research, monitoring, and resulting policy and management action in national parks for the foreseeable future. acknowledgements (back to top) we thank the many members of the sanparks conservation services division that contributed at various stages during the development of this framework, as well as steven chown, dian spear and tony rebelo for discussions and comments. references (back to top) bas, y., devictor, v., moussus, j.p. & jiguet, f., 2008, ‘accounting for weather and time-of-day parameters when analysing count data from monitoring programs’, biodiversity and conservation 17, 3403−3416. doi:10.1007/s10531-008-9420-6 bell, s., marzano, m., cent, j., kobierska, h., podjed, d., vandzinskaite, d., et al., 2008, ‘what counts? volunteers and their organisations in the recording and monitoring of biodiversity’, biodiversity and conservation 17, 3443−3454. doi:10.1007/s10531-008-9357-9 bengis, r.g., grant, r. & de vos, v., 2003, ‘wildlife diseases and veterinary controls: a savanna ecosystem perspective’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 349−369, island press, washington. biggs, h.c., 2003, ‘integration of science: successes, challenges, and the future’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 469−487, island press, washington. biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring, and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59−80, island press, washington. braschler, b., mahood, k., karenyi, n., gaston, k.j. & chown, s.l., 2010, ‘realizing a synergy between research and education: how participation in ant monitoring helps raise biodiversity awareness in a resource-poor country’, journal of insect conservation 14, 19−30. doi:10.1007/s10841-009-9221-6 bruner, a.g., gullison, r.e., rice, r.e. & da fonseca, g.a.b., 2001, ‘effectiveness of parks in protecting tropical biodiversity’, science 291, 125−128. doi:10.1126/science.291.5501.125, pmid:11141563 bryden, h.b. & de vos, v., 1994, ‘a scientific bibliography on the national parks of south africa’, koedoe, suppl. 1994. buckland, s.t., magurran, a.e., green, r.e. & fewster, r.m., 2005, ‘monitoring change in biodiversity through composite indices’, philosophical transactions of the royal society b 360, 243−254. doi:10.1098/rstb.2004.1589, pmid:15814343, pmcid:1569463 buckley, r., robinson, j., carmody, j. & king, n., 2008, ‘monitoring for management of conservation and recreation in australian protected areas’, biodiversity and conservation 17, 3589−3606. doi:10.1007/s10531-008-9448-7 butchart, s.h.m., walpole, m., collen, b., van strien, a., scharlemann, j.p.w., almond, r.e.a., et al., 2010, ‘global biodiversity: indicators of recent declines’, science 328, 1164−1168. doi:10.1126/science.1187512, pmcid:20430971 carey, c., dudley, n. & stolton, s., 2000, threats to protected areas. squandering paradise? the importance and vulnerability of the world’s protected areas, world wide fund for nature international, gland, switzerland. craigie, i.d., baillie, j.e.m., balmford, a., carbone, c., collen, b., green, r.e. & hutton, j.m., 2010, ‘ large mammal population declines in africa’s protected areas’, biological conservation 143, 2221−2228. doi:10.1016/j.biocon.2010.06.007 daszak, p., cunningham, a.a. & hyatt, a.d., 2000, ‘emerging infectious diseases of wildlife – threats to biodiversity and human health’, science 287, 443−449. doi:10.1126/science.287.5452.443, pmcid:10642539 department of environmental affairs and tourism (deat), 2002, environmental indicators for national state of the environment reporting south africa, department of environmental affairs and tourism, south african government, pretoria. department of environmental affairs and tourism (deat), 2005, south africa’s national biodiversity strategy and action plan, department of environmental affairs and tourism, south african government, pretoria. department of environmental affairs and tourism (deat), 2006, south african environment outlook. a report on the state of the environment, department of environmental affairs and tourism, south african government, pretoria. department of environmental affairs and tourism (deat), 2008, national protected area expansion strategy, department of environmental affairs and tourism, south african government, pretoria. department of environmental affairs and tourism (deat), 2009, south africa’s fourth national report to the convention on biodiversity, march 2009, south african national biodiversity institute, south african government, pretoria. department of water affairs and forestry (dwaf), 2005, ‘achieving sustainable forest management: the principles, criteria, indicators and standards framework’, a guide for forest managers & auditors to achieve sustainable forest management , department of water affairs and forestry, south african government, pretoria. department of water affairs and forestry (dwaf), 2007, south africa’s state of the forests report 2004–2006, department of water affairs and forestry, south african government, pretoria. de villiers, s. & thiart, c., 2007, ‘the nutrient status of south african rivers: concentrations, trends and fluxes from the 1970s to 2005’, south african journal of science 103, 343−349. driver, a., maze, k., lombard, a.t., nel, j., rouget, m., turpie, j.k., et al., 2005, ‘national spatial biodiversity assessment 2004: priorities for biodiversity conservation in south africa’, strelitzia 17, south african national biodiversity institute, pretoria. durrheim, g.p., 2009, ‘monitoring for sustainable forest management in the garden route’, south african national parks, scientific services, knysna. du toit, j., 2003, ‘large herbivores and savanna heterogeneity’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 292−309, island press, washington. du toit, j.t., rogers, k.h. & biggs, h.c. (eds.), 2003, the kruger experience: ecology and management of savanna heterogeneity, island press, washington. elzinga, c.l., salzer, d.w., willoughby, j.w. & gibbs, j.p., 2001, monitoring plant and animal populations, blackwell science, oxford. fabricius, c., 2004, ‘the fundamentals of community-based natural resource management’, in c. fabricius, e. koch, h. magome & s. turner (eds.), rights, resources and rural development: community-based natural resource management in southern africa, pp. 3−43, earthscan, london. field, s.a., o’connor, p.j., tyre, a.j. & possingham, h.p., 2007, ‘making monitoring meaningful’, austral ecology 32, 485−491. doi:10.1111/j.1442-9993.2007.01715.x foxcroft, l.c., 2009, ‘developing thresholds of potential concern for invasive alien species: hypotheses and concepts’, koedoe 51, 11−16. doi:10.4102/koedoe.v51i1.157 foxcroft, l.c., richardson, d.m., rouget, m. & macfadyen, s., 2009, ‘patterns of alien plant distribution at multiple spatial scales in a large national park: implications for ecology, management and monitoring’, diversity and distributions 15, 367−378. doi:10.1111/j.1472-4642.2008.00544.x gardner, t.a., barlow, j., araujo, i.s., avila-pires, t.c., bonaldo, a.b., costa, j.e., et al., 2008, ‘the c ost-effectiveness of biodiversity surveys in tropical forests’, ecology letters 11, 139−150. gaston, k.j., 1996, biodiversity: a biology of numbers and difference, blackwell science, oxford. gaston, k.j., 2010, ‘valuing common species’, science 327, 154−155. doi:10.1126/science.1182818, pmcid:20056880 gaston, k.j., charman, k., jackson, s.f., armsworth, p.r., bonn, a., briers, r.a., et al., 2006, ‘the ecological effectiveness of protected areas: the united kingdom’, biological conservation 132, 76−87. doi:10.1016/j.biocon.2006.03.013 gaston, k.j., jackson, s.f., cantú-salazar, l. & cruz-piñón, g., 2008, ‘the ecological performance of protected areas’, annual review of ecology, evolution, and systematics 39, 93−113. doi:10.1146/annurev.ecolsys.39.110707.173529 geldenhuys, c.j., 1992, ‘disjunctions and distribution limits of forest species in the southern cape’, south african forestry journal 161, 1−13. goodman, p.s., 2003, ‘assessing management effectiveness and setting priorities in protected areas in kwazulu-natal’, bioscience 53, 843−850. doi:10.1641/0006-3568(2003)053[0843:ameasp]2.0.co;2 grobler, j., 2009, biodiversity monitoring strategy and five-year action plan, version 2, south african national biodiversity institute, pretoria. hansen, a.j. & defries, r., 2007, ‘ecological mechanisms linking protected areas to surrounding lands’, ecological applications 17, 974–988. doi:10.1890/05-1098, pmcid:17555212 heink, u. & kowarik, i., 2010, ‘what are indicators? on the definition of indicators in ecology and environmental planning’, ecological indicators 10, 584−593. doi:10.1016/j.ecolind.2009.09.009 henry, p.y., lengyel, s., nowicki, p., julliard, r., clobert, j., čelik, t. et al., 2008, ‘ integrating ongoing biodiversity monitoring: potential benefits and methods’, biodiversity and conservation< /i> 17, 3357−3382. doi:10.1007/s10531-008-9417-1 hockings, m., 2003, ‘systems for assessing the effectiveness of management in protected areas’, bioscience 53, 823−832. doi:10.1641/00063568(2003)053[0823:sfateo]2.0.co;2 hockings, m., leverington, f. & james, r., 2006, ‘evaluating management effectiveness’, in m. lockwood, g.l. worboys & a. kothari (eds.), managing protected areas: a global guide, pp. 635−655, earthscan, london. hulme, p.e., 2009, ‘trade, transport and trouble: managing invasive species pathways in an era of globalization’, journal of applied ecology 46, 10−18. doi:10.1111/j.1365-2664.2008.01600.x iucn standards and petitions working group, 2009, guidelines for using the iucn red list categories and criteria, version 7.0., prepared by the standards and petitions working group of the iucn ssc biodiversity assessments sub-committee in august 2008, viewed 15 october 2009, from http://intranet.iucn.org/webfiles/doc/ssc/redlist/redlistguidelines.pdf. james, a., gaston, k.j. & balmford, a., 2001, ‘can we afford to conserve biodiversity?’, bioscience 51, 43−52. doi:10.1641/0006-3568(2001)051[0043:cwatcb]2.0.co;2 jones, k.e., patel, n.g., levy, m.a., storeygard, a., balk, d., gittleman, j.l., et al., 2008, ‘global trends in merging infectious diseases’, nature 451, 990−993. doi:10.1038/nature06536, pmcid:18288193 kapos, v., balmford, a., aveling, r., bubb, p., carey, p., entwistle, a., et al., 2008, ‘calibrating conservation: new tools for measures of success’, conservation letters 1, 155−164. doi:10.1111/j.1755-263x.2008.00025.x knight, m.h., 2007, sanparks state of biodiversity report (sob) background sheet: park planning & development, conservation services, unpublished report, sanparks, port elizabeth. leipold, m. & van zyl, c.j., 2008, ‘the economic impact of sport and recreational angling in the republic of south africa, 2007’, project report, pp. 48, development strategies international, cape town. lepetz, v., massot, m., schmeller, d.s. & clobert, j., 2009, ‘biodiversity monitoring: some proposals to adequately study species’ responses to climate change’, biodiversity and conservation 18, 3185−3203. doi:10.1007/s10531-009-9636-0 lombard, a.t., strauss, t., harris, j., sink, k., attwood, c. & hutchings, l., 2004, ‘marine component’, in south african national spatial biodiversity assessment technical report, chapter 4, south african national biodiversity institute, pretoria. lovett, g.m., burns, d.a., driscoll, c.t., jenkins, j.c., mitchell, m.j., rustad, l. et al., 2007, ‘ who needs environmental monitoring?’ frontiers in ecology and the environment 5, 253−260. doi:10.1890/1540-9295(2007)5[253:wnem]2.0.co;2 mace, g.m., delbaere, b., hanski, i., harrison, j.a., novo, f.g., pereira, h.m., et al., 2005, a user’s guide to biodiversity indicators, european academy of sciences advisory council, the royal society, london. margules, c.r., grant, c.c., poon, e., deacon, a., macfadyen, s., biggs, h.c., et al., 2003, a suggested biodiversity survey and monitoring program for kruger national park, scientific report 01/03, scientific services, skukuza, kruger national park. mcgeoch, m.a., butchart, s.h.m., spear, d., marais, e., kleynhans, e.j., symes, a., et al., 2010, ‘global indicators of biological invasion: species numbers, biodiversity impact and policy responses’, diversity & distributions 16, 95−108. doi:10.1111/j.1472-4642.2009.00633.x midgley, g.f., chown, s.l. & kgope, b.s., 2007, ‘monitoring effects of anthropogenic climate change on ecosystems: a role for systematic ecological observation?’, south african journal of science 103, 282−286. moloney, c.l. & shillington, f.a., 2007, ‘progress towards marine ecosystem observing systems in south africa’, south african journal of science 103, 301−305. naughton-treves, l., buck holland, m. & brandon, k., 2005, ‘the role of protected areas in conserving biodiversity and sustaining local livelihoods’, annual review of environment and resources 30, 219−252. doi:10.1146/annurev.energy.30.050504.164507 nichols, j.d. & williams, b.k., 2006, ‘monitoring for conservation’, trends in ecology and evolution 21, 668−673. doi:10.1016/j.tree.2006.08.007 nielsen, s.e., haughland, d.l., bayne, e. & schieck, j., 2009, ‘capacity of large-scale, long-term biodiversity monitoring programmes to detect trends in species prevalence’, biodiversity and conservation 18, 2961−2978. doi:10.1007/s10531-009-9619-1 noss, r.f., 1990, ‘indicators for monitoring biodiversity: a hierarchical approach’, conservation biology 4, 355−364. doi:10.1111/j.1523-1739.1990.tb00309.x podjed, d. & muršič, r., 2008, ‘dialectical relations between professionals and volunteers in a biodiversity monitoring organisation’, biodiversity and conservation 17, 3471−3483. doi:10.1007/s10531-008-9443-z pomeroy, r.s., watson, l.m., parks, j.e. & cid, g.a., 2005, ‘how is your mpa doing? a methodology for evaluating the management effectiveness of marine protected areas’, ocean & coastal management 48, 485−502. doi:10.1016/j.ocecoaman.2005.05.004 pringle, c.m. & collins, s.l., 2004, ‘needed: a unified infrastructure to support long-term scientific research on public lands’, ecological applications 14, 18−21. doi:10.1890/03-5106 regan, h.m., hierl, l.a., franklin, j., deutschman, d.h., schmalbach, h.l., winchell, c.s. et al., 2008, ‘species prioritization for monitoring and management in regional multiple species conservation plans’, diversity and distributions 14, 462−471. doi:10.1111/j.1472-4642.2007.00447.x republic of south africa, 2003, national environmental management: protected areas act, 57 of 2003, government printer, pretoria. republic of south africa, 2004, national environmental management: biodiversity act, 10 of 2004, government printer, pretoria. revenga, c., campbell, i., abell, r., de villiers, p. & bryer, m., 2005, ‘prospects for monitoring freshwater ecosystems towards the 2010 targets’, philosophical transactions of the royal society b 360, 397−413. doi:10.1098/rstb.2004.1595, pmcid:15814353, pmcid:1569454 reyers, b. & mcgeoch, m.a., 2007, ‘a biodiversity monitoring framework for south africa: progress and directions’, south african journal of science 103, 295−300. rogers, k.h. & o’keefe, j., 2003, ‘river heterogeneity: ecosystem structure, function and management’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 189−218, island press, washington. south african national parks (sanparks), 2006a, south african national parks coordinated policy framework governing park management plans, sanparks, pretoria, viewed 15 january 2010, from http://celtis.sanparks.org/conservation/park_man/cpfjanuary2010.pdf south african national parks (sanparks), 2006b, south african national parks project management policy v1.2., policy no. 13/1/p/01, south african national parks, pretoria. south african national parks (sanparks), 2008, south african national parks resource use policy, south african national parks, pretoria. schulze g.c., 2007, ‘atmospheric observations and numerical weather prediction’, south african journal of science 103, 3180−322. statistics south africa (ssa), 2008, south african statistical quality assessment framework (sasqaf), 1st edn., statistics south africa, pretoria. stolton, s., hockings, m., dudley, n., mackinnon, k., whitten, t. & leverington, f., 2007, reporting progress in protected areas: a site level management effectiveness tracking tool, 2nd edn., world bank/wwf forest alliance, wwf, gland, switzerland. soberón, j. & peterson, a.t., 2009, ‘monitoring biodiversity loss with primary species-occurrence data: toward national-level indicators for the 2010 target of the convention on biological diversity’, ambio 38, 29−34. doi:10.1579/0044-7447-38.1.29 spellerberg, i.f., 2005, monitoring ecological change, 2nd edn., cambridge university press, cambridge. doi:10.1017/cbo9780511614699 teder, t., moora, m., roosaluste, e., zobel, k., pärtel, m., kõljalg, u., et al., 2007, ‘monitoring of biological diversity: a common-ground approach’, conservation biology 21, 313−317. doi:10.1111/j.1523-1739.2006.00575.x, pmcid:17391181 the royal society, 2003, measuring biodiversity for conservation, policy document 11/03, the royal society, london. timko, j.a. & innes, j.l., 2009, ‘evaluating ecological integrity in national parks: case studies from canada and south africa’, biological conservation 142, 676−688. doi:10.1016/j.biocon.2008.11.022 unep-wcmc, 2008, state of the world’s protected areas: an annual review of global conservation progress, unep-wcmc, cambridge. van jaarsveld, a.s., biggs, r., scholes, r.j., bohensky, e., reyers, b., lynam, t., et al., 2005, ‘measuring conditions and trends in ecosystem services at multiple scales: the southern african millennium ecosystem assessment (safma) experience’, philosophical transactions of the royal society b 360, 425−441. doi:10.1098/rstb.2004.1594, pmcid:15814355, pmcid:1569451 van jaarsveld, a.s., pauw, j.c., mundree, s., mecenero, s., coetzee, b.w.t. & alard, g.f., 2007, ‘south african environmental observation network: vision, design and status’, south african journal of science 103, 289−294. van wilgen, b.w., biggs, h.c. & potgieter, a.l.f., 1998, ‘fire management and research in the kruger national park, with suggestions on the detection of thresholds of potential concern’, koedoe 41, 69−87. von maltitz, g.p., & shackleton, s.e., 2004, ‘use and management of forests and woodlands in south africa: stakeholders, institutions and processes from past to present’, in m.j. lawes, h.a.c. eeley, c.m. shackleton & b.g.s. geach (eds.), indigenous forests and woodlands in south africa: policy, people and practice, pp. 109−135, university of kwazulu-natal press, durban. walker, k.l., 2009, ‘protected-area monitoring dilemmas: a new tool to assess success’, conservation biology 23, 1294−1303. doi:10.1111/j.1523-1739.2009.01203.x, pmcid:19765040 walpole, m., almond, r.e.a., besançon, c., butchart, s.h.m., campbell-lendrum, d., carr, g.m., et al., 2009, ‘tracking progress toward the 2010 biodiversity target and beyond’, science 325, 1503−1504. doi:10.1126/science.1175466, pmcid:19762630 world wildlife fund (wwf), 2004, ‘are protected areas working? an analysis of forest protected areas by wwf’, world wildlife fund international, gland, switzerland. filelist convert a pdf file! page 1 page 2 page 3 page 4 terblanche 1.qxd the taxonomy, biogeography and conservation of the myrmecophilous chrysoritis butterflies (lepidoptera: lycaenidae) in south africa r.f. terblanche and h. van hamburg terblanche, r.f. and h. van hamburg. 2003. the taxonomy, biogeography and conservation of the myrmecophilous chrysoritis butterflies (lepidoptera: lycaenidae) in south africa. koedoe 46(2): 65–81. pretoria. issn 0075-6458. the relevance and integration of scientific knowledge to conservation management of the locally popular and highly endemic butterfly genus chrysoritis are investigated within the research fields of taxonomy and biogeography. the butterfly genus chrysoritis contains at least 41 species endemic to south africa. the taxonomy of chrysoritis has reached a state where revisions could easily result in a plethora of names between “lumping and splitting”. in practice, the state of the taxonomy of these butterflies on species level may alter their conservation priority. the two most species rich species groups in chrysoritis have different centres of endemism, however, a butterfly atlas becomes a necessity to reveal more about their biogeography. there is an absence of butterfly species lists in many of our national parks and nature reserves. legislation should facilitate rather than limit the valuable role of the amateur lepidopterist to add distribution records. in turn, the amateur lepidopterists should adapt and make an effort to explore unknown localities, apart from monitoring butterflies at their well-known localities. the red listing of localised butterflies in south africa, including a number of chrysoritis species, is in need of an urgent review in the light of the most recent iucn categories. a species such as chrysoritis dicksoni should be protected by law – but at its known localities. the scenario that real conservation action is only needed if the last known locality of a butterfly is threatened, should be abolished. a paradigm shift to conserve the metapopulations of the highly endemic chrysoritis genus and not merely a few of its species as items that appear on lists, seems necessary. key words: chrysoritis, myrmecophilous, endemic, conservation, research priorities, environmental management, south africa, taxonomy, biogeography, red list r.f. terblanche and h. van hamburg, school of environmental sciences and development, private bag x6001, potchefstroom university for che, potchefstroom, 2520 south africa. issn 0075-6458 65 koedoe 46/2 (2003) introduction the purpose of this paper is to consider the conservation of the genus chrysoritis as a real example of insect conservation in south africa. interactions between disciplines, such as taxonomy and legislation, as well as the role of various interested parties such as the amateur lepidopterists and professionals, have been poorly investigated to date. the main objective of this overview is to identify research needs and to create a framework along which research priorities could be directed towards sustainable environmental management with a view to butterfly conservation. all the members of the genus chrysoritis seem to be myrmecophilous (heath 1997a; 2001). myrmecophilous butterflies pose special challenges for conservation management due to their intricate habitat requirements and populations are often localised. a summary of the distribution of the chrysoritis species in lesotho, swaziland and the provinces of south africa is presented for the first time (table 1). biogeographic notes with regard to chrysoritis is given from a regional perspective on conservation priorities and will hopefully stimulate further research on the distribution of these butterflies. terblanche 1.qxd 2005/12/09 11:25 page 65 koedoe 46/2 (2003) 66 issn 0075-6458 table 1 the distribution of chrysoritis species in the provinces of south africa as well as the kingdom of lesotho and swaziland. a number 1 indicates that the chrysoritis species occurs in that area. the major regional literature sources used were clark & dickson (1971), terblanche (1991), keller, amodio & stänz (1997), kloppers & van son (1978), pringle, henning & ball (1994), duke, saunders & saunders (1999), and claassens (2000). chrysoritis species lesotho, swaziland and provinces of south africa ec fs gp kz le li mp nc nw wc sw chrysaor species group chrysoritis aethon (trimen & bowker, 1887) 1 1 1 chrysoritis aureus (van son, 1966) 1 1 chrysoritis chrysaor (trimen, 1864) 1 1 1 1 1 1 1 chrysoritis lycegenes (trimen, 1874) 1 1 1 chrysoritis lyncurium (trimen, 1868) 1 1 chrysoritis midas (pennington, 1962) 1 chrysoritis natalensis (van son, 1966) 1 1 chrysoritis phosphor (trimen, 1866) 1 1 1 chrysantas species group chrysoritis chrysantas (trimen, 1868) 1 1 oreas species group chrysoritis dicksoni (gabriel, 1946) 1 chrysoritis oreas (trimen, 1891) 1 zeuxo species group chrysoritis zeuxo (linnaeus, 1764) 1 1 chrysoritis zonarius (riley, 1938) 1 1 pyroeis species group chrysoritis felthami (trimen, 1904) 1 1 chrysoritis pyroeis (trimen, 1864) 1 1 1 thysbe species group chrysoritis adonis (pennington, 1962) 1 chrysoritis aridus (pennington, 1953) 1 1 chrysoritis azurius (swanepoel, 1975) 1 chrysoritis beaufortius (dickson, 1966) 1 chrysoritis beulah (quickelberge, 1966) 1 chrysoritis blencathrae (heath and ball, 1992) 1 chrysoritis braueri (pennington, 1967) 1 chrysoritis brooksi (riley, 1938) 1 chrysoritis daphne (dickson, 1975) 1 chrysoritis endymion (pennington, 1962) 1 chrysoritis irene (pennington, 1968) 1 chrysoritis nigricans (aurivillius, 1924) 1 chrysoritis orientalis (swanepoel, 1976) 1 chrysoritis palmus subsp. palmus (stoll, 1781) 1 1 chrysoritis pan (pennington, 1962) 1 1 1 chrysoritis pelion (pennington, 1953) 1 1 1 1 chrysoritis penningtoni (riley, 1938) 1 chrysoritis perseus (henning, 1977) 1 chrysoritis plutus (pennington, 1976) 1 1 chrysoritis pyramus (pennington, 1953) 1 chrysoritis rileyi (dickson, 1966) 1 chrysoritis swanepoeli (dickson, 1965) 1 chrysoritis thysbe (linnaeus, 1764) 1 1 chrysoritis trimeni (riley, 1938) 1 chrysoritis turneri (riley, 1938) 1 1 chrysoritis uranus (pennington, 1962) 1 chrysoritis violescens (dickson, 1971) 1 ec fs gp kz le li mp nc nw wc sw total 13 3 1 9 2 0 5 14 0 25 1 terblanche 1.qxd 2005/12/09 11:25 page 66 discussion taxonomy of the genus chrysoritis most of the present chrysoritis species were previously known as the poecilmitis species. based on wing pattern, genitalia features and larval morphology, heath (1997a) treated the genera chrysoritis, poecilmitis, bowkeria and oxychaeta as synonyms of chrysoritis. his treatment with regard to chrysoritis, poecilmitis and oxychaeta was supported by the phylogeny inferred from mitochondrial cytochrome oxidase i sequences (rand et al. 2000). rand et al. (2000) could not ascertain the phylogenetic position of bowkeria due to a lack of fresh specimens, being a rare canopy dweller and difficult to obtain. the inclusion of bowkeria in chrysoritis needs further study, especially since its life history remains unknown. the studies of heath (1997a) that led to the inclusion of chrysoritis, oxychaeta and poecilmitis being synonyms of chrysoritis, as well as rand et al. (2000) expanded the understanding of the genus chrysoritis considerably, including the ecological functional similarities among the species in this group. for instance, apart from chrysoritis dicksoni (previously known as oxychaeta dicksoni) that might have a derived aphytophagy, the rest of the chrysoritis species studied to date are phytophagous with a mutualism that includes myrmicine ants of the genera myrmicaria and crematogaster. figure 1 illustrates some members of the species groups of chrysoritis. heath (2001) is followed for the species groups within the chrysoritis genus, which in turn is largely based on heath (1997a) and rand et al. (2000) (see table 1 for the species and species groups). heath (2001) instated a number of synonymies within the genus chrysoritis, which reduced the total number of species from 59 to 42. it was suggested that there may be a number of synonymies among the remaining 42 species as well. even if the number of species was to be lowered further, the number of endemic species of this genus could still be regarded as very issn 0075-6458 67 koedoe 46/2 (2003) high. although the latest synonymies presented by heath (2001) have been followed in this paper and in the most recent checklist of southern african butterflies (vári et al. 2002), there is currently much debate among lepidopterists of africa whether all the synonymies suggested by heath (2001) should be accepted. a few factors that could have fig. 1. examples of chrysoritis species. top: c. aethon; middle: c. pelion; bottom: c. zeuxo. terblanche 1.qxd 2005/12/09 11:25 page 67 koedoe 46/2 (2003) 68 issn 0075-6458 led to the confusion in lepidopterists’ circles are briefly discussed below. heath (2001) stated that his conclusions rely in part on the paper by rand et al. (2000) and a revision of aphnaeini (heath 1997a) but did not specify to what extent for each of the synonymies. rand et al. (2000) investigated mitochondrial dna sequences of only 19 species of the former 59 species of chrysoritis species. although this investigation was to a great extent sufficient to clarify and confirm the species groups and other phylogenetic relationships within the genus chrysoritis, not enough information was gained to base synonymies of species on, with the possible exception of the chrysoritis zeuxo species group which is the only group of chrysoritis of which some dna information exists for all the species in the group. it seems that the synonymies presented by heath (2001) of most of the chrysoritis species are still largely based on morphological and life history information and not the dna study by rand et al. (2000). heath (1997a, 1997b) showed that the morphology of chrysoritis genitalia, even among the majority of the species in the genus, is very uniform in structure, and that this might be related to their myrmecophily. therefore, the separation of chrysoritis species relies on other morphological characteristics. unfortunately, other morphological details and life history information that were presented to sink species in the heath paper (2001), were not presented or illustrated in any specific detail for any of the synonymies. reference to characterisics specified in the original descriptions as well as the type series of all the entities that were synonomised, were lacking at large. heath (2001) states that rearing chrysoritis over several years from egg to adult from caterpillars collected in the field from various localities, has demonstrated the extent to which the environment could, during the larval stage, affect adult wing shape, size and the markings of both wing surfaces. no reference to controlled experiments in a climatic breeding room, were provided. therefore, in the comparisons of the life histories, the reader could not confirm or repeat the judgements. one example, chrysoritis lycia, has been synonimised with chrysoritis chrysaor. based on breeding experiments, of which no further details apart from the host plants and localities are presented, the conclusion is drawn that chrysoritis lycia could be considered an extreme and more yellowish form of chrysoritis chrysaor. however, important other morphological characteristics used to distinguish chrysoritis lycia from chrysoritis chrysaor, such as the series of markings on the hind wing underside margins are not discussed. the chrysoritis lycia material has also been bred from material at leipoldtville, which is a few hundred kilometres from the type locality near matjesfontein. even if the synonymies and heaths’ judgements based on his reference collection were correct in these cases, the above lack of details, illustrations as well as more specific details about the material and methods could have led to the present confusion. it is to be questioned if sufficient information is available to synonimise many of the chrysoritis species at present. a similar analogy exists in the plant genus haworthia that also contains variations between and within populations and which is also of considerable amateur and collectors’ interest (see terblanche et al. 1993; bayer 1999). heath (2001) suggested that other synonymies might be extant among more chrysoritis species, especially within the chrysoritis chrysaor species group. this follows the suggestion by owen-johnston (1991) that chrysoritis lycegenes and chrysoritis lyncurium and an unidentified chrysoritis entity that occurs at morgenzon in southern mpumalanga, might be a cline belonging to the same species (owen-johnston 1991). in addtion heath (2001) suggested that the relation between chrysoritis aethon and chrysoritis aureus should be investigated. the habitats of chrysoritis lyncurium seem to be in great danger, especially from alien invasion (woodhall 1996). this may butterfly escape as a conservation priority due to taxonomic uncertainties. terblanche 1.qxd 2005/12/09 11:25 page 68 issn 0075-6458 69 koedoe 46/2 (2003) table 2 a summary of the red data book status of chrysoritis species in the past. literature sources: s.f. & g.a. henning (1989) and g.a & s.f. henning (1995). the taxonomic status and taxonomic research recommendations according to heath (2001) are given in the last column. species group and species red data status proposed taxonomic status 1989 1995 taxonomic status (heath, 2001) chrysaor species group chrysoritis aureus (trimen, 1864) r r relation with c. aethon needs research chrysoritis lyncurium (trimen, 1868) r r relation with c. lycegenes needs research chrysoritis phosphor subsp. borealis r r status quo (quickelberge, 1972) oreas species group chrysoritis dicksoni (gabriel, 1946) v e status quo chrysoritis oreas (trimen, 1891) r r status quo zeuxo species group chrysoritis cottrelli (dickson, 1975) e r regarded as a synonym of c. zeuxo pyroeis species group chrysoritis pyroeis subsp. hersaleki r r status quo (dickson, 1970) thysbe species group chrysoritis adonis (pennington, 1962) r i status quo chrysoritis azurius (swanepoel, 1975) r i status quo chrysoritis balli r i synomym (dickson & g.a. henning, 1980) chrysoritis brooksi subsp. taerei r i status quo (dickson, 1966) chrysoritis daphne (dickson, 1975) r i relation with c. swanepoeli needs research chrysoritis endymion (pennington, 1962) r i status quo chrysoritis irene (pennington, 1968) r r status quo chrysoritis henningi (bampton, 1981) r r regarded as a synonym of c. pan chrysoritis hyperion(dickson, 1975) r i regarded as a synonym of c. swanepoeli chrysoritis kaplani (s.f. henning, 1980) r r regarded as a synonym of c. beaufortius chrysoritis lyndseyae (s.f. henning, 1979) i i regarded as synonym of c. thysbe subsp. bamptoni chrysoritis mithras (pringle, 1995) – r regarded a subsp. of c. thysbe by heath (2001) chrysoritis nigricans subsp. nigricans i i status quo (aurivillius, 1924) chrysoritis nigricans subsp. zwartbergae i i status quo (dickson, 1982) chrysoritis orientalis (swanepoel, 1976) i i relation with c. pelion needs research chrysoritis pan (pennington, 1962) i i contains several new synonymies chrysoritis penningtoni (riley, 1938) r i status quo chrysoritis pyramus (pennington, 1953) r i contains c. balli as a new synonymy. relation with c. thysbe needs research chrysoritis rileyi (dickson, 1966) r i status quo chrysoritis stepheni (dickson, 1978 r i regarded as synonym of c. beaufortius chrysoritis swanepoeli (dickson, 1965) r i contains c. hyperion as a new synonymy. relation with c. daphne needs research chrysoritis trimeni (riley, 1938) i i relation with c. pan needs research chrysoritis wykehami (dickson, 1980) r i regarded as a synonym of c. turneri by heath (2001) r =rare ; v = vulnerable; e = endangered; i = indeterminate terblanche 1.qxd 2005/12/09 11:25 page 69 koedoe 46/2 (2003) 70 issn 0075-6458 the taxonomic uncertainties regarding the chrysoritis species are of much more than academic importance, since the conservation of unique entities are at stake. for instance, most often a species has a higher priority in practice as a conservation priority than a subspecies or a form that is part of a “cline”. furthermore, species and subspecies appear on the protected or red lists and not unique habitats. more specifically, it means that the conservation priority for entities such as chrysoritis hyperion, chrysoritis lyndseyae, chrysoritis kaplani and chrysoritis stepheni, which were all regarded as rare or intermediate in the red data book of henning & henning (1989), could change considerably (see table 2). to name one more example: chrysoritis cottrelli was previously regarded as endangered (1989) but may dissapear from the revised list (table 2). these species are now all regarded as synonyms of other species according to heath (2001). a taxonomic impediment could lead directly to a conservation impediment (new 1997). the above illustrates the importance of taxonomy and its implications for conservation of these localised butterfly species. however, it is perhaps the perception of conservation of rare butterflies that must change. the genus chrysoritis in its entirety is highly endemic and contains a large number of populations and subpopulations of which most are very localised (even for some of the more widespread species). one is also struck by the variability within and among the local (sub) populations of chrysoritis species. with regard to wing pattern and wing morphology, some of them are seemingly on their way to become species, and other forms/entities perhaps becoming less plentiful – these life processes should be conserved. it is especially the mixture of chrysoritis thysbe forms along the western and southern coast of south africa as well as the adjacent inland areas, that reflects entities in a stage of speciation difficult to estimate. sympatry and allopatry have been poorly resolved in chrysoritis, especially for example along the western coast and the swartberg. it is often difficult to know whether different populations or variations are encountered. such localities include lambert’s bay, hondeklip bay, and port nolloth along the western coast in the northern cape. these localised populations are indicative of unique sets of habitat requirements. a small colony of a widespread chrysoritis that deserves conservation is the colony of chrysoritis thysbe at hout bay in the cape peninsula (claassens 1994). for insect conservation, the impact of exotic and invasive flora is of particular concern in such rare and restricted habitat types (mcgeoch 2002). the butterflies of chrysoritis with their either metallic orange or metallic orange and blue upper sides of wings could all be regarded as an aesthetic asset and much could be lost before science unravels the full significance of their phylogeny. distribution of chrysoritis species obviously, a better knowledge of the distribution of chrysoritis species will enhance research (taxonomy and ecology) and the applications of environmental impact assessments and management. despite many shortcomings in the study of the distribution of butterflies, proportionally far more information is available compared to many other groups of arthropods in africa. at least one knows to some extent that a butterfly is rare or not—this cannot apply to many other insects of which the information on distribution and the manpower to obtain these are very poor. furthermore, relatively few butterflies await discovery in south africa, which is not true for many other insect groups. despite the possibilities that such a well-known group presents in terms of distribution, it was is not yet been applied sufficiently in the african context. a butterfly atlas is becoming an increasing need to interpret the distribution of south african butterflies (terblanche & taylor 2000). table 1 gives a summary of the distribution of all the chrysoritis species in all the new provinces (since 1994) of south africa as well as other african countries where these butterflies have been recorded. political terblanche 1.qxd 2005/12/09 11:25 page 70 issn 0075-6458 71 koedoe 46/2 (2003) boundaries were used due to the lack of data in an atlas format. these political boundaries are used for practical reasons since the distribution in standard reference works on butterflies are commonly described according to provinces (cotrell 1985). these political boundaries are still useful, since it is known for example that much of the fynbos biome is confined to the western cape as well as smaller parts of the eastern and northern cape and that the most of the free state consists of grassland (see rutherford & westfall 1994). furthermore, the list is now available to conservation authorities in the different regions. much of the knowledge on the distribution of chrysoritis species has been accumulated by many amateur lepidopterists. the genus chrysoritis is almost entirely endemic to south africa, lesotho and swaziland (table 1). the only chrysoritis species collected outside the borders of south africa, lesotho or swaziland, is chrysoritis chrysantas. this was found in the southern parts of namibia, by plowes on the tiras mountains (pringle et al. 1994). the genera chrysoritis and poecilmitis, which are today only known as one genus, namely chrysoritis, had been described as cape province centred (an area now known as the eastern cape, western cape and northern cape) by cottrell (1985). it falls into the category of species mainly confined to the old cape province (although some have been to a limited extent recorded outside this area. the work of cottrell (1985) preceeded the synonymies suggested by heath (2001) and the new provinces in south africa so that the levels of endemism are briefly reviewed here. from table 1, containing the species list according to the revision by heath (2001), it can be seen that the highest number of chrysoritis species occurs in the western cape, namely 25 (59.5% of the total), followed by the 14 species from the northern cape (33%) and 13 species of the eastern cape (31%). this echoes cottrell (1985) that chrysoritis is cape-centred. the richest area of rare endemic lycaenidae is found in the cape fold mountains (samways 1993), a trend strongly followed by the chrysoritis species. there is also a high number of local varieties of subpopulations of chrysoritis species in the sandy (dune) areas of the western cape. the extent to which the entities are sympatric and allopatric has not been clarified sufficiently. more distribution records will add to the taxonomic and ecological insights. some gaps in the distribution of species might be real rather than artificial, warranting subspecific status of a number of “clines”. no chrysoritis species have yet been reported from the highlands of zimbabwe or further northwards in africa. this contrasts with species of other ant-associated lycaenid genera with high concentrations of endemic species in south africa, especially the western cape (fynbos). such genera include aloeides, lepidochrysops and even thestor. the latter has one recorded species from parts of zimbabwe. a few aloeides species and many more lepidochrysops species are also found in equatorial africa. these distributions of ant-associated genera are in contrast with the genus chrysoritis and highlight the high level of endemism of the genus. the distribution of the two most species rich groups, the chrysoritis chrysaor species group and the chrysoritis thysbe species group is discussed in more detail to identify further research priorities of the biogeography of the genus. all the species within the chrysoritis chrysaor species group (which include chrysoritis aureus) lack the metallic blue colour on the upper side of the wings. instead, they only have an orange upper side with black markings and a metallic sheen on the orange background on the upper side of the wings—from which the vernacular/trivial name “copper” originated (fig. 1). the metallic blue (opal) colouring with orange and black markings are most prominent in most members of the chrysoritis thysbe group (fig. 1). this group is largely confined to the fynbos biome, with a few exceptions. simplified, the distribution of chrysoritis species follows a pattern from the southern (western and eastern cape) part of south africa with northerly extensions to the westterblanche 1.qxd 2005/12/09 11:25 page 71 koedoe 46/2 (2003) 72 issn 0075-6458 ern escarpment to the coast and eastern escarpment to the coast. chrysoritis is poorly represented in the central grassland plateau. no chrysoritis species occur in the savanna biome, hence their absence in the north west and limpopo provinces of south africa (table 1; fig. 2). the most northern locality record for a species belonging to the chrysoritis thysbe species group, on the eastern (moister) side of south africa, is that of chrysoritis pelion. the northernmost locality records for chrysoritis pelion is at the golden gate highlands national park. m.c. williams discovered the locality (terblanche 1991). chrysoritis pelion was also collected by terblanche, in january 1991, near sentinel mountain, where the drakensberg and maluti mountains meet. the butterfly is more widespread to the south, in the highlands of lesotho (fig. 2). both the above localities of chrysoritis pelion are found at an altitude above 2700 m, in areas with rocks and shrubs, among the grasslands. the highlands, northwards of the golden gate highlands national park and the sentinel, drop considerably in height above see level (down to a maximum height of 2350 m). chrysoritis pelion represents the northernmost outlier of the chrysoritis thysbe group on the eastern side of southern africa, where the butterfly is found on the alti mountain grassland (granger & bredenkamp 1996). patches of afromontane vegetation, affiliated to fynbos, are scattered throughout this grassland (granger & bredenkamp 1996). this distribution of chrysoritis pelion represents a faunal reflection of extensions of such fynbos-affiliated vegetation. in contrast, chrysoritis aethon of the chrysoritis chrysaor species group, has been found much further north on mariepskop (fig. 2) (pringle et al. 1994) and represents at present the most northerly distributed chrysoritis species. mariepskop is 510 km nne of the golden gate highlands national park. the chrysoritis chrysaor species group is significant in the sense that it contains the most north-easterly recorded species of the chrysoritis genus, namely fig. 2. map of south africa showing the nine provinces and three localities of importance. terblanche 1.qxd 2005/12/09 11:25 page 72 issn 0075-6458 73 koedoe 46/2 (2003) chrysoritis phosphor subsp. phosphor, chrysoritis aethon, chrysoritis aureus and the widespread chrysoritis chrysaor. significantly, the northern extensions of the genus on the western side of south africa contain mostly members of the chrysoritis thysbe group. on the northern extension of their range and on the eastern side of the country, the chrysoritis chrysaor group is better represented. all the chrysoritis species that occur in the moister (eastern) side of south africa belong to the chrysoritis chrysaor species group and the centre of endemism of the chrysoritis chrysaor species group seems to be outside the fynbos biome. only two of the eight species (that is 25 %) that belongs to the chrysoritis chrysaor species group has been recorded in the western cape, of which one is also widespread in the eastern side of south africa. this is in contrast with the tendency of the rest of the genus (table 1). it seems, therefore, that the centre of endemism of the chrysoritis chrysaor group is in the highlands of kwazulu/ natal (6 species), mpumalanga (5 species) and the eastern cape (4) species. the exception is chrysoritis natalensis that occurs in the moist, subtropical, eastern coastal areas. nevertheless, the centre of endemism of the chrysoritis chrysaor group is not in the fynbos biome but in the eastern parts of the grassland biome. furthermore, when the northerly extensions of the chrysoritis genus is considered, the chrysoritis chrysaor group has its centre of endemism within the summer rainfall area, whereas the chrysoritis thysbe group has its centre of endemism in the winter rainfall area. the only chrysoritis associated with afromontane forest, namely chrysoritis phosphor, is also a member of the chrysoritis chrysaor species group, although chrysoritis phosphor, that superficially looks atypical to the genus might not be congeneric. interestingly, chrysoritis phosphor is often recorded at forest margins bordering grassland. the only chrysoritis species, apart from the ubiquitous chrysoritis chrysaor that is encountered sparsely in the rocky highveld grassland, is chrysoritis aureus. acocks (1975, 1988) interpreted the bankenveld (now rocky highveld grassland) as a fire-maintained grassland that would develop into savanna if fire were excluded. fire, as variable, plays at least an important role in the maintenance of grassland (bredenkamp & van rooyen 1996; o’connor & bredenkamp 1997). in the light of the above distribution of chrysoritis species, it is significant that chrysoritis aureus extends the distribution of chrysoritis to inland grassland areas to the west of the escarpment at altitudes between 1600 m and 1800 m. these distributions show that even among the species groups of chrysoritis different ecological pressures may have shaped the species. a study of the vegetation of three localities of chrysoritis aureus points to the possible importance of fire in maintaining a viable open grassland habitat for the butterfly (terblanche et al. 2003). since many colonies of species of the chrysoritis chrysaor group is often found in grassy areas not far from forests or well-wooded kloofs, fire and climate may have been important in shaping these species and their ant associations. after decades of exploration by a number of lepidopterists only five confirmed localities of chrysoritis aureus were reported (terblanche et al. 2003). a fortunate contribution towards the conservation of chrysoritis aureus was the discovery of the locality at the alice glockner nature reserve (near heidelberg, gauteng) in 1998 by p.s. roos and g.a. henning (roos & henning 2000). this discovery has increased the conservation priority of the, at that stage, undeveloped alice glockner nature reserve considerably, since this locality would probably have been destroyed by urbanisation. the future of chrysorities aureus has been secured by the co-operation between developers, government officials and concerned ngo’s such as the lepidopterists’ society of africa. chrysoritis aureus is very localised in all the areas where it occurs. most of the individuals in such a colony will for most of the time terblanche 1.qxd 2005/12/09 11:25 page 73 koedoe 46/2 (2003) 74 issn 0075-6458 fly within an area of 200 m2 to 400 m2. the size of these localities is being measured at present. migration behaviour is not associated with any of the chrysoritis species and the populations of every species tend to live in relatively small confined areas. the chrysoritis populations and sub populations are therefore referred to as colonies in literature. males of the chrysoritis aureus seem to choose an outpost (usually some branch of a shrub) from where it controls its small territory, whereas females often wander around within the borders of such a colony. despite the fact that confinement of chrysoritis species to its habitats is appreciated by all the lepidopterists experienced with the chrysoritis butterflies, scientific information on the dispersal of chrysoritis species is absent. two single males and two single females of chrysoritis aureus were recorded by r.f. terblanche on 1 october 2002 and 23 january 2002 at suikerbosrand nature reserve 10.5 km w and 6.5 km wsw of the known locality in this reserve. no host plant (clutia pulchella) occurs near these spots, and only a single specimen that was clearly staying in a small area of not more than 25 m2 was recorded each time. the two males were observed, each in an isolated spot more than 100 m apart, at perdekop (26º30's; 28º15'e). they were near the summit, defending their small area in the manner they would do in a normal colony. the females were recorded on separate dates, on the northern aspect below the summit of the hill south of diepkloof (26º29's; 28º12'e). all these specimens were recorded at an altitude of 1860–1900 m, which is higher than the normal elevation at which chrysoritis aureus occurs (see table 3). the markings of the undersides of these specimens left one in no doubt about the identification. this is the first time that such “isolated” chrysoritis aureus individuals were reported, which is especially significant in the light of the above-mentioned surveys that the lepidopterists society of southern africa had done in suikerbosrand nature reserve (henning 1986a, 1986b, 1987). the full significance of these findings is being further researched but may point to dispersal behaviour. the above outline of the knowledge on the distribution of chrysoritis aureus highlights a few important aspects of research and conservation management. the importance of metapopulation studies and the lack of knowledge on the dispersal of chrysoritis species become clear. corridors and linkages are important for the conservation of insect diversity (samways 1993; 1994; pryke & samways 2001) and the possible dispersal behaviour of chrysoritis should be investigated in more detail. table 3 the five confirmed and apparantly viable localities of chrysoritis aureus (heidelberg copper butterfly) locality province grid reference 1 altitude (m) alice glockner nature reserve gauteng 26º34's, 28º22e(2628cb) 1690 8 km s heidelberg hill n of greylingstad mpumalanga 26º44's, 28º44'e(2628da) 1810 malanskraal locality14 km w balfour mpumalanga 26º39's, 28º26'e(2628cb) 1780 suikerbosrand nature reserve gauteng 26º29's, 28º18'e(2628ad) 1720 type locality at the south african gauteng 26º29's, 28º21'e(2628ad) 1600 national force premises 1 the grid reference reading is from the north-western corner of each minute by minute square. terblanche 1.qxd 2005/12/09 11:25 page 74 issn 0075-6458 75 koedoe 46/2 (2003) conservation history (ngos, politics, law, education and environmental management) science is vital, but alone cannot curb the decline of the earth’s biodiversity (kohm et al. 2000). kohm et al. (2000) further state that new partnerships and improved communication should be forged among scientists, managers and policy makers so that conservation science could be available and understandable to all parties. whitten et al. (2001) that ask if conservation biology is merely another scientific discipline, safely nestled within the confines of academia, is very appropriate for butterfly conservation (and not only the forests of indonesia that are discussed in the relevant paper). the question is further asked whether conservation is not a mission-concerned with judicial reform, political economy, other people’s spatial planning, community participation, poverty alleviation, human and institutional capacity, consumption, population growth and agricultural planning (whitten et al. 2001). the great challenge of conserving butterfly habitats in south africa thus not only depends on the level of biological scientific information, but on how that information is presented and linked with management that takes into account socio-economic factors. conservation history the conservation of butterflies in south africa has a proud history, but has not been without difficulties. henning (1997) gave an outline of major events that enhanced the conservation of butterflies in south africa. four of sixteen butterflies that were added to schedule 2 of the list of protected wild animals in the cape province in 1976, due to motivation by c.g.c dickson, were chrysoritis species. these were chrysoritis endymion, chrysoritis lyncurium, chrysoritis nigricans and chrysoritis rileyi. in the northern areas (the old transvaal), chrysoritis aureus was added to the protected wild animal list of transvaal (now known as four provinces: gauteng, mpumalanga, northern province and north west) under ordinance 12 of 1983, since the type locality had been the only known locality of the butterfly for a long time (de wet 1987). unfortunately, all the members of the genus charaxes were also put on the protected wild animal list at the same time. many of the charaxes species are very widespread in the savanna and had not been under any threat at any stage. the placement of all the charaxes species on the protected list therefore compromised the validity of the above ordinance and did not help to highlight the importance of protecting chrysoritis aureus. this showed that it is essential that effective consultation with stakeholders should take place before laws are instated. an ngo that has played a very important role in lepidoptera conservation to date is the lepidopterists’ society of southern africa that was founded in 1983. one of the aims of the lepidopterists’ society of africa (previously of southern africa) is to promote the conservation of lepidoptera in the afrotropical region. although this society consists largely of amateur lepidopterists, major contributions towards butterfly (insect) conservation in africa was initiated by its members. the proclamation of the ruimsig entomological reserve for the threatened aloeides dentatis subsp. dentatis by the roodepoort city council in 1985 – the first of its kind in africa – was a landmark for conservation of localised butterflies in africa. this event draw attention to the urgency and significance of conserving the vulnerable habitats of localised myrmecophilous butterflies. no issue on insect conservation in africa entered the realm of government policies and public perceptions to such an extent as the saga that led to the protection of the last known locality of orachrysops niobe (the brenton blue butterfly). the “brenton blue saga” has been described in detail by steenkamp & stein (1999). the minister of environmental affairs and tourism invoked section 31a of the environment conservation act, 73 of 1989, whereby a landowner was deprived of developing his property in order to conserve the habitat of the brenton blue butterfly (orachrysops niobe) (steenkamp & stein 1999). any similar conflict to follow will be able to deal with the matter at hand with far terblanche 1.qxd 2005/12/09 11:25 page 75 koedoe 46/2 (2003) 76 issn 0075-6458 firmer legislative and procedural foundations at their disposal (steenkamp & stein 1999). the red data book of butterflies (henning & henning 1989; henning & henning 1995) was yet another major achievement for insect conservation in africa. again, without the endeavours of many professional entomologists but many more of the amateur fraternity over the years, the red data book would have remained only a dream. table 2 gives a summary of the chrysoritis species that were regarded as red data species. a number of synonymies are present for formerly red data categorised butterflies (table 2), which confirms that the taxonomy of the chrysoritis genus, should be more satisfactory resolved, especially if the aim was to compile species lists for red listing. the red listing of some of the chrysoritis species needs to be reviewed to an extent that one can follow conservation progress clearly. the latest categories by the iucn (2001)—adopted in 1994—will help to standardise and establish a red listing, especially in the case of chrysoritis, for those species that were previously in the rare and intermediate categories of the red data book. a review falls beyond the scope of this paper but a few examples are mentioned to illustrate some practical aspects of such a review. in the case of chrysoritis dicksoni one is sure at present that the butterfly is in the process of vanishing if proper conservation strategies were not implemented in the near future (this butterfly should be entered in the critically endangered category). as mentioned earlier in the paper, the numbers of chrysoritis aureus are also dwindling (it is almost absent at its type locality) but it seems that conservation efforts may be in time to save the butterfly, and especially much of its metapopulation structure, from the impact of urbanisation. localised chrysoritis species in remote farm areas such as those in the sutherland area (roggeveld escarpment) in the northern cape should probably be transferred from the old rare to the new least concern category or be abandoned from the red list if more localities become known (see iucn 2001). monitoring of these localised butterflies should be in place for such species that could become threatened in a relatively short time by alien vegetation and perhaps also global climate changes. according to the most recent iucn categories, the red listing is being addressed at present. blue lists of threatened species may later be introduced to motivate people in their conservation efforts (gigon et al. 2000). such lists could be of special value to the conservation of chrysoritis species. conservation, collecting and the law the legal regime for the conservation and sustainable use of biodiversity is centred on the 1992 convention on biological diversity. the primary focus of the 1992 convention is the conservation of biodiversity and equitable distribution of its benefits (glazewski 2000). the inclusion of an environmental clause, 108 of 1996, in the bill of rights chapter of the south african constitution was a major step in the development of substantial environmental law in south africa (glazewski 2000). not only does the environmental clause constitute statements that clearly promotes conservation, these have also been further concretised in the set of environmental management principles underpinning the national environmental management act, 107 of 1998 (nema) (glazewski 2000). in effect, these inclusions in law had and will have a substantial constructive effect on conservation of sensitive habitats and endangered species in south africa in future. some species, such as the red listed species, are protected in various ways in the schedules of ordinances of the provinces in south africa. such a system is thought to be regionally adaptable to local needs and ecological circumstances (glazewski 2000). the four nature conservation ordinances that applied to the former four provinces in south africa, still apply in many cases to the nine provinces of south africa (see glazewski 2000). in practice the above necessitates a constant vigilance by the scientific community to monitor the status of species in each province and thus terblanche 1.qxd 2005/12/09 11:25 page 76 issn 0075-6458 77 koedoe 46/2 (2003) demands a sophisticated administrative and technical infrastructure that many of the under-resourced provinces lack (glazewksi 2000). the particular challenges of the monitoring of insect species in the vast south african countryside should be appreciated here. in the case of chrysoritis (which already belongs to a well-known insect group, the butterflies) neither the scientific community nor the conservation authorities has the manpower and funds to monitor the status of the rare species regularly enough to follow progress or decline sufficiently, not even in nature reserves or national parks. for perspective, note that for many of the nature reserves and national parks in south africa a species list of butterflies (the most baseline information) does not exist. to complicate matters, some of the chrysoritis species, such as chrysoritis endymion, are not seen every year despite visits by lepidopterists at the right time of the year. to identify the more rare chrysoritis species in the field and even in the laboratory, takes a specialist – a skill that would be nearly impossible in most cases for the already overburdened conservation officials to acquire. one possible solution is to involve the insect collectors (in this case the butterfly collector) in the monitoring as well as the location of the insect (butterfly) habitats. an important issue, inevitably touched on in many of the above discussions regarding legislation that seriously needs to be addressed, is the role of the butterfly collector. therefore, this is an area that needs urgent attention. new (1997) stated that it is vital to sustain the symbiosis and co-operation between professional scientists and amateurs, as efficient assessment of many butterfly populations can only be enhanced in this way. armstrong (1998) gives a valuable perspective on the importance of the contribution of the amateur lepidopterist but also the vital role of conservation authorities. the information that could be made available by bona fide amateur butterfly collectors for properly conserving butterflies and ecosystems by conservation authorities have been very valuable in the past. this is relevant for the chrysoritis species, e.g., the new localities that have been found for chrysoritis dicksoni. in south africa, there is no evidence of any butterfly populations being destroyed by collecting. it is recommended that collection take place strictly for scientific purposes, such as for sem studies, at any of the known localities of chrysoritis dicksoni. collection of more specimens at these localities, for purposes of expanding collections, will be of no value. the same applies to other localised butterflies from the genus orachrysops (orachrysops niobe and orachrysops ariadne). it is not certain whether the conservation legislation would be better served by a national body than legislation by the different provinces or at least a streamlined coordination from a national level. the issuing of permits should be streamlined and a more pragmatic approach should be adapted for facilitating collectors who wants to make a contribution towards the knowledge on lepidoptera. as stated in glazewski (2000): “much remains to be done particularly as regards the co-ordination of the administration of a diffuse set of laws”. there remains, as pointed out by steenkamp & stein (1999), a number of strategic issues with which the conservation establishment will have to deal if they want to ensure ongoing civil society participation in environmental decision-making. the conservation efforts in connection with aloeides dentatis and orachrysops niobe have led to an awareness that will possibly enhance the conservation of all the localised lycaenid butterflies in south africa. personal experience is that since the application of the law in the case of orachrysops niobe, people, including landowners and developers, have been much more aware of possible red listed butterfly species and possible unique habitats. to what extent, remains to be seen. biodiversity and the quality of such data in environmental impact assessments deserve attention beyond the scope of this overview. another key problem in saving threatened butterfly species is the notion that the habitat must be the last known habitat before the protection really becomes urgent. conservation of butterblanche 1.qxd 2005/12/09 11:25 page 77 koedoe 46/2 (2003) 78 issn 0075-6458 terflies (and other insects) should be seen in a much wider integrated sense of conserving metapopulations. education and awareness without education and public awareness the loss of touch with the environment, in which most modern and developing societies seem to find themselves, would continue. conservation in south africa has shifted in recent years from a protectionist approach towards one that involves the community in conservation (wynberg 2002). butterflies such as chrysoritis should be taken to the community. education, and use of a more indigenous curriculum in schools is needed to appreciate the existence of biological intricacies such as the ecology of chrysoritis species in africa. nature reserves such as the alice glockner nature reserve and suikerbosrand nature reserve provide excellent opportunities for such educational programmes and should be managed as such. in the case of chrysoritis aureus, the efforts of gauteng nature conservation to enhance co-operation in research and conservation have been most valuable. hopefully, these initiatives will prove valuable in a country that has many socio-economic issues that have to be addressed urgently. encouraging is that the above conservation efforts are directed at the ‘small things’ in natural veld, a possible source for ecotourism and education – very poorly exploited up to now. synecological studies, which are conducted to conserve the butterfly, could also benefit the development of the alice glockner nature reserve as a whole, addressing other aspects of biodiversity apart from butterfly conservation. at present the management of urbanisation close to the alice glockner nature reserve deserves further attention, especially with regard to community involvement and public awareness. the involvement of the younger generation (school learners and students), if only to start collecting butterflies in a back yard, may mean much more to science in the end, if it results, for instance, in new locality records of butterflies—or even new scientists—than no involvement caused by too strict legislation. in the end such involvement will enhance the conservation of butterflies (or other insects) rather than decrease the chances of their survival. a fear that the layman collector might collect butterflies such as chrysoritis aureus and destroy colonies of the butterfly is probably overrated for the following three reasons. the butterfly is easy to recognise by anybody with a good reference guide and reasonable experience of lepidoptera, but unlikely to be found by a novice. furthermore, the confirmed localities are clearly protected in nature reserves, especially the established alice glockner nature reserve. any lepidopterist that would want to survey butterflies in such a nature reserve would have to submit a research proposal (which always foreruns the issuing of any permit), stating that they know about the butterfly and will avoid to collect any of these. integrating science,environmental management, policies, law and education the integration of the efforts of scientists, the public sector and the government can only be successful if proper communication takes place and therefore proper forums are available for such communication. the journal metamorphosis (journal of the lepidopterists’ society of southern africa) has acted as a forum for debate in the past (e.g. de wet 1990; vlok 1993; woodhall 1994) between the lepidopterists and conservation authorities, with regard to the issue of collecting butterflies and approaches in conservation. the discussions between conservation authorities, researchers and other stakeholders on the issuing of permits for collecting insects as well as the law governing the protection of material collected in south africa, was initiated at the 13th congress of the entomological society of southern africa in pietermaritzburg were encouraging. another important event that coincided with the 13th congress was a symposium on arthropod diversity and conservation in southern africa, so that emphasis was also terblanche 1.qxd 2005/12/09 11:25 page 78 issn 0075-6458 79 koedoe 46/2 (2003) on insect conservation and not only those insects of economic agricultural importance (not that it is not all part of environmental management). conclusions since the genus contains a number of localised populations and subpopulations of significant biological and aesthetic value, chrysoritis genus should be seen as endemic as a whole. it is clear that the genus contains bioindicators that should be verified according to the categories set out by mcgeoch (1998). since the butterfly populations of chrysoritis tend to be localised, it is especially their role as ecological indicators and biodiversity indicators that are worth considering. the scales within which the chrysoritis species could be used are important. on a larger geographical scale their dwindling number of populations is certainly indicative of anthropogenic pressures on sensitive habitats. despite the fact that butterflies are such a relatively well studied group of insects in south africa, limitations in our knowledge still exist even on the taxonomic and distributional level. being a relatively well-known invertebrate group, the taxonomy of butterflies such as chrysoritis has reached a state where revisions could easily result in a number of unnecessary names. it is recommended that the relationships between the entities are carefully analysed in such a manner that the work can be traced and repeated. more specific distribution records will be a valuable aid for resolving the taxonomy and responses of such habitat-sensitive butterflies to environmental changes. legislation should facilitate rather than limit the valuable role that the amateur may play to contribute in the gathering of distribution records. in turn, the amateur lepidopterists should adapt and make an effort to include unknown localities, apart from monitoring butterflies at their well-known localities. this information should also be readily available to conservation authorities. a system in which the amateur fraternity also takes responsibility— aided by codes of conduct—to maintain optimal co-operation between the government and the ngo should be in place. species such as chrysoritis dicksoni should be protected by law (also from collecting) at their known and conserved localities. distribution records are not in any atlas format, a fact which limits our understanding of especially a genus such as chrysoritis that contains a number of highly localised and endemic butterflies. a southern african butterfly atlas would be appropriate and in fact a necessity, since it would enhance the understanding of the biogeography of a group of butterflies that might be good bioindicators on a large geographical scale. it is to be hoped that butterflies, that seems to be ideal for atlas work, would be included as part of the biodiversity and conservation for south african invertebrates initiative (hamer & underhill 2001) or any other similar initiatives. the red listing of localised (myrmecophilous) butterflies such as the chrysoritis is in need of urgent review in the light of the most recent iucn categories (iucn 2001) that were adopted in 1994. the environmental management priorities are linked to applied ecology, but the infrastructure to build the necessary bridges in practice, is often lacking. a sound ecosystem approach in conservation biology lies in the intersection between ecological, socio-economic and institutional perspectives (meffe & carol 1997). this cannot be more true for the conservation of chrysoritis species. it is therefore clear that the prosperity of chrysoritis species not only depends on our understanding of its interactions in nature in the context of communities, but also of our human interactions as scientists, ngo members, policymakers, law makers, educators, environmentalists and citizens in whatever capacity— especially when it comes to the integration of our efforts. the above that the organisations/institutions involved with conservation strategies, the monitoring ideals and the legislation, should appreciate the challenges particular to insect conservation. terblanche 1.qxd 2005/12/09 11:25 page 79 koedoe 46/2 (2003) 80 issn 0075-6458 acknowledgements the comments of three anonymous reviewers greatly improved the manuscript. references acocks, j.p.h. 1975. veld types of south africa. 2nd ed. memoirs of the botanical survey of south africa no. 40. acocks, j.p.h. 1988. veld types of south africa. 3rd ed. memoirs of the botanical survey of south africa 57: 1–146. armstrong, a.j. 1998. letters to the editor. metamorphosis 9(3): 141–143. bayer, m.b. 1999. haworthia revisited: a revision of the genus. hatfield: umdaus press. bredenkamp, g.j. & n. van rooyen. 1996. rocky highveld grassland. p. 39. in: low, a.b. & a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. claassens, a.j.m. 1994. recent observations on lepidoptera. metamorphosis 5(2): 53–54. claassens, a.j.m. 2000. the butterflies of the cape peninsula: a comprehensive guide. cape town: tafelberg. clark, g.c. & dickson, c.g.c. 1971. life histories of the south african lycaenid butterflies. cape town: purnell. cottrell, c.b. 1985. the absence of coevolutionary associations with capensis floral element plants in the larval/plant relationships of south western cape butterflies. pp. 115–124. in: vrba, e.s. (ed.). species and speciation. transvaal museum monograph no. 4. cowling, r.m., d.m. richardson & s.m. pierce (eds.). 1997. vegetation of southern africa. cambridge: university press. de wet, j.i. 1987. die bewaring van skoenlappers van die genus charaxes en van erikssonia acraeina in die transvaal. proceedings of the first lepidoptera conservation symposium, roodepoort: lepidopterists’ society of southern africa: 5–7. de wet, j.i. 1990. surveys of lepidoptera on provincial nature reserves. metamorphosis 27: 8–10. duke, n., j. saunders & c. saunders. 1999. a checklist of insects of swaziland. mbabane: the conservation trust of swaziland and the natural history society of swaziland. gigon, a., r. langenauer, c. meier & b. nievergelt. 2000. blue lists of threatened species with stabilised or increasing abundance: a new instrument for conservation. conservation biology 14(2): 402–413. glazewski, j. 2000. environmental law in south africa. durban: butterworths. granger, j.e. & g.j. bredenkamp. 1996. alti mountain grassland. p. 49. in: low, a.b. & a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. hamer, m.l. & l. underhill. 2001. improved biodiversity conservation through survey, atlasing, and conservation evaluation of invertebrates in south africa. proceedings of the 13th entomological congress of the entomological society of southern africa. heath, a. 1997a. a review of african genera of the tribe aphnaeini (lepidoptera: lycaenidae). metamorphosis: occasional supplement 2: 1–60. heath, a. 1997b. myrmecophily and the male genitalia of african lycaenidae: a preliminary discussion. metamorphosis: occasional supplement 3: 89–97. heath, a. 2001. new synonymies and taxonomic notes on the genus chrysoritis (butler) (lepidoptera: lycaenidae). metamorphosis 12(3): 85–98. henning, g.a. 1986a. research at suikerbosrand nature reserve: 7 sept. unpublished report for the transvaal nature conservation division: lepidopterists’ society of southern africa. henning, g.a. 1986b. research at suikerbosrand nature reserve: 2 dec. unpublished report for the transvaal nature conservation division: lepidopterists’ society of southern africa. henning, g.a. 1987. research at suikerbosrand nature reserve: 1st quarter. unpublished report for the transvaal nature conservation division: lepidopterists’ society of southern africa. henning, g.a. 1997. butterfly conservation in south africa. metamorphosis: occasional supplement 3: 120–128. henning, g.a. & s.f. henning. 1995. updating the status of south african red data butterfly species. metamorphosis 6(3): 96–98. henning, s.f. & g.a. henning. 1989. south african red data book: butterflies. pretoria: council for scientific and industrial research, national scientific programmes unit. (south african national scientific programmes report; no. 158.) iucn. 2001. iucn red list categories and criteria: version 3.1. gland: iucn species survival commission. keller, w.c.f., c.r. amodio & s. stänz. 1997. a preliminary and annotated list of lepidoptera butterflies (papilionoidea) and moths (hesperiidae, sphingidae, saturniidae), recorded from the kingdom of lesotho, south africa. metamorphosis 8(3): 109–130. kloppers, j., & g. van son. 1978. butterflies of the kruger national park. pretoria : national parks board. terblanche 1.qxd 2005/12/09 11:25 page 80 issn 0075-6458 81 koedoe 46/2 (2003) kohm, k., p.d. boersma, g.k. meffe & r. noss. 2000. editorial: putting science into practice and practice into science. conservation biology 14 (3): 593–594. low, a.b. & a.g. rebelo. (eds.). 1996. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. mcgeoch, m.a. 1998. the selection, testing and application of terrestrial insects as bioindicators. biological reviews 73(1): 181–201. mcgeoch, m.a. 2002. insect conservation in south africa: an overview. african entomology 10(1): 1–10. meffe, g.k. & c.r. carol. 1997. principles of conservation biology. sunderland: sinauer. new, t.r. 1997. butterfly conservation. melbourne: oxford university press. o’connor, t.g. & g.j. bredenkamp. 1997. grassland. pp. 215–257. in: cowling, r.m., richardson, d.m. & pierce, s.m. (eds.) 1997. vegetation of southern africa. cambridge: university press. owen-johnston, n.k. 1991. thoughts and observations on the genus poecilmitis butler (lepidoptera: lycaenidae) in the transvaal. metamorphosis 2(3): 49–50. pringle, e.l., g.a. henning & j.b. ball. (eds.). 1994. pennington’s butterflies of southern africa. cape town: struik. pryke, s.r. & m.j. samways. 2001. width of grassland linkages for the conservation of butterflies in south african afforested areas. biological conservation 101: 85–96. rand, d.b., a. heath, t. suderman & n.e. pierce. 2000. phylogeny and life history evolution of the genus chrysoritis within the aphnaeini (lepidoptera: lycaenidae), inferred from mitochondrial cytochrome oxidase i sequences. molecular phylogenetics and evolution 17: 85–96. roos, p. & g.a. henning. 2000. the heidelberg copper butterfly. african wildlife 54(2): 24 –27. rutherford, m.c. & r.h. westfall. 1994. biomes of southern africa: an objective categorisation. memoirs of the botanical survey of south africa, no 63. samways, m.j. 1993. threatened lycaenidae of south africa. pp. 62–69. in: new, t.r. (ed.). conservation biology of lycaenidae (butterflies). gland: iucn species survival commission. (occasional paper of the iucn species survival commission; no. 8.) samways, m.j. 1994. insect conservation biology. london: chapman & hall. steenkamp, c. & stein, r. 1999. the brenton blue saga: a case study of south african biodiversity conservation. parkview: ewt. terblanche, r.f. 1991. trip to golden gate in the orange free state. metamorphosis 2(3): 46–48. terblanche, r.f., g.f. smith & j.d. theunissen. 1993. did scott typify names in haworthia (asphodelaceae: alooideae)? taxon 42(1) : 91– 95. terblanche, r.f., t.l. morgenthal & s.s. cilliers. 2003. the vegetation of three localities of the threatened butterfly species chrysoritis aureus (lepidoptera: lycaenidae). koedoe 46(1): 73–90. terblanche, r.f. & j.c. taylor. 2000. notes on the butterflies of witsand—a unique terrestrial island in the northern cape province, south africa—with special reference to two red data book butterfly species. metamorphosis 11(3): 122–131. vári, l., d.m. kroon & m. krüger. m. 2002. classification and checklist of the species of lepidoptera recorded in southern africa. chatswood: simple solutions. vlok, j. 1993. in defence of cape nature conservation – a reply to steve woodhall. metamorphosis 4(4): 180–181. whitten, t., d. holmes & k. mackinnon. 2001. conservation biology: a displacement behaviour for academia? conservation biology 15(1): 1–3. woodhall, s.e. 1994. cape nature conservation – a reply to j. vlok. metamorphosis 5(2): 77–78. woodhall, s.e. 1996. the plight of poecilmitis lyncurium (trimen, 1868). metamorphosis 7(1):34–36. wynberg, r. 2002. a decade of biodiversity conservation and use in africa: tracking progress from the rio earth summit to the johannesburg world summit on sustainable development. south african journal of science 98: 233–243. terblanche 1.qxd 2005/12/09 11:25 page 81 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <feff005500740069006c0069007a006500200065007300740061007300200063006f006e00660069006700750072006100e700f5006500730020007000610072006100200063007200690061007200200064006f00630075006d0065006e0074006f0073002000500044004600200063006f006d00200075006d00610020007200650073006f006c007500e700e3006f00200064006500200069006d006100670065006d0020007300750070006500720069006f0072002000700061007200610020006f006200740065007200200075006d00610020007100750061006c0069006400610064006500200064006500200069006d0070007200650073007300e3006f0020006d0065006c0068006f0072002e0020004f007300200064006f00630075006d0065006e0074006f0073002000500044004600200070006f00640065006d0020007300650072002000610062006500720074006f007300200063006f006d0020006f0020004100630072006f006200610074002c002000520065006100640065007200200035002e00300020006500200070006f00730074006500720069006f0072002e00200045007300740061007300200063006f006e00660069006700750072006100e700f50065007300200072006500710075006500720065006d00200069006e0063006f00720070006f0072006100e700e3006f00200064006500200066006f006e00740065002e> /dan <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> /nld <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> /esp <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> /suo <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> /ita <feff00550073006100720065002000710075006500730074006500200069006d0070006f007300740061007a0069006f006e00690020007000650072002000630072006500610072006500200064006f00630075006d0065006e00740069002000500044004600200063006f006e00200075006e00610020007200690073006f006c0075007a0069006f006e00650020006d0061006700670069006f00720065002000700065007200200075006e00610020007100750061006c0069007400e00020006400690020007000720065007300740061006d007000610020006d00690067006c0069006f00720065002e0020004900200064006f00630075006d0065006e00740069002000500044004600200070006f00730073006f006e006f0020006500730073006500720065002000610070006500720074006900200063006f006e0020004100630072006f00620061007400200065002000520065006100640065007200200035002e003000200065002000760065007200730069006f006e006900200073007500630063006500730073006900760065002e002000510075006500730074006500200069006d0070006f007300740061007a0069006f006e006900200072006900630068006900650064006f006e006f0020006c002700750073006f00200064006900200066006f006e007400200069006e0063006f00720070006f0072006100740069002e> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 dippenaar microstigmatidae.qxd habitat preferences and seasonal activity of the microstigmatidae from ngome state forest, south africa (arachnida: araneae) a.s. dippenaar-schoeman, m. van der merwe and a.m. van den berg dippenaar-schoeman, a.s., m. van der merwe and a.m. van den berg. 2006. habitat preferences and seasonal activity of the microstigmatidae from ngome state forest, south africa (arachnida: araneae). koedoe 49(1): 85–89. pretoria. issn 0075-6458. as part of the south african national survey of arachnida (sansa) inventories are underway to determine the arachnid biodiversity of various biomes. a survey of groundliving spiders was conducted over a year period (1992-1993) at ngome state forest, kwazulu-natal, south africa. spiders were collected with 180 pitfall traps from five different habitat types ranging from indigenous forest to a pine plantation and grassland. a total of 186 specimens of the rare mygalomorphae family microstigmatidae were sampled, representing 2 % of the total spider fauna collected during the study. two species, microstigmata longipes (lawrence) and m. zuluensis (lawrence), occur sympatrically. both species were more active in the indigenous forest, and were absent or present in low numbers in the open grass and pine plantation. microstigmata zuluensis was slightly more abundant (59 % of total) and was recorded from three of the five habitats, while m. longipes (41 %) was found in four of the habitat types. both species were active throughout the year with the lowest numbers recorded in winter (june to august). the males of m. longipes peaked in november while those of m. zuluensis peaked in april. key words: afromontane forest, araneae, biodiversity, microstigmatidae, microstigmata, spiders, south africa national survey of arachnida. a.s. dippenaar-schoeman, arc-plant protection research institute, private bag x134, queenswood 0121, republic of south africa / department of zoology and entomology, university of pretoria, 0002 pretoria, republic of south africa; a.m. van den berg, arc-plant protection research institute, private bag x134, queenswood 0121, republic of south africa; m. van der merwe, department of zoology and entomology, university of pretoria, 0002 pretoria, republic of south africa (present address: the joint science department of the claremont colleges, w.m. keck science centre, 925 n. mills ave, claremont, ca 91711-5916, usa). issn 0075-6458 85 koedoe 49/1 (2006) introduction the microstigmatidae is a small family of spiders represented by seven genera and 13 species known from africa, central and south america (platnick 2005). in africa it is represented by a single genus microstigmata, which is known only from south africa (dippenaar-schoeman 2002; griswold 1985). in a revision of the genus, griswold (1985) recognised six species with their distribution restricted mainly to the forest regions of the eastern parts of south africa. the microstigmata are small (4–13 mm), free-living mygalomorph spiders. they are the only members of the mygalomorphae in the afrotropical region that do not live in silk-lined burrows or retreats. they are wanderers and found when turning stones and logs or sifting leaf litter and humus. specimens are usually encrusted with earth due to sand particles adhering to their clavate body setae. although widely distributed throughout the eastern parts of south africa, they are shy animals and are rarely collected. little is known about their behaviour and natural history (griswold 1985). during a 12-month survey of the cryptofauna at ngome state forest in kwazulu-natal, south africa, 186 microstigmatids were samdippenaar microstigmatidae.qxd 2006/04/17 10:51 pm page 85 pled with pitfall traps. this represented about 2 % of the total number of spiders collected during the study (van der merwe et al. 1996). as very little information is available on the ecology of most mygalomorph spiders, data on the habitat preferences and seasonal activity of the two microstigmata species collected are presented here. this is the third paper on the ground-living spider fauna of the ngome state forest (van der merwe 1994; van der merwe et al. 1996) and the first quantitative survey of microstigmatids in south africa. this survey forms part of the south african national survey of arachnida (sansa) that was initiated in 1997. the main aim of this umbrella project is to make an inventory of the arachnid fauna of south africa (dippenaar-schoeman & craemer 2000). thrusts within sansa are focused on inventories of the arachnid fauna of the different floral biomes (here the forest biome), provinces, conserved areas and agro-ecosystems. study area ngome state forest (27º49's; 31º26'e) is situated on the escarpment of northern kwazulu-natal, south africa, at an altitude of 405–1365 m above sea level. rainfall averages 1507 mm/year, with january the wettest (227 mm) and june the driest (26 mm). the mean maximum temperatures are 26.6 ºc (january/february) and 18.6 ºc (june/july). the five different habitat types sampled were: ‘open grass’ consisting of grass surrounded by indigenous forest; ‘open forest’ comprising closed canopy indigenous forest characterised by the absence of significant undergrowth; ‘dense forest’ consisting of closed canopy indigenous forest with dense undergrowth; ‘ecotone pine’ with mature pine plantation surrounded and penetrated by indigenous forest vegetation, and a pine plantation not directly surrounded by indigenous forest. for more detail on the habitat types see van der merwe et al. (1996). methods pitfall traps (n = 180) consisting of a plastic container (10 cm diameter) sunk to ground level with a funnel leading to a honey jar, and filled with 70 % alcohol–5 % glycerol mixture, was used. the traps were spaced 5 m apart in a 3x3 grid pattern with four grids (36 pitfall traps) placed in each of the five habitats. traps were continuously open and emptied in the middle of each month, from january 1992 to january 1993. voucher specimens are deposited in the national collection of arachnida at the arc-plant protection research institute, pretoria. statistical analysis were conducted using the statistical program genstat (2000) to determine if significant differences existed in the abundance of the two species in the different habitat types, as well as during different seasons, using 2x3, 2x4 and one sample chi-squared tests. results species numbers a total of 9 369 spiders were collected during the survey, of which 186 belonged to the microstigmatidae. two species, m. longipes (lawrence) and m. zuluensis (lawrence), occur sympatrically. a total of 110 individuals of m. zuluensis were collected, representing 59 % of all microstigmatids collected, while 76 specimens of m. longipes were collected, representing 41 % of the total (table 1). habitat preferences microstigmata zuluensis was collected from three of the five habitats (table 1). they were absent from the open grass and pine plantation, and more active (n = 45) in the open forest. they represented 41 % of the total, followed by the dense forest (n = 42) with 38 %, and ecotone pine (n = 23) with 21 %. microstigmata longipes was active in four of the five habitats, being absent only from the open grass (table 1). they were more active in the dense forest (n = 37) representing 49 % of the total, followed by the open koedoe 49/1 (2006) 86 issn 0075-6458 dippenaar microstigmatidae.qxd 2006/04/17 10:51 pm page 86 forest (n = 32) with 42 % and 9 % in the ecotone pine and pine plantations. both species prefer closed canopy indigenous forest. a total of 79 % of microstigmatids were collected from the two indigenous forest habitats compared to the 21 % from the ecotone pine and pine plantation. the cover of the forest floor plays a less important role; 41 % in total were active in the forest habitat characterised by the absence of significant undergrowth, compared to the 43 % collected from forest with dense undergrowth. the chi-squared tests indicated that the relative activity of the two microstigmata spp. differed significantly in open forest, dense forest and ecotone pine at the 0.05 test level (χ² = 5.991, p <0.05), and that the distribution of the two species differed significantly between the dense forest and ecotone pine (χ² = 5.737, p <0.05). seasonal activity microstigmata zuluensis was active throughout the year (except in july), but their numbers fluctuated to show an increase in surface activity during autumn and early winter (april to june) when 42 % were collected, issn 0075-6458 87 koedoe 49/1 (2006) table 1 microstigmata spp. collected from 180 pitfall traps from five habitat types at the ngome state forest, kwazulu-natal, south africa during 1992-1993 species open grass pine plantation ecotone dense forest open forest total % m. zuluensis 0 0 23 42 45 110 59 m. longipes 0 2 5 37 32 76 41 total 0 2 28 79 77 186 100 fig. 1. seasonal activity of microstigmata zuluensis collected from 180 pitfall traps at ngome state forest, kwazulu-natal, south africa during 1992-1993. dippenaar microstigmatidae.qxd 2006/04/17 10:51 pm page 87 with a peak in april (n = 28). their numbers declined to a low of 11 % during the winter months (july-august) and increased again in the spring and summer to 26 %. microstigmata longipes was active throughout the year with peaks in february (n = 15) and november (n = 20). their activity fluctuated from autumn when 16 % were recorded, to a low (8 %) in winter. activity increased again to reach a high of 40 % in the spring and 37 % in summer. as expected of pitfall trapping, more males (55 %) were sampled in both species, followed by immature specimens (32 %) and females (13 %). males of m. zuluensis were active during eight months of the year (fig. 1). they were active in autumn and early winter with a peak (n = 21) in april when 19 % of the specimens were collected and second peaks in october (n = 9) and december (n = 8). juveniles were active throughout the year (except july) with a slight peak in april (n = 7). the females were collected irregularly during five months of the year. males of m. longipes were active five months of the year, and their numbers peaked in november (n = 19; 20 %) but were absent from samples between february and august. juveniles were absent during four months of the year with a peak in february (n = 10). female numbers peaked in february and march (n = 5; 17 %) (fig. 2). discussion and conclusion the two species m. longipes and m. zuluensis are typical cryptofauna of indigenous forest habitats (griswold 1985; lawrence 1952). although the two species occurred sympatrically and both were active throughout the year their reproductive periods differ. the males of m. zuluensis peaked in april, compared to the males of m. longipes that peaked in november. koedoe 49/1 (2006) 88 issn 0075-6458 fig. 2. seasonal activity of microstigmata longipes collected from 180 pitfall traps at ngome state forest, kwazulu-natal, south africa during 1992-1993. dippenaar microstigmatidae.qxd 2006/04/17 10:51 pm page 88 both species preferred the indigenous forest habitats with either dense or low undergrowth where 85 % of the specimens were collected above the open grass habitat or pine forest where only 15 % of the two species were found. based on general collection information and distribution records griswold (1985) identified m. zuluensis as a more tolerant species occurring also in habitats outside indigenous forests. this observation was confirmed in this study where 12.4 % of m. zuluensis was collected from ecotone pine compared to the 3.8 % of m. longipes found outside the indigenous forest. conservation biologists are starting to recognise the importance of the invertebrate component in the functioning of healthy ecosystems. of prime importance in any conservation scheme is the identification of species sensitive to changing environmental factors, as found here. in many areas of africa the planting of exotic trees has supplanted indigenous forest. this has an effect on plant structure at the ground level, which could have an effect on the community structure of the ground-living spider fauna. since spiders are an important component of the forest floor ecosystem this could have significant effects on other invertebrate groups and plants as well. as only a fraction of south africa’s area (0.2 %) is covered by high evergreen forest (huntley 1984) the conservation of these undisturbed indigenous forests with their cryptofauna is of utmost importance and should receive high priority for conservation efforts. acknowledgement we thank the forester michael peter at ngome state forest for his assistance as well as students who assisted with field work. helene muller of the arcbiometry unit was responsible for the statistical analysis of data. references dippenaar-schoeman, a.s. 2002. the baboon and trapdoor spiders of southern africa: an identification manual. pretoria: agricultural research institute. (plant protection research institute handbook no. 130). dippenaar-schoeman, a.s. & c. craemer. 2000. the south african national survey of arachnida (sansa). plant protection news 56: 11-12. genstat for windows. 2000. release 4.2. fifth edition. oxford: vsn international. griswold, c.e. 1985. a revision of the african spiders of the family microstigmatidae (araneae: mygalomorphae). annals of the natal museum 27: 1-37. huntley, b.j. 1984. characteristics of south african biomes. in: booysen p. de v & n.m. tainton (eds). ecological effects of fire in south african ecosystems. berlin: springer. lawrence, r.f. 1952. new spiders of the eastern half of south africa. annals of the natal museum 12: 183-226. platnick, n.i. 2005. the world spider catalog, version 5.5. american museum of natural history, online at http://research.amnh.org/entomology/ spiders/catalog/index.html van der merwe, m. 1994. a comparative survey of cursorial spider communities in indigenous afromontane forests and in pine plantations. m.sc. thesis, university of pretoria, pretoria. van der merwe, m., a.s. dippenaar-schoeman & c.h. scholtz. 1996. diversity of ground-living spiders of ngome state forest, kwazulu/natal: a comparative survey in indigenous forest and pine plantations. african journal of ecology 34: 342-350. issn 0075-6458 89 koedoe 49/1 (2006) dippenaar microstigmatidae.qxd 2006/04/17 10:51 pm page 89 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /all /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /warning /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /fra <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> /enu (use these settings to create pdf documents with higher image resolution for improved printing quality. the pdf documents can be opened with acrobat and reader 5.0 and later.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308000200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e30593002537052376642306e753b8cea3092670059279650306b4fdd306430533068304c3067304d307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e30593002> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 brown ingleside.qxd the vegetation of the farms ingleside and welgedacht of the mountain zebra national park, eastern cape l.r. brown and h bezuidenhout brown, l.r. and h bezuidenhout. 2005. the vegetation of the farms ingleside and welgedacht of the mountain zebra national park, eastern cape. koedoe 48(2): 23–42. pretoria. issn 0075-6458. south africa is well known for its semi-arid lowland areas that have a distinct flora and species composition. because ecosystems react differently to different management practices, it is important that a description and classification of the vegetation of an area be done. as part of a vegetation survey programme for the newly acquired farms incorporated into the mountain zebra national park, the vegetation of the ingleside and welgedacht sections were surveyed following the braun-blanquet approach. from a twinspan classification, refined by braun-blanquet procedures, 10 shrub and grassland plant communities, which can be grouped into seven major groups, were identified. a classification and description of these communities, as well as a vegetation map are presented. the diagnostic species as well as the prominent and less conspicuous species of the tree, shrub, herb and grass strata are outlined. the area generally comprises lowland communities and higher-lying communities. the lower-lying communities consist mainly of two communities and comprise the largest proportion of the area in hectares. in contrast, the higher-lying communities are more diverse with specific habitats. using the ecological index method the veld condition and grazing capacity were calculated for each community and the total study area. large sections of the lowland areas are overgrazed due to previous farming grazing practices while the higher-lying areas that were less accessible to the animals are in a slightly better condition. overall this has resulted in the area generally being degraded within a high grazing capacity of 30.1 ha/lsu. keywords: braun-blanquet, plant communities, twinspan, vegetation classification, veld condition, grazing capacity. l.r. brown , applied behavioural ecology & ecosystem research unit, department of nature conservation, unisa, private bag x6, florida, 1710 republic of south africa; h bezuidenhout, arid ecosystems research unit, conservation services, sanparks, p.o. box 110040, hadison park, kimberley, 8306 republic of south africa. issn 0075-6458 23 koedoe 48/2 (2005) introduction the prevention of the increasing loss of biodiversity is a daunting challenge facing the world today. humans continuously alter their environment to suit their needs resulting in a large number of ecosystems being destroyed every year. south africa is richly endowed with endemic plant and animal species and is the third most biologically diverse county in the world (hall-martin 2003). the establishment of protected areas within which large ecosystems are conserved, is increasingly seen as a solution to the prevention of ecosystem loss. however, to be able to conserve our natural resources for the current and future generations, it is important that the different ecosystems be studied and managed on a scientific basis. south africa has a well-established system of national parks situated within the various biomes to protect and manage the ecological integrity of the different ecosystems for present and future generations. one such park, the mountain zebra national park (mznp) was established in 1937 to protect the cape mountain zebra (equus zebra zebra) population that at that time numbered only 11 (brown & bezuidenhout 2000). the size of the park was too small to sustain a viable population of these animals resulting in the park area being extended in 1964 (carruthers 2003). since 1996, the park size was again extended and various farms adjacent to the park have been procured or are in the process of being bought and incorporated into the park. these new areas have not only resulted in the park more than doubling its size from its previous 6 536 ha, to approximately 21 000 ha, but include new ecosystems and make the re-introduction of various large mammals such as cape buffalo, black rhino and gemsbok possible. because ecosystems react differently to different management practices (bredenkamp 1982; bezuidenhout 1993), it is important that a description and classification of the vegetation of an area is undertaken (van rooyen et al. 1981). vegetation is the most physical representation of an ecosystem. any change in the ecosystem (pollution, development, grazing, droughts) is first observed in changes in the vegetation. it is therefore of utmost importance that vegetation is studied in detail to determine species and population composition in order to compile an inventory of the flora. this will not only enable us to sustainably manage and protect our natural resources, but also to determine linkages between ecosystems and detect any changes in the environment. it is widely recognised that a detailed description, identification, classification and mapping of the vegetation forms the basis for sound land-use planning and management (tueller 1988; fuls et al. 1992; fuls 1993; bezuidenhout 1996, brown 1997). although extensive studies on various aspects of the vegetation of the current mznp have been conducted by van der walt (1980), novellie (1990a, 1990b) and novellie & bezuidenhout (1994), little is known about the vegetation and habitat status of these newly acquired areas. the main aim of this study was therefore to describe and map the plant communities of the welgedacht and ingleside farms, which have already been acquired by the south african national parks. this study forms part of a larger and longterm research project undertaken by south african national parks and unisa. the aim of this project is to describe and map the vegetation of each of the new farms and assess the habitats of these areas for the establishment of large herbivores. this will enable management of the mznp to take scientifically based decisions on the management of each of these areas before incorporating it into the larger mznp area. all the data will then be incorporated into the existing management plan for the mznp. the study area the mznp is situated approximately 12 km west from the town of cradock in the eastern cape province. it extends from latitude 32º05'–32º20's and longitude 25º23'–25º 32'e (fig. 1). the park is situated within the nama-karoo biome. this biome is the second largest biome of south africa and occupies the hotter, drier, western half of the inland plateau of south africa. though extending to areas as low as 500 m asl, it mostly lies between 1000–4000 m asl (hallmartin 2003). the park is situated in a transitional zone between the arid karoo bushveld of the western and central karoo and the drier more palatable grassveld in the east (van der walt 1980). according to acocks (1988), the area can be classified as false karroid broken veld (37), whereas hoffman (1996) classifies it as eastern mixed nama karoo (52). some isolated patches of the south-eastern mountain grassland (grassland biome) (lubke et al. 1996) are also found in the park. the low rainfall of these areas ensures that the predominant soil type of this biome is lime-rich and prone to erosion (hall-martin 2003). the welgedacht and ingleside sections (study area) comprise 1554 ha and lie approximately 11 km north of the previously proclaimed section of mznp. the study area borders onto the previously described de koedoe 48/2 (2005) 24 issn 0075-6458 rust section (brown & bezuidenhout 2000) of the mznp in the west (fig. 1). the northern part is bound by the main road to graaff reinet. the dominant vegetation is a mixture of grassy, dwarf shrubland with trees usually occurring along drainage lines and dominated by only one or two species (van der walt 1980). the northern and eastern areas of the study area comprise lower lying valley bottomlands and dry riverbeds, together with flat to undulating plateau sections. the southwestern section comprises steep to moderately steep mountain slopes of the salpeterkop mountain which is also the most prominent geological feature of the study area. land types with references to the physiography, geology and soil three land types, namely ag, ib and fc occur in the study area. according to the land type survey staff (1986) “a land type denotes an area that can be shown at 1:250 000 scale and that displays a marked degree of uniformity with respect to terrain form, soil pattern and climate”. a remarkable association between the major plant communities and the different land types has been observed in other studies (kooij et al. 1990; bezuidenhout 1993; eckhardt 1993; brown 1997). the land type unit ag refers to red apedal, freely drained, high base status soil, which is shallower than 400 mm. the predominant geology of this land type is mudstone, shale and sandstone of the balfour formation, beaufort group of the karoo supergroup (toerien 1972). a dense subdendritic drainage and dissection pattern occurs in this land type with some dolerite intrusions. the slightly undulating plains are dominated by the valley bottomland terrain unit with hutton the dominant soil form (soil classification working group 1991). this soil is shallow (150–400 mm), rocky, and the sand grade is fine (land type survey staff 1999). issn 0075-6458 25 koedoe 48/2 (2005) fig. 1. location of the study area. the f land type refers to pedologically young landscapes which are predominantly rocky and not alluvial or aeolian. the fc unit refers to land where lime occurs regularly in upland and valley bottomland soil (land type survey staff 1986). soil forms that epitomise this land type are glenrosa and oakleaf. the geology consists of mudstone, shale and sandstone of the beaufort group of the karoo supergroup with rare dolerite intrusions (land type survey staff 1999). two topographical positions are prominent in this terraced landscape, namely the footslopes and valley bottomlands. the ib land type unit refers to exposed rocks that cover 60–80 % of the area. the rocky portions of ib may be underlain by soil which has qualified the unit for inclusion in another broad soil pattern had it not been for the surface rockiness. the midslopes are the most prominent topographical position in this mountainous landscape and the dominant soil-rock complex consists of rock while the glenrosa soil form is subdominant. the geology consists of dolerite with mudstone, shale and sandstone of the balfour formation, beaufort group of the karoo supergroup (land type survey staff 1999). climate the average annual rainfall for the area, as measured at the mznp weather station from july 1962 to june 2004, is 405 mm with the highest rainfall of 651 mm and lowest rainfall of 153 mm being recorded in 1976 and 1966, respectively. the general rainfall trend from 1962–2004 shows an increase in averkoedoe 48/2 (2005) 26 issn 0075-6458 fig. 2. percentage deviation from average annual rainfall at mountain zebra national park between 1962 and 2004. (rainfall year = jan. to dec.) age annual rainfall since 1962. there is no increase in rainfall amplitude but rather a decrease in rainfall extremes (fig. 2). this is also illustrated in the higher average annual rainfall of 424.5 mm for the past 10 years. the lowest rainfall recorded for this 10-year period was 213.4 mm (1992) and the highest 565.5 mm (1991). the average monthly rainfall varies from 11.3 mm in the dry winter season to 60.8 mm in the wet summer season (brown & bezuidenhout 2000). the average maximum temperature varies between 23.11–28.4 ºc in summer (september-march) and 16.2–22.7 ºc in winter (april-august), whereas the average minimum temperature varies between 5.6– 13.6 ºc in summer and 0.05–7.8 ºc in winter (fig. 3) (brown & bezuidenhout 2000). methods plant communities are conceived as types of vegetation recognised by their floristic composition. the full species composition of the different communities expresses their relationship to each other and the environment better than any other characteristic (westhoff & van der maarel 1978). in order to accomplish a detailed vegetation survey of the study site the area was stratified into physiognomic-physiographic units using 1:50 000 stereo aerial photographs. after a reconnaissance of the area sample plots were randomly located within the different homogeneous units identified from the aerial photographs. the number of sample plots allocated within each homogeneous unit depended on the size of the area, the larger the area the higher the number of sampling plots allocated to the unit. a total of 52 sample plots were randomly located within these units to ensure that all variations in the vegetation were considered and sampled. plot sizes were fixed at approximately 400 m² according to brown (1997) and brown & bezuidenhout (2000). veld condition and grazing capacity were determined for each community. the ecological index method (foran et al. 1978; vorster 1982; smit 1988), which has proved successful for formulating management practices, was used (orban 1995). after the percentage grass and forb species composition for each sample site was calculated, the species were classified into ecological categories, namely decreasers, increasers i, iia, iib and iic species, according to their reaction to utilisation (danckwerts 1989; trollope et al. 1990). with only few exceptions, the ecological status of the grasses conforms to that used by van oudtshoorn (1992) while all forbs and/or herbaceous weeds were classified as increaser iic species. the data was then incorporated into a model (graze) used to calculate the grazing capacity. this model was developed by bredenkamp (ecotrust cc ) and is described in brown (1997) who successfully applied it in the borakalalo nature reserve, north-west province. issn 0075-6458 27 koedoe 48/2 (2005) fig. 3. the mean average minimum and maximum temperatures for mountain zebra national park. sampling total floristic composition was noted for every sample plot. making visual estimates, sampling was done by means of the braun-blanquet method (mueller-dombois & ellenberg 1974). braun-blanquet cover abundance values were assigned to all plant species encountered in the relevés (table 1). plant species identification was done according to germishuizen & meyer (2003). soil classification is according to the soil classification work group (1991). trees and shrubs were distinguished from each other using the guidelines set by edwards (1983). trees are classified as rooted, woody, selfsupporting plants > 3 m high with one or a few definite trunks and shrubs are classified as rooted, woody, self-supporting, multi-stemmed or singlestemmed plants < 3 m high (adapted from edwards 1983). a minimum of 200 and a maximum of 600 step points depending on community size were done within each plant community to determine veld condition score. the habitat conditions including geology, soil forms and other edaphic factors, slope, aspect and the rockiness of the soil surface, of each relevé, were qualitatively described. this information was used in the description of the different plant communities. the coordinates of each relevé was determined and noted with the use of a gps. data processing the floristic data was analysed according to braunblanquet procedures using the bbpc suite (bezuidenhout et al. 1996). the numerical classification program twinspan (hill 1979), which is regarded as a successful approach for classification by several phytosociologists (mucina & van der maarel 1989; bredenkamp & bezuidenhout 1995; cilliers 1998) was used to derive a first approximation of the floristic data. further refinement of the classification was achieved by braun-blanquet procedures (bredenkamp et al. 1989; kooij et al. 1990; bezuidenhout 1993; eckhardt 1993; brown & bredenkamp 1994). using the phytosociological table and the habitat information gathered during the sampling period, the different plant communities were identified and described. plant communities were recognised by using diagnostic species as defined by westhoff & van der maarel (1978). diagnostic/ character species are those that are largely restricted to a community. these species do not necessarily have a high importance value. the different plant communities are described according to their dominant species. dominant species are those that are most conspicuous in the community and are high in one or more of the importance values (whittaker 1978), in this case cover and frequency. no attempt was made to formally fix syntaxa names as this is normally avoided in detailed local studies (coetzee 1983). this will only follow after extensive studies on savanna vegetation in south africa to prevent synonymy of syntaxa names. uncommon species not diagnostic for any of the communities are not included in the table, but are available upon request from the corresponding author. results classification the results are presented in a phytosociological table (table 1). the analysis resulted in the following ten plant communities (fig. 4), which may be grouped into seven major community types: 1. buddleja glomerata-rhus lucida shrubland 2. carissa macrocarpa-cenchrus ciliaris shrubland. 2.1 euclea undulata-carissa macrocarpa shrubland 2.1.1 portulacaria afra variant 2.1.2 euclea undulata variant 2.2 rhus lucida-carissa macrocarpa shrubland 3. setaria neglecta-becium burchellianum grassland 4. rhigozum obovatum-enneapogon scoparius grassland. 5. panicum maximum-acacia karroo woodland. 6. eragrostis obtusa-pentzia globosa shrubland 6.1 tragus koelerioides-pentzia globosa shrubland 6.2 cynodon incompletus-pentzia globosa shrubland 7. chloris virgata-cynodon incompletus grassland description of the plant communities in the descriptions of the different plant communities, all species groups refer to table 1 while the veld condition index and grazing capacity values are given in table 2. therefore no repeated reference will be made to tables 1 & 2 in the descriptions of the various plant communities. 1. buddleja glomerata-rhus lucida shrubland. this community is situated in the southwestern section on the escarpment and cliffs at the top of the salpeterkop mountain. the altitude was measured at 1 432 m above sea level, while the very steep eastern slopes ranges between 25–28º. rock cover varies between 55–90 % with the plant species growing between rocky outcrops and in rock crevices. this community is only 16 ha in extent. the cover of the woody layer varies between 15–40 %, and the herbaceous layer between 30–50 %. the woody layer comprises shrubs varying in height between 1.8–2.2 m while the herbaceous layer varies between 0.3–0.6 m in height. koedoe 48/2 (2005) 28 issn 0075-6458 issn 0075-6458 29 koedoe 48/2 (2005) fi g. 4 . v eg et at io n m ap o f th e in gl es id e an d w el ge da ch t s ec tio ns o f th e m ou nt ai n z eb ra n at io na l p ar k. the diagnostic species include the shrubs buddleja glomerata, ehretia rigida and the forbs ceterach cordata, hermannia vestita and hibiscus species (species group a). the vegetation is dominated by the shrubs rhus lucida (species group f) and buddleja glomerata (species group a). other species also prominent locally include the shrubs rhus longispina, grewia occidentalis (species group j), and the grasses aristida junciformis (species group f) and cenchrus ciliaris (species group j). this community shows affinity with the rhus lucida-buddleja glomerata shrubland described by brown & bezuidenhout (2000) in the de rust section and the buddleja glomerata-rhus lucida woodland described by de klerk et al. (2003) in the ebenaeser section of the mznp. no veld condition data were collected within this community due to the inaccessibility of the area to animals and because the largest section is situated on cliffs. 2. carissa macrocarpa-cenchrus ciliaris shrubland. this shrubland comprises 136 ha and is found on the steep northern and western midslopes of the salpeterkop mountain. the altitude varies between 1115–1400 m asl with large rocks covering the soil. rock cover is high varies between 50–70 %. tree cover is estimated at between 2–4 %, while the shrub stratum covers between 15–40 % of the area. the herbaceous cover is estimated at between 5–60 %. the average tree height varies between 3.5–4 m, while the average height of the shrubs is 2.5 m. the herbaceous layer is approximately 0.5 m tall. this community is characterised by the presence of the shrub carissa macrocarpa, the grass bothriochloa insculpta and the forb galenia species (species group b). although these species are diagnostic for this community, they are locally prominent in the rhigozum obovatum-enneapogon scoparius grassland (community 4). this community can be divided into the following two sub-communities: 2.1 euclea undulata-carissa macrocarpa shrubland the euclea undulata-carissa macrocarpa shrubland is situated on the steep northern and western midslopes of the salpeterkop mountain. altitude varies between 1124– 1304 m asl. the shrubland is covered with large rocks with an approximate 70 % coverage. the shrub layer is well developed and the most prominent layer within this sub-community, whereas the tree and herbaceous layers are not well developed with low cover. the shrub euclea undulata and the grass setaria verticilata (species group c) are diagnostic for this sub community. the vegetation is dominated by the shrub euclea undulata (species group c), whereas the shrubs rhigozum obovatum, rhus longispina, grewia occidentalis and the grass cenchrus ciliaris (species group j) are prominent locally. 2.1.1 portulacaria afra variant this variant (42 ha) is found on the higher northern midslopes of the salpeterkop mountain in the southern section of the study area. altitude varies between 1226–1304 m above sea level. the area comprises large to medium sized rocks covering approximately 70 % of the area. there is no soil between these rocks resulting in no herbaceous vegetation occurring between the rocks. the tree layer is approximately 3.5 m tall with a 3 % cover. the shrub layer is the most prominent layer varying in height between 2–2.5 m with a 35–40 % cover. the herbaceous layer (0.2 m) is poorly developed with a 5–18 % cover. the tree boscia oleoides and the tall shrub portulacaria afra (species group d) are diagnostic for this variant. although also present in the buddleja glomerata-rhus lucikoedoe 48/2 (2005) 30 issn 0075-6458 da shrubland (community 1), the presence of the forb cyphostemma quinatum (species group e) together with the diagnostic species (species group d) is characteristic for this community. the vegetation is dominated by the shrub euclea undulata (species group c), whereas the grass cenchrus ciliaris (species group j) is prominent in the herbaceous layer underneath trees where there is some soil present. the tall shrub portulacaria afra (species group d) is very dense and dominant locally. other species prominent locally include shrubs rhigozum obovatum and grewia occidentalis (species group j). the tree lycium oxicarpon (species group l), the dwarf shrub asparagus suaveolens and the grass tragus koelerioides (species group q) are also present. this community is inaccessible to most game species due to the large rocks with no soil in-between covering the area. thus no veld condition was calculated for this small community and it was not taken into account when determining the grazing capacity for the study area. 2.1.2 euclea undulata variant this variant is found on the higher western midslopes of the salpeterkop mountain in the southern section of the study area though slightly lower than the portulacaria afra variant (2.1.1) with the altitude varying between 1124–1140 m asl. this variant is only 14 ha in size and comprises large boulders to medium sized rocks covering approximately 70 % of the area. the trees are taller than 4 m and have an estimated 5 % coverage while the shrub layer (2.5–3 m tall) is the most prominent layer of this community with an 18–25 % cover. the herbaceous layer covers between 5–15 % of the area and is between 0.2–0.3 m tall. this variant is distinguished from the portulacaria afra variant (community 2.1.1) by the absence of species from species groups d and e. the vegetation is dominated by the shrubs carissa macrocarpa (species group b) and euclea undulata (species group c). the shrub rhus longispina is prominent locally. the tree acacia karroo (species group l) and the grass panicum maximum (species group k) are also present throughout this variant. the woody layer of this variant has previously been heavily utilised by angora goats with browsing line effects clearly visible. the veld condition and grazing capacity for this variant were calculated at 48 % and 41.4 ha/lsu, respectively. 2.2 rhus lucida-carissa macrocarpa shrubland this shrubland comprising 80 ha is found on the steep northern and north-western midslopes of the salpeterkop mountain with slopes ranging between 15º and 21º. altitude varies from 1115–1400 m asl with large rocks covering 50–60 % of the area. the woody layer is not very prominent with the trees (<4 m tall) covering only 2 % and the shrubs (1.5–2.5 m tall) covering between 15–20 % of the area. the herbaceous layer (0.5–0.7 m tall) is the most prominent layer covering between 50–60 % of the area. the presence of the shrub rhus lucida, the grasses themeda triandra, aristida junciformis, sporobolus fimbriatus and the forb pegolettia retrofracta (species group f) together with the shrub carissa macrocarpa, the grass bothriochloa insculpta and the forb galenia species (species group b) are characteristic for this community. the shrub rhus longispina (species group j) dominates the woody layer, while the grass cenchrus ciliaris (species group j) dominates the herbaceous layer. the shrubs rhus lucida (species group f), rhigozum obovatum (species group j) and the grasses themeda triandra and aristida junciformis (species group f) are also prominent. this community is in good condition with certain sections slightly degraded due to preissn 0075-6458 31 koedoe 48/2 (2005) koedoe 48/2 (2005) 32 issn 0075-6458 ta bl e 1 p hy to so ci ol og ic al ta bl e of th e ve ge ta tio n of th e in gl es id e an d w el ge da ch t s ec tio ns o f t he m ou nt ai n ze br a n at io na l p ar k issn 0075-6458 33 koedoe 48/2 (2005) ta bl e 1. (c on tin ue d) koedoe 48/2 (2005) 34 issn 0075-6458 ta bl e 1. (c on tin ue d) issn 0075-6458 35 koedoe 48/2 (2005) ta bl e 1. (c on tin ue d) vious grazing practices. owing to the abundance of the palatable grass cenchrus ciliaris this community achieved a high veld condition score of 93 % and a grazing capacity of 20 ha/lsu. the steep slopes on which it is located makes this community inaccessible to plains game species and it will therefore not be utilised frequently by game. if incorrectly managed this community could change into a dense thicket. 3. setaria neglecta-becium burchellianum grassland this grassland is found on the eastern midslope plateaus and lower midslopes of the salpeterkop mountain in the centre of the study area. the altitude varies between 1099–1144 m asl with large rocks covering 25–50 % of the area. the midslope plateaus are level though the lower midslopes are moderate to steep varying from 8º to 15º. this grassland is 148 ha in size. there is no tree layer. the shrub layer varies in height between 1.5–2.5 m with a 10 % cover. the herbaceous layer (0.4–0.6m tall) is the most prominent layer and covers between 30–60 % of the area. diagnostic species include the shrub diospyros lycioides, dwarf shrub rhus ciliata, the grass digitaria eriantha and the forb becium burchellianum (species group g). this grassland is dominated by the grasses setaria sphacelata and heteropogon contortus (species group i), and the forb becium burchelliatum (species group g). the shrubs rhus longispina, rhigozum obovatum (species group j) and eriocephalus ericoides (species group p) are prominent locally. other species also present include the dwarf shrubs asparagus striatus, felicia filifolia (species group i), pentzia globosa (species group q), the grass cymbopogon pospischilii (species group i) and the forbs helichrysum dregeanum and walafrida geniculata (species group p). this area has previously been mildly utilised by domestic animals and does not show signs of severe overgrazing. the veld condition score for this community is 55 % resulting in it having a grazing capacity of 15.3 ha/lsu. 4. rhigozum obovatum-enneapogon scoparius grassland the rhigozum obovatum-enneapogon scoparius grassland is the second largest community on the study area (383 ha). it is located on midslopes and footslopes and sometimes on mid-plateaus in the northern and western sections of the study area. the area has small to large rocks with a 30–60 % cover. altitude varies between 1053–1215 m above sea level. koedoe 48/2 (2005) 36 issn 0075-6458 table 2 veld condition and grazing capacity for ingleside and welgedacht farms, mountain zebra national park plant community 1.0 2.1.1 2.1.2 2.2 3.0 4.0 5.0 6.1 6.2 7.0 total size (ha) 16.0 42.0 14.0 80.0 148.0 383.0 129.0 272.0 415.0 55.0 1554.0 veld condition index (%) 0.0 0.0 48.4 93.1 55.4 35.7 28.5 22.2 32.5 19.7 grazing capacity (ha/lsu ) 0.0 0.0 41.2 20.0 15.3 28.1 25.6 68.3 34.1 64.8 number lsu game 0.0 0.0 0.3 4.0 9.7 13.6 5.0 4.0 12.2 0.8 49.7 total grazing capacity (ha/lsu) game 30.1 there are only a few trees (3.5 m tall) present covering approximately 3 % of the area. the shrub layer consists mainly of dwarf shrubs and a few larger shrubs ranging between 0.5 m and 2 m tall with a 15–30 % cover. the grasses are between 0.2–0.4 m tall with a 15–50 % cover. this community is characterised by the presence of species from species group h. these include the shrubs cadaba aphylla, ehretia rigida, the dwarf shrub eberlanzia ferox, chrysocoma ciliata, the grasses enneapogon scoparius, aristida diffusa, fingerhuthia africana, oropetium capense and the forbs tephrosia species, aptosimum decumbens, kleinia longiflorus, euphorbia decepta, pachypodium succulentum, polygala leptophylla, hibiscus trionum, crassula muscosa, solanum species, bulbine narcissifolia, nenax microphylla, pellaea calomelanos, sutera halimifolia, plexipus pumilus and sansevieria species. the vegetation is dominated by the shrubs rhigozum obovatum (species group j) and gymnosporia buxifolia (species group l) together with the grasses enneapogon scoparius (species group h) and heteropogon contortus (species group i). the following species are locally prominent: the tree acacia karroo (species group l), the shrub rhus longispina (species group j), the grasses aristida diffusa (species group h), cymbopogon pospischilii (species group i), cenchrus ciliaris (species group j), tragus koelerioides (species group q) and the forbs pachypodium succulentum (species group h) and blepharis integrifolia (species group p). this community is accessible to game and domestic animals. as a result it has been heavily overgrazed due to previous farming practices and has a veld condition index of only 36 % resulting in a grazing capacity of 28.1 ha/lsu. 5. panicum maximum-acacia karroo woodland the panicum maximum-acacia karroo woodland is situated in the valley bottomland drainage channels in the south-east and western sections of the study area comprising 129 ha. altitude varies between 1022– 1077 m asl. rock cover is low varying between 1–3 % though in one section it has been estimated at 20 %. the trees are taller than 5 m and cover between 10–25 % of the area, while the shrub layer (2 m tall) covers between 15–35 % of the area. the herbaceous layer with an average height of 0.7 m has a 30–60 % cover. the vegetation is characterised by the presence of the diagnostic species of species group k and include the grasses panicum maximum and the forbs felicia muricata, sutera virgulosa, lepidium bonariensis and atriplex semibaccata. the woody layer is dominated by the tree acacia karroo (species group l), while the herbaceous layer is dominated by the grass panicum maximum (species group k). the shrub lycium oxicarpon (species group l), the dwarf shrub psilocaulon absimile (species group s) and the grasses eragrostis lehmaniana (species group o) and cynodon incompletus (species group s) are locally prominent. this community consists of various drainage channels flowing into each other. the vegetation of this community is disturbed due to previous overgrazing and trampling resulting in various erosion gulleys in and along the drainage channels. this disturbance is evident in the poor condition of the veld (28.5 %) and low grazing capacity of 25.6 ha/lsu. animals prefer this community throughout the year because of the presence of the palatable tree acacia karroo and the palatable grass panicum maximum. this normally leads to this community being heavily utilised by animals. 6. eragrostis obtusa-pentzia globosa shrubland this community is found on the lower footand midslopes of the salpeterkop mountain in the eastern and northern section of the study area. altitude varies between 1017– issn 0075-6458 37 koedoe 48/2 (2005) 1099 m asl with small rocks and gravely soil. rock cover is low and varies between 0–30 %. this major community with a total size of 687 ha comprises 44 % of the study area. the vegetation consists mainly of grasses and dwarf shrubs with no tree layer present. the dwarf shrubs have an average height of 0.3 m and covers between 25–30 % of the area while the grasses are between 0.1–0.3 m tall with a 25–60 % coverage. this shrubland is characterised by species group m and include eragrostis obtusa, aristida congesta subsp. barbicollis, aristida adscensionis and the forbs lycium cinerium, talinum caffrum and thesium lineatum. the vegetation is dominated by the dwarf shrub pentzia globosa (species group q) while the grasses eragrostis obtusa and aristida congesta subsp. barbicollis (species group m) are also prominent. 6.1 tragus koelerioides-pentzia globosa shrubland this largest section of this sub-community is located in the eastern section and the north western sections of the study area on the lower midslopes of the salpeterkop mountain. altitude varies between 1063–1099 m above sea level. rock cover varies between 25–60 % with shallow soil. this is the third largest community of the study area and is 272 ha in size. this community is characterised by the absence of species from species group n. the vegetation has been heavily grazed resulting in an average height of 0.2 m for the dwarf shrubs covering between 15–20 % of the area while the grasses are between 0.1–0.2 m tall with a 5–10 % coverage. the vegetation is dominated by the dwarf shrub pentzia globosa and the grasses tragus koelerioides (species group q) and aristida congesta subsp. congesta (species group m). the grass enneapogon scoparius (species group h) are locally prominent while the dwarf shrub asparagus suaveolens (species group q) are present throughout the community. this shrubland is heavily overgrazed and degraded (the north western sections particularly) with a low species diversity mainly comprising pioneer species. this severe overgrazing of this sub-community is reflected in the very low veld condition of 22.2 % and grazing capacity of 68.3 ha/lsu. 6.2 cynodon incompletus-pentzia globosa shrubland the cynodon incompletus-pentzia globosa shrubland is the largest single community of the study area comprising 415 ha. this shrubland is located in the eastern footslopes and midslopes of the salpeterkop mountain on deeper and more fertile soil. altitude varies between 1017–1083 m asl while rock cover is low and varies between 0–10 %. this community is characterised by the presence of the diagnostic species of species group n and include the dwarf shrubs zygophyllum incrustatum, plinthus karooicus, the grasses panicum coloratum, sporobolus nitens, the forbs opuntia aurantiaca, galenia sarcophylla, berkeya species commelina africana and the geophyte ammocharis coranica. the vegetation is dominated by the dwarf shrub pentzia globosa (species group q) and the pioneer grass cynodon incompletus (species group s), whereas the grasses eragrostis obtusa, aristida congesta subsp. congesta (species group m), sporobolus nitens (species group n) and eragrostis lehmaniana (species group o) are also prominent. the declared invader opuntia aurantiaca (species group n) is present in large numbers in sections within this community and should be eradicated. a fairly large number of termite mounds characterise this shrubland. this sub-community has also been prone to overgrazing and has a veld condition score of 32.5 % and a grazing capacity of 34.1 ha/lsu. koedoe 48/2 (2005) 38 issn 0075-6458 7. chloris virgata-cynodon incompletus grassland this community is situated on old cultivated fields in the lower lying areas in the western section of the study area and is only 55 ha in size. the area is level and the altitude was measured at 1023 m above sea level. there are only a few small rocks present covering less than 1 % of the area. there is no woody layer present and the herbaceous layer is 0.2 m tall and covers between 40–80 % of the area. the grass chloris virgata and the forbs altenanthera species, amaranthus praetermissus, conyza bonariensis, schkuhria pinnata and hibiscus pusillus (species group r) are diagnostic for this community. the vegetation comprises a mixture of various pioneer species dominating sections locally. these include the grasses cynodon incompletus (species group s), chloris virgata (species group r) and the forb salsola kali (species group s) that are present throughout the community. the forb psilocaulon absimile (species group s) are dominant locally. this community comprises old fields of various ages in different stages of secondary succession. in its natural condition this community formed part of the panicum maximum-acacia karroo woodland (community 5) but has been cleared for cultivation purposes. once cultivation of these old fields has been discontinued the lands have been left fallow resulting in the increase of pioneer species. since many of these pioneer grass species (chloris virgata, cynodon incompletus, urochloa panicoides) are palatable it has resulted in the area being overgrazed by angora goats and sheep previously stocked on the farm. this is evident in the very low veld condition of 19.7 % and grazing capacity of 64.8 ha/lsu. in order for this community to return to its natural condition it is recommended that low stocking rates are applied to this community or that the overgrazed sections are restored to a more acceptable condition. discussion plant communities 1 and 2 are located at high altitudes on the escarpment with very steep (25º–28º) upper slopes and high rock cover ranging between 50–90 %. plant communities 3 and 4 are located at medium altitudes with rock cover ranging between 25–60 %. plant community 3 is restricted to midslope plateaus, while plant community 4 is mainly located on the lower midslopes and footslopes. the lower footslopes and midslopes are characterised by the eragrostis obtusa-pentzia globosa shrubland (community 6) at medium to low altitudes. compared to the other communities described within the study area, this major community has a relatively low rock cover varying between 0–30 %. the drainage channels are represented by plant communities 5 and 6. these low-lying communities are situated on more fertile soil with very low rock cover varying between 1–3 %. this study resulted in the identification of 195 different species representing 112 genera and 38 families being identified in ingleside-welgedacht section of the mountain zebra national park. this represents 28 % of all the species and 44 % of all the families previously identified within the park (pond et al. 2002). the poaceae has the largest number of species (31) reflecting 16 % of the species of the study area followed by the asteraceae with 15 species (8 %). these are followed by two much smaller though significant families, the mesembryanthemaceae and fabaceae with seven (4 %) and five (3 %) of the species total for the study area. the largest section of the study area has been overgrazed due to previous farming practices. this has resulted in an overall grazing capacity of 30.1 ha/lsu for the study area which is high compared to the agricultural norm of 14 ha/lsu for the area. the only communities not affected as much by the previous grazing practices are those situated on the higher lying more inaccessible areas (communities 1, 2 & 3). in order to improve the veld condition and grazing capacity of the study area it is recommended that the issn 0075-6458 39 koedoe 48/2 (2005) area is stocked at well below the carrying capacity for the next few years to ensure stabilisation of the soil and recovery of the vegetation. restoration of the drainage channels (community 5) and the old fields (community 7) could also be considered. this could be achieved by re-seeding these areas with natural grasses occurring in the area. field trials at the doornhoek farm of mznp yielded good results using a one tooth implement to rip the soil approximately 100 mm deep and sowing the seeds in the ripped line (pers. obs.). it is also important that no water points are placed in these communities since that would attract more game causing further trampling and overgrazing. conclusion the braun-blanquet approach proved to be an accurate and effective way whereby floristically defined plant communities could be classified and identified in the field. a total of 10 different clearly recognisable plant communities that can be grouped into seven major communities were identified and described for the study area. no similar vegetation descriptions have previously been completed on the vegetation of the study area and this research therefore provides valuable data on the ecosystems present. an understanding of the plant communities and their associated habitats are of fundamental importance for compiling sound management and conservation strategies. these vegetation surveys and descriptions provide baseline information that allows monitoring as well as similar surveys to be conducted in future. the expansion of mznp does not only contribute floristically to the diversity of the park but also faunally due to the inclusion of new habitats into the park which make it suitable for the introduction of larger herbivores that occurred historically in this region. the data obtained from this study will therefore be invaluable when compiling a new management plan for the mountain zebra national park as part of a larger project undertaken by unisa and south african national parks. acknowledgements the following individuals and institutions are sincerely thanked: national research foundation (nrf) and unisa for financing this study; sanparks, especially management of mountain zebra national park, for allowing and encouraging the research; mr johan de klerk for his dedication and support of this project; dr stephen holness for the assistance with the maps; mr ernest daemane (kimberley south african national parks herbarium–ksan) for verifying the plant species names; and gn dithlale (unisa) for assistance with the species data analysis. references acocks, j.p.h. 1988. veld types of south africa. 3rd ed. memoirs of the botanical survey of south africa 57: 1–146. bezuidenhout, h. 1993. syntaxonomy and synecology of western transvaal grasslands. ph.d dissertation, university of pretoria, pretoria. bezuidenhout, h. 1996. the major vegetation communities of the augrabies falls national park, northern cape. 1. the southern section. koedoe 39: 7–24. bezuidenhout, h., h.c. biggs, & g.j. bredenkamp. 1996. a process supported by the utility bbpc for analysing braun-blanquet data on a personal computer. koedoe 39: 107–112. bredenkamp, g.j. 1982. 'n plantekologiese studie van die manyeleti-wildtuin. d.sc. thesis. university of pretoria, pretoria. bredenkamp, g.j. & h. bezuidenhout. 1995. a proposed procedure for the analysis of large data sets in the classification of south african grasslands. koedoe 38(1): 33–39. bredenkamp, g.j., a.f. joubert & h. bezuidenhout. 1989. a reconnaissance survey of the vegetation of the plains in the potchefstroomfochville-parys area. south african journal of botany 55: 199–206. brown, l.r. & h. bezuidenhout. 2000. the phytosociology of the de rust section of the mountain zebra national park, eastern cape. koedoe 43(1): 1–18. brown, l.r. & g.j bredenkamp. 1994. the phytosociology of the southern section of the borakalalo nature reserve, south africa. koedoe 37: 59–72. brown, l.r. 1997. a plant ecological and wildlife management plan of the borakalalo nature reserve, north-west province. ph.d. dissertation. university of pretoria, pretoria. carruthers, j. 2003. changing philosophies & values. pp. 23–42. in: south african national koedoe 48/2 (2005) 40 issn 0075-6458 parks: a celebration. hall-martin, a. & j. carruthers (eds.). singapore: tien wah press. cilliers, s.s. 1998. phytosociological studies of urban open spaces in potchefstroom, north west province, south africa. ph.d thesis, potchefstroom university for che. coetzee, b.j. 1983. phytosociology, vegetation structure and landscapes of the central district, kruger national park. dissertationes botanicae 69: 1–456. danckwerts, j.e. 1989. monitoring vegetation and assessment of veld condition in grassveld. pp. 96–99. in: danckwerts, j.e. & w.r. teague (eds.). veld management in the eastern cape. pretoria: government printer. de klerk, j., brown, l.r., & h.bezuidenhout. 2003. plant communities of the ebenaeser section of the mountain zebra national park. koedoe 46(2): 1–13. eckhardt, h.c. 1993. a synecological study of the vegetation of the north-eastern orange free state. m.sc. thesis. university of pretoria, pretoria. edwards, d. 1983. a broad-scale structural classification of vegetation for practical purposes. bothalia 14: 705–712. foran, b.d., n.m.tainton & p. de v. booysen. 1978. the development of a method assessing veld conditions in three grassveld types in natal. proceedings of the grassland society of southern africa 13: 27–33. fuls, e.r. 1993. vegetation ecology of the northern orange free state. ph.d. dissertation. university of pretoria, pretoria. fuls, e.r., g.j. bredenkamp, & n van rooyen. 1992. the plant communities of the undulating grassland of the vredefort–kroonstad–lindley–heilbron area, northern orange free state. south african journal of botany 58: 224–230. germishuizen, g. & n.l. meyer (eds). 2003. plants of southern africa: an annotated checklist. pretoria: national botanical institute. (strelitzia 14). hall-martin, a. 2003. south africa’s biodiversity. pp. 1–19. in: south african national parks: a celebration. hall-martin, a. & j. carruthers (eds.). singapore: tien wah press. hill, m.o. 1979. twinspan–a fortran program for arranging multivariate data in an ordered twoway table by classification of the individuals and attributes. new york: cornell university. hoffman,t. 1996. eastern mixed nama karoo. p. 55. in: low, a.b. & a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. kooij, m.s., g.j. bredenkamp & g.k. theron. 1990. classification of the vegetation of the b land type in the north-western orange free state. south african journal of botany 56: 309–318. land type survey staff. 1986. land types of the maps se27/20 witdraai, 2720 noenieput, 2722 kuruman, 2724 christiana, 2820 upington, 2822 postmasburg. memoirs on the agricultural natural resources of southern africa 3: 1–185. land type survey staff. 1999. land types occurrence (maps) and areas of 3224 graaff-reinet. unpublished information obtained from institute for soil, climate and water, pretoria. lubke, r., g.j. bredenkamp & n. van rooyen. 1996. south-eastern mountain grassland. p. 55. in: low, a.b. & a.g. rebelo (eds.). vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs & tourism. mueller-dombois, d. & h. ellenberg. 1974. aims and methods of vegetation ecology. new york, ny: wiley. mucina, l. & e. van der maarel. 1989. twenty years of numerical syntaxonomy. vegetatio 81: 1–15. novellie, p. 1990a. habitat use by indigenous grazing ungulates in relation to sward structure and veld condition. south african journal of wildlife research 7: 16–23. novellie , p. 1990b. the impact of a controlled burn on karroid merxmuellera mountain veld in the mountain zebra national park. south african journal of ecology 1: 33–37. novellie, p.a. & h. bezuidenhout. 1994. the influence of rainfall and grazing on vegetation changes in the mountain zebra national park. south african journal of wildlife research 24: 60–71. pond, u., b.b. beesley, l.r. brown & h. bezuidenhout. 2002. floristic analysis of the mountain zebra national park, eastern cape. koedoe 45(1): 35–57. orban, b. 1995. an ecological management plan for the lionspruit game reserve. m.sc. (wildlife management) dissertation. university of pretoria, pretoria. smit, g.n. 1988. die invloed van langtermyn graslaag benuttingspraktyke op die suurgemengde bosveld. m.sc. dissertation. university of pretoria, pretoria. soil classification working group. 1991. soil classification: a taxonomic system for south africa. memoirs on the agricultural natural resources of south africa 15: 1–262. toerien, d.k. 1972. geologie van die bergkwagga nasionale park. koedoe 15: 77–82. trollope, w.s.w., l. trollope & o.j.h. bosch. 1990. veld and pasture management terminology in southern africa. journal of the grassland society of southern africa 7: 52–61. issn 0075-6458 41 koedoe 48/2 (2005) tueller, p.t. 1988. vegetation science applications for rangeland analysis and management. dordrecht: kluwer academic publishers. van der walt, p.t. 1980. a phytosociological reconnaissance of the mountain zebra national park. koedoe 23: 1–32. van oudtshoorn, f.p. 1992. guide to grasses of south africa. arcadia, pretoria: briza. van rooyen, n., g.k. theron & n. grobbelaar. 1981. a floristic description and structural analysis of the plant communities of the punda milia-pafuri-wambiya area in the kruger national park, republic of south africa. 1. the hygrophilous communities. south african journal of botany 47: 213–246. vorster, m. 1982. the development of the ecological index method for assessing veld condition in the karoo. proceedings of the grassland society of southern africa 17: 84–89. westhoff, v. & e. van der maarel. 1978. the braun-blanquet approach. pp. 287–399. in: whittaker, r.h. (ed.). classification of plant communities. the hague: junk. whittaker, r.h. 1978. dominance types. pp. 65–79. in: whittaker, r.h. (ed.). classification of plant communities. the hague: junk. koedoe 48/2 (2005) 42 issn 0075-6458 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 page 3 stalmans banhine.qxd plant communities, wetlands and landscapes of the parque nacional de banhine, moçambique m. stalmans and m. wishart stalmans, m. and m. wishart. 2005. plant communities, wetlands and landscapes of the parque nacional de banhine, moçambique. koedoe 48(2): 43–58. pretoria. issn 00756458. the parque nacional de banhine (banhine national park) was proclaimed during 1972. it covers 600 000 ha in moçambique to the east of the limpopo river. until recently, this park, originally and popularly known as the ‘serengeti of moçambique’, was characterised by neglect and illegal hunting that caused the demise of most of its large wildlife. new initiatives aimed at rehabilitating the park have been launched within the scope of the greater limpopo transfrontier park. a vegetation map was required as input to its management plan. the major objectives of the study were firstly to understand the environmental determinants of the vegetation, secondly to identify and describe individual plant communities in terms of species composition and structure and thirdly to delineate landscapes in terms of their plant community and wetland make-up, environmental determinants and distribution. a combination of fieldwork and analysis of landsat satellite imagery was used. a total of 115 sample plots were surveyed. another 222 sample points were briefly assessed from the air to establish the extent of the different landscapes. the ordination results clearly indicate the overriding importance of moisture availability in determining vegetation composition in the parque nacional de banhine. eleven distinct plant communities were recognised. they are described in terms of their structure, composition and distribution. these plant communities have strong affinities to a number of communities found in the limpopo national park to the west. the sandveld community is relatively the most species-rich of all communities. different combinations of these plant communities can be grouped in five major landscapes, namely the wetland, grassland, mopane, sandveld and androstachys landscape. these different landscapes hold six different wetland types as defined by the ramsar classification. the landscapes with their individual plant communities and wetland types represent a unique combination of habitats that have great intrinsic conservation value. these habitats are key to the long-term maintenance and survival of a diverse avifauna, including the original ostrich population, as well as wattled cranes. key words: parque nacional de banhine, greater limpopo transfrontier park, tfca, landscape, wetland. m. stalmans, international conservation services, p.o. box 19139, nelspruit, 1200 republic of south africa (stalmans@ics-consulting.co.za); m. wishart, freshwater research unit, department of zoology university of cape town, rhodes gift, 7701 republic of south africa. issn 0075-6458 43 koedoe 48/2 (2005) introduction the parque nacional de banhine (pnb) was proclaimed during 1972. it covers 600 000 ha in moçambique to the east of the limpopo river and forms one of the major components of the greater limpopo transfrontier park (fig. 1). a vegetation map was required as one of the essential building blocks for the drafting of its management plan (anonymous 2002). little or no botanical surveys were carried out in moçambique between 1980 and 1994 during the long period of internal conflict that affected much of the country (anonymous 1997). the extent of the area, its relative inaccessibility due to poor road infrastructure, and the limited time available prekoedoe 48/2 (2005) 44 issn 0075-6458 fig. 1. locality map of the parque nacional de banhine. cluded a traditional fine-scale vegetation description and map of the pnb. such a finescale input is in any case likely to be too detailed for the scale of management envisaged, certainly in the short to medium term. vegetation is often used as a surrogate or building block for the definition of habitats (timberlake et al. 1993). the use of broad habitat units defined by a combination of environmental factors and vegetation would probably represent the most useful input for the drafting of the management plan. this is because such units have relevance to different species of wildlife, they may differ in their ecological capacity and availability of water, they might require different fire regimes, and they will differ in their sensitivity to utilisation and development. as both the limpopo national park and the pnb form major components of the greater limpopo transfrontier park, it was decided to build on the work recently completed in the limpopo national park (stalmans et al. 2004). a similar conceptual approach was followed whereby landscapes were mapped explicitly whereas the embedded plant communities are implicit. a landscape is defined as ‘an area with a specific geomorphology, climate, soil vegetation pattern and associated fauna’ (gertenbach 1983). the objectives of this study were firstly to understand the environmental determinants of the vegetation, secondly to identify and describe individual plant communities in terms of species composition and structure, and thirdly to identify and delineate landscapes in terms of their plant community and wetland make-up, environmental determinants and occurrence. this is the second in a series of three articles on the vegetation of the limpopo, banhine and zinave national parks that are located in the moçambican component of the greater limpopo transfrontier park. difficult access, limited resources and a lack of previously published information have severely limited the sampling intensity, the extent of the descriptions and the detail of the mapping. nevertheless, it is hoped that these first published accounts of the vegetation and landscapes of these parks will provide a basis on which further research projects can expand, much as the level of knowledge of the kruger national park has been improved upon over the years. study area the physical environment largely determines the vegetation composition and structure. the parque nacional de banhine is situated between latitudes 22º30'–23º20's and longitudes 32º15'–33º25'e in the gaza province of moçambique (fig. 1). total area is ca. 600 000 ha. the pnb forms part of the greater limpopo transfrontier park with the limpopo national park to the west and the zinave national park to the north-east. according to the köppen classification the area has a warm arid climate with a dry winter and a mean annual temperature exceeding 18 ºc (van rooyen et al. 1981). a mean annual rainfall value of 399 mm applies to the western half of the pnb and 427 mm to the eastern half using the grid at 0.5 degree longitude/latitude resolution of the leemans & cramer (1991) database. considerable variations can be expected within and between seasons (kelly & walker 1976). the highest elevation is on the north-western boundary at 181 m above sea level. the area dips generally in a south-easterly direction with the lowest point being at 69 m asl on the eastern boundary. the area is underlain by arenaceous and argillaceous sediments with colluvium in the east (anonymous 1995). the pnb is covered with a pale grey sandy soil mantle that overlies a layer of calcrete about seven metres below the surface. very occasionally, there are broken lines of calcrete that reach the surface and extend for distances of up to a hundred metres. the pnb represents a converging endorheic delta-type system with limited overflow (tinley pers. comm. 2002). a number of ephemeral streams converge from the northwest into a delta-type system forming an endorheic basin with thin alluvial deposits. issn 0075-6458 45 koedoe 48/2 (2005) fig. 2. schematic representation of the endorheic wetland system of the parque nacional de banhine. an overflow is present on the eastern boundary, draining into the changane river (fig. 2). the pnb falls within the mopane vegetation of the sudano-zambezian region as described by werger & coetzee (1978). mopaneveld has been described for the hot, dry valley bottom of the limpopo river in moçambique (wild & barbosa 1967). the area was formerly often popularly referred to as the ‘serengeti of moçambique’ because of the large numbers of zebra, wildebeest and eland that occupied its wide-open grasslands. most of this diverse and numerous large herbivore component has been lost over the last decades, mostly through indiscriminate and illegal hunting. there are approximately 2000–3000 people living in and adjacent to the pnb in 11 villages. these people are highly dependent on the direct consumption of park resources for day-to-day survival, including basic household needs of shelter, water, food and medicines, crop and livestock production. methods sampling a total of 115 plots of 40 x 40 m were subjectively located in representative stands of vegetation to cover the variation in elevation, soils and terrain position. the limited road network was used during the ground survey (fig. 3). edwards’ (1983) structural classes were used to describe the overall structural properties of the sampled plots. overall cover was estimated for the woody, grass, forb and geophyte components, respectively, using the semi-quantitative measures of the braun blanquet approach (mueller-dombois & ellenberg 1974). cover and height classes were recorded for individual woody and grass species. records of environmental data included gps position, geology, landscape position (land type survey staff 1989), slope steepness, soil texture (using the sausage method (national working group for vegetation ecology 1986)) and rockiness. fieldwork was undertaken from may to june 2002. access to the study area can be very difficult due to flooding during summer. vegetation sampling subsequently had to be postponed till the end of the summer season into winter although this is not ideal in terms of plant species identification. in order to further increase sampling intensity, so-called ‘pseudo-plots’ were also used. these consisted of a gps point and subjective visual assessment relative to the formally surveyed sites. a total of 222 such pseudoplots were assessed during several fixedwing and microlight flights (fig. 3). analysis data were analysed through a combination of classification and ordination techniques. classification is used to identify groups and to impose structure to raw data. ordination aims at arranging species and samples in a low-dimensional space such that similar entities are close by and dissimilar entities far apart. a twinspan classification (hill 1979) was performed on the sample data. two-way indicator species analysis (twinspan) is a polythetic divisive technique based on reciprocal averaging ordination (gauch 1982). it is one of the preferred hierarchical techniques because of its effectiveness and robustness. it results in the definition of communities, each characterised by its own distinctive species combination. although more contemporary and sophisticated software packages are available for vegetation classification, twinspan’s simplicity and koedoe 48/2 (2005) 46 issn 0075-6458 fig. 3. position of vegetation sample plots and control points in the parque nacional de banhine. robustness were appealing within the constraints of this particular study. the canoco package (ter braak 1992) was selected to analyse relationships between the data set of 115 plots by 168 species and the underlying environmental factors. canoco allows for canonical ordination. this is an intermediate technique that combines aspects of regular ordination with aspects of regression (jongman et al. 1987). cca (canonical correspondence analysis) was used. the resulting ordination diagram expresses not only the pattern of variation in species composition but also the main features of species distributions along the gradient of environmental variables (ter braak 1986). delineation of landscapes the relevant landscapes for the pnb were identified using the knowledge gained through the ordination and classification of the pnb field data. a combined approach was used to map these landscapes using the available soils map, the existing land cover map and the landsat image. the land cover map (“carta de uso e cobertura da terra”) has been derived for moçambique from landsat satellite imagery at a scale of 1:250,000 (anonymous 1999). a ‘best fit’ map of the landscapes was subjectively drawn by matching the individual land cover polygons to the field sampling data and the 222 pseudoplots. furthermore, polygon boundaries were manually edited following visual inspection of the satellite image. the coverage of the polygons by the available data set was assessed in order to derive a measure of certainty or uncertainty regarding the quality of the final landscape map. results and discussion causal factors of vegetation pattern in the pnb the first ordination run with the full set of 115 vegetation plots resulted in two distinct ‘clouds’ of sample plots in ordination space (fig. 4). the environmental variables issn 0075-6458 47 koedoe 48/2 (2005) fig. 4. canoco ordination of 115 vegetation sample plots in the parque nacional de banhine. note dense cloud of sample plots on left side of diagram with outlying plots to the right that represent vegetation in areas with high moisture availability. ‘floodplain’ had a high canonical coefficient of 0.74 with the first axis and ‘drainage line’ had a canonical coefficient of 0.56. the 37 plots on the right of the diagram represent all the pans, riverbanks and reed beds that were sampled (fig. 4). the amount of available moisture as determined through landscape position therefore represents a major environmental determinant of vegetation composition along the first ordination axis. the different sample scores of a specific environmental variable are represented within the ordination diagram by a single point, called the ‘centroid’. the grasses cynodon dactylon and eragrostis cf. heteromera are typical of these floodplain conditions. the woody species mimusops obtusifolia, grewia sulcata, spirostachys africana and acacia robusta are typical for the fringes of the major drainage lines feeding the banhine system from the north. the 37 plots associated with the floodplain and drainage line were removed from the data set and a second ordination was run. the resulting diagram identifies an outlier in the mopane veld on the northern boundary with the grass layer being dominated by cymbopogon. the other outliers represent the bush clumps found in the floodplain that are dominated by xanthocercis zambesiaca in association with salvadora persica, allophylus africanus, ficus capreifolia, maclura africana and rhus guenzii. these bush clumps on termite mounds have a high clay content. the parameter ‘soil texture’ has a canonical coefficient of 0.89 and t-value of 10.3 for the first axis. this clearly indicates the increasing clay content of the soils. once again, the five outlying plots were removed and the remaining data set was ordinated. the split between ‘sandveld’ and non-sandveld sample plots is very clear along the gradient of decreasing clay content of the soil along the first ordination axis (fig. 5). the parameter ‘soil texture’ has a canonical coefficient of 0.76 and t-value of 9.6 for the first axis. there is a weaker gradient in terms of landscape position with crest and upper slope positions negatively correlated to the second axis (canonical coefficients of 0.66 and 0.46 respectively). the non-sandveld sample plots are characterised by mopane colophospermum mopane. sandveld has typically no mopane (or only a low cover percentage for this species) with typical sandveld species such as xerroderris stuhlmannii, strychnos madagascariensis and guibourtia conjugata. koedoe 48/2 (2005) 48 issn 0075-6458 fig. 5. canoco ordination of 73 vegetation sample plots in the parque nacional de banhine (excluding sample plots along major drainage lines, on floodplains and on termite mounds). the successive ordination runs clearly indicate the overriding importance of moisture availability (in conjunction with soil texture) in determining vegetation composition in the pnb. firstly, the obvious differences in moisture availability by virtue of sample positions along drainage lines and in the floodplain came out of the first ordination. thereafter, the gradient in soil clay content (as a result of underlying geological substrate and landscape position) and landscape position per se (in determining water flow) largely determine soil moisture and nutrient availability. the interplay of soil moisture and soil nutrient availability conforms to the current understanding of the determinants of savanna. the four-determinant model gives water availability and nutrient availability equal status in establishing the range of possible forms a issn 0075-6458 49 koedoe 48/2 (2005) savanna can assume (scholes & walker 1993). fire and herbivory then determine the actual form and function within that range. similarly, timberlake et al. (1993) consider soil moisture in this environment as a major determinant in the distribution of vegetation types. soil moisture availability results from the interaction between rainfall, topography, soil texture, soil depth, drainage and rooting habit. siebert et al. (2003) identified a gradient of decreasing soil moisture availability along the first axis of ordination of more than 2000 sample plots in mopane veld straddling south africa, namibia and botswana. stalmans (1994) and stalmans et al. (2004) identified water availability (as controlled by the position of the sample in the landscape and by its soil texture) as the major determinant of vegetation composition in an area adjacent to the gonarezhou national park in zimbabwe and in the limpopo national park respectively. twinspan dendrogram the results from the twinspan classification are presented by means of a hierarchical dendrogram (fig. 6). starting from the top of the dendrogram, the set of 115 sample plots is divided into two groups. the left group represents the bulk of the sample plots (n=83). the right group with indicator species cynodon dactylon, eragrostis heteromera, panicum coloratum and paspalidium obtusifolium represent vegetation associated with seasonally or permanently flooded conditions. these two main divisions are now described separately. fig. 6. twinspan dendrogram for 115 vegetation sample plots in the parque nacional de banhine. the androstachys woodlands and thickets already become a separate entity at level 2 of the diagram (community 1). the xanthocercis zambesiaca bushclumps on termitaria are split off from the remaining plots at level 3 (community 2). at level 4, the division is according to the clay content in the soil profile. the vegetation on more clayey soils is divided into two communities, namely closed woodlands along seasonal drainage lines (community 3) and typical mopane woodlands (community 4). the split at level 5 for the more sandy substrates produces a group of plots located on welldrained soils and community 7 that is found on sandy soils with a high water table. finally at level 6, the remaining plots on sandy soils are split into the sandy variant of the mopane community (community 5) and the true sandveld of community 6. within the wetland and floodplain vegetation, the split at level 2 is into units that are wooded or not wooded. at level 3, in the wooded units, the acacia borleae shrubland that is found encroaching from the mopane into the grasslands becomes distinct (community 8). community 9 represents wooded grasslands. within the non-wooded unit, the grasslands of the seasonally flooded flats (community 10) are separated from the pans and true wetland. the latter two communities are differentiated by cynodon dactylon on seasonally flooded grasslands and paspalidium obtusifolium as a typical (but not exclusive) exponent of the pans. definition of plant communities the community concept is applied in its broad sense and reflects a recurring assemblage of grass and woody species of characteristic composition and structure, growing in an area of essentially similar environmental conditions and land use history (adapted from gabriel & talbot (1984)). the classification outcome was evaluated subjectively against photographs of each sample plot. the main criterion applied was the need for each community to be identifiable in the field by an observer who is not necessarily a trained botanist. community names were chosen subjectively so as to have practical value in the field through the use of two species which are visually and/or diagnostically important. the communities broadly conform to the lower divisions of the dendrogram and they are therefore discussed from left to right following fig. 6. a total of 11 communities were identified. description of plant communities 1 androstachys johnsonii-croton pseudopulchellus closed woodlands this is a species-poor community found on red, deep sandy soils. it consists of an extremely dense, short (5–10 m) woodland dominated by androstachys johnsonii. other species have a very low percentage cover value and biomass. the woody species croton pseudopulchellus, guibourtia conjugata, xeroderris stuhlmannii and manilkara mochisia are also found in this community. the grass cover is very low with the perennial panicum maximum and the annual brachiaria deflexa being the most important species. this community is mostly found on the north-western boundary as well as in the southern corner of the park. it is generally sharply demarcated from other communities and occurs in patches ranging from a few dozen metres in diameter to large areas covering many hectares. this community corresponds to the androstachys johnsonii-croton pseudopulchellus dry forest described by van rooyen et al. (1981) in the northern part of the kruger national park and to the androstachys johnsonii-guibourtia conjugata short forest described by stalmans et al. (2004) for the limpopo national park. 2 xanthocercis zambesiaca salvadora persica thickets this community occurs as distinct bush clumps with a diameter of 20–30 m on termitaria, mostly on the transition from the grasslands to the wetlands. each clump is characterized by an emergent xanthocercis koedoe 48/2 (2005) 50 issn 0075-6458 zambesiaca (nyala tree) that is surrounded by a variety of other species forming a fringe around the core of the clump. these other species comprise allophylus africanus, grewia sulcata, rhus guenzii, ficus capreifolia, maclura africana and salvadora persica. the grass layer is poorly developed and consists mostly of panicum maximum. 3 mimusops obtusifolia-spirostachys africana closed woodlands this community is found along seasonal rivers, mostly on the northern boundary of the park. this riverine community occupies a narrow strip of 10–20 m wide along these drainage lines. the most important species apart from the dominant mimusops obtusifolia and spirostachys africana are grewia sulcata, acacia robusta, combretum imberbe, garcinia livingstonei and ziziphus mucronata. the most important grass species is panicum maximum. 4 colophospermum mopane urochloa mosambicensis woodlands this represents the most extensive ‘mopane’ vegetation found in the pnb . it occurs on slightly more clayey soils (sandy loam) than the mopane of community 5. prominent woody species are combretum apiculatum, sclerocarya birrea, dichrostachys cinerea, dalbergia melanoxylon and drypetes mossambicensis. a major difference with the community 5 is the marked absence of terminalia sericea and xeroderris stuhlmannii, as well as the lower frequency of occurrence and lower cover of combretum apiculatum. the most important grasses are panicum maximum, digitaria eriantha, schmidtia pappaphoroides, aristida adscencionis and heteropogon contortus. 5 colophospermum mopane eragrostis pallens woodlands this is still a mopane-dominated community but represents a more ‘sandy’ variant. in addition to mopane, the woody species combretum apiculatum, terminalia sericea, xeroderris stuhlmannii, strychnos madagascariensis, guibourtia conjugata, boscia albitrunca and sclerocarya birrea are important components. the grass component also reflects the more sandy nature of the soils with panicum maximum, digitaria eriantha, perotis patens, eragrostis pallens and aristida stipitata. heteropogon contortus is almost totally absent. 6 guibourtia conjugata eragrostis pallens woodlands this community is found on deep sandy soils. characteristic woody species of the ‘sandveld’ are hugonia orientalis, pteleopsis myrtifolia, cleistanthus schlechteri, hymenocardia ulmoides and xylotheca kraussiana. other important species are the more wide-spread combretum apiculatum, terminalia sericea, strychnos madagascariensis and s. spinosa. the grass layer is represented by eragrostis pallens, panicum maximum, tricholaena monachme, perotis patens and aristida spp. this particular community very closely resembles the terminalia sericea-pogonarthria squarrosa tree savanna identified by van rooyen et al. (1981) in the punda milia area of the kruger national park. it also encompasses the xeroderris stuhlmannii combretum apiculatum tree savanna of van rooyen et al. (1981). these communities in turn are similar to the terminalia sericea eragrostis pallens low woodland described for the limpopo national park by stalmans et al. (2004). 7 hyphaene petersiana eragrostis gummiflua bushlands this community consists of a generally fairly open, but sometimes closed vegetation with a mix of trees and shrubs on the transition from the sandveld to the wetland. in addition to the hyphaene palms, the trees terminalia sericea and garcinia livingstonei are prominent. the grass layer is dominated by eragrostis gummiflua, e. pallens, perotis patens and digitaria eriantha. the palms issn 0075-6458 51 koedoe 48/2 (2005) and eragrostis gummiflua are usually an indication of a high water table. 8 acacia borleae shrublands this community also occurs on ecotones, but this time between the mopane woodlands and the grasslands. besides acacia borleae and rhigozum zambesiacum, few other woody species occur in association with colophospermum mopane. acacia borleae is generally known to occur on clayey soils. the grass layer is characterised by the dominance of eragrostis cf. heteromera in association with sporobolus spp., paspalum sp. and eragrostis viscosa. 9 cynodon dactylon panicum coloratum wooded grasslands this is an open community with a low cover and diversity of woody species. the most prominent woody species are hyphaene petersiana, colophospermum mopane, acacia nilotica and a. xanthophloea. the grass layer is dominated by cynodon dactylon, panicum coloratum, digitaria eriantha and eragrostis cf. heteromera with occasionally setaria incrassata and ischaemum afrum on more clayey and poorly drained soils. the sedge component (family cyperaceae) is very prominent. a crust of detritus and algae was found on most sample plots. this metaphyton originates as floating algae populations that aggregate along the shore line through wave action and the recession of flood waters (wetzel 2001). die-back of trees (including acacia xanthoploea) as a result of long periods of flooding was observed. 10 cynodon dactylon open grasslands these are open grasslands with hardly any woody species present. they are found in seasonally flooded areas where the high water table prevents (probably in combination with recurrent fires) any permanent presence of trees. the grass layer is dominated by cynodon dactylon, eragrostis cf. heteromera and paspalidium obtusifolium. other species are cf. acroceras macrum, sporobolus consimilis, digitaria eriantha and eragrostis gummiflua. as in community 9, the sedge component is highly visible. a crust of detritus and algae which is a residue from the flooding occurs. this most likely plays a very important role in nutrient supplementation in a community on sandy soils with limited nutrient content. below the alluvial plug (fig. 2), along the banks of the changane on the eastern boundary, the vegetation indicates almost saline characteristics with the presence of succulent halophytes and sporobolus spp. (probably s. kentrophyllus). similar-looking grasslands were observed from the air, 15 km north-northwest of pio cabral. the cynodon dactylon open grasslands are considered to be ‘natural’ (given the edaphic control in combination with fires). they do, therefore, not represent a secondary degradation product of savannas that would be expected to occur under the prevailing climatic conditions. this viewpoint is in support of the analysis by matthews et al. (1999) for the coastal grasslands of maputaland. 11 paspalidium obtusifolium open grasslands these open grasslands are found in small depressions and in the larger wetland in areas that are flooded for long periods. the most obvious difference with community 10 is the absence of cynodon dactylon. the pans are sometimes fringed by the tree combretum imberbe. other important grass species are sporobolus consimilis, eragrostis cf. heteromera, cf. acroceras macrum and echinochloa colona. in deeper water, dense stands of phragmites australis or p. mauritianus reeds and bulrushes typha capensis are found. these could not be sampled because of problems in terms of accessibility. the blue water lily nymphaea nouchali, as well as the small yellow water lily nymphoides thunbergiana and the white-flowered nymphoides indica ssp. occidentalis are very conspicuous. the communities described above are very dynamic in nature. as waters recede, comkoedoe 48/2 (2005) 52 issn 0075-6458 munity 11 gives way to community 10. community 10 can become more wooded if flooding does not occur for a number of years, it also being encroached upon by expanding acacia borleae of community 8. stands of 5 m tall dead acacia xanthophloea and mopane have been observed. this indicates that trees could establish themselves and thrive for a number of years before being killed during a long period of inundation. wetland types of the pnb the wetlands of the pnb are characterised by episodic flooding from sporadic storm events that produce ephemeral flows followed by a gradual drying out. this process is determined not only by the amount of rainfall, but also by the intensity and duration of the event. these factors, along with levels of soil saturation, determine how much of the catchment runoff reaches the wetland. the right conditions mainly tend to occur when tropical cyclones move in over the pnb catchment from the mozambique channel, mainly during the period december to march. such a hydrological regime means that water flows in to the wetland system only as a result of intense and sporadic storm events. these driving pulses result in a high degree of diversity and productivity within the pnb’s wetland systems. the continuous transition between the terrestrial and aquatic condition as a result of the cyclic flooding and drying of the pnb system creates a dynamic ecotone supporting an array of diverse temporal habitats and rich fauna and flora. the pulsed nature of inflows from within the catchment brings with it a rich sediment load high in nutrient content. as a result of this pattern, and the concentration issn 0075-6458 53 koedoe 48/2 (2005) fig. 7. landscape map of the parque nacional de banhine. affected through high evaporation rates, nutrient levels within some of the smaller wetland systems would appear to be extremely high. the following six wetland types (based on the ramsar convention classification) occur in the park: l permanent inland deltas; m permanent rivers/streams/creeks; n seasonal/intermittent/irregular rivers/streams/ creeks; p seasonal / intermittent freshwater lakes (>8ha) including floodplain lakes; tp permanent freshwater marshes/pools; ponds (below 8ha), marshes and swamps on inorganic soils; with emergent vegetation water-logged for at least most of the growing season; ts seasonal/intermittent freshwater marshes/ pools on inorganic soils; includes sloughs, potholes, seasonally flooded meadows, sedge marshes. landscapes of the pnb not all of these 11 plant communities can be mapped individually, not even using the results of the satellite imagery analysis. they often occur in a fine-scaled mosaic that reflects small variations in topography and accompanying moisture regime. these factors cannot always be mapped in sufficient detail to allow for extrapolation and correlation with the vegetation. broader units have thus been mapped out of necessity. landscapes have been defined that have bearing on management requirements and development potential as they represent a composite of topographical factors, underlying soil, hydrology and current vegetation pattern. different combinations of the plant communities can be grouped in five major landscapes. extensive use was made of the existing “carta de uso e cobertura da terra” (anonymous 1999) for the mapping of the following individual landscapes (fig. 7): wetland landscape – comprises the seasonally to permanently flooded areas in the northeastern part of the park, particularly towards pio cabral. this landscape is characterised by community 11, although communities 9 koedoe 48/2 (2005) 54 issn 0075-6458 fig. 8. sampling intensity across original land cover polygons in the parque nacional de banhine. polygons with no or only one sample indicate greater uncertainty as to their correct landscape classification. & 10 occur on levees and patches of higher ground within the landscape. large areas of open water with dense stands of phragmites australis/mauritianus reeds, typha capensis and sedges occur. this landscape holds examples of wetland types l and tp as described earlier. it covers 6 588 ha or 1.1 % of the pnb. wattled cranes grus carunculatus have been observed in the wetland landscape during 2002, 2003 and 2004 (m. stalmans pers. obs.). grassland landscape – this landscape represents the seasonally flooded areas surrounding the wetland landscape. its characteristic feature is its open grasslands (community 10), sometimes with scattered trees (community 9), bush clumps on termitaria (community 2) and acacia borleae shrublands (community 8) and hyphaene petersiana terminalia sericea trees (community 7) on its margins. ‘islands’ of sandveld, colophospermum mopane and baobabs adansonia digitata with commiphora spp. occur on slightly higher ground within the extensive grasslands. this landscape represents the flooded meadows of wetland type ts with seasonal pans as well as larger seasonal floodplain lakes of wetland type p. it covers 85 170 ha or 13.6 % of the pnb. the grassland landscape is very important to the large population of ostriches struthio camelus that is found in the park. oribi ourebia ourebi occur in this landscape, particularly towards the ecotone with the wetland landscape. mopane landscape – this landscape is found on more loamy to clayey soils, specifically where the sand mantle has been eroded away, to the north-west of the wetland and grassland landscapes. its dominant feature is the closed woodlands of communities 4 & 5. embedded within the mopane are the seasonal streams representing wetland type n (with vegetation community 3). these streams channel water from the areas north of the park towards the wetland above the alluvial plug. further downstream these streams become ill-defined, with an almost ‘braided’ character. they are wide and grassy, covered by communities 9 & 10, with community 11 in small pans within those illdefined drainage lines. this landscape therefore holds elements of wetland types n and ts. it covers 212 226 ha or 33.9 % of the parque nacional de banhine. sandveld landscape – this landscape covers the larger part of the park, particularly to the west and south. it typically consists of community 6 with patches of community 5. pans in this landscape are covered by community 11. wetland type ts is therefore represented. issn 0075-6458 55 koedoe 48/2 (2005) table 1 relative species richness of plant communities in the pnb community no. of total expected no. actual no. relative sample species of species as % of diversity plots expected 1 5 9 18 50 medium 2 5 22 18 123 medium 3 3 29 11 259 high 4 28 81 96 85 medium 5 22 102 75 135 high 6 13 83 45 185 high 7 7 24 25 97 medium 8 2 9 8 115 medium 9 13 26 45 58 low 10 12 9 42 22 low 11 5 6 18 33 low this is the most extensive landscape and covers 292 350 ha or 46.7 % of the pnb. this is similar to the situation in the limpopo national park where the nwambia sandveld landscape covers 41 % of the park. sandveld is not well represented in the kruger national park on the south african side of the greater limpopo transfrontier park. androstachys (nsimbitsi) landscape – this landscape is very uniform and mostly consists of community 1. it is interleaved with the sandveld landscape in the extreme northwestern corner of the park and with the mopane and sandveld landscapes in the extreme south. it covers 29 688 ha or 4.7 % of the pnb. the map boundaries are drawn subjectively because of the fact that there is seldom a definite and visible border between landscapes, but rather a gradient in terms of vegetation and topography. the sampling did not cover all land cover polygons (fig. 8), with 12 % of the pnb not being assessed at all. the greatest uncertainties with regard to the landscape map are along the western boundary and in the south-eastern corner of the pnb. plant community and landscape diversity the available time frame and limited ground coverage did not allow for a detailed inventory of plant diversity. however, a relative measure of plant community diversity can be derived by comparing the total species count to the sampling intensity for each respective community. the number of species per community in relation to the expected species number (based on the average accumulated number of species per sample plot) is depicted in table 1. communities 3 (riverine), 5 (partly sandveld) and 6 (pure sandveld) are relatively more species-rich than expected from the number of samples assessed. in particular, the sandveld of community 6 much exceeds the expected value. the same applied for sandveld in the limpopo national park (stalmans et al. 2004). land use patterns and their influence on the vegetation of the pnb a number of people reside within the boundaries of the pnb. they practise mainly subsistence farming and rely on the fish resources of the wetland. subsistence farming takes mostly place on bushclumps, in mopane woodlands and in the wetland. bushclumps of community 2 are cleared and maize is planted in the fertile soil of the termitarium. the number of bushclumps that have been destroyed in this manner seems fairly limited. however, it can be anticipated that recovery will take many years. the whole functioning (and therefore recovery) may be affected by the loss of the emergent xanthocercis zambesiaca trees. manual clearing of mopane woodlands (communities 4 & 5): with stacking of the material around the bigger trees is followed by burning that kills those trees. crops are planted in the ashbed that increases fertility. crops can only be produced for a very few years before these lands must be left fallow to restore soil fertility. the total area that has been cleared and is in some stage of recovery is therefore much larger than the extent of land that is actively cultivated in any given year. as waters recede, crops are planted in the wetland (community 11). the extent of cultivated land is relatively limited and the impact is probably fairly small as the next flood cycle probably ‘resets’ the cultivated patch and surrounding areas to a similar state. use is made of trees for firewood, building of dwellings, livestock pens, and carving of various items. the impact of these activities is selective. whereas this use does not seem to impact significantly on the overall woody resource, no data are available on the impact on specific species. grass is cut and used for thatching (mostly heteropogon contortus). much use is made of medicinal plants and wild fruits (eg. strychnos spp.). no data are available regarding the impact thereof on specific species. the hyphaene palms are koedoe 48/2 (2005) 56 issn 0075-6458 heavily tapped for their sap. this involves the cutting of the stems. as a result very few tall palms have been observed in the field. the devil’s claw harpagophytum procumbens (family pedaliaceae) occurs in the pnb (dr a.b. (tony) cunningham pers. comm. 2003). the root tubers of this plant are traded internationally and concern has been expressed at the cites level regarding its situation (anonymous 2003). levels of herbivory (both grazing and browsing) seem presently very limited in the pnb. current numbers of game are certainly very low. livestock numbers are also relatively limited including cattle and goats. during the subjective scoring of the sample plots only 7 out of 115 plots had a medium level of grazing pressure and one had a medium browsing impact (old elephant utilisation). no heavy grazing was observed. the grass and tree layer are generally in a good condition. it offers suitable habitat for the herbivores that could be expected in this environment (both those occurring historically and those still present). the only alien plant species observed was the prickly pear (opuntia sp.). it occurs in a few dense stands near the village of tchove. fires are very prominent within the pnb with signs of old and recent fire events visible throughout the landscape. although a number of fires likely originate from lightning, the most probable cause of fire is through the activities of the inhabitants of the pnb, e.g., clearing of lands, producing a green flush for livestock grazing, smoking out of beehives. within the constraints of the present study it was not possible to assess the fire return period and its appropriateness. of interest is the relatively small-scale pattern that can be observed in the field. this pattern of spatially and temporally varying fire parameters (frequency, seasonality, intensity and type of fire) across the landscape is likely to be required for the maintenance of diversity (brockett et al. 2001; gill & mccarthy 1998). acknowledgements dr ken tinley provided unpublished data which he collected during the 1970s. these data proved invaluable in describing the functioning of this wetland system. warden armando nguenya provided logistical support and accompanied the survey team in the field. dr jeremy anderson assisted with access to the area, provided assistance with the vegetation sampling and organised over flights of the pnb. members of the bateleur group of pilots (in particular joe holmes and avroy shlain) graciously provided fixed-wing and micro-light flying time. craig beech of the peace parks foundation assisted with gis data. mervyn lötter, ernst schmidt, warren mcleland and john burrows of the plant specialist group assisted with species identification. the vegetation and hydrological research was undertaken as part of the planning work by development alternatives inc that was funded by usaid in support to the direcção nacional de áreas de conservação of the ministério do turismo, moçambique. more specifically, the management plan for the pnb was compiled by the ecologists dr d grossman and ms p holden. references anonymous. 1995. legenda da carta nacional de solos (escala 1:1.000.000). maputo, moçambique: compilada pelo departemento terra e agua. anonymous, 1997. first national report on the conservation of biological diversity in moçambique. ministry for the coordination of environmental affairs. maputo. anonymous. 1999. carta de uso e cobertura da terra. moçambique: joint venture ignfi cenacarta dinageca. anonymous. 2002. banhine national park, management and development plan. unpublished report to the national directorate for conservation areas, ministry of tourism, republic of moçambique. anonymous. 2003. devils’ claw (harpagophytum procumbens). fauna & flora 4 (april 2003): 28. brockett, b.h., h.c. biggs & b.w. van wilgen. 2001. a patch mosaic burning system for conservation areas in southern african savannas. international journal of wildland fire 10:169183. coetzee, b.j. 1983. phytosociology, vegetation structure and landscapes of the central district, kruger national park. dissertationes botanicae 69: 1-456. issn 0075-6458 57 koedoe 48/2 (2005) edwards, d. 1983. a broad scale structural classification of vegetation for practical purposes. bothalia 14: 705-712. gabriel, h.w. & s.s. talbot. 1984. glossary of landscape and vegetation ecology for alaska. blm-alaska technical report 10. u.s. department of the interior. gauch, h.g. 1982. multivariate analysis in community ecology. cambridge: university press. gertenbach, w.p.d. 1983. landscapes of the kruger national park. koedoe 26:9-121. gill, a.m. & m.a. mccarthy. 1998. intervals between prescribed fires in australia: what intrinsic variation should apply? biological conservation 85:161-169. hill, m.o. 1979. twinspan a fortran program for arranging multivariate data in an ordered twoway table by classification of the individuals and attributes. ithaca, new york: cornell university. jongman, r.h.g., c.f.j. ter braak & o.f.r. van tongeren. 1987. data analysis in community and landscape ecology. wageningen: pudoc. kelly, r.d. & b.h. walker. 1976. the effect of different forms of land use on the ecology of a semi-arid region in south-eastern rhodesia. journal of ecology 64:553-576. land type survey staff. 1989. land types of the map 2530 barberton. memoirs of the agricultural natural resources of south africa 13. leemans, r. & w. cramer. 1991. the iiasa database for mean monthly values of temperature, precipitation and cloudiness of a global terrestrial grid. international institute for applied systems analysis (iiasa). rr-91-18. matthews, w.s., a.e. van wyk & n. van rooyen. 1999. vegetation of the sileza nature reserve and neighbouring areas, south africa, and its importance in conserving the woody grasslands of the maputaland centre of endemism. bothalia 29(1):151-167. mueller dombois, d. & h. ellenberg. 1974. aims and methods of vegetation ecology. new york: john wiley. national working group for vegetation ecology. 1986. soil classification according to the binomial classification system. technical communication 3. scholes, r.j. & b.h. walker. 1993. an african savanna. cambridge cambridge: university press. siebert, f., g.j. bredenkamp & s.j. siebert. 2003. a comparison of mopaneveld vegetation in south africa, namibia and zimbabwe. bothalia 33(1):121-134. stalmans, m. 1994. vegetation survey of malilangwe. unpublished report to the malilangwe conservation trust. stalmans, m., w.p.d gertenbach & f. carvalhoserfontein. 2004. plant communities and landscapes of the parque nacional do limpopo, moçambique. koedoe 47(2): 61-81. ter braak, c.j.f. 1986. canonical correspondence analysis: a new eigenvector technique for multivariate direct gradient analysis. ecology 67(5): 1167-1179. ter braak, c.j.f. 1992. canoco a fortran program for canonical community ordination. ithaca, new york, usa: microcomputor power. timberlake, j.r., n. nobanda & i. mapaure. 1993. vegetation survey of the communal landsnorth and west zimbabwe. kirkia 14(2): 171270. van rooyen, n., g.k. theron & n. grobbelaar. 1981. a floristic description and structural analysis of the plant communities of the punda milia-pafuri-wambiya area in the kruger national park, republic of south africa: 2. the sandveld communities. journal of south african botany 47(3):405-449. werger, m.j.a. & b.j. coetzee. 1978. the sudano-zambezian region. in: werger, m.j.a. (ed.). biogeography and ecology of southern africa. the hague: w. junk. wetzel, r.g. 2001. limnology: lake and river ecosystems. 3rd edition. san diego: academic press. wild, h. & l.a.g. barbosa. 1967. supplement of the vegetation map of the flora zambesiaca area. pp. 1-71. in: wild, h. & a. fernandes. vegetation map of the flora zambesiaca region. harare, zimbabwe. koedoe 48/2 (2005) 58 issn 0075-6458 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe reviewer acknowledgement 101 koedoe african protected area conservation and science koedoe african protected area conservation and science recognises the value and importance of the peer reviewer in the overall publication process – not only in shaping the individual manuscript, but also in shaping the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely, productive peer review process. we would like to take this opportunity to thank all reviewers who participated in shaping this volume of koedoe: angela gaylard anna spenceley brian van wilgen charles haddad ed witkowski izak smit jane carruthers johann du preez kate matchett kelly scheepers we appreciate the time taken to perform your review successfully. in an effort to facilitate the selection of appropriate peer rievewers for koedoe, we ask that you take a moment to update your electronic portfolio on www.koedoe.co.za for our files, allowing us better access to your areas of interest and expertise, in order to match reviewers with submitted manuscripts. if you would like to become a reviewer, please visit the journal website and register as a reviewer. to access your details on the website, you will need to follow these steps: 1. log into the online journal at http://www. koedoe.co.za 2. in your ‘user home’ [http://www.koedoe. co.za/index.php/koedoe/ user] select ‘edit my profile’ under the heading ‘my account’ and insert all relevant details, bio statement and reviewing interest. 3. it is good practice as reviewer to update your personal details regularly to ensure contact with you throughout your professional term as reviewer to koedoe. please do not hesitate to contact me if you require assistance in performing this task. rochelle bronkhorst submissions@koedoe.co.za tel: +27 (0)21 914 5100 fax: +27 (0)21 914 5105 leon lotz lesley henderson leslie brown louise swemmer mahlomola daemane mathilda (tilly) van niekerk michael panagos sally archibald stefan siebert theo mostert whittington.qxd short note new breeding locality for crowned cormorant p.a. whittington issn 0075-6458 125 koedoe 47/2 (2004) the crowned cormorant phalacrocorax coronatus is endemic to the southern african subregion with an estimated population of about 2700 pairs, breeding at 48 localities between walvis bay, namibia and cape agulhas, south africa (crawford et al. 1994, crawford 1997). the easternmost point at which breeding has previously been recorded is 2 km west of aasfontein (34°46's, 19°50'e), where 35 nests were recorded in january 1981 (crawford et al. 1982). non-breeding birds have been seen further to the east as far as holkom meester se baai (34°23's, 21°49'e) (crawford et al. 1982). on 29 october 2003, counts were made of breeding seabirds at the number 12 stacks (33°59's, 23°35'e) in the de vasselot section of the tsitsikamma national park. counts were made of occupied nests and fledged young of all species present on three stacks visible from the mainland. these three stacks were thought to be those labelled as carbo, capensis and dominicanus by crawford (1983). while counting nests of white-breasted cormorants phalacrocorax carbo lucidus and cape cormorants phalacrocorax capensis, four occupied nests of crowned cormorants were located on the stack corresponding to that named carbo by crawford (1983). all four had a sitting bird and it was not possible to see whether the nests contained eggs or chicks. most occupied nests of nearby white-breasted and cape cormorants contained chicks. in addition, 20 fledged white-breasted cormorants and 71 fledged cape table 1 counts of occupied nests, old nests and fledged young of all species breeding at number 12 stacks, 29 october 2003 stack carbo capensis dominicanus white-breasted cormorant phalacrocorax carbo occupied nests (total) 15 0 0 nests with chicks visible 6 0 0 old nests 13 0 0 fledged young 20 0 0 cape cormorant phalacrocorax capensis occupied nests (total) 27 2 8 nests with chicks visible 25 0 6 old nests not not not counted counted counted fledged young 71 103 94 crowned cormorant phalacrocorax coronatus occupied nests (total) 4 0 0 nests with chicks visible 0 0 0 old nests 0 0 0 fledged young 0 0 0 little egret egretta garzetta occupied nests (total) 1 0 0 nests with chicks visible 0 0 0 old nests 0 0 0 fledged young 0 0 0 kelp gull larus dominicanus occupied nests (total) 1 11 5 nests with chicks visible 0 0 0 old nests 0 0 0 fledged young 0 0 0 cormorant chicks were counted on this stack (table 1). the solitary little egret egretta whittington.qxd 2004/10/05 12:57 page 125 garzetta nest contained one egg. crowned cormorants can breed throughout the year (rand 1960). at malgas island off the west coast of south africa, most breeding takes place between september and april with a peak in breeding activity between december and march (crawford et al. 1999). it is thought that the birds recorded at number 12 stacks were either incubating eggs or brooding small chicks. the potential for the crowned cormorant to extend its breeding range on finding suitable breeding sites is illustrated by the construction of the bird rock platform north of walvis bay, namibia, and the wreck of the meisho maru no. 8. these resulted in previous breeding range extensions of 415 km to the north and 16 km to the east respectively (crawford et al. 1994). this observation extends the breeding range of the crowned cormorant by approximately 355 km to the east and is about 169 km east of the nearest observations made of non-breeding birds. the species was not recorded by previous seabird surveys made in tsitsikamma national park (crawford 1983). acknowledgements support for this research from the university of port elizabeth and from the national research foundation is gratefully acknowledged. i am indebted to andre riley of south african national parks for arranging access to seabird colonies within tsitsikamma national park at very short notice, to karools peterse for guidance in the field and to peter and carol humphrey for their wonderful hospitality. r.j.m. crawford, john cooper and n.t.w. klages commented on the draft manuscript. references crawford, r.j.m. 1983. some observations on seabirds breeding in the tsitsikamma coastal national park. koedoe 26: 145-152. crawford, r.j.m. 1997. crowned cormorant phalacrocorax coronatus. pp 38-39. in: harrison, j.a., d.g. allan, l.g. underhill, m. herremans, a.j. tree, v. parker & c.j. brown. (eds.). the atlas of southern african birds. vol 1. non-passerines. johannesburg: birdlife south africa. crawford, r.j.m., p.a. shelton, r.k. brooke & j. cooper. 1982. taxonomy, distribution, population size and conservation of the crowned cormorant, phalacrocorax coronatus. le gerfaut 72: 3-30. crawford, r.j.m., b.m. dyer & r.k. brooke. 1994. breeding nomadism in southern african seabirds-constraints, causes and conservation. ostrich 65: 231-246. crawford, r.j.m., b.m. dyer & l. upfold. 1999. seasonal pattern of breeding by cape and crowned cormorants off western south africa. ostrich 70(3&4): 193-195. rand, r.w. 1960. biology of guano-producing seabirds 3. distribution, abundance and feeding habits of the cormorants phalacrocoracidae off the south western coast of the cape province. investigational report department of commerce and industries, division of sea fisheries. 42: 1-32. p.a. whittington, department of zoology, university of port elizabeth, box 1600, port elizabeth, 6000, south africa (philip.whittington@upe.ac.za) koedoe 47/2 (2004) 126 issn 0075-6458 whittington.qxd 2004/10/05 12:57 page 126 filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! article information authors: stefan w. grab1 andrew s. goudie1,2 heather a. viles1,3 nicola webb1 affiliations: 1school of geography, archaeology and environmental studies, university of the witwatersrand, south africa 2st cross college, university of oxford, united kingdom 3school of geography and the environment, university of oxford, united kingdom correspondence to: stefan grab email: stefan.grab@wits.ac.za postal address: private bag x3, witwatersrand university 2050, south africa dates: received: 04 jan. 2010 accepted: 12 nov. 2010 published: 14 mar. 2011 how to cite this article: grab, s.w., goudie, a.s., viles, h.a. & webb, n., 2011, ‘sandstone geomorphology of the golden gate highlands national park, south africa, in a global context’, koedoe 53(1), art. #985, 14 pages. doi:10.4102/koedoe.v53i1.985 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) sandstone geomorphology of the golden gate highlands national park, south africa, in a global context in this original research... open access • abstract • introduction    • study area • methodology • geological context    • ‘blueberries’ (concretions)    • case hardening    • surface spalling (exfoliation, flaking)       • sandstone geomorphological map of the gghnp       • arches (natural stone bridges)       • pedestal rocks (mushroom-shaped pillars)       • cavernous weathering       • rock basins (weathering or solutional pits, gnammas)       • rock doughnuts       • polygonal cracking (tessellation or alligator cracking)       • bedrock drainage microchannels (fluting, gutters, karren forms)       • lichens and weathering • discussion and conclusion • acknowledgements • references abstract (back to top) the golden gate highlands national park (gghnp) is well known for its impressive sandstone formations. while previous geoscience research in the park has focused on geology, palaeontology, slope forms and the prominent lichen weathering, remarkably little has been written on the diversity and possible origins of sandstone phenomena in the region. the objectives of this study were (1) to present a geomorphological map of prominent and interesting landforms for particular portions of the park and (2) to document the variety of macroand microscale sandstone formations observed. during field work, we undertook global positioning system measurements to map landforms and, in addition, measured the dimensions of several landform types. a schmidt hammer was used to conduct rock hardness tests at a variety of localities and lithologies for comparative purposes. we indentified and mapped 27 macroand microscale sandstone landforms, of which 17 are described in detail. it is demonstrated that for the most part, the landforms are a likely product of surface lithological reactions to a regional climate characterised by pronounced multitemporal temperature and moisture shifts, recently and in the past. however, many of the geomorphological processes producing landforms are controlled by microclimates set up by factors such as macroand microtopography. conservation implications: the gghnp is best known for its geological, geomorphological and palaeontological heritage. this paper highlights the diversity of sandstone geomorphological phenomena, many of them rare and ‘unique’ to the region. not only are these landforms of aesthetic interest to tourists, but they also provide microhabitats for biota. thus, conservation of biota requires associated conservation of geo-environments where they are established. introduction (back to top) the work sandstone landforms (young, wray & young 2009) provides a global view of our current understanding of macroand microscale sandstone landforms and their associated geomorphic processes. it provides numerous examples of sandstone landforms from around the world and highlights the many important new studies that have appeared from south america, australia, central europe and china, in particular. although the book identifies the clarens area of the northeastern free state as one of the world’s 61 major sandstone landscapes, relatively little is known about the diversity of sandstone phenomena and their process origins in southern africa. despite sandstones not generally being as well known for their distinctive landforms as are limestones and granites (robinson & williams 1994), the golden gate highlands national park (gghnp) hosts some of southern africa’s most spectacular scenery. the gghnp has been identified as a geotourism hotspot, with the defining feature being the ‘red, cream and yellow weathered sandstone of the clarens formation that is deeply eroded to form bluffs and cliffs’ (reimold, whitfield & wallmach 2005:55). in recent years several world heritage sites have been proclaimed in south africa, and more than half of those currently listed and proposed represent entirely natural geological heritage (reimold et al. 2005). the gghnp falls within the drakensberg world heritage region, which boasts mixed heritage status owing to the presence of both culturally and naturally significant features. it is thus not surprising that the gghnp has become a world-famous geotourism destination, with particular attractions including the brandwag ‘golden gate’ (figure 1), san rock art and vertebrate fossils. the gghnp was established in 1963 for the purpose of protecting a pristine area, but more specifically, to conserve the impressive sandstone formations (hence its name) and the montane and afro-alpine grassland biome (sanparks 2004). previous earth science research in the park has focused mainly on the geological and palaeontological heritage and macroscale geomorphological evolution (e.g. groenewald 1986; kitching & raath 1984; spies 1969), slope forms and processes (moon 1991; moon & munro-perry 1988; nicol 1973), biogenic weathering (büdel et al. 2004; wessels & büdel 1995; wessels & schoeman 1988; wessels & wessels 1991, 1995; wessels et al. 1995), weathering basins (cooks & pretorius 1987) and the origins of ‘cave’ formations (le roux 1978). despite these previous studies and the recognition of the regional geoheritage, the diversity of both macroand microscale sandstone landforms has not yet been comprehensively documented or mapped in the gghnp. to this end, our objective was, firstly, to provide a map of prominent and interesting sandstone phenomena for particular portions of the park, thus contributing to enhanced knowledge on geodiversity. secondly, we aimed to document the variety of macroand microscale sandstone morphologies (table 1) and their possible process origins within a global context. although we concentrate here on sandstone features, some of our observations may also be relevant to granitic terrains, which often have similar assemblages of weathering forms (see twidale & vidal romaní 2005). figure 1: the brandwag ‘golden gate’, displaying clarens formation sandstone cliffs. table 1: a scale hierarchy of sandstone landforms in the golden gate highlands national park. study area the gghnp is located in the eastern free state (28°31’s, 28°37’e) and covers an area of 120 km2 across the rooiberg mountain range (1892–2837 m a.s.l.), which forms part of the greater drakensberg–maluti mountain system (figure 2). although annual precipitation varies considerably across the region, it averages approximately 764 mm, of which most falls between november and april (groenewald 1986). summers are mild to warm (mean temperature ranges from 13 °c to 26 °c) and winters cold (mean temperature ranges from 1 °c to 15 °c). (cooks & pretorius 1987). frost is widespread during the winter months and snow occasionally falls on the higher peaks in the park. the vegetation is typically highlands sourveld and themeda–festuca alpine veld, with identified plant communities representative of the tristachya leucothrix – trachypogon spicatus grassland of moist mountain slopes and plateaus, and to some extent the merxmuellera drakensbergensis – festuca caprina high-altitude austro-afro alpine grassland (du preez & bredenkamp 1991; kay, bredenkamp & theron 1993). the area is located on the african surface of partridge and maud (1987) and its drainage systems have undergone subsequent incision (moon & munro-perry 1988). the observed landscape has been influenced by ever-changing climates, particularly during the quaternary period, which encountered an intensification of cold climate processes and also major changes in vegetation cover (henderson et al. 2006). the landforms of the area thus have a long and complex geomorphic history. figure 2: location of the golden gate highlands national park and map of the park showing the insets for the geomorphological maps in figure 3. cliffs. methodology (back to top) large portions of the sandstone sections of the gghnp were traversed, with sandstone phenomena photographed and recorded using a global positioning system device. the recorded phenomena could then be incorporated onto existing topographic and geological maps. for some landforms and topographic settings we undertook morphometric and rock hardness measurements using a tape measure and a schmidt hammer (classic n-type), respectively. the schmidt hammer indicates rebound values (r-values ranging from 0 to 100), which are proportional to the hardness of the rock. rock hardness as described here is based on the indentation produced when pressure is exerted on the rock mass (as opposed to individual mineral hardness, which indicates resistance to scratching). rock surfaces were first cleaned (smoothed) using a carborundum, after which the hammer was applied to the smoothed surface. mean rebound values were calculated from 30–50 readings per site. the objective with this was primarily to compare the hardness of clarens formation sandstone with that of other geological units in the park and consequently its potential strength variability in various spatial and geological contexts. young et al. (2009) classify sandstone phenomena into four hierarchical groupings, based on morphology rather than process origins. we classify the sandstone landforms observed in the gghnp according to size and morphological attributes. however, given that microsurface forms such as case hardening and spalling (flaking) contribute to the development of other sandstone features (e.g. rock basins and tafoni), these are dealt with first as part of the geological structure which invariably impacts on landform genesis. while a variety of landform types are discussed within separate subsections for ease of presentation, it should be acknowledged that several varieties of sandstone phenomena may be closely linked and have common developmental origins despite their different morphological surface expressions. such differences may be due to macroand microtopographic position, existing surface morphology and differences in underlying geological controls. for instance, rock doughnuts, rock basins and rock pans within a region may have common process geomorphic origins but the outcomes may spatially take on rather different visible forms. geological context (back to top) the sedimentary portion of the late carboniferous – middle jurassic karoo basin of southern africa includes glacial (dwyka group), marine, swamp and deltaic (ecca group), fluvial (beaufort group and parts of the stormberg group) and aeolian (upper part of the stormberg group) deposits, and is capped by the volcanic drakensberg group (catuneanu, hancox & rubidge 1998; catuneanu et al. 2002; smith et al. 1998). sills and dykes were emplaced in sedimentary strata around 183 ma (holzförster 2007) and in places may form up to 70% of the basin volume (rowsell & de swardt 1976; svensen et al. 2006). the stormberg group is made up of the molteno, elliot and clarens formations, all of which are represented in the gghnp (eriksson 1981, 1984, 1985; groenewald 1986). while the elliot formation has a distinct lower contact with the molteno formation, it has a more gradual upper contact with the clarens formation (bordy, hancox & rubidge 2004; groenewald 1986). the succession reflects changing palaeo-environmental conditions (climatic and basin foreland tectonics) during deposition; from dominantly fluvial (molteno formation) to increasingly aeolian (clarens formation) (bordy et al. 2004; eriksson 1981, 1984, 1985; groenewald 1986). sandstones are produced through the cementation or lithification of sand grains through processes of pressure melting and recrystallisation, compaction and consolidation of incorporated clay minerals, and/or by the precipitation of authigenic minerals from water migration through pores (robinson & williams 1994). although a wide variety of sandstones occur in the gghnp, primarily owing to a diversity of depositional mechanisms, they are, for the most part, a product of compaction and consolidation over time. the molteno formation varies in thickness from less than 10 m to more than 100 m and consists of light-coloured sandstones of which the grains can range from fine to very coarse (eriksson 1984). the molteno formation sandstones are primarily quartz-rich feldspathic wackes, have a high interstitial content and sometimes tend towards subordinate argillaceous and conglomeritic structures (eriksson 1984). the elliot formation varies in thickness from approximately 28 m to more than 150 m and consists of red-coloured, argillaceous sandstone with grains ranging from fine to coarse (eriksson 1985). the elliot formation is particularly well known in the gghnp for its vertebrate fossil content (e.g. groenewald 1986; kitching & raath 1984). the clarens formation is typically 115 m – 195 m thick and consists of light-coloured, fine-grained sandstones, sandy siltstones and mudstones, and subordinate coarser-grained components (eriksson 1981). fresh faces of the rock collected from boreholes and quarries are white, yet on exposure the rock weathers to give its characteristic tan or golden colour. however, there have been no detailed studies on the chemical and mechanical weathering processes causing such colour alterations. the sedimentary strata, which are generally approximately horizontally bedded, have been intruded by several fineto coarse-grained dolerite dykes and a few sills in the gghnp (groenewald 1986). some major faults occur in association with these intrusions and these are likely to have played a role in the genesis of the landforms, as has the presence of the bedding planes and joints within the sedimentary rocks. schmidt hammer testing of the rocks in the area provides estimates of the hardness characteristics of the various rock units (table 2). the mean schmidt hammer hardness r-value for the clarens formation sandstone, based on 30–50 readings from each of nine sites, was 48.91. this is marginally higher than the r-values that have been recorded at two other major sandstone geomorphological sites: approximately 45 for the navajo sandstone of utah (irwin et al. 2009) and 41–44.7 for the cambrian–ordovician sandstones of wadi rum in jordan (goudie et al. 2002). the r-value for a dolerite dyke piercing the clarens formation sandstone was 63.0, while that for drakensberg group basalt was 55.92. sandstone of the elliot formation had an r-value of 58.12. thus, all the main rock units in the area are relatively hard. however, the wide variety of rock hardness values (compressive strength) obtained across sandstone surfaces in the gghnp attests to both macroand microscale variability in porosity, composition and bonding of mineral grains, and the extent of surface weathering or case hardening. although not yet locally tested, it is thus likely that many of the sandstone morphologies at the gghnp are a product of differential rates and processes of weathering and erosion across microscales, owing at least in part to rock hardness variability. table 2: hardness values as measured with a schmidt hammer (classic n) for rock types in the golden gate highlands national park. ‘blueberries’ (concretions) calcareous and ferruginous concretions are particularly common to the sandstones of the clarens formation (groenewald 1986), where they impact on the fine-scale spatial dynamics of weathering processes. such concretions are sometimes informally referred to as ‘blueberries’, possibly owing to their nodular spheroidal structure (squyres & knoll 2005). according to mahaney et al. (2008), it is thought that such surface nodules are the product of subsurface fluid migration through porous sandstone. fluids become enriched with iron and calcite, with precipitates eventually forming around a central nucleus of fine sandstone particles (mahaney et al. 2008). these are released when the host sandstone is weathered and produce a surface lag. case hardening an important and common attribute of many sandstone surfaces is the development of a surface crust or ‘outer [shell] more resistant to erosion than interior material’ (dorn 2004:118). crust development is primarily caused by the deposition of solutes when pore water migrates towards the surface and evaporates (robinson & williams 1994). however, the processes causing case hardening may be polygenetic in origin and vary between sites. for example, it has been found that micro-organic biofilms may aid the cementation process to produce case-hardened surfaces in jordanian sandstones (viles & goudie 2004). examples of case hardening occur throughout the gghnp where the iron-rich crusts produce hardened, reddish surfaces. the importance of case hardening cannot be underestimated: on the one hand, it protects underlying rock from weathering and, on the other, it reduces localised infiltration and so facilitates enhanced surface runoff. both these functions consequently impact on weathering and erosion on the surrounding uncrusted rock surfaces. in addition, the disintegration of the surface crust provides new irregularities that may promote localised weathering and erosion and consequently lead to new phenomena such as rock basins (weathering pits) (goudie & migón 1997). surface spalling (exfoliation, flaking) in arid and semi-arid regions (or microsettings) with seasonal rainfall (humidity) patterns, sandstone may be susceptible to spalling or flaking (i.e. thin rock flakes peeling off). in such instances cracks develop parallel to the surface, typically at a depth of between 5 mm and 50 mm (robinson & williams 1987, 1992). spalling of rock surfaces is common where cracks develop below a case-hardened surface, or along the backwalls of tafoni caverns and rock shelters. it is understood that a primary mechanism for such sandstone spalling is the dissolution of silica and other cementing agents during wetting cycles and the translocation in the surface layers during drying cycles (robinson & williams 1992). crystallisation of salt during desiccation cycles has also been observed to cause spalling (goudie & viles 1997; holmer 1998). observations on the backwalls of cavernous weathering features show that in the gghnp there are salt efflorescences associated with spalling rock. it has also been suggested that contemporary sub-zero air temperatures at the gghnp during the winter period may contribute to frost action (spalling) where moisture is locally available (marker 1990); however, freeze-thaw processes are considered contentious given that rock thermal data required to prove such a mechanism have not been forthcoming (hall, marker & le roux 1991).a further mechanism causing widely distributed spalling on rock surfaces in the gghnp is substrate alkalisation during the photosynthesis of cryptoendolithic cyanobacteria (büdel et al. 2004). this process is particularly prominent on the surfaces of clarens formation sandstones. biofilms dominated by cryptoendolithic cyanobacteria are common here and the in situ ph values of the sandstone typically range between 9.5 and 10.5 along endolithic growth zones, which enhance dissolution of sio2 (büdel et al. 2004). sandstone geomorphological map of the gghnp we have mapped both macroand microscale sandstone phenomena on a common base map to indicate prominent and/or unique features that may be of particular interest to geotourists (figure 3). while the resolution does not permit detailed information at the metre scale or the mapping of all phenomena, it does allow for the identification of prominent or abundant forms within particular areas of the gghnp, which can guide visitors in making associations between local relief and/or geology. figure 3 provides a legend of the mapped sandstone phenomena in the gghnp, most of which are discussed in detail later. general observations suggest that although most macroand microscale sandstone phenomena occur widely within the various sedimentary formations of the karoo supergroup, the uppermost broad sandstone plateaus of the clarens formation host the highest density and greatest variety of sandstone landforms. notably, such plateaus are topographic positions capable of storing considerable quantities of surface water during the wet season (particularly where infiltration is relatively low or drainage is poor), while also being subjected to substantial heating and cooling cycles because of exposure. to this end, paradise (2002) found that aspect-controlled weathering processes and associated landform production are particularly pronounced in regions where wetting-drying and heating-cooling cycles are important weathering mechanisms, such as have been observed in petra, jordan. dolerite dyke intrusions are also likely to have influenced landform development within the adjacent sandstones in the gghnp, and we note phenomena such as flared slopes, a rock arch, caverns, honeycombs and rock basins adjacent to some of the dykes. figure 3: geomorphological map of sandstone phenomena in the golden gate highlands national park (geological overlays modified after geological series 2828 harrismith, council for geoscience, 1998; localities of subfigures a–c as shown in figure 2). slope forms 1. general slope forms and processes: as a consequence of fluvial incision by the little caledon and its tributaries, the gghnp has a variety of slope forms, ranging from horizontal sandstone pavements along broad interfluves to vertical cliffs along valley margins. the variable nature of bedrock forms (gradients) has to a large extent controlled the development of microscale sandstone phenomena through controlling the amount of solar radiation received at the rock surface (rock surface microclimatology), moisture retention, surface water movement and gravity. there is likely feedback as the microscale phenomena contribute to the further evolution of the larger slope forms. the prominent slope forms in the gghnp are rectilinear cliff–talus slope combinations and bedrock slopes inclined at 30° – 35° (moon & munro-perry 1988). the presence of a resistant and relatively massive caprock (normally clarens formation sandstone) permits parallel slope retreat and the preservation of rectilinear slopes where the rock mass strength values are higher than 70. however, where values are lower, the rock is too weak to preserve the richter slope forms and thus undergoes a process of ‘slope replacement’ (moon 1991; moon & munro-perry 1988). most valley sides typically have near-vertical cliffs and/or steeply inclined (> 35°) bedrock surfaces, below which is a zone of discontinuous debris cover (‘vegetated islands’) on somewhat gentler slopes (20° – 35°). these ‘vegetated islands’ (figure 4a) appear to be the result of peat rafts’ sliding over the underlying bedrock, although there is evidence that the movements are complex and involve diffusive, waterborne erosion of sediment as well as sliding. finally, there is a zone of debris accumulation on lower slope units approaching the valley floor (moon & munro-perry 1988; nicol 1973). the number and size of some of the blocks on slopes below the cliffs is impressive and shows the importance of rock falls from the cliffs above. prominent ‘cirque-shaped’ hollows frequent several of the south-facing slopes in the gghnp and these were initially thought to have originated from nival and frost-related processes during past pleistocene cold phases, as were rockfall debris and thick colluvial mantles along the valley margins (marker 1989; nicol 1973). however, such cold-process origins were subsequently contested (le roux & marker 1990) and have not been given much further consideration. 2. cliffs and pillars: cliffs and pillars represent steep bedrock exposures, especially along deeply incised valleys or around the periphery of plateaus (robinson & williams 1994). the conspicuous cliffs of the gghnp are most prominent along the caledon valley and are primarily associated with the massive and strongly cemented fine-grained clarens formation sandstones. although cliffs are also typical of the molteno and elliot formations, the generally thicker and more stable clarens formation displays the most impressive and extensive cliff development in the region. the height of the cliffs, which in places exceed 100 m to make the cliffs among the highest in the world (moon & munro-perry 1988), is to some extent controlled by the thickness of the sandstone formations (i.e. sedimentary beds) and the associated extent of basal weathering and erosion. the best-known cliff in the gghnp is the ‘brandwag buttress’ (figure 1), alongside which the clarens formation sandstone has been intruded by a dolerite dyke, and consequently has yielded a hard quartzitic sandstone with greater resistance to weathering (groenewald 1986). the recent collapse (i.e. during the last few hundred years) of joint-bounded blocks at a few localities also provides evidence that cliff development is an ongoing, contemporary process in the gghnp. the formation of pillars is limited in the gghnp as joints are relatively widely spaced and the sandstone structures are massive and coherent. nevertheless, block structures along cliffs have developed in places owing to accelerated weathering and erosion along more closely spaced joints. another feature of the valley side slopes is the local development of summit and buttress tors. arches (natural stone bridges) despite their unusual and aesthetic appeal, natural arches and bridges are among the least studied of sandstone landforms worldwide. fairbridge (1968) classified features into ‘arches’ (which do not span a stream channel) and ‘bridges’ (which span a stream channel). a photograph (figure 4b) and painting by johan van reenen, dating back to between 1930 and 1940, depict the former occurrence of a natural stone bridge spanning the zulu hoek stream in the park. although few known natural stone bridges exist in the region today, some small natural arches presently occur in the park and the surrounding area. elsewhere, such arches are said to be the product of enhanced weathering and water infiltration along parallel joints on cliff faces, eventually leading to material collapse and production of the ‘window area’ (leasure, kind & busby 2007). figure 4: (a) ‘vegetated islands’ overlying sandstone outcrops, (b) a natural stone bridge spanning the zulu hoek stream (since collapsed; electronic copy of original photo provided by johan taljaard, gghnp), (c) water seepage and associated black staining at the rear of a cavern and (d) large rock basins with distinct water level notches. pedestal rocks (mushroom-shaped pillars) relatively little research has focused on sandstone rock pedestals and mushroom-shaped pillars, also described elsewhere as rock columns and bulbous pinnacles (mueller & twidale 2002). nevertheless, it is understood that such forms are the likely result of differential rates of weathering, which are enhanced along existing fracture-defined blocks and sheets in bedrock. pedestals occasionally occur on slopes in the gghnp; these were most likely initiated beneath a former mantle of regolith, which has subsequently been eroded and left behind such bedrock landforms of etch origin, as suggested by mueller and twidale (2002) for similar landforms identified in new mexico, usa. pedestals often have a characteristic basal concavity, above which are broader bedrock cappings, resembling a mushroom (twidale & campbell 1992). although such bedrock forms may have been initiated through original subsurface etching processes, once they have been exhumed, differential rates of weathering and erosion would have continued along existing joints and areas of locally weaker rock strength, particularly from induced fracturing (ollier 1978). the formation of these forms may also have been controlled by spatial dynamics such as distance from the ground surface and aspect. cavernous weathering cavernous weathering has been described as ‘differential weathering of a rock surface’ which represents ‘areas of material loss surrounded by intervening areas of stable rock surface (with minimal material loss)’ (turkington & phillips 2004:665). international research on cavernous weathering has focused on both microcavernous forms (e.g. negative spheroidal weathering forms [wagner 1913], dewholes [demek 1964] and honeycombs [e.g. mcbride & picard 2004; rodriguez-navarro, doehne & sebastian 1999]), and macrocavernous forms such as tafoni (e.g. martini 1978; matsukura & tanaka 2000; mottershead & pye 1994), alcoves (lamb et al. 2006) and rock overhangs (locally also known as rock shelters or caves). to this end, there has been limited discussion concerning the likely successive evolution of forms from honeycombs (embryonic tafoni) to tafoni to so-called rock shelters (‘mature tafoni’ or ‘tafoni caverns’as described by mellor, short & kirkby 1997). cavernous weathering is particularly prominent in crystalline rocks with a medium to coarse grain structure, including granites and sandstones (jennings 1968; mellor et al. 1997), and is thus active within the sandstones of the gghnp. 1. honeycombs: honeycombs are the smallest of the cavernous weathering types (only few tens of centimetres across), consisting of closely spaced pits which are bounded by thin raised lips or walls (mcbride & picard 2004). such weathering of microforms have previously been described as stone lattices, stone lace or fretting (mustoe 1982), but are more commonly also known as alveoli (e.g. benito, machado & sancho 1993; grisez 1960). in some instances honeycombs may develop within larger cavernous forms such as tafoni, but are said to be affected more strongly by lithological heterogeneities (martini 1978). in fact, groenewald (1986) observed honeycombs in the gghnp and similarly ascribed their formation to differential weathering associated with calcareous concretions. international research suggests that honeycombs are typically a product of solutional processes and the crystallisation of salt in the pits (e.g. lambert 1980; mcbride & picard 2004; young 1987), while case hardening by iron oxide or calcium carbonate offers greater resistance to weathering along the surrounding, less denuded walls (campbell 1999; mcbride & picard 2004). the spatial occurrence of honeycombs in the gghnp is likely controlled by local sandstone permeability, the extent of sandstone cementation and mineralogy. 2. tafoni, caverns, alcoves (rock shelters, caves): tafoni are relatively large (several metres across) cavernous weathering forms with overhanging hoods, arch-shaped entrances, concave inner walls, overhanging margins and relatively smooth, gently sloping, debris-covered floors (mcbride & picard 2004; mellor et al. 1997). the formation of tafoni has been ascribed to differential weathering, granular disintegration and flaking associated with microlithological and microclimatological differences along rock walls. meteoric water migration through megapores and associated dissolution and reprecipitation of calcareous and saline cements, crystallisation pressure, hydrolysis, hydration and/or diffusion, and wind deflation have also been noted as particular contributors (cooke, warren & goudie 1993; martini 1978; matsukura & tanaka 2000; mcbride & picard 2000; mellor et al. 1997; mottershead & pye 1994). however, water-based weathering processes may be temporally and spatially constrained at some tafoni sites, such as those reported from arid regions of antarctica, where thermal stress may take on a more important weathering function (hall & andré 2006). the gghnp hosts a variety of variably sized tafoni forms (ranging from only a few to several tens of cubic metres), which most likely represent various stages of tafoni evolution, as described by mellor et al. (1997). it would appear that several of the elongated rock shelters (caves) observed in the gghnp (primarily in clarens formation sandstone) formed through the amalgamation of tafoni caverns that are in places separated by thin rock walls. multiple vertical zones of tafoni and cavern development may occur within rock walls and are likely controlled by zonal lithological (rock hardness) and moisture differences (matsukura & tanaka 2000). the caverns are thought to enlarge in an upward and backward direction, thus potentially capturing subsurface water and causing water flow down the backwall (mellor et al. 1997), as is frequently observed at sites in the gghnp. as the tafoni enlarge over time, so the microclimate within the cavern is altered (primarily through greater contrasts in solar radiation levels across rock surfaces), thus influencing rock characteristics such as moisture and temperature. consequently, different weathering processes may dominate on different parts of tafoni, such as case hardening on outer roof surfaces and granular disintegration on inner cavern surfaces (mellor et al. 1997). another type of cavern occasionally observed in the gghnp region is the alcove. an alcove has been defined as ‘a steep-sided, arcuate cavity on the flank of a rock outcrop, typically produced by water erosion processes such as solution and spring sapping or seepage’ (nash 2000:14). thus, several of the so-called caverns or caves in the gghnp, and possibly even large tafoni which owe their origin to seepage and dissolution processes, might be classified as alcoves. water is often found to be seeping out from the rear of these features, producing black staining on the backwalls (figure 4c). however, as lamb et al. (2006) point out, the origin of alcoves remains unclear given that the ability of seepage to erode bedrock is not yet comprehensively understood, and also because other mechanisms might have contributed to such arcuate cavities. according to le roux (1978), caves in the gghnp are a consequence of exudation, whereby weathering processes are accelerated at the base of cliffs, in part owing to the presence of shale or clay-bearing sandstone layers. many of the caverns observed in the gghnp are located at the lower boundary of the clarens formation where it merges with the siltstones and shale beds of the elliot formation. here we observed that brittle fractures are common phenomena, possibly not so much due to weathering in the clay-like rocks, but rather due to mass failure of bedrock below the recess and beyond the dripline of water coming over the cliff (see also young et al. 2009). rock basins (weathering or solutional pits, gnammas) rock basins are relatively circular or oval and usually flat-floored and steep-sided bedrock depressions ranging from a few centimetres to several metres in diameter and depth (domínguez-villar et al. 2009) (figure 4d). rock basins have also been referred to as gnammas (domínguez-villar et al. 2009), weathering pits (alexandrowicz 1989; domínguez-villar 2006; goudie & migón 1997), potholes (chan et al. 2005; graham & wirth 2008; johnsson 1988), solution pans (dzulynski & kotarba 1979), opferkessel (hedges 1969), pias (vidal romaní 1989) and, locally, as rock tanks (king 1942) or weathering basins (cooks & pretorius 1987). rock basins may contain standing water for short durations after rainfall or for several weeks to months during wet periods or seasons. rock basins, which most typically develop in granite and sandstone, are understood to be polygenetic in origin (domínguez-villar et al. 2009; dzulynski & kotarba 1979). processes include chemical weathering associated with dissolution (domínguez-villar et al. 2009; twidale 1982; wray 1997) and mechanical erosion associated with, for instance, convective (circulatory) currents within the standing water (dzulynski & kotarba 1979), wash and deflation (alexandrowicz 1989; graham & wirth 2008), undercutting of the rock basin sides (goudie & migón 1997), and the plucking action of lichen thalli during the drying phase (moses & smith 1993). complex biofilms may accumulate at the base of rock basins; these are known to dissolve the cement between sandstone grains and also act as biological sealants to water infiltration (chan et al. 2005). cooks and pretorius (1987) have identified blue-green algae below the rock basins’ seasonal water lines in the gghnp and suggested that associated biochemical activity would enhance weathering there. solutional weathering is prominent along joints and bedding planes and may cause intense etching, particularly in quartz-rich sandstone (young et al. 2009). cooks and pretorius (1987) similarly observed rock basin development in the gghnp at sites where joints intersect at the surface. in addition, some of the pans have well-developed water level notches which are probably the result of solutional attack. rock basins in the gghnp are commonly 15 cm – 150 cm wide and approximately 5 cm – 30 cm deep. however, cooks and pretorius (1987) report features up to 16 m in length and 1 m in depth. examples of isolated deep pits (> 1 m deep) and exceptionally wide (several metres in diameter), shallow bedrock basins (also referred to locally as ‘tarns’) occur elsewhere on clarens formation sandstones, such as in the sehlabathebe national park, southern drakensberg. in some cases spillways develop at the lowest point of the basin rim where erosion eventually produces a confined water exit route, below which drainage microchannels usually develop. the overflow of water during summer is an effective mechanism to remove weathered products accumulating in the rock basins. less frequently, a staircase of rock basins with connected drainage microchannels may develop down sandstone surfaces, while in other cases closely spaced basins may eventually coalesce. rock basins may also be associated with incision into polygonally cracked surfaces (robinson & williams 1992) or with raised conical forms such as rock doughnuts (netoff & shroba 2001; young et al. 2009). rock doughnuts rock doughnuts were first described by blank (1951:822), who described them as ‘weather pits of various sizes on granite’ which are ‘surrounded by raised annular rims’. although rock doughnuts are relatively rare phenomena which have received little research attention, they have also been reported from a variety of sandstone terrains in australia (twidale & campbell 1998; young et al. 2009) and from south-eastern utah, usa (netoff & shroba 2001). rock doughnuts belong to a group of conical-shaped raised landforms with a central summit ‘pit’ or rock basin, examples of which were found at one locality in the gghnp (figures 3 and 5a). rock doughnuts ranging between 5 cm and several metres high have been reported in australia and the usa and have pit diameters ranging from approximately 20 cm to 10 m (netoff & shroba 2001; twidale & campbell 1998; young et al. 2009). the ones in the gghnp are typically 5 cm – 50 cm high and have pit diameters ranging from 15 cm to 60 cm. although reasons for the formation of rock doughnuts remain uncertain, twidale and campbell (1998) suggest that subsurface water held in the regolith was preferentially drained from zones adjacent to basins (doughnuts). the drained areas consequently had less moisture and thus weathered less rapidly than adjacent areas where more water was retained. polygonal cracking (tessellation or alligator cracking) polygonal fracture patterns consist mainly of pentagonal or hexagonal cracks meeting at ca 120° angles where they make tri-radial junctions on predominantly curved rock surface pavements (robinson & williams 1989, 1992; young et al. 2009). polygonal crack diameters may vary from 5 cm to more than 50 cm and in some instances micropolygonal cracking has been observed within larger tessellation plates (robinson & williams 1989). although understanding the origin of tessellation has caused some debate (young et al. 2009) and may differ between sites, it is a particularly common phenomenon in case-hardened rocks associated with regions of seasonal precipitation patterns (robinson & williams 1989, 1992; twidale & campbell 1993). some of the likely causative mechanisms suggested include shrinkage of silica gel due to changing thermal and/or moisture conditions on the rock surface (robinson & williams 1989, 1992). these include thermal changes that set up differential surface stress (branagan 1983; croll 2009), expansion to accommodate precipitating solutions of iron, and thermal stresses or geochemical changes caused by emplacement of dolerite dykes (velázquez et al. 2008). polygonal cracking is a common phenomenon in the gghnp and in most instances occurs on horizontal case-hardened rock pavements, but may also be observed on vertical exposures such as at the base of cliffs along the echo ravine hiking trail and at baboon ridge. as observed in the field, the cracks may penetrate beyond the ca 5-mm thick case-hardened surface and continue further into the subsurface. at the oribi drive vulture site and also at baboon ridge, well-developed pavements occur in sandstones close to dolerite dykes. less common are polygonal ‘boxwork’ structures where the joints are positive features, possibly due to silica induration along the joints, while the polygonal cells have been preferentially eroded to form small rock basins (figure 5b). figure 5: (a) a rock doughnut, (b) polygonal ‘boxwork’ structures, (c) convex polygonal structures and (d) lichen etching into clarens formation sandstone in the golden gate highlands national park. bedrock drainage microchannels (fluting, gutters, karren forms) bedrock drainage microchannels most frequently develop on limestone, dolomite and other soluble rock types. they are, however, also known to form on basaltic, granitic and sandstone surfaces as a result of sheet or channelised water flow across bedrock surfaces (forming rillenkarren or ’solution flutes’ and rinnenkarren or rundkarren, respectively). the formation of such structures is primarily due to dissolution and biokarstic processes and/or the pre-existence of joints or surface irregularities (crowther 1997; dunkerley 1979; fiol, fornós & ginés 1996; wray 1997). in some instances long drainage microchannels are interrupted by shallow rock basins. sandstone surfaces with ferruginous concretions are common to the gghnp and often lead to intricate surface drainage across bedrock, producing sharp-crested and highly fragmented surfaces. trittkarren are much rarer and lesser-known weathering and erosion surface forms and have been described as ‘small arcuate impressions’ on sloping bedrock (vincent 1983:205). although examples of trittkarren have been observed in the gghnp, their precise process origins remain poorly understood. however, bögli (1960) suggested that such forms may develop where solution rates are influenced by the slow dissolution of gaseous co2 into the surface water in contact with bedrock and subsequently, when the microforms are established, they initiate self-sustaining processes such as concentrated overland flow towards the centre of each riser lip (bögli 1980). twidale (1980) first described incised channelling into sandstone in the drakensberg and suggested it to be the product of subsurface flow beneath a soil cover before the sandstone had been exposed. however, it is now more generally recognised that such sandstone channels (up to 100 mm wide and 10 mm deep) represent selective channelised drainage routes across rock surfaces (robinson & williams 1992; wray 1997). polygonally cracked surfaces permit drainage along the cracks, consequently accelerating erosion and eventually leading to well-incised bedrock microchannels, which grow in diameter and depth in the downflow direction. observations at a site in the gghnp indicate that originally fairly flat polygonally cracked surfaces may evolve into a more dramatically undulating micro-relief of incised channels (mean depth = 7 cm; n = 20) that separate broad convex polygonal tops with an average diameter of 33 cm (n = 40) (figure 5c). lichens and weathering although it has previously been argued that lichens may protect rocks from enhanced weathering (danin & garty 1983), the more common view is that lichens promote rock weathering through chemical and mechanical processes which include metallic cation chelation, dissolution, swelling of organic and inorganic salts, and hyphal penetration, where expansion and contraction cause the detachment, incorporation and expulsion of rock flakes (chen et al. 2000; mccarroll & viles 1995). wessels et al. (1995) have measured high strain on clarens formation sandstone surfaces in the gghnp in association with increasing moisture uptake by some endolithic lichens, particularly at the thallus rim. in another study undertaken in the gghnp, wessels and wessels (1991) identified bagworm larvae constructing their bags around lichen-weathered quartz crystals and thus demonstrated that lichen-induced weathering may operate synergistically with other biogenic processes on rock surfaces. studies, including some from the gghnp, have indicated that some lichens may form circular or crescentic depressions bounded by microscarps on rock surfaces (wessels & schoeman 1988; wessels et al. 1995) (figure 5d) owing to the combined actions of chemical weathering by chelation and mechanical disintegration associated with the penetration and expansion of hyphae (cooks & otto 1990; mckechnie, grab & drennan 2007; moses & smith 1993). many of the flat or gently sloping sandstone pavements in the gghnp are extensively covered with circular patches of endolithic lichen thalli. when these thalli merge, they can produce remarkable flower-like patterns of pits in the rock surface (figure 5d). broadly similar forms have been recorded from the high atlas mountains in morocco (edwards et al. 2002). a survey undertaken on a rock wall in the gghnp found that up to 29% of the colonisable surface was covered by thalli (wessels & schoeman 1988). our own surveys supported this observation. the associated thalli microdepressions observed in the gghnp include compound (i.e. microscarps), coalescent and crescentic forms. lichenometric dating has become a common method for determining the relative age of exposed rock surfaces and has assisted in providing indications of glacial advances (bradwell 2004), the timing of debris flows (innes 1983), rates of fluvial downcutting (thompson & jones 1986) and past rockfall events (bull 1996), to name a few. surface age is traditionally determined by the single largest lichen, but other methods such as using the gradient of the lichen size–frequency distribution have also been used to produce well-calibrated dating curves (bradwell 2004). wessels and schoeman (1988) examined freshly quarried clarens formation sandstone blocks used in the construction of a stone wall in 1942 to determine rates of lichen-induced weathering in the gghnp. their results suggest a theoretical mean surface denudation rate of 9.6 mm per 100 years where endolithic lichens are active on rock surfaces. observations at the same site (the glen reenen graveyard) in 2009 showed that such lichen attack still continues. we measured 29 lichens (lecidea aff. sarcogynoides) on 16 blocks and found that the average width of the lichens was 98.28 mm and that they penetrated into the rock surface to an average maximum depth of 4.8 mm. the average maximum rate of lichen denudation in the 67 years since 1942 was thus 7.16 mm per 100 years. such results may prove valuable for any future attempt to determine the relative lichenometric-derived ages of mass movement phenomena in the gghnp. the depths of lichen denudation at glen reenen are not unique. for example, at a clarens formation sandstone at baboon ridge we measured 30 lecidea pits and found that the average maximum width was 127 mm and the average maximum depth was 5.8 mm. at red mountain we measured 20 lecidea pits in elliot formation sandstone and found that the comparable values were 65.25 mm and 3.3 mm, respectively. discussion and conclusion (back to top) why does the gghnp have such a rich diversity of macroand microscale sandstone phenomena? warke, mckinley and smith (2006) have experimentally demonstrated that even minor structural and mineralogical differences between sandstone samples can result in substantially different weathering behaviours and consequent rock morphological outcomes. the gghnp and foothills to the east of the great escarpment along kwazulu-natal and the eastern cape host a particularly dynamic assortment of sandstones, ranging from fine-grained mudstones and siltstones to coarse-grained quartzitic sandstones. given the relative abundance of feldspathic sandstones in the gghnp, many of the macroand microscale phenomena observed resemble those commonly recorded in granitic terrain. the presence of ferruginous chloritic mud and calcitic cement (eriksson 1981) has provided the catalyst to the wide assortment of landforms associated with dissolution, which more typically occur in limestone or dolomitic terrain. further, the rich diversity of the sandstone depositional history and concurrent neotectonics have provided for an equally complex spatial diversity of sandstone porosity, degree of lithification, and bedding and jointing characteristics, all of which contribute to sedimentary constructions and destructions. finally, neither the past nor present climate can be underestimated for its all-important role in developing geomorphic phenomena through weathering, erosion and deposition of earth materials. the larger landforms (i.e. cliffs, pillars, arches, etc.) discussed in this paper are the likely product of past geological processes and controls and climate-driven geomorphic processes operating at geological timescales, while the smaller microforms (i.e. honeycombs, rock basins, polygonal cracks, etc.) are likely consequences of more recent climates (i.e. from the last few thousand years), characterised in particular by strong seasonal temperature and moisture differences. considerable work has focused on the deterioration of sandstone cultural rock, particularly in the context of monuments that belong to world cultural heritage (mohammadi & krumbein 2008) and sandstone buildings (viles 2002). the rapid deterioration of sandstone containing rock art both locally (hall, meiklejohn & arocena 2007; meiklejohn 1997) and internationally (benito et al. 1993; walderhaug 1998) has also been a matter of concern in the gghnp. the primary geomorphic causes for rock art deterioration are granular disintegration and flaking associated with salt crystallisation, hydration, wetting and drying, and chemical alterations (benito et al. 1993; meiklejohn 1997; meiklejohn, hall & davis 2009). however, it should be recognised that for the most part, rock weathering is a natural process, and as pope, meierding and paradise (2002:224) convey: ‘geomorphic processes are integral in the creation of the “ruined” aesthetic’. notwithstanding this, rock art requires preservation against unnecessary accelerated weathering, particularly where anthropogenic contributing factors are part of the cause. finally, an improved understanding of the mechanisms producing terrestrial sandstone phenomena, such as those described for the gghnp, offers opportunities to better understand sedimentary and weathering processes on other planets such as mars (chan et al. 2008). some of the common sandstone phenomena identified in the gghnp and on mars include surface flaking (exfoliation), case hardening, honeycombs, tafoni, ‘blueberries’ and polygonal cracking. the water-based mechanism for blueberry formation on earth has been used as a terrestrial analogue for martian blueberry formation and, more importantly, to provide possible evidence for the past or present occurrence of fluids on mars (squyres & knoll 2005). we hope that this work has extended scientific knowledge on sandstone phenomena in the gghnp and thus provides a contribution to geoconservation and the regional geotourism industry through the provision of educational materials. acknowledgements (back to top) we thank south african national parks for permission to carry out research at the gghnp. particular thanks to the manager of the gghnp, johan taljaard, for his kind assistance and permission to use the photographic image of the natural stone bridge. s.g. acknowledges financial support from the wits university research committee. wendy job kindly assisted in the cartographic production. we also appreciate the valuable comments and suggestions by the referees. references (back to top) alexandrowicz, z., 1989, ‘evolution and weathering of pits on sandstone tors in the polish carpathians’, zeitschrift für geomorphologie 33, 275−289. benito, g., machado, m.j. & sancho, c., 1993, ‘sandstone weathering processes damaging prehistoric rock paintings at the albarracin cultural park, ne spain’, environmental geology 22, 71−79. doi:10.1007/bf00775287 blank, h.r., 1951, ‘rock doughnuts, a product of granite weathering’, american journal of science 249, 822−829. doi:10.2475/ajs.249.11.822 bögli, a., 1960, ‘kalkösung und karrenbildung [calcification and solution flute development]’, zeitschrift für geomorphologie suppl. 2, 4–21. bögli, a., 1980, karst hydrology and physical speleology, springer-verlag, berlin. bordy, e.m., hancox, p.j. & rubidge, b.s., 2004, ‘fluvial style variations in the late triassic – early jurassic elliot formation, main karoo basin, south africa’, journal of african earth sciences 38, 383−400. doi:10.1016/j.jafrearsci.2004.02.004 bradwell, t., 2004, ‘lichenometric dating in southeast iceland: the size-frequency approach’, geografiska annaler 86a, 31−41, doi:10.1111/j.0435-3676.2004.00211.x branagan, d.f., 1983, ‘tesselated pavements’, in r.w. young & g.c. nanson (eds.), aspects of australian sandstone landscapes, pp. 11−20, australian and new zealand geomorphology group, wollongong. büdel, b., weber, b., kühl, m., pfanz, h., sültemeyer, d. & wessels, d., 2004, ‘reshaping of sandstone surfaces by cryptoendolithic cyanobacteria: bioalkalization causes chemical weathering in arid landscapes’, geobiology 2, 261−268. doi:10.1111/j.1472-4677.2004.00040.x bull, w.b., 1996, ‘dating san andreas fault earthquakes with lichenometry’, geology 24, 111−114. doi:10.1130/0091-7613(1996)024<0111:dsafew>2.3.co;2 campbell, s.w., 1999, ‘chemical weathering associated with tafoni at papago park, central arizona’, earth surface processes and landforms 24, 271−278. doi:10.1002/(sici)1096-9837(199903)24:3<271::aid-esp969>3.3.co;2-k, doi:10.1002/(sici)1096-9837(199903)24:3<271::aid-esp969>3.0.co;2-t catuneanu, o., hancox, p.j. & rubidge, b.s., 1998, ‘reciprocal flexural behaviour and contrasting stratigraphies: a new basin development model for the karoo retroarc foreland system, south africa’, basin research 10, 417−439. doi:10.1046/j.1365-2117.1998.00078.x catuneanu, o., hancox, p.j., cairncross, b. & rubidge, b.s., 2002, ‘foredeep submarine fans and forebulge deltas: orogenic off-loading in the underfilled karoo basin’, journal of african earth sciences 35, 489−502. doi:10.1016/s0899-5362(02)00154-9 chan, m.a., beitler bowen, b., parry, w.t., ormö, j. & komatsu, g., 2005, ‘red rock and red planet diagenesis: comparisons of earth and mars concretions’, gsa today 15, 4−10. doi:10.1130/1052-5173(2005)15[4:rrarpd]2.0.co;2, doi:10.1130/1052-5173(2005)015[4:rrarpd]2.0.co;2 chan, m.a., yonkee, w.a., netoff, d.i., seiler, w.m. & ford, r.l., 2008, ‘polygonal cracks in bedrock on earth and mars: implications for weathering’, icarus 194, 65−71. doi:10.1016/j.icarus.2007.09.026 chen, j., blume, h.p. & beyer, l., 2000, ‘weathering of rocks induced by lichen colonization – a review’, catena 39, 121−146. doi:10.1016/s0341-8162(99)00085-5 cooke, r., warren, a. & goudie, a., 1993, desert geomorphology, ucl press, london. cooks, j. & pretorius, j.r., 1987, ‘weathering basins in the clarens formation sandstone, south africa’, south african journal of geology 90, 147−154. cooks, j. & otto, r., 1990, ‘the weathering effects of the lichen lecidea aff. sarcogynoides (koreb.) on magaliesberg quartzite’, earth surface processes and landforms 15, 491−500. doi:10.1002/esp.3290150602 croll, j.g.a., 2009, ‘possible role of thermal ratcheting in alligator cracking of asphalt pavements’, international journal of pavement engineering 10, 447−453. doi:10.1080/10298430902730547 crowther, j., 1997, ‘surface roughness and the evolution of karren forms at lluc, serra de tramuntana, mallorca’, zeitschrift für geomorphologie 41, 393−407. danin, a. & garty, j., 1983, ‘distribution of cyanobacteria and lichens on hillsides of the negev highlands and their impact on biogenic weathering’, zeitschrift für geomorphologie 27, 423−444. demek, j., 1964, ‘slope development in granite areas of bohemian massif (czechoslovakia)’, zeitschrift für geomorphologie suppl. 5, 83−106. domínguez-villar, d., 2006, ‘early formation of gnammas (weathering pits) in a recently glaciated area of torres del paine, southern patagonia (chile)’, geomorphology 76, 137−147. doi:10.1016/j.geomorph.2005.10.006 domínguez-villar, d., razola, l., carrasco, r.m., jennings, c.e. & pedraza, j., 2009, ‘weathering phases recorded by gnammas developed since last glaciation at serra da estrela, portugal’, quaternary research 72, 218−228. doi:10.1016/j.yqres.2009.05.004 dorn, r.i., 2004, ‘case hardening’, in a.s. goudie (ed.), encyclopedia of geomorphology, vol. 1, pp. 118−119, routledge, london. du preez, p.j. & bredenkamp, g.j., 1991, ‘vegetation classes of the southern and eastern orange free state (republic of south africa) and the highlands of lesotho’, navorsinge van die nasionale museum bloemfontein 7, 477−526. dunkerley, d.l., 1979, ‘the morphology and development of rillenkarren’, zeitschrift für geomorphologie 23, 332−348. dzulynski, s. & kotarba, a., 1979, ‘solution pans and their bearing on the development of pediments and tors in granite’, zeitschrift für geomorphologie 23, 172−191. edwards, h.g.m., holder, j.m., seaward, m.r.d. & robinson, d.a., 2002, ‘a raman spectroscopic study of lichen-assisted weathering of sandstone outcrops in the high atlas mountains, morocco’, journal of raman spectroscopy 33, 449−454. doi:10.1002/jrs.859 ericksson, p.g., 1981, ‘a palaeoenvironmental analysis of the clarens formation in the natal drakensberg’, transactions of the geological society of south africa 84, 7−17. ericksson, p.g., 1984, ‘a palaeoenvironmental analysis of the molteno formation in the natal drakensberg’, transactions of the geological society of south africa 87, 237−244. ericksson, p.g., 1985, ‘the depositional environment of the elliot formation in the natal drakensberg and north-east orange free state’, transactions of the geological society of south africa 88, 19−26. fairbridge, r.w. (ed.), 1968, the encyclopedia of geomorphology, reinhold, new york. fiol, l., fornós, j.j. & ginés, a., 1996, ‘effects of biokarstic processes on the development of solutional rillenkarren in limestone rocks’, earth surface processes and landforms 21, 447−452. doi:10.1002/(sici)1096-9837(199605)21:5<447::aid-esp607>3.0.co;2-x goudie, a.s. & migón, p., 1997, ‘weathering pits in the spitzkoppe area, central namib desert’, zeitschrift für geomorphologie 41, 417−444. goudie, a.s. & viles, h.a., 1997, salt weathering hazard, john wiley & sons, chichester. goudie, a.s., migón, p., allison, r.j. & rosser, n., 2002, ‘sandstone geomorphology of the al quwayra area of south jordan’, zeitschrift für geomorphologie 46, 365−490. graham, t.b. & wirth, d., 2008, ‘dispersal of large branchiopod cysts: potential movement by wind from potholes on the colorado plateau’, hydrobiologia 600, 17−27. doi:10.1007/s10750-007-9171-5 grisez, l., 1960, ‘alveolisation littorale de schistes metamorphiques [alveolisation of coastal metamorphic schists]’, révue de géomorphologie dynamic 11, 164−167. groenewald, g.h., 1986, ‘geology of the golden gate highlands national park’, koedoe 29, 165−181. hall, k. & andré, m.-f., 2006, ‘temperature observations in antarctic tafoni: implications for weathering, biological colonization, and tafoni formation’, antarctic science 18, 377−384. doi:10.1017/s0954102006000423 hall, k., marker, m.e. & le roux, j.s., 1991, ‘the significance of periglacial geomorphology in southern africa: a discussion’, south african geographer 18, 134−139. hall, k., meiklejohn, i. & arocena, j., 2007, ‘the thermal responses of rock art pigments: implications for rock art weathering in southern africa’, geomorphology 91, 132−145. doi:10.1016/j.geomorph.2007.02.002 hedges, j., 1969, ‘opferkessel [rock basins],, zeitschrift für geomorphologie 13, 22−55. henderson, z., scott, l., rossouw, l. & jacobs, z., 2006, ‘dating, paleoenvironments, and archaeology: a progress report on the sunnyside 1 site, clarens, south africa’, archaeological papers of the american anthropological association 16, 139−149. doi:10.1525/ap3a.2006.16.1.139 holmer, b., 1998, ‘flaking by insolation drying and salt weathering on the swedish west coast’, zeitschrift für geomorphologie 42, 39−55. holzförster, f., 2007, ‘lithology and depositional environments of lower jurassic clarens formation in the eastern cape, south africa’, south african journal of geology 110, 543−560. doi:10.2113/gssajg.110.4.543 innes, j.l., 1983, ‘size-frequency distributions as a lichenometric technique: an assessment’, arctic and alpine research 15, 285−294. doi:10.2307/1550825 irwin, r.p., fortezzo, c.m., tooth, s.e., howard, a.d., zimbelman, j.r., barnhart, c.j., et al., 2009, ‘origin of theater-headed tributaries to escalalante and glen canyons, utah’, [poster] presentation at the 40th lunar and planetary science conference (lunar and planetary science xl), the woodlands, texas, 23−27th march. jennings, j.n., 1968, ‘tafoni’, in r.w. fairbridge (ed.), the encyclopedia of geomorphology, p. 1104, reinhold, new york. johnsson, g., 1988, ‘potholes-glacial and non-glacial cavities’, geografiska annaler 70a, 333−336. doi:10.2307/521266 kay, c., bredenkamp, g.c. & theron, g.k., 1993, ‘the plant communities of the golden gate highlands national park in the north-eastern orange free state’, south african journal of botany 59, 442−449. king, l.c., 1942, south african scenery, oliver and boyd, edinburgh. kitching, j.w. & raath, m.a., 1984, ‘fossils from the elliot and clarens formations (karoo sequence) of the northeastern cape, orange free state and lesotho, and a suggested biozonation based on tetrapods’, palaeoentologia africana 25, 111−125. lamb, m.p., howard, a.d., johnson, j., whipple, k.x., dietrich, w.e. & perron, j.t., 2006, ‘can springs cut canyons into rock?’, journal of geophysical research 111, eo7002. doi:10.1029/2005je002663 lambert, d., 1980, ‘the influence of ground water salts in the formation of sandstone shelters near gosford, nsw’, institute for conservation of cultural materials bulletin 6, 29−34. le roux, j.s., 1978, ‘die ontstaan van grotte en holtes in die kranse van die clarenssandsteenformasie in die golden gategebied [formation of caves and hollows in the cliffs of the clarens sandstone formation in the golden gate area]’, south african geographer 61, 23−29. le roux, j.s. & marker, m.e., 1990, ‘discussion of periglacial geomorphology at golden gate highlands national park, orange free state, south africa’, south african geographer 17, 129−131. leasure, v.l., kind, t.c. & busby, m.r., 2007, ‘formation of apex natural arch, kentucky’, journal of the kentucky academy of science 68, 96−101. doi:10.3101/1098-7096(2007)68[96:foanaa]2.0.co;2 mahaney, w.c., milner, m.w., netoff, d., dohn, j., kalm, v., krinsley, d., et al., 2008, ‘blueberries on earth and mars: correlations between concretions in navajo sandstone and terra meridiani on mars’, presentation at the american geophysical union fall meeting (abstract no. p33b-1440), san francisco, 15–19th december. marker, m.e., 1989, ‘periglacial geomorphology at golden gate highlands national park: a note on its fieldwork potential’, south african geographer 16, 147−153. marker, m.e., 1990, ‘nivation evidence from a north-facing slope? golden gate, eastern orange free state’, south african geographical journal 72, 15−18. martini, i.p., 1978, ‘tafoni weathering, with examples from tuscany, italy’, zeitschrift für geomorphologie 22, 44–67. matsukura, y. & tanaka, y., 2000, ‘effect of rock hardness and moisture content on tafoni weathering in the granite of mount doeg-sung, korea’, geografiska annaler 82a, 59−67. doi:10.1111/j.0435-3676.2000.00112.x mcbride, e.f. & picard, m.d., 2000, ‘origin and development of tafoni in tunnel spring tuff (oligocene), crystal peak, utah’, earth surface processes and landforms 25, 869−879. doi:10.1002/1096-9837(200008)25:8<869::aid-esp104>3.0.co;2-f mcbride, e.f. & picard, m.d., 2004, ‘origin of honeycombs and related weathering forms in oligocene macigno sandstone, tuscan coast near livorno, italy’, earth surface processes and landforms 29, 713−735. doi:10.1002/esp.1065 mccarroll, d. & viles, h., 1995, ‘rock-weathering by the lichen lecidea auriculata in an arctic alpine environment’, earth surface processes and landforms 20, 199−206. doi:10.1002/esp.3290200302 mckechnie, c., grab, s. & drennan, g., 2007, ‘documenting lichen-induced mechanical weathering of quartzitic sandstone at kaapsehoop, mpumalanga’, south african journal of science 103, 117−120. meiklejohn, k.i., 1997, ‘the role of moisture in the weathering of the clarens formation of the kwazulu-natal drakensberg: implications for the preservation of indigenous rock art’, south african geographical journal 79 (special issue), 199−206. meiklejohn, k.i., hall, k. & davis, j.k., 2009, ‘weathering of rock art at two sites in the kwazulu-natal drakensberg, southern africa’, journal of archaeological science 36, 973−979. doi:10.1016/j.jas.2008.11.020 mellor, a., short, j. & kirkby, s.j., 1997, ‘tafoni in the el chorro area, andalucia, southern spain’, earth surface processes and landforms 22, 817−833. doi:10.1002/(sici)1096-9837(199709)22:9<817::aid-esp768>3.0.co;2-t. mohammadi, p. & krumbein, w.e., 2008, ‘biodeterioration of ancient stone materials from the persepolis monuments (iran)’, aerobiologia 24, 27−33. doi:10.1007/s10453-007-9079-6 moon, b.p., 1991, ‘the significance of rectilinear bedrock slopes’, south african journal of science 87, 208−210. moon, b.p. & munro-perry, p.m., 1988, ‘slope development on the clarens sandstone formation in the northeastern orange free state’, the south african geographical journal 70, 57−68. moses, c.a. & smith, b.j., 1993, ‘a note on the role of the lichen collema auriforma in solution basin development on a carboniferous limestone substrate’, earth surface processes and landforms 18, 363−368. doi:10.1002/esp.3290180405 mottershead, d.n. & pye, k., 1994, ‘tafoni on coastal slopes, south devon, uk’, earth surface processes and landforms 19, 543−563. doi:10.1002/esp.3290190607 mueller, j.e. & twidale, c.r., 2002, ‘geomorphic development of the giants of the mimbres, grant county, new mexico’, new mexico geology 24, 39−48. mustoe, g.e., 1982, ‘the origin of honeycomb weathering’, bulletin of the geological society of america 93, 108−115. doi:10.1130/0016-7606(1982)93<108:toohw>2.0.co;2 nash, d.j., 2000, ‘alcoves’, in d.s.g. thomas & a. goudie (eds.), the dictionary of physical geography, 3rd edn., p. 14, blackwell, oxford. netoff, d.i. & shroba, r.r., 2001, ‘conical sandstone landforms cored with clastic pipes in glen canyon national recreation area, southeastern utah’, geomorphology 39, 99−110. doi:10.1016/s0169-555x(00)00096-9 nicol, i.g., 1973, ‘land forms in the little caledon valley, orange free state’, south african geographical journal 55, 56−68. ollier, c.d., 1978, ‘induced fracture and granite landforms’, zeitschrift für geomorphologie 22, 249−257. paradise, t.r., 2002, ‘sandstone weathering and aspect in petra, jordan’, zeitschrift für geomorphologie 46, 1−17. partridge, t.c. & maud, r.r., 1987, ‘geomorphic evolution of southern africa since the mesozoic’, south african journal of geology 90, 179−208. pope, g.a., meierding, t.c. & paradise, t.r., 2002, ‘geomorphology’s role in the study of weathering of cultural stone’, geomorphology 47, 211−225. doi:10.1016/s0169-555x(02)00098-3 reimold, w.u., whitfield, g. & wallmach, t., 2005, ‘geotourism potential of southern africa’, in r.k. dowling & d. newsome (eds.), geotourism: sustainability, impacts and management, pp. 40−62, elsevier, oxford. robinson, d.a. & williams, r.b.g., 1987, ‘surface crusting of sandstones in southern england and northern france’, in v. gardiner (ed.), international geomorphology 1986, vol. 2, pp. 623−635, wiley, chichester. robinson, d.a. & williams, r.b.g., 1989, ‘polygonal cracking of sandstone at fontainebleau, france’, zeitschrift für geomorphologie 33, 59−72. robinson, d.a. & williams, r.b.g., 1992, ‘sandstone weathering in the high atlas, morocco’, zeitschrift für geomorphologie 36, 413−429. robinson, d.a. & williams, r.b.g., 1994, ‘sandstone weathering and landforms in britain and europe’, in d.a. robinson & r.b.g. williams (eds.), rock weathering and landform evolution, pp. 371−392, john wiley and sons, chichester. rodriguez-navarro, c., doehne, e. & sebastian, e., 1999, ‘origins of honeycomb weathering: role of salts and wind’, geological society of america bulletin 111, 1250−1255. doi:10.1130/0016-7606(1999)111<1250:oohwtr>2.3.co;2 rowsell, d.m. & de swardt, a.m.j., 1976, ‘diagenesis in cape and karoo sediments, south africa and its bearing on their hydrocarbon potential’, transactions of the geological society of south africa 85, 203−210. sanparks, 2004, golden gate highlands national park, viewed 12 march 2007, from http://www.sanparks.co.za/parks/golden_gate/ smith, r.m.h., turner, b.r., hancox, p.j., rubidge, b.r. & catuneanu, o. (eds.), 1998, trans-karoo ii: 100 million years of changing terrestrial environments in the main karoo basin. guidebook, gondwana-10 international conference, university of cape town, cape town. spies, j.j., 1969, ‘die geologiese en geomorfologiese geskiedenis van golden gate hoogland nasionale park [the geological and geomorphological history of the golden gate highlands national park]’, koedoe 12, 184−198. squyres, s.w. & knoll, a.h., 2005, ‘sedimentary rocks at meridiani planum: origin, diagenesis, and implications for life on mars’, earth and planetary science letters 240, 1−10. doi:10.1016/j.epsl.2005.09.038 svensen, h., jamtveit, b., planke, s. & chevallier, l., 2006, ‘structure and evolution of hydrothermal vent complexes in the karoo basin, south africa’, journal of the geological society london 163, 671−682. doi:10.1144/1144-764905-037 thompson, a. & jones, a., 1986, ‘rates and causes of proglacial river terrace formation in southeast iceland: an application of lichenometric dating techniques’, boreas 15, 231−246. doi:10.1111/j.1502-3885.1986.tb00928.x turkington, a.v. & phillips, j.d., 2004, ‘cavernous weathering, dynamical instability and self-organization’, earth surface processes and landforms 29, 665−675. doi:10.1002/esp.1060 twidale, c.r., 1980, ‘origin of minor sandstone landforms’, erdkunde 34, 219−224. doi:10.3112/erdkunde.1980.03.07 twidale, c.r., 1982, granite landforms, elsevier, new york. twidale, c.r. & campbell, e.m., 1992, ‘on the origin of pedestal rocks’, zeitschrift für geomorphologie 36, 1−13. twidale, c.r. & campbell, e.m., 1993, ‘fractures: a double edged sword: a note on fracture density and its importance’, zeitschrift für geomorphologie 37, 459−475. twidale, c.r. & campbell, e.m., 1998, ‘development of a basin, doughnut and font assemblage on a sandstone coast, western eyre peninsula, south australia’, journal of coastal research 14, 1385−1394. twidale, c.r. & vidal romaní, j.r., 2005, landforms and geology of granite terrains, balkema, leiden. doi:10.1201/9781439833704 velázquez, a.f., giannini, p.c.f., riccomini, c., sallun, a.e.m., hachiro, j. & gomez, c. de b., 2008, ‘columnar joints in the patiño formation sandstones, eastern paraguay: a dynamic interaction between dyke intrusion, quartz dissolution and coolinginduced fractures’, episodes 31, 302−308. vidal romaní, j.r., 1989, ‘geomorfologia granitica en galicia (nw españa) [granitic geomorphology in galicia (nw spain)]’, laboratorio xeolóxico de laxe (coruña) 13, 89−163. viles, h.a., 2002, ‘implications of future climate change for stone deterioration’, in s. siegesund, t.n. weiss and a. vollbrecht (eds.), natural stone, weathering phenomena, conservation strategies and case studies, pp. 407−418, geological society, bath. viles, h.a. & goudie, a.s., 2004, ‘biofilms and case hardening on sandstones from al-quwayra, jordan’, earth surface processes and landforms 29, 1473−1485. doi:10.1002/esp.1134 vincent, p.j., 1983, ‘the morphology and morphometry of some arctic trittkarren’, zeitschrift für geomorphologie 27, 205−222. wagner, p.a., 1913, ‘negative spheroidal weathering and jointing in a granite of southern rhodesia’, transactions of the geological society of south africa 15, 155−163. walderhaug, o., 1998, ‘chemical weathering at rock art sites in western norway: which mechanisms are active and how can they be retarded?’, journal of archaeological science 25, 789−800. doi:10.1006/jasc.1997.0224 warke, p.a., mckinley, j. & smith, b.j., 2006, ‘variable weathering response in sandstone: factors controlling decay sequences’, earth surface processes and landforms 31, 715−735. doi:10.1002/esp.1284 wessels, d. & büdel, b., 1995, ‘epilithic and cryptoendolithic cyanobacteria of clarens sandstone cliffs in the golden g ate highlands national park, south africa’, botanica acta 108, 220−226. wessels, d. & schoeman, p., 1988, ‘mechanism and rate of weathering of clarens sandstone by an endolithic lichen’, south african journal of science 84, 274−277. wessels, d. & wessels, l., 1991, ‘erosion of biogenically weathered clarens sandstone by lichenophageous bagworm larvae (lepidoptera: psychidae)’, lichenologist 23, 283−291. wessels, d. & wessels, l., 1995, ‘biogenic weathering and microclimate of clarens sandstone in south africa’, cryptogamic botany 5, 288−298. wessels, d., venter, d., wessels, w. & wessels, l., 1995, ‘experimental strain analysis of clarens sandstone colonised by endolithic lichens’, koedoe 38, 35−47. wray, r.a.l., 1997, ‘a global review on solutional weathering forms on quartz sandstone’, earth science reviews 42, 137−160. doi:10.1016/s0012-8252(96)00056-6 young, a.r.m., 1987, ‘salt as an agent in the development of cavernous weathering’, geology 15, 962−966. doi:10.1130/0091-7613(1987)15<962:saaait>2.0.co;2 young, r.w., wray, r.a.l. &. young, a.r.m., 2009, sandstone landforms, cambridge university press, cambridge. filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 hooimeijer.qxd the diet of kudus in a mopane dominated area, south africa j.f. hooimeijer, f.a. jansen, w.f. de boer, d. wessels, c. van der waal, c.b de jong, n.d. otto and l. knoop hooimeijer, j.f., f.a. jansen, w.f. de boer, d. wessels, c. van der waal, c.b de jong, n.d. otto and l. knoop. 2005. the diet of kudus in a mopane dominated area, south africa. koedoe 48(2): 93-102. pretoria. issn 0075-6458. the composition of the plant species eaten by kudu (tragelaphus strepsiceros) determines the diet quality, which impacts on kudu condition and mortality levels. the yearround diet composition of kudus in the limpopo province, a mopane (colophospermum mopane) dominated area, was determined by faecal analysis. the most important dietary plant species were colophospermum mopane, grewia bicolor, terminalia prunioides, tinnea rhodesiana, boscia albitrunca and combretum apiculatum, with c. mopane comprising on average 39.2 % of diet per month. small amounts of herbs, grasses and seeds made up the remaining part of the diet. the contribution of c. mopane in the diet was negatively correlated with precipitation. colophospermum mopane was consumed, irrespective of its high condensed tannin load (5.2–9.8 % dw) for the majority of the months. no seasonally significant differences were detected for modelled kudu diet crude protein, tannin or phenol concentrations. colophospermum mopane showed significant seasonal differences with lowest values of protein, tannin and phenols in the late wet season. surprisingly, crude protein concentrations were positively correlated with high levels of tannins and phenols for c. mopane. the diet of kudus comprised of significantly more species during the wet season compared to the dry season. diet diversification, instead of protein maximization, seems a potential tool to satisfy protein requirements while reducing potential toxic effects associated with a high intake of secondary compounds. a significant positive correlation was therefore detected between the tannin concentration of c. mopane leaves and the number of plant species in the diet. key words: bushveld, condition, condensed tannin, crude protein, preference, habitat use. j.f. hooimeijer, f.a. jansen, w.f. de boer (fred.deboer@wur.nl), c. van der waal, c.b de jong, n.d. otto and l. knoop, resource ecology group, wageningen university, bornsesteeg 69, 6708 pd wageningen, the netherlands; d. wessels, university of limpopo, turfloop campus, private bag x1106, sovenga, 0727 republic of south africa. issn 0075-6458 93 koedoe 48/2 (2005) introduction due to large-scale kudu mortalities during the drought of 2002, kudu populations decreased critically in different nature areas and game ranches in northern south africa. mass mortality among kudus frequently occurs, and from 1981–1986 kudu mortalities were reported from numerous ranches in the dry winter months between july and september (van hoven 1991). kudu population crashes commonly occur in game fenced areas in the arid sweet bushveld of the limpopo province (van der waal & smit 2001). previous research (owen-smith 1982; owen-smith & cooper 1989) showed that the dietary quality and quantity of the vegetation in kudu habitats can differ strongly per season. this variation is mainly related to rainfall and consequently forage availability. owen-smith & cooper (1989) showed that food quality and availability decrease in the dry season (may-october), and that the estimated metabolisable energy intake falls below the nutritional requirements of the kudu during this period. the nutritional gain can be reduced by the amount of secondary plant compounds like poly-phenols which hooimeijer.qxd 2006/01/10 04:35 pm page 93 includes condensed tannins. diet quality in deciduous savannas is expected to differ significantly between the wet and the dry season, and might therefore offer an explanation for the observed mass mortality kudus suffered, rather than simple lack of food. the first hypothesis that will be tested in this study is that kudu diet quality (measured by crude protein, tannin, and phenol concentrations) is lower in the dry season compared to the wet season; do kudus maximise protein or minimise secondary compounds? the study area is situated in the mopani veld (acocks 1988), of the northern part of the limpopo province. mopani veld is generally regarded as valuable browse (walker 1980). colophospermum mopane, a winter deciduous tree species, has a variable leaf carriage period that may continue through the dry season into the next wet season. due to its dominance, c. mopane is expected to play an important role in the kudu diet. colophospermum mopane is known for its antifeedants, like polyphenolic compounds, e.g. condensed tannins (macala et al. 1992). tannins bind with dietary, enzymatic and microbial protein to form insoluble complexes that are not degraded in the rumen, resulting in a reduced digestibility and intake. information on the fate of tannin-protein complexes postrumen varies and kidney, liver and gastrointestinal tract damage in animals consuming tanniferous forage has been reported (bailey 1978). cooper & owen-smith (1985) found that plant species with condensed tannin content of > 5 % leaf dry weight are preferably avoided by kudu in a non-mopane area. van hoven (1991) investigated kudu mortalities during winter in savanna areas of south africa, covering drought conditions, and revealed significant correlations between kudu mortality, kudu density, and tannin content of the browse. it is therefore expected that in the rainy season, when forage availability is high, deciduous c. mopane is avoided by kudus due to its high tannin content. the high forage availability during the rainy season enables the kudu to select for high quality alternative browse. the kudu diet composition is therefore expected to consist only of high quality species. subsequently, species with a tannin content >5 % are not expected in the rainy season diet. rainfall influences forage availability, and diet choices, and thereby stocking rates (bothma et al. 2004). due to the high availability of c. mopane during the dry season, we hypothesise that the contribution of c. mopane to the kudu diet increases in the dry season, resulting in a negative relation between rainfall and percentage c. mopane material in the kudu faeces. in this study, the year-round diet of kudus in a mopane (colophospermum mopane) dominated area is analysed through faecal analysis. this microhistological examination of herbivore droppings provides an estimate of the ingested biomass per plant taxon (stewart 1967; sparks & malechek 1968; cid & brizuela 1990; bartolome et al. 1995). to this end, epidermis fragments of ingested plants in the faeces are compared to photomicrographs of epidermis fragments on reference slides. this is possible because the plant cuticle, an indigestible layer covering the epidermis, bears a specific pattern of underlying epidermal cells and hairs along with structures of its own (stace 1965). this pattern can be identified down to genus or species level, even after passage through the herbivore gut. study area the study was conducted on the messina experimental farm, situated along the limpopo river (22º12's and 29º50'e), within the central zone of the limpopo belt. the study area covers 6991 ha, and is divided by a game-proof fence into a northern game section (4605 ha) and a southern cattle section (2386 ha), with kudu occurring in both sections. the mean annual rainfall is 357 mm (measured over a 66-year period from 1927/1928 to 1993/1994). the coefficient of variation for the total annual rainfall is 36 %, indicating a high frequency of droughts. of the total annual rainfall, 75 % is recorded during the period november–march. the mean daily maximum temperature varies from 25 ºc in july to 34 ºc in january; winkoedoe 48/2 (2005) 94 issn 0075-6458 hooimeijer.qxd 2006/01/10 04:35 pm page 94 ter temperatures can be regarded as moderate (dekker & van rooyen 1995). the study area falls within the northern block of the mopani veld or colophospermum mopane-acacia nigrescens savanna type, where c. mopane is the dominant tree species (dekker & van rooyen 1995). the vegetation in the study area is relatively homogeneous. in total, 183 plant species were recorded in eight distinct plant communities (dekker & van rooyen 1995). the browse availability on the messina experimental farm was determined by dekker & smit (1996) in both the northern and southern section. the total leaf dm ranged between 1224 kg/ha and 2672 kg/ha (dekker & smit 1996). colophospermum mopane contributed substantially to the total leaf dm in all communities. the kudu population size in the game section, determined by a helicopter game count in 2002, approached a hundred. no major off-takes or mortalities occurred since then (cornelis van der waal pers. comm. 2005). materials and methods faecal analyses the leaves of over a 100 species of potential food plants occurring on the messina experimental farm were sampled for a reference collection. pieces of relevant parts (foliage, seeds and fruits) were cleaned in household bleach overnight, washed in water, and fragments of epidermis stripped off and mounted in glycerol. photographs of these slides were used to identify the fragments of cuticle observed in samples of kudu faeces. the magnification ranged from 200–400 times, depending on cell, hair or stomata size of the epidermis fragment. the cuticles on mature green plant parts show the pattern of the epidermis cells they cover. very young undeveloped cuticles present in the dung samples may show no imprints of the epidermis cells in which case no identification was possible. also, soft cuticles may get crinkled beyond recognition. however, often hairs, glands or specific patterns were present on the cuticle itself which allowed identification of a given species. digestibility of plant parts as such bore no relation to recognition of cuticle or epidermis fragments. due to its chemical composition, plant cuticle is indigestible in any animal’s guts (stace 1965). this applies to the leaves of trees and grasses as much as to forbs. the faecal analysis was carried out on monthly collected mixed faecal samples of kudu droppings from june 2003 to may 2004. individual pellet groups are not independent samples of the diet of a single animal (lewis 1994); the availability of food items changes with increasing consumption, thereby influencing diet composition. also, herbivores vary their choice of food plants between meals in order to maintain a varied diet, and to limit consumption of secondary chemicals (freeland 1991, mcarthur and others 1991); mcarthur et al. 1991). each mixed sample consisted of three or more sub-samples, which were randomly collected in the study area. every sub-sample consisted of five fresh (< 24 hours old) kudu pellets. all collected sub-samples from the same month were pooled and thoroughly mixed. the mixed pellet samples were heated under pressure to 115 ºc in water for ± 2 hours, and left to soak overnight. a sub-sample was washed in a blender, and strained over a plankton sieve, then preserved in 70 % ethanol. the samples were analysed by using a microscope, and at least 100 cuticle or epidermis fragments in each sample were identified by comparison with the reference collection. the surface area of individual fragments was estimated by counting the obstructed grid cells (0.01 mm² per cell) in the microscopic field of view (de jong et al. 2004). the abundance of each species was calculated as a percentage of the total area of the fragments measured (de jong et al. 2004). encountered grass species in the faeces were placed into one category as the amount of consumed grasses was relatively low and their nutritional value assumed to be similar. diet quality foliage samples of the most important dietary plant species (n = 5; comprising 66 % of the kudu diet) were collected in the field for quality analysis by the university of limpopo from december 2002 till june 2004. seven fresh leaves were randomly collected at kudu browse height from the canopies of 40 randomly selected trees, in each sample cycle of 55 days. the bias of this hand-sampling method is unknown. samples were transported in plastic bags to the laboratory in a cooler box where they were dried in plant presses in the shade at room temperature (mueller-harvey 2001; hagerman 2002). the dried leaves were thoroughly mixed, and a pooled sample of 10 g per treatment per sample cycle (55 days) was sent to the botany department, universiissn 0075-6458 95 koedoe 48/2 (2005) hooimeijer.qxd 2006/01/10 04:35 pm page 95 ty of cape town, for the determination of total phenolics, condensed tannins and crude protein. all parameter values were measured in %dw, phenol content was based on gae (gallic acid equivalents) and condensed tannin content on ste (sorghum tannin equivalents) as described by hagerman (2002). additional nutritional data (n = 5; comprising 24 % of the kudu diet) were obtained from other sources (aganga & adolga-bessa 1999; aganga & mosase 2001; fustenburg & van hoven 1994; lagendijk 2003) or retrieved from the animal feed recourses information system (afris) (fao 2004). dietary species were classified into four groups: deciduous woody, evergreen woody, forbs and others. a monthly class average per nutritional parameter was used for species for which no nutritional data was available (9 % of the kudu diet). the diet quality, in terms of condensed tannins crude protein, and total phenolics was calculated using ∑(xi * perci) with xi as the bi-monthly average value of the quality parameter (condensed tannin, crude protein, or phenol) per plant species i and perci as the mean monthly percentage of occurrence of a plant species i in the diet, similar to owen-smith & cooper (1989). climatic data were obtained from the south african weather service which were measured at the macuville weather station (no. 0809706x), situated on the messina experimental farm. statistical analysis anovas were carried out using as dependent variables, the mean values recorded per month for species richness (averaged from the different samples) or the monthly mean diet quality parameters. as dependent variables did not violate basic assumption for normality (kolmogorov-smirnov test) and equality of variances (levene’s test), anovas were carried out to test for seasonal difference, using spss (v12), followed by a tukey multiple comparison test (zar 1984). results kudu diet: species composition the fragments of 20 dicotyledonous species contributed 82 % of the analysed kudu faecal samples and, together with the grass species, comprised 96.6 % of a full year’s samples (table 1). the remaining 3.4 % consisted of species which were only encountered once and were therefore not included in the diet quality analyses. colophospermum mopane was the most prominent in the faecal samples; it was consumed each month in relatively large amounts, with an annual average koedoe 48/2 (2005) 96 issn 0075-6458 fig. 1. seasonal differences of kudu diet composition for the main (annual > 1.9 %) dietary species. σ hooimeijer.qxd 2006/01/10 04:35 pm page 96 issn 0075-6458 97 koedoe 48/2 (2005) ta bl e 1 a nn ua l p er ce nt ag e of s pe ci es o cc ur re nc e in th e ku du d ie t a nd th ei r av er ag e se as on al n ut ri tio na l c om po ne nt v al ue (e w = e ar ly w et ; lw = la te w et ; e d = e ar ly d ry , l d = la te d ry ) c la ss if ic at io n sp ec ie s % a nn ua l d ie t cr ud e pr ot ei n % d w co nd en se d ta nn in % d w ph en ol % d w e w lw e d l d e w lw e d l d e w lw e d l d d ec id uo us a ca ci a ni gr es ce ns 0. 9 13 .3 * 12 .9 * 11 .5 * 12 .0 * 3. 3* * 3. 3* * 3. 3* * 3. 3* * 10 .3 * 9. 5* 9. 8* 9. 6* w oo dy a ca ci a to rt ili s 1. 9 11 .6 ** 11 .6 ** 11 .6 ** 11 .6 ** 3. 8* * 4. 6* * 5. 0* * 4. 5* * 2. 6* * 3. 0* * 3. 3* * 3. 0* * c ol op ho sp er m um m op an e 39 .2 11 .8 12 .3 9. 9 7. 1 9. 4 7. 7 7. 4 4. 3 8. 2 6. 4 6. 7 5. 3 c om br et um a pi cu la tu m 4. 0 13 .2 12 .6 9. 6 11 .8 0. 9 0. 6 0. 5 0. 7 14 .3 13 .1 13 .6 13 .7 c om br et um im be rb e 2. 0 14 .2 ** 14 .2 ** 14 .2 ** 14 .2 ** 3. 1* * 3. 1* * 3. 1* * 3. 1* * 13 .3 ** 13 .3 ** 13 .3 ** 13 .3 ** c om br et um m os sa m bi ce ns e 0. 6 13 .3 * 12 .9 * 11 .5 * 12 .0 * 3. 6* 3. 5* 3. 5* 3. 0 * 10 .3 * 9. 5* 9. 8* 9. 6* g ar de ni a re si ni flu a 1. 3 13 .3 * 12 .9 * 11 .5 * 12 .0 * 3. 6* 3. 5* 3. 5* 3. 0 * 10 .3 * 9. 5* 9. 8* 9. 6* g re w ia b ic ol or 9. 4 16 .2 14 .5 12 .8 14 .5 4. 8 5. 2 5. 1 5. 0 3. 9 3. 5 3. 7 3. 7 sc le ro ca ry a bi rr ea 0. 9 13 .3 * 12 .9 * 11 .5 * 12 .0 * 3. 6* 3. 5* 3. 5* 3. 0 * 10 .3 * 9. 5* 9. 8* 9. 6* te rm in al ia p ru ni oi de s 7. 8 11 .6 11 .0 8. 2 10 .3 3. 6* 3. 5* 3. 5* 3. 0 * 19 .8 17 .9 18 .2 18 .6 ti nn ea r ho de si an a 6. 5 13 .3 * 12 .9 * 11 .5 * 12 .0 * 3. 6* 3. 5* 3. 5* 3. 0 * 10 .3 * 9. 5* 9. 8* 9. 6* zi zi ph us m uc ro na ta 0. 8 14 .3 ** 14 .3 ** 14 .3 ** 14 .3 ** 0. 2* * 0. 2* * 0. 2* * 0. 2* * 10 .3 * 9. 5* 9. 8* 9. 6* e ve rg re en b os ci a al bi tr un ca 5. 1 15 .6 14 .2 14 .6 12 .8 0. 4 0. 4 0. 4 0. 4 0. 6 0. 6 1. 0 1. 0 sc ho tia b ra ch yp et al a 0. 7 15 .6 * 14 .2 * 14 .6 * 12 .8 * 10 .3 ** 10 .3 ** 10 .3 ** 10 .3 ** 0. 6* 0. 6* 1. 0* 1. 0* fo rb s h ib is cu s m ic ra nt hu s 0. 9 n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. in di go fe ra h et er ot ri ch a 0. 7 n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. o th er s k yl lin ga a lb a 1. 3 n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. sc le ro ca ry a bi rr ea f ru it 1. 9 6. 2* * 6. 2* * 6. 2* * 6. 2* * 2. 6* * 2. 6* * 2. 6* * 2. 6* * n. a. n. a. n. a. n. a. g re w ia sp p. s ee d 5. 1 19 .0 ** 19 .0 ** 19 .0 ** 19 .0 ** n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. u nk no w n 1 1. 5 n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. u nk no w n 5 1. 2 n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. g ra ss s pe ci es ( al l) 2. 9 n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. n. a. *c la ss a ve ra ge ** d at a re tr ie ve d fr om li te ra tu re o r f a o d at ab as e (a nn ua l v al ue ) hooimeijer.qxd 2006/01/10 04:35 pm page 97 of almost 40 %. the percentage of mopane in the diet was the highest during the late dry season (august–november) (fig 1.), reaching a peak of 91 % in september. the contribution of c. mopane to the diet was always >16 %, except for march when it dropped to 3 % and was replaced by combretum apiculatum. besides c. mopane, only six species comprised >5 % of the kudu diet when averaged over the year; these were grewia bicolor (9.4 %), terminalia prunioides (7.8 %), tinnea rhodesiana (6.5 %), boscia albitrunca, and grewia spp. seeds (5.1 %). grass species were only eaten at the beginning of the wet season, with a maximum contribution of 13.5 % in samples in december. the percentage of grasses in samples declined as the wet season progressed. in february, when the fruits of sclerocarya birrea (marula) ripened (late wet season), large quantities were found in the dung samples which peaked at 16 % contribution in this month’s samples. the contribution of c. mopane in the kudu faecal samples could be best explained by a quadratic regression model in which the monthly percentage of c. mopane in the diet was significantly related to the monthly precipitation records (n = 12; f = 4.03; p < 0.05; r² = 0,472; fig. 2). the number of dietary species during the dry season was relatively stable and ranged from eight to 13 between months. during the wet season, the contribution of other species gradually increased from 15 species in december to 25 in april, including the species encountered only once. the average number of different species recorded in the faecal samples in the dry season was 10.8, which is significantly lower than the 18.2 consumed during the wet season (anova, f1,10 = 8.417; p < 0.02). diet quality the calculated crude protein content declined in the late dry season and reached its minimum (6.2 % dw) in october. in the late wet season (may 2004), the maximum crude protein values recorded were 12.6 % dw, corresponding to the month with the highest number of species found in the diet koedoe 48/2 (2005) 98 issn 0075-6458 fig 2. relationship between monthly amount of rainfall (mm) and % of colophospermum mopane leaf material in the diet of kudus. (table 1). surprisingly, no significant differences (anova, f3,8 < 3.583; p > 5 %) could be found in average cp, tannin or phenol concentrations between the wet and dry season, probably because high diet cp persisted quite long into the dry season, as shown in table 2. however, when the analysis was repeated for the dominant diet species (c. mopane) only, significant differences were detected, with significantly lower values of cp, tannins, and phenols in the late dry season compared to the wet seasons (anova, f3,8 > 6.664; p < 0.03; with a tukey post hoc test at 5 %). condensed tannin content, projected for the diet, generally followed the same trend as crude protein levels with low levels in the late wet and late dry seasons (table 2). no significant seasonal difference could be detected for average tannin levels. measured condensed tannin levels in c. mopane were only 3.8 % dw in october/november (annual lowest), but the tannin values more than doubled in december/january to 9.8 % dw, hooimeijer.qxd 2006/01/10 04:35 pm page 98 when the percentage of c. mopane in the diet declined to 16 %. condensed tannin levels in mopane leaves were between 7.2–9.8 % for the entire period between december and july. the other species were lower in tannin levels than c. mopane, normally < 5 % (table 1), except for schotia brachypetala, with estimated levels around 10 %. mopane tannin content increased significantly concomitant with crude protein levels (spearman-r = 0.657; p < 0.05; n =12). however, no significant correlation between tannin and crude protein, comprising all species, could be detected (spearman-r = 0.399; p > 0.05; n = 12). phenol content varied over the season; in november the estimated dietary phenol content reached a peak of 13.0 % dw, caused by a large amount of terminalia prunioides (62.7 %) in the diet. terminalia prunioides and combretum apiculatum had higher phenol levels than c. mopane throughout the year. lowest dietary phenol levels were recorded in february at 3.5 %. the phenol content of only c. mopane increased significantly with cp levels (spearman-r = 0.600; p < 0.05; n = 12). no significant correlation issn 0075-6458 99 koedoe 48/2 (2005) table 2 diet quality estimates in terms of crude protein, condensed tannins and phenol season month nutritional component (%) crude condensed phenol protein tannin early wet january 11.1 4.9 8.0 early wet february 10.1 3.7 3.5 late wet march 10.0 2.0 4.2 late wet april 10.6 3.3 5.2 late wet may 12.6 4.2 6.0 early dry june 11.0 5.8 5.6 early dry july 10.5 7.0 6.8 early dry august 10.6 4.7 6.3 late dry september 9.5 4.9 5.8 late dry october 6.2 2.5 3.6 late dry november 9.4 3.0 13.0 early wet december 11.3 3.7 7.5 could be found between the percentage occurrence of c. mopane foliage in the diet of kudu and any of the other three parameters of c. mopane foliage quality (spearman test). discussion owen-smith & cooper (1987) stated that the local soil nutrient status is an important factor in influencing plant defence mechanisms against herbivory. the majority of our forage quality estimates was obtained from field samples, but some literature data were used as well, which may not be representative for the quality of the species occurring in the study area. hence, it is possible that the results of the diet quality estimates might deviate from actual values. another point of discussion is the availability of the different forage species, which was not measured in this study, and could also influence kudu diet selection. so, further research is needed to support the general validity of our findings. we found that the diet quality between the wet season and dry season was not significant in terms of cp, tannin or phenol content. dekker & smit (1995) revealed that browse availability in the study area is lowest from september to november (late dry season). during the same period, estimated dietary crude protein levels reached a minimum with cp levels varying between 6.2–9.5 % between months. as was hypothesised, the contribution of c. mopane to the kudu diet peaked during this period of limited forage availability. the large contribution to the kudu diet and the long leaf carriage period of c. mopane, which often appears as early as september with leaf senescence starting as late as june (dekker & smit 1995), illustrates that c. mopane is the most important species in the kudu diet. dekker & smit (1995) also noticed that the long leaf carriage period of c. mopane, relahooimeijer.qxd 2006/01/10 04:35 pm page 99 tive to other tree species, underlies its potential value as a fodder resource in the study area. the expected correlation between increased intake of c. mopane foliage and decreasing rainfall was confirmed in this study, and followed a quadratic relationship. several parallels with research on kudu of the nylsvley nature reserve, where burkea africana and ochna pulchra are dominant tree/shrub species (owen-smith & cooper 1988), could be noticed. their study, conducted at the nylsvley nature reserve showed that during the late wet season forbs comprised circa half the diet, but during the course of the dry season, foliage of palatable evergreen species became the most important dietary component. our study also revealed that the species that were generally favoured became more important during the course of the dry season, forbs however showed to be less favoured in comparison to the nylsvley study. both studies showed that fruits and seeds were often eaten/selected, but comprised a minor part of the diet. grass consumption also revealed a similar pattern; grasses were only consumed at the beginning of the wet seasons, and grass intake decreased as the season progressed. kudu browse not only on foliage, but also on the supporting twigs and shoots. histological faecal analyses can only identify foliage, fruit and seed fragments with certainty. the woody twig material in the diet was therefore not considered. stems and twigs are of a lower quality than foliage and, as our diet quality calculations were based on leaf quality data only, we might have over-estimated the diet quality. other determinants such as trace elements, not included in this study, might also influence the browsing behaviour and nutritional gain of kudu in the mopani veld of the limpopo province. however, the quality of c. mopane was significantly different in the late dry period, with lowest values of cp, tannins and phenols. measured condensed tannin levels of c. mopane were above 7.2 % dw except between september and november (3.8– 5.2 % dw). these findings do not match the results of cooper & owen smith (1985), who found in their study at nylsvley that plant species with condensed tannin contents > 5 % of leaf dry mass are preferentially avoided by kudu, except in periods when little other food was available. at the onset of the wet season the contribution of c. mopane declined as expected but still formed a substantial part of the diet. apparently kudus do not reject c. mopane foliage due to high tannin concentrations. the question is of course, do they have a choice; is the forage availability sufficiently large to enable to switch diet to species with a lower tannin content? the fact that c. mopane formed a substantial part of the kudu diet during the wet season proves that tannin-rich species are not avoided when food availability is sufficiently large. high condensed tannin levels do not seem to pose a threshold limiting browse consumption by kudu in the study area. in fact, the coupling between crude protein levels and secondary compounds is remarkable, and the tannin content does not increase with the building-up of structural carbohydrates in the dry season. this atypical pattern in mopane is confirmed in recent studies (wessels et al., in prep.). sinclair (1975) and owen-smith (1982) showed that during the season of active vegetation growth, herbivores are surrounded by an abundance of potential food in the form of plant foliage. we found that the number of species comprising the kudu diet increased at the beginning of the wet season. kudus seem to select for diet quality, but are restricted in their choices by the availability of species in their habitat. however, during the dry or cold season when many deciduous plants are dormant, food availability decreases drastically in both quantity and quality. against a background of low forage availability and quality the timing of leaf flushes and the ripening of fruits therefore have an important influence on dietary composition, as could be seen in this study by the increased intake of grewia seeds, sclerocarya birrea fruits, and terminalia prunioides leaves the moment they became available. owen smith & novellie (1982) found that kudu exhibited a tendency to widen their diet as the dry season advanced. they suggested that kudus were koedoe 48/2 (2005) 100 issn 0075-6458 hooimeijer.qxd 2006/01/10 04:35 pm page 100 limited more by the availability of food than by the effects of food quality on digestive rates. however, our study showed that the number of species in the kudu diet increases significantly during the wet season compared to the dry season. hence, we can conclude that kudus do not restrict their diet to quality-rich species only in times of high forage availability. this pattern of a broad diet in times of plenty and a restricted diet when the food availability and quality is reduced, is not a pattern predicted by the optimal foraging theory in terms of nutrient or energy maximisation (stephens & krebs 1986), but seem to follow the idea, also illustrated in owen-smith & cooper (1987), that dietary thresholds are more important than optimisation. however, forage availability and quality are confounded in this study, and therefore calls for controlled feeding experiments in order to be able to separate the unique effect of forage availability and forage quality on kudu diet selection. except for the low forage availability, and therefore limited potential of alternative forage species in the dry season, an alternative hypothesis might be found in the positive correlation between protein levels and tannin and phenol concentration. choosing for high protein forage automatically increases the concentration of the secondary compounds in the diet. in order to avoid toxic effects, kudus seem to follow the satiety hypothesis (provenza et al. 2003), and diversify their diet in the wet season, leading to an increased number of species in the period that secondary compound concentrations are at its highest. owen smith & novellie (1982) also suggested that large herbivores need to restrict their intakes of various potential toxins below certain maximum thresholds, and that this could be an important factor promoting varied diets. in fact, we found a significant positive correlation between the condensed tannin concentration of c. mopane leaves and the total number of species in the diet (rs = 0.701, p < 0.01, n = 12); the higher the tannin concentration, the more forage species the kudus consumed. this diversification could also be an alternative explanation for the fact that the average diet crude protein concentration was not larger during the wet season. a diverse diet, spreading the risk of an excessive intake of secondary compounds, seems more important than maximising protein intake. acknowledgements special thanks go to chris kellermann for providing us with the data of the chemical analyses. furthermore we wish to thank norman owen-smith for his invaluable comments and advice on the article. willemien luus-powel and martin potgieter are thanked for various forms of assistance. this research was supported by grants from the delta scholarships, the fona foundation for research and nature conservation, eskom-tesp, nrf pretoria and the united kingdom department for international development (dfid) [reference number: r7822]. project title: mopane woodlands and the mopane worm: enhancing rural livelihoods and resource sustainability for the benefit of developing countries. the views expressed are not necessarily those of dfid. the university of limpopo is thanked for various forms of assistance and the department of agriculture, limpopo province for permission to conduct research on the messina experimental farm. references acocks, j.p.h. 1988. veld types of south africa. 2nd edition. memoirs of the botanical survey of south africa 57. aganga, a.a. & t. adogla-bessa. 1999. dry matter degradation, tannin and crude protein contents of some indigenous browse plants of botswana. archivos de zootecnica 48(181): 79–83. aganga, a.a. & k.w. mosase. 2001. tannin content, nutritive value and dry matter digestibility of lonchocarpus capassa, zizyphus mucronata, sclerocarya birrea, kirkia acuminata and rhus lancea seeds. animal feed science and technology 91: 107–113. bailey, e.m. 1978. physiologic response of livestock to toxic plants. journal of range management 31: 343–347. bartolome, j., j. franch, m. gutman & n.g. seligman. 1995 physical factors that influence fecal analysis estimates of herbivore diets. journal of range management 48: 267–270. bothma, j. du p., n. van rooyen & m.w. van rooyen. 2004 using diet and plant resources to issn 0075-6458 101 koedoe 48/2 (2005) hooimeijer.qxd 2006/01/10 04:35 pm page 101 set wildlife stocking densities in african savannas. wildlife-society-bulletin 32: 840–851. cid, m.s. & m.a. brizuela. 1990. grass blade and sheath quantification by microhistological analysis. journal of wildlife management 54: 349–352. cooper, s.m. & n. owen-smith. 1985. condensed tannins deter feeding by browsing ruminants in a south african savanna. oecologia 67: 142–146. de jong, c.b., s.e. van wieren, r.m.a. gill & r. munro. 2004. relationship between diet and liver carcinomas in roe deer in kielder forest and galloway forest. veterinary record 155: 197–200. dekker, b. & n. van rooyen. 1995. the physical environment and plant communities of the messina experimental farm. south african journal of botany 61(3): 158–167. dekker, b. & g.n. smit. 1996. browse production and leaf phenology of some trees and shrubs in different colophospermum mopane savanna communities. african journal of range & forage science 13(1): 15–23. food and agriculture organization of the united nations (fao), homepage. www.fao.org. http://www.fao.org/ag/aga/agap/frg/afris/i ndex_en.htm (15-12-04). furstenburg, d. & w. van hoven. 1994. condensed tannin as anti-defoliate agent against browsing by giraffe (giraffa cameleopardalis) in the kruger national park. comparative biochemistry and physiology 107a: 425–431. freeland, w. j. 1991. plant secondary metabolites: biochemical coevolution with herbivores. pp. 61–81. in: palo, r.t. & c.t. robbins (eds.). plant defenses against mammalian herbivory. boca raton: crc press. hagerman, a.e. 2002. tannin handbook. http:// www.users.muohio.edu/hagermae/ (05-06-05) lagendijk, d.d.g. 2003. mopane woodlands: a food source for elephants. msc thesis, wageningen university, wageningen. lewis, s.w. 1994. fecal and rumen analyses in relation to temporal variation in black-tailed deer diets. journal of wildlife management 58: 53–58. macala, j., b. sebolai & r.r. majinda. 1992. colophospermum mopane browse plant and sorghum stover as feed resources for ruminants during the dry season in botswana. pp. 151–161. in: stares, j.e.s., a.n. said & j.a. kategile (eds.). the complimentary of feed resources for animal production in africa. proceedings of the joint feed resources network workshop. ilca, addis ababa, ethiopia. mcarthur, c., a. hagerman & c.t. robbins. 1991. physiological strategies of mammalian herbivores against plant defenses. pp. 103–114. in: palo, r.t. & c.t. robbins (eds.). plant defenses against mammalian herbivory. boca raton: crc press. mueller-harvey, i. 2001. analysis of hydrolysable tannins. animal feed science and technology 91: 3–20. owen-smith, n. 1982. factors influencing the consumption of plant products by large herbivores. pp. 359–404. in: huntley, b.j. & b.h. walker (eds.). the ecology of tropical savannas. berlin, germany: springer-verlag. owen-smith, n. & s. cooper. 1987. palatability of woody plants to browsing ruminants in a south african savanna. ecology 68(2): 319–331. owen-smith, n. & s. cooper. 1989. nutritional ecology of a browsing ruminant, the kudu (tragelaphus strepsiceros), through the seasonal cycle. the zoological society of london 219: 29–43. owen-smith, n. & p. novellie. 1982. what should a clever ungulate eat? american naturalist 119: 151–178. owen-smith, n. 2002. adaptive herbivore ecology: from resources to populations in variable environments. cambridge: cambridge university press. provenza f.d, j.j. villalba & r.e. banner. 2003. linking herbivore experience, varied diets, and plant biochemical diversity. small ruminant research 49(3): 257–274. sinclair, a.r.e. 1975. the resource limitation of trophic levels in tropical grassland ecosystems. journal of animal ecology 44: 497–520. sparks, d.r. & c. malechek. 1968. estimating percentage dry weight in diets using a microscopic technique. journal of range management 21: 264–265. stace, c.a. 1965. cuticular studies as an aid to plant taxonomy. bulletin of the british museum (natural history) botany 4: 1–78. stewart, d.r.m. 1967. analysis of plant epidermis in faeces: a technique for studying the food preferences of grazing herbivores. journal of applied ecology 4: 83–111. stephens, d.w. & j.r. krebs. 1986. foraging theory. princeton: princeton university press. van hoven, w. 1991. mortalities in kudu (tragelaphus strepsiceros) populations related to chemical defence of trees. revue de zoologie africaine 105: 141–145. van der waal, c., & g.n. smit. 2001. relations between kudu mortalities and woody browse availability in a semi-arid savanna. pp. 38-43. in: ebedes, h., b. reilly, w. van hoven & b. penzhorn (eds.). sustainable utilisation – conservation in practice. pretoria: south african game ranchers' organisation. walker, b.h. 1980. a review of browse and its role in livestock production in southern africa. browse in africa, the current state of knowledge. addis ababa: ilca. zar, j.h. 1984. biostatistical analysis. london: prentice hall. koedoe 48/2 (2005) 102 issn 0075-6458 hooimeijer.qxd 2006/01/10 04:35 pm page 102 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 barrett.qxd phytosociology and plant community utilisation by vervet monkeys of the blydeberg conservancy, limpopo province a.s. barrett, l.r. brown, l. barrett and s.p. henzi barrett, a.s., l.r. brown, l. barrett and s.p. henzi. 2006. phytosociology and plant community utilisation by vervet monkeys of the blydeberg conservancy, limpopo province. koedoe 49(1): 49–68. pretoria. issn 0075-6458. the plant communities of the blydeberg conservancy were investigated as part of a research project on the foraging ecology of vervet monkeys cercopithecus aethiops pygerythrus (senso lato) in mixed lowveld bushveld and sour lowveld bushveld areas. to date there are no formal management plans for vervet monkeys. this is attributed to the limited knowledge of vervets and their utilisation of and impacts on ecosystems. from a twinspan classification refined by braun-blanquet procedures, ten plant communities that can be placed into four major groups were identified. a classification and description of these communities, including a vegetation map are presented. diagnostic species as well as prominent and less conspicuous species of tree, shrub, herb and grass strata are outlined. of the ten available plant communities, the vervets utilised only six during the study period. there was an abundant supply of various food sources throughout the year, with movement patterns mostly coinciding with the fruiting times of several tree and other plant species. keywords: braun-blanquet procedures, conservancy, plant communities, twinspan, vervet monkeys. a.s. barrett and l.r. brown, applied behavioural ecology and ecosystem research unit, university of south africa, florida campus, rsa; l. barrett and s.p. henzi, applied behavioural ecology and ecosystem research unit, university of south africa, florida campus, rsa (present address: department of psychology, university of central lancashire, lancashire, uk) issn 0075-6458 49 koedoe 49/1 (2006) introduction the blydeberg conservancy constitutes a group of privately owned farms belonging to individuals with varying interests. owners have diverse backgrounds including farming, hunting, research, business and property development. the intention of the conservancy was to pool resources and properties to form a large area containing various game species for tourism ventures and private utilisation. not many extensive vegetation studies have been carried out in the blydeberg area or surrounds and those that have been completed (van der schijff 1963; van der schijff & schoonraad 1971), were done only once, without any recorded follow-up surveys. due to the various land-use practices occurring within the area and their long-term effects on ecosystems, as well as the research project on vervet monkeys cercopithecus aethiops pygerythrus (senso lato) undertaken in the area (barrett 2005), it was deemed important to undertake a vegetation description and classification of the blydeberg conservancy. it is widely recognised that a detailed description, identification, classification and mapping of vegetation forms the basis of sound land-use planning and management (tueller 1988; fuls et al. 1992; fuls 1993; bezuidenhout 1996; brown 1997). the results obtained from this study could therefore be used to assist management in making decisions for the conservancy, whilst simultaneously providing data for the determination of the seasonal foraging ecology of vervet monkeys. the main aim of this study thus was to describe and map the plant communities of the study area within the blydebarrett.qxd 2006/04/17 01:47 pm page 49 berg conservancy in order to determine the seasonal foraging ecology of a troop of vervet monkeys living within the area. vegetation assessments are prerequisites to any ecological or habitat related research, forming a basis to any further studies. vervets are extremely adaptable opportunistic generalists (similar to baboons but, less so) that easily move into disturbed areas, including agricultural areas and lodges. they can even be found in specialised habitats such as mangrove swamps (oates 1996; kingdon 1997; fedigan & fedigan 1988; skinner & smithers 1990; booth 1979). vervets are capable of becoming a pest species in areas where they are forced into close proximity with humans, leading to conflict and ultimately their demise. the aim of the encompassing study on vervet foraging ecology is to elucidate some of their foraging behaviour. it is anticipated that results obtained from such a study would assist management in successfully implementing management plans for vervets. not many detailed ecological studies of vervets have been undertaken in temperate subtropical areas (struhsaker 1967; harrison 1983, 1984; whitten 1988; lee & hauser 1998; isbell et al. 1998; pruetz & isbell 2000). at these low latitudes, overall habitat productivity is high and seasonal variability is relatively constrained (caughley & sinclair 1994). a fuller understanding of vervet ecological flexibility, as well as the factors that might limit their distribution both broadly and locally, is likely to be derived from this study and will benefit the overall management of the species. no related vervet studies have been undertaken locally; however, a study on baboon resource utilisation has been done in the blyde canyon nature reserve (marais 2005), though they mostly feed at different strata. study area the blydeberg conservancy is situated approximately 19 km south of the town of hoedspruit in the limpopo province (fig. 1). it is located between longitude 30º27'e–25º56'e and latitude 24º23's– 24º28's. altitude ranges from 350–800 m above sea level (low & rebelo 1998). the study area constitutes the farms dunstable (farm number 230) and jongmanspruit (farm number 234). the current size of the blydeberg conservancy is 3000 ha with the study area being 816 ha. the geology of the conservancy ranges from a zone of fine to medium-grained quartzite, gritty in places, with pebble layers; basic lava, tuff, agglomerate and shale in the south; to a zone of light-grey, mediumgrained biolite gneiss with coarse-grained quartz-feldspar leucosomes, recrystallised in places in the north (visser 1989; walraven 1989). an intermediate zone of laminated micaceous and graphitic shale, locally interlayered with sandy shale, flagstone and quartzite seperates the southern and northern zones (visser 1989; walraven 1989). also, according to visser (1989) and walraven (1989), intermittently layered along the south of the northern zone and between such and the intermediate zone is another less distinct zone of greenish grey intermediate lava, amygdaloidal in places, interbedded porphyritic layers and layers of shale and quartzite. visser (1989) and walraven (1989), state that the southern, intermediate and the less distinct zone between the northern and intermediate zones are rocks of volcanic and sedimentary origin dating back to the vaalian quaternary. the southern zone originates from the black reef formation of the transvaal sequence; the intermediate zone originates from the selati formation of the undifferentiated upper part of the wolkberg group and the less distinct zone between the northern and intermediate zones originates from the abel erasmus formation of the undifferentiated lower part of the wolkberg group (visser 1989; walraven 1989; buckle 1992). the northern zone is rock of intrusive origin, dating back to the swazian quaternary (visser 1989; walraven 1989; buckle 1992). the topography is mostly mountainous with steep to moderately steep slopes gradually tapering off to a relatively flat mountain koedoe 49/1 (2006) 50 issn 0075-6458 barrett.qxd 2006/04/17 01:47 pm page 50 plateau. patches of montane forest occur at higher altitudes in kloofs and ravines. due to dense shade caused by an almost continuous tree canopy, very little ground cover is present in the forest patches, however many ferns and lianas including dalbergia armata, capparis tomentosa, strophanthus speciosus and hippocratea crenata prevail. larger canopy trees found in such patches include celtis africana, rauvolfia caffra, trichilia dregeana, nuxia oppositifolia and ficus sycomorus. the vegetation comprises a mixture of dense montane forest in kloofs; open tree savanna/sour bushveld on northern and western slopes; and high mountain sour grass veld and dense riverine thickets along drainage lines and streams. the area is rather patchy and vegetation varies from relatively open bushveld with long grass species on undulating hills and slopes, through dense shrub and scrub in dry ravines and dongas, to patches of semi-montane forest in sheltered kloofs. according to low & rebelo (1998), the area can be classified as intermediate between mixed lowveld bushveld (veld type 19) and sour lowveld bushveld (veld type 21). four land types, namely fa, fb, ib and ic, occur in the study area. according to land type survey staff (1989), “a land type denotes an area that can be shown at 1:250 000 scale and displays a marked degree of uniformity with respect to terrain form, soil pattern and climate”. the f land type refers to pedologically young landscapes that are not predominantly rock and alluvial or aerolian in which the main soil forming processes have been rock weathering (land type survey staff 1989). dominant soil forms are glenrosa and mispah. however, exposed rock and soils belonging to almost any of the other 39 soil forms may occur in type f land types. both shallow and deep soils of the oakleaf form are present in upland areas. in the fa land type lime in the soil is not commonly encountered and is rare or absent throughout the landscape. this land type consists of two terrain units, mid slopes comprising 95 % of the land type, and valley bottoms constituting the remaining 5 % of the land type. soils are mostly medium sandy loam to sandy clay loam and are shallower than 400 mm (land type survey staff 1989). in the fb land type, lime in the soil is rare or absent in upland areas but, generally, lime is present in low-lying areas. this land type consists of two terrain units, footslopes comissn 0075-6458 51 koedoe 49/1 (2006) fig. 1. location of blydeberg conservancy within the limpopo province of south africa. barrett.qxd 2006/04/17 01:47 pm page 51 prising 80 % of the land type, and valley bottoms constituting 20 % of the land type. soils are mostly medium to coarse sand to loamy sand and shallower than 1200 mm (land type survey staff 1989). the land type ib indicates land types with exposed rock (country rock, stones or boulders), which covers 60–80 % of the area. this land type consists of three terrain units, crests comprising 20 % of the land type, mid slopes comprising 75 % of the land type, and valley bottoms constituting the remaining 5 % of the land type. soils are mostly medium sandy clay loam to sandy clay and shallower than 900 mm (land type survey staff 1989). the land type ic refers to land types with exposed rock (country rock, stones or boulders), covering more than 80 % of the area. this land type consists of four terrain units, crests comprising 10 % of the land type, scarps comprising 20 % of the land type, mid slopes comprising 68 % of the land type, and valley bottoms constituting the remaining 2 % of the land type. soils are mostly fine to medium sandy loam to sandy clay loam and shallower than 300 mm (land type survey staff 1989). climate the average annual rainfall for the study area, as measured by a weather station situated on the jongmanspruit farm (234) for the period 05/1999 to 04/2004 was 561 mm, with a high of 953 mm and a low of 255 mm recorded in 2000 and 2002, respectively. for the period 05/1999 to 04/2004, average monthly rainfall varied from 0.7 mm during the dry winter season (may to october) to 106 mm in the wet summer season (november to april) (fig. 2). for the larger hoedspruit region, average annual rainfall for the period 05/1999 to 04/2004 was 900 mm with a high of 1463 mm and a low of 629 mm recorded in 2000 and 2002, respectively. for the period 05/1999 to 04/2004, average monthly rainfall varied from 2 mm during the dry winter season (may to october) to 189 mm in the wet summer season (november to april) (fig. 2). rainfall recorded for the study area is less than that of the larger hoedspruit region due to the study area’s location on the foothills of the drakensberg mountains along the great escarpment. the mountains form a barrier causing a rain shadow which could be responsible for less rainfall in the study area. however, the mountains do function as an important catchment for the study area and are the source of several small mountain koedoe 49/1 (2006) 52 issn 0075-6458 fig. 2. average monthly rainfall and temperatures for the blydeberg conservancy and hoedspruit from 05/1999 to 04/2004. barrett.qxd 2006/04/17 01:47 pm page 52 streams (van zyl 2003). thunderstorms and fog are the main sources of precipitation for the study area. average annual temperature for the study area for the period 05/1999 to 04/2004 was 22 ºc, with mean temperatures varying from 17 ºc during the dry winter season to 26 ºc in the wet summer season (fig. 2). a minimum temperature of 3 ºc and a maximum of 42 ºc were recorded in the 05/2003 to 04/2004 period. temperatures for the study area are higher than those for the larger hoedspruit region due to the study area lying along the northfacing footslopes of the drakensberg mountains along the great escarpment. the study area has more direct exposure to sunlight and is more sheltered from southerly winds than the surrounding areas (tyson & preston-whyte 2000). methods vegetation classification and description using a 1:10 000 digitised ortho photo for the blydeberg area, the study area was stratified into physiognomic-physiographic units (barbour et al. 1987; kent & coker 1997). a total of 49 sample plots were located in a randomly stratified manner within these units to ensure that all variations in the vegetation were considered and sampled. plot sizes were fixed at 400 m² (barbour et al. 1987; brown & bredenkamp 1994; brown 1997). in each sample plot, all species were recorded, all woody species counted, and cover abundance was assessed using the braunblanquet cover abundance scale (mueller-dombois & ellenberg 1974; barbour et al. 1987). fieldwork was undertaken during april, may, and june 2003 as well as in january 2004. in the light of taxon names constantly changing, this publication conforms to those of arnold & de wet (1993). structural terminology for the woody species is according to edwards (1983). environmental data collected included aspect, estimates of slope, percentage rockiness and percentage erosion. floristic data was analysed using the turboveg suite (hennekens 1996a), which includes the two-way indicator species analysis multivariate classification technique twinspan (hill 1979) for deriving an initial approximation of the main plant communities. this numerical classification technique is regarded as a successful approach to vegetation classification by various phytosociologists (brown & bredenkamp 1994; bredenkamp & bezuidenhout 1995; brown et al. 1996; cilliers 1998). the visual editor megatab by hennekens (1996b) was used to generate a phytosociological table. further refinement of the classification was undertaken through the application of braun-blanquet procedures (barbour et al. 1987; bredenkamp et al. 1989; kooij et al. 1990; bezuidenhout 1993; eckhardt 1993; brown & bredenkamp 1994; kent & coker 1997). using the final phytosociological table and habitat information collected during sampling in the field, different plant communities were identified, described and ecologically interpreted. erosion was estimated within the following classes (matthee & van schalkwayk 1984): class 1 = slight; class 2 = moderate loss of topsoil with slight soil cutting by run-off channels or gullies; class 3 = severe loss of topsoil with marked soil cutting by run-off channels or gullies; class 4 = total loss of topsoil and exposure of subsoil or deep intricate soil cutting by gullies. slope was estimated within the following categories: level = 0–3º; gentle = 3–8º; moderate = 8–16º; steep = 16–26º; very steep = 26–45º. a variation of the gradient analysis ordination techniques (barbour et al. 1987; kent & coker 1997) was used for community ordination. each community’s vegetation cover was split into four cover classes i.e. tree, shrub, herb and grass cover. using ms excel, the arithmetic mean for each cover class per community was calculated. average percentages were plotted for each community, and for each cover class a moving average was calculated using the least squares method. the moving averages were used to generate trend lines that predict potential future tendencies in cover classes across communities. the same procedure was followed for slope, erosion and rockiness. plant community selection electivity of available plant communities was calculated using ivlev’s electivity index, i.e., plant community electivity = ri-ni/ri+ ni where ri = proportion of community utilised and ni = proportion of community available in home range (krebs 1989). the proportion of a community utilised was calculated as a percentage of all the communities size; the proportion of a community’s availability within the home range was calculated as a percentage of the size of all communities. values for ivlev’s electivity index range between -1 and 1; >0 indicates positive selecissn 0075-6458 53 koedoe 49/1 (2006) barrett.qxd 2006/04/17 01:47 pm page 53 tion of a food item; <0 indicates selection against, or avoidance of a food item. results vegetation classification the analysis resulted in the identification of the following ten plant communities (fig. 3), which may be grouped into four major community types (table 1): 1. eragrostis lehmanniana-grewia flava shrubland. 1.1 heteropogon contortus-grewia flava shrubland. 1.2 ziziphus mucronata-grewia flava shrubland. 2. bridelia mollis-acacia nigrescens woodland. 2.1 berchemia zeyheri-acacia nigrescens woodland. 2.2 combretum imberbe-acacia nigrescens woodland. 3. acacia nigrescens-combretum apiculatum woodland. 4. gymnosporia glaucophylla-panicum maximum woodland. 4.1 balanites maughamii-panicum maximum woodland. 4.2 sclerocarya birrea-panicum maximum woodland. 4.3 combretum zeyheri-panicum maximum woodland. 4.4 asparagus setaceus-philenoptera violacea woodland. 4.4.1 pappea capensis variant. 4.4.2 diospyros mespiliformis variant. description of plant communities the study area is mostly open woodland with a depleted herbaceous layer due to a history of overgrazing and irregular agricultural practices leading to a low plant species diversity. there is a strong presence of the shrub species grewia flava and grewia flavescens throughout the study area. the general vegetation of the study area is characterised by the presence of species from species groups o, p and q occurring in communities 1, 2, 3 and 4; and with species from species group n found mostly in communities 1, 2 and 3. for all species groups refer to table 1. prominent species include: the shrubs grewia flavescens, grewia flava (species group q), dichrostachys cinerea (species group p); the grass elionurus muticus (species group q) which occur in all communities; and the trees combretum apiculatum and ziziphus mucronata (species group p) which occur in most communities, with the exception of the diospyros mespiliformis variant (4.4.2.). the tree sclerocarya birrea (species group o) is prominent in most communities except sub-community 4.4. while the tree acacia nigrescens (species group o) is dominant in communities 1, 2, 3 and subcommunity 4.1 (table 1). table 2 depicts additional plant species recorded for blydeberg but not significant enough to be reflected in table 1. the species names and the communities they occur in are presented. 1. eragrostis lehmanniana-grewia flava shrubland this shrubland plant community covers approximately 24 % of the study area (193 ha) and consists of flat rocky terrain with the vegetation in various stages of degradation (fig. 3). soils are sandy, gravelly and relatively well drained. rock cover for this community varies between 20–45 % and erosion is estimated at 15 %. species belonging to species group a are diagnostic for this community and include the tree sterculia rogersii; the grasses eragrostis lehmanniana, aristida congesta, a. stipitata; and the forbs sida cordifolia and corchorus kirkii. vegetation is dominated by the trees acacia nigrescens, sclerocarya birrea (species group o), ziziphus mucronata and combretum apiculatum (species group p), while the shrubs dichrostachys cinerea (species group p), grewia flava, grewia flavescens (species group q) are also prominent. the grass layer is not well developed and includes elionurus muticus (species group q), panicum maximum (species group f) and eragrostis lehmanniana (species group a). koedoe 49/1 (2006) 54 issn 0075-6458 barrett.qxd 2006/04/17 01:47 pm page 54 portions of this community were previously used for cultivation purposes and are in various stages of secondary succession as can be seen from the large number of different species that are prominent within this community. this community is divided into two sub communities. although they are delineated on the vegetation map, small sections of sub community 1.1 form a mosaic distribution pattern within sub community 2.2 and could therefore not be mapped. 1.1 heteropogon contortus-grewia flava shrubland the heteropogon contortus-grewia flava sub-community is situated in three small areas in the northwest and central sections of the study site (fig. 3). this sub-community covers 3 % of the study area (20 ha) and 10 % of community 1, and occurs on what appears to be more disturbed sites or pockets. the areas are relatively rocky (20 % rock cover) and fairly open. erosion is estimated at 15 % with gentle slopes. soils are mostly sandy and gravelly. the grasses heteropogon contortus, digitaria eriantha and pogonarthria squarrosa (species group b) are diagnostic for this subcommunity. the vegetation is dominated locally by the trees combretum apiculatum (species group p), acacia nigrescens and sclerocarya birrea (species group o) together with the shrubs dichrostachys cinerea (species group p), and grewia flavescens (species group q). the herbaceous layer is not well developed and include the grasses eragrostis lehmanniana (species group a), digitaria eriantha, heteropogon contortus (species group b) and panicum maximum (species group f). the average number of plant species per 400 m² in this sub-community is 17. the tree layer has a 10–60 % cover with an average of 37 %; the shrub layer has a 10–70 % cover with an average of 29 %; the herb layer has a 0–5 % cover with an average of 2 %; and the grass layer has a 1–70 % cover with an average of 26 %. the shrub dichrostachys cinerea has the highest density of 5333 individuals/ha followed by the tree acacia nigrescens with 2583 individuissn 0075-6458 55 koedoe 49/1 (2006) fig. 3. vegetation map of the study area within the blydeberg conservancy. map depicts the various communities, sub-communities and variants. als/ha and the shrub acacia karroo with 2167 individuals/ha. 1.2 ziziphus mucronata-grewia flava shrubland. this sub-community comprises two separate sections. one section is located on the northern boundary of the study area while the other is situated on the eastern boundary extending towards the southern boundary in a narrow strip, isolating sub-community 2.1 from the rest of the study area (fig. 3). this sub-community covers 21 % of the study area (173 ha) and 90 % of community 1, being flat and relatively open with signs of prior mismanagement in the form of bush encroachment and several stands of samesized and aged trees occurring within this sub-community. the areas are quite rocky (45 %). erosion is estimated at 15 % with gentle slopes. soils are mostly sandy, gravelly and relatively deep. barrett.qxd 2006/04/17 01:47 pm page 55 koedoe 49/1 (2006) 56 issn 0075-6458 ta bl e 1 p hy to so ci ol og ic al ta bl e of th e ve ge ta tio n fo r th e st ud y ar ea . barrett.qxd 2006/04/17 01:47 pm page 56 issn 0075-6458 57 koedoe 49/1 (2006) ta bl e 1 (c on tin ue d) barrett.qxd 2006/04/17 01:47 pm page 57 the trees acacia exuvialis, commiphora africana, the shrub balanites pedicellaris, the herbs cyclopia capensis and bidens pilosa, the grass anthephora pubescens, and the forb solanum panduriforme (species group c) are diagnostic species for this subcommunity. the vegetation is dominated by the trees acacia nigrescens (species group o), ziziphus mucronata, combretum apiculatum (species group p); and the shrub grewia flava (species group q). the tree sclerocarya birrea (species group o) is also very prominent within this sub-community. the herbaceous layer comprises the grasses elionurus muticus (species group q) and eragrostis lehmanniana (species group a) that are locally prominent. the pioneer forbs bidens pilosa and solanum panduriforme (species group c) are also present. the average number of plant species is 15/400 m² in this sub-community. the tree layer has a 5–80 % cover with an average of 46 %; the shrub layer has a 5–90 % cover with an average of 42 %; the herb layer has a 0–80 % cover with an average of 17 %; and the grass layer has a 1–70 % cover with an average of 17 %. the tree acacia exuvialis has the highest density (6000 ind/ha), followed closely by the shrub grewia flava (5222 ind/ha) and the tree ziziphus mucronata with 2 667 ind/ha. 2. bridelia mollis-acacia nigrescens woodland. the bridelia mollis-acacia nigrescens woodland comprises the largest proportion of the study area and covers 51 % (415 ha) of the conservancy (fig. 3). soils are mostly sandy and well drained with a non-perennial koedoe 49/1 (2006) 58 issn 0075-6458 table 2 table of less prevalent plant species occurring at blydeberg and not represented in table 1 species name community number species name community number acacia burkei 1.1, 2.1 melinis repens 2.2 acacia galpinii 1.1 nuxia oppositifolia 4.4 acacia tortilis 1.1 olea europaea 2.1, 4.1 aloe dyeri 1.2 opuntia ficus-indica 2.2 aloe marlothii 4.1, 4.2 ozoroa paniculosa 1.2, 2.2 asparagus cooperi 1.1, 3 ozoroa sphaerocarpa 4.2 bacchar adoensi v. kot 2.1 pavonia columella 4.2, 4.4 boscia albitrunca 1.2, 2.2 plectroniella armata 2.2 canthium ciliatum 2.1 rauvolfia caffra 4.3 capparis tormentosa 4.3 rhigozum obovatum 1.1 celtis africana 4.4 rhus dentata 4.4 chloris virgata 1.1 rhus tumulicola 4.3 cussonia spicata 1.1 schotia brachypetala 4.1 diheteropogo filifoliu 1.2 scolopia zeyheri 4.4 eragrostis rigidior 4.4 spirostachys africana 4.4 eragrostis superba 2.1 strelitzia nicolai 1.2 eustachys paspaloides 4.2 strophanthus gerrardii 4.2, 4.3 ficus sycomorus 4.3 strophanthus speciousus 4.3 garcinia livingstonei 4.3 tagetes minuta 1.1, 1.2 hippocratea crenata 4.3 terminalia sericea 1.1, 2.1 hyparrhenia hirta 1.2 trichilia emetica 4.4 hypoxis hemerocallidea 1.1 vernonia adoensis 2.1 leucosidea sericea 1.2 wachendorfia parviflor 4.2 maerua angolensis 4.1 xanthium spinosum 2.1 markhamia zanzibarica 4.2 barrett.qxd 2006/04/17 01:47 pm page 58 mountain stream passing through the centre of this community. rock cover and erosion for this community varies between 25–35 % and 10–15 %, respectively. species belonging to species group d are diagnostic for this community and include the trees ficus stuhlmannii, combretum imberbe and bridelia mollis. the vegetation is dominated by the trees acacia nigrescens (species group o), combretum apiculatum (species group p) and acacia karroo shrubs (species group n). the trees bridelia mollis (species group d) and sclerocarya birrea (species group o) are locally prominent. other species also present include the shrubs dichrostachys cinerea (species group p), grewia flava, grewia flavescens (species group q), and the grass elionurus muticus (species group q). the grass species anthephora pubescens (species group c) occurs sparsely in the community, being locally dominant. within this community there are signs of previous overgrazing with old kraals, dipping structures and various old farming implements found in various locations throughout the area. this community consists of two sub-communities. 2.1 berchemia zeyheri-acacia nigrescens woodland. the berchemia zeyheri-acacia nigrescens sub-community is located in two large areas in the southern and southeastern sections of the study area (fig. 3). this sub-community is relatively large—28 % of the study area (226 ha) and 54 % of community 2—and occurs in areas where there are signs of historical disturbance, particularly overgrazing with some relics of agricultural farming found within the area. the areas are flat with gentle slopes and sandy soils with intermittent rocks scattered throughout (35 % rockiness). there is slight evidence of erosion in the form of small furrows at places (10 % erosion). the trees berchemia zeyheri, commiphora mollis, lannea discolor, the shrubs mundulea sericea, euclea divinorum, and the grasses imperata cylindrica and brachiaria brizantha (species group e) are diagnostic for this sub-community. the woody layer comprises a mixture of species with the vegetation dominated by the trees acacia nigrescens (species group o), combretum apiculatum (species group p), and acacia karroo shrubs (species group n). the trees berchemia zeyheri (species group e), sclerocarya birrea (species group o) and ziziphus mucronata (species group p) are locally prominent, while the shrubs grewia flava, grewia flavescens (species group q), gymnosporia glaucophylla (species group g), the grass elionurus muticus (species group q) and the forb lobelia species (species group n) are also present. the average number of plant species is 16/ 400 m² in this sub-community. the tree layer has a 10–80 % cover with an average of 46 %; the shrub layer has a 5–80 % cover with an average of 34 %; the herb layer has a 1–30 % cover with an average of 8 %; and the grass layer has a 1–60 % cover with an average of 20 %. the shrub acacia karroo has the highest density of 5615 ind/ha followed by the trees acacia nigrescens with 3962 ind/ha and combretum apiculatum with 3308 ind/ha. 2.2 combretum imberbe-acacia nigrescens woodland. this sub-community is located centrally and towards the western boundary of the study area, east of community 3 that forms part of the western boundary of the study area (fig. 3). this woodland is relatively large covering 23 % of the study area (189 ha) and 46 % of community 2, and is mainly flat and open with signs of mismanagement in the form of previous overgrazing. rockiness is estimated at 25 % with most rocks occurring adjacent to a dry river bed. erosion is estimated at 15 %. the area has a gentle slope with soils being predominantly sandy and well drained. issn 0075-6458 59 koedoe 49/1 (2006) barrett.qxd 2006/04/17 01:47 pm page 59 koedoe 49/1 (2006) 60 issn 0075-6458 characteristic of this sub-community is the absence of species from species group e. the trees peltophorum africanum (species group g), combretum apiculatum, ziziphus mucronata (species group p) and the shrub dichrostachys cinerea (species group p) dominate the vegetation. other species also present are the tree bridelia mollis (species group d), the shrub corchorus kirkii (species group a) and the grass elionurus muticus (species group q) being locally prominent. the average of 18 plant species per 400 m² are found in this sub-community. the tree layer has a 30–90 % cover with an average of 60 %; the shrub layer has a 5–40 % cover with an average of 18 %; the herb layer has 0 % cover; and the grass layer has a 1–40 % cover with an average of 19 %. the shrub dichrostachys cinerea has the highest density of 5500 ind/ha, with the tree combretum apiculatum having 2500 ind/ha and the shrub grewia flavescens having 1667 ind/ha. 3. acacia nigrescens-combretum apiculatum woodland. this woodland community forms part of the western boundary of the study area comprising approximately 6 % (48 ha) of the study area (fig. 3). slope is moderate with shallow soils as most of the community is on a rocky protrusion. average erosion for this community is estimated at 10 %, with average rockiness estimated at 15 %. the acacia nigrescens-combretum apiculatum woodland is characterised by the absence of any diagnostic species. also characteristic of this woodland is the presence of species group f and the absence of species groups g to m. vegetation is dominated by the trees combretum apiculatum (species group p), acacia nigrescens, sclerocarya birrea (species group o), the shrub grewia flava (species group q), and the grass elionurus muticus (species group q). characteristic of this community is the absence of the shrubs dichrostachys cinerea (species group p) that are present in almost all the other communities of the study area. acacia karroo shrubs (species group n) are also present. this community has a more developed herbaceous layer with the increaser 2 grass eragrostis lehmanniana (species group a) and the decreaser grass panicum maximum (species group f) present throughout this community. there are signs of previous overgrazing by cattle throughout this community, with several old farm implements scattered throughout the community. the dominance of the increaser 3 grass elionurus muticus is indicative of the degraded condition of the herbaceous layer. however, the presence of the grasses panicum maximum and eragrostis lehmanniana is indicative that this community is recovering and in a secondary successional phase. the average number of plant species in this community is 13 per 400 m². the tree layer has an average estimated cover of 32 %; the shrub layer has an average estimated cover of 18 %; the herb layer has an average estimated cover of 1 %; and the grass layer has an average estimated cover of 33 %. the shrub grewia flava has a density of 4833 ind/ha; the tree combretum apiculatum has 4000 ind/ha and the shrub acacia karroo has 3333 ind/ha. 4. gymnosporia glaucophylla-panicum maximum woodland. this woodland plant community covers about 20 % of the study area (160 ha) and is mostly mountainous (fig. 3). soils are predominantly sandy and relatively well drained. rock cover for this community varies between 10–50 % and erosion is 10 %. species belonging to species groups f and g are diagnostic for this community and include: the trees peltophorum africanum, rhus leptodictya; the shrub gymnosporia glaucophylla; and the grass panicum maximum. the vegetation is dominated by the shrubs grewia flavescens and grewia flava (species group q), while the grass elionurus muticus (species group q) is also prominent. the barrett.qxd 2006/04/17 01:47 pm page 60 issn 0075-6458 61 koedoe 49/1 (2006) trees combretum apiculatum and ziziphus mucronata (species group p), and the shrub dichrostachys cinerea (species group p) are also dominant throughout, except in the diospyros mespiliformis variant (4.4.2). there are many signs of animal activity throughout this community, mainly game tracks, middens and foraging evidence. there are also signs of previous grazing by cattle. a few roads pass through the community and a few small dwellings are present. this community consists of four sub-communities and two variants. 4.1 balanites maughamii-panicum maximum woodland. this sub-community is located in the south western section of the study area and consists of two separate areas split by the pappea capensis variant (fig. 3). this sub-community covers 5 % of the study area (43 ha) and 27 % of community 4, and is found on the relatively steep, predominantly northfacing footslopes of the drakensberg mountain range constituting the southern boundary of the study area. there are signs of previous cattle grazing in the lower lying areas. the areas are relatively rocky (30 %). erosion is estimated at 10 % with a moderate to steep slope. soils are mostly sandy. the trees combretum erythrophyllum and balanites maughamii; the shrub gossypium herbaceum; and the perennial herb argyrolobium velutinum (species group h) are diagnostic for this community. the woody layer comprises a mixture of species with the vegetation dominated by the trees combretum apiculatum, ziziphus mucronata (species group p) and sclerocarya birrea (species group o), the shrubs dichrostachys cinerea (species group p) and grewia flavescens (species group q), and the grass elionurus muticus (species group q). the tree acacia nigrescens (species group o) and the shrub grewia flava (species group q) are also present. in this sub-community, the average number of plant species is 15 per 400 m². the tree layer has a 40–80 % cover with an average of 55 %, the shrub layer has a 20–50 % cover with an average of 30 %, the herb layer has 1 % cover, and the grass layer has a 40–60 % cover with an average of 49 %. the shrub dichrostachys cinerea has the highest density (8000 ind/ha), followed by the shrubs grewia flavescens (5750 ind/ha) and grewia flava (4125 ind/ha). 4.2 sclerocarya birrea-panicum maximum woodland. this is a small sub-community occupying the southwestern corner of the study area (fig. 3). this woodland covers 1 % of the study area (5 ha) and 3 % of community 4. the area is steep and has a large cliff-like rock face forming part of its eastern boundary. there are many signs of wild animal activity and several animal tracks traverse this sub-community. rockiness and erosion are estimated at 30 % and 10 %, respectively. soils are predominantly sandy and shallow. the grass cymbopogon excavatus and the herbaceous shrublet pelargonium sp. (species group i) are diagnostic for this subcommunity. the vegetation is dominated by the trees sclerocarya birrea (species group o) and combretum apiculatum (species group p). the shrub grewia flavescens (species group q) and the grass elionurus muticus (species group q) are also very prominent within this sub-community. the tree acacia exuvialis (species group a) is present. in this sub-community, the average number of plant species is 15 per 400 m². the tree layer has a 30–80 % cover (average 47 %); the shrub layer has a 5–80 % cover (average 42 %); the herb layer has 0–1 % cover (average 1 %); and the grass layer has a 40–90 % cover (average 70 %). the tree combretum apiculatum has the highest density (5000 ind/ ha), followed by the tree sclerocarya birrea (2333 ind/ha) and the shrub grewia flavescens (2333 ind/ha). barrett.qxd 2006/04/17 01:47 pm page 61 koedoe 49/1 (2006) 62 issn 0075-6458 4.3 combretum zeyheri-panicum maximum woodland. this sub-community is found in the south western section of the study area (fig. 3). this woodland covers 6 % of the study area (51 ha) and constitutes 32 % of community 4. the area is moderate to steep in the south, gradually becoming moderate towards the north. a few small dwellings are found within this sub-community. there are many signs of wild animal activity and some signs of previous human agricultural disturbance. old farm implements are lying in the veld and small unused man-made dams are found. open areas, with stands of similar-sized and aged dichrostachys cinerea shrubs occur throughout, indicating overgrazing by cattle. rockiness and erosion are estimated at 45 % and 10 %, respectively. soils are sandy and shallow in the south, becoming deeper towards the north. the trees acacia caffra and combretum zeyheri and the shrub ptaeroxylon obliquum (species group j) are diagnostic for this subcommunity. the vegetation is dominated by the trees sclerocarya birrea (species group o) and combretum apiculatum (species group p) and the shrub grewia flavescens (species group q). the shrub dichrostachys cinerea (species group p) and the grass elionurus muticus (species group q) are also prominent within this sub-community. there is also a strong presence of the tree bridelia mollis (species group d). the average number of plant species in this sub-community is 17 per 400 m². the tree layer has a 70–80 % cover (average of 73 %); the shrub layer has a 15–70 % cover (average of 35 %); the herb layer has a 0–1 % cover (average of 1 %); and the grass layer has a 1–80 % cover (average of 29 %). the tree combretum apiculatum has the highest density (4500 ind/ha), the shrub dichrostachys cinerea has 3833 ind/ha, and the shrub grewia flavescens has 3500 ind/ha. 4.4 asparagus setaceus-philenoptera violacea woodland. this woodland sub-community consists of two variants, the pappea capensis variant located in the south western section of the study area, and the diospyros mespiliformis variant located in the north eastern section (fig. 3). this sub-community covers approximately 8 % of the study area (61 ha) and 38 % of community 4. the terrain ranges from steep (the pappea capensis variant) to gentle (the diospyros mespiliformis variant). rockiness for this sub-community varies between 10–50% and erosion is 10 %. soils are predominantly sandy and well drained. the trees philenoptera violacea, combretum hereroense and combretum molle, the shrub dovyalis caffra, and the herbs asparagus setaceus and tacazzea apiculata (species group k) are diagnostic for this sub-community. the vegetation is dominated by the shrub grewia flavescens (species group q). the shrub grewia flava (species group q) and the grass elionurus muticus (species group q) are also prominent within this sub-community. there is also a strong presence of the tree berchemia zeyheri (species group e). there are signs of previous and current human disturbance in the form of old cattle dipping structures, old cattle kraals, recent vegetation removal for development, a few recently constructed log cabins and a new earthen dam. 4.4.1 pappea capensis variant this variant is located on the north-facing midslopes and footslopes of the drakensberg mountains in the southeastern section of the study area and divides the balanites maughamii-panicum maximum woodland subcommunity (4.1) into an eastern and western section (fig. 3). this variant covers 1 % of the study area (7 ha) and 11 % of subcommunity 4.4, and is found on the moderate to steep midslopes and footslopes of the drakensberg mountain range. signs of human disturbance include previous cattle barrett.qxd 2006/04/17 01:47 pm page 62 issn 0075-6458 63 koedoe 49/1 (2006) dipping structures and kraals. rockiness is estimated at 50 % and erosion is estimated at 10 %. soils are sandy. the tree pappea capensis and the shrubs bolusanthus speciosus, euclea crispa, ehretia rigida and grewia monticola (species group l) are diagnostic for this variant. the vegetation is dominated by the tree combretum apiculatum (species group p) and the shrub grewia flavescens (species group q). the tree commiphora mollis (species group e) and the grass elionurus muticus (species group q) are also prominent. the trees ziziphus mucronata (species group p), ficus stuhlmannii (species group d) and berchemia zeyheri (species group e); and the shrubs dichrostachys cinerea (species group p), grewia flava (species group q) and euclea divinorum (species group e) are also present. fig. 4. community ordination scatter diagrams depicting a) tree cover, shrub cover, herb cover, and grass cover with trend lines portraying tendencies in the aforementioned parameters from community 1 through community 4; and b) slope, erosion and rockiness with trend lines depicting inclinations of aforementioned parameters from community 1 through community 4. barrett.qxd 2006/04/17 01:47 pm page 63 koedoe 49/1 (2006) 64 issn 0075-6458 the average number of plant species/400 m² is 22. the tree layer covers 70–90 %, the shrub layer 15–60 %; the herb layer 2–5 %; and the grass layer 20–70 % of the area. the shrub species grewia flavescens has the highest density of 2500 ind/ha, the tree species combretum apiculatum having 1750 ind/ha and the shrub species grewia flava with 1500 ind/ha. 4.4.2 diospyros mespiliformis variant this variant is located in the north eastern section of the study area (fig. 3), covering 7 % of the study area (54 ha) and 89 % of subcommunity 4.4. slope is gentle. a dry river bed with several large riverine trees growing adjacent to the river bed runs through the area. there are many signs of recent human disturbance throughout this variant. several small wooden cabins and an earthen dam have recently been constructed along the river bed due to the shady and aesthetic nature of the area. a portion of the variants vegetation has also been removed for development. average erosion for this community is estimated at 10 %, with average rockiness also estimated at 10 %. soils are deep, sandy and well drained. species belonging to species group m are diagnostic for this variant and include: the trees diospyros mespiliformis, acacia schweinfurthii and dombeya rotundifolia; the shrubs indigofera arrecta and morella pilulifera; the herb asparagus virgatus; and the liana dalbergia armata. the vegetation is dominated by the shrubs grewia flavescens and grewia flava (species group q). the tree berchemia zeyheri (species group e) is also prominent. the grass layer is not well developed and includes the grasses panicum maximum (species group f) and elionurus muticus (species group q). the average number of plant species is 16/400 m². the tree layer has an 80–90 % cover with an average of 87 %; the shrub layer has a 30–60 % cover with an average of 40 %; the herb layer has a 2–3 % cover with an average of 2 %; and the grass layer has a 0–5 % cover with an average of 2 %. the shrub species grewia flava has the highest density of 2000 ind/ha, the tree species berchemia zeyheri has 2000 ind/ha, while fig. 5. community utilisation by vervets for wet and dry seasons in the blydeberg conservancy. barrett.qxd 2006/04/17 01:47 pm page 64 issn 0075-6458 65 koedoe 49/1 (2006) the tree species diospyros mespiliformis has 1667 ind/ha. ordination figure 4 depicts community vegetation parameters along gradients of slope, erosion and rockiness. there is a general increase in tree cover from community 1 through 4, with the exception of community 3 which occurs on a rocky protrusion with shallow soils which is more conducive to grasses. an increase in tree cover could be attributed to a decline in erosion and rockiness from community 1 through to community 4, reaching their respective troughs in community 3 due to the rocky protrusion. shrub and herb cover decrease from community 1 through 3, but increase slightly in community 4; decreases could be related to a trend of increasing tree cover which blocks sunlight and out competes non-woody vegetation. increases in slope and decreases in erosion and rockiness (with the exception of community 3 which occurs on a rocky outcrop as mentioned previously) from community 1 through 4 might also be partially responsible for increases in shrub cover for community 4. grass cover generally increases with increasing slope and decreasing erosion and rockiness from community 1 through 4, with a slight exception for community 2 which could be attributable to previous mismanagement and overgrazing. plant community selection of the ten available plant communities identified within their home range, the vervets only utilised the following six: community 2.2 made up 55.1 % of the study area. during the wet season vervets utilised 0.1 % of the community compared to the 1.6 % in the dry season. community 3 made up 14.0 % of the study area. during the wet season vervets did not use it and during the dry season they utilised only 0.2 %. community 4.1 made up 12.5 % of the study area. during the wet season vervets utilised 1.1 % of the community and during the dry season they utilised 3.0 %. community 4.2 made up 1.5 % of the study area. during the wet season vervets utilised 13.4 % of the community and during the dry season 13.1 %. community 4.3 made up 14.9 % of the study area. during the wet season vervets utilised 14.2 % of the community versus 13.1 % during the dry season. community 4.4.1 made up 2.0 % of the study area. vervets did not use this community during the wet season but during the dry season they utilised 1.2 %. figure 5 depicts community utilisation across seasons for the study area and study period. community utilisation was not significantly different for wet and dry seasons (u = 23.000, p = 0.4233). discussion and conclusions there are relatively clear distinctions between the various communities identified at the blydeberg conservancy. the braunblanquet approach proved to be an accurate and effective way whereby floristicallydefined plant communities could be classified and identified in the field. all the plant communities identified in the study area reflect some form of human related impact, mostly due to agricultural practices. with the exception of the diospyros mespiliformis variant (4.4.2) all the plant communities identified are in the process of self rectification to a more stable successional stage. in the diospyros mespiliformis variant (4.4.2) there are recent signs of largescale vegetation destruction through clearing actions by humans. as this is not consistent with the definition and objectives of a conservancy, i.e., the conservation of the natural resources occurring within the area, this would therefore need to be addressed by the members of the conservancy. a close association between the major plant communities and the different land types has been observed in this study. this is in accordance with the findings of kooij et al. (1990), bezuidenhout (1993), eckhardt (1993) and brown (1997). barrett.qxd 2006/04/17 01:47 pm page 65 koedoe 49/1 (2006) 66 issn 0075-6458 many of the low-lying areas within the study area have dense woodlands and a poorly developed herbaceous layer, which could be ascribed to previous long term overgrazing by cattle. the tree densities of all the low lying areas (communities 1.1; 1.2; 2.1; 2.2) are more than 5 000 ind/ha which are higher than the threshold value of 1500–1700 ind/ha after which veld condition will decline (orban 1995). overgrazing by cattle has compromised the herbaceous layer by removing the natural tree/grass competition factor resulting in areas that appear to be in a semi-degraded condition. a decline in the condition of the grass or herbaceous layer is typically accompanied by an increase in the density of trees and shrubs (smit et al. 1999). reduced competition from grass communities in overgrazed areas leads to an increase in the density of woody species. in most cases these changes are irreversible due to woody species out-competing grasses for moisture. over time, fuel loads are reduced in areas with a degraded grass layer and fire can no longer be used as an effective means of bush control. since vervets mostly feed in forest/woodland areas, these areas also become more suitable to these species. this study forms part of a larger study on the foraging ecology of a single troop of vervet monkeys in mixed lowveld bushveld and sour lowveld bushveld of the blydeberg conservancy. to date there are no formal management plans for vervet monkeys. this is attributed to the limited knowledge of vervets and their utilisation of and impacts on ecosystems. the study troop’s diet consisted of a number of staple food items, with a relatively wide seasonal variation for additional supplementation (barrett 2005). there was an abundant supply of various food sources throughout the year, with movement patterns mostly coinciding with the fruiting times of several tree and plant species. home range size appeared to be linked to food and particularly water availability, with the troops’ mean daily distance travelled increasing slightly during the dry season (barrett 2005). with regards to habitat utilisation, the vervets had ten plant communities available to them within their home range. of the ten available plant communities, they utilised only six during the study period namely sub-community 2.2 (combretum imberbe-acacia nigrescens woodland), community 3 (acacia nigrescens-combretum apiculatum woodland), sub-community 4.1 (balanites maughamii-panicum maximum woodland), sub-community 4.2 (sclerocarya birrea-panicum maximum woodland), sub-community 4.3 (combretum zeyheri-panicum maximum woodland), and the pappea capensis variant (4.4.1) (barrett 2005). of the six communities utilised, community 4.2 was used the most during both the wet and the dry season. the acacia nigrescens-combretum apiculatum woodland (community 3) and the pappea capensis variant (4.4.1) were only used during the dry season, whereas the other communities were randomly utilised across both seasons (barrett 2005). no similar vegetation descriptions have previously been completed on the vegetation of the study area and this research therefore provides valuable data on the ecosystems present. previous vegetation studies for the region were conducted on broader plant communities (deall 1985; deall et al. 1989; matthews 1991; matthews et al. 1994; marais 2005). data obtained from this initial assessment could be incorporated into a larger vegetation map for the entire blydeberg conservancy. setting up goals and objectives for the management of the area would be essential to its continued existence over the medium to long term and should be seen as a priority by all involved. by establishing a set of minimum objectives, a baseline management plan can be developed where the natural vegetation is managed and protected. a veld management and monitoring program needs to be implemented as a priority so that a thorough knowledge of the vegetation; the variability thereof; carrying capacities; quality of veld; reaction of the vegetation to various ecological impacts such as burning, grazing, browsing, bush clearing, invader species etc. and the growth processes of plants in different stages of their life cycles can be understood and managed to the benefit of all animals and humans in the area. barrett.qxd 2006/04/17 01:47 pm page 66 issn 0075-6458 67 koedoe 49/1 (2006) acknowledgements professor d. mitchell is thanked for allowing the research to take place on his privately owned portion of the blydeberg conservancy. this research was financially supported by the national research foundation. references arnold, t.h. & b.c. de wet. 1993. plants of southern africa: names and distribution. memoirs of the botanical survey of south africa 62: 1-825. barbour, m.g., j.h. burk & w.d. pitts. 1987. terrestrial plant ecology. 2nd ed. massachusetts: benjamin/cummings. barrett, a.s. 2005. foraging ecology of the vervet monkey (chlorocebus aethiops) in mixed lowveld bushveld and sour lowveld bushveld of the blydeberg conservancy, northern province, south africa. m.tech. nature conservation dissertation. university of south africa, pretoria. bezuidenhout, h. 1993. syntaxonomy and synecology of western transvaal grasslands. ph.d. dissertation, university of pretoria, pretoria. bezuidenhout, h. 1996. the major vegetation communities of the augrabies falls national park, northern cape. 1. the southern section. koedoe 39: 7-24. booth, a.h. 1979. the distribution of primates in the gold coast. pp. 139-154. in: sussman, r.w. (ed.). primate ecology. problem-oriented field studies. new york: wiley bredenkamp, g.j. & h. bezuidenhout. 1995. a proposed procedure for the analysis of large data sets in the classification of south african grasslands. koedoe 38(1): 33-39. bredenkamp, g.j., a.f. joubert & h. bezuidenhout. 1989. a reconnaissance survey of the vegetation of the plains in the potchefstroomfochville-parys area. south african journal of botany 55: 199-206. brown, l.r. 1997. a plant ecological and wildlife management plan of the borakalalo nature reserve, north-west province. ph.d. dissertation. university of pretoria, pretoria. brown, l.r. & g.j. bredenkamp. 1994. the phytosociology of the southern section of the borakalalo nature reserve, south africa. koedoe 37: 59-72. brown, l.r. & g.j. bredenkamp & n. van rooyen. 1996. the phytosociology of the northern section of the borakalalo nature reserve. koedoe 39(1): 9-24. buckle, c. 1992. landforms in africa: an introduction to geomorphology. essex: longman. caughley, g. & a.r.e. sinclair. 1994. wildlife ecology and management. canada: oxford university press. cilliers, s.s. 1998. phytosociological studies of urban open spaces in potchefstroom, north west province, south africa. ph.d. thesis, potchefstroom university for che, potchefstroom. deall, g.b. 1985. a plant-ecological study of the eastern transvaal escarpment in the sabie area. m.sc. thesis, university of pretoria, pretoria. deall, g.b., g.k. theron, r.h. westfall. 1989. the vegetation ecology of the eastern transvaal escarpment in the sabie area. 2. floristic classification. bothalia 19: 69-89. eckhardt, h.c. 1993. a synecological study of the vegetation of the north-eastern orange free state. m.sc. thesis. university of pretoria, pretoria. edwards, d. 1983. a broad-scale structural classification of vegetation for practical purposes. bothalia 14: 705-712. fedigan, l. & l.m. fedigan. 1988. cercopithecus aethiops: a review of field studies. pp 389-411. in: gautier-hion a., bourlière f., gautier j., kingdon j. (eds.). a primate radiation: evolutionary biology of the african guenons. new york: cambridge university press. fuls, e.r. 1993. vegetation ecology of the northern orange free state. ph.d. dissertation. university of pretoria, pretoria. fuls, e.r., g.j. bredenkamp & n. van rooyen. 1992. the plant communities of the undulating grassland of the vredefort–kroonstad–lindley– heilbron area, northern orange free state. south african journal of botany 58: 224-230. harrison, m.j.s. 1983. patterns of range use by the green monkey, cercopithecus sabaeus, at mt. assirik, senegal. folia primatol 41: 157-179. harrison, m.j.s. 1984. optimal foraging strategies in the diet of the green monkeys, cercopithecus sabaeus, at mt. assirik, senegal. international journal of primatology 5: 435-471. hennekens, s.m. 1996a. turboveg: a software package for input, processing and presentation of phytosociological data. user’s guide, version july 1996, ibn-dlo. lancaster: wageningen & lancaster university. hennekens, s.m. 1996b. megatab: a visual editor for phytosociological tables. ulft: giesen. hill, m.o. 1979. twinspan: a fortran program for arranging multivariate data in an ordered twoway table by classification of individuals and attributes. new york: cornell university. isbell, l.a., j.d. pruetz & t.p. young. 1998. movements of vervets (cercopithecus aethiops) and patas monkeys (erythrocebus patas) as estimators of food resource size, density and distribution. behavioral ecology and sociobiology 42: 123-133. barrett.qxd 2006/04/17 01:47 pm page 67 kent, m. & p. coker. 1997. vegetation description and analysis a practical approach. new york: wiley. kingdon, j. 1997. the kingdon field guide to african mammals. new york: naturalworld. kooij, m.s., g.j. bredenkamp & g.k. theron. 1990. classification of the vegetation of the b land type in the north-western orange free state. south african journal of botany 56: 309318. krebs, c. j. 1989. ecological methodology. new york: harper and row. land type survey staff. 1989. land types of the maps 2330 tzaneen, 2430 pilgrims rest. memoirs on the agricultural natural resources of south africa 12. lee, p.c. & m.d. hauser. 1998. long-term consequences of changes in territory quality on feeding and reproductive strategies of vervet monkeys. journal of animal ecology 67: 347-358. low, a.b. & a.g. rebelo (eds.). 1998. vegetation of south africa, lesotho and swaziland. 2nd ed. pretoria: department of environmental affairs and tourism. marais, a.j. 2005. resource utilisation of the chacma baboon in different vegetation types in northeastern mountain sour veld, blyde canyon nature reserve. m.tech. (nature conservation) dissertation. university of south africa, pretoria. matthee, j.f. & c.j. van schalkwyk. 1984. a primer on soil conservation. pretoria: government printer. (department of agriculture bulletin; no. 399.) matthews, w.s. 1991. phytosociology of the northeastern mountain sourveld. msc. thesis, university of pretoria, pretoria. matthews, w.s., g.j. bredenkamp & n. van rooyen. 1994. the phytosociology and syntaxonomy of relative low-altitude areas in the north-eastern mountain sourveld, in the eastern transvaal escarpment region. koedoe 37(2): 73–87. mueller-dombois, d. & h. ellenberg. 1974. aims and methods of vegetation ecology. new york: wiley. oates, j.f. 1996. african primates status survey and conservation action plan. gland, switzerland: iucn. (iucn/ssc primate specialist group. s.i.) orban, b. 1995. an ecological management plan for the lionspruit game reserve. m.sc (wildlife management) dissertation. university of pretoria, pretoria. pruetz, j.l. & l.a. isbell. 2000. correlations of food distribution and patch size with agonistic interactions in female vervets (cercopithecus aethiops pygerythrus senso lato) and patas monkeys (erythrocebus patas) living in simple habitats. behavioral ecology and sociobiology 49: 38-47. skinner, j.d. & r.h.n. smithers. 1990. the mammals of the southern african subregion. 2nd ed. south africa: university of pretoria. smit, g.n., c.g.f. richter & a.j. aucamp. 1999. bush encroachment: an approach to understanding and managing the problem. pp. 246-260. in: tainton r.m. (ed.). veld management in south africa. pietermaritzburg: university of natal press. struhsaker, t.t. 1967. ecology of vervet monkeys (cercopithecus aethiops). ecology 48: 891-904. tueller, p.t. 1988. vegetation science applications for rangeland analysis and management. dordrecht: kluwer academic publishers. tyson, p.d. & r.a. preston-whyte. 2000. the weather and climate of southern africa. cape town: oxford university press. van der schijff, h.p. 1963. a preliminary account of the vegetation of the mariepskop complex. transvaal provincial administration. fauna and flora 14: 42-53. van der schijff, h.p. & e. schoonraad. 1971. the flora of the mariepskop complex. bothalia 10(3): 461-500. van rooyen, n., g.k. theron & n. grobbelaar. 1981. a floristic description and structural analysis of the plant communities of the punda milia-pafuri-wambiya area in the kruger national park, republic of south africa. 1. the hygrophilous communities. south african journal of botany 47: 213-246. van zyl, d. 2003. south african weather and atmospheric phenomena. pretoria: briza. visser, d.j.l. (ed.). 1989. the geology of the republics of south africa, transkei, bophuthatswana, venda and ciskei and the kingdoms of lesotho and swaziland. geological survey, explanation of the 1: 1 000 000 geological map. 4th ed. pretoria: government printer. walraven, f. 1989. the geology of the pilgrim’s rest area. geological survey, explanation of sheet 2430. pretoria: government printer. whitten, p.l. 1988. effects of patch quality and feeding subgroup size on feeding success in vervet monkeys (cercopithecus aethiops). behaviour 105: 35-52. koedoe 49/1 (2006) 68 issn 0075-6458 barrett.qxd 2006/04/17 01:47 pm page 68 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <feff0041006e007600e4006e00640020006400650020006800e4007200200069006e0073007400e4006c006c006e0069006e006700610072006e00610020006e00e40072002000640075002000760069006c006c00200073006b0061007000610020005000440046002d0064006f006b0075006d0065006e00740020006d006500640020006800f6006700720065002000620069006c0064007500700070006c00f60073006e0069006e00670020006600f60072002000700072006500700072006500730073007500740073006b0072006900660074006500720020006100760020006800f600670020006b00760061006c0069007400650074002e0020005000440046002d0064006f006b0075006d0065006e00740065006e0020006b0061006e002000f600700070006e006100730020006d006500640020004100630072006f0062006100740020006f00630068002000520065006100640065007200200035002e003000200065006c006c00650072002000730065006e006100720065002e00200044006500730073006100200069006e0073007400e4006c006c006e0069006e0067006100720020006b007200e400760065007200200069006e006b006c00750064006500720069006e00670020006100760020007400650063006b0065006e0073006e006900740074002e> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice filelist convert a pdf file! page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 article information author: marc stalmans1 mike peel2 affiliation: 1international conservation services cc, nelspruit, south africa 2agricultural research council, nelspruit, south africa correspondance to: marc stalmans email: stalmans@ics-consulting.co.za postal address: international conservation services cc, po box 19139, nelspruit 1200, south africa keywords greater limpopo transfrontier park (gltp); parque nacional de zinave (pnz); save river; sandveld; mopane; miombo dates: received: 14 apr. 2009 accepted: 11 oct. 2009 published: 11 mar. 2010 how to cite this article: stalmans, m. & peel, m., 2010, ‘plant communities and landscapes of the parque nacional de zinave, mozambique’, koedoe 52(1), art. #703, 11 pages. doi: 10.4102/koedoe.v52i1.703 copyright notice: © 2010. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0771 (online) plant communities and landscapes of the parque nacional de zinave, mozambique in this original research article... open access • abstract • introduction • study area • method    • vegetation sampling    • analysis    • delineation of landscapes • results and discussion    • causal factors of vegetation pattern in the pnz    • twinspan classification    • definition of plant communities       • plant communities       • definition of landscapes       • landscapes of the pnz       • accuracy of the landscape map       • plant community diversity    • land-use patterns and their influence on the vegetation of the pnz • acknowledgements • references abstract (back to top) the parque nacional de zinave covers 400 000 ha in mozambique to the south of the save river. until recently, this park had been characterised by neglect and illegal hunting that caused the demise of most of its large wildlife. a recent initiative has been launched that aims at rehabilitating the park within the scope of the greater limpopo transfrontier park (gltp). a vegetation map was required as input to its management plan. the three primary objectives of the study were, firstly, to understand the environmental and biotic determinants of the vegetation, secondly, to identify and describe individual plant communities in terms of species composition and structure along the roads in the study area and, thirdly, to delineate landscapes in terms of their plant community make-up, environmental as well as biotic determinants and distribution. this is the third survey and description of the landscapes of the national parks located in the mozambique part of the gltp. a combination of fieldwork and analysis of landsat satellite imagery was used. a total of 75 sample plots were surveyed. a brief subjective visual assessment was undertaken for another 114 sample points. the accuracy of the landscape map was evaluated by means of 582 points assessed during an aerial game count. the ordination results clearly indicate the overriding importance of moisture availability in determining vegetation composition. ten distinct plant communities were recognised. different combinations of these plant communities can be grouped in six major landscapes, namely the save river channel and river banks, save riverine forest, acacia nigrescens woodland landscape, mopane landscape, miombo landscape and sandveld landscape. the landscapes with their individual plant communities represent habitats that are highly suitable for the reintroduction of many game species that were lost during the latter part of the last century. conservation implication: no formal description and mapping of the vegetation existed for zinave. the landscape map is a vital input for the management plan. the reintroduction of wildlife species that were exterminated during the civil war requires a selection of suitable habitats for the placement of the ‘sanctuary’ that will be used to ensure the initial security of the introduced animals. the landscape map of zinave fits into the broader mapping of the great limpopo transfrontier park and conservation area (gltfca) for conservation planning purposes. introduction (back to top) the parque nacional de zinave (zinave national park; pnz) has been said to comprise some of the best wildlife habitats in southern africa (duarte & oreste 1999; storer & dalquest 1967) and used to harbour a wide diversity of wildlife (see table 1). it was initially declared a hunting concession in 1962 and was run by mozambique safarilandia. it was upgraded to national park status in 1972. it covers some 400,000 ha in mozambique to the south of the save river and forms one of the major components of the greater limpopo transfrontier park (gltp) (see figure 1). a vegetation map was required as one of the essential building blocks for the drafting of its management plan (dnac 2002) and as a critical element for the planning of management actions. few botanical surveys were carried out in mozambique between 1980 and 1994, during the long period of internal conflict that affected much of the country (micoa 1997). the extent of the area, its relative inaccessibility due to poor road infrastructure and the limited time available precluded a traditional fine-scale vegetation description and map of the pnz. vegetation is often used as a surrogate or building block for the definition of habitats (timberlake, nobanda & mapaure 1993). the use of broad habitat units defined by a combination of environmental factors and vegetation would probably represent the most useful input for the drafting of the management plan. this is because such units are used differently by different species of wildlife. the units also usually differ in their ecological capacity and water availability and thus might require different fire regimes. moreover, the units tend to differ in their sensitivity to utilisation and development. this is the third survey and description of the landscapes of the national parks located in the mozambican part of the gltp. this follows the description of the landscapes of the limpopo national park (lnp) (stalmans, gertenbach & carvalho-serfontein 2004) and those of the banhine national park (bnp) (stalmans & wishart 2005). difficult access, limited resources and a lack of previously published information have severely limited the sampling intensity, the extent of the descriptions and the detail of the mapping. detailed vegetation sampling was, therefore, restricted to the areas adjacent to roads. nevertheless, it is hoped that the initial published accounts of the vegetation and landscapes of these parks will provide a basis on which further research projects can expand, much as the level of knowledge of the kruger national park (knp) has been improved upon over the years. similar landscape mapping exercises have been undertaken in the knp (gertenbach 1983) and in the protected areas to the west of the knp (peel, kruger & macfadyen 2007). a similar conceptual approach to the lnp and bnp studies was followed, whereby landscapes were mapped explicitly, whereas the embedded plant communities are implied. a landscape is defined as ‘an area with a specific geomorphology, climate, soil vegetation pattern and associated fauna’ (gertenbach 1983). the objectives of this study were, firstly, to understand the environmental determinants of the vegetation, secondly, to identify and describe individual plant communities in terms of species composition and structure and, thirdly, to identify and delineate landscapes in terms of their plant community make-up, environmental determinants and occurrence. study area (back to top) the physical environment largely determines the vegetation composition and structure. the pnz is situated between latitudes 21°21’ s to 21°54’ s and longitudes 33°2’ e to 34°2’ e, in the inhambane province of mozambique (see figure 1), encompassing an area of approximately 400 000 ha. the pnz forms part of the gltp with the lnp to the west and the bnp to the southwest. the highest elevation is on the southeastern boundary at 174 m above sea level. the area dips generally in a north-northeasterly direction with the lowest point at 110 m elevation on the northern boundary along the save river. the area is underlain by arenaceous and argillaceous sediments with alluvium along the save river (departemento terra e agua 1995). the pnz is mostly covered with sandy soils. clayey soils derived from alluvial sediments occur along the save river. these soils are susceptible to salination. the area is drained by numerous seasonal streams that run in an east-northeasterly direction towards the perennial save river. a number of seasonal pans (some of which are quite large) are scattered across the park. a mean annual rainfall value of 690 mm applies to the northeastern sector of the znp, 613 mm in the southeast and 571 mm in the western sector, using the grid at 0.5 degree longitude/latitude resolution of the leemans and cramer (1991) database. considerable variations can be expected within and between seasons (kelly & walker 1976). the pnz is located on the interface between the mopane and the miombo of the sudano-zambezian region, as described by werger and coetzee (1978) and bandeira, hatton, munisse and izidine (1994). the pnz was established specifically for the conservation of important species such as giraffe (presently extinct) that historically only occurred in mozambique south of the save river (smithers & lobão tello 1976). most of the original diverse and numerous large herbivores were largely decimated during the long civil war and its aftermath (see table 1). the pnz supports the subsistence of an estimated 4 200 people in about 700 households located within the official park boundaries (dnac 2002). figure 1: locality map of the parque nacional zinave in the greater limpopo transfrontier park method (back to top) vegetation sampling as access to the area can be very difficult due to flooding during summer, vegetation sampling was carried out at the end of the summer season and into the winter of june 2003. although this is not ideal in terms of plant species identification, it was unavoidable. a total of 75 plots of 40 m x 40 m (160 m2) were subjectively located in representative stands of vegetation to cover the altitudinal variation, soils and terrain position as determined from the topographical map and landsat imagery. sampling was limited to the areas adjacent to the road network during the ground survey (see figure 2). edwards (1983) structural classification was used to describe the overall structural properties of the sampled plots. overall cover was estimated for the woody, grass, forb and geophyte component respectively using the semi-quantitative measures of the braun‑blanquet approach (mueller-dombois & ellenberg 1974). cover and height classes were recorded for individual woody and grass species. records of environmental data included gps position, geology, landscape position (land type survey staff 1989), slope, soil texture (using the sausage method; national working group for vegetation ecology 1986) and rockiness. in order to further increase sampling intensity, 114 ground observation points were also used (see figure 2). these consisted of a gps point and subjective visual assessment relative to the formally surveyed sites. in 2007, during an aerial game count (stalmans & peel 2007), a further 582 gps points were assessed to improve the landscape map (see figure 2). figure 2: position of vegetation sample plots as well as ground and aerial control points in the parque nacional zinave analysis data were analysed through a combination of classification and ordination techniques. classification was used to identify groups and to give structure to the raw data. ordination of data was aimed at arranging species and samples in a low-dimensional space so that similar entities were close to one another and dissimilar entities far apart. although more contemporary and sophisticated software packages are available for vegetation classification, the same techniques were used as those applied in the lnp (stalmans et al. 2004) and bnp studies (stalmans & wishart 2005) in order to maintain a uniform approach that is ultimately aimed at providing a single description of the entire gltp. a twinspan (two-way indicator species analysis) classification (hill 1979) was performed on the 75 sample plots. this is a polythetic divisive technique that is based on reciprocal averaging ordination (gauch 1982). the simplicity and robustness of twinspan as a hierarchical technique were appealing within the constraints of this particular study. it results in the definition of communities each characterised by its own distinctive species combination. the canoco package (ter braak 1992) was selected to analyse relationships between the data set of 75 plots and 168 species and the underlying environmental factors. canoco allows for canonical ordination, which is an intermediate technique that combines aspects of regular ordination with aspects of regression (jongman, ter braak & van tongeren 1987). the resulting ordination diagram expresses not only the pattern of variation in species composition but also the main features of species distributions along the gradient of environmental variables (ter braak 1986). canonical coefficients, t-values of the regression coefficients and inter-set correlations of the environmental variables with axes are presented in the output of the canonical correspondence analysis (cca). canonical coefficients in a cca are the equivalent of regression coefficients in indirect gradient analysis. they are coefficients of a weighted multiple regression of the sample scores on the standardised environmental variables. they have a larger variance than regression coefficients. when the t-value of a variable is less than 2.1, the variable does not contribute much to the fit of the species data in addition to the contributions of the other variables in the analysis. the inter-set correlations of environmental variables with axes are the correlation coefficients between the environmental variables and the species axes consisting of the sample scores. delineation of landscapes the relevant landscapes for the pnz were identified using the knowledge gained through the ordination and classification of the pnz field data. a combined approach was used to map these landscapes using the available soils map, the existing land cover map and the landsat image. the land cover map (carta de uso e cobertura da terra) has been derived for mozambique from landsat satellite imagery at a scale of 1:250 000 (dinageca 1999). the pnz comprises a total of 166 land cover polygons. a ‘best fit’ map of the landscapes was subjectively drawn by matching the individual land cover polygons to the field sampling data, the 114 ground observation points and the 582 aerial observations. where required, polygon boundaries were manually edited following visual inspection of the satellite image and the recorded occurrence of the landscapes during the aerial survey. the coverage of the polygons by the available data set was assessed in order to derive a measure of certainty or uncertainty regarding the quality of the final landscape map. results and discussion (back to top) causal factors of vegetation pattern in the pnz the first ordination run with the full set of 75 vegetation plots resulted in three distinct ‘clouds’ of sample plots in ordination space (see figure 3). these ‘clouds’ represent respectively sample plots in riverine, floodplain or upland positions. the environmental variables ‘drainage line’ had a canonical coefficient of 0.97 on the second axis and ‘floodplain’ had a high canonical coefficient of 0.89 on the first axis. the different sample scores for these nominal environmental variables are represented within the ordination diagram by a single point, called the ‘centroid’.the 11 sample plots associated with the floodplain and drainage lines were removed from the data set and a new ordination was run. the resulting diagram identifies two outlying plots on the right of the diagram (see figure 4). these two outliers represent two previously cultivated areas. these old lands are characterised by a combination of melinis repens, perotis patens and tricholaena monachme in the grass layer. the parameter ‘old land’ has a canonical coefficient of 0.76 and t-value of 12.1 for the first axis. the parameter ‘soil texture’ has a canonical coefficient of 0.87 and t-value of 9.8 for the second axis. the sample plots are clearly spread in the diagram along the second axis with sandy substrates in the bottom-left corner (including miombo with androstachys johnsonii) and more clayey soil (acacia nigrescens woodlands) in the top-left corner of the diagram. the successive ordination runs clearly indicate the overriding importance of moisture availability (in conjunction with soil texture) in determining vegetation composition in the pnz. firstly, the obvious differences in soil moisture availability by virtue of sample positions along drainage lines and in the floodplain came out of the first ordination. thereafter, the gradient in soil clay content (as a result of underlying geological substrate and landscape position) and landscape position per se (in determining water flow) largely determine soil moisture and nutrient availability. the interplay of soil moisture and soil nutrient availability conforms to the current understanding of the determinants of savanna systems. the four-determinant model gives moisture and nutrient availability equal status in establishing the range of possible forms that a savanna can assume (scholes & walker 1993). fire and herbivory then determine the actual form and function within that range. similarly, timberlake et al. (1993) consider soil moisture in this environment a major determinant in the distribution of vegetation types. soil moisture availability results from the interaction between rainfall, topography, soil texture, soil depth, drainage and rooting habit of the plants. siebert, bredenkamp and siebert (2003) identified a gradient of decreasing soil moisture availability along the first axis of ordination of more than 2 000 sample plots in mopane veld straddling south africa, namibia and botswana. within the gltp, stalmans (1994), stalmans et al. (2004) and stalmans and wishart (2005) identified water availability (as controlled by the position of the sample in the landscape and by its soil texture) as the major determinant of vegetation composition in an area adjacent to the gonarezhou national park in zimbabwe, in the lnp and in the bnp respectively. figure 3: cca ordination biplot of 75 vegetation sample plots in the parque nacional zinave using nominal environmental variables, where triangle symbols represent the centroids of the environmental variables ‘floodplain’ and ‘drainage line’ position figure 4:cca ordination biplot of 64 vegetation sample plots in the parque nacional zinave, with the spread of samples aligned along a gradient of increasing clay content in the soil and where the cca excluded sample plots along major drainage lines, on floodplains and on termite mounds, by using one quantitative (soil texture) and one nominal (land-use history) environmental variable twinspan classification the results from the twinspan classification are presented by means of a hierarchical diagram (see figure 5). starting from the top of the diagram, the set of 75 sample plots was divided into two groups. the left group represents the bulk of the sample plots (n = 73). the right group of two sample plots has as indicator the synthetic species ‘no woody’, meaning that the group is characterised by an absence of trees and shrubs. this is the vegetation of seasonally inundated pans and grasslands (community 10) with as preferential species the grasses eragrostis cf. heteromera and paspalidium obtusifolium.< >a group of four plots at level 2 of the dendrogram represent acacia xanthophloea woodland with tall grasses. hyphaene petersiana and sporobolus consimilis are the indicator species (community 1). the remaining plots are split at level 3, in communities on fairly heavy soils (communities 2 to 5) and those on sandy soils (communities 6 to 9). the indicators for the left division (representing vegetation growing on clayey substrates) are urochloa mosambicencis, acacia nigrescens and colophospermum mopane. the indicators for the right group (indicating a habitat with sandy substrates) are hugonia orientalis and monodora junodii. the left group of level 3 (clayey substrates) is split into a riverine component (community 2) and a non-riverine assemblage at level 4. the indicator species for the non-riverine vegetation are urochloa mossabicensis, colophospermum mopane and digitaria eriantha. at level 5, the remaining plots on clayey soils are split into true mopane woodlands (with colophospermum mopane with a cover of 5% to 25% as indicator) (communities 3 and 4) and acacia nigrescens woodlands (community 5) (with acacia nigrescens and sclerocarya birrea as indicator). the right group of level 3 (sandy substrates) divides at level 4 into woodland and a near-forest (community 9) component. the near-forest component has acacia schweinfurtii as an indicator species and forest species such as cordyla africana and lecaniodiscus fraxinifolius as preferential species. the narrow fringe of vegetation found on the deep sands along the save river splits off the remaining sandy plots at level 5 (community 8) with ficus capreifolia being the indicator species. finally, the remaining sandy vegetation is split at level 6 into miombo (with julbernardia globiflora, heinsia crinita and hymenocardia ulmoides as indicator species) (community 6) and sandveld with guibourtia conjugata woodlands (community 7) (with guibourtia and alchornea laxiflora as indicator species). figure 5:hierarchical representation of the vegetation communities in the parque nacional zinave, where levels 1 to 6 indicate increasing levels of homogeneity and specific composition and indicator species, as illustrated by the twinspan classification, are underlined; communities that split off at an early level indicate a very distinctive and different assemblage from other communities definition of plant communities the community concept is applied in its broad sense and reflects a recurring assemblage of grass and woody species of characteristic composition and structure, growing in an area of essentially similar environmental conditions and land-use history (adapted from gabriel & talbot 1984). the classification outcome was evaluated subjectively against photographs of each sample plot. the main criterion applied was the need for each community to be identifiable in the field by an observer who is not necessarily a trained botanist. community names were chosen subjectively so as to have practical value in the field through the use of two species that are visually and/or diagnostically important. the 10 communities that were identified broadly conform to the lower divisions of the dendrogram. they are therefore discussed from left to right following figure 5. plant communities 1. hyphaene petersiana – acacia xanthophloea woodland: this community is situated to the south of the save river in its floodplain, mostly on fairly heavy clays. a large proportion of this community is probably seasonally inundated. this woodland is characterised by acacia xanthophloea, salvadora persica and hyphaene petersiana palms with a canopy height that is generally between 5 m and 10 m. the grass layer is very dense and tall with a high frequency of setaria incrassata and panicum maximum. in drier areas on the edge of this community, the vegetation becomes shorter, with the grass layer less dense and more saline. this community corresponds to the sporobolus consimilis – setaria incrassata tall grassland found in the lnp (stalmans et al. 2004). van rooyen, theron and grobbelaar (1981) similarly described sporobolus consimilis grassland for the limpopo floodplain in the knp. 2. acacia burkei – andropogon gayanus woodland: this community is found along seasonal rivers, occupying a narrow strip between 10 m and 20 m wide, along these drainage lines. the vegetation has affinities with the surrounding vegetation away from the drainage line and is therefore not homogeneous across its distribution range. the species composition is thus variable, making a characterisation on a purely plant composition basis difficult. acacia burkei and a. nigrescens are conspicuous as well as the grasses andropogon gayanus and eragrostis spp. it is the association of a wide range of species with the small drainage line conditions that sets this community apart. 3. xanthocercis zambesiaca – salvadora persica thickets: this community occurs as distinct bush clumps with a diameter of 20 m to 30 m on termitaria. these are embedded in the open acacia nigrescens woodlands of community 5. each clump is characterised by an emergent tree (often xanthocercis zambesiaca – nyala tree) that is surrounded by a variety of other species such as grewia bicolor, lecaniodiscus fraxinifolius, salvadora persica and maclura africana forming a ‘skirt’ around the core of the clump. this community is equivalent to the bush clumps found in the bnp that were assigned the same community name (stalmans & wishart 2005). the grass layer is poorly developed and comprises mostly panicum maximum. 4. colophospermum mopane – urochloa mosambicensis woodland: mopane is the dominant species in tall, closed woodland, which becomes low closed shrubland where drainage is impeded. other woody species have low cover values and include combretum apiculatum, dalbergia melanoxylon, drypetes mossambicensis, grewia bicolor and flueggea virosa. the grass layer is dominated by palatable grasses such as urochloa spp., heteropogon contortus, digitaria eriantha and panicum maximum. this community has much affinity with the similarly named mopane woodlands found in the bnp (stalmans & wishart 2005), the colophospermum mopane – panicum maximum woodland of the lnp (stalmans et al. 2004) and the digitaria milanjiana – colophospermum mopane vegetation type described by siebert et al. (2003). 5. acacia nigrescens – urochloa mosambicensis woodland: this is generally open (but sometimes closed) tall woodland with acacia nigrescens as an important and diagnostic species. other important woody species are sclerocarya birrea, combretum imberbe, c. apiculatum, c. hereroense and philenoptera violacea (previously lonchocarpus capassa). the grass layer is dominated by palatable urochloa spp., heteropogon spp., panicum maximum and digitaria eriantha. this community, which is found mostly in the northeastern part of the pnz, in a broad band between the save river in the north and the mopane woodlands of community 4 to the south, represents one of the most valuable grazing and browsing resources for wildlife in the park. 6. julbernardia globiflora – digitaria eriantha woodland: this is the typical ‘miombo’, consisting of a wide variation from julbernardia-dominated short closed woodlands to majestic brachystegia-dominated tall closed woodlands. characteristic ‘sandveld’ specialists such as hugonia orientalis, pteleopsis myrtifolia, cleistanthus schlechteri, hymenocardia ulmoides and xeroderris stuhlmannii are common. it includes patches of androstachys johnsonii. these are, however, not classified by the twinspan analysis as separate communities such as those found further west in bnp and the lnp. the androstachys patches are generally more species rich than further west. this species may be close to the edge of its distribution range as a result of increasing rainfall. this community also encloses numerous large termite mounds (5 m – 15 m diameter). these mounds are much more clayey than the surrounding sands on which the miombo is found. as a result species such as boscia foetida, manilkara mochisia and thilachium africanum occur here. in comparison, miombo was not found in either the lnp or bnp. 7. guibourtia conjugata – digitaria eriantha woodland: this community generally consists of fairly open, but sometimes closed, woodland with a very characteristic appearance due to the abundance and striking form of guibourtia conjugata and milletia stuhlmannii. other important woody species include strychnos madagascariensis, philenoptera bussei, alchornea laxiflora, brachystegia glaucescens and lannea schweinfurtii. panicum maximum is the most important grass after digitaria eriantha. this community has many affinities to the guibourtia conjugata – eragrostis pallens woodlands described in the bnp (stalmans & wishart 2005). 8. pluchea dioscoridis – digitaria eriantha thicket: this community occurs in a narrow strip on the banks of the save river where it is subject to flooding disturbance. a heterogeneous mix of species occurs with the shrub pluchea being the most frequent. other species include ficus capreifolia, ziziphus rivularis, grewia flavescens and g. sulcata. occasionally, large faidherbia albida form conspicuous copses on these banks. this community was relatively most affected by grazing and browsing by domestic goats. the regular deposition of fresh nutrients with the summer floods undoubtedly raises the quality of the grazing on these deep sands. 9. cordyla africana – philenoptera violacea forest: this closed community is found along the banks of the save river on levees and other stable substrates. this is a mature community that takes many years to develop. it is not prone to flood disturbance to the same degree as communities 1 and 8. it is characterised by a tall canopy between 10 m and 20 m in height with large cordyla africana, philenoptera violacea, sclerocarya birrea and trichilia emetica. the large-leaved grewia olukondae and the scrambling acacia schweinfurtii are diagnostic species for this community. species in zinave that are confined to this environment are craibia zimmermannii, cola natalensis, tapura fisheri and xylocarpus granatum. the grass layer consists mostly of panicum maximum with some setaria incrassata on the margins. 10. paspalidium obtusifolium open grasslands: these open grasslands are found in pans that are distributed across the different landscapes of the znp. the blue water lily, nymphaea nouchali, is often very conspicuous. the pans are often fringed by the tree combretum imberbe. this community is equivalent to the similarly named community recorded in the bnp (stalmans & wishart 2005). this represents the ramsar-defined wetland type ts: ‘seasonal/intermittent freshwater marshes/pools on inorganic soils; includes sloughs, potholes, seasonally flooded meadows, sedge marshes’. definition of landscapes not all of these 10 plant communities can be mapped individually, not even using the results of the satellite imagery analysis. they often occur in a fine-scaled mosaic that reflects small variations in topography and accompanying moisture regime. the current state of mapping of these environmental factors is not good enough to allow for the extrapolation and correlation of the vegetation into such fine-scaled mosaic. broader units therefore had to be mapped. landscapes have been defined that have bearing on management requirements and development potential as they represent a composite of topographical factors, underlying soil, hydrology and current vegetation pattern.different combinations of the plant communities can be grouped in six major landscapes (see figure 6). cultivated areas have been superimposed on the landscape map. patches of seasonally inundated pans and grasslands occur in the miombo and mopane landscapes. where they are sufficiently large, they have been individually mapped and are superimposed on the landscape map. landscapes of the pnz save river channel and river banks: this landscape is very dynamic, encompassing 1.7% of the pnz. when the river is in flood, most of this landscape would be under water. however, when the waters recede, large expanses of sand banks are exposed. these are clearly visible on the satellite image. the vegetated part of this landscape consists of community 8 (pluchea dioscorides – digitaria eriantha thickets). save riverine forest: this landscape, which occupies 0.8% of the pnz, occurs on the stable banks and levees south of the save river that are not much influenced by flood dynamics. the landscape is primarily made up by community 9 (cordyla africana – philenoptera violacea forest) and adjoining community 1 (hyphaene petersiana – acacia xanthophloea woodland). this forest type consists of a very diverse assemblage of trees reaching heights of over 20 m. this forest has been heavily impacted upon by shifting cultivation. acacia nigrescens woodland: this landscape, which covers 10.5% of the pnz, mostly occurs on the brown soils derived from calcaric sedimentary rocks in an east-west band that lies south of the save river. this landscape merges into the save riverine forest to the north and into the mopane and miombo landscapes in the south. large expanses of this landscape are found in the eastern half of the pnz. smaller patches were observed south of the save river in the western part but could not be mapped. this landscape is principally made up by community 5 acacia nigrescens – urochloa mosambicensis woodland, with bush clumps of community 3 (xanthocercis zambesiaca – salvadora persica thickets) and community 2 (acacia burkei – andropogon gayanus woodland) along the small seasonal rivers that drain this landscape towards the save river. large patches of mopane (community 4) are also found. this landscape has strong affinities to the sclerocarya birrea/acacia nigrescens landscape 17 described from the northern part of the knp by gertenbach (1983). mopane landscape: this landscape is found on heavier soils including the clayey colluvium derived from mananga sediments and portions of the brown soils derived from calcaric sedimentary rocks. this is the most extensive landscape found in the pnz; it covers 37.7% of the park. its dominant feature is the mopane-dominated closed woodlands of community 4 (colophospermum mopane – urochloa mosambicensis woodlands) with community 2 (acacia burkei – andropogon gayanus woodland) along the small seasonal rivers that drain this landscape towards the save river. seasonal pans with community 10 (paspalidium obtusifolium open grasslands) occur. wetland type ts (ramsar definition) is therefore represented here. sandveld landscape: this landscape was mostly encountered in the western sector of the park. it comprises mostly community 7 guibourtia conjugata – digitaria eriantha open woodlands on deep reddish sandy soils that are characterised by relatively high ca/mg ratios and high p values. it adjoins both the miombo and mopane landscapes. the sandveld landscape is the dominant feature of the lnp and bnp; it covers respectively 41% and 46.7% of these parks (stalmans et al. 2004; stalmans & wishart 2005). however, it covers only 16.7% of the pnz. within the gltp, the sandveld landscape appears to be gradually replaced in the east by the miombo landscape in response to a gradient of increasing rainfall towards the coast. miombo landscape: this landscape covers mostly the unspecified soils derived from sandy substrates and interior dunes as well as the unspecified soils derived from mananga sediments. this is the second-most extensive landscape of the pnz and covers 29.5% of the park. this landscape covers the higher lying part of the znp in the southeastern sector, representing the wetter part of the park. the miombo landscape extends in a northern and northwesterly direction where it becomes interwoven with the mopane and the sandveld landscape. the miombo landscape is primarily made up by community 6 julbernardia globiflora – digitaria eriantha woodland. seasonal pans comprising community 10 (paspalidium obtusifolium open grasslands) occur. wetland type ts (ramsar definition) is therefore represented here. the size, density and regularity of tall termitaria are one of the prominent features of miombo landscapes (desanker et al. 1997). the termitarium’s soils have higher levels of total nitrogen, acid-extractable phosphorus and cations than surrounding soils due to both the concentration in and subsequent decomposition of organic matter within the mounds and the concentration of minerals through the evaporation of soil water in chimneys within the mound. these nutrient-rich patches within an otherwise nutrient-poor landscape invariably support distinctively different species and are often the focus of activity for birds and other animals that might otherwise not thrive so well in such a largely nutrient-poor environment. accuracy of the landscape map the occurrence of sample plots and control points in the original land cover polygons of the carta de uso e cobertura da terra (dinageca 1999) provides a good idea of the level of certainty or uncertainty that can be associated with the landscape map (see figure 7). the highest level of uncertainty is associated with the boundary areas. although several land cover polygons were not sampled on the northern boundary, the well-known role of the riverine position in determining the vegetation make-up results in a much lower level of uncertainty than assumed from the map. analysis of the satellite image strongly suggests that the poorly sampled areas in the east and west are covered by the miombo and mopane/sandveld landscapes respectively as mapped in figure 6. figure 6:landscape map of the pnz figure 7:sampling intensity across original land cover polygons in the pnz plant community diversity time constraints and limited ground coverage did not allow for a detailed inventory of plant diversity. therefore no inventory of forbs and geophytes was undertaken. the pnz is very rich in tree species, with 208 woody species a recorded in the sample plots. additionally, 41 species of grasses were noted. this species richness is evident if one considers that only 178 woody species were recorded in the much greater sample of 175 plots that were surveyed in the much larger lnp. it is of interest to note that a number of species were recorded during this survey that according to the generally used literature (coates palgrave et al. 2002) only occur much further north along the zambezi and along the coast. these species include xylocarpus granatum and ficus bussei. this further strengthens the fact that very little is formally known about the vegetation of the pnz highlighting the need for further studies. land-use patterns and their influence on the vegetation of the pnz a number of communities reside within the boundaries of the pnz, living off subsistence farming and fishing. the subsistence farming is based on the chitimene system of cultivation (shifting cultivation) whereby fields are cleared manually with farmers often cutting only the branches off trees and leaving the stems standing. the cut material is gathered and burnt in the central parts of the field. crops are planted in the ash bed that provides nutrients. androstachys patches are cleared preferentially in the miombo landscape. julbernardia wood lands are also used for this subsistence farming. farmers abandon the fields after 3–4 years of cultivation, allowing the miombo woodland to regenerate while new fields are cleared and planted. cultivation of termitaria within the miombo was also observed whereby maize is planted in the fertile soil of the cleared/flattened termitarium. the edges of the pans are often also cultivated as the waters recede. the extent of the cultivated area is relatively limited and the impact is probably fairly small as the next flood cycle probably ‘resets’ the cultivated patch and surrounding areas to a comparable state. a significant impact was observed on the small extent of save riverine forest landscape where clearing for cultivation is commonplace.although no formal assessment was made, much use is clearly made of trees for the building of dwellings and livestock pens and the carving of various items. the impact of these activities is selective. whereas this use does not seem to impact significantly on the overall woody resource, no data are available on the impact on specific species. grass is cut and used for thatching. at least three species are cut and used for thatching (heteropogon contortus, eragrostis pallens and aristida cf. adscensionis). much use is made of medicinal plants and wild fruits (e.g. strychnos spp.) but no data are available regarding the impact thereof on specific species.the largest selective impact through resource use is probably as a result of the harvesting of bark for the making of beehives. long cylinders of bark are stripped from mostly julbernardia globiflora, resulting in the death of the tree. due to the current low diversity and numbers of game, levels of herbivory (both grazing and browsing) are currently very low in the pnz (see table 1). although both cattle and goats were observed grazing in the area, it appears that livestock numbers are also relatively low and with the exception of a single sample plot, only light grazing was observed. the grass and tree layer is generally in an excellent condition, offering a suitable habitat for the herbivores that could be expected in this environment (both those occurring historically and those still present; see table 1). during this vegetation survey, opuntia ficus-indica (prickly pear) was the worst invader species observed. it occurs in a few dense stands on the southeastern border in the miombo landscape. agave sisalana was also observed on the southeastern border as well as ricinus communis on the banks of the save river. it is supposed that because of the higher rainfall and extensive riverine fringe (that is affected by annual flood disturbance) the pnz is relatively susceptible to alien plant invasion. table 1: historical and current occurrence of wildlife species in the pnz acknowledgements (back to top) the vegetation research was undertaken during 2003, as part of the planning work by development alternatives inc. and funded by usaid, in support to the direcção nacional de áreas de conservação of the ministério do turismo, mozambique. the management plan for the pnz was compiled by ecologists dr d. grossman and ms p. holden. assistant park warden snr. sensao and his staff provided logistical support and accompanied the survey team in the field. j. burrows, e. schmidt and m. lötter of the plant specialist group assisted with species identification. the aerial control points were assessed during the aerial wildlife count that was funded by the projecto áreas de conservação transfronteira e desenvolvimento do turismo (actfdt) of the ministério do turismo. logistical support was provided during 2007 for the aerial survey by park warden snr a. abacar, pilot m. pingo, as well as j. peel, a. jacobs, b. cuthill, b. olbricht and o. büchner. references (back to top) bandeira, s.o., hatton, j.c., munisse, p. & izidine, s., 1994, ‘the ecology and conservation status of plant resources in mocambique’, strelitzia 1, 105–115. coates palgrave, k., coates palgrave, m., drummond, r.b. & moll e.j. (eds.), 2002, trees of southern africa, 3rd edn., struik publishers, johannesburg. departemento terra e agua, 1995, ‘legenda da carta nacional de solos (escala 1:1000000)’, compilada pelo departemento terra e agua, maputo, moçambique. desanker, p.v., frost, p.g.h, frost, c.o., justice, c.o & scholes, r.j. (eds.), 1997, ‘the miombo network: framework for a terrestrial transect study of land-use and land-cover change in the miombo ecosystems of central africa’, igbp report 41, the international geosphere-biosphere programme (igbp), stockholm, sweden. dinageca, 1999, ‘carta de uso e cobertura da terra’, joint venture ignfi cenacarta dinageca, moçambique. dnac, 2002, ‘zinave national park, management and development plan’, unpublished report to the national directorate for conservation areas, ministry of tourism, republic of moçambique. duarte, p.d. & oreste, m.n., 1999, ‘country brief on non-wood forest products: ec-fao partnership programme (1998–2000)’, project gcp/int/679/ec data collection and analysis for sustainable forest management in acp countries. edwards, d., 1983, ‘a broad‑scale structural classification of vegetation for practical purposes’, bothalia 14(3&4), 705–712. gabriel, h.w. & talbot, s.s., 1984, ‘glossary of landscape and vegetation ecology for alaska’, blm–alaska technical report 10, u.s. department of the interior. gauch, h.g., 1982, multivariate analysis in community ecology, cambridge university press, cambridge. gertenbach, w.p.d., 1983, ‘landscapes of the kruger national park’, koedoe 26, 9–121. hill, m.o., 1979, twinspan – a fortran program for arranging multivariate data in an ordered two-way table by classification of the individuals and attributes, cornell university, ithaca, new york. jongman, r.h.g., ter braak, c.f.j. & van tongeren, o.f.r., 1987, data analysis in community and landscape ecology, wageningen, pudoc. kelly, r.d. & walker, b.h., 1976, ‘the effect of different forms of land use on the ecology of a semi-arid region in south-eastern rhodesia’, journal of ecology 64, 553–576. land type survey staff, 1989, ‘land types of the map 2530 barberton’, memoirs of the agricultural natural resources of south africa 13. leemans, r. & cramer, w., 1991, ‘the iiasa database for mean monthly values of temperature, precipitation and cloudiness of a global terrestrial grid’, international institute for applied systems analysis (iiasa), rr-91-18. micoa, 1997, ‘first national report on the conservation of biological diversity in moçambique’, ministry for the coordination of environmental affairs, maputo. mueller‑dombois, d. & ellenberg, h., 1974, aims and methods of vegetation ecology, john wiley & sons, new york. national working group for vegetation ecology, 1986, ‘soil classification according to the binomial classification system’, technical communication 3. peel, m.j.s., kruger, j.m. & macfadyen, s., 2007, ‘woody vegetation of a mosaic of protected areas adjacent to the kruger national park, south africa’, journal of vegetation science 18(6), 807–814. scholes, r.j. & walker, b.h., 1993, an african savanna, cambridge university press, cambridge. siebert, f., bredenkamp, g.j. & siebert, s.j., 2003, ‘a comparison of mopaneveld vegetation in south africa, namibia and zimbabwe’, bothalia 33(1), 121–134. smithers, r.h.n. &. lobão tello, j.l.p., 1976, ‘check list and atlas of the mammals of mocambique’, museum memoir no. 8, trustees of the national museums and monuments of rhodesia, bulawayo, zimbabwe. stalmans, m., 1994, ‘vegetation survey of malilangwe’, unpublished report to the malilangwe conservation trust. stalmans, m., gertenbach, w.p.d & carvalho-serfontein, f., 2004, ‘plant communities and landscapes of the parque nacional do limpopo, moçambique’, koedoe 47(2), 61–81. stalmans, m. & peel, m., 2007, ‘wildlife survey – parque nacional de zinave, moçambique’, unpublished report by international conservation services and agricultural research council for áreas de conservação transfronteira e desenvolvimento do turismo – ministério do turismo, república de moçambique. stalmans, m. & wishart, m., 2005, ‘plant communities, wetlands and landscapes of the parque nacional de banhine, moçambique’, koedoe 48(2), 43–58. storer, r.w. & dalquest, w.w., 1967, ‘birds from the save river of mozambique’, occasional papers of the museum of zoology, university of michigan 62, 1–14. ter braak, c.j.f., 1986, ‘canonical correspondence analysis: a new eigenvector technique for multivariate direct gradient analysis’, ecology 67(5), 1167–1179. ter braak, c.j.f., 1992, canoco – a fortran program for canonical community ordination, microcomputer power, ithaca, new york. timberlake, j.r., nobanda, n. & mapaure, i., 1993, ‘vegetation survey of the communal lands – north and west zimbabwe’, kirkia 14(2), 171–270. van rooyen, n., theron g.k. & grobbelaar, n., 1981, ‘a floristic description and structural analysis of the plant communities of the punda milia-pafuri-wambiya area in the kruger national park, republic of south africa: 2. the sandveld communities’, journal of south african botany 47(3), 405–449. werger, m.j.a. & coetzee, b.j., 1978, ‘the sudano-zambezian region’, in m.j.a. werger (ed.), biogeography and ecology of southern africa, pp. 301–454, dr w. junk, the hague. a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 160 183 original research vegetation of the hantam-tanqua-roggeveld subregion, south africa part 2: succulent karoo biome related vegetation abstract the hantam-tanqua-roggeveld subregion lies within the succulent karoo hotspot that stretches along the western side of the republic of south africa and namibia. this project, carried out to document the botanical diversity in the hantam-tanqua-roggeveld subregion, was part of a project identified as a priority during the skep (succulent karoo ecosystem programme) initiative in this hotspot. botanical surveys were conducted in an area covering over three million hectares. satellite images of the area and topocadastral, land type and geology maps were used to stratify the area into relatively homogeneous units. an analysis of the floristic data of 390 sample plots identified two major floristic units, i.e. the fynbos biome related vegetation and the succulent karoo biome related vegetation. a description of the vegetation related to the succulent karoo biome is presented in this article. seven associations, 16 subassociations and several mosaic vegetation units, consisting of more than one vegetation unit, were identified and mapped. various threats to the vegetation in the region were identified during the survey and are briefly discussed. keywords: hantam, phytosociology, roggeveld, succulent karoo, tanqua karoo, vegetation map, western mountain karoo helga van der merwe margaretha w. van rooyen noel van rooyen department of plant science university of pretoria south africa correspondence to: helga van der merwe e-mail: soekop@hantam.co.za postal address: department of plant science, university of pretoria, pretoria, 0002, south africa 160 the succulent karoo (cepf 2003) stretches along the western side of south africa and namibia and is one of only two global hotspots that are entirely arid (conservation international 2006). within this area the succulent karoo ecosystem plan (skep) identified the hantam-tanqua-roggeveld subregion as an area for which there is little information on the floral diversity, and consequently the critical ecosystem partnership fund (cepf) funded botanical studies in the subregion. the hantam-tanqua-roggeveld subregion occurs where three biomes, namely the fynbos, succulent karoo and the nama karoo biomes (rutherford & westfall 1986) meet. due to the transitional nature of the area there has been some controversy as to whether the vegetation, especially that of the roggeveld, should be classified as fynbos or succulent karoo. the transitional nature of the subregion is supported in a recent study by born et al. (2007) who argue for the recognition of a greater cape floristic region. the fynbos related vegetation, or fire-prone vegetation within the subregion, is discussed by van der merwe et al. (2008). this article will focus on the succulent karoo related vegetation, which is not fire-prone. only a few studies have been conducted in the area, most of which were concentrated in the nieuwoudtville area. amongst these was a study by snijman and perry (1987) on the floristics of the nieuwoudtville wild flower reserve and the conservation farming project of the former national botanical institute (now the south african national biodiversity institute, sanbi), resulting in a host of research projects in the nieuwoudtville area (south african national biodiversity institute, 2006). the tanqua karoo has been less studied than the nieuwoudtville area, although a phytosociological study was conducted in the tankwa national park by rubin (1998). however, the park subsequently substantially extended its boundaries beyond the area included in rubin’s original study. hilton-taylor (1994) considered the subregion as part of the succulent karoo biome. he identified two centres of endemism within the western cape domain, viz. the western mountain karoo and tanqua karoo, both of which occur within the area covered in this article. the botanical importance of the hantam-roggeveld was also emphasised by van wyk and smith (2001). they identified the hantam-roggeveld as one of their 13 principal centres of plant endemism in southern africa and also classified it as part of the succulent karoo biome. in this study a broad-scale vegetation survey of the entire subregion of approximately 3 000 000 ha (30 000 km2) was undertaken. analysis of the data revealed two distinct vegetation clusters. the first cluster, predominantly mountain renosterveld, is related to the fynbos biome, and is described in van der merwe et al. (2008). the aim of the present paper is to present a phytosociological analysis and description of the second vegetation cluster, i.e. the vegetation related to the succulent karoo biome, and to create a map depicting the different vegetation units. a brief discussion of the various threats to the region as identified by the farming community and the northern cape department of agriculture, and through personal observations, is presented. study area the hantam-tanqua-roggeveld subregion (see figure of subregion in van der merwe et al. 2008) lies in the predominantly winter rainfall region of the northern and western cape provinces of south africa. in the south it stretches from the ceres karoo, where the swartrug and bontberg mountains meet, northwards into the tanqua basin. the eastern border includes the hantam, roggeveld, komsberg, klein roggeveld and nuweveld mountains to just southwest of fraserburg, while the cederberg and bokkeveld mountains to just north of loeriesfontein form the western boundary. the smaller study area discussed in this paper covers parts of the hantam region, the tanqua and ceres karoo regions 2008 original research van der merwe, van rooyen & van rooyen vol. 50 no. 1 pp. 160 183koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za as well as the area east of the roggeveld mountains. this area includes two of acocks’s veld types (acocks 1953, 1988), namely: succulent karoo (veld type 31) and western mountain karoo (veld type 28). these two veld types of acocks are equivalent to the lowland succulent karoo (unit 57) and upland succulent karoo (unit 56) of low and rebelo (1998), respectively. mucina et al. (2005) recognised the following large vegetation types in the study area: hantam karoo (skt 2), roggeveld karoo (skt 3), tanqua escarpment shrubland (skv 4), tanqua karoo (skv 5), tanqua wash riviere (azi 7) and namaqualand riviere (azi 1). smaller patches of nieuwoudtville roggeveld dolerite renosterveld (frd 1), hantam plateau dolerite renosterveld (frd 2), nieuwoudtville shale renosterveld (frs 2) and bushmanland vloere (azi 5) are also mapped in the area. the physical geography of the region differs greatly. from the level plains of the tanqua karoo the landscape in the east rises steeply up the escarpment to the plateau formed by the roggeveld, komsberg, koedoesberg and nuweveld mountains. the hantam is characterised by a gently undulating to a steeply rolling topography. geologically, the beaufort and ecca groups dominate the study area (rubidge & hancox 1999, council for geoscience 1973, 1983, 1989, 1991, 1997, 2001, 2008). the ecca group covers most of the study area, with the dwyka group (tillite, sandstone, mudstone and shale) cropping out in the west. the ecca group includes sediments of the tierberg (shale), prince albert (mudrock), kookfontein (shale, siltstone and stone) and skoorsteenberg formations (mudstone, siltstone and sandstone). the mudstones of the abrahamskraal formation in the beaufort group are found on the eastern side of the study area, while igneous intrusions of dolerite occur throughout the region. an array of land types is represented in the study area (agricultural research council 1986a, 1986b, 1995, 1999a, 1999b, 2002, 2003). the prominent land types include ag, da, dc, fb, fc, ia and ib (du plessis 1987). rainfall ranges from 50 to 300 mm a year, with a mean of 228 mm, as measured at calvinia (sa weather bureau 1998). although the rain falls mainly in winter it does include a few summer thunderstorms. the highest annual rainfall, measured over a 29-year period, at calvinia, was 472 mm in 1976 (sa weather bureau 1998). january and february have a mean maximum temperature of 30.8°c while an extreme maximum of 41.2°c was recorded in february 1990. the coldest months are june and july with a mean maximum of 17.1°c and 17.2°c respectively. the lowest temperature recorded was –6.5°c in june 1978 (sa weather bureau 1998). methods and materials on the basis of the colour, texture and topography, satellite images (bands: 4,5,3 (r,g,b)) of the study area were visually stratified into relatively homogeneous units. floristic surveys were carried out during the spring of 2004 at 390 sites selected in the stratified homogeneous units close to any national or provincial road or farm track. plot sizes 10 x 10 m were used in most of the surveys but larger plots (20 x 20 m) were used in more denuded areas (rubin 1998). global positioning system (gps) coordinates were taken at each site. a cover-abundance value, according to the braun-blanquet scale (werger 1974), was noted for each species present in a plot. aspect, slope, position in the landscape, soil type and colour, an estimation of rock cover, rock size and erosion were noted at each sampling point. rock size was categorised into gravel (< 10 mm), small stones (10–50 mm), stones (> 50–200 mm) and boulders (> 200 mm). trampling, small mammal activity and invasion by alien plants were some of the biotic effects also recorded. using the turboveg and megatab computer package an analysis of the floristic data was conducted (hennekens & schaminée 2001). turboveg software was used to capture the vegetation data and a twinspan (two way species indicator analysis (hill 1979)) was run in megatab as a first step to the classification of the vegetation data. the result of the twinspan (two way species indicator analysis (hill 1979)) on the entire data set of 390 relevés confirmed the presence of two distinct floristic groups, which enabled the data set to be split into two, and a twinspan was then run on each data set separately. the resulting differential tables were further refined using braun-blanquet procedures. the first data set (107 relevés) characterised the vegetation of the predominantly mountain renosterveld with fynbos affinities, and is described by van der merwe et al. (2008). the second data set (283 relevés) characterised the vegetation with predominantly succulent karoo affinities and is discussed further in this article. the basic unit used here in the classification of the vegetation is the association. an association, as defined by nelder et al. (2005), describes vegetation units on the basis of the presence and abundance of species, vegetation structure and spatial distribution of individuals in the dominant layer. species within a subdominant structural layer and their canopy cover are used to describe the subassociations. associations and subassociations were mapped using the botanical survey data as well as 1:250 000 topocadastral maps, land type maps, geology maps (and electronic information supplied by the council of geoscience, 2008), and satellite images. several mosaic units, which include more than one association or subassociation due to the transitional nature of the area, were also identified and mapped. (an electronic arcview compatible version (raster or vector format) of the vegetation map is available from the authors on request. the pdf version provided here should best be printed to a3.) herbarium specimens were sent to compton herbarium, kirstenbosch, for identification if identification of species in the field was in doubt. the collection code (hr) and numbers of the specimens were kept throughout the process. all the species, especially within the aizoaceae (mesembryanthemaceae), have not yet been positively identified. the h.g.w.j. schweickerdt herbarium (pru), university of pretoria, houses the voucher specimens collected. nomenclature follows that of germishuizen and meyer (2003). results the phytosociological analysis of the floristic data is given in tables 1a, 1b and 1c. in order to prevent confusion and to aid in the compilation of a vegetation map of the entire subregion the plant associations have been numbered sequentially following the three associations (1–3) described in part 1 of this series (van der merwe et al. 2008). seven associations were identified in the area under discussion and these were subdivided into 16 subassociations, as set out in the following scheme: 4. pteronia glauca – euphorbia decussata escarpment karoo 4.1 montina caryophyllacea – pteronia glauca roggeveld escarpment karoo 4.2 galenia africana – pteronia glauca escarpment karoo 4.3 euphorbia decussata – ruschia cradockensis escarpment karoo 5. eriocephalus purpureus hantam karoo 5.1 erodium cicutarium – eriocephalus purpureus hantam karoo 5.2 ruschia cradockensis – eriocephalus purpureus hantam karoo 5.3 leipoldtia schultzei – eriocephalus purpureus hantam karoo 6. pteronia glomerata roggeveld karoo 6.1 phyllobolus tenuiflorus – pteronia glomerata roggeveld karoo 6.2 eriocephalus pauperrimus – pteronia glomerata roggeveld karoo 161 succulent karoo biome related vegetation – part 2 original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 160 183 162 6.3 pentzia incana – pteronia glomerata roggeveld karoo 6.4 ruschia intricata – pteronia glomerata roggeveld karoo 6.5 eriocephalus ericoides – pteronia glomerata roggeveld karoo 7. aridaria noctiflora tanqua and loeriesfontein karoo 7.1 ruschia robusta – aridaria noctiflora tanqua karoo 7.2 drosanthemum (hr217) sp. – aridaria noctiflora tanqua karoo 7.3 malephora crassa – aridaria noctiflora tanqua karoo 7.4 atriplex lindleyi – aridaria noctiflora loeriesfontein karoo 7.5 ruschia intricata – aridaria noctiflora tanqua karoo 8. stipagrostis obtusa central tanqua grassy plains 9. mesembryanthemaceae (hrp359) sp. ceres karoo vygieveld 10. pteronia (hrp118) sp. tanqua brackish flats two vegetation units falling outside the present study area have been included in table 1a. with the exception of these units and associations 9 and 10 all vegetation units could be mapped (fig. 1). ten mosaics found in areas where the landscape is spatially diverse were also mapped. the vegetation units making up the mosaics and the most important environmental features distinguishing the units are set out below: the nieuwoudtville mosaic consists of vegetation • units 2.1.1, 2.1.4, 2.2 of the mountain renosterveld vegetation (van der merwe et al. 2008.) and 5.1 predominantly found on the dolerite outcrops; the grootfontein mosaic consists of vegetation units 4.2, • 4.3 on undulating ridges and 5.1 and 5.2 on dolerite outcrops and dolerite derived soils; the houhoek mosaic consists of vegetation unit 4.1 at a • higher altitude and higher rock cover and unit 4.3 at a lower altitude and lower rock cover; the adamsfontein/noodsaaklikheid mosaic with • vegetation unit 5.2 predominantly on dolerites and 6.5 on shales and dolerite intrusions that have been disturbed; the calvinia mosaic consists of vegetation units 6.5 on • ecca shales and dolerite intrusions and 7.4 on brackish soils (historically overutilised); the kalkgat mosaic consists of vegetation units 4.1 on • rocky ridges, 7.2 on shales and sedimentary deposits and 7.3 on brackish soils; the tanqua karoo inselberg mosaic with vegetation • unit 5.2 on dolerites and 7.5 on ecca shales; the tanqua pan mosaics consist of vegetation units 7.2 • on shales and alluvial deposits and 10 on brackish soils; the windheuwel/rooiheuwel mosaic consists of • vegetation units 4.1, 4.2 on the rocky ridges and 7.3 on the brackish plains; the naresie mosaic consists of elements of the • namaqualand brokenveld as defined by acocks (1953, 1988) and vegetation unit 6.4. the year in which the surveys were conducted (2004) was a poor rainfall year, and would have substantially reduced the annual and geophyte components of the vegetation. the resulting phytosociological tables thus poorly represent these two components. in normal or good rainfall years the contribution of the annuals and geophytes to the vegetation cover would be considerably higher. description of plant associations (tables 1a, 1b, 1c; fig. 1) two vegetation units, included in table 1a, have not been mapped as they fall outside the present study area and their linkages with the vegetation in the van rhynsdorp region still requires further study. these two units are related to what acocks (1953, 1988) described as succulent karoo in the van rhynsdorp region, and are found along the doorn river and its tributaries, the biedouw valley and the western extreme of the brackish system of the hantam river in the vicinity of van rhynsdorp. the first unit is characterised by a strong presence of species group a, and a weak presence of species group b, while the second unit comprises species group b. at present it is not clear whether the unmapped units form part of association 4 or whether they have closer affinities to associations outside the study area. 4. pteronia glauca – euphorbia decussata escarpment karoo this plant association is located along the roggeveld escarpment, the slopes of the hantam mountains, on the slopes alongside the hantam river in the agter-hantam and to the west of the stinkfontein, lang and boegoeberg mountains, and has links to associations along the western part of the doorn river and its tributaries (fig. 1). geologically, the ecca group dominates this association while tillite and dolerite intrusions are present at times. a variety of land types are found in this plant association, including fc, fb, ia, ib, da and ag. the association generally occurs in lower-lying areas on level terrain, gentle or moderate to steep slopes. the percentage rock cover varies tremendously (0–99%) and includes gravel, small stones, stones and boulders. shrub cover is usually high, while grass and annual cover is limited. prominent species included in this plant association are pteronia glauca (species group c, table 1a), euphorbia decussata (species group d), pentzia incana and ruschia cradockensis (species group n), as well as tripteris sinuata, drosanthemum (hr217) sp. and galenia africana (species group v). plant associations 4, 5 and 6 are closely related because they share the common species in species group n, yet associations 4 and 5 (table 1a) differ from association 6 (table 1b) since both associations 4 and 5 lack species group u. 4.1. montina caryophyllacea – pteronia glauca roggeveld escarpment karoo this subassociation characterises the vegetation of the roggeveld escarpment, i.e. the west-facing slopes of the roggeveld mountains (fig. 1), and also forms part of the houhoek, kalkgat as well as windheuwel/rooiheuwel mosaics. subassociation 4.1, excluding the various mosaic vegetation units, covers an area of 152 057 ha (9.2% of the total area in fig. 1). the subassociation occurs at intermediate altitudes of 700 to 1 100 m above sea level, on gentle to moderate, and sometimes steep, slopes. a high rock cover of generally more than 90%, comprising small stones, stones and boulders, is found in this subassociation. the land type is predominantly fc. the light brown to brown coloured loamy soils are derived from ecca shales. the vegetation is characterised by a high shrub cover (50–90%) and grasses are usually absent. a very low cover of annuals can be found in certain areas. subassociation 4.1 comprises species group b and there is a strong presence of the diagnostic species group c (table 1a). the vegetation is dominated by pteronia glauca (species group c) that is constantly present and often has a very high canopy cover. other prominent species present include tylecodon wallichii and montinia caryophyllacea (species group b). 4.2. galenia africana – pteronia glauca escarpment karoo plant subassociation 4.2 is located on the slopes of the hantam mountain, the undulating slopes of the escarpment in the platberg and surrounding area southwest of calvinia, and the slopes where the roggeveld and klein roggeveld mountains meet (fig. 1). subassociation 4.2, excluding the mosaic vegetation units, covers an area of 45 047 ha (2.7% of the total area in fig. 1). it is also found in the mosaic between the roggeveld and koedoesberg mountains in the vicinity of the farms windheuwel and rooiheuwel. elements of this subassociation are found on the ridges between the dolerite derived soils in the original research van der merwe, van rooyen & van rooyen vol. 50 no. 1 pp. 160 183koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za figure 1 vegetation map of the succulent karoo biome related vegetation of the hantam-tanqua-roggeveld subregion 163 succulent karoo biome related vegetation – part 2 original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 160 183 grootfontein mosaic in the hantam along the r27 national road, east of nieuwoudtville. land types are numerous, and include fb, ia, ib and da, at altitudes ranging from 700 to 1 200 m above sea level. ecca shales and dolerite intrusions predominate in this vegetation unit on gently sloping terrain. a low rock cover (< 10%) or a high rock cover (> 80%), consisting of gravel, small stones, stones and boulders covers the light brown or brown loamy soils. shrub cover in this subassociation is high (> 70%), while the grass and annual components are not well represented. pteronia glauca (species group c) is diagnostic of this subassociation (table 1a). other species present include pentzia incana, eriocephalus ericoides (species group n) and tripteris sinuata, drosanthemum (hr217) sp. and galenia africana (species group v). 4.3. euphorbia decussata – ruschia cradockensis escarpment karoo this small subassociation occurs at lower altitudes of 400 to 1 200 m above sea level, to the west of the hantam mountain and, excluding the mosaic vegetation units, covers an area of 5 196 ha (0.3% of the total area in fig. 1). elements of this subassociation can also be found closer to nieuwoudtville in the grootfontein mosaic as well as in the agter-hantam in the houhoek mosaic where it covers the lower slopes. the ecca group dominates in this subassociation, while tillite and dolerite intrusions are present in the mosaics. land types include fb, fc, da, ag and ib, resulting in the mosaic vegetation. the level, gently or moderately sloping ridges are usually covered with either very little rock (0–5%) or much rock (60–99%). small stones, stones and boulders are present in the light brown to brown loamy soils. shrub cover is high (45–95%), whereas the grass and annual components are generally weakly developed. the dominant species are euphorbia decussata (species group d, table 1a) and ruschia cradockensis (species group n), and both these species generally have a high cover. other species present include tetragonia fruticosa (species group n), tripteris sinuata, drosanthemum (hr217) sp. and galenia africana (species group v). 5. eriocephalus purpureus hantam karoo association 5 occurs in the vicinity of the farm matjiesfontein, close to calvinia, northwards to loeriesfontein and southwards to the farm klipbak. it also occurs as part of the grootfontein mosaic, westwards along the r27 national road to the nieuwoudtville mosaic, as described by van der merwe et al. (2008). additionally, it is found in the agter-hantam (adamsfontein/noodsaaklikheid mosaic) and on small inselbergs into the tanqua karoo, such as the nuwewater se berg, elandsberg, eselberg, leeuberg, potkleiberg and sterretjieberg (tanqua karoo inselbergs mosaic) (fig. 1). the altitude is lower than 1 000 m and an array of land types occur, including dc, fc, da and ag. the level to gently sloping ridges usually have a low rock cover. the brown, red brown or light brown loams and clays are derived from ecca shales and dwyka tillite, as well as dolerite intrusions that scatter the landscape. the shrub cover varies considerably (0–80%) and grasses are generally absent. the cover of the annual component ranges from very little to 95%. it is assumed that the annual component contributes substantially to the total aboveground biomass in good rainfall years since the area is renowned for its annual and geophyte spring displays. eriocephalus purpureus (species group i) and leipoldtia schultzei (species group j) characterise the vegetation of this association (table 1a). other prominent species include pentzia incana, ruschia cradockensis, asparagus capensis, tetragonia fruticosa and eriocephalus microphyllus from species group n, as well as aridaria noctiflora and galenia africana from species group v. plant association 5 is closely related to both associations 4 (table 1a) and 6 (table 1b) due to the common species in species group n. this association is subdivided into three subassociations. 5.1. erodium cicutarium – eriocephalus purpureus hantam karoo the two variations of this subassociation (table 1a) are mainly due to the presence or absence of annuals and/or geophytes in species groups e and f, which could be as a result of the drought year in which the surveys were conducted. these life forms are rainfall dependent and thus by conducting the same surveys in a good rainfall year it might not be possible to distinguish between the two variations. this subassociation occurs in the vicinity of the farm matjiesfontein, close to calvinia, southwards to the farm klipbak and northwards to loeriesfontein (fig. 1). subassociation 5.1, excluding the mosaic vegetation units, covers an area of 92 682 ha (5.6% of the total area in fig. 1). it also occurs as part of the grootfontein mosaic, as well as westwards to the nieuwoudtville mosaic where it occurs in combination with variants 2.1.1, 2.1.4 and subassociation 2.2 (van der merwe et al. 2008). the subassociation is found on the ecca group where dolerite intrusions occur in the landscape, on level to gently sloping areas, at an altitude from 700 to 1 000 m above sea level on land types fb, ea, dc or da. soils are clays or loams of red brown or dark brown colour, with little or no rocks. shrub cover is highly variable (< 5–75%) and the contribution of the grass component is very small. annual cover varies from < 1% to 95%, indicating its high variability. subassociation 5.1 differs from 5.2 by the presence of species groups e, f and g, which occur only in subassociation 5.1 (table 1a). common species include erodium cicutarium (species group g), ehrharta calycina (species group h), eriocephalus purpureus and tetragonia microptera (species group i), pentzia incana, ruschia cradockensis and asparagus capensis (species group n). 5.2 ruschia cradockensis – eriocephalus purpureus hantam karoo subassociation 5.2 occurs as a clearly defined vegetation unit as well as in various mosaic vegetation units. as a clearly defined unit it occurs in the agter-hantam near klipwerf (fig. 1) and, excluding the mosaic vegetation units, covers an area of 22 353 ha (1.3% of the total area in fig. 1). here the subassociation is found on the ecca group while dolerite intrusions are scattered throughout the landscape. as part of a mosaic, it occurs north and south of the hantam river in the adamsfontein/noodsaaklikheid mosaic. it also occurs on dolerites on the inselbergs south of calvinia into the tanqua karoo (tanqua karoo inselbergs mosaic). these inselbergs include the nuwewater se berg, elandsberg, eselberg, leeuberg, potkleiberg and sterretjieberg. additionally, subassociation 5.2 occurs in the grootfontein mosaic vegetation unit on the dolerite derived soils. this subassociation generally occurs on land types fb and fc at altitudes from 700 to 1 000 m above sea level. the rock cover varies considerably from zero to 90% and includes small stones, stones and boulders. the underlying red brown or brown soils are clays and loams, primarily derived from the dolerites and the ecca shales. shrub cover ranges from 40 to 90%, while the grass and annual components are usually absent or < 5%. ehrharta calycina (species group h), eriocephalus purpureus (species group i), pentzia incana and ruschia cradockensis (species group n), as well as aridaria noctiflora (species group v) are prominent in this subassociation (table 1a). subassociation 5.2 is closely related to subassociation 5.1 as they both include species groups h, i, j, n and v, but it differs from the latter by the absence of species groups e, f and g. subassociation 5.2 is also closely related to subassociation 5.3, from which it only differs by the absence of species group h in the latter. 164 original research van der merwe, van rooyen & van rooyen vol. 50 no. 1 pp. 160 183koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za r e l e v é n u m b e r 3 3 3 3 1 1 1 2 1 1 3 2 2 3 1 1 3 3 3 1 1 2 2 2 2 2 1 2 2 3 3 1 1 2 3 1 1 1 1 3 8 4 7 8 8 2 6 6 4 9 9 8 3 3 4 5 1 5 7 8 8 3 8 4 4 7 1 2 8 0 2 5 5 8 0 5 8 3 2 1 1 4 5 6 0 4 6 8 0 4 1 5 9 9 1 2 3 2 1 1 1 2 3 3 5 1 6 6 6 1 4 4 4 5 1 2 2 2 5 5 7 3 2 2 5 9 5 3 3 7 1 4 7 2 5 5 1 7 0 6 9 1 6 5 1 9 7 8 3 1 7 9 2 9 9 9 5 1 0 2 9 7 4 6 4 2 4 5 6 7 1 8 7 1 2 3 0 3 0 0 0 8 3 4 6 8 1 0 0 0 2 3 3 2 3 1 2 9 7 1 1 5 8 9 9 5 8 9 1 6 6 7 8 5 8 1 8 5 6 3 8 n ot m ap pe d n ot m ap pe d 4. 1 4. 2 4. 3 5. 1 5. 2 5. 3 s p e c ie s g r o u p a z yg op hy llu m fo et id um + . + + + . . . + . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . e up ho rb ia ( h r p3 5) s p. + 1 + . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s ta pe lia s p. + + . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . a sp ar ag us ( h r p3 78 ) sp . . . . + . + . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . d id el ta s pi no sa . . + + . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . r us ch ia ( h r p3 5) s p. + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v yg ie ( h r p3 78 ) sp . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v yg ie ( h r p3 82 ) sp . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p te ro ni a (h r p3 82 ) sp . . + . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v yg ie ( h r p3 8 4) s p. . 1 . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p b ty le co do n w al lic hi i . . . . . . . . + . . . + . + + . . . . + + . + . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . c ra ss ul a al pe st ri s + . . . . . . . . + + + . + + . . . . . + . + . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r hu s bu rc he lli i + . . . . . r r r . . . . + . . . + . . r . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p hy llo bo lu s sp . r . . . . . . . . . . . + . . . r . + r . . . + . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . m on tin ia c ar yo ph yl la ce a + . a . . . b 1 1 . . . . 1 . . . . . . . b . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p c p te ro ni a gl au ca . . . . . . . . . . . . . . . . r . . . 1 1 1 a 1 + + 1 1 + 1 + 4 b + + + 1 1 + . . . . + + . . . . . . . . r . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p d e up ho rb ia d ec us sa ta . . . . 1 r . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . 1 + 1 1 1 1 + + 1 1 + + 1 + + 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s ar co ca ul on s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a nt im im a cf . g ra ni tic a (h r 24 8) . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . + . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e up ho rb ia h am at a . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p e a rc to tis a ca ul is . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . + . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b ab ia na s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . t he si um c f. hy st ri x . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . r + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . tr ac hy an dr a fa lc at a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h er m an ni a cu ne ifo lia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . la ch en al ia v io la ce a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . o th on na fi lic au lis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . la pe ir ou si a m on ta na . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . la ch en al ia s pp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . + + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p f d or ot he an th us s p. . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . h el io ph ila c ol lin a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . ta b l e 1 a ph yt os oc io lo gi ca l t ab le o f t he s uc cu le nt k ar oo b io m e re la te d ve ge ta ti on o f t he h an ta m -t an qu ar og ge ve ld s ub re gi on – a ss oc ia ti on s 4 an d 5 165 succulent karoo biome related vegetation – part 2 original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 160 183 r e l e v é n u m b e r 3 3 3 3 1 1 1 2 1 1 3 2 2 3 1 1 3 3 3 1 1 2 2 2 2 2 1 2 2 3 3 1 1 2 3 1 1 1 1 3 8 4 7 8 8 2 6 6 4 9 9 8 3 3 4 5 1 5 7 8 8 3 8 4 4 7 1 2 8 0 2 5 5 8 0 5 8 3 2 1 1 4 5 6 0 4 6 8 0 4 1 5 9 9 1 2 3 2 1 1 1 2 3 3 5 1 6 6 6 1 4 4 4 5 1 2 2 2 5 5 7 3 2 2 5 9 5 3 3 7 1 4 7 2 5 5 1 7 0 6 9 1 6 5 1 9 7 8 3 1 7 9 2 9 9 9 5 1 0 2 9 7 4 6 4 2 4 5 6 7 1 8 7 1 2 3 0 3 0 0 0 8 3 4 6 8 1 0 0 0 2 3 3 2 3 1 2 9 7 1 1 5 8 9 9 5 8 9 1 6 6 7 8 5 8 1 8 5 6 3 8 n ot m ap pe d n ot m ap pe d 4. 1 4. 2 4. 3 5. 1 5. 2 5. 3 s en ec io c ak ile fo liu s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . m er xm ue lle ra s tr ic ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . m or ae a sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . z yg op hy llu m p yg m ae um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r + . . . . . . . . . . . . . . . . . . . . + . . . . . lo to no ni s hi rs ut a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . b + . + . . + . . . . . . . . . . . . . . . . . . . . . . . . . o st eo sp er m um ac an th os pe rm um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + + . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p g b ul bi ne lla s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . + . . . . . + . + + . . . . . . . . . . . . . . . . . . . . . . . . . . d im or ph ot he ca s in ua ta . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . + + . . . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . e ro di um c ic ut ar iu m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + + . . + + 1 . + + . . . . . r . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p h e hr ha rt a ca ly ci na . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . + + + . . . . . + + . . . . + + + . . . . . . . . . . . . . . . . . . . . . . c yp hi a di gi ta ta . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . + . . . . + . . . . . . . + + + r . . + + . . . . . . . . . . . . . . . . . m ic ro lo m a sa gi tt at um + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r + + + . + . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . b er kh ey a fr ut ic os a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . + . + . + . . + . . . . . . . . . + r . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p i e ri oc ep ha lu s pu rp ur eu s . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . + . . . . . a . . + . + 1 . a + . a . . . . . . . . 1 . . . . . . a . . . . . te tr ag on ia m ic ro pt er a + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . + + + + + + + . + + + + . . . . . + + . + + . . . . . + + . + + . + . . . . + s en ec io c ar da m in ifo liu s . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . + . . . + + + . . . + . + . . . . . . . . . + . . + . . . + + . + + . . . . . c ot ul a ba rb at a . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . + . + . . . . . . . . . . . . . . . . + . . . . . . . . + . . . . + . . + . + . . + . . . . + . . 1 . . . + b ul bi ne s uc cu le nt a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . + . . . . . . . . . + . . . . + + . . . . . . . . s p e c ie s g r o u p j le ip ol dt ia s ch ul tz ei 1 . + . . . . . . . . . . + 1 . . a . . . . . . . . . + . . . . . . . . . . + . + . 1 + + . . . . . . . . . . + . . . . . . . . . . . . . . r . . 3 . . . . . . + + 1 . + . . . . + . 1 a . . g or te ri a di ff us a + . + . . . . . . . . . + . . + . . . + . . . . . . . . . . . . . . . . + . . . . + + . . . . . . . . . . . . + + + + + + . . . . + . . + . . . . . + . . . . + . . . + + + . + + . . . . 1 . e up ho rb ia m au ri ta ni ca + . + 1 . . . . . . . . + r . . + . . . . . . + . . . . . . + . . . . . . . . . 1 + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . + + . + . + . . . . . . . . . . . s p e c ie s g r o u p k p hy llo bo lu s te nu ifl or us . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . + + . . . . + . . . . . . . . . . . . . . + . . + . . . . . . . b ro m us p ec tin at us . . . . . + . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . + . . . . . + . . + . . . . . . . . . . . . . . . . . . . . . . . . . o nc os ip ho n gr an di flo ru m . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e ri oc ep ha lu s na m aq ue ns is . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . lo to no ni s sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p l p en ta sc hi st is p at ul a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r os en ia g la nd ul os a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . o th on na a ur ic ul ifo lia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . + . . . . . . . . d ro sa nt he m um c f. la tip et al um . . . . . . . . . . . . + . . + . . . + + . . . . . . . . . . . . + . . + . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . + . . . . . . . . . . . + . . . . . . . . . . + . . . . h el ic hr ys um o bt us um . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . + ta b l e 1 a (c o n t. ..) 166 original research van der merwe, van rooyen & van rooyen vol. 50 no. 1 pp. 160 183koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za r e l e v é n u m b e r 3 3 3 3 1 1 1 2 1 1 3 2 2 3 1 1 3 3 3 1 1 2 2 2 2 2 1 2 2 3 3 1 1 2 3 1 1 1 1 3 8 4 7 8 8 2 6 6 4 9 9 8 3 3 4 5 1 5 7 8 8 3 8 4 4 7 1 2 8 0 2 5 5 8 0 5 8 3 2 1 1 4 5 6 0 4 6 8 0 4 1 5 9 9 1 2 3 2 1 1 1 2 3 3 5 1 6 6 6 1 4 4 4 5 1 2 2 2 5 5 7 3 2 2 5 9 5 3 3 7 1 4 7 2 5 5 1 7 0 6 9 1 6 5 1 9 7 8 3 1 7 9 2 9 9 9 5 1 0 2 9 7 4 6 4 2 4 5 6 7 1 8 7 1 2 3 0 3 0 0 0 8 3 4 6 8 1 0 0 0 2 3 3 2 3 1 2 9 7 1 1 5 8 9 9 5 8 9 1 6 6 7 8 5 8 1 8 5 6 3 8 n ot m ap pe d n ot m ap pe d 4. 1 4. 2 4. 3 5. 1 5. 2 5. 3 s p e c ie s g r o u p m e ri oc ep ha lu s pa up er ri m us . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . g az an ia r ig id a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . + . . + . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . e ur yo ps m ul tifi du s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . p en tz ia c f. sp ha er oc ep ha la . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . r os en ia o pp os iti fo lia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p n p en tz ia in ca na . . + . . . + . . . . . . . . . . . + . + . + . . . + + . 1 + . . + . . + + + + + + . . + + . . . . . . . . . . + + + + + . + . . + . . + . + + . + + . + . . . . . + + . . + + . + . . . . . r us ch ia c ra do ck en si s . . . . . . + . . b . . . . . . . . . . . . . . . . . . + . . . . . . . 1 . . . . + . . a a . a . . . 1 1 4 . + . 1 + . + . . . . + . . + . 3 + . + . 1 . . . . . . . . 1 . + . + . . + a . . p te ro ni a gl om er at a . . . . . + . . . . a 1 . . . . . . + . . . . . . . . . . . . . . . . + . . + + . . . . . . 1 . . . . . . . . . . . . . . r . . . . . . . . . . . . + . . . . . . . . . . r . . . . . . . . . m or ae a sp p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . + . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . + . + . . . . + . . . . . + + . . . . . . . . . . . + . . . . a e ri oc ep ha lu s er ic oi de s + . + . . . . . . . . . . . . . . . + . . . . + . 1 . . . . + b . + . 1 . . . 1 + . . . . . . . . . . . . . . . . . . . . . b . . . . . . 1 . . . . . . . . . . . . . . + + . . . . . . . . . le ys er a te ne lla . . + . . . . . . . . . + . . + + . . + . . . . . . . . . . . . . . . . . . + . . . + + . . . . . + + . + . + . . . . . . . . . . . . . + + . . . . . . + . . . + + + . . . . . + . . . . 1 + k ar ro oc hl oa s ch is m oi de s . . . . . . + . . . + . . . . . 1 . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a rc to th ec a ca le nd ul a . . . . . . . . . . + . . . + . + . . . . . . . . . . . . . . . . + + . . + . . . . . . . . . . . + . . . . . . + . + + + . + . + + + . . + . . . . + . + + . . . . + . . + . + + . + . . . + a sp ar ag us c ap en si s . . . . . . r . . . . . . . + . . . + . . + + . . + . . r . . + . . . . + . + . . . . . + . . . . . + . . + . . + . . + + + + . . + . + + . + . . r . r + + . . . . + . . . + . . . . . . . + h ir pi ci um a lie na tu m . . . . . . . . + . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . r . . . . . . . . + + . + . . . . . . . . . . . . . . . . . . . . . te tr ag on ia fr ut ic os a . . . . . + . . . . . . . . + + . . . . . . . . . . . + . + . . . . + . . + . . . . + . . . + . . . + . + . . r + . . . . . . . . . . + . . . + + + . . . r . + . . + . + . . . . . . . r . . e ri oc ep ha lu s m ic ro ph yl lu s . . . . . . . . . . . . . . . . . . . . . . r . . . . . + . . . . . . . + + . . . . . . + . . . . . . . . . . + . 1 . + . + . . . . . . + . . . + + . . + 1 . . . . . . . . + . 1 + . . . . 1 t he si um li ne at um + . . . . + r r . . . . . . . . . . . . . + . . . + r . . . . . . . . . . . . . . + . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . + . . r + . . . . . . . . . . . . . . . . . . c hr ys oc om a ci lia ta . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . + . . . . . . . . . . . . . . . . . . + . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . s p e c ie s g r o u p o r us ch ia r ob us ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m el ol ob iu m c an di ca ns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . tr ib ol iu m h is pi du m . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . + . . . . . + 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + a ug ea c ap en si s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e ur yo ps a nn uu s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . + . . . . . . . . s p e c ie s g r o u p p c ep ha lo ph yl lu m s p. . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . p te ro ni a vi llo sa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . f el ic ia ( h r p1 71 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . f el ic ia s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . + . . . . . . . . . . . . . . . p te ro ni a gl ab ra ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p q g az an ia li ch te ns te in ii . . . . . . . . . . . . . . + . + + . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . + . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . s p e c ie s g r o u p r d ro sa nt he m um ( h r 21 9) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . + . . . m al ep ho ra c ra ss a . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . + . . . . g al en ia fr ut ic os a . . . . . + . . r . . . . . + . . . . . . . . . . . . . . . . . . . . . . . + . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . ta b l e 1 a (c o n t. ..) 167 succulent karoo biome related vegetation – part 2 original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 160 183 r e l e v é n u m b e r 3 3 3 3 1 1 1 2 1 1 3 2 2 3 1 1 3 3 3 1 1 2 2 2 2 2 1 2 2 3 3 1 1 2 3 1 1 1 1 3 8 4 7 8 8 2 6 6 4 9 9 8 3 3 4 5 1 5 7 8 8 3 8 4 4 7 1 2 8 0 2 5 5 8 0 5 8 3 2 1 1 4 5 6 0 4 6 8 0 4 1 5 9 9 1 2 3 2 1 1 1 2 3 3 5 1 6 6 6 1 4 4 4 5 1 2 2 2 5 5 7 3 2 2 5 9 5 3 3 7 1 4 7 2 5 5 1 7 0 6 9 1 6 5 1 9 7 8 3 1 7 9 2 9 9 9 5 1 0 2 9 7 4 6 4 2 4 5 6 7 1 8 7 1 2 3 0 3 0 0 0 8 3 4 6 8 1 0 0 0 2 3 3 2 3 1 2 9 7 1 1 5 8 9 9 5 8 9 1 6 6 7 8 5 8 1 8 5 6 3 8 n ot m ap pe d n ot m ap pe d 4. 1 4. 2 4. 3 5. 1 5. 2 5. 3 s p e c ie s g r o u p s b ra un si a cf . a pi cu la ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b ro w na nt hu s cf . v ag in at us . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p te ro ni a sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cf . l am pr an th us ot ze ni an um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s al so la s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p en tz ia s pi ne sc en s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p t a tr ip le x lin dl ey i . . . . . . . . . . . . + . . + . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . + . . . . . + . + . . . . . . . . . . . . p hy llo bo lu s sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . p si lo ca ul on ( p9 1) s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . s al so la a ph yl la . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . e xo m is m ic ro ph yl la . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p u r us ch ia in tr ic at a . . . . . . . . . . . 1 . . . . . . 1 . . . + . . . + . . . . . . . . . . . . 1 . . . + . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . 1 + f el ic ia a us tr al is . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . + . . e ri oc ep ha lu s de cu ss at us . . . 1 . . . . . . + . . . . . + . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . + . . + . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . + . . . . . . . f ov eo lin a di ch ot om a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . z yg op hy llu m ( h r p6 5) s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . + . . . . . . . . . . . . . . . . . s p e c ie s g r o u p v a ri da ri a no ct ifl or a . . . . . . . . . . . . . . . . . + . + . . . . . . . . . + . . . + 1 + . . . . . + + . . . . . . . . . . + + . + . + . . . + . . . . . . . . + + . + + . + 1 . . . . + . + + . + 1 + . . . . g al en ia s ar co ph yl la . . + . . + . . . . . . . . . + . . . . . . . . . . . . . . . . . + . . . . . . . + + . . . . . . + . . . + . . . . . . + . . . . . b . . . . . . . . + . . . . + . . . + + . . . . + . . . + ly ci um s pp . . . . . . . . . . . r r . . . . . . . . . . . . . . . . . . + . . r + . . . . + . . . . . + . . . + . . . . . . . . . . + . r . . + . . + . . . . . . + . + . . + . . + + . + . . . . . . . . s al so la tu be rc ul at a . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . r + . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . + . . . . r . . . . tr ip te ri s si nu at a . . . . . + . . + . . . . . + + + . . . + . . + . . . + . . + . . . . + + . . + + + . + . . + + . . + . + + . . . . . . . . . . . . . . . . . . . . . . + . + . + . . + + + + + + . + . . . . d ro sa nt he m um (h r 21 7) s p. . . . . . . . . + + r . + . + + + . . . . . . . . . . . . + + . . + 1 + . . + . + + + . . + + . . . . . . + + + . . . . . . . . . + . . . . . . . . + . . + 1 + . + + + . + . + + . + + . . . p te ro ni a pa lle ns . . . . . . + . . . . . . . + . . . . . . . + . + . 3 . . . . . . . + . 1 . . . . . . . + . . . . . . . . . a . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . + 1 . . . . . . 1 . . g eo ph yt ic s pp . . . + . . + . . + . + + . . + + + + + + + . . + . . . . . . . . + + . + + . + . + + + + . . . . . + . . + . . . + + . + + + . 1 + + + + + . + . + . + + + . . + + . + + . . + + . + + r . . + o xa lis s pp . . . + . . . + . + . + + . . + . + + + . . + . + . . . . + + . + + + + + . . + . + . . + . . . . . + + . + . + + + + + + + + + . + + 1 . + . + + + + + . + + + + + + + + + . + + + . + + + . + g al en ia a fr ic an a + + + + + + . + . . r . + . + . + . . . + + . . . . + . + + + + . + + . . + + . . . + . . + . + + + . . + . . . + . . . r . . . . . . . . . + . . + . . . + . . . . . + + . . . + . + . . . . m es em br ya nt he m um gu er ic hi an um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . + . . . . . . . . . . . . . . + . . . + . . . . . . + + . + + . . . . + . . . . + . . . . . . . . . . . . . . . . . . . . o st eo sp er m um p in na tu m . . + . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . + . . . . + . . . . . . . . . + . + . . + . . . + . + + . . + . . + . . s p e c ie s g r o u p w c la do ra ph is s pi no sa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p x s tip ag ro st is o bt us a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . z yg op hy llu m r et ro fr ac tu m . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . + . . . . . . + . . . . . . . . . . . . . . . + . . . . . . . . + . . . . . . . . . . . s tip ag ro st is b re vi fo lia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . s tip ag ro st is c ili at a . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ta b l e 1 a (c o n t. ..) 168 original research van der merwe, van rooyen & van rooyen vol. 50 no. 1 pp. 160 183koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za r e l e v é n u m b e r 3 3 3 3 1 1 1 2 1 1 3 2 2 3 1 1 3 3 3 1 1 2 2 2 2 2 1 2 2 3 3 1 1 2 3 1 1 1 1 3 8 4 7 8 8 2 6 6 4 9 9 8 3 3 4 5 1 5 7 8 8 3 8 4 4 7 1 2 8 0 2 5 5 8 0 5 8 3 2 1 1 4 5 6 0 4 6 8 0 4 1 5 9 9 1 2 3 2 1 1 1 2 3 3 5 1 6 6 6 1 4 4 4 5 1 2 2 2 5 5 7 3 2 2 5 9 5 3 3 7 1 4 7 2 5 5 1 7 0 6 9 1 6 5 1 9 7 8 3 1 7 9 2 9 9 9 5 1 0 2 9 7 4 6 4 2 4 5 6 7 1 8 7 1 2 3 0 3 0 0 0 8 3 4 6 8 1 0 0 0 2 3 3 2 3 1 2 9 7 1 1 5 8 9 9 5 8 9 1 6 6 7 8 5 8 1 8 5 6 3 8 n ot m ap pe d n ot m ap pe d 4. 1 4. 2 4. 3 5. 1 5. 2 5. 3 b le ph ar is p ru in os a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . z yg op hy llu m ( h r p3 86 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p y m es em br ya nt he m ac ea e 1 (h r p3 59 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m es em br ya nt he m ac ea e 2 (h r p3 59 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p z p te ro ni a (h r p1 18 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ly ci um ( h r p1 18 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s al so la ( h r p1 18 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p te ro ni a (h r p1 18 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * n on -d ia gn os tic s pe ci es a re e xc lu de d * h r c ol le ct io n co de a nd n um be rs a re in cl ud ed fo r fu tu re r ef er en ce , i f n ec es sa ry * s pe ci m en s in ad eq ua te fo r id en tifi ca tio n ye t d iff er en t f ro m s pe ci es th at c ou ld b e id en tifi ed a re in di ca te d w ith a p lo t n um be r (p … ) * sp . = o ne s pe ci es in a ta xo no m ic g ro up * sp p. = m or e th an o ne s pe ci es in a ta xo no m ic g ro up . t he se s pe ci es , e ve n al th ou gh th ey a re g ro up ed to ge th er , a re in cl ud ed in th e ta bl e si nc e th ey o cc ur in d iff er en t s pe ci es g ro up s. h ow ev er , t he y ar e no t u se d in th e de sc ri pt io ns in th e te xt . ta b l e 1 a (c o n t. ..) 5.3 leipoldtia schultzei – eriocephalus purpureus hantam karoo this subassociation is located around the farms soetwater and leeuriet southwest to mensieskraal and stretches southeastwards to approximately the farm soutpan (fig. 1). it covers an area of 58 214 ha (3.5% of the total area in fig. 1). subassociation 5.3 occurs on dwyka tillites and ecca shales at an altitude ranging from 300 to 1 000 m, and the dominant land types are da and fc. the ridges of this subassociation are generally level, occasionally gentle, with a low rock cover. the loamy soils are red brown or light brown in colour. shrub cover in this subassociation varies from 45 to 80%. the grass component is absent while the annual component varies greatly, from < 1% to 60%. subassociation 5.3 shares species groups i and j with subassociations 5.1 and 5.2 but lacks its own defining group (table 1a). the dominant species are leipoldtia schultzei and gorteria diffusa (species group j), while other species describing this subassociation include pentzia incana, ruschia cradockensis and eriocephalus microphyllus (species group n), as well as aridaria noctiflora, tripteris sinuata, drosanthemum (hr217) sp. and occasionally a high cover of pteronia pallens (species group v). 6. pteronia glomerata roggeveld karoo this association is predominantly found along the eastern boundary of the study area (fig. 1) and has been termed roggeveld karoo. the ecca group, characterised by shales, and the beaufort group, characterised by mudstones, meet in this transitional area. additionally, this area is the transition between the winter rainfall region to the west and the summer rainfall region to the east. in general, this association occurs at an altitude greater than 1 000 m above sea level and predominantly on land types fc or da. the level to gently sloping ridges, plains and valleys commonly have light brown or brown loamy soils. a high shrub cover (usually > 60%) is found, whereas the grass and annual components are absent or < 5%. association 6 (table 1b) is related to associations 4 and 5 (table 1a) through common species such as pentzia incana, ruschia cradockensis and pteronia glomerata (species group n), but also related to association 7 (table 1c) through common species such as ruschia intricata and eriocephalus decussatus (species group u). 6.1. phyllobolus tenuiflorus – pteronia glomerata roggeveld karoo located in the vicinity of the farms rooiwal and weltevrede on dolerite soils, southeast of calvinia (fig. 1), subassociation 6.1 is found predominantly on land types fc and ag, and covers an area of 9 172 ha (0.6% of the total area in fig. 1). it is generally found on level terrain to gently sloping ridges and plains at an altitude varying from 1 000 to 1 300 m above sea level. soils are loamy and usually brown, with a rock cover that ranges from zero to 80%, comprising small stones and stones. shrub cover in this subassociation varies from 5 to 90%. the grass component is generally absent except in isolated cases where it can be as high as 25 to 40%. the annual component varies considerably from < 1% to 60%. subassociation 6.1 comprises species groups k, n, u and v but lacks species groups l and m, which are present in subassociations 6.2, 6.3 and 6.4 (table 1b). annual species include phyllobolus tenuiflorus (species group k), felicia australis and foveolina dichotoma (species group u), and galenia sarcophylla (species group v), while perennial species include pentzia incana, ruschia cradockensis and pteronia glomerata (species group n) (table 1b). 6.2. eriocephalus pauperrimus – pteronia glomerata roggeveld karoo this subassociation is located within the basterberg mountains (fig. 1) and covers an area of 50 790 ha (3.1% of the total area 169 succulent karoo biome related vegetation – part 2 original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 160 183 in fig. 1). it is predominantly found on land types da and fc at an altitude of 1 100 to 1 400 m. these level to gently sloping mudstone ridges of the beaufort group are covered with brown or light brown coloured loamy soils with a rock cover varying from zero to 95% stone. the shrub cover is high while the grasses and annuals make little contribution to the overall plant cover. conspicuous species in this subassociation include phyllobolus tenuiflorus (species group k) and eriocephalus pauperrimus (species group m) (table 1b). other common species are pentzia incana, ruschia cradockensis, pteronia glomerata, karroochloa schismoides (species group n), eriocephalus decussatus (species group u) and tripteris sinuata (species group v). 6.3. pentzia incana – pteronia glomerata roggeveld karoo subassociation 6.3 is located in the vicinity of elandsoog se berg, around the farm de puts, southwest of the droëberg mountains and northeast of loeriesfontein (fig. 1), and covers an area of 31 594 ha (1.9% of the total area in fig. 1). this subassociation is found on mudstones of the beaufort group and shales of the ecca group, and predominantly on land type da. the altitude ranges from 1 000 to 1 200 m and the subassociation is found on relatively level areas of ridges, plains and valleys. generally, the light brown loamy soils have little rock cover but when rock occurs it is in the form of stones or boulders. the shrubby component in this subassociation is well developed but grasses and annuals are usually absent or contribute very little to the overall cover. eriocephalus pauperrimus (species group m) has a high cover in this subassociation, which is closely related to subassociations 6.2 and 6.4 through the presence of species group m (table 1b). other species present include pentzia incana and eriocephalus ericoides (species group n) and ruschia intricata (species group u), as well as various lycium species (species group v). 6.4. ruschia intricata – pteronia glomerata roggeveld karoo subassociation 6.4 is located north to south along the eastern boundary of the study area, predominantly on mudstones of the beaufort group (fig. 1) and, excluding the mosaic vegetation units, covers an area of 265 394 ha (16% of the total area in fig. 1). it forms the transition with the arid karoo and false succulent karoo, as described by acocks (1953, 1988). elements of this subassociation are also found in the transition with the namaqualand broken veld (acocks 1953, 1988), west of the hantam mountain, in the naresie mosaic vegetation unit, predominantly on dwyka tillite. this subassociation is found on land types fc and da at an altitude ranging from 800 to 1 400 m above sea level. the level terrain and gentle slopes are usually covered in light brown or brown coloured soils with either a low rock cover (0–15%) or a high rock cover (80–95%). shrub cover usually ranges from 50 to 90%, while grass and annual cover is usually absent or < 5%. common species include rosenia glandulosa (species group l), eriocephalus pauperrimus (species group m), pentzia incana, ruschia cradockensis, pteronia glomerata, eriocephalus ericoides (species group n), ruschia intricata, eriocephalus decussatus (species group u), aridaria noctiflora and tripteris sinuata (species group v) (table 1b). 6.5. eriocephalus ericoides – pteronia glomerata roggeveld karoo this subassociation, located on land type fc, is found northeast and southwest of the hantam mountain (fig. 1) and, excluding the mosaic vegetation units, covers an area of 69 356 ha (4.2% of the total area in fig. 1). it is also located in a mosaic vegetation unit around the town of calvinia (calvinia mosaic) and in the adamsfontein/noodsaaklikheid mosaic. geologically, the ecca group dominates these areas, with dolerite intrusions interrupting the shales. the subassociation occurs at altitudes ranging from 900 to 1 200 m above sea level on level to gently sloping ridges, with generally a higher rock cover than in subassociations 6.1, 6.2 and 6.3. the shrub layer covers 40 to 80% of the light brown and brown loamy soils, while the grass and annual component is either absent or covers < 5%. subassociation 6.5 differs from the previous subassociations because it lacks species groups k, l and m (table 1b). dominant species in this subassociation are pentzia incana, pteronia glomerata, eriocephalus ericoides (species group n), ruschia intricata (species group u), aridaria noctiflora, tripteris sinuata and galenia africana (species group v). these species are all common to the entire association 6. 7. aridaria noctiflora tanqua karoo association 7 is located primarily in the tanqua karoo basin and the adjacent foothills of the koedoesberg mountains. large areas east and northeast of loeriesfontein, as well as on the western extreme of the hantam river, a brackish system, are also included in this association, as well as the vegetation mosaic located around the town of calvinia (fig. 1). geologically, shale of the ecca group and tillite of the dwyka group dominate association 7, while alluvial deposits are present at times. limited intrusions of dolerite also occur. this association is found predominantly on land types ia and fc at an altitude ranging from 200 to 1 100 m. the level to gently sloping ridges and plains occur on brown and light brown soils, with generally a low rock cover. shrub cover is intermediate (ca. 40%), while the grass and annual components are absent or poorly developed. the annual component is expected to be much higher in normal and good rainfall years. association 7 (table 1c) differs from associations 4 to 6 (table 1a, 1b) as it lacks species group n but shares species groups u and v with association 6 (table 1b). associations 4 to 7 (table 1a, 1b, 1c) are related through the shared presence of species groups v, and have no species groups in common with associations 8, 9 and 10 (table 1c). common species in association 7 are ruschia intricata (species group u), aridaria noctiflora, salsola tuberculata, tripteris sinuata and drosanthemum (hr217) sp. (species group v). this association is subdivided into five subassociations (table 1c). 7.1. ruschia robusta – aridaria noctiflora tanqua karoo subassociation 7.1 is located at the eastern edge of the tanqua basin at the foot of the roggeveld escarpment and in the vicinity of the farm twee damme, and covers an area of 79 395 ha (4.8% of the total area in fig. 1). this low-lying (300–800 m above sea level) subassociation is found on the ecca group on gently sloping ridges and plains with brown coloured loamy soils of land types ia and fc. rock cover varies considerably, from no rocks to 50 to 99% consisting of small stones (> 10–50 mm). shrub cover is generally low, but can be as high as 75%. grass cover is low or absent, while annual cover ranges from very low (the norm) to isolated dense patches (50–80%). this subassociation is characterised by perennials such as ruschia robusta, augea capensis and the annual euryops annuus (species group o) (table 1c). other perennial species are aridaria noctiflora, drosanthemum (hr217) sp. and pteronia pallens (species group v), and annual species include gazania lichtensteinii (species group q) and felicia australis (species group u). 7.2. drosanthemum (hr217) sp. – aridaria noctiflora tanqua karoo subassociation 7.2 is located on the alluviums in the vicinity of the farms weltevreden and platfontein, southwest of calvinia (fig. 1) and, excluding the mosaic vegetation units, covers an 170 original research van der merwe, van rooyen & van rooyen vol. 50 no. 1 pp. 160 183koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za r e l e v é n u m b e r 2 1 1 2 2 2 2 2 1 1 2 2 2 2 2 2 2 3 3 2 3 2 1 2 2 2 2 1 2 2 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 1 2 2 1 1 1 2 3 2 3 1 2 2 3 2 2 2 7 0 0 0 1 1 2 5 9 5 1 6 6 6 9 9 7 9 0 2 6 3 5 7 7 7 7 8 3 2 3 7 7 3 0 7 5 7 3 3 4 4 4 6 8 0 2 3 5 6 6 6 6 9 9 9 0 0 0 1 1 2 2 0 4 0 5 6 8 5 5 1 7 6 4 0 6 3 1 5 7 6 7 0 1 2 0 2 9 0 6 1 8 7 9 7 7 6 7 8 0 3 6 4 1 8 3 1 9 8 3 4 7 0 4 9 0 7 8 5 2 6 8 9 6 7 0 1 2 0 8 4 0 1 6 1 2 4 5 6 7 8 2 4 8 8 9 2 7 3 3 3 5 1 3 6 8 2 8 5 0 6 0 7 9 0 4 4 4 0 6 3 5 6. 1 6. 2 6. 3 6. 4 6. 5 s p e c ie s g r o u p a z yg op hy llu m fo et id um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e up ho rb ia ( h r p3 5) s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s ta pe lia s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a sp ar ag us ( h r p3 78 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . d id el ta s pi no sa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r us ch ia ( h r p3 5) s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v yg ie ( h r p3 78 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v yg ie ( h r p3 82 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p te ro ni a (h r p3 82 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v yg ie ( h r p3 8 4) s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p b ty le co do n w al lic hi i . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . c ra ss ul a al pe st ri s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . r hu s bu rc he lli i . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p hy llo bo lu s sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m on tin ia c ar yo ph yl la ce a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p c p te ro ni a gl au ca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p d e up ho rb ia d ec us sa ta . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . s ar co ca ul on s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a nt im im a cf . g ra ni tic a (h r 24 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e up ho rb ia h am at a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p e a rc to tis a ca ul is . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b ab ia na s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . t he si um c f. hy st ri x . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . tr ac hy an dr a fa lc at a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h er m an ni a cu ne ifo lia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . la ch en al ia v io la ce a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . o th on na fi lic au lis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . la pe ir ou si a m on ta na . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . la ch en al ia s pp . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p f d or ot he an th us s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h el io ph ila c ol lin a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s en ec io c ak ile fo liu s . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ta b l e 1 b ph yt os oc io lo gi ca l t ab le o f t he s uc cu le nt k ar oo b io m e re la te d ve ge ta ti on o f t he h an ta m -t an qu ar og ge ve ld s ub re gi on – a ss oc ia ti on 6 171 succulent karoo biome related vegetation – part 2 original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 160 183 r e l e v é n u m b e r 2 1 1 2 2 2 2 2 1 1 2 2 2 2 2 2 2 3 3 2 3 2 1 2 2 2 2 1 2 2 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 1 2 2 1 1 1 2 3 2 3 1 2 2 3 2 2 2 7 0 0 0 1 1 2 5 9 5 1 6 6 6 9 9 7 9 0 2 6 3 5 7 7 7 7 8 3 2 3 7 7 3 0 7 5 7 3 3 4 4 4 6 8 0 2 3 5 6 6 6 6 9 9 9 0 0 0 1 1 2 2 0 4 0 5 6 8 5 5 1 7 6 4 0 6 3 1 5 7 6 7 0 1 2 0 2 9 0 6 1 8 7 9 7 7 6 7 8 0 3 6 4 1 8 3 1 9 8 3 4 7 0 4 9 0 7 8 5 2 6 8 9 6 7 0 1 2 0 8 4 0 1 6 1 2 4 5 6 7 8 2 4 8 8 9 2 7 3 3 3 5 1 3 6 8 2 8 5 0 6 0 7 9 0 4 4 4 0 6 3 5 6. 1 6. 2 6. 3 6. 4 6. 5 m er xm ue lle ra s tr ic ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m or ae a sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . z yg op hy llu m p yg m ae um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lo to no ni s hi rs ut a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . o st eo sp er m um a ca nt ho sp er m um . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p g b ul bi ne lla s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . d im or ph ot he ca s in ua ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e ro di um c ic ut ar iu m . . . . . . + . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . s p e c ie s g r o u p h e hr ha rt a ca ly ci na . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . c yp hi a di gi ta ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m ic ro lo m a sa gi tt at um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b er kh ey a fr ut ic os a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p i e ri oc ep ha lu s pu rp ur eu s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . te tr ag on ia m ic ro pt er a + . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . s en ec io c ar da m in ifo liu s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . c ot ul a ba rb at a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . b ul bi ne s uc cu le nt a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p j le ip ol dt ia s ch ul tz ei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . g or te ri a di ff us a . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e up ho rb ia m au ri ta ni ca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . s p e c ie s g r o u p k p hy llo bo lu s te nu ifl or us + + + + + . + . + . + + + + + + + . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b ro m us p ec tin at us . . . . . . + . + + . . . + + . . + . . + . . . . . . . . . . + . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . o nc os ip ho n gr an di flo ru m . . + . . + . . . . . . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e ri oc ep ha lu s na m aq ue ns is . a . + . . . 1 . . . . a . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lo to no ni s sp . . . . . . . . . . . . + . + + + . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p l p en ta sc hi st is p at ul a . . . . . . . . . . . . . . . . . . . . . . . . + + . . + . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r os en ia g la nd ul os a . . . . . . . . + . . . . + . . . . . . . . . 1 . . . . . + + . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . o th on na a ur ic ul ifo lia . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . + . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . d ro sa nt he m um c f. la tip et al um . . . . . . . . . . . . . . . . . . . . . . . + + . + . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h el ic hr ys um o bt us um . + . . . . . . . + . . + . . . . . . . . . . + . . . . . . . + + . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ta b l e 1 b (c o n t. ..) 172 original research van der merwe, van rooyen & van rooyen vol. 50 no. 1 pp. 160 183koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za r e l e v é n u m b e r 2 1 1 2 2 2 2 2 1 1 2 2 2 2 2 2 2 3 3 2 3 2 1 2 2 2 2 1 2 2 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 1 2 2 1 1 1 2 3 2 3 1 2 2 3 2 2 2 7 0 0 0 1 1 2 5 9 5 1 6 6 6 9 9 7 9 0 2 6 3 5 7 7 7 7 8 3 2 3 7 7 3 0 7 5 7 3 3 4 4 4 6 8 0 2 3 5 6 6 6 6 9 9 9 0 0 0 1 1 2 2 0 4 0 5 6 8 5 5 1 7 6 4 0 6 3 1 5 7 6 7 0 1 2 0 2 9 0 6 1 8 7 9 7 7 6 7 8 0 3 6 4 1 8 3 1 9 8 3 4 7 0 4 9 0 7 8 5 2 6 8 9 6 7 0 1 2 0 8 4 0 1 6 1 2 4 5 6 7 8 2 4 8 8 9 2 7 3 3 3 5 1 3 6 8 2 8 5 0 6 0 7 9 0 4 4 4 0 6 3 5 6. 1 6. 2 6. 3 6. 4 6. 5 s p e c ie s g r o u p m e ri oc ep ha lu s pa up er ri m us . . . . . . . . . . . 1 . a a . . . + . 1 . 1 1 + + + . . a . + + . . . + . . . . + + + . . 1 . . 1 + . 1 . . . . . . . . . . + 1 . + . . . . . . . . . . . . . . . . . . . . g az an ia r ig id a . . . . . . . . . . . . + . + . + + . . . . . + . + + + + . . . . . . + . + . + . + . . + + . + . + + r . . . . + . . . . . . . + + . + . . + . . . . . . . . . . . . . . . . e ur yo ps m ul tifi du s . . . . . . . . . . . . . . . . . . + . . + + . . . . . . . . . . . . . . . + . . . . . . . . . + . . + . + + . . r + 1 r + . . . . . . . + . . . . . . . . . . . . . . . . . p en tz ia c f. sp ha er oc ep ha la . . . . . . . . . . . . . + . . . . . + . + + . . . . . . . + . . . . . . . . . . . 1 . . . . . . . . . + . . + . . + . . + + . . . . . . . . . . . . . . . . . . . . . . . . r os en ia o pp os iti fo lia . . . . . . . . . . . . . + . . + . . 1 . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . + . . . . + + + . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p n p en tz ia in ca na . . . + . . . + 1 . . . . . . . + + + . + . . + . . . + 1 + . + 1 . . . . 1 + + + + . + + + + + 1 + + + + + . . 1 1 . + a + . . + + . 1 . + . + . + . . + + + + + + . + + + . r us ch ia c ra do ck en si s . . . . . . . . . b 1 . . . . . . a 1 . + . + . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . 1 + + + . . . . . . . . . . . . + . . . . . . + . a + . . . . . . . . . p te ro ni a gl om er at a + . . . . . + + 1 . . . r + . . + + . + + + . . + + . + . + + a . . + + . . + + . . . . + + . . . + . + + . . + + . . + . + + . + . . . . + + + . . . 1 + . + . + . . + . + . m or ae a sp p. + . . . . + . + + + . + 1 1 + 1 + . . . + . . . . . . 1 + . . + + + . + . + . . + . . . . . . . + . + . + . + . + + . . . + . + + + . . + . . . . . . . . + . . . . . . + . . e ri oc ep ha lu s er ic oi de s + . . . . + . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . + + . . . . . . . . . . . + + . 1 . . 1 + + + . . . . . . . + + + . . . . . . 1 + + . . . . . 1 . le ys er a te ne lla . . . . . . . . . + . + . . + + . . + . . . . . . . . + + . . + + + . . . . . . . + + . . . . . + . . . . . . . . + . . . . . . . . . . . . . . . . . . . . + . . . + . . . . k ar ro oc hl oa s ch is m oi de s + . . . + . . . + . . . + + + + + + + . . + . . . . . . + . . . . + . . + . . . + . . . . . . . + . + . . . . . . + + + + . + . . . + . . . . . + . . . + . + . . . . . . . . a rc to th ec a ca le nd ul a . + . . . . + . . . . . + . . . . . . . . . . + . . . + . . . + + . . + . . . . + . . . . . . . . . + . . . . . . . . . . . . . + + . . . . . . . . . . . + . . . . . . . . . a sp ar ag us c ap en si s + . . . + . . . . . + . . . . . . . . . . . . . . . . . . . . . . r + . . . . + . . . + + . . . . . . + . . . + . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . + h ir pi ci um a lie na tu m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . + + . . . + . . . . . . + . . . . te tr ag on ia fr ut ic os a . . . . . . . . + . + . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . + . . . . . . . + . . . . . . . . . . . . . . . . . . + . . . . . . . . r . . . . e ri oc ep ha lu s m ic ro ph yl lu s . . . . . . . . . 1 . . . . . a . . . + . . . . . . . . . . . . . . 1 . . . . . 1 . . . . . . . . . . . . . . + . . . + . . . . . + . . . . . . . . . . . . . . . . . 1 . . . t he si um li ne at um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . + . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . + . . . + . c hr ys oc om a ci lia ta . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . + . . . . . . + + . . . . . . . s p e c ie s g r o u p o r us ch ia r ob us ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m el ol ob iu m c an di ca ns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . tr ib ol iu m h is pi du m . a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a ug ea c ap en si s . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e ur yo ps a nn uu s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p p c ep ha lo ph yl lu m s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p te ro ni a vi llo sa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . f el ic ia ( h r p1 71 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . f el ic ia s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p te ro ni a gl ab ra ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p q g az an ia li ch te ns te in ii . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p r d ro sa nt he m um ( h r 21 9) s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m al ep ho ra c ra ss a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . g al en ia fr ut ic os a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ta b l e 1 b (c o n t. ..) 173 succulent karoo biome related vegetation – part 2 original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 160 183 r e l e v é n u m b e r 2 1 1 2 2 2 2 2 1 1 2 2 2 2 2 2 2 3 3 2 3 2 1 2 2 2 2 1 2 2 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 1 2 2 1 1 1 2 3 2 3 1 2 2 3 2 2 2 7 0 0 0 1 1 2 5 9 5 1 6 6 6 9 9 7 9 0 2 6 3 5 7 7 7 7 8 3 2 3 7 7 3 0 7 5 7 3 3 4 4 4 6 8 0 2 3 5 6 6 6 6 9 9 9 0 0 0 1 1 2 2 0 4 0 5 6 8 5 5 1 7 6 4 0 6 3 1 5 7 6 7 0 1 2 0 2 9 0 6 1 8 7 9 7 7 6 7 8 0 3 6 4 1 8 3 1 9 8 3 4 7 0 4 9 0 7 8 5 2 6 8 9 6 7 0 1 2 0 8 4 0 1 6 1 2 4 5 6 7 8 2 4 8 8 9 2 7 3 3 3 5 1 3 6 8 2 8 5 0 6 0 7 9 0 4 4 4 0 6 3 5 6. 1 6. 2 6. 3 6. 4 6. 5 s p e c ie s g r o u p s b ra un si a cf . a pi cu la ta . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . + + . . + . . . . . . . . . . . . . + . . . . . + . . . . . . . + . . . . . . . . . . . . . . + . 1 1 . . . . . . b ro w na nt hu s cf . v ag in at us . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p te ro ni a sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . cf . l am pr an th us o tz en ia nu m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s al so la s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p en tz ia s pi ne sc en s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . s p e c ie s g r o u p t a tr ip le x lin dl ey i . . . . + . . . . . + . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . 1 . . . . . . . . . . . . p hy llo bo lu s sp . . . . . . . . + . . + . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p si lo ca ul on ( p9 1) s p. . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . + . . s al so la a ph yl la . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e xo m is m ic ro ph yl la . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . s p e c ie s g r o u p u r us ch ia in tr ic at a + . . . . . . + . . . . . . . . . . . . . . . + . . + + + . . . + . . . . + 1 1 . 1 . 1 1 1 . . . . a . . . + . + . 1 . . + . . + . . . a 1 1 a . . . . . . 1 . + 1 . + . . 1 f el ic ia a us tr al is . . + . . + + . + . . . . + . . + . . . + . . + . + + . + + . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . e ri oc ep ha lu s de cu ss at us . . . . . . . . . . . . . . . . 1 + + . . . . . . . 1 1 + . . . . + . 1 . . . . . + . . 1 + . + + + + . . + 1 . + . . . + a . . + . . . . . . + . . . . + . . + + + . . . . . f ov eo lin a di ch ot om a . . . + + + + . . . . + . . . . . . + . + . . . . . . . . . . . . . . . + . . . . . . . . . . . + . + . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . z yg op hy llu m ( h r p6 5) s p. . . . . . . + . . . . . . . . . . . . . . . . . . . . . . + + . . . + . 1 . . . . . . . . . . + + . . . . + . . . . . . . . . . . . + . . . . . . . . . . . . + . . . . . . + s p e c ie s g r o u p v a ri da ri a no ct ifl or a . . . . . . + . . . . . . . . . . + . . . . . . . + . . . . . . . r . + . r r . + . . . + + + + . . . . . . + + + . . . . + . . + . . + . . . . . + . + . . + . . + . . + + . g al en ia s ar co ph yl la . . . + . . + . + . + . + . + . . . . . . . . + . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . + + + . . . . . . . + + . . . ly ci um s pp . . r r . . + . . . . . . . + + + + + . . + + + + r . . + . + + + . + + + + + + + . + . + + + + + 1 + r + . + + . . + + + + . + . . 1 + . . . . . . + . . . + + + + . r r . . + s al so la tu be rc ul at a . . . . . . + . . . . . . . . . . . . + . . . + . . . . r r + + . . . . . . . . . + + . . . . . . . r . . . . . + . . . . . + . . 1 . r . . . . + + . . . . . + . . . . . . + tr ip te ri s si nu at a . . + . + + . . . . + . . . . . + + + . r . + . . . . r + + + . . . . . . . + + . . + . + 1 + + . + r + + . . . . + . . . . . . . . + + . + + + . + . . . . + + . + + + . + . d ro sa nt he m um ( h r 21 7) s p. . . . . . . a + . . . . . . . . . + . . . . + . + . . . + + . . . . . . . . . . + . . . . + . . + + . . . . . . . . . . . . . . . + + . . + . . . + . . . . . . . . . . + . . p te ro ni a pa lle ns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . g eo ph yt ic s pp . . + . . . + + . . + + . . . . . + . . . . + . + + + + + . + . . . + . + . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . + . . . . . . + . . . . . . . + . . + . o xa lis s pp . . . . . . . . . . . + . . . . . . . . + . . . + . . . + + . . . . . . + . + . + . . . . + . . . . . . . . . . . . . . . . . . . . . . . + + . + . + + . . + . . . . . + + + . g al en ia a fr ic an a . . . . . . . . . r r . . . . . . . . . . . . . . . . . . . . . . . . . . . + . + . + . . + . . . . . . . . . . . . . . . . . . . . . . . . . + + . . + . . . + . . . + + . . m es em br ya nt he m um gu er ic hi an um . . . + . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . + . . . . . . . . . . . . . . . + . . + . . . . + . . . . . . . . . . . . . . o st eo sp er m um p in na tu m . . . . + . + + . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . + . . . . s p e c ie s g r o u p w c la do ra ph is s pi no sa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p x s tip ag ro st is o bt us a . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . + . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . z yg op hy llu m r et ro fr ac tu m . . + . + + . . . . . + . . . . . . . . . + . . . . . . . . + . . + . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 + . . . . . + . . . . . . . + . . . . . s tip ag ro st is b re vi fo lia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ta b l e 1 b (c o n t. ..) 174 original research van der merwe, van rooyen & van rooyen vol. 50 no. 1 pp. 160 183koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za r e l e v é n u m b e r 2 1 1 2 2 2 2 2 1 1 2 2 2 2 2 2 2 3 3 2 3 2 1 2 2 2 2 1 2 2 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 1 2 2 1 1 1 2 3 2 3 1 2 2 3 2 2 2 7 0 0 0 1 1 2 5 9 5 1 6 6 6 9 9 7 9 0 2 6 3 5 7 7 7 7 8 3 2 3 7 7 3 0 7 5 7 3 3 4 4 4 6 8 0 2 3 5 6 6 6 6 9 9 9 0 0 0 1 1 2 2 0 4 0 5 6 8 5 5 1 7 6 4 0 6 3 1 5 7 6 7 0 1 2 0 2 9 0 6 1 8 7 9 7 7 6 7 8 0 3 6 4 1 8 3 1 9 8 3 4 7 0 4 9 0 7 8 5 2 6 8 9 6 7 0 1 2 0 8 4 0 1 6 1 2 4 5 6 7 8 2 4 8 8 9 2 7 3 3 3 5 1 3 6 8 2 8 5 0 6 0 7 9 0 4 4 4 0 6 3 5 6. 1 6. 2 6. 3 6. 4 6. 5 b le ph ar is p ru in os a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . z yg op hy llu m ( h r p3 86 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p y m es em br ya nt he m ac ea e 1 (h r p3 59 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m es em br ya nt he m ac ea e 2 (h r p3 59 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p z p te ro ni a (h r p1 18 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ly ci um ( h r p1 18 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s al so la ( h r p1 18 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p te ro ni a (h r p1 18 ) sp . . . . . . . . . . . . . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * n on -d ia gn os tic s pe ci es a re e xc lu de d * h r c ol le ct io n co de a nd n um be rs a re in cl ud ed fo r fu tu re r ef er en ce , i f n ec es sa ry * s pe ci m en s in ad eq ua te fo r id en tifi ca tio n ye t d iff er en t f ro m s pe ci es th at c ou ld b e id en tifi ed a re in di ca te d w ith a p lo t n um be r (p … ) * sp . = o ne s pe ci es in a ta xo no m ic g ro up * sp p. = m or e th an o ne s pe ci es in a ta xo no m ic g ro up . t he se s pe ci es , e ve n al th ou gh th ey a re g ro up ed to ge th er , a re in cl ud ed in th e ta bl e si nc e th ey o cc ur in d iff er en t s pe ci es g ro up s. h ow ev er , t he y ar e no t u se d in th e de sc ri pt io ns in th e te xt . ta b l e 1 b (c o n t. ..) area of 103 665 ha (6.2% of the total area in fig. 1). it is also found near kalkgat in the north of the tanqua basin (kalkgat mosaic) and in the pan system in the tanqua basin (tanqua pan mosaics). it occurs predominantly on land types ia and fc. geologically, alluvial deposits dominate, while shales of the ecca group are present at times. limited intrusions of dolerite also occur. this low-lying (400–800 m above sea level) subassociation is found on level to gently sloping ridges and plains with no rock cover or a 50 to 98% rock cover comprising small stones. the sands are brown, light brown or red brown in colour. the shrub cover is intermediate to high (30–80%), the grassy component is absent, and annual cover is very low. the subassociation is characterised by a weak species group p with species such as cephalophyllum sp., pteronia villosa and pteronia glabrata (table 1c). other common species include malephora crassa (species group r), ruschia intricata (species group u) as well as aridaria noctiflora, salsola tuberculata, tripteris sinuata and pteronia pallens (species group v). subassociations 7.1 and 7.2 share gazania lichtensteinii of species group q. 7.3. malephora crassa – aridaria noctiflora tanqua karoo this subassociation is found predominantly at the southern extreme of the tanqua basin, i.e. ceres karoo, as well as on the western extreme of the hantam river where the river system is brackish (fig. 1). subassociation 7.3, excluding the mosaic vegetation units, covers an area of 176 425 ha (10.6% of the total area in fig. 1). it also occurs in the kalkgat mosaic southwest of calvinia and in the windheuwel/rooiheuwel mosaic between the roggeveld and koedoesberg mountains. shales of the ecca group and tillite of the dwyka group are found in these areas. land types da, ia and fc dominate the area that ranges in altitude from 200 to 1 000 m above sea level. these generally level ridges and plains have a low rock cover on light brown loamy soils. shrub cover averages 30 to 40%, while the cover of grasses and annuals is absent or very low. there is no diagnostic species group defining subassociation 7.3 (table 1c). prominent species include malephora crassa (species group r), atriplex lindleyi (species group t), ruschia intricata (species group u), aridaria noctiflora, salsola tuberculata, drosanthemum (hr217) sp. and pteronia pallens (species group v). the absence of species groups o, p and q distinguishes this subassociation from subassociations 7.1 and 7.2, but the presence of species group r shows affinity with them. a relationship with subassociation 7.4 is indicated by the shared presence of species group t (table 1c). 7.4. atriplex lindleyi – aridaria noctiflora loeriesfontein karoo subassociation 7.4 is located around the town of loeriesfontein and east thereof (fig. 1) and, excluding the mosaic vegetation unit, covers an area of 184 612 ha (11.1% of the total area in fig. 1). in the calvinia mosaic subassociation 7.4 (table 1c) is found in combination with subassociation 6.5 (table 1b). these areas are closely associated with what acocks (1953, 1988) termed false succulent karoo. the vegetation has a desert character and is sparsely populated with mesembs (vygies) and relics of the arid karoo. the degradation of the vegetation has been ascribed to the excessive grazing pressure and, consequently, the species that are of value for grazing are precisely the ones that no longer appear in it (acocks 1953, 1988). this subassociation occurs on the shales of the ecca group and a network of karoo dolerites, predominantly on land types ia, fc and da, at an altitude ranging from 400 to 1 000 m. these usually level ridges generally have a low rock cover on the brown to light brown coloured loamy soils. an intermediate shrub cover (40–60%), and grass and annual components that are absent or very low, describe this subassociation. species group s with species such as braunsia cf. apiculata and brownanthus cf. vaginatus characterise this subassociation (table 1c). atriplex lindleyi (species group t), 175 succulent karoo biome related vegetation – part 2 original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 160 183 ruschia intricata (species group u), as well as aridaria noctiflora, salsola tuberculata and tripteris sinuata (species group v) are also abundant. 7.5 ruschia intricata – aridaria noctiflora tanqua karoo this subassociation is located at the foothills of the koedoesberg mountains (fig. 1) and, excluding the mosaic vegetation unit, covers an area of 73 290 ha (4.4% of the total area in fig. 1). it is also found in the tanqua karoo inselberg mosaic south of calvinia. these inselbergs include the nuwewater se berg, elandsberg, eselberg, leeuberg, potkleiberg and sterretjieberg. the subassociation occurs predominantly on the skoorsteenberg formation of the ecca group in land types fc and ia. in the mosaics it co-occurs with subassociation 5.2 (table 1a), which occupies the dolerites of these inselbergs. the level to occasionally gently sloping ridges on light brown or brown coloured loams have either a low rock cover of < 5% or a high rock cover of 70 to 90%. an intermediate (40–45%) shrub cover is present in this subassociation and the grass and annual components are usually absent. there is no diagnostic species group defining this subassociation and species groups u and v are the only links with the other subassociations in association 7 (table 1c). ruschia intricata (species group u) and aridaria noctiflora (species group v) dominate the subassociation. 8. stipagrostis obtusa central tanqua grassy plains association 8 is located in the central tanqua basin (fig. 1) and covers an area of 239 781 ha (14.5% of the total area in fig. 1). it occurs from 200 to 1 000 m above sea level, predominantly on land type fc. tillite of the dwyka group and mudrock and shales of the ecca group comprise the geology of this area. the sandy plains and ridges generally have no rock cover or have a high rock cover in localised patches, with a 60 to 99% cover of gravel, small stones and/or stones. the sandy soils vary in colour from light brown to brown to red brown. shrub cover is less than 20%, while grass cover ranges from 10 to 90%. annuals are generally absent or have < 5% cover. two variations can be distinguished (table 1c). the first variation occurs at a low altitude (200–600 m) on deep sandy plains without any rock cover. the second variation occurs at an altitude of 300 to 1 000 m above sea level on level ridges and plains, and in washes with no rock cover or on ridges with a rock cover of 60 to 99%. the first variation of association 8 is dominated by cladoraphis spinosa (species group w) but stipagrostis obtusa, stipagrostis brevifolia and stipagrostis ciliata (species group x) may be present, while the second variation lacks cladoraphis spinosa (species group w) and is characterised by stipagrostis species of species group x (table 1c). 9. mesembryantheceae (hrp359) sp. ceres karoo vygieveld a single relevé was surveyed defining this plant association that has not been mapped separately but occurs in isolated patches within subassociation 7.3 in the southern part of the tanqua basin on shales of the ecca group and tillites of the dwyka group. this vygieveld (succulent veld) is found on level ridges with a 99% cover of small stones. these small black stones and brown chips are responsible for creating a unique microhabitat in which these succulents grow. the succulent and shrub cover is less than 15%, with no grass or annual species present. species group y characterises this association (table 1c). 10. pteronia (hrp118) sp. tanqua karoo brackish flats association 10 occurs in combination with association 7.2 in the tanqua pan mosaic and is found in the central region of the tanqua karoo on alluvial deposits. the alluvial soils are brackish and depending on the salinity of the soils the vegetation differs. in some areas the pan systems contain no vegetation. this lowlying association occurs on level valley floors or on the edges of the pans. shrub cover can be as high as 50% while grasses and annuals are not usually present. species characterising this association are represented in species group z (table 1c). the three major river systems in the region, namely the hantam, tankwa and doorn rivers, were not sampled, but were mapped (fig. 1). these drainage systems occur on land type ia. many tributaries originating to the northwest of the hantam mountain converge to form the hantam river, which only flows after heavy rains. patches of salsola tuberculata and salsola cf. aphylla dominate this drainage system. other species include pentzia incana, ruschia cradockensis, atriplex lindleyi, aridaria noctiflora and galenia africana. there are numerous flood-irrigated lands as a result of the more favourable water conditions along the drainage system. other forms of transformation evident are the presence of the invasive alien prosopis species and the naturalised species atriplex lindleyi. the succulent karoo vegetation (acocks 1953, 1988) of the tankwa and doorn rivers comprises species such as acacia karoo and malephora crassa, and is usually dominated by salsola tuberculata. transformation of the natural vegetation into floodirrigated lands next to these drainage systems has occurred in the past. few of these lands are still utilised for cropping and now lie barren with limited vegetation cover to combat erosion. the invader prosopis species is also a serious problem, and has taken over vast areas of these drainage systems. discussion the clear difference in species composition of the fynbos related vegetation units and the succulent karoo vegetation units was supported by the twinspan on the entire data set of 390 relevés. this difference subsequently led to the final two phytosociological tables and associated maps as discussed in this article and in van der merwe et al. (2008). however, since the hantam-tanqua-roggeveld is an area where three biomes meet, namely the fynbos, succulent karoo and nama karoo biomes (rutherford & westfall 1986), it is expected that some vegetation units are transitional between two biomes. for example, rosenia oppositifolia is a prominent mountain renosterveld (fynbos) related species (van der merwe et al. 2008), but it also occurs in the roggeveld karoo subassociations 6.2, 6.3 and 6.4 (species group m, table 1b). furthermore, nama karoo elements such as euryops multifidus are also found in these subassociations. while the relationship between the tanqua karoo grasslands and grasslands of the bushmanland, nama karoo biome, was not investigated, common species with a high cover and constancy are found in both areas. further studies of these affinities are necessary before the relationship between these two areas can be quantified. comparisons between previous maps published by acocks (1953, 1988), low and rebelo (1998) and mucina et al. (2005) show similarities and some interesting dissimilarities. since the scale of these maps and the current map, and the reasons for compiling the maps, all differ; only generalisations can be made when comparing the various maps. the area covered in the present paper basically includes two of acocks’s veld types (acocks 1953, 1988). associations 4 (escarpment karoo), 5 (hantam karoo) and 6 (roggeveld karoo) combined form part of the western mountain karoo (veld type 28), which is incorporated into low and rebelo’s upland succulent karoo (unit 56) (low & rebelo 1998), while associations 7 (tanqua and loeriesfontein karoo) and 8 (central tanqua grassy plains) are both included into acocks’s succulent karoo (veld type 31) (acocks 1953, 1988), which relates to low and rebelo’s lowland 176 original research van der merwe, van rooyen & van rooyen vol. 50 no. 1 pp. 160 183koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za 37 r e l e v é n u m b e r 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 2 2 3 3 1 2 3 3 1 1 2 2 2 3 1 1 1 2 2 2 2 3 2 2 2 1 2 2 2 2 2 3 2 3 1 2 3 3 3 2 3 1 1 3 2 2 3 1 1 1 1 1 2 2 3 1 3 3 3 1 9 9 9 0 0 0 1 2 4 6 9 9 9 0 0 2 2 6 7 7 7 7 6 6 7 7 1 4 6 8 4 1 6 6 7 4 4 5 5 8 5 5 5 2 5 9 7 1 1 3 8 6 6 5 8 8 0 2 3 3 8 2 2 4 8 0 2 3 4 4 5 9 5 7 6 5 7 7 3 1 4 5 2 1 7 3 9 5 1 1 1 1 2 3 4 8 7 8 8 5 1 2 4 7 7 5 8 7 0 3 3 5 3 6 4 6 3 4 9 3 5 0 2 6 8 4 7 8 7 7 0 4 0 3 4 8 8 9 9 2 6 3 3 6 8 9 1 9 4 9 2 9 2 6 9 4 7 9 3 2 6 8 4 7 2 5 1 6 5 1 4 5 9 0 6 0 1 5 6 7 7 2 4 2 1 3 8 5 7 2 4 5 6 1 9 0 6 1 5 7 8 8 7. 1 7. 2 7. 3 7. 4 7. 5 8 9 10 s p e c ie s g r o u p a z yg op hy llu m fo et id um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e up ho rb ia ( h r p3 5) s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s ta pe lia s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a sp ar ag us ( h r p3 78 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . d id el ta s pi no sa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r us ch ia ( h r p3 5) s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . v yg ie ( h r p3 78 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v yg ie ( h r p3 82 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p te ro ni a (h r p3 82 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v yg ie ( h r p3 8 4) s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p b ty le co do n w al lic hi i . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . c ra ss ul a al pe st ri s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . r hu s bu rc he lli i . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p hy llo bo lu s sp . . . . . . . . . . . . . . . . . . . . . . . . . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m on tin ia c ar yo ph yl la ce a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p c p te ro ni a gl au ca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . 1 . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p d e up ho rb ia d ec us sa ta 1 . . . . . . . . . . . . . . . . . . . . . . . . a . . + . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . s ar co ca ul on s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a nt im im a cf . g ra ni tic a (h r 24 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e up ho rb ia h am at a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p e a rc to tis a ca ul is . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b ab ia na s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . t he si um c f. hy st ri x . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . tr ac hy an dr a fa lc at a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h er m an ni a cu ne ifo lia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . la ch en al ia v io la ce a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . o th on na fi lic au lis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . la pe ir ou si a m on ta na . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . la ch en al ia s pp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ta b l e 1 c ph yt os oc io lo gi ca l t ab le o f t he s uc cu le nt k ar oo b io m e re la te d ve ge ta ti on o f t he h an ta m -t an qu ar og ge ve ld s ub re gi on – a ss oc ia ti on s 7, 8 , 9 a nd 1 0 177 succulent karoo biome related vegetation – part 2 original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 160 183 r e l e v é n u m b e r 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 2 2 3 3 1 2 3 3 1 1 2 2 2 3 1 1 1 2 2 2 2 3 2 2 2 1 2 2 2 2 2 3 2 3 1 2 3 3 3 2 3 1 1 3 2 2 3 1 1 1 1 1 2 2 3 1 3 3 3 1 9 9 9 0 0 0 1 2 4 6 9 9 9 0 0 2 2 6 7 7 7 7 6 6 7 7 1 4 6 8 4 1 6 6 7 4 4 5 5 8 5 5 5 2 5 9 7 1 1 3 8 6 6 5 8 8 0 2 3 3 8 2 2 4 8 0 2 3 4 4 5 9 5 7 6 5 7 7 3 1 4 5 2 1 7 3 9 5 1 1 1 1 2 3 4 8 7 8 8 5 1 2 4 7 7 5 8 7 0 3 3 5 3 6 4 6 3 4 9 3 5 0 2 6 8 4 7 8 7 7 0 4 0 3 4 8 8 9 9 2 6 3 3 6 8 9 1 9 4 9 2 9 2 6 9 4 7 9 3 2 6 8 4 7 2 5 1 6 5 1 4 5 9 0 6 0 1 5 6 7 7 2 4 2 1 3 8 5 7 2 4 5 6 1 9 0 6 1 5 7 8 8 7. 1 7. 2 7. 3 7. 4 7. 5 8 9 10 s p e c ie s g r o u p f d or ot he an th us s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . h el io ph ila c ol lin a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s en ec io c ak ile fo liu s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m er xm ue lle ra s tr ic ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m or ae a sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . z yg op hy llu m p yg m ae um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lo to no ni s hi rs ut a . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . o st eo sp er m um a ca nt ho sp er m um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p g b ul bi ne lla s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . d im or ph ot he ca s in ua ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e ro di um c ic ut ar iu m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p h e hr ha rt a ca ly ci na . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . c yp hi a di gi ta ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m ic ro lo m a sa gi tt at um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b er kh ey a fr ut ic os a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p i e ri oc ep ha lu s pu rp ur eu s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . te tr ag on ia m ic ro pt er a . + . + . 1 . + + . + + + + . . . . . . . + . . . . . . . . . . . . . + . . . . . . . . + + . . . . . . . . . + . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . s en ec io c ar da m in ifo liu s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . c ot ul a ba rb at a . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b ul bi ne s uc cu le nt a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p j le ip ol dt ia s ch ul tz ei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . g or te ri a di ff us a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e up ho rb ia m au ri ta ni ca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p k p hy llo bo lu s te nu ifl or us . . . . . . . . . . . . . . . . . . . . . . . . . . . + . + + . . + . . . . . . . . . . . . . r . + . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b ro m us p ec tin at us . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . + . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . o nc os ip ho n gr an di flo ru m . . . . . . . . + . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . + . + . . . . . + . . . . . . . . + . . . . e ri oc ep ha lu s na m aq ue ns is . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lo to no ni s sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p l p en ta sc hi st is p at ul a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r os en ia g la nd ul os a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . o th on na a ur ic ul ifo lia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . d ro s a nt he m um c f. la ti p et al um . . . . . . . . . . . . + . . . . . . . . . . . . . . + . . + . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h el ic hr ys um o bt us um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ta b l e 1 c (c o n t. ..) 178 original research van der merwe, van rooyen & van rooyen vol. 50 no. 1 pp. 160 183koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za r e l e v é n u m b e r 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 2 2 3 3 1 2 3 3 1 1 2 2 2 3 1 1 1 2 2 2 2 3 2 2 2 1 2 2 2 2 2 3 2 3 1 2 3 3 3 2 3 1 1 3 2 2 3 1 1 1 1 1 2 2 3 1 3 3 3 1 9 9 9 0 0 0 1 2 4 6 9 9 9 0 0 2 2 6 7 7 7 7 6 6 7 7 1 4 6 8 4 1 6 6 7 4 4 5 5 8 5 5 5 2 5 9 7 1 1 3 8 6 6 5 8 8 0 2 3 3 8 2 2 4 8 0 2 3 4 4 5 9 5 7 6 5 7 7 3 1 4 5 2 1 7 3 9 5 1 1 1 1 2 3 4 8 7 8 8 5 1 2 4 7 7 5 8 7 0 3 3 5 3 6 4 6 3 4 9 3 5 0 2 6 8 4 7 8 7 7 0 4 0 3 4 8 8 9 9 2 6 3 3 6 8 9 1 9 4 9 2 9 2 6 9 4 7 9 3 2 6 8 4 7 2 5 1 6 5 1 4 5 9 0 6 0 1 5 6 7 7 2 4 2 1 3 8 5 7 2 4 5 6 1 9 0 6 1 5 7 8 8 7. 1 7. 2 7. 3 7. 4 7. 5 8 9 10 s p e c ie s g r o u p m e ri oc ep ha lu s pa up er ri m us . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . a . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . g az an ia r ig id a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . + . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e ur yo ps m ul tifi du s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p en tz ia c f. sp ha er oc ep ha la . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . r os en ia o pp os iti fo lia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p n p en tz ia in ca na . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r us ch ia c ra do ck en si s . . . . . . . . . . . . . . . . a . . . . . . . . . . . . . . . . . . a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . p te ro ni a gl om er at a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . r . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m or ae a sp p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e ri oc ep ha lu s er ic oi de s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . le ys er a te ne lla . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . k ar ro oc hl oa s ch is m oi de s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . + . . . . . . . . . . . . . . . . . . a rc to th ec a ca le nd ul a . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . + . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a sp ar ag us c ap en si s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . + . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . h ir pi ci um a lie na tu m . . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . te tr ag on ia fr ut ic os a . . . . + . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . e ri oc ep ha lu s m ic ro ph yl lu s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . t he si um li ne at um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . c hr ys oc om a ci lia ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p o r us ch ia r ob us ta a 1 r . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m el ol ob iu m c an di ca ns . + . . + . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . tr ib ol iu m h is pi du m . + . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a ug ea c ap en si s . . + . . + . . + . + . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . e ur yo ps a nn uu s . . + + . + + . . + . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p p c ep ha lo ph yl lu m s p. . . . . . . + . . . . . . . . . . . . . . . a 1 . . + . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p te ro ni a vi llo sa + . . . . . + . . . . . . . . . . + . . . . 1 + . . . . r . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . f el ic ia ( h r p1 71 ) sp . . . . . . . . . . . . . . . . . . . + + . + . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . f el ic ia s p. . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . p te ro ni a gl ab ra ta . . . . . . . . . . . . . . . . . . . 1 . . . . . . . 1 . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p q g az an ia li ch te ns te in ii . + + 1 + 1 + + . + + + + 1 + + + + . + . . . . + + + + + . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p r d ro sa nt he m um ( h r 21 9) s p. + + . . + . + + . . . + . + . . . + . . . . . + . + . . . + . . . + . + + + . . + + + . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m al ep ho ra c ra ss a . . a . . . + . . . . . . . 1 . . . 1 . . . . . . . . + . . + + . 1 . . . + + 1 . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . g al en ia fr ut ic os a + . . . . + . . . . . . . . . . + . . . . . . . . . . . + . . . 1 . . + . . . . . + . . . . + . a . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . ta b l e 1 c (c o n t. ..) 179 succulent karoo biome related vegetation – part 2 original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 160 183 r e l e v é n u m b e r 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 2 2 3 3 1 2 3 3 1 1 2 2 2 3 1 1 1 2 2 2 2 3 2 2 2 1 2 2 2 2 2 3 2 3 1 2 3 3 3 2 3 1 1 3 2 2 3 1 1 1 1 1 2 2 3 1 3 3 3 1 9 9 9 0 0 0 1 2 4 6 9 9 9 0 0 2 2 6 7 7 7 7 6 6 7 7 1 4 6 8 4 1 6 6 7 4 4 5 5 8 5 5 5 2 5 9 7 1 1 3 8 6 6 5 8 8 0 2 3 3 8 2 2 4 8 0 2 3 4 4 5 9 5 7 6 5 7 7 3 1 4 5 2 1 7 3 9 5 1 1 1 1 2 3 4 8 7 8 8 5 1 2 4 7 7 5 8 7 0 3 3 5 3 6 4 6 3 4 9 3 5 0 2 6 8 4 7 8 7 7 0 4 0 3 4 8 8 9 9 2 6 3 3 6 8 9 1 9 4 9 2 9 2 6 9 4 7 9 3 2 6 8 4 7 2 5 1 6 5 1 4 5 9 0 6 0 1 5 6 7 7 2 4 2 1 3 8 5 7 2 4 5 6 1 9 0 6 1 5 7 8 8 7. 1 7. 2 7. 3 7. 4 7. 5 8 9 10 s p e c ie s g r o u p s b ra un si a cf . a pi cu la ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 + + . . . 1 . . . . + . . . . 1 + . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . b ro w na nt hu s cf . v ag in at us . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . 1 . . . + . + + . . + . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p te ro ni a sp . . . . . + . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . + . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cf . l am pr an th us o tz en ia nu m . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + 1 . . . . . . . . . . . + . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s al so la s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . 1 . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p en tz ia s pi ne sc en s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p t a tr ip le x lin dl ey i . . . . . . . . . . . . . . + . . . . . . . . . . . . + . . + . . . . + . . . . + . . + + . . 1 + + + . . + 1 + + + + . + . . . + + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p hy llo bo lu s sp . . . . . + . . . . . . . . + . + . . . . . . . . . . . . . . . . . . + + + . . . + . . + . . . . . . + . . + . . . . + . + + . + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p si lo ca ul on ( p9 1) s p. + . . . . . . . a . + . . . . . . . . . . . . . . . . . . + . . . + . + . . . . . . . . . + 1 + + . . . . . + . . . + . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s al so la a ph yl la . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . 1 a 1 . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . e xo m is m ic ro ph yl la . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . + . . . a . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p u r us ch ia in tr ic at a . . . . 1 . . . . 1 . + 1 . . . . . . 1 . . 1 + 1 1 b . 1 + . . . . . . . 1 1 . . + . . . . . . + + . . . . . . + a . a . . . . . . . . . . + a a + a 1 1 + . . . . . . + . . . . . . . . . . . . . . . . f el ic ia a us tr al is . + . + + . . + . . + + . + . . . . . . . . . . . . . . . . . . . . a . . . . + . . . + . + . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . e ri oc ep ha lu s de cu ss at us . . . . . . . . . + . . + . . . . . . . . . . . . . + . . . . . . . . . . . . + . . . . . . . . . . . . . . . . + . . + . . . . . . . + . . . . + . + . 1 + . . . + . . + . . . . . . . . . . . . . . . . f ov eo lin a di ch ot om a + + . . . . . + + . + . + + . . . + + . . . . + . . . . . . . . . . . . . . . + . . . + . . + + + . . . . + . + . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . z yg op hy llu m ( h r p6 5) s p. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . + . . + . + + + . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . s p e c ie s g r o u p v a ri da ri a no ct ifl or a . + . . + . + + 1 . . a . + + 1 . + + + + 1 . r . + . 1 . . + . + + + + + + + . . + 1 + + . + + . . . . . + . . + . + . + + + + . + . + + + . + . + + + + + . + 1 + . . . . . . . . . . . . . . . . . . . g al en ia s ar co ph yl la . . . . + . . . + . + . . . . + . . + . . . . . . . . + . . . . . . + . . . . . + . . . . + . . . . . a . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ly ci um s pp . . . . . . . . . . . . . . . . + r . . r . . . . . . . . . . . . . . . . . . . + + . + . + . . + + . + + . . . + . . + . . + + + + + + . . . r r . . + . . r . . . . . . + . . + . . . . . . . . . . . . . s al so la tu be rc ul at a . + . . + . . . + . . + . + + . . + . + + + r . . . . + . . + 1 + . + . + + . . . + . . 1 . + + + . + + . + . . 1 . 1 . . + + + . . . . 1 + . . . + . r . . . . r + . . . . . . . . . r . . . . . . . . . tr ip te ri s si nu at a + + . . . + . . . + . . + . . + + . . . . . . . . + . + + + + . . . + . . . + + . . . . . . . . . . . . . + + . . r . + . + . + + . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . d ro sa nt he m um ( h r 21 7) s p. . . . . . . . . . 1 . + + . . . . + + + a . . + . . . + . + . . . + 1 . . . . + 1 . + + + 1 . + . . . . . + + 1 . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . p te ro ni a pa lle ns . 1 r . . . . 1 . . . . . . . . . r . . . . . 1 + + . . . 4 . . . . + + 1 . + . . a + 1 . + . . . . . . . . . . . . . . . . . . . . . . . . + . r a . + . . . . . . . . . . . . . . . . . . . . . . . . . g eo ph yt ic s pp . . . . + + . + . . + . . + . . . . + . . . + . . . . . . . . + . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . 1 . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . o xa lis s pp . + + + . . + . . . . . . . . . . . + . . . . . . . . . . . . + . . . + . . . . . . . . + + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . g al en ia a fr ic an a . . . + . r . . + . . . + . . . . . . . + . . . . + r . . . . . . . . + . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . + . + . . + . . . . . . . . . . . . . . . . + m es em br ya nt he m um gu er ic hi an um . . . . . . . . + . + + . . . + . . . . + . . . . . . . . + . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . + . . . . . . . . . . . . . . . . . . . . . o st eo sp er m um p in na tu m . + . . . . . + . . 1 + . . . . . . . . . . . . r . . + . . + . . . + . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . s p e c ie s g r o u p w c la do ra ph is s pi no sa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b 1 a 1 . 1 . . . + . . . . . . . . . . . . . s p e c ie s g r o u p x s tip ag ro st is o bt us a . . . + . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a . . . . . . . . . . . . . . . . . . . + 1 + 1 . . . + 1 + 1 + a + 1 1 + . . . z yg op hy llu m r et ro fr ac tu m . . . + . + . . + + . . . . . + r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . + + . + . . . . . . . . . . + . . . . + . . . + . 1 . . . + . + r + 1 . . . . . . s tip ag ro st is b re vi fo lia . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . + . 3 . . . . . . . . + . . . . ta b l e 1 c (c o n t. ..) 180 original research van der merwe, van rooyen & van rooyen vol. 50 no. 1 pp. 160 183koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za r e l e v é n u m b e r 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 2 2 3 3 1 2 3 3 1 1 2 2 2 3 1 1 1 2 2 2 2 3 2 2 2 1 2 2 2 2 2 3 2 3 1 2 3 3 3 2 3 1 1 3 2 2 3 1 1 1 1 1 2 2 3 1 3 3 3 1 9 9 9 0 0 0 1 2 4 6 9 9 9 0 0 2 2 6 7 7 7 7 6 6 7 7 1 4 6 8 4 1 6 6 7 4 4 5 5 8 5 5 5 2 5 9 7 1 1 3 8 6 6 5 8 8 0 2 3 3 8 2 2 4 8 0 2 3 4 4 5 9 5 7 6 5 7 7 3 1 4 5 2 1 7 3 9 5 1 1 1 1 2 3 4 8 7 8 8 5 1 2 4 7 7 5 8 7 0 3 3 5 3 6 4 6 3 4 9 3 5 0 2 6 8 4 7 8 7 7 0 4 0 3 4 8 8 9 9 2 6 3 3 6 8 9 1 9 4 9 2 9 2 6 9 4 7 9 3 2 6 8 4 7 2 5 1 6 5 1 4 5 9 0 6 0 1 5 6 7 7 2 4 2 1 3 8 5 7 2 4 5 6 1 9 0 6 1 5 7 8 8 7. 1 7. 2 7. 3 7. 4 7. 5 8 9 10 s tip ag ro st is c ili at a . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . + r . a . b . . . . . . a + . . . . . . b le ph ar is p ru in os a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . + . . . . . . z yg op hy llu m ( h r p3 86 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . s p e c ie s g r o u p y m es em br ya nt he m ac ea e 1 (h r p3 59 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . m es em br ya nt he m ac ea e 2 (h r p3 59 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . s p e c ie s g r o u p z p te ro ni a (h r p1 18 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 ly ci um ( h r p1 18 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + s al so la ( h r p1 18 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + p te ro ni a (h r p1 18 ) sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + * n on -d ia gn os tic s pe ci es a re e xc lu de d * h r c ol le ct io n co de a nd n um be rs a re in cl ud ed fo r fu tu re r ef er en ce , i f n ec es sa ry * s pe ci m en s in ad eq ua te fo r id en tifi ca tio n ye t d iff er en t f ro m s pe ci es th at c ou ld b e id en tifi ed a re in di ca te d w ith a p lo t n um be r (p … ) * sp . = o ne s pe ci es in a ta xo no m ic g ro up * sp p. = m or e th an o ne s pe ci es in a ta xo no m ic g ro up . t he se s pe ci es , e ve n al th ou gh th ey a re g ro up ed to ge th er , a re in cl ud ed in th e ta bl e si nc e th ey o cc ur in d iff er en t s pe ci es g ro up s. h ow ev er , t he y ar e no t u se d in th e de sc ri pt io ns in th e te xt . ta b l e 1 c (c o n t. ..) succulent karoo (unit 57) (low & rebelo 1998). mucina et al. (2005) mapped approximately ten vegetation units within the study area. there is good agreement between association 4 (escarpment karoo) and the tanqua escarpment shrubland (skv 4) of mucina et al. (2005). in general, the delineation of the hantam karoo (association 5), roggeveld karoo (association 6) and tanqua karoo (association 7) in this study is more restricted than that of the hantam karoo (skt 2), roggeveld karoo (skt 3) and tanqua karoo (skv 5) of mucina et al. (2005). comparisons of the acocks (1953, 1988), low and rebelo (1998) and mucina et al. (2005) maps with the map presented in this article reveal that the maps of acocks (1953, 1988) and low and rebelo (1998) differ appreciably in the van rhynsdorp/doorn river region while the mucina et al. (2005) map is more closely related to the map presented in this paper. the differences in this region require additional study to determine the boundaries of each different vegetation type since the region is situated in the transition between the tanqua karoo and van rhynsdorp succulent karoo and the fynbos biome. in the sutherland area, the region is classified by acocks (1953, 1988) as western mountain karoo and by low and rebelo (1998) as upland succulent karoo, yet the present study found this vegetation to be more closely related to the mountain renosterveld vegetation of the fynbos biome, and is described by van der merwe et al. (2008) as association 1. also, north and east of sutherland, parts of the mucina et al. (2005) roggeveld karoo (skt3) were incorporated into the mountain renosterveld (association 1 in van der merwe et al. 2008). two major threats to the vegetation in the study area were identified by the farming community and the northern cape department of agriculture, and through personal observation during field surveys. the first threat is that of invasive alien species, especially prosopis species. prosopis glandulosa was introduced to the karoo to aid in fodder production for small stock in the drier months of the year when little natural vegetation is available (zimmermann 1991). the pods of these trees are high in protein and many farmers rely on them to carry their stock through the dry months. however, more than one species of prosopis was introduced and these have now hybridised. these hybrids, with their hybrid strength, have invaded large tracts of land and are now a serious threat in many areas. other alien invasive species include nerium oleander and nicotiana glauca, which are usually restricted to drainage lines. these species are a serious problem in some areas, although they are not as widespread as the prosopis species that threaten not only drainage lines but also the natural vegetation between the drainage lines. the second serious threat is that of less than ideal farming practices. due to a lack of infrastructure, especially fencing, optimal farm management is not implemented. the main reason for this is that farms in the region have a low income as a result of the unfavourable and harsh environmental conditions. additionally, the monetary value of the land is low and the cost of infrastructure so high that it is not financially viable for a farmer to invest too much in infrastructure as it will not be possible to recover these costs. there is definitely willingness amongst farmers for improved farm management and infrastructure development; however, their financial means usually do not allow it. although damage can happen fast, recovery in the karoo is very slow, because it depends upon unpredictable rainfall events (esler et al. 2006). apart from the transformation as a result of alien invasive vegetation, transformation of natural vegetation into floodirrigated lands next to the major drainage systems has taken place in the past. few of these lands, especially in the tanqua 181 succulent karoo biome related vegetation – part 2 original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 160 183 karoo, are still utilised for cropping; they now lie barren, with little vegetation cover to combat erosion, or the lands are infested by invasive species, particularly prosopis species. formal conservation within the hantam-tanqua-roggeveld subregion is limited. there are two local municipal reserves, the nieuwoudtville wildflower reserve (115 ha) and the akkerendam nature reserve (230 ha) that qualify as formally protected areas. however, most of the akkerendam nature reserve is located on the plateau of the hantam mountain within the mountain renosterveld, fynbos related, vegetation of association 3 (van der merwe et al. 2008). the nieuwoudtville wildflower reserve is situated in the nieuwoudtville mosaic consisting of vegetation units 2.1.1, 2.1.4, 2.2. (van der merwe et al. 2008) and 5.1 (fig.1), which are a combination of mountain renosterveld (fynbos related vegetation) and succulent karoo vegetation. the conservation status of the tanqua karoo has increased considerably with the expansion of the tankwa karoo national park over the last few years. this national park currently protects 92 495 ha of land; however large tracts thereof are highly degraded. a few private nature reserves and conservancies are also found in the tanqua karoo, but these are generally located at the southern end of the tanqua basin, closer to ceres. in conclusion, this project aimed to classify and describe the vegetation units in the hantam-tanqua-roggeveld subregion using species composition, environmental parameters and vegetation unit relationships to one another to map their geographical distribution. this map could serve as a basis to aid future planning and biodiversity conservation through sustainable land use practices to reduce the impact on the land. acknowledgements the authors would like to thank the critical ecosystem partnership fund (cepf) through the skep (succulent karoo ecosystem plan/program) initiative for funding the project. the critical ecosystem partnership fund is a joint initiative of conservation international, the global environmental facility, the government of japan, the macarthur foundation and the world bank. its fundamental goal is to ensure that civil society is engaged in biodiversity conservation. capenature, department of tourism, environment and conservation (northern cape) as well as sanparks are thanked for the necessary permits and permission to conduct this research. the council for geoscience is thanked for providing geological data of the study area. the assistance of hennie van den berg of iris international for compiling the vegetation map is gratefully acknowledged. references acocks, j.p.h. 1953. veld types of south africa. memoirs of the botanical survey of south africa, 28: 1–192. acocks, j.p.h. 1988. veld types of south africa. 3rd ed. memoirs of the botanical survey of south africa, 57: 1–146. agricultural research council. 1986a. land type map 3018 loeriesfontein. pretoria, institute for soil, climate and water. agricultural research council. 1986b. land type map 3220 sutherland. pretoria, institute for soil, climate and water. agricultural research council. 1995. land type map 3118 calvinia. pretoria, institute for soil, climate and water. agricultural research council. 1999a. land type map 3120 williston. pretoria, institute for soil, climate and water. agricultural research council. 1999b. land type map 3218 clanwilliam. pretoria, institute for soil, climate and water. agricultural research council. 2002. land type map 3319 worcester. pretoria, institute for soil, climate and water. agricultural research council. 2003. land type map 3320 ladismith. pretoria, institute for soil, climate and water. born, j., linder, h.p. and desmet, p. 2007. the greater cape floristic region. journal of biogeography, 34: 147–162. cepf, 2003. ecosystem profile: the succulent karoo hotspot, namibia and south africa. critical ecosystem partnership fund report. c o n s e r v a t i o n i n t e r n a t i o n a l . 2 0 0 6 . h t t p://w w w. biodiversityhotspots.org [accessed 20 february 2006]. council for geoscience. 1973. geological map 3218 clanwillliam. pretoria, council for geoscience. council for geoscience. 1983. geological map 3220 sutherland. pretoria, council for geoscience. council for geoscience. 1989. geological map 3120 williston. pretoria, council for geoscience. council for geoscience. 1991. geological map 3320 ladismith. pretoria, council for geoscience. council for geoscience. 1997. geological map 3319 worcester. pretoria, council for geoscience. council for geoscience. 2001. geological map 3118 calvinia. pretoria, council for geoscience. council for geoscience. 2008. electronic data supplied by the council for geoscience, silverton, pretoria. du plessis, h.m. 1987. land types of the maps 2816 alexander bay, 2818 warmbad, 2916 springbok, 2918 pofadder, 3017 garies, 3018 loeriesfontein. memoirs on the agricultural natural resources of south africa, 9: 1–538. esler, k.j., milton, s.j. & dean, w.r.j. 2006. karoo veld ecology and management. pretoria, briza publications. germishuizen, g. & meyer, n.l. (eds.). 2003. plants of southern africa: an annotated checklist. strelitzia 14. pretoria: national botanical institute. hennekens, s.m. & schaminee, j.h.j. 2001. turboveg, a comprehensive data base management system for vegetation data. journal of vegetation science, 12: 589–591. hill, m.o. 1979. twinspan – a fortran program for arranging multivariate data in an ordered two-way table by classification of the individuals and attributes. ithaca, ny, ecology & systematics: cornell university. hilton-taylor, c. 1994. western cape domain (succulent karoo). in davis, s.d., heywood, v.h. & hamilton, a.c. (eds.), centres of plant diversity, a guide and strategy for their conservation, volume 1, cambridge: iucn publications unit, pp. 201–203. low, a.b. & rebelo, a.g. 1998. vegetation of south africa, lesotho and swaziland. pretoria: department of environmental affairs and tourism. mucina, l., rutherford, m.c. & powrie, l.w. (eds). 2005. vegetation map of south africa, lesotho and swaziland, 1 : 1 000 000 scale sheet maps. pretoria, south african national biodiversity institute. nelder v.j., wilson, b.a., thompson, e.j. & dillewaard, h.a. 2005. methodology for survey and mapping of regional ecosystems and vegetation communities in queensland. version 3.1. updated september 2005. brisbane, queensland herbarium: environmental protection agency. rubidge, b.s. & hancox p.j. 1999. the karoo – a palaeontological wonderland. in: viljoen, m.j. & reimold w.u. an introduction to south africa’s geological and mining heritage. pretoria: published by the geological society of south africa and mintek. rubin, f. 1998. the physical environment and major plant communities of the tankwa karoo national park. koedoe, 41: 61–94. rutherford, m.c. & westfall, r.h. 1986. biomes of southern africa. an objective characterisation. memoirs of the botanical survey of south africa, 54: 1–98. south african national biodiversity institute. 2006. http:// www.sanbi.org/consfarm/ [accessed 2 october 2006]. sa weather bureau 1998. climate of south africa. climate statistics up to 1990. wb 42. pretoria, government printer. snijman, d. & perry, p. 1987. a floristic analysis of the nieuwoudtville wild flower reserve, north-western cape. south african journal of botany, 53: 445–454. 182 original research van der merwe, van rooyen & van rooyen vol. 50 no. 1 pp. 160 183koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za van der merwe, h., van rooyen, m.w. & van rooyen, n. 2008. vegetation of the hantam-tanqua-roggeveld subregion, south africa. part 1. fynbos biome related vegetation. koedoe, 50: 61–71. van wyk, a.e. & smith, g.f. 2001. regions of floristic endemism in southern africa: a review with emphasis on succulents. pretoria, umdaus press. werger, m.j.a. 1974. on concepts and techniques applied in the zürich-montpellier method of vegetation survey. bothalia, 11: 309–323. zimmermann, h.g. 1991. biological control of mesquite, prosopis spp. (fabaceae), in south africa. agriculture, ecosystems and environment, 37: 175–186. 183 article information authors: brian w. van wilgen1 navashni govender2 gregory g. forsyth1 tineke kraaij3 affiliations: 1centre for invasion biology, csir natural resources and the environment, stellenbosch, south africa 2scientific services, kruger national park, skukuza, south africa 3scientific services, garden route, south africa correspondence to: brian van wilgen email: bvwilgen@csir.co.za postal address: po box 320, stellenbosch 7599, south africa dates: received: 03 dec. 2009 accepted: 13 july 2010 published: 13 may 2011 how to cite this article: van wilgen, b.w., govender, n., forsyth, g.g. & kraaij, t., 2011, ‘towards adaptive fire management for biodiversity conservation: experience in south african national parks’, koedoe 53(2), art. #982, 9 pages. doi:10.4102/koedoe.v53i2.982 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) towards adaptive fire management for biodiversity conservation: experience in south african national parks in this essay... open access • abstract • introduction • achieving management goals through the use of fire    • the goals of management    • the use of fire to achieve ecosystem goals    • fire and the achievement of societal goals    • adaptive management and fire • study areas • adaptive fire management in the kruger national park    • early development of adaptive fire management    • gaining new understanding    • moving from passive to active adaptive management    • remaining issues • expanding adaptive fire management to other national parks    • table mountain national park    • bontebok national park    • assessment of progress • discussion    • making use of modelling    • linking fire to conservation outcomes • acknowledgements • references abstract (back to top) this paper reviews the experience gained in three south african national parks (kruger, table mountain and bontebok) with regard to the adaptive management of fire for the conservation of biodiversity. in the kruger national park, adaptive approaches have evolved over the past 15 years, beginning initially as a form of ‘informed trial and error’, but progressing towards active adaptive management in which landscape-scale, experimental burning treatments are being applied in order to learn. in the process, significant advances in understanding regarding the role and management of fire have been made. attempts have been made to transfer the approaches developed in kruger national park to the other two national parks. however, little progress has been made to date, both because of a failure to provide an agreed context for the introduction of adaptive approaches, and because (in the case of bontebok national park) too little time has passed to be able to make an assessment. fire management interventions, ultimately, will manifest themselves in terms of biodiversity outcomes, but definite links between fire interventions and biodiversity outcomes have yet to be made. conservation implications: significant challenges face the managers of fire-prone and fire adapted ecosystems, where the attainment of ecosystem goals may require approaches (like encouraging high-intensity fires at hot and dry times of the year) that threaten societal goals related to safety. in addition, approaches to fire management have focused on encouraging particular fire patterns in the absence of a sound understanding of their ecological outcomes. adaptive management offers a framework for addressing these issues, but will require higher levels of agreement, monitoring and assessment than have been the case to date. introduction (back to top) south african national parks (sanparks) have a primary mandate to conserve, on the land that they manage, south africa’s biodiversity, landscapes and associated heritage assets. in line with this, broad conservation goals normally call for the maintenance of all elements of biodiversity over space and time (sanparks 2008). managers of most national parks have used a range of interventions to achieve the goals of biodiversity conservation, but in recent years there has been a growing recognition that top-down, ‘command-and-control’ management can be both ineffective and damaging (holling & meffe 1996). this, in turn, has led to the introduction of adaptive forms of management in national parks. adaptive management was introduced into the kruger national park (knp) in the early 1990s (biggs & rogers 2003) and, as practised by sanparks, has the following key features (sanparks 2008): • the development of a shared vision and desired state for the future of the park • the formulation of a hierarchy of objectives, agreed to by all stakeholders, that will provide a defendable purpose and clear focus for management for achieving the desired state • the formulation of targets (or thresholds of potential concern), with the help of experts, to describe the boundaries of the desired state that management aims to achieve • the implementation of agreed management actions, and the monitoring of outcomes in terms of agreed targets or thresholds • in cases where targets are not met, or thresholds are exceeded, that consideration be given to management interventions that will drive the system towards targets or thresholds, or, alternatively, targets or thresholds can be recalibrated. the above may be regarded as ‘passive adaptive management’ (wilhere 2002), which involves the formulation of predictive models, making policy decisions based on these models, and revising the models or the policy as monitoring data become available. active adaptive management, on the other hand, deliberately applies different management approaches, as in an experiment with replication (walters & green 1997; wilhere 2002). proponents of active adaptive management argue that it accelerates learning, which is required if sustainability goals are to be met in any reasonable timeframe. for example, the resilience alliance states that ‘adaptive management seeks to aggressively use management intervention as a tool to strategically probe the functioning of an ecosystem’ (resilience alliance 2010), both to change the system and to learn about the system. many of south africa’s national parks are fire-prone, and fire is an important process to consider when managing these ecosystems for biodiversity (bond 1997). managers therefore often seek to influence the timing, frequency, size and intensity of fires, and the evolution of fire management has a long history in south african ecology (edwards 1984; phillips 1930; van wilgen 2009a, 2009b). the knp also has a long history of fire management (joubert 2007), including a move to adaptive management of fire in the late 1990s (van wilgen, biggs & potgieter 1998; van wilgen, govender & macfadyen 2008). more recently, adaptive approaches to fire management have been proposed for two additional national parks (table mountain national park [tmnp] and bontebok national park [bnp]), and there is potential to expand these approaches further to other fire-prone national parks. this paper reviews the development and use of adaptive approaches to manage fire in the knp, tmnp and bnp. it provides an assessment of the degree to which these approaches have impacted on the conservation of biodiversity, and discusses the ongoing challenges that will face managers when expanding these approaches to other national parks. achieving management goals through the use of fire (back to top) the goals of management for the purposes of this discussion on the use of fire, we have divided management goals in national parks into two broad categories. these are, (1) ecosystem goals, which include the conservation of biodiversity and landscapes and (2) societal goals, which include the impacts of fire on people, assets and infrastructure. in the case of national parks, societal goals include the protection of park assets and the safety of inhabitants and visitors, as well as infrastructure, crops and people outside of the park’s borders. in the sections that follow, we provide a broad, but brief, background to the use of fire as a management tool to achieve a range of goals. the use of fire to achieve ecosystem goals ecosystem managers often use fire as an intervention to influence vegetation structure and composition by manipulating the timing and frequency of fire. these interventions are normally based on an understanding of the ecology and responses of species and ecosystems to fire. examples of such interventions include:• the improvement of grazing for wildlife in grassland, savanna and fynbos ecosystems by burning to maintain the vigour and palatability of the grass sward (bond 1997; kraaij & novellie 2010) • the maintenance of populations of long-lived, obligate reseeding shrubs (mainly in the family proteaceae) in fynbos ecosystems. in this case, increases in fire frequency or shifts in fire season are detrimental, and managers seek to burn in appropriate seasons, and at appropriate intervals, to prevent population declines (van wilgen, bond & richardson 1992) • the prevention of increases in the density of woody plants in savannas (‘bush encroachment’), brought about by the combined effects of historic reductions in fire frequency, overgrazing, reductions in browsing (especially where elephant populations have been reduced) and increases in atmospheric carbon dioxide (bond, woodward & midgley 2003; kraaij & ward 2006; roques, o’connor & watkinson 2001). the usual approach here is to apply frequent fires of high intensity that would increase woody plant mortality and favour grasses • the integration of fire with efforts to control invasive alien plants, especially where these are spread by fire. pines and hakeas in fynbos, and australian wattles in most fire-prone ecosystems, are good examples (holmes et al. 2000; van wilgen et al. 1992). despite the widespread use of fire to achieve specific ecosystem objectives, simple yet robust goals that can cater for the protection and maintenance of all aspects of biodiversity are not easily formulated. for this reason, managers sometimes express goals in terms of desired fire regimes. fire regimes describe the patterns of fires in a particular ecosystem, and include the season, frequency, intensity, severity and size distribution of historical fires in the area concerned (gill & allan 2008). management goals can be expressed in terms of achieving target ranges of each of these elements, which are in turn delimited, based on an understanding of ecological responses to fire. because there is little or no understanding of responses to fire for most species, the assumption is made that, by promoting a variable fire regime within agreed limits, the survival of such species will be ensured because the variation will capture the conditions under which the ecosystem had evolved historically, and to which its component species would be pre-adapted (brockett, biggs & van wilgen 2001; van wilgen et al. 1998). fire and the achievement of societal goals managers of fire-prone protected areas, unless these areas are very large or sufficiently remote, have to contend with the effects of fires within a landscape that includes developed areas, crops or plantations, often immediately adjacent to the protected area. achieving the goals of safety and the protection of assets in such landscapes often requires the application, prevention, suppression and containment of fire in ways that may not be optimal for biodiversity conservation. as a result, there are often serious conflicts of interest in fire management. for example, unplanned fires that would otherwise be regarded as beneficial, and even necessary, for the maintenance of healthy ecosystems, may pose serious threats and have to be suppressed or contained for safety reasons. in addition, prescribed burning at the height of the fire season (often the best time to burn for ecological reasons), or at the high intensities needed for reversing or containing bush encroachment, often cannot be carried out because of restrictions imposed from outside national parks (for example, by fire protection associations). suppressing and containing unplanned fires is also expensive, as it involves the use of labour, vehicles, aircraft and other specialised equipment, and can deplete funds that may otherwise have been available for fire management to achieve conservation objectives. burning to reduce fuel loads, and the preparation of firebreaks, are two additional approaches used to achieve safety goals, although they are not universally successful (van wilgen et al. 2010). adaptive management and fire various approaches to managing fires have been adopted, or proposed, to achieve the goals set by sanparks and managers of other protected areas. these include regular or flexible prescribed burning (bands 1977; edwards 1984), ‘adaptive interference’ (van wilgen, richardson & seydack 1994), natural (lightning) fires (biggs & potgieter 1999; seydack, becker & marshall 2007), patch mosaic burning (brockett et al. 2001) and range condition burning (biggs & potgieter 1999). none of these approaches are perfect, and each has shortcomings. common problems include the following, (1) unplanned fires that disrupt prescribed burning plans, (2) suitable safe weather in which to burn is scarce, (3) complications arise as a result of the presence of invasive alien plants and (4) public resistance to deliberate burning. furthermore, the long-term ecological outcomes of each approach cannot always be predicted accurately. the degree to which managers can influence long-term fire regimes in reality, and whether or not a significant degree of control is actually necessary, are two issues that need clarification. for example, recent reviews have revealed that managers have not exerted a significant degree of control over fire reg imes, despite policies that promote prescribed burning with this aim in mind (van wilgen et al. 2004, 2010). adaptive fire management, as practised by or proposed for sanparks, has sought to address some of these issues. the approach has been to set targets in terms of elements of the fire regime, and to monitor whether the fire regimes that arise from various forms of management remain within the specified ranges. initially this was carried out as an interim measure, until better information became available (van wilgen et al. 1998). the resilience alliance does not recognise this type of management as adaptive management, but rather as ‘informed trial and error’, where the best available knowledge is used to generate a ‘risk-averse, best-guess’ strategy, which can be changed when new information allows for the initial strategy to be modified (resilience alliance 2010). the sections that follow provide a brief history of the evolution of adaptive management approaches in sanparks. study areas (back to top) this assessment focused on the knp, tmnp and bnp. although the need to manage fires is common to all three of these parks, the parks differ in many other respects (table 1). the parks differ in size by orders of magnitude, which impacts on the options available for fire management. the vegetation of the knp is a well-wooded savanna, in which fires occur regularly during the dry winter at return intervals of about 2−4 years. as a typical, semi-arid savanna, it displays wide levels of variability. the predominant vegetation of the tmnp comprises evergreen, mountain fynbos shrublands, with well-developed afromontane forests occurring in sheltered areas. these forests tend to be fire-free, although forest margins are scorched by fire from time to time. the fire regime of the area is characterised by dry-season (summer) fires at intervals of between 10 and 20 years. the bnp is covered by a mixture of fynbos and renosterveld shrublands. the fire regimes of these lowland areas are not well documented, but fire return intervals are thought to be shorter than those typical of mountain fynbos areas. importantly, a different set of issues exists in each area, which complicates or constrains fire management in different ways. table 1: salient features of three national parks to which adaptive approaches to fire management have been introduced. adaptive fire management in the kruger national park (back to top) early development of adaptive fire management fire management policies in the knp have changed several times over the past century, and have included fire protection and suppression, various forms of deliberate prescribed burning on fixed areas, the promotion of a lightning-driven fire regime, and point-ignition patch burning. these changes are well documented (biggs and potgieter 1999; joubert 2007; van wilgen et al. 2004, 2008, 2010; venter et al. 2008) and are not repeated in detail here. the development of fire management has also been informed by long-term and ongoing fire research (brynard 1971; van der schijff 1958; van wilgen, govender & biggs 2007). considerable changes in fire management came about, following the adoption of a heterogeneity paradigm in the early 1990s, with variability as a central concept (mentis & bailey 1990; rogers 2003). this new thinking coincided with the introduction of adaptive management in the knp (biggs & rogers 2003) and in 1998, led to a set of objectives relating to fire management, embedded in the hierarchy of objectives, and the formulation of fire-specific thresholds of potential concern (van wilgen et al. 1998). in this hierarchy, the knp’s overall goal of maintaining biodiversity in all its natural facets and fluxes would be supported by the development of an understanding of the role of fire as an ecosystem process. specific objectives included research to understand the effects of point versus perimeter-ignited burns, the effects of spatial heterogeneity in fires, the effects of fire on vegetation structure and on animals, and historic fire patterns. ultimately, the goal was to use this understanding to develop an informed context for fire management. the original fire-related thresholds (formulated in 1998) included addressing the cause of fires. this was included to ensure that most fires originated from lightning. while the cause of fires has little biological significance, it was argued that, in order to effectively promote a ‘natural’ (lightning-driven) fire regime, most fires would need to be ignited by lightning. as it turned out, most fires were not started by lightning. the management approach was then changed to one that combined point ignitions (‘patch fires’, brockett et al. 2001) with unplanned and lightning fires. the approach aimed to burn an annual target area, determined by rainfall and fuel conditions, in point-ignition fires of different sizes. the number of fire-related thresholds was reduced from seven to two (van wilgen et al. 2008). these were, (1) a threshold that would be reached if the area covered by fires in any one of three intensity classes (very low and low combined, moderate, and high and very high combined) constituted < 20% or > 50% of total area burnt in a given year and (2) a threshold that would be reached if a heterogeneity score (which measured the spatial heterogeneity of fire patterns) dropped below a predetermined value for two consecutive years. gaining new understanding the process of formulating, evaluating and reformulating thresholds for fire management has resulted in several advances in our understanding. as a result, managers now have a more informed context for fire management. advances in understanding include:• the demonstration that the area burnt in any given year is driven by variation in annual rainfall, and is largely unaffected by attempts to impose an annual burning target through prescribed burning (van wilgen et al. 2004) • the further demonstration that, although fire management does not influence the total area burnt, it is able to influence both the spatial heterogeneity and seasonal distribution of areas burnt through prescribed burning (van wilgen et al. 2004) • the conclusion, from burning on experimental plots, that the knp’s savannas are remarkably resilient under a wide range of fire treatments, and that fire has few negative effects unless the fire regimes deviate markedly from the norm (these deviations included burning every year, burning in the wet summer season, or long periods of fire exclusion (van wilgen et al. 2007) • the demonstration of the importance of fire intensity (as opposed to season or return period) for determining the survival rate of wood plants after fire, and the balance between grasses and trees (higgins, bond & trollope 2000; trollope 1998). moving from passive to active adaptive management in response to this understanding, the focus of fire management has shifted from attempts to control return periods and seasons to influencing spatial heterogeneity in fire intensity. new thresholds have been formulated, and a threshold for area burnt in high-intensity fires (as opposed to moderate or low-intensity fires) has been exceeded more than once, suggesting a need to find a way to reduce the proportion of high-intensity fires. a landscape-scale experiment has been initiated to establish the effects of ignition patterns (point ignitions or perimeter ignitions) on the proportion of area that burns at different intensities. the hypothesis is that point ignitions will generate a greater diversity of fire intensities within a single management fire, as it is thought that most of the area burnt in a perimeter-ignited fire burns as a high-intensity headfire. the contrasting of perimeter and point ignitions is of particular interest to managers, who favour the perimeter ignitions, as they allow for more control of fire. in order to persuade managers to replace a perimeter-ignition system with one of point ignitions, it will be necessary for researchers to demonstrate that, (1) different ignition patterns will result in differences in patterns of fire intensity and (2) these differences in fire intensity are meaningful in terms of biodiversity outcomes. this short history suggests that the use of fire initially followed the ‘informed trial and error’ approach, but that, with the initiation of the landscape-scale experiment, it has entered a phase of active adaptive management in which large-scale treatments are applied across the landscape in order to learn. remaining issues despite many advances in the understanding of the role and management of fire and a recognition that fire, as an ecological driver of vegetation composition and structure, is perhaps not as important as was once thought (van wilgen & biggs 2010), the need for ongoing development of further understanding is frequently expressed. it is recognised as simplistic to treat the entire knp (approximately 2 million ha) as homogenous, and to apply a single set of fire-related thresholds over the entire area. mean annual rainfall varies from between approximately 350 mm in the north and approximately 750 mm in the south, and the effects of fire are far more marked in areas of higher rainfall (van wilgen et al. 2007). in addition, important differences in responses to fire are apparent on the two major geological substrates (granite and basalt), suggesting that at least four sets of targets or thresholds (low and high rainfall on granite and basalt, respectively) would be more appropriate, but these have yet to be developed. the interactions between elephants and fire are also not clearly understood, but are important. concern is often expressed about declining numbers of large trees in the knp, which is partly due to interactions between fire and herbivory by elephants (eckhardt, van wilgen & biggs 2000; edkins et al. 2007). where, whether, and how managers should intervene in this regard remains unclear. furthermore, rising levels of carbon dioxide in the atmosphere will favour trees over grasses (bond et al. 2003). managers may wish to counter the expected increases in woody plant density if losses of biodiversity are to be avoided. this, in turn, may require the application of high-intensity and possibly, dangerous, fires to control woody plant growth (bond & archibald 2003), and is likely to present significant challenges to managers. expanding adaptive fire management to other national parks (back to top) table mountain national park in 2000 a fire management plan was prepared, on contract, for the tmnp (forsyth et al. 2000), and revised in 2004 (forsyth & bridgett 2004) by the council for scientific and industrial research (csir). then, in 2007 the park requested the csir to formulate a set of fire-associated thresholds for assessing the condition of the park’s fire-dependant ecosystems (forsyth, van wilgen & schonegevel 2007). a set of preliminary thresholds had been developed by van wilgen & scott (2001), and these were used as a basis for the development of operational and ecosystem thresholds for the tmnp (table 2). table 2: proposed thresholds of potential concern relating to fire management in the table mountain national park. an analysis of recent fire records revealed that the threshold regarding the area of fynbos (36.4%) in the age class of 4–6 years was 16.4% greater than the desired 20% upper threshold (forsyth et al. 2007; forsyth & van wilgen 2008), mainly because of a large unplanned fire in january 2000. slower-growing serotinous proteaceae in the area were therefore vulnerable to subsequent unplanned fires, should they occur. as a result of the threshold being exceeded, it was proposed that the vegetation should not be burnt for at least a further 4 years, and that the management emphasis in this area should be on protecting the vegetation from fire and ensuring a rapid response to limit the extent thereof in the event of an unplanned fire. fire size distributions were also found to have exceeded thresholds. large fires accounted for only approximately 45% of the area of all fires. in addition, a few large fires in january 2000 exceeded the upper threshold of 2000 ha. the remaining (ecosystem) thresholds in table 2 have not yet been assessed, mainly because monitoring programmes to gather the necessary data are not yet in place. bontebok national park the bnp is a small protected area (table 1), originally proclaimed to protect the rare bontebok (damaliscus pygargus pygargus). vegetation types within the park are also rare and threatened (kraaij 2011), and the park’s managers have long been faced with the dilemma of reconciling the need for short-interval fires that promote grazing for bontebok, with that for longer-interval fires to maintain plant diversity (kraaij 2010). in the 1970s a short-interval burning programme was introduced to stimulate grazing for bontebok. under this programme renosterveld and fynbos vegetation were burnt at intervals of 4 and 12 years, respectively. in response to the increasing need to conserve the vegetation, prescribed burning schedules were changed in 2004, and the interval between fires was extended to not less than 8 years in renosterveld vegetation and 16 years in fynbos. as a result, mean fire return periods increased from 6.7 to 10.9 years (kraaij 2010). in order to monitor and improve fire management in the bnp a preliminary set of thresholds for fire (in terms of frequency, season and size distribution) has been developed, in line with those used in the tmnp (table 3). some of the thresholds used in the bnp were adopted from the tmnp set, and would apply, for example, to surviving populations of obligate reseeding protea repens shrubs. furthermore, thresholds were designed to address the effects of interacting fire and grazing (kraaij & novellie 2010; novellie & kraaij 2010) on the vegetation. table 3: proposed thresholds of potential concern relating to fire management and fire-herbivory interactions in the bontebok national park. assessment of progress despite the attempted introduction of adaptive approaches to fire management, our understanding has not yet been noticeably advanced, at least in the tmnp. in the case of the bnp, it is too early to expect that progress should have been made. in reality, the attempt to introduce an adaptive approach to fire management in the tmnp has suffered from a number of shortcomings, including the following:• a holistic adaptive management approach was not followed in these cases, in that that there was no formulation of a desired state and a hierarchy of objectives, agreed to by all stakeholders, that would have provided the purpose and focus of management • the purpose and use of thresholds were not universally well understood by the tmnp’s managers • several important problems relating to fire management in the tmnp were not addressed in the formulation of operational and ecosystem thresholds. these problems relate to how to deal with a critical public (including gaining consensus within a fire protection association whose members did not share biodiversity goals), how to use scarce resources to ensure that both safety and ecosystem goals are met, and how to deal effectively with invasive alien plants. because of these shortcomings, there has been little real buy-in regarding adaptive management approaches, very little monitoring, and almost no assessment. in the case of the bnp, the thresholds presented here are recent (developed in 2008) and have not yet been assessed. it would appear, therefore, that the well-intentioned attempt to introduce an element of adaptive management, addressing fire alone, has not yet delivered any tangible benefits. in order to make progress, attention to the development of a shared vision and desired future state, and the formulation of a hierarchy of objectives, agreed to by all stakeholders, will be required. this would provide a defendable purpose and clear focus for management for achieving the desired state. should this be done, fire management issues will be more appropriately embedded within an agreed hierarchy of objectives, and should lead to improvements in fire management over time. discussion (back to top) making use of modelling the operational thresholds proposed for tmnp and bnp in terms of fire return intervals, season and size, are first approximations, based on current understanding of what constitutes a fire regime under which biodiversity can be conserved. however, whether, under what circumstances, or how often these thresholds are likely to be exceeded is not clearly understood. issues of scale are also important here. in the knp, the area is probably large enough for fire patterns to remain within thresholds on average. however, a single fire could cover the whole of the bnp, resulting in most of the thresholds being exceeded. the appropriate management responses to such scenarios have not yet been thought through. the modern and historic fire regimes in fynbos and savanna ecosystems are well understood (forsyth & van wilgen 2008; seydack et al. 2007; van wilgen et al. 2000; van wilgen et al. 2010), and a range of plausible scenarios could be formulated and modelled relatively easily. in this way, the frequency with which thresholds are likely to be exceeded in individual protected areas could be rapidly generated. this could lead either to the formulation of appropriate responses in advance, or to the delimitation of more appropriate thresholds. either way, this approach would lead to more rapid learning, and should be considered. linking fire to conservation outcomes whether or not adaptive fire management (as practised by sanparks) is having the desired effect of conserving biodiversity is an important question. the adoption of thresholds in terms of fire patterns may appear at first glance to imply that the fire patterns themselves are the desired outcome. in the knp, the overall goal for ecosystem management in the objectives hierarchy is ‘to understand and manage the knp as part of the lowveld savanna … in such a manner as to conserve and restore its varied natural structure, function and composition over time and space … through an approach integrating the different scales and types of objectives’. this goal, ultimately, was sub-divided into 369 sub-objectives (13 for atmosphere, 116 for water, 204 for terrestrial, 30 for alien species, and 6 for threatened biota, respectively), only 6 of which relate to fire. the outcomes of fire management interventions, ultimately, will manifest themselves in terms of vegetation structure, function and composition, which are monitored in terms of non-fire thresholds set to address the other sub-objectives. it has not yet been possible to make any definite links between fire interventions and conservation (biodiversity) outcomes in the knp. in a critical assessment of adaptive management in the knp, van wilgen and biggs (2010) concluded that ‘because the conservation outcomes of management shifts will only manifest themselves in the longer term, the relative success of adaptive management should be measured by the degree to which management has been refocused onto priority issues, and by the rate at which new understanding is generated’. because of the difficulty of making links between fire interventions and biodiversity outcomes, the thresholds set for fire regimes are seen by managers as operational thresholds; that is, they guide managers towards targets that can be re latively easily understood. although the difference between operational and ecosystem thresholds was never framed as such in the knp, the distinction has been made explicit in the thresholds proposed for the tmnp and the bnp (tables 2 and 3). ultimately, however, the links need to be made between fire patterns and the ecological outcomes as embodied in the ecosystem thresholds, and this remains a major future challenge. acknowledgements (back to top) we thank the csir, the national research foundation, south african national parks and the dst/nrf centre for invasion biology for supporting the production of this paper. angela gaylard and philip prins commented on an earlier version of this paper. this paper was extensively revised following constructive criticism from two anonymous reviewers. references (back to top) bands, d.p., 1977, ‘prescribed burning in cape fynbos’ in h.a. mooney & c.e. conrad (eds.), proceedings of the symposium on the environmental consequences of fire and fuel management in mediterranean ecosystems, palo alto, california, 01–5 august, 1977, pp. 245–256, general technical report wo-3, usda forest service. biggs, h.c. & potgieter, a.l.f., 1999, ‘overview of the fire management policy of the kruger national park’, koedoe 42, 101–110. biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring and management in practice’ in j.t. du toit, k.h. rogers, h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59–80, island press, new york. bond, w.j., 1997, ‘fire’, in r.m. cowling, d.m. richardson & s.m pierce (eds.), vegetation of southern africa, pp. 421–446, cambridge university press, cambridge. bond, w.j. & archibald, s., 2003, ‘confronting complexity: fire policy choices in south african savanna parks’, international journal of wildland fire 12, 381–389. doi:10.1071/wf03024 bond, w.j., woodward, f.i. & midgley, g.f., 2003, ‘the importance of low atmospheric co2 and fire in promoting the spread of grasslands and savannas’, global change biology 9, 973–982. doi:10.1046/j.1365-2486.2003.00577.x brockett, b.h., biggs, h.c. & van wilgen, b.w., 2001, ‘a patch mosaic burning system for conservation areas in southern africa’, international journal of wildland fire 10, 169–183. doi:10.1071/wf01024 brynard, a.m., 1971, ‘controlled burning in the kruger national park: history and development of the range burning policy’, proceedings: tall timbers fire ecology conference, tallahassee, florida, 10–13 april, 1971, 219−231. eckhardt, h.c., van wilgen, b.w. & biggs, h.c., 2000, ‘trends in woody vegetation cover in the kruger national park, south africa, between 1940 and 1998’, african journal of ecology 38, 108–115. doi:10.1046/j.1365-2028.2000.00217.x edkins, m.t., kruger, l.m., harris, k. & midgley, j.j., 2007, ‘baobabs and elephants in kruger national park: nowhere to hide’, african journal of ecology 38, 108–115. edwards, p.j., 1984, ‘the use of fire and a management tool’, in p. de v. booysen & n.m. tainton (eds.), ecological effects of fire in south african ecosystem, pp. 349–362, springer, berlin. forsyth, g.g. & bridgett, j., 2004, table mountain national park fire management plan, report env-s-c-2004-043, csir, stellenbosch. forsyth, g.g. & van wilgen, b.w., 2008, ‘the recent fire history of the table mountain national park, and implications for fire management’, koedoe 50, 3−9. forsyth, g.g., van wilgen, b.w., ruddock, g., nel, j.l., le maitre, d.c., smith, n., et al., 2000, a fire management plan for the cape peninsula national park, report env-s-c 2000-111, csir, stellenbosch. forsyth, g.g., van wilgen, b.w. & schonegevel, l.y., 2007, an assessment of the current fire situation, proposed fire monitoring and recommendations for future fire management activities in the table mountain national park, report csir/nre/eco/er/2007/0056/c, csir, stellenbosch. gill, a.m. & allan, g., 2008, ‘large fires, fire effects and the fire regime concept’, international journal of wildland fire 17, 688–695. doi:10.1071/wf07145 higgins, s.i., bond, w.j. & trollope, w.s.w., 2000, ‘fire, resprouting and variability: a recipe for grass-tree coexistence in savanna’, journal of ecology 88, 213–229. doi:10.1046/j.1365-2745.2000.00435.x holling, c.s. & meffe, g.k., 1996, ‘command and control and the pathology of natural resource management’, conservation biology 10, 328–337. doi:10.1046/j.1523-1739.1996.10020328.x holmes, p.m., richardson, d.m., van wilgen, b.w. & gelderblom, c., 2000, ‘the recovery of south african fynbos vegetation following alien tree clearing and fire: implications for restoration’, austral ecology 25, 631–639. doi:10.1046/j.1442-9993.2000.01063.x, doi:10.1111/j.1442-9993.2000.tb00069.x joubert, s., 2007, the kruger national park: a history, 3 vols., high branching, johannesburg. kraaij, t., 2010, ‘changing the fire management regime in the renosterveld and lowland fynbos of the bontebok national park’, south african journal of botany 76, 550−557. doi:10.1016/j.sajb.2010.04.008 kraaij, t., 2011, ‘the flora of the bontebok national park in regional perspective’, south african journal of botany. doi:10.1016/j.sajb.2010.09.013 kraaij, t. & novellie, p.a., 2010, ‘habitat selection by large herbivores in relation to fire at the bontebok national park (1974–2009): the effects of management changes’, african journal of range and forage science 27(1), 21−27.doi:10.2989/10220111003703450 kraaij, t. & ward, d., 2006, ‘effects of rain, nitrogen, fire and grazing on tree recruitment and early survival in bush-encroached savanna, south africa’, plant ecology 186, 235–246. doi:10.1007/s11258-006-9125-4 mentis, m.t. & bailey, a.w., 1990, ‘changing perceptions of fire management in savanna parks’, journal of the grassland society of southern africa 7, 81–85. novellie, p. & kraaij, t., 2010, ‘evaluation of themeda triandra as an indicator for monitoring the effects of grazing and fire in the bontebok national park’, koedoe 52(1), 49-53. phillips, j.f.v., 1930, ‘fire: its influence on biotic communities and physical factors in south and east africa’, south african journal of science 27, 352–367. resilience alliance, 2010, adaptive management, viewed 03 june 2009, from http://www.resalliance.org/600.php rogers, k.h., 2003, ‘adopting a heterogeneity paradigm: implications for management of protected savannas’ in j. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 468−488, island press, new york. roques, k.g., o’connor, t.g. & watkinson, a.r., 2001, ‘dynamics of shrub encroachment in an african savanna: relative influences of fire, herbivory, rainfall and density dependence’, journal of applied ecology 38, 268–280.doi:10.1046/j.1365-2664.2001.00567.x sanparks, 2008, a framework for developing and implementing management plans for south african national parks, conservation services division, south african national parks, pretoria, 24 pp. seydack, a.h.w., bekker, s.j. & marshall, a.h., 2007, ‘shrubland fire regime scenarios in the swartberg mountain range, south africa: implications for fire management’, international journal of wildland fire 16, 81–95. doi:10.1071/wf06015 trollope, w.s.w., 1998, effect and use of fire in the savanna areas of southern africa, department of livestock and pasture science, university of fort hare. van der schijff, h.p., 1958, ‘inleidende verslag oor veldbrandnavorsing in die nationale krugerwildtuin [preliminary report on wildfire research in the kruger national park]’, koedoe 1, 60–93. van wilgen, b.w., 2009a, ‘the evolution of fire and invasive alien plant management practices in fynbos’, south african journal of science 105, 335−342. van wilgen, b.w., 2009b, ‘the evolution of fire management practices in savanna protected areas in south africa’, south african journal of science 105, 343−349. van wilgen, b.w. & biggs, h.c., 2010, ‘a critical assessment of adaptive ecosystem management in a large savanna protected area in south africa’, biological conservation. doi:10.1016/j.biocon.2010.05.006 van wilgen, b.w., biggs, h.c., o’regan, s. & mare, n., 2000, ‘a fire history of the savanna ecosystems in the kruger national park, south africa between 1941 and 1996’, south african journal of science 96, 167–178. van wilgen, b.w., biggs, h.c. & potgieter, a.l.f., 1998, ‘fire management and research in the kruger national park, with suggestions on the detection of thresholds of potential concern’, koedoe 41, 69−87. van wilgen, b.w., bond, w.j. & richardson, d.m., 1992, ‘ecosystem management’, in r.m. cowling (ed.), the ecology of fynbos: nutrients, fire and diversity, pp. 345−371, oxford university press, cape town. van wilgen, b.w., forsyth, g.g., de klerk, h., das, s., khuluse, s. & schmitz, p., 2010, ‘fire management in mediterranean-climate shrublands: a case study from the cape fynbos, south africa’, journal of applied ecology 47, 631–638. doi:10.1111/j.1365-2664.2010.01800.x van wilgen, b.w., govender, n. & biggs, h.c., 2007, ‘the contribution of fire research to fire management: a critical review of a long-term experiment in the kruger national park, south africa’, international journal of wildland fire 16, 519–530. doi:10.1071/wf06115 van wilgen, b.w., govender, n., biggs, h.c., ntsala, d. & funda, x.n., 2004, ‘response of savanna fire regimes to changing fire management policies in a large african national park’, conservation biology 18, 1533−1540. doi:10.1111/j.1523-1739.2004.00362.x van wilgen, b.w., govender, n. & macfadyen, s., 2008, ‘an assessment of the implementation and outcomes of recent changes to the fire management of the kruger national park’, koedoe 50, 22–31. van wilgen, b.w., richardson, d.m. & seydack, a., 1994, ‘managing fynbos for biodiversity: constraints and options in a fire-prone environment’, south african journal of science 90, 322−329. van wilgen, b.w. & scott, d.f., 2001, ‘managing fires on the cape peninsula: dealing with the inevitable’, journal of mediterranean ecology 2, 197−208. venter, f.j., naiman, r.j., biggs, h.c. & pienaar, d.j., 2008, ‘the evolution of conservation management philosophy: science, environmental change and social adjustments in kruger national park’, ecosystems 11, 173–192. doi:10.1007/s10021-007-9116-x walters, c.j. & green, r., 1997, ‘valuation of experimental management options for ecological systems’, journal of wildlife management 61, 987–1006. doi:10.2307/3802096 wilhere, g.f., 2002, ‘adaptive management in habitat conservation plans’, conservation biology 16, 20–29. doi:10.1046/j.1523-1739.2002.00350.x article information authors: robert j. scholes1 judith m. kruger2 affiliations: 1natural resources and environment, csir, pretoria, south africa 2south african national parks, skukuza, south africa correspondence to: judith kruger email: judith.kruger@sanparks.org postal address: private bag x402, skukuza 1350, south africa dates: received: 20 jan. 2010 accepted: 19 may 2010 published: 13 may 2011 how to cite this article: scholes, r.j. & kruger, j.m., 2011, ‘a framework for deriving and triggering thresholds for management intervention in uncertain, varying and time-lagged systems’, koedoe 53 (2), art. #987, 8 pages. doi:10.4102/koedoe.v53i2.987 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) a framework for deriving and triggering thresholds for management intervention in uncertain, varying and time-lagged systems in this essay... open access • abstract • introduction • a general approach to the problem    • define a sensitive, relevant and measurable indicator of the issue at hand    • set a limit of acceptable change in the indicator    • add a safety margin based on societal risk tolerance and technical certainty    • project the indicator into the future using a model, including an uncertainty range    • note the time at which the relevant uncertainty limit intercepts the safety margin    • subtract the system response time    • subtract the management response time    • subtract the monitoring interval       • is an action triggered? • example: elephants and big trees in the kruger national park    • choosing the indicator    • setting a safety-buffered limit    • projecting the dynamics forward    • when is the limit likely to be transgressed?    • subtracting the response times    • is an action triggered? • discussion and conclusion • acknowledgements • references abstract (back to top) ecosystems are characterised by complexity: high connectivity, the presence of positive and negative feedback loops, non-linear, abrupt and sometimes irreversible changes, delays between cause and effects, and uncertainties in observations, understanding and prediction. ‘adaptive management’ is the preferred approach for the rational management of such systems. where the management objective is to allow natural feedbacks and adaptive processes to operate as much as possible – as it is in many areas set aside for biodiversity conservation – a key issue is defining the thresholds that will trigger management intervention. this paper outlines and illustrates a logical process for doing so, taking into account the characteristics of complex, continuously changing ecosystems and the reality of information that is partial and understanding that is always provisional. after identifying a key ecological process that is believed to have an element of irreversibility beyond a certain point, the process has several steps, (1) define an indicator of the system state, (2) set a limit of acceptable change and add a safety margin, (3) project the indicator forward using a model, including uncertainty, (4) note the time when the indicator might transgress the safety-buffered limit and (5) subtract ecosystem and management response times. if the resultant time is at hand, an action is indicated – if not, the action is to continue to monitor the situation and refine the observations and models. conservation implications: ecosystems are characterized by abrupt and sometimes irreversible changes. the challenge that face conservationists and managers are to identify which of these changes are likely to be irreversible and at what levels this will occur. this paper describes a logical process that enable mangers to determine which ecological processes have levels of irreversibility and monitor their status at all times. once these processes are nearing the levels that are undesirable management actions can be invoked to prevent this from happening. introduction (back to top) the formal and informal management philosophy that has guided ecosystem management in state-owned, protected areas in south africa has evolved through various phases over the past century (carruthers 1995). the approach adapted in the early years (up to the mid-1950s) has been characterised as ‘command and control’. the first conservation officers were military men – decisive, action-orientated and used to being in charge. they assumed that the desired state was known, unchanging and achievable using the tools at hand. the desired state usually amounted to ‘preservation’: a perpetuation of the status quo or a return to some pre-existing, eden-like ideal. the underlying world view was the notion that nature is intrinsically in balance, but perturbed by human activities. therefore, self-regulation can be achieved by simply eliminating the human impacts, or, where this is impossible, by intervening to restore the balance. this first generation of conservators was replaced by a generation of technocrats, many trained as biological scientists, and backed up by a powerful, but not very consultative state apparatus (mid-1950s to about 1990). they were increasingly able to implement ‘grand schemes’ such as fencing, water provision, game translocation, culling and fire control. nevertheless, the ecological outcomes were often not what was anticipated or desired. these experiences, which were not unique to south africa, eventually taught a degree of humility regarding both our state of relative ignorance towards ecosystem functioning and in relation to how ineffective our interventions can be (holling & meffe 1996). the past three decades have seen a fundamental shift, worldwide, in the scientific perspective regarding ecosystem dynamics (a key paper in the african savanna context is ellis and swift 1988). it is now generally accepted that ‘natural’ ecosystems may be intrinsically variable due to their internal dynamics and as a result of the variability of external drivers, such as climate. thus, although the ecosystem may have a theoretical point of balance or equilibrium, it is seldom actually at that point, and it is neither desirable nor possible in the long term to suppress the variability. it is now thought to be necessary for ecosystems to vary in time and space in order to maintain their diversity and resilience in light of future challenges (folke et al. 2004). furthermore, many ecosystems may exhibit more than one equilibrium state, with transitions between them that may be somewhat abrupt and irreversible, and that very often are unanticipated. since both human and ecological systems are self-organising, the position of the threshold itself may change over time. finally, the distinction between ‘human’ and ‘natural’ disturbances is often spurious. for instance, people have been important shapers of ecosystem structure and function in africa for millennia and are internal to the system, rather than external forces impinging on it. these considerations, and a realisation of how expensive (and often unintentionally injurious) ongoing interventions on multiple fronts can be, have led to a much greater reluctance on the part of conservation managers to intervene in ecosystems. furthermore, the conservation authorities themselves, and interested parties outside the agencies, began to ask questions regarding what the desired state is, and who decides it. this trend started in the 1980s, and in south africa, accelerated after 1994 as a broader and more participatory style of governance took root in many facets of south african society. in this paper, we start from a position that there is probably no universally ‘right’ style of ecosystem management (although it is likely that there are some that are clearly ‘wrong’ in a given situation). there are circumstances in which one style is more likely to achieve the desired ends, at lower financial and ecological cost, than others. command and control is a reasonable approach when the level of understanding and the capacity to effect change are both high. where the understanding of how the system works is essentially no better than blind guesswork and the capacity to take directed action is very low, a non-interventionist stance (sometimes called a ‘laissez faire’ approach) is both theoretically defensible and practically unavoidable. in the case where understanding is high but the capacity or desire to intervene is low, ‘watchful tolerance’ might be a sensible philosophy. south african national parks (sanparks) are generally neither completely powerless nor completely ignorant. furthermore, there is a societal and legislated expectation that the conservation authorities act to meet various explicit and implicit objectives, including the avoiding of ecosystem degradation and the prevention of species extinction. the park managers have demonstrated a capacity to influence certain ecosystem outcomes, even on a large scale, and are relatively well resourced to do so. after more than half a century of research, some things are relatively well understood, and at least reasonable hypotheses as to what might be going on can be posed on most topics. under these circumstances, we argue that the most reasonable management philosophy is ‘adaptive management’ (biggs & rogers 2003). in this style of management, objectives are stated provisionally but explicitly – in other words, they are equivalent to ‘hypotheses’ in experimental research. management actions are treated as experiments from which learning can take place, even if the outcome is not the expected one. this requires that the interventions are designed and implemented with enough scientific rigour for the outcomes to be attributed to causes. the interventions are undertaken on a scale and with an intensity that permits the consequences to be distinguished against the background noise of inherent variation. the actions are documented and the outcomes are monitored. it is an axiom of this style of management that the actions and, if necessary, the objectives are subject to change on the basis of evidence provided by the experiment itself, or because changing societal expectations require it. gregory, ohlson and arvai (2006) highlight four criteria that must be met for adaptive management to be valid; (1) it must be able to cope with the spatial and temporal scale of the issue, (2) it must deal explicitly with the uncertainty associated with observing and making predictions, (3) it must evaluate costs, benefits and risks and (4) it must have stakeholder support. this is the context in which sanparks, starting in the kruger national park, began to define their management plan, based on the ‘thresholds of potential concern’ (tpc) (biggs & rogers 2003). the approach is summarised briefly as follows: a set of high-level goals is defined and then translated into more specific objectives. a threshold or thresholds are then set in relation to selected indicators of each objective. the thresholds are often expressed in the form of a traffic light analogy: an ‘amber light’ warns of an impending problem, and a ‘red light’ triggers an intervention. initially the tpcs were determined by expert judgement, following a debate among specialists and a review of the evidence that was available. the tpcs have subsequently been refined and reviewed in several iterations. problems have surfaced as the system has been implemented. for instance, the number of tpcs and indicators has proliferated, which makes the system burdensome and open to internal contradictions. secondly, in the cases where tpcs have been transgressed, the reaction has most often been to simply redefine the acceptable level of change, rather than to embark on an expensive and far-reaching intervention. this underscores the often arbitrary nature of the ‘thresholds’ that have been set. this paper suggests a conceptual framework for setting trigger points for management action, such as the sanparks tpcs, in time-varying ecosystems, taking into account uncertainty and the time lags inherent in both ecosystem processes and intervention actions. a general approach to the problem (back to top) the approach outlined below is analogous to a control-theory problem (brogan 1985), such as guiding a missile to a moving target or steering a vehicle down a twisty road. the control system consists of a sensor, which provides information (including a degree of error) about the position and motion of the object relative to the current location. this information is used to activate course adjustments, taking into account the time lag in the response of the system. in this paper, we assume that the highest-level objective has been set, that is, there is clear guidance regarding what the area is managed for. in the missile analogy, the target has been selected. in our view, the top-level goal is not principally a technical decision, but a societal one. a ‘political’ process – in other words, a legitimate, values driven consensus-based process – must be followed to decide on the desired state, taking into account the various tradeoffs, perceptions and power relations between the stakeholders. once the overarching purpose has been agreed upon (accepting that this social consensus process is non-trivial and will always be provisional and dynamic), the following, more ‘technical’ (i.e. expert-based) analysis can follow. the way in which the approach is developed is illustrated in figure 1 (panels [a] to [c]). figure 1: steps in establishing a management action trigger. define a sensitive, relevant and measurable indicator of the issue at hand although it typically is a system that is being managed, with many component parts, we need to settle on a single measure of system performance for tracking purposes, otherwise potentially contradictory signals will result. ‘indicator’ is a term that also embraces proxy measures, but the preferred indicators are the ones most closely linked to the actual factor of concern. for instance, if soil loss is the issue, a direct measure of soil export in tonnes per hectare is unambiguous; an indirect observation such as the area of bare ground is less preferred, while an index such as a calculation based on the universal soil loss equation (usle, see risse et al. 1993) is only indirectly based on observations. if complex indices are chosen, then those with physical units and a clear and established underlying logic (such as usle) are by far preferable to arbitrary combinations of indicators, such as averages of a number of variables that are thought to be related to the state of the system in some way. set a limit of acceptable change in the indicator in this paper, we avoid calling the limit of acceptable change a ‘threshold’ in order to prevent it from becoming confused with the notion of a system threshold separating one state from another; in other words, a boundary in system state-space at which negative feedbacks are overwhelmed by positive feedbacks, and thus, when crossed, causes the system to accelerate into a new domain of attraction under its own dynamics. many of the limits that are typically set for ecosystem management do not have this property of discontinuous change, but are simply agreed levels along a continuum, beyond which you do not wish to proceed, for whatever reason. nonetheless, it is very helpful to the identification of limits if they are true system thresholds, since this introduces a much stronger element of causality and non-arbitrariness into the definition, which usually cuts out a lot of unproductive debate. furthermore, we argue that these ‘true’ thresholds of relative irreversibility (very few things are truly irreversible, if you have enough time and money to fix them) are the ones that the managers of protected areas really need to worry about. therefore, selecting system thresholds as the limits, even if they are relatively rare, has the advantageous effect of reducing the number of criteria that have to be monitored. (see martin et al. [2009] for a discussion of the differences between ecological thresholds, utility thresholds and decision thresholds.) here we are concerned with decision thresholds, but it is helpful if they are founded on ecological thresholds. note that ecological thresholds are themselves dynamic, so the process of defining the limit needs to be revisited periodically. add a safety margin based on societal risk tolerance and technical certainty it is very rare that ecologists know the position of a threshold or limit exactly. neither is the present state of the system known with absolute precision. faced with a situation of risk, most people prefer to stay well away from the edge – especially if they are not too sure where the edge lies. our willingness to stay near to the limit depends on what is at stake. for instance, if a population were allowed to drop below some minimum viable level, would the result be a global extinction, or just a local extirpation? the level of risk taken also depends on the appetite for risk among the stakeholders. for example, people struggling to meet basic human needs are thought to be willing to take greater local environmental risks in meeting those needs than populations that are more comfortably off (goklany 2001). the central point is that the magnitude of the safety margin is not simply a technical or statistical issue, but a decision that is taken within a particular ecological and social context. figure 1(a) illustrates the steps so far. project the indicator into the future using a model, including an uncertainty range knowing that you have crossed a limit after the fact is not very helpful. it is much more useful to be able to predict well ahead of time that you are likely to cross the line, so that corrective action can be taken. therefore, decisions should generally be made on the basis of some sort of predictive ‘model’. models can range in sophistication from a simple extension of a trend line to a complex, multi-variable, non-linear systems model. expert judgement and experience are also ‘models’, but not very transparent ones. any model, regardless of its sophistication, has uncertainty associated with it. for instance, the 95% confidence interval can be calculated for a linear trend, as is suggested in figure 1(b). in the case of a complex systems model with many variables, each of which has some uncertainty, the joint uncertainty distribution is quite complex. this uncertainty is often approximated by running an ensemble of different models incorporating different assumptions, or else by running the same model but varying the key parameters in a statistically realistic way within their likely ranges (‘monte carlo simulation’, vose 1996). note the time at which the relevant uncertainty limit intercepts the safety margin the approach we describe aims to be conservative. therefore the key point is not where the mean of the forward projections breaches the safety-buffered limit that you have set, but the earliest moment when there is a reasonable probability of such a limit being exceeded. note that for ‘upper limits’ being approached from below, it is the higher uncertainty range that matters, whereas for ‘lower limits’ being approached from above it is the lower uncertainty range. what is ‘reasonable’ must be considered against the risk tolerance of the stakeholders, and it should be consistent with the implied risk tolerance used when setting the safety buffer, described above. in many cases, the default is the widely accepted scientific norm of the 95% confidence limit – in other words, there is a less than one in 20 chance that the safety margin will be breached earlier than the time projected. subtract the system response time from the time when the limit is crossed by the lower 95th percentile confidence limit, we subtract a period that reflects the inertia in the ecosystem – in other words, the response time following an intervention. in our car analogy, think of it as how responsive the vehicle is to a touch on the steering wheel. if the car is fast and heavy, the turning circle will be large and the response sluggish, but if the car is light and agile, last-minute course corrections are possible. ecosystem inertia can have many causes. one of the most common sources of ecosystem inertia is the longevity of the organisms involved. for example, if you try to alter the population dynamics of an elephant herd through contraception, you need to start a decade ahead of time, since that is more or less the period between birth and sexual maturity. if you are similarly intervening in an impala population, 2 years would be sufficient. sources of ecosystem inertia that are short-lived (weeks) are probably virtually irrelevant in this context, as are exceedingly slow processes – for instance, land-forming processes that take millennia. the most critical issues for management lie inbetween – processes where you need to act years to decades before the threshold of non-return is reached if you wish to materially affect the outcome. multiple runs of the projection model, starting at progressively earlier times and simulating the appropriate corrective action in the model until a starting moment is reached, that does not result in the safety limit being transgressed, is a way of assessing the inertial delay. bear in mind that the models are invariably imperfect – so an informed judgement based on a consideration of the key lags in the ecosystem may be nearly as good. subtract the management response time it is not only the natural world that has delays built into it – the human side of the system does as well. for instance, you may need to wait until the next management meeting, or the next budgetary cycle, to commence an action. or you may need to initiate a protracted stakeholder engagement process. while it is tempting to think that we have a bit more control over human delays than we have over the time lags in the ecosystem, experience teaches us that big decisions take time to implement. the uncertainties in the magnitude of the management response time underscore the futility of trying to be too precise in estimating the ecosystem response time, since the two numbers are added together. subtract the monitoring interval a further time lag must be subtracted to allow for the fact that observations of the system take place only occasionally. sometimes the period is very short – for instance, satellite-derived images of fire extent are obtained weekly. in other cases, the interval may be long: for example, full census data for large mammals in the kruger national park are only obtained once every 3 to 5 years because of the effort and expense of doing so. on average, you are only half a monitoring interval away from the next estimate, but in order to be conservative, this lag should be set as equal to the monitoring interval. is an action triggered? since the point at which we identified when the conservative limit of projected trend intercepts the safety-buffered threshold of concern we have been working in time units, backwards from that moment. the final steps in the procedure are illustrated in figure 1(c). we first subtracted the ecosystem response time, then the management response time, then the monitoring interval. if the result is a date that is at or before the current date, then an intervention is triggered. if the result is after the current date, then, in theory, you can safely wait until the next monitoring date, assuming that reanalysis will occur at that time on the basis of the updated data. if the indicated trigger date is between the current date and the next monitoring date, it is appropriate to indicate an ‘amber’ (caution) light, which would set off various actions aimed at increasing your confidence or proactively decreasing the various delays in the system. for instance, you could shorten the monitoring period or intensify the monitoring effort for the key variable involved; you could commission research to improve the confidence in the projection model; you could take actions to reduce the delays in the decision cycle; or you could take actions to reduce the ecological lag – for instance, you could proactively administer immuno-contraception primers to reduce population response delays to contraception. often you would undertake some combination of these actions. the practical application of these steps is illustrated in a worked example below. example: elephants and big trees in the kruger national park (back to top) the elephant population in what is now the kruger national park and surrounding private and state-owned wildlife areas had been reduced to close to zero at the beginning of the 20th century. small groups of elephant began to wander into the region from mozambique after the first reserves were established. by 1967, the elephant population had reached an estimated 6600 through a combination of in-migration and natural increase. at this time, on the basis of expert consideration of the elephant densities in a range of protected areas in africa, it was decided to limit the number of elephants in kruger park to a total of 7000 (which corresponded to ‘one per square mile’, the threshold above which several experienced persons at the time considered unacceptable damage to the vegetation to occur). the kruger park elephant population was kept near this level through annual culling until 1994, when a moratorium on culling was imposed in response to pressure from animal rights advocates. since then, the population has grown at a rate of approximately 6% per annum, and an intense debate is underway about what action to take (if any) in the future, and what the trigger points might be (carruthers et al. 2008, and other chapters in scholes & mennell 2008). we have chosen this example because it nicely illustrates several points made in the preceding section. our analysis of the topic is solely for the purposes of illustrating the procedure we propose for setting action thresholds. it does not represent the policy of the south african government, sanparks or the authorities in the kruger national park. while the worked example is broadly grounded in observed data, in some respects it involves our informed judgement (as will be the case in almost any real-life situation). if a management trigger were to be set in practice for the elephant-tree system using an approach based on the one we describe, it would need to be based on a process somewhat more inclusive than the opinion of these two authors alone. choosing the indicator the first thing that can be learned from the elephant-and-tree example is that taking an overly simple and purely technocratic view is likely to lead to implementation problems sooner or later. the culling moratorium was motivated partly by the arbitrariness of the ‘one elephant per square mile’ rule, the origin and validity of which was never properly researched or documented. as the scientific assessment of elephant management in south africa (scholes & mennell 2008) points out, elephant population density per se is not the problem. elephants are part of a system, which involves, at a minimum, elephants and the plants that they consume or damage. an even wider view would also consider indirect ecosystem effects (for instance, other species that are in competition with elephants for food, or are affected by the habitat changes elephants bring about). a yet wider view contextualises elephants within human society and the choices this society collectively makes regarding the desired appearance of landscapes and the relative importance of different species.therefore, the indicator variable on which the intervention trigger is set in this case should not be the size of the elephant population, but some variable that reflects our underlying concern for the effects the elephants have on the ecosystem. the issue that usually surfaces in consultation with a range of stakeholders in the context of the kruger park ecosystem is damage to trees (and, specifically, large and old trees) caused by elephants. this often has an aesthetic component – visitors have grown accustomed to seeing the kruger park landscape in a certain way, with scattered large trees and dense riparian forests, and do not wish to see a landscape consisting of bushy coppice and downed trees, regardless of whether the latter situation supports the same productivity or biodiversity. this is a legitimate stance, but raises difficult questions in terms of setting an operational limit – whose opinion counts in the poorly defined trade-off between the appearance of a landscape and the desire not to interfere with the lives of elephants? how much landscape transformation is acceptable, and to whom? setting a safety-buffered limit further unpacking of the ‘large tree issue’ reveals that a whole guild of species – the large raptors – are almost totally dependent on large trees for nesting opportunities. since there is well-established literature on the importance of maintaining top predators and scavengers in functional ecosystems (e.g. terborgh et al. 1999), conserving large raptors by providing nesting opportunities provides a somewhat more robust ecological basis for setting a limit. large raptors and vultures occur relatively sparsely and, in some cases, are rare outside large protected areas such as the greater kruger national park. some are arguably dependent for their regional population viability on having a core breeding population in the kruger park. the lower limit for a minimum viable population could therefore conceivably be reached if the breeding habitat was restricted. population viability analysis (shaffer 1990) could be applied to determining what the safe lower limit of breeding pairs might be. since the birds nest in the crown of live trees, typically at a height of 12 m or more above the ground, this minimum population level can be translated into a number of nesting sites, and, in turn, into a minimum density of large trees. we do not intend to do this full analysis here, but simply assume that it works out at about one tree per square kilometre, including a generous safety margin. (in practice, because the decline in large trees accelerates as they become rarer, the exact value of the threshold hardly matters in setting the trigger. this is a very desirable property in an indicator.) this can be considered a ‘ threshold of irreversibility’, since the regional or global extinction of some species could result if the large tree population falls below this threshold. projecting the dynamics forward reasonably reliable data exist to build simple models of both elephant population dynamics and tree demography. the two models can then be coupled to create a model of the elephant-tree system (for example, duffy et al. 1999; owen-smith 2002). a simple implementation of such a model is applied here, again for illustrative purposes only. because important time lags are introduced by both elephant maturation and the growth of large trees, the minimal model needs to include an age structure. our discrete-timestep elephant model has annual cohorts of female elephants (the number of bulls is simply assumed to equal the number of females) that reach sexual maturity at age 12, and thereafter produce one calf every 2.5 years until death at age 60. the basic rate of mortality is 3% per year for all size classes, and increases as food becomes scarce. the values are chosen to reproduce the observed 6% per annum growth rate in the elephant population, as well as the extensive research on elephant population dynamics in the kruger park system (van aarde et al. 2008). no direct effect on population growth rate resulting from food shortage or other density-dependent phenomena has been observed at the densities experienced to date in the kruger park ecosystem, but there are data from elsewhere in southern africa to suggest that the inter-calf interval extends when forage becomes scarce (van aarde et al. 2008) and, at some degree of forage inadequacy, mortality must occur. these effects should be included in the model if it is to be realistic when projected forward sufficiently to take into account the inertia of tree and elephant populations. forage is produced by both trees and grasses, in an inverse relationship, and a portion of it (set nominally at half) is consumed by other species of herbivores. the tree model is based on discrete size classes of stem diameter, in 5 cm increments up to a maximum diameter of 80 cm. all species are combined. there is a rough relationship between stem diameter (d, cm) and height (h, in m): h = -0.011 d2 + 0.2162 d+1.336 derived from a database of several thousand stems measured in the kruger national park (scholes, unpublished data). trees taller than 12 m have a diameter at breast height of 80 cm or more on average. the same database was used to calculate the size-based mortality rates for trees. on average, the number of trees in each size class is 32.3% fewer than the number in the next smallest diameter class, for all classes with a diameter greater than 5 cm. in order to allow the size-structured tree model to run on a common basis with the age-structured elephant model, it is necessary to know the stem diameter growth rate. based on multi-year repeated measurements on savanna trees throughout south africa (shackleton 1998), the percentage annual growth rate in stem cross-sectional area (da) is a function of both the diameter of the stem and the degree of competition to which it is exposed: da = (1.0 + 19.0 e-0.2d )*(1-b/bmax) where b is the basal area of woody plants (m2/ha) and bmax is the maximum basal area that can be attained given the rainfall (converted from the data used by sankaran et al. 2005 into basal area terms, bmax= 0.03 [map-100]), which in this example is about 12 m2/ha for the average kruger park rainfall of 500 mm/year. seed production was made proportional to stem cross-section, and began when the trees reached a third of their maximum height. a combined germination and survival rate for seedlings up to the 5-cm diameter class was estimated on the basis of the assumption that the tree population was initially in a steady state. coupling of the two models was achieved by assuming that elephants have two main impacts on trees. for trees larger than 15 cm in diameter but smaller than 40 cm, the stem is snapped or uprooted as a result of being pushed over by elephants in order to browse on the canopy foliage. for trees smaller than 15 cm in diameter, the canopy is entirely within reach and, although browsed, the main stem (if one exists) is not broken. trees larger than the upper limit are harder to break or uproot, but are subject to debarking. the resulting mortality is complex, depending on the fraction of bark circumference lost and the presence of other agents, such as wood-boring insects and pathogens. we assumed that tree death occurred when the equivalent of a full circumference was debarked. in the absence of data, we assumed that each elephant strips 1 m of bark circumference per day, distributed over the size classes in proportion to their circumference. the fraction of a size class that dies per year (fi) through this mechanism is thus: fmortality = [ni pdi /s(ni pdi )] [365 e /nipdi ] where ni is the number of trees in size class i (km-1), pdi is the midpoint circumference of size class i, snipdi is the sum over all size classes of the stem circumferences, and e is the number of elephants (km-1). pushed-over trees do not die, but revert to the smallest size class as coppice resprouts. female elephants push over an average of one tree a day and bulls, four times as many. when is the limit likely to be transgressed? starting this model with the observed tree size distribution and the observed elephant age distribution for the kruger park shows the trajectories illustrated in figure 2. note that our model projects a slow decline in the number of large trees even in the absence of elephants, suggesting that the observed tree size distributions are not in a steady state. the ensemble results presented in the same figure are the year in which the threshold is crossed in a hundred independent model runs, when both a key ‘tree’ parameter (the mortality due to elephant damage) and a key elephant parameters (mortality and its dependence on density) were systematically and independently varied according to a normal distribution using the mean and standard deviation reported in the literature. more of the assumptions in the model could be tested in this way, but this scheme is sufficient to demonstrate the principle of creating an uncertainty range. figure 2: an example of a model for a hypothetical elephant-tree system, based on data from the kruger national park. the lower 95th percentile of the predicted number of trees taller than 12 m intersects the chosen threshold of one tall tree per km2 at a time 60 years from the simulation start (which nominally represents year 2009). this is the starting date from which we begin the intervention calculation. subtracting the response times in this example, we have assumed that the chosen management intervention is to reduce the fertility of elephant cows using immuno-suppressant contraception. a 75% success rate is assumed (bertshinger et al. 2008). running the model using just the ‘best estimate’ values for the parameters but reducing the fertility of the cows by this fraction results in the response trajectories illustrated in figure 3, depending on when the intervention begins. by plotting the years of pre-emptive management against the extra years of non-transgression of the threshold thereby gained, we can deduce that the ‘inertia’ of the elephant-tree system as defined above is in the order of 30 years. figure 3: the effect of the year of commencement of elephant contraception on the decline of large trees. we further estimate that it would take about 3 years to launch a contraception programme of this magnitude, including making the decision, getting approval in terms of an elephant management plan approved by the minister concerned, consulting with stakeholders, manufacturing the vaccines, and conducting field operations to vaccinate all sexually mature elephant cows. finally, we added a monitoring interval for the key elements in the system (trees taller than 12 m, and elephant population) of one year. is an action triggered? the indicated time to trigger a decision regarding elephant contraception in order to prevent the limit of one tall tree per km2 being exceeded, thus, is the year in which more than 5% of the model runs indicate the limit being breeched (2069), minus the sum of the system response time (30 years), the management delay (3 years) and the monitoring interval (1 year). this suggests that a decision would need to have been made, at the latest, by 2035. we stress once again that this is a theoretical example only. discussion and conclusion (back to top) this paper has outlined a logical procedure for triggering management interventions in systems that are dynamic and prone to measurement error and theoretical uncertainty. the steps seem complicated, but we would argue that they are necessary and that the level of detail aimed for in any one of them needs to be matched to the uncertainty inherent in the full logical chain, which is determined by the data that are available or readily accessible. the effort involved should be guided by the consequences of getting the signal wrong, either as a result of initiating an action where no action was needed, or by failing to do so when intervention was necessary. the process we have described reveals where the key uncertainties lie and what can be done to address them. finally, it is clear that several of the steps (selecting the overall objective, setting the level of risk aversion) are not purely technocratic, but require input from stakeholders in a legitimate fashion. on the other hand, there are elements (particularly the forward projection of system trajectories) that do require specialised technical input, and the quality of those technical inputs can be increased through investment in monitoring and research. acknowledgements (back to top) two anonymous reviewers made useful comments that improved the paper. references (back to top) bertshinger, h., delsink, a., van altena, j.j., kirkpatrick, j., killian, h., ganswindt, a., et al., 2008, ‘reproductive control of elephant’, in r.j. scholes & k.g. mennell (eds.), elephant management: a scientific assessment for south africa , pp. 257−327, witwatersrand university press, johannesburg. biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience, pp. 59−80, island press, washington dc. brogan, w.l., 1985, modern control theory, prentice-hall, new york. carruthers, j., 1995, the kruger national park: a social and political history, university of natal press, pietermaritzburg. carruthers, j., boshoff, a., slotow, r., biggs, h.c., avery, g. & matthews, w., 2008, ‘the elephant in south africa: history and distribution’, in r.j. scholes & k.g. mennell (eds.), elephant management: a scientific assessment for south africa, pp. 23−83, witwatersrand university press, johannesburg. duffy, k.a., page, b.r., swart, j.h. & baji, v., 1999, ‘realistic parameter assessment for a well known elephant–tree ecosystem model reveals that limit cycles are unlikely’, ecological modelling 121, 115−125. doi:10.1016/s0304-3800(99)00091-5 ellis, j.e. & swift, d.m., 1988, ‘stability of african pastoral ecosystems: alternate paradigms and implications for development’, journal of range management 41, 450−459. doi:10.2307/3899515 folke, c., carpenter, s., walker, b., scheffer, m., elmqvist, t., gunderson, l., et al., 2004, ‘regime shifts, resilience and biodiversity in ecosystem management’, annual review of ecology and systematics 35, 557−581. doi:10.1146/annurev.ecolsys.35.021103.105711 goklany, i.m., 2001, the precautionary principle: a critical appraisal of environmental risk, cato institute, washington dc. gregory, r., ohlson, d. & arvai, j., 2006, ‘deconstructing adaptive management: criteria for application to environmental management’, ecological applications 16, 2411−2425. doi:10.1890/1051-0761(2006)016[2411:damcfa]2.0.co;2 holling, c.s. & meffe, g.k., 1996, ‘command and control and the pathology of natural resource management’, conservation biology 10, 328−337. doi:10.1046/j.1523-1739.1996.10020328.x martin, j., runge, m.c., nichols, j.d., lubow, b.c. & kendall, w.l., 2009, ‘structured decision making as a conceptual framework to identify thresholds for conservation and management’, ecological applications 19(5), 1079–1090. doi:10.1890/08-0255.1, pmid:19688917 owen-smith, n., 2002, ‘credible models for herbivore-vegetation systems: towards an ecology of equations’, south african journal of science 98, 445−449. risse, l.m., nearing, m.a., laflen, j.m. & nicks, a.d., 1993, ‘error estimation in the universal soil loss equation’, soil science society of america journal 57, 825−833. doi:10.2136/sssaj1993.03615995005700030032x sankaran, m., hanan, n.p., scholes, r.j., ratnam, d.j., augustine, b.s. cade, j., et al., 2005, ‘determinants of woody cover in african savannas’, nature 438(7069), 846−849. doi:10.1038/nature04070, pmid:16341012 shackleton, c.m., 1998, ‘annual production of harvestable deadwood in semi-arid savannas, south africa’, forest ecology and management 112, 139−144. doi:10.1016/s0378-1127(98)00321-1 shaffer, m.l., 1990, ‘population viability analysis’, conservation biology 4, 39−40. doi:10.1111/j.1523-1739.1990.tb00265.x scholes, r.j. & mennell, k.g. (eds.), 2008, elephant management: a scientific assessment for south africa, witwatersrand university press, johannesburg. terborgh, j., estes, j., paquet, p., ralls, k., boyd-heger, d., miller, b., et al., 1999, ‘the role of top carnivores in regulating terrestrial ecosystems’, wild earth 9, 42−56. van aarde, r., ferreira, s.m., jackson, t.p., page, b., de beer, y., junker, j., et al., 2008, ‘population biology and ecology’, in r.j scholes & k.g. mennell (eds.), elephant management: a scientific assessment for south africa, pp. 84−l45, wits university press, johannesburg. vose, d., 1996, quantitative risk analysis: a guide to monte carlo simulation modelling, john wiley & sons, new york. article information authors: craig a. mcloughlin1 andrew deacon1 hendrik sithole1 thomas gyedu-ababio2 affiliations: 1south african national parks, conservation services, kruger national park, south africa 2south african national parks, conservation management, kruger national park, south africa correspondence to: craig mcloughlin email: craig.mcloughlin@sanparks.org postal address: private bag x402, skukuza 1350, south africa dates: received: 15 apr. 2010 accepted: 02 nov. 2010 published: 13 may 2011 how to cite this article: mcloughlin, c.a., deacon, a., sithole, h. & gyedu-ababio, t., 2011, ‘history, rationale, and lessons learned: thresholds of potential concern in kruger national park river adaptive management’, koedoe 53(2), art. #996, 27 pages. doi:10.4102/koedoe.v53i2.996 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) history, rationale, and lessons learned: thresholds of potential concern in kruger national parriver adaptive management in this essay... open access • abstract • introduction    • heterogeneity and the knp river systems    • a strategic adaptive framework for managing knp rivers    • lags in implementation of sam • the operational thresholds of potential concern    • tpcs for river flow       • application, refinements and assessment post-knprrp    • tpcs for water quality       • rationale       • application, refinements and assessment post-knprrp       • rationale • the biodiversity thresholds of potential concern    • in-stream tpcs    • tpcs using fish indicators       • rationale       • application, refinements and assessment post-knprrp    • tpcs using macro-invertebrate indicators       • rationale       • application, refinements and assessment post-knprrp       • riparian tpcs       • tpcs for sedimentation and habitat loss    • tpcs for biotic response to sedimentation and habitat loss       • rationale       • application, refinements and assessment post-knprrp    • tpcs for biotic response to changes in flooding regime       • rationale       • application, refinements and assessment post-knprrp • lessons learned    • people: motivated, dedicated and adaptable    • suitable benchmarks against which to measure indicators    • information: generation and use    • implementation lags: key processes    • timing: between monitoring and management action    • feasibility: monitoring programmes    • refinements: ongoing tpc requirement    • adaptive feedback loops: identification and closing • kruger national park river tpcs – future directions    • the ecological reserve    • aquatic and riparian ecosystem monitoring • acknowledgements • references • appendix 1    • conservation services management committee meeting, kruger national park       • notification of river tpc reached    • background    • definition of the threshold of potential concern    • data or information supporting the threshold of potential concern • results    • consequences or potential impacts    • proposed management action • references abstract (back to top) the kruger national park’s (knp) adopted system of management, called strategic adaptive management (sam), originated during the kruger national park rivers research programme (knprrp) of the 1990s. an important concept in sam is the thresholds of potential concern (tpcs), representing end-points in a continuum of change. tpcs within the knp sam system guide management if or when reached, ‘red-flagging’ possible negative biodiversity impacts and catalysing consideration of management options. tpc-related monitoring generates the strategic information for ongoing evaluation, learning and adaptation within sam. post-knprrp, although river flow and water quality tpcs have been implemented partly, those designed to detect undesirable changes in biodiversity have not been implemented, until recently. this paper describes the history, rationale, application and ongoing developments associated with the knp river tpcs over the last decade, providing some key lessons for organisations utilising sam. the paper concludes with an overview of new thinking and future directions envisaged for the knp river tpcs, as part of the knp sam system. conservation implications: this paper documents important concepts of strategic adaptive management associated with the knp river systems. understanding, related to the rationale and justification for use and development or refinement of the thresholds of potential concern, lays an important foundation for ongoing work in managing these rivers adaptively. introduction (back to top) the paradigm shift in ecology from ‘balance’ to ‘flux’ of nature embraces notions about ecological systems as complex and dynamic, rather than homogeneous, linear systems (pickett, parker & fiedler 1992). furthermore, change in conservation focus, from species to explicit ecosystem management, emphasises spatial and temporal change or heterogeneity (rogers & bestbier 1997; rogers 2003). with trends leaning towards biodiversity management at the ecosystem level (poff et al. 1997) and a requirement for alternate methods to manage complexity in heterogeneous systems (pickett, cadenasso & benning 2003; biggs & rogers 2003), it became opportune to instigate adaptive management processes for rivers in and around the kruger national park (knp). this was fuelled by concerns about the deteriorating biophysical conditions of the knp’s aquatic and riparian ecosystems, due to upstream influences, for example, water abstractions, impoundments and pollution (breen et al. 2000). the kruger national park rivers research programme (knprrp) was initiated towards the late 1980s to further river conservation efforts (breen et al. 2000). a major thrust was the development of capacity to predict impacts of upstream influences on the biophysical condition of the rivers, for mitigation purposes. breen et al. (2000), rogers and biggs (1999) and rogers and o’keeffe (2003) provide comprehensive reviews of this approximately 10-year multidisciplinary research programme, ultimately leading to the adoption of a strategic adaptive management system for knp river management. the major challenges to scientists during the knprrp era included a need to (1) develop and describe a common understanding of the heterogeneity and dynamics of the knp’s river systems and then (2) to articulate this understanding to management within a strategic framework (rogers & biggs 1999). heterogeneity and the knp river systems the first (1988–1993) and second (1994–1996) phases of the knprrp provided a multi-disciplinary, scaled understanding and predictive base for better decision making involving the knp’s river systems. this was important because causal links between catchment processes (land-use, water abstractions and sediment production) and downstream biotic responses were previously fraught with problems of scale and inter-disciplinarity (rogers 1997). spatial-temporal heterogeneity in the physical template of the river landscape, encompassing a highly variable flow regime and geology, governing patterns of sediment erosion, transportation and deposition, were key concepts. the formation of a diverse set of both alluvialand bedrock-controlled geomorphic features within the river landscape leads to a diversity of habitat types (van niekerk, heritage & moon 1995; heritage, van niekerk & moon 1999), and biota that respond to these (e.g. van coller, rogers & heritage 1997; russell 1997; van coller, rogers & heritage 2000; rogers & o’keeffe 2003). knowledge of this type became the basis for predicting biodiversity responses to changing catchment conditions that affect and influence the river systems (rogers & o’keeffe 2003). a strategic adaptive framework for managing knp rivers managing complex systems characterised by uncertainty requires an adaptive, learning-by-doing approach (rogers 2003; stankey, clark & bormann 2005, see editorial by roux & foxcroft 2011). indeed, a key principle is that appropriate management actions should be implemented, even if based on imperfect knowledge, because complete understanding is never attainable realistically (rogers 2003). furthermore, management actions present potential learning opportunities, feeding back more reliable information to improve decision making (rogers & biggs 1999). the strategic adaptive management (sam) framework pioneered during the final phase (1997–1999) of the knprrp requires the articulation of a vision, a hierarchy of objectives, ultimately leading to specific end-points to guide management (see rogers & bestbier 1997). the development of purposeful monitoring programmes to audit against these end-points is important (see rogers & bestbier 1997; rogers and o’keeffe 2003). the end-points, known as thresholds of potential concern (tpcs) within sam, are upper and lower levels of acceptable change that when exceeded, set in motion a process of investigation as to the reasons why the tpc was exceeded (biggs & rogers 2003). possible management action, or revision of the actual tpc, is then considered (biggs & rogers 2003). the first tpcs developed for the knp rivers (from knprrp) were linked to a vision of maintaining biodiversity in all its natural facets and fluxes, via a hierarchy of objectives (see rogers & bestbier 1997; rogers & biggs 1999 for a detailed account). these tpcs addressed issues of river flow, water quality (physical-chemical guidelines), and sedimentation (geomorphic indictors). in addition, tpcs were developed to test biotic response to deteriorating river flows and unnatural increases in sedimentation (fish, macro-invertebrates and riparian vegetation indicators), and alterations to the flooding regime (riparian vegetation indicators), resulting from upstream human influences. although identified as a potential problem, a tpc for reed cover was not completed because of capacity issues. tpcs were also developed for alien vegetation invasions (foxcroft 2009), relating invasions to the introduction, spread and increase in abundance of alien species. lags in implementation of sam until recently, however, only the river flow and water quality tpcs had been implemented sufficiently within the knp river sam system, post-knprrp. many factors contributed to this outcome. within the knp, a full implementation of the monitoring and reporting systems did not get off the ground. this diminished the capacity to feed back information into evaluation and learning, critical within any adaptive management system. in the wider catchment context, lags in implementation of environmental flow requirements had been (and remain) a major setback for sam. this has diminished motivation for testing the biophysical response components. knp influences outside of the park have, however, been partially successful in improving river flows. in this paper, we discuss the river tpcs (operational and biodiversity-related) within the knp, involving pertinent monitoring, reporting and feedback processes. we focus firstly on the history and rationale for the specific tpcs, including application and ongoing developments. we then highlight key learning opportunities achieved so far, post-knprrp, pertinent to other organisations utilising sam. we conclude by providing an overview of new thinking and future directions envisaged for the knp river tpcs. the operational thresholds of potential concern (back to top) tpcs for river flow rationale under the south african national water act (act 36 of 1998), the quantity and quality of water required to protect the ecological functions on which humans depend (defined as the ecological reserve) must be determined and maintained in natural water resources (o’keeffe & rogers 2003). over the past decade, the knp has monitored environmental flows within all major rivers according to in-stream flow requirements (ifrs; the minimum flow required to sustain ecologically healthy rivers), determined during the 1990s using the ‘building-block’ methodology (see king & louw 1998) (see table 1). although the ifrs were well researched, they are considered minimum flows required to avoid unacceptable biodiversity loss in the rivers. in addition, unnaturally high flows are of concern, leading to biodiversity impacts too. for example, the practice of releasing high water volumes during low flow periods may provide breeding cues for fish at the wrong time of the year (russell 1997). table 1: in-stream flow requirements, as determined for the kruger national park rivers using the ‘building block’ methodology. for a full treatise, a number of sources provide detailed information on methods used in the development of ifrs for each of the knp rivers: crocodile river – csir (2002); olifants, letaba and shingwedzi rivers – dwaf (1991); sabie river – tharme (1997) and luvuvhu river – louw and o’keeffe (1999). application, refinements and assessment post-knprrp although high flows are incorporated within the ifr tables, only the low flows under the maintenance and the drought scenarios have been utilised as tpcs (see table 1). there has been frequent tabling (and un-tabling) of river flow tpcs, using the ifr levels as lower thresholds to river flow. notably, the knp river management have experienced many challenges associated with actual compliance with these ifrs (see figure 1). figure 1: history of in-stream flow requirement tpc exceedances in the major rivers of the kruger national park over the last decade. the implementation of the river flow tpcs over the past decade has provided key opportunities to improve the system within the sam framework. firstly, reporting of river flow conditions to scientists and managers was considered inadequate for making informed assessments to guide decision making. for example, information for comparing river flow situations between time periods was lacking. furthermore, feedback about management action outside of the knp was not made explicitly available to the relevant people, therefore making it difficult to assess and learn from past actions. in addition, river flow reporting relied heavily on the motivation and discretion of the river technician at that time, making this component highly vulnerable. changes to the river flow tpc system therefore occurred in 2008, and included (1) automation of tpc auditing procedures via the introduction of a simple excel spreadsheet program, where mean daily discharge values are entered; the program indicates if or when river flow tpcs are exceeded, for tabling at the bi-monthly knp conservation management committee meeting where management actions are considered, (2) the summarising of statistics to provide information on trends in river flow and (3) the introduction of a management log to record relevant management information, for later use in feedbacks (see table 2). table 2: the kruger national park (knp) management log, used within the knp strategic adaptive management system for rivers. secondly, the benchmarks against which to measure river flows (i.e. ifrs) have, at least conceptually, become increasingly difficult to use. this is because (1) determining when the maintenance or drought ifrs should be used is intricate, as rainfall influences cannot be predicted easily, (2) the static nature of the ifrs (within months, between years) does not mimic natural flow variability realistically and (3) there are high demands for water in the catchments. some catchments are under severe water demand stress (e.g. the crocodile river), with increasing pressure to allocate scarce water resources before the ecological reserve requirements are met. in essence, it has become difficult for the knp to demand more water without adequate benchmarks against which to measure. therefore, in 2009, changes were proposed to improve the river flow tpc process. this included recalibrating the river flow tpcs, that is, selecting a level between the maintenance and drought ifrs that could be used to initiate management action realistically, considering water demand in the catchments. subsequently however, major rethinking resulted in a decision to discontinue the tpc tabling process for river flow. the main reason was that the bi-monthly timeframe for tpc tabling at the knp conservation management committee meeting was inadequate to catalyse management action in time, prior to undesirable river flow conditions occurring. thus, during a meeting held in skukuza (river flow workshop), knp scientists and managers agreed to start strengthening the more pertinent rapid systems for dealing with deteriorating river flows. since july 2009, a ‘rapid response system’ has been in place, incorporating increasing levels of concern from management if or when river flows reach various critical levels relative to the ifrs. the spreadsheet program (described above) has been modified to indicate (with discharge data inputs) when river flows fall within different ‘worry zones’. this is so that pertinent action, by the individuals in the relevant management level, can take place in time, for example, by contacting the water resource manager to make releases from dams (mcloughlin 2009). for evaluation purposes, feedback is provided to the knp conservation management committee bi-monthly for assessment about the effectiveness of the ‘rapid response system’. this includes a graphic depiction of daily river flow per river, per month, indicating if/when management actions have been taken (see figure 2 [a]), as recorded in the management log. in addition, summary statistics about flows are still presented (see figure 2 [b]), providing information about management performance over a specified time period. initial feedback from managers and scientists suggests that this system does provide better information formats for interpretation and assessment, for improved decision making. figure 2: the new information format for river flows within the kruger national park, indicating rapid response (worry levels) zones. tpcs for water quality rationale according to the knp water provision policy (knp 1989), the ‘abuse’ of perennial rivers in the knp in the form of pollution and abstraction commenced in the 1920s with the advent of industrial and agricultural development in the lowveld and adjoining areas. silting of rivers had also increased from the mid-1940s. the stringent quality requirements set by the department of water affairs and forestry (dwaf) primarily for drinking water have been adopted by the knp as tpcs (table 3). these were then seen as stringent enough for ecosystem requirements as well (however, this is currently not the case). notably, these tpcs (see rogers & bestbier 1997) had been implemented within the knp sam system and integrated with the national water quality monitoring programme under the auspices of the national dwaf (now department of water affairs – dwa). the dwa supply and maintain the equipment and consumables, analyse the samples and maintain a national database. table 3: guidelines used to assess thresholds of potential concern for water quality, associated with the major kruger national park rivers. the present water quality guideline for ecosystem maintenance is incomplete. as an interim measure it was suggested by representatives of the institute for water quality studies, the freshwater research unit of the university of cape town and the institute for water research at rhodes university to use a combination of standards until more information becomes available (table 3). the sources for standards used at different sampling sites along the major knp rivers include: crocodile, letaba and luvuvhu rivers – dwaf (1996), rogers and bestbier (1997); sabie river – rogers and bestbier (1997). a combination of the south african water quality guideline for domestic use, the south african water quality guideline for aquatic ecosystems (dwaf 1996) and those proposed by kühn (1991) are used for the olifants river. application, refinements and assessment post-knprrp it was realised that the isolated efforts of knp towards the management of water quality of rivers that run through the park would not be enough to ensure the achievement of its objectives, since the impacts on the water quality originate upstream before entering the knp. the knp, therefore, decided to spearhead activities associated with integrated water resource management (iwrm) in relevant catchments that involve the park. stakeholder involvement became a priority, leading to the birth of the catchment forums. stakeholder involvement includes a number of activities: awareness creation and specialist inputs on water related issues –monitoring of the quality and quantity of water, waterrelated research, information sharing and knowledge transfer. the crocodile river forum and related activities is a good example of this process. water quality monitoring networks cannot be static entities, but require ongoing change, for example, objectives, technology and data users (sanders & ward 1993). in 2000, the technical committee of the crocodile river forum divided the catchment into four management units for the purpose of water quality monitoring, consisting of the upper kwena, nelspruit, kaap and the kruger national park (see box 1[a]) management units. water quality objectives (wqos) were then developed, based on the newly developed model for resource quality objectives (rqo). this incorporated the dwa target water quality range (dwaf 1996) and the desktop river classifications. the requirements of water users, historical water quality trends and existing standards (including knp tpcs) were considered. hence, a common set of wqos for the entire sub-catchment is being used as the guideline for monitoring of water quality (see box 1[b]), and relevant information is fed back at the forum meetings (see box 1[c] – for example, conductivity measurements). box 1: (a), (b), (c): water quality management units, guidelines and feedback system currently employed by the crocodile river forum. whenever there are non-conformances to the agreed objectives, the sources of non-compliance are investigated and action is taken. this has contributed immensely to improving the management of water quality in the crocodile river catchment. sewage pollution remains a problem however, but improvements are occurring via actions of the crocodile river catchment forum. the biodiversity thresholds of potential concern (back to top) the suite of river tpcs within the knp sam system incorporates a number of biophysical components as well, designed to test effectiveness of the environmental flows and water quality entering the knp river systems. these include (1) the in-stream tpcs – making use of the national river health programme monitoring tools for habitat, fish and macro-invertebrates and (2) the riparian tpcs – for sedimentation, loss of bedrock influence and vegetation changes. in-stream tpcs the river health programme (rhp) was designed in the early 1990s as a nationwide monitoring system to assess river health (see dwaf 1998) and is a component of the national aquatic ecosystem health monitoring programme (dwaf 2008a). the knp (and south african national parks) has been involved with the rhp from the onset, and is actively involved with the development of methodologies, and implementation of these.the aim of the rhp is to generate information for the national state-of-the-rivers reporting. as such, it involves the rapid monitoring of a comparatively large number of sites distributed over all catchments of concern (kleynhans et al. 2005). the rhp has various monitoring tools to generate relevant information that can be used to compile in-stream tpcs. the two sections below describe the history and development of the fishand macro-invertebrate-related monitoring and reporting within the knp, ultimately becoming an important contributor to the rhp. tpcs using fish indicators rationale broad surveys and ad hoc monitoring of fish started in the early 1950s, but from the 1960s more detailed and systematic surveys were undertaken. after surveys of all the perennial rivers were completed, pienaar (1961) published a list of fish species, and a guidebook was produced in 1969. riverine monitoring only started in earnest when the knprrp was launched in 1987. russell (1997) presented results from a study conducted from 1987 to 1989, and weeks et al. (1996) did extensive surveys for the sabie river. however, a structured monitoring programme was only initiated from the early 1990s when a full-time river scientist was employed in the knp. fish are considered suitable indicators for monitoring the aquatic system because as consumers they integrate the effects of detrimental environmental changes, being relatively high up in the aquatic food chain (hellawell 1978). fish have been used as indicators of environmental conditions on the basis of individuals, populations and indicator species. motivation for the monitoring of fish assemblages in the knp rivers stems from concerns regarding the effects that catchment development have on the quantity and quality of water affecting the integrity of aquatic biota. it was envisaged that a monitoring programme could provide a means of detecting incipient change in the diversity of fish assemblages. this would act as an early warning system of deterioration in the aquatic environment, and signify a need for management response. the first fish-related tpcs were developed during the knprrp (see rogers & bestbier 1997). however, after a phd study on monitoring and fish assemblages in knp (russell 1997), and a water research commission-associated workshop (fish tpcs january 2000), a decision was made to concentrate on the fish assemblage integrity index (faii) (see kleynhans 1999). the faii embraces sensitivity, frequency of occurrence and percentage anomalies, and has resulted in the upgraded tpcs of identified criteria for fish in the knp (see table 4). table 4: the developed and more focused fish thresholds of potential concern (tpcs) for the kruger national park rivers. application, refinements and assessment post-knprrp the fish rapid assessment index (frai) replaced the faii and formed the core of the fish tpc process (see kleynhans 2007). the frai is based on the categorisation of the fish community according to an intolerance rating. this includes trophic preference and specialisation, and requirement for flowing water during different life-stages and habitats with unmodified water quality. results from the frai are expressed numerically and as a ratio of observed conditions versus conditions that would have been expected in the absence of human impacts. the numerical scores indicate the direction of change when measured against an agreed upon, desired ecological category. there has been ongoing assessment of the fish monitoring data using the frai, however, very few outcomes had been evaluated successfully within a tpc framework. the main reason was that outputs from the frai model are not refined enough to differentiate between habitat, river flows and water quality influences that were impacting fish. this is exacerbated by river systems being highly complex. thus, implementing pertinent management response to the frai model outputs has proved difficult. although the rhp can provide the necessary tools for development of fish tpcs, the following issues have been raised (tpc workshop, skukuza, 2007), (1) the rhp does not provide sufficient insight into cause-and-effect relationships, (2) species sensitivity to influences should be unpacked and then (3) tpcs for all rivers should be developed using these findings. furthermore, ‘ red-flagging’ (tpc tabling) too many concerns would cause the tpc process to lose impetus, that is, an overload of monitoring, reporting and managing would be impractical. hence, a new suite of detailed fish tpcs was generated, based on methods developed during the ecological reserve determination studies for the letaba river (see dwaf 2006). it is important to note that specific fish species are chosen as indicators of change in the main drivers in the river system, for example, river flow and/or water quality, and should act as an early warning system when conditions begin to deteriorate. testing of this approach was completed in 2008 using sites along the olifants river (mcloughlin, deacon & sithole 2009a). this effort led to the first official tabling of a fish tpc within the knp’s sam system (mcloughlin et al. 2009a). it is notable that there are still gaps and shortcomings in the implementation of the knp fish tpc system. this is, however, being addressed currently. tpcs using macro-invertebrate indicators rationale gold mining along the upper sabie river in the 1920s prompted the first environmental study (1933) in the knp, related to invertebrates (moore & chutter 1988). the study indicated that the river was impacted severely by the discharge of waste into the river, with virtually no small aquatic life present. the second study investigated the diversity of benthic invertebrates, and was completed along the crocodile and sabie rivers in 1959 by the national institute for water research. new species were discovered comprising the ephemeroptera and trichoptera families, with the sabie river having more benthic invertebrate diversity than the crocodile river (moore & chutter 1988). a third study was conducted along the crocodile, sabie, olifants, letaba, luvuvhu and motale rivers in 1985/1986 because of concerns about mining impacts (moore & chutter 1988). the main objectives were to make monitoring recommendations for conservation purposes, based on the presence of benthic invertebrates. the sabie and luvuvhu rivers had the lowest chemical concentrations; sabie river had the highest invertebrate taxa diversity, followed by luvuvhu, mutale, crocodile, olifants and letaba rivers. moore and chutter (1988) recommended the implementation of a benthic invertebrate monitoring programme in the knp rivers to provide warnings of insidious changes in the benthic invertebrates, therefore of a broader river health. concerted macro-invertebrate-related monitoring along the five major rivers of the knp commenced in 1996 with initiation of the rhp, using the south african scoring system (sass) methods. sass is a standardised technique for monitoring macro-invertebrates, using a rapid bio-assessment method where macro-invertebrates are sampled to family level from three biotopes (i.e. stones, vegetation and gravel/sand/mud) (dickens & graham 2002; gerber & gabriel 2002). the ecological status at sites is determined by analysing sass scores and the number of taxa found (dickens & graham 2002). during the knprrp, tpcs for macro-invertebrates were developed, based on the abundance of macro-invertebrate taxa and habitat condition, to be sampled within four biotopes (mud/gravel, stones-in-current, stones-out-of-current, and marginal vegetation) (see rogers & bestbier 1997). a bi-annual monitoring programme was envisaged (march and august) at five sites within the knp’s boundary, in each perennial river. the tpc would be exceeded when the number of macro-invertebrate taxa declined by more than 50% compared to previous records, indicating potential deterioration in conditions (dickens & graham 2002; gerber & gabriel 2002; rogers & bestbier 1997). another tpc was associated with habitat loss, to be audited using the sass associated habitat index sampling method (see rogers & bestbier 1997). a loss of 20% or more would act as an early warning flag of detrimental physical and/or chemical impacts on macro-invertebrates. application, refinements and assessment post-knprrp although the five major rivers of the knp have been monitored every 2 to 3 years under the rhp since 1996, feeding of this information through the macro-invertebrate tpc framework was not implemented until 2008. perceptions about adequacy of sampling events achieved, and therefore ability to monitor trends, may have been responsible. since the inception of the rhp, the olifants river has been sampled four times (1998, 2002, 2008 and 2009), the letaba, luvuvhu and crocodile rivers twice (2007/2009, 2008/2009, 2005/2009 respectively) and the sabie river once in 2009. however, in 2008, a decision was made to investigate application of available data. data from moore and chutter (1988) were used as reference to compare data and assess the tpc for the olifants river in july 2008. the tpc was exceeded (and tabled) because three of five sites had 50% fewer taxa than in the mid-1980s (mcloughlin et al. 2009a). at the march 2009 knp conservation management committee meeting it was decided that a more intensive monitoring effort should be conducted, acting as an initial feedback loop. results indicated that those sites that had exceeded the tpc in 2008 had recovered by june 2009. higher river flows than those in 2008 may therefore have helped, and subsequently the tpc has been removed from the list of un-closed tpcs. at present, there is a requirement to incorporate all sass-derived monitoring information into the macro-invertebrate response assessment index (mirai) model, developed for south african rivers (see kleynhans et al. 2005). mirai was developed to determine the ecological category (class) of rivers, using macro-invertebrates as indicators of change in relevant environmental variables. mirai compares the present conditions of rivers (water flow/velocity, habitat, water quality) and macro-invertebrate presence (taxa diversity and abundance) to reference conditions (kleynhans et al. 2005), aiming to identify causes (drivers) of change. for these reasons, new tpcs will be derived that are associated with macro-invertebrates and the mirai model. to date, there has been an inadequate capacity within the knp to implement mirai modelling, and to derive relevant tpcs. this is, however, being addressed in 2010 via project collaboration with relevant macro-invertebrate specialists. riparian tpcs tpcs for sedimentation and habitat loss rationale: several of the larger rivers in the knp are characterised by an unusual mix of both bedrock (underlying geological) and sandy sedimentary influence within the channel. the variable mix of sandy and underlying rocky areas creates a high degree of morphological (‘habitat’) complexity, ranging from slow-flowing, very sandy areas to very fast bedrock rapids, creating many niches for biota (rountree, mcloughlin & deacon 2008). for example, the rivers of the knp have the highest fish species diversities in south africa; many species are critically dependent on the fast-flowing, well-oxygenated, in-stream bedrock habitats (rountree, mcloughlin & deacon 2008). research has shown that the crocodile, sabie, olifants and letaba rivers within the knp fluctuate in space and time between various rocky and sandy conditions (carter & rogers 1995; rountree, rogers & heritage 2000; rountree & rogers 2004). there has, however, never been a time when there was no bedrock influence in these rivers. under natural conditions, there is a tendency for some areas to become rockier whilst, simultaneously, other sections of the river become sandier (rountree, mcloughlin & deacon 2008). land-use changes within the upstream catchment (outside the knp) have resulted in reduced flows, reduced floods and increased sediment inputs. these are thought to be causing a reduction in the availability of bedrock habitat because of the increased sediment storage. this results from increased sediment inputs and decreased sediment transport because of the reduced flows (rountree, mcloughlin & deacon 2008). the original geomorphological indicators were generated along the sabie river to measure and monitor the structural components of biodiversity (rogers & bestbier 1997). here, ecosystem response to changes in flow regime and increased sediment storage were defined according to five channel types (out of nine), characterised by bedrock influence and susceptibility to sedimentation (table 5). it was envisaged that these channel types would be monitored by mapping changes observed on aerial photos, at various time intervals and after events such as droughts and floods of particular magnitudes (rogers & biggs 1999; rogers & o’ keeffe 2003) (see table 5). changes in the channel characteristics that were used to set the lower limits of the geomorphology tpcs included (see table 5), (1) directional loss in bedrock area at the whole river scale within the knp and (2) the proportion of bedrock influenced geomorphic features at the finer geomorphic unit scale (rogers & biggs 1999; rogers and o’keeffe 2003). table 5: the original geomorphology thresholds of potential concern proposed for the sabie river, kruger national park, monitoring for the structural biodiversity components. application, refinements and assessment post-knprrp: the geomorphology monitoring (described above) was tested within the knp sam system during 2007. although based on extensive research, the subjectivity of measuring various geomorphic units along representative reaches, and expert input required, was a major problem and hence was unsustainable (mcloughlin et al. 2009b). an early feedback loop within a sam system should ask the question whether the monitoring programme is feasible. if not, this must be reviewed and changed where appropriate. in addition, new thinking suggested that the geomorphology tpc was inadequate because it assumed very gradual, incremental change in the morphology of the river channel over time, which was not suitable for flagging problems before sedimentation problems arose (rountree et al. 2008). hence, using the best available knowledge, a new approach was initiated in 2008 for ‘red-flagging’ potential sedimentation problems, or for providing the relevant ecosystem information if rehabilitation is required. this was based on analysis using historical aerial photographs, involving a count of all rapid sections along the length of a bedrock-influenced river. trends would be established relative to the 1940s where conditions were assumed more pristine, and a tpc determined, based on the rate of rapid section (critical habitat) loss over time (rountree, mcloughlin & deacon 2008). it was soon realised, however, that a simple count of rapids was inadequate; more information was required to assess the condition of the channel in terms of degradation by sedimentation. thus, the geomorphic tpc was revised to monitor only in the most sensitive sites along the river where increases in sedimentation could be flagged the fastest (rountree, mcloughlin & deacon 2008). the process now includes (1) assessing change in the size (width) of the active channel (in-stream habitat availability) and (2) functioning of exposed bedrock habitats important for biota. in 2008, the tpc was tested successfully along the olifants river. pertinent sediment-sensitive sites were demarcated and geo-referenced using a geographic information system (example figure 3 [a]). mean width of the active channel is measured using pre-selected transects across the mapped active channels (figure 3 [a]). a geomorphic scoring system (developed using expert knowledge) (see figure 3 [b]) is used to calculate a rocky habitat index (rhi) in selected areas of the river (polygons, figure 3 [a]). a once-off historical analysis of trends is carried out to set first estimates for tpcs associated with active channel width (see figure 4 [a]) and the rhi (linked to an ecological category) (figure 4 [b]) that can then be monitored against, in future, to audit the tpcs (rountree, mcloughlin & deacon 2008). trends emanating from the interpretation of this monitoring will provide the broader-scale context involving geomorphic changes occurring within the rivers. additionally, pertinent decision support protocols, linked to this tpc system, are under development. figure 3: methodology for the new geomorphology thresholds of potential concern concept. (a) example of a site selection and demarcation, (b) rocky habitat index scoring system, as tested on the olifants river. figure 4: development of the new geomorphology thresholds of potential concern. tpcs for biotic response to sedimentation and habitat loss rationale the riparian tree species breonadia salicina was recognised as a good indictor of bedrock influence because it grows in close proximity to the active channel and in close association with bedrock-related habitats (van coller 1993). additionally, b. salicina germinates abundantly on any substrate type, however, it only establishes on exposed bedrock because this substrate allows for sufficient anchorage during flooding events, which is important for individual persistence. hence, the population structure exhibited by b. salicina will be affected detrimentally by the smothering of bedrock via increased sedimentation during the establishment phase of its life cycle, with adult populations progressively declining in abundance (mackenzie, van coller & rogers 1999). importantly, current knowledge suggests that for healthy b. salicina populations, the size-structure should display a negative j-shaped curve on bedrock-influenced, geomorphic habitats (figure 5). three parameters that denote characteristics of population structure are being used to assess the breonadia tpc (mackenzie, van coller & rogers 1999), (1) the goodness of fit (r2) of a linear regression of population structure, (2) the slope of the linear regression (x-coefficient) and (3) the y-intercept (constant) of the linear regression. mackenzie, van coller & rogers (1999) explain the biological interpretations for the initial parameter values set, and provide details of a population model (for predictive purposes) and monitoring requirements linked to this tpc. figure 5: the negative j-shaped population structure for all non-germinant, established individuals of breonadia salicina, used for auditing of thresholds of potential concern. application, refinements and assessment post-knprrp the first non-research-related monitoring effort associated with the breonadia tpc was completed only in august 2008. therefore, since the tpcs’ inception, major assessment opportunities have been lost. there was, however, a large, infrequent flood disturbance in the sabie river during february 2000, impacting the b. salicina populations. this resulted in further interest for post-graduate work. dowson (2009) refuted assumptions by mackenzie, van coller & rogers (1999) that the b. salicina populations would recover within 4 years after such a large flood event. this provided important information for further use of this tree as an indictor of bedrock influence, linked to biodiversity goals. pertinent refinements to the breonadia tpc were made, including an 8-year predicted population recovery period after a large flood event. in addition, although the pool-rapid channel type is most sensitive to increasing sedimentation, and was targeted for monitoring b. salicina (mackenzie, van coller & rogers 1999), new knowledge suggested a need to move the monitoring sites to the mixed anastomosing (see van niekerk et al. 1995) channel type. reasons included (1) there is a naturally occurring low abundance of b. salicina in the pool-rapid channel type and (2) the pool-rapid channel type undergoes more frequent state change (natural) between the sediment-dominated (braided) and bedrock-dominated (pool-rapid) states (rountree pers. com.). it is, therefore, difficult to tease out natural from unnatural sediment-related dynamics. the first tpc audit associated with the b. salicina indicator was completed in 2008 after the monitoring effort in august of that year, resulting in the tpc being tabled at the knp conservation management committee meeting of may 2009 (see appendix 1 for structure and content of the feedback report). the response was to resample the b. salicina populations in 2009, to further investigate population recovery after the 2000 flood (initial feedback loop). although the tpcs were still exceeded, there was evidence to suggest population recovery, that is, recruitment and seedling establishment (absent during 2008). it is important to note that there is a requirement, currently, to reassess the tpc parameters, because initial values were estimates based on available knowledge (mackenzie, van coller & rogers 1999) and monitoring sites were changed to the mixed anastomosing channel type. the re-assessment of threshold parameters are being tested under a current water research commission project titled ‘application and testing of a strategic adaptive management system for freshwater protection’ (wrc project number k5/1797), using all monitoring data collected to date. learning associated with the breonadia tpc has been largely biophysical in nature. for example, b. salicina populations require more than 10 years for recovery after a large flood disturbance (mcloughlin et al. 2009b). this refutes the 8-year prediction made (dowson 2009) subsequent to the initial estimate of 4 years by mackenzie, van coller & rogers (1999). furthermore, the breonadia tpc example highlights the need to select the most appropriate sites for monitoring the tpc indicators (mcloughlin et al. 2009b). tpcs for biotic response to changes in flooding regime rationale although terrestrial woody species do occur within the riparian zone, under natural conditions, there is a decreasing abundance closer to the active channel. an increasing terrestrial species abundance in riparian zones threatens biodiversity (mackenzie, jacoby & rogers 2003). the reasons for this increase include (mackenzie et al. 2003) (1) the loss of riparian species while terrestrial species abundance remains unaltered, because of a reduced recruitment of riparian individuals with less flooding, or loss of established riparian plants from drought, (2) an increase in terrestrial species while the riparian species abundance stays the same, because of reduced flooding frequency – terrestrial species usually not tolerant of flooding conditions may now establish closer to the active channel and (3) a decrease in riparian species abundance while terrestrial species numbers increase, because of severe reductions in flow over time, resulting from upstream abstractions and barriers to flow. the current tpc for terrestrialisation is framed as the ratio of the abundance of key terrestrial woody species, to the abundance of an equal number of key riparian species, along an index of flow frequency and availability of water from the water table (see mackenzie et al. 2003). four woody species used to assess the terrestrialisation tpc are combretum erythrophyllum and syzygium guineense (riparian); dichrostachys cinerea and acacia nigrescens (terrestrial). application, refinements and assessment post-knprrp the first non-research-related monitoring effort for the terrestrialisation tpc was completed in 2008, within four sites along the sabie river and follow-up monitoring took place in august 2009 (mcloughlin et al. 2009b). sites from knprrp (see mackenzie et al. 2003) were extended to 100 m (from 20 m)-wide belt transects to sample the relevant woody species adequately. the first tpc audit associated with the terrestrialisation tpc was achieved in 2008 and later in 2009, with no tpc exceedances being recorded. one site was, however, considered inadequate for monitoring and required replacing. an assessment about the future use of the terrestrialisation tpc in its present form is currently being carried out. newer techniques associated with riparian vegetation condition have been developed by specialists working for the dwa and can potentially be used (mcloughlin & mackenzie 2009). ultimately, it is envisaged that these methods will be simpler to implement, but will still provide necessary information within the tpc framework. lessons learned (back to top) the implementation of river tpcs within the knp sam system since the knprrp, and changes instigated to date, provide key lessons about the practical application of tpcs within sam, including the use of adaptive feedbacks loops that are important for evaluation and learning. it is envisaged that these ‘lessons learned’ may be beneficial to other organisations that seek to adopt sam. one example within the south african context is the catchment management agency. these institutions are required, under the national water act (36 of 1998), to practise de-centralised, stakeholder-engaged, adaptive water resource management to meet sustainability, equity and efficiency objectives. some of the knp river lessons include the following: people: motivated, dedicated and adaptable the success of the newly implemented ‘rapid response system’ within the knp (with integration into the broader catchment situations) is attributed to the dedication of the knp river technician. this person is essential as a catalyst for various feedbacks associated with the different ‘worry levels’ mentioned above. in addition, this person is critical in collecting, collating and distributing the information each month, so that relevant people can make use of it in decision making. the ‘management log’ is also updated monthly by this person; this provides critical information for evaluating effectiveness of the system. without such a person, it is doubtful whether this information would be made available on an ongoing basis, at the correct times and to the right people (without the luxury of an expensive, automated data system). the sam tpc system is highly dependent on generation of suitable data and information that is disseminated and actually used. therefore, employing motivated and dedicated people of this type is an important enabling factor, critical at the monitoring, information generation and dissemination stages of sam, for later evaluation and reflection processes. at the reflection and decision-making stages, motivated and ‘adaptive thinking’ individuals are also required to disseminate information in appropriate ways, so that suitable discussion on issues arising from this information is facilitated. suitable benchmarks against which to measure indicators the importance of a benchmark against which to monitor river flows is a critical enabling factor for adaptive feedback processes, as applied to river flow management (pollard & du toit 2009). with the implementation of a ‘rapid response system’ within the knp, defining which benchmarks to use that are suitable must also be emphasised. the crocodile river provides a good example, as the appropriateness of the ifrs for the river that were developed over 10 years ago is proving to be a major stumbling block. this is further exacerbated by the deteriorating river flows in the crocodile river. broader recognition of these ifrs as being outdated, exacerbated by lags in implementation of the newer, improved ecological reserve determination methods, has caused some confusion about further implementation of the knp ‘rapid response system’ system for this river – for example, when to initiate various responses from the knp to outside water resource managers and knowing when to make legitimate calls for improvements in river flow, rather than merely antagonising other stakeholders. this situation leads potentially to a loss of cooperative behaviour and trust between stakeholders (e.g. knp to catchment managers and other water users) and is exacerbated when there is disagreement between stakeholders on the merits of an existing benchmark, leading to frustrations about when actions must occur. a major risk is loss of motivation by key people (as described above), and a subsequent loss in momentum of the adaptive feedback processes. therefore, determining suitable and acceptable benchmarks against which to monitor system change, and acting at the appropriate times relative to these, must be encouraged and actively sought after. in addition, the acceptance and buy-in from all key stakeholders is important. information: generation and use adaptive management implies the production of knowledge through policy and management actions, and that this knowledge is disseminated among all interested groups, and is actually used (stankey, clark & bormann 2005). in this context, knowledge is continually being constructed and reconstructed as various people interact with each other and as new information becomes available. learning from the generation of new knowledge is, however, a complex process, and learning can be derived from both technical and social processes (stankey et al. 2005). on one hand, an increased understanding about systems, such as the socio-ecological and socio-economic systems and their interactions, is a form of technical knowledge. however, enhancing the social processes associated with how this new understanding may be better formulated, communicated and incorporated into organisational policies and management, is critical. although deficient learning can result from a lack of data and resultant information, in other instances the information is available, but not made accessible, or is presented in a manner not conducive to learning. it is important to note that the social processes needed to facilitate communication and debate among an organisation’s members may be lacking. here, feedback of information is essential for enhancing information flow within an adaptive management context (stankey et al. 2005).the issues highlighted by stankey, clark & bormann (2005) are also applicable to the implementation of the knp river sam system over the last decade. firstly, within sam, monitoring programmes where information is derived must be developed and prioritised according to the objectives hierarchy. this allows for explicit linkages between the higher-level, value-laden objectives and those on the ground, that is, the tpcs (see section on ‘knp river tpcs – future directions’, refinements to the river objectives hierarchy). generation of information, via implementation of these strategic monitoring programmes, then plays a critical role for effective evaluation and reflection. a lack of monitoring (prior to 2008) associated with the geomorphology, breonadia and terrestrialisation tpcs within the knp sam system resulted in no information having been derived to make assessments, thus opportunities for evaluation and reflection were lost. learning only occurred after 2008 once the first monitoring and information feedbacks were instigated, as discussed in this paper. secondly, once derived, information must then be made available to all pertinent people within an organisation, otherwise scarce resources spent on information collection are wasted. although monitoring associated with the in-stream tpcs (fish and macro-invertebrates) within the knp was carried out over the past decade, this information was not made available in an adequate form to decision makers. thus, managers had very little idea about the condition of the river ecosystems in terms of these indicators, limiting the urgency of responses that were made. in 2008, reporting and tabling of the fish and macro-invertebrate tpcs for the olifants river, using available historical information generated via monitoring, did increase awareness about the extent of deterioration of the olifants river over time. this set in motion the initial adaptive loops, that is, the instigation of more detailed monitoring to further investigate conditions within the river, for subsequent action. it is important to note that, because this information was not made wholly available in the past, earlier management opportunities resulting from this information were potentially lost. although stoppages in river flow occurred in 2005, it is notable that more concerted action commenced in 2008 with the onset of a crisis: the nile crocodile deaths in the olifants river gorge (mcloughlin, deacon & sithole 2009a). thirdly, within sam, pertinent information must actually be used within the decision-making process, relative to management objectives, so that adaptation of actions can be made where appropriate. importantly, the amount, detail and format of the information must be applicable at different organisational levels. for example, scientists within the knp may deal with the detailed data and information associated with biophysical conditions in the river (e.g. for the breonadia tpc), managers may require only broader information on this, but a clear indication of trends related to drivers, like river flows where management action can be applied. allowing for better visualisation of information, as occurs for the river flow data within the knp ‘rapid response system’, facilitates information transfer and enhances the assessment and evaluation environment for decision making. however, improving this feedback system is an ongoing process as further learning occurs. implementation lags: key processes within the sam tpc system, lags in implementation of various components (e.g. monitoring) should be expected. the key issue, then, is how to define and deal with these. important questions asked concerning the deteriorating knp river flow situation are: which lags are acceptable and which are not? importantly, contingency plans should be put in place to deal with unacceptable lags. the crocodile river situation, with lags in development of methods to calculate the new ecological reserve benchmarks provides a good example of this. the contingency in this case was implementation of the three-tiered management response system for the crocodile river (see figure 2), in relation to the older ifr benchmark levels, until availability of the newer ecological reserve benchmarks (see section on ‘knp river tpcs – future directions’). another example of a lag within the knp sam system is a delay in the implementation of the intended evaluation processes associated with the ‘rapid response system’. specifically, there should be discussion within the knp conservation management committee on how the ‘rapid response system’ has worked over the previous period, that is, what the associated river flow outcomes were, which actions worked or did not work, and how to adapt and improve the system for future application, to improve river flow conditions ultimately. an important lesson is to follow through on plans; although there may be general agreement on particular systems, this does not automatically translate into implementation of adaptive processes. implementation requires dedicated, motivated and adaptive people (champions), to move the processes forward. without such people, the adaptive feedback process is likely to stagnate, and future management adaptation, using past actions to learn from, will not be forthcoming. timing: between monitoring and management action the timing of feedback processes within an organisation, that is, from data and information collection to decisions made using this information, must be considered when implementing tpcs within sam, to increase strategic use of information. the ‘rapid response system’ within the knp was developed to deal with quick management responses that were required, associated with an indicator (in this case river flow that can change within short time periods). within the knp, the assessment and reflection opportunity once every two months, using the tpc tabling system, was not appropriate for this indicator. adapting past management responses, from within the knp to achieve the desired state, that is, compliance with ifrs, was therefore difficult. this situation generated frustration within the knp sam system, while ongoing deterioration of river flow conditions occurred within rivers, because actions could not be adequately assessed and adapted to improve future implementation of the ‘rapid response system’. essentially in practice, feedbacks within sam require different time frames, dependent on the type of drivers, and indicators used within the tpc framework, these issues must be considered. feasibility: monitoring programmes ongoing assessment about the adequacy of monitoring programmes, as used within the sam tpc framework, is important. feasibility of monitoring methods to assess tpcs, in terms of cost or expertise required for example, should be considered in detail before this is included within an organisation’s management plans. unfeasible monitoring programmes can lead to lags in implementation of the monitoring required to assess components of a tpc system. the result is a lack of information that should have been made available for decision making, to reflect on the effectiveness of the selected and implemented management actions. adaptation of future management actions, based on information about past management actions is then hampered. a good example is the geomorphology tpcs that were originally developed during the knprrp of the 1990s. although based on sound research and principles, the expertise required to implement these tpcs over the longer term may have resulted in no monitoring taking place over a decade since the tpcs’ inception. a simpler system was necessitated, with less expert input required, but that could still provide relevant information about the geomorphic state of the rivers. this has been achieved with the new geomorphology tpcs that have been developed recently for the knp rivers. refinements: ongoing tpc requirement tpc development within sam is not a once-off process, but rather, ongoing assessment and refinement over time is needed. because of the large amounts of uncertainty and lack of complete knowledge about systems, the development of tpcs often commences with basic first estimates involving the relevant indicators of change. as new information is generated, via monitoring efforts or further research, feedbacks must then focus on evaluating the appropriateness of the tpcs. using current knowledge, thresholds must then be refined to improve the tpc over time. ideally, ongoing learning about the system under consideration will then also occur. the breonadia tpc development and refinement process within the knp demonstrates this point well. a lesson is that changing management actions, as a consequence of new information generated via monitoring efforts, is not always an outcome in sam. rather, learning processes occur within the tpc process itself, that is, increased understanding about drivers of change, indictors of these drivers, and pertinent threshold parameters to set and apply within the tpc framework. adaptive feedback loops: identification and closing over more than a decade, one major focus of the knp sam system associated with river management has been on implementation of the ‘operational’ components, which are the monitoring and reporting related to river flows (ifrs) and water quality. although very important, under this situation there has been some loss of recognition for why these operational components were originally conceptualised under the national water act (related to water resource sustainability and the delivery of ecosystem goods and services, for human benefit). in the knp context, these benefits are associated with sanparks’ mandate to maintain biodiversity, with links to eco-tourism outcomes, among others. this provides an important adaptive management lesson, in the south african context, particularly for catchment management agencies that are tasked with giving effect to the national water act, and in this case particularly freshwater protection. implementing the ecological reserve in an adaptive manner requires furthering understanding about the effects of the ecological reserve on the biophysical components of the system, ultimately linked to maintaining the delivery of ecosystem goods and services, for human benefit. associated information must be fed back richly into learning, ultimately, to improve water resource management over the longer term. in essence, achieving agreed upon sustainability goals, over and above other (e.g. social and economic) objectives. the main lesson here is identifying and then actively attempting to close adaptive cycles within the sam framework, along with explicit learning. notably, various stakeholder sectors within any catchment context will have various priorities and perceptions associated with where ‘closing of the adaptive cycle’ occurs. there are potentially many nested adaptive cycles occurring simultaneously within a broader sam system. for instance, there has been a major focus at the catchment scale within south africa on the implementation of various strategies to allocate water adequately to different water use sectors, while meeting ecological reserve targets at strategic points along the river. refining and improving practices to achieve this is a key factor, involving learning about how to adapt (e.g. dam operating rules, validating water use and re-licensing, regulating use via water restrictions and so on) along the way to achieve this desirable outcome. however, in order to meet the requirements of the national water act, there is an obligation over longer time scales for directed learning within sam, that is, about how implementation of the recommended flow regimes (and water quality) generates the desired state (vision) of the resource under management. for example, achieving or maintaining the recommended management class for sustainability, or the equity/economic-related targets. it is this process of learning and subsequent adapting to meet these types of highlevel objectives that will result in a more active application of sam, as applied within the context of the south african national water act. kruger national park river tpcs – future directions (back to top) the river tpcs and associated monitoring plans have the longest history within the knp, since the inception of sam within the knp during the knprrp of the 1990s (see rogers & bestbier 1997). however, a lack of full implementation after a decade has now necessitated a broader re-evaluation. revision of rationale and a reflection on objectives and best available indicators and monitoring methods, in line with management goals, is currently underway. this reflection process is not within the scope of this paper; however, an overview of pertinent initiatives currently taking place is provided below. firstly, an assessment of the knp objectives hierarchy has identified the following potential problem areas related to the strategic objectives for river management, (1) too many levels associated with the hierarchy of objectives, generating undue complexity, (2) the proportion of value-laden and research objectives is too high, compared to the finer level of detail where ‘doing something’ is required, (3) a need to link the higher level, value-laden objectives higher up with more rigorous and explicit end-points (tpcs), thus to prioritise monitoring activities to guide management in the achievement of goals, this being a key component of sam and (4) no link to socio-ecological processes and understanding required, this now being an important focus for integrated catchment management and adoption of sam wider in the catchments. hence, a modified objectives hierarchy for river management has been recommended, (see figure 6), but still requires further reflection during the knp sam revision process. these changes include the following, (1) the functional processes and composition/ structure and pattern objectives are not explicitly stated as before, but integrated within the biodiversity-related objectives, for clearer interpretation, (2) the research objectives have been grouped separately, and broken down into relevant sub-objectives related to both biophysical and governance (social) aspects for more clarity on research needs (with necessary linkages), (3) research objectives, related to generating further understanding and maintenance of tpcs has been made more explicit, (4) there are more explicit links between higher level aquatic biodiversity objectives and the more detailed end-points (tpcs) to guide relevant monitoring in relation to goals, (5) tpcs are now explicitly determined per river, an important aspect because of differences between river characteristics and given knowledge per river system and (6) the ‘social understanding objective ’ has been incorporated, requiring linkages with the knp social science objectives hierarchy where appropriate (for further development). figure 6: recommended re-structuring related to the river-associated objectives. secondly, important to consolidation of the knp river tpc system is the recognition of and integration with relevant national processes associated with fostering sustainable water resources. specifically, the ecological reserve process, including implementation of environmental flows and monitoring initiatives where pertinent. the ecological reserve the current ecological reserve determination processes provide a more realistic estimate of environmental flow requirements compared to the ifr-based system, and these requirements are legally justifiable under the national water act (once signed off by the minister of water affairs). the ecological reserve (incorporating water quality also, but only referring to river flows in this instance) has been determined for the crocodile, sabie, olifants and letaba rivers, albeit at varying levels of detail and confidence. further studies are in progress (or are planned) to update older versions (see table 6). to date, effective implementation of this ecological reserve has, however, not taken place, a consequence of various catchment wide lags in implementing the national water act. a key factor is that most catchments associated with the knp are highly water-stressed (the sabie river is an exception) (see dwaf 2004a; dwaf 2004b; dwaf 2004c). it is believed that rapid implementation of the ecological reserve is dependent on significantly increased water restrictions on users, resulting in large economic impacts in the surrounding catchments, for example, on irrigation farming in the crocodile river catchment (jackson, pers. com.). table 6: status of the ecological reserve studies for major rivers of the kruger national park. hence, a longer-term, progressive realisation of ecological reserve implementation is envisaged for most major rivers of the knp, but implementing strategies to achieve this is now critical. the knp is playing an increasingly active role, for example, in the development of the inkomati catchment management strategy during march 2010 (as required by the water affairs ministry), after a series of public participation workshops run by the inkomati catchment management agency. although not discussed further in this paper, the authors acknowledge that water quality monitoring and management does require further improvement. the practicality of water resource managers delivering the ecological reserve has, however, also contributed to lags in implementation. it is encouraging that, to date, improved hydrological models and methodologies to implement the ecological reserve have been developed and tested, such as the spatial and time series information modelling (spatsim) software (see hughes, mallory & louw 2008) for the groot letaba river system and the mike flood watch and water resource modelling platform (wremp) (see dwaf, 2008b) for the crocodile east river system. the idea is to mimic natural flow variability as much as possible. dam releases and/or water restrictions will ideally be calculated and applied to meet both the ecological reserve requirements (progressively realised over time where pertinent) with adequate water supply to the various water user sectors. a requirement will be that these sectors need to become more efficient in their water use. the knp is collaborating with catchment management agencies on various implementation strategies associated with these developments. monitoring river flows associated with an adaptive feedback system is one important initiative being implemented, involving the integration of knp river sam processes wider into the catchment areas. aquatic and riparian ecosystem monitoring the knp is consolidating existing knowledge, using updated understanding about the physical drivers (water quality, flow and sediment load) from the upstream catchments that influence the biophysical conditions (at different temporal and spatial scales) of the rivers flowing through the knp (see rountree et al. 2008; mcloughlin et al. 2009b for details). in addition, integration of the knp’s tpc monitoring programmes with ecological reserve monitoring processes is being sought where appropriate. for example, kleynhans et al. (2009) propose integration of ecological reserve monitoring with that of the river health programme, forming the basis of an integrated ecological water resource monitoring system. it is envisaged that this should be undertaken within a structured decision-making framework, following the principles of sam (dwa 2009) (see figure 7). the main purpose is to provide a decision framework within which monitoring results can be interpreted in terms of the attainment of specified objectives, that is, ecological specifications (ecospecs) and tpcs linked to an ecological category. the ecological category is associated with river health conditions linked to a vision set for a particular river. the ecological water resource monitoring system uses the newly developed and available monitoring techniques called rapid habitat assessment methods (see dwa 2009). rapid habitat assessment methods were developed because capacity within south africa is a major stumbling block to longer-term implementation of the ecological reserve process. essentially, before biotic indicators are monitored (using river health programme tools: the frai, the mirai), data are collected using the more feasible rapid habitat assessment methods, and then analysed. the premise is that suitable habitat conditions will indicate the likely presence, abundance and frequency of occurrence of particular biota. the associated tpc system will act as a ‘red flag’ prior to (potential) requirements of the more resource-intensive biotic methods (see figure 7). although originated as a tool for in-stream conditions, application of rapid habitat assessment methods to riparian vegetation has also been developed (mackenzie, pers. com.) and will be used to make assessments associated with the knp terrestrialisation tpc and others. the knp is spearheading initiatives to implement this new monitoring and decision support system (figure 7). within the knp sam system (and wider), this will provide further learning opportunities associated with information generation and storage, interpretation, dissemination and use. figure 7: : envisaged structured decision-making framework to guide in-stream river biophysical monitoring and decision making associated with implementation of the ecological reserve acknowledgements (back to top) we thank llewellyn foxcroft for his critical inputs to this manuscript, and for useful suggestions. funding from the water research commission allowed further investigation into testing and refining the knp river tpcs and associated processes, with the many outcomes incorporated in this paper. we are also grateful for the enthusiastic inputs associated with ongoing knp tpc developments, provided by james mackenzie, mark rountree and jacques venter. in addition, the broader support given by rina grant, stefanie freitag-ronaldson, harry biggs, kevin rogers and freek venter has made this work possible. references (back to top) biggs, h. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59−80, island press, washington. breen, c.m., dent, m.c., jaganyi, j., madikizela, b., maganbeharie, j., ndlovu, a., et al., 2000, ‘the kruger national park rivers research programme’, water research commission report tt 130/00, wrc, pretoria. carter, a.j. & rogers, k.h., 1995, ‘a markovian approach to investigating landscape change in the knp rivers’, report no. 2/95, cwe, university of the witwatersrand, johannesburg. council for scientific and industrial research, 2002, ecological reserve determination for the crocodile river catchment, incomati system, mpumalanga, report no. env-p-c 2002 009, csir, pretoria. department of water affairs and forestry, south africa, 1991, proceedings of a workshop on the flow requirements of kruger national park rivers, report tr 149, department of water affairs and forestry, pretoria. department of water affairs and forestry, south africa, 1996, south african water quality guidelines: aquatic ecosystems, vol. 7, 1st edn., the government printer, pretoria. department of water affairs and forestry, 1998, river health programme, internal report, viewed n.d., from http://www.dwaf.gov.za/iwqs/rhp/state_of_rivers.html department of water affairs and forestry, south africa, 2004a, inkomati water management area: internal strategic perspective – version 1, ed. tlou and matji (pty) ltd. dwaf report no p wma 05/000/00/0303, dwaf, pretoria. department of water affairs and forestry, south africa, 2004b, olifants water management areas internal strategic perspective, ed. gmks tlou and matji and wmb on behalf of the directorate: national water resource planning, dwaf report no p wma 04/000/00/0304, dwarf, pretoria. department of water affairs and forestry, south africa, 2004c, luvuvhu/letaba water management areas internal strategic perspective, ed. goba moahloli keeve steyn (pty) ltd in association with tlou and matji, golder associates and bks on behalf of the directorate: national water resource planning, dwaf report no p wma 02/000/00/0304, dwaf, pretoria. department of water affairs and forestry, south africa, 2006, letaba catchment reserve determination study – main report final, ed. howard and de lange inc., on behalf of the directorate: resource directed measures, dwaf report no rdm/b800/00/con/comp/1304, dwaf, pretoria. department of water affairs and forestry, south africa, 2008a, national aquatic ecosystem health monitoring programme (naehmp): river health programme (rhp) implementation manual. version 2, dwaf, pretoria. department of water affairs and forestry, south africa, 2008b, the development of a real-time decision support system (dss) for the crocodile east river system – inception report, ed. clear pure water with water for africa and dormehl technology, on behalf of the directorate: water resource planning systems, project no wp 9429. department of water affairs, south africa, 2009, operationalise the reserve: rapid habitat assessment model manual, ed. d. louw & c.j. kleyhans, report no. rdm/nat/00/con/0709, dwaf, pretoria. dickens, c.w.s. & graham, p.m., 2002, ‘the south african scouring system (sass) version 5 rapid assessment method for rivers’, african journal of aquatic science, 1−10. dowson, l., 2009, ‘the effects of an extreme flood on breonadia salicina population structure: implications for modelling, monitoring and adaptive river management in the kruger national park’, unpublished m. sc. dissertation, animal, plant and environmental studies, university of the witwatersrand. foxcroft, l.c., 2009, ‘developing thresholds of potential concern for invasive alien species: hypotheses and concepts’, koedoe 51, 1−6. doi:10.4102/koedoe.v51i1.157 gerber, a. & gabriel, m.j.m., 2002, aquatic invertebrates of south african rivers: field guide, 1st edn., resource quality services, department of water affairs and forestry, pretoria. hellawell, j.m., 1978, biological surveillance of rivers, water research centre, stevenage, uk. heritage, g.l., van niekerk, a.w. & moon, b.p., 1999, ‘geomorphology of the sabie river, south africa: an incised bedrock-influenced channel’, in a.j. miller & a. gupta (eds.), varieties of fluvial forms, pp. 53−79, john wiley and sons, chichester, uk. hughes, d.a., mallory, s.j.l. & louw, d., 2008, methods and software for the real-time implementation of the ecological reserve – explanations and user manual, water research commission report no. 1582/1/08, water research commission, pretoria. king, j. & louw, d., 1998, ‘instream flow assessments for regulated rivers in south africa using the building-block methodology’, aquatic ecosystem health and management 1, 109−124. doi:10.1080/14634989808656909, doi:10.1016/s1463-4988(98)00018-9 kleynhans, c.j., 1999, ‘the development of a fish index to assess the biological integrity of south african rivers’, water sa 25, 265−278. kleynhans, c.j., louw, m.d., thirion, c., rossouw, n.j. & rowntree, k., 2005, river ecoclassification: manual for ecostatus determination (version 1), joint water research commission and department of water affairs and forestry report, wrc report no. kv 168/05, wrc, pretoria. kleynhans, c.j., 2007, module d: fish response assessment index in river ecoclassification: manual for ecostatus determination (version 2), joint water research commission and department of water affairs and forestry report, wrc report no. tt 330/08, wrc, pretoria. kleynhans, c.j., louw, m.d., birkhead, a., thirion, c., deacon, a., angliss, m., et al., 2009, ‘on the way to implementation: ecological reserve monitoring’, unpublished report. kruger national park, 1989, ‘kruger national park water provision policy’, unpublished report. kühn, a.l., 1991, ‘sensitiewe visspesie werkswinkel 1991’ [sensitive fish species workshop 1991], rivers research programme report, kruger nationale park. louw, d. & o’keeffe, j., 1999, luvuvhu river ifr: second refinement, institute for water research, grahamstown. place of publisher. mackenzie, j.a., van coller, a.l. & rogers, k.h., 1999, rule based modelling for management of riparian systems, water research commission report no. 813/1/99, wrc, pretoria. mackenzie, j.a., jacoby, d.l. & rogers, k.h., 2003, modelling of terrestrialisation and technology transfer to enable management of kruger national park rivers, water research commission report no. 1063/1/03, wrc, pretoria. mcloughlin, c.a., 2009, status and progress with the framework for prototyping catchment level adaptive management processes within the inkomati, olifants and letaba-luvuvhu water management areas, unpublished water research commission report. mcloughlin, c.a., deacon, a. & sithole, h., 2009a, ‘implementation status of river tpcs developed and tested within the kruger national park, including tpc audits achieved’, unpublished water research commission report. mcloughlin, c.a., mackenzie, j., rountree, m., deacon, a. & sithole, h., 2009b, ‘update on kruger national park river tpcs – roll-out of the geomorphology and in-stream tpc monitoring and data collection associated with the terrestrialisation and breonadia tpcs’, unpublished water research commission report. mcloughlin, c.a. & mackenzie, j., 2010, ‘update on new developments associated with riparian vegetation tpc requirements for the kruger national park’, unpublished water research commission report. moore, c.a. & chutter, f.m., 1988, a survey of the conservation status and benthic biota of the major rivers of the kruger national park, council for scientific and industrial research, national institute for water research, pretoria. o’keeffe, j. & rogers, k.h., 2003, ‘heterogeneity and management of the lowveld rivers’, in j.t.d toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 447− 468, island press, washington. pickett, s.t.a., parker, v.t. & fiedler, p.l., 1992, ‘the new paradigm in ecology: implications for conservation biology above the species level’, in p.l. fiedler & s.k. jain (eds.), conservation biology: the theory and practice of nature conservation, preservation and management, pp. 65–88, chapman and hall, new york. pickett, s.t.a., cadenasso, m.l. & benning, t.l., 2003, ‘biotic and abiotic variability as key determinants of savanna heterogeneity at multiple spatiotemporal scales’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 22–40, island press, washington. pienaar, u., 1961, ‘a supplementary checklist of decapoda, freshwater fish, amphibians, reptiles and and small mammals recorded in the kruger national park’, koedoe 4, 167−177. poff, n.j., allan, j.d., bain, m.b., karr, j.r., prestegaard, k.i., richter, d.b., sparks, r.e. & stromberg, j.c., 1997, ‘ the natural flow regime: a paradigm for river conservation and restoration’, bioscience 47, 769−784. doi:10.2307/1313099 pollard, s. & du toit, d., in review, towards the sustainability of freshwater systems in south africa: an exploration of factors that enable and constrain meeting the ecological reserve within the context of integrated water resource management in the catchments of the lowveld, water research commission report, wrc, pretoria. rogers, k.h., 1997, ‘operationalizing ecology under a new paradigm: an african perspective’, in s.t.a. pickett, r.s. ostfeld, m. shachak & g.e. likens (eds.), the ecological basis of conservation. heterogeneity, ecosystems and biodiversity, pp. 257–270, chapman and hall, new york. rogers, k.h., 2003, ‘adopting a heterogeneity paradigm: implications for management of protected areas’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 41−58, island press, washington. rogers, k.h. & bestbier, r., 1997, development of a protocol for the definition of the desired state of riverine systems in south africa, department of environmental affairs and tourism, pretoria. rogers, k.h. & biggs, h., 1999, ‘integrating indicators, end points and value systems in the strategic management of the rivers of the kruger national park’, freshwater biology 41, 439−451. doi:10.1046/j.1365-2427.1999.00441.x rogers, k.h. & o’keeffe, j., 2003, ‘river heterogeneity: ecosystem structure, function, and management’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 189−218, island press, washington. rountree, m.w., rogers, k.h. & heritage, g.l., 2000, ‘landscape state change in the semi-arid sabie river, kruger national park, in response to flood and drought’, southern african geographical journal 82, 173−181. rountree, m.w. & rogers, k.h., 2004, ‘channel pattern changes in the mixed bedrock/alluvial sabie river, south africa: response to and recovery from large infrequent floods’, in d.g. de jalon & p. vizcaino (eds.), proceedings of the fifth international symposium on ecohydraulics, pp. 318−324, iahr, madrid, spain. rountree, m., mcloughlin, c.a. & deacon, a., 2008, ‘integrating the naehmp monitoring data with the knp strategic adaptive management system: tpc refinement for rivers’, unpublished water research commission report. rountree, m., mcloughlin, c.a., mackenzie, j., deacon, a. & sithole, h., 2008, ‘kruger national park tpc data collection, testing and implementation’, unpublished water research commission report. russell, i.s., 1997, ‘monitoring the conservation status and diversity of fish assemblages in the major rivers of the kruger national park’, unpublished ph.d. thesis, zoology department, university of the witwatersrand. sanders, t.g. & ward, r.c., 1993, ‘water quality monitoring system design, simplified, pollution prevention’, journal of water and wastewater international 8, 44−49. stankey, g.h., clark, r.n. & bormann, b.t., 2005, adaptive management of natural resources: theory, concepts and management institutions, gen. tech. rep. pnw-gtr-654, us department of agriculture, forest service, pacific northwest research station, portland, or. tharme, r., 1997, sabie-sand river system: in-stream flow requirements. proceedings of the ifr workshop, report to the department of water affairs and forestry, southern waters ecological research and consulting cc, freshwater research unit, university of cape town, rondebosch. van coller, a.l., 1993, ‘riparian vegetation of the sabie river: relating spatial distribution patterns to characteristics of the physical environment’, unpublished m.sc. dissertation, centre for water in the environment, university of the witwatersrand. van coller, a.l., rogers, k.h. & heritage, g.l., 1997, ‘linking riparian vegetation types and fluvial geomorphology along the sabie river within the kruger national park, south africa’, african journal of ecology 35, 194−212. doi:10.1111/j.1365-2028.1997.090-89090.x van coller, a.l., rogers, k.h. & heritage, g.l., 2000, ‘riparian vegetation-environment relationships: complimentarity of gradients versus patch hierarchy approaches’, journal of vegetation science 11, 337−350. doi:10.2307/3236626 van niekerk, a.w., heritage, g.l. & moon, b.p., 1995, ‘river classification for management: the geomorphology of the sabie river in the eastern transvaal’, southern african geographical journal 77, 68−76. weeks, d.c., o’keeffe, j.o., fourie, a. & davies, b.r., 1996, the ecological status of the sabie-sand river system. volume 1: a pre-impoundment study of the sabie-sand river system, eastern transvaal with special references to predicted impacts on the kruger national park, water research commission report, wrc, pretoria. appendix 1 (back to top) audit report format used within the thresholds of potential concern (tpc) tabling process within the kruger national park strategic adaptive management (sam) system. example: breonadia tpc exceedances conservation services management committee meeting, kruger national park notification of river tpc reached breonadia salicina population structure in the sabie river, first tpc assessment background the loss of bedrock influence in the kruger national park (knp) river systems is associated with increasing sediment storage – from an increased sediment supply and/or reduced stream-flow ability to transport sediment. the result is a loss of bedrock habitats, because of the smothering of exposed rock by sediment, with a consequent reduction in available regeneration niches for various riparian tree species, leading to a decrease in riparian vegetation diversity. a useful indicator of this loss in bedrock influence is the population structure of the tree species breonadia salicina, because it grows in close proximity to the active channel and in close association with bedrock habitats (van coller 1993). b. salicina is a good indicator of the change in sediment regime and/or flow because although it can germinate abundantly on any substrate type, it establishes only on exposed bedrock; this allows for sufficient anchorage during high flow events, thus increasing chances of individual survival and persistence. importantly, current knowledge suggests that the b. salicina population displays a negative, j-shaped curve on geomorphic units with large amounts of bedrock influence (see figure 1) (de fonteine, unpublished; mackenzie, van coller & rogers 1999). the negatively skewed j-shaped curve represents a population where the abundance of breonadia individuals is highest in the smaller size classes, and diminishes as the size class increases (mackenzie, van coller & rogers 1999). this population structure results if there is (1) sufficient exposed bedrock for establishment of seedlings, including a suitable flow regime, (2) continuous, rather than punctuated, recruitment and (3) an increasing rate of mortality within cohorts because of high disturbance rates. changes to the negative, j-shape population structure exhibited by b. salicina will occur if there is increased sedimentation during the establishment phase of its lifecycle. figure 1: the negative, j-shape population structure for all non-germinant, established individuals of breonadia salicina, used for the auditing of thresholds of potential concern. definition of the threshold of potential concern ultimately, a loss of bedrock influence via sedimentation, and/or a sharp reduction in flow frequency will cause a reduction in seedling recruitment of b. salicina, resulting in the loss of the negative, j-shaped population structure (mackenzie et al. 1999). figure 2 shows in concept the ripple-effect changes that would occur in the population structure, following a marked increase in sediment storage and/or a reduction in flow frequency over time (mackenzie et al. 1999). figure 2: a hypothesis of the change over time in population size structure of non-germinant individuals of breonadia salicina on bedrock-dominated channel sections. an 8% level of bedrock proportion in pool-rapid channel types was considered by mackenzie et al. (1999) as a rough guide to critically low levels of bedrock in the river, and was considered a level of concern. characteristics of the negative, j-shaped curve, fitted to the population structure of all non-germinant size classes of b. salinica monitored in the field, are used for tpc auditing. assessment of the tpc is carried out using the log of this curve, because the log of a negative, jshaped curve outputs a straight line, and pertinent parameters can then be assigned to this line for monitoring change in the population structure. the values given in figure 3 represent the lower limits (thresholds) of the parameters associated with the logged, negative, j-shaped population structure curve of b. salicina (i.e. a straight line), derived from modelled populations where bedrock proportion was at 8% cover. it is important to note that these are first estimates and need refinement over time within the strategic adaptive management (sam) system. figure 3: parameters for the breonadia threshold of potential concern (tpc). data or information supporting the threshold of potential concern the breonadia tpc was developed by mackenzie et al. (1999) during the knp river research programme, using information collected within belt transects demarcated for research along the length of the sabie river. field data were used to estimate levels of concern for the degree of bedrock within critical sites along the sabie river.the current assessment of the breonadia tpc was completed using data sampled during august−september 2008 at dedicated monitoring sites along the sabie river. re-establishment of monitoring sites along the sabie river for the breonadia tpc was carried out as part of the water research commission-funded ‘adaptive management’ project. all individuals recorded were categorised into size classes, and size class frequencies were established in order to assess the tpc parameters associated with the b. salicina sub-populations occurring at each site. these parameters were compared against the tpc thresholds given above. results (back to top) interpretation of the results from the breonadia tpc audits undertaken in 2008 for the three monitoring sites along the sabie river (figure 4) indicated that all the tpc parameters associated with b. salicina population structure within each site have been exceeded. figure 4: breonadia salicina population structure at the three threshold of potential concern (tpc) monitoring sites along the sabie river. consequences or potential impacts a loss of bedrock influence along the riparian corridors of the knp results in diminished habitat diversity within the rivers. consequently, this leads to a decrease in regeneration niches for certain riparian vegetation within the knp, and a loss of riparian vegetation diversity. this is in contradiction to the mandate of the knp – that of maintaining biodiversity. however, the large flood of 2000 impacted greatly on the b. salicina population within the sabie river, causing thresholds to be exceeded naturally. the populations are still recovering, but the question arises as to when the population will recover for re-use within the tpc. understanding prior to the 2000 flood had suggested that the b. salicina sub-populations would recover within 4 years after such a large flood event occurred. however, a research project conducted in 2004 by the centre for water in the environment at the university of the witwatersrand showed that b. salicina sub-populations did not recover as expected. evidence found during this study indicated that the surviving adult b. salicina individuals put more resources into sprouting as a means of regeneration, than flowering and setting of seed. consequently, a smaller seed source results in fewer germinants, and less recruitment into the seedling stage class, with a subsequent loss in the negative, j-shape of the population structure. refined understanding during this research predicted that the b. salicina population would recover 8 years after a large flood event. the b. salicina monitoring data collected in 2008 contradicted this prediction, although there is increasing evidence to suggest that the adults have started to flower and set seed. proposed management action • the next action to be taken should be to re-sample the b. salicina population during 2009 within the monitoring sites demarcated along the sabie river, to investigate the potential rate of population recovery, for example, increased flowering, germination and numbers of seedlings. • further work is being undertaken as part of the water research commission funded ‘adaptive management’ project to test and refine this tpc, including monitoring site selection and the population modelling component. references (back to top) mackenzie, j.a., van coller, a.l. & rogers, k.h., 1999, rule based modelling for management of riparian systems, water research commission report no. 813/1/99, wrc, pretoria. van coller, a.l., 1993, ‘riparian vegetation of the sabie river: relating spatial distribution patterns to the physical environment’, msc thesis, centre for water in the environment, university of the witwatersrand. article information authors: mahlomola e. daemane1 sarel s. cilliers2 hugo bezuidenhout1 affiliations: 1conservation services department, south african national parks, south africa 2school of environmental sciences and development, north-west university, south africa correspondance to: mahlomola daemane email: ernestd@sanparks.org postal address: conservation services department, po box 110040, hadison park, kimberley 8306, south africa keywords braun-blanquet; conservation; degradation; grassland biome; ordination dates: received: 07 may 2009 accepted: 11 dec. 2009 published: 26 apr. 2010 how to cite this article: daemane, m.e., cilliers, s.s. & bezuidenhout, h., 2010, ‘an ecological study of the plant communities in the proposed highveld national park, in the peri-urban area of potchefstroom, south africa’, koedoe 52(1), art. #708, 8 pages. doi: 10.4102/koedoe.v52i1.708 copyright notice: © 2010. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0771 (online) an ecological study of the plant communities in the proposed highveld national park, in the peri-urban area of potchefstroom, south africa in this original research... open access • abstract • introduction • study area    • location    • geology, soil and land types    • climate    • vegetation • method    • stratification of sample sites    • data processing • results    • vegetation classification    • detailed description of plant communities for hnp    • ordination • discussion • conclusion • acknowledgements • references abstract (back to top) the proposed highveld national park (hnp) is an area of high conservation value in south africa, covering approximately 0.03% of the endangered grassland biome. the park is situated immediately adjacent to the town of potchefstroom in the north-west province. the objective of this study was to identify, classify, describe and map the plant communities in this park. vegetation sampling was done by means of the braun-blanquet method and a total of 88 stratified random relevés were sampled. a numerical classification technique (twinspan) was used and the results were refined by braun-blanquet procedures. the final results of the classification procedure were presented in the form of phytosociological tables and, thereafter, nine plant communities were described and mapped. a detrended correspondence analysis confirmed the presence of three structural vegetation units, namely woodland, shrubland and grassland. differences in floristic composition in the three vegetation units were found to be influenced by environmental factors, such as surface rockiness and altitude. incidences of harvesting trees for fuel, uncontrolled fires and overgrazing were found to have a significant effect on floristic and structural composition in the hnp. the ecological interpretation derived from this study can therefore be used as a tool for environmental planning and management of this grassland area. conservation implications: grasslands are amongst the most threatened and least conserved vegetation biomes in south africa, with less than 1.3% currently being conserved. the hnp has significant value for biodiversity conservation and the protection of this area will contribute to the preservation of the highly threatened highveld vegetation types. introduction (back to top) conservation strategies are being developed to expand the number and size of protected areas by incorporating communal lands. south africa stands at the threshold of important and difficult socio-political decisions with respect to land restitution and redistribution (fabricius, koch & magome 2001; hoffman & meadows 2002). new policies allow communities better access to natural resources and call for their participation in the management of protected areas, as well as facilitate the restitution of land from which they had been forcibly removed (fabricius et al. 2001). conservation agencies have, in some instances, been able to expand the size of the protected wildlife estate by entering into negotiations with local residents. this has resulted in a new category of protected areas called ‘contractual parks’. in south africa, examples of this co-management strategy include the richtersveld national park and the agreement with the makuleke people in the kruger national park (archer 1999; steenkamp 1999). although the highveld national park (hnp) is not regarded as a contractual park, all the stakeholders (south african national parks, the barolong community, north-west parks and tourism board, and the tlokwe city council of potchefstroom) constitute the ‘founding partners’, with a memorandum of understanding regulating their respective rights and obligations within the park. this agreement defines the roles and responsibilities of the different partners, including the management and administration of the hnp. the establishment of the hnp aimed at conserving a considerable area of the poorly conserved rocky highveld grassveld (bredenkamp & van rooyen 1996a) and dry sandy highveld grassveld (bredenkamp & van rooyen 1996b) of the western grassland biome in south africa; areas which have been considered to be the most scenic landscapes in the western grassland biome (mucina & rutherford 2006). according to fairbanks et al. (2000), almost 30% of the grassland biome of south africa has been permanently transformed, primarily as a result of cultivation (23%), plantation forestry (4%), urbanisation (2%) and mining (1%). a further 7% has been severely degraded by erosion and agricultural development. in order to maintain biodiversity, it is important to conserve elements such as vegetation and soil, because their loss enhances erosion and reduces the productive value of the land (hoffman & todd 2000; o’brien 2004). urbanisation is a major contributor to the loss of natural areas in certain parts of the grassland biome (cilliers, müller & drewes 2004), particularly in the potchefstroom area. although urbanisation in some provinces has been lower than south africa’s national figure of 53%, a rapid increase is expected in the future (cilliers et al. 2004), as a result of the higher levels of poverty and fewer job opportunities in rural areas that have been caused by continued droughts, specifically in the north-west province. the identification and understanding of plant communities, as well as their associated habitats, are of fundamental importance in the formulation and execution of conservation strategies for the hnp. study area (back to top) location the study focused on the area west of potchefstroom (figure 1), between 26°39’36” s – 26°46’12” s and 26°52’12” e – 27°04’12” e. the area for park development lies between the n12, running from potchefstroom to klerksdorp in the south and eleazer road in the north. the park occupies approximately 10 200 hectares, which consists of a core area of potchefstroom local authority land (5500 ha), modderfontein (defence force land, 2 700 ha), nooitverwacht (privately owned land, 1500 ha)and the agricultural college farm (500 ha). figure 1: location of the highveld national park in north-west province, south africa, with the insert placing the hnp in the context of the rest of south africa geology, soil and land types the relatively heterogeneous geology is represented by the witwatersrand and ventersdorp supergroups and the transvaal sequence, with isolated occurrences of old archaic granites and karoo sequence sediments (bezuidenhout 1993). soils in the study area are heterogeneous and vary from sandy to clayey, due to great variation in parent rock material (land type survey staff 1984). four land types – ba, bc, fa and fb – occur in the hnp (daemane 2007). a land type denotes an area that can be shown at a scale of 1:250 000, displaying marked degree of uniformity with respect to terrain form, soil pattern and climate (land type survey staff 1984). climate the rainfall is erratic and varies from an average of 600 mm per annum, to exceptional occurrences of more than 900 mm per annum (institute for soil, climate and water 2003). the area is characterised by large seasonal and daily variations in temperature, from very hot in summer (daily average temperatures may exceed 32 °c in january), to mild to cold in winter, with average minimum monthly temperatures of up to 12 °c (iscw 2003). vegetation the north-west province encloses two major biomes: the grassland and the savanna biome. the grassland biome in this study was represented by the vaal reefs dolomite sinkhole woodland (gh 12), the rand highveld grassland (gm 11), the klerksdorp thornveld (gh 13), and the carletonville dolomite grassland (gh 15) of the dry highveld grassland bioregion unit (mucina & rutherford 2006). the savanna biome, represented by the central bushveld bioregion unit, consists of andesite mountain bushveld (svcb 11). low & rebelo (1996) classified this area under the rocky highveld grassland and dry sandy highveld grassland. the bankenveld vegetation type (veld type 61), which acocks (1953) described as ‘false grassveld’, is dominant in the hnp; the climax of this veld type is open acacia caffra woodland. method (back to top) stratification of sample sites vegetation sampling was undertaken to represent the communities of the hnp and consisted of 88 relevés of 900 m2, each randomly placed within the stratified units. stratification was done on 1:50 000 scale aerial orthophotographs, on the basis of relatively homogeneous physiographic and physiognomic units. following bezuidenhout (1993), further stratification was carried out by recognising the terrain types associated with topographical positions, such as the crest, midslope, footslope, valley bottomlands and floodplains. sampling procedure plant species were identified in each relevé and the cover abundance of each species was visually estimated using the braun-blanquet cover abundance scale (mueller-dombois & ellenberg 1974). unknown specimens were identified in the kimberley south african national parks herbarium. germishuizen & meyer (2003) was used as reference to confirm the names of taxa. in addition, the average height and canopy cover of tree, shrub and herbaceous strata were estimated. features of the habitat were recorded for each relevé, including geology, soil forms (macvicar et al. 1991), and rockiness of the soil surface data processing the twinspan classification algorithm (hill 1979), which is regarded as a very successful approach by many phytosociologists (mucina & van der maarel 1989; bredenkamp & bezuidenhout 1995; cilliers 1998), was used as a first analysis for the floristic data. the twinspan classification was then refined further, by application of braun-blanquet procedures by means of the bbpc-program (bezuidenhout, biggs & bredenkamp 1996). detrended correspondence analysis (dca; hill 1979) was applied to the data set to confirm the phytosociological association and to assess floristic relations between communities. the stepwise ordination approach followed by mucina & van tongeren (1989), was adopted to cope with the heterogeneity of the data set. descriptive terms such as ‘differential’ and ‘dominant species’ were used following kent & coker (1992) and jennings et al. (2009). ‘differential species’ refers to species of medium to low constancy, which tends to occur together in a series of quadrats and can thus be used to characterise groups. ‘dominant species’ refers to species with the highest percentage of cover, usually in the uppermost dominant layer. results (back to top) vegetation classification the analysis of vegetation by phytosociological means, resulted in the identification of nine different plant communities (figure 2), brief descriptions of which can be found in table 1. the species groups within each plant community can be found in the phytosociological table (see online appendix). the presence of the municipal dumping site within the park is shown in the map (figure 2) and its implications are highlighted in the discussion section. however, the area indicated as ‘spitskop’ on the map is highly degraded; therefore, the plant communities found in this area do not form part of this discussion. figure 2: vegetation map of the plant communities of the highveld national park, north-west province, south africa table 1: a summary of the plant communities described for the highveld national park detailed description of plant communities for hnp 1. mystroxylon aethiopicum – pavetta zeyheri shrubland. this community is mainly limited to the rocky quartzite plateau and ridges of the study area. rocks cover up to 73% of the soil surface. glenrosa is the dominant soil form in this community. the differential species of this community include indigenous species such as pavetta zeyheri, mystroxylon aethiopicum, grewia occidentalis, vangueria parvifolia, pappea capensis, olea europaea subsp. africana and panicum maximum (species group a). dominant species include themeda triandra, elionurus muticus (species group u), vangueria infausta subsp. infausta (species group c), and acacia caffra (species group n). the average height of the tree stratum is 5.3 m, with a canopy cover of 19%. the average height of the herbaceous stratum is 0.9 m, with a canopy cover of 41%. on average, 33 species were recorded per sample plot, of which 1.2% was exotic species. exotic plants included species such as gomphrena celosioides and opuntia ficus–indica (species group v).a closely related shrubland community, pavetta zeyheri –vangueria infausta of the ba land type of the former western transvaal, was described by bezuidenhout, bredenkamp and theron (1994c) and corresponds with the dry sandy highveld grassland. the shrubland communities form a distinctive structural vegetation type within the matrix of the dry sandy highveld grassland. only 0.28% of the dry sandy highveld grassland is currently being conserved in south africa, while approximately 65% of this vegetation has been transformed (bredenkamp & van rooyen 1996b). 2. vangueria infausta subsp. infausta – loudetia simplex shrubland. this community is associated with the midslopes of the rocky quartzite outcrops of the study area. the mispah and glenrosa soil forms are dominant in this habitat. rocks cover up to 63% of the soil surface. the differential species of this community is species group b, which includes the grass loudetia simplex, the shrub mundulea sericea and a forb helichrysum kraussii. the dominant shrub is vangueria infausta subsp. infausta (species group c). the dominant grasses are trachypogon spicatus and andropogon schirensis (species group e), schizachyrium sanguineum, brachiaria serrata, diheteropogon amplectens, eragrostis racemosa, melinis nerviglumis, and aristida meridionalis (species group f), and setaria sphacelata var. sphacelata and eragrostis curvula (species group t). the average height of the tree stratum is 4.6 m and the average canopy cover is 6.8%, with acacia caffra (species group n) forming the major tree component. the average canopy cover of the herbaceous stratum is 53.6%, with an average height of 0.8 m. on average, 34 species were recorded per sample plot, of which 1.6% was exotic species. exotic plants included species such as tagetes minuta (species group l), verbena officinalis (species group s), and conyza bonariensis, achyranthes aspera and nicotiana longiflora (species group v). cilliers, schoeman & bredenkamp (1999) described a closely related shrubland community (vangueria infausta subsp. infausta – rhus pyroides) in the semi-urban area of potchefstroom. according to their findings, the lack of the acacia caffra and the dominance of rhus pyroides in the semi-urban area of potchefstroom is possibly a result of the encroachment of rhus pyroides from the adjacent dolomitic and chert grasslands of the fa land type. a similar community, the vangueria infausta subsp. infausta – acacia caffra woodland was described for the bc land type of the former western transvaal grassland (bezuidenhout & bredenkamp 1991). according to bezuidenhout & bredenkamp (1991), the bc land type can be divided into two broad physiognomic classes, namely woodland and grassland. the woodland competes with the grassland for more or less the same habitat. the grassland is often overgrazed and burned and the dominance of mundulea sericea is indicative of these disturbances. these shrubland communities, typical of the rocky highveld grassland (bredenkamp & van rooyen 1996a), are well worth conserving. 3. diheteropogon amplectens – trachypogon spicatus grassland. this grassland occurs mostly on the rocky plains and midslopes. the geology in this community is characterised by quartzite and andesitic lava. rocks cover 48% of the soil surface. the soil forms comprise the mispah and glenrosa. differential species of this community, indicated by species group d, include the grass – brachiaria nigropedata, the forbs – helichrysum nudifolium var. nudifolium and chascanum adenostachyum, and the succulent – aloe zebrina. the dominant grass species in this community are diheteropogon amplectens (species group f) and trachypogon spicatus (species group e). the average height of the herbaceous stratum is 0.9 m and the canopy cover is 65%. the average height of the tree stratum is 4.6 m, with a canopy cover of 3.8%; acacia caffra (species group n) is the main component of this stratum. the average height of the shrub stratum is 1.5 m, with a canopy cover of 9%. on average, 33 species were recorded per sample plot, with the only exotic species (0.2%) found in this community being conyza bonariensis (species group v). a similar plant community, the trachypogon spicatus –diheteropogon amplectens grassland from the dolomitic region in the potchefstroom–ventersdorp–randfontein area, was previously described by bezuidenhout & bredenkamp (1990). other closely related communities described by bezuidenhout, bredenkamp and theron (1994b) include the diheteropogon amplectens – schizachyrium sanguineum grassland from the fb land type. bezuidenhout et al. (1994b) also found that the diheteropogon amplectens – trachypogon spicatus grassland occurs in the fb land type and, although it is still relatively undisturbed, it is highly threatened by mining and industrial activities elsewhere and is therefore worth conserving. cilliers et al. (1999) described the schizachyrium sanguineum – diheteropogon amplectens grassland in the natural and semi-natural areas in the municipal area of potchefstroom, noting that the variation between these previously described communities could be contributed to factors such as altitude, terrain type, soil type, soil depth and rockiness of soil surface. 4. schizachyrium sanguineum – cymbopogon excavatus grassland. this community is associated with the midslopes of the chert outcrops, where mispah is the dominant soil form. rocks cover 37% of the soil surface. no differential species were identified in this community, but it is characterised by the absence of species within species groups a and c. the dominant species are schizachyrium sanguineum, cymbopogon excavatus, brachiaria serrata, diheteropogon amplectens, and dicoma anomala (species group f), heteropogon contortus (species group q), eragrostis curvula (species group t), and pseudognaphalium undulatum (species group s). the average height of the herbaceous stratum is 0.7 m and it has a canopy cover of 66.3%. the average height of the tree stratum is 6.8 m, with only a poor canopy cover of 1.5%. celtis africana and acacia erioloba (species group g) are the only tree species in this community and are very poorly represented. on average, 54 species were recorded per sample plot, of which 0.7% was exotic species. exotic plants included species such as conyza bonariensis and achyranthes aspera (species group v). bezuidenhout & bredenkamp (1990) described a closely related community, aristida diffusa – cymbopogon excavatus grassland, as also occurring on the stony chert areas of the dolomitic region in the potchefstroom–ventersdorp–randfontein. another related community, cymbopogon excavatus – diheteropogon amplectens grassland, found in the former western transvaal dolomite and chert grassland, was described by bezuidenhout, bredenkamp and theron (1994a). he associated this grassland with the high-lying areas of the fa land type, which has a high conservation status due to its floristic and habitat diversity. 5. rhus pyroides – acacia erioloba woodland. this woodland is limited to the valley bottomlands of the chert and dolomitic areas. mispah is the dominant soil form in this community, with rocks forming 30% of the soil surface. sinkholes, underlain by dolomite, are present in this habitat and mostly filled by deep sandy soil (> 1.2 m). differential species of this community, indicated by species group g, include eragrostis superba, sporobolus africana, eragrostis trichophora, celtis africana, acacia erioloba, diospyros lycioides subsp. guerkei, ipomoea oblongata, sida dregei, and striga asiatica. dominant species in this plant community are acacia erioloba (species group g), triraphis andropogonoides (species group h), ehretia rigida (species group l), rhus pyroides (species group m), and cymbopogon pospischilii (species group q). the average height of the tree stratum is 5.7 m, with a canopy cover of 4.8%. the trees are restricted to deep sandy soils in the sinkholes. the average height of the shrub stratum is 2.8 m, with a canopy cover of 15.6 m. the average height of the herbaceous stratum is 0.8 m, with a high canopy cover of 67%. the succulent lithops lesliei is also found in this community (species group v). on average, 40 species were recorded per sample plot, of which 4.5% was exotic species. exotic plants included tagetes minuta (species group l) and conyza bonariensis, achyranthes aspera, bidens bipinnata, and solanum nigrum (species group v). a closely related woodland community, rhus lancea – acacia erioloba in the former western transvaal (bezuidenhout et al. 1994b), shares a similar floristic composition with the rhus pyroides – acacia erioloba woodland, except for the dominance of rhus pyroides in the current study. the rhus – acacia woodland is aesthetically associated with the most scenic landscapes in the western grassland biome and certainly deserves high conservation priority (bezuidenhout 1993). this community also falls within the vaal reefs dolomite sinkhole woodland (gh 12), which occurs naturally in clumps around sinkholes, especially in places of dolomite outcrops (mucina & rutherford 2006). the hnp is supposed to conserve a considerable area of this vegetation unit (mucina & rutherford 2006). gathering of wood for fuel by rural poor communities is the most significant threat to this plant community. 6. acacia caffra – setaria sphacelata var. sphacelata woodland. the setaria sphacelata var. sphacelata – acacia caffra woodland is mainly associated with the andesitic lava and quartzite midslopes of the rocky outcrops in the study area. rocks cover 27% of the soil surface. glenrosa is the dominant soil form, with mispah and hutton to a lesser extent. the differential species in this community are found in species group i and include the grasses – digitaria eriantha, eragrostis rigidior and aristida transvaalensis, the forbs – crabbea angustifolia, blepharis integrifolia, indigofera heterotricha and chaetacanthus costatu, the tree euclea undulate, and the shrub rhus leptodictya. dominant species in this community include setaria sphacelata var. sphacelata (species group t), acacia caffra (species group n), pseudognaphalium undulatum, aloe greatheadii var. davyana, and clematis brachiata (species group n), ziziphus zeyheriana (species group q), and acacia karroo (species group s).the average height of the tree stratum is 5 m, with a canopy cover of 11.2%. the average height of the shrub stratum is 1.9 m, with a canopy cover of 17.9%. the average height of the herbaceous stratum is 0.9 m, with a canopy cover of 61.7%. on average, 40 species were recorded per sample plot, of which 6.6% was exotic species. exotic plants included species such as tagetes minuta (species group l), verbena officinalis (species group s), paspalum dilatatum, conyza bonariensis, achyranthes aspera, bidens bipinnata, gomphrena celosioides, schkuhria pinnata, verbena bonariensis, gulleminea densa, plantago lanceolata, solanum sisymbriifolium, chenopodium album, opuntia ficus-indica, oxalis corniculata, zinnia peruviana, oenothera tetraptera, and cirsium vulgare (species group v). bezuidenhout et al. (1994b) described a closely related community, rhus leptodictya – acacia caffra woodland of the fb land type in the former western transvaal grassland. another related community (vangueria infausta – acacia caffra woodland) of the bc land type in the former western transvaal grassland, was described by bezuidenhout & bredenkamp (1991). these communities correspond with the andesite mountain bushveld (svcb 11), described by mucina & rutherford (2006), which is dominated by species such as acacia karroo, acacia caffra and ziziphus zeyheriana. intensive mining of the gold-bearing quartzitic rocks of the witwatersrand supergroup has caused destruction and degradation to much of the vegetation within the fb land type (bezuidenhout 1993). uncontrolled fires, gathering of trees for fuel wood and accompanying disturbances, such as trampling and soil compaction, are also significant threats to these communities (cilliers et al. 1999). 7. acacia karroo – ziziphus zeyheriana woodland. the acacia karroo ziziphus zeyheriana woodland occurs on footslopes, valley bottomland areas and along the drainage lines and, occasionally, on the midslopes in degraded areas. rocks cover 3% of the soil surface, with glenrosa and hutton as the dominant soil forms in this community (although mispha and shortland occur in few relevès). the geology is mainly quartzitic, with andesitic lava in few relevès. the differential species in this community include delosperma herbeum, nidorella anomala, sida spinosa, pavonia burchellii, pollichia campestris, pentzia globosa and eragrostis lehmanniana (species group j). dominant species are ziziphus zeyheriana (species group q), acacia karroo (species group s), asparagus suaveolens (species group k), ehretia rigida (species group l), aloe greatheadii var. davyana, clematis brachiata, and acacia caffra (species group n), cymbopogon pospischilii and teucrium trifidum (species group q), cynodon dactylon and asparagus laricinus (species group s), and setaria sphacelata var. sphacelata (species group t). the average height of the tree stratum is 4.5 m and the canopy cover is 17.2%. the average height of the shrub stratum is 1.8 m, with a canopy cover of 21.9%. the average height of the herbaceous stratum is 0.9 m, with a canopy cover of 6.5%. on average, 41 species were recorded per sample plot, with 7.2% exotic species. exotic plants included species such as tagetes minuta (species group l), verbena officinalis (species group s), conyza bonariensis, achyranthes aspera, bidens bipinnata, gomphrena celosioides, lepidium bonariensis, schkuhria pinnata, solanum sisymbriifolium, opuntia ficus – indica, oxalis corniculata, and zinnia peruviana (species group v). bredenkamp, joubert and bezuidenhout (1989) described a similar community (acacia karroo woodland) in the potchefstroom–fochville–parys area, with ziziphus zeyheriana abundantly occurring in severely degraded areas. they also stated that this community is an encroachment of the acacia karroo woodland into grassland communities. the conservation value of this vegetation type, based on the variety in habitat and floristic diversity, was emphasised by bezuidenhout et al. (1994b) in the syntaxonomy of the vegetation of the fb land type in the former western transvaal grassland. 8. cymbopogon pospischilii – themeda triandra grassland. this grassland is associated with the rocky quartzite midslopes of the study area. rocks cover 3% of the soil surface, with glenrosa, hutton and shortland as the dominant soil forms. the differential species in this community are in species group o and include species such as hibiscus pusillus, convolvulus sagittatus, eragrostis chloromelas, menodora africana, gladiolus splendens, helichrysum rugulosum, gymnosporia buxifolia, and anthospermum hispidulum. dominant species in this community are cymbopogon pospischilii and ziziphus zeyheriana (species group q), themeda triandra (species group u) hermannia depressa (species group p), berkheya radula, hyparrhenia hirta, setaria incrassata, and helichrysum nudifolium var. nudifolium (species group r), and acacia karroo and conyza podocephala (species group s). the average height of the tree stratum is 5.4 m, with a canopy cover of 9.8%. the average height of the shrub stratum is 1.2 m, with a canopy cover of 8.2%. the average height of the herbaceous stratum is 0.9 m, with a canopy cover of 77%. on average, 31 species were recorded per sample plot, of which 6.3% was exotic species. exotic plants included species such as tagetes minuta (species group l) verbena officinalis (species group s) conyza bonariensis, achyranthes aspera, avena fatua, bidens bipinnata, gomphrena celosioides, physalis angulata, oenothera rosea, oenothera tetraptera, and cirsium vulgare (species group v). a closely related plant community (themeda triandra – elionurus muticus grassland) with an absolute dominance of themeda triandra, was described by bredenkamp, joubert and bezuidenhout (1989) in a reconnaissance survey of the vegetation of the plains in the potchefstroom–fochville–parys areas. in the bc land type, the cymbopogon pospischilii – themeda triandra grassland shares floristic characteristics with the themeda trianda, a variant described by bezuidenhout & bredenkamp (1991), who found vegetation associated with these plant communities to be in a state of degradation, due to continued overgrazing by domestic stock. 9. setaria incrassata – hyparrhenia hirta grassland. the setaria incrassata – hyparrhenia hirta grassland is restricted to the valley bottomlands and floodplains, and in or along drainage lines within the study area. the geology comprises mostly colluvial deposits from the upper slopes. rocks cover 1% of the soil surface. this habitat is fairly unstable due to seasonal flooding and drying. hutton and swartland are the dominant soil forms found in this habitat. no differential species were identified in this community. setaria incrassata and hyparrhenia hirta (species group r) are the dominant species. the average height of the herbaceous stratum is 0.9 m, with a high canopy cover of 69%. the average height of the shrub stratum is 0.8 m, with a poor canopy cover of 1.3%. the average height of the tree stratum is 3 m and a canopy cover of 6%. on average, 24 species were recorded per sample plot, of which 12% was exotic species. exotic plants included verbena officinalis (species group s), and paspalum dilatatum, conyza bonariensis, ciclospermum leptophyllum, persicaria lapathifolia, oenothera rosea and oenothera tetraptera (species group v). several researchers have done studies on similar wetland grassland communities. bredenkamp et al. (1994) described a hyparrhenia hirta grassland in the boskop dam nature reserve, bezuidenhout et al. (1994b) described a hyparrhenia hirta – eragrostis plana grassland in the fb land type in the fomer western transvaal, and cilliers et al. (1998) gave an account of a hyparrhenia hirta grassland in the potchefstroom municipality. the similarity found in all these communities was that their habitats were fairly unstable due to seasonal flooding and drying which, together with the frequent overgrazing of these sites, caused an advanced state of degradation of the vegetation. ordination dca was applied to the data set to confirm the existence of the different plant communities and to determine whether the gradients in the plant communities along different ordination axes could be explained by gradients in specific environmental characteristics. the gradient along ordination axis 1 (e = 0.65) could be explained by surface rockiness (figure 3). plant communities to the left of the scatter diagram (plant communities 1 and 2) are characterised by a largely rocky soil surface (> 50%), while those on the right (plant communities 7, 8 and 9) are characterised by fewer rocks on the soil surface (0% – 25%). plant communities at the centre of the scatter diagram (plant communities 3, 4, 5 and 6) are characterised by a moderately rocky surface (25% – 50%). the gradient along ordination axis 2 (e = 0.45) could be explained by altitude. plant communities at the top of the scatter diagram occurred mainly on the rocky quartzite plateau and crests in the hnp. plant communities found at the bottom of the scatter diagram are associated with relatively low altitudes and occur mainly in the valley bottomlands of the cherts and dolomitic areas. discussion (back to top) three structural vegetation units, namely woodland, shrubland and grassland, were identified in the hnp. the dca-ordination also confirmed the relationship between the plant communities and the environmental gradients. many of the plant communities previously described in the grassland of the former western transvaal also show floristic and habitat affinities with the plant communities described in the present study. the acacia karroo woodland, as well as several grassland communities, occurs on the plains, while acacia caffra woodland occurs on the rocky outcrops and hills (bezuidenhout & bredenkamp 1991; bezuidenhout et al. 1994a; bezuidenhout et al. 1994b). the plant communities described in the present study also fall within the poorly conserved rocky highveld and dry sandy highveld grasslands previously described by bredekamp & van rooyen (1996a; 1996b). these vegetation types are highly threatened by urbanisation, mining and agriculture. the hnp aims to conserve a considerable area of the vaal reefs dolomite sinkhole woodland (gh 12), because, according to mucina and rutherford (2006), this is one of the most scenic landscapes in the western grassland biome. mucina and rutherford (2006), also point out that almost a quarter of this vegetation unit has already been transformed by mining (the region contains possibly the highest concentration of mines in south africa), cultivation, urban sprawl, and road construction. figure 3: ordination of 87 relevés in highveld national park showing the relationship between surface rockiness and altitude five alien species belonging to category 1 plants were identified in the hnp. category 1 plants are described as prohibited species that should not be tolerated because they are able to invade undisturbed environments and transform or degrade natural plant communities (henderson 2001). these five species are opuntia ficus-indica (species group s), and achyranthes aspera, cirsium vulgare, solanum elaeagnifolium, and solanum sisymbrifolium (species group v). focus should be given to plant communities 6 and 7, as they both have the highest number of category 1 species, with 6.6% and 7.2 % per sample plot, respectively. even though plant community 9 has the highest number of alien species per sample plot (12%), no category 1 species were found in this community. control of these species should aim towards their long-term reduction in density and abundance. it is therefore crucial to prevent the further spread of alien plants into un-invaded areas by isolating the dense patches within the landscape through appropriate control measures and monitoring. the unique scenario of the existence of the municipal dumping site within the park is a cause for concern and it cannot be emphasised enough that proper management and control of the dump site is crucial. landfill exfiltration and water leakage can adversely affect the quality of the adjoining soils and consequently affect the flora and fauna in the adjoining areas. air pollution and wind-carried suspensions can also have direct effects in the park. although the current operations ensure that storm water drains away from the waste to prevent contamination of any runoff, the closure and rehabilitation of the dump is recommended. the aesthetics of the park are compromised by the dump, its associated infrastructure and heavy traffic. the hnp is also threatened by increasing urbanisation, which is why it is crucial to recognise the threats to biodiversity and to understand the factors that will influence the management of the park. for example, settlements along the park’s border pose a threat to the park on a number of levels, including giving rise to noise, air and land pollution, the loss of wilderness ambience, as well as representing a sustained pressure on natural resources. south african cities, including potchefstroom, have witnessed an increase in the number of informal settlements on their urban fringe. there has also been an increase in squatting and informal settlements in rural areas, where fewer job opportunities exist and higher poverty levels are prevalent (cilliers et al. 2004). incidences such as fence cutting, harvesting of trees for firewood, uncontrolled fires and overgrazing, are just a few of the challenges that face the hnp. therefore research on the hnp should focus not only on those problems that can be measured ecologically, but also those that allow for sustainable resource use. this highlights the importance of conservation to the economy in general, and to people’s lives, in particular. the rural and urban poor are those most affected because they depend directly on the natural environment for their livelihood. considerable areas covered by plant communities have not only been transformed by urban sprawl, but existing plant communities on the urban fringe of potchefstroom are also highly fragmented (cilliers et al. 2004). conclusion (back to top) the fundamental role of protected areas has been to separate elements of biodiversity from processes that threaten their existence, by establishing them in remote areas (margules & pressey 2000). recently, however, a more systematic approach to locating and designing protected areas has been evolving – one in which the influences of communities are taken into consideration. this approach needs to be implemented if a large proportion of today’s biodiversity is to endure in the future. the park has not yet been proclaimed, but is currently under the management of north-west parks and tourism board. it will therefore require a management plan that will provide an overarching spatial planning framework. such a framework should address issues regarding urban development close to the park, as well as its future expansion. the ecological interpretation derived from this study can be used as a tool for environmental planning, management and conservation of this grassland area. acknowledgements (back to top) the following individuals and institutions are sincerely thanked: south african national parks for financing this study, north-west university for supporting this research, dr. stephen holness and ms. phozisa mamfengu for assisting with maps, and mr. abbey legari for assisting with field work and the highveld national park’ founding partners for allowing us to undertake this study. editor: llewellyn c. foxcroft references (back to top) acocks, j.p.h., 1953, ‘veld types of south africa’, memoirs of the botanical survey of south africa 28, 1–192. archer, f., 1999, participatory decision-making at richtersveld contractual park: myth or reality?, draft report to the evaluating eden project, international institute for environment and development, london. bezuidenhout, h., & bredenkamp, g.j., 1991, ‘the vegetation of the bc land type in the western transvaal grassland, south africa’, phytocoenologia 19(4), 497–518. bezuidenhout, h., 1993, ‘syntaxonomy and synecology of western transvaal grassland’, phd thesis, department of botany, university of pretoria. bezuidenhout, h. & bredenkamp, g.j., 1990, ‘a reconnaissance survey of the vegetation of the dolomitic region in the potchefstroom–ventersdorp–randfontein area, south africa’, phytocoenologia 18 (2/3), 387–403. bezuidenhout, h., bredenkamp, g.j. & theron, g.k., 1994a, ‘a classification of the vegetation of the western transvaal dolomite and chert grassland, south africa’, south african journal of botany 60(3), 152–161. bezuidenhout, h., bredenkamp, g.j. & theron, g.k., 1994b, ‘the syntaxonomy of the vegetation of the fb land type in the western transvaal grassland, south africa’, south african journal of botany 60(3), 72–81. bezuidenhout, h., bredenkamp, g.j. & theron, g.k., 1994c, ‘the vegetation syntaxa of the ba land type in the western transvaal grassland, south africa’, south african journal of botany 60(4), 214−224. bezuidenhout, h., biggs h.c. & bredenkamp, g.j., 1996, ‘a process supported by the utility bbpc for analysis of braun-blanquet data on personal computer’, koedoe 39, 107–112. bredenkamp, g.j. & bezuidenhout, h., 1989, ‘a phytosociology of the faan meintjies nature reserve in the western transvaal grassland, south africa’, die suid afrikaanse tydskrif vir natuurwetenskap en tegnologie 56(1), 54−64. bredenkamp, g.j., joubert, a.f. & bezuidenhout, h., 1989, ‘a reconnaissance survey of the vegetation of the plains in the potchefstroom–fochville–parys areas’, south african journal of botany 55(2), 199−206. bredenkamp, g.j., bezuidenhout, h., joubert, h. & naude, c., 1994, ‘the vegetation of the boskop dam nature reserve, potchefstroom’, koedoe 37(1), 19−33. bredenkamp, g. j. & bezuidenhout, h., 1995, ‘a proposed procedure for the analysis of large data sets in the classification of south african grasslands’, koedoe 38(1), 33–39. bredenkamp, g.j. & van rooyen, n., 1996a, ‘rocky highveld grassland’, in a.b. low & a.g. rebelo (eds.), vegetation of south africa, lesotho and swaziland, p. 39, department of environmental affairs and tourism, pretoria. bredenkamp, g.j. & van rooyen, n., 1996b, ‘dry sandy highveld grassland’, in a.b. low & a.g. rebelo (eds.), vegetation of south africa, lesotho and swaziland, p. 41, department of environmental affairs and tourism, pretoria. cilliers, s.s., 1998, ‘a phytosociological study of urban open spaces in potchefstroom, north-west province, south africa’, phd thesis, school of environmental sciences and development, potchefstroom university for christian higher education. cilliers, s.s., schoeman, l.l. & bredenkamp, g.j., 1998, ‘wetland plant communities in the potchefstroom municipal area, north-west province, south africa’, bothalia 28(2), 213–229. cilliers, s.s. & bredenkamp, g.j., 1998, ‘vegetation analysis of railway reserves in the potchefstroom municipal area, north-west province, south africa’, south african journal of botany 64(5), 271–280. cilliers, s.s., van wyk, e. & bredenkamp, g.j., 1999, ‘urban nature conservation: vegetation of natural areas in the potchefstroom municipal area, north-west province, south africa’, koedoe 42(1), 1–30. cilliers, s.s, müller, n. & drewes, e., 2004, ‘overview on urban nature conservation: situation in the western-grassland biome of south africa’, urban forestry & urban greening 3, 49–62. daemane, m.e., 2007, ‘an ecological study of the plant communities and degraded areas of the highveld national park, north-west province, south africa’, msc thesis, school of environmental sciences and development, north-west university. edwards, d., 1983, ‘a broad-scale structural classification of vegetation for practical purposes’, bothalia 14(3/4), 705–712. fabricius, c., koch, e. & magome, h., 2001, ‘towards strengthening collaborative ecosystem management: lessons from environmental conflict and political changes in southern africa’, journal of the royal society of new zealand 31(4), 831–844. fairbanks, d.h.k., thomson, m.w., vink, d.e., newby, t.s., van den berg, h.m. & everard, d.a., 2000, ‘the south african land-cover characteristics database: a synopsis of the landscape’, south african journal of science 96, 69–82. germishuizen, g. & meyer, n.l. (eds.), 2003, ‘plants of southern africa: an annotated checklist’, strelitzia 14, 1–1231. hill, m.o., 1979, twinspan a fortran program for the arranging multivariate data in an ordered two-way table by classification of individuals and attributes, cornell university, new york. hoffman, t. & todd, s., 2000, ‘a national review of land degradation in south africa: the influence of biophysical and socio-economic factors’, journal of southern african studies 26(4), 743–758. hoffman, m.t. & meadows, m.e., 2002, ‘the nature, extent and causes of land degradation in south africa: legacy of the past, lessons for the future?’, area 34(4), 428–437. institute for soil, climate and water, 2003, ‘comprehensive report: brakspruit, potchefstroom’, agricultural research council, agromet, pretoria. jennings, m.d., faber-langendoen, d., loucks, o.l., peet, r.k. & roberts, d., 2009, ‘standards for associations and alliances of the u.s. national vegetation classification’, ecological monographs 79, 173–199. kent, m. & coker, p., 1992, vegetation description and analysis: a practical approach, john wiley & sons ltd., chichester. land type survey staff, 1984, ‘land types of the maps 2626 west rand and 2726 kroonstad’, memoirs of agricultural natural resources of south africa 4, 1–441. low, a.b. & rebelo, a.g., 1996, vegetation of south africa, lesotho and swaziland, department of environmental affairs and tourism, pretoria. macvicar, c.n., loxton, r.f., lambrechts, j.j.n. et al., 1991, soil classification a taxonomic system for south africa, department of agricultural development, soil and irrigation research institute, pretoria. memoirs on the agricultural natural resources of south africa 15. margules, c.r. & pressey r.l., 2000, ‘systematic conservation planning’ nature 405, 243–253. mucina, l. & van tongeren, o.f.r., 1989, ‘a coenocline of the high-ranked syntaxa of ruderal vegetation’, vegetation 81, 117–125. mucina, l. & rutherford, m.c. (eds.), 2006, ‘the vegetation of south africa, lesotho and swaziland’, strelitzia 19, 1–807. mucina, l. & van der maarel, e., 1989, ‘twenty years of numerical syntaxonomy’, vegetation 81, 1–15. mueller-dombois, d. & ellenberg, h., 1974, aims and methods of vegetation ecology, john wiley and sons, inc., new york. o’brien, j.w., 2004, ‘vegetation classification, mapping and condition assessment of shamwari game reserve, eastern cape’, msc thesis, terrestrial ecology research unit, university of port elizabeth. steenkamp, c., 1999, ‘the makuleke land claim power relations and cbnrm’, draft report to the evaluating eden project, international institute for environment and development, london. abstract introduction research method and design results discussion conclusion acknowledgements references appendix 1 appendix 2 about the author(s) eduardo m. arraut wildlife conservation research unit, department of zoology, oxford university, united kingdom department of water resources and environment, aeronautics institute of technology, brazil division of remote sensing, national institute for space research, brazil department of plant biology, state university of campinas, brazil andrew j. loveridge wildlife conservation research unit, department of zoology, oxford university, united kingdom lady margaret hall, oxford university, united kingdom simon chamaillé-jammes cefe, cnrs, univ. montpellier, univ. paul valéry montpellier 3, ephe, ird, montpellier, france ltser-france, zone atelier ‘hwange’, hwange national park, zimbabwe department of zoology & entomology, university of pretoria, south africa hugo valls-fox cefe, cnrs, univ. montpellier, univ. paul valéry montpellier 3, ephe, ird, montpellier, france ltser-france, zone atelier ‘hwange’, hwange national park, zimbabwe cirad, umr selmet, montpellier, france selmet, univ montpellier, cirad, inra, montpellier supagro, montpellier, france david w. macdonald wildlife conservation research unit, department of zoology, oxford university, united kingdom citation arraut, e.m., loveridge, a.j., chamaillé-jammes, s., valls-fox, h. & macdonald, d.w., 2018, ‘the 2013–2014 vegetation structure map of hwange national park, zimbabwe, produced using free satellite images and software’, koedoe 60(1), a1497. https://doi.org/10.4102/koedoe.v60i1.1497 original research the 2013–2014 vegetation structure map of hwange national park, zimbabwe, produced using free satellite images and software eduardo m. arraut, andrew j. loveridge, simon chamaillé-jammes, hugo valls-fox, david w. macdonald received: 05 oct. 2017; accepted: 06 aug. 2018; published: 27 sept. 2018 copyright: © 2018. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract vegetation mapping of protected areas is a cornerstone of conservation worldwide. established in 1928 and covering over 1.4 million hectares, hwange national park (hnp) is the largest natural reserve in zimbabwe. in 1993, the sole comprehensive map of its vegetation to date was produced and since then it has been used in numerous research and conservation endeavours. over the last two decades, however, the park’s vegetation changed, safari areas and forest reserves were created at its edge and high-positional accuracy data on a suite of species were collected. to tend to contemporary mapping needs, in this article, we present the 2013–2014 vegetation structure map of hnp and its surroundings. it was produced by supervised classification of landsat-8 operational land imager (oli) images, indices derived from these and the landsat tree cover continuous field product. its accuracy was assessed statistically using samples collected from high-resolution satellite imagery and basic ancillary field data. of its total pixels, 83.2% were correctly classified. mean omission and commission error were, respectively, 0.82 (0.74–0.90) and 0.82 (0.72–0.89), and this similarity held on a per class basis, indicating reliable area estimates. it was produced using only freely available imagery and software. conservation implications: in addition to providing researchers and conservationists working within and around hnp with an updated vegetation map, aiming at an even broader audience, we provide a step-by-step approach for using modern freely available imagery and software for cost-effectively mapping hnp in future or other protected savannas across africa. introduction vegetation mapping of protected areas and their surroundings has been a cornerstone of conservation planning and management worldwide (freemantle et al. 2013; pettorelli, safi & turner 2014; rose et al. 2015). in africa, for example, it has allowed for inferring the health of the threatened miombo woodlands of mozambique (sedano, gong & ferrão 2005), assessing the large-scale impacts of herbivores upon the structural diversity of kruger national park’s vegetation (asner et al. 2009), predicting the risk of african lion (panthera leo) attacks on humans in tanzania (kushnir et al. 2014) and planning of the great limpopo transfrontier conservation area (martini et al. 2016). such mapping has also been at the heart of conservation initiatives in south america (oliveira et al. 2017), asia (davies, murphy & bruce 2016), europe (palomo et al. 2013) and north america (wiens et al. 2009). hwange national park (hnp), covering an area of over 1.4 million hectares, is the largest protected area in zimbabwe and part of the kavango-zambezi transfrontier conservation area (the government of the republic of angola et al. 2011). it is home to rich biodiversity and adjacent to important land concessions (loveridge, reynolds & milner-gulland 2007a, loveridge et al. 2007b). to our knowledge, since hnp’s establishment in 1928, three maps of its vegetation has been produced. the first one, of the robins area, was based on aerial photography and the vegetation was manually classified (robinson, hill & rushworth 1973). twenty years later, similar data were used to produce the park’s first comprehensive map of major vegetation types and structures (rogers 1993). for the following two decades, this was the reference map in studies concerning lions (davidson et al. 2012; loveridge et al. 2009; valeix et al. 2009), jackals (canis mesomelas and canis adustus) (loveridge & macdonald 2002), elephants (loxodonta africana) and other herbivores (chamaillé-jammes et al. 2008; periquet et al. 2012; valeix et al. 2011). the more recent map, which covers a third of the park, was made using landsat thematic mapper images acquired in 2002–2003 and used to study the responses of zebras (equus quagga) to lion encounters (courbin et al. 2016). the map introduced in this study represents the status of the vegetation structure in 2013–2014, thus representing a 20-year update in the park’s mapping, and encompasses hnp plus a 50 km buffer around it, to which safari areas, forest reserves, communal lands and research and conservation endeavours have expanded in the last decade. in addition, the map herein was subject to a statistical accuracy assessment, making it particularly suitable for analyses involving the modern geo data sets that come with positional precision estimates, such as global positioning system (gps) telemetry, and which have been collected on a suite of species living in hnp over the last two decades. in addition to budgetary constraints, its production required overcoming challenges related to (1) difficulty of visiting remote areas on the ground, (2) marked differences in ground colour because of soil and fire and (3) asynchronicity in phenology. as explained earlier, we overcame these using free images, products and software. we believe that our procedure could be used for retrospective analyses of vegetation change in hnp and the production of future maps of the area, as well as for mapping similar classes of savanna vegetation structure within and around other protected areas. we hope the detailed information concerning the making of a vegetation structure map ‘from scratch’ provided here may represent a case study for practitioners with similar needs elsewhere. research method and design setting: study area the 46 207 km2 study area includes hnp (14 651 km2) and adjacent safari areas, forest reserves and communal lands in zimbabwe and botswana within a 50 km buffer around hnp (figure 1). the vegetation is typical of a highly heterogeneous dystrophic wooded savanna (figure 2). overall, woody cover increases with distance from water pans (chamaillé-jammes, fritz & madzikanda 2009). sandy soils, locally known as kalahari sands, that cover about two-thirds of hnp are generally dominated by baikiaea plurijuga woodlands or a mixed bushland community dominated by combretum spp., terminalia sericea and a few acacia spp. groves, with open grasslands along drainage lines. to the north, basaltic and clayey soils are dominated by woody species, such as colophospermum mopane bushland or woodlands (rogers 1993). outside the hnp, additional vegetation types include farmlands and extensive wetlands and seasonal riverbeds to the south-east. altitude throughout the study area varies from c. 835 m to c. 1200 m above sea level. annual rainfall is highly variable (coefficient of variation = 25%) around a mean of 600 mm. rains predominate between november and february (chamaillé-jammes, fritz & murindagomo 2006), with numerous water pans being formed during the rainy season and most drying out completely in the dry season. to sustain animals during the dry season, within hnp, underground water is pumped to some artificial waterholes (chamaillé-jammes, fritz & murindagomo 2007). figure 1: study area with hwange national park boundary (continuous black line) and the park’s location in southern africa. dashed black line within hwange national park limits outlines area shown in figure 6. underlying image is a mid-dry season landsat 8 operational land imager rgb654. figure 2: examples of the vegetation structure classes and an unclassified condition: (a) grassland, (b) bushed grassland, (c) bushland on kalahari sand, (d) deciduous woodland on kalahari sand, (e) scrubland on basalt, (f) deciduous woodland on basalt, (g) evergreen woodland and (h) buildings in the town of dete illustrating the unclassified condition. materials field data the field protocol consisted of visiting sites, making schematic diagrams and taking geotagged photos. field sites were representative of vegetation structure classes and underlying ground signals (see figure 2). their locations were chosen based on expert knowledge and accessibility by car or on foot. geotagged photos were taken of the 208 field sites visited, while 76 diagrams were made. the purpose of making diagrams was to oblige the researcher to carefully observe and record on paper the surroundings of a field site. this information would later aid interpretation of the photos. satellite image data and products only free images and products were used in the study. these included 30 m × 30 m resolution bands 2–6 of 27 landsat 8 (l8) operational land imager (oli) images (see appendix 1), downloaded from the earthexplorer website (https://earthexplorer.usgs.gov/), and the 2005 vegetation coninuous fields (vcf) tree cover product (sexton et al. 2013), available at nasa’s global land cover facility (http://glcf.umd.edu/data/landsattreecover/). thematic classes used in classification the vegetation structure classes were (1) grassland, (2) bushed grassland, (3) bushland on kalahari sand, (4) deciduous woodland on kalahari sand, (5) scrubland on basaltic soil, (6) deciduous woodland on basaltic soil and (7) evergreen woodland (these numbers are used to refer to these classes elsewhere in the article; classes with no associated soil class name occurred on both soil types). however, these were not the actual classes on which the classification was based because during fieldwork we identified three confounding factors: (1) difference in the reflective properties of the two major soil types, (2) fire and (3) asynchronicity in phenology. the effect of the difference in soil type was more prominent in the dry season images, when the weaker signal of the dry vegetation led to an increase in the relative contribution to pixel reflectance of the underlying soil signal. as the classification required multitemporal images and different vegetation structure classes had the same underlying soil type, this played towards reducing their spectral separability. burning reduced dramatically the reflectance of the target, and because it happened to different vegetation classes, it also contributed to reducing their separability. finally, asynchronicity in phenology implied that the same vegetation structure class appeared vigorous and dry in the same image, resulting in an increase in intra-class variance, and hence, a reduction in inter-class separability. to solve these issues, the classification was carried out considering a subdivision of the desired vegetation classes into ‘spectral classes’. these were composed of the above vegetation structure classes plus the classes ‘bare grassy areas near waterholes’, ‘grassland, burnt’, ‘deciduous woodland on kalahari soil, burnt’ and ‘scrubland on basaltic soil, burnt’. as will be explained later, these spectral classes were grouped into the desired vegetation structure classes during post-processing. procedure a summary of the steps for producing the final map is presented in figure 3. figure 3: overview of procedure for producing the vegetation structure map. image visual and digital processing were carried out using grass 7.0 (grass 2012) and qgis desktop 2.8 with the grass 6.4 plugin (development team qgis 2015). grass 7.0 was used mainly during pre-processing, and classification and initial post-processing steps (filtering), while qgis 2.8 with the grass 6.4 plugin was used mainly during late post-processing (manual editing and merging edited layer with original classification) and accuracy assessment (figure 3). geotagged photos were imported using the photo2shape plugin in qgis (bruy 2013), which creates a georeferenced point vector layer of each photo. the openlayers qgis plugin (sourcepole 2014) was used to visualise high-resolution quickbird images from google earth during the response sample definition (see ‘accuracy assessment’ section). a general introduction to the newest grass manual, which contains the explanations and algorithms regarding image classification, may be found on the internet – at time of this publication, it was grass 7.4 (grass 2017). pre-processing the first step was to convert each band of each scene to surface reflectance using the i.landsat.toar function with dos4 option enabled and parameter values set from the metadata file downloaded with each scene. this particular algorithm reports at-surface reflectance by, firstly, performing a radiometric calibration (using gain and bias parameters) and then correcting for acquisition geometry (using estimates for the distance from the sun, solar elevation angle and mean solar exoatmospheric irradiance) and, secondly, removing atmospheric path radiance (scattering) with the dark object method (chavez 1988; vincent 1972). the i.landsat.toar sets pixels in the image’s bounding square to zero, which is a problem for the mosaicking algorithm used later (it considers them meaningful values). to avoid this, the r.null function was applied to them. then, the r.patch function was used to produce three mosaics with nine scenes each, for early-dry, mid-dry and late-dry seasons. as at the time, the grass algorithms could not handle 16-bit images, all mosaics were converted from 16 to 8 bit. true colour rgb432 and false colour rgb645 composites were then produced. we then used the grass function i.vi to produce the enhanced vegetation index 2 (evi-2) (jiang et al. 2008) and the soil adjusted vegetation index (savi) (huete 1988), and i.pca to produce the first, second and third principal components of the late-dry season oli bands. the evi-2 and savi were used, respectively, to help differentiate classes during the earlyand mid-dry seasons, when they are useful for differentiating green vegetation with different metabolism and canopy structure, and during the late-dry season, when the soil signal from underneath the dry vegetation is more noticeable. the principal components were made to further aid the differentiation of classes during the dry season. to define the study region, a mask corresponding to hnp plus a 50 km boundary strip was applied using the r.mask function in grass. supervised classification image grouping: in grass geographic information system (gis), all training sample statistics and classifications are performed on image groups and subgroups, which are defined using the function i.group. an image group is a named collection of raster layers and a subgroup is a specific combination of the layers present in the group. by varying the subgroup used in each of a series of preliminary classifications, the user can test which combinations of layers within a group yield the best results. training sample: training sample was created using field information, high-resolution quickbird images available freely in google earth, the lvcf and the landsat-8 oli bands from wet, early-, midand late-dry seasons. to define the samples, a vector layer was created in grass 6.4 with a column ‘cover’ representing a code for each land-cover class. the function v.edit was used to create the polygons delimiting training sample acquisition. this vector layer was then converted to raster using v.to.rast, with the source of raster values (the land-cover class) coming from the column ‘cover’. this new raster layer was used as the reference map. it delimited areas in space from which sample statistics were derived from the image subgroups. the final training sample was defined after an iterative process in which classifications involving training samples with progressively greater per-class variance were visually contrasted, that is, one begins by classifying using samples obtained from pixels representing homogeneous areas of each vegetation type and at every step adds variance to each sample by including pixels located at class transition zones. supervised classification: supervised classification was carried out using the bayesian multiscale random field sequential maximum a posteriori (smap) classifier available in grass 7.0 (bouman & shapiro 1994; mccauley & engel 1995). when classifying a pixel, this classifier considers information from its contiguous neighbours and checks that the class assigned to it appears consistent by sequentially comparing which classes predominate in a set of larger neighbourhoods around this pixel. this effectively implies that it classifies a pixel based on information about the land-cover patch within which it is situated. sequential maximum a posteriori classifiers have been shown to perform better than classifiers based on maximum likelihood (ml) or other bayesian-based approaches (bouman & shapiro 1994; mccauley & engel 1995); we confirmed this by carrying out preliminary classifications using the ml algorithm also available in grass and using our field data and knowledge to visually compare its results with those produced by smap. to carry out the classification, firstly, class statistics were generated using the i.smap function and using the raster reference map and subgroups mentioned above. this created the signature file, which contained the reference statistics for each of the classes to be used in the classification. secondly, contextual image classification using smap estimation was performed by the function i.smap using the aforementioned groups and signature file. post-processing the function r.neighbors was used to apply 3 × 3, 5 × 5, 7 × 7, 9 × 9 and 11 × 11 median neighbourhood filters to the final classification. median filters reduce classification noise without altering class values (in contrast to, for example, a mean filter, which alters class values). to the best of our judgement, the 5 × 5 filter yielded the better results and therefore the classification to which we applied it was chosen as the final map. then, manual vector editing was performed to assign the correct class to some areas that had been so recently and intensely burnt that the classifier confounded them with water (which has extremely low reflectance in the l8 bands we were using). virtually all of the edited area fell outside hnp and it amounted to approximately 3% of the total mapped area and about 20% of the total recently burnt area (as estimated using qgis to manually delimit burnt areas in the late-dry season image). for this purpose, a vector layer was created in qgis and polygons with attribute values defining the new classes were created. the rasterise function in qgis was then used to merge these polygons with the unedited raster classification, thus producing the map to be subject to accuracy assessment. accuracy assessment accuracy assessment was carried out following nomenclature and recommendations by olofsson et al. (2014) and stehman and czaplewski (1998). the sampling unit was the vector point, and the sampling design, which is the method by which the sampling units are chosen from the statistical population (the map), consisted of placing 500 points on the map using a simple random sampling scheme (distribution of the points shown in appendix 2). the distribution of points across classes was then compared to the distribution of class areas and their similarity indicated that the scheme had not favoured any class. this illustrates a key property of a simple random sampling scheme, which is not to attribute importance to any class a priori. as a consequence, the interpretation of overall and class probabilities becomes straightforward: (1) global accuracy can be translated into the probability that a point on the map was correctly classified, (2) producer’s accuracy (omission error) to the probability that a patch of a certain vegetation structure type was correctly classified and (3) user’s accuracy (commission error) to the probability that a patch of a map class represents the correct cover type. the response design, which is the protocol by which the reference class is associated with each sampling unit, consisted of attributing, to the point, the class predominating within a 60 m radius centred on it (figure 4). this buffer size was chosen to minimise co-registration errors. the assessment of which class predominated within a buffer was made using high-resolution quickbird images (figure 4), aided by all other satellite and field data and information available (including a data set of images from camera-traps deployed throughout most of hnp). the computation of statistics was carried out using the r.kappa algorithm, which produced the confusion matrix and calculated classes’ accuracy statistics and overall percentage of correctly classified pixels. global, producer and user accuracies were then estimated from the confusion matrix. as several scientific studies are restricted to within hnp limits, in addition to assessing the accuracy of the entire mapped area, we also estimated each of the accuracy parameters for the area restricted to within the park’s limits (n = 198 points). figure 4: examples of high-resolution quickbird images used for creating validation samples. yellow circles indicate 120 m-buffers around locations of validation points (images shown vary in scale, hence variation in apparent buffer size). results the 2013–2014 vegetation structure map of hnp and its surroundings is shown in figure 5. figure 5: the 2013–2014 vegetation structure map of hwange national park and its surroundings. as an example of the map’s ability to capture areas that are vital for one of hnp’s most emblematic species, figure 6 shows how lion gps locations fell within the grassland and bushed grassland areas that these animals regularly use for resting, foraging and commuting between waterholes. figure 6: superimposing high-accuracy (< 30 m) lion global positioning system locations on the map provides visual indication of good mapping of grassland and bushed grassland classes. of the 500 samples used in validation, 83.2% were classified correctly (table 1), and when only the 198 validation points that fell within hnp were considered, this percentage increased to 89.9%. as this map was validated using a simple random sampling scheme, the measures of global accuracy can be interpreted as the chance that a randomly chosen pixel (0.09 ha) was correctly classified. table 1: quantification of classification error using confusion matrix produced from 500 validation points obtained using a simple random sampling scheme. producer accuracies (pas) varied between 0.90 and 0.74 (mean = 0.83; standard deviation [sd] = 0.06) and user accuracies (uas) between 0.89 and 0.72 (mean = 0.822; sd = 0.06), and the per-class difference (in modulus) between them varied between 0.15 (grassland) and 0.00 (deciduous woodland on kalahari), suggesting class area estimates were reliable (the smaller the difference, the more reliable the area estimate). the two classes that conformed less to this pattern were 1 (grassland), with pa versus ua = 0.15, and 6 (deciduous woodland on basalt), with pa versus ua = 0.14. in class 1, higher pa (0.87) than ua (0.72) indicated that most grassland patches were correctly identified, but overall class area was slightly overestimated by attribution of this class to patches covered by other types, namely, bushed grassland, bushland and deciduous woodland on basalt (table 1). these classes are characterised by a continuous underlying grass cover in places where shrubs or trees are sparse. in such areas, the signal from the lvcf gets weaker and that from the grasses becomes stronger, resulting in a preponderance of the latter and hence confusion with the grassland class. conversely, in class 6, the lower pa (0.75) and higher ua (0.89) suggested that this vegetation structure type was rarely wrongly mapped, but that patches where it occurs were sometimes assigned to other classes, implying that its area was marginally underestimated. final estimates for the area occupied by each vegetation structure class are presented in table 2. table 2: area and percentage cover of each vegetation structure class. discussion the map introduced in this article depicts the state of the vegetation in hnp and surrounding reserves and land concessions in 2013–2014 (figure 5), 20 years more recently than the previous map of the entire park (rogers 1993). the vegetation classes distinguished in it are rather broad classes defined on the basis of vegetation structure. in particular, the classes reflect variability in openness and vegetation height, which we consider to have important and numerous functional consequences. for instance, in our own work on large mammal ecology, we have previously shown that dense and bushy areas where visibility is low, rather than dense woodlands where visibility is greater, are selected for by lions, which are ambush predators (davidson et al. 2012). thus, we think that the classes identified should be useful to map and understand functional processes at the landscape scale. also, the map produced here offers baseline data to monitor, at a large scale, changes in these key variables (openness and height). in hnp, this is particularly important in the face of the persistence of a large elephant population that may affect vegetation structure in the long run (valeix et al. 2011). as this map was validated via a simple random sampling scheme, its classes can be combined to create other maps with global and per-class accuracy statistics easily calculable from the original confusion matrix. thus, if, for example, the grassland and bushed grassland classes should be grouped into the ‘open areas’ class, which has been shown to be strongly selected for by zebras (courbin et al. 2016), this new class would have pa = 0.89 and ua = 0.80 (n = 117). for a recent review of the applicabilities and limitations of other accuracy assessment schemes, most of which can be quite easily implemented using freely available gis (development team qgis 2015; grass 2012; see methods section), the readers are referred to stehman and wickham (2011). main confounding factors and possible solutions the principal confounding factors were ground colour and variation in phenology. with regard to ground cover, two signals were observed: a brighter one, produced by the kalahari sands in the central and southern parts, and a darker one, predominantly from basaltic rocks and soil to the north, north-west and extreme south-east. the contribution of this difference to classification uncertainty can be illustrated by an example. when green, grasses present the characteristic vegetation signal with a particularly high infrared reflectance (ceccato et al. 2001). in our study area, however, as the grasses dried and vegetation cover diminished, the contribution of soil to the overall signal became stronger and more variable, leading to an increase in class variance and overlap with other classes. this probably explains most of the commission error for grassland (table 1). in future classifications, a possible way of avoiding this would be to establish two subregions and classify them independently. prior to the accuracy assessment, the two subregions should ideally be merged back, as this would allow class statistics to apply to the entire map. by preventing confusion between current kalahari sand and basalt classes, this measure alone could increase map accuracy by up to about 6% (up to 31 of the 84 errors in table 1 could disappear). a secondary confounding factor was variation in phenology because of earlier drying out of the vegetation in the extreme south-west of the park (in figure 1, whitish area in the lower left), which was the product of a rainfall gradient (chamaillé-jammes et al. 2006). when dry, deciduous plants re-absorb most of the photosynthetic pigments, and, to save water and reduce gas exchange with the atmosphere, they lose leaves or the cell mesophylls of the leaves that remain collapse (asner 1998). for all vegetation structure types considered here, this flattens the response in the visual bands and decreases it in the near-infrared bands (asner 1998; asner & lobell 2000; ceccato et al. 2001; elvidge 1990). as with the soil cover issue, the net effect is the reduction in the ability to separate deciduous woodland from other classes that also lose their leaves or die, such as grassland, bushed grassland or bushland. this probably explains a better part of the confusion involving these three classes (table 1). a way to try to minimise this error would be to substitute the early-dry season image used here with one acquired prior to the vegetation in the south-west of the park drying out (i.e. an image from february or march). an added benefit would be that in such image burn scars from that year would likely be less frequent, which would further contribute to reducing class confusion and hence decrease the requirement for manual editing of the final classification. when this study was carried out, the landsat 8 satellite had been orbiting the earth for only one and a half years and no such image existed. as the oli sensor images the earth every 16 days, the chances of it acquiring an image free of cloud cover naturally increase every year. broader significance of this mapping endeavour the principle underlying vegetation mapping is to translate the spectral data acquired by a remote sensor into the desired vegetation categories. in the past four decades, the means to do so have evolved fast owing chiefly to advancement in the field of remote sensing, particularly regarding the diversification of the spatial, spectral, temporal and radiometric resolutions of sensors and of the ability to store, analyse and interpret the data they acquire (chuvieco 2016; jensen 2013). nevertheless, remote sensing remains highly underused by conservationists, mainly because of the cost of and access to imagery and software (turner et al. 2015). aiming at making such new technologies more easily accessible to the end user, the international research community, backed by governments and transnational institutions, has pushed forward an agenda to make satellite imagery, derived products (fonseca et al. 2014; wulder et al. 2012) and software (development team qgis 2015; gdal 2018; grass 2012; inglada & christophe 2009) freely available to all in easy-to-use repositories. given our requirement of a vegetation structure map to be used by several ongoing ecological and conservation studies within and around hnp, coupled with our limited budget for producing it, we opted to tackle the challenge of creating one using solely such freely available data and software. we hope that by providing a step-by-step description of our mapping (see the ‘materials’ and ‘procedure’ sections), our work may help researchers and conservationists produce future maps of hnp or similar maps of other protected savannas at a low cost. conclusion vegetation mapping of protected areas is a cornerstone of conservation and in this article we tend to the needs of researchers and conservationists working within and around hnp by providing a recent and validated vegetation structure map of the park and its surroundings. the map’s spatial resolution and accuracy are similar to those of data collected using other modern technology, such as gps telemetry, making it particularly suitable for analyses involving such data. aiming at making this work useful to an even broader audience, we provide a step-by-step approach for people with some field knowledge, but modest remote sensing experience, to use freely available data and software to produce future maps of hnp or similar maps of other protected savannas across africa. acknowledgements the authors thank zimbabwe parks and wildlife management authority for giving them permits and authorisation to conduct this research. they also deeply appreciate the fundamental support during fieldwork provided by brent stapelkamp, justin seymour-smith and andreas sibanda, the insightful comments during manuscript preparation by dr evlyn novo and dr marion valeix and the photo of dete, zimbabwe, supplied by and authorised for use by ms jennifer ailloud. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions e.m.a. planned the research, produced the classification and performed the accuracy assessment. e.m.a. and s.c.j. conducted the fieldwork and h.v.f assisted. a.j.l. and d.w.m. contributed with financial support and ancillary field data. all authors critically revised the manuscript. funding information this work was funded by instituto nacional de ciência e tecnologia para mudanças climáticas (inct-mc), conselho nacional de desenvolvimento científico e tecnológico (cnpq), uk natural environment research council and oxford university, grant anr-11-ceps-003 of the french ‘agence national de la recherche’, together with grants to d.w.m. from the kirk-turner, robertson and recanati-kaplan foundations. references asner, g.p., 1998, ‘biophysical and biochemical sources of variability in canopy reflectance’, remote sensing of environment 64, 234–253. https://doi.org/10.1016/s0034-4257(98)00014-5 asner, g.p., levick, s.r., kennedy-bowdoin, t., knapp, d.e., emerson, r., jacobson, j. et al., 2009, ‘large-scale impacts of herbivores on the structural diversity of african savannas’, proceedings of the national academy of sciences of the united states of america 106, 4947–4952. https://doi.org/10.1073/pnas.0810637106 asner, g.p. & lobell, d.b., 2000, ‘a biogeophysical approach for automated swir unmixing of soils and vegetation’, remote sensing of environment 74, 99–112. https://doi.org/10.1016/s0034-4257(00)00126-7 bouman, c.a. & shapiro, m., 1994, ‘a multiscale random field model for bayesian image segmentation’, ieee transactions on image processing 3, 162–177. https://doi.org/10.1109/83.277898 bruy, a., 2013, photo2shape v0.2.2 plugin for qgis, open source geospatial foundation project, viewed 01 september 2014 from http://qgis.osgeo.org ceccato, p., flasse, s., tarantola, s., jacquemoud, s. & grégoire, j.m., 2001, ‘detecting vegetation leaf water content using reflectance in the optical domain’, remote sensing of environment 77, 22–33. https://doi.org/10.1016/s0034-4257(01)00191-2 chamaillé-jammes, s., fritz, h. & madzikanda, h., 2009, ‘piosphere contribution to landscape heterogeneity: a case study of remote-sensed woody cover in a high elephant density landscape’, ecography 32, 871–880. https://doi.org/10.1111/j.1600-0587.2009.05785.x chamaillé-jammes, s., fritz, h. & murindagomo, f., 2006, ‘spatial patterns of the ndvi–rainfall relationship at the seasonal and interannual time scales in an african savanna’, international journal of remote sensing 27, 5185–5200. https://doi.org/10.1080/01431160600702392 chamaillé-jammes, s., fritz, h. & murindagomo, f., 2007, ‘climate-driven fluctuations in surface-water availability and the buffering role of artificial pumping in an african savanna: potential implication for herbivore dynamics’, austral ecology 32, 740–748. https://doi.org/10.1111/j.1442-9993.2007.01761.x chamaillé-jammes, s., fritz, h., valeix, m., murindagomo, f. & clobert, j., 2008, ‘resource variability, aggregation and direct density dependence in an open context: the local regulation of an african elephant population’, journal of animal ecology 77, 135–144. https://doi.org/10.1111/j.1365-2656.2007.01307.x chavez, p.s., 1988, ‘an improved dark-object subtraction technique for atmospheric scattering correction of multispectral data’, remote sensing of environment 24, 459–479. https://doi.org/10.1016/0034-4257(88)90019-3 chuvieco, e., 2016, fundamentals of satellite remote sensing, 2nd edn., crc press, boca raton, fl. courbin, n., loveridge, a.j., macdonald, d.w., fritz, h., valeix, m., makuwe, e.t. et al., 2016, ‘reactive responses of zebras to lion encounters shape their predator-prey space game at large scale’, oikos 125, 829–838. https://doi.org/10.1111/oik.02555 davidson, z., valeix, m., loveridge, a.j., hunt, j.e., johnson, p.j., madzikanda, h. et al., 2012, ‘environmental determinants of habitat and kill site selection in a large carnivore: scale matters’, journal of mammalogy 93, 677–685. https://doi.org/10.1644/10-mamm-a-424.1 davies, k.p., murphy, r.j. & bruce, e., 2016, ‘detecting historical changes to vegetation in a cambodian protected area using the landsat tm and etm+ sensors’, remote sensing of environment 187, 332–344. https://doi.org/10.1016/j.rse.2016.10.027 development team qgis, 2015, qgis geographic information system, open source geospatial foundation project, viewed 01 september 2014, from http://qgis.osgeo.org elvidge, c.d., 1990, ‘reflectance characteristics of dry plant materials’, international journal of remote sensing 11, 1775–1795. https://doi.org/10.1080/01431169008955129 fonseca, l., epiphanio, j., valeriano, d., soares, j., dalge, j. & alvarenga, m., 2014, ‘earth observation applications in brazil with focus on the cbers program’, ieee geoscience and remote sensing magazine 2, 53–55. https://doi.org/10.1109/mgrs.2014.2320924 freemantle, t.p., wacher, t., newby, j. & pettorelli, n., 2013, ‘earth observation: overlooked potential to support species reintroduction programmes’, african journal of ecology 51, 482–492. https://doi.org/10.1111/aje.12060 gdal, 2018, gdal – geospatial data abstraction library, viewed 01 april 2018, from http://www.gdal.org grass development team, 2012, geographic resources analysis support system (grass) software, viewed 01 september 2014, from http://grass.osgeo.org. grass development team, 2017, image processing in grass gis: 7.4.1svn reference manual, viewed 10 february 2017, from https://grass.osgeo.org/grass64/manuals/imageryintro.html. huete, a.r., 1988, ‘a soil-adjusted vegetation index (savi)’, remote sensing of environment 25, 295–309. https://doi.org/10.1016/0034-4257(88)90106-x inglada, j. & christophe, e., 2009, ‘the orfeo toolbox remote sensing image processing software’, 2009 ieee international geoscience and remote sensing symposium, iv-733–iv-736, cape town, south africa, july 12–17, 2009. https://doi.org/10.1109/igarss.2009.5417481 jensen, j.r., 2013, remote sensing of the environment: an earth resource perspective, 2nd edn., prentice hall, nj. jiang, z., huete, a., didan, k. & miura, t., 2008, ‘development of a two-band enhanced vegetation index without a blue band’, remote sensing of environment 112, 3833–3845. https://doi.org/10.1016/j.rse.2008.06.006 kushnir, h., weisberg, s., olson, e., juntunen, t., ikanda, d. & packer, c., 2014, ‘using landscape characteristics to predict risk of lion attacks on humans in south-eastern tanzania’, african journal of ecology, 52(4), 524–532. https://doi.org/10.1111/aje.12157 loveridge, a. & macdonald, d., 2002, ‘habitat ecology of two sympatric species of jackals in zimbabwe’, journal of mammalogy 83, 599–607. loveridge, a., reynolds, j.c. & milner-gulland, e.j., 2007a, ‘does sport hunting benefit conservation’, in d. macdonald & s. katerina (eds.), key topics in conservation biology, pp. 224–240, wiley-blackwell, oxford. loveridge, a., searle, a., murindagomo, f. & macdonald, d., 2007b, ‘the impact of sport-hunting on the population dynamics of an african lion population in a protected area’, biological conservation 134, 548–558. https://doi.org/10.1644/1545-1542(2002)083%3c0599:heotss%3e2.0.co;2 loveridge, a.j., valeix, m., davidson, z., murindagomo, f., fritz, h. & macdonald, d.w., 2009, ‘changes in home range size of african lions in relation to pride size and prey biomass in a semi-arid savanna’, ecography 32, 953–962. https://doi.org/10.1111/j.1600-0587.2009.05745.x martini, f., cunliffe, r., farcomeni, a., sanctis, m. de., d’ammando, g. & attore, f., 2016, ‘classification and mapping of the woody vegetation of gonarezhou national park, zimbabwe’, koedoe 58, 1–10. https://doi.org/10.4102/koedoe.v58i1.1388 mccauley, j.d. & engel, b.a., 1995, ‘comparison of scene segmentations: smap, echo, and maximum likelihood’, ieee transactions on geoscience and remote sensing 33, 1313–1316. https://doi.org/10.1109/83.277898 oliveira, u., silveira, b., filho, s., paglia, a.p., brescovit, a.d., de carvalho, c.j.b. et al., 2017, ‘biodiversity conservation gaps in the brazilian protected areas’, scientific reports 7, 1–9. https://doi.org/10.1038/s41598-017-08707-2 olofsson, p., foody, g.m., herold, m., stehman, s.v., woodcock, c.e. & wulder, m.a., 2014, ‘good practices for estimating area and assessing accuracy of land change’, remote sensing of environment 148, 42–57. https://doi.org/10.1016/j.rse.2014.02.015 palomo, i., martín-lópez, b., potschin, m., haines-young, r. & montes, c., 2013, ‘national parks, buffer zones and surrounding lands: mapping ecosystem service flows’, ecosystem services 4, 104–116. https://doi.org/10.1016/j.ecoser.2012.09.001 periquet, s., todd-jones, l., valeix, m., stapelkamp, b., elliot, n., wijers, m. et al., 2012, ‘influence of immediate predation risk by lions on the vigilance of prey of different body size’, behavioral ecology 23, 970–976. https://doi.org/10.1093/beheco/ars060 pettorelli, n., safi, k. & turner, w., 2014, ‘satellite remote sensing, biodiversity research and conservation of the future’, philosophical transactions of the royal society of london, series b, biological sciences 369, 20130190. https://doi.org/10.1098/rstb.2013.0190 robinson, j.c., hill, j.r.c. & rushworth, j., 1973, wankie national park, deka river catchment survey, zimbabwe parks and wildlife management authority, bulawayo, zimbabwe. rogers, k.m.l., 1993, a woody vegetation survey of hwange national park, zimbabwe parks and wildlife management authority, harare, zimbabwe. rose, r.a., byler, d., eastman, j.r., fleishman, e., geller, g., goetz, s. et al., 2015, ‘ten ways remote sensing can contribute to conservation’, conservation biology 29, 350–359. https://doi.org/10.1111/cobi.12397 sedano, f., gong, p. & ferrão, m., 2005, ‘land cover assessment with modis imagery in southern african miombo ecosystems’, remote sensing of environment 98, 429–441. https://doi.org/10.1016/j.rse.2005.08.009 sexton, j.o., song, x.p., feng, m., noojipady, p., anand, a., huang, c. et al., 2013, ‘global, 30-m resolution continuous fields of tree cover: landsat-based rescaling of modis vegetation continuous fields with lidar-based estimates of error’, international journal of digital earth 6, 427–448. https://doi.org/10.1080/17538947.2013.786146 sourcepole, 2014, openlayers plugin v1.3.4, open source geospatial foundation project, viewed 01 november 2014, from http://qgis.osgeo.org stehman, s. & czaplewski, r., 1998, ‘design and analysis for thematic map accuracy assessment: fundamental principles’, remote sensing of environment 64, 331–344, viewed 04 march 2003, from http://www.sciencedirect.com/science/article/pii/s0034425798000108 stehman, s.v. & wickham, j.d., 2011, ‘pixels, blocks of pixels, and polygons: choosing a spatial unit for thematic accuracy assessment’, remote sensing of environment 115, 3044–3055. https://doi.org/10.1016/j.rse.2011.06.007 the government of the republic of angola, the government of the republic of botswana, the government of the republic of namibia & the government of the republic of zimbabwe, 2011, kavango-zambezi transfrontier conservation area, p. 56, viewed 10 february 2015, from https://docs.google.com/viewer?url=https://www.kavangozambezi.org/index.php/en/publications/6-kaza-tfca-treaty/download?p=1 turner, w., rondinini, c., pettorelli, n., mora, b., leidner, a.k. et al., 2015, ‘free and open-access satellite data are key to biodiversity conservation’, biological conservation 182, 173–176. https://doi.org/10.1016/j.biocon.2014.11.048 valeix, m., fritz, h., sabatier, r., murindagomo, f., cumming, d. & duncan, p., 2011, ‘elephant-induced structural changes in the vegetation and habitat selection by large herbivores in an african savanna’, biological conservation 144, 902–912. https://doi.org/10.1016/j.biocon.2010.10.029 valeix, m., loveridge, a.j., davidson, z., madzikanda, h., fritz, h. & macdonald, d.w., 2009, ‘how key habitat features influence large terrestrial carnivore movements: waterholes and african lions in a semi-arid savanna of north-western zimbabwe’, landscape ecology 25, 337–351. https://doi.org/10.1007/s10980-009-9425-x vincent, r.k., 1972, ‘an erts multispectral scanner experiment for mapping iron compounds’, in proceedings of the 8th international symposium on remote sensing of environment, pp. 1239–1247, viewed 01 september 2017, from http://adsabs.harvard.edu/abs/1972rse..conf.1239v wiens, j., sutter, r., anderson, m., blanchard, j., barnett, a., aguilar-amuchastegui, n. et al., 2009, ‘selecting and conserving lands for biodiversity: the role of remote sensing’, remote sensing of environment 113, 1370–1381. https://doi.org/10.1016/j.rse.2008.06.020 wulder, m.a., masek, j.g., cohen, w.b., loveland, t.r. & woodcock, c.e., 2012, ‘opening the archive: how free data has enabled the science and monitoring promise of landsat’, remote sensing of environment 122, 2–10. https://doi.org/10.1016/j.rse.2012.01.010 appendix 1 table 1-a1: images used for creating study area mosaics. appendix 2 figure 1-a2: simple random sampling points used in accuracy assessment and visible burn scars. landsat 8 operational land imager (oli) image, bands rgb654, of the late 2013 dry season showing the 500 randomly distributed validations points (white dots), recent burn scars (indicated by red arrows), what is probably an agricultural area (purple arrow), and the location of small town of dete used as an example of unclassified condition in figure 2 (tiny red cross at edge of the north-east boundary of hwange national park). ryan.qxd activity patterns of african buffalo syncerus caffer in the lower sabie region, kruger national park, south africa s.j. ryan and w. jordaan ryan, s.j. and w. jordaan. 2005. activity patterns of african buffalo syncerus caffer in the lower sabie region, kruger national park, south africa. koedoe 48(2): 117–124. pretoria. issn 0075-6458. the activity budgets of three herds of african buffalo in the lower sabie region of kruger national park, recorded between 1991 and 1992 were quantified to examine both nocturnal and seasonal effects on feeding activity and 24-hr movement in an area of dense bushveld. we found that the average 24-hr distance traveled by herds (3.35 km) was shorter than that found in other studies and that there was no seasonal effect on this distance, which we attribute to ready availability of water in both seasons. we found that the buffalo spent a similar amount of time feeding (9.5 hrs) as in other studies, but that the proportional feeding and resting time was influenced by the time of day and the season. these herds spent proportionally more time feeding at night (44.5 % vs 32.0 %) and more time resting in the day (28.4 % vs 16.0 %). in addition, they appeared to rest more during the day and feed more at night during the wet season, when it was hotter. keywords: african buffalo, syncerus caffer, activity budgets, kruger national park s.j. ryan, department of environmental science, policy & management, 137 mulford hall, university of california at berkeley, berkeley, ca 94720, usa (sjryan@nature.berkeley.edu) (present address: museum of vertebrate zoology, 3101 valley life sciences building, university of california at berkeley, berkeley, ca 94720, usa); w. jordaan, 3 conservation services, sanparks, private bag x402, skukuza, 1350 republic of south africa. issn 0075-6458 117 koedoe 48/2 (2005) introduction mammalian herbivores spend the majority of their time feeding; as beekman & prins (1989) pointed out, horses spend more than 70 % of their time grazing and cattle, 50–60 %. ruminant ungulates must additionally allocate time to ruminating and resting. in savanna ecosystems, ruminants must not only obtain sufficient quality and quantity of food, but may also need to adjust their activity budget as seasonal changes constrain when the activity can take place in a 24-hour period. african buffalo syncerus caffer are large grazers, whose feeding ecology predicts greater selection of high biomass (owensmith 1988; redfern et al. 2003) than high quality grass swards. additionally, redfern et al. (2003) have shown empirically that buffalo select for areas close to water sources in kruger national park. thus the feeding and ranging behavior of buffalo is subject to constraints such as the abundance of grass and the availability of water. in previous studies, seasonal changes in vegetation quality and water availability have been shown to alter both ranging and feeding habits of buffalo (funston et al. 1994). in this study, we describe activities and behaviors of three breeding herds of buffalo in a semi-arid wooded savanna environment. this study contrasts with prior studies of buffalo which tend to focus on diurnal behaviour in open savanna habitats (funston et al. 1994; prins 1996; sinclair 1977; winterbach & bothma 1998), as herds were tracked in relatively dense bushveld. moreover, this study comprises 24-hour samples, meaning that nocturnal behavior of buffalo was also quantified. in addition, this data was collected during a severe drought period, such that there was less rainfall occurring than in prior studies at similar latitude (funston et al. 1994). buffalo groups were followed in the lower sabie region of kruger national park, south africa. tracking data from radio-collared individuals and visual observations were used to describe daily ranging activity, and scan sampling was used to describe daily herd activity between 1991 and 1992. differences in activity budgets, timing of activity during the day and ranging observed in buffalo are hypothesised to arise due to seasonal and habitat differences (funston et al. 1994). we explore both diurnal and nocturnal activities, seasonal differences and compare this study with others in different habitats. study site the data used in this study were collected in the lower sabie region of kruger national park (fig. 1). the study area spans two ecozones: knobthorn/marula savanna and lebombo bushveld (gertenbach 1983). the data was collected in an area bordered at (31.90°, -25.11°)–(32.00°, -25.35°); all data was recorded in the wgs84 datum. this region receives 400–500 mm of rain a year on average, and the rainfall and temperature follow a unimodal distribution, which leads to a warm wet season (oct–mar) and a cool dry season (apr–sep) (gertenbach 1980). methods herd activity data data was collected on three herds that were tracked using radio telemetry and visual observations. the herds were estimated at approximately 400, 200 and 250 animals, respectively. fifteen separate focal samples of activity were taken between march 1991 and august 1992. seven of these samples were ≥24 hrs in duration and eight were ≥11 hrs of daytime observations. main herd activity was sampled at 5-minute intervals during the day and night, where possible. herd activities recorded were: 1. feeding stand/walk and eat 2. standing standing with head up and ruminating 3. resting standing or lying with head bent downwards, not eating 4. moving walking but not grazing 5. drinking drinking water or standing < 10 m from water. koedoe 48/2 (2005) 118 issn 0075-6458 fig. 1. the lower sabie region of kruger national park. detail shows locations of observations (asterisks) on a coarse (5 km) grid; the ecozones and three perennial river courses are also shown. herd movement data using radio telemetry, locations of herds during 40 separate days were collected over the study period (dry season, n = 24 days; wet season, n = 16 days). these locations were used to calculate the distance traveled by the herd during each day’s sample. due to sampling time differences (range: 13–29 hrs), the distance estimate from each data collection was multiplied by 24÷(observation time (hrs)) to estimate a 24-hr distance. as this is simply a difference in location over an elapsed time, this represents a minimum distance estimate. we pooled all herds and types in the study to obtain an average daily movement estimate. these estimates were compared by season using a welch modified t-test for unequal variances. herd activity to examine diurnal activity of buffalo herds, we cropped the 24-hr samples to 12 hrs and analysed all 15 samples together. due to unequal observation times, we converted hours of activity into proportions of the total. an anova with post-hoc means tests (tukey-kramer’s honest significant difference test, α = 0.05, table 1) was used to determine both the order of times allocated to each activity and which activity occupied significantly more of the time than others. we also used the 24-hr sample data to quantify the mean time spent by the herd in each activity during the full day (table 2). we used the longer activity budgets to compare diurnal and nocturnal activity budgets of buffalo. due to differences in the hour of sunset in the datasets, we chose 19:00 (7 pm) as the division in the dataset. welch modified anovas for unequal variances were used to compare means. significant results were further tested to examine whether differential activity might occur as a result of seasonality. activity timing was quantified by pooling all herd activity budgets. the proportion of observations within each 5-minute scan corresponding to activities 1–5 were plotted against time (fig. 4). activity periods were defined for activities 1–3 as periods of time when the proportion exceeded 20 %. for activities 4 and 5, the activity periods were both of short duration and infrequent in occurrence, such that a single herd engaging in the activity might comprise the entire occurrence. furthermore, this might appear as five separate observations within a few hours and thus appear to be a repeated low proportion activity. we took clusters of activity observations, rather than the proportion to represent activity periods and descriptively reported the timing of these activities. results herd movement data the mean 24 hr estimated minimum distance traveled by all herds pooled was 3.35 ± 0.35 km (x ± se). a welch modified t-test did not detect a significant seasonal difference in issn 0075-6458 119 koedoe 48/2 (2005) table 1 an anova and tukey-kramer’s honest significant difference (hsd) post-hoc means test of the diurnal activity budgets of buffalo herds; the percent of observation time in each sample with the standard error (se) and the conversion of these means and errors into hours are included activity hsd* % observation approx time (se) hours (se) feeding a 37.47 (2.62) 4.50 (0.31) standing b 28.78 (2.14) 3.45 (0.26) resting b 25.86 (2.71) 3.10 (0.33) moving c 5.22 (1.56) 0.63 (0.19) drinking c 2.66 (0.64) 0.32 (0.08) *activities with different letters have significantly different means anova: f = 54.07, df = 4, p = 0.0001 table 2 an anova and tukey-kramer’s honest significant difference (hsd) post-hoc means test of the 24-hr activity budgets of buffalo herds; the percent of observation time in each sample with the standard error (se) and the conversion of these means and errors into hours are included activity hsd* % observation approx time (se) hours (se) feeding a 39.41 (2.21) 9.46 (0.53) standing b 31.49 (1.69) 7.56 (0.40) resting c 21.45 (2.36) 5.15 (0.57) moving d 4.76 (0.90) 1.14 (0.22) drinking d 2.88 (0.51) 0.69 (0.12) *activities with different letters have significantly different means anova: f = 90.21, df = 4, p = 0.0001 the 24-hr distance estimates (wet: 3.14km, dry: 3.49 km; t = 0.4547, df = 26.022, p = 0.6531). herd activity analysis of the diurnal activity of all herds showed that significantly more time was devoted to activity 1, i.e., feeding, than other activities. the herds spent an average of 37.5 % or 4.5 hrs of the day actively feeding, and another 28.8 % (approx. 3.5 hrs) to standing and ruminating; and resting took another 25.9 % (approx. 3.1 hrs) of the day. this shows us that 75 % or 9 hrs of the day is devoted to obtaining and ruminating food (table 1). using the seven 24-hr samples, we found a similar breakdown of activity pattern, but the relative proportion of time spent feeding (1) and standing (2) increased and the other activities showed a corresponding decrease in proportions (see table 2). when we compared diurnal and nocturnal components of the herd activity budget, we found that the buffalo spent a significantly greater proportion of time feeding (#1) during the night [44.5 % vs 32.0 % (f = 9.85, df = 1, p = 0.009)]; and a trend towards a greater proportion of time resting (#3) during the day [28.4 % vs 16.0 % (f = 3.61, df = 1, p = 0.08)]. we found no significant proportional differences in diurnal and nocturnal time budget for other activities (fig. 2). due to low sample size, we could not detect seasonal effects on overall activity budgets. however, in fig. 3 there is a trend suggesting that buffalo spend more time feeding during the dry season than the wet, more time moving around and less time resting. although the sample size was too small for rigorous testing, we sought to find seasonal differences in diurnal and nocturnal behaviour. the difference between day and night feeding and resting activities was not statistically significant, although the data suggest a trend toward a greater proportion of nocturnal feeding and resting in the wet season (feeding (#1), dry season: f = 2.61, p = 0.16; wet season: f = 8.48, p = 0.07; resting (#3), dry season f = 0.21, p = 0.67; wet season: koedoe 48/2 (2005) 120 issn 0075-6458 fig. 2. relative proportions of time allocated by the buffalo herds to activities, diurnal (white) and nocturnal (black) components, with their standard errors (se). fig. 3. relative proportions of time allocated by the buffalo herds to activities, dry season (white) and wet season (black) components, with their standard errors (se). f = 6.31, p = 0.07). all other potential seasonal effects were not statistically significant. figure 4 shows the relative proportions of each activity in each 5-minute interval in the upper panel; in the lower panel we show the corresponding number of observations in each 5-minute interval. due to differences in the start and end time of the diurnal observation sessions, the number of observations within the 5-minute time intervals increase in the early morning and decrease towards evening. the nocturnal observation sesissn 0075-6458 121 koedoe 48/2 (2005) fi g. 4 . r el at iv e pr op or tio ns o f ob se rv at io ns o f ea ch a ct iv ity ( se e m et ho ds ) in e ac h 5m in ut e in te rv al ( up pe r pa ne l) a nd th e co rr es po nd in g nu m be r of o bs er va tio ns in e ac h 5m in ut e in te rv al ( lo w er p an el ). t he 2 4ho ur c yc le r un s fr om 7 am -7 am ; w e m ar k 7p m b ot h as a r ef er en ce f or th e on se t o f no ct ur na l a ct iv ity a nd a s a no te o f th e re du ct io n in n um be r of o bs er va tio ns . sions, although fewer in number, have a consistent number of observations per scan. the 24-hour cycle runs from 7am-7am; we marked 7 pm both as an indicator of the onset of nocturnal activity and of the reduction in number of observations. we found that the buffalo had two defined feeding (#1) periods during the day and into the night (approximately 5:30-10:30; peak: 9:15 and 15:00-21:00; peak: 17:10), and less defined but moderate continuous feeding during the night hours and into the dawn. we see that standing (#2), contributes a large proportion of the observations during and following the feeding bouts (8:15-13:50 and 18:00-24:00), and as feeding does, it continues at moderate levels throughout. resting (#3) has a clear daytime peak, with a major bout occurring from 12:00-16:00, and another moderate bout occurring in the early morning (3:006:00). moving (#4) appears to occur in two clusters; one in the mid-morning to early afternoon between 10:00 and 13:00 and another cluster in the evening to night at 19:00-22:00; drinking (#5) occurs in an early morning cluster around 6:00-8:00 and another mid-morning cluster from 10:00-12:00; a couple of observations occur later in the day also and two toward the middle of the night. discussion in this study, the estimated minimum distance that the buffalo herds traveled in 24 hrs was 3–4 km. this did not change between seasons, suggesting that these buffalo have high site fidelity, perhaps due to a combination of the constraint of artificial water availability and the presence of reliable grazing resources. in cameroon, stark (1986) found that buffalo moved around 7 km in the wet season and 5.6 km in the dry; grimsdell & field (1976) reported 9.6 km on average in rwenzori national park, uganda; conybeare (1980) reported 6.1 km on average in the dry season in sengwa wildlife research area, zimbabwe. funston et al. (1994) found that in the early wet season the herds in the sabie sand reserve (on the western edge of kruger national park) tended to remain in an area around a watering hole for up to eight days and then move. they noted also that in the dry season, the herd would make long distance (10 km) movements at night in search of better grazing. the present study took place in a more closed habitat than a majority of these prior studies, and the high density of artificial water availability in knp (brits et al. 2002) might reduce the need for wide ranging in this habitat. moreover, there are three perennial river courses running through this study area (see fig. 1); and buffalo herds tend to maintain smaller home ranges where there are perrennial rivers hunter (1996). as this study was conducted during a drought period, these river courses may have dried out and served as areas retaining green grass swards, a reliable source of grazing, but not providing sufficient drinking areas. thus, the buffalo may have been constrained by artificial water presence instead and behaved similarly to the study animals of funston et al. (1994). we cannot compare our measure directly with the daily tracking data of prior studies, as it is not representative of a full grazing path route, rather the shortest distance between radio tracking locations. however, it provides us with a range approximation for the area. we found that feeding comprised around 9.5 hrs (39.41 %) on average of the 24-hr activity budget (table 2). this falls within the range for grazing found by both grimsdell & field (1976) (9 hrs) and sinclair’s (1977) bachelor bull study in the serengeti (9.7-10.1 hrs). this compares with the 40 % estimated by winterbach & bothma (1998). we found that across both seasons, the buffalo herds appeared to spend a greater proportion of time feeding at night than during the day, similar to the buffalo at lake manyara, tanzania (beekman et al. 1989) and as seen by sinclair (1977) in the serengeti, although his observations occurred in the wet season, not the dry. stark (1986) estimated that during dry season, his study herd of savanna buffalo (syncerus caffer brachyceros) grazed for an average of 3.1 hrs (~26 %) in a 12-hr period. this is koedoe 48/2 (2005) 122 issn 0075-6458 considerably less than the average for our study herds. in addition, as suggested by sinclair (1977), we saw a trend toward more time spent feeding in the dry season than the wet, perhaps in response to lower vegetation quality and quantity. we also saw a trend toward more time moving in the dry season, which may reflect searching for feeding patches; concomitant with this was less resting, perhaps a repercussion of spending more time looking for food. we saw that the buffalo tended to spend time feeding and standing in succession (fig. 4), with standing and ruminating occupying around 30 % of observed activity time, both diurnally and nocturnally (tables 1 & 2). while this provided time for rumination, it is likely a combination of both rumination and vigilance behaviour. buffalo exhibit standing, raised head vigilance; it is plausible that they are not only using visual cues, but also olfactory. we see that this activity occurs in greater proportion at night (fig. 2), although this is not statistically significant. we posit that with a reduction in visibility at night more vigilance will be necessary. as this data was collected in non-exclusive categories, we can assume that the proportion of time spent being vigilant is higher than simply the time allocated to the standing (#2) category. buffalo were recorded as resting (#3) while standing, if they were not feeding. this suggests that vigilant behaviour may comprise a significant proportion of buffalo activity. we found that the buffalo grazed in two major daytime periods, which is consistent with the findings of winterbach et al. (1998). we also found that they continued to graze moderately throughout the night. we saw that buffalo tended to rest more during the day than at night, particularly during the warm wet season. as found in other studies (stark 1986; winterbach et al. 1998), the herds had a long mid-day rest period, after the morning grazing peak as shown in fig. 4. sinclair (1977) showed that grazing activity ceased completely in buffalo at a certain elevated ambient temperature. thus we would expect the buffalo to cease grazing and to rest when the temperature is hottest—midday in the warm wet season. we found that the buffalo herds had two main periods of diurnal drinking, which occurred in the early morning and midmorning, after a grazing bout and while ruminating. there appears to be more time devoted to drinking during the day than at night, but we found no evidence to suggest that they spent more time drinking during the dry season than the wet. previous studies have suggested that buffalo only need to drink once a day (e.g. sinclair 1977). winterbach & bothma (1998) found that buffalo drank in the early afternoon, whereas grimsdell & field (1976) found that they drank in the mid-morning. since we did not distinguish between drinking and standing near water, it is not clear how often these herds drank each day. this system has a high density of available water sources, so buffalo may drink more than once in a day. qualitative observations (wj) suggest that buffalo will stand in the water, but only a few herd members will drink at one time. future directions this data represents only 15 sampling periods in total, which was not sufficient to demonstrate seasonal differences in buffalo herd activity budgets, but provided a baseline to inform future data collection. in future studies, we would suggest additional observation sessions, spread more evenly across a year to encompass both seasons appropriately. this study presented an opportunity to refine activity categories; in particular, the overlap in potentially important behaviours between categories, such as vigilance, can inform category definitions. the categories of activity such as standing and resting were both likely during ongoing rumination, but this distinction made it harder to compare with prior studies. in the future, the addition of samples across the year will provide more detailed comparisons and will allow further exploration of the flexibility of activity budgets as buffalo adjust to seasonal changes. the interest in issn 0075-6458 123 koedoe 48/2 (2005) nocturnal activity suggests that additional long observation sessions such as these will provide valuable information regarding buffalo ecology. furthermore, with the advent of more sophisticated tracking devices, such as gps collars, the combination of geographic locations and activity observations can be better used, in conjunction, to understand buffalo feeding ecology. acknowledgements thank you to kruger national park’s scientific services for permission to carry out this study. thanks to a.g. vaughan for help with data formatting, and to three reviewers for comments and edits. s.j. ryan’s research was supported by epa-star fellowship fp-916382. references beekman, j.h. & h.h.t. prins. 1989. feeding strategies of sedentary large herbivores in eastafrica, with emphasis on the african buffalo, syncerus caffer. african journal of ecology 27: 129–147. brits, j., m.w. van rooyen & n. van rooyen. 2002. ecological impact of large herbivores on the woody vegetation at selected watering points on the eastern basaltic soils in the kruger national park. african journal of ecology 40: 53–60. conybeare, a. 1980. buffalo numbers, home range and daily movement in the sengwa wildlife research area, zimbabwe. south african journal of wildlife research 10: 89–93. funston, p.j., j.d. skinner, & h.m. dott. 1994. seasonal-variation in movement patterns, homerange and habitat selection of buffalos in a semiarid habitat. african journal of ecology 32: 100–114. gertenbach, w.p.d. 1980. rainfall patterns in the kruger national park. koedoe 23: 35–44. gertenbach, w.p.d. 1983. landscapes of the kruger national park. koedoe 6: 9–122. grimsdell, j.j.r. & c.r. field. 1976. grazing patterns of buffaloes in the rwenzori national park, uganda. east african wildlife journal 14: 339–344. hunter, c.g. 1996. land uses on the botswana/ zimbabwe border and their effects on buffalo. south african journal of wildlife research 26: 136–150. owen-smith, r.n. 1988. megaherbivores: the influence of very large body size on ecology. cambridge: university press. prins, h.h.t. 1996 ecology and behaviour of the african buffalo: social inequality and decision making. new york, ny.: chapman & hall. redfern, j.v., r. grant, h. biggs, & w.m. getz. 2003. surface-water constraints on herbivore foraging in the kruger national park, south africa. ecology 84: 2092–2107. sinclair, a.r.e. 1977. the african buffalo: a study of resource limitation of populations. chicago: university of chicago press. stark, m.a. 1986. daily movement, grazing activity and diet of savanna buffalo, syncerus caffer brachyceros, in benoue-national-park, cameroon. african journal of ecology 24: 255–262. winterbach, h.e.k. & j. du p. bothma. 1998. activity patterns of the cape buffalo syncerus caffer caffer in the willem pretorius game reserve, free state. south african journal of wildlife research 28: 73–81. koedoe 48/2 (2005) 124 issn 0075-6458 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false /preserveoverprintsettings true /startpage 1 /subsetfonts true /transferfunctioninfo /apply /ucrandbginfo /preserve /useprologue false /colorsettingsfile () /alwaysembed [ true ] /neverembed [ true ] /antialiascolorimages false /downsamplecolorimages true /colorimagedownsampletype /bicubic /colorimageresolution 300 /colorimagedepth -1 /colorimagedownsamplethreshold 1.50000 /encodecolorimages true /colorimagefilter /dctencode /autofiltercolorimages true /colorimageautofilterstrategy /jpeg /coloracsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /enu (use these settings to create pdf documents with higher image resolution for high quality pre-press printing. the pdf documents can be opened with acrobat and reader 5.0 and later. these settings require font embedding.) /jpn <feff3053306e8a2d5b9a306f30019ad889e350cf5ea6753b50cf3092542b308030d730ea30d730ec30b9537052377528306e00200050004400460020658766f830924f5c62103059308b3068304d306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103057305f00200050004400460020658766f8306f0020004100630072006f0062006100740020304a30883073002000520065006100640065007200200035002e003000204ee5964d30678868793a3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002> /fra <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> /deu <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> /ptb <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> /dan <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> /nld <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> /esp <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> /suo <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> /ita <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> /nor <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> /sve <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> >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice article information authors: sam ferreira1 andrew deacon1 hendrik sithole1 hugo bezuidenhout2 mahlomola daemane2 marna herbst2 affiliations: 1savanna research unit, south african national parks, skukuza, south africa 2arid research unit, south african national parks, kimberley, south africa correspondence to: sam ferreira email: sam.ferreira@sanparks.org postal address: private bag x402, skukuza 1350, south africa dates: received: 18 apr. 2010 accepted: 02 feb. 2011 published: 13 may 2011 how to cite this article: ferreira, s., deacon, a., sithole, h., bezuidenhout, h., daemane, m. & herbst, m., 2011, ‘from numbers to ecosystems and biodiversity: a mechanistic approach to monitoring’, koedoe 53(2), art. #998, 12 pages. doi:10.4102/koedoe.v53i2.998 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) from numbers to ecosystems and biodiversity: a mechanistic approach to monitoring in this essay... open access • abstract • introduction • the sanparks context • thresholds of potential concern revisited    • linking thresholds of potential concerns • rare, endemic and cryptic species • selected species abundances, trends and impacts • ecosystem indicators: ecological, community-based assessments • the generality of monitoring across parks • conclusions • references abstract (back to top) diverse political, cultural and biological needs epitomise the contrasting demands impacting on the mandate of the south african national parks (sanparks) to maintain biological diversity. systems-based approaches and strategic adaptive management (learn by doing) enable sanparks to accommodate these demands. however, such a management strategy creates new information needs, which require an appropriate analytical approach. we use conceptual links between objectives, indicators, mechanisms and modulators to identify key concerns in the context of and related to management objectives. although our suggested monitoring designs are based mostly on defined or predicted underlying mechanisms of a concern, sanparks requires inventory monitoring to evaluate its key mandate. we therefore propose a predictive inventory approach based on species assemblages related to habitat preferences. inventories alone may not always adequately serve unpacking of mechanisms: in some cases population size needs to be estimated to meet the information needs of management strategies, but actual population sizes may indirectly affect how the species impact on other values. in addition, ecosystem objectives require multivariate assessments of key communities, which can be used in trend analysis. sanparks therefore needs to know how to detect and define trends efficiently, which, in turn, requires precision of measures of variables. conservation implications: current research needs with regard to monitoring should focus on defining designs to yield optimal precision whilst taking methodology, survey trade-offs and analytical approaches into account. use of these directives and research will guide monitoring during evaluation of sanparks objectives at various scales. introduction (back to top) monitoring the performance of protected areas on several scales is a common conservation challenge worldwide (noss 1990; noon, spies & raphael 1999; possingham et al. 2001; woodroffe & ginsberg 1998) and has been met with some success (bruner et al. 2001; knight et al. 2006; salafsky & margoluis 1999) and some failure (curran et al. 2004; liu et al. 2001). in many instances, monitoring the performance of conservation agencies with regard to achieving their key mandate of maintaining biodiversity has been, at best, haphazard in the past (mcgeogh et al. 2011). this may stem from (1) mechanisms (i.e. the biological effects of a factor and modulators in the system) being poorly defined, which may threaten conservationists’ achieving objectives, (2) ill-defined or nonexisting objectives of monitoring plans, (3) uncertainties regarding indicators (i.e. biological indicators of specific environmental conditions) and measurement parameters to address specific objectives, and (4) uncertainties and disagreements regarding measurement of response as well as explanatory variables (lindenmayer & likens 2009). in addition to nonrobust approaches to monitoring, political, cultural and religious beliefs influence conservation approaches (brandon, redford & sanderson 1998; doyle & mceachern 2008; lawton 1997; martello 2001). these contrasting demands force modern-day conservationists to embrace complexity as a leading value to recognise the social, economic and ecological interactions that epitomise cultural, historical and natural heritage (bruner et al. 2001; ghimire & pimbert 1997; scoones 1999). in such instances conservationists are vulnerable to effects of shifting conceptual approaches (e.g. lewis 1996; lindenmayer & likens 2009; morellet et al. 2007) that create confusion or direct conservation management into areas that have not been scientifically defined yet. the recognition of complexity requires a management style based on a systems approach (brussard, reed & tracy 1998; brandon et al. 2005; de leo & levin 1997; kay & schneider 1994). for example, because of the complexity of biodiversity, surrogates such as subsets of species, species assemblages and habitat types were proposed as measures of biodiversity. the locations of these surrogates within areas have to be plotted to allow an estimated comparison between the areas (margules & pressey 2000). whether correct or not, such styles impose information needs that exceed the traditional requirements on which conservationists base management decisions. ecosystem management is a topical systems paradigm (bocking 1994). the approach can allow for appropriate management of biological diversity of ecosystems, species and genetic resources – the primary mandate of conservation agencies – within the constraints of tourist, stakeholder and economic expectations (lindenmayer, franklin & fischer 2006; mcneely 1994; slocombe 1998). ecosystem management involves two essential elements, namely integration of scientific knowledge and maintenance of evolutionary potential (bocking 1994; parrish, braun & unnasch 2003; rodrigues et al. 2004; sale et al. 2005; simberloff 1999). the latter focuses on facilitating, maintaining or restoring ecological processes (e.g. wassenaar et al. 2005; wassenaar, ferreira & van aarde 2007). this can be addressed, to a large extent, by strategic adaptive management approaches (biggs & rogers 2003; tompkins & adger 2004): conservationists use knowledge of a system’s underlying mechanisms to develop a management strategy that predicts relevant system responses, robustly assess the outcomes, and then adapt management actions accordingly if needed (rogers 1997). protected areas are seldom pristine (mcneely 1994). they are either limited owing to features that impede spatial and temporal variability in ecological processes (e.g. du toit, rogers & biggs 2003) or degraded owing to a history of disturbed ecological processes often due to agricultural activity (agardi 2000; braaz 1992; noss 1999). this challenges traditional monitoring programmes directed at supporting management decisions focused on species (e.g. regan et al. 2008) and to some extent future monitoring may still have such a focus. in addition, only restorative actions can attempt to maintain evolutionary potential when large sections of the biota are locally extinct and/or have been replaced by exotic organisms (e.g. wassenaar et al. 2005), but this requires very different measures or interpretation of existing approaches. in this paper we provide a conceptual framework for defining monitoring parameters. we first establish the context specific to south african national parks (sanparks), especially as it pertains to adaptive management. we then challenge the static nature of thresholds of potential concern as applied at present and suggest a revised approach that flags potential ecosystem degradation through the development of a mechanisms-based unpacking of an ecological concern. the approach allows us to illustrate how information needs, albeit diverse, can have a common strategic design. the sanparks context (back to top) biological diversity results from a variety of processes and interactions on ecological and evolutionary time scales (wilson 1988). human activities and the organisms that travel with them (animal pests and weeds) have changed biological processes throughout the world (chapin et al. 2000; daszak, cunningham & hyatt 2000) and are collectively referred to as human-induced disturbance. a systems-based approach to improving the capacity of sanparks to achieve its key mandate requires the management of human-induced disturbances. according to such an approach management should view ecosystem problems as components of an overall system and therefore not act only on specific aspects, outcomes or events in order to reduce the contribution to unintended consequences (ackoff 2010). the key mandate of sanparks requires conservationists to maintain biological diversity within its jurisdiction (knight 2007; varghese 2008). sanparks therefore has two primary needs to evaluate whether it is achieving its mandate, namely determining species status and ecosystem health within the parks. determining species status within sanparks is particularly important with reference to rare species defined as not abundant or only locally abundant (gaston 2008; preston 1948). evaluating conservation outcomes for such species when using traditional approaches to define population trends is challenging (mcardle 1990). measuring rare species, however, may not reflect much on ecosystem dynamics (gaston 2008; smith & knapp 2003), despite important contributions to them. the wellness of ecosystems can be assessed by considering less common species, which act as keystone species, augment efficacy (e.g. invasion resistance) of the community and/or play critical roles in ecosystem resilience (lyons et al. 2005). these measurements require multivariate approaches and techniques that have been applied only to a limited extent in the conservation and restoration environment (spellerberg 2005). sanparks uses thresholds of potential concerns (tpcs) to define a range of possible states for species populations or ecosystems (see biggs & rogers 2003; foxcroft 2009). this approach presents challenges because it requires definition of these states (e.g. milton et al. 1994), a process which is not formally established in the sanparks planning environment. in addition, the tpc approach requires various types of information, for example a definition of (1) species diversity, (2) population size, (3) temporal and/or spatial changes in populations and (4) an assessment of whether such changes are within the (established or predicted) tpc. several species-focused tpcs were defined for the kruger national park in 2002 (table 1), but few of these have been addressed since the inception of this approach. at the time, the large-scale herbivore and fire interaction research experiment (lashfire) programme was about to be launched (trollope et al. 1999) and elements of the programme were built into most of the research and monitoring of tpcs. after the suspension of the lashfire programme, species-focused monitoring, particularly of small vertebrates, was implemented in pilot studies of different vegetation types. however, during drafting the 2007 tcps (south african national parks 2007), the approach was used extensively to describe the concept of homogenisation, which was considered as a loss of dissimilarity (table 2). the purpose was to create an indicator of system function and associated biodiversity. however, these were never fully implemented owing to the intense and cumbersome monitoring required. many of the constraints with regard to implementation of tpcs stem from prioritisation and evaluation uncertainty. a conceptual approach that links objectives, indicators, mechanisms and modulators in an effort to identify key concerns in the context of management objectives (gaylard & ferreira 2011) and a range of states and transitions (westoby, walker & noy-meir 1989; rietkerk & van de koppel 1997) can greatly assist in the development of appropriate, focused and robust monitoring plans. in such instances, monitoring designs should be based on defined or predicted underlying mechanisms of a conservation concern. however, sanparks also requires inventory monitoring to evaluate its key mandate (biggs & rogers 2003). in this case a conceptual approach should use predictive inventory techniques (i.e. evaluations of observed versus predicted species presence), because few species lend themselves to easy estimation of abundance (regan et al. 2008). in several cases, though, species such as large herbivores (e.g. kruger, reilly & whyte 2008) do lend themselves to estimation of numbers. management may require such information when population sizes may indirectly affect how a species impact on other values (sanparks 2010), given that spatial mechanisms, rather than abundance, are primary drivers of large mammal effects on ecosystems (van aarde, jackson & ferreira 2006). in addition, evaluating ecosystem objectives require multivariate assessments of key communities (wassenaar et al. 2007). nearly all measurement types and sanparks objectives require trend analysis to allow efficient detection and definition of trends. this requires precision of each measure (gerrodette 1987). current research needs to facilitate appropriate monitoring should focus on defining designs that allow optimal precision with regard to methodology, survey trade-offs and analytical techniques. such directives can then inform and focus monitoring to evaluate the sanparks objectives at various scales. in this paper we therefore also highlight constraints of traditional approaches and explore particularly why many existing sanparks tpcs have not been addressed. we propose three key approaches and define likely measures of outcome success, which relate to a more process-based approach for defining tpcs. as a first step we revisit the applicability of tpcs and propose an alternative conceptual tpc. we then introduce an approach to deal with rare, cryptic and difficultly sampled taxa. we evaluate the value of species abundance and trends, but also highlight the constraints. in the final step, we introduce ecosystem outcome indicators. based on these discussions we provide conceptual examples of park monitoring programmes directed at assessing the conservation outcomes of maintaining species and communities that are affected by large herbivores. large herbivores are the focus for this conceptual discussion because they often form a key part of the conservation management decisions of sanparks. table 1: thresholds of potential concern, with a species focus, as constructed for the kruger national park in 2002. table 2: thresholds of potential concern, with a species focus or components with a species focus, as revised for the kruger national park in 2007.a thresholds of potential concern revisited (back to top) the tpc approach adopted by sanparks (biggs & rogers 2003) uses a range of possible ecosystem states as guidelines. this acknowledges that ecosystems are in flux and tend to vary across space and time (biggs & rogers 2003). the challenge is to define acceptable variation. sanparks can expect that ecosystems will be bounded and may actually have some form of equilibrium dynamics. this is because different areas provide different environmental limitations (belsky 1990; john et al. 2007; walker 1987), species have physiological limits (weiner 1992) and they respond to other species (agrawal 2001). such limitations mean that a small, finite number of possible states exist (stringham, krueger & shave 2003). conservationists may consider systems to reflect the ‘balance’ versus the ‘flux’ of nature (pimm 1991), which depends on the temporal and spatial scale perceived, whether assumed or enforced (figure 1). the reality is that conservationists, irrespective of the paradigm they adhere to, guide their management strategies such that a system can adopt any state as long as it does not lose a key property. given that diversity and ecosystem resilience are associated with heterogeneity (christensen 1997), should sanparks manage for heterogeneity? if so, what would be an acceptable level of flux? the mechanisms underlying development and maintenance of heterogeneity give some direction (figure 2). agents that act on a specific feature to effect change are not the same as those factors that control and maintain the change once it has taken place. the balance paradigm changes controllers to agents and vice versa, which may force equilibrium values over scales that are inappropriate. the outcomes are likely to contrast conservation objectives. the realisation that the balance paradigm challenges achieving common biodiversity objectives means that sanparks needs measures and tpcs that reflect outcomes of heterogeneity. the heterogeneity paradigm is sensibly adopted by sanparks (du toit et al. 2003). for systems that are relatively intact, setting tpcs is a rational way of defining outcomes which a monitoring programme should evaluate and inform (e.g. foxcroft 2009). this is advanced by the fact that the number of states is limited and that the range depends on the biome and system responses to disturbances. for example, arid systems may change little in the face of disturbance (i.e. they are resistant) whereas more mesic systems may recover quickly in the face of disturbance (i.e. they are resilient), as described by pimm (1991). however, most of the areas that sanparks manages are conceivably degraded. many have an agricultural history prior to proclamation, or have an intense management history that embraced balance-paradigm approaches. within a heterogeneity paradigm the challenge is to define something as degraded as well as determining how such a system can be restored. in reality, sanparks cannot define the ’natural’ state and is left with providing opportunities for ecological processes to play out through restoration or, if impossible, mimic outcomes. the traditional use of tpcs to define a range of states may carry little value in such cases. sanparks thus needs a conceptual approach to set tpcs for degraded areas, which can then direct monitoring programmes within the constraints highlighted earlier. such tpcs may formally be defined as targets. degraded systems can be considered when a strong driver has changed the system to such an extent that the system has lost function, diversity or structure (aradóttir & arnalds 2001; hüttl & schneider 1998). driver changes are often human induced (aradóttir & arnalds 2001). subsequent restoration involves human activities directed at instituting a set of processes to guide the system towards the range of acceptable ecosystem states (society for ecological restoration 2004). it means that a tpc needs measures with direction and trends, not just a range of states. such measures and tpcs will be constrained by the limitations of detecting trends, much as the detection of changes in species abundance is constrained by several trade-off factors (gerrodette 1987). although restoration constitutes the process of assisting recovery, success ultimately depends on whether populations, communities and ecological functions attain limits typical of nondegraded reference systems (wassenaar et al. 2007). with evidence increasing that removing stressors is not always sufficient to guarantee restoration success, the true arbiters will be the organisms and processes at which restoration is aimed. stringent appraisal of these ecological criteria will be required if restored systems are to conserve biodiversity and deliver ecosystem services (ormerod 2003). as argued thus, sanparks can develop a generalisation of tpc. states are limited because of restricted species pools, specific niche needs and constrained species coexistence due to species interactions (wassenaar et al. 2005). however, the ‘natural state’ is unknown and is most likely dynamic (pimm 1991). for this very reason, the management objective should be simple: to provide opportunities for spatial and temporal variability in processes that will lead to dynamic states. in this context state-transition models (e.g. stringham et al. 2003) may hold great value in helping to prioritise which transitions are undesirable (usually human induced – see chapin et al. (2000) and daszak et al. (2000)) in the context of set objectives and allow conservationists to define tpcs to indicate such transitions or changes. measuring state change can be applied to intact or recovering degraded systems. in both intact and degraded cases thresholds reflect spatial variability through a distribution of changes, and temporal variability through a distribution of variance in changes. for example, changes in vegetation may be considered nondetrimental if study sites within landscapes comprising specific areas have different rates of change, that is, spatial heterogeneity. for intact systems, conservationists should anticipate that this distribution of rates of change centres on zero, whilst for degraded systems the distribution should centre on a value not equal to zero (biggs et al. 2011). setting tpcs according to this approach requires substantial assessment of trends at all levels. the guideline above is thus relevant to support sanparks fully in evaluating outcomes of management actions directed at achieving various objectives. figure 1: outcomes of the balance versus flux paradigms of ecosystem dynamics. figure 2: interactions between agents (e.g. elephants) that act on a substrate (e.g. woodlands) to change it from one state to an alternative state (e.g. shrublands). however, the maintenance of the shrublands state is controlled by something else (e.g. kudu, impala and mice) other than the agent (adapted from pickett, t.a., cadenasso, m. & benning, t., 2003, ‘biotic and abiotic variability as key determinants of savannah heterogeneity at multiple spatiotemporal scales’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 22–40, island press, washington dc). several other aspects of an ecosystem may respond (e.g. birds) to this sustained change. linking thresholds of potential concerns monitoring is directed at evaluating objectives at several scales, including at park and national level (dickens & graham 2002; noon et al. 1999; noss 1990; possingham et al. 2001). defining the linkages between management actions and objectives is a key step in deciding on the appropriate action, anticipated consequences and subsequent measurement (figure 3). in this paper we focus on direct measures of objectives. the unpacking of a concern, as described by gaylard and ferreira (2011), clearly illustrates how measuring sanparks objectives in a strategic adaptive management framework (biggs & rogers 2003) forces conservationists to monitor from individual species to ecosystem level. this allows determining whether the proposed mechanism of a concern has been adequately addressed through a management action. figure 3: sanparks uses a linkage framework that connects objectives, drivers, mechanisms and modulators. typically, conservationists seek nondirectional variance in biodiversity over time – a key objective. a hypothetical set of linkages relate ecosystem (biodiversity) objectives to key drivers and an indicator of such a driver. there are also linkages to the likely mechanism of how a driver affects biological diversity and factors that act as modulators on the mechanism. this example uses elephants and illustrates that management responses focus on modulators and that information requirements are diverse (adapted from gaylard, a. & ferreira, s.m., 2011, ‘modification and maturation of sanpark’s adaptive planning process – making critical linkages between conservation objectives and actions’, koedoe 53(2), art. #1005, 8 pages. doi:10.4102/koedoe.v53i2.1005). rare, endemic and cryptic species (back to top) most species are rare (cao, williams & williams et al. 1998; mcardle 1990), and rarity in itself poses significant problems for evaluating changes (cao et al. 1998). estimation of population sizes carries high uncertainty when populations are small (mackenzie et al. 2005; owen & rosentreter 1992). we propose that sanparks should focus on inventory techniques used in a predictive manner to evaluate its success with regard to achieving its biodiversity objectives. the process accommodates typical adaptive management approaches, with each cycle having a learning element. as shown in table 3, the initiation has two phases: • informed guessing based on expert knowledge because data are often too restrictive to construct formal landscape models • predicting species presence based on landscape characteristics. the proposed inventory monitoring requires some initial research. firstly, the degree of survey effort required to identify what species are present at a site (e.g. lewis & gould 2000; mackenzie & royle 2005) needs to be established. this will most likely take the form of a species accumulation curve with effort (colwell & coddington 1994), based on rarefaction methods (gotelli & colwell 2001) and field data. secondly, the required number of sites to develop predictive models regarding species absence or presence need to be determined. table 3: example of a predictive inventory approach for rare species and species of special concern inhabiting nondegraded areas. selected species abundances, trends and impacts (back to top) several species require and lend themselves to assessments of population sizes and associated trends (carter et al. 1995). this holds enormous value because population growth is a summary of a species’ response to the environment, other species and its own members (dann 1992; sibly et al. 2005). assessment of population sizes needs to recognise that conservationists can seldom, if ever, provide accurate (bias2 + precision) estimates (critchley & poulton 1998). however, in the past conservationists have seldom reported precision (standard deviation, confidence intervals, etc.) for estimates (e.g. bauer & van der merwe 2004; blanc et al. 2007). this may have influenced interpretation of information dramatically, because any technique for estimating population sizes entails several sources of error that introduce uncertainty to the estimate. these sources of error include sampling bias when a sample-based approach is used (walsh et al. 2001). however, even total counts commonly used by sanparks (e.g. van aarde, whyte & pimm 1999) or registration studies (e.g. gough & kerley 2006) may introduce biases relating to observers, availability and detectability (caughley 1974; redfern et al. 2002). we propose that these biases should be formally assessed, particularly with regard to the assessment of large mammal species. in addition to bias, sampling effort also affects precision of population estimates (ferreira & van aarde 2009). the precision of population estimates has a critical value because the detection of trends in a time series of population estimates depends on several factors (figure 4), one of which is the precisions of population estimates. estimates of precision help to minimise type i and type ii errors (concluding there is a trend when, in fact, there is none and concluding there is no trend when, in fact, there is, respectively), which may have dire consequences for conservation decisions if undetected (critchley & poulton 1998). monitoring designs have to consider trade-offs imposed by precision: for a desired level of precision, conservationists may be better off counting less often, with longer intervals between counts, but achieving better trend detection. in addition, monitoring selected species for trends regarding abundance usually is most valuable when linked explicitly to the impact on objectives. large mammals are of particular importance in this context (e.g. gordon, hester & festa-bianchet 2004). however, impacts by large mammals on other values are primarily associated with the degree to which mammals are using specific places (e.g. van aarde et al. 2006; van aarde & jackson 2007). intensity of use is primarily driven directly by the distribution of critical resources (e.g. harris et al. 2008; loarie, van aarde & pimm 2009) and indirectly by abundance (e.g. young, ferreira & van aarde 2009). detecting change with regard to impact and intensity of use carries the same trade-off as detecting trends in population estimates. these aspects need further development for defining monitoring of objectives associated with large mammals. given that intensity of use is the likely mechanism through which large mammals will affect ecosystems (table 4), information needs may substantially exceed what sanparks generates at present. using the ‘objective, indicators, mechanisms and modulators’ approach suggests that information relating to large mammals needs to include regional distribution, intensity of use and demography (table 5). figure 4: detecting population trends are constrained by type i and type ii errors. researchers traditionally accommodate type i errors and usually conclude that there is no trend when α > 0.05. however, type ii errors may have as important consequences. for example, a researcher may conclude that there is no declining trend when α > 0.05 for a threatened species, when, in fact, there was a high probability for a type ii error, which may ultimately lead to local extinction. the power to detect a trend carries several trade-offs illustrated in the diagram. this includes the magnitude of the trend to be detected, the variance of population estimates, the intervals between surveys and the number of surveys (gerrodette, t., 1987, ‘a power analysis for detecting trends’, ecology 68, 1364–1372. doi:10.2307/1939220). table 4: herbivory concerns unpacked across selected parks managed by sanparks, to illustrate how monitoring needs driven by linkages to objectives, mechanisms and modulators require a spectrum of monitoring information, but also many commonalities (sanparks unpublished records). table 5: example of information needs of sanparks based on the requirements of monitoring to support and evaluate management actions and their effects on objectives and summary of current data sets available for the specific example. ecosystem indicators: ecological, community-based assessments (back to top) community structure is the outcome of interactions between ecological processes (bradshaw 2000; prach et al. 2001). the evaluation of ecosystem management approaches (i.e. those actions that provide the opportunity for ecological processes to play out) may best be achieved by measuring temporal and spatial variation in key communities (i.e. those that are important for specific objectives of a national park). this could help sanparks to assess its ecosystem objectives and adhere to the complexity value that sanparks embraces (biggs & rogers 2003). evaluation of multivariate data sets that reflect various ecosystem processes may be a conceptual approach towards reflecting on ecosystem outcomes. such multivariate data sets can include abiotic, producer, decomposer and consumer aspects (e.g. van aarde et al. 1996). sanparks can gather several kinds of information to compile such data sets, including species composition, soil structures, vegetation cover assessments, landscape features and water quality assessments. we propose using multivariate data as the basis of ecosystem feature evaluation. sampling effort therefore needs to maximise the detection of species and species accumulation curves could potentially be used as an indicator (colwell & coddington 1994). sanparks should also be able to define differences and/or changes in multivariate indicators. biological communities for which data comprise species composition and a measure of abundance are used most often. differences and/or changes in multivariate data in the context of evaluating ecosystem objectives can be measured using either simple or complex parameters (figure 5). such measures are useful for two reasons. firstly, they allow sanparks to assess directional change in systems (e.g. wassenaar et al. 2005; wassenaar et al. 2007) and therefore can be used to direct maintenance as well as restoration conservation. secondly, these measures allow sanparks to relate biological community changes to other factors such as intensity of fires or the local density of large mammals, thereby supporting assessment of the links between management actions and objectives (see figure 3). figure 5: illustration of two scenarios of communities that differ and how simple measures can distinguish communities in some instances, but not so in other instances. to overcome this, we propose the use of dissimilarity measures such as an euclidean distance (d) illustrated here. the generality of monitoring across parks (back to top) the conceptual approach described here may present several challenges. some parks, for instance, may need case-by-case management decisions, which could impact differently across several management tiers in sanparks. even so, these may share mechanisms characteristic of a specific type of concern. to illustrate this, we collated all the herbivory concerns that have been expressed at various science–management interactions across several parks (table 4). for each case we unpacked the detail underlying the concern. for example, degrading vegetation results from elephants spending too much time in a specific area, which, in turn, is governed by critical resources such as water (van aarde et al. 2006). conservationists therefore need to understand the temporal and spatial distribution of water sources, the relationship between elephant spatial use and water, and vegetation changes related to elephant spatial use. although the concerns are varied, monitoring needs converge onto commonalities. the commonalities illustrate information needs of which the emphasis has shifted from a number focus to a systems and biodiversity approach. sanparks needs to find measures that evaluate whether the focus of a concern has changed. this may be well served through (1) a predictive inventory approach with residuals of observed versus expected species presence at sample sites as indicators of ecosystem health and (2) a multivariate approach to evaluate the response of biological communities to management-induced changes in herbivory. evaluating the ecological mechanisms leading to a herbivory concern converge onto defining spatial use and the effects of its direct modulators as well as those of indirect modulation resulting from abundance or density. these commonalities illustrate that the approach to use large mammal censuses as a primary basis of decision making in sanparks (e.g. kruger et al. 2008) requires additional information to assess fully whether management objectives have been reached within an adaptive management framework (biggs & rogers 2003). however, such approaches may be comparable across parks. the unpacking of a specific concern into mechanisms clearly illustrates a need to analyse how changes in the concern need to be associated with spatial use of large herbivores. this, in turn, needs to be associated with factors affecting spatial use to relate how management actions affect a concern. although the conceptual discussion used herbivory as a key example, the unpacking of concerns into mechanisms can easily be applied to all sanparks objectives (see gaylard & ferreira 2011). it is likely that other common concerns will have similar commonalities with regard to drivers, mechanisms and modulators across parks. in addition, unpacking concerns defines measurement parameters and inadvertently prioritises monitoring tasks. this approach forms the focus of one of the programmes of the biodiversity monitoring strategy of sanparks (mcgeogh et al. 2011). conclusions (back to top) sanparks adopted a strategic adaptive management approach for managing complex systems in the face of uncertainty by implementing tpcs. however, the success of defining and implementing tpcs across several parks has been variable, partly because of unfamiliar definitions, case-specific constraints on the applicability of state-based tpcs, logistical and methodological challenges with regard to measurement and, finally, challenges relating to prioritising the magnitude of tpc-generated tasks. given the dynamic nature of system processes, tpcs that define directional changes and associate with mechanisms may best serve the adaptive management approach that sanparks has embraced. such approaches share common features when concerns that may impose on the sanparks mandate are unpacked into drivers, mechanisms and associated modulators. in addition, they define the shift in information needs from numbers to ecosystems and biodiversity, and prioritise monitoring programmes that feed directly into key management issues, appropriate responses and consequences of the chosen management actions. references (back to top) ackoff, r.l., 2010, systems thinking for curious managers, triarchy press, devon. agardi, t., 2000, ‘information needs for marine protected areas: scientific and societal’, bulletin of marine science 66, 875–888. agrawal, a.a., 2001, ‘phenotypic plasticity in the interactions and evolution of species’, science 294, 321–326. doi:10.1126/science.1060701, pmid:11598291 aradóttir, á.l. & arnalds, o., 2001, ‘ecosystem degradation and restoration of birch woodlands in iceland’, in f.e. wielgolaski (ed.), nordic mountain birch ecosystems, pp. 293–306, parthenon, paris. bauer, h. & van der merwe, s., 2004, ‘inventory of free-ranging lions panthera leo in africa’, oryx 38, 26–31. doi:10.1017/s0030605304000055 belsky, a.j., 1990, ‘tree/grass ratios in east african savannas: a comparison of existing models’, journal of biogeography 17, 483–489. doi:10.2307/2845380 biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring, and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59–80, island press, washington dc. biggs, h.c., ferreira, s.m., freitag-ronaldson, s. & grant-biggs, r., 2011, ‘taking stock after a decade: does the “thresholds of potential concern” concept need a socio-ecological revamp?’, koedoe 53(2), art. #1002, 9 pages. doi:10.4102/koedoe.v53i2.1002 blanc, j.j., barnes, r.f.w., craig, g.c., dublin, h.t., thouless, c.r., douglas-hamilton, i., et al., 2007, african elephant status report 2007: an update from the african elephant database, iucn/ssc african elephant specialist group, gland. doi:10.2305/iucn.ch.2007.ssc-op.33.en bocking, s., 1994, ‘visions of nature and society: a history of the ecosystem concept’, alternatives 20, 12–19. braaz, s., 1992, conserving biological diversity: a strategy for protected areas in the asia-pacific region, world bank, washington dc. bradshaw, a.d., 2000, ‘the use of natural processes in reclamation – advantages and difficulties’, landscape and urban planning 51, 89–100. doi:10.1016/s0169-2046(00)00099-2 brandon, k., gorenflo, l.j., rodrigues, a.s.l. & waller, r.w., 2005, ‘reconciling biodiversity conservation, people, protected areas, and agricultural suitability in mexico’, world development 33, 1403–1418. doi:10.1016/j.worlddev.2004.10.005 brandon, k., redford, k.h. & sanderson, s.e., 1998, parks in peril: people, politics, and protected areas, island press, washington dc. bruner, a.g., gullison, r.e., rice, r.e. & da fonseca, g.a.b., 2001, ‘effectiveness of parks in protecting tropical biodiversity’, science 291, 125–128. doi:10.1126/science.291.5501.125, pmid:11141563 brussard, p.f., reed, j.m. & tracy, c.r., 1998, ‘ecosystem management: what is it really?’, landscape and urban planning 40, 9–20. doi:10.1016/s0169-2046(97)00094-7 cao, y., williams, d.d. & williams, n.e., 1998, ‘how important are rare species in aquatic community ecology and bioassessment?’, limnology and oceanography 43, 1403–1409. doi:10.4319/lo.1998.43.7.1403 carter, h.r., sowls, a.l., rodway, m.s., wilson, u.w., lowe, r.w., mcchesney., et al., 1995, ‘population size, trends, and conservation problems of the double-crested cormorant on the pacific coast of north america’, in d.n. nettleship & d.c. duffy (eds.), the double-crested cormorant: biology, conservation and management, pp. 189–215 (colonial waterbirds; vol. 18). caughley, g., 1974, ‘bias in aerial survey’, journal of wildlife management 38, 921–933. doi:10.2307/3800067 chapin, f.s., zavaleta, e.s., eviner, v.t., naylor, r.l., vitousek, p.m., reynolds, h.l., et al., 2000, ‘consequences of changing biodiversity’, nature 405, 234–242. doi:10.1038/35012241, pmid:10821284 christensen, n.l., 1997, ‘managing for heterogeneity and complexity on dynamic landscapes’, in s.t.a. pickett, r.s. ostfeld, m. shachak & g.e. likens (eds.), the ecological basis of conservation: heterogeneity, ecosystems and biodiversity, pp. 60–80, chapman & hall, new york. colwell, r.k. & coddington, j.a., 1994, ‘estimating terrestrial biodiversity through extrapolation’, philosophical transactions of the royal society of london b 345, 101–118. doi:10.1098/rstb.1994.0091, pmid:7972351 critchley, c.n.r. & poulton, s.m.c., 1998, ‘a method to optimize precision and scale in grassland monitoring’, journal of vegetation science 9, 837–846. doi:10.2307/3237049 curran, l.m., trigg, s.n., mcdonald, a.k., astiani, d., hardiono, y.m., siregar, p., et al., 2004, ‘lowland forest loss in protected areas of indonesian borneo’, science 303, 1000–1003. doi:10.1126/science.1091714, pmid:14963327 dann, p., 1992, ‘distribution, population trends and factors influencing the population size of little penguins eudyptula minor on phillip island, victoria’, emu australian ornithology 91, 263–272. doi:10.1071/mu9910263 daszak, p., cunningham, a.a. & hyatt, a.d., 2000, ‘emerging infectious diseases of wildlife – threats to biodiversity and human health’, science 287, 443–449. doi:10.1126/science.287.5452.443, pmid:10642539 de leo, g.a. & levin, s., 1997, ‘the multifaceted aspects of ecosystem integrity’, conservation ecology, vol. 1, viewed 02 february 2010, from http://www.consecol.org/vol1/iss1/art3 dickens, c.w.s. & graham, p.m., 2002, ‘the south african scouring system (sass) version 5 rapid assessment method for rivers’, african journal of aquatic science 27, 1–10. doi:10.2989/16085914.2002.9626569 doyle, t. & mceachern, d., 2008, environment and politics, routledge, new york. du toit, j.t., rogers, k.h. & biggs, h.c., 2003, the kruger experience: ecology and management of savanna heterogeneity, island press, washington dc. ferreira, s.m. & van aarde, r.j., 2009, ‘aerial survey intensity as a determinant of estimates of african elephant population sizes and trends’, south african journal of wildlife research 39, 181–191. doi:10.3957/056.039.0205 foxcroft, l.c., 2009, ‘developing thresholds of potential concern for invasive alien species: hypotheses and concepts’, koedoe 51, 1–6. doi:10.4102/koedoe.v51i1.157 gaston, k.j., 2008, ‘biodiversity and extinction: the importance of being common’, progress in physical geography 32, 73–79. doi:10.1177/0309133308089499 gaylard, a. & ferreira, s.m., 2011, ‘modification and maturation of sanpark’s adaptive planning process – making critical linkages between conservation objectives and actions’, koedoe 53(2), art. #1005, 8 pages. doi:10.4102/koedoe.v53i2.1005 gerrodette, t., 1987, ‘a power analysis for detecting trends’, ecology 68, 1364–1372. doi:10.2307/1939220 ghimire, k.b. & pimbert, m.p., 1997, social change and conservation, earthscan publications, london. gordon, i.j., hester, a.j. & festa-bianchet, m., 2004, ‘the management of wild large herbivores to meet economic, conservation and environmental objectives’, journal of applied ecology 41, 1021–1031. doi:10.1111/j.0021-8901.2004.00985.x gotelli, n.j. & colwell, r.k., 2001, ‘quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness’, ecological letters 4, 379–391. doi:10.1046/j.1461-0248.2001.00230.x gough, k. & kerley, g.i.h., 2006, ‘demography and population dynamics in the elephants loxodonta africana of addo elephant national park, south africa: is there evidence of density dependent regulation?’, oryx 40, 434–441. doi:10.1017/s0030605306001189 harris, g.m., russell, g.j., van aarde, r.j. & pimm, s.l., 2008, ‘habitat use of savanna elephants in southern africa’, oryx 42, 66–75. doi:10.1017/s0030605308000483 hüttl, r.f. & schneider, b.u., 1998, ‘forest ecosystem degradation and rehabilitation’, ecological engineering 10, 19–31. doi:10.1016/s0925-8574(97)10023-4 john, r., dalling, j.w., harms, k.e., yavitt, j.b., stallard, r.f., mirabello, m., et al., 2007, ‘soil nutrients influence spatial distributions of tropical tree species’, proceedings of the national academy of sciences of the united states of ameria 104, 864–869. doi:10.1073/pnas.0604666104, pmid:17215353, pmcid:1783405 kay, j.j. & schneider, e., 1994, ‘embracing complexity: the challenge of the ecosystem approach’, alternatives 20, 32–46. knight, a.t., driver, a., cowling, r.m., maze, k., desmet, p.g., lombard, a.t., et al., 2006, ‘designing systematic conservation assessments that promote effective implementation: best practice from south africa’, conservation biology 2, 203–217. knight, m.h., 2007, sanparks state of biodiversity report (sob) background sheet, sanparks, pretoria. kotliar, n.b. & wiens, j.a., 1990, ‘multiple scales of patchiness and patch structure: a hierachical framework for the study of heterogeneity’, oikos 59, 253–260. doi:10.2307/3545542 kruger, j.m., reilly, b.k. & whyte, i.j., 2008, ‘application of distance sampling to estimate population densities of large herbivores in kruger national park’, wildlife research 35, 371–376. doi:10.1071/wr07084 lawton, j.h., 1997, ‘the science and non-science of conservation’, oikos 79, 3–5. doi:10.2307/3546084 lewis, c., 1996, managing conflicts in protected areas, iucn, gland. lewis, s.a. & gould, w.r., 2000, ‘survey effort effects on power to detect trends in raptor migration counts’, wildlife society bulletin 28, 317–329. lindenmayer, d.b., franklin, j.f. & fischer, j., 2006, ‘general management principles and a checklist of strategies to guide forest biodiversity conservation’, biological conservation 131, 433–445. doi:10.1016/j.biocon.2006.02.019 lindenmayer, d.b. & likens, g.e., 2009, ‘adaptive monitoring: a new paradigm for long-term research and monitoring’, trends in ecology and evolution 24, 482–486. doi:10.1016/j.tree.2009.03.005 liu, j., linderman, m., ouyang, z., an, l., yang, j. & zhang, h., 2001, ‘ecological degradation in protected areas: the case of wolong nature reserve for giant pandas’, science 292, 98–101. doi:10.1126/science.1058104, pmid:11292872 loarie, s.r., van aarde, r.j. & pimm, s.l., 2009, ‘fences and artificial water affect african savannah elephant movement patterns’, biological conservation 142, 2086–3098. doi:10.1016/j.biocon.2009.08.008 lyons, k.g., brigham, c.a., traut, b.h. & schwartz, m.w., 2005, ‘rare species and ecosystem functioning’, conservation biology 19, 1019–1024. doi:10.1111/j.1523-1739.2005.00106.x mackenzie, d.i., nichols, j.d., sutton, n., kawanishi, k. & bailey, l.l., 2005, ‘improving inferences in population studies of rare species that are detected imperfectly’, ecology 86, 1101–1113. doi:10.1890/04-1060 mackenzie, d.i. & royle, j.a., 2005, ‘designing occupancy studies: general advice and allocating survey effort’, journal of applied ecology 42, 1105–1114. doi:10.1111/j.1365-2664.2005.01098.x margules, c.r. & pressey, r.l., 2000, ‘systematic conservation planning’, nature 405, 243–253. doi:10.1038/35012251, pmid:10821285 martello, m.l., 2001, ‘a paradox of virtue?: “other” knowledges and environment-development politics’, global environmental politics 1, 114–141. doi:10.1162/152638001316881430 mcardle, b.h., 1990, ‘when are rare species not there?’, oikos 57, 276–277. doi:10.2307/3565950 mcgeogh, m.a., dopolo, m., novellie, p., hendricks, h., freitag, s., ferreira, s.m., et al., 2011, ‘a strategic framework for biodiversity monitoring in sanparks’, koedoe 53(2), art. #991, 10 pages. doi:10.4102/koedoe.v53i2.991 mcneely, j.a., 1994, ‘protected areas for the 21st century: working to provide benefits to society’, biodiversity and conservation 3, 390–405. doi:10.1007/bf00057797 milton, s.j., dean, w.r.j., du plessis, m.a. & siegfried, w.r., 1994, ‘a conceptual model of arid rangeland degradation’, bioscience 44, 70–76. doi:10.2307/1312204 morellet, n., gaillard, j-m., hewison, a.j.m., ballon, p., boscardin, y., duncan, p., et al., 2007, ‘indicators of ecological change: new tools for managing populations of large herbivores’, journal of applied ecology 44, 634–643. doi:10.1111/j.1365-2664.2007.01307.x noon, b.r., spies, t.a. & raphael, m.g., 1999, ‘conceptual basis for designing an effectiveness monitoring program’, in b.s. mulder (ed.), the strategy and design of the effectiveness of a monitoring program for the northwest forest plan, pp. 21–48, us department of agricultural forest service, portland. noss, r.f., 1990, ‘indicators for monitoring biodiversity: a hierarchical approach’, conservation biology 4, 355–364. doi:10.1111/j.1523-1739.1990.tb00309.x noss, r.f., 1999, ‘assessing and monitoring forest biodiversity: a suggested framework and indicators’, forest ecology and management 115, 135–146. doi:10.1016/s0378-1127(98)00394-6 ormerod, s.j., 2003, ‘restoration in applied ecology: editor’s introduction’, journal of applied ecology 40, 44–50. doi:10.1046/j.1365-2664.2003.00799.x owen, w.r. & rosentreter, r., 1992, ‘monitoring rare perennial plants: techniques for demographic studies’, natural areas journal 12, 32–38. parrish, j.d., braun, d.p. & unnasch, r.s., 2003, ‘are we conserving what we say we are? measuring ecological integrity within protected areas’, bioscience 53, 851–860. doi:10.1641/0006-3568(2003)053[0851:awcwws]2.0.co;2 pickett, t.a., cadenasso, m. & benning, t., 2003, ‘biotic and abiotic variability as key determinants of savannah heterogeneity at multiple spatiotemporal scales’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 22–40, island press, washington dc. pimm, s.l., 1991, the balance of nature? ecological issues in the conservation of species and communities, university of chicago press, chicago. possingham, h.p., andelman, s.j., noon, b.r., trombulak, s. & pulliam, h.r., 2001, ‘making smart conservation decisions’, in m.e. soule & g.h. orians (eds.), conservation biology: research priorities for the next decade, pp. 225–244, island press, washington dc. prach, k., bartha, s., joyce, c.b., pyšek, p., van diggelen, r. & wiegleb, g., 2001, ‘the role of spontaneous vegetation succession in ecosystem restoration: a perspective’, applied vegetation science 4, 111–114. doi:10.1111/j.1654-109x.2001.tb00241.x preston, f.w., 1948, ‘the commonness and rarity of species’, ecology 29, 254–283. doi:10.2307/1930989 redfern, j.v., viljoen, p.c., kruger, j.m. & getz, w.m., 2002, ‘biases in estimating population size from an aerial census: a case study in the kruger national park, south africa’, south african journal of science 98, 455–461. regan, h.m., hierl, l.a., franklin, j., deutschman, d.h., schmalbach, h.l., winchell, c.s., et al., 2008, ‘species prioritization for monitoring and management in regional species conservation plans’, diversity and distributions 14, 462–471. doi:10.1111/j.1472-4642.2007.00447.x rietkerk, m. & van de koppel, j., 1997, ‘alternate stable states and threshold effects in semi-arid grazing systems’, oikos 79, 69–76. doi:10.2307/3546091 rodrigues, a.s.l., andelman, s.j., bakarr, m.i., boitani, l., brooks, t.m., cowling, r.m., et al., 2004, ‘effectiveness of the global protected area network in representing species diversity’, nature 428, 640–643. doi:10.1038/nature02422, pmid:15071592 rogers, k.h., 1997, ‘operationalizing ecology under a new paradigm: an african perspective’, in s.t.a. pickett, r.s. ostfeld, m. shachak & g.e. likens (eds.), the ecological basis of conservation: heterogeneity, ecosystems and biodiversity, pp. 60–80, chapman & hall, new york. salafsky, n. & margoluis, r., 1999, ‘threat reduction assessment: a practical and cost-effective approach to evaluating conservation and development projects’, conservation biology 13, 830–841. doi:10.1046/j.1523-1739.1999.98183.x sale, p.f., cowen, r.k., danilowicz, b.s., jones, g.p., kritzer, j.p. lindeman, k.c., et al., 2005, ‘critical sciences gaps impede use of no-take fisheries reserves’, trends in ecology and evolution 20, 74–80. doi:10.1016/j.tree.2004.11.007 sanparks. see south african national parks. south african national parks, 2007, kruger national park biodiversity management programme: strategic adaptive management, sanparks, skukuza. south african national parks, 2010, south african national parks: resource use policy, sanparks, pretoria. scoones, i., 1999, ‘new ecology and the social sciences: what prospects for a fruitful engagement?’, annual review of anthropology 28, 479–507. doi:10.1146/annurev.anthro.28.1.479 sibly, r.m., barker, d., denham, m.c., hone, j. & pagel, m., 2005, ’on the regulation of populations of mammals, birds, fish and insects’, science 309, 607–610. doi:10.1126/science.1110760, pmid:16040705 simberloff, d., 1999, ‘the role of science in the preservation of forest biodiversity’, forest ecology and management 115, 101–111. doi:10.1016/s0378-1127(98)00391-0 slocombe, d.s., 1998, ‘lessons from experience with ecosystem-based management’, landscape and urban planning 40, 31–39. doi:10.1016/s0169-2046(97)00096-0 smith, m.d. & knapp, a.k., 2003, ‘dominant species maintain ecosystem function with non-random species loss’, ecology letters 6, 509–517. doi:10.1046/j.1461-0248.2003.00454.x society of ecological restoration, 2004, the ser primer on ecological restoration, society for ecological restoration science and policy working group, viewed 02 february2010, from http://www.ser.org/reading_resources.asp spellerberg, i.f., 2005, monitoring ecological change, cambridge university press, cambridge. doi:10.1017/cbo9780511614699 stringham, t.k., krueger, w.c. & shave, p.l., 2003, ‘state and transition modeling: an ecological process approach’, journal of range management 56, 106–113. doi:10.2307/4003893 tompkins, e.l. & adger, w.n., 2004, ‘does adaptive management of natural resources enhance resilience to climate change?’, ecology and society 9, 10, viewed 02 february 2010, from http://www.ecologyandsociety.org/vol9/iss2/art10/ trollope, w.s.w., potgieter, a.l.f., biggs, h.c. & brockett, b., 1999, report on the necessity, design and implementation of the large scale herbivore fire interaction research experiment (lashfire) in the kruger national park in south africa, sanparks, skukuza. van aarde, r.j., ferreira, s.m., kritzinger, j.j., van dyk, p.j., vogt, m. & wassenaar, t.d., 1996, ‘an evaluation of habitat rehabilitation on coastal dune forests in northern kwazulu-natal, south africa’, restoration ecology 4, 334–345. doi:10.1111/j.1526-100x.1996.tb00186.x van aarde, r.j. & jackson, t.p., 2007, ‘megaparks for metapopulations: addressing the causes of locally high elephant numbers in southern africa’, biological conservation 134, 289–297. doi:10.1016/j.biocon.2006.08.027 van aarde, r.j., jackson, t.p. & ferreira, s.m., 2006, ‘conservation science and elephant management in southern africa’, south african journal of science 102, 385–388. van aarde, r.j., whyte, i. & pimm, s.l., 1999, ‘culling and the dynamics of the kruger national park african elephant population’, animal conservation 2, 287–294. doi:10.1111/j.1469-1795.1999.tb00075.x varghese, g., 2008, ‘public-private partnerships in south african national parks: rationale, benefits and lessons learned’, in a. spenceley (ed.), responsible tourism: critical issues for conservation and development, pp 69–84, earthscan publications, london. walker, b.h., 1987, ‘a general model of savanna structure and function’, in b.h. walker (ed.), determinants of tropical savanna, pp. 1–12, icsu press, miami. walsh, p.d., white, l.j.t., mbina, c., idiata, d., mihindou, y., maisels, f., et al., 2001, ‘estimates of forest elephant abundance: projecting the relationship between precision and effort’, journal of applied ecology 38, 217–228. doi:10.1046/j.1365-2664.2001.00578.x wassenaar, t.d., pimm, s.l., van aarde, r.j. & ferreira, s.m., 2005, ‘community convergence in disturbed sub-tropical dune forests’, ecology 86, 655–666. doi:10.1890/03-0836 wassenaar, t.d., ferreira, s.m. & van aarde, r.j., 2007, ‘flagging aberrant sites and assemblages in restoration projects’, restoration ecology 15, 68–76. doi:10.1111/j.1526-100x.2006.00191.x weiner, j., 1992, ‘physiological limits to sustainable energy budgets in birds and mammals: ecological implications’, trends in ecology and evolution 7, 384–388. doi:10.1016/0169-5347(92)90009-z westoby, m., walker, b. & noy-meir, i., 1989, ‘opportunistic management for rangelands not at equilibrium’, journal of range management 42, 266–274. doi:10.2307/3899492 wilson, e.o., 1988, the current state of biological diversity, biodiversity national academy press, washington dc. woodroffe, r. & ginsberg, j.r., 1998, ‘edge effects and the extinction of populations inside protected areas’, science 280, 2126–2128. doi:10.1126/science.280.5372.2126, pmid:9641920 young, k.d., ferreira, s.m. & van aarde, r.j., 2009, ‘the influence of increasing population size and vegetation productivity on elephant distribution in the kruger national park’, austral ecology 34, 329–342. doi:10.1111/j.1442-9993.2009.01934.x a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe original research a rticle #149 (page number not for citation purposes) 17 a checklist of the non-acarine arachnids (chelicerata: arachnida) of the de hoop nature reserve, western cape province, south africa authors: charles r. haddad1 ansie s. dippenaarschoeman2 affiliations: 1department of zoology & entomology university of the free state, south africa 2biosystematics: arachnology arc plant protection research institute south africa 2department of zoology & entomology university of pretoria south africa correspondence to: charles r. haddad e-mail: haddadcr.sci@ufs.ac.za postal address: department of zoology & entomology, university of the free state, p. o. box 339, bloemfontein, 9300, south africa keywords: de hoop nature reserve; fynbos biome; nonacarine arachnids; species; south african national survey of arachnida dates: received: 13 nov. 2007 accepted: 08 july 2008 published: 21 apr. 2009 how to cite this article: haddad, c.r. & dippenaar-schoeman, a.s., 2009, ‘a checklist of the non-acarine arachnids (chelicerata: arachnida) of the de hoop nature reserve, western cape province, south africa’, koedoe 51(1), art. #149, 9 page. doi: 10.4102/ koedoe.v51i1.149 this article is available at: http://www.koedoe.co.za © 2009. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. vol. 51 no. 1 page 1 of 9 abstract as part of the south african national survey of arachnida (sansa) in conserved areas, arachnids were collected in the de hoop nature reserve in the western cape province, south africa. the survey was carried out between 1999 and 2007, and consisted of five intensive surveys between two and 12 days in duration. arachnids were sampled in five broad habitat types, namely fynbos, wetlands, i.e. de hoop vlei, eucalyptus plantations at potberg and cupido’s kraal, coastal dunes near koppie alleen and the intertidal zone at koppie alleen. a total of 274 species representing five orders, 65 families and 191 determined genera were collected, of which spiders (araneae) were the dominant taxon (252 spp., 174 genera, 53 families). the most species rich families collected were the salticidae (32 spp.), thomisidae (26 spp.), gnaphosidae (21 spp.), araneidae (18 spp.), theridiidae (16 spp.) and corinnidae (15 spp.). notes are provided on the most commonly collected arachnids in each habitat. conservation implications: this study provides valuable baseline data on arachnids conserved in de hoop nature reserve, which can be used for future assessments of habitat transformation, alien invasive species and climate change on arachnid biodiversity. introduction the south african national survey of arachnida (sansa) was initiated in 1997 to record the biodiversity of arachnids in south africa (dippenaar-schoeman & craemer 2000). as part of this initiative, surveys are underway in various conservancies, agroecosystems, provinces and biomes. so far, only two long-term surveys have been carried out in western cape province conservancies, namely of the spiders of the karoo national park, falling within the nama karoo biome (dippenaar-schoeman et al. 1999), and the swartberg nature reserve, falling within the succulent karoo biome (dippenaar-schoeman et al. 2005). these two surveys indicate a moderately high diversity of spiders in these conservancies, with 116 species (38 families) and 186 species (45 families) recorded from the two reserves, respectively. the cape floristic region comprises unique vegetation types such as fynbos, which are characterised by high levels of plant endemism. according to linder (2005) some 9,000 species can be found in the region in an area of approximately 90,000 km2. although the factors influencing insect abundance and diversity in this biome have been well studied (e.g. giliomee 2003; procheş & cowling 2006; wright & samways 1996, 1999), little is known on the diversity of arachnids in the fynbos biome. coetzee et al. (1990) studied the spiders associated with five proteaceous plant species, visser et al. (1999) studied the arachnids associated with protea nitida mill., and sharratt (2000) included arachnids in their assessment of the conservation status of cave-dwelling arthropods of the cape peninsula. the general lack of information regarding arachnid diversity, as well as that for many other invertebrate groups in the western cape province, is a great hindrance to effective conservation planning. conservation strategies should not only take into account plants and vertebrates, but also need to recognise the role that invertebrates play in ecosystem functioning. arachnids, with the exception of some phytophagous and parasitic acari, form an important group of predatory terrestrial arthropods that feed on a wide variety of prey using a range of capture methods, including webs and active hunting strategies. arachnids are frequently regarded as suitable candidates for studying ecological processes, as 1) they are diverse and abundant, 2) they can be easily sampled, 3) they are functionally significant in ecosystems as predators, and as food for other predators, and 4) they interact with their abiotic and biotic environment in a manner that reflects ecological change (churchill 1997). therefore, arachnids can be used to monitor ecosystem stability and changes over time, making them useful organisms in long-term conservation planning. since fynbos vegetation, which is largely endemic to the western cape province, is under increasing threat from urbanisation, agriculture, alien invasive species and climate change (e.g. picker & samways 1996; richardson et al. 1996; mcneely 2001; midgley et al. 2003; witt & samways 2004), arachnids provide an alternative taxonomic group to monitor changes in this unique vegetation type. the present paper aims to report on the diversity of arachnids (excluding the acari) in the de hoop nature reserve (dhnr) in the western cape, which consists of large areas of pristine fynbos and protected marine habitats. apart from its value as a biodiversity and conservation tool, this checklist can thus be used as a baseline to assess impacts of the aforementioned effects on biodiversity in areas surrounding the reserve. this study forms part of the south african national survey of arachnida in conserved areas and the fynbos biome, and also contributes towards the checklists of species of the western cape province. study area dhnr is situated on the south coast of the western cape province, south africa, and covers an area of koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research haddad & dippenaar-schoeman a rt ic le # 14 9 (page number not for citation purposes) 18 vol. 51 no. 1 page 2 of 9 figure 1 location of the de hoop nature reserve along the south coast of south africa. enlarged map shows key sampling points in the reserve figures 2–7 habitats sampled in the de hoop nature reserve: 2) fynbos (fb); 3) eucalyptus plantation at potberg (ep); 4) wetland at de hoop vlei (wl); 5–6) coastal dunes at koppie alleen (cd), with natural vegetation (5) and dunes covered with invasive alien acacia species (6); 7) intertidal zone at koppie alleen (iz) 32,279 hectares terrestrially (figure 1). in addition, the coastline and adjacent marine areas are also included in the reserve for the protection of the marine environment and its diversity. for the purposes of this survey the reserve was divided into five broad sampling habitats (plant classification follows germishuizen et al. 2006): fynbos (fb) – the largest portion of the reserve contains 1. typical fynbos vegetation characteristic of this particular floral biome (figure 2). an upper vegetative layer consisting primarily of taller protea spp. (p. aurea potbergensis rourke, p. obtusifolia h.buek ex meisn. and p. repens (l.) l.) is found in certain areas, particularly near hills and mountains. the field layer comprises a high diversity of fynbos plants, including agathosma spp., cliffortia spp., leucodendron spp., phylica spp., serruria fasciflora salisb. ex knight and thamnochortus spp.. eucalyptus2. plantation (ep) – two large plantations at potberg and cupido’s kraal consist primarily of eucalyptus camaldulensis dehnh., with endemic low-growing shrubs (e.g. carissa bispinosa (l.) desf. ex brenan) and other short vegetation (agaranthus sp., asparagus falcatus l., bidens sp., cynodon dactylon (l.) pers. and sansevieria hyacinthoides (l.) druce) (figure 3). wetlands (wl) – a single inland wetland, i.e. the de hoop 3. vlei, is situated in the south-west of the reserve (figure 4). the wetland is separated from the ocean by coastal dunes, and therefore does not form a lagoon per se. the de hoop vlei is fed by water from the zout river, the catchment of which receives most of its rainfall during the winter rainfall season. the shores of the wetland are dominated by sarcocornia spp. and exomis microphylla (thunb.) aellen., with scattered patches of the reed phragmites australis (cav.) steud.. beyond the shoreline the dominant vegetation includes sideroxylon inerme l. trees and a variety of fynbos species. coastal dunes (cd) – coastal dune vegetation is found 4. along the entire coastline of the reserve (figure 5). seafacing dunes consist primarily of endemic shrub species, including carissa bispinosa, cynanchum obtusifolium l.f., euclea racemosa murray, passerina rigida wikstr., ptaeroxylon spp., robsonodendon sp., rhus glauca thunb. and secamone spp., interspersed with shorter species such as arctotheca populifolia (p.j.bergius) norl., asparagus falcatus, bassia diffusa (thunb.) kuntze, chironia baccifera l., dasispermum suffruticosum (p.j.bergius) b.l.burtt, gazania krebsiana less., limonium scabrum (thunb.) kuntze, plantago crassifolia forssk., silene primuliflora eckl. & zeyh., spirobolus sp., trachyandra ciliata (l.f) kunth and fynbos vegetation. many dunes are strongly overgrown with invasive alien plant species such as acacia cyclops a.cunn ex g.don and a. saligna (labill.) h.l.wendl. (figure 6), occasionally interspersed with fynbos elements. intertidal zone (iz) – this habitat includes all rocky shores 5. along the coastline and the vegetation immediately associated with the high tide breaker line (figure 7). on the rocky shores themselves, various marine algae dominate, while plants associated with the high tide mark include scattered fynbos insertions and coastal dune shrubs. sampling period and methods intensive sampling for arachnids was carried out during five visits to the reserve. three of the trips were carried out during early autumn (march 1999 – april 1999, 2004 and 2005) and lasted 10 – 12 days each, the fourth trip was undertaken during the middle of winter (july 2005) and lasted four days, and the last trip took place in spring (september 2007) for two days. sampling was undertaken ad hoc in each of the habitats by active searching under rocks, logs and in leaf litter, beating foliage, sifting leaf litter and sweeping low-growing vegetation. additional sampling was conducted by searching under bark in the ep, as this was the only habitat in which loose bark was available. material was preserved in 70% ethanol for sorting and identification. due to time and logistical constraints during the sampling trips, material was not collected quantitatively (i.e. according to a set sampling protocol). thus, the sampling intensity varied considerably between habitats with a bias towards collecting in fb and ep, as these were the easiest habitats to access. however, adequate sampling was conducted in the other three habitats using various methods to give a good indication of the arachnid diversity of each. guilds observed all arachnids were grouped into guilds based on the typical habits known for each family or genus, but also took into consideration the strata in which each species was sampled. potberg de hoop vlei cupido’s kraal western cape gauteng kwazulu-natal northern cape eastern cape limpopo north west free state mpumalanga koppie alleen checklist of the non-acarine arachnids original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #149 (page number not for citation purposes) 19vol. 51 no. 1 page 3 of 9 all arachnid orders collected, with the exception of spiders, can be classified as wanderers. spiders can be separated into wandering and web-building guilds. the wandering arachnids can be broadly separated into ground wanderers (gw) and plant wanderers (pw). for the latter group, distinction was made between spiders associated with foliage (pwf) of plants and those associated with the bark of trees (pwb). webbuilding spiders can be separated into various guilds based on the types of webs they construct, namely orb-web builders (owb), funnel-web builders (fwb), sheet-web builders (swb), space-web builders (spwb), hackle-web builders (hwb) and gum-foot-web builders (gwb). representative specimens of each species are deposited in the institutions of the various specialists listed in the acknowledgements, who provided identifications for their respective groups. material of all the remaining taxa is deposited in the national collection of arachnida at the plant protection research institute, pretoria, south africa. results & discussion diversity a total of 274 species of arachnids were collected in dhnr, representing five orders, 65 families and 191 determined genera (table 1, appendix 1). the most species rich order was the araneae, with 252 species in 54 families. this includes one published record of a species that was not collected in the current survey, nephila fenestrata thorell (nephilidae) (fromhage et al. 2007). the spider family diversity represents the highest from south africa, exceeding the 46 families collected in the western soutpansberg in limpopo province (foord et al. 2002) and ndumo game reserve in kwazulu-natal (haddad et al. 2006). the relatively high spider diversity from fynbos is impressive when compared to more structurally complex habitats such as savanna, where greater species diversity could be expected (see table 2). the remaining arachnid orders were relatively poorly represented, the most species rich being the pseudoscorpiones (nine species, five families), followed by opiliones (eight species, table 1 order composition of the non-acarine arachnids of the de hoop nature reserve, western cape province, south africa order common name families genera species araneae spiders 53 174 252 opiliones harvestmen 3 5 8 pseudoscorpiones false scorpions 5 7 9 scorpiones scorpions 3 4 4 solifugae sun spiders 1 1 1 total 65 191 274 table 2 guild composition of spiders collected in the de hoop nature reserve, compared to other surveys carried out in south african conservation areas. abbreviations: wa – wanderers; wb – web-builders conservancy biome spp. %wa %wb reference de hoop nature res. fynbos 252 70.6 29.4 current study karoo nat. park nama karoo 116 66.4 33.6 dippenaar-schoeman et al. (1999) kruger nat. park savanna 152 79.0 21.0 dippenaar-schoeman & leroy (2003) makalali game res. savanna 268 69.4 30.6 whitmore et al. (2002) mountain zebra nat. park nama karoo 76 53.9 46.1 dippenaar-schoeman (2006) ndumo game res. savanna 431 74.2 25.8 haddad et al. (2006) polokwane nature res. savanna 275 69.5 30.5 dippenaar et al. (2008) roodeplaat dam nature res. savanna 110 65.5 34.5 dippenaar-schoeman et al. (1989) sovenga hill savanna 76 83.9 16.1 modiba et al. (2005) swartberg nature res. succulent karoo 186 76.5 23.5 dippenaar-schoeman et al. (2005) western soutpansberg savanna 127 63.8 36.2 foord et al. (2002) three families), scorpiones (four species, three families), and solifugae (one species, one family). one published record of scorpiones, of parabuthus planicauda (pocock) (buthidae), was found in the literature (prendini 2004). as in several other south africa surveys, salticidae were the most species rich family (32 spp., 12.7% of spiders), followed by the thomisidae (26 spp., 10.3%) and gnaphosidae (21 spp., 8.3%). several other families contributed 5% or more of the spider species: araneidae (18 spp., 7.1%), theridiidae (16 spp., 6.3 %) and corinnidae (15 spp., 6.0%). in contrast to some other reserves previously sampled in south africa, such as the ndumo game reserve in kwazulu-natal, the family composition of spiders was considerably less skewed in the current study (figure 8). at ndumo, the five dominant spider families contributed 52% of the species, with the salticidae dominant (82 spp., 19.0%) (haddad et al. 2006). in contrast, the five families dominating the current study contributed 44.7% of the total spiders, with the dominant salticidae only contributing 12.7% of the total. guilds the majority of the arachnid species collected in dhnr are wanderers (73.0%), while web-builders comprise 27.0%. when spiders alone are considered, 70.6% are wanderers while 29.4% are web-builders. this compares well with several surveys completed in south africa (table 2). this indicates that fynbos and associated habitats sampled in this study are sufficiently heterogeneous to support a fauna similar to that found in more structurally complex habitat types, such as savanna. common taxa by stratum this study was qualitative in its entirety and thus there is no data available on the relative abundance of arachnids. however, based on the frequency of collection and observations made during the study the following species can be recognised as representative of each stratum and guild: ground wanderers: a large proportion of the species collected are wandering arachnids on the soil surface (appendix 1). the coastal dune (cd) fauna was largely dominated by pardosa and trabea spp. (lycosidae), griswoldia robusta (simon) (zoropsidae), opopaea speciosa (lawrence) (oonopidae), zelotes anchora tucker (gnaphosidae), natta spp. (salticidae), diores simoni o. p.-cambridge (zodariidae) and orthobula infima simon (corinnidae). in the eucalyptus plantation (ep), various gnaphosids (especially zelotes, camillina and xerophaeus spp.), caponia capensis purcell (caponiidae), opopaea speciosa, xysticus lucifugus lawrence (thomisidae), griswoldia robusta and phanotea digitata griswold (zoropsidae), lepthercus rattrayi hewitt (nemesiidae), various lycosids, fuchiba and fuchibotulus spp. (cori n n idae) a nd drassodella vasivulva tucker (gallieniellidae) were common. original research haddad & dippenaar-schoeman koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 14 9 (page number not for citation purposes) 20 vol. 51 no. 1 page 4 of 9 opistacanthus capensis thorell (liochelidae) and uroplectes lineatus (c. l. koch) (buthidae) were often collected under logs and rocks. the fynbos (fb) fauna was dominated primarily by lycosids (particularly pardosa, trabea and zenonina spp.), drassodella vasivulva, various gnaphosids (camillina, xerophaeus and zelotes spp.), philodromus guineensis millot and suemus punctatus lawrence (philodromidae) and afrilobus sp. (orsolobidae). large numbers of pseudoscorpiones were collected by sifting leaf litter of protea spp.. the fauna at de hoop vlei (wl) was strongly dominated by gnaphosids (zelotes and xerophaeus spp., and drassodes ereptor purcell), lycosids (geolycosa and pardosa spp.), and heliophanus spp. (salticidae). various gnaphosids, corinnids and pseudoscorpions were common in sifted leaf litter of sideroxylon inerme (milkwood) trees near to the wetland. in the intertidal zone (iz), only two species were particularly common. amaurobioides africanus hewitt (anyphaenidae) was commonly found in retreats constructed in sandstone formations at the back end of the intertidal zone, while desis formidabilis (o.p.-cambridge) (desidae) was occasionally collected from beneath limpet shells and between algae on the rocky shores. these two species are regarded as marine specialists, occurring only in association with the intertidal zone along rocky shores (lamoral 1968). ground web-builders: web-builders were generally uncommon on the ground surface, but several species can be singled out. in cd leaf litter, hahnia spp. (hahniidae) were frequently found in their sheet-webs, while in fb leaf litter, benoitia ocellata (pocock) (agelenidae) and various linyphiids were common. lamaika sp. and vidole capensis (pocock) (phyxelididae) were frequently collected in leaf litter and under logs in the ep. the most common web-builders in the wl were steatoda capensis hann and euryopis sp. 1 (theridiidae), while very few web-builders were collected from the ground level in iz. arachnids associated with bark: due to the vegetative structure of fynbos, very few large shrubs and trees are found in most of the habitats sampled. only the ep contained eucalyptus trees that were large enough to sample arachnids from under bark. common wandering arachnids collected include clubiona spp. (clubionidae), aneplasa sculpturata tucker, poecilochroa anomala (hewitt) and upognampa aplanita tucker (gnaphosidae), pseudicius spp. and menemerus bivittatus (dufour) (salticidae), platyoides quinquedentatus purcell (trochanteriidae), cetonana martini (simon) (corinnidae) and uroplectes lineatus (buthidae). dominant web-dwelling spiders include theridion spp. (theridiidae) and neoscona subfusca (c.l. koch) (araneidae). interestingly, several specimens of the tree trapdoor spider moggridgea peringueyi simon (migidae) were collected from their silken burrows under bark. foliage wanderers: the fauna of cd was dominated by massagris regina wesolowska and heliophanus sp. (salticidae) and predominantly immature palystes superciliosus l. koch (sparassidae). wandering spiders were quite rare in wl, comprising primarily of heliophanus spp., various philodromids, and ground-dwelling lycosids (particularly pardosa spp.) that had wandered onto short vegetation. in ep, various salticids (massagris regina, thyene and heliophanus spp.), oxyopes and hamataliwa spp. (oxyopidae), synema spp. (thomisidae), immature tibellus minor lessert (philodromidae) and clubiona spp. (clubionidae) were collected from short shrubs and creepers. the fb plant-dwellers were considerably more diverse. the most common species collected include chariobas spp. (zodariidae), various thomisids (tmarus, thomisus and misumena spp.), and salticids (thyene and menemerus spp.). foliage web-dwellers: web-dwellers in the cd and fb were particularly dominated by neoscona and cyclosa spp. (araneidae), theridion spp. and various linyphiids. several rare species were also collected in the fb and ep, particularly. the only common web-dweller near the figure 8 species diversity of spider families collected in the de hoop nature reserve as ranked from highest to lowest. black bars indicate wandering spiders and grey bars indicate web-builders checklist of the non-acarine arachnids original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #149 (page number not for citation purposes) 21vol. 51 no. 1 page 5 of 9 iz was larinia natalensis (grasshoff) (araneidae), which constructs its orb-web in creepers and other vegetation between rocky outcrops surrounding the intertidal zone. conclusion this study provides the first intensive data on spider diversity in the fynbos biome, although two studies have previously been conducted in this vegetation type (coetzee et al. 1990; visser et al. 1999). in total, 274 species of arachnids were collected, with spiders the dominant group (252 species). this diversity represents approximately 12.5% of the currently known south african fauna of approximately 2000 species (dippenaarschoeman & haddad, unpubl.). while the species diversity is slightly lower than surveys conducted in the savanna biome, it compares favourably with studies conducted in the succulent and nama karoo biomes. the relatively high number of arachnid species collected, and the presence of several fynbos endemics (e.g. 10 of the 15 corinnidae species), supports the generalised perception that fynbos contains a unique fauna and flora. the only spiders currently considered to be of conservation importance are the baboon spiders, harpactira cafreriana (walkenaer) and harpactirella sp. both species are relatively common under rocks and within tussocks of thamnochortis grasses and populations are unlikely to be threatened by occasional collecting. perhaps also worth noting was the unusual stasimopes sp. (trapdoor spider), of which only males were collected. these have unusual spine-like tubercles in the eye region, something which could not be traced to any described species in the literature. consequently, this species may possibly be new or an undescribed male of a described species. the scorpions collected all have a relatively broad distribution within the western cape province (prendini pers. comm.). for example, parabuthus planicauda (pocock) was recorded from dhnr by prendini (2004), but is widespread throughout the western and eastern cape provinces. the occurrence of these scorpions within a protected area such as dhnr can be considered important for the conservation of the species, particularly when the growing threats to the fynbos biome are considered. in this study several new species and three new genera were collected, some of which have recently been described (haddad 2006; haddad & lyle 2008). this study expanded the distribution ranges known for many species, and provided valuable material for future taxonomic studies. this emphasises the need to expand efforts to survey the arachnid faunas of conservancies throughout south africa, but particularly within the western cape province, where invertebrate endemism may be relatively high compared to other areas. acknowledgements capenature is thanked for permits to collect in the reserve. peter chadwick and keith spencer at dhnr are thanked for their support and interest in the project. the following taxonomic specialists are thanked for providing identifications of material of their respective taxa: lorenzo prendini, american museum of natural history, new york, u.s.a. (scorpiones); leon lotz, national museum, bloemfontein, south africa (miturgidae, opiliones, sicariidae); rudy jocqué and mark alderweireldt, royal museum for central africa, tervuren, belgium (linyphiidae and zodariidae, and lycosidae, respectively); mark harvey, western australian museum, perth, australia (some pseudoscorpiones); wanda wesołowska, wrocław university, wrocław, poland (salticidae); and bernhard huber, zoological institute and museum alexander koenig, bonn, germany (pholcidae). andor venter, johan venter and johan du preez (university of the free state, bloemfontein) kindly provided details of the floral composition of the habitats. several records, collected by norman larsen and listed on the capenature invertebrate species database for dhnr, are also acknowledged. references coetzee, j.h., dippenaar-schoeman, a.s. & van den berg, a., 1990, ‘spider assemblages on five species of proteaceous plants in the fynbos biome of south africa’, phytophylactica 22, 443–447. churchill, t.b., 1997, ‘spiders as ecological indicators: an overview for australia’, memoirs of the museum of victoria 56(2), 331–337. dippenaar, s.m., modiba, m.a., khoza, t.t. & dippenaarschoeman, a.s., 2008, ‘a checklist of the spiders (arachnida, araneae) of the polokwane nature reserve, limpopo province, south africa’, koedoe 50, 10–17. dippenaar-schoeman, a.s., 2006, ‘new records of 43 spider species from the mountain zebra national park, south africa (arachnida: araneae)’, koedoe 49, 23–28. dippenaar-schoeman, a.s. & craemer, c., 2000, ‘the south african national survey of arachnida’, plant protection news 56, 11–12. dippenaar-schoeman, a.s. & leroy, a., 2003, ‘a checklist of the spiders of the kruger national park, south africa (arachnida: araneae)’, koedoe, 46, 91–100. dippenaar-schoeman, a.s., leroy, a, de jager, m. & van den berg, a., 1999, ‘a checklist of the spider fauna of the karoo national park, south africa (arachnida: araneae)’, koedoe 42, 31–42. dippenaar-schoeman, a.s., van den berg, a.m. & van den berg, a., 1989, ‘species composition and relative seasonal abundance of spiders from the field and tree layers of the roodeplaat dam nature reserve’, koedoe 32, 25–38. dippenaar-schoeman, a.s., van der walt, a.e., de jager, m., le roux, e. & van den berg, a., 2005, ‘the spiders of the swartberg nature reserve in south africa (arachnida: araneae)’, koedoe 48, 77–86. foord, s.h., dippenaar-schoeman, a.s. & van der merwe, m., 2002, ‘a checklist of the spider fauna of the western soutpansberg, south africa (arachnida: araneae)’, koedoe 45, 35–43. fromhage, l., jacobs, k. & schneider, j.m., 2007, ‘monogynous mating behaviour and its ecological basis in the golden orb spider’, nephila fenestrata. ethology 113(8), 813–820. germishuizen, g., meyer, n.l., steenkamp, y. & keith, m., 2006, a checklist of south african plants, south african botanical diversity network, pretoria. giliomee, j.h., 2003, ‘insect diversity in the cape floristic region’, african journal of ecology 41, 237–244. haddad, c.r., 2006, ‘spinotrachelas, a new genus of tracheline sac spiders from south africa (araneae: corinnidae)’, african invertebrates 47, 85–93. haddad, c.r., dippenaar-schoeman, a.s. & wesolowska, w., 2006, ‘a checklist of the non-acarine arachnids (chelicerata: arachnida) of the ndumo game reserve, maputaland, south africa’, koedoe 49, 1–22. haddad, c.r. & lyle, r., 2008, ‘three new genera of tracheline sac spiders from southern africa (araneae: corinnidae)’, african invertebrates 49, 37–76. lamoral, b.h., 1968, ‘on the ecology and habitat adaptations of two intertidal spiders, desis formidabilis (o pcambridge) and amaurobioides africanus hewitt, at the island (kommetjie, cape peninsula), with notes on the occurrence of two other spiders’, annals of the natal museum 20, 151–193. linder, h.p., 2005, ‘evolution of diversity: the cape flora’, trends in plant science 10, 536–541. mcneely, j., 2001, ‘invasive species: a costly catastrophe for native biodiversity’, land use and water resources research 1(2), 1–10. midgley, g.f., hannah, l., millar, d., thuiller, w. & booth, a., 2003, ‘developing regional and species-level assessments of climatic change impacts on biodiversity in the cape floristic region’, biological conservation 112(1/2), 87–97. original research haddad & dippenaar-schoeman koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 14 9 (page number not for citation purposes) 22 vol. 51 no. 1 page 6 of 9 modiba, m.a., dippenaar, s.m. & dippenaar-schoeman, a.s., 2005, ‘a checklist of spiders from sovenga hill, an inselberg in the savanna biome, limpopo province, south africa (arachnida: araneae)’, koedoe 48(2), 109–115. picker, m.d. & samways, m.j., 1996, ‘faunal diversity and endemicity of the cape peninsula, south africa – a first assessment’, biodiversity and conservation 5(5), 591–606. prendini, l., 2004, ‘the systematics of southern african parabuthus pocock (scorpiones, buthidae): revisions to the taxonomy and key to the species’, journal of arachnology 32(1), 109–186. procheş, s. & cowling, r.m., 2006, ‘insect diversity in cape fynbos and neighbouring south african vegetation’, global ecology and biogeography 15(5), 445–451. richardson, d.m., van wilgen, b.w., higgins, s.i., trindersmith, t.h., cowling, r.m. & mckell, d.h., 1996, ‘current and future threats to plant biodiversity on the cape peninsula, south africa’, biodiversity and conservation 5(5), 607–647. sharratt, n.j., picker, m.d. & samways, m.j., 2000, ‘the invertebrate fauna of the sandstone caves of the cape peninsula (south africa): patterns of endemism and conservation priorities’, biodiversity and conservation 9(1), 107–143. visser, d., wright, m.g., van den berg, a. & giliomee, j.h., 1999, ‘species richness of arachnids associated with protea nitida (proteaceae) in the cape fynbos’, african journal of ecology, 37(3), 334–343. whitmore, c., slotow, r., crouch, t.e. & dippenaar-schoeman, a.s., 2002, ‘diversity of spiders (araneae) in a savanna reserve, northern province, south africa’, journal of arachnology 30(2), 344–356. witt, a.b.r. & samways, m.j., 2004, ‘influence of agricultural land transformation and pest management practices on the arthropod diversity of a biodiversity hotspot, and cape floristic region, south africa’, african entomology 12(1), 89–95. wright, m.g. & samways, m.j., 1996, ‘gall-insect species richness in african fynbos and karoo vegetation: the importance of plant species richness’, biodiversity letters 3(4/5), 151–155. wright, m.g. & samways, m.j., 1999, ‘plant characteristics determine insect borer assemblages on protea species in the cape fynbos, and the importance for conservation management’, biodiversity and conservation 8(8), 1089–1100. appendix 1 a checklist of the non-acarine arachnids of the de hoop nature reserve. guild abbreviations are provided in the text. habitat abbreviations: cd – coastal dunes; ep – eucalyptus plantation; fb – fynbos; iz – intertidal zone; wl – wetlands. symbols: † indicates a new species, ‡ indicates a possible new species, and ? indicates a dubious identification. family/genus/species guilds habitats order: araneae (spiders) family: agelenidae benoitia ocellata (pocock 1900) fwb fb family: anapidae crozetulus rhodesiensis (brignoli 1981) owb fb family: anyphaenidae amaurobioides africana (hewitt 1917) gw iz family/genus/species guilds habitats family: araneidae araneus apricus (karsch 1884) owb ep a. nigroquadratus (lawrence 1937) owb ep argiope trifasciata (forskål 1775) owb wl caerostris sexcuspidata (fabricius 1793) owb ep, wl cyclosa insulana (costa 1834) owb cd, ep, fb c. oculata (walckenaer 1802) owb fb cyrtophora citricola (forskål 1775) owb fb gea infuscata (tullgren 1910) owb wl ideocaira transversa (simon 1903) owb ep isoxya cicatricosa (c.l. koch 1844) owb fb kilima sp.‡ owb wl larinia natalensis (grasshoff 1971) owb fb, iz lipocrea longissima (simon 1881) owb fb, wl nemoscolus tubicola (simon 1887) owb wl neoscona rufipalpis (lucas 1858) owb wl n. subfusca (c.l. koch 1837) owb cd, ep, fb paralarinia bartelsi (lessert 1933) owb fb prasonica sp.? owb fb family: caponiidae caponia capensis (purcell 1904) gw/pwb cd, ep, fb, wl family: clubionidae clubiona abbajensis (strand 1906) gw/pwb ep, fb, wl clubiona sp. 2 pwb ep, fb family: corinnidae apochinomma sp.† gw fb castianeira fulvipes (simon 1896) gw cd, ep, fb cetonana martini (simon 1896) gw/pwb ep, fb cetonana sp. 2† gw ep cetonana sp. 3† gw fb cetonana sp. 4† gw fb copa flavoplumosa (simon 1885) gw cd, ep, fb fuchiba capensis (haddad & lyle 2008) gw ep, fb, wl fuchibotulus bicornis (haddad & lyle 2008) gw ep, fb, wl graptartia tropicalis (haddad 2004) gw cd, ep, fb orthobula infima (simon 1897) gw cd, ep, fb, wl pronophaea natalica (simon 1897) gw ep spinotrachelas capensis (haddad 2006) gw ep, fb, wl trachelas sp. 1† pwf fb trachelas sp. 2† pwf fb family: ctenidae thoriosa sp.‡ gw ep, fb family: ctenizidae stasimopus sp.‡ gw ep, fb family: cyatholipidae cyatholipus quadrimaculatus (simon 1894) gwb ep cyatholipus sp. 2‡ gwb ep, fb ulwembua denticulata (griswold 1987) owb ep family: cyrtaucheniidae homostola reticulata (purcell 1902) gw ep family: deinopidae avellopsis capensis (purcell 1904) mowb ep, fb menneus camelus (pocock 1902) mowb ep, fb family: desidae desis formidabilis (o.p.-cambridge 1890) gw iz family: dictynidae archaeodictyna sp. hwb fb dictyna sp. 1 hwb fb dictyna sp. 2 hwb fb family: eresidae dresserus collinus (pocock 1900) swb ep, fb gandanameno spenceri (pocock 1900) swb ep, fb appendix 1 (cont...) checklist of the non-acarine arachnids original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #149 (page number not for citation purposes) 23vol. 51 no. 1 page 7 of 9 appendix 1 (cont...) family/genus/species guilds habitats family: gallieniellidae drassodella quinquelabecula (tucker 1923) gw fb d. vasivulva (tucker 1923) gw cd, ep, fb family: gnaphosidae aneplasa sculpturata (tucker 1923) gw/pwb ep, fb aphantaulax stationis (tucker 1923) gw cd asemesthes sp. imm. gw cd camillina corrugata (purcell 1907) gw ep, fb c. pavesii (simon 1897) gw ep, fb, wl c. procurva (purcell 1908) gw ep, fb drassodes ereptor (purcell 1907) gw wl echeminae sp. indet. gw pw echemus sp. imm. gw wl megamyrmaekion schreineri (tucker 1923) gw wl micaria sp. gw cd, fb poecilochroa anomala (hewitt 1915) gw/pwb ep, wl setaphis subtilis (simon 1897) gw ep upognampa aplanita (tucker 1923) gw/pwb ep, wl xerophaeus capensis (purcell 1907) gw fb x. crusculus (tucker 1923) gw cd, ep, fb, wl x. phaseolus (tucker 1923) gw ep, fb zelotes anchora (tucker 1923) gw cd, ep, fb, wl z. capsula (tucker 1923) gw ep, wl z. fuligineus (purcell 1907) gw ep, fb, wl z. montanus (purcell 1907) gw ep, fb family: hahniidae hahnia clathrata (simon 1898) swb fb h. tabulicola (simon 1898) swb cd, ep, fb hahnia sp. 3‡ swb ep family: idiopidae idiopidae sp. gw ep family: liocranidae rhaeboctesis sp. gw fb family: linyphiidae callitirchia sp. swb cd, fb ceratinopsis dippenaari (jocqué, 1984?) swb cd, fb linyphiidae sp. 1 swb fb linyphiidae sp. 2 swb fb linyphiidae sp. 3 swb fb linyphiidae sp. 4 swb fb mecynidis sp.† swb fb meioneta sp. swb fb metaleptyphantes sp. swb fb microlinyphia sterilis (pavesi 1883) swb ep, fb ostearius melanopygius (o.p.-cambridge 1879) swb wl family: lycosidae arctosa sp. gw cd hogna sp. gw ep, fb, wl lycosa sp. gw ep pardosa sp. 1 gw cd pardosa sp. 2 gw cd proevippa albiventris (simon 1898) gw wl trabea purcelli (roewer 1951) gw cd, wl t. rubriceps (lawrence 1952) gw ep, fb, wl trochosa sp.? gw wl zenonina sp. gw ep, fb, wl family: migidae moggridgea peringueyi (simon 1903) pwb ep family: mimetidae ero sp. pwf ep mimetus sp. 1‡ pwf ep mimetus sp. 2‡ pwf wl family/genus/species guilds habitats family: miturgidae cheiramiona ansiae (lotz 2002) pwf fb family: nemesiidae lepthercus rattrayi (hewitt 1917) gw cd, ep, fb, wl pionothele sp.† gw ep family: nephilidae nephila fenestrata (thorell 1859) owb fb family: oecobiidae oecobius navus (blackwall 1859) pwb cd, fb family: oonopidae gamasomorpha humicola (lawrence 1947) gw fb oonopinae sp. gw ep, fb opopaea speciosa (lawrence 1952) gw cd, ep, fb, wl family: orsolobidae afrilobus sp.† gw cd, ep, fb family: oxyopidae hamataliwa kulczynski (lessert 1915) pwf ep, fb hamataliwa sp. 2 pwf ep, fb oxyopes russoi (caporiacco 1940?) pwf ep oxyopes sp. 2 imm. pwf ep family: palpimanidae palpimanus sp. 1 gw ep, fb, wl palpimanus sp. 2 gw ep family: philodromidae philodromus guineensis (millot 1941) gw fb suemus punctatus (lawrence 1938) gw cd, ep, fb, wl tibellus minor (lessert 1919) pwf ep, fb family: pholcidae quamtana sp. spwb cd, fb smeringopus sp. spwb ep, fb family: phyxelididae lamaika sp.† hwb ep, fb vidole capensis (pocock 1900) hwb ep, fb family: pisauridae chiasmopes sp. imm. pwf fb cispius sp. pwf fb euprosthenopsis sp. imm. pwf fb rothus purpurissatus (simon 1898) pwf ep, fb thallassius spinossissimus (karsch 1879) gw wl family: prodidomidae prodidomus capensis (purcell 1904) gw fb theuma ababensis (tucker 1923) gw ep t. capensis (purcell 1907) gw fb t. schreineri (purcell 1907?) gw fb family: salticidae asemonea sp. pwf ep baryphas ahenus (simon, 1902) pwf fb dendryphantes purcelli (peckham & peckham 1903) pwf ep euophrys purcelli (peckham & peckham 1903) gw fb euophrys sp. 2‡ gw ep, fb evarcha dotata (peckham & peckham 1903) pwf ep habrocestum sapiens (peckham & peckham 1903) gw fb habrocestum sp. 2 gw ep heliophanus claviger (simon 1901) pw fb h. modicus (peckham & peckham 1903) gw ep, fb, wl h. patellaris (simon 1901) gw wl heliophanus sp. 4 gw/pwf cd, iz massagris regina (wesolowska 1993) gw cd, ep, fb, iz, wl menemerus bivittatus (dufour 1831) pwb ep menemerus sp. 2 pwf fb myrmarachne leleupi (wanless 1978) gw cd, fb myrmarachne sp. 2 gw fb appendix 1 (cont...) table continues on the next column original research haddad & dippenaar-schoeman koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za a rt ic le # 14 9 (page number not for citation purposes) 24 vol. 51 no. 1 page 8 of 9 family/genus/species guilds habitats family: thomisidae avelis hystriculus (simon 1895)? pwf ep diaea sp.† pwf ep, fb firmicus abnormis (lessert 1923) pwf ep, fb f. bragantinus (brito capello 1866) pwf fb heterogriffus berlandi (lessert 1938) pwf ep, fb heterogriffus sp. 2‡ pwf fb holopelus almiae (dippenaar-schoeman 1986) pwf fb monaeses pustulosus (pavesi 1895) pwf fb oxytate argenteooculata (simon 1886) pwf ep, fb pactactes obesus (simon 1895) gw cd, ep, fb, wl pherecydes tuberculatus (o.p.-cambridge 1883) pwf fb pherecydes sp. 2† pwf ep, fb phrynarachne melloleitoa (lessert 1933) pwf ep p. rugosa (latreille 1804) gw ep runcinia aethiops (simon 1901) pwf ep, fb simorcus capensis (simon 1895) pwf fb stiphropus sp. gw fb synema abnorme (lessert 1923) pwf ep, fb s. decens (karsch 1878) pwf ep, fb s. nigrotibiale (lessert 1919) pwf ep, fb thomisus australis (comellini 1957) pwf fb t. stenningi (pocock 1900) pwf fb tmarus comellinii (garcia-neto 1989) pwf ep, fb t. foliatus (lessert 1928) pwf fb tmarus sp. 3‡ pwf ep, fb xysticus lucifugus (lawrence 1937) gw ep, fb family: trochanteriidae platyoides leppanae (pocock 1902) pwb ep p. quinquedentatus (purcell 1907) pwb ep family: uloboridae miagrammopes brevicaudus (o.p.-cambridge 1882) mowb ep uloborus sp. imm. owb cd, ep, fb family: zodariidae caesetius globicoxis (lawrence 1942) gw ep, fb chariobas cylindraceus (simon 1893)? pwf ep, fb chariobas sp. 2‡ pwf fb chariobas sp. 3‡ pwf fb cyrioctea griswoldorum (platnick & jocqué 1993) gw ep, fb diores simoni (o.p.-cambridge 1904)? gw cd, fb, wl heradida extima (jocqué 1987) gw wl procydrela procursor (jocqué 2000) gw fb psammorygma sp. gw fb ranops sp.? gw cd rotundrela rotunda (jocqué 2000) gw ep, fb systenoplacis sp.‡ gw ep, fb family: zoridae voraptus sp. gw/pwf ep, fb family: zoropsidae griswoldia robusta (simon 1898) gw cd, ep, fb, iz, wl machadoniinae sp. gw fb phanotea digitata (griswold 1994) gw cd, ep, fb order: opiliones (harvestmen) family: caddidae caddella sp.† gw cd family: phalangiidae rhampsinitus vittatus (lawrence 1931)? gw cd, ep, fb family: triaenonychidae adaeum spatulatum (lawrence 1931) gw ep, fb, wl ceratomontia annae (lawrence 1934) gw fb c. karooensis (lawrence 1931) gw fb, wl c. minor (lawrence 1931) gw fb, wl family/genus/species guilds habitats natta chionogastra (simon 1901) gw ep, fb n. horizontalis (karsch 1879) gw cd, ep, fb pellenes geniculatus (simon, 1868)? gw fb, wl phintella aequipes (peckham & peckham 1903) gw ep phlegra sp.? gw ep pignus sp.‡ gw ep pseudicius africanus (peckham & peckham 1903) pwb ep pseudicius sp. 2 pwf fb rhene sp. imm. pwf fb salticidae sp. indet. 1 pwf fb salticidae sp. indet. 2 gw cd thyene inflata (gerstaecker 1873) pwf ep, fb t. ogdeni (peckham & peckham 1903?) pwf ep, fb thyene sp. 3 pwf fb thyenula sp.? gw ep family: scytodidae scytodes cedri (purcell 1904) gw cd, ep, fb, wl scytodes sp. 2 gw ep family: segestriidae ariadna sp. twb fb family: selenopidae anyphops capensis (lawrence 1940) pwb ep, fb anyphops sp. 2 pwb ep, fb, wl family: sicariidae loxosceles spinulosa (purcell 1904) gw ep, fb loxosceles sp.‡ gw ep sicarius spatulatus (pocock 1901) gw ep, fb family: sparassidae olios sp. 1 pwf fb olios sp. 2 pwf fb palystes castaneus (latrielle 1819) pwf ep, fb p. superciliosus (l. koch 1875) pwf cd, ep, fb panaretella sp. pwf fb pseudomicrommata sp. pwf fb family: tetragnathidae leucauge festiva (blackwall 1866) owb ep, fb, wl l. levanderi (kulzcynski 1901) owb ep, fb, wl tetragnatha ceylonica (o.p.-cambridge 1869) owb ep, fb tetragnatha sp. 2 owb ep family: theraphosidae harpactira cafreriana (walkenaer 1837) gw ep, fb harpactirella sp. gw fb family: theridiidae achaearanea sp. gwb ep anelosimus sp. 1 gwb fb anelosimus sp. 2 gwb fb dipoena sp. 1 gwb cd, ep, fb dipoena sp. 2 gwb fb dipoenura sp. gwb fb euryopis sp. 1 gwb fb, wl euryopis sp. 2 gwb fb latrodectus geometricus (c.l. koch 1841) gwb ep, fb l. indistinctus (o.p.-cambridge 1904) gwb ep pholcomma sp.? gwb fb phoroncidia capensis (simon 1895)? gwb ep steatoda capensis (hann 1990) gwb ep, fb, iz theridion delicatum (o.p.-cambridge 1904) gwb ep, fb theridion sp. 2 gwb ep, fb theridion sp. 3 gwb ep family: theridiosomatidae theridiosomatidae sp. owb fb appendix 1 (cont...) appendix 1 (cont...) checklist of the non-acarine arachnids original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoe a rticle #149 (page number not for citation purposes) 25vol. 51 no. 1 page 9 of 9 appendix 1 (cont...) family/genus/species guilds habitats anaulacodithella angustimana (beier 1955) gw fb order: scorpiones (scorpions) family: buthidae parabuthus planicauda (pocock 1889) gw cd, ep, fb uroplectes lineatus (c.l. koch 1844) gw/pwb ep, fb, wl family: liochelidae opistacanthus capensis (thorell 1877) gw ep, fb family: scorpionidae opistophthalmus macer (thorell 1877) gw ep order: solifugae (sun-spiders) family: solpugidae solpugema sp. imm. gw fb family/genus/species guilds habitats larifuga granulosa (lawrence 1931) gw ep, fb triaenonychidae sp. imm. gw ep order: pseudoscorpiones (false scorpions) family: atemnidae cyclatemnus sp. gw iz family: cheliferidae beierius simplex (beier 1955) gw fb b. walliskewi (ellingsen 1912) gw fb hansenius sp. gw ep family: chernetidae caffrowithius biseriatus (mahnert 1983) gw fb c. natalensis (beier 1947) gw fb pselaphochernes natalensis (beier 1947) gw fb family: geogarypidae geogarypus purcelli (ellingsen 1912) gw ep, iz family: tridenchthoniidae article information authors: alan s. barrett1 leslie r. brown1 louise barrett1,2 peter henzi1,2 affiliations: 1applied behavioural ecology and ecosystem research unit, department of environmental sciences, university of south africa, south africa 2department of psychology, university of lethbridge, canada correspondence to: leslie brown email: lrbrown@unisa.ac.za postal address: applied behavioural ecology and ecosystem research unit, department of environmental sciences, unisa, florida campus, private bag x6, florida 1710, south africa keywords braun-blanquet; phenology; plant communities; resource energy; twinspan dates: received: 24 feb. 2010 accepted: 14 aug. 2010 published: 26 oct. 2010 how to cite this article: barrett, a.s., brown, l.r., barrett, l. & henzi, p., 2010, ‘a floristic description and utilisation of two home ranges by vervet monkeys at the loskop dam nature reserve, south africa’, koedoe 52(1), art. #990, 12 pages. doi: 10.4102/koedoe.v52i1.990 copyright notice: © 2010. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0771 (online) a floristic description and utilisation of two home ranges by vervet monkeys in loskop dam nature reserve, south africa in this original research... open access • abstract • introduction    • climate • methods    • study area       • vegetation classification       • phenology • results    • vegetation classification    • donga home range       • description of the donga home range plant communities    • picnic home range    • ordination    • floristic analysis       • donga home range       • picnic home range    • resource items    • resource availability • discussion • conclusion • acknowledgements • references abstract (back to top) the plant communities occurring in the home ranges of two vervet monkey (chlorocebus aethiops) troops (referred to as the donga and picnic troops) were investigated as part of a comprehensive research project on the spatial and temporal patterns in resource dispersion or distribution and range use. from two-way indicator species analysis (twinspan) multivariate classifications refined by braun-blanquet procedures, seven plant communities that can be placed into three major groups were identified in the home range of the donga troop, and eight plant communities that can be placed into four major groups were identified in the home range of the picnic troop. classifications and descriptions of the major groups, including vegetation maps, are presented. diagnostic as well as prominent species of tree, shrub, forb and grass strata are outlined. ordinations and floristic analyses were undertaken for the two home ranges to highlight vegetative differences between the areas. the phenology of vervet resources and the available resource energy were determined as a basis for the comprehensive study. conservation implications: this vegetation description and phenology provide valuable information on the utilisation of home ranges by vervets and demonstrate how various management applications can affect the home range dependence of troops. introduction (back to top) the loskop irrigation dam was completed in 1938 to provide water for the irrigation needs of agriculture downstream. to ensure that the water users received good-quality water, it was critical for the area surrounding the dam to be managed appropriately for water quality conservation. this led to the establishment of the loskop dam nature reserve (ldnr) in 1942, with subsequent proclamation in 1954 (barrett 2009). currently, the mpumalanga tourism and parks agency (mtpa), whose mandate is the conservation of natural biodiversity for current and future generations through the sustainable use of natural resources, manages the reserve. the reserve falls within the savanna biome of south africa and, because of climate and topography, there are a variety of habitats on the reserve, making it possible to stock a large variety of game species. to ensure efficient management and to assist the managers in making evidence-based decisions, various research projects on different animals and vegetation are undertaken and encouraged in the reserve. no ecologically based studies on vervet monkeys have been undertaken in the ldnr to date, and only a few vegetation studies have been conducted in the reserve. theron (1973) conducted the first detailed vegetation study of the reserve and did a broad-scale analysis of the overall vegetation, using what appears to be a scale of 1:50 000 for his delineation of the plant communities. the vegetation of areas that have been added to the north-eastern section of the reserve (two farms called rietfontein and parys) have been classified and mapped by götze et al. (1998) using 1:30 000 aerial photographs. vervets are extremely adaptable, opportunistic generalists that have a wide distribution in sub-saharan africa (barrett et al. 2006). not many detailed ecological studies of vervets have been undertaken in temperate areas (isbell, pruetz & young 1998; pruetz & isbell 2000). at low latitudes, overall habitat productivity is high and seasonal variability is relatively limited (caughley & sinclair 1994). a fuller understanding of the flexibility of vervet ecology and the factors that influence their utilisation of their habitat will benefit the overall management of the species. no related vervet studies have been conducted locally. according to singh and kushwaha (2006), seasonal duration of leafing or flushing, flowering and fruiting determine phenological behaviour in tropical and subtropical trees. major phenological events may occur at various times during both the wet and the dry season (fensham & bowman 1992). sarmiento and monasterio (1983) and van schaik et al. (1993) observe that there may be considerable variation between years, species and life forms in the timing of such events. van schaik et al. (1993) state that most savanna woody plants produce new leaves and flowers in bursts rather than continuously, displaying seasonal variations in the presence of new leaves, flowers and fruits. since leaves, fruits and flowers comprise the bulk of primate diets, investigating the dynamics of these resources is valuable when attempting to understand the spatial and temporal habitat utilisation and ranging patterns of vervet monkeys. for many primates, and particularly for the vervet monkeys living in the ldnr, activity budgets, ranging patterns, timing of births and weaning are influenced by the spatial and temporal availability of food resources (barrett 2000). to date, no phenological data has been collected or reported for the ldnr. this study provides the basic framework for a comprehensive research project on how two troops of vervet monkeys (the donga troop (n = 14) and the picnic troop (n = 15)) use their home ranges to obtain temporally and spatially distributed resources through their ranging patterns. of interest is how the two home ranges vary floristically and in terms of available resources. in this article, the various plant communities occurring in the two troops’s home ranges are identified, described and mapped, and the general vegetation structure of the areas is determined. environmental factors influencing the plant communities are explored to determine whether there are specific associations between these factors and the respective plant communities. a floristic analysis of both the home ranges is undertaken and the results compared. the phenology of resource tree species used by the two troops is also determined and presented. the home ranges of the two troops that were studied are geographically separated and approximately 4.5 km apart. the donga home range is 224 ha in size and the picnic home range is 138 ha. the underlying geology of the donga home range comprises the sedimentary loskop formation and the selonsrivier formation, while the picnic home range consists only of the loskop formation. there are conspicuous topographical differences between the two home ranges, with the donga area being more rugged and hilly than the picnic area, which is more level and uniform in appearance. the average slope for the donga area is 13.6° compared to 10° for the picnic area. the donga plant communities occur at a wider range of altitudes than those of the picnic home range, with the donga altitudes ranging from 1005 m to 1105 m and the picnic altitudes ranging from 1031 m to 1097 m. according to mucina and rutherford (2006), the vegetation occurring on the mountains of the ldnr and surrounding areas belongs to the loskop mountain bushveld vegetation unit (svcb 13), whereas the vegetation occurring in the valley of the olifants river to the loskop dam and surrounding areas belongs to the loskop thornveld vegetation unit (svcb 14). the loskop mountain bushveld consists of open, broad-leaved woodland and the loskop thornveld of deciduous, open woodland (mucina & rutherford 2006). climate the ldnr is located in a summer rainfall area with moderate to very hot summers and mild to cold winters, and with frost occurring on the mountain tops and in the low-lying valley bottoms (theron 1973). temperatures range from a minimum of -8 °c to a maximum of 40 °c, with an average of 21 °c (low & rebelo 1998). rainfall occurs mainly in the form of short-lived, high-intensity thunderstorms associated with strong, south-westerly winds. the majority of the rainfall occurs in the summer (november–april). the average monthly rainfall for the wet season (november 2006 –april 2007) during the study period was 55 mm, compared to 87 mm for the period november 2002–april 2007 (figure 1). during the study period, the mean temperature for the dry season was 18.5 °c, compared to 24 °c for the wet season. a mean minimum temperature of 8.5 °c (july 2006) and a mean maximum of 32 °c (february 2007) were recorded for the study period. the average annual rainfall for the study area, as measured by weather stations located at various monitoring points on the reserve, was 654 mm for the period may 2002 to april 2007. the study period was drier for both the dry and the wet seasons compared to the average for the previous 5 years. average monthly rainfall for the dry season (may 2006–october 2006) during the study period was 6 mm, compared to 11 mm for may 2002–october 2006. the average climate of the two home ranges is similar due to their close proximity, yet the effects of climate and particularly of rainfall on the areas are not the same, due to topographical distinctions between the areas, such as slope and altitudinal differences. the picnic home range is less steep than the donga home range, resulting in surface water being stored for longer periods in a series of natural pools and small, man-made dams. there are no impoundments in the donga home range and the run-off rate into the loskop dam is rapid after rain has fallen. figure 1: average monthly rainfall and temperatures for the loskop dam nature reserve from may 2002 to april 2007 compared to those for the study period from may 2006 to april 2007 methods (back to top) study area the ldnr is located 52 km north of middelburg on the n11 national road to groblersdal in mpumalanga province (figure 2). according to emery, lotter and williamson (2004) and ferrar and lotter (2007), the reserve is currently 23 175 ha in size, of which 2350 ha is water surface. it is located between longitude 29°15’00” and 29°40’00” east and latitude 25°34’00” and 25°56’00” south. at the heart of the reserve is the loskop irrigation dam (approximately 24 km long), situated in the olifants river valley. the dam is fed predominantly by the perennial olifants river and the krantz spruit, and their respective tributaries (theron 1973). a network of small mountain streams and drainage channels originating from both inside and outside the boundaries of the reserve also drain into the dam. vegetation classification a series of 1:10 000 digitised orthophotos of the ldnr was used to stratify the study area into physiognomic-physiographic units, as described by barbour, burk and pitts (1987) and kent and coker (1997). to ensure that all variations in the vegetation were captured during the sampling, 69 sample plots (43 in the donga home range and 26 in the picnic home range) were placed in a random manner within the stratified units. plot sizes were fixed at 400 m2 (barbour et al. 1987; brown 1997, brown & bredenkamp 1994). vegetation-related fieldwork was undertaken during november and december 2005, january 2006 and december 2006. the two troops’s home range boundaries were determined from all gps co-ordinates collected for the troops over the entire study period (june 2005–june 2007). taxon names conform to those of germishuizen and meyer (2003). environmental data collected included altitude, latitude and longitude, aspect, slope, percentage surface rock cover, percentage erosion, basic soil type, evidence of animal activity and the effects of fire. all plant species growing in the sample plots were recorded, and cover abundance was assessed using a modified braun-blanquet cover abundance scale (barbour et al. 1987, mueller-dombois & ellenberg 1974). erosion was estimated using erosion classes from matthee and van schalkwyk (1984), and slope was estimated using the categories specified by barrett et al. (2006). floristic data were recorded according to braun-blanquet techniques and analysed using the turboveg suite (hennekens 1998), which includes the two-way indicator species analysis multivariate classification technique (twinspan) for deriving an initial approximation of the main plant communities (hill 1979). this technique is regarded as a successful approach to vegetation classification by a number of phytosociologists (bredenkamp & bezuidenhout 1995; brown, bredenkamp & van rooyen 1996; brown & bredenkamp 1994; cilliers 1998). the visual editor megatab (hennekens 1996) was used to generate phytosociological tables and further refinement of the classification was undertaken through the application of braun-blanquet procedures (barbour et al. 1987; kent & coker 1997; werger 1974). using the final phytosociological tables and habitat information collected during sampling in the field, different plant communities were described and ecologically interpreted. plant communities were identified using ‘diagnostic species’ as defined by westhoff & van der maarel (1978), who define species as diagnostic if they are present in at least 20% of the relevés of a particular plant community, whilst simultaneously being absent or less frequent in other communities. an adaptation of this definition was used in this study, where a species was considered diagnostic for a particular community if it was present within the following constancy parameters (grobler, bredenkamp & brown 2006): • ≥ 70% and with a constancy of ≤ 35% in all other communities • ≥ 50% and with a constancy of ≤ 25% in all other communities • ≥ 20% and with a constancy of ≤ 13% in all other communities. ‘dominant species’ are defined as all species that have cover values exceeding a predetermined cover threshold value in any relevé in a selected vegetation unit (westhoff & van der maarel 1978). the vegetation of the study area is relatively heterogeneous, with dominant species not restricted to specific threshold values, but rather to the species with the highest overall cover abundance in a community. figure 2: location of the loskop dam nature reserve in the mpumalanga province of south africa phenology resource trees (utilised for food by the vervets) were identified by following the two troops during habituation and during data collection for this comprehensive study (barrett 2009). to determine the number of vervet resource trees required for phenological monitoring, density figures for the tree species were used. ten per cent of the total density figure for each tree species was calculated and used as the number of trees to tag for that species. representative samples of each species for tagging were selected from plant communities that occurred in both the troops’s home ranges. in the field, these species were randomly selected and identified, locations were recorded with a gps device and individual trees were tagged with a numbered aluminium disc for data-collection purposes. initial phenology data were collected from all tagged trees; thereafter data were collected on a monthly basis for the duration of the study period. the quantification of available resources has been undertaken in tropical forests, where the focus was on the ecology of fruiting trees and the behaviour of frugivorous consumers (chapman 1990; chapman et al. 1992; clutton-brock 1977; leighton & leighton 1982; raemakers 1980; terborgh 1983). a combination of quantification methods used in the forest environments of uganda (chapman et al. 1992) and in the savanna environments of south africa (smit 1989a, 1989b) were used for this study. direct measurement techniques were not an option for collecting reliable tree dimension estimates quickly and accurately because of the dense nature of the vegetation and the inaccessibility of many trees sampled in the areas frequented by the troops. the inexpensive, relatively quick ‘one-man’ method employed by smit (1996) was modified and linear tree dimension data were collected using improvised equipment. instead of using a calibrated 35-mm photographic slide, as suggested by smit (1996), a clear fibreglass reader with markings matching the markings on a 3-m measuring stick was used. within the 3-m markings, the reader was subdivided into 0.25-m segments for refined measurements. luminous yellow tape was used to mark the bottom and the 3-m point on the measuring stick. to measure a particular tree, the measuring stick was placed vertically adjacent to the trunk of the tree at ground level. the observer then paced away from the stick until the stick’s markings lined up to the 3-m markings on the reader when held at arm’s length from the observer. at this point, the reader was used to measure all linear tree dimensions in 3-m and/or 0.25-m intervals, whichever was required. if a tree was on a slope, its height was calculated on the uphill side. initially, a clinometer was used for calibration testing of the modified ‘one-man’ method, particularly for tree heights. the measurements corresponded well and the results obtained from the fibreglass reader were constantly within 0.5 cm of the clinometer readings (barrett 2009). according to chapman et al. (1992), the various methods of estimating resource abundance (particularly fruit) for biological studies are all susceptible to potential errors and no particular method is more suitable than any other. various authors have a predisposition towards specific methods based on personal preference, the objectives of the research being undertaken or the repeatability of the method(s) being employed. as the ldnr falls within the savanna biome, and because the nature of savanna trees differs significantly from those occurring in forest environments (hoffman & franco 2003; hoffman et al. 2005), a combination of visual estimates, actual fruit counts and canopy volume extrapolation was used in this study. only one observer collected all the phenological data over the study period, hence there was no need to do any inter-observer variability calculations or adjustments. results (back to top) vegetation classification for the purposes of this paper, only the major plant communities of the home ranges of the study troops are described. more detailed classification and descriptions of the sub-communities and their associated habitats can be found in barrett (2009). all references to species groups refer to tables 1 and 2 (see online supplement). donga home range the donga home range could typically be classified as acacia caffra–rhus leptodictya woodland; however, for the purposes of the comprehensive vervet study, it was necessary to do a more detailed vegetative analysis of the area. noteworthy species occurring within the home range include the small, erect, deciduous tree dombeya rotundifolia, the tree faurea saligna, the deciduous shrub dichrostachys cinerea, the erect shrub lippia javanica, the spiny, multi-stemmed shrub rhus pyroides, the multi-stemmed grewia spp. shrubs, the shrub euclea crispa, and the tufted, perennial grasses setaria sphacelata var. sphacelata and panicum maximum.seven plant communities that can be grouped into the following three major communities were identified in the donga troop’s home range, which are shown in figure 3 (figure 3 and table 1, see online supplement): 1. lippia javanica–loudetia simplex shrubland 2. acacia nilotica–acacia caffra woodland 3. olea europea subsp. africana–rhus leptodictya woodland. figure 3: vegetation map of the donga home range depicting the main plant communities description of the donga home range plant communities the donga home range is characterised by a large donga (gully) cutting through its centre. the large donga has several, smaller, subsidiary dongas that merge into it at various locations along its course as it winds down towards the loskop dam. vegetation in and adjacent to the donga was characterised by large riverine trees and shrubs with a sparse to non-existent grass and forb layer. areas between the dongas were covered by open woodland. 1. lippia javanica–loudetia simplex shrubland: this shrubland community covers approximately 25% (55 ha) of the donga home range and consists of relatively rocky terrain that varies from gentle, low-lying valley bottoms to steeper midslopes. there were signs of trampling, and several animal tracks were observed throughout this community. evidence of fire damage could be seen in places, but this was not recent and was likely due to the burning programme that is in place in the reserve. soils were sandy and well drained. rock cover ranged from 10% to 90%, with an average of 59%, and erosion was estimated at 22% (slight). altitude ranged from 1018 m to 1105 m, with the average slope being 16°. species belonging to species group a were diagnostic for this community (table 1, see online supplement). the woody vegetation is dominated by the large shrub ximenia caffra (species group a). the tree f. saligna and the erect shrub l. javanica (species group j) are prominent. the grass layer is dominated by the tufted perennial grasses s. sphacelata var. sphacelata (species group j), loudetia simplex (species group a) and hyparrhenia hirta (species group e). the average number of plant species recorded in this community was 24 per 400 m2. the tree layer has a canopy cover ranging between 2% and 50%, the shrub layer 4% and 15%, the grass layer between 21% and 70%, and the forb layer, between 1% and 2%. the tree f. saligna had the highest density of 472 individuals per ha, followed by the shrub d. cinerea with 405 individuals per ha and the tree a. caffra with 215 individuals per ha. two sub-communities were distinguished, namely the lippia javanica–loudetia simplex–sclerocarya birrea woodland and lippia javanica–loudetia simplex–acacia caffra woodland. 2. acacia nilotica–acacia caffra woodland: this woodland community covers approximately 52% (117 ha) of the donga home range and had relatively little rock cover, although there were sections that had a high cover. rock cover ranged from 1% to 90%, with an average of 17%. the erosion was estimated at 15% (slight), and altitude ranged from 1015 m to 1090 m, with an average slope of 8°. species belonging to species group f are diagnostic for this community (table 1, see online supplement). the woody vegetation is dominated by the smallto medium-sized deciduous tree a. caffra and the shrub e. crispa (species group m). the medium-sized tree r. leptodictya (species group m), the small, erect, deciduous tree d. rotundifolia (species group j), the shrub acacia nilotica (species group f), and the deciduous shrub d. cinerea (species group j) are all prominent. in the grass layer, the tufted perennial grass p. maximum (species group l) is locally prominent. the average number of plant species was 27 per 400 m2 in this community. the canopy cover for the tree layer ranged between 17% and 60%, the shrub layer between 17% and 57%, with the grass layer relatively high, ranging between 25% and 77%. the forb layer is not well developed and covers between 1% and 5% of the area. the tree a. caffra had the highest density of 1358 individuals per ha, followed by the shrub e. crispa with 1251 individuals per ha, and the shrub a. nilotica with 1205 individuals per ha. three sub-communities were distinguished, namely the acacia nilotica–acacia caffra–cymbopogon pospischilii shrubland, the acacia nilotica–acacia caffra–tragus berteronianus woodland and the acacia nilotica–acacia caffra–elionurus muticus woodland. 3. olea europea subsp. africana–rhus leptodictya woodland: this woodland community covers approximately 23% (52 ha) of the donga home range, with little rock cover and erosion. there are large trees throughout the area, with not much forb cover. rock cover ranged from 1% to 70%, with an average of 9%, and erosion was estimated at 22% (slight). altitude ranged from 1005 m to 1093 m, with the average slope being 20°. species belonging to species group k were diagnostic for this community (table 1, see online supplement). the woody vegetation is dominated by the smallto medium-sized evergreen tree olea europea subsp. africana, the deciduous, round-crowned, small to large tree celtis africana (species group k), the tree a. caffra, the medium-sized tree r. leptodictya and the largeto medium-sized tree berchemia zeyheri (species group m). the smallto medium-sized trees pappea capensis and ziziphus mucronata (species group m), and the spiny, multi-stemmed shrub r. pyroides (species group l) are prominent. the grass layer is dominated by the tufted perennial grass, p. maximum, with the perennial herb gerbera jamesonii (species group l) locally prominent. the average number of plant species was 18 per 400 m2 in this community. the tree and shrub layers covered between 5% to 95% and 1% to 12% respectively, compared to the 1% to 42% and 1% to 7% cover of the grass and forbs layers. the tree a. caffra had the highest density, with 236 individuals per ha, followed by the shrub e. crispa with 186 individuals per ha and the tree o. europaea subsp. africana with 136 individuals per ha. two sub-communities were distinguished, namely the olea europea subsp. africana–rhus leptodictya–carissa bispinosa woodland and the olea europea subsp. africana–celtis africana–mimusops zeyheri woodland. picnic home range the picnic home range could typically be classified as an acacia karroo–dichrostachys cinerea shrubland. in the same way that a detailed vegetative analysis was undertaken for the donga home range, the picnic home range was also analysed. noteworthy species occurring within the picnic home range include the smallto medium-sized deciduous tree a. caffra, the medium-sized tree r. leptodictya, the shrub solanum panduriforme, the erect shrub l. javanica and the tufted, perennial grasses p. maximum and heteropogon contortus. the picnic home range comprises eight plant communities (shown in figure 4), which can be grouped into the following four major community types (table 2, see online supplement): 1. acacia caffra–setaria sphacelata var. sphacelata woodland 2. acacia karroo–dichrostachys cinerea shrubland 3. heteropogon contortus–sclerocarya birrea woodland 4. hyperthelia dissoluta grassland figure 4: vegetation map of the picnic home range depicting the main plant communities description of the picnic home range plant communities the picnic home range is characterised by a series of dry streambeds cutting through it. there are two small, man-made dams, previously used for livestock and agricultural purposes. the vegetation structure is mostly open shrubland and woodland, with a herbaceous layer generally depleted due to overgrazing with cattle and irregular agricultural practices that took place prior to the establishment of the ldnr. the grass layer is represented by p. maximum, which occurs throughout the home range, while h. contortus is prominent. forbs are locally abundant throughout the home range, with several annual and geophytic species recorded seasonally.1. acacia caffra–setaria sphacelata var. sphacelata woodland: this woodland community covers approximately 31% (42 ha) of the picnic home range, with rock cover and erosion being relatively low. rock cover ranges from 1% to 20% with an average of 8%, and erosion was estimated at 15% (slight). altitude ranged from 1032 m to 1064 m, with the average slope being 12°. species belonging to species group a are diagnostic for this community (table 2, see online supplement). the woody vegetation is dominated by the smallto medium-sized deciduous tree a. caffra (species group g), the evergreen shrub a. karroo and the deciduous shrub d. cinerea (species group n). the small, erect, deciduous tree d. rotundifolia, the spiny, multi-stemmed shrub r. pyroides, the shrub e. crispa (species group g), the erect shrub l. javanica (species group n) and the multi-stemmed shrub grewia flava (species group f) are prominent. the tufted, perennial grasses p. maximum and h. contortus (species group n) dominate the grass layer. the average number of plant species recorded for this community was 26 per 400 m2. the tree layer had a 13% – 60% cover, the shrub layer 5% – 60%, the grass layer 50% – 75%, and the forb layer had a 1% – 3% cover. the shrub a. karroo had the highest density of 806 individuals per ha, followed by the shrub gymnosporia buxifolia with 750 individuals per ha and the tree a. caffra with 625 individuals per ha. two sub-communities were distinguished, namely the acacia caffra–setaria sphacelata var. sphacelata–ziziphus mucronata woodland and the acacia caffra–setaria sphacelata var. sphacelata–dombeya rotundifolia woodland. 2. acacia karroo–dichrostachys cinerea shrubland: this shrubland community occurs in two stands, one forming the eastern boundary and the other, smaller section located in the central part of the home range. this shrubland covers approximately 22% (31 ha) of the picnic home range (see figure 4). rock cover ranged from 1% to 5% with an average of 3%, and erosion was estimated at 38% (slight to moderate). altitude ranged from 1031 m to 1052 m, with an average slope of 8°. species belonging to species group d are diagnostic for this community (table 2, see online supplement). the woody vegetation is dominated by the evergreen shrub a. karroo and the deciduous shrub d. cinerea (species group n). the medium-sized tree r. leptodictya and the shrub s. panduriforme (species group g) are prominent. the grass layer is dominated by the tufted, perennial grasses aristida stipitata and eragrostis lehmanniana (species group g), with the tufted, perennial grasses h. contortus and p. maximum (species group n) being locally prominent. the average number of plant species was 20 per 400 m2, while the tree layer covers between 10% and 80% and the shrub layer 10% – 80% of the area. as expected, the grass layer has a relatively low cover ranging between 5% and 40%, while the forb layer has a 1% cover. the shrub a. karroo had the highest density of 558 individuals per ha, followed by the shrub d. cinerea with 483 individuals per ha and the shrub g. buxifolia with 408 individuals per ha. 3. heteropogon contortus–sclerocarya birrea woodland: this woodland community covers 46% (63 ha) of the picnic home range and has relatively high rock cover and soil erosion. rock cover ranged from 2% to 50%, with an average of 22%, and erosion was estimated at 21% (slight to moderate). altitude ranged from 1045 m to 1097 m, with the average slope being 12°. species belonging to species group h are diagnostic for this community (table 2, see online supplement). the woody vegetation is dominated by the evergreen shrub a. karroo and the deciduous shrub d. cinerea (species group n), with the erect shrub l. javanica (species group n) being locally prominent. the tufted, perennial grass h. contortus (species group n) dominates the herbaceous layer, with p. maximum (species group n) prominent locally. the average number of plant species was 24 per 400 m2 in this sub-community. the tree layer had 28% – 36% cover, the shrub layer 11% – 26%, the grass layer 39% – 58% and the forb layer 1%. the shrub d. cinerea had the highest density of 1133 individuals per ha, followed by the tree combretum apiculatum with 1075 individuals per ha and the shrub a. karroo with 783 individuals per ha. four sub-communities were distinguished for this community, namely the heteropogon contortus–sclerocarya birrea–urochloa oligotricha shrubland, the heteropogon contortus–sclerocarya birrea–ledebouria apertiflora shrubland, the heteropogon contortus–sclerocarya birrea–cenchrus ciliaris shrubland and the heteropogon contortus–sclerocarya birrea–faurea saligna woodland. 4. hyperthelia dissoluta grassland: this small grassland community covers 1% (2 ha) of the picnic home range (see figure 4). there were no surface rocks present and erosion was estimated at 45% (moderate). altitude was 1052 m, with the average slope being 2°. this community is characterised by the absence of diagnostic species and is distinguished by presence of species from species group m and the absence of species from species group l (table 2, see online supplement). the small, round-crowned tree ozoroa sphaerocarpa (species group h), the evergreen shrub a. karroo (species group n) and the tufted, perennial grass p. maximum (species group n) are prominent. the grass layer is dominated by the tufted, perennial grass hyperthelia dissoluta (species group m). the average number of plant species recorded in this community was 4 per 400 m2. the shrub layer had a 5% cover, while the grass layer covered 90% of the area. the shrub a. karroo had the highest density of 100 individuals per ha, followed by the shrub o. sphaerocarpa, with 50 individuals per ha. ordination for both the donga home range (see figure 5a) and the picnic home range (see figure 5b) there was an overall positive association between rock cover and altitude (with the exception of community four in the picnic troop), with higher lying (high altitude) areas being rockier than lower lying (low altitude) areas. erosion was more evident in the lower lying areas, varying at intermediate altitudes, with the exception of picnic community four, which occurred in a dry riverbed forming the confluence of two non-perennial streams. erosion in the picnic home range can largely be attributed to historical overgrazing by livestock and mismanagement of the natural vegetation. for the donga home range there was an inverse relationship between tree cover and altitude, resulting in higher tree cover in lower lying areas than in higher lying areas (see figure 6a). for the picnic home range, tree cover was highest at intermediate altitudes, with lower tree cover recorded in highand low-lying areas (see figure 6b). in the donga home range, the shrub layer was predominantly represented in community two. community two occurred at an intermediate altitude between communities one and three (see figure 6a). the recorded canopy cover for the shrub layer in donga community two was high in both the high-lying and low-lying areas, despite high tree-cover values and high exposure to wind, respectively. shrub cover in the picnic home range was highest at lower altitudes (see figure 6b). figure 5: rock cover and erosion in relation to altitude for (a) donga and (b) picnic home range communities; communities are arranged according to a gradient of increasing altitude figure 6: tree, shrub, forb and grass cover in relation to altitude for (a) donga and (b) picnic home range communities; communities are arranged according to a gradient of increasing altitude donga home range the flora in the donga home range is represented by 43 families, constituting 94 genera with 127 species. the dominant plant family is poaceae, constituting 22.8% (29 species) of the total flora, followed by fabaceae, with 9.4% (12 species). subsidiary families include the anacardiaceae, with seven species representing 5.5% of the total flora, combretaceae, with six species representing 4.7%, and asparagaceae and tiliaceae, both with five species each representing 3.9%. combined, these families represent 50.4% of the flora in the donga home range, while the remaining 36 families constitute 49.6%, comprising 63 different species (see figure 7). picnic home range the flora of the picnic home range is represented by 34 families, constituting 72 genera with 94 species. the dominant plant families are poaceae, constituting 26.6% of the total flora, followed by the fabaceae (6.4%) and anacardiaceae (6.4%). the poaceae comprise 25 species, with the fabaceae and anacardiaceae comprising six species each. other families include combretaceae, with five species representing 5.3% of the total flora, and asparagaceae and asteraceae, both with four species each representing 4.3%. combined, these families represent 53.2% of the flora in the picnic home range, while the remaining 28 families constitute 46.8%, comprising 44 different plant species (see figure 8). differences between the donga and picnic home range plant family representation were not significant (t46 = 0.037, p = 0.97). figure 7 & 8: 7: dominant plant families in the donga home range; percentage contributions to total flora for the area are depicted 8: dominant plant families in the picnic home range; percentage contributions to total flora for the area are depicted resource items the phenology of vervet resource items on resource tree species for the study period is shown in figure 9.vervet foods in the ldnr fell into two main seasonality categories. some showed year-round availability, such as euclea undulata, mimusops zeyheri and z. mucronata, which are keystone or mainstay species, while others are highly seasonal. these latter species are available only at certain times of the year, such as the various acacia sp. pods, b. zeyheri fruits, c africana fruits, o. europaea subsp. africana fruits, lannea discolor fruits and sclerocarya birrea fruits (see figure 9). figure 9: leafing, fruiting and flowering phenology of vervet resource trees for the two vervet home ranges resource availability although vervets have eclectic diets and generally eat whatever is available to them (lee & hauser 1998), predicting what they will consume at different times of the year is difficult because of the considerable seasonality of plant productivity in savannas, driven by variable rainfall and temperature (solbrig 1996). some trees require specific environmental conditions to synchronise before they produce fruit or resources (chapman et al. 2005), resulting in the vervets not being able to achieve complete dietary stability and their diets thus varying based on current availability. the main forage species showed heterogeneity across months in their contribution to the vervets’s diets. as a result, while the vervets consumed different plant specie resource items in different months, there was a constant supply of resource items throughout the year on resource tree species, especially leaves and fruit or pods (figure 10). fruits, however, showed a more complex pattern of availability across species, but fruits of some sort were also available and consumed in every month.comparisons of the available tree resources used by the two troops over the study period showed that there was a significant difference between the two home ranges for leaves (u = 27, p = 0.01), with trees in the donga home range bearing more leaves than those in the picnic home range. there were no significant differences for fruits or pods (u = 46, p = 0.13) and flowers (u = 65, p = 0.69). discussion (back to top) in terms of plant species composition, the donga home range shows similarity to the faurea saligna–setaria perennis, acacia caffra–setaria sphacelata and combretum apiculatum–heteropogon contortus tree savannas described by theron (1973). there is also a similarity to the acacia caffra–setaria sphacelata community described for the two newly proclaimed areas (rietfontein and parys) by götze et al. (1998). the picnic home range has broad floristic similarity to the acacia caffra–combretum apiculatum–themeda triandra and combretum apiculatum–heteropogon contortus tree savannas described by theron (1973). there also are similarities to the combretum apiculatum–themeda triandra community described by götze et al. (1998). dominant picnic home range tree species found in the combretum apiculatum–themeda triandra community include a. caffra, a. karroo and d. cinerea. the vegetation of the two home ranges has general similarities and could be combined into a single, large acacia caffra–dichrostachys cinerea–panicum maximum shrubland community. prominent species include r. leptodictya, d. rotundifolia, b. zeyheri and eragrostis chloromelas. ordination results for the home ranges show that tree and grass cover are negatively correlated, with increasing tree cover leading to decreased grass cover. traditionally it was thought that trees in temperate savannas reduced understory plant productivity through competition for light, water and nutrients (mcmurtrie & wolf 1983; walker & noy-meir 1982). recent studies, however, suggest that the relationship between trees and understory plants is more intricate, and that isolated trees occurring in communities with low tree densities may in fact improve understory productivity (belsky et al. 1993; knoop & walker 1985; stuart-hill, tainton & barnard 1987; weltzin & coughenour 1990). thus the relationship between trees and understory plants is dynamic and various site-specific factors can influence interactions among them. from the ordination, donga community one has noticeably lower tree cover values than communities two and three, which are located at higher altitudes. this most probably could be explained by the lower temperatures at higher altitudes, while the effect of soil type and depth cannot be excluded. in the picnic home range, community three occurs on footand midslopes and is signified by large s. birrea specimens. these footand midslopes provide a highly suitable habitat for large marula trees. for both home ranges, the higher lying areas had higher grass cover than the lower lying areas, with the exception of community three in the picnic home range. as mentioned above, community three had large s. birrea trees, providing too much shade for the grass layer to establish a dense sward; however, the shade-loving grass p. maximum occurs in dense patches in some localities. the forb layer is not well developed in either of the two home ranges. herbaceous plants that do occur are restricted to the shaded and sheltered areas under trees and shrubs, where soil moisture conditions are favourable. forbs flourished after short periods (4−5 days) of abundant rainfall, but their tenure was transient, as the soils dried out rapidly when sunny conditions returned. the results indicate higher numbers of large trees in the donga home range than in the picnic home range. historically, farmers removed many large trees from the picnic area for crop planting. it is speculated that the number of mature, seed-producing trees found in the donga home range are a reason for the large quantity of young trees found in the area, because of the sheltered environment they provide for seedlings, and the seeds they produce. however, in most instances these saplings did not survive to become adults due to competition for light and resources, particularly in the densely shaded areas that comprise extensive portions of the donga home range. the picnic area had more medium-sized trees than the donga area. this could be attributed to previous human interference that resulted in large, homogeneous stands of trees that were similar in size and age, including a. karroo and terminalia brachystemma. the invasive, weedy annual herbs bidens pilosa and tagetes minuta occurred in both home ranges, but densities in the picnic area exceeded those in the donga area. floristic analyses showed that the donga and picnic home ranges have 30 plant families in common. the donga home range has 13 families not represented in the picnic home range, with the picnic home range containing four families not represented in the donga home range. overall, the two home ranges show no significant differences when comparing results for plant family representation. the largest plant family represented in both home ranges is the poaceae, which is the most dominant and economically important family of flowering plants worldwide (walter 1979). the effects of seasonality on this family suggest that many species provide only temporary food supplies, especially during the dry period when grasses die back and store resources underground until favourable conditions return (gibbs russel et al. 1990). fabaceae is the second largest plant family represented in the two home ranges. this family shows high nitrogen metabolism capabilities, often with root nodules containing nitrogen-fixing bacteria. this suggests resources with high protein and high energy content, which are sought after by foraging animals, including vervets (grobbelaar & rosch 1981). the plant families anacardiaceae (the mango family), which contain mostly trees and shrubs, and asteraceae (the daisy family), which occur mostly as herbs and sometimes as shrubs, are well represented in the picnic home range. from a structural perspective, both home ranges had similar tree cover values, with the donga area containing more trees in the > 3 m class than the picnic area, which had more trees in the > 1 m – 3 m class. in the donga home range, tree cover was 46.7%, shrub 16.4%, forb 3.5% and grass 41.2%. picnic area tree cover was 31%, shrub 25.7%, forb 1% and grass 51%. overall vegetation cover in the donga home range was 75.6%, compared to 70.6% in the picnic home range. the donga home range was more heterogeneous compared to the picnic area and provided a larger number of plant species for the vervets to feed on. the donga troop exploited 36 different woody plant species, six of which constituted 50% of the total used. the picnic troop used 21 different species, of which only two species were responsible for 50% of the total usage. the loskop vervet troops preferred riparian areas with large trees, venturing into less-preferred areas only when there was sufficient incentive to do so, such as the availability of various fruits. highly used areas had relatively dense canopy cover, with trees that were sufficiently high to provide protection, resources and sleeping sites for the troops. for the donga troop, community three, the acacia nilotica–acacia caffra woodland provided all of their requirements and the troop spent the majority of their time in this community during both the wet and the dry season (barrett 2009). community three contained the bulk of resource trees that the donga troop utilised. for the picnic troop, community one, the acacia caffra–setaria sphacelata var. sphacelata woodland, and community two, the acacia karroo–dichrostachys cinerea shrubland, were used most. both these communities were adjacent to a non-perennial stream that had two man-made farm dams built in it. trees growing along the edges of the stream provided food, safety and sleeping sites for the troop and they spent most of their time in these trees. community two also contained a picnic site that was only frequented during school holidays or weekends, when visitor numbers increased. the troop generally spent less time in the area and preferred the natural food resources of communities one and two. figure 10: monthly proportions of available resource items consumed (n = 36) on all trees sampled (n = 113) for the main vervet resource species (n = 25) by home range, and combined conclusion (back to top) it is important that the different plant communities of the home range of animals are studied when conducting research on behavioural patterns. the braun-blanquet approach to classifying and identifying plant communities in the field provided an accurate and appropriate description of the vegetation for the purposes of this study. the results showed variations in the vegetation occurring in the two home ranges, with topographical differences and the effects of environmental factors influencing plant species availability. the acacia nilotica–acacia caffra woodland provided most of the food sources for the donga troop, while the acacia caffra–setaria sphacelata var. sphacelata woodland and the acacia karroo–dichrostachys cinerea shrubland were utilised most for food by the picnic group, suggesting that acacia woodlands play an important role in providing habitat for vervets at loskop dam nature reserve. similar communities have been described in other parts of the reserve (götze et al. 1998; theron 1973) and could be studied as possible habitats for vervet monkeys. overall resource plant availability and utilisation suggests that there is a large base of resources for both vervet troops to use in their respective home ranges. the effects of seasonality, with accompanying shifts in spatial and temporal resource abundance, could affect troop ranging patterns, particularly during the dry season when resources are less available. the phenology of resources used by the two troops showed that both the donga and picnic home ranges had sufficient plant resources to meet the minimum habitat and energy requirements of vervet monkeys. there are sufficient food species in the home ranges to sustain viable troops, even during the drier periods when resources are less abundant, hence vervet presence in the area. the larger tree species provide protection from predators while the vervets feed and sleep. both areas have access to drinking water. fire in the two troops’s home ranges will have less of an impact on the donga troop, as their resource trees occur mostly in or adjacent to the donga areas in their home range. these areas are relatively shady and there is not much fuel to sustain a fire under the trees. in addition, the large resource trees are sufficiently high to prevent excessive resource loss in the event of a fire going through the area. however, the grassy areas between the dongas will be affected by fire, although the donga troop is not dependent on the resources found in these areas. the picnic troop is more prone to resource loss from fire due to the homogenous nature of their home range and the general structure of the area. a large proportion of resource trees occurs interspersed throughout the area, with high fuel loads adjacent to and under them, making them susceptible to fire damage. a fire during the dry season could have a severe impact on vervet resources, and this would be exacerbated during a drought period. in the event of a fire occurring during such conditions, the resources that the vervets depend on might be insufficient to sustain the troop under natural conditions. acknowledgements (back to top) we thank the staff and management of loskop dam nature reserve for allowing and encouraging the research. this research received financial support from the national research foundation. editor: llewellyn c. foxcroft references (back to top) barbour, m.g., burk, j.h. & pitts, w.d., 1987, terrestrial plant ecology, 2nd edn., benjamin cummings, massachusetts. barrett, a.s., 2009, ‘spatial and temporal patterns in resource dispersion and the structure of range use and co-existence in a social omnivore chlorocebus aethiops’, phd dissertation, department of environmental sciences, university of south africa. barrett, a.s., brown, l.r., barrett, l. & henzi, s.p., 2006, ‘phytosociology and plant community utilisation by vervet monkeys of the blydeberg conservancy, limpopo province’, koedoe 49(1), 49−68. barrett, l., 2000, baboons: survivors of the african continent, london, toucan books. belsky, a.j., mwonga, s.m., amundson, r.g., duxbury, j.m. & ali, a.r., 1993, ‘comparative effects of isolated trees on their undercanopy environments in highand low-rainfall savannas’, journal of applied ecology 30, 143−155. bredenkamp, g.j. & bezuidenhout, h., 1995, ‘a proposed 6variation in feeding and ranging behavior in primates’, in t.h. clutton-brock (ed.), primate ecology, pp. 539–556, academic press, london. emery, a.j., lotter, m. & williamson, s.d., 2004, determining the conservation value of land in mpumalanga, mpumalanga parks board, nelspruit. fensham, r.j. & bowman, d.m.j.s., 1992, ‘stand structure and the influence of overwood on regeneration in tropical eucalypt forest on melville island’, australian journal of botany 40, 335−352. ferrar, t.a. & lotter, m.c., 2007, mpumalanga biodiversity conservation plan handbook, mpumalanga parks board, nelspruit. germishuizen, g. & meyer, n.l. (eds.), 2003, ‘plants of southern africa: an annotated checklist’, strelitzia 14, national botanical institute, pretoria. gibbs russell, g.e., watson, l., koekemoer, m., smook, l., barker, n.p., anderson, h.m. & dallwitz, m.j., 1990, ‘grasses of southern africa’, memoirs of the botanical survey of south africa, no. 58, botanical research institute, south africa. götze, a.r., meyer, s., cilliers, s.s. & kellner, k., 1998, ‘the classification of plant communities and range condition assessment of two new parts of loskop dam nature reserve, mpumalanga’, honours study, school of environmental sciences and development, potchefstroom university for christian higher education. grobbelaar, n. & rosch, m.w., 1981, ‘biological nitrogen fixation in a northern transvaal savanna’, journal of south african botany 47, 493−506. grobler, c.h., bredenkamp, g.j. & brown, l.r., 2006, ‘primary grassland communities of urban open spaces in gauteng, south africa’, south african journal of botany 72, 367−377. hennekens, s.m., 1996, megatab: a visual editor for phytosociological tables, giesen, ulft. hennekens s.m., 1996, turboveg: software package for input, processing and presentation of phytosociological data, version 1.9, lancaster university, lancaster.    hill, m.o., 1979, twinspan: a fortran program for arranging multivariate data in an ordered two-way table by classification of individuals and attributes, cornell university, new york. hoffmann, w.a. & franco, a.c., 2003, ‘comparative growth analysis of tropical savanna and forest trees using phylogenetically-independent contrasts’, journal of ecology 91, 475−484. hoffmann, w.a., franco, a.c., moreira, m.z. & haridasan, m., 2005, ‘specific leaf area explains differences in leaf traits between congeneric savanna and forest trees’, functional ecology 19, 932−940. isbell, l.a., pruetz, j.d. & young, t.p., 1998, ‘movements of vervets (cercopithecus aethiops) and patas monkeys (erythrocebus patas) as estimators of food resource size, density and distribution’, behavioral ecology and sociobiology 42, 123−133. kent, m. & coker, p., 1997, vegetation description and analysis – a practical approach, john wiley & sons, new york. knoop, w.t. & walker, b.h., 1985, ‘interactions of woody and herbaceous vegetation in a southern african savanna’, journal of ecology 73, 235−254. lee, p.c. & hauser, m.d., 1998, ‘long-term consequences of changes in territory quality on feeding and reproductive strategies of vervet monkeys’, journal of animal ecology 67, 347−358. leighton, m. & leighton, d.r., 1982, ‘the relationship of size and feeding aggregate to size of food patch: howler monkey alouatta palliate feeding in trichilia cipo trees on baro colorado island’, biotropica 14, 81−90. low, a.b. & rebelo, a.g. (eds.), 1996, vegetation of south africa, lesotho and swaziland, a companion to the vegetation map of south africa, lesotho and swaziland, department of environmental affairs and tourism, pretoria. matthee, j.f. & van schalkwyk, c.j., 1984, a primer on soil conservation, bulletin no. 399, department of agriculture, government printer, pretoria. mcmurtrie, r. & wolf, l., 1983, ‘a model of competition between trees and grass for radiation, water and nutrients’, annals of botany 52, 449−458. mucina, l. & rutherford, m.c. (eds.), 2006, the vegetation of south africa, lesotho and swaziland, strelitzia, 19, south african national biodiversity institute, pretoria. mueller-dombois, d. & ellenberg, h., 1974, aims and methods of vegetation ecology, wiley & sons, new york. pruetz, j.l. & isbell, l.a., 2000, ‘correlations of food distribution and patch size with agonistic interactions in female vervets (cercopithecus aethiops pygerythrus senso lato) and patas monkeys (erythrocebus patas) living in simple habitats’, behavioral ecology and sociobiology 49, 38−47. raemakers, j., 1980, ‘causes of variation between months in the distance travelled daily by gibbons’, folia primatologica 34, 46−60. sarmiento, g. & monasterio, m., 1983, ‘life-forms and phenology’ in f. brouliere (ed.), ecosystems of the world, tropical savannas 13, pp. 79−108, elsevier, amsterdam. singh, k.p., & kushwaha, c.p., 2006, ‘diversity of flowering and fruiting phenology of trees in a tropical deciduous forest in india’, annals of botany 97, 265–276. smit, g.n., 1989a, ‘quantitative description of woody plant communities: part i. an approach’, journal of the grassland society of southern africa 6, 186−191. smit, g.n., 1989b, ‘quantitative description of woody plant communities: part ii. computerized calculation procedures’, journal of the grassland society of southern africa 6, 192−194. smit, g.n., 1996, ‘becvol: biomass estimates from canopy volume (version 2) – users guide’, university of the free state, bloemfontein. solbrig, o.t., 1996, ‘the diversity of the savanna ecosystem’, in o.t. solbrig, e. medina & j.f. silva (eds.), biodiversity and savanna ecosystem processes: a global perspective, ecological studies, vol. 21, pp. 1–27, springer verlag, berlin. stuart-hill, g.c., tainton, n.m. & barnard, h.j., 1987, ‘the influence of an acacia karoo tree on grass production in its vicinity’, journal of the grassland society of south africa 4, 83−88. terborgh, j., 1983, five new world primates: a study in comparative ecology, princeton university press, princeton. theron, g.k., 1973, ’n ekologiese studie van die plantegroei van die loskopdam natuurreservaat [an ecological study of the vegetation of the loskop dam nature reserve]’, phd dissertation, department of general botany, university of pretoria. van schaik, c.p., terborgh, j.w. & wright, s.j., 1993, ‘the phenology of tropical forests: adaptive significance and consequences for primary consumers’, annual review of ecology and systematics 24, 353−377. walker, b.h. & noy-meir, i., 1982, ‘aspects of the stability and resilience of savanna ecosystem’, in b.j. huntley & b.h. walker (eds.), ecology of tropical savannas, pp. 556–590, springer verlag, new york. walter, h., 1979, vegetation of the earth, springer, new york. weltzin, j.r. & coughenour, m.b., 1990, ‘savanna tree influence on understory vegetation and soil nutrients in northwestern kenya’, journal of vegetation science 1, 325−332. werger, m.j.a., 1974, ‘on concepts and techniques applied in the zurich-montpellier method of vegetation survey’, bothalia 11, 309−323. westhoff, v. & van der maarel, e., 1978, ‘the braun-blanquet approach’, in r.h. whittaker (ed.), classification of plant communities, pp. 289−399, dr w junk, the hague. article information authors: kelly scheepers1 louise swemmer2 wessel j. vermeulen3 affiliations: 1csiro ecosystem sciences, winnellie, australia 2savanna and arid parks, south african national parks, south africa 3scientific services (knysna), south african national parks, south africa correspondence to: kelly scheepers email: kelly.scheepers@csiro.au postal address: csiro ecosystem sciences, pmb 44 winnellie, nt 0822, australia dates: received: 19 apr. 2010 accepted: 08 mar. 2011 published: 13 may 2011 how to cite this article: scheepers, k., swemmer, l. & vermeulen, w.j., 2011, ‘applying adaptive management in resource use in south african national parks: a case study approach’, koedoe 53(2), art. #999, 14 pages. doi:10.4102/koedoe.v53i2.999 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) applying adaptive management in resource use in south african national parks: a case study approach in this essay... open access • abstract • introduction    • setting the policy scene for south africa   • the sanparks resource use policy as a framework for case study comparison • case study 1: seven-weeks fern (rumohra adiantiformis) harvesting in the garden route national park    • introduction    • socio-economics aspects    • ecological studies       • community association and delineation of the harvest area       • population structure       • plant demography and reproductive phenology       • impact of harvesting    • adaptive management and refinement of harvest prescriptions    • lessons learnt from case study • case study 2: pepper-bark tree (warburgia salutaris) harvesting in the kruger national park    • introduction    • socio-economic aspects    • ecological studies    • adaptive management       • ecological tpcs       • discussion of existing ecological tpcs       • managing the demand    • lessons learnt from the case study • case study 3: sour fig (carpobrotus acinaciformis) harvesting in agulhas national park    • introduction    • socio-economic aspects    • ecological studies    • application of adaptive management to monitor sour figs    • integration of local and scientific knowledge    • flexible institutions    • lessons learnt from case study • conclusions • references abstract (back to top) south african national parks (sanparks) has a history of formal and informal natural resource use that is characterised by polarised views on national conservation interests and benefits to communities. current efforts aim to determine the sustainability of existing resource use in parks and to formalise these activities through the development of resource use protocols. the resource use policy of sanparks outlines principles for sustainable resource use, including greater involvement of local communities in management of protected areas and an adaptive management approach to determining sustainable use levels. this paper examines three case studies on plant use in national parks with regard to the development of criteria and indicators for monitoring resource use, and the role of thresholds of potential concern in measuring effectiveness of managing for sustainable use levels. opportunities and challenges for resource use management are identified. findings show that platforms for discussion and knowledge sharing, including research committees and community associations, are critical to building relationships, trust and a shared vision of sustainable resource use between stakeholders. however, additional capacity building is needed to enable local community structures to manage internal social conflicts and jealousy, and to participate fully in monitoring efforts. long-term monitoring is essential for developing flexible harvest prescriptions for plant use, but this is a time-consuming and resource-intensive exercise. flexible management strategies are difficult to implement and sometimes command-and-control measures are necessary to protect rare or endangered species. a holistic approach that considers resource use in national parks as a complement to broader community development initiatives offers a way forward. conservation implications: there is no blueprint for the development of sustainable resource use systems and resource use is often addressed according to multiple approaches in national parks. however, the sanparks resource use policy provides a necessary set of guiding principles for resource use management across the national park system that allows for monitoring progress. introduction (back to top) most protected areas are established with the primary goal to conserve biodiversity; however, the other roles protected areas play in people’s livelihoods are increasingly recognised. these include the provision of ecosystem services and support of socio-economic development opportunities (scherl et al. 2004). in south africa, as elsewhere in the world, the establishment of many protected areas have been associated with colonial processes of displacement and dispossession, and protected area management has been subject to exclusionary policies (abbot & mace 1999; nagothu 2001). consequently, strong arguments are also made for the provision of benefits from conservation to historically disadvantaged groups (scherl et al. 2004; von maltitz & shackleton 2004; willis 2004). in response to the ‘fortress conservation’ model adopted widely across africa to protect remnants of pristine habitats and recover declining wildlife populations, more socially responsible models have emerged as alternatives in recent years (hoole & berkes 2010; scherl et al. 2004). these models are more inclusive of the values and aspirations of local peoples and encourage greater involvement by local communities in protected area management (scherl et al. 2004). various community-based conservation initiatives, such as wildlife management areas in zimbabwe and community conservancies in namibia, have aimed to diversify the conservation economy to include nature-based tourism and the commercialisation of natural resource use (hoole & berkes 2010; lindsey, roulet & romañach 2007). zimbabwe’s communal areas management programme for indigenous resources (campfire) is perhaps the most famous african example of how local communities can benefit from being granted commercial rights to use wildlife. however, more recently, major socio-economic factors related to the fast track of land reform in zimbabwe, such as high unemployment, rampant inflation, food insecurity and social unrest, have threatened the long-term sustainability of the wildlife industry (clover & eriksen 2009). in south africa, contractual national parks offer a potential win-win solution for conservation and communities through joint management of the land and its resources (reid & turner 2004). contractual parks provide a way to expand the national network of protected areas with the inclusion of land that cannot be bought and/or on which mining rights will not be released. the concept was first developed when the government could not expropriate powerful commercial land owners from land wanted for conservation but has more recently been applied in the context of land claims on national parks by communal land owners (reid et al. 2004). for communities, contractual parks represent a way to enforce their land ownership rights and gain access to conservation, tourism and natural resource use benefits through co-management arrangements with south african national parks (sanparks). however, most contractual parks are of little biodiversity conservation significance, as compared to the greater national park area of which they form part (reid et al. 2004). contractual parks, as a model for social redress, have also been criticised for reasserting state or elite control over resources, perpetuating the dominance of western management frameworks, contributing inadequately to capacity building, and exacerbating the skewed distribution of benefits in south africa (murphree 2000; reid & turner 2004). yet, as with most recent initiatives, these issues perhaps represent the to-be-expected ‘teething problems’ of a more participatory approach to management compared to the strict ‘fines and fences’ policies of the past (reid et al. 2004). in addition, the sanparks resource use policy (2008) allows a wide range of stakeholders to use park resources for everyday purposes. there are many current examples of both informal and formal resource use in national parks. certain of these practices are being allowed, and in some cases supported by sanparks, with the long-term view towards developing monitoring systems and establishing local governance arrangements for sustainable resource use that incorporate local and scientific knowledge. a number of flagship projects, which span varied stakeholder groups’ use of multiple resources across the national park network, serve as case studies of the implementation of the resource use policy. three such case studies are presented in this paper. this study examines the role of sustainable resource use as a strategy for providing ‘benefits beyond boundaries’ in south african national parks. a brief description of the current context and driving factors of resource use in national parks is provided. using the national resource use policy as a framework for comparing case studies on plant resource use across three national parks, we then examine the effectiveness of the sanparks management strategy for ensuring sustainable use and providing resource users with socio-economic benefits. setting the policy scene for south africa sanparks has a long history of natural resource use. however, whereas incidents of illegal poaching and resource use were largely isolated, opportunistic and for customary use purposes in the past, high demands for improved access to park resources, driven by an increasingly complex set of socio-political factors under the new political dispensation in south africa (after 1994), have recently shaped policy development and implementation. resource use can no longer be managed on a case-by-case basis because of the growing diversity of resource users, natural resources and resource use strategies that support human livelihoods. resource harvesting currently takes place in national parks as part of core business, nature-based tourism activities, research, and in the spirit of ‘benefits beyond boundaries’. the resource users comprise several stakeholder groups, including the public, staff, conservation partners, researchers, local entrepreneurs and small business owners, tourists and neighbouring communities. in particular, there has been strong emphasis on the development of smallto medium-scale enterprises for sustainable resource use, such as community nurseries and arts-and-crafts centres. this marks a shift in resource use strategies, moving from customary use towards greater reliance on the market economy. past experiences have shown that once resource use entitlements have been granted they are difficult to amend. sanparks has therefore also had to implement a management system that promotes learning by doing (cundill & fabricius 2009; stankey, clark & bormann 2005). strategic adaptive management entails a participatory process to define management challenges, strategise future directions, implement management actions and evaluate their effectiveness in meeting management objectives. however, because national-scale drivers such as social redress and poverty alleviation influence resource use in national parks, localised management solutions alone cannot bridge the divide between conservation policy and management for ecosystem integrity. resource use management requires a systems approach that recognises linkages between national parks and their surrounds and acknowledges a complex definition of sustainability with ecological and socio-economic components (fabricius 2004; hansen & defries 2007). despite the best efforts of national parks to conserve biodiversity and ecosystem integrity, threats such as unsustainable resource use persist because ecological functions and processes often occur over larger spatial scales (hansen & defries 2007). to ensure that ecosystems retain the ability to renew themselves, additional land is needed for the expansion of national parks given limited sites suitable for conservation, competing land development options and growing tensions over the national conservation budget. consequently, creative ways to supplement formal conservation efforts are required. this involves the implementation of policy reforms and marketbased incentives to encourage private and communal land owners to invest in conservation as a competitive land use option (bulte, van kooten & swanson 2003). bioregional plans, contractual national parks and stewardship arrangements are examples of such initiatives. however, as secure land ownership rights are central to these community-based conservation models, benefit sharing is tied to a narrow definition of the legitimate resource users as land title holders. ultimately, there is a need for a wider-reaching policy that addresses people’s everyday resource use needs and which can complement land management solutions. the sanparks resource use policy as a framework for case study comparison the sanparks resource use policy (sanparks 2008) deals exclusively with resource use rights rather than land rights. this is significantly different from the application of contractual park agreements. the policy first aims to regulate existing resource use in national parks but also allows for the identification of new opportunities to provide socio-economic benefits to people. legitimate resource users are defined as all stakeholders who influence or are influenced by management decisions pertaining to resource use in national parks. a complex systems view of sustainable resource use acknowledges that the direct, consumptive use of park resources alone cannot meet people’s demands. consequently, sanparks supports investments in collaborative partnerships and networks that link local resource users to wider socio-economic development initiatives. in particular, the current policy highlights opportunities for sustainable resource use associated with the eradication of invasive alien species in national parks through the expanded public works programme. sustainability also takes on a complex meaning, as the policy calls for an adaptive management approach that links monitoring to management, combines scientific and local knowledge, and allows for flexible rules governing resource use. adaptive management requires change indicators to monitor trends in resource use, where ecological sustainability is defined by a rate of use that is less than that of the natural rate of replenishment of the resource. however, rather than using fixed targets, sanparks measures sustainable resource use against thresholds of potential concern (tpcs). this approach allows for a certain amount of ecosystem change brought about by resource use, provided it remains within the natural limits of variation. consequently, tpcs create a feedback loop between monitoring and management, which serves as an early detection system for undesirable changes to ecosystems and prompts informed management interventions to promote cyclical learning (biggs & rogers 2003; parr, woinarski & pienaar 2009). there is also a socio-economic component to sustainable resource use, as the process of developing monitoring indicators and tpcs is resource intensive and time consuming. this process entails the active involvement and support of all stakeholders to ensure effective monitoring of sustainable resource use and adherence to the rules. the sanparks policy does not explicitly call for the co-management of resources; however, greater stakeholder participation in resource use management is encouraged, including the incorporation of local knowledge in monitoring and evaluation systems. the roles and responsibilities of stakeholders vary according to the local context and purpose of the resource use. for example, community projects that address historical disadvantage through local economic development and capacity building are distinguished from commercial big business and afforded special recognition and greater ‘hands-on’ support by sanparks. on the contrary, commercial developers, while expected to adhere to sustainable resource use practices, are also required to contribute financially and otherwise to the development and implementation of monitoring systems. flexible rules that are informed by the zonation of harvest and control areas, multiple indicators of change, tpcs for different harvest intensities and provisional harvesting permits also support sustainable resource use. the zonation of use and no-use areas within national parks allow for greater flexibility when setting tpcs. multiple overlapping sets of rules make it more difficult for opportunits and outsiders to abuse the system. provisional harvesting permits enable issues that arise from the management process to be dealt with in a timely manner and facilitate feedbacks between monitoring and management. the following case studies discuss the implementation of these principles for sustainable resource use. in particular, they focus on the use of an adaptive management approach to monitoring sustainable resource use, which incorporates scientific and local knowledge and informs flexible rules of management. case study 1: seven-weeks fern (rumohra adiantiformis) harvesting in the garden route national park (back to top) introduction the fronds (leaves) of the seven-weeks fern (rumohra adiantiformis [g. forst.] ching) are extensively used in flower arrangements, both locally and abroad. harvesting of r. adiantiformis fronds from natural forests in the southern cape started as far back as 1970 when private land owners were issued permits to harvest small quantities on their lands for the local flower market (milton & moll 1988). however, owing to rising demands and the high economic value of the resource, which outweigh the risk of prosecution and fines, illegal harvesting escalated (geldenhuys & van der merwe 1988). in 1982, approximately 4000 ha of indigenous state forest land was released for the commercial harvesting of r. adiantiformis fronds as a means to address the high demand and to counteract illegal harvesting. however, too little was known about the species’ ecology, dynamics and productivity to inform sustainable harvest practices. following an adaptive management approach, harvest prescriptions for the species were developed and refined as research results became available. socio-economics aspects the species is solely harvested for commercial purposes and the sale of r. adiantiformis fronds from state forest land is managed through an open tender process. given the commercial value of the product and growing demand for access, entrepreneurs with a business drive, financial capacity and a vested interest in the resource became the key stakeholders. state-funded research was initiated to advise managers of sustainable harvesting practices while fern harvesting contractors commissioned additional research and studies on the ecology and dynamics of the species (geldenhuys & van der merwe 1986; milton & moll 1987). the industry dictates minimum requirements for the commercial sale of the resources and only fully matured, hardened fronds without necroses or other blemishes are considered usable for the florist trade. frond size, form and colour also constitute important criteria for product quality (geldenhuys & van der merwe 1988). however, as these criteria were not always clearly defined, fern populations were initially harvested every five weeks with no restrictions on the number of fronds that could be harvested. as stricter harvest prescriptions were implemented, the harvesting of fronds from state forests was insufficient to meet industry demands for a maximum sustainable yield. the industry had to find alternatives to meet the demand, which resulted in the establishment of commercial nurseries in addition to fern stands that had been established under commercial pine plantations. the alternatives to harvesting wild populations of the resource ensured not only the economic sustainability of the industry but also the extension of socio-economic benefits through the creation of additional jobs (kok 1998). with harvesting from the wild becoming less attractive to large commercial players, a community fern harvesting project was initiated to supply commercial operators (vermeulen, du plessis & herd 2005). in this way, poor and unemployed individuals (especially women) living close to the forest (the target group) had become direct beneficiaries from fern harvesting. at its peak, the income from fern sales exceeded that of indigenous timber (vermeulen 2009), but, more importantly, a new industry had been developed with associated large socio-economic benefits. forest management had to adapt to the harvesting of a non-timber forest product during a time when commercial timber harvesting was the primary focus of resource management. similarly, the industry had to shift from the harvesting of wild populations to alternative resources to meet demand and maintain benefits. ecological studies the generic process of determining sustained yield for forest products and developing harvest prescriptions has been well documented (cunningham 2001; geldenhuys 2000; peters 1996). this process has also guided research and monitoring to inform harvest prescriptions for r. adiantiformis (figure 1). two key components of the process are: • delineating the potential resource area following studies on the habitat and distribution of the target species, and • studying the population dynamics, demography and reproductive phenology of the species. figure 1: flow diagram indicating the generic process for the development of harvest systems and management prescriptions for resource use (adapted from vermeulen, w.j., 2009, ‘the sustainable harvesting of non-timber forest products from natural forests in the southern cape, south africa: development of harvest systems and management prescriptions’, phd thesis, department of conservation ecology and entomology, stellenbosch university). the section that follows is a summary and review of research results as they became available and shows how these impacted on the formulation of management prescriptions through a process of adaptive management. community association and delineation of the harvest area to identify and accurately map the potential harvest area for a species, knowledge of its habitat and distribution and insight into its community association are required (figure 1). based on earlier forest type classification (von breitenbach 1968) and with consideration to local knowledge, forest areas with potential for commercial fern harvesting were identified and mapped. the species was found to have a patchy distribution and to occur in fairly dense clusters in wet and moist high forest, which made the accurate mapping of the potential harvest area difficult. a detailed phytosociological study of the forest vegetation in the southern cape, including a description of both tree and ground flora communities was concluded in 1993 (geldenhuys 1993). r. adiantiformis was found to be an important component of several ground flora communities and to occur in scarp, platform and mountain forests. of major significance was that r. adiantiformis is frequently associated with specific understorey communities in tall to high, moist regrowth forest, on a variety of sites in mountain, foothill, platform and coastal scrub vegetation. it represents an early regrowth phase (after disturbance by, for example, fire) and should eventually be suppressed by shrub understorey and tree regeneration (geldenhuys & van der merwe 1994). population structure knowledge of the population structure and demography of the target species is essential in the development of a system for sustainable use and in assessing potential yield (figure 1). this was also a focus point of research. the results showed that the population structure and dynamics of the species vary temporally and spatially (within and between populations) and are affected by the density of the tree and shrub layers (geldenhuys & van der merwe 1988; milton & moll 1987). plant density ranges between zero and 14 plants per square metre, with an average of four plants per square metre. the number of fronds per plant, which proved to be an important variable in the formulation of harvest prescriptions for the species, varies between two and three, depending on site and season (geldenhuys & van der merwe 1988, 1994; milton & moll 1987). this resulted in a harvest prescription that restricted harvesting to only 50% of the harvestable fronds per plant. frond size is another critical component for setting harvest prescriptions and is affected by factors such as habitat, stand density, season of bud production and rainfall (geldenhuys & van der merwe 1988, 1994; milton & moll 1987). an average frond length of 48 cm was recorded for floodplain populations, compared to 27 cm for ridgetop populations. together with market demands, this informed the decision to set the minimum frond length for harvesting at 25 cm. plant demography and reproductive phenology apart from population structure, information on frond development, rate of production and frond longevity was essential in refining harvest prescriptions, especially with regard to the harvest cycle. study of the reproductive phenology of r. adiantiformis allowed the optimum harvest season to be identified. the rate of frond production by r. adiantiformis was low compared to the rate of exploitation. long-term monitoring indicated an annual production of between one and three fronds per plant (milton & moll 1987) or two fronds per square metre, depending on stand density (geldenhuys & van der merwe 1994). it was also found that bud initiation began during july and august and peaked between september and november (geldenhuys & van der merwe 1988, 1994). considering the above, the period from january to march (i.e. before the onset of bud formation) could be regarded as the least sensitive to harvesting, and the opposite for the period between september and november. monitoring also showed that mature fronds increase in proportion to dominate from january to march, with a peak between december and january, and the lowest number in july (geldenhuys & van der merwe 1994). this finding was critical for assessing the potential yield and scheduling harvest operations. phenological events are not always predictable because they are synchronised with rainfall and temperature fluctuations. however, because a reliable supply of the resource is required for a viable socio-economic industry, a moratorium on harvesting during certain times of the year would not have been a suitable management option. the problem was partly addressed through a longer harvest cycle of 15 months, which would ensure that a particular area was not harvested consistently during the same time of year. impact of harvesting assessing the impact of harvesting on individual plants and the resource as a whole (cunningham 2001) is an essential part of refining harvest prescriptions through an adaptive management approach. with an industry dependent on the r. adiantiformis resource, research into the impact of frond harvesting was a priority. experimental harvesting at different intensities and frequencies was subsequently initiated (geldenhuys & van der merwe 1986, 1988; milton 1987a, 1987b, 1991; milton & moll 1987). monitoring results indicated a significant decline in mature frond size for partial and total defoliation treatments at a harvest rotation of less than 12 months. overharvesting also led to plant mortality and a reduction in harvestable fronds and impacted negatively on rhizome growth, the timing of budding and spore production. in addition, plant recovery after overharvesting was slow. milton (1987a) found no significant increase in the average length of fronds on defoliated plants (two cycles, nine-month interval) during a 22-month rest period. adaptive management and refinement of harvest prescriptions preliminary research results to inform harvest prescriptions for the species were presented at rumohra research committee meetings attended by researchers, managers and representatives from the industry in 1984 and 1985, followed by formal reports and publications (viz. geldenhuys & van der merwe 1986, 1988, 1994; milton 1987a, 1987b, 1991; milton & moll 1987, 1988). the interpretation of the findings and refinement of harvest prescriptions were complicated by spatiotemporal variation in the research, drought during parts of the research period (geldenhuys & van der merwe 1994) and the disturbance of some research sites due to uncontrolled fern picking, which demonstrated the importance of setting aside unharvested areas for control purposes. the sensitivity of the species to overutilisation, as reflected especially by the decline in frond size and the slow recovery rate, could be regarded as the most important trigger for the revision of the harvest prescriptions (vermeulen 2009). the long-term implications for the industry, if harvest rates were not aligned with the natural rate of production for the species, were therefore realised. through the adaptive management approach, and based on results of population dynamics and plant demography research and monitoring, the initial harvest cycle of 5 weeks was increased to 8 weeks, then to 16, 26 and 52 weeks, and is currently set at 15 months. total defoliation was prohibited and harvest intensity restricted to 50% of mature fronds per plant, with a minimum frond length of 25 cm. phytosociological studies indicated that r. adiantiformis is often associated with regrowth forest. the potential harvest area can therefore not be fixed for the long term but should be considered at a landscape level, taking into account the natural ecological processes and disturbance regimens. this further emphasises the relevance of an adaptive management approach to r. adiantiformis harvesting. the formulation of a clear definition for a usable frond, based on market demands as well as ecological considerations, proved to be an important management tool in controlling harvest operations, enforcing harvest prescriptions and minimising wastage. the need for continued monitoring to assess the impact of harvesting and to gather further information on fern stand development was stressed by geldenhuys (1994). to this end, and to refine harvest prescriptions further, various long-term monitoring projects were initiated in and outside fern picking areas (geldenhuys & van der merwe 1994; kok 1998). the results indicated that the current harvest prescriptions provide for the sustainable harvesting of the species. however, to ensure optimum use given spatiotemporal variation in the distribution of harvestable fern stands harvest areas should be redefined where required (vermeulen 2009). specific indicators of change are frond size, number of usable fronds and yield from picking areas. specific tpcs were not historically developed for the species but a consistent decline in fern frond size in picking areas triggered the extension of the harvest rotation. the gradual decline in harvest potential of the resource in picking areas, detected through continued monitoring (vermeulen 2009), was also of concern. the same trend, however, was recorded in control areas outside of picking areas, supporting the argument for a natural decline in fern densities in regrowth forest. tpcs for the harvesting of the species need to take such spatiotemporal variation into consideration. however, other drivers of change (e.g. climate) also need to be considered in future research. lessons learnt from case study key lessons were learnt from the development of a harvest system for r. adiantiformis. • the precautionary principle (cooney 2004) should be followed with the development of interim harvest prescriptions. given the negative impact of harvesting on fern populations in accordance with initial prescriptions, it was apparent that the precautionary principle was not applied effectively. this was exacerbated by the slow recovery rate following overharvesting. • knowledge gaps should be identified and research objectives clearly set. with the complexities of plant populations and the dynamic nature of forest ecosystems, it is difficult to predict long-term yield or the impact of resource use. the development and refinement of harvest prescriptions should be ongoing, which require continued monitoring at different levels. • the identification of indicators of change and the development of tpcs could support sustainable resource use. instead of fixed measures, tpcs should be developed and refined as the knowledge base on the resource and ecosystem functioning is broadened through research and long-term monitoring. • the costs of developing, implementing and monitoring sustainable harvest systems should not be underestimated. management agencies should ensure that the necessary resources are available when initiating resource use programmes. • where resources are harvested for commercial use, the developer should be prepared to contribute towards scientific studies on the target species to inform management prescriptions. in the case of r. adiantiformis, research was conducted in a complementary way, funded and initiated by the state and the commercial developers. • close cooperation and knowledge sharing between the different role players are essential. the r. adiantiformis research committee was indispensable as a platform for feedback and discussions between researchers, managers and industry. the needs of the different role players also had to be understood. resource managers had to appreciate the economic potential of the resource and the loss of potential business opportunities if access to the resource was restricted. similarly, businesses had to appreciate the importance of ecological and economic sustainability and the complexities and time span associated with the development of scientifically based harvest prescriptions. • applying the zonation principle whereby use and no-use areas are defined is essential for integrated resource management. not only does it allow for assessing the sustainability of the resource being harvested but also for the spatial separation of incompatible management practices to accommodate multiple forest management objectives. • the case study demonstrated the importance of the development of alternative resources where desirable, not only to meet demand and reduce pressure on resources from the wild, but also to optimise socio-economic benefits through the commercialisation of high-value products. case study 2: pepper-bark tree (warburgia salutaris) harvesting in the kruger national park (back to top) introduction it is estimated that approximately 27 million south africans rely on traditional medicine (mander 1998). considering that 34% of the population are between the ages of 15 and 65 years and have no fixed employment (botha, witkowski & shackleton 2004), the medicinal plant trade makes an important contribution towards human livelihoods. tissue from the pepper-bark tree (warburgia salutaris [bertol.f.] chiov.) is a popular traditional medicine that is used and traded throughout southern africa (botha, witkowski & shackleton 2004). the bark, stems, roots and leaves are used to treat numerous health complaints, including colds, respiratory complaints, fever, malaria, influenza, coughs, venereal diseases, abdominal pain, constipation, cancer, rheumatism, stomach ulcers and headaches (diederichs 2006). as such, the demand for w. salutaris products is high and most tree populations have been intensely impacted. the historical distribution of w. salutaris stretches from kenya and tanzania in east africa, down through mozambique and zimbabwe, to south africa and swaziland in the south. it is also found in madagascar. in south africa, the species was recorded widely in kwazulu-natal, mpumalanga and the limpopo province, but is thought to have been extirpated from much of this area as a result of high harvesting intensities and habitat destruction. w. salutaris is listed as both nationally and internationally endangered, according to the sanbi red list (sanbi 2009) and the iucn red list (iucn 2010), respectively. the kruger national park (knp) is host to what is believed to be the largest remaining wild populations of w. salutaris in the limpopo province. although the plants are located within the boundaries of the park, illegal harvesting of bark from w. salutaris trees occurs. this threatens the park’s mandate of biodiversity protection. on the contrary, denying communities access to this valuable resource contributes towards negative park–neighbour relations, which, in turn, threaten to undermine other park management objectives that pertain to access and benefit sharing. other potential threats to the plants include the possible negative effects of climate change, limited reproductive behaviour and genetic isolation of subpopulations. in response, sanparks has embarked on a multidisciplinary programme that aims to promote and facilitate the conservation of w. salutaris in the knp and the surrounding areas, in order to benefit both current and future generations (swemmer 2010). socio-economic aspects traditional health practitioners (thps), who operate locally as well as in urban areas, are the primary users of w. salutaris tissue. in the past, medicinal plant harvesting was subsistence based, and regulated by local customary laws and harvesting methods that promoted ecological sustainability. however, more recently these traditional practices have been threatened by outsiders using commercial approaches which adopt high-intensity, high-impact harvesting strategies which are less ecologically viable and sustainable in the long term (natural justice 2010). in the limpopo province, w. salutaris is in high demand by local thps and commercial herbalists or ’muti hunters’ (natural justice 2010). most thps interviewed in the province indicated that they had never seen a w. salutaris tree despite their use of its bark (anon., pers. comm.). this is mainly because the tree is so rare that thps are forced to purchase its bark either as whole pieces or as powder on the local markets, often at a high price (anon., pers. comm.). as a result of this scarcity, the market value of w. salutaris tissue has increased remarkably over the past decade. many of the local thps have not heard of using the leaves as opposed to the bark, although studies have shown that the chemical and healing properties of these two tree parts are similar. w. salutaris leaves are not readily available in the local markets (anon., pers. comm.). ecological studies as a result of the threatened status of w. salutaris, and an increase in illegal harvesting of the population within the knp, an active research, management and monitoring strategy was needed to promote the long-term survival of the species. a number of plant surveys were conducted over the past decade. the first survey was completed in 2000 and found that human impact had affected approximately 12% of the total number of stems recorded (botha, witkowski & shackleton 2004). very few specimens of the smaller size classes (< 1.5 m) were found (less than 10% of the population). in 2006, a second survey found that the situation had worsened, with 57% of the trees showing signs of human damage (figure 2). the survey again reported very few individuals (3%) in the smaller size classes, with most plants (90%) recorded as between 3 m and 7 m tall. almost two-thirds (59%) of the plants were multi-stemmed. figure 2: percentage of damaged and undamaged individuals for each height class of warburgia salutaris recorded in the kruger national park in 2006. adaptive management ecological tpcs these results inspired the development of the first suite of tpcs for w. salutaris (zambatis 2006). the aim of this set of tpcs was to provide managers and researchers with indicators and acceptable limits they could then use to monitor and detect changes in the population, especially in the event that these changes were human induced. tpcs 1 and 2 related to population dynamics, and tpcs 3 and 4 to harvest impact. tpc 1: no or very few (< 30%) adults recorded. (an adult is defined as a specimen with a primary stem diameter of at least 10 cm. stem diameter is measured at a point just above the basal swelling.) tpc 2: no or very few (< 10%) juvenile plants recorded in the population. (a juvenile is defined as a specimen that is less than 1.5 m tall.) tpc 3: the primary stems of at least 20% of all specimens have been ring barked and the entire plant is dead. tpc 4: one or more primary stems of more than 60% of specimens have been debarked and/or broken but are not dead, even though the individual may be re-sprouting from the main stem below the injury point or coppicing from ground level. the results of the 2006 survey showed that tpc 2 had been exceeded by about 7% (table 1). it was also noted that tpc 4 was close to being exceeded. towards the end of 2008, a number of illegal harvesting incidents took place including extensive ring barking of more than 13 stems on a single day (a. lumbe, pers. comm.). this highlighted the need for immediate management action. funding was obtained to implement an integrated conservation effort, which included employing a team of field rangers to patrol the area daily. table 1: the suite of thresholds of potential concern tested using data from surveys conducted in 2006 and 2009. in 2009, a third survey was conducted and the results showed a further 6% increase in the proportion of damaged plants. this exceeded the limits set by tpc 4. again, only a small percentage (about 5%) of individuals was taller than 1.5 m (table 1). in addition, there was a drop of approximately 13% in the proportion of adults recorded. discussion of existing ecological tpcs no or very few adults recorded (tpc 1): this tpc was designed to detect a change in the proportion of adult trees in the population. if the observed change was a reduction in proportions, as was seen between 2006 (56%) and 2009 (43%), it would suggest that the number of large w. salutaris plants is declining and that there may be cause for concern. however, since the tpc is based on proportions as opposed to total counts, it is not possible to ascertain whether there has been a drop in the total number of large plants or whether there has been a drop in the total proportion of large plants. the latter could be true if a large recruitment event took place, resulting in a large number of small plants, which, in turn, would reduce the proportion of large plants in the population as a whole, but not necessarily the numbers. since the population size is small, it may be more useful to use total population counts or density counts in permanent plots to pick up more subtle changes. these options are feasible temporary alternatives when the entire population of plants is reasonably small and contained as is seen in the w. salutaris population. it is important to keep the point at which the stem diameter is measured consistent to ensure repeatability. stems rather than individuals should be measured. no or very few juvenile plants (tpc 2): this tpc was designed to detect changes in recruitment patterns of w. salutaris. it is assumed that in healthy populations, the ‘reverse j’ population curve would be observed with large numbers of plants in the smaller size classes, decreasing for the bigger size classes. low proportions of juvenile plants would suggest little recruitment, which would be indicative of an unhealthy population. however, this application is not suitable for some savanna tree species that recruit sporadically during infrequent and opportunistic recruitment events. in this case, a bell-shaped distribution of size classes does not necessarily indicate a problem with the species. unfortunately, the natural population dynamics of w. salutaris are not well documented. for the w. salutaris population in the knp, low proportions of small plants have been recorded, namely 3% in 2006 and 5% in 2009. in kwazulu-natal, it was found that there was a very small seed set of w. salutaris. no seedlings were recorded and all plants had reproduced vegetatively (hilton-taylor et al. 1998). similar observations were made in the knp population. very few seeds were found and most plants were either coppicing or joined by coppice shoots to one another. this suggests that the plants are reproducing asexually from root suckers. a measure of small individuals is useful as an indication of recruitment but, as in tpc 1, it would be more useful to use total counts or plot densities as opposed to proportions of individuals at this stage. furthermore, using stem diameter as a measure of size classes rather than height is suggested for the modified version of this tpc. the primary stems of individuals have been ring barked and the entire plant is dead (tpc 3): although ring barking causes death of the crown above the site of damage of the effected stem, it may not be the case with a multi-stemmed plant with one or more undamaged stems. in the w. salutaris population in the knp, most plants (59%) are multi-stemmed (zambatis 2006) and coppice regularly. botha, witkowski and shackleton (2004) report an increased coppicing rate in individual plants that have been subjected to uncontrolled harvesting. this tpc could be modified to monitor numbers as opposed to proportions of dead stems. monitoring proportions will be better in the long term, but since the population is small, it may be possible and preferable to do total counts. one or more primary stems of most individuals have been debarked and/or broken but are not dead (tpc 4): this tpc was designed to use as an indicator of the proportion of damaged plants. the increasing proportion of damaged plants between 2000, 2006 and 2009 (12%, 55% and 61%, respectively) is a real management concern. however, once again, a measure of numbers or plot densities as opposed to proportion of damaged stems may be a more accurate measure of impact. a measure using percentage of stems as opposed to percentage of plants may also yield more accurate results. the acceptable limit of 60% may also be too high and may have to be reduced. managing the demand it is acknowledged that to conserve the w. salutaris population in the long term, an integrated approach involving biophysical and social research, outreach and awareness, and conservation and protection will be necessary. in 2004, the people and conservation department of the knp implemented an awareness education programme focused on rare and invasive species, funded by the development bank of south africa. a survey was conducted amongst local communities adjacent to the south-western boundary of the knp with the aim of encouraging youth development in conservation challenges. the results regarding people’s perceptions of the status of w. salutaris showed that 13% of the respondents did not know the tree and 67% of respondents have never seen one before. a further 34% of respondents did not know the tree was rare, but 24% said that it was not easy to find. only 16% of the respondents said it was rare because it was used for medicine. additional perceptions given by respondents for it’s rarity included the belief that w. salutaris is used to make paper, is being chopped down and that it needs more water than other trees (swemmer 2006). another issue addressed through the programme is the development of alternative sources to wild populations (swemmer 2010). this is being done through a number of workshops aimed at training local thps in the propagation of w. salutaris. these workshops are also used to market the use of w. salutaris leaves rather than bark in traditional medicine practices. the programme aims to promote access to w. salutaris trees in home gardens for medicinal use, as well as to enable the selling of surpluses to local and urban markets. well-established associations of traditional healers (e.g. the vukuzenzele nursery and medicinal garden in bushbuckridge and the makuya traditional healer association) could consider a more commercial approach to w. salutaris propagation. so far, workshop participants have been supportive of using w. salutaris leaves. they have also propagated more than 600 w. salutaris cuttings successfully. although the cuttings are being cared for by the propagation and training nursery, once established, they will be given over to the thps to plant at home. local resistance to using the artificially propagated plants was cited as a potential challenge, but so far this has not been the case. further challenges arise when defining which user groups should be targeted in awareness initiatives. only a small percentage of the thps who work in the communities neighbouring the knp can be directly involved in the workshops owing to resource limitations, and so selecting the participants in a transparent and fair way is challenging. furthermore, since it may be that illegal harvesting by outsiders supplies commercial markets in the large cities, targeting only local thps may not be effective in protecting wild populations of w. salutaris in the knp. identifying the demand and supply chain to commercial markets will play an important role in the identification of target groups for outreach initiatives. during the 2009 field trip, it was speculated that the harvesting methods originally used were different from those used to cause more recent damage, as older tree scars were made by striprather than ring barking. the former resulted in better bark recovery than the latter, as the main part of the stem remained untouched (swemmer 2010). lessons learnt from the case study key lessons have been learnt from the adaptive management approach. • a thorough understanding of the reproductive biology and regeneration mechanisms of w. salutaris is needed to refine the monitoring programme and the associated set of tpcs further. • understanding the drivers of the demand for the resource is essential if sanparks wants to reduce pressure on wild populations. employing field rangers to protect the trees may be effective in the short term, but without addressing the larger drivers of demand pressure on the plants will not be reduced. • linking management interventions to the appropriate drivers at appropriate scales should be a priority during the adaptive planning and management phases. • continuous engagement with user groups is essential to monitor and adapt to long-term social and economic impacts of harvesting programmes. • in the case of w. salutaris, a multi-pronged, multi-scaled approach seems to be most effective. • although w. salutaris is an endangered plant, an approach that involves a small-scale harvesting programme (controlled access) may be more effective than a non-harvesting approach (no access). • a clear conceptual framework of how the system works, what the drivers are and what response variables can be measured and monitored is essential to guide the development of clear programme objectives. the objectives should be linked to clear measurables and deliverables and associated time frames. • identifying and getting agreement on roles and responsibilities in multidisciplinary projects can be challenging, and requires an overarching strategic plan that is developed, implemented and monitored together with all relevant stakeholders. case study 3: sour fig (carpobrotus acinaciformis) harvesting in agulhas national park (back to top) introduction the agulhas region has a history of sour fig harvesting (cowling & richardson 1995; ed. hassan 2002; hudson 2009). two species, hottentot fig (carpobrotus edulis [l.] n.e. br.) and sour fig (carpobrotus acinaciformis [l.] l. bol.), occur here. however, local resource users prefer the suurvy for its sweeter fruits and greater yield (scheepers 2008a). the high demand for a single species highlights the potential for overuse yet the quantities harvested and the associated impacts on biodiversity have never been monitored. with the establishment of the agulhas national park (anp), the challenge of sustainable resource use was inherited by park management. legitimate resource users are recognised as those stakeholder groups that have traditionally harvested sour figs in the agulhas region. they include the poor and unemployed, schoolgoing children, pensioners, the public, tourists and local farm labourers. however, most resource users are residents of the local settlements of struisbaai and buffelsjag, where sour fig harvesting underpins subsistence and small-scale commercial use. faced with shortfalls in local law enforcement capacity, it has proven difficult to provide legitimate users with secure resource use rights and to exclude outsiders and opportunists (oström 1990; sandström 2009). large-scale commercial traders operating at the regional level often rely on local labour pools to create a monopoly on trade that undercuts small-scale local business (scheepers 2008a). consequently, the challenge of providing for a fair and equitable distribution of benefits from sour fig harvesting requires a multifaceted approach. faced with high current demands for the resource and a historical precedent of use, sanparks must make management decisions based on the best available science (biggs & rogers 2003). adaptive management is therefore necessary. because people’s livelihoods are affected by changes in the resource use management of the park, a sense of urgency drives the need for management experiments to trial different monitoring methods, governance arrangements and ways to integrate local and scientific knowledge to foster learning. socio-economic aspects fresh sour figs are used to make jams and preserves or are eaten as dried fruits. fresh figs are sold at r15/kg while dried figs are worth r10/kg on the local market. during a study of sour fig harvesting as a potential community development initiative, farabi and associates (2008) estimated that 3 kg of sour figs are currently harvested per hectare on between 3000 ha and 5000 ha of the park, producing an annual yield of 9000 kg – 15 000 kg. the annual contribution to the local economy (fresh and dried figs, combined) was estimated at r90 000–r225 000 (farabi & associates 2008). at the household level, an average annual income of r738 per person from sour fig harvesting was calculated for the period between 2005 and 2009. although only a seasonal resource, one benefit of sour fig harvesting is that people can bolster their cash reserves for the start of another year. the additional income is particularly important as most sour fig harvesters are pensioners (57%). only 29% of local resource users were income earners. ecological studies little monitoring of sour fig harvesting in the park has taken place. however, certain areas are preferentially selected by local resource users while others are avoided, which introduces the risk that some areas will be overexploited. although not of concern in itself while ecosystem integrity appears to be maintained, a lack of feedbacks in the socio-ecological system to prompt management interventions poses a greater, more long-term problem. during the 2009 harvesting season, the first monitoring plots for sour fig harvesting were established to generate baseline data. the survey results showed no significant differences in sour fig numbers across the park. this finding indicated that people’s resource use patterns were likely not driven by resource availability. however, further monitoring across a more representative sample of plots is required to test this hypothesis. the survey also did not find signs of sour fig harvesting for which there were many possible explanations. monitoring was undertaken at the beginning of the season while harvesting began later owing to a late fruiting season caused by unusual weather conditions. sour figs may also occur within a patchier distribution than expected, as indicated by gaps in people’s local knowledge of the best harvesting areas. household surveys amongst local resource users showed that only 2169 kg sour figs were harvested, supporting the theory that the harvesting season produced a low yield. however, this figure did not take into account illegal sour fig harvesting. the reliability of the survey data was also dependent on the local resource users reporting the actual quantities harvested. continued monitoring is required to gain a better understanding of current resource use patterns. however, interviews indicated that people’s resource use patterns are driven by a cultural preference for c. acinaciformis over c. edulis and resource accessibility, which is likely a complex function of multiple entry points to the park, the limited road network, the method of travel (i.e. whether walking or driving), and the distance covered by resource users in search of sour figs. most users (57%) stated that they harvested the preferred species but travelled greater distances now than in the past to harvest the same quantities. application of adaptive management to monitor sour figs in response to the overexploitation of sour figs in some areas, sanparks initially established a uniform harvest prescription of no more than 30% of the available yield across the park (farabi & associates 2008). this was never successfully implemented. without adequate stakeholder involvement and support, the harvest prescription was largely imposed by sanparks on the local resource users. local people questioned the legitimacy of the decision and perceived the restriction on harvesting as unfair, particularly as poor law enforcement failed to curb illegal use. local users also found the percentage of available yield harvested as an indicator for sustainable use difficult to understand. adaptive management emerged as an alternative. it entailed a process of collaborative problem framing, planning, implementation and evaluation together with local resource users and other relevant stakeholders (cundill & fabricius 2009). this participatory approach to management enhanced stakeholder engagement through the creation of platforms for dialogue concerning the need to monitor sour fig harvesting, and the creation of incentives for sustainable use. it also fostered better relations between the community and law enforcement authorities such that the increase in social capital helped buffer local resource users and law enforcement against future conflicts; in fact, registered resource users have started to report incidents of illegal use to the authorities. an adaptive management approach also allowed for the recognition of different intensities of use by local resource users across the park. consequently, separate harvesting areas were assigned to each of the two major resource user groups at struisbaai and buffelsjag. designated areas of high, moderate and no use also continue to allow for a diversity of impacts from sour fig harvesting across the park. this mosaic of harvesting intensities, in turn, informs an experimental design for the development of sustainable resource use tpcs across the park in future. however, indicators more appropriate to the local context are needed in order for resource users to understand and support the implementation of a monitoring system. as local users harvest sour figs in standard-sized shopping bags, the number of bags harvested and their mass represent potential alternatives to the percentage yield harvested. with continued monitoring, the quantities of sour figs harvested from the park can be compared with household surveys to identify discrepancies between observed and expected levels of use, monitor trends and inform research priorities. the calculation of preference indices (scheepers 2008b; shackleton 1993) for the different harvesting areas across the park can help highlight areas of overexploitation. the income generated from sales can be used to track financial benefits from sour fig harvesting for local people. the use of multiple indicators also makes it more difficult for opportunists and outsiders to break the rules (fabricius et al. 2007; oström 1990). integration of local and scientific knowledge in response to an unsuccessful first attempt to manage sour fig harvesting, sanparks adopted a more participatory approach, which aimed to link stakeholder aspirations to management actions (cundill & fabricius 2009). current stakeholder participation is consultative, as local resource users make inputs into the management process at every stage (fabricius 2004). participatory research techniques such as transect walks, questionnaire surveys, user group discussions and workshops capture inputs from local resource users to inform management decisions about appropriate monitoring indicators and local governance arrangements. creating more opportunities for dialogue has also improved communication between the park and local resource users. whereas communication mostly occurred during the build-up to the harvesting season in the past, year-round activities that include planning, monitoring, data analysis, reporting and reflection promote co-learning through the management process. as part of the new resource use protocols, for example, local resource users are required to make inputs into an annual report, working with sanparks scientists and the staff from the people and conservation division. in turn, high levels of support by sanparks and others have encouraged local resource users to become complaisant. while sanparks has played a greater advisory role to resource users in the past, poor local leadership and inadequate capacity building may explain the current lack of community ownership of the project. transition strategies are needed if greater responsibility is to be given to local resource users to monitor their own use in the long term. these strategies could include mentorship opportunities, leadership training, sharing experiences, reciprocal knowledge transfers and management workshops. flexible institutions local resource users were initially organised into respective community associations for buffelsjag and struisbaai. each group developed their own constitution and code of conduct, which sanparks recognised (fabricius et al. 2004). however, these rules were never revisited as member numbers increased and social cohesion within the groups became stressed. as conflicts arose, the community associations were ill equipped to resolve them. successive mergers and splits from the original groups occurred and as communication networks broke down, confusion resulted over membership lists and examples of nepotism were observed within the management structure. consequently, an investment in conflict resolution mechanisms, including the use of an independent arbitrator on a case-by-case basis, is necessary to strengthen local governance. over the long term, processes of deliberation, negotiation, and the development of social capital and trust amongst resource users must be in place as part of everyday management (sandström 2009). flexible rules for governing sour fig harvesting were developed to be sensitive to local conditions (fabricius 2004; oström 1990). these include harvesting times that are outside the hottest time of the day. the time restrictions also help local law enforcers to distinguish between legitimate and unlawful resource users, as illegal harvesting mostly takes place at night. however, as all offences are equally punishable, many registered users (43%) said that there is little incentive for sustainable resource use. instead, people stated that they were disadvantaged by ‘doing the right thing’, as this required the payment of an administration fee for a harvesting permit while illegal resource users were seldom caught. to influence people’s attitudes and actions in favour of sustainable use, graduated sanctions for offences and improved law enforcement efforts are required at the anp. lessons learnt from case study several lessons emerged from the implementation and ongoing management as described in this case study. • an adaptive management approach is necessary where faced with a historical precedent of use, high demands for the resource and little baseline data. as sour figs were harvested from the area prior to the establishment of the park, it was difficult for sanparks to implement restrictions on use pending the necessary research and monitoring to set sustainable use levels without adequate involvement from the local resource users. • as shown by shackleton and shackleton (2004), moderate socio-economic benefits from sustainable resource use can make a significant contribution to people’s livelihoods when available at strategic times of the year. the seasonal income from sour fig harvesting supplements household budgets over the december holiday season and helps kick-start the new year. • long-term monitoring data and appropriate ecological and socio-economic indicators of change are needed to inform tpcs for the sustainable use of sour figs at the anp. the number of bags harvested by local resource users, identification of preferred harvesting areas, and the income generated from sales may be good indicators of sustainable use but should be tested and refined through ongoing research and monitoring. • a participatory approach supports the legitimacy of management decisions regarding sustainable resource use (cundill & fabricius 2009). communication between the park, local resource users and other stakeholders was improved through the creation of additional platforms for dialogue, enhanced stakeholder relationships, and the incorporation of local knowledge into management. • a mosaic of harvesting intensities across the park promotes habitat heterogeneity, while the designation of a no-use area allows for ecosystem renewal (biggs & rogers 2003). the experimental design of high, moderate and no-use areas also allows for the development of multiple tpcs for sustainable use. • responsible facilitation and the co-management of resources require transitional strategies that will empower local resource users to become self-governing (fabricius et al. 2004). in the case of sour fig harvesters in the anp, these transitional strategies should focus on leadership development, training in monitoring resource use and conflict resolution. • there need to be adequate incentives for local resource users to support sustainable use practices. improved law enforcement and graduated sanctions for offences that distinguish between the actions of registered and unlawful resource users can create this incentive for sustainable sour fig harvesting in the anp. conclusions (back to top) the sanparks policy on resource use (sanparks 2008) embraces a systems view of sustainable resource use with ecological and socio-economic drivers that link the management of national parks to their surrounds. the case studies in this paper inform the implementation of the policy and promote learning through an adaptive management framework. one lesson is that the identification of appropriate indicators and the setting of acceptable limits of change should be part of a dynamic process informed by ongoing monitoring and research. however, faced with incomplete knowledge of the ecological and socio-economic factors driving resource use, the precautionary principle must always be applied (cooney 2004). in the first two case studies, the impacts of harvesting were initially underestimated, whereas they were overestimated in the third, with potentially large implications for socio-economic benefits from resource use. the zonation of different areas for use, as in the first and third case studies, sets up an experimental design to develop tpcs that support flexible rules for management. multiple overlapping sets of rules facilitate effective monitoring, and make it more difficult for oppportunists and outsiders to break the rules (fabricius et al. 2007; oström 1990). this applies particularly where incentives for sustainable use and resource protection are linked to strict law enforcement, as in the case of haversting pepper-bark tree and sour figs. a participatory research approach in all case studies enabled the incorporation of local knowledge into the management process. however, as shown for the sour fig case study, responsible facilitation and the co-management of resources require transitional strategies that empower local resource users to become self-governing (fabricius et al. 2004). without adequate capacity building, new dependencies can be created and local resource users may become complaisant. other challenges raised by the case studies include the development of too few ecological tpcs, with the exception of those for pepper-bark tree harvesting. a common explanation is the lack of long-term monitoring data and a poor understanding of the ecological drivers of resource use. further research into the population dynamics, demography and reproductive phenology of plant resources are required to refine indicators for monitoring and determine tpcs for their sustainable use. however, with the exception of the knp, using tpcs as part of the management of south african national parks is new. therefore, it will be some time before the usefulness of the concept can be evaluated at the national scale. no socio-economic tpcs for sustainable resource use have been developed. possible explanations include a historical bias towards ecological over social science research and limited in-house capacity to develop the necessary monitoring and evaluation systems. methods to quantify and monitor intangible benefits from sustainable resource use should also be developed. the case studies suggest multiple drivers of sustainable resource use across various scales. consequently, sanparks will need to invest in the necessary resources and capacity building to ensure effective management systems. partnerships with the private sector and local resource users, as in the case of seven-weeks fern harvesting, represent a possible mechanism for leveraging opportunities for sustainable resource use. references (back to top) abbot, j.i.o. & mace, r., 1999, ‘managing protected woodlands: fuelwood collection and law enforcement in lake malawi national park’, conservation biology 13(2), 418−421. doi:10.1046/j.1523-1739.1999.013002418.x biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring, and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 59−80, island press, washington dc. botha, j., witkowski, e.t.f. & shackleton, c.m., 2004, ‘the impact of commercial harvesting on warburgia salutaris (‘pepper-bark tree’) in mpumalanga, south africa’, biodiversity and conservation 13, 1675−1690. doi:10.1023/b:bioc.0000029333.72945.b0 bulte, e.h., van kooten, g.c. & swanson, t., 2003, ‘economic incentives and wildlife conservation’, viewed 21 july 2010, from http://www.cites.org/eng/prog/economics/cites-draft6-final.pdf clover, j. & eriksen, s., 2009, ‘the effects of land tenure change on sustainability: human security and environmental change in southern african savannas’, environmental science and policy 12, 53−70. doi:10.1016/j.envsci.2008.10.012 cooney, r., 2004, ‘the precautionary principle in biodiversity conservation and natural resource management: an issues paper for policy-makers, researchers and practitioners’, iucn, cambridge. cowling, r. & richardson, d., 1995, fynbos: south africa’s unique floral kingdom, fernwood press, vlaeberg. cundill, g. & fabricius, c., 2009, ‘monitoring in adaptive co-management: toward a learning based approach’, journal of environmental management 90, 3205−3211. doi:10.1016/j.jenvman.2009.05.012, pmid:19520488 cunningham, a.b., 2001, applied ethnobotany: people, wild plant use and conservation, earthscan publications, london. diederichs, n., 2006, commercializing medicinal plants: a southern african guide, sun press, stellenbosch. fabricius, c., 2004, ‘the fundamentals of community-based natural resource management’, in c. fabricius, e. koch, h. magome& s. turner (eds.), rights, resources and rural development: community-based natural resource management in southern africa, pp. 3−43, earthscan, london. fabricius, c., folke, c., cundill, g. & schultz, l., 2007, ‘powerless spectators, coping actors, and adaptive co-managers: a synthesis of the role of communities in ecosystem management’, ecology and society 12(1), 29, viewed 17 august 2010, from http://www.ecologyandsociety.org/vol12/iss1/art 29 fabricius, c., koch, e., magome, h. & turner, s., 2004, rights, resources and rural development: community-based natural resource management in southern africa, earthscan, london. farabi and associates, 2008, ‘agulhas national park: suurvye harvesting’, consultants’ report, farabi and associates, pretoria. geldenhuys, c.j., 1993, ‘composition and dynamics of plant communities in the southern cape forests’, deliverable report no. for-dea 612, division of forest science and technology, council for scientific and industrial research, pretoria. geldenhuys, c.j., 1994, ‘review of studies of the fern rumohra adiantiformis in the southern cape forests: implications for sustainable management’, deliverable report no. for-dea 758, division of forest science and technology, council for scientific and industrial research, pretoria. geldenhuys, c.j., 2000, ‘sustained yield determination of forest and woodland produce’, in d.l. owen & w.j. vermeulen (eds.), south african forestry handbook 2000, pp. 643−650, southern african institute of forestry, pretoria. geldenhuys, c.j. & van der merwe, c.j., 1986, ‘effects of frond harvesting on growth of the fern rumohra adiantiformis in the southern cape forests’, research report s. 30/85, saasveld forestry research centre, george. geldenhuys, c.j. & van der merwe, c.j., 1988, ‘population structure and growth of the fern rumohra adiantiformis in relation to frond harvesting in the southern cape forests’, south african journal of botany 54(4), 351−362. geldenhuys, c.j. & van der merwe, c.j., 1994, ‘site relations and performance of rumohra adiantiformis in the southern cape forests’, deliverable report for-dea 759, division of forest science and technology, council for scientific and industrial research, pretoria. hansen, a.j. & defries, r., 2007, ‘ecological mechanisms linking protected areas to surrounding lands’, ecological applications 17, 974−988. doi:10.1890/05-1098, pmid:17555212 hassan, r.m. (ed.), 2002, accounting for stock and flow values of woody land resources: methods and results from south africa, ceepa, university of pretoria, pretoria. hilton-taylor, c., scott-shaw, r., burrows, j. & hahn, n., 1998, ‘warburgia salutaris’ in iucn red list of threatened species, viewed 13 april 2010, from http://www.iucnredlist.org hoole, a. & berkes, f., 2010, ‘breaking down fences: recoupling social-ecological systems for biodiversity conservation in namibia’, geoforum 41, 304−317. doi:10.1016/j.geoforum.2009.10.009 hudson, v., 2009, ‘the harvesting of sour figs (carpobrotus edulis) on capenature reserves on the agulhas plain’, paper presented at botanical society of south africa conference, bredasdorp, 4–7th august. international union for conservation of nature, 2010, iucn red list of threatened species, viewed 01 april 2010, from http://www.iucnredlist.org iucn. see international union for conservation of nature. kok, h.r., 1998, ‘studie na die volhoubare benutting van seweweeksvaring (rumohra adiantiformis) uit die inheemse woude van die suid-kaap en tsitsikamma’, msc dissertation, department of environmental management and ecology, university of the orange free state. lindsey, p.a., roulet, p.a. & romañach, s.s., 2007, ‘economic and conservation significance of the trophy hunting industry in sub-saharan africa’, biological conservation 134, 455−469. doi:10.1016/j.biocon.2006.09.005 lumbe, a., 2008, e-mail, 13 november 2008, andrewl@sanparks.org mander, m., 1998, marketing of indigenous medicinal plants in south africa. a case study in kwazulu-natal, food and agricultural organisation, united nations, rome. milton, s.j., 1987a, ‘growth of seven-weeks fern (rumohra adiantiformis) in the southern cape forests: implications for management’, south african forestry journal 143, 1−4. milton, s.j., 1987b, ‘effects of harvesting on four species of forest ferns in south africa’, biological conservation 41, 133−146. doi:10.1016/0006-3207(87)90116-9 milton, s.j., 1991, ‘slow recovery of defoliated seven-weeks fern rumohra adiantiformis in harkerville forest’, south african forestry journal 158, 23−27. milton, s.j. & moll, e.j., 1987, ‘utilisation potential of rumohra adiantiformis in the southern cape forests’, forest biome project report, ecosystem programmes, foundation for research development/council for scientific and industrial research, pretoria. milton, s.j. & moll, e.j., 1988, ‘effects of harvesting on frond production of rumohra adiantiformis (pteridophyta: aspidiaceae) in south africa’, journal of applied ecology 24, 725−743. doi:10.2307/2403858 murphree, m.w., 2000, ‘community-based conservation: old ways, new myths and enduring challenges’, key address at the conference on african wildlife management in the new millennium, college of african wildlife management, mweka, 13–15th december. nagothu, u.s., 2001, ‘fuelwood and fodder extraction and deforestation: mainstream views in india discussed on the basis of data from the semi-arid region of rajasthan’, geoforum 32, 319−332. doi:10.1016/s0016-7185(00)00034-8 natural justice, 2010, ‘biocultural protocol of the traditional health practitioners of bushbuckridge’, natural justice, mpumalanga tourism and parks agency and the traditional health practitioners of bushbuckridge, viewed 19 april 2011, from http://www.naturaljustice.org/images/naturaljustice/bushbuckridge%20k2c%20bcp.pdf oström, e., 1990, governing the commons: the evolution of institutions for collective action, cambridge university press, cambridge. parr, c.l., woinarski, j.c.z. & pienaar, d.j., 2009, ‘cornerstones of biodiversity conservation? comparing the management effectiveness of kruger and kakadu national parks, two key savanna reserves’, biodiversity conservation 18, 3643−3662. doi:10.1007/s10531-009-9669-4 peters, c.m., 1996, ‘the ecology and management of non-timber forest resources’, technical paper no. 322, world bank, washington dc. reid, h. & turner, s., 2004, ‘the richtersveld and makuleke contractual parks in south africa: win-win for communities and conservation?’, in c. fabricius, e. koch, h. magome & s. turner (eds.), rights, resources and rural development: community-based natural resource management in southern africa, pp. 223−234, earthscan, london. reid, h., fig, d., magome, h. & leader-williams, n., 2004, ‘co-management of contractual national parks in south africa: lessons from australia’, conservation and society 2(2), 377−409. sanbi. see south african national biodivesity institute. sanparks. see south african national parks. sandström, c., 2009, ‘institutional dimensions of comanagement: participation, power, and process’, society and natural resources 22(3), 230−244. doi:10.1080/08941920802183354 scheepers, k., 2008a, ‘research trip to agulhas np: towards formalizing suurvye harvesting in the park’, internal report, cape research centre, south african national parks. scheepers, k., 2008b, ‘harvesting strategies of fuelwood and kraalwood users at machibi: identifying the driving factors and feedbacks’, phd thesis, department of environmental science, rhodes university. scherl, l.m., wilson, a., wild, r., blockhus, j., franks, p., mcneely, j.a., et al., 2004, ‘can protected areas contribute to poverty reduction? opportunities and limitations’, iucn, cambridge, uk. doi:10.2305/iucn.ch.2005.6.en shackleton, c.m., 1993, ‘fuelwood harvesting and sustainable utilisation in a communal grazing land and protected area of the eastern transvaal lowveld’, biological conservation 63, 247−254. doi:10.1016/0006-3207(93)90720-l shackleton, s. & shackleton, c., 2004, ‘everyday resources are valuable enough for community-based natural resource management programme support: evidence from south africa’, in c. fabricius, e. koch, h. magome & s. turner (eds.), rights, resources and rural development: community-based natural resource management in southern africa, pp. 135−146, earthscan, london. south african national biodivesity institute, 2009, national red list of south african plants, south african national biodivesity institute, pretoria. south african national parks, 2008, sanparks resource use policy, south african national parks, pretoria. stankey, g.h., clark, r.n. & bormann, b.t., 2005, ‘adaptive management of natural resources: theory, concepts, and management institutions’, general technical report pnw-gtr-654, united states department of agriculture, forest service, pacific northwest research station. swemmer, l.k., 2006, ‘report on rare and invasive species programme’, internal report, kruger national park, south african national parks. swemmer, l.k., 2010, ‘warburgia salutaris conservation programme’, internal report, scientific services, south african national parks. vermeulen, w.j., 2009, ‘the sustainable harvesting of non-timber forest products from natural forests in the southern cape, south africa: development of harvest systems and management prescriptions’, phd thesis, department of conservation ecology and entomology, stellenbosch university. vermeulen, w.j., du plessis, l. & herd, h., 2005, ‘harvesting of seven weeks fern (rumohra adiantiformis) from natural forests of the southern cape’, in participatory forest management. case studies in south africa 2005, pp. 83−88, department of water affairs and forestry, pretoria. von breitenbach, f., 1968, southern cape indigenous forest management manual, foundations of management, department of forestry, george. von maltitz, g.p. & shackleton, s.e., 2004, ‘use and management of forests and woodlands in south africa: stakeholders, institutions and processes from past to present’, in m.j. lawes, h.a.c. eeley, c.m. shackleton & b.g.s. geach (eds.), indigenous forests and woodlands in south africa: policy, people and practice, pp. 109−135, university of kwazulu-natal press, durban. willis, c.b., 2004, ‘policy frameworks pertaining to the conservation and sustainable use of forests and woodlands in south africa’, in m.j. lawes, h.a.c. eeley, c.m. shackleton & b.g.s. geach (eds.), indigenous forests and woodlands in south africa: policy, people and practice, pp. 77−107, university of kwazulu-natal press, durban. zambatis, n., 2006, ‘report on the state of the pepper-bark tree (warburgia salutaris) in the knp’, internal report, kruger national park, south african national parks.